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 Stable URL: http://www.jstor.org/stable/25027869 Accessed: 08/06/2009 11:27 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=aspt. 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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 . :. A ::;<l:y:s : : /|WS ;w S ( ilv; r .. . . .. A.^.<SA.:.XtF : -i .;: . .4A_.i iW;^,S,.: j; ii rsu_ __ _ l_ tffl - l - rx - - __ _ _ _ | - l _ - I _l!!! _MiA * o]R_ I_I _ __ _ __ _ _ I _ U_ _- I il * :e: _ __ _ _ _ | w | _l l _I r - l w - - m>.. | | * l_ l l | __ - l == _ | _ Photo by RobynnK. Shannon L. 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_ | * __ __ III __ _ _ _11I-|_-_ | _ w _ _ _ w _r| . _ , . . w . / . s . . . . .. . t _ _b . . _ . . . I Ii . .. _ I s , 1_ 0 | |I j|| || || li8 I I |i X | ..111 || |I _ l o_ w * I I . i .. w .* i . i i ls I IE . iI |,E | .. ..- - ._ ., *__ VOLUME 50 | . . | | I . . _ - _ * _ _ I ! . _S_ ,^_ _ t_ - _ | -. t 1E * _ _|1 __ _-! _ S. I _ w \ _J | l _-:s== \ S S | _'- - - | .,_ S_ - . s . __ *s \ I | 111|_ I L .... _ _ l _ ,. _ _ -| _ _/ - _ _ |' | _ __ __s 1-I -i _ *1 s_ . \ ___ / s _\ _| _ / . _ | I hypothetiods \ | zoestor / | _ 1 | _ _ ! Nplastome IIJ __ . . _ l It __ _ // ./ ' I - _: _ ..._ _ /. i_ . , _|_ . _ . | _ | SS * x J _ _l _. / _ _ __ _i / s . s hypotheXcalw _t t | I _____ _ . . . _ J _ t _! 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. LITERATURECITED Aiton, W. 1789. Hortus or a catalogue kewensis; of the plants in the Royal cultivated London:George Nicol. Bartlett, H. H. 1914. 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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