oirs

of

The Torrey Botanical Club

Volume 23 Number 1 December 1972

Editor

William Louis Stem

A REVISION OF THE GENUS GAURA
(ONAGRACEAE)

Peter H. Raven and David P. Gregory

LI

JAN 31 1973

RARY

NEW YORK

fiOTAKiCAL ;»Ar?DECM '

Memoirs

of the

Torrey Botanical Club

Volume 23 Number 1
December 1972

Manuscripts intended for the Memoirs and related
editorial correspondence should be sent to Professor
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A REVISION OF THE GENUS GAURA
(ONAGRACEAE)i

Peter PI. Raven and David P. Gregory

Missouri Botanical Garden and Washington University, St. Louis, Missouri
and

Keene State College, Keene, New Hampshire

1 This paper is respectfully dedicated to Dr. Philip A. Munz, who ha

soft. K°?i eidge °f Athe.,fa™>y Onagraceae during a lifetime of devoted stuc
80th birthday, 1 April 1972.

contributed so much to
on the occasion of his

A REVISION OK THE GENUS GAURA
(ONAGRACEAE)

Among the 11 genera of Onagraceae, tribe Onagreae, Gaura can be
recognized by the divided sporogenous tissue in its anthers; its predominant¬
ly vespertine flowering; nutlike, indehiscent capsules; and, in most species,
the presence of a flap of sterile tissue near the base of each filament
(Raven, Brittonia 16: 276-288. 1964; Carlquist and Raven, Amer. J. Bot.
53: 378-390. 1966). It is probably most closely related to the other genera
with divided sporogenous tissue: Calylophus, Clarkia, Heterogaura, and
perhaps Hauya. Like Calylophus, the genus Gaura centers in the southern
plains of the United States and adjacent Mexico, especially in and around
Texas, with a few mostly derivative species endemic beyond this region.

Gaura first became known to science through one of these derivative
species, G. biennis. Linnaeus grew this species in the botanical garden at
Uppsala from seeds that were collected in eastern Pennsylvania and
brought to him by his student Peter Kalm, who collected in the area from
1748 to 1750. Linnaeus described the new genus in a thesis defended by
another of his students, Leonhard Joh. Chenon, and published it in 1751.
Here and in his later references to Gaura, Linnaeus erroneously included
as a synonym Plukenet’s “Lysimachia lutea angustifolia virginiana, flore
minore,” a concept that Plukenet had presented in his Almagestum botan -
icum ... in 1696. The Virginian plants that Plukenet had were actually
the plant that we now know as Oenothera biennis L. or a related species,
and it is clear that Linnaeus based his concept of Gaura entirely on the
living plants that he had grown himself from Kalin’s seeds.

Additional species of Gaura were discovered along the eastern seaboard
of the United States, and by the expeditions associated with the names of
Alexander von Humboldt and Martin de Sesse y Lacasta and Jose Mariano
Mocino in Mexico. Still more were added by David Douglas, Thomas
Nuttall, and Thomas Drummond, among others, in their botanical explor¬
ations west of the Mississippi. By 1864, Rothrock was able to list 13 species
in his synopsis of the group (Proc. Amer. Acad. Arts 6: 347-354), in which
he established the genus with its present circumscription bv separating
the monotypic Californian Heterogaura and reaffirming the distinctive
characteristics of Spach’s genus Stenosiphon. The latter comprises a single
species of the southern plains of the United States which we consider more
closely allied to Oenothera than to Gaura (Raven, Brittonia 16: 276-288.
1964), whereas Heterogaura is now usually considered a derivative of

2 A REVISION OF THE GENUS CAURA ( ONAGRACEAE )

Clarkia. They were confused with Gaura at one time only because of their
nutlike, indehiscent fruits, which are in fact entirely different in structure.

Joseph Rothrock was a prespicacious observer. The material he had
has now been increased many hundredfold, and yet the 13 species he
recognized are all included in the present revision, all but one with the
same names. In addition, the main groupings of species he set up appear
reasonable, although we have subdivided two of them in the light of a
more complete understanding of the species involved. After the publication
of Rothrock’s synopsis, a few more species of Gaura were discovered, and
a great deal of information was accumulated about the species that he
knew. In 1938, P. A. Munz provided a much more extensive revision as
one of his invaluable “Studies in Onagraceae” (Bull. Torrey Bot. Club 65:
105-122, 211-228. 1938), presenting in convenient form a critical review
of what had been accomplished in the more than 70 years since the publi¬
cation of Rothrock’s brief synopsis. In 1965, Dr. Munz emended his 1938
treatment slightly in reviewing the genus Gaura again, this time for his
monograph of the North American species of the family (N. Amer. FI.
II. 5: 1-231).

Our own studies of the genus Gaura have occupied much of our atten¬
tion for the past 15 years. During the course of our work, we have studied
all of the species in the field, usually at several or in some cases up to
hundreds of localities. We have observed their pollination systems and
floral biology at one or more localities. All of the species have been grown
in the experimental garden, and hybrids have been made in many of the
possible combinations. Hundreds of determinations of chromosome number
and meiotic configuration have been made ( Raven and Gregory, Brittonia
24: 71-86. 1972). All of this information has been brought to bear in the
present paper upon the taxonomy of the genus, and it has naturally made
possible some consolidation and reorganization of the taxa, as well as the
recognition of a few hitherto undetected species. Our experimental data will
not, however, be presented in the current paper, which is designed to pro¬
vide a basis for subsequent publications on different aspects of the genus
Gaura.

PHYLOGENETIC RELATIONSHIPS

We have in the current paper divided the 21 species of Gaura we
recognize into eight sections. We visualize these sections as homogeneous
groups of closely related species, and important evolutionary units. Insofar
as possible, we have set them up so as to be directly comparable with the
sections recognized by Lewis and Lewis in their outstanding monograph of
the genus Clarkia (Univ. Calif. Publ. Bot. 20: 241-392. 1955). In his

MEMOIRS OF THE TORREY BOTANICAL CLUB

3

1938 revision, Munz grouped the species of Gaura into only two sections,
one including only G. mutabilis. Although this species has yellow flowers,
a long floral tube, and is pollinated by hawkmoths, we do not consider it
much more distinct than some of the other sections we have recognized
here, and we believe that sections based upon small, closely related as¬
semblages of species provide a useful basis for discussions of a genus for
which sufficient information is available to make the recognition of such
sections possible.

W ithin Gaura, a clumped, perennial habit is clearly primitive, and is
shared with the more generalized members of the relatively closely related
genus Calylophus. Rhizomatous perennials and also annuals are specialized
in this respect. The fruits of all species of Gaura are fundamentally similar,
with thick, tough walls and usually four ovules (occasionally up to eight)
and very thin partitions between the loeules that do not persist to maturity.
^ ithin this fundamental ground plan, we have assumed that more complex
morphologies have been derived: ornate shape (sections Campo gaura,
Xenogaura), an evident stipe (section Stipogaura ), or obvious wings on
the angles (section Pterogaura ). In addition, the obviously 4-angled fruits
of section Gaura are held to be more specialized than the more nearly terete
ones of what we would regard as generalized species.

These two trends— in habit and fruit shape— are correlated not only with
one another but also with other morphological features and with the
geographical distribution of the respective groups in suggesting a pattern
of phylogeny for the genus as a whole. Only a few species have elongate,
subterete fruits lacking wings, stipes, or other obvious elaborations (figs.
1-4). Three of these-Gm/ra mutabilis (section Gauridium ) and G. macro-
carpa and G. boquillensis (section Xero gaura)— are clumped perennials.
These three species are colonial, occurring in scattered localities in west
Texas, Nuevo Leon, and Coahuila, and south through Chihuahua to the
Sierra Madre Occidental and the Trans-Mexican Volcanic Belt. Gaura
mutabilis is the only species of the genus with yellow flowers. It has a
floral tube that is very narrow and 26-42 mm long, in contrast to the other
species of the genus, in which the floral tube is 1.5-15 mm long. At the
base of each filament, there is no scale closing the mouth of the floral tube,
as there is in all other species of the genus. This species is the only member
of the genus that is regularly pollinated by hawkmoths, and it may be
primitive in its long floral tube and lack of scales, as well as in its yellow

flowers. All of these features are found in some species of the related
genus Calylophus.

The two species of section Xerogaura differ from Gaura mutabilis in
their white flowers, relatively short floral tube, and scale at the base of each

4

A REVISION OF THE GENUS GAURA (ONAGRACEAE)

FIGURES 1-15. Mature fruits of species of Gaura. Fig. 1. G. mutabilis.— Fig. 2.
G. macrocarpa.— Fig. 3. G. boquillemis.— Fig. 4. G. parvifiora.— Fig. 5. G. coccinea.—
Fig. 6. G. villosa subsp. villosa.— Fig. 7. G. villosa subsp. parksii— Fig. 8. G. calcicola.
—Fig. 9. G. filipes.— Fig. 10. G. mckelveyac.— Fig. 11. G. sf nuata— Fig. 12. G. drum-
mnndii (G. odorata of many authors).— Fig. 13. G. lindheimeri.— Fig. 14. G. angusti-
folia, 4-sided fruit.— Fig. 15. G. angmtifolia, 3-sided fmit.

MEMOIRS OF THE TORREY BOTANICAL CLUB

5

FIGURES 16—30. Mature fruits of species of Gaura, and of Stenosiphon. Fig. 16.
G. neomexicana subsp. neomexicana.- Fig. 17. G. neomexicana subsp. coloradensis.—
Fig. 18. G. longiflora (Marion Co., Illinois).- Fig. 19. G. longiffora (Harris Co., Texas).
-Fig. 20. G. biennis.- Fig. 21. G. demareei.— Fig. 22. G. suffulta subsp. suffulta — Fig.
23. G. suffulta subsp. nealleyi (G. suffulta var. terrellensis ) .—Fig. 24. G. suffulta
nealletji.— Fig. 25. G. hexandra subsp. gracilis.— Fig. 26. G. hexandra subsp. hexandra. —
Fig. 27. G. brachycarpa— Fig. 28. G. triangulata, 3-sided fruit.-Fig. 29. G. triangulata,
4-sided fruit.-Fig. 30. Stenosiphon linifolius.

6

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

filament. Otherwise, they resemble G. mutabilis vegetatively and in fruit
morphology and, like it, have nearly actinomorphic flowers. It is tempting
to speculate that, with the transition from pollination by hawkmoths to
pollination by noctuids and other medium-sized moths, predominant
throughout the entire genus except for G. mutabilis, the floral tube was
shortened and then the scales evolved because of their adaptive role in
closing the mouth of the shortened floral tube against possible “illegitimate”
visitors such as bees or flies. The change to white petals seems to have
occurred at this time also, but its adaptive significance is not clear.

One of the most interesting points of similarity among these three
species, primitive on other grounds, is that they have flowers that are
actinomorphic or essentially so (figs. 31-33). With one exception, all other
species of the genus have zygomorphic flowers (e.g., figs. 35-36, 58-63),
a floral trend that represents an obvious specialization. The exceptional
species is the autogamous annual Gaura parviflora, the only member of
section Schizocarya. The very small flowers of this species (fig. 34) do not
have well developed scales at the base of the anthers, but, as pointed out by
Rothrock (Proc. Amer. Acad. Arts 6: 348. 1864), they do have papillae in
the position of the scales in other species. This, taken together with its
very generalized fruits (fig. 4) suggests strongly that Gaura parviflora
was derived from an ancestor comparable to G. macrocarpa. It grows
in moist places, especially along streams in plains country, and is a rank
weed that produces thousands of fruits under favorable circumstances,
when the plants may be up to 3 m tall, with juicy stems 3 cm or more
in diameter.

Specialization of the flowers and a progressive modification of the fruit
toward a more or less stipe-like base residted in the evolution of Gaura
coccinea (section Campogaura ), widespread and variable throughout the
plains from southern Canada to south-central Mexico. Morphologically,
G. coccinea is similar to G. boquillensis; but unlike all species mentioned
previously, it is a polyploid complex, comprising populations with n = 7,
n — 14, n = 21, and n = 28. The extensive recombination associated with
polyploidy may have played a role in the success of this species in achieving
the widest range of any member of the genus.

Section Stipogaura consists of five closely related species in which the
base of the fruit is attenuated into a stripe, and in which the fruit is some¬
times also winged at the angles (figs. 6-11). Four species are clumped
diploid perennials, and of these G. villosa is a large, densely pubescent
perennial of the sand dunes of western Texas and adjacent states. One of
the others, G. filipes, is characteristic of drv, sandy habitats in the south¬
eastern United States, thus occurring beyond the range of any member

MEMOIRS OF THE TORREY BOTANICAL CLUB

7

of the genus mentioned up to this point. The fifth species of section
Stipogaura, G. sinuata, is an aggressively rhizomatous tetraploid (n=rl4)
that often becomes a weed in relatively rich, light soils, as in central Texas.

The single species of section Xenogaura, G. drummondii (G. odorata
of many authors) is uniformly tetraploid and also an aggressive rhizomatous
weed. 1 o a certain extent, its fruits ( fig. 12 ) combine the characteristics
of those of G. coccinea with those of section Stipogaura, and we visualize
an allotetraploid origin for it between these two groups. The diploids of
section Stipogaura are less tightly clumped than are the more generalized
species of the genus, and some (e.g., G. mckelveyae, G. villosa ) have a
tendency to become rhizomatous in light soils.

The remaining two sections are relatively large and clearly specialized
within the genus. Section Gaura has six species. Of these, only one—
G. lindheimeri of the black-soil prairies of east Texas and adjacent Louisiana
—is perennial. It may even be secondarily perennial, and one could readily
imagine how its loose clumps could be derived from the persistence and
budding of the sort of taprooted annual or biennial plants that make up the
rest of the section. Of these, the self-incompatible G. longiflora (G. fili-
formis) is widespread in the the central United States. Populations similar
to it seem to have given rise to G. neomexicana, a self-compatible relict
species found in several disjunct areas along the east flank of the Rocky
Mountains from southeastern Wyoming to New Mexico; to G. biennis, a
complex structural heterozygote that occurs in openings in the deciduous
forest of eastern North America; and to G. demareei, a localized species of
southwestern Arkansas. Less closely related is the autogamous G. angusti-

folia, a very common species in sandy areas of the Coastal Plain of the
southeastern United States.

Gaura lindheimeri and G. demareei have departed from the usual mode
of pollination in the genus. Their flowers open near sunrise, instead of
near sunset, and they are pollinated by bees instead of by the medium-
sized moths that visit the flowers of other white-flowered outcrossing species
of the genus. Their flowers are larger than those of their immediate rel¬
atives, a factor that is probably correlated with the increased caloric reward
that is necessary for their pollinators to be active at the lower temperatures
of early morning (Heinrich and Raven, Science 176: 597-602. 1972).

The final section of Gaura, section Pterogaura, consists entirely of an¬
nual species with winged fruits and some tendency (especially in G. suff ulta
subspecies nealleyi) for the formation of a stipe also. Of the four species
of this group, two have relatively large flowers and are outcrossing, and
two are autogamous. The former center in and around Texas, the latter
extend far beyond its borders, one (G. hexandra subspecies hexandra) as

8

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

far as Guatemala and the other (G. triangulata) into Oklahoma. Gaura
triangulata is a complex structural heterozygote, and, with G. biennis (sec¬
tion Gaura), one of two that has originated independently within the genus.

In both of the widespread autogamous taxa in section Pterogaura (G.
hexandra subspecies hexandra, G. triangulata), 3-merous flowers predom¬
inate (figs. 26, 28, 29, 61, 63), and they are likewise common in G. angusti-
folia (section Gaura). In our experience, these plants are virtually never
visited by insects, and thus agree with other populations in Onagraceae
( Camissonia , Ludwigia; cf., Raven, Contr. U.S. Natl. Herb. 37: 376, 377.
1969 ) in which the evolution of flowers with reduced numbers of parts seems
to have followed the elimination of insect visitors and rather strict autogamy.

ACKNOWLEDGMENTS

We are most grateful to Rancho Santa Ana Botanic Garden and to the Plant
Biology Station of the Carnegie Institution of Washington, Stanford, for the use
of their facilities in cultivating selected strains of Gaura during the past 15 years.
This study was initially suggested by Philip A. Munz, whose encouragement and
counsel have been invaluable. Harlan Lewis has kindly reviewed the entire manuscript.
The fruits (figs. 1—30) were drawn by Marilyn Wright, and the maps were mostly
prepared by Pamela Radford, who also provided expert assistance in other aspects
of the project. Others who helped considerably with the plants in cultivation and
with the processing of many thousand herbarium specimens were Michael Huxtable
and Sarah Hornby. William Tai contributed a great deal of the information on
chromosome numbers and interspecific hybridization; although these results are not
reviewed in detail in this paper, they have affected the conclusions we are presenting
here. The work has been supported generously by a series of grants from the
U.S. National Science Foundation.

Many individuals have contributed special materials for this study, comprising
either herbarium specimens, seeds, or fixed buds for eytological study. Others have
given us the benefit of their advice in critical matters of nomenclature or in the

interpretation of the literature. Among those who helped us in these and other

ways, we would like especially to mention H. E. Allies, D. M. Bates, D. E. Breed¬
love, E. T. Browne, Jr., K. L. Chambers, W. J. Cody, T. S. Cooperrider, E. L. Core,

D. S. Correll, J. E. Dandy, D. G. DeLisle, D. Demaree, D. B. Dunn, R. A. Evers,
J. Ewan, J. M. Fogg, Jr., R. K. Godfrey, C. G. Gunn, V. J. Hoff, N. Hotchkiss, A.
Hunziker, M. C. Johnston, S. B. Jones, Jr., W. M. Klein, Jr., R. W. Long, R.
McVaugh, R. H. Mohlenbrock, Jr., T. Mosquin, C. Nackejima, A. M. Powell, C. F.
Reed, V. E. Rudd, J. Rzedowski, S. G. Shetler, L. G. Shinners, P. D. Sorenson,
W. T. Steam, J. W. Thieret, H. F. Towner, D. B. Ward, W. A. Weber, and G. S.
Winterringer.

We are most grateful to the curators of the following herbaria, from which
material has been made available for this study: Sid Ross State College, Alpine,
Texas (SRSC); Iowa State College, Ames (ISC); University of Michigan, Ann Arbor
(MICH); University of Georgia, Athens (GA); University of Texas, Austin (TEX);
Indiana University, Bloomington (IND); University of Colorado, Boulder (COLO);
Jardin Botanique de l’Etat, Bruxelles (BR); Pringle Herbarium of the University of

MEMOIRS OF THE TORREY BOTANICAL CLUB

9

Vermont, Burlington (VI ); University of Cambridge (CGE); Gray Herbarium, Cam¬
bridge (GH); New England Botanical Club, Cambridge ( NEBC); Southern Illinois
University, Carbondale (SIU); University of North Carolina, Chapel Hill (NCU);
Field Museum of Natural History, Chicago (F); University of Cincinnati (CINC);
Pomona College, Claremont (POM); Rancho Santa Ana Botanic Garden, Clare¬
mont (RSA); Texas A & M University, College Station (TAES); Botanical Museum
and Herbarium, Copenhagen (C); Oregon State University, Corvallis (OCS);
Southern Methodist University, Dallas (SMU); Duke University, Durham, North
Carolina (DUKE); Michigan State University, East Lansing (MSC); Royal Botanic
Garden, Edinburgh (E); North Dakota State University, Fargo (NDA); University
of Arkansas, Fayetteville (UARK); Herbarium Universitatis Florentinae, Firenze
(F); Colorado State University, Ft. Collins ( CS ) ; Agricultural Experiment Station,
Gainesville (FLAS); Conservatoire et Jardin Botaniques, Geneve (G); State Uni¬
versity of Iowa, Iowa City (IA); L. H. Bailey Hortorium, Ithaca, New York (BH);
Cornell University, Ithaca (CU); Kent State University, Kent, Ohio (KE); Royal
Botanic Gardens, Kew (K); University of Tennessee, Knoxville (TENN); University
of Southwestern Louisiana, Lafayette (LAF); University of Kansas, Lawrence
(KANU); Komarov Botanical Institute of the Academy of Sciences of the U.S.S.R.,
Leningrad (LE); University of Kentucky, Lexington (KY); University of Nebraska
State Museum, Lincoln (NEB); British Museum (Natural History), London (BM);
University of California, Los Angeles (LA); Linnaean Herbarium, London (LINN);
Texas Technical University, Lubbock (TTC); University of Wisconsin, Madison
(WIS); Kansas State University, Manhattan (KSC); Institute de Biologia, Mexico,
D. F. (MEXU); Institute Politecnico Nacional, Mexico, D. F. (IPN); University
of Minnesota, Minneapolis (MIN); West Virginia University, Morgantown (WVA);
Botanische Staatssammlung, Munchen (M); Tulane University, New Orleans,
Louisiana (NO); New York Botanical Garden, Bronx (NY); University of Oklahoma,
Norman (OKL); University of Notre Dame, South Bend, Indiana (ND); National
Museum of Canada, Ottawa (CAN); Plant Research Institute, Ottawa (DAO);
Museum National d’Histoire Naturelle, Paris (P); University of Pennsylvania,
Philadelphia (PENN); Academy of Natural Sciences, Philadelphia (PH); Car¬
negie Institute, Pittsburgh (CM); Botanical Research Institute, Pretoria (PRE);
Texas Research Foundation, Renner (LL); Missouri Botanical Garden, St. Louis
(MO); California Academy of Sciences, San Francisco (CAS); Stanford University,
Stanford, California (DS); National Herbarium of New South Wales, Sydney
(NSW); Florida State University, Tallahassee (FSU); University of South Florida,
Tampa (USF); Pennsylvania State University, University Park (PAC); University
of Illinois, Urbana (ILL); Illinois State Natural History Survey, Urbana (ILLS);
US. National Museum, Washington, D.C. (US); Naturhistorisches Museum, Wien

(W); Botanischer Garten und Institut fiir Systematische Botanik der Universitat
Zurich (Z).

SYSTEMATIC TREATMENT
GAURA L.

Gaura L., [Diss. Bot. Nova Plant. Gen. (p.p. L. H. Chenon) 47 1751]

Sp. PI. 1: 347. 1753; Gen. PI. ed. 5. 163. 1754.

Gauridium Spach, Hist. Veg. 4: 379. 1835. Lectotype: Gauridium mutabile
( Cav. ) Spach = Gaura mutabilis Spach.

10

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Schizocarya Spach, Nouv. Ann. Mus. Hist. Nat. 4: 325, 381. 1835; Ann.
Sci. Nat. (Paris) II. 4: 170, 283. 1835. Lectotype: Schizocarya micrantha
Spach = Gaura parviflora Dough

Annual, biennial, or perennial herbs, sometimes woody near the base.
Leaves gradually decreasing in size upward, the basal or rosette leaves
largest, lyrate, and gradually narrowed to a winged petiole, the cauline
leaves subsessile, usually with an acuminate apex; pubescence on leaves
thickest or present only along the veins, especially the midrib, and around
the margins. Inflorescence a spicate raceme, sharply delimited, not leafy,
more or less conspicuously pedunculate, the flowers bracteate. Flowers
home laterally, usually strongly zygomorphic, the petals opening to ap¬
proximately a flat plane and all held in the upper half of the flower, the
stamens and style projecting slightly forward and mostly confined to the
lower quarter or half of the flower, or more rarely the flowers actinomorphic.
opening either near sunset or near sunrise and fading within a single day,
usually 4-merous but less commonly 3-merous. Floral tube narrow, mostly
0.75-1.5 mm in diameter at its apex, usually lanate in upper half within,
the lower half swollen and secreting nectar. Petals usually white, flushed
with reddish after pollination, usually sharply clawed. Stamens twice as
many as the sepals, subequal. Each filament usually with a scale ca. 0.3-0. 5
mm long near its base, these scales nearly closing the mouth of the hvpanth-
ium; anthers usually reddish, the sporogenous tissue divided by sterile tissue
into discrete packets. Style with the same sort of pubescence in the same
area as the floral tube, the hairs interlocking. Stigma deeply 4-(3-)lobed,
usually held beyond the anthers in outcrossing species. Fruit an indehiscent,
nutlike capsule with hard, woody walls, basically 4- (3-) locular but the
septa incomplete and fragile, not evident at maturity; fruit with 1 ovule per
locule basically but the whole fruit often with only 1 ovule and occasionally
as many as 8. Seeds ovoid, mostly 2-3 mm long and 1-1.5 mm thick, soft,
smooth, usually flattened on some or all of the sides, top, and bottom by
crowding within the fruit, yellowish to pale brown. Basic chromosome
number, x = 7. Self-incompatibility characteristic of 13 of the 21 species.

Type Species: Gaura biennis L.

The sympatric occurrence of species of Gaura is indicated in table 1,
which incorporates our own observations as well as those of Bryan (Southw.
Naturalist 8: 10-14. 1963). Naturally occurring hybrids have been observed
only between G. lindheimeri and G. longiflora (rather frequently) and be¬
tween G. calcicola and G. coccinea (one plant; Raven and Gregory, Brit-
tonia 24: 71-86. 1972). They also probably occur at some localities where G.
drummondii and G. coccinea grow together.

The distribution maps presented in this paper generally include onlv

TABLE 1. Sympatric occurrence of species of Gaura.1

m

cc

a

u

-a

oj

f-

dddddddddddddd ddddd

G. biennis

•

G. boquillensis

o

•

G,' brachycarpa

0

o

•

G. calcicola

0

o

o

•

G. coccinea

X

X

X

2

•

G. drummondii

0

0

X

X

3

•

G. hexandra
gracilis

0

0

o

0

X

0

•

G. h. hexandra

o

0

0

0

X

X

4

•

G. lindheimeri

o

0

o

0

o

0

0

0

•

G. longiflora

5

0

o

0

X

0

o

0

6

•

G. macrocarpa

0

0

o

0

X

0

0

0

0

o

•

G. mckelveyae

0

0

o

0

o

0

o

0

0

o

0

•

G. mutabilis

o

0

0

0

o

0

o

X

0

o

o

o

•

G. neomexicana
coloradensis

0

0

o

0

X

0

o

0

0

0

0

0

0

•

G. n. neomexicana

0

0

o

0

X

0

o

o

o

o

o

o

o

7

•

G. parviflora

X

0

X

X

X

X

0

o

0

X

o

X

o

X

o

•

G. sinuata

0

o

X

0

X

X

o

0

0

0

0

o

o

o

0

X

•

G. suffulta
nealleyi

0

0

o

o

X

0

0

o

o

0

X

o

0

o

o

o

0

•

G. s. suffulta

o

0

X

X

X

X

o

0

0

o

o

o

o

0

o

X

X

8

•

G. triangulata

o

0

o

o

X

0

0

0

0

o

o

0

0

0

0

o

X

0

X

•

G. villosa parksii

o

0

X

o

o

0

0

0

0

0

0

0

0

0

o

X

o

0

0

0

•

G. v. villosa

0

0

o

0

X

o

o |

o

0

0

0

0

0

0

0

0

0

o

X

0

9

•

1 Gaura a/ngustifolia, G. demareei, and G. filipes, which we have not observed growing with
other species of the genera, are not included in the table. Numbers in the table refer to the
notes. Unless otherwise noted, s signifies sympatric occurrence without hybridization.

2 One hybrid individual found; Raven & Gregory, Brittonia 24; 71-86. 1972.

3 Hybridization probable but difficult to confirm owing to close similarity of parents.

4 Autogamous and occurring in mixed colonies.

6 Sympatric occurrence and hybridization almost certain but difficult to confirm owin°- to
very close similarity of parents.

® Hybridization frequent but species remain largely distinct.

7 Allopatric subspecies.

8 Broadly intergrading subspecies.

9 Allopatric subspecies.

G. triangulata
G. villosa parksii
G. v. villosa

12

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

one record per county. Records from particular counties not included
among the lists of cited specimens are available from Raven on request.

The floral characters of Gaura have not been utilized to any great ex¬
tent in previous systems of classification, largely because the flowers wither
and fade approximately 12-16 hours after they first open. The photographs
of each taxon in this paper were taken of plants grown in the experimental
garden at Stanford University within a few hours after the flowers first
opened. We hope that these photographs and the drawings of mature
fruits (figs. 1-30) will prove useful in conjunction with the key.

KEY TO THE SPECIES

Floral tube 26-42 mm long; petals yellow . 1. G. mutabilis

Floral tube 1.5-15 mm long; petals white.

Fruits with a slender stipe 0.5—10 mm long.

Plants suffrutescent, conspicuously and densely soft-villous with long

hairs . 6. G. villosa

Plants herbaceous, not conspicuously and densely soft-villous with long
hairs.

Leaves deeply sinuate; plants sparsely villous and strigulose; south

Texas and adjacent Mexico . 0. G. mckelveyae

Leaves slightly to markedly sinuate; pubescence variable.

Clumped perennials; apex of the inflorescence in bud slender, the
buds small and well-spaced.

Leafy portion subglabrous to sparsely strigulose, sometimes with
admixture of longer hairs; stamens held in lower half of flower;

west Texas and south into Mexico . 7. G. calcicola

Leafy portion either strigulose or villous; stamens distributed
around the perimeter of the flower; east Louisiana to Florida and

South Carolina . 8. G. filipes

Annual or rhizomatous perennial; apex of inflorescence in bud stout,
the buds large and crowded.

Coarse annual; inflorescence glandular-pubescent; stipe 0.2-2.2

mm long . 18a. G. suffulta subsp. nealleyi

Mat-forming, rhizomatous perennial; inflorescence subglabrous or

strigulose; stipe 3—8 mm long . 10- G. sinuata

Fruits subsessile or with a thick, cylindrical stipe.

Sepals 2-3.5 mm long; petals 1.5-3 mm long . 4. G. parviflora

Sepals (2.5-) 4.5-17 mm long; petals 3.5-15 mm long.

Flowers nearly actinomorphic; fruits fusiform, often curved, 8—17
mm long, 1-2.5 mm thick; Trans-Pecos Texas to central Chi¬
huahua, Coahuila, and western Nuevo Leon.

Floral tube 9-13 mm long; flowers actinomorphic. 2. G. macrocarpa

Floral tube 5-8.5 mm long; flowers somewhat zygomorphic

3. G. BOQUIELENSIS

MEMOIRS OF THE TORREY BOTANICAL CLUB

13

Flowers zygomorphic, usually strongly so; fruits variable, 4-13
mm long, often thicker; widespread.

F ruits more or less abruptly constricted to a thick cylindrical
stipe nearly half the length of the body; perennial species.

Plants from a branching woody caudex; fruits not conspicuously
bulging above, 4— 8.5(-9) mm long, (1-)1.5— 3 mm thick;

bracts 2—5 mm long, 0.4-1. 1 mm wide . 5. G. coccinea

Plants aggressively rhizomatous; fruits conspicuously bulging
above, 7—13 mm long, 3—5 mm thick; bracts 2—8 mm long,

0.8-2 mm wide . 11. G. drummondii

Fruits not abruptly constricted to a thick cylindrical stipe; mostly
annual or biennial species from a taproot.

Flowers 3-merous; fruits 3-angled.

Plant usually simple below, 6—18 dm tall; fruit narrowly
ellipsoid or ellipsoid; coastal plain from Florida to
North Carolina . 13. Q. angustifolia

Plant usually branched at the base, 1. 5-6.5 (-10) dm
tall; fruits ellipsoid or ovoid.

Sepals very finely strigulose or subglabrous; pollen ca.

50 percent aborted; north-central Texas and Oklahoma

. 21. G. TRIANGULATA

Sepals strigulose or glandular pubescent; pollen mostly
fertile.

Sepals 3—10 mm long; inflorescence usually gland¬
ular-pubescent; Mexico and Guatemala

. 19b G. HEXANDRA Subsp. HEXANDRA

Sepals 10-15 mm long; inflorescence strigulose;

Texas and adjacent states . 20. G. brachycarpa

Flowers 4-merous; fruits 4-angled.

Sepals with long erect hairs; flowers opening near sun¬
rise; petals 10.5-15 mm long; perennial; southeast

Texas and Louisiana . 12. G. lindheimeri

Sepals with appressed pubescence, or flowers opening
near sunset and much smaller; annual or biennial.
Biennial from a stout taproot; meadows of Colorado,

Wyoming, and New Mexico . 14. G. neomexicana

Annual, occasionally persisting through mild winters.

Fruits broadly winged on the angles and deeply
furrowed between the angles; plants rarely to
1 m tall, usually well branched from the base.

Sepals 6-12 mm long, glandular-pubescent, sub-
glabrous, or strigulose; Mexico to Arizona
and New Mexico. .19. G. hexandra subsp. gracilis
Sepals 10-21 mm long, glabrous or strigulose;

Texas and adjacent states, but not known
from Mexico or New Mexico.

Sepals glabrous; bracts 2.5-6.5 mm long
. 18. G. SUFFULTA subsp. SUFFULTA

14 A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Sepals strigulose; bracts 2-4 mm long

. 20. G. BRACHYCARPA

Fruits angled but not winged; plants often 1-4 m tall,
usually unbranched from the base.

Sepals 2.5-8 mm long; flowers often 3-merous and
4-merous on the same plant; coastal plain
from Florida to North Carolina

. 13. G. ANGUSTIFOLIA

Sepals 8-20 mm long; flowers 4-merous; not on
coastal plain.

Flowers opening near sunrise; sepals 13—20 mm
long; pubescence appressed; southwestern
Arkansas . 17. G. demareei

Flowers opening near sunset ( rarely near sun¬
rise in Louisiana and Alabama ) ; sepals 8—

18 mm long; pubescence appressed or
spreading.

Stems densely villous; inflorescence glandular-
pubescent; pollen about 50 percent fertile
(usually 35-65 percent) . 16. G. biennis

Stems strigulose or villous; inflorescence ap¬
pressed- or glandular-pubescent; pollen
usually more than 90 percent fertile

. 15. G. LONGIFLORA

SECTION I. GAURIDIUM

Gaura sect. Gauridium (Spach) Endl., Gen. 1195. 1840.

Gauridium Spach, Hist. Veg. 4: 379. 1835.

Clumped perennial herb, the steins villous, hirtellous, and glandular-
pubescent. Inflorescences erect. Flowers nearly actinomorphic, the petals
spreading at right angles, the stamens and style projecting outward from
the flower; flowers opening near sunset and withering rapidly tit next
morning. Floral tube sparsely pubescent within. Petals yellow, flushed
with red after anthesis, not clawed. Filaments lacking scales. Fruit not
reflexing, narrowly ovoid or fusiform, rounded at the base and gradually
tapering to a point near the apex, mostly terete but occasionally somewhat
4-angled, the ribs obscure but the 4 main ones broader. Self-compatible.
Gametic chromosome number, n = 7.

Type Species: Gaura mutabilis Cav. [Gauridium mutabile (Cav.)

Distribution: Middle elevations in the Sierra Madre Occidental and
Trans-Mexican Volcanic Belt of Mexico, south to the vicinity of Oaxaca;

usually in pine or oak forest. ,

Gaura mutabilis, the single species of this section, has an array of what

have been interpreted as generalized features within the genus: large,
nearly actinomorphic, yellow flowers that lack scales at the base of the

MEMOIRS OF THE TORREY BOTANICAL CLUB

15

35 36

FIGURES 31-36. Flowers of species of Ganra. Fig. 31. G. mutabilis. -Fig. 32.
G. macrocarpa.- Fig. 33. G. boquiUensis. -Fig. 34. G. pam/lora.-Fig. 35. G. coccinea.
-Fig. 36. G. drummondii (G. odorata of many authors).

16

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

filaments and have the longest floral tube in Gaura; elongate fruits that
are superficially similar to the capsules from which they have been derived;
and a Mexican distribution. Viewed in an ecological context, however, the
interpretation of these characteristics becomes more difficult, for G. muta¬
bilis is the only hawkmoth-pollinated species of the genus, and all or most
of the floral features could be associated with this. Furthermore, the species
is self-compatible, an obviously secondary characteristic. Despite these
cautionary notes, there is enough overall similarity in fruit morphology and
habit to G. macrocarpa to suggest that G. mutabilis is at the very least an
ancient offshoot of a primitive group of species of Gaura, and to leave it in
its customary position at the start of the genus.

1. Gaura mutabilis Cav., Ic. 3: 30, pi. 258. 1795. Munz, Bull. Torrey Bot.
Club 65: 107. 1938; N. Amer. FI. II. 5: 183. 1965. Figs. 1, 31.

Oenothera anomala Curtis, Bot. Mag. 11: pi. 388. 1809. Based on Gaura
mutabilis Cav.

Gaura oenotheriflora Zucc., in Roem., Coll. 140. 1809. Lectotype: culti¬
vated, herb. Zuccagni (FI).

Gaura mollis H.B.K., Nov. Gen. et Sp. 6: 93, pi. 529. 1823. Type:
Mexico, 1803-1804, Alexander von Humboldt (P).

Gauridium mutabile (Cav.) Spach, Hist. Veg. 4: 380. 1835.

Gauridium molle (H.B.K.) Spach, Hist. Veg. 4: 380. 1835.

Gauridium kunthii Spach, Nouv. Ann. Mus. Hist. Nat. 4: 375. 1835.
Based on Gaura mollis H.B.K.

Gaura grandiflora Rose, Contr. U. S. Natl. Herb. 12: 293. 1909. Type:
Papasquiaro, Durango, Mexico, 7 August 1898, E. W. Nelson 4671
(US 332725. Isotype, K).

Gaura gentri/i Standley, Field Mus. Nat. Hist. Bot. Ser. 17: 203. 1937.
Type: On rocky outcroppings, Guasaremos, Rio Mayo, Chihuahua,
Mexico, 26 September 1935, II. S. Gentry 1859 (F. Isotypes, GH, K,
MO, PH, US).

Gaura mutabilis Cav. f. glabra Munz, Bull. Torrey Bot. Club 65: 108.
1938. Based on Gaura grandiflora Rose. Munz, N. Amer. Fl. II. 5:
184. 1965.

Clumped perennial herb with a heavy, woody, twisted rootstock; stems
erect, branching near the base and again just below the inflorescence, 2.5-
8.5 dm tall. Pubescence of very long hairs 3.5-4.5 mm long near the base,
scattered upward; shorter erect hairs on the stems, leaves (especially along
midrib and margins), and sometimes in the inflorescence; and erect glan¬
dular hairs found all over the plant. Leaves narrowly lanceolate to lanceolate,
sometimes narrowly elliptic, 1 —6 ( —8 ) cm long, 0.2-1.8 cm wide, sinuate-

MEMOIRS OF THE TORREY BOTANICAL CLUB

17

dentate to subentire. Inflorescence unbranched, 5-25 cm long, glandular-
pubescent with varying amounts of short erect eglandular hairs; bracts
narrowly lanceolate to lanceolate, 2-9 mm long, 1-1.75 mm wide. Floral
tube 26-42 mm long. Sepals 11-24 mm long, 1-2 mm wide. Petals 12-25
mm long, 6-15 mm wide. Filaments 5-18 mm long; anthers 3-5 mm long.
Style 38-67 mm long. Fruit 7-13 mm long, 2-6 mm thick. Seeds almost
always 4, 2.5-4.5 mm long, 1.5-2 mm thick, light reddish brown. Self¬
compatible, the stigma surrounded by the anthers at anthesis or held well
above them. Gametic chromosome number, n = 7.

Type: Described from plants cultivated in the Botanical Garden at
Madrid, the seeds of Mexican origin.

Distribution ( fig. 37 ) : Rocky slopes and outcrops in pine or oak
forest, rarely in matorral, 2000—2800 m elevation, from the vicinity of
Chihuahua and the upper Rio Mayo in Sonora scattered southward in the

18 A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Sierra Madre Occidental to the Trans-Mexican Volcanic Belt as far as
southern Hidalgo and Puebla; collected once near Oaxaca. Flowers July
to September.

Representative Specimens Examined:

Mexico, chihuahua. Colonia Juarez, Jones in 1903 (DS, POM); hills near
Chihuahua, Pringle 1125 (BM, BR, COCO, F, G, CH, LE, MSC, RSA, US); high
plains between Cusihuiriachic and Guerrero, Pringle 1244 (COCO, F, GH, MEXU,
MIN, NO, NY, PH, RSA, US); Culebra Mts., Le Sueur in 1936 (GH, MO, SMU);
e foothills of Sierra Madre 13 mi w of General Trias, 5900 ft, Ripley & Barneby
13887 ( DS, NY), durango. 3 mi ne of Otinapa, Maysilles 7305 (MICH, RSA);
ca. 60 air miles sw of Durango, 2500-2650 m, Maysilles 7792 (MICH); ca. 50 km
w of Durango near the highway to Mazatlan, 2550 m, McVaugh 21677 (MICH);
near El Salto, Nelson 4570 (POM, US); Papasquiaro, Nelson 4671 (GH); Durango
and vicinity, Palmer 270 ( BM, F, G, GH, MO, NY, US); Palos Colorados on railroad
w of Durango, 2400 m, Pennell 18233 (PH, US); Sierra de Registro 13 mi nw of
Nombre de Dios, 6300 ft, Ripley &■ Barneby 14165 (DS); 9 m w of Durango, Water¬
fall 12560 (GH, MICH, OKLA, US): 28 mi sw of Durango, Waterfall is Wallis
13563 (IA, OKLA, SMU); 5 mi sw of Durango, Waterfall 15416 (OKL, OKLA,
SMU). Guanajuato. Guanajuato, Dtiges 303 (GH); Montapues, Guanajuato and
vicinity, Duges 303 A (GH); Cerro de la Batea, Guillemin-T ardyre (K, LE, P); near
summit of Montana de Cubilete, ne of Silao, 8000 ft, Straw 6- Forman 1450 (MICH,
RSA). hidalgo. Hills near Marquez, 8000 ft, Pringle 7635 (POM, US); Cerro
Ventoso, between Pachuca and Real del Monte, 2600 m, Rzeclowski 20622 ( DS, IPN,
ISC, MICH, TEX); Cerro Santa Ana, 10 mi ne of Apam, West B-l (DS). Mexico.
Santa Fe, Valley of Mexico, Bourgeati 296 (C, C»H, NY, RSA, L^S); 12 mi s of
Tlapam, D.F., W. E. dr M. S. Manning 531051 (GH); lomas de Mixcoae, D.F.,
Lyonnet 1471 (US); Villa Obregon, w of San Angel, 8000 ft, D.F., Munz 15056
(GH, MO); Pedrigal, Valley of Mexico, Pringle 6661 (BM, BR, CM, F, G, C,H, IPN,
ISC, LE, M, MEXU, MIN, MO, ND, NMC, NY, PH, US), morelos. Hwy. to
Cuernevaca near Morelos Monument, Straw 6- Gregory 1076 (DAO, GY, MEXU,
MICH, RSA). oaxaca. Near Oaxaca, Anclricux 384 (CH). puehla. Santa Barbara,
Nicolas 259 (BY, F, GH, MO, NY, P, US); San Martin, Nelson in 1893 (US); vicinity
of San Luis Tultitlanapa, near Oaxaca, Purpus 3388 (BM, F, CH, MO, NY, US);
Puenta de Emperador near La Venta, 8300 ft., Sharp 44393 (RSA); Cerro Amalucan,
8 km al e de Puebla, Rzedowski 24135 (DS, IPN). sonora. Alamos Dist., Arroyo,
Algodones valley, Gentry 468 ( DS, MICH); Arroyo del Auga Blanca, Distrito Alamos,
Gentry 514 (MICH); Sierra de las Papas, Gentry 622 ( DS, MICH).

Gaura mutabilis is sometimes called “noche buena,” this name con¬
trasting the late opening of its flowers (in full dark after sunset) with those
of G. coccinea “linda tarde.” It has been in cultivation in botanical gardens
ever since its first introduction in the 1790’s. In Mexico, it tends to be
colonial, with individual populations widely scattered. There is a small
amount of interpopulational variability, especially in pubescence, style
length, and fruit shape, but no consistent patterns that appear to require
formal taxonomic recognition.

MEMOIRS OF THE TORREY BOTANICAL CLUB

19

SECTION II. XEROGAURA

Herbae perennes confertae, caulibus strigulosis, scabris, glandulosis,
vel villosis. Inflorescentiae subnutantes. Flores subactinomorphi, stylo
subdeclinato, vespertini. Frueti fusiformes, saepe subcurvati, tereti, costibus
4 majoribus, 4 angustioribus.

Clumped perennial herbs, the stems strigulose, scabrous, glandular, or
villous, or with various mixtures of these pubescence types. Inflorescences
slightly nodding. Flowers nearly actinomorphic, the style slightly declinate;
flowers opening near sunset and withering the next morning. Fruit reflexing
or not, fusiform, often slightly curved, terete, with 4 main ribs and 4 narrower
ones. Self-incompatible. Gametic chromosome number, n = 7.

Type Species: Gaura macrocarpa Rothr.

Distribution: Washes and sandy flats from the Davis Mountains of
west Texas to central Chihuahua, Coahuila, and western Nuevo Leon.

The white, nearly actinomorphic flowers and elongate fruits of the two
species of this section seem clearly to be generalized features within the
genus.

2. Gaura macrocarpa Rothr., Proc. Amer. Acad. Arts 6: 353. 1864-1865.

Munz, Bull. Torrey Bot. Club 65: 115. 1938; N. Amer. FI. II. 5: 186.

1965. Figs. 2, 32.

Clumped perennial herb with a narrow woody rootstock usually branch¬
ing several centimeters below the ground level; stems with several branches
from the base and occasional branches above, arching slightly outward from
the base but then strictly erect, 2-10 dm tall. Pubescence of four kinds
of hairs: 1. long (1-1.5 mm) erect hairs near the base of the plant; 2.
short, appressed hairs everywhere but heaviest on the ovaries and usually
least dense on the branches of the inflorescence; 3. short, scabrous hairs,
thick at the base, common around edges of leaves but sometimes dense on
stems also; and 4. short, erect, glandular hairs all over the plant, usually
heaviest on the branches of the inflorescence. Leaves very narrowly elliptic
or usually narrowly lanceolate, 0.5-8 cm long, 0. 1-0.8 cm wide, subentire to
sinuate-denticulate, the apex mucronate. Inflorescence 10-25 cm long,
often with a few branches near the base; bracts narrowly ovate, 1.5-3 mm
long, 0.85-1.25 mm wide. Many flowers in a single inflorescence open at
one time. Floral tube 9-13 mm long. Sepals 7-9 mm long, 1-2 mm wide.
Petals 7-8 mm long, 3-6 mm wide. Stamens projecting forward from the
flower. Filaments 4-4 mm long; anthers 4-5 mm long. Style 18-22 mm
long. Fruit not reflexing, (9-) 13-17 mm long, 2-3 mm thick, with 8 ribs
of about equal prominence, although 4 are slightly narrower; fruits sessile.

20

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

MEMOIRS OF THE TORREY BOTANICAL CLUB

21

Seeds (l-)3-4, 2-3.5 mm long, 1-2 mm thick, yellowish or light brown.
Self-incompatible. Gametic chromosome number, n = 7.

Lectotype: Limpia Canyon (“Valley of the Limpio”), Jeff Davis Co.,
Texas, 21-26 August 1849, C. Wright 1079 (US 59209. Isotypes, BM,
GH, LE, M, P, PH); Munz, Bull. Torrey Bot. Club 65: 115. 1938.

Distribution (Fig. 38): Sandy flats and washes, very local, vicinity of
the Davis Mountains in eastern Jeff Davis Co., northeastern Presidio Co.,
and northern Brewster Co., Texas; near Gallego and Chihuahua, Chihuahua.
Flowers, May to July.

Representative Specimens Examined:

Mexico, chihuahua. 7 mi n of Gallego, Breedlove 15729 (DS); Chihuahua,
Palmer 195 (NY, US).

United States, texas. Brewster Co.: Haley Ranch, Parks ir Cory 9278 (POM,
TAES); Flats to Sunny Glen, Sperry T604 (GH, TAES, US); 2 mi e of Alpine,
Warnock 8468 (LL, SMU); Bois d’Arc Spring, Warnock 127 (PENN). Jeff Davis
Co.: Near Fort Davis, Palmer 34558 (GH, MO, NY, PH, US); 18.3 mi s of Toyahvale,
0.7 mi n of Wild Rose Pass, Gregory 165 (RSA); 5.8 mi s of Toyahvale, Gregory 252
( RSA). Presidio Co.: 5 mi w of Alpine, Gregory 202 ( DS, RSA); 2.7 mi n of
Marfa, T. 6- L. Mosquin 5651 (DAO).

Gaura macrocarpa is known only from three disjunct regions and it
seems likely, in view of its generalized features, that it is a relatively old
species that has survived locally in a few favorable situations. It has been
collected only twice in Mexico.

3. Gaura boquillensis Raven & Gregory, sp. nov. Figs. 3, 33.

Herba perennis sublignosa copiose multiramosa, 2.5-10 dm alta, stri-
gulosa saepe etiam glandulosa scaberulaque. Folia 0.5-6.5(-13) cm longa,
0.1-1 cm lata, basalia anguste oblanceolata raro spatulata, superiora angus-
tissime elliptica vel anguste lanceolata, saepe linearia, subsessilia, sinuato-
dentata vel subintegria, plerumque sinuata. Inflorescentia 8-37 cm longa,
stricta, bracteis lanceolatis vel linearibus 1.5-3.5 mm longis, 0.25-0.5 mm
latis. Tubus floris 3-8.5 mm longus, intus costibus hispidulus. Sepala 3-9
mm longa, 1-2.5 mm lata. Petala 4-10 mm longa, 2-4.5 mm lata. Filamenta
2-4.5 mm longa; antherae 2-4 mm longae. Stylus 6.5-15 mm longus, basin
versus hispidulus. Fructus fusiformis, subarcuatus, 5.5-13 mm longus, 1-2.5
crassus, interdum parte tertia inferiora subattenuata stipaformaque; semina
( 1- ) 2-4, 1.5-2.5 mm longa, 0.75-1.25 mm crassa, obovoidea, straminea vel
pallide brunnea. Numerus chromosomaticus gameticus, n = 7.

Clumped perennial herb with a narrow, woody rootstock usually branch¬
ing several centimeters below the ground level; stems well branched at the
base and above, often somewhat woody, the plant very full, 2.5-10 dm tall.

22 A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Pubescence very variable, of three kinds of hairs: 1. short, erect glandular
hairs, absent or fairly dense on stems and leaves; 2. strigulose pubescence,
common everywhere except the branches of the inflorescence; 3. short,
scabrous hairs, thick at base, usually sparse on stems and leaves, sometimes
dense, absent in inflorescence. Leaves usually narrowly oblanceolate near
the base, becoming very narrowly elliptic or narrowly lanceolate upward,
or often linear, 0.5-6.5(-13) cm long, 0.1-1 cm wide, sinuate-dentate to sub¬
entire. Inflorescence unbranched, 8-37 cm long; bracts lanceolate to linear,
1.5-3. 5 mm long, 0.25-0.5 mm wide. Floral tube 3-8.5 mm long. Sepals
3-9 mm long, 1-2.5 mm wide. Petals 4-10 mm long, 2-4.5 mm wide.
Filaments 2-4.5 mm long; anthers 2-4 mm long. Style 6.5-15 mm long.
Fruit often reflexing at maturity, 5.5-13 mm long, 1-2.5 mm thick, some¬
times slightly narrowed in the lower third and then approaching a stipitate
condition. Seeds (l-)2-4, 1.5-2.5 mm long, 0.75-1.25 mm thick, yellowish
or light brown. Self-incompatible. Gametic chromosome number, n - 7.

Type: Mouth of Boquillas Canyon, Big Bend National Park, Brewster
Co., Texas, 20 April 1960, D. P. Gregory 232 (RSA 145239. Isotype, NY).

Distribution (fig. 38): Washes and sandy canyonsides in the drv moun¬
tains near the Rio Grande in southern Brewster Co., Texas, southward to
central Chihuahua, Coahuila, and western Nuevo Leon. Flowers, May and
June.

Representative Specimens Examined:

Mexico, chihuahua. Plains near Chihuahua, Pringle 1497 (GH, MEXU); 5 mi
s of Gallego, Shreve 7930 (US), coahuila. 1 mi e of Muzquiz, Gould 11230 (DS);
3 mi above Socorro, Johnston 8840 (GII); Monclova, Marsh 1679 ( F, GH, OKLA,
TEX), 1704 (GH); Muzquiz, Santa Anna Canyon, Marsh 483 (OKLA, SMU, TEX)
5.6 mi e of Saltillo, Towner 55 (DS); 12 mi e of Saltillo, Canedo et al. 9072 ( DS,
DUKE, OKLA, SMU, TEX, TTC); El Puerto de San Lazaro, Wynal & Mueller 145
(A, FSU, ILL, MICH, MO, MSC, US); w slope of Sierra de San Vicente, Schroeder
132 (GH). nuevo leon. Sierra Madre Oriental, Monterrey, Mueller 218 (A, POM);
79.9 mi s of Tamaulipas border on road from Nuevo Leon to Monterrey, Towner 37
(DS).

United States, texas. Brewster Co.: Boquillas Canyon, Harrison in 1919 (WISC),
Warnock in 1937 (TAES), Cory 2139 (POM); Big Bend National Park, Webster
4481 (DUKE, G, OKLA, SMU, W); mouth of Santa Elena Canyon, A. 6- R. A.
Nelson 5052 (GH), Innes et al. 503 (GH); banks of the Rio Grande, in Grand
Canyon, Palmer 34224 (MIN, NY); Pinnacle Mts., Chisos Mts. area, Warnock 1055
(US).

This distinctive new species has been confused with G. macrocarpa,
but differs markedly in its smaller, more irregular flowers and shorter,
somewhat stipitate fruits. It grows in the same general area as G. macro-
carpa, and the two species may ultimately be found growing together near
Gallego or Chihuahua.

MEMOIRS OF THE TORREY BOTANICAL CLUB

23

SECTION III. SCHIZOCARYA
Gaura sect. Schizocarya (Spach) Raven & Gregory, comb. nov.

Schizocarya Spach, Nouv. Ann. Mus. Hist. Nat. 4: 325, 381. 1835; Ann.

Sei. Nat. (Paris) II. 4: 170, 283. 1835.

Rank annual herb, the stems glandular-pubescent with an admixture
of villous pubescence. Inflorescences slender, nodding. Flowers actino-
morphic, the petals open only to about 90°, the stamens and stigma erect;
flowers opening near sunset and withering the next morning. Filaments
with minute scales only. Fruit reflexing at maturity, fusiform, tapering
more or less abruptly toward the base, terete, weakly 4-ridged and furrowed
in the upper third, 8-ribbed below. Autogamous. Gametic chromosome
number, n = 7.

Lectotype Species: Gaura parviflora Dough ( Schizocarya micrantha
Spach ) .

Distribution: Weedy places and along streams throughout the central
United States, south to central Mexico.

Despite Rothrock’s accurate observation in 1864 (Proc. Amer. Acad.
Arts 6: 348), the papillae to which the scales at the base of the filaments
in Gaura parviflora have been reduced have generally been overlooked
since. Their presence makes very likely the derivation of this rank annual
weed from plants similar to those currently assigned to sect. Xerogaura
highly probable, in view of the very similar fruits. Presumably the scales,
which serve in most species of Gaura to exclude all but long-tongued in¬
sects from the nectar in the floral tube, have been greatly reduced in G.
parviflora owing to its strict autogamy, very small flowers, and absence
of nectar.

4. Gaura parviflora Dougl., in Hook., FI. Bor.-Amer. 1:208. 1833. Hook.,
Bot. Mag. 63: t. 3506. 1836. Munz, Bull. Torrey Bot. Club 65: 108.
1938; N. Amer. FI. II. 5: 184. 1965. Figs. 4, 34.

Schizocarya micrantha Spach, Ann. Sci. Nat. (Paris) II. 4: 283. 1835.
Lectotype: Austin, Travis Co., Texas, 1833-1834, Thomas Drummond
46 (G. Isolectotypes, GH, NY); Munz, Bull. Torrey Bot. Club 65- 110
1938.

Gaura micrantha (Spach) D. Dietr., Enum. PI. 2: 1297. 1840.

Gaura hirsuta Scheele, Linnaea 21: 580. 1848. Type: Prairies between
Bastrop and Austin, Bastrop or Travis Co., Texas, 1846, Ferdinand
Roemer (not seen).

Gaura australis Griseb., Abh. Konigl. Ges. Wiss. Gottingen. 24: 132. 1879.

24

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Lectotype: Entre Mina Argentina y Yerba Buena, en las cercanias del
Cerro de Orcosu, Dpto. Minas, Cordoba, Argentina, 17 Feb. 1876,
Hieronymus 399 (GOET. Isolectotype, CORD. Neither seen).

Gaura parviflora Dougl. var. lachnocarpa Weath., Rhodora 27: 14. 1925.
Type: Roadside, Austin, Travis Co., Texas, 18 April 1918, M. S. Young
95 (GH. Isotype, TEX).

Gaura parviflora Dougl. var. tyjnca Munz, Bull. Torrey Bot. Club 65:
109. 1938.

Gaura parviflora Dougl. var. typica Munz f. glabra Munz, Bull. Torrey
Bot. Club 65: 110. 1938. Type: Deer Run, 4700 ft, between Kannah
and Delta, Meso Co., Colorado, 11 June 1901, C. F. Baker 94 (POM
32745. Isotypes, GH, NY, US).

Rank annual herb with a heavy taproot up to 3 cm or more in diameter;
stems erect, usually unbranched or branching just below the inflorescence,
thick and juicy, tapering gradually to the inflorescences, (2-)3-20(-30) dm
tall, the basal leaves usually deciduous by the time of flowering and the
lower stems then naked. Pubescence up to the base of the inflorescences
dense, glandular, with fewer long, spreading hairs, both of which also occur
along the main veins and margins of the leaves; surface of leaves strigulose.
Leaves narrowly elliptic to narrowly ovate, mostly commonly acuminate,
2-12.5 cm long, 0.5-4 cm wide, slightly sinuate-dentate. Rosette leaves often
broadly oblanceolate, up to 15 cm long and 3 cm wide, the blade gradually
narrowed into a winged petiole. Inflorescence a series of subterminal dense,
spicate racemes 5-45 cm long, glabrous (except for long hairs on edges of
floral bracts) to densely covered with a mixture of glandular and eglandular,
long hairs, then also with short erect hairs on the ovaries and loosely ap-
pressed pubescence on the calyx; bracts narrowly lanceolate to linear, 1.5-
5.5 mm long, 0.25-0.5 mm wide. Floral tube 1.5-5 mm long. Sepals 2-3.5
mm long, 0.5-1 mm wide. Filaments 1.5-3 mm long; anthers 0.5-1 mm
long, shedding pollen directly on the stigma at or before anthesis. Style 3-9
mm long; lobes of the stigma short. Fruit 5-11 mm long, 1.5-3 mm thick.
Seeds 3-4, 2-3 mm long, 1-1.5 mm thick, reddish brown. Autogamous.
Gametic chromosome number, n = 7.

Type: Sandy banks of the Walla Walla River, Walla Walla Co., Wash¬
ington, 10-16 June 1826, David Douglas (K. Isotype, BM).

Distribution (fig. 39): Weed of cultivated fields, pastures, waste places,
and along streams, throughout the interior of North America; in the United
States from Indiana, Iowa, southern South Dakota, Montana (very rare),
and southeastern Washington to the lower Mississippi River; in Mexico,
Nuevo Leon, Zacatecas, Durango, Sinaloa, and Baja California. Locally

MEMOIRS OF THE TORREY BOTANICAL CLUB

25

FIGURE 39. Range of Gaura parviflora in the United States and Mexico.

established in the United States in Alabama, California, Florida, Georgia,
Massachusetts, and Tennessee. Introduced in Argentina (Cordoba, San
Luis), China, Okinawa, and northern Australia. Flowers throughout the
warm part of the year. In Argentina, Gaura parviflora grows in disturbed
ground, near roads and houses, in ravines, and along intermittently dry
streams at 500 to 1000 m elevation. The earliest collection we have seen
from South America is that of Hieronymus, cited above, which was made
in 1876.

Representative Specimens Examined (outside United States only):

Argentina, cordoba. Dpto. Calamuchita: Valle de los Reartes, Castellanos 48
(GH); La Cruz, ca. 850 m, Gutierrez 109 (BH). Dpto. Capital: Alrededores de la
Ciudad de Cordoba, A. T. Hunziker 8108 (RSA); alrededores del cementerio, Stuckert
3610 (G, GH). Dpto. Isehilin: Quilino, Villafafio 76 (NY, TEX, US). Dpto. Punilla:
Valle Hermoso, Carlson in 1940 (RSA); Sierra de Cordoba, Capilla del Monte, Barros
in 1925 (GH); San Francisco, Sierra de Cordoba, Hieronymus 578 (NY); Sierra
Chica, Losser 226 (BY, F, GH, MO, PH); Dolores, Valle de Punilla, Nicora 1673
(POM); San Salvador, 970 m, Rodrigo 364 (US). Dpto. San Alberto: Sierra Grande,
Mina Clavero Burkhart 7328 (F, POM); Sierra Grande, 10 km below El Condor, 1000
m, Goodspeed 17242 (GH); Nono, O’Donnell 6- Rodriguez 714 (A). Dpto. San
Javier: La Barranca, Castellanos in 1939 (RSA); Quebrada de Yacanto, Castellanos in
1927 (RSA); La Poblacion entre Yacanto y Luyaba, A. T. Hunziker 11462 (RSA).
Dpto. Santa Maria: Alta Gracia, ca. 600 m, ]. H. Hunziker 1158 (RSA).

26

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Australia, new south wales. Wiangaree, shire, of Kyogle, Mackatj in 1967
(NSW); banks of Richmond River, nw of Kyogle, Salasoo in 1963 (NSW'). Queens¬
land. Pittsworth, White 6662 (BY, GH, NY); Darling Downs District, Jandowae to
Dalby road at Pirrinauri, Doherty in 1959 (K); Millars Vale Creek, Marvvale, Glen-
gallen Shire Council in 1958 (K).

China, chekianc. Chienteh, Ling 2738 (NY).

Japan. Okinawa. Kadekaru, Kume-jima Island, 1966, Nakejima 1682 (DS)
(seen since about 1954).

Mexico, baja California. San Jose del Cabo, Anthony 330 ( BM, DS, F, GH, K,
MENU, MIN, MO, POM, US); Miraflores, Jones in 1928 ( DS, POM); Todos Santos,
Jones 24137 (DAO, DS, DUKE, F, MICH, MO, NY, POM, PH, RSA, SMU, TEX);
Mision Guadalupe, near 26°54'N, 112°26'W, ca. 700 m, Moran 11788 (DS); Mulege,
Palmer 11 (GH, US); Santiago, Thomas 7804 ( DS, RSA); ca. 19 mi w of Tecate
on road to Tijuana, Wiggins 6- Thomas 471 (DS). chihuahua. 6 mi s of Ciudad
Juarez, Breedlove 15727 (DS; 20 km s of Ciudad Camargo, 1300 m, White 2228

(GH, MENU, MICH); near Lake Santa Maria, Nelson 6406 (US); vicinity of

Chihuahua, 1300 m. Palmer 185 ( F, GH, MO, NY, US); Valle de Rosario, Pennington
411 (TEX); Jimenez, Pringle 7529 (MEXU); 45 mi s of Ahumada, Moldenke 2088
(DS). coahuila. Viesca, Fisher 3725 (NY); near Saltillo, 1500 m, Hinton 16798
(GH); 1 mi se of Ocampo, Johnston 8884 (GH); Muzquiz, Marsh 946 ( F, GH,
TEX); Monclova, Gloria Mts., Marsh 1667 ( F, GH, LL, OKLA, TEX); 6 mi w of
San Pedro, 4000 ft, Munz 15041 (GH, MO, POM), durango. 10 mi sw of Ciudad
Lerdo, Munz 15037 (C.H, MO, POM); Durango and vicinity, Palmer 102 ( F, GH,
MO, NY, US), nuevo leon. Monterrey, 550 m, Arsene et al. in 1911 (NY, RSA);
Poblado de Higueras, Hernandez et al 16M508 (TEX); 16 km w of Sabinas Hidalgo,
Dominguez ir McCart 8266 (TEX), sinaloa. Culiacan, Rose et al. 14867 (NY, US);
Las Palmas, Sinaloa, Gonzalez Ortega 4542 (MEXU). sonora. 7-9 mi w of Navojoa,
Gentry 7976 ( DS, MEXU, MICH, US); El Desierto, Rio Mayo, Gentry 3044 ( F,
GH, MEXU, MO, US); Cochuto, 5100 ft, Hartman 96 ( C.H, NY, PENN, PH, US);
35 mi w of Sonoyta, Keck 4227 ( DS, POM); Hermosillo, Maltby 254 (US); Alamos,
Roe et al. 12961 (NY, US); Arroyo del Pulpito, near Colonia Oaxaca, White 777

( DS, MICH); Bavispe, Rio Bavispe, White 2902 (GH, MICH); Horconcitos, Rio

Huachinera, White 2968 (GH, MICH, PH); Colonia Morelos, 2600 ft, White 4419
(GH, MICH, NY, US); Moctezuma, White 299 (GH, MICH); 42 mi e of Caborca,
Moldenke 1536 (DS). zacatecas. 26 mi se of Sombrerete, Breedlove 14341 (DS).

Gaura jtarviflora was probably originally a native of the shortgrass
prairie in the interior of the United States, and spread widely from there
as a weed of cultivated and waste areas. Owing to its strict autogamy,
it easily becomes established from a single fruit. It has been cultivated
as a curiosity in botanical gardens since at least the 1840s. The small dif¬
ferences in pubescence that have been reflected in the naming in infra¬
specific taxa are consistent with what would be expected in an autogamous
species and do not deserve fonnal taxonomic recognition.

According to information kindly sent by Mr. S. L. Everist this species
is now extensively naturalized in southeastern Queensland, Australia. The
first known collection was made* at Charter’s Towers in the North Kennedy

MEMOIRS OF THE TORREY BOTANICAL CLUB

27

District, where it has not persisted, in 1929. The Pittsworth collection,
cited above, was obtained in 1930, and from this locality the plant has
spread out over an extensive area. In 1950 it was eollected at Roma, in
the Maranoa District, its westernmost station. It was first noted at Goon-
diwindi in the Darling Downs District, its southernmost locality in Queens¬
land, in 1956, and had spread into the vicinity of Kyogle in New South
Wales by 1963, as shown by specimens cited above. All of the collections
from the north end of the range of this species in Queensland, excluding
the 1921 record (where the plant failed to persist), have been made
since 1960 and a collection from Mulgildie, now the northernmost limit
of the plant in Australia, was made for the first time in 1969. Mr. Everist
has suggested that it is quite possible that Gaura parviflora has still not
reached the northern limits of its range in Queensland.

SECTION IV. CAMPOGAURA

Gaura sect. Campogaura Raven & Gregory, sect. nov.

Herbae perennes confertae, caulibus subglabris vel strigulosis, saepe
etiam villosis. Inflorescentiae nutantes. Flores valde zygomorphi, vesper-
tini. Fructi 4-angulati, parte superiore 4-costato, 4-angulato, ad basin at¬
tenuate, tereto.

Clumped perennial herb, the stems subglabrous or strigulose, often
also villous. Inflorescence nodding. Flowers strongly zygomorphic, the
stamens in the lower half; flowers opening near sunset and withering late
the next morning. Fruit not reflexing, 4-angled, with 4 sharp ridges and
4 furrows alternating on the pyramidal upper half, constricted sharply
below, the base terete, narrowing gradually toward the bottom but at
least half the diameter of the top throughout. Self-incompatible. Gametic
chromosome numbers, n = 7, 14, 21, 28.

Type Species: Gaura coccinea Pursh

This distinctive species of the prairies and plains (hence the sectional
name) is clearly related to the species of sections Stipogaura and Xero-
gaura. It is probably one parent of the tetraploid Gaura drummondii.

5. Gaura coccinea Pursh, FI. Amer. Sept. 2: 733. 1814. (Norn, nud.,
Frasers Cat. 1813.) Munz, Bull. Torrey Bot. Club 65: 220. 1938; N.
Amer. FI. II. 5: 194. 1965. Figs. 5, 35.

Gaura odorata Sesse ex Lag., Gen. et Sp. Nov. 14. 1816. Type: Described
from plants cultivated in the Royal Botanical Garden at Madrid, the
seed brought from Mexico in 1804 by Martin de Sesse y Lacasta. See
discussion on pp. 54, 55.

28

A REVISION OF THE GENUS GAURA ( ONACRACEAE )

Gaura epilobioides H.B.K., Nov. Gen. et Sp. 6: 93. 1823. Type: Near
Actopan, Hidalgo, Mexico, May 1803, Alexander von Humboldt ( P.
Isotype, B. Photographs, F, GH, POM, US).

Gaura coccinea Pursh (S? integerrima Torr., Ann. Lyceum Nat. Hist.
New York 2: 200. 1828. Type: Major Long’s Creek, near Stead,
Union Co., New Mexico, 1 August 1820, Edivin James (NY).

Gaura parvifolia Torr., Ann. Lyceum Nat. Hist. New York 2: 201. 1828.
Type: Major Long’s Creek, near Stead, Union Co., New Mexico, 1
August 1820, Edwin James (NY).

Gaura? suffrutescens Moc. & Sesse ex Ser., in DC., Prod. 3: 45. 1828.

Type: FI. Mex. Ic. (ined.), t. 374 (G-DC).

Gaura bracteata Ser., in DC., Prod. 3: 45. 1828. Type: In gardens,
San Angel, Mexico, Mexico, Fl. Ic. (ined.), t. 373 (G-DC).

Gaura multicaulis Raf., Atl. Jour. 146. 1832. Based on Gaura coccinea
Pursh (}? integerrima Torr.

Gaura marginata Lehm., in Hook., Fl., Amer.-Bor. 1: 208. 1834. Type:
Plains of the Saskatchewan River, Canada, T. Drummond (Isotvpes,
BM, BR, NMC).

Gaura glabra Lehm., in Hook., Fl. Amer.-Bor. 1: 209. 1834. Type:
Vicinity of Fort Carleton, Saskatchewan, T. Drummond (Isotypes,
BM, BR, LE).

Schizocarya kunthii Spach, Nouv. Ann. Mus. Hist. Nat. 4: 383. 1835.
Based on Gaura epilobioides II.B.K.

Gaura coccinea Pursh var. glabra (Lehm.) Munz, Bull. Torrey Bot. Club
65: 224. 1938.

Gaura spicata Sesse & Moc., PI. Nov. Hisp. ed. 1. 56. 1888. Type: Vicinity
of Mexico City, D. F., Mexico, June.

Gaura induta Wooton & Standi., Contr. U. S. Natl. Herb. 16: 153. 1913.
Type: Dry, clay hills near Pecos, 2010 m, San Miguel Co., New Mexico,
15 August 1908, P. C. Standley 4933 (US 498956. Isotypes, CH, NY).
Gaura linearis Wooton & Standi., Contr. U. S. Natl. Herb. 16: 154. 1913.
Type: Gypsum soil near Lakewood, Eddy Co., New Mexico, 6 August
1909, E. O. Wooton (US 564593).

Gaura coccinea Pursh var. parvifolia (Torr.) Rickett, Kew Bull. 1934:
57. 1934. Munz, Bull. Torrey Bot. Club 65: 223. 1938; N. Amer. Fl.
II. 5: 196. 1965.

Gaura coccinea Pursh var. typica Munz, Bull. Torrey Bot. Club 65: 221.
1938.

Gaura coccinea Pursh var. epilobioides (II.B.K.) Munz, Bull. Torrey Bot.

Club 65: 222. 1938. Munz, N. Amer. Fl. II. 5: 195. 1965.

Gaura coccinea Pursh var. arizonica Munz, Bull. Torrey Bot. Club 65:

MEMOIRS OF THE TORREY BOTANICAL CLUB

29

225. 1938. Type: Globe, Gila Co., Arizona, 17 May 1919, Alice East-
wood 8657 (GH). Munz, N. Amer. FI. II. 5: 196. 1965.

Clumped perennial herb, from a deep, thick taproot, often branching
several centimeters below the ground and giving rise to underground
stems, or branching only at the surface or not at all; underground stems
often becoming horizontal or nearly so and giving rise to new plants; stems
1-12 dm tall, the plants varying from full and copiously branched at the
base, little above, to strict and little branched at the base, copiously above.
Entire plant densely strigulose with long spreading trichomes often present
near the base to subglabrous with strigulose pubescence on ovary, out¬
side of floral tube, and sepals, and sometimes at edges of lower leaves also.
Leaves linear to narrowly elliptic, 0.7-6.5 cm long, 0. 1-1.5 cm wide, entire
to remotely and coarsely serrate. Inflorescence 8-47 (-65) cm long, the
peduncle 1-6.5 cm long, strict or well branched; bracts subulate, 2-5 mm
long, 0.4-1. 1 mm wide. Floral tube 4-11 (-13) mm long. Sepals 5-9 (-10)
mm long, (0.75— ) 1-1. 5(-2) mm wide. Petals white fading to orange-red
to deep maroon, very rarely pale cream, 3-7 (-8) mm long, 2-4 mm wide.
Filaments cream, 3-6.5(-7) mm long; anthers yellow to light or deep red,
(2.5-) 3-5 (-5.5) mm long. Style (9.5-) 10-21 (-21.5) mm long. Fruit 4-
8.5 (-9) mm long, (1-) 1.5-3 mm thick at the thickest point. Seeds (1-)
3-4, 1.5-3 mm long, 1-1.5 mm thick, light to reddish brown. Self-incom¬
patible. Gametic chromosome numbers, n = 7, 14, 21, 28 (with intermediate
numbers and supernumerary chromosomes occasionally found).

Type: Near Mandan, Morton Co., North Dakota, 22 June to 5 July 1811,
John Bradbury (PH).

Distribution (fig. 40): Frequent in light, sandy soil, usually at middle
elevations and ascending to about 3000 m in Mexico; local in the Kootenay
District of British Columbia, and from the valley of the North Fork of the
Saskatchewan River at about 53°40'N lat. near Edmonton, Alberta, and
about 52°40'N lat. near Battleford, Saskatchewan south throughout southern
Alberta, Saskatchewan, and Manitoba, and through Montana, North Dakota,
and western Minnesota south, mainly east of the Continental Divide to
central Colorado, northwestern Missouri, western Oklahoma, central and
western Texas, southern Utah, southernmost Nevada, and northeastern San
Bernardino County, California, then southward through northeastern Sonora
and the mountains of Chihuahua, Coahuila, Nuevo Leon, and Tamaulipas
to the Trans-Mexican Volcanic Belt (where abundant); rare in southern
Puebla and northern Oaxaca at about 17°N lat. Introduced and collected
once each in Chiapas, Mexico, in Rio Grande do Sul, Brazil, and in Wales.
Also a sporadic weed in Wisconsin, Illinois, Indiana, Michigan, New York,

MEMOIRS OF THE TORREY BOTANICAL CLUB

31

coastal southern California, and in southern Ontario. Possibly native at
Piute Creek, Kern County, California (6 June 1893, N. C. Wilson, DS, NO),
but probably introduced. Flowers April to August.

Representative Specimens Examined (from outside of the United States):

Brazil, rio grande do sul. Porto Alegre, Reinick in 1899 (GH).

Great Britain, wales. Carmathenshire: Adventive, Pembrey Burrows, Hollis
in 1935 (K).

Mexico, aguascalientes. Aguascalientes, Kuntze 23484 (NY). chiapas. Es-
cuintla, Matuda 676 (MEXU, MICH, US), chihuahua. Santa Eulalia, pass between
Velardena and Cristo mines, ca. 28°37'N, 105°53'W, Hewitt 294 (GH); near Charco,
Johnston 8134 (GH); mountains w of Chihuahua, LeSueur 805 (F, NO, SMU, TEX);
foothills of the Sierra Madre near Colonia Juarez, Nelson 6336 (GH, US); end of
Sierra del Diablo, Stewart 1943c (GH); 7 mi s of junction of Casas Grandes Hwy. on
Mexico Hwy. 45, Straw 6- Forman 1842 (MICH, RSA); Mpio. de Janos, 4800 ft,
White 2572 (GH, MEXU, MINN); 12 mi s of Camargo, 400 ft, White 2229 ( GH,
MICH), coahuila. Saltillo, 1600 m, Arsene 6- Adola 6346 (US); Parras, Kenoijer
ir Crum in 1948 (A); Viesca, 3725 ft, Fisher 44108 (GH, MO, NSC, NY, SMU);
Arteaga Road, Kenoijer 6- Crum 2659 (A, MICH); 10 mi e of Arteaga (ca. 20 mi e
of Saltillo), Straw 6- Forman 1349 (MICH, RSA); 6 mi n of La Ventura on road to
Saltillo, Johnston 7645 (GH); Muzquiz, Marsh 2119 ( F, GH); Monclova, Marsh 1704
(TEX); Sierra de Santa Rosa, s of Muzquiz, Marsh 2272 (GH, TEX); 13 mi s of
Saltillo, McGregor 16713 ( DS, KANU); 6 mi w of San Pedro, Munz 15040 (C.H,
MO, POM ) ; end of road from T. Armendaiz N. into the Sierra del Pino, Johnston ir
Mueller 690 (GY); Sierra de las Cruces, Johnston 6- Mueller 231 (GH); 9 km s of
Parras on Sierra Negra, Stanford et al. 240 (GH, ILL, DS, MEXU, MO, NY); Fraile,
59 km s of Saltillo, Stanford et al. 251 ( DS, GH, ILL, MEXU, MO, NY); 11 km
ne of Jimuleo, 2100 m, Stanford et al. 20 ( DS, GH, MO, NY); 15 km w of Con¬
cepcion de Oro, Stanford et al. 537 ( DS, GH, MO, NY); 25 km s of Piedras Negras,
Rinehart 216 (OKL); Carneros Pass, Pringle 3105 ( BR, CM, COCO, F, G, GH, LE,
MICH, MEXU, MO, MSC, ND, NY, RSA, US); Canon del Indio Felipe, Sierra
Hechiceros, Stewart 19 (GH); n of La Ventura, 6000 ft, Shreve 6- Tinkhorn 9610
(GH); Paila, Km 570, 1200 m, Aguirre & Reko 42 (NY); 8 km ne of Santa Elena,
Stewart 1122 (GH); n slope of Sierra Planchada, 5 mi w of El Oro, White 1998
(MEXU, MICH); 60 mi w of Cuatro Cienegas, White 1955 (DS, GH, MICH).
durango. Chupaderos n of Durango, Pennell 18181 (MEXU, PH, US); near Yerbanis,
Shreve 9148 (GH); 11 mi n of La Zarca, Waterfall 12541 (GH, MICH, OKLA, US);
34-38 mi ne of Durango, Waterfall 13359 ( IKLA, SMU); 0.5 mi w of El Pino, ca.
10 mi w of Durango, Waterfall ir Wallis 13359 (IA, OKLA); 1 mi s of La Zarca,
Wiens 3524 (COLO). Guanajuato. Acambaro, Duges 303 (GH); San Miguel Allende,
Kenoijer 1913 (GH); Leon, Matuda M 19057 (F); Irapuato, Woodruf 144 (NY).
hidalgo. Vicinity of Zimapan, Kenoijer 1058 (GH); Tulancingo, Cuautepec, Moore
1601 (BH); near Tequixquiac, Rose 6- Painter 6636 ( GH, US ) ; near El Salto,
Rose 6- Painter 8031 (US); Hacienda Palmar near Pachuca, Rose i? Painter 8820
(NY, US), jalisco. Near Km 31 sw of Ojuelos on road to Aguascalientes, above Presa
de Valerio, McVaugh 16891 (MICH, LIS); 1.5 mi s of Ojeulos, Winetraub 6- Roller 33
(MICH). Mexico. Alrededores de Atlahutenco, Mpio. de Ecatepec Morelos, 2250 m,
Cruz 508 ( DS, MICH, TEX); Teotihuacan, 2200 m, Froderstrom <b Hulten 1198
(NY); Los Reyes, on road to Puebla, Longman 2470 (PH); near Texcoco, Mac-

32

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Daniels 577 (BY); Tlamanalco, 7000 ft, Munz 15051 (GH, MO, POM)- field above

mKr'mTf''™ (COL°' DA°’ DUKE' F' CH' IA- LL- KANU, MIN,
, ; ISC, OKI. A, SMU, TEX, US); Lomas de Chapultepec, D. F. Barkley i?

xv £? iFLAS> MICH- TEX); Vera Cruz’ 8300 ft, Guadalupe Victoria,

f, oES' Vs l NUE™ LE6N' Near Cerro GrantIe" ca' 3 mi s» “f Ascencion,
7000-8000 ft, Straw tr Forman 1400 ( MEXU, RSA); 67 km nw of San Roberto on

^ ,Rne 6 Mor‘ 50 (WISC>1 Sabinas Hidalgo, Ports in 1941

( L); 38 mi se of Saltillo, Waterfall 16615 (OKL, OKLA, RSA, SMU). Oaxaca,
Ua. o mi s of I amazulapan, Straw 6- Gregory 1038 (GH, MICH, MEXU, RSA).
plebla. Rancho Posada, 2194 m, vicinity of Puebla, Arsetw 95 (US)- Cholula

DTnm(<Gw t10i); TehuaCi‘n’ 1 N ■ ** L S' Rose 9960 <GH’ US); near Tepeaca!
Tool ™ ' E; 6 M ' S' ManninS 53900 (FLAS, GH, TEX); 45 mi e of Puebla,

ft’ UU f/ 6 M' S- Manning 53708 (GH); Acatzingo, 7500 ft, Munz 15068
(t-H MO); Hwy. from Tehuacan to Orizaba just above Azumbilla, 1500-1800 m

™lt\et al 9959 (F’ CH’ NY> US). SAN LUIS potosi. 16 mi e of San Luis Potosi,’
Breedlove 14364A (DS); 13 mi nw of San Luis Potosi, Breedlove 15471 (DS)-
Charcas, Landed 5137 (MICH, NY, US), sonora. 23.5 mi ne of Bocoachic on road
to Esqueda 4480 ft, Wiggins 11742 (DAO, DS, TEX, US), tamaulipas. Between
Marcela and Hermosa, Stanford et al. 2662 ( DS, GH, MO, NY, RSA). tlaxcala.
Hwy. to Veracruz ca. 9 km e of Mexico, Lambert 41-23 (PENN, PH); Km. 185
on road from Huamantla to El Carmen, Sohns 624 (US); 10 mi n of San Martin
Texmelucan, Sohns 659 (US), veracruz. Mt. Orizaba, 4000 ft, Seaton 266 (GH)-
Maltrata, Matuda 1219 (MEXU, MICH); Jalapa, Coulter 164 (GH). Zacatecas’.
32 mi nw of San Luis Potosi, Breedlove 15476 (DS); Pedernalillo, near Guadalupe,
Dressier 306 (MO); Fresnillo, Knobloch 1030 (MICH, MSC); near Concepcion del
Oro, Palmer 318 (GH, NY, US); 22 mi n of Zacatecas, Straw 6 ■ Forman 1493 (GH,
MEXU, RSA); 2 mi se of Sombrereto, Waterfall 13804A (OKLA, IA).

Gaura coccinea is commonly known in Mexico as “linda tarde” or
hierba de and in Texas as wild honeysuckle. There appears to

be nothing to be gained by ascribing this species to “Nutt, ex Pursh,” as
Nuttall is not mentioned either by Pursh or in the Fraser Catalogue of
1813, even though we now know that Nuttall was the author of this
anonymous catalogue (cf. Reveal, Rhodora 70 : 25-54. 1968). Pursh clearly
based his species upon the Bradbury specimen cited above, and not on any
Nuttall specimen; indeed he remarks that the habit of his plant differs from
that indicated (by a sign) in the Fraser Catalogue, where the name is, in
any case, a nomen nudum. Pursh did include in his protologue a reference

to the Catalogue. We are indebted to J. E. Dandy for his remarks on this
matter.

As attested by its rather complex synonymy, Gaura coccinea is a variable
species. Over much of its wide range, it is relatively uniform, with the
plants low, usually 2-5 dm tall, branching mostly at and below ground
and not much above, and often forming large colonies partly by means of
vegetative reproduction. The inflorescence in such plants is usually less than
5 cm long. Plants from southern Nevada and northwestern Arizona are

MEMOIRS OF TIIE TORREY BOTANICAL CLUB

33

taller but usually do not differ in habit, although they occasionally branch
more than usual above. Such plants represent one end of a very gradual
cline which begins in north-central Arizona.

From west and central Texas north into the panhandle and southern
Oklahoma, the plants are taller, usually 5-10 dm tall, unbranched or with
a very few branches from the base, but well branched above, and occur
in clumps, spreading very little by rhizomes. The more numerous branches
are long, with the central leader generally more than 5 cm long and longer
than that of the widespread race at a comparable stage of growth. There
is a very gradual transition from this race to the more widespread one,
with many populations and individual plants impossible to assign to one
or the other. Under the circumstances, it does not appear desirable to
reflect these differences in the formal taxonomy.

Chromosoinally, Gaura coccinea is very complex, with diploid (n = 7),
tetraploid (n=14), hexaploid (n = 21), and octoploid (n = 28) popula¬
tions. The most extreme expression of the “Texas” race is found in the
diploids, which are confined to Trans-Pecos Texas and adjacent New
Mexico, and are relatively uniform morphologically. In general, the diploids
have the smallest flowers found in the species, although the ranges of
measurements of flower parts overlap broadly. Tetraploids and hexaploids
found in adjacent regions are in general intermediate in their characteristics
between the diploids and the more widespread expression of the species.
The transitions are often not gradual. For example, in the area of the
Davis Mountains in west Texas, populations of hexaploids growing within a
few miles of one another may represent one or the other extreme or various
intermediates. Around Carrizozo in south-central New Mexico the same
sort of mixture of distinctive populations is found among the tetraploids.
On the other hand, north of the central Texas panhandle and south of the
Rio Grande (the sampling is relatively poor in Mexico), the populations
seem relatively uniform and to change gradually over vast areas.

The octoploids occur locally as populations in predominantly tetraploid
areas. They were presumably derived from tetraploid plants directly.
Both tetraploids and hexaploids occur widely in the central and southern
parts of the range of the species, and are, like the octoploids, indistinguish¬
able morphologically except occasionally on a very local basis. Neither
pollen size nor pore number is useful in distinguishing chromosomal races
in Gaura coccinea.

The consistent occurrence of autopolyploid pairing in all polyploids of
this species indicates a lack of chromosomal differentiation. The simplest
explanation for the observed pairing would seem to be an initial stabilization
of a diploid race of plants similar to those occurring in west Texas and a

34

A REVISION OF THE GENUS GAURA (ONAGRACEAE)

tetraploid race with the characteristics of populations found in the central
and northern parts of the species area. There is no reason to postulate an
allotetraploid origin for the latter; indeed, the consistent autotetraploid
association of its chromosomes and the presence of but one diploid in the
complex militate against such an hypothesis. Nor can it confidently be
asserted that the tetraploid originated from the diploid; cytogenetic evidence
would be equally consistent with the opposite point of view ( cf., Raven
and rhomson, Amer. Naturalist 98: 251, 252. 1964), and there appears to be
no logical basis for distinguishing between these two theories. The re¬
striction of the diploid to an area where many related diploid species occur,
however, lends credence to the idea that it may be ancestral rather than
derived.

Given the initial presence of these two races, it is a simple matter to
visualize the development of the pattern seen at present. The origin of
the tetraploids from diploids and of octoploids from tetraploids would be
consistent with the geographical patterns observed and the lack of morpho¬
logical differentiation. The opposite processes may also have taken place
occasionally. With the sort of chromosomal equivalence that prevails in
Gaura coccinea, hexaploids can originate directly from the functioning of
unreduced tetraploid gamete, from tetraploid-octoploid hybridization, or
in the conventional way. Similarly, tetraploids can probablv be derived in
one step from hexaploid-diploid crosses. The proliferation of polyploid
races and the subsequent hybridization within and doubtless also between
chromosomal levels is attested by the pattern displayed from this complex
species and the impossibility of finding morphological discontinuities in it.
In our survey of approximately 300 wild plants, approximately 5 apparent
interploid hybrids were encountered (Raven and Gregorv, Brittonia 24: 71-
86. 1972).

SEGTION V. STIPOGAURA
Gaura sect. Stipogaura Raven & Gregory, sect. nov.

Herbae perennes confertae vel rhizomatosae, caulibus strigulosis vel
glandulosis, saepe etiam villosis. Infiorescentiae erectae. Flores valde
zygomorphi, vespertini. Fructorum corpus ovoideum, lateribus concavibus,
longe abrupteque stipitatum.

Clumped or rhizomatous perennial herbs, the stems strigulose or glan¬
dular, and usually villous also. Inflorescence erect. Flowers strongly zygo-
morphic, the stamens in the lower quarter or half or well distributed;
flowers opening near sunset and withering early the next morning. Fruit
reflexing at maturity, with the bodv ovoid, winged along the angles, the

MEMOIRS OF THE TORREY BOTANICAL CLUB

35

sides concave with the upper half of each concavity forming a shallow
trough, abruptly narrowed into a long stipe. Self-incompatible. Gametic
chromosome numbers, n = 7, 14.

Type Species: Gaura villosa Torr.

Distribution: Sandy and rocky slopes and flats from southeasternmost
Colorado and Texas south to eastern Durango, Coahuila, Nuevo Leon, and
western Tamaulipas, Mexico.

The five species of section Stipoguara are closely related to one another
and present an interesting pattern of relationship. Three— Gaura calcicola,
G. mckelveyae, and G. filipes— are medium-sized to small clumped diploids.
They replace one another geographically, with G. calcicola on rocky slopes
at moderate elevations in west Texas and southward into Mexico, G.
mckelveyae on sandy plains at low elevations in and near the Rio Grande
valley, and G. filipes in the southeastern United States. Gaura sinuata,
an aggressively rhizomatous weedy tetraploid, was very probably derived
following hybridization between G. calcicola (or the common ancestor of
G. calcicola and G. filipes, which are extremely similar morphologically)
and G. mckelveyae. It is somewhat intermediate ecologically and has spread
rather widely as a weed of cultivated fields. The fifth species, Gaura villosa,
is a large, handsome plant of the dunes and sandy flats in south and west
Texas and adjacent states, apparently separated from the others by its
extreme ecological specialization. It is listed first in the section not because
woodiness is thought to be primitive, a dubious proposition, but rather
because it is thought to have been a rather early offshoot of the common
ancestor of the other species.

6. Gaura villosa Torr., Ann. Lyceum Nat. Hist. New York 2: 200. 1827.

Munz, Bull. Torrey Bot. Club 65: 213. 1938; N. Amer. FI. II. 5: 192.
1965.

Clumped woody perennial of dunes and sandy flats, from a deep,
twisted, woody rootstock; stems sending up several branches near ground
level and branching again where the inflorescences originate to form at
maturity a full, leafy plant with long inflorescences rising above; stems
6-18 dm tall. Plants densely villous with hairs 2-3 mm long all over below
the inflorescence, sometimes more sparsely so, and with a dense under¬
story of strigulose or glandular hairs; inflorescence strigulose, glandular-
pubescent, or hirtellous; leaves densely covered with loose, strigose pu¬
bescence, rarely with a few glandular hairs below. Leaves oblanceolate to
very narrowly elliptic or linear, 0,5-8 cm long, 0.15-2 cm wide, subentire or
shallowly sinuate-dentate, occasionally markedly so, often undulate, acum¬
inate at the tip, sessile. Inflorescence generally well branched, 2o-127 cm

36

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

44 45 46

FIGURES 41-46. Flowers of the species of Gaura sect. Stipogaura. Fig. 41. G.
villosa subsp. villosa.— Fig. 42. G. villosa subsp. parksii.— Fig. 43. G. calcicola.— Fig.
44. G. filipes.— Fig. 45. G. mckelveyae.— Fig. 46. G. sinuata.

MEMOIRS OF THE TORREY BOTANICAL CLUB

37

long; bracts narrowly lanceolate to narrowly ovate, 1-6 (-10) mm long, 0.75-

1.5 mm wide. Flowers with stamens in the lower quarter. Floral tube 1.5-5
mm long. Sepals 6-14 mm long, 1-2.5 mm wide. Petals 7-13 mm long,
4-6 mm wide. Filaments 4.5-11 mm long; anthers 2-4.5 mm long. Style
9-18.5 mm long. Body of the fruit 9-19 mm long, 1-3.5 mm thick, the
stipe 2-10 mm long. Seeds (l-)2-4, 2-3(-4) mm long, 0.75-1.25 mm
thick, yellowish to light brown or rarely reddish brown. Self-incompatible.
Gametic chromosome number, n = 7.

Type: Along the upper Canadian River, in Hartley, Oldham, or Potter
Co., Texas, 4-9 August 1820, Edwin James (NY).

Distribution (fig. 38): Dunes and sandy flats, Rio Grande Plain of
south Texas; High Plains and Rolling Plains of northwestern Texas, eastern
New Mexico, southeastemmost Colorado, southwestern Kansas (one station
in Ellis Co.), and the western half of Oklahoma; locally naturalized in
New Jersey.

The two subspecies recognized in Gaura villosa have a disjunct distribu¬
tion but are so closely similar that it seems best to emphasize these sim¬
ilarities by maintaining them within a single species.

6a. Gaura villosa subsp. villosa. Figs. 6, 41.

Gaura cinerea Wooton & Standi., Contr. U. S. Natl. Herb. 16: 152. 1913.
Type: 20 mi s of Roswell, 1080 m, Chaves Co., New Mexico, August
1900, F. S. 6- E. S. Earle 533 (US 382592. Isotype, NY).

Gaura villosa Torr. var. typica Munz, Bull. Torrey Bot. Club 65- 214
1938.

Gaura villosa Torr. var. arenicola Munz, Bull. Torrey Bot. Club 65: 215
1938. Type: 5 mi ne of Portales, on sand dunes near U.S. Hwy. 70,
Roosevelt Co., New Mexico, 14 June 1930, G. /. Goodman & C. L.
Hitchcock 1124 (POM 171881. Isotypes, MICH, NY). Munz, N.
Amer. FI. II. 5: 192. 1965.

Plants with an understory of either strigulose or glandular hairs, the
corresponding kind covering the ovaries, outside of the floral tube, and
sepals more densely; branches of the inflorescence either glabrous or sparse¬
ly glandular-pubescent; leaves rarely with a few glandular hairs beneath.
Leaves narrowly lanceolate to very narrowly elliptic or linear, occasionally
narrowly elliptic to lanceolate or even narrowly oblanceolate toward the
base of the plant, subentire or shallowly sinuate-dentate, occasionally mark¬
edly so. Inflorescence whiplash-like, comparatively stout. Floral tube 2-5
mm long. Petals 8.5-13 mm long. Filaments 5-11 mm long; anthers 2.5-

4.5 mm long. Style 10-18.5 mm long. Body of the fruit 9-18 mm long.

38

A REVISION OF THE GENUS GAURA (ONAGRACEAE)

2-3.5 mm wide, the stipe 2-8 mm long. Self-incompatible. Gametic chro¬
mosome number, n — 7.

Distribution (fig. 38): Sand dunes and sandy flats, High Plains and
Rolling Plains of northwestern Texas, eastern New Mexico, southeastern-
most Colorado, southwestern Kansas (one station farther north, in Ellis
County), and the western half of Oklahoma; locally naturalized in New
Jersey.

Representative Specimens Examined:

United States. Colorado. Baca Co.: 23 mi s of Walsh, Stephens 6- Brooks
21792 ( DS, KANU); s bank of Cimarron River at se comer of county, Weber 5142
(COCO, COLO, DAO, DS, GH, KANU, NSC, OKL, RSA, SMU, TAES, TEX, US).
Kansas. Barber Co.: 3 mi s of Medicine Lodge, Horr ir Franklin in 1940 (KANU).
Clark Co.: 10 mi s of Ashland, Rydberg ir Imler 749 (KANU, NY). Comanche Co.:
Coldwater, Rydberg ir Imler 678 (NY). Ellis Co.: Hays, Bondy in 1937 (OKL,
UMO). Kiowa Co.: 5 mi n of Greensburg, McGregor 10634 (KANU). Meade
Co.: 2 mi w of Meade Co. State Park, Horr 3668 (KANU). Morton Co.: 9
mi n of Elkhart, McGregor 5105 (KANU). Pratt Co.: Turkey Creek, Norris 193
(MO). Seward Co.: 20 mi ne of Liberal, Rydberg ir Imler 852 (KANU, NY).
Stevens Co.: Hitchcock in 1897 (POM), new jersey. Cape May Co.: S of New
England Road, Cold Spring, Brown in 1921 (PENN), in 1922 (PH), new Mexico.
Chaves Co.: 30 mi s of Roswell, F. S. 6- E. S. Earle 533 (MIN, MO, NCU). Colfax
Co.: W of Willow Bar, Fendler 213 (MO). Curry Co.: 9 mi s of Broadview,
Stephens ir Brooks 25624 ( DS, KANU). De Baca Co.: Fort Sumner, Shinners
20924 (SMU). Guadalupe Co.: Santa Rosa, Cooley 3 (OKLA). Lea Co.: 3.3 mi
e-ne of Hobbs, Shinners 20071 (SMU). Quay Co.: 3 mi s of Sand Springs, Wagen-
knecht in 1957 (KANU). Roosevelt Co.: 5 mi ne of Portales, Goodman L~ Hitchcock
1124 ( GH, MIN, POM). Oklahoma. Alfalfa Co.: 1 mi nw of Gonot Salt Plains Dam,
Waterfall 7834 (OKL). Beaver Co.: 2 mi e of Headquarters, Hindman 216 (NCU,
OKLA). Blaine Co.: 3 mi s, 1.5 mi w of Watonga, Cox in 1954 (OKLA). Caddo
Co.: E of Bridgeport, Munz 13576 (POM). Cimarron Co.: S of Walsh, Weber 5210
(COLO, NSC). Cleveland Co.: 6 mi nw of Norman, Bruner in 1924 (MIN).
Comanche Co.: Wichita Mts., Marcy’s Expedition. Cotton Co.: 4 mi s of Randletter,
Waterfall 7283 (OKL, OKLA). Custer Co.: 1/4 mi s, 2 mi e of Weatherford, Water¬
fall 2238 (OKL, OKLA). Dewey Co.: Near Canton, Stevens 871 ( DS, GH, ILL, MO,
MIN, OKL, OKLA). Ellis Co.: Pack Saddle Bridge, Canadian River valley, Goodman
2591 (ISC, MO, OKL, POM). Grady Co.: 3/4 mi e of Juttle, Pearce 690 (OKL).
Grant Co.: 20 mi w of Medford, Waterfall 7366 (OKL, OKLA). Greer Co.: 1 mi e
of Magnum, Waterfall 7309 (OKI,, OKLA). Hannon Co.: 1.7 mi w of Vinson,
Waterfall 7767 (OKL). Harper Co.: 4 mi s of Rosstown, A. ir R. A. Nelon ir
Goodman 5319 (OKI,, TEX). Jackson Co.: 3 mi w of Altus, Hoj)kins 1010 (OKI,).
Jefferson Co.: S of Terral, Goodman 7204 (OKL). Kingfisher Co.: 2 mi se of Dover,
Byers 71 (OKLA). Kiowa Co.: 8 mi w of Snyder, Waterfall 13101 (OKLA). Logan
Co.: Near Guthrie, Stevens 3262 1/2 (GH). Murray Co.: Davis, Arbuckle Mts.,
Emig 746 (CM). Payne Co.: 3 mi w of Stillwater, 5.5 mi s, Byler 141 (OKI \
Roger Mills Co.: Engleman 1624 (OKL). Texas Co.: Goodwell, Butler 119 (OKLA).
Woods Co.: Salt Fork River near Alva, Stevens 2876 (ILL, MIN, OKLA, POM).
Woodward Co.: Fort Supply, Potter in 1882 (GH). texas. Bailey Co.: 1.5 mi s of

MEMOIRS OF THE TORREY BOTANICAL CLUB

39

Muleshoe, Cory 27499 (TAES). Baylor Co.: Seymour, Reverchon in 1879 ( F, GH,
NY). Briscoe Co.: 3 mi s of Quitaque, Whitehouse 10010 (MICH, NY, SMU).
Carson Co.: 3 mi w of Skellytown, Stephens 17341 ( DS, KANU). Collingsworth Co.:
5.5 mi n of Wellington, Parks <b Cory 16140 (TAES). Crane Co.: 9 1/3 mi n of
Crane, Cory 37642 (TAES, TEX). Crosby Co.: 5 mi e of Crosbyton, Baker 53
(OKLA). Dickens Co.: 1 mi w, 2 mi n of Dickens, Rowell 10292 ( DS, RSA).
Dallam Co.: Just n of Dalhart, York ir Roclgers 221 (TEX, SMU, TTC). Donley Co.:
11.8 mi nw of Clarendon, Shinners 15213 (SMU). Ector Co.: 14 mi e of Monahans.
Raven ir Gregory 19235 (DS). Gaines Co.: Stateline, Tharp &■ Grimhrede 51-1013
(COLO, KANU, MIN, PENN, TAES, TEX). Garza Co.: 12 mi n of Post, McNabb
150 (OKLA). Gray Co.: 3 mi e of McLean, Munz ir Gregory 23512 (RSA). Hall
Co.: Estelline, Reverchon 3847 (MO). Hardeman Co.: Quanah, Tharp 4486
(TEX). Hartley Co.: 5.4 mi sw of Middlewater, Parks 6 - Cory 16317 (POM, TAES).
Hemphill Co.: Canadian, Bondy in 1938 (CU, DUKE, WVA). Hutchinson Co.: 10
mi e, 1 mi s of Pringle, Cutter 67 (KANU, MIN). Kent Co.: 2.9 mi sw of Jayton,
Shinners 30046 (SMU). Lipscomb Co.: Howell 42 (US). Loving Co.: Mentone
to Wink, Wamock 10722 (LL, SMU). Lubbock Co.: 16 mi e of Lubbock, Adams 94
(OKLA). Martin Co.: 1.5 mi ne of Stanton, Parks &■ Cory 13878 (TAES). Mitchell
Co.: 5 mi nw of Loraine, Pohl 4324 (SMU). Motley Co.: N of Matador, Parks &
Cory 16045 (TAES). Ochiltree Co.: 12 mi se of Perryton, 5 mi e of highway,
Wallis 4860 (OKLA, SMU). Oldham Co.: 20 mi n of Adrian, Rowell 8012 (OKLA,
TEX, TTC). Parmer Co.: 1 mi e of Bovina, Hicks in 1940 (GH). Potter Co.: N
of Amarillo, Brenkle 48175 (DAO). Randall Co.: Palo Duro Canyon, Palmer 14067
(MO, WISC). Roberts Co.: 27 mi s of Perryton, Wallis 7170 (OKLA, TEX).
Taylor Co.: 10 mi sw of Tuscola, Mahler 1321 (OKLA). Terry Co.: Wellman,
Tharp in 1941 ( GH, TEX). Ward Co.: 1 mi e of Monahans, Munz 23408 (DAO,
RSA). Wheller Co.: 5 mi e of Tela, Fogg 18790 (PENN). Wichita Co.: Red River
above Burkbumett, Tharp 574 (NY). Winkler Co.: 8 mi ne of Kermit, Irving 91
( IPN, SMU).

Gaura villosa var. arenicola Munz was separated by virtue of the glan¬
dular, not strigulose, pubescence in the inflorescences. Similar differences
in pubescence are common in many species of Onagraceae, and, by them¬
selves, are insufficient for taxonomic recognition.

6b. Gaura villosa subsp. parksii (Munz) Raven & Gregory, comb. nov.
Figs. 7, 42.

Gaura villosa Torr. var. parksii Munz, Bull. Torrey Bot. Club 65: 215.
1938. Munz, N. Amer. FI. II. 5: 192. 1965.

Plants with an understory of hispidulous pubescence which also covers
the ovaries, outside of the floral tube, and sepals more densely; branches
of the inflorescence glabrous; glandular pubescence absent. Leaves narrow¬
ly oblanceolate or oblanceolate to narrowly elliptic, shallowly sinuate-
dentate. Inflorescence delicate, well branched. Floral tube 1.5-4 mm long.
Petals 7-12 mm long. Filaments 4.5-9 mm long; anthers 2-3.5 mm long.
Style 9-16 mm long. Body of the fruit 11-19 mm long, 1-2 mm wide, the

40 A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

stripe 5-10 mm long. Self-incompatible. Gametic chromosome number,
n - 7.

Type: Open sandy places, Terrell Hill, near Kaicaster, Wilson Co.,
Texas, 24 June 1935, P. A. Munz 13334 (POM 212450. Isotvpes, GH, POM,
US).

Distribution (fig. 38): Flats and hills of red sand, local, Rio Grande
Plain of south Texas.

Specimens Examined:

United States, texas. Atascosa Co.: Poteet, Parks R1282 (MO, SMU).
Bexar Co.: 17 mi s of San Antonio, Metz 656 (NY). Dimmit Co.: 3/4 mi se of
Carrizo Springs, Parks & Cory 29407 (POM, TAES); e of Carrizo Springs, Jones
in 1931 ( BM, POM). Frio Co.: Near Pearsall, Penfound in 1936 (NO). Guadalupe
Co.: C. 10 mi s of Seguin, Webster ir Rowell 7087 (IPN). Jim Hogg Co.: 5 mi ne
of Hebbronville, Garcia 93 (OKLA). Maverick Co.: 31 mi e of Eagle Pass, Gregory
282 (RSA). Medina Co.: 3 mi s of Devine, Correll 15210 (LL); 2 3/4 mi sw of
Devine, Parks ir Cory 12737 (POM, TAES); 2 mi sw of Devine, Turner et al. 3419
(OKLA, TEX); Devine, Tharp in 1931 (TEX). Wilson Co.: Kaicaster School, Parks
6- Cory 15106 (TAES); Bexar Co. and western Wilson Co., Burr 194 (KANU). Zapata
Co.: San Ygnacio, Thaqj 3622 (TEX, US). Zavala Co.: Stony plains of the Nueces,
Wright in 1848 (GH).

7. Gaura calcicola Raven & Gregory, sp. nov. Figs. 8, 43.

Gama sinuata sensu Munz, Bull. Torrey Bot. Club 65: 218. 1938, pro
parte ; sensu Munz, N. Amer. FI. II. 5: 194. 1965, pro parte.

Herba perennis basi sublignosi multiramosaque ramis numerosis erectis
pauciramosis 0.2-1. 5 m alta sparse stigulosa. Folia linearia vel anguste
oblanceolata, 1-12 cm longa, 0.1-1 cm lata, basalia lyrata vel oblanceolata
conspicue petiolata, superiora subsessilia. Inflorescentia 14-112 cm longa,
virgata, bracteis lanceolatis vel ovatis, 1-3 mm longis, 0.5-1 mm latis.
Flores zygomorphi, vespertini. Tubus floris 3-9 mm longus, intus lanatus.
Sepala 6-12 mm longa, 1-2 mm lata. Petala 7-10.5 mm longa, 3-6.5 mm
lata. Filamenta 3-7 mm longa; antherae 2.5-5 mm longae. Stylus 9.5-19
mm longus, basin versus lanatus. Fructus clavatus, parte crassiore 7-12 mm
longo, 1. 5-2.5 mm crasso, stipite 2-5 mm longo; semina (2-)3-4(-5),
L. 5-2.5 mm longa, 0.75-1.25 mm lata, obovoidea, straminea vel pallide
castanea. Numerus chromosomaticus gameticus, n = 7.

Clumped perennial herb, from a heavy, twisted, woody rootstock; stems
branching below and just above the surface of the ground to form at
maturity a clumped leafy plant from which long, wand-like inflorescences
arise; stems also branching in a whorl within the leafy portion below the
inflorescences, 2-15 dm tall. Plants subglabrous or with more or less dense,
erect hairs 1.5-2 mm long in leafy portion, in either case the plants also

MEMOIRS OF THE TORREY BOTANICAL CLUB

41

42

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

sparsely strigulose on the lower portions and sometimes on the leaves,
and more or less heavily strigulose on the pedicels, outside of the floral
tube and sepals, and ovary; inflorescence subglabrous or occasionally
sparsely strigulose, rarely glandular-pubescent. Leaves linear to narrowly
oblanceolate, 1-12 cm long, 0.1-1 cm wide, slightly to markedly sinuate-
dentate. Inflorescence 14-112 cm long, the naked lower portion about as
long as the portion with flowers and fruits above; bracts lanceolate to
ovate, 1-3 mm long, 0.5-1 mm wide. Flowers with the stamens in lower
half. Floral tube 3-9 mm long. Sepals 6-12 mm long, 1-2 mm wide.
Petals 7-10.5 mm long, 2-6.5 mm wide. Filaments 3-7 mm long; anthers

2.5- 5 mm long. Style 9.5-19 mm long. Body of the fruit 7-12 mm long,

1.5- 2.5 mm thick, the stipe 2-5 mm long. Seeds (2-)3-4(-5), 1.5-2. 5 mm
long, 0.75-1.25 mm thick, light brown or reddish brown. Self-incompatible.
Gametic chromosome number, n = 7.

Type: Limey, rocky soil in a fairly rich mesquite range, 15 mi e of Big
Lake on U.S. Hwy. 67, Irion Co., Texas, 3 June 1964, P. II. Raven 6- D. P.
Gregory 19242 (DS 574029. Isotypes, BM, GH, US).

Distribution (fig. 47): Dry limestone, gypsum, or caliche slopes and
banks in the southern Trans-Pecos and Edwards Plateau regions of Texas,
southward in Mexico into the mountains of eastern Durango and Coahuila,
Nuevo Leon, and western Tamaulipas; northeastern Durango.

Representative Specimens Examined:

Mexico, coahuila. 23 mi se of Monterrey, Barkley iz Warnock 14849 (DUKE,
F, GH, NY, TEX, US); Ojo Caliente, Barkley ir Gutierrez 17M114 ( F, MICH, TEX);

39 mi n\v of Muzquiz, Gould 11156 (DS); near Saltillo, Hinton et al. 1663/ (FI,
GH, US); Muzquiz, Marsh 2131 ( F, GH, OKLA, SMU, TEX); Sierra de Santa Rosa,
s of Muzquiz, Marsh 1229 ( F, GH, SMU, TEX); Palo del Diamente near Saltillo,
Munz 15026 (GH, MO, POM); Saltillo, Palmer 131 ( C, F, G, GH, MO, NY, US),
233 (C, GH, NY, POM, US); Rio Grande valley near Diaz, Pringle 8320 (BM, CM,
F, G, GH, IPN, ISC, LE, M, MEXU, MICH, MIN, MO, MSG, ND, PH, POM, US);

40 km nw of Hacienda de la Encantada, Stewart 1621 (GH); e slope of Sierra de
Puerto Santa Ana, Wyrnl ir Mueller 249 (GH, K, MO, NY, US), duranco. 95 mi
ne of Durango, Waterfall 6 Wallis 13221 (IA, OKL, SMU). nuevo leon. Galeana,
Chase 7722 ( F, GH, ILL, MO, NCU, NY); Mpio. Galeana, Chase 7647 (F); 5 km
s of Sabinas Hidalgo, T. C. 6 E. M. Fry 2401 ( DS, C,H, IND, MO, NY, RSA,
SMU, US); 11 mi ne of Sabinas Hidalgo, 5 mi se of Vallecillos, John ton 6 Graham
4608 (IPN, MICH, TEX); 15 mi sw of Galeana, C. 11. 6 M. T. Mueller 1040 (A, F,
MICH, TEX, NY); 67 km nw of San Roberto, Roe et al. 51 ( DS, WISC); Hacienda
Pablillo, Galeana, Taylor 23 ( DS, F, MEXU, MICH, MO, OKLA, RSA, SMU, TEX);
35 mi s of Monterrey, White 1635 (GH, MICH); La Escondida, Roybal 276 (US).
tamaulipas. Cerro del Chino, vicinity of San Miguel, Sierra de San Carlos, Barthit
10717 (MEXU, MICH, US).

United States, texas. Brewster Go.: Hess Hill, Glass Mts., Warnock W303
(DS, FSU, GH, OKLA, TEX, WVA); s of Marathon, Steiger 586 (NY); Pena Blanca,

MEMOIRS OF THE TORREY BOTANICAL CLUB

43

Sperry T852 (US); Haley Ranch, Parks ir Cory 9 275 (TAES); Reagan Canyon,
Cory 2148 (POM); 4 mi w of Alpine, Davis Mts. Warnock 5962 ( LL, SMU, TEX).
Crockett Co.: Ozona, Jones 25876 ( BM, CU, DS, MO, POM). Edwards Co.: Rock
Springs, Reed ir Cory 33847 (TAES). Irion Co.: Mertzon, Palmer 12415 ( BM, GH,
MO, NT US). Jeff Davis Co.: 3 mi n of Fort Davis, Munz 6- Gregory 23383 (DAO,
RSA). Kendall Co.: Spanish Pass, Clemens ( BH, MIN, POM). Kerr Co.: Kerrville,
Heller 1692 (BM, C, CU, F, GH, ILL, ISC, LE, MICH, MIN, MSC, NY, PH, POM,
US). Kinney Co.: 4 mi ne of Bracketville, Strouther 267 (COLO, DS, IPN, SMU,
TEX). Maverick Co.: 13 mi n of Eagle Pass, Gould 11310 (DS). McCulloch Co.:
4.5 mi se of Brady, Munz ix Gregory 23430 (DAO, RSA). Menard Co.: Beede 7116
(IND). Pecos Co.: Tunis Creek, Tharp 43-737 (F, MO, NY, OKL, PH, POM, TEX).
Presidio Co.: 18 mi sw of Marfa, Hinckley 3525 (RSA). Sterling Co.: Tharp 3620
(TEX, US). Ten ell Co.: 13 mi s of Sheffield, Webs ter 417 (TEX). Tom Green Co.:
San Angelo, Brady in 1899 (TEX); 7 2/3 mi s of Christoval, Cory 50571 ( DS, GH,
NY, SMU, TEX, US). Uvalde Co.: 16 mi n of Uvalde, T. 6- L. Mosquin 5620
(DAO). Val Verde Co.: 13 3/4 mi ne of Comstock, Parks 6- Cory 31693 (TAES).

Goura calcicola has heretofore been grouped with G. sinuata, a tetra-
ploid (n = 14) species that is strongly rhizomatous and lower in stature.
The large clumps of G. calcicola, with their numerous virgate branches,
are often a conspicuous feature of limestone-covered hillsides in western
Texas; hence the specific epithet. In contrast, G. sinuata, a noxious weed,
often forms patches of considerable size, spreading by means of rhizomes.
In bud, the tips of the inflorescences in G. calcicola are very slender, with
small, well-spaced buds; those of G. sinuata are truncate, with numerous
well-developed buds crowded right at the apex. Despite these fundamental
differences, the two species are often difficult to distinguish as herbarium
specimens. Gaura mckeheyae is likewise similar, but may be distinguished
from both by its more deeply divided leaves and conspicuous villous pu¬
bescence; scattered villous pubescence does, however, occur in G. calcicola.

8. Gaura filipes Spach, Nouv. Ann. Mus. Mist. Nat. 4: 379. 1835. Munz.
Bull. Torrey Bot. Club 65: 216. 1938. N. Amer. FI. II. 5: 193. 1965.
Figs. 9, 44.

Gaura michauxii Spach, Nouv. Ann. Mus. Paris 4: 379. 1835. Type:
Cherokee Territory ( = eastern Tennessee or an adjacent state), Palisot
de Beauvois (G. Photographs, DS. POM).

Gaura filipes Spach 0 major Torr. & Gray, FI. N. Amer. 1: 517. 1840.
Lectotype: Barrens of Kentucky (prairies in w part of the state), 1840,
C. W. Short (NY. Isotype, PH). Munz, Bull. Torrey Bot. Club 65: 217.
1938; N. Amer. FI. 5: 193. 1965.

Gaura filipes Spach var. ti/pica Munz, Bull. Torrey Bot. Club 65- 216
1938.

44

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Clumped perennial herb, from a heavy, twisted, woody rootstock; stems
branching below and just above the surface of the ground to form at
maturity a clumped leafy plant from which long, wand-like inflorescences
arise; stems also branching in a whorl within the leafy portion below the
inflorescence, 6-15(-20) dm tall. Plants densely strigulose on ovary, out¬
side of floral tube, and sepals, sometimes also strigulose on the branches
of the inflorescence and the stems, but usually the lower inflorescence
glabrous; in some plants the leafy portion densely covered with erect hairs
1-2 mm long, in others strigulose with the very base of the inflorescence
glandular-pubescent; leaves in plants with the leafy portion with long,
erect hairs strigulose, especially along the veins and margins; leaves in
plants with the stems of the leafy portion strigulose subglabrous. Leaves
linear or narrowly lanceolate to narrowly oblanceolate, 1-9 cm long, 0.1-
1.3 cm wide, slightly to coarsely sinuate-dentate. Inflorescence 50-115 cm
long, often well branched; bracts linear, 1-2 mm long, 0.2-0.5 mm wide.
Flowers with stamens well distributed around their diameter. Floral tube
2.5-6 mm long. Sepals 5-12.5 mm long, 1-2.1 mm wide. Petals 5-10 mm
long, 1. 6-4.5 mm wide. Filaments 3-8.5 mm long; anthers 1.4-4 mm long.
Styles 8.5-19 mm long. Body of the fruit 5-10 mm long, 1.5-2 mm thick,
the stipe 0.5-4. 5 mm long. Seeds 1-2, 1. 5-3.5 mm long, 1-1.5 mm thick,
yellowish to reddish brown. Self-incompatible. Gametic chromosome num¬
ber, n — 7.

Lectotype: Jacksonville, near present-day Sheridan, Washington Parish,
Louisiana, autumn 1832, Thomas Drummond (P. Isolectotypes, CAMB,
G, GH, K); Munz, Bull. Torrey Bot. Club 65; 216. 1938, who incorrectly
assumes that Drummond’s “Jacksonville” was in Florida, as does McKelvey
( Botanical Exploration of the Trans-Mississippi West 1790-1850, p. 494,
1955).

Distribution (fig- 47): Sandy hills and flats, often in open woods,
southernmost Illinois and southern Indiana to South Carolina, northern
Florida, southeastern Mississippi, and Washington Parish, Louisiana. In
Florida, we have seen no collections from east of the Oehlockonee River
(near Tallahassee) made since 1895. A specimen said to have been col¬
lected at Tampa Bay by Asa Gray in 1840 (NY) is probably mis-labelled,
as the species is otherwise unknown from south of Alachua County. 1* lowers
August to October.

Representative Specimens Examined:

United States, alahama. Baldwin Co.: 2 mi e of Foley, Godfrey 65958 ( DS,
FSU). Barbour Co.: Spring Hill, Grave 1043 (MO). Bibb Co.: 6.5 mi e-ne of
Centerville, Jones 1673 (NCU). Blount Co.: Banks of Mulberry River, Expert in
1898 (MO). Clay Co.: Earle 997 (NY). Henry Co.: Near Headland, Wiegand 6

MEMOIRS OF THE TORREY BOTANICAL CLUB

45

Maiming 2165 (CU). Lawrence Co.: Decatur, Smith in 1884 NY, US). Lee Co.:
Auburn, Earle 6- Baker in 1898 (F). Madison Co.: Near Huntsville, Canhy 48 (MO,
US). Mobile Co.: Mobile, Mohr in 1878 (GH, NY, PH). Pike Co.: Springhill, Mohr
in 1886 (US). Shelby Co.: Calera, Everts in 1878 (CINC, NY). Florida. Alachua
Co : Hitchcock 672 (F). Bay Co.: Vicinity of Youngstown, Goclfreij 69327 (FSU).
Calhoun Co.: Dead Lakes, Godfrey 64611 ( DS, FSU). Columbia Co.: Lake City,
Xash 2498 (PH, US). Cadsden Co.: River Junction, Curtiss 6003 (CU, DS, F, FLAS,
CH, G, GA, ILL, ISC, KANU, LE, MIN, MO, NCU, PAC, PH, US); Aspalaga!
Biltmore Herb. ( Curtis ) 3929c (C, GH, MIN, NY, US). Jackson Co.: 2 mi n of
Sneads, Godfrey 67509a ( DS, FSU). Okaloosa Co.: C. 8.5 mi ne of Niceville,
Ward 5975 (DS, FLAS). Santa Rosa Co.: 4 mi n of Milton, Davis 090362-5
(LAF). Walton Co.: N of De Funiak Springs, West 6 • Arnold in 1954 (US). Wash¬
ington Co.: S of Chipley, Hood 2860 (FLAS). Georgia. Baker Co.: 2 mi sw of

^?n^°dfrey 67492 (DS’ FSU)- Barlow Co-: 61 mi e 56° of Adairsville, Duncan
GH’ M°’ SMU’ US)- Brantley Co.: 10 mi s-se of Waycross, Kuns 93
C Catoosa Co.: 1.4 mi e 18° of Fort Oglethorpe, Harris 6- Duncan 13022
(GH, NSC, US). Chattahoochee Co.: E from Columbus, Donnell Smith in 1883
(N\). Clarke Co.: Athens, Miller & Maguire 1055 (BH, CU). Cobb Co.: Archean
\\ ilson 34 (GH, MO, NY, US). Emanuel Co.: 9 mi e of Adrian, Wilson 2911 (FSU*
.A, IA, NSC). Floyd Co.: Rome, Chapman (MO, US). Grady Co.: 5 mi e of
Climax, Cronquist 5471 ( GA, NY). Gwinette Co.: Yellow River, Eggert in 1897
(CM, MO, N\, US). Habersham Co.: Currahee Mt., Small in 1894 (F, MICH,
MIN, MO, MSC, NY, PH). Jasper Co.: Montieello, Porter 5 (PH). Jefferson Co.:
Calhura, Hopkins 25 (NY). Liberty Co.: 4 mi sw of Hinesville, Wiegand & Manning
“ 164 (CU). McDuffie Co.: 6 mi ne of Thomson, Duncan 10650 ( GA MO) Rich¬
mond Co.: Augusta, Cuthbert in 1897 (FLAS). Screven Co.: Southeastemmost part,
Eyles 7553 (GA). Stephens Co.: 1 mi n-nw of summit of Gurrahee Mt. Duncan
11774 (FLAS, GA, IA, IND, ISC, MO, SMU, TENN). Sumter Co.: Ca. 2 mi s of
Leslie, Duncan 1 841 (GA). Thomas Co.: Thomasville, Harrison in 1904 (GH).
Toombs Co.: 7.5 mi e 39° s of Lyons, Hardin 6- Duncan 14514 (GA). Tromp Co •

3 mi ne of Hogansville, Wiegand & Manning 2166 (GH). Illinois. Hardin Co.:
Saline Cieek, W. D. in 1916 (Ill.) Indiana. Pt. Commerce, 1846, A. Wood (NY).
Daviess Co.: 4 mi n of Washington, Deam 47902 (GH, IND US)- 15 mi sw of
Plainville, Kriebel 5804 (GH, NCU, SMU). Kentucky. Barren Co.: Diamond Cavern
Braun 3639 (Braun herb.); e of Glasgow Junction, O. E. 6- G. K. Jennings (CM)
Edmondson Co.: Mammoth Cave Park National Forest, Aides 7550 (ILL) Hart
Co.: M unford ville, Rhoades in 1925 (WISC). Nelson Co.: Balltown, Braun 3212
(Braun herb.). Pulaski Co.: Alpine, McFarland <& James 28 ( BH F GA IND
MIN, MO, NY, PENN, PH, SMU, TENN, WISC, WVA). Wayne ’ Co.:’ N of
Montieello, Braun 1030 (Braun herb.). Louisiana. Washington Parish: Ca. 5 mi w

Thieret 32676 (DS)- Mississippi. Forrest Co.: Hattiesburg, Jones 3557
(FSU). Harrison Co.: Long Beach, Joor in 1891 (MO). Jackson Co., Ocean Springs
Tracy in 1889 (MSC). Lauderdale Co.: Meridian, Schuchert in 1896 (US), south
Carolina. Aiken Co.: 4.3 mi s of Montmorenci, Aides 6- Crutchford 55578 (NCU)
Allendale Co.: 1.6 mi nw of Fairfax, Aides ir Bell 15887 (NCU). Bamberg Co.: 0.2
mi n of junction of County Hwy. 27 and State Hwy. 64, Aides 37647 (CM, NCU).
Beaufort Co.: 0.5 mi sw of New River Crossing, Raven 18707a (DS). Calhoun
o.. Junction of County Hwy. 157 with County Hwy. 72, Aides 35341 (NCU, SMU).
Colleton Co.: Junction of County Hwy. 35 with State Hwy. 61, Aides 6 Bell 17936

46

A REVISION OF THE GENUS GAURA (ONAGRACEAE)

(GH, NCY). Dorchester Co.: 1 mi w-sw of Reevesville, Ahles <b Leisner 31924
(NCU). Edgefield Co.: 1.2 mi sw of Trenton, Radford 30158 GA, NCU). Jasper
Co.: 3 mi e of Grahamville, Ahles 6- Bell 18072 (NCU). Kershaw Co.: Junction
Hwys. 602 and 12, Duke 2973 (NCU). Lexington Co.: 25 mi w of Columbia, Munz
13359 (POM). Oconee Co.: Hwy. 123 at Tugaloo River, Freeman 58507 (NCU).
Orangeburg Co.: 2.3 mi sw of Orangeburg, Leisner 6- Ahles 31806 (DAO, NCU).
Richland Co.: Columbia, Crawford iti 1890 (F, GH). Saluda Co.: 2 mi s of Ridge
Spring, Radford 30424 ( FSU, NCU). Sumter Co.: 2.5 mi w of Pinewood, Radford
29682 (NCU). Williamsburg Co.: 3.5 mi sw of Mouson, Ravlford 31248 (NCU).
Tennessee. Bledsoe Co.: Mountain e of Pikeville, llarger 7721 (TENN). Bradley Co.:
Chattanooga, Engelmann in 1876 (MO). Decatur Co.: Ames in 1855 (MICH). Frank¬
lin Co.: Cowan, Biltmore Herb. 3929a (GH, NY). Hamilton Co.: Lookout Mt.,
Biltmore Herb. 3929b (CH, NY, US). Lewis Co.: Hohenwald, King 414 (US).
Loudon Co.: Sharp et al. 1948-8 (TENN). Monroe Co.: 1.5 mi n of Vernon, Raven
20449 (DS). Rhea Co.: Waldens Ridge, Shanks et al. 8908 ( RSA, TENN). Ruther¬
ford Co.: 6 mi e-se of Murfreesboro, DeSelm 718 (TENN). White Co.: Caney Fork
near Webbs Camp, Sharp et al. 5030 ( KY, TENN).

The characters used by Munz (Bull. Torrey Bot. Club 65: 216, 217.
1938) to divide Gaura filipes into two varieties vary within populations
and are not geographically correlated; furthermore, as pointed out by E.
Lucy Braun (Annotated Catalogue of the Spermatophytes of Kentucky
99, 100. 1943), flower size varies considerably from year to year.

9. Gaura mckelveyae (Munz) Raven & Gregory, comb. nov. Figs. 10, 45.

Gaura villosa Torr. var. mckelveyi Munz, Bull. Torrey Bot. Club 65:
214. 1938; N. Amer. Fl. II. 5: i92. 1965.

Clumped perennial herb, from a twisted, woody rootstock; stems branch¬
ing below or just above the surface of the ground; stems well branched,
2-6 dm tall. Stems more or less densely covered with long, erect hairs 2-3.5
mm long, also strigulose; leaves, sepals, ovary, and outside of floral tube
strigulose; inflorescence glabrous or often with glandular pubescence from
the base of the peduncle upward for 2-3 cm or more, very rarely with a
few long or strigulose hairs. Leaves exceptionally variable in size and
margins, elliptic to more commonly narrowly oblanceolate, 1-6.5 cm long,
0.1-1. 5 cm wide, deeply sinuate-dentate or less markedly so, often undulate,
the basal ones up to 17 cm long and 2 cm wide. Inflorescence 10-75 cm
long, well branched; bracts lanceolate to narrowly ovate, 1-5 mm long,
0.5-1 mm wide. Flowers with stamens in the lower half. Floral tube 2-3.5
mm long. Sepals 6-12 mm long, 1-2 mm wide. Petals 7-10.5 mm long,
2-5.5 mm wide. Filaments 5-9 mm long, lanate at the very base; anthers
2-4 mm long. Style 9-16 mm long. Body of the fruit 8-19 mm long, 1.5-2
mm thick, the stipe 3-9 mm long. Seeds ( 1— ) 2—4, 2-3 mm long, ca. 1 mm

MEMOIRS OF THE TORREY BOTANICAL CLUB

47

thick, yellowish to reddish brown. Self-incompatible. Gametic chromosome
number, n = 7.

Type: Near Mathis, San Patricio Co., Texas, 5 April 1931, Susan Delano
McKelvey 1718 (GH. Isotype, POM).

Distribution ( fig. 47 ) : Sandy flats, sand hills, and waste places in the
Rio Grande Plain of south Texas, northwesternmost Tamaulipas, and ad¬
jacent Nuevo Leon. Flowers, March to June.

Representative Specimens Examined:

Mexico, nuevo leon. 50 mi s of Laredo on Hwy. 85, Hess ir Hall 640 (OKL);
7 mi ne of Vallecillo, Rodriguez 60 (SMU, TAES). tamaulipas. Nuevo Laredo,
Arizmendi 68 (DS, LL); 24 km s of Nuevo Laredo, Dominguez 6- AlcCart 8291
(TEX); 50 km s of Nuevo Laredo on Hwy. 85, Dominguez 6- AlcCart 8234
(COLO, DS, MSC).

United States. Texas. Aransas Co.: 4.6 mi ne of Rockport, AlcCart 5576 ( DS,
SMU). Bee Co.: 4 mi se of Normanna, Cortj 54120 ( GA, LL, SMU). Brooks Co.:
15 mi e of Hebbronville, Chavez et al. 54 (SMU). Cameron Co.: National Wildlife
Refuge, Traverse 1128 (F, SMU, US). De Witt Co.: Cuero, Howell 323 (US).
Dimmit Co.: Carrizo Spring, Cuellar 41 (LL). Duval Co.: San Diego, Croft in 1885
(MICH, NY). Hidalgo Co.: Near San Manuel, C. L. ir A. A. Lundell 10834 (MICH,
POM, SMU). Jim Hogg Co.: 5 mi ne of Hebbronville, Gutierrez Garcia 93 (SMU,
TEX). Jim Wells Co.: 5.5 mi e of Alice, Skinners 24109 (SMU). Karnes Co.: 5
mi se of Choate, Johnson 1554 (RSA, SMU). Kennedy Co.: Sand hills below Sarito,
Tharp in 1928 (TEX). Kleberg Co.: Santa Gertrudis Division of King Ranch,
Johnston 53224.19 ( LL, MSC, SMU, TEX); Kingville, Tracy 9275 (CU, DUKE, F,
G, GH, MIN, MO, MSC, NY, PENN, TAES, TEX, US, WISC). La Salle Co.: Encinal,
Barrera 13 (DS, DUKE, SMU, TEX); 5 mi ne of Encinal, Canter 70 (IKLA, SMU,
TAES, TEX). Nueces Co.: Corpus Christi, Tracy 9273 ( F, G, GH, MIN, MO, MSC,
NY, TAES, TEX, US). Refugio Co.: Benke 5401 ( F, POM). San Patricio Co.:
Welder Wildlife Foundation n of Sinton, Traverse 1202 ( F, MO, SMU, TEX, US).
Starr Co.: 3 mi w of Sullivan City, C. L. 6- A. A. Lundell 9889 ( LL, MICH, POM,
US). Webb Co.: 6 mi n of Bruni, Chaves et al. 32 ( LL, OKLA, TTC); 13 mi n of
Webb, Correll 20752 (LL); Laredo, Dickey 134 (OKLA). Willacy Co.: Redfish Bay,
C. L. ir A. A. Lundell 8769 (DS, GH, LL, MICH, NY, POM, SMU). Zapata Co.:
4 mi e of Zapata, Araiza 52 (COLO, DS, DUKE, NCU, SMU); 6 mi s of San Ignacio,
Garcia 15 (DS, DUKE, SMU).

A better understanding of this entity has made it necessary to accord
it specific rank. It is coordinate in status with the other diploid species of
section Stipogaura.

10. Gaura sinuata Nutt, ex Ser., in DC., Prod. 3: 44. 1828. Munz, Bull.
Torrey Bot. Club 65: 218. 1938, pro parte ; Munz, N. Amer. FI. II. 5:
194. 1965, pro parte. Figs. 11, 46.

Aggressively rhizomatous perennial herb, forming extensive mats with
well branched stems 2-6 dm tall. Plants subglabrous or sparsely pubescent.

48

A REVISION OF THE GENUS GAURA (ONACRACEAE)

the stems strigulose and with long spreading hairs; ovary, outside of floral
tube, and sepals strigulose, the leaves densely strigulose in relatively hairy
plants. Leaves linear to narrowly oblanceolate, 1-11 cm long, 0.1-2 cm
wide, sparsely sinuate-dentate, rarely subentire, often undulate. Inflorescence
10—102 cm long, simple or branched; bracts lanceolate to narrowly ovate,
1-5 mm long, 0.5-2 mm wide. Flowers with stamens in lower half. Floral
tube 2.5-5 mm long. Sepals 7-14 mm long, 1.25-2.5 mm wide. Petals 7-14.5
mm long, 3-/ mm wide. Filaments 5-11 mm long, lanate at the very base;
anthers 3—5 mm long. Style 12—18.5 mm long. Body of the fruit 8—15 mm
long, 1. 5-3.5 mm thick, the stipe 2-8 mm long. Seeds (l-)2-4, 2-3 mm
long, 1-1.5 mm thick, light to reddish brown. Self-incompatible. Gametic
chromosome number, n = 14.

Type: Near the Red River, in what is now southeastern Oklahoma, 16
May to 21 June 1819, Thomas Nuttall (G-DC. Isotvpes, BVI CAMP K
PH).

Distribution (fig. 47): Flats and washes in light sandy loam, central
and southern Oklahoma and southward in the Rolling Plains of Texas,
whence eastward and southward throughout the eastern half of the State
below the Edwards Plateau, but especially common in the Blackland
Prairies and probably introduced following agricultural clearing in the East
Texas Timberlands and on the Rio Grande Plain; conceivably native in
Franklin County, Arkansas, but widely introduced elsewhere in the light
agricultural soils of Alabama, Arkansas, California, Florida, Georgia, Mis¬
souri ( Muehlenbach, Ann. Missouri Bot. Gard. 56: 168. 1969), New York
(House, N. Y. State Mus. Bull. 254: 57-59. 1924), Italy, and South Africa.
A noxious weed, but fortunately limited by its self-incompatibility. The
species was probably originally native to the prairies and rolling plains of
south-central Oklahoma and central and north-central Texas.

Representative Specimens Examined:

Italy. Viareggio, 15 Oct. 1961, Montelucci 8034 (FI), “copiosamente naturalissata
su tutte le dune della localita . . . dune alte presso le lame elevante.”

South Africa. Cape Province. Port Elizabeth District: Van Sladens Pass, 500
ft, 4 Jan. 1936, Theron 1837 (M), “a small patch of 5 square yards, spreading by
rhizomes ... no mature fruit seen,” 1 Dec. 1965, Wells 3192 (G, M).

United States. Alabama. Lee Co.: Auburn, Jones 614 (NCU). Baldwin Co.:
Ft. Morgan fortifications, litis et al. 21478 (DS). Tuscaloosa Co.: University Campus,
Harper 3192 (G1I, MO, NY, PI I, US, WISC). Arkansas. Franklin Co.: Valley of
the Arkansas River s of Ozark, ca. 500 ft elevation, Moore 410105 ( F, WISC). Lonoke,
Co.: Rock Island Railroad right-of-way, near ballast, P.O. Carlisle, Demaree 55604
(DS). Pulaski Co.: Along railway embankment, near Little Rock, Palmer 30332 (MO,
UMO). California. Alameda Co.: From an infestation on fallow land a few mi ne
of Livermore, Bellue in 1949 (RSA). Fresno Co.: Fresno, Quibell in 1960 ( RSA).
Kern Co.: Tehachapi Mts., Hutchinson 3609 (DS). Los Angeles Co.: Calleano

MEMOIRS OF THE TORREY BOTANICAL CLUB

49

Orchard near Walnut, Johnson in 1939 (POM). Riverside Co.: Near Temecula,
Rowntree in 1937 ( I EX). Sacramento Co.: 8.5 mi ne of Sacramento, Fuller 2413
(DS, RSA, US). San Bernardino Co.: Below Mill Creek Power Station on road to
Yucaipa, Craig et al. 325 (POM, US). San Diego Co.: Near Ramona, Roos 513 (RSA).
San Mateo Co.: Junction of Los Trancos and San Francisquito creeks, Abrams 13687
(DS, N\, POM). Santa Barbara Co.: E of Carpenteria, Pollard in 1957 (CAS, DAO,
MIN, TEX). Ventura Co.: 1.5 mi ne of Somis, Johnson in 1940 (POM). Florida.’
Alachua Co.: Gainesville, Beckner 1246 ( DS, FLAS). Marion Co.: McIntosh, 1936,
Gist 112 (US). Georgia. Dougherty Co.: 4 mi s of Albany, Thorne 3199 (CU, Ga’
GH, IA). Greene Co.: 1.5 mi n of Greensboro, Pyron ir McVaugh 1647 (GA). Jack-
son Co.: 2.7 mi se of Jefferson, Duncan 18550 (GA, KY, LAF, NCU, NSC, SMU).
Macon Co.: 4 mi s of Reynolds, Pyron ir McVaugh 2807 (GA). Schley Co.: Ellaville,
Sargent 7431 ( GA, NSC). Missouri. St. Louis Co.: Baden Freight Yard, Muhlenbach
2760 (DS). Oklahoma. Atoka Co.: 15 mi s of Atoka, Hopkins ir Nelson 1133
(MO, IKL, RSA). Blaine Co.: 8 mi sw of Okeene, Cutter in 1951 (OKL). Bryan
Co.: Vicinity of Durant, Blain 108 ( US). Cleveland Co.: Norman, Crabb 96 (PENN).
Custer Co.: 6 mi sw of Butler, Mericle 1628 (OKL). Grady Co.: Chiekasha Prairie,
McGuire in 1921 (OKL). Harmon Co.: 2.6 mi e of Gould, Shinners 31403 (SMU)!
Hughes Co.: 6 mi e of Holdenville, Hopkins 1621 (BH, IND, OKL, OKLA, POM).
Jefferson Co.: Ringling, Pryor 258 (OKL). Kay Co.: Blackwell, Byler 327 (OKLA).
Kiowa Co.: N of Snyder, Eggert in 1903 (MIN). Marshall Co.: 1.5 mi e of Lebanon,
Goodman 6500 (OKL, OKLA, RSA). Murray Co.: 8 mi n of Springs, Cory 59093
(OKLA, SMU). Oklahoma Co.: Oklahoma City, Waterfall 2801 (OKLA). Payne
Co.: 914 mi w, 1.8 mi n of Stillwater, Mueller 1901 (OKLA). Pittsburgh Co • 7 mi
w of McAlester, Hopkins 1635 (OKL, POM). Tillman Co.: 4 mi n Gandfield,
Stephens 20353 ( DS, KANU). Washita Co.: 3 1/3 mi s of Rocky, Waterfall 2019
(GH, OKL). south CAROLINA. Charleston Co.: Navy Yards, Robinson 196 (GH).
Edgefield Co.: 2.5 mi s of Trenton, Radford 22559 (DAO, FSU, GA, GH, NCU,
USE, VDB). Lexington Co.: Leesville, Radford 27169 (NCU). Richland Co.:’
Ridgewood, near Columbia, Philson in 1936 ( GA, NCU). Union Co.: 4 mi n of
Union, Freeman 56146 (NCU). Texas. Anderson Co.: Palestine, Tharp 863 (US)
Bexas Co.: Bandera Road, Schulz 303 (US). Borden Co.: 19 mi nw of Gail, Parks
C Cory 30112 (TAES). Brazos Co.: College Station, Reeves 819 (POM). Cameron
Co.: Gunnery Range Unit at Stover Point, Traverse 1128 (MO, TEX) Childress
Co.: C.H.S. School, Biology Class 15 (TEX). Collin Co.: 3.7 mi’ e of Farmersville,
Simmers 14317 (SMU). Colorado Co.: Perkins 6- Hall 2464 (CU). Coleman Co.:
1 mi se of Santa Anna, Shinners 26394 (SMU). Cooke Co.: 7.5 mi n of Gainsville,
Shinners 12433 (SMU). Duval Co.: San Diego, Croft in 1886 (NY). Erath Co.:
Dublin, Maxwell 46 (F). Fayette Co.: Muldoon area, Ripple 51-588 (TEX) Frio
Co.: Wolcott & Barkley 16T257 (ILL, RSA, TEX). Garza Co.: 1 mi s, 1 mi w of
Post, Rowell 10164 (OKLA). Goliad Co.: Goliad, Williams 35 (PH, TEX). Gonzales
Co.: 3/4 mi w of Monthalia, Parks 6 Cory 5912 (POM, TAES). Hays Co.: 5 mi w
of San Marcos, Gregory 418 (RSA). Jack Co.: 2 mi w of Jacksboro, Whitehouse
15305 (MICH, SMU). Jackson Co.: Edna, Drushel 2435 (ILL, MO). Lamar Co.:
Paris, McMullen in 1927 (TEX). Lampasas Co.: 5 mi s of Evant, Mayler 1239

(OKLA). La Salle Co.: Carrizo Spring to Cotulla, - in 1898 (TEX). Mitchell

Co.: 3 mi n-nw of Colorado City, Pohl 5020 (PENN, SMU). Montgomery Co-
Near Harmons Creek, Dixon 448 ( F, GH, NY, WISC). Nacogdoches Co.: Nacog¬
doches, Banks 3180 (DS). Navarro Co.: 8 mi se of Corsicana, Shinners 14427 (SMU).

50

A REVISION OF THE GENUS GAURA (ONAGRACEAE)

Orange Co.: Curtis School, Wild Wood Club 8 (TEX). Parker Co.: 8 mi se of
Weatherford, Timmons in 1939 (SMU). Rains Co.: 2.5 mi nw of Point, Skinners
14342 (SMU). San Patricio Co.: 3 mi n of Sinton, Willige s 328 (TEX). Shackleford
Co.: 5 mi w of Albany, McCoy 3762 (DAO, NCU, OKL, USL). Tarrant Co.: Fort
Worth, A. Ruth 37 (GH, MIN, US). Taylor Co.: Near Abilene, Tolstead 7183
(ILL, OKLA, SMU). Uvalde Co.: Utopia, Kincaid ir Johnston 54638 (TEX).
Victoria Co.: Victoria, Eggert in 1900 (MIN). Waller Co.: Hempstead, Hall 214
(F, C.H, ILL, MO). Wichita Co.: Division Lake, Parks ir Cory 13221 (TAES).
Williamson Co.: Near Taylor, Innes 844 (GH). Wise Co.: 4.1 mi nw of Bridgeport,
Hennen 558 (SMU).

SECTION VI. XENOGAURA
Gaura sect. Xenogaura Raven & Gregory, sect. nov.

Herba perennis rhizomatosa, caulibus strigulosis interdum etiam villosis.
Inflorescentiae subnutantes. Flores valde zygomorphi, vespertini. Fructus
4-angulatus, parte superiore crassiore pyrimidali basi turgida, demum acu-
tissime constricta, parte basali tereti.

Rhizomatous perennial herb, the stems strignlose and sometimes also
villous. Inflorescence slightly nodding at the tip. Flowers strongly zygo-
morphic, the stamens in the lower quarter of the flower; flowers opening
near sunset and withering the next morning. Fruit not reflexing, 4-angled,
with 4 distinct ridges and 4 furrows alternating at the top of the pyramidal
upper half, the base of the pyramidal portion distinctly bulging, then im¬
mediately and sharply constricted, the basal portion terete and about a
quarter of the diameter of the swollen base of the upper portion. Self¬
incompatible. Gametic chromosome number, n = 14.

Type Species: Gaura drummondii (Spach) T. & G.

The single species of this section, Gaura drummondii, is consistently
tetraploid. Its morphological characteristics and geographical and eco¬
logical intermediacy suggest strongly that it is of polyploid derivation be¬
tween G. coccinea and a species of section Stipogaura somewhat resembling
G. mckelveijae. Because it is impossible to confirm this hypothesis at present,
we have given the section a name based on the Greek work zivos, stranger,
indicating the uncertain origin of the species concerned.

11. Gaura drummondii (Spach) T. & G., FI. N. Amer. 1: 519. 1838. Figs.
12, 36.

Schizocartja drummondii Spach, Nouv. Ann. Mus. Hist. Nat. 4: 382.
1835. Type: Austin, Travis Co., Texas. 1833-1834, Thomas Drummond
III. 36 (G. Isotypes, BM, CAMB, GH, K, NY, P).

Scldzocan/a crispa Spach, Nouv. Ann. Mus. Paris 4: 384. 1835. Type:

MEMOIRS OF THE TORREY BOTANICAL CLUB

51

Matamoros, Tamaulipas, Mexico, April 1831, /. L. Berlandier 2313
(G. Isotypes, BM, K, P).

Gaura hispida Benth., PI. Hartw. 288. 1840. Type: In fields near Leon,
Guanajuato, June 1837, Thomas Hartweg 1603 (K. Isotypes, BM,
CAMB, G).

Gaura crispa (Spach) D. Dietr., Syn. Pi. 2: 1298. 1840.

Guara roemeriana Scheele, Linnaea 21: 579. 1848. Type: New Braunfels,
Comal Co., Texas, 1846, Ferdinand Roemer ( B, destroyed. Isotypes,
CAS, MO).

Gaura odorata sensu Munz, Bull. Torrey Bot. Club 65: 219. 1938; N.
Amer. FI. II. 5: 194. 1965; non Sesse ex Lag. 1816.

Perennial herb from extremely aggressive horizontal rhizomes of about
the same diameter as the lower stem, the plants spreading to form large
patches; stems 2-6 (-12) dm tall, sometimes strict with a single unbranched
main stem but usually somewhat decumbent with several branches from
the base and usually much rather irregular branching above also. Entire
plant strigulose, the pubescence often denser on the ovary, outside of the
floral tube, and sepals, and the main stems sometimes with an admixture
of long, villous pubescence. Leaves narrowly lanceolate to elliptic, 0.5-7 .5
(-9.5) cm long, 0. 1-2.2 cm wide, subentire to shallowly sinuate-dentate.
Inflorescence 14-36 cm long, the peduncle 3-10 cm long, sometimes
branched; bracts narrowly lanceolate, 2-8 mm long, 0.8-2 mm wide. Floral
tube 4-14 mm long. Sepals 7-11 (-14) mm long, 0.75-2 mm wide. Petals
fading reddish, 6-10 mm long, 2.5-5 mm wide. Filaments 4-8.5 mm long;
anthers 3-6 mm long. Style 12-26 mm long. Fruit 4-angled, with 4 dis¬
tinct ridges and 4 furrows alternating at the top of the pyramidal upper
half, the base of the pyramidal portion distinctly bulging, then immediately
and sharply constricted, the basal portion terete and about one fourth the
diameter of the swollen base of the upper portion, 7-13 mm long, 3-5 mm
thick at the thickest point. Seeds (2-) 3-4 (-8), 2-2.5 mm long, 1-1.25 mm
thick, reddish biown. Self-incompatible. Gametic chromosome number.
n = 14.

Type: Austin, Travis Co., Texas, 1833-1834, Thomas Drummond III.
36. (G. Isotypes, BM, CAMB, GH, K, NY, P).

Distribution (fig. 48): Sandy loam at middle and lower elevations,
and spreading to become a weed of light agricultural soils, from central
Texas south through the Rio Grande Plain to eastern Coahuila, Nuevo Leon,
and Tamaulipas, then southward to southeastern Durango, southern San
Luis Potosi, Guanajuato, Queretaro, and Hidalgo, Mexico; absent in the
Trans-Mexican Volcanic Belt, but reappearing in southernmost Puebla and

52

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

MEMOIRS OF THE TORREY BOTANICAL CLUB

53

abundant in and around the Valley of Oaxaca. Occasional as a weed of
cultivated fields in coastal southern California, and collected once in Santa
Cruz County, California; locally established (persistent from 1923 to 1948,

at least) on Jekyll Island, Glynn County, Georgia and in Sevier County
Arkansas (in 1937).

MICH, MO, NY, US);
Wislizenius 299 (MO);
TEX); Muzquiz, Marsh
Durango and vicinity,

Representative Specimens Examined:

Mexico, coahuila. Saltillo, Palmer 23 (C, F, G, GH,
between La Varquieria and San Juan, 30 mi w of Saltillo!

Sierra de la Gloria, se of Monclova, Marsh 1916 (GH, SMU

111 (TEX); Monclova, White 1717 (MICH), durango. _ ^ miu ViUiimy

vc GH’ M°’ NY’ US^’ 71 mi ne of Durang°. Waterfall ir Wallis
11 (OKLA’ RSAh Guanajuato. 33 mi s of San Luis Potosi on road to Queretaro,
Jackson 7008 (DS); 34 mi n of Queretaro, Waterfall 16554 (OKL, OKLA, RSA
SMU). hidalgo. Between Actopan and Ixmiquilpan on highway to’ Laredo’, Gillu
^ Camp 6 (MICH, MSC); near El Salto, Rose & Painter 7101 (US); Ixmiquilpan
Rose et al. 9079 (GH, US); near Tula, Rose et al. 8323 (US), michoacan. Vicinity
of Morelia Arsene 5891 (ILL, LE, MO, MY, US), nuevo leon. 10 mi se of Las
Tortillas, Garcia & Garcia 64 ( DS, FSU, DUKE, NCU, SMU, TEX); 15 mi n of
Monterrey, Villanueva 57 (OKLA, TEX, TTC); Montemorelos, Kelley 17 (TEX)-

?lToT\R°Unal 34 (US)' OAXACA- 5 mi ne of Oaxaca on road to Ixtlan de Juarez’
6500 ft, Breedlove 12208 (DS); San Felipe del Agua, 1750 m, Conzatti 573 (GH)’

Nochitlan, 2000 m, Conzatti in 1882 (F); Monte Alban, Longman 3461 (PH)- near
Oaxaca, 5500 ft, Pringle 6715 (BR, CM, F, G, GH, IPN, ISC, LE, MEXU MIN

NY- P, PH, us, W); Huitzo, /. N. * /. S. Rose 11364 (GH,
, LS), Etla, Seler 83 (GH, US); 1 mi e of El Tide, Webster et al. 17M426 (MICH,
TEX), puebla. San Luis Tultitlanapa, Purpus 3387 (BM, F GH MO NY US)
queretaro. Queretaro, Arsene 10527 (F. MO, US); 11 mi ne of San Tuan del Rio'
Waterfall C Wall « 3937 (F, IA, OKL, OKLA, SMU), ,*N um potosi. Gauscama,
Minas de San Rafael, Purpus 5383 ( BM, F, GH, MO, NY, US); Teneria, 3 km sw of
San Luis Potosi, Rzedowski 3332 (IPN). tamaulipas. 24 km s of Nuevo Laredo
Dominguez Is McCart 8214 (SMU, TEX); Soto la Maria, Viereck 1124 (US)- vie, mu’
of Victoria, Palmer 152 (F, GH, MY, US). ’

United States. Arkansas. Sevier Co.: Horatio, Brinkley 96 (F). California
Los Angeles Co.: Puente, Johnson in 1939 (POM); Avalon, Santa Catalina Island’
lorne 36478 ( DS, RSA). Orange Co.: 1 mi n of Placentia, Johnson in 1940 (POM)
Santa Barbara Co.: Goleta Point, Raven 14045 (DS). Santa Cruz Co - Ca 2 mi n

IvnTt’JfZ 5m ^ RSA)- San Dieg° C°-: Fletcher Hills- Zander 8787
(POM, RSA). Ventura Co.: Meiners Oaks, Pollard in 1946 (CAS COLO RS\)

Jeky11 Ls,land’ McMee in 1923 (US>’ Th°™ b' Muenscher 8905
(in 948) (CU). Texas. Archer Co.: 2 mi w of Archer City, Shinners 18559 (SMU)

Austin Co.: Industry, Wurzlow in 1894 (ILL). Bastrop Co.: Bastrop, Duval in 1928
(U3). Bee Co.: 5 mi ne of Beeville, Rios Cavazos 198 (LL). Bell Co.: Near
Temple, Wolf 3555 ( F, TAES). Bexar Co.: San Antonio, Heller 1590 (CU F GH
MICH, MIN, MSC, MO, NY, PH, US). Brazos Co.: Lindheimer in 1843 (MO)’
Brown Co.: Brownwood, Ewing 47 (SMU, TEX). Burnet Co.: 13 mi w of Burnet!
Jones 8 (LL). Calhoun Co.: La Feria, Cannon in 1926 (MICH). Cherokee Co •
Old Frankston Crossing, Stripling (FSU). Comal Co.: New Braunfels, Lindheimer

54

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

in 1848 (MIN, SMU). Crockett Co.: Ozona, Parks ir Cory 29669 (TAES). Dallas
Co.: Oak Cliff, Mervin in 1953 (SMU, TEX). Dewitt Co.: Cuero, Howell 296
(US). Dimmit Co.: 5 mi se of Catarina, C. 6- V. C. Flores 171 (SMU, TAES, TEX).
Duval Co.: San Diego, Croft in 1886 (NY). Erath Co.: Dublin, Maxwell 28 (F).
Fayette Co.: Muldoon area, Ripple 51-589 (TEX). Frio Co.: Dilley, Drews 7 (TEX).
Gillespie Co.: 30 mi s of Mason, Munz ir Gregory 23433 (RSA). Guadalupe Co.:
Seguin, Gregory 286 (RSA). Harris Co.: N of Harrisburg, Eggert in 1899 (MIN,
MO). Hay Co.: 3 mi s of Dripping Springs, Knodel <Lr Warnock 46511 (SMU).
Hidalgo Co.: Los Ebanos, Runyon 1582 (POM). Hood Co.: Center Mills, Blackwell
49 (SMU). Jim Hogg Co.: 2 mi ne of Hebbronville, Saenz 92 (OKLA, TEX).
Kendall Co.: 1.1 mi n of Boerne, Raven ir Gregory 19343 (DS). Kimble Co.: 8
mi s of Junction, E. R. 6- E. M. Bognusch 4719 (POM). Kleberg Co.: Kingsville,
Sinclair G-16 (GH, TEX). La Salle Co.: 5 mi n of Encinal, Cabrera 91 (TEX).
Live Oak Co.: 5 mi sw of George West, Rios Cavazos 214 ( LL, SMU). McCulloch
Co.: Brady, Henderson 62-267 (FSU). McLennan Co.: Speegleville, Smith 1103
(TEX). Menard Co.: 10.3 mi n of Menard, Raven 6- Gregory 19267 (DS). Milam
Co.: Ben Arnold, Sliinners 16662 (SMU). Nueces Co.: Corpus Christi, Orcutt 5936
(MO). San Patricio Co.: 1 mi nw of Mathis, Raven i? Gregory 19384 (DS). Shackle¬
ford Co.: Griffen State Park, Rohrbaugh 475 (COLO, OKL). Starr Co.: 4 mi e of
Rio Grande City, Navarro 96 (DS). Tarrant Co.: Fort Worth, Ruth 276 (US).
Tom Green Co.: San Angelo, Tharp 44184 (ISC, TEX). Travis Co.: Austin, Hall
213 (F, GH, MO, NY, PH, POM, US). Uvalde Co.: Sabinal, Palmer 10101 ( DS,
MIN, PH, US). Val Verde Co.: San Felipe Creek near Del Rio, Correll 14943
( LL, US). Webb Co.: 6 mi s of Laredo, Cisneros 16 (DS, FSU, NCU). Willacy
Co.: 1 mi n of Sebastian, Runyon 2509 (POM). Williamson Co.: Georgetown, Bodin
158 (US). Wilson Co.: Floresville, Metz in 1931 (US). Zapata Co.: 6 mi s of San
Ignacio, Garcia 16 (DUKE, FSU, NCU, SMU).

In the absence of mature fruit, Gouro drummondii can be separated
only with great difficulty from G. coccinea. Although G. drummondii
generally occurs at lower elevations and in somewhat more fertile soil, the
two species do occur together and appear to hybridize rather freely when
they do. Nevertheless, they are strikingly different in fruit and modally also
in habit ( Goura drummondii is strongly rhizomatous) and maintain their
distinctiveness over very wide areas.

It is unfortunate that the name Goura odorato, applied to this species
for more than 30 years, must now be abandoned in favor of the name that
was generally associated with it earlier, as for example by Rothrock (Proc.
Amer. Acad. Arts 6: 353. 1864-1865). No authentic material of G. odorato
seems to have survived in the herbarium at Madrid, but Dr. Munz ex¬
amined a specimen at Geneva, labeled “G. odorata Lag., Ilort. reg (“veg.”)
Matrit 1814? Reuter ( Boissier ) comparing it with Bourgeon 38 from the
Valley of Mexico. Bourgeon 38 is G. coccinea, which is extremely common
in and about Mexico City, where G. drummondii is unknown. Lagasca
tells us that Martin de Sesse y Lagasta brought him the seeds of his G.
odorato when he returned from Mexico City to Madrid in 1804. It is well

MEMOIRS OF THE TORREY BOTANICAL CLUB

55

known that Sesse made no extensive trips during his last year or two in
Mexico City, and G. drummondii does not occur around the city, the closest
station being some 80 km to the north near Pachuca. Although Sesse,
Mocino, Castillo, and Maldonado collected G. coccinea several times and
illustrated it twice, they are not known ever to have collected G. drum¬
mondii. Although circumstantial in part, the evidence appears overwhelm¬
ing that G. odorata is in fact a synonym of G. coccinea, and that the name
has been misapplied to G. drummondii in recent years.

SECTION VII. GAURA

Gaura sect. Eugaura Munz, Bull. Torrey Bot. Club 65: 106. 1938.

Annual, biennial, or perennial herbs, the stems villous, glandular, or
strigulose. Inflorescence erect. Flowers strongly zygomorphic, the stamens
in the lower quarter or third; flowers opening near sunset and withering
the next morning, or opening near sunrise and withering in the afternoon.
Fruit not reflexing, narrowly ellipsoid or ellipsoid, sharply 4-angled, with
fairly deep furrows alternating with the angles for 2-3 mm from the apex,
ribbed from base of furrow to base of the fruit. Self-incompatible or self¬
compatible. Gametic chromosome number, n = 7; one species (G. biennis )
a complex structural heterozygote.

Type Species: Gaura biennis L.

Distribution: Nearly throughout in eastern half of the United States,
but rarer northward, and barely entering southernmost Canada, in more or
less open sandy places, with one species (G. lindheimeri ) in black prairie
soil.

With six species, section Gaura is the largest in the genus. Of these,
G. longiflora and its recent derivative G. demareei are self-incompatible,
as is the morning-opening G. lindheimeri. The self-compatible, complex
structural heterozygote G. biennis has doubtless been derived from northern
populations of G. longiflora, and has become the only species of the genus
throughout most of the northeastern United States. Somewhat more dis¬
tantly related are the self compatible G. neomexicana, obviously a relict
restricted to its present three disjunct areas in Colorado and New Mexico
by the elimination of summer rainfall over much of the western United
States; and the essentially autogamous G. angustifolia, the most common
and often only species of the genus in the Coastal Plain of the southeastern
United States.

12. Gaura lindheimeri Engelm. & Gray, Bost. J. Nat. Hist. 5: 217. 1845.
Munz, Bull. Torrey Bot. Club 65: 115. 1938; N. Amer. FI II 5- 187
1965. Figs. 13, 49.

56

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

2 53 54

FIGURES 49-55. Flowers of the species of Gaura sect. Guura. Fig. 49. G. lind-
heimeri.— Fig. 50. G. a ngmtifolia .—Fig. 51. G. neomexicana subsp. neomexicana.—
Fig. 52. G. longi flora.— Fig. 53. G. biennis.— Fig. 54. G. demareei.

MEMOIRS OF THE TORREY BOTANICAL CLUB

57

Gaura filiformis Small var. munzii Cory, Rhodora 39: 420. 1937. Type:
About 6.5 mi n of High Island, Chambers Co., Texas, 17 September
1936, V. L. Cory 20179 (GH). Munz, Bull. Torrey Bot. Club 65: 120.
1938; N. Amer. FI. II. 5: 187. 1965.

Robust clumped perennial herb, usually branching copiously from a
heavy underground crown, the stems erect or more often arching outward
from the crown to form a full, clumped plant 5-15 dm tall. Entire plants
villous, somewhat more densely so below, the trichomes mostly 1-2 mm
long, subappressed on the densely pubescent leaves, or the plants more
rarely subglabrous; inflorescence with a more or less dense admixture of
glandular trichomes less than 1 mm long. Leaves very narrowly elliptic
(to occasionally narrowly oblanceolate), 0.5-9 cm long, 0.1-1.3 mm wide,
coarsely and remotely serrate, occasionally subsinuately so, diminishing in
size upward. Inflorescence well branched or simple, 10-80 cm long; bracts
narrowly to broadly elliptic, 4-11 mm long, 1.5-6 mm wide. Flowers open¬
ing near sunrise. Floral tube 4-9 mm long, densely lanate in the upper
two-thirds or half within. Sepals 8.5-17 mm long, 1-2 mm wide. Petals
white, fading to light or deep pink, 10.5-15 mm long, 5-10 mm wide.
Filaments 7.5-12 mm long; anthers usually dull red, 3.5-4.5 mm long. Style
16-26.5 mm long, lanate near the base. Fruit 6-9 mm long, 2-3.5 mm
thick. Seeds 1-4, 2-3 mm long, 1-1.5 mm thick, yellowish to light brown.
Self -incompatible. Gametic chromosome number, n ~ 7.

Type: Prairies from Houston to the Brazos, Harris and Fort Bend
counties, Texas, April to May 1843, Ferdinand Jakob Lindheimer 1.61 ( MO.
Isotypes, BM, CAMB, GH, LE, P, PH, US, W).

Distribution (fig. 55): Black prairie soil in the coastal prairie of south¬
eastern Texas and southwestern Louisiana; Tangipahoa Parish, Louisiana.
Flowers April-July.

Representative Specimens Examined:

United States. Louisiana. Acadia Parish: 3 mi w of Raye, Thieret 22847 ( DS,
LAF). Allen Parish: Road from Keman to Kinder, Crockett 8643 (LL). Beauregard
Parish: Near Longville, Cooleij <b Brass 4075 (GH, USF). Calcasieu Parish: 1 mi
e of Lake Charles, D. S. ir H. D. Correll 9636 (DUKE, F, GH, NY). Jefferson Davis
Parish: Jennings, Palmer 7612 (MO, NY, US); Munz 13340 (F, G, NY, POM).
Lafayette Parish: Without other data (NY). St. Mary Parish: Near Centerville,
Munz 1588 (POM). Tangipahoa Parish: Hammond, Cocks in 1903 (NO); 1 mi n
of Ponchatoula, Wilson 1070 (FSU). Vermillion Parish: 3.5 mi sw of Kaplan, Thieret
& Tuselier 19787 (DS, LAF). Texas. Brazoria Co.: 2.2 mi n of Alvin, Shinners 24398
(SMU). Brazos Co.: 8-10 mi s of College Station, Reeves 1846 (POM). Chambers
Co.: 14.5 mi nw of White Ranch, Cory 22459 (POM). Fort Bend Co.: Areola,
Raven 20363 (DS). Galveston Co.: Kemah, Fisher in 1937 (C, OKLA, US).
Hardin Co.: 2.6 mi e of Camp Jackson, Parks 6- Cory 19762 (TAES). Harris Co.:

58

A REVISION OF THE GENUS CAURA ( ONACRACEAE )

FIGURE 55. Ranges of some species of Gaura sect. Gaura. Gaura lindheimeri
(circles), G. angustifolia (open diamonds), G. neomexicana subsp. ncornexicana (filled
diamonds), G. demareei (dots), G. neomexicana subsp. coloradensis (filled triangles),
G. biennis (open triangles). Outline of range of G. longiflora, the remaining species
of the section, indicated by the dotted line.

5 mi n Houston, Correll 14161 ( LL, SMU); Sheldon prairie, Reverchon 3842 (GH,
MIN, MO, POM, US). Jasper Co.: 6 mi s of Buna, Correll et al. 22302 (LL).
Jefferson Co.: 6 mi e of Port Arthur, Cory 50820 (SMU). Liberty Co.: 3 mi e of
Liberty, Munz 13342 (BII, G, GH, POM). Orange Co.: 10 mi e of Beaumont,
Munz <Lr Gregory 23460 (RSA). Victoria Co.: McFadden Beach, Tharp in 1937
(TEX). Wharton Co.: 35 mi w of East Bernard, Raven 20362 (DS).

In addition to these presumably native localities, Gaura lindheimeri
has been popular in private and public gardens since it was introduced into
cultivation in the mid-1840’s. It is cultivated throughout the temperate
regions of the world but not, as far as we know, ever truly spontaneous
outside of Texas and Louisiana. The following are a few cultivated speci¬
mens to indicate something of the range:

Argentina. Mendoza, Jorgenson 172 (C). Australia, new south wales.
Camden, “Spontaneous,” McRarron 10365 (NSW); Hazelbrook, 2200 ft. Constable in
1949 (NSW). Canada. Ontario, 3 mi se of Thomas, James 2018 ( DAO). China,
chekianc (UC). England. Cambridge, Leabitt in 1898 (MO). Germany. Stadt-

MEMOIRS OF THE TORREY BOTANICAL CLUB

59

gaiten, Miinchen, Alayer in 1896 ( MICH). Italy. Botanical Gardens, Firenze, in
1857 (FI). Mexico, coahuila, near Saltillo, Nicolas in 1910 (BR, G, GH, K,
LE, NY, P); Remate, near Monterrey, Nuevo Leon, Arsene in 1911 (NY, P). puebla,
vicinity of Puebla, Nicolas (G). Sweden. Kolmar, Anderson in 1929 (DS, OKL,
SMU). United States. California. Berkeley, University of California Botanic
Garden, E. B. Babcock in 1903 (UC, US); iowa. Marshalltown, Harmon in 1929
(ISC); new jersey. Passaic, Rusby in 1877 (MICH).

There seems no reason to regard the collections from Mexico as being
from native populations, since the areas are completely out of line with
those within the native range of the species ecologically and there are
no undoubtedly spontaneous collections of Gaura lindheimeri from Mexico.
The type material of G. filiformis var. munzii is typical G. lindheimeri.
Hybridization between G. lindheimeri and G. longiflora is discussed under
the latter species.

13. Gaura angustifolia Michx., FI., Bor.-Amer. 1: 226. 1803. Munz, Bull.
Torrey Bot. Club 65: 116. 1938; N. Amer. FI. II. 5: 187. 1965 ’ Figs
14, 15, 50.

Gciuro fruticosa Jacq., Coll. 1: 142, pi. 181. 1786; non G. fruticosa Lofl.,
Iter Hisp. 248. 1758. Type: Cultivated in the garden at Schonbrunn
Palace in \ ienna from seeds collected by Boos and Bredemeyer in the
vicinity of Charleston, South Carolina, June-September 1785, N.
J acqiiin.

Gaura simulates Small, Bull. New York Bot. Gard. 3: 432. 1905. Type:
In pinelands near the unfinished railroad grade, between Coconut
Grove and Cutler, Dade Co., Florida, 31 October to 4 November 1903,
J. K. Small & J. J. Carter 766 (NY. Isotypes, F, GH, MIN).

Gaura eatonii Small, FI. S. E. U. S. ed. 2, 1353, 1375. 1913. Type: Sandy
beach, Punta Rassa, Lee Co., Florida, 4 March 1905, A. A. Eaton 1289
(NY).

Gaura angustifolia Michx. var. typica Munz, Bull. Torrey Bot Club 65-

117. 1938.

Gaura angustifolia Michx. var. strigosa Munz, Bull. Torrey Bot. Club 65:

118. 1938. Type: Dry sandy soil, vicinity of Eustis, Lake Co., Florida,
16-31 July 1894, G. V. Nash 1464 (NY. Isotypes, BY, CU, DAO, F,
G, GH, LE, MICH, MIN, MO, MSC, PH, US). Munz, N. Amer FI
II. 5: 188. 1965.

Gaura angustifolia Michx. var. simulates (Small) Munz, Bull. Torrey Bot.

Club 65: 118. 1938; N. Amer. FI. II. 5: 188. 1965.

Gaura angustifolia Michx. var. eatonii (Small) Munz, Bull. Torrey Bot.
Club 65: 118. 1938; N. Amer. FI. II. 5: 188. 1965.

60

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Annual herb, occasionally persisting through mild winters in the southern
parts of its range, from a taproot; stem usually single, but occasionally with
several from the base, unbranched to branched near the base or above,
6-18 dm tall. Plants subglabrous, strigulose, or hirtellous, or with a mix¬
ture of strigulose and hirtellous pubescence. Leaves narrowly lanceolate
to narrowly oblanceolate, often red-blotched, 0.8-13 cm long, 0.1-1.25 cm
wide, slightly to markedly sinuate-dentate. Inflorescence simple or fairly
well branched, 10-70 cm long; bracts lanceolate, 2-3.5 mm long, 0.75-1
mm wide. Flowers 4-merous or 3-merous, often both types mixed on a
single plant, opening near sunset. Floral tube 3-8 mm long. Sepals 2.5-8
mm long, 0.5-1. 5 mm wide. Petals 4.5-8 mm long, 0.5-1. 5 mm wide.
Petals 4.5-8 mm long, 1-3 mm wide. Filaments 2.5-6 mm long; anthers
0.5-2 mm long, shedding pollen directly on the stigma at anthesis. Style
7.5-19 mm long. Fruit 5-9 mm long, 2-3 mm thick. Seeds 2-4, 1.5-2.25
mm long, ca. 1 mm thick, yellowish to light brown. Self-compatible and
largely autogamous. Gametic chromosome number, n = 7.

Type: Vicinity of Charleston, South Carolina, 1787-1802, Andre Michaux
(P. Isotype, F).

Distribution ( fig. 55 ) : Sand hills and sandy flats, dunes and waste
places, on the Coastal Plain from Cape Hatteras, North Carolina to and
throughout Florida. Flowers chiefly May to August, but sporadically in all
months, particularly in southern Florida.

Representative Specimens Examined:

United States. Florida. Alachua Co.: Millhopper Reserve, mv of Gainesville,
Wiggins 19675 ( DS, FLAS, USF). Bay Co.: 15 mi s of Round Lake Common,
T. 6- L. Mosquin 5868 (DAO). Bradford Co.: 6 mi e of Lake Butler, Godfrey
63899 ( DS, FSU). Brevard Co.: Cape Canaveral, Burgas 654 ( F, NY). Broward
Co.: Pompano Beach area, Lakela 28294 (FLAS, RSA, USF). Calhoun Co.: 8 mi n
of Wewahitchka, Ford 3325 (FLAS). Charlotte Co.: Chadwick Beach, Deam 57722
( F, IND). Citrus Co.: 5 mi s of Homosassa, Krai 7760 (FLAS, FSU, GA, GH, USF).
Clay Co.: 9 mi s of Creen Cover Springs, Rossbach 2106 (FLAS, WVA). Collier
Co.: Marco Island, Ward 5387 (DS, FLAS). Columbia Co.: 7.8 mi w of Baker Co.
line on U.S. Hwy. 90, Ward 4800 ( DS, FLAS). Dade Co.: Miami, Garber in 1877
(F, FLAS, GH, NY, PAC, PH, US). De Soto Co.: 14 mi e of Arcadia, Adams 136
(GA, GH, NCU, NSC, USF). Dixie Co.: 5 mi s of Cross City, Murrill in 1940
(FLAS). Duval Co.: Near Jacksonville, Curtis 895 ( F, FLAS, GA, IA, LE, MO,
NMC, PENN, PH, US). Flagler Co.: 10 mi e of county line on Hwy. 28, West
& Arnold in 1940 (FLAS). Franklin Co.: 3 mi sw of Panacea, Krai 4688 (FSU,
GH, LAF, SMU). Glades Co.: 10 mi n of Lakeport, Will in 1960 (FLAS). Hamilton
Co.: 2.7 mi w of Jasper, Godfrey 63687 ( DS, FSU). Hardee Co.: Cooley et al.
6107 (FSU, GA, GH, NY, USF). Hendry Co.: S of Labelle, Lakela 26935 (USF).
Hernando Co.: Brooksville, Murrill in 1939 (FLAS). Highlands Co.: 8 mi w of
Childs, Deam 61188 (IND). Hillsborough Co.: N of Gibsonton, Lakela 25183 (DS,

MEMOIRS OF THE TORREY BOTANICAL CLUB

61

FLAS, GH, NCU, SMU, USF, VDB). Indian River Co.: Near Fellsmere, Small
et al. 1113 (GH, NY, US). Jefferson Co.: Hitchcock 670 ( F, MO). Lafayette Co.: 2.7
mi sw of Mayo, Ward 4485 (I1 LAS). Lee Co.: W of Buckingham, Moldenke 1008
(DUKE, MIN, MO, N^ , PENN, US). Leon Co.: Near Tallahassee, Godfreij 53488
(DUKE, ESU, GH, NY). Levy Co.: 5 mi s of Williston, Krai 4855 (FSU, SMU).
Liberty Co.: Sargent in 1950 (KANU, MIN, OKL, WVA). Madison Co.: 6.8 mi e
of Greenville, Raven 18633 (DS). Manatee Co.: Bradenton, Longboat Key, Long ir
Lakela 2i581 ( DS, FLAS, OKLA, USF). Marion Co.: Ocala National Forest, 1.5
mi e of Juniper Springs Road, Mather M-296 (USF). Martin Co.: 2 mi s of Stuart,
West in 1944 (FLAS). Monroe Co.: Big Pine Key, Killip 43200 (FLAS, MICH,
NO, MSC, NY). Nassau Co.: Yulee, Raven 18699 (DS). Okaloosa Co.: 2.7 mi e
of Destin Bridge, Raven 18592 (DS). Okeechobee Co.: Zigzaggy, Demaree 50228
(DS, NO). Orange Co.: N of Winter Park, Bright 3676 (CM). Osceola Co.: Road
to Gables Golf Course, Singeltary in 1938 (DUKE, MSC). Palm Beach Co - Near
Palm Beach, von Reis 201 (MICH, US). Pasco Co.: 10 mi n of Hillsborough Co
line, Wiggins 19595 ( DS, FLAS, USF). Pinellas Co.: Mullet Key, Lakela 27625
(DS, USF). Polk Co.: 10 mi e of Lake Wales, Demaree 49442 (DS, SMU).
Putnam Co.: 3.6 mi nw of Pomona Park, T. 6- L. Mosquin 5911 (DAO). St. Johns
Co.: San Sebastian River, Godfrey 63965 (DS, FSU). St. Lucie Co.: Fort Pierce
Burgess 735 ( F, NY). Santa Rosa Co..- Santa Rosa Island, Mohr in 1872 (US) Sara¬
sota Co.: Longboat Key, Lakela ir Laker 28698 (DS, FSU). Seminole Co.: Cooley
et al. 7422 (USF). Sumter Co.: 4 mi s of Bushnell, Krai 6847 (FLAS, FSU GA
CH, USF). Suwannee Co.: 5 mi w of O’Brien, West 6- Arnold in 1946 (FLAS)’
Union Co.: Lake Butler, Godfrey 63859 ( DS, FSU). Volusia Co.: 6 mi e of Geneva
Kra! 5170 (FSU, GH, SMU). Walton Co.: 16 mi e of Destin, Dress & Read 7656
(BH). Georgia. Camden Co.: 10 mi w of Kingsland, Gregory 540 (DS). Chatham
Co.: Tybee Island, Duncan 21178 ( GA, KY, LAF, TENN). Decatur Co- Near
Hutchinsons Ferry, Thorne 4746 (CU, IA). Glynn Co.: 10 mi n of Brunswick
Munz 6 Gregory 23490 ( RSA). Lanier Co.: 4.9 mi e of Lakeland, Krai 24257 FSU)!
McIntosh Co.: Sapelo Island, Duncan 20164 (DUKE, F, GA, GH, ISC, NCU, SMu!
TEX, US, USF). Seminole Co.: 3 mi e of Donaldsonville, Thorne 4430 (CU GA)
Ware Co.: Waycross Airport, Runs 279 (WISC). Wayne Co.: 2 mi n of Jessup,
Fitzgerald 72 (GA). north Carolina. Beaufort Co.: Near Bayview, Correll 1653
(DUKE). Brunswick Co.: Southport, Wells in 1949 (NSC). Cartaret’ Co • Shackle
ford Banks, Anderson 359 (DUKE, FSU, SMU). Craven Co.: Havelock Radford
37495 (NCU). Dare Co.: Buxton, Pence 45091 (CM, KY, LAF, NCU) Hyde Co •
Swan Quarter, Radford & Stewart 797 (NCU). Jones Co.: Maysville, Radford 37229
(NCU). New Hanover Co.: 0.5 mi n of bridge into Wrightsville Bridge, Gregory 527
(DS). Onslow Co.: 4.8 mi w-sw of Haw, Allies 6- Haesloop 28203 (KANU NCU)
Pamlico Co.: Front of Bay River, L. F. <b F. R. Randolph 856 ( BH, CU GH)
Pender Co.: Aides & Haesloop 27988 (FLAS, NCU, SMU). south Carolina’ Beau¬
fort Co.: Beaufort, Churchill in 1917 (C.H, MIN, MO). Berkeley Co.: 10 mi nw of
Charleston, Godfrey <x Try on 712 (F, CH, MO, MY, POM, US). Charleston Co-
Sullivans Island, Gibbs in 1856 (NY). Colleton Co.: 14 mi e of Walterboro, Wiegand

(CU’ GH)' Georgetown Co-: 3 mi 11 of Litchfield Beach, Gregory
(DS). Hampton Co.: 1.7 mi se of Early Branch, BeU 6- Aides 15719 (GA NCU)

Horry Co.: Myrtle JBeach , Baker in 1954 (CM). Jasper Co.: 3 mi „e of Coosawhatchie;
Aides 6- Bell 15687 (NCU, USF).

62

A REVISION OF THE GENUS CAURA ( ONAGRACEAE )

The four varieties into which Gaura angustifolia was divided by Munz
(1938) differ in the character of the pubescence in the inflorescence only.
Although they are weakly correlated geographically, there are numerous
intermediates between them. The pattern of variation in this species is
not extraordinary for such a strictly autogamous taxon, and does not appear
to warrant taxonomic subdivision. Gaura angustifolia was collected once
as a weed on ballast in the Navy Yard at Philadelphia ( Parker , PPI), cur¬
iously enough on 30 September 1865; one may wonder how it became es¬
tablished at Philadelphia during the Civil War in view of its known range.

14. Gaura neomexicana Wooton, Bull. Torrey Bot. Club 25: 307. 1898.
Munz, Bull. Torrey Bot. Club 65: 113. 1938; N. Amer. FI. II. 5: 186.
1965.

Biennial herb sending up several stems from the top of a fleshy, carrot¬
like taproot; stems erect, 5-12.5 dm tall, unbranched to moderately branched.
Pubescence villous, glandular, or strigulose. Leaves subglabrous or strigu-
lose, very narrowly elliptic, 2-13 cm long, 1-4 cm wide, subentire to repand-
denticulate, those of the rosette to 33 cm long. Sepals 9.5-15 mm long,
1.5-2. 5 mm wide. Petals 7-13.5 mm long, 4-8 mm wide. Filaments 6.5-9
mm long; anthers 2.5-4 mm long. Style 19-28 mm long. Fruit 6-11 mm
long, 2-5 mm thick. Seeds 2-4, 2-3 mm long, 1-1.5 mm thick, yellowish to
light brown. Self-compatible. Gametic chromosome number, n.= 7.

Type: White Mts., 6500 ft elevation, Lincoln Co., New Mexico, 26
July 1897, E. O. Wooton 204 (US. Isotypes, G, LE, MIN, MO, ND, NY, P).

Distribution (fig. 55): Heavy sod in mountain meadows and openings,
very local: southern Larimie County, Wyoming, and adjacent Weld County,
Colorado; near Fort Collins, Larimer County, Colorado (probably ex¬
tinct); western foothills of San Juan Mountains, Archuleta County, Colorado,
and Rio Arriba County, New Mexico, and perhaps also near Durango, La
Plata County, Colorado; Sierra Blanca and Sacramento Mountains in
Lincoln and Otero counties, south-central New Mexico. Flowers July to
September.

In view of the remarkably disjunct range, the differences between the
northernmost and two southern populations of this species appear to war¬
rant taxonomic recognition.

14a. Gaura neomexicana subsp. neomexicana. Figs. 16, 51.

Gaura neomexicana Wooton var. typica Munz, Bull. Torrey Bot. Club
65: 114. 1938.

Stems with both villous and strigulose pubescence, the inflorescences
occasionally with a few long hairs. Sepals 11-15 mm long. 1. 6-2.5 mm wide.

MEMOIRS OF THE TORREY BOTANICAL CLUB

63

Petals 10.8-13.5 mm long, 5-8 mm wide. Style 22-28 mm long. Fruit
8.5-11 mm long, 4.5 mm thick. Self-compatible. Gametic chromosome
number, n = 7.

Distribution (fig. 56): Mountain meadows in openings in coniferous
forest, western foothills of the San Tuan Mountains in Archuleta County,
Colorado, and Rio Arriba County, New Mexico; Sierra Blanca and Sacra¬
mento Mountains in Lincoln and Otero counties, south-central New Mexico.
Collected once at Durango, La Plata County, Colorado, but the locality
needs to be confirmed.

Representative Specimens Examined:

United States. Colorado. Archuleta Co.: Pagosa Springs, Baker 490 (BM, G,
GH, M, MO, ND, N\, POM, US), Bethel in 1917 (COCO); 18 mi s of Pagosa Springs,
Penland 4656 (COCO, COLO); Piedra, Baker 491 (MO, ND, NO, NY, POM, US).
La Plata Co.: Durango, Livingston in 1936 (COCO), new Mexico. Otero Co.: Apache
Indian Reservation, Sacramento Mts., Huber in 1932 (PH); James Canyon, Sacra¬
mento Mts., Wooton in 1899 (US); vicinity of Cloudcroft, Wooton in 1899 (COLO,
DS, CH, ILL, NY, POM, US); High Rolls, Young in 1916 (TEX); Tularosa Creek,
3 mi s of Mescalero Agency, Wolf 2768 ( BH, DS, GH, POM, RSA); 2.8 mi w of
Cloudcroft, Harms 1833 ( KANU, SMU). Rio Arriba Co.: Vicinity of Chama Standley
6783 (NY, US).

The populations in southern Colorado have much less villous pubescence
than those in the White Mountains of New Mexico, thus in some sense
being transitional to the more northerly subsp. coloradensis.

14b. Gaura neomexicana subsp. coloradensis (Rydb.) Raven & Gregory,

comb. nov. Fig. 17.

Gaura coloradensis Rydb., Bull. Torrey Bot. Club 31: 572. 1904.

Gaura neomexicana Wooton var. coloradensis (Rydb.) Munz, Bull.

Torrey Bot. Club 65: 114. 1938; N. Amer. FI. II. 5: 186. 1965.

Stems strigulose, the inflorescences with glandular pubescence only.
Sepals 9.5-13 mm long, 1.5-2 mm wide. Style 19-25 mm long. Fruit 6-8,5

mm long, 2-3 mm thick. Self-compatible. Gametic chromosome number
n = 7.

Type: Fort Collins, Larimer Co., Colorado, 8 July 1895, ]. H. Cowen
( NY. Isotype, GH ) .

Distribution (fig. 56): Mountain meadows, in heavy soil, southern
Laramie County, Wyoming, and northern Weld County, Colorado; once
near Fort Collins, Larimer County, Colorado, but apparently now extinct
there (not collected since 1897).

64

A REVISION OF THE GENUS GAURA ( ONACRACEAE )

Specimens Examined:

United States. Colorado. Larimer Co.: Meadows east of college, Fort Collins,
Crandall 1308 (ISC, NY); Fort Collins, Cowen in 1895 (US); e of Poudre, Cotven
1632 (COCO); 30 mi n of Fort Collins, 6200 ft, Harrington in 1944 (COCO).
Weld Co.: 3.2 mi s of Wyoming line on U.S. Hwy. 87, Porter 9690 (DS, MSC,
RSA). Wyoming. Laramie Co.: Pine Bluffs, Bodin in 1889 (MIN).

15. Gaura longiHora Spach, Nouv. Ann. Mus. Hist. Nat. 4: 380. 1835.
Figs. 18, 19, 53.

Gaura biennis L. var. pitched T. & G., FI. N. Amer. 1: 517. 1840. Type:
Red River, presumably near the site of Fort Towson, Chocktaw Co.,
Oklahoma, 1831-1834, Zina Pitcher (NY). Munz, Bull. Torrey Bot.
Club 65: 113. 1938; pro parte; N. Amer. FI. II 5: 185. 1965; pro parte.

Gaura filiformis Small, Bull. Torrey Bot. Club 25: 617. 1898. Type:
Near Texarkana, Miller Co., Arkansas, 23 August 1898, A. A. i? E. G.
Heller 4138 (NY. Isotypes, CU, F, G, GH, ISC, LE, MO, NY, P,
PH, US). Munz, Bull/ Torrey Bot. Club 65: 118. 1938; N. Amer. FI.
II. 5: 188. 1938.

Gaura filiformis Small var. kearneyi Munz, Bull. Torrey Bot. Club 65:
119. 1938. Type: Along railroad track, Starkville, Oktibbeha Co.,
Mississippi, 28 Sept. 1896, T. H. Kearney, Jr. (NY).

Gaura biennis sensu Munz, Bull. Torrey Bot. Club 65: 111. 1938; N.
Amer. FI. II. 5: 185. 1965; et auct. mult., pro parte.

Robust winter annual or perhaps occasionally biennial from a fleshv
taproot; stems usually well branched above the base, 5-40 dm tall. Stems
and leaves more or less densely strigulose with an admixture of glandular
or villous pubescence in northern populations; inflorescence densely strigu¬
lose, hirtellous, or glandular, the latter two hair types often intermixed.
Leaves usually narrowly elliptic, 1.5-13 cm long, 0.2-2.75 cm wide, sub¬
entire or shallowly undulate-denticulate, often left in short axillary fascicles
after main leaves have dropped; rosette leaves (withered by time of
flowering) up to 40 cm long and to 3 cm or more wide, often highly ir¬
regular in outline. Inflorescence well branched; bracts lanceolate or ovate,
1-6 mm long, 0.5-2 mm wide. Flowers opening near sunset. Floral tube
4-13.5(-15) mm long. Sepals 7-18 mm long, 1-2.3 mm wide. Petals 6.5-15
mm long, 2.5-7 mm wide. Filaments 5-13 mm long; anthers 1.5-5 mm
long. Style 12-33.5 mm long. Fruit 4.5-7 mm long, 1.5-2.5 mm thick.
Seeds 2-4, 1.25-3 mm long, 0.75-1.25 mm thick, yellowish to reddish
brown. Self-incompatible. Gametic chromosome number, n = 7.

Type: Jacksonville, near present-day Sheridan, Washington Parish,
Louisiana, autumn 1832, Thomas Drummond (F. Isotypes, GH, K, P).

MEMOIRS OF THE TORREY BOTANICAL CLUB

65

FIGURE 56. Range of Gaura longiflora ( G . filiformis ). Plants with strigulose
pubescence on the sepals are indicated by solid triangles, those with glandular
pubescence by circles.

The type locality of this species has generally (and erroneously) been
considered to have been Jacksonville, Florida, one factor in its having been
relegated to the synonymy of G. angustifolia.

Distribution (fig. 56): Open grassland and waste places, often along
roadsides and railway embankments, southeastern Nebraska, Iowa, Illinois,
and western and northern Indiana south to eastern and southern Texas,
Louisiana, Mississippi, and Alabama; rare and scattered in Colorado, cen¬
tral and western Nebraska, western Kansas, Minnesota, Wisconsin, Mich¬
igan, and Ohio, with one station each in southern Ontario, Massachusetts,
Connecticut, Pennsylvania, and Maryland. Probably introduced at most
stations outside of its main area of distribution. Flowers July to October.

66

A REVISION OF THE GENUS C.AURA (ONAGRACEAE)

Representative Specimens Examined:

Canada. Ontario. Scarborough Twp., Ontario, Scott in 1905 (DAO).

United States. Alabama. Bullock Co.: Flora, 3929g (MIN). Choctaw Co.:
Cocoa, Schuchert in 1896 (NY). Greene Co.: 1 mi n of Forkland, Whitehou.se
24360° (SMU). Hale Co.: 1 mi nw of Rosemary, Harper 3253 (BY, GH, MO, PH).
Lee Co.: Auburn, Earle & Baker 0 (ILL). Mobile Co.: Monroe Park, Mobile, Drushel
2577° (ILL, MO); 11.5 mi s of Mobile, Shinners 12579° (SMU); 0.6 mi n of Dog
River, Raven 22110° (DS); South Dog River at State Hwy. 163, Raven 22112° (DS).
Pickens Co.: Pickensville, Demaree 51291° ( DS, SMU). Pike Co.: Spring Hill,
Graves 1043a (MIN). Arkansas. Arkansas Co.: Nady P.O., Demaree 21657 (MO,
N^ ). Benton Co. Claytonville, Demaree 4678° (SMU); Siloan Springs, Demaree
22389 (MO, POM, SMU). Bradley Co.: Demaree 24639 (MO, SMU). Craighead
Co.: P. O. Jonesboro, Demaree 26552° ( RSA, SMU). Drew Co.: Lone Pine Prairie,
Demaree 21592 (ISC, MIN, NY, OKL, OKLA, SMU). Faulkner Co.: Demaree
10966 (GH, SMU, US). Madison Co.: 2 mi w of Cliffty, Hoff 210-4 (DS). Marion
Co.: 4.5 mi sw of Flippin, Raven 20559 (DS); Jimmie Creek, P.O. Bull Shoals,
Demaree 301/2° (RSA, SMU). Newton Co.: 1.4 mi n of Print, Raven 20574 (DS).
Pope Co.: P.O. Hector, Demaree 19898 (MO, NY, SMU). Pulaski Co.: Arkansas
River, Demaree 8741 (CM, IND, NY, US). Randolph Co.: Noland P. O., Demaree
31403 (ISC, RSA, SMU, UMO). Searcy Co.: 4.8 mi se of Marion Co. line, Raven
20560 (DS). Sebastian Co.: Fort Smith, Raven 20430 (DS). Stone Co.: P. O.
Optimus, Demaree 29929° ( KANU, ISC, MIN, RSA, SMU, TEX). Washington Co.:
Near Fayetteville, Isely 4782 (ISC, OKLA); Mt. Sequoyah, Fayetteville, Beck 4°
(OKL). Yell Co.: N of Centerville, Raven 20572 (DS). Colorado. Douglas Co.:
Sedalia, Ginter in 1942 (COCO). Connecticut. Hartford Co.: Southington,
Bissell in 1906° (GH). Illinois. Bond Co.: E of Smithbow, Evers 14334° (ILLS).
Cass Co.: N of Bluff Springs, Evers 21362° (ILLS). Clay Co.: 3 mi e of Flora,
Boewe in 1938 (ILLS). Coles Co.: Lincoln Log Cabin State Park, Voigt 959°
(MIN). Cook Co.: Des Plaines River, Leyden, Bennett in 1956° (ILLS). De Witt
Co.: Nw of Farmer City, Evers 31664° (ILLS). Edgar Co.: N of Chrisman, Evers
7425° (ILLS). Franklin Co.: 2 mi s of Sesser, Evers 52327° (ILLS). Hamilton
Co.: 1 mi e of Piopolis, Evers 13144° (ILLS). Henderson Co.: Near Oquawka,
Patterson ° (ND). Iroquois Co.: 1 mi w of Woodworth, Evers 6321° (ILLS). Jersey
Co.: 2 mi se of Jerseyville, Boewe in 1944° (ILLS). Kankakee Co.: Lansing ir
Sherff 14° (GH, MIN, PENN). Lawrence Co.: 8 mi w of Lawrenceville, Henderson
62-1070° (FSU). Logan Co.: 1 mi e of Burton View, Evers 5858° (ILLS). Marion
Co.: Nw of Kinmundy, Evers 81476° (DS). McHenry Co.: Rattray Hill near Al¬
gonquin, Nason in 1913° (F). Peoria Co.: 3 mi e of Edwards, Bates 2665° (DS).
Pike Co.: Nw corner of C. F. Gurhrie’s peach orchard, Campbell <L~ Alexopoulos 4404
(ILLS). Pope Co.: Ramsey Branch, Hopkins 1017 (SIU). Rock Island Co.: W of
Andalusia, Evers 31926° (ILLS). St. Clare Co.: East St. Louis, Eggert in 1896
(MIN). Sangamon Co.: 2 mi se of Springfield, Winterringer 22375° (DS). Taze¬
well Co.: E of Tremont, Evers 31763° (ILLS). Washington Co.: 2 mi n of New
Minden, Boewe in 1946° (ILLS). Winnebago Co.: Rockford, Puller 3048° (GH).
Indiana. Jasper Co.: Remington, Barnes 18 (GH); 3 mi n, 5 mi w of Remington.
Deam 42089 (IND). Newton Co.: 5 mi nw of Goodland, Deam 57322 (IND).
Vermillion Co.: 1 mi n of Rileysburg, Deam 51339 (IND). Wells Co.: W of
Murray Bridge along river, Deam 480 (US), iowa. Allamakee Co.: Yellow River,
* Specimens with glandular pubescence on the buds; all others were entirely strigulose.

MEMOIRS OF THE TORRE Y BOTANICAL CLUB

67

Pummel in 1927* (PH). Appanoose Co.: E of Moravia, Shimek in 1923 * (IA).
Black Hawk Co.: Bennington Township, Burk 729 * (ILL). Calhoun Co.: Carroll,
Pummel 23* (CINC, FLAS, GH, MO, NY). Clinton Co.: Clinton, Blake 53* ( ISC )’.
Crawford Co.: Butler in 1928 * (ISC). Davis Co.: 0.5 mi w of Bloomfield, Davidson
3715* (IA, SMU, US). Des Moines Co.: Davidson 3245* ( IA, TEX). Dubuque
Co.: Anderson in 1900* (IA). Emmet Co.: Wolden in 1925* (ISC). Fremont Co.:
Hamburg, Shimek in 1898* (IA). Hardin Co.: Near Cave in Rock, Bailey & Swayne
1668 (ISC). Greene Co.: Cratty in 1924 * (ISC). Lee Co.: Skunk River Valley,
Bartsch in 1895* (IA). Louisa Co.: 3/4 mi e of Wapello, Davidson 6* (IA, ISC,
SMU). Poweshiek Co.: 8 mi w of Grinnell, Russell 85556 * IA, TENN). Ringgold
Co.: 2 mi sw of Tingley, Fay 4494* (IA). Shelby Co.: Se of Harlan, Shimek in
1913* (ISC). Warren Co.: Indianola, Pammel in 1913 * (ISC). Webster Co.:
Olsson in 1903* ( ISC). Kansas. Butler Co.: 4.5 mi s of Leon, Stevens 6618 (KANU).
Chase Co.: 4 mi s of Matfield Green, Stephens 8706* (KANU). Chautauqua Co.:
1 mi w of Wauneta, Raven 20590* (DS); Niotaze, Raven 20587 (DS). Cherokee
Co.: Riverton Dam, Wagenknecht 2075 * (KANU); 5 mi e of Baxter Springs,
McCregoi 3/95 (KANU). Cowely Co.: Junction of Hwy. 166 with Hwy. 15, Raven
20590* (DS). Dickinson Co.: 3 mi ne of Detroit, Hoover 1239* (PAC). Doniphan
Co.: Agrelius in 1918* (KANU). Douglas Co.: 4 mi sw of Clinton, McGregor 12634
(US). Jefferson Co.: 2 mi n of Oskaloosa, Horr 4222* (KANU, NY, US). Linn Co.:
1 mi s of Parker, Richardson 6- Robertson 1036 (KANU, SMU). Montgomery Co.:
5.4 mi w of Hwy. 169 on Hwy. 166, Raven 20586* (DS). Morris Co.: Horr in 1930
(KANU), 4 mi e of Coimcil Grove, McGregor 14154 * (KANU). Nemaha Co.: 3.3 mi w
of Wetmore, Stephens 9449 * (KANU). Neosha Co.: Horr in 1929 (KANU). Republic
Co.: 2 mi s and 0.5 mi w of Munden, Morley 943 * (SMU). Scott Co.: Scott,
Thompson in 1936* (KANU). Shawnee Co.: 2.5 mi ne of Topeka, Voile 444 ( KANU)!
Washington Co.: 1 mi n and 3 mi w of Washington, Horr 4636 (KANU). Wood-
son Co.: 1 mi sw of Yates Center, Lathrop 1366 (MIN, NY); Latlirop 1469* ( RSA).
Kentucky. Calloway Co.: Spann in 1932 (KANU). Louisiana. Acadia Parish: 1.5
mi w of Midland, Raven 22093 (DS). Beauregard Parish: 2 mi e of De Ridder,
Ewan 20060 (DS). Boissier Parish: 7 mi n of Plain Dealing, Penn 951* (NO). Caddo
Parish: 6 mi n of Springridge, Raven 20383 (DS). East Baton Rouge Parish: Brown

7204* (GH). Evangeline Parish: 5 mi e of Beaver, Ewan 20019 (DS). Grant

Parish: 4.5 mi n of Dry Prong, Shinners 24941 (SMU). Jefferson Davis Parish:
Turn to Jennings off Interstate 10, Raven 22092 (DS). Lafourche Parish: 7 mi s of
Leeville, Thieret 25272 * (DS). La Salle Parish: 1.5 mi e-se of Tullos, Shinners
24730 (SMU). Ouachita Parish: 7 mi w of Monroe, Krai 8148 (FSU, LAF); banks
of Bartholomew Bayou, P.O. Bastrop, Demaree 14128 * (DS, GH, NY, OKLA,
POM, SMU). St. Landry Parish: U.S. Hwy. 190 4.6 mi w of State Hwy. 741,

Raven 22098 (DS). St. Mary Parish: Franklin, Langlois in 1884 (PENN). St.

Tammany Parish: Mandeville, Cooks in 1902 (NO); Slidell, Raven 22106 * (DS);
point s of Slidell at Lake Pontchartrain, Raven 22117* (DS). Tangipahoa Parish:
Hammond, Cooks in 1903 (NO). Vernon Parish: 1.3 mi n of Rosepine, Shinners
21593 (SMU). Washington Parish: Choctaw Trail, 3816 mi s of Pine, Ewan 19453*
(DS; glandular and with some long hairs on the sepals). Webster Parish: 2 mi s
of Doyline, Thieret 24344 (DS). West Feliciana Parish: Catalpa, Pennell 4297 *
(PENN). MARYLAND. Prince Georges Co.: Patuxent Wildlife Research Center,
Hotchkiss in 1964 * (DS). Massachusetts. Plymouth Co.: Wareham, Hunnewell
4912* (NEBS). Michigan. Jackson Co.: 7 mi s of Jackson, Raven 20508 * (DS).

05 A revision of tiie genus caura (onagraceae)

Monroe Co.: Pointe Mouillee State Game Area, mouth of Huron River, McDonald
5351° ( MSC). Minnesota: Without definite locality, Young in 1925 (CM).

MISSISSIPPI. Adams Co.: 6 mi e of Natchez, Gillis 5971 (MSC). Bolivar Co.: Shelton,
Rhoades in 1931 (GH). Clarke Co.: 3.2 mi s-sw of Pachuta, Skinners 27569 (SMU).
Covington Co.: 1 mi nw of Covington, Oosting 1964 ° (DUKE). Forest Co.: 11 mi
s of Hattiesburg, Ray in 1958 ° (USE). Hancock Co.: 7.5 mi s of Kiln, Jones 9498
(DS); Magnolia Ridge, Diener 436 ° (ILL); Bay St. Louis, Demaree 32986° (DUKE,
LAF, RSA, SMU, VDB); 4 mi s of Santa Rosa, S argeant 8330° ( OKLA, SMU,
WISC). Harrison Co.: Tracys Point, MacFarlane <b Goertz in 1905 (PENN). Hinds
Co.: Riverside Park, Dunaway in 1962 (BY). Jackson Co.: Ocean Springs, Pollard
1021° (CU, F, MIN, ND, NY), Seymour in 1891° (DUKE, FSU, GH, IA, ILL, K,
KANU, MIN, MSC, NCU, SMU, UMO, USF, WISC); Biloxi, Tracy 5071° CM, CU,
DAO, F, GH, KANU, MICH, MO, MSC, NCU, PENN, TAES). Jeff Davis’ Co.:
Near Carson, Jones 9867 ° (DS). Jefferson Co.: McDougaU 1080° (US). Lamar Co.:
5 mi sw of Hattiesburg, Ray 5954° (GA, GH, NCU, NY, USF). Leake or Madison
Co.: McDougall 1421° (US). Lowndes Co.: P.O. Columbus, Demaree 51280 ( DS,
KY, SMU). Montgomery Co.: P.O. Duck Hill, Demaree 31318° (KANU, OKl’
RSA, SMU). Oktibbeha Co.: Agricultural College, Pollard 1320 (MIN, NO, NY,
US). Smith/Scott County line: Road between Marathon Lake and Otho, Jones 2407
(FSU, GA, NCU). Warren Co.: Near Vicksburg, Schallert 13126° (SMU). Wayne
Co.: Chickawawhay River ca. 4 mi nw of Waynesboro, Jones 9665° (DS). Missouri.
Atchison Co.: 6 mi ne of Watson, Stcyermark 4451° (F). Barton Co.: 2 mi ne of
Milford, Palmer 53119° (UMO). Boone Co.: Hallsville, Daniels in 1897° (UMO).
Camden Co.: Zebra, Zirkle & Drouet in 1928° (UMO). Carter Co.: 2 mi w of Van
Buren, Munz 13589° (POM). Cass Co.: Near Pleasant Hill, Palmer 43804° (MO).
Clarke Co.: 2 mi w of Alexandria, Drones 1788° (GH, UMO). Davies Co.: 5 mi
sw of Gullatin, Evers 68081° (ILLS). Greene Co.: 8 mi w of Springfield, Munz
13553° (BY, F, NY); e of Fellows, Redfearn 3789 (LAF). Jasper Co.: Carthage,
Palmer 3147° (MIN); Prosperity, Palmer 15798 (UMO). Jefferson Co.: 2 mi w of

Hunter Ford, Steyermark 345 (MIN); Hillsbow, - in 1885° (MIN). Lafayette

Co.: Wellington, Rush 11519° (UMO). Lawrence Co.: 4 mi w of Chesapeake,
Raven 20580° (DS). Lincoln Co.: Whiteside, Davis 139 (MIN); near Whiteside,
Davis 6031 ° (MIN, MO). Madison Co.: Hwy. 67 at Wayne Co. line. Raven 20533°
(DS); Mine La Motte, Monell in 1890 (MO). Marion Co.: Hannibal, Davis 6146°
(MIN, MO). McDonald Co.: McNatt, Palmer 60875 (UMO); Butler Creek, Noel,
Palmer 4223° (US). Oregon Co.: Just n of Gregg, Raven 20552 (DS). Ozark Co.:
Near top of “Bald Jesse,” near Gainesville, Palmer 33-59° (MO, USF). Phelps
Co.: Little Piney River, Yancey Mills, Rickett in 1927° (UMO); Rolla, Coe in 1903
(GH). Reynolds Co.: Route M near Walker Branch School, Shacklette 6760° (MICH).
St. Genevieve Co.: Bloomsdale, Kellogg 1897 (MO). Shannon Co.: 9 mi w of Birch
Tree, Munz 13549 (POM). Stone Co.: 0.8 mi w of Taney Co. line on Hwy. 65,
Raven 20577° (DS). Taney Co.: Spencer 135 (UMO). Wayne Co.: Piedmont,
Raven 20546 (DS). Nebraska. Cass Co.: Weeping Water, Williams0 (US). Caster
Co.: Broken Bow, Webber in 1889° (NY). Douglas Co.: Omaha, Eastman in 1915°
(ILL). Gage Co.: Beatrice, Washburn in 1899° (US). Jefferson Co.: 1 1/4 mi s
of Hevley, Rolirbaugh 155° (OKL, TEX). Kearny Co.: Minden, Rydberg 546° (NY).
Lancaster Co.: Near Lincoln, Mulford in 1892° (MIN). Namaha Co.: Nemaha,
Winter in 1933° (US). Otoe Co.: 5 mi s, 1 mi e of Syracuse, Strjdiens 17610°
(DS, KANU). Saline Co.: Crete, Drcisbach 6038° (PH). Sioux Co.: Winter in

MEMOIRS OF THE TORREY BOTANICAL CLUB

69

1931 0 (CU). Thomas Co.: Middle Loup River near Seneca, Rydberg in 1893 0 (US).
ohio. Lorrain Co.: 2 mi nw of Oberlin, Cooperrider 8047 0 (KENT). Oklahoma.
Adair Co.: 3 mi w of Stilwell, Stephens 28027 ( DS, KANU); 25.3 mi ne of Tahle-
quahon, Wallis 2667 0 (OKLA). Bryan Co.: 0.5 mi e of Bokchito, Stephens 27575
( DS, KANU). Cherokee Co.: 3 mi ne of Fort Gibson, Wallis 8417 0 (KANU, NCU,
VDB). Cleveland Co.: Little River, Norman, Brewer 2150 (ISC). Comanche Co.:
Wichita Mts. Wildlife Refuge, McMurnj 1274 (OKL). Delaware Co.: 15 mi s of
Jay, Stephens 28137 (DS, KANU); Flint, Wallis 18780 (OKLA). Haskell Co.: 6 mi
vv, 1 mi s of Kinta, Stephens 27888 (DS, KANU). Kay Co.: 9 mi ne of Newkirk,
Btyler 337 (OKLA). McCurtain Co.: 10 mi n of Broken Bow, Stephens 27701
( DS KANU). Murray Co.: Back of Cowper Canyon, Conglomerate, Arbuckle Mts.,
Hopkins b Nelson 705 0 (DS, F, MO, OKLA, SMU). Oklahoma Co.: 4.5 mi e of
Arcadia, Wateifall 17790 (GH, OKL, PH). Osage Co.: 1 mi w, 5 mi n of Burbank,
Ste'fihens 27038 ( DS, KANU); 10 mi w of Pawhusta, Price 69 0 (OKLA). Ottawa
Co.: Near Miami, Stevens 2254 0 ( DS, CH, ILL, MIN, SMU). Payne Co.: 4 mi e
of Stillwater, Fry 490 (PH, POM, TEX). Seminole Co.: 6 mi se of Wowoka, Smith
114 0 (OKL). Sequoyah Co.: 1 1/4 mi se of Gore, Wallis 83320 (FSU, KANU,
OKL, VDB). Tulsa Co.: 5 mi n of Tulsa, Scully 51O0 (POM). Pennsylvania.
Montgomery Co.: Roslyn, Redles in 192O0 (PH; 92 percent stainable pollen). Tennessee.
Fayette Co.: S of Lakeland, Goolsby et al. 6677 (TENN). Hardeman Co.: 1 mi e of
Grand Junction, Rogers 40035 (TENN). Madison Co.: Bain in 1892 (US). Shelby
Co.: 0.5 mi n of Shelby, Gregory 558 (DS). Texas. Anderson Co.: Slocum, Hoff-
378 (DS). Angelina Co.: 7 mi w of Lufkin, Cory 10633 (POM). Aransas Co.:
N shores of Capano Bay, Johnston 53224.21 (MIN, OKLA, SMU, TEX). Austin Co.:
2.4 mi w of Belleville, Cory 17711 (POM). Bowie Co.: 5.3 mi n of Corley, Raven

20395 (DS). Brazoria Co.: 2.4 mi w of Alvin, Raven 20366 (DS). Brazos Co.:

College Station, Parks in 1946 ( RSA, TAES). Chambers Co.: 6.5 mi n of High
Island, Raven b Gregory 19420 (DS). Cherokee Co.: 4.6 mi e of Summerfield,
Shinners 19075 (SMU). Colorado Co.: 10.5 mi e of Alleyton, Cory 19505 (POM).
Dallas Co.: White Rock Lake, Longnecker 89 (some glandular pubescence; LL,
MICH, NY, SMU, TEX). Fort Bend Co.: Richmond, Palmer 6655 ( CU, F, MIN,
POM, US). Galveston Co.: Evergreen Ranch, Joar in 1884 (MIN). Hardin Co.:
Near Village Creek, Watson H-10 (LL). Harris Co.: Houston, Fisher 47O460 (SMU),
Traverse 217 ( F, CH, SMU). Henderson Co.: 3 mi ne of Frankston, Hoff 385
(DS). Hidalgo Co.: Donna, Parke 2942 (F). Hopkins Co.: Midland, Tharp 2435

(US). Houston Co.: 6 mi w of Crockett, Gregory 573 (DS). Hunt Co.: 6 mi sw of

Commerce, Turner 16770 (DAO, SMU). Lamar Co.: 3.7 mi s of Paris, Shinners
163280 (SMU). Leon Co.: Centerville, Gregory 574 (DS). Madison Co.: Trinity
Bottom, Tabenhaus 2639 (TAES, US). Morris Co.: 10 mi s of Daingerfield, Cory
25741 (POM). Nacogdoches Co.: Nacogdoches, Francis b Warnick in 1950 (SMU).
Newton Co.: 2 mi s of Newton, Cory 10906 (POM, TAES). San Augustine Co.:
3 mi n of San Augustine, Gregory 569 (DS). Shelby Co.: Timpson, Crausby in
1944 (TEX). Smith Co.: Amigo, Moore 1025 ( BH, GH). Upshur Co.: Ore City.
Shinners 16267 (SMU, TAES); 10 mi s of Daingerfield, Parks b Cory 25741 0 (TAES).
Van Zandt Co.: 15 mi n of Edgewood, Whitehouse 16443 (GA, MICH, SMU.
US). Victoria Co.: Mission Valley, Schott in 1851 (NY). Waller Co.: Waller,
Whitehouse in 1936 (TEX). Washington Co.: Brackett in 1939 (GH, TEX).
Wise Co.: Ferris b Duncan 3324 ( DS, NY). Wisconsin. Taylor Co.: Rib Lake
Goessl in 19150 (WISC).

70

A REVISION OF THE GENUS GAURA (ONACRACEAE)

In tlle ailtumn of 1832, while on a trip from Covington, Louisiana to
Columbia, Mississippi, Thomas Drummond gathered the specimens that
were to become the type material of Gama filipes Spach and G. longiflora
paeh in the vicinity of Jacksonville, Louisiana. This town no longer exists
but was located near present-day Sheridan, in Washington Parish/ Curious¬
ly, the locality has been confused in the present century with Jacksonville,
Florida (e.g., by Munz, Amer. J. Bot. 65: 117. 1938), and Gama longiflora
regarded as a synonym of G. angustifolia Michx., a species not known from
w est of Santa Rosa County, Florida. Even though Rothrock used the name
G. longiflora correctly in his monograph of the Gaurineae in 1864 (Proe.
Amer. Acad. Arts 6: 347-354), the much later name G. filiformis Small has
generally been associated with this species since 1900.

As partly documented by the map (fig. 56), Gama longiflora exhibits
a complex pattern of morphological features over a wide geographical area.
Fiom noithem Arkansas and eastern Oklahoma northward, populations of
this species usually consist of plants that have glandular pubescence on the
inflorescence, whereas most southern ones consist of plants in which the
pubescence is strigulose. In addition, northern plants tend to be more
coarsely branched and lower (often about 1-1.5 m tall), southern ones to
be virgate and often over 2 mm tall. Nevertheless, these differences are
recombined over a very broad zone of intergradation, and exceptions to
these geographical trends are numerous. For several years we intended to
separate the two series of populations as subspecies, but this no longer
seems desirable in view of the overall pattern of variation.

The northern populations of Gama longiflora were grouped by Munz
(Bull. Torrey Bot. Club 65: 105-122, 211-228. 1938) with G. biennis, a
self-compatible complex structural heterozygote that was probably derived
from similar populations of the self-incompatible G. longiflora , in which
bivalents or small rings of chromosomes are formed at meiotic metaphase I.
In G. longiflora, the pollen is usually more than 80 percent stainable, where¬
as in G. biennis, as is the case in most other complex structural hetero-
zygotes in Onagraceae, the pollen is only about 50 percent stainable
(usually 35-65 percent). The chromosomal complex that is not trans¬
mitted through the pollen is eliminated in this way.

In many areas, Gaura longiflora and G. biennis can be separated by
the long, villous pubescence on the main stems of the latter. Where their
ranges overlap broadly in Illinois and Iowa, and to a lesser extent eastward
(where G. longiflora becomes rare), the situation is more complex, and
phenotypic characteristics (except for pollen fertility) do not always
provide a reliable guide to the identity of a particular plant. These two
species share the same pollinators (mostly medium-sized to small moths).

MEMOIRS OF THE TORREY BOTANICAL CLUB

71

bloom at the same time of year, and can be hybridized readily in the ex¬
perimental garden to produce an F, generation that ranges from 30-90
percent pollen fertility. Undoubtedly they also hybridize where they come
together in the field.

Caura longiflora is a species of light prairie soils; its derivative G. biennis
occurs mainly in openings in the eastern deciduous woodland. With the
ad\ ('lit of European man, the habitats of both have been profoundly dis¬
turbed, and they have spread along roadsides to occur in vacant lots and
other waste places. Their populations are very probably much larger and
more extensive now than they were in the past. Their presumed hybrid¬
ization, as well as the broad zone of overlap that was a necessary pre¬
requisite for this hybridization, are probably phenomena of the last 150
years. Although considerable modification of the populations of the two
species by intiogressive hybridization seems to be going on at present, it
still appears best to reflect the profound genetic discontinuity between them
by recognizing them as distinct species.

In the southern portion of its range, Gaura longiflora forms hybrids
with a much more distinct species, G. lindheimeri. In eastern Texas and
the Gulf States, as we have noted above, most populations of G. longiflora
have appressed, strigulose pubescence on the inflorescences. Gaura lind¬
heimeri, a handsome species with long, spreading hairs on the inflorescences
and large flowers that open in the morning, is restricted to black clay
prairie soils in a relatively small area of southeastern Texas and south¬
western Louisiana, with disjunct stations in Tangipahoa Parish of eastern
Louisiana. Gaura longiflora occurs in the same area, but grows in light,
sandy soils as it does throughout its range. It has very probably entered
the area of distribution of G. lindheimeri to a large extent along roadside
banks and in other habitats that have resulted from disturbance bv man.

At some localities, the two species coexist without hybridizing. For
example, in the low prairie along U.S. Highway 90, 1.5 miles west of
Midland, Acadia Parish, Louisiana, in 1967, Gaura lindheimeri was abun¬
dant, whereas G. longiflora was represented by scattered plants along the
roadside and canalside diggings. No hybrids were observed. At other
localities, hybrids were uncommon. For example, obvious F, individuals
were seen in mixed populations as follows: 0.6 mi w of Manvel, Brazoria
Co., Texas, in 1965, Raven 20371 (one plant with many of both parents,
at the edge of a rich, boggy pasture; pollen stainability 39 percent); edge
of pine woods along U.S. Highway 59, 4.8 mi s of Humble, Harris Co.,
Texas, 1965, Raven 20375 (two hybrids, with many plants of the parental
species; one had 1 chain of 4, 1 chain of 6, and 2 pairs, with a pollen stain-
ability of 55 percent; the other had 68 percent); large field at east limits

A REVISION OF THE GENUS GAURA (ONAGRACEAE)

of Areola, Fort Bend Co., Texas, 1965, Raven 20365. In the last locality,
there seemed to be about one hybrid individual per 10,000 of the parents.
These were easily seen owing to their combination of the height of G.
longiflora, nearly 2 m, with the basal, virgate branching pattern of G. lind-
heimeri. One of them had a chain of 10 chromosomes and 2 ring bivalents,
with a pollen stainability of 63 percent; another had a chain of 14 chromo¬
somes, with a pollen stainability of 40 percent; and a third had a pollen
stainability of 37 percent. The species could easily be hybridized in the
experimental garden, with about 75 percent capsule set, and their hybrids
had a cytology compatible with that reported above and a fertility that
ranged from 13-76 percent (40-70 percent in most hybrid individuals).

In addition to their gross ecology, there are at least two other factors
tending to restrict hybridization between these two species. Gaura lind¬
heimeri blooms earlier, from May to early July, for the most part, whereas
Gaura longiflora blooms from July to late summer. Thus their period of
overlap is restricted. In addition, and in line with their different daily times
of bloom, they are pollinated by different insects: medium-sized to small
moths in G. longiflora, bees in G. lindheimeri. Although bees occasionally
visit the withered flowers of G. longiflora the morning after they have first
opened (and may then carry pollen from one of these species to the other),
the moths probably never visit flowers of G. lindheimeri.

Despite these barriers, populations are also encountered which seem
to consist largely or entirely of hybrids. For instance, 2.5 mi n of Talisheek,
St. Tammany Parish, Louisiana, in 1967, an extremely variable population
( Raven 2210. 5) was encountered that appeared referable to G. longiflora,
but with individual plants having variable amounts of the long spreading
pubescence of G. lindheimeri. Even more direct evidence of past hybrid¬
ization is provided by a series of populations that occur in eastern Louisiana
and southern Alabama which resemble G. longiflora but have large flowers
that open in the morning. Populations of this nature were studied in
southern Mobile County, Alabama, where they are the only representatives
of the genus Gaura: abundant along edge of salt marsh near Cedar Point,
2 mi s of turn to Bayou la Batre on State Hwy. 163, Raven 22116; 4 mi s,
Raven 22115; Hildebrand 65 (TEX); Dauphin Island, Deramus D.1074
(DS); State Hwy, 163 at South Deer River, Raven 22112. They likewise
occur in Orleans Parish, Louisiana, where they may have had an inde-
pendent origin: Ft. Pike at the Rigolets, along U.S. Hwy. 90 Ewan 221H2
(DS); edge of marsh among tall grasses and Baccharis, between Rigolets
and mouth of Pearl River, 7 mi s of junction of State Hwy. 433 on U.S.
Hwy. 90, Chef Menteur Hwy., Ewan 22170 (DS), Lemaire 95 7 (NO),
Raven. 22107. The pollen stainability of three individuals from the last-

MEMOIRS OF THE TORREY BOTANICAL CLUB

73

mentioned locality was studied, with the following results: 99 percent, 44
percent, and 74 percent. Some of the individuals resembled one or the
other parent closely, and there was considerable variability. Similar plants
were once collected "on shell islands near the Gulf, Plaquemines Parish,”
Louisiana, Aug. 1879, Langlois 97 (R, NY, PENN), and were confused by
Munz (Bull. Torrey Bot. Club 65: 217. 1938) with G. filipes. Another
collection is from Jefferson Parish, Louisiana, Langlois in 1884 (M).

Certain collections from Texas, from populations that we have not
studied in the field, are morphologically similar and may also represent
morning-opening plants of the derivation just discussed. Examples are:
Texas. Aransas Co.: Goose Island, shell beach, Whitehouse 18302 (NY).
Galveston Co.: Near Galveston, Perkins & Hall 2458 (POM). Harris Co.:
Morgans Point, Cory 8140 (POM). Nueces Co.: Corpus Christi, Picken-
pangh in 1936 (POM). San Patricio Co.: Ransom Island, Parks 21272
(POM).

It might be argued that this series of large-flowered, relatively glabrous,
morning-opening plants ought to be recognized as a distinct species, and
we considered this possibility for several years. The chief reasons that we
decided not to adopt this course of action were 1). populations of this
nature may well have originated more than once along the Gulf Coast, and
further study might well reveal significant differences between them; 2.
even locally, the situation is extremely dynamic and the populations are
highly variable, a condition that does not lend itself well to formal classifi¬
cation; 3. as indicated by the discussion above, plants of this nature are
very difficult to identify except from living plants, and it is therefore im¬
possible at this point to give an accurate idea of their distribution in
nature; 4. they are connected with typical populations of G. longiflora by
a series of intermediate populations without any abrupt morphological gaps.
For example, a population that was growing along the borders of the salt
marsh south of Slidell, St. Tammany Parish, Louisiana, in 1967 ( Raven
22117), was similar in aspect and pubescence to the population we studied
between Rigolets and the mouth of the Pearl River, farther east, but con¬
sisted of plants in which the petals were only 10-12 mm long (11-17 mm
in the Rigolets population) and the flowers opened in the evening. Along
the highway 0.6 mi n of Dog River, Mobile Co., Alabama, at the margins
of the woods, we studied a population ( Raven 22110 ) in which the petals
were 11-16 mm long, but were not able to determine whether the flowers
opened in the morning, as in the more glabrous plants found along the
borders of the salt marshes farther south, or in the evening.

As may be seen from the map (fig. 56), a number of the populations
in the southern part of the range of G. longiflora consist of plants that have

74

A REVISION OF THE GENUS CAURA ( ONACRACEAE )

glandular pubescence (and rarely also longer hairs) in their inflorescences,
n many cases these plants, which are similar to those just discussed from
og River. Alabama have probably resulted from introgression of Gaum

" , ?■ lon£‘fl°ra- They were treated by Munz (Bull. Torrev

Bot. Club 65: 18-120. 1938) as G. filiform* Small var. kearneui Munz.

ey occur considerably beyond the present range of G. lindheimeri, hut
It is likely, ,n view of the pattern reported here and considering the iso¬
lated stations for G. lindheimeri in eastern Louisiana, that that species

once had a wider range, perhaps accompanied by a greater ecological
amplitude. b

16. Gaura biennis L„ Sp. PI. .347. 1753. Munz, Bull. Torrev Bot. Club
65: 111. 1938; N. Amer. FI. II. 5: 185. 1965; pro parte. Figs. 20. 53.

Similar to G. longiftora in all except the following characteristics: plants
up to IS dm tall; stems conspicuously and densely villous, rarelv short-
vi ous, the leaves short-villous along the veins and margins; inflorescence
Ml ous, also densely glandular pubescent; plants entirely lacking strigulose
pubescence; bracts of inflorescence 1-2.25 mm long, 0.5-1 mm wide; floral
tube 6-12,5 mm long; sepals 8-1.3 mm long, 1-2 mm wide; petals 6 5-12
mm long, 2-6 mm wide; filaments 5-10 mm long; anthers 2-4 mm long-
pollen fertility ca. 50 percent; style 1.3.5 mm long; fruit 5-9 mm long
2-3 mm thick; seeds 3-6, 2-2.5 mm long, 1-1.25 mm thick, light to reddish
brown. Self-compatible, often self-pollinating. Gametic chromosome num-
ber, n = 7 (ring of 14 at meiotic metaphase I).

Lectotype: Cultivated in Uppsala, Sweden, prior to 1751, from seeds
from eastern Pennsylvania collected by Peter Kalm, 1748-1750. C. Linnaeus
(LINN 485.1). Linnaeus’ reference 'to “Virginia” in his protologue con¬
cerns his reference to Plukenet, who was discussing some member of
Oenothera subgenus Oenothera.

Distribution (fig. 55): Clearings in woodland and waste places, eastern
Iowa, southeastemmost Minnesota, southern Wisconsin, southern Michigan
southernmost Ontario, southern New York, and Massachusetts, south to
\ irginia, western North Carolina, central and eastern Tennessee, and Il¬
linois. Flowers July to September.

Representative Specimens Examined:

Canada. Ontario. Essex Co.: Windsor, McCrae 7845 (NMC); Amsterburjjh,
Macoun 44459 (CM, GH, NMC); Leamington district, Eroh in 1924 (DAO). Lin¬
coln Co.: Queenston, Scott 20506 (NMC). Welland Co.: Fort Erie, Sharp 4 (DAO);
Erie Beach, Bertie Twp., Miller in 1954 (DAO).

United States. Connecticut. Fairfield Co.: Oxford, Harder 6 Weatherby 2287
(NSC). Hartford Co.: Bristol, Bishop in 1897 (GH). New Haven Co.: Eaton

MEMOIRS OF THE TORREY BOTANICAL CLUB

75

(GH, POM). Delaware. New Castle Co.: Holly Oak, near Wilmington, Canby in
1878 ( LE, NY), district of columhia. Washington, Ward in 1882 (GH). Illinois.
Adams Co.: S of Burton, Evers 89613 (ILLS). Carroll Co.: Palisades State Park,
Hartley 9931 (IA). Champaign Co.: Sangamon River near Mahomet, Jones 12717
( GH, ILL). Cook Co.: Chicago, Greenman in 1876 (MO). Dupage Co.: Naperville,
Umbach in 1897 (MIN). Grundy Co.: 1.5 mi s of Morris, Evers 6647 (ILLS).
Hancock Co.: Augusta, Mead (GH). Jo Davies Co.: W of Galena, Weber 1953
(COLO). Kane Co.: Vicinity of Elgin, Gronemann 101 (F). Kankakee Co.: 7 mi
nvv of Kankakee, Sherff 1625 (MIN). La Salle Co.: Utica, Clemens in 1909 (POM).
Marshall Co.: S of Jacon, Evers 6232 (ILLS). McHenry Co.: Algonquin, Nason in
1878 (ILLS). Menard Co.: Hall in 1861 (NY). Ogle Co.: Cratty in 1877 (ISC). Peoria
Co.: Near Tubercular Santarium, Chase 13011 (DAO, NY, OKL). Platt Co.: Deland,
Seymour in 1889 (DUKE). Richmond Co.: 3.5 mi n of Olney, Scherer 449 (ILL). Rock
Island Co.: N of Reynolds, Evers 6988 (ILLS). St. Clair Co.: Eggert in 1877 (MO).
Sangamon Co.: 0.5 mi e of Curran, Winterringer 22374 (DS). Tazewell Co., Near
East Peoria, Chase 3286 (BH, ILL, MIN, PAC, PH, POM). Vermillion Co.: Between
Oakwood and Collison, Jones 13045 (SMU). Whiteside Co.: 3 mi w of Garden
Plain, Evers 6893 (ILLS). Will Co.: Romeo, Umbach 1991 MICH). Winnebago Co.:
Bebb in 1858 (PH). Indiana. Adams Co.: 2 mi nw of Decatur, Deam 5315 (IND).
Fayette Co.: 1 mi s of Connersville, Buser 1643 (ILL). Franklin Co.: 1/8 mi n
of Brookville, Deam in 1903 (IND). Grant Co.: 1/4 mi nw of Matthews, Deam
49439 (IND). Harrison Co.: State Forest, Friesner 19579 (DAO, FLAS, KANU,
K\ ). Howard Co.: 4 mi n of Kokomo, Ek 42 (US). Knox Co.: 3 mi sw of Vincennes,
Deam 29302 (IND). Lake Co.: Dyer, Peattie 2026 (F). Marion Co.: Indianapolis’
Friesner 7892 (DUKE, PENN, POM, SMU, TENN, TEX, WVA). Perry Co.: 2 mi
s of Oriole, Deam 47991 (IND). Ripley Co.: About 3 mi ne of Morris, Deam 58094
(IND). Steuben Co.: Near Lake James, Blatcliley in 1902 (IND). Tippecanoe Co.:
3 mi nw of Westpoint, Deam 54407 (IND). Vermillion Co.: About 0.5 mi e of
Dana, Deam 54372 (WVA). White Co.: 2 mi e of Seafield, Deam 15225 (IND).
iowa. Allamakee Co.: Yellow River, Pommel in 1927 (ISC, OKLA). Benton Co.:
Vinton, Davis 6 (US). Clinton Co.: Elk River Township, Cooperrider 2708 (MIN).
Dallas Co.: Union Township, Van Bruggen 906 (SMU). Guthrie Co.: 4 mi n of
Panora, Hayden 3633 (ISC). Jackson Co.: Union Township, Cooperrider 2610
(IA). Johnson Co.: S of Iowa City, Shimek in 1915 (ISC). Jones Co.: Clay Town¬
ship, Brown in 1948 (ISC). Lee Co.: Skunk River Valley, Bartsch in 1894 (ISC).
Linn Co.: Cedar Rapids, Buchanan in 1902 (ISC). Makaska Co.: 3 mi nw of Fre¬
mont, Rohrbaugh 99 (COLO, TEX). Muscatine Co.: 8 mi nw of Muscatine, Shimek
in 1925 (SMU, USF). Poweshiek Co.: Grinnell, Jones (POM). Scott Co.: Parry
(ISC). Van Buren Co.: 4 mi e of Willets, Shimek in 1932 (ISC). Washington Co.:

3 mi s of Wellman Fine Creek, Wagenknecht 1239 (SMU). Webster Co.: Fort
Dodge, Somes in 1906 (ISC). Kentucky. Anderson Co.: Floodplain of Gilbert
Creek, Wharton 9387 (KY). Bullitt Co.: Wilson Creek Road, Gunn 841 (KY).
Clark Co.: 3 mi s of Indian Fields, Wharton 5104 ( KY, USF). Fayette Co.: Lexing¬
ton, Short (KY). Grayson Co.: Leitchfield, Rose in 1895 (US). Jefferson Co.:
Jennings Lane, Gunn 2265 (KY). Jessamine Co.: Wilmore, Anderson 585 (GH,
IA). Mercer Co.: Burgin, Kind 232 (F). Nelson Co.: Agnes (IA). Robertson Co.-
Blue Licks State Park, Smith 1748 (KY). Woodford Co.: Near Hifner Road, Wharton
8660 (KY). Maryland. Alleghany Co.: Wills Creek, Shreve 6- Jones 995 (US).
Anne Arundel Co.: Near Portland, Otis in 1916 (PH). Baltimore Co.: Powhatan!

A revision of the genus gaura (onacraceae)

Plett 866 (GH). Cecil Co.: Porters Bridge, Octararo Creek, Long 28503 (GH, PH).
Frederick Co.: Emmitsburgh, Aikin (PH). Harford Co.: Shores Landing, Leeds 2056
(PH). Montgomery Co.: Great Potomac Falls, Schott in 1858 (F). Washington
Co.: Potomac River near Hancock, Rose in 1910 (US). Massachusetts. Berk¬
shire Co.: Hoffman in 1940 ( NEBC). Bristol Co.: Ware 4163 ( NEBC). Middlesex
Co.: Morong in 1886 (NY). Plymouth Co.: Knowlton in 1925 (NEBC, USF).
Suffolk Co.: Nepomset, Knowlton in 1935 (IND, NEBC). Michigan. Berrien Co.:
Benton Harbor, E.B.M. in 1891 (F). Houghton Co.: W.W.C. (MICH). Kent Co.:
Plaster Creek s of Grand Rapids, Skul in 1895 ( MSC, NMC). Monroe Co.: Pointe
Mouillee State Game Area, McDonald 5487 (MSC). Oakland Co.: Oakwood, Sutton
in 1914 (MICH). Wayne Co.: West Detroit, Chandler in 1941 (MSC). Minnesota.
Houston Co.: Jefferson, Wheeler 574 (MIN), new jersey. Burlington Co.: Borden-
town, Long In 97 (PH). Camden Co.: Camden, Boice in 1888 (PH). Hunterdon
Co.: Near Stanton Station, Benner 6873 (PH). Mercer Co.: Princeton, Peters in
1885 (NY). Middlesex Co.: Along Raritan Road, McKenzie 2863 (IND, MO, US).
Morris Co.: Whippany, Moldenke 21675 (OKLA, SMU). Passaic Co.: Vicinity of
Clifton, Nash in 1891 (MO). Warren Co.: 1 1/4 mi nw of Broadway, Schaeffer
29978 (PH), new york. Albany Co.: N end of Van Schaick Island, Burnham in
1911 (GH). Bronx Co.: Van Cortland Park, Monachino 655 (NY). Broome Co.:
Front Street swamp, Millspaugh in 1886 (F). Cattaraugus Co.: Near Irving, Smith
in 1944 (MO). Chemung Co.: Ashland, Lucy 728 (NY). Erie Co.: Buffalo, Clinton
(MICH). Herkimer Co.: Bank of Mohawk River at Little Falls, Burnham in 1905
(GH). Madison Co.: Vicinity of Hamilton, Longworthy 51 (CU). Niagara Co.:
Goat Island, Johnson in 1927 (US). Rensselaer Co.: Troy, Hall in 1828-1834 (F).
Rockland Co.: Nyack, Aides 842/48 (ILL). Schenectady Co.: Bank of Mohawk
River, Foxworthy in 1902 (CU, NY). Schuyler Co.: Watkins Glen, Schuster A-6923
(DUKE). Seneca Co.: Seneca Lake, Sartwell (NY). Steuben Co.: 1 mi n of Bath,
Clausen ir Trapido 2706 ( BH, CU). Tioga Co.: MiUspaugh 189 (F). Tompkins
Co.: Renwick Park, Eames ir MacDaniels 4634 (CU, GH, IND). Tompkins Co.:
Ithaca, Munz 767 (POM). Yates Co.: Dundee, Wright (POM), north Carolina.
Alleghany Co.: 0.9 mi n of Twin Oaks, Radford 38403 (NCU). Cherokee Co.:
2 mi e of Murphy, Radford 17479 (GH, NCU). Davidson Co.: Yadkin River bottom
land, near Yadkin College, Totten in 1926 (NCU). Haywood Co.: 0.5 mi s of
Sunburst, Aides ir Duke 46679 (COLO, MIN, NCU, RSA). Jackson Co.: Near
Tuckaseegee, Godfrey 52025 (GA, NSC, PI I, TENN). Macon Co.: Cartoogechaye,
Johnson 894 (F). Mitchell Co.: 2.3 mi ne of Huntdale, Aides ir Duke 47237 (NCU).
Person Co.: 2 mi e of Woodsdale, Bell 14647 ( KY, NCU). Rockingham Co.: Spray,
De Chalmot (US). Surry Co.: Yadkin River near Elkin, Radford 18379 (FSU, GA,
NCU, USF). Swain Co.: Cherokee, Munz 13522 (POM), ohio. Coshocton Co.:
Near Coshocton, Moldenke 13378 (CM, NSC, OKLA, VDB). Cuyahoga Co.:
Cleveland, Ashcroft in 1897 (MO). Erie Co.: Vermillion River e of Birmingham,
Cooperrider 7966 (KENT). Franklin Co.: Reynoldsburg, Raven 18733 (DS). Green
Co.: Yellow Springs, Demaree 11541 (MIN, MO, NY, OSU, SMU, US). Hocking
Co.: Laurelville, Demaree 11592 ( DS, MU, SMU). Lucas Co.: Toledo, Young in
1884 (NY). Muskingum Co.: 3 mi s of Adamsville, Bilker in 1957 (CM). Pickaway
Co.: Pickaway Township, Cooperrider ct al. 9415 (DS). Richland Co.: Mansfield,
Wilkinson 3425 (F, ISC, MICH, MIN, MSC). Ross Co.: 2 mi sw of Kingston,
Cooperrider et al. 9396 ( DS, KENT). Wayne Co.: Wooster, Duvel 833 (NY).
Pennsylvania. Allegheny Co.: Harmarville, Van Dcrsal 557 (NY). Bedford Co.:

MEMOIRS OF THE TORREY BOTANICAL CLUB

77

Bedford Springs, Berkheimer 1363 (PENN). Bucks Co.: 2 mi above Yardley,
Meredith in 1923 (GH). Center Co.: 2 mi w of Spring Mills, Wahl 339 ( FLAS,
ISC, C.H, PAC, PENN). Columbia Co.: Bersick, Small in 1889 (F). Delaware Co.:
Tenni, Pennell in 1906 (PH). Greene Co.: Monongahela River near Marion, Bright
7651 (MIN). Juniata Co.: Open banks above Juniata River, Fogg 15546 (GH,
PENN). Lehigh Co.: 1 mi n of Lowhill, Schaffer 32317 (PH). Mercer Co.: Sharon,
Aschman in 1886 (PH). Montgomery Co.: Perkiomen Creek, Long in 1908 (GH).
Perry Co.: 5 mi s of Liverpool, Munz 13408 (POM). Pike Co.: 2.5 mi ne of Mil¬
ford, Fogg 19229 (PENN). Union Co.: 0.5 mi se of Millmont, Westerfield in 1949
(PAC). Westmoreland Co.: Ligonier, Gress 766 (PENN). Tennessee. Blount Co.:
Little Tennessee River above Chilhowee Dam, Sharp <Lr Veloira 21000 (TENN).
Campbell Co.: Near Chaska, McFarland 4863 (IND, POM). Carter Co.: Elizabethton,
Wiegand 6- Manning 2162 (CU, GH). Cheatham Co.: Craggie Hope, Svens on 328
(GH). Coffee Co.: Between Manchester and Tullahoma, Woodruff in 1941 (TENN).
Hamblen Co.: Morristown, Smith in 1880 (US). Hancock Co.: Clinch River bank
near Narrows, Kyles Ford, Sharp 3074 (IND, TENN). Knox Co.: Knoxville, Ruth
2782 (NY). Loudon Co.: Clinch River at corner of county, Hatcher 29803 ( GA,
TENN). Roane Co.: White Oak Lake, Nease in 1950 (TENN, US). Rutherford
Co.: N of Lavergne, Eggert in 1897 (MO). Sevier Co.: Near Sevierville, Rugel in
1842 (DAO, RSA). White Co.: Caney Fork near Webbs Camp, below Grand Falls
Dam, Sharp et al. 5030 ( KY, TENN). Virginia. Augusta Co.: Near Augusta Springs,
Steele in 1908 (US). Bath Co.: Hot Springs Valley, Steele 6- Whealer in 1907
(US). Bedford Co.: Blue Ridge Parkway, Freer 1844 (GH). Charlotte Co.: 2.1
mi s of Harrisburg, Ahles 6- James 61658 (NCU). Fairfax Co.: Great F alls of the
Potomac, Ball 263 (ISC). Fauquier Co.: W slope of Bull Run Mt;., Allard 5462
(GH). Frederick Co.: Middletown, Hunnerwell 11338 (GH). Giles Co.: Near
Newport, Thorne 17847 (FSU). Grayson Co.: 3.8 mi sw of Elk Creek, Ahles i?
Tucker 60286 (NCU). Halifax Co.: Just n of Halifax, Ahle, 6- James 60561 (NCU).
Henrico Co.: Three Shopt Road, Canada 2074 (GH). Isle of Wight Co.: W of old
Fort Bavkin, James River, Fernald 6- Long 13709 (GH). Loudoun Co.: 1.3 mi nw
of junction of U.S. Hwy. 15 with Co. Route 672, Ahles 6- James 61316 (NCU).
Madison Co.: 4 mi se of Marksville, Fosherg 36118 (GH, US). Montgomery Co.:
2 mi nw Blacksburg, Krai 11468 (USF). Orange Co.: Gordonsville, Blanchard in
1890 (MO). Page Co.: Luray, Heritage in 1887 (PENN). Pittsylvania Co.: Staunton
River on County Route 668, Ahles 6- James 62470 (NCU). Prince Edward Co.:
Appomattox River at f armville, Ahles Is James 62937 (NCU). Prince George Co.:
City Point, Fernald Is Long 11094 (GH). Prince William Co.: E slope of Bull Run
Mts., Allard 8171 (GH). Roanoke Co.: Wood 5308 (GH, PENN, PH). Rockbridge
Co.: Near Lexington, Churchill in 1924 (C.H). Rockingham Co.: Mt. Crawford,
Heller is Halback in 1893 (CU, DAO, DS, F, C.H, MO, PENN, PH). Smyth Co.:
Marion, Small in 1892 (MO). Surry Co.: Claremont, Fernald t? Long 13708 ( C.H,
US). Warren Co.: Front Royal, Oldfield, Sargent in 1949 (NY). Washington Co.:
Wallace Switch, Small in 1892 (NY), west Virginia. Brooke Co.: 3 mi above Bethany,
Bartholomew B193t -15 (WVA). Jefferson Co.: Charleston, Harper in 1894 (POM).
Marshall Co.: Near Majorsville, Bartholomew 85 (WVA). McDowell Co.: Red
Chesnut Mt., Music in 1961 (WVA). Mercer Co.: Shawnee Lake, Core 1860 (DS,
WVA). Mineral Co.: Burlington, Core 3333 (NY, WVA). Monongalia Co.: Valley
Crossing, Morgantown and Kingwood Railroad, Sheldon in 1918 (CM). Morgan
Co.: Near Berkeley Springs, Alexander et al. in 1933 (NY). Randolph Co.: Above

78

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Huttons ville, Greenman 167 (GH). Tyler Co.: Davenport, Barkley 957 (MO). Wirt
Co.: 1 mi e of Elizabeth, Bartholomew in 1933 (WVA). Wisconsin. Dane Co.:
Mazomanie to Sauk City, Greene in 1948 (WISC). Grant Co.: Opposite Dubuque
Iowa, Fassett 13338 (WISC). Green Co.: 8 mi w of New Clams, Shinners ir
Catenhusen 4608 (WISC). Iowa Co.: 1.5 mi e of Arena, Mason 1371 (WISC).
Lafayette Co.: Wagner in 1959 (WISC). Milwaukee Co.: Milwaukee, Shinners &
Stearns 4131 (GH, WISC). Outgamie Co.: Binghampton Bog, Melchert in 1957

(WISC). Rock Co.: Beloit, Shinners 4642 (WISC). Walworth Co.: Wadmoncl 2605
(WISC).

The type specimen, like all other cultivated material from Europe, has
about 50 percent sterile pollen and is referable to the complex structural
heterozygote. G aura biennis has been in cultivation in the botanical gardens
of Europe and, later, North America, since the mid-18th Century. For
example, we have seen a specimen collected in the botanical garden at
Leningrad in 1779, from seeds received in 1771 (LE).

17. Gaura demareei Raven & Gregory, sp. nov. Figs. 21, 54.

A G. longifiora differt: calibus inflorescentiis dense cinereo-strigulosis
eglandulosis; bracteis 3-10 mm longis, 0.5-1.5 mm latis; floribus diurnalibus;
tubo floris 4-13 mm longo; sepalis 13-20 mm longis, 1-2.5 mm latis; petalis
(10— ) 11-17 mm longis, 4. 5-8. 5 mm latis; filamentis 8—17 mm longis; anthe-
ribus 3-7 mm longis; stylo 18-31 mm longo.

Similar to G. longifiora in all except the following characteristics: stems
and inflorescences densely gray-strigulose, with no spreading or glandular
hairs; bracts of inflorescence 3-10 mm long, 0.5-1. 5 mm wide; flowers
opening near sunrise; floral tube 4-13 mm long; sepals 13-20 mm long;
1-2.5 mm wide; petals ( 10— ) 11—17 mm long, 4.5-S.5 mm wide; filaments
8-17 mm long; anthers 3-7 mm long; style 18-31 mm long. Self-incom¬
patible. Gametic chromosome number, n = 7.

Types 4.6 mi ne of Hot Springs on U.S. Highway 270, Garland Co.,
Arkansas, 3 August 1965, P. II. Raven 20435 (DS 574399. Isotvpes, BM,
GH, NY, RSA, US).

Distribution (fig. 56): Open meadows in sandy loam, southwestern
Arkansas. Flowers July to October.

Specimens Examined:

United States. Arkansas. Chalybete Mt., Demaree 54824 (DS); P.O. Arkadelphia,
Demaree 15639 (MO, NY, POM), 40476 (G, RSA, SMU). Garland Co.: Hot
Springs, Beer in 1947 (CM); Hot Springs National Park, Demaree 49027 (DS),
53971 A (DS); 6 mi n of Hot Springs, Scully 377 (POM); Crystal Springs, Demaree
54256 (DS), 54255 (I)S); Hot Springs National Park. Demaree 53971 A (DS);
Mountain Valley P.O., Demaree 37652 (KANU, OKL, SMU); Eggert in 1896 (MO);
2.5 mi w of Hwy. 5, Haven 20570 (DS); Hot Springs Mts., Runyon 2275 ( BH, US).

MEMOIRS OF THE TORREY BOTANICAL CLUB

79

Hempstead Co.: Bingen, Demaree 54098 (DS); McNab, Demaree 44670 (DS);
P.O. Ozan, Demaree 15966 (MIN, MO), 40517 ( NCU, SMU); P.O. Magnet Cove,
Demaree 17462 (SMU). Howard Co.: P.O. Nashville, Demaree 9858 (MO). Mont¬
gomery Co.: Caddo River, Demaree 9583 (US); Mt. Ida, Demaree 54266 (DS);
South Fork of Ouachita River, Demaree 44872 ( DS, RSA). Pike Co.: Delight,
Demaree 54119 (DS). Saline Co.: 3.7 mi s of Paron, Raven 20568 (DS); 1.5 mi w
of Owensville on Hwy. 5, Raven 20569 (DS). Sevier Co.: P.O. Ben Lomond,
Demaree 41625 (NCU).

It is a great pleasure to name this attractive, large-flowered, morning-
opening derivative of Gaura longiflora in honor of Dr. Delzie Demaree
(1889- ), whose active botanical work in Arkansas and in other states

is attested by his many collections of this local species cited above. Dr.
Demaree’s collections are best represented in the herbarium of Southern
Methodist University, but large sets are deposited in many other institutions.

SECTION VIII. PTEROGAURA
Gaura sect. Pterogaura Raven & Gregory, sect. nov.

Heibae annua infra inflorescentiam villosae. Inflorescentiae erectae.
Flores valde zygomorphi, vespertini. Fructus ellipsoidalis vel ovoideus,
late alatus.

Annual herbs, villous below the inflorescence. Inflorescence erect.
Flowers strongly zygomorphic, the stamens in the lower quarter; flowers
opening near sunset and withering the next morning. Fruit not deflexing,
ellipsoid or ovoid, broadly winged on the 4 (or 3) angles and deeply
furrowed between them, sessile or short-stipitate and with pronounced
comers at the bottom or these absent. Self -incompatible or self-com¬
patible. Gametic chromosome number, n = 7; one species (G. triangulata)
a complex structural heterozygote.

Type Species: Gaura suffulta Engehn.

Distribution: Sandy flats and waste places, or as a weed of cultivated
fields, from Arizona to Guatemala, west through Oklahoma and Texas to
southern Mississippi.

This highly specialized section consists of four species, one— Gaura
stiff ulta-se lf-incompatible, another-G. bracliycarpa-se lf-compatible but
outcrossing. Of the remaining two highly autogamous species, G. hexandra ,
which ranges throughout Arizona, New Mexico, and western Mexico to
Guatemala, is a derivative of G. suffulta subspecies nealleyi. The remaining
species, G. triangulata, which is found mainly in north-central Texas and
south-central Oklahoma, is a complex structural heterozygote probably de¬
rived from hybrids between G. suffulta subspecies suffulta and G. brachy-

80

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

carpa, between which it is intermediate in its morphological character¬
istics.

Some plants of all species of this section, except the self-incompatible
G. suffulta, and most plants of G. triangulata have 3-merous flowers, as is
often also the case in G. angustifolia. As discussed earlier (Raven, Contr.
U.S. Natl. Herb. 37: 376-377. 1969), such a reduction in number of floral
parts is apparently closely correlated with self-pollination in the family
Onagraceae. As circumscribed by Munz (Bull. Torrey Bot. Club 65: 211-
213. 1938; and with increasing doubts in N. Amer. FI. II. 5: 191. 1965),
Gaura hexandra ( G . tripetala Cav. ) included as subspecies elements here
treated as G. hexandra, G. hrachycarpa, and G. triangulata which have
independently attained the 3-merous condition in connection with self-
pollination.

18. Gaura suffulta Engelm. ex Gray, Bost. J. Nat. Hist. 6: 190. 1850.

Munz, Bull. Torrey Bot. Club 65: 120. 1938; N. Amer. FI. II. 5: 189.

1965.

Annual herb from a stout taproot; stems usually well branched, 2-12
dm tall. Plants villous (with hairs 1.5-2. 5 mm long) below the inflorescence,
the leaves subglabrous or sparsely villous along the veins and around the
margins; inflorescence glabrous or glandular-pubescent, especially on the
outside of the floral tube and the sepals. Rosette leaves lyrate, 3.5-11 cm
long, 0.5-2 cm wide, gradually narrowed to the petiole; cauline leaves very
narrowly elliptic or linear to lanceolate, 1-9.5 cm long, 0. 1-2.3 cm wide,
sinuate-dentate to subentire, subsessile. Inflorescence dense in bud, un¬
branched to sparsely branched, 2-80 cm long; bracts lanceolate to ovate,
2. 5-6.5 mm long, 1-2.5 mm wide. Floral tube 6.5-20 mm long. Sepals
11-21 mm long, 2-3 mm wide. Petals 10-15 mm long, 4-8 mm wide. Fil¬
aments 6-13 mm long; anthers 2-6 mm long. Style 16-36 mm long. Fruit
with or without prominent lower comers or projections of the wings at or
above the middle, 4.5-8 mm long, 2-5 mm thick, sessile or on a stripe up to
2.2 mm long. Seeds ( l-)2-4( -5), 2-3 ( -4 ) mm long, 1-1.5 mm thick,
yellowish to light brown. Self-incompatible. Gametic chromosome num¬
ber, n = 7.

Type: Cedar woods, in sandy and rocky soil, New Braunfels, Comal
Co., Texas, May 1847, F. Lindheimer IV. 611 (GH. Isotypes, BM, CAMB,
F, LE, M, MO, NY, PH, US).

Distribution (fig. 57): Open sandy places and washes, Tulsa and
Woodward counties, Oklahoma, south to Nacogdoches, Goliad, and Cam¬
eron counties, Texas, to northern Coahuila, Mexico, northward to Bernalillo

MEMOIRS OF THE TORREY BOTANICAL CLUB

81

and Torrance counties, New Mexico, and the central Panhandle in Texas.
Flowers April to June.

In characters of the pubescence of the inflorescence, bract size, and fruit
morphology, the two subspecies of Gaura suffulta intergrade unevenly, with
various recombinants being characteristic of different populations in the
zone of contact. One such recombinant is G. suffulta variety terrellensis
Munz, which has the glandular pubescence of subspecies nealleyi but the
bracts and non-stipitate fruits of subspecies suffulta. The length of the
floral tube appears to increase along a cline from east to west, being 12-
16.5 mm in populations of New Mexico, 4.5-7.5 mm in those of eastern
Texas.

18a. Gaura suffulta Engelm. ex Gray subsp. nealleyi (Coult.) Raven &
Gregory, comb. nov. Figs. 23, 24, 59.

Gaura nealleyi Coult., Contr. U.S. Natl. Herb. 1: 38. 1890. Munz, Bull.

Torrey Bot. Club 65: 121. 1938; N. Amer. Fl. II. 5: 193. 1965.

Gaura suffulta Engelm. ex Gray var. terrellensis Munz, Bull. Torrey Bot.
Club 65: 121. 1928. Type: Sanderson, Terrell Co., Texas, 1 May

82

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

FIGURES 58-63. Flowers of the speeies of Gaura seet. Ptcrogaura. Fig. 58. G.
suffulta subsp. suffult a.— Fig. 59. G. suffulta subsp. nealleyi.— Fig. 60. G. hexandra
subsp. gracilis.— Fig. 61. G. hexandra subsp. hexandra.— Fig. 62. G. hrachycarpa.—
Fig. 63. G. triangulata.

MEMOIRS OF THE TORREY BOTANICAL CLUB

83

1934, V. L. Cory 8469 (POM 200873. Isotype, TAES). Munz, N.

Amer. FI. II. 5: 190. 1965.

Stems usually branched from the base, 2-7 (-10) dm tall. Inflorescence
glandular-pubescent, especially on the outside of the floral tube and on
the sepals, the ovary subglabrous. Rosette leaves 3.5-9 cm long, 0.5-1. 5
cm wide; cauline leaves very narrowly elliptic or narrowly lanceolate to
linear, 1.5-7 cm long, 0.1-0.6 cm wide, sinuate dentate, undulate. Inflores¬
cence 10-50 cm long; bracts lanceolate to ovate, 2.5-3.5 mm long, 1-2.5
mm wide. Floral tube 10-20 mm long. Petals 5-8 mm wide. Filaments
8—13 mm long. Style 22-36 mm long. Fruit with more prominent lower
comers than in subspecies suffulta, the stipe 0.2-2. 2 mm long. Seeds .3-4
(~5), 2-3 (-4) mm long. Self-incompatible. Gametic chromosome number,
n = 7.

Type: Chenate Mts., Presidio Co., Texas, 1889, G. C. Nealley 545 (US
15158. Isotype, F).

Distribution (fig. 57): Trans-Pecos Texas and northern Coahuila,
Mexico, north to Bernalillo and Torrance counties, New Mexico, in washes
and other sandy places. Flowers April to June.

Representative Specimens Examined:

Mexico, coahuila. Canon de Indio Felipe, Stewart 18 (GH).

United States, new Mexico. Bernalillo Co.: Above Tijeras, Barneby 2414
(US). Chaves Co.: 6 mi se of Elk, Munz b Gregory 23347 (DAO, RSA). Eddy
Co.: Artesia, Gregory 306 (NY, RSA). Lincoln Co.: 15 mi e of Capitan, Hitchcock
et al. 4232 ( DS, MY, POM, RSA, US). Socorro Co.: Weather Station 5, Chupadera
Mesa, 6500 ft, Dunn 4856 (RSA). Torrance Co.: 8 mi se of Cedarvale, Stephens
b Brooks 26008 (DS). texas. Brewster Co.: 5 mi w of Alpine, Cory 53523 ( DS,
FSU, GH, NY, SMU); 20 mi s of Alpine, Sperry T-981 (TAES, US). Culberson
Co.: Guadalupe Mts., Havard in 1881 (US); 14.5 mi s of Van Horn, Gregory 80
(RSA); 4 mi w of Kent, Parks b Cory 30875 (POM, TAES). El Paso Co.: Gueco
Tanks, Whitehouse 8584 (F). Hudspeth Co.: 43 mi w of Salt Flat, Raven b Gregory
19151 (DS). Jeff Davis Co.: Limpia Canyon, Tracy 294 ( BM, G, GH, MIN, MSC,
PENN, TAES, WISC). Pecos Co.: 24 mi nw of Sheffield, McVaugh 8210 ( DS, F,
G, LL, MICH, SMU, TEX). Presidio Co.: Marfa, Hinkley 2952 (BH, TAES).
Terrell Co.: Sanderson, Cory 53602 ( GH, NY, RSA, SMU, TAES); 22 mi se of
Dryden, Whitehouse 18604 (SMU). Val Verde Co.: Near mouth of Devils River
Correll 27130 (LL).

Gaura suffulta subspecies nealleyi has, in general, been a rather poorlv
understood entity. Because of the stipe present in some individuals it has
been loosely associated with the members of section Stipogaura and not
seen in proper perspective as an intergrading geographical race within the
species G. suffulta. The many collections made in recent years have helped
greatly to clarify its affinities.

84

A REVISION OF THE GENUS GAURA (ONAGRACEAE)

18b. Gaura suffulta suffulta. subsp. Figs. 22, 58.

Caura suffulta Engelm. ex Gray var. typica Munz, Bull. Torrey Bot.
Club 65: 120. 1938.

Stems moderately well branched, but without any definite pattern,
2.5—12 dm tall. Inflorescence glabrous or rarely with a very few long
hairs on the bracts. Rosette leaves 7—11 cm long, 1.5—2 cm wide; cauline
leaves narrowly lanceolate to lanceolate, 1-9.5 cm long, 0. 1-2.3 cm wide,
sinuate-dentate to subentire, plane. Inflorescence 6—80 cm long; bracts lan¬
ceolate to narrowly ovate, 2-6.5 mm long, 1.5-2 mm wide. Floral tube 6.5-
14 mm long. Petals 4—6 mm wide. Filaments 6-9 mm long. Style 16—32 mm
long. Fruit without prominent lower corners but usually with projections of
the wings at or above the middle, sessile, or in western populations, on a
stipe up to 1 mm long. Seeds (l-)2-4, 2-2.5 mm long. Self-incompatible.
Gametic chromosome number, n = 7.

Distribution (fig. 57): Open, sandy places, common in Texas but rare
in southern, eastern, and northern portions of the state and absent in the
Trans-Pecos; southern Oklahoma, east to Tulsa, Okfuskee, and Coal coun¬
ties, and in Woodward County.

Representative Specimens Examined:

United States. Oklahoma. Caddo Co.: 10 mi w of Apache, Waterfall 11932
(OKLA). Carter Co.: 6 mi s of Turner Falls, Hopkins 4859 (OKLA). Coal Co.:
Camp Simpson near Bromide, Goodman 5464 (OKL). Comanche Co.: 11 mi n of
Medicine Park, Stephens 20336 ( DS, KANU). Jefferson Co.: W of Wanrika, Rohr-
haugh 498 (OKL). Greer Co.: Below Lake Altus, Goodman 4400 (OKL, OKLA).
Kiowa Co.: S of Lake Altus, Granite Mts., Waterfall 9456 (KANU, OKLA, RSA,
SMU). Marshall Co.: 4 mi n of Medill, Goodman 6539 (OKL, OKLA, RSA, SMU).
Murray Co.: Davis, Emig in 1917 (CM). Okfuskee Co.: 1 mi s of Paden, Ray 2

(OKLA). Oklahoma Co.: Oklahoma City, Waterfall 1938 (OKLA). Pontotoc Co.:

1/4 mi e of Ada, McCoy 582 (OKLA). Tillman Co.: Skinners 26064 (SMU).
Tulsa Co.: Clark 339 (OKL). Woodward Co.: 5 mi ne of Woodward, Crockett 32
(OKL). texas. Anderson Co.: Palestine, Thar)) 864 (US). Andrews Co.: Scudday

in 1960 (LL). Armstrong Co.: Palo Duro State Park, Parks 6- Cory 13521 (TAES).

Baylor Co.: 13 1/3 mi s of Seymour, Cory 37247 (TAES). Bell Co.: C. L. ir G.
York 55233 (KANU, OKL). Bexar Co.: 6.8 mi n of San Antonio, Klein 1719 (DS).
Bosque Co.: 5 mi ne of Clifton, Shinners 14214 (SMU). Brown Co.: 18 mi w of
Brownwood, Atkerson 8 (TEX). Burnet Co.: Marble Falls, Carsiets 6- Studhalter
4345 (TEX). Callahan Co.: 15 mi se of Abilene, Henderson 64-555 (DS). Cameron
Co.: 8.4 mi w of Brownsville, Blankinship 6 (SMU). Coke Co.: Smith et al. 21640
(TAES). Comal Co.: Comanche Spring, Lindheimer 42 (GH). Coryell Co.: 13 mi
e of Gatesville, Jackson 5 ( LL, SMU, TEX). Cottle Co.: Meek 60 ( TTC ) . Crockett
Co.: Ozona, Jones 25878 (POM). Crosby Co.: 5 mi e of Crosbytown, Pittman 104
(OKLA). Dallas Co.: Prairies, Reverchon 4334 A (MIN, NY). Dawson Co.: 15 mi
w of Gail, C. L. ix A. A. Lundell 16960 (LL). Denton Co.: 5 mi w of Denton,
Whitehouse 15564 (SMU). Dickens Co.: 11.4 mi w of Guthrie, Roberts 32 (OKLA).

MEMOIRS OF THE TORREY BOTANICAL CLUB

85

Eastland Co.: Rising Star, Crossley in 1931 (TEX, mixed with G. triangulata) .
Ector Co.: 3 mi e of Odessa, McCracken 53 (TTC). Edwards Co.: Yard Ranch
Exper. Station, Parks ir Cory 15264 (TAES). Ellis Co.: 4 mi n of Midlothian, Perkins
16 (SMU, TAES). Erath Co.: 3.5 mi s of Hico Hwy., lloisington 57 (OKL). Fayette
Co.: La Grange to Schulenburg, H. L. S. i? K. O. L. in 1935 (MO). Galveston Co.:
0.5 mi w of Arcadia Park, Parks & Cory 32087 (TAES). Garza Co.: 12 mi n of
Post, McNabb 125 (OKLA). Gillespie Co.: 10 mi w of Frederiekberg, Barkley
1452B ( MSC, TEX). Goliad Co.: Goliad to Bexar, Berlarulier 2578 (PH). Grayson
Co.: Near Sherman, Maples in 1963 (VDB). Hamilton Co.: N of Edson, Wolff 2090
(TAES). Harp Co.: Speck Hall, Cheaser in 1964 (SMU). Hays Co.: San Marcos and
vicinity, Stanfield (NY). Hill Co.: Itasca, Smith 27 (TEX). Hood Co.: 7.5 mi ne of
Granbury, Skinners 11058 (SMU). Jack Co.: 4.1 mi ne of Jacksboro, Shinners 12349
(SMU). Kendall Co.: 2.1 mi n of Boeme, Raven ir Gregory 19344 (DS). Kent Co.:
t mi sw of Clairemont, Cory 13680 (POM). King Co.: 4 mi n of Guthrie, Sullivan
43 (OKLA). Lampasas Co.: 5 mi nw of Lampasas, Shinners 28407 (SMU). Lubbock
Co.: 1 mi w of Lubbock, Stewart 125 (OKLA). Martin Co.: W of Tarzan, Gregory
300 ( RSA, US). McCulloch Co.: 12.3 mi w of Brady, Shinners 30130 (SMU).
McLennan Co.: N of Mart, Smith 955 (OKLA). Menard Co.: 1 mi e of Menard,
Anders 132 (TAES). Midland Co.: 4 mi e of Midland, Raven 6- Gregory 19223
(DS). Mills Co.: 6.5 mi s of Goldthwaite, Parks <b Cory 13030 (POM, TAES).
Mitchell Co.: Lone Wolf Creek, Colorado, Pohl 4924 (ISC, SMU). Montague Co.:
Brooks 13223 (POM). Nacogdoches Co.: Powell 27 (DS). Navarro Co.: Frost,
McCard 2 (TEX). Parker Co.: 9.5 mi e of Weatherford, Shinners 10161 (SMU).
Randall Co.: Palo Duro State Park, Cory 13522 (POM). San Saba Co.: 2.2 mi s of
San Saba, Shinners 28339 (SMU). Shackelford Co.: 3 mi w of Albany, Mahler 1628
(OKLA). Somerwell Co.: 1 mi s of Glen Rose, Shinners 11279 (SMU). Sutton
Co.: 4 mi s of Sonora, Rohrbaugh 382 (COLO, OKL). Tarrant Co.: 2 mi sw of
Benbrooke, Cory 54407 (GA, KANU, LL, NSC, SMU). Taylor Co.: 1 mi e of
Needle Dome, Mahler 3192 (SMU). Travis Co.: Austin, Raven <Lr Gregory 19369
(DS). Upton Co.: Rankin, Cory 5136 (POM). Uvalde Co.: 14 mi n of Uvalde,
T. & L. Mosquin 5618 (DAO). Valverde Co.: N of Del Rio, Pilsbry (PH). Wilbarger
Co.: 11.8 mi w of Electra, Whitehouse 9808 (SMU). Williamson Co.: 2 mi e of
Georgetown, Gregory 288 (NY, RSA, US). Winkler Co.: 15 mi ne of Kermit
Collins in 1965 (TTC).

The distribution and characteristics of this entity were first clearly de¬
lineated by Mathias (Ann. Missouri Bot. Gard. 16: 399-401. 1929).

19. Gaura hexandra Gomez Ortega, Dec. 14. 1797. Munz, N. Amer. FI.
II. 5: 191. 1965, pro parte.

Annual herb from a stout taproot; stems usually well branched at the
base and above, 1.5-10 dm tall. Plants villous with hairs 1-2 mm long
below the inflorescence, the leaves subglabrous to more or less densely
short-villous; inflorescence subglabrous, strigulose, or glandular-pubescent,
or with a mixture of strigulose and glandular pubescence. Rosette leaves
lyrate, 3-15 cm long, 0.5-1 cm wide, gradually narrowed to the petiole;
cauline leaves linear to very narrowly elliptic or narrowly lanceolate, 1-9

A revision of the genus gaura (onagraceae)

cm long, 0. 1-0.8 cm wide, sinuate-dentate to subentire, subsessile. In¬
florescence strict to somewhat branched, 7-53 cm long; bracts narrowly
lanceolate to ovate, 2—5 cm long, 1—2 mm wide. Flowers 3— merous or 4—
merous. Floral tube 4.5—10 mm long. Sepals 3—12 mm long, 0.7.5— 2 mm
wide. Petals 4.5-9.5 mm long, 2-4 mm wide. Filaments 3-6 mm long;
anthers 1.5-3 mm long. Style 9-18.5 mm long. Fruit ellopsoid or narrowly
obovoid, narrowed at the base but not stipitate, 4.5—8 mm long, 2 — 1.5 mm
thick. Seeds 3 or 4, 1.75-3 mm long, 1-1.25 mm thick, yellowish to reddish
brown. Self-compatible and mostly autogamous. Gametic chromosome num¬
ber, n = 7.

Type: Cultivated in the botanical garden at Madrid, and flowering

FIGURE 64. Range of Gaura hcxarulra subsp. gracilis (filled triangles) and
G. hcxarulra subsp. hcxarulra (open triangles).

MEMOIRS OF THE TORREY BOTANICAL CLUB

87

in August 1797, from seeds sent from Mexico by Martin de Sesse y Lacasta;
incorrectly stated to be from Cuba.

Distribution (fig. 64): Washes and other sandy places at middle
elevations in the mountains, from central Arizona and southwestern New
Mexico south in the Sierra Madre Occidental to the Trans-Mexican Vol¬
canic Belt, and in Chiapas and southern Guatemala. Flowers July to
October.

Mixed populations involving 3- and 4-merous plants occur in the region
where the two subspecies come together in northern Mexico; very rarely
3- and 4-merous flowers are found on the same plant. Both sorts of plants
are found at various localities in western Chihuahua, where subspecies
gracilis is more common, and in Durango, where subspecies hexandra is
more common. They are maintained by their rather strict autogamy.

19a. Gaura hexandra subsp. gracilis (Wooton & Standi.) Raven &

Gregory, comb. nov. Figs. 25, 6.

Gaura gracilis Wooton & Standi., Contr. U.S. Natl. Herb. 16: 153. 1913.
Munz, Bull. Torrey Bot. Club 65: 121. 1938; N. Amer. FI. II. 5: 190.
1965.

Gaura podocarpa Wooton & Standi., Contr. U.S. Natl. Herb. 16: 154.
1913. Type: Bear Mt., 1500 m, near Silver City, Grant Co., New
Mexico, 17 June 1903, O. B. Metcalfe 166 (US 495277. Isotypes, DS,
G, GH, LE, MIN, MO, NY, POM).

Gaura strigillosa Wooton & Standi., Contr. U.S. Natl. Herb. 16: 154.
1913. Type: Wingfields Ranch on Ruidosa Creek, White Mts., Lincoln
Co., New Mexico, 8 July 1895, E. O. Wooton (US 561073).

Gaura brassicacea Wooton & Standi., Contr. U.S. Natl. Herb. 16: 152.
1913. Type: Socorro, Socorro Co., New Mexico, May 1881, G. R. Vasey
(US 45764).

Gaura glandulosa Wooton & Standi., Contr. U.S. Natl. Herb. 16: 153.
1913. Type: Reserve, Catron Co., New Mexico, 9 July 1906, E. O.
Wooton (US 561072).

Gaura gracilis Wooton & Standi, f. glandulosa (Wooton & Standi.) Munz,
Bull. Torrey Bot. Club 65: 122. 1938. Munz, N. Amer. FI. II. 5: 190.
1965.

Rosette leaves persistent. Flowers 4-merous. Floral tube 6-10 mm
long. Sepals 6-12 mm long, 1-2 mm wide. Petals 5.5-9.5 mm long, 2.25
mm wide. Filaments 4-6 mm long; anthers 2-3 mm long. Style 11.5-18.5
mm long. Fruit 6-8 mm long, 2-3 mm thick. Seeds 4. Self-compatible and
often autogamous. Gametic chromosome number, n = 7.

88

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Type: Forest Nursery, Fort Bayard, Grant Co., New Mexico, 29 August
1905, J. C. Blumer 44 (US 499693. Isotypes, GH, NY).

Distribution ( fig. 64 ) : Sandy washes at middle elevations, from eastern
Mohave County south through the mountains of central Arizona to eastern
Pima County and the southwestern quarter of New Mexico, and in Mexico
southward in the Sierra Madre Occidental to eastern Sonora and throughout
the western halves of Chihuahua and Durango, but much rarer than sub¬
species hexandra south of central Chihuahua.

Representative Specimens Examined:

Mexico, chihuahua. 3 mi e of La Junta, Freytag b Baxter 29 (MO); moun¬
tains near Pilares, Hartman 796 (F, GH, NY, PENN, US); Loreto, Rio Mayo, Gentry
2557 (GH, MO); Memelichi, Rio Mayo, Gentry 2741 (F, GH, MO, MENU, US);
San Juanito, Knobloch 5415 ( F, MSC); Sierra Madre foothills, Nelson 6110 (GH,
K, US); vicinity of Chihuahua, Palmer 204 ( F, GH, MO, NY, US); Strawberry
Creek, ne of Colonia Pacheco, Pennell 19195 (PH, US); El Cima, Le Seuer 806
(GH, IND, MENU, MO, OKL, PH, SMU, TEX) ; mesas near Carretas, Pringle 2002
( BM, BR, CM, COCO, G, LE, MEXU, MO, MSC, NY, US); Broad Canyon, 10 mi
w of Cuautemoc, Straw b Forman 1680 (GH, MEXU, RSA); 1 mi w of Aguaje,
Straw b Forman 1849 (RSA); 1 mi s of Guadalupe, Straw b Forman 1912 (MICH,
RSA); 11 mi s of Matamoros, Ripley b Barneby 13942 (DS); Colonia Garcia, Sierra
Madre Occidental, Townsend b Barber 124 (G, K, MEXU, ND, NY, P, POM, US);
7 mi s of Parral, Waterfall 16130 (OKL, OKLA, RSA, SMU); 40 mi nw of Chihuahua,
White 2399 (GH, MEXU, MICH), durango. Near La Providencia, Nelson 4982
( GH, US); Otinapa, Palmer 552 (US); Inde, Reko 5171 (US); 28 mi sw of Durango,
Waterfall b Wallis 13410 (OKLA). sonora. San Bernardo, along Rio Curahui,
Pennell 19595 (PH, US); Puerto de los Aserrados, Phillips 405 (GH, MICH);
Las Tierritas del Temblor, White 3442 (GH, MICH).

United States. Arizona. Apache Co.: McNary, Deaver 1664 (RSA); 3 mi e
of Alpine, Krai 21397 (VDB). Cochise Co.: Cave Creek Canyon, Barkley 14A683
(GH, MIN, NY, TEX); Mule Mts., Bisbee, Henderson in 1933 (TEX); Huachuca
Mts., Goodding 250 (GII, NY). Coconino Co.: Flagstaff, Hanson A113 (CU, F, ILL,
MO, MSC, NY, PENN, TEX); U.S. Army Navajo Ordnance Depot, Schallert in
1943 (F, ILL, NCU, NY, PENN). Gila Co.: Matzatzal Mts, Collom 15 (GII, MICH,
MO, NY, US); Workman Creek Canyon, Gould 3648 (NY, RSA). Graham Co.:
San Carlos Indian Reservation, A. b R. A. Nelson 1835 (GH, MICH, NY, PH, SMU,
US). Greenlee Co.: 37 mi n of Metcalf Range, Hutchinson 6866 (NO, RSA). Maricopa
Co.: W of Phoenix, Harrison 1823 (US). Mohave Co.: S top of Blue Mt, Coronado
Trail, Ramsey 1105 (POM). Navajo Co.: 2 mi w of McNary, Ferris 10102 ( DS,
MICH). Pima Co.: Manning Camp, Blunter 3378 ( DS, F, CH, ISC, MIN); Santa
Rita Mts, Peebles b Harrison 2950 (US). Pinal Co.: 7 mi w of Superior, Gregory
578 (DS). Santa Cruz Co.: White House Canyon, Shreve 7681 ( F, MICH, MO).
Yavapai Co.: 0.5 mi w of Prescott, Wolf 2340 (BH, DS, CH, POM, RSA). new
Mexico. Catron Co.: Beaverhead, Eggleston 20374 (US); Mangas Mts, 8 mi s of
Horse Spring, 10 mi n of Aragon, Weber b Salamun 12778 (COLO). Chaves Co.:
5 mi ne of Roswell, Palmer 6 (F). Dona Ana Co.: Organ Mts, Wooton b Standley
in 1906 (US). Granite Co.: 15 mi n of Silver City, ca. 7000 ft, Hitchcock et al.
4394 (COLO, DS). Grant Co.: Mimbres River, Metcalfe 1033 ( F, CH, MIN, MO,

MEMOIRS OF THE TORREY BOTANICAL CLUB

89

NY, POM). Lincoln Co.: White Mts., Wooton 339 (ILL, MIN, MO, NY, US).
Sierra Co.: Lake Valley, Beals in 1904 (MICH). Socorro Co.: Pinos Altos Mts.,
Greene (GH, MO, NY, PH, POM); Mogollon Mts., Rushy 137 ( F, MICH, MO,
NY, PENN).

This taxon is modally self-pollinating but closely similar to the self¬
incompatible Gaura suffulta subspecies nealleyi, from which it was cer¬
tainly derived. In turn, G. hexandra subspecies gracilis seems to have given
rise to subspecies hexandra, which ranges much farther to the south and is
almost exclusively self-pollinating.

19b. Gaura hexandra subsp. hexandra. Figs. 26, 61.

Gaura tripetala Cav., Ic. 4: 66, pi. 396. 1798. Type: Flowering in the
Royal Botanical Garden, Madrid, from May to August 1797, the
seeds gathered along the roadside near Pachuca and San Agustin de
las Cuevas, Hidalgo, Mexico, and sent to Madrid by Louis Nee
(wild gathering, F; cultivated material from Madrid, F. Fragment,
GH). Munz, Bull. Torrey Bot. Club 65: 211. 1938.

Gaura tripetala Cav. var. coryi sensu Munz, Bull. Torrey Bot. Club 65:
212. 1938, pro parte.

Gaura hexandra Gomez Ortega var. coryi sensu Munz, N. Amer. FI. II.
5: 191. 1956, pro parte.

Rosette leaves not as persistent as in subspecies gracilis. Flowers 3-
merous. Floral tube 4. 5-7.5 mm long. Sepals 3-10 mm long, 0.75-1.5 mm
wide. Petals 4.5-7 mm long, 2.3 mm wide. Filaments 3-5.5 mm long;
anthers 1.5-2 mm long. Style 9-14.5 mm long. Fruit 4.5-8 mm long,
2.5-4. 5 mm thick. Seeds 3. Self-compatible and highly autogamous.
Gametic chromosome number, n = 7.

Distribution (fig. 64): Sandy and weedy places, and in cornfields; in
Mexico from Durango south in the Sierra Madre Occidental to the Trans-
Mexican Volcanic Belt, where abundant, and in Chiapas and Guatemala.

Representative Specimens Examined:

Guatemala. Near Teepan, Goodman 3500 ( F, ISC, OKL).

Mexico, aguascalientes. Near Aguasealientes, Rose 6- Painter 7786 (NY, US).
chiapas. Cerro San Cristobal, Breedlove 10245 (DS). chihuahua. 3 mi e of La
Junta, Freytag i? Baxter 29 (US); 10 mi e of Madera, Breedlove 15541 (DS);
Basigochi, se of Creel, Knobloch 480 (MSC). durango. 5 mi e of El Salto, Breed¬
love 15501 (DS); 3 mi ne of Otinapa, Maysilles 7308 (MICH); Durango, Palmer
257 (C, F, G, GH, MO, NY, US); 57 mi sw of Durango, Waterfall 6- Wallis 13713
(F, IA, IKL, OKLA, SMU); 18 mi w of Santiago Papasquiaro, Wiens 3515 (COLO,
DS); San Luis del Rio, 51 road mi nw of Coyotes, Maysilles 7226 (MICH), hidalgo.
2 mi w of Tulancingo, Moore 2047 (BH, GH). jalisco. 3 mi s of Mazamitla,

90

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

McVaugh 13074 (MEXU, MICH, SMU, US); near Km 27, sw of Ojuelos, 6 mi
ne of Paso de la Froje, McVaugh 16902 (MICH, RSA, TEX, US). Mexico. Com-
munidad Temascaltepec, Hinton 7688 (C, F, G, GH, LL, MICH, MO, NY, US);
near Pyramid of Calixtlahuaca, Barkley et al. 664 (F, TEX); Rancho de Rosario,
10 mi n of Mexico, D.F., Happ 153 (MO), michoacan. Just n of San Juan Viejo,
Beaman 4397 ( DS, MSC); near Volcan Paricutin, Eggler 36 (US); Llano, Hinton
11920 (DS, F, GH, MICH, NY, POM, US); Zitacuaro, Hinton 11863 ( BM, BR,
DS, F, G, GH, LE, MICH, NY, POM, TEX, US); 2 mi w of Tancitaro, Leaven¬
worth 294 ( F, GH, ILL, MICH, MO, NY), morelos. Cuernevaea, Kenotjer A419
(F). puebla. Guadalupe, Arsene 1113 (BR, US), tlaxcala. 5 mi nw of Huamantla,
Breedlove 15877 (DS). zacatecas. Villa Nueva, Seler 4790 (GH); 5 mi w of
Sanches Roman, Straw 6- Forman 1809 (RSA).

Gaura hexandra was described from plants that flowered in the Royal
Botanical Garden in Madrid in 1797, and it has been in cultivation in
botanical gardens ever since. Probably both it and Gaura tripetala , the
better-known name for this species, were described from the same culti¬
vated strain. We have seen numerous specimens collected from European
gardens throughout the 19th Century.

20. Gaura brachycarpa Small, FI. S.E. U.S. 848, 1335. 1903. Munz, Bull.
Torrey Bot. Club 65: 211. 1938; N. Amer. FI. II. 5: 190. 1965. Figs.
27, 62.

Gaura tripetala Cav. var. cortji Munz, Bull. Torrey Bot. Club 65: 212.
1938. Type: Dry banks, Hempstead, Waller Co., Texas, 25 April 1872,
E. Hall 211 (POM 39283. Isotvpes, GH, NY, US). The Mexican
specimens cited are G. hexandra Gomez Ortega subsp. hexandra.

Gaura hexandra Gomez Ortega var. coriji (Munz) Munz, N. Amer. FI.
II. 5: 191. 1965. The Mexican portion of the range given rests upon
specimens of G. hexandra Gomez Ortega subsp. hexandra.

Annual herb from a taproot; stems usually with several to many branches
from the base, very rarely with a single stem, 1.5-6.5(-8.5) dm tall. Plants
villous with hairs 1.5-2 mm long below the inflorescence, the leaves sub-
glabrous or sparsely villous along the veins and around the margins; in¬
florescence subglabrous or the ovary, outside of the floral tube, and sepals
strigulose. Rosette leaves lyrate, 6-9.5 cm long, 1—1.5 cm wide, gradually
narrowed to a petiole; cauline leaves narrowly lanceolate to oblanceolate,
1-7 cm long, 0. 1-1.6 cm wide, shallowly sinuate-denticulate to subentire.
Inflorescence usually unbranched, 10-51 cm long; bracts lanceolate, 2-4
mm long, 0.5-1 mm wide. Flowers 4-merous, very rarely 3-merous. Floral
tube 6. .5-12 mm long. Sepals 10-15 mm long, 1.5-2 mm wide. Petals 8-12.5
mm long, 2.5-5 mm wide. Filaments 5-8 mm long; anthers 2-4 mm long.

MEMOIRS OF THE TORREY BOTANICAL CLUB

91

Style 15-24 mm long. Fruit ellipsoid, with or without prominent lower
“comers,” 5.5-10 mm long, 2-3 mm thick, sessile or on a stripe up to 1
mm long. Seeds 3-4, 2-2.5 mm long, 1-1.5 mm thick, yellowish to light
brown. Self-compatible but mostly outcrossing. Gametic chromosome num¬
ber, n = 7.

Type: San Diego, Duval Co., Texas, 1885-1886, Alary B. Croft 109
(NY).

Distribution (fig. 65): Open sandy places, eastern Texas except for
the East Texas Timberlands, and in Bryan and Love counties, Oklahoma;
introduced near Tulsa, Oklahoma, and at one station each in Arkansas and
Mississippi. Flowering March to June, or occasionally longer at the southern
end of its range.

92

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Representative Specimens Examined:

United States. Louisiana. Acadia Parish: Trackway, Crowley, Penfound in
1936 (NO). Mississippi. Amite Co.: W of Pike Co. line on Hwy. 98, Jones 5345
(DS). Oklahoma. Bryan Co.: Vicinity of Durant, Blain 109 (US); 2 mi e, 1.25 mi s of
Roosevelt Bridge, J. ir C. Taylor 1578 (OKL). Love Co.: Near Marietta, Stevens 95
(NY). Tulsa Co.: 1.5 mi n, 2.5 mi e of State Hwy. 33, Clark 383 (OKLA). texas.
Anderson Co.: N of Palestine, Eggert in 1899 (MO). Aransas Co.: Rockport, Tharp
5599 (MICH, TEX, US). Atascosa Co.: 2.5 mi s of Campbellton, Correll 29233
(LL). Austin Co.: Industry, Wurzlow in 1891 (ILL). Bastrop Co.: Smithville,
Mans in 1930 (TEX). Bee Co.: Munz ir Gregory 23448 (RSA). Bexar Co.: San
Antonio, Behb 2264 (OKL). Brazos Co.: College Station, Crockett 8488 (LL).
Brooks Co.: 7 mi e of Falfurrias, Gould ir llycka 8114 (SMU, TEX, TTC). Burnet
Co.: 10 mi w of Burnet, Wolff 6- Dana 1550 (TAES, US). Caldwell Co.: 6 mi s
of Lockhart, McCart 6452 (SMU). Colorado Co.: 1 mi s of Eagle Lake, Harris in
1958 ( FSU, GH, LAF). Dallas Co.: Rylie, Krai 761 (FSU). Delta Co.: 5 2/3 mi
sw of Cooper, Cory 55830 (SMU, US). Denton Co.: 2 mi sw of Denton, McCart 1983
(ISC, SMU, TEX). DeWitt Co.: 3 mi w of Cuero, McCart 5992 (SMU). Dimmit Co.:
Catarina, Cory 28062 (TAES). Duval Co.: S of Sejita, Johnston 5487 (SMU, TEX).
Fayette Co.: Near La Grange, Albers 224 (NY). Fort Bend Co.: 21 mi w of Richmond,
L. N. i? G. B. Upton in 1930 (CU). Freestone Co.: 15 mi se of Fairfield, Krai
872 (FSU). Frio Co.: 4 mi w of Dilley, Alvarez et al. 8016 (FSU, OKLA, TEX).
Gillespie Co.: 18 mi ne of Fredricksburg, Raven 6- Gregory 19339 (DS). Goliad Co.:
N of Goliad, Eggert in 1900 (MIN, MO). Gonzales Co.: McCaleb 38 (OKLA).
Grayson Co.: Denison, Leas in 1932 (TEX). Harris Co.: Houston, S chneck in 1894
(ILL). Hays Co.: 5 mi w of San Marcos, Gregory 420 (RSA). Hidalgo Co.: McAllen,
Fleetwood 8067 (TEX). Hill Co.: 7 mi sw of Hillsboro, Shinners 14170 (SMU).
Hood Co.: Granbury, Waynles School in 1929 (TEX). Hunt Co.: Bank of Greenville
River, Bebb 2649 (OKL). Jim Wells Co.: 7.7 mi n of Fremont, Shinners 17922
(SMU). Johnson Co.: Cleburne, Whitehouse in 1930 (TEX). Karnes Co.: 7 3/4
mi n of Karnes City, Shinners 9834 (SMU). Kaufman Co.: Gentry 53-221 (TEX).
Kleberg Co.: 2.7 mi s of Ricardo, Shinners 17843 (SMU). La Salle Co.: 6.5 mi w of
Fowlerton, Parks ir Cory 28534 (TAES). Leon Co.: 5.5 mi e of Marquez, Shinners
30983 ( DS, DUKE, SMU). Limestone Co.: Near Delia, Matthews 145 (TTC).
Llano Co.: Llano, Bray 320 (TEX). Mason Co.: 4 mi w-nw of Mason, Shinners
26295 (SMU). McLennan Co.: East Waco, Smith 406 (ILL, OKLA). Medina
Co.: 3 mi ne of Natalia, Raven <Lr Gregory 19348 (DS). Milam Co.: 3.2 mi e of
Mayfield, Shinners 29450 (SMU, TEX). Mills Co.: 8 mi sw of Goldthwaite, Shinners
28320 (SMU). Navarro Co.: Dawson, Reverchon 3843 (MO). Nueces Co.: Corpus
Christi, Heller 1391 ( F, GII, ILL, ISC, LE, MICH, MIN, MO, MSC, NY, PH, RSA,
SMU). Rains Co.: 2.5 mi nw of Point, Shinners 14341 (SMU). Refugio Co.: 6
mi n of Bayside, Whitehouse 21231 (SMU). San Patricio Co.: 4 mi n of Sinton,
Goodman 7469 (OKL, RSA). San Saba Co.: 2.2 mi s of San Saba, Shinners 28342
(SMU). Tarrant Co.: Fort Worth, Moldenke 6968 (NY). Travis Co.: Pedemales
River w of Hamilton Pool, Webster 74 (TEX). Uvalde Co.: W of Uvalde, Jones in
1931 (RSA). Van Zandt Co.: Wills Point, Henderson 5693 (TEX). Victoria Co.:
Victoria, Lcwton 92 (PH, US). Waller Co.: Hempstead, Hall in 1872 ( F, GH, ILL,
MO, NMC, NY, PH, POM, US). Webb Co.: Laredo, Palmer 11264 (GH, MO,
NY, US). Wharton Co.: 1 mi e of Hungerford, Turner 4515 (TEX). Wichita

MEMOIRS OF THE TORREY BOTANICAL CLUB

93

Co.: 12.5 mi nw of Iowa Park, Mertins 836 (OKLA). Willacy Co.: 4 mi s of
Raymond ville, Goodman 7475 (OKL, RSA). Wilson Co.: 1.4 mi n of Stockdale,
Skinners 19525 (SMU). Zavala Co.: 5 mi n of La Pryor, Serna et al. 125 (DUKE,
OKLA, SMU).

We have seen one specimen of Gaura brachycarpa labeled Sabinal,
Mexico, 26 March 1932, M. E. Jones (POM). Although the species is to be
expected in Mexico, its occurrence in northern Chihuahua would be in¬
consistent with the rest of its range. Jones collected at one place called
Sabinal in this area in earlier years (cf., Leaf!. W. Bot. 10: 222. 1965),
but perhaps his 1932 locality was different; thus far we have been unable
to trace it.

21. Gaura triangulata Buckl., Proc. Acad. Nat. Sci. Philadelphia 1861:

454. 1862. Figs. 28, 29, 63.

Gaura tripetala Cav. var. triangulata (Buckl.) Munz, Bull. Torrey Bot.

Club 65: 213. 1938.

Gaura hexandra Gomez Ortega var. triangulata ( Buckl. ) Munz, N. Amer.

FI. II. 5: 191. 1965.

Slender, erect annual herb from a taproot; stems usually with several
branches from the base, sometimes unbranched, 1.5-6 dm tall. Plants
villous with hairs 1. 5-2.5 mm long below the inflorescence, the leaves sub-
glabrous or sparsely villous along the veins and around the margins; in¬
florescence glabrous or the ovary, outside of the floral tube, and sepals
lightly strigulose. Leaves very narrowly elliptic to oblanceolate, 1.5-8 cm
long, 0.2-1. 5 cm wide, entire to very slightly sinuate-dentate, subsessile.
Inflorescence dense in bud but fairly well spaced just before the flowers
open, 10-37 cm long; bracts lanceolate or narrowly ovate, 2-4 mm long,
1-1.75 mm wide. Flowers 3-merous, very rarely 4-merous. Floral tube
4-5.5 mm long. Sepals 4.5-6 mm long, 0.75-1 mm wide. Petals 3.5-5 mm
long, 1-2 mm wide. Filaments 2-3.5 mm long; anthers 1.5-3 mm long.
Pollen about 50 percent fertile. Style 9-10 mm long. Fruit ovoid, 7-9 mm
long, 3-5 mm thick, sessile. Seeds (l-)2-5, 1.5-3.5 mm long, 1-1.5 mm
thick, yellowish to light brown. Autogamous. Gametic chromosome num¬
ber, n — 7 ( ring of 14 at meiotic metaphase I ) .

Type: “Prairies, northern Texas”; i.e., near Fort Bellknap, Young Co.,
Texas, May, S. B. Buckley (PH).

Distribution (fig. 57): Open sandy places, north-central Texas and
south-central Oklahoma, with outlying stations near Tulsa, Oklahoma, and
in Tom Green and Crosby counties, Texas. Flowers May and June.

94

A REVISION OF THE GENUS GAURA ( ONAGRACEAE )

Representative Specimens Examined:

l^ITCD States- Oklahoma. Cleveland Co.: 5 mi nw of Norman, Engelmann
in 1935 (ISC, MO, OKL). Comanche Co.: Near Cache, Stevens 1325 (DS, MIN,
MO, NY). Cotton Co.: 1.7 mi n of Randlett, Skinners 25966 (SMU). Grady Co.:
Chickasha, McElreath 94 (OKL). Oklahoma Co.: Oklahoma City, Waterfall 1938
(GH, NY, OKLA). Rogers Co.: Catoosa, Bu h 1131 (MIN, MO, ND, NY, OKL,
WVA ) . Stephens Co.: 0.5 mi e of Comanche, Waterfall 3649 (MO, OKL). Tulsa
Co.: 15 mi e of Tulsa, Engelmann in 1935 (OKL, POM), texas. Archer Co.: 3 mi
n of Olney, McCart 834 (SMU). Baylor Co.: 8 mi e-ne of Mabelle, Shinners 18752
(SMU). Callahan Co.: 8 mi e of Abilene, Henderson 62-181 (DS). Clay Co.: 3
mi se of Henrietta, Stephens 20600 (DS). Coleman Co.: 1 mi se of Santa Anna
Shinners 26395 (ILL, SMU). Crosby Co.: 3 mi e of Crosbyton, Graham in 1960
(TEX). Eastland Co.: Eastland, Harris 51 (TEX). Erath Co.: 5 mi ne of Stephen-
ville, Shinners 11080 (SMU). Jones Co.: 6.5 mi ne of Anson, Cory 37214 (TAES).
Montague Co.: 8.5 mi w of Nocona, Shinners 13291 (SMU). Taylor Co.: 4 mi s of
Buffalo Gap Mahler 3166 (SMU). Throckmorton Co.: 5.4 mi n-nw of Throckmorton,
Shinners 18795 (SMU). Tom Greene Co.: 3 mi e of San Angelo, Ragsdale (SMU).
Wichita Co.: Wichita Falls, Shinners 18744 (SMU). Wilbarger Co.: 11.8 mi w of
Electra, Whitehorse 9808 (MICH). Young Co.: 3 mi se of Olney, Parks 6 Cary
13141 (MICH, SMU).

This distinctive and rather local complex structural heterozygote con¬
sists of an extremely uniform, highly autogamous series of populations.
As mentioned earlier, the species was almost certainly derived following
hybridization of the self-incompatible Gaura suffulta subspecies suffulta
with the self-compatible G. brachycarpa.

EXCLUDED NAMES

Gaura chinensis Lour., FI. Cochin. 225. 1790. Definitely not Gaura, and
probably not Onagraceae.

Gaura P epilobia Moc. & Sesse ex Scr., in DC., Prod. 3: 45. 1828 = Oeno¬
thera rosea L’Her. ex Ait., Ilort. Kew. ed. 1, 2: 3. 1789.

Gaura fruticosa Loefl., Inter I lisp. 248. 1758 = Combretum fruticosum
( Loefl. ) Stuntz, U.S. Dept. Agrie. Bur. PI. Ind. Invent. Seeds and
PI. Import. No. 31: 86. 1914; see J. Linn. Soc. Bot. 55: 117. 1953.
Gaura? fruticulosa Benth., Bot. Voy. Sulph. 15. 1844 =r Gongylocarpus
fruticulosus (Benth.) T. S. Brandcgee, Proc. Calif. Acad. Sci. II.
2: 158. 1889.

Gaura heterandra Torr., Pac. R.R. Rep. 4: 87. 1857 = Heterogaura het¬
erandra (Torr.) Coville, Contr. U.S. Natl. Herb. 4: 106. 1893.

Gaura humilis (Juss.) D. Dietr. Linnaea 21. 580. 1848 = Gayopliytum
humile Juss., Ann. Sci. Nat. (Paris) I. 25: 18. 1832.

Gaura linifolia Nutt., in James, Exp. Long 2: 355. 1823 = Stenosiphon
linifolius (Nutt.) Heynh., Norn. 2: 704. 1840.

INDEX*

Calylophus, 1, 3
Camissonia, 8
Clarkia, 1, 2

Combretum fruticosum, 94
Gaura, 9-12

section Campogaura, 27-34, 3, 6
section Eugaura, 55
section Gaura, 55-79, 3, 7, 8
section Guaridium, 14-18
section Pterogaura, 79-94, 3, 7, 8
section Schizocarya, 23-27, 6
section Stipogaura, 34-50, 3, 6,
7, 27, 83

section Xenogaura, 50-55, 3, 7
section Xerogaura, 19-22, 3, 27
angustifolia, 59-62, 4, 7, 8, 11,
55, 56, 58, 65, 70, 80
angustifolia var. eatonii, 59
angustifolia var. simulans , 59
angustifolia var. strigosa, 59
angustifolia var. typica, 59
australis, 23

biennis, 74-78, 1, 5, 7, 8, 10, 11,
55, 56, 64, 70, 71
biennis var. pitcheri, 64
boquillensis, 21-22, 3, 4, 6, 11,
15, 20

brachycarpa, 90-93, 5, 11, 79, 80,
82, 94
bracteata, 28
brassicacea, 87

calcicola, 40-43, 4, 10, 11, 35, 36
chinensis, 94
cinerea, 37

coccinea, 27-34, 4, 6, 7, 10, 11,
15, 18, 50, 54, 55
coccinea var. arizonica, 28
coccinea var. e/nloboides, 28
coccinea var. glabra, 28
coccinea p? integerrima, 28
coccinea var. parviflora, 28
coccinea var. ttjpica, 28
coloradensis, 63
crispa, 51

demareei, 78-79, 5, 7, 11, 55, 56,
58

drummondii, 50-55, 4, 7, 10, 11,
15, 27
eatonii, 59
epilobia, 94
epilobioides, 28
filiformis, 7, 64, 65, 70
filiformis var. kearneiji, 64
filiformis var. munzii, 57, 59
filipes, 43-46, 4, 6, 11, 35, 36,
41, 70

filipes p major, 43
filipes var. typica, 43
fruticosa, 59, 94
fruticulosa, 94
gentryi, 16
glabra, 28
glandulosa, 87
gracilis, 87

gracilis f. glandulosa, 87
grandiflora, 16
heterandra, 94
hexandra, 85-90, 79, 80, 90
hexandra var. coryi, 89, 90
hexandra subsp. gracilis, 87-89,
5, 11, 82, 86

hexandra subsp. hexandra, 89-90,
5, 7, 8, 11, 82, 86
hexandra var. triangulata, 93
hirsuta, 23
hispida, 51
humilis, 94
induta, 28

lindheimeri, 55-59, 4, 7, 10, 11,
71, 72, 74
linearis, 28
linifolia, 94

longiflora, 64-74, 5, 7, 10, 11, 55,
56, 58, 59, 78, 79
macrocarpa, 19-21, 3, 4, 6, 11,
15, 16, 22
marginata, 28

*Synonyms in italics.

96

INDEX

mckelveyac, 46-47, 7, 11, 35, 36,
41, 43, 50
michauxii, 43
micrantha , 23
mollis, 16
multicaulis, 28

mutabilis, 16-18, 3, 4, 6, 9, 11,
14, 15

mutabilis f. glabra, 16
nealleiji, 81

neomexicana, 62-64, 7, 55
neomexicana subsp. coloradensis,
63-64, 5, 11, 58
neomexicana var. coloradensis, 63
neomexicana subsp. neomexicana,
62-63, 5, 11, 56, 58
neomexicana var. tijpica, 62
odorata , 4. 7, 15, 27, 51, 52, 54
oenotheriflora, 16
parviflora. 23-27, 4, 6, 10, 11, 15,
23

parviflora var. lachnocarpa, 24

parviflora var. tijpica, 24

parviflora var. tijpica f. glabra, 24

parvifolia, 28

podocarpa, 87

roemeriana, 51

simulans, 59

sinuata, 47-50, 4, 7, 11, 35, 36,
40, 41, 43
spicata, 28
strigillosa, 87
s uffrutescens, 28
sufFulta, 80-85, 79
suffulta subsp. nealleyi, 81-83, 5,
7, 11, 79

sufFulta subsp. sufFulta, 84-85, 5,
11, 79, 81, 82, 83, 94

suffulta, var. terrellensis, 5, 81
suffulta var. tijpica, 84
triangulata, 93-94, 5, 8, 11, 79,
80, 81

tripetala, 80, 89, 90
tripetala var. coryi, 89, 90
tripetala var. triangulata, 93
villosa, 35-40, 6, 7
villosa var. arenicola, 37, 39
villosa var. mckelveyi, 46
villosa subsp. parksii, 39-40, 4,
11, 20, 36

villosa var. parksii, 39
villosa, var. tijpica, 37
villosa subsp. villosa, 37-39, 4,
11, 20, 36
Gauridium, 9, 14
kunthii, 16
molle, 16

mutabile, 9, 14, 16
Gayophytum humile, 94
Gongylocarpus fruticulosus, 94
Hauya, 1
Heterogaura, 1

heterandra, 94
Ludwigia, 8
Oenothera, 1

section Oenothera, 74
anomala, 16
biennis, 1
rosea, 94

Schizocarya, 10, 23
crispa, 50
drummondii, 50
kunthii, 28
micrantha, 10, 23
Stenosiphon, 1, 4

linifolius, 5, 94

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