Front Matter
Source: Systematic Botany Monographs, Vol. 50, Systematics of Oenothera Section Oenothera
Subsection Oenothera (Onagraceae) (Mar. 17, 1997)
Published by: American Society of Plant Taxonomists
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SYSTEMATIC
BOTANY
MONOGRAPHS
VOLUME 50
Systematics of Oenothera Section Oenothera
Subsection Oenothera (Onagraceae)
WernerDietrich
Warren L. Wagner
Peter H. Raven
THE AMERICAN
SOCIETY OF PLANT TAXONOMISTS
17 March
1997
SYSTEMATIC BOTANYMONOGRAPHS
ISSN 0737-8211
Copyright ? 1997
The American Society of Plant Taxonomists
All rights reserved
ISBN 0-912861-50-9
Printed
in the United
States of America
Editor
CHRISTIANEANDERSON
University
North University
of Michigan
Building,
Herbarium
Ann Arbor, Michigan
Editorial
Iowa State University
Garden
WALTER S. JUDD
University
Montana
State University
MELISSALUCKOW
PETER C. HOCH
Botanical
Committee
MATTLAVIN
LYNN G. CLARK
Missouri
48109-1057
of Florida
JACQUELYNA. KALLUNKI
New York Botanical Garden
Cornell University
NORTON G. MILLER
New York State Museum
WARREN L. WAGNER
Smithsonian
Institution
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BIENNIS
OENOTHERA
SYSTEMATICS OF OENOTHERA SECTION OENOTHERA
SUBSECTION OENOTHERA (ONAGRACEAE)
WernerDietrich
Botanisches InstitutderUniversitatDusseldorf
Universitatsstr.
1
D-40225 Dusseldorf, Germany
Warren L. Wagner
Department of Botany, MRC
166
of Natural History
National Museum
Smithsonian Institution
Washington, DC 20560
Peter H. Raven
Missouri
Botanical Garden
P. 0. Box 299
St. Louis, MO 63166-0299
ABSTRACT.
plex species
had a long history
of permanent
that is consistent
phology
names
cluding many
the 388 validly
PTH
widespread
many
with
(156),
published
species
for naturalized
specific
European
and eight PTH
ferentiated
species.
into species
crospecies)
genomes
There
0.
approach
the greatest
bivalent
to reflect
are three basic
number
herbarium
(562) were
lineages within
clonal
of their genomic
history
Oenothera
PTH
wolfii
appear
and the polymorphic
sumably
derived
ica) appears
0. villosa,
from 0. elata
tained sporadically
presumably
legheny Mountains
are both PTH
while
from 0. elata
with BB genomes
in 0. grandiflora,
derived
directly
subsp. hookeri,
to have been derived
of eastern North America
species
to have been derived
directly
from 0.
species.
with
relating
0.
argillicola,
The
in the western
second
lineage
III. Self-incompatibility,
elsewhere
from 0. grandiflora.
shale barren endemic,
(widespread
The
which
in the subsection.
third lineage
consists
has a CC genomic
type, and
a reliable
to genome
longissima,
0.
outcrossed
longissima
I species,
plastome
coastal
dif
(or mi
plastome
I (0. elata, 0.
two AA
the ple
homozygosity,
and to provide
former, a rare Pacific
The
0. villosa
not occur
does
to the
into perceptibly
subsect. Oenothera,
elata. The other
subsp. hirsutissima.
and plastome
published.
populations
complexes,
of the group
and plastome
genomes
and genomic
and fix variation
the essentially
the evolutionary
are delimited
species
0. wofii, and 0. villosa). The first three species have plesiomorphic
characteristics
of mostly
large flowers and formation of bivalents or small rings of chromosomes
during meiosis. Oenothera
jamesii
in
analyzed,
jamesii,
and 0.
and mor
plastome,
(292) have been applied
in meiosis,
serve to partition
aggregates
have AA
names
func
as a phar
0. parviflora,
and 0. villosa),
including
there are three major genomes, designated A,
formation
to the composition
according
and extensive
garden,
that have never been validly
I, II, III, IV, and V). Five
of the PTH system
and recently
of the genome,
published
has
system
oakesiana,
the subsection
flowers,
Five of the species
composition.
All
literature,
names
(plastomes
outcrossed
characters
classifications.
Within
common
genetic
in studies of chloroplast
and plastome,
features
using
com
as "Euoenothera,"
the anomalous
significant
genome
the most
representing
known
subsection,
that elucidated
cytological,
system
glazioviana,
populations.
delimited
for identification.
and plastome
0.
strains. Our
morphological
acid. New
taxonomic
and infraspecific
The properties
true-breeding
This
between
from the cytogenetics
(0. biennis,
features of primarily
siomorphic
means
interactions
other angiosperm
especially
B, and C, and five basic plastid
associated
genetic
into a revised
incorporated
subsect. Oenothera,
(PTH). The group has been
heterozygosity
crop for the fatty acid y-linolenic
studies were
of Oenothera
are recognized.
and genetic work
cytogenetic
self-incompatibility,
macological
revision
13 species
of study, notably
translocation
tion, genetics,
Of
In this comprehensive
in the Onagraceae,
group
0.
is pre
endemic,
half of North Amer
consists
of two species
a plesiomorphic
Oenothera
nutans
feature
of the very distinctive
composition
re
is a PTH
Al
and is the only
2
SYSTEMATICBOTANYMONOGRAPHS
species
with
V. Three
plastome
ranges had hybrid origins:
additional
0. biennis
PTH
species
with wide,
(AB or BA with plastome
primarily
VOLUME 50
eastern North
IIor III); 0. oakesiana
natural
American
(AC with plastome
IV);
in Europe. Hybrids
and 0. parviflora
(BC with plastome
IV). All three species are widely naturalized, especially
the PTH species. When
occur between many of the species (19 known combinations),
especially
they represent
a more widespread
species
and plastome
of recent hybrid origin
in Europe
via hybridization
the latter in Italy. Oenothera
in the taxon they phenotypically
they are included
phenotype,
has the same genome
composition.
are recognized:
outside
Exceptionally,
0. glazioviana
the indigenous
glazioviana
(AB-III),
and 0.
range of the subsection,
has achieved
most
two morphologically
a nearly worldwide
closely
distinctive
stucchii
resemble
(AA-I). Both
the former possibly
and that
additional
PTH
originated
in England
and
distribution.
INTRODUCTION
in a series treating the five subsections of
is the fourth and final publication
sect. Oenothera
(70 species), divided into its present arrangement based on
studies and on genome and plastome relationships inferred
comparative morphological
from experimental analysis, especially crossing behavior (Stubbe & Raven 1979a). The
other papers in this series are: Dietrich (1977) on Oenothera subsect. Munzia (45 species,
This
Oenothera
study by Dietrich, unpubl., suggests there are only 39 species); Dietrich,
Raven, and Wagner (1985) on Oenothera
subsect. Emersonia
(4 species); Dietrich and
Wagner (1988) on Oenothera
subsect. Raimannia
subsect.
(11 species) and Oenothera
Nutantigemma
(3 species). The detailed assessment presented here basically follows the
and extended and outlined
taxonomic philosophy suggested by Cleland (1972: 316-318),
in 1979 by Raven, Dietrich, and Stubbe. This paper presents the first worldwide compre
but subsequent
hensive
revision of Oenothera sect. Oenothera subsect. Oenothera (13 species), the most
complex group of species in the Onagraceae. We reviewed the copious literature on this
group of species and searched for all of the scientific names ever applied to any entity we
here include within subsect. Oenothera. All names (562) are formally treated; those
validly published (388) are included in the taxonomic treatment. Names not validly pub
lished as well
as those applied to hybrids and experimental
strains are listed in a separate
section.
Species of Oenothera subsect. Oenothera have been useful experimental organisms
in studies of chloroplast
function, self-incompatibility,
genetic interactions between
genome and plastome, and, especially, complex heterozygosity, which have occupied nu
merous workers for over a century of genetic and cytogenetic analyses. Species of the sub
section also have recently become a pharmacological
crop for the extraction of the fatty
acid from the seeds (Wolf et al. 1983; Bosisio 1990).
Although hundreds of papers have been published on themembers of Oenothera sub
sect. Oenothera, there has not been an overall philosophy on which a taxonomic treatment
acid y-linolenic
could be developed
and the Onagraceae.
known genetically
that is consistent with those applied throughout the rest of Oenothera
As a result, subsect. Oenothera presents a paradox in that it is as well
and biologically as almost any group of plants, but despite this, its tax
onomy has been piecemeal and there have been widely dissimilar approaches to its clas
sification. Because subsect. Oenothera is well studied and represents a group of historical
importance to the development of biological thought in Hugo de Vries's theory of muta
tion (see Mayr, 1982), as well as constituting a group of current importance tomolecular
biology, and because this group represents the best-studied example of the uncommon and
anomalous phenomenon of permanent translocation heterozygosity
(PTH), it is a com
monly used example in general evolutionary texts (e. g., Dobzhansky
et al. 1977; Futuyma
1979; Grant 1975, 1981). In fact, Grant includes the evolution of this group under a spe
1997
OENOTHERA
3
cial category of the heterogamic complex, which consists of "heterogamic microspecies"
(series of true-breeding forms) and their bivalent-forming
ancestors.
Permanent translocation heterozygosity
(PTH) occurs in only about 59 species in
seven families of plants (cf. Holsinger & Ellstrand 1984), including Onagraceae
(49
species: Oenothera, 45 spp.; Gaura, 2 spp.; Gayophytum,
1 sp.; and Calylophus,
1 sp.;
Raven
1979), Campanulaceae
(2 spp.), Commelinaceae
(2 spp.), Clusiaceae
(2 spp.), Iri
daceae (3 spp.), Paeoniaceae (2 spp.), and Papaveraceae (1 sp.). There are variations in the
features of PTH in these families, and in some cases it is not clear if they are truly PTH;
they are included only tentatively in this list (for discussion
theoretical model has been proposed to account
1984). A
see Holsinger & Ellstrand,
for the evolution of PTH
(Holsinger & Feldman
1981).
Species of Oenothera subsect. Oenothera are. known nearly worldwide, especially in
Europe, but they are indigenous only toNorth America, with a few populations of 0. elata
subsp. elata extending as far south as Panama. Several of the species of subsect.
were introduced to Europe at least three centuries ago, and, largely through hy
bridization, numerous new phenotypes have originated there. Renner (1942) recognized
at least 18 PTH species in Europe, and many others have been described as species since
then (e.g., Hudziok 1964, 1968; Rostan'ski 1985). This approach of giving formal names
to every different true-breeding phenotype discovered also has been used inNorth Amer
ica (e.g., Gates 1936). If this approach were taken to its extreme, hundreds or perhaps
Oenothera
thousands of specific names would result. The study needed to begin to identify the re
sulting entities with anything but rudimentary accuracy would be extraordinary. Because
there has been no overall taxonomic treatment of the group, many names have persisted
in the literature, leaving no one with a clear idea of what classification and what names to
use.
The lifelong cytogenetic study of the group by Ralph Cleland (summarized in 1972;
see also Harte, 1994), followed byWilfried Stubbe (1953, 1959, 1960, 1964, 1980; Stubbe
& Raven 1979a) and several other workers, such as C. D. Darlington and E. Steiner, cou
pled with the taxonomic studies attempting to put the cytogenetic results into perspective,
culminated in bringing much order to the classification of the group. Philip Munz (1949,
1965) provided
a complete treatment for Oenothera subsect. Oenothera for North Amer
ica, but did not include any material or names published outside of the indigenous area of
distribution. Raven et al. (1979) provided an overall outline of the classification,
taking
full consideration of the cytogenetic
and genetic work and the very large naturalized
ranges. The present work provides the details and justification of this system and in a few
places modifies it.
For this revision of the taxonomy of Oenothera subsect. Oenothera, new information
was gathered from cultivation of strains from 344 localities throughout the natural and
naturalized ranges, made possible largely through the initiative of Peter Raven. We also
examined over 30,000 herbarium specimens. New cytological observations are provided
for 562 individuals, representing all 16 taxa. A major effort was made to locate and ana
lyze every name published in the widely-scattered
literature. We have made a complete
analysis of the extensive nomenclature of this complex group, including those names that
cannot be considered validly published, primarily from the genetics literature. A total of
562 names are included.
Previous taxonomic studies (Seringe 1828; Fischer & Meyer 1835; Spach 1835; Tor
rey & Gray 1840; Rose 1905; Munz 1949, 1965; Raven 1968; Rostan'ski 1985) are either
out-of-date, use a different taxonomic philosophy, or are incomplete. The results of Cle
SYSTEMATICBOTANYMONOGRAPHS
4
land's, Renner's,
1972), coupled with
summary, see Cleland,
crossing
and cytological
studies (for
recent crossing
and cytological
analyses
earlier experimental
and Stubbe's
VOLUME 50
by
(1987), Schumacher and
Steiner (1993), Wasmund and Stubbe (1986), Steiner and Stubbe (1984, 1986), Raven et
al. (1979), Stubbe and Raven (1979a), and Werner Dietrich (at the Botanical Institute of
the University of Dusseldorf) have been used to develop the classification presented here,
Drillisch
(1975), Wasmund
(1980,
1984,
and to understand better the relationships
1990),
Schumacher
and origins of the species of Oenothera
subsect.
Oenothera.
studies
of the taxa were based on extensive comparative morphological
on plants grown in the experimental garden in Dusseldorf and
Descriptions
made by Werner Dietrich
studies of herbarium
on extensive
principally
at theMissouri
Numerous
specimens
Botanical
strains from within
by Werner
Dietrich
and Warren Wagner,
Garden.
the naturalized
distributions
of all of the species from
various countries were studied, obtained from colleagues or by the international seed ex
change of the botanical gardens. These strains were cultivated at Dusseldorf. During the
past 20 years collections of the 13 species from a total of 440 localities have been stud
ied, of which 344 were examined cytologically. An additional 161 strains from R. Cle
land's research collection and 37 from 0. Renner's collection were cultivated at Dussel
dorf. In all, we have analyzed 638 strains of this group of Oenothera. These experimental
garden and herbarium
studies,
including
cytogenetics,
experimental
crossing
behavior,
of breeding systems, coupled with the cytoge
by Cleland and Renner, form the basis of the
and assessment
comparative morphology,
netic and evolutionary studies, especially
taxonomy presented here.
Material from at least 186 herbaria representing
about 12,000
localities worldwide
by at least one of us over the past 16 years. Records provided by Rostan'ski
from an additional 37 herbaria are incorporated for Oenothera biennis. In 1981, 1986, and
was examined
1990, Werner Dietrich
Garden,
examined
at the herbarium of the Missouri
and Warren Wagner,
approximately
can herbaria. Werner Dietrich
17,000
loaned from primarily North Ameri
specimens
travelled
subsequently
Botanical
to the United
States herbaria of F,
GH, MT, NA, NY, PH, and US, and the European herbaria of BM, BR, G, K, KRAM,
KTU, L, LY, M, P,WRSL, and Z to study an additional 8,000 specimens. After compil
ing these specimen
dorf to complete
The herbarium
borrowed
data Werner Dietrich
the worldwide
investigations
distribution
were
tion of the hundreds of strains cultivated
a further 5,000
specimens
at Duissel
studies.
supplemented with data gathered by the examina
in the common garden component of the project.
important, because many of the useful diag
This aspect of the project was particularly
nostic characteristics, particularly of the PTH species, are obscured by pressing and dry
ing. The collection data from all sources were combined in the preparation of distribution
maps for the worldwide
range of each species.
EXPERIMENTAL
STUDY OF OENOTHERA
SUBSECTION
OENOTHERA
Oenothera subsect. Oenothera, known in the literature as "Euoenothera," has had a
long history of scientific study resulting in hundreds of research papers and several books,
including
the excellent
(1989a) and Harte
group. Other
summary by Cleland
(1994), which
(1972)
and more
recent ones by Stubbe
recount nearly a century of experimental
than a few descriptive works,
investigations
of Oenothera
studies of the
began with Hugo
OENOTHERA
1997
5
era of study of mutation and its relationship
to evolution and speciation (see Cleland 1972; Stubbe 1972; Mayr 1982; Nei 1987). De
that new
Vries believed, based on his study of Oenothera lamarckiana (= 0. glazioviana),
species could be formed by single mutations. He conducted breeding experiments for
many generations of these plants and found that they continually produced small numbers
de Vries's
studies that opened up the modem
of aberrant forms. De Vries's studies were followed by decades of experimental work on
the group, especially by Renner and later Cleland, that elucidated many unique properties
of these plants, including the anomalous PTH genetic system, and by extensive cytologi
cal, genetic, and more recently molecular
investigations. This body of work showed that
subsect. Oenothera, which played the central role in the development of de Vries's ideas
on speciation, exhibits a complex variation pattern because of the unique properties of
PTH, features that could not be generalized for evolutionary theory. Despite this, the basic
ideas put forth by de Vries on mutation were correct, and true cases were discovered
shortly afterward (Stubbe 1972), making
his theory an important one in evolutionary
bi
ology.
Oenothera subsect. Oenothera is the most studied group of species in the genus and
the Onagraceae. Hundreds of papers have been published since de Vries's first publication
in 1895 on the introduced 0. glazioviana
in the Netherlands. The focus of most of these
papers is on genetics and cytogenetics, and in the past few decades also on other subjects,
such as evolution of the group, behavior of plastids, genetic variation in populations, mol
ecular genetics, chemistry, and taxonomy. One reason why Oenothera has been so inten
sively studied is that it is one of the few taxa inwhich the obscuring effect of genetic vari
ance is naturally limited (Mulcahy 1995). Phenomena which are often difficult to detect,
such as selective fertilization (Schwemmle
1968), nonrandom interactions between dif
ferent ovule and pollen genotypes, style/pollen interaction, and competition between de
can all be studied effectively in Oenothera (Mulcahy 1995).
summary of the literature is provided to give access by topic to the
most important publications and the relevant researchers in each area of investigation. The
summary is arranged by author in chronological order within each topic. Publications up
veloping microspores
The following
toCleland's classic book, Oenothera: cytogeneticsand evolution (1972) are not cited in
the usual format here, because he included an essentially complete bibliography of the lit
erature up to that time. For authors cited by Cleland only the name is listed; for more re
cent work citations are given.
Anatomy:
Chemistry:
Carlquist
Bosisio
(flavonoid
(1975).
(1990) (y-linolenic acid); Cretti (1996); Howard et al. (1972)
studies); Kawano et al. (1995) (floral volatiles); Wolf et al. (1983) ('y
linolenic acid); Zinsmeister
and Bartl (1971)
(flavonoid
studies).
Cytology: Catcheside; Cleland; Darlington; Davis; Gates; Lutz; Shull.
Developmental
biology: Harte (1994).
Embryology: Harte (1994); Noher de Halac and Harte (1977); Renner; Sniezko and
Harte (1984); Tobe and Raven (1985).
Genetics: Bartlett; Cleland; Davis; Emerson; Gates; Harte (1994); Oehlkers; Shull;
Renner; Steiner; Stubbe (1989b); de Vries.
Genetics and plastid behavior: Chiu et al. (1988); Cleland; Epp (1973); Epp and
Parthasarathy (1987); Gordon et al. (1980); Grun (1976); Kutzelnigg and Stubbe
(1974); Kutzelnigg et al. (1975a, 1975b); Renner; Schotz; Stubbe and Herrmann
(1982); Winter and Herrmann (1988).
SYSTEMATICBOTANYMONOGRAPHS
6
Genetic
variation
structure: Levin
and population
VOLUME 50
et al. (1972), Levin
(1975), Levy
and Levin (1975), Levy et al. (1975), Levy and Winterheimer
(1977); summa
rized in Holsinger and Ellstrand (1984).
Molecular genetics and phylogeny: Gordon et al. (1981, 1982); Hachtel et al. (1991);
Hildebrandt et al. (1984); Sears and Herrmann (1985); Winter and Herrmann
(1988).
Pollen
development
and morphology:
Praglowski
et al. (1987);
Takahashi
and
Skvarla (1990).
Seed coat anatomy: Tobe et al. (1987).
Seed ecology: Gross and Kromer (1986).
Taxonomy: Bartlett; Gates; Hudziok
(1964,
Raven
et al. (1979); Renner; Rostan'ski
1974); Munz; Raven (1968);
in 1985); Soldano (1979,
1968,
(summarized
1983).
PERMANENTTRANSLOCATIONHETEROZYGOSITY
Permanent
translocation heterozygosity
tion of the genus Oenothera
(PTH) has been very important in the evolu
It occurs in sev
and several other genera of the Onagraceae.
eral sections of the genus, but nowhere
is it better developed
than in Oenothera
subsect.
The genera of tribe Onagreae have chromosomes with highly pycnotic, con
densed proximal regions that are flanked by less densely contracted distal segments; the
are characteristically metacentric
chromosomes
(Kurabayashi et al. 1962; Cleland 1972;
Raven 1979). This chromosome morphology
is associated with the regular occurrence of
Oenothera.
rings of chromosomes,
resulting from reciprocal translocations. These rings, variable in
occur widely in the tribe and are
size and involving variable numbers of chromosomes,
frequently associated with outcrossing species. They thus form linkage groups involving
more
than one bivalent.
The phenomenon
the specialized
of reciprocal
translocations
reaches an endpoint of development
system known as PTH. The most well-known
tem are the members
of Oenothera
anisms were worked
out. This
(Futuyma 1979).
subsect. Oenothera,
species possessing
in which
in
this sys
the structure and mech
system represents the ultimate in linkage disequilibrium
In these plants each of the seven haploid chromosome complements
is
connected
through reciprocal translocations, making the entire genome behave as a single
linkage group. A major feature of this evolutionary curiosity is related to the way recom
bination is restricted (Cleland 1972; Raven 1979; Harte 1994). The reproduction of es
sentially identical genotypes and phenotypes results in populations in which a relatively
high proportion of the individuals are suited for a particular set of ecological parameters.
The habitats of the PTH species are usually marginal relative to those of the outcrossing
species to which they are most closely related. This system allows only one or two basic
genotypes
to be reproduced virtually unchanged
also allowed, through hybridization,
genetic features that appear tomake
vironments. A PTH species
populations
having
in each generation.
the immediate and permanent
them well able to colonize
in the sense employed
similar morphological
This mechanism
and persist inmarginal
here is an aggregation
and genetic
has
fixation of new sets of
en
of true-breeding
attributes.
The genetic mechanisms
that control the formation of the PTH system were largely
discovered and worked out by Otto Renner (see Cleland, 1972). In addition to the translo
cations, the system requires balanced lethals, which prevent the formation of the ho
OENOTHERA
1997
mozygous
(most easily observed
combinations
7
as ca. 50% infertile pollen),
self-pollina
tion, and alternate disjunction of the chromosomes during meiosis. Cleland and his stu
dents studied in detail the end arrangements of the chromosomes
through experimental
in
hybridization of hundreds of wild strains throughout North America
(summarized
1972). The attributes, evolution,
Holsinger
and Ellstrand
and systematic occurrence of this system are reviewed by
(1984).
in Oenothera
The taxonomic quandary
such as Crepis, Hieracium,
Rubus,
is similar to those in agamospermous
Taraxacum,
or Alchemilla,
where
genera,
numerous
mi
crospecies have been described. In these apomictic taxa some of the most intricate pat
terns of variation in the flowering plants are known (see Fryxell 1957; Grant 1981). Lit
erally hundreds of the variants have been given scientific names inmany of these genera.
The breeding system in species of Oenothera subsect. Oenothera
is exclusively
sexual;
this, the PTH forms actually behave like clonal organisms. Each new phenotype
that arises via occasional mutation, recombination, or frequent hybridization results in a
despite
new true-breedingform.
In the revision presented here, we accept 13 species, 8 of which have PTH. The pe
culiarities of the PTH species have led to the description of hundreds of new species, and
further study would yield hundreds more. The recognition of so much essentially individ
ual variation would neither contribute to our understanding of the group as a whole nor
result in the creation of taxonomic units even approximately
other parts of the genus. Therefore we delimit
equivalent
to those found in
in a broad sense, based on the
five plastome types. We agree with
the species
fundamental three genomic types and the associated
Cleland, who cautioned taxonomists against the immense splitting (1972, p. 316), which
would result in a taxonomic system in which even the specialist would lose the overview.
Because
of the very specialized nature of the biology of Oenothera subsect. Oeno
a
thera,
considerable number of terms have been invented or modified from other defini
tions to describe various aspects of the PTH mechanisms
and behavior. Many of these are
they are relevant to understanding our taxo
specific to the genetics literature. Because
nomic philosophy we briefly characterize the most important ones here. Whole chapters
could be developed to examine most of them; however, largely because this is already
available (Cleland 1972; Holsinger & Ellstrand 1984; Harte 1994), we provide only short
definitionshere.
a genetically controlled system in
lethals (Muller 1917): in Oenothera,
which homozygosity
of nonallelic recessive lethal genes results inmortality, ei
ther sporophytic or gametophytic, when the parent plant is autogamous. This sys
tem prevents the formation of the homozygous
combinations
or, in the PTH
Balanced
species, makes the young embryos with homozygous complexes lethal.
Complex heterozygote
(Renner 1917): a plant with its two genomes differing in the
chromosome segmental end arrangements through reciprocal translocations. The
end point situation is a ring of all chromosomes
(in Oenothera, 14 chromosomes,
written 0)14) formed at meiotic metaphase I followed by alternate disjunction of
the paternal and maternal chromosomes, producing only two classes of gametes,
each one identical to one of the parental types. This system effectively links the
whole chromosome complement together as though there were only one super
chromosome pair. These are known in the literature as "Renner complexes." The
genic constitution of the two genomes is identical or only slightly different in
those species with both complexes
of the same type. In subsect. Oenothera,
these
SYSTEMATICBOTANYMONOGRAPHS
8
VOLUME 50
are 0. wolfii (AA), 0. villosa (AA), and 0. nutans (BB). The genic constitution
is distinctly different in those species with two types of complexes, as are found
in 0. biennis
gosity
(AC), and 0. parvifiora (BC). The heterozy
combinations.
of the homozygous
(AB), 0. oakesiana
through the prevention
is maintained
smaller ring configurations are known in some individuals of otherwise
PTH populations, but these are apparently associated with recent cases of hy
bridization with another entity and are not stable.
a plant with two genomes of the same genic constitution and
Complex homozygote:
Other
segmental end arrangement of the chromosomes, forming only bivalents (7 in the
I.
case of Oenothera, written 711) at meiotic metaphase
as "complex
Permanent structural heterozygote:
essentially the same phenomenon
heterozygote."
It is a term created by Renner
(genomes)
by Renner, Cleland,
and Stubbe.
Permanent
translocation
heterozygote
the genic heterozygos
used in the genetics literature
(PTH; see Holsinger
& Ellstrand
but with
the emphasis
same as "permanent structural heterozygote,"
manent
to describe
and subsequently
ity of the complexes
translocations
fixation of the chromosomal
genetically
1984):
the
on the per
locking
in the
clonal nature of these plants.
Plastome (Renner 1934): a term proposed
for the plastid DNA.
Structural heterozygote: condition inwhich the two chromosome sets of a species dif
the chromo
fer by one or more reciprocal translocations of the chromosomes,
somes thus forming circles or chains in meiotic metaphase I.
sets of a species
Structural homozygote:
condition in which the two chromosome
have no reciprocal
meiotic metaphase
Isogamous
(de Vries
thus forming bivalents
the chromosomes
translocations,
in
I.
1911): a complex
heterozygous
in which
species
plexes are transmitted through egg (a) and pollen (f3).
species
Heterogamous
(de Vries 1911): a complex heterozygous
in which
both com
one com
plex is transmitted through the egg (oc), the other through the pollen (13).
species
Halfheterogamous
(Renner 1918): the condition in a complex heterozygous
in which
both complexes
are transmitted through the egg (a), whereas
only one
(13), or the reverse situation with both
transmitted through the pollen and only one through the egg.
complex
is transmitted through the pollen
GENOME
The crossing
AND
analysis by Stubbe
PLASTOME
ANALYSIS
(1959, 1964) established
that there are three major
differentiatedgenomes inOenothera subsect.Oenothera,which were designatedA, B,
and C, and five basic plastid genomes, designated as plastomes I, II, III, IV, and V (Table
relationships worked out by
1; Figs. 1, 2). Figure 1 shows the detailed compatibility
Stubbe. The molecular studies by Gordon et al. (1981, 1982) confirm the distinctiveness
of the five basic plastome types. Detailed mapping of the two rDNA regions of the five
plastomes indicate that a deletion in plastome III and an insertion in plastome V relative
to plastome IV had occurred. Gordon et al. (1982) believed these changes were located in
the spacer region between the genes for the 16S and 23S rRNA.
The taxonomy detailed here delimits
nomically
homozygous
species,
five mostly
and eight PTH
outcrossing,
species
bivalent-forming,
based on the combination
ge
of
OENOTHERA
1997
genomes
(Table 1; Fig. 2). The properties of the PTH system serve to par
and plastomes
tition and fix variation
9
into perceptibly
differentiated true-breeding strains; i.e., essentially
clonal organisms. Many European taxonomists, without consideration of the patterns of
variation found within the indigenous populations of the group in North America, have
used morphological
features to delimit very large numbers of species. The units that we
recognize are comparable to those employed in the classification of the other subsections
of Oenothera sect. Oenothera and other parts of the genus, which we believe is a much
more useful way of formally reflecting the variation patterns inmorphology, genome, and
plastome of subsect. Oenothera. There are no limits to the number of individual patterns
of variation
that could be described within
the units that we have recognized as species,
in no way helps to un
derstand the situation in nature or to provide useful reference points for discussing signif
and excessive
icant evolutionary
reflects
means
of binomials
application
to the pattern of variation
units. We have devised
the knowledge
of the evolutionary
a comprehensive
taxonomic
system that both
history of the group and provides
a reliable
for identification
and for information synthesis and retrieval. Our approach aggre
into species delimited ac
gates the essentially clonal PTH populations (or microspecies)
to
of
the composition
their genomic complexes (A, B, or C), plastome type (I, II,
cording
characters.
III, IV, or V), and associated morphological
GENOMICPHENOTYPES
Each of the three fundamental
1964) in Oenothera
Figs.
1, 2). All
genomes
subsect. Oenothera
species
(A, B, and C) elucidated
exhibits certain phenotypic
by Stubbe
expressions
(1959,
(Table 2;
have one or a combination of two of these
of the species are given in Table 1.
of the subsection
The genomic combinations
There are two different kinds of PTH species within Oenothera subsect. Oenothera:
1) Species having both genomes of the same type (AA, BB, or CC). The genetic con
stitution of the two genomes, which is expressed phenotypically,
is often similar, and in
genomes.
some cases almost identical, as in 0. wolfii (AA) and 0. nutans (BB) (Wasmund & Stubbe
1990). In 0. villosa (AA) for example, the complexes may be nearly
identical, or they may differ by minor characters, such as length of floral tube, color of
1986; Wasmund
sepals, or density of inflorescence, but in all cases they clearly represent A genomes.
2) Species having two different genomes (AB, BA, AC, or BC). Oenothera biennis
(AB), 0. oakesiana (AC), and 0. parviflora (BC), which have arisen by hybridization,
show characters of the two genomes in their phenotypic expression. For example, pustu
late hairs, which in general result from genes associated with the A genotype, occur usu
ally only in 0. biennis (AB) and 0. oakesiana (AC); strigillose pubescence, which is typ
ical for a number of 0. elata strains as well as 0. longissima and 0. jamesii, also occurs
in 0. biennis and 0. oakesiana. In 0. biennis, however, this character can be suppressed
by the dominance
illose pubescence,
of genes associated with the B complex, which does not exhibit strig
at least in the region of the inflorescence. Subterminal free sepal tips
and recurved inflorescence
tip, which
are characters associated with genes
located in the
C complex, occur in0. oakesiana (AC),0. parviflora (BC), and0. argillicola (CC).Sub
glabrous leaf surfaces are typical in strainsof 0. grandiflora (BB) and in 0. argillicola
(CC), so that forms of 0. biennis (AB) and 0. parviflora (BC) sometimes also express this
character. Pale and deciduous bracts, characters associated with genes located on the B
genome, from time to timeoccur in0. biennis and0. parviflora.
Table
1. Summary
SI = self-incompatible;
PTH
species
usually
of attributes
of taxa of Oenothera
SC = self-compatible;
prove
not
to be stable
sect. Oenothera
A = autogamous;
and indicate
MO
spontaneous
subsect.
= modally
hybridization
The
Oenothera.
outcrossing.
following
Configurations
in an otherwise
symbol
including
permanent
translo
here for completeness.
Summaryof
Meiotic Chromosome
Taxon
Breeding
1. Oenothera elata
subsp. elata
subsp.hirsutissima
System
SC,MO
SC,MO
Configurations
subsp.hookeri
2. Oenothera
jamesii
7ii; 04+511; (06+411;08+31l;
0)6+OD4+2u;
Genome
No
AA
No
AA
010+211
SC,MO
71I;0)4+5I1;(36+4II
No
AA
SC, MO
71,; 04+511;
No
AA
No
AA
Yes
AA
0310+211;
3. Oenothera longissima
Heterozygote
711
204+311;
Permanent
Translocation
SC,MO
0)6+4II; 08+31,;
(014
71I;04+511; G)6+411;(08+311;
204+311
4. Oenotherawolfii
SC, A
0)14
5. Oenothera
SC, A
villosa
014;
012+111
Yes
AA
Yes
AA
subsp. villosa
subsp. strigosa
6. Oenothera
stucchii
7. Oenothera grandiflora
SC, A
(0 14
SC, A
012+11l;
SC (SI),MO
AA*
Yes
014
7II;04+511;06+41I;
No
BB
Yes
BB
204+311; 08+3I1;
014
012+11I;
036+(04+211;
010+21j;
8. Oenothera
nutans
SC, A
014;
9. Oenothera
biennis
SC, A
0)14; 012+11I;
10. Oenothera
glazioviana
SC, MO
11. Oenothera
argillicola
SC, MO
012+111
012+11I
711;
04+510;
08+311;
12. Oenothera
oakesiana
SC, A
13. Oenothera
parviflora
SC, A
*The genome
of 0.
stucchii
appears
to have
Yes
010+21j;
014;
(D6+4,,; 204+311;
and B
Yes
AB
No
CC
Yes
AC
Yes
BC
(D10+2Ij
0D12+1,1;
0 10+04
(014
some B genome
AB
characteristics
based
on crossing
experiments
described
12
SYSTEMATICBOTANYMONOGRAPHS
f ne| I
I
Gcnotypc
|
__ _
,__
II
|
III
_ _.
.
VOLUME 50
| ~~~IV |
l
V
...
fl@
AB
l
BBgff
0
BC00
Icc
**|
I+I+
1
AC *~0@4
*
= nornal green
*
= green to grayish
green
= yellow-green
(lutescent)
()
oD = periodically
lutescent
) = yellow-green to
yellow
o
E
+
= white, with inhibitionof
growth andger
on
= lethal, but white
as an exception
if occuning
= slightlyyellowing
= periodicallypale
(diversivirescent)
= periodicallypale
(virescent)
- white or yellow
sect.
and plastome
of major diploid genomes
relationships
types in Oenothera
There are three primary genome types, A, B, and C. These can be combined with
The symbols in each of
the five basic plastome
types, 1,H, II,,
IV, and V, with various degrees of compatibility.
en
indicate the development
and function of the plastids in a particular genomic
the 30 possible combinations
of plastid pigments. When more than one symbol appears in a
vironment,
including synthesis and degradation
FIG.
Oenothera
1. Compatibility
subsect. Oenothera.
cell, different
Only
plastome
the most
IV is compatible
plesiomorphic
were
observed
resulting from genetic differences
and therefore
all of the genomic combinations
from Stubbe (1959, 1964).
type. Redrawn
kinds of interactions
with
forms.
among A-genome
to be
has been considered
OENOTHERA
1997
HG2 Vabendno-vabe
symbols Vibetente
setubbenohr
us(1964).a
omiaton
o mjo
adnnvalcobntosomargemendplastome
h
pcesaealiclddi
gnoe
13
ndplstm
rereen
rltvagesinssoechpstome
the
types
h fgr,adinsm
type in thvaaoeohr
type Redanotfrom
ae
tedsga
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
14
Table
2. Morphological
characteristics
associated
with
the genomes
A, B, and C.
Character
Genome A
Genome B
Genome C
Stem
Erect
Erect
Suberect to oblique
Tip of inflorescence
Straight,
not curved
not curved
Straight,
but the ultimate
Recurved,
tip incurved
again
(S
shaped)
Shape
of rosette
leaves
oblanceolate
Narrowly
Oblanceolate
Very narrowly
oblanceo
late to linear
Bracts
Same
of leaves
Constitution
Leaf color
color
as other
Often
Same
pale and
color as other
leaves,persistent
deciduous
leaves,persistent
Somewhat
Thin,
Somewhat
coriaceous
"hygromorphic"
"teromorphic"
Dark green
Grayish green
Green
Lanceolate
Narrowly
Sepal tips
Erect, terminal,"thick"
Erect, terminal,tiny
Divergent, subterminal,
"thick"
Stem pubescence
Strigillose or villous,
glandularhairs present
or absent
Strigillose to glabrous,
glandularhairs usually
present
Short-strigillose to
glabrous
Scattered
villous
Glandular
puberulent
glabrous,
glandular
glabrous,
rarely scattered
Outline
of mature
(excluding
ovary
buds
Lanceolate
lanceolate
and
floral tube)
Ovary,
floral tube, and
sepal pubescence
or villous,
Strigillose
hairs present
glandular
to
to
or absent
hairs usually present
villous
Pustulatehairs
Absent or present,when
present,pustules red
Rarely present,pustules
translucent,not red
Rarely present,pustules
red
Capsule
Gradually tapering
Gradually tapering
Attenuate towardthe apex
toward
Inflorescence
the apex
Simple
toward
the apex
Often with secondary
spikes just below
main
Plastome
I
Simple
the
one
III
V
CYTOLOGY
New cytological observations are provided in this paper for all 16 taxa (562 individ
of chromosome number, meiotic configurations,
uals) from 344 localities. Determination
and compatibility by repeated self-pollination was made on all strains that have been
brought into cultivation for this project. Our results are completely consistent with the co
pious cytogenetic literature for this group of species. All determinations were diploid, n =
7, with no polyploidy
or aneuploidy
detected. The association
of chromosome
configura
OENOTHERA
1997
tions with particular collections
can be determined
15
from the section
listing these under
each species.
as is the
All species of Oenothera
subsect. Oenothera have 2n = 14 chromosomes,
case throughout the genus, with only a few exceptions. A summary of the meiotic config
urations found, both from our studies and in the literature, are presented in Table 1. The
structural homozygous
species, which are large-flowered and predominantly
outcross,
I.Within populations of homozygous
usually form 7II inmeiotic metaphase
species, con
figurations range from small rings up to 08, indicating that the chromosomal end arrange
ments are not uniform within populations. The strains of 0. grandifiora from Alabama
collected by E. Steiner in 1983 and analyzed inDusseldorf
(Steiner & Stubbe 1984, 1986;
Schumacher
1987; Schumacher & Steiner 1993), contained plants exhibiting 0 10 and 211;
is believed to represent
012 and 1II; and 014. The latter configuration
that are not completely pure 0. grandiflora, but rather ones influenced by hy
and 04;
0)10
plants
bridization with
the sympatric
the A genome,
with
as well
0. biennis, which
The permanent structural heterozygous
0314 chromosomes
contributed
characteristics
associated
as an altered chromosomal
atmeiotic metaphase
configuration.
species of Oenothera subsect. Oenothera
I, themost
specialized
form
situation in the subsection.
a stable configuration of 08 and 06 occurs, such as in the European forms of
0. biennis. This configuration has not been found inNorth American strains of 0. biennis.
Another stable configuration, 0)12 and III, is typical for 0. glazioviana worldwide. Other
Sometimes
configurations, e.g., 08, 04 and 1II; (I10 and 2II; or (010 and 04, usually prove to be un
stable and indicate spontaneous hybridization in an otherwise PTH population.
In spite of Cleland's thousands of diakinesis examinations and determinations of end
arrangements for hundreds of strains, the examinations performed at Dusseldorf presented
here have considerably
widened
the spectrum of known
chromosomal
configurations
in
populations
of Oenothera subsect. Oenothera (Table 1), especially in structural homozy
gous species such as 0. argillicola, 0. grandiflora, and 0. elata, and give new insights
into the population
discussed
structure of these species. The specific details of these situations
are
under each of the species.
THE IMPORTANCE
OF HYBRIDIZATION
The essentially clonal PTH populations (or microspecies)
are, as a system, exceed
ingly important in the evolution of this subsection. The properties of these PTH organisms
have resulted in a great amount of variability and recent evolution in the complex, in large
part due to hybridization and segregation of new, essentially clonal, phenotypes. Cleland
(1972, p. 228) pointed out that the high degree of autogamy found in the PTH species se
limited the extent and frequency of hybridization, but at the same time the proper
ties of the PTH system fix any result of hybridization. Therefore, hybridization, although
verely
not frequent, has played amajor role in the evolution of the group. We have considered the
majority of this new variation as intraspecific within four of the PTH species (0. biennis,
0. oakesiana, 0. parvifiora, and 0. villosa). Hybrids occur between many of the species
(19 known combinations, Table 3), especially the PTH species, and they are treated as such
when the hybridization appears to represent a local phenomenon and when the phenotype
is intermediate. They are grouped with the taxon they most closely resemble when the hy
brids or their derivatives represent amore widespread phenotype, such as the intermediates
between the subspecies of 0. villosa or between 0. oakesiana and 0. parviflora.
16
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
Table 3. Known naturallyoccurringhybrids.
Viable hybridphenotype
genome/plastome
Combination
0. argillicola x 0. parviflora
CC-V/IV
0. biennis x 0. glazioviana
AB-II/III
0. biennis x 0. grandiflora
BB-III orAB-IJI
0. biennis x 0. jamesii
AA-I
0. biennis x 0. nutans
AB-IJI or BB-III*
0. biennis x 0. oakesiana
AC-IV
0. biennis x 0. parviflora
AC-IV
0. biennis x 0. villosa subsp. strigosa
AA-II
x 0. villosa
0. biennis
subsp. villosa
AA-1/II,
0. elata subsp.hirsutissimax 0. longissima
0. elata
x 0. villosa
subsp. hirsutissima
subsp. strigosa
AB
AA-I
AA-I*
0. glazioviana x 0. villosa subsp. strigosa
AA-I/III
0. glazioviana x 0. villosa subsp.villosa
AA-I/Ill
0. glazioviana x 0. wolfii
x 0. villosa
0. jamesii
0. nutans
AB-III or II
subsp. villosa
AA-I
x 0. parviflora
BB-IV
0. oakesiana
x 0. parviflora
0. parvifiora
x 0. villosa
0. villosa
subsp. villosa
I/II
AC-IV
subsp. villosa
x 0. villosa
and BC-IV
subsp. strigosa
AB-I/IV
AA-I
*Not confirmed
Several of the species of subsect. Oenothera were introduced to Europe at least three
centuries ago, and as a result of hybridization, numerous new phenotypes have originated.
Many of these almost exclusively European entities have been given formal names (Table
4). Renner (1942) recognized at least 18 PTH species in Europe. Many others have been
described as species since then (e.g., Hudziok 1964, 1968; Rostan'ski 1985). This ap
proach of giving formal names to every different true-breeding phenotype discovered also
has been used in North America (e.g., Gates 1936). If this approach were taken to its ex
treme, hundreds or perhaps thousands of specific names would result. The amount of
to begin to identify the resulting entities with anything but rudimentary ac
be extraordinary.
Some of the new European hybrid forms have proven to be stable and true-breeding
study needed
curacy would
1997
OENOTHERA
(Table 4), although none of these hybrids
17
[0. biennis x 0. glazioviana],
0. xalbipercurva
[0. biennis x 0. oakesiana], 0. xhoelscheri
[0. biennis x 0. villosa
subsp. villosa]) have become widely established. Rather, they arise anew where the
(e.g., 0. xfallax
parental species grow together. Some other "hybrids" are formed between
elements within
what we consider
the limits of a single taxonomic species, especially 0. biennis (e.g., 0.
rubricaulis or 0. suaveolens). Because of the unique properties of PTH, once hybrid prog
eny are formed they can persist and reproduce themselves, at least locally. This seems to
be the case in a number of the relatively better-studied European hybrids. For example,
what has been known as 0. xfallax has become established in scattered localities in Eu
rope in places where
near Dusseldorf,
ulations because
the parents are sympatric.
this hybrid is common.
In certain places,
vigorous
combination
it breeds true. It has a named genomic
from 0. glazioviana)
and rubens (B genome from 0. biennis),
chromosomes and one bivalent, like 0. glazioviana.
Concerning
North America
hybridization
such as along the Rhine
It has established
and stable local pop
of velans
(A genome
and forms a ring of 12
and the new combinations
that can arise, the situation in
to that in Europe, but even more complex. The
number of different phenotypes found inmost of the species within their indigenous range
is considerably greater than that observed in European populations (Cleland 1972, p. 227).
is, of course, comparable
the European species concept in North America, especially in areas of recent sym
patric contact (e.g., Oregon, Washington, British Columbia), would lead to a chaotic situ
ation in which numerous microspecies
could be described. Such a proliferation of formal
names, however, would not improve our understanding of these plants, their origins, or
Using
their variation.
ECOLOGYAND GEOGRAPHY
All species of Oenothera subsect. Oenothera,
like most species of the genus, occur in
primarily or secondarily open habitats, including old fields and roadsides, and often, es
pecially in arid regions such as the southwestern United States, in at least seasonally wet
sites, such as stream sides, arroyos, and dunes. They grow from sea level along the At
lantic (0. oakesiana) and Pacific coasts (0. elata subsp. hookeri, 0. wolfii) to elevations
over 3000 m in the Rocky Mountains
(0. villosa subsp. strigosa).
Recent studies (Gross & Werner 1982; Gross 1985) have shown that seeds of 0. bi
ennis require light to germinate, and that seedlings establish only on bare soil. Provided
that these requirements are met, 0. biennis can grow on a wide range of soil types. An
other study (Gross & Kromer 1986) indicates that seed weight has a transitory effect on
seedling and rosette diameter, but that soil type has an increasing effect after four weeks
on growth rate, final plant size, and reproductive output. These ecological traits are among
the specialized features of subsect. Oenothera, which include robust biennial or short
lived perennial habit with stems up to 3 cm in diameter basally and seed production many
times higher than that of other sections of Oenothera.
Part of the specialized growth form of all species of subsect. Oenothera
is that they
are facultative biennials or rarely winter annuals (0. jamesii; Munz 1965). It has been
shown that under unfavorable conditions, such as low levels of water and/or nutrients, 0.
glazioviana may stay in the rosette stage for several years, depending on rosette size
(Kachi & Hirose 1983). In cultivation, all species have the useful quality that they can be
grown as annuals when
sown in January or February
in the greenhouse
and kept there until
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
18
Table 4. The 79 scientific
names
in Oenothera
described
sect. Oenothera
subsect. Oenothera
based
on
wild-collected types frompopulations inEurope (excludingexperimentalhybrids and experimentalstrainsof
H. de Vries;
also excluded
are the numerous
names
applied
to European
populations
that have never been
validly published).
PresentDisposition
Name
0. biennis
0. biennisL. var. ieptomeresBartlett
0. brevispicataHudziok
0. cambrica
Rostaiiski
0. cambrica
Rostadski
0. carinthiaca
var. impunctata
Rostaiiski
Rostanski
0. chicaginensis
de Vries
ex Renner & Cleland
var. bartlettii
0. chicaginensis
de Vries
ex Renner & Cleland
var. minutiflora
Rostatiski& Jehlik
0. compacta
Hudziok
0. editicaulisHidziok
0. ersteinensisLinder& Jean
0. flaeminginaHudziok
0. inconspectaHudziok
0. jueterbogensisHudziok
0. macrosperma (Hudziok)Hudziok
0. mediomarchicaHudziok
0. muricata
L. var. latifolia Ascherson
0. nissensis Rostaniski
0. obscurifoliaHudziok
0. octolineataHudziok
0. paradoxa
Hudziok
0. punctulata
Rostan'ski
& Gttte
0. pyramidifloraHudziok
0. rostanskiiJehlik
0. rubricaulisKlebahn
0.
rubricaulis
0.
rubrica-ulis Klebahn
0.
sesitensis
0.
suaveolens
Klebahn
var. dentifolia
Jehlik & Rostatiski
var. longistylis
Gutte & Rostatiski
Soldano
Persoon
var. latipetala
Soklano
0. tacikiiRostatiski
0. bienmisor possible hybrid
0. marinellae Soldano
0. pedemontana
Soldano
0. pellegrinii Soldano
0. glazioviana
0. bipartitaLutz
0.
coronifera
(Borbas) Borbas
0. erythrosepala
(Borbas) Borbas
0. multiflora
0. multiflora
0. oakesiana
Renner
0. eryt-hrosepala
var. elliptica
Gates
0.
rubrinervoides
0.
rubritincta
0.
tardiflora Gates
Gates
Gates
0. ammophilaFocke
0. gerinanica
var. azorica
Gates
Boedijn
Gates
Rostaniski
Soldano
OENOTHERA
1997
19
Table 4. cont.
PresentDisposition
0. parviflora
Name
0. lipsiensisRostadiski& Gutte
0. pachycarpa
ex Rudloff
Renner
0. rubricuspisRenner ex Rostanski
0.
silesiaca
Renner
0. turoviensisRostariski
0. stucchiiSoldano
0. stucchii
0. villosa
subsp. villosa
0. bauri Boedijn
0. canovertexHudziok
0. depressa
E. Greene
f. angustifolia
0. depressa
E. Greene
f. latibracteata
0. hungarica
0.
RostaAski
Rostaniski
(Borbas) Borbas
renneri H. Scholz
0. velutinifoliaHudziok
0. biennis
x 0. glazioviana
0. xadriatica
Soldano
0. xbritannica
Rostanski
0. xcoloratissimaHudziok
0. xconferta
Renner & Hirmer
0. xfallacoides
Soldano
0. xfallax
Renner
0. xfallax
Renner
0. xoehlkersii
0. biennis
x 0. oakesiana
f. rubrinervis
Rostaiski
ex Rostaiski
Kappus
0. xalbipercurva
Renner
ex Hudziok
0. xalbipercurva
Renner
ex Hudziok
0. xbraunii
0. xclavifera
Renner
Hudziok
0. xheiniana
Teyber
0. xindivisa
Hudziok
0. xissleri
var. impunctata
Doll
Renner
var. silesiacoides
Rostanski
& Jehlik
0. xpseudocernuaHudziok
0. biennis x 0. parviflora
0. biennis
x 0. villosa
subsp.villosa
0. nissensis Rostaniskinothovar.fiedleriGutte & Rostatiski
0. xpseudochicaginensisRostan'ski
0. xdrawertii
0. xwienii
0. glazioviana
x 0. villosa
Renner
ex Rostanski
0. xpolgari Rostatiski
Renner
0. xpurpurans
ex Rostatiski
Borbas
subsp.villosa
0. parviflora
x 0. villosa
subsp.villosa
0. xslovaca
Jehlik & Rostaniski
ex Hudziok
SYSTEMATICBOTANYMONOGRAPHS
20
VOLUME 50
the rosettes are large enough for planting in the field in April. In contrast, plants can be
held in the rosette stage for several years when planted in pots and held relatively dry and
with only a very little fertilizer.
The natural distribution of subsect. Oenothera extends from southern Canada nearly
throughout the United States, southward and increasingly sporadically through Mexico
to
Costa Rica and Panama. Some species, especially those having PTH, have become ex
in temperate and subtropical regions. The most widely
tensively naturalized worldwide
naturalized species is 0. biennis; however, 0. oakesiana, 0. parviflora, and 0. villosa
subsp. villosa
also have moderately
large naturalized
distributions.
Oenothera
villosa
in Eurasia and South Africa, whereas 0. oakesiana and
0. parvifiora have become naturalized primarily in Europe with sporadic occurrences
elsewhere. Oenothera
in South
jamesii has a more sporadic naturalized distribution
subsp. villosa occurs commonly
Japan, and the Canary
Africa,
Islands although it is well established locally. Two of the
that we recognize here at the level of species, 0. glazioviana and 0. stucchii, have
arisen during the past few hundred years outside of the natural range of the subsection,
and of the two only 0. glazioviana,
the earlier to arise, has an extensive distribution on all
entities
continents
except Antarctica.
range of the genus Oenothera
is entirely confined to North and South
of 0. biennis in Europe and Asia has led Rostan'ski (e.g.,
to hypothesize
that species he treats as distinct, such as 0. rubricaulis (included
The native
The wide distribution
America.
1975)
here in 0. biennis),
ennis a Eurasian
originated
in the Old World, whereas
species without
further comment,
number of European botanists. Although
in Europe,
there is no evidence
Jehllfk (1989, p. 260) calls 0. bi
suggesting
that this idea is held by a
unique minor phenotypes
have indeed originated
for an occurrence
of 0. biennis in the Old World before
1492. There are no fossils known, nor any hint in the literature or art before 1492. We be
lieve that such a conspicuous plant as 0. biennis would certainly have been depicted by
one of the great artists of the late Middle Ages (or the period immediately after), such as
Albrecht Durer (1471-1528),
or would have been treated in one of the early herbal books.
Although
Jehllk (1989, p. 260) thought that 0. glazioviana
may be of North Ameri
can-Eurasian
origin, a glance at the distribution map of 0. glazioviana
(Fig. 35) shows it
to have the sort of distribution that is characteristic of weedy naturalized herbs. Also, the
dates of first collection make
it virtually certain that it did not exist before the middle of
the 19th century. The earliest collection known to us is from 1868 from Brazil; the next,
1869, cultivated in Germany; Great Britain in 1871; Uruguay in 1874; France, Austria,
in 1876; Poland in 1879; Argentina in 1880; Switzerland
in 1883; and Japan and the United States in 1884.
and the Netherlands
Canada
in 1882;
BREEDING SYSTEMSAND POLLINATION
All species of Oenothera subsect. Oenothera reproduce exclusively
sexually. There
is no vegetative propagation nor any other kind of asexual reproduction like apomixis.
Large flowers that are open-pollinated
represent a plesiomorphic character in the subsec
tion, as for the genus. Also, as is typical for most of the other sections of Oenothera,
the
evolution
of autogamy from large-flowered progenitors has occurred repeatedly within
subsect. Oenothera. The breeding systems of all of the species are summarized in Table 1.
There is usually some degree of outcrossing whenever
the flower is open, and all mem
bers of subsect. Oenothera are visited by hawkmoths when they are flowering, although
1997
OENOTHERA
studies have been conducted
detailed
21
only for 0. elata
(Gregory 1963, 1964). Because
there are pollinators present, hybridization between
sympatric species (or phenotypic forms) is relatively common.
is genetic self-incompatibil
Another plesiomorphic
feature in the genus Oenothera
not
have any known self-incom
ity. Only three of the fourteen sections of the genus do
open pollination
occurs whenever
[0. sect. Contortae W. L. Wagner, 0. sect. Gauropsis (Torr. & Frem.)
and 0. sect. Hartmannia
(Spach) Endl.]. All others have at least some in
dividuals or species that exhibit genetic self-incompatibility. Within sect. Oenothera, all
are self-compatible,
and until re
members of both subsections Munzia and Nutantigemma
of
cently, subsect. Oenothera was believed to be entirely self-compatible. Determination
patible
individuals
W. L. Wagner,
compatibility
by repeated self-pollination was made on all strains that have been brought
for this project. All were self-compatible
except for several strains of 0.
from populations in Alabama that have retained self-incompatibility
(Stubbe
into cultivation
grandiflora
& Raven 1979b; Steiner & Stubbe 1984, 1986; Schumacher & Steiner 1993). This species
is far more diverse than previously
thought. Some populations seem to be entirely or
mostly composed of self-incompatible
individuals, whereas others consist of self-com
in Oenothera; 0. primiveris
patible plants. This is an extremely uncommon phenomenon
A. Gray (Wagner unpubl.) is the only other species known to occur inmixed populations
and self-compatible
of self-incompatible
individuals. The discovery of self-incompatibil
ity in subsect. Oenothera was important, because Steiner (1956, 1957, 1961, 1964) sug
gested that Si-alleles were still present in the group in an unbalanced condition and acted
as male gametophytic
lethals in the formation of a PTH form.
A detailed study of pollination biology of 0. elata (as 0. hookeri) was conducted in
southern California (Gregory 1963, 1964). No other species of subsect. Oenothera have
been the subject of detailed studies of pollination. Eight colonies of 0. elata subsp. hir
sutissima were studied by Gregory over a period of time, often a full season. As is typi
cal for hawkmoth-pollinated
species of Oenothera,
the flowers open near sunset during a
brief span of a few minutes. Visitors included Eumorpha achemon, Hyles lineata, Man
duca quinquemaculata, M. sexta, Sphinx chersis, and S. perelegans asellus. The diversity
and abundance of the visitors varied over time, but overall the most common ones were:
Eumorpha
achemon, Hyles lineata, andManduca sexta. Gregory also noted three species
of bees as common visitors: Apis mellifera, Xylocopa brazilianorum varipunctata, and X.
tabaniformis orpifex. All of these insects were pollinators to some degree. Oenothera
elata subsp. hirsutissima
bees gather residual pollen
1964; Linsley et al. 1973).
is pollinated
almost entirely by the hawkmoth
from the flowers and effect some pollination
visitors, but the
(Gregory 1963,
that because 0. elata is self-compatible,
Gregory mentioned
a substantial amount of
autogamy occurs. This happens in two ways: 1) the insects often visit more than one
flower per individual plant, and 2) some pollen is transferred from anther to stigma in a
single flower. He estimated that over 50% of the pollination is by selfing.
A recent study (Kawano et al. 1995) showed that linalool, a monoterpene,
is the pri
mary constituent of the floral volatiles of 0. glazioviana and 0. biennis (based on popu
lations in Japan). Other substances were present but not identified. Both of these species
strong UV-absorbant
spots near the center of the corolla. Kawano and collabo
rators also found that local Japanese hawkmoths visited and effected pollination, includ
exhibited
ing Agrius
convolvulii, Deilephila
elphenor lewisii, and Theretra japonica; however, 0.
is primarily self-pollinating
and therefore cross-pollination
by moths is of lesser
importance. They concluded that the UV patterns coupled with the floral volatiles served
biennis
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
22
to the flowers. One of the other compounds,
to attract all but Agrius
presumably
responsible
as yet unidentified,
is
for the attraction of Agrius.
ORIGINS
The early diversification
inMadrean
vegetation
of Onagraceae
tribe Onagreae
of western North America
typical of this pattern. Twelve
appears to have taken place
1978). Oenothera is
(Raven & Axelrod
of the 14 sections
of Oenothera
are represented
in the
southern half of Texas south to northern Mexico, mostly associated with Madrean wood
land or closely related derivative vegetation types (Tobe et al. 1987). Arizona and New
are nearly as diverse with representatives of 10 sections each; however, diversity
the genus very quickly decreases outside the area including Texas to Arizona and
northern Mexico. The high sectional diversity in this region and the occurrence in Texas
and Mexico of species with greater numbers of plesiomorphic characters, such as self-in
Mexico
within
compatibility, strongly indicate that Oenothera originated inMadrean vegetation in this
region or adjacent regions formerly occupied by vegetation similar to that of this area,
probably by the early Neogene.
The genus has subsequently
ing from low-elevation
diversified
hot deserts
into a wide variety of habitats,
greatly
rang
to montane
temperate and subtropical forests, sub
eastern
deciduous
conifer
and
alpine
forests. Species inhabit open, sandy, rocky,
forests,
or clay sites to occasionally wet soils at stream or wetland margins. The geographical
range of Oenothera
America.
includes most
of North America
Judging from the patterns of distribution,
pears to have occurred
areas. Oenothera
in conjunction
subsect. Oenothera
with
as well
as all of temperate South
the origin of the various
sections ap
shifts into new ecological
is representative
or geographical
of this overall pattern in the genus,
with
the shift being primarily ecological rather than geographic. The principal shift in this
group appears to have been development of a robust upright habit coupled with increased
flowering and seed production.
The species grouped here as Oenothera
nized
subsect. Oenothera
have always been recog
as a monophyletic
group. Among
the principal features indicating that subsect.
is a monophyletic
Oenothera
group are: 1) seeds prismatic and angled; 2) seeds with
mesotesta nearly crushed (Tobe et al. 1987) [this character apparently was also indepen
dently derived in the common ancestor of the clade comprising Oenothera sects. Lavauxia
(Spach) Endl., Gauropsis
pleurum
(Torr. & Frem.) W. L. Wagner,
(Spach) Endl., and Hartmannia
perennial habit with
manifold
over
Kneiffia (Spach) Endl., Xylo
(Spach) Endl.]; 3) robust biennial or short-lived
stems up to 3 cm in diameter basally; 4) seed production
that of other members
of Oenothera
increased
and
than a hundred capsules per plant). Oenothera subsect. Oenothera appears
on morphological
grounds to be most closely related to subsect. Emersonia. This hypoth
esis is supported by extensive experimental
hybridization
studies (Stubbe & Raven
dozens
(300-400
seeds per capsule
tomore
1979a). These crossing analyses suggest that 0. maysillesii Munz most closely resembles
the common ancestor of the section, and that subsect. Oenothera is the group most closely
related to subsect. Emersonia, with subsect. Munzia less closely related to either of these
subsections, but presumably directly derived from sect. Emersonia. Oenothera
subsec
tions Raimannia
phylogenetic
Emersonia)
and Nutantigemma
are apparently more highly derived, but on the same
branch as subsect. Oenothera. The seeds of 0. organensis Munz (subsect.
are similar to, but not identical with, those of subsect. Oenothera (Dietrich et
OENOTHERA
1997
23
larger, have a thicker endotesta, and have a thin, rather than
(Tobe et al. 1987). These features suggest that subsect. Oenothera may
related to 0. organensis.
al. 1985). They are somewhat
crushed, mesotesta
be most
closely
Emersonia
subsect.
Oenothera
(Dietrich
are self-incompatible.
in the genus (Raven 1979). Oenothera subsect. Emersonia, although possibly
as presently delimited, appears to occupy a basal position within sect.
paraphyletic
Oenothera (Dietrich et al. 1985). The resolution of the overall relationships in the genus
species,
three of which
et al. 1985) consists of four perennial
Both of these features are considered ple
siomorphic
is currently under investigation
by both morphological
and molecular
phylogenetic
analy
ses (Wagner unpubl.; Sytsma et al. unpubl.).
that appears to be
(Herrmann et al. 1983; Hachtel et al. 1991;
Sytsma et al. 1993). The inversion is approximately 45kb and appears to be absent from
all of the species tested from three of the four other subsections of sect. Oenothera
Molecular
studies of chloroplast DNA
to Oenothera
restricted
have revealed an inversion
subsect. Oenothera
(Hachtel et al. 1991).
There are three basic
lineages within
subsect. Oenothera
Oenothera
relating to
we
genome and plastome composition. Below
discuss the probable origins of the species
that have both their genomes of the same type (AA, BB, CC); the more complex origins
of those PTH species with mixed
that arose via hybridization (0.
genomic composition
biennis, 0. oakesiana, and 0. parvifiora); and the possible origins of the two species, 0.
glazioviana and 0. stucchii, that have originated recently in Europe, outside of the natural
distribution
of the subsection.
AA GENOME SPECIES
Five species have AA genomes and all of them have plastome I (Fig. 3). Oenothera
and 0. longissima have retained several plesiomorphic
characters, in
cluding large mostly outcrossed flowers, and formation of bivalents or variably sized rings
of chromosomes during meiosis. Oenothera elata is widely distributed in western North
elata, 0. jamesii,
America
and south to Panama, whereas
uted inwestern North America.
rectly and independently
that the latter two may have been derived di
from 0. elata (Munz 1949; Raven et al. 1979). Like a number of
other species of Onagraceae
jamesii and 0.
longissima
by hawkmoths with
the other two species are more narrowly distrib
It is possible
in the southwestern United
appear to have diverged
longer proboscides,
States and northern Mexico,
in response
to specialized
such as species of Manduca
0.
pollination
(Raven et al. 1979).
The other two AA-I species are both PIH species. Oenothera woffii, formerly treated
as 0. hookeri subsp. wolfii (Munz 1949), is a coastal endemic in northern California and ad
jacent Oregon. Oenothera wolfii appears to have evolved from populations of 0. elata
subsp. hookeri to the south in coastal California in the recent past by the accumulation of
reciprocal translocations and the acquisition of balanced
lethals (Wasmund & Stubbe 1986).
villosa, occurring widely in North America, appears to have been derived
from populations of 0. elata subsp. hirsutissima. Morphology
suggests that the two sub
species of 0. villosa were independently derived from different ancestral populations, fol
Oenothera
lowed by extensive, apparently secondary, intergradation between them. Oenothera vil
losa subsp. villosa is very similar in pubescence type and pattern and in other vegetative
features to populations at the southeastern periphery of the distribution of 0. elata subsp.
hirsutissima
in Texas, Kansas, and eastern New Mexico.
losa arose from one of them, perhaps more than once.
Presumably
0. villosa
subsp. vil
24
SYSTEMATICBOTANYMONOGRAPHS
. N.
.
. . . ->
. .
|
.
K
.
_
.
/
.,
{
* . _ | _ _.
. *-- I_
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III
__ _
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w
_
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w
_r|
.
_
,
.
.
w
.
/
.
s
.
.
.
.
..
.
t
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_b
.
.
_
.
.
.
I
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.
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s
,
1_
0
| |I
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li8
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VOLUME 50
|
. .
| |
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!
.
_S_
,^_
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t_
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1E
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\
_J | l
_-:s== \ S S | _'- - - |
.,_ S_ - . s . __ *s \ I | 111|_
I
L ....
_
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l
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-|
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1-I -i _ *1
s_
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/
s
_\
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hypothetiods
\
| zoestor / |
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Nplastome
IIJ
__ . . _ l It
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//
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' I
- _:
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..._
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i_
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,
_|_
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SS
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FIG. 3. Schematic diagram of the origin of the genomically homozygous species of Oenothera sect.
Oenothera subsect.Oenothera showing geographical relationshipsinNorth America, and thegenome-plastome
combinationsand theirpresumed relationshipto each other (arrows).
*
,
.
w
_
.
OENOTHERA
1997
Based on morphological
25
especially pubescence patterns, the populations
subsp. strigosa probably arose on several independent
occasions from 0. elata subsp. hirsutissima to the north and west of those giving rise to
0. villosa subsp. villosa. Evidence for this is based on the fact that populations of 0. elata
subsp. hirsutissima occurring in a particular geographical area often closely resemble the
similarities,
that we group together as 0. villosa
this pattern
subsp. strigosa populations in the same area. Alternatively,
could have resulted from secondary contact followed by hybridization between 0. elata
subsp. hirsutissima and 0. villosa subsp. strigosa. It also is possible that 0. villosa subsp.
derived 0. villosa
strigosa may have arisen only once from 0. elata subsp. hirsutissima, and that the two
taxa subsequently evolved several characters in parallel, or converged via a pattern of
local hybridization events. Current information does not allow discrimination between
thesehypotheses.
The populations grouped by Munz as 0. strigosa subsp. cheradophila also probably
arose independently from other populations of 0. elata subsp. hirsutissima in the Pacific
Northwest. The form of 0. elata subsp. hirsutissima with densely appressed pubescence
(referred by Munz to 0. hookeri subsp. ornata) grows adjacent to the cheradophila pop
ulations and is likely to have given rise to the cheradophila phenotype. Because all of the
here assigned to 0. villosa subsp. strigosa intergrade to a considerable extent
and all presumably arose from populations of 0. elata subsp. hirsutissima, they are best
phenotypes
grouped
in one taxon.
BB GENOME SPECIES
The
genomes
second
consists of two species of eastern North America with BB
III (Fig. 3). Oenothera grandiflora has a scattered and evidently
in the southeastern United States. Self-incompatibility,
retained in
lineage
and plastome
relictual distribution
some populations of 0. grandiflora, does not occur elsewhere in the subsection (Stubbe
& Raven 1979b; Steiner & Stubbe 1984, 1986; Schumacher & Steiner 1993). Although
less specialized than 0. argillicola, 0. grandiflora presumably evolved early in the diver
sification of the subsection as ancestral populations migrated to the southeastern United
States from further west and south.
The second species of this lineage, presumably
derived from 0. grandiflora, is 0. nu
is a PTH species known in the literature as Biennis-II,
0. biennis subsp. aus
or
austromontana.
It
tromontana,
0.
occurs throughout the eastern United States, espe
tans, which
cially
in the Appalachian
Mountains
Cleland (1958, 1972) suggested
the two races of 0. biennis, Biennis-I
and surrounding areas.
that 0. nutans originated via hybridization between
(BA) inwhich the B genome is transmitted through
the a (egg) complex, and Biennis-I1 (AB) in which the B genome is transmitted through
the f3 (pollen) complex. Biennis-I also has plastome III. By means of the sort of hy
bridization event Cleland suggested, an entity with BB genome and plastome III would be
created, having many grandiflora-like
Wasmund
phenotypically
characteristics.
(1984, 1990), by contrast, showed that both BB complexes of 0. nutans are
very similar to each other. He hypothesized that 0. nutans evolved directly
from 0. grandiflora by accumulation of reciprocal translocations and the simultaneous ac
quisition of sporophytic lethals that had lost the self-compatibility
character.
Recent work (Steiner & Stubbe 1984, 1986; Schumacher & Steiner 1993) has demon
strated that 0. grandifiora has considerable translocation variation, suggesting that 0. nu
tans could have been derived directly from it. Analysis of the phenotypes of the com
SYSTEMATICBOTANYMONOGRAPHS
26
VOLUME 50
(1990) indicated that the two complexes of 0. nutans are very simi
lar to those of 0. grandiflora, especially in populations from the southern part of the range
of 0. nutans, closest to adjacent populations of 0. grandiflora. This also suggests that the
complexes may have had a common origin. There was, however, phenotypic variation
plexes by Wasmund
among complexes
different
of different
phenotype
grandifiora.
represents
Furthermore,
strains, which
ledWasmund
an independent
themost common
origin
segmental
to speculate
that each slightly
populations of 0.
arrangement occurring in Biennis
from different
I was not found in any of the 0. nutans strains studied. Because Wasmund did not have
many strains from the northern part of the range of 0. nutans, it is not possible to refute
Cleland's hypothesis completely. Yet, Wasmund's
results suggest a scenario that virtually
reverses Cleland's
hypothesis:
the two races of 0. biennis arose via hybridization between
of the segmental arrangements in the
0. nutans and 0. villosa followed by diversification
newly formed 0. biennis (Biennis-I).
CC GENOME SPECIES
The third lineage consists of the very distinctive
shale barrens in the Allegheny
United
States.
Mountains
It has a CC genomic
0. argillicola,
which
is restricted to
region of the eastern
and is the only species with plastome V
of the mid-Appalachian
composition
(Fig. 3).
Oenothera argillicola is a distinctive species that appears to have been derived early
in the evolution of subsect. Oenothera.
It is the only outcrossing species of those having
a C genome. This genome may have concomitantly
arisen with the evolution of this
species or its immediate ancestor. It is also the only bivalent-forming
species of subsect.
in the northeastern United States. It may have been more widespread
in the
Oenothera
for other shale barren endemics, such as Trifolium vir
past, as has been hypothesized
Small
The
immediate progenitor of 0. argillicola, which has been
ginicum
(Isley 1990).
hypothesized
(Cleland 1972) to have had plastome IV rather than V, appears to have been
involved, at least indirectly, in the formation of two PTH species, 0. oakesiana
0. parviflora
(AC) and
(BC).
MIXED GENOME SPECIES
Three additional species, all PTH species, had hybrid origins: 1) Oenothera biennis,
including Biennis-I and Biennis-II of Cleland (1972) (AB or BA genomes with plastome
II or III); 2) 0. oakesiana, formerly treated as Parviflora-II of Cleland or as a variety of
0. parviflora of Munz (1965) (AC genomes with plastome IV); and 3) 0. parviflora, for
merly treated as Parviflora-I (BC genomes with plastome IV). All three species are widely
and naturalized in many areas of the
distributed, primarily in eastern North America,
world, especially in Europe. Based on current data, we have summarized the likely ori
gins for these mixed
genomic species in Figure 4.
Populations with AB or BA genomic combination with plastome II or III grouped to
gether as 0. biennis have evolved through a number of hybridization events in a similar
manner as that discussed for the PTH homozygous
species 0. villosa and 0. nutans. Cle
land (1972) suggested an origin of the two primary races of 0. biennis through hy
bridization of his hypothetical Population 2 (B genome ancestor of 0. grandiflora) with
Population 3 (A genome ancestor of 0. elata). A more plausible hypothesis, based on our
better understanding of the origin of certain species, such as 0. nutans, is that all of the
PTH species with a mixed genomic composition (AB, BA, AC, BC) have arisen through
OENOTHERA
1997
27
'C
heterozygous
diagram showing the most likely hybrid origins of the genomically
sect. Oenothera
subsect. Oenothera
and 0. stucchii).
(except for 0. glazioviana
FIG. 4. Schematic
of Oenothera
taxa
hybridization events where at least one or both of the parents were small-flowered autog
amous PTH taxa. These may have had either homozygous or heterozygous genomic com
positions. In the case of 0. biennis, this means that there are several alternative hypothe
ses for the separate origins of Biennis-I and Biennis-11 from other PTH taxa. The studies
of the segmental arrangements by Cleland and others (summarized in 1972) helped gain
an understanding of how the races of 0. biennis came into being.
These studies coupled with more recent work led to the hypothesis that the Biennis
I and Biennis-11 races of 0. biennis arose independently via hybridization between differ
ent strains of 0. villosa subsp. villosa for the A genome and from the aXand f3complexes
states that Biennis-111 (0. nu
of 0. nutans for the B genome. Cleland (1972, p. 282-283)
tans) originated via hybridization between Biennis-I and Biennis-11 because of the simi
larity between the end arrangements of their complexes; however, the additional informa
tion reviewed here has led us to the conclusion that just the reverse occurred.
Cleland pointed out that plants with the A genome were associated with dry habitats
and those with the B genome with more mesic habitats, and that 0. biennis was interme
diate in this respect. Once initial hybridization had taken place, additional biennis pheno
types may have evolved from backcrossing of 0. biennis with 0. nutans and 0. villosa,
or from hybrids of 0. biennis with 0. oakesiana and 0. parviflora.
The Biennis-I group is strikingly uniform in the segmental arrangements of the chro
mosomes
of its ax complexes or B genome (Cleland 1972, p. 272). The majority of the
strains studied by Cleland (56/78 strains) share a single arrangement that differs from the
presumed original arrangement of subsect. Qenothera by one translocation. This common
arrangement differs from most of the arrangements in the ax complexes of 0. nutans by
two translocations, suggesting that they could share a common origin. Most of the other
SYSTEMATICBOTANYMONOGRAPHS
28
arrangements
of 48 Biennis-I
found in cxcomplexes
VOLUME 50
strains were only one (32), two (13),
or three (3) translocations different, and exhibited no apparent geographical pattern.
The ,Bcomplexes (A genome) of Biennis-I represent the opposite extreme. Of the 91
kinds of chromosome end arrangements possible, 67 have been found among the strains of
studied (Cleland 1972, p. 277). There are two arrangements that predominate in
the central part of the geographical range, with an additional 33 out of 67 strains studied
that have other arrangements. These other arrangements predominantly occur in moun
Biennis-I
or in the southeastern United States.
tainous regions of the Ozarks and the Appalachians,
The occomplexes (A genome) of Biennis-II are more homogeneous
than those in Bi
ennis-I. The most prevalent arrangement (26/39 strains) occurs throughout the geograph
ical range of Biennis-II.
translocations different
The few other arrangements that are known are only one or two
from the primary one. On the whole Cleland felt that these com
plexes were similar to, but more primitive than, the f complexes (A genome) of Biennis
I. He concluded that the ox complexes of Biennis-II most likely shared a common origin
with
the oxcomplexes
villosa
of Oenothera oakesiana (Parviflora-I1) and the , complexes of 0.
(Cleland 1972, p. 277), whereas the D complexes of Biennis-I have been derived
from another variant of 0. villosa
subsp. villosa.
The
,Bcomplexes (B genome) of Biennis-II are also a relatively homogeneous group,
but differ on average by four translocations from the ct complexes of Biennis-I, and there
3 complex in 0. nutans is
fore appear to have had different origins. The most common
identical
to two of the most
common
arrangements
found
in Biennis-II,
suggesting
a
common origin.
Oenothera oakesiana originated via hybridization, but a number of possible paths
exist to create an AC genomic combination with plastome IV. These hypotheses can be
summarized as follows:
x 0. parviflora (BC-IV)
x 0. parviflora
3) 0. villosa subsp. villosa (AA-L) x 0. parviflora
4) 0. biennis (Biennis-I1, AB-II) x 0. argillicola ancestor
5) 0. argillicola ancestor x 0. villosa subsp. villosa
1) 0. biennis
(Biennis-II, AB-II)
2) 0. biennis
(Biennis-I, BA-Ill)
(CC-IV)
Each of the hypotheses require somewhat different assumptions, and some are more
than others. Moreover,
these hypotheses differ in some respects from Cleland's
(1972). He proposed an origin from a hybridization between two ancestral populations,
likely
i.e., the 0. argillicola ancestor (Population 1) and a xerophytic A genome population
(Population 3). The hypotheses stated here are more parsimonious, because they do not
require a hypothetical ancestral Population 3 that no longer exists in eastern North Amer
ica, but instead suggest that the A genome came from another PTH entity. As such, these
all require one or more of the other PTH species (0. biennis, 0. parviflora, or
to have evolved first, and all but one requires that 0. parviflora evolved first.
all of the hypotheses involving 0. parvifiora as one of the parents require that
Moreover,
the pollen parent contributes plastids and that those plastids dominate in the hybrid. This
is not a problem in the second hypothesis, because plastome III is not particularly viable
in a BC genomic combination; however, the second hypothesis requires the A genome to
be transmitted through the egg (ox),which is a relatively rare event, but certainly possible
hypotheses
0. villosa)
inOenothera.The thirdhypothesis,which involves0. villosa subsp.villosa, is supported
by the overlapping
distributions
in the Great Lakes
region of that species and a recently
OENOTHERA
1997
29
formed 0. parviflora. In this hypothesis it is easy to see how plastome IV would domi
nate, since plastome I obtained from 0. villosa subsp. villosa would not be particularly vi
able in a BC combination.
In contrast, a hypothesis involving 0. argillicola directly appears unlikely, because
plastome V clearly evolved from plastome IV, but an alternative origin involving the pre
sumed ancestor of 0. argillicola would be possible. This hybridization would be unlikely
unless the ancestor of 0. argillicola had a considerably more western distribution than the
present range of 0. argillicola. If plastome IV arose only once, then the fifth hypothesis
would also require that 0. parviflora, the only other species with plastome IV,must have
subsequently via hybridization with 0. oakesiana. The evidence does not
favor any one path of origin over another; however, hybridization between
0. biennis or 0. villosa subsp. villosa and 0. parvifiora appears to be slightly more likely
been derived
conclusively
than the other hypotheses
(see Fig. 4).
parvifiora originated via hybridization, but a number of pathways are pos
sible that would result in a BC genomic combination with plastome IV. These hypotheses
can be summarized as follows (the first hypothesis the most likely):
Oenothera
1) 0. nutans (BB-III) x 0. argillicola ancestor
2) 0. biennis (AB-I1) x 0. argillicola ancestor
3) 0. biennis x 0. oakesiana (AC-IV)
4) 0. nutans x 0. oakesiana
As in the case of 0. oakesiana,
(CC-IV)
each of these hypotheses
requires somewhat different
and some are more
assumptions,
likely than others. If 0. oakesiana had been derived first,
then it seems likely that 0. parviflora would have arisen via the third or fourth hypothe
sis; however, it ismore plausible that 0. parvifiora was derived first given that hybridiza
tion between 0. biennis and 0. parvifiora seemed to be a likely path of origin for 0. oake
siana. Therefore, hypotheses 3 and 4 appear less likely than the first two hypotheses.
3 is unlikely for the additional reason that hybridization between 0. biennis
Hypothesis
and 0. oakesiana is usually not successful in either direction, but possible only when the
C genome and plastome IV of 0. oakesiana are transmitted exceptionally through the egg.
The first two hypotheses are both similar to Cleland's hypothesis for the origin of 0.
parvifiora (Parviflora-1), except that he invoked a hypothetical B genome-contributing
an
cestor that in turn also gave rise to 0. grandifiora. That ancestor would therefore have
been a large-flowered outcrossing entity, which, according to recent work on 0. grandi
flora, could well have been a more widespread 0. grandifiora. It is simpler to suggest an
origin for 0. parviflora directly via hybridization with another PTH species, such as 0.
nutans. This hypothesis also seems more feasible in terms of the distribution of those
species, and thus hypothesis 1 appears to be most likely (Fig. 4).
RECENT ANOMALOUS SPECIES
Exceptionally we recognize two additional PTH species of hybrid origin: 0. glazio
viana (formerly known as 0. lamarckiana auct. or 0. erythrosepala; AB-III); and 0. stuc
chii (recently described in Italy; AA-I). These differ from all other cases of hybridization
outside the natural range of the subsection in that they have unique, highly distinctive fea
tures. Both have originated outside the indigenous range, the former possibly in England
and the latter in Italy. Oenothera glazioviana has achieved a nearly worldwide distribution.
SYSTEMATICBOTANYMONOGRAPHS
30
VOLUME 50
The origin of 0. glazioviana
is not exactly clear, but it almost certainly arose via hy
bridization between two garden-grown or escaped species introduced to Europe sometime
before 1860, when it appeared in the garden trade. Davis (1911) suggested that it arose
sometime in the late 1700's. Cleland (1972) summarizes much of the history of 0. glazio
viana. Here we provide a few additional comments.
Three competing hypotheses
Currently, we cannot differentiate
be summarized
have been proposed for the origin of 0. glazioviana.
between the different hypotheses. The hypotheses can
as follows:
1) 0. elata subsp. hookeri (AA-I) x 0. biennis (BA-IIl)
2) 0. elata subsp. hookeri x 0. grandifiora (BB-III)
3) 0. biennis (AB-IL or BA-III) x 0. grandiflora
between 0. biennis and 0. elata would provide the correct genomic
Hybridization
and plastome, if the strain of 0. biennis were Biennis-I. This hypothesis was
combination
(1916a, 1916b, 1924), who synthesized his neolamarckiana by ex
two species (summary in Cleland 1972, chapter 17). Cleland
these
perimentally crossing
supported
(1972)
this hypothesis as the most likely, and thought that the event took place
between the two species introduced in ballast dumped in a European port, perhaps Liver
first proposed by Davis
pool (Davis 1912). Several points of evidence support this hypothesis. First, the P com
plex of 0. glazioviana
(gaudens) has the same chromosome end arrangement as several
European a complexes of 0. biennis: rubens, Poznan I, Poznan II, and Poznan III (Cle
land 1972, pp. 339-340). Moreover, all of the European races of 0. biennis known to have
this arrangement have plastome II, not III. The other parent in this hypothesis would be
0. elata subsp. hookeri. It would provide an A genome as well as many of the morpho
logical characteristics of 0. glazioviana,
including the large, outcrossed flowers. More
over, the A complex velans of 0. glazioviana differs from known complexes of 0. elata
subsp. hookeri (franciscana and hookeri) by a single reciprocal translocation. The only
problem with this idea is that 0. elata was apparently only occasionally cultivated in Eu
rope at that time, and we have seen no evidence of naturalized populations.
The weakness of the second hypothesis
is that the only chromosome end arrange
ments known in 0. grandiflora (Steiner 1951; Steiner & Stubbe 1984, 1986) differ from
the B genome of 0. glazioviana by several reciprocal translocations. In 1979, Raven et al.
commented that new arrangements were being discovered in 0. grandiflora by the work
of Steiner and Stubbe; however, none have turned out to be the same as the B genome of
0. glazioviana.
In the third hypothesis, the A genome would have been contributed from 0. biennis
and the B genome
see summary
from 0. grandiflora.
in Cleland
This idea was
1972, chapter 17), but was
the first suggested by Davis
soon displaced
(1911;
by the hypothesis
of
hybridization between 0. biennis and 0. elata. Both 0. biennis and 0. grandiflora were
certainly in cultivation in Europe at the appropriate time, but the proper chromosome end
arrangements are not known.
The origin of 0. stucchii appears to have been similar to that of 0. glazioviana.
It is
a species that arose via hybridization between two naturalized populations. Oenothera
glazioviana, which presumably arose in the 19th century, is now distributed nearly world
wide. In contrast, 0. stucchii appears to have evolved very recently, and is beginning to
spread from northern Italy, where it probably arose, to southern France. The earliest col
lection known is from 1952 (Soldano 1979).
OENOTHERA
1997
31
stucchii is probably a stabilized hybrid between 0. jamesii and one of the
European phenotypes of 0. biennis. The probable 0. jamesii parentage is supported by the
0. jamesii-like
features of long floral tubes, quadrangular buds, and the short-petiolate
Oenothera
analysis indicates that 0. stucchii has plastome I. In 1993, Dietrich
crosses to investigate the origins of 0. stucchii, which included 0.
stucchii with 0. argillicola
(CC-I), 0. biennis (AB-II), 0. elata (AA-I), 0. glazioviana
(AB-II), 0. jamesii (AA-I), 0. nutans (BB-III), 0. oakesiana
(AC-IV), 0. parviflora
rosette leaves. Recent
made
experimental
(AA-I). Analyses of the resulting hybrids demon
I and genomic composition AA. One of the A
genomes is from 0. jamesii and the other is from an 0. biennis-like strain, which exhibits
some B genome characters, including large petals, short floral tube, and dense glandular
pubescence on the floral tube and sepals. The A genome characteristics from 0. jamesii
(BC-IV), and 0. villosa
strated that 0. stucchii
include small petals,
sepals. Oenothera
subsp. villosa
has plastome
long floral tube, and appressed
stucchii
is phenotypically
but with a long floral tube. Hybrids
more
pubescence
like 0. villosa
between 0. stucchii
on the floral tube and
than any other species,
A genome)
(with the B-infected
are pale green or variegated. In contrast, a hybrid with a pure
constitution would be normal green, whereas an AB with plastome I in the
above hybrid combination would also be variegated or pale. Thus, we have concluded that
the A genome of 0. stucchii from 0. biennis also appears to have some B genome genetic
and 0. elata or 0. jamesii
AA genomic
characteristics.
Although we do not formally recognize the numerous unique European phenotypes
of 0. biennis or other hybrid combinations, we accord 0. stucchii specific rank, because
it represents a stabilized entity with a distinctive new genomic phenotype (AA) unknown
It has a true-breeding unique phenotype (as has 0. glazioviana),
is intermedi
two very different species with different genomic compositions,
and cannot
be
accommodated
within either of these taxa. The numerous European forms of 0.
easily
elsewhere.
ate between
biennis are much
this species
easier to accommodate
within
the broad variation pattern expressed
by
in its wide natural range in North America.
TAXONOMICHISTORY
The taxonomic history of Oenothera began when Linnaeus described 0. biennis, the
type of the genus, in Species plantarum (1753). Linnaeus also described two other species
in Oenothera subsect. Oenothera, 0. parviflora in the 1759 edition of Systema naturae,
and 0. muricata
(=O. biennis)
in the 1767 edition. During
century a number. of other species were described,
rently recognized:
although
0. grandiflora
L'Her.
in Aiton
the closing decades of the 18th
including two additional species cur
(1789), and 0. villosa Thunb. (1794),
this species went under other names until 1976, themost common of which were
0.
strigosa, 0. depressa, and 0. canovirens. All of the other names published in the
1700's are now considered to represent one of these four species: 0. angustifolia and 0.
glabra by P.Miller in 1768 (both considered to represent 0. parviflora, although in the
latter case there is doubt about the application to 0. parviflora); and 0. grandiflora by
Lamarck in 1798, which represents 0. grandiflora L'Her.
Detailed studies of Oenothera were initiated when, in 1886, Hugo de Vries discov
ered populations of 0. glazioviana
(as 0. lamarckiana) in the Netherlands
(de Vries
which
1895),
became the subject for lifetime study by a considerable number of scientists
during
the following
century. Currently,
intensive
study of Oenothera,
which
has con
SYSTEMATICBOTANYMONOGRAPHS
32
VOLUME 50
has a considerably lower level of
laboratories, especially Dusseldorf
(Stubbe and
Dietrich), Munich
(Herrmann and collaborators), and Amherst (Mulcahy and collabora
tors). In recent years, molecular studies have contributed to a resurgence in the study of
subsect. Oenothera. Many notable geneticists have studied species of subsect. Oenothera,
tributed much,
especially
activity, but work
to the study of cytogenetics,
continues
in several
including H. de Vries, R. R. Gates, B. M. Davis, 0. Renner, H. H. Bartlett, F. Oehlkers,
G. H. Shull, R. E. Cleland, D. G. Catcheside, S. H. Emerson, E. E. Steiner, W. Stubbe, and
F. Schotz. The special meiotic behavior occurring in PTH plants has been studied inten
sively by R. R. Gates, B. M. Davis, and, especially, R. E. Cleland. The transmission of
plastids through pollen, which is unusual in higher plants because the heredity of plastids
is usually exclusively maternal, and the interactions between plastome and genome, have
been studied by Renner, Schotz, and Stubbe. The results of these studies supply the foun
dation for the taxonomy of subsect. Oenothera presented here.
Concerning study of overall patterns of variation in subsect. Oenothera, the morpho
logical studies by Munz (1949, 1965) are the only comprehensive ones for the species in
is the only one that has been useful for experimental re
search efforts such as Cleland's as well as for general-purpose
identification. For the most
part, Munz's concepts with few exceptions have been widely adopted in regional North
North America.
His classification
American
treatments (e.g., Gleason & Cronquist
in Europe have differed greatly.
The history of Oenothera subsect. Oenothera
1991; Welsh
et al. 1987), but taxonomic
schemes
in Europe has been summarized by Cle
treated taxonomically by Raven (1968), although the lat
in utilizing a European concept for the taxa recog
ter treatment was very conservative
land (1972, chapter 19) and was
nized. By contrast, the revision presented here is a comprehensive
attempt to include all
of the many described species, especially from Europe, into a single taxonomic system
with a uniform
that allows information retrieval, gives predictive value,
treatment of the North American
phylogenetic
information. Munz's
Oenothera species (1965), which represented a tremendous step forward in comprehend
ing this group, is altered in this revision in several respects. Our taxonomy is compared to
Munz's and to Cleland's (1972) in Table 5. The reasons for these changes are discussed
under the respective species.
species concept
and incorporates
there has been a strong tendency
During the intensive work on subsect. Oenothera,
by those primarily concerned with genetics to describe each of their true-breeding exper
imental strains as distinct species. The majority of the names have been published for PTH
taxa, especially the five species with wide natural and naturalized distributions: 0. bien
nis (96 names), 0. parviflora (60), 0. villosa (46), 0. glazioviana
(45), and 0. oakesiana
(45). This practice has continued up to the present time, especially in Europe, where a
large number of species, hybridogenous
species (those originating as hybrids, but existing
and spreading from their place of origin), and hybrids are recognized (e.g., Jehlik & Ros
tan'ski 1995). In North America
the attempts by geneticists Bartlett (1914) and Gates
(1936) to create classifications,
especially for the eastern North American species of sub
sect. Oenothera, were not particularly useful, because their approach was to name selected
phenotypes among the thousands occurring in the natural ranges of the subsection. The
splitting done by Bartlett and especially Gates has never been accepted by North Ameri
can taxonomists and is of historical interest only.
Renner
(1937, p. 206; 1938, p. 102; 1942, p. 465), a physiologist and geneticist, ex
promoted the idea that a reasonable taxonomy in Oenothera
is gained only
through genetic studies in which the genetic constitution of the phenotype is examined.
plicitly
OENOTHERA
1997
Table
5. Comparison
sect. Oenothera,
1972)
of the major
the system
including
and Munz
(1965).
treatments
presented
The genomic
of taxonomic
33
systems
in this paper compared
and plastome
combination
for Oenothera
sect. Oenothera
to those of Cleland
type are given
(summarized
in parentheses
in the first column.
Dietrich,Wagner, andRaven
0. elata
Munz (1965)
Cleland (1972)
0. elata
0. hookeri
subsp.angustifolia
subsp.grisea
subsp.hewettii
subsp.hirsutissima
0. elata
0. hookeri
(AA-I)
subsp. elata
subsp.hirsutissima
subsp. ornata
subsp.venusta
subsp.hookeri
subsp.montereyensis
subsp.hookeri
0. hookeri
0. jamesii (AA-I)
0. jamesii
0. hookeri
0. longissima (AA-I)
0. longissima
subsp. longissima
subsp.clutei
0. hookeri
0. wolfii (AA-I)
0. hookeri subsp.wolfii
0. hookeri
0. villosa (AA-I)
subsp.villosa
subsp. strigosa
0. strigosa
subsp. canovirens
subsp. strigosa
subsp.cheradophila
0. strigosa
0. stucchii (AA-I)
-
-
0. grandiflora (BB-III)
0. grandiflora
0. grandiflora
0. nutans (BB-III)
0. biennis
0. biennis-III
subsp. austromontana
0. biennis
(AB-I1 or BA-I1)
0. biennis
subsp. biennis
(European)
subsp.centralis
subsp.caeciarum
0. glazioviana
(AB-I1 or -III)
0. argillicola (CC-V)
0. biennis-I
0. biennis-II
0. erythrosepala
0.
0. argillicola
0. argillicola
lamarckiana
var. argillicola
var. pubescens
0. oakesiana (AC-IV)
0. parviflora subsp.
parviflora
0. parviflora (BC-IV)
0. parviflora-II
var. oakesiana
0. parviflora
subsp.parviflora
var. parviflora
subsp. angustissima
0. parviflora-I
sub
in
after the taxon
34
SYSTEMATICBOTANYMONOGRAPHS
This premise
has been advocated
by Cleland
(1972, p. 304), who
VOLUME 50
said that Renner was
in Europe at the time, because he "has made the only exten
sive study of the European Oenothera flora that is based on cytogenetic as well as pheno
typic criteria, his conclusions possess a validity not found in purely taxonomic studies."
unique among
the scientists
the PTH populations
approach follows this philosophy
by aggregating
(or mi
into species delimited according to the composition of their genomic com
crospecies)
plexes, plastome type, and associated morphological
characters. Such a system reflects the
knowledge of the evolutionary history of the group and provides a reliable means for iden
Our
tification.
The papers by Renner
1975, 1982,
(1937,
1938, 1942, 1950,
1956) and Rostan'ski
(1965,
1966,
1982; Jehllk & Ros
taniski 1979) were attempts to provide a comprehensive
taxonomy for the populations in
Europe as well as the new combinations resulting from hybridization in Europe, those of
Renner
1985; Rostan'ski & Ellis
1979; Rostan'ski & Forstner
from a genetic point of view, and those of Rostan'ski
morphological
one. Unfortunately,
ropean species of Oenothera
and others from a primarily
a system for the Eu
were done apart from almost any consideration of the situa
most of these attempts to develop
tion within
the indigenous range of subsect. Oenothera in North America.
This narrow interpretation of species used in Europe has resulted in the description of
80 taxa (Table 4) for strains of naturalized PTH species or hybrids between them. The mu
tations of "Oenothera lamarckiana," described by de Vries as newly evolved species, are
not included in this list, nor are the names based on artificial hybrids, nor those not validly
published for European populations. These 80 taxa, based on wild-collected
populations,
are described
at the species, variety, or forma level. Many of these names (32) are con
sidered here to represent additional phenotypes of 0. biennis, whereas some are assigned
to other PTH taxa, including9 to 0. glazioviana, 2 to 0. oakesiana, 5 to 0. parviflora,
and 7 to0. villosa subsp.villosa. In addition to0. glazioviana only one of these taxade
scribed in Europe, 0. stucchii, is recognized as a species. The list also includes 23 hybrids
that have been reported from Europe and assigned formal names.
Only two hybrids are accepted here as species, 0. glazioviana and 0. stucchii, be
cause these particular hybrid combinations have resulted in plants with divergent pheno
types and unique genetic attributes coupled with their spread and establishment. Another
entity of this kind, described from Germany, is 0. purpurata, an AA genomic phenotype
with plastome II (Stubbe 1959), but it is known only in cultivation. It is apparently a bi
valent-forming
segregate
most likely the European
that occasionally
phenotype
occurs
rubricaulis.
tity and the fact that it does not occur
in populations of an 0. biennis strain,
In view of the unique origin of this en
in nature, we have chosen not to regard it as a
species and have included its name in the synonymy
of 0. biennis.
INFRASECTIONALCLASSIFICATION
Oenothera
subsect. Oenothera
is a group of very closely related species; additional
of the group, such as proposed by Rostan'ski (1985), does not seem ap
propriate. Itmight be argued that further division is possible, if groups could be delimited
to correspond with the three major genomes. This is not possible because hybridization
formal subdivision
has played a major
role in the formation of the PTH species within subsect. Oenothera,
and these mixed genomic species are impossible to place in the system.
We do not accept Rostan'ski's subdivision of subsect. Oenothera
into five series for
1997
OENOTHERA
35
several reasons. First, the series of Rostan'ski
are not equivalent to series in other parts of
or the remainder of the genus in their level of morphological
and genetic
distinctness (see Dietrich 1977; Wagner et al. 1985; Dietrich & Wagner 1988). The taxo
nomic level of series has been used in Oenothera and elsewhere in the Onagraceae to des
sect. Oenothera
ignate distinctive groups of very closely related species within more heterogeneous
sec
tions, such as sect. Oenothera. The series are characterized by distinct morphological
features, crossing relationships, and often plastome type or geographical distribution. If
the situation with
the PTH species did not involve hybridization, then the formal recogni
subsect. Oenothera that corresponded to the major genomes and as
tion of series within
sociated plastomes
would
only partially follow
be possible.
The problem with Rostan'ski's
series is that they
this concept.
In addition, Rostan'ski's treatment is based on a species concept that splits the species
accepted here into many microspecies.
Finally, his treatment does not convincingly reflect
the natural relationships of the species of subsect. Oenothera. For example, series II, se
ries Devriesia, which includes primarily species with AA genomic combinations, also in
cludes 0. insignis, which is an AC type, and 0. pedemontana, which has an AB compo
sition. In his series III, series Oenothera, he treats predominantly
the AB combinations,
but includes 0. rubricapitata (BC), 0. wolfii (AA), and 0. strigosa (AA). Moreover, 0.
ersteinensis (AB) and 0. perangusta
(AC) are listed as synonyms of O. strigosa (AA). In
series V, series Rugglesia, he assembles all of the C genome combinations, but also in
cludes 0. nutans (BB) through misinterpretation
of the epithet "nutans." Overall, there
fore, Rostan'ski's classification
is unnatural and uninformative, and we urge its abandon
ment.
NOMENCLATURE
Given the wide geographical distribution of species of Oenothera subsect. Oenothera
and the array of scientists studying the group, especially non-taxonomists,
we felt it es
sential to make a special effort to locate and analyze every name published in the widely
scattered literature. A corollary to compiling a comprehensive
synonymy was to analyze
each name for compliance with the Botanical Code and to fix the usage via lec
totypification of each name without a holotype. Because many of the Oenothera
re
carefully
searchers were
the PTH system rather than doing descriptive
investigating
taxonomy,
they published a significant number of problematic names. We present a thorough analy
sis of the extensive nomenclature of this complex group, including those names that can
not be considered validly published (156), primarily from the genetics literature. They are
predominantly names given to experimental strains in cytogenetic studies, but some are
from taxonomic work. Certain authors investigating Oenothera, especially performing ex
perimental rather than taxonomic analyses, did not modify their methods to comply with
changes in the Botanical Code; thus they did not fulfill one or more of the ICBN articles
(Greuter et al. 1994), and hence the names are not validly published. In fact, a number of
these experimental workers, such as 0. Renner, were merely giving binomials to their ex
perimental strains and did not really intend to propose new species. When the Code be
came more explicit in the requirements for valid publication (e.g., Latin diagnosis re
quired) a large number of the names subsequently assigned to experimental Oenothera
strains were not validly published. A good example of this problem is seen in Renner's
papers inwhich he gave binomials to experimental strains. None of Renner's names pub
SYSTEMATICBOTANYMONOGRAPHS
36
VOLUME 50
lished after 1935, when it became mandatory to include a Latin diagnosis or description
when publishing new names, are valid.
Rostan'ski did not indicate holotypes for many of his names after it became necessary
under the Code to do so. Names not validly published are listed in an appendix. In most
cases we have listed nomina nuda in the appendix of names not validly published; if a
nomen nudum was subsequently validated by another author we have given the place of
publication for the nomen nudum in square brackets preceding the validating author and
the bibliographic
citation.
(1901a; 1901b, 1903) described and gave binomials to numerous phe
the course of his experimental work on cultivated strains of 0.
glazioviana. These names were based on cultivated plants, and we have been unable to lo
cate any material preserved of most of these entities. He probably did not make vouchers
Hugo de Vries
notypes
arising during
for them. We have designated lectotypes based on the photographs in the original publi
cations. In other cases we have designated a neotype based on a photograph in a subse
quent publication or merely given an indication of photographs representing the entity in
one of his publications. De Vries also published numerous other names, especially "mu
tations," that are not validly published. These are listed in the appendix of names not
validly published.
A
Bartlett
similar situation is encountered with the publications of both H. H.
(especially Bartlett 1914) and R. R. Gates (1936). The new taxa in these papers
somewhat
were based on cultivated
plants from wild-collected
seeds or rosettes. The experimental
strains were often cultivated
through several generations. Sometimes vouchers were pre
served, and other times apparently they were not. An additional difficulty with Gates's
material was that an original set of his experimental strains grown in Regent's Park Gar
dens in London was lost or discarded along with the entire collection at King's College
shortly after World War II (A. 0. Chater, pers. comm.). Gates stated (1936) ". . .complete
plant specimens from many of the cultures are now at Kew, and several other sets of spec
imens are being prepared for circulation to other leading herbaria [BM, GH], an original
set being retained in the herbarium at King's College." Because the original set of mate
rials at King's College, which would have represented holotypes of Gates's names, is not
extant, we have designated many of the collections at K, BM, or GH as lectotypes.
In their publications both Bartlett and Gates often cited a number of strains or gener
ations with different numbers for each. We have treated these as syntypes and have se
lected lectotypes from among them focusing especially on those for which there are pre
served collections in herbaria rather than photographs in the publications, but photographs
were used when they were the only option. We have not cited the numerous culture num
bers of Gates
for each name. The interested reader can find them in his publication.
(1968) also presents a special problem. Until recently his col
lections were held in a personal herbarium. In the 1970's he moved from East Germany
toWest Germany, but was prevented from taking his library and herbarium with him. In
stead they were returned to his former house. The director of HAL was subsequently suc
The work by Hudziok
cessful in obtaining the Hudziok herbarium. We learned of these events in early 1996 and
have been able to examine the type material of his names. Unfortunately,
some of the
types were apparently not among the collections obtained by HAL. Rostan'ski has the sit
uation under study and plans tomake a number of lectotypifications and neotypifications.
We have indicated the material we have seen and the situation as we know it for the oth
ers. We have made two neotypification but have avoided making other typifications that
will be published by Rostan'ski independently.
1997
OENOTHERA
37
TAXONOMY
Many of the species in Oenothera subsect. Oenothera are very common and have ex
tremely broad geographical distributions, and the corresponding representative specimen
citations are extensive. Therefore all specimens examined are cited in an appendix.
Oenothera
L., Sp. pl. 346. 1753. Onagra Miller, Gard. dict. abr., ed. 4, vol. 2. 1754;
Adanson, Fam. pl. 2: 85. 1763, nom. superfl. Oenothera sect. Onagra Fischer &
Meyer, Index secundus sem. hort. petrop. 45. 1835 [authorship following provi
sions of ICBN (1994) Art. 58.3]. Oenothera
subg. Onagra (Fischer & Meyer)
Jepson, Man.
fl. pl. California 679. 1925. Brunyera Bubani, Fl. pyren. 2: 648.
1900, nom. superfl. (based on Oenothera). Usoricum Lunell, Amer. Midl. Natu
ralist 4: 481. 1916, nom. superfl. (based on Brunyera).-LECTOTYPE,
designated
by Rose, 1905: Oenothera biennis L.
Pseudo-oenothera
Ruprecht, Fl. ingr. 1: 365. 1860.-TYPE:
giniana Ruprecht [=Oenothera biennis L.].
Pseudo-oenothera
vir
Annual, biennial, or perennial herbs, caulescent or acaulescent, with erect, ascending,
or occasionally decumbent stems, when decumbent sometimes rooting at the nodes, with
a taproot or fibrous roots, occasionally with shoots arising from spreading lateral roots,
rarely with rhizomes. Leaves alternate, entire, toothed to pinnatifid, often irregularly so;
stipules absent; immature plants usually also with a basal rosette (often absent in mature
plants). Flowers perfect, actinomorphic,
in axils of the apical leaves, when numerous
forming leafy terminal spikes, racemes or corymbs, usually ephemeral, opening near sun
set (and usually wilting in direct sunlight the following day) or near sunrise. Floral tube
well developed, cylindrical and somewhat flared near the mouth, usually deciduous soon
after anthesis. Sepals 4, reflexed, green or tinged red or purple. Petals 4, yellow, purple,
or white, rarely pink, red or merely with a red basal spot, usually aging orange, purple,
pale yellow, or whitish, usually obcordate or obovate. Stamens 8, subequal or the ante
petalous ones shorter; anthers versatile, the sporogenous tissue in each locule undivided;
pollen shed singly, connected by viscin threads. Ovary with 4 locules, ovules numerous;
stigma deeply divided into 4 linear lobes, entire surface of lobes receptive. Fruit a capsule,
usually
loculicidally
dehiscent,
sometimes
tardily so, rarely indehiscent,
straight or
terete to 4-angled or -winged, sessile or the basal portion sterile and stipelike.
Seeds numerous, in 1-2 (-3) rows or in clusters in each locule. Chromosome
numbers: n
curved,
= 7, 14, 21, 28. Self-incompatible
or self-compatible.
is a genus of 119 species of temperate to subtropical areas of North and
Oenothera
South America with a few species in Central America, usually of open, often disturbed
habitats, with the center of diversity in the southwestern U.S.A. and northern Mexico;
sev
eral species are widely naturalized. Oenothera is currently divided into 14 sections, 12 of
which have distributions that include Texas and northern Mexico
(Wagner in Praglowski
et al. 1987). The largest section by far is sect. Oenothera with 70 species, subdivided into
five subsections. Most PTH species exhibit 30-60 (-70)% pollen fertility (Cleland 1972),
except the species of subsect. Munzia, which exhibit pollen fertility of over 90% and are
maintained
by selective fertilization (Schwemmle
1968; Dietrich
1977). Most of the
of
Oenothera
that have become naturalized outside their natural range are PTH,
species
and all of the naturalized
PTH.
species
that have achieved
a wide
naturalized
distribution
are
SYSTEMATICBOTANYMONOGRAPHS
38
VOLUME 50
Oenothera sectionOenothera.
or rarely subacaulescent, with erect,
stems, when decumbent sometimes rooting at the
nodes, with a taproot, rarely with fibrous roots or with shoots arising from spreading lat
eral roots. Basal rosette present, rarely few-leaved and fugaceous. Flowers usually in
Annual,
or perennial herbs, caulescent
biennial,
decumbent
or occasionally
ascending,
leaf axils, opening near
to lax terminal spikes, sometimes solitary in well-spaced
and wilting the following day with direct sunlight. Mature buds with
free sepal tips. Petals yellow, rarely with a red basal spot or entirely red, usually aging or
suborbicular.
ange, obcordate, obovate, broadly elliptic or rhombic-elliptic,
occasionally
dense
sunset, ephemeral,
Capsules
to narrowly
cylindrical
or ovoid, bluntly 4-angled
lanceoloid
or terete, straight
or curved, sessile, rarely (0. stubbei) basally with a short, sterile and stipelike portion, de
hiscent nearly throughout capsule length. Seeds numerous, in (1-) 2 rows per locule, pris
matic
and angled, or ellipsoid
to subglobose,
rarely obovoid
angled, the testa
number: n = 7. Self
and obtusely
reticulate and regularly or irregularly pitted, rarely flat. Chromosome
some species or populations
compatible,
TO THE
KEY
1. Perennial
herbs with
or plant with
nodes,
AND
SUBSECTIONS
self-incompatible.
SERIES
or weakly
long decumbent
a multistemmed
these
stems,
ascending
floral tube 5.5-19
habit;
SECTION
OF OENOTHERA
OENOTHERA
sometimes
at the
rooting
cm long; northern Mexico
and Organ
Mountains, New Mexico (U.S.A.).
subsect. Emersonia
Oenothera
1. Annual,
or short-lived
biennial,
forming matlike
America)
2. Seeds
prismatic,
the surface
erect
herbs with
stems never
clumps,
angled,
P. H. Raven & W. L. Wagner
(Munz) W. Dietrich,
perennial
to ascending
at the nodes;
rooting
pitted; Canada
irregularly
stems,
floral tube 1-5
2. Seeds
ellipsoid
South
cm long.
several
species
Oenothera subsect.Oenothera.
to globose,
not angled,
flower buds with
3. Young
(-16)
to Central America;
widely naturalized.
et al. 1985).
(Dietrich
rarely (Andes,
the surface usually
floral tube curved
downward,
regularly
pitted.
nodding;
southwestern
U.S.A.
toMex
ico and South America.
flower buds with
4. Apex
of petals
4. Apex
acute
floral tube curved upward
to rounded;
subsect. Raimannia
Oenothera
of petals
truncate
central
to southeastern
ser. Candela
flower buds with
subsect. Raimannia
4-9 mm
capsule
valves
slightly
narrowed
spreading
ward
mm
in diameter
the apex; bract not or only
spreading
7. Capsules
their entire
length,
slightly
incurved
narrowed
gradually
section Oenothera
and Mexico.
(Dietrich & Wagner
1988).
slightly
adnate
or recurved
enlarged
W. Dietrich
(Dietrich
1977).
toward the apex; bract ad
after dehiscence.
ser. Renneria
W. Dietrich.
tapering or not at all tapering
to capsule;
capsule
valves
to
distinctly
after dehiscence.
in the apical
third; bract not at all adnate
Oenothera subsect.Munzia ser.AllochroaW. Dietrich.
tance to the capsule.
Oenothera
at base,
terete, rarely somewhat
to capsule.
7. Capsules
1988).
subsect. Munzia
Oenothera
1.5-3.5
(Dietrich & Wagner
to eastern U.S.A.
ser. Raimannia
subsect. Munzia
at base, gradually
in diameter
nate to base of capsule;
6. Capsules
1988).
floral tube straight; South America.
Oenothera
6. Capsules
(Dietrich & Wagner
U.S.A.
& W. L. Wagner
W. Dietrich
floral tube curved upward; central
Oenothera
5. Young
& W. L. Wagner
or straight and erect.
to emarginate.
flower buds with
5. Young
W. Dietrich
subsect. Nutantigemma
Oenothera
3. Young
toward the apex; bract weakly
Oenothera
subsect. Munzia
adnate for a short dis
ser. Clelandia
W. Dietrich.
subsection Oenothera.
sect. Strigosae Rostan'ski, Fragm. Florist. Geobot. 11: 509. 1965.-TYPE:
Oenothera depressa E. Greene [=Oenothera villosa Thunberg subsp. villosa].
Oenothera
1997
OENOTHERA
39
sect. Parviflorae Rostan'ski, Fragm. Florist. Geobot.
11: 512. 1965.
TYPE: Oenothera parviflora L.
Oenothera
sect. Oenothera
subsect. Oenothera
ser. Linderia Rostan'ski, Feddes
Repert. 96: 4. 1985.-TYPE:
Oenothera hookeri Torrey & A. Gray subsp. hook
Oenothera
elata subsp. hookeri Torrey & A. Gray].
eri [=Oenothera
sect. Oenothera
Oenothera
subsect. Oenothera
ser. Devriesia
Rostan'ski,
Feddes
Rostan'ski,
Feddes
grandiflora L'Heritier.
ser. Rugglesia Rostan'ski,
sect. Oenothera
Oenothera
subsect. Oenothera
Repert. 96: 10. 1985.-TYPE:
Oenothera parviflora L.
Feddes
Repert.
96: 5. 1985.-TYPE:
Oenothera
sect. Oenothera
subsect.
Repert. 96: 9. 1985.-TYPE:
ser. Stubbia
Oenothera
Oenothera
biennial or short-lived perennial herbs, rarely annual, caulescent, with
Falcultatively
erect, ascending,
or occasionally
rosette well
Basal
elata Kunth.
Oenothera
developed;
stems 4-25
decumbent
bracts usually persistent.
dm tall, from a taproot.
(-40)
in dense, usually
Flowers
erect,
terminal spikes, spreading at an acute angle to the stem, opening near sunset, ephemeral.
Floral
tube well developed,
usually deciduous
section,
except bluntly quadrangular
species
subterminal. Petals yellow
somewhat
in 0.
or pale yellow,
opaque, or rarely yellowish
cordate or obovate. Capsules
usually erect to somewhat
cola), usually
after anthesis. Mature
jamesii;
white
narrowly
usually
aging orange, pale yellow
and somewhat
lanceoloid
buds terete in cross
free sepal tips terminal or in three
translucent
(0. nutans),
or ovoid, bluntly 4-angled,
and
ob
dehiscent,
rarely at nearly a right angle to the stem (0. argilli
spreading,
straight, or rarely arcuate (0. argillicola),
sessile.
Seeds numerous,
in 2
rows per locule, prismatic and angled, the testa reticulate and irregularly pitted. Self-com
patible,
rarely (0. grandiflora)
self-incompatible;
outcrossing
or regularly outcrossing PTH (0. glazioviana).
(5 species),
autogamous
PTH (7 species),
The species of Oenothera
pecially
near permanent
subsect. Oenothera
tion extends
in North America
and Atlantic
coasts to elevations
central Mexico,
occur in open, often disturbed sites, es
or seasonally wet habitats. The
from southern Canada
up to 3200 m in the Rocky Mountains
El Salvador, Costa Rica,
Guatemala,
indigenous
range of the subsec
from sea level on both the Pacific
southward through
and Panama. The range has been
greatly extended with several of the PTH species
(0. biennis, 0. oakesiana,
and 0. villosa
naturalized
subsp. villosa)
becoming
widely
in many
0. parviflora,
parts of the world.
One other species,
the mostly outcrossing bivalent-forming
0. jamesii, is sparingly natu
ralized in South Africa, Canary Islands, and Japan. Two additional species, 0. glaziovana
and 0. stucchii, apparently have arisen recently via stabilized hybridization and PTH for
mation;
the former is now widely
Departement
Bouches-du-Rhone,
In the following
are used somewhat
the pubescence
distributed
descriptions
subjectively
around the world,
and the latter in Italy and
France.
we use several terms for vesture. Because
in the literature, we provide
the following
these terms
definitions
for
terms used in this monograph.
Glandular-puberulent:
minute,
erect,
long that exude a drop of fluid; on ovaries
cal region of inflorescence,
Long-strigillose:
transparent, bluntly
(capsules),
tipped hairs 0.1-0.2
mm
floral tubes, sepals, stems in the api
and bracts.
appressed
to somewhat
spreading, white,
unicellular
hairs, 0.5-1.7
SYSTEMATICBOTANYMONOGRAPHS
40
mm
at base; on stems,
long, not or slightly broadened
VOLUME 50
floral tubes, and
leaves, ovaries,
sepals.
short, appressed or recurved, white, unicellular hairs, ca. 0.2 mm long, not
at base; occurrence as in long-strigillose.
Villous: erect or somewhat spreading, white or translucent hairs, 1-2 mm long, not or
slightly broadened at base; on stems, leaves, ovaries, floral tubes, and sepals.
Strigillose:
broadened
Short-villous:
Pustulate
like villous, but hairs only 0.5-1 mm
hairs:
long.
long unicellular hairs arising from a reddish
pustule; on stems, ovaries, floral tubes, and sepals.
but with
like villous,
purple to translucent multi-cellular
KEY TO THE SPECIES OF OENOTHERA SUBSECTIONOENOTHERA
Note: The differences
between
fertile pollen and smaller, often shriv
larger, plump,
elled, sterile pollen are easily observed under lOx magnification,
making pollen fertility a
is prepared with standard stains, the
less or no stain, indicating the absence of cytoplasm.
useful character for field identification. When
takes up much
sterile pollen
1. Stigma
elevated
long; pollen
2. Floral
the anthers
above
fertile
90-100%
tube 6-13
3. Mature
at anthesis,
the flowers mostly
(in 0. glazioviana
ca. 50% fertile).
outcrossed;
petals
(2.5-)
cm
3-6.5
cm long.
(-16)
buds 7-12 mm
on ovary
pollen
in diameter,
long; at low elevations
bluntly
in cross
quadrangular
capsules
6-12 mm
in Kansas,
Oklahoma,
after anthesis;
in diameter,
and Coahuila
Texas,
floral tube persistent
section;
the free tips of the valves
and Nuevo
mm
2.5-5
Le6n.
2. 0. jamesii.
3. Mature
buds 5-9 mm
capsules
4-9 mm
in diameter,
terete in cross
floral tube deciduous
section;
1-3 mm
the free tips of the valves
in diameter,
after anthesis;
long; montane
2. Floral
tube 2-5.5
half of plant appearing
4. Apical
half of plant with
of
the inflorescence
cm wide;
leaves 0.4-1
wide,
curved;
but mostly
cent,
sepals often
7. Leaves
at least
flushed with
red-pustulate
lanceolate
dark to bright
usually
mm
spreading,
strongly
elliptic,
dense
always
hairs present;
11. 0. argillicola.
leaves
gradually
green,
10-12
mm
oblanceolate,
cm
to
or a few scattered
oblanceolate
inconspicuous
crinkled;
long red-pus
then stem conspicuously
long; pollen
flushed
90-100%
fertile;
and Mexico.
1. 0.
leaves narrowly
elliptic
to lanceolate,
2.5-4
stem in apical part of plant green or mostly
numerous
long red-pustulate
long; pollen
of apical half of plant, ovary,
naked eye, pustulate
pubes
persistent;
not strongly
hairs;
elata.
cm
green and
free sepal
tips 5-8
ca. 50% fertile; up to ca. 50% seeds abor
tive;widespread.
6. Stems
1-6.5
attenuate
conspicuously
bracts
(-4) cm wide,
and conspicuous,
cauline
crinkled;
with
narrowly
1-2.5
long; anthers 7-23 mm
U.S.A.
green;
covered
long; anthers
and ovary
leaves
cauline
or narrowly
few seeds abortive; western
conspicuously
cauline
red or red-striped.
to gray-green;
dull green
red; free sepal tips 1-7 mm
wide,
spreading;
and erect; cauline
erect or slightly
floral tube, sepals,
some
tulate hairs, or if pubescence
7. Leaves
elata.
to the stem, long-attenuate
eastern U.S.A.
Mts,
tips terminal
stem in apical part of plant with many
with
1. 0.
or ap
towards apex.
with
to narrowly
usually
a right angle
arcuate; Allegheny
capsules
of apical half of plant,
usually
tips subterminal,
at nearly
erect; free sepal
cm wide;
>1.5
narrowed
6. Stems
appressed-pubescent.
and longer erect pubescence,
pubescence
free sepal
spreading
conspicuously
of the inflorescence
gradually
of appressed
capsules
toward apex, usually
5. Apex
to the naked eye exclusively
amixture
to the naked eye.
glabrous
5. Apex
3. 0. longissima.
cm long.
(-6)
4. Apical
pearing
in
habitats
Nevada, Utah, Colorado,Arizona, andCalifornia.
10. 0. glazioviana.
floral tube and sepals often appearing
hairs absent or sometimes
present
and translucent,
glabrous
the pustules
to the
never
1997
41
OENOTHERA
red in fresh material;
pale green and deciduous;
bracts sometimes
1. Stigma
by or below
surrounded
0. wolfii
sometimes
elevated
sepals; petals 0.7-2.5
at anthesis
the stigma
tube 5-6
8. Floral
tube 1.5-4.6
or in
shorter than the
ca. 50% fertile.
cm long; pollen
(-3.5)
onto
directly
anthers and then the petals conspicuously
(-7) cm long; Italy and France.
8. Floral
6. 0.
stucchii.
cm long.
to the naked eye exclusively
9. Plant appearing
10. Apex
above
7. 0. grandiflora.
shed pollen
anthers, which
slightly
green
sepals yellowish
or flushedwith some red; southeasternU.S.A.
of the inflorescence
appressed-pubescent.
in bud; dry capsules
tips erect
erect; free sepal
green or
grayish
dull green.
11. Leaves
dull green
to grayish
11. Leaves
stems,
and ovary densely
floral tube, sepals,
naturalized.
and southern
to eastern U.S.A.
central
to eastern Canada;
ap
5. 0. villosa.
to pale green; stems, ovary, floral tube, and sepals sparsely
green
bescent;
green;
and southern Canada; widely
U.S.A.
pressed-pubescent;
appressed-pu
naturalized.
widely
9. 0.
10. Apex
of the inflorescence
capsules
biennis.
dry
to eastern Canada.
and southern
eastern U.S.A.
rusty brown; mostly
usually
in bud, erect to spreading;
free sepal tips subterminal
curved;
12. 0. oakesiana.
9. Plant obviously
glabrous
12. Apex
a mixture
with
hairs and appressed
of long pustulate
or appearing
pubescence,
to the naked eye.
of inflorescence
13. Plant,
dull green;
predominantly
leaves grayish
strigillose;
eastern U.S.A.
rusty brown; mostly
dry capsules
in bud.
free sepal tips subterminal
curved;
at least in the lower portions,
12. 0. oakesiana.
Canada.
13. Plant predominantly
usually
green;
bright
or appearing
erect-pubescent
usually
dry capsules
to the naked
glabrous
or black;
dark green
eye;
14.
15. Free
sepal
in bud; petals
tips terminal
pale green; capsules
sometimes
and somewhat
1.4-2.5
cm
(-3)
dull green when
southeastern
translucent;
in bud.
so to the naked eye.
or appearing
glabrous
and
13. O. parviflora.
erect; free sepal tips terminal or subterminal
of inflorescence
Inflorescence
leaves
eastern U.S.A.
southernto easternCanada.
12. Apex
to
green
to eastern
and southern
long; bracts
dry; petals
toMaine
U.S.A.
caducous,
white
fading yellowish
and Ontario,
Canada.
8. 0. nutans.
15. Free sepal tips subterminal
capsules
black or dark green when
usually
ally somewhat
opaque;
(-2) cm long; bracts persistent,
in bud; 0.8-1.5
fading pale yellow
dry; petals
and southern
eastern U.S.A.
green;
and usu
to eastern Canada.
13. O. parviflora.
14. Inflorescenceconspicuouslypubescent.
16. Mature
buds 1.7-3
17. Plant
cm long.
densely
petals
with
covered
hairs of several
types;
than the sepals;
shorter
conspicuously
anthers
7-12
southern Oregon
mm
long;
and northern
California, in coastal areas.
17. Plant usually
mm
long; petals
buds 0.8-2
18. Sepals
hairs of several
with
equalling
approximately
and southern
ern U.S.A.
16. Mature
4. 0. wolfii.
covered
sparsely
the sepals
to eastern Canada;
widely
types; anthers 3-6
in length; central
9. 0. biennis.
naturalized.
cm long.
yellowish
green;
to eastern
central
U.S.A.
and southern
to eastern
9. 0. biennis.
Canada;widely naturalized.
18. Sepals
green,
19. Ovary
variously
19. Ovary with
20.
red or red-striped.
flushed with
ern U.S.A.
20.
to eastern Canada; widely
hairs and often also with
pustulate
Inflorescence
and
ovary
also
appressed-
sometimes
U.S.A.
and southern Canada;
Inflorescence
hairs; central
but never with pustulate
pubescent,
and southern
ovary
U.S.A.
(-9)
to east
and
and southern
other
ovary
9. 0. biennis.
types of hairs.
to subappressed-pubescent,
glandular-puberulent
widely
to east
naturalized.
and with
erect
naturalized.
glandular-puberulent;
to eastern Canada;
widely
hairs;
5. 0.
central
naturalized.
villosa.
to eastern
9. 0.
biennis.
SYSTEMATICBOTANYMONOGRAPHS
42
1. Oenothera
elata Kunth
1823. Onagra
VOLUME 50
inHumboldt,
Bonpland & Kunth, Nov. gen. sp. (quarto) 6: 90.
Spach, Nouv. Ann. Mus. Hist. Nat., Paris 4(4): 353.
kunthiana
1836 ["1835"], nom. superfl.-TYPE:
1803, von Humboldt
Mexico,
& Bonpland
4040 (holotype:P-HBK!).
Erect biennial
4-25
to short-lived
dm tall, green, flushed with
perennial herbs with a taproot, forming a rosette; stems
red below or entirely red, unbranched or with branches
obliquely arising from the rosette and secondary
with one of the following patterns of pubescence:
arising from the main stem,
a) exclusively densely strigillose with
strigillose and with a few to many appressed to
some longer appressed hairs; b) densely
to erect hairs, some with
spreading
also sparsely
with white
to densely
veins,
illose or villous
densely
spaced,
leaves 10-43
sparsely
sometimes
cm long, 1.2-4
to densely
strig
also sparsely
(-6) cm wide,
to
nar
or narrowly elliptic, margin bluntly dentate to suben
in lower part rarely sinuate-dentate,
to the petiole. Cauline
to oblanceolate,
the bracts
and margins,
Rosette
to oblanceolate
tire, the teeth widely
margin
Leaves
rarely red, flat, rarely with undulate margins,
on both surfaces
rowly oblanceolate
oblanceolate
bases; or c) as in (b), but inflorescence
dull green to gray green, usually
red-pustulate
glandular-puberulent.
glandular-puberulent.
ally narrowed
branches
leaves 4-25
cm long, 1-2.5
apex acute, base gradu
(-4) cm wide, narrowly
lanceolate
or very narrowly to narrowly elliptic,
the teeth sometimes widely spaced, the lower ones
toward the base, apex acute to long-acute, base acute to atten
narrowly
bluntly dentate or subentire,
sometimes
sinuate-dentate
uate, short-petiolate or sessile. Bracts 1.5-9 cm long, 0.5-2.8 cm wide, forming an acute
or right angle to the stem, narrowly lanceolate to lanceolate or narrowly elliptic to ellip
tic, green to gray-green, margin bluntly dentate to subentire, sometimes undulate or
slightly twisted, apex acute to long-acute,
cuneate.
Inflorescence
Floral tube (2-) 3-5
with
sometimes recurved, base rounded to narrowly
the flowers at an acute to obtuse angle to the stem.
(-5.5) cm long, 1.2-1.8 mm in diameter, yellowish green or flushed
unbranched,
red, with one of the following
sometimes
0.2-1.7
with
scattered pustulate
pubescence
patterns: a) densely
strigillose
to sparsely
strigillose,
hairs; b) densely
to villous,
the hairs
mm
long, and densely to sparsely glandular-puberulent;
or c) densely to sparsely
villous and densely to sparsely glandular-puberulent. Mature buds 2.5-4.5 cm long, 6-10
mm in diameter, narrowly lanceoloid to lanceoloid. Sepals 2.7-5 cm long, 4-8 mm wide,
yellowish
green, red-striped or strongly flushed with red, pubescence same as the floral
tube; free sepal tips 1-7 mm long, strigillose, sometimes also glandular-puberulent,
erect
in bud. Petals (2.5-) 3-5.5 cm long, (2.7-) 3-5.3 cm wide, yellow to pale yellow, very
broadly obovate, apex retuse. Filaments 17-25 mm long; anthers 7-23 mm long; pollen
90-100% fertile. Ovary 1.2-1.8 cm long, ca. 1.5 mm in diameter, densely strigillose and
with some longer appressed hairs or densely to sparsely villous and densely to sparsely
glandular-puberulent
throughout or only at apex, often also with many to a few pustulate
hairs; style 5-9 cm long, the exserted part 2.1-4 cm long; stigma elevated above the an
thers at anthesis, the lobes 4-11 mm long. Capsules 2-6.5 cm long, 4-7 mm in diameter,
narrowly lanceoloid, tapering toward the apex, pubescence
like that of ovary but less
dense, bright to dull green when fresh, dull green or gray-green when dry; free tips of the
valves ca. 0.5-2.5 mm long, truncate or emarginate. Seeds 1-1.9 mm long, 0.6-1.2 mm
in diameter, brown to almost black. Chromosome
number: n = 7
and
06 and
(711;04
511;
or
2 04 and 3II; based on 147 individuals from 68 lo
411;08 and 31I; 06, 04, and 2II;
calities). Self-compatible,mostly outcrossing.
1997
OENOTHERA
43
Phenology. Flowering as early as April, more frequently from June to September, but
in the southern part of the range until February.
Distribution. Widely distributed in open, mesic sites from the coast to a variety of
montane habitats, sea level to 3200 m, in the western United States and Mexico,
from the
Pacific coast in southern Oregon south to northern Baja California, Durango, and Sinaloa,
east throughout the Rocky Mountains
from Idaho towestern Texas, and into the Plains re
gion in southern Kansas, Oklahoma, and eastern Texas; scattered southward through cen
tral and northwestern Mexico,
El Salvador, Costa Rica, and Panama.
Guatemala,
Oenothera elata commonly occurs along streams, inmeadows, on rocky slopes or scree,
in arroyos, and in other disturbed habitats, such as along highways and ditchbanks or on
fallow agricultural land.
Oenothera elata is the most polymorphic and widespread of the outcrossing bivalent
forming species of subsect. Oenothera. It is an AA genomic combination, and it has plas
tome I (Stubbe 1959, 1963, 1964). In the first meiotic metaphase 0. elata plants usually
form 7 bivalents, but floating translocations associated with small rings of four, six, or
are also frequent in certain populations (see also Cleland 1944, 1972;
small translocation rings indicate some degree of diversity in the
chromosomal end arrangements. Cleland (1944, 1972) indicated that the diversity of end
arrangements was greatest in the Rocky Mountain area (0. elata subsp. hirsutissima), and
eight chromosomes
Steiner
1951). These
less diverse on the California
coast (0. elata subsp. hookeri) and inMexico. The new re
here bear this out to some extent, but indicate greater diversity of end
arrangements in 0. elata subsp. hookeri than previously thought.
Munz (1949, 1965) presented a classification of the outcrossing AA genome species
in which he recognized 0. elata, 0. hookeri (subdivided into 9
of subsect. Oenothera,
subspecies), 0. jamesii, and 0. longissima. He also treated 0. glazioviana
(as 0. ery
sults presented
throsepala). Our taxonomy of the Oenothera hookeri group, as Munz referred to it, dif
fers from his classification in a number of respects (Table 5). Most important, many of the
populations referred by Munz to 0. hookeri subsp. wolfii have been given specific status
as 0. wolfii (Raven et al. 1979; Wasmund & Stubbe 1986), because they represent a PTH
entity derived from 0. elata subsp. hookeri. The remaining populations referred by Munz
to 0. hookeri subsp. wolfli were misplaced collections of 0. villosa subsp. strigosa, 0.
elata subsp. hookeri, or an odd hybrid between 0. villosa subsp. strigosa and 0. glazio
viana; they are discussed further under 0. wolfli.
Aside from 0. jamesii and 0. longissima, which we circumscribe in the same fash
ion as Munz, Munz divided the remaining populations of the 0. hookeri group into two
species, 0. elata and the polymorphic 0. hookeri. Munz (1949) stated that "[0. elata] is
questionably distinct from the hookeri assemblage from farther north . . . It intergrades
most definitely with 0. hookeri through ssp. hewettii (irrigua) especially in the plants re
ferred to that subspecies in this paper coming from Coahuila and Texas." Most recent
floristic accounts have maintained Munz's
taxonomy.
More recently Raven et al. (1979) examined an ample series of populations in culti
vation and determined that the characters used by Munz (1949) and Steiner (1951) to sep
arate 0. hookeri from the allopatric 0. elata fail to delineate strongly distinctive groups;
because of the absence of any significant gaps in the variation pattern, we treat 0. elata
and 0. hookeri of Munz as a single species. Studies of living plants during the past 20
years in Dusseldorf by Dietrich, coupled with the study of extensive herbarium material
amassed atMO in 1981 by Wagner and Dietrich, confirmed that 0. elata and 0. hookeri
should be treated as a single species. There are, however, suites of characters that segre
SYSTEMATICBOTANYMONOGRAPHS
44
of this complex
gate populations
VOLUME 50
into three, although not discretely
sub
distinguished,
species. Of the many characters studied, the only features that differentiate populations of
0. elata subsp. elata are the mature buds, free sepal tips (in bud), petal size, free tips of
the capsule valves, and leaf texture. The most notable distinguishing characters of 0. elata
free tips of the capsule
subsp. elata are the indistinct
flowers, broader
smaller
valves,
shape of the buds, and the short free tips of the sepals. The modal morphological
ences,
level at which
taxonomic
After 0. wolfli
indicate
remain. Our studies
hookeri
trapopulational,
or petals. Moreover,
is not uncommon
herbarium specimens,
common
the field suggest
of these subspecies
entities.
The remaining
and 0. hookeri
garden studies in Dusseldorf,
two subspecies
subsp. montereyensis,
ences used by Munz
environment
to differentiate
tions in these characters; moreover,
one essentially
Brazos
ation with
of the phenotype
the degree of di
in a common
grown
among and within
intergradation
garden
in its natural habitat.
expressed
they often vary independently.
Therefore,
popula
we have
coastal entity as 0. elata subsp. hookeri.
as 0. elata
subsp. texensis
a disjunct population
(Dietrich & Wagner
of 0. elata from
1987), known only
indicated a pattern of clinal vari
entity at one endpoint, and grading toward 0. elata subsp.
the newly described
Therefore,
from subsp. hookeri only ap
however, when
from a single locality. Field studies in 1991 by Wagner
hirsutissima.
in
differentiated
trends that probably correlate with
the course of this project we described
Co., Texas,
which
subsp. hookeri
represent a moderately
subsp. montereyensis
environment;
to the maritime
each strain retains much
During
re
treat as 0. elata subsp. hookeri. The differ
Our studies have shown that there is extensive
recognized
this
States and northern
0. hookeri
by Munz,
recognized
collectively
which we
pear to represent weak morphological
rect exposure
as subspecies,
is 0. elata subsp. hirsutissima,
It occurs throughout the western United
in coastal California,
subspecies
in
and study of populations
series can be recognized within
of 0. elata into three subspecies.
The most polymorphic
Mexico.
or not at all geographically
separated.
Our extensive study of hundreds of
in Oenothera.
to 0. elata subsp. elata. They are here accepted
sulting in the subdivision
cludes six of Munz's
in
color, and lengths of the sepals, free sepal tips,
that two nearly distinct population
in addition
complex,
often
of these are based on minor,
most of them were only weakly
This type of variation
be
taxa within 0.
(1949, 1965) infraspecific
that many
in pubescence,
variations
status would
that subspecific
these populations.
to recognize
eight of Munz's
is excluded,
differ
in the chromo
arrangements
segmental
(1951) suggest
in the analysis by Steiner
somes detected
an appropriate
and the different
distribution,
allopatric
we here combine
0. elata subsp. texensis with 0. elata subsp.
hirsutissima.
Two other species with AA genomic
Raven
et al. 1979).
species,
combinations
and plastome
I, 0. jamesii and 0.
related to and perhaps derived directly from 0. elata (Munz 1949;
are closely
longissima,
to the PTH 0. wolfii, discussed above, another PTH
genomic constitution and plastome I, presumably was de
In addition
0. villosa, with AA
rived from 0. elata or its immediate ancestors.
KEY TO THE SUBSPECIES OF OENOTHERA ELATA
1. Stem
and ovary exclusively
2. Mature
mm
buds
long
leaves
(excluding
(in bud); petals
somewhat
strigillose;
stem rarely with
floral tube and ovary)
2.5-3.5
leathery; plant
scattered
lanceoloid,
2-3
pustulate
cm long; free tips of the capsule
in cultivation
<10 dm tall.
hairs (muricate).
cm long; free sepal
valves
la.
tips 1-2
indistinct;
0.
bracts
elata
(-3)
flat;
subsp. elata.
45
OENOTHERA
1997
2. Mature
buds
(excluding
flat or undulate;
bracts
floral tube and ovary)
long (in bud); petals
2-6 mm
narrowly
lanceoloid,
leaves membranous;
plant
in cultivation
cm long; free sepal tips
2.5-5
cm long; free tips of the capsule
3-5.5
valves
usually
conspicuous;
?10 dm tall.
subsp. hirsutissima.
lb. 0. elata
1. Stem
and ovary predominantly
with
spreading
pubescence
(short- and long-villous),
stem often with
pustulatehairs.
3. Sepals
green or flushed with
strigillose
mm
and usually with
long; plant
red, without
in cultivation
indistinct
or with
long pustulate
pustulate
hairs,
hairs, but not glandular-puberulent;
?10 dm tall.
sparsely
villous;
anthers 8-15
lb. 0. elata
stem
(-22)
subsp. hirsutissima.
3. Sepals always flushedwith red,with distinct red-pustulatehairs, usually densely long-villous;
stems
strigillose
puberulent;
with
anthers
la. Oenothera
numerous
12-23 mm
pustulate
long; plant
hairs
and,
in cultivation
especially
toward
<8 dm tall.
the apex,
glandular
Ic. 0. elata
subsp. hookeri.
elata subsp. elata.
[Tabl. ecole bot., ed. 2, 271. 1815, nomen nudum]
salicifolia Desfontaines
ex Seringe inDC., Prodr. 3: 47. 1828, non Oenothera salicifolia J. Lehmann, 1824,
ex G. Don, 1832. Onagra salicifolia (Des
nec Oenothera salicifolia Desfontaines
Oenothera
fontaines ex Seringe) Spach, Hist. nat. veg. 4: 361. 1835.-TYPE: "Oenothera sali
h. p. [hort. Paris], 28 Jul 1815" (holotype: G-DC!).
cifolia Desfontaines,
green, rarely red-flushed below or entirely red, strig
to
the plant. Leaves somewhat leathery, strigillose; bracts
illose, giving a gray appearance
flat, spreading horizontally from the stem, giving the stem apex a broadly obtuse appear
Stems 6-10
dm tall, usually
tube 3.2-4.5 cm long. Mature buds 2.5-3 cm long, lanceoloid, green, rarely
flushed with red, strigillose; free tips 1-2 (-3) mm long. Petals 2.5-3.5 cm long. Anthers
7-12 mm long. Ovary strigillose. Capsules 2.5-4 cm long; free tips of the valves indis
tinct. Chromosome number: n = 7 (7II; based on 9 individuals from 8 localities). Fig. 5.
Phenology. Flowering nearly throughout the year, from July to February, but some
ance. Floral
times as early as April.
Distribution
(Fig. 6). Oenothera elata subsp. elata has a disjunct distribution south of
the other two subspecies. It occurs in scattered localities, in open, often sandy sites such
and in openings in pine forest, 1100-2300
as fields, along streams and other watercourses,
m, ranging from the highlands
of central Mexico,
ico, Michoacain, Puebla, Queretaro,
Rica, and Panama.
and Veracruz,
including Guanajuato, Hidalgo, Mex
south to Guatemala, El Salvador, Costa
elata subsp. hirsutissima
(A. Gray ex S. Watson) W. Dietrich inW. L.
Wagner, Ann. Missouri Bot. Gard. 70: 195. 1983. Oenothera biennis var. hir
sutissima A. Gray [Mem. Amer. Acad. Arts, ser. 2, 4: 43. 1849, nomen nudum]
lb. Oenothera
ex S.Watson, Proc. Amer. Acad. Arts 8: 579, 603. 1873. Oenothera hirsutissima
1: 327. 1901 [combination
(A. Gray ex S. Watson) de Vries, Mutationstheorie
also proposed by Rydberg, Bull. Torrey Bot. Club 40: 66. 1913]. Oenothera
hookeri var. hirsutissima (A. Gray ex S. Watson) Munz, Leafl. W. Bot. 2: 157.
1939. Oenothera hookeri subsp. hirsutissima
(A. Gray ex S. Watson) Munz,
Aliso 2: 18. 1949. Oenothera elata var. hirsutissima
(A. Gray ex S. Watson)
Cronquist, Intermountain fl. 3A: 202. 1997.-TYPE: U.S.A. New Mexico: Santa
Fe Co., valley of Santa Fe Creek,
by Munz,
Oenothera
1949: GH!;
isolectotypes:
Jun 1847, Fendler
218 (lectotype,
designated
BM! FL! G! K! 2 sheets, MIN! MO!).
corymbosa Sims, Bot. Mag. 45: t. 1974. 1818, non Oenothera corymbosa
1798. Oenothera simsiana Seringe in DC., Prodr. 3: 47. 1828, nom.
Lamarck,
SYSTEMATICBOTANYMONOGRAPHS
46
a,b,CL
VOLUME 50
mm
30
.
c
FIG. 5. Oenothera
c. Mid-cauline
elata
leaf. d. Capsule.
subsp. elata
(Munz 15048,
e. Inflorescence
cult. DUSS-88-2006).
pubescence.
a. Inflorescence.
b. Rosette
leaf.
47
OENOTHERA
1997
0. elata subsp. elata
0~~~~90
*00
500 km
0
FIG.
nov. Onagra
1836
6. Distribution
spectabilis
["1835"],
nom.
of Oenothera
Spach, Nouv.
illeg. Oenothera
elata
subsp. elata.
Hist. Nat.,
Paris 4(4): 352.
var. simsiana
(Seringe) Gates,
Ann. Mus.
hookeri
[MEXICO.] cultivated from seeds in theMarquis
Rhodora 59: 16. 1957.-TYPE:
of Bath's garden at Longleats, Wiltshire, England, 1816 or 1817; no authentic
material located. [The hirsute pubescence and the distinct free tips of the valves
of the capsule in the illustration suggest that this entity represents 0. elata subsp.
hirsutissima,
and it is therefore tentatively included here.]
E. Greene, Fl. fran. 211. 1891.-TYPE: U.S.A.
Oenotherajepsonii
Co., Rio Vista,
California:
Solano
along Sacramento River, Sep 1891, Jepson s.n. (lectotype, here
[We have not seen the other collection cited by Greene.]
designated: NDG!).
Onagra macbrideae A. Nelson, Bot. Gaz. (Crawfordsville) 52: 269. 1911. Oenothera
macbrideae
(A. Nelson) Gates, Trans. Linn. Soc. London, Bot. 8: 11. 13 Jan
9: 68. 30 Jun
1913 [combination also proposed by A. Heller, Muhlenbergia
Idaho: Owyhee Co., 8 mi W of Silver City, Twilight
473 (holotype:
slopes, 1650 m, 27 Jul 1910, Macbride
RM-67650!;
isotypes: DS! F! GH! MIN! MO! NY! P! RM! 2 sheets, UC! US!
WS! WTU!; photo of NY isotype: BH!).
52: 268. 1911. Oenothera or
Onagra ornata A. Nelson, Bot. Gaz. (Crawfordsville)
1913].-TYPE:
Gulch, moist
U.S.A.
grassy
nata (A. Nelson) Gates, Trans. Linn. Soc. London, Bot. 8: 11. 13 Jan 1913 [com
bination also proposed by Rydberg, Bull. Torrey Bot. Club 40: 66. 18Mar 1913].
var. ornata (A. Nelson) Gates, Rhodora 59: 15. 1957.
Oenothera macbrideae
Oenothera hookeri subsp. et var. ornata (A. Nelson) Munz, Aliso 2: 25. 1949.
Idaho: Ada Co., Boise, dry sandy soil, 850 m, 18 Jun 1910,
262 (holotype: RM-67228!;
isotypes: GH! MIN! MO! NY!).
hookeri subsp. hewettii Cockerell, Proc. Biol. Soc. Wash. 26: 203. 1913.
TYPE: U.S.A.
Macbride
Oenothera
SYSTEMATICBOTANYMONOGRAPHS
48
Proc. Biol. Soc. Wash.
hewettii Cockerell,
Oenothera
VOLUME 50
26: 204. 1913. Oenothera
hookeri var. hewettii
(Cockerell) Gates, Rhodora 59: 16. 1957 [not validly pub
(Mutation factor in evolution 29. 1915), as assumed by Munz,
lished by Gates
Aliso 2: 31. 1949].-TYPE:
U.S.A. New Mexico: Sandoval Co., Abbott Ranch,
El Rito de los Frijoles [now Bandelier National Monument],
growing in a grove
of Populus angustifolia, Aug 1912; described from a single plant transplanted by
Cockerell
and grown in his garden at Boulder, Colorado. Cockerell states that his
to flower profusely the next [1913] season. We have seen two
collections
resulting from these seeds: cultivated in 1914, Cockerell s.n. (US
693275!); cultivated atMissouri Botanical Garden, 1914, Emig s.n. (MIN! MO!).
plant overwintered
There
is another
sheet (MO-713239)
on which
a small packet of
to represent the only
it as the lectotype.
is mounted
seeds from the original plant, and thus it can be considered
preserved
type material.
irrigua Wooton
Oenothera
Oenothera
evolution
Therefore we here designate
& Standley,
Contr. U.S. Natl. Herb. 16: 155. 1913.
hookeri var. irrigua (Wooton & Standley) Gates, Mutation factor in
29. 1915.-TYPE: U.S.A. New Mexico: Dona Ana Co., Mesilla Valley,
Jun 1906, Wooton & Standley s.n. (holotype: US-561366!).
Oenothera venusta Bartlett, Rhodora 16: 36. 1914. Oenothera
venusta
hookeri subsp. et var.
U.S.A. California: San
1949.-TYPE:
(cultivated from seeds collected by S. B. Parish
13-23
Davis
here designated: MO
(lectotype,
(Bartlett) Munz, Aliso
Bernardino Co., San Bernardino
on
16 Sep
1912),
2: 21.
1913,
8 sheets). The Davis material was found among R. Cleland's
originally at IND, now atMO. Another sheet (MO-3838403!)
contains
3838395-3838402!
vouchers,
two letters from Parish (25 Aug 1912 and 20 Sep 1912) and notes indicating that
strain 13-23 was grown from Parish's seeds.
Oenothera
venusta var. grisea Bartlett, Rhodora 16: 36. 1914. Oenothera hookeri
subsp. et var. grisea (Bartlett) Munz, Aliso 2: 29. 1949 [combination also pro
posed by Gates, Rhodora 59: 16. 1957]. Oenothera grisea (Bartlett) Rostaniski,
Feddes Repert. 96: 5. 1985.-TYPE: U.S.A. California: Riverside Co., Riverside
from seeds; "plant 358 from F. M. Reed"), 1913, Bartlett 3599 (lec
totype, here designated: MICH! mounted on 3 sheets; isolectotype: US!).
Oenothera hookeri var. angustifolia Gates, Mutation
factor in evolution 30. 1915.
Oenothera hookeri subsp. angustifolia (Gates) Munz, Aliso 2: 26. 1949.-TYPE:
U.S.A. Utah: Utah Co., Asphalt[um],
12 Jul 1894, Jones 5624 (holotype: BM!;
(cultivated
2 sheets, NY! POM! RM! US!; photo of POM isotype: BH!).
hookeri var. semiglabra Gates, Mutation factor in evolution 30. 1915.
TYPE: U.S.A. California: without further locality, 1875, Lemmon s.n. (holotype:
isotypes: DS! MO!
Oenothera
BM!).
Oenothera elata subsp. texensis W. Dietrich & W. L. Wagner, Ann. Missouri Bot.
Gard. 74: 152. 1987.-TYPE: U.S.A. Texas: Brazos Co., ca. 17 km NW of Nava
sota River Bridge on Hwy 6 in vicinity of Peach Tree Cutoff (cultivated from
seeds collected by P. M. Ortling & K. L. Intosh, 25 Oct 1978), 12 Sep 1984,
Stubbe s.n., DUSS-84-204
(holotype: MO-3326507!;
isotype: M!).
Stems 3-25 dm tall, red below or entirely red, sometimes green, strigillose with a few
appressed pustulate hairs, or strigillose with a few to numerous spreading or erect pustu
late hairs. Leaves membranous,
strigillose; bracts flat or undulate. Floral tube strigillose
or villous and glandular-puberulent,
sometimes with indistinct pustulate hairs. Floral tube
1997
OENOTHERA
49
2.5-5
(-5.5) cm long. Mature buds 2.5-5 cm long, usually narrowly lanceoloid. Sepals
green to yellowish green, red-striped or entirely red, pubescence like floral tube; free sepal
tips 2-7 mm long. Petals 3-4.7 (-5.5) cm long. Anthers 8-15 (-22) mm long. Ovary strig
or with a few pustulate hairs. Capsules 2.5-4.5
illose or villous, glandular-puberulent
(-6.5) cm long; free tips of the valves usually distinct, 0.5-2 mm long. Chromosome num
ber: n = 7
04 and 5II; 06 and 4II; 2 04 and
(71;
311;08 and 311;06, 04, and 211;010
on 122 individuals from 52 localities). Fig. 7.
and
based
211;
Phenology. Flowering primarily from July through September, but sometimes as
early as April or as late as October.
Distribution (Fig. 8). Scattered to locally common inmontane sites along streams and
in mesic meadows or along roadsides or, at lower elevations, near permanent or season
ally wet sites such as ditch banks, river banks, or fallow agricultural land, 15-3000 m,
throughout much of the western United States from Washington
and Idaho southeast to
Kansas and western Texas, and south to California, northern Baja California, Chihuahua,
Coahuila, Durango, Sinaloa, and Sonora, Mexico, with scattered populations from Custer,
Logan, and McCurtain counties, Oklahoma, and from Anderson, Brazos, and Leon coun
ties in eastern Texas.
The majority of the populations of 0. elata are here grouped into the polymorphic 0.
elata subsp. hirsutissima. Munz (1949, 1965) subdivided these plants into six subspecies
under 0. hookeri. His taxa were largely based on minor characters, such as pubescence
types: glandular-puberulent,pustulate-hirsute,and strigillose.The distributionof these
pubescence
types exhibits a geographical gradient, with one type predominating
in one
area, another in other areas, or several types occurring together in certain populations. To
delimit some of his taxa,Munz used, in addition to the hair characters, the variation in the
bracts from completely flat to undulate. He also used several size characteristics, espe
cially the length of the sepals, free sepal tips, seeds, and floral tube, as well as bract width.
By examining a much larger set of specimens than was available toMunz from through
out the entire range of this complex, we found that none of these features distinguish
highly coherent groups, and that the features often exhibit considerable intrapopulational
variation. Even inMunz's own view, several of his 0. hookeri subspecies intergraded con
siderably. We found almost complete intergradation between certain pairs of his sub
species of 0. hookeri: hirsutissima with hewettii and angustifolia; grisea with venusta;
and ornata with angustifolia. This intergradation is so great that these taxa appear to be
largely artificial. Yet, there is an imperfect clinal pattern over a considerable part of the
geographical and ecological range, with sparsely pubescent plants at higher elevations and
in the northern part of the range, and plants with denser grayish pubescence predominat
ing in the southern part of the range and at lower elevations.
The more broadly circumscribed 0. elata subsp. hirsutissima intergrades with both
of the other subspecies recognized here. The few collections of 0. elata from Durango,
are transitional between subspecies elata and hirsutissima. They have the free
Mexico,
sepal tips, reddish green stems, and capsule valve tips comparable to those of subsp. hir
sutissima, but have the more compact bud shape and pubescence of subsp. elata. In
tergradation with 0. elata subsp. hirsutissima
is discussed in the notes under 0. elata
subsp. hookeri.
The scattered populations of 0. elata subsp. hirsutissima from Oklahoma have few
to none of the longer hairs characteristic throughout much of the range and only scattered
glandular hairs, petals up to 4.7 cm long, and a taller habit, with plants up to 25 dm tall.
The extreme form in these populations occurs in the southeastern-most
location of the
SYSTEMATICBOTANYMONOGRAPHS
50
VOLUME 50
mm
a,b,c
?304
d 3O
FIG. 7. Oenothera elata subsp.hirsutissima (Ind.Sem.Vancouver1972 no. 121, Oregon, GrantCo., Blue
Mountains, cult.DUSS-88-2009). a. Inflorescence.b. Rosette leaf. c.Mid-cauline leaf. d. Capsule. e. Inflores
cence pubescence.
1997
OENOTHERA
0
~
51
0
~
0~~~~~
0
90
0
0
*
00
*
0
0
*
0
0~
00
0~~~0ol
0
*00
0.
0
0
00
0
0
0
0
0
0
0
0
00
0 ,0
0
00
:0
*00
00
0~~~0
0
0
0
0
00
000
0
0
~~~~~
0
0~~~~~~~~~~~~
ett
0.
suap
hirutism
FIG. 8.Distribution of Qenothera elata subsp.hirsutissima.
species inBrazos Co., Texas.When firstdiscovered in 1981,we described it asQenothera
elata subsp. texensis (Dietrich& Wagner 1987). This populationwas distinctive in its
largepale yellow petals (?5.5 cm), longercapsules (?6.5 cm), and leaf texturethat in cul
tivation resembles thatof 0. grandiflora.The tallhabit of theseplants also is sharedwith
0. grandiff!ora;
however,W. Stubbe's (unpubl.)studies in theexperimentalgarden atDuis
seldorf showed thattheseplants form7I andhave theAA genome andplastome I, clearly
allying themwith 0. elata. Fieldwork in 1991 byWagner revealed thatpopulationsof 0.
elata subsp.hirsutissima to thenortheast (AndersonandLeon counties) of the type local
ity of subsp. texensis also grow to over 25 dm in height.Moreover, the field studies
SYSTEMATICBOTANYMONOGRAPHS
52
VOLUME 50
that subsp.
texensis differed only from these nearby populations of subsp. hir
in having petals ca. 5 mm longer and capsules up to 2 cm longer. In fact, this
for the Brazos Co.
may be an overestimation of the differences, because themeasurements
plants were made on cultivated plants, whereas the measurements made from populations
inAnderson and Leon counties were taken in the field. Unfortunately,
the 1991 field work
showed
sutissima
failed to reveal any Oenothera populations at the type locality of 0. elata subsp. texensis
(due to habitat alteration) that could be compared in the field to these newly discovered
populations in Anderson and Leon counties. In summary, this new information has led us
to conclude that, although the plants described as 0. elata subsp. texensis represent one
end of the morphological
in petal and capsule
spectrum of 0. elata subsp. hirsutissima
size, they are no more distinctive than some of the populations elsewhere in the geo
graphical range of subsp. hirsutissima formerly given taxonomic recognition by Munz.
the adjacent populations from Leon and Anderson counties form a connecting
Moreover,
link to populations northward and westward. Therefore
morphological
and geographical
we here include 0. elata subsp. texensis within the polymorphic 0. elata subsp. hirsutis
sima.
The discovery of these populations of 0. elata in eastern Texas may provide a con
necting morphological
link between the AA genome taxa in western North America and
the BB genome, represented by 0. grandiflora,
in the southeastern United States. This
link gives additional credence to the suggestion that subsect. Oenothera originated some
where in the region of Texas to northern Mexico,
as hypothesized by Cleland (1972), or
at least in vegetation similar to that presently occupying this region (Raven & Axelrod
1978; Tobe et al. 1987; Dietrich & Wagner 1988).
lc. Oenothera
elata subsp. hookeri (Torrey & A. Gray) W. Dietrich & W. L. Wagner,
Ann. Missouri Bot. Gard. 74: 152. 1987. Oenothera hookeri Torrey & A. Gray,
Fl. N. Amer. 1: 493. 1840. Onagra hookeri (Torrey & A. Gray) Small, Bull. Tor
rey Bot. Club 23: 171. 1896. O[e]nothera
communis race biennis var. hookeri
(Torrey & A. Gray) H. Leveille, Bull. Acad. Int. Geogr. Bot. 19: 330. 1909.
biennis f. hookeri (Torrey & A. Gray) J. Boivin, Naturaliste Canad.
93: 644. 1966. Oenothera biennis var. hookeri (Torrey & A. Gray) J.Boivin, Nat
Oenothera
uraliste Canad.
94: 654. 1967.-TYPE:
U.S.A. California: 1833, Douglas
s.n.
(holotype: GH!; isotypes: BM! K!). [The GH specimen has "(15.)" after the
name D. Douglas.]
Oenothera franciscana Bartlett, Rhodora 16: 35. 1914. Oenothera hookeri var. fran
ciscana (Bartlett) Gates, Rhodora 59: 16. 1957.-TYPE: U.S.A. California: Mon
terey Co., Carmel Beach (cultivated from seeds taken from a herbarium sheet,
Smith 1063 in herb. Bartlett, collected 30 Jul 1905) (lectotype, here designated:
MICH! 3 sheets). [In the protologue Bartlett mentions that he grew this entity for
three years beginning in 1910, and that B. M. Davis at Philadelphia also grew the
Smith strain.]
Oenothera
hookeri subsp. et var. montereyensis
montereyensis
California: Monterey
Wolf 6223
Munz, Aliso 2: 14. 1949. Oenothera
Feddes Repert. 96: 5. 1985.-TYPE:
U.S.A.
Co., 0.2 mi S of mouth of Alder Creek, 100 ft, 6 Nov 1934,
(Munz) Rostan'ski,
(holotype: RSA-12778!;
isotypes: GH! 2 sheets, NY! POM! UC!).
Stems usually less than 8 dm tall, flushed with red below or entirely red, strigillose,
villous and with numerous pustulate hairs, the inflorescence and young growth also glan
1997
OENOTHERA
53
Leaves not leathery, strigillose to villous; bracts flat, villous and glan
and with some pustulate hairs.
dular-puberulent. Floral tube villous, glandular-puberulent,
Mature buds 2-4 cm long, lanceoloid. Sepals flushed with red, pubescence like floral tube
but usually with many distinct pustulate hairs; free sepal tips 1-5 mm long. Petals 2.5-4
dular-puberulent.
cm long. Anthers
12-23 mm
and with many
villous, glandular-puberulent
2.5-4.5 cm long; free tips of the valves distinct, 1-2.5
mm long. Chromosome
number: n = 7 (71I;04 and 5SI;06 and 4II; based on 16 individ
uals from 8 localities). Fig. 9.
Phenology. Flowering mostly inAugust through October, but as early as June and as
late as November.
long. Ovary
distinct pustulate hairs. Capsules
Distribution
(Fig. 10). Occurring inmoist coastal and slightly inland sandy and bluff
habitats, sea level to about 200 m, in California around San Francisco Bay along the coast
from the vicinity of Petaluma, Sonoma Co., and Point Reyes south to Santa Barbara Co.,
including Santa Cruz Island, and possibly south to San Diego Co. Other localites included
in the specimens cited probably represent introduced populations or intermediates with 0.
elata subsp. hirsutissima. The ones most similar to 0. elata subsp. hookeri are included
it pending further study [inland sites in Contra Costa (Mt. Diablo) and Napa coun
ties], while those intermediate to 0. elata subsp. hirsutissima are included there (sites in
with
Madera, Sacramento, and Yuba counties).
Our concept of 0. elata subsp. hookeri includes the strictly coastal plants with a
bushy habit, blunt buds, free sepal tips 1-2.5 mm long, and sepals usually 2-2.5 cm long,
recognized by Munz (1949, 1965) as 0. hookeri subsp. montereyensis.
Study of extensive
series of specimens in the herbarium and common garden indicate that these plants repre
sent themorphological
endpoints of a clinal variation pattern from the slightly inland pop
ulations with narrower, more attenuate buds, free sepal tips 2-4 mm long, and sepals
3-3.5
cm long, assigned by Munz
to 0. hookeri subsp. hookeri. There
tergradation represented both as intrapopulational
tain them as distinct subspecies.
and interpopulational
is far too much
in
variation tomain
Intergradation of a somewhat lesser degree was noted between 0. elata subsp. hook
eri and 0. elata subsp. hirsutissima. For example, the inland plants from Sacramento,
Madera, and Yuba counties, California, were extremely difficult to place, and are listed
here with the specimens examined of 0. elata subsp. hirsutissima. Further study of pop
in these areas is needed. Another example of intermediate plants that are difficult
to place is the experimental strain Johansen from Sutter Co., California,
in cultivation
since the early 1930's, here also listed under 0. elata subsp. hirsutissima.
ulations
2. Oenothera
jamesii Torrey & A. Gray, Fl. N. Amer. 1: 493. 1840. Onagra jamesii (Tor
rey & A. Gray) Small, Bull. Torrey Bot. Club 23: 171. 1896. O[e]nothera
com
munis race biennis var. jamesii (Torrey & A. Gray) H. Leveille, Bull. Acad. Int.
U.S.A. Oklahoma: Canadian River [some
Geogr. Bot. 19: 330. 1909.-TYPE:
where in northeastern Custer Co., downstream to near the mouth of Bear Creek,
Blaine Co.], 23 Aug 1820, James s.n. (holotype: NY!, photo BH!). [This species
is not known from Blaine Co., and thus the collection was most likely made in
Custer Co.
(locality
and date reconstructed with
aid of Goodman
and Lawson,
1995).]
Erect biennial (or winter annual) herb with a long taproot, forming a rosette; stems to
18 dm in cultivation, usually green, rarely red-flushed, unbranched or with branches aris
SYSTEMATICBOTANYMONOGRAPHS
54
VOLUME 50
*
b
?uwe& 1ce4mrJf
FIG. 9. Oenothera
leaf. c. Mid-cauline
elata
subsp. hookeri
leaf. d. Capsule.
(Hardham
e. Inflorescence
s.n., cult. DUSS-88-2010).
pubescence.
a. Inflorescence.
b. Rosette
55
OENOTHERA
1997
1250W
0. elata
/
J
subsp. hookeri
0
FIG.
50
100
10. Distribution
A
200 km
of Oenothera
elata
subsp. hookeri.
SYSTEMATICBOTANYMONOGRAPHS
VOLUME 50
from the rosette and secondary branches arising from the main
stem, exclu
56
ing obliquely
sively and densely strigillose or with some additional longer appressed hairs, rarely with
a few pustulate hairs, stem in the apical part of the inflorescence sometimes also glandu
lar-puberulent. Leaves dull green with paler veins, densely strigillose on both surfaces and
along the margins. Rosette leaves 10-30 cm long, 2.5-5 cm wide, narrowly oblanceolate
to oblanceolate, margin bluntly dentate, the teeth widely spaced, apex acute, base gradu
ally narrowed to the petiole. Cauline leaves 4-20 cm long, 1-5 cm wide, narrowly lance
olate to lanceolate or narrowly elliptic to elliptic, margin bluntly dentate, apex acute to
long-acute, base narrowly cuneate to attenuate, short-petiolate to subsessile. Bracts 3-10
cm long, 0.9-2.8 cm wide, narrowly lanceolate, margin bluntly dentate, the teeth widely
spaced, apex acute to long-acute, base obtuse to narrowly cuneate, sessile, the apical
bracts often recurved. Inflorescence unbranched or rarely interrupted by side branches.
Floral tube (6-) 8-12 (in cultivation up to 16) cm long, 1.8-2.5 mm in diameter, yellow
ish green or flushed with red, sometimes also red-maculate, exclusively
densely strig
illose, or sparsely strigillose and sparsely to densely glandular-puberulent,
persistent in
the withered state on the ovary. Mature buds 3-5 cm long, 7-12 mm in diameter, narrowly
lanceoloid to lanceoloid, bluntly quadrangular in cross section. Sepals 3-5.5 cm long,
6-10 mm wide, greenish to yellowish green, red-striped or entirely red, pubescence like
that of the floral tube; free sepal tips 0.5-3 mm long, straight in bud, strigillose. Petals 4-5
cm long, 4-5.5 cm wide, yellow, very broadly obovate, retuse. Filaments 23-30 mm long;
anthers 12-22 mm long; pollen 90-100% fertile. Ovary 1-1.5 cm long, 2.5-3 mm in di
and densely strigillose, sometimes also glandular-puberulent
ameter, exclusively
at the
apex or throughout. Style 9-17 (-20) cm long, the exserted part 3-5.2 cm long; stigma el
evated above the anthers at anthesis, the lobes 5-15 mm long. Capsules 2-5 cm long, 6-12
mm
in diameter at the base, lanceoloid,
tapering toward the apex, green, the valves with
the same as the ovary but less dense, green; free tips of the
valves conspicuous,
2.5-5 mm long, apex rounded to retuse. Seeds 1-1.2 mm long,
0.7-1.3 mm in diameter, dark brown to almost black. Chromosome number: n = 7
(711;04
and 5I; 06 and
and 311;010 and
08
based
on
11
individuals
4
from
local
0)14;
411;
211;
ities). Self-compatible, mostly outcrossing. Fig. 11.
Phenology. Flowering principally from August through October, but sometimes in
as early as July and as late as November.
populations in northern Mexico
whitish midvein,
Distribution
and other moist
pubescence
(Figs. 12, 13, 14). Occurring on sandy stream banks and along ditches,
in cultivated areas or along disturbed roadsides,
areas, or occasionally
(30-) 300-1750
m, from southern Kansas
through central Oklahoma
and Texas to
Coahuila, west-central Nuevo Le6n, and Puebla, Mexico. Oenothera jamesii is natural
ized in the Canary Islands, Japan, and South Africa.
Oenothera jamesii, like 0. elata, is a bivalent-forming
species with an AA genomic
constitution and plastome I (Stubbe 1959, 1963, 1964). All plants studied were self-com
patible, but the large flowers of this species and its elevated stigma suggest that it is usu
ally outcrossing. One of the primary autapomorphies of 0. jamesii is the long floral tube
6-16 cm long. This feature suggests that 0. jamesii was derived from 0. elata via a pol
linator shift to longer-tongued hawkmoths, such as those of the genus Manduca
(Raven et
al. 1979). Additional autapomorphies
include a floral tube persistent after anthesis, stout
capsules, and free tips of the capsule valves up to 5 mm long. Oenothera jamesii resem
bles populations of 0. elata from the same geographical area in its appressed pubescence
and conspicuous
leaf venation and margins, but the leaves are wider in 0. jamesii.
Oenothera villosa subsp. villosa has similar vegetative features.
1997
OENOTHERA
57
mm
,30
a,b,c
d
e
30
5
f:t~~~~~~~~~~~~~~~~~~~~~~f
FIG.
cauline
11. Oenothera
leaf. d. Capsule.
jamesii
(Munz 15077,
e. Inflorescence
cult. DUSS-88-2013).
pubescence.
a. Inflorescence.
b. Rosette
leaf. c. Mid
SYSTEMATICBOTANYMONOGRAPHS
58
VOLUME 50
0~~~~~~~~~~~~~~~
A
V
~ ~
~
0 500
~~~
~A
AAA
AA
A AA
A
0~~~~~~~~~~~~~~~~~
A
A
A
A
Iongissim
O.
*
AA
A
A 0.
* 0.
jamesi4
longissima
Co0
0
FIG.
12. Indigenous
distribution
500km
of Oenothera
jamesii
and distribution
of 0.
longissima.
In Kansas, Oklahoma, Texas, and Coahuila the range of 0. jamesii overlaps with that
of 0. elata subsp. hirsutissima; however, hybrids have not been detected. Its range also
overlaps with that of 0. villosa subsp. villosa; a single intermediate from near Oklahoma
City (Meyers 80, OKL) with petals within the size range of 0. villosa and a floral tube 6.2
cm long presumably represents a hybrid.
is the only bivalent-forming
subsect.
Oenothera
jamesii
species of Oenothera
Oenothera to be naturalized outside of its indigenous range. It iswell established in Japan,
itwas described as 0. suzukiana, as well as in the Canary Islands and South Africa.
The earliest collections we have seen from these areas are: 1889 in Japan (Faurie 700);
1899 in South Africa (Galpin 2585); and 1969 in the Canary Islands (Hansen s.n.). The
where
1997
OENOTHERA
//
59
C
*r'--
4 _
Dw
7'-'
;,,
-
l---
<
r~~~~~~- - - -- - -- - - - - - - - -
55?15'E 55?30'
''
j si,~~~~~~~~~~~~~~~~--------t,
-'i
' ' 1'a
'
- - - - --- - - - - - - - - - - - - - - - - - - - - - -
, j
- -- - - - - - - -
55?45'
R unio
21'S
21
300W
2t
23
Azores
10 20 30 Km
0
*
800
1200 Km
400
0 ________________
40\S \
FIG.
Africa,
13. Distribution
the Azores,
of Oenothera
biennis,
0.
jamesii,
0. parviflora,
0. biennis
*
0.
A
0. parviflora
jamesii
0. villosa subsp. villosa
and 0.
villosa
subsp.
villosa
in
and Reunion.
only strain that we have studied in the experimental garden from the Canary Islands had
a 0)14 at meiotic metaphase
I. It is not entirely clear what this represents, but it is pre
a
PTH
and the configuration
is presumably not stable. Similar plants from
sumably not
Japan that formed large rings during meiosis were selfed by Jean and Linder (1979). The
SYSTEMATICBOTANYMONOGRAPHS
60
VOLUME 50
1800E
* .jamesil
A .parvif
lora
* 0. oakesiana
0
FIG.
14. Distribution
of Oenothera
jamesii,
2000 Km
0. oakesiana,
and 0. parviflora
in Asia.
OENOTHERA
1997
cytological
configurations
showed a lack of stability; some plants formed
and others formed 7II.
ring configurations
3. Oenothera
in the progeny
61
longissima
Rydberg, Bull. Torrey Bot. Club 40: 65. 1913.-TYPE:
U.S.A.
Utah: San Juan Co., near the Natural Bridges, Armstrong orWhite Canyon, 1600
m, 4-6 Aug 1911, Rydberg & Garrett 9410 (holotype: NY!, photo BH!; isotype:
US!).
28: 22. 1922. Oenothera
Oenothera
clutei A. Nelson, Amer. Bot. (Binghamton)
longissima subsp. et var. clutei (A. Nelson) Munz, Aliso 2: 46. 1949.-TYPE:
U.S.A. Arizona: Coconino Co., War God Spring, Navajo Mtn, 2130 m, 9 Jul-24
Aug 1919, Clute 4 (holotype: RM-98480!).
Erect biennial to probably short-lived perennial herb from a taproot, forming a
rosette; stem 6-30 dm tall, unbranched or with branches arising obliquely from the rosette
and secondary branches arising from themain stem, usually flushed with red, rarely green,
exclusively densely to sparsely strigillose, or strigillose and with pustulate hairs, in the re
gion of the inflorescence
sively strigillose
sometimes
Leaves dull green, exclu
also with some erect hairs, in
also glandular-puberulent. Rosette leaves
also glandular-puberulent.
on both surfaces and margins,
sometimes
the basal region of the inflorescence sometimes
9-40 cm long, 1.4-5 cm wide, very narrowly oblanceolate to oblanceolate, margin bluntly
dentate to subentire, the teeth widely spaced, apex acute, base gradually narrowed to the
to nar
petiole. Cauline leaves 5-22 cm long, 0.8-2.5 cm wide, narrowly oblanceolate
the
or
dentate
to
very narrowly elliptic, margin bluntly
subentire,
teeth
rowly lanceolate
widely spaced, apex acute, the lower ones gradually narrowed to the petiole, the middle
and apical ones narrowly cuneate to attenuate at base, short-petiolate to sessile. Bracts 2-5
cm long, 0.3-1 cm wide, narrowly lanceolate, margin bluntly dentate to subentire, apex
acute, base acute to narrowly cuneate. Inflorescence unbranched, lax. Floral tube 6-13.5
cm long, 1.3-2 mm
in diameter, yellowish
green, flushed with
some red to entirely red,
and sparsely villous, sometimes also with
exclusively
strigillose, or glandular-puberulent
some pustulate hairs. Mature buds 2.3-4.7 cm long, 5-9 mm in diameter, narrowly lance
oloid or cultrate to narrowly oblong. Sepals 2.5-5.5 cm long, 4-8 mm wide, yellowish
like floral tube; free
green, flushed with some red or entirely red to dark red, pubescence
sepal tips 2-6 mm long, strigillose to villous, erect in bud. Petals 2.8-6.5 cm long, 3.2-6
cm wide, very broadly obovate, retuse, pale yellow to yellow. Filaments 20-40 mm long;
anthers 14-20 mm long; pollen 90-100% fertile. Ovary 1.2-2 cm long, 2-2.5 mm in di
ameter, pubescent in one of three ways: a) densely strigillose; b) strigillose, glandular-pu
berulent,
and with pustulate
longer hairs; or c) glandular-puberulent
and with pustulate
longer hairs. Style 9-18 cm long, the exserted part 3-5.5 cm long; stigma elevated above
the anthers at anthesis, the lobes 5-9 mm long. Capsules 2.5-5.5 cm long, 4-9 mm in di
ameter, narrowly lanceoloid to lanceoloid, tapering toward the apex, pubescence like that
of ovary but less dense, green, often red-striped; free tips of the valves distinct or indis
tinct, 1-2 (-3) mm long, truncate to emarginate. Seeds 1.1-1.9 mm long, 0.6-1.2 mm in
04 and 5II; 06
number: n = 7
diameter, dark brown to almost black. Chromosome
(711;
and
08 and
or 2 04 and
based on 28 individuals from 7 localities). Self-com
411;
311;
311;
patible, mostly outcrossing. Fig. 15.
Phenology. Flowering from July through September, rarely in October.
Distribution (Fig. 12). Occurring in at least seasonally moist sites, usually in sandy or
sandy loam soils, sometimes
in sites with high alkalinity
or associated
with
limestone,
SYSTEMATICBOTANYMONOGRAPHS
62
VOLUME 50
mm
a,b,c
d
Q ... I.
t- ---X -------
id
FIG.
Mid-cauline
15. Oenothera
leaf d. Capsule.
longissima
(Keliher
e. Inflorescence
s.n., cult. DUSS-88-2014).
pubescence.
a. Inflorescence.
b. Rosette
leaf. c.
OENOTHERA
1997
63
typically along desert washes, streams, seeps, and roadsides, 850-2800 m, from western
Colorado (Delta and Montezuma
counties) through southern Utah, northern and western
Arizona, to southeastern and eastern Nevada and southern California (Inyo, Los Angeles,
and San Bernardino counties).
Oenothera
is one of five outcrossing
longissima
bivalent-forming
species of
Oenothera subsect. Oenothera. Like 0. elata and 0. jamesii, it has an AA genome and
plastome I (Stubbe 1959, 1963, 1964). Itmay have been derived from populations of 0.
elata subsp. hirsutissima via a shift to long-tongued pollinators, such as those of the genus
Manduca
(Raven et al. 1979).
In contrast
to 0. jamesii, which
has several autapomor
phies, the only autapomorphy of 0. longissima is its long floral tube (6-13.5 cm long) and
associated pollinator spectrum; otherwise, it differs in no essential way from certain pop
ulations of O., elata subsp. hirsutissima.
Munz (1949, 1965) distinguished
two subspecies of 0. longissima, the eastern subsp.
longissima and the western subsp. clutei. The former taxon was characterized by ap
pressed pubescence, whereas the latter featured erect to spreading hairs as well as short
glandular
hairs. Our study of considerably more material than was available to Munz
that all pubescence
types occur in both the eastern and western portions of the
range, sometimes expressed as intrapopulational variation. The situation is analogous to
the pubescence variation in 0. elata subsp. hirsutissima; both possibly represent rather
shows
For example, a collection
from San Juan Co., Utah (Welsh
20844, MO), which represents the eastern part of the range (appressed pubescence),
ex
pressed all three pubescence types in cultivation.
simple genetic
situations.
Because
0. longissima could grow sympatrically with 0. elata subsp. hirsutissima
are expected; however, because of the great morphological
similarity between
these taxa, it would be difficult to detect hybridization. The only known case of putative
hybridization is one in which seeds of plants from Coconino Co., Arizona, yielded con
hybrids
siderable variation in floral tube length (DUSS-77-092,
DUSS-77-091,
DUSS-76-0104):
13.2 cm, 9.0 cm, and 6.0 cm long respectively. Another collection of 0. elata subsp. hir
sutissima from the same county (DUSS-76-065)
consistently yielded plants with floral
tubes ca. 5 cm long. The plants with 6 cm long floral tubes may represent hybrids. More
over, cytological investigations would be of little help since there is a diversity of config
urations in both taxa. Additional
field studies may resolve the delimitation of these taxa
more
clearly.
wolfii (Munz) P. H. Raven, W. Dietrich & Stubbe, Syst. Bot. 4: 244. 1980
["1979"]. Oenothera hookeri subsp. et var. wolfli Munz, Aliso 2: 16. 1949.
TYPE: U.S.A. California: Humboldt Co., Redwood Hwy roadside just S of
Trinidad, 11 Oct 1934, Wolf & Johnson 6172 (holotype: RSA-12706!;
isotypes:
NY! POM! US!).
4. Oenothera
Erect biennial herb with a taproot, forming a rosette; stems 5-10 dm tall, rarely taller,
unbranched or branched from the rosette, the branches arcuating or obliquely arising from
the rosette, themain stem sometimes with additional secondary branches, flushed with red
or green in the region of the inflorescence, densely strigillose and with many spreading to
subappressed pustulate hairs, in the region of the inflorescence
puberulent. Leaves dull green, densely strigillose to villous on
Rosette leaves 13-35 cm long, narrowly oblanceolate, margin
part of the leaf, and bluntly dentate with widely spaced teeth
also villous and glandular
both surfaces and margins.
irregularly dentate in distal
to sinuate in the proximal
SYSTEMATICBOTANYMONOGRAPHS
64
VOLUME 50
part, apex acute, base gradually narrowed to the petiole. Cauline leaves 5-18 cm long, 1-4
cm wide, narrowly lanceolate or very narrowly elliptic to elliptic, the lower leaves with
margins like rosette leaves, margins of the apical half of plant dentate to subentire, apex
acute, base narrowly cuneate to attenuate, short-petiolate to sessile. Bracts 2-9 cm long,
0.5-3
cm wide,
on
narrowly lanceolate to narrowly ovate, often glandular-puberulent
entire to weakly dentate, apex acute, base obtuse to narrowly
Inflorescence unbranched. Floral tube 3-4.6 cm long, 0.8-1.1 mm in di
lower surface, margin
sessile.
cuneate,
flushed with red, densely long-villous, often some of these hairs pustulate,
also glandular-puberulent. Mature buds 1.7-2.5 (-3) cm long, 5-8 mm in diameter, lance
oloid. Sepals 1.7-2.8 cm long, 4-6.5 mm wide, yellowish green, and usually flushed with
red or red-striped, pubescence
like that of the floral tube; free sepal tips 1-3 mm long,
ameter, usually
erect in bud. Petals 1.3-2.3 cm
strigillose, sometimes also glandular-puberulent,
shorter than
long, 1.4-2.5 cm wide, very broadly obovate, retuse, yellow, conspicuously
the sepals. Filaments 12-20 mm long; anthers 7-12 mm long; pollen ca. 50% fertile.
densely
Ovary 0.7-1.2 cm long, 1.5-1.8 mm in diameter, very densely long-villous, some of the
cm long, the
hairs pustulate, also strigillose and glandular-puberulent.
Style 4.3-5.8
exserted part 1.4-2 cm long; stigma usually slightly elevated above the anthers or sur
rounded by them, which shed pollen directly onto the lobes at anthesis, the lobes 3-9 mm
long. Capsules 3-4.8 cm long, 5-7 mm in diameter, narrowly lanceoloid to lanceoloid, ta
pering toward the apex, dark dull green when fresh, usually red-striped, pubescence like
that of ovary but less dense; free tips of the valves distinct, 0.9-1.5 mm long, rounded to
slightly retuse. Seeds 0.9-2 mm long, 0.9-1.3 mm in diameter, dark brown. Chromosome
number: n = 7 (014; based on 11 individuals
autogamous, PTH. Fig. 16.
from 9 localities).
Self-compatible,
usually
Phenology. Flowering from June through October.
Distribution
(Fig. 17). Rare in coarse-textured
sandy or, in Oregon, rocky sites, on
coastal dunes and bluffs, or loose rocky sites, and sometimes sandy sites along roads;
along the Pacific coast from the vicinity of Port Orford, Curry Co., Oregon (currently ap
parently only as far north as Otter Rock), south in a scattered distribution through Del
Norte Co. to the mouth of Mattole River, Humboldt Co., California. The distribution, at
least in California, according to D. Imper (pers. comm.) is closely associated with small
patches of Cenozoic-age marine sediments,
and metamorphic
rocks. Moreover,
mentary
mouths
isolated from each other by Franciscan
most populations
or to the south of a headland. The largest populations
sedi
appear to occur near river
center in the area about 11
km long in the vicinity of Crescent City in Del Norte Co., between Point George and En
derts Beach in Redwood National Park. There are collections from two inland California
localities, one at the eastern border of Humboldt Co., California (Willow Creek, Trinity
and the other at Carville, Trinity Co., thatmay be 0. wolfii. If so, they would
represent recent introductions and should be studied further.
Oenothera wolfii is a rare endemic of coastal habitats and known from about 20 dif
ferent sites (Skinner & Pavlik 1994). The total number of individuals of 0. wolfii appar
River Valley)
presumably
ently fluctuates, with perhaps no more than about 5000 individuals total at any one time.
It is threatened by any potential development
and alteration of its habitat, presently by
road maintenance
and foot traffic (Skinner & Pavlik 1994). Another threat comes from
the recent spread of 0. glazioviana
ulations
observed,
probably
through hybridization
to this area. Oenothera
glazioviana
and perhaps by direct competition.
could swamp pop
D. Imper (pers. comm.)
during a detailed field survey of 0. wolfii in 1987, that the greatest threat
comes from hybridization with 0. glazioviana
rather than habitat alteration.
1997
OENOTHERA
65
mm
a,bsc XI30Q_ J
e~
~
~
~
.
a
FIG.
cauline
16. Oenothera
leaf d. Capsule.
wolfii (Hoch
e. Inflorescence
1853,
cult. DUSS-88-2025).
pubescence.
~~~~~~b
a. Inflorescence.
b. Rosette
leaf. c. Mid
SYSTEMATICBOTANYMONOGRAPHS
66
+0
VOLUME 50
0
1250W
0. wolfii
FIG. 17.Distribution of Oenotherawolfi.
Oenothera wolfi hybridizes with 0. glazioviana
throughout most of its range in northern
California. The hybrid populations occur adjacent to populations of both parents, but in
more disturbed sites, preferring gravelly roadsides. Hybrids rarely occur in the same ex
posed strand or bluff habitats as 0. wolfii. The hybrid appears to be more aggressive than
0. wolfii, and large populations have developed in the median strip of Highway 101 south
of Trinidad near Clam Beach, the south end of the Klamath River bridge along Highway
101, and between Smith River and the Oregon border. In 1982, Imper (pers. comm.) esti
mated the first two populations at fewer than 100 individuals, but by 1984 the Clam Beach
OENOTHERA
1997
population
had greatly
increased
to a population
67
in the thousands. The hybrid may have
represent a newly evolved phenotype that
stabilized, and in effect may
itself. Further study is needed to examine this hypothesis. There is urgent
need to preserve the integrity of 0. wolfii by collecting seeds from as many populations
as possible and preserving them in seed banks.
become
somewhat
can propagate
Oenothera wolfii is ranked by the California Native Plant Society as a category lB
species (Skinner & Pavlik 1994). This category was established for plants that are rare,
threatened, or endangered in California and elsewhere. All lB species meet the definitions
of Sec. 1901, chapter 10 (Native Plant Protection) of the California Department of Fish
and Game Code, and are eligible for State listing. Oenothera wolfii was given the highest
ranking for rarity and endangerment, but only the second highest ranking for distribution,
it is not endemic to California. This species ought to be listed as an endangered
species federally, under the provisions of the Endangered Species Act.
because
Oenothera wolfii is a PTH species with an AA genomic constitution and plastome I
(Wasmund & Stubbe 1986). It was originally described as a subspecies of 0. hookeri by
Munz (1949, 1965), but in the 1970's it was discovered to be a PTH species (Wasmund
1980; Wasmund & Stubbe 1986). It is therefore treated at the species level in parallel with
the other AA genome PTH species, 0. villosa.
0. wolfii is very similar to 0. elata subsp. hookeri, but differs from
Morphologically,
it primarily in several features of the flower, most of which correspond to the evolution of
PTH. The petals are conspicuously
smaller in 0. woflfi (1.3-2.3 cm vs. 2.5-4 cm long),
but the sepals are disproportionately
long (1.7-2.8 cm vs. 2-4 cm long) relative to the
petals. This feature gives the flowers a unique appearance with the petals as little as half
as long as the sepals. Typically the stigma is surrounded by the anthers in an autogamous
PTH plant, but the lobes are slightly elevated in 0. wolfii. Despite this the pollen is still
shed more or less directly onto the stigma. The anthers of 0. wolfii are also smaller than
those of 0. elata subsp. hookeri (7-12 mm vs. 12-23 mm long).
Genetically, both of the A complexes of 0. wolfii are nearly identical. They both are
closely related to the neighboring maritime ecotype of 0. elata subsp. hookeri (Wasmund
& Stubbe 1986). The work by Wasmund and Stubbe showed that both complexes, when
crossed to any other genotype, produced phenotypes so much alike that the twin hybrids
can only be distinguished by their diakinesis. The a (egg) complex is not transmitted by
the pollen. From 49% to 62% of the pollen consists of empty grains. This indicates an in
activation of one of the two complexes by an Si-allele acting as a gametophytic
lethal.
By contrast, the , (pollen) complex is transmitted by both the pollen and the ovule.
The ,B,3homozygotes
are believed to be eliminated on selfing by the evolution of a sporo
phytic lethal. This is presumably the reason that 14-45% abortive seeds are produced
when plants of 0. wolfli are selfed. The variation in the percentages of abortive seed may
be explained by differences in the degree of embryo sac competition between the com
plexes.
The
study by Wasmund and Stubbe also investigated the end arrangements of the
in both complexes by analysis of diakinesis configurations in hybrids with
chromosomes
a standard. The f3complex of the strain Luffenholtz differs from the 3 complex of strain
Mendocino
by two reciprocal interchanges. The other strains in experimental cultivation
have not yet been studied.
Munz (1949) gives a wider range for 0. wolfii than we do here. The collections from
Washington
confused him, and he only tentatively included them under 0. wolfii. He in
cluded several collections that we have assigned elsewhere: Marin Co., California, How
SYSTEMATICBOTANYMONOGRAPHS
68
VOLUME 50
Co., California, Butler 1799 and Brown
485 (=O. villosa subsp. strigosa); Trinity Co., California, Hall 8695 (=O. villosa subsp.
strigosa); Jackson Co., Oregon, Hammond 142 (=O. villosa subsp. strigosa); Klickitat
Co., Washington,
Suksdorf 2066, 4765, 5859, 7807, 10603 (=O. biennis x 0. villosa
subsp. strigosa; not 0. villosa as suggested by Raven et al., 1979). Individuals from the
two inland California localities discussed in the paragraph on distribution may also prove
ell 23027
(=O. elata subsp. hookeri);
Siskiyou
to be like these, and further studies are necessary
to determine unequivocally
what entity
they represent.
SOUTH AFRICA. "e Cap.
5. Oenothera
villosa Thunberg, Prodr. fl. cap. 75. 1794.-TYPE:
b. Spei," between Apr 1772 and Mar 1775, Thunberg s.n. (holotype: UPS!). [The
mention
of the holotype of this species at S by Dietrich
and Raven
(1976) was
in
error.]
Erect biennial herb with a taproot, forming a rosette; stems 5-20 dm tall, unbranched
or with branches obliquely arising from the rosette or from themain stem, green or flushed
with red in lower part or entirely red, pubescent with one of the following patterns: a)
densely strigillose with numerous to a few long appressed to subappressed hairs; b) same
as in (a) but with numerous to few subappressed to erect pustulate hairs; or c) strigillose
in the
long appressed to spreading red-pustulate hairs, also glandular-puberulent
inflorescence. Leaves dull green to grayish green, veins inconspicuous or pale green,
sometimes red, margins dentate to denticulate or subentire, the teeth sometimes widely
and with
spaced, the lower part sometimes
surfaces
and margins,
sinuate-dentate,
rarely villous,
sometimes
undulate,
the apical bracts usually
strigillose
on both
in
glandular-puberulent
subsp. strigosa. Rosette leaves 10-30 cm long, 1.2-4 (-5) cm wide, narrowly oblanceo
late to oblanceolate,
apex acute, base gradually narrowed to the petiole. Cauline leaves
5-20 cm long, 1-2.5 (-4) cm wide, the lower ones similar in shape to the rosette leaves,
those toward the apex narrowly lanceolate to lanceolate, narrowly elliptic or elliptic, apex
acute, base obtuse
to narrowly cuneate, sessile. Bracts 2-7 cm long, 0.5-1.5
(-2.8) cm
(or up to 3 cm wide in cultivation), in an oblique or right angle to the stem, some
times the tips bent down, narrowly lanceolate to narrowly ovate or narrowly elliptic, mar
gins conspicuously
dentate to subentire, apex acute to narrowly acute, base rounded to
wide
acute, sessile. Inflorescence
mm in diameter, yellowish
unbranched, dense to lax. Floral tube 2.3-4.4 cm long, ca. 1
or flushed with red, very densely to sparsely strigillose and
with numerous to few longer appressed to subappressed hairs, and usually only in subsp.
strigosa also glandular-puberulent
and often with some pustulate hairs. Mature buds
cm long, 3-5 mm in diameter, lanceoloid to narrowly oblong or oblong. Sepals
cm long, 2.5-4.5 mm wide, yellowish green, red-striped or flushed with red, pu
bescence the same as the floral tube; free sepal tips 0.5-3 mm long, strigillose, erect in
bud. Petals 0.7-2 cm long, 0.8-2.1 cm wide, very broadly obovate, retuse to emarginate,
0.8-1.8
0.9-1.8
yellow
to pale yellow. Filaments
7-15 mm
long; anthers 4-10 mm long; pollen ca. 50%
fertile. Ovary 0.7-1.4 cm long, 1.5-2 mm in diameter, very densely to densely strigillose,
also with longer appressed to subappressed hairs, and in subsp. strigosa usually also glan
and with subappressed red-pustulate hairs. Style 3-5.5 cm long, the
dular-puberulent
exserted part 0.3-1.4 cm long; stigma surrounded by the anthers, which shed pollen di
rectly onto the lobes at anthesis, the lobes 3-9 mm long. Capsules 2-4.3 cm long, 4-7 mm
in diameter, lanceoloid, tapering toward the apex, pubescence
like that of ovary but less
dense, grayish green to dull green, red-striped or with whitish green midvein; free tips of
OENOTHERA
1997
the valves usually
69
less than 0.5-1 mm
long, truncate to retuse. Seeds 1-2 mm
to almost black. Chromosome number: n = 7 (014
indistinct,
mm
in diameter, brown
and 1II; based on 39 individuals
long, 0.5-1.2
or rarely 012
ally autogamous
Phenology.
from 36 localities). Self-compatible,
usu
in 0. villosa subsp. villosa), PTH.
(and often cleistogamous
Flowering in July and August, sometimes into September, rarely as early
as June.
Distribution. Occurring in at least seasonally moist open or disturbed sites, such as
stream or ditch banks, meadows,
bottom lands, fields, and roadsides, 30-3150 m. The
original natural range of this species was presumably from southern British Columbia
south to California and east through the Rocky Mountain and the Great Plains regions. It
now occurs eastward as far as eastern Quebec south throughout most of the eastern half
of the United
States, except for extreme southern and southeastern parts. The occurrences
in the eastern and southern portions of the range appear to represent extensions of the dis
tribution during the past several hundred years. Oenothera villosa is subdivided into two
subsp. strigosa occurring primarily in the Pacific Northwest southeast through
and subsp. villosa primarily found from the eastern foothills of the
subspecies,
the Rocky Mountains,
Rocky Mountains
eastward
throughout
the Great Plains region. Both
these regions, and subsp. villosa
cally beyond
is naturalized
in many
taxa occur sporadi
other parts of the
world.
Oenothera villosa is a PTH species with an AA genomic constitution and plastome I
(Stubbe 1959, 1963, 1964). The stigma is always surrounded by the anthers at anthesis in
0. villosa, and the pollen is shed directly onto the lobes. In fact, 0. villosa subsp. villosa
is the only taxon in subsect. Oenothera that occasionally has cleistogamous
flowers. De
spite the strong autogamy, outcrossing
ation patterns observed
sometimes occurs and has contributed
in this very widespread
species.
to the vari
Oenothera
villosa is heterogamous. The ox (egg) complex is transmitted only through
the v (pollen) complex only through the pollen. Out of 42 strains analyzed by
the ovules,
Cleland
often
(1972, p. 287) only 9 were
transmitted
through the pollen. Crossing
Cleland
complexes
there were
for which
ments.
exceptions.
In the exceptions,
both complexes
through the egg, and in two strains both complexes
also made
studies by Dietrich
extensive
analyses
inDusseldorf
of the segmental
were
agree with Cleland's
arrangements
were
transmitted
results.
found in both
(summarized in 1972, pp. 288-293). Among the 42 strains that he analyzed
14 different end arrangements in the f3complexes. By contrast, the 39 strains
the oc complexes were analyzed expressed far more variation in end arrange
In all, Cleland
discovered
potheses of the relationships
27 different arrangements
among them and the migration
figures 18.18 and 18.19 of his 1972 book. Cleland
tions from a center of origin inMexico
hybridization
between members
complex, while Population
gin for 0. villosa. We
suggest
and Central America,
of Populations
3 provided
believed
in the ox complexes.
pattern followed
that there were
His hy
are given
in
four migra
and that 0. villosa arose via
3 and 4. Population
4 contributed the cx
the ,Bcomplex. Our studies suggest a different ori
that the various phenotypes
within
0. villosa had several
independent origins from 0. elata.
The oc and ,3complexes of 0. villosa are genetically somewhat different, resulting in
phenotypes when they are present in homozygous conditions; both conditions
represent distinct A-genomic
complexes. Morphological
comparison suggests that 0. vil
losa has been directly derived from 0. elata. The accumulation of reciprocal transloca
tions and the acquisition of balanced lethals, similar to that proposed by Wasmund and
distinctive
70
SYSTEMATICBOTANYMONOGRAPHS
VOLUME 50
(1986) for 0. wolfii, may have resulted in the evolution of PTH populations from
0. elata in several different geographical areas.
The AA genome plastome I PTH forms derived from 0. elata, with the exception of
the very distinctive 0. wolfii, are treated here as two subspecies of 0. villosa. These enti
ties have been treated under various names in the past, but most authors have considered
Stubbe
them, as we do here, as a single taxonomic species. The name most frequently used for
this species has been 0. strigosa, which was delimited in the same sense as 0. villosa is
here. There are actually three older specific names for this complex than 0. strigosa: 0.
(1794), 0. erosa (1824), and 0. depressa (1891). The identity of 0. villosa, the
subsect. Raimannia by
name, was discovered only during study of Oenothera
Dietrich and subsequently aligned with the current group (Dietrich & Raven 1976). Sur
villosa
oldest
this species was originally described from a population naturalized and well es
tablished by 1820 in the Cape region of South Africa (Dietrich & Raven 1976). Oenothera
villosa also has been named 0. canovirens inNorth America, and 0. bauri, 0. hungarica,
prisingly,
0. renneri, and 0. salicifolia in Europe. At times 0. villosa has been included in 0. bien
nis without infraspecific recognition (e.g., Welsh 1986), or treated as an infraspecific
taxon of 0. biennis, most commonly as 0. biennis var. canescens (e.g., Gleason & Cron
quist 1963, 1991), even though the application of the name had not been clarified by lec
totypification until this publication.
In 1965, Munz established a new classification
for 0. villosa, under the name 0.
strigosa. He subdivided it into three subspecies: strigosa, canovirens, and cheradophila.
His subdivisions are basically parallel to the three major derivations discussed in the chap
terOrigins above. The treatment presented here accepts the second subspecies in the same
sense as Munz, under the name 0. villosa subsp. villosa, whereas the other two are com
bined under the name 0. villosa subsp. strigosa.
Our study of a full series of herbarium specimens from Oregon and Washington,
as
well as field studies by Wagner, indicates that 0. strigosa subsp. cheradophila should not
be maintained. Oenothera strigosa subsp. cheradophila was characterized by Munz as
plants with only appressed hairs, none of the hairs pustulate, and weakly angled or terete
stems. Other than these differences,
the plants treated by him as 0. strigosa
are very similar to other populations we treat as 0. villosa subsp.
subsp. cheradophila
younger
strigosa. The principal reasons for grouping plants with this phenotype with 0. villosa
subsp. strigosa instead of according them formal recognition are: 1) plants with this phe
notype do not have a geographical or ecological
range distinct from that of 0. villosa
subsp. strigosa, but rather represent an east-west clinal trend in Oregon and Washington;
2) plants with a very similar phenotype also occur in Nevada and British Columbia; 3)
within the full geographical area of these two entities there is extensive intergradation be
tween individuals and populations assigned by Munz to 0. strigosa subsp. strigosa and 0.
strigosa subsp. cheradophila
(e.g., Peck 9758 from Deschutes Co., Oregon).
the distinctive morphological
Given
features and presumed independent origins
within different populations of 0. elata of the two subspecies of 0. villosa, itwould seem
at first logical to treat them in parallel fashion to the other AA genome plastome I PTH
taxon in western North America, 0. wolfii. There are two reasons why they are here
grouped as subspecies of 0. villosa. First, these two entities apparently have evolved from
different populations of a single subspecies, 0. elata subsp. hirsutissima. Moreover,
there
has been very extensive secondary intergradation and evolution of intermediate pheno
types that cover a large geographical area across a broad contact zone along the eastern
foothills of the Rocky Mountains
and adjacent western plains. In contrast, 0. wolfii
71
OENOTHERA
1997
evolved from 0. elata subsp. hookeri and does not have any intermediate phenotypes with
other AA combination PTH taxa. The intergradation between the subspecies of 0. villosa
is also different from the more localized and patchy pattern resulting from hybridization
between 0. villosa and0. biennis, 0. oakesiana, or 0. parviflora.Within 0. villosa there
two very closely related taxa that share the same genome
the
intergradation
between 0. villosa and other PTH species is lim
and plastome, whereas
different
genome and plastome. Our delimitation of
ited in extent and between taxa with
0. villosa follows our basic approach of giving specific status to groups throughout
is broad intergradation between
that share derived morphological
features, a common
subsect. Oenothera
genome, and plastome type.
The extensive intergradation between the two subspecies of 0. villosa occurs across
and in their foothills. Despite
the area of high plains to the east of the Rocky Mountains
Oenothera
their usual autogamy both taxa occasionally
bridization,
termediate forms occupy
In subsequent generations
outcross.
the ecologically
and the plains habitat of subsp. villosa. The
of subsp. strigosa
after hy
have apparently arisen. These in
transitional areas between the montane habitat
intermediate phenotypes
new true-breeding
intermediate
phenotypes
and pustulate
usually have the oblong bud, reddish green sepals, glandular-puberulence,
hairs of subsp. strigosa; the congestion of the inflorescence, and later the infructescence,
ismore like that
is intermediate between the subspecies; the dense appressed pubescence
of subsp. villosa, except with the addition of the hair types of subsp. strigosa as noted
above; and the free tips of the capsule valves are somewhat intermediate, but nearly as
short as in subsp. villosa. The intermediate forms occur everywhere the two subspecies
and often they are more frequent; however, they seem to be the only
present in the Colorado counties of Boulder, Denver, Douglas, El Paso, and
and eastward across much of North and South
Larimer, north to southeastern Wyoming,
subspecies they appear
Dakota. These intermediates have been grouped with whichever
come
into contact,
phenotype
to resemble most
closely.
KEY TO THE SUBSPECIES OF OENOTHERA VILLOSA
1. Sepals
green
shorter
bracts
truncate;
narrowly
Plains
green; plants
than the capsules;
(1.5-)
leaf venation
naturalized
region, widely
to dull green,
gray
a few subappressed
with
hairs on the floral tube; margins
glandular
apex
to yellowish
sometimes
illose,
of leaves
2 times
conspicuously
than wide;
longer
prominent,
pubescence
pale green,
in eastern North America,
of
the
Europe,
South America,
and South
5a. 0. villosa subsp.villosa.
usually
strigillose,
yellow
of pustulate
teeth often widely
longer
dense,
the infructescence
on the lower surface; Great
especially
Asia,
type, strig
parts or a few
inflorescence
dentate;
internodes
Africa.
1. Sepals
of one
primarily
long hairs on the vegetative
to spreading
than wide;
tion inconspicuous;
flushed with
hairs,
spaced;
red, or red; plants
and of glandular
inflorescence
intemodes
relatively
of the infructescence
Rocky Mountain
region
to dull green, pubescence
green
hairs; margins
open,
of leaves
the apex broadly
longer
denticulate
obtuse;
of three types:
to subentire,
bracts (2.5-)
than or as long as the capsules;
and Pacific Northwest,
the
3 times
leaf vena
not naturalized.
5b. 0. villosa
subsp.
strigosa.
5a. Oenothera
villosa subsp. villosa.
erosa J. Lehmann, Sem. hort. bot. Hamburg 1824: 20. 1824; Linnaea 3
(Litt.): 8. 1828. Onagra lehmanniana Spach, Nouv. Ann. Mus. Hist. Nat., Paris
communis race erosa (J.
4(4): 354. 1836 ["1835"], nom. superfl. O[e]nothera
de
330. 1909.-TYPE:
Bull.
Acad.
Int.
Bot.
19:
H.
Geogr.
Leveille,
Lehmann)
Oenothera
scribed from plants cultivated
in the Botanical Garden
atHamburg,
the seeds col
SYSTEMATICBOTANYMONOGRAPHS
72
lected in the Cape region of South Africa
by M. Thalwitzer.
Onagraceae
No
authentic material
VOLUME 50
and said to have been sent to Hamburg
located; it is not known to whom the
of the Lehmann Herbarium were
sold.
salicifolia Desfontaines
[Tabl. ecole bot., ed. 2: 271. 1815, nomen nudum]
ex G. Don, Gen. Syst. 2: 685. 1832, non Oenothera salicifolia J. Lehmann, 1824,
nec Oenothera salicifolia Desfontaines
ex Seringe, 1828.-TYPE:
No authentic
Oenothera
material
Oenothera
located; disposition
var. canescens
biennis
Oenothera muricata
based on description.
Torrey & A. Gray, Fl. N. Amer.
1: 492. 1840.
(Torrey & A. Gray) B. L. Robinson, Rhodora
communis race biennis f. canescens
(Torrey & A.
var. canescens
10: 34. 1908. O[e]nothera
Bull. Acad.
Gray) H. Leveille,
Int. Geogr. Bot.
19: 330. 1909. Oenothera
parvi
flora var. canescens
(Torrey & A. Gray) Farwell, Amer. Midl. Naturalist 8: 274.
1923.-TYPE:
[U.S.A. Missouri: Jackson Co.], Independence, [1050 ft, after 24
Sep 1846], Fendler 220 (neotype, here designated: MO-2529730!).
No locality
or material was cited by Torrey and Gray, and we have not been able to locate
any apparently authentic material at BM, GH, NY, or PH. There are only two col
lections that we have seen labelled "O. biennis var. canescens." The first is a
sheet of 0. villosa subsp. strigosa collected by the Rev. H. H. Spalding (GH)
from what
is now western
Idaho. This collection
almost certainly was not made
to information inMcKelvey
(1955, pp. 824-829). The
is the one selected here as the neotype. It is the only specimen lo
cated that Gray annotated and cited (1849) as 0. biennis var. canescens, and it
before
1846 according
other specimen
also corresponds
Oenothera
closely
to the original description.
depressa E. Greene,
Pittonia 2: 216. 1891. Onagra depressa (E. Greene)
Small, Bull. Torrey Bot. Club 23: 170. 1896. Oenothera strigosa var. depressa
(E. Greene) Gates, Monogr. Biol. 7: 46. 1958, nom. illeg. [when combining these
two taxa, both originally published at the species level, the older epithet "de
pressa" should have been used for the species, ICBN (1994) Art. 11.4].-TYPE:
U.S.A. Montana:
Co., near Custer (cultivated at Berkeley, Califor
from J. W. Blankenship),
1891, Greene s.n. (holotype: UC
20459!; isotype: US!). [B. Hellenthal
(pers. comm.) indicates that there is no
specimen of this collection inGreene's herbarium in NDG.]
Yellowstone
nia, from seeds
Onagra
hungarica
Borba's, Magyar
Borba's, Kert 1902: 204. 1902. Oenothera hungarica
(Borbas)
Bot. Lapok 2: 246. 1903. Oenothera muricata subsp. hungar
ica (Borba's) Soo, Acta Biol. Acad. Sci. Hung.
3: 226. 1952. Oenothera strigosa
(Borba's) Love & L6ve, Opera Bot. 5: 258. 1961.-TYPE:
HUNGARY. Budapest,
in sandy places, 10 Jul 1902, de Borbas s.n. (lectotype,
here designated: BP-67336!;
isolectotype: BP!; photo of destroyed B sheet at
subsp. hungarica
MO!).
Oenothera
canovirens Steele, Contr. U.S. Natl. Herb. 13: 365. 1911. Oenothera
strigosa subsp. canovirens (Steele) Munz, N. Amer. Fl., ser. 2, 5: 136. 1965.
TYPE: U.S.A. Illinois: Morgan Co., along St. Louis division of Chicago, Burling
ton and Quincy Railroad, 2 mi S of Concord, 14 Aug 1910, Steele s.n. (holotype:
[The publication indicated the date as "20 Aug."]
US-618797!).
Oenothera
cockerellii
Oenothera
Bartlett
ex de Vries, Gruppenweise
56. 1913. Oenothera
Gates, Rhodora
strigosa
59: 15. 1957.-TYPE:
var. cockerellii
U.S.A.
Colorado:
Artbildung
d. Gattung
(Bartlett ex de Vries)
Boulder
(cultivated
at
OENOTHERA
1997
73
from seeds sent by T. D. A. Cockerell);
fig. 19, p. 53 in de Vries,
1913 (lectotype, here designated). [No authentic material located.]
Oenothera hookeri var. parviflora Gates, Mutation
factor in evolution 29. 1915.
TYPE: CANADA. British Columbia: Kamloops,
19 Jun 1889, Macoun s.n. (holo
Amsterdam
type:BM!).
Oenothera bauri Boedijn, Z. Indukt. Abstammungs-Vererbungsl.
32: 360. 1924.
TYPE: GERMANY. Brandenburg: Berlin-Friedrichshagen
(cultivated from seeds
collected by E. Baur in 1918). No authentic material located; disposition based
on description.
Oenothera
albinervis
Gates, Philos. Trans., Ser. B, 226: 339. 1936. Oenothera
var. albinervis (Gates) Gates, Rhodora 59: 15. 1957.-TYPE:
U.S.A.
North Dakota: Cass Co., Fargo (cultivated from seeds collected by R. R. Gates
strigosa
in 1932), 1935, Gates 99.35
(lectotype,
here designated: BM! 2 sheets; isolecto
type:GH!).
Oenothera renneri H. Scholz, Wiss. Z. Padagog. Hochschule Potsdam, Math.-Natur
wiss. Reihe 2: 206. 1956.-TYPE:
GERMANY. Brandenburg: Berlin, Tiergarten,
"Kronprinzen" shore, 18 Aug 1955, Scholz s.n. (holotype: B!; isotype: B!).
Oenothera
f. angustifolia Rostan'ski,
POLAND.Wroclaw: Wroclaw,
depressa
1965.-TYPE:
11: 509.
Fragm. Florist. Geobot.
Port Miejski, 3 Jul 1959, Rostan'ski
s.n. (holotype:WRSL!).
Oenothera
depressa f. latibracteata Rostan'ski, Fragm. Florist. Geobot.
11: 510.
1965.-TYPE:
SWEDEN: Skane, Hyby, Klagerup, 28 Aug 1924, Ander s.n. (holo
type: LD!;
Oenothera
isotype: LD!).
canovertex
Hudziok, Verh. Bot. Vereins Prov. Brandenburg
105: 104.
GERMANY. Brandenburg: Teupitz, Gross Koris, 14 Jul 1965,
s.n. (holotype: HAL, not located).
1968.-TYPE:
Hudziok
Oenothera velutinifolia Hudziok, Verh. Bot. Vereins Prov. Brandenburg
105: 103.
1968.-TYPE:
GERMANY. Brandenburg:
Juterbog, sandy site at railway near
Tiefenbrunnen,
15 Jul 1967/A., Hudziok s.n. (holotype: HAL, not located; iso
type:LZ!).
Stems usually green or red in the lower part, exclusively densely strigillose, some
times also with few appressed to subappressed pustulate hairs, the pustules red or green.
Leaves gray-green to dull green, with prominent pale green, rarely red venation, espe
cially on the lower surface, margins conspicuously dentate, the lower part sometimes sin
uate-dentate.Bracts exclusively densely strigillose. Inflorescence relatively dense, the
apex narrowly
truncate, the infructescence with internodes conspicuously
shorter than the
capsules. Floral tube strigillose, rarely sparsely glandular-puberulent
or with a few longer
spreading to subappressed hairs. Sepals green to yellowish green, densely strigillose.
Ovary strigillose and with longer appressed hairs, sometimes with a few longer spreading
to subappressed hairs. Free tips of the capsule valves erect. Chromosome
number: n = 7
(0)14; or 0)12 and
Phenology.
III;based on 29 individuals from 26 localities).
Flowering during July and August, rarely later.
Fig. 18.
Distribution (Figs. 13, 19,20, 21, 22). Occurring at low elevations up to 1500 (-1650)
m, primarily in the Great Plains region of North America, but now established throughout
much of North America. The full North American
range encompasses
the area from
southern British Columbia
east to Quebec and New Brunswick, Canada, south throughout
the eastern two-thirds of the United States from the eastern foothills of the Rocky Moun
SYSTEMATICBOTANYMONOGRAPHS
74
VOLUME 50
mm
,
13
~~~~~a,b,c
P,~~~~~~~~~~~~~/
e
W-0-
?
I~~~~~~~~~~~~~~~~~~~~~~~~~~~~...
)~~~~~
FIG. 18. Oenothera villosa subsp. villosa (Barkley 045-7, cult. DUSS-88-2024). a. Inflorescence.b.
Rosette leaf. c.Mid-cauline leaf. d. Capsule. e. Inflorescencepubescence.
1997
OENOTHERA
75
0~~~~~~~~~
*
*
~~~~00.
:.
0. villosa subsp. villosa
FIG.
tains inMontana
Arkansas,
19. Indigenous
southeastern
0
distribution
states. Oenothera
villosa
Great Plains, which has subsequently,
villosa
foothills
a few additional
and Virginia;
400km
of Oenothera
to the eastern Rocky Mountain
Tennessee,
*
subsp. villosa
subsp. villosa.
in Colorado,
collections
in historic
times. It grows
habitats, primarily prairies, along streams or lakes, open woodlands,
sites. It is also widely
ported as 0. villosa
in 1977), Europe, Asia,
subsp. strigosa by Dietrich
(erroneously
cited by Rostan'ski (1975) under the names Oenothera
strigosa,
and 0. depressa were used in the production of the distribution maps.
The flowers of 0. villosa
on weather
haps depending
subsp. villosa
The most
Those
conditions.
intergrades extensively
in the easternmost
the pollen
level of autogamy,
subsp. strigosa
shed
per
0. villosa
as discussed
above,
(AA genome,
III, Biennis-I),
the result of the presumed
the plains states. Hybrids
the direction of the cross. When
plastome
I) is crossed
an AA combination
area in the
south to Arkansas.
to Ohio and Wisconsin,
part of the range are probably
subsp. villosa beyond
and 0. biennis differ somewhat with
plastome
the high
with 0. villosa
States from Iowa and Missouri
invasion of 0. villosa
genome,
Despite
with
renneri,
the flowers are cleistogamous,
hybrids are with 0. biennis. They occur across a wide
common
losa as the female
are highly autogamous,
0.
several other species of subsect. Oenothera.
and hybridizes with
central United
subsp. villosa
the flower opens. Sometimes
re
and South Africa.
Specimens
onto the stigma before
spread to
in a variety of
old fields, and other
in southern South America
naturalized
in the
appears to be largely a taxon of the
both naturally and with human assistance,
the north and east of the plains region, primarily
disturbed
northeastern Texas,
have been made
0. villosa
0. villosa
subsp. vil
to 0. biennis
with plastome
recent
between
as the male
(BA
1/III results, similar
SYSTEMATICBOTANYMONOGRAPHS
76
VOLUME 50
+
40;N
1800 E
0 0. biennis
0. villosa
subsp.
villosa
0
FIG. 20. Distribution
of Oenothera
to the female parent but less pubescent.
distinguish
from the phenotype
ographical
area, because
biennis
and 0. villosa
2000 Km
subsp. villosa
in Asia.
In this respect these hybrids are often difficult
to
of 0. biennis [Biennis-I of Cleland (1972)] in the same ge
they usually have few, if any, glandular hairs and no pustulate
hairs in the inflorescence.
The reciprocal cross with 0. biennis as the female (BA-Ill)
losa as the male
more
densely
pubescent.
They
and 0. villosa
subsp. vil
hybrids. These
also resemble 0. biennis, but are
can often be distinguished
from 0. biennis by the more
(AA-I) yields BA-III/I
silky aspect of the pubescence derived from 0. villosa subsp. villosa.
Oenothera villosa subsp. villosa apparently also hybridizes with 0. jamesii where
their ranges overlap. The situation is discussed
under 0. jamesii
(no. 2). Apparent
past hy
OENOTHERA
1997
'I *
77
1
D
|
400
800 Km
* 0. biennis
* 0. villosa subsp. villosa
t
+~
;
60W
tk
FHG. 21. Distribution
0
~
~~~~50
of Oenothera
biennis
and 0. villosa
subsp. villosa
in southern
South America.
SYSTEMATICBOTANYMONOGRAPHS
78
VOLUME 50
40N
W5E
*0. oakesiana
* 0. villosa subsp. villosa
400
0
FIG. 22. Distribution
of Oenothera
800
oakesiana
Kmo
and 0.
villosa
subsp. villosa
in Europe.
bridization
between these two species has resulted in novel phenotypes within 0. villosa
such as one with longer floral tubes. These plants occur in Kansas and Ne
braska (e.g., Ottawa Co., Kansas, Brooks 18319; cultivated material from Riley Co.,
Kansas, Wetter 041, 049 and Barkley 045-77; and Kearney Co., Nebraska, Hapeman s.n.).
Their phenotype is characterized by floral tubes usually 4-5 cm long, vigorous rosettes
subsp. villosa,
short-petiolate leaves, and very dense apically truncate inflorescences.
Oenothera villosa subsp. villosa is naturalized in Asia, Europe, South Africa, and
southern South America, and it hybridizes with 0. biennis, 0. glazioviana,
and 0. jame
sii in these areas where they co-occur. The combinations involved, and the names ascribed
with
to some of them, are given
in the Introduction
(Table 4).
5b. Oenothera
villosa subsp. strigosa (Rydberg) W. Dietrich & P. H. Raven, Ann. Mis
souri Bot. Gard. 63: 383. 1976. Onagra strigosa Rydberg, Mem. New York Bot.
Gard. 1: 278. 1900. Oenothera strigosa (Rydberg) Mackenzie & Bush, Fl. Jack
son Co., Missouri
139. 1902 [Mackenzie and Bush in error gave "Oenothera bi
ennis Linnaeus
var. strigosa Rydberg"
as the basionym for their combination].
(Rydberg) Lunell, Amer. Midl. Naturalist 4: 481. 1916.
Oenothera rydbergii House, New York State Mus. Bull. 233: 61. 1921, nom. su
perfl. Onagra biennis var. strigosa (Rydberg) Piper in Piper & Beattie, Fl.
Palouse Region 124. 1901. Oenothera
biennis var. strigosa (Rydberg) Piper,
Contr. U.S. Natl. Herb. 11: 407. 1906 [combination also proposed by Cronquist,
Usoricum
strigosum
Great Basin Naturalist
52: 77. 1992]. Oenothera
villosa
var. strigosa
(Rydberg)
OENOTHERA
1997
79
298. 1988.-TYPE:
U.S.A. Montana: [Madi
son Co.], near Pony, 7000 ft, 12 Jul 1897, Rydberg & Bessey 4584 (lectotype,
designated by Dietrich & Raven, 1976: NY!; isolectotypes: CAN! F! K! MIN!
PH! US!). [Tiehm and Stafleu (Mem. New York Bot. Gard. 58: 57. 1990) incor
Dorn, Vascular plants of Wyoming
rectly indicate the lectotype here chosen as a holotype "designated in herb."]
strigosa [var.] subulata Rydberg, Mem. New York Bot. Gard. 1: 279. 1900.
Oenothera subulifera Rydberg, Bull. Torrey Bot. Club 40: 66. 1913, nom. nov.
Onagra
strigosa var. subulifera Gates, Rhodora 59: 15. 1957, nom. superfl.
TYPE: U.S.A. Montana: [Madison Co.], forks of theMadison
[River], 7000 ft, 26
Jul 1897, Rydberg & Bessey 4588 (holotype: NY!; isotypes: F! K! US!).
44: 302. 1907.
Oenothera
cheradophila
Bartlett, Bot. Gaz.
(Crawfordsville)
Oenothera
Oenothera
strigosa var. cheradophila
(Bartlett) Gates, Rhodora 59: 15. 1957.
Oenothera strigosa subsp. cheradophila (Bartlett) Munz, N. Amer. Fl., ser. 2, 5:
136. 1965. Oenothera villosa subsp. cheradophila (Bartlett) W. Dietrich & P. H.
Raven, Ann. Missouri Bot. Gard. 63: 383. 1976.-TYPE:
U.S.A. Washington:
Klickitat
Co., Bingen, river bank, 20 Aug 1906, Suksdorf 5860 (holotype: GH!;
isotypes: BH! BM! CAS! DS! F! ISC! MICH! 2 sheets, MIN! MO! NY! ORE!
RSA! US! WS! 3 sheets).
procera Wooton & Standley, Contr. U.S. Natl. Herb. 16: 156. 1913.
Oenothera strigosa var. procera (Wooton & Standley) Gates, Rhodora 59: 15.
1957.-TYPE: U.S.A. New Mexico: San Miguel Co., Pecos River National For
est, Winsor Creek, 2550 m, 5 Jul 1908, Standley 4212 (holotype: US-498579!;
Oenothera
isotype:GH!).
Stems flushed with red at least below, and often entirely red, pubescence usually con
sisting of three types mixed together: a) strigillose, rarely exclusively
so; b) of long erect
to ascending or subappressed red-pustulate hairs; and c) at least in the inflorescence glan
dular-puberulent. Leaves green to dull green, venation not especially prominent, margins
denticulate to subentire, sometimes moderately dentate, the teeth usually widely spaced.
Bracts
strigillose, the apical ones often also glandular-puberulent.
Inflorescence relatively
open, the infructescence with internodes as long as or longer than the capsules. Floral tube
strigillose, and usually also appressed short-villous, glandular-puberulent,
and with some
longer pustulate hairs. Sepals red-striped or flushed with red. Ovary pubescence same as
that of floral tube. Free tips of the capsule valves conspicuous,
spreading. Chromosome
number: n = 7 (014; based on 10 individuals from 10 localities). Fig. 23.
Phenology.
Flowering
primarily during July, but rarely as early as June and as late as
September.
Distribution
(Fig. 24). Occurring primarily in open, often wet sites such as stream
sides, fields, and roadsides, 30-3150 m, in the Rocky Mountains
to the Pacific Northwest
and eastward to the Great Lakes, including the southern portions of British Columbia east
ward to Manitoba,
south to northern California,
the northern half of Nevada and Utah,
Apache, Coconino, and Yavapai counties, Arizona, New Mexico,
northeast to Nebraska
and Minnesota. Known from a few scattered sites in northern Wisconsin, Michigan,
and
Ontario, which appear to represent introductions.
There are instances of presumed hybridization
involving 0. villosa subsp. strigosa
that require discussion. One involves plants from central British Columbia, which ap
peared initially to be very similar to 0. wolfii; a second is a case of possible hybridization
between
0. villosa
subsp. strigosa
and 0. elata subsp. hirsutissima
reported by Cleland
SYSTEMATICBOTANYMONOGRAPHS
80
VOLUME 50
mm
-du3Q..a
e,f u2
e
FIG. 23. Oenothera
1934,
cult. DUSS-88-2023
rescencepubescence.
villosa
subsp.
strigosa
[f]). a. Inflorescence.
(Munz s.n. in 1936,
b. Rosette
leaves.
cult. DUSS-88-2022
c. Mid-cauline
[a-e]; Munz
leaf. d. Capsule.
e-f.
s.n.
in
Inflo
81
OENOTHERA
1997
0
@0.00
X
.000
0
K~~~~~~~~~~~~~~~
0
villosa
sbspF
strigsaX
FJG. 24. Distribution
(1972); and the last concerns
of Oenothera
villosa
the proper placement
rather frequent at higher elevations
0. biennis
One of the specimens
t3
O 0 9
.
0
termediate between
,4SOkm
primarily
subsp.
of plants with
in Colorado
and 0. villosa
strigosa.
a phenotype,
and New Mexico,
which
is
in
somewhat
subsp. strigosa.
studied for this project was an odd specimen very similar in ap
to 0. wolfii, but without glandular hairs, from Botanie Valley, British Columbia
very far to the north of the range of this rare local coastal species. Field
pearance
(Perry 4506),
study of this locality by Wagner
in 1981 revealed only very densely pubescent forms of
at Dusseldorf
from seeds of Wagner
subsp. strigosa. Plants grown subsequently
0. villosa
4547 yielded
two different phenotypes.
notype. They
The first consisted
represent hybrids between
0. villosa
of plants with AB- or BA-phe
subsp. strigosa
and 0.
biennis
(the
"ersteinensis"). These individuals closely resembled 0. biennis, but
were more variable, especially in flower size. The other phenotype represented was an AA
combination, similar to 0. villosa subsp. strigosa but with somewhat larger flowers; these
western
phenotype
correspond
to the Perry 4506 collection.
small-flowered
phenotype
notype of 0. villosa
cially to the south.
selfing
these plants yielded
only a single
that corresponds closely to the normal densely pubescent phe
in this region. Thus, the Perry collection and the orig
subsp. strigosa
inal progeny of Wagner 4547
biennis. Oenothera
Upon
biennis
represent hybrids between 0. villosa
is common
in more mesic
subsp. strigosa
sites throughout
and 0.
this region, espe
SYSTEMATICBOTANYMONOGRAPHS
82
case of hybridization
Another
VOLUME 50
was reported by Cleland
(1972, pp. 294-295). He grew
plants from a single collection from Hesperus, Colorado, that yielded plants with several
phenotypes and chromosomal configurations from 014 to 04 and 5I. His further analy
sis of these plants with experimental crosses to standards suggested that one of the com
a hookeri and the other a strigosa. His conclusion was that the plants repre
plexes was
sented a case of recent hybridization between 0. villosa subsp. strigosa and 0. elata
subsp. hirsutissima. He also pointed out that such cases would probably be with 0. villosa
subsp. strigosa
pollen
as the male,
because
from 0. elata would
0. villosa
rarely have a chance
is highly autogamous
subsp. strigosa
to function.
and
The final instance is particularly difficult, and has not been fully explored, but demon
strates the very active evolution of Oenothera subsect. Oenothera. This situation involves
a series of plants, primarily
from Colorado
and New Mexico
but also from scattered lo
in Oregon, Washington,
and Wyoming,
that are intermediate between 0. villosa
subsp. strigosa and 0. biennis. The question is whether these populations represent intro
calities
ductions of 0. biennis, or the evolution of a new AA combination phenotype, either di
rectly from 0. elata subsp. hirsutissima or from a large-flowered form of 0. villosa, which
is exceedingly
difficult to distinguish from 0. biennis, at least in pressed material. The
plants in question
are generally
lations of 0. villosa
greener, more
robust and less pubescent
subsp. strigosa. They have the somewhat
tulate hairs and the glandular hairs that characterize
vegetative
parts as well
they have only sparse, sometimes
on the ovary). This pubescence
type of 0. biennis,
but when
nally, these plants have midcauline
usually
cm wide,
about 2.2-2.6
leaf margins
in 0. villosa
dentate midcauline
leaves wider
both taxa.
again characterizes
very sparse, strigillose
it is characteristic
on the younger
feature of these plants is
pattern does not occur in the western
denser
than most popu
to spreading pus
both taxa, especially
as on the floral tube and ovary. Another
the red flush on the stems and often on flower parts, which
Moreover,
appressed
of 0. villosa
pubescence
ersteinensis
(at least
pheno
subsp. strigosa. Fi
than usual in 0. villosa
subsp. strigosa,
but have subentire
subsp. strigosa.
to denticulate margins, similar to the
In contrast, 0. biennis in theWest usually has
leaves that are over 2.5 cm wide.
Thus far only one collection of this kind has been grown in the experimental garden.
It is from central New Mexico
(Bernalillo Co., Sandia Mts, Cienaga Canyon, 1975, Wag
ner s.n.). Our crosses with this strain indicate that it is an AB combination with plastome
II, i.e., 0. biennis.
Itwas collected
a parking lot, and could well
in a relatively disturbed
site along a stream adjacent to
represent an introduction. The other collections
we have seen
(indicated by an asterisk in the specimens
cited) usually have been collected
ently relatively
are characteristic
To complicate
less-disturbed
the matter
subsp. hirsutissima,
sites, which
further, populations
particularly
of 0. villosa
from appar
subsp. strigosa.
in the southern part of the range of 0. elata
from Colorado
(e.g., Raven 26551 from Chaffee Co.,
Colorado), represent a large-flowered outcrossing phenotype very similar to the odd in
termediate one described above, and could have given rise directly to it. Therefore, in this
treatment these plants are tentatively included in 0. villosa subsp. strigosa pending more
definitive studies. The strain from Bernalillo Co., New Mexico, has been placed in 0. bi
ennis because of its genomic constitution.
6. Oenothera
stucchii
Soldano,
["1979"].-TYPE:
Ist. Bot. Univ. Lab. Crittog. Pavia,
ITALY.Region
Lombardy:
Prov. Milano,
Cuggiono, Sep 1954, Stucchi s.n. (holotype:Fl!).
ser. 6, 13: 151. 1980
at river Ticino
near
OENOTHERA
1997
83
Erect biennial herb with a taproot, forming a rosette; stems 12-20 (-30) dm tall, green
or in the lower parts flushed with red, usually branched obliquely from the rosette and
with secondary branches, densely strigillose and with scattered long appressed hairs, these
sometime with
both
surfaces
oblanceolate
red-pustulate
bases. Leaves
bright green, veins pale green, strigillose on
15-25 cm long, 2-4 cm wide, narrowly
bluntly dentate, the teeth widely spaced, near the
apex acute, base gradually narrowed to the petiole.
and margins. Rosette
to oblanceolate, margin
leaves
base becoming
sinuate-dentate,
Cauline leaves 6-15 cm long, 2-3 cm wide, narrowly elliptic, margins dentate, the teeth
sometimes widely
spaced, or subentire, apex acute, base narrowly cuneate, sessile to
short-petiolate. Bracts 2-6 cm long, 1.5-2.4 cm wide, lanceolate to narrowly ovate, mar
gin remotely denticulate, apex acute, base acute to rounded, sessile. Inflorescence un
branched. Floral tube 5-6 (-7) cm long, ca. 1.5 mm in diameter, densely strigillose to ap
pressed short-villous, also sparsely glandular-puberulent. Mature buds 1.5-2 cm long, 5-7
mm in diameter, narrowly oblong to oblong, obtusely quadrangular in cross section.
Sepals
1.7-2.5
cm long, 4-5 mm wide,
green to yellowish green, pubescence like floral
long, strigillose, erect in bud. Petals 2-3.5 cm long, 2-3.4 cm
wide, yellow, broadly obovate to very broadly obovate, retuse to emarginate. Filaments
1.5-2 cm long; anthers 8-13 mm long; pollen ca. 50% fertile. Ovary 8-10 mm long, ca.
2 mm in diameter, densely strigillose or with longer appressed hairs, also glandular-pu
tube; free sepal tips 2-4 mm
berulent toward the apex. Style 5.5-6.5 (-7.5) cm long, the exserted part 0.5-1.3 cm long;
stigma surrounded by or slightly below the anthers, which shed pollen directly onto the
lobes at anthesis, the lobes 4-6 mm long, erect and appressed or spreading in open flower.
Capsules 2.5-3.5 cm long, 5-8 mm in diameter, lanceoloid, tapering toward the apex,
green with pale green midvein, dull green when dry; free tips of the valves 1.8-2.5 mm
long, obtuse to rounded. Seeds 1.2-1.8 mm long, 0.8-1.2 mm in diameter, brown. Chro
mosome number: n = 7 (0314; 0)12 and 1II;based on 2 individuals from 2 localities). Self
compatible,
usually autogamous,
Phenology.
Flowering
PTH. Fig. 25.
from August
through October, rarely as early as July.
(Fig. 26). Occurring along rivers, in fallow fields, and along roadsides,
in northwestern Italy in the regions of Liguria, Molise, Piedmont, and Tuscany; also re
ported (Soldano 1979) from two other regions, Emilia-Romagna
(province of Piacenza),
and Lombardy (provinces of Milano, Pavia, and Varese). In 1978 it was discovered to
occur also in Bouches-du-Rhone,
France.
Distribution
Oenothera stucchii is phenotypically
an AA genomic combination with plastome I. It
is a highly autogamous PTH species with about 50% pollen sterility. Distinguishing
fea
tures of this species are the long floral tubes (5-7 cm long) in combination with petals
cm long, and obtusely quadrangular buds. It appears to have originated
between 0. biennis and 0. jamesii, as discussed in the chapter Origins.
2-3.5
as a hybrid
7. Oenothera
grandiflora
L'Heritier in Aiton, Hortus kew. 2: 2. 1789, non Oenothera
grandiflora Lamarck, 1798. Oenothera grandiflora var. glabra Seringe in DC.,
Prodr. 3: 46. 1828. Oenothera biennis var. grandiflora (L'Heritier) Lindley, Ed
wards's Bot. Reg. 19: t. 1604. 1833 [combination also proposed by Torrey & A.
Gray, Fl. N. Amer. 1: 492. 1840]. Oenothera biennis f. grandiflora (L'Heritier)
Carpenter in Dole, Fl. Vermont, ed. 3, 198. 1937.-TYPE:
U.S.A. Alabama:
Baldwin Co. "a few miles above Tensaw" [modern spelling], which according to
F. Harper (1958, p. 405) is "along the east channel between Hall's Creek and the
Alabama
River" of the Tensaw River
(cultivated
at Upton, West Ham, England
SYSTEMATICBOTANYMONOGRAPHS
84
VOLUME 50
mm
a L.Q
e b
tAf~
~
~~
aI
e
FIG. 25. Oenothera
c. Mid-cauline
stucchii
leaf. d. Capsule.
(Soldano
s.n. in 1983,
e. Inflorescence
cult. DUSS-88-2021).
pubescence.
a. Inflorescence.
b. Rosette
leaf.
85
OENOTHERA
1997
* . stucchBii
O., parviflora
J
FIG. 26. Distribution
X
of Oenothera
40
parviflora
?
00 Km/-5
and 0.
stucchii
in Europe.
from seeds collected by W. Bartram after 5 Aug 1778) Fothergill s.n. (neotype,
here designated: BM!). We have not located any authentic material in G-DC
where, according to Stafleu and Cowan (1981), the specimens used by L'Heritier
in preparing descriptions for his contributions toHortus Kewensis are deposited.
in the protologue that Bartram sent seeds to Fothergill. The
L'Heritier mentions
Fothergill specimen at BM has written on the back of the sheet "Hort. Fothergill
1778 "; it is the only specimen located that was derived from the original collec
tion of Bartram, and is therefore designated the neotype.
Oenothera grandifiora Lamarck, Encycl. 4(2): 554. 1798, non Oenothera grandifiora
lamarckiana Seringe in DC., Prodr. 3: 47. 1828,
L'Heritier, 1789. Oenothera
nom. nov.-TYPE:
cultivated from seeds at the Botanical Garden of the Natural
at Paris, Lamarck s.n. (lectotype, here designated: P-LA!).
History Museum
[There are two sheets in Lamarck's herbarium, both labeled Oenothera grandi
flora by Lamarck; the one labelled "sheet A" by H. de Vries is selected as the lec
totype.]
1824: 20. 1824.
J. Lehmann,
Sem. Hort. bot. Hamburg
spectabilis
Oenothera grandifiora var. pubescens Seringe in DC., Prodr. 3: 46. 1828.
TYPE: "O. grandiflora , pubescent in Sims, Bot. Mag. 46: t. 2068" (1819). [No
authentic material seen.]
Oenothera
Erect biennial herb with a taproot, forming a rosette; stems 10-30 (-40) dm tall, un
branched or obliquely branched from the rosette and the main stem, stems toward apex of
plant usually green, the lower ones red, rarely red throughout the plant, often appearing
SYSTEMATICBOTANYMONOGRAPHS
86
glabrous
to the naked eye, but usually
translucent hairs below
rescence
glabrous,
numerous
and with
strigillose
VOLUME 50
or few pustulate
the pustules
never red in fresh material, the inflo
or only glandular-puberulent.
and glandular-puberulent,
inflorescence,
strigillose
Leaves
soft and thin, bright green, the rosette leaves sometimes with reddish brown flecks,
rosette and cauline leaves glabrous or sparsely short-villous on upper surface and sparsely
cm wide,
tic, margins
or deeply
and on veins of lower surface. Rosette
on margins
strigillose
3-6.5
to narrowly obovate,
narrowly oblanceolate
the teeth widely
bluntly dentate,
cm wide,
cm long, (2-)
to ellip
in lower half often sinuate-dentate
spaced,
to the petiole. Cauline
lobed, apex acute, base gradually narrowed
cm long, 1.5-6.5
leaves 18-32
or narrowly elliptic
leaves 6-20
to elliptic, narrowly ovate, narrowly lanceo
late or in lower parts also narrowly oblong, margins, apex, and base the same as in rosette
narrowly elliptic
leaves, except base often more
pale green and deciduous,
margins,
attenuate. Bracts 2-6.5
glabrous,
or glandular-puberulent,
sometimes
narrowly
cm long, 0.5-1.6 cm wide, usually
some appressed hairs at apex and
with
to lanceolate,
lanceolate
narrowly ovate, margin bluntly dentate, often with widely
narrowly elliptic or
spaced teeth, or subentire, apex
acute to long-acute, base obtuse to cuneate. Inflorescence unbranched, often with sec
ondary or tertiary spikes just below the main spike, flowers at an acute to an obtuse angle
to the stem, in the latter case somewhat curved upward. Floral tube 3.5-5.5 cm long, 1-1.3
mm
in diameter, yellowish
green or flushed with
berulent and sparsely long-villous,
Mature
buds 2-4.5
Sepals 2.2-4.6
bescence
sometimes
cm long, 5-9 mm
in diameter,
cm long, 3.5-7 mm wide,
yellowish
lanceoloid
to lanceoloid.
green or flushed with
some red, pu
narrowly
the same as the floral tube; free sepal tips 2-9 mm
illose, erect in bud. Petals
3-4.5
(2.5-)
truncate to retuse. Filaments
cm long, 3-4.8
18-27 mm
to sparsely glandular-pu
red, densely
only one hair type present or tube glabrous.
long, strigillose
cm wide,
long; anthers 10-15 mm
fertile. Ovary 0.8-1.5 cm long, 1.2-2 mm
some of the hairs pustulate, and sometimes
in diameter,
sparsely
to substrig
very broadly obovate,
long; pollen 90-100%
long-villous,
also sparsely strigillose
sometimes
and sparsely covered
with
longer appressed hairs, or glabrous. Style 5.7-9 cm long, the exserted part 2.4-3.8
cm long; stigma elevated above the anthers at anthesis, the lobes 5-10 mm long. Capsules
1.5-3.5
cm long, 3.5-5.5 mm
the apex, bright green when
in diameter,
lanceoloid
to narrowly ovoid,
tapering toward
fresh, dull green and the valves with a whitish
green midvein
when dry; free tips of the valves 0.7-1.5 mm long, rounded to truncate. Seeds 1-1.7 mm
long, 0.6-1 mm in diameter, brown to dark brown. Chromosome
number: n = 7
(711;04
or
and 5I; 06 and 4II; 06, 04, and 21I; 2 04 and
012
and
and
311;08
311; (0)10 and 211;
or rarely self-in
III; or (0 14; based on 85 individuals from 16 localities). Self-compatible
compatible, mostly
Phenology.
outcrossing.
Flowering
Fig. 27.
usually
from August
through October,
as early as July and as
late as November.
Distribution
(Fig. 28). Occurring in scattered, presumably relictual populations on
loose sand over limestone, along streams, marshes or ditches, but some
times as a colonizer in disturbed sites such as along roads, from widely scattered locali
ties in the southeastern United States, ranging from the eastern half of Mississippi
and Al
chalky bluffs,
abama, east to Tennessee
Macon,
Martin, Moore,
(Oconee, Spartanburg,
(Franklin and Marion
counties),
North Carolina
(Cherokee,
New Hanover, Sampson, and Swain counties), South Carolina
and Sumter counties), and Florida (Alachua, Escambia, Franklin,
Lake, Leon, Polk, Putnam, and Santa Rosa counties). Collections
from New York, Penn
sylvania, Vermont, and West Virginia almost certainly represent cultivated plants, garden
87
OENOTHERA
1997
a,b,,c
~~~L)
~
~
L
mm
30
I
e
FIG. 27. Oenothera
c. Mid-cauline
grandi;flora (Kral s.n. in 1979, cult. DUSS-87-st-279).
e. Inflorescence
pubescence.
leaf. d. Capsule.
a. Inflorescence.
b. Rosette
leaf.
SYSTEMATICBOTANYMONOGRAPHS
88
VOLUME 50
a A,
X AA
0
500
km
FIG. 28. Distribution
escapes,
or adventive
populations,
of Oenothera
and
the
s'ingle
grandiflora
and O. nutans.
from
locality
central
also
Kentucky
may
be an introduction.
1959,
Oenothera
grandiflora
1963,
The
1964).
flora. Oenothera
other
grandiflora
is one of two BB genome
species,
O.
nutans,
is a large-flowered
may
species with plastome
be
a direct
bivalent-forming
derivative
species
III (Stubbe
of 0.
grandi
that is presum
OENOTHERA
1997
89
It has long been thought to be an entirely self-compatible
outcrossing.
species, as are all other members of subsect. Oenothera. Self-incompatibility
was discov
ered in plants from York and Bellamy, Alabama (Stubbe & Raven 1979b). This discovery
ably mostly
led to more
studies of this species (Steiner & Stubbe 1984, 1986; Schumacher
that have shown that 0. grandiflora is far more diverse than previously
thought. Some populations seem to be entirely or mostly composed of self-incompatible
individuals, whereas others consist of self-compatible
plants. This is an extremely un
intensive
& Steiner 1993)
common phenomenon
in Oenothera; the only other species known to exhibit mixed pop
ulations of self-incompatible
and self-compatible
individuals is 0. primiveris A. Gray (W.
L. Wagner unpubl.).
The discovery
of self-incompatibility
in 0. grandiflora
is significant, because the
for immediate establishment of a PTH entity following hybridization requires
in the chromosome complexes
the presence of Si-alleles
(Steiner 1956; Cleland 1972;
mechanism
Stubbe & Raven
1979b). Oenothera grandiflora or its common ancestor has been hy
as one of the presumed progenitors of 0. biennis (Steiner 1952; Cleland 1972),
and the absence of Si-alleles in any known population led Cleland to hypothesize that the
present 0. grandiflora is a remnant for his Population 2 in which the Si-genes necessary
pothesized
for the formation of 0. biennis were present. The discovery of the alleles in a number of
extant populations suggests that 0. grandifiora could have been one of the direct ances
tors of 0. biennis (Stubbe & Raven 1979b; Steiner & Stubbe 1984); however, our current
favored hypothesis is that 0. biennis originated directly from hybridization with 0. nutans
as one of the parents as indicated in the section on origins above.
0. grandiflora exhibits far more diversity than previously
Cytologically,
thought
(Steiner & Stubbe 1984, 1986; Schumacher & Steiner 1993). A wide range of configura
tions are known, from 7II to 0 12 and III, or rarely a 0 14. Moreover,
the recent work in
cluding the extensive study of herbarium material has revealed a much greater geograph
ical range of 0. grandiflora than previously known.
Despite
the chromosomal
diversity, morphological
variation is considerably less than
is variation in pubescence and the lobing of the basal
leaves. Plants from Florida have deeply lobed basal and lower cauline leaves, whereas
those from populations to the west are usually merely dentate. An unusual characteristic
for subsect. Oenothera
is that inflorescences of 0. grandiflora often have secondary lat
eral inflorescences below the central one. This phenotypic characteristic is clearly associ
that exhibited
by 0. elata. There
ated with the B genome. Oenothera grandiflora has several features that suggest adaption
to more mesic conditions. Among them are the broad membranous
leaves, sparse pubes
cence, the tall habit up to 4 m, and perhaps the quickly deciduous bracts.
Both of the species believed to be derived, at least in part, from 0. grandiflora, 0. bi
ennis and 0. nutans, grow sympatrically with it. Like 0. grandiflora, the form of 0. bi
ennis referred to as Biennis-I by Cleland (1972) and known taxonomically as 0. biennis
subsp. centralis by Munz (1965) has plastome III. Hybrids or hybrid derivatives between
0. grandiflora and 0. biennis have been found in cultures of seed from Alabama collected
by E. Steiner in 1983 (Steiner & Stubbe 1986). These hybrids, as would be expected, are
normal green. They exhibit a wide range of phenotypes extending from very similar to ei
ther parent to intermediates. Apparently, the seeds produced an array of hybrid types, in
cluding Fl and various backcross types.
A recent study of these plants by Schumacher and Steiner (1993) has shown that the
Alabama Castleberry and Chastang strains with (010, 012, or 0314 chromosomes during
meiosis clearly consist of two different complexes, one a typical B genome of 0. grandi
90
SYSTEMATICBOTANYMONOGRAPHS
VOLUME 50
flora, and the second amodified B genome. The modified complex has an influence of the
A genome and appears to have been derived from an unstable hybrid with 0. grandiflora
as the pistillate parent and 0. biennis as the staminate parent. The study did not determine
of the A and B genomes to this modified complex, nor what se
lective advantage may have been obtained, which could influence the further evolution of
these populations.
the relative contributions
Oenothera grandiflora has been cultivated in botanical gardens and in the horticul
tural trade since the beginning of the 19th century. About a hundred years ago, however,
it was largely displaced in cultivation by the more easily grown 0. glazioviana.
In culti
vation, 0. grandiflora is a short-day plant. This causes considerable difficulty when grow
ing the species as far north as Dusseldorf, where it does not flower in gardens before Sep
tember. Therefore, in the studies of this species, plants were first grown to the flowering
stage in the greenhouse under short-day conditions.
8. Oenothera
nutans Atkinson & Bartlett in Bartlett, Rhodora 15: 83. 1913. Oenothera
biennis var. nutans (Atkinson & Bartlett) Wiegand,
Rhodora 26: 3. 1924.
TYPE: U.S.A. New York: Tompkins Co., near Ithaca (cultivated from seeds),
Atkinson 2 (holotype: CU!; isotypes: CU! 10 sheets, MICH! 3 sheets).
Oenothera
biennis subsp. austromontana Munz, N. Amer. Fl., ser. 2, 5: 134. 1965.
Oenothera austromontana (Munz) P. H. Raven, W. Dietrich & Stubbe, Syst. Bot.
4: 244. 1980 ["1979"]. Oenothera biennis var. austromontana (Munz) Cronquist
inGleason & Cronquist, Man. vasc. pl. North. U.S. and Can., ed. 2, 864. 199 1.
TYPE: U.S.A. Virginia: Washington
Co., White Top Mtn, blackberry thicket,
1400 m, 21 Aug
1935, Munz 13510
(holotype: POM-212478!;
isotypes: POM! 2
sheets).
Erect biennial
herb with a taproot, forming a rosette; stems 3-20 dm tall, usually
from the rosette and with secondary branches from the main stem, often ap
pearing glabrous to the naked eye, but usually pubescent with either: a) older parts strig
illose and with scattered pustulate hairs, strigillose and sparsely glandular-puberulent
in
the inflorescence; or b) older parts densely strigillose and with longer appressed hairs, the
branched
inflorescence densely glandular-puberulent
to glabrous. Leaves dark green, sparsely strig
illose to villous on both surfaces, or glabrous, except strigillose on themidrib of lower sur
face, the bracts often glabrous or glandular-puberulent. Rosette leaves 10-32 cm long, 3-7
cm wide, narrowly oblanceolate
to narrowly obovate, margin bluntly dentate, the lower
half sinuate-dentate, apex acute, base gradually narrowed to the petiole. Cauline leaves
6-20 cm long, 2-8 cm wide, narrowly elliptic, narrowly lanceolate to lanceolate, or nar
rowly oblanceolate
to oblanceolate,
apex acute to long-acute, margins of the lower cauline
the teeth widely spaced, the lower half of the blade usually sinu
of cauline leaves toward apex of plant dentate to subentire, apex
base gradually narrowed to a short petiole. Bracts 1-2.5 cm long,
leaves bluntly dentate,
ate-dentate, margins
acute to long-acute,
0.2-0.8 cm wide, pale green and caducous, narrowly lanceolate to lanceolate or narrowly
ovate, margin denticulate to subentire, apex acute to long-acute, base acute to narrowly
cuneate, sessile. Inflorescence unbranched, or often with secondary lateral inflorescences
just below the main one. Floral tube 3-4.3 cm long, 0.8-1 mm in diameter, yellowish
green, sparsely villous or glandular-puberulent
or glabrate. Mature buds 0.9-2 cm long,
4-6 mm in diameter, lanceoloid or narrowly oblong to oblong. Sepals 1-2.3 cm long, 3-5
mm wide, yellowish green, rarely red toward apex, pubescence like floral tube; free sepal
OENOTHERA
1997
91
tips 1.5-6 mm long, erect in bud, strigillose. Petals 1.4-2.5 (-3) cm long, 1.5-2.8 cm
wide, very broadly obovate, retuse to emarginate, yellow, fading yellowish white and
translucent. Filaments 10-25 mm long; anthers 4-10 mm long; pollen 16-86% fertile.
Ovary 0.9-1.2 cm long, 1.1-1.5 mm in diameter, with a sparse covering of long appressed
or spreading hairs, and sparsely to densely glandular-puberulent.
Style 3.5-6.3 cm long,
the exserted part 0.5-2 cm long; stigma surrounded by the anthers, which shed pollen di
rectly onto the lobes at anthesis, the lobes 3-7 mm long. Capsules 1.2-3.6 cm long, 3-6
mm in diameter, lanceoloid to narrowly ovoid, tapering toward the apex, sometimes base
like that of ovary but
constricted to a very short stipe, dull green when dry, pubescence
less dense, often glabrate; free tips of the valves 1-1.5 mm long, rounded to retuse. Seeds
1.1-1.9 mm long, 0.6-0.9 mm in diameter, brown to nearly black. Chromosome number:
n = 7 (014 or 012 and 1II;based on 28 individuals from 25 localities). Self-compatible,
usually autogamous, PTH. Fig. 29.
Phenology. Flowering from June through August.
Distribution (Fig. 28). Occurring inmostly open, often disturbed sites, such as stream
beds and flood plains, slopes and margins of mixed deciduous forest, roadsides and old
fields, (240-) 400-1700 m, in eastern North America, primarily in the Appalachian Moun
tains, but extending from Maine west through southeastern Ontario and Michigan,
south
to Mississippi,
in Missouri
Alabama, Georgia, and Florida; disjunct occurrences
and
Arkansas probably represent unintentional introductions by humans.
is a PTH species with a BB genomic constitution and plastome III
in the cyto
1990). It long has been known as Biennis-III
genetics literature (Cleland 1972), and taxonomically
it was treated as 0. biennis subsp.
austromontana
(Munz 1965). More recently, it was elevated to specific status as 0. aus
Oenothera
nutans
(Stubbe 1963, 1964; Wasmund
tromontana by Raven et al. (1979) to give it parallel status to, but distinct from, the AB
and BA genome species, 0. biennis. During the herbarium study for this paper, we dis
covered the earlier name, 0. nutans, which had never been used for this taxon since its
original publication. The epithet does not refer to a nodding inflorescence as in 0. parvi
flora or 0. oakesiana, but rather to the fact that the flowers were thought by the original
authors to wither very quickly after anthesis. Despite this epithet the flowers of 0. nutans
do not wither more quickly than those of other species of subsect. Oenothera.
Wasmund (1984, 1990) studied strains from 31 different populations from New York,
Pennsylvania, Tennessee, Virginia, West Virginia, and North Carolina. His results showed
that all but seven strains had a self-incompatibility
factor in one of the complexes. The cX
(egg) complex generally possesses the Si-gene with each population exhibiting only one
Si-gene, and different populations with different alleles. Between 16% and 52% of the
pollen consists of empty grains. These results indicate an inactivation of one of the two
complexes by the Si-allele acting as a gametophytic
lethal. In 7 of the 31 strains studied
by Wasmund, a pollen sublethal or lethal factor is operative in the cXcomplex, preventing
the transmission of the cxcomplex by the pollen, rather than by Si-alleles.
The great range of pollen fertility in 0. nutans can partly be explained by aberrant
separation of the chromosomes during meiosis. Wasmund also suggested that 0. nutans is
a recently derived species that has not yet reached an optimal stage in development of the
PTH system where about 50% of the pollen would be sterile, as in other heterogamous
PTH taxa. If this is true then 0. nutans was probably not involved in the formation of 0.
biennis, as suggested elsewhere in this monograph.
The ,3 homozygotes
in 0. nutans are prevented by sporophytic lethals, expressed by
abortive seeds (up to 50%). The percentage of abortive seeds varies depending on the
SYSTEMATICBOTANYMONOGRAPHS
92
30
a,b,c
e
-
9
FIG. 29. Oenothera
cauline
VOLUME 50
leaf. d. Capsule.
b
nutans
(Cleland
e. Inflorescence
200,
cult. DUSS-88-2016).
a. Inflorescence.
b. Rosette
leaf. c. Mid
pubescence.
amount of embryo sac competition. As with the Si-alleles, the various strains possess dif
ferent sporophytic lethals. The variation in percentage of viable seeds formed is a further
indication of an imperfectly formed system in a recently derived species.
similar to 0. grandiflora, especially in its dark
Oenothera nutans ismorphologically
OENOTHERA
1997
93
lateral inflorescences, and the pale and
soft leaves, sparse pubescence,
0. nutans is, in general, adapted to cooler, more montane
bracts. Ecologically,
habitats than the other PTH species of the eastern United States, 0. biennis, 0. parviflora,
and 0. oakesiana, as well as its presumed progenitor, 0. grandifiora. Despite this differ
ence in habitat preference, 0. nutans at times grows sympatrically with both 0. biennis
green, somewhat
caducous
and 0. parvifiora. Hybridization between 0. nutans and 0. biennis is possible, because
there are no known incompatibility barriers. Hybrids between these species would be BB,
AB, or BA combinations with corresponding phenotypes. These are the same combina
tions that are present in the parental species; therefore, the hybrids could not be easily de
tected as pressed specimens. Hybridization between 0. nutans and 0. parviflora (BC ge
nomic combination, plastome IV) is also possible with 0. nutans as the pollen parent.
According
to the compatibility analysis of Stubbe (1959), these hybrids (BB combination
with plastome IV) would be normal green and viable, but would not be phenotypically de
tectable. The reciprocal hybrid (BC combination with plastome III) is lethal.
9. Oenothera
biennis L., Sp. pl. 346. 1753. Onagra biennis (L.) Scopoli, Fl. carniol., ed.
1:
2,
269. 1772. Oenothera graveolens Gilibert, Fl. lit. inch. 2:186. 1782, nom.
superfl. Onagra europaea Spach, Hist. nat. veg. 4: 359. 1835, nom. superfl. On
agra vulgaris Spach, Nouv. Ann. Mus. Hist. Nat., Paris 4(4): 353. 1836 ["1835"],
nom. superfl. Oenothera biennis var. vulgaris Torrey & A. Gray, Fl. N. Amer. 1:
492. 1840. Pseudo-oenothera
virginiana Ruprecht, Fl. ingr. 1: 365. 1860, nom.
superfl. Brunyera biennis (L.) Bubani, Fl. Pyrenaea 2: 649. 1900. O[e]nothera
communis H. Leveille, Bull. Acad. Int. Geogr. Bot. 19: 328. 1909, nom. illeg.
[Leveille assembled under this name all of the species of Oenothera
subsect.
Oenothera, and his name is included here under the oldest name, 0. biennis].
TYPE: "Habitat in Virginia, unde 1614, nunc vulgaris Europae," (lectotype, des
ignated by Gates, 1911: LINN-484. 1). Original material included: Herb. Clifford
144 (BM! 2 sheets); Herb. Burser XIV: 4 (UPS); Morison
(1680: 271, s. 3, t.Il,
f. 7); LINN-484. 1. Linnaeus mentioned both Virginia and Europe in the proto
logue. The lectotype matches a phenotype of 0. biennis inNorth America; how
ever, Rostan'ski (1982) believes that this specimen represents a non-North Amer
icanphenotype.
Oenothera muricata
L., Syst. nat., ed. 12, 263. 1767. Onagra muricata (L.) Moench,
675. 1794. Onagra chrysantha var. grandiflora Spach, Nouv. Ann.
Methodus
Mus. Hist. Nat., Paris 4(4): 355. 1836 ["1835"]. Oenothera biennis var. muricata
(L.) Torrey & A. Gray, Fl. N. Amer. 1: 492. 1840. Oenothera parviflora var.
muricata (L.) Farwell, Amer. Midl. Naturalist 8: 274. 1923. Oenothera biennis f.
muricata
(L.) J. Boivin,
Naturaliste
Canad.
93: 644.
1966.-TYPE:
CANADA
(holotype:LINN-484.3!).
Oenothera
suaveolens
Persoon, Syn. pl. 1: 408. 1805. [Persoon perhaps obtained this
from Desfontaines,
Tabl. ecole bot. 169. 1804, nomen nudum.]
O[e]nothera
biennis subsp. suaveolens (Persoon) Rouy & Camus, Fl. France 7:
200. 1901 [combination also proposed by Love & Love, Opera Bot. 5: 258.
name
1961]. O[e]nothera
communis race biennis f. suaveolens (Persoon) H. Leveille,
Bull. Acad. Int. Geogr. Bot. 19: 330. 1909. Oenothera biennis subsp. grandiflora
Stomps, Receuil Trav. Bot. Neerl. 41: 135. 1948, nom. superfl. [illegitimate
stitution for 0. biennis subsp. suaveolens].-TYPE:
Hort. Paris, Desfontaines
(neotype, here designated: FI-W!). [No original material located.]
sub
s.n.
94
SYSTEMATICBOTANYMONOGRAPHS
VOLUME 50
Oenothera media Link, Enum. hort. Berol. alt. 1: 377. 1821. Onagra linkiana Spach,
Nouv. Ann. Mus. Hist. Nat., Paris 4(4): 354. 1836 ["1835"], nom. superfl.
TYPE: No definite authentic material located. A photo atMO of a specimen cul
tivated at the Berlin Botanical Garden from Link's Herbarium
(B, now de
is labeled "Oenothera media"
and fits his description. The specimen is
form of 0. biennis known in Europe as Oenothera suaveolens.
Onagra chrysantha var. latifolia Spach, Nouv. Ann. Mus. Hist. Nat., Paris 4(4): 355.
1836 ["1835"]. No authentic material located. The tentative disposition here is
stroyed)
a large-flowered
based on Spach's description.
muricata var. latifolia Ascherson, Fl. Brandenburg 1: 213. 1864.-TYPE:
GERMANY. Brandenburg: Prietzen s.n. (holotype: B, destroyed). [This entity was
analyzed by Renner (1937); he referred it to 0. rubricaulis because it represents
Oenothera
the rubricaulisphenotype.]
2: 309. 1903.-TYPE:
NETHERLANDS.
from seeds in a nursery; fig. 53, p. 310 (lectotype, here designated).
[No authentic material seen.]
Oenothera
hirtella de Vries, Mutationstheorie
Cultivated
Oenothera
rubiennis
de Vries, Mutationstheorie
LANDS. Cultivated
atAmsterdam
is an artificial hybrid between
2: 102.
1903.-TYPE:
NETHER
by H. de Vries; no authentic material
0. biennis
seen. This
(as 0. cruciata varia) and 0. biennis.
[AB combination]
Oenothera
tracyi Bartlett, Rhodora
(Bartlett) Gates, Rhodora
Dixie
Landing
Bartlett 2749
Oenothera
near Tensaw
grandiflora var. tracyi
Alabama: Baldwin Co.,
from seeds collected
4 sheets).
& Bartlett
biennis var. pycnocarpa
3. 1924.-TYPE:
in Bartlett,
by S. M. Tracy),
Rhodora
15: 83.
(Atkinson & Bartlett) Wiegand,
salicastrum
de Vries,
NETHERLANDS. Cultivated
type, here designated).
Gruppenweise
at Amsterdam
Artbildung
by H. de Vries;
[No authentic material
seen. This
304.
1913.
Rhodora
U.S.A. New York: Tompkins Co., near Ithaca (cultivated
1 (holotype: CU! 6 sheets; isotype: MICH!).
seeds), Atkinson
Oenothera
Atkinson
U.S.A.
(cultivated
(holotype: MICH!
pycnocarpa
Oenothera
13: 210. 1911. Oenothera
59: 13. 1957.-TYPE:
26:
from
1913.-TYPE:
fig. 111, p. 305 (lecto
is a mutant of 0. bien
nis "Chicago."]
Oenothera
reynoldsii Bartlett, Cybele Columb.
1: 39. 1914.-TYPE:
U.S.A. Ten
nessee: Knox Co., Knoxville
(cultivated from seeds collected by E. S. Reynolds
in 1910), Bartlett 3171 (lectotype, here designated: MICH! 2 sheets).
Oenothera
pratincola
Bartlett, Cybele
tucky: Fayette Co., Lexington
Columb.
1: 40. 1914.-TYPE:
(cultivated
from seeds collected
U.S.A.
Ken
by Bartlett
in
3499 or 3542. No material located at MICH or anywhere else;
among R. Cleland's vouchers (now at MO) is a single sheet, MO
of Bartlett material, which represents this entity. This sheet, other than
1912), Bartlett
however,
3838404!
the collector
and taxon, has no identifying marks;
it could represent missing
syn
type material.
Oenothera
numismatica
var. numismatica
Bartlett, Cybele Columb.
(Bartlett) Gates, Rhodora
tucky: Fayette Co., Lexington
1912), Bartlett
scription.
(cultivated
3498 or 3544. No material
1: 41. 1914. Oenothera
59: 16. 1957.-TYPE:
from seeds collected
located; disposition
pratincola
U.S.A.
Ken
by Bartlett
in
based on the de
1997
OENOTHERA
95
brevicapsula Bartlett, Cybele Columb. 1: 42. 1914. Oenothera gauroides
U.S.A. Mary
var. brevicapsula
(Bartlett) Gates, Rhodora 59: 16. 1957.-TYPE:
branch of
land: Montgomery
Co., Chevy Chase, embankment of Georgetown
Baltimore
and Ohio Rivers (cultivated from seeds from Bartlett 2247, Aug
Oenothera
1910), Bartlett
2714 (lectotype,
here designated: MICH!
2 sheets).
Oenothera
ruderalis Bartlett, Cybele Columb. 1: 44. 1914.-TYPE:
U.S.A. Mary
land: Montgomery
Co., vic. of Chevy Chase Lake (cultivated from rosette col
lected by Bartlett in 1910), Bartlett 3149 (lectotype, here designated: MICH! 2
sheets; isolectotypes:BM! RSA!).
biennis var. leptomeres Bartlett, Amer. J. Bot. 1: 242. 1914.-TYPE:
NETHERLANDS. Santpoort (cultivated from seeds obtained from E. de Vries),
Bartlett s.n. (holotype: MICH! 2 sheets).
U.S.A. Mary
Oenothera stenomeres Bartlett, Amer. J. Bot. 1: 242. 1914.-TYPE:
Oenothera
Co., Chevy Chase Lake (cultivated from seeds collected by
in 1910), 1913 (?), Bartlett 3146 (holotype: US-693733-35!
3 sheets).
Oenothera
rubricaulis Klebahn,
31: 12. 1914.
Jahrb. Hamburg. Wiss. Anst.
biennis subsp. rubricaulis (Klebahn) Stomps, Recueil Trav. Bot.
Oenothera
Neerl. 41: 136. 1948.-TYPE:
GERMANY. Niedersachsen: Bevensen near Uelzen,
land: Montgomery
Bartlett
16 Jul 1967, Walther 6702 (neotype, here designated: HBG!;
located; we have designated
[No authentic material
lected at the original
isoneotype: KTU!).
a neotype, which was
col
locality.]
var. rubricaulis Farwell, Pap. Michigan Acad. Sci. 1: 95. 1921
[1923]. Oenothera biennis var. rubricaulis (Farwell) Farwell, Amer. Midl. Natu
ralist 8: 275. 1923.-TYPE: U.S.A. Michigan: Wayne Co., fields near Dearborn,
15 Aug 1920, Billington & Farwell 5597 (lectotype, here designated: BLH;
Oenothera
muricata
isolectotype:MICH!).
Oenothera furca Boedijn, Z. Indukt. Abstammungs-Vererbungsl.
32: 361. 1924.
TYPE: U.S.A. Minnesota: Hennepin Co., North Town Junction (cultivated from
seeds collected by H. de Vries in Aug 1904). No material located; disposition
based on the description.
Oenothera
purpurata
Niedersachsen:
bahn's
Klebahn,
Bevensen
t. 2 (lectotype,
phenotype
represents
Z. Vererbungsl.
(cultivated
39: 19. 1925.-TYPE:
from seeds from H. Klebahn
GERMANY.
in 1914); Kle
here designated).
[No authentic material located.] This
an apparently bivalent-forming
segregate known only in
cultivation; we include it in the synonymy under 0. biennis based on its origin.
A recent reference specimen of this entity is DUSS-88-2020
(MO)
(711).
Oenothera sabulosa Farwell, Amer. Midl. Naturalist 12: 69. 1930.-TYPE:
U.S.A.
Michigan: Keweenaw Co., sandy places, 1 Sep 1889, Farwell 721 (holotype:
BLH;
Oenothera
isotype: MICH!).
shulliana
1931.-TYPE:
Sturtevant, Z. Indukt. Abstammungs-Vererbungsl.
59: 367.
U.S.A. New Jersey: Morris Co., in a garden atMorristown
(culti
vated from seeds collected
by A. H. Sturtevant
(neotype, here designated: MO-3838393!).
Oenothera
in 1926),
1934, Cleland
[No authentic material
34-22
located.]
victorinii
Gates & Catcheside
in Gates, J. Linn. Soc., Bot. 49: 182.
1933.-TYPE:
CANADA. Quebec: Rouville Co., St. Hubert near Montreal (culti
vated in Regent's Park, England from seeds collected by Marie-Victorin
in Sep
1930), 1934, Gates
cated.]
s.n. (neotype, here designated: K!).
[No authentic material
lo
SYSTEMATICBOTANYMONOGRAPHS
96
Oenothera
de Vries
chicaginensis
ex Renner & Cleland,
VOLUME 50
Z. Indukt. Abstammungs
(de Vries ex
U.S.A. Illi
66: 275. 1933. Oenothera biennis subsp. chicaginensis
Renner & Cleland) Love & Love, Opera Bot. 5: 258. 1961.-TYPE:
Vererbungsl.
of Chicago
nois: Cook Co., vicinity
Vries
grandifolia
CANADA. Nova
Oenothera
by H. de
and Cleland
(lectotype,
Ser. B, 226: 282. 1936.-TYPE:
Wentworth
Station (cultivated at Re
Co.,
from seeds collected by R. R. Gates on 30 Sep 1932), 1935,
Philos.
Gates,
Trans.,
Scotia: Cumberland
gent's Park, England,
Gates 58.35.13
from seeds collected
1913, cited by Renner
[No authentic material located.]
here designated).
Oenothera
(cultivated
t. 6 in de Vries,
in 1904);
(lectotype, here designated: BM!).
var. serratifolia Gates, Philos.
Trans., Ser. B, 226: 259.
Scotia: Kings Co., Kentville, Gates 9.33. No au
novae-scotiae
1936.-TYPE:
CANADA. Nova
thentic material
seems
located. Gates
it seems very doubtful
to have made
voucher
specimens
seeds from this strain did not germinate
1934 and 1935, and because
that any authentic material
only
in
after 1933,
exists. This name is therefore
only tentatively placed here.
Oenothera
paralamarckiana
U.S.A. Massachusetts:
Philos. Trans., Ser. B, 226: 245.
Gates,
Barmstable Co., Penzance, Woods
Hole
1936.-TYPE:
(cultivated at Re
gent's Park, England, from seeds collected by R. R. Gates in Sep 1932), 28 Aug
1934, Gates 124.34 (lectotype, here designated: K! 3 sheets).
Oenothera
pycnocarpa
U.S.A.
1936.-TYPE:
Park, England,
1934, Gates
Oenothera
var. cleistogama
from seeds collected
119.34
Philos.
Gates,
Ser. B, 226: 250.
Trans.,
New York: Oneida
(lectotype,
Co., Clinton (cultivated at Regent's
by G. L. Stebbins, Jr., in Sep 1932), 31 Aug
here designated: K! 3 sheets).
var. parviflora Gates, Philos. Trans., Ser. B, 226: 250.
U.S.A. New York: Madison Co., Hamilton (cultivated at Regent's
pycnocarpa
1936.-TYPE:
Park, England,
from seeds collected
by G. L. Stebbins,
Jr., in 1932),
1934, Gates
113.34 (lectotype,here designated:K!).
Oenothera
royfraseri Gates,
sackvillensis
Philos.
var. royfraseri
CANADA. New Brunswick:
England,
sackvillensis
Gates,
CANADA. New Brunswick:
226:
Rhodora
285.
1936. Oenothera
12. 1957.-TYPE:
59:
Albert Co., Sackville
from seeds collected
type, here designated: BM!;
Oenothera
Ser. B,
Trans.,
(Gates) Gates,
by R. Fraser
isolectotype:
Philos.
(cultivated at Regent's Park,
in 1933), 1934, Gates 3.34 (lecto
GH!).
Trans.,
Ser. B,
Albert Co., Sackville,
226:
287.
1936.-TYPE:
behind power house of Uni
versity and adjacent vegetable garden (cultivated at Regent's Park, England,
from seeds collected by R. R. Gates on 29 Sep 1932), 22 Sep 1934, Gates 39.34
(lectotype,
Oenothera
here designated: K! 3 sheets).
sackvillensis
1936.-TYPE:
var. albiviridis
Gates,
CANADA. New Brunswick:
house of University
and adjacent vegetable
Philos.
Trans.,
Ser. B,
Albert Co., Sackville,
garden (cultivated
226:
290.
behind power
at Regent's
Park,
England, from seeds collected by R. R. Gates on 29 Sep 1932); fig. 35 of culture
40.34 (lectotype, here designated).
[No authentic specimens located.]
Oenothera
victorinii var. intermedia Gates, Philos. Trans., Ser. B, 226: 321. 1936.
TYPE: CANADA. Quebec: Jacques Cartier Co., Ste. Anne de Bellevue (cultivated
at Regent's Park, England, from seeds collected by W. G. Dore on 5 Nov 1933),
1934, Gates 4.34 (lectotype,
here designated: BM!;
isolectotype:
GH!).
1997
OENOTHERA
97
Oenothera
victorinii var. parviflora Gates, Philos. Trans., Ser. B, 226: 320. 1936.
TYPE: CANADA. Quebec: Kamouraska Co., Ste. Anne (cultivated at Regent's
on 12 Oct 1931). No au
Park, England, from seeds collected by Marie-Victorin
thentic material
located. Cultures cited are R. R. Gates 49.33, 56.33, 57.33,
73.34, 80.34, 81.34, 82.34, 82.35, 83.34, 88.35, 89.35, 90.35. Dis
position based on description of cultivated strains.
Oenothera victorinii var. undulata Gates, Philos. Trans., Ser. B, 226: 322. 1936.
58.33, 59.33,
TYPE: CANADA. Ontario: Regional Municipality
Co., Toronto, York Mills Road
(cultivated at Regent's Park, England, from seeds collected by R. R. Gates on 7
Oct 1932), 1935, Gates 91.35 (lectotype, here designated: BM!).
Oenothera
101: 47.
Hudziok, Verh. Bot. Vereins Prov. Brandenburg
GERMANY. Brandenburg:
in western part of Luckenwalde,
on
editicaulis
1964.-TYPE:
25 Jul 1962, Hudziok
sandy dry grassland,
s.n. (holotype:
HAL-075235!
3
sheets).
Oenothera
jueterbogensis
Hudziok,
Verh. Bot. Vereins
Prov. Brandenburg
101: 47.
1964.-TYPE: GERMANY.
Brandenburg:Julterbog,"AltesLager," 15Aug 1962,
2 sheets).
s.n. (holotype: HAL-075233!
var. macrosperma Hudziok,
101: 48. 1964. Oenothera macrosperma
Hudziok
Oenotherajueterbogensis
denburg
Martin-Luther-Univ.
Halle-Wittenberg,
Verh. Bot. Vereins Prov. Bran
(Hudziok) Hudziok, Wiss. Z.
Math. Naturwiss. Reihe 14: 490. 1965.
TYPE:GERMANY.
Brandenburg:Luckenwalde, dumping ground near forester's
house "Lindhorst," 22 Jul 1962, Hudziok
s.n. (holotype: HAL-075232!
2 sheets).
Oenothera
tacikii Rostan'ski,
11: 503. 1965.-TYPE:
Fragm. Florist. Geobot.
POLAND. Wroclaw: Wroclaw, between railroad and Robotnicza
Street, 8 Aug
1961, Rostan'ski s.n. (holotype: WRSL! 5 sheets). [Rostan'ski indicates that this
specimen
represents a hybrid between
two phenotypes
of 0. biennis: suaveolens
x rubricaulis.]
Oenothera
nissensis Rostan'ski, Fragm. Florist. Geobot.
POLAND.Wroclaw: Nysa (cultivated from wild-collected
Rostan'ski s.n. (holotype: WRSL! 3 sheets).
11: 508.
rosettes),
1965.-TYPE:
17 Jul 1962,
Oenothera biennis subsp. caeciarum Munz, N. Amer. Fl., ser. 2, 5: 133. 1965.
TYPE: U.S.A. New Hampshire: Hillsborough Co., Hollis (cultivated from seeds
collected on 3 Aug 1935), 24 Jul 1936, Munz 14219 (holotype: POM-224307!;
isotypes: POM! 2 sheets).
biennis subsp. centralis Munz, N. Amer. Fl., ser. 2, 5: 134. 1965.-TYPE:
U.S.A. Kentucky: McCracken Co., 11mi W of Paducah, old field, 29 Aug 1935,
Munz 13542 (holotype: POM-212756!;
isotypes: BH! IND! NY!).
Oenothera
Oenothera
brevispicata Hudziok, Verh. Bot. Vereins Prov. Brandenburg 105: 101.
1968.-TYPE:
GERMANY. Brandenburg: on sandy places in Potsdam, 20 Aug
1965,Hudziok s.n. (holotype:HAL, not located).
Oenothera
compacta
Hudziok,
Verh.
Bot. Vereins
Prov. Brandenburg
105: 96.
1968.-TYPE: GERMANY.
Brandenburg:Teupitz, nearGross Koris, 24 Jul 1967,
Hudziok s.n. (not located, destroyed?).GERMANY. Brandenburg: Motzen, east
of Zossen, 24 Jul 1967, Hudziok s.n. (neotype, here designated: HAL-076600!
3
sheets; isoneotypes, HAL! 3 sheets).
Oenothera
flaemingina
Hudziok,
Verh. Bot. Vereins
Prov. Brandenburg
105: 88.
1968.-TYPE: GERMANY.
Brandenburg:Juiterbog,
W of Tiefenbrunnen, 17 Jul
1967/A.,
Hudziok
s.n.
(holotype:
HAL-076596!
3 sheets;
isotype: HAL!
3
SYSTEMATICBOTANYMONOGRAPHS
98
sheets).
[Described
as a hybrid between
two phenotypes
VOLUME 50
of 0. biennis:
rubri
caulis x jueterbogensis.]
105: 89.
Oenothera
inconspecta Hudziok, Verh. Bot. Vereins Prov. Brandenburg
1968.-TYPE:
GERMANY. Brandenburg: Ludwigsfelde,
sandy places at railway,
31 Jul 1965, Hudziok s.n. (holotype: HAL, not located).
Oenothera mediomarchica
Hudziok, Verh. Bot. Vereins Prov. Brandenburg 105: 90.
roadside near Kummers
1968.-TYPE:
GERMANY. Brandenburg: Luckenwalde,
dorf, 1Aug 1967/A., Hudziok
HAL! 2 sheets).
Oenothera
s.n. (holotype: HAL-075225!
105: 94.
Hudziok, Verh. Bot. Vereins Prov. Brandenburg
GERMANY. Brandenburg: Luckenwalde, on disturbed sandy place,
obscurifolia
1968.-TYPE:
5 Jul 1967, Hudziok
s.n. (holotype: HAL-075214!
octolineata Hudziok, Verh. Bot. Vereins
Oenothera
1968.-TYPE:
GERMANY. Brandenburg: Teltow,
Aug
2 sheets; isotype:
1966, Hudziok
s.n. (holotype: HAL,
3 sheets).
Prov. Brandenburg
105: 89.
sandy place in Stahnsdorf, 5
not located).
105: 93.
Oenothera paradoxa Hudziok, Verh. Bot. Vereins Prov. Brandenburg
in
11 Jul
1968.-TYPE: GERMANY. Brandenburg: Zossen, sandy place Wunsdorf,
3 sheets; isotype: HAL! 2
s.n. (holotype: HAL-075228!
1967/A., Hudziok
sheets).
Oenothera pyramidiflora Hudziok, Verh. Bot. Vereins Prov. Brandenburg 105: 95.
1968.-TYPE:
GERMANY. Brandenburg: Zossen, roadside near Neuhof, 24 Jul
1967/A.,
Hudziok
s.n. (holotype:
HAL-076599!
3 sheets;
isotype: HAL!,
3
sheets).
Oenothera
ersteinensis
Linder & Jean, Bull. Soc. Bot. France 116: 523. 1969.
Erstein, at junction of national rd 83 with 426 (culti
TYPE: FRANCE. Bas-Rhin:
vated from seeds), Jul 1968, Jean s.n. (holotype: LILLE! 2 sheets, photo: MO!).
Sachs. Bot. 9: 71.
Oenothera punctulata Rostan'ski & Gutte, Ber. Arbeitsgem.
POLAND.Wroclaw: Nysa, 10 Jul 1963, Rostan'ski s.n. (holotype:
1971.-TYPE:
WRSL! 2 sheets; isotype: LZ!). [Described as a hybrid between two phenotypes
of 0. biennis: Biennis-II x chicaginensis
(=Biennis-I).]
cambrica Rostan'ski, Fragm. Florist. Geobot. 23: 285.
Oenothera
UNITED KINGDOM. WALES. Pembry, sandy areas at Carmarthen,
s.n. (holotype: KTU!; isotype: BM!).
McClintock
1977.-TYPE:
25 Sep 1970,
cambrica var. impunctata Rostan'ski, Fragm. Florist. Geobot. 23: 287.
Oenothera
1977.-TYPE: UNITED KINGDOM. WALES. Jersey Marine, Swansea (cultivated at
Botanical
Garden of Katowice,
Poland,
from seeds collected
in 1973 by D. Mc
Clintock), 15 Jul 1975, Rostan'ski 8/74 (holotype: KTU!).
carinthiaca Rostaniski, Fragm. Florist. Geobot. 23: 287. 1977.-TYPE:
AUSTRIA. Karnten, Villach, 16 Dec 1970, Melzer s.n. (holotype: KTU!).
rubricaulis var. dentifolia Jehll'k & Rostan'ski, Folia Geobot. Phytotax.
Oenothera
(Praha) 14: 398. 1979.-TYPE:
CZECH REPUBLIC. Central Bohemia, railway sta
Oenothera
tion of Neratovice,
165 m, 3 Jul 1972, Jehllk s.n. (holotype: PR).
Oenothera sesitensis Soldano, Atti Ist. Bot. Univ. Pavia 6, 13: 147. 1980 ['"1979"].
TYPE: ITALY.Region Piedmont: Prov. Vercelli, at river Sesia near Vercelli, 13
Aug & 16 Sep 1976, Soldano 1137 (holotype: PAy!).
chicaginensis var. minutiflora Rostan'ski & Jehlik, Folia Geobot. Phytotax.
CZECH REPUBLIC. Northern Bohemia, at River
(Praha) 14: 401. 1979.-TYPE:
Upa, 24 Aug 1973, Jehli'k 6649 (holotype: PR!; isotype: KTU!).
Oenothera
1997
OENOTHERA
99
var. fiedleri Gutte & Rostan'ski, Ber. Arbeitsgem. Sachs. Bot. 11:
GERMANY. Sachsen: Gundorf near Leipzig (cultivated from
seeds), 4 Sep 1970, Gutte s.n. (holotype: LZ-2397! 2 sheets). [A hybrid that is
phenotypically most similar to 0. biennis (BA).]
Oenothera
nissensis
185. 1981.-TYPE:
rubricaulis var. longistylis Gutte & Rostan'ski, Ber. Arbeitsgem.
Sachs.
Bot. 11: 187. 1981.-TYPE:
GERMANY. Sachsen: Dresden (cultivated from wild
Oenothera
collected
13 Jul 1968, Rostan'ski
rosettes),
13/67
(holotype: WRSL!;
isotype:
LZ!).
Oenothera
var. latipetala Soldano, Riv. Piem. St. Nat. 2: 237. 1981.
Piedmont: Prov. Vercelli, Arborio at river Sesia, 180 m, 4
suaveolens
TYPE: ITALY.Region
Jul 1979, Soldano
12677
(holotype: TO!).
[The type has the suaveolens
pheno
type.]
Oenothera
Soldano, Arch. Bot. Biogeogr.
Liguria: Prov. La Spezia, Marinella
Ital. 58: 178. 1982.-TYPE:
marinellae
ITALY.Region
di Sarzana, 6 Aug 1980, Sol
dano 3259 (holotype: FH!). [The type is phenotypically
like 0. biennis, but itmay
represent a hybrid between 0. biennis and another species of subsect. Oeno
thera.]
Oenothera
pellegrinii
Soldano, Arch. Bot. Biogeogr.
Ital. 58: 181. 1982.-TYPE:
ITALY.Region Tuscany: Prov. Massa-Carrara, Cinquale near Montignoso,
29 Jul
1976, Soldano s.n. (holotype: Fl!). [This entity is phenotypically
like 0. biennis
(AB), but itmay
Oenothera
Region
TO!).
represent a hybrid involving 0. biennis.]
Soldano, Riv. Piem. St. Nat. 4: 131. 1983.-TYPE:
pedemontana
Piedmont:
Prov. Torino, Saluggia,
[This entity is phenotypically
6 Sep 1980, Soldano
ITALY.
1617 (holotype:
like 0. biennis, but itmay represent a hybrid
involving0. biennis.]
rostanskii Jehlik, Fol. Geobot. Phytotax. 20: 439. 1985. Oenothera vic
Oenothera
torinii f. rostanski (Jehllk) Jehlik & Rostan'ski, Folia Geobot. Phytotax. (Praha)
30: 437.
1995.-TYPE:
CZECH REPUBLIC. Moravia: Thinec (cultivated from
seeds collected by V. Jehlifk in 1978), 14 Aug 1980, Jehlik 4484 (holotype: PR!
4 sheets; isotypes: BRNU, KTU).
Oenothera
chicaginensis
["1992"].-TYPE:
var. bartlettii
ITALY.Region
Grande, presso Calavorno,
Soldano,
Tuscany:
Nat.
Bresciana
Garfagnana,
15 Sep 1986, Marchetti
28:
105.
tra di Fegana
1993
e Piano
s.n. (holotype: PI).
Erect biennial herb, forming a rosette; stems 3-20 dm tall (taller in cultivation), un
branched or with side branches obliquely arising from the rosette or the main stem, green
or flushed with red in lower parts, sometimes the axis of the inflorescence red, densely to
sparsely strigillose and with longer somewhat appressed to spreading hairs which are
often pustulate, the axis of the inflorescence sometimes also glandular-puberulent
and/or
with pustulate hairs. Leaves usually green to pale green, both surfaces and margin strig
illose, the apical bracts sometimes also with spreading hairs and glandular-puberulent.
Rosette leaves 10-30 cm long, 2-5 cm wide, narrowly oblanceolate to oblanceolate, mar
gin dentate, sometimes bluntly so, the teeth sometimes widely spaced, the lower part
sometimes sinuate-dentate to somewhat lobed, or subentire, apex acute to narrowly ob
tuse, base gradually narrowed to the petiole. Cauline leaves 5-22 cm long, (1-) 1.5-5 (-6)
cm wide, narrowly oblanceolate
to oblanceolate or narrowly elliptic to elliptic, margin
dentate,
sometimes
bluntly
so, the teeth sometimes widely
spaced,
the lower part some
100
SYSTEMATICBOTANYMONOGRAPHS
VOLUME 50
to lobed, or subentire, apex acute to long-acute, base acute to atten
uate, short-petiolate or sessile. Bracts 1.2-5 cm long, 0.8-2.5 cm wide, narrowly lanceo
late to narrowly ovate, or narrowly elliptic, margin bluntly dentate to subentire, apex acute
times sinuate-dentate
to long-acute, base acute to narrowly cuneate, sessile, sometimes deciduous. Inflorescence
unbranched, or often with secondary lateral inflorescences below the main one. Floral
tube (2-) 2.5-4 cm long, 1-1.2 mm in diameter, yellowish green or flushed with red,
or sometimes also with long
sparsely villous and sparsely to densely glandular-puberulent
appressed hairs or with pustulate hairs. Mature buds 1-1.8 (-2.5) cm long, 3.5-6 mm in
diameter, narrowly lanceoloid to lanceoloid, or narrowly oblong. Sepals 1.2-2.2 (-2.8) cm
long, 3-5 mm wide, yellowish green, rarely flushed with red or red-striped, pubescence
like that of floral tube; free sepal tips 1.5-3 mm long, usually erect and appressed in bud,
sometimes somewhat divergent, strigillose or strigillose and glandular-puberulent.
Petals
1.2-2.5
(-3) cm long, 1.4-2.7
yellow,
(-3.2) cm wide, very broadly obovate, retuse to emarginate,
8-15 (-20) mm long; anthers 3-6 (-9) mm long;
cm long, 1.5-2 mm in diameter, pubescence vari
rarely pale yellow. Filaments
pollen ca. 50% fertile. Ovary 0.9-1.3
able: a) densely glandular-puberulent
and sparsely villous; b) densely glandular-puberu
lent and with scattered erect to somewhat appressed pustulate hairs; c) densely strigillose,
the hairs 0.2-1.5 mm long, and glandular-puberulent
toward apex; or d) strigillose, also
with erect to somewhat appressed pustulate hairs and glandular-puberulent.
Style 3-5.5
cm long, the exserted part 0.3-1.5 cm long; stigma surrounded by the anthers, which shed
pollen directly onto the lobes at anthesis, the lobes 3-6 mm long. Capsules 2-4 cm long,
4-6 mm in diameter, narrowly lanceoloid to lanceoloid, straight, tapering toward the apex,
fresh capsules green or red-striped, dry ones dull green, pubescence like that of ovary but
less dense; free tips of the valves distinct, 0.8-1.5 mm long, rounded to retuse. Seeds
1.1-2 mm long, 0.6-1.1 mm in diameter, brown to dark brown or nearly black. Chromo
some number: n = 7 (014; 012 and 1II;08 and 06; other configurations are not stable
and include 010 and
211;010 and 04; 08, 04, and 1II;08 and 311; 3 04 and 1II;based
on 133 individuals from 123 localities). Self-compatible,
usually autogamous, PTH. Fron
tispiece, Fig. 30.
Phenology. Flowering from July through September, and sometimes into October,
rarely as early as June.
Distribution
(Figs. 13, 20, 21, 31, 32, 33). Locally common, usually in open, dis
turbed sites, in Canada from southern Alberta, east to New Brunswick, Newfoundland,
and Nova Scotia, and in the United States from North Dakota south to eastern Texas, and
east to the Atlantic coast; occurrences inwestern North America west of the plains region
may
represent naturalized populations, either from the natural range or reintroductions
from naturalized populations from other parts of the world. Oenothera biennis is the most
common and widespread species of subsect. Oenothera. It has been introduced and is nat
uralized virtually worldwide
in temperate and subtropical regions. It is the only wide
spread species of Oenothera naturalized inRussia, Ukraine, and other countries of the for
mer Soviet Union.
Oenothera biennis is a PTH species with an AB or BA genomic constitution and plas
tome II or III. All strains investigated so far are autogamous and heterogamous;
i.e., they
transmit only one complex through the egg (oc) or pollen (f). Rarely both complexes are
transmitted as female, as happens occasionally
in certain strains of the suaveolens pheno
type (Stubbe 1959) and more often in the European phenotype rubricaulis (Renner 1950).
This species has been known as Biennis-I and Biennis-II in the cytogenetics literature
(Cleland
1972),
and most
recently
treated by Munz
(1965)
as comprising
two North
1997
101
OENOTHERA
mm
elff-
2.
3tb~~~~~~~~~~~~~~
e
f
I"I
_I?
FIG. 30. Oenothera
a. Inflorescence.
b. Rosette
;
(Cleland s.n., cult. DUSS-88-2005
[a-e]; Hoch 1843, cult. DUSS-88-2003
e-f. Inflorescence
leaf. c. Mid-cauline
leaf. d. Capsule.
pubescence.
biennis
'gp
[11).
102
SYSTEMATICBOTANYMONOGRAPHS
?e _
FIG. 31. North American
American
subspecies:
0. biennis
distribution
subsp. centralis
VOLUME 50
= km
of Oenothera
(Biennis-I)
biennis.
and 0. biennis
subsp. cae
ciarum (Biennis-II).
Previously the BB combination entity that we treat as 0. nutans was
included within 0. biennis, and referred to as Biennis-I1I by Cleland. Munz treated it as a
third North American
subspecies (0. biennis subsp. austromontana).
differences between Biennis-I and Biennis-II primarily relate to a re
and paternal transmission of the complexes.
In Biennis-I, which has a
the B (grandiflora) genome is transmitted through the egg,
combination,
The principal
versal of maternal
BA genomic
in the other race, Biennis-I1, which has an AB genomic constitution,
the B
through the pollen. Biennis-I has plastome III (Stubbe 1959) and
is only
apparently also sometimes plastome II (Drillisch 1975). By contrast, Biennis-I1
whereas
genome
is transmitted
known to have plastome II (Stubbe 1959). However, Cleland's studies (1972) suggested
that the type II and type III plastids could scarcely be differentiated. From his study of
over 3000 combinations
involving 0. biennis complexes he concluded
that, although
in plastid behavior are found among the various races, all 0. biennis plas
tids show very similar behavior.
minor variations
and Biennis-II
are weakly differentiated phenotypically
garden, but are recognizable only with difficulty or not at all
under natural conditions in the field (Cleland 1972, p. 279; Raven et al. 1979). The dif
ferences, as summarized by Munz (1965), are that his 0. biennis subsp. caeciarum is glan
Populations
when
grown
of Biennis-I
in a common
dular-puberulent in the inflorescence, the lower leaves are merely dentate, capsule valves
are retuse or entire, lower bracts are narrowly lanceolate and persistent to deciduous, and
stems are often flushed with red, whereas 0. biennis subsp. centralis is not glandular-pu
berulent
valves
in the inflorescence,
are entire,
the lower leaves are often lobed toward the base, capsule
lower bracts are narrowly ovate and persistent, and stems are usually
OENOTHERA
1997
103
5
O. biennis
0
FIG.
In general,
green.
ern United
ennis-I
Munz
States
from
ranges
Alberta
throughout the southeastern United
in Europe.
Munz
form
branched
has
beaklike
further
capsules
subtle features
pritnarily
and
leaves,
Oenothera.
in our
nent,
and
that
never
diverse
of
strains.
or
the other
the
has
between
inflorescence
these
features
stated
p. 227)
from
seeds
found
strains
from
the
stems,
variations
to find
impossible
forms.
biennis
within
collected
coworkers
from
different
and
localities,
grade
clear-cut
into
lines
one
of
the
species
"have
grown
rarely
from
contain
another
separation,
capsules,
of
subsect.
hundreds
across
localities
many
are
of Munz
subspecies
pubescence
and
and
red on
emarginate
are
variable
One
less
capsules with usually
two O.
and
pubescence,
and shape of mature
that he
often
two
into
inflorescences,
in appearance. A single locality may
These
Texas
and
form naturalized
sparse
simpler
Bi
whereas
Iowa,
caeciarum
subsp.
northeast
the
Carolina,
also is the common
on
red
as
(caeciarum)
to North
to Nebraska,
and scarcely-beaked
derived
identical
it is difficult
of
(1972,
garden,
have
ities, that were
cally
All
bracts.
Cleland
lines
and
valves;
in the phenotype
differences
biennis
abundant
the stems, abundant pubescence,
valves.
The
O.
in Europe.
south
and Michigan
his
subdivided
entire
and
States. Biennis-11
inflorescences,
with
biennis
for Biennis-11
range
to Ontario
west
Canada
of Oenothera
the
describes
and
(centralis)
32. Distribution
K
Km0*0_
the
nearby
of
conti
local
several phenotypi
so gradually,
and
so
however,
to be
able
to
one taxon from another."
distinguish
These morphological
races
are
so
slight,
especially
differences
in
among
the wild,
even
these genetically
though
each
similar
is maintained
sneries of clonal
by
autogamy
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
104
400S
+
17CPE
f f0
<
*
A_
p *
FIG. 33. Distribution
and PTH,
of Oenothera
400 Km
200
0. parviflora
0. biennis
biennis
that it does not seem appropriate
and 0. parviflora
to recognize
in New
Zealand.
them in the formal taxonomic
system.
treated here, 0. biennis consists of literally hundreds of minutely differing phe
races that are true-breeding, and recognizing any of them in the taxonomic sys
tem would lead to a never-ending description of them. We combine here all of these true
breeding strains that share common genome, plastome, and certain related morphological
traits into one polymorphic entity without further subdivision. We also include within the
widespread and polymorphic 0. biennis the many phenotypes, including many of those in
As
notypic
that have arisen subsequently through hybridization with other species of subsect.
Oenothera as long as those hybrids are BA or AB phenotypes with plastome II or III.
interesting series of populations of 0. biennis, which has not been
An exceptionally
this study, occurs in western North America. During the herbar
all
at
before
understood
Europe,
1997
OENOTHERA
105
ium study for this revision we detected several specimens, primarily identified by others
as 0. villosa subsp. strigosa (or some alternative names for this taxon) that were clearly
not typical 0. villosa subsp. strigosa, but we believed them to be an AA genomic combi
nation morphologically
resembling 0. biennis. During 1981-1987 we investigated these
phenotypes, and eventually determined that they closely resembled a phenotype recently
described from France, 0. ersteinensis. This taxon was described by Linder and Jean in
since
(dep. Bas-Rhin). It had been in cultivation in Dusseldorf
(from the type locality Erstein). Upon further investigation we learned that
from North
a similar phenotype was already in cultivation from an earlier collection
the North American
America
(Hood River Co., Oregon, Hoch 1843a-c). Comparing
1969 from eastern France
the late 1970's
the plants from Erstein, we realized that their phenotypes were extremely sim
ilar. Further study showed that other American strains were at least very similar to er
steinensis. Field studies were conducted by Wagner, and an experimental garden study
was started in 1983 at Dusseldorf by growing up all putative strains in order to perform a
plants with
full comparative analysis of these strains morphologically,
and genetically.
cytologically,
Surprisingly there were a considerable number of other strains of American origin that
were the same as the original ersteinensis strain. These strains are from British Columbia
(Wagner 4545, 4546, Merchant s.n. in 1981), Washington
(Wagner 4540, 4542, 4543, An
derson 3632), Oregon
(Wagner 4535, 4537, 4538, 4539, Stubbe s.n. in 1980), and Col
orado (Wagner 4532). Several other earlier strains also fit here, including ones from Utah
(Nye s.n. in 1975), New Mexico
(Wagner s.n. in 1975), and Idaho (Hoch s.n. in 1975, Cle
land "Cceur d'Alene"). Experimental hybridizations, evaluated in 1984, showed that all of
the strains had the AB genome rather than AA, which would have been suspected from
their phenotype, and thus represented a form of 0. biennis. This was demonstrated by the
following experiments. When one of the ersteinensis strains is crossed with 0. elata (AA),
the progeny have an AA phenotype, whereas when 0. grandiflora (BB) is crossed with
the descendants have a BB phenotype. The crossing experiments further
ersteinensis,
showed that all strains of ersteinensis have plastome II, a result which does not agree with
the BA genomic combination suggested by Linder and Jean (1969). In their publication
0. ersteinensis is also said to be associated with plastome III. But all hybrids between er
steinensis and 0. grandiflora (BB-III) proved to be pale green with white margins, which
not occur if ersteinensis had plastome III. Another strain that Cleland (1972, p.
340) listed as unclassified appears to represent this ersteinensis form of Oenothera bien
nis (Portland, Oregon). The segmental arrangement of the I complex of this strain is the
would
same as one complex of ersteinensis.
The most distinctive morphological
characters of ersteinensis are the dense pustulate
pubescence on stems and the intense, often dark red color of the stems and sepals, char
acters which are not typical for the eastern North America forms of 0. biennis. The pus
tulate pubescence of this form is also a characteristic feature of many 0. elata forms, sug
that this feature may have originated via hybridization with 0. elata or 0. villosa
subsp. strigosa. Our crossing studies confirm that one of these species was most likely in
volved. For example, 0. grandiflora (BB-III) crossed with 0. villosa subsp. strigosa (AA
I) yields a hybrid that, other than in its larger flowers, is a close match for an ersteinensis
gesting
phenotype.
Similarly, hybridization of 0. elata subsp. hookeri (AA) with ersteinensis
(AB) yields descendants with an ersteinensis-like
phenotype. The morphological
differ
ences between ersteinensis
and 0. biennis are not significant enough to consider er
steinensis a distinct species, especially considering the overall taxonomic philosophy that
we apply to the subsection. Thus, it is treated here as yet another part of 0. biennis. This
SYSTEMATICBOTANYMONOGRAPHS
106
is consistent with
decision
except
the distinctive
VOLUME 50
the inclusion of all other AB and BA genomic combinations,
has unusual features of its PTH system and
0. glazioviana, which
very different morphological
characteristics.
What was the origin of this form of 0. biennis? One of the seemingly odd features of
this form is its scattered distribution. At first this suggested a series of introductions to the
Pacific Northwest and other areas (Idaho, Colorado, New Mexico,
and Utah) of a form
that had arisen in Europe. Although the collections made during our study came from dis
turbed sites, it did not seem likely that the very scattered distribution would result from a
reintroduction from Europe. More likely, we think that this phenotype has arisen several
times independently through hybridization of 0. villosa subsp. strigosa with introduced
0. biennis. This scattered distribution of ersteinensis in the western United States and ad
jacent Canada may represent a hybridized form of 0. biennis derived from more typical
eastern North American 0. biennis. This could have occurred if 0. biennis had a wider
inwestern North America during the recent past, such as the last glacial max
imum, which now is only represented by relictual populations maintained at scattered sites
in a hybridized form. These former populations would have come into contact with the
more common and xerophytic 0. villosa subsp. strigosa, and hybrids between them prob
ably evolved into the ersteinensis forms growing in such sites as along rivers. They also
distribution
are known
from disturbed roadsides (which are more mesic than surrounding sites). The
form does not appear to be a recent product of hybridization, because, if it
were, there would be some evidence of more typical forms of 0. biennis inwestern North
America. We have tentatively included the specimens of this entity in the list of specimens
from the indigenous range.
ersteinensis
Oenothera
biennis
hybridizes with the other species of subsect. Oenothera with
in contact. Hybrids or intergrading forms with 0. villosa subsp. villosa
have been discussed under 0. villosa. Hybrids also are formed with 0. grandiflora, 0. nu
tans, 0. oakesiana, 0. parviflora, and 0. villosa subsp. strigosa.
There is a series of intermediate phenotypes between 0. biennis and 0. nutans in
which
it comes
and South Carolina. We have seen herbarium specimens of probable hybrids with
0. nutans from North Carolina (e.g., Stanley Co., Ahles 57147; Swain Co., Munz 13523),
which are similar in the type of pubescence present (a few appressed hairs), but are more
North
like 0. nutans. We place them under 0. biennis because they repre
and, as dried specimens, are not determinable as hybrids with cer
example of an intermediate between 0. nutans and 0. biennis, which we
inDusseldorf,
is from West Virginia, Randolph Co. (DUSS-79-0563).
The
glandular-puberulent
sent BA phenotypes
tainty. Another
have cultivated
is again similar to that of 0. nutans, but the bracts are not deciduous and the
leaves have the shape of those of 0. biennis. This strain breeds true as a PTH.
In Florida there are forms of 0. biennis with the deeply parted basal leaves more typ
ical of 0. grandiflora from the same region. As in the example above, these plants repre
sent BA phenotypes and are presumably PTH populations. Other hybrids of this combi
pubescence
nation have been found in Alabama
and are discussed in the notes under 0. grandiflora.
When hybrids between 0. biennis and 0. oakesiana (AC-IV) or 0. parvifiora (BC
IV) are preserved as herbarium specimens, they are virtually impossible to detect as hy
brids, and therefore are included under the species they most closely represent. As the pre
ceding paragraphs demonstrate,
hybridization
undoubtedly
represents an important
for the origin of new fixed phenotypic variations in the PTH species of sub
mechanism
sect. Oenothera.
The Biennis-I
and Biennis-II
forms of 0. biennis have been introduced
to Europe.
OENOTHERA
1997
Plants that have been referred to 0. biennis
(Biennis-II). Likewise,
(AB-II), as is the Linnaean
confused with
considerably
can genetics
Oenothera
placed
0.
this, the application of the name 0. muricata has been
the misapplication
of the name in the European and Ameri
for 0.
oakesiana, and sometimes for 0. parvifiora.
is a typical Biennis-I strain (BA-IlI) and would be
in 0. biennis subsp. centralis.
literature, sometimes
of Renner
chicaginensis
inMunz's
largely toMunz's 0. bi
rubricaulis Klebahn is another pheno
type of 0. muricata (LINN-484.3)
(Ros
in Europe correspond
ennis subsp. caeciarum
type of Biennis-II
tan'ski & Ellis 1979). Despite
107
taxonomy
In the wake of the introduction of several PTH species of subsect. Oenothera
rope, new stable PTH phenotypes
arose by hybridization
into Eu
that do not occur within
the na
tive North American
ranges of these species. The taxa that appear to have been involved
are primarily 0. biennis, 0. oakesiana, 0. parvifiora, and 0. villosa subsp. villosa (Table
4). Many of these have been named and are included in the section on hybrids. Only 0.
glazioviana
and 0. stucchii, which
represent such new combinations,
tive new features that we accord them species
sect. Oenothera.
In general
the chromosomal
configurations
are very variable compared with
nous range in North America.
and 2II; 010
and 04;
08,
be seen in the specimen
ual collections
04,
between
have such distinc
the formal taxonomy of sub
found in European
strains of 0. biennis
of a 014 within
the nearly stable occurrence
The variation
in the diakinesis
and 1II;08
and 3II; and 3 04
citations of cultivated plants where
are given. Many
result of hybridization
status within
configurations
the indige
include 010
and 11. This variation can
the configurations of individ
of these configurations
different 0. biennis
are not stable and probably are the
strains or even with other species re
sulting in an 0. biennis phenotype.
The presumably
true breeding
hybrids among the four species mentioned often ex
the range of variation defined by us for these species, and are thus
taxonomically. The sexual breeding system of many species of the genus
press phenotypes within
not relevant
Oenothera, characterized by complex heterozygosity
and a high level of self-fertilization
results in essentially clonal reproduction. If coupled with a narrow species concept, there
would be a flood of countless species, making practical application of the taxonomic sys
tem a puzzle and consequently
impossible. With the exception of 0. glazioviana,
there
fore, we assemble under 0. biennis all permanent structural heterozygous and autogamous
AB and BA combinations with plastome II or III.
10. Oenothera
glazioviana
Micheli
in Martius,
BRAZIL. Rio de Janeiro, Tijuca,
Fl. brasil.
7 Feb 1868, Glaziou
13(2):
178. 1875.-TYPE:
2568
(holotype: P!; iso
types: BR! Fl! G!).
Oenothera
albida de Vries, Rev. gen. bot. 13: 11. 1901.-TYPE:
Amsterdam
(cultivated). No authentic material
laterMutationstheorie
(1903) represents
NETHERLANDS. Near
seen; t. 3 opposite
p. 192 of the
this entity.
Oenothera
brevistylis de Vries, Mutationstheorie
1: 153, 223. 1901.-TYPE: NETHER
LANDS. Near Amsterdam
(cultivated). No authentic material seen; fig. 80 on p.
430 of Mutationstheorie
(vol. 2, 1903) represents this entity.
Oenothera
elliptica de Vries, Mutationstheorie
1: 156, 280. 1901.-TYPE:
NETHER
LANDS. Near Amsterdam (cultivated); fig. 83, p. 281 (lectotype, here designated).
[No authentic material
seen.]
108
SYSTEMATICBOTANYMONOGRAPHS
VOLUME 50
Oenothera
1: 301. 1901.-TYPE:
fatua de Vries, Mutationstheorie
NETHERLANDS.
Near Amsterdam
(cultivated); fig. 94, p. 301 (lectotype, here designated).
[No
authentic material seen.]
Oenothera gigas de Vries, Rev. gen. bot. 13: 11. 1901; Mutationstheorie
1: 158, 225.
1901.-TYPE:
NETHERLANDS. Near Amsterdam (cultivated). No authentic mate
rial seen; t. 2 inMutationstheorie
represents this entity.
Oenothera
1: 153, 218. 1901.-TYPE: NETHER
laevifolia de Vries, Mutationstheorie
LANDS. Near Hilversum
(cultivated); fig. 56, p. 218 (lectotype, here designated).
[No authentic material seen.]
lata de Vries, Rev. gen. bot. 13: 11. 1901; Mutationstheorie
1: 155, 168,
287. 1901.-TYPE:
NETHERLANDS. Near Amsterdam
(cultivated). No authentic
material seen; fig. 88 on p. 288 of Mutationstheorie
represents this entity.
Oenothera
Oenothera
de Vries, Mutationstheorie
1: 156, 250. 1901.-TYPE:
leptocarpa
NETHERLANDS. Near Amsterdam
(cultivated); no authentic material seen.
Oenothera nanella de Vries, Rev. gen. bot. 13: 12. 1901; Mutationstheorie
1: 155,
165, 225. 1901.-TYPE:
NETHERLANDS. Near Amsterdam
No
au
(cultivated).
thentic material
seen; fig. 45 on p. 165 of Mutationstheorie
represents this entity.
oblonga de Vries, Rev. gen. bot. 13: 11. 1901; Mutationstheorie
1: 163,
238. 1901.-TYPE:
NETHERLANDS. Near Amsterdam
(cultivated). No authentic
material seen; fig. 44 on p. 163 of Mutationstheorie
represents this entity.
Oenothera
Oenothera
rubrinervis de Vries, Rev. gen. bot. 13: 11. 1901; Mutationstheorie
1: 155,
1901.-TYPE:
NETHERLANDS. Near Amsterdam
(cultivated). No au
thentic material seen; fig. 70 on p. 237 of Mutationstheorie
represents this entity.
Oenothera scintillans de Vries, Rev. gen. bot. 13: 12. 1901; Mutationstheorie
1: 170,
268. 1901.-TYPE:
NETHERLANDS. Near Amsterdam
(cultivated). No authentic
161, 231.
material seen; fig. 47 on p. 171 of Mutationstheorie
represents this entity.
Oenothera semilata de Vries, Mutationstheorie
1: 156, 254. 1901.-TYPE:
NETHER
LANDS. Near Amsterdam
(cultivated); no authentic material seen.
Oenothera
sublinearis
de Vries, Mutationstheorie
1: 156, 285. 1901.-TYPE:
NETHERLANDS. Near Amsterdam
(cultivated); fig. 85, p. 285 (lectotype, here
[No authentic material seen.]
designated).
Oenothera subovata de Vries, Mutationstheorie
1: 156, 301, 303. 1901. Fig. 86, p.
303 (lectotype, here designated). [No authentic material seen.]
Onagra erythrosepala Borbas, Kert 1902: 202. 1902. Oenothera erythrosepala (Bor
ba's) Borbas, Magyar Bot. Lap. 2: 245. 1903. Oenothera
suaveolens
f. ery
throsepala (Borbas) Javorka, Magyar
fl. 2: 748. 1924. Oenothera grandiflora
subsp. erythrosepala
(Borba's) Love & Love, Opera Bot. 5: 258. 1961.-TYPE:
HUNGARY. Budapest, Rakos, near the new cemetery, in sandy fields, 22 Jun
1899, de Borba's s.n. (lectotype, here designated: BP!).)
Oenothera
rubricalyx Gates, Annual Rep. Missouri Bot. Gard. 20: 133. 1909.
TYPE: No original material seen; this entity, which usually breeds true, originated
inGates's cultures of 0. rubrinervis obtained in 1907 from de Vries, 1934, Gates
s.n. (neotype, here designated: K!).)
multiflora Gates, Ann. Missouri
LAND. Cheshire: vicinity of Birkenhead
Oenothera
Bot. Gard.
(cultivated
1: 386. 1914.-TYPE:
ENG
from seeds collected by D. T.
in 1907); fig. 3 of plate 20 (lectotype, here designated). [No authen
seen. Gates indicated that there were specimens deposited atMO col
lected in 1909 and at BM collected in 1912.]
MacDougal
ticmaterial
OENOTHERA
1997
109
Oenothera multiflora var. elliptica Gates, Ann. Missouri Bot. Gard. 1: 387. 1914.
TYPE: ENGLAND. Cheshire: vicinity of Birkenhead
(cultivated from seeds col
lected by D. T. MacDougal
in 1907); fig. 4 of plate 20 (lectotype, here desig
nated).
[No authentic material seen.]
rubrinervoides Gates, Ann. Missouri
ENGLAND. Cheshire: vicinity of Birkenhead
Oenothera
D. T. MacDougal
authentic material
Bot. Gard. 1: 389. 1914.-TYPE:
(cultivated from seeds collected by
in 1907); fig. 10 of plate 21 (lectotype, here designated).
[No
seen.]
Oenothera
tardiflora Gates, Ann. Missouri Bot. Gard. 1: 391. 1914.-TYPE:
ENG
LAND. Cheshire: vicinity of Birkenhead (cultivated from seeds collected by D. T.
in 1907; fig. 17 of plate 22 (lectotype, here designated).
MacDougal
[No au
thentic material seen.]
Oenothera rubritincta Gates, Ann. Missouri Bot. Gard. 1: 391. 1914.-TYPE:
ENG
LAND. Cheshire: vicinity of Birkenhead (cultivated from seeds collected by D. T.
MacDougal
in 1907); fig. 16 of plate 22 (lectotype, here designated).
[No au
thentic material seen.]
cana de Vries,
Oenothera
Bot.
NETHERLANDS. Near Hilversum
Gaz.
62: 250. 1916.-TYPE:
(Crawfordsville)
(cultivated); the plant on the right-hand side of
fig. 2, p. 253 (lectotype, here designated). [No authentic material seen.]
Oenothera pallescens
de Vries, Bot. Gaz. (Crawfordsville)
62: 260. 1916.-TYPE:
NETHERLANDS.Without locality (cultivated); fig. 3, p. 261 (lectotype, here des
ignated).
[No authentic material
seen.]
Oenothera superfluade Vries, Bot. Gaz. (Crawfordsville)62: 270. 1916.-TYPE:
NETHERLANDS.Without locality (cultivated). No authentic material seen; no fig
ure provided; disposition based on description.
Oenothera aberrans Lutz, Amer. J. Bot. 3: 512. 1916.-TYPE:
U.S.A. New York:
Suffolk Co., cultivated at Cold Springs Harbor by A. M. Lutz; fig. 5, p. 512 (lec
totype, here designated).
[No authentic material seen.]
Oenothera plicatula Lutz, Amer. J. Bot. 3: 505. 1916.-TYPE:
U.S.A. New York:
Suffolk Co., cultivated
at Cold Springs Harbor by A. M. Lutz; fig. 1, p. 506 (lec
[No authentic material seen.]
Oenothera bipartita Lutz, Amer. J. Bot. 4: 62. 1917.-TYPE: BELGIUM. Brabant: Cul
tivated at theUniversity of Louvain [now Universite Catholique de Louvain] by A.
M. Lutz; fig. 1, p. 63 (lectotype, here designated). [No authentic material seen.]
totype, here designated).
Oenothera
blandina de Vries, Bot. Gaz. (Crawfordsville)
63: 2. 1917.-TYPE:
NETHERLANDS. Near Amsterdam
(cultivated); the color plate, t. 1, right-hand
plant opposite p. 24 (lectotype, here designated).
[No authentic material seen.
Oenothera blandina is a complex homozygote AA combination derived from 0.
glazioviana found in deVries's experimental field. To avoid confusion it is placed
here under synonymy of 0. glazioviana.]
Oenothera simplex de Vries, Ber. Deutsch. Bot. Ges. 37: 65. 1919.-TYPE:
NETHER
LANDS.Without locality (cultivated); t. 2 after p. 351 in de Vries, Z. Indukt. Ab
31: 1923 (neotype, here designated).
stammungs.-Vererbungsl.
[No authentic
material seen.]
Oenothera
liquida de Vries, Bot. Gaz. (Crawfordsville)
62: 268. 1916 [nomen
nudum];Z. Indukt.Abstammungs-Vererbungsl.35: 212. 1924 [description].
TYPE: NETHERLANDS.Without
designated).
locality (cultivated);
[No authentic material
seen.]
fig. 4, p. 205 (lectotype, here
110
SYSTEMATIC
BOTANY
MONOGRAPHS
VOLUME50
Oenothera hamata de Vries, Z. Indukt. Abstammungs-Vererbungsl.
35: 216. 1924;
fig. 10, p. 217 (lectotype, here designated).
[No authentic material seen.]
candicans de Vries, Z. Indukt. Abstammungs-Vererbungsl.
Oenothera
35: 216.
NETHERLANDS.Without locality (cultivated); fig. 9, p. 214 (lecto
1924.-TYPE:
type, here designated). [No authentic material seen.]
Oenothera militaris de Vries, Z. Bot. 17: 194. 1925.-TYPE: NETHERLANDS.Without
locality (cultivated). No authentic material seen; no figure provided; disposition
based on description.
pulla de Vries, Z. Bot. 17: 206. 1925.-TYPE: NETHERLANDS.Without lo
(cultivated); fig. 2, p. 266 in de Vries & Boedijn, Bot. Gaz. (Craw
fordsville) 78: 1924 (lectotype, here designated).
[No authentic material seen.
The figure was not explicitly cited by de Vries, but he did specifically refer to the
Oenothera
cality
1924 article.]
scindens de Vries, Bot. Gaz. (Crawfordsville)
80: 265. 1925.-TYPE:
NETHERLANDS. No authentic material seen; no figure provided; disposition based
Oenothera
on description.
tarda de Vries, Bot. Gaz. (Crawfordsville)
80: 272. 1925.-TYPE:
NETHERLANDS.Without locality (cultivated). No authentic material seen; no fig
ure provided; disposition based on description.
Oenothera
fusiformis Munz & I.M. Johnston, Contr. Gray Herb. 75: 21. 1925.
TYPE: ECUADOR. Loja: between El Tambo and La Toma, 1000-2000 m, 3 Sep
Oenothera
1923,Hitchcock 21350 (holotype:US-1196309!; isotypes:GH! NY!)
coronifera Renner, Planta 47: 239. 1956. Oenothera grandiflora subsp.
coronifera (Renner) Weihe in Garcke, Ill. Fl. Deutschland
23 ed., 982. 1972.
TYPE: GERMANY. Brandenburg:
Distr. Potsdam,
near railway station of
Monastery Zinna (cultivated from seeds originally collected by Renner in Jul
Oenothera
1936), 1967, Rossmann 91/66 (neotype, here designated: M!). [No authentic ma
terial seen. Renner apparently did not prepare a voucher nor did he designate a
type. The Rossmann collection, which derives from the original material, is des
ignated as neotype.]
Oenothera
var. azorica Rostan'ski, Bol. Soc. Brot. 64: 28. 1991.
erythrosepala
TYPE: PORTUGAL.Azores:
hos, 1964, Dansereau,
Santa Maria
[Faial], Capelo,
da Silva & Rainha
near Farol dos Capelin
483 (holotype: LISE-70343;
isotype:
NY!).
Erect biennial to short-lived perennial herb with a taproot, forming a rosette; stems
dm tall, usually obliquely branched from the rosette and with secondary branches
from main stem, densely to sparsely strigillose and with numerous long erect to suberect
5-15
red-pustulate hairs, and in the region of the inflorescence also glandular-puberulent
and
with only a few appressed hairs. Leaves dark to bright green, white- or red-veined, sur
face usually conspicuously
crinkled, villous to strigillose on both surfaces and margins,
bracts in apical part of inflorescence also glandular-puberulent. Rosette leaves 13-30 cm
long, 3-5 cm wide, narrowly oblanceolate
to oblanceolate, margin remotely and bluntly
dentate, toward the base usually sinuate-dentate, apex acute to subobtuse, base attenuate
to the petiole. Cauline leaves 5-15 cm long, 2.5-4 cm wide, narrowly elliptic to lanceo
late, margin remotely and bluntly dentate to regularly dentate, apex acute to subobtuse,
base usually abruptly narrowed to the petiole, those toward the apical part of plant nar
rowly cuneate, sessile. Bracts 1-3 cm long, 0.7-3.2
cm wide, lanceolate to narrowly
1997
OENOTHERA
ovate, green, margin
rowly cuneate.
111
remotely and indistinctly dentate, apex acute, base rounded to nar
unbranched. Floral tube 3.5-5 cm long, 1-1.2 mm in diam
Inflorescence
eter, sparsely villous with some pustulate hairs and densely glandular-puberulent. Mature
buds (2.5-) 3-4 cm long, 7-9 mm in diameter, narrowly lanceoloid to lanceoloid. Sepals
2.8-4.5 cm long, 5-7 mm wide, yellowish green, usually flushed with red or red-striped,
sometimes entirely very dark red, pubescence
like that of floral tube; free sepal tips 5-8
mm long, densely villous. Petals 3.5-5 cm long, 3.5-5.3 cm wide, very broadly obovate,
1.7-2.5 cm long; anthers 10-12 mm long; pollen ca. 50% fer
tile. Ovary 0.7-1.2 cm long, 1.5-2 mm in diameter, densely to moderately villous with
many long red- to dark red-pustulate hairs and densely glandular-puberulent.
Style 5-8 cm
long, the exserted part 2-3.5 cm long; stigma elevated above the anthers at anthesis, the
lobes 5-8 mm long. Capsules 2-3.5 cm long, 5-6 mm in diameter, narrowly lanceoloid,
retuse, yellow. Filaments
tapering toward the apex, green or with a red stripe on each valve when fresh, pubescence
like that of ovary but less dense; free tips of the valves 1-1.5 mm long, truncate to retuse.
Seeds 1.3-2 mm long, 1-1.5 mm in diameter, brown to dark brown, up to ca. 50%
abortive. Chromosome
number: n = 7 (012 and 1II; one other configuration found [010
is not a stable one; based on 25 individuals
211]
regularly outcrossing, PTH. Fig. 34.
and
Phenology.
Flowering
from 20 localities).
from July through September,
and sometimes
Self-compatible,
into October.
Distribution (Fig. 35). Oenothera glazioviana originatedvia hybridizationbetween
two cultivated or naturalized species in Europe, and was introduced into the horticultural
trade by Carter and Company (England) in 1860 (Cleland 1972; Raven et al. 1979). The
oldest name applied to this entity was based on plants cultivated inRio de Janeiro in 1868,
and thus clearly 0. glazioviana must have spread very rapidly. Now it is widely dispersed
inNorth and South America, Europe, Asia, Africa, and Australia. It is found in open dis
turbed sites such as roadsides, gardens, fallow fields, and along railroad right-of-ways.
Oenothera glazioviana, better known as 0. erythrosepala, or by the misapplied name
lamarckiana, has an AB genomic constitution and plastome III (Stubbe 1959, 1964). It
in
nearly invariably forms 0)12 and
I. It is the only regularly out
III meiotic metaphase
crossing PTH species in the family Onagraceae
(Raven et al. 1979). Although
it has the
same genome and plastome composition as 0. biennis (Biennis-I), it is quite distinct from
0.
it, especially
in its larger flowers, elevated stigma, sepals usually strongly red or flushed
red, and usually conspicuously
crinkled leaves toward the apex of plant and bracts.
Superficially, it ismost similar morphologically
to 0. grandiflora and 0. elata, but it can
be distinguished from both by its crinkled leaves, from the latter by its broader leaves and
with
the absence of strigillose pubescence in the inflorescence, and from the former in its pus
tulate hairs on the ovary and floral tube, and reddish green to red sepals.
Oenothera glazioviana
is accorded specific status, despite its unusual origin as a sta
bilized hybrid in Europe, because
it has very distinctive morphological
features, and has,
through cultivation, become widely spread around the world. In contrast to the
other numerous situations of perpetuated stabilized hybrids or unique phenotypes that we
do not give formal taxonomic recognition, we give species status to 0. glazioviana be
cause its strikingly distinctive morphological
features would be hard to accommodate
in
the other AB combination species, 0. biennis. The PTH system of 0. glazioviana also has
features not otherwise occurring in 0. biennis, including regularly outcrossing flowers, an
invariant meiotic configuration of 0)12 and
and nearly 50% seed abortion.
1Iil
This species is well known in the genetics literature under the name Oenothera
partly
lamarckiana.
In fact, it was
this species
that Hugo
de Vries
found in a potato field near
112
SYSTEMATICBOTANYMONOGRAPHS
mm
1 3
a,b,c
dw
30
VOLUME 50
--
d
FIG. 34. Cenothera glazioviana (Ld. Sem.Bot. Gard. Salburg 84 no.
J1OS, cult.DUSS-88-2012). a. In
florescence.b. Rosette leaf. c.Mid-cauline leaf. d. Capsule. e. Inflorescencepubescence.
1997
113
OENOTHERA
0. glazioviana
0
FIG. 35. World
Hilversum,
2000
distribution
4000
6000 Km
of Oenothera
glazioviana.
formed the basis of his life-long work on his mu
that work he provided many "mutants" with names, as
in 1886, and which
Holland,
tation theory. In conjunction with
list above and in the many
can be seen in the synonymy
names pre
invalidly published
sented in a section on this topic at the end of the paper.
Oenothera
complex
is half-heterogamous.
glazioviana
(velans).
It is also occasionally
AA phenotype
selfed the structural homozygous
Oenothera
so that when
of de Vries) may
(deserens or decipiens
transmitted only through the pollen, because
is successfully
glazioviana
of 0. glazio
the structural
is lethal as abortive seeds (W. Stubbe, pers. comm.).
BB combination
homozygote
is transmitted as the at
through the pollen,
is similar to 0. elata subsp. hookeri. The B genome
arise. This phenotype
viana (gaudens)
The A genome
transmitted
is known
to hybridize with a number of the other taxa: 0. bi
and 0. wolfii. Hybrids between 0. glazioviana
and 0.
oakesiana or 0. parviflora were made experimentally by de Vries, but do not occur in nat
ural situations. Some of these hybrids have been named either as species or as hybrids, es
ennis, 0. villosa
subsp. villosa,
in Europe. Oenothera
pecially
xconferta,
are three of the
and 0. xbritannica
0. xfallax,
three are hybrids between 0. biennis and 0. glazioviana. Full
listing of the hybrids is given in the section on hybrids later in this paper. Most of the hy
brids exhibit some intermediate characteristics, but many are very difficult to distinguish,
better known examples. All
especially
posed
as pressed
It is significant
specimens.
a threat to the continued
existence
that hybridization
of that rare species
with
0. wolfii has
under 0.
(see discussion
wolfii, no. 4).
11. Oenothera
cola
West
Mackenzie,
argillicola
(Mackenzie)
Virginia:
Mackenzie
J. Boivin,
Greenbrier
Torreya 4: 56. 1904. Oenothera
Naturaliste
Co.,
373 (holotype: NY!,
Canad.
near White
photo BH!;
biennis
93: 644. 1966.-TYPE:
f. argilli
U.S.A.
Sulphur Springs, 27 Aug 1903,
isotypes: GH! IND! MO! POM!).
SYSTEMATICBOTANYMONOGRAPHS
114
Oenothera
argillicola
var. pubescens
VOLUME 50
Core & H. A. Davis, Castanea 18: 31. 1953.
Co., Great Cacapon, 27 Jul 1939, Davis &
TYPE: U.S.A. West Virginia: Morgan
Davis
3084
(holotype: WVA!).
Erect to suberect bushy biennial (or short-lived perennial) herb with a taproot, form
ing a rosette; stems up to 4 dm tall, usually obliquely ascending, green or red, recurved
toward the apex, but the tip ascending, unbranched or with branches obliquely arising
from the rosette and with side branches arising in the apical half of the main stem, these
sometimes branched again, the branches widely spreading, densely strigillose, also with
some longer subappressed hairs, these sometimes with a red or green pustulate base,
sometimes sparsely pubescent to glabrous toward and in the inflorescence. Leaves dark
green, upper surface somewhat glossy, both surfaces and margins strigillose, usually
glabrate with age. Rosette
late to narrowly
leaves 7-25 cm long, 0.7-2 cm wide, very narrowly oblanceo
or nearly linear, margins remotely and bluntly dentate,
oblanceolate
larger teeth near base, apex acute, base gradually narrowed to the petiole.
Cauline leaves 6-13 cm long, 0.4-1 cm wide, narrowly lanceolate to linear-oblanceolate,
linear-elliptic or linear, margins remotely and bluntly dentate to subentire, often sinuate
sometimes with
dentate near base, apex acute, base gradually narrowed to a short petiole or subsessile.
1.5-5.5 cm long, 0.3-0.7 cm wide, narrowly lanceolate to linear-lanceolate or lin
ear-elliptic, margins strigillose or glabrous, remotely and bluntly denticulate to subentire,
Bracts
apex acute to narrowly acute, base obtuse to cuneate, sessile. Inflorescence unbranched,
apical part recurved with the tip ascending, the flowers widely spreading from or perpen
dicular to the stem. Floral tube 3.2-5.2 cm long, 1-1.3 mm in diameter, yellow or flushed
with red to entirely red, glabrous or sparsely glandular-puberulent,
sometimes also with
long spreading hairs. Mature
oloid
to lanceoloid.
buds 2.5-3.5
Sepals 2.7-3.8
cm long, 4-8 mm
in diameter, narrowly
cm long, 3-7 mm wide,
yellowish
green
lance
to yellow,
sometimes flushed with red especially at apex, glabrous or sparsely glandular-puberulent
and sparsely long-villous; free sepal tips 3-9 mm long, subterminal in bud, divergent and
hornlike, glabrous or strigillose. Petals 2.5-4.2 cm long, 2.7-4.5 cm wide, very broadly
obovate, retuse to truncate, yellow to pale yellow. Filaments 2-2.7 cm long; anthers 9-13
mm long; pollen 90-100% fertile. Ovary 0.8-1.3 cm long, 1.5-2 mm in diameter, either
and densely to sparsely strigillose and with
a) glabrous, b) sparsely glandular-puberulent
a few pustulate hairs, c) sparsely strigillose and with a few longer appressed hairs, or d)
with
scattered longer appressed hairs. Style 6-8.5 cm long, the exserted part 2.5-4 cm
long; stigma elevated above the anthers at anthesis, the lobes 3-6 mm long. Capsules 2-4
cm long, 4-6 mm in diameter, narrowly lanceoloid to lanceoloid, spreading at an acute or
right angle from the stem, arcuate upward, sometimes secund, long-attenuate toward the
apex, pubescence
like that of ovary but less dense, green or red-striped when
green or rusty brown when
dry; free tips of the valves distinct,
1-2 mm
fresh, dull
long, truncate to
emarginate. Seeds 1.3-1.9 mm long, 0.7-1.1 mm in diameter, dark brown. Chromosome
number: n = 7
04 and 5II; 06 and
2 04 and
(711;
411;
311;08 and 311;010 and 21, [Stin
son 1953]; based on 18 individuals from 5 localities). Self-compatible,
mostly outcross
ing. Fig. 36.
Phenology.
Flowering
from July through October,
rarely as early as June.
Distribution
(Fig. 37). Occurring on open Brallier shale slopes, barrens, outcrops, or
from south-central
adjacent roadsides in the mid-Appalachian
Allegheny Mountains,
Pennsylvania
through western Maryland, western Virginia, and eastern West Virginia.
Oenothera argillicola is one of eight angiosperm species restricted to these shale barrens,
/'
~~~~mm
/
.
.
I~~~~~~~tg
Ad
a.~~~~~~~~~~~~~~~
f
.b~~~~~:
FIG. 6. Oeothea argllicoa
FiG.auin36a. Qenothrac.e
Cargillcol
(Subbe
(Sfbbgs.
.n. i 197,
Infl
cul. DU
1979,ecultDub
S-86/8-101a
[ad,
g] Brow
s.n
s
S-68-02cadegnBncein17,cl.DS-8-01[-..a.Ifoecne
in 176,
clt.
U
SYSTEMATICBOTANYMONOGRAPHS
116
0
VOLUME 50
5~~~~~~~~
je
0S
*.\
I
}\/^'.
g/
4r\;K
*
O. argillicola
V* oakesiana
355N
755W
500km
0
FIG. 37. Distribution
but among
of Oenothera
4_
~~~~~~~5
them only
0. argillicola
argillicola
and indigenous
distribution
and Trifolium virginicum
of 0. oakesiana.
occur throughout
/
the shale
barren region (Platt 1951).
is a distinctive, outcrossing, bivalent-forming
species. It has a
Oenothera argillicola
CC genomic combination, and it is the only species with plastome V. Most plants form 7II
has been
in meiotic metaphase
I, but some diversity of chromosomal end-arrangements
detected. We have observed all of the arrangements reported except (010 and 211,which
variabil
was reported in Stinson's (1953) extensive study of this species. Morphological
In
to
in
0.
in
similar
that
general,
and
is
extensive
range
occurring
grandifiora.
ity is not
from glabrous to densely strigillose inflores
is primarily in pubescence:
the
occasional presence throughout the range of
cences. Another variable characteristic is
the species of scattered long hairs on the ovary, sepals, and floral tube, which was the basis
the variation
var. pubescens.
very distinctive within subsect. Oenothera,
Oenothera argillicola ismorphologically
in its bushy habit, obliquely ascending stems, very narrow glossy leaves,
especially
widely spreading branches, sigmoid inflorescence apex, subterminal free sepal tips 3-9
mm long, and the arcuate, attenuate, widely spreading capsules. It is likewise ecologically
of 0. argillicola
on Devonian
shales of the Brallier Formation. These
specialized, growing exclusively
poor-soil habitats have open vegetation, and are exposed to much higher sunlight levels
than adjacent habitats (Platt 1951; Wherry 1930, 1933).
t'7?
OENOTHERA
1997
117
Three other species of subsect. Oenothera, 0. biennis (both Biennis-I and Biennis-II
forms), 0. nutans, and 0. parviflora, occur within the range of 0. argillicola. Of these,
only 0. parviflora actually grows at the same general localities with 0. argillicola,
al
though it usually grows on adjacent non-shale sites. No hybrids between 0. argillicola
and either 0. biennis or 0. nutans have been detected. Part of the explanation is that they
do not grow closely together. More importantly, however, crosses in both directions be
tween these species yield hybrids exhibiting incompatibilities between genome and plas
tome (Stubbe 1959), which are difficult to grow under optimal conditions, and would not
be expected to survive in the wild. The only exceptions to this pattern are experimental
crosses between 0. argillicola
(CC-V) and the Biennis-I1 form of 0. biennis (AB-II),
which yield viable hybrids with AC-I/V
genome/plastome
constitution; none have been
observed in the wild.
In Dusseldorf we have
Only 0. parviflora hybridizes regularly with 0. argillicola.
for several years wild-collected
small-flowered CC-phenotypes
from Highland
Co., Virginia, and Mineral Co., West Virginia, which exhibit various small chromosome
rings inmeiosis. These apparently represent natural hybrids between 0. parviflora and 0.
argillicola. Their flowers are intermediate in size. These plants can be interpreted only as
cultivated
hybrids with 0. argillicola
as the female parent and 0. parviflora as the male parent, be
cause this is the only way to obtain the CC combination. These plants would have plas
tome IV or V, and either would be a normal green in this combination. In addition to these
putative hybrids, the wild-collected
seeds sown in Dusseldorf yielded several individuals
larger flowers, which presumably represent natural back-crosses
to 0. argillicola.
The small-flowered CC combination hybrids yield exclusively
de
small-flowered
scendants. Currently, we do not know if the chromosome rings observed in the first gen
with
eration would be stable in subsequent generations or if self-pollinated
individuals would
yield some plants with
It is possible that plants such as these could represent an evo
711.
lutionary trend toward a new CC combination PTH species.
Reciprocal hybrids with 0. parviflora as the female apparently do not occur. We have
never observed anywhere within the range of 0. argillicola any BC combination individ
uals that formed small rings of chromosomes. All individuals tested had 014.
12. Oenothera
oakesiana
(A. Gray) J.W. Robbins ex S. Watson & Coulter, Manual, ed.
6. 190. 1890. Oenothera biennis var. oakesiana A. Gray, Manual, ed. 5. 178.
1867. Onagra oakesiana (A. Gray) Britton, Mem. Torrey Bot. Club 5: 233. 1894.
Oenothera parviflora var. oakesiana (A. Gray) Fernald, Rhodora 51: 66. 1949.
TYPE: U.S.A. Massachusetts: Bristol Co., Norton (cultivated from seeds collected
at Apponaganset),
Aug 1865, Robbins s.n. (lectotype, here designated: GH!).
[This specimen annotated by Robbins is surely the basis for A. Gray's parenthet
ical comment "Oe. oakesiana Robbins" and thus is selected as the lectotype.]
Oenothera
ammophila Focke, Abh. Naturwiss. Vereine Bremen
18: 183. 1906.
Oenothera muricata
subsp. ammophila
(Focke) Stomps, Receuil Trav. Bot.
Neerl. 41: 142. 1948 [combination
also proposed by Tischler, Chromos.
Gefaisspfl. Mitteleur. 58. 1950]. Oenothera parviflora subsp. ammophila (Focke)
Janchen, Phyton
(Horn) 3: 7. 1951.-TYPE:
GERMANY. Niedersachsen:
Isle of
Wangerooge,
Jul 1902, Focke s.n. (lectotype, here designated: BREM!). This en
tity is the coastal form in Europe.
Oenothera millersii
de Vries, Gruppenweise Artbildung 59. 1913.-TYPE: U.S.A. In
diana: Lake, La Porte, or Porter Co., Miller's Station at Lake Michigan
(culti
SYSTEMATIC
BOTANY
MONOGRAPHS
118
VOLUME50
inWashington
from seeds collected by H. de Vries), 1915, Bartlett s.n.
(neotype, here designated: MO-3838394!).
[No authentic material located; there
fore the Bartlett specimen (from Cleland's material now housed atMO), which
derives from the original collection, is designated here as a neotype.]
Oenothera cymatilis Bartlett, Cybele Columb. 1: 51. 1914. Oenothera canovirens var.
vated
cymatilis (Bartlett) Gates, Rhodora 59: 14. 1957.-TYPE:
U.S.A. Michigan:
Berrien Co., Sawyer (cultivated from seeds collected by W. Pfeiffer), 1913,
Bartlett 3665 (lectotype, here designated: MICH! 3 sheets).
Oenothera insignis Bartlett, Cybele Columb. 1: 52. 1914.-TYPE: U.S.A. Minnesota:
St. Louis Co., along sandy shore of Lake Superior atMinnesota Point near Du
luth (cultivated
from seeds collected
here designated: MICH!
by C. A. Davis),
Bartlett
3583
(lectotype,
2 sheets).
1: 48. 1914. Oenothera
litorea Bartlett, Cybele Columb.
syrticola var.
litorea (Bartlett) Gates, Rhodora 59: 16. 1957.-TYPE: U.S.A. Connecticut: New
Haven Co., seashore of Orange (cultivated from seeds collected by G. E.
Nichols), Bartlett 3606 (lectotype, here designated: MICH! 2 sheets).
Oenothera
rubescens Bartlett, Cybele Columb. 1: 50. 1914.-TYPE:
U.S.A. Massa
Oenothera
chusetts: Nantucket
B. Gardner), Bartlett
Co., Nantucket Island (cultivated from seeds collected by G.
3594 (holotype: MICH! 2 sheets; isotype: RSA!).
stenopetala Bicknell, Bull. Torrey Bot. Club 41: 79. 1914. Oenothera cru
ciata var. stenopetala (Bicknell) Fernald, Rhodora 51: 67. 1949. Oenothera bi
ennis f. stenopetala (Bicknell) J. Boivin, Naturaliste Canad. 93: 644. 1966.
Oenothera
TYPE: U.S.A.
embankment
Massachusetts:
Nantucket
beyond Orange Street, 15 Aug
Co.,
Nantucket
1906, Bicknell
Island, railroad
s.n. (holotype: NY!;
isotype:GH!).
Oenothera
tidestromii Bartlett, Cybele Columb. 1: 54. 1914. Oenothera oakesiana
var. tidestromii (Bartlett) Gates, Rhodora 59: 14. 1957.-TYPE:
U.S.A. Mary
land: St. Mary's Co., mouth of Patuxent River between Millstone & Piney Points
(cultivated from seeds collected by Bartlett in 1911), Bartlett 3672 (lectotype,
here designated: MICH!).
Oenothera muricata var. parvifiora Gates, Mutation factor in evolution 25. 1915.
TYPE:CANADA.
Quebec: Anticosti, JupiterRiver, 1883,Macoun s.n. (holotype:
BM).
Boedijn, Z. Indukt. Abstammungs-Vererbungsl.
32: 360. 1924.
ammophila var. germanica (Boedijn) Renner, Flora 131: 222. 1937.
Oenothera muricata
subsp. germanica
(Boedijn) Stomps, Receuil Trav. Bot.
Neerl. 41: 140. 1948. Oenothera parviflora subsp. germanica (Boedijn) Janchen,
Phyton (Horn) 3: 7. 1951. Oenothera ammophila subsp. germanica
(Boedijn)
Oenothera
germanica
Oenothera
Renner, Planta 47: 223. 1956.-TYPE:
GERMANY. Brandenburg: Berlin-Rahns
dorf (cultivated from seeds collected by E. Baur in 1918). No authentic material
located; disposition based on description.
Oenothera
Z. Indukt. Abstammungs-Vererbungsl.
disjuncta Boedijn,
32: 361.
1924.-TYPE: U.S.A. Minnesota: Hennepin Co., North Town Junction (culti
vated from seeds collected
disposition
by H. de Vries
in Aug
1904). No material
located;
based on description.
Oenothera eriensis Gates, Canad. Field-Naturalist41: 26. 1927.-TYPE: CANADA.
Ontario: Essex Co., Colchester, Lake Erie (cultivated from seeds collected by R.
R. Gates on 24 Aug 1924), 1935, Gates 15.35 (neotype, here designated: BM!).
1997
OENOTHERA
[No original material
119
thus we have designated a specimen cultivated
in a later year as a neotype.]
59: 367. 1931.
Oenothera nobska Sturtevant, Z. Indukt. Abstammungs-Vererbungsl.
Oenothera oakesiana var. nobska (Sturtevant) Gates, Rhodora 59: 14. 1957.
located,
from the original material
Barnstable Co., on sandy beach N of Nobska
TYPE: U.S.A. Massachusetts:
from seeds collected
(cultivated
34-20
(neotype,
by A. H. Sturtevant
here designated:
cated. The Cleland
MO-3838392!).
(now housed atMO), which derives from
is here designated as a neotype.]
cultivated material
the original collection,
ostreae Sturtevant, Z. Indukt. Abstammungs-Vererbungsl.
Oenothera atrovirens var. ostreae (Sturtevant) Gates, Rhodora
Oenothera
TYPE: U.S.A. Massachusetts:
Barmstable Co., Falmouth,
by A. H. Sturtevant
tivated from seeds collected
(neotype, here designated: MO-3838391!).
Cleland
cultivated material
nal collection,
Oenothera
(now housed
is here designated
59: 367. 1931.
59: 13. 1957.
near Oyster Pond (cul
in 1927),
atMO),
which
34-23
1934, Cleland
[No authentic material
located. The
from the origi
derives
as a neotype.]
Gates & Catcheside
ammophiloides
1933.-TYPE:
Point
in 1926), 1934, Cleland
[No authentic material
lo
in Gates,
CANADA. Nova Scotia: Guysborough
J. Linn. Soc., Bot. 49: 180.
(cultivated at Regent's
Park,
England, from seeds collected by J. Rousseau and transmitted by Marie-Victorin
on 21 Aug 1930), Aug 1934, Gates s.n. (neotype, here designated: K!). [No orig
inal material
located. The neotype
is a specimen cultivated
from the original ma
terial in a later year.]
Oenothera
var. laurensis Gates, Philos. Trans., Ser. B, 226: 292.
CANADA. New Brunswick: Westmoreland Co., shore at Cape Tor
near Port Elgin (cultivated from seeds collected in 1932 by R. R. Gates),
ammophiloides
1936.-TYPE:
mentine
1934,Gates 45.34 (lectotype,here designated:BM!; isolectotype:GH!).
Oenothera
laevigata
1936.-TYPE:
var. rubripunctata
CANADA. Quebec:
shore of St. Lawrence
Philos.
Gates,
Bellechasse
[River] (cultivated
Trans.,
Co., mouth
at Regent's
Ser. B, 226:
317.
of River Boyer,
Park, England,
S
from seeds
collected by Marie-Victorin
and J. Rousseau on 31 Sep 1932 or by R. R. Gates
on 2 Oct 1932), 1934, Gates 53.34 (lectotype, here designated: BM!).
Oenothera
leucophylla Gates, Philos. Trans., Ser. B, 226: 301. 1936.-TYPE:
CANADA. Quebec: Bellechasse
land, from seeds collected
(lectotype,
Oenothera
eriensis
here designated: BM!;
niagarensis
Philos.
Gates,
var. niagarensis
New York: Niagara
(cultivated at Regent's
on 30 Sep 1932),
Trans.,
Ser. B, 226: 326.
(Gates) Gates, Rhodora
76.35
isolectotype:
1936. Oenothera
59: 13. 1957.-TYPE:
(cultivated
by R. R. Gates on 28 Aug
type, here designated: BM!;
Park, Eng
1935, Gates
isolectotype: GH!).
Co., Niagara Gorge
from seeds collected
Oenothera
Co., St. Vallier
by R; R. Gates
at Regent's
U.S.A.
Park, England,
1932), 1935, Gates 49.35
(lecto
GH!).
repandodentata
Gates, Philos. Trans., Ser. B, 226: 328. 1936. Oenothera
var. repandodentata
(Gates) Gates, Rhodora 59: 13. 1957.-TYPE:
CANADA. Ontario: Essex Co., Colchester (cultivated at Regent's Park, England,
eriensis
from seeds collected
type, here designated:
Oenothera
by R. R. Gates
BM!;
on 9 Oct
isolectotypes:
1932),
1935, Gates
97.35
(lecto
GH! 2 sheets).
deflexa var. bracteata Gates, Philos. Trans., Ser. B, 226: 335. 1936.
TYPE: CANADA. Ontario: Essex Co., Sandwich, vicinity of Windsor
(cultivated
SYSTEMATICBOTANYMONOGRAPHS
120
VOLUME 50
Park, England, from seeds collected by R. R. Gates on 9 Oct 1932),
96.35 (lectotype, here designated: BM!; isolectotype: GH!).
64: 142. 1950.-TYPE:
Oenothera
perangusta
Gates, Canad. Field-Naturalist
CANADA. Ontario: sandy beach of Lake Huron at Stokes Bay, Bruce Peninsula,
at Regent's
1935, Gates
11 Jul 1934, Krotkov 9252 (holotype: TRT!; isotypes: GH! US!).
var. rubricalyx Gates, Canad. Field-Naturalist
Oenothera
perangusta
64: 143.
1950.-TYPE:
CANADA. Ontario: Thunder Bay Dist., Canadian Pacific Railway
station at Jackfish Station, Lake Superior, 7 Jul 1933, Pease & Bean 23526 (holo
type:GH!).
Oenothera
magdalena
Gates,
Canad.
Field-Naturalist
65:
196.
1951.-TYPE:
Islands, Coffin Island, dry crevices or talus of East
Cape, 17 Aug 1912, Fernald, Long & St. John 7834, pro parte (lectotype, here
designated: GH!). Gates based his description on plants grown from seeds col
lected by M. Gauvreau in 1934 on the Magdalen
Islands. We have not located
CANADA. Quebec: Magdalen
any vouchers from these cultivated plants; however, Gates also presented in a
table morphology
on eight other specimens in GH collected by Fernald, Long,
and St. John in 1912 on theMagdalen
Islands with lengths of the bud, free sepal
tips, mid leaf, and a few miscellaneous
additional comments on hairs, petioles
and leaf spots given. We have located six of these specimens at GH with assis
tance of D. Boufford. The material represents two species: 0. oakesiana
(4
sheets [Fernald et al. 7833 (1 sheet) and Fernald et al. 7834 (3 sheets)], plus one
plant on the fifth sheet [Fernald et al. 7834a,
amixed collection of 0. oakesiana
and 0. biennis]); and 0. biennis [showing some evidence of hybridization with
0. oakesiana]
(the sixth sheet, another sheet of Fernald et al. 7834a, plus the
other plant on the first sheet of Fernald et al. 7834a). All of these collections can
be associated with measurements
in the table; we have selected as the lectotype
the specimen (0. oakesiana) that most closely fits the original description.
Oenothera
var. angustifolia Gates, Monogr. Biol. 7: 74: 1958.
ammophiloides
TYPE: CANADA. Quebec: Montmorency
Co., St. Joachim (cultivated from seeds
presumably collected by R. R. Gates); fig. 17 (culture 80.39), p. 75 (lectotype,
here designated).
Erect to procumbent
(taller in cultivation),
[No authentic material
seen.]
biennial herb with a taproot, forming a rosette; stems 1-6 dm tall
green or flushed with
red in the lower parts or throughout, un
branched or bushy-branched
from the base with side branches arising obliquely or arcu
ately from the rosette, either a) densely silky strigillose and with scattered long appressed
hairs, b) as in (a) but also with subappressed to erect pustulate hairs in the apical parts, or
in lower parts, in the region of the inflorescence only glandu
c) densely silky-strigillose
lar-puberulent and with pustulate hairs. Leaves grayish green to dull green and silky,
densely strigillose on both surfaces and margins, the apical bracts sometimes also glan
dular-puberulent. Rosette leaves 8-30 cm long, 0.5-3 cm wide, very narrowly oblanceo
late to narrowly oblanceolate, margin remotely dentate, the teeth sometimes blunt, at base
sometimes also sinuate-dentate, apex acute to narrowly acute, base gradually narrowed to
the petiole. Cauline leaves 3.5-20 cm long, 0.5-2.7 cm wide, very narrowly oblanceolate
or very narrowly elliptic
teeth blunt, to subentire,
to narrowly
elliptic, margin remotely dentate, sometimes the
the base sometimes sinuate-dentate,
apex acute to narrowly
acute, base narrowly cuneate to attenuate, short-petiolate to sessile. Bracts 2-10 cm long,
0.3-2 cm wide, narrowly lanceolate to narrowly ovate or narrowly elliptic, margin bluntly
OENOTHERA
1997
121
dentate to subentire, apex acute to narrowly acute, base acute to narrowly cuneate, sessile,
usually longer than the capsules they subtend. Inflorescence unbranched, the apical part
usually recurved with the tip ascending, rarely suberect. Floral tube 1.5-4 cm long, 1-1.5
mm in diameter, yellowish green, often flushed with red and/or flecked with red to dark
red spots, silky-strigillose with long and short hairs, also with pustulate hairs, and glan
dular-puberulent. Mature buds 0.8-1.5 cm long, 3-5 mm in diameter, narrowly oblong to
broadly oblong, lanceoloid or narrowly ovoid to ovoid. Sepals 0.9-1.7 cm long, 2.5-4 mm
wide, green to yellow, flushed with red and dark red-flecked or red-striped, pubescence
like floral tube; free sepal tips 2.5-4 mm long, subterminal in bud, erect to spreading,
strigillose. Petals 0.7-2 cm long, 0.8-2 cm wide, very broadly obovate, retuse, yellow to
pale yellow. Filaments 6-15 mm long; anthers 3-7 mm long; pollen ca. 50% fertile. Ovary
cm long, 1.4-2 mm in diameter, densely silky-strigillose with long and short hairs,
villous, glandular-puberulent
and densely to sparsely pustulate-pubescent.
Style 2-4.5 cm
long, the exserted part 0.3-0.8 cm long; stigma surrounded by the anthers, which shed
pollen directly onto the lobes at anthesis, the lobes 3-5 mm long. Capsules 1.54 cm long,
0.7-1.2
in diameter, narrowly lanceoloid to lanceoloid, attenuate to the apex, when fresh
dark to dull green, sometimes red-striped or red-flecked, when dry usually rusty brown;
free tips of the valves not more than 0.5 mm long, obtuse to truncate. Seeds 1.1-1.2 mm
number: n = 7
long, 0.8-1.1 mm in diameter, dark brown to almost black. Chromosome
4-8 mm
(014; 012 and 1II; 010 and 04 [Cleland 1972, p. 339]; based on 5 individuals
localities). Self-compatible,
usually autogamous, PTH. Fig. 38.
from 5
Phenology.Flowering in July throughSeptember,sometimesOctober.
Distribution
(Figs. 14, 22, 37). Occurring
in sandy coastal meadows
and dunes, or on
gravelly or rocky sites along rivers, also in disturbed sites such as roadsides. Ranging in
Canada from southeastern Manitoba
through southern Ontario and Quebec to Newfound
land, south into the United States from northern and eastern Minnesota,
southeast to
northern Illinois and Indiana east to the Atlantic coast, and thence south toNorth Carolina;
naturalized in Europe and sporadically in Asia.
widely
Oenothera oakesiana is a PTH species with an AC genomic constitution and plas
tome IV (Stubbe 1959, 1963, 1964). It is almost exclusively autogamous. This species has
only recently been recognized
under the name 0. oakesiana (Raven et al. 1979). In the ex
literature, the plants referred here to 0. oakesiana were designated as Parvi
flora-Il by Cleland (1972). InNorth America this species most commonly has been treated
as a variety of 0. parviflora following Munz (1965). Naturalized populations in Europe
have been known under the name 0. muricata (misapplied, the type is a specimen of 0.
biennis) during the 19th century and the first half of the 20th century. More recently, plants
perimental
from inland parts of Europe, which
a name that despite its wide
cola,
have a similar phenotype,
have been called 0. syrti
use has never been validly
published,
whereas
the
coastal plants have been referred to 0. ammophila. A number of segregate species have
been described for AC genome plastome IV plants, all of which we refer to 0. oakesiana.
Two names, 0. ostreae and 0. stenopetala, were based on plants with the cruciate petal
character. This character represents a mutation in which the petals are modified into nar
row ligulate structures, typically with irregular margins, and often greenish yellow rather
than normal yellow. It has been studied in some detail; the work is summarized by Cle
land (1972).
In 1979, Raven et al. argued that the strains traditionally genetically recognized as
Parviflora-II
morphological
ought to be recognized
distinctness
as a distinct
and especially
species, 0. oakesiana, because of their
their unique genomic constitution (AC), which
SYSTEMATICBOTANYMONOGRAPHS
122
VOLUME 50
mm
b
dC
FIG. 38. Oenothera
cauline
leaf d. Capsules.
oakesiana
(Hall 3436,
e. Inflorescence
cult. DUSS-88-2017).
pubescence.
a. Inflorescence.
b. Rosette
A4
leaf. c. Mid
1997
OENOTHERA
123
is different from Parviflora-I (BC, = 0. parviflora). These two species share the C genome
as their f (pollen) complex, which contributes the sigmoid inflorescence apex, the subter
minal free sepal tips, and the narrow leaves (Table 2), but they have different aX (egg)
complexes and thus different origins. These are the only two species that have plastome
IV, presumably indicating a common ancestry. The more detailed studies presented here
support the continuation of the taxonomic philosophy proposed by Raven et al. (1979),
which is to treat as different species PTH entities that either arose from different sources
(e.g., 0. wolfi, from 0. elata subsp. hookeri, is treated as distinct from 0. villosa, from
0. elata subsp. hirsutissima) or that combine genomes from different sources, as is the
case in the separation of 0. nutans (BB) from 0. biennis (AB or BA). The X complex of
0. oakesiana contributes A genome characteristics, such as the dense pubescence and in
part the narrow leaves (Table 2). Unique features of 0. oakesiana
dry capsule color and the silkiness of the pubescence.
include the rusty brown
Individuals of 0. oakesiana nearly always form a 014 during meiotic metaphase
I,
but some individuals of the ammophila (coastal) phenotype have (0 12 and I1, and one of
Cleland's strains (1972, p. 339) from Michigan
(Manistique) had a 010 and 04 configu
ration. The Cleland strain may represent an F1 hybrid between 0. oakesiana and 0. par
vifiora.
There are basically two forms of 0. oakesiana: a coastal form, characterized by a
short habit, conspicuous pustulate pubescence, and a strongly sigmoid inflorescence, and
an inland form characterized by a larger and robust habit, less conspicuous pustulate pu
bescence, and a slightly recurved inflorescence apex weakly ascending at the tip. As men
tioned above, these two forms have long been recognized in Europe as 0. ammophila and
0. syrticola, respectively. A similar pattern with more robust plants occurring at inland
sites also was observed in our studies throughout the indigenous North American range of
the species. Bartlett (1914) and Gates (1936) both published names for a number of addi
tional minor variants; all have AC genome combination with plastome IV, and thus are not
formally recognized
There
here.
is conspicuous
but local intergradation between 0. parviflora and 0. oake
siana. Despite extensive overlap in their ranges and the fact that the hybrids are fully vi
able, the area of intergradation is primarily only in the Great Lakes region. For example,
inMichigan
and Ontario many intermediates have the silky pubescence of 0. oakesiana,
but leaves and capsules more characteristic of 0. parvifiora. The intermediates perpetuate
themselves faithfully because the plants are PTH. Their predominance at inland localities
suggests that certain of the hybrids have been perpetuated, because the new phenotype
was somehow adaptive in northern inland environments. Throughout much of their range
0. oakesiana and 0. parviflora grow in adjacent, but different, habitats, without visible
signs of intergradation. However,
it should be noted that we have not studied this phe
nomena extensively
cult to detect many
in the field, but primarily from herbarium specimens. It is very diffi
features of subsect. Oenothera species from pressed specimens, and
therefore, we may have somewhat underestimated the extent of intergradation between 0.
parviflora and 0. oakesiana. We have dealt with the extensive herbarium material by an
notating many of the putative intermediates as the species which they most closely re
semble.
Oenothera oakesiana (AC-IV) also hybridizes with 0. biennis (AB-IT), both within
the overlap of their indigenous ranges and in Europe, where hybridization has been more
thoroughly studied. In both directions the hybrids are viable (Stubbe 1959; Stubbe in Cle
land 1972). When 0. oakesiana is the female parent the hybrids are an AB-IV/II combi
SYSTEMATICBOTANYMONOGRAPHS
124
nation, while
are AC-II/IV.
the reciprocals
In Europe, a number of these hybrids have been
as species,
sometimes
formally described,
also hybrids section). Hybrids with
at other times as named hybrids
this genomic
combination
ther parent, and the flowers are intermediate
in size. In the context of mixed
of both parents, the hybrids usually can be detected
true of herbarium specimens,
most
even when
populations
the reverse is
relatively easily. Quite
a full suite of collections
we have annotated most
Therefore,
closely
(Table 4; see
are phenotypic
essentially
of the parents. In nature these hybrids have a more vigorous habit than ei
reconstructions
population.
VOLUME 50
from amixed
are made
of these hybrids as the parent that they
resemble.
13. Oenothera
Methodus
Moench,
Mus.
L., Syst.
parvifiora
Hist. Nat.
nat., ed. 10. 998.
1802. Onagra
suppl. 287.
355.
4(4):
1836
1759. Onagra
(L.)
parviflora
Spach, Nouv. Ann.
chrysantha
non Onagra
["1835"],
chrysantha Michaux
var. parvifiora (L.) Spach, Nouv. Ann. Mus. Hist.
Nat., Paris 4(4): 355. 1836 ["1835"]. Oenothera biennis var. parviflora (L.) Tor
(1803). Onagra
chrysantha
rey & A. Gray, Fl. N. Amer.
(L.) H.
parvifiora
Oenothera
1: 492. 1840. O[e]nothera
Bull.
Leveille,
muricata
Int. Geogr.
Acad.
subsp. parviflora
communis
Bot.
Chromos.
(L.) Tischler,
race biennis
f.
19: 330.
1909.
Gefasspfl.
Mit
teleur. 58. 1950. Oenothera
parviflora var. oakesiana f. parviflora (L.) Scoggan,
4: 1143. 1979.-TYPE:
[possibly from Europe] (holotype: LINN
Fl. Canada
484.2!).
Oenothera
angustifolia
here designated:
var. cruciata
355.
s.n. in Herb. Sloane,
U.S.A.
ex G. Don, Gen. hist. 2: 686. 1832. Onagra
1836 ["1835"]. Oenothera
G. Don) Britton, Mem.
biennis var. cruciata
1: 492. 1840. Onagra
(Nuttall ex G. Don) Torrey
biennis var. cruciata
Torrey Bot. Club 5: 223. 1894. Onagra
19: 330.
1909.-TYPE:
said to be from North America
cruciata,
from
onward,
not located. Cultivated
the Botanical
(1827, E!); Cannonmills
Gardens,
Cambridge
Lodge, Edinburgh
(1825, CGE!);
from Cambridge,
cruciata Nuttall"
original
Oenothera
angustissima
Brown
Edinburgh
parvifiora.
The collection
by
is labelled "Oenothera
This collection may be the
var. varia de Vries, Bull. Torrey Bot. Club 30: 76. 1903.-TYPE:
angustissima
gustissima
New
the name
from 1824
source of the seeds sent to England.
cruciata
No authentic material
Oenothera
Int. Geogr.
(1828, E!); and Harvard University
(BM! PH!)
Massachusetts
and is also Oenothera
(Nuttall
communis
plants bearing
and in cultivation
(1856 & 1875, GH!) are all the cruciate form of 0. parviflora.
T. Nuttall
(Nuttall ex
cruciata
ex G. Don) Small, Bull. Torrey Bot. Club 23: 169. 1896. O[ejnothera
race biennis var. cruciata (Nuttall ex G. Don) H. Leveille, Bull. Acad.
Oenothera
chrysantha
(Nuttall ex G. Don) Spach, Nouv. Ann. Mus. Hist. Nat., Paris 4(4):
& A. Gray, Fl. N. Amer.
Bot.
Virginia.
vol. 295: 69 (lectotype,
BM-SL!).
cruciata Nuttall
Oenothera
Gard. dict., ed. 8: 2. 1768.-TYPE:
Miller,
in Europe; Miller
Seeds cultivated
seen; no figure provided; disposition
Gates, Rhodora
(Gates) Wiegand,
(Gates) Munz,
York: Tompkins
Co.,
in 1909), 1913, Gates
based on description.
15: 46. 1913. Oenothera
parviflora
var. an
Rhodora 26: 3. 1924. Oenothera parviflora subsp.
N. Amer. Fl., ser. 2, 5: 123. 1965.-TYPE:
U.S.A.
Ithaca (cultivated
13.35
from seeds collected
(holotype: BM!).
by H. B.
1997
OENOTHERA
125
Oenothera
U.S.A. West
l[a]evigata Bartlett, Cybele Columb. 1: 47. 1914.-TYPE:
Virginia: Greenbrier Co., White Sulphur Springs (cultivated from rosettes col
lected by Bartlett in 1912), Bartlett 3504 (holotype: MICH! 2 sheets).
Oenothera scitula Bartlett, Cybele Columb. 1: 45. 1914. Oenothera
laevigata var.
scitula (Bartlett) Gates, Rhodora 59: 13. 1957.-TYPE:
U.S.A. West Virginia:
Greenbrier Co., White Sulphur Springs (cultivated from rosettes collected by
Bartlett in 1912), Bartlett 3559 (lectotype, here designated: MICH! 2 sheets).
atrovirens Shull & Bartlett in Bartlett, Amer. J. Bot. 1: 239. 1914.
Oenothera
Oenothera
Bot.
muricata
5: 257.
(Sandy Hill)
subsp. atrovirens
(Shull & Bartlett) Love & Love, Opera
York: Washington
Co., Hudson Falls
form seeds from D. T.MacDougal),
1913?, Bartlett 3500
1961.-TYPE:
(cultivated
U.S.A.
New
(holotype:US-693736!, US-693737! 2 sheets).
venosa Shull & Bartlett in Bartlett, Amer. J. Bot. 1: 241. 1914.-TYPE:
New York: Washington Co., Hudson Falls (Sandy Hill) (cultivated from
seeds from D. T. MacDougal),
1913?, Bartlett 3501 (holotype: US-393738-40!
3 sheets).
Oenothera cleistantha Shull & Bartlett in Bartlett, Rhodora 17: 43. 1915.-TYPE:
Oenothera
U.S.A.
U.S.A.
New York: Suffolk Co., Huntington
(cultivated from seeds collected by
G. H. Shull), Bartlett 3646 (lectotype, here designated: MICH!;
isolectotypes:
BH! UC!).
Oenothera
robinsonii Bartlett, Rhodora 17: 42. 1915.-TYPE:
U.S.A. New Hamp
shire: Cheshire Co., Jaffrey (cultivated from seeds collected by B. L. Robinson),
Sep 1913, Bartlett 3505 (lectotype, here designated: MICH!).
Oenothera
novae-scotiae Gates, pre-print of Trans. Nova Scotia Lit. Soc. 14: 142.
1916 Uournal published in 1918].-TYPE:
CANADA. Nova Scotia: North Mtn Rd
above reservoir near Middleton
(cultivated from seeds collected by R. R. Gates
in 1914), 12 Jul 1916, Gates s.n. (holotype: UC-193440!).
Oenothera
pachycarpa Renner ex Rudloff, Gartenbauwissenschaft
3: 499. 1930.
Oenothera parviflora subsp. pachycarpa
(Renner ex Rudloff) Janchen, Phyton
(Horn) 3: 7. 1951. Oenothera muricata subsp. pachycarpa
(Renner ex Rudloff)
Love & Love, Opera Bot. 5: 257. 1961.-TYPE:
GERMANY. Sachsen: sponta
in Botanical Garden of Jena; fig. 2, p. 500 (lectotype, here designated).
[No authentic material seen.]
Oenothera
angustissima
var. quebecensis Gates, Philos. Trans., Ser. B, 226: 324.
1936.-TYPE:
CANADA. Quebec: Montmorency
Co., north shore of the St.
Lawrence
[River] at Cap Tourmente
(cultivated from seeds collected by F.
neous
Michel
& M. L. Chollet),
1935, Gates 44.35
(lectotype, here designated:
BM! 2
sheets; isolectotype:GH!).
Oenothera
biformiflora Gates, Philos. Trans., Ser. B, 226: 303. 1936.-TYPE:
CANADA. Quebec: Bellechasse Co., St. Vallier (cultivated at Regent's Park, Eng
land, from seeds collected on 1Oct 1932 by Marie-Victorin
and J. Rousseau),
1934, Gates 61.34 (lectotype, here designated: K!).
Oenothera biformiflora var. cruciata Gates, Philos. Trans., Ser. B, 226: 305. 1936.
TYPE: CANADA. Quebec: Bellechasse
Co., St. Vallier (cultivated at Regent's
Park, England, from seeds collected on 1 Oct 1932 by Marie-Victorin
and J.
Rousseau),
1934, Gates 60.34 (lectotype, here designated: K! 2 sheets).
comosa Gates, Philos. Trans., Ser. B, 226: 262. 1936.-TYPE:
CANADA.
Nova Scotia: Wilmot, ca. 2 mi from Middleton
(cultivated at Regent's Park, Eng
Oenothera
126
SYSTEMATICBOTANYMONOGRAPHS
land, from seeds collected
VOLUME 50
on 6 Sep 1932),
by R. R. Gates
1934, Gates
23.34
(lectotype,here designated:BM!; isolectotypes:GH! 2 sheets,K!).
Oenothera
flecticaulis Gates, Philos. Trans., Ser. B, 226: 269. 1936. Oenothera am
mophiloides
var. flecticaulis
(Gates) Gates, Rhodora 59: 11. 1957.-TYPE:
CANADA. Nova Scotia: Lunenburg Co., Beach near mouth of Lahave River (cul
tivated at Regent's
1932);
from seeds collected
Park, England,
fig. 17 of culture 103.34
terial located, but cultures 78.33,
Nova
102.35 were
103.34,
in Oct
[No authentic ma
cited. Disposition
based
of cultivated plant 103.34.]
on photograph
Oenothera
by Mrs. W. Bell
here designated).
(lectotype,
hazelae Gates, Philos. Trans., Ser. B, 226: 272. 1936.-TYPE: CANADA.
Co., near Lockeport (cultivated at Regent's Park, Eng
Scotia: Shelbume
by Mrs. W. Bell
land, from seeds collected
107.34
Oenothera
var. parviflora
hazelae
TYPE: CANADA. Nova
Park, England,
Gates
in Oct 1932),
31 Aug
1934, Gates
(holotype: K! 2 sheets).
110.34
Philos. Trans., Ser. B, 226: 275. 1936.
Gates,
Co., Port Mouton
Scotia: Queen's
by Mrs. W. Bell
from seeds collected
here designated:
(lectotype,
(cultivated
at Regent's
in Oct 1932), 31 Aug
1934,
K! 2 sheets).
Philos.
Trans., Ser. B, 226: 266. 1936. Oenothera
(Gates) Gates, Rhodora 59: 11. 1957.-TYPE:
CANADA. Nova Scotia: Digby Co., Bear River (cultivated at Regent's Park, Eng
Oenothera
intermnedia Gates,
novae-scotiae
var. intermedia
land, from seeds collected
50.35
determined
by R. R. Gates
here designated).
(lectotype,
on 22 Sep 1932);
[No authentic material
(fig. 15), but cultures
from photograph
fig. 15 of culture
located. Disposition
3.33,
were
50.35
21.34,
cited.]
Oenothera
var. similis Gates,
laevigata
Trans., Ser. B, 226: 312. 1936.
Mouth
of River Boyer, S shore of St.
Co.,
Philos.
TYPE: CANADA. Quebec: Bellechasse
Lawrence
[River] (cultivated at Regent's
Marie-Victorin
1934, Gates 54.34
Oenothera
var. distantifolia
CANADA. Nova
gent's Park, England,
Gates 29.34
Trans.,
Ser. B, 226: 296.
at Re
by R. R. Gates
(cultivated at Regent's
on 2 Oct 1932),
am
1936. Oenothera
59: 12. 1957.-TYPE:
(Gates) Gates, Rhodora
Co., Bic, by old wharf
from seeds collected
(cultivated
here designated: K! 3 sheets).
Philos.
var. parva
Quebec: Rimouski
Philos. Trans., Ser. B, 226: 260.
Gates,
Scotia: Hants Co., Newport
from seeds collected by R. R. Gates on 27 Sep 1932), 1934,
(lectotype,
parva Gates,
mophiloides
here designated: BM!).
(lectotype,
novae-scotiae
1936.-TYPE:
Oenothera
Park, England, from seeds collected by
on 31 Sep 1935 or R. R. Gates on 2 Oct 1932),
and J. Rousseau
CANADA.
Park, England,
1934, Gates 47.34
(lecto
type, here designated: BM!).
Oenothera
U.S.A.
rubricapitata Gates, Philos. Trans., Ser. B, 226: 343. 1936.-TYPE:
North Dakota: Cass Co., Kindred, wooded area by a pond (cultivated at
Regent's Park, England, from seeds collected by R. R. Gates
1935, Gates 100.35 (lectotype, here designated: BM!).
Oenothera
hazelae
subterminalis
Gates,
var. subterminalis
CANADA. Nova
vated at Regent's
1932), 28 Aug
Philos. Trans., Ser. B, 226: 278.
(Gates) Gates,
Scotia: Colchester
Park, England,
1934, Gates 34.34
Rhodora
Co., Higgins
Brook
from seeds collected
(lectotype,
59:
on 15 Oct 1932),
1936. Oenothera
11.
1957.-TYPE:
near Wentworth
(culti
by R. R. Gates on 28 Sep
here designated:
K! 3 sheets).
OENOTHERA
1997
127
deflexa Gates, Philos. Trans., Ser. B, 226: 332. 1936.-TYPE:
CANADA.
Ontario: Essex Co., Ojibway, vicinity of Windsor
(cultivated at Regent's Park,
England, from seeds collected by R. R. Gates on 9 Oct 1932), 31 Aug 1934,
Oenothera
Gates 89.34 (lectotype, here designated: K! 3 sheets).
Oenothera silesiaca Renner, Ber. Deutsch. Bot. Ges. 60: 455. 1942. Oenothera muri
cata subsp. silesiaca (Renner) Tischler, Chromos. Gefasspfl. Mitteleur. 58. 1950.
Oenothera parviflora subsp. silesiaca (Renner) Janchen, Phyton (Horn) 3: 7.
1951.-TYPE:
POLAND. Wroclaw:
at railway station of Krzystkowice
near
Nowogrod
(cultivated from seeds collected by 0. Renner in 1937), 1967, Ross
mann 286/66 (neotype, here designated: M!). [Renner never made any specimens
of his strain, nor did he designate a type. The material used for the neotype is de
scendent from Renner's
Oenothera
cruciata
CANADA. Nova
original material.]
var. sabulonensis
51: 67. 1949.-TYPE:
Fernald, Rhodora
Scotia: Sand dunes, Sable Island, 18 Aug 1913, St. John 1283
(holotype:GH!).
Oenothera
apicaborta
Gates, Canad. Field-Naturalist
65: 194. 1951.-TYPE:
CANADA. Quebec: Champlain Co., Les Piles between railway and St. Maurice,
12 Aug 1936, Marie-Victorin
& Rolland-Germain
51 (lectotype, here desig
nated: MT!;
isolectotypes:
51 of Marie-Victorin
CU! DAO!
& Rolland-Germain,
FSU! TRT! US!).
from which
[Gates referred to no.
he also took seeds for his
cultures, but did not designate a type. The collection atMT, where the first set of
their collections is housed, is here designated as the lectotype.]
Oenothera rubricuspis Renner [Ber. Deutsch. Bot. Ges. 63: 131. 1950 (without Latin
ex Rostan'ski, Fragm. Florist. Geobot.
diagnosis)]
11: 512. 1965. Oenothera
muricata
subsp. rubricuspis (Renner ex Rostan'ski) Weihe
in Garcke, Ill. Fl.
Deutschland,
Neu-Isenburg
ed. 23, 979. 1972.-TYPE:
GERMANY. Hessen: at railroad between
and Luisa near Frankfurt (cultivated from seeds sent by F. Schotz
[Munich] from Renner's
strain), 15 Jul 1964, Rostan'ski 20/63
(holotype: WRSL!
5 sheets).
turoviensis Rostaniski, Fragm. Florist. Geobot.
Oenothera
11: 514. 1965.-TYPE:
POLAND.Wroclaw: Turoszow, 12 Sep 1963, Rostan'ski s.n. (holotype: WRSL! 4
sheets; isotype: KTU!).
Oenothera
lipsiensis Rostan'ski & Gutte, Ber. Arbeitsgem.
1971.-TYPE:
GERMANY. Sachsen: Leipzig-M6ckern,
Sachs. Bot., n.s., 9: 69.
"neuer Miillberg," 26 Jul
1965, Gutte s.n. (holotype: LZ-2410!).
Erect biennial herb with a taproot, forming a rosette; stems 3-15 dm tall, unbranched
branched from the base or only from the apical half of main stem, green or red in
the lower part or throughout, sparsely strigillose, glandular-puberulent, and with pustulate
or mostly
hairs, sometimes strigillose only in the lower part, or with pustulate hairs only in apical parts
and other long spreading hairs in the inflorescence, other times glabrous in the region of the
inflorescence.
Leaves
usually
bright
green,
veins
white
or red,
sparsely
strigillose on both surfaces and margins, the upper surface sometimes subglabrous, bracts
also glandular-puberulent and with long spreading hairs or with scattered appressed hairs
near apex. Rosette leaves 10-30 cm long, 1-4 cm wide, very narrowly to narrowly oblance
olate or narrowly elliptic, margin regularly dentate to remotely denticulate, apex acute, base
attenuate to the petiole. Cauline leaves 4-18 cm long, 1-3 cm wide, lanceolate to narrowly
ovate, very narrowly to narrowly elliptic, or narrowly oblong, margin usually regularly den
SYSTEMATICBOTANYMONOGRAPHS
128
VOLUME 50
tate, apex acute to long-acute, base attenuate to acute, sessile or short-petiolate. Bracts 2-8
cm wide, sometimes pale green, narrowly lanceolate to narrowly ovate,
cm long, 0.3-2.5
regularly to remotely dentate or subentire, apex acute to long-acute, base acute to
narrowly cuneate. Inflorescence erect or curved, unbranched or often with secondary spikes
below the main one. Floral tube 2.2-4 cm long, ca. 1mm in diameter, yellowish, glabrous
and sparsely villous. Mature buds 0.6-1.5 cm long, 3-5
to densely glandular-puberulent,
margin
in diameter, narrowly oblong to lanceoloid. Sepals 0.7-1.7 cm long, 2.5-4 mm wide,
green to yellowish green or flushed with red or dark red, sometimes only red-flecked, pu
bescence like that of floral tube; free sepal tips 0.5-5 mm long, distinctly to indistinctly sub
mm
terminal, ca. 0.5-1 mm apart in bud, strigillose or with spreading hairs. Petals 0.8-1.5 (-2)
cm long, 0.9-2 cm wide, very broadly obovate, retuse to emarginate, yellow to pale yellow.
Filaments 7-13 mm long; anthers 3.5-6 mm long; pollen ca. 50% fertile. Ovary 0.9-1.3 cm
long, 1.5-1.8 mm
in diameter, strigillose, sparsely villous and glandular-puberulent, some
times glandular-puberulent and either strigillose, villous, sparsely pustulate-pubescent,
or
sparsely appressed pubescent near the apex, occasionally glabrous. Style 2.5-5 cm long, the
exserted part 0.1-1 cm long; stigma below or surrounded by the anthers, which shed pollen
directly onto the lobes at anthesis, the lobes 2.5-6 mm long. Capsules 2-4 cm long, 3.5-5
cm in diameter, narrowly lanceoloid to lanceoloid, attenuate toward the apex, dark green
fresh, often becoming black when dry, pubescence like that of ovary but less dense,
often becoming glabrous. Seeds 1.1-1.8 mm long, 0.5-1 mm in diameter, brown to dark
brown. Chromosome number: n = 7 (014; based on 15 individuals from 14 localities). Self
when
compatible,
usually autogamous,
Phenology.
Flowering
PTH. Fig. 39.
from July to September,
and occasionally
into October.
Distribution
(Figs. 13, 14, 26, 33, 40). Occurring usually in open or disturbed, sandy
or gravelly sites, such as along roadsides, fallow fields, clearings, river banks or along
other water courses, salt marshes, and coastal meadows,
in eastern North America, from
southern Ontario and Minnesota
east to Newfoundland,
and south through Iowa, Indiana
and eastern Tennessee, to the Atlantic coast south to North Carolina, with a few scattered
collections
from Illinois and Missouri;
now widely naturalized
and South Africa.
Oenothera parviflora is a PTH species with a BC genomic
IV (Stubbe 1959, 1963, 1964). It is nearly always autogamous,
during meiosis. This species is strongly heterogamous. The a
in northeastern China, Eu
rope, Japan, New Zealand,
or rarely the C genome, whereas
the i complex
is always
constitution and plastome
and it always forms a 014
complex is the B genome
the C genome. The relatively
sparse pubescence characteristic of this species is a feature of both the B and C genomes.
the most distinctive features of 0. parviflora are those conferred by genes lo
cated on the C genome: relatively narrow leaves, curved inflorescence apex, and subter
However,
minal free sepal tips (Table 2), the latter two may
land 1972, p. 249).
result from closely
linked genes
(Cle
This specieswas subdividedinto two subspecies,0. parviflora subsp.parviflora and
0. parviflora subsp.angustissima,and the formersubspecieswas furthersubdivided into
two varieties (Munz 1965).We have elevated 0. parviflora subsp.parviflora var. oake
siana to the rank of species, because of its different genomic
constitution
and origin. Thus,
theplants referredhere toOenotheraparviflora corresponddirectly to thosedesignatedas
Parviflora-I
in the experimental
We also do not subdivide
literature (Cleland 1972).
the here more narrowly delimited 0. parviflora,
because the
features used by Munz for the recognition of two subspecies, primarily pubescence, petal
length, and capsule length, vary relatively continually from one extreme to the other. Also,
1997
OENOTHERA
129
mm
A
L.'A
d d30
ej~~~~~~~~~~~V
b
e
FIG. 39. Oenothera
DUSS-88-W867
cence.
c
parviflora
[f]). a. Inflorescence.
(Munz 17517,
b. Rosette
cult. DUSS-86/88-1018
leaf. c. Mid-cauline
[a-e]; Stoutamire
leaf. d. Capsule.
e-f.
s.n. in 1952, cult.
Inflorescence
pubes
SYSTEMATICBOTANYMONOGRAPHS
130
0
0
h
*
,~~~~
0*
00~~~~
0
_X 4
0
'
027~~~~~~~~~~~~
@00 ~
0
00
0.
0
0
00
,g/ 0
~~~~~~~~0
:.:.
~~~~~0
0
~
?<
0
000
0
0. parviflora
\
\ <S
0- e
VOLUME 50
t \
~~+30?N
80?W
500 km
FIG. 40. Indigenousdistributionof Qenothera parvifltora.
thedensity of the varioushair types and size of thepetals and capsules all appear to vary
independentlyof one another.Plants from the southernpart of the range are less pubes
cent than those in thenorth,and generally correspondtoMunz's concept of 0. parviflora
subsp. angustissima.The increase in pubescence density in the northmay be caused, at
least inpart,by hybridizationwith 0. biennis and0. oakesiana and incorporationof their
A-genome pubescence characters.
A number of segregate species have been described for BC genome plastome IV
plants, all of which we recognize here as Qenothera parvfiJora.Some of them,0. atro
virens, 0.cruciata,
0. cleistantha,0. robinsonii,and 0. venosa, were based on plants
with thecruciatepetal character.This characterrepresentsamutation inwhich each petal
1997
OENOTHERA
131
is narrowed into a ligulate structure, typically with irregular margins, and often greenish
yellow rather than normal yellow. It has been studied in some detail, and the work is sum
marized by Cleland (1972). The cruciate character segregates genetically, and therefore
we do not recognize it taxonomically.
Most of the other names included in 0. parviflora were published by Gates. His
species were based on variations in leaf coloration, shape, and texture, color and pubes
cence of the stem, and degree of bowing and density of the inflorescence. Although they
represent true-breeding variants, it is not useful to recognize them formally. The differ
ences among them are trivial, and hundreds more variants separated by similar degrees of
difference
could be described
from within
the PTH populations
here referred to 0. parv
iflora.
Oenothera parviflora
(BC-IV) grows sympatrically with a number of the other
species of subsect. Oenothera. Hybrids have been documented with 0. argillicola (local),
0. biennis, 0. nutans, 0. oakesiana, and 0. villosa subsp. villosa (Table 3). Hybridization
involving 0. parviflora occurs within the indigenous range in North America, especially
with 0. oakesiana (discussed under that species, no. 12), and also in Europe. Most of the
characteristics that would allow hybrids to be recognized are not evident in dried speci
mens, and consequently we have doubtless often annotated such hybrids with the name of
the parent they most closely resemble.
Hybrids between 0. oakesiana and 0. parviflora are possible in both directions and
essentially conform to the maternal phenotype. Because
the chromosomal
formula of
these two species differ, the hybrids will not form (0 14 chromosomes
at meiotic
metaphase
I, but rather form small to large rings along with one to several bivalents. It
should be expected that following self-pollination or backcrossing of one of these hybrids
with either parent, different forms with intermediate phenotypes between 0. oakesiana
and 0. parviflora may arise. Cleland (1972, pp. 338-339) demonstrated an experimental
example to verify this by combining the ax complexes and the a complexes of a race of 0.
parviflora
(Parviflora-I, Clifton Forge) with those of the three races of 0. oakesiana (Parv
iflora-II, rigens-curvans, Manistique, Ashland A), all races for which the chromosomal
formula of both complexes is known. Only the combination a Clifton Forge (B complex)
f^Ashland A (C complex) gives a 0)14 at meiotic metaphase
I. All other combina
tions show unstable configurations such as 08 and
and 211,08, 04 and III, 012
010
311(
and III, and 0)10 and 04.
The particular strains that Cleland hybridized originated at widely separate localities
with
inMichigan, Virginia, Wisconsin,
and Europe, and we presume that the cytological and
results in hybridizing them are similar to those that occur in nature where
morphological
the ranges of 0. oakesiana and 0. parviflora overlap. Naturally occurring intermediate
forms may approach either parental type phenotypically;
however, as dried herbarium
specimens they are a punishment for a taxonomist who demands from himself the most
scrupulous order up to the most distant nook of his system.
European hybrids between 0. biennis (AB-II) and 0. parviflora (BC-IV) have been
described. According
to Stubbe (1959) they are viable and normal green AC combinations
with plastome II or IV.One such hybrid has been referred to 0. xalbisubcurva Renner (but
the name not validly published) with the complexes albicans from 0. biennis (A genome)
and subcurvans from 0. parviflora (silesiaca; C genome). The distribution of 0. oake
siana in the Appalachian Mountains
and areas adjacent to the east and south of them ap
parently represent such AC hybrids between 0. biennis and 0. parviflora; if so, they rep
resent independent origins of plants with the morphological
features and chromosomal
SYSTEMATICBOTANYMONOGRAPHS
132
constitution
VOLUME 50
of 0. oakesiana.
Similarly, BB genomic combinations between 0. parviflora
are as BB-IV/II normal green and viable, have
been discussed under 0. nutans. They have presumably contributed to enlarging the dis
tribution of 0. nutans further north, but as yet there is no proof for this assumption, be
and 0. biennis
(BC-IV)
cause BB phenotypes
(AB-II), which
II, derived
with plastome
from natural sites, have never been ob
served in cultivation.
HYBRIDS
In this section we have arranged all of the known hybrids, which are usually mor
intermediate to the parents. Hybrids that are very close to one of the parental
phologically
included under the taxon that they most closely resemble. The hy
brids are arranged alphabetically by taxon without regard to the direction of the cross.
phenotypes
are usually
of any formally described taxa
examined. Nearly all of the hybrids involve PTH
taxa, and therefore we have indicated and grouped the names and material according to
Within
we have included nomenclature
each combination
followed by additional non-type material
the genomic
Not
combination.
included here are names
that have not been validly pub
lished; these are listed in a separate section.
Oenothera
argillicola
x Oenothera
parviflora.
SPECIMENS FROM CULTIVATED PLANTS. U.S.A.
vated DUSS-86-1012b,
cultivated
87-349
86-1007
DUSS-77-0171,
Oenothera
biennis
Calvados:
x Oenothera
Germany.
Dietrich
The neotype
Hirmer comment
as O. xconferta,
at Dusseldorf
the pass, Stubbe
that when
s.n.,
Garden
FRANCE.
from
(cultivated
of University
of Dusseldorf,
in 1942), 7 Aug
1985,
(neotype, here designated: MO!).
is cultivated material
as a species
20, culti
1976, Brown
Co.,
75: 513. 1956.-TYPE:
sent to 0. Renner by F. Hilpert
Seeds originally
descendent
from Renner's
original
strain. This en
and appears to represent a stabilized hybrid. Renner
selfed it yields
two phenotypes,
and
one like the plants described
and another with a lower habit, shorter leaves, and smaller flowers. Study
suggests
that one complex
(A), the two complexes
of 0. glazioviana;
intermediate between
A collection
Calais: Collection
0. glazioviana
of this combination
is similar to gaudens
however,
and 0. biennis.
in cultivation
1968.-TYPE:
Hudziok,
13 Jul 1967/A., Hudziok
Zossen,
85-1049
on 3 sheets]). This
types: 0. biennis
(rubricaulis)
and
IId).
3 sheets; isotypes: HAL!
is a hybrid between
and 0. glazioviana
FRANCE: Pas-de
(MO) (012
Prov. Brandenburg
105: 87.
at railway N of Blankenfelde,
s.n. (holotype: HAL-076604!
[2, both mounted
some
[AB combination]
has been examined:
Verh. Bot. Vereins
GERMANY. Brandenburg:
(B) and the other to velans
the plants are phenotypically
of 0. Renner, cultivated DUSS-77-0255,
xcoloratissima
tion]
behind
glazioviana.
strain at the Botanical
DUSS-85-1049
tity was described
Co.,
coast of the channel NE of Caen, dune near Cabourg
seeds of Renner's
Oenothera
Highland
and
xconferta Renner & Hirmer, Biol. Zentralbl.
Oenothera
what
Virginia:
and 211).-West
Virginia: Mineral
41,; 0(10
(MO) (06 and
4,I). [CC combination]
(MO) (06
two European
(coronifera).
pheno
[AB combina
1997
OENOTHERA
Oenothera
xatra de Vries, Gruppenweise
Artbildung
133
NETHERLANDS.
152. 1913.-TYPE:
Amsterdam, cultivated by H. de Vries. No authentic material seen; fig. 66 on p.
152 is here designated as the lectotype. This is an artificial hybrid between
Oenothera biennis "Chicago" and Oenothera glazioviana.
[BB combination]
Oenothera
xlaeta de Vries, Bot. Gaz. (Crawfordsville) 44: 403. 1907.-TYPE:
LANDS. Amsterdam, cultivated by H. de Vries. No authentic material
is an artificial hybrid between
Oenothera
0. biennis and 0. glazioviana.
xlaxa de Vries, Gruppenweise
[BB combination]
144. 1913.-TYPE:
Artbildung
NETHER
seen. This
NETHERLANDS.
cultivated by H. de Vries. No authentic material
Amsterdam,
seen; fig. 61 on p.
as the lectotype. This is an artificial hybrid between O.bi
146 is here designated
ennis "Chicago"
and 0. glazioviana.
[BB combination]
97: 373. 1966] ex Rostan'ski, Feddes
Kappus [Z. Vererbungsl.
9.
GERMANY.
Altenheim
at
Repert. 96:
1985.-TYPE:
Baden-Wuirttemberg:
River Rhine, 24 Aug 1980, Kappus & Rostan'ski 39 (holotype: KTU!). As first
published by Kappus, no type was designated, but Rostan'ski later chose one. The
Oenothera
xoehlkersii
at Kappus's original locality. It is phenotypically
similar to 0.
but was described as a new species that arose via hybridization in
type was collected
glazioviana,
the suaveolens
volving
of 0. biennis. Kappus also observed both pu
to Harte (1994, p. 135), the com
are gaudens (B) from 0. glazioviana
and the other
phenotype
tative parents at the type locality. According
of 0. xoehlkersii
plexes
similar toflavens (B) from0. biennis. [BB combination]
Certain widely scattered plants from Brazil, Germany, India, Italy, Nepal, Portugal,
South Africa, Spain, and Tanzania appear to represent a BB genomic phenotype. Instead
of being 0. grandiflora, however, they most likely represent hybrids between 0. biennis
(or suaveolens phenotype) and 0. glazioviana. The fact that they have different chromo
some configurations from 0. grandiflora strongly suggests that they are of hybrid origin.
Further corroborative evidence is that both 0. glazioviana and 0. biennis are known from
the same or nearby localities. We have grown one strain from Portugal [Porto: Douro
Litoral, between Porto and Lordelo, Ind. Sem. Bot. Gard. Porto 1989 no. 382, cultivated
(08 and 3I1;O 14) (MO) ],which had variegated leaves (green/lutescens).
DUSS-90-2024
to the experimental work of Stubbe (1959), this suggests that the plant may be
combination with plastome II and III, thus representing a hybrid between
According
a BB genomic
0. biennis
(plastome
II) and 0. glazioviana
(plastome
SPECIMENS EXAMINED. BRAZIL. Parana: Guaratuba,
INDIA. Punjab, Nachar
Region
Tuscany.
(BM, DS);
Prov. Lucca:
Taplejung,
SOUTH AFRICA. Natal,
Sennen
Rungwe,
[Himachal
Hara
Pradesh]:
Forte dei Marmi,
et al. 6306586
Umzinto,
Shelley
s.n. (BC, BM, LY).-Orense:
Masoko
Oenothera
Rd,
Kinnaur:
1957, Richards
III).
Boa Vista, Hatschbach
Nachar
(31?33' N, 77?59'
1907, Sprenger
Beach,
Strey 7285, pro parte
9820, pro parte
E), Koelz
s.n. (FI). NEPAL. Panchasi,
(BM). PORTUGAL. Coimbra,
Sierra Santa Eufemia,
& Ramamoorthy
Lorios,
Choupal,
9281
(MO).
(NY).
ITALY.
Staunton
1953, Matos
(M). SPAIN. Barcelona:
Castroviejo
37995
7352
et al. 8321
s.n. (BM, COI).
Manlleu,
1914,
(MA). TANZANIA. Distr.
(BR).
xfallax Renner, Z. Indukt. Abstammungs-Vererbungsl.
18: 176. 1917.-TYPE:
on an artificial hybrid between a strain of 0. biennis referred to as
Munchen
from the Botanical Garden Munich
in Germany and a strain of 0.
glazioviana (as 0. lamarckiana) referred to as Heribert-Nielson
from Sweden. In
based
SYSTEMATIC
BOTANY
MONOGRAPHS
134
a naturally occurring
1965 Rostan'ski designated
VOLUME50
from Poland
specimen
(Silesia,
Street near Olawska
in ruins of Podwale
Street, 16 Jul 1962, Rostan'ski
as a type; it can be considered a neotype.
s.n.; WRSL-07371!)
This hybrid is stable, usually breeding true, and occurring inmixed populations of 0.
Wroclaw,
glazioviana
It also occasionally may be found by itself. According to Ren
constitution of velans (A; 0. glazioviana) and rubens
[AB combination, = 0. velutirubata]
and 0. biennis.
ner (1942), 0. xfallax has a genomic
(B; 0. biennis).
Oenothera
f. rubrinervis Rostan'ski,
xfallax
TYPE: POLAND. "Silesia," Wroclaw
11: 508. 1965.
Fragm. Florist. Geobot.
(Breslau), in the ruins of Chetmonskiego
Street, 16 Jul 1962, Rostan'ski s.n. (holotype: WRSL!).
is a red-nerved phenotype having the same genomic constitution
This
as 0. xfallax.
[AB combination]
Oenothera
14: 19. 1982.-TYPE:
Rostan'ski, Watsonia
xbritannica
South Wales: Glamorgan,
s.n. (holotype: KTU!).
UNITED KINGDOM.
dunes, 14 Sep 1977, Rostan'ski & Ellis
Gower Oxwich
[AB combination]
This hybrid is similar to that described
as 0. xfallax,
and grows
inmixed
populations
of the parental species.
Oenothera
Soldano & Rostan'ski, Riv. Peim St. Nat. 4: 127. 1983.-TYPE:
Prov. Alessandria, Camino, at river Po near the bridge
xfallacoides
ITALY.Region
Piedmont:
of the rd to Trino Vercellese,
This may
represent the hybrid combination
instead of 0. biennis x 0. glazioviana.
biennis,
Soldano, Nat. Bresciana
xadriatica
Oenothera
gion Veneto:
Argenti
Prov. Belluno,
0. biennis
(suaveolens
28: 103. 1992 [1993].-TYPE:
Greto del Piave a Socchieva,
ITALY.Re
300 m, 23 Sep 1989,
s.n. (holotype: PAV; isotype: PAV).
moravica
Jehlifk & Rostan'ski, Folia Geobot. Phytotax.
(Praha) 30: 440.
CZECH REPUBLIC. Jihomoravsky: Mohelno,
at lake Dukovany
above valley
of river Jihlava, ca. 330 m, 25 Jul 1989, Jeli'k & Rostan'ski
(holotype: PR-11398;
The authors believe
The
following
this taxon to be a hybrid between
collections
0. biennis
("victorinii")
glazioviana
231 (C).
x 0.
[AB combination]
represent additional
hybrids between
0. biennis
and 0.
not associated with any formally described names. They fall into two genomic
and phenotypic categories: 1) AA and 2) AB.
1 (AA). This difficult specimen, which apparently arose from hybridization
biennis
s.n.
isotype: KTU).
(0. biennis x 0. glazioviana).
glazioviana
0.
x 0.
phenotype)
1995.-TYPE:
Oenothera
xfallax
s.n. (holotype: TO!).
6 Sep 1976, Soldano
and 0.
glazioviana,
probably represents an AA-combination
and A from 0. biennis): PORTUGAL.Azores: Terceira, Angra,
2 (AB). The
phenotype more
West-Vlaanderen:
following
cultivated
similar to 0. biennis
Ostende,
specimens
represent
than to 0. glazioviana
1982, Wasmund
an AB
between
(A from 0.
1972, Hansen
combination
[i.e., 0. xfallax].
s.n., cultivated DUSS-83-0147
with
a
BELGIUM.
(MO) (012
OENOTHERA
1997
135
and 11) FRANCE. Lot: Ind. Sem. Bot. Gard. Bordeaux 1974, no. 1066, cultivated DUSS
1977-0340 (MO) (012 and
1975 no.
111).HUNGARY. Pest: Ind. Sem. Bot. Gard. Vdcrat6t
2242, cultivated DUSS-85-1050
(MO) (012 and II,), Ind. Sem. Bot. Gard. Vdcrat6t 1975
no. 2241, cultivated DUSS-77-0374
(MO) (012 and III). ITALY.Region Tuscany: Prov.
Pisa: Migliarino,
Ind. Sem. Bot. Gard. Pisa 1974, cultivated DUSS-79-0611
(MO).
The
following
AUSTRIA. Tirol:
1900, Godding
brecht 2529
1460-2,
Sachsen:
Kern
Leiden,
Jehltk 6681
Antwerpen,
Lyon,
Les Marins,
Plzeii
Brunner
s.n. (K).-Lancashire:
1550
s.n. (FR).-Vendee:
Abrantes,
s.n. (K); Southport,
Bayonne,
La Fonte-sur
(Breslau),
s.n. (LZ).
s.n. (B). NETHERLANDS.
Aniol
Brasil,
759
(FI, GZU,
s.n. (LISU).
s.n.
SWITZER
1968, Ed
Muldsworth,
s.n. (BM).-Nottingham:
1933, Foggitt
LD,
1979, Skorgaard
1889, da Cunha
(BR). UNITED KINGDOM. ENGLAND. Cheshire:
1907, Bailey
(PR). DEN
1967, Gutte & Rostan'ski
Terceira, Monte
(BR);
(PR).-Stre
et al. 6676
Jehltk
(Pilsen),
(BR), Rob
1646
Dubois
(Bruinn), Jehli'k 7126
1920, Bornmiller
(L). POLAND. Wroclaw
s.n. (PO).-Santarem:
8032
(C). FRANCE. Basses-Pyrenees:
Juterbog,
s.n. (LZ). PORTUGAL. Azores:
1972, Serra
Massarelos,
Brno
s.n.
Halle,
Lawalree
s.n. (BR); Koksijde,
1912, Reverchon
s.n. (LZ).-Sachsen-Anhalt:
19449
Gent,
1929, Dewildeman
(PR).-Zdpad6esky:
1967, Anderson
Kalvebod,
& Reichgelt
& Rostatiski
LAND. Vaud: Lausanne,
monton
glazioviana).
biennis
(BR).-Oost-Vlaanderen:
Duinbergen,
(P).-Rh6ne:
1971, Gutte
1969, Gutte
(C).-Porto:
s.n.
1942, pro parte (BR). GERMANY. Brandenburg:
Leipzig,
Zuid-Holland:
SOM),
near Praha,
(Copenhagen):
pro parte
Mer, Geerinck
and B from 0.
(W). BELGIUM. Antwerpen:
(A from 0.
Polatschek
s.n. (BR). CZECH REPUBLIC. Jihomoravsky:
1934, Stand
Pruhonice
1968,
13185
Lawalr&e
(BR).-West-Vlaanderen:
MARK. K0bnhavn
Hibon
represent AB-combinations
N of Landeck,
s.n. (BR); Postel,
Middelkerke,
do6esky:
specimens
Stanzerleiten
1963,
McClintock s.n. (BM).-Essex: Colchester, 1881, Gray s.n. (BM).-West-Sussex: Worthing, 1928, Crosfield
s.n. (K).
Oenothera
x [Oenothera
biennis
Oenothera
xvelutina
Bot.
de Vries,
x Oenothera
oakesiana
NETHERLANDS. Amsterdam,
Gaz.
glazioviana].
44:
(Crawfordsville)
cultivated
403.
by H. de Vries. No
1907.-TYPE:
authentic material
seen.
This
is an artificial hybrid between
glazioviana
Oenothera
(as 0.
lamarckiana)].
biennis
x Oenothera
Four genomic
material
and AA;
(as 0. muricata)
x 0.
oakesiana.
are possible
combinations
species: AC, BC, AB,
0. biennis x [0. oakesiana
[AA combination]
from crosses
between
these
two PTH
the first two are by far the more common. The names and
are grouped by these combinations.
AC Combination
Oenothera
xbraunii Doll,
Fl. Baden
berg: at River Dreisam
3: 1077. 1862.-TYPE:
near Freiburg,
GERMANY. Baden-Wulrttem
Jul 1849, Braun
s.n. (lectotype,
here des
ignated: K!).
Doll did not designate
mentions
in his description
cality. Therefore
a type, nor could any authentic specimen be located, but Doll
that A. Braun had seen and collected
this specimen
a hybrid between 0. oakesiana
notype as the plants described
0. oakesiana
is designated
(muricata)
this hybrid at the type lo
here as lectotype. This appears to represent
(C) and 0. biennis
as 0. xalbipercurva
(see Renner
(A), and has the same phe
1956) as a hybrid between
and 0. biennis. Both putative parents grow at the type locality.
SYSTEMATICBOTANYMONOGRAPHS
136
Oenothera
[Flora 131: 196. 1937, nomen
Renner
xalbipercurva
VOLUME 50
ex Hudziok,
nudum]
Bran
Verh.Bot. Vereins Prov.Brandenburg105: 105. 1968.-TYPE: GERMANY.
Jiuterbog, railway W
denburg:
of Hauschteckslust,
The holotype was not among the Hudziok
from Renner's
(C) and 0. biennis
to be a hybrid between
(A). This phenotype
forms a 0 12 and
orous habitat than 0. oakesiana.
(C). Renner obtained
Oenothera
occurs
The genomic
constitution
1968.-TYPE:
[AC combination]
[Ber. Deutsch.
Bot. Ges. 60: 462. 1942,
Prov. Brandenburg
at railway between
105: 106.
Juterbog and Griuna,
2 sheets).
s.n. (holotype: HAL-075231!
is the same hybrid as 0. xalbipercurva,
(A) and percurvans
hybrids and from hybrids gen
Verh. Bot. Vereins
GERMANY. Brandenburg:
21 Jul 1965, Hudziok
This
is albicans
of the putative parents.
ex Hudziok,
of the
populations
and has larger flowers and a more vig
var. impunctata Renner
nudum]
in mixed
regularly
I inmeiosis,
hybridization
xalbipercurva
nomen
the ammophila
these results both with wild-collected
erated by experimental
and therefore we have
to what he (1937, 1938, 1942,
phenotype of 0. oakesiana
these plants appear to conform
discussion
parental species,
at HAL,
collections
a European
involving
1950, 1956) considered
s.n.
1967, Hudziok
annotated by him as this entity as a neotype. These plants rep
phenotype of 0. oakesiana, ammophila. Judging
selected another collection
resent a hybrid
15 Aug
2 sheets).
(neotype, here designated: HAL-075229!
that the stems do not have red
except
spots.
The
following
additional
AUSTRIA. Wien
oakesiana).
specimens
(Vienna),
also represent AC-combinations
1888, Rechinger
(A from 0.
s.n. (LD). BELGIUM. Liege:
and C from 0.
biennis
Chaudfontaine,
1943,
Isadc
son s.n. (BR). CZECHREPUBLIC.
Severocesky:Chomutov, Jehlfk6263 (PR), Jehltk 6757 (PR);Ustf (Aussig),
1899, Schubert (BR).-Vychodocesky: Chaceni,1900, Fleischer s.n. (BR).GERMANY.
Hamburg:Harburg,Wil
helmsburg,
1909, Mohr
s.n. (HBG); Reiherstieg,
(LZ). FRANCE. Bas-Rhin:
Strasbourg,
1909, Schmidt
1873, Stahl
s.n. (HBG).-Sachsen:
Leipzig,
1977, Gutte
s.n.
s.n. (FI).
BC Combination
xindivisa
Oenothera
Hudziok,
Reihe
Naturwisse
Wiss.
12: 709.
Z. Martin-Luther-Univ.
1963.-TYPE:
railway near "Millionenbruicke,"
075234!
10 Aug
Math.
Jiuterbog,
s.n. (holotype: HAL
1962, Hudziok
2 sheets).
The disposition
combination.
Halle-Wittenberg,
GERMANY. Brandenburg:
reflects the opinion of Hudziok.
Hudziok
This collection
appears to be a BC
it as a species of hybrid origin between
described
two European
phenotypes:jueterbogensisand ammophila.
Oenothera
xissleri
var. silesiacoides
14: 424. 1979.-TYPE:
Rostan'ski & Jehllfk, Folia Geobot. Phytotax. (Praha)
CZECH REPUBLIC. Zaipad6esk9: Plzeni-Koterov (Pilsen),
320 m, 5 Jul 1972, Jehlik 6260
This hybrid involves
with
a European
(holotype: PR!).
phenotype
of 0. oakesiana,
red stems in the lower part of the plant and red-nerved
Additional
collections
of this cross
Jihocesky:
Chodovo,
Jehl(k 6261
berg-Ring,
Rostaniski
74 (HBG).
(B from 0. biennis
(PR). GERMANY. Bavaria:
POLAND. Wroclaw
syrticola.
and C from 0. oakesiana)
Sonthofen,
(Breslau),
It is a form
leaves.
1966, Dorr
1959, Rostatiski
are: CZECH REPUBLIC.
s.n. (M).-Hamburg:
s.n. (KTU).
Wins
OENOTHERA
1997
Oenothera
xpseudocernua
Hudziok,
137
Verh. Bot. Vereins
Prov. Brandenburg
Potsdam,
1974.-TYPE:
GERMANY. Brandenburg:
(holotype: HAL, not located).
111: 100.
25 Sep 1971, Hudziok
s.n.
like 0. parviflora; it was described
as a hybrid involving a European phenotype of 0. oakesiana, ammophila, and the chicagi
nensis phenotype of 0. biennis. The description suggests a BC combination phenotype.
From the description
this plant is phenotypically
AB Combination
xheiniana Teyber, Verh. K. K. Zool.-Bot. Ges. Wien 46: 469. 1896. Oenothera
biennis subsp. heiniana (Teyber) Love & Love, Opera Bot. 5: 258. 1961.-TYPE:
AUSTRIA. Inundation dam at river Donau (Danube) near Wien
(Vienna), Sep
Oenothera
1896, Teyber's.n. (lectotype, designated by Rostan'ski & Forstner, 1982: WU!).
The phenotype of this plant is biennis-like. Renner examined the type and another
collection made by Teyber in 1900 (WU); he commented that it is similar to the chicagi
nensis phenotype of 0. biennis, but has larger flowers. Teyber observed both putative par
ents at the type locality, and considered this taxon to be a hybrid between the two species.
The following specimens also appear to represent an AB-combination
(A from 0.
and B from 0. biennis: NORWAY. Telemark: Kragero, 1889, Landmark s.n. (0).
(Vienna), Prater, 1895, Fritsch s.n. (GZU), 1901, Arbesser s.n. (GZU).
oakesiana
AUSTRIA:Wien
AA Combination
xclavifera Hudziok, Verh. Bot. Vereins Prov. Brandenburg 105: 104. 1968.
TYPE: GERMANY. Brandenburg: on sandy places in Luckenwalde,
17 Aug 1967,
4 sheets).
Hudziok s.n. (holotype: HAL-076601!
Oenothera
This
Hudziok
Oenothera
hybrid has
the phenotype
as a hybrid between
biennis
Four genomic
of 0.
villosa
subsp. villosa.
two European phenotypes:
x Oenothera
ammophila
It was
described
x editicaulis.
by
parviflora.
combinations
are possible
from crosses
between
species: AC, BC, AB, and BB; the first two are by far more common.
terial are grouped by these combinations.
these
two PTH
The names and ma
AC Combination
The following
specimens
represent AC-combinations
(A from 0. biennis and C from
0. parviflora).GERMANY.
Berlin: Charlottenburg,SpandauerBerg, 1911, Schulz s.n. (B);
Westend, 1886, Sydow s.n. (FR);Berlin-Zehlendorf,1922, Sydow s.n. (GZU). POLAND.
Wroclaw
(Breslau),
1958, Rostan'ski
s.n. (KTU).
BC Combination
Oenothera
11: 504. 1965.
xpseudochicaginensis
Rostan'ski, Fragm. Florist. Geobot.
TYPE: POLAND.Wroclaw
(Breslau), at Szczesliwa Rd, 17 Jul 1961, Rostan'ski
s.n. (holotype: WRSL!).
Rostan'ski
suggests
that this entity is of hybrid origin between
two European
pheno
SYSTEMATICBOTANYMONOGRAPHS
138
VOLUME 50
types, silesiaca and rubricaulis;however, ithas subterminalfree sepal tips like0. parvi
flora.
AB Combination
The following
represent an AB-combination
specimens
(A from 0. biennis
and B
Sachsen:Leipzig, 1978,Gutte s.n. (LZ), 1979,Gutte s.n.
from0. parviflora):GERMANY.
(LD).
BB Combination
The
following
specimen
most
probably
a BB-combination
represents
(B from
0. parviflora andB from0. biennis): BELGIUM.
Limburg: St. Trond, Thielens 231, pro
parte (FH).
Oenothera
x Oenothera
biennis
All of the collections
villosa
subsp. strigosa.
of this cross appear to represent an AB combination
(A from 0.
villosa andB from0. biennis).
CULTIVATED SPECIMEN. CANADA. British
4547, DUSS-82-0389,
SPECIMENS EXAMINED. CANADA. British
ton: Garfield
4765
Co.,
(MICH),
(MICH
Creek,
Ilia, St. John et al. 9237
5859
(ARIZ, BH, DS,
2 sheets, WS),
(WS); Vila,
Kamilehe
7615
10603
Suksdorf
Rd, Freer
(MICH),
(BH),
81 (WTU).
Botanie
from seeds
cultivated
Valley,
from Wagner
Columbia:
(WS). Klickitat
F, GH, MICH,
7652
(WS),
Botanie
NY, ORE,
7653
from Suksdorf
grown
Snohomish
Valley,
Co., Bingen,
Co., Marysville,
PH, RM),
7654
(MICH),
5859,
Perry
Bartlett
Bartlett
4506
3102
7576
(WS),
2691
(UBC). U.S.A. Washing
(BH), Suksdorf
(MICH
7807
(BH),
(BH, RSA).
7891
Mason
s.n. (WS). Spokane
1922, Sprague
2066
(GH),
2 sheets, WS),
7577
(WS),
7915
Co., Elma
Co., Hangman
s.n. (WS).
1889, Suksdorf
Oenothera
Columbia:
pro parte (MO).
biennis
x Oenothera
Both of the possible
genomic
observed. They are AA and AB
villosa
subsp. villosa.
combinations
in a cross between
(A from 0. villosa
these taxa have been
and B from 0. biennis).
AA Combination
Oenothera
xpolgari
GARY. At
Rostan'ski, Acta Bot. Acad.
river Danube
Sci. Hung.
near Dunaharaszti,
12: 347. 1966.-TYPE:
19 Aug
HUN
1946, Vajda & Boros
s.n.
(holotype:BP!).
This taxon was described as a hybrid between the suaveolens phenotype (0. biennis)
and the depressa phenotype (0. villosa subsp. villosa). Study of the description and the
type suggest a phenotype similar to 0. villosa subsp. villosa, but with larger flowers.
Oenothera
xwienii Renner
Florist.
Geobot.
[Flora 131: 198. 1937, nomen nudum] ex Rostan'ski, Fragm.
23: 289.
1977.-TYPE:
POLAND. "Heubude" near Gdansk
7 Sep 1974, Rostan'ski s.n. (holotype: KTU!).
Rostan'ski selected the holotype from plants collected at the original locality of Ren
i rubricaulis (0. biennis) phe
ner's strain. Rostan'ski indicated it to be a hybrid between
(Danzig),
1997
OENOTHERA
(0. villosa
notype and a depressa
subsp. villosa)
139
Renner's
phenotype.
analysis
rubricaulis
that it is a hybrid between
indicates
and bauri with a
two of the European phenotypes:
tingens (A) and undans (A). This conforms with our observations in
field. This combination is a stable hybrid, but it occurs only at the type
constitution
genomic
the experimental
locality.
AB Combination
Oenothera
Renner
xdrawertii
Bot. Ges. 63: 135. 1950, nomen nudum] ex
HUNGARY. River
Sci. Hung. 12: 341. 1966.-TYPE:
[Ber. Deutsch
Rostan'ski, Acta Bot. Acad.
Tisza
in Szolnok,
13 Aug
This taxon was described
(A) and suaveolens
s.n. (holotype: WRSL!).
1964, Rostan'ski
two of the European phenotypes,
as a hybrid between
(B). The phenotype
to what Renner
conforms
analyzed under this name from a collection
in France with
bauri
(1950) described
the genomic
and
lax
constitution
ans (A;depressa) andflavens (B; suaveolens).
The following
from Hungary,
are morphologically
collections
Szolnok,
very similar to the phenotype
of the
HUNGARY. Cult. at Botanic Garden Wroclaw,
as 0. xdrawertii:
specimen described
3/65
Rostan'ski
(KTU).
seeds
SLOVAKIA. Vaipadoslovensky:
Nove
Zamky, ?tenkow,Jehlik 7214 (PR).
The following
villosa
represent a hybrid between
collections
subsp. villosa
and 0. biennis
(depressa)
(rubricaulis),
nation. They are what has been called 0. xhoelscheri
ever validly published
villosa
(1942, 1950, 1956) described
(bauri) and 0. biennis
0.
phenotypes,
in an AB combi
resulting
(but neither Renner nor Rostan'ski
description of 0. xhoelscheri
this name). Rostan'ski's
form to what Renner
two European
seems to con
as a hybrid between 0. villosa
subsp.
(rubricaulis).
SPECIMENS
EXAMINED:
AUSTRIA.Tirol: Brixlegg, 1936, Schneider s.n. (W). CZECHREPUBLIC.
near Praha, Jehlfk 6795
Pruhonice
Stredocesky:
HUNGARY. Tunde
Street
8 Aug
Jehlik & Kreilovd
Rychnov,
(PR).-Vychodo6esky:
in Budapest-Kobanya,
s.n.
1964, Rostanski
(WRSL).
GERMANY.
7128
(PR).
Sachsen:
Leipzig-Mockern, 1970, Gutte s.n. (LZ).-Sachsen-Anhalt: Coswig nearWittenberg, 1967, Gutte & Rostanski
s.n. (LZ, WRSL).
NORWAY. Oslo,
sent by F. Schotz
seeds
Aleksandrow
1064
(GZU, LD,
taniski s.n. (KTU, LZ).
s.n. (0). POLAND. Leslau
SOM);
Poznani
at river Wisla
26 Jul 1966, Rostaniski
collection),
10 Jul 1963, Olesinski
Kujawski,
(LZ), Koziol
1889, Mol
from 0. Renner's
s.n. (WRSL); Wroclaw
(Posen),
SLOVAKIA. Stredoslovensky:
Lwowek,
Martin,
(Breslau),
Koziol
Jehlik 6688
910
(Weichsel)
s.n. (WRSL);
(cultivated
Ciechocinek,
from
distr.
1969, Gutte & Rostan'ski
(LD, SOM),
s.n.
1969, Gutte & Ros
(PR).
Specimens cited by Rostaniski (1975) (as 0. xhoelscheri): RUSSIA.Kursk, ShchigryTim, 1897,Widusi
askaja
s.n. (LE); Tambov,
zoski s.n. (LE); Kiyev,
Oenothera
1924, Wasiliew
1956, Senczenko
glazioviana
As far as is known
Oenothera
xmurinella
s.n. (LE). UKRAINE. Kherson
at River Dnepr
(Dnieper),
1906, Pac
s.n. (LE).
x Oenothera
oakesiana.
this hybrid combination
de Vries,
Ber. Deutsch.
NETHERLANDS. Amsterdam,
cultivated
does not occur in nature.
Bot.
Ges.
26a:
669.
by H. de Vries. No
1908.-TYPE:
authentic material
seen.
This
(as 0.
is an artificial hybrid between
lamarckiana nanella).
0. oakesiana
[AB combination]
(as 0. muricata)
x 0. glazioviana
SYSTEMATICBOTANYMONOGRAPHS
140
Oenothera
x Oenothera
glazioviana
As far as is known
parviflora.
xdensa
de Vries,
LANDS. Amsterdam,
155. 1913.-TYPE:
Artbildung
by H. de Vries. No authentic material
Gruppenweise
NETHER
cultivated
seen; fig.
69 on p. 155 is here designated
is an artificial hybrid
This
does not occur in nature, but two genomic
this hybrid combination
have been formed, BB and AC.
combinations
Oenothera
VOLUME 50
as the lectotype.
the phenotype
involving
cruciata
of 0. parviflora.
[BB
combination]
Oenothera
163. 1913.-TYPE:
xgracilis de Vries, Gruppenweise
Artbildung
LANDS. Amsterdam,
cultivated by H. de Vries. No authentic material
78 on p. 165 is here designated as the lectotype.
is an artificial hybrid between 0. glazioviana
This
ata phenotype.
Oenothera
seen; fig.
the cruci
[AC combination]
xhero de Vries, Gruppenweise
Amsterdam,
NETHERLANDS:
Artbildung 327. 1913.-TYPE:
cultivated by H. de Vries. No authentic material seen; fig. 120 on p.
330 is here designated
This
and 0. parviflora with
NETHER
as the lectotype.
is an artificial hybrid
the phenotype
involving
cruciata
of 0. parviflora.
[AC
combination]
Oenothera
xpercruciata
de Vries, Gruppenweise
LANDS: Amsterdam,
Artbildung 320. 1913.-TYPE:
by H. de Vries. No authentic material
cultivated
117 on p. 320 is here designated
is an artificial hybrid
This
involving
NETHER
seen; fig.
as the lectotype.
the phenotype
cruciata
of 0. parviflora.
[AC
combination]
Oenothera
glazioviana
x Oenothera
villosa
subsp. villosa.
xpurpurans Borba's, Kert 1902: 204. 1902.-TYPE:
HUNGARY. Budapest,
22 Jul 1901, de Borbas s.n. (BP, presumably destroyed).
This taxon was described as a hybrid involving one of the European phenotypes of 0.
Oenothera
Kobanya,
villosa
subsp. villosa
(hungarica).
Borbas observed
the putative parents at the type local
ity. [AA combination]
Oenothera
glazioviana
x Oenothera
wolfii.
The naturally occurring hybrids of this combination
can have either an AB or an AA
genomic constitution. The following collections have the approriate phenotypes
combinations but have not been studied experimentally.
for these
AA Combination
SPECIMENS EXAMINED. U.S.A.
sheets). Humboldt
Hwy.
101, 1986,
Co., S of Trinidad,
Imper s.n. (HSC),
California:
Del Norte
near Moonstone
1987,
Beach,
Imper s.n. (HSC).
Co., N of Smith River,
1987,
Imper
s.n. (HSC,
1987, Imper s.n. (HSC); S side of Klamath
River,
3
1997
OENOTHERA
141
AB Combination
SPECIMENS EXAMINED. U.S.A.
boldt Co.,
S of Clam Beach
Oenothera
California:
exit, Hwy.
Del Norte
101, 1986,
x Oenothera
grandiflora
Co., Crescent
Imper s.n. (HSC);
City,
1987,
S of Trinidad,
Imper s.n. (HSC). Hum
1986,
Imper s.n. (HSC).
biennis.
The following specimens are phenotypically
similar to 0. grandiflora, but show some
influence of 0. biennis and are PTH plants (Steiner & Stubbe 1984; Schumacher 1987;
Schumacher et al. 1992; Schumacher & Steiner 1993). Some of them are also cited under
0. grandiflora, because they are exceedingly similar to it.
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. U.S.A.
Steiner
307
s.n., cultivated
(MO)
Mobile
(0 14; 0)10
Co., Chastang,
4, 1983, Steiner
DUSS-84-304
and 211; 0)10 and 04),
1983, Steiner s.n., cultivated
s.n., cultivated
DUSS-84-357,
jamesii x Oenothera
Oenothera
DUSS-84-350
84-358
villosa
(MO)
s.n. (MO). U.S.A.
Conecuh
Co., Castleberry
(A), 1983,
Oklahoma:
(MO)
(012
and 11I).Washington
Co., Route
(014).
subsp. villosa.
SPECIMENS EXAMINED. SOUTH AFRICA. Cape, Trandkei,
Gorden-Gray
Alabama:
and 211;012
and 1il), 1983, Steiner s.n., cultivated DUSS-84
1983, Steiner s.n., cultivated DUSS-84-309
(MO) (0310 and 211).
(MO) (010
Oklahoma
Co.,
Haven,
Bashee
S of Oklahoma
River mouth
City, Meyers
near Lagoon,
1966,
80 (OKL).
Oenothera parviflora x Oenothera villosa subsp.villosa.
Oenothera
xslovaca
1979.-TYPE:
Jehlik & Rostan'ski, Folia Geobot. Phytotax.
(Praha) 14: 413.
SLOVAKIA.Vapadoslovensky:
Nove Zamky, 15 Jul 1974, Jehl[k
6750 (holotype: PR; isotype: KTU!).
This name was published earlier by these authors (Preslia 49: 94. 1977) as a no
men nudum. The taxon was described as a hybrid between two of the European pheno
types, depressa and turoviensis; it is phenotypically
like 0. biennis. [AB combination]
DOUBTFUL AND EXCLUDED NAMES
communis race japonica Guffroy ex H. Leveille, Bull. Acad. Int. Geogr. Bot.
O[elnothera
19: 329. 1909.-TYPE:
JAPAN. Sapporo, 2 Sept 1886, Faurie 1302; no material
located.
Oenothera
corymbosa Lamarck, Encycl. 4 (2): 554. 1798, non Oenothera
corymbosa
Sims. No authentic material
located in the Lamarck Herbarium.-Because
Lamarck states that this species has the floral tube equal in length to the ovary,
the capsule ovate to oblong with sparse pubescence, a flexuous stem with some
short hairs, and a nodding stem apex, flowers crowded toward tips of stems form
ing a corymbose
inflorescence, each flower pedunculate, and yellow petals, it
does not appear to be a member of Oenothera
subsect. Oenothera,
and more
likely represents a member of sect. Kneiffia.
Oenothera
dubia E. H. Krause
Oenothera
dubia f. latifolia E. H. Krause
in Sturm, Flora Deutschl.
ed. 2, 9: 192. 1901.
in Sturm, Flora Deutschl.
ed. 2, 9: 192. 1901.
SYSTEMATICBOTANYMONOGRAPHS
142
Oenothera
gauroides Hornemann,
Hort. bot. Hafn.
cultivated plants at the Botanical
cultivated material
(without data, 1829), and GOET!
they are a mixture
Oenothera
P. Miller,
glabra
at B!
Gard.
Miller
unsuccessful.
Oenothera
seen.
seeds sent by Homemann),
C!
but
(1811,
and 0. biennis
(4 specimens)
(3 specimens).
this name may apply to 0. parviflora.
dict., ed. 8, 3. 1768.-TYPE:
U.S.A.
associated with
expanded his diagnosis
by comparison
Virginia.-A
this name has been
to his other species,
from which he says it differs "in having smooth stalks which
pale green color." This name cannot be associated
may
from
first intro
No definite authentic material
search by C. Jarvis for extant Miller material
0. angustifolia,
described
which were
(1810), are labeled "Oenothera gauroides,"
of 0. parviflora
To judge from the description,
1: 362. 1813.-TYPE:
Garden of Copenhagen,
duced in 1808 from Baltimore, Maryland.
Contemporary
VOLUME 50
are
definitely with any taxon; it
represent 0. parviflora.
J. Lehmann,
salicifolia
Oenothera
salicifolia
thentic material
ex Seringe
(1832).-TYPE:
from Hort. Dresd.
lia Desfontaines
hort. bot. Hamburg
Desfontaines
ex G. Don
Desfontaines
Sem.
1824:
Lehmann
located from either source.-The
1824,
non
salicifolia
refers to Oenothera
[Dresden] and Hort. Vindob.
is unknown. The description mentions
20.
(1828) nec Oenothera
salicifo
[Vienna]; no au
fate of the Lehmann herbarium
that the stems and leaves are glabrous, and
thus thenamemay apply to0. biennis or 0. parviflora.
NAMES NOT VALIDLY PUBLISHED
In order to avoid repetitive explanations
in the text and below, this part is divided
into
four sections. The first lists names not validly published by authors other than H. de Vries,
0. Renner
maining
(except names based on wild-collected
three sections
ner, only names based on experimental
Oenothera
Oenothera
according
sect. Oenothera
subsect. Euoenothera
Rostan'ski,
under the provisions
Fragm. Florist. Geobot.
as syntypes:
"Wagonownia Wroclaw
P. H. Raven, W. Dietrich
Oenothera
& Stubbe,
biennis L. Although
allow
this name is now contrary to the ICBN (1994) Art.
21.3, and not validly published
acutifolia
1: 492. 1840. Not validly
to the ICBN (1994) Art. 21.3.
able in 1980 when published,
two specimens
and the re
hybridizations).
Syst. Bot. 4: 252. 1980 ["1979"].-TYPE:
Oenothera
and J. P. Lotsy,
by these three authors (for Ren
Torrey & A. Gray, Fl. N. Amer.
subg. Euoenothera
published
plants),
list names not validly published
of Art. 32.lb.
11: 501. 1965. Rostan'ski
1) POLAND. Silesia, Wroclaw
Glowny,"
9 Aug
1958, Rostan'ski
cited
(Breslau), railroad near
s.n. (WRSL!); 2) cul
tivated from seeds of the cited Rostan'ski collection. Published without
tion of the type [ICBN (1994) Art. 37.1]. = 0. biennis.
designa
OENOTHERA
1997
Oenothera
ammophila
lished because
143
var. rhodoneura Renner, Flora 131: 222. 1937. Not validly pub
a Latin diagnosis was not included [ICBN (1994) Art. 36.1]. = 0.
oakesiana.
Oenothera
beckeri Renner, Ber. Deutsch. Bot. Ges. 60: 457. 1942; Baerecke, Flora 138:
81. 1944. Not validly published because a Latin diagnosis was not included in ei
ther publication [ICBN (1994) Art. 36.1]. = 0. villosa subsp. villosa.
Oenothera
biennis var. angustifolia Renner, Planta 47: 244. 1956. Not validly published be
cause a Latin diagnosis was not included [ICBN (1994) Art. 36.1]. = 0. biennis.
biennis var. cruciata Klebahn, Jahrb. Hamburg. Wiss. Anst. Beih. 31: 5. 1914,
nomen nudum, non Oenothera biennis var. cruciata (Nuttall ex G. Don) Torrey
& A. Gray (1840). = 0. biennis.
Oenothera
Oenothera biennis subsp. nuda Love & Love, Opera Bot. 5: 258. 1961, nomen nudum.
This name is neither a new combination, as Love & L6ve intended, nor a new
name. The presumed basionym, Oenothera nuda, was not validly published by
Renner (1956) or Rostan'ski (1968). = 0. biennis.
Oenothera
biennis
f. ochroleuca
Gaiyer, Magyar
Bot. Lap. 16: 59. 1917, nomen nudum. =
0. biennis.
Oenothera
biennis var. sulphurea Klebahn,
1914, nomen nudum. = 0. biennis.
Oenothera
cantabrigiana B. M. Davis, Genetics 25: 433. 1940. Oenothera biennis var.
cantabrigiana B. M. Davis, Rhodora 59: 16. 1957. Neither name is validly pub
lished because a Latin diagnosis was not included [ICBN (1994) Art. 36.1]. = 0.
biennis.
Oenothera
var. parviflora Renner, Planta 47: 233. 1956. Not validly pub
chicaginensis
lished because a Latin diagnosis was not included [ICBN (1994) Art. 36.1]. = 0.
biennis.
Oenothera
chigagoensis
Renner
Jahrb. Hamburg, Wiss.
ex Cleland & Blakeslee,
Anst. Beih.
31: 23.
Proc. Natl. Acad. U.S.A.
16:
189. 1930, nomen nudum.
communis race vrieseana H. Leveille, Bull. Acad. Int. Geogr. Bot. 19: 329.
1909, nomen nudum. Leveille included two validly published, earlier names, 0.
grandifiora and 0. glazioviana.
O[e]nothera
Onagra
guttata E. Greene ex Gates, Amer. Naturalist 45: 589. 1911; Mutation factor in
evolution 30. 1915. Oenothera guttata Cockerell, Gard. Chron., ser. 3, 55: 84.
1914. In the 1911 publication Gates merely discussed Greene's unpublished name
written on a specimen from Sierra Co., New Mexico
[Kingston, 1Aug 1904, Met
calfe 1193 (BM, CAS, GH, NY, MO 2 sheets, POM) ], indicating that it is an in
teresting specimen, which he briefly characterized. In the 1915 publication Gates
SYSTEMATICBOTANYMONOGRAPHS
144
cited the BM sheet of this collection,
indicating
VOLUME 50
that he tentatively refers it to 0.
hookeri var. angustifolia, and mentioned
that itmay be worthy of specific status.
Cockerell mentioned the 1911 Gates publication and the binomial, Oenothera gut
tata; then he tentatively commented that it probably represents 0. irrigua Wooten
& Standley. None of these publications can be considered
name, because none fulfill ICBN (1994) Art. 34.1(a).
to have validated either
xhoelscheri Renner [Ber. Deutsch. Bot. Ges. 60: 460. 1942, nomen nudum] ex
Rostan'ski, Fragm. Florist. Geobot. 14: 189. 1968. Published without indication
of a type [ICBN (1994) Art. 37.1]. = 0. biennis x 0. villosa subsp. villosa.
Oenothera
Oenothera
xhoelscheri
var. albinervis Rostan'ski, Fragm. Florist. Geobot.
indication of a type [ICBN (1994) Art. 37.1].
14: 192. 1968.
var. rubricalyx Rostaniski, Fragm. Florist. Geobot.
indication of a type [ICBN (1994) Art. 37.1].
14: 191. 1968.
Published without
Oenothera
xhoelscheri
Published without
Oenothera
issleri Renner [Ber. Deutsch Bot. Ges 63: 134. 1950, nomen nudum] ex Ros
tan'ski, Fragm. Florist. Geobot. 11: 514. 1965. Rostan'ski cited three collections
as syntypes: 1) grown from seeds of Renner's collection sent by F. Schotz, 1964,
from: France, Detp. Haut-Rhin, Colmar, 15 Jul 1963, Rostan'ski 26/63 (WRSL!);
2) Poland, Breslau, 1958, Rostan'ski s.n.; 3) cultivated from seeds from the Ros
tan'ski collection of 1958. Published without indication of a type [ICBN (1994)
Art. 37.1]. Described as a hybrid involving a European phenotype of 0. oake
siana (syrticola). Renner studied this entity both as artificially created hybrids
and as wild-collected
hybrids. It has a 0 14 inmeiosis and breeds true. It seems
to arise whenever
the parents grow together and has the genomic
(C). = 0. biennis x 0. oakesiana.
constitution
of
rubens (B) and curvans
Oenothera
italica Rostan'ski & Soldano, Fragm. Florist. Geobot. 27: 376. 1981. Rostan'ski
cited two specimens as syntypes: 1) Italy, Fossa dell'Abate near Viareggio, 1977,
Soldano s.n.; 2) cultivated from seeds of the Soldano collection. Published with
out indication of a type [ICBN (1994) Art. 37.1]. = 0. biennis.
Oenothera
lamarckiana var. brevistylis de Vries ex Pohl, Oesterr. Bot. Z. 45: 206. 1895,
nomen nudum. A short-styled phenotype that arose in experiments. = 0. glazio
viana.
Oenothera
lamarckiana var. lata de Vries ex Pohl, Oesterr. Bot. Z. 45: 209. 1895, nomen
nudum. = 0. glazioviana.
Oenothera
mississippensis
Bartlett
in Klaphaak & Bartlett, Amer.
J. Bot. 9: 458.
1922,
nomen nudum.
Oenothera
mollis Renner, Planta 47: 328. 1956. Not validly published. Renner gave this
name to a strain ("O. nova von Juterbog") without description in 1937 (Flora 131:
194). In the 1956 publication he did not give a Latin diagnosis; therefore this
name is not validly published [ICBN (1994) Art. 36.1]. = 0. villosa subsp. villosa.
OENOTHERA
1997
Oenothera muricata
145
var. ammophila (Focke) Stomps, Receuil Trav. Bot.
Neerl. 41: 142. 1948. This name is not validly published, because Stomps men
tions it only in discussion but does not accept it himself [ICN (1994) Art. 34.1].
subsp. germanica
subsp. issleri Love & Love, Opera Bot. 5: 257. 1961, nomen nudum.
as the authors intended, nor a new
is neither a new combination,
name. The presumed basionym, 0. issleri Renner (1956), was not validly pub
lished at the time.
Oenothera muricata
This name
Oenothera
muricata
subsp. syrticola Tischler, Chromos.
nomen nudum. = 0. oakesiana.
Gefasspf.
Mitteleur.
58. 1950,
var. rhodoneura Renner, Flora 131: 222. 1937. Not validly published
because Renner did not include a Latin diagnosis [ICBN (1994) Art. 36.1]. = 0.
Oenothera muricata
oakesiana.
Oenothera
nervosa Hornemann ex Sweet, Hort. Brit. ed. 2: 199. 1830, nomen nudum [see
ICBN (1994) Art. 32.1 ex. 3]. = 0. villosa subsp. villosa.
Oenothera
[Planta 47: 244, fig. 17. 1956 (without Latin diagnosis)] ex Ros
Florist.
tan'ski, Fragm.
Geobot. 14: 192. 1968. Rostan'ski cited two specimens,
both cultivated from seeds of Renner's original collection sent by F. Schotz: 1)
nuda Renner
France, Dep. Bas Dauphine,
Saint-Laurent-du-Pont
(cultivated from seeds), 21
Jun 1966, Rostan'ski 14/65 (WRSL!); 2) 25 Jul 1966, Rostan'ski 14/65. Published
by Rostan'ski without indication of a type [ICBN (1994) Art. 37.1].
(Horn) 3: 7. 1951, nomen nudum.
Oenothera
parviflora subsp. syrticola
= 0. oakesiana.
Oenothera
renneri f. mollis Renner ex Rostan'ski, Fragm. Florist Geobot. 11: 510. 1965.
he was making a new combination based on Oenothera mol
lis Renner, but Renner never validly published this name. This forma is also not
validly published because no type was designated [ICBN (1994) Art. 37.1]. = 0.
Janchen, Phyton
Rostan'ski believed
villosa subsp.villosa.
Oenothera
Renner [Flora 136: 146. 1942] ex Gutte & Rostan'ski, Ber. Ar
Sachs. Bot. 11: 187. 1981. No Latin diagnosis was provided, and there
fore the name is not validly published [ICBN (1994) Art. 36.1]. = 0. oakesiana
(as 0. ammophila) x 0. biennis.
xrigirubata
beitsgem.
Oenothera
strigosa subsp. mollis Renner ex Weihe in Garcke, Ill. Fl. Deutschland ed. 23,
982. 1972. This trinomial cannot be considered a new combination
(the pre
sumed basionym was never validly published) nor a new taxon (neither a Latin
diagnosis [ICBN (1994) Art. 36.1] nor designation
37.1] were given). = 0. villosa subsp. villosa.
Oenothera
of a type [ICBN (1994) Art.
suzukiana Jean & Linder, Cytologia 44: 775. 1979. Not validly published
cause no type was indicated [ICBN (1994) Art. 37.1]. = 0. jamesii.
be
SYSTEMATICBOTANYMONOGRAPHS
146
syrticola Bartlett, Cybele Columb.
Oenothera
1: 38. 1914, nomen nudum. = 0. oakesiana.
vrieseana H. Leveille, Bull. Acad. Int. Geogr. Bot.
O[e]nothera
nudum. = 0. glazioviana or 0. grandiflora.
Oenothera
VOLUME 50
19: 329. 1909, nomen
wratislaviensis
Rostan'ski, Fragm. Florist. Geobot. 11: 510. 1965. Rostan'ski
cited two specimens as syntypes: 1) Poland. Silesia, Wroclaw
(Breslau), Piekna
Street near railway station, 9 Jul 1959, Rostan'ski s.n. (WRSL!); 2) cultivated
from seeds from the Rostan'ski collection.
Published without
indication of a type
[ICBN (1994) Art. 37.1].
Names Not Validly Published
- H. de Vries
The names of H. de Vries that were never validly published are listed below in al
phabetical order. These names were either published without a description or without in
dication of rank. Most of the plants on which the names are based are simply mutants of
de Vries's Oenothera
lamarckiana (=O. glazioviana). He gave them new names in his
publications,
Oenothera
because he believed
they were newly evolved
blandina mut. spiralis de Vries, Bot. Gaz.
entities.
(Crawfordsville)
63: 24. 1917. = 0.
glazioviana.
Oenothera
grandiflora mut. gigas de Vries, Bot. Gaz.
(Crawfordsville)
65: 385. 1918. =
(Crawfordsville)
65: 384. 1918. =
0. grandiflora.
Oenothera
grandiflora
mut.
lorea de Vries, Bot. Gaz.
0. grandiflora.
Oenothera
grandiflora mut. ochracea
= 0. grandiflora.
de Vries, Bot. Gaz.
(Crawfordsville)
65: 382. 1918.
Oenothera grandifloramut. semigigasdeVries, Bot. Gaz. (Crawfordsville)65: 387. 1918.
= 0. grandiflora.
Oenothera
lamarckiana mut. ablata de Vries, Z. Indukt. Abstammungs-Vererbungsl.
129. 1929. = 0. glazioviana.
52:
Oenothera lamarckiana
mut. angustifoliadeVries, Z. Indukt.Abstammungs-Vererbungsl.
52: 123. 1929. = 0. glazioviana.
Oenothera
lamarckiana mut. auricula de Vries, Z. Bot. 15: 376. 1923. = 0. glazioviana.
Oenothera
lamarckiana mut. aurita de Vries, Z. Bot.
15: 376. 1923. = 0. glazioviana.
Oenothera lamarckianamut. cinerea deVries, Z. Indukt.Abstammungs-Vererbungsl.52:
134. 1929. = 0. glazioviana.
Oenothera
lamarckiana mut. compacta de Vries, Z. Bot.
15: 400. 1923. = 0. glazioviana.
OENOTHERA
147
Oenothera
lamarckiana mut. crinita de Vries, Z. Indukt. Abstammungs-Vererbungsl.
136. 1929. = 0. glazioviana.
52:
Oenothera
lamarckiana mut. cucumis de Vries, Z. Indukt. Abstammungs-Vererbungsl.
130. 1929. = 0. glazioviana.
52:
Oenothera
lamarckiana mut. decipiens
0. glazioviana.
1997
de Vries, Ber. Deutsch.
37: 70. 1919. =
15: 403. 1923. = 0. glazioviana.
lamarckiana mut. delata de Vries, Z. Bot.
Oenothera
Bot. Ges.
Oenothera
lamarckiana mut. deserens de Vries,
16: 262. 1916. = 0. glazioviana.
Oenothera
lamarckiana mut. detruncata de Vries, Z. Indukt. Abstammungs-Vererbungsl.
59: 127. 1929. = 0. glazioviana.
Oenothera
Z. Indukt. Abstammungs-Vererbungsl.
lamarckiana mut. diluta de Vries & Boedijn,
Genetics
8: 235.
1923. = 0.
glazioviana.
Oenothera
lamarckiana mut. distans de Vries, Z. Bot.
Oenothera
lamarckiana mut. elongata
Oenothera
15: 389. 1923. = 0. glazioviana.
de Vries, Z. Bot.
15: 394. 1923. = 0. glazioviana.
lamarckiana mut. erythrina de Vries, Ber. Deutsch.
Bot. Ges.
37: 70. 1919. =
0. glazioviana.
Oenothera
lamarckiana mut. favilla
de Vries & Boedijn,
Genetics
8: 235.
1923. = 0.
glazioviana.
Oenothera
lamarckiana mut. flava de Vries, Z. Bot.
Oenothera
lamarckiana mut. fragilis
15: 403. 1923. = 0. glazioviana.
de Vries & Boedijn,
Genetics
8: 235.
1923. = 0.
glazioviana.
Oenothera
lamarckiana mut. ingeminans de Vries, Z. Indukt. Abstammungs-Vererbungsl.
52: 136. 1929. = 0. glazioviana.
Oenothera
lamarckiana mut.
= 0. glazioviana.
Oenothera
lamarckiana mut.
lactuca de Vries, Bot. Gaz.
(Crawfordsville)
linearis de Vries, Ber. Deutsch.
62: 266. 1916.
Bot. Ges. 37: 69. 1919. = 0.
glazioviana.
Oenothera
lamarckiana mut. metallica
de Vries, Ber. Deutsch.
Bot. Ges.
37: 71. 1919. =
0. glazioviana.
Oenothera
lamarckiana mut. nitens de Vries, Z. Bot.
15: 384. 1923. = 0. glazioviana.
SYSTEMATICBOTANYMONOGRAPHS
148
VOLUME 50
Oenothera
lamarckiana mut. opaca de Vries, Z. Indukt. Abstammungs-Vererbungsl.
133. 1929. = 0. glazioviana.
52:
Oenothera
lamarckiana mut. pallida de Vries, Z. Indukt. Abstammungs-Vererbungsl.
178. 1925. = 0. glazioviana.
38:
Oenothera
lamarckiana mut. perennis
de Vries, Flora 116: 336. 1923. = 0. glazioviana.
Oenothera
lamarckiana mut. persicaria de Vries, Z. Indukt. Abstammungs-Vererbungsl.
52: 130. 1929. = 0. glazioviana.
Oenothera
lamarckiana mut. planifolia de Vries, Z. Indukt. Abstammungs-Vererbungsl.
52: 135. 1929. = 0. glazioviana.
Oenothera
lamarckiana mut. proxima de Vries, Z. Indukt. Abstammungs-Vererbungsl.
125. 1929. = 0. glazioviana.
Qenothera
lamarckiana mut. pustulata de Vries, Z. Indukt. Abstammungs-Vererbungsl.
52: 136. 1929. = 0. glazioviana.
Oenothera
lamarckiana mut. retardata de Vries, Z. Indukt. Abstammungs-Vererbungsl.
52: 125. 1929. = 0. glazioviana.
Oenothera
lamarckiana mut. secunda de Vries, Ber. Deutsch
52:
Bot. Ges. 37: 71. 1919. = 0.
glazioviana.
Oenothera
lamarckiana mut. semigigas
1918. = 0. glazioviana.
de Vries, Bot. Gaz.
Oenothera
lamarckiana mut. stenophylla
52: 124. 1929. = 0. glazioviana.
de Vries, Z. Indukt. Abstammungs-Vererbungsl.
Oenothera
Oenothera
Oenothera
lamarckiana mut.
65: 387.
tardescens de Vries, Z. Bot. 17: 198. 1924. = 0. glazioviana.
lamarckiana mut. truncata de Vries,
52: 127. 1929. = 0. glazioviana.
de Vries, Mutationstheorie
oxypetala
(Crawfordsville)
Z. Indukt. Abstammungs-Vererbungsl.
1: 153. 1901, nomen nudum. = 0. glazio
viana.
Oenothera
pohliana
de Vries, Mutationstheorie
2: 435. 1903, nomen nudum. = 0. glazio
viana.
Oenothera
spathulata
de Vries, Mutationstheorie
1: 156. 1901, nomen
nudum.
glazioviana.
Oenothera
suaveolens mut. apetala de Vries, Genetics
3: 4. 1918. = 0. biennis.
= 0.
OENOTHERA
1997
149
Oenothera
suaveolens mut. fastigiata
de Vries, Genetics
3: 10. 1918. = 0. biennis.
Oenothera
suaveolens mut. jaculatrix
de Vries, Genetics
3: 13. 1918. = 0. biennis.
Oenothera
suaveolens mut.
lata de Vries, Genetics
Oenothera
suaveolens mut.
lutescens de Vries, Genetics
Oenothera
suaveolens mut. sulphurea de Vries, Genetics
Oenothera
subrobusta
de Vries, Gruppenweise
3: 5. 1918. = 0. biennis.
3: 9. 1918. = 0. biennis.
3: 8. 1918. = 0. biennis.
Artbildung
192. 1913, nomen nudum. =
0. glazioviana.
Names Not Validly Published
(Based on Experimental
Hybrids)
- 0.
Renner
The names listed below were applied by Renner to experimental hybrids. Although
he used binomials, it is clear that he did not intend to publish new taxa, but was merely
using Latin designations to provide names for ease of reference to his experimental strains
and a means to discuss them. We include them here for completeness of our revision, so
that the status of every name in the very extensive
literature on subsect. Oenothera
is an
alyzed. The names were not validly published, because Renner did not fulfill the require
ments of the ICBN in one or more of the following ways: they were not provided with de
scriptions (thus are nomina nuda), Latin diagnoses [after 1934; ICBN (1994) Art. 36.1],
or designation of types [after 1957; ICBN (1994) Art. 37.1].
Oenothera
Oenothera
albata Hoeppner & Renner, Bot. Abh.
subsp. hookeri (as 0. hookeri).
15: 61. 1929. = 0. biennis x 0. elata
albicurva Renner, Flora 131: 187. 1937. = 0. biennis x 0. oakesiana
(as 0.
muricata).
albiflexa Renner, Flora 131: 224. 1937. = 0. biennis x 0. parviflora
Oenothera
(as 0.
atrovirens).
Oenothera
Oenothera
albifranciscana Renner, Flora 136: 154. 1942. = 0. biennis x 0. elata subsp.
hookeri (as 0. franciscana).
albihookeri Renner, Flora 136: 156. 1942. = 0. biennis x 0. elata subsp. hook
eri (as 0. hookeri).
Oenothera
albilaeta Renner, Flora 131: 223. 1937. = 0. biennis x 0. glazioviana
(as 0.
lamarckiana).
Oenothera
Oenothera
xalbisubcurva
0. silesiaca).
Renner, Flora 136: 326. 1942. = 0. biennis x 0. parviflora
albiundata Renner, Flora 131: 199. 1937. = 0. biennis x 0. villosa
losa (as 0. bauri).
(as
subsp. vil
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
150
Renner, Flora 131: 223. 1934. = 0. biennis x 0. glazioviana
albivelutina
Oenothera
0.
auctirubata
Oenothera
curvilaeta Renner, Flora 136: 142. 1942. = 0. oakesiana
(as 0.
glazioviana
excelsihookeri
Oenothera
(as 0. muricata)
x 0.
lamarckiana).
Renner, Flora 136: 165. 1942. = 0. biennis
sis) x 0. elata subsp. hookeri
Oenotheraflavicurva
oakesiana
x 0. biennis.
Renner, Planta 47: 221. 1956. = 0. parviflora
Oenothera
Oenothera
(as
lamarckiana).
(as 0. chicaginen
(as 0. hookeri).
Renner, Flora 127: 217. 1938. = 0. biennis
(as 0. suaveolens)
x 0.
(as 0. muricata).
flavihookeri Renner, Flora 136: 166. 1942. = 0. biennis
0. elata subsp. hookeri (as 0. hookeri).
Renner, Flora 136: 144. 1942. = 0. biennis
Oenotheraflavirubata
x
(as 0. suaveolens)
(as 0. suaveolens)
x 0.
(as 0.
x 0.
biennis.
Renner, Flora 131: 192. 1937. = 0. biennis
Oenotheraflavitincta
biennis
rubricaulis)
(as 0. suaveolens).
Renner, Flora 136: 151. 1942. = 0. biennis
Oenotherafiavivelutina
0. glazioviana
(as 0.
Renner & Sensenhauer, Z. Indukt. Abstammungs-Vererbungsl.
=
576. 1942.
0. parviflora (as 0. atrovirens) x 0. biennis.
80:
Oenothera flexirubata
Oenothera
laetiflava Renner, Flora 136: 142. 1942. = 0. glazioviana
x 0. biennis
Oenothera
Oenothera
(as 0.
lamarckiana)
(as 0. suaveolens).
laetihookeri Renner, Flora 136: 167. 1942. = 0. glazioviana
iana) x 0. elata subsp. hookeri
Oenothera
x
(as 0. suaveolens)
lamarckiana).
laxiflava Renner, Ber. Deutsch.
villosa (as 0. bauri) x 0. biennis
laxirubata Renner,
(as 0.
lamarck
(as 0. hookeri).
Bot. Ges.
63: 135. 1950. = 0. villosa
subsp.
(as 0. suaveolens).
Planta 47: 236.
1956. = 0. villosa
subsp. villosa
(as 0.
bauri) x 0. biennis.
Oenothera
Oenothera
80:
pictiflava Renner & Sensenhauer, Z. Indukt. Abstammungs-Vererbungsl.
576. 1942. = 0. parviflora (as 0. atrovirens) x 0. biennis (as 0. suaveolens).
pictilaeta
Renner, Flora 136: 143. 1942. = 0. parviflora
0. glazioviana
(as 0.
lamarckiana).
(as 0. atrovirens)
x
1997
OENOTHERA
Oenothera
pictirubata
151
Renner, Flora 133: 218. 1939. = 0. parviflora
(as 0. atrovirens)
x
0. biennis.
Oenothera
Renner, Z. Indukt. Abstammungs-Vererbungsl.
pictivelutina
parviflora
(as 0. atrovirens)
x 0. glazioviana
(as 0.
rigilaeta Renner, Flora 136: 143. 1942. = 0. oakesiana
Oenothera
(as 0.
glazioviana
74: 94. 1937. = 0.
lamarckiana).
(as 0. muricata)
x 0.
lamarckiana).
Oenothera
rubiaucta Renner, Flora 131: 223. 1937. = 0. parviflora
x 0. biennis.
Oenothera
rubiflava Renner, Flora 135: 211. 1941. = 0. biennis x 0. biennis
(as 0. suave
olens).
rubipercurva Renner, Flora 136: 145. 1942. = 0. biennis x 0. oakesiana
Oenothera
(as
0. ammophila).
Oenothera
rubipicta Renner, Flora 131: 223. 1937. = 0. parviflora
biennis.
rubiplana Renner, Ber. Deutsch.
Oenothera
glazioviana
Bot. Ges.
(as 0. atrovirens) x 0.
60: 460. 1942. = 0. biennis x 0.
(as 0. coronifera).
rubirigida Renner, Flora 131: 187. 1937. = 0. oakesiana
Oenothera
(as 0. muricata)
x 0.
(as 0. rubricaulis)
x 0.
biennis.
rubiundata Renner, Planta 47: 237. 1956. = 0. biennis
Oenothera
villosa
subsp. villosa
(as 0. hungarica).
rubivelutina Renner, Flora 136: 146. 1942. = 0. biennis x 0. glazioviana
Oenothera
0.
Oenothera
(as
lamarckiana).
subpictirubata
Renner, Flora 136: 327. 1942. = 0. parviflora
(as 0. silesiaca)
x 0. biennis.
Oenothera
tinctiundata Renner, Flora 131: 205. 1937. = 0. biennis
0. villosa
Oenothera
subsp. villosa
veluticurva
rubricaulis)
x
(as 0.
rubricaulis)
x
(as 0. bauri)] x 0. biennis
(as 0.
rubricaulis).
Renner, Flora 136: 152. 1942. = 0. glazioviana
iana) x 0. oakesiana
Oenothera
(as 0.
(as 0. bauri).
undirubata Renner, Flora 134: 155. 1940. = [0. biennis
0. villosa
Oenothera
subsp. villosa
(as 0.
lamarck
(as 0. muricata).
velutiflava Renner, Flora 136: 151. 1942. = 0. glazioviana
x 0. biennis (as 0. suaveolens).
(as 0.
lamarckiana)
SYSTEMATICBOTANYMONOGRAPHS
152
VOLUME 50
velutiflexa Renner, Flora 136: 152. 1942. = 0. glazioviana
Oenothera
x 0. parviflora
Oenothera
Renner, Ber. Deutsch.
velutirubata
(as 0.
(as 0.
lamarckiana)
(as 0. atrovirens).
Bot. Ges. 60: 461. 1942. = 0. glazioviana
lamarckiana) x 0. biennis.
Names Not Validly Published
These binomials,
- J. P. Lotsy
order, were not explicitly
listed in alphabetical
applied to species or
hybrids but to "nucleus chimeras." They are all based on artificial hybrids. As
it seems quite obvious
of Renner,
of reference
to these entities, and that he did not intend to use these Latin names formally
in the taxonomic
Oenothera
system.
xbiennoides
xbiennivelutina
xcoerulea
muricata
Oenothera
Oenothera
x (0. biennis x 0.
lamarckiana x 0. xfallax).
(as 0.
0. biennis x 0.
Oenothera
1: 36. 1919. = 0. glazioviana
Lotsy, Genetica
glazioviana)
Oenothera
in the case
that Lotsy merely was supplying Latin formulas for ease
1: 23. 1919. = 0. biennis x 0. glazioviana
Lotsy, Genetica
(as
lamarckiana).
1: 46. 1919. = 0. oakesiana
Lotsy, Genetica
x 0. oakesiana
(as 0.
x 0. muricata).
xepilobioides
Lotsy, Genetica
1: 47. 1919. = (0. oakesiana
x 0. oakesiana
(as [0. muricata
x 0.
x 0. glazioviana)
x 0. muricata).
1: 41. 1919. = 0. biennis x 0. glazioviana
Lotsy, Genetica
xfalloides
lamarckiana]
(as mu
tant of 0. xfallax).
xintermedia
Oenothera
Oenothera
1: 44. 1919. = (0. oakesiana
Lotsy, Genetica
(0. biennis x 0. glazioviana)
xlanceolata
(as [0. muricata
x 0.
x 0. glazioviana)
lamarckiana]
1: 41. 1919. = 0. biennis x 0. glazioviana
Lotsy, Genetica
x
x 0. xfallax).
(as mu
tant of 0. xfallax).
Oenothera
xlinearis
ennis x 0.
Oenothera
Lotsy, Genetica
xmuricatoides
0. oakesiana
Oenothera
Oenothera
Lotsy, Genetica
(as [0. biennis x 0.
xmurilaeta
0. muricata
1: 24. 1919. = 0. biennis x 0. glazioviana
(as 0. bi
lamarckiana).
Lotsy, Genetica
x 0.
xmurivelutina
(as 0. muricata
1: 48. 1919. = (0. biennis x 0. glazioviana)
lamarckiana]
x
x 0. muricata).
1: 25. 1919. = 0. oakesiana
x 0. glazioviana
(as
lamarckiana).
Lotsy, Genetica
x 0.
lamarckiana).
1: 26. 1919. = 0. oakesiana
x 0. glazioviana
OENOTHERA
1997
1: 20. 1919. = 0. biennis x 0. glazioviana
xnova Lotsy, Genetica
Oenothera
153
(as mutant
of 0. xfallax).
Oenothera xsubfalloides Lotsy, Genetica
mutant of 0. xfallax).
Oenothera
1: 41. 1919. = 0. biennis x 0. glazioviana
(as
xsublamarckiana
Lotsy, Genetica 1: 42. 1919. = (0. biennis x 0. glazioviana)
x (0. biennis x 0. glazioviana)
(as 0. xfallax x [0. biennis x 0. lamarckiana]).
ACKNOWLEDGMENTS
We
grants
gratefully
and Catherine
the Missouri
We
number
Botanical
T. MacArthur
appreciate
for support
Foundation
for field work
was
Funding
and in the preparation
ica) in 1981. Thanks
also to Chris Sherman
tion of the manuscript,
review
helpful
C. Hoch
comments
made
the entire manuscript,
who
Sisson
for help with
by Harlan
including
and considerable
of the manuscript
from 1983
of the illustrations
from living plants
also extend
von Berg
garden), G. Linne
bers (0. wolfii
logs of 1823),
on 0. wolfii),
nese
labels), L. Mencke
tion of 0. biennis), W. Rogmann
tions), W. Stubbe
Chinese
and W. A. Weber
(search
for Cockerell
for preparation
thus avoiding
to borrow
Botanical
ing visits
We
ALTA,
Garden
the need
for providing
from 1981
also
AMD,
through
work
names
of specimens.
space and kind assistance,
We
(assistance
with
U. Kleingarn
experimental
(assistance
(translation
J. Reveal
determina
W. Tai (translation
0. Wasmund
assistance
N.
(typifica
(cytological
Dietrich),
in
of Chi
determinations),
manuscript),
(helpful
of
discus
from the staff at A, GH,
after annotation
are grateful
var.
seed cata
D. K. Imper (information
E. Schumacher
especially
K. Cham
and Vienna
and cytological
appreciate
of the
experimental
of 0. biennis
labels), K. Mading
toWerner
label information
for the
aspects
of 0. magdalena),
(Dresden
and specimens),
We
specimens).
of specimen
large numbers
garden),
facilities
of
of the
by Werner
Dietrich
in
to the staff of theMissouri
to the herbarium
and library staff dur
1994.
thank the staff of
ARAN,
working
the course
with
(typification
J. Haubenreich
garden
to Peter
review
and S. Lechner
(assistance
(typification
(proof-reading
experimental
a critical
on specific
for assistance
specimen
the very
indebted
at DUsseldorf.
R. Jean (0. suzukiana),
S. Rehm
(Desfontaines
MT, NA, NY, PH, and US
1981,
names),
and com
appreciate
during
to R. Lechner
J. Flanagan
in the experimental
and providing
to mention
(type of 0. depressa),
of Japanese
labels),
with
(assistance
labels), K. Vetschera
specimen
sions),
specimen
(helpful discussion
from 1824),
of Linnaean
(translations
(assistance
of Japanese
(translation
labels),
B. Hellenthal
of 0. cruciata),
garden), H. Koyama
specimen
specimen
we
to St. Louis,
types), A. Cronquist
(Gates
list of Hamburg
(seed
(typification
C. E. Jarvis (typification
the experimental
Murakami
of Russian
(translation
D. Boufford
determinations),
the final ver
for reviewing
visits
F. Baum
names),
in Oenothera
in preparing
are particularly
garden
of people
locality
for the names
We
are grateful
to a number
Spanish
A. 0. Chater
information),
I. Friederichsen
I. C. Hedge
garden),
(cytological
population
W. Daugsch
canescens),
with
(assistance
the prepara
are ours. Likewise
in the experimental
our thanks and appreciation
P. Acevedo
including
1996. We
through
for publica
with
assistance
provided
too numerous
of the large
(North Amer
and illustrations
to Dan Nicolson
for the many
in various ways
help
preparation
We
Shannon
are grateful
logistics
maps
of a database
creation
on the entire manuscript.
Lewis
ranges of the species
for considerable
any errors remaining
however,
preparation
project
Fund at the Smithson
in the handling
Rudolph
completed
of the superb help Robynn
in diverse ways,
for assistance
and Betsy
and Michael
and the index. We
1-4,
figures
on all of the nomenclature;
menting
and a support grant to
Opportunity
for the indigenous
and Peter Hoch,
Gowing
are appreciative
sion of the manuscript,
to Peter Raven
by the Research
Arey
of the maps
Arey, Gloria Hoch,
and to Donald
We
subsect. Oenothera.
and to the John D.
recently BSR-8906848,
through a fellowship
from Susan Mill
the fine assistance
to Susan Mill
PI), most
also provided
through a series of
for support of this work
Foundation
in 1991.
of specimens
tion. Thanks
Science
(Peter H. Raven,
Garden
Garden.
Botanical
ian Institution
the National
acknowledge
to the Missouri
ARIZ,
the following
ASC,
CGE, CHR, CM, COI, COLO,
ASU,
CORD,
herbaria
CS, CTES,
FM, FR, FSU, G, G-BOIS,
G-DC,
IND, ISC, ISL, ISU, JACA,
JEPS, K, KANU,
CU, DAO,
GA, GH, GOET,
LL, LP, LY, LZ, M, MA, MAF, MAK,
for making
their material
B, BC, BCF, BH, BM, BOL,
MASS,
GRA,
KR, KRAM,
DS, DUKE,
GZU,
MEL, MERL,
DUSS,
H, HAL,
KTU, KUN,
MICH,
available
BP, BR, BREM,
E, ENCB,
HB, HBG,
KYO,
for this study: A,
BRY, C, CAN,
HGAS,
HSC,
L, LA, LAM, LD, LILLE,
MIL, MIN,
CAS,
F, FI, FI-W, FLAS,
MJG, MNA,
ID, ILL,
LISU,
MO, MONTU,
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
154
MSC,
MUB,
MT, MU,
NSMC,
NSW,
POM,
NU,
NA,
MUC,
NY, 0, OKL,
NBG,
NCSC,
OKLA,
PR, PRC, PRE, RM, RSA,
ORE,
S, SAM,
NCU,
OSC,
ND,
SD, SEV, SIU,
TEX, TI, TO, TRT, TTC, UAC,
UBC,
UWM,
V, VDB,
WIS, WRSL,
UWSP,
are most
VPI, W, WH, WILLU,
to Prof. Dr. K. Rostanski
grateful
nis from the following
LOD,
LW, MHA,
Szczecin
who
WTU,
J.Madalski
MW,
0, POZ, PR, RIG, RV, Zemaljski
(Poland),
TOR, TUR, WA, WI, WRSL,
Mus.
SRSC,
(Yugoslavia),
STE,
US, UT, UWL,
YU, Z. We
Chrzan6w
bien
(Poland),
(Poland), H, HAL,
Poland),
NO,
PH, PO,
records of 0.
specimen
Gdanisk
(Wroclaw,
Sarajevo
WRSP,
SP, SRGH,
BIL, Herb. Mus.
Pharm. Bot. Acad. Med.
(Gij6n, Spain), Herb.
NMC,
WU, WVA, WWB,
has provided
(Poland),
NH,
PE, PENN,
UNLV, UNM,
UNCC,
WS, WTS,
Poland),
NESH,
PAV, PDA,
SMY, SOM,
SMU,
UMO,
Stat. Bialowieza
(Russia), GB, Hertb.
KW, Herb. M. Lafnz
Acad.
(Katowice,
herbaria: Herb. Geobot.
CL, Herb. Bot. Gard. Donetsk
KTC, KTU,
UC, UMN,
NEBC,
PAC,
P, P-LA,
SMS,
TAES, TENN,
NEB,
NDG,
OSH,
KRAM,
LATV, LBL, LE, LGU,
Herb.
Bot. Dept.
Agric.
Botanic
Garden
at Kew.
ZA.
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Phytochemistry
APPENDIX
SPECIMENS EXAMINED
For each species we divided
our cytological
including
vouchers
and cultivation
number);
of the species;
and 4) historic
diakinesis
the specimens
for each taxon and give
ically at the end of each description
The
cultivation
or material
collections
numbers
examined;
the number
in the section,
2) the year of cultivation;
and 3) the cultivation
the progeny
from one capsule
and most
even
national
seed exchange
herbarium
akinesis
acronym
of botanical
in most
gardens).
examined
from cultivated
plants,"
that would
Two
number
cytolog
additional
a binomial
the strain was
is given
obtained
in the 0. grandiflora
or from more
individuals)
of
be
strains of E.
than one capsule,
from seeds obtained
items may
through
or
the inter
included
in parentheses
after the
for this project
are deposited:
1) the di
for: the name under which
the taxonomy
consist
of Dtissel
Institute, University
for that year. The strains cultivated may
(as is the case
of the strains studied
apply using
the name under which
(Botanical
of the strains cultivated
the vouchers
and 2) occasionally
the name
the known
examined
and other
by colleagues
(as is the case
indicating where
configuration;
in the literature;
case of strains,
of the seeds
of the strains obtained
from several plants
the natural range
summarize
and populations
acronym DUSS
represent
Steiner
from outside
gardens. We
plants,
information
of individuals
"specimens
1) the institutional
dorf, Germany);
either
after the collection
3) specimens
in botanical
cultivated
from cultivated
examined
in parenthesis
in the section Taxonomy.
listed
by hyphens:
1) specimens
are given
or all material
2) representative
configurations
three parts separated
into four groups:
(the configurations
of European
the strain was
Oenothera
from the botanical
garden
treated
researchers;
or in the
seed exchange
or used
in that index seminum.
la. Oenothera
elata
subsp. elata.
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. Costa
inas, cult.
DUSS-83-072
from
seeds
from Pohl
s.n., cult.DUSS-81-698
Rica.
CARTAGO (9050'N,
(MO); Moravia,
cathedral
83052'W):
yard, Pohl
Las Ru
14069,
cult.
(MO)
San Jose (9059'N, 84?04'W), Avenida Central, Pohl 14075, cult. DUSS-83-070
(MO)
(711);
San
de
Pablo,
Cant6n
Dota, Pohl 14108, cult. DUSS-83-03
(MO) (711); San Pedro, NE telephone building
(711);
Los Yoses, Pohl 14078, cult. DUSS-83-071
(MO)
Cartago, Tierra Blanca, Pohl 14123, cult. DUSS-83-02
(7,I);
(MO)
Monasterio
antiguo, 1967, Mittak
s.n., cult. DUSS-76-09
(MO)
Mexico.
HI
(711). Guatemala.
(711).
DALGO: Zimapan,
cult. DUSS-76-07
(MO) (7II).-MICHOACAN:
Morelia
(19040'N,
1977, Re
101011'W),
hwinkel s.n., cult. DUSS-78-070,
DUSS-78-071
(MO) (7II).-QUERtTARO:
Pifias, at rd 120 between Tamazun
chale and Vizarr6n,
1985, LJischper
s.n., cult. DUSS-86-2275
REPRESENTATIVE SPECIMENS. Costa
Colegio
La Granja
(10?02'N,
84?27'W),
Rica.
Weston
Ruinas
(MO)
de Cartago
et al. 3004
(711).
(9?52'N,
83055'W),
(DS); San Jose, Khan
Huhn
et al. 1018
115 (MO); near
(MO). El Salvador.
160
SYSTEMATICBOTANYMONOGRAPHS
San Salvador,
21823
del Aguila
(13?54'N,
(15?07'N,
90?05'W),
Contreras
Chilasco
huetenango
(15?20'N,
can San Pedro
Guanajuato
(14?39'N,
Zimapan
(20?45'N,
POM), Munz
98?54'W),
15309
POM, UC), Munz
15307
IND, NY,
from Munz
(BH, GH, NY, POM, US),
15067
Munz
15304
(BH,
15070: Munz
15242
(BH, IND, POM), Munz
& Ugent
2090
(F, MO).
Alto
(WIS).-VERACRUZ:
Panama.
5858
66172
F,MO);
(ENCB,
Robert
(MO); Nueva
cult.
from Munz
15070
Munz
Quiel-Boquete,
194
Beliz
et al. 692
(MO); Cerro
& Amable
15067: Munz
3585
15255
(18?54'N,
98?26'W),
96?55'W),
Calzada
Punta
(8?51'N,
& Todzia 2037
Schmalzel
& Schmalzel
(F,
from Munz
cult.
(MO); Cerro
Punta, Robert
(F,
City
(GH, POM, RSA, US),
(19?32'N,
above Cerro Campana,
1926
from Mexico
(BH, POM, UC),
de las Palmas
(19?29'N,
15253
Munz
Arsene
(BH, IND, NY, POM, US); Atlixco
Guadalupe,
& Schmalzel
15254
98?12'W),
Jardin Paseo
a Alto
Suiza, Hamilton
15050: Munz
(MO);
(BH, GH,
Texcoco
City, hwy
(19003'N,
15886
15252
99?39'W):
aereo near Mexico
Puebla
15310
Jalapa,
CHIRIQUi: Camino
above Boquete,
Quiel,
Croat
1293
A. Gentry
82?34'W),
Puerto
Acatzingo,
Breedlove
15048: Munz
(19?18'N,
(14?28'N,
45 (WIS); Vol
GUANAJUATO: cult. from
del Chico,
2 km S of Chapingo,
POM, US, WTU),
IND, NY, POM, US);
US). Mexico.
from Toluca
cult. from Munz
(GH, NY,
POM), Munz
Ugent
15075
Mathewson
91010'W),
Mineral
POM, US, WTU).-MExico:
cult.
near Hue
de Agua
Volcan
cult. from Munz
(GH, NY, POM),
& Montalvo
BAJA VERAPAZ:
(LL).-HUEHUETENANGO:
(14?44'N,
(ENCB, MICH,
IND, NY, POM, US).-PUEBLA:
Texmelucan,
10955
(GH).-HIDALGO:
(BH, IND, POM, US, WTU);
15050
(BH, DS,
(MO, US);
15048
Munz
(ENCB);
15308
to Puebla, Munz
Munz
(BH,
550
et al. 26654
Molina
88'20'W),
(GH, US).-SACATEPEQUEZ:
Panajachel
302
Dugas
(13?48'N,
(RSA, UC, US). Guatemala.
(MO), Contreras
3672
Molina
101?15'W),
99?21'W),
Garcia
10954
s.n. (RSA).-SOLOLA:
91?16'W),
(21?01'N,
of El Boquer6n
Tucker 1268
89?42'W),
Nelson
91?28'W),
1905, Kellennan
90?45'W),
NE
El Picacho
San Salvador,
Volcan
(NY); Cerro
VOLUME 50
(MO);
1682
(MO),
(MO).
SPECIMEN CULTIVATED IN BOTANICAL GARDEN. Germany.
Frankfurt,
1832
(FR).
lb. Oenothera elata subsp.hirsutisissima.
SPECIMENS FROM CULTIVATED PLANTS. Mexico.
1875 m, Moran
25118,
24474,
cult. DUSS-82-0337
cult. DUSS-82-0338
Valle de Las Palmas,
(711);
San Jose (32?33'N,
116?33'W)
(MO) (7II); Rancho
cult. DUSS-82-0336
(MO) (7II); Rio San Vicente
cult. DUSS-82-0342,
82-0343
Moran
26372,
2, Spellenberg
& Soreng
rd to Little Morman
13260,
(MO)
(MO) (7II; 04
(2 04
Co., Forestdale
and 3II). Coconino
Co.,
18389,
Alpine
Co.,
cult. DUSS-77-070
s.n., cult. DUSS-76-035
(MO)
DUSS-76-034
(MO) (04
DUSS-77-072
(MO)
(04
ca. 2400 m,
ca. 1730 m,
Lake,
(MO) (7II; 06
60, ca. 2000 m, Pinkava
Park,
in front of Natl.
(MO) (7II( 04
house
and SII); Tuttle Creek
and 5II). Mono
1146, cult. DUSS-76-029
Riverside
(MO)
Cardiff,
strong
04
Co.,
(7II; 04
Pine,
and SII); 4 mi
1975, Armstrong
Co., Sacramento
River, Dog
River
Island,
Hot Creek
along Hwy
S Temecula,
(MO)
ca. 200 m,
S Red Bluff,
ca. 80 m, Devine
395,
04
(MO)
(7II); San Diego,
0461,
van Wagtendonk
(MO)
Co., Yosemite
National
airport, ca. 2300 m, Stock
Taylor 5205,
cult. DUSS-87-2024
(MO)
and 5II).
DUSS-76-037
(7II). San Diego
ca. 240 m,
cult. DUSS-76-036
(MO)
(MO).
(MO) (7II; 04
s.n., cult. DUSS-76-038,
and SII); Escondido,
(MO)
and
(7II; 04
s.n., cult. DUSS-76-093
area, E Mammoth
cult. DUSS-79-0527
cult.
(MO)
1144, cult. DUSS-76
and 5II). Lassen Co., S end of
(7II; 04
1505,
s.n., cult.
18387,
s.n., cult. DUSS-76-033
(7II). Mariposa
ca. 3 m, Wedberg
(2 04
1975, Trowbridge
ca. 2500 m, Thomas
(MO)
s.n., cult. DUSS-76-041
cult. DUSS-79-0528
(MO)
4, ca. 2300 m,
1969, Stubbe
s.n., cult. DUSS-76-043
(7II; 04
(MO) (7II;
and 2II). Navajo
along Hwy
near Bishop,
cult. DUSS-77-073
Munz
cult. DUSS
s.n., cult. DUSS-76-065
cult. DUSS-76-068
Pass
(MO)
1975, Armstrong
Canyon,
4524,
1143, cult. DUSS-76-030
ca. 1700 m, Stockhouse
picnic
750902-1,
1975, Armstrong
0357,
250 m,
116046'W),
Co., Ramsey
and 3II; 06,
1969, Stubbe
Co., Pt. Cabrillo,
Clarke
ca. 80 m, Devine
Ranch
1975,
70, 1975, Reveal
s.n., cult. DUSS-76-040
s.n., cult. DUSS-76-039
and SII); San Marcos,
Building,
(MO) (7II).Monterey
3 mi N of Rainbow
and 4II; 08
ca. 18 mi NW Bishop,
near Lone
Administration
along Hwy
(32?31'N,
Theroux
et al. 18992,
s.n., cult. DUSS-77-069
Co., Sierra Nevada,
Plumas Co., Feather River Canyon
27857,
and 4II); Grover Hot Spring near Markleeville,
and SII); Rock Creek Canyon,
Park Service
21124,
15 m, Moran
s.n., cult. DUSS-76-066
ca. 10.4 mi E of Ebbetts
and SII). Inyo Co., Hotel
1976, Schlising
1975,
1975, Purvis
3 mi NE of Kennedy
(7II); Kennedy Meadows,
Peak, Stockhouse
5II); Sierra Nevada,
along S Fork Kern River, NE Kennedy Peak,
031 (MO) (7II; 04 and 5II), Stockhouse
1145, cult. DUSS-76-032
Eagle
116?26'W),
475 m, Moran
650 m, Moran
ca. 1800 m, Reeves
Monument,
Flagstaff,
(MO) (06
Station
and SII). Cochise
(7II; 04
10 mi N of Sedona,
Sierra Nevada,
(7II; 04
(Meling),
4 km SE Erendira,
116?16'W),
National
s.n., cult. DUSS-76-064
Post along Hwy
Trading
and 311).-CALIFORNIA:
(MO)
(7II); Chiricahua
and SII); Oak Creek Canyon,
(31038'N,
Caniada Cesma,
(MO) (7II).-CHIHUAHUA:
N end of San Luis Mtns on Hwy
(MO). U.S.A. ARIZONA: Apache Co., jct. AZ Hwy 260 with
cult. DUSS-87-2021
(MO)
and SII), 1975, Theroux
Thomas
DUSS-82-0341
cult. DUSS-76-067
Lake,
cult. DUSS-76-084
82-0326
04
6853,
(31?19'N,
and 4II); 1 km E Valle Redondo
(MO) (06
cult. DUSS-82-0340,
BAJA CALIFORNIA: La Grulla
(MO)
(7II; 04
(MO) (08
(71d).-COLORADO:
Co.,
1975, Arn
(MO)
(7II;
and
Tehama
%II).
and 3II); Sacramento
Chaffee
Co., Raven
OENOTHERA
1997
cult. DUSS-77-0145
26550,
cult. DUSS-82-0330
051
(7II).-NEVADA:
and 51).-NEW
MExIco:
(MO) (711;04
(7,I). Otero Co., Apache
76-073
Summit
Co., S Bonita Lake, Northington
1085, cult. DUSS-76-072
70, 12 mi SE Ruidoso,
1975, Zimmernann
s.n., cult. DUSS
Lincoln
along Hwy
1975, Halley
(711). Sandoval Co., Alamo Canyon, Bandelier Natl. Monument,
and 5II). San Miguel Co., Montezuma,
1975, Sch. s.n., cult. DUSS-77-0214
(MO)
(MO)
and SII)- San Juan Co., Animas River near Needleton, Wagner 4532,
Washoe
Co., Carson Valley, 1975, Robertson
s.n., cult. DUSS-76
(MO)
(MO)
070
04
(711;
(MO)
161
(04
s.n., cult. DUSS-76
(MO) (7II). Socorro
Baker Co.,
(MO) (7II).-OREGON:
Legal-Sec.
2, T75, R47E, North Pine Campground,
1975, Strickler s.n., cult. DUSS-76-015
(MO) (711). Grant
Co., Blue Mtns, Ind. Sem. bot. Gard. Vancouver 1972 no. 121, cult. DUSS-76-013,
DUSS-76-014
(MO) (711).
Co., Magdalena
Mtns, Water
TEXAS: Anderson
Co.,
(MO)
along Hwy
Town Creek,
and
Brazos
2,,).
& Intosh s.n., cult. DUSS-82-0504
6, 1978, Catling
645 along Hwy
2.9 mi NE of Hwy
and 3II; 010
and 4II; 08
(06
s.n., cult. DUSS-76-071
1975, Wagner
SW of Palestine,
6500, DUSS-92-2043
bridge
Canyon,
(MO), DUSS-87-2025
ca. 2200 m, Higgins
204 (MO) (711).-UTAH: Kane Co., 4 mi N of Glendale,
10912,
Grove Campground,
1979, Clark & Stenger s.n., cult. DUSS-82-0333
04
Red Butte Canyon,
Mtns,
ca. 2830 m,
1975, Arnow
River
(MO), DUSS-84
cult. DUSS-82-0332
Ponderosa
Wasatch
& Sisson
79, Wagner
17 km NW Navasota
Co.,
(MO);
Salt Lake Co.,
(711).
(MO) (711;05 and 4II; 2
(MO)
s.n., cult. DUSS-76-056
and
Sevier Co., 3.5 mi W of Fremont jct. along Red Creek, Albee 2119, cult. DUSS-76-054
(MO) (04
311).
and 5II); between Clear Creek and Red Creek, 1974, Garrett s.n., cult. DUSS-76-055
(MO) (711;04 and
511);
Wasatch
Plateau, Jack Addelyx Monument,
Albee 3350 p.p., cult. DUSS-82-0386
(MO). Uintah Co., Sweetwa
ter Canyon,
Neese
near Bogart,
6560,
cult. DUSS-82-0334
1975, Hecht
(MO)
s.n., cult. DUSS-76-092
338
(DS, US),
Broder
582
(DS, US);
Valley, Mearns
3768
royo La Grulla
5 km SW of La Grulla
Valle
de las Palmas
(32?19.5'N,
Moran
116?45'W),
26372
NY, POM, RSA,
Garcia
(29?59'N,
445
GH, NY, POM, US),
Pedro, Munz
15042
RANGO: 43.6
mi W
rahui,
Sierra
1 10?18'W),
NE
(DS, GH,
293
Surotato
(F, GH, MO,
(25?53'N,
4187
1244
Breedlove
15726
MIN,
NY, RM). Coconino
140c
(ARIZ, NY, WTU);
(MO). Cochise
(ARIZ, DS, US).
Mtns, McGill
6465
& Lehto L20582
Coronado
mi above Fort Apache,
Mtns,
Rita Mtns,
Clark
12401
Carson
River,
Meadows,
Nevada,
Johnson
149
s.n. (NY, UC, US).
Austin
Mono
445
Creek
(COLO, MT,
1104
(MO, UC).
RM, UC, WS,
El Dorado
WTU).
Lake
Co.,
Pinaleno
S of Alpine,
Deaver
Co., S top
4883
from 10
Pima Co., Spud Ranch,
Sabino Canyon,
Wetzel
Rincon
0.5 mi
Canyon,
Coconino
475
Natl.
Forest,
3475
(DS). Alpine
Co., E of
Station,
(UC). Calaveras
s.n. (UC, US).
Fresno
Inyo Mtns,
Lake Co., Whispering
(A,
1910,
Co., Big
Co., Sierra
rd, Everett & Johnson
Canyon,
S of
Santa
H. S. Gentry
(Wylie's)
19458
rd, jct. with Hot Springs
Smith 905 (RM, UC, WTU).
Toumey
Forest,
lower Madera
ISC,
Oak Creek
(MO). Mohave
Co., Cook's
Inyo Co., Barrel Springs, Mazourka
SD, UC). Kern Co., Kernville,
4291
Mtns,
of Alpine,
F, GH,
cult. from Harrison
Jepson
1913, Brandegee
Natl.
Meadows,
Natl. Mon.,
Bridge,
Amador
del Roble,
Creek, White
(ARIZ, DS,
SE side of Coyote Mtn,
near Oroville,
El Rancho
NA, WS, WTU);
Coronado
Santa Cruz Co.,
Well
Ocu
(30?57'N,
(MO); 4 mi NW
1420
Pinal Co., Upper
E of Niles
Co., Riverton,
Lake-Florence
OSC, RSA).
UC).
(24?02'N,
(MO). Gila Co., Pinal Mtns,
POM, UC, WTU).
Yuma Co.,
Canyon,
3809
Campground,
Co., Montezuma
Butte Co., Feather River
rd toHuntington
(GH, UC);
Park, Blumer
(POM). Navajo
(BH,
Cananea
Co., Bonita
Sec, 28), Fletcher
F, NY, ORE,
ASU).
Co., Niles
899
3790
15039
5 mi E of San
(MO).-SINALOA:
(MO).-SONORA:
2 mi N of Hannegan
s.n. (ARIZ, ENCB).
Yavapai
(ARIZ, ASC,
(NY, POM,
Co.,
ISC, MIN, MO,
1951, Caldwell
Alameda
Canyon
Munz
San Miguel,
1102
(BH, DS, GH, MO,
89A, Wagner
(T12S, R14E,
(BH, CAS,
(GH, OKL).
(DS, LAM, MO, MT, OKLA,
3990
14214
41966
NY).-CALIFORNIA:
Brandegee
Greenlee
& Ramsey
Trail, Ramsey
Munz
Demaree
McGuireville,
NMC,
(ASU, NY).
Mt. Lemmon,
Summerhaven,
on US Hwy
Co., Pima Canyon
Barfoot
NY, POM, RM, US),
(MO); Cd. Durango
ARIZONA: Apache
16974
MO,
(BH, IND, NY, POM, US).-Du
Gonzales
6177
27857
near Colonia
Sierra Madre
NDG,
(32?31'N,
Moran
F, GH, MICH,
103?36'W),
3964
(GH), Hartman
Mtns,
Munz
Co., Swift Trail of Wet
s.n. (ARIZ, DS, GH,
1909, Blumer
887
MO,
E of Luna Lake, Wagner
Co., Chiricahua
Co., Moencopi,
Graham
(ASC, DS). Maricopa
of Blue Mtn,
S. Gentry
(NY, US);
IND, POM, US);
Est. Guadiana,
SD); Ar
S side of
Station
116?20'W),
4111 (CAS, DS,
15312
(CAS); Tecate
(DS, RSA,
(SD); Caniada Cesma,
(25?48'N,
& Brown
U.S.A.
campground
1mi. N of Pine Flat Campground
Canyon,
H.
107?32'W),
12382
14601
(31?17.5'N,
F, MICH,
15042: Munz
NY, UC, US);
(ARIZ, GH, MICH).
Goodding
(BH,
40, Wagner
20 (UC); San Pedro, Hartman
Donnelly
of El Tigre, White
MO,
cult. from Munz
along Hwy
24474
6049
of San Pedro
15311
64 mi SE of Ensenada,
Eastwood
Moran
& Gillespie
107 (ARIZ, DS,
15039: Munz
POM),
of Durango
Wiggins
6 mi W
Park
Co., Boger
1 km E of Valle Redondo
(SD);
Juarez, Nelson
& Barber
cult. from Munz
E. Palmer
104?40'W),
near Colonia
(POM).-COAHUILA:
Moran
4 km NE of Erendira
on rd to Neji Rancho,
Townsend
108?20'W),
25118
116?37'W),
115?36'W),
115?31'W),
Moran
116'38.5W'),
US).-CHIHUAHUA:
& Barber
Townsend
(31?52'N,
(31?03'N,
(30?51.5'N,
(SD); Rfo San Vicente,
15 mi NE of Ojos Negros,
(SD);
Ensenada
San Pedro Martir
(US); Rancho
Whitman
and
511).
BAJA CALIFORNIA: S end of Santa Catarina,
REPRESENTATIVE SPECIMENS. Mexico.
Broder
and 41I).-WASHINGTON:
(06
(MO) (04
7442
Langenheim
Pines Re
SYSTEMATICBOTANYMONOGRAPHS
162
sort, Baker
Angeles
3685
(UC). Lassen
2327a
Co., Arroyo
(ARIZ, CAS,
18, 39-117k
Yosemite
F, GH, LL, MICH,
(LA); Reds Meadow
209
Co., Lee Vining
(DS, UC, US). Mono
ISC, NA, NY, POM, UC, WTU).
Sulphur
Creek,
Springs
Monterey
14492
Jepson
Co., Rancho
WTU).
Placer Co., near Brockway,
& Kennedy
s.n.
Munz
Santa Ana,
(NY, POM, UC), Munz
13997
San Diego
s.n. (GH). San Joaquin Co., Union
man
3814
Santa Clara Co., Guadalupe
from Wolf 6234
from Capitola,
Munz
1913, Smith
s.n. (CAS, GH, MO).
near Mount
Shasta City, Cooe
City, Sacramento
Valley, Wolf 1435
14062
UC), Munz
Ventura Co., Griffins,
Pagosa
Elmer 3819
Hall
2639
Springs,
Baker
SW of Kirkwell,
mi NW
Weber
13973
14104
(POM), Munz
(KANU).
16978
RSA,
& Norby
City, Norby
Co., 32 mi E of Gunnison,
Munz
13915
(MO). Hinsdale
Huerfano
CAS,
Canyon, Mackay
Munz
13015: Munz
14181
973
Saguache,
131-141
River, RR
Silverton,
Wolfe
13014
Teller Co., Divide
on Hwy
NY, ORE,
POM, WTU).
C. L. Hitchcock
26501
3728
(MO). Montezuma
Co., Monte
San Juan Co.,
& Norby
10411
Camas
510
(RSA).-IDAHO:
14546: Munz
14702
Co., Corral Creek,
14551: Munz
Co., Caldwell,
14500
14039
13013
16614
Vista,
422
Co.,
(BH,
Co.,
(COLO).
Munz
13011
cult. from
near the Rio Grande Natl.
15019
(IND).
Wagner
4532
14162
Saguache
along
Co.,
the Animas
(MO); 20 mi
S of
(BH, POM, UC).
7 mi above Boise, Munz
14546
(CAS, GH, NY, POM, UC, US), Munz
15273
14551
GH, NY, POM, US), Munz
1935, Tucker s.n. (ID). Elmore
(BH, CAS, NA, NY, UC, WTU).
Munz
Ramaley
(POM), Munz
Ada Co.,
(POM),
s.n. (GH). Las Animas
3 mi N of Corral, Munz
(BH, CAS,
13904
Butte, Hoch
s.n. (UNM). Huerfano
ca. 3 mi. N of Needleton
vic. of Needleton,
13014: Munz
s.n. (MIL). Gunnison
(BH, F, GH, POM),
Springs, Munz
Co., High
(RM). Fremont Co.,
0.5 mi E of Hesperus,
10 mi W
13369
(BRY, COLO,
13018: Munz
Co., Wasson,
POM, US). Rio Grande
of Steamboat
4624
670
Co.,
2
Ramaley
McGregor
and Crested
13015
5 mi
Co.,
(POM), Munz
Co., Antonito,
7 mi N of Durango,
Co.,
Co.,
Creek,
13921
1903, Finger
1910, Hunnewell
Mineral
San Jac
Archuleta
s.n. (NY). Dolores
1967, Barr
River,
137 (ND).
Valley,
s.n. (MO). Chaffee
Osterhout
cult. from Munz
Creeks,
& Yuba
(NY, POM,
(GH, NA, NY).
(COLO).
of Russell,
& Greene
Ouray Co., Ouray, Munz
p.p., WTU).
cult from Munz
Canyon
& Muhlick
(DS, KANU).
cult. from Munz
24, Norby
cult from Munz
Co., Delta,
Springs,
Co., Estes Park,
(MO); 33 mi. N of Durango,
(BH, IND, NY, POM, US, WTU).
NY, POM, UC),
13993
5343
13021: Munz
(POM); vic. of Gunnison
& Henson
Lake Rd,
Nicolaus
Baca Co., East Carrizo
7 mi NE
1877, McCosh
14082
Larimer
Routt Co.,
(CU, MICH
(BH, GH, NY, POM, UC),
Delta
La Plata Co., Animas
(UNM).
(CAS, NY, ORE,
4512
& Martin
(MO). Conejos
Co.,
Co., Glenwood
jct. Nellie
Mtn, Wagner
(F, POM),
26550
Costilla
(CAS, GH, NY, POM, US),
& Brooks
(NA, NY).
tracks, Wagner
Munz
from Denver,
NY, POM, RM, US, WS).
Forest, Gierisch
Shasta Co., Castella,
Co., Castle
Co., Mather, Mahurin
cult. from Munz
118 (MO, NY).
(POM), Munz
6C-236
Co., E side of Douglas
s.n.
cult.
between
14129
Twissel
Ranch,
Santa Cruz Co.,
(POM).
1913, Cockerell
Co., Boulder,
of Salida, Raven
(RSA). Garfield
F, GH, NY, POM, RSA, UC, US).
(BH, GH, MIN, MO,
1866, Bolander
1958, Pollard
(POM), Munz
Ramaley
13968
(POM), Munz
Siskiyou
s.n. (COLO). El Paso Co., Palmer Lake, Osterhout
of Lake City,
Stephens
13982
NY, POM, RM, US).
UC, WS).
Redfield
13018
14042
6 mi W
Co.,
6.5 mi E of Stonewall,
Moffat
496
Munz
(POM), Munz
13929
s.n. (CAS).
Tuolumne
Co., Alamosa,
Boulder
TEX,
20 mi
River,
plateau, Lone Cone Ranch, Brewster
Canon
Alamosa
2 mi NW
(POM);
(CAS, COLO,
Co., Arkansas
(CAS, F, GH, MIN,
Co., Twisselman
City, C. L. Hitchcock
(BH, GH, POM, US, WTU),
Custer Co., Wet Mt. Valley,
RM). Denver
(CAS,
4201
(DS, F, GH, MIN, MO, NY, POM, US). Yuba Co., Strawberry
(COLO, WTU).
13021
14066
14690
Sutter Co.,
POM).
(ARIZ, GH, POM).
(ARIZ, F, GH, MO, NDG,
5123
of Salida, Munz
& Gambill
4555
14009
San Francisco,
(CAS).
Co., Mt.
from Alder Creek, Munz
14057
7917
SD,
(POM), Munz
in Santa Ynez Mtns,
1435: Munz
10 mi E of Douglas
(SD).
1907, Heller
(F). Riverside
13956
Parish
1896, Cannon
NY, OSC,
cult. from Wolf
(UC).-COLORADO:
495
Pass
Eastwood
7339
(POM, US), Munz
Co.,
(CAS, DS, NY, POM), Munz
(RSA),
Culbertson
Creek,
near Agnew,
(DS, GH, NA,
(POM). Trinity Co.,
Tulare Co., Nine Mile
into Mtns,
13994
San Francisco
San Marcos
Sierra Co., Loyalton,
15325
14610
717 (UC). San Luis Obispo
Ranch,
Creek
8861
from Julian, Munz
13021
(POM).
Island, Hoover
Santa Barbara Co., Kinevan
(CAS).
(CAS, MO).
14097
& Collethon
near Genessee,
13238: Munz
vic. of San Bernardino,
cult. from Hall
Co.,
Manning
(CU, DS, GH,
10 (DAO). Napa Co., White
Valley
Co., cult. from Wolf 6223
Co.,
5469
(ARIZ, CAS, DS, NY, PH, RSA,
(CAS, GH, NY, POM), Munz
(NY, POM, US), Munz
& Martin
& Kennedy
Heller
39
Co.,
2.9 mi E of Santa Rita
Collaland
Co., Genessee
cult. from Munz
Hood
near Fort Bidwell,
Graham
12133
Baker
(HSC). Mariposa
Co.,
Pass, Hitchcock
Munz
PH, POM),
San Bernardino
133
Co.,
Co., Soda Springs,
Canyon,
Sacramento
14603
RSA). Modoc
from Carmel Valley,
(UC). Plumas
MT,
MO,
(POM), Munz
NY, UC, US, WS).
(POM), Munz
5 mi
(GH, UC). Nevada
(NY, POM).
GH, NY, POM, UC, US, WTU).
MO,
E of Tioga
(BH, NY, POM, UC, US),
14179
14274
(POM), Munz
Canyon,
Yates 3902
(CAS, DS, MIN,
13238
(DS, OSC,
Co.,
Co., Red's Meadow,
11), Chavez
Los
(RSA, UC).
near Claremont,
Canyon
(GH, RM, UC, US). Merced
20696
Santa Ana River
Orange
Roubidoux,
9029
& Wiggins
Sec.
(T3S, R26E,
Hall
Bridge,
Park, San Joaquin River, Wiggins
San Antonio
NY, POM, RM, UC, US). Madera
MO,
59-147
Hot Creek, MacSwain
Camp,
s.n. (LAM);
Axtman
Campground
near Stoneman
Valley,
2 mi N of Bridge
Co.,
South Pasadena,
Seco,
VOLUME 50
(BH, CAS,
15271
F, GH,
(BH, IND,
ca. 10 mi E of Featherville,
Gem Co., Sweet, Nelson
& Macbride
1631
1997
OENOTHERA
(BH, GH, MIN, MO, NA, NY, RM, SMU, UC, WTU).
berg 3216
(CAN, COLO,
Anderson
s.n. (UWM).
NY, RSA),
Creek
Lewis
6166
state line, McGregor
Edwards
27587
(KANU).
Hamilton
Hitchcock
687
Kingman,
Stephen & Brooks
SW corner
(KANU,
of Meade
gor 27689
Bentley,
(KANU).
OKLA,
WIS,
Stafford
ID, MIN,
meralda
Co., Chiatovitch
Gridley
Lake,
Range,
NY, POM, WS, WTU).
Pershing
5 mi
S of Virginia
Ormsby
Pine Co.,
Chaves
Co., Roswell,
bad, unknown
Mtns,
Wooton
Hidalgo
& Standley
Co., Clanton
3713
7, 53 (UNM). McKinley
onmound,
Stephens
13281: Munz
Wagner
near Pecos,
Co.,
37978
SMU).
Socorro
(CU, DAO,
Baca
(POM), Munz
Bedker
16863
586
del Apache
(BH, DS,
14001
910
(OKL).-OREGON:
4703
NDG,
Natl. Wildlife
Co., Coos
13910
14049
River,
unknown
(UNM).
Los Alamos
Torrance
Waterfall
Rec. Area
1883, Carpenter
collector
014
2978
3 mi
(ARIZ,
Co., White
Canyon,
Fox
S of Mescalero
cult from Munz
Co., San Juan Mesa,
(MIN, UNM).
1904, Beals
Castetter
Co.,
(OKLA).
Mtns,
Co.,
&
s.n. (MICH,
(UNM).
Taos
(POM), Munz
Fourth
of July
ca. 2 mi N of Los Lunas,
Logan
Natl.
6562
13922
Co., without
Forest,
Taylor
s.n. (NA). Curry Co., Rogue
(MO). Harney
1321
9 mi E of Santa Fe, Heller
Co., Manzano
in Ouchita
Hess
Lincoln
58080701
13279: Munz
(MO). Valencia
(POM).
Eddy Co., Carls
(F, POM),
Sandoval
S of San Antonio,
71837
(CAS, F,
14048
Co., 0.5 mi N of Wag
Creek,
13281
Santa Fe Co.,
cult. from Munz
13278
Co., Water
(COLO). Mora
Sierra Co., Lake Valley,
(POM).
Godfrey
Co., Bokhoma
1075
(F,
Co.,
390 (CAN, NY,
Munz
OSC, WS, WTU).
Munz
914
(CAS, NY, POM), Munz
(POM), Munz
Co., Tularosa
Storey Co.,
& Franklin
Mtns, Metcalfe
POM, UC, US).
Refuge,
Custer Co., Weatherford,
(US). McCurtain
NY, POM, RM, UC,
MEXICO: Bernalillo
Pass, Chuska Mtns, McKnight
NY, UC, US).
Co., Capulin,
(ID). Grant Co., Blue Mtns,
Otero
Co., Embudo,
(MO, NY, US, WTU).
(NY, POM), Munz
Coos
UWL).
Co., Sher
(DS, ISC, NA,
NY, US).
River, Moore
14030
Jones
069
Camazine
F, GH, NY, POM, UC, US),
(UNM). Union
24 (UNM).-OKLAHOMA:
locality, Gephardt
Baker
13279
Mtns,
(BH, GH, NY, ORE,
4010
NDG,
of Cimarron,
(POM), Munz
ORE, US, WIS, WS).
3 mi S of Washington
Standley
ca. 15 mi NW
Peloncillo
Rio Arriba
14159
5695
SE of
4 mi up
Co.,
PH, POM). Nye
& Martin
13282: Munz
Co., near
10 mi E of
Co.,
State Park, 4 mi
12 mi NE of Silver City, Cherry Creek Canyon,
Canyon,
(DS, KANU,
GH, MIN, MO,
Co., Bosque
Co., Taos, Munz
13974
(POM), Munz
(MO). San Juan Co.,
Heller
Canyon,
26150
Lander
Ryan
NA,
Truckee
(COLO, DS, MIN,
Co., Zuni, Blackrock,
(CAS, DS, GH, RSA).
13959
3852
San Miguel
& Brooks
2794
13913
3332
(COLO, DS, MIN,
& Tierney
Wolf
& Standley
7 (UNM). Grant Co.,
collector
SMU, UNCC).
MO,
(MO). Es
POM). Humboldt
s.n. (NSMC). Mineral
fork of Gila River, Mogollon
Co.,
& Macbride
4476
Wagner
(NA, POM).-NEw
cult from Munz
13278: Munz
Valley, Wooton
463
US, UT, UWM,
Nelson
(CAS, DS, GH, MO,
Co., Mogul,
2 (RM). Colfax
Cockerell
LA, UNCC,
(CAS, F, GH, MO,
10616
of Ely, Henrichs
(POM, UC),
cult from Munz
Dona Ana Co., Mesilla
OKL,
1/4 mi W
1258
Heller
(NA, POM). Washoe
(POM). Catron Co., W
14103
GH, NY, POM, US),
Baker
Mtns,
BH, BRY,
Canyon,
Peak, C. L. Hitchcock
of
262
NY, OKL,
Canyon,
LAM, MICH,
of Sherman
Allen
(ARIZ, ASU,
(BRY, DS, NY, WTU).
1967, Mathis
S of
7542
10 mi N & 7 mi W
Co.,
Old River Dist.,
(ARIZ, NA,
Co., Kershaw
(DAO, DUKE,
Humboldt
13282
Munz
(POM), Munz
Lincoln
Co., Eagle Valley,
497
1333
7187
2.5 mi
Co.,
MO, MT, NA, ND,
TEX, UC,
Lamoille
3487
& McGregor
(UC). Elko Co., Rowland,
Ruby Mtns,
Smith Valley,
3 mi NW
West
City, Allen
2 mi S of Albuquerque,
Agency,
6884
TENN,
MIN,
of
Co.,
3 mi E of Pratt, McGre
8039
Clokey
A. S.
7 mi W
Co.,
Sedgwick
(DS). Stevens
Mtns,
Ranch, Archer
27622
(KANU). Meade
Brooks
8.5 mi N of Fallon,
4102
& Reveal
Lyon Co.,
Archer
Co., Humboldt,
POM, UC). White
RM).
of Kellog
Holmgren
S end of Ruby Range,
US).
14031
Range,
(NA, POM).
Cory Creek, Wassuk
NA,
SMU, US, WTU);
2 mi W
Creek,
SD, SMU,
[Okla
(KANU).
of Elkart, Richards
(KANU).
of Kismet,
ISC, LAM, MICH,
Kennedy
(CAS, POM, UC, WTU).
Train 2438
man Creek,
PH, RM, RSA,
Kingman
Pratt Co.,
3108
Charleston
27673
City, McGregor
27661
McGregor
SW
19259
Co.,
River, Davis
S edge of Cimarron, McGregor
(KANU).
(KANU).
5 mi
Co.,
Stephens
F, GH,
Co., Curters,
NY, RM,
28066
Churchill
DS, DUKE,
5185
Wagenknecht
Seward
Clark Co., Cold Creek,
Douglas
Pine Forest
16325
of Nickerson,
(KANU).
(KANU).-NEVADA:
PAC, PENN,
(BH, GH,
Munz
4 mi W
(DS,
Clark Co.,
McGregor
9 mi N & 2 mi W
Co.,
1001
NY, UC, US). Harper Co., Sand Hills,
Co., E of Bevidere,
s.n. (MO). Morton
Perce
11.8 mi SSE of Rock
SMU).
S edge of Dodge
Gray Co.,
163 (GH, MO,
S of Lamed, McGregor
34822
CU, DAO,
Ford Co.,
(KANU).
of Jetmore, McGregor
Kiowa
(KANU).
Co.,
NY, POM).
WVA).
Caliente,
42496
1944, Harr
87523
OSC,
10 mi W
Co.,
5 mi E & 7 mi N of Hudson,
Co.,
COLO,
ORE,
Austin,
Thompson
Co.,
WS, WTU,
2150
Co., Syracuse,
Reno Co.,
Stephens
CAS,
73895
& Brooks
(ARIZ, DS, MO,
CAN,
(KANU).
Pawnee
(KANU).
Stephens
Hugoton,
50556
(KANU,
1928,
Nez
& Grayson
S Fork of Salmon
Co.,
14759
10 mi SE of Coldwater,
Co.,
City, Krucke
(ID, NY).
Bartlett
RSA),
NY, US). Valley
Stephens
Stephens
(YU). Hodgeman
NY, SMU).
Comanche
12967
Twin Falls Co., South Hills,
S of Sun City, McGregor
S of Ulysses,
11 mi
Grant Co.,
Christ
999 (DS, MICH,
NY).
BRY, KANU,
1mi
(KANU).
2 mi E of Kinsley,
Co.,
(KANU).
4.5 mi S of Craigmont,
(DS, MICH,
just N of Riggins
Latah Co., Cedar Mtn, Moscow,
UC, WS, WTU).
& Grayson
1003
(ARIZ, ASU,
27608
Idaho Co., Salmon River,
RSA,
Canyon,
Bartlett
Barber Co.,
(POM).-KANSAS:
homa]
Co., Lawyer
& Grayson
Bartlett
(town), Holmgren
2637
DS, NY, OSC,
River N of Lewiston,
Co., Clearwater
MICH,
DAO,
163
S of Mann's
further
& Taylor
River Canyon,
Lake Ranch,
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
164
"Camp Sange,"
Williamson,
s.n. (ORE). Klamath
Ireland
1954,
Munz
14415
near Plush, Hart Mtn, Antelope
5767
(WILLU). Wallowa
on Hwy
19, above
(MO), 14437
Vista, W
of Alpine,
Hemphill
Co., Canadian
Kokernot
Ranch,
Cache
River
Valley,
Co., Logan
Canyon,
6256
2.25 mi NW
of Roosevelt,
15010
Midway,
TAES, US).
Presidio
Co., Lower Musgrave
Beaver
15262
14575: Munz
& Garrett
Carlton
sec. 19, J. L. Gentry
6704
(BRY).
to Burr Trail
Canyon,
Ogden,
to Notom
Pammel
elata
Emery
Garrett
15245
Foster
(BH, CAS,
(BRY, DS, DAO,
7640
& Blackwood
3737
Foster
4774
& Foster
(BRY, NY).
Co.,
5136
(BRY).
Kane Co., Glen
14575
(F, GH, NY,
Wasatch
Santa Clara River,
KANU,
TEX, UNCC).
1272
(MO, UT).
near Grouse
& Foster
Co., Pine Valley Mtns,
Harrison
Tierra
Canyon,
Canyon,
IND, NY, POM, UC, US, WTU).
NY, RM, RSA,
Canyon,
79, 0.8 mi E of
9072
(F). Utah Co., Payson, Munz
Rd, SE of Oak Creek Campground,
173 (MICH).
s.n. (F). Duchesne
(UT), Garrett
Co., Huntington,
5316
Cottam
0.8 mi down Grouse
Co.,
Garrett
Canyon,
(NY). Washington
& Jensen 2207
der Mtn
lc. Oenothera
NY, RSA,
US).-UTAH.
7 (NY). Daggett
Co., Buena
Limpia
along Hwy
Co., Wah Wah Mine,
Brazos
TAES, US).
Leon Co.,
(F). Salt Lake Co., Garfield,
cult. from Munz
POM),
NY, RSA,
SRSC).
head of Proctor Canyon,
Co., Paunsaugunt-Sevier,
dale, Munz
12811
Town
SMU,
Co., Farmington
Goodrich
E. J. Palmer
10 mi N of Fort Davis,
Co.,
Creek
SW of Palestine,
Co., Frijole, Grassl
Culberson
SRSC, TEX, US).
(LL, MICH,
& Corey
Gessel
TAES, US);
(GH, MO,
Peck
River,
8 mi E of Service
& Intosh s.n. (TRT). Brewster
6, 1978, Catling
s.n. (US). Jeff Davis
7943
(GH, MO,
(BRY). Davis
6500
Co.,
Co., Palestine,
NY, RSA,
& Sisson
79, Wagner
bank of Columbia
97 near
Lake Co.,
(BH, IND, NY, US).
(US). Wheeler
Anderson
(GH, MO,
of Hwy
1mi W
Lake,
15272
Co., Umatilla,
8697
(MO).-TEXAS:
(GH, PENN,
(GH, TEX,
8685,
6499
along Hwy
Warnock
1902
Creek, Neese
Garfield
4470
1853, Bigelow
6501
& Sisson
Hinckley
bridge
T142
Warnock
Davis Mtns,
Jewett, Wagner
87 (NA). Umatilla
Sheldon
645 along Hwy
17 km NW Navasota
Vieja Mtns,
Antrsi
& Sisson
Wagner
point of Klamath
14415: Munz
River,
John Day River, Wagner
(MIN, US, WIS),
2.9 mi NE of Hwy
Creek,
Co.,
Refuge,
Co., Wallowa
Co., NE
cult. from Munz
(F, GH, NY, US),
near
Co.,
T39S,
R14W,
Co., Boul
Wayne
(BRY). Weber
Co., Ogden
(GH, ISC, MO).
subsp. hookeri.
FROM
PLANTS.
SPECIMENS
CULTIVATED
U.S.A. CALIFORNIA:
Alameda Co., Berkeley, cult.DUSS-76-075,
Co., Salmon Creek, 1975, Hardham
s.n., cult. DUSS-76-077
06 and 41I). Monterey
(71I;
DUSS-88-2010;
Willow
Springs, Hwy 1, 1973, Stubbe s.n., cult. DUSS-76-021
(MO)
51,),
Co., Pescadero, Moldenke
3418, cult. DUSS-76-026
(MO) (04 and 5SI); Pigeon Point, Mold
DUSS-76-076
(MO)
(7II; 04
(MO)
and
San Mateo
(71I).
enke 3419, cult. DUSS-76-027
s.n., cult. DUSS-76-079,
DUSS-77-075
(MO), DUSS-77-076
stabilized
Creek,
(MO)
(7II); 04
(MO)
1975, Price
(MO)
(711);
(MO)
Thomas
line,
cult.
18390,
04 and SII) Santa Cruz Co., Woddell
(711;
(MO), 1975, Price s.n., cult. DUSS-76-078
cult. DUSS-76-025
3417,
Point Light House,
Pigeon
(MO), DUSS-76-080
sand dune, Moldenke
1, 14.8 mi N of Santa Cruz County
and SII); along Hwy
DUSS-76-077
(7II).
REPRESENTATIVE
SPECIMENS.
Alameda Co., Alvarado, Jepson 14496 (UC);Berkeley,
U.S.A. CALIFORNIA:
424 (UC). Contra Costa Co., Lower
Walker
Island, San Francisco
(ND). Marin
School,
Tomales,
Stinson
Co.,
Howell
23027
Munz
14304
14763
Munz
(CAS, RSA);
5704
1892, Cannon
Wheeler
& Wiggins
to Santa Maria,
Cruz
Island, Smith 6505
Thomas
Eastwood
(DS, MICH,
839
RSA,
ca. 4.2 mi
lard s.n. (MO).
Munz
3713
308
UC,
UT, WTU);
Thomas
Santa Ynez
Sonoma
Co., Petaluma,
Steele
6313
1981, Jones
5 mi
Bay,
S of
(CAS, POM, UC, US),
1599
Co.,
Spreck
(CAS); Carmel,
San Diego
of Arroyo
LA); Oceano,
Co.,
Valley,
RSA,
3 mi S of Pismo Beach,
Colma
Point Light
Canyon,
(MO). Santa Barbara
1955, Secrest
(DS, OKLA);
Creek,
s.n. (DS, NY, POM).
US),
1926, Dud
Grande,
2 mi N of Pigeon
San Bruno Mtn,
18390
Co.,
1mi N of San Onofre, Munz
s.n. (CU, DS, GH, MO,
120 (DS); Stevens
1880, Congdon
53
s.n. (CAS). Napa Co., St. Helena,
et al. s.n. (LA);
(CAS, RSA,
Wheeler
line E of Aurora
(GH, POM, UC, US).
s.n. (UC); mouth
(MO). Santa Clara Co., Sargents, Abrams
campus,
15569
(UC). San Mateo
Dam,
14695
Creek, Howitt
1902, Heller
1888, K.C.
S of Pescadero,
(CAS); Cachuma
(DS). Santa Cruz Co., Santa Cruz,
(KANU).
of Avilla
1932, Eplinger
Hardham
Co.
IND, NY, POM, UC, US). Monterey
Lake Merced,
Bay, Summers
s.n. (DS); Stanford University
4301
2520
438
s.n. (CAS);
cent
Palmer
Co.,
edge of Brooks
of Claremont,
s.n. (UC); Tomales
14304: Munz
s.n. (DS); Big Sur, 1937, Winblad
Co., San Simeon,
(RM, UC); Morro
& McClintock
1885, Rattan
San Francisco
& Bioletti
near mouth
1mi below Capistrano,
creek near San Luis Obispo,
& Smith 823
house, Abrams
(BH, F, GH,
1mi W
Co.,
LA); Marin-Sonoma
cult. from Munz
s.n. (CAS); El Cajon,
(F). San Luis Obispo
ley s.n. (CAS);
Nobs
15241
& Wiltz
Co.,
RSA,
1892, Michener
397 (UC); NW
Bowerman
(UC). Los Angeles
(BH, CAS,
Tocaloma,
Munz
1907, Patterson
(BH, POM, UC).
Heller
21254
(CAS, DS, GH, POM, US),
s.n. (RSA). Orange
Spring Valley,
4030
s.n. (UC); ca. 80 mi S of Monterey
near Pacific Grove,
1881, Jones
Silver Spring, Mt. Diablo,
Robbins
Howell
Beach,
(BH, F, POM, WTU),
els, 1908, Condit
15568
Bay, Richmond,
Co.,
1967,
adja
s.n. (MO); Santa
near Alma,
San Jose RR,
ca. 3 mi SE of Los Altos,
Siskiyou
Co., Mt.
Shasta, E.
s.n. (MIN). Ventura Co., Foster Park, 1970, Pol
1997
OENOTHERA
2. Oenothera
165
jamesii.
SPECIMENS FROM CULTIVATED PLANTS. U.S.A.
TExAS: Kerr Co., Kerrville
s.n. cult. DUSS-76-0100
(7II); Kerrville
(II) 1973, Galley
River, 2.5 mi W of Hunt, 1975, Galley s.n., cult. DUSS-76-0101
DUSS-76-099
(MO)
of the Guadalupe
(I), 1973, Galley
(MO)
s.n., cult.
and 5SI); South Fork
(04
(MO), DUSS-76-0102
(MO)
in valley of Kino, 1968, Linder s.n., cult. DUSS-87
Pref., station Koyaguchi
(7II). Japan. HONSHU: Wakayama
2037 (MO) (71I), DUSS-87-2036
(010 and
(MO) (08 and 3LI), DUSS-87-2038
(06 and 4II)' DUSS-87-2037
also Jean & Linder, 1979). Mexico.
COAHUILA: Parras, Munz 15077, cult. DUSS-88-2013
(MO)
(see
(M,
211)
MO)
Ind. Sem. Bot. Gard.,
CANARY ISLANDS: La Paz,
(7II). Spain.
Puerto
1975 no. 96,
de la Cruz
(Tenerife)
(29'03'N,
102?35'W),
cult.DUSS-77-0358 (MO) (014).
REPRESENTATIVE SPECIMENS. Mexico.
(OKLA,
15251
cult. from Munz
TEX);
15077
(CAS, GH, NY, POM, US); Villa Acuna
US).-NUEVO
LE6N: 2.3 km W
of Laguna
et al. 10133
(LL, MO);
(US). U.S.A.
KANSAS: Clark Co., Clark
of Beckham
534
COAHUILA: Sierra del Carmen
from Parras, Munz
Monterrey,
Co., Eskew
1380
(MO); E of Bridgeport,
(POM, UC, US), Munz
POM, UC, WTU),
Custer
Co.,
MICH,
OKLA,
RSA,
tuck, Stevens
OKLA,
2899
Jackson Co.,
Lake Supply
on Wolf
(TEX). Comal
Creek,
1mi W
ell 1098
of Gruver,
(ARIZ, F, NY, SMU).
82558
Sec. 34, Pohl 4659
ton on Hwy
River
Bryant
(SMU). Nolan
on Hwy
Reed & Morton
(FSU, MICH,
3995
OKLA,
3478
11490
s.n. (NY). Randall
(TAES, US). Randolph
Cory 33387
(POM). Victoria
TEX).
City,
Verdun
Pref., Maeyama
Farm,
1899, Galpin
Shinomachi,
1904, Bononi
2585
Sakiya
(GRA, PRE); Warrenton,
Adams
(POM); Hunt,
8
Fryx
1mi E of Dar
Creek,
12 mi SE of Perry
1mi N of Canadian
15 mi N of Amarillo
Co., Sierra Tierra Vieja, Lower Mus
s.n. (MIN, MO);
Lindheimer
502
(TEX). Tom
(GH).
700
(P).-HON
CAPE PROVINCE: French Hoek,
distr., Modderrivier,
du Plessis
Palo Duro
Young 224
1889, Faurie
Kirosaki,
97 (MO). South Africa.
s.n. (FH, LY); Middelburg,
Hope
Co.,
Co., branch of Pal Duro Canyon,
s.n. (GRA, PRE, STE); Kimberley
1962, Stadler
Cape Town, Meistland,
Poort,
Hirota
(MICH,
Co., Borge,
Co.,
Potter Co.,
1900, Eggert
Co., Guadalupe,
Cory 50262
Co., Hackberry
Oldham
NY).
(BRY). Presidio
SPECIMENS EXAMINED FROM OUTSIDE NATURAL AREA. Japan.
SHU: Kyoto
Deaf
63 (F, GH). Hansford
(OKLA, TTC);
s.n. (US). Ochiltree
SMU,
Co., Canyon
MO,
Cory 23967
T-7 (FSU). Mitchell
(CAS, DS, MO,
326
Co.,
F, GH, MIN,
Hutchinson
10583
(MO,
Stratton
s.n. (GH, SMU).
Thurber
TTC).
Rowell
1881, Harvard
& Duncan
DS,
1868, Engelmann
TEX,
(OKL,
404
further locality, Tharp 224
14.5 mi S of Canadian,
Co.,
973
Stratton
OKL). Woodward
(ARIZ, CAN,
Co., Fredericksburg,
Bottimer
Co., Menard,
river bottom, Higgins
1946, Hinckley
Green Co., Christoval,
7925
(KANU,
13.75 mi SW of Kerrville,
Co., Champion,
Rd, Ferris
87, then 3 mi E along
grave Canyon,
Canyon,
Menard
83 and 5 mi E, Wallis
on Amarillo-Dalhart
808
Co., 6 mi S of Lipscomb,
Lipscomb
(KANU).
s.n. (ARIZ,
near Lugert Dam,
1253
808:
(MICH, OKLA,
100 (OKL, UC);
POM, US).
Ellis Co., near Shat
Springs,
Bell Co., without
Lindheimer
from Lindheimer
4327
(CAS,
(DS, KANU).
Co.,
14712
(BH, CAS, GH,
1951, Waterfall
Co., Doby
Nighswonger
s.n. (OKL).-TEXAS:
Braunfels,
13575
6 mi E & 2.7 mi N of Tuttle, Pearce
s.n. (OKL). Kiowa
229 (OKL). Hemphill
Rowell
27217
Stephens
& Musselman
13580: Munz
14769
POM), Munz
(OKL). Harper
(SMU, TTC). Gillespie
Cutter
5 mi E of Canadian,
rouzett, Stephens
384
1936, Hopkins
cult. probably
1170
Waller
(NA). Kerr Co., Kerrville,
Bull
Spring, New
S of Dawn,
8 mi S &
(CAS, GH,
0.5 mi SE of Weatherford,
Co., 0.5 mi N of Taloga,
1974, Springer
Co., Comanche
Smith Co.,
SMU);
14711
(OKL);
7 mi W & 8.5 mi N of Alva,
Co.,
PH, TEX, UC, US),
Co.,
926
of Altus,
NY, OKL,
NY,
13575: Munz
s.n.
SE part
Co.,
37, Lawson
Munz
Chiang
1907, Arsene
cult. from Munz
3 mi SW of Norman,
Co.,
(ARIZ, GH, MO,
Beckham
on Hwy
Munz
100?18'W),
de Cholula,
(MO).-OKLAHOMA:
0.25 mi N of Cogar
Co.,
Cleveland
4 mi N of Reed,
3 mi W
Woods
(MO, OKLA).
26559
(DS, GH, MIN, MO, OKL, US). Grady Co.,
Co.,
Route
(CAS, F, GH, NY, POM, US),
Mericle
SMU). Dewey
Greer
SMU).
OKLA).
from Munz
3 mi SE of Thomas,
584
764
Marsh
IND, NY, POM, US, WTU),
Wynd & Mueller
100?55'W),
(BR, MO).-PUEBLA:
Caddo
13580
(BH, GH,
& 0.5 km E of San Isidro (25?21'N,
State Lake, Raven
(POM, WTU).
14768
cult.
6115
(MO, OKL).
Munz
(29?18'N,
de Sanchez
Arsene
15313
250
15 (GRA).-NATAL:
1952, Mostert
s.n. (PRE);
(PRE); Queenstown,
Pietermaritzburg,
Bongolo
Ward 7 (NU);
Durban, Springfield,Akitt 17 (NU).-ORANGEFREESTATE:
Bloemfontein, Loch Logan, 1968,Hanekom 1044
(K).-TRANSVAAL:
aanspoort,
Smith
Potgietersrus,
Burt-Davy
P, PRE, SRGH,
Repton
Evans
m,
1373
4015
Pretoria,
1965, Gerber
9789
Z); Pretoria,
(PRE); Van Wyksrust,
1961, Devenish
633
Dist.
Moss
Roodeplaat,
Codd 2756
18378
(BM); Wakkerstroom,
Orotawa
unknown:
Louw
1375
s.n. (GH); Pretoria,
Dam,
Leendertz
1703
Pretoria, Burgers
11072
Bavi
(PRE);
Park, 1942,
(PRE); Tsaneen,
spruit, near Dirkiesdurp,
(PRE); Wondsfontein
Raven
1969, Hansen
Theron
(BM, BR, G, K, M, MO, NH, NY,
(SRGH);
Mabola
near Steynsdorp,
and Agua Manza,
1938, Louw
Strey 3171
12365
(PRE); Standerton,
(K, PRE, SRGH); Welverdiend,
between
TVI,
s.n. (PRE); Loskop
s.n. (K); Pretoria, Mogg
110 (GH, PRE).-Province
CANARY ISLANDS: Tenerife,
Pretoria,
1941, Phillips
(PRE); Potchefstroom,
1928, Louw
(K); 19 km NE of Pretoria,
end, Liebenberg
s.n.(US);
117 (NH, PRE); Krugersdorp,
& Raven
s.n. (C).
Pole
ca. 1700
Farm near Welverdi
26109
(MO). Spain.
SYSTEMATICBOTANYMONOGRAPHS
166
SPECIMENS CULTIVATED IN BOTANICAL GARDENS. Switzerland.
MASSACHUSETrs:
Hort. Cantab.
Botanical
[Cambridge
Geneva,
seeds
Garden],
VOLUME 50
s.c. s.n. (G-BOIS).
1856,
from Texas,
1848
U.S.A.
(NY).
3. Oenothera longissima.
SPECIMENS FROM CULTIVATED PLANTS. U.S.A.
turnoff on the rd to Point
2014
062
(MO)
(7II; 04
074
(MO)
(7II; 04
from seeds of BRY,
16348,
Canyon W
near Virgin
River
along moist
cult. DUSS-82-0346
roadside
(MO)
near Rockville,
at the
(7II)' DUSS-88
(MO)
and SII; 204 and 311).-COLORADO: Delta Co., E
of Delta, Weber & Steward 15240, cult. DUSS-76
(04
50, W
Co., McElmo
and SII). Montezuma
Co., N rim of the Grand Canyon
s.n., cult. DUSS-76-0104
of Cortez,
and 511).-UTAH. Garfield Co., 12 mi NE Escalante,
and 3II). Grand Co., 2 mi NNE of Moab, Negro Bill Canyon,
(2 04
(MO)
Welsh
(MO)
of jct. with Hwy
10 mi W
Plateau,
ARIZONA: Coconino
1975, Keliher
(MO), DUSS-77-091
(MO), DUSS-77-092
side Uncompahgre
ca. 2800 m,
Imperial,
s.n., cult. DUSS-77
1976, Kelly
Welsh
cult. DUSS-82-0345-2
19283,
Castle Valley
Co., Zion Natl.
s.n., cult. DUSS-76-053
1975, Harrison
Sec. 29),
(T25S, R22E,
and 4II). Washington
(06
Park,
(7II; 06
(MO)
08 and 3II); Zion Natl. Park, along main rdjust W of tunnel, 1975, Reveal s.n., cult. DUSS-76-052
(MO)
4I,;
(MO) (7II).
(7II; 2 04 and 3II; 08 and 3II); Zion Natl. Park, Foster 5333, cult. DUSS-0348
10200 (MICH 5
REPRESENTATIVE SPECIMENS. U.S.A. ARIZONA: Coconino
Co., Big Springs, Eggleston
and
& Lehto 203
sheets); 4 mi N of Cape Royal, Halvorson
(RM);
14 mi NNE
3185
of Kaibito,
(BH, CAS, DS, RSA). Mohave
NIA: Inyo Co., Antelope
(HSC).
Co., Eastern Mojave
IND, KANU,
Mtns,
Charleston
MICH,
MIN,
3591
& White
19283
(BRY). Kane
(BRY); 7 mi NW
Juan River,
Canyon,
confluence
1mi N of Bluff, Munz
13008
(DS, NY, POM, UC), Munz
14035
NY);
Mon.,
Welsh
between
& Moore
Oak Grove
2458
15, Atwood
533 (ARIZ, UT); Hurricane,
ter & Foster
5353
Wayne
Co.,
(BRY);
15009
Zion Natl.
on Hwy
Atwood
between
Kachina
Co.,
(BRY, RM);
E Bench
Park,
IND, ISC, LA, LAM,
LL,
Creek,
Co., Colorado
(T35S, R4E,
Sec.
1),
& Allen
R5W,
3156
2816
Cutler
from East, Gra
(T25S, R22E,
Sec.
26),
Sec. 29), Welsh
1977, Foster
137 (UT). San Juan Co., W
Lindsay
(BRY);
Dark
cult. from Munz
Canyon,
Cataract
Pine Valley Mtns
Campground
SW
053
R20W, Welsh
(T29S, R7E,
29,
Natl.
rd
1/4 Sec.
15 (T41S, RIOW),
Harrison
T42S,
13909
17-20,
Bridges
(T42S, R16W,
to Hwy
near Rockville,
Secs.
Natural
along Oak Grove
3 mi SE of Santa Clara
Wash,
13008: Munz
95 (T37S, R20E,
& Sipapu Bridge,
S cen. Henry Mtns
s.n.
side of San
(BH, DS, GH, LAM, MO, NA, ND,
via Hwy
Bridge
River,
Garfield
Calf Creek
of Zion Natl. Park adjacent
Park near Virginia
at Pleasant
Clark Co., Charleston
GH,
Valley
89 (T43S,
(DS, F, GH, NY, POM); Lytle Ranch, Beaverdam
2 mi due N of Notum
(ARIZ, CAS, COLO,
tributary to Green River
Castle
11 mi SW of Blanding
5394
15240
& Steward
Weber
7940
Calf Creek Falls & Calf Creek Recreation
Canyon,
in the Rock,
River,
Escalante,
Lower
Bill Canyon,
(BRY, NY). Washington
& Hwy
26), Christian
Verken, Munz
between
Negro
Canyon,
s.n.
(POM).-UTAH:
& Lund 574
(RSA, UT);
slope of Navajo Mtn,
(POM);
Lloyd
(CAS, GH).-COLORADO:
of Delta,
F, FLAS,
(BH, CAS, CU, F, GH, NY, POM, UC),
74 (COLO); White
30, 32), Northcutt
50, W
Mts],
rd, 1967, Apperson
Nichols
14785
Colorado
(UT); W
168, White
TEX, TRT, UC, US, WS, WVA, WTU).
729 (NA); Hole
with
& Jotter 2147
14187
Munz
DS, DUKE,
9 mi E of Kanab
Co., Vermillion,
of Kanab, Hester
above
Clover
of Moab,
52 (DS, POM).-CALIFOR
Lake Canyon
Wolf
POM, SMU, TENN,
PENN,
Cottam
Braem
TEX, WS, WTU).-NEVADA:
CU, DAO,
Ruby Mtns,
Trail,
of jct. with Cortez Rd, Weber
(BRY). Grand Co., Florence
2 mi NNE
(MO, POM);
York Mtns,
740
Indian Gardens,
Angel
Pass, N of Hwy
of jct. with Hwy
12 mi NE of Escalante
(BRY);
(T34S, R4E), Welsh
16348
Canyon,
15 mi below Hite,
Creek,
Canyon,
from Elizabeth
New
RM, RSA,
NA, ND, NY, OKL, OKLA,
River, Warm
ham 9961
OKLA,
(CAS, COLO,
Bright
E of Westward
17 mi W
Canyon,
NCSC,
5542
5 mi S in Lamoille
Elko Co.,
Neese
MIN,
Clokey
10 mi W
Plateau,
Co., McElmo
Desert,
(MNA);
Sawmill
Mtns,
[Deep Springs Valley,
Co., E side Uncompahgre
DAO,
Co., Hualapai
735
below Tovar, Hanson
Grand Canyon
(ASU, NCSC);
& McDougall
0.5 mi down Ruby Canyon
Co.,
San Bernardino
(COLO). Montezuma
Area
Springs
(UC). Los Angeles
3015
Delta
Theroux
Fos
(DUSS);
23654
Sec. 25), Neese
La
(MO).
& White
3751 (BRY).
4. Oenothera
wolfii.
SPECIMENS FROM CULTIVATED PLANTS (diakinesis
lished
Hoch
inWasmund,
1853,
1980,
1855a,
and Wasmund
1855b,
& Stubbe,
cult. DUSS-83-036
cult. DUSS-81-602
(MO) (014); Wilson
(0 14). Humboldt
Co., S of Cape Mendocino,
cult. DUSS-76-081
(MO), DUSS-83-605
Luffenholtz
Beach
cult. DUSS-76-047
County
(MO)
Park, 2-2.5
DUSS-85-125 (MO) (014).
Stubbe
Stubbe
(MO)
were made
(MO)
mi S of Trinidad
Stubbe
(MO) (014);
(T8N, RIE,
Sec. 31), Montalvo
along Hwy
(014);
9,
(MO)
Petrolia, Hoch
(MO)
City,
Stubbe
(MO), DUSS-81-604
14, cult. DUSS-81-605
Curry Co., Pistol River
and pub
Co., Crescent
(0)14), DUSS-88-2025,
10, 11, cult. DUSS-81-603
14a, cult. DUSS-81-606
(014),
by 0. Wasmund,
CALIFORNIA: Del Norte
(MO), DUSS-78-061
Creek,
(014).-OREGON:
examinations
1986). U.S.A.
1365,
S end of
& Ackernan
747 p.p.,
101, 1984, Stansell
s.n., cult.
OENOTHERA
1997
CALIFORNIA: Del Norte Co., beach mtns, Crescent
ADDITIONAL SPECIMENS EXAMINED. U.S.A.
223
Hitchcock
19510
Radio
between
City,
Rd,
Cove
S of Crescent
City,
toWilson
City, Keck
(DS); Sister Rocks
jct. with Hwy
northern
488
Thompson
Co., Willow
(DS, WTU);
Crescent
6 mi S of Crescent
City, Tracy 15602
s.n. (HSC);
(NY, POM, US), Munz
cult.
(BH, GH, MO,
1931,
(UC); Willow
Van Dyke
Peck 8663
NY),
14609
moist
(CAS, F, GH, NY, POM, UC, US, WTU);
(UC), Tracy 8302
just S of point on bluffs
s.n. (MO); beach, Harbor,
1921,
14698
(CAS,
3113
(UC); Crescent
ca. 100 ft. N of Pistol River
City,
Humboldt
Beach
County
(MO); 2 mi S of Cape Men
13977
(POM), Munz
14050
POM), Munz
14692
(UC), Tracy 18401
Port Orford,
the Heads,
8662
sec. 13), on bare weathered
4 mi E of Gold
River,
the beach, Brookings,
rock
Kildale
Beach,
bridge on E side (T38 N, R14 W,
bank above
(UC).
Peck
Imper s.n. (HSC); Otter Pt.,
1985,
(T36 S, R15 W,
(WILLU);
Requa,
Tracy 13598
s.n. (MO); Rogue
Peck 5777
River,
14386: Munz
3 mi S of point of Cape Mendocino,
Curry Co.,
and slopes
Stansell
1985,
101, on sandy roadbank
(DS); Hwy
1984, Stansell
local in small patch,
S of Otter Pt., N of Gold Beach,
(WILLU);
individuals,
6057
14643
1mi
(GH, NY,
(type), Munz
Trinity River Valley,
Creek,
and loose
ca. 100
6172
s.n. (CAS).-OREGON:
of swamp
(JEPS, UC, WTU).
747 p.p.
(NY, POM), Munz
bluff,
ca. 3 mi N of Gold Beach,
rubble,
& Ackerman
from Wolf & Johnson
Imper s.n. (HSC);
(HSC); S end of Luffenholtz
2246
Sec. 31), Montalvo
Imper
Co., Carrville,
Anderson
(DS); Luffenholtz,
(T8N, R1E,
City, Parks
along
Jetty, Imper
Lester & Yearout 224
Creek,
up Klamath
IND, POM, WTU);
City,
City,
1987,
cult. from Munz
bluff
14262
Tracy 4968
3 mi
(NY);
(POM, UC),
(BH,
1987,
Trinity
to Nickel
coastal
(POM), Munz
docino,
788
101, Lowry
14386
City, Munz
of Crescent
Imper s.n. (HSC); margin
Enderts Beach
(MO); Cres
1987, Imper s.n. (HSC);
City,
of Smith River,
1987,
15244
7168
Creek, Abrams
Park, 2-2.5 mi S. of Trinidad
City,
City, C. L.
1855
Imper s.n. (HSC); Crescent
Pt., 1987,
Quad.,
(MO), Hoch
s.n. (HSC); NW
Imper
s.n. (HSC); mouth
Imper
1853
City, Baker
S of Crescent
1mi
Island near Crescent
1987,
15 mi S of Crescent
s.n. (DS); beach, Crescent
NY, POM, UC, US), Munz
101,
City, Hall
1987,
Creek,
5613
SW of Klamath,
(HSC);
McGregor
and Hwy
Aquarium
tide mark
City, Hoch
1987, Imper s.n. (HSC); Preston
Blue
S side of Crescent
s.n. (HSC);
False Klamath
S edge of Crescent
Imper s.n. (HSC); Crescent
1987,
just above high
beach
(UNM);
(COLO, RSA, WS, WTU);
cent City, S of Pt. St. George,
Crescent
3561
City, Fuller
(JEPS); Crescent
167
sec. 19),
Peck
20440
(WILLU).
5a. Oenothera
villosa
subsp. villosa.
SPECIMENS
FROMCULTIVATED
PLANTS.
Canada. QUEBEC:
Carleton Co., Huntley, 1972, Forstner s.n.,
cult. DUSS-76-R19
Finland.
(014).
(0 14). France.
Tdpi6szeli
1978,
Roma
tatiski s.n., cult. DUSS-76-R7
84-220
(MO). Russia.
1982, Skvortsov
1981 no. 1655,
cult. DUSS-77-0201
(MO)
(MO) (014).-KRAK6w:
s.n., cult. DUSS-84-235
(MO)
(0314; Renner,
Jaworzno,
(MO)
(MO)
Hun
Ind. Sem. Bot. Gard.
(014).
near Rim
Poland.
GDANSK:
1942); Gdanisk-Stogi,
1983, Rostatiski
s.n., cult. DUSS-78-0157
(MO). Slovakia.
cult. DUSS-77-0220
(MO). Ger
(MO) (014).
(MO);
cult. DUSS-82-0452
(MO)
cult. DUSS-82-0380
s.n., cult. DUSS-92-2013
cult. DUSS-77-0221
(MO)
cult. DUSS-84-182
Italy. EMILIA-ROMAGNA: Prov. Forli: Miramare
no. 852,
1978
BRYANSK: 1975, Alexeev
1975 no. 437,
Leipzig
no. 3299,
(MO) (014).
(Rome)
of 0. Renner,
collection
1990, Wolfstetter
1975
Vd'crdt6t
cult. DUSS-79-0616
ini, Ind. Sem. Bot. Gard.
Heubude,
215 m, Ind. Sem. Bot. Gard. Basel
near Aschaffenburg,
PEST: Ind. Sem. Bot. Gard.
no. 433,
1974
Ind. Sem. Bot. Gard. Lyon 1982 no. 67, cult. DUSS-84-249
(MO).-RH6NE:
BAYERN: Nickheim
gary.
Ind. Sem. Bot. Gard. Helsinki
HAUT-RHIN: Chalampe,
(MO), DUSS-83-0142
many.
Ind. Sem. Bot. Gard. Tallin 1975 no. 670, cult. DUSS-77-0378
(MO) (0 14). Estonia.
UUSIMAA: Tenhola,
& Dietrich
1974, Ros
s.n., cult. DUSS
KRAY:
(014).-PRIMORSKIY
ZAPADOSLOVENSKY: Pie?t'sny,
Ind. Sem. Bot. Gard.
(014).
UNVOUCHERED
WITHDIAKINESIS
STRAINS
CONFIGURATION
EXAMINED.
Hungary. BUDAPEST:
Ind. Sem.
Bot. Gard. Budapest
zokowa,
1974 no. 2467,
1983, Rostatiski
Ind. Sem. Bot. Gard. Kosice
Natl.
Forest,
top of Magazine
der Co., Boulder,
der, collection
Weber
cult. DUSS-77-0369
& Dietrich
1975
no. 1815,
cult. DUSS-77-0178
31, 32, cult. DUSS-77-0210,
0212,
DUSS-77-0213
0371
(0) 14); 2 mi E of Manhattan,
ter 049,
cult. DUSS-82-0374
82-0377
(0)14).-MINNESOTA:
WYOMING: Laramie
(014).
Shawnee
Crow Wing
Co., Laramie,
1973, Stubbe
Co.,
Co.,
1976, Crawford
REPRESENTATIVE SPECIMENS. Canada.
(014),
along
ARKANSAS: Logan
Co., Ozark
rd to Flagstaff
Creek, Weber
Mtn,
Manhattan,
932,
s.n., cult. DUSS-77-0218,
039,
cult. DUSS-82
ALBERTA: near banks
of Oldman
N Bushnell
24, Wetter
043,
cult. DUSS-82-0435
77-0219
River,
Boul
near Boulder,
59, 60, cult. DUSS-77
Wetter
along Hwy
Clemants
Boul
(014);
(MO) (0) 14). Riley Co., Manhattan,
DUSS-88-2024;
and Szc
(014).-COLORADO:
6.6 mi E Waemego,
0.7 mi E Rossville
Crosslake,
Jaworzno
VfCHODOSLOVENSKY:
s.n., cult. DUSS-77-0182
South Boulder
Co.,
041, cult. DUSS-82-0373
DUSS-77-0217
U.S.A.
cult. DUSS-78-069
Pitkin Co.,
(014);
Pottawatomie
Wetter
cult. DUSS-77-0216,
045-4,
1650 m,
KRAKOW: between
and 111). Slovakia.
(014).
72303,
(014).
DUSS-77-0211
(014).-KANSAS:
045-77,
Complex,
Poland.
(012
cult. DUSS-77-0359
ca. 830 m, Demaree
Mtn,
UC campus, Kittredge
of 0. Renner,
(014).
s.n., cult. DUSS-84-219
Barkley
Hall, Wet
cult. DUSS
(0 14).
(014).
N of Pincher, Moss
283
SYSTEMATICBOTANYMONOGRAPHS
168
COLUMBIA: Okanagan
(GH).-BRMTISH
Brandon, Macoun
WTU);
12884
NEW BRUNSWICK: Kent Co.,
5 mi
Marie
1966, Squires
167
Island,
River,
Preston,
1932, Groh
(TRT); Center
(DAO):
8541
Garton
et al. 3383
ley Twp., Soper
Erie, Soper 337
(MT); Charlevoix,
& Knowlton
David,
s.n.
Rouleau
Vercheres,
Marie-Victorin
14134
a Langlade,
Rivire
Craighead
Co.,
ORADO: Adams
(BH, DS, NY, POM).
& Brooks
Stephens
6317
Osterhout
42852
dum 400
(Lombard),
3 mi W
(IND). Floyd Co.,
Deam
50756
Deam
Brownstown,
17449
1.5 mi SE of Oswego,
(POM). Lawrence
31856
2 mi W
(IND). Monroe
(IND). Ripley
Co., 4 mi NW
Sullivan
ner s.n. (IND). Wabash
Lebanon,
Deam
kee Co., New
Black
Hawk
Co.,
50634
Albin,
Co.,
POM).
Cherokee
55899
Shimeck
Monson
Co.,
bluffs N of Marquette,
Co.,
near Milton,
52251
3 mi NW
Co.,
Co., W
s.n. (SMU).
Co., Orange
Co., Fitzpatrick
Tama Beach,
Fayette
1894, Fink s.n. (GH). Floyd Co., E of Nora
Lammers
474
(ISC). Dickinson
Co.,
Co.,
49866
Deam
14008
2.5 mi
(IND).-IoWA:
1923,
Shimeck
et al. 3867
City, Munz
Twp., Cooperrider
37829
(BH, POM,
1941, Zel
SW of W
Allama
s.n. (ISC).
(ISC). Calhoun
17537
(BH, IND,
s.n. (ISC). Clayton
62 (MIN). Davis
61 (MO, RM). Des Moines
Co., N of Miller's
Springs,
SW of
Deam
Co., Evansville,
1926, Spiker
& Fitzpatrick
2 mi S of
4 mi
(IND). Kosciusko
Munz
Pammel
Co.,
Co.,
Co.,
s.n.
52599
S side of Lake Maxinkuckee,
55432a
5 mi N of Mason
Chickasaw
Co., N of Burlington,
Co., Fayette,
Deam
Deam
2 mi SW of Morton,
of Unionville,
Co., Ledges,
Co.,
s.n. (NY). Clinton
s.n. (ISC). Decatur
Co.,
1932, Edwards
near Liverpool,
(IND, POM). Warren
of Bluffton,
(MO). Boone
1955, Carter
River,
(IND, POM). Vanderburgh
s.n. (WIS). Appanoose
1927, Shimek
1931, Shimek
51013
20307
s.n. (F).
(F).-INDIANA:
Franklin
Jackson
just E of Friendship,
Creek
of Treaty, Deam
716 (ISC). Cerro Gordo
Co., Cedar Twp.,
(IND, ND).
(IND). Marshall
1913, Nason
64858a
S of Fredonia,
(MICH).
159 (IND). Putnam Co.,
from Laughery
Deam
of Grayville,
2.5 mi W
17234
Ellis
no further locality, Burk 561
1mi S of Lohrville,
Co.,
Deam
of Mitchell,
(IND, POM). Wells
1917,
52585
Co.,
border, Fox & Wee
of Fayette,
0.5 mi
or
(DAO).
(POM). DuPage
36134
Co., 2.5 mi S of N Vernon, Deam
Co., Bloomington,
cult. from Deam
Co.,
Co.,
13276
Fulton
11309
Chase
Co., Algonquin,
(IND, POM). Lake Co., N side of Deep
55322
(CAS).
Co., Windsor,
(MIN).-GEORGIA:
Peattie
11437
Deam
of jct. of
14514
0.5 mi SW of Beulah,
of Ashland,
Co., 0.25 mi W
state rd 62, Deam
Ewan
(COLO, RSA). Weld
2580
(US). McHenry
Co., Camby,
(DS, IND). Jennings
Deam
Co.,
Hendrick
Ouachita
(MO).-COL
3.8 mi W
Forest,
quad., near Missouri
s.n. (ISC). Crawford
along
Albany,
POM).
7199
Moyer
Cass Co., W
(IND). Boone
city
El Paso Co., Palmer Lake, Munz
Cook Co., Glenview,
410
65392
Plume,
Co., 2 mi SE Custer Park, Steyermark
1907, Urban
of New
Co., Wilton,
Belle
S of Mobile
s.n. (KANU).
(IND). Pueblo Co.,
Jersey Co., Brussels
34587
Deam
Green,
(IND,
(GH, NY).
s.n. (CU). Will
of Ft. Wayne,
13024
Julesburg, Weber
Co.,
Fairfield
Crampton
Co., Kankakee,
Clay Co., Bowling
Laurel,
of Conifer, Munz
s.n. (TEX).-ILLINOIS:
12685
Natl.
(MT);
(MO, NY, POM, UC).
Jasper, 1955, Demaree
Co., Roosevelt
428
and Miquelon.
22394
(MO). Clear Creek Co., Silver
Portage
Pontiac,
& Bibbey
Co., Bald Knob, Demaree
(BH, IND, NY, POM, US).
(ASU, UNCC).
1913, Cottolu
8 mi NW
Co.,
Jones
Keil 520
4021
Sedgwick
1926, McClung
(NEB). Kankakee
Pike Co., Shepherd,
(PENN).
13273
(RM, UC).-CONNECTICUT:
Co., near Urbana,
Champaign
York Center
RSA).
(KANU).
Co.,
96 (DAO, MT, US);
Pierre
Demaree
Val
(US); Terrebonne,
0.6 mi N of Dog River,
Co.,
Springs,
(POM). Boulder
4 mi W
Co.,
Siloam
OKL, RSA). White
13428
Munz
Co.,
(RSA). Newton
26519
(GH, MO,
Munz
Jefferson
Co., near Atlanta,
DeKalb
Benton
St. Vrain Creek, Wagner
Co., 5 mi S of Littleton,
Douglas
Deam
37726
Co., Brighton,
36 & 7, above
Allen
Demaree
Demaree
ALABAMA: Mobile
(GH). U.S.A.
612
& Cyprien
1923, Chamberlain
Frankton
8Q0 (NY). Saint
Co., Conc.
Simcoe
Allyre
119 (DAO, MT);
Lake,
Hunt
Turkey Point, Lake
95 (DAO);
& Rolland-Germain
Manitoba
s.n.
1896, Pollard
(TRT); Carleton,
11121
s.n.
561
from seed from Hi
dump, Asbestos,
118 (DAO, US),
Plain, Macoun
Louis
(MO).-ARKANSAS:
Jonesboro,
Co., Eagle Mills,
Hwys
350
Arsene
8156
limits, Lelong
Marie-Victorin
(MT).-SASKATCHEWAN:
(GH); Assiniboia,
Mine
(DAO);
Minshall
Elmbrook,
Lac Megantic,
Frontenac,
0.75 mi W
6496
1939, Krotkov
s.n. (DAO); Norfolk,
Islington, Welch
Ste.
& Grayson
River,
Lake,
628
(CAN).
Gloucester,
Gillett
(US); grown
Twp., Shields
1938, Urquhart
& Rolland-Germain
et al. 2004
10111
(DAO); Port Albino,
(MT); La Ferme, Morton
11633
Cleonique
Eagle
11690
695.5
(VPI); Richmond,
(MT); Lac St.-Jean, Mistassini,
Contrecceur,
Gates
1576
(BH, IND, POM, US,
Scoggan
Prince Edward,
S Crosby
(MICH).-QUEBEC:
Cayouette
(GH); Laprairie,
2495
du-Fort, Marie-Victorin
Saskatoon,
3113
Saint-Hilarion,
Owens
s.n. (TRT); York,
1978, Varga
Twp., Bobbette
12, Vespra
Point Pelee,
(DAO); Essex,
(DAO); Toronto,
15285
of Marmora,
Frontenac,
(UBC);
Lake Opinicon,
E of
s.n. (TRT); Rainy
3.5 mi NE
Point, Morton
(POM); Alderdale,
1890, Scott s.n. (TRT); Leeds,
(US); Ottawa,
V, Lot
13451
Munz
s.n.
(DAO, GH);
(BH, IND, NY, POM);
& Mayall
Co.,
62248
& Brayshaw
1889, Macoun
6530
2 mi NE of Pine River, Bartlett
s.n. (DAO);
1156 (DAO); Dawson
Finch, Minshall
Stormont,
Park, near Guelph,
pot-lo
17514
1936, Cormack
Hastings
Kamloops,
& Breitung
GH, MT); Melita,
s.n. (DAO);
& Squires
(UBC);
from Haskett, Munz
Munz
1912, Herriot
Merriles
Petawawa,
s.n.
Lake, Boivin
(ALTA, CAN,
(DAO).-ONTARIO:
s.n. (DAO); Galt,
Lake,
Clemens
of Max
S of Kouchibouguac,
Bay, Pardee Twp.,
Thunder
(MICH, RSA);
of Rainy
8015
& Baldwin
Scoggan
Rd, Shippegan
1935,
cult. from seed from Stevens
(CAN);
Mtn,
Porcupine
Mission,
Turtle Mtns, W
Souris District,
(GH).-MANITOBA:
VOLUME 50
1931, Shimek
Bay, Shimek
222
s.n. (ISC). Greene
(NY).
Co.,
1997
OENOTHERA
Grand
17536
Junction, Munz
Co., 6 mi SE of Modamin,
& 6 mi W
of Ida Grove,
Twp., Brown
Hayden
(ISC, TEX). Marion
(ISC). Muscatine
TEX).
Co., Knoxville
7 mi NW
Co.,
Pocahontas
Sac Co., Albertson
of Ames,
351-113
2126
Hayden
rill 1132
(ISC). Worth
Co., Fort Dodge,
Marsh
Barton
3 mi E of Claflin, McGregor
Co.,
Butler
(KANU).
Green,
8705
29527
Stephens
83288
& Miller
s.n. (KANU).
Geary
59396
4386
Co., Meade
Morris
& Hauser
10763
Stephens
50556a
(KANU).
59130
(KANU).
of Waemego,
046
59077
W
(KANU).
Scott Co.,
of Richland,
(KANU).
Raven
(KANU,
3687
11488
Co.,
Stephens
2 mi
26526
(MO). Wallace
2 mi NW
of Yates,
(IND). Calloway
UNCC).
Meade
Between
S&
1mi W
18843
14201
glas Lake,
1850
Ehlers
Co.,
Palmer
Co., Sleepy
gorge
just below
bridge,
J.A.S. 409
outlet
1955, Marlett
13473
Kent Co.,
2528
Fowl
Earth Co.,
Lake, Burns
Anoka
& Hendrickson
324
Co., Lake Christina,
481
Deam
588
Cen
Co.,
7431
Lake,
(GH). Wayne
G-3
& Ranzinger
(MIN). Bel
509
(MIN).
(MIN). Cook Co., Pigeon
S3470
Co.,
s.n.
2 mi NE of Dou
Co., Canadaiqua
Cottrell
cult.
Fernald
1927, Beattie
Co.,
(MIN). Dakota
(NY,
Trigg Co., Land
Co., Cumberland
Farwell
Fridley,
(RSA). Wood
Co., Horwich,
Lenawee
Sheldon
609
of Bandana,
Co., Dracut,
3001
1mi
J. L. Gentry
Cheboygan
Park, Jacobs
Ballard
Co.,
Montgomery
Barnstable
Co.,
near Minneopa
(MIN). Clay Co., Glyndon,
s.n. (MIN). Douglas
& Norby
Cumberland
(MICH).
049
Stephens
Ungar
Shawnee
141 (UWL).
(US). St. Clair Co., Algonac,
s.n. (MICH).-MINNESOTA:
Co.,
Republic
Co., 43 mi E of Snokomo,
(MO).-MARYLAND:
s.n. (NEBC). Middlesex
Coles
Osborne
Wetter
Salt Marsh,
8 mi NW
& Fuller
s.n. (MICH).-MICHIGAN:
1910
(KANU).
Co., Manhattan,
Co., Big
Co.,
(NY).-MASSACHUSErTS:
Ferginnes,
Sabetha,
Pottawatomie
(KANU, NY).
40, Norby
Ballard
Fuller
1917
31855
Co.,
(KANU).
87750
Riley
1105
(UNCC).-MAINE:
Cole
1898, Hunnewell
5980
of South
02243
1910, Bicknell
(MIN). Blue
Eye, Sheldon
of Muldraugh,
Meade
1 mi E of Bennington,
Henry Co., 0.5 mi N of Berea Church,
Co., Union,
Co., Chaney
park, 1917, Billington
Brown
Snelling
(MICH).
s.n. (MIN). Mason
trami Co., Bemidji,
(UNCC).
Munz
Co., Magnolia,
Co., Nantucket,
1893, Durand
1067
204
(KANU).
(KANU). Wabaunsee
(KANU).-KENTUCKY:
Lake, Forrester
from Kensington,
(GH). Essex
1902
s.n. (US). Knox
(POM). Nantucket
16128
Richard
(KANU).
Saline Co., 3.5 mi NE of Bavaria,
(UC). Stafford
188
4545
Holland
Stephens
(KANU).
of Scott City, Harms
Co., Weber
of 31W on 1638 W
near Barkley
670
(KANU).
2.5 mi
Leaven
(KANU).
(SMU). Nemaha
30470
3.5 mi SW of Arlington,
30462
Johnson Co.,
1mi N of Mantey,
5 mi N &
Co.,
area, McGregor
Co.,
11820
SE of Elk City, McGregor
of Walnut,
Ottawa
0.5 mi S of
Hamilton
Brooks
30594
Barker
14153
jct. of N Smoky Hill River & U.S. Hwy
Lathrop
2 mi W
Co.,
the Lakes,
Co.,
Co., Austin
ter, 1909, Chamberlain
from Munz
11.7 mi N & 0.5 mi W
3 mi
Co.,
(KANU).
Reservoir
Co.,
Sheridan
SMU, UNCC).
Linn Co.,
Grove, McGregor
59232
Co., Kirwin
(KANU).
(KANU,
(KANU).
Montgomery
1912, Wilson
Co.,
(KANU).
S of Lakin, McGregor
3 mi N & 0.5 mi E of Frankfort,
(KANU).
(MO). Reno
1mi
Co.,
Brooks
Co.,
Neosho
14992
McGregor
Kearny
et al. s.n.
of Fall River,
3 mi E & 2 mi N of Holton,
Jackson Co.,
NY).
63669
Stephens
(KANU).
1913, Marconey
2 mi S & S mi W
Elk Co.,
1.5 mi E of Neal,
1mi N of Saunders, McGregor
Stanton Co.,
son Co.,
60140
Volle 652
Co., Orchard,
Co., Cowley
Graham
Co.,
(KANU).
50887
Finney Co.,
5 mi SW of Stockton, McGregor
Co.,
Cowley
Stephens
(KANU).
Co., 0.5 mi E & 0.5 mi N of the SW corner of county, Morley
(MO). Rooks
of Opolis,
(KANU).
Marshall
Phillips
Wetter
31317
McGregor
82827
S of Matfield
58965
(GH, KANU).
(KANU).
4 mi
Co.,
59898
1mi N & 0.3 mi E of Covert,
mi W
Stephens
4 mi E & 0.25 mi S of Council
Co.,
Chase
& Bare 338 (KANU).
Doniphan
Greenwood
State Park, Horr
Co.
s.n. (KANU).
S of Ft. Scott, Stephens
of Melrose,
0.5 mi W
(KANU).
1mi E of Wilson,
2.5 mi S & 2 mi
Co.,
& McGregor
Stephens
1001
1011 (KANU).
McGregor
1902,
Bluff, Mor
Clark Co., SE corner of Clark State Lake,
(KANU).
Co.,
Allen
(KANU).
59048
4 mi W & 2.5 mi N of Tongonoxie,
Co.,
son & Robertson
88108
3 mi NW
Co., Kalena,
Co., N of Sergeant's
3.5 mi
Co.,
(ISC,
765 (ISC).
Monson
s.n. (ISC). Story Co.,
1946, Morr
& 4 mi N
24024
Crawford
5 mi SW of McLouth,
Co.,
S of Stanley, McGregor
(KANU).
(KANU).
1929,
8655
Twp., Hayden
8 mi S of Junction City, Stephens
Co.,
Co.,
& Miller
Rydberg
Jefferson
(KANU).
Ellsworth
(KANU).
Hill City, Stephens
S of Kendall,
2 mi E of Kinsley,
Co.,
Co., E of Sutherland,
(ISC).-KANSAS:
Bourbon
Stephens
of Miltonvale,
Stephens
2929
3 mi W
(KANU).
1301
s.n. (F). Washington
S of Muscotah,
(KANU).
Co.,
Co., 2.2 mi W
Co., 2 mi W of Abilene,
Edwards
Brooks
Cherokee
(KANU).
(MO). Cloud
(KANU).
Co.,
30534
S & 5.5 mi E of Elbing,
State Park, McGregor
Dickinson
worth
1mi
65
Morrill
of Des Moines,
s.n. (ISC). Woodbury
Monson
Co.,
Co., 0.5 mi SW of St. Frances, McGregor
26562
County
Atchison
(KANU).
1mi
Co.,
Stephens
Cheyenne
Raven
476
1903, Oleson
s.n. (MO).
& Blosser
Co., SW of Glenwood,
1927, Shimeck
1911, Shimek
Junction,
1mi N of Northwood,
Co.,
E of Moran,
Blosser
s.n. (ISC). O'Brien
2 mi E of Walcort,
Co., Afton
near Patterson,
Co.,
(ISC). Polk Co., W
1368
Jones Co., Hale
(WVA).
1892, Bush
s.n. (MO). Palo Alto Co., Nevada
Prairie, Brotherson
(ISC). Scott Co.,
810
(ISC). Lee Co.,
s.n. (ISC). Mills
1927, Shimeck
1890, Barkley
(ISC). Union
s.n. (ISC). Webster
Arnold
1949, Moorman
Twp.,
(BH, IND, POM). Harrison
s.n. (ISC). Ida Co., 2 mi NW
Easterly
4432
(ISC). Madison
3621
of Muscatine,
Co., Kalsow
of Homestead,
Co., Lu Verue, Blumer
State Park, Hayden
17532
1929, Pummel
Co., Cresco,
(ISC). Iowa Co., W
3622
s.n. (ISC). Page Co., Coin,
Shimeck
Howard
(UNCC).
149 (ISC). Kossuth/Humboldt
Lyon Co., Gitche Manito
20 mi S of Iowa Falls, Munz
(BH). Hardin Co.,
Fay 3518
169
1mi
River
S of Fort
(MIN). Freeborn
Co.,
1
SYSTEMATICBOTANYMONOGRAPHS
170
5.6 mi W
of Oakland,
(MIN). Houston
Kandiyohi
(MIN). Morrison
3 mi NW
Co.,
of Monticello,
s.n. (POM).
TENN, WIS).
Madison
of Birch Tree, Munz
Wright
2 mi W
Co.,
Busby,
E of Flaxville,
(KANU).
Rosebud,
Stephens
Stephens
69343
612
(KANU).
17535
24962
1578
Co., Rydberg
1573
(US). Hamilton
(NY). Howard
s.n. (ND, TENN,
Hapeman
UC).
Seward
Stephens
6339
(KANU). Webster
roll Co., Conway,
1927,
(PENN). Mercer
POM). Warren
Ferry, Eames
1.25 mi
& Wiegand
Hempstead,
Ferguson
Co., Valley
10497
Co., Mandan,
Sarvis
Sourris Wildlife
Co.,
Fourmile
DAO,
Creek,
DS, MU,
near Oxford,
s.n.
Allegheny
(GH).-OKLAHOMA:
ground,
Johnson
& Goodman
tle, Pearce
600
967
Greene
UC).
1934, Hutchinson
unglaciated
Refuge,
221
(CU).
Plateau,
Adair Co.,
(OKL). Hughes
& Waters
Co., Clifton,
Co.,
Long
Russia,
Silberhorn
(OKL). Cleveland
(OKL, UNCC).
1785
of Stilwell,
Co.,
616
10504
Seiler
1040
SMU);
(NY). Hamilton
MacDaniels
(UNCC).
Stephens
1mi
Cleverdon
S mi E of Calvin, Munz
2479
27
of Robinson,
Airy
(CU). Morgan
Co.,
28026
(KANU).
S of Hollywood
Braceville
Corner
DAKOTA:
1189
1mi W &
Co.,
(KANU).
Butler
Cobb
Forest
Emmons
Co., Devil's
61497
Co.,
84 (COLO,
Park, Cincinnati,
Twp.,
York
Twp.,
flora of
1917, Rood
Bryan Co., Kansas
3 (OKLA). Grady Co.,
Island, W
Williams
Sioux
Stephens
Co.,
(BH,
Co., Fisher's
(KANU, MO).-OHIO:
Trumbell
13573
Co., Long
near Collinsville,
3938
Co., N of King's
s.n. (GH, US).
(KANU).
Co., Mt.
14218
(CU).-NORTH
14 mi S of Jamestown,
(MICH,
Munz
1911, s.c. 173 (RM). Ramsey
Dam, Ward
143
7 mi E &
Craig Co., Vinita,
Co.,
Co.,
Lake Darling
Wehmeyer
10 mi W
of Lisbon,
Stutsman
s.n. (NEBC).
Inlet, Fender
(CU). Suffolk
1905, Lunell
Co., 6 mi S & 1.5 mi W
Co., Stump Lake,
5 mi NW
Co.,
below
Lorain
Ithaca, Wiegand
Co., Leeds,
HAMPSHIRE: Car
Brunswick,
10500
58516
24 mi S of Rushville,
Co.,
(CU). Nassau
Richard
Stephens
YORK: Cayuga
Lake, Gershoy
Heights,
Kidder
(KANU).
Co., Townsends
Smith 1014
of Cayuta
1932,
s.n. (MIN, MO).
7800
(ISC).-NEw
Co., New
near Mullen,
Co.,
Co., Minden,
1926, Batchelder
s.n. (PH). Cape May
s.n. (RM). Benson
(KANU).
Brooks
411270
Co., Rock
(BH, IND, POM). Merrick
Sheridan
Co., Park Hill,
Co., W
95 (US). Nelson
33678
Tolstead
S of
& Bare
s.n. (MIN). Grant
1899, Hedgcock
Co., Peterborough,
3 mi
Dundy
Saline Co., 2 mi E of Dorchester,
(PH).-NEW
(KANU, MO).
s.n. (NY). Ransom
1891, Waldron
15 mi N of Fort Yates, Brooks
Ward
1246
Co., Ruderal,
49885
Co., Cayuga
1896, Rerrivue
17530
(SMU, TEX, UWL).
Schaeffer
(CU). Chemung
Munz
11 mi N of Osceola,
s.n. (WVA). Middlesex
(NY). Schuyler
(GH). Tompkins
City,
4355
(DS). Kearney
of Landers,
Co.,
Co., Gothenburg,
(KANU).
15808
68485
1mi E of
McGregor
(F, GH). Hooker
(KANU).
1900, Brown
1903, Logan
Co., 2 mi N of Linton, Williams
(MO). Morton
Koch
S of Port Murray,
1919
Island, St. John 2830
58066
of Valentine,
60712
15021
of Sutherland,
Polk Co.,
3 mi W
Arthur
(UWM).
Kiener
Stephens
s.n. (US). Hillsborough
Co., Medford,
Co., Trenton,
Co.,
Stephens
Co.,
198
1893, Laybourue
Co., Farmers Creek Valley,
Johnson
NEW JERSEY: Burlington
Lake,
(CAS, GH).
6 mi E of Seward,
Co.,
of Aurora,
5 mi W
Co.,
116 (MO). Treasure
of
s.n. (NY).
Stephens
1mi S &
Co., Franklin,
(BH, IND, POM). Nuckolls
5250
1.5 mi S & 2 mi E of Rulo,
(KANU).
Barnes
Lincoln
17534
City, Bates
(ISC). Franklin
6 mi W
Co.,
Co.,
Stephens
(MO).
Co., 2 mi
1883, Scribner
(BH, IND, POM). Dawson
17529
Co.,
15071
Daniels
(KANU).
Rosebud
of Valley,
5.5 mi SE of Dannebrog,
Co.,
2 mi E of Clarks, Munz
Otol Co., Nebraska
son Co.,
411271
Tolstead
Grounds,
Pole, Munz
3.5 mi NW
Co.,
NY,
Co., N of Hanni
SW of Havre,
20 mi S & 6 mi W
Cherry Co.,
Co., 4 mi E of Lodge
Cheyenne
9 mi
Warnecke
Co., Royal,
1926,
Big Horn Co., 0.25 mi W
Teton River,
(KANU).
Larsen
Co., Westby,
(DS, KANU).
69641
Stephens
Co., Oakville,
159 (MIN). Shannon
68345
(RM). Hill Co.,
14955
Co., Willard,
F, ISC, LAM, MO,
Davis
Stephens
Spring Creek,
8019
Antelope
(BH, IND, POM). Douglas
Creek Recreation
Rydberg
Sheridan
(KANU).
& Brooks
Co.,
of Broadus,
(KANU).-NEBRASKA:
Arthur, Stephens
Munz
69169
Dawson
of Zurich,
Cole
Clark
Steyermark
1mi S of Irena, Steyermark
Co.,
Co.,
Creek,
Greene
(DS). Marion
(BH, IND, NY, POM).-MoNTANA:
Park, Osterhout
35 mi W
of Amity,
Jefferson
(ISC). Ralls Co., Oakwood,
Co., 0.25 mi W
67944
of Glacier
River Co.,
Powder
13551
Munz
Blaine
Stephens
351
Steams
IND, NY, POM).
(DAO,
20535
s.n.
s.n.
(UNCC). Wright
72 at Yellow
UNCC).
24661
line, Raven
Co.
(BH, IND, NY, POM). Worth
(KANU).
Co., E entrance
(KANU).
13546
of Norwood,
69507
Stephens
Glacier
67 atWayne
s.n. (RM). Pettis Co., Riggins
bal, 1913, Davis
7 mi W
on U.S. Hwy
2 mi W
(MO, NY, US).
Steyermark
160
(BH, DS,
(CAN, MO,
21690
(BH, IND, POM).
RSA).
96, Lindayen
Co., Hwy
Co.,
26265
11504
City,
(MIN).
& Skimmer
17538
Munz
13555
Munz
MacMillan
(DS, IND, MICH,
Tichomingo
(F). DeKalb
Bush
1 mi E of Bates
1372
2.7 mi E of Hwy
30 & 47, Raven
Jackson Co., Vale,
Co.,
Co.,
68865
Steyernark
jct. of Hwys
s.n. (MO). Lafayette
5 mi N of Northfield,
7 mi S of Monett,
731101
315
1912, Chandonnet
Co., Perham,
Polk Co., Crookston,
(MIN).-MISSISSIPPI:
Wertman
of Isabella Lake, Lakela & Davidson
& Grayson
Co.,
Ballard
Co., Fumbrota,
estate, Westside,
79 (MIN). Ottertail
16, Bartlett
Barry Co.,
St. Clair,
Co.,
1919, Blankinship
Mathias
Co.,
Smith 878
SE of St. Francisville,
(MO). Franklin
Trail, W
s.n. (MIN).
s.n. (DAO). Washington
(TENN).-MISSOURI:
9.5 mi
Co.,
along U.S.
Goodhue
RSA).
Isanti Co., Cornea's
(MIN). Rice Co.,
Taylor 864
of Luverne
1933, Kuehne
man 50314
1961, Morea
S of Hinkley,
11.3 mi W
Collegeville,
(DS, MICH,
(MO).
s.n. (US). Lake Co., Tomahawk
(MIN). Pope Co., Glenwood,
Rock Co.,
2179
15 mi SE of Little Falls, Siemers
Co.,
Pine Co.,
1384
& Grayson
& Leykom
1892, Frost
Co.,
(PENN).
Bartlett
Co., Ziegler
VOLUME 50
s.n.
Creek Camp
near Little River,
Lawson
6 mi E & 2.7 mi N of Tut
(BH, IND, POM). Kingfisher
Co.,
0.5 mi S
1997
OENOTHERA
of Dover,
Waterfall
Co.,
Byers
75 (TEX). LeFlore
Co., Heavener,
Munz
8478
(GH, OKL,
TEX). Mayes
Co.,
3 mi E of Muskogee,
Ottawa
Little 269
near Miami,
Co.,
Haileyville,
Stevens
13570
Munz
OKLA,
(OKL). Nowata
2253
601
of Spring Valley,
Co., W
11449
Pretz
of Brookings,
Johnson
Visher 4091
Harding
Co., Long
8082
Stephens
& Brooks
2198
(DS, KANU).
41580
(F, NY).
(TENN).
boro, DeSelm
Reverchon
689
2758
Mecklenburg
(WVA). Hampshire
Botanical
Mick
191
(WIS). Ashland
(WIS). Burnett
Baru
Smith 7472
s.n. (WIS). Green
Anderson
Boy
Wickham
Lake,
1925, Allen
Genoa
City, Linderud
17 in Fremont
Sorensen
Door
Campbell
83
Hwy
Stephens
Platte Co., Uva, Nelson
RSA). Weston
70941
8577
(KANU).
Co.,
Degener
313
Sublis
Calingasta,
Arbesser
745 (CORD, MO).-MENDOZA:
Fabris
Lake,
Monument,
16189
Pamonima
(CTES,
LP, M). Austria.
s.n. (GZU).-OBEROSTERREICH:
Wels,
1977, Forstner
s.n. (GZU).-WIEN:
(BR). China.
704 (PE).-JILIN:
Breitenlee,
HEILONGJIANG: Yilan
Long White
HOMORAVSKY: Brno
Mt.,
[Briinn],
Chien
426
1946, Rechinger
Co., Zhang
1899
6730
s.n.
Co.,
15 (GH).
1956,
1.8 mi SE of Rome,
Pope
Co., Argyle,
7
s.d.,
Co., Tembina,
s.n. (WIS). Rock Co.,
5 mi E
(MIL). Walworth
Co.,
12 mi W
Sorensen
Wagner
2822
4035
(WIS). Wood
(MO). Goshen
Stephens
Bartlett
Co.,
70187
of
sect.
Co.,
Co.,
4
(KANU).
& Grayson
1308
(DS,
(GH, POM, NY).
80, near Sta.
C6RDOBA: Ruta 9, Km
near La Paz, Leal 8536
(MERL).-SAN
KARNTEN: WeiBenstein
JUAN:
near Villach,
s.n. (W).-STEIERMARK:
Liebenau
s.n. (W). Belgium.
BRABANT: Brussels,
Cangbei
Tsintao,
Tsui 437
(PR).-VYCHODOCESKY:
Mt.,
(PE). Czech
Rychnov,
1912,
near Graz,
(PE).-LIAONING:
(PE).-SHANDONG:
Jehl(k & Rostaniski
Bay, Greenman
14.5 mi E of Lance Creek,
2391
1925, Arbesser
r6e 1303
& Marchionni
Villa
1952, Byle
(BH, IND, POM). Waupaca
SPECIMENS EXAMINED FROM OUTSIDE NATURAL AREA. Argentina.
Marfa,
Twp.,
s.n. (WIS). Sawyer Co., N shore Sand
Co., 6.9 mi N of Worland,
& Peiler
Co., 6
s.n. (WIS). Crawford
Co.,
s.n.
1929, WVU
of Briggsville,
s.n. (WIS). Marinette
17520
Co., Virginia
(ARIZ, RM). Washakie
Co., Newcastle,
6 mi NW
s.n. (WIS). Lafayette
1925, Davis
vic. Lincoln
Co.,
of Elk
1953, Boggs
School,
Suamico
Inn, Ellison
Jefferson
State Park, Fuller
Niobrora
(BH, IND).
s.n. (VWVA).Morgan
Turner
22, 1965, Kust
Spooner, Munz
(OSH). Waushara
Albany
14250
Co., 4 mi W
near Miller
Co.,
s.n.
s.d., Carpenter
Rockingham
Co.,
1988,
1966, Huempfner
72 (WIS). St. Croix Co., Woodville,
Co.,
Brown
1961, Longenecker
Co., Andral
Springs,
Irwin s.n. (WIS). Green Co., N of Monticello,
14 mi S of Montello
114 (WIS). Washburn
Tarrant Co.,
Adams
Poynette,
s.n. (WIS).
near Reedsville,
Co.,
(WIS).-WYOMING:
mi N of Meriden,
1966,
Co., Dallas,
TEX).
1972, White
Co., near Garrett Bay
Point,
Dallas
S of Gassaway,
Co.,
Hollow,
(BH, POM).
Co.,
1925, Martin
s.n. (WIS). Sheboygan
Twp.,
2538
17542
Scout Camp,
Co.,
s.n. (WIS). Marquette
of Jamesville,
MICH,
Rock
s.n. (WIS). Manitowoc
1819, Sykes
Munz
(UWM, WIS).
Lincoln
s.n. (TEX).-WISCONSIN:
(WIS). Columbia
Lake Co., Dalton,
Juneau Co., Castle
(WVA).
(DS, MIN).
Lake, Munz
(UNCC).
et
(FSU, SMU,
atMountain
Sugar Creek
1mi up Melville
along Lake Superior, Wisconsin
Co.,
(WIS).
1724
10722
27802
(OKL).
Park, Heidecker
County
10.4 mi ESE of Murfrees
Co.,
Co., Middletown
4002
Co.,
1062c
Frye
Co., Ashland,
Co., Mauritz
du Chien,
Prairie
Douglas
Rock,
1939, Strausbaugh
Springs,
Seaman
VIRGINIA: Braxton
s.n. (MIN). Logan Co.,
Expedition
mi E of Berkeley
Rutland
seed col. by McNeil
on island in Lake Gaston,
Co., Hanging
Palmer
Knee Creek,
13136
Co., Cove Creek, Hesler
Campbell
Shinners
Co., Pactola Lake,
Todd Co., Over
Rutherford
(TENN).
Co.,
(KANU).
(CAN, MIN,
Co., Wounded
Co., Union
(MIN).-TENNESSEE:
(KANU).
Deuel
2738
160
157 (US). Pennington
Union
SE
Co., 22.5 mi N of Howes,
(RM). Shannon
(KANU).
Co., Palestine,
cult. from 1935
(WVA).-WEST
626
7827
Harms
Rydberg
s.n. (NY). Meade
mi
(MIN). Brookings
Stephens
Co., Deadwood,
Creek, Carr
River, Clark
1-1.25
Co.,
384
of
Se
(OKL).
s.n. (MICH, NY).
Lawrence
SE of Bellevue,
3 mi
1940, Johnson
& Swanson
s.n. (MIN, US).-VERMONT:
Co.,
Co., Savannah
s.c. 107
ton, 1918,
Anderson
Moore
Co., Mastin,
1mi S of Alexandria,
34102
2170
(UC). Lehigh
SMU).
2 mi W
Co.,
Hopkins
26 mi S of Camp Crook,
et al. 9593
Sharp
Co.,
NR-2297
Lake,
Visher
86 (OKLA,
Pittsburg
of Lexington,
75 (MIN, MT, POM, UC).
694
Moyer
(TENN).-TEXAS:
Giles
(LA).-VIRGINIA:
& Brooks
Co., Emville,
1902, Reverchon
Handley,
(RM, WTU).
Co., N of Idabel,
(OKLA). Muskogee
Bennett
US).
7 mi W &
& Todd Co., Horse
Stephens
(US). Montague
Co.,
Ricksecker
2614
of Sand Springs, N of Arkansas
Russell
DAKOTA: Bennett
Perkins Co., Bixby,
Co., Mobridge,
Chester
Rock,
1969, Swanson
Spink Co., Redfield,
131 (KANU). Walworth
Tulsa Co., W
Slippery
Hanson
266
Mellette
(KANU).
2 mi E of Colome,
Tripp Co.,
al. 2278
Visher
Lyman Co., near Kennebec,
Stephens
Visher
Pine Hills,
12 mi NW
Co.,
s.n. (MIN). Butte Co.,
(KANU).
Wallis
NY, OKL, OKLA,
(MO). Custer Co., Sylvan
8 mi S of Clear Lake, Croat 2588
MO, WIS).
MO,
(OKLA).
Co.,
(MICH).-SOUTH
1903,
Clay Co., Vermillion,
2787
Butler
(OKLA).-PENNSYLVANIA:
of Peggs,
6 mi E & 3 mi N of Wann,
Co.,
Pottawattomie
quoyah Co., 0.5 mi SE of Gore, Wallis
(BH, IND, POM). McCurtain
1.8 mi NW
(DS, GH, MIN,
(IND, POM).
13568
171
Lawal
1400 m, Qian
Republic.
Jehli'k 6736
Ji
(PR).
ZAPADCESKY:
Plzeri [Pilsen], Jehlik 6256 (PR).Denmark. Sjaelland:Copenhagen, 1922,Andersen s.n. (C).
Finland.
UUSIMAA:
(RSA).-GIRONDE:
Helsinki,
Bassens,
VAR: St. Cassien-des-Bois,
DENBURG: Senftenberg,
1951,
Buchon
Gavelle
1980,
Gutte
Haysen
2400
5804
s.n.
(LD).
France.
(BC, DS, P).-HAuT-RHIN:
(COI). Germany.
& Jentsch
s.n.
ALLIER: Moulins,
Ottmarsheim,
BAYERN: Schongau,
(LZ).-BREMEN:
railway
1952, Deschatres
Rastetter
1971, Dorr
3886
s.n.
(BR, L).
s.n. (M).-BRAN
station Bremen-Weser,
1980,
SYSTEMATICBOTANYMONOGRAPHS
172
s.n. (BREM).-HAMBURG:
Jehlik
2005
Wagenitz
s.n. (BP).-CSONGRAD:
Thaisz
(BP).-KISKUN:
Bacsa,
MACHAL PRADESH: Kangra
Prov. Ancona:
Viareggio,
Fiori
1315
13426
Hyogo
Kern & Reichgelt
12182
1922, Steenis
way.
1923, Stiirmer
OSLO: Oslo,
vika, Lyche
[Breslau],
& 29269
15477
434
Rostaniski
KRAY: Smolenskoye,
2195
BP, BR, C, DS,
FI, HBG,
Merwe
Kalmar,
(LD).-KRONOBERG:
Botanical
2380
Gdanisk,
Scotland.
Kingdom.
10191
1836
s.n.
(LD). Malmo,
Fushiebridge,
(as 0. biennis
1762
France.
salicifolia).
(as Onagra
(C, Fl, FR, M,
5b. Oenothera
villosa
and Thompson
Germany.
(as 0. erosa).
s.n., cult. DUSS-84-140
(MO)
4529,
cult. DUSS-82-0382
Hoch
s.n.*, cult. DUSS-76-015
4547,
W
Co., Fountain
Co.,
Geneve,
1826
(NY).
(MO)
cult. DUSS-82-0331
of Logan,
(MO)
1mi S of Woods
Co.,
Lakes
Natl.
DAO,
GH, NY); Waterton
combe, Dixon
Carmangay,
Park, Blais
1209
Groh
Dumais
865
2254
Hat,
coun
s.n. (NA); Little
(ALTA);
Smoky
1914, Fyles
River
Fish Lake
10644
Lake, Dumais
(ALTA); Bow
River,
s.n. (DAO);
at the Goodwin
1975,
Hwy,
phenotype
Nest
Pass,
Boivin
Brook,
1785
Park, SW of Drumheller,
& Watson
along Columbia
(MO) (0 14).
(MO)
s.n. (CAN,
East Gate, Waterton
& Perron
Breitung
12319
15782
Dumais
473
5827
1748
La
just W
(ALTA);
of
John
(UC); Peace River, Grande
Island, Calgary, Mc
(DAO); Athabasca
McPherson
(ALTA,
(F, RSA);
(DAO); Little Bow River,
Goddard
(014).
in the notes).
1938, Anderson
S side of St. Patrick's
708
cult. DUSS
discussed
(ALTA, DAO);
near Carseland,
(DAO);
1803,
Co., S of Vantage
du Vieux,
& Watson
Schultz
Bird
(MO) (014).
cult. DUSS-82-0391
132
of Blakiston
Ferry, McNary
(CAN, GH); Red Deer, Malte
Pine,
near Sumpter, Hoch
728, cult. DUSS-84-136
Bird
Sarcee Reserve,
1601
City, Wagner
and Yellow
10 mi E of Salt Lake City, ca. 2160
Yakima
2618,
Riviere
SE of Calgary
Park, Jenkins
Provincial
View
s.c. s.n. (ALTA); Crows
(CAN); Macleod,
(014).
1973, Stubbe
(MO), DUSS-77-0207
an odd biennis-like
Park, near crossing
50 to Canon
McCall
Baker Co.,
Range,
2280 m, Hammel
indicates
(MO)
dis
Fraser
Box Elder Co., Bear River Migratory
on Valley
10 mi SSE of Vermilion,
Lakes Natl.
(DAO); Banff Natl.
9212
River, Macoun
& Hagy
5821
Calla
Milk
1916,
s.n. (CAS);
(CU); SE end of Buck
son's Garden, Medicine
Prairie,
Edmonton,
1918, Bickle
S of Hwy
(MO), DUSS-76-095
Landing,
between
(MO), DUSS-84-103
between
(014).-UTAH:
(014).-WASHINGTON:
REPRESENTATIVE SPECIMENS (an asterisk
ALBERTA: Campus,
1mi
Salt Lake Co., Wasatch
(014).
cult. DUSS-82-0385
p.p.
phenotype
Valley
near Estes Park, ca. 2300 m,
River
4695,
Calive,
& Wynberg,
an odd biennis-like
12 in Botanie
(MO) (014).-OREGON:
2.1 mi E of "boat landing"
(MO)
indicates
Creek,
Salmon
shore of river above Priest Papias Dam, Mastrogiuseppe
Canada.
11 Sep
(as 0. kunthiana),
(from Rodebosch
Park, headquarters
River, W
MT);
Glamorgan,
1820
jct. of Hwy
m, Arnow
WYOMING: Albany
s.n.
1930, Probst
1832, Vahl ex herb. Fischer
(Fl, herb. Webb)
cult. DUSS-82-0389,
Valley
(014).
(MO), DUSS-84-127
(MO), DUSS-77-0208
1mi above
(MO), DUSS-84-129
et al. 16088,
Co., marshes
76-094
Pueblo
(014).
(MO)
926, cult. DUSS-76-012
Cache
(LD).-SODERMAN
Hamburg,
Switzerland.
BRITISH COLUMBIA:
rivers at Lytton, Wagner
Holmgren
s.n.
s.n.
s.n.
Lange
subsp. strigosa.
COLORADO: Larimer Co., Rocky Mtn Natl.
Refuge,
1925,
(BM).-WALES:
Copenhagen,
1836 & 1837
Paris, Aug
lehmanniana).
P, SAM)
in the notes). Canada.
U.S.A.
s.n.
Lousley
SPECIMENS EXAMINED FROM CULTIVATED PLANTS (an asterisk
cussed
Kristianstad,
SOLOTHURN: Oberdorf,
1964,
Sweden.
(PRE).
1948, Andersson
Halland,
1929, Norrman
(BC,
(PR). South Africa.
s.n.
Strey
1452
Smejkal
s.n. (K).
(Fl, herb. Webb)
Cape),
KRAY: Shko
?turovo,
1961,
(LD).-KRISTIANSTAD:
ALTAYSKIY
(DS).-PRIMORSKIu
Jehlik 5354
Tygerport,
s.n. (LD). Switzerland.
Lothian,
Bu
WROCLAW: Wroclaw
1942, Tollinen
Zamky,
s.n.
Johnsson
s.n. (L). Nor
(Viipuri),
s.n. (LD).-HALLAND:
SPECIMENS CULTIVATED IN BOTANICAL GARDENS. Denmark.
(GOET)
Vyborg
Trebisov,
(PRE).-TRANSVAAL:
1933, Hakanson
s.n. (L).-NOORDBRA
s.n. (M). Russia.
1928, Mayer
Skvortsov
Garden,
Frederiksen
Pref., Kat
Netherlands.
(KYO).
Poland.
(0).
VAPADOSLOVENSKY: Nove
1946, Fries
1931,
Almhult,
1926, Melville
Cardiff,
1946,
Hiistholmen,
(G). United
24567
Ouren
L, LD, M, MA).-VYCHODOSLOVENSKY:
CAPE PROVINCE: Knysna,
s.n. (FI). Japan.
Kyoto
1939, de Bruyn
Schiedam,
s.n. (A, NA).-KARELIA:
& Zwierewa
GOTEBORG OCH BOHUS: Hultmansholme,
(K). Italy. REGION
Jel0y Isle, 1973, Suirlye s.n. (O).-S0R-TRONDELAG:
Porsgrunn,
(DS). Slovakia.
10445
1920, Kloos
Weert,
s.n.
India. HI
TUSCANY. Prov. Lucca:
(MAK).-HONSHU:
Fukuoka
Havendijk,
15539
1944, Minio
Alberoni,
City,
(L).-LIMBURG:
s.n. (O).-0STFOLD:
in University
spont.
Schreter
LAND: Nacka,
Kobe
Pref.,
(0).-TELEMARK:
Lido,
& Sako 27019
(FH,LD, RSA, WRSL).-GDATNSK:
tovo-Nakhodka,
(LD).-KALMAR:
s.n. (FI).-REGION
s.n. (L).-ZUIDHOLLAND:
1957, Kuminova
s.n. (L).-MOSKVA:
and Spiti dist., Bor
1923,
1938, Polgar
s.n. (BP).
1936, Polgar
S of Lahul
Hatusima
Airagun,
Esztergeto,
1952, Pierfaoli
VENETO. Prov. Venezia:
(MAK);
BANT: Ginneken,
Esztergom,
Isle Baltrum,
BUDAPEST: Budapest,
s.n. (BP).-GYOR-SOPRON:
N of Nagar,
di Senigallia,
Pref., Kamo-cho,
GELDERLAND: Nijmegen,
70 (HBG).-NIEDERSACHSEN:
(LZ). Hungary.
s.n. (BP).-KoMAROM:
dist., Manali
Spiaggia
Murata
Tauchritz,
1943, Polgar
Szeged,
(K).-REGION
KYUSHU: Kagoshima
sure, Kyoto-city,
Lausitz,
1933, Polgar
MARCHES.
Rostan'ski
Otto 4995
Schneckenburg-Allee,
(GOET).-SACHSEN:
VOLUME 50
485b
(CAN); Bow
Plains,
(ALTA);
1872, Ma
Sage Creek,
Valley, Moodie
75 (F,
1997
NY);
OENOTHERA
Rosedale,
Red Deer
Fort Assinaboine,
ton, Moss
1870
SW
(ALTA);
Valley, Moodie
(ALTA, DAO,
of Edson,
10597
Oldenburg
(ORE);
Park, Scammon
Reservation,
1894, Shortt
River, N Pincher,
1mi W
7853
(RSA);
de Vries
Samos,
2087
Beamish
630211
ner ferry, Bird
rd to Alexis
4198
Hill
s.n. (UBC); Botanie
4507
Valley,
8393
1914, Malte
Prairie,
Sandall
(DAO); Coquitlam,
nipeg, Bernard
North West
Savona,
5529
Lake, Dugle
lier,Macoun
9040
1412
& Scoggan
Pipestone
ISC); 4 mi E,
River
3409
Scoggan
Lake,
win
(CAN);
53-61
(DAO,
1953, Mosquin
(MIN),
3644
3297
Scoggan
140 mi NW
Gypsumville,
8 mi
Lake,
Buffalo
Bay
s.n.
s.n. (CAN); Fort El
to Bird Lake,
Gillam,
Schofield
3229
Scoggan
(CAN);
45 mi N of Lake Winnipeg,
Lake,
9460
Scoggan
s.n. (TRT);
1935, Groh
2 mi S of Hwy
(CAN); Duck Mtn,
of Winnipeg,
between
(DAO); Churchill,
S of Pipestone
695.5
Win
Matlock,
s.n. (CAN); Red Deer
1951, Love & Love
Cross
1912, Taylor
Warren
1873, Burgess
N of Lake Winnipeg,
River,
Lake,
Lake,
s.n. (DAO);
rd to Pinaiva,
(CAN);
4114
on Okanagan
s.n. (DAO); Arborg,
R1555
Ronald
Rapids,
(CAN); Minago
off N end of Lake Winnipeg,
campus,
& Wood
end of Kamloops
(UBC).-MANITOBA:
1930, Dudley
9039
Sica
s.n. (V); New West
1911, Criddle
1922, Groh
12,
N of Gray
(CAN);
Kelowna,
(UBC); Turtle Mtn,
s.n. (DAO);
1mi N of Morden,
Scoggan
591
ISC); Aweme,
(OKL); Birtle,
at McCall
(NY, SMU, UNCC);
Siding, Wilson
(ISC); Stoney Mtn,
35 mi N of Lake Winnipeg,
Playgreen
7906
694
64605
1937, C. T s.n. (V); Popcum,
Chilliwack,
47 (MIN); Fort Garry, University
(DAO); Nelson
Lake,
Sand Bay,
4409
(CAN); Brokenhead,
(DAO,
1350
Chunys
River, Crook
& Dugle
Guess
et al. 3818
(UBC);
1896,
jct. with Hwy
of Vernon, Mulligan
(VPI); W
1174
on
1955, Krajina
Sproat, Macoun
1917, Newcombe
Straley
1961, F Vr cc. 610211
s.n. (UBC); Tappen's
Lake,
(CAN);
Macoun
Smith 3 (DAO); Trout Creek Point
(DAO); Big Eddy, The Pas, Bryant
S of Moose
(DAO); Minnedosa,
Rogge
1658
Straley
above
9017
s.n. (V); Lillooet,
s.n.
(UBC); Pitt Mead
UNCC, WTU);
of Cascade,
24.5 mi NNW
s.n. (CAN);
(UBC); Creston,
S Okanagan,
24 mi S of Overflowing
River,
& Moose
1918, Malte
s.n. (UBC); N of Huntingdon,
1915, Welch
Angle,
(CAN, US); W
4508
1mi
(SMU, UBC,
Macoun
Kawloops,
64604
903
Tran
s.n. (DAO); Nelson,
Island near Agassiz,
above Lytton,
Krajina
(UBC);
1938, Bucklond
s.n. (UBC); Appledale,
Eastham
1930, Groh
Gardner
area,
S of Lad
(DS); 2 mi E of Redstone
11516
Nelson,
Rivers,
Vancouver,
1919, Newcombe
11160
1937, Stonor
2 mi W of Oliver,
and Thompson
Drive,
Macoun
Bridge,
Fraser Valley,
The Reservation,
(ALTA, UBC);
S of Rossland,
Campground,
s.n. (UBC);
Fraser
between
(MO); Marine
Spences
S of Summerland,
Steelhead
(GH, UBC);
s.n. (CAN); Vernon,
(DAO);
Sumas
minster,
496
Longbeach,
(UBC);
48151
Balfour,
1913, Cormish
107 (UBC); Agassiz,
Falls, Barr
Lake, Pavilion
Brayshaw
NY, US);
Prov.
COLUMBIA:
Shuswap
(UBC); Crown
& Savile
Hills
Cypress
(ALTA).-BRITISH
Kamloops,
(DS); Hedley,
District,
Fodor
Gate,
River,
Turner
s.n. (UBC); Lumby,
(GH, MO,
Calder
Lake,
et al. 20469
(UBC); Kootenay
Krajina
4547
Sheep Creek,
bury 1974
area, Calder
733
Piegan
113 (DAO); Oldman
shore of Blue Lake, Beil 67-07-17
s.n. (UBC);
Brown
side of Osoyoos
s.n. (UBC); Kings
dry slopes, Wagner
mous,
1949, Brayshaw
F9
Raymond
(ALTA);
55 mi SW of Edmonton,
s.n.
et al. 750251
3219
319-63
N Saskatchewan
(RSA); Twin Lakes,
& E.H.M.
1913, Baggs
above Soda Creek,
Beavermouth,
on W
Chilcotin
s.n. (UBC); Hatzic,
Creek, McCalla
zone,
(UBC); Hope,
Eastham
1958, Evans
(CAN);
Island,
s.n. (UBC);
Creek,
Slocan Valley,
ows,
S side of Savary
J.H.W
12 mi E of Slocan Lake, Beamish
1mi S of Osoyoos
(UBC);
1950,
(UBC); Caribou
Beach,
Turner 8400
near Blairmore,
2557
Rd, Sudol
736 (ALTA, DAO),
(ALTA); Ma Me-Ce
60 mi SW of Edmonton,
Lake,
Creek,
1962, Brink
quille,
Turner 2813
Edmonton,
Park, Red Rock
Moss
Newsome
Lake,
Shaw
(ALTA);
N of Edmon
& Macleod,
Cardston
SW Alberta,
Turner
1191
River,
of Lacombe,
at Reesor
between
River,
Lakes Natl.
Sturgeon
(ALTA); Alix, W
near dam
River,
(GH); St. Mary's
Patricia, Moss
(DAO, GH);
2578
Hills,
St. Mary's
(MIN);
3389
NY, UC, US);
922
Moss
Cypress
(ALTA); Fort Saskatchewan,
(DAO);
(UBC); Wilson
(ALTA);
40-44
s.c. s.n. (DAO);
1942,
9856
283
near Battle
Gap, Moss
s.n. (DAO); Waterton
Survey
of C.P.R. Bridge,
Park,
(DS, F, GH, MO,
N of Kinsilla,
PENN);
Moss
(DAO); Waterways,
Jasper Natl.
1086
s.n. (ALTA); Whisky
Jul, Moss
173
Scoggan
Blue Lake,
(CAN,
GH);
3892
Scoggan
Neepawa,
(CAN);
& Bald
Scoggan
10521 (CAN);Virden, Scoggan 11154 (CAN).-NORTHWEST
TERRITORIES:
Athabasca River, Preble & Cary
158
(US).-ONTARIO:
1912, Herriot
Galt,
SASKATCHEWAN: Saskatoon,
Val Marie
Reservoir,
Bird
et al. 9992
Mosquin
River, Breitung
4770
N Battleford,
(DAO);
11440
(DAO); McKague,
1427
(DAO); Patience
Sunny Brow,
s.n. (DAO);
(DAO);
1mi N of Yonker,
Boivin
& Dunbar
1934, Breitung
10439
of Wallwort,
Breitung
(V); Lipton,
1911, Clokey
s.n. (DAO); Manito
Harns
s.n. (TRT); Kindersley,
29 (DAO); Marriott,
1935, H. WM.
16999
Lake,
Grankton
(DAO); Moose
near Ternan's
Lake,
52
(DAO).
(CAN);
S side of Lake
near Souris
River,
Boivin
& Dore
9799
(DAO);
s.n. (DAO);
Lake, McMillan
11221
& Alex
2 mi NE
S of Beauval,
of Cedar
Lake, Baldwin
S of Halbrit,
(OKLA); Weyburn,
Boivin
1 mi N
s.n. (CAN); Meadow
Lake, Child 322
(DAO);
1941, Estate
28 mi
1938, Hutcheson
Jenkins
1328
Lac du Diable,
(DAO); Battlefords,
s.n.
1933, Anderson
Prince
4 mi NE
1452
(DAO);
Albert,
of Nipawin
& Bibbey
1930,
Duck
428
Jenkins
s.n. (DAO, TRT); Assiniboia,
Milk
Lake,
Boivin
&
shore of Saskatchewan
(DAO, GH); Cypress
s.n. (UC); Bjorkdale,
Jaw, Mortlach,
7905
Swift Current Creek, Boivin
Hills
Park, Breitung
1941, Colk s.n. (DAO);
(DAO); Weyburn,
Hudson
s.n. (DAO);
375
1933, Groh
(DAO); Yonkers,
0.5 mi E of Hoosier,
River, Macoun
10644
(ND-G);
174
SYSTEMATICBOTANYMONOGRAPHS
Bredenbury,
Head,
& Herriot
Macoun
Langbank,
(DAO);
1936,
1947,
Apache
Co., White
Canyon,
Whiting
Mtns,
Wickes
s.n. (CAS).
Trinity
Hall
8695
Boulder
Co.,
1mi
26551
(MO). Costilla
Payson
Norby
512
Fremont
(RSA). El Paso Co., Engelmann
1916, Comstock
1934, Stigall
s.n. (COLO).
0.5 mi E of Hesperus,
Brooks
26487
(DS, KANU).
13023
Munz
Shawnee,
mi W
2083
Creek,
18908
Emigrant
Brown
Canyon,
Custer Co.,
mont Co.,
884
15 mi E of Stanley,
Kootenai
WS, WTU).
(US). Minidoka
IND, NY, POM, US, WTU).
Hitchcock
467
(NY). Valley
ley, Smith 1513
Co.,
7 mi
1mi N of jct. of Hwys.
Ely,
Jones
POM). Broadwater
354
Spragg
Co.,
Creek,
between
14561
(BH,
of Virginia
in Blackfoot
(MICH).
Deer,
Valley,
C. L. Hitchcock
Munz
& Grayson
1917, Garnell
on Hwy
Forks, Munz
& Rock Creek,
City, C. L. Hitchcock
17 mi S of Livingston,
lier, Bartlett
in Glenwood,
420
14571
Liberty
15812
1233
C. L. Hitchcock
Powell
Falls, Rogers
Co., Deer
15283
Christ
Lodge,
RSA).
(MIN).
of Haw
1891, Bur
Pope Co.,
6 mi W
Lake,
14555
near
(IND,
Chouteau
Co., Square
1068
(MO, NY, WS).
Co.,
1mi
Granite
from Rock
(WTU). Lake Co., Polson, Munz
11362
(DAO). Madison
Missoula
Co., Cupuyer
Hall
C. L.
5461
(BH, US).
3002
7.5 mi WSW
(MIN).
s.n.
(BH,
Co., Twilight
11573
Silver
(CAN, NY, RSA, WS, WTU).
Creek,
6 mi
Bow Co.,
Park Co.,
5.8 mi W
(UC). Rosebud
Stillwater
Co.,
Co., N of Bonner
NY, PH, POM, RM, WS, WTU).
Pondera
1574
1894, Kirlley
Co., Armstrong
& Rogers
14735
ID, NY,
(BH, DS, MO, NY, PH, POM,
UC, WS, WTU).
(BH, MO,
Co., Bull River, Kaul
17057
1738
15284
DAO,
Co., Minneapolis,
13644
Co., Les Trois Buttes, Mosquin
11457
Davis
S of
(MIN). Fre
8 mi N of Avery,
Clay Co.,
(DS, MONTU,
(DS, GH, NY, RSA,
74-160
5 mi N of Lima, Munz
& Muhlick
& Muhlick
(ID, NY).
14578: Munz
City,
Co.,
(BH, IND, POM, US, WTU).
C. L. Hitchcock
14398
(ID). Owyhee
St. Louis
up Fall
Co.,
Teton Co., Tetonia,
& Massey
Co.,
14726
1mi E of Columbia
& Muhlick
(DS, MICH).
9351
S of Weiser,
(MIN).
Beaverhead
(BH, IND, NY, POM, US).
s.n. (PH). Sanders
10, SE of Butte,
Richards
Smith
Thomas
14570
5 mi
NY, RM,
Cassia
14548: Munz
Shoshone
104 (UC). Hennepin
C. L. Hitchcock
Co., near Cascade,
IND, NY, POM, US).
Co.,
Itasca Park, Mayle
Co.,
Baker
NA,
(CAN, COLO,
Salmon
Co.,
(NA).-MINNESOTA:
16 mi N of Grand Rapids,
Flathead
of Three
Phillipsburg
7057
MT, NY).-MONTANA:
Cascade
Reservoir,
(ID). Washington
Bright
cult. from Munz
DS, NY, RM, RSA, WS, WTU).
Perdue
river, Baker
20363
Lehmi
2
s.n.*
S of Chatcolet,
MO,
cult. from Munz
NY, RM, US, WS, WTU).
Co., 20 mi E of Townsend,
1mi W
Gallatin
Co.,
29 & 104
(MIN, MO,
RM, UC, WS, WTU).
Butte,
(POM),
2011
from Alpena,
Itasca Co.,
and
18512
(F,MO).
14548
(COLO, DAO,
(MIN). Clearwater
s.n. (MO).
1558
Munz
below Weston
Co.,
along
6 mi SW of Preston,
Co.,
2.5 mi
Co.,
1142 (BRY, NY, UNCC).
IND, NY, POM),
Leiberg
of Heyburn,
Co., Cougar Mtn, Aller
MICHIGAN: Alpena
(BH, DS,
Co.,
& Greene
7 mi SW of Swan Valley,
Co.,
S of Elk River
stream, Wag
Summit
(COLO).
(BH, GH, MIN,
of Elk City, C. L. Hitchcock
(DS, F, GH, MIN, MO,
21715
& Muhlick
2 mi
(ID). Franklin
20 mi W
Oneida
Co.,
14578
Co., Caeur d'Alene,
2 mi NE
Co.,
Gulch, Macbride
Munz
Idaho Co.,
POM, NY).
along Lost River, Atwood
253
Jenson
(ID). Bonneville
small
Ada Co., 7 mi N of Boise, Munz
Benewah
1265
&
1mi E of
and Indepen
Aspen
1877, McCosh
Pass,
(POM, WS).
& Macbride
16026
(ID). Clearwater
3 mi N of St. Anthony,
(BH, WTU,
glehaus
Butte Co.,
(ID, NY).
2424
Nelson
Co., Priest Lake, Baker
Bonner
Christ
Co., Ketchum,
14411
(BH, IND, US).-IDAHO:
Davis
(BH, IND,
Stephens
Park Co.,
between
Ramaley
(MO). Teller Co., Ute
13026
Munz
Sand Dunes,
Co., Cuchara
La Plata Co.,
13027
Munz
Ear Pass along
of Rabbit
NY, RM).
Jones 36034
(RSA).
s.n. (COLO).
Campground,
of summit
Great
426*
Co., Evergreen,
Blaine
(OKLA).
UC, US, WTU).
W
Creek, Hoch
along Tenmile
& Clark
Pitkin Co., Weller
1.5 mi W
San Luis Valley,
Co.,
1980, Fenton
&
City, Norby
9 mi E of Stonewall,
Co.,
2
near Platte
Co.,
Huerfano
555
Co., Livermore,
(US). Las Animas
SE of Mack,
(MO). Routt Co.,
(NY, POM, UC). Adams
Mahler
Co.,
Co., 0.5 mi S of Platteville,
(NY). Weld
14552
Mesa
(MO). Saguache
of Frisco
(RSA, US).
& Norby
Co.,
Raven
(COLO, RM). Hins
3574
Norby
s.n.*
Chaffee
of Rollinsville,
650-62
(BH, IND, NY, POM).
26545
Pass, Raven
ner 4043*
1898, Crandall
9 mi W
& Robbins
1917,
Munz
of Salida,
s.n. (CU, DS, MIN,
& Spongberg
(BH, IND, NY, POM, US). Larimer
13012
Munz
Ft. Collins,
POM, US);
(UC, WS).
near Brighton,
Denver
2 mi N of Leadville,
Barrell
Creeks,
40, Sanders
8074
5 mi N of Palmer
& Clements
Ramaley
on Hwy
2 mi NW
(COLO).
Co.,
Co., East Portal,
Springs,
Oak Creek
(IND, US).
River
15218
Ramaley
ARIZONA:
Co.,
Co., Red Bluff,
Co.,
13033
Munz
(US). Douglas
s.n. (CU). Gilpin
Lake Co.,
Adams
(CAN);
et al. 2801
Senn
& Schneider
Tehama
9018
U.S.A.
Coconino
of Ashfork
Grant
near Arkansas
1901, Clements
Canyon,
Park, Hot Sulphur
Grand Co., Middle
S of jct. of Fish & Bebolla
dale Co.,
S of Longmont,
s.n.*
Smith
1888,
5 mi W
Co.,
Indian
(CAN, NY);
(DAO).
(MNA).
(UC).-COLORADO:
near Fort Garland,
Creek
263
239
(CAS, NY, RSA).
IND, NY, POM, US);
(RM); Denver,
Co., Keating,
Valley,
(BH,
Co., Ute
1622
(ARIZ, UBC).
dence
13020
of Salida, Munz
de Vries
S of Alturas,
28197
Howell
72378
Jaw Creek, Macoun
1 mi N of St. Louis,
(DAO);
Yavapai
Co., Pit Valley,
Co., Carrville,
(BH, IND, NY, POM, US).
S1191
UNM).
& Herriot
(CAN); Moose
Creek, Marshall
(ARIZ, ASU,
Modoc
4953
Fort Qu'Appelle,
Diamond
Indian Valley,
Sierra Nevada,
River,
NY,
5100
Macoun
Cherryfield,
Sallans
(DAO);
Little
along
& Sanders
Co.,
13025
s.n.
Shevkenek
(MO).-CALIFORNIA:
Plumas
mi NW
(CAN, NY);
s.n. (TRT); Canora,
Sallans
Outram,
& West 3886
72379
s.n. (CAN); Crane Lake, Macoun
1911, Malte
VOLUME 50
of Va
Co., Lame
10 mi E of divide
Co., W
Fork Camp,
1997
Stillwater
Elliot
River,
Yellowstone
POM).
McGregor
& Bare
533
4262
1919, Tidestrom
10693
(MO). Catron
Mogollon
along Willow
sula of Lake
Co., Devil's
Steele
(KANU).
Sauvie
Island, Trainer
63-19
River, Munz
14470
1mi NE of Central
Jackson Co.,
Klamath
Creek
3 mi down Willamette
20 mi
S of Nyssa,
Co., Umatilla,
Peck 4381
Canyon,
Umatilla
(DS, IND, MICH,
(KANU).
Custer
City, Moore
715
Iron Creek
Lake,
1905, White
2 mi E & 6 mi
(MIN). Harding
13732
UTAH: Box
Elder Co., Bear River Migratory
3 mi W
of Logan,
S of Asotin,
Baker
US, WS).
15216
14527: Munz
from Munz
(ID). Benton
14707
Island Co., Seattle,
14468
bottom, Munz
14770
US), Munz
(BH,
168
of Yakima
on Naches
Township,
0.25 mi W
Co., Drummon
Polk Co.,
5.7 mi
Goshen
103
Co.,
Falls,
Co.,
Bartlett
Crook
Nelson
3.5 mi W
(RM). Park Co., Yellowstone
RM).
Sheridan
MO,
Co., Businga
near Ranchester,
ND-G,
Ranch,
Hoch
427*
63 on country
Co., Radisson,
Canyon,
(RM). Hot
Porter
Natl.
NY, POM, RM, US).
Story, Uttal
5173
(MO). Sweetwater
397
1948, Scheffer
Co., Bingen,
river
IND, NY, POM,
Okanogan
Co., Nancy
Horner
1896,
Creek,
(GH,
R181B
40 (UWSP).
Porter
& Porter
Co.,
TEX, WTU).
RSA).
8381
Hot
Carbon
Co.,
Platte Co., Whalen
(RM). Natrona
Canyon,
Co.,
Natl. Wildlife
Nelson
Refuge,
UC).
Freylag
Co., Garden
Nelson
along U.S.
Al
Slater,
(DS, RM, RSA,
Springs,
247
Big Horn Co.,
20 mi N of Shoshoni,
10535
Sublette
Langlade
Kelsey
3 (WIS).-WYOMING:
(DS, MICH,
Park, Mammoth
Co., Seedskadee
Stevens
rd N, Allen
McFerson
Springs
(TENN, VPI).
cult.
(BH, NY, UC,
0.5 mi E of Camp Kawaga,
PH, RM, RSA,
1298
Co., 4 mi
(BH, IND, NY, RSA, US).-WISCON
5 mi N of Sundance,
2350
1976, Greenwood
Pierce Co., Tacoma,
Co., Waitsburg,
14511
8161
Co., Silver Creek,
(BH,
14709
(BH, WS, WTU).
S of Minocqua,
& Grayson
Co.,
179
of Granite
Klickitat
of
(MO).
& Goodrich
(BH, IND, NY, POM, US).
Walla Walla
of Hwy
Sawyer
Co.,
(CU, DS, GH, MIN,
7706
River, Munz
1mi
(POM).
Asotin
& Leiberg
14468: Munz
15250
(DAO, DS, GH, MO, MT,
2.8 mi N of Basin,
6029
Dayton,
4329
1-1.5 mi SW of Torrington,
Jozurk
(UC, NY, WS).
s.n. (MIN).
(COLO, NY, RM, US).
(RM). Laramie
Creek
region, Porter
S of Bray Bull,
1744
(WIS). Oneida
1892, Burglehaus
bany Co., Pole Mtn
Goodding
194
14403
(BH, IND, NY, POM),
s.n. (NA, SMU, WS).
Suksdorf
& Weldert
Jack Lake, Kelsey
(WIS).
Co., Prindle,
2 mi W
(WS). Grant Co., Dry Falls Coulee,
Munz
Stone
Co.,
et al. 16088
Canyon,
Sandberg
859
51675
Stephens
Over
(UT). Summit
14527
(NY).
(OSC).-SOUTH
Neese
(MIN, NY, POM, RM, US, WS).
15 mi NW
SIN: Bayfield
Spring,
cult. from Munz
Falls, Boner
Co.,
2259
1941, Eyerdam
495
of Neola,
Munz
s.n.
& Grayson
Grant Co., Big
of hdqtrs., Holmgren
of Kennewick,
& Chapman
Island,
Bartlett
(BH, IND, POM).-WASHINGTON:
Co., near Egbert
City, Elmer
Skamania
US). Yakima
Co.,
6 mi W
Co.,
Co., Sucker
1894, Lloyd
Lake,
Co., Whiterocks
(BH, IND).
Co., near Lake Ewana,
(RM). Lawrence
4695
River Co.,
15246
S of Hecla,
Stone
7.5 mi NW
Uintah
15013
IND, NY, POM, US, WTU),
Okanogan
3 mi W
Co.,
Salt Lake City, Arnow
IND, NY, POM, US);
s.n. (CU, WS, WTU).
Co., Big
(BRY, NY).
(US). Douglas
Harbor
(BH, DS,
Co., near Lake Chelan,
2 mi N of Kettle
12548
along RR, Beatie
Ferry Co.,
s.n. (WS).
& Welsh
Roberts
1684
cabin, Head
(ID). Malheur
(KANU).
267
Visher
Bird Refuge,
6 mi
35052
MIN,
36556
Stephens
Peck 4382
River,
Co.,
& Brooks
(MO). Duchesne
3 mi SW of Heber, Munz
Co.,
of Deschutes
Slim Buttes,
(KANU).
10 mi E of downtown
SE edge of Park City, Welsh
s.n. (BRY). Wasatch
1803
Shultz
Salt Lake Co.,
(BRY, MO).
Stephens
Co., Shady Valley,
Cache
Co.,
at mouth
s.n. (MO). Brown
S of Custer,
& Brooks
Stephens
3360
Co., S of Imbler, 3.4 mi NE of Alice,
River
Uttal
Co., Penin
s.n. (OSC). Hood
near Jefferson,
Co.,
(MO).
s.n. (OSC). Colum
14470: Munz
Baker
4212*
s.n. (CAN, DAO,
(BH, IND, NY, POM, US). Klamath
of Coburg,
Mtn,
further lo
(NY). Benson
1952, Mosier
1932, Tucker
(OSC). Marion
(OSC). Union
Co., Brookings,
Co.,
22308
Co., Columbia
RSA). Wasco
DAKOTA: Brookings
Peck
inWhite
Standley
near Smidts'
Co.,
cult. from Munz
Lane Co., W
Lake
Mtns,
3 mi E of Questa,
1935, Stevens
Baker
Co., Prineville,
14411
(DS, OSC, US).
280
(MO);
Mogollon
Co., 0.5 mi S of Ft. Ransom,
from Corvallis,
(BH, IND, NY, POM),
Point, Munz
2145
Falls, Lawrence
River
(OSC). Crook
near Cowles,
Fargo,
s.n. (OKL).-OREGON:
1891, Wright
Co.,
S of Hood
bia Co.,
18 mi
Benton
Co., Bonita
Lake, Mabbott
(MIN). Ransom
Park, Ellis
4824*
Fletcher
(F). Rio Arriba Co., without
Creek,
Cass Co.,
US).
2635
Co., Wadsworth,
east side, Balsam
(MO). Taos Co., Red River,
Eckelson
Sanborn,
Lake, Bergman
Co., Hope,
(GH, OSC, WS).
Co.,
s.n. (F, MIN,
1909, Lunell
Ibsen,
Ramsey
7537*
Worthington
(MO). Elko Co., Elko,
(MO, UNM);
Lincoln
18573
Co., Winsor
(US). San Miguel
DAKOTA: Barnes
7623*.
Area, Worthington
17016
13 mi N of Hay
(US). Wadsworth
R. 5420*
(CAS,
of Johnstown,
& Brooks
Co.,
Indian Creeks,
and
Fletcher,
de las Casas, Arsene
Co., Canyon
11145
Canyon,
627
Sandia Mtns,
Co.,
of Gilita
Stephens
Tiehn 3610
Ranch,
A. S. Hitchcock
2419
3 mi W
Co.,
Sheridan
(KANU).
Co., ditch near Mori
confluence
Mtns,
C. L. Hitchcock
Brown
of Chadron,
20033
MEXICO: Bernalillo
side of Sandia Mtns,
(VPI).-NORTH
WIS).
Churchill
Co., Battle Mtn,
at Ben Lilly Recreation
& Bollman
Co., W
15 mi W
McGregor
at jct. of Whitetail
Creek
(UNM). Mora
cality, Standley
9930
S &
Co., Mogollon
Creek
1530
Sandoval
13 mi
Co.,
(GH, US).-NEw
Mineral
Mtns,
Hutchins
Dawes
Lander
(DS, NESH).
of Harlowton,
(RM).-NEBRASKA:
S of McCook,
1 mi
10 mi W
Co.,
5155
(KANU).
Co.,
Wheatland
Bartholomew
186 (RM).-NEVADA:
Springs, Nixon
Kennedy
137*
70 (POM, WIS).
Co., Billings,
Red Willow
(KANU).
Flett
175
OENOTHERA
& Nelson
519
(MIN,
14, just SW of
Welsh & Welsh
176
SYSTEMATICBOTANYMONOGRAPHS
(MO). Teton Co., Grand
19114
Stockade
Co.,
6. Oenothera
Teton Natl.
Nelson
Beaver,
9483
965
Park, Williams
(GH, MIN,
(CAS, GH,
VOLUME 50
NY, RM, UC). Weston
IND, MO,
RM).
stucchii.
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. Italy. REGION LIGURIA. Prov. Genova:
Ind. Sem. Bot. Gard. Genova,
s.n., cult. DUSS-86-2312
1983, Soldano
bano,
1991, cult. DUSS-92-2005
and 111), cult. DUSS-88-2021.
BOUCHES-DU RHONE: Aries,
1978, Geerinck
Novara:
Isernia: Sessano
Ponte Ticino,
Galliate,
del Molise,
1982, Tammaro
(MO). Prov. Vercelli:
3624
Soldano
GION TUSCANY. Prov. Lucca: Viareggio,
Al
(MO) (012
ADDITIONAL SPECIMENS EXAMINED. France.
Italy. REGION MOLISE. Prov.
Porto di Pre,
PIEDMONT. Prov. Vercelli:
(MO) (014).-REGION
1913, Sevelli
Cerrione,
4178
Soldano
Elvo,
s.n. (FI). Prov. Pisa: mouth
1794
(BR).
PIEDMONT. Prov.
s.n. (FI).-REGION
(TO).-RE
of river Serchio,
1971, Seipka
s.n. (KTU).
7. Oenothera
grandiflora.
SPECIMENS EXAMINED FROM CULTIVATED PLANTS (diakinesis
by E. Schumacher
made
et al., 1992; Schumacher
1986; Schumacher
1983, Steiner
84-342
s.n., cult. DUSS-84-337
(MO); Bay Minette
1983, Steiner
Bolinger,
Conecuh
berry
(2 04
and 3II), DUSS-84-317
(MO)
DUSS-84-325
and 5II), DUSS-84-326
and 3II) (MO). Sumter Co., Bellamy,
(04
and 5II; 06
bee,
1983, Steiner
and 4II); York,
and 3II); Frankville,
Steiner
1983,
s.n., cult. DUSS-84-368
cult. DUSS-84-359
DUSS-84-331
(MO)
(MO)
cult. DUSS-81-024
(MO)
rd 87, Godfrey
Florida
(2 04
and 5II; 2 04
(7II; 04
(MO)
(both 04
strain
(MO)
and 5II).Washington
(04
Co., Big
and III), DUSS-84-373
(7II; 04
and 5II); county
and 5II); Sims Chapel,
Escambia
(MO),
and 5II; one
(MO)
rd 6, 1983
1983, Steiner
1983, Steiner
Co., Cantonment,
s.n.,
s.n., cult.
and
and 3II). Santa Rosa
(MO), DUSS-84-334
06, 04 and 211), DUSS-84-333
31;
Co., Avalon Beach Rd, vicinity of Milton, Godfrey 76808,
and 3II; 06,
cult. DUSS-80-064
76815,
(MO). Escam
s.n., cult. DUSS-76-0110
(7II; 04
(MO)
(both 04
(MO)
and 311).-FLORIDA:
(7II), DUSS-84-332
(MO)
(MO)
(7II), DUSS-84-374
(all 7II),
(MO), DUSS-84-324
1974, Jones & Arrington
s.n., cult. DUSS-84-370
(MO)
and 5II), DUSS-84-313
(MO), DUSS-84-314
(MO), DUSS-84-328
(MO), DUSS-84-369
Co.,
and 5II)
(MO) (04
(MO), DUSS-84-319
(MO) (7II; 04
(MO) (7II), DUSS-84-323
cult. DUSS-76-0105
(MO)
Steiner
(7II; 2 04
and 5II; 2 04
(both 04
(MO)
Jones 15345,
s.n., cult. DUSS-84-372
and 5II). Choctaw
(04
(0 10 and 211; 0 12 and 1i), 1983,
(MO) (010 and 21I); Castle
(MO)
and 4II), DUSS-84-312
(all 04
(MO)
(326) also 2 04
(2 04
(all 06
(A),
and 5II)' DUSS
s.n., cult. DUSS-84-309
and 5II), DUSS-84-316
s.n., cult. DUSS-84-330
1983, Steiner
Co., Bay Minette
(MO) (all 04
(MO)
strains were
1987; Steiner & Stubbe,
(MO), DUSS-84-365
(MO), DUSS-84-318
(MO) (all 04
(MO)
cult. DUSS-84-347
s.n., cult. DUSS-84-304
s.n., cult. DUSS-84-315
(MO), DUSS-84-320
bia Co., Flomaton,
s.n.,
1983, Steiner
(MO) (0)14),
DUSS-84-311
(MO), DUSS-84-344
(MO), DUSS-84-364
(A), 1983, Steiner
Steiner
1983,
Steiner
1983,
in Schumacher,
ALABAMA: Baldwin
1993). U.S.A.
(MO), DUSS-84-339
(B),
s.n., cult. DUSS-84-307
(B),
& Steiner,
of the E. Steiner
examinations
and some are published
s.n., cult. DUSS-84-361
Co., Castleberry
Steiner
von Berg,
and G. Linne
and 211); 0.5 mi E of jct. US Hwy
04
(7II; 04
(MO)
and 5II)' DUSS-80-210
90 with
(MO), DUSS-80-211
(MO),DUSS-80-212 (MO),DUSS-80-213 (MO),DUSS-80-215 (MO).-MISSISSIPPI:
County andcollectorun
cult. DUSS-76-0105
known,
1979, Kral
(MO)
s.n., cult. DUSS-81-021
(04
and 511).-TENNESSEE:
(MO)
(7II), DUSS-87-ST279.
Marion
Co., Monteagle,
along
24,
Interstate
CULTIVATED STRAINS WITH DIAKINESIS CONFIGURATION EXAMINED BUT WITHOUT VOUCHER (all exami
by E. Schumacher
nations made
Steiner
1983,
s.n.,
DUSS-84-335
(7II; 04
DUSS-84-346,
DUSS-84-349
363
and G. Linne
cult. DUSS-84-340
(both 7II). Conecuh
and 5II); Bay Minette
(all 04
von Berg).
Co., Castlebeffy
Co.,
(A), 1983, Steiner
and 111),DUSS-84-351
(012
and 3II)' DUSS-84-354
(08
496
(CU, F, GH, MIN,
(US). Mobile
Tunnel
Clark
Springs,
17280
MO,
Co., Mobile,
Harper
(NCU).
2999
NY
of Alabama
Gainesville,
Arnold
(7II), DUSS-84-345,
s.n., cult. DUSS-84-360,
DUSS-84
s.n., cult. DUSS-84-301,
(7II)' DUSS-84-353
(04
ALABAMA: Baldwin
5 sheets, PENN,
Bartlett
& Grayson
(BH, MO,
Sumter Co.,
University
(A),
and 5II)'
DUSS-84-302
DUSS-84-329
(both 010
(both 7II), DUSS-84
Co., Chastang,
1983, Steiner
and 5II), DUSS-84-356
(04
s.n., cult.
and 5II; 08
and 3II)
REPRESENTATIVE SPECIMENS. U.S.A.
8001
(all 04
s.n., cult. DUSS-84-348
1983, Steiner
and 211). Escambia Co., Flomaton,
1983, Steiner s.n., cult. DUSS-84-321,
322 (04 and III), DUSS-84-327
(7II; 04 and SII; 2 04 and 3II). Mobile
DUSS-84-350
Co., Bay Minette
DUSS-84-342
DUSS-84-341,
(B), 1983, Steiner
and SII). Choctaw
ALABAMA: Baldwin
U.S.A.
(7II), DUSS-84-336,
NY).
171 (FLAS). Escambia
3976
3214
Pickens
1.7 mi S of York,
campus, Harper
TAES,
Co.,
(MICH),
Co.,
Jones
3214a
8.2 mi
15345
(FLAS, GEO,
Co., Pensacola,
12 mi N of Tensaw,
TEX, US, WIS).
(FLAS,
Dixie
Co.,
Tracy
Howell
ca. 1.5 mi S of
bank of Tombigbee
SMU, TENN, WVA).
NCU,
Landing,
Co., Castleberry,
(MICH). Monroe
S of Aliceville,
GH, MO,
Brinker
Conecuh
US).-FLORIDA:
River,
Tuscaloosa
Co.,
Alachua
Co.,
411 (MO). Franklin Co., Apalachicola,
God
1997
OENOTHERA
frey 65863
58330
(DS, FSU).
Lake Co.,
Leesburg,
(FSU). Polk Co., Bartow, McFarlin
Rosa
vic. of Milton,
Co.,
PH).-MIssIssIPPI:
rest General
Clay Co., Kilgore
Hospital,
Hattiesburg,
Jones
14583
POM, US).
Tishomingo
Co., near Holcut,
ham 20056
(CU).-NORTH
Leonard
Moore
et al. 2535
3 mi N of Southern
Co.,
& Randolph
1005
(PENN).-SOUTH
burg Co.,
2688
Bell
SSE of Monteagle,
Franklin
Kral
VIRGINIA: Monongalia
Co., without
(UNCC).
47557
(MO, US).-VERMONT:
3339
Sheldon
30162
Windsor
(UNCC).
Spartan
27488
Radford
s.n. (MO). Marion
Dale
Pin
11559
Glowenke
of Shiloh,
Co., Chester,
Randolph
Swain Co.,
(UNCC).
4 mi ENE
1898, Eggert
17,
SMU, UC, UNCC, WTU).
& Sake 62121a
locality, Rogers
Mtn,
(TENN).
Co., 2 mi S of Wilmington,
Jordon Swamp
Co.,
42495
NY,
Burn
Co., Apiary,
Co., E of Mt. Mocono,
Monroe
(NE
(CU, F, GH, MO,
19, Rogers
RSA,
& Haesloop
(NY,
64 ca. 1mi E of jct. Hwy
along Hwy
New Hanover
further
Sumter
Co., Cumberlin
Co., Morgantown,
Co.,
S of Ingold, Ahles
(TENN).-PENNSYLVANIA:
8325
on Hwy
NY, OKL, OKLA,
593 (UNCC).
3.7 mi
Co.,
MIN,
1285
YORK: Tompkins
(TENN).-NEW
(TENN). Martin
2736
Pines, Carter
(COLO, UNCC).-TENNESSEE:
mi
50315
Santa
s.n.
18 mi E of Crawford
Co.,
Pollard
College,
Short
Forrest Co., E of For
(TENN).
Lowndes
SMU).
Godfrey
s.n. (FLAS).
Co., Lexington,
45602
Co., S of Marble Meadow
FSU, GH, MICH,
CAROLINA: Oconee
Spartanburg,
(FSU, NCSC,
Co., Agriculture
Coleman
Wilson
(CU). Sampson
Smith & Jennison
Oak Gap,
14176
CAROLINA: Cherokee
(COLO, FLAS,
2 mi E of Tallahassee,
1940, Laessle
Fayette
Sec. 8), Rogers
(T15S, R3E,
(MO). Oktibbeha
Co., Hall Place, Highlands,
Macon
(FSU, MO).-KENTUCKY:
Hills
1/4 Sec. 33), McDaniel
Leon Co.,
(FLAS).
(MICH). Putnam Co., Welaka,
6119
76808
Godfrey
188
Baltzell
177
Co.,
822
1-24 E,
1
(DAO).-WEST
(WVA).
SPECIMENS CULTIVATED IN BOTANICAL GARDEN. Germany.
Erlangen,
s.n. (M); Frankfurt,
1798, Schreber
1823 (FR).
8. Oenothera
nutans.
SPECIMENS EXAMINED FROM CULTIVATED PLANTS (diakinesis
and most
Wasmund
look, Stubbe
(MO)
571
DUSS-81-578
(MO)
(014),
1978, Pitillo
0579
(MO)
3928,
cult. DUSS-79-0571
Boone
woods,
ca. 1200 m,
hagone
Overlook,
DUSS-81-580
along
1978, Rossbach
forest
Harman,
mi W
Sevier
service
rel Fork Bridge
E of Elkins,
of Whitmer
(II), along Hwy
Klingman's
Natl.
Parkway,
(I), ca. 1340 m,
13768,
Dome
(012
(014). Wilkes
Cleland
Morton
9024
(CAN);
ABAMA: Baldwin
Creek,
& Rossbach
Randolph
Stubbe
44,
1978, Bell
s.n.,
(MO) (014);
Boone
1I,);
Co., Doughton
Lick Gap,
(IV), Campus
Park near Ma
Stubbe
37, cult.
200, cult. DUSS-78-0138
(MO)
441,
ca. 1500 m, Solomon
1215,
1216,
Co., Dolly
(MO)
Sods
(014);
Co., Wymer,
1978, Stubbe
7142
(CAN);
cult. DUSS-79-0565
(MO)
(014);
ONTARIO: Jct. of Moose,
1927, Ricker
Loading,
Bartlett
(MICH). Calhoun
along
3 mi
3 mi NW
(014);
rd 33, Mid
& Missinaibi
Lake, Harton
1934, Wood
Co.,
VOUCHER. U.S.A.
(014).
Mattagomi
11, Postagoni
s.n. (TRT); Strathroy,
2750
(014);
rd 33, Lau
(MO) (014);
(MO)
area,
(I) and
(MO) (014).
s.n., cult. DUSS-84-474
1.5 mi E of Hwy
along
s.n., cult. DUSS-79-0558
s.n., cult. DUSS-79-0556
Elkins
(MO)
(MO) (014);
cult. DUSS-79-0564
recreation
between
s.n., cult. DUSS-79-0557
cult. DUSS-79-0561
& Rossbach
ca. 1060 m, Evans
29, ca. 1000 m, Evans
(014).
& Rossbach
1214,
ca. 930 m, 1978, Glencoe
Port Credit,
Co., Dixie
(MO)
ca. 1650 m,
Gap,
ca. 830 m,
and
Co., Deer
cult. DUSS-79-0562
1978, Glencoe
REPRESENTATIVE SPECIMENS. Canada.
& Porsild
cult. DUSS
Rd,
and Newfound
CULTIVATED STRAIN WITH DIAKINESIS CONFIGURATION EXAMINED BUT WITHOUT
Island, Baldwin
S Inckasegee,
cult. DUSS-79-0576
Hospital,
Park, Park jct. with US
(MO)
2118,
33, ca. 660 m, Evans
1978, Glencoe
WEST VIRGINIA: Randolph
ca. 1650 m,
107,
s.n., cult. DUSS-79-0584
Yancey
Porter Co., Coudersport,
Smoky Mtn
Ridge
(II), Hardin
Dome
(MO)
(MO)
(014).
38, 39, cult. DUSS-81
along NC
13765,
ca. 1000 m,
Parkway,
Stubbe
Rd, E of Highlands
County
s.n., cult. DUSS-79-0585
(MO)
32, 33, cult. DUSS-81-577
441 along Klingman's
(I), opposite
s.n., cult. DUSS-79-0553
(I), along Gandee
atWymer,
Cove
of Highlands
(II), ca. 1100 m, 1978, Morron
rd 70, ca. 1330 m, Evans
(II), along US
E of Whitmer
along Blue
Irassy Branch
by 0.
made
Co., Bear Den Over
(MO) (0)12 and 1II).-VIRGINIA: Carrol Co., Pipers Gap, 1978, Bell s.n., cult. DUSS
VIRGINIA: Pendleton Co., Seneca Creek, between falls of Seneca and White Run,
rd 33 at top of Rich Mtn,
along
of Elkins
dle Mtn,
Co., Great
Rd Gap
Co.,
of US
s.n., cult. DUSS-79-0586
primarily
Ridge
Valley,
ca. 1000 m, Hardin
between
cult. DUSS-81-575
Stubbe
on Blue
Co., Maggies
rd 64 W
Park, 6 mi W
(014);
Mtn,
Pisgah
Co., Horse
along
rd 1603,
(MO) (G)14).-PENNSYLVANIA:
(MO) (0 14).-WEST
ca. 760 m,
Boone
1978, Morron
28a,
Jackson
Co., Boone
1978, Morron
cult. DUSS-79-0597
3926,
along
(MO)
(014);
421,
Stubbe
(014).-TENNESSEE:
79-0552
Cove
(014). Watauga
(MO)
(III), along US
(014).
Smoky Mtn Natl.
(MO)
S of Wagon
(014);
(MO). Macon
cult. DUSS-79-0591
cult. DUSS-81-584
Co., Mt.
Haywood
(MO) (014);
SE of Horse
(014);
(014).
were
examinations
NORTH CAROLINA: Avery
and 111);Grandfather
(012
(MO)
cult. DUSS-79-0575
Swain Co., Great
Solomon
(MO)
s.n., cult. DUSS-79-0592
13764,
(014).
(MO)
DUSS-81-581
1990). U.S.A.
Buncombe
(014).
cult. DUSS-79-0580
13769,
Hardin
(MO)
(MO)
cult. DUSS-79-581
13770,
Hardin
inWasmund,
34, cult. DUSS-81-579
(014),
Hardin
are published
7864
Rivers,
Portage
(DAO); Wingham,
s.n. (DAO).-U.S.A.
6 mi N of Jacksonville,
Clark
AL
7026
1
SYSTEMATICBOTANYMONOGRAPHS
178
(UNCC).
Conecuh
on Hwy
Auburn,
1896, Baker
Tuscaloosa
69, Whetstone
Lake, Apple
Munz
from Tallahassee:
13357
Sec. 20),
Alachua
4.5 mi SE of Adairsville,
Greear
8 mi S of Fort Gaines,
Co.,
Co., Croy Creek
30223
berland
River
Lincoln
Valley,
Neck,
Fernald
gomery Co., Chevy
3659
shire Co., Amherst,
191 1,Hasselbring
1935, Long
Dutchess
Co., Brady's
2172
cult.
NY, POM),
1900, Morss
(BH, MICH,
Pawling,
Co., Holden,
Palmer
Station, Randolph
& Wiegand
13400: Munz
10472
5780
(LAM). Erie Co., Buffalo,
Baldwin
10475
14209
(CU). Tompkins
(BH,
Hamp
HAMPSHIRE:
1mi E of Cherryville,
13389
1131
Sawada
Co., 5 mi W
(BH, IND, POM).
(KYU). Greene
Lake, Gershoy
10468
1mi E of Etna, Munz
Co.,
of Patux
Genesee
(CU). Broome
S of Cayuta
Co.,
mouth
s.n. (NEBC).
s.n. (POM).-MISSOURI:
Co.,
Falls, Munz
IND, POM). Washington
(BH,
(BH). Mont
(MIN, NY).-NEw
1mi W
of Cook's
3161
(OKL).--MICHIGAN:
Co.,
s.n.
Pistaquis
13471
Munz
Barlett
1890, Baker
3452
Knowlton
1941,
Co., Millstone,
236
Bar
Cum
Co., Foxcraft,
Baltimore,
JERSEY:Hunterdon
(NEBC).-NEW
Clay
Deam
Co., Black Mtn,
1913, Forbes
Piper
College,
Co., Alfred
Pistaquis
St. Marys
Co.,
(CU). Early
(KANU).-MAINE:
de Grace,
Co., Backland,
1891, Vail s.n. (NY). Schuyler
Eames
from Munz
UC, US).
Bartow
1073
(F).-INDIANA:
Chesterville,
Co.,
Co., Havre
Ingham Co., Agrielo
YORK: Allegany
Swamp,
Co.,
s.n. (NEBC).
Jasper Co., Carthage,
12857
Pease
Harlan
66-1013
Baltimore
Franklin
(IND, POM). Delaware
of Cayuga,
Henderson
Franklin
(PENN).
20 (TIS, R4W,
Co., Hwy
Smith 2411
267
cult. from
Leon Co.,
ca. 1/4 mi SW of Vallonia,
(IND).-KENTUCKY:
(WVA). Harford
2806
(MO).
East Rill Valley,
Co., W
Seneca
9897
s.n. (UC). Worchester
near Riverton,
13390
Co., Tannersville,
44
Steele
s.n. (MICH).
Bush
s.n. (PH).-NEw
Munz
s.n.
(CU).-MASSACHUSETTS:
Jackson Co., Courtney,
of Deposit,
Deam
Holmer
Pleasant,
& Maguire
Miller
Jackson Co.,
(IND).
s.n. (GH, NEBC).-MARYLAND:
1909, Brooks
Jefferson
13854
Bayville,
Chase Lake, Bartlett
ent River, Bartlett
Coos Co.,
Harbour,
Liberty
(UNCC).
vic. of Waterbury,
Co.,
s.n. (MICH).-GEORGIA:
1900, Pieters
5 mi NE of Richmond,
Co.,
Haven
Jannin Co., Blue Ridge Mtns,
Deam
of Hillsdale,
(IND, NY).
22111
s.n. (MICH, UNCC).
(FLAS, GA, MO, WIS).
Clarke Co., Winterville,
(MICH).
1967, Moldenke
Co., Boothbay
1897,
14580
(UNCC).
6117
Co., N of Oakland,
IND). Garrett
Co., Flint,
Thorne
182 (CU). Madison
Prouts
Co.,
(NEBC).
63261
Co., NW
(NY). Vermillion
(IND), Munz
Co., Clarcona,
1mi E of Harmony,
ca.
bour & Barbour
14239
Munz
s.n. (FLAS). Orange
1967, Smith
2209
of
(MO). Lee Co.,
163, Raven
Co., Mount
Castle
NE
Church,
36857
Jones
1960,
s.n. (US). New
DArcy
Kral
on Hwy
River
New
(NY).-DELAWARE:
Co., Gainesville,
SE of Concord
Co., Geneva,
Co., Camden,
1902, Dowell
Preston,
107
1.5 mi
Co.,
Geneva
(UNCC).
0.6 mi N of Dog
Co.,
near New
Co.,
(CU). Cullman
5238
Iltis s.n. (WIS). Wilcox
1943,
Hill Rd, Lucian
(CU, MIN).-FLORIDA:
3650
& Wagner
s.n. (POM). Mobile
Co., Tuscaloosa,
CONNECTICUT: Litchfield
Bantum
Bartlett
Co., Castleberry,
Cullman
VOLUME 50
(CU).
13400
(DS,
N of Hudson
Co., Vaughns,
Falls, 1903, Burnham s.n. (CU).-NORTHCAROLINA:
Ashe Co., near Jefferson,Correll 4029 (DUKE).Avery
6 mi NE of Linville,
Co.,
14618
(POM), Munz
115, Daggy
(UNCC).
4177
4176,
(UNCC).
13520
(IND, POM). Mitchell
(DAO).
Blowing
Munz
Rock,
Munz
Cuyahoga
Rowan
14614
Co.,
Mtn,
s.n. (NY). Swain
Co.,
13512: Munz
13516: Munz
14128
Co., Unity
14587
3 mi
14270
(BH,
SW of
(BH, DS,
Yancey
Co.,
(POM), Munz
township
1895, Stoir s.n. (MO). Ottawa
650
& Leisner
Ahles
1958, Burk s.n. (UNCC).
Columbiana
Pittillo
(POM), Munz
s.n. (DUKE). Watauga
cult. from Munz
73 &
16123
3 mi W of Old Fort, Munz
1894, Small
14259
Rds.
Gap, Radford
Co., Bald-top
13523: Munz
cult. from Munz
(POM).-OHIO:
between
Co., N of Silverdale,
1mi ESE of Dudley,
Co., Cleveland,
(IND, POM), Munz
River
of Black
Co., Dunnis Mtn,
1963, Mowbray
(BY, IND, NY, POM, US),
(UNCC).
ISC). Henderson
cult. from Munz
Co.,
64, W
(NCSC). McDowell
(BH, IND, NY, POM),
(POM). Wayne
(BH, IND, NY, POM, WTU),
7842
13512
area near Hwy
121 (US). Onslow
14277
Co., Rocky
13764
(DUKE).
Gorge,
13514: Munz
Cabarrus
(DUKE,
Cannon
3207
(BH, IND, POM),
14617
Creek
Hardin
Co., Bearwallow
13516
Munz
Cooperrider
Museum,
Co., Roan Mtn,
Blue Ridge, Munz
IND, NY, POM), Munz
cult. from Munz
1022
Sterling, Murley
13523
Transylvania
Mt. Mitchell,
Co., Buck
Polk Co., Tryon, Correll
Co., Cherokee,
POM).
Clay
Co., E of Highlands
32593
NY,
(NY, POM),
(BH, IND, NY, POM, US, WTU).
Co., Mt.
Haywood
(WVA). Macon
13514
Munz
14688
14253
(E 1/2 Sec. 35),
Co., Put-in-Bay,
Clarkson
2273 (WVA).Richland Co., Mansfield, 1897,Wilkinson s.n. (CU,DUKE).-PENNSYLVANIA:
Allegheny Co.,
Pittsburgh,
1950, Buker
DS,
IND, POM).
Chester
Bucks
Co., Berwyn,
ford Co., Meadville,
campus, Whitehead
Co.,
s.n. (KANU).
1mi E of Centerville,
ford Co.,
Co., Plumsteadville,
McKean
Co.,
Long 6763
Pue 464
(PAC). Fulton
Co., Slippery
3 mi S of Myrtle,
(PH). Northhampton
(PENN).
Sullivan
0.5 mi SE of Craigsville,
Moyer
Co.,
s.n. (US). Centre
1.3 mi W
39 (POM). Elk Co., Fairview,
1.25 mi SW of Petersburg,
(MIN). Lawrence
Co.,
s.n. (WVA). Bradford
10 (US). Clearfield
Steele
Curtis
46
Armstrong
1971, Duppstadt
Co.,
Westerfield
Rock,
0.5 mi SE of Fort Littleton,
6632
1946, Russel
1949, Wherry
Co., Bethlehem,
Co., High Knob
(PAC). Lancaster
s.n. (PENN).
1889, Caifrey
(PAC, PENN).
Ehrle
Wahl
208
Bed
13402
(BH,
(CU, GH, PAC).
& Wahl 2880
(PAC). Craw
(PAC). Erie Co., Edinboro
State College
Westerfield
Co., mouth
Lycoming
s.n. (PENN). Montgomery
near Eaglesmere,
Co., Waddle,
of Shawville,
Wahl 3347
Wahl 3973
10 mi S of Sayre, Munz
Co.,
12646
of Tucquan,
Co., Barbours,
Co.,
near Tacony
s.n. (PAC). Pike Co.,
1938, Westerfield
(PAC). Huntingdon
Heller
Wahl
Creek,
& Bach
13635
Ashbourne,
2.5 mi NE of Milford,
s.n. (PAC). Venango
548
(PAC).
De
Co., 2.5 mi
1997
NW
OENOTHERA
of Reno,
Baltzell
OLINA: Berkeley
Winnsboro,
9939
Bell
(PAC). Wayne
lace & Jennison
(UNCC).
1695
Oconee
MONT: Caledonia
(TENN).
458
Sts., Allard
1662
S of Petersburg,
Fernald
Co., White
Grayson
13481
(DS,
Stony Man Mtn,
near Luray,
2.6 mi S of Wabun,
13707
Steele
5655
Wood
VIRGINIA: Calhoun
s.n. (MIN). Greenbrier
13497
14693
IND, NY, POM, US), Munz
POM). Morgan
Girl Scout Camp,
Preston
1300
2 mi W
Co.,
1934, Bartholomew
(WVA). Webster
9. Oenothera
3 mi W
Co.,
& Steele
Co., without
along Hwy
near Castlegar
1, Wagner
cult. DUSS-82-0501
(0) 14); Slocan
(MO)
exact
82-0445
(MO); without
(MO).-QUEBEC:
Jones
Forsyth
Cienaga
Street, Dorr
Canyon,
look, Hardin
13767,
Stubbe
13494
(IND,
Co., Wheeling
1939, Brown
s.n. (MIN).
Clarkson
Creek
Picnic Grounds
(II), 1978, Morron
(MO)
(MO)
along US
441, Solomon
84-126
(MO), DUSS-84-124
Multnomah
(MO)
(014).
4537,
Co., Portland, Wagner
cult. DUSS-82-0483
1980, Stubbe
13766,
(MO)
(0)14).-TENNESSEE:
Hamilton
1706, DUSS-79-0595
(MO)
3939,
Co., Chattanooga,
Walden,
cult. DUSS-79-0549
(MO)
(014).
Jones
22858,
s.n., cult. DUSS-76-028
Co., Columbia,
(MO)
Park
Park, 5.8 mi NW
Natl.
(MO). Watauga
Point, Hoch
River,
Stubbe
cult. DUSS-82-0384
(MO).-SOUTH
1656,
1729,
Irassy
Falls Over
226a with Blue Ridge
jct. Hwy
Smoky Mtn
Sandy
(014);
4535,
Van Horn
(014).
of
Co., Boone
Co., parking place "Devils Garden Overlook,"
& Halley
Wagner
Van Horn
St. Louis
Co., San
(MO)
(MO); Whitewater
Hood River Co., Dalton
& Halley
(MO). Richland
(014);
Co.,
4546,
(MO). Jackson Co.,
cult. DUSS-79-0590
3932,
(MO)
s.n., cult. DUSS-82-384a
cult. DUSS-79-0577
Solomon
(014),
1, Wagner
605, cult. DUSS
(MO); Flatwoods,
(MO), DUSS-77-0215
Co., Great
(MO) (0)14). Wilkes
(MO) (014).-OREGON:
Halley
McDowell
1698,
no. 162, cult. DUSS
1983
cult. DUSS-79-0582
13771,
Swain
Stubbe 29, cult. DUSS-81-576
(MO), DUSS-78-058
Toronto
s.n., cult. DUSS-79-0592
(014).
(014).
s.n., cult. DUSS-79-0582
at Hwy
Shelton
(MO) (0 12, 111).-MISSOURI:
MEXICO: Bernalillo
(MO) ((0 14).-NEW
Hardin
1978, Pitillo
cult. DUSS-79-0578
Straley
3, 1981, Merchant
s.n., cult. DUSS-82-0512
1975, Whitaker
Co., Hayward,
Lake,
at Hwy
s.n., cult. DUSS-77-0256
cult. DUSS-76-069
s.n.,
Co., Swannanoa,
Shuswap
78?44'W),
Hall,
(49?09'N,
Ootischenia,
cult. DUSS-76-024
3416,
1975, Wagner
36, cult. DUSS-81-570
(44?56'N,
cult. DUSS-77-0162a
280, cult. DUSS-82-0438
on NC Rd 107, S of Inckasegee,
Monteagle,
(CU,
1961,
near Laneville,
(0 14); S Yale
1959, Stubbe
22857,
Alameda
(014);
and Nelson
(MO)
Co., Minden
near Montreal,
(MO).-CALIFORNIA:
NORTH CAROLINA: Buncombe
Troutdale,
Ohio
(MO) (014);
Rd, 1981, Merchant
Ind. Sem. Bot. Gard. Glendon
locality,
Lake Opinion
(MO). Santa Cruz Co., Santa Cruz, Moldenke
Co., Clayton,
(MO)
Castlegar
Park, Cunningham
Haliburton
ALABAMA: Sumter Co., Bellamy,
cult. DUSS-77-0162b
between
s.n., cult. DUSS-82-0508
(MO).-ONTARIO:
84-231
Hardin
14691
Twin,
BRITISH COLUMBIA: Chilliwack
(MO), DUSS-84-101
s.n., cult. DUSS-82-0510
1981, Merchant
(MO) (014); Winlaw,
Collins
1966,
1938,
Co., Bluefield,
Springs, Munz
Tucker Co.,
&
(CU, IND, NY,
s.n. (WVA).
1957, Hinkle
(MO) (0)14); Slocan River Bridge,
s.n., cult. DUSS-82-0511
way,
Co.,
Fernald
Co., Pughtown,
(UNCC).
Co., near Thomwood,
cult. DUSS-82-0500
4545,
1981, Merchant
Airport,
cult. DUSS-84-105
Branch
60049
biennis.
121?54'W)
dia Mtns,
5238
Downs
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. Canada.
U.S.A.
13509
right fork of Upper
of Salt Sulphur
crossroads,
s.n. (MICH, NY, US).
further locality,
& Clark
Co., E of Thurmond,
Fayette
Co.,
Park,
Page Co.,
(GH). Roanoke
(CU, IND, NY, POM, US), Munz
POM, US). Mingo
at Orleans
Natl.
(POM).
ca. 1mi N of Henry,
Munz
(GH).
(CU, IND,
Shenandoah
1940, Yates s.n. (WVA). Mercer
14251
s.n. (WVA). Pocahontas
1898, Steele
(WVA).
gar
Bed
2233
13498
14205
(CU, IND, POM). Hancock
further locality,
IND, NY,
of RR crossing
vic. of Aurora,
Co.,
(CU,
Freer
Rock,
et al. 6844
Siding,
head of
Co., Allard's
near Stuart, Ahles
Co.,
Co.,
4 (MICH, POM).
Skyline,
to Konnarock,
153
13493
13497: Munz
s.n. (WVA). Monroe
& Mansenheimer
McPherson
Harris
Co., Grantsville,
Co., without
Patrick
rd from Chilhowie
Roane
Arlington
(POM), Munz
near Mason's
Co.,
Co., Cades Cove, Wal
Steele
at head of Poo Run, Fernald
Co.,
cult. from Munz
(NY, POM),
Co.,
of
65-89
(BH, IND, POM).-VER
13526
Twp., Black
14143
(GH, US).
& Patton
Co., Mt. Lake, Munz
(US). Madison
CAR
5.4 mi NE
Co.,
(TENN).
Springs,
(CU, PH). Giles
Co., Organ Cove, Munz
s.n. (WVA). Kanawha
Sumpstine
Munz
Stribling
(PH).-SOUTH
Powell
Blount
33099
Co., Appalachian
222
Sussex
(PH, VPI).
(GH, NY, PH, US). Washington
POM).-WEST
Phillips
& Steele
Co.,
13481: Munz
3 mi SE of Petersburg
Co.,
of Gatlinburg,
14224
from Munz
River,
True 52 (PENN).-VIRGINIA:
& Long 9604
Top Mtn, Munz
cult.
IND, POM),
Prince George
(UNCC).
5 mi W
& Frodin
1810
(CU). Fairfield
(UNCC).-TENNESSEE:
(GH). Beaufort
of Otter, Freer
Co.,
NY, POM).
30521
(US, VPI). Augusta
Peaks
212c
Fork & Whitewater
3584
ford Co.,
Lake, Harper
Hunt
Springs,
Devil's
Sevier Co.,
Dinwiddie
Munz
Co.,
Barnet & East Barnet,
Co., between
den, 7th & Garfield
Long
side of Miawatha
Eutaw
Carter Co., Roan Mtn, Norris
(TENN).
Oak Lake, Nease
Munz
Co., W
Area,
Reservoir
Saluda Co., 4.2 mi N of Saluda, Radford
(UNCC).
White
5-79
Co., Pinopolis
179
4538,
(MO)
CAROLINA: Oconee
Van Horn
1704,
cult. DUSS-80-0259
Cache
cult. DUSS
Wagner
(014),
cult. DUSS-82-0487
cult. DUSS-80-0263
(014).-UTAH:
1843,
1, cult. DUSS-81-595
(MO)
Co., Ellicott
Rock
cult. DUSS-80-0258
&
(014);
Rd,
(MO)
(MO) (0)14), Van Horn
(MO)
(014).
Co., Greenville
Marion
Co.,
Farm, N Logan,
180
SYSTEMATICBOTANYMONOGRAPHS
1975, Nye
s.n., cult. DUSS-76-082
DUSS-81-567
(MO) (014).
INGTON:Cowlitz
Co., Woodland,
Gray's Harbor Co., Westport,
4542,
Wagner
s.n., cult. DUSS-79-0559
(MO)
cult. DUSS-82-0499
4543,
Co., Sultan, Wagner
1978, Glencoe
Montrose,
Co., Fort Alkinson,
Jefferson
VIR
(MO) (0) 14).-WEST
(MO) (014);
(014).-WISCONSIN:
10 (MO) (0 14).
(0)14), DUSS-84-1
(MO)
23, cult.
(MO) (014).-WASH
(MO) (0 14). Snohomish
cult. DUSS-79-0563
1217,
Stubbe
Forge,
15, cult. DUSS-81-566
cult. DUSS-76-011
Startup, Wagner
Evans
Creek,
Co., Clifton
Alleghany
Stubbe
cult. DUSS-82-0496
4540,
3632,
(MO) (014);
Co., Seneca
(014).-VIRGINIA:
Co., Strasburg,
Anderson
cult. DUSS-82-0489
GINIA: Randolph
& Rossbach
(MO)
Shenandoah
VOLUME 50
Nee
18069,
cult.DUSS-82-0440 (MO) (014).
KARNTEN: Villach,
Austria.
Ind. Sem. Bot. Gard.
Sem. Bot. Gard.
Liege
cult. DUSS-82-450
0617
(MO).
(MO). Czech
France.
0345
(MO)
(010
DUSS-77-0253
no. 589,
1974
(MO)
Renner,
cult. DUSS-77-0259
TEMBERG: between
(08
DUSS-85-0145
DENBURG: Berlin,
0613
(08
and 06);
(Hirschgarten),
near Beeskow,
Ind. Sem. Bot. Gard.
Dannenberg,
Sem. Bot. Gard. Oldenburg
no. 413,
1982
Oldenburg
no. 486,
DUSS-77-0252
Berlin-Dahlem
cult. DUSS-84-413
1967, Gutte
cult. DUSS-79-0615
no. 992,
(MO)
s.n., cult. DUSS-76-R26,
PtCS: Pecs
and 0)6).-BRAN
(MO) (012
Univ. 1976,
Liichow
(MO); Neuscharrel,
s.n., cult. DUSS-R30
DUSS-84-198
(MO);
(Fuinfkirchen),
collection
Ind.
Ind. Sem. Bot. Gard.
Ind. Sem. Bot. Gard. Braunschweig
Senne
and
cult. DUSS-79
(MO).-HAMBURG:
Oldenburg,
Bielefeld,
1967, Rostanski
(MO)
and 06),
(MO).-NIEDERSACHSEN:
and 06);
(MO); Wolfenbuittel,
Dresden,
(MO). Hungary.
(MO) (08
cult. DUSS-84-252
(08
of 0.
cult. DUSS-77-0251
Renner,
s.n., cult. DUSS-84-247
(MO).-NORDRHEIN-WESTFALEN:
(MO).-SACHSEN:
near Leipzig,
1982
of 0.
collection
(MO) (08
cult. DUSS-77-0260
cult. DUSS-84-239
cult. DUSS-79-0608
1978,
cult. DUSS-84-241
1983, Graberg
1982 no. 333),
collection
s.n., cult. DUSS-83-0146
of 0.
of 0. Renner,
cult.
BADEN-WURT
Germany.
Ind. Sem. Bot. Gard. Berlin Humboldt
near Gransee,
Tarnow
Ind. Sem. Bot. Gard. Hamburg
Boberg,
Standard"),
cult. DUSS-77-0250
collection
of 0. Renner,
Ind. Sem. Bot. Gard. Nantes
cult. DUSS-77-0249
collection
of 0. Renner,
s.n.,
cult. DUSS-77
1975,
Fontainebleau,
1982, Foerster
("O. biennis Miinchen"),
(MO)
(014).-HAuT-RHIN:
s.n., collection
Standard Weissherz").
of 0. Renner,
collection
collection
cult. DUSS-77-0350
Nantes,
III) ("O. suaveolens
near Karlsruhe,
(Munich),
Friedrichshagen
1953); Lieberose
("O. suaveolens
and Wiesental
and 06)
and
(012
(014)
Ind. Sem. Bot. Gard. Basel
Rumersheim,
1947, Gagnieu
(MO)
cult. DUSS-82-0506
(MO)
Ind. Sem. Bot. Gard. Montpellier
(MO)
s.n., cult. DUSS-79
1981,
s.n.,
and 111).-SEINE-ET-MARNE:
(012
(014)
cruciata"),
1II; Stubbe,
(MO)
(MO)
(MO)
(08
("O. biennis
Ldschper
(MO) (014).-LoIRE-ATLANTIQuE:
Muinchen
(MO)
(MO) (0)14)
(MO) (014);
Saint-Laurent-du-Pont,
DUSS-85-1046
Philippsburg
and 0)6).-BAYERN:
1975,
Ind.
no 3567,
1978
DUSS-84-176
cult. DUSS-82-0477
1965,
cult. LILLE-1983
cult. DUSS-77-0265
cult. DUSS-77-0258,
Le Grou,
Bordeaux
1976, Burdovd
Univ. Strasbourg
Linz,
LIEGE: Amay,
Ind. Sem. Liege
Huy,
cult. DUSS-83-0164,
s.n.,
and 06).-HERAULT:
(08
(014);
Ind. Sem. Bot. Gard.
Camargue,
and 211).-IStRE:
(MO), DUSS-85-1048
Renner,
Jean
1983,
ersteinensis),
of 0. Renner,
collection
(MO)
(014).-OBEROSTERREICH:
(MO). Belgium.
SEVEROCESKY: Jetledsk6 Mtns,
Erstein,
Ind. Sem. Bot. Gard.
cult. DUSS-83-0140
(MO)
cult. DUSS-86-2277
cult. DUSS-77-0351
(014).-BOUCHES-DU-RHONE:
(014).-GIRONDE:
Colmar,
no. 1049,
Republic.
BAS-RHIN:
(from seeds of the type of 0.
(MO)
1984
no. 2812,
1974
s.n., cult. DUSS-R3
1971, Melzer
Univ. Salzburg
cemetary,
Stubbe
(MO)
and
(08
Ind. Sem. Bot. Gard.
1982/83
s.n.,
cult.
311);Gundorf
Leipzig 1976,
of 0. Renner, cult. DUSS-77-0264
(MO) (0)10 and
Fiinfkirchen"),
collection of 0. Renner, cult. DUSS-77
211; Stubbe, 1953) ("O. s-xa-suaveolens
0262 (MO) (010 and 2I1, Stubbe 1953) ("O. k-suaveolens
Funfkirchen"),
collection
of 0. Renner, cult. DUSS
77-0263
(MO) (0)10 and 211) ("O. xa-k-suaveolens
Italy. REGION FRIULI-VENEZIA GIULIA. Prov.
Fiinfkirchen").
Goriza:
Grado,
collection
collection
Grado"),
Prov. Undine:
of 0.
of 0. Renner,
Renner,
Ind. Sem. Bot. Gard.
Udine
LIGURIA. Prov. La Spezia: Marinella
MONT. Prov. Alessandria:
Torino:
Torino,
DUSS-84-225
1975
1969, Rostatiski
s.n.,
(MO)
Prov. Pisa: Migliarino
(MO) (0 14). REGION VENETO. Prov. Venezia:
Lithuania.
Pref.,
Frunze,
Lazdijai,
spont.
Sendai
City
in Botanical
near Yorita,
Garden,
Ind. Sem. Bot. Gard. Kaunas
GOSZSZ: Torun
(Thorn),
caulis Thorn"),
1969, Rostaniski
collection
Jesolo,
City, Boufford
of 0. Renner,
1973, Hiibl
& Wood
1978
Skvortsov
no. 1295,
cult. DUSS-77-0254
s.n., cult. DUSS-76-R32
1983,
(MO)
and 111). Prov.
Soldano
(MO)
(014);
(MO) (0) 14).
cult. DUSS-78-0160
(MO) (0)14).
(MO)
(014).
(MO)
(014).
(MO). Poland.
BYD
cult. DUSS-82-0454
cult. DUSS-82-453
(MO)
(014;
(014).-KATOWICE:
1983,
s.n., cult. DUSS
s.n., cult. DUSS-78-0162
s.n.,
s.n., cult.
Oldenico,
Calambrone,
1982, Soldano
s.n., cult. DUSS-77-0430
19859,
Grado").
and li).-REGION
(MO).-REGION
PIED
(MO) (012
Avigliana,
Pisano,
("O. suaveolens
(012
TUSCANY. Prov. Livomo:
1977, Boufford
1979,
(MO)
s.n., cult. DUSS-84-229
(014).-REGION
HOKKAIDO: Ishikari Pref., Sapporo
KYUSHU: Kagoshima
Soldano
(014);
(MO)
Japan.
Kyrgyzstan.
(MO)
1953)
("O. s-xa-suaveolens
s.n., cult. DUSS-84-224
(014).
84-226
Stubbe,
1953)
s.n., cult. DUSS-84-222
cult. DUSS-84-200
s.n., cult. DUSS-84-227
s.n., cult. DUSS-84-223
1983, Soldano
1983,
(014;
Stubbe
cult. DUSS-77-0357
1983, Soldano
Albano,
(MO)
(MO) (014,
no. 993,
de Sarzana,
Isola S. Antonio,
(MO). Prov. Vercelli:
1983, Soldano
Soldano
cult. DUSS-85-1047
cult. DUSS-77-0261
Renner,
1942)
Brzezinka
("O. rubri
near Myslow
1997
OENOTHERA
ice, 1983, Rostaniski
DUSS-76-R13
& Dietrich
(MO)
1975, Rostatiski
Baltic
s.n., cult. DUSS-76-R15
cult. DUSS-84-201
DUSS-84-206
s.n., cult. DUSS-84-204
s.n., cult. DUSS-84-207,
(MO); Jaworzyna,
VAPADOSLOVENSKY: Mail
1105,
cult. DUSS-84-255
(MO); Mail
DUSS-84-256
(MO). South Africa.
77-0419
(08,
1976,
(MO)
1984
no. 535,
s.n., cult. DUSS
Ind. Sem. Bot. Gard. Bratislava
Ind. Sem. Bratislava
dolina,
CAPE PROVINCE: Roadside
(MO)
near Tokai,
and 06).-SODERMANLAND:
(08
cult. DUSS-86-2276
no. 1355, cult. DUSS-79-0661
Clintock
1970, Rostatiski
lII);Wroclaw,
1983, Rostatiski & Dietrich
Prievaly,
Karpaty, Mlynska
Botanic
(MO).-WROCLAW:
and
s.n.,
s.n., cult.
1982 no.
no. 1106,
1982
cult.
s.n., cult. DUSS
1974, Goldblatt
and 11) Spain. TARRAGONA: Macanet
de La Selva, Ind. Sem. Bot. Gard. Barcelona
(MO). Sweden. MALMOHUS: Halsingborg,
Ind. Sem. Bot. Gard. Lund 1976 no. 84,
04,
cult. DUSS-79-0612
cult. DUSS-79-0609
Karpaty,
(014),
& Dietrich
& Dietrich
(MO), Rostaniski
(012
s.n., cult.
(MO)
1983, Rostaniski
DUSS-84-208
(MO)
60 km SW of Wroclaw,
s.n., cult. DUSS-84-199
84-203
(MO). Slovakia.
Jaworzno,
s.n., cult. DUSS-76-R25
1968, Rostatiski
s.n., cult. DUSS-84-189
1975, Rostaniski
(MO) (014).-KRAKOW:
& Dietrich
(MO), Rostaniski
1975, Rostan'ski
(MO); Katowice-Piotrowice,
Ustka,
& Dietrich
(MO), Rostaniski
of Wroclaw,
Garden
s.n., cult. DUSS-84-209
(014).-KoszALIN:
181
(MO) (014).
s.n., cult. DUSS-84-192
s.n., cult. DUSS-76-R2
(MO)
(MO)
United
Wales.
Kingdom.
West
(014).
Vardinge
Ind. Sem. Bot. Gard.
Kers,
BASEL: Briiglingen,
(MO). Switzerland.
Glamorgan:
Stockholm
Ind. Sem. Bot. Gard. Basel
Dyfed:
Carmarthen,
Swansea,
1976
1970, Mc
Pembrey,
1973, McClintock
Jersey Marine,
(014).
CULTIVATED STRAINS WITH DIAKINESIS CONFIGURATION EXAMINED BUT WITHOUT VOUCHER. Canada.
ALBERTA: Ind. Sem. Bot. Gard. Edmonton
Montreal
no. 902,
(012
0347
(012
Germany.
and 111).-BAYERN:
no. 269,
08
(014,
1974
no. 1168,
1169,
cult. DUSS-77-0320
DUSS-77-0324
Golmsdorf
Jena, Ind. Sem. Bot. Gard.
and 06);
cult. DUSS-77-0336
(08
cult. DUSS-77-0335
DUSS-77-0443
Sem. Budakaldsz
and 06);
1974
Horsmar
near Miihlhausen,
cult. DUSS-77-0370
1974 no. 1172, cult. DUSS-77-0372
cult. DUSS-76-R34
(010
(08
Pref., Samani,
Boufford
1975 no. 305,
and 06).-PEST:
Tapi6szele,
1975 no. 2231,
& Wood
19696,
near Wisla,
Ind. Sem. Bot. Gard. Lublin
1975
no. 2610,
Ind. Sem. Bot. Gard. Lublin
1974
no. 2282,
Gard. Dresden
no. 1827,
0368
(014).
(08
and 06).
Union
cult. DUSS-77-0367
1975 no. 152, cult. DUSS-77-0318
cult. DUSS-77-0361
DUSS-77-0363
and 06);
1975 no. 2609,
(014);
Russia.
Dzierby
near Sokolow
Lange Erlen,
Co., Mountainside,
BASEL: Hard,
Ind. Sem. Basel
Moldenke
1975 no. 1474,
31578,
(Moscow)
Ind. Sem. Bot. Gard. Basel
cult. DUSS-82-0378
Krzywa
Warszawa
1974
Gora,
(08
(0) 14).-NORTH
1975
and
Poland.
(012
and
(014);
Je
Ind. Sem. Bot.
no.
1975
1541,
cult.
cult. DUSS-77
cult. DUSS-77-0379
cult. DUSS-77-0339
and 06).
s.n.,
(010
(014).
no. 1510,
1974 no. 2826,
1975 no. 1473,
cult. DUSS-77-0337
1964, Rostaniski
cult. DUSS-77-0375
cult. DUSS-77-0366
Ind. Sem. Warszawa
VOLGOGRAD: Ind. Sem. Bot. Gard. Moskva
Switzerland.
Ind.
Ind. Sem. Bot. Gard.
Ind. Sem. Bot. Gard. Poznan
Ind. Sem. Bot. Gard.
Podlaski,
cult.
and 06),
cult. DUSS-77-0364
(0)14).-OPOLE:
and 111).-POZNAN:
(012
(014).-WARSZAWA:
no. 2580,
(08
cult. DUSS-78-0159
near Deblin,
Ind. Sem. Lublin
1974 no. 337,
1975
spont. Bot. Gard. Vacrdt6t,
Golab
siero Biale,
(014);
Ind. Sem. Bot. Gard. Miihlhausen
KATOWICE: Kasimierz
111).-LUBLIN:
and
(08
Ind. Sem. Bot.
no. 971, cult. DUSS-77-0371
and 06);
(08
no.
cult.
(3 04 and
Rudolstadt,
111);
Ind. Sem. Bot. Gard. Muihlhausen
and 211), Ind. Sem. Bot. Gard. Vdcrdt6t
HOKKAIDO: Hidaka
Japan.
1975
1974
1974 no. 1172, cult. DUSS-77-0321
BUDAPEST: Ind. Sem. Bot. Gard. Budapest
and 111).Hungary.
1975
1974 no. 1170,
Jena 1974 no. 765, cult. DUSS-77-0326
Erfurt,
Ind. Sem. Bot. Gard. Budakalasz
no. 691,
Ind. Sem. Bot. Gard. Halle
no. 766, cult. DUSS-77-0329
(014);
Jena 1974
Ind. Sem. Bot. Gard. Halle
Ind. Sem. Bot. Gard. Halle
near Jena, Ind. Sem. Bot. Gard.
Jena 1974
(012
(014),
near Rosslau,
cult. DUSS
Ind. Sem. Bot. Gard. Olden
Aschersleben,
near Wittenberg,
1974 no. 1171, cult. DUSS-77-0322
Gard. Halle
Gribo
1975 no. 306,
Ind. Sem. Bot. Gard. Leipzig
Leipzig,
Ind. Sem. Bot. Gard. Halle
Sangerhausen,
(014);
06).-THURINGEN:
(08
Kremberg
Jena 1974 no. 764, cult.
Ind. Sem. Bot. Gard.
Oldenburg,
and 06).-SACHSEN:
(08
cult. DUSS-77-0323
(014);
near Usedom,
and 06).-SACHSEN-ANHALT:
(08
1974
University
Jena 1974 no. 763, cult. DUSS-77
Ind. Sem. Bot. Gard. Muihlhausen
and 06).-NIEDERSACHSEN:
cult. DUSS-77-0332
cult. DUSS-77-0331
Tdpioszele
Ind. Sem. Bot. Gard.
Wolgast
and 111).
(012
cult. DUSS-77
Ind. Sem. Bot. Gard. Berlin-Dahlem
Ind. Sem. Bot. Gard.
Frankfurt,
near Potsdam,
near Konigs-Wusterhausen,
no. 761, cult. DUSS-77-0330
04).
and 06);
(08
Eichwald
1975
Ind. Sem. Bot. Gard. Humboldt
Berlin,
and (0)6).-MECKLENBURG-VORPOMMERN:
(08
no. 241,
near Regensburg,
Donaustauf
(014).-BRANDENBURG:
and 06);
(08
(014); Wildau
burg 1975
1975
cult. DUSS-77
012
Caen
Strasbourg
and 111).-LOIRE-ATLANTIQUE: Nantes,
Ind. Sem. Bot. Gard. Nantes 1975 no. 564,
and
Ind. Sem. Bot. Gard. Bordeaux
1975 no. 1446, cult. DUSS-77-0341
1I,).-LOT:
BADEN-WURTTEMBERG: Karlsruhe,
Ind. Sem. Bot. Gard. Karlsruhe
1975 no. 1042,
DUSS-77-0328
0325
Ind. Sem. Bot. Gard.
cult. DUSS-77
cult. DUSS-77-0315
77-0334
Ind. Sem. Bot. Gard.
(014).-QUEBEC:
BAS-RHIN:
Ind. Sem. Bot. Gard.
(014).-CALVADOS:
1974 no. 623, cult. DUSS-77-0317
Berlin
France.
(014).
no. 475,
(014,
no. 2501,
no. 136, cult. DUSS-77-0312
cult. DUSS-77-0313
cult. DUSS-77-0349
0342
0319
no. 332,
1975
U.S.A.
(08
NEW JERSEY:
CAROLINA: Jackson Co., Bal
SYSTEMATICBOTANYMONOGRAPHS
182
sam Gap,
1978, Pitillo
cult. DUSS-81-595
s.n., cult. DUSS-79-0593
2552
Lake,
Boissevain,
Morine
W
181
Bossenmaier
of Fredericton
(GH); Shediac
ramichi River,
of Cape Bald, Bassett
(CU, GH,
PAC, PENN);
St. George's,
Donovans,
Rouleau
(MT).-NOvA
verly Rd,
1830, Forward
& Mulligan
Bassett
Brook,
Mollichicgeck
Intervale, Margaree,
(CAN); Halifax,
Liscom's
TRT); Kenora,
Forest Reserve,
near Rainy
& Baldwin
City, Comack
Ferry, Rideau
Fletcher
Murillo,
(DAO);
Garton
1121
(DAO); Thunder
Bay, Gillett
ham 266
(DAO),
Tremblay
Lake, Harrison
Waterloo,
& Graham
2309
477
Donald
(DAO);
s.n. (CAN); Hamilton,
Macoun
44464
133
(NY); Port Colborne,
3-6 mi SE of Hawkesburg,
1814, Moss
Munz
17523
(TRT); 66 mi NNE
(TRT); Queenston,
ers Lake,
Skelton
Dale
4223
Bay,
Spradboron
Tozer 532
Minshall
on Rte.
11 (OSH);
396
605
(CAN);
Frontenac,
(DAO);
Peel, Hart House
WARD ISLAND: Queens,
Fernald
PH); Queens,
N. Rustico,
& St. John 7827
Mt. Albion,
Farm, Vickers
1967, Winterholder
Fernald
Erskine
1267a
(GH); Queens,
et al. 7830
Minshall
of Kitchener,
1870,
Sandwich,
& Rolland
(MT); Lennox
Montgomery
Bay, Porsild
of Orillia,
Senn 1274
Haldimand
263
7961
Reznicek
(GH); Mid
Taylor
Lake, Soper 5259
et al. 142
s.n. (TRT); Scugog,
3 (TRT); Bruce
20 mi
& Reznicek
4780
(TRT); Moose
143
Fernald
Erskine
S of Soy
North
2 mi E of Burketon,
Watson
3173
s.n. (CU).-PRINCE
1497
et al. 7828
Soper &
factory, James
(MT); Nipissing,
Howdenvale,
1919, Young
(DAO); Charlottetown,
1898, Scott s.n.
Forest Reserve,
Tozer 141 (DAO);
Peninsula,
s.n. (CAN); Cobourg,
Charlottetown,
(MT),
Park,
(CAN);
(DAO); Haliburton,
Twp., E North
&
955 (DAO); Prescott,
1894, Scott s.n. (TRT); Toronto,
2 mi SW Brouget,
Ganonoque,
(CAN, NY);
144
Mc
Islands, Mc
Alfred, Marie-Victorin
Kemptville,
11 mi WSW
S end of Beaver
1931, Taylor
Bay
et al. s.n. (CAN); Belleville,
(ND); Constance
Simcoe,
(TRT); Northumberland,
(DAO, GH, TRT, US); Barry Sound,
62471
393
Cobalt,
(GH, TRT); Lincoln,
(BH, IND, POM); Kent, Lake Erie, Rondeau
(DAO); Victoria,
s.n. (TRT); Russell,
1898, Scott
& Skelton
17522
Jennings
(CU); Timiskaming,
& Rolland-Germain
1mi W
(DAO); Waterloo,
Island,
SE of Nipigon,
21704
near
Ivery s.n. (TRT);
1911,
Parry Sound, Georgian
Prescott,
(DAO); Grenville,
Island,
F.S., Gra
Sydney
(NA); Cunningham
10736
44465
St. John's, Nieuwland
101, Reardon
15003
35, 1971, McIntosh
E of Jackfish,
Haycock
(DAO); Lake Superior,
(DAO);
Park, Macoun
691
(DAO);
James 3049
8287
Stewarton,
SW of Kinkamp,
2.5 mi
467
(CAN,
83 (TRT); Leeds,
(GH, TRT, US);
303
(CAN); Algonquin
(GH, NY);
3132
1614
Cruise
(DAO); Carleton,
Lake, Krotkov
s.n. (CAN);
of Hwy
2037
& Jennings
Jennings
Port Hope, Marie-Victorin
1469
of Port Arthur, Roske
Sudbury, Worthington,
Alberton,
Macoun
Minshall
Bay, Taylor et al. 146 (MT); Muskoka,
TRT);
10815
s.n. (DAO);
Haddow
Jenkins
(CAN,
& Bragg
s.n. (CAN); London,
s.n. (TRT); Port Sydney,
Queenleen
s.n. (UAC); Kent, Chatham, Munz
(BH, IND, POM);
ESE of Chapleau
Lake, W
Bassett
(DAO);
6669
s.n. (DAO); Ottawa,
6848
1.5 mi E of St. Thomas,
(DAO).
1885
& Porsild
Marburg,
Hill, Gogo
(DAO); Midland,
Superior,
Durham,
1-2 mi SW of Morven,
Addington,
dlesex,
Austin
(DAO, MT, US),
Sugarbush
1870, McConnell
Macoun
7, Gillett
1930, Hartley
Peninsula,
Chatham,
(CAN); Haliburton,
Macoun
Germain
of Lake
(MT); Bruce
Anderson
(DAO); Curran, Dunbar
61, Garton
on Hwy
2611
Smith et al. 3399
1883, Dawson
River,
1971, Gad
2 mi S of Newington,
Stormont,
NW
(WTU);
10866
(ALTA, CU, US);
286
Groh
118 (TRT); Norwich,
Island, Ma
(CAN); Pic
s.n. (TRT); York, 2.1 mi S of King
1878, Fletcher
S of Hwy
(DAO); Glengarry,
James 263 (DAO); Elgin,
Bridgeport,
Jenkins & Ilman 3280
& Jennings
9798
28 (DAO); Beaucage,
Cleonique-Joseph
0.5 mi
Bay,
5 mi E of Marmora
Sable
2051
s.n. (TRT); Norfolk,
Pool,
(PH); Wa
s.n. (DAO); Auburn,
Shchepanek
Sault Ste. Marie,
673
339
Distr.,
(CAN, GA);
Baldwin
Ogoki,
& Coleman
(NY); Ottawa,
E shore of Gloucester
s.n. (GH); Hastings,
19125
Village,
127 (MT); Ingolf, Denike
1931, Ayre
& Long
et al. 21995
Fernald
1936, Groh
(TRT); Kapuskasing,
Old Man's
Pond,
(MT); St. John's Northern
near Cape North
1802, Cosens
Murray
(CAN); West
Island, Fernald
Lake,
1938, Coleman
1256
NW Mi
12208
Scoggan
S end of Seloam
(CAN, TRT); Algoma,
14198
1111 (DAO); Retigouche,
Port Montou,
5481
& Long
Bathurst,
(GH); Nappan,
(UBC); Cochrane,
45 (DAO); Ottawa,
Dearness
Simcoe,
Shirley's
Calla
Davidson
Park, Edmondson
695.5
1953, Gates
8418
(TRT); Port Carling,
71 (MT); Oakland,
Rideau
2524
Bailey
Baldwin
River
1096
(DAO);
6477
Cape Breton, Macoun
Island, Krotkov
16 (TRT); Brampton,
CU, TRT); St. Catherines,
Dearness
River,
Lookout,
Cain 436
Bear
Rouleau
1894, Mc
BRUNSWICK: 18 mi
Jct., Fernald
Park, Monro
& Bassett
Central
(NY); Stettler Dis
(DAO); Whitewater
Portage-la-Prairie,
(DAO); Governor's
51.35
Sanctuary,
N Aspy
s.n. (TRT); Victoria,
Sioux
(DAO); Otterville,
Game
236
(DAO).-NEWFOUNDLAND:
Brook,
Gates
s.n. (DAO); Big
Nat'l.
1,
(014).
(MO).-NEw
Cape Tourmentine,
681
SCOTIA: Queen's,
s.n. (UBC); Middleton,
1937, Groh
Land Reserve
2935
(CAN);
1810
(DAO); McAdam
Mulligan
& Breitung
coun 78527
Wild
9038
Stubbe
Sandy River,
LYve & Love 5133
Clokey
2852
(TRT); 50 mi E of Shediac,
10 mi E of Corner
tou, 1891, Sheraton
Dawson
& Mulligan
Co.,
Brinkman
District,
campus,
Sifton,
River
s.n., cult. DUSS-81-547
s.n. (GH); Kouchibouguac
s.n. (GH);
ONTARIO: Temagami
(DAO);
2.5 mi S of E Florenceville,
5804
7181
339
Bassett
1914, Hubbard
Sandercock
6 mi W
morland,
(MIN); Turtle Mtn,
Peak, Rowe
2 Hwy,
near No.
Cape,
717 (CAN); Carleton,
Malte
Fort Gary University
(US).-MANITOBA:
s.n. (DAO); Riding Mtn
1979, Kral
Co.,
ALBERTA: Craigmyle
REPRESENTATIVE SPECIMENS. Canada.
trict, Brinkman
Hood
(014).-OREGON:
Marion
(014).-TENNESSEE:
VOLUME 50
(DAO, MT);
ED
Prince,
(CU, GH, MICH,
(CAN, CU, GH, MIN, MT, NY, PH, US); Bideford,
(NY,
MT,
1948, Taylor
OENOTHERA
1997
s.n. (DAO).-QUEBEC:
(MT); Arthabaska,
Val Morin,
Bassett
Bromptonville
Black
et al. 10448
Joliette,
Fabius
(CAN);
Shefford
Cap Tour Mente,
(GH, MT);
de Bellevue,
Groh
Co., Granby
Gates
41.35
s.n. (DAO);
Notre-Dame
de Ham, Hamel
Kucryniak
tawa, Laferrrre
120 (DAO);
(MT); Quebec
Marie-Victorin
St.-Vallier,
s.n. (KYO); Le Bic
Marie-Victorin
40
63 (DAO,
land-Germain
MIN,
FSU, MT,
MT, US);
de la Perade, Marie-Victorin
reville, Marie-Victorin
(DAO, MT,
US);
& Rolland-Germain
St.-Maurice,
10806
(GH); Quebec,
pedia, Marie-Victorin
Trois-Riviers,
33825
land-Germain
Minshall
Greenville,
Anne,
Munz
144
Terrebonne,
(MT); Angers,
25572
(CAN, GH, MT);
14735
from
seeds
col.
Munz
17510
(BH); Matapedia,
of St. Vallier, Munz
St-Jerome,
17512
(DAO); Gaspe,
Rigand,
Marie-Victorin
Rousseau
(DAO, MT);
31256
Gatineau
(MT); Argenteuil,
430
(MT); cult. from
Munz
Munz
Compton,
des Rapides,
Longueuil,
17511
(POM);
(BH, IND,
Bishopton,
Robert
7867
80 (MT); Mont
Pleureuse,
Park, Skyline
from Sainte
14761
Rolland-Germain
Rouleau
(MT); Anse
Mata
& Rol
Marie-Victorin
St. Vallier,
Routhierville,
(MT); Laval
121
124
US),
& Rolland-Ger
et al. 45686
(BH, IND, NY, POM);
Pier
(MT); Bonaventure,
St. Bruno, Morin
from
(DAO); Chambly,
Grande Vallee,
Roy 3905
(DAO, MT,
cult. from seeds col. by Gates
by Gates
6025
19285
Rolland-Germain
Vaudreiul,
(IND, POM);
Rolland-Germain
122
28321
Marie-Victorin
s.n. (CAN);
1913, Momalte
Ste.-Anne
& Rolland-Germain
Petite, Cascapedia,
Munz
cult.
de la Pocatiere,
3.5 mi W
Bellechasse,
Head, Rousseau
St. Vallier,
POM);
Sainte Anne
(DAO);
(MT), 7869
(IND,
iles de laMadeleine,
Bellechasse,
(DAO); Cascades,
Marie-Victorin
Riviere
Champlain,
MIN, MT, NY, US); Yamaska,
& Rolland-Germain
(DAO, MT);
Asbestos,
& Rolland-Germain
113 (DAO, MT, US); Que
& Rolland-Germain
28667
from Cape Tourmente,
14741
Kamouraska,
POM);
(CU, DAO);
2626
seeds col. by Gates
(DAO, MT),
& Rol
Richmond,
115 (DAO, MT);
Nicolet,
Marie-Victorin
Marie-Victorin
(DAO, MT);
& Rolland-Germain
(MT, US);
Ile de l'Etang-du-Nord,
Sainte-Foy,
28478
120
Saint
104 (DAO, FSU, MICH,
Marie-Victorin
116 (DAO, GH, MICH,
73
Soulanges,
Marie-Victorin
Caledonia,
102
Baie Saint-Paul,
Luskville,
& Rolland
& Rolland-Germain
70 (DAO, MT, US);
114 (DAO, MT),
& Rolland-Germain
Hull,
Marie-Victorin
& Rolland-Germain
Marie-Victorin
(DAO, MT, US);
(DAO);
Marie-Victorin
Prescott,
Ste.-Th6rese,
& Rolland-Ger
41
46
(MT),
Saint-Michel,
& Rolland-Germain
& Rolland-Germain
(DAO, MT, US); Cap-aux-Meules,
main
81 (DAO, MT, US);
Lanoraie, Marie-Victorin
St.-Michel,
Marie-Victorin
C6te
de Saint-Sulpice,
107 (MT); Charlevoix,
(DAO, MT, US); Bellechasse,
44
10137
1929, Marie-Victorin
Terrebonne,
& Rolland-Germain
& Rolland-Germain
Marie-Victorin
Meilleur
(MT); Bellechasse,
Marie-Victorin
Chichester,
Marie-Victorin
Domaine
& Rolland-Germain
Pointe-Gatineau,
Pontiac,
TRT, UC);
La Trappe,
Cabano,
s.n. (MT); Ot
(DAO); Rimouski
2049
25 (DAO, MT);
& Rolland-Germain
Thurso, Marie-Victorin
& Rolland-Germain
bec, Cap-Rouge,
8 (DAO); Deux-Montagnes,
60 (DAO, MT,
TRT, US);
Michel
(DAO);
Lake, Hasem
1934, Lanouette
286.2234
s.n.
1927,
1445
Lake, Groh
(DAO); Temiscouata,
Lemieux
St. Antoine,
& Rolland-Germain
(d'Abitibi),
St. F6licien,
(DAO);
(CAN); Gaspe, Mont-St.-Pierre,
(MT); Lotbiniere,
Marie-Victorin
86 (DAO, US); Berthier,
Marie-Victorin
108
1504
Villemontel
Papineau,
Marie-Victorin
Clement,
60320
29 (DAO, MT, US);
& Rolland-Germain
(DAO, MT, US);
33 (MT); Quebec,
Marie-Victorin
Pointe-a-la-Garde,
Marie-Victorin
Laurent
16922
(MT); Ste. Anne
(UC); Wakefield
3321
(CAN); Mont-Royal,
Macoun
& Rolland-Germain
(Rimouski),
(DAO, MT, US);
Germain
Michel
& Rolland-Germain
Bonaventure,
Lacass&e A33
Granby,
18275
& Brisson
13360
Brompton,
1932, Gates
569
Gauthier
14228
Boivin
Forest
de Tilly,
Johnson
Jenkins & Bayly
(DAO);
(BH); Rouville,
Lac Brompton,
de Brandon,
Gillett
Chemin
13846
& Ernest
near Lascellas,
2121
(MT); Megantic,
Cody & Kemp
(DAO);
Saint-Antoine
Mill,
187
Les Eboulements,
22636
Dutilly
(NY); Laviolette,
& Hamel
Lac Saguay,
Richmond,
St. Gabriel
679
Hendley
Charlevoix,
Lotbiniere,
River, Massham
(BH); Wakefield,
Co., Cape-Rouge,
(MT, NY);
(DAO); Lac Sylver, Hamel
Shefford,
3303
Lapage
(DAO);
of the center of district,
& Tardif 135 (MT); Saint Placide,
Co., Rimouski,
main
14816
Ungava,
s.c. s.n.
1978,
(DAO); Terrebonne,
Bissonnet
Dore
5752
s.n. (DAO);
1932, Groh
14 mi NW
(DAO); Gatineau,
TRT, UBC);
2005
Bassett
Mont-Louis,
Factory,
(GH); Berthier,
(GH); along Lapeche
Barnhart,
of Dolbeau,
Ste.-Marie,
Grenville,
Anderson
s.n. (DAO); Labelle,
Fabius
Frere 292
10.34
Bromptonville,
MT,
(MT); Mont
de Route,
Fabius,
Gates
DAO,
(UC); Moose
Bord
(DAO);
Royal,
9 mi NE
s.n. (UBC); Gaspe,
135
147
& Beaulieu
5548
Mount
1951, Cayouette
Coiteux
(MT); Argenteuil,
St. John, Mistassini,
Roberval,
(CAN, COLO,
St.-Jovite,
Ste.-Beatrix,
Doucet
Sherbrooke,
(DAO);
1956, Beaulieu
(Richmond),
Champ,
398
1982
(MT); Cite Terrebonne,
(DAO);
(RSA); Lake
s.n. (MT); Montreal,
& Hamel
Lake, Blais
1416
2366
Allyre
1940, Major-Bornabe
La Tuque,
Smith et al. 9988
South River,
Antigonish,
Nicolet,
183
Owls
Rousseau
Trail, Senn 2097
32280
(DAO);
Sherbrooke,Lennoxville, Tessier 601 (DAO).-SASKATCHEWAN:
Carlyle Lake, 1831,Anderson s.n. (CAN);
McKague,
Breitung
Rd, Harms
18249
ALABAMA: Baldwin
(SMU). Dallas
Garden's
Co.,
entrance,
Co., Fields,
Graves
8593
(DAO);
(CAN, DAO,
Co.,
3 mi E of Beauval,
GH); Weyburn,
near Loxley,
Burkhalter
5 mi E of Selma, Demaree
Clark & Landers
1918
8056
(MO). Lauderdale
Co.,
172a
6230
52971
(UNCC).
17896
Harms
Sausan
(MO). Cherokee
(DS). Franklin
Greene
11 mi N of jct. of Ile-a-La-Crosse
(DAO);
(NY); Assiniboia,
Co., W
1903, White
Co.,
Co., E of Dismols
Creek,
of Smith Lake, Harper
cult. from seed of Sheffield,
s.n. (TRT). U.S.A.
2 mi N Leesburg,
Bartlett
3653
4522
Kral
upstream
3372
from
(MO). Jackson
(CU). Mobile
Co.,
184
SYSTEMATICBOTANYMONOGRAPHS
Citronelle,
13351
Munz
Montgomery,
(CU, IND, NY, POM, US, WTU).
3521
Bartlett
Tuscaloosa
Co., Tuscaloosa,
Farm, White
POM,
SMU, WTU).
20339
(BH, MO,
18367
Jefferson Co., Pine Bluff, Demaree
Demaree
36085
Demaree
well,
s.n. (TEX).
IND, POM, US).-CALIFORNIA:
Alameda
1985,
Imper s.n. (HSC); McKinleyville,
TEX),
1343
(TEX). Riverside
s.n. (CAS).
coy, 1965, Granert
Danielson,
3 mi N of Broadkill
Louis
5123
Larsen
Beach,
1.5 mi W
(US, WTU).
s.n. (FLAS).
Seminole
Holmes
5 mi
on W Hwy
of Wakulla,
82, 1970, Shaw & Whipple
Co.,
15.1 mi NNW
ning 2209
(CU, GH). DeKalb
nah River,
Coile
Hall Co.,
et al. 1139
5.1 mi NW
Co.,
Grady
Co.,
449121
(PH). Jackson Co., Cloverhurst
Thomson,
Bartlett
1029
Co., Augusta,
der Co., NW
Prese
55
Daviess
(MICH). Rabun Co.,
1907, Cuthbert
of Cairo,
Elias
(US). Gallatin
Co., Canyon
Co.,
Camp,
nois State Park, Thone
103 (F,MO).
son Co., Edwardsville,
Demaree
Co., Athens,
Rapp
Lansing
Lawrence
51110
2 mi. SE of Custer
Co.,
Stark Co., Valley
Twp.,
Park, Steyermark
64858
Faircloth
1929, Pyron
3301
Blackford
Nest,
Friesner
7193
(FLAS).
Lane
1085
Wiegand
& Man
3 mi
Savan
(UNCC).
S of Wellston,
(UNCC).
19591
LaSalle
62-980
1912, Nason
Co.,
Richmond
Alexan
Co., Lisle,
(ASU).
(MIN, TAES).
Jo
(MO).
Co.,
Illi
(FSU). Madi
s.n. (F). Menard
Pope Co., Glendale,
Co., Athens,
Co., Fuller & Fisher
Co., NE of Glycerine
7599
578
Palmer
Henderson
line
(ID).
Jones 878 (UNCC).
14 (GA, MICH).
(POM). Sangamon
s.n. (PH). Union
(F).-INDIANA:
Co., 0.25 mi E of Whippoor-Will's
15427
(CAS).
from county
(POM). DuPage
2439
Co., Algonquin,
Chase
73117
6585
& Vickers
Co.,
Co., Rosiclair,
1.5 mi E of Lawrenceville,
Ridgway
21.3 mi
7654
36154
Keil
Curtiss
1960, Moore
s.n. (DUKE). McDuffie
Wehmeyer
Peattie
Co.,
Pollard,
s.n. (NY). Elbert Co.,
along GA 28, Bozoman
McHenry
Plymouth,
City, Hood
(TEX). Houston
(MO). Hardin
Long Bridge
Clarke Co., Athens,
Co., Quincy,
Co.,
Sussex
(NA, VPI).
2 mi NE of Moreland,
Co., E of Galesburg,
1896, Chase
Co.,
of Cochran,
1894, Small
19178
Peoria Co., Peoria,
Bridge,
Sati
State Park, Godfrey
Bibb Co.,
SE of Whigham,
Adams
17045
(DS, UNCC).
Co., near Cowford
71 (IND, MO, US). Brown
of Stone Mt.,
Co., Glenview,
(WIS). Knox
& Sherif 80 (F, GH).
88 (MT). Richland
Sherf 715 (PH, UCLA).
Co.,
Palmer
314
Co.,
Golf Club, Athens,
Cook
(PH,
1922,
Ranch,
Lake, McFarlin
Co., Orange
(UNCC).
& Duncan
391
near Jacksonville,
Co.,
3.5 mi WSW
Co.,
S of Big Creek
(A, MO).
Equality,
Peters
Volusia
21485
Blake
San Francisco,
(FLAS, GH). Citrus Co., 4 mi E
Co., Torreya
(TEX).-GEORGIA:
4.75 mi
s.n. (FLAS).-ILLINOIS:
1522
Liberty
(GA). Coweta
Adams
Branch,
Ainsworth
et al. 11093
Polk Co., Crooked
Bleckley
1mi S of Orleans,
(NY, POM). Hartford
250
Lake Co., Mt.
and summit
slopes
(FLAS).
of Flowery
103
Bridges
Small
(FLAS).
(CU,
(IND, POM). Windham
(FLAS).
Canal, Duncan
of Valdosta,
13363
Ford 3548
s.n. (UNCC).
3 mi SE of Old Ogeechee
Cook Co.,
13386
Duval
63-1696
De
14255
(MO). Ventura Co., Hawley
(FLAS).
4509
Baltzell
Henderson
Co.,
2270
(FLAS).
14242
Munz
Munz
Co.,
OF COLUMBIA: Southend
(FSU, NA).
5350
Baltzell
(MO,
Crow
Sevier Co., Horatio,
near Winslow:
Kent Co., Smyrna, Holmer
Baltzell
52462
Godfrey
from Waldron:
Co., Los Angeles,
London, Munz
of Hallandale,
River,
15503
Junction,
(MO). Van Buren Co., Clinton,
Co., Danbury, Munz
Co., New
3 mi SW of Bonifay,
Co.,
3416
(PENN).-DISTRICT
Co., W
SW of Sanford,
2 mi W
Co.,
Chatham
Broward
Co., E of Lake Weir,
Co.,
Wakulla
1217
of Withlacoochee
Leon Co., Tallahassee,
(FSU). Marion
London
Demaree
Munz
s.n. (MIN). Santa Barbara Co., Santa Barbara,
1902, Congdon
(CU, IND, POM).-DELAWARE:
764 (NY).-FLORIDA:
of Inverness,
New
(NEBC).
13561
Co.,
Polk Co., Howard,
s.n. (POM). San Francisco
Fairfield
(DS). Nevada
56975
Co., Madison
from Mt. Gaylord
Santa Cruz Co., Santa Cruz, Moldenke
267
13383
Munz
Island,
Logan Co., Blue Mt.,
(CU, IND, POM).
17 (WVA). Los Angeles
1934, Reed
s.n. (WIS).-CONNECTICUT:
Andrews
Southington,
Co., Riverside,
Demaree
179
De
OKL, RSA,
NY, POM, UC). Miller
s.d., s.c. s.n. (UC). Humboldt
Co., Berkeley,
Maxfield
San Joaquin, Bouldin
s.n. (CAS).
13560
(GH, KANU,
Prairie Co., Hazen,
13563
De
(DS, SMU, UNCC).
SMU, TEX).
cult. from Munz
Desha
(UNCC).
Jackson Co., Newport,
24774
St. Francis
Sharp Co., Hardy, Emig
cult. from Munz
Co.,
46774
(TENN, UNCC).
(SMU).
12 mi S of Ft. Smith, Munz
Co.,
1935, Brinkley
(MO). Washington
9423
Demaree
OKL, RSA,
13 mi NE Monticello,
VPI).
RSA,
51 (SMU).
31379
Tucker 6115
Co.,
(GH, ISC, MIN, MO,
Co., Norman,
Scott Co.,
(MO, NY, POM).
Sebastian
Hill
Demaree
Co., Noland,
21566
(CU, IND, POM, US).
20648
Demaree
Pope Co., Russellville,
Randolph
SMU, UNCC,
(ISC, MIN,
(CAS, SMU). Montgomery
79 (US). Pike Co., Murfreesboro,
(CU, IND, NY, POM).
1972, Pollard
25237
Co., Flippin, Demaree
(GH, SMU). Marion
NY, POM, WTU).
Walther
Demaree
Co., Sedgewick,
OKL,
(GH, KANU,
(GH, KANU,
Pine, Demaree
Co., Mountain
(GH, KANU,
Lawrence
73981
Garland
25806
SMU, UC). Drew
NY, POM,
22644
Clay Co., Coming,
near Lake Fort Smith,
Springs,
(BH, MIN, MO,
Demaree
Jonesboro,
42914
Prescott, Hollister
Henderson's
Co.,
Island Chute
Co., Big
Demaree
WVA).
SMU).
Co., Doddridge,
maree
Craighead
from
(UNCC).
(DS, IND, NY, POM). Bradley
Co., Tinsman,
(SMU, UNCC,
Demaree
41869
ley's Ridge,
54822
Co., Lake Shepherd
13776
Calhoun
SMU).
13558
Munz
collection
5950
& Wagner
Arkansas
NY, US).-ARKANSAS:
(CAS, GH, MIN, MO,
SMU, UNCC).
13567
POM,
Whetstone
Springs,
Co., Bestwater,
Demaree
(MIN, NY, POM, US, WTU).
20405
Demaree
(CAS, MO,
Crawford
Hot Springs Co., Malvern,
maree
(BH, FLAS,
NY, POM, UC, WTU).
Demaree
13650
3982
Co., cult. from Bartlett's
Montgomery
of Lacey
117 (NA). Benton
Clark Co., Amity,
SMU, UC, UNCC).
Co., McGehee,
maree
Harper
River Refuge, Miller
Jersey, Demaree
Co.,
Co., N
(CU). Morgan
VOLUME 50
(CAS, DUKE,
Lansing
204
&
(F). Will
Works,
Deam
PENN,
POM,
1997
RM,
OENOTHERA
SD, SMU,
16147
TENN,
(CAS, UC).
of West
Fork, Deam
Elkhart, Demaree
Gibson
2628
ington,
7178
Deam
49832
(POM).
Friesner
View,
49493
ftW
13908
(GH, NY).
51308
man Lake,
Arnolds
2354
Co.,
Island near Electric
1892, Stewart
11498
Lake, McGregor
31991
s.n. (KANU).
Stephens
TUCKY: Ballard
Co.,
Junction, Munz
13539
(CU, DS,
Hickman
1.2 mi W
Co.,
tween the Lakes, Ellis
from Munz
MIN,
NCSC,
POM). Warren
ND,
(UNCC).
Calcasieu
Parish,
merfield,
Thieret 24655
Feliciana
Parish,
Moore
Parish,
& Moore
Bastrop,
(SMU). Richland
Vincent
& Erbe
stook Co., NW
Furbish
6422
Parish,
2730
14115
Co., W
Hancock
Co., Somerset,
Rockcastle
13540
East Baton Rouge
11531
Lafayette
Shinners
Seymour
Co., Orland,
Ogden
& Moldenke
Parish,
4938
South Poland,
25971
(DAO, TEX).
30998
et al. s.n. (CAS,
(DS,
SE Gate Rd, Balogh
Parish,
1565
(NY). East
1mi SE of Clay,
Shinners
22008
Parish, 0.5 mi S of Varnardo,
1893, Furbish
Lincoln
211
3.5 mi E of Sum
32 (PAC). Morehouse
Desselle
of Natchitoches,
(MO). Franklin
(TEX, UWL).
IND,
13531
1mi N of Williams
Co.,
Jackson Parish,
(GH, SMU). Washington
Co.,
(CU, IND, NY, POM).
2 mi S of Baker, Brown
5.2 mi S of Lafayette,
& Svenson
be
cult.
(BH, POM, WTU).-LOuIslANA:
WTU).
SSE
4645
Lyon Co., Land
Vernon, Munz
(ND). Claiborne
2.2 mi
13537
Braun
IND, NY, POM),
13541
4 mi N of Mt.
Parish,
Parish,
24642
Androscoggin
9586
(TENN, UNCC,
Parish,
UWL).
(CU, IND, POM). Whitley
& Correll
Correll
(GH, US).
IND, NY, POM, US).
(CU, DS,
Munz
14260
4113
(CU, DS, POM).
Covington,
1941, McFarland
Bossier
21231
(POM). Natchitoches
of Newry, Moldenke
Co.,
(KANU).-KEN
13533
Co.,
(UNCC,
10 mi E of Benton,
Co.,
Pulaski
(MO).
1mi E of Paola,
S mi SW of Glasgow
(BH, DS,
13532
(GH, SMU).
Thomas
(FLAS).-MAINE:
375
13530: Munz
FSU).
S side of Delhi,
& Fuller
S of
State
Hart Co., Conmer, Munz
14278
Shinners
branch of St. John's River,
s.n. (NEBC).
(NY). Oxford
of Clinton,
Munz
3 mi
Co.,
1326
Croat
29729a
Munz
Kenton
cult. from Munz
near Holmwood,
(GH, MT, UC).
Demaree
93, Fuller
Green, Munz
of Gibbsland,
(DS, DUKE,
10 mi W
(MO, US).
Co., 9 mi S of Berea, Munz
UC, WVA).
(CU, IND, NY, POM),
4.1 mi W
Parish,
Brown
Smith & Hodgdon
251 (UNCC, UWL).
215
Story Co.,
Co.,
s.n. (UNCC). Doniphan
Barren Co.,
8 mi S of Lexington,
col. from Magnolia:
(TEX, US).
of Bowling
(UNCC).
of Murray,
SW of Os
(UNCC).
2 mi N &
Co.,
Hill, McGregor
317
Co., W
Fayette Co.,
(POM). Marshall
1259
1750
(US).-KANSAS:
Miami
(KANU).
SE of Maple
Calloway
453 on Hwy
POM, TENN,
2 mi W
Co.,
13530
burg, Munz
Beinville
14221
Runyon
Douglas
jct., Fuller
McFarland
797 (UNCC). Madison
13532: Munz
Co., Rockdale,
MO,
of Hwy
12807
border, Fuller
Co., Clinton,
Carter
Co., N of Baldwin,
NY).
1938, Loufek
Co.,
s.n. (ISC). Wapello
1963, Clarkson
1 mi
(SMU).
La Rue Co., cult. from seed Munz
(POM, US).
Owen
27677
G3871
Co.,
Co., Eilers
s.n. (ISC). Palo Alto Co., S beach
1.5 mi SE of Pittsburg,
64/60
IND, POM).
Shinners
(ISC). Floyd
(UNCC). Mahaska
Co., LeMars,
Somes
(ISC).
(ISC). Dickinson
(KANU,
Co.,
of Hwy
628
1911, Shimeck
Junction,
Brooks
313, 6.3 mi from Tennessee
IND, POM).
Livingston
Wabaunsee
0.5 mi N of NW
Co., Bardwell,
Fulton Co., Hwy
(CU,
Co.,
23490
4.5 mi E & 4 mi N of DeSoto,
Co.,
Carlisle
Crawford
Cloud, McGregor
1729
8656
(MO). Johnson Co., Lake Macbride,
1929, Cratty & Aikwau
Fort Dodge,
Co.,
8654
Wolden
in Free
1919, Bode
Co., Lamout,
of Floris, Hayden
Co., Davidson
(ISC). Plymouth
Co., Afton
(ISC). Webster
(KANU).
1mi N of White
1974,
10101
Park, Hayden
s.n. (MIN). Union
1875
island
Co.,
s.n. (MIN). Audubon
Lake, Hayden
1mi W
Deam
Deam
3 mi SW of Lebanon,
Co.,
shore of Trumbull
Co., Estherville,
Co., Tracey,
1mi E of Ashley,
Co.,
(ISC). Buchanan
St.
(IND).
(IND). Tipton Co., Goldsmith,
1919, Shimek
(TEX). Louisa
s.n. (ISC). Marion
of Lost
Hayden
2875
7431
1.7 mi SE of Pleasant
Co., NE of Postville,
263
of Marshall,
of Porter, Deam
Deam
(POM). White
Co.,
Emmet
5 mi W
Co.,
(IND, POM). Warren
Co., Getty
Deam
of Thayer, Deam
4 mi NW
55423a
Co., 7 mi NE of Jefferson, Davidse
Ames,
Johnson
s.n. (MIN, WIS).
52841
Deam
s.n. (ISC). Clay Co., N
s.n. (UT). Davis
Lee Co., Davidson
1920, Shimeck
Cliffland,
52814
Allamakee
IND). Shelby
s.n. (IND). Steuben
Deam
of Bluffton,
s.n. (ISC). Boone
(MIN). Parke Co.,
(DUKE,
49446
Henderson
8 mi N of Bloom
Co.,
4 mi SE of Russellville,
Co.,
63030
Deam
of Howe,
Co., W
(IND). Newton
(DS, IND). Porter Co.,
1975, Zemelko
3 mi NW
Co.,
1938, Cottam
(RSA). Greene
s.n. (ISC, NY).
Co.,
Co.,
1925, Sfiker
1901, Shimek
Park,
kaloosa,
Spencer
1918, Shimeck
Co., De Witt,
Clinton
line, Deam
1945, Ward s.n. (FLAS).-IowA:
S of Brayton,
Putnam
(IND).
county
(IND, POM).
3 mi NE of Tunneltown,
Co.,
(CAS). Monroe
21016
2544
51506
Deam
Elkhart Co.,
51642
2.5 mi NW
Co.,
Lawrence
Tryon 2031
Baden,
1mi E of Dana, Deam
Co.,
(IND, POM). Wells
s.n. (ISC). Chickasaw
Deam
Deam
Co., 4 mi SW of Lafayette,
Tippecanoe
(MICH). Vermillion
Deam
57128
of Elkhart
17089
(IND, POM).
of West
1.5 mi S of Oatville,
Co., Deam
Pulaski
200
4 mi W
Co.,
(IND). Pike Co.,
Joseph Co.,
(IND). Lagrange
(IND, MIN).
Co., Blue Bluffs,
(MICH).
Deam
6 mi E of Huntington,
Co.,
Friesner
2 mi NW
Co.,
0.25 mi NE of Matthews,
s.n. (DS). Huntington
Co., E of Lake of theWoods,
Orange
12510
Floyd Co., 2 mi E of Byrnville,
29237
Crawford
POM).
2 mi N of St. Paul, Deam
Co.,
Deam
of Decker,
s.n. (OKL). Morgan
(IND, ND,
(IND, POM). Grant Co.,
50945
SW corner of Lake Cicotte,
Co.,
52584
3 mi ENE of Otis, Tryon 2652
(ND). Marshall
(IND, NY).
Deam
SMU).
1910, Morrison
7 mi W
Co.,
1914, Lipps
21392
Deam
Co., Noblesville,
(FSU). Knox
RSA,
Cass
UT, WVA).
IND). Decatur
(FLAS,
(GH, KANU,
(CAN). La Porte Co.,
Potzger
TRT, UC, UCLA,
0.25 mi N of Memphis,
6 mi NE of Griffin,
Co.,
14968
Co.,
63745
40428
(IND). Hamilton
61-618
TEX,
Clark Co.,
185
Co.,
Piscataquis
s.n. (NEBC).
Co., Farmington,
South Bristol,
Aroo
1894,
Wilson
Co., Penobscot
61
Valley,
SYSTEMATICBOTANYMONOGRAPHS
186
& Fernald
1897, Foxcroft
s.n. (NEBC).
1968, Bustamonte
ington Co., Wesley,
(SMU, TEX).-MARYLAND:
Munz
Baltimore,
Allard
3882
13472
Prince George's
(US). Washington
13376
den Co., Holyoke,
US, WTU).
(WIS).
Nantucket
Suffolk
13372
Munz
L'Anse,
207
of Wakefield,
17544
Munz
2, Turner 348
835
3588
Bazlun
Co.,
Ballard
lake, Clemants
913
& Grayson
Rock Co.,
& Moore
1393
505
11273
of Afton
of Roxie,
Shinners
Bridge
in Belzoni,
Shelton
92 (OKLA).
man 50312
(MIN, TRT).
Lake
of the Woods
7 mi SE of Cassville,
(UMO). Buchanan
Co., Zumbro
Harrison
4611
Magrath
(MIN, SMU, UWL).
13545
1871, Krause
(BH, DS,
s.n. (OKL). Greene
Franklin,
Co.,
67-1561
Cape Girardeau
Jasper Co., Oronogo,
Lincoln
Co., W
Morgan
Co., Bush
of Foley,
13115
Co.,
Palmer
Steyermark
807
(MO). Pike Co., Aberdeen,
14266
s.n. (MO). Howell
8118
Davis
1141
(MO). McDonald
Co., NW
Steyermark
Co.,
Jefferson
(MO). St. Clair Co., Tiffin, Bush
Brooks
1892, Bush
et al. 1091
13554
River
25 N of Luka, Cole
(KANU).
(UNCC).
(KANU).
Butler Creek,
Franklin Co., Gray
Palmer
(MO). St. Francois
Ripley
4221
(MIN, US).
Steyermark
Co.,
(IND,
s.n. (MO).
s.n. (KANU).
of Sedalia,
(MIN, UNCC).
13550
1910, Craig
1956, Steyermark
Carter
Jefferson
IND, POM). Howard
Springs, Munz
Springs,
216
9 mi E of Preston,
7539
(CU, DS,
Barry
& Drouet
s.n. (MO). Cole Co.,
(F). Pettis Co., 4 mi W
14381
Co., Yazoo
Co., Hwy
5 mi E of Willow
4777
1888,
733 (FSU). Lee Co., Tupelo,
Co.,
Co., Noel,
(MO). Ralls Co., Davis
Co., Winona,
(CAS, FSU). Camden
Sulphur
St. Joseph,
87 (MIN). Washing
Benson
Co.,
s.n. (NY).
Co.,
2 mi E of Columbia,
of New Madrid,
67022
Steams
(MICH).
16, Bartlett
1891, Taylor
14984
Munz
346
(MO). Franklin Co., 4.6 mi WNW
Co., Cape Girardeau,
of Springfield,
(MIN, MO).
(MO). New Madrid
cot Co., 3-4 mi. SE of Caruthersville,
8 mi W
1377
on US Hwy
& McGregor
IND, NY, POM). Clark Co.,
1934, Steyermark
POM).
nett, Steyermark
Boone
(OKL,
Frost
Spicer,
& Grayson
Falls, Gustitus
(DS). Tishomingo
Brooks
(IND, MICH).
Island near Fort St.
(POM). Humphreys
s.n. (MO). Dent Co., along Current River, Redfearn
1954, Hall
Co., N of New
(KANU).
Co.,
(MIN). Winona
4032
13585
1415
1961,
of Cross
Co., Bebb 5692
(MICH).
of Purvis, Barnes
10301
Sargent
3 mi S of St. Joseph, Henderson
Co.,
Munz
3 mi W
Co., 3 mi S of Langdon,
Atchison
Bartlett
near Zumbro
Rogers
s.n. (BH).
(MIN). Carver
Co., W
on Magnuson
1361
4380
&
557
& Grayson
Hubbard
Pope Co., Glenwood,
Hills,
Co., Biloxi,
LaMar Co.,
(UNCC).
Taylor & Taylor 12004
Summit,
River,
75
1910, Kauffman
Crow Wing
Co., Deer Creek
& Grayson
trail (Rte 21), Ownbey
Forrest Co., Ragland
Co., Van Buren, Munz
City,
s.n. (MIN).
16, Bartlett
Rd, Bunce
1880, Bailey
Kandiyohi
Co.,
Mower
NY, WS).
1977, Lightfoot
Hwy
UCLA).
3.8 mi E of Sherburn,
Co.,
s.n. (F,MIN,
Pearl River Co., Picayune,
(TENN).-MISSOURI:
Bartlett
(GH, MO,
Ham
Estates,
(MIN). Clay Co., Moarhead,
(MO, NY).
2032
along WS
(SMU).
65328
and Whalan,
of
1951, Bartlett
Co., Holland,
NY, POM, US).
2204
Lake,
SE of Rice, Dorio
1704
25 mi NE
Co., Pte. Mouillee
of Amble
South Haven,
Ballard
8, Swanson
Co., Rednalis
Rd & W
1927
on Rte.
S of Engadine,
Co.,
(PH). Monroe
Co., Walled
5 mi
Johnson
on Stagecoach
28782
Pullen
Co.,
Co.,
City, Dieterle
Mackinac
(MICH). Ottawa
MO,
Lancast
(MIN). Martin
Pine Co.,
of Luverne
(GH, MIN,
Pool
Sand Lake,
s.n. (MIN).-MISSISSwI:
Holzinger
3049
s.n. (MIN). Wabasha
1 mi N
640
Van Buren
(MICH).
Co., between
1892, Sheldon
(RSA).
1970, Westkaemper
ton Co.,
Co., Grant
Ingersand,
11.3 mi W
Nimke
s.n. (MIN). Benton
River, Navigation
Lacs Co., Milaca,
Oakland
(KYO).
(MIN). Cass Co., Lake Kilpatrick,
Lake Co., Ahlgren
Charles, Moore
992
Co., N of Briggs
5414
Co.,
Co., 20 mi SE
Gogebic
(POM). Kent Co.,
5250
5 mi S
Co.,
Iron Co., Brule River
(MICH). Mason
8400
Lake, Dreisbach
(MICH). Montcalm
Sandberg
(MIN). Filmore
Itasca Co.,
(MIN, WS).
Co.,
B646
Park, Umbach
Co., Chippewa
950
721 (GH, MICH).
286
Fitchburg,
(MICH). Baraga Co.,
10 mi SE of Traverse
Co.,
IND, POM,
(MICH, NY). Crawford
(MICH, WIS).
Twp., Hanes
Co., Farwell
Gillis
424
and
Dukes
Wiegand
1 mi N of West
Voss 4607
2656
1789
(POM). Hamp
(CU, DS,
117 (SD). Cheboygan
(BH, IND, NY, POM).
36 Kalamazoo
Co., 4 mi NE of Mio,
1981,
Co., Mississippi
SMU).
Mille
5397
(MIN). Clearwater
Houston
Sect.
Co., Henson
Schoolcraft
Co., Chaska,
Co.,
N Muskegon,
Co.,
s.n. (MICH). Oscoda
s.n. (MICH).
Erlanson
17539
Munz
Co., Portage
Area, McDonald
MINNESOTA: Aitkin
Co., SE Standish,
Co., St. Johns, Brown
of Petoskey,
of Lansing,
153 (POM). Mecosta
(MICH). Muskegon
s.c. 2528
Arenac
14681
Co., Wellesley,
Co.,
side of Platt River, Kempers
(MICH). Keweenaw
(MICH). Manistee
lin Lake, Silander
Co., N
Norfolk
Co., Baltars
s.n. (PH, TENN).
13373
Co.,
13476
Provincetown
Co.,
Co., Peabody, Munz
(MICH, RSA). Worchester
(BH, IND, POM, US). Grand Traverse
10 mi NW
Barnstable
(BH, NY, PENN).
Co.,
Munz
Co., Kensington,
Munz
s.n.
Baltimore
s.n. (PH). Charles
1937, Popowsky
Co., Lexington,
578
2780
2 mi W
Co.,
(OSH). Kalamazoo
Grand Rapids,
Richards
MacKeever
1907, Hubbard
(UNCC).
1922, Meredith
Essex
(DUKE).
PH). Wash
(NEBC,
s.n. (DS, MT, PH). Queen Anne's
Middlesex
(BH, IND, POM). Clinton
(US). Emmet
Ingham Co.,
State Game
4671
Seymour
Plain, Bartlett
17546
point,
8686
(MIN). Montgomery
Co., Mt. Washington,
(ISC, UNCC).
(MICH). Benzie
28598
Downs
(UNCC).-MASSACHUSETTs:
(CU, IND, NY, POM, US).-MICHIGAN:
Mell
Hyypio
87706
Jamaica
of Indian River, Munz
(MICH).
9813
Co., Siasconset,
Co.,
Bourdo
Grayling,
Vineyard,
Ahles
104
1904, Steele
Co., Hyattsville,
(CU, POM). Berkshire
Martha's
Co., Edgartown,
Co., Town
Park, Steele
Pearre, Downs
Co.,
North Truro, Munz
Cecil
(DS, IND, NY, POM).
14203
& Long
1.5 mi SW of Cumberland,
Co.,
Lake
Fernald
Co., Fairfield,
s.n. (MO). York Co., Brave Boat Harbor,
& Zegarra
Allegany
(US). Garret Co., Mt.
(IND, POM).
Somerset
VOLUME 50
Pemis
20345
5 mi E of Ben
Co., Bismarck,
Dewart
85
OENOTHERA
1997
City, St. Louis,
(MO). St. Louis
Louis Co., Clayton,
Birch Tree, Munz
RR overpass
280
Dorr
13547
1mi N of Cedar Gap, Munz
ter 118 (MO). Wright
Co.,
(DS). Sanders Co.,
NEBRASKA: Brown
Co., Long Pine,
S of Nemaha,
1mi
Co.,
Coos
(CU, DS, NY, POM).
1320
10238
Co., NE
Bergen
(MIN). Cumberland
Passaic
Co., Allwood,
Warren
Russell
selaerville,
17436
Munz
Britton
(CU). Nassau
Townsend
Co., Long
s.n. (CU). Oneida
Park, 1905, Bicknell
Lehr 496
land, Cold
s.n. (NY). Rensselaer
Co.,
7142
Ashe
883 (NCSC).
20476
24, Wilson
5.9 mi W
of Wade,
14448
(UNCC).
Haywood
Co.,
Co.,
14298
39416
Onslow
erty, Bell
Salisbury,
Radford
Sears & Ahles
N
Co.,
Parkway
327
on Hwy
(UNCC).
Horton
334
NW
of N Wilkesboro,
Co.,
S of Micaville,
Munz
12320
13519
MIN,
s.n. (DUKE).
of Rich
Square
17, Ahles
Nash
(DS,
Co.,
US, WIS).
River,
1mi NW
Walsh
Ahles
Hyde
3.7 mi
Co.,
(UNCC).
Cum
Radford
14757
of Clemmons,
Co., S of Friend
Scranton,
17082
Radford
in 1935 from
(UNCC). McDowell
50137
Person
of Sims, Radford
& Bell
45750
Co.,
Co.,
16738
(UNCC).
3 mi SW of
Randolph
Rowan
(UNCC).
17327
(MICH, UNCC).
Ahles
& Duke
(UNCC).
(DUKE).
1450
Raven
1.5 mi SW of Robersonville,
& Duke
(UNCC).
33419
73
Blair
& Leisner
Co.,
15246
Co.,
39766
& Leisner
IND, NY, POM, US).--NoRTH
UC, UNCC,
32510
Morgan
Ahles
Chatham
6.3 mi NE
Co.,
305, Ahles
Radford
Oosting
7 mi NE of Albemarle,
(NCSC). Wilson
Ahles
Keever
2 mi N of NC Hwy
37283
Co., Little Easonburg,
& Leisner
Co., Lake
Co., E of Asheville,
line, Bell
on NC Hwy
0.7 mi E of Pactolus,
Co., Marl
Crossroads,
Lee Co., cult. from Munz
(UNCC). Martin
of Huntersville,
Vance Co., 4 mi SW of Henderson,
Domes
(CAN, DAO, MICH,
on Roanoke
(UNCC).
IND, POM).
CAROLINA: Alamance
(NCSC, US). Guilford
(BH, IND, NY, POM).
Co., E of Reidsville,
Stanly Co.,
Forsyth
Is
York, Long
Co., E of Mocksville,
Rapids
1929
on US Hwy
Rockingham
(RSA).
(UNCC).
Wells
Pitt Co.,
(UNCC).
80, Bell 4469
s.n. (ARIZ).
s.n. (GH). Wayne
Bern, Radford
Davie
s.n.
1895,
Island, Forest
(CU, DS,
s.n. (UNCC).
1958, Bell
Co., 4.2 mi S of county
7 mi W
Co.,
Rd,
Co., Clarkstown,
Co., New
Carteret Co.,
2139
13521
Farm, Ridge
13388
Buncombe
Godfrey
1935, Correll
1.2 mi NNW
(UNCC).
2036
Taylor 1046
Co., N of Taylorsville,
(UNCC).
1914,
Island,
1891, Burnett
1mi SE of Leggets
Co., New
(UNCC).
of Roanoke
Lincoln
of Uwarrie,
Co.,
33554
Falls,
River, Grand
York, Long
Suffolk
of Rens
Co., N of Horseheads,
(FLAS).-NORTH
Co.,
14910
Craven
Co., Creedmoor,
(IND, NY, POM).
2 mi NW
Duke
14097
18315
& Leisner
(PH).
(GH). Rockland
Munz
s.n. (UNCC).
Radford
(UNCC).
of Clyde, Munz
4.9 mi SW of Deppe
Gordonsville,
1087
Halifax
1mi W
North Hampton
Co.,
Ahles
(UNCC). Mecklenburg
Co.,
24203
Alexander
Beaufort
2 mi SW of Catawba,
2440
of Durham,
1.3 mi S of Blue Ridge
(UNCC).
Co.,
7901
1932, Phillips
Co., New
1912, Burnham
(UNCC).
1958, Bell
Fogg
(CU, IND, NY, POM, US). Ulster
Falls,
(UNCC).
(NY).
Co., Windham,
Co., Naples,
Queens
House
13398
16925
2 mi SE of Araco,
Jackson Co., near Glenville,
(UNCC).
Montgomery
38454
(UNCC). Granville
(UNCC).
Radford
& Bell
Catawba
ship, Bell
39061
Ahles
Radford
1451
Co., Copake
Co., Townsend
Co., Bloomingburg,
and Etna, Munz
Co.,
Taylor
Co., Webster,
713 (CAN, CU, GH, US).
Co., N of Hudson
& Hammond
Co., NE
& Duke 48973
Sanford: Munz
S of Castleton,
of Wilmington,
Co.,
(UNCC).
Ramseur
Durham
(UNCC).
Co., W
Caldwell
3043
berland Co.,
Ahles
Jefferson,
Brunswick
of Bonsal,
NY).
s.n. (LAM). Yates Co., Potler Swamp, Ward 586
Co., W
(NY, RSA).
(GH). Ontario
(GH, MICH,
Phelps
Heights
62 N of Burlington,
Co., US Hwy
(DUKE).
Cayuga
(CU). Greene
(CU). Monroe
(NY). Niagara
112.35
(PH). Middlesex
s.n. (NY). Erie Co., Niagara
21933
14902
(OKLA). Hud
Co., 2 mi NW
s.n. (POM). Columbia
3448
6634
(NY). Sullivan
(POM). Washington
1883, Hankins
Bluff,
3097
(NY). Chemung
Swamp, Munz
7695
Co., Canton,
457
1915, Wilson
Gates
Co., Clinton,
2 mi SW of Harrisonville,
Park, Nash
Pennell
404
Co., Belmar,
(MO).
Moldenke
Co., Hammonton,
Hymowitz
Kral
789
Chandler
Co., Peermont,
YORK: Albany
1888, Fitch
Bicknell
13368
5 mi N of Enfield,
Co.,
s.n. (PH).-NEW
& Lindsey
Raup
between
Co.,
boro, Benson
Co., Bronx
Co.,
(NY). Sioux
28231
Island, Long 38267
Monmouth
(LAM, MO).
3250
Co., East Jaffrey, Munz
Co., Hampton,
Co., Clifton,
VPI, WVA).
(CU). Salem
Co., Cedar
Spring Harbor, Banker
Tompkins
737
Muenscher
Reservoir,
St. Lawrence
(NY).
PH). Essex
54194
Stephens
JERSEY: Atlantic
Co., Treasure
PENN,
Lake Co., Pol
of Thedford,
s.n. (FLAS). Cape May
1950, Schaeffer
Island, Woodmere,
Co., near Upper
Orange
(TEX). Rockingham
(PENN,
Co., Arkville,
above Wells,
Co.,
555
Co., Norwich,
s.n. (NY). Genessee
(NY). Hamilton
NW
Chenango
et al. s.n. (NY). Delaware
1875, Morong
(CU, GH, RM). Grafton
472
(MIN). Bronx
775413
(BH, OKL).
Moore
(NEBC, OKL).-NEW
& Clausen
Co., 0.25 mi N of Summerfield,
Demaree
Cheshire
of
Sut
(CAS, DS, GH, RSA, UC).
1mi W
Co.,
(MO). St.
8 mi W
Jacks Fork River,
City, Bates
s.n. (NEBC).
s.n. (CU, FLAS,
Marold
Thomas
s.n. (BH). Hunterdon
1920, Bailey
1891, Halsted
(KANU).
1888, Cuthbert
Adams
Co., Millville,
son Co., Hoboken,
Park,
2743
Co., River Vale,
460
1911, Farlow
Blake
Sharon,
county, Hodgdon
(BH, NY).
Menlo
Co.,
Co.,
(CU, IND, POM).-MONTANA:
Falls, Gregory
Otoe Co., Nebraska
4863
4521
758 (MO). Texas Co.,
(KANU).
16403
Stephens
& Mill
Shannon
Wagner
(IND, POM).
s.n. (MIN). Cass Co., Louisville,
58149
Co., Randolph,
(FSU). Hillsborough
Strafford
1898, Bates
HAMPSHIRE: Carroll Co.,
(KANU).-NEw
13552
9 mi E of Thompson
Stephens
Co., 4 mi E & 3 mi N of Harrison,
13543
Taney Co., Swan, Bush
(CU, IND, POM).
Nemaha
and Vandeventer,
Scott Co., Sikeston, Munz
(UNCC).
son, Thomas
14779E
at Tower Grove
187
Co., Lib
0.5 mi N of
Surry Co., near Elkin,
(UNCC). Wilkes
Co.,
1mi
(UNCC).
Yancey
DAKOTA: Cass Co., Kindred,
Stevens
Co., 9 mi W
37920
of Park River,
Stephens
28996
(DS,
KANU).-OHIO:
Co., Demaree
Champaign
(MIN). Erie Co., East Harbor
655
(LAM, MO).
Greene
Co.,
Portage
Co., Garrettsville,
(OKLA,
leyville,
13569
Munz
Bartlesville,
River
River Co., Columbia
0.4 mi W
1691
(MICH);
Sauvie
Polk Co., W
(PENN).
of Salem, Nelson
Allegheny
Berks
15673
(PENN).
13403
(IND, POM). Bucks
1950, Wohl
5477
Chester
(PAC, PENN).
mi WSW
s.n. (PAC). Erie Co.,
Jefferson
(PH).
Presque
2127
1.5 mi N of Sharon,
cer Co.,
3258
s.n. (NEBC).
1964, Richardson
ington Co., Westerly,
Colleton
(UNCC).
Providence
Ahles
Co.,
1.9 mi E of Smoaks
on Rte
of Providence,
River
& Tryon 1613
(UNCC).
Clarendon
& Leisner
of Cold
Munz
Co.,
(PENN). Mer
s.n.
Philadelphia
Co.,
s.n. (PENN).
Pot
(BH, IND, POM).
14799
(PENN).
(PAC). Wash
508a
14422
Spring, Wahl
ISLAND: Bristol
13382
Co., Slip
Lycoming
Stone
Aharrah
and Hwy
(PAC).
Co., Warren,
(BH, IND, POM). Wash
Co., Anderson,
170, Raven
Davis
18708
7742
(DAO, DS,
(CAS, GH, POM, UC, US). Cherokee
Co.,
217, Ahles
31925
13406
Shady Nook,
CAROLINA: Anderson
of the New
on SC Hwy
18, Ahles
Co.,
(US).-RHODE
8194
Godfrey
30931
& Haesloop
of Reevesville
5 mi W
Co.,
IND, POM).
Co., Oleopolis,
Mill,
3 mi SE of Cherryville,
Co.,
Ferry, 1938, DePue
2 mi NNE
Co.,
s.n. (GH).-SOUTH
Co., 4 mi W of Bonneau,
RSA, US). Berkeley
of Blacksburg,
(MO). Wayne
0.5 mi SW of crossing
Co.,
Venango
107
Donley
1890, Britton
Co., Launersville,
(BH, DS,
(PAC). Sullivan
s.n. (NCSC).
699
(KANU).
Snyder Co., Sunbury, Munz
(NY, RM). York Co., Rose
1912, Woodward
Beaufort
(MICH, ND).
6172
Westerfeld
13407
23
of Doyles
et al. 19965
Co., Fogg
1.5
Saxe
S of Damascus
Lawrence
NY, US).
36869
Schaeffer
1mi
Co.,
Station Dams,
0.75 mi ESE
(PAC). Northhampton
Munz
(PENN).
Fulton
Co.,
1959, Yacobucci
of St. Mary's,
Pump
Wahl
Crawford
of Shippensburg,
SSE
s.n. (F, NDG,
2 mi SW of Dingmans
Pike Co.,
et al. 7875
Mathias
7167
Co., Osterhout
8748
PAC, PENN,
(CU, FLAS,
3 mi
Brave
s.n. (PENN). Monroe
Adams
S of Sayre, Munz
PENN).
Juaniata Co.,
(PAC).
Berkheimer
(GH, NA,
s.n. (PENN).
Creek,
21567
Fogg
1mi S of Van Ormer,
Co.,
206
Sheldon
1.5 mi E of Williamsburg,
Co.,
POM). McKean
S of Liverpool,
1950, Larson
Lewisburg,
ington Co., E of Washington,
mi WSW
Wahl
1mi SE of Fairhope,
Co.,
Co., West
Wyoming
5 mi
10 mi
Co.,
Wahl
Co.,
1891, Small
(DS, IND, ORE,
(MICH, PENN).
0.5 mi N of Germania,
Somerset
Union
1689
3651
1939, Whittenberger
(PAC). Perry Co.,
Olney, Dreisbach
ter Co.,
13404
Bradford
1mi N of Mosserville,
Co.,
Co., SE of Ft. Washington,
(NY). Montgomery
Schaeffer
Wahl
Co., Conewago,
(WVA). Lehigh
5 mi NE of Picture Rocks, Munz
& Jennings
along Dunkard
Co.,
of Rathmel,
Lancaster
(PENN).
pery Rock, Russell
Cumberland
(MO);
Suksdorf
NY, POM, US, WS).
0.4 mi E of Cessna,
134 (PAC). Elk Co., 4 mi W
Cady
Jennings
Isle, 1935,
(PAC). Greene
0.5 mi ESE
Co.,
et al. 7084
Wherry
1mi E of Harrisburg,
11755
Westerfeld
(PENN).
4537
s.n. (WS); Portland,
(GH, MO,
Co.,
15953
Fosberg
of Scio, Peck
& Halley
1.5 mi NE of New Oxford,
Co.,
Bedford
(NY). Clarion
Co., 0.25 mi N of Central,
SE of
Co.,
(IND, POM). Hood
(MO); near Portland,
S.11009
and Milesburg,
6383
Wallis
of Hai
(DAO, DS, GH, NY, OSC, RSA, UNCC,
(GH, PENN).
Bellefonte
s.n.
14431
1925, True s.n. (MICH). Cambria
Edmondson
1949, Wherry
Co.,
Adams
(US).
(UNCC).
2 mi W
Co.,
(OKLA). Washington
1885, Suksdorf
(UC, MT).
3394
Eddy,
between
Co.,
Chester,
Columbia
of Teepleville,
(PENN). Dauphin
Church,
Centre
779
Berkheimer
Co., upper Black
Co., West
2619
(MO); St. Johns, Sheldon
Sumstine
Pittsburg
4538
& Halley
& Dorris
s.n.
(MO). Linn Co., 6 mi W
1843
Bonneville,
(PH).-PENNSYLVANIA:
Co., Reading,
s.n. (PENN).
TENN, WS).
4090
618
Jones
s.n. (PAC).-OKLAHOMA:
30 in E Portland, Wagner
street on Hwy
Rd, Dennis
5499
Co., Wilkinsburg,
Clark
River, Hoch
of Hood
156 (UNLV);
Epling
(OKLA).
61345
16999
6.7 mi NE of Talequah,
Co.,
Co., 6 mi N of Bend, Munz
in E Portland, Wagner
Island, along Reeder
(ORE); near Portland,
S.10103
Point, W
E side of Mt. Tabor, Leary
WS, WTU);
S of Jenks,
Rose
1966,
1911, Overholls
2641
Means
Deshutes
just E of 205th
Co.,
street on Sand Blvd.
of 238th
1.5 mi
Co.,
at Dalton
(WILLU). Multnomah
24366
Tulsa
S of Hughes,
39 (OKLA).-OREGON:
2 mi E of Norwalk,
Elkton,
City, Demaree
Co., Cincinnati,
(OKL). Cherokee
Greenman
Co., Cleveland,
Co., Grove
(US). Hamilton
Co.,
Taylor 978
2.6 mi
Co.,
(POM).
McDonald
s.n.
Cuyahoga
Franklin
(MIN).
(GH). Preble Co., West
Webb 244
LeFlore
94
(MO). Huron
s.n.
1914, Drushel
SMU).
(MO, SMU, UC, US).
1886, Wooton
1.5 mi S & 4 mi E of Bennington,
Bryan Co.,
5980
Friburg,
11701
State Park, Albright
Jamestown,
Co.,
Holmes
NW
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
188
3 mi ENE
& Bell
(KANU,
of Turberville,
17851
UNCC).
Radford
(UNCC). Dorchester
Edgefield
Co.,
28230
Co.,
1.6
Co., N of Augusta,
Bartlett 2384 (MICH).Georgetown Co., Maryville, Radford 28547 (UNCC).Greenville Co., nearGreenville,
& Mullens
Rodgers
67132
Bell 5296
sawhatchie,
berry Co., NW
of Brown's
Leisner
(UNCC).
31627
din, Radford
Custer
Mt.
28356
Tazewell,
Bartlett
s.n. (TENN).
Harmon
Crossroads,
Pickens
Co., W
Anderson
15 (TENN).
Bell 9793
(UNCC).
of Pickens,
1326
Elizabethton,
Coffee
Co.,
239
(UNCC).
Williamson
Orangeburg
Radford
(MICH).
Co., near Norris,
Co.,
Duke
8 mi NE of Kershaw,
s.n. (ISC). Lawrence
& Grayson
Carter
Co.,
16657
DAKOTA: Codington
(UNCC).-SOUTH
(KANU).-TENNESSEE:
Thomas
Lancaster
1909, Whitham
Co., Custer,
Rushmore,
Horry Co., Nixonville,
(UNCC).
(UNCC).
Co.,
8 mi N of Pelham,
Bennett
s.n. (TENN).
Godfrey
(UNCC).
Co., 0.4 mi N of US Hwy
3.5 mi W
& Manning
1.1 mi SE of Coo
2191
1312
of Northville,
Blount
2210
69784
Dugle
(DAO).
310
(FSU, UC).
(DAO).
Stephens
Claiborne
Fayette
&
of Gour
Pennington
Co., Chilhowee
(CU, GH).
New
301, Ahles
Co., 4 mi W
shore of Lake Kampeska,
Co.,
1940, Cole
Wiegand
& Ahles
(SMU). Williamsburg
Co., Spearfish,
Spink
Jasper Co.,
Mtn,
Co.,
61318
1964,
Co., New
Co.,
near
OENOTHERA
1997
Loosahatchie,
Grundy
et al. 13325
Shanks
State Forest,
Knox
Co.,
5 mi NW
Lawrenceburg,
of Knoxville,
Kral
kee National
32943
1102
Munz
13527
Co.,
52979
(TENN). Montgomery
46340
Co., Embreeville,
(DS, SMU).-TEXAS:
Thorp 2798
Pittsburg,
& Watson
s.n. (SMU). Morris
Douglass,
Shinners
24888
Amerson
Pleasant,
Manchester,
(NEBC).
262
Day
Orange
ington Co.,
POM, US).
UNCC).
Co., Fairlee,
Amelia
Rodman
Co., Lewis
(UNCC).
Co.,
13478
(BH, IND, NY, POM, US).
Chesterfield
Co.,
James City Co.,
& James 61967
from Munz
13360: Munz
Island,
& Waggoner
Wytheville,
5 mi W
TON: Clark Co.,
near Cape Horn,
696
& Grayson
Longview,
ID, MICH,
3632
635
Skamania
Co.,
of Monroe,
lor 3544
(DS, MICH,
Suksdorf
& Grayson
625
(WWB); Lummi
(US). Yakima
Falling Waters,
Shore Drive
Co., Naches,
Downs
3673
Jones 8614
(UNCC).
616
MIN,
(CU, DAO);
Lummi
& Grayson
(CAS, OSC,
Sauk River
Co.,
and Goldbar,
MIN,
Bartlett
POM, WTU).-WEST
Bartlett
698
(MO);
(BH, GH, CAN,
and M
Wagner
8292
3632
Bay, 2.4 mi NW
COLO,
of Toke
Street S in Takoma,
Sundquist
along RR
4543
5.3 mi W
& Grayson
Bartlett
s.n. (WTU). Kit
near Rockport,
NY);
626
509
tracks,
(MO); 0.4 mi W
of Sultan,
2.5 mi E
(DS, RSA). What
(CU); Bellingham,
C. Begert
Indian Reservation,
Tay
further locality, Benson
2362
VIRGINIA: Berkeley
S of Gassaway,
& Grayson
(DS, IND, MICH,
E side of Sultan
s.n. (WWB); without
Co., Sugar Creek,
1893,
Anderson
6269
Suksdorf
Island, Muenscher
1966, King
Vancouver,
T C. Frye
River,
290 (WWB); Red River Rd, near Lummi
at Bel Bay,
Braxton
NY, RSA),
701
14706
830, SE of Longview,
Pacific Co., Willapa
Sultan
& Grayson
14481: Munz
IND, POM);
from near Westport,
(DS, MICH,
Co.,
14, 7 mi E of 32nd St. inWashougal,
of Duwamish
Snohomish
Southamp
(UNCC). Wythe
(IND, NY, POM).-WASHING
Jct. of S Takoma Way
2 between
& Grayson
(DS, MICH,
8291
(WTU).
757
cult. from Munz
Bartlett
Skagit Co.,
Co.,
SE of Harrisonburg,
(BH, IND, NY, POM).
Appler
1968,
Pulaski
(NY, US).
5 mi
Co.,
(MO). Northamp
Co., Co. Rd 924,
1mi N of Woodland,
Co., Bingen,
Pierce Co.,
11516
SE part of Sand Point, Sutherland
cult.
Co.,
near mouth
NY, RSA).
B2625
3538
(BH, DS,
99,
US, WS, WTU).
NY).
MIN,
cult.
Co.,
(MO); along US Hwy
(DS, IND, MICH),
(WS). Klickitat
NY, RM, UBC,
Prindle,
4540
Harbor
(POM). Gray's
(DS, MICH,
com Co., Birch Bay, Muenscher
52 (WWB);
697
& Grayson
1.5 mi E of Startup, Bartlett
Bartlett
15248
along Hwy
along US
Co.,
(MO); 0.7 mi E of Startup on Hwy
4542
of Goldbar,
(OSC). Cowlitz
(NY, POM),
7.9 mi N of Rock Creek, Bartlett
(WTU);
just off 1-5, Wagner
Inlet, Otis 2164
686
& Grayson
(WWB).
9301
835
Thompson
& James
SW of Lunen
Co.,
(GH). Nelson
14269
(BH, IND, NY, POM, RSA),
(POM). King Co., Seattle,
MIN, MO,
& Grayson
land, Bartlett
Wagner
Abrams
head of Sinclair
sap Co.,
DS, GH,
Bartlett
Woodland
(MICH), Bartlett
Anderson
Westport,
13511: Munz
(BH, IND, NY, POM), Munz
4699
Chambers
700 (DS, IND, MICH);
NY);
14773
s.n. (WS); near Vancouver,
News,
501, Ahles
7551
Allard
13507
Munz
Munz
1.5 mi NW
Co.,
13360
(US). Pittsylvania
Co., Newport
(POM). Din
of Warrenton,
Munz
Hill, Hubricht
2647
903
& James
& Long
s.n. (US). Rockingham
cult. from Munz
14481
Lunenburg
of Bowers
99, 3.6 mi S of Johnson Creek,
along US Hwy
on US Hwy
1mi N of Anbirch,
Co.,
1904, Steele
(IND, POM, US),
IND, POM), Munz
Suksdorf
0.5 mi W
(US). Warwick
3 mi W of Camas, Munz
(MICH, RSA);
(BH,
13511
s.n.
River
Fernald
Ahles
(VPI). Giles
(GH, MICH).
S of Suffolk,
Co.,
Smyth Co., 4 mi SW of Marion,
Steele
1916,
174
s.n. (US). Page Co., Walker
Co.,
2070a
& Condit
(BH,
Co., Custip
14296
Munz
3 mi W
Co.,
10 mi SW of LaCrosse,
Co.,
Prince William
Fauquier
(MIN,
17593
Fosberg
Co., Herman,
40 mi N of Richmond,
9896
(BH, IND, NY,
(VPI). Bedford
Charlotte
3 mi S of Hyco
Co.,
Co.,
12431
28425
s.n. (GH). Wash
13488
Munz
Co.,
Pease
& Moldenke
SE of Arlington,
(UNCC).
Porter
of Toano, Menzel
(VPI). Norfolk
1897, Palmer
mi
Freer
(IND, NY, POM).
(POM). Nansemond
(DS, IND, POM).
Munz
13362
Munz
1909, F S. H. s.n. (VPI). Rockbridge
13483
7544
Floyd Co., Buffalo Mtn,
s.n. (UNCC).
ton Co., Riverdale,
James
(UNCC). Mecklenburg
14294
et al. 9310
Tye River, Ramsey
Munz
0.3-0.5
Preserve,
(BH, IND, POM). Halifax
13495
Munz
burg, Ahles
Ruska
Co.,
cult. from seed col. by Munz
10 mi SW of Petersburg,
ton Co., Smith
(VPI). Arlington
Game
s.n. (GH). Bennington
Moldenke
5 mi NE of Covington,
4.7 mi E of
6.5 mi E of Mt.
1896, Williams
Co., Willoughby,
Co.,
of Big Levels
1970, Amerson
Co.,
(TEX). Essex Co., Concord,
Co., Townshend,
s.n.
De
s.n. (POM). Camp Co.,
(NY). Titus Co.,
Alleghany
2.1 mi N of Amsterdam,
Co.,
60845
(UNCC).
(CU). Orleans
1934, Kelley
Co., Silsbee,
1908, Williams
21527
Area, Blum 2978
(POM). Nacogdoches
1927
(UNCC).
Co., Gladeville,
1937, Cory
Ruth 997
Co., Ripton,
Junction, Blake
(PENN). Windham
1006
entrance
widdie
60481
1913
True 145b
Co., NW
Botetourt
of Narrows,
Co., Essex
POM, WTU).-VIRGINIA:
Augusta
(KANU).
Addison
(SMU).-VERMONT:
S of Waterburg,
MO, NA, OKLA,
Tarrant Co., Lake Worth,
(SMU, TEX).
73 (US). Chittenden
Cory 25740
1816
Lake,
Wilson
Co., Chero
Demaree
Recreational
s.n. (ISC). Hardin
8.3 mi N of
Co.,
Chester
Co., Norris
of Montalba,
s.n. (TENN).
(DS). Monroe
Co., Memphis,
(SMU, UNCC).
s.d., Rolfs
8 mi S of Daingerfield,
Co.,
Shelby
Dam, Rockland
4576
4 mi NW
Co.,
Co., Navasota,
Lawrence
20490
(SMU). Union
& Mahler
Anderson
(TEX). Grimes
Raven
Co.,
(SMU). Grundy
1955, Wolfe
6 mi SE of Clarksville,
Co.,
27668
Shinners
Mahler
8967
Valley,
IND, NY, POM, US).
Sumner Co., Old Hickory
NY, POM, SMU, WS).
Nichols
Standley
s.n. (SMU, TENN).
1964, Thomas
(FSU). Tipton Co., 9.7 mi SW of Covington,
maree
(BH, DS,
Co.,
7 mi SE of Loudon,
Loudon
Forest, Malter
(TENN). Washington
3 mi S of Greeneville,
Hawkins
(UNCC).
(SMU).
Sevier Co., N slope of Sugarloaf Mtn,
(GH, ISC, MO,
Co.,
(RSA). Greene
Clark & Stevens
189
1953, Boggs
Co., 2 mi SE of
s.n. (WVA). Ca
SYSTEMATICBOTANYMONOGRAPHS
190
bell Co., near Milton,
1933, Harris
Co.,
(WVA). Fayette
Co., Nuttallburg,
Co., Tomlinson's
Co.,
Run Park, West
Co., Fairmont,
Springs,
3 mi SW of Pisgah,
Co., Bluestone
WO-194
(WVA).-WISCONSIN:
land, Munz
17543
Iltis & Noamesi
Mondovi,
Co., New
MIN,
Rhodes
384
Thomson
Co., Argonne,
s.n. (WIS).
9 (WIS). Jefferson Co., Fort Atkinson,
Co., Christensen
Seymour
1471
(BH, IND, POM, US).
7657
Co.,
Co., Hull Twp.,
Portage
River,
17 (WIS). Sawyer
Shawano,
s.n. (MIN, WIS).
(WIS). Laglade
378
17521
of Hwy
SPECIMENS EXAMINED FROM OUTSIDE NATURAL AREA
by K. Rostaiiski).
and submitted
AIRES: La Paternal, Morfino
menia.
GORIS (Sgorits):
(PH); Villach,
Humpata,
(AMD).-MENDOZA:
s.n. (GZU);
1971, Holzner
63
Co., near
(WIS). Trempealeau
Poltzger
Co., 0.5 mi W
8702
(ND, WIS).
of West
Bend,
Theis
36 (WIS). Waushara
Co.,
1956,
by an asterisk were
followed
& Moreno
9987
BURGENLAND: Andau,
1896,
1926, Arbesser
Cavas
1968, Hubl
1970, Melzer
s.n. (GZU);
BUENOS
18525
(FI). Ar
s.n. (W); Ober
s.c. s.n. (US); Schwertberg,
Zeile,
examined
(K). Argentina.
Barrio Gov. Cano,
s.n. (W); Ossiacher
near Villach,
Seebach
17519
70, Iltis et al.
(WIS). Shawano
Twp., White
(citations
Barbosa
Hoecken,
at Hwy
45 (WIS).
Las Heras,
s.n. (WU). Austria.
Petrop.
s.n. (GZU).-KARNTEN:
s.n. (W),
Seipka
1929, Eggler
1476
Hort.
1935, Miischl
schiitzen,
Huila:
Angola.
Co., Three Lakes,
St. Croix Co., N of Glenwood
(WIS).
Iltis et al. 7405
83, Derrwaldt
1mi SW of Fisk, Hein
Co.,
Winnebago
1mi W
Co.,
s.n. (WIS). Milwaukee
1964, Goff
(BH, IND, POM). Washington
59 &
Duffie
s.n. (WIS). Lin
1965, Gates
near Trout Lake,
Co.,
of Mercer,
(WIS). Kewaunee
s.n. (CU). Oneida
Co., Westboro
(UWL). Vilas
119 (WIS).
(WIS). Forest
25 mi E of St. Croix Falls, Munz
40, Iltis et al. 7840
(DS,
Irving s.n. (POM). Marinette
(WIS). Price Co., Pike Lake
Lake Dam,
1440
s.n. (WIS). Green
Iltis 15833
Co.,
1929,
1934, Honey
228
s.n. (WIS). Taylor
1mi S of Hwy
Co.,
Juneau Co., Point Bluff,
191 (WIS). Polk Co.,
on Hwy
Demaske
Mtn,
85-I
1963, Ackerman
1.5 mi N of Moose
Co., N end of Spooner, Munz
10 (OSH). Waukesha
Hagene
Creek
Co.,
1958, Melchert
S end of Trempealeau
Washburn
Lake, Bell & Colvin
Co., Wausau,
2 mi E of
Co., Reeseville,
s.n. (WTU). Dunn Co., Duffie
Iron Co., 2 mi NW
Tessene
Co., Ash
Co.,
& Grayson
(WIS). Dodge
bottoms,
Co., Keshena,
Cutler
Bartlett
161
6 (WIS).
1674
s.n.
& Wilson
1957, s.c. s.n. (WIS). Chippewa
River
(WIS).
11426
Allard
s.n. (WIS). Ashland
of Portage,
Keller
Co.,
Summers
1957, Hinkle
Co.,
Iowa Co., Beal
Co., Hartley
(WIS). Oconto
(WVA).
near Davis,
Co.,
Beach,
1956, Bennett
(SMU). Marathon
Co., Cedarburg,
(WIS). Rusk Co., N of Weiger
City, Radcliffe
18069
RO-202
s.n. (WIS). Buffalo
Jack Pine Stand near Sand Lake Lookout,
Co.,
Shinners
Smith 42 (WIS). Ozaukee
Wolf
15501
90 (WIS). Menominee
Co., Milwaukee,
Nee
(WIS). La Crosse
3947
coln Co., Tomahawk,
& Wilson
(WIS). Fond Du Lac Co., Mauthe
1960, Ugent
s.n. (WVA). Randolph
1970, Jarrell
Waverly
Sterling,
77 (ISC). Grant Co., Mississippi
Lake Co., near Fox River,
6 mi E of Berkeley
(WVA). Preston Co.,
1952, Byle
Co., 4.4 mi W
of Mt.
(WIS). Door Co., 2.5 mi ESE of Gills Rock,
Eau Claire Co., Fall Creek, Kunz 264
Martz
1 mi W
Co.,
Co.,
2307
1948, Brown
Twp.,
(BH, NY, POM).
3 mi SW of Slate, Bartholomew
Co.,
Co., Appleton,
s.n. (WIS). Columbia
Crawford
TEX, WIS).
Co., Suamico
(WVA). Mercer
14226
s.n. (WVA). Webster
2 mi N of Friendship,
Co.,
Brown
(WIS). Calumet
1950, Helm
Auburn,
MICH,
Adams
8115
611 (WVA). Wood
(IND).
Clarkson
s.n. (WVA). Tucker
1972, Wilcox
579
s.n. (WVA).
(WVA). Morgan
Bartholomew
1967, Phillips
1063
252
Hancock
(UNCC).
McCauley
13501: Munz
Springs,
Co., Rock Creek,
Co., Liverpool,
Haught
(BH, IND, POM).
MO-6
& Wilson
1961, Music
Co., Anawalt,
cult. from Munz
Co., Dunmore
3 mi NE of Middlebourne,
Co., E of Littleton,
(WVA). Wetzel
4959
Co., Point Pleasant, Millspaugh
& Shoulders
River Valley,
Downs
Co., Romney,
Calhoun
US, WIS).
Bartholomew
Co., Park, suburb of Clarksburg,
(BH, IND, POM),
s.n. (WVA). Raleigh
New
Tyler Co.,
PH, SMU, TENN,
Post Office,
(WVA). McDowell
s.n. (WVA). Mason
s.n. (WVA). Roane
Reservoir,
(GH, US, WVA).
J-50
s.n. (PH). Pocahontas
1937, Myers
1937, Perine
(WVA). Hampshire
Bartholomew
Co., Morgantown,
1939, Strausbaugh
Pickens,
623
NY, OKL,
S of Blandville
729 (WVA). Harrison
13501
Munz
MIN, MO,
7 mi
Co.,
Bartholomew
1953, Steele
3 mi S of Princeton,
Monongalia
(BH, DUKE,
Nuttal
2 mi W of Garfield,
Jackson Co.,
Marion
653
Williams
s.n. (WVA). Doddridge
VOLUME 50
1876, Kelk
s.n. (GZU);
St. Magdalena
s.n.
Feistritz,
near Villach,
1929,
Arbesser s.n. (GZU).-NIEDEROSTERREICH:
Altenburg, 1889,A. s.n. (LY);Boheimkirchen, 1975,Kaisergruber
s.n. (WU); Florisdorf
near Wien,
Kienberg
near Gaming,
Arbesser
s.n. (GZU);
between
s.n. (W); Pressbaum,
s.n.
Zerny
Lungau,
Graz,
Ramingstein,
1898, Janchen
near Bruck,
Tragoss
Schwarz
Sulm
s.n.
(W); Linz,
1884,
1850, Rauscher
Steininger
1900, Vierhapper
s.n. (WU).-STEIERMARK:
Gstatterboden,
Preissmann
(W); Bruck
s.n. (GZU); Frojach NE of Murau,
valley,
Gleinstatten,
Strohmeyer
(PR); Pfennigberg
s.n. (Fl, PR); Schweitberg,
s.n. (Z); Gesause,
s.n.
s.n.
1942,
249
750 m,
1932, Wyatt
at river Mur,
1930, Buch
(GZU); Knittelfeld
1917,
(B); Kramesau,
1969,
s.n.
near Linz,
1874, Keck
Gleichenberg,
1885,
Topitz
s.n.
s.n. (W);
1913, Sabransky
s.n. (GZU);
s.n. (GZU); Gaisfeld,
(BC);
s.n. (UC).-SALZBURG:
1875, Preissmann
s.n. (LY); Sochau,
at river Mur,
1899,
1911, Kobb
s.n. (WU); Zeiselmauer,
Baschant
1877, Fritsch
s.n. (GZU);
near Wien,
s.n. (W); Kalksburg,
1862, Sonklar
Neustadt,
1898, Kerner
s.n. (GZU); Stadlau
1969, Polatschek
near Linz,
Klein-Munchen
near Mautern,
1896, Kerner
and Reichenau,
s.n. (ND); Wiener
1878, Vederetz
at river Enns,
s.n. (HBG); Baumgarten
s.n. (GZU); Mautern,
Hirschwang
(W).-OBEROSTERREICH:
Wurm-Zochbauer
Reichraming
1903, Krebs
1878, Przyb.
Feldkirchen,
1900, Fritsch
1969, Mayerwieser
s.n. (W);
1901,
s.n. (GZU);
s.n.
(GZU);
OENOTHERA
1997
Leoben,
1917, Widder
Lorenzen
Troyer
at river Preg,
Insterberg,
1968, Polatschek
Freudenau,
s.n. (W).-WIEN:
Litovsk,
Skvortsov
& Chalovsky
(BR); Brasschaat,
8078
Lawalree
(BR).-BRABANT:
s.n. (CAS); Bruxelles-Forest,
Etterbeek,
1923, Michiels
s.n. (BR),
1933, Vandevelde
1961, Feller
(BR).-LUXEMBOURG:
Anseremme,
s.n. (BR); Gent
Sloover
GABROVO: Gabrovo,
DIV: Plovdiv,
1898, Sorkpil
river Yantra,
Mrkvicka
s.n.
752
1898,
(GH, V);
Lake, Dawson
Shuswap
(CAN, GH, NY); W
Raven
S of Yale
1, 5.8 mi
Lessauer
(55?17'N,
27899
daihe, Hou
10411
out further
locality,
& Morse
(PE); Guiren,
Cui & Zhu 226
7317
MORAVSKY: Brno
(Brunn),
Tonghua,
between
Bor
Nanking
Nachod,
between
Ceske
(Plau)
and Tachov
Fangshan,
s.c. 3381
1926, Chiao
(PE).-LIAONING:
(Tachau),
1910,
and Loubi,
Skalice,
s.n.
Lake,
(MO); Hwy
1889,
s.c. s.n. (PE); Bei
with
s.n. (UC).-JILIN:
Andong,
Wang
Kunming,
s.n. (FI); Zagreb,
Treboii,
Urban
248
16 (54?8'N,
4545
(PE).-HuNAN:
(BM).-YUNNAN:
and Mala
C249
1917, Newcombe
in a garden,
1915,
Jehli'k 6826
1032
1955,
1913, Rossi
(PR).-JIHO
Jehlik & Rostanuski 6645
s.n. (SOM).-STAEDCESKKY:
Skalice
s.n.
(SOM).
Macoun
on BC Rte
(Nanjing),
Dedin
s.n.
Lake, Carter
1, Wagner
Beijing,
1902, Malegari
1911, Petrak
Kuzmanov
s.n. (MO); N of Appledale
of Endako
and Hradec,
11112 (NY).-SEVERCESK?Y:
(Weigkirchen),
1905,
1971,
s.n. (MO); Shawnigan
on Hwy
Chu 3839
(Parenzo),
Jindfichiv
valley
VALPARAiSO: Quilpu6,
et al. 479
Zhongde
Pao-hsing-hsing,
(MO).
9 (SOM, WU);
s.n. (V); Victoria,
(MO); 16 mi NW
55 (PE).-HUBEI:
Istra (Istria), Porec
(PR).-VYCHOI)CESK'Y:
(PR).-ZAPADCESKY:
Guan
JIHCESKS': between
Hranice
6, 1981, Marchant
(MO). Chile.
43526
66 (WU).-PLov
Balabanova
1981, Marchant
exit of Chilliwack
4546
(BR); St. Ides
(LY); Samokov,
(NY); Harrison
1914, Newcombe
2530
(BR).
s.n. (L). Bhutan.
(V); New Westminister,
2-3-13
Bridge,
s.n.
1927,
85894
2824
Hatschbach
Sofiya,
840 m, s.c. 900 (HGAS).-HEBEI:
(PE).-SICHUAN:
1874, Schur
Macoun
Robbrecht
1633
1989, Urumoff
(BR);
1883, Mag
Robbrecht
Urumoff
Stfibrny
(LY).-SOFIYA:
Long
27898
s.n. (US).-JIANGSU:
(POM, US);
Republic.
(PR).-SEVEROMORAVSK'Y:
Jehli'k 6241
Xinyang,
s.n. (KUN). Croatia.
(ZA). Czech
Raven
Sevlievo,
1899,
s.n. (BR);
s.n.
Bellem,
1952, Herhelst
YAMBOL: Elkhovo,
and Yale,
of western
tracks, Wagner
Steward
Jilin, Dorsett
Liou Sheu-o
Hope
GuIZHOU: Zunyi,
1925,
(SOM).
E side of Hwy
House,
127?38'W),
(PE).-HENAN:
s.n.
1 km N of Winlaw
(MO); 3.8 mi W
along RR
s.n. (M). China.
Charrel
(NY); Agassiz,
s.n. (V); C.P.R. between
s.n. (V); Old Hazelton
125?22'W),
Comfort
Kostenec,
Island, Koksilah,
55017
side of Slocan River,
turnoff, outside Valley
1912, Newcombe
s.n.*
s.n.
1886, Pietquin
Scheidweiler
s.n.
(BR).-NAMUR:
and Zelzate,
PARANA: Palmeira,
s.n. (SOM);
at river Yantra,
Tirnovo,
Stribmy
Gent
and Brugge,
(E, GH). Brazil.
1888,
1960, Vannerom
1933, Masseray
(BR); Zelzate,
s.n. (UC); St. Denijs,
1907, Davidoff
Sliven,
BRITISH COLUMBIA: Vancouver
Canada.
1055
1863, Thie
1908, Mathieu
12499
and Taillis,
s.n. (BR);
1920, Vermoessen
(Tirlemont),
Lawalree
(BR); between
Robbrecht
(SOM).-SAMOKOv:
(SOM).-SLIVEN:
Varna,
2929
St. Michiels
& Long 2702
at river Staz,
s.n. (PRC).-RUSE:
Jurkoeskij
(SOM).-VARNA:
Bridge
(BR); between
(Louvain),
s.n. (BM).-OOST-VLAANDEREN:
& Jongepier
s.n. (BR); Brux
1951, Rogier
s.n. (BR); Eprave,
Houx
between
1929, Dewildeman
2581
2400 m, Grierson
Taba, Thimphu,
(BR);
s.n.
near Postel,
1868, s.c. s.n. (BR);
(BR); Stokrooie,
Lahage,
1889, Cluysenaar
s.n.
s.n. (BR); Oudegem,
Duinbergen,
bald near Koksijde,
(BR);
1882, Hennen
s.n. (BR); Val-Benoit,
8010
Lawalree
s.n.
1914, Boulanger
(Gand), Robbrecht
1863, Thielens
WEST-VLAANDEREN:
of
1938, Gras
s.n. (BR); Wavreille,
(BR); Langerloo,
Genk,
Legrain
s.n. (BR); Belvaux,
near Eprave,
Crein
Bulgaria.
1946,
s.n.
1920, Vermoessen
(BR); Tienen
720
1892, Sladden
of
1957, Leothard
(BR); Moeren
s.n. (BR); Deurne,
Lawalree
Liege,
s.n. (Z).-LIMBURG:
Poncelle,
1867, Guismot
de Han
Rochefort,
nel
s.n. (BR); Pare near Leuven,
123
s.n. (BR); Leuven
1861, Pire
1930, Vits s.n. (BR, UC);
lens s.n. (FI); Vilvoorde,
(Louvain),
1866, Coomans
s.n. (BR); Groenendael,
Peeters
1974,
at river Dnepr,
(WA); Turov W
s.n.*
s.n. (BR); Auderghem,
and Leuven
Blandput
Pushcha,"
(LE); Mogilev
s.n. (BR); Kalmthout,
and Postel,
1937 & 1939, Michiels
Aarschot,
s.n. (BR); between
1956, Lawalree
1920, Lamberts
Mol
s.n. (BR); between
Pire
s.n. (LD);
s.n. (W). Belarus.
"Belovezhskaia
1921, Ptaszcki
and
Schonwies
1980, Merbeck
ANTWERPEN: Antwerpen,
(LE). Belgium.
1978,
s.n. (W); Worgl,
1915, Korb
s.n.*
1923, Savicz
near Lachwa,
marshland
s.n.*
s.n. (W); between
natural
reservatio
Brest,
1973, Po
s.n. (W); Lienz,
1970, Polatschek
s.n. (W); Praterspitz,
near Gomel',
s.n. (BR); Herentals,
Spas
1874,
1894, Michel
s.n. (BR); Vise,
(M). Prov.
1893, Pachoski
1896,
(Lierre),
Blandput,
Moule
s.n.
at river Dnepr
near Lienz,
1975, Polatschek
s.n. (W); Vill,
1921,
s.n. (GZU).
s.n. (W); Jenbach,
s.c. s.n. (WU); Donau-Auen,
1810,
s.n.* (LE); Poles'ye
at river Pripyat,
(BR); Lier
elles,
Skvortsov
1974,
s.n. (MO); Rogachev
1852, Pabo
Mozyr'
Augarten,
(GZU); Stainz,
s.n. (W); Amlach
1973, Polatschek
s.n. (LD); Prater, 1857, Mllner
1899, Rechinger
4595
1980, Polatschek
s.n. (W); Landeck,
1978, Polatschek
s.n. (GZU); St.
1986, Melzer
Valley,
s.n.
Schwarz
1902,
1927, Strohmeyer
Salzmann
upper Mur
1976, Polatschek
and Norsach,
Nikolsdorf
s.n. (W); Strass,
1972, Polatschek
ob Ebiswald,
s.n. (W); Innsbruck,
1980, Polatschek
Kranebitten,
s.n. (W); between
1976, Polatschek
Brest
1978, Polatschek
s.n. (W); Vols,
Polatschek
s.n. (W); Ainet,
1976, Polatschek
ob Drassling,
s.n. (GZU); Zeltweg,
1933, Leopold
near Graz,
s.n. (GZU); Puntigau
near St. Nikolai
s.n. (GZU); St. Oswald
1963, Melzer
s.n. (W); Brixlegg,
latschek
in Hainsdorf
Schwarzau
s.n. (GZU); Wies,
TIROL: Aguntum,
1897, Palla
s.n. (GZU); Liebach,
(GZU); distr. Leibnitz,
191
Melnik,
Kralupy,
Jehl('k & Rostaniski
(M). Denmark.
6647
ABENRA
SYSTEMATICBOTANYMONOGRAPHS
192
SODERBORG: Lundtoft,
B. s.n. (C).-ALBORG:
s.n. (C).-BORNHOLM:
Luthen
s.n. (C).-K0BNHAVN
Hansen
s.n.
(C).-PRMST0:
US,
Z).-RIBE:
Hede,
BORG: Loret
(C).-VEJLE:
1962, Hansen
Aast,
UUSIMAA: Helsinki,
523
Pasilan,
AIN: Nernay
France.
(BM, BR,
1964, Oinonen
HAUT-RHIN:
1927, Soest
near Perigeux,
near Avignon,
s.c.
Colmar,
s.n.
1888, Beauverd
HERAULT: Palavas,
1877, Verrier-Lirardery
NY, P).-ISERE:
between
1890, Perris
l'Oc6an, Masson
de-Ce, Bioret
a-Mousson,
867
St. Denis-en-Val,
(LY). Georgia.
s.n. (HBG); Hohentwiel
1909, Braun
Wasseralfingen,
603
Doppelbauer
near Miihldorf,
s.n. (GOET),
(B); Dahlem,
Roebes
Marschner
Altensteinstrasse,
5134
Scheppig
Hudziok
Hegi
1980, Gutte
near Cottbus,
lier s.n. (HBG);
Finkenwerder,
143 (HBG).-HESSEN:
(FR); Frankfurt,
s.n. (FR); Kassel,
between
s.n. (FR); Ostpark,
Tyroffs.n.
between Munden
Fuckel
1878, Chanin
s.n. (LZ).-NIEDERSACHSEN:
376
&
1911, Schulz
s.n.
1906, Gross
& Hanelt
6820
(FR); Grafenhausen
and Hedemunden
near Alfeld,
s.n. (LZ);
(B); Rangsdorf,
1862, Hoppe
Nolde
1888, Diirer
near Darmstadt,
s.n.
1892, Duirer
at river Werra,
1908, Schmidt
1901,
1900, Hal
s.n. (M); Reinbeck,
near Gottingen
1963,
Jtiterbog,
s.n. (HBG); Blankenese,
1964, Dietrich
(FR); Sachsenhausen,
s.n. (POM).
s.n. (SOM);
Bornmuller
1868,
s.n. (GZU);
1967, Pietsch
s.n. (GOET); Frankfurter Wald,
s.n. (G); Boizenburg,
Siebenberge
1957, Wagenitz
s.n. (GOET); Hasenheide,
Kopenick,
1866, Schmidt
s.n. (HBG); Hamburg,
1848, Bartlinz
near Wiesbaden,
BURG-VORPOMMERN: Lychen,
& Henker
and Bmrnsen,
bridge
(M); Wald
1968, Sukopp & Poelt
s.n. (LZ); Ruhland,
1957, Fouquet
at river Werra,
s.c. s.n. (GOET);
s.n. (GOET); Ostrich
Bergedorf
1927, Vogeler
Allendorf
1909, Reipers
s.n. (C); Berlin,
s.n.
at
1962,
at Aisinger
Berg,
Spandauer
1957, Wagenitz
s.n. (KTU); Potsdam,
1965, Hudziok
at river
s.n. (GOET); Mtihldorf
1957, Griitzmann
s.n. (GZU); Wannsee,
1985, Nilsen
s.n. (B);
Gunzburg
station Beusselstrasse,
1980, Gutte & Jentsch
1960, Hiugin
1905, Zimmermann
s.n. (M); Mangfall
1956, Wild
s.n.
1859,
s.n. (M); Muinchen-Obermenzing,
s.n. (GOET); Charlottenburg,
(B); Grunewald,
1869, Gobert
s.n. (GOET).-BAYERN:
Railway
near
1874 &
Heidelberg,
and Freiburg,
Lorrach
near Berchtesgaden,
s.n. (LZ); Dahlwitz,
& Jentsch
s.n. (KTU); Luckenwalde,
s.n. (Z).-HAMBURG:
484
St. Hilaire-de-Riez,
1949, Merxmiiller
1979, Scholz & Poelt
Bonsecours
and Bezons,
Argenteuil
at river Isar, 1885, Peter
(M).-BERLIN:
de
St. Laurent-d'Audenay,
BADEN-WURTTEMBERG:
s.n. (Z); between
s.n. (M); Passau,
(HBG); Tiergarten,
BRANDENBURG: Greifenhain
Senftenberg,
173
between
s.n. (M); Schwetzingen,
Hollriegelskreuth
& Scholz
Meyer
s.n. (HBG); Schoneberg,
Westend,
1889, Kdser
Pont
Bois
s.n. (CAS).-PYRENEES-ORIENTALES:
s.n. (B).-VENDEE:
Schneizelreuth
(MJG);
1957, Wagenitz
de Champions
Agen,
Les Pont
s.n. (LY).-PARIS:
s.n. (LY).-SEINE-ET-MARNE:
at river Saal, Armburg
1900, Renner
Troll 6050
s.n. (DS).-MAINE-ET-LoIRE:
(MA).-SA6NE-ET-LoIRE:
(LE). Germany.
s.n. (M); Mtinchen-Laim,
St. Brevin
s.n. (FI).-LOT-ET-GARONNE:
1932, Alleizette
371
(BR).
Mont-de
LY, P, Z).-LANDES:
s.n. (LY).-LoIRE-ATLANTIQUE:
1880, Magnier
1885, Forster
s.n. (L); Weyn
s.n. (M); Neu-Ulm,
near Rosenheim,
Scholz
s.n.*
near Singen,
(M); Griinwald,
river Inn, 1976, Maschner
kraiburg
Vasilev
s.n. (W); Mannheim-Muihlau,
& Neumann
Roessler
1916, Beger
Montrieux,
1949,
Klukhovi,
s.n.
(B, BR, C, F, LY,
s.n. (BR).-MEuRTHE-ET-MosELLE:
1892, Chabert
Bois
553
near Noyon,
Lyon, Mutel
(POM).-VAL-D'OISE:
(FI).-HAUTE
1915, Berger
Issoire,
Chamb6ry,
s.n.
Liveilli
1892,
(DS, Fl, LISU,
Geirard
s.n. (AMD).
Tours, Delaunay
1896, Gautier
Carlefesse
s.n. (DS).-RH6NE:
at river Doubs,
Bertrand
1857,
1902, Girandias
Ste. Marie-a-Py,
s.n. (LY).-OISE:
s.n. (LY).-SAVOIE:
13416
4097
de St. Georges-de-Cher,
1962, Stuurman
Aixe-sur-Vienne,
1847, Chirat
Le Collet-de-Deze,
(BC, P).-MARNE:
s.n. (LY).-VAR:
1898, Rouy
Donau,
Lombard
(G).-LOIRET:
1881, Gautier
Schmidt
and Tencin,
Montbrison,
Jordan
Tidestrom
Scey-sur-Sa6ne,
s.n.
1887, Autheman
(BC, BM).-DORDOGNE:
Baume-les-Dames
s.n. (LY).-INDRE-ET-LoIRE:
1860, E. s.n. (LY).-PUY-DE-DoME:
1891, Gandoger
Rouen,
Goncelin
Bais
(BR).-GARONNE:
(G).-HAUTE-VEENNE:
s.n. (LY).-LOIRE:
1048
1847,
Vincennes,
Millas,
s.n.
s.n. (LY).-LozERE:
1860, Debeaux
1803
& Magnier
Sennen
1904,
Bayonne,
6117 p.p.
Desplantes
s.n. (L).-DOUBS:
Geerinck
(M).-HAUTE-SAONE:
SAVOIE: Gaillard,
Marsan,
Semur-en-Auxois,
184 (BRY).
Chailvet, Martin
Isle Ste. Lucie,
AHVE
s.n. (MA).
Jackson
(Pietarsaari),
s.n. (LAM).-CHARENTE:
1974, Ledoux
Orgon,
Remoulins
787 (LY).-GARD:
Jokobstad
iEr0,
1955, Hansen
1876, Hollmen
s.n. (P).-AISNE:
s.n. (LY).-AUDE:
Contr
1872,
s.n. (BR).-C6TE-D'OR:
de Connezac
St. Vincent
JAPoRI: Turku,
1836, Martel
s.n. (C).-SKANDER
s.n. (C). Finland.
Fredakild
1949,
s.n.
(A, BR, L, LD,
Bredebro,
route 988, 1968, Jean s.n. (LILLE).-BASSES-PYRENEES:
Erstein,
1896, Felix
Funder,
s.n. (H, LD).-VAASA:
Vichy,
s.n. (FI).-BoucHEs-Du-RHdNE:
Vierzon,
s.n. (C).-T0NDER:
s.n. (H).-TURKU
at river Rh6ne,
near Lagnieu
F1, LY).-ALLIER:
(LY).-BAs-RHIN:
s.n. (C).-VIBORG:
1963, Haakana
Kattby,
388
s.n. (C).-SVENDBORG:
1969, Hansen
1962, Jensen
Tjerristsbo,
1968,
1965, Hansen
& Pedersen
1967, Hansen
Herming,
Kirkehavn,
s.n.
1928, Os
Vester0havn,
Fyn, Agernmes,
Larsen
Djursland,
s.n. (C).-RINGK0BING:
s.n. (C).-SOR0:
Thorming
s.n. (C).-LAEs0:
(C).-ODENSE:
1962,
Arhus,
1862, Leth
Farum,
s.n. (C).-HOLBAEK:
1935, Lund
1840, Benzon
s.n.
s.n. (C).-RANDERS:
s.n. (C).-THISTED:
NANMAA: Hammarland,
Jensen
s.n. (C).-ARHus:
(LD).-FREDERIKSBORG:
Bagtorp,
Federiksdal,
1962,
1955, Vesterager
1974,
1959, Hansen
Soby,
s.n.
Skor,
Nabskod,
1961, Jensen
R0dveg,
s.n.
s.n. (C).-HJ0RING:
(Copenhagen):
(C).-LOLLAND:
1965, Christensen
Hyllebjerg,
1885, Henningen
1937, Andersen
(DS).-HADERSLEV:
tenfeld
Hasle,
VOLUME 50
1852, s.c.
s.n. (M).-MECKLEN
s.n. (B); Gustrow,
s.c. s.n. (GOET); Braunschweig,
1981, Gutte
1906, Ferrez
1896, Peter
s.n. (LY); Celle,
s.c. s.n. (GOET); Hildesheim,
Nienburg
at river Weser,
Wunstorf,
1865, Freund
23993
burg, Soest
Dahner
(GOET); Mainzer
191 1, Beger
1966, Gutte
near Rosslau,
1978, Gutte
et al. 170
al. 10460
s.n. (HBG); Kiel,
Boros
gymanyos,
s.n.
Szurdk
BRca,
s.n. (BP).-KoMAROM:
Dorog,
1949, Boros
s.n. (M).-ZALA:
Jdvorka
1900,
1882, Hermann
PEST: Czepel,
s.n. (BP).-VAS:
Szeged,
Polgar
1650
Luzzara,
336
Prov. Trieste:
s.n.
LATIUM. Prov. Roma: Roma,
1954, Cacciato
LOMBARDY. Prov. Brescia:
s.n. (FI).-REGION
ico at Lago
Pavia:
Pavia,
s.n.
1856, Rampaldi
Prov. Ancona:
MARCHES.
near Casale Monferrato,
Soldano
(TO). Prov. Torino: Castagnole
Prov. Nuoro:
Sorgono,
s.c. s.n.*
1833,
dano 4941
s.n. (M). Prov. Verona:
Angerer
ford & Wood
19696
1904, Rigo
Cervino,
s.n. (KYO); Hidaka
1958, Okamoto
Sohma & Takahashi
614
Boufford
& Wood
19760
Akita
Pref., Mt.
Chiba
Pref., Kazusa,
Pref., Nishi
gun,
Kurihara
nikusen,
11034
Ohashi
& Ueno
11029
Pref., Fukuzumi,
s.n. (MAK);
Naito
(GH, US); Nagasaki
Ishikawa
Pref., Nikko
Marugama
Taki-gun,
& Wood
Iwaki-shi,
Pref., Kawauchi
1967, Hosomi
s.n. (MAK);
Ogawa-machi,
Satou-cho,
s.n. (KYO);
Tatsunokuchi-cho,
Mieno
395
Migazu-shi,
1969, Katsuhiro
247
Asa
Ibaraki
N6mi
(TUS); Kamagawa
Horie
near Karuizaea,
Shinano-Oiwake
City, 700 m, Takenaka
s.n. (KYO);
9.2 km S of Shibecha,
1957, Mori
Pref.,
Osawa,
19656
Pref., Furano City,
654 (KYO).-HONSHU:
Mutsu,
Fukushima
Sol
1981,
Pref., Rubeshibe,
1962, Nitta
Pref.,
Pref., Amanohashidate,
Waniura,
1907, Som
Bibione,
City, Boufford
Pref., Shimotokuyama,
Pref.,
SARDINIA.
236 at Samani, Bouf
(MO); Kamikawa
Hamakoshimizu,
Mt. Goyozan,
(FSU); Nagano
Pref., Kamitsusima
15 (KYO); Tochigi
19859
(TUS); Hiroshima
Taki-cho,
s.n. (MAK); Kyoto
72630
8 km N of Hwy
Shimokita-gun,
s.n. (KYO);
s.n.
(BR).-REGION
HOKKAIDO: Abashiri
Hakodate-shi,
Pref.,
Iwate Pref., Ofunato-shi,
1962, Kobayashi
Ito & Kinoshita
Pref., Yukawa,
s.n.
1984, Soldano
Forte dei Marmi,
s.c. s.n. (MAK); Nemuro
(TUS); Aomori
Sesia,
Terranova
& Forneris
s.n. (FI).Prov. Venezia:
Okada,
Koshinizu-cho,
1967,
1938, Tagawa-Motozi
1929, Murata
Shiogama-shi,
City,
Oshima
635
s.n. (TI); Hyogo
s.n. (MAK);
Pref., Kamakura-shi,
Pref.,
Endo
Itinomiya,
ibaragi-gun,
1968, Fukui
Miyagi
(KYO, MO);
Ushiogawa,
1971, Enomoto
Pref., Shari-gun,
702
E of Tomakomai
& Wood
City, Boufford
beach, Kushiro
Taihei-zan,
Natsuigawa-keikoku,
gun,
(H); Kitami
Pref., Otanoshike
Samanicho,
Iburi Pref., Hidaka,
(KYO, LAM, MO);
Ishikari Pref., Sapporo
Kushiro
s.n. (LY). Japan.
Pref., Samani-gun,
(KYO, LAM, MO);
at River
Col
s.n. (FI). Prov.
s.n. (MO). Prov. Pisa: Migliarino,
1893, Fiori
Piove,
1838,
(PAV).-REGION
1981, Montacchini
Hausmann
1980, Marchesetti
di Massa,
La Spezia,
s.n.
Lenga
TUSCANY. Prov. Lucca:
(HAL).-REGION
Marina
VENETO. Prov. Padova:
(MO).-REGION
Bozen,
Evers
s.n. (FI).-REGION
PIEDMONT. Prov. Alessandria:
San Marzanotto,
Prov. Bolzano-Bozen:
s.n. (FI). Prov. Massa-Carrara:
Prov. Reggio
Grado,
at river Po, 1889, Fiori
(MO). Prov. Vercelli:
4856
Soldano
Piemonte,
(L). Italy. RE
s.c. s.n. (M). Prov. Como:
at river Adda,
s.n. (FI).-REGION
(MO). Prov. Asti:
3654
1968, Rieger
37595
1888, Tacconi
1912,
Sermide
Castello
Sondrio:
1890, Profete
TRENTINO-ALTo.
(MO).-REGION
mier
(FI). Prov.
Ancona,
Balaton,
Backe
LIGURIA. Prov. La Spezia:
di Garda,
Lago
s.n. (BP).
EMILIA-RoMAGNA.
(FI).-REGION
s.n. (Z). Prov. Mantova:
1901, Geilinger
di Como,
s.n. (BP).-VESZPREN:
s.n. (SMU).-REGION
s.n.
1956, Lengyel
s.n. (BP).-SZABOLCS-SZATMAR:
s.n. (FI). Prov. Udine:
Parlatore
La
Tauscher
Erosi,
1943, Boros
(Baranya): Keskend,
et
near Stari-Slankamen,
(BP).-BUDAPEST:
of H6rvolgy,
valley
Larsen
MAKEDHO
FRIULI-VENEZIA GIULIA. Prov. Gorizia:
1874, Marchesetti
Trieste,
Bukk,
Z);
s.n. (C);
near Lauenburg,
s.n.
JAVATIMUR: Pasuruan,
1843, Avellino
s.n. (FI).-REGION
Flens
WVA,
1955, Hansen
(BP).-FEJER:
1964, Kdrolyi
Ortilos,
s.n. (BP). Indonesia.
1881, Pirotta
nell'Emilia:
Mt.
1936, Kovdcs
Koszeg,
1943, Boros
Kotor,
GION CAMPANIA. Prov. Caserta: Matese,
(GZU).
1907, Budal
s.n. (BP).-PECS
SMU, WTU,
Titel
BAcs-KISKUN:
Zsolca,
s.n. (BP).-SOMOGY:
1979, Gutte
s.n. (FR). Greece.
1894, Goldschmidt
s.n. (BP).-HEVES:
1930, Polgar
(BP).-GYOR-SOPRON:
Kisvarda,
(BP).-CSONGRAD:
LD,
Felso
(BP).-BoRsoD-ABATJ-ZEMPLIN:
1908,
M, MA,
near Eckernforde,
s.n. (LY). Hungary.
1889, Charrel
(Salonique),
s.n.
KYO,
&
Rodleben
s.n. (C).-SCHLESWIG-HOLSTEIN:
s.n. (HBG); Sandkrug
1967, Straka
at river Werra,
Vacha
s.n. (LZ);
1975, Gutte
s.n. (HBG); Haldensleben,
1922, Schmitz
GZU,
Wolf 2254
1967, Gutte
near Leipzig,
1937,
1949, Mi
Bautzen,
s.n. (WRSL).-SAcHsEN-ANHALT:
s.n. (HBG); Hemmelmark
1920, Schmidt
(MJG); Oppenheim,
s.n. (LZ); Leipzig-Connewitz,
1894, Gelert
GOET,
C, COLO,
(BR, C, FI).-THURINGEN:
NIA: Tessaloniki
1943,
(BM, BR,
1895, Ohl
Kiel-Friedrichsort,
at river Elbe,
s.n. (GOET).
s.n. (B); Weinbohla,
1922, Beger
& Gutte
1967, Rostaniski
s.n. (LZ); Domitz
near Hamburg,
Geesthacht
1968, Gutte
s.n. (LZ); Leipzig,
s.n. (HBG);
s.n. (M); Ludwigswinkel,
& 2135
s.n. (Z); Altenburg
1875, Wolf s.n. (FI); Tangermuinde,
s.n. (LZ); Magdeburg,
burg, Pedersen
Klotz
(GZU);
s.n. (C); Duis
C.B.
1837,
s.n. (CAS).-SACHSEN:
near Pirna,
s.n. (B); Mugeln
s.n. (LD); Leipzig-Knautkleeberg,
(Cologne),
2134
2188
s.c. s.fn. (GOET);
1890, Schmidt
1950, Heine
Sand, Hecker
1852,
1884, Wilshusen
near Bonn,
(L); Koiln
1871, Mettzbaer
s.n. (LZ); Zwickau,
& Heier
1967, Muller
12183
Meyer
Gartow,
near Hemmoor,
Beuel
(MJG); Ludwigshafen,
near Bad Kreuznach,
titzer s.n. (LZ); Dresden,
(LA); Chemnitz,
1053 & 1054
3232
Wagenitz
s.n. (Z); Wolistein
Brandis,
Kern & Reichgelt
Magin
s.n. (GOET); Hannover,
s.n. (GOET); Luneburg,
s.n. (GOET); Warstade
1858, Ndldeke
(L); Emmerich,
Felsenland,
1911, Joesting
s.n. (GZU).-NORDRHEIN-WESTFALEN:
RHEINLANDPFALZ: Leeheim,
Ddniker
1851, Stieg
s.n. (GOET); Gottingen,
Heersum,
Zahn
193
OENOTHERA
1997
s.n. (TI); Nara
(TI); Tokyo
42 (KYO);
Beattie
&
Pref., Nin
Pref., Tama
194
SYSTEMATICBOTANYMONOGRAPHS
gawa,
s.n. (CAS, UC, WVA);
1936, Makino
(KYO); Wakayama
Pref.,
Pref., Higashine-shi,
Inosawa,
(KYO); Yamanashi
s.n. (TI).-KyuSHU:
Kumamoto
43885
(KYO). Korea
near Riga,
1947, Wikiele
s.n.*
s.n.
[South].
(WI). Luxemburg.
Madagascar.
5229
1580-1590
at river Dnjestr,
DRENTE: Noord
tema s.n. (L).-GELDERLAND:
megen,
Winschoten,
Duiben
(L); Valkenburg,
2582
s.n.
deko
(L).-LIMBURG:
(L); Wieringermeer,
s.n. (L); Rotterdam,
59/170
Ashwin
69a
AGDER: Lillesand,
Ouren
Halden,
s.n.*
(WA).-BYDGOSZCZ:
Gdansk);
Bialowdzka
Ruda,
G6rna,
1976, Chycki
Stary, 1971, Sendek
chowice,
s.n.*
Wielka
Pulawy
s.n.*
(Fl, GZU,
s.n. (KTU); Czestochowa-Aniolowa,
s.n.* (KTU).-KIELCE:
Wadowice,
s.n.
tatiski
Losice,
1972, Rostaniski
near Zamsc,
(LBL); Klemens6w
(KTU);
1968,
s.n.*
Rostaniski
s.n. (HBG); Boleslawiec,
gau), 1901, Schmidt
285
(Z). Reunion.
Talmaciu
(Talmatsch)
Cadet
s.n.*
(DAO,
2152
Althof
Rainha
s.n.*
Stara
1979, Latowski
KRAM,
KTO,
Lubelski,
Nysa
L);
1964,
(Neisse),
Aniol
(WRSL).-POZNAN:
432
Cybulski
1855,
Walbrzych
s.n.* (WRSL).
1975, Ros
(BM); Korczew
s.n.
Sadebeck
(Waldenburg),
Sta. Anna,
(Hermannstadt),
1909-11,
Jacobsen
Schur
s.n.
(Glo
AzORES: Pico,
1871, s.c. s.n.
at river Tejo, da Cunha
1903,
near
(HBG);
GORA: Glog6w
Portugal.
Fi
760
Trzcianka
(LOD).-SZCZECIN:
(LD).-MADEIRA:
Sibiu
s.n. (KTU);
(LOD); Chorz6w
s.n.* (LOD).-LUBLIN:
s.n.* (WRSL).-ZIELONA
ARGES: Calimanesti,
(P).-BRASOV:
(BR, Fl, GZU,
s.n.*
Abrantes,
1928,
1958, Tacik s.n.* (KRAM).-L6DZ:
near Wroclaw,
4851
(Pharm.
s.n.* (LOD); Kielce,
(LBL).-OPOLE:
1961, Rostaniski
s.n.*
Heubude,
s.n. (KTU); Pila,
near Tomasz6w
LD, SOM, WRSL);
s.n. (PO).-SANTAREM:
near Sibiu, Schur 6271
s.n.*
1889,
(TOR).-KATOwICE:
s.n.*
1955, Warcholinska
Belzec
(POZ).-WARSZAWA:
at river Odra,
(K). Romania.
346
Czyzewska
1962, Rostaniski
Pinhal Novo,
1965, Costa
1956, Jedras
1962, Rostatiski
1984,
Fl, GZU,
at river Bobr,
s.n. (HBG); Krosno
da Gaia,
(LD); Racib6rz,
Szmajda
Tryb.,
Larvik,
& Rostanski
1973, Piasecki
and Wisla,
(KTU);
1976, Lysiak
(WSRP).-WROCLAW:
96 (PH, US).-EsTREMADURA:
PORTO: Vila Nova
s.n.
s.n. (B); Kalisz-Winiary,
Pyrzyce,
Glowacki
Wroclaw-Midolajow,
911
Rostan'ski
1906, Bothe
1974,
Knorn
s.n.* (LOD); Stok near Piotrk6w
(Weichsel),
s.n.*
Rau
1921, Hryniewiecki
1960, Giers
1969, Szotdowski
Lancucka
of rivers Nidzica
Park,
s.n. (0).-AUST
(TOR).-GDAN4SK:
1975, Rostatiski
near Andrych6w,
Wieprz
confluence
s.n.*
(Gleiwitz),
Ustka,
Hagley
s.n. (K).-0stfold:
(BIL); Lake Wigry,
near Wloszowa,
Chrzast6w
s.n.* (KTC).-KOSZALIN:
Co., Piotrowice
SOM),
Gliwice
s.n.
1941, de Bruyn
Collett
1885, Klinggraeff
s.n. (KRAM);
1960, Sowa
LD,
(Schonlanke),
near "Mattembleuch,"
1949,
1971, Beck
(O).-VESTFOLD:
near Tuchola,
1885, Hohnfeldt
& Rostaniski
at river Wista
jalkowski
Forst"
s.n.
(L); Groenekan,
Bodegraven,
1864,
34479
s.n.*
1963, Koteja
(POZ).-KRAK6w:
L6dz, Rokicie,
PO).
"Oliver
(L).-OVERIJSSEL:
1951, Andressen
(Christiania),
Ouren
s.n. (M); Plaskosz
(Schwetz),
2834
s.n. (CHR, POM).-WELLINGTON:
Drobak,
1971, Sokolowski
Torun' (Thorn), Renner
near Swiecie
1978, Maciejczak
Kazimierza
Brown
BIALYSTOK: Bialystok,
Terespol
Oslo
Vallberg,
1920,
1895, Be
Velsen,
CANTERBURY: Christchurch,
1938, Hodgkins
(O).-OSLO:
Kragero,
(L); Valkenswaard,
s.n. (L); Maassluis,
Jongh
Zealand.
21395
5341
s.n. (L).-ZUID-HoLLAND:
1955,
near
Ooststroom
1924, Kern & Reichgelt
Ooststroom
Dolder,
AKERSHUS: Akershus,
s.n.
Leenhouts
s.n. (L); Nij
Zuiderveen
WVA);
s.n. (L); Tubbergen,
(L); Leiden,
s.n. (L). New
(CHR). Norway.
s.n. (M); Gdaiisk,
Mayer
2329
(0).-TELEMARK:
s.n. (0). Poland.
(L); Den
AUCKLAND: Tauranga,
1921, Hesselberg
27420
Dyring
Bot.
s.n.
1900, Jansen & Wachter
(CHR).-SouTH
Beach,
Quadgras
(L); Zandvoort,
1928, Kurseman
1887, Lako
Renesse,
(Den Haag),
s.n.
Harshagen
1896, Bondam
Amersfoort,
(L); s'Gravenhage
mati
1966,
260
s.n.
1969, Bein
Rust
s.n. (L); Meerssen,
s.n. (SMU, UC, WTU,
1957,
1916, Pando
Tetouan,
Schiermonnikoog,
Van den Houten
Breda,
Jonkes
del Carmen,
near Kalarash,
s.n. (L); Gendringen,
1961, Spaargaren
1951,
s.n. (L); Hengelo,
1946, Hooglund
s.n. (UC).-ZEELAND:
Glerum
Healy
Den Helder,
s.n.*
s.n. (BR).
1831, Lejeune
s.n. (UC).-GRONINGEN:
1948, Roosje
Heerlen,
(WI);
1965, Natkevicaite
(DS); Bachtut
(L).-FRIESLAND:
1942,
s.n.*
COAHUILA: Sierra Madre
(LW). Morocco.
Veth & Koopmans
1956,
1910, Kupfer
(Memel),
Gronskaite
1954,
near Diekirch,
et al. 929
s.n.*
6017
s.c. s.n. (L).-NooRD-BRABANT:
1901,
(L).-UTRECHT:
Waalkes
s.n. (L); Wageningen,
s.c. s.n. (L).-NoORD-HOLLAND:
Denekamp,
Iwanow
(Wolmar),
near Ignalina,
(MO). Mexico.
Borisova
1898,
Drente,
Apeldoom,
1954, Kern & Reichgelt
29971
Korneshty,
(K). Latvia.
at river Lielupe,
near Klaip6da
at river Neman,
(KYO);
Yamamoto
4423
Jelgava
of Valmiera
Bachmann
s.n. (BR); Bettendorf
27886
Kochi-shi,
s.n.* (LATV);
(Pemigel) W
(KYO); Miyazaki
Seoul, Dunn
(MICH);
Oka 35204
& Kobayashi
Yamamoto
Pref., Asakura,
s.n.* (WI); Pusilieskies
m, Croat
(ENCB). Moldavia.
7551
(Z). Lithuania.
near Vilkija
1856, Prarets
d'Arval,
(MW); Pirasajal
(BC). Netherlands.
639
1902, Mobimienko
Fianarantsoa,
s.n.*
at the sea near Liepupe
1914, Dieterlen
12020
13309
Yamagata
Yoshiki-gun,
Shimada
1977, Tabaka
(KYO);
Mizushima
Isoura, Niihama-shi,
In-Cho
Kyongsi-Do,
Apanagusiasa-Nevezio
Abbaye
& Robert
Cheban
Leribe,
Pref., Ogouri,
31 (TUS); Kochi
s.n.* (LATV); Daugavpils,
at river Neman,
Druskininkai
Passini
Kangnung,
on cliffs
(LATV);
(HBG);
Kusaka
Pref.,
592
Kitakoma-gun,
Kumamoto-shi,
Ehime
Tanaka
Hino-gun,
Hatsuyama
(TUS); Yamaguchi
Ima, Demizu-cho,
Asahishin-machi,
1979, Fatare
(RIG). Lesotho.
Magnus
Pref.,
Mizogu-chi,
Nishimuro-gun,
station, Takane-cho,
(TUS).-SHIKOKU:
Pref., Takamatsu-shi,
Pref., Kanaya-dani,
et al. 10762
Ohashi
Oka 45270
Brasas
s.n.*
Tottori
Shirahoma-cho,
Pref., Kiyosato-ryo-Kiyosato
Pref., Nichinan-shi,
Kagawa
Sandan-peki,
VOLUME 50
1397
(LISU,
(G).-BACAU:
s.n. (L).-CLUJ:
Aiud
1997
OENOTHERA
(Nagy-Enyed),
1892, Csato
(Kara-Orman),
1874, Sintenis
atra Closanilor
Muresul
Mtns,
(Maros),
lau), 1940, Schiitt
s.n. (BREM);
(Juchnov),
Skvortsov
1977,
Garden,
1913, Kozo-Polyanskiy
& Preobrazenski
dimova
1232
& Konovalova
Schreter
& Barkalov
Kozachenko
19383
& Rostaniski
Skvortsov
s.n.*
s.n.* (MHA). Rwanda.
Slovakia.
1902, Waldhaus
(Pressburg),
River,
1974,
9311
(PRE). Spain.
Castroviejo
Tavira & Tormo
s.n.
s.n. (JACA).-JAEN:
273
1941,
Salto
Ojeda
Monaragala
Dist.,
s.n.
de Miller,
1899, Gyllenkrok
(LD); Augerum,
657
s.n. (LD).-GUNNARSNAS:
s.n. (LD); Vasta
Fries
(LD).-GOTLAND:
1903, Bolger
Blixt
(LD);
Tryde and Salsborg,
Ryden
1896, Kurch
1907, Gorton
s.n. (LD).-MALMOHUS:
Persson
Brunnby,
1860,
Hogestad,
1910, Ahlin
s.n.
Steijern
s.n. (LD); Skanor,
narslov,
1943, H&ikanson
(LD); Koping,
1951, Malmstrom
s.n. (LD); Vomb,
1951, Olsson
(LD).-OSTERGOTLAND:
Stenstorp
Hastbacken,
1906, Lundvall
& Nelini
1886, Ekdal
(LD),
s.n. (LD); Knislinge,
1947, Oredsson
s.n.
1926, Bjornstrom
s.n. (LD).-OLAND:
1891, Hamnstrom
s.n. (LD); Krokek,
Glbmminge,
1915, Ander
Ljungby,
and
1916,
s.n. (LD);
s.n.
s.n. (LD); Rang,
s.n. (LD); Vastra
1930, Almborn
s.n. (LD); Kumla,
Get'a, 1944, Kjellmert
Borlange,
Osterslov
1926, Nilsson
1952, Brandt
(LD);
s.n. (LD);
1953; Dege
1976, Wieslander
(LD); Hallestad,
s.n. (LD); Lund,
1965,
s.n. (LD); between
s.n. (LD).-KRONOBERG:
s.n. (LD); Barkakra,
s.n. (LD); Sodra Sandby,
Grave,
Thornm s.n.
s.n. (LD); between
s.n. (LD); Stoby,
s.n.
1905, Slott
1941, Bjornstrom
1940, Gunnarson
1946, Linders
Olsson
1882, Hammstrom
1936, Johansson
1939, Wallin
1897,
s.n.
s.n. (LD),
s.n. (LD); Tofta,
s.n. (LD).-KOPPARBERG:
1939,
Agered,
s.n. (LD); Tors,
1896, Wigforss
1891, Soderberg
Angelholm,
1941, Weimarck
s.n. (LD).-OREBRO:
Alvestad,
s.n. (LD); Halmstad,
s.n.
s.n. (LD).
1925, Hylander
s.n. (LD); St'anga, 1911, Fries
s.n. (LD); Locknevi,
s.n. (LD); Vinslov,
(LD); Dalby,
(LD); Romelanda,
and Skottorp,
s.n.
1922, Holmgren
OCH BOHUS:
(LD).-GOTEBORGS
s.n.
Guecho,
Sri Lanka.
s.n. (LD); Kdllunga,
Asarum,
1916, Kuhler
Norra Akarp,
s.n. (LD); Loderup,
s.n.
Palmer
(MA).
1914, Rasmusson
Hogsby,
1974,
1975, Montserrat
1896, Waldenstrom
Anderstorp,
s.n. (LD); L'angebro,
s.n. (LD); Simrishamn,
Sandholmen,
1893, Ortengren
Listershuvud,
La
1986,
Cape Plana, Cots &
s.n.
(LD).-JONKOPING:
(LD).-KRISTIANSTAD:
Santonia,
(BC, BM, G, MA).
Arbeca,
s.n.
s.n. (LD); Virserum,
s.n.
4403
s.n. (LD).-BLEKINGE:
1953, Lundegren
Stenberg
Crocodile
s.n. (MUB).-ASTURIAS:
Rios
1992, Aldosoro
Lindstorp,
1950, Fries
Skummeslov
s.n. (LD); Hassleholm,
1930, Stenberg
1889, Olin
Alvshog,
1903,
1934, Bergtsson
1927,
Ward 6173
1987, Lafnz s.n.* (herb. Lafnz).-VIZCAYA:
Westerlund
s.n. (LD); Hellvi,
s.n. (LD).-KALMAR:
s.n. (LD); Kverrestad,
Ekestad,
1951,
(Windis
(ZA). South Africa.
Santa Cruz de La Seros,
ALVSBORG: Amal,
s.n.
(Z).-VAPA
Pietermaritzburg,
1150 m, Sennen
s.n. (LD); Mjallby,
s.n. (LD); between
1867, Gustafsson
Dommarvet,
berga,
1882, Berg
Vinberg,
Nodstedt
Jonkoping,
Hudiksvall,
s.n.
110 (PRE).-TRANSVAAL:
s.n. (MUC).-LERIDA:
1944, Willen
(LD); Foss, Mundekal,
s.n. (LD).-HALLAND:
1882, Berg
Olmevalla,
s.n.
Tunhem,
1926, Rundkwist
s.n.
Bro,
(PDA). Sweden.
Usman',
1895, Movi
Slovenjgradec
1704*
s.n.
1962,
(MHA).-VOL
near Nov
s.n. (MA).-CANTABRIA:
de S. Francisco,
Barrio
1951, Nilsson
Kummeln,
1942,
Rios
Santonia and Raos,
(MAF).-ZAMoRA:
GAVLEBORG: Halsingland,
Rosered,
Ukna,
between
1982, Rico
s.n.*
1898, Kupeok
Slovenia.
1972,
Sloboda,
San Marino.
Paulin
1988, Segundo
Puigcerda,
1987, Segundo
Skvortsov
(MO).-NATAL:
s.n. (LY).-HUESCA:
1882, Gil
Passara, Austin
(LD); Gamlebyn,
1394
Jerte,
La Cerdafia,
de Jajufica,
(LD).
1700 m, Jarman
River,
(MA).-CACERES:
(BC).-SANTANDER:
Pozo
Goldblatt
s.n.*
1988,
Rogachevka
near Ljubljana,
ALBACETE: La Graya,
s.n. (MA).-GERONA:
GUADALAJARA: Soranca
Bold.
Cape Peninsula,
1951, Torelowa
delta of river Don,
Zabere'e,
(Grau),
s.n.
Chrtek
1950,
Bay, near Ugol
1951, Libarskij
(K, MO).
at river Hron
1963,
FREE STATE: bank of Calodon
Burt-Davy
Arena,
Zohor,
on Iturup Island, Pobe
at river Suchan,
1976,
4540, p.p.
s.n. (GZU); Lake Notranje-gorice
CAPE PROVINCE: Kalk Bay Mtn,
(K, MO).-ORANGE
Auquier
Ruhengeri,
(MO);
s.n.* (RV).-RYAZAN:
(MHA).-VORONEZH:
STREDOSLOVENSKEY: Brezno,
DOSLOVENSKEY: Bratislava
chgraz),
s.n.*
s.n.
KRAY: Nakhodka,
(DS, GH).-SMOLENSK:
at river Kuban,
Karachayevsk
Lowelius
1968,
s.n.
s.n.
1972,
at river Fars near Maykop,
Dugino,
(LE); S-Sakhalin
1902, Riskina
Alagir,
(MHA).-STAVROPOL':
1945, Voroshilov
(MIN, NY).
Caucasus,
Akhtuba,
1988, Fedajewa
414
Be?ekov
1915,
near Sebezk,
Krupova
s.n.* (LGU).-ROSTOv:
Suchoboskij
Palatki,
(LE); marsh E side of Uglovoy
(MO).-PSKOV:
1960, Shmidt
Chekhov,
Nikolinski,
(DS).-PRIMORSKIY
17621
Sdorovska
1940, Schutt
Yukhnov,
OSTROVA: Kuril'sk
10186
s.n.* (LGU); Romanovskaya,
(A).-SEVERO-OSETINSKAYA:
GOGRAD: Srednyaya
Skvortsov
19417
(MO),
Kuybyshev,
(MO);
s.n.
(Pil
Baltiysk
s.n. (HBG); Rad
(Rossiten),
s.n. (DAO). -KRASNODAR:
(RV).-KURIL'SKIYE
at river Tinok,
s.n. (FI). Sakhalin,
Seroseova
s.n.*
near Pushkinskiye,
(LGU); Selichnovo
(M, MO);
(LE).-MOSKVA:
s.n. (DS); Partisanskiy
naya, Solomon
s.n.
1940, Drosdova
1892, Grutter
s.n.
Skvortsov
Pi
at river
1968, Skvortsov
Novoborisovka,
s.n. (GOET); Rybacij
1973,
Vadu
distr. Mehedinti,
et al. s.n. (M).-KALININGRAD:
near Dabrovaol'sk,
1870, Peter
Ozerna,
s.n. (M).-DOBROGEA:
UNKNOWN: Transsylvania,
BELGOROD: Borisovka,
(Pillkallen)
(Gumbinnen),
Yukhnov
(MO),
s.n. * (CL).-PROVINCE
1928, Nyarady
Tulpeningken
Renner
(Karlsburg),
1965, Toma s.n. (H, M).-TIMIA:
at river Iput', 1980, Skvortsov
near Gusev
spont. Botanical
Leningrad,
lulia
Probata,
s.n. (MIN). Russia.
Klintsy,
(BREM).-KALUGA:
Alba
(LD).-IASI:
m,
1889, Csato
lauken at river Pissa
s.n.
953
1200-1427
(MHA, MO).-BRYANSK:
Skvortsov
s.n. (RSA).-DEVA:
195
1913, Hjort
(LD);
1927,
Son
s.n.
s.n.
s.n. (LD); Linkoping,
SYSTEMATICBOTANYMONOGRAPHS
196
holm City,
s.n.
s.n. (LD).-UPPSALA:
(LD); Sunne,
(LD).-VASTMANLAND:
1882, Fischer-Sigwart
Amandus
s.n. (Z).-BERN:
1921, Rohrer
Gallen,
Rhone,
terthur, Lutz 1251
Arnold
Oaklands,
s.n.*
(MW).-DONETSK:
Donetsk).-KAMENETS-PODOL'SKY:
KHAR'KOV: Borki
1901, Pachoski
railway
s.n.*
(Bot. Gard. Donetsk).-LUTSK:
(MW).-ROVNO:
Latoriza,
1947, Bila
lenkoye,
1930, Kukush
BEDFORD: Luton,
men
4187
s.n.*
(LW).-VINNITSA:
Bay,
s.n. (BM).-LINCOLN:
232
(BM).-OxFoRD:
s.n. (BM, BREM).-SUFFOLK:
(BM). Scotland.
Zhitomir,
1915, Marshall
Banbury,
Liverpool,
1926, Corstorphine
Kingdom.
at river
London,
Bristol,
1869, Tri
1851, Dugdale
Regent's
Park,
1934,
Park,
Bow
1906, Marshall
Berrow,
Hurst
s.n. (BM).-MIDLOTHIAN:
ISLANDS:
Brake near Ollerton,
Boughton
s.n. (BM).-Surrey:
Be
England.
s.n. (K).-Cornwall:
Southport,
s.n. (BM).-SOMERSET:
1872, French
1935, Evans
(MW).
s.n. (BM).-CHANNEL
(Middlesex):
(herb.
(MW).-ZAPORSH'YE:
1968, Edmondson
s.n. (K).-NOTTINGHAM:
s.n.*
Mukachevo
s.n. (BM).-GLOUCESTER:
s.n. (BM).-LONDON
Kessingland,
ANGUS: Invergowrie,
1895, Jackson
Chester,
1605*
1938, Osadcha
1938, Osadcha
s.n.* (KW). United
s.n. (BM).-LANCASHIRE:
1834, Mann
Sambor, Madalski
s.n.*
s.n.*
s.n.*
1936, Grin
Iznik at river Vorskla,
Akhtyrka,
(LW).
at river Dnepr,
1929, Ganeschin
(KRAM).-UZHGOROD:
1907, Landa
Enbome,
Burrows,
1862, Lowe
Thorpe,
s.n.*
near
(Bot. Gard.
s.n.*
Kherson
Kiyev,
1927, Gniatnovskaja
Vinnitsa,
1974, Lousley
Laceby,
s.n. (NY).-NORFOLK:
(KW).-SuMY:
Slendzinski
s.n.*
Kuzniecova
at river Donets,
s.n.* (KW).-L'vov:
s.n. (BM).-CHESHIRE:
Braunton
Stone,
(LW).-KIyEv:
near Lisichansk
(MW).-POLTAvA:
s.n. (BM).-BERKS:
1884, Hanbury
(BM).-DEVON:
den & Hillman
s.n.*
1976,
s.n.* (MW).-ZHITOMIR:
s.n. (BM).-KENT:
Gates
s.n.*
1899, Puring
near Borshchev,
1901, Higgins
Jersey, St. Authin's
1916, Ilwinskiy
Janata
1906,
Dubno,
TERNOPOL: Dzwinogrod
s.n.*
1960, Kozub
s.n.*
Banilov
at river Samara,
Gornonog
1919,
1830
1966, Mrynski
Volnoje
s.n.* (MW).-KHERSON:
s.n. (MO).-LUGANSK:
Manevichi,
Nikolajew,
1975,
Staroushejeck,
& Marinskaya
Vyzkov,
1968, Skvortsov
Madalski).-ODESSA:
Marshall
1922, Kotov
station,
near
Biereg
Dist., Mufindi,
1970, Ruffo 353 (BR);
supply dam,
(KW).-DNEPROPETROVSK:
at
s.n. (KYO).
1936, Koidzumi
(KW).-CHRNOVTSY:
near Yenakiyevo,
Olchovatka
Bolshoj
(LE).-KHUST:
(LE); near Snitynka,
s.n.*
1952, Z. B. s.n.*
of river Cheremos,
s.n.*
1932, Dziubenko
s.n. (Z); Wyla,
1883, Siegfried
near Zolotonosha,
CHERKASSY: "Bilchi"
near Gorodnya,
at Lake
1875, Buol
Nidelbad,
Ngwasi
near
km N of Win
s.n. (Z); Herrliberg
Manzhou,
water
s.n.
at River
Illarsaz
Altikon,9
Frei
(K).-MTWARA:
165 (K); Lushoto
s.n. (Z); St.
1912, Fueter
s.n. (BR); Les Pierrettes
s.n. (Z); Rtischlikon,
s.n. (Z); Winterthur,
Biberist
1981, Geitler
1911, Visher
s.n. (Z).-VALAIS:
s.n. (Z); Glattfelden,
9820, p.p.
between
Marstetten,
1948, Brunner
s.n. (Z). Taiwan.
Rd, Richards
(K). Ukraine.
Tupichewy
at Lake Walen,
1882, Hug
Frick
Lushoto Dist., Mgaza
996
Batty
at mouth
Kotov
Dist., Masoka
(MO).-TANGA:
(MW).-CHERNIGOV:
Waniekov
(Z); at River Toss,
1554
MBEYA: Rungwe
Usambara
1881, Fries
and Weisslingen,
s.n. (Z); Zuirich, 1915, Rohrer
1896, Schinz
m, Lovett 1569
Kollbrunn
Luzern,1885,
GALLEN: Bad Ragaz,
Vetter s.n. (Z).-ZURICH:
at River Broye,
s.n.
Schaffhausen,
s.n. (Z).-SOLOTHURN:
1905, Rohrer
15 km SW Yverdon,
Bavois,
s.n. (Z).-LUZERN:
s.n. (Z).-THURGAU:
s.n. (Z); Locarno,
at River Rhine,
(Z); Eglisau
s.n. (Z); Schlieren,
River T6ss,
1910, Karl
1938,
1913, Ruffieuse
(Z).-SCHAFFHAUSEN:
s.n. (Z).-ST.
s.n. (Z); Quinten
1869, Stadler
s.n. (Z); Payerne
s.n. (Z); between
1913, Werndli
1907, Probst
s.n. (Z);
Reinach,
s.n. (Z).-GRAUBUNDEN:
1904, Schinz
s.n.
1943, Stecher
Fribourg,
s.n.
Rohrerschachen,
s.n. (Z).-BASEL:
1916, Candriani
Thielke
1916,
at Lake Zurich,
Seeger
1907,
1831, Bertschinger
Zurich, Egli
Tanzania.
at river Thur,
s.n. (Z).-VAUD:
1890, Wolf
Lausanne,
Hurden
s.n. (Z); at Lake Walen,
Bellinzona,
(Z).-TIcINo:
de Neuchatel,
1936, Frick s.n. (Z); Solothum,
s.n. (G); Henau
Garsans
1926,
Lac
s.n. (Z).-SCHWYZ:
1878, Lulger
and Derendingen,
s.n. (Z); Thusis,
1921, Schibler
s.n. (Z).-NEUCHATEL:
near Weesen
Gasi
Karlstad,
1907, Gredin
Sund,
near Brugg,
1901, Kaser
Teufen,
s.n. (LD); Stock
AARGAU: Aarau,
1881, Tavel s.n. (Z).-FRBOURG:
s.n. (G).-GLARUS:
s.n. (Z); Landquart,
Neumann
near Bern,
Belpmoos
Skon,
Schachen
s.n. (Z); Umiker
Johansson
s.n. (LD).-VARMLAND:
Switzerland.
(LD).
s.n. (Z).-APPENZELL:
1907, Beauverd
(Z).-GENEVE:
s.n.
Elmqvist
s.n. (LD).
1898, Stackelberg
1917,
(LD).-VASTERNORRLAND:
1910, Zehnder
s.n. (Z); Dottingen,
Zofingen,
Chur, Heer
1876,
Alvsjo,
1853, Gothe
Norrsunda,
s.n.
Hilphers
Nasby,
s.n. (LD); Skonberga,
s.n. (LD).-STOCKHOLM:
1883, Sederholm
1885, Thedenius
1898, Hiilphers
1908, Zurcher
1949, Lundevall
s.n. (LD); Norrkoping,
1867, Westerlund
SODERMANLAND: Kila,
VOLUME 50
1963, Lousley
Webster
8323
s.n.
(K). Wales.
BuRRows:Pembrey,Carmarthen,1899,Marshall s.n. (BM).-GLAMORGAN:
Cardiff, 1906,Gregor s.n. (MA).
MERIONETH: Aberdovey,
CRNAGORA: Ulcinj,
346
1875, Fox
1973, Sverepova
(WU). Zimbabwe.
Salisbury,
s.n. (BM).-PEMBROKE:
Tenby Burrows,
s.n. (PRC); Donji Milanovac,
Gordon
245147
herb. Fischer
(GOET)
(as 0. gauroides);
treuse (hort. Carthusia Majoris),
(as Onagra
manni),
Aug
1829, Bauer
linkiana),
1844
1836
(DS)
1758, Biond.
(Fl, herb. Webb)
(as 0. media).
s.n. (CORD);
Erlangen,
1831,
(as Onagra
Berlin,
1781, Schreber
Wien,
herb. Fischer
37 (DS); Paris,
Germany.
s.c. s.n. (BM). Yugoslavia.
(M); Predejana
near Vranje, Niceic
(K).
SPECIMENS CULTIVATED IN BOTANICAL GARDENS. Austria.
1810,
1867,
57067
Bleeie
(W). Denmark.
(as 0.
1834, Weinkauff
media),
1826
1849
(GOET)
s.n. (M), 1836,
1839, herb. Fischer
(B, destroyed;
s.n. (M); Frankfurt,
serotina).
photo
1823
Char
(Fl, herb. Webb)
(GOET)
at MO)
Copenhagen,
France.
(as Onagra
(as 0.
(FR); Hamburg,
sick
spectabilis),
1834
(GOET)
(as 0.
Pirna, Abendrothe,
comeniana);
(as 0. gauroides).
(GOET)
from Rostanski
0.
197
OENOTHERA
1997
1840, Besser
1975). Kiyev,
1836
Zurich,
10. Oenothera
& Coveny
1983 no. 1436,
1981 no. 1657,
Gard.
Liege
1975
Gard.
Paris
1975,
cult. DUSS-82-232
83-141
1822
(MA).
cult. DUSS-77-0348
(010
cult. DUSS-78-0161
(MO)
(012
cult. DUSS-77-0446
no. 459,
1974
Sem. Bot. Gard. Barcelona
cult. DUSS-77-0441
Gard. Basel
(MO)
cult. DUSS-77-0447
1975
1979 no. 1798,
and 19).-SOMME:
Ind. Sem. Bot.
Pisa
Ind. Sem. Bot. Gard.
Lido,
and 21i). Japan. HONSHU: Miyagi
Pref., Mt. Fubo, Boufford &
and 111). Portugal.
COIMBRA: Ind. Sem. Bot. Gard. Coimbra
cult. DUSS-82-406
and
(012
7, cult. DUSS-81-600
de la Selva,
Ind.
of 0. Renner,
and 111). Sweden. Collection
BASEL: Pratteln, Ind.
Switzerland.
Schweden").
(012
Ind. Sem. Bot.
450 m,
Senzach,
(MO).-ZURICH:
and 11) U.S.A. CALIFORNIA: Del Norte Co., Smith
(MO) (012 and 111).Humboldt Co., Arcata Bottoms near
(012
749, cult. DUSS-76-045,
748,
Ind. Sem. Bot. Gard.
de Gaia,
TARRAGONA: Macanet
III). Spain.
(MO)
(MO)
747, p.p.,
Vila Nova
and 111).-PORTO:
(012
(MO)
and 111) ("O. R-r-lamarckiana
cult. DUSS-77-0338
& Ackermann
Ind. Sem. Bot.
Etaples,
La Molliere,
BRANDENBURG: Berlin, Lichterfelde,
III).Germany.
Ind. Sem. Bot. Gard. Leipzig 1974 no. 194,
Leipzig,
no. 1472,
101, Stubbe
along Hwy
River, Montalvo
and
(012
cult. DUSS-78-0158
1977,
(MO) (012
Sem. Bot. Gard. Basel
and 111).-PAS-DE-CALAIS:
(012
(MO).-SACHSEN:
(MO)
19864,
(MO)
(MO)
no. 171, cult. DUSS-83-0143
no. 1564,
Ind. Sem. Bot. Gard. Copenhagen
Italy. REGION TUSCANY. Prov. Pisa: Migliarino
(MO).
near Jind
UBC Cam
BRITISH COLUMBIA: Vancouver,
HOLBAK: Kundby,
(MO) (012
cult. DUSS-77-0445
42, cult. DUSS-84-254
cult. DUSS-77-0442
River
(MA),
(cited
Ind. Sem. Bot.
and 111).France. C6TE-D'OR: Soisson-sur-Nacey,
Ind. Sem. Bot.
77-0344
(MO) (012 and 11).-HAUTE-SAVOIE:
270 m, Ind. Sem. Bot. Gard.
Bartenheim,
(MO).-HAUT-RHIN:
(MO) (012
cult. DUSS-82-451,
no 3088,
Nowak-Krawietz
Mad
1801
(MA),
NEW SOUTH WALES: Pinch River
(MO). Canada.
(MO). Denmark.
cult. DUSS-77-0343,
1975 no. 1974,
Gard. Geneva
1984
Salzburg
cult. DUSS-82-0405
1502,
Basel
Porto
(LE) (as 0. media;
SALZBURG: Lehen,
(MO) (0)12 and 1 ,). Austria.
WEST-VLAANDEREN:
no. 1050, DUSS-88-2012
(MO). Belgium.
s.n., cult. DUSS-83-0161
1975 no. 794, cult. DUSS-77-0353
1974
1835
79-0637
cult. DUSS-77-0448,
2757,
1982, Wasmund
Gard. Dijon
Wood
(LE). Russia
(Z).
Ind. Sem. Bot. Gard.
pus, Straley
1974
1798
(LE). Spain. Madrid,
herb. Fischer
1832,
1834
glazioviana.
abyne, Pickard
460 m,
(Warsaw),
(h.b. Petropolitani),
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. Australia.
Ostende,
Leiden,
Netherlands.
1975). Warszawa
s.n. (KW); Leningrad
1820, herb. Stephanianum
Moscow,
suaveolens);
Switzerland.
s.n. (CORD).
1819, Bauer
(cited from Rostaniski
Poland
76-046,
76-049,
76-047,
76-050
(MO).
and 111).Mendocino
Co., Fort Bragg along Hwy 1, 1975, Hoch s.n., cult. DUSS-76-044
(MO) (012 and III).Douglas
OREGON: Curry Co., Nesika Beach along Hwy 101, Stubbe 6, cult. DUSS-81-599
Snohomish
Co., Monroe,
Stubbe 5, cult. DUSS-81-598
(MO) (012 and II,).-WASHINGTON:
Co., Reedsport,
(MO) (0 12 and 111).
Wagner 4544, cult. DUSS-82-401
(MO) (0)12 and 1II), DUSS-82-404
CULTIVATED STRAINS EXAMINED, BUTWITHOUT VOUCHERS. Chile. Province unknown: Casa Pangue, cult.
(MO)
(012
DUSS-74-035
ZEELAND: Domburg,
and 111).Netherlands.
(012
(M); Khanabad,
(NA); Kabul,
Munz
Agronomfa,
363;
(NSW);
Jindabyne
s.n. (NSW);
Johnston
Dist.,
Peuguin,
Pedley
Raven
1971, Meares
(PERTH);
Ferry,
1165
& Engelhorn
(NSW).-WESTERN
15 mi NNE
Ibm, Oberwinkler
Royce
8506
1985, Polatschek
(M).-TIROL:
Snowy Mtns,
W
(K, NSW,
Gulch
s.n. (W).-NIEDEROSTERREICH:
at River Ybbs,
at River
1973, Schulz
Inn, 1936,
Merkersdorf,
AUSTRALIA: Largs
Sulphur Creek
1967, Henshall
239
1983
1951,
191
Dar
Bay,
near
s.n.
(NY); Bridgetown,
Beach
Rd, Green
(PERTH). Austria.
1878, Oborny
s.n. (B).-OBEROSTERREICH:
Schneider
s.n.
1962, Evans
ca. 1000 m,
Isle, Balcatta
Park, Scrymgeour
1880,
Park, Raven
RSA).-QUEENSLAND:
Anway
de
s.n. (NSW); Weaver's,
near Orbost,
s.n. (PERTH); Osbume
National
River,
(CHR, NSW);
25998
coast of SW Australia,
Yanchep
Pitt Water,
1959, Burgers
751
Gilbert's
1948, Hamburg
Brixlegg
Thredbo
National
1575 m, Thompson
Chase,
s.n. (NSW).-SOUTH
Raven & Engelhorn
Quinta Wessel,
Innes, 1949, Noonan
Creek,
s.n.
114 (POM), Calder6n
to Kosciusko
Ku-Ring-Gai
Coveny
Gelsii
11572
1937, Koelz
of the Facultad
Garden
(Z); Glen
Park, Wragger
1966, Everist
(CHR).-VICTORIA:
s.n. (WU); Waidhofen
5161
entrance
Sydney,
of Windsor,
Somerset,
(PERTH);
1669
s.n. (NSW); Wentworth,
(K); Stanthorpe,
25982
National
Sharif,
del Unruguay,
Concepci6n
Kaspiew
s.n. (NSW);
1962, Mort
"Las Chacras,"
Jindabyne,
(NSW);
AUSTRALIA: SW of Busselton,
Pinjarra,
1902, Teyber
2754
1951, Clerk
s.n. (PERTH); Manjimup,
KARNTEN: Gmiind,
Retz,
near Bathurst,
Tumut Dist.,
(MEL);
Kosciusko
NSW);
s.n. (RSA).-TASMANIA:
1949, Cleland
3291
& Coveny
S of NSW Wiseman's
ling Downs
River, Muir
Pickard
Perthville
Rios:
(POM).-ENTRE
(CHR, MO,
Creek,
s.n. (NSW);
9977
del Plata, near Arroya
NEW SOUTH WALES: Armidale,
at Snowy
25804
(NSW); Wilson
379
Parodi
s.n. (PH). Australia.
& Engelhorn
6 mi
(GH, POM); Mar
s.n. (KYO); Mazar-e
BUENOS AIRES: spont., Botanical
(NA, US). Argentina.
15455
Pergamino,
(BAA);
Lorentz
13494
Koelz
and
III).
2400 m, Podlech
(012
Paghman,
Mtns,
1955, Kitamura
s.n. (NA); Kundury,
1939, Harlan
cult. DUSS-74-031
KABUL: Paghman
REPRESENTATIVE SPECIMENS. Afghanistan.
s.n. (W); Landeck,
s.n. (PRC);
Innviertel,
1982,
Po
SYSTEMATICBOTANYMONOGRAPHS
198
s.n. (W); V6ls,
latschek
Klause,
1970,
s.n.
latschek
s.n. (BR).-BRABANT:
(BR); Kessel-Lo,
Pelgrins
gleur, Lawalrie
16675
1822
Grama, Mexia
4232
PAULO: Ubatuba,
11700
34 (UBC);
Saturna
(SP). Bulgaria.
ley 1519
House,
(UBC).-MANITOBA:
Annapolis
Burgess
Royal,
St. Laurent, Montreal,
(MO). China.
(C, G, K, PE, UC).-JIANGSU:
ard 2499
Chen
80-72
1984
(PE). Colombia.
(NA). Czech
between
Vy?kow,
Hancand
Jaroslav,
Arhus,
delsslagteriet,
1968, Lorenzen
206
s.n. (C).-SOR0:
s.n. (C).-VEJLE:
LOJA: Catacocha,
(P).-ALPES-MARITIMES:
BASSES-PYRENEES:
3887
3069
(BR).-HAuT-RHIN:
Merl0se,
Vonsild,
Grasse,
SAoNE-ET-LoIRE:
s.n.
Merxmuller
10428
1895, Brachet
125
Alderlyst,
s.n.
(G).-SEINE-ET-OISE:
(C).-RIBE:
Augusten
1978, Holst
Hibon
(Z).-GIRONDE:
s.n.
Les-Essarts-le-Roi,
La Fonte-sur-Mer,
Gerrinck
1916,
Foucaud
Alleizette
Geerinck
p.p.
(Z).
Chatelaillon,
Hourtin,
SAVOIE: Lac Dunas,
1881,
s.n. (C).
1460-2,
1893, Hausser
Mimbaste,
K0bn
AEr0 Island, Aroskobing,
Barr,
6117, p.p.
s.n.
An
UNCC).-LOL
(HBG).-CHARENTE-MARITIME:
(M).-LANDES:
s.n. (LY).-VENDEE:
SOM, UBC,
AISNE: St. Quentin,
s.n. (DAO).-HAUTE
68/54
1971, Kaae
M0lle,
1959, Hansen
(DAO).-BAs-RHIN:
Desplantes
1962, Gavelle
Charpin
Macon,
Gavelle
Fl, HBG, M, MA, MO,
s.n. (C).-HJ0RRING:
s.n. (C).-SVENBORG:
France.
9339
(PR).-JIHOMORAVSKY:
s.n. (C).-S0NDERBORG:
s.n. (C).-VIBORG:
(RSA).
Semur-en-Auxois,
Noyant,
VAR: La Garde-Freinet,
622
and Biarritz, Walther
Bayonne
Habsheim,
(G).-INDRE-ET-LOIRE:
Jensen
1961,
& Fosberg
s.n. (C).-K0BNHAVN:
RM,
Assens,
1983, Svennimgsen
1962, Christiansen
1957,
(BR, L).-C6TE-D'OR:
(C).-ODENSE:
Nissum,
2050 m, Espinosa
between
Rallet
s.n.
GA, M, MO,
2000 m,
Kunming,
2100 m, Grant
1964, Hansen
Kalundborg,
Lu Shan, Stew
Kuling,
(PE).-YUNNAN:
1963, Pedersen
Lago
Steward
Kikungshan,
ALBORG: Marens
Genner,
(C, BM, FLAS,
1967, Hansen
s.c. s.n. (C).-RINGK0BING:
1965, Hansen
1963, Hansen
& Svendsen
Island, Grusgrur,
1967,
Ecuador.
s.n. (C).-HOLBnEK:
Jacobsen
team 1438
(PR). Denmark.
1962, Luithen s.n. (C).-HADERSLEV:
havn, Orstedverket,
borg,
Jehll'k 6669
1883,
s.c. s.n. (HBG).-OSORNO:
5 km S Gachala,
San Isidro,
Hillsdale
near London,
(PE).-HENAN:
s.n. (KYO).-JIANGXI:
survey
Stra
of Sechelt,
SCOTIA: opposite
Jehlik 6236
(Budweis),
Hradynear Budejovice
Unar & Unarova
1542 (A, B, BC, BR, C, DS,
Chalkovice,
(C).-ARHUS:
LAND: Falster
Jinfushan
CUNDINAMARCA:
P, US, WIS).-VYCHODCESKY':
Laeborg,
Co.,
JIHCESKY:Nove
Republic.
1932, Kimura
Shang-hai,
Nanchuan
(K, UC).-SICHUAN:
Canada.
1893, Scott s.n. (TRT).-QUEBEC:
1902,
75333
de
s.n. (DAO); Vancouver
PH, US).-ONTARIO:
ACONCAGUA: Jahuel,
ANHUI: Xie Co., Guan
to Fazenda
s.n. (V); W
s.n. (DAO).-NOvA
(CAN, GH, MICH,
and
(BR,
(C, NY, P).-SAo
s.n. (SOM).
1930, Groh
s.n. (CAN, TRT); Casselman,
1883, Burgess
Gent
13112
s.n. (V); 2 mi E of Kelowa,
1952, Melbourn
Savona,
1929, Neatby
s.c. s.n. (MT). Chile.
1891,
Schmid
Beach,
et al. 24212
Fernald
s.n. (CAN); Woodstock,
Llanquihue,
9676
Victoria
(DAO);
8343
1971, Kuzmanov
An
PARANA: Curitiba,
Araponga
Glaziou
1959, Ashlee
Harbour,
s.n. (UBC); Courtenay,
6740
Dist.,
Villa Theresa,
SOFIYA: Pavlovo,
1951, Edgar
Park, Hainault
between
Lawalree
s.n. (BR). Brazil.
1947, Vise
s.n.
s.n. (BR).-LIEGE:
(BR).-OOST-VLAANDEREN:
GERAIS: Carangola
Island, head of Fulford
Island,
1mi S of Goldstream
Island,
(BR); Ostende,
(US).-MINAs
s.n.
(Anvers),
1909, Michiels
(BR); Heverlee,
1930, Culot
St. Idesbald near Koksyde,
DE JANEIRO: Petr6polis,
(F, GH, UC).-RiO
Loefgren
8033
Lawalree
1066
& Joly 373
BRITISH COLUMBIA: Salt Spring
Brayshaw
Genk,
(BR).-WEST-VLAANDEREN:
1963, Loots
Moreira
1591
Montigny-sur-Sambre,
Po
Dombirn,
1897, Arbesser
Donau,
ANTWERPEN: Antwerpen
Belgium.
Geerinck
Brussel-Auderghem,
(BR).-LIMBURG:
Nieuwpoort,
Jard. Fac. Farmacia,
erythrosepala).
Gleichenberger
(GZU).-VORARLBERG:
bank of River
s.n. (W).-WIEN:
(BR).-HAINANT:
2528 & 2829
Robbrecht
s.n.
1986, Melzer
under 0.
1975,
s.n. (WU).-STEIERMARK:
1897, Handel-Mazetti
Polatschek
1971,
(see Rostaniski
1967, Rompaey
UC, UNCC);
Innsbruck,
(W); GroBwilfersdorf,
(W); Feldkirch,
(GZU). Belarus
Zelzate,
6 km W
s.n.
Seipka
VOLUME 50
s.c. s.n.
(LY).
s.n.
568
1942, p.p.
(BR, P).
(BR). Geor
gia (see Rostatiski 1975, under 0. erythrosepala).Germany. BADEN-WURTrEMBERG:
Schopfheim, s.c. s.n.
(NY).-BAYERN:
Berlin,
Neustadter
(HBG).-SACHSEN:
Nerlich
Leipzig,
WRSL).-SAcHSEN-ANHALT:
NISiA: Pilion Mtns,
(AMD).-FEJtR:
Halle,
Nagylang,
s.n. (BP).-SOMOGY:
(BM); Malashahi
Iraq. Baghdad,
Picentini,
Caresti
Sahira
Moraldo
1685 m, Dar
1061
1967, Koller
3331
(MO).-UTTAR
Carpesica,
between
PIEDMONT. Prov. Torino:
Fra Carisio
1896, Pampanini
and Balocco,
s.n. (FI). Japan.
between
1978, Abbd
Valley,
MAG
s.n.
1955, Csapody
s.n. (BP). India.
Ludlow
Avellino
Boccioleto
(LZ,
1906, Richter
PRADESH: Naini, Duthie
between
s.n.
s.n.
s.n. (LZ). Greece.
Tatabanya,
Desu
Klimmek
1967, Gutte
1910, Gayer
Celldomolk,
(K); Rajparan,
s.n. (HBG).
1948,
BUDAPEST: Klozsvar,
s.n. (BP).-KoMAROM:
s.n. (BP).-VAS:
Stewart
near Leer,
s.n. (LZ); Wiederitzsch,
(G). Hungary.
& Kummerle
(M).-BRANDENBURG:
s.n. (GOET).-BREMEN:
1958, Haase
Fuhlsbuittel,
Ihrhove
s.n. (B); Weissenfels,
s.n.
1962, Kdrolyi
et al. 69 (FI).-REGION
Cervada
1965 Gutte
13315
1879, Heiland
(K). Italy. REGION CAMPANIA. Prov. Avellino:
s.n. (TO). Prov. Vercelli:
Prov. Venezia:
Topali
near Srinagar,
Bag Gandorbal,
235
Street,
1923, Filarszky
Balatonbereny,
Oderberg,
(FR).-NIEDERSACHSEN:
1920, Bernau
1973,
Tsangaradha,
Leuze & Doppelbauer
s.n. (BREM).-HAMBURG:
63406
Semmelweiss
JAMMU AND KASHMIR: Dal Lake
8208
& Jehlik
1980, Kuhbier
Homburg,
s.n. (M); Ulm,
42 (BR, C, LD, MA);
Nowak-Krawietz
Hafen,
HESSEN: Bad
1969, Dorr
Kaufbeuren,
Lichterfelde,
& Sherriff
4032
(BM).
and Serino, Monti
and Valsesia,
s.n. (TO).-REGION
HOKKAIDO: Pref. Unk.,
1889,
VENETO.
South-Hokkaido,
1997
OENOTHERA
1884, Brooks
(UC). HONSHU: Chiba
s.n.
Kokuyurin,
1966,
Daigo-machi,
Okamoto
Mt. Yamizo-san,
et al. 5903
Murate
s.n. (KYO);
Shiga
633, p.p.
stroom
18363
Dietrich
6679
(L).-NoORD-HOLLAND:
cash: Huaraz,
57/21
Healy
53/362
Degener
BRA: Beira Litoral, Choupal,
1953, Matos
hamstown,
Belmont
Serfontein
Bridge,
Winterton
1330
Kwangnung:
Uribe-Echebaria
Strey 3870
s.n. (MA).-CIUDAD
Ladero
s.n. (LD).-GOTEBORG
s.n. (LD); Kiivlinge,
s.n. (LD).-OLAND:
(Wallis): Viege,
1908, Hohn
head,
gen
1905, Green
s.n.
m,
at Lake Zurich,
J.L. Wang
1947, Blom
1882,
Peng
8348
et al. 3588
(TAI).
s.n. (BM).-DERBY:
Cheltenham,
Poole,
1951,
1927,
Grevithick
Townsend
s.n.
s.n.
s.n. (G).-LA
RIOJA: Man
s.n. (MA).-MADRID:
Sierra del Eje,
del Ebro,
Ihnisia.
HSIEN: Ssuyuan,
Medinine,
Louis-Arsene
s.n. (M).-DEVON:
(K).-ESSEX:
(K).-HAMPSHIRE:
s.n. (Z); Re
1916, Frymann
710 Truck Rd
Braunton,
Hawkwell,
von
1913, Jakob
s.n.
(Nan
s.n. (G). United
Birken
(BM).-CHESHIRE:
Brockenhurst,
1979,
1967,
(BR).-VALAIS
1912, Cuenod
6627
1935,
Barkakra,
Ndgeli
s.n. (Z); Wil,
s.n.
Kummeln,
16628
Lawalre'e
s.n. (Z); Greifensee,
Mon
1974, Balade
AARGAU: Biinzen,
1918, Thellung
1960, Ball
Bakewell,
Fuenteheridos,
s.n. (LD).-MALMOHUS:
(MO).-TAICHUNG
s.n.
1980,
1904, An
Figueras,
1908, Traigairdh s.n. (LD); Skanor,
s.n. (LD). Switzerland.
station Tiefenbrunnen,
1982,
Sevillano
San Vicente,
BLEKINGE: Augerum,
4, 1914, Thellung
South
s.n. (MA).-SALAMANCA:
Itsher s.n. (Z); Rapperswil,
Zurich
PRE).
s.n. (ARAN).-ORENSE:
Delta
(K);
3673
del Ebro,
1970,
rivers, Zubia
Txalet,
s.n. (LD); Malmo,
CHANNEL ISLANDS: Jersey, Saint-Sauveur,
(BM).-DORSET:
GLOUCESTER:
1983, Cataldn
s.n. (MA). Sweden.
1951, Olsson
s.n. (G).-ZURICH:
Ramblas
(BM).-GERONA:
1974, Lainz
(K, PRE);
(BR, K, PRE).
(K, NY,
s.n. (ARAN).-HUELVA:
1969, Valdes-Bernejo
OCH BOHUS: Goteborg,
s.n. (Z); railway
1900-2350
England.
Lesaca,
(MA).-TARRAGONA:
1949, Lange
ILAN HSIEN: Szuyuanyakoa,
(Z). Taiwan.
hu-ta-shan),
Hurden
Thomas
195
& Cataldn
CAPE
Z); Gra
& Vahrmeyer
(W); Mieres,
of Iregua and Ebro
Portonovo,
Borgholm,
s.n.
s.n.
5841
Strey 9528
7445
s.n.
ALAVA: Elciego,
NA,
Theron 598
1700 m, Killick
Turner
Serra
Africa.
(NH); Harrismith,
1940,
CORUNA: Puentedeume,
confluence
1050 m, 1992, Aldosoro
Thorvinger
& Petty
Leadley
1990, Aizpum
et al. s.n.
South
(BR).
(BR, HBG, MO,
Park, Liebenberg
Polatschek
1977,
s.n. (PO).-LIS
1984,
1958, Michel
(BC, BM, G, LD, LISU, MA).-CANTABRIA:
Fuentearriba,
Rundkwist
Kingdom.
5066
s.n. (MAF).-NAVARRA:
Reguejo,
s.n. (Z).-SCHWYZ:
2671
Avil6s,
s.n. (MA, SEV).-LA
1981,
(BC).-ZAMORA:
gensdorf,
(F). Spain.
s.n. (MA).-PONTEVEDRA:
del Rio,
Gate Highland
In-Cho
Chung
462
Umkomaas,
Buffelspoort,
s.n. (MA).-LOGRONO:
1982, Gonzdles
1991, Aldosoro
Bothmer
Golden
REAL: Sierra Morena,
& Silvestre
silla, 1935, Cdmara
Upper
& Castro
1140 (PRE); Knysna,
King
(K); Rustenburg,
Gonzalo
s.n. (LY).-GUIPUZCOA:
Johnson
(LD). Portugal.
92 (GH, US).-COIM
Rubona,
p.p.
8487
(CHR).
92 (ISL). Peru. An
s.n. (PO); Port6,
4540,
Bayliss
Babanango,
Impendhle
s.n. (ARAN).-ASTURIAS:
Manlleu,
1974, Auquier
(GRA); Hogsback,
(K, NH);
Kyonggi-Do,
(MA).-BARCELONA:
temay6r
489
1944, Rozeira
s.n. (PO). Rwanda.
1118 (PRE); Blaaukranz,
FREE STATE: Bethlehem,
(K, PRE).-ORANGE
Madrid,
1870 m,
108 (C), Brown
77/36
Healy
122/89
s.c. 1879
(Bauerwitz),
1967, Costa
Carneiro
(K, PRE).-NATAL:
Strey 2528
TRANSVAAL: Tygerpoort,
1980, Rivera
1945, Barros
River,
Sykes
Chinar bagh, Maqsordand
Nazare,
Arrabida,
1988,
(CHR).-CANTERBURY:
Taylor
Alexandria,
Hansen
(MO); Sao Miguel,
to Uganda,
Valley, Britten
Werger
Gilgit:
(L).
1899, Lin
s.n. (L); Rotterdam,
Sykes 545/81
OPOLE: Baborow
s.n. (UT).-LEIRIA:
Johnson
Forest,
Settlement,
36506
at Rio Ave.,
near border
(BR); Ruhengeri,
PROVINCE: Alexandria
(P). Poland.
6334
s.n. (B); Vlissingen,
Blenheim,
(CHR).-OTAGO:
s.n. (LISU).-P6RTO:
1916, Coutinho
(PO); Santo Tirso, Lousado
855
(CHR). Pakistan.
Hu
AZORES: Ponta do Varadoura,
BOA: Lisboa,
Bay, near Mangonui,
(CHR).-MARLBOROUGH:
Brownlie
Healy
1932, Koster
(Den Haag),
s.n. (L).-LIM
s.n. (L); Petten, Oost
von Ruynen
Haaksbergen,
1960, Buchheim
Shi
Lesotho.
78 (BM); Klebang,
Paude
1898, Bedeke
Hilversum,
s.n. (TI);
(KYO).
Veth & Koopmans
1958,
Domburg,
Ootamigama,
(TUS); Yamanashi
16027
(BM); Katmandu,
(L).-OVERIJSSEL:
'sGravenhage
m, Proaash:
2600-2650
18307
s.n. (L).-ZEELAND:
Collingwood,
404
Ilam, Williams
1960, Koyama
Pref., Demizu-cho,
Yamamoto
Houjyo-cho,
(KYO); Nagano
1930, Nakajima
Oka 45854
Kumamoto
s.n. (TI).-KYUSHU:
19864
Garden
Tottori Pref., Mt. Dai Sen
Nishimurou-gun,
Pref., Abu-gun,
GELDERLAND: Apeldoorn,
AUCKLAND: Doubtless
Zealand.
WELLINGTON: Greytown,
gustin
1947, Ono
Pref., Botanic
Pref., Nikko
Tochigi
m,
1100-1400
Pref., Sanekawa,
s.n. (BR, CAS);
Pref., Tanabe-cho,
Pref., Befu-mura,
Ooststroom
near Rangiora,
River
1936, Makino
s.n. (KYO); Yamaguchi
(PRE, SAM). Nepal.
1976, Wolters
(CHR).-NELSON:
Rico
(KYO);
Nerima-Ku,
(M).-ZUID-HOLLAND:
den s.n. (L). New
Korea.
5607
Ehime
(L); Zandvoort,
4711
Ashley
Hashimoto
Ibaraki Pref.,
near Odagoe,
& Wood
Boufford
Pref., Mikiyama,
(KYO);
(CAS); Kyoto
(TI); Niigata
tsuru-gun,
Balhuizen
UTRECHT: Maarssen,
Pref., Mt. Fubo-san,
Hyogo
6991
Mtns
1030
et al. s.n. (K). Netherlands.
BURG: St. Pietersberg,
s.n.
1931, Makino
(KYO);
Hosomi
700 m, Midorikawa
Pref., Ooizumi,
Minami
s.n.
Taki-gum,
Iwate Pref., Hayachine
Fujisawa,
Higashiura,
(KYO).-SHIKOKU:
1963, Hara
(TUS);
s.n. (KYO); Miyagi
1962, Tanaka
Pref., Yamanakakohan,
Leribe, Dieterlen
Jyotou-cho,
s.n. (KYO); Wakayama
Pref., Kinpo,
12019
Pref.,
Pref., Otsu-shi,
1978, Terabayachi
mada
466
Lake Nojiriko,
1931, Ito s.n. (TI); Tokyo
Oki,
Sougi,
Nagayama
1924, Okeo
Pref., Kamiminochi-gun,
1961, Murata
Pref., Asahi,
(KYO);
(TI); Kanagawa
Kyoto University,
Yamagata
s.n.
199
1931, Meinertsha
1933,
Vine
1933,
s.n.
Hanbury
(K).
s.n.
SYSTEMATICBOTANYMONOGRAPHS
200
(BM).-HERTFORD:
Dony
Hitchin,
s.n. (BM).-KENT:
wick
82/52710
(DAO);
SEX: Twickenham,
Thames,
1935,
s.n.
Chapple
430819B
(BM, K); Weybridge,
Ross
cle in western
Little Rock,
14314: Munz
cult.
from
Golden
14694
Hawaii
Co., Nonquitt,
Bristol
shire Co., Amherst,
Co., Bath,
1880, Bailey
ton s.n. (RSA).-NEw
land,
Moldenke
s.n. (NEBC).
s.n.
8593
(GH).-NEW
sap Co., Bremerton,
14510: Munz
14697
Monongalia
Co.,
13th Ave.
Wight
entrance
Hot Springs,
Uttal
(BH, F, POM), Munz
Co., Morgantown,
Snyder Co.,
& Muenscher
1942, Ammons
10633
&
S of Johnson Creek
on
206
(WTU). Kit
s.n. (WS), Suksdorf
(DS, POM, WTU),
Dane
IS
Seymour
5868
cult. from Munz
15243
(BH, CAS, DS,
20, Woodland
1300
(CU, DS, GH, WTU).-WEST
s.n. (WVA).-WISCONSIN:
Isle of
s.n. (NY).-RHODE
3.6 mi
14765
Lake, Maguire
jct. of Hwy
(VPI). Lane Co.,
Co., Fletcher,
1912, Suksdorf
Co.,
s.n. (CU).
Co.,
10580
1913, Carter
on Hwy
Somerset
1924, Wiegand
(BH, CAS, GH, NY, POM, US), Munz
2 mi E of Sedro Woolley
1956, Den
s.n. (CU). Mon
Clackamas
14510
Panghorn
(NA).
s.n. (ORE).-PENNSYLVANIA:
Co., Bingen,
Co.,
Clinton
Coos Co., Northumber
Munz
Skagit Co.,
s.n. (GH). Hamp
Island Co., near San Juan de Fuca, McElvaine
s.n. (NY). Klickitat
(NEBC).
s.n. (GH). Norfolk
2155
Franklin
York
(NEBC).
8889
1882, Kilborne
College,
inlet, Seaside,
s.n. (NEBC).-VERMONT:
Cum
s.n. (NEBC).
s.n. (MO).-MICHIGAN:
to Fort Lewis,
GH, NY, POM, UC, US, WTU).
NY). Whatcom
1906, Williams
1880, Faxon
Clark Co., 7.9 mi N of Rock Creek,
703 (IND, MICH).
3 mi E of main
Co.,
Tower
s.n. (NA).-OREGON:
1955, Bierly
1884, Leland
1954, Monschino
(WS). Pierce
Co.,
s.n. (F).-MAINE:
627403
Co., Moravia,
(OKLA). York Co., near Pleasant Grove,
(MO).-WASHINGTON:
& Grayson
99, Bartlett
Hwy
Rutherford,
Co., bank of Veterinary
1910, Crayton
Co., Newport,
3222
Co.,
& De
2 (F).-INDI
1894, Furbish
s.n. (OKLA).
1910, Cheever
YORK: Cayuga
(MO). Clatsop
Co., Providence,
22587
Countryman
4465
(DS). Lincoln
Moldenke
LAND: Providence
Co., Biltmore,
garden, Degener
12 mi SW near Bozeman
Co.,
Jaffrey,
(UC). Santa
1909, Woodward
Wheeler
1896, Churchill
Co.,
Arboretum,
4156
Provincetown,
Co., Georgetown,
Riverside
Stybing
Park, Gardner
1886, Rose
Co., Bethel,
Co.,
cult. from Munz
Jepson
in Degener's
Farmington,
Co., Cambridge,
JERSEY: Essex
s.n. (CU). Tompkins
Wagner
344
Nelson
Gallatin
(BH, ND, NY).-NEw
1909, Arnold
26 & rd toWelches,
Eugene,
Oxford
Co., Dorchester,
Co.,
Co.,
Barnstable
s.n. (MIN). Essex
Suffolk
[cult.]
(BH,
1907,
Co., Angush,
cult. in Lincoln
Co., Liberty,
Franklin
s.n. (NEBC).
s.n. (BH).-MONTANA:
NORTH CAROLINA: Buncombe
Que, Selinsgrove,
s.n. (NEBC).
HAMPSHIRE: Cheshire
1906, Williams
roe Co., Greece,
(MICH). Union
6774
Creek,
Cir
(NA). Hum
14696
POM),
Co.,
(BM).
Lake Park,
IND, NY, POM).
London
New
Sts., Volcano,
(MT).-MASSACHUSETTS:
1889, Sturtevant
1884, Fuller
Co., Purissima
Co., Chicago,
1948, Torrey s.n. (MASS). Middlesex
Needham,
Watchung,
Cook
1873, Schribner
3079
Cleonique
Deam
1898, Furbish
Co., Manchester,
Co., Alfred,
San Mateo
s.n.
1834
West
San Francisco
(RSA).
1971, s.c. s.n.
14384: Munz
(BH, CAS,
F, GH,
(POM).
(MICH).-CONNECTICUT:
Co., Tiwi & Kilauea
Co., Wollcottville,
Co., Brunswick,
14080
Munz
16316
Co., Overlook
Wight
cult. from Munz
(BH, CAS,
Wilmott
Lewis
1943,
Co., Berkeley,
14314
Bay, Munz
Raven
Pulaski
Co., Los Angeles,
Los Angeles
15238
s.n. (CAS, RSA).
Smith 2320
(BR, C, CHR, W, Z).-ILLINOIS:
ANA: Lagrange
Kennebec
from Riverside,
1957, Howell
Alameda
(CAS, GH, POM, UC, US),
0.5 mi N of Anchor
Co.,
Creek,
s.n. (NEBC).-HAwAII:
berland
14384
Tintern,
on
Henley
FLINT: Rhuddlan,
(MO).-ARKANSAS:
212 (MO).-CALIFORNIA:
(CAS, NY, POM, UC), Munz
Gate Park,
667
& Be
(BM).-MIDDLE
(BM).-SURREY:
ca. 7 mi N of Hereford,
(BM).-MONMOUTH:
(CAS, NY, POM, UC, US).
seed of Reedgarden
34678
s.n.
near Spring Hill, Graves
s.n.
Babington
s.n. (BM). Wales.
1978, Webster
Bannister
1951,
Shepard
14764
Cruz Co., Boulder
gener
s.n. (K); Dinmore,
Co.,
s.n. (NA). Mendocino
Wight
1924, Fraser
1871,
699
s.n. (CHR).-OXFORD:
Sykes
1948,
Rosser
Ainsdale,
on the Sea, Bailey
1950,
Cockfield,
7 mi E of Arcata, Munz
boldt Co.,
F, POM, UC), Munz
Brandon,
(BM).-SUFFOLK:
Kenfig,
ALABAMA: Barbour
s.n. (K).-LANCASHIRE:
s.n. (MA); St. Anne's
(DS).-NORFOLK:
AND CROMARTY: Barbaraville,
(BM).-GLAMORGAN:
AND PETERBOROUGH: Conington,
(BM).-HUNTINGDON
1915, Crossfield
1965, Valdei
16493
Scotland.
U.S.A.
on Sea,
Freshfield,
Raven
s.n.
1923, Little
Littlestone
VOLUME 50
Co., Madison,
(DS, MIN,
VIRGINIA:
of
University
Wisconsin Arboretum, iltis28457 (MO,WIS). OutagamieCo., towardsKaukauna,s.c. 16 (DUKE).Uruguay.
MONTEVIDEO:
Miguelito, 1874, Fruchard s.n. (P).Yugoslavia. CRNAGORA:
Cetinje (Tsettinie),1889, Sommier
s.n. (FI).
SPECIMENS CULTIVATED INBOTANICAL GARDENS. Brazil.
Sao Paulo,
Jard. da Comissao,
1896, Edwall
s.n.
(POM,SP); Bot. Garden Sao Paulo, 1902, L&fgren11898 (SP).Germany. Berlin, 1869 (HBG), 1874,Dumas
s.n. (GOET).
11.Oenothera argillicola.
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. U.S.A.
Crossing
81-620
0164
Co.,
State Park, Ind. Sem. Bowman's
(MO)
(MO)
(7II; 08
and 3II; 2 04
(7II); Fort Lewis
1979, Stubbe
Hill Wild Flower
and 311).-VIRGINIA:
along Rd 678,
s.n., cult. DUSS-81-587
1979, Stubbe
(MO)
(7II; 06
PENNSYLVANIA: Washington
Preserve,
1975,
Co., Washington
no. 61, cult. DUSS-77-0166
Bath Co., Williamsville,
s.n., cult. DUSS-81-588
Wurdack
(MO)
and 4II), DUSS-86/88-1012a.-WEST
(04
(MO),
s.n. cult. DUSS-77
and SII). Highland
VIRGINIA: Min
OENOTHERA
1997
201
1976, Brown s.n., cult. DUSS-77-0167
(MO) (711), 1976, Brown s.n., cult. DUSS-77-0168
(MO) (06
and 41), 1976, Brown s.n., cult. DUSS-77-0169
(MO) (04 and 51), 1976, Brown s.n., cult. DUSS-77-0170
(MO)
(04 and 511), 1976, Brown s.n., cult. DUSS-77-0172
(MO) (04 and 511; 06 and 411)' 1976, Brown s.n., cult.
(MO) (04 and II)' 1976, Brown s.n.,
DUSS-77-0173
(MO) (04 and 51), 1976, Brown s.n., cult. DUSS-77-0174
cult. DUSS-77-0175
(MO) (04 and 51), 1976, Brown s.n., cult. DUSS-77-0176
(MO) (2 04 and 311).
REPRESENTATIVE SPECIMENS. U.S.A. MARYLAND: Allegany
Co., Little Orleans, Downs 4600 (UNCC).
eral Co.,
Co., 0.4 mi NNW
PENNSYLVANIA: Bedford
s.n. (PENN).
burg, 1921, Ward
of Huntington,
s.n. (PH).-VIRGINIA:
Hunnewell
SMU). Botetourt
1.5 mi WNW
Co.,
s.n. (PENN). Highland
Erlanson
MT, NCSC,
densville,
NY, OKLA,
PENN,
roe Co., N of Sweet
et al. sn.
Springs,
(NY). Pendleton
Rowlesburg,
tonsville,
54
Steele
Clarkson
SMU,
TENN,
TEX,
MIN).
Mineral
Steele & Steele
Co.,
(FSU, KANU,
2667
(WVA).
1954, Clarkson
s.n. (WVA).
12. Oenothera
oakesiana.
328
220
along Hwy
US).
Co., near White
UNCC,
(GH, MIN,
8 mi
no. 3090,
cult. DUSS-77-0422
Co.,
ner, 1942)
("O. syrticola
ner, cult. DUSS-77-0399
(MO)
Renner,
cult. DUSS-77-0404
enburg
1982
burg 1976
1974
no. 410,
no. 271,
no. 241,
cult. DUSS-84-244
Acer Wood
to Lake Michigan,
Portland,
1980,
(MO)
Caughlan
et al. 4479,
Scoggan
12174
Scoggan
Fernald
(DAO);
AAAS Excursion
Mt. McKay
in Calvert
NY,
Ingonish
Beach,
35326
(CAN); Linden,
near New
149 (V); Willard
Twp., Baldwin
MT, TRT); Mammamattawa,
5030
Fernald
Gorham
Zinck
1941, Anderson
30 mi W
4096
River, Hudson
13355
3
of Cove,
1775
1190
(TRT);
Island, Smith
et
1896,
Squaw Bay Rd near jct. with
Bay, Bailey
(50?25'N,
1284
(DAO);
Port Moutou,
Klawe
Scatarie
Bay St.
& Long
Lake Erie, Point Abino,
2884
N end at Dawson
lowlands
Tusket,
Bay
to Bonne
St. George,
(GH, PH); Central
s.n. (TRT); Ft. William,
of Vermilion
entrance
Park, Bassett
(DAO); NW
St.
Island,
s.n. (GH, MT);
Island Natl.
(DAO);
(DAO).
(CAN); Campobello
PH); near southern
& Long 21991
3733
1137 (DAO); Moncton,
1931, Dove
40 (DAO).-ONTARIO:
Bay
River
(GH); Carleton,
Bay, Howe
et al. 451316
(GH, TRT); Lake Timiskaming,
Kenogami
67135
Conception
Topsail,
Breton
Cape
Schofield
Germany,
Lake,
tailrace, Stardom
Gates
(F, GH, MIN, MT, NY, PH, WIS);
PH, US);
s.n. (PENN); Meteghan,
s.n. (NY); Kashabowie,
Rd, Bailey
of 0.
Pepin Co., Chippewa
Park, Munro
Scoggan
(BH, GH, PENN,
et al. 1885
of Lawrencetown,
Pinehurst,
Natl.
(CAN); Chatham,
et al. 337
(BH, GH,
Co.,
Sisters
Port Elgin,
1933, Ayre s.n. (GH); Brigus,
(PH); W
Rousseau
Guysborough,
W
& Long
et al. 21996
al. 5226
5930
along Seven
River, Kouchibouguac
13260
Fernald
Brook,
& Wiegand
1910, Fernald
collection
(014).
s.n. (UBC, US);
Clarenville,
Island, Fernald
SCOTIA: Victoria
(GH).-NOVA
Digby,
(or Tweed)
of Wallace's
Fernald
Ren
of 0. Ren
collection
Standard"),
(MO) (014);
(MO)
MANITOBA:
4.8 mi S of Poutage
Parish,
(CAN); Bathurst,
Cay, near mouth
George,
("O. ammophila
176, cult. DUSS-82-0467
1900, Fowler
Smith 870 (US).-NEWFOUNDLAND:
of Islands, French
and Luckenwalde,
cult. DUSS-88
(MO) (014,
(012
cult. DUSS-82-0469
REPRESENTATIVE SPECIMENS. Canada.
St. Louis
3436,
cult. DUSS-77-0406
and 1I) ("O. syrticola Venedig").
U.S.A. MAINE:
(MO) (012
s.n., cult. DUSS-82-0503
(MO).-WISCONSIN:
Manitowoc
Co.,
Friedrich
NEW BRUNSWICK: St. Andrew's,
Charles,
Ind. Sem. Bot. Gard. Liege
(MO) (012
cult. DUSS-77-0407
Renner,
Co.,
Hansen
Co.,
S of Hut
and
Isle of Mellum,
Ind. Sem. Bot. Gard. Oldenburg
1I,);
and 111).-NORDRHEIN-WESTFALEN:
Buchholz
near Bergheim,
(MO) (014).
Italy. REGION VENETO. Prov. Venezia: Venezia
(Venice),
cult. DUSS-79-0660
Cumberland
mi NW Durand,
Preston
up Cheat Mtn,
LIEGE: Angleur,
Juterbog
1942)
1933, Alexander
41 (WVA).
Isle of Borkum, Ind. Sem. Bot. Gard. Old
11).-NIEDERSACHSEN:
Groden,
Ind. Sem. Bot. Gard. Olden
(MO); Jadebusen, Riistersieler
s.n., cult. DUSS-82-0481
of 0.
1II, Renner,
(WVA). Mon
511 (WVA). Tucker Co., N of Parson City,
Boone
between
Co., War
Hardy
1933A
Co., Largent,
& Bartlay
&
FSU, GA, MO,
(MO) (0)12 and
cult. DUSS-77-0401
1980, Dietrich
collection
and
(012
WVA).
250, 0.5 mi
along Hwy
of 0. Renner,
collection
Ulm").-BRANDENBURG:
Hiebs
Co., Pow
Springs, Martin
FLAS,
Wherry
1937,
(DUKE, MO,
Shenandoah
NOVA SCOTIA: John River, Hall
(MO). Canada.
(POM). Au
13377
Sulphur
VPI, WS,
NY, US). Morgan
S of Franklin,
Co., near Hinton,
BADEN-WURTTEMBERG: Ulm,
2017. Germany.
US,
3 mi S of Ridgley,
Co.,
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. Belgium.
1975
14285
(CAS, LAM, OKLA,
& Pennell
Wherry
734 (CAS, DS, DUKE,
Frye
UC,
Randolph
Summers
(CAS, CU, GH, NY,
6883
(BH, NY, POM, US).
VIRGINIA: Greenbrier
s.n. (GA, KANU,
1949, Sargent
13484
Sargent
1mi E
Co.,
opposite Mt. Union,
Hamilton,
Munz
Springs,
13489
Springs, Munz
Co., 4 mi E of Springfield,
62 (NA). Hampshire
of Newton
E of Head Waters,
Ridge,
s.n. (POM, US).-WEST
ell's Fort, 1934, Artz
(PAC). Huntington
(GH). Craig Co., 0.5 mi SE of New Castle,
of Eagle Rock, Wood 6802
Co., Shenandoah
2449
(BH, CAS, CU, GH, NY, POM, US), Munz
(VPI). Bath Co., Millboro
Co., Stuart Run, Millboro
PH). Rockbridge
2 mi W
Co.,
Co., Harris
(PAC, PH, UC). Dauphin
5 mi SW of Covington,
Co.,
14204
19038
gusta Co., Headwater,
Fogg
Alleghany
13484: Munz
cult. from Munz
4050
of Pogue, Keener
s.n. (NY, PENN). Mifflin
1932, Wherry
1932, Wherry
POM, US),
Berkheimer
of Saxton,
0.5 mi ENE
Fulton Co.,
(V); Iroquois Falls,
Point, Baldwin
84?23'W),
Baldwin
5231
5 mi
(CAN,
et al. 6431
SYSTEMATICBOTANYMONOGRAPHS
202
(CAN);
Sibley
of Long
TTwp. (48?20'N,
Point, Brassard
Stonecliff,
Brierly
Park, Brown
& Hodge
Lake Ontario,
Brighton,
Adam's
Hurkett,
Garton
ton 15103
9612
TRT); Dryden,
of Steel River,
Harbour
perior, Michipicoten
lan's Point
(43?38'N,
1972, Johnson
80?15'W),
mouth, Manitoulin
(48?00'N,
79?23'W),
s.n. (TRT); Temagami
4781
near Northwood,
Munz
1967, Reynolds
s.n. (TRT); 25 mi NE
14637
1898, Scott
(CAN); Niagara,
Outer Duck
(48?20'N,
139
1338
(DAO, NY);
(CAN).-QUEBEC:
Oka,
Bouchard
Chicoutimi,
OKL,
70-663
a Claude,
Anse
& Bonin
ture, Marie-Victorin
WIS);
Desbiens,
main
123
du Golfe
(CU); Paspebiac
mi W
of Danbury,
IND, NY,
POM,
Larsen
1902, Williams
(DS, POM),
(NY), Munz
(RSA),
3606
Nichols
1237
(PENN,
PH).-DISTRICT
Vassey
s.n. (F); Riverside,
(MIN).
Franklin
4284
(UNCC).
nald
s.n. (NEBC). Waldo
Lincoln
enke & Moldenke
Mary's
14254
6233
Co., Millstone
(POM).-MAINE:
Island, 1919, Jenney
Springs,
(NY).-MARYLAND:
Montgomery
Blake
Co.,
Co., Barnstable,
Hartford
Nichols
& Tidestrom
between
Munz
Quantico
13378
Co.,
cult.
Co.,
Plainville,
St.
C6te Nord
PH, RM,
GH, MT,
Wiegand
& Wiegand
(POM), Munz
Nichols
Dewey
Co.,
14167
3
(BH,
1916, Bissell
6
s.n.
367
(MICH),
(NA).-ILLINOIS:
Cook
Crampton
Bartlett
Berkshire
339 (US).-IN
Greenman
Co., Madison,
1766
1892, Fer
Park, Mold
(MICH,
& Tidestrom
7432
3457
Pond, Rossbach
York Co., Ocean
from Blake
8
River,
Co., Old Harbor
(UNCC).
Nichols
of Mispillion
Co., S of Harpswell,
Tidestrom
IND, POM).
& Rolland-Ger
Co., cult. from seed col. by Nichols
Knox
Bethesda,
(NA),
& Rol
CONNECTICUT: Fairfield
14156
Co., Kankakee,
6081
& Salisbury,
(BH, DS,
des Monts,
et al. s.n. (MO). Somerset
Rossbach
11966
(CAN, CAS,
4 (MICH),
Cumberland
Rivers,
(GH, NY, UC, WS);
Kent Co., 2 mi N of mouth
s.n. (MO, NEBC).
Co., N of Stockton
(IND, NY). Wicomico
SETTS: Barnstable
(POM).
8 (MICH). Kankakee
1913, Knowlton
Co., Monhegan
Landing,
28180
s.n. (NEBC). New Haven
3 (MICH),
Steele
15882
13387: Munz
(BH,
(DAO,
(VPI); Bonaven
(DAO, GH, MT, NY, US);
OF COLUMBIA: N of Long Bridge,
Bay, Buhl F64
Co., Farmington,
14649
& Matapedia
7338
Marie-Victorin
s.n. (CU). U.S.A.
17 (MICH).-DELAWARE:
Co., Chicago,
DIANA: La Porte Co., Pinook
& Fernald
15295
26562
83 (DAO, GH, MT, NY, UC,
(MT); near Ste. Anne
cult. from Munz
1863, Robbins
16 (MICH), Nichols
17715
& Rolland-Germain
49649
(DAO); Havre
de Pabos, Marie-Victorin
Marie-Victorin
& Rolland-Germain
Marie-Victorin
14193
Sainte-Adelaide
du Loup, Marie-Victorin
Riages,
127 (DAO);
Hamel
of Restigouche
confluence
Cte.
(BH, CU, MIN,
765 (NY); Bic, Rousseau
89 (MT); Trois-Rivieres,
Lac St.-Jean,
14171
St. Leon,
286.1998
of La Ville Cap Chat, Uttal
45 (DAO, MT, US);
& Rolland-Germain
Co., Medford,
Bartlett
(MICH), Nichols
(DAO, MT);
"Cap Chat," W
Grandtnter
715 (MT); St. Vallier, Munz
Islands, Robinson
a Persil, Riviere
Marie-Victorin
Munz
US),
(NEBC). Middlesex
from Orange,
31259
Lemieux
Is
Little
2909
de Riv., Kamouraska,
Morin
(NY); Wood
(MT);
& Trapido
Saint-Jean-Port-Joli,
s.n. (MT); Dobleau,
(US); Seven
& Rolland-Germain
13387
St. Pierre, Clausen
1181
Bay
PH, WS);
MIN,
63?12'W),
Boivin
Scoggan
(DAO, GH);
& Rolland-Ger
1267
Erskine
(46?26'N,
Eboulements,
et al. s.n. (DAO); Quai
77 (CAN, GH);
590
ile Dumais,
Lighthouse,
Munz
Les
near summit of Mt.
Rousseau
Natashquan,
Marie-Victorin
(MT);
78?51'W),
Taylor et al. 2529
Bay, Marie-Victorin
Georgian
on
Park
Island, Dawson
84?45'W),
Beach
& Rolland-Germain
St.-Laurent,
US); Dolbeau,
s.n. (TRT); Pig
(47?00'N,
of Brackley
s.n.
(CAN, DAO, MT);
near Cap des Rosiers,
Alban,
et al. 2240
(CAN, GH); W
Barabe
Potter
Marie-Victorin
Beach,
(45?09'N,
Southampton,
Soper
(CAN, CU, GH, MICH,
1934, Morin
(DAO, MT, US);
Point
(IND); Rondeau
Twp.
et al. 7832
67 (DAO, MT, US); Anse
land-Germain
Stroud
1937,
(42?34'N,
(TRT); South Bay
vic. of Camp Hermosa
(DAO);
EDWARD ISLAND: North Rustico,
1932, Gates
Pleureuse,
32063
5683
Fernald
& Jordan
Kelsey
James Bay, Scoast,
GH, MO, MT);
Rousseau
(TRT); Wasaga
Tignish,
(DAO);
Iles de laMadeleine,
NY, US);
236
1940,
PAC, UC); Trois-Pistoles,
E of Riviere
Aubert,
(TRT); Guelph,
(DS, TRT, UC); Mamainse
Taylor 8366
& St. John 11133
Fernald
Sands,
7389
Taylor et al. 832
s.n.
Island, Han
1, last 0.5 mi
14750
Ridout
Lake Huron,
Beach,
1927, Hosie
TRT); Toronto
Munz
193
to
Gar
(CAN, MT, TRT); Lake Su
6 mi N of Goderich,
Twp., Ropke
s.n. (TRT); Ipperwash
(DAO, MT, NY, UC, US).-PRINCE
land, Erskine
Racine
Is
3 mi NE of Black
97?37'W),
Island, Krotkov
from Windsor,
Sable
1mi E of
Bay,
89?17'W),
49 (DAO);
Area
Point,
7 mi S of Dorset,
IND, POM, WTU);
of Chapleau,
& Grevatt
Island, Soper
88?50'W),
(CAN, TRT, UC); Midland,
main
(BH, GH,
Bear
Twp.,
cult. from seed of Gates
(DAO);
17524
Forest Reserve,
Groh
S of
Twp.,
Rd at turnoff
(48?25'N,
(CAN, DAO,
(TRT); Long
S in Lambton
s.n. (TRT); crest of
on Armstrong
Bay City
et al. 1712
et al. 1909
s.n.
96, 98 (TRT); Colborne
Island, Maher
Lake Huron, Minshall
et al.
4 mi
on Black
Bay
(48?45'N,
Hosie
87?15'W),
Hosie
85?00'W),
Centre
(TRT);
(DAO, TRT, UNCC);
Station
Thunder
s.n. (DAO); Nipigon,
(48?45'N,
1974, Hoy
6655
of Hurkett
Ranger
s.n. (TRT); Mine
Hincks
Jackfish
Sturgeon
Rd at Expressway,
1932, Groh
near Toronto,
67 (CAN);
(TRT); near mouth
23 mi N of Black
(CAN, UC); Olwir
S side
3116
Pt., Prov. Park, Brighton
Park, 1949, Edmund
mouth
(DAO);
& Hainault
(TRT); near Grand Bend
Presqu'ile
Park, Garton
(DAO, UBC);
3110
& Bragg
Island, Brassard
Lake, Algonquin
120, Quetico
8698
Garton
1066
(DAO, NY);
(TRT); Brewer
(DAO, TRT);
10425
Brown
603
1mi N of Hwy
Cape,
(CAN, MICH,
stock, Haber
350
of theWoods,
Bay, Garton
Chief
Ingolf, Denike
Dumais
Thunder
Apple,
land, S end of Lake
s.n. (GH); Blind River, Bassett
(TRT); Yorkshire
2994
2672,
(TRT); Point Pelee,
629
(TRT); Rocks
5677
Bannan
88?50'W),
& Hainault
VOLUME 50
POM).
St.
11966: Munz
(NA, US).-MASSACHU
Co., Pittsfield, Mazzeo
2327
OENOTHERA
1997
(NA, WTU).
14682
Dukes
Co., Chappaquiddick
(POM). Hampshire
s.n. (BRY).
Plymouth
POM).-MICHIGAN:
legan Co.,
Arenac
Co., Point Lookout,
Park, Bay City, R.R.D.
17549
Munz
Experiment
899
City, Gereau
land, Voss
4515
& Hsi 23424
MICH,
MIN,
& Grayson
s.n. (MIN). Van Buren
Pease
(MIN). Washington
511
Co.,
(CU, GH).
Co.,
Sodus
Erie Co.,
Presque
Northumberland
14194
487
Mearns
1422
Beaver
Dam,
Co., 0.25 mi
9512
17540
(BH,
Meyer
265
(WIS).
s.n. (WIS). Buffalo
Iowa Co.,
s.n. (MIN). Manitowoc
Co., without
waukee
Ave.,
Co., Lake
shore S of Grange
Outagamie
Co., Appleton,
Washington,
Iltis 8264
Tiffany, Green
Sec. 30, Kuhlman
1959, Rill
363
further locality,
Cudahy
(T6N, R22E,
(Springfield
Co.,
Hill),
Island
Monroe
Co.,
(T22N, R6E,
9940
Sec.
3), Skroch B169
(MIL). Dodge
381
on
Co.,
(WIS). Kewaunee
1932, Wadmond
(T27N, R7E,
& Leykom
SE 1/4 Sec.
s.n. (WIS). Mil
3345
(MIL, WIS).
4 mi NNE
(MIL). Rock Co.,
Sec. 6), Ziegler
(WIS). Ash
s.n. (WIS). Grant Co., Castle
(MIL). Sheboygan
Farm, Sorensen
2430
& Christensen
23240
Island,
1.4 mi E of
s.n. (WIS 6 sheets). Dunn Co.,
1/4 Sec. 36), Shinners
s.n. (MIN). Waushara
on the Hammerstrom
7151
shore of Lake Michgan,
Expedition
11 (T14N, R7W,
1932, Wadmond
terminal moraine
s.n. (WIS). Wood
of theWisconsin,
NW
Co.,
Co.,
41, 6 mi N of Suamico,
Co.,
1959, Patman
s.n. (PH).
(GH, PH).-WISCONSIN:
Co., Kenosha,
(WIS). Marathon
(PH).
13405: Munz
Prudence
Sorensen
State Park, Cross
(WIS). Kenosha
Iltis 14434
(WIS). Pierce Co., Prescott, Museum
50 (WIS). Vernon
consin & the adjacent
& Hansen
Co.,
Greene
Smith
1915, Goessl
Tower Hill
s.n. (WIS). Ozaukee
179 (WIS). Sauk Co., Dells
worth Co., Geneva
tonville,
1933, Rogers
City,
(PENN).
s.n.
1896, Crawford
et al. 5391
of Hwy
s.n. (CU).
Taylor
from Munz
Poin,
Sec. 23), 1961, Ugent
River,
farm, Newton,
5), 1965, Torin s.n. (WIS). Marquette
Fountain
Co., Eau Claire,
inWisconsin
shore, Hansen
Co., Polster
Co.,
(CU). Wayne
Rood 247
1914,
cult.
terminal moraine,
1mi W
Co., Staten
6359
1918, Gershoy
s.n. (NY).-VIRGINIA:
Co.,
(T27N, R26E,
97 (WIS). Eau Claire
from Lake Michigan
Brown
US),
Fernald
Co.,
Richmond
ISLAND: Newport
the adjacent
Co.,
s.n. (NY). Oswego
& Curtis
Fort Hill,
Co.,
s.c. s.n.
1898,
(CU). Monroe
of Ridgeway,
Co., Grand
1896, Bicknell
IND, POM).
1899, Bicknell
POM,
Co., W of Kiptopeke,
s.c. s.n. (UC). Door Co.,
1893,
IND, NY,
Philadelphia
&
2400
Island, Bronx,
(PENN).-RHODE
Lake Wisconsin
& Maguire
Elk Co., W
Co.,
Co.,
s.n. (GA).-NEW
Co., Lakewood,
Island, Muenscher
Co., City
(BH, DS,
IND, NY).
& McCoffrey
(BH, IND, NY, POM, US, WTU).
Co., Shelter
(US). Coos Co., Ran
JERSEY: Atlantic
s.n. (PH). Cape May
1912, Stone
1977, Ahles
(WIS). Montgomery
Co., Providence,
line, 1968, Page
SW of Elk Mound,
Rock, Hemphill
(BH,
(VPI). Northampton
Munz
Co.
13405
Glowenke
Co., Old Bed of Glacial
Oconto-Brown
5853
96 & 61, N of Lake Mas
298
Wright
(CU). Chautauqua
(GH).-PENNSYLVANIA:
Bright
14281
Hallsteads,
land Co., Ashland,
6073
Sunbury, Munz
(US). Providence
Harlow
Quinque,
Adams
Co.,
14271
s.n. (NDG). Westchester
Werner
(IND, POM), Munz
Susquehanna
10498
Bartlett
Scott Co.,
s.n. (MN).
s.n. (PH).-NEW
Bay, Lake Ontario, Muenscher
s.n. (NY). Suffolk
Isle Peninsula,
Co.,
& Wiegand
at Selkirk, Munz
1897, Greene
OHIO: Lake Co., Painesville,
on Island A12,
Co.,
1925, Fryblund
96 from jct. Hwys
Twp.,
188 (DS,
Point,
1-2 mi NE of Atsion,
Co.,
& Grayson
5 mi NE of Split Rock River,
1918, Knowlton
Co.,
along Le Sueur River, Decoria
vic. of Weirs,
Co.,
St. Clair Co.,
Point,
s.n. (NY). New York Co., Fort Schuyler,
1895, Britton
Point,
Ocean
shore of Snowshoe
cult. from seed col. by Wiegand
Island, New Dorp,
Co., Durham,
Point, Eames
1784, Bartram
Beach,
ca. 5.6 mi E of Hwy
Co.,
s.n. (PH). Burlington
Co., Long
Co., Rocky
Co., Hol
s.n. (MICH). Wayne
& 3 mi NE of Brule River, Bartlett
HAMPSHIRE: Belknap
Strafford
(NEBC).
Gershoy
Jefferson
Blue Earth Co.,
Lake of theWoods
RSA).
792 (MICH).
Co.,
& outlet
lighthouse
s.n. (NA). Ottawa
Zimmerman
Is
(CU). Oakland
Little Point Sable
1910, Kauffnan
Lake Co., near Lighthouse
TEX, WTU).
1913, Long
YORK: Cayuga
Jack Pine Region,
South Haven,
s.n. (WIS).-MINNEsoTA:
139 (MIN).-NEw
37353
Island Beach,
Co.,
Co.,
Iosco Co., Tawas
Co., North Manitou
s.n.
1965, Perdue
City,
Co., Rose
Co., Big Stone Bay,
472 (MICH).
& Wiegand
between
1.6 mi N of Waters,
(MICH).
Co., Mackinaw
(SMU). Clinton
s.n. (MICH). Leelanau
Co., Lake Michigan,
(MICH). Al
725
Garlitz
111 (KYO). Emmet
1937, Wiegand
(MIN). Cook Co., SW of Hoveland
B249
terman, Lindayen
1895, Fallas
Co., Marquette,
Co.,
13585
Shinners
13), Brooks
s.c.
1929,
(IND, MICH,
edge of Lake Huron, Waterworks
Co., The Entry, Hermann
(MICH). Houghton
876
Voss 2476
Co., Alpena,
(MICH). Cheboygan
of Paradise,
SE 1/4 Sec.
(T5N, RIW,
1646
(MICH). Otsego
(DS, MICH,
Jordan, Bollard
0.5 mi W
Co.,
s.n. (MICH). Roscommon
NY, RSA,
217
Cape May,
Chippewa
1936, Schmidt
Moore
Rapp 502
(LAM). Cass Co., Vicksburg,
1/2 (GH). Oceana
Voss 5316
Twp.,
Windsor
(MICH). Bay Co.,
Station
1901, Cooper
(POM).
12485
Ehlers
land, 1910, Kauffman
Livonia
E of Au Gres, McVaugh
(MICH). Marquette
Farwell
Algonac,
(MICH). Alpena
Island, Munz
Co., Lowell,
of Grand Marais,
12656
Plum
Co.,
Bartlett
Co., Revire,
ca. 2.5 mi W
Sable Lake
(MICH). Kent Co., Grand Rapids,
5154
of Silver Lake,
s.n. (UC). Suffolk
McVaugh
5149
Lake Region,
Rochester,
of Holland,
(BH, NY, POM).
Lake Wildlife
Douglas
dolph,
Co., N of Grand
Alger
Essex
s.n. (MICH).
s.n. (UNCC). Middlesex
1932, Manning
1926, Blake
Co., Rochester,
S & 4 mi W
3 mi
1915, Bicknell
Island,
Co., Northampton,
203
Co., T13N,
2890
of
R23E,
(MIN). Wal
Co., Old Bed of Glacial
4205
of Port
1.5 mi W
(WIS). Waupaca
Lake Wis
Co., Clin
(WIS).
REPRESENTATIVE
SPECIMENS
FROM
AREASWHERE
NATURALIZED.
Austria. NIEDER-OSTERREICH:
Airport
Schwechat
near Wien,
1968, Forstner
s.n. (W).-WIEN:
railway
station Breitenlee,
1968, Forstner
s.n. (W);
SYSTEMATICBOTANYMONOGRAPHS
204
Praterspitz,
1915, Korb
Winterhafen,
s.n. (W); between
Tscherning
BANT: Bruxelles-Nord,
Lawalree
SEVERCESK'Y: Liberec,
Jehli'k 2773
8210
Dubitsky
France.
s.n. (P); Ottmarsheim,
Mannheim
ten, 1867, Richter
s.n.
4372
438
BURG: Hamburg,
(G, LD, M);
Isle Juist, 1931, Schiitt
Beuel
s.n. (BREM);
near Bonn,
(GOET);
1900,
Niederwartha
at River Elbe,
derstedt,
at River Elbe,
s.n.
1978, Gutte
Zeche
(M).-SACHSEN:
BADEN-WURTTEMBERG:
1049
(L).-NOORD-HOLLAND:
(L). Norway.
1854, Thaeder
Netherlands.
& Reichgelt
Oudemans
Haarlem,
ROGALAND: Madla
s.n. (KTU).
Rostanski
Kern
566
near Stavanger,
s.n. (AMD).-SCHLESWIG-HOLSTEIN:
Lauenburg,
1877, Ndldeke
s.n. (BREM).
Switzerland.
Budapest,
1917, Aellen
1935, Kloos
near
s.n. (L).
1958, Vervoort
Oostvoorne,
s.n.
1949, Hoogland
KRAKOW: Jaworzno,
s.n. (Z). Ukraine
& Thellung
River
s.n. (M). Lithuania
Isle Rottum,
s.n. (0). Poland.
Ei
Lagymanyos,
bed of Tagliamento
1992, Lippert
FRIESLAND: Isle Ameland,
(BR).-ZUID-HOLLAND:
BASEL: Basel,
s.n. (Z); Peninsula
Hungary.
near San Vito,
1962, Wischmann
s.n. (LZ, WRSL);
s.n. (GOET).-SACH
1884, Hippe
(L).-GRONINGEN:
12194
of River Elbe,
s.n. (M).-RHEINLAND-PFALZ:
& Rostanski
s.n. (BP). Italy. REGION FRIULI-VENEZIA GIULIA. Prov. Pordenone:
cited as 0. ammophila).
at mouth
s.n. (BREM);
s.n. (BREM).-NORDRHEIN-WESTFALEN:
1992, Lippert
1975,
1905, Focke
s.n. (FR); Isle Scharhorn
1941, Penzes
(see Rostanski
(HBG).-HESSEN:
Cux
at River Elbe,
s.n. (M); bed of river Tagliamento
s.n.
Lenzen
Isle Baltrum,
1967, Gutte
1903, Focke
&
s.n. (C, LD, NY).-HAM
Prignitz,
1988, Dettmar
s.n. (BREM);
Berlin,
1875, Ascherson
at River
Spilimbergo,
GELDERLAND: Millingerwaard,
(M).-BRANDENBURG:
s.n. (G, GOET).-NIEDERSACHSEN:
Prosper,
s.n. (LZ); Magdeburg,
1850, Steinoorth
between
s.n. (M); Donauschuit
1971, Rostatiski
Coswig,
s.n. (LA); Konigstein
s.n. (C).
1838, Lenormand
Colmar,
1890, Scheppig
1899, Lemmermann
s.n. (HBG); Helgoland,
1929, Vogeler
1959, Pedersen
Koldby,
s.n. (LZ); Oderberg,
s.c. s.n. (GOET);
1961, Conert
Isle Mellum,
1898, Wolf
near Lauenburg,
Holler
1967, Pietsch
1878, Poscharsky
s.n. (B); Bottrop,
Schellenberg
SEN-ANHALT: Rosslau,
Artlenburg
Elbe,
HJ0RRING: Em
Ringk0bing-Fjord,
1895, Krdnzle
Schnackenburg-Allee,
Isle Wangerooge,
1891, Wirtgen
Bingerbruick,
Dillingen,
Donau,
BRA
Republic.
(BR, C, L, MA).-MEuRTHE-ET-MoSELLE:
s.n. (CAS, LD); Westend,
at River
Priesnitz
s.n. (HBG);
at River
Ruhland,
(NY). Czech
s.n. (B). Denmark.
L. s.n. (WRSL).-MECKLENBURG-VORPOMMERN:
1913, M. G. s.n. (FR); Hitzacker
1963, Mathiesen
6737
s.n. (W); Prater,
P, Z). Belgium.
(GOET).-RINGK0BING:
s.c. s.n. (P). Germany.
s.n. (B).-BAYERN:
1930, Gorz
1909,
s.n. (HBG);
Jaap
Rastetter
Nevers,
Hochstadt
s.n.
2860
5858
5468,
s.n. (F, GOET, WRSL).-HAuT-RHIN:
s.n. (AMD);
1866, Andre
near Mainz,
Gustavsburg
Prov., Chung
1852, Winkler
(Teplitz),
(BR); Rixheim,
1934, Baschant
s.n. (HBG); Quanzsee,
Fujian
LA, LD, LY, M,
1978, Luitken s.n. (LD).-TONDER:
(P).-NItVRE:
1887, Scheppig
C, DS, G, GZU,
Isle of Fan0, Wagenitz
1873, Stahl
Rastetter
and Neckarau,
Friedrichshagen,
haven,
(PR); Teplice
Agger,
Strasbourg,
Sont de Frouard, Godron
1888,
(BR). China.
(MJG).-THISTED:
BAS-RHIN:
Lange
1627
1897, Arbesser
and Stadlauer Bridge,
Bridge
(B, BM, BREM,
1933, Lund s.n. (C).-RIBE:
mersbeek,
Rudolph
5056
(Dorfler)
VOLUME 50
s.n.
1983,
(see Rostatiski
1975, cited as 0. ammophila).
SPECIMEN CULTIVATED IN BOTANICAL GARDEN. France.
13. Oenothera
no. 3569,
France.
cult. DUSS-82-0473
nd. Sem. Bot. Gard.
(Bober),
Switzerland.
Co.,
(014).
(Fl, herb. Webb)
(as Onagra
Garden,
and Augusta,
Stubbe
Stubbe
(MO) (0 14).-WEST
(MO)
1981,
(MO)
(0 14).-HAuT-RHIN:
Dietrich
s.n.,
5415,
Ownbey
Ridge
s.n., cult. DUSS-82-0475
Parkway,
cult. DUSS-82-0461
(MO) (014)
Highland
Shenandoah
VIRGINIA: Pendleton
(MO)
(Naumburg)
(MO)
cult. DUSS-79-0661
Staunton,
Co., Highland-Augusta
Stubbe
Augusta
CAROLINA:
(MO).
Co.,
Botetourt
line along Hwy
9 mi NW Franklin
(MO)
YORK: Wash
between
19, cult. DUSS-81-591
678, Stubbe
1942).
Washington
cult. DUSS-79-0588
(MO) (014).
county
along Hwy
Co., SE North Fork Mtn,
(0)14).-NEW
(0314).-VIRGINIA:
28, cult. DUSS-81-593
Co., Fort Valley
3979,
at River
Renner,
(as 0. atrovirens).-NORTH
ca. 1330 m, Solomon
NORD
between
(014).-SACHSEN:
silesiaca,
1976 no. 1355,
(MO).
KRAKOW: Jaworzno,
cult. DUSS-86/88-1018.-MINNESOTA:
(MO) (014),
97, Stubbe
(as 0.
(014)
(MO);
(MO)
Poland.
(014).
GORA: Novogr6d
(MO)
30, cult. DUSS-81-594
along Hwy
cult. DUSS-82-0474
(MO)
no. 3570
(MO). Germany.
cult. DUSS-82-476-1
(MO).-ZIELONA
cult. DUSS-77-0383
18, cult. DUSS-590
(014).
cult. DUSS-77-0416
cult. DUSS-77-0385
17517,
Liege
1978
cult. DUSS-79-0614
Munz
Ind. Sem. Bot. Gard.
Liege
280 m, Ind. Sem. Bot. Gard. Basel
Blue
s.n., cult. DUSS-88-W867.
cult. DUSS-81-592
no. 35,
LIEGE: Amay,
Ind. Sem. Bot. Gard.
1979 no 1797,
1980, Dietrich
Co., Camden,
221 with
Carrol Co., Pipers Gap
Stoutamire
Renner,
of 0. Renner,
jct. Hwy
Co., Devils
Staunton
1975
s.n., cult. DUSS-84-212
of 0.
MAINE: Knox
collection
B9,
no. 214,
State Park, St. Croix River,
ington Co.,
Wilkes
collection
1976
BASEL: Briigglingen,
U.S.A.
Co., Afton
along Hwy
Leipzig
& Dietrich
1983, Rostaniski
(014).
Lyon
near Grevenbroich,
and Dormagen
Sturzelberg
Angleur,
(014);
215 m, Itd. Sem. Bot. Gard. Basel
RHEIN-WESTFALEN: Allrath
Avery
(MO)
ALLIER: Ind. Sem. Bot. Gard.
Rumersheim,
B6br
1836
parviflora.
SPECIMENS EXAMINED FROM CULTIVATED PLANTS. Belgium.
1978
Sep
Paris,
v. parviflora).
chrysantha
(MO)
Co., Gala,
250, Stubbe 21,
17, cult. DUSS-81-589
along rd 33, 1978, Glen
1997
OENOTHERA
coe & Rossbach
Glencoe
s.n., cult. DUSS-79-0555
& Rossbach
(MO)
s.n., cult. DUSS-79-0554
rd 33, 1978, Glencoe
& Rossbach
s.n.
6016
Chrysler
(GH); Sun Co., Acadia
Dore
Woodstock,
Sunnybrae,
& Gorham
& Bassett
Mulligan
5 mi W
1080
17508
River, Munz
Cape Bald,
(MT); Moncton,
12282
Bassett
Valley,
1674
John's, Greer
Mills,
(DAO);
Camb, Wentworth,
476
Grant
Lunenburg,
(IND, POM),
Munz
mile River, Pease
749 (CAN);
Ingonish
578
Light House,
Schofield
3702
1961,
s.c. 1324
Otter Lake,
135
(TRT); Cochrane,
Forest
Experiment
Sound,
Portage
10-104
Partridge
Lake,
SMU); Norval,
5103
1mi NW
of Norval,
32 mi W of Rossport,
Osnabruck
Stormont,
Temagami,
Esquesing
1973, Drecktrah
water Bay, Lake of the Woods,
1946, Gordon
Bay
5224
near Constance
(CAN); Durham,
Midland,
Haddow
Wagner's
(DAO); Guelph,
E of Jellicoe,
& Jennings
Jennings
(TRT); Sibley, Mazokama
14518
(DAO);
Harcourt
800
1959, Lloyd & Frith
(UBC); Constance
Lake,
McKay
Lake, Macoun
468/22
(CAN, MICH,
wardsburgh
2406
Twp.,
(RM); Vespra
Lake Ontario,
Ramsay
85924
MIN,
Twp., Blakeney,
Park, Moore
Brighton,
Munz
(TRT); Stewart's
Minshall
951
17554
Minshall
861
3700
(DAO);
Ropke
(DAO);
(TRT); Rouge
499
Bay, Hosie
Swamp,
Lake,
Sturgeon
23700
River,
Port Hope,
(DAO);
3371
1898,
(MICH);
Lake
(GH, TRT); Moose
(DAO); Wong
River
Bay, Krotkov
9253
Lemon
s.n. (TRT); Renfrew,
(CAN);
548
Survey,
s.n. (TRT);
1926, McVittie
85927
Inlet,
Clear
Island, Groh
s.n. (TRT); Avonmore,
of Little Rouge
s.n. (TRT);
Casselman,
Malte
(TRT); Grenville,
Nelson
and Rouge
387
Ed
& Nelson
Rivers
S to
(DAO); Lanarck,
s.n. (CAN); Barron River Canyon,
17552
Lake Ontario, Munz
Cartwright,
1890, Scott s.n. (TRT); Ottawa,
et al. 1715
Scott
(GH,
17555
17,
(NY);
Sandy
4008
Service, Wye
near Lake Nipissing,
1968, Miron
2606
(DAO);
of
PH,
along Rte
(DAO); Kenora,
of Port Arthur, Ropke
Munz
Bay,
(ILL); Stokes
(CAN),
confluence
of Gananoque,
(KANU, MIN,
Park, 1944, Lloyd
Falls,
60 mi NNE
(TRT); Westworth,
Cody 99 (DAO); W
1921, Harvness
s.n. (TRT); Hespeler,
85925
Petawawa
(MO); Parry
Cunningham
Jenkins
Buck
Toronto,
s.n. (TRT); Simcoe,
Island, Grassl
2 mi SW of St. David, Miller
(DAO); Moosonee,
25 mi NE
(BH, IND, POM);
Ste. Marie,
Bush,
Highland,
s.n. (TRT); Muskrat
2540
Macoun
18741
3039
466
(DAO); Ottawa,
s.n. (TRT); Rockcliffe
Rockcliffe,
TRT); Lincoln,
Twp., Reznicek
1973, Riley
of Sault
(CAN);
(CAN, TRT);
7940
Gogo
1972, Leadbeater
1928, Lloyd
(CAN).-ON
Station, Edmondson
Gillett
River,
Station,
Jones
near Wilberforce,
s.n. (TRT); Hawthorne,
Spencerville,
gonquin
15 mi W
Bay,
1928, Lloyd
Twp.,
S of Kenora,
Prince
223
(DAO, TRT); Thunder
Dow's
near 5
Wilson
56795
s.n. (TRT); Lake Nipigon,
111 (CAN); Carleton,
(GH, TRT, US); Haliburton,
Mink
1161
Cliffs
Economy,
(GH); Cumberland,
Boufford
92 (CAN); Canadian Wildlife
s.d., Hamilton
145
Hillsdon
& Tuomikoski
Hustich
Groh
(TRT);
35584
5129
Falls, Demaree
Summerstown
River,
(GH); Hants,
1929, Brown
Lake, Haley's
s.n.
Middleton,
s.n. (TRT); North
s.n. (DAO); Gore Bay, Manitoulin
on Ottawa
5 mi E of Port Perry, Haber
400
Lake, Uxbridge,
River, Renison,
W
1156
Clawe
Annapolis,
(TRT); St. Davids,
612
Island,
(BH, NY, POM, US, WTU);
857a
Lake, Brown
St. Lawrence
Island,
s.n. (MIN); Glengarry,
Buckhams
Bay
Twp., Dickinson
s.n. (OSH); near Jeffrey's
SE shore of Steen
Twp.,
1927, Gilmour
15289
1901, Anderson
(TRT); near Neston
of Riley
1938, s.c. s.n. (UBC); Addington,
5362
St.
(CAN, MT);
beach
Wedgeport,
10 (TRT); Peel, Credit Forks,
335
Connolly
73-263
Rd, near Glen
Rousseau
61 (TRT); York, Musselman
Is
4 mi NE of Tomp
1915, Fyles
Point, Baldwin
Brown
65-6366
(CAN); Kentville,
E of Batchawana,
Algoma,
Haultain,
& Hay
Street, Wolfville,
Point Pelee,
shore of Long
Bay, Britton
(TRT); Peterborough,
Twp.,
Pleasant
(DAO); West
& Pugh
s.n. (UAC); Colchester,
Ile Madame,
Jacquet
(CAN); Grand Manan
(CAN, NY);
(POM),
PH);
(GH); Long Beach
Island, Nichols
Arichat,
(DAO);
Brown
Stanhope
21193
14749
(TRT); Essex,
W
12365
Yarmouth,
1921, Prince
(CAN); Kings,
Lake Abitibi,
Lake, N of Gordon
Brown
105.35
1883, J.X. s.n. (TRT); Aylmer,
Farlain Lake near Penetanguishene,
Waterdown,
s.n. (DAO);
Munz
(NY); Richmond,
Blair
Station,
55748
Gates
Ferry, Cape Breton
64-1952
Roberts
SCOTIA: Guys,
Erskine
Island, Macoun
(PH); Kings, Wolfville,
s.c. s.n. (TRT); London,
shot, 1932,
(V); Sable
Restigouche,
VPI).-NEWFOUNDLAND:
s.n. (MT).-NovA
Oxford,
(GH, PENN,
(BH, IND, POM); Charlotte,
Spencer,
S Dis., Deer Arm, Bouchard
1930, Groh
24
Pictou, Robinson
Isle au Haute,
(FSU, UNCC,
(POM); Lockeport,
(POM);
& Long 21992
TARIO:Kingston,
Baldwin
14758
14752
7530
Bay,
Carleton,
8 mi S of Grand Falls,
& Bateman
Scoggan
Quoddy
s.n. (DAO);
IND, POM);
17515
Roberts
of Sussex,
(GH); Port Mouton,
Sydney,
Lemkow
(BH,
1933,
Cloverdale,
Island,
(MT); Victoria,
St. Jean, Cape
15 mi W
(CAN); Cumberland,
(DAO); North
303129
17507
Munz
Co.,
& Long 14197
Jct., Fernald
Malte
(DAO);
St. Barbe
57.35
along
1960, Cunningham
Jemseg marshes,
1939, Penson
Lake,
Gates
Beach Meadows,
14734
Munz
51521
Erskine
Uttal
793 (DAO);
1978,
and Onego
Deer
(DAO);
4 mi E of Sussex, Munz
64-2659
(CAN); Norton,
(DAO); Deer
Liscomb
Queens,
Bathurst,
(BH); Queens,
s.n. (DAO); Hampton,
1308
Station,
St. Andrews,
Kings,
& Bateman
Anderson
York, McAdam
(DAO, MT);
s.n. (DAO);
17516
Roberts
3 mi NW Onego,
Harman
(MO) (014).
Experiment
(DAO); Gloucester,
Scoggan
land, 1953, Sharp
kins, Codroy
Forest
(BH, IND, POM, WTU);
of St. George, Munz
morland,
mont,
45885
1935, Groh
Moncton,
Run
Co., between
Randolph
BRITISH COLUMBIA: New Westminster
BRUNSWICK: Edmundston,
(TRT).-NEw
rd 33, McIntosh
along
(014).
s.n., cult. DUSS-79-0560
REPRESENTATIVE SPECIMENS. Canada.
Brown
(014);
(MO)
205
IND, POM);
Al
18 mi E of
(BH, IND, POM); Gros Cap,
s.n. (TRT); Wick,
1890, Scott s.n. (TRT); Russell,
Bourget,
1898,
Scott
Clarence
s.n.
Twp.,
206
SYSTEMATICBOTANYMONOGRAPHS
Senn
1275
1276
(DAO),
Soper
s.n. (TRT); Halton,
Twp.,
along Maitland
(DAO); Muskoka
Distr.,
George's
Waterdown,
River
opposite
et al. 14 (DAO); Aldbourough
12149
56
(SMU);
Ontario,
Marie-Victorin
land-Germain
UNCC);
Snelgrove,
GH); Black
End,
1935, Adams
1247
Smoky
9028
near Kapuskasing,
Falls,
s.n. (DAO); Arthabaska,
Amqui,
(CAN, MT);
1958,
Lac Aylmer,
Calder M-221
11418
(MT); Quebec,
St. Vallies
NW
658
Desmarais
s.n. (DAO); Lachine,
Hamel
Saint-Paul,
Laprairie,
1932, Fr. Euph.-Jos.
1173
(MT), 1087
MT);
St.-Vallier,
457
Gauthier
(CAN); Bromptonville,
St. Lawrence
(CAN); Grosse-ile,
76-1109
Preston,
364
1149
Ste.
(MICH);
& Lepage
1mi
15308
Parc, Nat. Forillon,
1043
Montcalm,
Ste-Anne-de
Thetford
(DAO); Nicolet,
3493
Jolicwur
Joyal 1304
Car
s.n. (DAO); Mt. Orford,
(MT, TRT); Megantic,
River, Hanson
ile de Montreal,
Lamoureaux
(MT); Chemin
de l'aqueduc,
Reservoir
(CAN); St.-Calixte,
s.n. (MT); Iles des Sceurs,
(DAO); Lake
(MT); Mont-Royal,
et al. s.n. (DAO, GH, MT,
Gates
12536
Parrish,
1182 (MICH, MT);
1932, Groh
Hamel
Stratford,
s.n.
Templeton
Dutilly
(MT); Gaspe-Sud,
s.n. (DAO); Kamouraska,
(CAN),
Desmarais
James, Harricanaiw,
(CAN); Bellechasse,
311
(MT); St. Jean, Desmarais
Boucherville,
Baie
(CAN);
1971, Bouchard
51-245
Sillery, Cayouette
Deschamps
Lac Aylmer,
Lac Simon,
(DAO);
Alphonse
155
Bergeron
Papineau,
(MT); Ile Ste.-Therese,
702
G1517
Lac Massawippi,
10693
St. Nicholas,
76-865
s.n. (CAN); Papineau,
Lacasse
328
du Mont-Orford,
(CU,
Island
Ste. Anne,
Bellemare
Clkonique
(DAO, MICH,
1936, Groh
C66129
Houle
658
Perce, Mt.
Baie Missisquoi,
(CAN, COLO);
(TRT,
7835
Fernald
Point-du-Lac,
1162 (CAN, DAO, MT);
(DAO); Wolfe,
Hamel
Becancour,
5705
Grandtner
la-Pocatiere,
d'Irlande,
Cayouette
Gallo
1935, Groh
Mines,
Roberval,
(GH); Levis,
riere du Cap-des-Rosiers-Est,
59
13162
1194
Matapedia,
70.35
Gates
LIslet,
& Hamel
& Rol
4036
(DAO); Cochrane,
Ste.-Foy,
Parc
Sherbrooke,
(MT); Stanstead,
Sillery, Desmarais
Desmarais
(RSA); Bonaventure,
US);
66455
Deschamps
(MT, WIS);
pointe Amos,
of Champlain,
(DAO);
s.n. (MT); St. Maurice,
1914, Churchill
Foy, Desmarais
(DAO); Louis-Hebert,
Brisson
Garthby,
(DAO); Pointe-Bleue,
Memphremagog,
Cleonique
Ile Perrot, Bernard
Vaudreuil,
(DAO); Wolfe,
s.n.
Smith 266a
(MT); Missisquoi,
13, Vick
on Lake
Lake, Watson
s.n. (TRT).-QUEBEC:
1037
Allyre
2000
& Gervais
Belzile
just SW of Tignish,
1938, Whelan
Toronto
Gananoque,
Marie-Victorin
Colborne,
Park, Canoe
Spicer
Taylor & Taylor
s.n. (TRT); Birchcliff,
EDWARD ISLAND: Cape Aylesbury,
Victoriaville,
Anderson
Rd S of Ottawa,
84 (DAO, MT, US);
Algonquin
1949,
Goderich
(TRT); Huron,
(TRT); Borden
(DAO); Northumberland,
Prince Co.,
(CAN);
along East River,
945
Soper
28 mi N of Sault Ste. Marie,
1978, Varga
Nipissing,
s.n. (TRT).-PRINCE
(MT); Lake St. John, Nomandin,
Matapedia,
Island,
143
near Huntsville
5008
(CAN);
& Rolland-Germain
151 (DAO, MT);
1908, White
River, Macoun
N of Rock
part of Ward's
& Rolland-Germain
150 (DAO, MT),
Bear
near Hamilton,
2618
Stewart
Marie-Victorin
Caledonia,
Billie
Camp
Soper & Shields
Benmiller,
Twp., Elgin,
Islands, NE
Toronto
(TRT); Prescott,
Woods,
VOLUME 50
(MT); Saint-Placide,
(CAN); Lac Saint-Jean
Lac
(MT, UC);
1925,
Est, Saint-Gideon,
1950, Laverdiere s.n. (MT);Charlevoix, Baie St.-Paul,Lemieux 286.2577 (DAO);L'Assomption, St.-Liboire,
Briscotte
& Riv.
a Pensis,
below
Cap
a L'Aigle,
& Rolland-Germain
Grosse-Ile,
main
& Durand
Marie-Victorin
UC, US),
(CAN);
(CAN); Gasp&Nord,
Macoun
67933
US),
La Mauricie
Nat.
Cap des Rosiers,
(CAN, GH);
& Rolland-Germain
I (DAO,
34 (DAO, MT, US);
et al. 1026
71-43-35
1932, Marie-Victorin
Saint-Bruno,
Victorin
Louis-Marie
l'Achigan,
River, Lamoureaux
St. Felicien,
Marie-Anselm
3 (DAO,
US),
& Rolland-Germain
16 (MT); Lac Temiscamingue,
Chambly,
Marie-Victorin
39 (DAO, GH, MT, US),
125
(DAO, FSU, MT),
US),
St.-Joachim,
Marie-Victorin
(DAO, MT),
(PAC); Port
114 (DAO); Chambly,
Marie
6 (DAO, US); Montmagny,
& Rolland-Germain
126
from St.-Maurice
40117
s.n. (DAO, TRT); Montmorency,
2 (DAO, MT,
Longueuil,
1mi
Park,
Louis-Marie
127
& Rolland-Ger
38 (DAO, MT, NY, TRT,
(DAO,
FSU, MT, US);
Fort
Temiscamingue,Marie-Victorin& Rolland-Germain42 (DAO);Deux-Montagnes, St.-Augustin,Marie-Victorin
& Rolland-Germain
80
MT),
53 (DAO,
(DAO, MT);
cheres, Contrecour,
FSU, MT);
Lac St.-Jean,
Marie-Victorin
(MT); Portneuf, Marie-Victorin
land-Germain
27158
las, Marie-Victorin
14751
Munz
60020
(IND,
IND, POM); Montreal,
Vaudreuil,
350
Laval,
Rigaud,
(MT), 604
Munz
Parnis
Montcalm,
419
1741
(BH),
17550
(BH,
15291b
St.-Jerome,
Rolland-Germain
Tremblant,
et al. 40128
14744
Cap Tourmente,
bank of St. Lawrence
St.-Adolphe,
125-4-5
7868
Petite,
de Lachine,
(MT), 6041
14755
& Coutu
7896
St.-Nico
St. Vallier,
(POM); Ri
17513
(BH,
(BH, IND, POM);
Ranger
(CAN, DAO,
(MT); Saint-Francois,
2291
& Rol
(MT); Levis,
17551
&
3171
Cascapedia,
(IND, POM);
Munz
Munz
iles des Rapides
Rolland-Germain
Rolland-Germain
Marie-Victorin
River, Lavoraie, Munz
25 mi SW of Quebec,
765 (CAN, DAO),
(MT); Ver
(MT); L'Islet, Marie-Victorin
(DAO, GH); Riviere
Marie-Victorin
Iles aux Herons,
79 (DAO,
109
sur les platieres, Marie-Victorin
28321
Rolland-Germain
Lac Monroe,
Mont-Laurier,
et al. 2392
(BH, NY, US);
IND, POM);
(DAO); Verdun,
(DAO, TRT); Argenteuil,
Parc du Mont
(MT); Labelle,
(IND, POM); Charny, Munz
(BH, POM); Berthier,
(MT); Terrebonne,
Laval-des-Rapides,
land-Germain
15291a
Grosse-ile,
14742
& Rolland-Germain
& Rolland-Germain
(MT); Riviere Macdonald,
(CU); Montmagny,
17509
117
Lorette, Marie-Victorin
(MT); Bic, Munz
POM),
St. Fabien, Munz
mouski,
Marie-Victorin
Marie-Victorin
Venise, Marie-Victorin
et al. 3703
(PH); Ancienne
et al. 33825
Marie-Victorin
Brackenridge,
d'Alma,
& Rolland-Germain
189 (MT); Missisquoi,
Rolland-Germnain
Hull,
St. Joseph
231
ile d'Orleans,
(FSU, SMU),
(CAN, MT);
(MT),
FSU, US);
36551
Brome,
Rol
(MT);
Foster,
OENOTHERA
1997
8520
Rolland-Germain
Parc de la Verendrye,
(MT); Gaspe-Sud,
Rossbach
near Wagon
epic, Rousseau
50278
ieres, Stanislas
574
(MT); Brandypot
245
(MT); Westmount,
Victorin-Lavoie
U.S.A.
field Co., Canoon,
cemetery,
land Co., Coventry,
Munz
Champaign
3 mi E of Urbana,
Co.,
Co., W
gomery
1mi SE of the Shades,
Co.,
(IND). Washington
(WIS). Clayton
Co.,
son 3436
(SMU). Webster
(IND). Hickman
1939, Bean
5222
1 mi
E of Westemport,
Cecil Co., Grove
Co.,
Club, Downs
Co.,
Churchill
1919,
Co.,
Thies
et al. s.n. (DUKE).
SMU,
TENN,
UBC,
Indian River, Munz
17547
361
(DAO, MICH).
195d
of Mt.
528
2932
(DS). Pope Co., Glenwood
543
Highland,
(MIN). Winona
1912, Bartram
Jaffrey, Munz
13364
Long
4763
(MICH). Ocean
Co.,
10458
2 mi
Co.,
Co., Winona,
SW of Carpentersville,
Grafton
line, Ahles
Co., Elwood,
Co., Cape May,
Co., Toms River,
Ballard
Co.,
69120
s.d., Burk
1873, Martindale
Schaeffer
59523
s.n. (PENN).
Co., Grand
of Mio
along
Co., Detroit,
Portage
Co.,
path to
International
& Grayson
Bartlett
1403
3 mi N of Stillwater,
Gun
St. Louis Co., Merewee
133 (CU). Cheshire
(PENN).
Burlington
Co.,
Co., New Mtn,
Co.,
Hillsborough
Co.,
1915, Taylor
Co., Crowleytown,
Cecil,
Co., Belmar,
YORK: Cayuga
&
Shore Drive,
s.n. (RM). Houston
Co., W Claremont,
Gloucester
s.n. (NY). Monmouth
(PH).-NEW
on Bay
(MIN). Koochiching
Sullivan
5 mi S of
St. Ignace, Bartlett
1897, Moore
s.n. (TENN).
s.n. (WIS).
Co., Flint, Has
4 mi NE
Co.,
Cook
360-B
1940,
PAC, PH,
Co.,
20 (MICH). Wayne
Intervale, Moore
s.n. (NA). Passaic
1889, Wiegand
Co.,
St. Croix River
(OKLA).
1966, Magee
Palmer,
& Bill
Creek
& Frankford,
Bowen
Berkshire
s.n. (MICH).
(BH, IND, NY, POM, US).
s.n. (MIN, NY).-MISSOURI:
Co., Hanover,
1891, Lighrhipe
Co., Truxton,
2749
HAMPSHIRE: Carroll Co.,
(BH, IND, POM).
Co., Woodbridge,
s.n. (DAO).
Grand Meadow
1886, Holzinger
county
Thompson
FSU, MICH,
ISC, MIN,
(BH). Genesee
of Birch
&
(BH, IND, NY,
(FLAS,
(MICH). Cheboygan
Co., Minneapolis,
Co., Hallock,
13475
1943, Gates
s.n. (ISC). Oscoda
1965, Disrud
31 (POM). Washington
s.n. (PH).-NEw
JERSEY: Atlantic
(CU, GH). Cortland
1920, Smith
5.5 mi NE of Vassar,
(PH). Frederick
Forest Lake,
(POM). Mackinac
near mouth
Tidestrom
6190
(DAO, DS, GA,
17548
US).
s.n. (GH).-MARY
1910, Bicknell
14686
Co., Top
NEBC,
(BH, IND, POM).
Co.,
1903,
17517
Sorrento,
Munz
Island,
Co., Paxton,
900
Bay
Co., Deer Creek between
Park, Moore
(PH). Cape May
(PENN). Middlesex
Lake, Hanes
Co.,
13380
Hampden
Munz
Sagadahoe
et al. 3707
Co., Avon, Munz
Munz
Munz
(GH, MO,
River, Benner
Island, Voss 4797
(MIN); Hennipin
(MO). Kittson
0.7 mi NE of Hillsboro/Cheshire
s.n. (PENN).-NEW
Co.,
Anoka
5018
Ownbey
76 (ISC). Mower
Falls, Kellogg
Norfolk
445
1891, Fernald
Munz
76114
1923,
Co., Auburn,
Co., Camden,
Baltimore
Co., Chappaquiddick
Co., Green
Co., Whitehall,
(MICH).-MINNESOTA:
& Leykom
Co., Ziegler
Co., Harrison
(MICH). Tuscola
Josephine,
Swansea,
3 mi N of Escanaba,
Menominee
(MICH). Muskegon
summit
WVA).
8043
Co., Lexington,
(NEBC, VPI).
Co., Kensington,
(BH, NY, POM, US).
Co.,
Knox
Top Mtn, Windler
Ahles
7094
Twp., Hartley
Androscoggin
& Strong
(UNCC).
5 (BH, PH). Worchester
(BH). Delta
11 (RSA). Kalamazoo
Farwell
14685
30501
Point, Sassafras
Co.,
Co., N shore of Fox Lake, Beaver
selbring
3517
& Round
Co., Williamsburg,
UWL, WTU,
Richards
Perry Creek, Nimke
1434
Deam
Deam
Franklin Co., Farmington,
(NCSC).
Fernald
(UNCC). Montgomery
(MIN). Dukes
Munz
Fayette
(MT).-MAINE:
Knowlton
Grove
Barnstable
Station, Kennedy
MICHIGAN: Charlevoix
Downs
Neck,
(POM).-INDI
s.n. (ISC). Van Buren Co., David
(VPI). York Co., Cornish,
Hancock
Hampshire
UNCC,
Co., Marion
Plymouth
Grassl
s.n.
3 mi S of Newbury,
Essex
East
2585
between
PAC, UNCC).-MASSACHUSETTS:
Sheffield,
dersen
30502
Buhl F51
Co., Center
(BH, POM).
Lexington,
s.n.
NY, US).-ILLINOIS:
(WVA). Lee Co., SW of Ft. Madison,
2287
17518
MIN,
Tol
1921, Otis
1875, Young s.n. (NY). Mont
s.n. (COLO).-KENTUCKY:
Munz
(CU).
s.n. (RM, UWM).
0.5 mi N of Versailles,
& Aikman
215
237
(SMU). Litch
along Lime Creek, NE part of county,
Co.,
1929, Crally
Trois-Riv
1.5 mi NE of Martinsville,
Co.,
Co.,
Allamakee
1064
& Anderson
Co.,
(MICH, UC).
POM, US). Washington
Co.,
Knowlton
65319
Co., Bryn Mawr,
s.n. (WS). Hancock
Lake, Easterly
Point, Bristol,
Somerset
& Wiegand
Co., Hanover,
s.n. (IND).-IOwA:
Co., near Mars Hill, Downs
Co., Pemaquid
Bluffs,
Cook
Jefferson
(MICH). Ripley
Somes
Wiegand
state line, Ahles
s.n. (F, GH, MICH,
(IND, POM). Morgan
4500
Co., Ellsworth,
s.n. (NEBC).
Co., River Rd Country
OKLA,
Ft. Dodge,
Co.,
Hancock
Co., Roque
LAND: Allegany
Bartlett
51015
3 mi SW of Polk City,
Lincoln
Furbish
1912,
Washington
Deam
Co., Clinton, McFarland
s.n. (NEBC).
(BH, POM, WTU).
sham,
(IND).
1924, Shimek
s.n. (NEBC). Aroostook
Knowlton
5074
1930, Brooks
of Giard,
s.n. (ISC). Polk Co.,
13536
s.n. (SMU).
Deam
Pat
Ste.-Anne-de-la-Pocatiere,
Castle Co., no further locality,
New
1948, Storm
St. Maurice,
(DAO);
1928, Jaussan
Co., Groton,
Steele
1902,
Park, Umbach
Salem,
Co., W
Park,
s.n. (ISC). Iowa Co., above Dutch
1927, Shimek
Shimek
of Greentown,
Porter Co., Dune
(MIN).
London
25725
near Lac Porc
Saint-Fabien,
Kamouraska,
(TRT); Rimouski,
(IND, POM).-DELAWARE:
ANA: Howard
13963
82 (PH). New
Lambert
OF COLUMBIA: Tacoma
(PH).-DISTRICT
s.n.
1902, Wilkes
et al. 818
(CAN);
Rousseau
(CAN, DAO, MO, MT);
Rimouski,
Senn
6895
Terrill
Island,
32064
jct. of US Rte 5 & 1-91, just S of MA
Co.,
13384
(CAN, GH, MT);
4 mi E of Wakefield,
(DAO); Gatineau,
CONNECTICUT: Hartford
32445
& Bonin
Le Domaine,
(MT); Pontiac,
de l'Ile d'Orleans,
Ste.-Petronville
Rousseau
Rivers,
Rousseau
Branch,
9630
Rolland-Germain
Coin-du-Banc,
(CAN); Montmagny,
& Matapedia
of Restigouche
(MT); confluence
apedia River,
7628
207
Fosberg
14459
1910, Alsberg
s.n.
True 5 (PENN). Warren
Co., Union
s.n. (CU). Franklin Co., upper Saranac,
Springs,
Eames
1926, Lloyd
s.n.
SYSTEMATICBOTANYMONOGRAPHS
208
Co., Bergen
Swamp, Muenscher
burg s.n. (NY). Montgomery
(PH). Genesee
Co., Amsterdam,
9 (UAC). Oswego
office, Hyduke
(COLO).
Saratoga
Co., Waterford,
US). Warren
Big Hollow,
OLINA: Ashe
Poebles
Co., Highlands-Walhalla
Spruce
Perdue
(UNCC).
Yancey
& Blum
ferson Co., Broadacre,
NIA: Allegheny
(PH). Berks
1877, Moyer
SW
Creek,
973
Lehigh
6171
dence Co., Blackstone,
4749
Sohmer
Co.,
ISLAND: Newport
terford, Seymour
21864
& Seymour
(MO). Lamoille
s.n. (TRT). Rutland
1924, MacMasters
5 mi NE
Amherst
of Covington,
on US
Co.,
60 along Huffs
(BH, IND, NY, POM).
13480
Fauquier
19770
Co., Allard
of Monterey,
James
13487
Munz
7665
William
17333
3 mi NW
Co.,
(IND, POM).
(US). Greene
James City Co., College
2 mi
Co., N slope
Morgan
290
Bartholomew
279
2, Stahmaun
341a
Fassett
Hertel,
(WIS). Dane
Dunn
facing Potomac
7252
SW of Laona,
Twp., Pucker
088
F with
Bell & Colvin
S of Hwy
238
2-3 mi NW
Jackson Co.,
(WIS). Marathon
185
River
Co.,
Co.,
284
Co.,
Creek,
Rd
Ridge
1954, Wagner
198 (OSH). Manitowoc
Co.,
Park,
Schmidt
297
in Sect.
7111 (WIS).
s.n.
(MO).
70, 1957, Snell
on Hwy
Iltis & Noamesi
Co., Hwy
Co., Viola
1958,
s.n. (WIS). Kewaunee
(WIS). Marquette
1967,
of Reedy
Co., Strandberg
Spaulding
Steele
Reservoir,
Adams
s.n. (WIS). Burnett
Co.,
(WTU, WVA).
Co., Terra Alta,
Ferry, Bluestone
near Tippler
of Merrillon,
(OSH). Lafayette
Co., Bushnell
Preston
6616
3 (WIS). Clark Co.,
1967, Rosing
Twp., Peters
VIR
3 mi S of Princeton,
Co.,
& Davis
23 (WIS). Door
s.n. (WIS). Greenlake
54 (WIS). Kenosha
64, Wolf
Wollangk
s.n. (WIS). Florence
St. Rte. 42 on F, Novotny
along RR, 0.25 mi
Forest,
(OSH).
Burger
(UNCC).
1965, Baker
Co., Faber
13490
1mi NE of Bayard, Downs
s.n. (UNCC).-WISCONSIN:
1964, Evans
36 (UT). Dodge
1964, Knight
Davis
Co., Shanklins
S side of Lake Chetek,
1928, Bachmann
Co., Menomonie,
of jct of Co.-
Bell
Springs, Munz
PENN).-WEST
(F). Wirt Co., 0.5 mi above mouth
Co., N Stockbridge,
(WIS). Calumet
Co., Eagle Heights,
Smith 1571
Valley,
Co., Herndon,
(WIS). Barron Co.,
son Co., 2 mi S of Sullivan,
Co.,
Co., Hacker
(WVA). Summers
flood plain,
s.n. (WVA). Hardy Co., E of Wardensville,
10271
Downs
SE
et al. s.n. (PH). Prince
(BH, NA,
s.n. (WVA). Mercer
Co., Morgantown,
at Brosius,
Core 6407
(WVA). Wyoming
(WIS). Forest Co.,
Brooklyn
River
Co.,
(UNCC). Mont
Co., Rte 680, Tye River
1936, Fernald
17355
Munz
s.n. (US).
Highland
544
Co.,
(IND).
of Goshen,
(BH, NA).
s.n. (WVA). Grant Co.,
1934, Greenlee
Hill,
14286
1915, Steele
Co.,
Co., Lick Run, Millboro
State Park, 1938, Sumpstine
Co., Watts
IND, NY, POM). Monongalia
s.n. (WVA). Webster
Phillips
& Triplett
20 mi NE
Barans
and Mary,
Fosberg
Co., near Wa
Alleghany
13487: Munz
17163
Fosberg
Co., Wa
Co., E Corinth,
Washington
Co.,
(VPI). Nelson
SW of Swift Run Gap,
Co., Tomlinion
Putnam Co., Nitro,
(NY, US).
209
Augusta
Rd,
of Crossville,
(PENN). Orange
s.n. (MO). Fairfax
West
(PH). Provi
Caledonia
(UNCC).
(NEBC).-VIRGINIA:
3 mi SW of Petersburg,
1958, Nestor
(WVA). Kanawha
(BH, DS,
628, Smyth 3903
(NA). Rockbridge
Co.,
Hancock
13502
on Hwy
6-8 mi NW
Co.,
from Munz
of William
of Joplin, Rudd
7277
Munz
(UNCC).
Co.,
Co., near Moatsville,
8104
12206
1907, Steele
5 mi E of Furnace,
Co.,
2512
cult.
POM),
(VPI). Prince George
Rockingham
(UNCC).
IND, ORE,
James
Creek,
(UNCC).
GINIA: Barbour
Rossbach
(BH,
et al. 10058
s.n. (DS, DUKE).
Underwood
Co., Hartland,
Bath Co., Millboro,
gomery Co., 6 mi NE of Blacksburg
Frees & Ramsey
1924, Dutton
Co., Brandon,
terburg, True 185 (MIN, PENN). Windsor
3331
Dam
Indian Rock
Co.,
1321
1mi
Co.,
Schaeffer
1mi SE of Fairhope,
Co.,
True 6942
Co., Eden Mills,
Westerfield
(MICH). Montgomery
Cumberland
Co.,
s.n. (PH). Greene
1.5 mi S of Raerhersville,
Island, Fernald
Co., S Shaftsbury, Marshall
Bennington
Co., 0.5 mi E
(PAC). Delaware
SE of Treichler,
(NY). York
Salt Pond, Block
Co.,
mi
Somerset
7168
Osterhout
s.n. (NEBC).-TENNESSEE:
1923, Leland
(UWL).-VERMONT:
1-1.5
Jef
Co., Quakertown,
SE of Denholm,
Co.,
11450
Pretz
Co.,
s.n. (PENN).
1953, Wherry
Tunkhannock,
Lancaster
(PENN).
Northampton
1304
0.5 mi
(PH). Juniata Co.,
10294
Wahl
(PENN).
of Bushkill,
11 (PAC).-RHODE
Zarfoss
s.n.
(PAC). Wyoming
108
Donley
(POM).
from Georgetown,
Bucks
near state line, 1918, Bright
Creek
1.25 mi SE of Spring Valley,
Co.,
1954, Wherry
Pike Co., 0.5 mi W
(PENN).
erfield
Station,
299
s.n. (PAC). Chester
Hoover
Co., Grantham,
of
38751
Radford
s.n. (MICH).--PENNSYLVA
(FLAS).
1941, Westerfield
PH). Cumberland
162
2.7 mi W
DAKOTA: cult. from
8 mi up Ohio River
416-32
CAR
Blomquist
Co.,
Benson
Co., Bueyrus,
Co.,
POM,
region,
(NY).-NoRTH
SW of Plymouth,
1902, Grover
Brumbach
S mi SE of Philipsburg,
Co., N of Richardville,
(PENN).
of Hatfield,
of Birdsboro,
1489
Co., Old Fields,
(UNCC).-NORTH
Crawford
s.n. (ISC). Beaver
1907, Sumstine
Pump
46881
IND, NY, ORE,
Co., Lake George
(DUKE). Mitchell
3.3 mi
Lorain Co., Oak Point,
(PH). Fulton Co., Big Tonoloway
Brave
(PAC). Jefferson
& Duke
Ahles
913
Co.,
(IND).-OHIO:
(UNCC).
s.n. (PENN,
1942, Stone
Co., Dunkard
14746
0.5 mi NE
Co.,
upper Darby, Long 60077
Brubaker
1362
s.n. (NA). Centre Co.,
of Downington,
12452
of Day Brook,
Co., Wilkinsburg,
34-A
Bright
Cusick
Falls, Keever
(FSU, TEX). Washington
from Barrie, Munz
Durham
(NCSC).
1126
Island, Cold
Co., Long
(BH, DS,
Gleason
Co., Yonkers,
S.U. Bursar's
Co., Todt Hill, Weber
73 (WVU). Washington
11463RMD
at Satula
Hwy
4164
4 mi SSW
Co.,
seed sent by Gates
Downs
Jefferson,
13391
behind
Co.,
s.n. (CU). Suffolk
Ithaca, Munz.
s.n. (CU, GH). Westchester
(GH). Macon
Pine,
Co.,
Margolin
1885, Poggen
Co., Ridgewood,
s.n. (IND). Richmond
1911, Burnham
Island,
(MIN). Tompkins
5 mi N of W
(CU). Kings
s.n. (MICH). Onondaga
1935, Wiegand
Co., Tripp Pond, Chestertown,
1896, Burnham
Co.,
21934
1939, Silva
Co., Selkirk,
C. S. H. C14229
Spring Harbor,
WTU,
& Brown
VOLUME 50
s.n.
23 of
Jeffer
Co., 2.2 mi NE
s.n. (WIS). Langlade
Co., Point Beach
State
1.5 mi N of Germania,
OENOTHERA
1997
Vowles
22 (WIS). Menominee
5 mi W
Co.,
Co., Little Chute,
Co., Clam
Twp.,
(WIS). Trempealeau
(WIS). Walworth
Waushara
of Keshena,
Co.,
Perrot
above Virginia
Co.,
s.n.
1906
Sorensen
4950
Lake,
s.n.
s.n. (LD); Kalksburg
Rechinger
s.n.
(W).-STEIERMARK:
(WIS). Vilas
(WIS). Waukesha
15859
162
(BR).-LIEGE:
(BM, BR,
Anseremme,
(PR).-ZAPADCESK'Y:
1891, Spindler
Rastetter
9476
Bois
LY, P).-PARIS:
s.n.
Hertel
(M).-BRANDENBURG:
Berlin,
1891, Hirte
pig
s.n. (AMD,
Dettmar
7945
Baschant
1934,
(B).-SCHLESWIG-HOLSTEIN:
Potamo
on Isle of Kerkyra
Valbusa
1451-2
(TO). Japan.
HONSHU: Hyogo
1961,
s.c. s.n.
Nukui,
Koganei
(MAK);
City,
Shiga
& Kobayshi
Mizushima
bicht, Mennema
1737
Te Puke,
"Glenmark
Sykes 50/85
tariski s.n. (KTU, WRSL);
Schur
1272
NBG,
PRE, S). Sweden.
cal Garden
(P); Nove
1974, Lousley
Philcox
Glendive,
2157
Lw6wek
1946, Becherer
Zurich,
specimens
Park, London,
(RSA). Wales.
Ouren
1969,
1925, Nilsson
Adema
1933, Lademann
s.n.
1882, Schep
"Prosper,"
s.n.
collections
Tokyo
Kita Koma-gun,
(L).-LIMBURG:
Buriel
s.n. (NY).-CHESHIRE:
v.c. 41, 1905, Riddelsdell
1964,
Greven
Maarssen,
Lincoln,
D.S.I.R.
1976,
near Rheineck,
v.c. 58, Edmondson
D2
(MO,
in Botani
1968, Geissler
Kingdom.
2, 1950, Miller
s.n. (K).-SURREY:
s.n. (BM). U.S.A.
Bratislava,
1435
Spont.
GENtVE:
Nov
Campus,
1958, Ros
s.n. (LZ). Slovakia.
1975). United
at BM, London,
Pref.,
NORTH ISLAND: Bay of Plenty,
s.n. (LD). Switzerland.
(see Rostaniski,
Pref.,
Chiisatata-gun,
CAPE PROVINCE: Tokai, Goldblatt
GALLEN: Fuchsloch,
At
La Loggia,
WROCLAW: Wroclaw,
& Rostaniski
Ka
(C). Greece.
s.n. (L).-UTRECHT:
(0). Poland.
Gutte
1988,
Wittenberg,
s.n. (KYO);
1153
ISLAND: Canterbury,
26328
HAMPTONSHIRE: v.c. 32, 1962, Mayes
Glamorgan:
s.n.
s.n. (KYO); Nagano
Takane-cho,
(L). New Zealand.
s.n. (Z). Ukraine
of Philip Miller
1934, Gates
L);
1893, Meyer
Sanada-cho,
1962, Murata
1875, Groll
2326
(PR). South Africa.
s.n. (G).-ST.
1915, Thellung
s.n. (BM).-NORTH
1880, Ward
6782
(GOET, HBG,
mine
Lange
Sugadaira,
Pref., Kiyosato,
Aerdenhout,
Quadgras
1884,
ISU, LD,
ISU).-MECKLENBURG-VOR
Bottrop,
1845,
BRABANT: Lieshout,
near Wroclaw,
Jehltl
s.n.
s.n. (M); Ulm,
1971, Murata
Honshu,
1890,
Dammartin,
s.n. (LZ).-SACHSEN-ANHALT:
(KYO);
70 (CHR).-SOUTH
Skane, Landskrona,
Photo of type & historic
cult. at Regents
to Kitahira,
VESTFOLD: Larvik,
Zamky,
of Geneva,
s.n. (G).-ZURICH:
Voorburg,
1989, deLange
Rd,"
(CHR). Norway.
s.n.
(BM, G,
Lyon,
1930, Sack s.n. (M).-SACHSEN:
Pref., Ohio, Himeji-shi,
s.n. (MAK); Yamanashi
(L).-NOORD-HOLLAND:
Braun
BAS
(MA).
(B, BM, CAS, G, GOET,
s.n. (FR, GOET,
1978, Gutte
1958, Murata
s.n. (KYO). Netherlands.
s.n. (L).-ZUID-HOLLAND:
biennis).-RHONE:
s.n. (LD). Italy. REGION PIEDMONT. Prov. Torino:
Pref., Omimaiko
1960, Teruya
2264
s.n. (AMD); Westend,
Kiel-Duistern-brook,
1887, Gerold
(Korfu),
to Jyumo; Minomisaku-gun,
Arusayama
Wolters
near Bautzen,
s.n.
Billot
near Guben,
Kiesgruben
near Ludwigshafen,
Hundersheim
SJm-L
Rumer
Bertrand
Liverdun,
1965, Dorr
1857, Scheppig
1888, Darer
Frankfurt,
(LZ); Kleinsaubernitz
s.n.
(M); Schlagsdorfer
Denmark.
s.c. s.n. (LY).-HAUT-RHIN:
near Freiberg,
Lindau,
s.n.
SEVERCESKY: Usti
s.n. (C). France.
1897,
&
Venhecke
1967,
s.n. (DAO, WS).
s.n. (GZU).-NORDRHEIN-WESTFALEN:
1876, Grantzer
s.n. (M).-RHEINLAND-PFALZ:
Otto 4750-2
menz,
Schwaben,
Soignies,
Heurck
(BR).-NAMUR:
Republic.
1916, Andersen
Ge,rard 1175
and Chatel,
s.n. (LD); Charlottenburg,
B, COLO).-HESSEN:
POMMERN: Prenzlau,
(PE). Czech
(BR).-SEINE-ET-MARNE:
at river Dreisam
s.n. (M).-BAYERN:
(B, LD). Straussberg,
325
12864
and Gent,
Langerloo
18225
Lawalree
s.n. (BR); Kessel
(St. Truiden),
(BR),
Scey-sur-Sa6ne,
Chassignol
Vaxancourt
between
BADEN-WURTTEMBERG:
1912, Oltmanns
12863
St. Troy, 1891,
1909, Keller
(BR, C, FI).-HAINAUT:
St. Trond
s.n. (G).-LOIRE-ET-MoSELLE:
Vindecy,
near Wien,
Auderghem,
1860, E. s.n. (LY) (on same sheet also 0.
de Vincennes,
et al. 62 (WIS).
1948, Michiels
1592
Odense,
s.n.
(WIS).
1893,
Kritzendorf
1900, Fleischer
MA).-HAUTE-SAONE:
1863, Ayasse
(Z).-SA6NE-ET-LoIRE:
LY, P). Germany.
Freiburg,
(Pilsen),
s.n. (C).-FYN:
s.n. (GOET).-GIRONDE:
C, M,
(BR,
s.n (LY).-VOSGES:
Gandoger
Plzen
91
1984,
Lawalree
between
s.n.
199
Steams
1940,
& Colvin
Fremdenau
Evere,
et al. 1413
Wang
1925, Allen
near Wien,
1118 (BR).-LIMBURG:
Hsien,
1953, Hansen
LOIRE-ET-CHER: Cour-Chevernie,
Lardiere
Benxi
1961,
NIEDER-OSTERREICH: Amstetten,
(BR).-BRABANT:
Mirwart,
Lake,
Bell
Sussex,
(W).-WIEN:
and Bever, Robbrecht
Wechuysen
Polk
Co.,
of Rib Lake, Anderson
Found
Co.,
Co.,
(BR, UNCC);
(BR).-OOST-VLAANDEREN:
near Copenhagen,
RHIN: Lauterburg,
sheim,
4640
Jehl(k 6754
LAND: K0ge
1647
Strombeek
LIAONING: Cao Houdo,
1mi NW
s.n. (B, MA);
4260
Vanhecke
Ixelles, Dubois
Herstal,
s.n.
1883, Wiesbaur
G, K, L).-LUXEMBURG:
Lawalree
(BR). China.
(AuBig),
Brussels,
s.n. (BR); between
Lo, 1951, Pelgrins
Martins
near Wien,
et al. s.n. (ASU).
(WIS). Racine
s.n.
Co., Birch Bluff, Adney
(WIS). Wood
Schneider
1901,
ANTWERPEN: Mechelen,
(Z). Belgium.
(BR, C, G, L, LD, MA);
Lawalree
Hieflau,
(WIS). Monroe
Co., N shore of Saud Lake,
SPECIMENS EXAMINED FROM OUTSIDE NATURAL AREA. Austria.
Malicky
Z21
s.n. (WIS). Outagamie
1968, Fredrich
1965, LeClair
s.n. (WIS). Taylor Co.,
1926, Almon
Zimmerman
1965, Liesner
Co.,
Saukville,
Co.,
s.n. (WIS). Sawyer
1934, Honey
State Park, Hartley
E Troy,
Co.,
Co.,
s.n. (WIS). Portage
1963, Cochrane
Co.,
Co., W
(WIS). Shawano
Ozaukee
(MO, WIS).
Johnson
1960,
and 2 Lakes Bridges,
s.n. (WIS). Oconto
1969, McKinney
10815
Seymour
Falls
s.n. (WIS). Rock
Carter
Co., between Mishawquit
of Camp Douglas,
209
England.
s.n. (NY);
(K).-KENT:
Walton
v.c. 16,
Common,
MONTANA: Dawson
Co.,
s.n. (US).
SPECIMENS CULTIVATED IN BOTANICAL GARDENS. Denmark.
Copenhagen,
1810, Fischer
s.n. (GOET)
(as
210
SYSTEMATICBOTANYMONOGRAPHS
0. gauroides).
France.
1981, Schreber
Fischer
Paris, Sep 1836,
(as Onagra
herb. Webb)
s.n. (M) (as 0. muricata);
s.n. (GOET)
(M); Munchen,
(NY)
(as 0.
(Fl, herb. Webb)
v. parviflora).
chrysantha
(as 0. muricata);
1832, Martius
Erlangen,
Hamburg,
s.n. (BR). Poland.
la. 0.
1982, Schreber
Wroclaw,
6. 0.
7. 0.
8. 0.
9. 0.
10. 0.
11. 0.
subsp. elata
wolfi
5a. 0. villosa
The numbers
in parentheses
presented
species
the specimen
601
Abrams
(1c); 7168
6313
& Wiggins
438
15 (2); 472
(9); 1002
(1c).
19178
(5a); 8748
Allyre
(13); 76114
(9); 16925
(13); 87706
(8).
& Duke
(9); 46881
Ahles
Ahles
45750
(9); 48973
& Haesloop
(9); 17851
(9); 18380
& James 60481
(13); 47893
(9); 49718
29146
(9); 30162
(9); 60845
(9);
(9); 50137
(9).
(7); 30931
(9); 61688
(9).
(9);
61967 (9); 62828 (8).
Ahles
& Leisner
17082
(9); 17327
(9); 18635
(9);
(9); 32593
(8);
Albee
Aniol
Anway
239
Appler
757
Archer
6884
624
Arnow 4695 (5b).
350
Arsene
& Amable
Ashwin
Atkinson
Atwood
(9).
(12).
(lb); 7187
171 (7); 1554
Arsene
Atwood
(9); 909
(9).
(13).
160 (9).
(13);
(4); 3632
(10).
& Gendlin
(9); 1798
(Sb, pro parte).
(9); 1983
(10); 2246
87 (lb).
Arnold
(lb, pro parte); 3350
(13).
(5b).
(13); 1885
(9); 2152
(9 x 10); 760
759
Antrsi
(9).
3350
(13); 1675
Andrews 267 (9).
(9); 32510
Albertson 35 1-113 (5a).
Albright 94 (9).
Aldous
(13); 2005
(9); 33554
Albee 2119 (lb).
51-A-70
199 (13); 1308
(9); 31925
17 (2).
(13); 3739
(9).
et al. 34 (9).
Anderson
Appel
Ainsworth 449121 (9).
Akitt
(lb).
(13); 1667
(13); 3607
31627
505
(9);
(5a).
(13); 1477
33026
Ahlgren
(9); 11426
(9).
612
& Gilkland
Anderson
2000
& Clark 60049
48732a
(13); 2366
1247
Anastasion
16738
between
Amerson 262 (9).
(9).
Ahles
1037
(13); 3179
(9).
& Bell
List of
intermediate
(8); 3882
(lb); 497
& Cyprien
Alphonse
(9).
(13); 69120
(9); 3584
40 (5b); 262
Allyre
(8); 508a
two taxa (when
Aller 2011 (5b).
1153 (13).
65319
3538
Allen
(9).
(9).
et al. 62 (13).
340
between
19770 (13).
Alleizette 568 (10).
(9).
Aerts 76-48 (13).
Ahles
1832
EXAMINED
in the text and in the Numerical
species
intermediate
Allard
(4); 9301
Adams
& Duncan
a specimen
(9).
Abrams
Ahles
1832
Basel,
Switzerland.
a hybrid).
is usually
Abbe & Abbe
Ahles
Muinchen,
stucchii
grandiflora
nutans
biennis
glazioviana
argillicola
COLLECTIONS
refer to the corresponding
above. An x indicates
Species
Aharrah
(B) (as 0. cruciata).
1832,
Gottingen,
cruciata);
13. O. parviflora
INDEX TO NUMBERED
Adema
(as 0.
(Fl,
(M); Erlangen,
12. 0. oakesiana
subsp. villosa
5b. 0. villosa subsp. strigosa
Adney
s.n.
LIST OF SPECIES
elata subsp.hirsutissima
elata subsp.hookeri
jamesii
longissima
Adams
Paris, Sep 1836
Schreber
s.n. (M) (as 0. muricata);
s.n. (HBG)
1826
cruciata);
1768,
simsiana).
elata
0.
0.
0.
0.
0.
chrysantha
Erlangen,
1832, Frolisch
NUMERICAL
lb.
Ic.
2.
3.
4.
(as Onagra
Germany.
VOLUME 50
(lb).
(9).
(5a); 6115
(2); 6627
3585
69a (9); 694
(la).
(9).
1 (9); 2 (8).
1142
(5b); 5394
& Allen
3156
Augustine 447 (5a).
(3).
(3).
(9); 18573
(Sb).
211
OENOTHERA
1997
Auquier
(10); 3198
1566
4540
Austin
445
657
(lb);
20624
Axtman
(5a); 4540
3650
(10, pro parte);
(9); 1067
(5a); 1075
(12); 420
Baca
24 (lb).
Backe
37595
Bailey
149 (12); 699
(9).
(9); 2884
(10); 2524
Baker
223
(9).
(Sb); 495
(4); 460
3360
(lb); 2327a
1258
(lb);
(lb); 4478
(Sb); 3685
(lb); 9351
Baldwin
& Porsild
et al. 6313
(12); 5129
(13); 8418
(8); 6313
& Breitung
Baldwin
1704
B563
Balogh
3184
(13); 5231
(9); 11221
(9); 3876
(9); 7142
6669
(13); 6431
(12);
(9); 2749
(9); B646
226
Baltzell
(8).
(12).
(Sa); 731101
(13); 3001
Banker
3097
Barans
544
(9); 2498
(Sa);
(9); 5350
(13).
9987
(9).
112 (8).
Barbour
& Barbour
182 (8).
Barclay
37 (9); 623
(8); 4545
045-4
Barnhart
(Sa); 045-77
(5a).
(Sa).
1662
(9); 703
(10); 955
(Sb); 1100
(Sb); 1127
(Sb); 1298
1307
(Sb); 1308
(Sa); 1326
(9); 1350
(9); 1372
(Sa); 1393
(5b);
279
793
Bassett
& Bragg
Bassett
& Hamel
Bassett
(Sb).
(13); SiSS
82 (9); RI-33
1130
Batty
996
(13); 1775
3110
(7).
(13); 753
(12);
(9).
(12); 3132
(9).
(9); 2121
(9).
1982
2852
(Sa); 3250
(9); 2922
(9); 5250
(13); 2935
(9).
(Sa).
(9).
282
(12).
8487
(10).
Bazlun
3588
(9).
6 (9).
Beamish
630211
Beamish
et al. 750251
(Sb).
Bean
16762
Beatie
& Chapman
Beattie
& Kurihara
(Sb); J-S0 (9); M1940
(Sb).
(13).
5692
2259
(5b).
11034
(9).
8511
(Sa).
(9).
& Wiegand
(8).
& Wilson
MO-6
252
36 (13); 4469
(9).
9939
(9); WO
(9); RO-202
(9); 1766
(12);
2247
(9); 2384
(9); 2691
(9 x Sb); 2709
2711
(9); 2714
(9); 2749
(9); 2750
(9); 2806
(8); 2867
3053
(9); 3100
(9); 3102
(9 x Sb); 3114
(9); 3149
(9); 3161
(8); 3163
(9); 3265
(12); 3498
(9); 14097
(9); 14448
(9); 15093
(9); 16004
(9); 16354
(9).
85-I (9); 88 (13); 91 (13); 238
et al. 50128
(9);
Bennett
86 (Sa); 1312
Benson
299
Benson
& Drouet
326
(9);
(13); 2362
(13); 3504
(13); 3505
(13);
Berdau
(12); 3509
(13); 3511
(13); 3517
(13);
Bergeron
311 (13).
3518
(12); 3519
(13); 3521
(9); 3542
Bergman
2635
(9); 3544
(12); 3583
(12); 3584
(12); 3592
(13); 3594
Berkheimer
(lb);
(12); 3604
(12); 3606
(9).
(12); 3613
(12); 3630
(13); 3649
216
(9).
(9); 7142
(9).
(9).
(9).
(Sb).
Berkeley 1675 (9).
(13); 3582
(12); 3599
(8); 3646
(13).
155 (13).
(13); 3501
3601
(8);
Benner 6190 (13).
3506
(13); 3565
(9);
(9);
(13).
3500
(13); 3559
(7); 9793
(8); 10518
13972
(9); 3168
(9); 3499
(9); 8325
(9); 10518
15246
Bellemare
(8);
3146
Bell
(9); 5296
(8); 10048
Bell & Colvin
(9);
(8); 2777
(8); 2958
(Sb).
194 (la).
194 (9).
(Sb); 876 (12); 1029
52 (9).
67-07-17
Beliz
Bell
Bartholomew
(9); 3557
(9);
(13); 1415
(12).
681
Bayliss
Beil
(9).
& Shoulders
(9); 3171
(7); 3214a
361 (13); 585
& Mulligan
Bates
Begert
650-62
Bartholomew
(9); 2780
(Sa); 1403
Beckett 1000 (9).
Bedker 272 (Sb);586 (lb).
(Sa).
Bartholomew
(Sb);
(Sa); 1356
(Sa); 1377
(Sa); 1375
(9); 1401
(9); 3214
& Breitung
Beck C-301
(9).
143-73
695
(Sb); 900
(9); 625
(9); 696
1056
Bechtel
(9).
Barrell & Spongberg
Bartlett
(9); 701
Bassett
Bebb
Barr 9856 (Sb).
Barrell
(9); 700
(10); 859
Bassett
Beal
(9).
& Moreno
733
(9); 4441
(9); 2540
(9).
Barbour
Barnes
(9); 698
Baumann
(9).
(8); 188 (7); 2270
5-79
Barkley
(9); 697
Bartlett & Richards
(Sb).
(9).
(9).
(9); 1789
(9); 4509
Barbosa
(10); 686
799 (12).
211 (9).
Baltars
(9); 664
1384
(Sb); 16026
Balhuizen 6679 (10).
Ballard
(10); 616
(9); 635
(9); 1440
135 (13); 5030
5780
Baldwin
(5b); 614
(9); 626
(Sa); 1361
(10); 4703
(Sb); 15216
(Sb); 14398
(Sb).
Baldwin
(12); 3669
(13).
999 (lb); 1001 (lb); 1003 (lb); 1022 (Sb);
217
(lb);
(8); 3665
(12); 3683
103 (12); 167 (Sa); 188 (12); 217
(Sb); 453
(10); 705
(12); 6603
(10).
Bainath
(13); 3672
Bartlett & Grayson
(9).
(lb).
(9); 3659
(8); 3653
(13); 3670
(9, pro parte).
(9);
Bernard
5529
3394
(9); 3431
(5b); 66455
(13); 4050
(13).
(11); 15673
50
MONOGRAPHSVOLUME
SYSTEMATIC
BOTANY
212
Bicknell 6621 (10); 6634 (9).
Billington
& Farwell
Brassard
(9); 5625
5597
& Hainault
2672
Braun
(13).
(9); 4645
4073
(9).
Billot
2264
Biond
37 (9).
Brayshaw
Bioret
867
Breedlove 15726 (lb); 15886 (la).
Bird
(9).
132 (Sb); 1328
(Sb); 4198
Blair
(13); 883
Blais & Hagy
Blais
2254
(9).
555 (9); 391
Blake
(9).
(Sb).
et al. 10448
(9); 1343
(9); 4688
(13); 3145
1427
(5b); 1452
(9); 1927
(13); 7031
Brierly
243
Brierly
& Hodge
Bright
74-160
629
Brinkman
(13); 7898
236
(9); 2552
Blake & Tidestrom
Brisson
& Hamel
11966
(12).
Blecic 57067 (9).
Blum
162 (13).
2978
Blumer
Boivin
(lb); 4432
3113
Boettcher
481
(12); 1181
9799
Boivin
& Breitung
Boivin
& Dore
& Dunbar
Boivin
& Mosquin
Boivin
& Perron
Boivin
et al. 9992
B249
Bonar
1 (9).
6530
(9).
(Sa).
Borisova
11440
12319
(9); 250-S
Bouchard
& Hay
73-263
Boufford
18741
Boufford
& Wood
Braem
Brain
Brandt
(13).
397 (1c).
(9).
52 (3).
6054
(13).
782
6 (Sb).
et al. 8164
(9); 8260
(13).
(9); 19696
24 (13).
(Sa).
(Sa);
(9); 13363
(Sa);
(5a).
14984
(lb);
(5a); 8317
(9).
(5a).
(5a).
21 (9); 35c (9); 61 (13); 92 (10); 96 (9); 149
(5b); 733
(9); 2656
(5b); 1066
(5b); 884
(9); 3038
(13); 5362
973
(10); 3039
(13); 5429
(12);
(13); 3565
(13); 5677
(13).
416-32
(9).
(13).
1550 (9 x 10).
53-61
Buchon
2400
(Sb); 100 (2).
(Sa).
39-10
(Sa); 31809
Buhl
FS1 (13); F64
Bull
384
(13).
(12).
(2).
Bunce
75 (9).
Burger
54 (13).
(5a); 697
(9).
Burkhalter6230 (9).
(9); 19760
(9);
Bums
20056
9311
Burt-Davy
Bush
(7).
& Hendrickson
758
Bushnell
Butler
(Sb).
Byers
75 (5a).
Cadet
2152
Cady
134 (9).
(9).
Cain 436
(9).
Calalmgh
100 (9).
(13).
(5b); 324
(Sa); 420
(9).
(2).
(9); 2172
(9); 15118
(13); 284
250
1799
Calder M-221
200
(9); 9789
(9); 2160
(9); 14381
232
10763
1368
Bryant
(9).
7654
Brandegee
Brooks
Burmham
19656
& Hillman
Bowerman
(8).
(13).
& Chapleau
Bozoman
7542
Burk 561
19859 (9); 19864 (10).
360-B
& Hauser
Buckstaff
181 (9).
(12).
Bowen
(Sa).
Brumbach
(9).
70-663
Bowden
(Sa); 33678
(13); 5103
(Sa).
28598
18319
Brunner
T-7 (2).
Bourdo
(Sa); 12807
(12).
Brownlie 855 (10).
Bornmuller6820 (9).
Bourassa
(5a); 11820
& McGregor
(9); 111 (12);
(5a); 8041
11488
Brubaker
Bouchard
(9); 11373
(5a); 8006
Brooks
(Sb).
Bottimer
(9); 7800
1565
168 (Sb).
6 (9); 243-21
(lb).
(9); 11256
Brooks
(Sb).
(12).
Bossenmaier
11180
(5a); 485
(Sb).
et al. 929
Bostick
338 (lb); 582
Brooks
Brown
(Sb).
511 (11).
15539
Broder
(13).
(13).
(10).
Brotherson
(Sb)
104 (9).
Boone
Bor
(12); 1416
(Sb); 7905
10439
Boner & Weldert
Bonin
10 (13).
7539
(Sb).
7897
Boivin
Bollard
489
(5a).
(13); 76780
13162
Britten
(9).
& Alex
(12).
(9).
(13); 5571
Britton
(Sa).
70-663
Boivin
65 (Sa).
(9).
1420
Bobbette
(13); 5853
(9);
Brinker 411 (7).
(9); 2023
5326
& Blosser
(5b); 8593
(12).
(5b); 34-A
Brisson
Blomquist
(5b); 4770
(9).
(13).
Blosser
(Sb).
15782 (5b).
Bridges 103 (9).
187 (9).
10-104
34 (10); 48151
Breitung
(Sb).
Birkholz 2576 (Sb).
Bissonnet
(13); 2994
(12); 2888
(12); 3116 (12).
(9).
(8); 11504
(5a).
(13); 320
(12).
(Sa); 13115
213
OENOTHERA
1997
11516
Calder & Savile
et al. 20469
Calder
(lb).
Camazine
069
Campbell
83 (Sa).
6704
Carlton & Garrett
224
Carnes
Canf 157 (Sa).
(7); 1750
(9); 593
(9); 1799
(9).
523
(Sa).
(13); 51-245
50-115
Cayouette
(13); 1576
(Sa); 5705
(13).
789
Chandler
(9).
& Smith
Chapman
10428
Charpin
Chen
Core
(Sa).
Child
322
(Sb).
Christ
2897
(Sb); 12967
(5b); 5461
(lb);
18908
(Sb).
Christensen 3947 (9).
RC35c
& Christensen
Christensen
533
Christian
(12); 5858
601
(9); 10181
(9).
(2); 25740
(9); 33387
(2); 50262
(5a); 618
273
(9); 7026
(8); 7507
(9); 7902
(8); 12401 (lb); 17280 (7).
14785
(3).
(10); 2320
(10).
(lb);
Cottrell
G-3
(Sa).
Couch
113 (Sa).
Coveny 751 (10).
Cowell 153 (13).
2319
(12); 358 (13); 410
339
Crane
2437
(9).
Croat
1326
(9); 2588
(Sa); 3572b
Cruise
8056
(9).
Culbertson4555 (lb).
1102
(9).
Cull 741
Clarke
750902-1
(lb).
Clarks
1953
(9).
1251
Claude-Joseph
& Trapido
Clausen
1156
Clawe
Clemants
(8); 2307
(9); 2667
(11).
(10).
(12).
2909
(13).
(12); 34-22
34-20
Cleland
Curtis
(8); 2273
(9); 932
913
(9); 34-23
(12); 200
(8).
Cleverdon
Clokey
(13); 11633
(Sa).
3 (Sa).
1810
(9); 5542
(3); 8039
(lb).
(12).
39 (8).
Curtiss 5123 (9).
Cusick 1362 (13).
(9).
Custip
903
Cutler
191 (9); 2816
Cutter
229
(3).
(2).
Cybulski 432 (9).
da Cunha
(Sa).
Cleonique 3079 (10); 10693 (13); 10866 (9); 10867
(9); 11418
(Sa); 3620
(9).
Clark & Stevens
1300
(Sa).
(9).
(9); 10659
1614
Cui & Zhu 226
Clark & Landers
Clarkson
(2).
(9).
9072
29971 (9); 66172 (la).
Crook 694 (Sb).
Cross 381 (12).
(12).
Chung In-Cho2671 (10).
Chunys 1065 (9); 1350 (Sb).
Clark
(8).
9586
Cottam
Crampton
(9).
5468
Chung
(13).
(8); 4029
23967
(8).
(13).
1238
(13); 6771
3207
Cowgill
(9).
(3).
Chrysler 6016 (13).
Chu 3839
62 (5a); 7842
& Diggs
Cots & Bold
(2).
Chichester 453 (Sb).
426
(la).
Coryell 1094 (5a).
(9).
Chien
10955
(la);
10954
Correll & Correll
(9).
Cory
et al. 10133
Chiang
(7).
16 (9).
& Baldwin
6407
Correll
(Sa).
1816
Chester
(5a); 50315
(9); 50314
Coles 2528 (Sa).
Collaland& Collethon 7339 (lb).
Collins 9884 (10).
Corcoran
(9).
(Sa); 15427
(10); 89366
6782
Cheny
(Sa).
50312
Cooperrider
(10).
133 (1c).
1984
(13).
(9).
Cooe 15325 (lb).
(9).
Chassignol 325 (13).
Chavez
1917
Contreras
(Sa).
1253
(9); 11309
10526
Chase
et al. 1139
(9).
Comins 1872 (10).
Connolly 335 (13).
(9).
129 (12); 1910
Chaney
22749
Comack
4699
Chambers
13360
Cody & Munro
Coleman
& Koeppen
Cawley
99 (13).
Cody & Kemp
Cole
176 (12).
18525
Cavas
84 (Sa).
Coiteux 135 (9).
(9 x 10).
9281
Caughlan
Cobb
Coile
(Sa).
Castetter6562 (lb).
Castroviejo
4 (3).
Cody
(lb).
(9).
Carter C249
Clute
Cockerell 2 (lb.)
Codd 2756 (2).
121 (8).
Cannon
(3).
& Jotter 2147
Clover
(la).
2090
Calzada
Clover 164 (12).
(Sb).
(Sb).
Daggy
1397
4176
Dahl
27 (12).
Dale
822 (7).
(9).
(8); 4177
(8).
Dansereau& Leclerk 600809-D156 (13).
(Sa);
Dansereau
Dar
et al. 483
1061
Darby
(10); 892
(Sa); 68033
65392
(10).
Demaske
148 (9).
378
603
Dennis
& Dorris
Davidse
Derrwaldt 36 (9).
(9).
1875
45 (9); 628
(9); 2875
(13); 6222
(9); 3436
499
Davis
13-23
(lb); 3002
(5b); 2637
(9); 7742
(5b); 3079
(9);
(11); 6616
de Vries
(9); 55017
Deam
(10); 7094
(13); 7178
11437
13854
(8); 13908
(13); 6774
(9); 5074
(9); 7431
(5a); 12510
(9); 13963
(9); 9897
(8);
(9); 13699
(8);
(13); 14968
15384
(9); 17234
(Sa); 17449
17623
(9); 18155
(9); 20307
(Sa); 21016
21392
(9); 24049
(9); 29237
(9); 30223
31856
(Sa); 34587
(9); 49493
(9); 49525
(Sa); 50634
(Sa); 50752
50836
(9); 50940
51015
(13); 51308
51642
(9); 52251
(9);
(Sa); 17552
(Sa);
(9);
(8);
(13); 37829
(Sa); 37501
(Sa); 40003
(Sa); 39080
(9); 41437
(9); 49832
(Sa); 49446
(9); 49866
(Sa); 50756
(Sa); 50945
(9); 51393
(Sa); 52523
(Sa);
(9); 51013
(Sa);
(9); 51506
(9);
(Sa); 52584
(9);
52585 (Sa);52599 (Sa);52638 (9); 52814 (9);
52841
(9); 53109
55322
(Sa); 55348
(9); 53224
(9); 54432
(9);
(9); 55432a
(9); 55423a
(5a); 55709
(9); 55809
(9); 55899
(5a); 56530
(5a); 56759
(9); 57128
(9); 60140
(Sa); 63030
(9); 63745 (9).
1 (13).
Dean
Dearness
Deaver
& Degener
& Peiler
553
Demaree
8233
Dieterlen
(10, pro parte); 633
633
Dixon
(Sb); 1581
1209
12320
Donley
107 (9); 108 (13).
(9).
Donnelly 20 (lb).
603
Doppelbauer
Dore & Gorham
Dorio
557
Dorr
280
Dorn
3314
45885
(9).
(Sa).
Dorsett
& Morse
Doucet
& Beaulieu
1434
Dreisbach
7317
(13); 1886
(9);
2028
du Plessis
(9); 2579
250
(9); 20526
(9);
Dubois
(9); 22394
(Sa);
Duffie
(9); 23750
(9); 23963
(9); 24774
(9);
Dugas
302 (la).
(9); 25679
(9); 25806
(9); 26519
(Sa);
Dugle
310
28567
(9); 31379
(9); 31509
(9); 36085
(9);
Dugle
& Dugle
37726
(Sa); 38237
(9);
(9);
(9);
(9); 332
Duke
239
(9); 327
54368 (9); 54822 (9); 56795 (13); 56934 (9);
Dumais
& Watson
(9); 61003
DuMond
1290
(9);
(9).
(13); 1648
(10).
(Sb); 4409
(Sb).
Duiben 2582 (9).
350
(9); 61345
(9); 5250
(Sb).
4392
Dumais
(9); 58692
(Sa); 4428
(9 x 10); 1647
1646
22644
(9); 46774
(9);
9 (9); 119 (9).
25237
(9); 54194
(13); 8686
(13); 11463RMD
(2).
(9); 20405
(9); 46340
(13);
(9); 5238
(9).
(9); 21566
(9); 52971
(13); 2507
(11); 4959
(13); 8502
(9); 10271
(9); 21527
(lb); 42914
(8); 2287
(9); 4600
(13); 7277
(9); 9813
(9); 20339
(9); 41869
(9).
147 (9).
(13); 3673
1689
Drinker
(9); 40428
(13).
(9).
(13).
(5b).
(9); 11701
(9).
Dore 22636 (9).
(10).
(9); 10911
(9, pro parte); 639
(5b).
Domes
(8); 6848
(5a); 16785
(9).
(9).
Dietrich 4711 (10).
18367
(9); 51591
(9, pro parte).
Diehl 6916 (13).
Dieterle 1927 (9).
20648
51110
(13);
(lb).
545
& Huber
Dickinson
Dubitsky 8210 (12).
41966
(13); 1149
(13).
Dewey 367 (12).
Dickinson 612 (13).
8791
(9).
(9); 9423
(13); 1194
(lb); 0461
13650 (9); 13776 (9); 14115 (9); 15503 (9);
56975
(13);
(13);
(9).
(13); 1087
(10, pro parte); 6117
6117
2585
34678
16189
(13); 1182
85 (9).
(lb).
Degener
29905
(13); 702
0357
Degener 36506 (10).
Degener
658
Dewart
Downs
Delaunay
(13); 1193
(13); 1866
(Sa).
Devine
71 (9); 127 (9).
6465
(9); 1173
(13); 1497
Desselle 32 (9).
Devenish 633 (2).
(Sb).
(5a); 2544
(Sa); 11962
& McGinnis
Desplantes
(9).
(5b); 2087
71 (9); 2462
689
DeSelm
1162
(8).
239
DeSelm
(13).
73 (9).
De Pue 464
(13); 1154
(13); 1439
Desmarais
(9).
3084
9038
(9); 1141
105 (9); 159 (5a); 401
(lb);
& Davis
Dawson
(9).
1879 (13).
(9).
4777
(9).
2619
658
1400
(9); 2424
Davis
(12); 673
Deschamps
(Sb).
Davies
(Sa);
(9).
Denike
Davidson
(9); 72303
(9); 68865
73981 (9).
(10).
D'Arcy 2209 (8).
Dausen 5060 (9).
Day
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
214
(9).
(12); 424
(8).
(9); 5821
1785
(Sb).
(Sb); 5827
(Sb).
215
OENOTHERA
1997
Dunbar
83 (9).
Duncan
10319
Dunn
Fernald & Long
(9); 21485
57 (9); 4423
(9); 21991
(9).
Duthie 4032 (10).
Dutilly & Ernest 13846 (9).
& Lepage
Dutilly
10458
Eames
Earle
(8); 10481
10475
& Wiegand
10497
(5a); 10498
1747
(12); 10499
7833
(9).
(13); 10486
(9);
(12).
810
Eastham
4505
(5a); 1064
Eaton
(5b); 4509
(5b); 4508
(Sb);
12382
(12, pro parte); 7834a
parte);
10058
Fink 200
40-270
(13); 1256
(9); 2606
(13); 6383
(9).
Ehrle
2725
1315
(9); 3189
(Sa).
(9); R. 4291
(Sb).
680
Fodor 107 (Sb).
(13).
7901
et al. 19965
Elias
Ford 3548 (9).
Forest 16922 (9).
(8).
(9).
Elliot 70 (5b).
137 (5b); 159 (5a); 797
Elmer
495
(5b); 3819
Emig
179 (9).
Forest
(9).
424
Erskine
1267
(9).
(12); 1267a
(9); 1338
(12); 1497
(9);
1497a (13); 51521 (13); 52731 (13); 531088
(13); 55748 (13).
Eskew
1380
1214
Evans
(10).
(8); 1215
(8); 1216
(8); 1217
(9); 2118
Evers
336
Ewan
14514
7442
517
23 (13).
(9); 5752
(9).
2391
& Vickers
578
700
Fernald
Freeland
F-444
5267
(13).
(2).
3938
218 (lb); 220
1919
7835
(5a).
(5a).
7599
Fritsch
26 (9).
(Sa).
428
(13); 18843
17333
(8); 2233
(8); 3325a
(9);
(13).
(9); 16147
(9).
Frye 734
(11); 1062c
(9); 4515
1/2 (12);
Fu 1006
(9); 17089
(9).
(5a).
(5a).
376
Fuller
(9).
10445
202
(5a).
(9); 251
(9); 313
Fuller & Fisher
204
Fuller & Fuller
141 (Sa); 375
328 (13).
2585
Gandoger
(5a).
(5a).
(9).
Frost 346
Galpin
(Sa).
(3).
103 (5b).
Friesner
Gallo
(5a).
5353
(9).
Fukuoka
(13).
1151 (5a); 3518
Ferguson
& Bibbey
Fuckel
Fassett & Wilson
Fendler
(9); 721
(lb);
Fryxell 1098 (2).
2 (10); 16 (10).
(5a).
7252
Fender
(5a).
(9).
(13); 1992
195d
7431
Fay
400
Frankton
Freylag
Farrooq & Ayyaz
Faurie
(lb); 5136
7 (lb); 53 (lb).
Fox & Tiemey
Frere 292
Fabris & Marchionni
Fassett
4774
Fourcade2048 (10).
Frees & Ramsey
(9).
5548
Farwell
(13).
12431 (9).
(5a).
Faber
Faircloth
Foster & Foster
(lb).
(9).
Ewer
Fabius
et al. 11604
Freer 81 (9 x 5b); 1662
(8).
Everett & Johnson
(9).
(Sa); 21567
(9).
Fox & Weedum
(2).
622
Espinosa
(9); 15282
Forrester02243 (Sa).
Fosberg 14459 (13); 15953 (9); 17161 (9); 17163
(13); 17355 (13); 17593 (9).
Foster 5333 (3).
(lb).
Epling 5499 (9).
Erlanson
(Sb); R.
(Sb).
Fogg
Ellis
(lb); R. 4824
et al. 194 (Sb).
Flowers
Fogg
1522
61 (5a).
& Fitzpatrick
695.5
Fletcher
Eilers 2354 (9); 2458 (13).
2880
(2).
3478
110 (9).
Flodman
(12); 7946
(5a); 4137
(12); 1850
Ehrle & Wahl
(12);
5420 (5b).
(9).
1646
Ehlers
(9 x 12, pro
(9); 21996
(13); 21995
(12);
(9, pro parte);
(9).
Fitzpatrick
D2
Eggleston 10200 (3); 15412 (Sa).
Egler
(12); 6844
(9); 7832
(12); 7834a
(12); 7834
Fishman
(lb).
(9).
5327
(12); 5391
(9); 7830
(9); 7829
7834a
Fiori
(lb);
839 (1c); 7917
Edmondson
(9).
(12).
(12); 1885
Ferris & Duncan
(13).
(5b); 4507
4510b (5b);4515 (5b).
Eastwood
(12); 11135
24212 (10).
(9).
Easterly
et al. 337
(8); 7828
(8); 10492
(9); 11133
(13).
5930
Fernald & Wiegand
Femald
(13); 10488
445
Fernald & Strong
Duvigneaud 813 (13).
Dzubin 52 (5b).
Eames
(8); 13707
(12).
Fernald & St. John 7827
(13).
15308
(9); 9604
(9); 7551
339
(8); 14197 (13); 14198 (9); 14202 (12); 14203
(9).
Garcia
550
(2).
399
(la).
(9).
(5a); 317 (9); 3561
(9).
(9).
(4).
Gardner
2 (10); 407
107.35 (13); 108.35 (9); 109.35 (9); 110.34
(13); 111.34 (13); 112.34 (13); 112.35 (9);
113.34 (9); 114.34 (9); 114.35 (9); 116.34 (9);
117.34 (9); 117.35 (9); 119.34 (9); 124.34 (9);
133.35 (12); 134.35 (12); 136.35 (12); 137.35
(12); 14134 (5a);67135 (12).
(Sb).
Garidi 8089 (9).
Garlitz
(12); 774
725
(9).
Garrett5316 (lb); 7640 (lb).
Garton
(9); 2037
1121
(12); 7354
9468
(9); 4308
(5a); 7864
(9); 9612
(13); 5234
(9); 8541
(9); 6655
(5a); 8698
(12); 12040
(12); 10425
(12);
(9);
15103 (12).
Garton
Gates
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
216
et al. 14326
(9); 4.34
(12); 2.34
(13); 9.33
(9); 3.33
(13); 5.34
(9); 5.33
(9); 9.34
(9); 6.34
(9); 10.33
(9); 7.33
(13); 10.34
15.33
(9); 15.35
(12); 16.33
(9); 17.33
(9);
18.33
(9); 19.33
(12); 20.33
(12); 20.34
(9);
(12);
21.33 (9); 21.34 (13); 22.33 (12); 23.33 (12);
23.34 (13); 25.33 (13); 26.33 (13); 26.34 (13);
27.37
(13); 28.33
(13); 29.33
(13); 29.34
30.33
(12); 31.33
(13); 31.34
(9); 32.33
32.34
34.34
35.35
37.34
(9); 33.33 (12); 33.34 (13); 34.33 (13);
(13); 34.35 (13); 35.33 (12); 35.34 (13);
(13); 36.33 (12); 36.34 (9); 37.33 (13);
(9); 38.33 (13); 38.34 (9); 38.35 (9);
39.34
(9); 39.35
(9); 43.14
(9); 40.34
(9); 42.33
(12); 43.33
(9); 40.35
(9); 41.34
(12); 42.35
(12); 42.34
(9); 44.33
(13);
(12);
(13); 44.34
(9); 44.35 (13); 45.33 (13); 45.34 (12); 45.35
(9); 46.33
(13); 47.35
(13); 46.34
(12); 48.33
(12); 47.33
(13); 47.34
(13); 48.34
(13); 48.35
(12); 49.33 (9); 49.34 (13); 49.35 (12); 50.34
(13); 50.35 (13); 51.34 (13); 51.35 (9); 52.34
(13); 53.34 (12); 54.34 (13); 54.35 (13); 55.34
(12); 55.35 (13); 56.33 (9); 56.34 (12); 57.33
(9); 57.34 (13); 57.35 (13); 58.33 (9); 58.34
(12); 58.35.13 (9); 59.33 (9); 59.34 (12); 59.35
(9); 60.34
(13); 63.33
5804
(13); 61.34
(9); 63.35
(13); 61.35
(9); 64.35
65.33
(13); 65.34
(12); 65.35
66.33
(13); 66.34
(9); 66.35
67.33
(12); 67.35
(12); 68.33
(9); 64.36
(9); 66.36
(9);
(9);
(9);
(12); 68.34
1591 (10); 1794
H. S. 3475
Gentry,
J. L. 588
Gentry,
J. L. & Jensen 2207
(lb); 6177
Gerard
511 (12); 512
Gershoy
Gillett
6496
Gillett
& Graham
(5a); 6848
Glaziou
2568
Glowenke
7464
(9); 71837
69784
(13); 9512
(12);
11559
(7).
(lb);
73117
(9); 76808
(7);
(lb);
(Sb).
76815 (7).
Godfrey
& Tryon
Godron
438
466
1613
(13); 467
(13); 466a
1394
Goldblatt
(9).
(12).
(9); 1435
(13); 72.34
(13); 72.33
(Sa);
Gonzalo
5066
74.35
78.33 (13); 79.35 (13); 80.34 (9); 80.35 (13);
Goodrich 15262 (lb).
Gordon 245147 (9).
80.39
(12); 81.34
Gordon-Gray
82.35
(9); 83.34
88.33
(13); 88.34
(13); 88.35
89.34
(13); 89.35
(9); 90.33
(9); 90.34
271
(Sb); 271
407
(Sa); 1244
et al. 451316
Graham
10 (lb);
28 (9); 266
(12); G1517
(12);
Grandtner
14171
& Bibbey
90.35
(9); 91.34
(12); 91.35
(9); 92.33
(9);
Grankton
93.33
(9); 93.35
(13); 94.33
(9); 94.34
(12);
Grant, M. L. 476
94.35
(13); 95.34
(13); 95.35
(12); 96.34
(Sa);
Grant
96.35
(12); 97.34
(Sa); 97.35
(12); 98.34
(Sa);
Grant & Fosberg
98.35
(13); 99.34
(13); 99.35
(Sa); 100.34
(12).
428
(9); 9961
(13).
(Sb).
(13).
(9).
Grant & Schneider
Grassl
173 (lb);
Graves
667
1744
(10).
Gorham
3049
(9).
(13).
(10).
Goodding
100.35 (13); 101.35 (Sa); 102.35 (13); 103.34
(13); 105.35 (13); 106.35 (13); 107.34 (13);
(10).
(9); 7881
(13); 71.35
(Sa);
(9).
Goddard473 (Sb).
Goddijn 663 (9).
Godfrey 2139 (9); 52462 (9); 58330 (7); 65863 (7);
73.33 (Sa);73.34 (9); 73.35 (13); 74.33 (13);
(13);
(13); 14228
(9).
(10); 8343
71.34
(9); 89.33
(Sa).
Gleason 1489 (13).
25 (9).
(9);
(8); 10500
Gillis 5414 (9).
Gilson 35 (12).
1102 (lb).
(13);
(9); 7940
9798
Goncalves
(13); 82.34
(12); 10468
Getty 263 (9).
Gierisch 973 (ib).
Gonzales
(9); 87.33
(13).
7 (lb).
& Corey
Gessel
(13);
(9); 81.35
(lb).
(9); 1175 (13); 2734
787
(13); 71.33
(9); 85.34
(lb).
Gerardin5168 (13).
Gereau 899 (12).
(13); 70.35
(9);
(10).
George 33 (9).
Gephardt910 (lb).
(12); 70.34
(12); 77.35
(10,
(Sa).
68.35 (12); 69.33 (13); 69.34 (13); 69.35 (13);
(Sa); 76.35
(9); 1942
(la).
Gentry,
70.33
(13); 75.33
(6); 1803
(9 x 10, pro parte); 3069
1942
Gentry, A. 5858
Gogo
(13);
(9).
(5a).
Gelsii 114 (10).
(13); 62.35
(9); 65.36
(13); 569
pro parte);
(13); 3.34
12.33 (9); 13.33 (13); 13.35 (13); 14.33 (13);
(12); 41.35
457
Gavelle
Geerinck
(12).
1.34 (9); 2.33
Gauthier
9339
(10).
8074
3517
(10); 1859
(5b).
(13); 4008
(5a); 1918
(13).
(9).
(3).
Greear 63261 (8).
Harton
179 (12); 379
Green
(9); 3457
(9); 1700
(12);
3521 (13).
Greer
1674
Gregory
460
Grether
6352
Hatschbach43526 (9).
(9).
(13).
Hatschbach
& Ramamoorthy
(9).
Hatsuyama
592
(9); 8184
(12); 6557
Hattum
(5a).
Gretz 10642 (5b).
Grierson
& Long
2702
Hausmann
(9); 865 (5b); 1161 (13); 1445
Groh 49 (12); 695.5
(9); 5224
(9); 2611
55 (9); 75333
Guan
(13).
93 (9).
Hayden
2126
87 (9).
Gutte & Jentsch
31808
Haddow
303
(9); 400
Hainault
6740
(10).
42 (9); 2639
(5b); 9029
179 (12).
& Lehto
& Brisson
53/362
203
728
(3).
(13); 14816
(9);
14839
(13); 15065
Hanes
900
(9).
Heller
5704
Heller
& Bach
(1c); 10616
Heller
& Heller
Heller
& Kennedy
548
(lb).
(8).
37978
(lb).
8861
(lb).
(9); 62-980
61-618
& Swisher
Henderson
(la).
(2).
(Sa); 950
(13); 940
Henson
(9).
01 (10); 45 (10); 108 (10); 231
35 (Sb).
(9 x 10).
(9); 1277
992
Hepburn
264
(10).
472
(12).
(9); 1878
Hansen
& Hansen
363
(12).
Hermann
Hansen
& Merkle
292
(5b).
Hertel 7945 (13).
Hansen
et al. 4479
Heruck
& Martins
162 (13).
Hanson
740
(13).
Hesler
et al. 2278
(Sa).
(12).
(3); 1043
et al. 6306586
Hardham
Hardin
3713
13763
13767
(9 x 10).
(1c); 3907
(9); 13764
(9); 13768
Hess
(10).
(8); 13765
(8); 13769
(8); 13766
(8); 13770
(9);
(8);
13771 (9).
Harding
540
(5a).
Harlow
1422
(12).
Harmon
Harms
17764
1810
Harper
(5a); 2992
(5a); 16999
(9); 17896
(9); 18249
(9).
Harrison
053
9390
Hartley
(7); 3976
(7); 3982
(5b);
(9); 4522
Hartman
(12).
(3); 118 (9); 1272
(lb); 4883
(9); 1906
(5a).
Heurck
& Martins
Hibon
1460-2
162 (13).
(10, pro parte);
& Bartlay
1807
Hill
1460-2
(9 x 10, pro
11490
(2).
(lb);
248
(13); 8043
126 (Sb); 887 (lb);
(13).
899 (lb).
41 (11).
(9); 10912
51 (9); 9388
(lb);
(13).
(9).
Hillsdon 145 (13).
Hinckley 1902 (lb).
Hitchcock,
A. S. 627
Hitchcock,
C. L. 2419
(5b); 687
(lb); 21350
(5b); 15812
(10).
(5b); 17057
(5b);
19510 (4); 20363 (5b).
Hitchcock,
(5b).
1674
379
Hillier
(9).
Hamfis 153 (8); 20977
(3).
Hetzer
Hiebs
(5a); 2738
(8); 2999
(lb).
729
Higgins
7 (Sb).
Harner
1321
Hester
(9).
parte).
15 (9).
1105
(9); 66
Hendley 679 (9).
Hendricks 505 (9).
Henrichs 463 (lb).
Hansen
Hara
(9); 63-1696
1013 (8); 67-1561 (9).
(Sb).
1044
(9); 2135
Heitlinger 327 (Sa).
Henderson
(13); 18275
142 (Sb).
Hammond
Hanekom
(9); 77/36
Hemphill 265 (12).
et al. 692
Hamilton
(10); 59/170
(10); 57/21
2134
(13); 18275 (9).
Hammel
(5b).
45 (9).
Hein
(12); 8695
C66129 (13).
Hamel
1684
Healy
Hecker
59 (13); 127 (12); 12536
Hamel
(Sa);
(9);
Heidecker 131 (Sa).
(13).
(lb); 11573 (5b).
Halvorson
(9); 10101
(10); 80/16 (10).
(lb); 3436
Hall & Hagenah
Head
(13).
67 (12); 92 (13).
Haber
(Sa); 7682a
(Sa); 3622
(5a); 8656
11498 (9).
Gunderson 543 (13).
Gustitus
(5a); 3621
(9); 8655
8654
(10).
(5a).
(13).
(9).
702
Hawyer
47 (5b).
Guess
& Sako 27019
611 (9); 707
Haught
(9).
(9 x 10).
37995
(9).
541 (9).
Hatusima
108 (9).
Grimm
11 (13); 38 (9).
Hasselbring
3736
& Greenman
Greenman
(8).
7864
Hasem 398 (9).
Hashimoto 5607 (10).
(10).
15 (5a); 655
Greenman
Hall
217
OENOTHERA
1997
5695
Hitchcock,
C. L. & Martin
5343
(lb); 5469
(lb);
(lb).
C. L. & Muhlick
9934
(Sb); 10411
(lb);
50
MONOGRAPHSVOLUME
BOTANY
SYSTEMATIC
218
11457 (5b); 13212 (5b); 13530 (5b); 13644
(Sb); 14735 (Sb);21715 (5b).
Hoch
422
(lb); 426
(5b); 427
(5b); 926
Jarman 110 (9).
(12); 4484
Jehli'k 2773
(10); 6241
(5b); 1365
(9); 5354
(9); 6256
(Sa); 6230
(9 x 12); 6263
(9 x 12); 6640
1855b (4).
Hodgdon 2743 (9).
6650
(9); 6661
128 (10).
6746
(9); 1853
(4); 1843
Hofmeyer
Holland
204
(4); 1855
(4); 1855a
(4);
565
Holmer
(Sb).
250
(9); 267
& Reveal
Holmgren
1333
et al. 16088
2170
(lb); 39-18
(lb); 7193
(9).
Homer
3687
Horton
334
Hou
(9).
76-865
House
2733
(13); 1909
(13); 76-1109
Jones 404
(9).
(12).
Hudson
22857
(5b); 2690
510
4040
Kaul
(la).
1574
111 (13).
& Tuomikoski
Keck
Hutchins 1530 (5b).
9 (13).
Hymowitz 404 (9).
Hyypio 835 (9).
In-Cho 7551
(12); 14434
(12); 14591
(13);
Jackson
Jacobsen
(13); 8115
(9); 7657
(9).
(9); 7840
(9).
73 (9); 913
520
& Svendsen
(9); 3049
(Sb).
39 (9 x 10).
(4).
(11).
(13).
(9); 2462
(5a); 2439
(9).
161 (9).
76 (13); 152 (9).
Kellogg
194 (5b); 247
Kempers
117 (9).
Kennedy
5 (13); 4102
206
(5a).
(13); 12206 (13).
12194
(10).
(9); 3204
(9); 7544
(9); 7665
2 (9).
Khan
et al. 1018
15021
10270
(12); 19449
Khan
Kiener
(lb); 4262
(Sb).
(9).
Kern & Reichgelt
509
(Sb).
& Jordan 77 (12).
Kern 5063
184 (9).
James 263
(13); 36034
(9).
Keever
Kelsey
15 (9).
Jacobs & Ranzinger
(9); 23700
(9).
(5b); 5613
2449
Kelsey
(9).
Ito & Kinoshita
(5a);
Keinholz 151 (9).
15833 (9); 28457 (10).
Iltis et al. 7405
816
Keller
7111
(lb); 5624
(9); 12685
(12).
Keener
Keil
(9).
Iltis & Noamesi
(9); 1075
(9); 9248
(5b).
725
(5a); 8264
(9); 1118 (10); 1140
(9).
(5b); 8614
& Rostaiiski
Kearny
(8).
508
(10).
Kaspiew 1669 (10).
(5b).
Hunt
Iltis 6848
(1c);
(13).
797
Ikegami
14496
Jurkoeskij752 (9).
(9).
Kappus
& Bonpland
Hyduke
(9); 2823
(9); 22858
Kayser
Hustich
(lb);
14176 (7); 15345 (7); 18512 (Sb);22237 (9);
Hunnewell 19038 (11).
212c
(13); 15003
Jozurk 179 (5b).
192 (5a); 375
Humboldt
8554
Joyal 1304
(12).
& Spence
(10); 14492
221 (5a); 454
Jones & Jones 9534
115 (la).
Hulbert
(9); 14518
(9); 829 (5a); 878
(lb);
HubrichtB2625 (9).
Huhn
(9);
Jolicceur3493 (13).
(13).
(13); 24203
(9).
15-41
2309
team 1438
149 (lb);
(10); 2032
Howell 496 (7); 21254 (1c); 23027 (1c); 28197 (Sb).
Howitt 1599 (1c).
Huber
(Sb); 1899
(9).
(9).
(5b); 4156
survey
Johnson
1284
(9);
14497 (10); 19458 (lb).
(12); 1912
(10).
& Long
Hsi 586
et al. 3919
Jenson 253
(12); 1715
(9); 7553
(9); 6647
(9).
(9).
Houle
(13); 8287
(9).
Jinfushan
10411
Howe
(9); 3371
3321
(5a).
(9).
6991
(9 x 5a).
(9); 6645
Jenkins & Ilman 3280
Jenkins
(9).
Hosomi
(13); 6795
(5a); 7126
(9 x 5a).
2786
Jennings & Jennings
(5b).
et al. 1712
Hosie
(9); 6782
(9 x 10).
Jenkins & Bayly
(9).
(5a); 4176
(5a);
(9); 6829
29 (5b); 1005 (Sb); 1601
3249
(5a).
R181B
Hoff
Jenkins
(13).
42 (9).
Horie
(9); 7214
7128
Jehlf'k et al. 6676
(5b).
1304
8 (2); 4782
Hope
Hopkins
(9 x 10); 7131
(9); 6736
6730 (5a).
(lb).
1192 (5a); 39-117k
717 (lb);
(10); 6826
Jehlik & Rostainski
Holmgren
Hoover
(9 x 12); 6773
(9);
(10); 6681
(13 x Sa); 6754
(5a); 6750
(9 x 5a); 6814
Jehllk & Krcilova
(8).
Holmgren 6166 (lb).
Hood
(5a); 6747
(13); 6757
(5a).
(9); 6649
(9); 6669
(9 x Sa); 6714
(9 x 10); 6688
Hollister 79 (9).
Holm
(9); 6659
(9); 6236
(9 x 12); 6261
(Sa); 6260
(la).
(Sa).
(12); 12182
(9 x 10).
(Sa); 12183
(9);
Lawson
& Goodman
Kildahl
42 (5a).
Lawson
& Musselman
Kildale
6057
Lazor 4798 (9).
et al. 6166
Kiesling
Killick
(4).
179 (9); 264
Kisha
3673
(10).
Leadley
(9); 462
(10).
Leal
1190
Klein
(5a).
et al. 34 (9).
Kluhsmann
Knight
& Knight
Knowlton
495
Koch
4355
Koelz
7352
(9 x 10); 13494
(10).
118 (8).
(9 x 5a); 1064
910
Koziol
Krajina
496
1320
(Sb); 903
(9); 3372
(9 x Sa).
5481
(9); 32943
(9); 36857
(8); 47557
(9); 5482
Kuhlman
50 (12).
Kunming
herbarium
Kunz
264
Kurtto
(9); 5683
(12); 9252
(12);
& Vallee
(7).
224
(4).
Leuze & Doppelbauer
13315
& Lassier
Lichvar
644
(Sa).
110 (2); 7445
team 206
(10).
Lindayen
Linderud 114 (5a).
Lindheimer502 (2); 808 (2).
(5a).
Lindsay
La Follette
Lingenfelter736 (Sb).
Lisil 60 (13).
170 (13).
A33
Lakela
(9).
120 (9).
Laferrere
Little
351 (5b); 9194
Lakela & Davidson
Lambert
(12); 12123
21690
Lammers
474
Lamoureaux
& Durand
346
Lancucka
1085
(5a).
71-43-35
& Sherff
3303
(Sb).
Larsen
116 (Sa); 207
80 (9); 715
Larsen
et al. 10460
LaRue
108 (9).
1902
Laurent
33 (9).
Lawalree
720
(9).
(Sb); 1217
388
(9); 1237
(12).
(Sa); 4763
(10); 1066
(13); 6763
(8);
Louis
764
(9).
(12, pro parte).
(10).
1039
(9); 1065
(9); 1237
24341 (9); 40117 (13).
(9).
1378
(9).
Louis-Marie
& Lamarre
341 (9).
Louis-Marie
et al. 1026
(13).
Louw
(5a).
1375
(2).
Love & Love
5133
(9).
Lovett 1569 (9).
(9); 1303
(5a); 1627
(12); 2529
(9 x
(10); 8010
(9); 8032
10); 8033
(10); 8078
(9); 11018
(10); 12499
(9 x
(9); 12863 (13); 12864 (13); 13112 (10); 13185
(9 x 10); 15859
(13); 16628
(13); 18225
2145
571
Louis-Marie
(13); 7779
Lawrence
(9); 616
Loots
(9).
10); 4640
17972
2-3-13
Louis-Arsene 6627 (10).
& Hardwick
Lathrop
(Sa).
Long & St. John 7834
(9).
Larsen & Pedersen
(Sa).
13353 (9); 28860 (9); 38267 (9); 58711 (10);
60077 (13); 69573 (10); 75610 (10).
(lb).
Lansing
235
Long
(12).
Lapage
Latartaine
(13).
(9).
3990
(9); 269
(9).
Lloyd 3015 (3).
Loefgren 11700 (10); 11898 (10).
Lombard4097 (9).
Lane & Letho MAL1607
Langenheim
264
Liwhe
(13).
(9); 2582
137 (3).
Liu 1573
(5a).
364
Lane
(13).
82 (13).
Lamoureaux
(10).
139 (12); 160 (Sa).
Kytovuori 10216 (9).
Lacassee
(10).
98 (9).
Lewis 1006 (9).
Liebenberg
(Sa); 4419
188 (13).
et al. 2535
Levasseur
(9).
4281
(5a).
Lemkow 24 (13).
Lemon 800 (13).
Leonard
135 (9).
& Tardif
(5a); 14024
(9).
Lester & Yearout
9253 (13); 10736 (9).
Kruckeberg3216 (lb).
Kucryniak
195 (10).
156 (9).
Lemieux
(Sb).
(7).
Krivda 1402 (5b); 1800 (5b).
Krotkov
(2).
Leiberg 1520 (Sb); 1558 (Sb).
Lelong 8156 (Sa).
Lemieux 286.1998 (12); 286.2111 (13); 286.2234
(9); 286.2577 (13); 14379 (13).
(13).
(Sa).
Komarek
(5a).
534
(5a); 8536
Lehr 496
(10).
(13); 30502
30501
600
Leclerc 57 (13).
Leendertz 11072 (2); 11504 (9).
Leenhouts 2834 (9).
(12).
1129
& Petty
1204
Leary
59 (9).
Klawe
Kral
(Sa).
& Vahrmeyer
King
219
OENOTHERA
1997
(Sb).
(13).
(10); 16675
(10);
788
Lowry
(4).
Lucian
107 (8).
Ludlow
& Sherriff
8208
(10).
Lundell 8307 (13).
Luteyn
Lutz
Lyche
2791
1251
(9).
(9).
15477
(Sa); 29269
(Sa).
(9); 24149
(9);
50
MONOGRAPHSVOLUME
BOTANY
SYSTEMATIC
220
Mabbott
280
136 (13); 137 (12); 139 (12); 143 (13); 144 (9);
(5b).
262 (lb); 467
Macbride
27 (5a); 4633
MacDaniels
Mackay
(5b); 473
733
(lb);
146 (9); 150 (13); 151 (13); 152 (13); 189 (13);
(5b).
1249
(8).
(11).
373
Macklin
47 (13).
Macoun
234
(13); 248
(9); 800
& Rouleau
Marie-Victorin
et al. 2004
(13); 9017
(5b); 9018
(Sb); 9028
Marloth
(5b); 9040
(5b); 9042
(5b); 10644
Marold
(5a); 19125
(9); 12884
(9); 21193
(13); 21704
(9); 44467
(9); 44468
(9); 44464
(9); 60320
(9); 19126
(9); 44465
(13); 85925
(9); 80585
(13); 85927
(13).
& Herriot
Macoun
MacSwain
72378
59-147
(Sb).
(Sb); 72379
(lb).
Madalski 1605 (9); 2558 (9); 23052 (9); 23888 (9);
24893 (9); 25504 (9).
Magin
1053
4611
Maguire
& Muenscher
(9).
10633
(10).
96 (12); 98 (12).
Maher
Mahler
2083
Mahler
& Mahler
Mahurin
4576
Malte
& Watson
Malter
(13); 717
52979
Manning
209
Margolin
(9); 773/29
1218
(Sb); 1748
(9); 303129
(Sb); 2680
(13).
(lb).
83 (12); 934
(9); 10803
(9); 3163
(9); 10804
15882
(12); 15883
28667
(9); 60020
Marie-Victorin
(13); 3703
(13).
737
(9).
(5a); 764
263
(2); 975
& Poulin
92 (9).
& Erlanson
62 (11).
Martin
& Magnier
523
Martz
90 (9).
Marugama
654
1048
(9); 4187
(9).
134 (13).
Martin
Masson
(5a).
(13); 1728
(5b); 1321
(9).
(9).
(9).
45 (la).
Mathias
699
(9).
Mauritz
843
(9); 1724
(5a).
17 (9).
104 (5b).
Mazzeo
2327
McCalla
477
9656
(12).
(9); 8393
(5b); 10223
(5b); 9212
(5b); 8396
(5b); 10448
(5b);
(5b); 10496
(5b);
11181 (5b).
Marie-Anselm 114 (13).
8326
(9).
(5a); 2392
Martin
(9).
73 (13); 158 (9).
Marie-Victorin
476
Martel
(Sb).
92 (10).
Maqsordand
Marsh
Mayle
(9).
137 (lb).
468/22
49541
173 (9).
Maxfield
(Sb).
Malte
Marschner
Mathewson
Magrath
(9);
Mastrogiuseppe 2618 (Sb).
(9).
(9); 1054
86 (13); 8287
& Clausen
Marshall
(9); 64604
(13); 78527
(Sb); 67933
(9); 85924
(9); 85894
(13);
(9); 33982
(13); 33825 (13);40128 (13); 45686 (9).
(5b); 9014
(5a); 4953
(13); 9039
(5b); 64605
(12); 27158
(9); 33825
Marie-Victorin
(13); 9016
(Sb); 10815
(9); 17715
(12); 28321
49649 (12).
MacKeever 578 (9).
Mackenzie
(9); 10806
28180
(lb).
6C-236
(9); 28321
(9); 3171
(9); 15881
(13);
(13);
(13); 28478
(9);
(13).
& Rolland-Germain
1 (13); 2 (13); 3
(13); 6 (13); 7 (13); 8 (9); 9 (13); 12 (13); 13
McCauley
579
McDaniel
14583
(9).
McDonald
39 (9); 286
McDougal
1733
(7).
(9); 3037
(5a); 5397
(9).
(9).
McElvaine 206 (10).
McFarland
215
McFarland
& Anderson
McFarland
& James 20 (5a).
McFarlin
(9).
6119
McFerson
215
(7); 6585
(13).
(9).
3 (Sb).
& Letho L20582
(13); 15 (9); 16 (13); 18 (13); 19 (9); 25 (9); 28
McGill
(9); 29 (9); 32 (13); 34 (13); 35 (9); 36 (9); 37
(9); 38 (13); 39 (13); 40 (9); 41 (9); 42 (13); 44
McGregor
292
13369
(9); 45 (12); 46 (9); 51 (13); 52 (13); 53 (13);
(lb);
54 (9); 55 (9); 56 (13); 57 (9); 59 (9); 60 (9);
17084
(lb).
(5a); 4386
(lb);
14992
13522
(5a); 5022
(Sa); 14153
(5a); 15522
(5a); 18679
(5a); 5185
(Sa); 16128
(5a); 20033
(Sa);
(5b); 20274
61 (9); 62 (9); 63 (9); 64 (9); 65 (9); 66 (9); 67
(5b); 23455
(12); 68 (9); 70 (9); 71 (9); 73 (9); 74 (9); 75
(Sa);27587 (lb); 27608 (lb); 27622 (lb);
(9); 77 (9); 78 (9); 79 (13); 80 (13); 81 (9); 82
27661
(9); 83 (9); 84 (13); 85 (13); 86 (9); 87 (9); 89
(lb);
(12); 93 (9); 94 (9); 95 (9); 96 (Sa); 97 (9); 98
(9); 99 (9); 100 (9); 101 (9); 102 (9); 103 (9);
30454
104 (9); 106 (13); 107 (9); 108 (9); 109 (13);
30642
(lb); 27673
29527
McGregor
(9); 121 (9); 122 (9); 123 (12); 124 (9); 125
McIver
(lb); 27689
(13); 126 (13); 127 (13); 128 (13); 129 (13);
McKnight 58080701 (lb).
130 (13); 131 (13); 133 (9); 134 (9); 135 (9);
McLaren
V90
(10).
(9);
(5a); 30525
(5a);
(5a); 31889
(9); 39454
(5a); 533
14 (9).
(lb); 28066
(5a); 30628
(Sa); 31855
(9); 31991
& Bare 338
(Sa); 24049
(9); 29729a
(5a); 30470
(5a); 30594
(5a); 31317
(Sa); 31931
(9); 116 (9); 117 (13); 118 (Sa); 119 (Sa); 120
(9); 24024
(5a); 29638a
(5a); 30462
(5a); 30534
110 (9); 111 (9); 112 (13); 113 (9); 114 (9); 115
(9); 23490
(lb);
(Sa); 14759
(5a).
(5b); 612
(5a).
52 (5b).
McMillan
McNary
708
(5b).
McPherson 485b (5b).
McVaugh
9592
(12); 12613
(5a); 12485
(12); 12656
(12).
Means
2257b
Mearns
487
(Sa); 2641
207
Menzel
& Franklin
Moore
& Hsi 23424
Moore
& Huff
Moore
& Moore
Moore
& Ownbey
914
(lb).
(12).
18057
(9).
(9); 11273
(13); 6422
249
22249
et al. 69 (10).
Moreira
& Joly 373
(10).
Morfino 1476 (9).
1737
(13).
1450
Morgan
Morin
174 (9).
Mericle 926 (2).
Morley
& Brayshaw
62248
(9).
(9); 715
430
670
(12).
(5a).
Morrill
1041
(5a); 1132
(5a); 1301
Merrill
1355
(Sa).
Morton
9024
(8); 11121
(Sa); 11690
Merwe
2380
(Sa).
Mosquin 11362 (Sb).
Merriles
(5a).
Merxmuller 68/54 (10).
Moss
& Grau 21218
Merxmuller
Metcalfe
379
(10).
(Sb); 390 (lb).
Mexia
4232
Meyer
97 (12); 484
Moyer
(Sb); 1191
(Sa); 922
694
(Sb); 10597
(Sa); 2580
& Brown
& Curtis
Muenscher
& Lindsey
3448
(9).
Muenscher
& Maguire
2400
(12).
Michel
1504
(9); 601
(Sa); 2188
(9).
(9).
(9); 5841
(9); 2049
1808
Muhlenbach
(10).
191 (Sa).
Muir
3291
(10).
& Bassett
Milan
(30)761
Mulligan
& Woodbury
Miller
117 (9); 548
Munro
(13).
1073
(8); 1076
Munz
(9).
Millspaugh 1063 (9).
Minshall
561
(Sa); 605
(9); 983
1469
(13);
Moffet
(13); 1966
(13); 4781
& Zinck
(9); 861 (13); 955
1050 (9); 1156
(9); 1679
(9); 3700
Minshall
(9); 691
(5a); 1360
(13); 2498
(13);
(9); 2626
(12).
64 (10).
12365
Mogg
Moldenke
(2).
1361
3418
(9); 3222
(1c); 3419
(10); 3416
(1c); 8593
(9); 3417
(10); 10238
(1c);
(9);
18875 (9); 31578 (9).
Moldenke
& Moldenke
6233
(12); 9896
(9); 30998
(9); 31573 (Sa).
1080
(13); 1111 (9).
1828
(Sb); 1974
(Sb).
1137 (12).
12133
(lb);
13008
(Sb); 13013
(lb);
13018
13020
(lb);
(3); 13011
13014
(lb);
(lb);
13015
(Sb); 13021
(lb);
13012
(lb);
13023
(5b); 13024 (5a); 13025 (5b); 13026 (5b);
13027 (5b); 13033 (5b); 13238 (lb); 13260
(lb);
23 (9); 104 (13).
(13).
(12).
(9).
Mulligan
(9).
21934
6359
Midorikawa 1030 (10).
& Maguire
(Sb); 2578
(2).
(Sa).
Muenscher
2646
Miller
(5b); 1870
Muenscher
165 (9).
(5a).
(Sb); 18378
80 (2 x Sa).
Meylan
(5a).
Mrkos 350 (9).
Muenscher 8291 (9); 8292 (9).
(10).
Mgaza
Mick
283
(Sb); 3219
Methews 21/467327 (10).
Meyers
(9).
(9).
Moran 14601 (lb); 21124 (lb); 24474 (lb); 25118
(lb); 26372 (lb); 27857 (lb).
(lb).
(9).
Mennema
Moore
Moraldo
(9).
(12); 3768
Meilleur 10137 (9); 10185 (9).
Meisner 87a/67 (13).
Mell
221
OENOTHERA
1997
13273
(5a); 13276
(5a); 13278
(lb);
13281
13282
13351
13279
(lb);
13357
(8); 13360
(lb);
(9); 13362
(9); 13372
(lb);
(9); 13363
(9);
(9);
13364
(13); 13368
13376
(9); 13378
(12); 13380
(13); 13382
13383
(9); 13384
(13); 13385
(9); 13386
13387
(12); 13388
(9); 13389
(8); 13390
(8);
13391
(13); 13398
(9); 13400
(8); 13402
(8);
13403
(9); 13404
(9); 13405
(12); 13406
(9);
13407
(9); 13428
(5a); 13451
(5a); 13471
(9); 13373
(9);
(9);
(9);
Molina
& Montalvo
21823
Molina
et. al 26654
(la).
13472
(9); 13473
(5a); 13475
(13); 13476
Monachino 11592 (13).
13478
(9); 13480
(13); 13481
(8); 13483
Monro
13484 (11); 13487 (13); 13488 (9); 13489 (11);
(9); 1242
1096
Monroe
9940
Monson
716
2846
Montalvo
(Sa); 765
(Sa); 2929
pro parte); 748
Moodie
Moore
(9).
(12).
& Ackerman
Montgomery
(la).
263
13490
(13); 13493
(8); 13494
(8); 13495
(9);
(5a); 1257
(13);
13496
(13); 13497
(8); 13498
(8); 13501
(9);
(Sa); 3210
(5a); 3610
(5a).
13502
(13); 13507
13511
(9); 13512
747
(4, pro parte); 747
(10); 749
(10,
(10).
(Sb).
31 (13); 133 (13); 384
2540
(13); 4863
(9);
(9);
(Sa); 917
(9).
75 (Sb); 1086
(8);
(5a); 715
(9); 22576
(5b); 900
(5a); 26433
(13).
(9);
(9); 13509
(8); 13514
(8); 13510
(8); 13516
(8);
(8);
13519 (9); 13520 (8); 13521 (9); 13523 (8);
13526 (8); 13527 (9); 13529 (9); 13530 (9);
13531 (9); 13532 (9); 13533 (9); 13536 (13);
13537 (9); 13538 (9); 13539 (9); 13540 (9);
13541 (9); 13542 (9); 13543 (9); 13544 (9);
MONOGRAPHS VOLUME
50
SYSTEMATIC
BOTANY
222
15310 (la); 15311 (lb); 15312 (lb); 15313 (2);
15455 (10); 15568 (1c); 15569 (1c); 16325
13545 (9); 13546 (5a); 13547 (9); 13550 (9);
13551
(5a); 13552
13558
(9); 13560
(9); 13554
(9); 13561
(9); 13555
(Sa);
(9); 13563
16974
(lb);
(9);
17436
(lb);
(13); 17508
(9); 17507
13564 (Sa); 13567 (9); 13568 (Sa); 13569 (9);
(13); 17509 (13); 17510 (9); 17511 (9); 17512
13570
(9); 17513
(Sa); 13575
(Sa); 13573
(2); 13580
(2);
13585 (9); 13904 (lb); 13909 (3); 13910 (lb);
13913 (lb); 13915 (lb); 13921 (lb); 13922
(lb); 13929 (lb); 13956 (lb); 13959 (lb);
13965 (lb); 13968 (lb); 13973 (lb); 13974
(lb);
13977
(4); 13982
13993
(lb);
13994
(lb);
14009
(lb);
(lb); 4031
14030
14035
(lb);
(3);
14039 (lb); 14042 (lb); 14048 (lb); 14049
(lb); 14050 (4); 14057 (lb); 14062 (1c); 14066
(1c); 14080 (10); 14082 (lb); 14097 (lb);
14103
(lb);
14104
(lb);
14128
(8); 14143
(8);
(Sa); 17521
(9); 17522
(12); 17529
(Sa); 17530
(Sa); 17535
(Sa); 17539
(9); 17544
(13); 17548
(13); 17549
17551
(13); 17552
13426
Murata
et al. 5903
72630
Nash
457
(Sa); 14219
(8); 14242
(9); 14250
(8); 14253
(8); 14254
(12); 14255
(8); 14260
(9); 14262
(4); 14269
(8); 14271
(12); 14274
(Sa); 14277
(8); 14278
(11); 14286
(13);
(8); 14226
Nation
(Sa); 14251
Nease
(9); 14259
(9); 14281
(1c); 14314
Nee
(8).
18069
(9).
Neese
& Goodrich
(12); 14285
Neese
& White
(9); 14298
Neilsen
(10); 14386
Nelson
(4); 14411 (Sb); 14415 (lb); 14431 (9); 14468
(Sb); 14470
(Sb); 14481
(9); 14500
(10); 14511
(Sb); 14512
(Sb); 14527
14614
(8); 14617
14649
(12);
14681
(9); 14682
(12);
14685
(13);
14686
(13);
14688
(8); 14690
(1c);
14691
(8);
14692
(4); 14693
(8); 14694
(8); 14643
(10);
(4);
14695
(1c);
14696
(10);
14706
(9); 14707
(5b);
14709
(Sb); 14711
14712
(2); 14734
(13);
14735
(9); 14741
14697
(10);
14698
(4); 14702
(12); 14751
(13);
14752
15048
15070
(la);
(la);
15243
(Sb); 15248
(la);
15253
15271
(lb);
(Sb); 15284
15291a
(la);
(la);
15050
15238
(10);
15241
15067
(1c);
(10); 15244
(4); 15245
(9); 15250
(5b); 15251
(la);
15254
15272
(5b);
(lb);
15285
(13); 15291b
15307
(la);
(la);
(la);
15273
15308
(lb);
(Sa); 15289
(13);
15295
(la);
15242
15246
(la);
15309
& Macbride
1265
Nelson
& Nelson
6029
(Sb); 1323
(Sb); 1631
(lb);
3 (12); 4 (12); 6 (12); 8 (12); 16 (12); 17
Nichols
(12); 857a
Nichols
Ni6it
(13); 8967
& Lund
346
393
(9).
528
Nixon
186 (Sb).
(13); 640
(9).
823 (1c).
143 (9).
Norby
& Norby
(Sb); 534
Norris
481
(Sa); 496
& Frodin
33099
(8).
74 (3).
185 (13).
Novotny
42 (10).
Nowak-Krawietz
623
(9).
Ogden
4938
Oka 35204
Oldenburg
(9).
(9).
40-44
(Sb).
(lb); 510
(Sb); 555
(Sb); 554
Northington 1085 (lb).
Nuttal
(3).
(9).
Nimke
Nolde
(9).
(Sb); 574
518
Oberwinkler5161 (10).
(la);
(Sa);
(Sa); 9483
(Sb).
15283
15304
8577
Nerlich 63406 (10).
Newsome 319-63 (Sb).
(13);
(12);
(Sb); 2697
(lb);
2150 (lb); 2406 (13).
Northcutt
(2); 15252
15255
(9); 6049
(Sb).
Nobs & Smith
(la);
(lb);
(10); 519 (Sb); 2350
(la); 4090
Nighswonger 1253 (2).
(13);
(3).
(Sa).
Nieuwland
(9);
(Sb).
(3); 3751
(lb);
14755 (13); 14758 (13); 14761 (9); 14763 (1c);
14764 (10); 14765 (10); 14768 (2); 14769 (2);
14770 (5b); 14773 (9); 15009 (3); 15010 (lb);
15013 (Sb); 15019 (lb); 15039 (lb); 15042
(lb);
3591
(2);
14742 (13); 14744 (13); 14746 (13); 14749
(13); 14750
344
(lb).
8161
et al. 3498
Nelson
(Sb);
(lb); 6560
3672
14510
(lb);
14546 (lb); 14548 (5b); 14551 (lb); 14552
(Sb); 14555 (Sb); 14561 (5b); 14570 (Sb);
14571 (5b); 14575 (lb); 14578 (Sb); 14580 (8);
14587 (8); 14603(c); 14609 (4); 14610 (1c);
(8); 14618
30 (10).
458
(9); 14276
(10); 14384
(13);
(13); 17555
(9).
6256
(9); 14296
(12); 17550
(13); 17554
(9).
Neese
(9); 14270
(1c); 14275
14294
14218
(lb);
(9); 14224
(9); 14221
(9); 14239
(9); 14304
(Sa); 14204
(8); 14214
(12); 17542
(9); 17547
(10).
Naito
(12); 14201
14194
(9); 17540
(9); 17546
(Sa).
(3); 14193
(8); 14209
(5a);
(Sa); 17537
(13); 21933 (9).
Murata
Murley 1022 (8).
Mutel 371 (9).
(12);
(9); 17520
(9); 17524
(Sa); 17532
(5a); 17536
(Sa); 17543
(13); 17516
(13); 17519
(9); 17523
(Sa); 17538
14156 (12); 14159 (lb); 14162 (lb); 14167
(12); 14170 (9); 14179 (lb); 14181 (lb); 14187
(11); 14205
(5a); 17515
(13); 17518
17534
(lb); 13997 (lb); 14001 (lb); 14008 (Sa);
(13); 17514
(13); 17517
(Sb).
(lb); 512
Oosting 33419 (9).
Ooststroom 5341 (9); 17429 (10); 18307 (10); 18363
(12); 21395
(10); 19738
670
Osterhout
7168
Otis
223
OENOTHERA
1997
(13); 8019
2164
(9).
(lb); 6317
(lb); 4624
(Sa); 7167
(9);
27430
(9); 28666
(9); 27494
(13); 27420
(9); 31135
(9);
6152
Owens
& Hecht
Pailin
1704
(13); 5415
3167
10746
Pabst & Sick
(10).
Palmer, E. J. 807
(1c).
(8); 4221
(9); 3452
(Sa); 12811
14437
(lb);
(9); 9001
(lb);
17045
18546 (Sa); 19591 (9); 33046 (9).
Pammel
& Blackwood
Pammel
et al. 3867
Parish 4201
3737
2070a
(9).
Porter & Porter
10535
(Sb).
(9);
(9);
& Patton
Powell
Preble & Cary
65-89
Prese
55 (9).
Pretz
11449
Prince
749
(13).
749
(13).
(Sa); 11450
5229
Qian
(9).
704
(Sa).
2326
(13); 2329
Quadgras
Paulin
1704
Radcliffe 17 (9).
(9).
1081
967
Pease
12857
(Sb); 1622
(Sa); 973
(13).
166 (9).
78 (10).
Pearce
(8).
158 (Sb).
Paude
Payson
(Sb).
Pucker 088 (13).
Pullen 65328 (9).
Purer7794 (Sb).
(4).
(10).
Passini & Robert
(Sb); 9675
(Sb); 8381
Porter & Condit
Pryor
(9).
(13).
9977
(5a).
(7).
7 (Sa).
Proace
(lb).
Parks 78 (9); 3113
Parodi
(lb).
(Sa).
Parker & Parker 69220
Parnis 419
(la);
Potter590 (12).
Potzger 2628 (9).
(9).
10722
14078
(la);
(la).
(9); 2689c
1285
Porter 4329
(13).
(Sb).
E. 293 (lb); 2520
Palmer,
14075
(la);
14123
Porsild 7961 (9).
(9); 5018
4380
Ownbey
1650
Pope
(Sb).
(Sa).
Ownbey
(la);
Poltzger 8702 (9).
(5a); 14403
(5a); 13136
695.5
Polgar
Pollard
(9);
34479 (9).
Over
(9).
Pole-Evans 4015 (2).
(9); 26328
(5a); 26217
24567
(2); 14069
14108
(5a).
Oudemans 566 (12).
Ouren
1232
& Konovalova
Pobedimova
Pohl 4659
(13); 4995
(lb).
Piper 236 (8).
Pittillo 650 (8).
Podlech 11572 (10).
(5a).
(9).
Otto 4750-2
et al. 18992
Pinkava
Radford
(Sb).
(2).
14757
(9).
(9); 15386
(9); 14910
16123
(8); 16657
(9); 17801
(9); 16045
(8);
(9); 18315
(9);
27488
(7); 28230
(9); 28356
(9); 28547
(9);
Pease & Bean
23526
(12).
29023
(9); 30075
(9); 30521
(8); 36686
(9);
Pease & Long
21992
(13).
37283
(9); 37775
(9); 37920
(9); 38162
(9);
Peattie
(Sa); 36154
38454
(9); 38751
(13); 39061
36134
Peck 4381
(9); 37353
(8); 28425
(Sb); 4382
8662
(4); 8663
22308
(9).
(Sb); 5767
(4); 9758
(Sb); 23759
Pedersen
411
(9); 9753
Pedersen
et al. 170 (9).
(12).
(lb); 5777
(Sb); 20440
(Sb); 24366
(9).
14902
Perry 4506
4164
(13).
(9 x Sb).
198 (13); 314
Philcox
713
(Sb); 14129
(lb);
Ramaley
& Gambill
16978
Ramaley
& Robbins
3574
& Ramsey
Ramsey
et al. 8873
(9).
Rapp
(9); 2156
(10); 2157
2754
V-452
10472
14411
(Sb); 15218
(13).
(10); 2757
(Sb).
(10).
88 (9); 502
Rastetter
Raup
Raven
(12); 9310
(13); 350
(13); 604
(13).
(12).
4876
(12); 6737
7695
(9).
16316
(9).
(9).
1005 (7).
(Sa); 4372
3886
(9).
(10); 1104 (lb).
(8); 10473
(13); 231
A-36
(lb).
(5b).
2440
838
& Randolph
Randolph
Ranger
Pickard & Coveny
& Hammond
Ramsey
Phillips 249 (9).
Pinel & Wallis
(8).
(10).
1356
Randolph
(9).
2155
(13).
6862
(9).
3887
Ramseur
(9).
194 (9).
Phelps
2581
(Sb); 16614 (lb).
(9); 16291
Perdue & Blum
Petit
4851
123 (9).
Perdue 7057 (5b).
Peters
& Whetsone
Ramaley
Pelgrins 1822 (10).
Peng 8348 (10).
Pennell
& Radford
Rallet
Pedley 1165 (10).
Peeters
Radford
Radford
Rainha
(12).
(9);
39766 (9); 40349 (9).
(4);
(4);
(9); 39416
(12); 4373
(12); 9476
(10); 16493
(13); 4374
(13);
(13).
(10); 18708
(9); 20476
(9);
50
BOTANY
MONOGRAPHSVOLUME
SYSTEMATIC
224
20490
(Sa); 22111
(9); 20535
(8); 26265
(Sa);
26526 (Sa);26545 (5b); 26550 (lb); 26551
(5b); 26559
(2); 26562
(9).
Raven & Engelhorn
(10); 25998
(10); 25982
(10).
26109
(5b).
74 (9 x 12); 285
Redfearn
et al. 1091
Reed & Morton
3995
2047
(2).
2758
(9); 31259
Rousseau
& Reznicek
4780
Rowe
(9).
Rowell
Rice
Roy
(9).
(9 x 10, pro
(9, pro parte); 9820
parte).
Richards
& Massey
& Robertson
Ricksecker
75 (Sa).
3301
(Sa); 1738
1554
Richardson
Ridgway
1011 (Sa); 1471
& Jongepier
& Mullens
Rodman
1913
4032
Rogers
& Rogers
1068
& Sake 62121a
et al. 3818
Romans
2-2
R1555
247
Ropke
Rose
(3).
284
(13); 64
(9).
(9).
(7).
Sakiya
97 (2).
S 1191 (Sb).
(9).
(5b).
Sandberg
542
Sandberg
& Leiberg
Sarvis
(9); 8520
7896
(13); 1741
(13);
(9); 7867
(13); 9630
(9); 6041
(13).
(9).
5804
6883
(13);
(13).
(5b).
3886
(5b).
(11); 10301
(9).
95 (Sa).
172a (9).
Sawada
Saxe
397
(9).
1131 (8).
23 (9).
Scammon
3389
(Sb).
Schaeffer
3258
(9); 3331
(Sa); 59523
(13); 36869
(13); J9613
Schaeftlein 1148-35 (9).
Scheidweiler 1633 (9).
(5b).
Scheppig
(13); 499
(13).
(8).
(5b); 8194
4595
11160
Sanders & West
(13); 49654
Schmalzel
Schmid
(9); 16999
(9).
(5b).
(5b).
(9).
Salzmann
Sargent
(13); 7642
(5a); 4583
(5b); 4588
(10).
Sandercock
(7).
(9); 6041
4584
Sallans
Sandall
(12).
3162
(9).
(Sa).
Saif 9 (10).
(la).
(12); 826 (5a).
(13); 765
(Sa); NR-2297
(5a); 1578
(5a).
Salle 440
(9).
193 (12); 387
Rosback
160 (Sa); 1573
9410
(5b).
& Coutu
(9); NNR815236
1001
(13); 64-2233
(7); 45602
(13); 7869
Rolland-Germain
(9); 3905
(9).
& Bessey
Sausan
(9); 36551
Ronald
775413
& Garrett
1926
(13); 765
125-4-5
(9); 7868
(9); 3187
(9).
& Miller
(9).
(13); 6025
19285
(13).
(13).
(9).
Rydberg
(5b).
Rolland-Germain
2291
1259
2127
Rydberg
19 (5a).
41569
(9).
(10).
(13).
(9); 42495
Rogers
Roland
(13); 50278
(2).
(9); 3006
(9); 2829
(13); 64-3978
67132
Rogge
Rohrbaugh
(12); 31256
(9); 32445
(12); 32064
Rydberg
(9).
Rogers
32063
(10);
(9).
(la);
64-1952
150 (13); 578
Rodgers
(12); 35584
(13);
(9); 7181
93 (8).
Sahira 235
(13); 64-3850
Robinson
(9); 32280
(13); 2528
(9); 2824
2929
& Pugh 65-6366
Roberts
Rood
(9); 1592
1682
& Bateman
2659
(13).
353
Rydberg
(1c).
(9); 1055
Robert & Schmalzel
Roberts
209
Ruth 997
144 (9).
Robert
(9); 1715
(9); 6477
(13); 26562
(2); 10583
Ruffo
Russell
(9 x 10); 2530
2529
Robbrecht
(9); 810 (13); 1107
(13); 1162
(5a); 4841
(9); 1642
Rudd
Russel
(5b); 4030
638
1249
Runyon
(Sa).
(9).
1178
Robbrecht
(13); 434
(9).
4327
Ruggles
(Sb).
Riggins 351 (Sa).
Robbins
(9); 288
Royce 8506 (10).
Richards 3487 (lb).
9820
(9);
(9).
(9); 25725
& Bonin
339
Rhodes 384 (9).
Richards
(9); 286
(9); 13/67
(9 x 12); 70 (5a);
(12); 31269
(13); 35326
(Sa).
Reznicek 951 (13).
1953
(9); 2495
(9).
Rousseau
25572
(2).
(9); 9/79
(13); 26/63
(13); 1151
(13); 1150
Reeves 4524 (lb).
Rensburg 407 (13).
Reznicek
(9); 20/63
80 (9); 166 (9); 507
Rouleau
(9).
Redfield 118 (lb).
Reverchon
(9 x 5a); 8/74
3/65
14/65
(Sa); 911 (9); 912
11 (9).
1373
(13).
(2).
F9 (5b); F87
Reardon
Repton
(13); 8104
Rosser & Bewick 82/52710 (10).
Rossmann 91/66 (10); 286/66 (13).
Rostaniski
Raven & Raven
Raymond
7628
(9); 27899
(5a); 27898
25804
Roske 396 (9).
Rossbach 2879 (13); 4284 (12); 6081 (12); 6503 (9);
5134
(9).
& Todzia
80-72
2037
(10).
Schneider8074 (Sb).
(la).
(12).
(9); 49885
Skorepa6654 (9).
Skvortsov 10186 (9); 10191 (5a); 10232 (9).
SkrochB169 (12).
Schofield 3702 (13);4096 (12).
1412
& Scoggan
Schofield
(5b).
Schreter2195 (5a).
Schultz 1803 (5b).
1272
Schur
37729
Scoggan
3229
(Sb); 3409
(Sb); 3644
(Sb); 10111
(5b); 9460
(9); 12282
(12); 13355
(12); 13803
(Sb);
(Sa);
(13); 12365
7906
(Sb); 8015
(12);
(9).
Smith & Jennison
57 (5b).
4002
2036
4859
1274
(9); 1275
2097
(9).
Senn
Senn & Zinck
Senn
4403
Seymour
(13); 1276
(13); 2006
(13);
Soldano
(13).
(9); 10815
(13);
(Sb).
15501
Seymour
& Countryman
Seymour
& Seymour
21864
(13).
Seymour
& Svenson
25971
(9).
431
1898
Sharp
Shaw
2557
(5b).
Shchepanek
2051
(9); 17263
(10).
Somes G3871
3339
(7); 5511
(9); 1418
(13).
(Sa); 8685
(lb);
(9).
Spragg
(9); 3308
(13); 3345
(12); 11686
(12); 21231
(9); 22008
(9); 24642
(9); 27668
(9); 27677
(8).
(9); 27802
79 (Sa).
Silander
153 (9).
Silberhorn1785 (Sa).
Sintenis 953 (9).
Skelton
& Skelton
605
(9); 693
(lb).
186 (Sa).
(9);
(9);
(Sa);
354
(13).
(Sb).
(13); 1284
St. John 1283
St. John et al. 9237
Stackler
Stahmaun
Shultz 1803 (Sb).
Siemers
et al. 14 (13).
6853
(9).
Spradboron62471 (9).
(Sa).
(9); 33399
(5a); 4205
(Sa).
388
Spillman
24888
(Sa).
(Sa); 2822
(Sa).
& Soreng
Spicer
28782
(9).
(12); 3383
Spellenberg
(10).
13585
(9).
(13).
(12); 2538
11274
212
Shields 628 (Sa).
1471
(9).
(12).
2879
Spegazzini
Shepard
Shinners
(9); 19417
(13).
288
Sparling
(9); 92 (9).
(10); 12020
(9);
Spalding 2055 (8).
(9).
605
12019
(9); 3939
(13); 5259
5008
2430
4950
Shelton
222
(8); 3932
(9).
7389
et al. 2240
Sorensen
Sperry
Shimada
4223
Soper & Shields
(8).
(9); 5980
(9);
(9).
Soper & Minshall
1009
(9); 3624
(9); 4941
19387
(Sa); 945
Soper & Dale
8697 (lb); S.10103 (9); S.11009 (9); S3470
(Sa).
Shimek
337
Soper
Soper
(13); 11874
(9); 3551
(6); 4856
Soltis 380 (9).
(9).
& Dougal
10566
(8); 3928
Soper & Grevatt
(Sa); 16996
Shea
Sheldon
22587
(9); 3259
(13).
& Barkalov
Solomon
(9).
Shchepanek
(9).
(9).
(9); 4178
3926
3979
(8).
Sharp et al. 9593
614
(9); 1617
1137
Solomon
(9).
et al. 13325
Shanks
(12); 9988
(9).
12677 (9).
(13); 18335 (9).
Shabani
(Sa); 8183
Sohmer 4749 (13).
(9).
4671
(7).
Sohma & Takahashi
(13); 818 (13); 2801
et al. 575
Sennen
(9).
(Sa).
(6); 3654
927
2688
(12); 7472
(9); 5226
(Sb);
(9); 6505
Sodiro 333 (10).
Soest 23993 (9).
(9).
(Sa); 2479
(9).
(8); 4792
55 (9).
Snyder
(Sa); 7866
2347
Seiler
4113
1014
Smyth 3903 (13).
(Sa).
Sears & Ahles
et al. 3399
Smith
Scrymgeour 191 (10).
Sdorovska 17621 (9).
Seigler
(Sa); 7151
Smith & Hodgdon
(9); 281
(lb);
(9); 1513
(Sb); 1439
(10); 2411
(13); 2320
1571
(Sa).
(13); 276
(5a); 905
(9); 878
(1c); 1233
(5a); 1063
(1c); 6875
& Baldwin
Seaman
(12); 877
(9); 870
15274 (10).
Scott 588
(12).
3 (Sb); 42 (9); 117 (2); 266a
Smith
(13); 13260
(13); 14637
(9, pro parte).
(9).
(Sa).
Smit et al. 5226
(5b); 12174
(5b); 11237
(12); 12208
Scribner
1452
Smejkal
(Sb); 11154
10521
et al. 11093
Small
(Sa).
(Sb); 3297
(Sb); 4114
3892
(9).
(13, pro parte); 2581
2581
Sloover
(9); 11112
(13); 6271
Schwab
Scoggan
225
OENOTHERA
1997
1758
(13); 2830
(Sa).
(9 x Sb).
(Sa).
341a
(13).
(lb); 4212
Standley
4010
Standley
& Bollman
Stanislas
574
Stardom
3733
Staunton
et al. 8321
Stebbins
237
11145
(Sb); 4246
(Sb).
(13).
(12).
(10).
(9 x 10).
(Sb).
(12);
226
SYSTEMATICBOTANYMONOGRAPHS
4 (8); 8 (12); 10 (8); 20 (8); 44 (8); 54 (11);
Steele
71 (9); 97 (13); 104 (9); 120 (1c); 290
17 (13); 222
Steele & Steele
Stephens
(Sa); 7827
6339
8705
(8); 328
(13).
(Sa);
(Sa); 15808
(Sa); 16403
(9); 19259
(lb);
(2); 28026
(Sa); 28231
(9); 28996
(9);
36556
(Sb); 49661
(lb); 50556a
(Sa); 50438
(Sa); 50635
(9); 58066
58516
(5a); 58847
(Sa); 58965
(Sa); 59077
59396
(Sa); 59130
(Sa); 59898
(9); 61497
(5a); 68345
(5a); 68485
(5a); 69169
(Sa); 69641
(2); 82827
83175
(Sa); 83288
(Sa); 87523
(Sa); 88399
& Brooks
(Sa); 17016
13732
(Sb); 26501
26487
35174
(Sb); 41580
(Sa); 43014
Stevens
1087
(9); 1907
Steward
2499
(10); 9676
Stewart
2618
(13); 2204
(Sa); 978
(13); 3544
(9); 832
(13); 1233
(12); 834
(5b);
(lb); 42852
Terrill 4629
(13); 6895
(13); 4630
(5a); 2899
(2).
Tharp 224
(2); 2437
(Sa).
Theis 10 (9).
(10).
Theron
(10); 1703
598
Theroux
(2).
& McDougall
735
(3).
(Sa); 64858
(9); 64858a
(Sa);
Thieret 24391 (9); 24655 (9).
(9); 68865
(Sa); 70214
1143 (lb);
1144 (lb);
(Sa).
1146
1063
(Sb).
Stone
14799
(9).
28480
(Sb); 1502
(9); 1856
297
(10); 1519
(10); 1658
(Sb);
(9).
(2); 404
parte); 7285
(2).
Thone
103 (9).
Thorne
6117
345
(8); 14653
(lb);
(10); 7285
(9 x 10, pro parte); 9528
(10, pro
(10).
(Sa).
Tolstead
411270
(Sa); 411271
(13); 28a (8); 29 (9); 30 (13); 32 (8); 33 (8); 34
Townsend
(8); 36 (9); 37 (8); 38 (8); 39 (8); 44 (8).
Tozer
(4); 14 (4); 14a (4); 15 (9); 17 (13); 18 (13); 19
745
(5a).
414
Train 2438
(9); 2066
(9 x Sb); 4765
5859
(9 x 5b); 5860
(5b); 5868
7576
(9 x 5b); 7577
(9 x 5b); 7615
(10); 6269
(4).
(lb).
Trainer63-19 (Sb).
(9 x 5b);
(9);
(9 x Sb);
Troll 6050
(9).
TroyKPP158 (13).
(lb).
(9).
(4); 8001
(4); 18401
(Sa).
107 (lb); 445
& Barber
141 (9); 532
Tracy 4968
(9).
113 (5b).
1691
(4); 835
Tidestrom7432 (12); 10693 (Sb); 13416 (9).
Tidestrom& Bartlett 5222 (13).
Tiehn 3610 (Sb).
(13); 20 (11 x 13); 21 (13); 23 (9); 27 (8); 28
Suchoboskij
(13); 488
(Sa).
Tonghua expedition 373 (Sa).
Toumey 140c (lb).
Tower 8889 (10).
Suksdorf
(Sb);
(9); 18387
63 (2).
Thurber
(2); 3870
1 (9); 5 (10); 6 (10); 7 (10); 9 (4); 10 (4); 11
Stubbe
(1c); 11357
(Sb); 14779E
Thorp 2798 (9).
(13).
(10); 3171
8715
(10); 4301
(9 x 13, pro parte).
(9); 1983 (10).
Thomson 77 (9).
Strohmeyer249 (9).
Strong
(9); 2654
(9); 14726
20 (13); 163 (lb);
Thompson
(13).
1174
326
219
11531
(13, pro parte); 231
18389 (lb); 18390 (1c).
Stolze
Strey 2528
231
Thomas
1145 (lb);
(lb).
Stratton
(13).
Tessene 228 (9).
Tessier 601 (9).
(Sa); 24661
Strandberg
(9);
(13);
2529 (12).
Temp 61007 (9).
(Sb); 34102
Thielens
1698
(12).
(lb).
(13); 1230
(13); 2527
(9); 2526
(9); 24170
Straley
8366
(13); 16863
et al. 86 (9); 87 (9); 132 (9); 142 (9); 143 (9);
(Sa); 20345
Strahler
(12);
(9); 1451
(lb);
(9); 12149
12004
(Sa); 15071
67022
(9); 2896
(9); 1046
(9); 5205
(Sa); 14984
Stockhouse
Sudol
(Sa); 864
1231
Steyermark8118 (9); 14266 (9); 14831 (9); 14955
Sublis
(9).
726
146 (9); 830
(lb);
(10).
(13); 3331
(9).
(9); 2528
(Sa); 16342
(Sa).
(Sa); 2253
247
13309
Taylor & Taylor
115 (5b).
Stemnberg
(9).
83 (8).
Taylor
(Sa); 42496
(Sa); 43669
1950 (Sa); 2182
Taggart
(Sa);
(Sa).
(lb); 35052
(9 x 5b); 11516
(Sb); 10603
118 (9); 164 (9).
Taylor
(lb); 87750
(lb); 26768
(5a); 34822
(Sa); 34568
Sutter
Tanaka
(Sa);
(Sa); 26150
(9 x Sb);
(9 x 5b); 7915
Summers 308 (1c).
Sumstine 779 (9).
Sundquist509 (9).
Survey 283 (5b).
Sutherland290 (9).
Takenaka
(5a); 69507
(Sb); 14486
(Sb); 24962
(Sa);
(Sa); 83121
(lb); 82558
(Sa); 88108
(5a); 67944
(Sa); 70941
(9 x Sb); 7654
(9 x Sb); 7891
3165 (9).
Sykes 50/85 (13); 122/89 (10); 545/81 (10).
(Sa);
(Sa); 68596
(5a); 69343
(Sb); 7807
Swanson
(Sa); 60712
(5a); 63669
(Sa); 70187
(9);
(Sa); 59048
(Sa); 59232
73895
Stephens
(Sa);
(Sa); 58149
(Sa); 60030
(5a); 61318
68967
(Sa); 50556
(Sa); 50887
(5b); 51758
(9 x Sb); 7653
7706
(9 x Sb); 8910
(Sa); 8352
27217
51675
7652
(11).
(5a); 8082
VOLUME 50
(7); 8302
(4); 13598
(4); 15602
OENOTHERA
1997
True 5 (13); 52 (8); 145b
(9); 185 (13); 752
(13);
6942 (13).
Truscott IA (10).
Tryon
2031
Tsui 437
Wahl
(9); 2652
Turner 348
14422
(9).
(Sb); 1979
(Sb); 2558
(Sb); 2579
Walker
208
& Jennison
(9); 424
Waller
1170
Wallis
825
(Sb); 2631
(Sb); 2645
(Sb); 2650
(Sb); 2693
(Sb); 2703
(Sb); 2710
(Sb); 2763
(Sb); 2813
Walther
(Sb); 4175
(Sb); 7255
(Sb); 7853
(Sb); 7859
Wan & Chow
(Sb); 5096
(Sa).
Twisselmann 3814 (lb).
& Ugent
Umbach
1293
4283
(la).
(9); 4500
Unar & Unarova
(13); 8400
1542
(9).
(10).
609
Ungar
(9); 6588
Van Horn
(9); 7338
(12); 7530
(13).
Wang
Zhong-tao
et al. 870007
Ward
7 (2); 585
(13); 586
(9); 1704
266
T142
7943
(lb);
2978
1061
(9); 1706
(9); 3173
(9); 1729
(9);
Weber
Weber
(lb); 6869
(9).
(Sb); 626
(Sa); 2198
(Sa); 4091
& Steward
15240
Wehmeyer
14 (9).
Wehmeyer
& Waters
Wells
1104
16348
(10).
(9); 4797
(13); 5154
(12); 5316
(12); 7738 (12).
6017
1469
(10); 1568
(12); 3232
Wagenknecht
Wagner,
1622
(lb); 4021
(12); 2005
(9); 3428
(lb);
(lb);
(10); 4470
(lb); 4476
(lb); 4512
(Sb); 4532
(lb); 4540
(9); 4542
(10); 4545
pro parte); 4547
(Sa); 2860
(9); 3108
3790
(Sa); 4043
(9); 4546
3809
Welsh
& Welsh
12548
315
West
(9).
(3); 23654
(3).
(Sb); 19114
(Sb).
3852
2578
3964
(lb).
Wagner,
W. L., & Halley
4535
(9); 4537
(9).
(9).
(9); 6172
(13); 6632
Westerfield
Weston
(lb); 4529
(9); 4547
(12).
(Sa).
(9); 11755
(8); 9721
(9); 5566
(13); 12291
(8);
(8);
12452 (13); 12646 (8).
(lb);
Wetter
(9 x Sb,
et al. 5838
et al. 3004
039
(8).
(la).
(Sa); 041
(Sa); 043
(Sa); 046
(Sa); 049
(Sa).
(9);
(Sb, pro parte).
4521
729
(lb).
(9); 4543
W. L., & Brown
W. L., & Mill
(3); 20844
(3).
89 (Sa); 6073
Wertman
Westerfeld
(Sb); 4465
Wagner,
Wagner,
2458
6171
(Sa); 4035
4544
(3).
(9).
(3); 19283
& Moore
(12).
(Sa); 1989a
W. L. 3728
(10); 1929
Welsh
Werner
(9).
Wadmond 3276 (9).
Wagenitz
(Sa);
Werger 1330 (10).
22 (13).
Vowles
Waalkes
(Sb); 7199
143 (Sa).
Welsh
(Sa).
(12); 4607
(9);
Webster 8323 (9).
Wechuysen 1118 (13).
Wedberg 1505 (lb).
Volle
2476
(13); 6910
(3).
57 (13); 95 (Sa).
Voss
(Sa); 10405
31 (Sa); 32 (Sa); 59 (Sa); 60 (Sa); 188 (Sa);
7940
(9); 40 (9).
6334
(13).
(9).
Welch
von Ruynen
(9).
(9).
(Sa).
Viviers 196 (13).
652
(Sb); 8478
(9); 4036
1126 (13); 5123
(Sa); 267
(Sa); 6173
(lb); 21391
(lb); 3461
220
244
Webb
Vick 56 (13).
Vickers 3 (9).
Victorin-Lavoie 245 (13).
Visher
(9).
(9); 1040
198 (Sa).
Watson
(13).
1-223
(9).
6911 (9).
(9).
& Erbe 2730
(9).
(Sa).
1032
1794 (9).
Vincent
81070
et al. 1413
Weatherby
260
Vincent
(Sa); 5980
Warren 695.5 (Sb).
(13).
4260
(9); 2787
(Sa).
Van den Houten
Vanhecke
(9).
(5a); 2738
Wang
Waterfall
(Sb); 6577
1656
(12).
(8).
1 (13); 125 (10); 6702
Warnock
(Sa).
1451-2
(9); 18024
(8).
Wang
(13); 9930 (Sb); 10580 (10).
Valbusa
(9);
(8);
(9); 7925 (2).
9 (9); 66 (9).
5173
(9); 3651
(13); 13635
(ic); 2647
1695
(Sa); 2614
Warnecke
Underwood 2512 (13).
Urumoff
(lb); 6501
(2).
(Sb); 2629
(Sb);8400 (Sb).
(8); 3445
(13); 14848
(9); 80030
Wallace
(Sb); 2622
Ugent
(lb); 6500
(9); 10294
(9); 14423
et al. 7875
(Sb); 2606
236
6499
(8); 3347
(8); 5477
(5b); 2605
Tweedy
Uttal
(9); 208
3973
Wahl
(la); 6115
(9); 736
206
(9).
(5a).
1268
W. L., & Sisson
Wagner,
(lb).
Tscherning5056 (12).
Tsiang 10717 (10).
Tucker
227
Wetzel
475
Wheeler
(lb).
53 (1c); 627403
(10).
Wherry 1933A (11).
(9); 4538
Wherry
et al. 7084
Wherry
& Pennell
Whetstone
(9).
13377
& Wagner,
(11).
W. H. 5238
(8); 5950
(9).
SYSTEMATICBOTANYMONOGRAPHS
228
White
63 (9); 4187
Wolfe 131-141 (lb).
Wollangk 3 (13).
(lb).
Whitehead 46 (8).
Whiting
& Sanders
5100
Wood
(Sb).
Whittacker 260 (9).
Wickham
Wooton
72 (Sa).
(9); 1717
(Sa); 10503
286
Wiegand
& Manning
2209
Wiegand
& Wiegand
236
Wiggins
& Gillespie
4111
Wiggins
& Wiggins
20696
(10); 2155
1834
(Sa); 10504
(9); 2210
(12); 237
(Sa).
(9).
(13); 238
(13).
(10); 653
(9); 792
(9); 965
& Ahles
(8).
& Mueller
Wolden
1307
(13); 2736
(7); 3043
1435 (lb);
3185
(13).
(Sa); 1729
2254
(3); 6223
Yates
3902
224
2834
Zhang
1899
mellifera
(9); 2634
(Sb); 2794
(lb);
are in roman type; the main
(lb).
(Sa).
(Sa).
(9).
2179
(12); 2932
(12); Z21
Zimmerman
792
Zinck
(9); 40 (12); 410
1043
(13).
(13); 954
entry for each
is in boldface.
are in italics.
Synonyms
subg. Onagra
(Adans.)
sect. Contortae
W. L. Wagner
Jeps. 37
21
(Torr. & Frem.) W. L. Wagner
sect. Kneiffia
93
(Spach) Endl.
sect. Lavauxia
(Spach) Endl.
sect. Oenothera
22
sect. Onagra
3
sect.ParvifloraeRostafiski 39
3
3
subsect. Raimannia
3
W. Dietr.
23, 56, 63
ser. Allochroa
21
L. 37
ser. Devriesia
Torr. & A. Gray
38-39
(Rose ex Britton & A. Brown)
2, 22, 38, 70
W. Dietr.
38
ser. Candela W. Dietr. & W. L. Wagner
ser. Clelandia
subg. Euoenothera
2, 21, 22, 37, 38
W. Dietr. & W. L. Wagner
2, 21, 22, 38
subsect. Oenothera
sexta 21
Oenothera
W. Dietr.
subsect. Nutantigemma
lineata 21
quinquemaculata
P. H. Raven, W. Dietr. &
4, 142
subsect. Munzia
Gayophytum 3
Hieracium 7
Hyles
P. H. Raven
2, 22, 23, 38
subsect. Euoenothera
Stubbe
22
(Munz) W. Dietr.,
& W. L. Wagner
37
38
(Spach) Endl.
subsect. Emersonia
21
Fischer & Meyer
Rostafiski
sect. Xylopleurum
Deilephila
elphenor lewisii 21
Eumorpha
Manduca
(Adanson)
sect. Strigosae
7
21, 22
38
3
Iridaceae
21,
22, 142
(Spach) Endl.
Commelinaceae
Gaura
(Sa).
NAMES
sect. Hartmannia
37
(L.) Bubani
achemon
(12);
(13).
Campanulaceae
Crepis
(9); 2819
(Sa); 2661
(13); 3043
22
Calylophus 3
Clusiaceae
(9).
(2).
sect. Gauropsis
1
biennis
(9); 43885
(4).
names
Bubani
(2).
(10); 27886
et al. 479
2890
(9).
Agrius 22
convolvulii 21
Alchemilla 7
Apis
Brunyera
(lb).
(9).
& Leykom
INDEX TO SCIENTIFIC
Accepted
3713
(5b).
11 (13).
Zeyher
Ziegler
(1c).
6172
& Johnson
584
16027
419
Zhongde
et al. 3707
(lb);
Zeitler 233 (12).
(9).
(9).
Wolf
(12).
et al. 12610
Zarfoss
2191
591 (Sb); 61 (9); 223
Windler
Wolf
60 (Sb); 298
Wright
Young
(Sa).
Wilmott 430819B (10).
Wilson
Wright
Yamell
(Sa); 1246
3332
(Sb); 7623
7537
Yamamoto
Williamson 1283 (13).
Williamson
(11).
Wynhoff W-49 (Sb).
(lb).
(10).
23 (5a); 404
(Sb); 1189
(8); 6802
& Standley
Worthington
Wynd
(lb).
Wilczek 1532 (13).
Williams
5655
Woodland 1300 (10).
Wiegand
Wight
VOLUME 50
142
ser. Linderia
W. Dietr.
Rostafiski
Rostafiski
38
35, 39
39
38
OENOTHERA
1997
ser.Oenothera 35
ser. Raimannia
ser. Renneria
W. Dietr.
Lutz
38
subsp. caeciarum
38
subsp. grandifiora
subsp. xheiniana
19, 134
albicurvaRenner 149
subsp.
albiflexaRenner 149
albifranciscanaRenner 149
albihookeriRenner 149
albilaeta Renner 149
albinervisGates 73
xalbipercurvaRenner ex Hudziok 17, 19, 135,
136
var. impunctata Renner
ex Hudziok
19, 136
93
Stomps
(Teyber) Love & Love
rubricaulis
(Kleb.) Stomps
var. angustifolia
Renner
var. austromontana
var. cantabrigiana
var. cruciata
ex G. Don)
(Nuttall
var. cruciata
143
Kleb.
var. hirsutissima
A. Gray
albivelutinaRenner 150
var. hookeri
ammophila
var. leptomeres
var. germanica
(Boedijn)
Renner
var. angustifolia
Gates
var. flecticaulis
(Gates) Gates
var. laurensis Gates
var. parva
angustissima
124
var. quebecensis
Gates
Mack.
argillicola
var. pubescens
9, 11, 13, 15, 24, 25, 26 ,28, 29,
131, 132, 200, 210
33
Core & H. A. Davis
33, 114,
116
xatra
atrovirens
Shull & Bartlett
125, 130, 149, 150,
151, 152
var. ostreae
auctirubata
austromontana
119
(Munz)
P. H. Raven, W. Dietr. &
beckeri Renner
19, 70, 73, 139, 149, 150, 151
(L.) J. Boivin
21, 23, 25, 26, 27, 28, 29, 30, 31, 32, 33, 34,
mut.
125
Gates
de Vries
109
de Vries
spiralis
brevicapsula
Bartlett
brevispicata
Hudziok
cambrica
146
19, 135
Doll
brevistylis
125
18, 109
de Vries
95
18, 97
107
19, 113, 134
Rostafiski
Rostafiski
18, 98
var. impunctata Rostanski
cana de Vries
82, 83, 89, 90, 91, 93-107,
candicans
de Vries
canovertex
Hudziok
canovirens
Steele
111, 113, 117,
J. Boivin
(Bicknell)
37,39,40,59,68,70,71,75,76,77,78,81,
120, 121, 123, 130, 131, 132, 133, 134, 135,
136, 137, 138, 139, 141, 142, 143, 144, 145,
148, 149, 150, 151, 152, 153, 179, 210
J. Boivin
93
143
Gayer
Gates
Lutz
bipartita
xbritannica
143
L. 4, 8, 9, 11, 13, 15, 16, 17, 18, 19, 20,
113
83
xbiennivelutinaLotsy 152
xbiennoidesLotsy 152
xbraunii
150
25, 90, 91
bauri Boedijn
biennis
(Torr. & A. Gray)
f. ochroleuca
93
J. Boivin,
(L'Her.) Carp.
f. stenopetala
blandina
(Sturtev.) Gates
Renner
Stubbe
(Mack.)
var. cruciata
133
143
f. argillicola
biformifora
de Vries
Kleb.
Torr. & A. Gray
f.muricata
31, 33, 39, 40, 113-117,
var. argillicola
(Rydb.) Piper 78
var. vulgaris
f. hookeri
125
124
(Farw.) Farw. 95
f. grandiflora
127
Gates
apicaborta
90
117
(Atk. & Bartlett) Wiegand
var. sulphurea
126
93
(L.) Torr. & A. Gray
var. strigosa
126
31, 124, 141
Gates
A. Gray
var. rubricaulis
119
52
18, 95
(L.) Torr. & A. Gray
var. oakesiana
45
J. Boivin
(Atk. & Bartlett) Wiegand
var. pycnocarpa
119
120
(Gates) Gates
Miller
angustifolia
var. nutans
Bartlett
var. parviflora
143
Gates & Catches.
ammophiloides
118
118
Renner
(Boedijn)
var. rhodoneura
Renner
83
ex S. Watson
(Torr. & A. Gray)
var. muricata
145, 151
subsp. germanica
Torr. & A.
124, 143
Gray
albiundata
18, 117, 121, 123, 136, 137,
B. M. Davis
(B. M. Davis)
(L'Her.) Lindl.
Focke
70, 72, 153
143
var. grandiflora
149
93
90
(Munz) Cronquist
xalbisubcurvaRenner 131, 149
Renner
95
143
Torr. & A. Gray
var. canescens
137
143
(Pers.) Rouy & Camus
subsp. suaveolens
107
de Vries
96
(Renner) Love & Love
subsp. nuda
149
ex Renner &
(de Vries
Love & Love
acutifoliaRostanski 142
albida
33, 97, 102, 103, 107
subsp. chicaginensis
xadriatica
& Renner
33, 97, 102, 103, 107
Munz
subsp. centralis Munz
Cleland)
Soldano
25, 33, 90, 91, 102
subsp.biennis 33
109
albata Hoeppner
Munz
subsp. austromontana
W. Dietr. & W. L. Wagner
ser.Rugglesia Rostafnski35, 39
ser.StubbiaRostafnski39
aberrans
229
var. cymatilis
18, 98
109
110
19, 73
31, 70, 72
(Bartlett) Gates
118
118
52
94
VOLUME 50
SYSTEMATICBOTANYMONOGRAPHS
230
B. M. Davis
cantabrigiana
45, 47-52, 53, 54, 58, 63, 70, 79, 82, 123,
160, 210
143
carinthiacaRostafiski 18, 98
cheradophilaBartlett 79
W. L. Wagner
96,
Soldano
var. minutiflora
var. parviflora
Renner
Renner
chicagoensis
18, 99
ex Cleland
Shull & Bartlett
cleistantha
cockerellii
& Blakeslee
143
var. hirsutissima
eriensis
72
erosa
(Torr. & A. Gray) H. Lv.
f. parviflora
(L.) H. Lv.
(Torr. & A. Gray) H. Lv.
(Torr. & A. Gray) H. Lv.
H. Lv.
(H. Lehm.)
race japonica
Guffroy
race vrieseana
comosa Gates
124
53
71
Hudziok
Renner & Hirmer
Renner
corymbosa
Lam.
45, 141
corymbosa
Sims
45, 141
var. sabulonensis
19, 113, 132
cymatilis
Bartlett
deflexa Gates
124, 130, 140, 153
127
depressa
94, 124
Gates
119
150
150
fiecticaulis
Gates
Renner
126
150
Bartlett
52, 149
95
fusiformis
Munz
gauroides
Homem.
gigas
140
& I.M.
(Bartlett) Gates
glazioviana
95
108
31, 142
glabra Miller
31, 38, 70, 72, 75, 138, 139,
110
18, 118
Boedijn
de Vries
Johnst.
142
Micheli
5, 11, 13, 15, 16, 17, 18, 19,
20,21,23,27,29,30,31,32,33,34,36,39,
f. angustifolia
Rostafiski
f. latibracteata
Boedijn
Rostanski
19, 73
40, 43, 63, 66, 78, 90, 106, 107-113,
19, 73
118
xgracilis
dubia E. H. Krause
grandiflora
141
f. latifolia E. H. Krause
141
L'H6r. 9, 11, 13, 15, 16, 21, 24, 25, 26,
9, 13, 15, 21, 23, 30, 31, 33, 39, 40,
159, 176, 210
subsp. coronifera
subsp. elata 3, 10, 24, 33, 44, 45, 46, 47, 49,
159, 210
(Renner) Weihe
subsp. erythrosepala
108
(A. Gray
92,
102, 105, 106, 111, 116, 133, 141, 143, 146,
41-53, 56, 63, 69, 70, 82, 89, 105, 111
Dietr.
140
de Vries
29, 30, 31, 33, 39, 40, 51, 52, 83, 85-90,
editicaulisHudziok 18, 97, 137
subsp. hirsutissima
132,
133, 134, 135, 139, 140, 143, 146, 147, 148,
149, 150, 151, 152, 153, 197, 210
xdrawertiiRenner ex Rostafiski 19, 139
elata Kunth
18, 97
var. brevicapsula
153
disjuncta
Renner
flexirubata
germanica
E. Greene
108
Renner
furca Boedijn
118
de Vries
19, 134
19, 134
Rostafiski
Hudziok
franciscana
127
var. bracteata
xdensa
f. rubrinervis
flavirubata
150
Renner
& Rostafiski
17, 19, 113, 133, 134, 152, 153
fiavivelutinaRenner 150
Fern.
var. varia de Vries
Soldano
Renner
flavitincta
var. stenopetala (Bicknell)Fern. 118
curvilaeta
xfallax
flavicurvaRenner 150
flavihookeriRenner 150
18, 110, 151
ex G. Don
Nutt.
xfallacoides
flaemingina
18, 97
coronifera
excelsihookeriRenner 150
fatua de Vries
143
125
compacta
18, 110
Rostafiski
xfalloides Lotsy 152
ex H. LMv. 141
H. Lv.
xconferta
cruciata
52
18, 29, 33, 43, 108, 111
Borbas
var. azorica
(Nutt. ex G. Don) H. Lv.
119
Linder & Jean 18, 35, 81, 98, 105,
erythrosepala
93
119
(Gates) Gates
106
72
124
(Pers.) H. Lv.
var. jamesii
race erosa
(Gates) Gates
70, 71
J. Lehm.
ersteinensis
f. canescens
f. suaveolens
118
Gates
var. repandodentata
93
var. hookeri
Cron
107
var. niagarensis
(L.) H. Lv.
var. cruciata
ex S. Watson)
xepilobioides Lotsy 152
ex de Vries
H. Lv.
race biennis
de Vries
elliptica
125, 130
xcoerulea Lotsy 152
xcoloratissimaHudziok 19, 132
communis
(A. Gray
quist 45
61
Bartlett
44,
48, 51, 52
143
xclaviferaHudziok 19, 137
clutei A. Nelson
texensis W. Dietr. & W. L. Wagner
subsp.
& Jehluk 18, 98
Rostafiski
10, 17, 23, 24, 30, 33, 39,
43, 44, 45, 49, 52-53, 54, 55, 67, 68, 71,
105, 113, 123, 149, 150, 164, 210
98, 107, 137, 150
var. bartlettii
(Torr. & A. Gray) W. Dietr. &
subsp. hookeri
ex Renner & Cleland
de Vries
chicaginensis
ex S. Watson)
W.
10, 16, 21, 23, 24, 25, 33, 43, 44,
var. glabra
Ser. 83
var. pubescens
Ser. 85
110
(Borbas) Love & Love
OENOTHERA
1997
var. tracyi (Bartlett) Gates
mut. gigas
de Vries
146
mut.
lorea de Vries
146
mut.
ochracea
mut.
semigigas
grandiflora
grandifolia
de Vries
146
31, 83, 85
Gates
96
var. macrosperma
graveolensGilib. 93
grisea (Bartlett)Rostafiski 48
hamata
de Vries
Gates
xhero
laevifolia
Gates
126
ex S. Watson)
de Vries
45
mut.
auricula
Torr. & A. Gray
mut.
aurita
mut.
cinerea
mut.
compacta
33, 48, 49
mut.
crinita
33, 43, 47, 49
mut.
cucumis
mut.
decipiens
mut.
delata
21, 33, 39, 43, 44, 49, 52,
subsp. angustifolia
(Gates) Munz
(Bartlett) Munz
Cockerell
subsp. hirsutissima
ex S. Watson)
(A. Gray
33, 45, 49
subsp. ornata
Munz
Munz
(A. Nelson)
subsp. venusta
33, 44, 52, 53
(Bartlett) Munz
subsp. wolfii Munz
var. angustifolia
33, 48, 49
var. ornata
(A. Gray
Munz
147
ex S. Watson)
Munz
48
52
47
Gates
48
var. venusta
(Bartlett) Munz
47
48
19, 70, 72, 140, 151
147
mut.
ingeminans
mut.
lactuca de Vries
147
mut.
linearis
de Vries
147
de Vries
de Vries
nitens
de Vries
147
mut.
opaca
de Vries
148
de Vries
147
147
mut.
mut. perennis
var.wolfiiMunz 63
(Borbas) Borbas
de Vries
mut. pallida
(Ser.) Gates
147
147
de Vries
mut. metallica
Munz
147
147
de Vries
mut. fiava
147
147
de Vries
mut. distans
mut. fragilis
48
73
Gates
Hudziok
de Vries
mut. favilla
48
Gates
var. simsiana
hungarica
147
de Vries
de Vries
(A. Nelson)
var. parvifiora
de Vries
erythrina
52
147
147
mut.
(Wooton & Standl.) Gates
var. semiglabra
de Vries
48, 144
45
var. irrigua
147
de Vries
de Vries
diluta
146
147
de Vries
(Cockerell)
var. montereyensis
de Vries
elongata
(Bartlett) Gates
var. hirsutissima
146
de Vries
mut. detruncata
mut.
146
de Vries
mut.
(Bartlett) Munz
var. hewettii
de Vries
146
146
23, 33, 43, 63
Gates
var. franciscana
25, 47, 49
146
de Vries
de Vries
mut. deserens
33, 44, 53
subsp. montereyensis
var. grisea
33, 48, 49
144
ex Pohl
de Vries
ablata
angustifolia
144
ex Pohl
de Vries
mut.
Rostafiski
subsp. hookeri
var. brevistylis
mut.
67, 149, 150
Munz
Ser. 4, 29, 31, 33, 34, 85, 111, 133,
var. lata de Vries
94
subsp. hewettii
125
126
135, 149, 150, 151, 152, 153
(A. Gray
subsp. grisea
119
Gates
(Bartlett) Gates
lamarckiana
48
xhoelscheri Renner ex Rostafiski 17, 139
var.albinervisRostafiski 144
hookeri
108
var. similis Gates
140
var. rubricalyx
de Vries
var. scitula
126
19, 137
de Vries
150
var. rubripunctata
(Gates) Gates
Cockerell
hirtella
97
l[a]evigataBartlett 125
Teyber
hirsutissima
Hudziok
133
laetihookeriRenner 150
110
de Vries
hewettii
de Vries
laetiflava Renner
126
var. subtenninalis
xheiniana
xlaeta
143, 144
var. parviflora
144
9, 10, 13, 16, 17, 20, 23,
24, 31, 33, 39, 40, 43, 44, 53, 56-61, 63, 76,
78, 83, 141, 145, 165, 210
jepsonii E. Greene 47
jueterbogensisHudziok 18, 97, 98, 136
146
Lam.
hazelae
& Soldano
italica Rostafiski
94
jamesii Torr. & A. Gray
de Vries
guttata Cockerell
231
148
de Vries
148
mut. persicaria
de Vries
148
mut. planifolia
de Vries
148
mut. proxima
de Vries
148
mut. pustulata
de Vries
148
xindivisaHudziok 19, 136
mut.
retardata
de Vries
148
insignis Bartlett
35, 118
mut.
secunda
126
mut.
semigigas
mut.
stenophylla
mut.
tardescens
mut.
truncata de Vries
inconspecta
intermedia Gates
18, 98
xintermediaLotsy 152
irrigua Wooton
& Standl.
issleri Renner
ex Rostafiski
var. silesiacoides
43, 48, 144
Rostafiski
144, 145
& Jehlik
19, 136
lamarckiana
de Vries
148
de Vries
nanella
148
de Vries
de Vries
139
148
148
148
144
232
SYSTEMATICBOTANYMONOGRAPHS
xlanceolata
Lotsy
lata de Vries
xlaxa
133
laxiflava Renner
laxirubata
xmurivelutinaLotsy 152
152
108
de Vries
Renner
leucophylla
Gates
108
liquida de Vries
longissima
9, 10, 13, 23, 24, 33, 40, 43, 44,
58, 61-63, 166, 210
subsp. clutei
subsp.
macrosperma
(Hudziok)
magdalena
Gates
marinellae
Soldano
var. serratifolia
Gates
47
47
Hudziok
mediomarchica
110
millersii
de Vries
117
18, 98
var. nobska
octolineata
144, 145
Jehlifk & Rostafiski
multifiora
Gates
134
18, 109
L. 31, 93, 107, 121, 135, 139, 149, 150,
151, 152
(Focke)
(Shull & Bartlett)
(Boedijn)
var. ammophila
issleri
paradoxa
118
(Focke)
Stomps
So6 72
145
(Renner
Love &
ex Rudloff)
(Renner
Love &
L. 8, 9, 11, 13, 15, 16, 19, 20, 23, 26,
124-132, 137, 138, 140, 141, 142, 149, 150,
151, 152, 204, 210
subsp. germanica
(Torr. & A. Gray)
var. latifolia Asch.
18, 94
Gates
118
var. rhodoneura
Renner
var. rubricaulis
Farw. 95
145
127
72
subsp. parviflora
var. parviflora
33, 124,
Janch.
(Boedijn)
(Renner
118
ex Rudloff)
Janch.
128
33
subsp. silesiaca
(Renner)
Janch.
subsp. syrticola
(Bartlett)
Janch. 145
var. angustissima
var. muricata
xmurinella
var. oakesiana
139
(Gates) Munz
125
145
B. L. Rob.
117
Janch.
(Focke)
subsp. pachycarpa
xmuricatoidesLotsy 152
xmurilaetaLotsy 152
de Vries
96
126
128, 130
ex Rostafiski)
(Bartlett) Tischler
subsp. syrticola
var. parviflora
18, 98
Gates
subsp. angustissima
127
var. canescens
Hudziok
91, 93, 104, 106, 107, 113, 117, 121, 123,
124
(Renner) Tischler
19, 125
109
subsp. ammophila
(L.) Tischler
rubricuspis
subsp. silesica
ex Rudloff
de Vries
27,28,29,31,32,33,34,39,40,59,60,71,
ex Rostafiski)
(Renner
parviflora
148
Renner
parva Gates
47
Gates
119, 121
de Vries
oxypetala
Love &
125
Weihe
Sturtev.
pallescens
145
subsp. parviflora
subsp.
ostreae
117
Stomps
(Borbas)
subsp. pachycarpa
Love
(A. Nelson)
paralamarckiana
subsp. hungarica
Love
Stomps
125
subsp. germanica
18, 98
18, 98
22, 23
ornata
pachycarpa
subsp. ammophila
subsp.
Hudziok
Hudziok
Munz
organensis
18, 108
Gates
118
108
xoehlkersiiKappus ex Rostafiski 19, 133
52
(Munz) Rostaniski
moravica
119
(Bartlett) Gates
de Vries
oblonga
obscurifolia
Love
(Sturtev.) Gates
var. tidestromii
mollis
subsp. atrovirens
ex S. Watson
23, 26, 27, 28, 29, 31, 32, 33, 34, 40, 60, 71,
mississippensis Bartlett 144
muricata
J. W. Robbins
8, 9, 11, 13, 15, 16, 17, 18, 19, 20,
130, 131, 132, 135, 136, 137, 139, 143, 144,
145, 146, 149, 150, 151, 152, 201, 210
Hudziok
var. elliptica
8, 9, 11, 13, 16, 24, 25, 26,
78, 91, 93, 106, 107, 113, 116, 117-124,
22
de Vries
montereyensis
143, 145
94
(A. Gray)
& Coult.
18, 99
militaris
Renner
126
96
ex Rostafiski
Bartlett
oakesiana
18, 97
Link 94
media
126
27,28,29,31,33,35,39,40,88,89,90-93,
102, 106, 117, 123, 131, 132, 177, 210
120, 153
Munz
maysillesii
(Gates) Gates
61
Gates
(A. Nelson)
var. intermedia
nutans Atk. & Bartlett
Munz
Gates
(A. Nelson)
var. ornata
Gates
numismatica
33, 61, 63
33, 63
(A. Nelson)
macbrideae
19, 99
125
Gates
var. distantifolia
nuda Renner
Munz
(A. Nelson)
longissima
var. clutei
145
119
153
novae-scotiae
118
Rydb.
ex Sweet
Gutte & Rostafiski
Lotsy
xnova
19, 127
109
litorea Bartlett
108
Gates
var.fiedleri
& Gutte
Rostafiski
Hornem.
nobska Sturtev.119
119
xlinearis Lotsy 152
lipsiensis
de Vries
nissensis Rostafnski18, 97
150
de Vries
nanella
nervosa
niagarensis
150
leptocarpa
VOLUME 50
var. canescens
(Gates) Wiegand
(Torr. & A. Gray)
127
124
Farw. 72
(L.) Farw. 93
(A. Gray)
Fern. 33, 117, 128
f. parviflora
pedemontana
Soldano
perangusta
Gates
sackvillensisGates 96
35, 120
var. albiviridis
Gates
Gates
var. royfraseri
(Gates) Gates
de Vries
120
pictilaeta Renner 150
Renner
pictirubata
151
Renner
pictivelutina
plicatula
Lutz
pohliana
de Vries
de Vries
salicastrum
140
150
Renner
pictiflava
Farw. 95
sabulosa
18, 35, 99
18, 99
var. rubricalyx
xpercruciata
rydbergiiHouse 78
124
(L.) Scoggan
Soldano
pellegrinii
Desf.
ex Ser. 45, 70, 72, 142
Desf.
ex G. Don
salicifolia
J. Lehm.
110
de Vries
scitula Bartlett
semilata
sesitensis
Soldano
shulliana
Sturtev.
var. numismatica
(Bartlett) Gates
A. Gray
primiveris
procera Wooton
94
21, 89
& Standl.
79
xpseudocernuaHudziok 19, 137
xpseudochicaginensisRostafiski 19, 137
de Vries
punctulata
110
& Gutte
Rostafiski
xpurpurans
Borbas
purpurata
Kleb.
Gates
var. parviflora
Gates
f. mollis
rigilaeta
19, 127, 131, 138, 149, 151
109
simsiana
Ser. 45
xslovaca
Jehlik & Rostafnski
spathulata
de Vries
148
spectabilis
J. Lehm.
85
ex Rostafiski
145
subsp. mollis
ex Gutte & Rostafiski
robinsonii
Bartlett
125, 130
rostanskii
Jehllk
145
33, 70
(Gates) Gates
var. cheradophila
151
stubbei W. Dietr.,
de Vries
94
stucchii
151
151
rubivelutina
rubricalyx
Gates
rubricapitata
rubricaulis
mut.
151
35, 126
107, 138, 139, 150, 151
var. dentifolia
Jehlik & Rostafiski
18, 98
var. longistylis
Gutte & Rostafiski
18, 99
rubricuspis
Renner
rubrinervis
de Vries
rubrinervoides
rubritincta
ruderalis
ex Rostaniski
Gates
Gates
Bartlett
P. H. Raven & W. L. Wagner
Pers.
108
18, 109
38
17, 93, 94, 97, 99, 138, 139, 15o,
Soldano
f. erythrosepala
17, 18, 20, 94, 95, 97, 98, 99,
Kleb.
79
79
11, 13, 16, 19, 20, 23, 27, 29, 30,
var. latipetala
108
Gates
72
72
151
151
Renner
(Rydb.) Gates
Soldano
suaveolens
151
Renner
Gates
31, 33, 34, 39, 40, 82-83, 84, 85, 107, 176,
210
151
Renner
rubiundata
79
(Wooton & Standl.) Gates
var. subulifera
Renner
rubirigida
73
(Bartlett) Gates
(E. Greene)
rubiennis
151
72
145
(Bartlett ex de Vries) Gates
rubiaucta
Renner
ex Weihe
subsp. strigosa
var. procera
96
rubiplana
25, 33, 70,
(Borbas) Love & Love
var. albinervis
var. depressa
rubescensBartlett 118
rubipicta Renner
33, 70, 72
(Bartlett) Munz
Renner
var. cockerellii
18, 99
Renner
31, 33, 35, 70, 75,
(Steele) Munz
subsp. hungarica
119
Renner
rubiflava Renner
& Bush
79
rigirubata
rubipercurva
19, 141
118, 121
Bicknell
(Rydb.) Mack.
subsp. canovirens
151
royfraseri Gates
18, 98
95
subsp. cheradophila
94
Renner
108
78
96
96
Gates
Bartlett
reynoldsii
de Vries
19, 70, 73, 75
Renner
repandodentata
Renner
simplex
strigosa
94
pyramidifioraHudziok 18, 98
renneri H. Scholz
silesiaca
stenopetala
34, 95
var. cleistogama
de Vries
stenomeresBartlett 95
19, 140
Atk. & Bartlett
pycnocarpa
18, 98
108
125
Bartlett
94
45, 72, 142
45, 72, 142
de Vries
scintillans
148
96
94
salicifolia
scindens
151
109
96
salicifolia
xpolgari Rostafiski 19, 138
pratincola
pulla
233
OENOTHERA
1997
19, 127
apetala
18, 99
Javorka
(Borbas)
de Vries
148
mut.fastigiata
de Vries
149
mut.
jaculatrix
de Vries
149
mut.
lata de Vries
mut.
lutescens
mut.
sulphurea
149
de Vries
149
de Vries
xsubfalloidesLotsy 153
xsublamarckiana
sublinearis
subovata
Lotsy
de Vries
de Vries
18, 109
subpictirubata
95
subrobusta
108
Renner
de Vries
153
108
151
149
149
108
234
SYSTEMATIC
BOTANY
MONOGRAPHSVOLUME
50
subterminalis
Gates
126
subuliferaRydb. 79
superflua
de Vries
suzukiana
109
Jean & Linder
syrticola
Bartlett
var. litorea
tarda de Vries
(Rydb.) Piper 78
Mchx.
124
chrysantha
Spach
124
var. cruciata
118
18, 97
var. latifolia
Bartlett
cruciata
118
depressa
erythrosepala
europaea
94
turoviensis
Rostanski
Renner
151
hookeri
veluticurva
Renner
151
hungarica
velutiflavaRenner 151
velutiflexaRenner 152
kunthiana
guttata
135
venosa
Renner
venusta Bartlett
var. grisea
48
var. internedia
Gates
var. undulata
Gates
f. rostanskii
(Jehlik)
Thunb.
95
96
97
Jehli'k & Rostanski
33, 40, 44, 58, 67, 68-82, 106, 123, 138
subsp. cheradophila
Raven
subsp.
vrieseana
H. Lev.
Renner
78
210
Miller
Rubus
7
chersis
21
37
(L.) Scop.
asellus
perelegans
21
Trifolium
virginicum
Usoricum
146
Small
Lunell
26, 116
37
(Rydb.) Lunell
78
Xylocopa
brazilianorum
93
21
7
strigosum
Rostanski
Rupr. 37
Rupr. 37, 93
Sphinx
japonica
8,
70, 79, 81, 113, 123, 140, 153, 166,
wratislaviensis
48
Theretra
19, 138
9, 10, 13, 16, 17, 23, 24, 33, 35, 40, 43, 44,
63-68,
virginiana
Taraxacum
146
ex Rostanski
P. H. Raven, W. Dietr. & Stubbe
(Munz)
biennis
3
angustifolia
11, 16, 19, 20, 23, 24, 25, 27, 28,
79
3
Papaveraceae
Pseudo-oenothera
105, 106, 138, 172,
Rydb.
93
2, 3, 5, 6, 22, 23, 35, 72, 111
71,72,75,78-82,
210
(Rydb.) Dorn
78
6, 22
Onagraeae
Populus
var. strigosa
Onagra
Spach
Paeoniaceae
(Rydb.) W. Dietr. & P. H. Raven
29,31,33,34,38,56,58,59,69,70,
71-78, 106, 107, 113, 137, 138, 139, 140,
141, 143, 144, 145, 149, 150, 151, 167,
210
wolfii
vulgaris
124
ex Ser.) Spach 45
Spach 47
Rydb.
11, 16, 17, 23, 24, 25, 33, 43, 68, 69, 70,
subsp. villosa
xwienii
(Desf.
Onagraceae
(Bartlett) W. Dietr. & P. H.
79
strigosa
(L.) Moench
var. subulata
8, 9, 13, 15, 23, 26, 27, 28, 31, 32,
117
47
salicifolia
99
47
93
(A. Gray) Britton
parviflora
strigosa
53
94
A. Nelson
spectabilis
97
Small
71
Spach
ornata A. Nelson
Gates
var. parviflora
villosa
48
52
42
(L.) Moench
oakesiana
Bartlett
Gates & Catches.
victorinii
Spach
muricata
Small
72
(Torr. & A. Gray)
macbrideae
125, 130
133
Borbas
linkiana Spach
134, 152
Shull & Bartlett
108
(Torr. & A. Gray)
lehmanniana
velutinifoliaHudziok 19, 73
124
72
93
E. Greene
jamesii
124
Small
Small
Borbas
Spach
undirubata
de Vries
19, 127
94
(L.) Spach
(E. Greene)
tracyi Bartlett
velutirubata
Spach
Spach
93
(Nutt. ex G. Don)
tinctiundataRenner 151
xvelutina
Spach
var. parviflora
18, 109
Britton
(Nutt. ex G. Don)
var. grandifora
110
tardiflora Gates
(Nutt. ex G. Don)
var. strigosa
chrysantha
120, 123, 136, 144, 146
(Bartlett) Gates
tacikii Rostaniski
tidestromii
56, 146, 153
var. cruciata
abaniformis
varipunctata
orpifex
21
21
124
124