PHYTOLOGIA

_An international journal to expedite botanical and phytoecological publication

Vol. 66 | May 1989 No. 3

CONTENTS

/REVEAL, J.L., Notes on selected genera related to Chorizanthe
{ (Polygonaceae: Eriogonoideae) 0000000000000 0000080S0OSSOSSOOSOSOOOESOSOOOEOOO0E9 199

J REVEAL, J.L., A review of the genus Harfordia (Polygonaceae:

E riogonoideae) SOOOCOOOOO STOO OOOO OOO OOS OOOOSOS ODES OOS OOOOH OOO SOS OSOOESOOSOSOSESOESSOOES 2?1

, REVEAL, J.L., Remarks on the genus Pterostegia (Polygonaceae:

Eriogonoideae) OOOO OOOOOOOOOE SOOO OOOO OOSSOSOOOSEOO OOO OOOOOOSSOOOOSOSOSOOOOOOOSOOSOOOOS 228

“REVEAL, J.L., Notes on selected genera related to Eriogonum
j (Polygonaceae: Eriogonoideae) OOCOCOOSSOOOOO OOOO SSOOOOOOOTOOSOSSOOSOOOOOCOSOOSOS 236

REVEAL, J.L., A new variety ‘of Eriogonum nudum (Polygonaceae:
Eriogonoideae) from the southern Sierra Nevada escsceccccccecssceseseves 246

BEVEAL,. Jul: and: J/R; SHEVOCK, A new variety of Eriogonum
_ _prattenianum (Polygonaceae: Eriogonoideae) from the southern Sierra

N evad a SOSSHOSSSSOSSSOSSHSSOSSSSSSSSSSSOSSSHSSSHHSSSSHOSSSHSSSSSSSSSOSSSSSSSSSSSSSSSOSEOESOSS 249

q REVEAL, J.L., New combinations and novelties in Eriogonum
(Polygonaceae: Eriogonoideae) COCCCCOO OOOO SOOOOOOOO OOO OOS SOSOOOEOOOOOESOOSOOESES 251

/ REVEAL, J.L., A checklist of the Eriogonoideae (Polygonaceae) seseeer+ 266

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Phytologia (May 1989) 66(3):199-220

NOTES ON SELECTED GENERA RELATED TO CHORIZANTHE
(POLYGONACEAE: ERIOGONOIDEAE)

JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

ABSTRACT

In a revision of Chorizanthe prepared by Reveal and Hardham (1989b) several species were
excluded. Three species were referred to new genera, Aristocapsa, Dodecahema and Systenotheca,
all removed from the previously excluded genus Censrostegia which itself is now considered to
be monospecific. The genus Mucronea with it two species and Lastarriaea with its three, along
with the monospecific genus Hollisteria, have been maintained as distinct from Chorizanthe. Four
of these excluded genera, Muconea, Censrostegia, Hollisteria and Lastarriaea are reviewed with
the genera and its species characterized and representative specimens listed documenting their
known distribution. Lastarriaea ptilota, endemic to the Desierto el Vizcaino and the Cedros
Island region of Baja California, México, is described as new. All of these genera are placed in
the subtribe Hollisteriineae, which is defined to include those genera of the Eriogoneae having

One to six flowers in an awned narrowly tubular involucre or subtended by bractlike involucral
lobes.

KEY WORDS: Polygonaceae, taxonomy, Mucronea, Centrostegia, Hollisteria, Lastarriaea, Cali-
fornia, México, Chile.

INTRODUCTION

In a recent review of the annual species of Chorizanthe (Reveal & Hardham 1989b), several
species were excluded and referred to other genera. Three species generally included in
Chorizanthe or Centrostegia were each placed in a monospecific genus: Aristocapsa,
Dodecahema and Systenotheca (Reveal & Hardham 1989a). Four other species were removed
from Chorizanthe as defined by some authors (Torrey & Gray 1870; Watson 1877; Jepson
1913, 1925; Abrams 1944; Munz 1959, 1974). These include Chorizanthe californica and C.
perfoliata which we, like Goodman (1934) the last monographer of Chorizanthe, prefer to
keep in Mucronea as defined by Bentham (1836, 1856) and followed by Hoover (1966). In
addition, two other species often placed in Chorizanthe (Parry 1884, 1885; Goodman 1934;
Munz 1959, 1974), C. coriacea of North America and the superfluous C. lastarriaea of South
America, are returned to Lastarriaea, a genus that has long been considered to be distinct
(Rémy 1851-1852; Torrey & Gray 1870; Watson 1877; Parry 1886; Jepson 1913, 1925; Abrams
1944; Hoover 1966; Wiggins 1980). Significantly, a new species is added to Lastarnaea, a
central Baja California endemic currently known only from two collections. Two additional
monospecific genera are also recognized, Hollisteria and Centrostegia. With the removal of
three species previously placed in Centrostegia as noted above, Centrostegia now becomes
monospecific but unlike the other segregates in North America whose species are essentially
restricted to California and Baja California, Centrostegia thurberi is distributed over a wide

199

200 PHYTOLOGIA volume 66(3):199-220 May 1989

area of the American Southwest. The genus Hollisteria has uniformly been regarded as dis-
tinct from Chorizanthe or Eriogonum except indirectly by Jones (1908) who proposed a new
species of Chorizanthe that proved to be based on specimens of H. lanata., and by Roberty
and Vautier (1964) who, in their bizarre review of the Polygonaceae, transferred H. lanata
to Eriogonum. Shields and Reveal (1988) excluded Hollisteria from the Chorizanthe complex
retaining it with Eriogonum where it had been traditionally placed (Curran 1885; Dammer
1892; Gross 1913a). That view is now rejected and Hollisteria is placed in the Chonzanthe
complex in an isolated position.

The purpose of this paper is to present a series of descriptions and discussions of the four
already established genera that are related to but distinct from Chorizanthe.

TAXONOMY

The subf. Eriogonoideae was established by Meisner (1856) in the introduction to the
Polygonaceae published in de Candolle’s Prodromus. The name was used by numerous au-
thors up to Melchior (1964) although, for some unknown reason, the taxon was reproposed
by Roberty and Vautier (1964). In all instances, the taxon was circumscribed to include those
genera of Polygonaceae that lacked a well defined ochrea. Dumort (1829) proposed the tribe
Eriogoneae, placing it in the Chenopodiaceae; Bentham (1836) would later transfer the taxon
to the Polygonaceae where it has remained essentially unchanged. Walpers (1852) elevated
the group to the family rank, but the name has rarely been used. Torrey and Gray (1870)
renamed the tribe Eueriogoneae. According to the /nternational Code of Botanical Nomencla-

ture (Greuter 1988), this last name is not only superfluous but such names are not permitted
(Art. 21.3).

Within the subfamily, Torrey and Gray (1870) distinguished two tribes, Eriogoneae and
Pterostegieae, and both are still recognized. The latter is restricted to Pterostegia and Har-
fordia (Reveal 1989a, b). Dammer (1892) had included these two genera, along with Lastar-
niaea, Nemacaulis and Hollisteria, in the subtribe Koenigiinae. Its type (Art. 22.4; Greuter
1988) is the genus Koenigia L., a taxon that is not now regarded to be a member of the
subfamily (Roberty & Vautier 1964; Melchior 1964).

In his review of the Polygonaceae, Gross (1913a) recognized the subf. Eriogonoideae and
divided it into two tribes, the Eriogoneae and the Hollisterieae. The latter was itself divided
into two subtribes, Hollisteriinae and Harfordiinae. The latter consisted of Pterostegia and
Harfordia, while the former included not only Hollisteria but Gilmania (as Phyllogonum) and
Nemacaulis along with Lastarriaea. Gross defined the Eriogoneae to include Enogonum,
Pterogonum (a new genus he proposed), Oxytheca, Centrostegia and Chorizanthe.

Reveal and Howell (1976) recognized 13 genera in Eriogonoideae. With the publication
of Goodmania (Reveal & Ertter 1977) the number increased to 14. In Shields and Reveal
(1988), the general phylogenetic scheme called for both Hollisteria and Nemacaulis to be
associated with Eriogonum, with the placement of Hollisteria the most problematic. Now it
is proposed that Hollisteria be placed in the Chorizanthe complex.

As here defined, the tribe Eriogoneae is divided into two subtribes, Eriogonineae (Roberty
& Vautier 1964) and the Hollisteriineae (Gross 1913a). The two taxa differ mainly in the
number of flowers per involucre (mostly numerous in Eriogonineae versus 1-6 in the Hol-
listeriineae) but the two may also be differentiated by the general shape and construction of
their involucres and by the presence or absence of uncinate or short involucral awns. A review
of genera belonging to the Eriogonineae is presented elsewhere (Reveal 1989c).

Reveal: Genera related to Chorizanthe (Polygonaceae) 201

Polygonaceae Juss. subtribe Hollisteriineae H. Gross, Bot. Jahrb. Syst. 49: 329. 1913.
Polygonaceae tribe Hollisterieae H. Gross, Bot. Jahrb. Syst. 49: 239. 1913.— TYPE: Hol-
listeria §. Wats. according to Art. 22.4 of the ICBN.

Polygonaceae chort Chorizanthastreae Roberty & Vautier, Boissiera 10: 84. 1964.— TYPE:
Chorizanthe R. Br. ex Benth. according to Art. 22.4 of the ICBN.

Plants annual or perennial herbs to subshrubs; /eaves basal or cauline, mostly narrow,
thinly to densely pubescent; inflorescences mostly cymose with the secondary branches typi-
cally suppressed; peduncles typically lacking or short and rather stout; involucres narrow and
cylindrical or if broader then the tube cylindric and the teeth broad, divergent and thickish,
the teeth typically armed with short, straight or more often uncinate awns, occasionally the
involucre reduced to a series of awned involucral bracts; flowers 1-6 per involucre, usually
short pedicellate, the tepals mostly narrow and thinly pubescent at least along the midrib;
achenes narrow or globose with a straight or curved embryo; n= mostly 14, 17 or 19-24, 40.

Widespread in western North America from eastern Washington and Idaho south through California, Nevada
and southern Utah to Arizona and northwestern México, and in southwestern South America in the arid regions
of extreme southern Perd south to central Chile, with the greatest concentration in coastal and arid regions of
California and northern Baja California.

The use of the name Hollisteriineae for the subtribe rather than one based on (and therefore
typified by) Chorizanthe is unfortunate especially given the controversal positioning of Hol-
listeria within the Chorizanthe complex. Gross (1913a) initially failed to describe the subtribe
(p. 239), but in his concluding remarks (p. 329) he provided a key to the higher taxa in
Polygonaceae and there effectively published the name.

As here defined, the subtribe Hollisteriineae is composed of eight genera, the largest being
Chorizanthe with some 50 species of annuals and perennials. This genus is the most widely
distributed of the taxon being found throughout the range of the subtribe in North and South
America (Reveal & Hardham 1989b). The majority of genera are endemic to California:
Aristocapsa, Dodecahema, Systenotheca, Mucronea and Hollisteria. Centrostegia is found in
southwestern North America from California east to southern Utah south into north- western
México. The genus Lastarriaea is encountered along the coast and in the more arid inland
regions of central and southern California southward to central Baja California, México, and
then like Chorizanthe, it reappears again in the deserts of northern and central Chile. Except
for Mucronea and Lastarriaea, all of these smaller genera are monospecific.

The vast majority of the species assigned to the subtribe are annuals; the ten or so Chilean
perennial species of Chorizanthe being the only exception. The flowers are generally solitary
in the involucre or at a node, but some may have two or three, with only one genus, Ars-
tocapsa, having six flowers in the involucre. Flowers are in well defined involucres in all
genera except Hollisteria and Lastarriaea. In these genera the involucre is reduced to a whorl
of awned involucral bracts subtending the flower. The general shape of the involucre is long
and slender with a generally cylindric shape. The involucral teeth are mostly short with short,
straight or uncinate awns, but in several species the teeth are longer and stout with the
divergent teeth thickened basally so that the teeth are often more pronounced than the tube.
In such instances not all of the teeth may be awned for the smaller alternating three may
be obscure and awnless. In a few instances, the number of lobes is truely three and in
Mucronea californica it can be reduced to two. The majority of the involucral tubes are
distinctly angled with six teeth. Major exceptions are the involucres of Arstocapsa insigne

202 PHYTOL OGIA volume 66(3):199-220 May 1989

and Chonzanthe spinosa where five involucral teeth and a smooth surface more typical of
those scen in the Eriogonineae are encountered.

The flowers of Lastarriaea are coriaceous with each tepal bearing a small, terminal, un-
cinate awn similar to that found on involucres elsewhere in the subtribe. It and Hollisteria
are the most advanced genera of the subtribe, but neither is all that closely related to the
other. In Centrostegia, the five involucral lobes are fused so as to create a highly modified,
3-angled, prismatic tube.

Key to the Genera

A. Involucres distinctly tubular and well defined at least basally.

B. Bracts opposite or whorled, not 3-lobed or parted; flowers 1 (2) in the involucre; plants annual or
perennial; widespread in western North America and southwestern South America. ...... 1. Chorizanthe

BB. Bracts alternate, 3-lobed or parted; flowers (1) 2-6 in the involucre; strictly annual.

C. Involucres 24- or 6-lobed or -toothed; flowers 1-2 per involucre.
D. _Involucres not awned basally, California.

E. Flowers pubescent, perfect; bracts conspicuous; involucral teeth unequal; Coast Ranges
of central and southern California eastward onto the western edge of the Mojave Desert.

Beas cidetevonecbonespsaupsuatiaaeededswadens seins stuatievahtuvuceessateiecedecscshsbsuaastesett eee REE 2. Mucronea
EE. Flowers glabrous but densely papillate, polygamo-dioecious with the upper one perfect
and the lower one female; bracts small; involucral teeth equal; Santa Lucia Mountains
of Monterey and San Luis Obispo COS. .......-.-.csecsecsecsessesessesnsssssesnsensesnenneeeee 3. Systenotheca

DD. Involucres awned basally, southwestern North America.
E. Basal awns 3, curved on saccate basal lobes; terminal awns 5, straight; involucres strongly
prismatic and 3-angled; flowers 2; California east to Utah and south to northwestern

MEXICO (i hasteacaossscptescsssesacedusdapvatossonscdectcsestravic tues cocecezelesncane ivoutetee redeem 4. Centrostegia
EE. Basal awns 6, uncinate on smooth basal lobes; terminal awns 6, uncinate; involucres

strongly 6-angled; flowers 3; southwestern California. ...........:cceseeeceeeee 5. Dodecahema
CC. Involucres 5-toothed; flowers 4-6 per involucre, pubescent; Coast Ranges of Monterey and San Luis

OTE ESS. ccrcereecosreecr er cor ere oreroercer oreo rrcee thor rorec oo rine barre occ ccbeerherecostrcersnthonnctern coer 6. Aristocapsa
AA. Involucres reduced to a series of 3 (4) involucral bracts.

B. Flowers bright yellow and densely tomentose with mucronate petaloid tepals; stamens 6-9; branches
tomentose; central! California... <.: 22:22. ssscacss2easc-cecscessenenscecessnsussaasicsasisuessneveseudcoeoouea str ttere eee EEE 7. Hollisteria

BB. Flowers light green to greenish-white, thinly pubescent with distinctly coriaceous awned tepals; stamens
3; branches glabrous or sparsely pubescent; widespread in central and southern California south to central
Baja California; disjunct*to: Chiles. occ svvccssc-cscececcersscenecstasesseccossascoeccsnds<ceuacivceset sce tte eee 8. Lastarriaea

Three of the above genera were described as new in this volume of Phytologia (Reveal &
Hardham 1989a) and a revision of Chorizanthe is also presented Reveal & Hardham 1989b).
Accordingly those genera are not treated again and the following remarks are restricted to
Mucronea, Centrostegia, Hollisteria and Lastarriaea.

2. Mucronea Benth., Trans. Linn. Soc. London 17: 419. 1836. Chorizanthe R. Br. ex Benth.

sect. Mucronea (Benth.) A. Gray, Proc. Boston Soc. Nat. Hist. 7: 148. 1859.—TYPE:
Mucronea californica Benth.

Erect to spreading, sparsely glandular-pubescent annual herbs arising from a thin taproot;
leaves basal, spatulate to obovate, sparsely hirsute or glabrous except for a ciliated margin
and midvein, tapering to a slightly winged, ciliated or sparsely hirsute, sometimes indistinct
petiole; flowering stems erect, slender to stoutish; branches dichotomously branched through-
out; inflorescences cymose with the secondaries suppressed; bracts 3 (5), triangular to ovate
or oblong, acute to obtuse, spreading to nearly erect, awn-tipped, hispid, hirsute or glandular

Reveal: Genera related to Chorizanthe (Polygonaceae) 203

as well as ciliated along the margin, united nearly throughout and positioned to one side of
the node or perfoliate and completely but unequally surrounding the node; peduncles lacking;
involucres 1-3 at each node, cylindric, glandular-pubescent, (2) 3-4-lobed, the tube round to
triangular or quadrangular, fused nearly throughout except for the glandular or slightly pubes-
cent lobes, each lobe terminated by an awn, the awns of unequal length and divergent, the
base of the tube slightly ventricose on the angles in some; flowers 1 (2) per involucre, white
to pink, pubescent without along the midrib, on long non-stipitate glabrous pedicels, the
tepals oblanceolate to oblong, entire or erose to fimbriate, united about a third of their
length; stamens 6-9, mostly arranged in two whorls, the filaments glabrous, the anthers
oblong; achenes brown to black, glabrous, the narrow to slightly globose base tapering to a
long, slightly 3-angled beak, the embryo straight, in abundant endosperm; n= 19 (Hardham
1989).

A bispecific genus of California ranging from Monterey and Stanislaus cos. south to San Diego Co. eastward
onto the northwestern edge of the Mojave Desert in Kern and Inyo cos. and in western San Bernardino and
Riverside cos., from near sea level to 5000 (6000) ft elev; flowering from Mar-Jul (Aug).

Key to the Species

A. Bracts unilateral; involucres 1-3 per node, the lobes (2) 3 (4), 25-5 (7) mm long with the awns (0.5) 1-25
(3) mm long; flowers 1-2, the tepals entire; mostly coastal and in the coastal ranges from Monterey Co. south
to San Diego Co. eastward to western San Bernardino and Riverside COS. ........csscssereernene 1. M. californica

AA. Bracts orbicular-perfoliate; involucres 1 per node, the lobes 4, 3-S (6) mm long with the awns (0.3) 0.5-1.2
mm long; flowers 1, the tepals entire or more often erose to fimbriate; coastal ranges and Central Valley
from Stanislaus Co. south to the western edge of the Mojave Desert in Kern and perhaps Inyo cos. ...........

are wasa arte venencoversctanessececesaterseucc conshsnvecsonbavdvacasasantecueasdateosoeas ecsesecoasassurasonte sox itay nconratanernaperassnnrancsansaneeaaseaerss 2. M. perfoliata

1. Mucronea californica Benth., Trans. Linn. Soc. London 17: 419. 1836. Chorizanthe califor-
nica (Benth.) A. Gray, Proc. Boston Soc. Nat. Hist. 7: 149. 1859.—TYPE: without location
data, possibly near Monterey, Monterey Co., or near Santa Barbara, Santa Barbara Co.,
California, “1833,” Douglas s.n. (holotype: K!; isotypes: CGE, GH, K, MO, OXF').

Chonzanthe californica (Benth.) A. Gray var. suksdorfii Macbride, Contr. Gray Herb. 53:
6. 1918. Mucronea californica Benth. var. suksdorfii (Macbride) Goodman, Ann. Missouri
Bot. Gard. 21: 91. 1934.-TyYPE: Surf, Santa Barbara Co., California, 12 Jun 1913,
Suksdorf 146 (holotype: GH!; isotypes, NY, WTU!).

Plants (0.3) 0.5-3 (5) dm high and 1-6 (8) dm across; leaves with the blade (0.5) 1-5 cm
long, (1) 2-8 (12) mm wide, narrowly spatulate to obovate, the petiole 0.5-3 cm long; bracts
3 (5), unilateral, triangular to ovate or oblong, 0.5-1 (2) cm long, acute to obtuse, spreading
to nearly erect, united at least half its length, becoming acerose only at the terminal nodes
and then linear to linear-lanceolate, mostly hirsute or glandular at least on the lower surface
as well as ciliated along the margin, sometimes glabrous on the upper surface, each lobed
tipped with an awn 1-2.5 (3) mm long; involucres 1-3 at each node, cylindric, 2.5-5 (7) mm
long, glandular-pubescent and occasionally hirsute, the tube round to triangular, obscurely
ribbed, the segments united about three-quarters of their length with the (2) 3 (4) free ter-
minal lobes spreading to strongly divergent, glandular or slightly hirsute, bearing divergent
awns of unequal lengths, (0.5) 1-2.5 (3) mm long, the tube not ventricose basally; flowers 1
(2), 1.5-2.5 (3) mm long, white to pink with reddish or greenish midribs, pubescent without
along the midribs or at least at the base, the tepals oblong, entire, united about a third of

204 PHYTOLOGIA volume 66(3):199-220 May 1989

their length; stamens 9, the filaments 1-2 (2.5) mm long, glabrous but minutely papillose, the
anthers 0.5-0.7 mm long, pink to red; achenes 2-3 mm long; n= 19 (Hardham 1989).

Locally common mainly in sandy places along the Pacific Coast from San Luis Obispo Co. south to San Diego
Co. and inland in the southern Coast and Transverse ranges from Monterey Co. to Los Angeles, extreme western
San Bernardino and northwestern Riverside cos., California, from sea level to 3200 (4500) ft elev; flowering from
(Mar) Apr-Jul (Aug).

Representative Specimens. - UNITED STATES. Catirornia: Kern Co.: 0.2 mi N of the confluence of Cedar and
Lumreau creeks on the Farnsworth Ranch near Glennville, 5 Jul 1978, Farnsworth s.n. (CAS). Los Angeles Co.:
Playa del Rey, 10 Jun 1902, Abrams 2507 (BM, DS, E, F, G, GH, K, MO, NEB, NMC, NY, ORE, PH, POM, US,
Z); Cajon Pass, San Gabriel Mountains, 16 May 1931, Epling et al. s.n. (BM, BR, DS, K, LA, MARY, MICH,
MIN, ND, NO, NY, RSA, UCR, WTU); Arcadia, 1901-1905, Grant 173 (E, F, G, GH, ILL, MIN, MO, NMC, PH,
RM, SMU, UC, US, W, WIS); Pasadena, 3 May 1882, M.E. Jones 3221 (BM, BR, CAS, DAO, ILL, LE, MO, MSC,
NY, POM, RM, UC, US, UTC); Point Dume, 4 Jun 1959, Raven & Thompson 14318 (CAS, GH, RSA); San Gabriel
Wash at Arrow Highway, 4 Jun 1932, Wheeler 798 (F, GH, ILL, MIN, NY, OKL, UC, US, Z). Monterey Co.: Bald
Mountain at SE edge of Fort Hunter Liggett Army Base, along the Bryson-Hesperia Road 4.3 km NW of Sapaque
Road, 11 Jul 1978, Broome et al. 2265 (BRY, CAS, MARY, RSA, WIS); Lowes Canyon, 15 mi NE of San Miguel,
15 May 1958, Hardham 3334 (CAS, DAO, RSA); near Pleyto, 29 May 1958, Hardham 3498 (CAS, RSA, SBBG);
10 mi above Carmel on Carmel River, 23 Jun 1923, Mason 544 (DS, GH, NY, UC). Riverside Co.: Wilder’s Canyon,
near Riverside, 15 May 1904, Wilder s.n. (POM). San Bernardino Co.: San Bernardino, May 1913, Jepson 5522
(JEPS, RM, WIS); San Bernardino, Apr 1876, Lemmon s.n. (BM, F, G, JEPS, NEB, OKL); near San Bernardino,
May 1894, Parish 2822 (ILL, MICH, MIN, NY, POM, VT, WIS); Colton, 30 May 1882, Pringle s.n. (CAN, F, G,
GH, K, LE, MIN, MPU, NY, PENN, PH, US, VT). San Diego Co.: Ocean Beach, 27 May 1902, Brandegee 1603
(CAN, CAS, E, G, GH, K, L, LE, MICH, MO, MSC, NY, POM, UC, US, WS); Coronado, 7 May 1891, Dunn
s.n. (B, DS, G, MIN, UC, US); Torrey Pines Park, near La Jolla, 30 Apr 1947, Munz & Everett 11706 (NY, RSA,
WTU). San Luis Obispo Co.: 2.3 mi SE of Genesco School on the road to Creston, 23 May 1955, Ferris 12794
(DS, NY, RSA, WS, WTU); North Traffic Way, Atascadero, 31 May 1958, Hardham 3465 (CAS, DAO, SBBG);
summit at head of Carpenter Canyon, N of Arroyo Grande, 23 Jun 1946, Hoover 6188 (CAS, OBI, OKL, UC);
Yaro Creek, near junction with Toro Creek, 24 May 1947, Hoover 7203 (CAS, DAO, OBI, RSA); 6 mi N of
Guadalupe, 20 Jul 1933, Purer 5088 (DS, GH, POM, SD); along California Highway 58, 05 mi W of Shell Creek
Road, 26 May 1988, Reveal 6911 (BRY, CAS, MARY, MO, NY, OSC, RM, RSA, UTC, WIS); along Los Osos
Valley Road, 02 mi NW of the entrance to Montafia de Oro State Park, San Luis Range, 18 Jun 1987, Reveal et
al. 6480 (ARIZ, BM, BRY, CAS, MARY, MICH, MO, NY, RM, RSA, TCD, US, WIS); Nipomo Mesa N of
Union Oil Refinery and S of the Stauffer Chemical Plant, just off California Highway 1, 7 mi W of Nipomo, 18
Jun 1987, Reveal et al. 6485 (ARIZ, CAS, MARY, MO, NY, RSA, WIS); Jack Lake, 3 mi S of Oceano, 7 Jun 1968,
Thorne 37782 (BRY, NY, RSA, UCR). Santa Barbara Co.: 0.5 mi NW of La Purisima Mission, near Lompoc, 15
Jul 1958, Balls 23538 (BM, CAS, E, RSA, UC, WTU); Burton Mesa, N of Lompoc, 15 Jun 1960, E.R. Blakley 3480
(CAS, RSA, SBBG, UCSB); Surf, May 1902, Elmer 3837 (BKL, CAS, COLO, DS, E, F, G, GH, K, MICH, MIN,
MO, NY, POM, SMU, UC, US, VT, WIS, Z); 5 mi S of Surf, 14 Apr 1929, Ferris 7560 (DS, F, GH, POM, UC);
Purisima Hills, Harris Grade, along California Highway 1, 1.2 mi N of Burton Hill Mesa Road, 22 Jun 1987, Reveal
& Broome 6524 (ARIZ, CAS, MARY, MO, NY, RSA, TCD, WIS); 20 mi E of Santa Maria on California Highway
166 along the Cuyama River at N end of Sierra Madre Mountains, 6 Jun 1954, Sharsmith 4418a (ARIZ, BR, DAO,
ID, IDS, ILL, MIN, OKL, RM, RSA, SMU, UC, US, UTC, WIS, WTU); Graciosa Ridge, SE of Orcutt, W of Mt.
Solomon, 13 Jun 1973, Tilforth & Dourley 841 (MICH, NY, RSA, SD, UC). Ventura Co.: Upper Cuyama River,
on the road to Frazier Mountain, 30 May 1950, Kappler 2067 (LA). A total of 300 collections were examined.

It is not easy to exactly determine where Douglas made his type collection of the genus
Mucronea (Latin mucronis, sharp point, alluding to the awn-tipped involucral lobes). It is
suggested that it likely came from near Monterey in Monterey Co., California, and this is
meant to imply the area along the Carmel River to the south and east. The plants, nonethe-
less, are not so distinctive as to exclude the possibility that Douglas found them near Santa
Barbara where he visited in May of 1832. All of Douglas’ eriogonoid types belonging to the
subtribe Hollisteriineae are dated 1833, but most were likely gathered in California in 1831
or 1832. Douglas was in California on his most extended visit from January 1831 until August
of 1832, a total of nineteen months by his own estimation (McKelvey 1955). He took his

Reveal: Genera related to Chorizanthe (Polygonaceac) 205

California collections with him when he sailed for the Hawaiian (Sandwich) Islands; it was
from here the lot was sent to England arriving there in 1833 which accounts for the date.

The large coastal expression found from Surf, Santa Barbara Co., to the Los Angeles area
has been distinguished as the var. suksdorfii. It is nothing more than a seasonal variation that
may be seen when sufficient moisture has made it possible for individual plants to reach
considerable size.

Mucronea californica was once fairly widely distributed in southern California, but given
the extensive urbanization of the Los Angeles and San Diego areas, the species is now rela-
tively rare in the more southerly counties. In San Luis Obispo and Santa Barbara cos.,
however, the species is common in deep sandy soil and might even be weedy. In general, if
the habitat is badly disturbed the species will gradually disappear. Off-road vehicular activities
on some coastal dunes have certainly resulted in the loss of local populations.

On the eastern foothills of the coast ranges, Mucronea californica can be found growing
near or with M. perfoliata on the distributional or ecological edge of both species. There has
been no indication of hybridization or intergradation of the two although occasionally old
and fragmentary material can be difficult to assign to a species.

2. Mucronea perfoliata (A. Gray) A. Heller, Muhlenburgia 2: 23. 1905. Chorizanthe perfoliata
A. Gray, Proc. Boston Soc. Nat. Hist. 7: 148. 1859.-TYPE: near Fort Tejon, Kern Co.,
California, 1857-1858, Xantus 108 (holotype: GH!; isotypes: DS, K, NY, PH TCD, US!).

Mucronea perfoliata (A. Gray) A. Heller var. opaca Hoover, Leafl. W. Bot. 10: 343. 1966.-
TYPE: Navajo Creek, E side of the La Panza Range, San Luis Obispo Co., California,
31 May 1947, Hoover 7248 (holotype: CAS!; isotypes, OBI, OKL, SD, UTC!).

Plants (0.2) 0.3-2 (3) dm high and 0.5-5 dm across; /eaves with the blade (1) 2-5 cm long,
(2) 3-12 (20) mm wide, spatulate, the petiole inconspicuous; bracts 3, perfoliate, orbicular to
triangular, 0.5-1 (2) cm long, acute to obtuse, spreading to nearly erect, united nearly their
entire length, becoming exceedingly one sided and weakly perfoliate apically, glabrous or
sparsely hirsute and often glandular, ciliated along the margin, each lobe tipped with an awn
0.3-1.2 (1.5) mm long; involucres solitary, cylindric, 3-5 (6) mm long, glandular-pubescent
and usually hirsute especially along the margins of the angles, the tube 4-angled, often curved,
distinctly ribbed and usually corrugate, the segments united about three-quarters of their
length with the 4 free terminal lobes spreading to strongly divergent, glandular or slightly
hirsute, the awns straight, (0.3) 0.5-1.2 mm long, the tube often strongly ventricose basally,
flowers 1, 1.5-3 (3.5) mm long, white to pink with reddish or greenish midribs, pubescent
without along the midribs, the tepals narrowly oblanceolate, entire or more commonly erose
or fimbriate apically, infrequently bilobed, united about a third of their length; stamens 9,
the filaments 1-2.5 (3) mm long, glabrous but minutely papillose, the anthers 0.5-0.8 (0.9)
mm long, pink to red; achenes 2-3 mm long; n= 19 (20) (Hardham 1989).

Locally common in sandy to gravelly places in the Coast and Transverse ranges from Stanislaus Co. south to
Ventura Co. and in the San Joaquin Valley from Kings Co. southward to the Tehachapi Mountains and the

northwestern edge of the Mojave Desert Kern and perhaps Inyo cos., California, from 600-5000 (6000) ft elev,
flowering from Mar-Jun (Jul).

Representative Specimens. - UNITED STATES. CALiFornia: Fresno Co.: S of Curry Mountain on the road from
Coalinga to Parkfield, 13 Jun 1938, Eastwood & Howell 5844 (CAS, ISC, PENN, UC, WTU). Inyo Co.: Hunter
Mountain, Panamint Mountains, Aug 1964, B.D. Rogers s.n. (HSC). Kern Co.: Tehachapi, 1884, Curran s.n. (F,
UC); Maricopa Hills, 15 May 1913, Eastwood 3268 (BM, CAS, GH, K, US); near Taft, 12 Apr 1935, Esau s.n.
(CAS, DAV); Keene, 22 May 1903, M.E. Jones s.n. (CAS, POM, US); 0.6 mi E of California City Boulevard, 35

206 PHYTOL OG1IA volume 66(3):199-220 May 1989

mi SE of California City, 13 Apr 1982, Luckow & Riggins 783-1 (OBI); 1.4 mi N of McKittrick along California
Highway 33, 24 May 1962, Tavares 1249 (CM, ISC, MSC, OBI, POM, RM, TEX, UC); Breckenridge Mountain,
8 Jun 1967, Vasek s.n. (UCR). Kings Co.: Kettleman Hills, near Avenal, 1 May 1938, Hoover 3320 (DS, K, LL,
MICH, NY, UC, US, UTC). Los Angeles Co.: near Fort Tejon Road, 0.5 mi W of Longview Road, 3.5 mi S of
Pearblossom, 19 May 1983, Ballmer s.n. (TEX, UCR); W side of Bob’s Gap Road, N of Bob's Gap, 11 May 1973,
DeBuhr 1031 (1SC); 4 mi E of Palmdale, Antelope Valley, 8 May 1926, Peirson 6726 (NO, RSA); along Mt. Emma
Road, 0.5 mi E of Little Rock Creek, 11 May 1973, Thorne 43549 (MICH, NY, RSA). Merced Co.: Ortigalita
Canyon, 2 mi NE of Ortigalita Peak, 3 May 1940, Mason 12262 (UC). Monterey Co.: Hope Bagby Hidden Valley
Ranch, 28 May 1965, Hardham 12631b (CAS). San Benito Co.: 4.6 mi S of Panoche Road on Idria Road, 20 Apr
1958, Hesse 2454 (UC); San Carlos Creek, San Carlos Range, 13 May 1907, Jepson 2737 (JEPS); 17.6 mi from New
Idria on road to Panoche, 6 May 1956, Raven et al. 9229 (CAS, NY). San Bernardino Co.: Kramer, Jun 1910, K.
Brandegee s.n. (UC); Mojave River, 23 May 1876, Palmer 468 (B, GH, LE, NY, US); Mojave Desert, May 1882,
Parish & Parish 1515 (BM, DS, F, LE, RSA, WU); hills bordering the Mojave Desert, 25 May 1882, Pringle s.n.
(CAN, E, F, G, LE, MPU, NY, PENN, PH, US, VT, WU). San Luis Obispo Co.: Black Mountain, 21 May 1960,
Hardham 5848 (CAS, RSA, SBBG); Pilitas Creek, 24 May 1947, Hoover 7191 (CAS, NY, OBI, OKL); Santa Mar-
garita, 1889, Parry s.n. (ISC, MO, NY); along California Highway 58, 0.6 mi E of the San Juan River bridge, 25
Jun 1978, Reveal 4763 (CAS, GH, MARY, MO, NY, OKL, RSA, TEX); at the junction of the roads E of Navajo
Campground, 17 Jun 1987, Reveal & Broome 6477 (ARIZ, CAS, MARY, MICH, MO, NY, RM, RSA, US, WIS),
summit of Palo Prieta Creek, 7 mi S of Cholame, 7 Jun 1954, Twisselmann 1339 (CAS). Santa Barbara Co.: White
Ledge, Hurricane Deck, San Rafael Mountains, 8 Jun 1969, E.R Blakley 6951 (SBBG); 8 mi N of Manzana
Campground, San Rafael Mountains, 29 May 1959, Hardham 4677 (CAS, DAO); along the Ballinger Canyon Road,
2.3 mi E of California Highway 33, 24 May 1988, Reveal 6895 (BM, BRY, CAS, G, MARY, MO, NY, OSC, RM,
RSA, US, UTC, WIS); 3 mi S of California Highway 166 on California Highway 33, 8 May 1977, RE. Steele 42
(UCR, UCSB). Stanislaus Co.: 24 mi above Camp 74, Puerto Cajion, 11 Jun 1862, Brewer 1261 (CAS, GH, K,
MO, UC, US, WS); Del Puerto Canyon, 17 Apr 1971, Thiers 27168 (SFSU). Ventura Co.: Ozena Valley, 4.5 mi E
of U.S. Highway 399 on the Lockwood Valley Road, 19 Apr 1962, Breedlove 2429 (DS, UCSB); along California
Highway 33, 3 mi below Pine Mountain summit, 17 Jun 1967, Chandler 3545 (SBBG); Quatal Canyon, 5.3 mi E
of California Highway 399, 7 Jul 1963, Piehl 63651 (SBBG); Dry Canyon, San Emigdio Range, 14 Jun 1956, Twis-
selmann 2992 (CAS, DS). A total of 194 collections examined.

The var. opaca was applied by Hoover (1966) to those populations of Mucronea perfoliata
found in the chaparral of the inner Coast Range in San Luis Obispo Co., California. The
bracts, in this expression, are thinly hirsute while those of the plants more to the east and
in more arid situations tend to be glabrous. While this is generally true, the Xantus type of
the species has hirsute bracts, and as this feature is not a consistent one, no taxonomic
differentiation is proposed.

Of perhaps greater interest is the variation in the tepal fimbriation and size. There are
two distinct size classes in Mucronea perfoliata, with one bearing much larger flowers than
the other. In the field the two can be found in close proximity (compare the Ballinger Canyon,
Santa Barbara Co. populations, Reveal 6895 and 6901) and the difference is obvious. Not
only are the flowers much larger but the extent of segmentation of the upper tepal is greater.
In some populations the tepals are entire and therefore similar to those found in M. califor-

nica. No geographic or ecological setting is associated with any one type of flower or tepal
shape.

Hardham (1989) found at least one population of Mucronea perfoliata with a chromosome
number of n= 20. Given the consistency of n= 19 she otherwise reported for the species,
and a similar number in M. californica, and other related genera such as Centrostegia and
very likely Systenotheca, | am reserving judgment on accepting the odd count as common.

The questionable inclusion of Inyo Co. in the distribution of Mucronea perfoliata is based
on a collection made at Hunter Mountain in the Panamint Range (see above). This is well
outside the expected distribution of the species and until it is confirmed, the range extension
must be regarded with some suspicion.

Reveal: Genera related to Chonzanthe (Polygonaceae) 207

4. Centrostegia A. Gray ex Benth. in A. DC., Prodr. 14: 27. 1856. Chonizanthe sect. Centro-
stegia (A. Gray ex Benth. in A. DC.) C. Parry, Proc. Davenport Acad. Nat. Sci. 4: 50.
1884.—TyYPE: Centrostegia thurben A. Gray ex Benth. in A. DC.

Erect to spreading, sparsely glandular annual herbs arising from a thin taproot; leaves
basal, oblong to broadly spatulate, glabrous, tapering to a glabrous indistinctly winged petiole;
flowering stems erect, slender; branches dichotomously branched throughout; inflorescences
cymose, often with the secondaries suppressed; bracts mostly 3, linear to linear-lanceolate,
spreading, awn-tipped, glabrous except for ciliated marginal hairs, united less than a quarter
of their length and positioned to one side of the node; peduncles lacking; involucres solitary,
prismatic and strongly 5-lobed and 3-angled, glabrous, the lobes variously united with each
terminated by a short awn, the base enlarged with 3 saccate, awn-tipped spurs; flowers 2 per
involucre, white to pink, pubescent without at least basally, on long, non-stipitate glabrous
pedicels, the tepals oblanceolate and apically bilobed, united only at the very base; stamens
9, mostly arranged in two whorls, the filaments glabrous, the anthers oblong; achenes brown,
glabrous, the globose base tapering to a long, 3-angled beak, the embryo curved, in abundant
endosperm; n= 19 (Hardham 1989).

A widespread monospecific genus of arid regions in southern and eastern California eastward across western
and southern Nevada to southern Utah, northern and western Arizona, and southward into northwestern Sonora
and northern Baja California Norte, México, from 1250-7700 ft elev, flowering from (Mar) Apr-Jun (Jul).

The genus Centrostegia (Greek kentron, spur, and stegion, roof, alluding to the arched saccate
spurs of the involucre) is now considered to be monospecific. Torrey and Gray (1870) were
the first to add to the genus, proposing C. /eptoceras, while Heller (1910) would add
Chorizanthe insignis after Watson (1877) had reduced Centrostegia to Chorizanthe. Goodman

(1934) transferred C. insignis to Oxytheca while at the same time he moved Chonzanthe
vortriedei to Centrostegia.

The continued association of Centrostegia leptoceras with C. thurberni, the type of the genus,
can no longer be maintained and the former is now referred to a new genus, Dodecahema
(Reveal & Hardham 1989a). Both basal and terminal awns are found in Centrostegia and
Dodecahema. The spinelike awns on the involucre of D. leptoceras represents a fusion of two
sets of bracts. The innermost formed the erect, 6-lobed involucral body with each segment
free or at best only slightly fused by a thin membrane. The basal set of bracts appear to
represent an additional whorl. The six awns of this outermost whorl are long, uncinate and
alternate with the six inner terminal ones. The basal ones arise from a thickened rim at the
base of involucral tube, but this is not in any way an enlargement or inflation.

The spurlike awns found at the base of the involucre in Centrostegia thurberi, as well as
the involucre itself, had an origin independent of that found in any other genus of the Hol-
listeriineae. The five lobes of the involucres are fused nearly throughout their entire length
forming a 3-angled body. Two of the resulting keeled segments of the involucre are con-
structed of two wholly fused involucral lobes so that the apex of each of the resultant keeled
segments is itself slightly lobed with each of the two fused lobes terminated by a short awn.
The remaining keeled segment of the involucral body, specifically the one opposite its point
of attachment at the node, is constructed of only a single lobe. This one may be readily
identified as it is terminated by a single awn.

208 PHYTOLOG1IA volume 66(3):199-220 May 1989

The enlarged saccate base of the involucre ends in a spurlike projection that is terminated
by a short awn. Its origin is not of a second whorl of bracts but rather of a basal expansion
of the involucral lobes themselves. Thus, the involucre of each genus is individually unique
and of totally different origins. The distinctly saccate base of the involucre of Centrostegia
thurber is hinted at in Mucronea and in scattered species of Chorizanthe where ventricose
involucral tubes may be seen, but none is as well developed as in C. thurber.

1. Centrostegia thurberi A. Gray ex Benth. in A. DC., Prodr. 14: 27. 1856. Chorizanthe thur-
beri (A. Gray ex Benth. in A. DC.) S. Wats., Proc. Amer. Acad. Sci. Arts 12: 269.
1877.—TyPE: hillsides near San Felipe, Imperial Co., California, May 1852, Thurber 630
(holotype: GH!; isotypes: F, K, NY!).

Chorizanthe thurberni (A. Gray ex Benth. in A. DC.) S. Wats. var. cryptantha Curran, Bull.
Calif. Acad. Sci. 1: 275. 1886. Centrostegia cryptantha (Curran) Goodding, Bot. Gaz. 37:
53. 1904.-TyPE: Lancaster Station, Los Angeles Co., California, Jul 1884, Curran s.n.
(holotype: CAS!; isotypes, ISC, NY!).

Chorizanthe thurberi (A. Gray ex Benth. in A. DC.) S. Wats. var. macrotheca J.T. Howell,
Leafl. W. Bot. 3: 205. 1943. Centrostegia thurberi A. Gray ex Benth. in A. DC. var. mac-
rotheca (J.T. Howell) Goodman, Leafl. W. Bot. 8: 128. 1957.-TyPE: Alcalde, Fresno
Co., California, 9 May 1893, Eastwood s.n. (holotype: CAS!).

Plants 0.3-2 (3) dm high and (0.6) 1-4 (5) dm across; /eaves with the blade (0.5) 1-3.5 (4)
cm long, 3-8 (10) mm wide, oblong to broadly spatulate, the petiole indistinct; bracts 3 (4),
unilateral, (1) 2-6 (10) mm long, linear to linear-lanceolate, mostly spreading, united less
than a quarter of its length, commonly acerose, glabrous except for the ciliated margins,
awn-tipped, the awns 1-2 mm long; involucres solitary, prismatic and strongly 3-angled, (2)
3-6 (8) mm long, glabrous, the 5 lobes fused along their margins more than four-fifths of
their length into three flattened keeled segments, two of the three keeled segments formed
by the complete fusion of two lobes with the third keeled segment composed of a single lobe,
each lobe terminated by a short, erect awn 0.3-1 mm long, the base of the tube enlarged at
each of the three keeled segments into a saccate, awned horn, the straight awns 0.2-2 mm
long; flowers 2, 2-3 (3.5) mm long, white to pink with reddish or greenish midribs, pubescent
without at the base with short thin hairs, on glabrous pedicels up to 2.5 mm long, the tepals
oblanceolate, apically bilobed, united only at the very base; stamens 9, the outer three on
filaments 2-3 mm long with the anthers soon deciduous, the inner six on filaments mostly
1-2 mm long with the anthers persistent and maturing as a unit after the outer three, the
anthers 0.4-0.5 mm long, pink to red; achenes 2-2.5 mm long; n= 19 (Hardham 1989).

Widespread and common on sandy to gravelly soils in arid regions mainly on the Mojave and Sonoran deserts
of eastern and southern California and in the mountains and foothills bordering the San Joaquin Valley from San
Benito and Fresno cos. southward and from Mono Co. eastward into western and southern Nevada from Washoe
Co. to Clark and Lincoln cos. and across the southern tier of counties in Utah into northern and western Arizona

south to northwestern Sonora and northern Baja California Norte, México, from 1250-7700 ft elev, flowering from
(Mar) Apr-Jun (Jul).

Representative Specimens. - MEXICO. BAJA CALIFORNIA Norte: NE base of Cerro Chichi de la India, Sierra
Juarez, 31 May 1982, Moran 30890 (ARIZ, ASU, CAS, COLO, DAV, ENSB, MARY, MICH, RSA, SBBG, SD,
TEX). SONORA: without location data, without date, Parry et al. 1170 (LE). UNITED STATES. Arizona: Coconino
Co.: N end of Coyote Valley, 0.3 mi S of the Utah line, 14 May 1978, Brown & Parfitt 552 (ASU, BRY); 25 mi
SE of Page along the Page-Kaibito Highway, 28 May 1973, Hevly s.n. (CM, NY). Gila Co.: W side of San Carlos
River near its mouth, 4 Apr 1935, Maguire et al. 10477 (ARIZ, BRY, NY, UTC, WTU); Apache Trail between

Reveal: Genera related to Chonzanthe (Polygonaceae) 209

Roosevelt Lake and Miami, 6 May 1935, Nelson & Nelson 1797 (BRY, G, GH, K, MICH, MO, NY, OKL, PH,
RM, SMU, UC, US, UTC, WTU). Graham Co.: near Fort Thomas, 17 Apr 1940, Peebles 14581 (ARIZ). Maricopa
Co.: W Sycamore Creek, Tonto Basin, 5 May 1931, GJ. Harrison 7799 (ARIZ, US); 12 mi E of Fish Creek, along
Arizona Highway 88, 19 May 1962, Lehto 824 (ASU, OBI); 3 mi W of Wickenburg, 20 Apr 1932, Peebles 8480
(ARIZ, US). Mohave Co.: 8 mi S of Arizona Highway 389, 4 mi W of Fredonia, 19 May 1972, Anvood & Higgins
3925 (BRY, HSC, US, UT); E entrance to Hualape Mountain Park, 6 Apr 1972, Brown et al. 689 (ARIZ, ASU,
ENCB, ID, OBI); Peacock Mountains, 3.5 air mi SE of Hackberry, 7 May 1973, Holmgren & Holmgren 7140 (ASU,
BRY, ID, MONTU, NY, UTC, WTU); Hackberry, 23 May 1884, ME. Jones 4712 (ARIZ, BM, BR, DS, F, G,
MIN, MSC, NY, POM, RM, US, UTC); W of Kingman, 14 Apr 1937, Maguire s.n. (BRY, DAO, NY, UTC, WTU);
Moseby’s Redrock Spring, E side of the Virgin Mountains, 6 Jun 1941, Munz 16770 (POM, WTU). Pima Co.:
along Pontatoc Road near de Grazia’s Mission, N of Tucson, 3 Apr 1958, Matsuda s.n. (ARIZ). Pinal Co.: 46 mi
N of Tucson, 26 Mar 1934, Maguire 10242 (NY, UTC). Yavapai Co.: hills on the E side of Black Mountains, W
of U.S. Highway 93, 6 May 1979, P.C. Fischer 6417 (ARIZ, ASU); 1.6 mi SE of Santa Maria River, U.S. Highway
93, Joshua Tree Parkway, 3 May 1969, Pinkava & Lehto 6188 (ASU, SD). CALIFORNIA: Fresno Co.: Alcalde, May
1891, Brandegee s.n. (UC); W of Coalinga, 13 Jun 1910, Condit 47 (UC). Imperial Co.: San Felipe, May 1855,
Anitisell s.n. (NY). Inyo Co.: Darwin Mesa, 20 May 1891, Coville & Funston 798 (DS, K, NY, US); 3 mi W of Laws,
8 May 1906, Heller 8206 (BKL, DS, E, F, G, GH, ISC, L, MO, NY, PH, UC, US, UTC, WU); E of Wyman Creek,
3.1 mi NE of Deep Springs College, 27 Apr 1986, Morefield & McCarty 3524 (BRY, MARY, MO, RSA, TEX,
UCR); Argus Mountains, Apr-Sep 1897, Purpus 5353 (E, GH, K, MO, US, Z); 2 mi NE of Willow Springs, Last
Chance Mountains, 18 Jun 1955, Roos 6422 (DS, RM, RSA, SD, UC, UCR, US). Kern Co.: 1 mi SW of Bodfish
on road to Havilah, Piute Mountains, 14 Jul 1962, Breedlove 3828 (CAS, DS, RSA, SMU); Kern River Valley, 23
Jun 1891, Coville & Funston 1045 (CAS, DS, K, NY, US); Mojave, 11 May 1917, M_E. Jones s.n. (CAS, DS, MIN,
PH, UC, US); Erskine Creek, Apr-Sep 1897, Purpus 5300 (E, GH, K, MO, US, Z); 1.2 mi S of Eugene Grade, S
of Pine Springs below trail to Mine Spring, Greenhorn Mountains, 26 Jul 1962, C.N. Smith 1115 (SJEPS, RM, UTC,
WTU); Temblor Range, Cedar Canyon, 0.5 mi above the forks, 5 Jul 1971, Twisselmann 17816 (CAS, DAO, OSC,
RSA). Kings Co.: Kettleman Hills near Avenal, 1 May 1938, Hoover 3314 (NY, UC, US). Los Angeles Co.: near
Lancaster, 23 May 1933, Duran 3417 (BKL, BM, BR, BRY, CAN, DAV, DS, F, G, GH, ISC, K, LE, MARY, MIN,
MO, MSC, NO, NY, RM, RSA, UC, US, UTC, WIS, WS, WTU); Acton, Jun 1902, Elmer 3679 (BKL, CAS, COLO,
DS, E, F, G, GH, K, MICH, MIN, NY, POM, US, VT, WIS, Z); 0.2 mi above North Fork Camp, North Fork of
Tujunga Creek, San Gabriel Mountains, 12 Jun 1932, Ewan 7368 (MIN, NO, OKL); Arraster Creek, San Gabriel
Mountains, 10 May 1919, Peirson 28 (CAS, DS, NO, RSA); Portal Ridge N of Elizabeth Lake, 2.1 mi E of Lake
Hughes, 16 May 1988, Reveal 6776 (BM, BRY, CAS, G, MARY, MEXU, MO, NY, OSC, RM, RSA, US, UTC,
WIS); Bob’s Gap, 2 mi ENE of Valyermo, San Gabriel Mountains, 24 May 1973, Tilforth & Dourley 824 (MICH,
NY, RSA, UC). Merced Co.: Piedra Azul Canyon, 4 May 1940, Hoover 4373 (DS, K, NY, UC, US, UTC). Mono
Co.: 1.5 mi below mouth of Pellisier Creek near road, 1.5 mi SW of Mt. View and Proctor Mine, White Mountains,
19 Apr 1986, Morefield & McCarty 3457 (BRY, MARY, MO, RSA, TEX, UCR). Riverside Co.: N base of Eagle
Mountains, 15 May 1941, Alexander & Kellogg 2176 (DS, GH, ISC, POM, UC, UTC, WS, WTU); 15 mi SW of
Twentynine Palms, 25 mi N of Salton View, Joshua Tree National Monument, 29 May 1971, Henrickson 5573
(ASU, CHSC, HSC, UNLV); Kenworthy, San Jacinto Mountains, 21 May 1922, Munz & Johnston 5508 (POM).
San Benito Co.: Griswold Hills at junction with Griswold Creek and Vallecitos Road, 10 May 1936, Lyon 796
(CAS, UC). San Bernardino Co.: S of Victorville, 5 Jun 1941, Alexander & Kellogg 2301 (GH, MO, NY, RM, UC,
UTC, WS); Horsethief Canyon, 15 May 1935, Clokey & Anderson 6598 (BKL, IDS, ILL, NY, POM, RENO, UC,
WIS, WS, WTU); 5 mi E of Cima, 4 May 1935, Munz 13948 (DS, F, POM, UC, UTC, WTU); Mojave Desert,
May 1881, Parish & Parish 895 (B, BR, DS, F, G, ISC, LE, MIN, PH, US, WS, WU); Mojave Desert, 24 May 1882,
Pringle s.n. (CAS, F, G, LE, MPU, NY, PENN, PH, US, VT, WU); summit of the Barstow-Cave Springs Road,
29 Apr 1935, Wolf 6584 (GH, RM, RSA, TEX, WS, WTU). San Diego Co.: Jacumba, 31 May 1903, Abrams 3655
(BM, DS, E, F, G, GH, K, MO, NEB, NMC, NY, PH, US, Z); Wagon Wash, near Sentenac Canyon, below San
Felipe Valley, 20 Apr 1928, Jepson 12488 (JEPS); San Felipe Valley, 17 May 1925, Keck & McCully 68 (POM, UC);
plains E of Campo, 2 Aug 1898, Stokes s.n. (MIN, NEB). San Luis Obispo Co.: between San Juan River and
Carriza Plain, 2 Jun 1946, Hoover 6137 (CAS, DAO, OBI, OKL, RSA); Caliente Range, Padrones Spring Canyon,
3 May 1957, Twisselmann 3466 (CAS). Santa Barbara Co.: Sisquoc River below Lower Bear Camp, San Rafael
Mountains, 30 Jun 1961, E.R. Blakley 4537 (CAS, RSA, SBBG, UCSB); Hurricane Deck, 20 May 1957, Hardham
1974 (CAS, RSA); Sierra Madre Mountains, along the lower slopes and ridges of Newsome Canyon, 24 May 1988,
Reveal 6888 (BRY, CAS, MARY, MO, OSC, RSA, WIS). Tulare Co.: 15 mi from Lamont Meadow on road to
Long Valley, Kern Plateau, 30 May 1967, Howell & True 42452 (CAS, DAV); Kern River N of Kernville, Sequoia
National Forest, 1 May 1935, Mason 8367 (UC, UTC, WIS, WS, WTU). Ventura Co.: near the Frazier Borax Mine,
Mt. Pinos, 12-14 Jun 1908, Abrams & McGregor 224 (DS, E, G, NY, US); Sandstone Camp, upper Sespe Creek,
9 Jun 1963, E.R. Blakley 6007 (OKL, RSA, SBBG); Apache Creek, upper Cuyama River Valley, 13 May 1960,

210 PHYTOL OGIA volume 66(3):199-220 May 1989

Hardham 5683 (CAS, RSA, UC); Long Grade Road N of Thorn Meadow, 16 Jul 1962, Pollard s.n. (CAS, G, ISC,
RM); Santa Ynez Mountains, along California Highway 33, 3 mi S of the turnoff to Pine Mountain, 24 May 1988,
Reveal 6873 (BM, BRY, CAS, G, MARY, MO, NY, OSC, RM, RSA, UTC, WIS); Lockwood Valley, 24 Jun 1987,
Reveal & Broome 6569 (ARIZ, CAS, MARY, MO, NY, RSA, TCD, WIS); Quatal Canyon, San Emigdio Range,
7 Jun 1955, Twisselmann 2109 (CAS, RSA, US). Nevapa: Clark Co.: W slope of McCullough Mountains, 16 mi
W of Searchlight, 11 May 1964, Cronquist 9975 (GH, NY, UTC, WTU); Timber Mountain, 15 mi NW of Searchlight,
24 Apr 1938, Train 1480 (GH, RENO, UC). Esmeralda Co.: Roosevelt Wells, 24 Jun 1973, MJ. Williams 73-G-2
(RENO). Lincoln Co.: Mormon Mountains, Jul 1906, Kennedy & Goodding 83 (MO, NY, UC, US). N end of the
Meadow Valley Mountains, 17 May 1983, Tiehm 7656 (BRY, COLO, DAO, NEB, NY, RSA, UTC); Delamar
Valley, 0.8 mi from pole line road to Delamar, 25 May 1980, Tiehm & Williams 5713 (CAS, MO, NESH, NY,
RSA, UTC). Nye Co.: Saddle Road, 1.5 mi N of Schocken Road, Jackass Flat, 23 Apr 1966, Beatley 3682 (DS,
NTS, RENO); NE Bullfrog Hills, 1.5 mi NW of Springdale, 28 Apr 1971, Beatley & Reveal 12339 (NTS, RENO,
RSA); SW of Pinyon Butte, E of Pahute Mesa Road and 1.5 mi S of Airport Road, 8 Jun 1968, Reveal 1188 (DS,
NTS, NY, RENO); S end of the Quinn Canyon Range on N side of Highway 25 at Queen City Summit, 16 May
1982, Tiehm 6925 (CAS, MO, NY, RM, RSA, UTC). Washoe Co.: on road from Pyramid Lake to Reno, Jun 1927,
Eastwood 14738 (CAS). UTAH: Kane Co.: W face of Johnson Canyon, NE of Kanab, 5 Jun 1969, Atwood 1789
(BRY, NY); Kanab, 1872, E.P. Thompson s.n. (GH); left fork of Last Chance Canyon along the Escalante Road,
10 mi E of Four Mile Bench, 23 May 1975, Welsh 12721 (BRY, CPH). San Juan Co.: Surprise Valley along Nasja
Creek, N side of Navajo Mountain along the Navajo Mountain-Rainbow Bridge Trail, 10 May 1979, Albee 4459
(UT). Unknown Co.: without location data, 1877, Palmer 423 (LE, MO, NY, US); without location data, 1874,
Parry 232 (CM, F, G, K, MIN, NY, PH, US). Washington Co.: E slope of Pine Valley Mountains, 3 mi NW of
Leeds, 19 May 1973, Atwood 4963 (BRY, RM); 20 mi SE of Hurricane, 16 May 1965, Cronquist 10095 (ARIZ, BR,
CAS, COLO, DAO, DS, GH, ID, ILL, ISC, MIN, NY, OKL, OSC, RM, RSA, TEX, UC, W, WTU); Diamond
Valley, 16 May 1902, Goodding 817 (G, GH, MO, NEB, NY, RM, US); below Anderson’s Ranch, 3 May 1932,
Maguire & Blood 1325 (GH, MO, POM, RM, UTC); East Zion Plateau, 22 May 1973, R.A. Nelson 10293 (BRY);
Lytle Ranch Preserve, Beaverdam Wash, 4 May 1986, Welsh 23719 (BRY). A total of 651 collections examined.

In the herbarium a distinction has been made between the var. thurberi mainly of the Mojave
and Sonoran deserts and the var. macrotheca restricted to the eastern edge of the inner Coast
Ranges and in the southwestern San Joaquin Valley of central California. At the extreme
the two are markedly distinct, with the involucres of var. thurber’ being generally 2-6 mm
long and 3-S mm wide while those of the var. macrotheca are mostly (4) 5-8 mm long and
5-8 mm wide. A total of sixteen collections were distinguished as the latter variety, but of

that number only twelve are unequivocal. Other than the type cited above, these are as fol-
lows:

Fresno Co.: Alcalde, May 1891, Brandegee s.n. (UC); W of Coalinga, 13 Jun 1910, Condit 47 (UC). Kem Co.: 5
mi W of Toad Springs, 22 Jun 1967, Philbrick s.n. (SBBG). Kings Co.: Kettleman Hills near Avenal, 1 May 1938,
Hoover 3314 (NY, UC, US). Merced Co.: Piedra Azul Canyon, 4 May 1940, Hoover 4373 (DS, K, NY, UC, US,
UTC); San Benito Co.: Griswold Hills at junction with Griswold Creek and Vallecitos Road, 10 May 1936, Lyon
796 (CAS, UC); Griswold Hills, 10 May 1936, G.S. Lyon 796 (CAS); Panoche Hills, 19 Apr 1945, G.S. Lyon 1686
(CAS). San Luis Obispo Co.: hills between Carriza and Cuyama, 13 Jun 1902, Eastwood s.n. (CAS); between San
Juan River and Carmza Plain, 2 Jun 1946, Hoover 6137 (CAS, DAO, OBI, OKL, RSA); Caliente Mountain, 1 Nov
1952, Hoover 8268 (CAS, OBI); Caliente Range, Padrones Spring Canyon, 3 May 1957, Twisselmann 3466 (CAS).

Several collections from northern Ventura and extreme northwestern Los Angeles cos. are
intermediate in involucral size and tend to break down the distinctiveness of the var. mac-
rotheca. Likewise, a few from eastern San Luis Obispo Co. approach the var. macrotheca but
not to the extent found in the mountains to the south. It is because of the tendency for the
two extremes to merge that no taxonomic distinction is now made.

7. Hollisteria S. Wats., Proc. Amer. Acad. Arts 14: 296. 1879. Eriogonum Michx. sect. Hol-

listeria (S. Wats.) Roberty & Vautier, Boissiera 10: 92. 1964.—TyYPE: Hollisteria lanata
S. Wats.

Reveal: Genera related to Chonzanthe (Polygonaceae) 211

Spreading, white-wooly tomentose annual herbs arising from a thin taproot; leaves basal
and cauline, the basal ones oblanceolate, few to many, mostly erect to spreading, tapering
to an indistinct petiole, the cauline ones elliptic to ovate, cuspidate, solitary at each node,
sessile or essentially so, gradually reduced above; flowering stems few to many arising from
the basal rosette, wiry, dichotomous; branches numerous and often diffuse, slender, variously
tomentose, dichotomously branched throughout; inflorescences cymose, open to diffuse, often
with the secondaries suppressed, each node bearing a sessile or short-pedicellate flower;
bracts 3, the lateral two distinct, linear to linear-lanceolate, thinly pubescent, usually several
times longer than the obscured third, this linear and typically densely tomentose, situated at
the base of the cauline leaf and obscured by it, all awn-tipped; peduncles lacking; involucres
1, reduced to a series of 3 (4) involucral bracts, linear, pubescent with long curly hairs,
awn-tipped; flowers solitary, yellowish with a yellowish-green to green midrib, densely tomen-
tose without, glabrous within, the pedicel short and peglike, the tepals 6, petaloid, lanceolate
to ovate, mucronate, united only at the base; stamens 6, included and attached at the base
of the sinsus of the united lobes and alternating with the tepals or if 9 then with the outer
3 opposite the alternating tepals and situated on the fused portion of the floral tube, the
filaments glabrous, the anthers oval to oblong, yellow; achenes brown to dark black, glabrous,

the globose base tapering to a short stout, 3-angled beak, the embryo curved, in abundant
endosperm; n= 21.

A locally common monospecific genus of the San Joaquin Valley and Coast Ranges of central California, from
50-3200 ft elev; flowering from (Mar) Apr-Jun (Jul).

Hollisteria (William Welles Hollister, 1818-1886, California sheepman and rancher for whom
the city of Hollister, San Benito Co., California, was named in 1868) has long been difficult
to place in any subfamilial scheme. The genus is similar to Chorizanthe and Jones (1908)
inadvertently described a specimen of H. /anata as a new species of Chorizanthe. Dammer
(1892) and Gross (1913a) associated Hollisteria with Nemacaulis, a view sustained by Jaretzky
(1925) in his doctoral dissertation on the genera of Polygonaceae. Curran (1885) felt the
genus was probably most closely related to Eriogonum. At that time she had submerged
Nemacaulis in Eriogonum, and her conclusion was most logical. Stokes (1936) retained
Nemacaulis in Eniogonum, but like Curran did not transfer Hollisteria. In 1964, Roberty and
Vautier made the combination, E. /anatum, thus completing the reduction of the genus. Hol-
listeria has been retained in the major California floras (Jepson 1913, 1925; Abrams 1944;
Munz 1959) and was recognized by Shields and Reveal (1988) who associated the taxon with
Eriogonum and placed it closest to Nemacaulis.

The transfer of Hollisteria from the Eriogonineae to a taxon that includes such genera as
Chorizanthe (and in a subtribe that must be called Hollisteriineae) is. not suggested without
some trepidation. What is emphasized here is the solitary flower subtended by a whorl of
awned involucral bracts. In Nemacaulis each flower is subtended by a single non-awned in-
volucral bract (not several) and the numerous flowers are congested into clusters. The flower
clusters (glomerules) of Nemacaulis are concentrated at each node of the axises along the
dichotomous branches (Reveal & Ertter 1980) whereas in Hollisteria there is only a single
flower at each node. The aneuploid chromosome number of n= 21 is further evidence of
Hollisteria being more closely related to Chorizanthe where the aneuploid number is known
rather than Eriogonum where it is currently unknown (Shields & Reveal 1988).

<

212 PHYTOL OG1IA volume 66(3):199-220 May 1989

1. Hollisteria lanata S. Wats., Proc. Amer. Acad. Arts 12: 296. 1879. Eriogonum lanatum (S.
Wats.) Roberty & Vautier, Boisseria 10: 96. 1964.—TyPE: Hollister Ranch, Cholame
Valley, Monterey Co., California, 8 Jul 1878, Lemmon s.n. (holotype: GH!; isotypes:
BM, CAS, CM, DS, F, G, GH, ISC, JEPS, K, MIN, MO, NY, P, PH, RSA, UC, US!).

Chonzanthe floccosa M.E. Jones, Contr. W. Bot. 12: 74. 1908.-TyPE: Bakersfield, Kern
Co., California, 23 May 1903, M.E. Jones s.n. (holotype: POM!; isotype, DS!).

Plants 0.3-0.8 (1) dm high and (0.5) 0.8-3 (5) dm across; leaves basal and cauline, the basal
ones oblanceolate, (1.5) 2-5 (6) cm long including the winged petiole, (2) 3-7 (9) mm wide,
sparsely pubescent on both surfaces, green but soon turning brown, quickly deciduous, the
cauline ones elliptic to ovate, 0.5-2.5 (3) cm long, (1) 3-8 mm wide, sparsely to densely
tomentose, green to white or tannish, solitary at each node, sessile or nearly so, (0.5) 1-3.5
(4) cm long, 3-8 (10) mm wide; bracts 3, the lateral two linear to linear-lanceolate, 2-5 mm
long, thinly pubescent, the third one linear, 1-2 (3) mm long, densely tomentose; involucres
reduced to a series of 3 (4) involucral bracts, linear, 1.5-2 mm long, mucronate, densely
pubescent with long curly hairs on the outer surface, glabrous within or at most with only a
few scattered hairs; flowers solitary, 1.5-2 mm long, yellowish with a yellowish-green to green
midribs, densely pubescent without, glabrous within, on short glabrous pedicels up to 1 mm
long, the tepals lanceolate to ovate, apically mucronate, united about a quarter of their length,
the floral tube campanulate with a minute nectiferous disk at the base of the inner whorl of
stamens; stamens 6-9, the filaments 0.8-1.2 mm long, the anthers 0.4-0.5 mm long, yellow;
achenes 1.7-2 mm long; n= 21.

Locally rare to common on sandy to gravelly or clayey soils in arid regions in the San Joaquin Valley and on
the slopes of the eastern Coast Ranges from Merced and Monterey cos., California, south to the northern edge

of the Tehachapi and Transverse ranges from Kern to Santa Barbara cos., from 50-3200 ft elev, flowering from
(Mar) Apr-Jun (Jul).

Representative Specimens. - UNITED STATES. CaLiForniA: Fresno Co.: 9 mi S of Kerman, 27 May 1937, Hoover
2327 (DS, GH, K, LL, NY, UC, US, UTC, WS); between Arroyo Hondo and Cantua Creek, 1 May 1938, Hoover
3289 (DS, K, NY, UC, US, UTC, WS); Zapato Chino Creek, 12 Apr 1930, Jepson 15361 (CAS, JEPS, WS); 3 mi
E of Mercy Hot Springs, 12 Jun 1967, Twisselmann 13362 (CAS, SBBG, RSA). Kern Co.: Maricopa Hills, 15 May
1913, Eastwood 3267 (BM, CAS, GH, K, NY, US); near Edison, 27 Apr 1937, Eastwood & Howell 4004 (CAS,
MIN, PENN, RSA, UC); 3 mi SE of Bakersfield, 16 Apr 1932, Ferris & Bacigalupi 8047 (CAS, DS, F, MICH,
MONTU, ND, PH, RSA, SMU, UC, US); Oil City near Bakersfield, 22 Apr 1905, Heller 7741 (DS, E, F, G, GH,
ISC, L, MICH, MO, NY, P, PH, RSA, UC, US, WIS); Edison, Greenhorn Mountains, 11 Apr 1937, Hoover 1818
(DS, K, LL, MICH, NY, UC, US, UTC, WS); along California Highway 58, 1.2 mi W of California Highway 33
near McKittrick, 25 Jun 1978, Reveal 4771 (CAS, K, MARY, MICH, NY, TEX); 7 mi E of Bakersfield, 21 May
1958, Rose 58037 (ARIZ, BRY, COLO, DAO, E, G, GB, MIN, RM, RSA, SMU, US, UC); 4 mi E of Bakersfield
along California Highway 178, 20 May 1962, Rose 62030 (BR, CAS, COLO, DAV, DS, ENCB, GB, HSC, MICH,
MO, OBI, RSA, TEX); summit of Devil’s Water Canyon, Temblor Range, 5 Jul 1971, Twisselmann 17811 (CAS,
NY, OSC, RSA). Kings Co.: Kettleman Hills near Avenal, 1 May 1938, Hoover 3310 (DS, GH, K, NY, UC, US,
UTC, WS). Merced Co.: Los Banos Hills, 28 May 1938, J.T. Howell 13827 (CAS, ISC, LA, RSA, SD); Ortigalita
Canyon, 2 mi NE of Ortigalita Peak, 4 May 1940, Mason 12266 (ARIZ, BRY, CAN, CAS, DAO, DAV, DS, F,
GH, ISC, K, MICH, MIN, MO, NO, NY, ORE, PENN, POM, RM, SBBG, SMU, UC, US, UTC, W, WS, WTU).
Monterey Co.: bed of Salinas River at San Miguel, without date, Norton s.n. (CAS). San Benito Co.: along California
Highway 180, 16.5 mi SE of Panoche, 6 May 1956, McCaskill 436 (ARIZ, DAV, OSC, UTC, WTU); 5 mi NE of
Llanada, Little Panoche Pass, 19 Apr 1938, Rose 38183 (CM, RM, SD, UC). San Luis Obispo Co.: 2 mi E of La
Panza, 8 May 1936, Eastwood & Howell 2326 (CAS, F, K, MICH, NY, PENN, RSA, US); Recruit Canyon, Temblor
Range, 4 Jun 1967, Hoover 10615 (CAS, ENCB, OBI, OSC); 2 mi E of La Panza, 8 May 1936, Rose 36257 (GH,
L, LE, MO, NY, US, W, WTU). Santa Barbara Co.: Cottonwood Creek Wash, near Bates Canyon Road crossing,
S side of Cuyama Canyon, 18 May 1965, Chandler 2289a (SBBG, UCSB); along the Ballinger Canyon Road, 0.5
mi E of California Highway 33, 24 May 1988, Reveal 6893 (BM, BRY, CAS, G, MARY, MEXU, MO, NY, OSC,

Reveal: Genera related to Chonzanthe (Polygonaceac) 213

RM, RSA, US, UTC, WIS). Tulare Co.: 4.5 mi ENE of Pixley, Pixley Nature Area, 15 May 1968, J.T. Howell 44474
(CAS). A total of 129 collections were examined.

Hollisteria lanata is a \ow, annual that spreads on the ground from a thin taproot. The size
of the mature plant is determined by the amount of available moisture so that individuals
may be a few centimeters across to several decimeters. The flowers are numerous, and es-
pecially so on the larger plants, with each one producing an achene that apparently is not
widely dispersed. While indirect evidence would point to some bird dispersal, the plant is
often local and while common, it is not weedy.

Hardham (1989) found Hollisteria lanata to be n= 21 or occasionally n= 22. My own
counts of the species were consistently of the former number (voucher by Reveal 6893) and
therefore only the one number is accepted here.

8. Lastarriaea Rémy in Gay, Fl. Chil. 5: 289. 1851-1852.—TyYPE: Lastarriaea chilensis Rémy
in Gay.

Prostrate to ascending, thinly pubescent annual herbs with several stems arising from a
thin taproot; /eaves basal or slightly sheathing, the blades linear, hirsute and sometimes rather
densely so along the margins, on an indistinct petiole, often with one or more flowers inter-
mixed with leaves; flowering stems few or more often numerous and diffuse; branches slender
and often brittle, readily disarticulating at the nodes especially in age, yellowish-green to
green or red, glabrous or more often thinly pubescent with slightly curled glandular hairs,
dichotomously branched throughout; inflorescences cymose, mostly diffuse, often with the
secondaries suppressed, each node bearing a sessile flower; bracts 2, lanceolate to narrowly
ovate, erect to spreading, the awns uncinate; peduncles lacking; involucres solitary, reduced
to a series of 3 linear to broadly lanceolate involucral bracts, pubescent on the margin, the
awns uncinate; flowers 1, light green to greenish-white, thinly pubescent without, glabrous
within, the pedicels lacking, the tepals 5, coriaceous, lanceolate, acute with a recurved, hooked
awn, united about half to three-quarters of their length; stamens 3, the filaments short,
glabrous, the anthers cream to white, oval; achenes brownish, glabrous, the narrow base
tapering to a narrow 3-angled beak, the embryo straight, in abundant endosperm.

A small genus of three species on the off-shore islands, along the coast and in the coastal mountains of south-
western California south to central Baja California, then disjunct to the deserts of northerm and central Chile,
from near sea level to 4600 ft elev, flowering from Sep-Dec or (Feb) Mar-Jun.

Key to the Species

A. Bracts 0.4-2 cm long, lanceolate; involucral bracts linear to lanceolate, 0.3-1 (1.2) cm long, 0.3-1 mm wide;
North America.
B. Bracts narrowly lanceolate, 0.4-1.5 (2) cm long, 0.8-1.5 (2) mm wide, distinctly hirsute, the awns (0.5)
1-2 (2.5) mm long; involucral bracts linear, 0.3-1 (1.2) cm long; California and northern Baja California
as) far’south vas ithe: Sterral San) BOLaS cccserresscxcccrsnsessastarsecosonsorcncirarcerassdeavessnncserzusescereeevoscercseseteres 1. L. coriacea
BB. Bracts lanceolate, 05-2 cm long, 1-2 mm wide, spreading pubescent, the awns 0.3-0.6 mm long; involucral
bracts lanceolate, 0.15-0.4 cm long; central Baja Califormia. ............::csessessesessesseseeseeseseesenesneees 2. L. ptilota
AA. Bracts 0.2-0.6 (1.5) cm long, 0.2-15 (3) mm wide, broadly lanceolate to narrowly ovate; involucral bracts
broadly lanceolate, 0.2-05 (1.5) cm long, 0.8-2 (2.5) mm wide; South America. ..........:ccceceseesess 3. L. chilensis

The taxonomic distinctiveness of Lastarriaea (José Victorino Lastarria Santander, 1817-1888,
lawyer and founder of the Liberal party in Chile, author and political leader who served as
a government minister, ambassador, delegate and senator, and as a member of Chile’s su-

214 PHYTOLOGIA volume 66(3):199-220 May 1989

preme court) has been greatly debated since Parry (1884) briefly reduced it to synonymy
under Chorizanthe, proposing a superfluous name C. Jastarriaea for the one species that he
recognized. Lastarriaea was originally described by Rémy (1851-1852) who established the
name on a South American species he called L. chilensis. He was not the first to find the
plant or even to recognize that it represented a new genus. A Poeppig collection gathered
in Chile from 1827-1829 was annotated with a new name by Kunze while a Guliemin collection
made in 1830 was given a different name. In North America, plants had been found by Nuttall
in the mid 1830s during his visit to the California coast to which he proposed various her-
barium names. Like the others none of Nuttall’s names were ever published (Parry 1885).

Bentham (1856) was unable to place Lastarriaea within Polygonaceae for some argued (at
least on herbarium labels) that the genus might be better placed in the Caryophyllaceae near
the genus Paronychia than in the Polygonaceae.

Gross (1913a) was the first to suggest that the North American expression might be dif-
ferent from its South American relative. He proposed Lastarriaea chilensis subsp. californica
but did not indicate if his new taxon was native to California or not. Goodman (1934) main-
tained the North American plants as distinct from those in Chile, calling it Chorizanthe las-
tarriaea var. californica. Soon thereafter, Goodman (1943) was able to study Chilean
specimens in some detail and came to the opinion the North American plants represented
a distinct species which he named C. coriacea. Hoover (1966) would eventually transfer that
epithet to Lastarriaea.

In 1885, Parry received a suite of chorizanthoid specimens from Rudolph A. Philippi, the
noted Chilean botanist. Philippi sent him a series of specimens belonging to Lastarriaea that
he had divided into three species, L. chilensis, L. stricta and L. linearis. Neither of the new
species had been described so that when Parry (1886) reversed his view of the previous two
years and recognized Lastarriaea as a good genus, he validly published the new species before
Philippi was able to propose them.

Various interpretations of the involucre and flower of Lastarriaea have been put forward.
Parry (1884) considered the flower equivalent to the involucre of Chorizanthe but failed to
explain how a 6-lobed involucre could be identical to a 5-lobed coriaceous flower. Curran
(1885) almost immediately challenged this notion. Goodman (1934) maintained Lastarriaea
in Chonizanthe contending that C. interposita was a true intermediate stage between the two
genera. Since then it has been possible to see more material than the single specimen of C.
interposita available to Goodman and the perianth of that species is not at all “involucriform.”
Rather, it is a typical 6-lobed flower similar to the kind found throughout Chorizanthe.

In the present treatment the bractlike structures subtending the flowers of Lastarriaea are
interrepted as involucral bracts. This is a major change but given the placement of the two
bracts around the node and three additional ones (“involucral bracts”) immediately subtend-
ing the flower, this appears not to be unduly foreign considering that a similar sequence is
seen in Hollisteria and even Dedeckera.

In our evaluation of characters prepared in concert with the revision of the annual species
of Chorizanthe (Reveal & Hardham 1989b), the distinctiveness of Lastarriaea from
Chorizanthe was called into question. Lastarriaea coriacea proved not to be that closely related
to C. interposita, as suggested by Goodman (1934), but in fact to be more similar to C.
brevicomu. Nonetheless, Lastarriaea is maintained as emphasis is placed on the reduction of
the involucre to a highly modified series of involucral bracts that resemble cauline leaves or
the foliar bracts rather than a true tube as in Chorizanthe. In addition, the fundamental
differences in the flowers is considered significant. It is not suggested, however, that Lastar-
riaea evolved from something other than Chorizanthe nor is it proposed that the genus nec-

Reveal: Genera related to Chorizanthe (Polygonaceac) 215

essarily came from a pre-Chonzanthe ancestor. The taxon is clearly a recently differentiated
genus and it certainly evolved from Chonzanthe. Its placement in the linear sequence close
to Hollisteria is because Lastarriaca represents a single, isolated branch off Chonizanthe while
Hollisteria is representative of another independent and isolated branch.

1. Lastarriaea coriacea (Goodman) Hoover, Leafl. W. Bot. 10: 342. 1966. L. chilensis Rémy
in Gay subsp. califomica H. Gross, Bot. Jahrb. Syst. 49: 435. 1913, non Chorizanthe
californica (Benth.) A. Gray. Chorizanthe lastarnaea C. Parry var. californica (H. Gross)
Goodman, Ann. Missouri Bot. Gard. 21: 33. 1934. Chorizanthe coriacea Goodman, Leafl.
W. Bot. 3: 230. 1943.—-LECTOTYPE: without location data but perhaps from San Diego,
San Diego Co., California, 1882, Parry s.n. (lectotype: B!; duplicates of the lectotype: F,
GOK EE NY. ORES US!):

Plants 0.2-1.5 dm high and 0.5-3 (5) dm across; /eaves basal, 0.5-3 cm long, 0.2-0.8 (1) mm
wide, linear, hirsute along the margins; branches slender and brittle, readily disarticulating,
yellowish-green to green or red, pubescent with curled hairs; bracts 2, 0.4-1.5 (2) cm long,
0.8-1.5 (2) mm wide, narrowly lanceolate, hirsute, erect or nearly so, the uncinate awns (0.5)
1-2 (2.5) mm long; involucral bracts 0.3-1 (1.2) cm long, 0.3-1 mm wide, linear, hirsute, erect,
the uncinate awns 0.5-2 mm long; flowers 2-3.5 mm long (including the awns), light green to
greenish-white, thinly pubescent without, the tepals narrowly lanceolate, awn-tipped, united
more than three-quarters of their length; stamens 3, the filaments 0.5-1 mm long, the anthers
0.2-0.3 mm long; achenes 2.5-3 mm long; n= 21, 30 (Hardham 1989).

Infrequent to locally common on sandy to gravelly soils along the coast of California and Baja California Norte,
México, from Sonoma Co. south to the El Rosario area, inland into the southern Coast Ranges, the Central Valley
and the foothills of the Sierra Nevada from Calavaras Co. south to western Riverside Co., and to the Sierra Borja
of central Baja California Norte, from near sea level to 2750 ft elev, flowering from (Feb) Apr-Jun.

Representative Specimens. - MEXICO. Basa CALIFORNIA Norte: Cuesta de la Piedra Parada, Sierra San Boma,
20 Mar 1966, Moran 12838 (SD, UC); 2 mi NE of Las Escobas, 21 Apr 1975, Moran 21848 (ASU, CAS, LL, RSA,
SD); Arroyo Nueva York, 2 mi SE of the ranch site, 22 Apr 1975, Moran 21877 (ENCB, MO, RM, SD, UC); 11
km E of Tecate, 7 Jun 1980, Moran 28700 (ARIZ, SD); 10 mi E of El Rosario, 24 Feb 1973, Moran & Reveal
20272 (COLO, DAV, SD, UC, US); Arroyo el Socorro, 4 mi from its mouth, 25 Feb 1973, Moran & Reveal 20279
(MICH, RSA, SD, US); Todos Santos, 7 Apr 1886, Orcutt s.n. (MO); San Quintin Bay, Feb 1889, Palmer 722 (GH,
K, MEXU, US, WU); 3 mi E of Uruapan, 21 May 1988, Reveal 6838 (BM, BRY, CAS, MARY, MEXU, MO, NY,
OSC, RM, RSA, UTC, WIS); 7 mi W and S of México Highway 1 at San Quintin, 25 Mar 1988, Reveal et al. 6760
(CAS, MARY, MO, NY, RM, RSA, US, WIS). UNITED STATES. Cauirornia: Calaveras Co.: 1.7 mi SW of
Burson, 20 May 1935, Roseberry 19] (RSA, UC). Contra Costa Co.: Antioch, 1868-1869, Kellogg & Harford 865
(BM, CAS, G, MO, NY, US). Fresno Co.: Mill Creek Valley, off Wonder Valley Dude Ranch Road, 9 May 1959,
Bacigalupi et al. 7055 (JEPS, RM, UTC, WS); 1 mi SE of Big Panoche Creek near PG&E Line Road, 12 Apr
1936, Lyon 795 (UC). Imperial Co.: Pigeon Pass in the Box Spring Mountains, 2 May 1967, Clarke s.n. (UCR).
Kern Co.: Poso Flat, Greenhorn Range, 2-10 Jun 1904, Hall & Babcock 5022 (DS, UC); near White River, 13 May
1973, Hardham 19131 (CAS); 8 mi NW of Lockern, 26 Mar 1937, Johannsen 1406 (RSA, UC); The Crater, Adobe
Canyon, 26 Apr 1965, Twisselmann 10618 (CAS, RSA). Kings Co.: Kettleman City, 1 May 1938, Hoover 3332 (K,
NY, UC, US). Los Angeles Co.: Ballona Harbor, 1 Apr 1901, Abrams 1218 (DS, MO, POM, Z); upper end of
Bulrush Canyon, Santa Catalina Island, 12 May 1965, Thorne & Everett 34600 (RSA); San Gabriel Wash at Arrow
Highway, 26 May 1932, Wheeler 745 (CPH, DS, LA, MO, ND, OKL, RSA). Madera Co.: 8 mi W of Chowchilla,
13 Apr 1935, Hoover 525 (K, NY, UC, US). Monterey Co.: 1 mi S of San Lucas, 29 Apr 1957, J.M. Anderson et
al. 436 (COLO, F, GH, NY, OKL, UC); along Del Venturi Road, Fort Hunter Liggett Reservation, 17 Jun 1978,
Broome & Cagle 2168 (CAS, MARY, RSA, WIS); Monterey, 18 Jul 1882, Pringle s.n. (E, F, G, LE, MIN, MO,
NY, PENN, PH, US, VT, WS, WU); 3 mi NW of McKay, 9 Apr 1938, RC. Wilson 530 (DAV, DS, POM, UC,
USFS). Orange Co.: along California Highway 74, 0.6 km E of San Juan Guard Station, Santa Ana Mountains, 3
May 1980, F.M. Roberts 98 (UCSB); Dana Point near Doheney Beach, 16 Jun 1965, Roos s.n. (UCSB). Riverside

216 PHYTOL O GTA volume 66(3):199-220 May 1989

Co.: E side of Indian Wash, Temescal Canyon, 19 Apr 1986, Boyd 1697 (HSC, NY, RSA, UCR); near Riverside,
29 May 1904, H.M. Hall 4964 (DS, PH, RM, UC); Whitewater, 11 May 1903, M.E. Jones s.n. (DS, MO, POM, US);
3 mi E of Valle Vista, 23 May 1988, Reveal 6865 (CAS, MARY, MO, RM, RSA, UTC). San Benito Co.: San
Carlos Creek, 13 May 1907, Jepson 2733 (JEPS). San Bernardino Co.: Barstow, 1 May 1932, Eastwood s.n. (CAS);
San Bernardino Valley, 2 Jun 1906, Parish 5777 (L, MIN, ORE, RM); Highland, 9 May 1919, Spencer 1110 (CAS,
GH, NY, POM). San Diego Co.: Point Loma, 7 May 1902, Brandegee 1623 (G, GH, K, LE, LL, MICH, MIN, NY,
POM, UC, US); near San Diego, Mar-Jun 1906, K. Brandegee s.n. (DS, MIN, UC, US); 4 mi E of Pala, 11 May
1930, J.T. Howell 4856a (CAS). San Joaquin Co.: Lathrop, 17 May 1896, Congdon s.n. (MIN). San Luis Obispo
Co.: ridge N of Nacimiento River, 3 Jun 1956, Hardham 849 (CAS); Black Lake Canyon, SE of Oceano, 5 May
1985, Keil 18755 (OBI, UCR); 1 mi S of San Miguel, 11 May 1937, Lee 933 (UC). Santa Barbara Co.: bottom of
La Cascada, N of Central Valley Road, 2 mi W of Stanton Ranch, Santa Cruz Island, 15 Jun 1969, M.R. Benedict
s.n. (SBBG); at California Highway 1 and Signature Road N of Lompoc, 15 Jun 1960, E.R Blakley 3483 (RSA,
SBBG); Cottonwood Canyon Wash at Bates Canyon Road crossing, 18 May 1965, Chandler 2282 (OKL, SBBG);
East Point, Santa Rosa Island, 9 Apr 1930, Munz & Hoffmann 11734 (GH, POM); N of Fox Canyon and E of Las
Positas Road, Santa Barbara, 24 May 1962, Pollard s.n. (ARIZ, DS, ENCB, MIN, NO, OKL, RSA, SBBG, SD,
TEX, UTC). Santa Cruz Co.: Pear Tree Ranch in lane leading to Isbell Grove, near Santa Cruz, 1887, C.L. Anderson
s.n. (UC). Sonoma Co.: Petaluma, 1880, Congdon s.n. (MIN). Stanislaus Co.: Gobin Ranch, 13 mi E of Waterford,
14 Apr 1935, Hoover 539 (NY, UC, US). Tulare Co.: 2.5 mi above Springville on North Tulare River, 8 May 1954,
Barneby & Howell 11373 (CAS). Unknown Co.: without location data, 1876, Parry & Lemmon 371 (BM, F, G, ISC,
MIN, NY, US, WIS). Ventura Co.: top of Piru Hill, 2 May 1931, Hoffmann s.n. (SBBG); floodplain of the Ventura

River N of and near Baldwin Road, 10 Apr 1970, Pollard s.n. (DAO, DS, RSA, SBBG). A total of 317 collections
examined.

When Gross (1913b) proposed the subsp. californica he cited four syntypes. He mentioned
a Parry collection without location data that was dated 1882, a Parish collection numbered
819 and dated Apr 1882, a Stokes collection from near San Diego dated Jun 1895 and a
collection from Monterey in John Ball’s herbarium at E. As a result of the herbarium studies,
the following probable syntypes were found:

Monterey Co.: Monterey, 18 Jul 1882, Pringle s.n. (E, F, G, LE, MIN, MO, NY, PENN, PH, US, VT, WS, WU).
San Bernardino Co.: mesas, San Bernardino, Mar 1881, Parish & Parish 819 (CM); plains, San Bernardino Valley,
Apr 1881, Parish & Parish 819 (F); mesas, San Bernardino Valley, Apr 1882, Parish & Parish 819 (BR, LE, MPU,
PH, US, WU); mesas, near San Bernardino, May 1882, Parish & Parish 819 (US); San Bernardino Valley, Apr
1884, Parish’ & Parish 819 (UC); San Bernardino Valley, Jun 1884, Parish & Parish 819 (DS); mesas, near San
Bernardino, Apr 1886, Parish & Parish 819 (MO, NY, US). San Diego Co.: hills near San Diego, Jun 1895, Stokes
s.n. (B). Unknown Co.: without location data, 1882, Parry s.n. (B, F, G, K, LE, NY, ORE, US).

The only collection from Monterey that was in the John Ball Herbarium is a Pringle collec-
tion. Although Pringle is not mentioned specifically, there is no question that this was the
collection Gross examined. Both the Stokes and Parry collections are at B and these sheets
were certainly examined by Gross. As for the Parish material, all of the material numbered
819 is cited. This is presented to indicate the kind of problem one often encounters when
working with Parish material in general.

The Parish Herbarium is now at DS and an examination of its holdings show that many
of the Parish collections found elsewhere are not represented. It is not known, for example,
if the various dates were actual dates the collections were made or the dates the specimens
were distributed to other herbaria. At DS, for example, the only specimen numbered 819
was collected in Jun 1884 and none of the remaining six collections found in this study bearing
number 819 is at DS. The collection specifically dated Apr 1882 is found mainly in European
herbaria (BR, LE, MPU and WU) with only a single sheet now in the United States (PH).

No Parish specimen with the date and number cited by Gross (1913b) was found at B
although the others examined by him are present. It is unknown if the sheet was destroyed
or is now misplaced. The Parry collection of 1882 is without location data, but two sheets
were found which were gathered at San Diego that were dated 1882 (MO, NEB). It is likely

Reveal: Genera related to Chonzanthe (Polygonaceae) 217

that all of these 1882 collections were taken at San Diego. Because Gross did not specifically
mention Pringle, and there is no Parish specimen at B, the Parry collection is selected here
as the lectotype and suggested to have been found in San Diego.

In the field, Lastarriaea coriacea is a remarkable plant. The stems readily disarticulate so
that at maturity a mere touch of the branches will cause the plant to crumble into pieces.
Because of the numerous recurved awns on the leaves, bracts, involucral bracts and even the
tepals, the individual fragments tend to hang together so that large pieces of the plant may
be carried away. This factor has certainly been a significant factor in the widespread distri-
bution of the species and genus.

The unusual chromosome numbers reported by Hardham (1989) indicate that the cytologi-
cal story of this species has yet to be fully examined.

2. Lastarriaea ptilota Rev., spec. nov.- TYPE: deflated mesa 24 km SE of Bahia Tortugas,
east of Bahia Thurloe, Baja California Sur, México, 150 m elev, 7 Mar 1985, Breedlove
62343 (holotype: MARY!; isotypes: CAS, RSA!).

A L. coriacea caulibus glabris cum bracteis lanceolatis, 0.5-2 cm longis et 1-2 mm latis,
involucris bractearum lanceolatis, 0.15-0.4 cm longis et 0.3-1 mm latis differt.

Plants 0.3-1 dm high and 0.5-2 dm across; /eaves basal, 0.5-2 cm long, 0.2-0.8 mm wide,
linear, sparsely hirsute along the margins; branches slender and brittle, weakly disarticulating,
mostly red, glabrous; bracts 2, 0.5-2 cm long, 1-2 mm wide, lanceolate, sparsely pubescent
mainly along the margins and midrib, spreading, the uncinate awns 0.2-0.6 mm long; in-
volucral bracts 0.15-0.4 cm long, 0.3-1 mm wide, lanceolate, sparsely pubescent, mostly spread-
ing, the uncinate awns 0.3-0.6 mm long; flowers 2-2.5 (3) mm long (including the awns), green
to greenish-white, thinly pubescent without, the tepals lanceolate, awn-tipped, united about
half of their length; stamens 3, the filaments 0.5-1 mm long, the anthers 0.2 mm long; achenes
1.5-2 mm long.

Rare and local on sandy to gravelly soils on Cedros Island and on the western Desierto el Vizcaino on the
adjacent mainland, Baja California, México, 150-500 ft elev, flowering from Mar-Apr.

Specimens Examined. - MEXICO. Basa CALIFORNIA Norte: 0.5 mi W of the village, Cedros Island, 16 Mar 1939,
Haines & Hale s.n. (LA, UC).

The recognition of the southern extreme as a new species is perhaps daring given the limited
material upon which it is based. The aspect of the plant is distinctive and when looking at
individuals one notes immediately the recurved or arched bracts that, as the specific name
implies, resemble the wings of seabirds in flight when viewed from a great distance. The
species is well isolated from the more northern populations of Lastarriaea coriacea which
does not extend as far south as the Desierto el Vizcaino.

The morphological features used to distinguish Lastarriaea ptilota from L. coriacea are not
all that significant at first glance. The differences in overall size of parts and the general lack
of hairs seems sufficient especially when compared with the features that are used to recog-
nize the Chilean species.

3. Lastarriaea chilensis Rémy in Gay, Fl. Chil. 5: 290. 1851-1852. Chorizanthe lastarriaea C.
Parry, Proc. Davenport Acad. Nat. Sci. 4: 63. 1884, nom. illeg. superfl.—- TYPE: without
location data or date, Gay s.n. (holotype: P!; isotypes: F, K!).

218 PHYTOLOG1IA volume 66(3):199-220 May 1989

Lastarnaea stricta Philippi ex C. Parry, Proc. Davenport Acad. Nat. Sci. 5: 36. 1886.— TYPE:
Coquimbo, Coquimbo, Chile, Sep 1885, Philippi s.n. (holotype: ISC!; isotypes: B, E, ISC,
SGO, WU!).

Lastariaea linearis Philippi ex C. Parry, Proc. Davenport Acad. Nat. Sci. 5: 36. 1886.—TYPE:
Coquimbo, Coquimbo, Chile, 18 Sep 1885, Philippi s.n. (holotype: ISC!; isotypes: B,
SGO!).

Plants 0.2-0.5 (1.2) dm high and 0.5-1 dm across; /eaves basal and sheathing up the stem,
0.5-1 (3) cm long, 0.2-0.4 mm wide, linear, hirsute along the margins; branches slender and
brittle, readily disarticulating, yellowish-green to green, sparsely pubescent with curled hairs;
bracts 2, 0.2-0.6 (1.5) cm long, 0.2-1.5 (3) mm wide, broadly lanceolate to narrowly ovate,
sparsely hirsute, erect or nearly so, the uncinate awns 0.2-0.6 mm long; involucral bracts
0.2-0.5 (1.5) cm long, (0.5) 0.8-2 (2.5) mm wide, broadly lanceolate to ovate, sparsely hirsute,
erect, the uncinate awns 0.2-0.5 mm long; flowers 2-3.5 mm long (including the awns), light
green to greenish-white, thinly pubescent without, the tepals narrowly lanceolate, awn-tipped,
united about half of their length; stamens 3, the filaments 0.5-1 mm long, the anthers 0.2
mm long; achenes 2.5-3 mm long.

Infrequent to locally common on sandy to gravelly soils along the coast and in the foothills of the mountains
of northern and central Chile, from 100-4600 ft elev; flowering from Sep-Dec.

Representative Specimens. - CHILE. ACONCAGUA: Valle de Marga-Marga, SE of Valparaiso, 1931, Jaffuel & Pirion
3112 (GH). ANTOFAGASTA: 2 km S of Chincolco on road to Fundo “El Sobrante,” 9 Nov 1970, Simon 292 (RSA).
CoLcHAGuA: 15 km W of Alcones on road to Pichilemu, Oct 1973, Stebbins 8636 (UC). Coquimso: Illapel, Oct
1898, Geisse s.n. (SGO, Z); Fray Jorge, 26 Sep 1935, Mufioz 110 (GH, SGO); Corral de Julio, Exclusion C. Jiles,
7 Nov 1976, Muroz-S. 977 (SGO); Vallenar, 13 Oct 1914, Rose & Rose 19332 (NY, US); Pichidangui, 14 Nov 1976,
Weber & Johnston 966 (COLO); La Serena-Vallenar Road, Cuesta Buenos Aires, 50 km N of La Serena, 1 Nov
1938, Worth & Morrison 16290 (GH, UC); dunes of Tongoy, 19 Sep 1975, Zéliner 8277 (MO, NY). SANTIAGO:
Santiago, 1855, Germain s.n. (BM, G, K); Valle del Clarillo, Nov 1933, Grandjot s.n. (MO); Santiago, without date,
Hohenocker 616 (BM, G, GOET, K, LE, TCD); La Obec, 20 Sep 1927, Montero 478 (F); 8 km W of Tiltil on E
slope of Cuesta de la Dormida, 9 Nov 1976, Weber & Johnston 762 (BRY, COLO, NY, TEX). UNKNown: without
location data, without date, Bertero 228 (B, F, G, GH, NY); without location data, without date, Gillies s.n. (GH,
ISC, NY); Concon, without date, Poeppig 50 (B, BR, K, LE). VALPARAISO: Quillota, Oct-Nov 1829, Bertero 959 (F,
G, GH, LE, NY); Cuesta de Zapata, between Curacavi and Casablanca, 3 Nov 1948, Killip & Pisano 39696 (US);
dunes of Cochoa near Vina del Mar, 21 Oct 1982, Zoliner 11356 (MO). A total of 62 collections examined.

Past workers have long associated the North and South American populations of Lastarriaea
under a single name. Both Gross (1913a) and Goodman (1934) considered the two to be
representatives of the same species, but at that time it was felt that the species had been
introduced into North America. That was not true, and when Goodman (1943) was finally
able to review a large suite of specimens from Chile, he concluded the two were distinct
species. As a result of the present review, there is no question that the two are distinct species
and each is unique to its own range. The dispersal pattern was from north to south and
occurred about the same time during the Pleistocene that the ancestral populations of
Chonzanthe commissuralis arfived in Chile from western North America. As both of the
Chilean species are well adapted for long distance dispersal (as is Oxytheca dendroidea which
is also in South America; Ertter 1980), it seems reasonable to conclude that birds were the
probable agent although migrating mammals cannot be discounted.

Reveal: Genera related to Chonizanthe (Polygonaceae) 219

ACKNOWLEDGMENTS

I wish to thank the curators and staff of the following herbaria for loans or for arranging
access to their collections during visits: A, ARIZ, ASC, ASU, B, BKL, BM, BR, BRY, CAN,
CAS. CGE, CHSC, CM. COLO; CPH; CS, DAG; DAY, DS, E, ENCEAE; G,.GBy GH,
GOET, HSC, ID, IDS, ILL, IND, ISC, JEPS, K, KANS, KSC, L, LL, M, MARY, MICH,
MIN, MNA, MO, MONTU, MPU, MSC, ND, NDG, NEB, NESH, NMC, NO, NTS, NY,
OBI, OKL, OKLA, ORE, OSC, OXF, P, PENN, PH, POM, RENO, RM, RSA, SBBG, SD,
SFSU, TCD, TEX, UC, UCR, UCSB, UNLV, US, USFS, UT, UTC, VT, W, WILLU, WIS,
WS, WTU, WU, and Z. Individuals wishing a complete list of herbarium specimens examined
may request a copy. Funding for field work has been made possible through a variety of
sources. This paper was reviewed by Dr. Edward E. Terrell and Clare B. Hardham. This is
Scientific Article A-4937, Contribution 7980, Maryland Agricultural Experiment Station.

LITERATURE CITED

Abrams, L. 1944. Illustrated flora of the Pacific States. Vol. 2. Stanford Univ. Press, Stanford.
Bentham, G. 1836. On the Eriogoneae, a tribe of the order Polygonaceae. Trans. Linn. Soc. London 17: 401-420.

. 1856. “Eriogoneae,”’ pp. 5-28. In: A. de Candolle (edit.), Prodromus systematis naturalis regni
vegetabilis. Vol. 14. Victor Masson, Paris.

Curran, M. K. 1895. Botanical notes. Bull. Calif. Acad. Sci. 1: 272-275.

Dammer, U. 1892. Polygonaceae. Nat. Pflanzenfam. 3(1a): 1-36.

Dumortier, B. C. J. 1829. Analyse des familles des plantes. J. Casterman, Tournay.

Ertter, B. J. 1980. A revision of the genus Oxytheca Nutt. (Polygonaceae). Brittonia 32: 70-102.
Goodding, L. N. 1904. Southwestern plants. Bot. Gaz. 37: 53-59.

Goodman, G. J. 1934. A revision of the North American species of the genus Chorizanthe. Ann. Missouri Bot.
Gard. 21: 1-102.

. 1943. Notes on Chorizanthe. Leafl. W. Bot. 3: 230.
. 1957. The genus Centrostegia, tribe Eriogoneae. Leafl. W. Bot. 8: 125-128.

Gray, A. 1859. List of a collection of dried plants made by L. J. Xantus, at Fort Tejon, and vicinity, California,
near Lat. 35°, and Long. 119,° 1857-8. Proc. Boston Soc. Nat. Hist. 7: 145-149.

Greuter, W. (edit.). 1988. International code of botanical nomenclature. Regnum Veget. 118: 1-328.
Gross, H. 1913a. Beitrage zur Kenntnis der Polygonaceen. Bot. Jahrb. Syst. 49: 234-339.
. 1913b. Polygonaceae nonullae novae. Bot. Jahrb. Syst. 49: 340-348.

Hardham, C. B. 1989. Chromosome numbers of some annual species of Chorizanthe and related genera (Polygon-
aceae: Eriogonoideae). Phytologia 66: 89-94.

Heller, A. A. 1905. Botanical exploration in California, season of 1905. Muhlenbergia 2: 1-176.
. 1910. Catalogue of North American plants north of Mexico. Ed. 3. Published by the author, Lancaster.
Hoover, R. F. 1966. Miscellaneous new names for California plants. Leafl. W. Bot. 10: 337-350.
Jaretzky, R. 1925. Beitrage zur systematik der Polygonaceae unter beriicksichtigung des oxymethyi-anthrachinon-
vorkommens. Repert. Spec. Nov. Regni Veg. 22: 49-83.
Jepson, W. L. 1913. Polygonaceae. Fl. Calif. 4: 376-428.
. 1923. Manual of the flowering plants of California. Student Store, Berkeley.
Jones, M. E. 1908. Contribution to western botany—no. 12. Contr. W. Bot. 12: 1-100.
Macbride, J. F. 1918. New or otherwise interesting plants, mostly North American Liliaceae and Chenopodiaceae.
Contr. Gray Herb. 53: 1-22.

McKelvey, S. D. 1955. Botanical explorations in the trans-Mississippi west, 1790-1850. Arnold Arboretum, Jamaica
Plains.

Meisner, C. F. 1856. “Polygonaceae,” pp. 1-5, 29-185. In: A. de Candolle (edit.), Prodromus systematis naturalis
regni vegetabilis. Vol. 14. Victor Masson, Paris.
Melchior, H. 1964. A. Engler’s syllabus der Pflanzenfamilien. Gebriider Borntraeger, Berlin.
Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.
. 1974. A flora of southern California. Univ. California Press, Berkeley.

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Parry, C. C. 1884. Chorizanthe RK. Brown. Revision of the genus, and arrangement of the annual species—with one
exception, all North American. Proc. Davenport Acad. Nat. Sci. 4: 45-63.
- 1885. Notes on Chorizanthe lastarriwa, Parry. W.-Amer. Sci. 1: 29-31.
- 1886. Lastarriaa, Rémy. Confirmation of the genus, with characters extended. Proc. Davenport Acad.
Nat. Sci. 5: 35-36.
. 1889. Chorizanthe, R. Brown. Review of certain species heretofore improperly characterized or wrongly
referred; with two new species. Proc. Davenport Acad. Nat. Sci. 5: 174-184.
Rémy, J. 1851-1852. “Poligoneas,” pp. 263-293. Jn: C. Gay, Historia fisca y politica de Chile ... BotAnica. Vol. 5,
part 3. E. Thunot y Ca., Paris.
Reveal, J. L. 1978. Distribution and phylogeny of Eriogonoideae (Polygonaceae). Great Basin Naturalist Mem. 2:
169-190.
. 1989a. A review of the genus Harfordia (Polygonaceae: Eriogonoideae). Phytologia 66: 221-227.
. 1989b. Remarks on the genus Prerostegia (Polygonaceae: Eriogonoideae). Phytologia 66: 228-235.
. 1989c. Notes on selected genera related to Eriogonum (Polygonaceae: Eriogonoideae). Phytologia 66:
236-245.
& B. J. Ertter. 1977. Goodmania (Polygonaceae), a new genus from California. Brittonia 28: 427-429.
- 1980. The genus Nemacaulis (Polygonaceae). Madrofo 27: 101-109.
& C. B. Hardham. 1989a. Three new monospecific genera of Polygonaceae subfamily Eriogonoideae
from California. Phytologia 66: 83-88.
. 1989b. A revision of the annual species of Chorizanthe (Polygonaceae: Eriogonoideae). Phytologia
66: 98-198.
& J. T. Howell. 1976. Dedeckera (Polygonaceae), a new genus from California. Brittonia 28: 245-251.
Roberty, C. & S. Vautier. 1964. Les genres de Polygonacees. Boissiera 10: 7-128.
Shields, O. & J. L. Reveal. 1988. Sequential evolution of Euphilotes (Lycaenidae: Scolitantidini) on their plant host
Eriogonum (Polygonaceae: Eriogonoideae). Biol. J. Linn. Soc. 33: 51-93.
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Press, San Francisco.
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Walpers, G. G. 1852. Ordo CLXXIV. Eriogonaceae Wiprs. Ann. Bot. Syst. 3: 297.
Watson, S. 1877. Descriptions of new species of plants, with revisions of Lychnis, Eriogonum and Chorizanthe. Proc.
Amer. Acad. Arts 12: 246-278. f
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Wiggins, I. L. 1980. Flora of Baja California. Stanford Univ. Press, Stanford.

Phytologia (May 1989) 66(3):221-227

A REVIEW OF THE GENUS HARFORDIA
(POLYGONACEAE: ERIOGONOIDEAE)

JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

ABSTRACT

The genus Harfordia has been generally defined to consist of two species, H. macroptera of the
mainland and the Cedros Island endemic, H. fruticosa. The revised genus is now considered to
be monospecific but composed of three varieties: the var. miacroptera restricted mainly to the
Magdalena Bay region of Baja California Sur, the insular var. fruticosa and the more northern
and common var. galioides (n= 20) mainly in Baja California Norte, México.

KEY WORDS: Polygonaceae, taxonomy, Harfordia, México.

INTRODUCTION

The genus Harfordia was proposed by Greene and Parry in an article published by Parry
(1886) shortly after Greene (1885) had recognized Pterostegia galioides and P. fruticosa as
distinct from the previously described P. macroptera (Bentham 1844). At the time, Pterostegia
was defined as a genus of Polygonaceae Juss. (tribe Pterostegieae Torr. & A. Gray) charac-
terized by a remarkably winged involucre surrounding the achene. When Bentham proposed
P. macroptera he described the species as a “much stronger and stiffer plant” than the type
of the genus, P. drymarioides, but it is obvious he was uncertain if his new species was an
annual (like the type) or a perennial. The realization that P. macroptera and its related species
in Baja California were shrubby perennials and that the large, bladdery inflated involucral
bracts were unique, Greene and Parry decided to establish Harfordia and restrict it to just
two species, H. macroptera of the mainland and the Cedros Island endemic H. fruticosa.

Harfordia has been generally defined since 1886 to consist of two species. When Wiggins
(1964) prepared a flora of the Sonoran Desert, he accounted for only H. macroptera, citing
Greene’s Pterostegia galioides in synonymy. At that time he did not have to consider the
species endemic to Cedros Island and therefore did not mention H. fnuticosa. It is suggested
that his failure to dispose of H. fruticosa when he wrote the flora of Baja California (Wiggins
1980) was simply an oversight.

While reviewing Harfordia (Polygonaceae Juss. subf. Eriogonoideae Meisner), it became
obvious that the three entities recognized by Greene were distinct. The type of the genus,
H. macroptera, is a low, spreading perennial that generally forms distinct shrubs with broad
leaves. The Cedros Island endemic, H. fruticosa is most closely related to var. macroptera
but is a more sprawling shrub with shorter and broader leaves. To the west and north is a
larger and an even more sprawling shrub; it was this expression readily distinguished by its
narrow leaves that Greene (1885) called Prerostegia galioides. When Harfordia was proposed,
P. galioides was submerged into H. macroptera. These expressions do not overlap except on

221

222 PHYTOL OG1A volume 66(3):221-227 May 1989

the extreme western tip of the Desierto el Vizcafno in northwestern Baja California Sur.
Here the distinction between all three is marred by the existence of a series of morphological
intermediates. For this reason, the formerly bispecific genus is now considered to be
monospecific with three varieties.

TAXONOMY

Harfordia E. Greene & C. Parry in C. Parry, Proc. Davenport Acad. Nat. Sci. 5: 26. 1886.
Polygonaceae Juss. subtribe Harfordiineae H. Gross, Bot. Jahrb. Syst. 49: 329. 1913.
Pterostegia Fischer & C. Meyer sect. Harfordia (E. Greene & C. Parry in C. Parry)
Roberty & Vautier, Boissiera 10: 108. 1964.- LECTOTYPE: Harfordia macroptera (Benth.
in Hinds) E. Greene & Parry in Parry, selected here.

Sprawling to spreading, suffruticose to shrubby, sparsely pubescent, monoecious or more
commonly dioecious woody perennials with numerous slender to stout stems arising from a
woody taproot; /eaves cauline, opposite, exstipulate, short petiolate, the blades linear to
oblanceolate or elliptic to obovate-spatulate, entire, glabrous or pubescent with slightly curled
hairs, thin to thickish and more or less succulent, the petioles basally connate at the slightly
swollen nodes; flowering stems slender to thick, the new growth arising from the axils of
previous year’s leaves, dichotomously branched, green to red or gray, often densely pubescent
with appressed curly hairs, becoming glabrous with age and bearing an exfoliating grayish
bark; branches numerous, dichotomously branched throughout; inflorescences cymose with
the flowers arranged in an axillary cluster or, on perfect or female plants, occasionally on
short, stout peduncles; bracts lacking; peduncles when present, stout, erect, usually densely
pubescent, restricted to perfect or female plants, arising from the lower nodes in the upper
part of the inflorescence; involucres composed of a highly modified involucral bract, this
situated lateral of and basal to a slender to stout pedicel, surrounding but basal to the im-
mature female or perfect flower with the immature modified sheath of the bract reflexed
until after fertilization then elongating and surrounding the achene, the lateral saccate wings
basal to the immature sheath, gradually inflating with age, and the whole structure greatly
enlarging and becoming laterally bilobed, gibbously bisaccate and accrescent in fruit, hyaline
and reticulated with deep red to red-purple veins, glabrous; flowers imperfect or perfect,
sparsely pubescent along the midribs and base without, glabrous within, the tepals 6, petaloid,
essentially monomorphic, lanceolate, united about one-third of their length; male flowers 1-5
per node, pale yellow, smaller than the female ones, occasionally with an aborted, imbedded
ovary; female flowers 1-2 (3) per node, initially pale yellow but becoming red to rose, always
with aborted stamens; perfect flowers mostly 2 per node, pale yellow to rose; stamens 9, those
of the male flower slightly exserted on elongated, glabrous filaments bearing yellow oval
anthers, those of the female flower on minute, glabrous filaments bearing minute aborted
anthers; achenes strongly 3-angled, yellowish-brown to brown, glabrous, the narrow base
tapering to a slightly winged apex, the embryo straight but with an excentric radicle, in abun-
dant endosperm.

A monospecific genus with three essentially allopatric varieties in western Baja California, México, from San
Vicente south to Magdalena Bay, and on the adjacent off-shore islands.

The remarkably altered involucral bract of the genus Harfordia is the one feature that almost
immediately catches the attention of even the most causal collector. This large, bisaccate and
bladdery inflated structure with deep red to red-purple veins is unmistakable. The sprawling

Reveal: Review of Harfordia (Polygonaceae) 223

plants are often covered with these colorful bracts.

The bract undergoes a scries of developmental changes. Initially the immature flower is
atop a short, slender to stout pedicel with the immature bract subtending the pedicel. The
bract is initially reflexed and composed of three individual segments joined only at the point
of attachment on the top of the peduncle but at the base of the pedicel. As the bract matures
the anterior segment expands from the base to form a single sheath surrounding the pedicel
with the two lateral edges expanding so that they nearly touch. The two lateral laminar regions
of the bract near the base of the sheath soon begin to swell between the single dorsal vein
and the two lateral veins near the margins. The two lateral flaplike edges beyond the lateral
veins begin to extend outwardly so as to produce the two distinctive thin flaps on each side
of the pedicel and maturing flower. At this point the flower is still well exserted but once
fertilization has occurred in the perfect or female flower, the two lateral laminar regions of
the involucral bract rapidly expand both outwardly and upwardly.

When the two laminar expansions of the bract differentiate and begin to form the precur-
sors of the inflated bladders, the inner lateral flaps around the pedicel begin to elongate so
that as the fruit expands, these central flaps soon enclose the entire achene. The flaps,
however, remain free of the pedicellate flower and fruit at the base of the involucral bract.
Since the pedicel never elongates, while the lateral flaps soon overtop it and extend far beyond
the fruit, the whole structure is now completely overtopping and encompassing the achene.
The two laminar expansions continue to expand resulting in the characteristic enlarged, in-
flated and bladdery lateral portion of the involucral bract.

This modified involucral bract is certainly an aid in the dispersal of the achene. The inflated
papery bract with its enclosed achene is easily detached from the parent plant and blown in
the wind. In such instances the fully mature achene is readily transported to new areas. As
the bract dries, it tends to deflate; but because it is vein-filled, the bract does not disintegrate
rapidly. After a rain, the bract reinflates and in the winds associated with such convection
storms, the inflated bracts can be moved once again. It appears that the bracts and their
associated achenes can remain associated two to three years before the achene is finally
liberated.

An aspect of dispersal that was considered but never observed is the possibility that birds
might be an effective agent. The achene is often nearly full size before the involucral bract
begans to enlarge significantly to obscure the fruit. In an immature state, the bract is typically

a bright red-purple to blood red, in short an attractive “berry” that just might be taken by
birds.

1. Harfordia macroptera (Benth. in Hinds) E. Greene & C. Parry in C. Parry

Dioecious or infrequently monoecious sprawling perennial shrubs or subshrubs 2-6 (10)
dm high and 3-8 (14) dm across, dichotomously branched throughout, green to red or gray,
the new growth often densely pubescent with appressed curled hairs, becoming glabrous with
age; /eaves opposite, linear to oblanceolate or elliptic to obovate-spatulate, 0.2-2 cm long,
0.5-6 mm wide, glabrous or pubescent with slightly curled hairs, thin to thickish and more
or less succulent; inflorescences cymose with flowers in axillary clusters or in perfect or female
plants on stout peduncles; peduncles stout, erect, 0.7-2 cm long, usually densely pubescent;
involucres lacking in male plants, those of perfect or female plants reduced to a highly
modified involucral bract, this up to 3 cm across, laterally bilobed, gibbously bisaccate and
accrescent at maturity, hyaline and reticulated with deep red to red-purple veins, glabrous;
flowers imperfect or perfect, sparsely pubescent without, the tepals monomorphic, lanceolate,

224 PHYTOLOG1IA volume 66(3):221-227 May 1989

the male ones 1-5 per node, pale yellow, 1.2-1.7 mm long, the perfect and female ones 1-2
(3) per node, pale yellow to rose or red, 0.5-2 mm long; stamens 9, mostly exserted, the
filaments 0.8-1 mm long, glabrous, the anthers 0.2-0.3 mm long, oval, yellow; achenes
yellowish-brown to brown, 3-5 mm long, the narrowly globose base tapering to a narrowly
winged, strongly 3-angled beak.

Local and often common in sandy to gravelly soil on the western coast of Baja California from San Vicente to
Magdalena Bay and on Cedros Island, associated with scattered shrubs from 15-2200 ft elev, flowering from Oct-
Apr.

Key to the Varieties

A. Leaves linear to narrowly oblanceolate, 0.5-2 (2.5) mm wide, (0.4) 0.8-2 cm long, pubescent; involucral bracts
2-3 mm wide at maturity; western coast of Baja California from San Vicente to the Desierto el Vizcaino. ..
Per asic ea fer tacuccee sot eacooncatonte ssunteres servants sua ncetaeraesho seen rstov usosenearserr=deecenes erect tspecnete renee aenteeeee teen la. var. galioides

AA. Leaves broadly oblanceolate to spatulate, 2-6 mm wide, 0.2-1.5 cm long, glabrous; involucral bracts 1-2 cm
wide at maturity; western coast of Baja California and adjacent islands from Desierto el Vizcaino south to
Magdalena Bay.
B: Leaves: 2-4immiwide; ‘Cedrosi Island -ciesc55.c0:sccccva-scvecee- asses svoa0eacesnancou-versctadeesaeneee coerce oe 1b. var. fruticosa
BBs Weavesi2-Gimim wide Mapdalenian Way. Wernecvccsesenescepecescxet-tocrsncaccraspuce concerts cates sucertetenreecteneae lc. var. macroptera

As here defined Harfordia macroptera is divided into three morphologically weakly defined,
yet geographically distinct varieties.

1a. Harfordia macroptera (Benth. in Hinds) E. Greene & C. Parry in C. Parry var. galioides
(E. Greene) Rev., comb. et stat. nov. Pterostegia galioides E. Greene, Bull. Calif. Acad.
Sci. 1: 213. 1885.-TyYPE: on the bluffs of Cape San Quintin, Baja California Norte,
México, 10 May 1885, Greene s.n. (lectotype: US!; duplicates of the lectotype: UC!,
selected here).

Large arid often sprawling shrubs mostly 3-6 (10) dm high and 4-8 (14) dm across; leaves
linear to narrowly oblanceolate, (0.4) 0.8-2 cm long, 0.5-2 (2.5) mm wide, pubescent with
slightly curled hairs, thin; involucral bracts 2-3 cm across at maturity; n= 20.

Local and common in sandy to gravelly soil on the western coast of central Baja California from San Vicente
to Desierto el Vizcaino, from 15-2200 feet elev, flowering from Oct-Apr.

Representative Specimens. - MEXICO. BAJA CALIFORNIA Norte: 2 mi N of San Sim6n, near San Quintin Bay,
22 Mar 1949, Bacigalupi 3036 (DS, JEPS, RSA, UC); San Borja, 1889, Brandegee s.n. (GH, PH, UC, US); near
Rio del Rosario, 11 km E of El Rosario, 6 Nov 1947, Carter et al. 1863 (DS, F, GH, K, MEXU, US); NW slope
of Las Amarillas, 6 mi N of the American Smelter Company mine, 4 Feb 1935, Chisaki & Bell 518 (CAS, COLO,
DS, F, GH, K, MEXU, MICH, MO, NY, OKL, RM, SD, SMU, UC, US, UTC, W, WS); 15 mi SW of Punta
Prieta, 10 Feb 1935, Epling & Robison s.n. (DS, GH, LA, MICH, NY, UC); S of San Augustin, 1 Feb 1935, Epling
& Robison s.n. (DS, F, GH, K, LA, MICH, POM, RM, SMU, UC); canyon in the foothills of Sierra Lino, 25 mi
S of Punta Prieta, 6 Mar 1947, Gentry 7343 (ARIZ, ASU, DS, MICH, RSA, SD, UC, US); 13 mi E of El Rosario,
30 Nov 1957, Klein et al. 3 (DS, GH, RSA, UC, UT); 2 mi S of Rancho Ibarra, 23 Mar 1970, Moran 16821 (ARIZ,
ENCB, LL, MICH, MSC, OKL, SD); N slope of Cerro San Juan de Dios, 1 May 1973, Moran 20692 (MO, MSC,
POM, SD, UC); 2 mi N of San Antonio del Mar, 19 May 1975, Moran 22012 (DS, PH, SD); San Telmo, Apr 1886,
Orcua 1374 (B, DS, F, G, GH, ISC, K, LE, MIN, MO, NDG, NY, PH, UC, US, VT); San Quintin Bay, Jan 1889,
Palmer 696 (ARIZ, F, GH, K, LE, LL, MICH, NY, PH, UC, US); 14 km NW of Colonia Guerrero, 4 Apr 1958,
Raven et al. 12205 (CAS, DS, GH, K, LA, SMU, UC); Arroyo Socorro, 20 May 1988, Reveal 6829 (BM, BR, BRY,
CAS, CLEM, G, K, KW, LE, MARY, MEXU, MO, NY, OKL, OSC, RM, RSA, SMU, US, UTC, VT, WIS, Z);
Jaraguay Grade, 23 Mar 1988, Reveal et al. 6728 (CAS, MARY, MO, NY, RSA, US); Cafiada las Palomas, 24 Mar
1988, Reveal et al. 6754 (CAS, MARY, MO, NY, RSA); near Hamilton Ranch, 3 Feb 1935, Shreve 6837 (ARIZ,

Reveal: Review of Harfordia (Polygonaceac) 225

F, GH, MICH, MO); 9.8 mi N of Colonia Guerrero, 25 Mar 1954, Straw & Ownbey 527 (COLO, MEXU, RSA);
near Aguajito Ranch, 18 mi from El Rosario, 23 Feb 1963, Thorne & Henrickson 32545 (MICH, RSA, Z); 10 mi
S of El Rosario, 9 Mar 1983, West et al. 383-52 (ARIZ, COLO, OBI, SD, TEX, UCR); between San Telmo and
the road to San Quintin, 1 Mar 1930, Wiggins 4282 (ARIZ, CAS, DS, GH, K, MICH, NO, NY, POM, SMU, UC,
US, WTU); Playa Santa Catarina, 10 Mar 1930, Wiggins 4459 (CAS, DS, GH, LA, MICH, NY, POM, UC, US);
near San Vicente, 5 Apr 1950, F. Wylie s.n. (SD). BasA CALIFORNIA Sur: Las Huevitas, 1889, Brandegee s.n. (GH,
PH, UC, US); Rinconado de San Hipdélito, Vizcaino Desert, 15-17 Nov 1947, Gentry 7804 (ARIZ, DS, MICH,
RSA, UC); Arroyo del Portezuelo, 9.5 mi S of San Jose de Castro, 5 Feb 1973, Moran & Reveal 19818 (SD, US);
at the pass at the head of Arroyo Largo, 8 Feb 1973, Moran & Reveal 19948 (ARIZ, ENCB, RSA, SD, US). A
total of 110 collections were examined.

The var. galioides is the more common expression of the species, being found along the west-
ern coast of peninsular Baja California from the San Vicente area just below Ensenada south-
ward on the coastal mesas and in the adjacent mountains to the Desierto el Vizcaino of
northwestern Baja California Sur. Here the generally pubescent, long, narrow leaved phase
of var. galioides gives way to a slightly broader phase that approaches that seen in var. mac-
roptera.

Plants of var. galioides are widely scattered and often not obvious except when the inflated
involucral bracts are fully expanded and numerous. The size of the individual is often de-
pendent upon what species of plants the var. galioides is able to sprawl over. When not on
another plant, most individuals are less than 5 dm high, and if subjected to onshore winds,
the plants can be less than 1 dm high. The numerous, thin branches make for intricate and
highly branched individuals. The flowering portions tend to be on the exterior of the shrub
so that below this level the plant can seem to be a never ending array of red to grayish twigs.

At present, only the var. galioides has been counted. The voucher for the number, n= 20,
is Reveal et al. 6728. The count is the first for the genus.

1b. Harfordia macroptera (Benth. in Hinds) E. Greene & C. Parry in C. Parry var. fruticosa
(E. Greene) Rev., comb. et stat. nov. Pterostegia fruticosa E. Greene, Bull. Calif. Acad.
Sci. 1: 212. 1885. Harfordia fruticosa (E. Greene) E. Greene & C. Parry in C. Parry,
Proc. Davenport Acad. Nat. Sci. 5: 28. 1886.-TyPE: Cedros Island, Baja California
Norte, México, 1 May 1885, Greene s.n. (lectotype: US!; duplicates of the lectotype: ISC,
MIN, MO, UC!, selected here).

Large sprawling shrubs mostly 1-3 (6) dm high and 2-4 (8) dm across; /eaves broadly
oblanceolate to obovate-spatulate, 0.2-0.8 (1) cm long, 2-4 mm wide, essentially glabrous,
thick and succulent; involucral bracts 1-2 cm across at maturity.

Local and common in gravelly soil on Cedros Island, Baja California Norte, México, from 15-200 feet elev,
flowering from Dec-Mar.

Representative Specimens. - MEXICO. BAJA CALIFORNIA Norte: all Cedros Island: Mar-Jun 1897, Anthony 285
(BM, COLO, DS, E, F, GH, K, MIN, NESH, POM, SD, UC, US, VT); 2 mi S of the lighthouse on the E side,
23 Feb 1977, C. Davidson 5487 (F, NY, RSA, SD); canyon in San Agustin Mountains, 22 Feb 1939, Haines & Hale
s.n. (ARIZ, CAS, COLO, K, LA, LL, MICH, MO, NY, POM, SD, SMU, UC, US, WTU); South Bay, 17 Aug
1932, J.T. Howell 10704 (CAS, DS, F, GH, POM, US); arroyo in middle of E coast, 16 Apr 1963, Moran 10683
(DS, RSA, SD, UC); canyon on E side, 4 mi from N end, 25 Jun 1968, Moran 15148 (MICH, RSA, SD, UC); pine
ridge at the head of Gran Cafion, 18 Jun 1960, C.H. Muller 10805 (SBBG, SD, UCSB); 18-20 Mar 1889, Palmer
704 (ARIZ, F, GH, K, MEXU, MICH, NY, UC, US, VT); Punta Prieta, 28 Nov 1976, Philbrick & Benedict B7674
(SBBG); 10 Mar 1911, Rose 16114 (NY, US). A total of 28 collections were examined.

The isolated nature of the var. fniticosa has certainly contributed to the evolution of this

226 PHYTOLOG1I1A volume 66(3):221-227 May 1989

expression. It is a more compact phase of the species compared to the southern var. mac-
roptera.

The recent discovery of several of the Cedros Island endemics on the mainland indicates
that the previously assumed marked isolation of that island is not as significant as once
thought. The weak morphological distinction between the var. fruticosa and the var. macro-
ptera is probably a manifestation of both the recentness of the isolation of Cedros as well as
the possibility that even the minor differences are due more to the insular habitat than to
any real biological difference. Additional collections of Harfordia from the Bahia de Tortugas
region would be useful. The mainland plants are all from a relative restricted area in the
mountains east of Bahia de Tortugas. Further study of these plants may reveal that they
might better be referred to var. fruticosa.

The accepted date the var. fruticosa was collected is 1 May 1885 as this is the date com-
monly found on specimens. The holotype at US, however, is dated 30 Apr 1885. This is
regarded as an error and all the duplicates cited above are considered to be isotypes.

1c. Harfordia macroptera (Benth. in Hinds) E. Greene & C. Parry in C. Parry, Proc. Daven-
port Acad. Nat. Sci. 5: 28. 1886, var. macroptera. Pterostegia macroptera Benth. in Hinds,
Bot. Voy. Sulphur 44. 1844.- LECTOTYPE: Magdalena Bay, Baja California Sur, México,
Nov 1839, Hinds s.n. (lectotype: K!; duplicates of the lectotype: BM, GH!, selected here).

Large sprawling shrubs mostly 2-4 (6) dm high and 4-6 (8) dm across; leaves broadly
oblanceolate to spatulate, 0.5-1.5 cm long, 2-6 mm wide, essentially glabrous, thick and suc-
culent; involucral bracts 1-2 cm across at maturity.

Local and common on gravelly soil in the Magdalena Bay region, Baja California Sur, from 15-200 feet elev;
flowering from Dec-Mar.

Specimens Examined. - MEXICO. BasA CALIFORNIA Sur: Magdalena Bay, Oct-Nov 1839, Barclay 3083 (BM, K);
Magdalena Bay, 12 Jan 1889, Brandegee s.n. (DS, GH, NY, PH, RSA, UC, US); Santa Magdalena Island, Cabo
San Lazaro, 26 Apr 1973, C. Davidson 2044 (RSA); Magdalena Bay, without date, Lung 29 (UC); Santa Maria
Bay, 30 Mar 1952, Moran 3544 (DS); hill near Santa Maria Lagoon, Magdalena Island, 20 Apr 1979, Nakai 480
(CAS, MEXU); Margarita Island, 29 Nov 1905, Nelson & Goldman 7301 (GH, US); Magdalena Island, Mar 1917,
Orcutt 16 (NY, US).

On the voyage of H.M.S. Sulphur were two major plant collectors, Richard Brinsley Hinds,
ship’s surgeon, and George Barclay, botanist sent by the Royal Botanic Garden at Kew. The
ship traveled southward from San Diego arriving at Magdalena Bay on 2 November. Here
the collectors were greeted by a variety of plants in full flower and they made the best of
the some three weeks the ship remained at the Bay (Lindsay 1955). Bentham (1844) had
access to both the Hinds and Barclay collections (McKelvey 1955) so it is not always possible
to determine exactly upon whose specimens he established his names. In this instance a
lectotype has been selected on that element that is most in agreement with Bentham’s descrip-
tion.

No specimens of var. macroptera have yet been found on the mainland. A lack of botanical
activity is likely the cause of this rather than a natural restriction of the taxon to Isla Santa
Margarita and Isla Magdalena.

ACKNOWLEDGMENTS

I wish to thank the curators and staff of the following herbaria for loans or for arranging

Reveal: Review of Harfordia (Polygonaccac) 227

access to their collections during visits: A, ARIZ, ASC, ASU, B, BKL, BM, BR, BRY, CAN,
CAS, CGE, CHSC, CM, COLO, CPH, CS, DAO, DAV, DS, E, ENCB, F, G, GB, GH,
GOET, HSC, ID, IDS, ILL, IND, ISC, JEPS, K, KANS, KSC, L, LL, M, MARY, MICH,
MIN, MNA, MO, MONTU, MPU, MSC, ND, NDG, NEB, NESH, NMC, NO, NTS, NY,
OBI, OKL, OKLA, ORE, OSC, OXF, P, PENN, PH, POM, RENO, RM, RSA, SBBG, SD,
SFSU; TED; TEX, UC, UGR, UCSB; UNEV; US, USES, UT, UTIG VT, W; WIELU; WIS;
WS, WTU, WU, and Z. Individuals wishing a complete list of herbarium specimens examined
may request a copy. Funding for field work has been made possible through a variety of
sources. I appreciate the comments of Dr. Edward E. Terrell and Clare B. Hardham who

reviewed the article. This is Scientific Article A-4938, Contribution 7981, Maryland Agricul-
tural Experiment Station.

LITERATURE CITED

Bentham, G. 1844-1845. “The botanical descriptions,” pp. 5-195. Jn: R. B. Hinds (edit.), The botany of the voyage
of H.M.S. Sulphur. Smith, Elder and Co., London.

Greene, E. L. 1885. Studies in the botany of California and parts adjacent. II. Bull. Calif. Acad. Sci. 1: 179-214.

Lindsay, G. 1955. Notes concerning the botanical explorers and exploration of lower California, Mexico. Published
by the author, Stanford.

McKelvey, S. D. 1955. Botanical explorations in the trans-Mississippi west, 1790-1850. Arnold Arboretum, Jamaica
Plains.

Parry, C. C. 1886. Harfordia Greene and Parry. A new genus of Eriogoneae from Lower California. Proc. Davenport
Acad. Nat. Sci. 5: 26-28.
Wiggins, I. L. 1964. “Flora of the Sonoran desert,” pp. 189-1740. Jn: F. Shreve & I. L. Wiggins, Vegetation and
flora of the Sonoran desert. Stanford Univ. Press, Stanford.
- 1980. Flora of Baja California. Stanford Univ. Press, Stanford.

Phytologia (May 1989) 66(3):228-235

REMARKS ON THE GENUS PTEROSTEGIA
(POLYGONACEAE: ERIOGONOIDEAE)

JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

ABSTRACT

The genus Pterostegia is a monospecific taxon ranging from California, eastward to southern
Nevada and adjacent southwestern Utah, western Arizona and northwestern México. An annual,
it is related to the Baja California perennial Harfordia, and together they compose the tribe
Pterostegieae of the subf. Eriogonoideae (Polygonaceae). The genus and its single species are
described, a lectotype selected for the species, and representative specimens are listed document-
ing its known distribution.

KEY WORDS: Polygonaceae, taxonomy, Pterostegia, California, México.

INTRODUCTION

The genus Pterostegia was proposed by Fischer and Meyer in a work devoted to plants
grown in the botanical garden in St. Petersburg, later Petrograd (1914-1924) and now
Leningrad. Friedrich Ernst Ludwig von Fischer (1782-1854) was director of the garden from
1823 until 1850. From 1835 until late 1846, Fischer, along with various coauthors, published
numerous new species and genera found from throughout the world that were in cultivation
at the garden. Many of the western North American species were described jointly with Carl
Anton von Meyer (1795-1855). Meyer was a botanical explorer who traveled widely, mainly
in Russia, became an assistant to Fischer in 1832, and replaced him as director when Fischer
resigned in 1850.

The history of Russian botanical efforts in California have been documented elsewhere
(Essig 1933; Howell 1937; Alden & Ifft 1943; McKelvey 1955; Thomas 1969).

The Russians had come early to the California coast, with the surgeon-naturalist Georg
Heinrich von Langsdorff (1774-1852) on the Krusenstern expedition (1803-1807) visiting the
San Francisco region from late March to mid June of 1806. Langsdorff was able to make a
small collection in spite of numerous difficulties (Langsdorff 1813-1814), but only a few of
his North American species were described early enough to be nomenclaturally significant.
The Russians returned in 1812 to establish an ice-free, warm-weather outpost near Bodega
Bay and at Fort Ross north of San Francisco to assist in maintaining their operations to the
north in Alaska. Four years later Johann Friedrich Gustav von Eschscholtz (1793-1831) and
Ludolf Adelbert von Chamisso (1781-1838) arrived in California aboard the Rurick to begin
one of the most intense period of natural history investigation in California during the first
half of the nineteenth century (Jepson 1929; Mahr 1932; Eastwood 1939, 1944; Lincoln 1966;
McClintock 1967). Not only did Eschscholtz and Chamisso discover numerous species of
vascular plants that would be published in various articles appearing in the first ten volumes

228

Reveal: Remarks on Pterostegia (Polygonaceae) 229

of Linnaea, of which several were from California, but they also established the concept of
intense biological studies at Fort Ross that was followed by a host of naturalists up to the
point when the Fort was abandoned by the Russians in December of 1841.

The place in this story of the plant that Fischer and Meyer (1836) would eventually name
Pterostegia drymarioides began after the adventures of Eschscholtz and Chamisso, the best
known of the Russian botanical collectors in California. The Baron Ferdinand Petrovitch
Wrangel (or Wrangell) arrived in California in 1829 where he eventually assumed the position
of governor of the Russian colony (1831-1836). Brewer (1880) stated that the governor main-
tain his residence at Bodega, but he did visit Fort Ross from time to time (Ogden 1933).
While Wrangel was governor he was an active collector sending specimens and seeds to the
botanic garden in St. Petersburg. Also during his term of office, several naturalists visited
northern California, especially entomologists. Even Eschscholtz returned in 1829 to collect
insects (Essig 1933). Ilya G. Vosnesensky (or Wosnessensky) was one of the few en-
tomologists who would make extensive botanical collections.

TAXONOMY

Pterostegia Fischer & C. Meyer, Index Sem. Hort. Petrop. 2: 48. 1836.—TYPE: Pterostegia
drymarioides Fischer & C. Meyer.

Sprawling and spreading, mostly thinly pubescent, monoecious annual herbs arising from
a slender taproot; /eaves cauline, opposite, exstipulate, petiolate, the blades broadly elliptic
to fan-shaped, entire to variously lobed, glabrous or pubescent, thin, the petioles basally
connate at the non-swollen nodes; flowering stems thin and wiry, dichotomous throughout,
green to red, hirsute; branches numerous, dichotomously branched throughout, the secon-
daries suppressed at the upper nodes, those at the lower nodes short and filiform, often
peduncle-like in position but terminated by a pair of opposite leaves at the terminal node;
inflorescences cymose with the flowers arranged in axillary clusters; bracts lacking; peduncles
lacking; involucres composed of a highly modified involucral bract, this situated lateral of
and basal to a short, slender pedicel, surrounding but basal to the immature female flower
with the immature modified sheath of the bract spreading until after fertilization then be-
coming erect and surrounding the achene, the upper portion of the vertical sheath entire or
variously lobed or notched along its margin, the two lateral wings basal to the immature
sheath, these becoming slightly gibbous on the abaxial surface with age with the adaxial
surface invaginated and its outer margins slightly enlarged, the whole cream or tinged with
pink or rose, hyaline and reticulated with faint cream colored veins, glabrous or pubescent;
flowers imperfect, 2-3 per node, pale yellow to pink or rose, sparsely pubescent along the
midribs and base without, the tepals (5) 6, petaloid, essentially monomorphic, lanceolate,
united about one-third of their length; stamens 6, mostly included, the filaments glabrous,
the anthers yellow, oval; achenes strongly 3-angled, yellowish-brown to brown, glabrous,
winged throughout, the narrow base tapering to a 3-angled beak, the embryo straight, in
abundant endosperm; 2n= 28 (fide Munz 1958).

A monospecific genus of western North America ranging from northern California south to northern Baja
California, México, eastward across southern Nevada to southwestern Utah and Arizona.

Torrey and Gray (1870) segregated Pterostegia from the remainder of the eriogonoid genera
referring it to a new tribe, Pterostegieae. Although they failed to give a formal description
of the taxon, it is diagnostically characterized in their key to the tribes and genera so that

230 PHYTOLOG1A volume 66(3):228-235 May 1989

the name is validly published (Art. 32; Greuter 1988). The tribe now includes two genera,
Pterostegia and Harfordia E. Greene & C. Parry in C. Parry (Reveal 1989), both monospecific.

Like Harfordia, the involucral bract of Pterostegia is highly modified into a slightly gibbous,
reticulated, winged structure that encloses the mature achene. Its development is similar to
that of its perennial relative, but it never obtains the size, prominence or bright colorization
of the involucral bract seen in Harfordia. The ventral side of the bract in Pterostegia is not
keeled nor enlarged as is the case in Harfordia, nor are the two lateral wings truly inflated.
The dorsal opening resembles that found in Harfordia, but the apex of the vertical sheath is
elongated and entire or notched rather than low and flat. The two wings are not inflated as
just noted. Instead in Prerostegia the outer margin of the body of each wing is invaginated
so that into the cavity created by the slightly gibbose abaxial surface, the enrolled margin
forms a short flap that encloses a small portion of that cavity.

Like in Harfordia, the modified involucral bracts must aid in the dispersal of the achenes.
Both the bract and achene are taken by birds, which undoubtly accounts for the plants wide
distribution, but to what extent mature bracts are blown about with achenes inside is un-
known.

1. Pterostegia drymarioides Fischer & C. Meyer, Index Sem. Hort. Petrop. 2: 48.
1836.—LECTOTYPE: “in Portu Bodega,” but more likely near Fort Ross, Sonoma Co.,
California, 1833 (but labelled 1834), Wrangel s.n. (holotype: LE!, selected here.)

Pterostegia diphylla Nutt., Proc. Acad. Nat. Sci. Philadelphia 4: 18. 1848.-TYPE: near Santa
Barbara, Santa Barbara Co., California, 1836, Nuttall s.n. (holotype: PH!; isotype: K!).

Pterostegia diphylla Nutt. var. biloba Nutt., Proc. Acad. Nat. Sci. Philadelphia 4: 18.
1848.—TYPE: near Santa Barbara, Santa Barbara Co., California, 1836, Nuttall s.n.
(holotype: BM!; isotype: GH!).

Pterostegia microphylla Nutt., Proc. Acad. Nat. Sci. Philadelphia 4: 18. 1848.-TYPE: near

Santa Barbara, Santa Barbara Co., California, 1836, Nuttall s.n. (holotype: BM!; isotypes:
GH, K!).

Monoecious, spreading and sprawling, thinly pubescent annual herb 0.1-10 (12) dm across,
dichotomously branched throughout, green to red; leaves opposite, broadly elliptic to fan-
shaped, entire or variously bilobed, 0.3-2 cm long, 0.5-2.5 (3) cm wide, glabrous or pubescent,
the petiole slender and usually pubescent, 0.2-0.6 (1) cm long; inflorescences cymose with
flowers in axillary clusters; involucres reduced to a highly modified involucral bract, 1-1.5 (3)
mm long, (1.5) 2-3 (3.5) mm wide, the vertical sheath entire or variously erose to shallowly
lobed, the two lateral wings slightly gibbous abaxially, invaginated adaxially, reticulate,
glabrous or pubescent; flowers imperfect, 0.9-1.2 mm long, pale yellow to pink or rose,
sparsely pubescent, the tepals (5) 6, lanceolate; stamens 6, included, the filaments 0.5-0.6 mm
long, glabrous, the anthers 0.2-0.3 mm long, oval, yellow; achenes yellowish-brown to brown,

1.2-1.5 mm long, the narrowly globose base tapering to a winged, strongly 3-angled beak;
n= 14. ‘

Widespread and common mainly in shady places from northern California southward throughout much of the
state to northern Baja California, México, including the off-shore islands from Angel Island south to Guadalupe
Island, and eastward across Lincoln and Clark cos., Nevada, to Washington Co., southwestern Utah, southward in
western Arizona from Mohave and Coconino cos. south to Gila, Pima, Pinal and Yuma cos., from near sea level
to 5000 ft elev; flowering from Mar-Jul.

Reveal: Remarks on Pterostegia (Polygonaceae) 231

Representative Specimens. — MEXICO. BAJA CALIFORNIA Norte: Los Arbolitos, Punta Banda, SW of Ensenada,
27 Apr 1978, Ertter et al. 2694 (BRY, MARY, NY); South Todos Santos Island, 7 Apr 1948, Moran 2786 (UC);
Guadalupe Island, NE anchorage, 12 Apr 1948, Moran 2897 (DS, SD); San Martin Island, Hassler Cove, 10 Apr
1963, Moran 10511 (ARIZ, COLO, DS, MICH, RSA, SD, UC); San Martin Island, summit crater, 11 Apr 1963,
Moran 10562 (DS, LL, SBBG, SD, UC, W); W slope of Cerro San Juan de Dios, 1 May 1973, Moran 20682 (SD);
canyon E of San Isidro, Sierra San Pedro Martir, 1 Jun 1975, Moran 22205 (SD); 4 km NE of Jacomun, Sierra
Judrez, 14 May 1977, Moran 24024 (SD); Guadalupe Island, 1875, Palmer 84 (BM, G, GH, K, MO, NY, PH); San
Quintin Bay, Jan 1889, Palmer 667 (ARIZ, BR, CAS, DS, MICH, UC, US); Cedros Island, 18-20 Mar 1889, Palmer
710 (F, GH, K, NY, US); along México Highway 3, 12.6 mi S of México Highway 2 in Tecate, 21 May 1988, Reveal
6845 (BRY, CAS, MARY, MEXU, MO, RSA, WIS); Escondido Creek, 13 mi S of Punta Prieta, 23 Feb 1935,
Shreve 6916a (ARIZ, US); 11 mi E of San Telmo, 4 May 1941, Wiggins 9741 (DS, US). MiCHOACAN: Morelia, near
the prison, Aug 1927, Arsené s.n. (Z). UNITED STATES. Arizona: Coconino Co.: side canyon at Colorado River
Mile 184, Grand Canyon, Apr 1975, Theroux 1199 (ARIZ, MNA). Gila Co.: San Carlos Indian Reservation, Cas-
sadore Spring Canyon, 20 mi N of San Carlos, 29 Mar 1935, Maguire et al. 10321 (ARIZ, BRY, DAO, GH, NY,
UTC, WTUV); W end of Canyon Lake, Apache Trail, 2 May 1935, Nelson & Nelson 1731 (G, K, MO, NY, RM,
SMU, UC, UTC). Maricopa Co.: Tempe, Papago Park, 19 Feb 1932, Gillespie 8811 (DS, GH, NY, POM, UC, US);
Canyon Lake, 16 Mar 1935, Peebles 10757 (ARIZ, MICH, POM). Mohave Co.: Virgin Narrows, 18 mi S of St.
George along Interstate Highway 15, 24 Mar 1973, Atwood 4396 (BRY, COLO, MO, US, UT). Pima Co.: canyons
of the Santa Catalina Mountains, 11 Apr 1881, Pringle s.n. (G, GH, LE, NY, PENN, PH, VT, WS); Sabino Canyon,
Santa Catalina Mountains, 3 Apr 1941, Shreve 10118 (ARIZ, COLO, DS, MICH, UC); Organ Pipe Cactus National
Monument, Alamo Canyon, 2 Apr 1982, Siplivinsky et al. 2720 (CAN, COLO, RM, TEX, UCR, UTC). Pinal Co.:
Superstition Mountains, 26 Mar 1932, Gillespie 5495 (DS, US); 2 mi below Coolidge Dam along the Gila River,
3 Apr 1935, Maguire 10435 (BRY, DAO, NY, UTC). Yavapai Co.: 5 mi N of Congress Junction, 9 Apr 1947, Gould
& Darrow 3642 (ARIZ); Hillside, 1 May 1903, M.E. Jones s.n. (BKL, CAS, DS, POM); Lake Pleasant Regional
Park, Cottonwood Creek, 31 Mar 1976, Lehto 19728 (ASU). Yuma Co.: Havasu National Wildlife Refuge on low
hills S of Bill Williams River, 1.8 mi E of Arizona Highway 95, 25 Feb 1973, Russell & Kobetich 73-10] (ASU).
CALIFORNIA: Alameda Co.: west base of Mt. Diablo, 1868, Gibbons s.n. (GOET, MPU, WU); Grizzly Peak, Berkeley
Hills, 29 Apr 1933, Tracy 12077 (DAO, UC). Amador Co.: Fisher’s Cabin, 17 Apr 1893, Hansen 316 (E, G, MIN,
NY, US). Butte Co.: Chico, Mar 1883, Austin s.n. (NY, US); Stilson Canyon E of Chico, 13 Apr 1916, Heller s.n.
(F, PENN). Calaveras Co.: Angeles Camp, 11 Apr 1923, Eastwood 11650 (CAS); E edge of North Shore Camp,
New Hogan Reservoir, 20 Apr 1972, McNeal 770 (CPH, NY). Colusa Co.: along Bear Valley Road 1.5 mi from
California Highway 20, 22 Apr 1978, Logan 63 (HSC). Contra Costa Co.: slopes of Mt. Diablo, 27 Apr 1940, H.S.
Reed 346 (DAO, OKL). El Dorado Co.: 0.5-1 mi NW of Chili Bar on the South Fork of the American River, 8
Apr 1978, Smith & Stebbins 7807 (CAS). Fresno Co.: 5 mi NE of Academy, 10 May 1954, Howell & Barneby
29328 (CAS); Alcalde Hills, 12 mi W of Coalinga Canyon of Los Gatos Creek E of White Creek, 24 Mar 1986,
Janeway & Janeway 1398 (CHSC); along Kings River in upper Pine Flat, 0.7 mi above old Pine Flat School, 24
Mar 1953, Quibell & Quibell 1720 (RM, RSA). Glenn Co.: 8 mi S of Elk Creek on Stony Ford Road, 13 Apr 1971,
Maze et al. 931 (NY, WTU). Humboldt Co.: Cape Mendocino, near ocean shore, 20 May 1933, Tracy 12226 (DS,
GH, NY, UC, UTC, WS, WTU). Imperial Co.: Mountain Springs Road exit off Interstate 8, 0.3 mi E of Mountain
Springs at head of Myer Creek, 10 Mar 1986, Sanders et al. 6275 (UCR). Inyo Co.: Surprise Canyon, Panamint
Range, 15 Apr 1891, Coville & Funston 646 (DS, F, GH, K, LE, MIN, MO, MSC, NY, PH, US); Alabama Hills,
S of Lone Pine Creek, 1 May 1974, DeDecker 3421 (CAS, RSA, WS); Goler Canyon, Panamint Range, 25 Mar
1937, Train s.n. (ARIZ, COLO, DS, ILL, NEB, OSC, RSA, US). Kern Co.: Sunset, 20 Apr 1902, Heller 7731
(ARIZ, BKL, E, G, GH, ISC, MO, NY, PH, US); Kern Canyon, 1 mi above its mouth, 26 Apr 1905, Heller 7780
(BKL, E, G, ISC, MIN, MO, NY, PH, US); Long Tom Mine, Greenhorn Mountains, 26 Apr 1965, Twisselmann
10615 (CAS, NY, RSA). Lake Co.: on the shore of Clear Lake, 3 mi S of Lucerne, 20 Apr 1963, Breedlove 4581
(SMU). Los Angeles Co.: Mandeville Canyon, Santa Monica Mountains, Mar 1929, Clokey & Templeton 4523 (CM,
F, G, GH, ILL, K, LE, MARY, MICH, MIN, MONTU, ND, NO, OKL, PENN, POM, RM, RSA, UC, US, UTC,
WS); Glendora, 7 May 1904, Grant & Wheeler 1057 (BKL, F, GH, ILL, NESH, NMC, PH, RM, UC); San Clemente
Island, Middle Ranch, 17 Feb 1941, Moran 576 (DS, MARY, MO, NY, RSA, SBBG, SD, WTU); Renton Mine,
Santa Catalina Island, 22 Jun 1965, Thorne & Everett 34844 (RSA, SBBG, SD). Madera Co.: Raymond, 8 May
1925, Eastwood 12511 (CAS). Marin Co.: San Rafael Hills, 6 May 1945, J.T. Howell 20806 (CAS, UC); NW side
of Angel Island, 2 May 1967, Raven & Johnson 21331 (DS); Point Reyes Bird Observatory, Arroyo Hondo, 3 mi
NW of Bolinas, 20 Feb 1970, True & Ross 5178 (CAS). Mariposa Co.: mountains above Exchequer Dam on the
Merced River, 30 Apr 1949, Rodin 4913 (BR, CAN, COLO, UC, WS); Bear Mountain, 1865, Torrey 431 (GH,
MIN, NY, US). Mendocino Co.: trail around Leonard Lake, head of Reeves Canyon, 21 May 1980, G.L. Smuth
5767 (CAS); Monterey Co.: Tassajara Hot Springs, Jun 1901, Elmer 3252 (DS, G, K, MIN, ORE, US); Pacific
Grove, Apr 1902, Elmer 3557 (BKL, CAS, COLO, DS, E, G, GH, K, MICH, MIN, MO, NY, POM, SBBG, SMU,

232 PHYTOL O GIA volume 66(3):228-235 May 1989

UC, US, VT, WTU, Z); Arroyo Seco Road, W of Greenfield and just W of the junction with Carmel Valley Road,
11 May 1980, Ertter & Strachan 3343 (CAS, MARY, NY, SD); 8 mi N of Carmel Valley, Santa Lucia Mountains,
27 Apr 1957, J.T. Howell 32037 (CAS, RSA, W). Napa Co.: Wooden Valley Grade, W side of Napa Valley, 2 Apr
1931, Keck 1024 (DS, POM). Nevada Co.: Rock Creek, 14 mi E of Wheatland, 25 Mar 1965, True 1759 (CAS).
Orange Co.: Silverado Canyon, Santa Ana Mountains, 20 Apr 1920, Munz & Harwood 3724 (PH, POM, RM); 0.5
km W of Dana Point Harbor, 0.8 km SSW of the R-H. Dana School, Dana Point Headlands, 3 Apr 1982, Roberts
& Fritzke 498 (UCSB). Riverside Co.: Oasis de los Osos Preserve, off Snow Creek Canyon at the N foot of the
San Jacinto Mountains, 1 mi W of Snow Creek Village, 9 Apr 1985, Fellows & Sanders 125 (HSC, OBI, UCR); 8
mi SE of Corona along road to Elsinore, 17 Mar 1964, Hitchcock & Muhlick 23079 (F, HSC, NY, WS); 1 mi below
Cottonwood Spring, Joshua Tree National Monument, 1 Apr 1940, C.L. Hitchcock 5909 (DS, MO, NY, RSA, UC,
UTC, WTU); San Jacinto Mountains, 1 Jul 1934, Stokes 313 (CAS, DS, G, GH, K, MICH, MIN, NY, POM, UC,
US, UTC, Z). Sacramento Co.: near Folsom, 22 Apr 1928, Copeland 531 (MARY, POM). San Benito Co.: 1.2 mi
SW of Gabilan School, 20 May 1936, Belshaw 2165 (UC). San Bernardino Co.: 4 mi E of Victorville, 18 Apr 1941,
Alexander & Kellogg 2024 (DS, GH, UC, UTC); Cucamonga Wash, NE of Upland, 1 Mar 1917, ILM. Johnston 1352
(DS, MONTU, POM, UC, WTU); near Bonanza, Providence Mountains, 30 Mar 1920, Munz & Harwood 3536
(BKL, DS, POM, RM); Sacramento Mountains, South Pass, 20 mi W of Needles along U.S. Highway 66, 1 Apr
1958, Smith & Hansen 95 (RSA); Arrowhead Canyon, 7 May 1919, Spencer 1105 (CAS, GH, NY, POM); Willow
Spring Canyon, Old Dad-Granite Mountain Range, 28 Apr 1941, Wolf 10053 (DS, NY, RSA). San Diego Co.:
Mission Hills, San Diego, 6 May 1903, Abrams 3414 (BM, DS, E, F, G, GH, K, MO, NMC, NY, PH, POM, US,
Z); Del Mar, 4 Apr 1914, Clements & Clements 55 (COLO, F, GH, ILL, MICH, MO, NEB, NY, PENN, PH,
UCSB); Palomar Mountains, 2.6 mi NE of California Highway 76, 28 Jun 1987, Reveal & Broome 6616 (CAS,
MARY, MICH, MO, NY, RM, RSA, US, WIS); San Felipe Ranch between Julian and Imperial Valley, 20 Mar
1926, Wiggins 2011 (DS, SD, WTU). San Francisco Co.: Obsolescent Valley, Apr 1892, Michener & Bioletti s.n.
(BKL, MICH, MIN, MSC, NEB, POM); between Lands End and the Golden Gate Bridge, 18 Mar 1956, Raven
8866 (CAS). San Joaquin Co.: hill between Castle Rock and Black Butte, Corral Hollow, 9 Apr 1938, Hoover 3046
(K, NY, UC, US). San Luis Obispo Co.: Baywood Park at Sweet Springs Marsh, S end of Morro Bay, 25 Apr
1981, Keil 14580 (OBI, UCR); Arroyo de la Cruz 1 mi E of California Highway 1, 18 May 1980, Keil et al. 13996
(ASU, OBI); La Panza Range, 0.4 mi W of La Panza Summit along the road to La Panza, 26 May 1988, Reveal
6902 (BM, BRY, CAS, G, MARY, MO, NY, OSC, RM, RSA, US, WIS). San Mateo Co.: Saratoga-Big Basin
Road, 23 Jun 1915, Abrams 5261 (DS, MICH); Franklin Point, 13 May 1961, Thomas 9333 (DS, OSC, RSA, US).
Santa Barbara Co.: W of Canyon del Mar and S of Cuyler Harbor, San Miguel Island, 23 May 1963, E.R. Blakley
5857 (SBBG); ravine W of Pelican Bay, Santa Cruz Island, 6 Jun 1930, Clokey 4904 (GH, K, MARY, ND, NY,
OKL, PENN, POM, UC, US); Dix Canyon, Santa Barbara Island, 26 May 1930, Clokey 4906 (ILL, MIN, MO, ND,
NY, PENN, RSA, UC, WTU); Santa Rosa Island, 24 May 1931, N. Dunn s.n. (LA); West Camino Cielo, 2-3 mi
W of California Highway 154, Santa Ynez Mountains, 17 May 1980, Ertter & Strachan 3438 (ASU, BRY, CAS,
COLO, CPH, ISC, MARY, MONTU, NESH, NY, RM, SD, TEX, US, UTC, WS, WTU); 4 mi S of Surf, 14 Apr
1929, Wiggins 3517 (DS, IDS, POM, WTU). Santa Clara Co.: foothills near Stanford University, 15 Apr 1902, C.F.
Baker 615 (G, GH, K, MICH, MO, NDG, NY, POM, UC, US); Stevens Creek, near Soda Rock, 18 May 1907,
Heller 8549 (BKL, DS, E, F, G, GH, ISC, MIN, NY, PH, US, WTU, WU). Santa Cruz Co.: Santa Cruz, 24 Jun
1881, M.E. Jones 2239 (BM, BR, CAS, DS, G, GB, LE, MIN, MSC, NY, POM, RM, US, UTC); Ben Lomand, 18
Jul 1962, Rose 62074 (ARIZ, B, BR, CAS, DAO, DS, G, GB, GH, MICH, MIN, NY, OBI, RM, RSA, SMU, TEX,
WS, WTU). Shasta Co.: Morley’s Station, 22 May 1894, Baker & Nutting s.n. (POM, RM, UC). Solano Co.: Gates
Canyon, near Vacaville, 27 Apr 1902, Heller & Brown 5386 (BKL, DS, F, GH, MO, NY, PH, US). Sonoma Co.:
Bodega Point, 4 Jul 1900, Eastwood s.n. (DS, GH, RM, US, Z); Bodega Bay, 27 May 1902, Heller & Brown 5612
(B, DS, E, F, G, GH, ILL, LE, MO, NY, PH, POM, US). Stanislaus Co.: Lower Crow Creek Canyon, 7 Apr 1940,
Hoover 4317 (NY, UC, US); 4 mi up Arroyo del Puerto Canyon, 21 Apr 1935, Sharsmith 1760 (DS, GH, UC, WS).
Sutter Co.: 2 mi SE of the Dean Cabin, 2 mi N of Sutter, Sutter Buttes, 5 May 1985, Ahart 4897 (CHSC, MO);
summit of the ridge E of South Butte, Marysville Buttes, 22 Mar 1905, Heller 7571 (BKL, E, ISC, MIN, PH, NY,
US). Tehama Co.: 5 mi W of Paskentain foothills, 26 Apr 1953, M.O. Baker 12561 (RSA). Tulare Co.: Lindsay, 20
Mar 1925, Munz 9051 (POM); Kern Canyon, flume road at Corral Creek above Corral Flat, 25 Apr 1971, Twis-
selmann 17597 (CAS). Tuolumne Co.: Pefion Blanco Mine near Indian Creek, 11-16 Apr 1919, Ferris 1526 (CAS,
DS, NY, UC); above Rawhide Road, 0.3 mi SW of the Jamestown-Tuttletown Road, NW of Jamestown, 31 Mar
1972, McNeal 707 (CPH, NY). Unknown Co.: without location data, without date, Coulter s.n. (BM, CGE, E, G,
GH, K, NY); without location data, without date, Douglas s.n. (BM, CGE, GH, K, NY, OXF); without location
data, 1876, Parry & Lemmon 365 (F, ISC, MO, NEB, US). Ventura Co.: S of the lighthouse, East Anacapa Island,
25 Apr 1959, E.R. Blakley B2776 (SBBG); summit of East Casitas Pass, Santa Ynez Mountains, 19 Apr 1962,
Breedlove 2457 (DS, UCSB); Matilija Canyon, 15 Apr 1945, Pollard s.n. (GB). Yuba Co.: near Marysville, May
1854, Bigelow s.n. (NY). NEVADA: Clark Co.: near Dripping Springs Canyon, 35 mi SE of Searchlight, 2 Apr 1966,

Reveal: Remarks on Pterostegia (Polygonaceae) 233

Cronquist 10606 (BRY, NY, UTC, WTU); Valley of Fire, Atalta Park, 6 Apr 1934, Maguire et al. 4770 (GH, ND,
RM, WS, UTC); Muddy Mountains, N end of Hidden Valley, 11 May 1980, Swearingen 1433 (ARIZ, RSA, UNLV);
S end of Arrow Canyon Range, N of Dry Lake Valley, 15 May 1983, Tiehm 7616 (BRY, CM, IDS, NY, RSA,
UTC). Lincoln Co.: SW side of Meadow Valley Mountains, 11 Apr 1979, Tiehm & Williams 4784 (MARY, NY,
UTC). NEw Mexico: Unknown Co.: Rio Grande Valley, below Donafia, without date, Parry et al. 1171 (US).
OreGon: Unknown Co.: Columbia River, without date, Tolmie s.n. (GH). UTAH: Washington Co.: St. George, S
slope of Black Hill, 16 Apr 1942, Gould 1563 (ARIZ, CAS, DS, F, ND, UC, US); St. George, 1 Apr 1880, M.E.
Jones 1618 (ARIZ, BKL, BM, BR, F, G, GB, LE, MICH, MSC, NY, POM, US, UT, UTC); without location data,
1876, Parry 229 (CM, F, G, MO, NY, PH, US); Cedar Pockets Wash, 5 May 1986, Welsh & Atwood 23740 (NY).
A total of 1259 collections were examined.

The interesting question regarding the typification of Pterostegia drymarioides (from the
genus name Drymaria Willd. ex Schultes (Caryophylloides) and -oides, resembling, alluding
to the spreading nature of the species) was whether one should select the voucher of the
seed collection gathered in the wild by Wrangel or one of the garden specimens made by
Fischer and sent to various institution with a variety of dates. The name of the taxon appeared
in a treatment of plants grown at the botanic garden in St. Petersburg and there is a garden
voucher at LE collected in 1835.

The Wrangel specimen selected here as the lectotype was supposedly collected near Fort
Ross and dated 1834. The sheet was named Pterostegia minutiflora with the actual location
data given as “Lect: in California prope colonoam Ross. Acc. a D. Wrangel, 1834.” Fischer
and Meyer (1836) give the type location of Bodega Point. It is uncertain which is correct,
but given the general statement given to the entire series of species by Fischer and Meyer,
it is likely that the reference to Bodega Point was a general one and that the actual place
where the type was gathered is near Fort Ross. It is also uncertain if the type specimen was
actually gathered in 1834 or if that was the year the voucher arrived at St. Petersburg. Ac-
cording to Ogden (1933), Wrangel was definitely at Fort Ross in 1833 and because it is
doubtful a specimen gathered in the summer of 1834 could have made it to St. Petersburg
early enough for the seeds to be planted for the 1835 growing season, it is suggested that
the type was actually gathered in 1833.

The 1835 garden specimen at LE is here considered to be a syntype but it is unclear how
other garden specimens should be regarded. At BM and G are two collections stating “Fis-
cher” and “Bodega.” The G sheet is dated 1840 and it is uncertain if that is the year the
plants were collected in the garden at St. Petersburg or that was the year the sheet was sent
to G. Other specimens at BM and GOET are simply labelled “California.” At GOET one
sheet is from the Fischer Herbarium with something like “Petrib 36” written on it. Does this
mean it is a garden collection gathered at St. Petersburg in 1836? It is assumed here that
the Fischer specimens alluding to California and Bodega are not Wrangel isotypes, but rather
garden specimens grown from seeds gathered in California.

At K are three sheets of garden specimens grown at Luxembourg. One is dated 25 Aug
1836 and is accompanied by a detail description. On the same sheet is another collection
that was apparently gathered in 1837. The second sheet is dated 28 Aug 1836 and associated
with it are two sketches; a third sketch was apparently made from a Douglas specimen
mounted on the same sheet.

The specimen gathered in 1837 is dated 10 Aug 1837 and the label data state that it was
grown from seeds sent to Luxembourg by Fisher. Furthermore, seeds were gathered at
Luxembourg on 16 Jun 1837. A third Luxembourg sheet, dated 19 Jul 1836, is also at K.

A Meisner collection (BM) is labelled “Hort. Basil” and dated Sep 1839. It refers to the
type location as “Hab in Portu Bodega Nova California,” this being taken from a report by
Hooker (1836). It is uncertain what the seed source of this collection might have been, but

234 PHYTOL O GIA volume 66(3):228-235 May 1989

most likely seeds were sent to Meisner at Basel by Fischer.

It is clear that Fischer widely distributed seeds of Pterostegia drymarioides, but it is uncer-
tain at this time if the seeds that went to Luxembourg, for example, were some originally
gathered by Wrangel or were taken from the specimens grown at St. Petersburg.

According to Peck (1942), Pterostegia drymarioides is found in southeastern Oregon. The
only specimen from Oregon discovered during this study is a Tolmie collection supposedly
gathered along the Columbia River.

William Frazer Tolmie (1812-1886), a surgeon-naturalist for the Hudson Bay Company,
came to the coast of North America in 1833 traveling overland from Fort Vancouver, past
Mount Rainier, toward the Puget Sound where he was to remain until the spring of 1836.
He stayed on the Columbia River until 1841 when he walked to Fort York on Hudson Bay
and sailed for England. In 1843 he returned to Fort Vancouver but almost immediately went
to the Puget Sound again to oversee operations at Fort Nisqually. When the international
boundary between the United States and Canada was settled, Tolmie moved to Victoria in
1859 where he lived until his retirement in 1870. McKelvey (1955) alludes to an 1840 trip by
Tolmie into the Willamette Valley, but there is no indication that Tolmie ever collected in
southeastern Oregon.

Thanks to McKelvey (1955) information now exists about a “friend of Mr. Tolmie” who
collected specimens for him that eventually found their way into the hands of William Jackson
Hooker who, with George Arnott Walker-Arnott, described several as new species in their
report on the botany of the Beechey voyage. This was John McLeod who went to the Green
River rendezvous representing the Hudson Bay Company. He traveled from Fort Vancouver
to the mid July meeting of trappers and fur traders, and returned sometime prior to the end
of October in 1837. McLeod was a trapper himself and was certainly inspired by Tolmie to
make a small collection during his travels. McKelvey concludes that the trapper went up the
Columbia River to Fort Walla Walla, across northeastern Oregon to the Umatilla River and
on to the Snake River which he followed to Fort Boisé. On his return trip, McLeod apparently
retraced his original path.

It is possible Peck was aware of the Tolmie collection at GH and felt that it could not
have been collected on the Columbia River. If he reasoned the specimen was collected by
Tolmie’s “friend” then perhaps he reasoned it must have come from southeastern Oregon.

An additional possibility is that Tolmie encouraged others to collect for him and somehow
he obtained a specimen of Pterostegia from someone else. There is no Tolmie specimen
among the Hooker specimens at K, E or TCD so there is an additional possibility that the
sheet at GH might have been mislabelled.

New Mexico is not included in the distribution of Pterostegia even though a specimen is
cited in the above exsiccata. The specimen is associated with a label from the United States-
México boundary survey, and it is strongly suspected the sheet was actually taken near San
Diego in southern California rather than near the Rio Grande in southern New Mexico.

As for the Morelia collection of Pterostegia made by Arsené in the state of Michoacan, I
cannot account for it. No other specimens of Arsené of the genus are known, but it is known
that he collected in the state. There is no indication that the species has been found anywhere
else on the mainland of México.

There is considerable variation within Pterostegia drymarioides in the size of its parts and
the pubescence. No geographic significance has been associated with this variation except
that the involucral bracts on plants found on the Mojave Desert tend to be slightly larger
than those encountered in the coastal mountains. The species is generally found in cool,
moist places in the shade were it spreads over the ground under shrubs. The degree of

Reveal: Remarks on Pterostegia (Polygonaceae) 235

available moisture and shade seem to influence the size of individual plants.

ACKNOWLEDGMENTS

I wish to thank the curators and staff of the following herbaria for loans or for arranging
access to their collections during visits: A, ARIZ, ASC, ASU, B, BKL, BM, BR, BRY, CAN,
CAS, ‘CGE, CHSC, CM; COLO; CPH,:CS, DAO; DAV, DS, E, F) G, GB, GH, GOET,
HSC, ID, IDS, ILL, IND, ISC, JEPS, K, KANS, KSC, L, LL, M, MARY, MICH, MIN,
MNA, MO, MONTU, MPU, MSC, ND, NDG, NEB, NESH, NMC, NO, NTS, NY, OBI,
OKL, OKLA, ORE, OSC, OXF, P, PENN, PH, POM, RENO, RM, RSA, SBBG, SD, SFSU,
TCD; TEX, UC, UCR, UCSB, UNLV, US, USFS, UT, UTC, VT; W, WILLU, WIS, WS,
WTU, WU, and Z. Individuals wishing a complete list of herbarium specimens examined
may request a copy. Funding for field work has been made possible through a variety of
sources. I would like to thank Dr. Edward E. Terrell and Clare B. Hardham for their review
comments. This is Scientific Article A-4945, Contribution 7988, Maryland Agricultural Ex-
periment Station.

LITERATURE CITED

Alden, R. H. & J. D. Ifft. 1943. Early naturalists in the far west. Occas. Pap. Calif. Acad. Sci. 20: 1-59.

Brewer, W. H. 1880. “List of persons who have made botanical collections in California,” pp. 553-559. In: S. Watson,
Botany of California. Vol. 2. John Wjlson and Son, Cambridge.

Eastwood, A. 1939. Early botanical explorers on the Pacific coast and the trees they found there. Quart. Calif.
Hist. Soc. 18: 335-346.

- 1944. The botanical collections of Chamisso and Eschscholtz in California. Leafl. W. Bot. 4: 17-21.

Essig, E. O. 1933. The Russian settlement at Ross. Quart. Calif. Hist. Soc. 12: 191-209.

Fischer, F. E. L. von & C. A. von Meyer. 1836. Index seminum, quae Hortus botanicus imperialis petropolitanus
pro mutua commutatione offert. Index secundus. Published by the authors, St. Petersburg.

Greuter, W. (edit). 1988. International code of botanical nomenclature. Regnum Veget. 118: 1-328.

Hooker, W. J. 1836. Botanical information. Comp. Bot. Mag. 2: 6-12.

Howell, J. T. 1937. A Russian collection of botanical plants. Leafl. W. Bot. 2: 17-20.

Jepson, W. L. 1929. Johann Friederich Eschscholtz. Madrofio 1: 253.

Langsdorff, G. H. von. 1813-1814. Voyages and travels in various parts of the world, during the years 1803, 1804,
1805, 1806, and 1807. 2 vols. London. [Translation of the original 1812 German work Bemerkungen auf einer
Reise um die Welt in den Jahren 1803-1807. 2 vols. Frankfurt am Main.]

Lincoln, A. 1966. Discoveries of the Rurik expedition. Pacific Discovery 19: 14-19.

Lindsay, G. 1955. Notes concerning the botanical explorers and exploration of lower California, México. Published
by the author, Stanford.

Manhr, A. C. 1932. The visit of the “Rurik” to San Francisco in 1816. Stanford Univ. Publ. Univ. Ser., Hist., Econ.
& Political Sci. 2(2): 1-194.

McClintock, E. 1967. Early plant exploration in the West. Part I. Calif. Hort. Soc. 28: 114-121.

McKelvey, S. D. 1955. Botanical explorations in the trans-Mississippi west, 1790-1850. Arnold Arboretum, Jamaica
Plains.

Munz, P. A. 1958. A California flora. University of California Press, Berkeley.

Nuttall, T. 1848. Descriptions of plants collected by Mr. William Gambel in the Rocky Mountains and upper
California. Proc. Acad. Nat. Sci. Philadelphia 4: 7-26.

Ogden, A. 1933. Russian sea-otter and seal hunting on the California coast, 1803-1841. Quart. Calif. Hist. Soc. 12:
217-239.

Reveal, J. L. 1989. A revision of the genus Harfordia (Polygonaceae: Eriogonoideae). Phytologia 66: 221-227.

Thomas, J. H. 1969. Botanical explorations in Washington, Oregon, California and adjacent regions. Huntia 3: 5-62.

Phytologia (May 1989) 66(3):236-245

NOTES ON SELECTED GENERA RELATED TO ERIOGONUM
(POLYGONACEAE: ERIOGONOIDEAE)

JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

ABSTRACT

The subtribe Eriogonineae consists of seven genera including the largest genus of the subf.
Eriogonoideae, Eriogonum. Revisions have been published on four of the genera and two more
modest treatments are presented here for Dedeckera and Gilmania.

KEY WORDS: Polygonaceae, taxonomy, Dedeckera, Gilmania, California.

INTRODUCTION

Current revisionary studies of the eriogonoid genera of Polygonaceae began in 1961 with
the discovery of a series of wild buckwheat species that would eventually prove to be un-
described. It was as a naive undergraduate that the initial efforts were made to comprehend
the genus Eriogonum under the tutelage of John Thomas Howell, George J. Goodman, Ar-
thur Cronquist and Arthur H. Holmgren. Upon entering the exalted air of graduate school,
first in a master’s and then a doctoral program, the same group continued to help with
Stanley L. Welsh assuming the principal role of advisor and Conrad V. Morton chief critic.
The result was a revision of Eriogonum that, much to the disgruntlement of many, is still
unpublished. Those years, as almost any professor will rightly claim later, were golden ones
in terms of productivity.

Since 1964 numerous papers were published treating Eriogonum. With the discovery of
the genus Dedeckera (Reveal & Howell 1976), greater attention was paid to the genera related
to it, and a series of revisions were published with Barbara J. Ertter (Reveal & Ertter 1977a,
b, 1980; Ertter 1980). An administrative change above the departmental level in the late
1970s made it impossible to work on western American botany and most efforts on the
Eriogonoideae came to an end. Another change in the mid 1980s made it possible once again
to concentrate on western plants and this time it is being done with a vengeance!

The current remarks are devoted to those genera related to Enogonum belonging to the
subtribe Eriogonineae. Revisionary treatments of Chorizanthe and its related genera (subtribe
Hollisteriineae) are presented elsewhere in this volume of Phytologia (Reveal 1989c; Reveal
& Hardham 1989) as are comments on the two genera of the tribe Pterostegeae (Reveal
1989a, b). With this paper, and those mentioned above, all of the eriogonoid genera will have
been revised except Eriogonum and the perennial species of Chorizanthe which are funda-
mental to any understanding of the evolutionary history of the subfamily.

Reveal: Genera related to Eriogonum (Polygonaceae) 237

TAXONOMY

The subfamily Eriogonoideae is divided into two tribes, Eriogoneae (fifteen genera) and
Pterostegeae (two genera). The former is further divided into subtribes, Eriogonineae (seven
genera) and Hollisteriineae (eight genera). The Eriogonineae may be characterized as those
genera of the tribe with numerous flowers in turbinate to campanulate involucres or at a
node. In this taxon the involucres are infrequently awned (Oxytheca, Goodmania and then
the awns are long, slender and always straight). Although not consistent in either subtribe,
the embryo is curved in the Eriogonineae (the exception being Eniogonum subgenus
Oligogonum) and typically straight in the Hollisteriineae (the exceptions being three species
in Chorizanthe and the species of Systenotheca, Centrostegia, Dodecahema, Aristocapsa and
Hollisteria). The Hollisteriineae is reviewed elsewhere (Reveal 1989c).

Polygonaceae Juss. subtribe Eriogonineae Roberty & Vautier, Boissiera 10: 84. 1964. — TYPE:
Eriogonum Michx., according to Art. 22.4 of the ICBN.
Polygonaceae cohort Eriogonastreae Roberty & Vautier, Boissiera 10: 92. 1964.—TyYPE:
Eriogonum Michx.
Polygonaceae Juss. cohort Pterogonastreae Roberty & Vautier, Boissiera 10: 107. 1964.-
TYPE: Pterogonum H. Gross.

Plants annual, biennial or perennial herbs, subshrubs or shrubs; /eaves basal or basal and
cauline, mostly broad, glabrous to densely tomentose on one or both surfaces; inflorescences
mostly cymose to umbellate or capitate, the secondaries rarely suppressed; peduncles typically
present, usually slender to stout and often elongated; involucres turbinate to campanulate or
sometimes cylindrical, the teeth short and erect or lobed and typically reflexed, infrequently
awned with long slender spines, occasionally the involucre reduced to a series of (2-3) 5-6
awned or non-awned involucral bracts; flowers (2) 7-200 per involucre or 5-30 at a node,
rarely some solitary in Dedeckera, usually long pedicellate, the tepals narrow to broad,
glabrous to tomentose; achenes mostly globose with a curved (rarely straight) embryo; n=
Orit 2,16; 1:7; 18,20), 22540.

Widespread in North America from eastern Alaska southward to central México eastward to the Applachian
Mountains southward to central Florida, with one species disjunct in southwestern South America, the greatest
concentration in the western United States and especially in California.

As here defined, the subtribe Eriogonineae is composed of seven genera, the largest being
Eriogonum with some 240 species of annuals, biennials or perennials. This genus is widely
distributed in North America being encountered from Alaska to central México and from
coast to coast. It may also be found from well below sea level to above timberline. The next
largest genus in the subtribe is Oxytheca, a taxon of seven annual species of the western
United States and northern Baja California Norte, México, with one variant disjunct in Chile
and Argentina. Of the five remaining genera only the monospecific genus Dedeckera is a
perennial, and it is a well formed shrub of the desert ranges of Inyo and Mono cos., California.
Two of the monospecific annual genera, Gilmania and Goodmania, are found mainly in the
Mojave Desert with the former endemic to Death Valley. The latter is more widespread. It
extends from the edge of the San Gabriel and San Bernardino mountains northward into the
Central Valley and along the eastern edge of the Sierra Nevada into the Great Basin of
west-central Nevada. To the south along the coast and in the Sonoran Desert of southern
California and northwestern México the monospecific genus Nemacaulis is found on deep

238 PHYTOLOG1IA volume 66(3):236-245 May 1989

sandy soil. To the east in the Colorado River drainage system of Wyoming southward through
eastern Utah and adjacent Colorado into the Four-Corners area of New Mexico and Arizona
is the bispecific annual genus Stenogonum. Of the fifteen genera of Eriogoneae only
Stenogonum is not found in California.

Key to the Genera

A. Plants perennials, or if annual then the well defined involucral tube lacking awn-tipped teeth.
B. Involucres with a distinctly tubular involucre, the lobes or teeth sometimes united only at the base;
annuals to perennials widely distributed in North America. ...........:::sccsesesecseseesesreseeesseeneeneaees 1. Eriogonum
BB. Involucres reduced to a series of obscure 2-5 involucral bracts; strictly perennial; endemic to Inyo and
Mono C0S., ‘Califia, oo ienieectceac. cnscdetnssseosisecassedsasessatsosiisespatcnncsancenca sataeaumsaeiecnepera eaten 2. Dedeckera
AA. Plants annual; involucres lacking or reduced to a series of involucral bracts or if present then with awn-tipped
teeth.
B. Involucres tubular with long straight spines or the bracts arranged in two whorls of three; deserts of
western North America and southwestern South America.
C. Involucral bracts arranged in 2 series of 3; flowers yellow, pubescent; southwestern Wyoming to

northwestern New Mexico and northeastern APrizOma. .........ccsssessssessesessessereeseeeereseeseesess 3. Stenogonum
CC. Involucres tubular, all the teeth in a single whorl and awn-tipped; deserts of western North America
and southwestern) South tA mericas | Sorccceccecercscecesmeceensanssessascerseceseevesenstesteeeneeeens eee =eee teense 4. Oxytheca

BB. Involucres lacking or composed of a series of 5 bracts in a single whorl; plants mainly of California and
northwestern México.

C. Flowers pubescent; stamens 9; Mojave Desert northward.
D. Involucral bracts 5, linear-lanceolate; Kern and Los Angeles cos. northward to Madera Co.,

California; and) Mineral! 'Co:, Nevada. 2.2...::.0c.<cecssscsoococeccendsscetacosteetontaes mies ee Eee 5. Goodmania

DD. Involucral bracts lacking; Death Valley, Inyo Co., Califormia. ..........csssseseseecseeese 6. Gilmania

CC. Flowers glabrous; stamens 3; coastal California from Los Angeles Co. south to central Baja Cali-
fornia Norte eastward in the Sonoran Desert to northwestern Sonora, México. ...... 7. Nemacaulis

1. Eriogonum Michx.

Efforts to complete a revision of this genus continue and hopefully can be completed as
part of the overall Flora North America project. State and regional treatments will be done
for California (Reveal 1989d), for a new flora of Arizona and volume 2 of the Intermountain
Flora. The latter two projects are currently in the writing stage.

2. Dedeckera Rev. & J. T. Howell, Brittonia 28: 245. 1976.—TyPE: Dedeckera eurekensis Rev.
& J. T. Howell.

Large perennial shrubs with numerous spreading woody branches arising from a woody
taproot; /eaves cauline, alternate, exstipulate, petiolate, the leaf-blades elliptic, hirsutulous;
flowering stems slender, annually produced from the axils of the previous year’s leaves;
branches green and hirsutulous; inflorescences cymose, compact to rather open, with a single,
short-pedicellate flower and a single peduncle at each node, and a cluster of 5-10 sessile or
subsessile flowers atop the 2-5-bracted peduncle; bracts foliaceous, similar to the leaves only
reduced, 3-4 at the lowest node, 2-3 above, these ultimately scale-like; peduncles erect,
slender, restricted to the axils of the branches, hirsutulous; involucres reduced to a series of
2-5 bracts, the lower ones arranged in two whorls (3 plus 2), the upper ones reduced to 2
bracts only; flowers yellowish, sessile to subsessile or short-pedicellate, not stipitate,
hispidulous, the tepals 6, petaloid, slightly united basally, essentially monomorphic, narrowly
lanceolate with the outer whorl of 3 slightly broader and longer than the inner; stamens 9,

Reveal: Genera related to Eriogonum (Polygonaceae) 239

slightly exserted at anthesis, the filaments pilose basally, the anthers yellow, oblong; achenes
3-angled, light reddish-brown, slightly pubescent apically, the embryo curved, in abundant
mealy endosperm; n= ca 12 (Wiens et al. 1989).

A rare, monospecific genus endemic to the desert ranges of Mono and Inyo cos., California, in the White
Mountains and in the Last Chance Range south to the northern Panamint Range, reportedly in the Inyo Mountains.

Over the decade following the initial discovery of Dedeckera (named for Mary Caroline
DeDecker, 1909- , field botanist and noted California conservationist) in the Last Chance
Range (Reveal & Howell 1976), its distribution has been expanded to the north onto the
western slopes of the White Mountains and to the east onto the Panamint Range.

The original interpretation that Dedeckera lacked an involucre is now changed. The bracts
subtending the cluster of flowers (five at the lower nodes, two to three above) are here
considered to be involucral bracts similar to those found in Stenogonum. The lower ones,
for example, are arranged in two whorls of three subtending two. In the upper flowers the
bracts are reduced to just two and it appears that the outer whorl of three has been lost. Its
placement in the tribe Eriogoneae has not been altered, and it is considered to be derived
from a basal subgenus of Eriogonum and thus a member of the subtribe Eriogonineae. None-
theless, Dedeckera is a highly specialized paleoendemic member of the tribe and essentially
phylogenetically isolated from the remainer of the taxon. Its unusually low number of flowers
per inflorescence (which can be solitary in some instances) and a chromosome number of
about n= 12 adds to its isolation.

The genus may be readily recognized by its numerous dense inflorescence of involucrally
bracted, pale-yellow, clustered, densely hispidulous flowers, its hirsutulous vegetation and its
shrubby, perennial habit.

1. Dedeckera eurekensis Rev. & J. T. Howell, Brittonia 28: 246. 1976.—TyPE: Last Chance
Range, in a rocky canyon (now Dedeckera Canyon) ca 3 airline mi SE of Eureka Valley
sand dunes and 3.5 airline mi NW of Marble VABM 7559, in T10S, R40E, on steep
limestone, north-facing, rocky slopes associated with Atriplex, Eriogonum and Prunus at
about 4000 ft elev, Inyo Co., California, 29 Jul 1975, Reveal et al. 3909 (holotype: US!;
isotypes: ASU, B, BRY, CAS, DAO, DEDECKER, F, GH, JEPS, K, LA, LE, MARY,
MICH, MIN, MO, MONTU, NEB, NESH, NY, OKL, RENO, TEX, UT, UTC!).

Large, densely branched shrubs 0.2-0.7 (1) m tall, 0.5-2 m across, with numerous, gray,
woody branches arising from a stout, woody root; /eaves green to yellowish-green, hirsutulous,
the leaf-blade narrowly to broadly elliptic, (0.7) 1-1.5 cm long, (4) 5-8 (13) mm wide, thinly
pubescent on both surfaces, the hairs longer and more dense along the entire margin, the
apex acute to obtuse, the base tapering to a 2-5 mm long petiole; flowering stems greenish,
erect or nearly so, leafy, 2-7 (10) cm long, hirsutulous; branches numerous, mostly alternately
arranged; inflorescences densely cymose, 1-4 (6) cm long, mostly trichotomously branched
with a single, short-pedicellate flower and a single peduncle at each node; bracts of the in-
florescence foliaceous and similar to the leaves except reduced in size toward the apex, the
3-4 bracts of the lowermost node 0.5-1.2 (1.7) cm long, the 2-3 bracts of the upper nodes
much reduced and ultimately scale-like, up to 6 mm long; peduncles erect, slender, greenish,
the lower one 4-6 (7) mm long, the upper ones greatly reduced; involucres reduced to a series
of 2-5 involucral bracts, these lanceolate to broadly lanceolate, 1-2 mm long, 0.4-0.6 mm
wide, the 5 bracts common on all peduncles except for 2-3 on the uppermost peduncles, the
lower ones arranged in 2 whorls with the outer ones broader and longer than the inner ones;

240 PHYTOL O GIA volume 66(3):236-245 May 1989

flowers yellow at anthesis, becoming paler after fertilization and before darkening to a
reddish-yellow in fruit, 1.8-3 (3.5) mm long at anthesis, becoming 2.5-4 mm long in fruit,
sessile or subsessile on peg-like pedicels in clusters of 7-10 atop the lower peduncles and 5-7
at the tips of the ultimate peduncles, the single flower at the base of the peduncle on a short,
slender pedicel 0.3-1 mm long, the tepals narrowly lanceolate, the inner 3 slightly narrower
and shorter than the outer 3, thickened and keeled along the midrib, densely hispidulous
without, sparsely hairy within mainly along the midrib, united only at the very base; stamens
9, the filaments 1-2 mm long, pilose basally, the anthers oblong, 0.2-0.3 mm long; achenes
light reddish-brown, 2-3 (3.5) mm long, the narrowly globose, distinctly 3-angled base tapering
to a slightly hispid 3-angled beak; n= ca 12 (Wiens et al. 1989).

Local and infrequent on limestone talus slopes along the western slope of the White Mountains from Coldwater
Canyon in Mono Co. south to near the Poleta Canyon area of Inyo Co., disjunct to the Last Chance Mountains
mainly southeast of the Eureka Valley sand dunes and onto the northern ridges of the Panamint Range, and
reportedly in the Inyo Mountains, from 4000-7200 ft elev, flowering from Jun-Aug, fruiting until Oct.

Specimens Examined.- UNITED STATES. Ca.iFornia: Inyo Co.: unnamed canyon, E of E. Line Street in Bis-
hop, 20 May 1982, Chamberlain 458a (CAS); canyon 5 mi E of Bishop, 14 Jul 1982, Chamberlain 498a (CAS);
Last Chance Mountains, limestone canyon (“Dedeckera Canyon’) S of Eureka Dunes, 6 Jun 1976, C. Davidson
4249 (ENCB, F, RSA, SD); Last Chance Mountains, SE of Eureka Valley sand dunes, 4 Jul 1975, DeDecker 3892
(CAS, DEDECKER, RSA); Dedeckera Canyon, Last Chance Mountains, SE of Eureka Valley sand dunes, 10
May 1977, DeDecker 4400 (DEDECKER); Dedeckera Canyon, Last Chance Mountains, 3 mi SE of Eureka Valley
sand dunes, 9 May 1978, DeDecker 4657 (DEDECKER); Last Chance Mountains, 3 mi SE of Eureka Valley sand
dunes, 27 Jun 1978, DeDecker 4740 (CAN, DEDECKER, NY, SBBG); Last Chance Mountains, along the summit,
2.25 mi W of NW corner of Death Valley National Monument, 7 Jul 1978, DeDecker 4745 (DEDECKER, NY,
RSA); Last Chance Mountains, 3 mi SE of Eureka Valley sand dunes, 13 Sep 1978, DeDecker 4794 (CAS,
DEDECKER, GH, UC, UT); unnamed canyon E of Bishop, White Mountains, 24 Aug 1984, DeDecker 5700
(UNLV, UT); canyon E of Bishop on or above the Forest Service line, 26 Jun 1980, DeDecker & Strohm 5073
(DEDECKER, UC); Panamint Range, Eureka Valley drainage, E of jeep corridor from Saline Valley, 10 Aug
1980, Kay 5144 (DEDECKER, UT); S end of Eureka Valley, 5S May 1977, A. Major s.n. (UCR); White Mountains,
1.6 mi NW of Poleta Mine at the base of the canyon slope, 18 Jun 1984, Morefield 2118 (ASC, MNA, NY, RSA,
UNLV); White Mountains, 1.2 mi SE of Southern Belle Mine in Gunter Creek Canyon, 18 Jun 1984, Morefield
2124 (ASC, MICH, MNA, NESH, NY, RSA, UNLV); 0.7 mi up Dedeckera Canyon, 3 mi SE of the main Eureka
Valley Sand Dunes, 2 Jun 1987, Morefield & Ehrendorfer 4499 (MARY); NE flank of Dry Mountain, Last Chance
Range, 4 Nov 1982, Rowlands s.n. (UCR); 2.4 km NE of E. Line Street in Bishop, between Silver Canyon and
Poleta Canyon, 24 Jun 1980, Strohm 560 (UC); first canyon N from Poleta Canyon, White Mountains, 3 Jul 1981,
D.W. Taylor 7886 (UC); canyon 2 mi NE of the E end of E. Line Street in Bishop, 7 Aug 1983, Vasek & DeDecker
s.n. (UCR). Mono Co.: Gunter Canyon, White Mountains, 28 Jun 1984, DeDecker 5596 (DEDECKER, UNLV);
Coldwater Canyon, White Mountains, 5 Jul 1985, DeDecker 5821 (DEDECKER, SBBG, UNLV, UT); White Moun-
tains, 1 km up Coldwater Canyon, near the water intake, 6 Jul 1985, DeDecker & Wiens 5981 (CPH, NY, UT);
Cold Water Canyon, White Mountains, 5 Jul 1987, Elias & Morefield 10369 (RSA); White Mountains, Coldwater
Canyon, 15 Jun 1985, Giuliani s.n. (UT); Lower Coldwater Canyon, White Mountains, 24 Aug 1984, Junak &
DeDecker 2264 (SBBG, UNLV); White Mountains, 3.5 mi up Coldwater Canyon, 19 Jun 1984, Morefield 2126 (ASC,
MICH, NY, RSA); White Mountains, 0.5 mi up Coldwater Canyon, 19 Jun 1984, Morefield 2132 (ASC, MICH,
NESH, NY, RSA).

The geographical range of Dedeckera, as predicted originally (Reveal & Howell 1976), has
expanded beyond the narrow confines of a single canyon on the western slope of the Last
Chance Range. That canyon, now officially called “Dedeckera Canyon,” is still the home of
a large population. Novak and Strohm (1981) reported the species from the Poleta Canyon
area of the White Mountains in Inyo County, with Morefield (1985) extending the range
northward to the Coldwater Canyon area in Mono County. In his note, Morefield reported
Dedeckera from the Panamint Range but did not cite specimens. He learned of the range
extension from Mary DeDecker who saw shrubs while flying in a helicopter. Since then, one

Reveal: Genera related to Enogonum (Polygonaceae) 241

of her associates has managed to document one of the sites and it is reported above.

More recently (Wiens et al. 1986) observed pollination in Dedeckera. They described the
normal maturation of flowers as it occurs in the majority of the Eriogonoideae, which is
protandrous with the anther releasing pollen typically a full day before the stigma is receptive.
Cross-pollination is possible during the second day as the open flowers display the anthers
in an outward array away from the erect stigmas. That night, however, when the flowers
close, the anthers come in direct contact with the stigma, and if pollen grains are still present,
self-pollination may occur. It is not uncommon for the tepals in the Eriogoneae to change
in color after fertilization (a feature seen in Dedeckera), following which the sepals then
tightly enclosing the maturing achene.

Seed production is exceedingly low according to Wiens and his colleagues (1989) and they
reported that successful seed set is only 2.5%. The cause of the low seed set is probably
“genetically mediated embryonic abnormalities.” This is reduced even further by low viability,
poor germinability and post-germination development failure. Nonetheless, populations of
Dedeckera show a high degree of heterozygosity (27%) probably due to the extreme habitats
where the species occurs. These researchers report that individual shrubs are long lived (140
years or more) and that the vegetative fitness seen is due to the highly heterotic genotypes
found both intra- and interpopulationally.

The genetic factors associated with the low successful seed set in Dedeckera (Wiens et al.
1989) might be augmented by environmental conditions. Given the large number of flowers
produced on each shrub (several thousand each year), the relatively large number of shrubs
in most of the populations, and the restricted habitat where this plant can exist, it might be
that a seed set of 2.5% is actually sufficient for this calcareous plant to survive long term.
The arid habitat where Dedeckera is found is only rarely subjected to natural fire, flash flood-
ing or other natural events so that given the long life of an individual shrub, dramatic pop-
ulation losses might well be rare. Also, consider the numbers. If an individual shrubs produced
1000 flowers year, and an individual lived an average of 100 years, then over that time some
100,000 flowers would have been formed. If only 2.5% of these set seed, some 2500 seeds
would have been formed. Assuming only a ten percent survivability (250 seeds left) and of
those only one percent grew into a plant of reproductive age, then there would be a positive
replacement rate of two and a half plants per existing plant. Given the nature of the desert,
this rate of long-term reproduction is not unacceptable. The actual number of flowers
produced per mature shrub is probably closer to 10,000 per plant than 1000 as used here. It
is proposed that when the reproductive biology of other long-lived desert shrubs is inves-
tigated low seed set in such species will not be found to be rare.

Nonetheless, from these studies, it is clear that the original “home” of Dedeckera was likely
in an environment unlike that where it is found presently. If the genus arose from some early
ancestral basal group within Eriogonum, then that original habitat was probably more akin
to that found presently in the northern part of the Intermountain Region or in the moun-
tanous regions of central Idaho.

Wiens et al. (1986) observed several insect visitors on the flowers of Dedeckera and several
were found with pollen. They concluded that sarcophagid flies and other short-tongued
scavenger flies “are probably the most numerous insect visitors that could effect pollination.”
Secondary pollinators are wasps and syrphid flies. Similar observations and conclusions have
been reached on pollination of Enriogonum, although ants are a significant factor in alpine
situations.

242 PHYTOL O GIA volume 66(3):236-245 May 1989

3. Stenogonum Nutt.

Since this genus was reestablished (Reveal & Ertter 1977a), Stenogonum flexum has been
found in New Mexico (Spellenberg & Corrall 8222 (NMC)). Additional isotypes of S. fleaum
(M.E. Jones s.n., near Cameron, Coconino Co., Arizona, 10 Jun 1890) have been seen at DS,
F, G and US!

4. Oxytheca Nutt.

The revision of this genus by Ertter (1980) is excellent and only a few additional notes
relating to the distribution of type material is presented here.

Additional isotypes of Oxytheca dendroidea var. hillmanii (M.E. Jones 4044), a synonym of
subsp. dendroidea, were found at BR, DS, LE and SBBG! Isotypes of Brisegnoa chilensis,
the basionym of O. dendroidea subsp. chilensis, beyond those reported by Ertter (1980) were
seen at B, BR, G, and LE! Two additional isotypes of the var. tonsiflora are at B and BA!
The type of O. parishii is the widely distributed Parish & Parish 993; the following additional
isotypes were found: BR, CM, G, LE, MIN, NEB, NDG and WS!

The isotypes of Oxytheca parishii var. cienegensis, Reveal & Reveal 4781, have only recently
been distributed to ARIZ, B, BM, BRY, CAS, COLO, F, GH, K, LA, MARY, MICH, MO,
OSC, RM, RSA, TEX, US, UTC and WIS. The holotype is at NY! Additional isotypes of
the var. goodmania, Parish & Parish 1241, were noted at BR, G, ISC, LE, WS and WU!

Two additional isotypes of Oxytheca abramsii, the basionym of O. parishii var. abramsii,
were found at E and G, with one isotype of O. trilobata seen at ISC! A second isotype of
O. emarginata was seen at DS!

Since Ertter (1980) published her thesis, specimens of Oxytheca watsonii have been found
in Inyo Co., California, all from near Santa Rosa Hills near the junction of California Highway
190 and Saline Valley Road: Dedecker 5337 and 5932 (DEDECKER); Dedecker et al. 5226
(DAV, DEDECKER); and Stone 316 (CAS, MARY, RSA).

Ertter (1980) discussed the problems with the type of Oxytheca caryophylloides noting that
two different dates are associated with Parish & Parish 1097. During the present study two
additional labels were found. Ertter properly concluded that the one dated simply “Aug 1881”
must be considered the type. The four variations are: San Bernardino Mountains, Aug 1881,
Parish & Parish 1097 (CM, F, G, GH, ISC-holotype, MIN, MPU, NY, PH, RSA, US, W,
WU); 15 Aug 1882 (BM, DS, F, ISC, MIN, NY, P, PH, UC, US, VT, WS); Aug 1882 (B,
BR); without a date (NDG).

5. Goodmania Rev. & Ertter

Ertter (1981) corrected my mistake of associating the wrong herbarium abbreviation with
Parry’s types (ISC, not IA). Since our segregation of Goodmania from Oxytheca, the following
collections have come to my attention, bring the total number of known collections to 72.

UNITED STATES. CALIFORNIA: Inyo Co.: E end of Twenty Mule Team Canyon, Furnace Creek Wash, Death
Valley National Monument, 26 Apr 1978, J.T. Ball 8-228 (UT). Kern Co.: Lancaster Boulevard, S of Rosamond,
28 Apr 1974, T. Gordon 214 (DAV). Los Angeles Co.: 1 mi W of California Highway 14 along California Highway
138, 18 Apr 1980, Keil & Holland s.n. (OBI); Antelope Valley, along G Street 0.2 mi E of 60th Avenue, 17 May
1988, Reveal 6797 (MARY); Antelope Valley, along G Street at 40th Avenue, 17 May 1988, Reveal 6799 (BM,
BRY, CAS, MARY, MO, NY, OSC, RM, RSA, US, UTC, WIS); corner of G and Division Street W of California
Highway 14 in Lancaster, 13 Apr 1988, Villasenor s.n. (MARY). NEVADA: Mineral Co.: Alkali Valley, N side of

Reveal: Genera related to Eriogonum (Polygonaceae) 243

Alkali Lake, 12 Jul 1983, Tiehm & Lavin 8140 (ASU, MIN, NEB, NESH, NY, RSA, UT, UTC).

The range extension reported by Tichm (1984) is the most important addition. According to
a note, Samuel B. Parish stated that “Parry’s type was collected in the drip of the railroad
water tank at Lancaster, Antelope Valley.” The GH isotype is dated Sep 1881 while the ISC
holotype is dated 188- and bears a label with “Gymnogonum spinescens, Parry n. gen.” on
it. An isotype is at CAN as well as at ISC.

6. Gilmania Cov., J. Wash. Acad. Sci. 26: 210. 1936. Phyllogonum Cov., Contr. U.S. Natl.
Herb. 4: 190. 1893, non Bridel (1827).—TyPE: Phyllogonum luteolum Cov.

Spreading annual herbs with several stems arising from a thin taproot; /eaves basal and
cauline, laminar and arranged in threes throughout, exstipulate, narrowly petiolate below,
becoming sessile above, the leaf-blade oblong to broadly elliptic or obovate, glabrous or with
only a few scattered hairs mainly on the entire margin; flowering stems mostly trichotomous
branched; branches slender, yellowish-green, glabrous or thinly pubescent with slightly curled
hairs; inflorescences cymose, dichotomously branched with a cluster of flowers at each node,
the secondaries often suppressed; bracts lacking; peduncles lacking; involucres lacking; flowers
on elongated pubescent non-stipulate pedicels, yellow, thinly pubescent with long curly hairs,
the tepals 6, petaloid, lanceolate, united only at their very base; stamens 9, mostly included,
the filaments yellow, pilose only at the very base, the anthers yellow, oval; achenes brownish,
3-angled, glabrous, with a short, stout beak, the embryo curved, in abundant mealy en-
dosperm.

A narrowly restricted monospecific genus known only from the Death Valley area in Death Valley National
Monument, Inyo Co., California, from -50 ft below sea level to 1500 ft elev; flowering from Feb-Apr (May).

When initially described by Coville (1893), he knew the genus only from two collections: a
type that consisted of two individuals, and a second collection of a similar size. When he
realized Phyllogonum (Greek phyllon, leaf, and gony, knee, alluding to the cauline leaves)
was a homonym and proposed Gilmania (M. French Gilman, 1871-1944, Death Valley
naturalist), Coville (1936) concluded that Gilmania was a genus that likely had a limited
evolutionary future. He did recognize that individuals of Gilmania managed to flower and
fruit every year, but that yearly, millions of seeds were lost from the seedbank. While this is
likely true (albeit to a lesser degree than suggested by Coville), there is no doubt that when
conditions are right, hundreds of thousands of individuals of Gilmania carpet the foothills
surrounding Death Valley, each in turn producing thousands of seeds. In this fashion, millions
of seeds are reintroduced into the seedbank.

The origin of Gilmania must lie within the confines of Eriogonum subgenus Ganysma.
There is no modern species or species complex within that subgenus, however, that appears
to be its point of origin. It is suggested that Gilmania and Goodmania may have come from
a closely related complex, but what, if any relationship might exist between the two is unclear.

1. Gilmania luteola (Cov.) Cov., J. Wash. Acad. Sci. 26: 210. 1936. Phyllogonum luteolum
Cov., Contr. U.S. Natl. Herb. 4: 190. 1893. Eriogonum luteolum (Cov.) M.E. Jones, Con-
tr. W. Bot. 11: 15. 1903, non E. Greene (1896).—TYPE: Furnace Creek Canyon, Funeral
Mountains, Death Valley National Monument, Inyo Co., California, 7 Apr 1891, Coville
584 (holotype: US!).

244 PHYTOLOGIA volume 66(3):236-245 May 1989

Spreading annual herbs 3-12 (15) cm high; /eaves broadly elliptic to obovate, 0.5-1.5 cm
long, 3-8 (10) mm wide, glabrous or sparsely pubescent with long hairs at least on the lower
surface or near the point of attachment on the petiole, the petiole glabrous or thinly pubes-
cent, 0.5-2 cm long in the basal rosette, 0.3-1.5 cm long at the lower nodes and gradually
becoming sessile in the uppermost clusters, expanding basally and clasping the branch, the
3 petiole bases fused around the branch; branches slender, yellowish-green, glabrous or
sparsely pubescent; inflorescences cymose and often with the secondaries suppressed, with a
cluster of flowers at the node, becoming rather densely clustered apically with nearly sessile
leaves and flowers; flowers on long, slender thinly pubescent pedicels 2-5 (7) mm long, yellow,
1-1.5 mm long in anthesis, 1.5-2 mm in fruit, thinly pubescent without mainly along the
obscure midrib and margin, densely so within at the point of fusion of the tepals forming a
small disklike nectary at the base of the filaments, the tepals lanceolate, all of equal size at
anthesis, the inner three elongating in fruit, united only at the very base; stamens 9, the
filaments 1-1.2 mm long, pilose basally, the anthers oval, 0.2-0.3 mm long; achenes yellowish-
brown to reddish-brown, shining, glabrous, 1.5-2 mm long, the globose base tapering gradually
to a stout, slightly 3-angled beak.

Locally rare to common on alkaline, often barren slopes and flats in Death Valley, Death Valley National
Monument, Inyo Co., California, from -S0 ft below sea level to 1500 ft elev, flowering from February-April (May).

Specimens Examined.- UNITED STATES. Catirornia: Inyo Co., Death Valley National Monument: Funeral
Mountains, 15 mi SE of Travertine Springs, 27 Apr 1983, Annable 609 (RSA, UNLV); mud hills S of Golden
Canyon, 2 Apr 1980, Armstrong s.n. (RSA, SD); Furnace Creek, 1940, Carpenter s.n. (JEPS, SD); Death Valley, 17
Apr 1939, Clokey 5852 (ASU, BKL, BM, BR, BRY, CAN, CAS, CM, DS, F, G, GB, GH, IDS, ISC, K, MARY,
MICH, MIN, MO, MONTU, NO, NY, OKL, ORE, OSC, RENO, RM, RSA, SD, SMU, TEX, UC, US, USFS,
UT, UTC, W, WILLU, WIS, WS, WTU); Furnace Creek, 2.5 km E of Furnace Creek Ranch, Funeral Mountains,
7 Apr 1891, Coville & Funston 584 (DS, US); Death Valley, Golden Canyon, 27 Mar 1940, Eastwood & Howell
7720 (CAS); Volcanic Drive, 27 Mar 1940, Eastwood & Howell 7726 (CAS, DS, F, GH, NY); foot of Furnace
Mountains, S of Furnace Creek, 20 Apr 1935, Gilman s.n. (US); foot of Furnace Mountains, near mouth of Golden
Canyon, 20 Apr 1935, Gilman 1375 (US); near mouth of Golden Canyon, 20 Apr 1935, Gilman 1377 (RSA); hill
E of Teck Springs, 27 Apr 1935, Gilman 1424.5 (DS); Death Valley, Artists Drive, 4 May 1935, Gilman 1520 (US);
Golden Canyon area, 10 Dec 1939, Gilman 4005 (CAS, POM); Artists Drive, 10 Dec 1939, Gilman 4006 (CAS,
POM); Artists Drive near the turnoff to Artists Palette, Black Mountains, 25 Apr 1983, Gustafson & Herbst 2531
(RSA); Artists Drive, Black Mountains, 16 Mar 1973, Holmgren & Holmgren 6309 (BRY, NY, UTC); 20 Mule
Team Canyon, 17 Mar 1940, Hood 40-20k (LA); small canyon E of Furnace Creek, 26 Mar 1939, Jaeger s.n. (DS);
Furnace Creek, Funeral Mountains, 2 May 1917, Jepson 6921 (JEPS); Artists Drive, 6 mi S of Furnace Creek, 24
Mar 1947, Keck 5740 (DS, UC); Volcanic Drive, 23 Feb 1940, Leach s.n. (ORE); Golden Canyon, 3 Apr 1939, H.
Lewis s.n. (LA); Gold Canyon, 24 Apr 1969, Myrick 2091 (SBBG); S of Furnace Creek Ranch, 4 Apr 1939, Nelson
& Nelson 3443 (DS, RM, UC); Furnace Creek, 18 May 1915, Parish 10008 (GH, MO, NY, RSA, SBBG, UC, US,
USFS); along California Highway 190, 0.2 mi N of Golden Canyon Road, 12 Apr 1968, Pinkava et al. 12734 (ASU);
Breakfast Canyon, 30 Apr 1976, Reveal 4368 (CAS, GH, MICH, MO, OKL, TEX); Desolation Canyon, 30 Apr
1976, Reveal 4369 (CAS, GH, MO, TEX); Artists Drive, Funeral Mountains, 19 Apr 1940, Ripley & Howell 2955
(NY); Zabriskie Point, 20 Apr 1947, Roos 3502 (UCR); 2 mi above Zabriskie Point, 30 Mar 1976, Schramm et al.
233 (UCR, UNLV); Golden Canyon, 10 Apr 1940, Shanteau 28 (ORE, UC); near Artists Palette, 30 Apr 1976,
Tiehm & Mason 2034 (RENO); Artists Drive, Black Mountains, 18 Apr 1973, Trowbridge 3191 (SFSU); Zabriskie
Point, 5 Apr 1983, Welsh & Welsh 21505 (BRY); Gowers Gulch, Corkscrew Canyon, 10 Apr 1980, Welsh et al.
19422 (BRY, NY); Artists Drive Road, Black Mountains, 18 Apr 1973, Wester 302 (WIS).

Gilmania luteola, often called “golden carpet,” is a reliable indicator of precipitation in the
Death Valley area. In a “good” year the slopes and canyons of the Funeral Mountains are
in fact carpeted with individuals of Gilmania.

Reveal: Genera related to Eniogonum (Polygonaceae) 245

7. Nemacaulis Nutt.

No significant new data needs to be added to the earlier treatment of this genus (Reveal
& Ertter 1977b) although numerous collections have been added to institutional holdings so
that some 270 collections are now known. Additional isotypes of Nemacaulis denudata var.
gracilis were found at BM, K, MIN, MSC, OKL, and US! The LAM isotype is now at RSA.

AKNOWLEDGMENTS

I wish to thank the curators and staff of the following herbaria for loans or for arranging
access to their collections during visits: A, ARIZ, ASC, ASU, B, BKL, BM, BR, BRY, CAN,
CAS, CGE, CHSC, CM, COLO, CPH, CS, DAO, DAV, DEDECKER, DS, E, ENCB, F,
G, GB, GH, GOET, HSC, ID, IDS, ILL, IND, ISC, JEPS, K, KANS, KSC, L, LL, M, MARY,
MICH, MIN, MNA, MO, MONTU, MPU, MSC, ND, NDG, NEB, NESH, NMC, NO, NTS,
NY, OBI, OKL, OKLA, ORE, OSC, OXF, P, PENN, PH, POM, RENO, RM, RSA, SBBG,
SD, SFSU, TCD, TEX, UC, UCR, UCSB, UNLV, US, USFS, UT, UTC, VT, W, WILLU,
WIS, WS, WTU, WU, and Z. Individuals wishing a complete list of herbarium specimens
examined may request a copy. Funding for field work has been made possible through a
variety of sources. I am grateful to Dr. Edward E. Terrell and Clare B. Hardham for their
comments. This is Scientific Artiele A-4939, Contribution 7982, Maryland Agricultural Ex-
periment Station.

LITERATURE CITED

Coville, C. V. 1893. Botany of the Death Valley expedition. Contr. U.S. Natl. Herb. 4: 1-363.
. 1936. Gilmania, a new name for Phyllogonum, a very rare genus of plants from Death Valley, Cali-
fornia, apparently in process of extinction. J. Wash. Acad. Sci. 26: 209-213.
Ertter, B. J. 1980. A revision of the genus Oxytheca Nutt. (Polygonaceae). Brittonia 32: 70-102.
. 1981. Notes on Goodmania and Oxytheca (Polygonaceae: Eriogonoideae). Brittonia 33: 37-38.
Jones, M. E. 1903. Eriogonum. Contr. W. Bot. 11: 4-18.
Michaux, A. 1803. Flora boreali-americana. 2 vols. Published by the author, Paris.
Morefield, J. D. 1985. Noteworthy collections. Madrofio 32: 122-123.
Novak, P. J. & KL. Strohm. 1981. Noteworthy collections. Madrofio 28: 86-87.
Reveal, J. L. 1989a. A review of the genus Harfordia (Polygonaceae: Eriogonoideae). Phytologia 66: 221-227.
. 1989b. Remarks on the genus Pterostegia (Polygonaceae: Eriogonoideae). Phytologia 66:228-235S.
. 1989c. Notes on selected genera related to Chorizanthe (Polygonaceae: Eriogonoideae). Phytologia 66:
199-220.
. 1989d. The eriogonoid flora of California (Polygonaceae: Eriogonoideae). Phytologia 66: 295-414.
& B. J. Ertter. 1977a. Re-establishment of Stenogonum (Polygonaceae). Great Basin Naturalist 36:
272-280.
. 19770. Goodmania (Polygonaceae), a new genus from California. Brittonia 28: 427-429.
. 1980. The genus Nemacaulis (Polygonaceae). Madrofo 27: 101-109.
& C. B. Hardham. 1989a. Three new monospecific genera of Polygonaceae subfamily Eriogonoideae
from California. Phytologia 66: 83-88.
. 1989b. A revision of the annual species of Chorizanthe (Polygonaceae: Eriogonoideac). Phytologia
66: 98-198.
Reveal, J. L. & J. T. Howell. 1976. Dedeckera (Polygonaceae), a new genus from California. Brittonia 28: 245-251.
Roberty, C. & S. Vautier. 1964. Les genres de Polygonacees. Boissiera 10: 7-128.
Tiehm, A. 1984. Noteworthy collections. Madrono 31: 192-193.
Wiens, D., M. DeDecker & C. D. Wiens. 1986. Observations on the pollination of Dedeckera eurekensis
(Polygonaceae). Madrofio 33: 302-305.
, D. L. Nickrent, C. I. Davern, C. L. Calvin, & N. J. Vivrette. 1989. Developmental failure and loss
of reproductive capacity in the rare palaeoendemic shrub Dedeckera eurekensis. Nature 338: 65-67.

Phytologia (May 1989) 66(3):246-248

A NEW VARIETY OF ERIOGONUM NUDUM (POLYGONACEAE:
ERIOGONOIDEAE) FROM THE SOUTHERN SIERRA NEVADA

JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

AND
JOHN C. STEBBINS AND JOANNA M. CLINES

Department of Biology, California State University
Fresno, California 93740-0073

ABSTRACT

A new variety of Eriogonum nudum Douglas ex Benth. (subg. Eucyla), the var. regirivum, is
proposed for a plant found on limestone slopes along Kings River in Fresno Co.. It may be
recognized by its strict habit, densely tomentose stems and branches, and pubescent creamy
white flowers arranged in 3-6 congested involucres disposed at the upper nodes.

KEY WORDS: Polygonaceae, taxonomy, Eriogonum, California.

INTRODUCTION

Soon after Munz and Reveal (Munz 1968) revised the California species of Eriogonum it
became obvious that additional expression within E. nudum Douglas ex Benth. deserved
recognition. In 1970, two narrowly endemic varieties were added to the species, the var.
decurrens (S. Stokes) Bowerman of Santa Cruz Co. and the var. murinum Rev. of Tulare Co.
During this latter review, an additional variant was noted from Fresno Co. that seemed worthy
of recognition, but was then known only from a single collection gathered at a site that was
extirpated by the construction of Pine Flat Reservoir. Additional collections of the taxon
were seen during herbarium visits in the 1970s, but all seemed to be from the same general
area covered by the Reservoir. Various collectors interested in the southern Sierra Nevada
were told of the variant but none was able to find the plant. In 1987, James R. Shevock told
the senior author that he was aware of a new variety of Eriogonum nudum in the southern
Sierra Nevada which flowered late in the year, was markedly distinct, but one he had not yet
been able to collect. Interestingly, this proved to be the same plant that the junior authors

were examining in the field. Specimens were gathered in 1988 for a type collection and the
taxon is now established.

246

Reveal, Stebbins & Clines: A new variety of Eriogonum nudum (Polygonaceae) 247

TAXONOMY

Eriogonum nudum Douglas ex Benth. var. regirivum Rev. & J. Stebbins, var. nov.— TYPE:
UNITED STATES. CALIFORNIA: Fresno Co.: 0.75 mi E of road’s end, Kings River
Canyon, Sierra National Forest, T12S, R27E, NWS26, 487 m elev, 17 Sep 1988, Stebbins
et al. 88-106 (holotype: US!; isotypes: BM, CAS, FSC, MARY, MO, NY, RM, RSA,
UC).

A var. decurrens foliis basi et var. oblongifolium floribus dense pubescens et foliis longior
differt.

Plants tall strict herbaceous perennials (5) 8-14 (18) dm high; /eaves sheathing slightly up
the stem but not at the nodes, elliptic to oblong, the leaf blade 2.5-4.5 (5.5) cm long, (1.5)
2-3.5 cm wide, densely white-tomentose below, grayish-tomentose to floccose or subglabrous
and greenish above, the margins slightly crisped, the tomentose to floccose petiole (2) 4-6
(7) cm long; flowering stems erect, slender, 1-3 (4) dm long, lanate to tomentose; inflorescences
cymose, strict, 3-10 (14) dm long, the branches tomentose; bracts scalelike, ternate, linear to
triangular, mostly 1-3 (5) mm long, densely tomentose except for the very tip, connate basally;
peduncles lacking; involucres congested, 3-6 per head, turbinate, 3-4 mm long, thinly tomen-
tose, the 5-8 erect teeth 0.5-0.7 mm long; flowers creamy white, 1.5-3 mm long, densely pubes-
cent without, less so but still distinctly pubescent along the midrib within, the tepals
monomorphic, oblong, the outer tepals shorter than the inner ones at maturity; stamens
exserted, 2.5-3.5 mm long, the filaments pilose basally, the anthers 0.4-0.5 mm long, white
with a reddish tinge to rose; achenes light brown, 3-3.5 mm long, glabrous; n= 20.

Specimens Examined. - UNITED STATES. Ca.irornia: Fresno Co.: Pine Flat, 6 mi below Trimmer, Kings River,
27 Sep 1942, Carter 39 (UC); 0.75 mi E of road’s end, Kings River Canyon, 15 Nov 1988, Clines 201 (CAS, FSC,
MARY, RSA, US), voucher for chromosome count; Dunlap, 19 Oct 1940, Hoover 4691 (UC, US); Kings River
Canyon, 26 May 1988, Shevock 11820 (CAS, FSC, MARY, RSA, UC).

Eriogonum nudum var. regirivum (Latin regis, king, and rivus, river, stream, for the Kings
River) is a localized endemic restricted to limestone slopes within the Digger Pine-Oak
Woodland community (Holland 1986) which prevails in the Kings River Canyon at lower
elevations. The type collection was obtained from steep unstable slopes that support few
woody species. These slopes are mostly south-facing and are characterized by loose calcium
carbonate soils interspersed with marble outcrops. The most common associates are Lupinus
albifrons, Artemisia dracunculus, Stephanomeria virgata and Eriogonum roseum.

The var. regirivum may be readily distinguished from the Santa Cruz Co. endemic, var.
decurrens, by its pubescent flowers and basally sheathing elliptic to oblong leaves. In var.
decurrens the roundish leaves are only 1-3 cm long and are frequently found at well defined
nodes. Probably the most closely related variant is the more northern var. oblongifolium S.
Wats. which occurs in the Sierra Nevada as far south as Tuolumne Co. It has white or yellow
flowers that are somewhat less pubescent than those of the new variety, and while the habit
is erect in the var. oblongifolium, it is not as strict as that in var. regirivum. Further, the
tomentum on the stems and branches of the new variety is dense to the point of being nearly
lanate, and while the tomentum of the stems and branches of var. oblongifolium may be
dense, it may only be floccose in some populations. Unlike the var. oblongifolium which
flowers mainly from June to September, the var. regirivum does not start to flower until
September and may continue well into November.

248 PHYTOL O GIA volume 66(3):246-248 May 1989

Cytologically, the new variety is a tetraploid (n= 20) and is therefore similar to such
variants as var. nudum, pauciflorum S. Wats., oblongifolium and others. Only the var.
auriculatum (Benth.) Tracy ex Jepson and its near relative, var. indictum (Jepson) Rev. in
Munz are octoploids (Stokes & Stebbins 1955).

ACKNOWLEDGMENTS

We acknowledge the assistance of John Lorenzana of the U.S. Forest Service, Sierra National
Forest and James R. Shevock of the regional office in San Francisco. We appreciate the
comments of Dr. Edward E. Terrell, Dr. James R. Shevock and Clare B. Hardham on this

paper. This is Scientific Article A-4940, Contribution No. 7983, Maryland Agricultural Ex-
periment Station. .

LITERATURE CITED

Holland, R. F. 1986. Preliminary descriptions of the terrestrial natural communities of California. California De-
partment of Fish and Game, Sacramento.

Munz, P. A. 1968. Supplement to A California Flora. Univ. California Press, Berkeley.

Reveal, J. L. 1970. Additional notes on the California buckwheats (Eriogonum, Polygonaceae). Aliso 7: 217-230.

Stokes, S. G. & G. L. Stebbins. 1955. Chromosome numbers in the genus Eriogonum. Leafl. W. Bot. 7: 228-233.

Phytologia (May 1989) 66(3):249-250

A NEW VARIETY OF ERIOGONUM PRATTENIANUM (POLYGONACEAE:
ERIOGONOIDEAE) FROM THE SOUTHERN SIERRA NEVADA

JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

JAMES R. SHEVOCK

Department of Botany, California Academy of Sciences
San Francisco, California 94118-4599

ABSTRACT

A new variety of Eriogonum prattenianum Durand (subg. Oligogonum), the var. avium, is pro-
posed for plants found on subalpine granitic rock outcrops in the southern Sierra Nevada. It
differs from the more northern, lower elevation, generally larger var. prattenianum of volcanic
flats and slopes in having subglabrate upper leaf surfaces.

KEY WORDS: Polygonaceae, taxonomy, Eriogonum, California.

INTRODUCTION

In 1968, Munz and Reveal (Munz 1968) noted the existence of an isolated high elevation
population of Eriogonum prattenianum in Fresno Co., California. At that time the plant was
known only from a single collection. In 1981, Tim McLaughlin, then with Kings Canyon
National Park, recollected the plant at Kettle Dome overlooking the Middle Fork of the
Kings River, but its significance was not immediately noted. In 1983, Larry L. Norris, also
with the Park Service, reexamined the McLaughlin collection and concluded that it
represented the southern expression of E. prattenianum. At that time he informed the junior
author and in 1984 Shevock and McLaughlin hiked to Kettle Dome to collect adequate ma-
terial for a proper analysis. In the intervening years the senior author found additional her-
barium material indicating that the plant occurs in Madera Co.

TAXONOMY

Eriogonum prattenianum Durand var. avium Rev. & Shevock, var. nov.-TYPE: UNITED
STATES. CALIFORNIA: Fresno Co.: East face of Kettle Dome overlooking Tehipite Val-
ley, Kings Canyon National Park, T19S, R29E, SEYANW%S26, 9400 ft elev, 5 Aug 1984,
Shevock & McLaughlin 10977 (holotype: US!; isotypes: BM, CAS, GH, MARY, MO,
NY, RM, RSA, UC!).

A var. prattenianum foliis subglabris super differt.

249

250 PHYTOLOGIA volume 66(3):249-250 May 1989

Plants low rounded subshrubs, 1-2 dm high and 1.5-3 dm across; leaves elliptic, 0.5-1 cm
long, 0.2-0.5 cm wide, densely white-tomentose below, subglabrous and bright green above,
the petiole 0.1-0.4 cm long; flowering stems erect, slender, 3-7.5 cm long, thinly pubescent,
bracteated about midlength; inflorescences capitate, 1-1.5 cm across; bracts restricted to the
middle part of the stem (technically at the apex of the stem and base of the peduncle), 4-6,
foliaceous, narrowly elliptic, 3-8 mm long, 1.5-3 mm wide, tomentose below, subglabrous and
green above; peduncles erect, slender, 2.5-5 (6) cm long, thinly tomentose; involucres solitary,
campanulate, the tube 3-4 mm long, 3-5 mm wide, thinly tomentose, the 8-10 reflexed lobes
2-3 mm long; flowers yellow, 3-6 mm long including the 1-1.5 mm long stipe, glabrous, the
tepals spatulate to obovate; stamens exserted, 3-4 mm long, the filaments pilose basally, the
anthers 0.5-0.6 mm long, yellow; achenes light brown, 4-5 mm long, glabrous.

Specimens Examined.- UNITED STATES. Ca.iFornia: Fresno Co.: 7 mi E of Hume Lake in the Lockwood
Grove area, Sequoia National Forest, 8 Aug 1963, Breedlove 5730 (DUKE, US); Tehipite Valley, 6-10 Jul 1900,
Hall & Chandler 512 (DS, MIN, MO, UC); E face of Kettle Dome, 13 Aug 1981, McLaughlin 002 (THRI); W face
of Kettle Dome, John Muir Wilderness, Sierra National Forest, 5 Aug 1984, Shevock & McLaughlin 10973 (CAS,
FSC, MARY, RSA, US). Madera Co.: Iron Creek, Sierra National Forest, 6 Aug 1951, Raven 3807 (CAS).

Eriogonum prattenianum var. avium (Latin avius, out of the way, alluding to the isolated lo-
cation of the new variety) is lithospecific to granitic rock outcrops where, on Kettle Dome,
it is found with Cryptogramma acrostichoides, Cystopteris fragilis, Pallaea bridgesii, Arctosta-
phylos nevadensis, Cassiope mertensiana, Chrysolepis sempervirens, Eriogonum wrightii var.
subcaposum, Heuchera rubescens subsp. alpicola, Holodiscus dumosus, Juncus parryi,
Penstemon newberryi, Sedum obtusatum, Senecio fremontii, Spiraea densiflora and Streptanthus
tortulosus. Habitats similar to that at Kettle Dome are widely scattered and access rather
limited so that the distribution of var. avium may be greater than indicated here although
no new populations have been found in the vicinity of Kettle Dome.

The new variety differs from the more northern and lower elevation var. prattenianum in
having subglabrous and bright green upper leaf surfaces and a compact, rounded habit. In
the strict sense the var. prattenianum is a large upright subshrub mostly 3-5 dm high with
stout branches terminated by a large terminal capitate inflorescence. This expression is found
mainly in Nevada and Placer cos. In Calaveras, Tuolumne and Mariposa cos. is a decidedly
more matted herbaceous expression with slender branches and smaller inflorescences.

ACKNOWLEDGMENTS
We wish to thank Dr. Edward E. Terrell and Clare B. Hardham for reviewing the paper.
This is Scientific Article A-4941, Contribution No. 7984, Maryland Agricultural Experiment

Station.

LITERATURE CITED

Munz, P. A. 1968. Supplement to A California Flora. Univ. California Press, Berkeley.

Phytologia (May 1989) 66(3):251-265

NEW COMBINATIONS AND NOVELTIES IN ERIOGONUM
(POLYGONACEAE: ERIOGONOIDEAE)

JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

ABSTRACT

Several new species, varieties and combinations are proposed in the genus Eriogonum. Three
combinations are proposed: E. thompsonae vat. matthewsae and E. lonchophyllum var. intermon-
tanum for narrowly restricted Utah endemics, and E. lonchophyllum var. fendlerianum for a plant
of southern Colorado, northern New Mexico and extreme northern Texas. Eriogonum ver-
rucosum, E. meledonum, E. capistratum (including the newly proposed var. muhlickii and var.
welshii) are new species of cespitose yellow flowered perennials related to E. chrysops Rydb. in
the subg. Eucycla. All are narrowly distributed in Idaho and western Montana. New varieties
are described in E. ochrocephalum (the var. sceptrum), E. shockleyi (the var. packardae) and E.
ovalifolium (the var. pansun); all are members of the subg. Eucycla and are found only in Idaho,
save the latter which is in adjatent southwestern Montana as well. In California, and in some
cases in adjacent Oregon as well, are new varieties of E. (subg. Eucycla) nudum (the coastal
var. paralinum) and three new varieties of E. (subg. Oligogonum) umbellatum all of serpentine
places in the northern Coast Ranges (var. goodmanii, var. humistratum and var. argus). Addi-
tional varieties of annual species (subg. Ganysma and Oregonium), all from California, are pro-
posed in E. trichopes (the var. hooveri) of the southern Coast Range, in E. deflecum (the var.
rectum of the Mojave and Sonora deserts previously attributed to the more eastern endemic E.
insigne), in E. luteolur (the var. saltuarium of granitic sands in the Sierra Nevada), and in E.
gracile (the glabrous var. incultum of Orange, Riverside and San Diego cos.). A new species of
the subg. Pterogonum from the Chihuahuan Desert of northern México, E. henricksonii, is also
proposed. The subg. Pferogonum sect. Pterogonum is formally established in accordance with
recent changes in the International Code of Botanical Nomenclature.

KEY WORDS: Polygonaceae, Eriogonum, taxonomy, California, Idaho, Montana, Oregon, Utah,
Colorado, México.

INTRODUCTION

Avsmall number of miscellaneous species and varieties of Eriogonum Michx. (Polygon-
aceae: Eriogonoideae) have gone unpublished for several years awaiting critical material, an
opportunity, or sufficient pressure to make them available. The rediscovery of E. chrysops
Rydb. in eastern Oregon now makes it possible to describe a series of species long felt to
be distinct from that species but included within it by various authors (Hitchcock 1964; Reveal
1973). A few new species and varieties in Eriogonum still remain to be described awaiting
adequate type material, but the opportunity to present a review of the eriogonoid genera of
California and Arizona for new state floras as well as forthcoming regional treatments for
the Chihuahuan Desert, the Intermountain West and for North America north of México
makes it mandatory that several names now receive their formal baptism.

251

252 PHYTOLOGIA volume 66(3):251-265 May 1989

TAXONOMY

Eriogonum thompsonae S. Wats. var. matthewsae (Rev.) Rev., comb. nov. E. corymbosum
Benth. in A. DC. var. matthewsae Rev., Phytologia 35: 441. 1976.

Welsh (1984) properly pointed out that the var. matthewsae was misplaced and my asso-
ciation of it near Eriogonum corymbosum var. aureum (M.E. Jones) Rev. was an error. The
variety is now transferred to its proper species.

Eriogonum lonchophyllum Torr. & A. Gray var. fendlerianum (Benth. in A. DC.) Rev.,
comb. nov. E. microthecum Nutt. var. fendlerianum Benth. in A. DC., Prodr. 14: 18.
1856. E. fendlerianum (Benth. in A. DC.) Small, Bull. Torrey Bot. Club 33: 55. 1906. E.
effusum Nutt. subsp. fendlerianum (Benth. in A. DC.) S. Stokes, Gen. Eriogonum 79.
1936.

Eriogonum lonchophyllum Torr. & A. Gray var. intermontanum (Rev.) Rev., comb et stat.
nov. E. intermontanum Rev., Madrofio 19:293. 1969.

As now defined, Eriogonum lonchophyllum is an exceedingly variable species ranging from
northeastern Utah and adjacent western Colorado south to northern New Mexico eastward
to northern Texas. Welsh (1984) reduced E. saurinum Rev. to a variety of the species and
E. humivagans Rev. [E. corymbosum Benth. in A. DC. var. humivagans (Rev.) Welsh] is now
considered to be a synonym of var. lonchophyllum. As indicated previously (Reveal 1976),
E. fendlerianum might well be considered a variant of E. lonchophyllum and that view is now
proposed.

Eriogonum ochrocephalum S. Wats. var. sceptrum Rev., var. nov.-TYPE. UNITED
STATES. IDAHO: Owyhee Co.: 11 mi SW of Bruneau along Idaho Highway 51, in clay
hills associated with Atriplex at about 2800 ft elev, 7 Jul 1974, Reveal 3687 (holotype:
US; isotypes: BRY, NY, RSA, UC, UTC and elsewhere).

A var. ochrocephalum scapis 2-3.5 dm longis (nec 0.5-2 dm) et involucris 3.5-4.5 (5) mm
longis (nec 3-4 mm) differt.

Plants erect clumped herbaceous perennials 2.5-4 dm tall; eaves narrowly elliptic to oblong,
the leaf-blade 1.5-2.5 cm long; scapes 2-3.5 dm long, glabrous or thinly floccose when imma-
ture in some; involucres narrowly turbinate to turbinate, 3.5-4.5 (5) mm long, 1.5-2.5 mm
wide, tomentose or sparsely floccose initially but soon glabrous; flowers yellow, 1-1.5 (2) mm
long; achenes 1.5-2 mm long.

Representative Specimens: - UNITED STATES. IDAHO: Elmore Co.: near Glenn Ferry, 5 mi W of King Hill, 13
Jul 1975, Reveal 3898 (BRY, CAS, DUKE, F, MICH, MO, NY, OKL, RSA, TEX, UTC); above Rosevear Gulch,
S of Glenn Ferry, 13 May 1980, Grimes et al. 1544 (CAS). Lemhi Co.: 3 mi S of Baker, 23 Jun 1947, Ripley &
Barneby 8852 (CAS). Owyhee Co.: 10 mi S of Bruneau, 29 May 1946, Maguire & Holmgren 26236 (BRY, CAS, DS,
UC, UTC); above Pickett Creek, 4 mi W of Oreana, 5 Jul 1980, Grimes et al. 1759 (CAS). Payette Co.: SE of
Payette, 3 Jun 1945, Ripley & Barneby 6546 (CAS). Twin Falls Co.: 10 mi N of Twin Falls, 30 May 1946, Maguire
& Holmgren 26239 (BRY, CAS, DS, MO, TEX, UC, UTC).

The var. sceptrum (Latin sceptrum, wand, as to the long naked scape) is the common form
of the species in southwestern Idaho and is most closely related to the var. calcareum (S.
Stokes) M.E. Peck of Malheur and Baker cos., Oregon. The leaves and the scapes of var.

Reveal: Combinations and novelties in Eriogonum (Polygonaceac) 253

calcareum are shorter while the flowers and achenes are decidedly longer (2-3 mm long)
than those of the var. sceptrim. The plants doubtfully placed in the new variety from eastern
Idaho (Ripley & Bameby 8852) differ in having sparsely floccose rather than glabrous scapes
and tomentose rather than thinly floccose to glabrous involucres. Efforts to relocate plants
in Lemhi Co. were unsuccessful.

Eriogonum verrucosum Rev., spec. nov.- TYPE. UNITED STATES. IDAHO: Custer Co.: On
a low ridge and slopes west of the road up the East Fork of the Salmon River, 1.3 mi
S of U.S. Highway 93, 6 mi E of Clayton and 20 mi SW of Challis, on steep clay slopes
associated with Artemisia, Atriplex and Phlox at about 5700 ft elev, 6 Jul 1974, Reveal
3678 (holotype: US!; isotypes: BRY, NY, RSA, UC, UTC and elsewhere).

A Eriogonum chrysops floribus verrucosis differt.

Plants low herbaceous perennials forming mats 1-3 dm across; /eaves basal, the leaf-blades
oblanceolate to spatulate, 10-15 mm long, 3-6 mm wide, densely white-tomentose below, less
so and greenish-white above, the tomentose petiole 8-15 mm long; flowering stems scapose,
+ erect, 3-7 cm long, glabrous or thinly pubescent with scattered hairs; inflorescences capitate,
0.7-1.2 cm across; bracts scalelike, ternate, triangular, 1-1.5 mm long, 0.8-1 mm wide, tomen-
tose; involucres congested, (4) 6-8 per head, turbinate, rigid, (2) 2.5-3 mm long, 1.7-2 mm
wide, floccose without especially on the upper half of the tube, the 5 triangular teeth 0.5 mm
long, not membranous-margined, the bractlets linear, 1-1.5 mm long, minutely fringed with
gland-tipped cells, the pedicels 3-4 mm long, glabrous; flowers yellow with greenish-yellow
to reddish-brown bases and reddish midribs, 1.5-2 mm long in anthesis and up to 2.5 mm
long in fruit, glabrous except for minute glands along the midribs within, the outer surface
distinctly pustulose especially along the midribs without forming a pronounced keel, the tepals
monomorphic, oblong to oblanceolate, those of the outer whorl 0.8-1.1 mm wide, those of
the inner whorl 0.5-0.7 mm wide, united about a third of their length; stamens slightly ex-
serted, the filaments sparsely pilose basally, the anthers yellow, 0.5-0.6 mm long, narrowly

oblong; achenes light brown, 1.5-2 mm long, glabrous, the subglobose base tapering to a long,
3-angled beak.

Representative Specimens.- UNITED STATES. IDAHO: Custer Co.: 5 mi above Challis along Garden Creek, 17
Jun 1969, Barneby 15079 (US); Lost River Range, Pahsimeroi Mountains, Lime Creek Canyon, 10 mi SE of Challis,
21 Jun 1982, Cholewa 1016 (ID); near top of divide on Spar Canyon Road, 4 Jul 1948, Christ & Christ 17707 (ID);
Morgan Creek, 9 mi N of Challis, 2 Jun 1951, Christ 51-239 and 51-241 (ID); Spar Canyon, Bear Wallow Road, 7
mi from the East Fork of the Salmon River Road, 2 Jul 1981, Grimes & Yakana 2195 (CAS, MARY); near mouth
of Pats Creek along Challis Creek Road, 13 Jun 1978, Henderson 4416 (MARY); E of summit of Spar Canyon, 14
Jun 1978, Henderson 4459 (MARY); Lake Creek, 15 mi W of Herd Lake, 21 Jun 1980, Henderson 5629 (ID); 0.2
mi from U.S. Highway 93 along W side of Morgan Creek, 13 Jun 1978, Henderson & Brunsfeld 4421 (ID); E side
of Salmon River, 0.2 mi below Deadman Hole and adjacent to Tunnel Rock at milepost 234 along U.S. Highway
93 between Challis and Clayton, 17 Jun 1980, Henderson & Henderson 5604 (ID); 20 mi S of Challis, 15 Jun 1944,
Hitchcock & Muhlick 8988 (CAS, DS, RM, UC, UTC, WTU); 8 mi S of Challis, 28 Jun 1946, Hitchcock & Muhlick
14118 (WTU); Challis Creek, 19 Jul 1916, Macbride & Payson 3328 (CAS, DS, MO, POM, RM, UC, US); 10 mi
SE of Challis, 22 Jun 1947, Ripley & Barneby 8822 (CAS). Lemhi Co.: 4.5 mi SW of Salmon, 7 Jun 1941, Chrise
12226 (ID); Salmon, 3 Jul 1920, Payson & Payson 1879 (CAS, RM).

Eriogonum verrucosum (Latin vernucosus, warty, alluding to the pustulose tepals) is restricted
to clay shale banks in the Challis area of central Idaho. The species is readily distinguished
by its densely pustulose tepals on flowers arranged in a rigid involucre. The narrow leaves
and glabrous to (and especially when young) sparsely pubescent stems are similar to those
seen in E. capistratum var. welshii which has membranaceous involucres.

254 PHYTOLO GIA volume 66(3):251-265 May 1989

The inclusion of low elevation plants from near Salmon within Eniogonum verrucosum must
be considered tentative. The specimens cited above from Lehmi Co. have been rather poorly
preserved and the features need to be confirmed. Attempts to recollect similar plants in the
area have been unsuccessful to date.

The new species has long been included within the broad definition of Enogonum chrysops.

Eriogonum meledonum Rev., spec. nov.-TYPE. UNITED STATES. IDAHO: Custer Co.:
Along U.S. Highway 93, 9 mi SSE of Stanley and 2 mi N of Obsidian, on steep sandy
clay slopes associated with Artemisia at about 6950 ft elev, 5 Jul 1974, Reveal 3674
(holotype: US!; isotypes: BRY, NY, RSA, UC, UTC and elsewhere).

A Eriogonum chrysops scapis 4-10 cm longis et rare tomentosis cum foliis (0.8) 1-1.5 cm
longis, floribus glabris differt.

Plants low herbaceous perennials forming mats (0.5) 1-2 dm across; leaves basal, the leaf-
blades spatulate to narrowly elliptic, (8) 10-15 mm long, 4-8 mm wide, densely grayish-
tomentose on both surfaces, sometimes less so above, the tomentose petiole (5) 10-15 (18)
mm long; flowering stems scapose, erect, 4-10 cm long, thinly tomentose; inflorescences
capitate, 1-1.5 cm across; bracts scalelike, ternate, linear-lanceolate, 2-3 mm long, 1-1.3 mm
wide, tomentose; involucres congested, (4) 6-8 per head, campanulate, membranaceous, 2.5-3
mm long, 3-3.5 mm wide, tomentose, the 5-6 lanceolate teeth 1-1.5 mm long, the pedicels
3-4 mm long, glabrous; flowers yellow, becoming rosy-yellow in fruit, 2.5-3 mm long, glabrous,
the tepals monomorphic, oblanceolate, united about a third of their length; stamens long
exserted, the filaments glabrous, the anthers yellow, 0.3-0.4 mm long, oval; achenes light
brown, 2.5 mm long, glabrous, the subglobose base tapering to a long, 3-angled beak.

Representative Specimens.- UNITED STATES. IDAHO: Custer Co.: 9 mi S of Stanley, 25 Jun 1941, Cronquist

2694 (DS, MO, UTC); Sawtooth Valley, bluffs along Valley Creek, 2 mi NW of Stanley, 3 Jul 1988, Moseley 1353
(MARY).

Eriogonum meledonum (Greek meledonos, guardian or caretaker, a fanciful name to honor
Mabel Allread Cronquist, 1916- , who collected with a famous botanist the summer of 1941
and has watched over him ever since) is a narrowly endemic species known only from a few
scattered locations in the Stanley area. It differs from E. capistratum in having thinly tomen-
tose rather than glabrous, glandular or densely tomentose scapes and glabrous rather than
generally glandular flowers. The leaves of the new species are broader than those of E.
chrysops and the scape is not densely tomentose as that seen in the Oregon endemic.

Eriogonum capistratum Rev., spec. nov.-TYPE: UNITED STATES. IDAHO: Custer Co.:
Antelope Pass, upper end of Copper Basin on a ridge SW of the pass, on rocky volcanic
slopes associated with scattered Artemisia and grasses at about 9000 ft elev, 11 Jul 1975,
Reveal & Ertter 3876 (holotype, US!; isotypes: BRY, CAS, DUKE, F, MARY, MEXU,
MICH, MO, NY, OKL, RSA, TEX, UTC and elsewhere).

A Eriogonum chrysops scapis 1-10 cm longis et glabris vel glandulosis, raro rare floccosis
cum involucris tomentosis vel glandulosis, floribus glabris vel glandulosis differt.

Plants low often compact herbaceous perennials forming mats (0.5) 1-2.5 (3) dm across;
leaves basal, the leaf-blades spatulate to narrowly elliptic or elliptic, 5-10 (15) mm long, 2-7
(11) mm wide, densely white-tomentose on both surfaces, sometimes slightly less so and
greenish above, with some glandular hairs in var. muhlickii, the tomentose petiole 2-6 (10)

Reveal: Combinations and novelties in Enogonum (Polygonaceae) 255

mm long; flowering stems scapose, + erect to erect, 1-10 cm long, glabrous to glandular or
sparsely tomentose to floccose; inflorescences capitate, 0.7-1.2 cm across; bracts scalelike,
ternate, triangular to oblanceolate, 1.5-3 mm long, 0.6-2 mm wide, sparsely to densely tomen-
tose or glandular without; involucres congested, 3-6 per head, turbinate-campanulate to cam-
panulate, membranaceous, 2-4 mm long, 2-3.5 mm wide, sparsely to densely tomentose or
glandular and sparsely floccose, the 5-7 triangular teeth 0.3-1.2 mm long, the bractlets linear-
oblanceolate, 1-2.5 mm long, minutely fringed with gland-tipped cells, the pedicels 2.5-4.5
mm long, glabrous except for a few glands near the tip in some; flowers yellow with greenish-
brown to reddish-brown midribs and bases, becoming rosy-yellow in fruit, 2-3.5 (4) mm long,
glabrous or sparsely glandular, the tepals monomorphic, oblanceolate to oblong, those of the
outer whorl slightly wider than those of the inner whorl, united about a quarter of their
length; stamens mostly exserted, the filaments sparsely pilose basally, the anthers yellow,
0.3-0.4 mm long, oblong; achenes brown, 2-3 mm long, glabrous, the subglobose base tapering
to a long, 3-angled beak.

Widespread in the valley and adjacent mountain ranges of central Idaho eastward to extreme western Montana
mostly above 6000 ft elev.

This species has long been known in the botanical literature as Eriogonum chrysops Rydb. In
its typical expression, the var. capistratum, the stems are glabrous or sometime glandular
with long, narrow leaves. This variety is found in the higher elevations of central Idaho usually
above 8000 feet elevation. On the eastern edge of its range this taxon gives way to the more
densely compact and decidedly glandular var. muhlickii in Montana while to the south it is
replaced to some degree by the generally lower elevation var. welshii which has a tomentose
to floccose scape. An unusual and probably undescribed relative to these taxa is represented
by an expression with cream colored glandular flowers on erect glabrous stems with long
narrowly elliptic leaves having blades 1-1.5 cm long and 1.5-5 mm wide on petioles 0.5-2 cm
long. At present the only specimen seen is a collection from the White Knob Mountains
along Alder Creek Road (Caicco & Civille 107, ID).

All of the expressions in the Eriogonum chrysops complex are morphologically similar dif-
fering mainly in having a distinctly rigid, firm involucral tube or a membranaceous, lax tube,
with or without glandular hairs, leaves, scapes and floral parts of various sizes, and slight
modifications on the tepals in terms of length, width, surface structures and pubescence.
Each of these entities is geographically isolated and locally consistent.

Eriogonum capistratum Rev. var. capistratum.

Low cespitose mats up to 3 dm across; /eaves tomentose, the leaf-blades (5) 10-15 mm
long on a petiole 4-10 mm long, the tomentum not glandular; scapes 3-10 cm long, glabrous
or glandular; involucres tomentose or glandular; flowers (2.5) 3-3.5 (4) mm long.

Representative Specimens. - UNITED STATES. IDAHo: Blaine Co.: above North Fork, Little Cottonwood Creek,
Craters of the Moon National Monument, 29 Jun 1956, W.H. Baker 14224 (ID); W of Cobb Peak and SW of
Hyndman Peak, Pioneer Mountains, 6 Aug 1977, Ertter & Lockwood 2166 (MARY, NY); S end of Soldier Mountain,
26 Jun 1916, Macbride & Payson 2896 (CAS, DS, MO, POM, RM, UC, US); base of Devils Bedstead, 28 Jul 1936,
Thompson 133546 (CAS, MO, US, WTU); Iron Bog, Pioneer Mountains, 24 Jun 1978, Wellner 1339 (ID). Custer
Co.: Cabin Creek Peak, 15 Aug 1980, Brunsfeld & Brunsfeld 1696 (ID); Antelope Pass, 29 Jul 1946, Christ 15986
(UC); head of Copper Basin, 7 Jul 1939, Davis 1188 (IDS); Mt. Hyndman, 11 Aug 1939, Davis 1712 (IDS, UC);
head of Slate Creek near Ocalkens Lake in the White Clouds, 24 Aug 1980, Ertter & Ertter 4023 and 4036 (MARY,
NY); Mount Parks, 15 Aug 1895, Evermann 375 (US); head of Pettit Lake, 29 Jul 1895, Henderson 3591 (US); 0.2
mi NW of Snowyslide Peak, 10 mi SW of Obsidian, 11 Aug 1939, Hitchcock & Martin 5786 (DS, OKL, UC, UTC,

256 PHYTOL OGIA volume 66(3):251-265 May 1989

WTU); Stanley Lake, 16 Jul 1936, Mann & Mann 36-131 (KSC); 3 mi SE of Stanley Lake, 16 Jul 1936, Mann &
Worley 70 (ID, US); Railroad Ridge, 5 Aug 1971, Schlatterer 200 (OGDF); near Stanley Lake, 28 Jul 1937, Thompson
14010 (CAS, WTU). Valley Co.: Monument Peak, 12 Aug 1924, Helm 234 (OGDF).

Eriogonum capistratum (from the Latin capistratus, tie with a halter or hitch, here fancifully
applied to honor Dr. Charles Leo “Hitchy” Hitchcock, 1902-1986, monographer of numerous
genera, author with others of the Vascular plants of the Pacific Northwest, long-time professor
of botany at the University of Washington, and a student’s best friend; see Denton 1986)
was long confused with E. chrysops and was so treated by Hitchcock (1964) where the fol-
lowing variety was illustrated.

The var. capistratum is restricted to the higher elevations mainly in the Sawtooth Mountains
of central Idaho where it is commonly found from 8000 to 10000 feet elevation on granitic
slopes.

Eriogonum capistratum Rev. var. muhlickii Rev., var. nov. - TyPE: UNITED STATES.

MONANA: Ravalli Co.: summit of St. Mary’s Peak, 7 air mi W of Stevensville, Bitterroot

Range, associated with Phlox, Dryas and Astragalus at about 9300 ft elev, 22 Aug 1974,

Reveal 3814 (holotype, US!; isotypes: BRY, MARY, MO, NY, OKL, RSA, UC, UTC
and elsewhere).

A var. capistratum foliis brevioribus (5-8 mm longis) cum scapis et involucris glandularis
differt.

Low compact cespitose mats up to 1.5 dm across; leaves tomentose, the leaf-blades 5-8
mm long on a petiole 2-5 mm long, the tomentum interspersed with glandular hairs; scapes
1-3 cm long, glandular; involucres glandular; flowers 2-2.5 (3) mm long.

Representative Specimens.- UNITED STATES. Montana: Deer Lodge Co.: Mt. Tiny, Pintlar Range, 24 Aug
1972, Lackschewitz 3976 (MONTU); Green Mountain, Pintlar Range, 20 Jul 1974, Lackschewitz 5302 (MONTU);
summit of Pagoda Mountain, 0.5 mi E of Pintlar Pass, Pintlar Range, 28 Jul 11974, Lackschewitz 5141 (MONTU).
Granite Go.: above Warren Lake, 21 Aug 1974, Lackschewitz 5603 (MONTU). Ravalli Co.: top of St. Mary’s Peak,
10 Aug 1946, Hitchcock & Muhlick 15320 (MO, RSA, UC, WTU); top of St. Mary’s Peak, 5 Aug 1947, Hitchcock
& Muhlick 17100 (CAN, DS, RSA, UC, WTU); Ward Mountain, Bitterroot Range, 17 Jul 1966, Lackschewitz &
Fageraas 52 (MONTU); Castle Crag, Bitterroot Range, 18 Aug 1970, Lackschewitz & Stuart 2376 (MONTUV).

For nearly the whole of their professional life times the names of C. Leo Hitchcock and
Clarence Muhlick were linked on numerous herbarium labels and in an entertaining array
of adventures that were frequently told from two rather divergent points of view. To a young
graduate student the interplay between these two men was wonderous. Now that both are
gone it is only appropriate that their association continue in at least one of their interests.

The var. muhlickii is the eastern component of the species being isolated on the Bitterroot
Range of Montana and on other high mountain ranges just to the east. The plant is decidedly
glandular, as compared to var. capistratum, as well as much more compact. So far as known,
the two do not share overlapping geographic ranges.

Eriogonum capistratum Rev. var. welshii Rev., var. nov.- TYPE: UNITED STATES. IDAHO:
Custer Co.: along the road from Ellis to Howe, 38 mi S of May and 46 mi N of Howe,
6 mi S of Summit Reservoir and 8.9 mi N of the Butte Co. line, on low windswept
barren gravelly clay upper slopes and ridgetops associated with Artemisia and Atriplex
at about 6400 ft elev, 16 Jul 1976, Reveal & Welsh 4501 (holotype, US!; isotypes: BRY,
CAS, DUKE, MEXU, MICH, MO, NY, OKL, RSA, TEX, UTC and elsewhere).

Reveal: Combinations and novelties in Eriogonum (Polygonaceae) 257

A var. capistratum scapis tomentosis vel floccosis differt.

Low compact cespitose mats up to 1.5 dm across; /eaves tomentose, the leaf-blades 5-12
mm long on a petiole 4-9 mm long, the tomentum not interspersed with glandular hairs;
scapes 2-8 (10) cm long, tomentose to densely floccose; involucres sparsely to densely tomen-
tose; flowers 2-3 mm long.

Representative Specimens. - UNITED STATES. IDAHO: Custer Co.: Lost River Range, Lime Creek, 10 mi SE of
Challis, 19 Jun 1979, Brunsfeld & Brunsfeld 1017 (ID); White Knob Mountains, W side of Burma Road N of Corral
Creek Pass, 8 Jul 1981, Caicco & Civille 165 (ID); White Knob Mountains, 0.5 mi E of Corral Creek Summit on
the Burma Road, 24 Jun 1979, Henderson 5161 (ID, MARY); 42 mi SE of May, 4 Jun 1964, Holmgren & Reveal
863 (CAS, NY, RSA, UC, UTC, US); 38 mi S of May, 31 Jul 1976, Reveal & Reveal 4606 (BRY, MO, US).

The var. welshii (for Dr. Stanley Larsen Welsh, 1929- , professor and curator of the her-
barium at Brigham Young University, and author of an Alaskan and a Utah flora) is a
narrowly restricted species known only from a few scattered sites in Custer Co., Idaho. This
expression is perhaps the most similar to the Oregon endemic, E. chrysops. The leaves of
the new variety are shorter and mostly broader than those of E. chrysops, the tomentose
scapes are shorter, and the flowers are glabrous. In E. chrysops, the flowers are weakly
glandular-puberulent with broader tepals. Another undescribed entity is known from the top
of War Eagle Peak in Owyhee Co. This plant has short narrow leaves (the blades 5-8 mm
long and 1-3 mm wide with a petiole 2-6 mm long), sparsely floccose scapes 4-6 cm long
and membranaceous 5-6-toothed campanulate involucres 3.5-4 mm long that are minutely
glandular without except for a few scattered hairs on the teeth. The bright yellow glandular
flowers are 2-3 mm long.

Eriogonum shockleyi S. Wats. var. packardae Rev., var. nov.- TYPE: UNITED STATES.
IDAHO: Owyhee Co.: above the upper sand dune of upper Halverson Lake, on “blow”
areas on the lava rim rock on the northern bank of the Snake River in the Bird of Prey
Santuary SE of Melba, associated with Artemisia and Atriplex at about 2500 ft elev, 7
Jul 1974, Reveal et al. 3686 (holotype, US!; isotypes: BRY, NY, RSA, UC, UTC and
elsewhere).

A var. shockleyi scapis + nullis differt.

Plants low cespitose herbaceous perennials forming mats 0.5-4 dm across; /eaves in dense
compact rosettes, the leaf-blades elliptic, 1-3 (3.5) mm long, 1-1.5 mm wide, densely tomen-
tose on both surfaces, the petioles 0.5-1 mm long; flowering stems lacking, or if present then
scapose and less than 5 mm long, tomentose; inflorescences capitate and terminal; bracts
scalelike, ternate, lanceolate, 0.5-1 mm long, tomentose; involucres congested, 2-4 per head,
broadly campanulate, + membranaceous, 2.5-3 mm long, tomentose, the 4-5 (6) lanceolate
teeth 0.8-1 mm long, the pedicels 0.3-0.5 mm long, glabrous; flowers 2-5 per involucre, pale
yellow, 2.5-3 mm long, densely tomentose, the tepals monomorphic, united about a third of
their length; stamens exserted, the filaments 3-4 mm long, glabrous, the anthers yellow, 0.3
mm long, oval; achenes light brown, 2.5-3 mm long, densely tomentose, the subglobose base
tapering to a short, 3-angled beak.

Representative Specimens. - UNITED STATES. IDAHO: Owyhee Co.: 10 mi S of Bruneau near Devil’s Bathtub,
9 Jun 1975, Bright s.n. (MIN); 15 mi ENE of Bruneau along Idaho Highway 51, 9 Jul 1975, Reveal 3852 (BRY,
MICH, NY, US).

258 PHYTOL O GIA volume 66(3):251-265 May 1989

The highly compact cespitose nature of the var. packardae and its essentially sessile cluster
of involucres covering the whole of the mat quickly differentiates this taxon from the more
open, less tightly matted, scapose var. shockleyi found to the south. The most densely cespitose
forms of var. packardae seen are those from the type area where the plant take on the
aspect — as the local common name implies — of dried “cow pies.” To the south near Bruneau,
the plants are less densely matted and here short scapes may be found.

The new variety is named for Dr. Patricia L. Packard, professor of biology and curator of
the Harold M. Tucker Herbarium at the College of Idaho in Caldwell, Idaho. Her enthusiasm
for systematic botany has been infectious to numerous undergraduate students, and many
such as James Henrickson, Barbara Ertter and James Grimes have gone on in the field.

Eriogonum nudum Douglas ex Benth. var. paralinum Rev., var. nov.-TYPE: UNITED
STATES. OREGON: Curry Co.: along U.S. Highway 101, 1.4 mi S of the Pistol
River —- Carpenterville turnoff, in sandy to gravelly soil on a low mesa above the Pacific
Ocean associated with scattered Pinus at about 150 ft elev, 16 Aug 1976, Reveal & Reveal
3961 (holotype, US!; isotypes: BRY, CAS, MEXU, MO, NY, OKL, TEX, UTC and
elsewhere).

A var. nudum involucris 5-10 in quoque fasciculo differt.

Plants spreading herbaceous perennials 1-5 dm high; /eaves essentially basal, the leaf-blades
1-2 cm long, tomentose below, glabrous and green above, the margins entire, on tomentose
petioles 4-10 cm long; flowering stems erect, slender, 0.8-3.5 dm long, glabrous, not fistulose;
inflorescences cymose, 1-2 dm long, branched 1-3 times, the branches glabrous; involucres
congested, 5-10 per head, 3-5 mm long, 2-2.5 (3) mm wide, glabrous, the 5-6 erect teeth
0.5-0.8 mm long; flowers white, 2-4 mm long, glabrous, the tepals + monomorphic, oblong,
those of the outer whorl slightly wider and shorter than those of the inner whorl; stamens
exserted, the filaments slightly pilose basally, the anthers dark red to maroon or purple,
0.3-0.4 mm long, oval; achenes light brown, 2.5-3 mm long, glabrous.

Representative Specimens.- UNITED STATES. Cauirornia: Del Norte Co.: 5 mi S of Crescent City, 17 Jun
1931, J.T. Howell 6729 (CAS); mouth of the Klamath River, 6 Aug 1947, Baker 11876 (CAS). OREGON: Curry Co.:
Ocean View Camp, Humbug State Park, 21 Jul 1940, Demaree 21445 (ISC, MIN, MO); 5 mi S of Port Orford, 19
Jul 1939, Maguire 17201 (RM, UTC); The Heads, Port Orford, 21 Jun 1919, M.E. Peck 8462 (MO, WILLU); mouth
of Wincheck River, 15 Jul 1919, M.E. Peck 8852 (MIN, WILLU); 0.8 mi S of Brush Creek, 2.7 mi S of Humbug
State Park, 16 Aug 1975, Reveal & Reveal 3963 (BRY, CAS, MO, NY, OKL, US); Harris Beach State Park, NW
of Brookings, 8 Jun 1955, Rossback 171 (DAO, UC); N of Brookings, 7 Aug 1935, Thompson 12544 (CAS, IND,
MIN, OKL, UC, WILLU, WTU); mouth of Pistol River, 21 Jun 1936, Thompson 12826 (CAS, MO, RSA, WILLU,
WTU).

The var. paralinum (Greek paralios, by or near the sea, alluding to its coastal habitat) is the
coastal expression of Eriogonum nudum, it being found on the mesas and bluffs above the
Pacific Ocean as well as on the immediate coast just above the high water mark. It differs
from the more inland var. nudum in its having more involucres in a cluster, shorter stems
and inflorescences, and in general its more crowded caudex. In many respects the var.
paralinum is similar to E. latifolium Smith in Rees, a densely tomentose coastal plant that
occurs from Del Norte Co., California, southward. There are glabrous forms of this species
mainly in the San Francisco area (these have been called E. oblongifolium Douglas ex Benth.),
but these plants still have the compact aspect of E. /atifolium rather than the more open and
erect aspect of E. nudum.

Reveal: Combinations and novelties in Eriogonum (Polygonaceae) 259

Eriogonum ovalifolium Nutt. var. pansum Rev., var. nov.- TYPE: UNITED STATES. IDAHO:
Boise Co.: along Idaho Highway 21, about 12.5 mi S of Lowman near the West Fork
Creek at milepost 59.5, on sandy-loam soil on slopes associated Pinus, Purshia, Artemisia,
Kelloggia and Lupinus at about 6000 ft elev, 12 Jul 1975, Reveal 3883 (holotype, US!;
isotypes: BRY, CAS, DUKE, F, MEXU, MICH, MO, NEB, NY, OKL, RSA, TEX,
UTC and elsewhere).

A var. ovalifolium inflorescentibus umbellatis differt.

Plants loosely matted herbaceous perennials 1-3 dm high and 2-5 dm across; /eaves essen-
tially basal, the leaf-blades elliptic or nearly so, mostly 1-2 cm long, densely tomentose on
both surfaces or with the upper surface slightly less so; flowering stems erect or nearly so,
0.7-2 dm long, thinly tomentose; inflorescences umbellate but usually not obviously so until
after fruit set, up to 7 cm long, the branches pedunculate and without bracts subtending the
involucres; flowers white.

Representative Specimens. - UNITED STATES. IDAHO: Boise Co.: 1 mi E of Lowman, 1 Jun 1944, Hitchcock &
Muhlick 8593 (OKL, RM, UC, WTU); E of Elk Lake above Sacajawea Hot Spring, 11-12 Jul 1944, Hitchcock &
Muhlick 9818 (RM, WTU). Custer Co.: Bonanza, 28 Jul 1916, Macbride & Payson 3506 (CAS, MIN, POM, RM,
UC); Cape Horn, 6 Aug 1916, Macbride & Payson 3628 (MIN, RM); 1 mi W of Sunbeam at Sunbeam Hot Springs,
6 Jul 1974, Reveal 3679 (BRY, MO, NY, OKL, RM, RSA, UC, US, UTC). Elmore Co.: 26 mi NE of Mountain
Home, 1 Jun 1966, Collotzi & Davidse 670 (UC, UTC); 3 mi NW of Pine, S of Dog Mountain, 5 Jun 1944, Hitchcock
& Muhlick 8728 (CAN, DS, IDS, OKL, RM, UC, WTU); Snowslide Creek, near Atlanta, 26 Jun 1942, MacFadden
25460 (CAS). Montana: Missoula Co.: above Upper Jocco Lake, E of Arlee, 10 Jul 1948, Hitchcock 18167 (RM,
RSA, UC, WTU). Powell Co.: 6 mi W of Ovando, Blackfoot Valley, 25 Jun 1945, Hitchcock & Muhlick 11524
(CAS, WTU). Silver Bow Co.: Durant, 6 Jun 1906, M.E. Jones s.n. (POM).

The var. pansum (from the Latin pansus, spreading out, extended, alluding to the umbellate
rather than capitate inflorescences) is not always readily distinguishable as the expanding
peduncles are often not obvious until fruit formation. Nonetheless, being aware of the variant,
one can quickly recognize by carefully examining the immature inflorescences.

Eriogonum umbellatum Torr. var. goodmanii Rev., var. nov.- TYPE: UNITED STATES.
OREGON: Josephine Co.: along the O’Brien-Happy Camp Road, 2.7 mi E of U.S. High-
way 199 near Waldo, on an open serpentine flat associated with scattered Pinus and
Arctostaphylos at about 1550 ft elev, 9 Aug 1978, Reveal & Reveal 4815 (holotype, US!;
isotypes: BRY, CAS, OKL, TEX!).

A var. polyanthum (Benth. in A. DC.) M.E. Jones plantis tegetes formantes 2-4 dm alatis
et 4-7 dm latis, foliis tomentosis, involucris brevioribus (2-3 mm longis nec (3) 4-6 mm longis)
cum floribus longioribus (6-8 (9) mm longis nec (4) 6-7 mm longis) differt.

Plants low spreading or prostrate matted herbaceous perennials 2-4 dm high and 4-7 dm
across; /eaves in loose rosettes, the leaf-blade elliptic to ovate, 0.5-2 (3.5) cm long, 0.5-1 (1.5)
cm wide, densely white lanate below, only slightly less densely tomentose above, rarely merely
densely floccose, the slender tomentose petiole 0.3-1.5 cm long; flowering stems stoutish, erect,
1-2.5 (3) dm long, floccose; inflorescences umbellate, 3-5 (8) cm long, the branches floccose;
involucres campanulate, the tube 2-3 mm long, the 6-8 (10) reflexed lobes 2.5-4 mm long,
floccose; flowers yellow, often becoming reddish with age, 6-8 (9) mm long including the stipe.

Representative Specimens.- UNITED STATES. CALIFORNIA: Del Norte Co.: Poker Flat-Twin Valley Trail, 31
Jul 1938, Van Deventer 204 (MSC, UTC). Humboldt Co.: Grouse Mountain, 25 Jul 1933, Tracy 12881 (UTC);

260 PHYTOL O GIA volume 66(3):251-265 May 1989

Horse Mountain, 6 Jul 1971, Anderson & Smith J-1281 (HSC). Siskiyou Co.: between Kings Castle and Black
Mountain, 9 Jul 1939, Hitchcock & Martin 5312 (OKL, WTU); 1.4 mi E of the summit of the O’Brien-Happy Camp
Road, 18 Aug 1974, Reveal & Reveal 3799 (BRY, MO, OKL, RM, UC, US, UTC); Marble Mountains, 27 Jun
1976, Stillman 147 (HSC). Trinity Co.: Senteney Rock, 28 Jul 1976, Nelson & Nelson 3051 (HSC). OREGON: Jose-
phine Co.: 5 mi N of the California line along the Redwood Highway, 28 Aug 1930, Goodman & Hitchcock 1820
(BKL, DS, MO, OKL, UC); 3 mi E of O’Brien, 8 Jun 1966, Holmgren & Reveal 2666 (BRY, CAS, DS, IDS, MSC,
NY, RSA, TEX, US, UTC); 10 mi W of U.S. Highway 99 along the Galice-Almeda Road, 22 Jun 1950, Kruckeberg
1900 (CAN, COLO, RM, RSA, UC); 7 mi E of O’Brien, 18 Aug 1974, Reveal & Reveal 3803 (BRY, MSC, UC,
UTC); 9 mi E of U.S. Highway 199 on the Happy Camp Road, 9 Aug 1978, Reveal & Reveal 4814 (BRY, CAS,
MO, RSA, US); Rough and Ready Creek, 7 Jun 1972, Rose 72022 (CAS); 1 mi E of O’Brien, 11 Jul 1965, White
7116559 (HSC).

The var. goodmanii is named for George Jones Goodman (1904— ), professor of botany at
the University of Oklahoma and long-time student of eriogonoid genera best known for his
work on Chorizanthe. He and C. Leo Hitchcock collected this plant while on their famed
1930 western botanical excursion. The new variety belongs to the var. polyanthum complex.
It occurs on serpentine soil where it forms large mats of densely tomentose leaves with long

erect flowering stems bearing rather short and compact umbellate inflorescences of bright
yellow flowers.

Eriogonum umbellatum Torr. var. humistratum Rev., var. nov.—TYPE: UNITED STATES.
CALIFORNIA: Trinity Co.: along the south and southwestern ridge of Scott Mountain,
associated with scattered Pinus and numerous shrubs at 6700-6800 ft elev, 17 Aug 1974,
Reveal & Reveal 3794 (holotype, US!; isotypes: BRY, MARY, NY, RSA, UC, UTC and
elsewhere).

A var. polyanthum (Benth. in A. DC.) M.E. Jones plantis tegetes formantes 1-1.5 dm alatis
et 1-3 dm latis, foliis tomentosis, involucris brevioribus (1.5-2 mm longis nec (3) 46 mm
longis) differt.

Plants low prostrate matted herbaceous perennials 1-1.5 dm high and 1-3 dm across; leaves
in + compact rosettes, the leaf-blade broadly elliptic, 0.5-1.5 cm long, 0.5-1 cm wide, densely
white tomentose on both surfaces or slightly less so and grayish-white tomentose above, the
tomentose petiole 0.3-1 cm long; flowering stems slender, erect, 0.5-1 dm long, floccose; in-
florescences umbellate, 2-5 cm long, the branches floccose; involucres campanulate, the tube
1.5-2 mm long, the 5-8 reflexed lobes 1.5-3 mm long, floccose; flowers bright yellow, 3-6 mm
long including the stipe.

Representative Specimens.- UNITED STATES. CaALiFornia: Siskiyou Co.: trail from Scott Mountain to Carmen
Lake, 8 Jul 1949, Balls 13914 (RM, RSA); N side of Mt. Shasta, 15-30 Jun 1897, Brown 433 (MIN, UC); Mt. Eddy,
16 Jul 1918, Heller 13027 (CAS, DS, F, MO, PENN, WIS, UC); summit of Salmon Mountain, 16 Aug 1948, Parker
182 (DS, RSA, UC); ridge SE of Mt. Eddy, 20 Aug 1976, Whipple 1713 (HSC). Trinity Co.: near Deadfall Lake,
Mt. Eddy, 4 Aug 1976, Whipple 1646 (HSC).

The name, var. humistratuyn (Latin humus, ground, and stratum, prostrate, as to the densely
matted habit), has appeared in various state and national listings of species, including some
rare species lists, even though the name had not been published. The plant is a low, densely
matted expression with tomentose leaves and a short stature. A member of the var. polyan-
thum complex, it occurs at the higher elevation mainly on serpentine outcrops in the northern
Coast Ranges. The Brown specimen from Mt. Shasta cited above should probably be assigned
here, but the Mt. Shasta material differs slightly in several features.

Reveal: Combinations and novelties in Eriogonum (Polygonaceae) 261

Eriogonum umbellatum Torr. var. argus Rev., var. nov.- TYPE: UNITED STATES. CALI-
FORNIA: Siskiyou Co.: along the Dry Lake Lookout Road, 0.5 mi W of Dry Lake Lookout
near the Buckthorn Trail, Klamath National Forest, on sandy-gravelly soil associated
with Abies and Lupinus at 6700 ft elev, 15 Aug 1975, Reveal & Reveal 3955 (holotype,
US!; isotypes: BRY, CAS, F, MARY, MEXU, MICH, MO, NY, OKL, RSA, TEX, UC,
UTC)).

A var. furcosum Rev. foliis subtiliter crenatis cum caulibus parce floccosis ad glabris differt.

Plants low spreading matted herbaceous perennials 2-4 dm high and 5-15 dm across; /eaves
in loose rosettes, the leaf-blade oblanceolate to elliptic, (0.7) 1-2 (2.5) cm long, 0.4-1 cm
wide, thinly to moderately tomentose below, sparsely floccose to glabrous and green above,
the thinly tomentose petiole 0.3-1.5 cm long; flowering stems slender, erect, (0.8) 1-2 dm long,
sparsely floccose to glabrous, often with a single foliaceous leaflike bract near the middle;
inflorescences compound umbellate, 2-5 (9) cm long, the branches sparsely floccose to
glabrous; involucres campanulate, the tube 2-3 mm long, the 5-8 reflexed lobes 2-3.5 mm
long, floccose; flowers bright yellow, 3-8 mm long including the stipe.

Representative Specimens. - UNITED STATES. CA.iFornia: Del Norte Co.: summit of the O’Brien-Happy Camp
Road, 19 Aug 1974, Reveal & Reveal 3798, (BRY, MO, MSC, NY, OKL, RM, RSA, UC, US, UTC); Little Greyback,
14 Aug 1934, Lee 1103 (UC). Glenn Co.: summit of Black Butte, 10 Jul 1982, Wheeler 3174 (CAS). Humboldt Co.:
Trinity Summit, 25 Jul 1902, Jepson 2119 (JEPS); North Trinity Mountain, 31 Jul 1973, Klipfel 224 (HSC); head
of Devil’s Hole, 28 Jul 1935, Tracy 14329 (UC, UTC). Siskiyou Co.: Log Lake, Shackelford Creek, 3 Aug 1908,
Butler 209 (MO, UC, UTC); E side of Mt. Eddy, 28 Aug 1914, Heller 11742 (CAS, DS, F, ILL, MIN, NEB, PAC,
PENN); Scott Mountain Campground, 8-9 Aug 1953, Kellogg 114 (DUKE, IDS, MSC, OKL, UC, UTC); ridge S
of English Peak, Marble Mountain, 13 Aug 1969, Oettinger 1353 (HSC, RSA, UC); 1.4 mi E of the summit of the
O'Brien-Happy Camp Road on the road to Bolan Mountain Lookout, 18 Aug 1974, Reveal & Reveal 3801 (BRY,
MO, US, UTC); Jaynes Canyon, 1 Sep 1934, Wheeler 3234 (CAS, DS, POM); 3.6 mi from Dry Lake Lookout on
road to Horse Creek, 5 Jul 1934, Wolf 5966 (CAS, RM, RSA, TEX, UC). Trinity Co.: Canyon Creek, Trinity
Mountains, 6 Aug 1948, Alexander & Kellogg 543] (UC); summit of Scott Mountain, 19 Aug 1948, Parker 223 (DS,
RM, RSA, UC); Devil’s Canyon Mountains, above White’s Creek Lake, 8 Aug 1935, Tracy 14711 (CAN, UC); 1
mi W of Mud Springs, 5 Jul 1947, Tracy 17781 (OKL, UC, UTC). OrEGon: Jackson Co.: near Pilot Rock, Siskiyou
Mountains, 18 Jun 1928, Applegate 5530 (DS); Mt. Ashland, 11 Jul 1935, Jackson s.n. (CAS, DS). Josephine Co.:
top of Lake Mountains, Oregon Cave National Monument, 6 Aug 1936. Applegate 10770 (DS); 15 mi SE of Cave
Junction on Happy Camp Road, 18 Aug 1965, White 8186597 (HSC).

The var. argus (from the Latin argus, eye, alluding to the small clusters of bright yellow flowers
against the bright green branches of the inflorescences) is the expression of the var. stellatum
(Benth. in A. DC.) M.E. Jones complex found mainly on serpentine soils in the northern
Coast Ranges of northwestern California and southwestern Oregon. The var. stellatum is a
more northern taxon mainly of the Cascade Ranges of Oregon and Washington eastward to
Idaho. The Sierra Nevada expression is the var. furcosum Rev. while the var. munzii Rev. is
found in the Transverse Ranges of southern California. All of these variants differ slightly
morphologically but are consistent within their geographic ranges.

Eriogonum henricksonii Rev., spec. nov.- TYPE: MEXICO. COAHUILA: 1.5 mi SW of Las
Delicias, ca 68 airmiles SW of Cuatro Cienegas, E side of the Sierra de las Delicias, in
a limestone arroyo above the main spring, 26°14°N, 102°49°W, 4000 ft elev, 15 Aug
1973, Henrickson 12464a (holotype, US!; isotype: TEX!).

A Enogonum fimbriatum Hess & Rev. floribus flavus differt.

262 PHYTOLOGIA volume 66(3):251-265 May 1989

Plants erect herbaceous perennials 4-5 dm high arising from a small compact slightly woody
caudex; leaves basal, the leaf-blades elliptic, 1.5-2.5 cm long, 0.8-1.5 cm wide, glabrous and
green on both surfaces except for the slender straight ciliate hairs 1-2 mm long along the
margin and midrib, the base tapering abruptly to a ciliate winged petiole 1-1.5 cm long;
flowering stems erect, slender, 2-3 dm high, glabrous throughout; inflorescences open, cymose,
divided 3-6 times, glabrous except for a few scattered small, nearly sessile, white, capitate
glands just above the node; bracts scalelike, ternate, 1-2 mm long, 0.5-1 mm wide, glabrous;
peduncles slender, erect, straight, the first 3-5.5 cm long, the succeeding ones (0.5) 1-2 cm
long, glabrous except for a few scattered glands at the very base; involucres broadly campan-
ulate, 2-2.5 mm long, 3-4 mm wide, glabrous within and without except for a few erect hairs
within at the top of the throat, the 5 rounded teeth 0.5-0.7 mm long, the bractlets linear,
1.5-2 mm long, densely villous with long hyaline non-glandular marginal cells up to 0.4 mm
long, the pedicels 1.5-2 mm long, glabrous; flowers yellow with reddish-brown midribs and
bases, 1.5-2.5 mm long in anthesis, 3-4 mm long in fruit, glabrous except for the minute
glands along the midrib and base, the tepals slightly dimorphic, those of the outer whorl
obovate and shorter than the longer and narrower oblanceolate inner ones, united about a
quarter of their length; stamens included, 1.5-2 mm long, the filaments glabrous, the anthers
yellowish, 1-1.2 mm long, oblong; achenes light brown, 2.5-3.5 (4) mm long, ovate, glabrous,
the broadly globose base tapering abruptly to a slightly 3-angled beak.

Specimens Examined. — MEXICO. CoaHulILa: 1.5 mi SW of Las Delicias, 12 Aug 1973, Henrickson & Wendt 12300
(US).

Eriogonum henricksonii (James Henrickson, 1940- , noted collector and authority on the
flora of the Chihuahuan Desert) has been known since just after a revision of the subg.
Pterogonum (H. Gross) Rev. was published by Hess and Reveal (1977). It was hoped that
additional material might be collected prior to the need to describe the species for the
Chihuahuan flora, but no one has been back to the area. The new species belongs to the
subsect. Adenogonum Hess & Rev. of the Eriogonum subg. Pterogonum sect. Pterogonum
(H. Gross) Rev., comb. nov., based on Pterogonum H. Gross, Bot. Jahrb. Syst. 49: 239. 1913;
this combination is now required by the ICBN. The species is most closely related to E.
fimbriatum of Nuevo Le6n but differs from the remainder of the species in the subsection
in having distinctly yellow flowers. The new species occurs on gypsum outcrops in association
with Agave, Mortonia, Acacia and Fouquieria shrevet.

Eriogonum trichopes Torr. in Emory var. hooveri Rev., var. nov.- TYPE: UNITED STATES.
CALIFORNIA: San Benito Co.: Griswold Hills, along the New Idria Road near Griswold
Creek, 6.4 mi SW of Panoche Road and 3.7 mi N of the summit of the canyon at the
Syncline Ranch, on clay hills and road cuts associated with grassland species at about
1550 ft elev, 23 Jun 1978, Reveal 4748 (holotype: US!; isotypes: BM, BRY, CAS, GH,
MARY, MO, NY, RSA, UTC and elsewhere).

A var. trichopes plantis altjoribus cum glandulosis basi, peduculatis et involucris longioribus
differt.

Plants erect herbaceous annuals 4-15 (18) dm high; /eaves basal, the leaf-blades reniform,
(0.7) 1-2 (3) cm long and wide, hirsute and green on both surfaces, the petiole 1-3 cm long;
flowering stems erect, 0.5-2 (5) dm long, glabrous except for the glandular base above the
leaves, slightly fistulose in most near the apex; inflorescences cymose, rather strict and upright,
3-10 (12) dm long, glabrous except for a few scattered glandular hairs above the first node,

Reveal: Combinations and novelties in Eriogonwn (Polygonaceae) 263

rarely so at the second or third, the branches of the first node often slightly fistulose, rarely
so above in larger plants; peduncles capillary, + erect, 1-4 cm long, glabrous; involucres tur-
binate, 1-1.5 (1.8) mm long, glabrous, the 4 (5) triangular teeth 0.5-0.7 mm long; flowers
yellow, 1.5-2 mm long at anthesis, 2-2.5 mm long and often reddish at maturity, densely and
conspicuously short-hirsute with coarse white curved hairs, the tepals monomorphic, narrowly
ovate; stamens included, 1-1.5 mm long, the filaments subglabrous basally, the anthers yellow,
0.3-0.5 mm long, oval; achenes brown, 2-2.5 mm long, glabrous, the subglobose base tapering
to a short, 3-angled beak; n= 16.

Representative Specimens. - UNITED STATES. CALIFORNIA: Fresno Co.: 9 mi N of Coalinga, 26 Apr 1934, Keck
2790 (CI, DS). Kern Co.: Cuyama Valley, 28 Apr 1937, Eastwood & Howell 4065 (CAS, PENN, UC); Microwave
Fork, Cedar Canyon, Temblor Range, 8 Jul 1969, Twisselmann 15632 (CAS). Monterey Co.: N of Parkfield, 13
Jun 1938, Eastwood & Howell 5887 (BKL, CAS, RSA); 11 mi S of Parkfield Junction, 28 May 1941, Ferris &
Bacigalupi 10371 (CAS, DS, LA, MICH, RM, RSA, UC, UTC); Parkfield Grade, N of Parkfield at milepost 7.4,
19 Jun 1987, Reveal & Broome 6489 (BRY, CAS, MARY, MO, NY, RM, RSA, US); Parkfield Grade, 8.2 mi above
Parkfield, 16 Jun 1967, Twisselmann 1337] (CAS, MARY, MSU, NY, RSA, UC, US, UTC). San Benito Co.: New
Idria, 10 Jul 1861, Brewer 763 (CAS, DS, MO, POM, UC); 5.6 mi S of Panoche Pass above Griswold Creek, 14
Jul 1956, Ferris et al. 13033 (CAS, DS, JEPS); 18 mi from New Idna on road to Panoche, 11 May 1957, Raven
10850 (BRY, ILL, KANS, LL, OKL, OKLA, RM, UC, UTC); 3.1 mi below New Idria along the Panoche Road,
4 Oct 1966, Twisselmann 12881 (CAS, MARY, RSA, UTC). San Luis Obispo Co.: Palo Prieta Canyon, 14 Sep
1946, Hoover 6389 (CAS, OBI, UC); Chalk Mountain, Cuyama Valley, 27 Mar 1967, Hoover 10281 (CAS, OBI);
Pinole Canyon near Shandon, 20 May 1952, McMillan 127 (CAS, OBI); Palo Prieta Canyon summit, 25 Apr 1965,
Twisselmann 10549 (CAS, RSA). Santa Barbara Co.: Cuyama Valley, 5S mi E of Santa Maria, 17 May 1929, Munz
11422 (POM). Ventura Co.: Lockwood Valley, 19 Jun 1896, Dudley & Lamb 4614 (DS, POM, US); 22 Jun 1949,
Pollard s.n. (CAS); Ballinger Canyon, San Emigdio Range, 25 May 1962, Twisselmann 7183 (CAS).

The var. hooveri has long been included within the concept of Eriogonum inflatum Torr. &
Frém. which, in California at least, can flower its first year on the Mojave and Sonora deserts.
The new variety is strictly an annual and while some individuals will have 5-toothed involucres
similar to E. inflatum, the majority are 4-toothed and thus more like the condition in E.
trichopes. The new variety differs from var. trichopes in being a taller, rather strict plant with
longer peduncles and involucres. The two variants overlap only slightly in portions of Kern
and Ventura cos., California. The new variety is named for Dr. Robert Francis Hoover (1913-
1970), one of Willis Jepson’s last students and long-time professor of botany at California
State Polytechnic University in San Luis Obispo.

Eriogonum deflexum Torr. in Ives var. rectum Rev., var. nov.- TYPE: UNITED STATES.
CALIFORNIA: San Bernardino Co.: Bristol Mountains, along the west bound lane of In-
terstate 40, 16.8 mi E of Ludlow and about 66.8 mi E of Interstate 15 at Barstow, on
gravelly slopes and road shoulder associated with Larrea, 5 Jun 1988, Reveal & Broome
6385 (holotype: US!; isotypes: BM, BRY, CAS, GH, MARY, MO, NY, RSA, UTC and
elsewhere).

A var. baratum (Elmer) Rev. involucris erectis et turbinatis ad pedunculatis nullis differt.

Plants erect herbaceous annuals (3) 5-10 (20) dm high; /eaves basal, the leaf-blades cordate,
1-3 cm long and wide, densely white-tomentose below, thinly floccose to subglabrate and
green above, the tomentose petiole 2-5 cm long; flowering stems erect, mostly solitary, 1-2
(4) dm long, slender, glabrous and glaucous; inflorescences strict and narrow, (2) 3-9 (18)
dm long, slender, glabrous; bracts scalelike, ternate, triangular, 1-2 mm long, glabrous;
peduncles lacking; involucres erect, turbinate, 1.5-2 mm long, 1-2 mm wide, glabrous, the 5
rounded teeth 0.2-0.4 mm long; flowers white to pinkish, 1-2 mm long, the tepals dimorphic,

264 PHYTOL OG1A volume 66(3):251-265 May 1989

those of the outer whorl oblong with with truncate bases, those of the inner whorl lanceolate;
stamens included, 1-1.5 mm long, the filaments subglabrous basally, the anthers red to
maroon, 0.3-0.5 mm long, oblong; achenes brown, 1.5-2 mm long, glabrous, the subglobose
base tapering a short, 3-angled beak; n= 20.

Representative Specimens.- UNITED STATES. Ca.irornia: Imperial Co.: Fish Creek, Split Mountain Canyon,
31 Mar 1939, Jaeger s.n. (DS); Split Mountain Canyon, 1 Apr 1939, Jaeger s.n. (DS). San Bernardino Co.: Rodman
Mountains and Lava Bed Mountains, 0.3 mi S of powerline road to Ford’s Knob microwave relay station, 25 Mar
1980, Emmel 697 (MARY); 10 mi E of Baker, 1 Aug 1939, Jaeger s.n. (POM); 18 mi E of Ludlow, Old Dad
Mountains, 15 Jul 1975, Norris 2703 (RSA). San Diego Co.: Fish Creek, Nov 1936, Gentry 2962 (AHFH); 5 mi NW
of Carrizo Station, 27 Jan 1940, Munz 16835 (POM); Colorado Desert, Nov 1890, Orcutt 1466 (B, GH, US).

In California this taxon has, in the past (Reveal 1968), been known as Enogonum insigne and
is reported as such in Munz (1968, 1974). That error is mine and due largely to my having
failed to fully appreciate the differences I was observing and trying to take into account. The
var. rectum (Latin rectus, erect, alluding to the erect sessile involucres) differs from E. insigne
in having truncate rather than deeply cordate outer tepals and sessile rather than shortly
peduncled involucres. Plants from Inyo Co. previously placed in E. insigne seem better in E.
deflexum var. deflexum. The new variety is most closely related to the var. baratum.

Eriogonum luteolum E. Greene var. saltuarium Rev., var. nov.—TYPE: UNITED STATES.
CALIFORNIA: Tuolumne Co.: along California Highway 108, 0.8 mi E of Dardanelle, in
deep granitic sandy soil near the highway above the Middle Fork of the Stanislaus River,
associated with Artemisia and various grasses under Pinus at about 6200 ft elev, 22 Jul
1975, Reveal 3967 (holotype: US; isotypes: BRY, CAS, MARY, MO, NY, OKL, UTC!).

A var. pedunculatum (S. Stokes) Rev. involucris brevioribus et floribus et fructibus longi-
oribus differt.

Plants erect herbaceous annuals 1-4.5 dm high; /eaves strictly basal, the leaf-blades ovate
to reniform, 0.5-1.5 cm long, (0.7) 1-1.5 (1.8) cm wide, densely white-tomentose below, floc-
cose and green above, the tomentose petiole 0.5-1.5 cm long; flowering stems erect, 0.2-1.5
dm long, slender, glabrous; inflorescences strict and narrow, 1.5-4 dm long, the branches
slender, glabrous; bracts scalelike, ternate, triangular, 1-2 mm long, glabrous; peduncles lack-
ing; involucres sessile along the branches and closely appressed, cylindric, 2-3 mm long, 0.8-1
(1.2) mm wide, glabrous, the 5 acute teeth 0.2-0.5 mm long; flowers white to red, often with
a yellowish hue, 2-3 mm long, the tepals monomorphic, obovate; stamens included, 1.5-1.8
mm long, the filaments glabrous, the anthers red, 0.2-0.3 mm long, oval; achenes brown, 2-2.5
mm long, glabrous, the subglobose base tapering a short, 3-angled beak.

Representative Specimens. - UNITED STATES. CauiFornia: Alpine Co.: along California Highway 89, 1.5 mi SE
of Luther Pass, 23 Aug 1975, Reveal 3968 (BRY, CAS, MARY, MO, MICH, OKL, RSA, TEX, US, UTC).
Tuolumne Co.: Eureka Valley, 22 Jul 1957, Hesse 2369 (CAS); Brightman Flat, 15 Jul 1938, Hoover 3652 (DS,
MICH, UC, UTC); near Dardanelle, 21 Jul 1939, Eastwood & Howell 7611 (CAS, ISC); 0.5 mi W of Dardanelle,
28 Jul 1972, Reveal & Reveal 2814 (BM, BRY, CAS, GH, MARY, MO, NY, RSA, US, UTC); 0.8 mi east of
Dardanelle, 22 Sep 1933, Wolf & Stark 5473 (BKL, CAS, DS, F, IDS, ISC, MARY, MICH, MIN, MO, POM, RM,
UC, UTC).

The var. saltuarium (Latin saltuarius, ranger or forester, in honor of my father, Jack L. Reveal
(1912-1988), district ranger for the United States Forest Service on the Summit District of
the Stanislaus National from 1948 until 1960 where the type was collected) is unique within
Eriogonum luteolum as it is not found on serpentine soils. It is most closely related to the

Reveal: Combinations and novelties in Eriogonum (Polygonaceae) 265

lower elevation var. pedunculatum found along the foothills of the Sierra Nevada from Butte
Co. to Tulare Co. The involucres of var. saltuarium are 2-3 mm long while those of var.
pedunculatum are 3-3.5 mm long. The flowers of the new variety are longer (2-3 mm long
compared to 1-1.8 mm long) as are the achenes (2-3 mm versus 1-1.4 mm). In general the
leaves of var. saltuartum are smaller and broader than var. pedunculatum.

Eriogonum gracile Benth. var. incultum Rev., var. nov.-TYyPE: UNITED STATES. CALI-
FORNIA: San Diego Co.: Palomar Mountains, along East Grade Road (County Road S-7),

1.6 mi SE of Palomar Mountain (townsite) near the junction of County Road S-6, in
sandy soil associated with Quercus and chaparral at about 5100 ft elev, 28 Jun 1987,
Reveal & Broome 6620 (holotype: US!; isotypes: BM, BRY, CAS, GH, MARY, MO,
NY, RSA, UTC and elsewhere).

A var. gracile plantis glabris differt.

Plants erect herbaceous annuals 2-5 dm high; /eaves basal and cauline, the leaf-blades
oblanceolate to oblong, 1-3 (4) cm long, 0.5-1 (1.5) cm wide, densely white-tomentose below,
less so to floccose and green above, often with wavy or crisped margins, the tomentose petiole
1-2 cm long; flowering stems erect, 0.5-2 dm long, mostly slender, glabrous; inflorescences
open but strict and generally narrow, 1.5-4.5 dm long, the branches slender, smooth glabrous;
bracts scalelike, ternate, triangular, 1-2 mm long, glabrous; peduncles lacking; involucres ses-
sile along the branches and closely appressed, cylindric, 1.8-2 mm long, 0.6-1 mm wide,
glabrous, the 5 acute teeth 0.1-0.2 mm long; flowers white to pale yellow and often reddish,
1-1.5 mm long, the tepals monomorphic, oblong; stamens included, 0.8-1.2 mm long, the
filaments pilose basally, the anthers yellowish, 0.2-0.3 mm long, oval; achenes light brown,
1-1.5 mm long, glabrous, the subglobose base tapering a narrow, 3-angled beak.

The var. incultum (Latin incultus, untilled, here fancifully referring to the lack of hairs) is
found in Orange, Riverside and San Diego cos., California, where it can occur with the var.
gracile but typically not in mixed populations. At present, my herbarium records do not fully
differentiate between the two varieties and thus no representative specimens are cited.

ACKNOWLEDGMENTS

I am grateful for the comments by Dr. Edward E. Terrell and Clare B. Hardham who
reviewed the manuscript. This is Scientific Article A-4942, Contribution No. 7985, Maryland
Agricultural Experiment Station.

LITERATURE CITED

Denton, M. L. 1986. C. Leo Hitchcock. Taxon 35: 642-643.
Hess, W. J. & J. L. Reveal. 1977. A revision of Eriogonum (Polygonaceae) subgenus Pterogonum. Great Basin
Naturalist 36: 281-333.
Hitchcock, C. L. 1964. Eriogonum. Univ. Wash. Publ. Biol. 17(2): 104-138.
Munz, P. A. 1968. Supplement to A California Flora. Univ. California Press, Berkeley.
. 1974. A flora of southern California. Univ. California Press, Berkeley.
Reveal, J. L. 1968. Notes on Eriogonum -IV. A revision of the Eriogonum deflexum complex. Brittonia 20: 13-33.
. 1973. “Eriogonum,” pp. 79-84. In: C. L. Hitchcock & A. Cronquist, Flora of the Pacific Northwest.
Univ. Washington Press, Seattle.
. 1976. Eriogonum (Polygonaceae) of Arizona and New Mexico. Phytologia 34: 409-484.
Welsh, S.L. 1984. Utah flora: Polygonaceae. Great Basin Naturalist 44: 519-557.

Phytologia (May 1989) 66(3):266-294

A CHECKLIST OF THE ERIOGONOIDEAE (POLYGONACEAE)
JAMES L. REVEAL

Department of Botany, University of Maryland,
College Park, Maryland 20742-5815

ABSTRACT

A putative phylogenetic scheme of the generic and major infrageneric entities of the subf.
Eriogonoideae (Polygonaceae), along with an index to the names, is given. More than 1500
names are accounted for. An index of important systematic literature is appended.

KEY WORDS: Polygonaceae, Acanthogonum, Acanthoscyphus, Aristocapsa, Brisegnoa, Centro-
stegia, Chorizanthe, Dedeckera, Dodecahema, Eriogonella, Eriogonum, Eucycla, Gilmania, Good-
mania, Harfordia, Hollisteria, Lastarriaea, Mucronea, Nemacaulis, Oxytheca, Phyllogonum,
Pterogonum, Pterostegia, Sanmartinia, Stenogonum, Systenotheca.

INTRODUCTION

In the series of papers on Polygonaceae subf. Eriogonoideae published in this volume of
Phytologia, attempts have been made to summarize the present taxonomic knowledge on the
genera that make up the subfamily. Although no complete revision of Eriogonum has been
published yet, the general overall scheme seems clear. Some undescribed entities are noted
here, and I am aware of one species not yet placed. The lack of type material is currently
preventing these entities from being proposed.

The treatment of the perennial species of Chorizanthe (species numbers 254-263) must be
regarded as highly tentative. Authentic type material of some of the species has not yet been
seen, and the lack of field experience compounds my overall ignorance of the group.

Synonyms are listed by date of publication with the basionym cited first followed by its
nomenclatural synonyms, if any. Nomenclatural synonyms of the accepted names are given
immediately following those names. Full author citations are given in all instances. The largest
genus is Eriogonum with 240 species, followed by Chorizanthe with 51. A total of 17 genera
and 316 species are listed.

LIST OF ENTITIES, INDEX AND LITERATURE
The following listing is divided into three parts. The first is a numeric summary of the
Eriogonoideae; the second is an index to all published names; the third is a listing of the
more important systematic literature on the Eriogonoideae.

ACKNOWLEDGMENTS

This is Scientific Article A-4943, Contribution Number 7986, Maryland Agricultural Ex-
periment Station.

266

Reveal: Checklist of the Eriogonoideae (Polygonaceae) 267

1. ERIOGONUM Michx.
I. Subg. Eucycla Nutt.
1. E. microthecum Nutt.
la. var. laxiflorum Hook.
E. microthecum subsp. laxiflorum (Hook.) S.
Stokes
E. confertiflorum Benth. in A.DC.-E. micro-
thecum var. confertiflorum (Benth. in A.DC.)
Torr. & A. Gray-E. microthecum subsp. con-
fertiflorum (Benth. in A.DC.) S. Stokes
E. spathulare Gandoger, nom. illeg.—E. micro-
thecum var. spathulare S. Stokes
E. intricatum Gandoger, nom. illeg., non Benth. in
Hinds.
E. tenellum Tort. var. grandiflorum Gandoger
E. tenellum Torr. var. sessiliflorum Gandoger
1b. var. simpsonii (Benth. in A.DC.) Rev.
E. simpsonii Benth. in A.DC.-E. effusum Nutt.
subsp. simpsoni (Benth. in A-.DC.) S. Stokes
E. effusum Nutt. var. foliosum Torr. & A. Gray-E.
microthecum var. foliosum (Torr. & A. Gray)
Rev.
E. microthecum var. rigidum Eastw.-E. micro-
thecum subsp. rigidum (Eastw.) S, Stokes
E. friscanum MLE. Jones—E. microthecum var. fris-
canum (M.E. Jones) S. Stokes
E. macdougalii Gandoger, nom. illeg.-E. micro-
thecum var. macdougalii S. Stokes
E. nelsoni L.O. Williams —E. effusum Nutt. subsp.
nelsonu (L.O. Williams) S. Stokes
E. microthecum subsp. intermedium S. Stokes
lc. var. panamintense S. Stokes
E. effusum Nutt. var. limbatum S. Stokes
1d. var. corymbosoides Rev.
le. var. johnstonii Rev.
If. var. lapidicola Rev.
1g. var. alpinum Rev.
lh. var. ambiguum (M.E. Jones) Rev. in Munz
E. aureum M.E. Jones var. ambiguum M.E.
Jones-E. fruticosum A. Nelson var. ambiguum
(M.E. Jones) A. Nelson—E. corymbosum Benth.
in A.DC. var. ambiguum (MLE. Jones) S. Stokes
E. tenellum Torr. var. erianthum Gandoger
E. microthecum var. expansum S. Stokes
1i. var. microthecum
E. microthecum subsp. typicum S. Stokes
E. idahoense Rydb.-E. microthecum var. idaho-
ense (Rydb.) S. Stokes
2. E. effusum Nutt.
2a. var. effusum
E. microthecum Nutt. var. effusum (Nutt.) Torr &
A. Gray-E. effusum subsp. typicum S. Stokes
E. myrianthum Gandoger, nom. illeg.
2b. var. rosmarnioides Benth. in A.DC.
E. helichrysoides Gandoger, nom. illeg.-E.
helichrysoides Rydb.-E. effusum subsp. heli-
chrysoides (Rydb.) S. Stokes

3. E. leptocladon Torr. & A. Gray
3a. var. ramossimum (Eastw.) Rev.
E. ramosissimum Eastw.—E. eastwoodae M.E.
Jones
E. pallidum Small-E. effusum Nutt. var. pallidum
(Small) S. Stokes
3b. var. papiliunculi Rev.
3c. var. leptocladon
E. microthecum Nutt. var. leptocladon (Torr. & A.
Gray) Torr. & A. Gray-E. effusum Nutt. subsp.
leptocladon (Torr. & A. Gray) S. Stokes
E. effusum Nutt. var. shandsii S. Stokes
4. E. nummulare M.E. Jones
E. kearneyi Tidestrom-E. nodosum Small var.
kearneyi (Tidestrom) S. Stokes
E. dudleyanum §. Stokes
E. nodosum Small subsp. monoense S. Stokes —E.
kearneyi var. monoense (S. Stokes) Rev.-E.

kearneyi subsp. monoense (S. Stokes) Munz ex
Rev.

5. E. ammophilum Rey.

E. nummulare M.E. Jones var. ammophilum

(Rev.) Welsh
6. E. leptophyllum (Torr. in Sitgreaves) Wooton &
Standley

E. effusum Nutt. var. leptophyllum Torr. in
Sitgreaves —E. microthecum Nutt. var. leptophyl-
lum (Torr. in Sitgreaves) Torr. & A. Gray

7. E. clavellatum Small
8. E. bicolor M.E. Jones

E. microthecum Nutt. subsp. bicolor (M.E. Jones)

S. Stokes
9. E. riplelyi J.T. Howell
10. E. ericifolium Torr. & A. Gray

10a. var. pulchrum (Eastw.) Rev.

E. pulchrum Eastw.-E. microthecum Nutt. subsp.
pulchrum (Eastw.) S. Stokes—E. mearnsii Parry
in Bnitton var. pulchrum (Eastw.) Kearney &
Peebles

10b. var. ericifolium

E. fasciculatum Benth. var. ericifolium (Torr. & A.
Gray) M.E. Jones—E. microthecum Nutt. subsp.
ericifolium (Torr. & A. Gray) S. Stokes

E. mearnsii Parry in Britton-E. microthecum Nutt.
var. mearnsii (Parry in Britton) S. Stokes

10c. var. thornei Rev. & Henrickson

E. ericifolium subsp. thornei (Rev. & Henrickson)
Thorne

11. E. smithii Rev.

E. corymbosum Benth. in A.DC. var. smitiii (Rev.)

Welsh
12. E. mortonianum Rev.
13. E. corymbosum Benth. in A.DC.

13a. var. corymbosum

E. effusum Nutt. subsp. corymbosum (Benth. in
A.DC.) S. Stokes

268 PHYTOLO GIA volume 66(3):266-294 May 1989

E. corymbosum var. divaricatum Torr. & A. Gray-
E. divergens Small-E. effusum Nutt. subsp.
divaricatum (Torr. & A. Gray) S. Stokes

13b. var. erectum Rev. & Brotherson

E. salinum A. Nelson-E. effusum Nutt. subsp.
salinum (A. Nelson) S. Stokes

13c. var. davidsei Rev.

13d. var. revealianum (Welsh) Rev.

E. revealianum Welsh

13e. var. orbiculatum (S. Stokes) Rev. & Brother-
son

E. effusum Nutt. subsp. orbiculatum S. Stokes

13f. var. velutinum Rev.

13g. var. aureum (M.E. Jones) Rev.

E. aureum M.E. Jones—E. fruticosum A. Nelson,
nom. superfl.—E. microthecum subsp. aureum
(M.E. Jones) S. Stokes

E. aureum M.E. Jones var. glutinosum M.E.
Jones-—E. fruticosum A. Nelson var. glutinosum
(M.E. Jones) A. Nelson—E. corymbosum var.
glutinosum (M.E. Jones) M.E. Jones

E. crispum L.O. Williams—E. microthecum Nutt.
var. crispum (L.O. Williams) S. Stokes

14. E. lancifolium Rev. & Brotherson
E. effusum Nutt. subsp. durum S. Stokes
15. E. hylophilum Rev. & Brotherson
16. E. jonesii S. Watson
E. lanosum Eastw.
17. E. lonchophyllum Torr. & A. Gray

17a. var. lonchophyllum

E. salicinum E. Greene-E. effusum Nutt. subsp.
salicinum (E. Greene) S. Stokes

E. scoparium Small-E£. nudicaule (Torr. in
Whipple) Small subsp. scoparium (Small) S.
Stokes

E. tristichum Small-E. nudicaule (Torr. in
Whipple) Small subsp. sristichum (Small) S.
Stokes

E. sarothriforme Gandoger, nom. illeg.

E. humivagans Rev.-E. corymbosum Benth. in
A.DC. var. humivagans (Rev.) Welsh

17b. var. fendlerianum (Benth. in A.DC.) Rev.

E. microthecum Nutt. var. fendlerianum Benth. in
A.DC.-E. fendlerianum (Benth. in A.DC.)
Small-E. effusum Nutt. subsp. fendlerianum
(Benth. in ADC.) S. Stokes

E. ainliei Wooton & Standley-E. effusum Nutt.
subsp. ainliei (Wooton & Standley) S. Stokes

17c. var. nudicaule (Torr. in Whipple) Rev.

E. effusum Nutt. var. nudicaule Torr. in Whip-
ple -E. nudicaule (Torr. in Whipple) Small -E.
nudicaule (Torr. in Whipple) Small subsp.
typicurn S. Stokes

17d. var. saurinum (Rev.) Welsh

E. saurinum Rev.

17e. var. intermontanum (Rev.) Rev.

E. intermontanum Rev.
18. E. thompsonae S. Watson

18a. var. thompsonae

18b. var. albiflorum Rev.

18c. var. atwoodii Rev.

18d. var. matthewsae (Rev.) Rev.

E. corymbosum Benth. in A.DC. var. matthewsae
Rev.

19. E. fasciculatum Benth.

19a. var. polifolium (Benth. in A.DC.) Torr. & A.
Gray

E. polifolium Benth. in A.DC.-E. fasciculatum
subsp. polifolium (Benth. in A-.DC.) S. Stokes

E. revolutum Goodd.-E. fasciculatum var.
revolutum (Goodd.) S. Stokes

19b. var. flavovirde Munz & I.M. Johnston

E. fasciculatum subsp. flavovirde (Munz & I.M.
Johnston) S. Stokes

19c. var. emphereium Rev.

19d. var. fasciculatum

E. fasciculatum subsp. typicum S. Stokes

E. rosmarinifolium Nutt.

E. fasciculatum var. oleifolium Gandoger

E. fasciculatum var. aspalathoides Gandoger-E.
fasciculatum subsp. aspalathoides (Gandoger)
S. Stokes

E. fasciculatum var. maritimum Parish

19e. var. foliolosum (Nutt.) S. Stokes ex Abrams

E. rosmarinifolium Nutt. var. foliolosum Nutt.—E.
fasciculatum subsp. foliolosum (Nutt.) S. Stokes

E. fasciculatum var. obtusifolium S. Stokes

20. E. cinereum Benth. in Hinds
21. E. parvifolium Smith in Rees

E. parvifolium subsp. typicum S. Stokes

E. parvifolium vat. crassifolium Benth.

E. parvifolium subsp. lucidum J.T. Howell ex S.
Stokes-E. parvifolium var. lucidum (J.T.
Howell ex S. Stokes) Rev. in Munz

E. parvifolium subsp. paynei C. Wolf ex Munz-—E.
parvifolium var. paynei (C. Wolf ex Munz) Rev.
in Munz

22. E. giganteum S. Watson

22a. var. giganteum

22b. var. compactum Dunkle

22c. var. formosum K. Brandegee

E. formosum (K. Brandegee) Brandegee -E.
giganteum subsp. formosum (K. Brandegec)
Raven

23. E. zapatoense Moran

24. E. encelioides Rev. & Hanson
25. E. molle Brandegee

26. E. arborescens E. Greene

E. x blissianum Mason, a named hybrid between
E. arborescens and varieties of E. giganteum

27. E. pondii E. Greene
E. pondii var. gentryi Rev. & Hanson

Reveal: Checklist of the Eriogonoideae (Polygonaceae) 269

28. E. orcuttianum S. Watson
E. peninsulare K. Brandegee
29. E. suffruticosum S. Watson
30. E. fastigiatum C. Parry
31. E. deserticola S. Watson
32. E. pendulum S. Watson
33. E. heermannii Durand & Hilg.
33a. var. humilius (S. Stokes) Rev.
E. heermannut subsp. humilius S. Stokes
33b. var. heermannii
E. geniculatum Durand & Hilg., non Nutt.-E.
heermanntt subsp. typicum S. Stokes
33c. var. clokeyi Rev.
33d. var. occidentale S. Stokes
E. heermannii subsp. occidentale (S. Stokes) S.
Stokes
33e. var. floccosum Munz
E. heermannii subsp. floccosum (Munz) Munz
33f. var. subracemosum (S. Stokes) Rev.
E. howellii S. Stokes var. subracemosum S. Stokes
33g. var. argense (M.E. Jones) Munz
E. sulcatum S. Watson var. argense M.E. Jones—E.
howellit S. Stokes var. argense (M.E. Jones) S.
Stokes—E. heermannii subsp. argense (M.E.
Jones) Munz
E. howellii S. Stokes
33h. var. sulcatum (S. Watson) Munz & Rev. in
Munz
E. sulcatum S. Watson-E. heermannii subsp. sul-
catum (S. Watson) S. Stokes
34. E. apachense Rev.
35. E. plumatella Durand & Hilg.
E. plumatella var. typicum S. Stokes
E. palmeri S. Watson
E. nodosum Small var. jaegeri Munz & I.M.
Johnston -E. plumatella var. jaegeri (Munz &
I.M. Johnston) S. Stokes ex Munz
36. E. apricum J.T. Howell
36a. var. apricum
36b. var. prostratum Myatt
37. E. brevicaule Nutt.
37a. var. brevicaule
E. campanulatum Nutt. subsp. brevicaule (Nutt.)
S. Stokes—E. brevicaule subsp. typicum S.
Stokes
E. campanulatum Nutt.-E. campanulatum subsp.
Qpicum S. Stokes—E. brevicaule subsp. cam-
panulatum (Nutt.) S. Stokes
E. confertiflonim Benth. in A.DC. var. stansburyi
Benth. in A.DC.
E. brevicaule var. aureum Benth. in A.DC.
E. sabulosum M.E. Jones
E. grangerense M.E. Jones—E. campanulatum
Nutt. subsp. grangerense (M.E. Jones) S.
Stokes -E. brevicaule subsp. grangerense (M.E.
Jones) S. Stokes

E. campanulatum Nutt. subsp. leptothecum S.
Stokes—E. brevicaule subsp. leptothecum (S.
Stokes) S. Stokes—E. brevicaule var. leptot-
hecum (S. Stokes) Rev.

E. nudicaule Torr. in Whipple subsp. garrewii S.
Stokes

E. nudicaule Torr. in Whipple subsp. paraleyense
S. Stokes

E. x duchesnense Rev., a hybrid complex involving
variants of E. brevicaule and E. corymbosum

E. corymbosum Benth. in A.DC. var. albogilum
Rev., part of the above hybrid complex

37b. var. micranthum (Nutt.) Rev. in Rev. & Spevak

E. micranthum Nutt.

E. orendense A. Nelson-E. campanulatum Nutt.
subsp. orendense (A. Nelson) S. Stokes—E.
brevicaule subsp. orendense (A. Nelson) S.
Stokes

37¢c. var. wasatchense (M.E. Jones) Rev.

E. wasatchense M.E. Jones

37d. var. cottamii (S. Stokes) Rev.

E. tenellum Torr. var. cottamii S. Stokes

37e. var. laxifolium (Torr. & A. Gray) Rev.

E. kingtt Torr. & A. Gray var. laxifolium Torr. &
A. Gray-E. chrysocephalum A. Gray-E. laxi-
folium (Torr. & A. Gray) A. Nelson-E.
chrysocephalum subsp. typicum S. Stokes

E. ochrocephalum S. Watson var. angustum M.E.
Jones—E. nudicaule (Torr. in Whipple) Small
subsp. angustum (M.E. Jones) S. Stokes

E. brevicaule var. pumilum S. Stokes ex M.E.
Jones-E. nudicaule (Torr. in Whipple) Small
subsp. pumilum (S. Stokes ex M.E. Jones) S.
Stokes

E. medium Rydb.

E. chrysocephalum A. Gray subsp. bannockense S.
Stokes -E. bannockense (S. Stokes) RJ. Davis

E. grayi Rev.

E. nanum Rev.-E. brevicaule var. nanum (Rev.)
Welsh

E. brevicaule var. promiscuum Welsh

38. E. lagopus Rydb.

E. multiceps Nees in Wied-Neuw. subsp. canum S.
Stokes-E. pauciflorum Pursh var. canum (S.
Stokes) Rev.-E. brevicaule Nutt. var. canum (S.
Stokes) Dorn

39. E. viridulum Rev.

E. brevicaule Nutt. var. viridulum (Rev.) Welsh

40. E. ephedroides Rev.

E. brevicaule Nutt. var. ephedroides (Rev.) Welsh

41. E. spathulatum A. Gray

E. spathulatum var. typicum S. Stokes
E. nudicaule (Tort. in Whipple) Small subsp.
ochroflorum S. Stokes

42. E. sp. (undescribed)
43. E. natum Rev.

270 PH YTOLOGIA volume 66(3):266-294 May 1989

E. spathulatum A. Gray var. natum (Rev.) Welsh
44. E. loganum A. Nelson
44a. var. loganum

E. chrysocephalum A. Gray subsp. loganum (A.
Nelson) S. Stokes—E. brevicaule Nutt. var.
loganum (A. Nelson) Welsh

44b. var. (undescribed).
45. E. brandegei Rydb.

E. spathulatum A. Gray var. brandegei (Rydb.) S.

Stokes
46. E. contortum Small ex Rydb.

E. effusum Nutt. subsp. contortum (Small ex

Rydb.) S. Stokes
47. E. pelinophilum Rev.
48. E. acaule Nutt.

E. caespitosum Nutt. subsp. acaule (Nutt.) S.
Stokes—E. caespitosum var. acaule (Nutt.) R.
Davis

49. E. gypsophilum Wooton & Standley
50. E. ostlundii M-E. Jones

E. tenellum Torr. subsp. ostlundii (M.E. Jones) S.
Stokes—E. batemanti M.E. Jones var. ostlundii
(M.E. Jones) Welsh

E. spathuiforme Rydb.—-E. spathulatum A. Gray
subsp. spathuiforme (Rydb.) S. Stokes

51. E. eremicum Rey.

E. batemanii M.E. Jones var. eremicum (Rev.)

Welsh
52. E. batemanii M.E. Jones
53. E. cronquistii Rev.

E. corymbosum Benth. in A.DC. var. cronquistii

(Rev.) Welsh
54. E. panguicense (M.E. Jones) Rev.

54a. var. panguicense

E. pauciflorum Pursh var. panguicense M.E.
Jones— E. spathulatum A. Gray var. pan-
guicense (M.E. Jones) S. Stokes

54b. var. alpestre (S. Stokes) Rev.

E. chrysocephalum A. Gray subsp. alpestre S.
Stokes

55, E. lachnogynum Torr. ex Benth. in A.DC.

E. lachnogynum subsp. typicum S. Stokes

E. tetraneuris Small-E. lachnogynum subsp.
tetraneuris (Small) S. Stokes

56. E. havardii S. Watson

E. leucophyllum Wooton & Standley —E. leucophyl-

lum subsp. typicum S. Stokes
57. E. desertorum (Maguire) R. Davis

E. chrysocephalum subsp. desertorum Maguire - E.
brevicaule Nutt. var. desertorum (Maguire)
Welsh

58. E. lewisii Rev.
59. E. ochrocephalum S. Watson
59a. var. ochrocephalum
E. nevadense Gandoger
59b. var. calcareum (S. Stokes) M.E. Peck

E. ochrocephalum subsp. calcareum S. Stokes
59c. var. sceptrum Rev.
59d. var. alexandreae Rev.
60. E. novonudum M.E. Peck
61. E. cusickii M.E. Jones non Gandoger
E. chrysocephalum A. Gray subsp. cusickit (M.E.
Jones) S. Stokes
E. anemophilum E. Greene var. cusickii Gandoger
62. E. tiehmii Rev.
63. E. prociduum Rev.
64. E. crosbyae Rev.
65. E. rosense A. Nelson & Kennedy
E. ochrcephalum S. Watson var. agnellum Jepson —
E. ochrocephalum subsp. agnellum (Jepson) S.
Stokes
66. E. beatleyae Rev.
67. E. anemophilum E. Greene
68. E. breedlovei (J.T. Howell) Rev.
68a. var. breedlovei
E. ochrocephalum S. Watson var. breedlovet J.T.
Howell
68b. var. shevockii J.T. Howell
69. E. scopulorum Rev.
70. E. chrysops Rydb.
E. ochrocephalum S. Watson subsp. chrysops
(Rydb.) S. Stokes
71. E. sp. (undescribed)
72. E. verrucosum Rev.
73. E. meledonum Rev.
74. E. capistratum Rev.
74a. var. capistratum
74b. var. muhlickii Rev.
74c. var. welshii Rev.
75. E. mancum Rydb.
E. chrysocephalum A. Gray subsp. mancum
(Rydb.) S. Stokes
76. E. kingii S. Watson
77. E. argophyllum Rev.
78. E. gracilipes S. Watson
E. kennedyi Porter ex S. Watson subsp. gracilipes
(S. Watson) S. Stokes—E. ochrocephalum S.
Watson var. gracilipes (S. Watson) J.T. Howell
79. E. holmgrenii Rev.
80. E. exilifolium Rev.
81. E. coloradense Small
E. multiceps Nees in Wied.-Neuw. subsp. colo-
radense (Small) S. Stokes
82. E. pauciflorum Pursh
82a. var. pauciflorum
E. parviflorum Nutt., nom. superfl.
E. dioecium Raf.
E. multiceps Nees in Wied-Neuw.-E. mutlticeps
subsp. typicum S. Stokes
E. depauperatum Small
82b. var. gnaphaloides (Benth. in Hook.) Rev.
E. gnaphaloides Benth. in Hook.

Reveal: Checklist of the Eriogonoidcae (Polygonaceae) Dall

83. E. villiflorum A. Gray

E, villiflorum var. typicum S. Stokes
84. E. tumulosum (Barneby) Rev.

E. villiflorum A. Gray var. tumulosum Barneby
85. E. soredium Rev.
86. E. aretioides Barneby
87. E. shockleyi S. Watson

87a. var. shockleyi

E. acaule Nutt. var. shockleyi (S. Watson) M_E.
Jones— E. shockleyi subsp. typicum S. Stokes

E. villiflorum A. Gray var. candidum M.E.
Jones-E. shockleyi subsp. candidum (M.E.
Jones) S. Stokes

E. longilobum M.E. Jones—E. acaule Nutt. var.
longilobum (M.E. Jones) M.E. Jones—E. shock-
leyi subsp. longilobum (ME. Jones) S. Stokes —
E. shockleyi var. longilobum (M.E. Jones) Rev.

87b. var. packardae Rev.
88. E. elongatum Benth. in Hinds
88a. var. elongatum
E. denudatum Nutt., non Curran
88b. var. areorivum Rev.
88c. var. vollmeri (Wiggins) Rev.
E. vollmeri Wiggins ‘
89. E. wrightii Torr. ex Benth. in A.DC.
89a. var. wrightii

E. trachygonum Torr. ex Benth. in A.DC. subsp.
wrightii (Torr. ex Benth. in A.DC.) S. Stokes —
E. wrightii subsp. typicum S. Stokes

E. wrightii var. floccosum Benth. in A.DC.

E. helianthemifolium Benth. in A.DC.-E. wright
var. helianthemifolium (Benth. in A.DC.) Torr.
in Emory

E. trachygonum Torr. ex Benth. in A.DC. subsp.
glomerulum S. Stokes - E. wrightit subsp. glomer-
ulum (S. Stokes) S. Stokes

89b. var. trachygonum (Torr. ex Benth. in A.DC.)
Jepson

E. trachygonum Torr. ex Benth. in A-DC.-E. tra-

chygonum subsp. typicum S. Stokes
89c. var. membranaceum S. Stokes ex Jepson

E. trachygonum Torr. ex Benth. in A-DC. subsp.
membranaceum (S. Stokes ex Jepson) S.
Stokes—E. wrightit subsp. membranaceum (S.
Stokes ex Jepson) S. Stokes

89d. var. dentatum (S. Stokes) Rev.

E. trachygonum Torr. ex Benth. in A.DC. subsp.
dentatum S. Stokes-E. wrighttt subsp. dentanum
S. Stokes

89e. var. taxifolium (E. Greene) Parish

E. taxifolium E. Greene -E. trachygonum Torr. ex
Benth. in A.DC. subsp. taxifolium (E. Greene)
S. Stokes—E. wrightiti subsp. taxifolium (E.
Greene) S. Stokes

89f. var. pringlei (Coulter & Fisher) Rev.

E. pringlei Coulter & Fisher-E. trachygonum Torr.
ex Benth. in A.DC. subsp. pringlei (Coulter &
Fisher) S. Stokes—E. wright subsp. pringlet

(Coulter & Fisher) S. Stokes
89g. var. nodosum (Small) Rev.
E. nodosum Small
89h. var. subscaposum S. Watson
E. trachygonum Torr. ex Benth. in A.DC. var. sub-
scaposum (S. Watson) M.E. Jones-E. tra-
chygonum Torr. ex Benth. in A.DC. subsp. sub-
scaposum (S. Watson) S. Stokes—E. wrightii
subsp. subscaposum (S. Watson) S. Stokes
E. curvatum Small-E. wrightit var. curvatum
(Small) Munz
E. kennedyi Porter ex S. Watson subsp. pinorum S.
Stokes
E. junceum E. Greene
89i. var. olanchense (J.T. Howell) Rev. in Munz
E. kennedyi Porter ex S. Watson var. olanchense
J.T. Howell
89j. var. oresbium Rev.
90. E. kennedyi Porter ex S. Watson
90a. var. kennedyi
E. kennedyi subsp. typicum S. Stokes
90b. var. austromontanum Munz & Johnston
E. kennedyi subsp. austromontanum (Munz &
Johnston) S. Stokes
90c. var. alpigeum (Munz & Johnston) Munz &
Johnston in Munz
E. kennedyi f. alpigeurm Munz & Johnston -E. ken-
nedyi subsp. alpigeum (Munz & Johnston) S.
Stokes
90d. var. purpursii (Brandegee) Rev.
E. purpursii Brandegee-E. kennedyi subsp. pur-
pusti (Brandegee) Munz
90e. var. pinicola Rev.
91. E. butterworthianum J.T. Howell
92. E. panamintense C. Morton
92a. var. panamintense
E. reliquum S. Stokes
E. racemosum Nutt. var. desertorum S. Stokes
92b. var. mensicola (S. Stokes) Rev. in Munz
E. mensicola S. Stokes—E. panamintense subsp.
mensicola (S. Stokes) Munz
93. E. rupinum Rev.
94. E. racemosum Nutt.
E. racemosum vat. typicum S. Stokes
E. orthocladon Torr. in Sitgreaves-E. racemosum
var. orthocladon (Torr. in Sitgreaves) S. Stokes
E. obtusum Benth. in A.DC.-E. racemosum var.
obtusum (Benth. in A.DC.) S. Stokes
E. racemosum var. sagittatum Gandoger
E. racemosum var. cordifolium Gandoger
95. E. zionis J.T. Howell
95a. var. zionis
E. racemosum Nutt. var. zionis (J.T. Howell)
Welsh
95b. var. coccineum J.T. Howell
E. racemosum Nutt. var. coccineum (J.T. Howell)
Welsh

272 PHYTOL O G1A volume 66(3):266-294 May 1989

96. E. nudum Douglas ex Benth.
96a. var. nudum
E. latifolium Smith in Rees subsp. nudum (Doug-
las ex Benth.) S. Stokes
E. longulum E. Greene
E. oblanceolatum E. Greene
E. latifolium Smith in Rees var. parvulum S. Stokes
96b. var. paralinum Rev.
96c. var. deductum (E. Greene) Jepson
E. deductum E. Greene
96d. var. scapigerum (Eastw.) Jepson
E. scapigerum Eastw.-E. latifolium Smith in Rees
var. scapigerum (Eastw.) S. Stokes
96e. var. pauciflorum S. Watson
E. nudum var. perturbum M.E. Jones, nom. super-
fl.—E. latifolium Smith in Rees subsp. pauct-
florum (S. Watson ) S. Stokes
96f. var. pubiflorum Benth. in A.DC.
96g. var. westonii (S. Stokes) J.T. Howell
E. latifoliurn Smith in Rees subsp. westonii S.
Stokes
E. saxicola A. Heller—E. latifolium Smith in Rees
subsp. saxicola (A. Heller) S. Stokes—E. nudum
subsp. saxicola (A. Heller) Munz-E. nudum
var. saxicola (A. Heller) Rev.
E. gramineum S. Stokes-E. nudum vat. gramin-
eum (S. Stokes) Rev.
96h. var. murinum Rev.
96i. var. auriculatum (Benth.) Tracy ex Jepson
E. auriculatum Benth.—E. latifolium Smith in
Rees subsp. auriculatum (Benth.) S. Stokes
E. latifolium Smith in Rees var. alternans S. Stokes
96j. var. indictum (Jepson) Rev. in Munz
E. indictum Jepson-—E. latifolium Smith in Rees
var. indictum (Jepson) S. Stokes
96k. var. regirivum Rev. & J. Stebbins
961. var. decurrens (S. Stokes) Bowerman
E. latifolium Smith in Rees subsp. decurrens S.
Stokes
96m. var. oblongifolium S. Watson
E. affine Benth. in A.DC.-E. latifolium Smith in
Rees var. affine (Benth. in A.DC.) S. Stokes
E. harfordti Smail—E. latifolium Smith in Rees var.
harfordti (Small) S. Stokes
E. sulphureum E. Greene-E. nudum var. sul-
phureum (E. Greene) Jepson-E. latifolium
Smith in Rees subsp. su/phureum (E. Greene)
S. Stokes
E. capitatum A. Heller
97. E. latifolium Smith in Rees
E. latifolium subsp. typicum S. Stokes
E. arachnoideum Eschsch.
E. oblongifolium Benth.
98. E. grande E. Greene
98a. var. grande
E. nudum Douglas ex Benth. var. grande Jep-
son-E. latifolium subsp. grande (E. Greene) S.
Stokes

98b. var. testudinum Rev.
98c. var. timorum Rev.

E. grande subsp. timorum (Rev.) Munz

98d. var. rubescens (E. Greene) Munz

E. rubescens E. Greene -E. latifolium Smith in
Rees subsp. rubescens (E. Greene) S. Stokes —
E. latifolium Smith in Rees var. rubescens (E.
Greene) Munz

E. grande var. dunklei Rev.

99. E. elatum Douglas ex Benth.
99a. var. elatum

E. elatum vat. limonifolium Gandoger

E. elatum var. erianthum Gandoger

99b. var. villosum Jepson

E. elatum subsp. villosum (Jepson) Munz ex Rev.

E. elatum var. incurvum Jepson
100. E. gilmanii S. Stokes
101. E. ovalifolium Nutt.

101a. var. purpureum (Nutt.) Durand

E. tenellum Nutt., non Torr.-E. nuttallii Gambel
in Nutt.—E. purpureum Nutt. var. tenius Benth.
in A.DC.

Eucycla purpurea Nutt.-—Eriogonum purpureum
(Nutt.) Benth. in A.DC.-Eriogonum ovali-
foliur subsp. purpureurm (Nutt.) S. Stokes

E. oblongifolium Benth. var. minus Benth. in
A.DC.

E. orthocaulon Small-E. ovalifolium var. celsum
A. Nelson-E. ovalifolium var. orthocaulon
(Small) C.L. Hitche.

. roseiflorum Gandoger, nom. illeg.

. ovalifolium var. utahense Gandoger

. ovalifolium var. cyclophyllum Gandoger

ovalifolium var. multiscapum Gandoger

. ovalifolium var. cerastoides Gandoger

. davisianum S. Stokes

101b. var. ovalifolium

Eucycla ovalifolia (Nutt.) Nutt.-Eriogonum
ovalifolium var. typicum Gandoger—Enogonum
ovalifoliur subsp. typicum S. Stokes

E. ovalifolium var. nevadense Gandoger

E. ovalifolium var. deltoideum Gandoger

101c. var. eximium (Tidestrom) J.T. Howell

E. eximium Tidestrom-E. ovalifolium subsp. ex-
imium (Tidestrom) S. Stokes

101d. var. williamsae Rev.

101e. var. vineum (Small) Jepson

E. vineum Small-E. ovalifolium subsp. vineum

(Small) S. Stokes
101f. pansum Rev.
101g. var. ochroleucum (Small ex Rydb.) M.E. Peck
E. ochroleucum Small ex Rydb.-E. ovalifolium

subsp. ochroleucum (Small ex Rydb.) S. Stokes
E. ochroleucum Small ex Rydb. var. macropodum

Gandoger-E. ovalifolium var. macropodum

(Gandoger) Rev.

moh hhh

Reveal: Checklist of the Eriogonoidcac (Polygonaceac) 273

E. ochroleucum Small ex Rydb. var. decalvans
Gandoger

101h. var. depessum Blank.

E. depressum (Blank.) Rydb.

E. rubidum Gandoger, nom. illeg.

E. rubidum Gandoger var. frigidum Gandoger,
nom. illeg.

101i. var. nivale (Canby in Cov.) M.E. Jones

E. nivale Canby in Cov.

E. rhodanthum A. Nelson & Kennedy

101j. var. caelestinum Rev.

102. E. strictum Benth.
102A. subsp. proliferum (Torr. & A. Gray) S. Stokes
102a. var. proliferum (Torr. & A. Gray) Rev.

E. proliferum Torr. & A. Gray-E. ovalifolium
Nutt. var. proliferum (Torr. & A. Gray) S. Wat-
son

E. cusickit Gandoger, nom. illeg., non M.E.
Jones—E. strictum var. cusickii S. Stokes

E. cusickii Gandoger var. californicum Gandoger

E. ovalifolium Nutt. var. bellum S. Stokes - E. stric-
tum subsp. bellum (S. Stokes) S. Stokes —E. bel-
lum (S. Stokes) R. Davis

E. strictum var. argenteum S. Stokes

E. fulvum S. Stokes

102b. var. greenei (A. Gray) Rev.

E. greenet A. Gray—E. niveum Douglas ex Benth.
var. greenei (A. Gray) S. Stokes

102c. var. glabrum C.L. Hitche.

102d. var. anserinum (E. Greene) R. Davis

E. anserinum E. Greene -E. ovalifolium Nutt. var.
anserinum (E. Greene) M.E. Jones—FE. pro-
liferum Torr. & A. Gray subsp. anserinum (E.
Greene) Munz

E. flavissimum Gandoger, nom. illeg.-E. ovali-
folium Nutt. subsp. flavissimum S. Stokes-E.
Strictum var. flavissimum (S. Stokes) C.L.
Hitche.

102B. subsp. strictum
E. strictum subsp. typicum S. Stokes
103. E. niveum Douglas ex Benth.

E. niveum subsp. typicum S. Stokes

E. dichotomum Douglas ex Benth. -E. niveum var.
dichotomum (Douglas ex Benth.) S. Stokes ex
M.E. Jones-E. niveum subsp. dichotomum
(Douglas ex Benth.) S. Stokes

E. decumbens Benth.—E. niveurn var. decumbens
(Benth.) Torr. & A. Gray-F. ntveum subsp.
decumbens (Benth.) S. Stokes

E. album Nutt.- E. dichotomum Douglas ex Benth.
var. humile Benth. in A.DC.

E. strictuum Benth. var. lachnostegium Benth. in
A.DC.-E. lachnostegium (Benth. in A.DC.)
Rydb.

E. niveum var. suksdorfti Gandoger

E. niveurm var. candelabrum Gandoger

104. E. saxatile S. Watson
E. bloomert Parish
E. stokesae M.E. Jones—E. saxatile var. stokesae
(M.E. Jones) S. Stokes ex M.E. Jones
E. saxaule subsp. multicaule S. Stokes
105. E. crocatum Davidson
E. saxatile S. Watson var. crocatum (Davidson)
Munz
II. Subg. Micrantha (Benth.) Rev. in Gunckel
106. E. annuum Nutt.
E. annuum subsp. typicum S. Stokes
E. annuum Raf., non Nutt.
E. lindheimerianum Scheele
E. stmpsoni Benth. in A.DC. var. floccoso-lanatum
Benth. in A.DC.
E. cymosum Benth. in A.DC.—E. annuum subsp.
cymosum (Benth. in A.DC.) S. Stokes
E. annuum var. pauciflorum Gandoger
E. hitchcockii Gandoger, nom. illeg.—E. annuum
subsp. hitchcockii S. Stokes
E. annuum f. roseum E. Kelso
E. annuum subsp. chihuahuaense S. Stokes
107. E. multiflorum Benth.
IIL Subg. Clastomyelon Cov. & C. Morton
108. E. intrafractum Cov. & C. Morton
IV. Subg. Eriogonum
109. E. longifolium Nutt.
109a. var. longifolium
E. longifolium subsp. typicum S. Stokes
E. longifolium var. plantagineum Engelm. & A.
Gray
E. texanum Scheele, non Coulter & Fisher
E. texanum Coulter & Fisher, non Scheele-E.
coriaceum Coulter & Fisher
E. longifoltum var. lindheimeri Gandoger
E. longifolium var. caput-felis Gandoger
E. longifolium subsp. diffusum S. Stokes
E. vespinum Shinners
109b. var. gnaphalifolium Gandoger
E. harper: Goodman
109b. var. harperi (Goodman) Rev.
E. floridanum Small
E. longifolium var. longidens Gandoger
E. longifolium var. floridanum Gandoger
110. E. tomentosum Michx.
V. Subg. Oligogonum Nutt.
111. E. umbellatum Torr.
111a. var. umbellatum
E. umbellatum subsp. typicum S. Stokes
E. lateum Small ex Rydb.
E. cupreum Gandoger, nom. illeg.
E. umbellatum var. crandalit Gandoger
111b. var. cladophorum Gandoger
E. subalpinum E. Greene var. cladophorum (Gan-
doger) S. Stokes
E. rydbergti E. Greene

274 PHYTOLO GIA volume 66(3):266-294 May 1989

Illc. var. aureum (Gandoger) Rev.
E. glaberrimum Gandoger var. aureum Gandoger
E. neglectum E. Greene
E. azaleastrum E. Greene
E. umbelliferum Small—E. umbellatum var. intec-
tum A. Nelson in Coulter & Nelson
E. marginale Gandoger
E. umbellatum var. glabratum S. Stokes
111d. var. porteri (Small) S. Stokes
E. portert Small
lle. var. hypoleium (Piper) C.L. Hitche.
E. umbellauum subsp. hypoletum Piper
11l1f. var. nevadense Gandoger
E. reclinatum E. Greene
E. heracleoides Nutt. var. virde Gandoger
E. umbellatum var. californicum Gandoger
1llg. var. vernum Rev.
111h. var. covillei (Small) Munz & Rev. in Munz
E. covillei Small-E. ursinum S. Watson var. covil-
lei (Small) S. Stokes—E. umbellatum subsp.
covillei (Small) Munz
111i var. hausknechtii (Dammer) M.E. Jones
E. hausknechti Dammer-E. umbellatum subsp.
hausknechni (Dammer) S. Stokes
E. montanum Howell
111j. var. dichrocephalum Gandoger
E. aridum E. Greene-E. umbellatum subsp.
aridum (E. Greene) S. Stokes—E. umbellatum
var. aridum (E. Greene) C.L. Hitchc.
E. subalpinum E. Greene vat. arachnoideum Gan-
doger
111k. var. desereticum Rev.
111L var. versicolor S. Stokes
E. umbellatum subsp. versicolor (S. Stokes) Munz
111m. var. minus I.M. Johnston
E. minus (1.M. Johnston) Ewan-E. umbellatum
subsp. minus (1.M. Johnston) Munz
llin. var. polyanthum (Benth. in A.DC.) M.E.
Jones
E. polyanthum Benth. in A.DC.—E. umbellatum
subsp. polyanthum (Benth. in A.DC.) S. Stokes
E. dumosum E. Greene-E. umbellatum subsp.
dumosum (E. Greene) S. Stokes
E. modocense E. Greene-E. umbellatum var.
modocense (E. Greene) S. Stokes
1llo. var. goodmanii Rev.
1llp. var. humistratum Rev.
111q. var. speciosum (Drew) S. Stokes
E. speciosum Drew
lllr. var. majus Hook.
E. umbellatum subsp. majus (Hook.) Piper
E. subalpinum EB. Greene - E. umbellauum var. sub-
alpinum (BE. Greene) M.E. Jones—E. umbel-
latum subsp. subalpinum (E. Greene) S.
Stokes— E. heracleoides var. subalpinum (E.
Greene) R. Davis

E. subalpinum E. Greene var. vulcanicum Gan-
doger
E. subalpinum E. Greene var. stenophyllum Gan-
doger
E. subalpinum E. Greene var. subnivale Gandoger
111s. var. stellatum (Benth.) M.E. Jones
E. stellatum Benth.—E. umbellatum subsp. stel-
latum (Benth.) S. Stokes
E. ellipticum Nutt.
E. croceum Small ex Rydb.-E. umbellatum var.
croceum (Small ex Rydb.) S. Stokes
E. umbellatum var. chrysanthum Gandoger
111t. var. devestivum Rev.
1llu. var. furcosum Rev.
11lv. var. argus Rev.
1llw. var. munzii Rev.
E. umbellatum subsp. munzi (Rev.) Thorne ex
Munz
111x. var. subaridum S. Stokes
E. umbellatum subsp. subaridum (S. Stokes) Munz
E. biumbellatum Rydb.
E. ferrissii A. Nelson-E. umbellatum subsp. ferris-
sti (A. Nelson) S. Stokes
1lly. var. chlorothamnus Rev. in Munz
111z. var. bahiiforme (Torr. & A. Gray) Jepson
E. polyanthum Benth. in A.DC. var. bahiiforme
Torr. & A. Gray-E. stellatum Benth. var.
bahiiforme (Torr. & A. Gray) S. Watson -E.
umbellatum subsp. bahiiforme (Torr. & A.
Gray) S. Stokes
E. trichotomum Small
E. smallianum A. Heller—E. umbellatum var.
smallianum (A. Heller) S. Stokes
l1llaa. var. juniporinum Rev.
111bb. var. cognatum (E. Greene) Rev.
E. cognatum E. Greene-E. umbellatum subsp.
cognatum (E. Greene) S. Stokes
lllcc. var. torreyanum (A. Gray in Torr. & A.
Gray) M.E. Jones
E. torreyanum A. Gray in Torr. & A. Gray
1lidd. var. glaberrimum (Gandoger) Rev.
E. glaberrimum Gandoger
112. E. heracleoides Nutt.
112a. var. heracleoides
E. heracleoides subsp. typicum S. Stokes
E. gyrophyllum Nutt.
E. heracleoides var. minus Benth. in A.DC.
E. heracleoides var. multuiceps Gandoger
E. heracleoides var. utahense Gandoger
E. heracleoides var. rydbergu Gandoger
112b. var. angustifolium (Nutt.) Torr. & A. Gray
E. angustifolium Nutt.-E. heracleoides aubsp. an-
gustifolium (Nutt.) S. Stokes
E. douglasii Benth. in A.DC. subsp. ramosum
Piper—E. caespitosum Nutt. subsp. ramosum
(Piper) S. Stokes

Reveal: Checklist of the Eriogonoideae (Polygonaceac) 215

E. heracleoides var. micranthum Gandoger

112c. var. leucophaeum Rev.

113. E. compositum Douglas ex Benth. in Lindl.
113a. var. compositum

E. compositum subsp. typicum S. Stokes

E. compositum var. pilicaule St. John & Warren—-
E. pilicaule (St. John & Warren) G.N. Jones

E. johnstonu S. Stokes

E. compositum var. citrinum S. Stokes

113b. var. leianthum Hook.

E. tolmieanum Hook.-E. umbellatum Torr. var.
monocephalum Torr. & A. Gray-E. umbel-
latum var. tolmieanum (Hook.) M.E. Jones

E. heracleoides Nutt. var. simplex S. Watson ex
Piper—E. compositum var. simplex (S. Watson
ex Piper) St. John & Warren

113c. var. lancifolium St. John & Warren

E. compositum subsp. lancifolium (St. John &
Warren) S. Stokes

114. E. siskiyouense Small
115. E. ternatum Howell

E. umbellatum Torr. var. ternatum (Howell) S.
Stokes

E. ursinum S. Watson var. confine S. Stokes

116. E. libertini Rev. :
117. E. congdonii (S. Stokes) Rev.

E. ursinum S. Watson var. congdonit S. Stokes - E.
ternatum Howell var. congdont (S. Stokes) J.T.
Howell

118. E. ursinum S. Watson

E. ursinum var. typicum S. Stokes

E. ovatum E. Greene
119. E. nervulosum (S. Stokes) Rev.

E. ursinum S. Watson var. nervulosum S. Stokes
120. E. prattenianum Durand

120a. var. prattenianum

E. umbellatum Torr. subsp. serratum S. Stokes

120b. var. avium Rev. & Shevock

121. E. tripodum E. Greene

E. sphaerocephalum Douglas ex Benth. var. brevi-

folium S. Stokes ex M.E. Jones
122. E. thymoides Benth. in A.DC.

E. thymoides subsp. typicum S. Stokes

E. minimum Small—E. sphaerocephalum Douglas
ex Benth. subsp. minimum (Small) S. Stokes - E.
sphaerocephalum var. minimum (Small) R.
Davis

E. thymoides var. pallens Gandoger

E. thymoides subsp. congestum S. Stokes

123. E. kelloggii A. Gray

E. caespitosum Nutt. var. kelloggit (A. Gray) M.E.

Jones
124. E. sphaerocephalum Douglas ex Benth.

124a. var. sphaerocephalum

E. sphaerocephalum subsp. typicum S. Stokes

E. ellipticum Nutt. var. megacephalum Nutt.-E.
sphaerocephalum var. megacephalum (Nutt.) S.
Stokes ex M.E. Jones -E. sphaerocephalum sub-

sp. megacephalum (Nutt.) S. Stokes
E. geniculatum Nutt.—E. sphaerocephalum var.
geniculatim (Nutt.) S. Stokes
124b. var. fasciculifolium (A. Nelson) S. Stokes
E. fasciculifolium A. Nelson
E. fruticulosum S. Stokes
124c. var. halimioides S. Stokes
E. halimioides Gandoger, nom. illeg.
125. E. douglasii Benth. in A.DC.
12Sa. var. douglasii
E. caespitosum Nutt. var. douglasit (Benth. in A.
DC.) M.E. Jones—E. caespitosum Nutt. subsp.
douglasit (Benth. in A.DC.) S. Stokes
125b. var. sublineare (S. Stokes) Rev.
E. caespitosum Nutt. var. sublineare S. Stokes
E. tenue Small-E. sphaerocephalum Douglas ex
Benth. subsp. tenue (Small) Piper—E. stricturm
Benth. var. tenue (Small) R. Davis—E. douglasii
var. tenue (Small) C.L. Hitchc.
126. E. twisselmannii (J.T. Howell) Rev. in Munz
E. douglasti Benth. in A.DC. var. twisselmannii J.T.
Howell
127. E. caespitosum Nutt.
E. caespitosum subsp. typicum S. Stokes
E. andinum Nutt.
E. caespitosum Nutt. var. alyssoides Gandoger
E. sericoleucum E. Greene ex Tidestrom-E.
sphaerocephalum Douglas ex Benth. var.
sericoleucum (E. Greene ex Tidestrom) S.
Stokes
128. E. marifolium Torr. & A. Gray
E. cupulatum S. Stokes
E. marifolium var. apertum S. Stokes
129. E. incanum Torr. & A. Gray
E. marifolium Torr. & A. Gray var. incanum
(Torr. & A. Gray) M_E. Jones
E. rosulatum Small—-E. ursinum S. Watson var.
rosulatum (Small) S. Stokes
130. E. diclinum Rev.
131. E. polypodum Small
E. umbellatum Torr. var. polypodum (Small) S.
Stokes
E. ursinum S. Watson var. venosum S. Stokes ex
Smiley
132. E. jamesii Benth. in A.DC.
132a. var. flavescens S. Watson
E. jamesii subsp. flavescens (S. Watson) S. Stokes
E. arcuatum E. Greene -E. jamesii var. arcuatum
(E. Greene) S. Stokes
E. bakeri E. Greene -E. jamesii subsp. bakeri (E.
Greene) S. Stokes
E. flavum var. foliatum Gandoger
E. vegetius A. Nelson
132b. var. xantum (Small) Rev.
E. xanthum Small-E. flavum Nutt. in Fraser var.
xanthum (Small) S. Stokes

276 PHYTOLOGIA volume 66(3):266-294 May 1989

E. chloranthum E. Greene —E. flavum Nutt. in Fra-
ser subsp. chloranthum (E. Greene) S. Stokes

132c. var. wootonii Rev.

132d. var. rupicola Rev.

132e. var. jamesii

E. jamesii subsp. typicum S. Stokes

E. jamesii var. neomexicanum Gandoger

132f. var. simplex Gandoger

132g. var. undulatum (Benth. in A.DC.) S. Stokes
ex M.E. Jones

E. undulatum Benth. in A.DC.-E. jamesti subsp.
undulatum (Benth. in A.DC.) S. Stokes

133. E. turneri Rev.
134. E. flavum Nutt. in Fraser
134a. var. flavum

E. flavum subsp. typicum S. Stokes

E. sericeum Pursh-E. flavum var. sericeum
(Pursh) S. Stokes

E. lateriflorum Raf.

E. crassifolium Benth.-—E. flavum var. crassifolium
(Benth.) Benth. in A.DC.-E. flavum subsp.
crassifolium (Benth.) S. Stokes

E. crassifolium Benth. var. tectum A. Nelson-E.
flavum var. tectum (A. Nelson) S. Stokes

E. flavum var. linguifolium Gandoger

E. flavum var. incisum S. Stokes

E. flavum var. multicum S. Stokes

E.. flavum var. caudiciferum S. Stokes

134b. var. aquilinum Rev. in Hultén

134c. var. polyphyllum (Small ex Rydb.) M.E. Jones
E. polyphyllum Small ex Rydb.

134d. var. piperi (E. Greene) M.E. Jones

E. piperi E. Greene-E. androsaceum Benth. in
A.DC. var. piperi (E. Greene) M.E. Jones-E.
flavum subsp. piperi (E. Greene) S. Stokes

E. piperi E. Greene var. ochrocephalum Gandoger

E. piperi E. Greene var. longiflorum Gandoger

135, E. androsaceum Benth. in A.DC.

E. flavum Nutt. in Fraser var. androsaceum
(Benth. in A.DC.) M.E. Jones—E. flavum subsp.
androsaceum (Benth. in A.DC.) S. Stokes

136. E. correllii Rev.
137. E. allenii S. Watson
138. E. latens Jepson

E. elatum Douglas ex Benth. subsp. glabrescens S.
Stokes

E. monticola S. Stokes

139. E. hirtellum Howell & Bacig.
140. E. pyroliifolium Hook.
140a. var. coryphaeum Torr. & A. Gray

E. coryphaeum (Torr. & A. Gray) G.N. Jones

E. pyroliifolium var. bellingeranum M.E. Peck

140b. var. pyroliifolium

E. pyroliifolium var. typicum S. Stokes

141. E. lobbii Torr. & A. Gray
14la. var. lobbii

E. lobbii var. minus Torr. & A. Gray
141b. var. robustum (E. Greene) M.E. Jones
E. robustum M.E. Jones
142. E. alpinum Engelm.
VI. Subg. Pterogonum (H. Gross) Rev.
143. E. atrorubens Engelm. in Wisliz.
143a. var. atrorubens
Pterogonum atrorubens (Engelm. in Wisliz.) H.
Gross
143b. var. pseudociliatum Rev.
143c. var. intonsum Rev.
143d. var. auritulum Hess & Rev.
143e. var. nemorosum Hess & Rev.
143f. var. rupestre (S. Stokes) Hess & Rev.
E. rupestre S. Stokes
144. E. clivosum Hess & Rev.
145. E. fimbriatum Hess & Rev.
146. E. henricksonii Rev.
147. E. viscanum Hess & Rev.
148. E. ciliatum Torr. ex Benth. in A.DC.
149. E. greggii Torr. & A. Gray
E. ciliatum Torr. ex Benth. in A.DC. var. foliosum
Torr. in Emory
150. E. hemipterum (Torr. & A. Gray) S. Stokes
150a. var. hemipterum
E. hieracifolium Benth. in A.DC. var. hemipteruim
Torr. & A. Gray
E. hieracifolium Benth. in A.DC. f. at@opurpureum
Standley
150b. var. griseum I.M. Johnston
151. E. nealleyi Coulter
152. E. hieracifolium Benth. in A.DC.
Pterogonum hieracifolium (Benth. in A.DC.) H.
Gross
E. pannosum Wooton & Standley-E. leucophyl-
lum Wooton & Standley subsp. pannosum
(Wooton & Standley) S. Stokes
153. E. alatum Torr. in Sitgreaves
153a. var. alatum
Pterogonum alatum (Torr. in Sitgreaves) H.
Gross- E. alatum subsp. typicum S. Stokes
E. alatum var. elatuum Geyer ex Benth. in A.DC.
E. triste S. Watson -E. alatum subsp. triste (S. Wat-
son ) S. Stokes
E. alatum var. brevifolium Gandoger
153b. var. mogollense S. Stokes ex M.E. Jones
E. alatum subsp. mogollense (S. Stokes ex M.E.
Jones) S. Stokes
E. alatum var. macdougalii Gandoger
153c. var. glabriusculum Torr. in Whipple
VII. Subg. Ganysma (S. Watson) E. Greene
154. E. preclarum Rev.
155. E. galioides I.M. Johnston
156. E. arizonicum S. Stokes ex M.E. Jones
157. E. angelense Moran
158. E. inflatum Torr. & Frém in Frém.

Reveal: Checklist of the Eriogonoideae (Polygonaceae)

158a. var. deflatum I.M. Johnston
E. glaucum Small-E. trichopes Torr. in Emory
subsp. glaucum (Small) S. Stokes
E. lagunense M.E. Jones
158b. var. inflatum
E. inflatum subsp. typicum S. Stokes
E. clutei Rydb.
158c. var. fusiforme (Small) Rev.
E. fusiforme Small
159. E. pilosum S. Stokes
160. E. repens (S. Stokes) Rev.
E. trichopes Torr. in Emory subsp. repens S. Stokes
161. E. trichopes Torr. in Emory
161a. var. trichopes
E. trichopodum Torr. ex Benth. in A.DC., nom. su-
perfl.—E. trichopes subsp. yypicum S. Stokes
E. trichopodum Torr. ex Benth. in A.DC. var.
minor Benth. in A.DC.—E. trichopes subsp. mi-
nus (Benth. in A.DC.) S. Stokes
E. clavatum Small—-E. tichopes subsp. clavatum
(Small) S. Stokes
161b. var. hooveri Rev.
162. E. contiguum (Rev.) Rev.
E. inflatum Torr. & Frém. in Frém. var. contiguum
Rev.
163. E. scalare S. Watson
E. irretitum Brandegee
164. E. intricatum Benth. in Hinds
165. E. howellianum Rev.
166. E. glandulosum (Nutt.) Nutt. ex Benth. in A.DC.
Oxytheca glandulosa Nutt.—E. trichopes Torr. in
Emory subsp. glandulosum (Nutt.) S. Stokes
2E. cordatum Torr. & Frém. in Frém.-E. trichopes
Torr. in Emory var. cordatum (Torr. & Frém.
in Frém.) Davidson & Moxley-E. trichopes
Torr. in Emory subsp. cordatum (Torr. & Frém.
in Frém.) S. Stokes
E. glandulosum var. cameum J.T. Howell-E. car-
neum (J.T. Howell) Rev. in Munz
167. E. apiculatum S. Watson
E. apiculatum var. subvirgatum S. Stokes
168. E. parishii S. Watson
169. E. ordii S. Watson
E. tenuissimum Eastw.
170. E. visheri A. Nelson
171. E. aliquantum Rev.
172. E. gordonii Benth. in A.DC.
E. trinervatum Small
173. E. capillare Small
174. E. esmeraldense S. Watson
174a. var. esmeraldense
174b. var. toiyabense J.T. Howell
175. E. concinnum Rev.
176. E. rubicaule Tidestrom
E. trichopes Torr. in Emory var. rubicaule (Tide-
strom) S. Stokes

277

E. lactum S. Stokes
177. E. lemmonii S. Watson
178. E. moranii Rey.
179. E. austrinum (S. Stokes) Rev.
E. deflexum Torr. in Ives subsp. austrinum S.
Stokes
180. E. deflexum Torr. in Ives
180a. var. deflexum
E. deflexum subsp. typicum S. Stokes
E. deflexum f. stenopetale H. Gross
180b. var. nevadense Rev.
180c. var. turbinatum (Small) Rev.
E. turbinatum Small
180d. var. rectum Rev.
180e. var. baratum (Elmer) Rev.
E. baratum Elmer-E. deflecum subsp. baratum
(Elmer) Munz
181. E. rixfordii S. Stokes
E. deflexum Torr. in Ives subsp. rixfordii (S.
Stokes) Munz
182. E. hookeri S. Watson
E. deflexum Torr. in Ives subsp. hookeri (S. Wat-
son) S. Stokes
E. deflexum Torr. in Ives var. gitvum S. Stokes
183. E. brachypodum Torr. & A. Gray
E. deflexum Torr. in Ives var. brachypodum (Torr.
& A. Gray) Munz-E. deflexum subsp. bra-
chypodum (Torr. & A. Gray) S. Stokes
E. parryi A. Gray-E. deflexum Torr. in Ives subsp.
parryt (A. Gray) S. Stokes
184. E. bifurcatum Rey.
185. E. insigne S. Watson
E. deflexum Torr. in Ives var. insigne (S. Watson)
M.E. Jones-E. deflexum subsp. insigne (S. Wat-
son) S. Stokes
E. exaltatum M.E. Jones—E. defleaum Torr. in Ives
subsp. exaltatum (M.E. Jones) S. Stokes
186. E. hoffmannii S. Stokes
186a. var. hoffmannii
186b. var. robustius S. Stokes
E. hoffmannii subsp. robustius (S. Stokes) Munz
187. E. watsonii Torr. & A. Gray
E. deflecum Torr. in Ives subsp. watsonii (Torr. &
A. Gray) S. Stokes—E. deflexur subsp. watsonii
(Torr. & A. Gray) R. Davis
E. cernuum Nutt. var. multipedunculatum S.
Stokes—E. deflexum Torr. in Ives var. mult-
pedunculatum (S. Stokes) C.L. Hitchce.
188. E. scabrellum Rev.
189. E. eremicola J.T. Howell & Rev.
190. E. tenellum Torr.
190a. var. tenellum
E. tenellurn subsp. typicum S. Stokes
E. tenellum var. leptocladon Benth. in A.DC.
190b. var. ramosissimum Benth. in A.DC.
E. tenellum var. caulescens Torr. & A. Gray, nom.
superfl.

278 PHYTOLOG1I1A volume 66(3):266-294 May 1989

190c. var. platyphyllum (Torr. ex Benth. in A.DC.)
Torr.

E. platyphyllum Torr. ex Benth. in A.DC. —E. tenel-
lum subsp. platyphyllum (Torr. ex Benth. in
A.DC.) S. Stokes

191. E. cernuum Nutt.
19la. var. cernuum

E. cernuum subsp. typicum S. Stokes

E. cernuum var. tenue Torr. & A. Gray-E. cer-
nuum subsp. tenue (Torr. & A. Gray) S. Stokes

E. cernuum var. umbraticum Eastw.

191b. var. viminale (S. Stokes) Rev.
E. cernuum subsp. viminale S. Stokes
192. E. rotundifolium Benth. in A.DC.

E. cernuum Nutt. subsp. rotundifolium (Benth. in
A.DC.) S. Stokes —E. rotundifolium var. typicum
Goodman

E. rotundifolium subsp. glaucescens S. Stokes

E. rotundifolium var. angustum Goodman

193. E. nutans Torr. & A. Gray

193a. var. nutans

E. cernuum Nutt. var. purpurascens Torr. & A.
Gray in Beckwith

E. rubiflorum M.E. Jones

E. nutans var. brevipedicellatum S. Stokes

E. deflexum subsp. ultrum S. Stokes

193b. var. glabratum Rev.

194. E. thurberi Torr. in Emory

E. cernuum Nutt. subsp. thurberi (Torr. in Emory)
S. Stokes

E. panduratum S. Watson

E. thurberi var. parishii Gandoger

E. cernuum Nutt. subsp. viscosum S. Stokes

195. E. thomasii Torr.
E. minutiflorum S. Watson
196. E. pusillum Torr. & A. Gray

E. reniforme Torr. & Frém. in Frém. subsp. pusil-
lum (Torr. & A. Gray) S. Stokes

E. comosum (M.E. Jones) M.E. Jones var. play-
anum MLE. Jones—E. reniforme Torr. & Frém.
in Frém. var. playanum (M.E. Jones) S. Stokes

E. reniforme Torr. & Frém. in Frém. var. asari-
folum Gandoger

197. E. reniforme Torr. & Frém. in Frém.

E. reniforme subsp. typicum S. Stokes

E. reniforme var. comosum M.E. Jones—E. com-
osum (M.E. Jones) M.E. Jones

198. E. wetherillii Eastw.
E. sessile S. Stokes ex M.E. Jones
E. filiforme L.O. Willams
199. E. subreniforme S. Watson
E. filicaule S. Stokes
200. E. viscidulum J.T. Howell
201. E. collinum S. Stokes ex M.E. Jones
202. E. salicornioides Gandoger

E. vimineum Douglas ex Benth. var. salicorniotdes

(Gandoger) S. Stokes

E. demissum S. Stokes
E. demissum S. Stokes var. romanum S. Stokes
203. E. abertianum Torr. in Emory
E. abertianum subsp. typicum S. Stokes
E. abertianum var. neomexicanum Gandoger
E. abertianum var. ruberrimum Gandoger
E. cyclosepalum E. Greene-E. abertianum var.
cyclosepalum (E. Greene) Fosb.
E. pinetorum E. Greene-E. abertianum subsp.
pinetorum (E. Greene) S. Stokes
E. lappulaceum E. Greene-E. abertianum subsp.
lappulaceum (E. Greene) S. Stokes—E. aber-
tianum var. lappulaceum (E. Greene) Fosb.
E. abertianum var. villosum Fosb.
E. abertianum var. gillespei Fosb.—E. cyclosepalum
E. Greene var. gillespei (Fosb.) I.M. Johnston
E. abertianum var. bracteatum Fosb.
204. E. pharnaceoides Torr. in Sitgreaves
24a. var. pharnaceoides
E. arizonicum Gandoger, nom. illeg., non M.E.
Jones
204b. var. cervinum Rev.
205. E. spergulinum A. Gray
205a. var. reddingianum (M.E. Jones) J.T. Howell
Oxytheca reddingiana M.E. Jones—E. spergulinum
subsp. reddingianum (M.E. Jones) Munz ex
Rev.
205b. var. spergulinum
Oxytheca spergulina (A. Gray) E. Greene -E. sper-
gulinum var. typicum J.T. Howell
205c. var. pratense (S. Stokes) J.T. Howell
E. pratense S. Stokes
206. E. hirtiflorum A. Gray ex S. Watson
Oxytheca hirtiflora (A. Gray ex S. Watson) E.
Greene
207. E. inerme (S. Watson) Jepson
207a. var. inerme
Oxytheca inermis S. Watson-E. vagans S. Watson,
nom. superfl.—E. inerme var. typicum Goodman
207b. var. hispidulum Goodman
E. inerme subsp. hispidulur (Goodman) Munz
208. E. angulosum Benth.
E. angulosum subsp. typicum S. Stokes
209. E. maculatum A. Heller
E. angulosum Benth. var. maculatum (A. Heller)
Jepson-E. angulosum Benth. subsp. macu-
latum (A. Heller) S. Stokes
. angulosum Benth. var. rectipes Gandoger
. angulosum Benth. var. pauciflorum Gandoger
angulosum Benth. var. flabellauaim Gandoger
angulosum Benth. var. patens Gandoger
thurberi Torr. in Emory var. acutangulum Gan-
doger—E. cernuum Nutt. subsp. acutangulum
(Gandoger) S. Stokes
210. E. viridescens A. Heller
E. angulosum Benth. var. viridescens (A. Heller)
Jepson -E. angulosum Benth. subsp. viridescens
(A. Heller) S. Stokes

mm mmm

Reveal: Checklist of the Eriogonoideae (Polygonaceac) 279

E. bidentatum Jepson-E. angulosum Benth. sub-
sp. bidentatum (Jepson) S. Stokes
211. E. gracillimum S. Watson
E. angulosum Benth. var. gracillimum (S. Watson)
M.E. Jones -E. angulosum Benth. subsp. gracil-
limum (S. Watson ) S. Stokes
E. variabile A. Heller—E. angulosum Benth. var.
variabile (A. Heller) Parish
E. angulosum Benth. var. victorense M.E. Jones—
E. angulosum Benth. subsp. victorense (M.E.
Jones) S. Stokes
212. E. gossypinum Curran
VIII. Subg. Oregonium (S. Watson) E. Greene
213. E. mohavense S. Watson
E. mohavense subsp. typicum S. Stokes
E. delicatulum S. Watson
214. E. ampullaceum J.T. Howell
E. mohavense S. Watson subsp. ampullaceum (J.T.
Howell) S. Stokes
215. E. eastwoodianum J.T. Howell
E. truncatum Torr. & A. Gray var. adsurgens Jep-
son-E. vimineum Douglas ex Benth. subsp. ad-
surgens (Jepson) S. Stokes—E. covilleanum
Eastw. subsp. adsurgens (Jepson) Abrams
216. E. temblorense J.T. Howell
217. E. vestitum J.T. Howell
218. E. truncatum Torr. & A. Gray
219. E. argillosum J.T. Howell
220. E. covilleanum Eastw.
E. vimineum Douglas ex Benth. var. covilleanum
(Eastw.) S. Stokes
221. E. nortonii E. Greene
E. vimineum Douglas ex Benth. subsp. nortonii (E.
Greene) S. Stokes
222. E. luteolum E. Greene
222a. var. luteolum
E. vimineum Douglas ex Benth. var. luteolum (E.
Greene) S. Stokes
222b. var. pedunculatum (S. Stokes) Rev.
E. pedunculatum S. Stokes
222c. var. saltuarium Rey.
222d. var. caninum (E. Greene) Rev.
E. vimineum Douglas ex Benth. var. caninum E.
Greene -E. caninum (E. Greene) Munz
E. vimineum Douglas ex Benth. var. californicum
Gandoger
223. E. vimineum Douglas ex Benth.
E. vimineum subsp. typicum S. Stokes
E. vimineum var. oregonense Gandoger
E. vimineum var. rigescens Gandoger
E. vimineum var. divergens Gandoger
E. shoshoense A. Nelson-E. vimineum var. sho-
shoense (A. Nelson) S. Stokes
224. E. davidsonii E. Greene
E. baileyi S. Watson var. davidsonii (E. Greene)
MLE. Jones-E. molestum S. Watson var. david-
soni (E. Greene) Jepson -E. vimineum Douglas
ex Benth. var. davidsonii (E. Greene) S. Stokes

E. juncinellum Gandoger—E. vimineum Douglas
ex Benth. subsp. jucinellum (Gandoger) S.
Stokes

E. vimineum Douglas ex Benth. var. glabrum S.
Stokes

E. vimineum Douglas ex Benth. var. aviculare S.
Stokes

225. E. molestum S. Watson

E. vimineurn Douglas ex Benth. subsp. molestum

(S. Watson ) S. Stokes
226. E. roseum Durand & Hilg.

E. virgatum Benth. in A.DC. var. roseum (Durand
& Hilg.) Torr. & A. Gray

E. virgatum Benth. in A.DC.-E. vimineum
Douglas ex Benth. subsp. virgatum (Benth. in
A.DC.) S. Stokes

E. virgatum Benth. in A.DC. var. rubidum Jepson
ex Bauer

227. E. gracile Benth. in Hinds
227a. var. gracile

E. vimineum Douglas ex Benth. subsp. gracile
(Benth. in Hinds) S. Stokes

E. verticillatum Nutt.

E. acetoselloides Torr. ex Benth. in A.DC.-E.
gracile var. acetoselloides (Torr. ex Benth. in
A.DC.) Torr. & A. Gray

E. leucocladon Benth.-E. gracile var. leuco-
cladon (Benth.) Torr. & A. Gray-E. roseum
Durand & Hilg. var. leucocladon (Benth.)
Hoover

227b. var. incultum Rev.

228. E. cithariforme S. Watson

228a var. cithariforme

E. vimineum Douglas ex Benth. var. cithariforme
(S. Watson) S. Stokes-E. gracile Benth. in
Hinds var. cithariforme (S. Watson ) Munz

228b. var. agninum (E. Greene) Rev.

E. vimineum Douglas ex Benth. var. agninum (E.
Greene) S. Stokes

E. vimineum Douglas ex Benth. subsp. polygon-
oides S. Stokes -E. gracile Benth. in Hinds var.
polygonoides (S. Watson ) Munz

229. E. baileyi S. Watson

229a. var. baileyi

E. vimineum Douglas ex Benth. subsp. baileyi (S.
Watson) S. Stokes-—E. vimineum Douglas ex
Benth. var. baileyi (S. Watson ) R. Davis

E. gracile Benth. in Hinds var. effusum Torr. & A.
Gray

E. baileyi var. porphyreticum S. Stokes ex M.E.
Jones —E. vimineum Douglas ex Benth. var. por-
phyreticum (S. Stokes ex M.E. Jones) S. Stokes

E. restioides Gandoger-E. vimineum Douglas ex
Benth. var. restioides (Gandoger) S. Stokes

E. vimineum Douglas ex Benth. var. multiradiatum
S. Stokes

280 PHYTOLOGTIA volume 66(3):266-294 May 1989

229b. var. praebens (Gandoger) Rev.

E. praebens Gandoger

E. praebens Gandoger var. divaricatum Gan-
doger—E. baileyi var. divaricatum (Gandoger)
Rev. in Munz

E. leucocladum Gandoger

E. commixtum E. Greene ex Tidestrom—E.
vimineum Douglas ex Benth. var. commixtum
(E. Greene ex Tidestrom) S. Stokes

230. E. elegans E. Greene

E. vimineum Douglas ex Benth. var. elegans (E.
Greene) Jepson-E. baileyi S. Watson subsp.
elegans (E. Greene) Munz

231. E. brachyanthum Cov.

E. baileyi S. Watson var. brachyanthum (Cov.) Jep-
son-E. vimineum Douglas ex Benth. var. bra-
chyanthum (Cov.) S. Stokes

232. E. foliosum S. Watson
233. E. hastatum Wiggins

E. foliosum S. Watson var. hastatum (Wiggins)

Rev. in Munz
234. E. nidularium Cov.

E. vimineum Douglas ex Benth. subsp. nidularium
(Cov.) S. Stokes

E. nidularium var. luciense M.E. Jones

235. E. palmerianum Rev. in Munz

E. plumatella Durand & Hilg. var. palmeri Tort.
& A. Gray-E. baileyi S. Watson var. tomen-
tosum S. Watson, nom. superfl.

236. E. polycladon Benth. in A.DC.

E. vimineum Douglas ex Benth. subsp. polycladon
(Benth. in A.DC.) S. Stokes

E. densum E. Greene-E. virmineurn Douglas ex
Benth. var. densum (E. Greene) S. Stokes

E. polycladon Benth. in A.DC. var. mexicanum
Gandoger

E. polycladon Benth. in A.DC. var. crispum Gan-
doger

237. E. dasyanthemum Torr. & A. Gray

E. vimineum Douglas ex Benth. var. eriocladon
Benth. in A-.DC.

E. dasyanthemum vat. jepsonii E. Greene

238. E. divaricatum Hook.
E. ameghinoi Speg.—Sanmartinia ameghinot
(Speg.) Buchinger
239. E. puberulum S. Watson
E. puberulum var. venosum S. Stokes
240. E. darrovii Kearney

2. Depeckera Rev. & J.T. Howell ~
2A1. D. eurekensis Rev. & J.T. Howell

3. STENOGONUM Nutt.
2A2. S. flexum (M.E. Jones) Rev. & J.T. Howell
Eriogonum flexum M.E. Jones
Eriogonum fleaim M.E. Jones var. ferronis M.E.
Jones

243. S. salsuginosum Nutt.
Eriogonum salsuginosum (Nutt.) Hook.

4. OXYTHECA Nutt.
41. Sect. Oxytheca
244. O. dendroidea Nutt.
244A. subsp. dendroidea
Eriogonum dendroideum (Nutt.) S. Stokes-E.
dendroideum vat. typicum S. Stokes
O. foliosa Nutt.—O. dendroidea var. foliosa (Nutt.)
M.E. Jones
O. dendroidea var. hillmanii S. Stokes ex M.E.
Jones
Eriogonum dendroideum vat. hillmanii S. Stokes
244B. subsp. chilensis (Rémy in Gay) Ertter
Brisegnoa chilensis Rémy in Gay
O. spiculata Miers
O. dendroidea var. tonsiflora 1.M. Johnston
24S. O. watsonii Torr. & A. Gray
Eriogonum cuspidatum S. Stokes
246. O. perfoliata Torr. & A. Gray
Eriogonum perfoliatum (Torr. & A. Gray) S.
Stokes
4II. Sect. Acanthoscyphus (Small) Ertter
247. O. parishii C. Parry
247a. var. parishii
Acanthoscyphus parishit (C. Parry) Small —Erio-
gonum abramsii (McGregor) S. Stokes subsp.
acanthoscyphus S. Stokes
247b. var. cienegensis Ertter
247c. var. goodmaniana Ertter
247d. var. abramsii (McGregor) Munz
Oxytheca abramsii McGregor -Eriogonum abram-
sii (McGregor) S. Stokes—E. abramsii subsp.
typicum S. Stokes-O. parishit subsp. abramsti
(McGregor) Munz
4IIL. Sect. Neoxytheca Ertter
248. O. caryophylloides C. Parry
Eriogonum caryophylloides (C. Parry) S. Stokes
249. O. trilobata A. Gray
Eriogonum trilobatum (A. Gray) S. Stokes
250. O. emarginata H.M. Hall
Eriogonum emarginatum (H.M. Hall) S. Stokes

5. GOODMANIA Rev. & Ertter
251. G. luteola (C. Parry) Rev. & Ertter

Oxytheca luteola C. Parry-Eriogonum spinescens
S. Stokes

6. GILMANIA Cov.
252. G. luteola (Cov.) Cov.
Phyllogonum luteolum Cov.—Eriogonum luteolum
(Cov.) M.E. Jones, non E. Greene

7. NEMACAULIS Nutt.
253. N. denudata Nutt.

Reveal: Checklist of the Eriogonoideae (Polygonaceae) 281

253a. var. denudata

N. nutallii Benth. in A.DC.-Erigonum denud-
atum (Nutt.) Curran, non Nutt.—E. nemacaulis
S. Stokes

N. foliosa Nutt.

253b. var. gracilis Goodman & L. Benson

8. CHORIZANTHE R.Br. ex Benth.
81. Subg. Chorizanthe
254. C. paniculata Benth.
C. ramosissimus Benth.
C. macraet Benth.
C. macraei Benth. var. humilis Benth. in A-.DC.
C. flavescens Philippi
C. rosea Philippi
C. densa Philippi
255. C. glabrescens Benth
C. vaginata var. arida Rémy in Gay
C. tenuis Philippi
256. C. dasyantha Philippi
C. parviflora Philippi
C. intricata Philippi
C. tenuis Philippi, non Philippi
C. fasciculata Philippi
C. humulis Philippi
C. illapelina Philippi
257. C. kingii Philippi

C. pauciflora Philippi
258. C. deserticola Philippi
259. C. virgata Benth.

C. virgata var. tomentosa Benth. in A.DC.
260. C. viridis Philippi
261. C. peduncularis Benth.

C. umbellata Philippi
262. C. vaginata Benth.

C. vaginata var. maritima Rémy in Gay
263. C. frankenoides Rémy in Gay
8II. Subg. Quintaria Rey. & Hardham
264. C. spinosa S. Watson

Erigonella spinosa (S. Watson) Goodman
S8III. Subg. Eriogonella (Goodman) Rev. & Hardham
265. C. membranacea Benth.

Eriogonella membranacea (Benth.) Goodman
8IV. Subg. Amphietes Rev. & Hardham
266. C. stellulata Benth. in A.DC.
267. C. douglasii Benth.

C. nortonii E. Greene
268. C. diffusa Benth. in A.DC.

C. pungens Benth. var. nivea Curran-C. nivea
(Curran) A. Heller—C. diffusa var. ntvea (Cur-
ran) Hoover

C. andersonii C. Parry

269. C. pungens Benth.
269a. var. pungens

C. douglasii Benth. var. albens C. Parry

269b. var. hartwegiana Rev. & Hardham

270. C. angustifolia Nutt.
C. angustifolia var. eastwwoodae Goodman
271. C. cuspidata S. Watson
271a. var. cuspidata
C. pungens Benth. var. cuspidata (S. Watson) C.
Parry
C. cuspidata var. marginata Goodman
271b. var. villosa (Eastw.) Munz
C. villosa Eastw.
272. C. robusta C. Parry
C. pungens Benth. var. robusta (C. Parry) Jepson
C. douglastt Benth. var. hartwegit Benth. in
A.DC.-C. pungens Benth. var. hartwegii (Benth.
in A.DC.) Goodman
273. C. howellii Goodman
274. C. valida S. Watson
275. C. palmeri S. Watson
276. C. biloba Goodman
276a. var. biloba
C. palmer S. Watson var. biloba (Goodman) Munz
276b. var. immemora Rev. & Hardham
277. C. ventricosa Goodman
C. palmeri S. Watson var. venticosa (Goodman)
Munz
278. C. obovata Goodman
C. obovata f. prostrata Goodman
279. C. blakleyi Hardham
280. C. rectispina Goodman
281. C. clevelandii C. Parry
282. C. uniaristata Torr. & A. Gray
283. C. parryi S. Watson
283a. var. parryi
283b. var. fernandina (S. Watson) Jepson
C. fernandina S. Watson
284. C. inequalis S. Stokes
285. C. procumbens Nutt.
C. uncinata Nutt.
C. procumbens var. albiflora Goodman
C. procumbens var. mexicana Goodman
C. jonestana Goodman
C. chaetophora Goodman
286. C. staticoides Benth.
C. nudicaule Nutt.-C. staticoides var. nudicaule
(Nutt.) Jepson
C. discolor Nutt.
C. staticoides f. bracteata Goodman
C. staticoides var. brevispina Goodman
C. staticoides var. elata Goodman
C. staticoides var. latiloba Goodman
C. chrysacantha Goodman -C. staticotdes subsp.
chrysacantha (Goodman) Munz
C. chrysacantha Goodman var. compacta Good-
man
287. C. leptotheca Goodman
288. C. xantii S. Watson
288a. var. xantii

282 PHYTOLOGIA volume 66(3):266-294 May 1989

288b. var. leucotheca Goodman
C. xantii subsp. leucotheca (Goodman) Munz
289. C. breweri S. Watson
290. C. wheeleri S. Watson
C. insularis R. Hoffmann
291. C. turbinata Wiggins
292. C. mutabilis Brandegee
293. C. rosulenta Rev.
294. C. pulchella Brandegee
295. C. flava Brandegee
C. vaseyi Rose
296. C. fimbriata Nutt.
296a. var. fimbriata
296b. var. laciniata (Torr. in Parke) Jepson
C. laciniata Torr. in Parke
297. C. polygonoides Torr. & A. Gray
297a. var. polygonoides
Acanthogonum polygonoides (Torr. & A. Gray)
Goodman
297b. var. longispina Goodman
Acanthogonum polygonoides (Torr. & A. Gray)
Goodman var. longispinum Goodman -C. poly-
gonoides subsp. longispina (Goodman) Munz
298. C. rigida (Torr. in Whipple) Torr. & A. Gray
Acanthogonum rigidum Torr. in Whipple
299. C. orcuttiana C. Parry
300. C. watsonii Torr. & A. Gray
301. C. corrugata (Torr. in Blake) Torr. & A. Gray
Acanthogonum corrugatum Torr. in Blake
302. C. brevicornu Torr. in Emory
302a. var. brevicornu
302b. var. spathulata (Small ex Rydb.) C.L. Hitche.
C. spathulata Small ex Rydb.-C. brevicornu subsp.
spathulata (Small ex Rydb.) Munz
303. C. commissuralis Rémy in Gay
304. C. interposita Goodman

9. MucRONEA Benth.
305. M. californica Benth.
Chorizanthe californica (Benth.) A. Gray
Chorizanthe californica (Benth.) A. Gray var.
suksdorfii Macbride — M. californica var. suksdor-
fii (Macbride) Goodman
306. M. perfoliata (A. Gray) A. Heller
Chonizanthe perfoliata A. Gray
M. perfoliata var. opaca Hoover

10. SYSTENOTHECA Rev. & Hardham
307. S. vortriedei (Brandegee) Rev. & Hardham
Chorizanthe vortriedei Brandegee — Centrostegia
vortriedei (Brandegee) Goodman

11. CENTROSTEGIA A. Gray ex Benth. in A.DC.
308. C. thurberi A. Gray ex Benth. in A.DC.
Chorizanthe thurberi (A. Gray ex Benth. in A.DC.)
S. Watson

Chorizanthe thurbert (A. Gray ex Benth. in A.DC.)
S. Watson var. cryptantha Curran — Chorizanthe
cryptantha (Curran) Goodd.

Chorizanthe thurbert (A. Gray ex Benth. in A.DC.)
S. Watson var. macrotheca J.T. Howell—Centro-
stegia thurbert var. macrothera (J.T. Howell)
Goodman

12. DoDECAHEMA Rev. & Hardham
309. D. leptoceras (A. Gray in Torr. & A. Gray) Rev.
& Hardham
Centrostegia leptoceras A. Gray in Torr. & A.
Gray — Chorizanthe leptoceras (A. Gray in Torr.
& A. Gray) S. Watson

13. Aristocapsa Rev. & Hardham
310. A. insigis (Curran ) Rev. & Hardham
Chorizanthe insignis Curran-Centrostegia insignis
(Curran) A. Heller—Oxytheca insignis (Curran)
Goodman

14. HOLLIsTERIA S. Watson
311. H. lanata S. Watson
Eriogonum lanatum (S. Watson) Roberty & Vau-
tier
Chorizanthe floccosa M.E. Jones

15. LASTARRIAEA Rémy in Gay
312. L. coriacea (Goodman) Hoover
L. chilensis Rémy in Gay subsp. californica H.
Gross -—Chorizanthe lastarriaea C. Parry var.
californica (H. Gross) Goodman — Chorizanthe
coriacea Goodman
313. L. ptilota Rev.
314. L. chilensis Rémy in Gay
Chorizanthe lastarriaea C. Parry, nom. superfl.
L. stricta Philippi ex C. Parry
L. linearis Philippi ex C. Parry

16. Harrorpia E. Greene & C. Parry in C. Parry
315. H. macroptera (Benth. in Hinds) E. Greene &
C. Parry in C. Parry
315a. var. galioides (E. Greene) Rev.
H. galioides E. Greene
315b. var. fruticosa (E. Greene) Rev.
H. fruticosa E. Greene
315c. var. macroptera
Pterostegia macroptera Benth. in Hinds

17. Prerostecia Fischer & C. Meyer
316. P. drymarioides Fischer & C. Meyer
P. diphylla Nutt.
P. diphylla Nutt. var. biloba Nutt.
P. microphylla Nutt.

Reveal: Checklist of the Eriogonoideae (Polygonaccae)

INDEX TO NAMES

Acanthogonum
corrugatum, 301
polygonoides, 297a

longispinum, 297b
rigidum, 298

Acanthoscyphus
parishii, 247a

Aristocapsa, 13
insigis, 310

Brisegnoa
chilensis, 244B

Centrostegia, 11
insignis, 310
leptoceras, 309
thurberi, 308

macrothera, 308
vortriedei, 307

Chorizanthe, 8
Amphietes, 8IV
andersonii, 268
angustifolia, 270

eastwoodae, 270
biloba, 276
biloba, 276a
immermora, 276b
blakleyi, 279
brevicornu, 302
brevicornu, 302a
spathulata, 302b
breweri, 289
californica, 305
suksdorfii, 305
chaetophora, 285
Chorizanthe, 81
chrysacantha, 286
compacta, 286
clevelandii, 281
commissuralis, 303
coriacea, 312
corrugata, 301
cryptantha, 308
cuspidata, 271
cuspidata, 271a
marginata, 27la
villosa, 271b
dasyantha, 256
densa, 254
deserticola, 258
diffusa, 268
nivea, 268
discolor, 286
douglasii, 257
albens, 269a
hartwegii, 272

Eriogonella, 8III
fasciculata, 256
fernandina, 283b
fimbriata, 296
fimbriata, 296a
laciniata, 296b
flava, 295
flavescens, 254
floccosa, 311
frankenoides, 263
glabrescens, 25S
howelli, 273
humulis, 256
illapelina, 256
inequalis, 284
insignis, 310
insularis, 290
interposita, 304
intricata, 256
jonesiana, 285
kingii, 257
laciniata, 296b
lastarriaea, 314
californica, 312
leptoceras, 309
leptotheca, 287
macraei, 254
humilis, 254
membranacea, 265
mutabilis, 292
nivea, 268
nortonii, 267
nudicaule, 286
obovata, 278
prostrata, 278
orcuttiana, 299
palmeri, 275
biloba, 276a
ventricosa, 276b
paniculata, 254
parryi, 283
fernandina, 283b
parryi, 2834
parviflora, 256
pauciflora, 257
pedunculanis, 261
perfoliata, 306
polygonoides, 297
longispina, 297b
polygonoides, 297a
procumbens, 285
albiflora, 285
mexicana, 285
pulchella, 294
pungens, 269
cuspidata, 27la

hartwegiana, 269b
hartwegii, 272
nivea, 268
pungens, 269a
robusta, 272
Quintana, 8II
ramosissimus,254
rectispina, 280
rigida, 298
robusta, 272
rosea, 254
rosulenta, 293
spathulata, 302b
spinosa, 264
Staticoides, 286
bracteata, 286
brevispina, 286
chrysacantha, 286
elata, 286
latiloba, 286
nudicaule, 286
stellulata, 266
tenuis, 255, 256
thurberi, 308
cryptantha, 308
macrotheca, 308
turbinata, 291
umbellata, 261
uncinata, 285
uniaristata, 282
vaginata, 262
arida, 255
maritima, 262
valida, 274
vaseyi, 295
ventricosa, 277
villosa, 271b
virgata, 259
tomentosa, 259
viridis, 260
vortriedei, 307
watsonii, 300
wheeleri, 290
xantii, 288
leucotheca, 288b
xantii, 288a
Dedeckera, 2
eurekensis, 241
Dodecahema, 12
leptoceras, 309
Eriogonella
spinosa, 264
membranacea, 265
Eriogonum, 1
abertianum, 203
bracteatum, 203

283

284

cyclosepalum, 203
gillespei, 203
lappulaceum, 203
neomexicanum, 203
pinetorum, 203
ruberrimum, 203
typicum, 203
villosum, 203
abramsii, 247d
acanthoscyphus, 247a
typicum, 247d
acaule, 48
longilobum, 87a
shockleyi, 87a
acetoselloides, 227a
affine, 96m
ainliei, 17b
alatum, 153
alatum, 153a
brevifolium, 153a
elatum, 153a
glabriusculum, 153c
macdougalii, 153b
mogollense, 153b
triste, 153a
typicum, 153a
album, 103
aliquantum, 171
allenti, 137
alpinum, 142
ameghinoi, 238
ammophilum, 5
ampullaceum, 214
andinum, 127
androsaceum, 135
piperi, 134d
anemophilum, 67
cusickii, 61
angelense, 157
angulosum, 208
bidentatum, 210
flabellatum, 209
gracillimum, 211
maculatum, 209
patens, 209
pauciflorum, 209
rectipes, 209
typicum, 208
variabile, 211
victorense, 211
viridescens, 210
angustifolium, 112b
annuum, 106
chihuahuaense, 106
cymosum, 106
hitchcockii, 106

PHYTOLOG1IA volume 66(3):266-294 May 1989

pauciflorum, 106
roseum, 106
typicum, 106
anserinum, 102d
apachense, 34
apiculatum, 167
subvirgatum, 167
apricum, 36
apricum, 36a
prostratum, 36b
arachnoideum, 97
arborescens, 26
arcuatum, 132a
aretioides, 86
argillosum, 219
argophyllum, 77
aridum, 111]j
arizonicum, 156, 204a
atrorubens, 143
atrorubens, 143a
auritulum, 143d
intonsum, 143c
nemorosum, 143¢
pseudociliatum, 143b
rupestre, 143f
aureum, 13g
ambiguum, lh
glutinosum, 13g
auriculatum, 96i
austrinum, 179
azaleastrum, 1llc
baileyi, 229
baileyi, 229a
brachyanthum, 231
davidsonii, 224
divaricatum, 229b
elegans, 230
porphyreticum, 229a
praebens, 229b
tomentosum, 235
baker, 132a
bannockense, 37e
baratum, 180e
batemanii, 52
eremicum, 51
ostlundii, 50
beatleyae, 66
bellum, 102a
bicolor, 8
bidentatum, 210
bifurcatum, 184
biumbellatum, 111x
blissianum, 26
bloomeri, 104
brachyanthum, 231
brachypodum, 183

brandegei, 45
breedlovei, 68
breedlovei, 68a
shevockii, 68b
brevicaule, 37
aureum, 37a
brevicaule, 37a
campanulatum, 37a
canum, 38
cottamii, 37d
desertorum, 57
ephedroides, 40
grangerense, 37a
laxifolium, 37e
leptothecum, 37a
loganum, 44
micranthum, 37b
nanum, 37e
orendense, 37b
promiscuum, 37e
pumilum, 37e
typicum, 37a
viridulum, 39
wasatchense, 37c
butterworthianum, 91
caespitosum, 127
acaule, 48
alyssoides, 127
douglasii, 125a
kelloggii, 123
ramosum, 112b
sublineare, 125b
typicum, 127
campanulatum, 37a
aureum, 37a
grangerense, 37a
leptothecum, 37a
orendense, 37b
typicum, 37a
caninum, 222d
capillare, 173
capistratum, 74
capistratum, 74a
muhlickii, 74b
welshii, 74c
capitatum, 96m
carneum, 166
caryophylloides, 248
cernuum, 191
acutangulum, 209
cernuum, 191la
multipedunculatum, 187
purpurascens, 193a
rotundifolium, 192
tenue, 19la
thurberi, 194

Reveal: Checklist of the Eriogonoideae (Polygonaccac)

typicum, 19la
umbraticum, 19la
viminale, 191b
viscosum, 194
chloranthum, 132b
chrysocephalum, 37e
alpestre, S4b
bannockense, 37e
cusickii, 61
desertorum, 57
loganum, 44
mancum, 75
typicum, 37e
chrysops, 70
ciliatum, 148
foliosum, 149
cinereum, 20
cithariforme, 228
agninum, 228b
cithariforme, 228a
Clastomyelon 1III
clavatum, 16la
clavellatum, 7
clivosum, 144
clutei, 1S8b
cognatum, 111bb
collinum, 201
coloradense, 81
commixtum, 229b
comosum, 197
playanum, 196
compositum, 113
compositum, 113a
citrinum, 113a
lancifolium, 113c
leianthum, 113b
pilicaule, 113a
simplex, 113b
typicum, 113a
concinnum, 175
confertiflorum, la
Stansburyi, 37a
congdonii, 117
contiguum, 162
contortum, 46
cordatum, 166
coriaceum, 109a
correllii, 136
corymbosum, 13
albogilvum, 37a
ambiguum, 1h
aureum, 13g
corymbosum, 13a
cronquistii, 53
davidsei, 13c
divaricatum, 13a

erectum, 13b
glutinosum, 13g
humivagans, 17a
matthewsae, 18d
orbiculatum, 13e
revealianum, 13d
smithit, 11
velutinum, 13f
coryphaeum, 140a
covilleanum, 220
adsurgens, 215
covillei, 111h
crassifolium, 134a
tectum, 134a
crispum, 13g
crocatum, 105
croceum, 111s
cronquistii, 53
crosbyae, 64
cupreum, llla
cupulatum, 128
curvatum, 89h
cusickii, 61, 102a
californicum, 102a
cuspidatum, 245
cyclosepalum, 203
gillespei, 203
cymosum, 106
darrovii, 240
dasyanthemum, 237
jepsonii, 237
davidsonii, 224
davisianum, 10la
decumbens, 103
deductum, 96c
deflexum, 180
austrinum, 179
baratum, 180e
brachypodum, 183
deflexum, 180a
exaltatum, 185
gilvum, 182
hooken, 182
insigne, 185
multipedunculatum, 187
nevadense, 180b
parryi, 183
rectum, 180d
rixfordii, 181
stenopetale, 180a
turbinatum, 180c
typicum, 180a
ultrum, 193a
watsonii, 187
delicatulum, 213
demissum, 202

romanum, 202
dendroideum, 244A
hillmanii, 244A
typicum, 244A

densum, 236
denudatum, 88a, 253a
depauperatum, 82a
depressum, 10th
deserticola, 31
desertorum, 57
dichotomum, 103
humile, 103
diclinum, 130
dioecium, 82a
divaricatum, 238
divergens, 13a
douglasii, 125
douglasii, 125a
ramosum, 112b
sublineare, 125b
tenue, 125b
twisselmannii, 126
duchesnense, 37a
dudleyanum, 4
dumosum, 111n
eastwoodae, 3a
eastwoodianum, 215
effusum, 2
ainliei, 17b
contortum, 46
corymbosum, 13a
divaricatum, 13a
durum, 14
effusum, 2a
fendlenanum, 17b
foliosum, 1b
helichrysoides, 2b
leptocladon, 3c
leptophyllum, 6
limbatum, Ic
nelsonii, 1b
nudicaule, 17c
orbiculatum, 13e
pallidum, 3a
rosmarnioides, 2b
salicinum, 17a
salinum, 13b
shandsii, 3c
simpsonii, 1b
typicum, 2a
elatum, 99
elatum, 99a
enanthum, 99a
glabrescens, 138
incurvum, 99b
limonifolium, 99a

285

286

villosum, 99b
elegans, 230
ellipticum, 111s
megacephalum, 124a
elongatum, 88
areorivum, 88b
elongatum, 88a
vollmen, 88c
emarginatum, 250
encelioides, 24
ephedroides, 40
eremicola, 189
eremicum, 51
ericifolium, 10
ericifolium, 10b
pulchrum, 10a
thornei, 10c
Eriogonum, 11V
esmeraldense, 174
esmeraldense, 174a
toiyabense, 174b
Eucycla, 11
exaltatum, 185
exilifolium, 80
eximium, 101c
fasciculatum, 19
aspalathoides, 19d
emphereium, 19c
ericifolium, 10b
fasciculatum, 19d
flavovirde, 19b
foliolosum, 19e
maritimum, 19d
obtusifolium, 19e
oleifolium, 19d
polifolium, 19a
revolutum, 19a
typicum, 19d
fasciculifolium, 124b
fastigiatum, 30
fendlerianum, 17b
ferrissii, 111x
filicaule, 199
filiforme, 198
fimbriatum, 145
flavissimum, 102d
flavum, 134
androsaceum, 135
aquilinum, 134b
caudiciferum, 134a
chloranthum, 132b
crassifolium, 134a
flavum, 134a
foliatum, 132a
incisum, 134a
linguifolium, 134a

PHYTOLOGIA volume 66(3):266-294 May 1989

multicum, 134a
piperi, 134d
polyphyllum, 134c
sericeum, 134a
tectum, 134a
typicum, 134a
xanthum, 132b
flexum, 242
ferronis, 242
floridanum, 109c
foliosum, 232
hastatum, 233
formosum, 22c
friscanum, 1b
fruticosum, 13g
ambiguum, lh
glutinosum, 13g
fruticulosum, 124b
fulvum, 102a
fusiforme, 158c
galioides, 155
Ganysma 1VII
geniculatum, 33b, 124a
giganteum, 22
compactum, 22b
formosum, 22c
giganteum, 22a
gilmanii, 100
glaberrimum, 111dd
aureum, 11lc
glandulosum, 166
carneum , 166
glaucum, 158a
gnaphaloides, 82b
gordonii, 172
gossypinum, 212
gracile, 227
acetoselloides, 227a
cithariforme, 228a
effusum, 229a
gracile, 227a
incultum, 227b
leucocladon, 227a
polygonoides, 228b
gracilipes, 78
gracillimum, 211
gramineum, 96g
grande, 98
dunklei, 98d
grande, 98a
rubescens, 98d
testudinum, 98b
timorum, 98c
grangerense, 37a
grayi, 37e
greggii, 149

gypsophilum, 49
gyrophyllum, 112a
halimioides, 124c
harfordii, 96m
harperi, 109b
hastatum, 233
hausknechtii, 111i
havardii, 56
heermannii, 33
argense, 33g
clokeyi, 33c
floccosum, 33¢
heermannii, 33b
humilius, 33a
occidentale, 33d
subracemosum, 33f
sulcatum, 33h
typicum, 33b
helianthemifolium, 89a
helichrysoides, 2b
hemipterum, 150
griseum, 150b
hemipterum, 150a
henricksonii, 146
heracleoides, 112
angustifolium, 112b
heracleoides, 112a
leucophaeum, 112c
micranthum, 112b
minus, 112a
multiceps, 112a
rydbergii, 112a
simplex, 113b
subalpinum, 111r
typicum, 112a
utahense, 112a
virde, 111f
hieracifolium, 152
atropurpureum 150a
hemipterum, 150a
hirtellum, 139
hirtiflorum, 206
hitchcockii, 106
hoffmannii, 186
hoffmannii, 186a
robustius, 186b
holmgrenii, 79
hookeri, 182
howellianum, 165
howellii, 33g
argense, 33g
subracemosum, 33f
humivagans, 17a
hylophilum, 15
idahoense, li
incanum, 129

Reveal: Checklist of the Eriogonoideae (Polygonaceae)

indictum, 96)
inerme, 207
hispidulum, 207b
inerme, 207a
typicum, 207a
inflatum, 158
contiguum, 162
deflatum, 158a
fusiforme, 158c
inflatum, 158b
typicum, 158b
insigne, 185
intermontanum, 17e
intrafractum, 108
intricatum, la, 164
irretitum, 163
jamesii, 132
arcuatum, 132a
bakeri, 132a
flavescens, 132a
jamesii, 132e
neomexicanum, 132e
rupicola, 132d
simplex, 132f
typicum, 132e
undulatum, 132g
wootonii, 132c
xantum, 132b
johnstonii, 113a
jonesii, 16
junceum, 89h
juncinellum, 224
kearneyi, 4
monoense, 4
kelloggii, 123
kennedyi, 90
alpigeum, 90c
austromontanum, 90b
gracilipes, 78
kennedyi, 90a
olanchense, 89i
Pinicola, 90e
pinorum, 89h
purpursii, 90d
typicum, 90a
kingii, 76
laxifolium, 37e
lachnogynum, 55
tetraneunris, 55
typicum, 55
lachnostegium, 103
laetum, 176
lagopus, 38
lagunense, 158a
lanatum, 311
lancifolium, 14

lanosum, 16
lappulaceum, 203
latens, 138
lateriflorum, 134a
lateum, llla
latifolium, 97
affine, 96m
alternans, 96i
auriculatum, 96i
decurrens, 961
grande, 98a
harfordii, 96m
indictum, 96j
nudum, 96a
parvulum, 96a
pauciflorum, 96¢
rubescens, 98d
saxicola, 96g
scapigerum, 96d
sulphureum, 96m
typicum, 97
westonii, 96g
laxifolium, 37e
lemmonii, 177
leptocladon, 3
leptocladon, 3c
papiliunculi, 3b
ramossimum, 3a
leptophyllum, 6
leucocladon, 227a
leucocladum, 227b
leucophyilum, 56
pannosum, 152
typicum, 56
lewisii, 58
libertini, 116
lindheimerianum, 106
lobbii, 141
lobbii, 141a
minus, 14la
robustum, 141b
loganum, 44
lonchophyilum, 17
fendlerianum, 17b
intermontanum, 17e
lonchophyllum, 17a
nudicaule, 17c
saurinum, 17d
longifolium, 109
caput-felis, 109a
diffusum, 109a
floridanum, 109c
gnaphalifolium, 109c
harperi, 109b
lindheimen, 109a
longidens, 109c

longifolium, 109a
plantagineum, 109a
typicum, 109a
longilobum, 87a
longulum, 96a
luteolum, 222, 252
caninum, 222d
luteolum, 222a
pedunculatum, 222b
saltuarium, 222c
macdougalii, 1b
maculatum, 209
mancum, 75
marginale, 1lic
manrifolium, 128
apertum, 128
incanum, 129
mearnsii, 10b
pulchrum, 10a
medium, 37e
meledonum, 73
mensicola, 92b
Micrantha, 1II
micranthum, 37b
microthecum, 1
alpinum, 1g
ambiguum, 1h
aureum, 13g
bicolor, 8
confertiflorum, la
corymbosoides, 1d
crispum, 13g
effusum, 2a
ericifolium, 10b
expansum, lh
fendlerianum, 17b
foliosum, 1b
friscanum, 1b
idahoense, li
intermedium, 1b
johnstonii, le
lapidicola, 1f
laxiflorum, la
leptocladon, 3c
leptophyllum, 6
macdougalii, 1b
mearnsii, 10b
microthecum, 1i
panamintense, Ic
pulchrum, 10a
rigidum, 1b
simpsonii, 1b
spathulare, la
typicum, li
minimum, 122
minus, 111m

287

288

PHYTOLOGIA volume 66(3):266-294 May 1989

minutiflorum, 195 grande, 98a flavissimum, 102d

modocense, 111n indictum, 96j macropodum, 101g

mohavense, 213 murinum, 96h multiscapum, 101la
ampullaceum, 214 nudum, 96a nevadense, 101b

typicum, 213 oblongifolium, 96m nivale, 101i
molestum, 225 paralinum, 96b ochroleucum, 101g

davidsonii, 224 pauciflorum, 96e orthocaulon, 101a
molle, 25 perturbum, 96e ovalifolium, 101b

montanum, 111i
monticola, 138
moranii, 178
mortonianum, 12
multiceps, 82a

canum, 38 westonii, 96g vineum, 10le
coloradense, 81 nummulare, 4 williamsae, 101d
typicum, 82a ammophilum, 5 ovatum, 118
multiflorum, 107 nutans, 193 pallidum, 3a
myrianthum, 2a brevipedicellatum, 193a palmeri, 35
nanum, 37e glabratum, 193b palmerianum, 235
natum, 43 nutans, 193a panamintense, 92
nealleyi, 151 nuttallii, 101a mensicola, 92b
neglectum, 111c oblanceolatum, 96a panamintense, 92a
nelsonii, 1b oblongifolium, 97 panduratum, 194
nemacaulis, 253a minus, 10la panguicense, 54
nervulosum, 119 obtusum, 94 alpestre, 54b
nevadense, 59a ochrocephalum, 59 panguicense, 54a
nidularium, 234 agnellum, 65 pannosum, 152
luciense, 234 alexandreae, 59d parishii, 168
nivale, 101i angustum, 37e parryi, 183
niveum, 103 breedlovei, 68a parviflorum, 82a
candelabrum, 103 calcareum, S9b parvifolium, 21
decumbens, 103 chrysops, 70 crassifolium, 21
dichotomum, 103 gracilipes, 78 lucidum, 21
greenei, 102b ochrocephalum, 59a paynei, 21
suksdorfii, 103 sceptrum, 59c typicum, 21
typicum, 103 ochroleucum, 101g pauciflorum, 82
nodosum, 89g decalvans, 101g canum, 38
jaegeri, 35 macropodum, 101g gnaphaloides, 82b
kearneyi, 4 Oligogonum, 1V panguicense, 54a
monoense, 4 orcuttianum, 28 pauciflorum, 82a
nortonii, 221 ordii, 169 pedunculatum, 222b
novonudum, 60 Oregonium, 1VIII pelinophilum, 47
nudicaule, 17c orendense, 37b pendulum, 32
angustum, 37e orthocaulon, 101a peninsulare, 28
garrettii, 37a orthocladon, 94 perfoliatum, 246
ochroflorum, 41 ostlundii, 50 pharnaceoides, 204
paraleyense, 37a ovalifolium, 101 cervinum, 204b
pumilum, 37e anserinum, 102d pharnaceoides, 204a
scoparium, 17a bellum, 102a pilicaule, 113a
tristichum, 17a caelestinum, 101j pilosum, 159
typicum, 17c celsum, 10la pinetorum, 203
nudum, 96 cerastoides, 101la piperi, 134d

auriculatum, 961
decurrens, 961
deductum, 96c
gramineum, 96g

pubiflorum, 96f
regirivum, 96k
saxicola, 96g
scapigerum, 96d
sulphureum, 96m

cyclophyllum, 101a
deltoideum, 101b
depessum, 101h
eximium, 101c

pansum, 101f
proliferum, 102a
purpureum, 10la
typicum, 101b
utahense, 10la

longiflorum, 134d
ochrocephalum, 134d
platyphyllum, 190c

plumatella, 35

Reveal: Checklist of the Eriogonoideae (Polygonaceac)

jaegeri, 35
palmeri, 235
typicum, 35
polifolium, 19a
polyanthum, 111n
bahiiforme, 111z
polycladon, 236
crispum, 236
mexicanum, 236
polyphyllum, 134c
polypodum, 131
pondii, 27
gentryi, 27
porter, 111d
praebens, 229b
divaricatum, 229b
pratense, 20Sc
prattenianum, 120
avium, 120b
prattenianum, 120a
preclarum, 154
pringlei, 89f
prociduum, 63
proliferum, 102a
anserinum, 102d
Pterogonum, 1VI
puberulum, 239
venosum, 239
pulchrum, 10a
purpureum, 10la
tenius, 10la
purpursii, 90d
pusillum, 196
pyroliifolium, 140
bellingeranum, 140a
coryphaeum, 140a
pyroliifolium, 140b
typicum, 140b
racemosum, 94
coccineum, 95b
cordifolium, 94
desertorum, 92a
obtusum, 94
orthocladon, 94
Sagittatum, 94
typicum, 94
zionis, 95a
ramosissimum, 3a
reclinatum, 111f
reliquum, 92a
reniforme, 197
asarifolum, 196
comosum, 197
playanum, 196
pusillum, 196
typicum, 197

repens, 160

revealianum, 13d
revolutum, 19a
rhodanthum, 101i
riplelyi, 9
rixfordii, 181
robustum, 141b
roseiflorum, 101a
rosense, 65
roseum, 226
leucocladon, 227a
rosmarinifolium, 19d
foliolosum, 19e
rosulatum, 129
rotundifolium, 192
angustum, 192
glaucescens, 192
typicum, 192
rubescens, 98d
tubicaule, 176
rubidum, 101h
frigidum, 101h
rubiflorum, 193a
Tupestre, 143f
rupinum, 93
rydbergii, 111b
sabulosum, 37a
salicinum, 17a
salicornioides, 202
salinum, 13b
salsuginosum, 243
sarothriforme, 17a
saurinum, 17d
saxatile, 104
crocatum, 105
multicaule, 104
stokesae, 104
saxicola, 96g
scabrellum, 188
scalare, 163
scapigerum, 96d
scoparium, 17a
scopulorum, 69
sericeum, 134a
sericoleucum, 127
sessile, 198
shockleyi, 87
shockleyi candidum, 87a
longilobum, 87a
packardae, 87b
shockleyi, 87a
typicum, 87a
shoshoense, 223
simpsonii, 1b
floccoso-lanatum, 106

siskiyouense, 114
smithii, 11
soredium, 85
spathuiforme, 50
spathulare, la
spathulatum, 41
brandegei, 45
natum, 43
panguicense, 54a
spathuiforme, 50
typicum, 41
speciosum, 111q
spergulinum, 205
pratense, 20Sc
reddingianum, 205Sa
spergulinum, 205b
typicum, 20Sb
sphaerocephalum, 124
brevifolium, 121
fasciculifolium, 124b
geniculatum, 124a
halimioides, 124c
megacephalum, 124a
minimum, 122
sericoleucum, 127
sphaerocephalum, 124a
tenue, 125b
typicum, 124a
spinescens, 251
stellatum, 111s
bahiiforme, 111z
stokesae, 104
Strictum, 102
anserinum, 102d
argenteum, 102a
bellum, 102a
cusickii, 102a
flavissimum, 102d
glabrum, 102c
greenei, 102b
lachnostegium, 103
proliferum, 102A, 102a
Strictum, 102B
tenue, 12Sb
typicum, 102B
subalpinum, 111r
arachnoideum, 111j
cladophorum, 111b
stenophyllum, 111r
subnivale, 111r
vulcanicum, 111r
subreniforme, 199
suffruticosum, 29
sulcatum, 33h
argense, 33g
sulphureum, 96m

289

290

taxifolium, 89c
temblorense, 216
tenellum, 10la, 190
caulescens, 190b
cottamii, 37d
erianthum, 1h
grandiflorum, la
leptocladon, 190a
ostlundii, 50
platyphyllum, 190c
ramosissimum, 190b
sessiliflorum, la
tenellum, 190a
typicum, 190a
tenue, 12Sb
tenuissimum, 169
ternatum, 115
congdonii, 117
tetraneuris, 55
texanum, 109a
thomasii, 195
thompsonae, 18
albiflorum, 18b
atwoodii, 18c
matthewsae, 18d
thompsonae, 18a
thurberi, 194
acutangulum, 209
parishii, 194
thymoides, 122
congestum, 122
pallens, 122
typicum, 122
tiehmii, 62
tolmieanum, 113b
tomentosum, 110
torreyanum, 111cc
trachygonum, 89b
dentatum, 89d
glomerulum, 89a
membranaceum, 89c
pringlei, 89f
subscaposum, 89h
taxifolium, 89e
typicum, 89b
wrightii, 89a
trichopes, 161
clavatum, 16la
cordatum, 166
glandulosum, 166
glaucum, 158a
hooveri, 161b
minus, 16la
repens, 160
rubicaule, 176
trichopes, 161a

PHYTOLOG1A volume 66(3):266-294 May 1989

typicum, 1l6la
trichopodum, 16la
minor, 1l6la
trichotomum, 111z
trilobatum, 249
trinervatum, 172
tripodum, 121
triste, 153a
tristichum, 17a
truncatum, 218
adsurgens, 215
tumulosum, 84
turbinatum, 180c
turnen, 133
twisselmannii, 126
umbellatum, 111
argus, 1llv
aridum, 111j
aureum, 1llc
bahiiforme, 111z
californicum, 111f
chlorothamnus, 11ly
chrysanthum, 111s
cladophorum, 111b
cognatum, 111bb
covillei, 111h
crandalii, 1lla
croceum, 111s
desereticum, 111k
devestivum, 111t
dichrocephalum, 111j
dumosum, 111n
ferrissii, 111x
furcosum, 11lu
glaberrimum, 111dd
glabratum, 111c
goodmanii, 11lo
hausknechtii, 1111
humistratum, 111p
hypoleium, 11le
smallianum, 111z
munzii, 1llw
intectum, 11llc
juniporinum, 11laa
majus, 111r
minus, 111m
modocense, 111n
monocephalum, 113b
nevadense, 111f
polyanthum, 111n
polypodum, 131
porteri, 111d
serratum, 120a
speciosum, 111q
stellatum, 111s
subalpinum, 111r

subaridum, 111x
ternatum, 115
tolmieanum, 113b
torreyanum, 111cc
typicum, 1lla
umbellatum, 1lla
vernum, 111g
versicolor, 1111
umbelliferum, 111c
undulatum, 132g
ursinum, 118
confine, 115
congdonii, 117
covillei, 111h
nervulosum, 119
rosulatum, 129
typicum, 118
venosum, 131
vagans, 207a
variabile, 211
vegetius, 132a
verrucosum, 72
verticillatum, 227a
vespinum, 109a
vestitum, 217
villiflorum, 83
candidum, 87a
tumulosum, 84
typicum, 83
vimineum, 223
adsurgens, 215
agninum, 228b
aviculare, 224
baileyi, 229a
brachyanthum, 231
californicum, 222d
caninum, 222d
cithariforme, 228a
commixtum, 229b
covilleanum, 220
davidsonii, 224
densum, 236
divergens, 223
elegans, 230
eriocladon, 237
glabrum, 224
gracile, 227a
jucinellum, 224
luteolum, 22a
molestum, 225
multiradiatum, 229a
nidularium, 234
nortonii, 221
oregonense, 223
polycladon, 236
polygonoides, 228b

Reveal: Checklist of the Eriogonoideae (Polygonaceae)

porphyreticum, 229a
restioides, 229a
rigescens, 223
salicornioides, 202
shoshoense, 223
typicum, 223
virgatum, 226
vineum, 10le
virgatum, 226
roseum, 226
rubidum, 226
viridescens, 210
viridulum, 39
viscanum, rae
viscidulum, 200
vishen, 170
vollmeri, 88c
wasatchense, 37c
watsonii, 187
wethernillii, 198
wrightii, 89
curvatum, 89h
dentatum, 89d
floccosum, 89a
glomerulum, 89a
helianthemifolium, 89a
membranaceum, 89c
nodosum, 89g
olanchense, 89i
oresbium, 89j
pringlei, 89f
subscaposum, 89h
taxifolium, 89e
trachygonum, 89b
typicum, 89a
wrightii, 89a
xanthum, 132b
zapatoense, 23
zionis, 95
coccineum, 95b
zionis, 9Sa
Eucycla
ovalifolia, 101b
purpurea, 10la
Gilmania, 6
luteola, 252
Goodmania, 5
luteola, 251
Harfordia, 16
fruticosa, 315b
galioides, 31Sa
macroptera, 315
fruticosa, 315b
galioides, 315a
macroptera, 315c
Hollisteria, 14

lanata, 311
Lastarnaea, 15
chilensis, 314
californica, 312
coriacea, 312
linearis, 314
ptilota, 313
Stricta, 314
Mucronea, 9
californica, 305
suksdorfii, 305
perfoliata, 306
opaca, 306
Nemacaulis, 7
denudata, 253
denudata, 253a
gracilis, 253b
foliosa, 253a
nuttallii, 253a
Oxytheca, 4
abramsii, 247d

Acanthoscyphus, 4II
caryophylloides, 248

dendroidea, 244
chilensis, 244B

dendroidea, 244A

foliosa, 244A
hillmanii, 244A

tonsiflora, 244B

emarginata, 250
foliosa, 244A
glandulosa, 166
hirtiflora, 206
inermis, 207a
insignis, 310
luteola, 251
Oxytheca, 41
Neoxytheca, 4III
parishii, 247
abramsii, 247d

cienegensis, 247b
goodmaniana, 247c

parishii, 247a
perfoliata, 246

reddingiana, p 20Sa

spergulina, 20Sb
spiculata, 244B
trilobata, 249
watsonii, 245
Phyllogonum
luteolum, 252
Pterogonum
alatum, 153a
atrorubens, 143a
hieracifolium, 152
Pterostegia, 17

291

diphyila, 316
biloba, 316
drymanioides, 316
macroptera, 31Sc
microphylla, 316
Sanmartinia
ameghinoi, 238
Stenogonum, 3
flexum, 242
salsuginosum, 243
Systenotheca, 10
vortriedei, 307

297 PHYTOLO GIA volume 66(3):266-294 May 1989

IMPORTANT SYSTEMATIC LITERATURE ON THE ERIOGONOIDEAE

Abrams, L. 1944. Illustrated flora of the Pacific States. Vol. 2. Stanford: Stanford Univ. Press.
Barneby, R. C. 1949. Eriogonum villiflorum and its near relatives in the Great Basin. Leafl. W. Bot. 5: 151-154.
Bentham, G. 1836. On the Eriogoneae, a tribe of the order Polygonaceae. Trans. Linn. Soc. London 17: 401420.

. 1856. “Eriogoneae,” pp. 5-28. In: A. de Candolle (edit.), Prodromus systematis naturalist regni
vegetabilis. Vol. 14. Paris: Victor Masson.

Brandegee, M. K. Notes on Eriogoneae. Erythea S: 79-81.
Coville, F. V. 1936. Gilmania, a new name for Phyllogonum, a very rare genus of plants from Death Valley,
California, apparently in process of extinction. J. Wash. Acad. Sci. 26: 209-213.
& C. V. Morton. 1936. Eriogonum intrafractum, a new species and new subgenus from Death Valley,
California. J. Wash. Acad. Sci. 26: 303-306.
Curran, M. K. 1885. Classification of the Eriogoneae as affected by some connecting forms. Bull. Calif. Acad. Sci.
1: 272-275.
Dammer, U. 1892. Polygonaceae. Nat. Pflanzenfam. 3(1a): 1-36.
Dumortier, B. C. J. 1829. Analyse des familles des plantes. Tournay: J. Casterman.
Ertter, B. J. 1980. A revision of the genus Oxytheca Nutt. (Polygonaceae). Brittonia 32: 70-102.
. Notes on Goodmania and Oxytheca (Polygonaceae: Eriogonoideae). Brittonia 33: 37-38.
Fosberg, F. R. 1938. Eriogonum abertianum and its varieties. Madrofio 4: 189-194.
Gandoger, M. 1906. Le genere Eriogonum (Polygonaceae). Bull. Soc. Bot. Belgique 42: 183-200.

Goodman, G, J. 1934. A revision of the North American species of the genus Chorizanthe. Ann. Missouri Bot.
Gard. 21: 1-102.

. 1939. A new species of Chorizanthe. Leafl. W. Bot. 2: 193-195.
. 1943. Notes on Chorizanthe. Leafl. W. Bot. 3: 230.
. 1947. A new Eriogonum from the southeast. Bull. Torrey Bot. Club 74: 329-331.
. 1948. Notes on Eriogonum. Amer. Midl. Naturalist 39: 498-504.
. 1955. The genus Acanthogonum, tribe Eriogoneae. Leafl. W. Bot. 7: 234-235.
. 1957. The genus Centrostegia, tribe Eriogoneae. Leafl. W. Bot. 8: 125-128.
Greene, E. L. 1897. New species of Eriogonum. Pittonia 3: 199-201.
. 1902. New species of Eriogonum. Pittonia 5: 67-71.
. 1910. Three new eriogonum. Muhlenbergia 6: 1-3.
Gross, H. 1913a. Beitrage zur Kenntnis der Polygonaceen. Bot. Jahrb. Syst. 49: 234-339.
. 1913b. Polygonaceae nonullae novae. Bot. Jahrb. Syst. 49: 340-348.
Heller, A. A. 1907. Compilations. The genus Eriogonum by Michael Gandoger. Muhlenbergia 3: 83-96.
Hardham, C. B. 1964. A new California Chorizanthe. Leafl. W. Bot. 10: 95-96.
. 1989. Chromosome numbers of some annual species of Chorizanthe and related genera (Polygon-
aceae: Eriogonoideae). Phytologia 66: 89-94.
Hess, W.J. & J.L. Reveal. 1977. A revision of Eriogonum (Polygonaceae) subgenus Prerogonum. Great Basin
Naturalist 36: 281-333.
Hitchcock, C. L. 1964. Polygonaceae. Univ. Wash. Publ. Biol. 17(2): 102-182.
& A. Cronquist. Flora of the Pacific Northwest. Seattle: Univ. Washington Press.

Hooker, J. D. 1880. “Polygonaceae,” pp. 88-105. In: G. Bentham & J. D. Hooker, Genera plantarum. Vol. 3.
London: L. Reeves and Co.

Howell, J. T. 1950. Variation in Eriogonum spergulinum. Leafl. W. Bot. 6: 75-81.
. 1952a. Eriogonum notes. Leafl. W. Bot. 6: 177-170.
. 1952b. Eriogonum notes II. E. polypodum. Leafl. W. Bot. 6: 177-170.
. 1952c. Eriogonum notes III: Confusions in E. ursinum. Leafl. W. Bot. 6: 225-229.
. 1955. Eriogonum notes IV; A new species from California. Leafl. W. Bot. 7: 237-238.
. 1956. Eriogonum notes V: E. glandulosum, with a new variety. Leafl. W. Bot. 8: 37-39.
. 1963. Eriogonum notes VI: Varieties of E. douglasii and E. ochrocephalum. Leafl. W. Bot. 10: 13-15.
. 1976. Eriogonum notes VII. Mentzelia 1: 17-22

Jaretzky, R. 1925. Beitrage zur systematik der Polygonaceae unter beriicksichtigung des oxymethyl-anthrachinon-
vorkommens. Repert. Spec. Nov. Regni Veg. 22: 49-83.
Jepson, W. L. 1913. Polygonaceae. Fl. Calif. 1: 376-428.
. 1923. A new species of Eriogonum from California. Madrono 1: 115.

Reveal: Checklist of the Eriogonoideae (Polygonaceae) 293

. 1925. Manual of the flowering plants of California. Berkeley: Student Store.
Jones, M. E. 1898. Eriogonum. Contr. W. Bot. 8: 39.
. 1903. Eriogonum. Contr. W. Bot. 11: 4-18.
. 1908. Eriogonum. Contr. W. Bot. 12: 73-74.
. 1933. Eriogonum. Contr. W. Bot. 18: 34-35.
Mason, H. L. 1938. A hybrid Ertogonum. Madrono 4: 290.
Michaux, A. 1803. Flora boreali-americana. Paris: C. Crapelet.
Munz, P. A. 1949. California miscellany. I. Aliso 2: 77-86.
. 1958. California miscellany. IV. Aliso 4: 87-100.
. 1959. A California flora. Berkeley: Univ. California Press.
Nuttall, T. 1817. Observations on the genus Eriogonum and the natural order Polygonaceae of Jussieu. J. Acad.
Nat. Sci. Philadephia 1: 24-31, 33-37.
- 1848. Descriptions of plants collected by Mr. William Gambel in the Rocky Mountains and Upper
California. Proc. Acad. Nat. Sci. Philadelphia 4: 7-26.
Parry, C. C. 1884. Chorizanthe R. Brown. Revision of the genus, and rearrangement of the annual species—with
one exception, all North American. Proc. Davenport Acad. Nat. Sci. 4: 45-63.
. 1885. Notes on Chorizanthe lastarriea, Parry. W.-Amer. Sci. 1: 29-31.
. 1886a. Harforida Greene and Parry. A new genus of Eriogoneae from Lower California. Proc. Daven-
port Acad. Nat. Sci. S: 26-28.
. 1886b. Lastarriwa, Rémy. Confirmation of the genus, with characters extended. Proc. Davenport Acad.
Nat. Sci. 5: 35-36.
. 1889. Chorizanthe, R. Brown. Review of certain species heretofore improperly characterized or wrongly
referred; with two new species. Proc. Davenport Acad. Nat. Sci. 5: 174-184.
Rémy, J. 1851-1852. “Polygoneas,” pp. 263-293. Jn: C. Gay, Historia fisca y politica de Chile ... Botanica. Vol. 5,
part 3. Paris: E. Thunot y Ca.
Reveal, J.L. 1965. Eriogonum. In: Documented chromosome numbers of plants. Madrofo 18: 124.
. 1965. A new alpine Eriogonum from Nevada. Leafl. W. Bot. 10: 183-186.
. 1966. On the specific distinction of Eriogonum nutans and collinum. Madrofo 18: 167-173.
. 1966. Notes on Eriogonum — I. Leafl. W. Bot. 10: 334-335.
. 1967. Notes on Eriogonum — Il. Variations in Eriogonum atrorubens. Sida 3: 82-86.
- 1967. Notes on Eriogonum — III. On the status of Eriogonum pauciflorum Pursh. Great Basin
Naturalist 27: 102-117.
- 1967. Eriogonum. In: Documented chromosome numbers of plants. Madrofio 19: 134-136.
. 1968. Notes on Ertogonum — IV. A revision of the Eriogonum deflecum complex. Brittonia 20: 13-33.

. 1968. Notes on Eriogonum — V. A revision of Eriogonum corymbosum complex. Great Basin Naturalist
27: 183-229.

. 1968. Notes on the Texas eriogonums. Sida 3: 195-205.
- 1969. “The subgeneric concept in Eriogonum (Polygonaceae),” pp. 229-249. In: J. Gunckel (ed.), Cur-
rent topics in plant science. New York: Academic Press.
. 1969. New species of Eriogonum from Utah. Madrono 19: 289-300.
. 1969. A new perennial buckwheat (Eriogonum, Polygonaceae) from southeastern Arizona. J. Ariz.
Acad. Sci. 5: 222-225.
- 1970. Additional notes on the California buckwheats (Eriogonum, Polygonaceae). Aliso 7: 217-230.
. 1971. Notes on Eriogonum — VI. A revision of the Eriogonum microthecum complex (Polygonaceae).
Brigham Young Univ. Sci. Bull., Biol. Ser. 13(1): 1-45.
. 1972. Two new species of Eriogonum (Polygonaceae) from California and adjacent states. Aliso 7:
415-519.
. 1972. Descriptions of new species and combinations in Eriogonum (Polygonaceae). Phytologia 23:
163-178.
. 1973. Eriogonum (Polygonaceae) of Utah. Phytologia 25: 169-217.
. 1974. Two shrubby novelties in Eriogonum (Polygonaceae) from the deserts of Utah and Arizona.
Brittonia 26: 90-94.
. 1975. Eriogonum (Polygonaceae) novelties from Baja California, Mexico. Brittonia 28: 337-340.
. 1976. Eriogonum (Polygonaceae) of Arizona and New Mexico. Phytologia 34: 409-484.
. 1978. “Distribution and phylogeny of Eriogonoideae (Polygonaceae),” pp. 169-190. In: K-T. Harper,
& J.L. Reveal (eds.), Intermountain biogeography: A symposium. Great Basin Naturalist Memoirs, No. 2
Provo, Utah: Brigham Young University Press.

294 PHYTOLO GIA volume 66(3):266-294 May 1989

. 1980. The genus Eriogonum Michx. (Polygonaceae) and Michel Gandoger. Great Basin Naturalist 40:
143-148.

. 1981. Eriogonum divaricatum Hook. (Polygonaceae), an Intermountain species in Argentina. Great
Basin Naturalist 41: 143-146.

. 1981. Notes on endangered buckwheats (Eriogonum: Polygonaceae) with three newly described from
the western United States. Brittonia 33: 441-448.

. 1983. The Demoulin Rule and newly mandated combinations in Eriogonum (Polygonaceae). Taxon
32: 292-295.

. 1985. New Nevada entities and combinations in Eriogonum (Polygonaceae). Great Basin Naturalist
45: 276-280.

. 1985. Types of Nevada buckwheats (Eriogonum: Polygonaceae). Great Basin Naturalist 45: 488-492.

. 1985. An annotated key to Eriogonum (Polygonaceae) of Nevada. Great Basin Naturalist 45: 493-519.

. 1989. Notes on selected genera related to Chorizanthe (Polygonaceae: Eriogonoideae). Phytologia 66:

. 1989. A review of the genus Harfordia (Polygonaceae: Eriogonoideae). Phytologia 66: 221-227.
. 1989. Remarks on the genus Pterostegia (Polygonaceae: Eriogonoideae). Phytologia 66: 228-235.
. 1989. Notes on selected genera related to Eriogonum (Polygonaceae: Eriogonoideae). Phytologia 66:
236-245.
. 1989. New combinations and novelties in Eriogonum (Polygonaceae: Eriogonoideae). Phytologia 66:
251-265.
. 1989. The eriogonoid flora of California (Polygonaceae: Eriogonoideae). Phytologia 66: 295-416.
. & BJ. Ertter. 1977. Goodmania (Polygonaceae), a new genus from California. Brittonia 28: 427-429.
. 1977. Re-establishment of Stenogonum (Polygonaceae). Great Basin Naturalist 36: 272-280.
. 1980. The genus Nemacaulis (Polygonaceae). Madrofio 27: 101-109.
& C.B. Hardham. 1989. Three new monospecific genera of Polygonaceae subfamily Eriogonoideae
from California. Phytologia 66: 83-88.

. 1989. A revision of the annual species of Chorizanthe (Polygonaceae: Eriogonoideae). Phytologia
66: 98-198.

& J.T. Howell. 1976. Dedeckera (Polygonaceae), a new genus from California. Brittonia 28: 245-251.

& PA. Munz. 1968. “Eriogonum,” pp. 33-72. In: P.A. Munz, Supplement to A California Flora.
Berkeley: University of California Press.

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Shield, O. & J.L. Reveal. 1988. Sequential evolution of Eupholotes (Lycaenidae: Scolitantidini) on their plant host
Eriogonum (Polygonaceae: Eriogonoideae). J. Linn. Soc., Biol. 33: 51-93.
St. John, H. & F. A. Warren. 1929. Eriogonum compositum and its variations. Res. Stud. State Coll. Wash. 1: 84-89.
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. 1932. New western eriogonums. Leafl. W. Bot. 1: 29-30.
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. 1936. The genus Eriogonum, a preliminary study based on geographical distribution. San Francisco:
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. 1937a. Further studies in Eriogonum. 1. Leafl. W. Bot. 2: 45-48.

. 1937b. Further studies in Eriogonum. II. Leafl. W. Bot. 2: 52-53.

. 1938. Further studies in Eriogonum. III. Leafl. W. Bot. 2: 72.

. 1941. Further studies in Eriogonum. IV. Leafl. W. Bot. 3: 15-18.

. 1943. Further studies in Eriogonum. V. Leafl. W. Bot. 3: 200-202.

& G. L. Stebbins. 1955. Chromosome numbers in the genus Eriogonum. Leafl. W. Bot. 7: 228-233.
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i [

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