(Polemoniaceae) Section Gilmania

Brigham Young University BYU ScholarsArchive

Theses and Dissertations

1965-08-01

Revision of gilia (polemoniaceae) section gilmania

M. Eileen Matthews Brigham Young University - Provo

Follow this and additional works at: https://scholarsarchive.byu.edu/etd

BYU ScholarsArchive Citation Matthews, M. Eileen, "Revision of gilia (polemoniaceae) section gilmania" (1965). Theses and Dissertations. 8086. https://scholarsarchive.byu.edu/etd/8086

This Thesis is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. REVISIONOF GILIA (POLEMONIACEAE)

SECTIONGILMANIA

A Thesis

Presented to the

Department of Botany and Range Science

Brigham Young University

In Partial Fulfillment

of the Requirements for the Degree

Master of Science

M. Eileen Matthews

May, 1971 •

ACKNOWLEDGMENTS

Grateful acknowledgment is made for the valuable suggestions and

help given by the chairman of my advisory committee, Dr. Stanley L.

Welsh, the chairman of the Department of Botany and Range Science, Dr.

Dayna L. Stocks, and other members of my committee, Drs. Glen Moore and

Dorald M. Allred. The author is also indebted to Daniel K. Johnson for

photographic work, Dr. James L. Reveal and N. Duane Atwood for assis-

tance in collection, and to Dr. Janice C. Beatley who made it possible

for the author to pursue field work in the desert mountains of the

United States Atomic Energy Commission's Nevada Test Site. Special

is also given to the curntors of the hcrbaria who made spcciwcns

available for study.

Financial assistance was provided (in part) by a fellowship from

the National Science Foundation (no. 999-00-8118). The Department of

Botany and Range Science, Brigham Young University, supplied laboratory

space, equipment, supplies, and financial support.

iii TABLEOF CONTENTS

Page

ACKNOWLEDGMENTS iii

LIST OF ILLUSTRATIONS V

INTRODUCTION•...•. 1

Statement of Problem 1

Methods and Procedures 2

History of the Section 4

Infrageneric Relationships 10

General Morphology 32

Distribution and Ecology 43

Phylogeny 49

Cytology 51

TAXONOMY. 54

Key to the species of Gilia section Gilrnania 55

Gilia latifolia 55

Gilia ripleyi 63

SUMMARY 71

REFERENCES 72

APPENDIXA 78

APPENDIXB. 84

iv LIST OF ILLUSTRATIONS

Figure Page

1. Holotype of Q. latifolia .. 5

2. Q. scopulorum collected from type locality 6

3. Isotype of G. stellata 7

4. Isotype of G. ripleyi 8

5. Leaf variation in G. latifolia 14

6. Leaf variation in G. la.tifolia 14

7. Leaf variation in G. ripleyi 15

8. Leaf variation in G. scopulorum 17

9. Leaf variation in G. stellata 17

10. Herbage pubescence in G. latifolia 19

11. Herbage pubescence in G. stellata 19

12. External flower form in G. latifolia and G. ripleyi 21

13. External flower form in G. scopulorum and G. stellata 21

14. Internal flower form in G. latifolia, G. ripleyi, Q. scopulorum, and Q. stellata 24

15. Capsule form in G. latifolia and Q. ripleyi 26

16. Capsule form in G. scopulorum and G. stellata 26

17. Seed form in G. latifolia, G. ripleyi, Q. scopulorum, and G. stellata . . . . . •.....• 28

18. Seed coat reaction to water in G. latifolia 30

19. Seed coat reaction to water in G. stellata. 30

20. Epidermal pattern in section Gilmania 34

21. Trichome form in section Gilmania 36 V Figure Page

22. Inflorescence in G. latifolia 38

23. Inflorescence in G. ripleyi .. 38

24. Petal venation in section Gilmania 40

25. Pollen form in section Gilmania ..• 42

26. Physiographic map of the southwestern United States . 45

27. Habitat of G. latifolia and~- ripleyi . . . 47

28. Pollen cell of G. ripleyi 52

29. G. latifolia 57

30. Southwestern United States. Distribution of G. latifolia 61

31. G. ripleyi 65

32. Southern Nevada and adjacent south~astern California. Distribution of G. ripleyi ...•...... 69

vi INTRODUCTION

Statement of Problem

The need for a revision of section Gilmania became apparent while the author was making routine identifications within the genus Gilia.

Within Gilia, the taxa occupying the same general area have a strong tendency to exhibit parallel characteristics in response to common habitat. This is especially true among the small-flowered species inhabiting the arid regions of the southwestern United States. In this region, gaps in variation between sympatric species are oftentimes less apparent than the gaps between related subspecies, making definition of infrageneric categories diffi~ilt. Subgenus Gilmania, as constituted by Mason and

Grant (1948), although numbered among the small-flowered, arid region taxa, has been one of the most distinctive groups within the genus. The morphological and ecological relationships between its two members,

G. latifolia and Gilia ripleyi are very close. A precursory study of contemporary keys and descriptions showed this closeness was no longer apparent beca•1se of the inclusion of additional species since 1956. With expansion of the group by Grant and Grant (1956a), two highly divergent groups were recognized within the section Gilmania (so reduced by Grant and Grant [1956a]). Later as a final modification, the original complex was completely obliterated (Grant, 1959) by its inclusion within the highly heterogeneous section Giliastrum, a group which combines a suffrutescent, broad-leaved, spiny, relict perennial with highly evolved, exceed- ingly delicate, linear-leaved annuals. This grouping alone warranted 2 closer analysis. Inasmuch as no explanation was given by Grant and Grant

(1956a) or by Grant (1959) for the modifications other than "in view of the evident relationship. ." in the former publication, this study was undertaken to clarify the morphological, ecological, phylogenetic and cytological relationships of Q. latifolia and Q. ripleyi and to establish possible reasons behind the changes in classication.

Methods and Procedures

Data were obtained from herbarium specimens and from living plants in their natural ranges in Arizona, California, Nevada and Utah.

Type specimens of both Q. latifolia and Q. ripleyi were examined.

The measurement of large structures such as stem, leaf, and inflorescence was with a metric ruler. Plant height was measured from the base of the stem to the tip of the inflorescence. Leaf measurement was from the base of the petiole to the tip of the blade. Measurement of inflorescence was from the node of the lowest flowering branch to the apex of the highest flower. The measurement of small structures such as calyces, flowers, and capsules was by ocular micrometer fitted to the eyepiece of a dissecting microscope. When given, the extremes of dimen- sions were separated from the normal range by parentheses. All ratios are the quotients of the first measurement divided into the second.

Paraffin sections (Gray, 1964) were used to make a series of slides from leaf, stem, root, flower and fruit. Pollen material was incidental on flower slides. Celluloidin peels (Sinclair, 1961) of the ab- and adaxial surfaces of larger leaves were also made. Flowers and epidermal peels taken from pressed plant materials were first softened overnight in Pohlstoffe (Pohl, 1954) before they were mounted.

Cytological material was collected during the summer of 1970 from 3

representative localities in California, Nevada, and Arizona. Chromo-

some number determinations were made from pollen mother cells of flower

buds (ca. 3.5 nun long from tip to base of calyx in Q. ripleyi, ca. 3.2 mm in G. latifolia) possessing anthers in a stage just prior to the in-

itiation of chlorophyll production, the latter an indication of tetrad

formation and overmaturity. Immediately upon collection, the huds·were

fixed in a mixture of one part glacial acetic acid and three parts ab-

solute alcohol and then refrigerated. After twenty-four hours in the

above solution, the buds were stored in seventy per cent alcohol to pre-

vent the brittleness associated with prolonged storage in fixative. The

buds were then pre-stained for a variable period of time in acetocarmine

dye, the anthers removed, and the pollen mother cells liberated. Stan-

dard procedures .(Gray, 1964) were followed in mounting.

A list of herbaria from which specimens were examined is given

below. The abbreviations are, with a few exceptions, the standard ones

given by Lanjouw and Staffleu (1964).

ARIZ University of Arizona, Tucson

ASC Northern Arizona University, Flagstaff

BRY Brigham Young University, Provo (Utah)

CAS California Academy of Sciences, San Francisco

DIX Dixie Junior College, St. George (Utah)

DS Dudley Herbarium, Stanford (California)

GCT Grant Canyon National Monument Herbarium, Toroweap (Arizona)

GH Gray Herbarium, Cambridge (Massachusetts)

JEPS Jepson Herbarium, Berkeley (California)

MO Missouri Botanical Garden, St. Louis

NTS Atomic Energy Commission, Nevada Test Site, Mercury NY New York Botanical Garden, New York 4

ORE University of Oregon, Eugene

POM Pomona College, Claremont (California)

RM Rocky Mountain Herbarium, Laramie (Wyoming)

RSA Rancho Santa Ana Botanic Garden, Claremont

UC University of Ca1iforrtia, Berkeiey

US United States National Museum, Washington, D. C.

UT University of Utah, Salt Lake City

lITC Intermountain Herbarium, Logan (Utah)

WTU University of Washington, Seattle

History of the Section Interpretation of the generic limits within the family Polemoni- aceae has had a long and difficult history. Gilia, the largest genus, has proved especially problematical to taxonomists. With the exception of Phlox and Polemonium, at some time or other, all of the herbaceous genera in the Polemoniaceae have been included in Gilia. The first species of section Gilmania to be described was Q. latifolia. Collected by Dr. C. C. Parry at the "Valley of the Virgen [sic] near St. George," Utah in 1874, and described by Watson (1875) the following year, the type specimen (Fig. 1) was included by its author in section Gilia (=Eugilia). Watson stated, however, that it was of "pecu- liar habit ... not approaching closely any of the other species." Three years later, Q. scopulorum (Fig. 2) was named by M. E. Jones

(1881) from his collections on the lava hills at St. George. It was also placed in the highly heterogeneous subgenus Gilia (elevated from section in 1870 by Gray) as was Q. stellata (Fig, 3), named by A. A. Heller in

1906 from collections made the previous year near Randsburg, California. The treatment of G. latifolia, Q. scopulorum, Q. stellata within subgenus Gilia stood until the specimens (Fig. 4) of G. ripleyi were 5

•.

Fig. 1. Holotype of fr. latifolia collected by Parry (no. 188) near St. Georee (Washington Co., Utah) in 1874 (GH). 6

Fig. 2. �- scopulorum collected by Galway (s.n.) at the type locality near St. George in 1935 (BRY). The original collection was made by Jones in 1881 (MO). 7

Fig. 3. Isotype of G. stellata collected by Heller (no. 7698) at Rands burg (Kern Co., California) in April, 1905 (CAS). 8

Fig. 4. Isotype of Q. ripleyi collected by Ripleyi & Barneby (no. 3992) on the Specter Range (Nye Co., Nevada) in July, 1941 (RSA). 9 discovered in 1940 (Barneby, 1942) from the "limestone cliffs at the south end of the Specter Range," Nye County, Nevada. This discovery prompted

Mason and Grant (1948) to erect a separate subgenus Gilmania for Q. latifolia and Q. ripleyi. Characterized by aristately toothed, obovate to orbicular leaf blades, many, small, brilliant rose, salverform corollas, basally inserted, stout-filamented stamens, and 30 to 100 small, reddish- brown, oval seeds per locule, the subgenus was well separated from the other groups. The section Gilmania was thought to be most closely related to the Q. leptomeria complex (Barneby, 1942; Mason and Grant, 1948) because of the cylindrical capsules, small, non-mucilaginous seeds, and entire, dentately toothed leaves of both groups. However, closer exam- ination indicates that the Q, leptomeria complex has other affinities, and the disposition of that complex is not treated herein.

The subgenus Gilia was redefined and sections Gilia, Arachnion, and Saltugilia formed (Grant, 1950-1954; Grant and Grant, 1954, 1956a,

1956b). The subgenus Gilmania was reduced to sectional level under the newly formed subgenus Gilia, and Q. scopulorum and Q. stellata added

(Grant and Grant, 1956a). The relationship between Gilmania and the rest of the genus then stressed by Grant and Grant was not to Q. leptomeria

(inasmuch as that entity was placed in a separate subgenus Giliandra) but to sections Arachnion, Gilia, and Saltugilia, the other members of the subgenus Gilia~ i.e., plants possessing highly dissected leaves, funnelform corollas, and apically inserted stamens--characters diverging widely from those of G. latifolia and G. ripleyi.

Departing radically from the above classification scheme, Grant

(1959) recognized only sectional rank within the genus--the subgenus

Gilia was rescinded and only its component parts recognized. The newly formed Giliandra was reduced to sectional rank and all other groups were 10 combined into a single section Giliastrum. This latter section as origi- nally described (Brand, 1907) included only Q. rigidula and its relatives, needle-leaved subshrubs and robust, heavy-textured herbs of Texas, adjacent states, and Mexico, characterized by winged seeds, rotate corollas, and prominent stamens (the filaments long and thread-like, the anthers large and sagittate). As expanded, however, the group included plants having in common only filaments inserted low in the corolla rather than in or near the sinuses. Opposing extremes were represented in many other characters, i.e. robust, broad-leaved, spiny perennials combined with delicate, linear-leaved, soft-textured annuals. No other revisions have been attempted until the present work.

Infrageneric Relationships

Although Q. scopulorum and Q. stellata were placed in Saltugilia with the expansion of section Giliastrum to include subgenus Campanulas- trum (Brand) Mason and Grant, subgenus Tintabulum Mason and Grant, and subgenus Gilia section Gilmania Grant and Grant; the section Gilmania was never redefined to exclude Q. scopulorum and G. stellata or reasons for the modification given. In order to formally recognize and justify the segregation of section Gilmania from extraneous species included within it by previous authors, the following key and accompanying discussion are given.

1. STAMENSinserted on the lower corolla tube; anthers sagittate; pollen

white; seeds ovoid, not mucilaginous when wet,~- 0.5 mm long, 30-

100 per locule; leaf blade simple to basally pinnatifid, broadly ob-

long to obovate to .orbicular in outline, margin coarsely serrate-

dentat~ vein system prominently elevated abaxially, lateral veins

strong, continuing past margin as rigid spines (section Gilmania, 11

sens. str.) .... • 2

2. ANNUAL;leaf teeth moderate to short aristate; corolla lobes

brilliant rose within, changing to yellowish or buff without;

stamens 2 long and 3 unequally shorter . . • G. latifolia

2. PERENNIAL; leaf teeth long aristate; corolla lobes brilliant

rose within and without; stamens 1 long and 4 shorter ...

. . . Q. ripleyi

1. STAMENSinserted in corolla lobe sinuses; anthers orbicular; pollen

blue; seeds angular, mucilaginous when wet, ca. 1.5 - 2.0 m.'11 long, 4-

10 per locule; leaf blade pinnatifid to tripinnate, narrowly oblong

to obovate in outline, margin coarsely serrate to pinnately divided

with obovate to linear lobes·, vein system not elevated abaxially,

lateral veins weak, inconspicuously forming an apiculate apex (sec-

tion Saltugilia pro. parte.) .. , ..... • • • • • 3

3. TRICHOMESof the herbage translucent, slender, straight, many

glandular-tipped; corolla 10 - 14.S mm long, tube twice as long

as throat, long exserted . • • . . • Q_. scopulorum

3. TRICHOMESof the herbage white opaque, stout, geniculate, not

glandular-tipped; corolla 6 - 10.5 mm long, tube equaling

throat, included to slightly exserted ... G. stellata

As can be seen from the above key, Q. latifolia and Q_. ripleyi differ from the other members of the section on several distinct points.

These points are as follows: (a) leaves simple, the basal obovate to

O;t"bicular · in G. latifolia and G. ripleyi versus leaves dissected, the basal narrowly to widely oblong to obovate in Q_. scopulorum and Q_. stellata

(Q_. scopulorum under certain conditions produces lower cauline leaves which superficially resemble those of G. latifolia; the lowermost, however, although obscured are once or more times pinnate [Figs. 5-9]); 12

(b) vein system of leaf abaxially elevated and prominent in G. latifolia and Q_. ripleyi versus vein system flush with surface and inconspicuous in G. scopulorum and Q. stellata (Figs. 5-9); (c) teeth of leaf aristate in G. latifolia and G. ripleyi versus teeth apiculate in Q.. scopulorum and G. stellata (Figs. 5-9); (d) trichomes of herbage and lower stem erect, long stipitate-glandular and translucent in G. latifolia and G. ripleyi versus trichomes genicular, non-glandular and white-opaque in

G. stellata (Figs. 10-11); (e) calyx lobes linear and the apex strongly aristate in G. latifolia and Q_. ripleyi versus lobes narrowly ~lliptical and the apex apiculate in Q_. scopulorum and G. stellata (Figs. 12-13);

(£) corolla short salverform in G. latifolia and Q_. ripleyi versus corolla funnelform in Q. scopulorum and G. stellata (Figs. 12-13); (g) corolla lobes long, narrowly ovate to elliptical in Q. latifolia and Q.. ripleyi versus lobes short, suborbicular in Q_. scopulorum and G. stellata (Figs.

12-13); (h) corolla throat indistinct in G. latifolia and Q_. ripleyi versus throat prominent in Q_. scopulorum and Q_. stellata (Figs. 12-13);

(1) corolla tube short, stout, and its outer surface papillate in Q_. latifolia and Q. ripleyi versus tube long, slender, and its outer surface naked in Q_. scopulorum and Q.. stellata (Figs. 12-13); (j) stamens inserted at the base of the corolla tube in G. latifolia and G. riple~i versus stamens inserted in the sinuses of the corolla lobes in Q.. scopulorum and

Q_. stellata (Fig. 14); (k) filaments long, their surfaces papillate in Q_. latifolia and Q.. ripleyi versus filaments short, their surfaces naked in

Q_. scopulorum and Q_. stellata (Fig. 14); (1) anthers sagittate in Q_. latifolia and G. ripleyi versus anthers orbicular in G. scopulorum and G. stellata (Fig. 14); (m) pollen white in G. latifolia and Q. ripleyi versus pollen blue in G. scopulorum and G. stellata; (n) stigma shallow to 13

Fig. 5. Leaf variation in G. latifolia. a. Inyo Co., California~ Inyo, Rixford s.n. (UC). b. Inyo Co., California, Shoshone, Welsh, Matthews & Atwood 9563 (BRY). Fig. 6. Leaf variation in G. latifolia. -. - . ---~-'--- a. I:-:.yo Co., California, Eureka Valley, Alexander & Kellogg 5659 (UC). b. Inyo Co., California, White Mountains, Munz 12719 (RSA). 14

Fi.g. 5

a b Fig. 6 15

==·

__ C"")

==J - -:; :;

--..

a b

Fig. 7. Leaf vAriation in Q. ripleyj, a, Nye Co., Nevada, Spotted Range, Matthews & Reveal 128 (BRY). b, Nye Co., Nevada, Spotted Range, Matthews & Reveal 126 (BRY). 16

Fig. 8. Leaf variation in Q. scopulorum. a. Washington Co., Beaver Dam Wash, Higgins 407 (BRY). b. Washington Co., Black Hill, St. George, Gould 1772 (DS). Fig. ~- Leaf variation in G. stellata. a. San Bernardino Co., California, Twenty-nine Palms, Alexander & Kellogg 93J (TTC). b. San Bernardino Co., California, Providence Mountains, Ferris & Bacigalupi 13298 (DS). c. Inyo Co., California, Shoshone, Welsh, Matthews & Atwood 9560 (BRY). 17

(.)

.0 18

Fig. 10. Herbage pubescence inf. latifolia. Inyo Co., California, Titus Canyon, Welsh, Matthews & Atwood 9637 (BRY). Fig. 11. Herbage pubescence in G. stellata. Inyo Co., California, Shoshone, Welsh, Matthews & Atwood 9560-(RRY). l 9

Fig. 10

Fig. 11 20

Fig. 12. External flower form. a. Q. ripleyi. Nye Co., Nevada, Spotted Range, Matthews & Reveal 126 (BRY). b. G. latifol:ia. Inyo Co., California, Titus Canyon, Welsh, Matthews & Atwood 9637 (BRY). Fig. 13. External flower form. a. Q. scopulorum. Washington Co., Utah, Beaver Dam Wash, Higgins 407 (BRY). b. G. stellata. Inyo Co., California, Daylight Pass, Welsh et. al. 9611 (BRY). 21

a h

Fig. 12

Fig. 13 22 more deeply lobed in Q. latifolia and Q. ripleyi versus stigma deeply

cleft in G. ~copulorum and G. stellata (Fjg. 14); (o) capsules cylindric

to narrowly ovate with 30 to 100 seeds per locule in G. latifolia and G.

ripleyi versus capsules globular to broadly ovate with 4 to 10 seeds per

locule in Q. scopulorum and Q. stellata (Figs. 15-16); (p) seeds oval,

small (barely 0.5 mm long axis, 0.3 - 0.4 mm short axis), and nonmuci-

laginous in Q. latifolia and Q. ripleyi versus seeds oblong-angular,

large (over 1.5 mm long axis, 0.8 - 0.9 mm short axis), and densely muc-

ilaginous in Q. scopulorum and Q. stellata (Figs. 17-19).

The shared characteristics are few and were found to be ecologic-

ally rather than morphologically controlled. It was perhaps their joint

membership in the spring flora of the Larrea divaricata-Ambrosia dumosa

communities of the Mojave and Colorado deserts of the southwestern United

States which first brought about their comparison. For in a poor water-

year (as was found in a 1970 collecting trip through Titus Canyon, Inyo

County, California), many of the specific differences between sympatric

populations of the two species are masked by depauperate growth, i.e.,

both plants were dwarfed, spindly, and sparsely leaved. Even the resem-

blance between the general leaf form of the lower cauline leaves of G.

scopulorum and the basal ones of G. latifolia was apparently dependent

on environmental rather than genetic factors. Although the lower cau-

line leaves of Q. scopulorum are typically of greater breadth than are

the basal, the cauline are shallowly toothed only on less vigorous spec-

imens. Plants which grow to full stature are readily separated from Q. latifolia even in a vegetative state by their greater height (to 6 dm),

djssected, narrowly eJongateleaves, stouter stems, and strict branching. The relationship of Q. stellata to Q. latifolia and G. ripleyi is even more obscure, and apart from its duration as a winter annual and its 23

Fig. 14. Internal flower form. a. G. latifolia. Nye Co., Nevada, Ash Meadows, Matthews & Reveal 2154 (BRY). b. Q. ripleyi. Nye Co., Nevada, Spotted Range, Matthews & Reveal 127 (BRY). c. G. scopulorum. Washington Co., Utah, Beaver Dam Wash, Higgins 407 (BRY). d. G. stellata. Clark Co., Nevada, Searchlight, Cronquist 9968 (BRY). 24

---~ I .I I u I ,,,.,,.------______J -✓ 26

Fig. 15. Capsule form. a. G. latifolia. San Bernardino Co., California, Shoshone, Welsh, Matthews & Atwood 9564 (BRY). b. ~- riplex~- Nye Co., Nevada, Spotted Range, Reveal 1768 (NTS). Fig. 16. Capsule form. a. Q. scopulorum. Mojave Co., Arizona, Toroweap Valley, Matthews 112 (BRY). b. G. stellata. Inyo Co., California, Shoshone, Welsh et. al. 9560 (BRY). 26

a b

Fig. 15

.,._

Fig. 16 27

Fig. 17. Seed form. a. G. latifolia. Nye Co., Nevada, Ash Meadows, Reveal & Matthews 2154 (BRY). b, G. ripleyi. Nye Co., Nevada, Spotted Range, Reveal 1768 (NTS). c. G. scopulorum. Clark Co., Nevada, Overton, Clokey 5909 (DS). d. G. stellata. Inyo Co., California, Daylight Pass, Welsh, Matthews & Atwood 9611 (BRY). N 00 d C - 17 Fig. b , , , a • • 29

Fig. 18. Seed coat reaction to water in Q. latifolia taken at 125 magnification after a 5 minute immersion. Nye Co., Nevada, Ash Meadows, Reveal & Matthews 2154 (BRY). Fig. 19. Seed coat reaction to water in G. stellata taken at 60 magnification after a 5 minute immersion. Inyo Co., California, Daylight Pass, Welsh, Matthews & Atwood 9611 (BRY). 30

Fig. 18

Fie. 19 31 placement in the creosote bush association (its range, however is much more extensive--covering the entire Sonoran Desert area of Baja, California and Sonora, Mexico), it has little in common with Q. latifolia. Gilia ripleyi is related to Q, scopulorum and Q. stellata only through generic equivalence--they would be placed at the opposite ends of a contin- uum. Thus, based on the above differences, the section Gilmania is redefined to include only Q. latifolia and Q. ripleyi.

Grant (1959) reviewed the infrageneric taxa within Gilia and placed

G. latifolia and Q. ripleyi within the section Giliastrum. This section, however, is not a taxon founded on phylogenetic proximity in the opinion of the author, but rather a depository for poorly known forms. At least two distinct species complexes compose the group.

The delicate, linear-leaved Gilias of the G. campanulata complex have been variously separated on the basis that they represent the reduc- tion-series end-products of several distinct lines rather than a commonly derived group. It is not within the scope of this paper to consider the relative merits of such a theory; however, whether closely related or not, the species placed within the complex definitely do not belong with Q. latifolia and G. ripleyi. Their spreading habit, delicate, soft-textured herbage, diffuse axillary inflorescences, and few seeded, globose capsules are in direct opposition to the ancestral rather than derived condition of the Q. latifolia complex and indicate other affinities.

The Q. rigidula complex, although more closely related to G. latifolia and Q. ripleyi than is the Q. campanulata complex by virtue of their often suffrutescent habit and heavy-textured leaves, are well separated from Q. latifolia and Q. ripleyi by their broadly campanulate calyces, broadly lobed, rotate corollas, thread-like, equal filaments, acerose to deeply palmately or pinnately dissected leaves, and winged 32 seeds.

Thus, based on the above differences G. latifolia and G. ripleyi are removed from section Giliastrum and set apart in a separate section

Gilmania.

General Morphology

Habit

The plant consists of a taproot, usually quite twisted and variable in degree of branching, and an erect stem, simple or with several branches from near the base. In Q. ripleyi, a perennial, the root is corky and extends into a caudex, whose apex bears a rosette of leaves and gives rise to the flowering stem.

Herbage

The stems are herbaceous (the persistent base of Q. ripleyi and the lower portions of robust fonus of Q. latifolia are ligneous), moderately to densely glandular-stipitate, and rise from a cluster of basal leaves (Q. latifolia under shade conditions is leafy to the inflorescence). In Q. ripleyi the flowering stems of previous years are often persistent as straw-colored, fragile axes covered by chartaceous, dried leaves at the base.

The leaves are alternate and simple. The basal leaves are strongly compact and erect in the case of Q. ripleyi to loose and ascending in

G. latifolia. In shape, the basal leaves are broadly oblong to obovate to orbicular (less ~ommonly rhombic) in Q. latifolia to more uniformly obovate to orbicular in G. ripleyi, with a flat upper surface (the lamina slightly puckered near the veins in G. latifolia), and a prominent, elevated, pinnate vein system beneath. The margin on both forms is coarsely serrate-dentate (to incised or basally pinnatifid with 1 to 3 33 pairs of oblong lobes in G. latifolia) with 3-4 triangular to acuminate teeth on each side (minor teeth are often formed upon the major by subsequent branching of the lateral veins). The teeth of both species are coursed by strong veins which extend past the margins as rigid spines.

The leaves are slightly succulent, modetately to densely glandular- stipitate (especially prominent on the leaf margins and abaxial veins) and bright green in Q. ripleyi to deeper and more muted green in Q. latifolia. The cuneate (to truncate in basally pinatifid forms of Q. latifolia) base of the blade is persistent as a narrow wing on both sides of the adaxially grooved, slender petiole. The cauline leaves are greatly r·educed (except in shade forms of G. latifolia) and subsessile. Their lateral' teeth, if present, decrease in number until in the inflorescence, the bracts become narrowly laminate, entire or basally toothed, glandular-stipitate spines.

The epidermal cells of the ad- and abaxial leaf surfaces are extremely small, 50-60 micrometers long and 40-50 micrometers wide, oblong to orbicular or even irregular in outline and prominently wavy-margined.

Over the vein systems and on the axes of the plant, the cells elongate parallel to the long axis of the clothed structure, and their margins straighten (Fig. 20).

The stomata are elliptical in outline, 25-30 micrometers long and ca. 15 micrometers wide, numerous and crowded,~- 50 micrometers between adjacent stomata. No subsidiary cells are present, with 3 to 5 epidermal cells bordering on each stomata.

Pubescence

The trichomes are multicellular and glandular-tipped, with the glands, themselves, several-celled. The hairs change from long-tapering 34

J = 10 micrometers

a b

Fig. 20. Epidermal pattern in section Gilmania. a, G. latifolia, Inyo Co., California, Shoshone, Welsh, Matthews & Atwood 9559 (BRY). b, Q, ripleyi, Spotted Range, Reveal & Matthews 126 (BRY). 35 and slender on the herbage and lower stem to short and stout in the inflorescence. They are borne in abundance perpendicular to the epidermal surface, and give to Q, latifolia and Q. yipleyi their characteristic rank odor and fragile, viscid appearance (Fig. 21).

Inflorescence

The inflorescence is compound cymose, with the primary and secondary flo:wering branches alternately disposed (monochasia), and the ul- timate units opposite branched (dichasia). Each division is subtended by a spinose bract. The numerous flowers are borne singly on delicate, moderately to densely stipitate-glandular, widely divaricate pedicels of various lengths. The longer stalks greatly exceed the flower length and

the shorter are almost lacking. The peduncle is characteristically branched only above in Q. Fipleyi, or to much branched from near the base

in G. latifolia, The lower. portion of the peduncle is naked or endowed with one to four widely spaced, bract-like cauline leaves in G. ripleyi, while in G. latifolia it is highly variable although commonly the cauline leaves are present and highly reduced (Figs. 22-23).

Flower

The flower is hermaphroditic, radially symmetrical and pentamerous, with a three-carpellate ovary and three-lobed stigma.

The narrowly campanulate calyx is five-parted to the base. The

linear lobes are united in the lower one-half by hyaline membranes in the sinuses. The lobes are all equal and moderately to densely glandular-stipitate. Their midribs are prominent and continue as aristate spines apically. In fruit, the calyx is accrescent by distention of the membranes.

The corolla is convolute in bud, expanding to short salverform in 36

= 1 mi 11 imeter

Fig. 21. Trichome form in section Gilmania as represented by G. latifolia, Inyo Co., California, Shoshone, Welsh, Matthews & Atwood 9559 (BRY). 37

Fig. 22. Inflorescence fonn in G. latifolia. Nye Co., Nevada, Ash Meadows, Reveal & MatthPW~ 215u (:RRV), Fig. 23. Inflorescence form in G. ripleyi. Nye Co., Nevada, Spotted Range, Matthews & Reveal 126 (BRY). 38

N N

bl) •r( r:..

'1111;111lnn1111111111 1111111,-rrtm,li,,, 1,,,i fhntmr� 39 flower. The long, scarcely-overlapping lobes are elliptical to narrowly

oval in outline, and are constricted to a short neck at the base (promi-

nent in Q_. ripleyi). The stout tube is minutely papillate without and

included in the calyx. Coloration of the lobes within is brilliant rose, with the outer surface changing to yellowish or buff in G. latifolia and

remaining constant in Q. ripleyi. The inconspicuous throat is buff

(often faint in G. ripleyi) and the tube, white.

The petal venation pattern of Q. latifolia and Q. Fipleyi is

strikingly similar and does not closely approach any of the other species

studied by Alva Grant (Grant, 1959). The primary vein of each petal divides upon leaving the receptacle into three prominent traces which run

to the base of each lobe. There the two lateral traces steeply incurve and unite with an ascending branch from the midvein and a peak is formed on each side of the petal axis. Five to six closely spaced secondaries

arise from the side and apex of these peaks and ascend, slightly radiat-

ing, to the apex, rebranching once in G. ripleyi, remaining straight to

the margin in Q. latifolia (Fig. 24).

The nectary is a greenish, obscurely five-lobed disc surrounding

the ovary base. It is believed by Dawson (1936) to be the enlarged, fused bases of a now vestigial second whorl of stamens--the vascular supply of which is still present in the more primitive tropical members of the

family.

The stamens are briefly epipetalous, their anthers sagittate-

erect and four chambered, and their filaments stout, minutely papillate,

and unequal--four short, one long in G. ripleyi and three short, two

long in G. latifolia.

The pollen grains are white, slightly flattened spheres with four

circular pores in opposing pairs on the equator. They are unusually 40

I I I I I I I I I I I I I I l I I I I I I I I I I I I I I I I I I I I ., I I I I I, I I I I I I I I I I I i I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I mm I I I I I I I I •------I -----.1I L______t, I•

a b

Fig. 24. Petal venation in section Gilmania. a, G. latifolia, Nye Co., Ash Meadows, Matthews & Reveal 2154 (BRY). b, G. ripleyi, Nye Co., Spotted Range, Matthews & Reveal 127 (BRY). 41 small, 25-30 micrometers in diameter, pollen less than 50 micrometers

typically associated with the low growing delicate members of the genus.

The exine is two layered--the round pores of the inner layer are over- hung by the smaller aperatures of the outer, thicker layer. Conspicuous

ornamentation is lacking, with the surface appearing only finely grainy

(Fig. 25).

The style is minutely papillate, included, and persistent on the

capsule. Stigmatic reflection is at or slightly below the level of the

anther of the longest stamen. The stigma is introrse (the inner surface

of the lobes stigmatic) and cleft in G. latifolia to less deeply lobed in

G. ripleyi.

The floral vascularization of~- latifolia and Q. ripleyi follows

the general pattern of many of the less-advanced members of the family as determined by Dawson (1936). The vascularization develops in the

following sequence: (a) sepal traces are three per lobe and are d~riv~d

from the ten traces initially given off by the sol-id cylindrical stele of the pedicel--with five forming the midribs and five, the compound lateral

traces (these latter dividing into the two lateral veins of two adjacent,

rather than of a single, sepal); (b) petal traces are one per lobe and are derived from the compound lateral sepal traces; (c) stamen traces are

one per filament and are derived from a portion of the remaining vascular tissue (moving outward slowly, they appear to be alternating members of the petal whorl until mid-tube where they become distinct); (d) carpel

traces are four per locule and are derived from the remaining recep- tacular tissue, with six strands forming the dorsal and compound lateral carpel traces. The three fused ventral traces remain as a three-corn~red

central core of vascular tissue; (e) ovule traces are one per locule,

and are derived from the ventral carpel core; (f) style traces are three, 42

I• 10 micrometers

a b

Fig. 25. Pollen form in section Gilmania as represented by Q. latifolia, Nye Co., Nevada, Ash Meadows, Reveal & Matthews 2154 (BRY). a, Equatorial view. b, Polar view. 43 and are derived from a continuation of the dorsal carpel traces.

Fruit

The capsule is three-loculed and axillary in placentation. Cylin- dric to narrowly ovate in shape, it is commonly less than or equal in length to the aristate calyx lobes in Q. latifolia and is commonly much surpassed in length by the calyx lobes in Q. ripleyi.

Seed

The seeds are numerous (30 to 100 per locule), small, and reddish- brown. Their coats are finely tuberculate and unaffected by water, i.e., spiracles or slime are not produced.

Embryo

The embryo is green, oblong to slightly spatulate, and rests in the middle of a thin endosperm.

Distribution and Ecology

The genus Gilia occurs from southern British Columbia southward into Mexico and eastward across Texas to the south Atlantic Coast. There it is represented by one species, Q. rubra, adventive as far north as

Massachusetts, with Gilia being absent·from Eurasia, Alaska, and north- western Canda. It reoccurs disjunctly in a secondary center of distribution along the west coast of South America, where on the coastal plains and higher mountain zones of Peru, Hip;lito L6pez Ruiz and J6se Pav;n y

Jimenez found and described Q. laciniata, the type of the genus. The primary distribution center of section Gilmania and the genus as a whole, however, is in the arid western United States (Fig. 26).

Gilia latifolia is a widespread winter annual of southern California, western Arizona, southern Nevada and extreme southwestern Utah. Gilia 44

Fig. 26. Physiographic map of the southwestern United States by Ruiz (1957). 45

• ------.&..--.,__-:200 miles o 50 46 ripleyi is a narrow endemic perennial of the limestone outcrops of the stark Paleozoic mountain ranges of southern Nevada and adjacent southeastern California. Both members occupy xerophytic habitats--Q. latifolia at low to middle elevations, -250 to +7000 feet, and Q. ripleyi at the narrow interposed elevationai band of 3100 to 5000 feet.

The Mojave Desert, inhabitated by both species, is characterized by extreme aridity and great temperature fluctuations. Little precipitation falls during the year (2-8 inches mean annual), with some years de- void of precipitation. The sources of moisture are effectively cut off by the South Coast and Peninsular ranges of the coastline and the Sierra

Nevadas to the interior. Most moisture falls as sporadic thunderstorms during August and September and gentler, general rains during the winter months. Summer rains are lacking in the west and south and highly localized and light in the northeastern portion of the desert.

The summers are hot (mean maximum 100-110° F. annually) and the winters are mild (mean minimum 30-42° F. annually) (Munz, 1959). Diur- nal temperature fluctuations are great, as the parched, unshaded soil quickly absorbs heat in the morning and rapidly dissipates it into the cloudless sky at night.

Where their ranges overlap in southern Nevada and adjacent southeastern California, the two species are often found in close proximity,

G. latifolia in the Larrea divaricata-Ambrosia dumosa connnunity of the valley floor and deeply cut washes, and Q. ripleyi directly above on the near-sterile limestone outcrops, commonly shared only by Buddlei.a utahensis, Eriogonum sulcatum, !• intrafractum, Cryptantha confertifolia, and a few strongly xerophytic shrubs of the Compositae (Fig. 27). Although closely tied geographically, Q. latifolia and Q. ripleyi represent contrasting extremes in adaptation to the arid environment Fig. 27. Habitat of Q_. latifolia .::md Q_. riplcyi (Nye Co., Nev.-ida, Red Mountain, Spotted Range). Q_. latifoJ.ia grows in the Larrea divaricnta-Ambrosia du�osa cormnunity of the valley floor and bajadns, ,,1'ith Q_. ripleyi inhabiting the limestone formations above. � -.., 48 which surrounds them. Gi.lia ripleyi is a "chasmophyte," a member of the calciphile flora whose combined ranges reach from the chasms and preci- pices of the mountain ranges near Darwin (Inyo County, California) east to St. George (Washington County, Utah) and south to Kelso (San Bernar- dino County., California). Barneby (1942) stated:

To this region belong such memorable species as Penstemon petiolatus . . . Phacelia perityloides . . . and the beautiful Maurandia petrophila ... all chasmophytes with broad, toothed leaves, taxonomically isolated and evident relicts of a mes- ophytic vegetation of immense antiquity and it is believed that Gilia [ripleyi] is an unrecognized member of this august and ancient company.

These "chasmophyte" species apparently arose prior to the massive Plio- cene upheavals which fonned the Sierra Nevada and Peninsular ranges, effective barriers to the flow of cooling winds and rains into the interior of southern California. Adapted to a moist climate both morphologically and physiologically, they have adjusted in various ways to the altered environmencal conditions. To che normal life cycle of G. ripleyi, a period of dormancy has been added. The short vegetative growth arises very early during the cool winter months from suffrutescent caudices; with the plants falling dormant during the normal desert blooming period beginning in early March and April. Finally, in the late surrnner months beginning in mid-June and proceeding until September, a slender, naked flowering stem is sent up in aburst of growth and abundant seed is set.

Rapid growth and maturation of reproductive structures are solely dependent on the occurrence of sporadic, summer rains. With the vegetative portion already partially deteriorated, the plant rapidly dies back until winter and the beginning of a new cycle of growth.

In contrast, Q. latifolia is a winter annual, an adaptation to the aridity of post-Pleistocene southern California. Unlike Q. ripleyi which requires both winter and summer rains to complete its full life 49 cycle, flowering and fruiting in£. latifolia is spontaneous under the stimulus of the rising temperatures and falling soil moisture of late spring. Thus,£. latifolia is able to extend into the Sonoran Desert of southern California, a region characterized by its lack of summer showers.

Q. ripleyi, in contrast, remains in the northeastern boundaries of the

Mojave Desert where summer rains, although sporadic, occur.

Furthermore, although typical of gravelly washes with sand fillings, G. latifolia grows on any well-drained coarse soil. Q. ripleyi occurs only in the talus dust of the harshest exposures of near vertical limestone, an indication of its inability to compete with better adapted forms in more favorable soil (and it may well be obligate on calcareous soils as the seedlings grown for cytological material of£. ripleyi all suddenly perished in rosette stage, while those of Q. latifolia plants in similar sterile, pre-moistened sand lots flowered and set viable seed).

Due to the small size of seeds, their lack of adhesive ornamentation and mucilage, and the fragile nature of their pedicels, wind is the logical means of seed distribution.

Phylogeny

The family Polernoniaceae as a member of subclass Asteridae is considered to have evolved from tropical, woody Ranalian ancestors through either subclass Rosidae (Cronquist, 1968) or Dilleniidae (Grant,

1959) to its present stage of development. The latter theory is based on the similarities between the emerging characters of certain members of

Dilleniidae (Theales, Ericales and Ebenales) and those of Asteridae, i.e., unitegmic, tenuinucellate ovules, sympetalous corollas, isomerous stamens, and axile placentation. The former theory is based on a like, but less prominent transition series, repeated development of trinucleate 50 pollen characteristic of Asterales, and centripetal stamen development

(centrifugal stamen development is limited only to subclass Dilleniidae

and Caryophyllidae with no member of Asteridae having stamens opposite

the corolla lobes). Other schemes focusing on Caryophyllidae have been

proposed (Bessey, 1915), but the basal or free central placentation,

betalain pigmentation, and staminal nectaries of this subclass exclude

it from consideration.

Within the subclass Asteridae, the closest associate to Polemon-

iaceae is Hydrophyllaceae (the Waterleaf family) by virtue of their sim-

ilar habit (erect herbs and shrubs), floral plan (radially symmetrical,

pentamerous flowers, deeply cleft calyces, tubular to campanulate corol-

las, alternate, epipetalous stamens, 2-3 carpellate ovaries [the former

only rarely developed in Polemoniaceae], and capsulate fruit), and dis-

tribution (centered in western North America).

Qithin the family, trends in evolucion are distinct. The

tropical species, although specialized in certain respects, generally

simulate the primitive condition by (a) woody, perennial habit; (b)

simple broad, heavy-textured leaves; {c) herbaceous, little fused calyces;

(d) large, symmetrical, open throated corollas; (e) basally inserted stamens; (f) large, elongate, versatile anthers; (g) thread-like, equal filaments; (h) prominently lobed, vascularized nectaries; (i) elongate,

trilocular, dehiscent capsules; (j) numerous seeds; and (k) nine pairs

of chromosomes. The advanced condition is found in the temperate species

(a) herbaceous, annual h_abit; {b) compound or simple and greatly reduced

leaves; (c) partially scarious, united calyces; (d) small or irregular, narrow throated corollas; (e) apically inserted stamens; (f) small, orbicular, stationary anthers; (g) stout, unequal filaments; (h) incon-

spicuous, non-vascularized nectaries; (i) globose, bilocular, indehiscent 51

capsules; (j) few seeds; and (k) six pairs of chromosomes.

Within Gilia, the primitive condition is best represented in two sections, Gilmania and Giliastrum (as defined by Brand). Their primitive and advanced characteristics, however, are for the most part reversed.

Whereas section Gilmania departs from the primitive condition in possessing well-united, partially scarious calyces, salverfonn corollas, stout, unequal filaments and stationary anthers, section Giliastrum departs from the ancestral characteristics in possessing acicular to pinnately or palmately parted leaves, globose, often few seeded capsules, and winged, mucilaginous seeds (although the two lattermost characters are often found well developed in the tropical species, such characteristics are logical- ly examples of an independent rate of organ evolution rather than primitive conditions).

The sections Gilmania and Giliastrum probably arose from members of the Madro-Tertiary flora, a woodland vegetation of subtropical trees and shrubs which migrated across Mexico into southern California during the mid-Miocene in response to a drying climate. Subsequent isolation of the western and eastern members of the flora by more severe climatic changes of the post-Miocene period formed the basis for sectional diver- gence.

Cytology

Based on the meiotic chromosome number determinations of G. ripleyi made by the author and of Q. latifolia made by Alva Grant (Grant,

1959), the chromosome numbers of Gilia, section Gilmania are as follows:

G. ripleyi N=9 (Fig. 28)

Nevada, Nye Co., Spotted Range, 1 mile north of Mercury,

Matthews & Reveal 126, 10 June 1970 (BRY). 52

{ = 1 micrometer

------Fig. 28. Pollen cell of G. ripleyi enlarged from a camera lucida drawing made under an oil immersion magnification of 1000. 53 G. latifolia N=l8

California, Riverside Co., 9 miles east of Mecca, Grant &

Grant 17619, 1 April 1952 (RSA, CAS).

The basic chromosome number of Gilia is nine. The basic chromosome number of eight is found only in a few species of section Giliandra.

Polyploidy is varied in occurrence within Gilia and is limited primarily to tetraploidy. Higher polyploid numbers are recorded only twice: N=

50 for G. micromeria of section Giliandra and N=72 for G. crassifolia of section Arachnion. Polyploidy is distributed in the sections of Gilia as follows: Arachnion 41%, Giliandra 20%, Giliastrum 20%, Gilmania 50%, and Saltugilia 12%.

The close morphological similarity of G. latifolia and G. ripleyi, the relict nature and basic chromosome number of the latter, and the tetraploidy of the former seem to indicated that G. latifolia is a direct allo- or autopolyploidic descendent of Q. ripleyi. The karyotype anal- yses and breeding experiments needed to substantiate this hypothesis, however, are not tenably pursued within the normal time allotted a

Master's research program and will be completed at some future time. 54

TAXONOMY

Gilia Ruiz & Pav~n section Gilmania (Mason & Grant) Grant & Grant.

Section Gilmania (Mason & Grant) Grant & Grant, El Aliso 3: 299.

1956.

Subgenus Gilmania Mason & Grant, Madroio 9: 205. 1948.

Erect or slightly ascending annual or perennial herbs, branched

from base or above. Leaves heavy-textured, alternate, borne chiefly be-

low mid-stem, usually in basal tuft, widely oblong to obovate to orbicu-

lar in outline, gradually tapering at base into narrowly winged, adaxial-

ly grooved petiole, teeth serrate-dentate to basally pinnatifid, moder-

ate to 1ong aristate, abaxial surface of leaf flat, abaxial surface con-

spicuously pinnate-nerved, lateral veins strong, both surfaces moderate-

ly to densely stipitate-glandular, the cauline leaves much reduced and

bract-like, rarely leafy to inflorescence in shade forms. Inflorescence

cymose, loosely paniculate to subglomerate, flowers numerous, borne on

filiform, divaricate pedicels of various lengths, from much exceeding

flower length to obsolete, 1-2 per node subtended by bract. Calyx nar-

rowly campanulate, lobes linear, equal, apex aristate, joined in lower

one-half by hyaline membranes, accrescent in fruit. Corolla salverform,

· lobes long, elliptical to narrowly oval, apex acute to slightly rounded,

margin entire or slightly irregular, inner surface of lobe brilliant

rose, outer surface same or buff, base constricted, sinuses prominent,

throat inconspicuous buff, tube stout, only slightly exceeding lobes,

minutely papillate without, white. Stamens unequal, inserted subequally 55 on lower tube, included, filaments stout, minutely papillate, anthers sagittate, white, subequal to less than half the length of shortest filament. Capsules oblong to narrowly ovate, hard walls finely reticulate without, smooth and glossy within, dehiscing loculidically from top to bottom, remaining attached basally, slightly recurving apically, many seeded, 30-100 per locule, equaling to much exceeded by aristate calyx lobes. Seeds ovoid, small, reddish-brown, inconspicuously tuberculate, not producing spiracles or slime when moistened.

Sectional type: G. latifolia S. Wats

Key to the species of Gilia section Gilmania

1. ANNUAL;leaf teeth moderate to short aristate; corolla lobes bril-

liant rose within changing to yellowish or buff without; stamens 2

long and 3 unequally shorter .•.. G. latifolia

1. PERENNIAL; leaf teeth long aristate; corolla lobes brilliant rose

within and without; stamens 1 long and 4 equally shorter ..

G. ripleyi

Gilia latifolia S. Wats. (Fig. 29)

G. latifolia S. Wats., Amer. Natur. 9: 347. 1875.

Navarretia latifolia Kuntze, Rev. Gen. Pl. 2: 433. 1891.

Annual, erect or ascending, (0.3) 0,5-2.8 (4.5) dm tall, rank odored. Stems slender to much thickened and rigid, branching only from above to much branched from near base, crisply pubescent with multicellular, translucent, glandular-tipped hairs 0.4 - 4.0 mm long. Leaves alternate, leafy below middle to subrosulate, rarely leafy to inflorescence in shade forms, ascending or less frequently erect; lower leaf blades ovate to oblong to orbicular (rarely rhombic) in outline (1.0)

2.5 - 5.0 (12.9) cm long, (0.8) 1.0 - 4.0 (7.6) cm wide including spines, 56

Fig. 29. G. latifolia. Habit drawing is actual size. Isolated fruit and fluwer are twic~ actual size. 57

/ ,,l: ·, L

•,J 58 ratio blade length to width (0.8) 1.0 - 1.5 (2.8), abruptly to gradually

tapering below to slightly winged, adaxially grooved petiole (0.0) 1.0 -

2.5 (4.8) cm long, leaf margin inconspicuously undulate, coarsely serrate-

. dentate to irregularly incised, teeth deltoid, moderately to shortly aris-

tate, spines extensions of lateral veins (0.5) 1.0 - 2.0 (2.5) mm long,

often alternating long and short by development of secondary teeth, up-

per surface of leaf flat to slightly puckered at veins, lower surface of

leaf coursed by prominently elevated, pinnate vein system, the midrib

giving off acutely 3 to 5 pairs of parallel, alternating, thick, straight

secondaries, pubescence as on stems, more dense adaxially; upper cauline

leaves much reduced, subsessile; bracts aristate-lanceolate, entire to

basally toothed, 0.5 - 5.0 mm long, abaxial veins prominent. Inflo-

rescence (0.8) 2.0 - 8.0 (14.5) cm long, peduncle (0.5) 2.5 - 9.0 (15.7)

cm lon~, ratio inflorescence to peduncle length (0.5) 0.7 - 2.5 (3.6);

flowers numerous, commonly two per node borne singly on moderately to

densely stipitate-glandular, divaricate, filiform pedicels (0.0) 0.2 -

2.5 (4.2) cm long, lengths are often various on same plant. Calyx

narrowly campanulate (3.3) 3.5 - 6.0 (7.1) mm long, 1.5 - 2.5 mm wide

in flower, moderately to densely stipitate-glandular, cleft to base,

united in lower half by hyaline membranes; lobes linear (1.4) 2.0 -

3.0 (4.0) mm long, apex long aristate, equal to or exceeding the corolla

tube; calyx expanding in fruit by distention of membranes, subequal

to greatly overtopping fruit. Corolla short salverform, convolute in

bud, (4.6) 5.6 - 8.6 (11.8) mm long; the lobes narrowly oval, incon-

spicuously constricted to parallel basal portion, apex acute, margins

smooth, (1.1) 2.0 - 3.5 (4.3) mm long, 1.2 - 4.4 mmwide, ratio lobe

length to total corolla length (1.9) 2.3 - 3.6 (5.0), lobes bright rose 59 within, buff or yellowish without; throat indistinct, buff; tube stout,

(3.4) 4.0 - 5.5 (6.7) nun long, equaling calyx width, minutely papillate without, white. Stamens included, unequally affixed to lower part of corolla tube, unequal in length, two longer, 4.4 - 4.8 mm, two intermediate, 3.7 - 3.8 mm, one short~r, 3.4 - 3.5 mm; filaments stout, cream or white, minutely papillate; anthers sagittate, 1.1 - 1.5 mm long; pollen spherical, very small, 25 - 30 microns in diameter, exine finely granular with four equatorial, orbicular pores in opposing pairs.

Ovary ovate-oblong, minutely stipitate-glandular, _£!, 2.0 mm long. Style

(0.8) 1.1 - 2.5 (3.5) mm long, minutely papillate, often persistent on capsule; stigma three-cleft (0.5) 0.7 - 1.1 (1.8) mm long, introrse, reflexing at or slightly below anther level, ratio stigma to style length (0.9) 1.5 - 3.0 (4,8). Capsule cylindric to narrowly ovate,

(2.2) 3.8 - 6.8 (7.5) mm long, (1.0) 2.1 - 3.8 (4.4) nun wide, ratio capsule length to width (1.1) 1.5 - 2.0 (3.5); walls hard, £i1~ly reticulate without, smooth, glossy within, dehiscing loculicidally from top to bottom, remaining attached below, slightly recurving above; the capsule equal to or greatly exceeded by aristate calyx lobes. Seeds ovoid, deep reddish-brown, inconspicuously tuberculate, very small,

0.6 - 0.7 mm long axis, 0.4 - 0.5 nun short axis, 30 - 100 per locule, producing neither spiracles nor slime when moistened. N = 18.

Holotype: Utah, Washington County, "Valley of the Virgen [sic] near St. George," Parry 188, 1874, GH. Isotypes: NY, MO.

Distribution: ARIZONA, Mojave and Yuma counties; CALIFORNIA,

Kern, I~perial, Inyo, Riverside, San Bernardino, San Diego, and Ventura counties; NEVADA,Clark, Esmeralda, Nye, and Washoe counties; UTAH,

Washington County. Sandy or gravelly washes of valley floors and deep canyons, limestone or metamorphic talus, -250 to +7000 feet (Fig. 30). 60

Fig. 30. Southwestern United States. Distribution of G. latifolia. 61

-.·------. ------,------·-·r·------. . GJ? I V '-·-·-·-·-i I I I ;• I ! I I I I ! I I ___,. __ _ :.,. ·-·-·-·-·-·-·-·-i • • I • • i I .I • • • I • I • • I • i I •• ••• • I • • • • • •• • --~----·-· • ...... t_ • ·,.,., ·,., ...... -·-·-·-- 62 Associated with: Larrea divaricata, Ambrosia dumosa, Atriplex confertifolia, ~. hymenelytra, Fouquieria splendens, Buddleia utahensis,

Narna demissum, Phacelia crenulata, f. pedicellata, f. fremontii, Calan- drinia ambigua, Yucca schidigera, Stanleya pinnata, Malvastrum rotundi- folium, Oenothera brevipes, Gilia scopulorum; Salvia funerea, Dalea cal- ifornica, Chenopodium botrys.

Flowering time: Precocious flowering (January to February) is associated with very low elevations, -250 to +500 feet, in the southern portion of the range, retarded flowering (June to September), with the higher elevations, 5000 to 7000 feet, in the north. The normal flowering period of Q. latifolia, however, is early March to mid-May, with the seed set, and most plants withered by mid-June.

Discussion: As a widespread annual of great elevational di- mensions, Q. latifolia occurs in a variety of habitats. In response, the species has become varied morphologically. 'i'he differences, however, are often as striking between plants a few feet away as are those of plants hundreds of miles apart. No set of character combinations worthy of infraspecific rank are known to be present on a populational level.

Representatives of the types of variation found within G. latifolia are as follows: (a) extremely robust forms exceeding 4 dm in height and intensely woody at the base; (b) strongly rosetted forms, leaves basally dissected into 2 to 3 oblong, aristate-tipped lobes, inflorescence glomerate, pedicels little developed to lacking; (c) much-branched forms, leafy to middle, leaves small, deltoid to rhombic, inflorescence exceed- ingly dense glandular-stipitate, capsules much reduced, prominently over- topped by calyx; (d) orbicular leaved forms, leaves alternate, widely spaced to inflorescenc.e; (e) near decumbent forms; (f) depauperate forms, stem under 1 dm tall, spindly, leaves small, long pedicellate and sparse; 63

(g) common forms, branching only above or with 1-2 stems from near the base, leaves tufted, semi-erect, inflorescence loose and pedicels prominently divaricate.

Collections: total examined 354, including 11 collected by author; representative: AR!ZONA, 'Yuma Co, Papago Wells, 26 mi w Tule

Mtns, Bens·on & Darrow 10795, 15 Apr 1941 (DS, POM, ARIZ, UC). CALIFORNIA,

Inyo Co, Darwin, Argus Mtns, Ferris 7930, 11 June 1930 (DS, POM, MO, UC);

Riverside Co, Salton Sea 1 min Imperial Co line, Wiggins 9587, 24 March

1941 (RM, POM, WTU, DS, NY, UC); San Bernardino Co, Needles, Jones

3826, 5 May 1884 (DS, NY, US, POM, UTC, CAS).

Gilia ripleyi Barneby (Fig. 31) • G. ripleyi Barneby, Leafl. West. Bot. 3: 129. 1942.

G. gilmanii Jepson, Fl. Calif. 3: 192. 1943.

Perennial, erect, (0.3) 0.9 - 2.6 (3.2) dm tall, rank odored.

Stems several, arising from woody caudex and robust, corky, twisted root, covered basally with previous years persistent, chartaceous leaves, crisply pubescent with multicellular, translucent, glandular-tipped hairs 0.2 - 2.0 mm long. Leaves alternate, strongly compact to rosetted, erect; lower leaf blades broadly obovate to orbicular in outline (1.0)

1.8 - 5.0 (6.1) cm long, (0.9) 1.2 - 3.6 (4.6) cm wide including spines, ratio blade length to width (0.9) 1.0 - 1.5 (2.3), gradually tapering below to slightly winged, adaxially grooved petiole (0.2) 0.5 - 2.0 (4.4) cm long, leaf margin coarsely serrate-dentate, teeth deltoid, long aristate, spines extensions of lateral veins (3.0) 6.0 - 7.5 (9.0) mm long, infrequently alternating long and short by development of secondary teeth, upper surface of leaf flat, lower surface of leaf coursed by prominently elevated, pinnate vein system, the midrib giving off acutely 64

Fig. 31. Q. ripleyi. Habit drawing is actual size. Isolated fruit and flower are twice actual size. 65 66

3 to 5 pairs of parallel, alternating, thick, straight secondaries, pubescence as on sterns, more dense adaxially, cauline leaves none or limited to 1-4 sessile, aristate-lanceolate (rarely ovate to orbicular, and if so then less reduced), entire or basally toothed leaves, 0.4 -

1.1 cm long, of the peduncle; bracts aristate-lanceolate, entire to basally toothed, 0.5 - 5.0 mm long, abaxial veins prominent. Inflores- cence cymose, open paniculate, never glomerate, (1.8) 4.5 - 10.5 (12.5) cm long, peduncle (2.3) 4.0 - 7.5 (8.3) cm long, ratio inflorescence to peduncle length (0.6) 1.0 - 1.5 (5.5); flowers numerous, commonly two per node borne solitary on moderately to densely stipitate-glandular, divaricate, filiform pedicels (0.2) 0.5 - 1.5 (2.8) cm long, lengths often varied on same plant. Calyx narrowly campanulate (3.2) 3.5 - 5.0

(6.5) mm long, 1.0 - 2.5 mm wide in flower, moderately to densely stipitate-glandular, cleft to base, united in lower half by hyaline membranes; lobes linear (1.4) 1.6 - 2.8 (3.1) mm long, apex long aristate, equal to or exceeding the corolla tube; calyx expanding in fruit by distention of membranes, pro~inently overtopping fruit. Corolla short salverforrn, convolute in bud, (5.6) 7.0 - 9.5 (11.0) mm long; the lobes elliptical to narrowly ovate, conspicuouslyconstricted to short, parallel basal portion, apex rounded acute, margins slightly irregular,

(2.5) 3.5 - 4.5 (5.3) mm long, 1.5 - 2.5 mm wide, ratio lobe to total corolla length (1.9) 2.2 - 2.9 (3.3), lobes bright rose within and without; throat indistinct, white to faintly buff; tube stout, (3.6) 4.0 -

5.5 (5.7) mm long, equaling calyx width, minutely papillate without, white, Stamens included, unequally affixed to lower part of corolla tube, very unequal in length, one longer, 4.2 - 5.0 mm, four shorter,

2.6 - 3.0 mm (rarely one of shorter stamens, intermediate, 3.5 - 3.7 mm); filaments stout, cream or white, minutely papillate; anthers sagittate 67 1.0 - 1.8 mm long; pollen spherical, very small, 25 - 30 microns in

diameter, exine finely granular with four equatorial orbicular pores in

opposing pairs. Ovary ovate-oblong, minutely stipitate-glandular,

~- 2.0 mm long. Style (1.4) 1.8 - 2.5 (2.7) mm long, minutely papillate,

often persistent on capsule; stigma three-lobed (0.3) 0.4 - 0.5 (0.6)

mm long, introrse, reflexing at or slightly below anther level of the

longest stamen, ratio stigma to style length (3.2) 4 - 7.5 (9.0).

Capsule cylindric, (2.2) 3.0 - 5.0 (6.5) mm long, (1.2) 1.8 - 2.5 (4.0)

mm wide, ratio capsule length to width (1.3) 1.5 - 1.9 (2.1); walls

hard, finely reticulate without, smooth and glossy within, dehiscing loc-

ulidically from top to bottom, remaining attached below, slightly re-

curving above; the capsule equai to or commonly greatly exceeded by

aristate calyx lobes. Seeds ovoid, deep reddish-brown, inconspicuously

tuberculate, very small, 0.4 - 0.5 rrnn long axis, 0.3 - 0.4 mm short

axis, 30-100 per locule, producing neither spiracles nor slime when moistened. N = 9.

Holotype: Nevada, Nye County, "in fissures of hard, dry, lime-

stone cliffs in the south end of the Specter Range," Ripley & Barneby

3992, 1941, CAS. Isotypes: POM, NY, K, UC.

Distribution: CALIFORNIA, Inyo County; NEVADA,Clark, Lincoln,

and Nye counties. Talus dust and crevices of calcareous mountain ranges,

3100 to 5000 feet (Fig. 32).

Associated with: Atriplex confertifolia, Buddleia utahensis,

Cryptantha confertiflora, Penstemon petiolatus, Phacelia perityloides, f. geraniifolia, Tetracoccus ilicifolius, Maurandia petrophila, Cymopteris

aboriginum, Eucnide urens, Eriogonum sulcatum, and E. intrafractum.

Flowering time: The earliest occurence of flowering recorded

for G. ripleyi is the 31st of May; the latest, the 1st of October. The 68

Fig. 32. Southern Nevada and adjacent southeastern California. Distribution of G. rirley1. 69

' '

' ••

1-·--- • I •• I .. • 70 plant nonnally blooms from mid-June to mid-August, sets seed rapidly, and dies back until the beginning of a new growth period in late winter.

Discussion: G. ripleyi, unlike Q_. latifolia, is highly unifonn in appearance throughout its range. Although distributed in many of the low, calcareous ranges of southernmost Nevada and adjacent southeastern

California, Q. ripleyi is highly localized in occurrence within a given area. Populations are composed of a few well-developed plants of previous years germinations surrounded by new seedlings. Occupying an isolated ledge or rock crevice, the populations are nowhere very widespread or abundant.

Collections: total examined 51, those collected by author 8, representative: CALIFORNIA, Inyo Co, Emigrant Cyn, Panamint Mtns, Roos

6107, 9 May 1954 (NY, RSA, UC), Johnson Cyn, Panamint Mtns, Gilman 4277,

22 June 1940 (POM, CAS). NEVADA,Clark Co, Spotted Rng, 5 min hywy 95,

Re~eal & Holmgren 1912, 24 Aug 1968 (NTS, NY); Nye Co, Specter Rng, s slope w of Barricade, Beatley & Reveal 1090, 31 May 1968 (NTS, NY, UTC). 71

SUMMARY

The results of this study show that section Gilmania (genus Gilia) consists of only two species .Q_. latifolia and .Q_. ripleyi, which are well separated _from the rest of the members of genus Gilia by their simple, aristately toothed, obovate to orbicular leaf blades, many, small, brilliant rose, salverform corollas, basally inserted, stout-filamented stamens, and 30 to 100 small, reddish-brown, oval seeds per locule.

The members of section Gilmania occur in the arid, western United

States. G. latifolia is a widespread winter annual of southern Californ- ia, western Arizona, southern Nevada and extreme southwestern Utah, and G. 'ripleyi is a narrow endemic perennial of the limestone outcrops of the stark Paleozoic mountain ranges of southern Nevada and adjacent southeastern California. Where their ranges overlap in southern Nevada and adjacent southeastern California, the two species are often found in close proximity, G. latifolia in the Larrea divaricata-Ambrosia dumosa community of the valley floor and deep~y cut washes and .Q_. ripleyi above on the near-sterile limestone outcrops commonly shared by only a few strongly xerophytic herbs and shrubs.

Considering this close morphological and ecological relationship between G. latifolia and G. ripleyi together with the relict perennial nature and basic chromosome number of the latter as contrasted to the more advanced annual habit and tetraploidy of the former, Q. latifolia may well be a direct allo- or autopolyploidic descendent of G. ripleyi. 72

REFERENCES

Abrahms, L. 1951. Illustrated flora of the Pacific States. Vol. III:

Geraniaceae to Schrophulariaceae. Stanford, Calif: Stanford

University Press. 866 p.

Axelrod, D. 1950. Evolution of desert vegetation in western North

America. Carnegie Inst. Wash. Publ. 590: 215-306.

Barneby, R. 1942. A suffrutescent Gilia from southern Nevada. Leaf.

West. Bot. 3: 129-32.

Beatley, J. 1967a. Survival of winter annuals in the northern Mojave

Desert. Ecology 48: 745-50.

1967b. Ecology of the Nevada Test Site. V. Winter annual

survival data. UCLA 12-650. Los Angeles: University of Calif-

ornia School of Medicine. 36 p.

1969. Vascular Plants of the Nevada Test Site, Nellis Air

Force Range, and Ash Meadows. UCLA 12-705. Los Angeles: Uni-

versity of California School of Medicine. 112 p.

Beeks, R. 1955. Improvements in the squash technique for plant chromo-

somes. El Aliso 3: 131-33.

Benson, L. 1957. Plant Classification. Boston, Mass: Heath and Co.

688 p.

Bentham, G. 1845. Polernoniaceae, pp. 302-22. In: Candolle, A. de.

Prodromus systematis naturalis regni vegetabilis. Vol. IX.

Parisiis: Treutel.

Bessey, C. 1915. Phylogenetic taxonomy of flowering plants. Ann. Mo. 73 Bot. Gard. 2: 109-164.

Brand, A. 1907. Polemoniaceae, pp. 1-203. In: Engler, A. Das Pflan-

zenreich. Vol. XX.VII. Berstrabe, Germany: von H. R. Englemann.

Brandegee, T. New species of plants mainly from Baja, California. Zoe

5: ],05-109.

Cronquist, A. 1968. The Evolution and Classification of Flowering

Plants. Boston, Mass: Houghton Mifflin Co. 396 p.

Dawson, M. 1936. The floral morphology of the Polemoniaceae. Amer.

Jour. Bot. 23: 501-511.

Day, A. 1965. The Evolution of a pair of sibling allotetraploid spe-

cies of Cobwebby Gilias (Polemoniaceae). El Ali.so 6: 25-75.

Grant, V. 1949a. Seed germination in Gilia capitata and its relatives.

Madrono 10: 87-93.

1949b. Pollination as an isolating mechanism. Evolution 3:

82-97.

1950. Genetic and taxonomic studies in Gilia. I. Gilia

capitata. El Aliso 2: 239-316.

1952. Genetic and taxonomic studies in Gilia. III. Gilia

tricolor complex. El Aliso 2: 375-88.

1953. The role of hybridization in the evolution of the

Leafy-stemmed Gilias. Evolution 7: 51-64.

1954a. Genetic and taxonomic studies in Gilia. IV. Gilia

achilleaefolia. El Aliso 3: 1-18.

1954b. Genettc and taxonomic studies in Gilia. V. Gilia

clivorum. El Aliso 3: 19-34.

1954c. Genetic and taxonomic studies in Gilia. VI. Inter-

specific relationships in the Leafy-stemmed Gilias. El Aliso 3:

35-49. 74 1956a. The influence of breeding habit on the outcome of nat-

ural hybridization in plants. Amer. Natur. 90: 319-22.

1956b. A synopsis of Ipomopsis. El Aliso 3: 351-62.

1956c. The genetic structure of races and species in Gilia.

Advances Genetics 8: 55-87.

1959. Natural History of the Phlox Family. Vol. I: System-

atic Botany. The Hague: Martinus Nijhoff. 280 p.

1966. The selective origin of incompatibility barriers in the

genus Gilia. Amer. Natur. 100: 99-118.

Grant, V., Beeks, R., and Latimer, H. 1956. Genetic and taxonomic

studies in Gilia. IX. Chromosome numbers in the Cobwebby Gilias.

El Aliso 3: 289-96.

Grant, V. and Grant, A. 1954. Genetic and taxonomic studies in Gilia.

VII. The Woodland Gilias. El Aliso 3: 59-91.

1955. The genus Allophyllum (Polemoni&ceae). El Aliso 3:

93-110.

1956a. Genetic and taxonomic studies in Gilia. X. Conspec-

tus of the subgenus Gilia. El Aliso 3: 297-300.

1956b. Genetic and taxonomic studies in Gilia. VIII. The

Cobwebby Gilias. El Aliso 3: 203-287.

Gra½ A. 1870. Revision of the North American Polemoniaceae. Proc.

Amer. Acad. Sci. 8: 247-282.

1878. Synoptical Flora of North America. 1st ed. Vol. II.

Pt. I. New York: Ivison, Blakeman, Taylor. 494 p.

1886. Synoptical Flora of North America 2nd ed. Vol. II.

Pt. I. New York: Ivison, Blakeman, Taylor. 494 p.

Gray, P. 1964. Handbook of Basic Microtechnique. New York: McGraw-

Hill Book Co. 302 p. 75

Heller, A. 1906. New and interesting plants from western North America.

Muhlenbergia 2: 117.

Hitchcock, C., Cronquist, A., Ownbey, M. and Thompson, J. 1959. Vascu-

lar plants of the Pacific Northwest. Pt. IV: Ericaceae through

Campanulaceae. Seattle, Wash: University of Washington Press.

510 p.

Jepson, W. 1925. A Manual of the flowering plants of California.

Berkely, Calif: University of California Press. 1238 p.

1943. A Flora of California. Vol. III: Polemoniaceae to

Solanaceae. Berkeley, Calif: University of California Press.

402 p.

Johnson, I. 1925. Some undescribed American spermatophytes. Contrib.

Gray Herb. n.s. 75: 39-40.

Jones, M. 1881. Notes from Utah. Bull. Torrey Bot. Club 8: 70-1.

Kearney, T. and Peebles, R. 1951. Arizona Flora. Berkeley, Calif:

University of California Press. 1032 p.

Lanjouw, J. and Stafleu, F. 1964. Index Herbariorum. Pt. I: The Her-

baria of the world. 5th ed. Utrecht, Netherlands: Kemink and

Zoon.

Lundell C. and collaborators. 1966. Flora of Texas. Vol. I. Renner,

Tex: Texas Research Foundation. 407 p.

Macbride, F. 1918. Various American spermatophytes. Contrib. Gray

Herb. n.s. 56: 50-61.

Mason, H. 1945. The genus Eriastrum and the influence of Bentham and

Gray upon the problem of generic confusion in Polemoniaceae.

Madro~o 8: 65-91.

Mason, H. and Grant, A. 1948. Some problems in the genus Gilia. Mad- 76

rorio 9: 201-20.

Munz, P. and Keck, D. 1959. A California Flora. Berkeley: University

of California Press. 1681 p.

Parry, C. 1875. Botanical observations in southern Utah. Amer. Natur.

9: 347-8.

Pohl, R. 1954. A rapid softening agent for dried plant structures.

Proc. Iowa Acad. Sci. 61: 149-50.

Raisz, Erw.in. 195 7. Landforms of the United States. 6th ed. No place:

by author.

Rydberg, P. 1922. Flora of the Rocky Mountain States and adjacent plains.

2nd ed. Lancaster, Penn: Intelligencer Printing Co. 1143 p.

Shinners, L. 1963. Gilia and Ipomopsis (Polemoniaceae) in Texas. Sida

1: 171-9.

Shreve, F. and Wiggins, I. 1964. Vegetation and Flora of the Sonoran

Desert. Vol. II: Flora of the Sonoran Desert. Stanford, Calif:

Stanford University Press. pp. 841-1740.

Sinclair, C. and Dunn, D. 1961. Surface printing of plant leaves for

phylogenetic studies. Stain Tech. 36: 299-304.

Small, J. 1933. Manual of the Southeastern Flora. Lancaster, Penn:

The Science Press. 1554 p.

Standley, P. 1913. Trees and shrubs of Mexico (Passifloraceae to Scroph-

ulariaceae). Contrib. U. S. Natl. Herb. 23: 849-1312.

Tidestrom, I. and Kittel, T. 1941. A Flora of Arizona and New Mexico.

Ann Arbor, Mich: Edwards Brothers Inc. 879 p.

Welsh, S., Treshow, M. and Moore, G. 1965. Common Utah Plants. Provo,

Utah: Brigham Young University Press. 312 p.

Went, F. and Lewis, H. 1945. Plant growth under controlled conditions. 77

IV. Response of California annuals to photoperiod and temperature.

Amer. Jour. Bot. 32: 1-12.

Went, F. and Westergaard, M. 1949. Ecology of desert plants III. De-

velopment of plants in the Death Valley National Monument, Calif-

ornia. Ecology 30: 26-38.

Wherry, E. 1940. A provisional key to the Polemoniaceae. Bartonia 20:

14-17.

1959. Contributions toward a flora of Nevada. No. 43:

Polemoniaceae. Beltsville, Maryland: United States Plant

Industry Station. 103 p. 78

APPENDIXA

Specimen Citations G, latifolia

The following list is arranged alphabetically according to state and county. The citation is given in the following order: locality, collector(s), field ntnnber, date, herbaria. Abbreviations used within the citations are: Apr (April), Aug (August), Feb (February), Crk

(Creek), Cyn (Canyon), Dsrt (Desert), e (east), hywy (highway), Jan

(January), Mtns (Mountains), n (north), Oct (October), Pk (Peak), rd

(road), Rng (Range), s (south), Sept (September), Sprgs (Springs), w

(west).

Arizona

MOJAVECOUNTY: Hoover Dam, Ripley & Barneby 2902, 9 Apr 1940 (NY),

Alexander 65, 10 May 1933 (UC), Beck sn, 27 Apr 1935 (DIX), Lake Mead,

Grapevine Wash, Kellogg 12, 4 Apr 1942 (US), Beaver Dam, Palmer 322,

1877 (NY, US), Kearney & Peebles 13224, 17 Apr 1937 (ARIZ); YUMACOUNTY:

Mohawk, Kearney 3925, 1 Apr 1927 (US, ARIZ), Harrison 3568, 1 March 1927

(ARIZ), Harrison & Kearney 5015, 8 March 1928 (ARIZ), McLellan & Stilt sn, 6 Apr 1941 (ASU), Yuma, Jones sn, 25 Apr 1906 (POM), Beard sn, 1911

(MO), Brown sn, Apr - May 1901 (ARIZ), Brown sn, 16 Apr 1905 (ARIZ),

Yuma to Quartsite rd, wash s Gila R, Mason 14216, 8 Apr 1952 (RSA, UC),

Papago well, 25 mi w Tule Mtns, Benson & Darrow 10795, 15 Apr 1941 (DS,

POM, UC, ARIZ), Tule Wells, 5 mi e Tule Mtns, Darrow sn, 15 Apr 1941

(UC), Dome, Peebles & Harrison 5046, 10 March 1928 (US, ARIZ), Wellton,

7 mis, Munz 16616, 30 March 1941 (WTU, POM). 79

California

KERNCOUNTY: Red Rock Cyn, Thorne 31791, 17 May 1963 (RSA), Iron Cyn,

El Paso Rng, Twisselmann 11892, 12 Apr 1966 (RSA); IMPERIALCOUNTY:

Split Mtn, Fish Creek Rng, Ferris 969613~ 29 March 1939 (DS, US), Brande- gee sn, Apr 1905 (UC), Painted Gorge, Wiggins 14048A, 24 March 1957 (DS),

Anderson 267, 18 Feb 1964 (CAS), Picacho Pk, 3 mi from Imperial Co line,

Ferris sn~ 20 Apr 1928 (DS), Salton Sea, Lehto 5683, 11 March 1965 (ASU),

Truckhaven, Wolk 8444, 6 Apr 1937 (RSA), Imperial Gables, 3 mi w, Choc- olate Mtns, Rutherford & Thorne 33605, 11 Apr 1964 (RSA), Glamis, 6 mi e,

Alexander & Kellogg 1930, March 1941 (GH, UC), Ogilby, 25 mine, Munz &

Hitchcock 12182, 6 Apr 1932 (MO, UC), Araz Wash, Cargo Muchacho Mtns,

Thorne & Rutherford 33620, 11 Apr 1964 (RSA, UC), Signal Mtn, Abrams 3165,

2 Apr 1903 (DS, NY, PO:M), Bard, Goodding 13-44, 23 Feb 1944 (NY), Cyn 2 mis hywy 98, Wiggins 14086, 25 Marc.h 1957 (WTU, DS); INYO COUNTY:

Darwin, Hitchcock 6218, 28 Apr 1940 (WTU, NY), Ferris 7930, 11 June 1930

(DS, POM, MO, UC), MacFadden 13295, 15 Apr 1935 (CAS, NY), Stove Pipe

Wells, MacFadden 13583, 15 Apr 1935 (CAS), Everett & Balls 23216 (UC,

RSA), Surprise Cyn, Panamint Mtns, Howell 3976, 16 June 1928 (CAS),

Panamint Mtns ne of Trona, Alexander &"Kellogg 1140, 18 Apr 1940 (POM),

Panamint Cyn, Jones sn, 3 May 1897 (US, POM), Panamint Sprgs, East- wood & Howell 7655, 26 March 1940 (CAS), Crum 2071, 1 June 1938 (MO),

Panamint Valley, Welsh, Matthews & Atwood 9647, 12 Apr 1970 (BRY), Twis- selmann 11947, 13 Apr 1966 (POM), Mathias 831, 18 Apr 1931 (MO), Train

523, 7 March 1937 (UC), Coville & Funston 672, 17 Apr 1891 (NY, GH, US,

MO), Jones sn, 1 May 1897 (POM), Clokey & Templeton 5707, 4 Apr 1935 (UC),

Townes Pass Cyn, Miller sn, 4 May 1963 (OSU, POM), Bacigalupi 4832, 7

Apr 1955 (JEPS), Welsh, et. al. 9644, 12 Apr 1970 (BRY), Owens Valley, 80

Rixford sn, 1890 (CAS), Keller, Coville & Funston 862, 15 May 1891

(DS, US, DAS), Brandegee sn, 4 Apr 1891 (UC, CAS), Thorne & Everett 33793, 18 Apr 1964 (POM), Kerr sn, 2 July 1939 (CAS), Funeral Mtns, Covil-

le & Funston 456, 21-22 March 1891 (US), Jones sn, 8 Apr 1907 (POM),

Jones sn, 9 Apr 1907 (POM); Furnace Crk~ Peebles 266, 11 June 1930

(ARIZ), Rose 40060, 9 March 1940 (MO), Eastwood & Howell 7749, 27 March

1940 (POM, CAS), Grant & Grant 17538, 4 Apr 1952 (NY, POM), Coville &

Gilman 76, 17 Apr 1931 (US), Coville & Funston 570, 8 Apr 1891 (US),

Pinkava, Batchelder, Lewis, Valasco, Keil, Lehto 12232, 12 Apr 1968 (ASU),

Furnace Crk, 20 min, Welsh et. al 9606, 11 Apr 1970 (BRY), Emigrant

Sprgs, Parish 10043, 14 May 1915 (GH, DS, UC), Gould 968, 24 March 1940

{DS, NY, MO, UC), Reveal & Holmgren 1796, 15 Aug 1968 (NTS), Golden Cyn,

Pinkava et. al. 12733, 12 Apr 1968 (ASU), Coral Reef Cyn, Clary 960,

12 Apr 1932 (JEPS), Texas Sprgs, Harrison 5440, 20 May 1932 (MO), Grotto

Cyn1 Epling, Robbon & Haines sn, 20 Apr 1935 (DS, MO, UC), Warm ::>prings,

Parish & Parish 253 (locale inconsistent), May 1882 (DS, US), Shoshone,

s of, Welsh et. al. 9559, 10 Apr 1970 (BRY), Pinkava et. al. 12547, 11

Apr 1968 (ASU), Mosaic Cyn, Templeton & Clokey 5752, 6 Apr 1935 (WTU,

BY, UC), Scottie's Castle, Gould 949, 23 March 1940 (ARIZ, DIX),

Ubehebe Crater, Pinkava et. al. 12130, 13 Apr 1968 (ASU), Devil's Golf

Course, Pinkava et. al. 12731A, 12 Apr 1968 (ASU), Bad Water, Welsh et. al.

9604, 11 Apr 1970 (BRY), Bailey sn, 23 March 1940 (JEPS), Death Valley

Esmond sn, Spring 1935 (CAS), Jones sn, 12 March 1924 (POM), Bradbury

Well, Munz 16453, 9 Apr 1940 (POM), Tule Sprgs, Nopah Mtns, Wolf 10582,

15 May 1941 (NY RSA), Titus Cyn, Kirby 1077, March 1958 (CAS), Raven 12093,

31 March 1958 (CAS), Welsh et. al. 9631, 9637, 11 Apr 1970 (BRY), Ramsey

1822, 24 Feb 1940 (POM), Grant 1014A, 23 Apr 1958 (POM), White Mtns,

Munz 12719, 6 Oct 1948 (NY, POM, CAS), Brandegee sn, July 1912, (GH, DS, 81

US, UC), Genson 5947, 28 Apr 1934 (POM), Mitchell & Lloyd 1763, 13 July

1963 (NY, ARIZ, UC), Deep Sprgs Lake Raven 6978, 14 June 1954 (CAS, NY),

Saline Valley, Kellogg & Alexander 2678, 21 Apr 1942 (DS, US, UC),

Brandegee sn, Apr 1901 (UC), Munz 18032, 8 Apr 1967 (RSA, ASU), Reveal &

Reveal 379, 5 May 1963 (NY, BRY, CAS, Ute:), Kerr sn. 8 May 1935 (CAS),

Jubilee Pass, Black Mtns, Train sn, 7 March 1937 (OSU, DS, ARIZ),

Pinkava et. al. 12633, 11 Apr 1968 (ASU), Welsh et. al. 9568, 11 Apr 1970

(BRY), Carpenter sn, 1 Apr 1940 (JEPS), Black Cyn, Duran 2737, 8 June

1930 (UC), Eureka Valley, Kellogg & Alexander 5659, 25 May 1949 (UC),

Roos 6334, 13 May 1955 (POM, UC), Inyo Mtns, Kerr sn, 2 July 1939 (POM,

CAS), Kerr sn, 5 May 1937 (CAS), Inyo, Rixford sn, 1890 (UC); RIVERSIDE

COUNTY: Mecca, Davidson 3365, Apr 1920 (US), Spencer sn, Apr 1917 (GR),

Spender 1571, 24 March 1922 (GH, US), Hiland sn, 2 Apr 1927 (US, POM),

Munz & Keck 4764, 9 Apr 1922 (POM), Grant & Grant 17519, 1 Apr 1952 (CAS),

Wolf 3003, 27 Apr 1932 (RSA), Painted CyLL, 3tason sn, 12 i•iarch 1927 (UC),

Peirson 7168, 12 Apr 1927 (RSA), Kelly sn, Apr 1926 (CAS), Ray 1385A, 12

March 1949 (JEPS), Bacigalupi sn, April-May (DS), Jaeger 1129, Apr 1921

(POM), McGregor 788, 18 March 1917 (DS), Ballou sn, 27 Feb 1926 (DS),

Indio, Jones sn, 27 Apr 1906 (US, DS), Duran sn, 18 Apr 1935 (UC), Braun- ton 2019, 2045, 3 Apr 1940 (RSA), Rose sn, 10 Apr 1937 (CAS), Dos Palmos

Sprgs Hall 5850, Apr 1905 (UC), Orcutt sn, 26 June 1888 (US), Munz 9967,

31 Jan 1926 (POM), Peirson 6498, 31 Jan 1926 (RSA, UC), Palm Sprgs,

Parish sn, 4-13 Apr 1896 (NY), Peirson 2385, 22 March 1921 (RSA),

Coachella Mtns, Mason 4203, 10 March 1928 (UC), Greata sn, April 1905

(UC), Whitewater, Munz & Keck 4972, 14 Apr 1922 (POM), Parish & Parish

253 (inconsistent locale), Apr 1880 (GH, NY), Chocolate Mtns, Winblad sn, 22 March 1937 (CAS), Santa Rosa Mtns, Perison 2871, 11 Apr 1922 (RSA),

Milpitas Wash, Hitchcock 24311, 20 March 1966 (WTU), Salton Sea, Wiggins 82 9587, 24 March 1941 (RM, P~~, WTU, DS, NY, UC), Colorado Dsrt, Orcutt sn, 24 Apr 1890 (US); SAN BERNARDINOCOUNTY: Shoshone, 20 mis of, Welsh et. al. 9563, 9564, 10 Apr 1970 (BRY), Saratoga Sprgs, Hitchcock 24351,

22 March 1966 (WTU), Hitchcock & Muhlick 23302, 24 March 1964 (WTU, DS,

NY), Valley Sprgs, Wolf 6618, 30 Apr 1935 (RSA), Searles Lake, Mason

8227, 8287, 29 March 1935 (DS, UC), Mason 8288, 30 Apr 1935 (DS, UC),

Ferris 3905, 17-24 March 1924 (DS), Kelso, Wolf 10845 (NY, UC, RSA),

Jaeger 1086, Apr 1921 (POM), Jones sn, 2 May 1906 (POM), Alva & Baci- galupi 1923, 4 Apr 1958 (JEPS), Barstow, Brandegee, June 1884 (UC),

Curran, June 1884 (CAS, UC), Bagdad, Hall 6083, May 1905 (UC), Calico,

Lemmon sn, May 1884 (UC), Calico Mtns, Shreve sn, 24 Apr 1915 (ARIZ),

Pisgah Crater, Robbins 3470, 27 Apr 1952 (ARIZ, JEPS), Trona, 11 mis,

Hitchcock 12275, 15 Apr 1932 (POM, MO UC), Lone Willow Sprgs, Parish

10076, 9 May 1915 (DS), Needles, Jones 3826, 5 May 1884 (DS, NY, US,

POM, UTC, CAS), Denning Sprgs, Wolf 6608, 30 Apr i935 (RSA), Sait Sprgs,

Parish 10061, 21 May 1915 (DS), Copper Basin Lake, Whipple Mtns,

Alexander 708, 6 May 1939 (UC), Baker, 34 min of, Howell 3596, 1 Apr

1928 (CAS), San Bernardino, Parish sn, 1880 (MO); SAN DIEGO COUNTY:

Borrego Sprgs, Jones sn, 19 Apr 1906 (GH, DS, POM), Brandegee sn, 17

Apr 1895 (UC, ASU), San Felipe Crk, Eastwood 2715, 14 Apr 1913 (GH, US,

UC CAS), Vallecito, 13 mi se, Munz 15812, 28 Jan 1940 (POM), Del Mar,

Nelson 1183A, 10 March 1930 (RM, UC), Palm Wash, Howell 3508, 24 March

1928 (UC, US, NY, JEPS, CAS), Pilot Knob, Carlson sn, 21 Apr 1905 (CAS):

VENTURACOUNTY: El Rio, Leamon 171, May 1881 (UC).

Nevada

CLARKCOUNTY: Spotted Rng, n-central, Beatley sn, 18 May 1969 (NTS),

Beatley sn, 27 Apr 1969 (NTS), Spotted Rng, s, Beatley sn, 14 Apr 1969 83

(NTS), Las Vegas Wash, Goodding 2262, 26 Apr, 1905 (GR, RM, MO), Las

Vegas, 18 min of, Train 1801, 22 May 1938 (UC), Indian Sprgs, Clokey

8211, 8056, 23 May 1938 (NY, UC), Glendale to St. Thomas rd, Maguire &

Blood 1498, 29 Apr 1932 (GH, UC, UTC): ESMERALDACOUNTY: Fish Lake

Valley, Archer 7258, 26 Sept 1938 (ARI?., UC): NYE COUNTY: Specter Rng, central, Beatley sn, 21 Apr 1968 (NTS), Specter Rng se, Beatley sn, 20

Apr 1968 (NTS), Specter Rng e, Beatley sn, 16 June 1969 (NTS), Specter

Rng, w, Beatley sn, 17 Apr 1969 (NTS), Timpahute Rng, Bare Mtn, Beatley sn, 19 June 1969 (NTS), Ash Meadows, Beatley sn, 17 June 1969 (NTS),

Reveal & Matthews 2154, 3 June 1969 (BRY), Amargosa Narrows, Beatley sn,

29 Apr 1969 (NTS), Amargosa Dsrt, Jones s~ 27 Apr 1907 (POM), Lathrop wells, Cronquist 10656, 21 Apr 1966 (WTU, BRY, UTC), Thirsty Cyn, Reveal

7 Beatley 1709, 28 July 1968 (NTS): WASHOECOUNTY: Reno, Shore 53,

Summer 1939 (NY).

Utah

WASHINGTONCOUNTY: Valley of the Virgin, near St. George, Parry 188,

1874 (NY, MO, GH), Price Bench, Hardy sn, 27 Apr 1941 (DIX), s Utah,

Jones sn, n.d. (POM). 84

APPENDIXB

Specimen Citations G. ripleyi

California

INYO COUNTY: Titus Cyn, Roos & Roos 6105, 8 May 1954 (RSA, CAS), Gilman

4262, 4263, 21 June 1940 (POM, JEPS), Gilman sn, Apr 1940 (JEPS), Ripley

& Barneby 2951, 18 Apr 1940 (CAS), Johnson Cyn, Gilman 4277, 25 June 19-

40 (POM, CAS), Jepson 19789, 25 Apr 1925 (JEPS), Emigrant Cyn Roos &

Roos 6107, 9 May 1954 (NY, RSA, UC).

Nevada

CLARKCOUNTY: Spotted Rng, Matthews & Reveal 128, 10 June 1970 (BRY),

Reveal & Holmgren 1912, 24 Aug 1968 (NTS, Ni), Pintwater Rng, Kipiey &

Barneby sn, 12 Apr 1940 (CAS), Desert Rng, Ripley & Barneby sn, 10 Apr

1940 (CAS); LINCOLNCOUNTY: Desert Rng, Ripley & Barneby sn (CAS); NYE

COUNTY: Ranger Mtns, Bostick sn, 24 Apr 1967 (NTS), Tungsten Cyn, Timp- ahute Rng, Beatley sn, 19 Apr 1969 (NTS), Gold Ace Mine area, Timpahute

Rng, Beatley sn, 2 Oct 1969 (NTS), Beatley sn, 19 June 1969 (NTS), Red

Mtn, Spotted Rng, Reveal & Matthews 2159, 4 June 1969, Matthews & Reveal

126, 10 June 1970 (BRY, NTS), Spotted Rng, sw end, Matthews & Reveal

127, 10 June 1970 (BRY), Beatley sn, 18 Aug 1968 (NTS, NY), Ripley &

Barneby 3420, 14 May 1941 (CAS), Specter Rng, Ripley & Barneby 2873,

5 Apr 1940 (RSA), Beatley sn, Oct 1968 (NTS, BRY), Beatley sn, 24 June

1968 (N'I'S, NY), Ripley & Barneby 3992, 18 July 1941 (NY, POM, UC, CAS,

K), Beatley, Hill, Kaaz sn, 6 May 1968 (NTS, NY), Beatley & Reveal 1090,

31 May 1968 (NTS, NY UTC).