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Onagraceae - The Evening Primrose Family

Xylonagra arborea Chylismia claviformis subsp. lancifolia Epilobium ciliatum watsonii Oenothera riskindii Gayophytum diffusum subsp. diffusum Chamerion angustifolium subsp. circumvagum Taraxia subacaulis Fuchsia pringsheimii Camissoniopsis bistorta Lopezia grandiflora subsp. macrophylla Hauya elegans subsp. cornuta Clarkia breweri Megacorax gracielanus Ludwigia peploides subsp. glabrescens Circaea lutetiana subsp. canadensis Eulobus californicus Gongylocarpus fructiculosus Tribe Hauyeae Tribe Circaeeae Tribe Lopezieae Tribe Gongylocarpeae Tribe Epilobieae Tribe Onagreae Holmgrenia Camissoniopsis Tetrapteron Camissonia Eremothera Oenothera Chylismia Subfamily Ludwigioideae Subfamily Onagroideae Eulobus Taraxia Chylismiella Gayophytum Clarkia Xylonagra Epilobium Chamerion Gongylocarpus Megacorax Lopezia Circaea Fuchsia Hauya Ludwigia
Oenothera glaucifolia

Introduction

In the site we present a full checklist of all taxa within each of the 18 recognized genera as well as diagnostic images for as many taxa as possible.

The new classification is based on a number of phylogenetic and revisionary publications and results of a recent project initiated at the Smithsonian Institution. A group of Onagraceae researchers worked together to gain a better understanding of the evolution of the family and to derive an appropriate classification. Using molecular, morphological, biochemical, and biogeographic tools, results have provided new insights into family relationships and monophyletic groups at the tribal and generic levels.

Four papers were published from this project. A more complete bibliography for the Onagraceae family is provided.

    Katinas, L., J. Crisci, W.L. Wagner, & P.C. Hoch. 2004. Geographical diversification of tribes Epilobieae, Gongylocarpeae, and Onagreae (Onagraceae) in North America, based on parsimony analysis of endemicity and track compatibility analysis. Ann. Missouri Bot. Gard. 91: 159-185. (pdf)

    Levin, R.A., W.L. Wagner, P.C. Hoch, M. Nepokroeff, J.C. Pires, E.A. Zimmer, & K.J. Sytsma. 2003. Family-level relationships of Onagraceae based on chloroplast rbcL and ndhL data. Amer. J. Bot. 90: 107-115. (pdf)

    Levin, R.A., W.L. Wagner, P.C. Hoch, W.J. Hahn, A. Rodriguez, D.A. Baum, L. Katinas, E.A. Zimmer, & K.J. Sytsma. 2004. Paraphyly in Tribe Onagreae: Insights into phylogenetic relationships of Onagraceae based on nuclear and chloroplast sequence data. Syst. Bot. 29: 147-164. (pdf)

    Wagner, W.L. 2004. Resolving a nomenclatural and taxonomic problem in Mexican Oenothera sect. Hartmannia (Tribe Onagreae, Onagraceae). Novon 14: 124-133. (pdf)

Smithsonian Mellon Fellows:
  Jorge V. Crisci, Museo de La Plata
  Peter C. Hoch, Missouri Botanical Garden
  Liliana Katinas, Museo de La Plata
  Rachel A. Levin, Smithsonian postdoc
  Robert A. Raguso, University of South Carolina -Columbia
  Kenneth J. Sytsma, University of Wisconsin -Madison

Smithsonian Scientists:
  Warren L. Wagner, Curator of Pacific Botany
  Elizabeth A. Zimmer, Curator, Laboratories of Analytical Biology

Junior Fellows:
  Richard Clinebell, Missouri Botanical Garden

This web site was conceived by Warren L. Wagner and co-written with Peter C. Hoch. Site development was by Denise Mix with assistance from Jessica Braun in image preparation and database maintenance. Click here for information on how to reference this Web site.
Fuchsia splendens

Phylogeny

Despite intensive morphological and molecular studies of Onagraceae, relationships within the family have not been fully understood. One drawback of previous molecular analyses was limited sampling within the large tribe Onagreae. In addition, the monophyly of two species-rich genera in Onagreae, Camissonia and Oenothera, has never been adequately tested. To understand relationships within Onagraceae, test the monophyly of these two genera, and ascertain the affinities of the newly discovered genus Megacorax, parsimony and maximum likelihood analyses with rbcL and ndhF sequence data for 24 taxa representing all 17 Onagraceae genera and two outgroup Lythraceae were conducted [Levin et al. 2003 (pdf)]. Results strongly support a monophyletic Onagraceae, with Ludwigia as the basal lineage and a sister-taxon relationship between Megacorax and Lopezia. Gongylocarpus is supported as sister to Epilobieae plus the rest of Onagreae, although relationships within the latter clade have limited resolution. Thus, we advocate placement of Gongylocarpus in a monogeneric tribe, Gongylocarpeae. Most relationships within Onagreae are weakly resolved, suggesting a rapid diversification of this group in western North America. Neither Camissonia nor Oenothera appears to be monophyletic. Morphological characters generally agree with the molecular data, providing further support for relationships.

In a second study study [Levin et al. 2004 (pdf)], DNA sequence data from one nuclear region (ITS) and two chloroplast regions (trnL-trnF and rps16) were used to infer phylogenetic relationships among 93 taxa, with concentrated sampling in the large tribe Onagreae. Results strongly support tribe Gongylocarpeae as sister to tribes Epilobieae + Onagreae, both of which are monophyletic. Within Onagreae, Camissonia appears to be broadly paraphyletic, as does Oenothera. In Oenothera there appear to be two lineages, one of which has Calylophus and Gaura + Stenosiphon nested within it. At the base of the Onagraceae phylogeny, we have clarified previous confusion regarding conflicting placements of Hauya and Lopezia based on nuclear versus chloroplast data. Results of these analyses suggest the need for new taxonomic delimitations that will be presented in comprehensive revised classification of the family Onagraceae based on molecular sequence data and morphology by W. L. Wagner, P. C. Hoch and P. H. Raven.

Onagraceae Bibliography
 

Baum, D. A., K. J. Sytsma & P. C. Hoch. 1994. The phylogeny of Epilobium (Onagraceae) based on nuclear ribosomal DNA sequences. Syst. Bot. 19: 363-388.

Berry, P. E. 1982. The systematics and evolution of Fuchsia sect. Fuchsia (Onagraceae). Ann. Missouri Bot. Gard. 69: 1-198.

Berry, P. E. 1985. The systematics of the apetalous Fuchsias of South America, Fuchsia sect. Hemsleyella (Onagraceae). Ann. Missouri Bot. Gard. 72: 213—251.

Berry, P. E. 1989. A systematic revision of Fuchsia sect. Quelusia (Onagraceae). Ann. Missouri Bot. Gard. 76: 532-584.

Berry, P. E., W. J. Hahn, K. J. Sytsma, J. C. Hall & A. Mast. 2004. Phylogenetic relationships and biogeography of Fuchsia (Onagraceae) based on noncoding nuclear and chloroplast DNA data. Amer. J. Bot. 91: 601-614.

Berry, P. E., J. J. Skvarla, A. D. Partridge, & M. K. Macphail. 1990. Fuchsia pollen from the Tertiary of Austral. Australian Syst. Bot. 3: 739–744.

Berry, P. E., B. A. Stein, S. Carlquist & J. W. Nowicke. 1988. Fuchsia pachyrrhiza, a new tuberous species and section of Fuchsia from northwestern Peru. Syst. Bot. 13: 483—492.

Boufford, D. E. 1982[1983]. The systematics and evolution of Circaea (Onagraceae). Ann. Missouri Bot. Gard. 69: 804-994.

Boufford, D. E., J. V. Crisci, H. Tobe & P. C. Hoch. 1990. A cladistic analysis of Circaea (Onagraceae). Cladistics 6: 171-182.

Breedlove, D. E. 1969. The systematics of Fuchsia section Encliandra (Onagraceae). Univ. Calif. Publ. Bot. 53: 1-69.

Breedlove, D. E., P. E. Berry & P. H. Raven. 1982. The Mexican and Central American species of Fuchsia (Onagraceae) except for sect. Encliandra. Ann. Missouri Bot. Gard. 69: 209-234.

Bult, C. J. & E. A. Zimmer. 1993. Nuclear ribosomal RNA sequences for inferring tribal relationships within Onagraceae. Syst. Bot. 18: 48-63.

Carlquist, S. 1975. Wood anatomy of Onagraceae, with notes on alternative modes of photosynthate movement in dicotyledon woods. Ann. Missouri Bot. Gard. 62: 386—424.

Carlquist, S. 1977 [1978]. Wood anatomy of Onagraceae: additional species and concepts. Ann. Missouri Bot. Gard. 64: 627-637.

Carlquist, S. 1982 [1983]. Wood anatomy of Onagraceae: further species; root anatomy; significance of vestured pits and allied structures in dicotyledons. Ann. Missouri Bot. Gard. 69: 755-769.

Carlquist, S. & P. H. Raven. 1966. The systematics and anatomy of Gongylocarpus (Onagraceae). Amer. J. Bot. 53: 378-390.

Carr, B. L., J. V. Crisci & P. C. Hoch. 1990. A cladistic analysis of the genus Gaura (Onagraceae). Syst. Bot. 15: 454-461.

Chen, C. J., P. C. Hoch & P. H. Raven. 1992. Systematics of Epilobium (Onagraceae) in China. Syst. Bot. Monogr. 34: 1-209.

Cleland, R. E. 1972. Oenothera: Cytogenetics and Evolution. London: Academic Press.

Clinebell, R. R., A. Crowe, D. P. Gregory & P. C. Hoch. 2004. Pollination ecology of Gaura and Calylophus (Onagraceae, tribe Onagreae) in western Texas, U.S.A. Ann. Missouri Bot. Gard. 91: 369-400.

Conti, E., A. Fischbach & K. J. Sytsma. 1993. Tribal relationships in Onagraceae: implications from rbcL sequence data. Ann. Missouri Bot. Gard. 80: 672-685.

Conti, E., A. Litt & K.J. Sytsma. 1996. Circumscription of Myrtales and their relationships to other rosids: evidence from rbcL sequence data. Amer. J. Bot. 83: 221-233.

Conti, E., A. Litt, P. G. Wilson, S. A. Graham, B. G. Briggs, L. A. S. Johnson & K. J. Sytsma. 1997. Interfamilial relationships in Myrtales: molecular phylogeny and patterns of morphological evolution. Systematic Botany 22: 629-647.

Crisci, J. V., E .A. Zimmer, P. C. Hoch, G. B. Johnson, C. Mudd & N. Pan. 1990. Phylogenetic implications of ribosomal DNA restriction site variation in the plant family Onagraceae. Ann. Missouri Bot. Gard. 77: 523-538.

Dahlgren, R. & R. F. Thorne. 1984. The order Myrtales: circumscription, variation, and relationships. Ann. Missouri Bot. Gard. 71:633-699.

Dietrich, W. 1977 [1978]. The South American species of Oenothera sect. Oenothera (Raimannia, Renneria; Onagraceae). Ann. Missouri Bot. Gard. 64: 425-626.

Dietrich, W., P. H. Raven & W. L. Wagner. 1985. Revision of Oenothera sect. Oenothera subsect. Emersonia (Onagraceae). Syst. Bot. 10: 29-48.

Dietrich, W. & W.L. Wagner. 1988. Systematics of Oenothera section Oenothera subsection Raimannia and subsection Nutantigemma (Onagraceae). Syst. Bot. Monogr. 24: 1-91.

Dietrich, W., W. L. Wagner & P. H. Raven. 1997. Systematics of Oenothera sect. Oenothera subsect. Oenothera (Onagraceae). Syst. Bot. Monogr. 50: 1-234.

Eyde, R. H. 1977 [1978]. Reproductive structures and evolution in Ludwigia (Onagraceae). I. Androecium, placentation, meristem. Ann. Missouri Bot. Gard. 64: 644-655.

Eyde, R.H. 1978 [1979]. Reproductive structures and evolution in Ludwigia (Onagraceae). II. Fruit and seed. Ann. Missouri Bot. Gard. 65: 656-675.

Eyde, R.H. 1981 [1982]. Reproductive structures and evolution in Ludwigia (Onagraceae). III. Vasculature, nectaries, conclusions. Ann. Missouri Bot. Gard. 68: 379-412.

Eyde, R. H. 1982 [1983].  Evolution and systematics of the Onagraceae:  floral anatomy. Ann. Missouri Bot. Gard. 69: 735-747.

Eyde, R. H. & J. T. Morgan. 1973. Floral structure and evolution in Lopezieae (Onagraceae). Amer. J. Bot. 60: 771-787.

Ford, V. S. & L. D. Gottlieb. 2003. Reassessment of phylogenetic relationships in Clarkia sect. Sympherica. Amer. J. Bot. 90: 284-292.

González Elizondo, M. S., I. L. López Enriquez & W. L. Wagner. 2002. Megacorax gracielanus (Onagraceae), a new genus and species from Durango, México. Novon 12: 360-365.

Gordon, K. H. J., E. Crouse, E. Bohnert, and H. Herrmann.  1981.  Restriction  endonuclease cleavage site maps of chloroplast DNA from Oenothera parviflora (Euoenothera plastome IV).  Theor. Appl. Genet. 59: 281-296.

_____.  1982.  Physical mapping of differences in chloroplast DNA of the five wild-type plastomes in Oenothera subsect. Euoenothera.  Theor. Appl. Genet. 61: 373-384.

Gottlieb, L. D. 1986. Genetic differentiation, speciation and phylogeny in Clarkia (Onagraceae). In K. Iwaksuki, P. H. Raven, and W. J. Bock [eds.], Modern aspects of species, 145-160. University of Tokyo Press, Tokyo, Japan.

Gottlieb, L. D. & V. S. Ford. 1996. Phylogenetic relationships among the sections of Clarkia (Onagraceae) inferred from the nucleotide sequences of PgiC. Syst. Bot. 21: 45-62.

Gottlieb, L. D., and N. F. Weeden. 1979. Gene duplication and phylogeny in Clarkia. Evolution 33:1024-1039.

Graham, S. A., and T. B. Cavalcanti. 2001. New chromosome counts in the Lythraceae and a review of chromosome numbers in the family. Syst. Bot. 26: 445-458.

Gregory, D. P. 1963. Hawkmoth pollination in the genus Oenothera. Aliso 5: 375-384.

Gregory, D. P. 1964. Hawkmoth pollination in the genus Oenothera. Aliso 5: 385-419.

Heslop-Harrison, Y. 1990. Stigma form and surface in relation to self-incompatibility in the Onagraceae. Nordic Journal of Botany 10: 1-19.

Hoch, P. C., J. V. Crisci & H. Tobe 1993. A cladistic analysis of the genus Lopezia (Onagraceae). Bot. J. Linn. Soc. 111: 103-116.

Hoch, P. C., J. V. Crisci, H. Tobe & P. E. Berry. 1993. A cladistic analysis of the plant family Onagraceae. Syst. Bot. 18: 31-47.

Hoch, P. C. & P. H. Raven. 1992. Boisduvalia, a coma-less Epilobium (Onagraceae). Phytologia 73: 456-459.

Hoggard, G. D., P. J. Kores, M. Molvray & R. K. Hoggard. 2004. The phylogeny of Gaura (Onagraceae) based on ITS, ETS and TrnL-F sequence data. Amer. J. Bot. 91: 139-148.

Holsinger, K. E., & N. C. Ellstrand.  1984.  The evolution and ecology of permanent translocation heterozygotes.  Amer. Naturalist 124: 48‑71.

Holsinger, K. E., & M. W. Feldman.  1981.  A single locus model of selection in permanent translocation heterozygotes.  Theoretical Population Biology 20: 218-240.

Holsinger, K. E. & H. Lewis. 1986. Description of a new section and subsection in Clarkia (Onagraceae). Ann. Missouri Bot. Gard. 73; 491-494.

Holub, J. 1972. Taxonomic and nomenclatural remarks on Chamaenerion auct. Folia Geobot. Phytotax. 7: 81—90.

Hupfer, H., M. Swiatek, S. Hornung, R .G. Herrmann, R. M. Maier, W.-L. Chiu & B. Sears. 2000. Complete nucleotide sequence of the Oenothera elata plastid chromosome, representing plastome I of the five distinguishable Euoenothera plastomes. Molecular and General Genetics 263: 581-585.

Johnson, L. A. S. & B. G. Briggs. 1984. Myrtales and Myrtaceae: A phylogenetic analysis. Ann. Missouri Bot. Gard. 71: 700-756.

Katinas, L., J. Crisci, W. L. Wagner & P. C. Hoch. 2004. Geographical diversification of tribes Epilobieae, Gongylocarpeae, and Onagreae (Onagraceae) in North America, based on parsimony analysis of endemicity and track compatibility analysis. Ann. Missouri Bot. Gard. 91: 159-185.

Keating, R. C. 1982 [1983]. The evolution and systematics of Onagraceae: leaf anatomy. Ann. Missouri Bot. Gard. 69: 770-803.

Klein, W. M. 1964. A biosystematic study of four species of Oenothera subgenus Anogra. University of Michigan Dissertation Services, Ann Arbor, Michigan. pp. 1-191.

Klein, W. M. 1970. The evolution of three diploid species of Oenothera subgenus Anogra (Onagraceae). Evolution. 24: 578-597.

Kurabayashi, M., H. Lewis & P. H. Raven. 1962. A comparative study of mitosis in the Onagraceae. Amer. J. Bot. 9: 1003-1026.

Levin, R. A., W. L. Wagner, P. C. Hoch, W. J. Hahn, A. Rodriguez, D. A. Baum, L. Katinas, E. A. Zimmer & K. J. Sytsma. 2004. Paraphyly in Tribe Onagreae: Insights into phylogenetic relationships of Onagraceae based on nuclear and chloroplast sequence data. Syst. Bot. 29: 147-164.

Levin, R. A., W. L. Wagner, P. C. Hoch, M. Nepokroeff, J. C. Pires, E. A. Zimmer & K. J. Sytsma. 2003. Family-level relationships of Onagraceae based on chloroplast rbcL and ndhF data. Amer. J. Bot. 90: 107-115.

Lewis H. & M. E. Lewis. 1955. The genus Clarkia. Univ. Calif. Publ. Bot. 20: 241-392.

Lewis, H. & P.H. Raven. 1992. New combinations in the genus Clarkia (Onagraceae). Madroño 39: 163-169.

Lewis H. & J. Szweykowski. 1964. The genus Gayophytum (Onagraceae). Brittonia. 16: 343-391.

Linsley, E. G.,  J. W. MacSwain, and P. H. Raven.  1964. Comparative behavior of bees and Onagraceae.  III. Oenothera bees of the Mojave Desert, California.  Univ. California Publ. Entomol. 33: 59-98.

Linsley, E. G., J. W. MacSwain, P. H. Raven, and R. W. Thorp.  1973.  Comparative behavior of bees and Onagraceae.  V.  Camissonia and Oenothera bees of cismontane California and Baja California.  Univ. Calif. Publ. Entomol.  71: xxx-xxx.

Martin, H. A. 2003.  The history of the family Onagraceae in Australia and its relevance to biogeography. Austral. J. Bot. 51: 585-598.

Martin, P. G. & J. M. Dowd. 1986. Phylogenetic studies using protein sequences within the order Myrtales. Annals of the Missouri Botanical Garden 73: 442-448.

MacSwain, J. W., P. H. Raven, and R. W. Thorp. 1973. Comparative behavior of bees and Onagraceae. IV. Clarkia bees of the western United States. Univ. Calif. Publ. Entomol. 70: 1-80.

Munz, P.A. 1931. Studies in Onagraceae VI. The subgenus Anogra of the genus Oenothera. Amer. J. Bot. 18: 309-327.

Munz, P.A.  1931.  Studies in Onagraceae VII.  The subgenus Pachylophus of the genus Oenothera.  Amer. J. Bot. 18: 728-738.

Munz, P. A. 1932. Studies in Onagraceae VIII. The subgenera Hartmannia and Gauropsis of the genus Oenothera. The genus Gayophytum. Amer. J. Bot. 19: 755-778.

Munz, P. A.  1949.  The Oenothera hookeri group.  Aliso 2: 1‑47.

Munz, P. A. 1965. Onagraceae. Flora North America. II. 5: 1-278.

Nowicke, J. W. J. J. Skvarla, P. H. Raven, & P. E. Berry.  1984. A palynological study of the genus Fuchsia (Onagraceae). Ann. Missouri Bot. Gard. 71: 35-91.

O'Kane, S. L. & B. A. Schaal. 1998. Phylogenetics of Lopezia (Onagraceae): evidence from chloroplast DNA restriction sites. 23: 5-20.

Peng, C.-I. 1989. The systematics and evolution of Ludwigia sect. Microcarpium (Onagraceae). Ann. Missouri Bot. Gard. 76: 221-302.

Peng, C.-I. 1988. The biosystematics of Ludwigia sect. Microcarpium (Onagraceae). Ann. Missouri Bot. Gard. 75: 970-1009.

Peng, C.-I. and H. Tobe. 1987. Capsule wall anatomy in relation to capsular dehiscence in Ludwigia sect. Microcarpium (Onagraceae). Amer. J. Botany 74: 1102-1110.

Plitmann, U., P. H. Raven & D. E. Breedlove. 1973. The systematics of Lopezia (Onagraceae). Ann. Missouri Bot. Gard. 60: 478-563.

Plitmann, U., P. H. Raven, W. Tai & D. E. Breedlove. 1975. Cytological studies in Lopezieae (Onagraceae). Bot. Gaz. (Crawfordsville) 136: 322-332.

Praglowski, J., J. Nowicke, P. H. Raven, J. Skvarla & W. L. Wagner; taxonomy by P.H. Raven and W. L. Wagner. 1987. Onagraceae Juss.: Onagreae R. Raimann pro parte. World Pollen and Spore Flora 15: 1-55.

Praglowski, J., J. Nowicke, P.H. Raven, J.J. Skvarla & P.C. Hoch. 1988. Onagraceae Juss.: Onagreae R. Raimann pro parte, Lopezieae Spach. World Pollen and Spore Flora 16: 1—35.

Praglowski, J., J. W. Nowicke, J. J. Skvarla, P. C. Hoch, P. H. Raven & M. Takahashi. 1994. Onagraceae Juss.: Circaeeae DC., Hauyeae Raimann, Epilobieae Spach. World Pollen and Spore Flora 19: 1-38.

Praglowski, J., J. J. Skvarla, P. H. Raven & J. Nowicke. 1983. Onagraceae Juss.: Fuchsiae L./ Jussiaeeae L. World Pollen and Spore Flora 12: 1—41.

Raguso, R. A. & E. Pichersky. 1995. Floral volatiles of Clarkia breweri and C. concinna (Onagraceae): recent evolution of floral aroma and moth pollination. Plant Syst. Evol. 194: 55-67.

Raguso, R. A. & M. A. Willis. Submitted. Behavioral responses of wild hawkmoths to floral advertisements and rewards. Oikos.

Ramamoorthy, T.P. 1979. A sectional revision of Ludwigia sect. Myrtocarpus s. lat. (Onagraceae). Ann. Missouri Bot. Gard. 66: 893-896.

Ramamoorthy, T. P. & E. M. Zardini. 1987. The systematics and evolution of Ludwigia sect. Myrtocarpus sensu lato (Onagraceae). Monogr. Syst. Bot. Missouri Bot. Gard. 19: 1-120.

Raven, P. H. 1962. The systematics of Oenothera subgenus Chylismia. Univ. Calif. Publ. Bot. 34: 1-122.

Raven, P. H. 1963. The old world species of Ludwigia (including Jussiaea), with a synopsis of the genus (Onagraceae). Reinwardtia. 6: 327-427.

Raven, P. H. 1964. The generic subdivision of Onagraceae, tribe Onagreae. Brittonia 16: 276-288.

Raven, P. H. 1969. A revision of the genus Camissonia (Onagraceae). Contr. U. S. Natl. Herb. 37: 161-396.

Raven, P. H. 1976. Generic and sectional delimitation in Onagraceae, tribe Epilobieae. Ann. Missouri Bot. Gard. 63: 326-340.

Raven, P. H. 1977. A new species of Lopezia (Onagraceae) from Sinaloa, Mexico. Ann. Missouri Bot. Gard. 64: 638-641.

Raven, P. H. 1979. A survey of reproductive biology in Onagraceae. New Zealand J. Bot. 17: 575-593.

Raven, P. H. 1988. Onagraceae as a model of plant evolution. Pp. 85-107 in L.D. Gottlieb & S.K. Jain (eds.), Plant Evolutionary Biology. A symposium honoring G. Ledyard Stebbins. London: Chapman, and Hall.

Raven, P. H. & D. I. Axelrod. 1978. Origin and relationships of the California flora. Univ. Calif. Publ. Bot. 72: 1-134.

Raven, P. H. & D. E. Breedlove. 1975. New combinations in the genus Hauya (Onagraceae). Ann. Missouri Bot. Gard. 62: 510.

Raven, P. H., W. Dietrich & W. Stubbe. 1979 [1980]. An outline of the systematics of Oenothera subsect. Euoenothera (Onagraceae). Syst. Bot. 4: 242-252.

Raven, P. H. & D. P. Gregory. 1972. A revision of the genus Gaura (Onagraceae). Mem. Torr. Bot. Club. 23: 1-96.

Raven, P. H. & D. M. Moore. 1965. A revision of Boisduvalia (Onagraceae). Brittonia 17: 238-254.

Raven, P. H. & D. R. Parnell. 1970. Two species and some nomenclatural changes in Oenothera subg. Hartmannia (Onagraceae). Madroño. 20: 246-249.

Raven, P. H. & T. E. Raven. 1976. The genus Epilobium (Onagraceae) in Australasia: a systematic and evolutionary study. New Zealand Dept. Sci. Industr. Res. Bull. 216: 1-321.

Raven, P. H. & W. Tai. 1979. Observations of chromosomes in Ludwigia (Onagraceae). Ann. Missouri Bot. Gard. 66: 862-879.

Seavey, S. R., R. E. Magill & P. H. Raven. 1977. Evolution of seed size, shape, and surface architecture in the tribe Epilobieae (Onagraceae). Ann. Missouri Bot. Gard. 64: 18-47.

Skvarla, J. J., P. H. Raven, W. F. Chissoe & M. Sharp. 1978. An ultrastructural study of viscin threads in Onagraceae pollen. Pollen & Spores 20: 5-143.

Skvarla, J. J., P. H. Raven & J. Praglowski. 1975. The evolution of pollen tetrads in Onagraceae. Amer. J. Bot. 62: 6-35.

Skvarla, J.J., P.H. Raven, & J. Praglowski. 1976. Ultrastructural survey of Onagraceae pollen. Pp. 447-479 in Ferguson, I.K. & J. Muller (eds.), The evolutionary significance of the exine. Linn. Soc. Symp. Ser. 1.

Small, E. 1971. The systematics of Clarkia, section Myxocarpa. Canad. J. Bot. 49: 1211-1217.

Smith, J. D. & J. N. Rose. 1913. A monograph of the Hauyeae and Gongylocarpeae, tribes of the Onagraceae. Contr. U. S. Natl. Herb. 16: 287-298.

Solomon, J.C. 1982. The systematics and evolution of Epilobium (Onagraceae) in South America. Ann. Missouri Bot. Gard. 69: 239-335.

Spach, E.  1835a.  Histoire Naturelle des Végétaux. Phanérogames. 4: 1-446.

Spach, E.  1835b.  Synopsis Monographia Onagrearum.  Annales des Sciences Naturelles, Botanique ser. 2,  4: 161-178.

Spach, E.  1835c [1836].  Monographia Onagrearum.  Nouvelles Annales du Museum d'Histoire Naturelle 4: 320-407 [408], pl.30-31.

Straley, G. B. 1977 [1978]. Systematics of Oenothera sect. Kneiffia (Onagraceae). Ann. Missouri Bot. Gard. 64: 381-424.

Stubbe, W. 1964.  The role of the plastome in the evolution of the genus Oenothera.  Genetica 35: 28‑33.

Stubbe, W. and P. H. Raven.  1979.  A genetic contribution to the taxonomy of Oenothera sect. Oenothera (including subsects. Euothera, Emersonia, Raimannia, and Munzia).  Plant Systematics and Evolution 133: 39-59.

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