Phytologia

PHYTOLOGIA

An international journal to expedite plant systematic, phytogeographical
and ecological publication

Vol. 75 ! December 1993 No. 6

CONTENTS

i TURNER, B. L. Nyaa Hinton: Letter alu a rabid plant collector in México.
9 AAW Ase keeps es AGN eb ee ee wm peeeersece ‘eee eee Pe 417

,OCHOA, C.M., Karyoreneavetic studies on wild Ecuadorian tuber-bearing

Solanum, sect. lige PEI CNP DALLA HRM EO ILA Sih ila\ atari 422
TURNER, B. i Texas species of Mirabilis SSAC AAA ANA 432
) -NESOM, GAL. ‘Taxonomy’ of Doellingeria (Asteraceae: Astereae). ..... 452

)-MACROBERTS, M.H. & BR. MACROBERTS, Vascular flora of sandstone

outcrop communities in western Louisiana, with notes on rare and note-

worthy species. ‘ : ace WARE SARC EAC FTW E ALY DY dN 463

) , TURNER, B.L., anche gypsostrata B.L. Turner, a new name for A. hin-
4 toniorum B. L. Turner, not A. hintoniorum B.L. Turner. .......... 481
COSHECIOUS and MAGILIANE MOO Omen ely AYU WL Lyk VAN NY, 482
Pex POA Us OTS) tad Volar’ 7. GV \cicia\ersve ieyeicicl evel Neigveleels\elake/clel\espletah ol elo/eveual als) 483

Index to taxa in volume 75. .............8.. . , A. RY ANH 484
| R RA U

MAY ~ 3 1994

NEW YORK
scTanmiCar GARDEN

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Phytologia (December 1993) 75(6):417-421.

JAIME HINTON: LETTER FROM A RABID PLANT COLLECTOR IN MEXICO
Billie L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

Excerpts of a letter from Jaime Hinton to B.L. Turner are repro-
duced to illustrate some recent plant collecting experiences in México.

KEY WORDS: México, plant collecting, Hinton

The literature is replete with accounts of early plant collectors in North
America, especially México, along with their trials and tribulations (e.g.,
Berlandier 1805-1851; Seemann 1825-1871; Pringle 1838-1911; etc.). Indeed,
George B. Hiriton (1882-1943), the father of Jaime Hinton whose exploits are
touted here, was a renowned collector of Mexican plants; much of the senior
Hinton’s activity has been chronicled by Hinton & Rzedowski (1972; J. Arnold
Arb. 53:141-181).

These early Mexican collectors were an unusual breed, often risking (and
sometimes losing!) their lives in the hope or realization that their discoveries
in the field might enrich all of botanical science, to say nothing of the long-time
legacy of their exploits, resulting eponymy, or whatever. In those bygone days
when practically every plant collection stood at least a fifty-fifty chance of
being undescribed, the impetus for collecting in remote, previously unvisited
areas, must have been irresistible to many, if not most.

But what about modern collectors? Have they outlived the perils of collect-
ing, the sense adventure and discovery that accompanies the field worker on
a sortie to some out-of-the-way site (albeit only 10-50 kilometers along a dirt
road from some paved major highway)? Obviously not, to judge by a recent
letter written to me by Jaime Hinton, giving an account of his and his son’s
attempt to collect in the remote mountainous regions of Nuevo Leon, México.
One might argue that “the hunt” of present day collectors has never been more
exciting, simply because what is expected in the way of novelties is drastically
reduced. Thus the reward of discovery is vastly enhanced, not to mention the
knowledge and sense of intellectual responsibility that the educated, environ-
mentally informed collector must feel as he treads the few remaining wilderness

417

418 PHYTOLOGIA volume 75(6):417-421 December 1993

areas looking for a last survivor of man’s pernicious onslaught upon pristine
habitats, most of this brought on by the senseless rampant reproduction of
mankind and the consumptive consumerism that accompanies such activity, a
consumption eating at the well springs of biodiversity everywhere.

George Hinton, his son Jaime and his grandson George represent ‘three
generations of plant collectors in Mexico, all avid students of that nation’s
flora. I never met the deceased member of this trio, but I know personally and
correspond erratically with the second and third generations, both zealous
collectors like their forefather George. Since this “introduction” is largely
meant to accompany excerpts from a letter written by Jaime, I will digress here
to attempt some encapsulation of Jaime Hinton’s physiognomy, personality,
character, and style.

Jaime is a wiry, resilient man about 5 feet 10 inches tall with the gait
of a western cowhand, what with his certain, unobtrusive, strides and his
ambience of belonging to his particular territory. A Mexican citizen, but of
British parents, green-eyed and greying at the temples, he first walked up to
me wearing a large Tarascan sombrero and a wisp of smile, extending his hand,
“Prof. Turner, I assume,” eyeing my newly married quite lovely wife Gayle (25
years or more younger than either of us) as if she might be a remarkable flower
to be plucked precariously off some Mexican bluff given the odd discovery,
wherever. Good sensible man, I thought, excellent tastes. And, later, settling
down as his guest at Rancho Aguililla, I marveled at his conversational abilities
about plants, architecture, peoples, commerce, and world affairs. He was a
consummate scholar and litterateur, and as to bearing he reminded me right
off as a protagonist from one of John Huston’s westerns, “Treasure of the
Sierra Madre”, perhaps. Whatever; I was enthralled. Later my wife said, “An
attractive man, Jaime”, I knew then that his peripheral glances were properly
catalogued.

The day after our first meeting Jaime insisted that the two of us take
a short field trip to the mountains east of Cerro Potosi. He took off in his
souped-up Ford at 100 plus miles per hour. Truly, the fastest I’d ever traveled
in an automobile. When I tactfully complained at the speed (“What’s the
rush?”) he grinned, like Socrates might have, given the same admonition from
his friends about sipping too fast his extract of hemlock, and responded “Hell,
I can’t wait to get in the field, not much time left in the day”, or something
like that, as if I too were wrapped up in his provocative enthusiasm. Anyway,
he slowed down to 95 or so for the rest of the paved road, then down to 50
on dirt, and finally lurched to a stop high up in the hills along an overgrown
semitropical gully, taking off upslope like a botanist bewildered, collecting
what was in flower or fruit, commenting on environmental degradation, the
catholic condition, confessions, confusions, whatever. Ten years older than
me, perhaps, he was clearly better shod with a better bridle.

But on to a single long excerpt, from his most recent letter, which I repro-

Turner: Letter from J. Hinton 419

duce here with his permission. And only over protestations of a sort: he would
not wish ostentation or advertisement. I responded, “Me neither, but future
generations ought to know the tribulations of plant collectors working in this
part of our century, how they knew absolutely that this was their last chance
to do something meaningful for mankind’s intellectual pursuits, that someone
cared about what.once was here, please ....”

He relented and the excerpts follow.

Dear Billie: June 25, 1993
. ... We’re hard at work in El Viejo, where we notice quite a few
species that were new when we collected them at other places not
so long ago. But we still hope to find some interesting things, espe-
cially some of those intriguing little orchids named by Carol Todzia.
Once done with El Viejo, then I can sell the four-wheel drive, and
buy a delightful smooth-riding turbo. Would you believe it that
our roughriding gasguzzling fourwheeldrivingsonofabitch got stuck
up at Agua Leon last week-for a mere eighteen galling hours. Fi-
nally, through slipping and skidding, the sob wound up at the edge
of an abyss, and I was sorely tempted to pull out the stones we
had under the other three wheels, and let the sob go. However,
we were out in the middle of no where, and the insurance people
had recently paid me for a total-loss on a four-wheel-driving Ram-
charger, so we finally got a tackle with three woodsmen, and tied
to a treetrunk actually pulled the damn truck sidewise from the
abyss, until I could coast down to a niche and turn around. But I
think it does an old fart good to have the shit scared out of him
now and then, Billie, don’t you agree? Afterwards, at least for a
time, an ordinary life seems by comparison quite enchanting.
This last trip, from which I returned last night, showed me
the colossal difference adequate chains can make on a sob. (Four-
wheeldrive = sob). Due to rampaging rainstorms, no lumber trucks
had been on the move for eleven days, so the whole range of El Viejo
was my preserve. I hate meeting those trucks coming down the
mountain, and having to back up a mile or two on the steep scary
tracks before they can pass, with thousand-meter drops nudging
me. Then, if C’s with me, I turn the truck over to him for a while.
Incidentally, before the rains began, while El Viejo was dry
as tinder, a forest fire broke out on the summit, burned fifteen
days, and consumed the whole top of the mountain (utterly free of
grazing) before it was finally put out by a hundred men, includ-
ing Federal Troops. Supposedly, the fire was started by lightening,
which is often blamed for our forest fires. But as you well know,
we don’t have forests like those of Oregon and British Columbia,

420 PHY TODO Ga volume 75(6):417-421 December 1993

where you have impossible jumbles twenty-feet deep of new and
ancient humus. Our forests are open, park-like, and I think they
are almost always deliberately set on fire by one Miguelito, who
invariably blames “un trueno” for the fire. As you may recall
Mathiasella bupleuroides was a dominant species on the heights
of El Viejo, which led me to solemnly promise Dr. Constance some
seed. But now that it’s utterly gone, what’ll I do about my solemn
promise? As the Jamaicans say—sheeeeeit, man? Maybe you could
tell Dr. Constance that I didn’t get his seed because I broke my
ass, or something. (Kidding aside, though, I’ve found a few Math-
iasella’s down below, and hope they’ll produce some seed for Dr.
Constance-if the peripatetic asshole goat don’t beat me to them.)

We’ve been trying to get a permit to collect and send herbar-
ium specimens abroad, and we seem, strangely enough, to be on
the right track. Among other things, I pointed out to the Lords of
Inexorable Reason, that there are only about thirty botanical col-
lectors in all Mexico. Assuming that each collector makes twenty
trips a year, which is a lot, and assuming that at each trip each
collector takes ten kilos of specimens, which is again a lot, (and
without dwelling upon the fact that much collecting is a matter
of pruning, which increases growth), we have 6,000 kilos of veg-
etation, a mere six tons of herbarium specimens a year. On the
other hand, we have thirty-six million head of cattle, three mil-
lion horses, and twenty-one million goats, sheep and pigs, for a
total of sixty million grazing beasties. Assuming that half of these
are properly taken care of in adequate grazing lands, which is a
lot, and that the other half are turned out to graze the national
territory helterskelter, we have thirty million cows, horses, goats,
sheep and pigs eating not forage crops but everything in sight, in-
cluding a coupe of tourists from Topeka, Kansas. Each of these
miserable mangy starving shambling slutty slattern shabby shitty
shiftless shameful sore-assed animals consumes at least ten kilos of
vegetation per day, or a yearly 3,650 kilos, for a total of a hundred
and eighty two million tons a year. Now, I ask you, Billie, with
the aid of your trusty computer and other secret methods you no
doubt have at your distinguished disposal, if you were a bush, a
tree, a terrestrial orchid, or even an untouchable German tourist,
what do you think would do more damage to our flora, thirty col-
lectors bringing home, along with a moldy piece of pork crackling
they gnawed at but didn’t finish for lunch because of three broken
teeth, six selected tons of herbarium specimens a year, or having
thirty million mangy starving shitty shabby shady screwly sheddy
sore-assed cattle chomping their way, just prior to dropping dead

Turner: Letter from J. Hinton 421

of inanition, blind staggers, aids and Almyer’s disease, chomping
their way through a hundred and eighty-two million tons of as-
sorted but unsustaining vegetation? I can tell you honestly, Billie,
that at this question, rhetorical as it might seem to you and Guy, I
could see a blush of shame mingled with a new and corruscating en-
lightenment dawning upon the faces of our honorably distinguished
bureaucrats. Now, before you accuse me of slovenly thinking, by
acidly pointing out that I’ve skipped both the not inconsiderable
multitudes of donkeys and mules ravaging our countryside, let me
hasten to assure you, Billie, that I’m saving both donkeys and
mules as weapons of last resort. In case I ever find myself on the
losing end of the argumentative stick, supposing some enlightened
bureaucrat were to advance a disquisition to the effect that botan-
ical collectors consume not ten but ten thousand kilos a trip, I
could providentially throw the donkeys and mules into the gap,
and still come out a winner of the scrap. But where would you
place the emphasis? With the six tons of herbarium specimens we
discriminating mortals collect each year, or with the hundred and
eighty-two million tons devoured by our wretched scurvy scroung-
ing scurrilous shitty cattle? Put a starving cow into a mixed forest,
and what chance of survival does anything lower than a tree have?
Off some trees, they’ll even eat off the bark, girdling the trees as
they die of hunger. ....

Kindest personal regards to you both.
Jaime

Phytolog1a (December 1993) 75(6):422-431.

KARYOTAXONOMIC STUDIES ON WILD ECUADORIAN TUBER-BEARING
SOLANUM, SECT. PETOTA

C.M. Ochoa

Genetic Resources Department, International Potato Center (CIP), P.O. Box
5969, Lima, PERU

ABSTRACT

A taxonomic and distributional summary is presented for the tuber-
bearing potatoes of Ecuador.

KEY WORDS: Solanum, Solanaceae, Ecuador, karyotaxonomy

After the recent monographic publications on Bolivian potatoes (Hawkes
& Hjerting 1989; Ochoa 1990), the tuber-bearing Solanum from Ecuador are
some of the least known in sect. Petota, subsect. Potatoe. In the present
paper, the author gives a brief summary of the Ecuadorian wild potato species
as a result of his explorations and field works made in Ecuador, as well as
his observations of living plants in CIP’s experimental plots, and laboratory
research. Exceptions, however, are S. baezense Ochoa (series Conictbaccata),
S. andreanum Baker, and S. serratoris Ochoa (series Tuberosa), of which I
did not have living material. Likewise, although I have made some herbarium
collections, I have not included in this treatment and will not include in any
of my further work, S. juglandifolium Dun. and S. ochranthum Dun. (series
Juglandifolia).

Data given are mainly on the morphology of the species, habitat, geograph-
ical distribution, and my determinations on the chromosome number (2n) and
the Endosperm Balance Number (EBN).

These studies have also been complemented by examinations of exsiccatae
collected in the past by other authors and presently housed in European, and
North and South American herbaria.

The wild tuber-bearing species studied here have been taxonomically grouped
in series. If identified synonyms are known, these are given for each species.

422

Ochoa: Ecuadorian tuber-bearing Solanum 423

I. Solanum series Acaulia Juz., Bull. Acad. Sci. U.R.S.S., ser. Biol. 2:316.
1937 (nom. nud.); er Buk. & Kameraz, Bases of Potato Breeding. 21.
1959.

Solanum albicans (Ochoa) Ochoa, Phytologia 54(5):392. 1983. BA-
SIONYM: Solanum acaule Bitt. var. albicans Ochoa, Agronomia,
Lima 27:363-364. 1960. Solanum acaule Bitt. subsp. albicans (Ochoa)
Hawkes, Scott. Pl. Breed. Rec. 117. 1963.

Plant small, rosette, short stem, very hairy, white hairs.
Leaves 3-4 pairs of leaflets without or with few interjected
leaflets. Corolla rotate, white or violet. Tubers round to elon-
gate, 2-3 cm long, white.

Distribution: This species was found for the first time in
Atocsaico, located in the Jalcas of Porcén at 3450 m alt.,
Province and Department of Cajamarca, northern Peru. Col-
lections in Ecuador were made in Cerro Quilua, 3600 m alt. in
route from Cerro Colorado to Carihuayrazo, Province Chimb-
orazo and in Romerillo, ca. 3900 m alt., Canton Ambato,
Province Tungurahua, under the V.n. of Curiquinga. Both
collections have, as do the Peruvian Solanum albicans, 2n =
72 chromosomes and EBN = 4. This species is highly resistant
to frost (-5°C).

II. Solanum series Conicibaccata Bitt. in DC., Prodr. 13(1):33. 1852.

Solanum albornozw Correll, Wrightia 2:178-179. 1961.

Leaves with numerous interstitial leaflets and (4-)5-6
pairs of folioles shortly petiolulate, glabrous or glabres-
cent, dark green and subvernicose above, puberulent in the
lower surface, margins slightly revoluted. Corolla rotate-
pentagonal, white above, white with a pale violet strip on
the back of each petal. Berry typically long-conical. Chro-
mosome number: 2n = 24, EBN = 2.

Distribution: So far it is collected only on the route
from Loja to Catamayo, 2300-2600 m alt., Province Loja,
Ecuador; mostly in humid thickets or bushes.

Solanum calacalnum Ochoa, Darwiniana 23(1):227-231. 1981.

This rare species is principally characterized by its small
branched plant, very long stolons (1.5-2.0 m); small tubers
2-4 cm), white, oval to round. Leaves glabrous, 3-4 pairs

424 PAY T+Ouhn® Gaia volume 75(6):422-431 December 1993

of leaflets with long petiolules (15-20 mm), 0-1(-2) pairs
of interjected leaflets. Corolla rotate to rotate-pentagonal,
very showy, large (4 cm). Berry long-conical with obtuse
apex, 2.5 cm long. It is very susceptible to the attack
of Phytophthora infestans and to the potato leafroll virus
(PLRV). Chromosome number: 2n = 24.

Distribution: Very restricted, so far it has been found
only on Mount La Sirena, 3000 m alt. and Sillacunga, 2900
m alt., a few km from Calacali, Province Pichincha, on
slopes of stony soil, with very poor vegetation.

Solanum colombianum Dun. in DC. Prodr. 13(1):33. 1852.

Solanum colombianum Dun. in DC. var. trianae Bitt., Fedde Repert.

Sp. Nov. 11:382-383. 1912.

Solanum dolichocarpum Bitt., Fedde Repert. Sp. Nov. 12:4-5.
1913.

Solanum colombianum Dun. in DC. var. trianae Bitt. f. quind:-
uense Buk., Suppl. 47, Bull. Appl. Bot., Genet., Pl. Breed.
225-226. 1930.

Solanum colombianum Dun. in DC. f. ztpaquiranum Hawkes, Bull.
Imp. Bur. Pl. Breed. & Genet., Cambridge. 112. 1944.

Solanum colombianum Dun. in DC. var. meridionale Hawkes, Bull.
Imp. Bur. Pl. Breed. & Genet., Cambridge. 112-113. 1944.

Solanum filamentum Correll, Wrightia 2:174-175. 1961.
Solanum caquetanum Ochoa, Phytologia 46(7):495-497. 1980.

Although the type lecality of Solanum colombianum is
Tovar, Estado de Mérida, Venezuela, in the time of Dunal,
author of this species, the present territory of Venezuela,
Colombia, and Ecuador were integrated under one nation
named La Gran Colombia; hence the epithet of colom-
bianum. This species has 3-5 pairs of leaflets and 2-4 (-
6) pairs of interjected leaflets. Corolla rotate to rotate-
pentagonal, white to light bluish or to light purple. Tubers
usually long-cylindrical or subcylindtical, up to 8 cm long
and 2 cm thick. Berries long-conical to ovoid-conical, 3.5
cm long.

En route from Leito to Rio Chico, Cordillera de Los
Leones, Province Tungurahua, at 2870 m alt., in the edges
of woods and shrubs, I found an abundant colony of Solanum
colombianum (2n = 48) locally called Papa de Monte which
must have great resistance to the attack of Phytophthora

Ochoa:

Ecuadorian tuber-bearing Solanum

infestans. Its leaves showed a type of hypersensitive reac-
tion proper for hosts with the major genes of resistance (2)
against late-blight. Small areas with cultivated potatoes
in the vicinities, on the contrary, were almost destroyed by
this fungus. Chromosome number: 2n = 48, EBN = 2.

Distribution: More in Colombia than in Venezuela or
Ecuador, especially in the provinces of Cundinamarca and
Boyaca. In Ecuador, the author found this species mostly
in Tungurahua Province. Living in cloud forest at 2500-
3500 m alt.

425

Solanum chomatophilum Bitt. f. angustifolium Correll, Wrightia 2:180.

1961.

Leaves 4-5 pairs of folioles and numerous interstitial
leaflets. Folioles narrowly elliptic-lanceolate to lanceolate
with subacute apex. Calyx asymmetric with longer lobes
than the typical form. Berries ovoid. Resistant to Phy-
tophthora infestans. Chromosome number: 2n = 24, EBN
= 2)

Distribution: Provinces Napo-Pastaza, Azuay, and Car-
chi, Ecuador, and the highlands of Department La Liber-
tad, Peru, occurs at elevations ranging from 2500-3200 m
alt., usually in cold and wet shrubby areas.

Solanum pauciyugum Bitt., Fedde Repert. Sp. Nov. 11:431. 1912.

Plant dwarf and bushy, 20-30(-50) cm tall, sparsely pi-
lose throughout. Tubers white, ovoid, 2-3 cm long. Leaves
2-3(-4) pairs of leaflets with (1-)2-3(-5) pairs of interjected
leaflets, terminal leaflet much longer than the lateral. Corol-
la rotate-pentagonal, 2.5-2.8 cm in diameter, lilac to pur-
ple. Berries long-conical, light green with 2-3 vertical darker
stripes, 2 cm long. Although it has some affinities with
Solanum flahaulti from Colombia, both species are quite
different in plant habit, leaf shape, and dissection and de-
tails of flowers. Chromosome number: 2n = 48, EBN =
ze

Distribution: Central Ecuador, mostly in the provinces
of Bolivar, Cotopaxi, Tungurahua, and Chimborazo be-
tween 3000-4000 m alt., in cloud forest, wet thickets and
grassy slopes of paramos.

Solanum tundalomense Ochoa, Ann. Cient., Univ. Agr., Lima 1(1):106-

109. 1963.

426 PHY TOL:O'G TA volume 75(6):422-431 December 1993

Plant usually very tall, 3-4 m high, branched very sparse-
ly pilose. Tubers small, 3-5 cm long, white, ovoid to long
subcylindrical. Leaves (3-)4-5 pairs of leaflets, (0-)2-5(-
7) pairs of interjected leaflets, leaflets elliptic-lanceolate or
narrowly elliptic-lanceolate with acute or acuminate apex.
Corolla rotate, white or white with pale violet stripes.
Berries long-conical, 3.5 cm long. Although this species
has affinities with Solanum colombianum, I consider them
to be different species. Besides the ploidy level, they have
substantial differences both in the shape of the corolla and
calyx morphology. It is resistant to Phytophthora infes-
tans but very susceptible to Synchytrium endobtoticum.
The chromosome number, cited formerly by the author for
Solanum tundalomense (see Ochoa 1972, p. 75) as 2n = 24,
unfortunately was mistyped. Counts made in more than
20 accessions of Solanum tundalomense from Ecuador have
given 2n = 72, EBN = 4.

Distribution: Widely distributed in Ecuador (in 10 of
20 provinces) with the highest concentrations in the provin-
ces of Azuay and Canar, occurs at elevations between 1900-
3600 m alt. In shrubby and forest vegetation.

III. Solanum series Olmosiana Ochoa, An. Cient. Univ. Agr. 3:33. 1965.

Solanum olmostanum Ochoa, An. Cient. Univ. Agr. 3:34-37. 1965.

So far, this is the only representative species of the se-
ries. Its main morphological characteristics are the shape
and dissection of the leaf, 1-3 pairs of leaflets, the irregular
and wide wings of the rachis extended all the way down
to the petiole, leaves glabrous, corolla deeply stellate and
white-cream, 2.0 cm in diameter, tubers white, oblong, 2-3
cm long, usually smooth. Solanum olmosianum was found
for the first time in the margins of Olmos River, near El
Sauce at 1640 m alt. in the Province and Department of
Lambayeque, Peru. However, I have also found it in Tabla
Rumi, at 2500 m alt., in the Province of Loja, Ecuador.
The two mentioned collections have 2n = 24 chromosomes,
EBN = 2.

Distribution: Ecuador and Peru, in the lower inter-
Andean valleys between 1600-2500 m alt., in shrubby thick-
ets.

Ochoa: Ecuadorian tuber-bearing Solanum 427

IV. Solanum series Tuberosa Rydberg, Bull. Torrey Bot. Club 51:146-147.
1924. nom. nud.

Tuberosa (Rydberg) Buk. (sensu stricto), er Buk. & Kameraz, Bases of
Potato Breeding. 18. 1959.

Andigena Buk. er Buk. & Kameraz, Bases of Potato Breeding. 24.

1959.

Transaequatorialia Buk. er Buk. & Kameraz, Bases of Potato Breeding.
21. 1959.

Vaviloviana Buk. ez Buk. & Kameraz, Bases of Potato Breeding. 18.
1959.

Andreana Hawkes, Bull. Imp. Bur. Pl. Breed. & Genet., Cambridge.
2:50. 1944. nom. nud.

Minuttfolia Correll, Texas Res. Found. Contrib. 4:216-218. 1962.

Solanum burtontt Ochoa, American Potato J. 59(6):263-266. 1982.

Plant to near 1 m tall, sparsely pilose throughout. Tu-
bers white, 2-3 cm long, ovoid. Leaves, 3-4 pairs of orbic-
ular interstitial leaflets. Leaflets rugose, cordate at base

shortly petiolulate, terminal leaflet broad ovate to elliptic-

lanceolate. Corolla rotate, small, 2 cm in diameter, light
purple-lilac outside with central petal streaks from the petal
base to tip of acumens. Berry unknown but the ovary is
pyriform. This hybridogenic species known with the ver-
nacular name of Papa de Monte or Papa Chavela has 2n =
36 chromosomes.

Distribution: Found only in Montes de Nahuasu, at
3400 m alt., between Monte Negro and Salado, just above
and behind the small waterfall in front of the village of
Banos. Living in cloud forest associated with trees (Ce-
drela, Cecropia, Juglans) and shrubs (Chusquea, Lupinus,
several species of Melastomataceae, orchids and ferns).

Solanum correlli Ochoa, American Potato J. 58(5):223-225. 1981.

Plant tall, up to 2 m high, suffrutescent. Tubers usually
moniliform. Leaves sparsely pilose, 3-4 pairs of leaflets,
shortly petiolulate, (1-)2-3(-4) pairs of interjected sessile
leaflets. Leaflets ovate to ovate-lanceolate. Calyx 5.5-6.0
mm with linear acumens 1.5-2.0 mm long. Corolla rather
rotate-pentagonal than rotate, lilac, 3.0-3.5 cm in diameter.
Berries ovoid to globose. Chromosome number: 2n = 24,

EBN 2.

428 Pa YT OUEsONG eA volume 75(6):422-431 December 1993

Distribution: So far found only near the shores of the
Angas River, to an altitude of 2700 m, Chimborazo Province.
In margins of humid forests or shrubby thickets.

Solanum minutifoliolum Correll, Wrightia 2:191. 1961.

Plant stout, erect, usually 30-60 cm tall, densely pi-
lose. Tubers ovate, white. Leaves subcoriaceous, dark
green and coarsely pubescent on upper surface, pale green,
finely pubescent on lower surface, 1-2(-3) pairs of elliptic-
lanceolate shortly petiolulate leaflets and numerous to mul-
tiple several sizes of interjected leaflets, from (6-9-)11-20 (-
26) pairs often minute, subimbricated and mostly suborbic-
ular. Terminal leaflet broader and longer than the lateral.
Peduncle densely hirsute. Corolla substellate, deep purple,
2.5 cm in diameter. Berries globose to slightly ovoid, 1.5
cm in diameter. It is quite resistant to late blight caused
by Phytophthora infestans. Chromosome number: 2n =
24, EBN = 1.

Distribution: Found in the provinces of Canar, Chimb-
orazo, and Tungurahua, occurs at elevations between 2800-
3100 m alt., mainly in cloud forest, in shrubby thickets or
margins of woods associated with ferns, orchids, Fuchsia,
Ozalis, Calceolaria, Melastomataceae, and many Composi-
tae.

Solanum regularifolium Correll, Wrightia 2:194. 1961.

Plant very simple, 50-70 cm tall, sparsely pubescent
throughout. Tubers white-yellowish, round to ovate, 3-4
cm. Leaves 3-4(-5) pairs of leaflets without interjected
leaflets, leaflets sessile to shortly petiolulate, elliptic to
elliptic-lanceolate, apex subacute to obtuse, base mostly
rounded and oblique. Corolla pentagonal, light blue with
white acumens, 2.5-3.0 cm in diameter. Calyx strongly
asymmetric, very pubescent, 7 mm long, linear acumens.
Chromosome number: 2n = 24. Very susceptible to Phy-
tophthora infestans in plant and tubers.

Distribution: Very limited, I found it only near the type
locality, south of Guasuntos, Iltus, en route Riobamba to-
wards Canar, 2400 m alt., Chimborazo Province. The col-
lection Correll & Smith P827, made near Olmos on road to
Jaen, Department Lambayeque, Peru, determined by Cor-
rell as Solanum regularifolium, in my opinion, belongs to S.

Ochoa:

Ecuadorian tuber-bearing Solanum

huancabambense Ochoa. The habitat of S. regularifolium
is a narrow and very dry valley, with poor vegetation. I
saw there only a few Gramineae and some trees of Schinus

molle L.

Solanum suffrutescens Correll, Wrightia 2:183-184. 1961.

Solanum cyanophyllum Correll, Wrightia 2:180. 1961.

Plant shrubby, very branched, slightly pubescent. Stem
subterete, slightly woody, strongly pigmented with reddish-
brown. Tubers round or long subcylindrical up to 8 cm
long and 1.5 cm thick. Leaves with narrow wings on the
rachis, 3-4(-5) pairs of leaflets and (2-)5-7(-8) pairs of in-
terjected decurrent leaflets, leaflets sessile to shortly peti-
olulate, elliptic-lanceolate with acute or shortly acuminate
apex, base obliquely rounded. Calyx asymmetrical, nar-
rowed lobes, linear acumens. Corolla rotate-pentagonal,
2.5-2.8 cm in diameter, deep purple to lilac, 2.5-3.0 cm in
diameter. Berries ovoid to subglobose. Chromosome num-

ber: 2n = 24, EBN = 2.

429

Distribution: In Ecuador, between Magdalena and Balza-

pampa, mainly in the hills of Samosurco and Pisco-urco,
also in Panjor and Guamote, at 2600-3700 m alt., Bolivar
Province. In wet thickets of valleys and near paramos in
edges of woods, frequently associated with Salvia, Calceo-
laria, Chusquea, Rubus, Compositae, and several species of
grasses.

V. Solanum series Piurana Hawkes, Ann. Mag. Nat. Hist., Ser. 12. 7:693.

1954.

Solanum chilliasense Ochoa, Lorentzia 4:9-11. 1981.

Plant about 1 m tall, glabrous or glabrescent through-
out. Tubers small, round to ovate, 1.0-2.0 cm long, white.
Leaves dark green and subvernicose above, light green and
opaque below, 2-3 pairs of shortly petiolulate leaflets and
(1-)2-3 pairs of interjected sessile leaflets; terminal leaflet
widely elliptic to elliptic-lanceolate with acuminate apex,
much larger than the laterals. Corolla rotate, lilac with
white acumens, 1.8-2.5 cm in diameter. Berries ovoid, 1.5-
2.0 cm long. This species presents a type of hypersensitive

430 PVT OM OG kA volume 75(6):422-431 December 1993

reaction to the attack of Phytophthora infestans, therefore,
it is highly valuable for potato breeding programs dealing
with major genes of resistance (R). Chromosome number:
on — ea, DN "2.

Distribution: So far has been found only in the vicini-
ties of Cordillera de Chilla, between Burro Urco and Chilola,
at 3450 m alt., El Oro Province. Usually in cold foggy
places or cloud forest among shrubby thickets or edges of
woods.

Solanum solisii Hawkes, Bull. Imp. Bur. Pl. Breed. & Genet., Cam-
bridge. 125-156. 1944.

Plant small, 30-40 cm tall, bushy, branched and rosette
near base, glabrescent to sparsely pilose throughout. Tu-
bers small, 1.0-3.0 cm, round and white. Leaves with little
shine, 1-2(-3) pairs of sessile leaflets, usually without in-
terjected leaflets, terminal leaflet larger than the laterals,
elliptic to broadly elliptic-lanceolate, lateral leaflets ellip-
tic. Corolla lilac (2.0-)2.5-3.5 cm in diameter, rotate with
short and wide acumens with deep interpetalar notches giv-
ing an outline of multilobulate aspect. Berries ovoid to
long-ovoid, 1.5-2.0 cm long.

Distribution: From central to south Ecuador, in the
provinces of Tungurahua, Canar, and Azuay, between 3500-
4000 m alt., especially in thickets of high altitude paramos
and grassy meadows.

Solanum tuquerrense Hawkes, Ann. Mag. Nat. Hist., Ser. 12. 7:693-697.
1954.

Plant robust, 50-60(-80) cm tall, glabrous or glabres-
cent throughout. Tubers long, cylindrical or subcylindri-
cal up to 8 cm long, whitish. Leaves olive-green verni-
cose above, pale green and opaque below, (2-)3-5 pairs of
slightly revolute leaflets, interjected leaflets few to many
(1-2-)4-8(-11) pairs, sessile or decurrent on the narrowly
winged rachis. Lateral leaflets broadly elliptic or ovate-
lanceolate to narrowly elliptic-lanceolate with acute or short-
ly acuminate apex, subsessile. Terminal leaflet larger than
laterals. Corolla rotate to rotate-pentagonal, blue-purple
or violet purple, 2.5-3.5 cm in diameter. Berries long-ovoid
to long-conical, 3 cm long and 1.7 cm broad. Chromosome
number: 2n = 48. EBN = 2.

Ochoa: Ecuadorian tuber-bearing Solanum 431

Distribution: From Department Narino, south of Colom-
bia to the provinces of Carchi, Imbabura, Pichincha, Co-
topaxi, and Napo in northern Ecuador at elevations be-
tween 3000-3450 m. Occurs in cold places, grassy mead-
ows, wet thickets or edges of woods.

LITERATURE REFERENCES

Correll, D.S. 1962. The Potato and its Wild Relatives. Texas Research
Foundation, Renner, Texas, 606 pp.

Hawkes, J.G. 1990. The Potato, Evolution, Biodiversity and Genetic Re-
sources. Belhaven Press, London, Great Britain. 259 pp.

Hawkes, J.G. & J.P. Hjerting. 1989. The Potatoes of Bolivia. Their Breeding
Value and Evolutionary Relationships. Clarendon Press, Oxford, Great
Britain. 472 pp.

Ochoa, C.M. 1972. El Germoplasma de Papa en Sud America. pp. 68-86
in E.R. French (ed.). Prospects for the Potato in the Developing World.
CIP. lima,-Peras 3 °°: 7

. 1983. A new taxon and name changes in Solanum (sect. Petota).

Phytologia 54(5):391-392.

. 1990. The Potatoes of South America: Bolivia. Cambridge Uni-
versity Press, New York, New York. 512 pp.

Spooner, D.M. & T.R. Castillo. 1993. Synonymy within wild potatoes
(Solanum sect. Petota: Solanaceae): The case of Solanum andreanum.

Syst. Bot. 18(2):209-217.

Spooner, D.M. & R.G. van den Berg. 1992. An analysis of recent taxonomic
concepts in wild potatoes (Solanum sect. Petota). Genetic Resources &

Crop Evaluation 39:23-37.

Phytologia (December 1993) 75(6):432-451.

TEXAS SPECIES OF MIRABILIS (NYCTAGINACEAE)
Billie L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

A taxonomic treatment of the Texas species of Mirabilis (s.1.) is ren-
dered. Thirteen species are recognized: M. albtda, M. austrotexana
B.L. Turner, spec. nov., M. comata, M. gigantea, M. glabra, M. hirsuta,
M. jalapa, M. linearis, M. longtflora, M. multiflora, M. nyctaginea, M.
otybaphoides, and M. texensis (Coulter) B.L. Turner, comb. et stat.
nov. This stands in marked contrast with the most recent accounts
of the Texas species rendered by Reed (1969) and Correll & Johnston
(1970), both treatments recognizing 29 species. All of the names used by
these authors are appropriately accounted for in the taxonomic treat-
ment, and a key to the Texas species is provided, along with maps
showing distributions.

KEY WORDS: Nyctaginaceae, Mtrabths, Orybaphus, Texas

Mirabilis (sensu lato) is a New World genus of perhaps some 50 or more
species, mostly confined to North America (Heimer! 1934). Standley (1909,
1911, 1918) and others after him, segregated from Mirabilis several natu-
ral groupings such as Aliionia L., Hesperonia Standl., Orybaphus L’Herit.,
and Quamoclidion Choisy, treating these as genera. But Standley (1931) re-
canted and reverted to Heimerl’s generic concept, and most recent workers
have tended to accept Mirabilis in the broad sense (e.g., Pilz 1978; Le Duc
1993).

Mirabilis (s.1.) is well represented in the Texas flora, the most recent treat-
ments recognizing 29 species (Reed 1969; Correll & Johnston 1970). Attempts
to use either of the latter contributions is certain to induce taxonomic con-
sternation of the most severe sort. This is largely due to the very superficial
treatment accorded the group by Reed. His treatment placed considerable
emphasis upon habit, leaf shape, and vestiture, characters which are very
variable both within and between populations. He did little, if any, field work
in connection with his study.

432

Turner: Texas Mirabilis 433

Indeed, Reed’s treatment of Mirabilis for Texas is essentially unusable; his
keys and annotations make little biological sense and, as noted in my comments
under M. austroterana B.L. Turner, one is left with the impression that he was
not deeply involved with the taxonomic process in this instance, or else had
little interest in providing a meaningful treatment with biological merit. It is
unfortunate that Correll & Johnston chose to follow his treatment; this has
caused a generation of workers, both professional and amateur, to throw up
their hands in despair, myself included.

After many years of frustration in my attempts to identify Mirabilis species
in Texas and northern Mexico, I decided to start from scratch and work up
the genus in this region based upon my own field experience, taxonomic con-
cepts, and character analysis. In this I emphasized mainly fruit characters and
placed relatively little emphasis upon leaf shape and vestiture. In addition,
I attempted to relate morphological characters, whatever their nature, with
ecogeographical variables. In short, an effort was made to recognize morpho-
geographical populational units that represent my best estimates of biological
species. I was surprised and pleased by the results obtained. Instead of the
29 species proposed for Texas by Reed, only thirteen species seem deserving
of specific status. This number might be increased to fourteen if one opts to
recognize Mirabilis dumetorum Shinners, but if the latter is to be accepted
it must bear a newly constructed name, M. latifolia (= Allionia latifolia (A.
Gray] Standl.). Mirabilis dumetorum appears to be a broad-leafed form of the
widespread excéedingly variable M: albida (Walt.) Heimer!, as noted under the
latter. “

The following key should prove useful in attempts to identify the thirteen
species recognized here. In combination with the maps provided, relatively
little difficulty should be encountered in understanding my taxonomic views
regarding this group in Texas, or elsewhere.

KEY TO TEXAS MIRABILIS

L. Penanthe:3-L7 cm. longoh toe. ibist -sBaseeee « CS. Se eee (2)
Pre CRI AAR BL 2 CREMONA o.5es 5 aiasyadate s ana'e Aa iees Cee al ele oho RIE Ve ee aa (4)
Ze involucres with d-LO"NOWEIS: o-cc ccs sressiwc oe «see weeps M. multiflora
2. cavoluctes. with. | Aowers, . seecastev s.00 Seiten veerite cee eee ae (3)
3. Perianths 10-17 cm long, mostly white. ................... M. longtflora

3. Perianths 3-6 cm long, variously pink to purple, rarely white. . M. jalapa

4. Anthocarps ovoid, ribless, glabrous and essentially smooth; trans-
Recoge tare mira nomena msn eneceting ae eae ers) SOR M. ozrybaphoides

434 POH YT .O1beO'G: EA volume 75(6):432-451 December 1993

4. Anthocarps mostly ellipsoid, variously pubescent, or if glabrous then
clearly ornate with ribs or tubercles. ©... ; <1 w.0-sashieees ome (5)

5. Anthocarps glabrous; stems stiffly erect, mostly glabrous and 1-2 m high;
mostly sandy soils of northwestern Texas. .................. M. glabra

5. Anthocarps to some extent pubescent, either pilose or short-glandular,
mostly in silty or silty-clay soils (in sandy soils mainly in trans-Pecos,
central, and southern Texas). ............--200q0s5 eee eee (6)

6. Leaves linear to linear-lanceolate, mostly 2-10 mm wide; anthocarps
conspicuously and rather evenly short-pilose, only a smattering of
much shorter glandular hairs present, if at all. ........ M. linearis

6. Leaves lanceolate to cordate, mostly 10-80 mm wide; anthocarps
variously pubescent, but if so, the leaves ovate to cordate. ....(7)

7. Stiffly erect, simple-stemmed, robust herbs mostly 1-2 m high; mostly
deep sandy soils of northcentral and southern Texas. .............. (8)

7. Sprawling to erect herbs mostly 0.3-0.8 m high; mostly alluvial, silty clay
in, calcareous soil... ...0<+..j05+ 22+ ++ 902: ome op ee (9)

8. Anthocarps conspicuously pubescent with a mosaic of mostly tufted
hairs ca. 0.5 mm long; stems strigo-puberulent, hairs strongly up-
curved and eglandular; northcentral Texas. ........ .. M. gigantea

8. Anthocarps faintly pubescent with scattered pilose hairs ca. 0.3 mm
long or less; stems pilose, hairs often_glandular, or stems glabrous
or glabrate; southern ‘Texas:..ic2 2 .c::.:a250 ee Peee M. austroterana

9. Anthocarps densely glandular-pubescent throughout with very short hairs;
trans-Pecos. s .6640<2 Peed SookGckowt.cekoed 1 ooo eee M. tezensis

9. Anthocarps variously pubescent with well-developed eglandular pilose
hairs, any glandular hairs much shorter and of secondary notability. (10)

10. Stem leaves sessile or nearly so, densely hirsute; northwestern Texas.
i Fale ctarcarlavoanals Ste aa ae ee hiss aw ./ekie ohalte oe Da hears M. hirsuta

10. Stem leaves various but usually to some considerable extent petio-
late, glabrous to sparsely or moderately hirsute. ............. (11)

11. Midstem leaves mostly 4-8 cm wide, the blades broadly obtuse, truncate
or cordate at base; flowers mostly arranged in rather congested terminal
clusters, northern: lexas.) 2:0... sess 0ss ata oe ee M. nyctaginea

Turner: Texas Mirabilis 435

11. Midstem leaves mostly 1-4 cm wide, the blades gradually tapering upon
the petioles, or abruptly truncate to cordate; flowers variously arranged
but often in open divaricate corymbose panicles. ................. (12)

12. Leaves mostly cordate; involucres melanic, pubescent with unis-
eriate multiseptate trichomes, at least the cross-walls purplish or
blackish in color; Franklin Mts., El Paso Co., rare. ....M. comata

12. Leaves mostly lanceolate, broadly ovate to rarely cordate; involucres
mostly not melanic, the trichomes with + white or tawny cross-
walls: widespread and Common... <..c00 dee. -+ =e even ven M. albida

Mirabilis albida (Walt.) Heimerl, Ann. Cons. Jard. Geneve 5:182. 1901. BA-
SIONYM: Allionia albida Walt. Mirabilis nyctaginea (Michx.) MacMil-
lan var. albida (Walt.) Heimerl, Ozybaphus albidus (Walt.) Sweet

Allionia coahuilensis Standl. Mirabilis coahutlensis (Standl.) Stand.
Ozybaphus coahuilensis (Standl.) Weatherby

Allionia grayana Standl. Mirabilis grayana (Standl.) Standl.

Alhonia latifolia (A. Gray) Standl. Ozybaphus nyctagineus (Michx.)
Sweet var. latifolius A. Gray

Allionia oblongifolia (A. Gray) Small. Mirabilis oblongifolia (A. Gray)
Heimerl. Ozybaphus nyctagineus (Michx.) Sweet var. oblongifohus
A. Gray

Allionia pseudaggregata (Heimerl) Weatherby. Mtrabils pseudaggregata
Heimerl. Orybaphus pseudaggregata (Heimer!) Standl.

Alhionia rotata Standl. Mirabilis rotata (Standl.) I.M. Johnst.

Mirabilis albida (Walt.) Heimer! var. /ata Shinners

Mirabilis dumetorum Shinners

Mirabils entricha Shinners

Mirabihs muellert Stand.

Mirabilis pauctflora (Buckl.) Standl. Orybaphus pauciflorus Buckl.

As indicated by the above partial synonymy, and many more names not
listed (cf. Reed 1969), Mirabilis albida is the most widespread highly variable
species of Mirabilis in North America. This is probably due to its phenotypic
plasticity and in large measure to its proclivity towards cleistogamic repro-
duction, presumably compounded by occasional hybridization with the many
species with which it is sympatric. In any case, I accept a wide range of habit
forms, leaf types, and vestiture in the complex. These various forms have been
keyed and recognized as this or that species by Reed and yet others. But if

436 PHY TjOL.O-GIA volume 75(6):432-451 December 1993

one examines carefully such plants they are very uniform as regards anthocarp
shape, ornamentation, and vestiture. Characteristically, their anthocarps are
markedly tuberculate, usually including the 4-5 ribs; at least to some degree,
they are irregularly pubescent with tufted white hairs ca. 0.5 mm long; be-
neath the latter there is nearly always a minute layer of much shorter glandular
hairs. Hairs of the latter type are not normally found in any large numbers
on yet other species from Texas (for example, on anthocarps of M. nyctaginea,
which has otherwise similar fruits to those of M. albida, nor are they found
on fruits of M. linearis (Pursh) Heimerl, M. glabra (S. Wats.) Standl., or M.
austrotezana, all of which might be confused with M. albida (given the aber-
rant individual among these). I am reasonably confident about my judgment
with respect to the above treatment. I am, however, not especially sure of
my relegation of M. dumetorum to synonymy. In spite of Shinners’ certainty
about its specific status, I believe what he has done is to select broad-leafed,
pubescent-stemmed forms of otherwise typical M. albida, dubbing these M.
dumetorum. For example, Travis County contains numerous sheets assignable
to both M. dumetorum and M. albida by use of Shinners’ (1951) key to species,
but these do not appear to form discrete populational units. Indeed, various
intermediate conditions in those characters states which purportedly distin-
guish between the species are found, suggesting that only a single variable
taxon is concerned. Nevertheless, I might be wrong in this conjecture and, be-
cause of this, I have shown in Figure 1 the distribution of those leaf forms (by
closed circles) which seem to conform to Shinners’ concept of M. dumetorum.
It will be seen that such plants occur over a broad region, but always confined
within the broad distribution of M. albida.

In any case, if one accepts the biological reality of Mirabils dumetorum,
its correct name must be M. latifolia, as noted in my introduction to the
present paper. The latter is based upon Orybaphus nyctagineus var. latifolius
A. Gray in Torr., U.S. and Mez. Bound. Surv. Bot. 174. 1859. TYPE: USA.
Texas: Travis Co., near Austin, May 1849, C. Wright 603 (LECTOTYPE
(designated here]: GH!). Several collections were cited or referred to by Gray in
his protologue. I have selected as lectotype one of two sheets bearing Wright’s
collection number 603, both collected in the vicinity of Austin, Texas. The
isolectotype is essentially sterile, while the lectotype itself has excellent fruiting
material, the anthocarps are almost exactly like those of M. albtda, both as to
ornamentation and vestiture.

Mirabilis entricha Shinners appears to be a form of M. albtda with some-
what longer stem-hairs than is typical for the species. I believe that most of
the other names listed in the above synonymy are reasonably certain, although
I suspect that names applied to some of the Mexican collections might ulti-
mately prove worthy of at least varietal recognition. Indeed, M. comata is very
closely related to M. albida, and might be treated as a regional morphogeo-
graphical variety of the latter without much ado; I have retained the former

Turner: Texas Mirabilis

437

Figure 1. Distribution of Mirabilis albida and M. comata in Texas and closely

adjacent areas: M. albida, leaves lanceolate to ovate (open circles); leaves ovate
to cordate (closed circles); M. comata (open triangles)

438 PHY TOL OCIA volume 75(6):432-451 December 1993

as a species because it is largely allopatric with M. albida and undeniable in-
termediates at the periphery of their distributions have not been found so as
to suggest varietal status.

Mirabilis austrotexana B.L. Turner, spec. nov. TYPE: U.S.A. Texas:
Cameron Co.: Port Isabel, near the coast in sandy soil, 20 Nov 1964,
Robert Runyon 5831 (HOLOTYPE: TEX; Isotype: TEX).

Mirabili giganteae (Standl.) Shinners similis sed differt caulibus
glabris vel pilosis trichomatibus patentibus saepe glandulosis (vs.
rigide strigosis trichomatibus incurvatis nonglandulosisque et an-
thocarpis costis laevibus, inter costas sparsim pubescentibus tri-
chomatibus minutis non caespitosisque (vs. costis nodosis, inter
costas moderate pubescentibus trichomatibus caespitosis).

Stiffy erect robust perennial herbs mostly 0.8-1.5 m high. Stems mostly
reddish brown, sparsely to densely pilose with spreading, often glandular,
trichomes, rarely glabrous throughout. Midstem or lower leaves succulent,
broadly lanceolate to ovate, sparsely pubescent to glabrous, mostly 6-12 cm
long, 2-5 cm wide; petioles 0.3-2.0 cm long. Flowers arranged in terminal
corymbose panicles 10-30 cm long, 10-15 cm wide. Fruiting involucres 8-12 cm
across, 5-lobed, the lobes united for 1/2 their length or more. Flowers mostly
3 per involucre. Corollas rotate, mostly described as pink. Anthocarps mostly
4.5-5.5 mm long, 2.0-2.5 mm wide, about equally tapering at both ends, the
5 ribs mostly smooth and glabrous to sparsely short-pilose, between these the
surface variously tuberculate, but nearly always bearing a collection of thin
short-pilose hairs readily observable at 30-40.

REPRESENTATIVE SPECIMENS (from among 40+ collections): U.S.A.
Texas. Aransas Co.: dunes, ca. 300 yards back from Gulf, 31 Apr 1965,
Turner 5164 (NY,TEX). Atascosa Co.: ca. 10 mi N of Pleasanton in deep
Carrizo sand, 6 Oct 1985, Nesom 5203 (TEX); 4 mi NE of Pleasanton, 19 May
1980, Turner 80-56M (TEX). Bexar Co.: Essar Ranch, W of San Antonio,
2 Jan 1948, Burr 227 (NY). Brooks Co.: 10 mi N of Encino, 16 Apr 1954,
Johnston 54500 (TEX); between Encino and United Carbon Black Plant, 16
Apr 1954, Johnston 54500 (TEX); Falfurrias, 30 Nov 1951, Tharp 52-561
(TEX). Cameron Co.: South Padre Isle, 3 Jun 1966, Burlage s.n. (TEX);
dunes at mouth of Rio Grande, 10 Feb 1969, Correll 36778 (LL); 5 mi W of
Boca Chica, 2 May 1940, Lundell & Lundell (LL); Brazos Island State Park,
27 Aug 1977, Richardson 2545 (TEX); same locality, 26 Nov 1977, Richardson
2606 (TEX); clay dunes along Boca Chica Road near coast, 16 Jul 1935,
Runyon 3507 (TEX); Point Isabel, 29 Apr 1959, Runyon 4669 (TEX). Jim
Wells Co.: 2 mi S of Premont, 1-5 Aug 1921, Ferris & Duncan 3249 (MO).

Turner: Texas Mirabilis 439

Kennedy Co.: near Rudolph, S of Norias, 3 Jan 1963, Correll 26919 (TEX).
Lavaca Co.: ca. 18 mi SE of Yoakum, 16 Jul 1949, Tharp 49211 (TEX).
Medina Co.: ca. 3 mi S of Devine, 28 Oct 1952, Correll 15709 (LL). Willacy
Co.: Yturria Station, 8 May 1949, Runyon 4321 (TEX).

Reed (1969), both by citation and annotation, inexplicably treated this
very natural populational complex from southern Texas (Figure 2) as belong-
ing to six disparate species: M. albida, M. dumetorum, M. ezaltata (Standl).
Standl., M. gigantea, M. nyctaginea, and M. oblongifolia. As already noted,
this was largely due to his emphasis upon habit, leaf shape, and vestiture. In
short, he keyed and recognized states of these characters as representing species
irrespective of their morphogeographical correlation with other characters.

Mirabilis comata (Small) Standl., Publ. Field Mus. Bot 8:306. 1931. BA-
SIONYM: Alkonia comata Small. Ozybaphus comatus (Small) Weath-
erby.

Reed (1969) positioned this taxon in synonymy under his concept of Mirabilis
oblongifolia. I treat the latter as synonymous with the widespread, highly vari-
able, M. albida. The type of M. comata is from southwestern New Mexico and
is part of a populational complex largely confined to Arizona, New Mexico
and closely adjacent states, including México ‘Figure 1). The taxon is closely
related to M. albida but is seemingly readily distinguished by its usually cor-
date, long-petiolate leaves, sprawling habit and involucral vestiture of mostly
darkened trichomes, as noted in the key to species. Only a single collection
has been examined from Texas (Franklin Mountains, E] Paso Co., Worthington
8472 |TEX)).

Mirabils gigantea (Standl.) Shinners, Field & Lab. 19:177. 1951. BA-
SIONYM: Allionza gigantea Standl. Orybaphus giganteus (Standl.) Weath-
erby.

As noted by Shinners (1951) this is a well-marked taxon largely confined
to loose sandy soils of north-central Texas (Figure 2). It was also retained by
Reed (1969) who confounded its distribution by citation of specimens of yet
other taxa. Mirabilis gigantea has the habit of M. austrotezana but the latter
is readily distinguished by its vestiture and anthocarps, as noted in the key to
species.

Mirabilis glabra (S. Wats.) Standl., Publ. Field Mus. Bot. 8:304. 1931. BA-
SIONYM: Ozybaphus glaber S. Wats. Allionia glabra (S. Wats.) Kuntze

440 PH Y¥eT.O L.0;G. LA volume 75(6):432-451 December 1993

Figure 2. Distribution of Mirabilis austroterana (open circles); M. gigantea
(closed circles); and M. glabra (open triangles).

Turner: Texas Mirabilis 44]

Alhonta carletona Standl.
Allionia ciliata Standl. Mirabilis ciliata (Standl.) Shinners.

Alhonia eraltata Standl. Mirabilis ezaltata (Standl.) Standl. Ozybaphus
ezaltatus (Standl.) Weatherby.

I cannot distinguish Mirabilis ezaltata from M. glabra, although Reed (1969)
and Correll & Johnston (1970) maintained both of these, distinguishing among
them by relatively trivial features (mainly leaf shape and vestiture). Shinners
(1951) also maintained M. carletonii and M. eraltata but notes that some of
the former may “have pubescent fruits instead of glabrous ones”. I presume
that this observation was due to his misidentification of robust forms of M. lin-
earis with M. carletonii (= M. glabra). In the Flora of the Great Plains (1986)
it is noted that “Some specimens [of M. glabra] are difficult to distinguish from
M. ezaltata and we suspect intergradation.” As already noted, I believe the
two are indistinguishable. Its distribution in Texas and closely adjacent areas
is shown in Figure 4.

Mirabilis hirsuta (Pursh) MacMillan, Metasp. Minn. Valley 217. 1892. BA-
SIONYM: Allionia hirsuta Pursh. Mirabilis nyctaginea (Michx.) MacMil-
lan var. hirsuta (Pursh) Heimerl. Ozybaphus hirsutus (Pursh) Sweet.

This taxon is recognized as a species with some reservation. Heimerl, as
noted in the above (only partial) synonymy, treated it as a variety of Mirabilis
nyctaginea, but I suspect that as treated by most American workers, it is
a hodge-podge of hirsute specimens belonging to several species, mainly M.
albida and M. nyctaginea. For example, Steyermark (1963), in his Flora of
Missouri retained the species, but it seems clear from his key and distribution
maps that it might be better treated as a leaf form of M. albida.

In the treatment of Mirabilis for the Flora of the Great Plains (Great Plains
Flora Association 1986) M. hirsuta is said to be rare in Kansas and Missouri,
and unreported from Oklahoma, but from my own map (Figure 4), it can be
seen that forms referable to this taxon, as identifiable by their key, occur as
far south as northern Texas and adjacent Oklahoma. In truth, I take such
plants to be hirsute forms of M. albida but have mapped these as M. hirsuta.
It should be noted that the specimens of M. hirsuta cited by Reed from Jeff
Davis County, Texas are almost certainly hirsute forms of M. albida, both
taxa occurring at the same site and apparently “intergrading” (Hanson 506a-
6 (LL,TEX)).

In short, Mirabilis hirsuta, if accepted as a biological entity, might best be
treated within the M. albida complex, but its regional distribution, interpop-
ulational variability, and typification needs additional study.

442 PHY TOLOGIA volume 75(6):432-451 December 1993

Figure 3. Distribution of Mirabilis glabrifolia (open circles) and the closely
related M. tezensis (closed circles).

Turner: Texas Mirabilis 443

Figure 4. Distribution of Mirabilis hirsuta (open circles) and M. jalapa (closed
circles) in Texas and closely adjacent areas.

444 PHY TOTO Gir volume 75(6):432-451 December 1993

Mirabilis jalapa L., Sp. Pl. 177. 1753.

Mirabilis jalapa L. var. lindhermerz (Standl.) Cory. BASIONYM:
Mirabilis jalapa L. subsp. lindhetmerz Standl. Mirabilis indhetmert
(Standl.) Shinners.

Shinners (1951), Reed (1969), and Correll & Johnston (1970) recognized
both Mirabils jalapa and M. lindheimeri as distinct species. Le Duc (1993),
who monographed the subgenus Mirabilis, did not recognize infraspecific taxa
under this widespread (Figure 4) highly variable, commonly cultivated species,
many clones of which escape cultivation and persist.

Mirabilts linearis (Pursh) Heimerl, Ann. Cons. Jard. Bot. Geneve 5:186. 1900.
BASIONYM: Allionia linearis Pursh. Ozybaphus linearis (Pursh) B.L.
Robins.

Allionia decumbens (Nutt.) Spreng. Calymenia decumbens Nutt. Mirabi-
lis decumbens (Nutt.) Daniels.

Allionia diffusa Heller. Mirabilis diffusa (Heller) Reed.
Allionia gausapoides Standl. Mirabilis gausapotdes (Standl.) Standl.
Allionia vaseyi Standl.

As conceived here, this is a widespread highly variable taxon occurring
over a broad region (Figure 5). It is sympatric with a number of other taxa
and possibly forms the occasional hybrid with them. Reed (1969) and Correll
& Johnston (1970) maintained Mirabilis decumbens, M. gausapotdes, and M.
ezaltata; the first two appear to be decumbent and erect forms of M. linearis
respectively, while M. ezaltata (the type from Kansas) appears to be an un-
usually broad-leaved, pubescent-stemmed form with achenes essentially the
same as found in typical M. linearis. It is likely that M. ezaltata is of hybrid
origin between M. linearis and M. hirsuta, the two taxa presumably occurring
in close proximity upon occasion. Regardless, the anthocarps of all of these
reputed species are seemingly identical, and are distinguished from those of M.
albida (with which it might be confused in habit) by their relatively uniform
short pilosity, and few, if any, much shorter glandular hairs beneath the pilose
vestiture.

Mirabilis longiflora L., Kongl. Svenska Vetenska Acad. Handl. 176. t.6, 1755.
Jalapa longiflora (L.) Moench

Texas material of this species belongs to the widespread Mirabilis longtflora
var. wrightiana (A. Gray) Kearney & Peebles. The var. longzflora is largely

Turner:

Texas Mirabilis

445

Figure 5. Distribution of Mirabilis linearis (open circles) and the superficially
similar M. nesomii (closed circles).

446 PHYTOLOGTIA volume 75(6):432-451 December 1993

restricted to southern México (Figure 6). Various workers have treated these
two varieties as good species, but Le Duc (1993) maintained their varietal
status.

Mirabilis multiflora (Torr.) A. Gray., Bot. Mez. Bound. Surv. 173. 1859.
BASIONYM: Ozybaphus multiflorus Torr. Quamoclidion multiflorum
(Torr.) Torr. & A. Gray.

Standley (1911) recognized this taxon, along with three others, as belonging
to the genus Quamoclidion. In 1931, however, Standley repositioned the taxon
in Mirabilis, following the treatment of Heimer! (1889). Heimer! (1934), in a
definitive monograph, retained Quamoclidion in Mirabilis, as did Reed (1969).

Pilz (1978) has provided the most recent account of Quamoclidion, treating
this as a subgenus of Mirabilis with six species, only one of which occurs in
Texas, M. multiflora. Three more or less regional allopatric varieties of the
latter were recognized by Pilz, ours belonging to the var. multiflora, which is
confined to the trans-Pecos regions (Figure 7).

Mirabilis nyctaginea (Michx.) MacMillan, Vetasp. Minn. Valley 217. 1892.
BASIONYM: Allionia nyctaginea Michx. Ozybaphus nyctagineus (Michx.)
Sweet.

Mirabilis collina Shinners.

This widespread highly variable species, with its large somewhat sprawl-
ing habit, very large broadly ovate to subcordate leaves. and subfasciculate
terminal inflorescence is rather easily recognized. In floral and fruit charac-
ters, however, it is very similar to Mirabilis albida, with which it is partially
sympatric (cf. Figures 1 and 8).

Shinners (1951) thought Mirabilis collina to be “A very restricted endemic
of the northwestern limits of the East Texas Pine Belt, suggesting a more
delicate, more showy, and finely pubescent equivalent of M. nyctaginea; flow-
ering rather early in the spring.” Reed (1969) and Correll & Johnston (1970)
retained the species. Shinners distinguished (in key form) M. collina from
M. nyctaginea by vestiture (upper internodes pubescent or glabrous, lower in-
ternodes glabrous in M. nyctaginea vs. all internodes pubescent in M. collina),
while Reed (1969) attempted to distinguish between these by mainly fruit char-
acters (4 ribs in M. collina vs. 5 ribs in M. nyctaginea) and root-branching. In
view of the considerable variation found in these characters, both within and
between populations of M. nyctaginea, I have little hesitancy in treating M.
colina as but a populational variant of the latter.

Turner: Texas Mirabilis 447

+——
200 Km

Figure 6. Distribution of Mirabilis longiflora: var. longtflora (open circles);
var. wright (closed circles).

448 Poo. Y TE OL, O:Guge volume 75(6):432-451 December 1993

Figure 7. Distribution of Mirabilis multiflora.

Turner: Texas Mirabilis 449

Mirabilis orybaphoides (A. Gray) A. Gray, Bot. Mez. Bound. Surv. 173.
1859. BASIONYM: Quamoclidion orybaphoides A. Gray. Alhoniella
orybaphoides (A. Gray) Rydb.

Ozybaphus wrighti Hemsl.

This taxon in Texas occurs only in the trans-Pecos (Figure 8). Early work-
ers positioned it in the monotypic genus Allioniella, but most subsequent work-
ers have positioned the species in Mirabilis (Heimerl 1934; Reed 1969; Correll
& Johnston 1970).

Mirabilis orybaphoides, in vegetative features, superficially resembles sev-
eral species of Mirabilis in Texas, but is readily distinguished from all such by
its smooth, glabrous ovoid achenes.

Mirabilis texensis (Coulter) B.L. Turner, comb. et stat. nov. BASIONYM:
Allionia corymbosa Cav. var. terensis Coulter, Contr. U.S. Natl. Herb.
2:351. 1894. Allionia tezensis (Coulter) Small, Fl. Se. U.S. 406. 1903.
TYPE: U.S.A. Texas. Hudspeth Co.: north base of Eagle Mountains, 3
Sep 1849, C. Wright 605 (HOLOTYPE: US!; Isotype: GH!).

Reed (1969) placed this taxon in synonymy with his concept of M. glabrifo-
lia (G. Ortega) I.M. Johnst., to which it is closely related. Except for the type,
all of the specimens cited by him belong to yet other taxa. Mirabilis glabrifo-
lia is relatively widespread in México but does not occur in Texas. Mirabilis
tezensis is readily distinguished from the latter by its thicker, more uniformly
cordate leaves. It is restricted to the Chihuahuan desert regions of western
Texas and closely adjacent Coahuila, México, as shown in Figure 3.

ADDITIONAL NAMES RECOGNIZED BY REED FOR TEXAS AND
NOT ACCOUNTED FOR IN THE ABOVE ACCOUNT

Mirabils aggregata (Ort.) Cav.

This name was originally applied to Mexican material which I treat as
synonymous with Mirabilis glabriflora Ort. The latter does not occur in Texas.

Mirabilis coccinea (Torr.) Benth. & Hook.

Reed listed this plant for Texas, but saw no specimens, nor have I. It is
native to more western areas, mainly Arizona and closely adjacent states.

450 PHY TO bOrG PA volume 75(6):432-451 December 1993

Figure 8. Distribution of Mirabilis nyctaginea (open circles) and M. ozy-
baphoides (closed circles) in Texas and closely adjacent regions.

Turner: Texas Mirabilis 451

ACKNOWLEDGMENTS

This study is based upon the study of approximately 1,000 specimens,
mostly on file at LL, TEX. It was supplemented by the examination of critical
collections and type specimens from GH and US. I am grateful to the latter
institutions for the loan of these materials. Guy Nesom kindly provided the
Latin diagnosis, and both he and Mark Mayfield reviewed the manuscript.
Jackie Poole also read the paper and made helpful suggestions.

LITERATURE CITED

Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of
Tezas. Texas Research Foundation, Renner, Texas.

Great Plains Flora Association. 1986. Flora of the Great Plains. Univ. Press
of Kansas, Lawrence, Kansas.

Heimerl, A. 1934. Nyctaginaceae, in Engler & Prantl, Die Naturlichen
Pflanzen. 2, 16C:86-134.

Le Duc, F.A. 1993. Systematic study of Mirabilis section Mirabilis (Nyctag-
inaceae). Doctoral Dissertation, Univ. of Texas, Austin, Texas.

Pilz, G.E. 1978. Systematics of Mirabilis subgenus Quamoclidion (Nyctagi-
naceae). Madrono 25:113-132.

Reed, C.F. 1969. Mirabilis, in Flora of Texas. 2:160-188.

Shinners, L.H. 1951. The North Texas species of Mirabilis (Nyctaginaceae).
Field & Lab. 19:173-182.

Standley, P.C. 1909. The Allioniaceae of the United States with notes on
Mexican species. Contr. U.S. Natl. Herb. 13:372-430.

. 1911. The Allioniaceae of Mexico and Central America. Contr.
U.S. Natl. Herb. 13:377-430.

—______.. 1918. Allioniaceae, in N. Amer. Fl. 21:171-254.

. 1931. Studies of American plants V. Field Mus. Nat. Hist., Bot.
Ser. 8:293-298.

Steyermark, J. 1963. Flora of Missouri. Iowa State Univ. Press, Ames, Iowa.

Phytologia (December 1993) 75(6):452-462.

TAXONOMY OF DOELLINGERIA (ASTERACEAE: ASTEREAE)
Guy L. Nesom
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

Doellingeria has most recently been treated within Aster, but it
is here regarded as a distinct genus comprising eleven species. The
five species of sect. Doellingeria are divided between eastern Asia (two
species) and eastern North America (three species), while the six species
of sect. Cordifolium are restricted to eastern Asia. The genus is hypoth-
esized to be as closely related to Solidago and its relatives as to Aster.

KEY WORDS: Doellingeria, Aster, Astereae, Asteraceae

The genus Doellingeria was established by Nees (1832) and recognized by
him as a group divided between North America and Asia. DeCandolle (1836)
accepted Doellingeria as a distinct genus but restricted it to Asian species,
inexplicably relegating the type (D. umbellata [Mill.] Nees) and other North
American species to the genus Diplostephium Kunth. Doellingeria was ac-
cepted for a period during the 19th century, until Bentham (in Bentham &
Hooker 1873) included it within a greatly expanded, heterogeneous Aster. Asa
Gray maintained Doellingeria as a distinct genus in various treatments but fi-
nally submerged it within Aster in his Synoptical Flora (1884), deciding to
adopt Bentham’s view. Most North American botanists subsequently have
subscribed in some degree to the concept of a conglomerated Aster advocated
by Bentham and Gray, but some have continued to recognize Doellingerza as
distinct (e.g., Greene 1896; Rydberg 1917; Small 1933; Correll & Johnston
1970). Two recent studies of Aster in a relatively broad perspective (Jones
1980; Semple & Brouillet 1980) retained Doellingeria within Aster, although
their justification for including it was not explicit.

The revisional study of Aster subg. Doellingeria (Semple et al. 1991) clari-
fied the variation patterns of the North American taxa and their corresponding
taxonomy, but the Old World taxa were not considered. Following an early
judgment by Asa Gray (1884), recent treatments by Jones (1980), Semple &

452

Nesom: Taxonomy of Doellingeria 453

Brouillet (1980), and Semple et al. (1983) have included A. retzculatus Pursh
in subg. Doellingeria, but that species is here considered to lie outside the
bounds of Doellingeria (see comments below).

Some Asian taxonomists have recently recognized the distinctiveness of
Doellingerta (e.g., Ling et al. 1985), but a number of “doellingerioid” Asian
species have been retained within Aster. Tamamschyan (1959), apparently
following DeCandolle, regarded the genus as monotypic, comprising only the
Old World D. scabra (Thunb.) Nees. In China and Japan, where the greatest
number of Doellingeria species occur, they have been treated either as Aster
or Kalimeris Cass. (Kitamura 1936, 1937; Ohwi 1965; and literature citations
below). Thus, Doellingeria as a genus has never been consolidated. The
nature of the relationship between the Asian and American species of the
genus apparently has only been considered by Bentham (in Bentham & Hooker
1873), who observed a strong relationship between the Asian D. scabra and
the American D. infirma (Michx.) E. Greene.

As interpreted here, the boundaries of Aster do not encompass Doellinge-
ria, which has ancestry closer to Solidago and related genera (comments be-
low). Doellingeria comprises eleven species in two main groups: those of sect.
Doellingeria have lanceolate, entire to serrulate, essentially epetiolate leaves,
while those of sect. Cordifolium have ovate, coarsely toothed leaves with a dis-
tinct, narrowly winged petiole. There is a named hybrid (see below) between
species of the two sections. Three species of sect. Cordifolium ser. Cordifolium
have strongly foreshortened pappus and have been treated within the genus
Kalimeris. Gu (1987, in press) excluded these species from Kalimerzs but has
not suggested an alternate placement for the group.

The five species of sect. Doellingeria are divided between eastern Asia and
eastern North America, while the six of sect. Cordifolium are restricted to
eastern Asia. Doellingeria scabra (sect. Cordifolitum) occurs widely in eastern
China, Japan, Korea, and northward into the Manchurian region of China
and Russia; D. marchandu (Levl.) Ling and D. longipetzolata (Chang) Nesom
(sect. Cordifolium) are endemic to southeastern China; and all of the other
Old World species are restricted to Japan.

TAXONOMY OF DOELLINGERIA

Complete synonymy for the New World species is found in Semple et al.
(1991); also see comments on nomenclature and typification in Jones (1980)

and Reveal (1991).

Doellingeria Nees, Gen. Sp. Aster. 177. 1832. (TYPE: Doellingeria umbel-
lata |Mill.| Nees).

454 PHY TO LOG TA volume 75(6):452-462 December 1993

Perennial, rhizomatous herbs, glabrous to sparsely strigose, eglandular.
Leaves elliptic-oblanceolate or oblong-oblanceolate without an evident petiole
to ovate-cordate with a long petiole, margins entire to coarsely toothed. Heads
solitary on leafy peduncles, borne in a corymboid capitulescence; phyllaries in
2-4 weakly to strongly graduated series, broadly elliptic-oblong to ovate with a
blunt or rounded apex, without a distinctly differentiated herbaceous tip, the
midvein commonly slightly raised and resinous, often with conspicuous lateral
nerves. Disc corollas abruptly broadened at the tube-throat junction, with
long, reflexing-coiling lobes. Ray flowers few, the ligules white, not coiling
with maturity. Achenes eglandular, otherwise sparsely strigose to glabrous,
obovoid with 5-9, raised, broad, sometimes orange-resinous, equally spaced
nerves or ribs, the achenes elongating at maturity to 3-4 mm long, nearly the
length of the involucral bracts, raising the pappus almost completely above the
involucre. Pappus 2-3-seriate, an outer series of setae or slender bristles much
shorter than the inner, and much longer bristles in one or two inner series,
all elements highly reduced in length in the three species of ser. Cordtfolium;
inner bristles with dilated apices. Chromosome number in all reported species
of Doellingerta, n=9; secondary constriction of NOR chromosome in the middle
of the short arm, the “primitive” type among various groups of Aster sensu
lato according to Semple et al. (1983).

Key to the sections of Doellingeria

Pappus 3-seriate or 2(-3)-seriate; leaves entire, lanceolate, epetiolate or nearly
so; eastern North America and eastern Asia. ........ sect. Doellingeria

Pappus 2(-3)-seriate, sometimes prominently reduced in length; leaves coarsely
toothed, ovate with relatively long, winged petioles; eastern Asia. .....
4: ua ela Grote Bias are. erties alia laine: dian0 pie a Seppe ee sect. Cordifolium

A. Doellingeria sect. Doellingeria

Aster subg. Doellingeria (Nees) A. Gray, Synopt. Fl. N. Amer. 1(2):196.
1884. Aster sect. Doellingeria (Nees) Kitamura, J. Jap. Bot. 12:721.
1936.

Diplopappus sect. Triplopappus Torr. & Gray, Fl. N. Amer. 2:182. 1841.
(TYPE: Aster umbellatus Mill.). Aster subg. Doellingeria sect.
Triplopappus (Torr. & Gray) A.G. Jones, Brittonia 32:237. 1980.

Aster ser. Sohayaktenses Kitamura, J. Jap. Bot. 12:722. 1936. (TYPE:

Aster sohayakiensts Koidzumi).

Nesom: Taxonomy of Doellingeria 455

a. Doellingeria ser. Doellingeria

1. Doellingeria infirma (Michx.) E. Greene, Pittonia 3:52. 1896.
BASIONYM: Aster infirmus Michx., Fl. Bor.-Amer. 2:109.
1803.

Doellingeria humilis (Willd.) Britt., Bratt. & Br. Illus. Fl.
3:392. 1898.
Aster cornifolius Muhl. ez Willd., Sp. Pl. 3:2039. 1803.

Appalachian Mountains in eastern United States, north-
ern Florida to New York and Massachusetts (see Semple et
al. 1991, Fig. 16).

2. Doellingeria sericocarpoides Small, Bull. Torrey Bot. Club 25:620.
1898. Aster sericocarpoides (Small) K. Schum., Just. Bot. Jahresb.
26(1):375. 1900.

Southeastern to south-central United States, North Car-
olina to Arkansas, southeastern Oklahoma, and east Texas
(see Semple et al. 1991, Fig. 15).

3. Doellingerta umbellata (Miller) Nees, Gen. Sp. Aster. 178.
1832. BASIONYM: Aster umbellatus Miller, Gard. Dict., ed.
8, no. 22. 1768.

Aster amygdalinus Lam., Encycl. Meth. 1:305. 1783. Doellinge-
ria amygdalina (Lam.) Nees, Gen. Sp. Aster. 179. 1832.

Doellingeria umbellata (Miller) Nees var. umbellata

Northeastern to east-central United States and im-
mediately adjacent Canada (see Semple et al. 1991, Fig.
3):

Doellingeria umbellata (Miller) Nees var. pubens (A. Gray)
Britt., Britt. & Br. Illus. Fl. 3:392. 1898. BASIONYM:
Aster umbellatus Miller var. pubens A. Gray, Synopt. FI.
N. Amer. 1(2):197. 1884. Doellingerta pubens (A. Gray)
Rydb., Bull. Torrey Bot. Club 37:147. 1910. Doellngeria
umbellata (Miller) Nees subsp. pubens (A. Gray ) Love &
Love, Taxon 31:357. 1982.

Aster pubentior Cronq., Bull. Torrey Bot. Club 74:147.
1947.

Northeast-central United States and immediately
adjacent Canada, completely sympatric with var. um-
bellata (see Semple et al. 1991, Figs. 13 and 14).

456

PHY TO gO Gas volume 75(6):452-462 December 1993

4. Doellingeria sohayakiensis (Koidzumi) Nesom, comb. nov.
BASIONYM: Aster sohayakiensis Koidzumi, Tokyo Bot. Mag.
37:56. 1923.

Japan.

5. Doellingeria rugulosa (Maxim.) Nesom, comb. nov. BA-
SIONYM: Aster rugulosus Maxim., Mel. Biol. 7:333. 1870.

Japan.

B. Doellingeria sect. Cordifolium (Kitamura) Nesom, comb. nov.
BASIONYM: Kalimeris sect. Cordifolium Kitamura, Mem. Coll.
Sci. Kyoto Univ., ser. B. 8:312. 1937. (LECTOTYPE, designated
here: Biotia japonica Miq.).

Aster sect. Teretiachaenitum Kitamura, Mem. Coll. Sci. Kyoto
Univ., ser. B. 8:357. 1937. (LECTOTYPE, designated here:
Aster scaber Thunb.).

b. Doellingeria ser. Cordifolium (Kitamura) Nesom, comb. et
stat. nov. BASIONYM: Kalimeris sect. Cordifolium Kitamura,
Mem. Coll. Sci. Kyoto Univ., ser. B. 8:312. 1937. LECTO-
TYPE: Doellingeria japonica (Miq.) Nesom.

6. Doellingeria japonica (Miq.) Nesom, comb. nov. BA-
SIONYM: Biotia japonica Miq., Ann. Mus. Bot. Lugduno-
Batavum 2:170. 1866. Boltonia japonica (Miq.) Franch. &
Sav., Enum. Pl. Japon. 1:226. 1875. Asteromoea japonica
(Miq.) Matsum., Shokub. Mei-1ed. 2:41. 1895. Aster japon-
icus (Miq.) Franch. & Sav., Enum. Pl. Japon. 2:398. 1876.
Not Less. ez Nees 1832. Aster miquelianus Hara (nom.
nov.|, J. Jap. Bot. 12:338. 1936. Kalmeris miqueliana
(Hara) Kitamura, Mem. Coll. Sci. Kyoto Univ., ser. B.
8:312. 1937.

Japan.

7. Doellingerta marchandit (Levl.) Ling, Icon. Cormorph. Sin.
4:423. 1975. BASIONYM: Aster marchandii Levl., Fedde
Repert. Sp. Nov. 11:306. 1912. Kalimeris marchandit
(Levl.) Kitamura, Acta Phytotax. Geobot. 33:195. 1982.

Widespread in southeastern China.

Nesom: Taxonomy of Doellingeria 457

8. Doellingeria longipetiolata (Chang) Nesom, comb. nov.
BASIONYM: Aster longipetiolatus Chang, Sunyatsenia 6:22.
1941. Kalmeris longipetiolata (Chang) Ling, Fl. Reipubl.
Pop. Sin. 74:108. 1985.

Aster trichanthus Hand.-Mazz., Oesterr. Bot. Zeit. 90:125.
1941.

China, Szechuan province.

c. Doellingeria ser. Papposae Nesom, ser. nov.

Setae pappi longitudine corollas disci aequantes.
TYPE: Doellingerta scabra (Thunb.) Nees.

9. Doellingeria scabra (Thunb.) Nees, Gen. Sp. Aster. 183.
1832. BASIONYM: Aster scaber Thunb., Fl. Jap. 316.
1784. Eucephalus scaber (Thunb.) Gandoger, Bull. Soc.
Bot. France 65:40. 1918.

Biotia discolor Maxim., Prim. Fl. Amur. 146. 1859.

Widespread in eastern China, to Japan, Korea, and
the Manchurian region of China and Russia.

10. Doellingeria komonoensis (Makino) Nesom, comb.
nov. BASIONYM: Aster komonoensis Makino, Tokyo Bot.
Mag. 12:65. 1898.

Japan.

11. Doellingeria dimorphophylla (Franch. & Sav.) Nesom,
comb. nov. BASIONYM: Aster dimorphophyllus Franch. &
Sav., Enum. Pl. Japon. 1:224. 1875.

Japan.

HYBRIDS:

Doellingeria sekimotoi (Makino) Nesom, comb. nov. BASIONYM: Aster
sekimotot Makino, J. Jap. Bot. 7:10. 1931. Aster hashtmotoi Kitamura,
Acta Phytotax. Geobot. 3:130. 1934. [D. rugulosa (Maxim.) Nesom x
D. scabra (Thunb.) Nees; see Kitamura 1937, Ohwi 1965]

Japan.

458 PHY TOs OrGTa volume 75(6):452-462 December 1993

EXCLUDED SPECIES:

1. Doellingerta reticulata (Pursh) E. Greene = Aster reticulatus Pursh.
2. Doellingeria obovata (Nutt.) Nees = Aster reticulatus Pursh.

The alliance of Aster reticulatus with Doellingeria appar-
ently has been on the basis of its corymboid capitulescence and
other habital similarity and its tendency to produce a triseriate
pappus. In A. reticulatus, however, the peduncles, phyllaries,
and sometimes the leaves are glandular, the disc corolla lobes
are erect and relatively more shallow, the achenes are fusiform
and densely glandular, and the pappus bristles are apically
acute. The species is an integral member of the group that
includes A. acuminatus Michx. and A. nemoralis Sol. (Nesom
in prep.).

3. Doellingeria trichocarpa DC., Prodr. 5:263. 1836. =? Aster striatus
Champ. ea Benth. [Fl. Hongkong.|, Hooker’s J. Bot. Kew Gard. Misc.
4:233. 1852.

Doellingeria trichocarpa was noted in Inder Kewensis to be
a synonym of Aster striatus Benth. The rationale for this is
not clear, because Bentham (in Bentham & Hooker 1873) ap-
parently accepted both species within the Doellingeria group
of Aster. Judging from their descriptions, however, neither
species can be interpreted as Doellingeria in the present view.
Neither name has been included in Aster in relatively recent
bibliographic and taxonomic accounts of the Chinese Composi-
tae, but specimens at US originally identified as A. striatus
have been annotated as A. panduratus Walp.

4. Doellingeria ptarmicoides Nees = Oligoneuron album (Nutt.) Nesom (Ne-
som 1993).

DEFINITION OF DOELLINGERIA

Doellingeria is recognized by its (1) corymboid capitulescence, (2) strongly
graduated phyllaries with a blunt or rounded apex, without a distinctly dif-
ferentiated herbaceous tip, with the midvein commonly raised and resinous,
“and often with conspicuous lateral nerves, (3) few ray flowers, the ligules not
coiling with maturity, or at least coiling very little, (4) large, terete achenes
with broad, often resinous ribs, and (5) a 2- or 3-seriate pappus of bristles with

Nesom: Taxonomy of Doellingerza 459

dilated apices. The pappus in Doellinger1a comprises one or two inner series of
long bristles and an outer series of setae or slender bristles much shorter than
the inner. The North American species have a consistently triseriate pappus,
but within sect. Doellingeria, the pappus of the Asian D. rugulosa and D.
sohayaktensis tends to be biseriate. The pappus in sect. Cordifolium also is
mostly biseriate but the inner series tends to be congested or biseriate; the
pappus is strongly reduced in length in ser. Cordifoltum. The pappus bristles
of the inner series in all species of both sections have dilated apices.

Doellingerta dimorphophylla and D. japonica differ between themselves pri-
marily in relatively technical features of vestiture and the nature of their pap-
pus. The pappus of the former (ser. Papposae) is composed of slender, apically
dilated bristles 4-5 mm long in 2(-3) series; the pappus of D. japonica (ser.
Cordifolium) is reduced to broad, flat, barbellate bristles 0.5-1.0 mm long,
mostly lanceolate but sometimes with a distinctly clavellate apex. Doellinge-
ria marchandi and D. longipetiolata have similarly reduced pappus, but the
similarity between D. japonica and D. dimorphophylla in their particularly
long stylar collecting appendages, which form 1/2-3/4 the length of the style
branches, suggests that reduction of pappus may not be a reliable indicator of
relationship among these species.

SUBTRIBAL PLACEMENT OF DOELLINGERIA

The phyletic position of Doellingeria is here hypothesized to lie near the
base of the Solidagininae, near its point of divergence both from an Old World
ancestor similar to Aster sensu stricto and from one group of New World Aster
apparently closely related to the Solidagininae (7.e., the “Biotian lineage”, Ne-
som in prep.). The white rays and multiseriate pappus of Doellingeria are
similar to true Aster, but the small number of ray flowers and eglandular,
multinerved and more or less terete achenes are characteristic of the Solidagin-
inae. White rays occur in other genera unequivocally placed among yellow-
rayed Solidagininae (Nesom 1993) and they are invariably characteristic of the
Biotian lineage. Disc corollas with deeply cut, reflexing-coiling lobes and pap-
pus bristles with dilated apices occur in basal, yellow-rayed elements of the
Solidagininae as well as the Biotian lineage. Correspondingly, the distinctive
phyllaries of Doellingerta markedly resemble those of Solidago L., Oligoneuron
Small, and the small group of species that has been treated as Aster sect.
Biotia (DC.) Torr. & Gray (e.g., Jones 1980).

Doellingerta was not included in the overview of the subtribe Solidagininae
(Nesom 1993), but its morphology as well as its occurrence in eastern North
America, with other primitive members of that subtribe, also suggest that the
phyletic position of Doellingeria is in the same area. Although the radiation
of the Solidagininae was primarily in North America, one of its most primitive

460 PHY TOLO GIA volume 75(6):452-462 December 1993

members (Solidago) has a distribution disjunct between North America and
Asia. An analogous disjunction is hypothesized to occur between the southeast
Asian endemic genus Nannoglottis Maxim., which also appears to be a prim-
itive member of the Solidagininae, and the closely related, monotypic genus
Oreochrysum Nutt. of the western United States (Nesom in prep.).

Jones & Young (1983, Figs. 4 and 5) placed Doellingeria as the sister group
to the Eurasian genera Galatella DC. and Crinitaria Cass. (=Linosyris Cass.),
but the latter two have glandular, flattened, primarily 2-ribbed, and obovate
achenes and are more closely related to typical Aster. Plants of Galatella and
Crinitaria also have a strong tendency to produce glandular-punctate leaves.

ACKNOWLEDGMENTS

I thank Mark Mayfield, Marshall Johnston, and Billie Turner for their
review and comments on the manuscript, Lindsay Woodruff (MO) for help in
securing critical literature, Zai-ming Zhao (TEX) for translations of Chinese
literature, the staffs of MO and US for their help during recent visits, and the
staff of GH for a loan of specimens.

LITERATURE CITED

Bentham, G. & J.D. Hooker. 1873. Genera Plantarum. Reeve & Co., Lon-

don, Great Britain.

Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of
Tezas. Texas Research Foundation, Renner, Texas.

DeCandolle, A.P. 1836. Doellingeria. Prodr. 5:263.

Gray, A. 1884. Synoptical Flora of North America. Ivison, Blakeman, Taylor
& Co., New York, New York.

Greene, E.L. 1896. Studies in the Compositae. —III. Pittonia 3:43-63.

Gu, H.-y. 1987. A biosystematic study of the genus Kalimeris. Ph.D. disser-
tation, Washington Univ., St. Louis, Missouri.

. In press. Systematics of Kalimeris (Astereae, Asteraceae). Ann.

Missouri Bot. Gard.

Nesom: Taxonomy of Doellingeria 461

Jones, A.G. 1980. A classification of the New World species of Aster (Aster-
aceae). Brittonia 32:230-239.

Jones, A.G. & D.A. Young. 1983. Generic concepts of Aster (Asteraceae):
A comparison of cladistic, phenetic, and cytological approaches. Syst.
Bot. 8:71-84.

Kitamura, S. 1936. Les Aster du Japon; Leur classification et leur distribu-
tion (I). J. Jap. Bot. 12:529-536; (II), 640-652; (III), 12:721-729.

. 1937. Compositae Japonicae [Astereae]. Mem. Coll. Sci. Kyoto
Univ., ser. B. 8:299-399.

Ling, Y., Y.-l. Chen, & Z. Shi. 1985. Compositae (1), [Astereae]. Flora
Retpublicae Popularis Sinicae 74:73-353.

Nees, von Esenbeck, C.G. [1832] 1833. Genera et Species Asterearum.
Leonard Schrag., Nuremberg, Germany.

Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and Oligoneuron'
(Asteraceae: Astereae) with a hypothesis of their phylogenetic position.
Phytologia 75:1-44.

Ohwi, J. 1965. Flora of Japan (J.G. Meyer & E.H. Walker, eds.). Smithso-
nian Institution, Washington, D.C.

Raven, P.H. & D.I. Axelrod. 1974. Angiosperm biogeography and past con-
tinental movement. Ann. Missouri Bot. Gard. 61:539-673.

Reveal, J.L. 1991. On the lectotypification of Aster infirmus Michx. (Aster-
aceae). Phytologia 70:234-235.

Rydberg, P.A. 1917. Flora of the Rocky Mountains and Adjacent Plains.
Published by the author, New York, New York.

Semple, J.C. & L. Brouillet. 1980. A synopsis of North American Asters:
the subgenera, sections and subsections of Aster and Lasallea. Amer. J.
Bot. 67:1010-1026.

Semple, J.C., J.G. Chmielewski, & C.C. Chinnappa. 1983. Chromosome
number determinations in Aster L. (Compositae) with comments on cy-
togeography, phylogeny and chromosome morphology. Amer. J. Bot.
70:1432-1443.

Semple, J.C., J.G. Chmielewski, & C. Leeder. 1991. A multivariate morpho-
metric study and revision of Aster subg. Doellingeria sect. Triplopappus
(Compositae: Astereae): the Aster umbellatus complex. Canad. J. Bot.
69:256-276.

462 PHY T OF OGrA volume 75(6):452-462 December 1993

Small, J.K. 1933. Manual of the Southeastern Flora. Univ. North Carolina
Press, Chapel Hill, North Carolina.

Tamamschyan, S.G. 1959. Doellingeria. Fl. U.R.S.S. (ed. V.L. Komarov)
25:126-128.

Phytologta (December 1993) 75(6):463-480.

VASCULAR FLORA OF SANDSTONE OUTCROP COMMUNITIES IN
WESTERN LOUISIANA, WITH NOTES ON RARE AND NOTEWORTHY
SPECIES

M.H. MacRoberts & B.R. MacRoberts
Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A.

ABSTRACT

The floristics and edaphic factors of west Louisiana sandstone out-
crop communities are described. The soils of this open xeric community
are moderately rich in calcium and support a number of calciphiles.
Lichens and mosses are common, especially on the open rock pavement
that characterizes this community. A number of rare species occur: Tal-
inum parviflorum, Schoenolirion wrightiu, Carer meadu, and Selaginella
arentcola var. riddellit.

KEY WORDS: Sandstone outcrop, sandstone glade, calcareous
prairie, cedar glade, calciphile, Kisatchie National Forest, floristics,
Louisiana

INTRODUCTION

The eastern and southeastern United States is — or at least until recently
was — heavily forested. Nonetheless, there were natural openings, usually of
small size, scattered throughout. The more xeric of these openings — vari-
ously referred to as prairies, glades, and barrens — have long attracted the
attention of naturalists, ecologists, and botanists, and there is a fairly large
literature dealing with them (e.g., Ebinger 1979; Perkins 1981; DeSelm 1986,
1990; Greller 1988; Baskin & Baskin 1989; Bartgis 1993).

In two previous papers, we have described sandstone glades in western
Louisiana (MacRoberts & MacRoberts 1992, 1993). As our studies of open
xeric communities in this area have expanded, we have become aware that
there are at least two different types of sandstone related communities (Mac-
Roberts & MacRoberts 1993). The type studied previously - referred to

463

464 Pir y, FO. OXG A volume 75(6):463-480 December 1993

as glade or sandstone glade - is an open area, often mesa-like, with acidic
low-nutrient soils strewn with boulders and scattered with old, slow growing,
stunted trees. The sandstone community described in this paper — referred to
as sandstone outcrop or simply outcrop — while superficially similar to glades,
is floristically and edaphically quite distinct. Among other things, these com-
munities have a rock pavement or ledge, not boulders, upslope from which is
open calcareous prairie-like habitat. An examination of the literature suggests
that these openings most resemble cedar glades of Tennessee and Kentucky,
and barrens in southeastern Texas (Baskin & Baskin 1975, 1985; Marietta &
Nixon 1984; Bridges & Orzell 1989; Mohlenbrock 1993).

In this paper we describe outcrop communities in the Kisatchie National
Forest in western Louisiana, an area for which such communities have not
yet been described. We also compare these communities with the sandstone
glades that we have studied previously, and briefly discuss calcareous prairies
and forests in this part of Louisiana.

STUDY SITES AND METHODS

Three outcrops were selected for detailed study. All occur within 1 km
of each other in T6N R8W, about 5 km north of Kisatchie, Louisiana, in the
Kisatchie Ranger District of the Kisatchie National Forest (Caldwell 1991;
Martin & Smith 1991). Two of these (KG30-3 and KG30-8) have large ex-
panses of sandstone pavement. The third (KG30-2) does not, and while un-
derlain by sandstone bedrock, has not eroded down to it except in a few small
areas. Consequently, KG30-2 represents what can be considered an earlier
stage in the evolution of this community. KG30-8 is about 0.4 ha, KG30-2
about 0.6 ha, and KG30-3 about 1.2 ha. All occur at approximately 75 meters
above sea level.

Following Perkins (1981) we divide outcrops into life zones (Figure 1).
These are 1) eroded area below the lip of the sandstone bedrock, 2) bare rock
pavement, 3) pockets of shallow soil on rock pavement, 4) sloping prairie above
pavement with soils of varying depth depending on distance from exposed rock
and degree of slope, and 5) tree/shrub zone uphill.

Not all outcrop communities have all zones. In the three we studied, KG30-
3 and KG30-8 had all zones; KG30-2 consisted almost exclusively of zones 4
and 5, with only small areas of 2 and 3. Also, zone 1 at KG30-2 was heavily
treed and shaded the very narrow zones 2 and 3. Other outcrop sites in the
Kisatchie District consisted of only zones 1, 2, and 3; zones 4 and 5 had been
eroded away (Figure 1). Zone 1 is perhaps the most variable, as we attempt
to depict in Figure 1. Almost all of these communities are on hillsides, not on
hilltops. The few we encountered on hilltops were entirely eroded to bedrock,
as indicated in Figure 1.

MacRoberts & MacRoberts: Sandstone outcrop flora 465

0 50 100

METERS

ZONE: ) 1 2-3 4 5

Figure 1. Profiles of typical outcrops
with floristic zones indicated.

466 PHY ROG PA volume 75(6):463-480 December 1993

Table 1. Taxa of three sandstone outcrops.
AGAVACEAE - Manfreda virginica (L.) Rose.
AMARYLLIDACEAE - Hypozis hirsuta (L.) Cov.

CYPERACEAE - Carez caroliniana Schwein., C. flaccosperma Dewey (2), C.
meadit Dewey (2, 3), Fimbristylis puberula (Michx.) Vahl., Rhynchospora +
tnezpansa (Michx.) Vahl., R. globularis (Chapm.) Small, Sclerta cthata
Michx., S. oligantha Michx.

IRIDACEAE - Sisyrinchium sagttiferum Bickn. (2, 3).
JUNCACEAE - Juncus marginatus Rostk. (8).

LILIACEAE - Aletris aurea Walt., Allium canadense L., Nothoscordum bi-
valve (L.) Britt., Schoenolirion wrighti Sherman (3, 8), Smilaz sp.

ORCHIDACEAE - Platanthera nivea (Nutt.) Luer (3), Sptranthes lacera
(Raf.) Raf., S. praecoz (Walt.) S. Wats.

POACEAE - Agrostis elliottiana Schultes (3, 8), Andropogon tenarius Michx.,
Aristida longespica Poir., Aristida oligantha Michx., Aristida purpuras-
cens Poir. (2, 3), Azonopus affinis Chase (8), Chasmanthium sessiliflo-
rum (Poir.) Yates (3), Dicanthelium aciculare (Desv. ex Poir.) Gould &
Clark, D. acuminatum (Sw.) Gould & Clark (3), D. sphaerocarpon ( Ell.)
Gould, Eragrostis elliott: S. Wats. (8), E. spectabilis (Pursh) Steud.
(2), Muhlenbergia capillaris (Lam.) Trin. (3), Panicum anceps Michx.
(2, 8), Paspalum notatum Flugge (3), Schizachyrium scoparium (Michx.)
Nash, Schtizachyrium tenerum Nees, Setaria geniculata (Lam.) Beauv.
(3), Sporobolus junceus (Michx.) Kunth (2, 3), Vulpta octoflora (Walt.)
Rydb. (2, 3).

CUPRESSACEAE - Juniperus virginiana L.
PINACEAE - Pinus echinata P. Mill., P. palustris P. Mill., P. taeda L.

SELAGINELLACEAE - Selaginella arenicola Underw. var. riddellit (Eselt.)
Waterfall (3).

ACANTHACEAE - Ruellia humilis Nutt.
APIACEAE - Eryngium yuccifolium Michx. (2).
AQUIFOLIACEAE - Ilez decidua Walt. (8), J. vomitoria Ait.

ASCLEPIADACEAE - Asclepias longifolia Michx., A. viridiflora Raf. (2,
3):

MacRoberts & MacRoberts: Sandstone outcrop flora 467

Table 1 (continued).

ASTERACEAE - Aster dumosus L., A. linaritfolius L., A. oolentangien-
sis Ridd. (3), A. paludosus Dryand. ez Ait. ssp. hemisphericus ( Alex.)
Cronq., A. patens Ait. (2, 3), A. sertceus Vent., Bigelowia nuttallit An-
derson, Cirsium carolintanum (Walt.) Fern. & Schub. (2, 3), Coreopsis
lanceolata L., Erigeron strigosus Muhl. ez Willd. (2, 3), Gnaphalium
purpureum L. (2, 3), Helianthus angustifolius L., Heterotheca graminifo-
lia (Michx.) Shinners, Krigta virginica (L.) Willd. (3, 8), Liatris aspera
Michx. (2), L. earle: (E. Greene) Schum. (2), L. squarrosa (L.) Michx.,
Pyrrhopappus carolinianus ( Walt.) DC. (2), Silphium laciniatum L., Sol-
idago nitida Torr. & Gray (2, 3), Vernonia tezana (A. Gray) Small (2).

BIGNONIACEAE - Campsis radicans (L.) Seem. ez Bureau (3).

CAMPANULACEAE - Lobelia appendiculata A.DC., Triodanis perfoliata
(L.) Nieuwl. (2).

CISTACEAE - Lechea tenuifolia Michx. (3).

CLUSIACEAE - Hypericum gentianoides (L.) B.S.P. (3, 8), H. hypericoides
(L.) Crantz.

CONVOLVULACEAE - Evolvulus sericeus Sw.

CORNACEAE - Cornus florida L. (3).

DROSERACEAE - Drosera brevifolia Pursh (8).

ERICACEAE - Vaccinium arboreum Marsh., V. corymbosum L. (3, 8).

EUPHORBIACEAE - Croton capitatus Michx. (2), Crotonopsis elliptica Willd.,
Euphorbia corollata L., Tragia urticifolia Michx. (3).

FABACEAE - Baptisia leucophaea Nutt., Crotalaria sagittalis L. (2), Dalea
candida (Michx.) Willd. (3), D. purpurea Vent. (3), Galactia volubilis
(L.) Britt. (2, 3), Medicago lupulina L. (2), Schrankia microphylla (Dry.)
J.F. Macbr. (2, 8), Stylosanthes biflora (L.) B.S.P., Tephrosia virginiana
(L.) Pers.

FAGACEAE - Quercus falcata Michx. (2), Q. marilandica Muenchh., Q.
stellata Wang.

GENTIANACEAE - Sabatia campestris Nutt.
HAMAMELIDACEAE - Liguidambar styraciflua L.

468 Poa YT OF 0G FA volume 75(6):463-480 December 1993

Table 1 (continued).
JUGLANDACEAE - Carya sp. (3).

LAMIACEAE - Hedeoma hispidum Pursh (2, 3), Prunella vulgaris L., Salvia
lyrata L., Scutellaria integrifolia L. (2, 8), Scutellaria parvula Michx. (3).

LENTIBULARIACEAE - Pinguicula pumila Michx. (8).

LINACEAE - Linum medium (Planch.) Britt.

LOGANIACEAE - Gelsemium sempervirens (L.) St. Hil.

MYRICACEAE - Myrica cerifera L.

ONAGRACEAE - Gaura sp. (2), Oenothera linifolia Nutt. (3, 8).
OXALIDACEAE - Ozalis stricta L.

PLANTAGINACEAE - Plantago aristata Michx. (3), P. virginica L. (2, 3).
POLEMONIACEAE - Phloz pilosa L.

POLYGALACEAE - Polygala nana (Michx.) DC., P. verticillata L. (2, 3).
PORTULACACBEAE - Talinum parviflorum Nutt. ez Torr. & Gray (3, 8).
RANUNCULACEAE - Delphinitum carolintianum Walt. (3).
RHAMNACEAE - Berchemia scandens (Hill) K. Koch.

ROSACEAE - Crataegus marshallii Eggleston, C. spathulata Michx., Prunus
sp. (3), Rubus sp. (2).

RUBIACEAE - Diodia teres Walt., Hedyotis crassifolia Raf., H. nigricans
(Lam.) Fosberg (2, 3).

SCROPHULARIACEAE - Agalinis fasciculata (Ell.) Raf. (8), Agalinis pluke-
netti (Ell.) Raf. (3, 8), Agalinis skinneriana (Wood.) Britt. (2, 3), Au-
reolaria pectinata (Nutt.) Penn. (2, 3).

VERBENACEAE - Callicarpa americana L. (2), Verbena halet Small (2).
VIOLACEAE - Viola pedata L.

MacRoberts & MacRoberts: Sandstone outcrop flora 469

Table | is a list of the vascular plants found in zones 2, 3, and 4 of KG30-2,
3, and 8. The number “2” following the species indicates presence at KG30-2,
“3” presence at KG30-3, and “8” presence at KG30-8. Absence of a letter
indicates presence at all three sites.

We recorded a total of 136 taxa, representing 102 genera and 48 families for
the three outcrops. KG30-3 had 110 species and 84 genera, KG30-2 had 101
species and 78 genera, and KG30-8 had 82 species and 65 genera, which makes
these communities as rich in species as bogs (MacRoberts & MacRoberts 1992).
Plant families with the greatest representation are Asteraceae, Fabaceae, and
Poaceae, which account for 37% of the total. However, lichens and mosses,
important components of the outcrop communities especially in zones 2 and
3, are not included here.

The three outcrops are similar. Among them, Sorensen’s Index of Sim-
ilarity ranges from 74 to 78. Combining all plants from sandstone glades
(MacRoberts & MacRoberts 1992, 1993) and from sandstone outcrops, and
comparing these lists, shows that glades and outcrops are not the same com-
munity. Sorensen’s Index of Similarity between them is 49.

We visited all three study sites every two weeks from March to mid-
November 1993 to collect and identify plants. Although these communities
are rich in lichens and mosses, we did not attempt to identify them. We fol-
low MacRoberts (1984, 1989), Gandhi & Thomas (1989), and Allen (1992) in
most instances for botanical nomenclature. Voucher specimens of many of the
species collected are deposited in the Vanderbilt University Herbarium (VDB).
While the specific fire history of outcrop communities is uncertain, they are
embedded in the pyrogenic longleaf pine community and thus probably burned
with regularity in the past (Martin & Smith 1991; Smith 1991). The study
sites had not burned in several years. Soil samples were taken from all zones at
each study site and from all zones of a number of other outcrop communities
from several calcareous prairies, and from one calcareous forest. The samples
were analyzed by A & L Analytical Laboratories, Memphis, Tennessee.

To compare the spatial distribution and size of trees in outcrops with those
in other communities, we ran transects through the middle of KG30-2, 3, and
8. This totaled an area 195 meters long and 3 meters wide (585 square meters).
Within this area we mapped all trees over 1.5 meters tall and measured their
diameter at breast height (dbh).

We cut at ground level four small pines (3 loblolly and 1 shortleaf) from
zone 4 of KG30-3 to examine growth rings and thus growth rate.

We randomly selected ten temporary one meter square plots each in KG30-
2, 3, and 8. Ten plots were in the thin soils on the pavement area (zone 3)
and twenty in the deeper soils upslope (zone 4). In each we counted pine
seedlings (first and second year trees) to see if pine establishment differed
between glades and outcrops, and to determine why these communities remain

open (see MacRoberts & MacRoberts 1993).

470 PHY TOLOGIA volume 75(6):463-480 December 1993

Using aerial photographs, we located 33 additional outcrop communities
and surveyed each of these at least once, noting extent of sandstone pavement,
erosion, flora, condition, typical and rare species, size, and other features.
These surveys extended from February 1992 until December 1993.

Climatic data are given in Martin et al. (1990). Annual precipitation av-
erages about 125 cm and is fairly evenly distributed throughout the year. In
summer, temperatures rise to 35° C, which, combined with short droughts,
translates into very hot and dry conditions, especially in open areas.

RESULTS

No vascular plant grew entirely on bare rock (zone 2); these areas were
either bare or lichen covered. Lichens, mosses, and vascular plants occurred
in zone 3. Depending on soil depth, there might also be a few very stunted
pines or oaks. Lichens and mosses were found almost entirely in zones 2 and 3,
and in the shallow soils between 3 and 4. When soil depth increased, lichens
dropped out and were replaced by forbs and grasses, and by an occasional
shrub. The few trees and shrubs growing in zone 4 usually occurred in scat-
tered clumps. Zone 5 typically began abruptly as dense woods with heavy
mid- and understory.

Table 2 gives soil characteristics of the various zones. We collected soil
samples from nine outcrops. These represent all zones, but especially 3 and
4, notably near rare species such as Schoenolirion and Talinum (both occur
in zone 3). In Table 2 we have combined and averaged also, soils from several
outcrops. Soils for zone 4 are divided into two groups: 4a is the upper 15 cm;
4b is 0.5 m deep or deeper. The upper layer of zone 4 is dark grey to black,
but changes to light grey or buff between 0.25 and 0.5 m.

It was abundantly clear prior to soil analysis that the vegetation in the
outcrop openings was usually calciphilous, and that almost always in the im-
mediate vicinity of outcrops there was calcareous forest and very occasionally
remnant calcareous prairie. Species characterizing calcareous forest and prairie
are Aesculus pavia L., Andropogon spp., Apocynum cannabinum L., Aristida
spp., Berchemia scandens, Bumelia lycioides (L.) Pers., Crataegus spp., Dalea
spp., Gleditsta triacanthos L., Helianthus hirsutus Raf., Juniperus virginiana,
Neptunia lutea (Leavenw.) Benth., Prunus spp., Ratibida pinnata ( Vent.) Barn-
hart, Salvia azurea Lam., Schizachyrium spp., Schrankia microphylla (Sm.)
Macbr., and Viburnum dentatum L.

To have a standard by which to judge their soil properties and those of
associated communities, we collected and analyzed soils from two well stud-
ied calcareous prairies (Carpenter Road Prairie and Coldwater Road Prairie,
Smith et al. 1989) in the Winn Ranger District of the Kisatchie National For-
est about 65 km northeast of our study sites. We also had soils analyzed from

MacRoberts & MacRoberts: Sandstone outcrop flora

Table 2. Soil characteristics.

Exchangeable ions (ppm)
pH 4P) «KS Ca Mg

OM%

All Outcrops (Kisatchie District)

Zone | (3) 5.5 5 91 2223 285 1.0
Zone 3(15) 5.3 15 83 1193 250 1.6
Zone 4a (11) 5.4 6 102 2535 281 3.2
Zone 4b (2) 7.8 1 117 4780 346 1.0
Zone 5 (2) 5.3 14 134 3590 376 9.1

Specific Outcrops (Kisatchie District)
KG30-2

Zone 4 (2) 5.4 4 198 4290 459 3.8
KG30-3
Zone 4 (1) 5.9 3 151 3910 326 (3.3

Zone 5 (2) 5.3 14 134 3590 376 (9.1
KG30-8
Zone 4 (1) 48 7 87 720 272) A233

Prairies (Winn District)
Carpenter (3) 7.8 3 137 3667 ol: 57-4
Coldwater (2) 7.7. 1 183 5145 73° 7.0
Prairies (Kisatchie District)

Ratibida (3) 7.7 3 182 7330 90 6.7
K50H (2) 7.8 1 174 6485 60 4.6
Calcareous Forest (Kisatchie District)
K50C (1) 5.9 3 234 6530

Table 3. Tree species number and size on outcrops.

Species No. on outcrops Average dbh (cm)

(range)
Pinus palustris 149°" (5.122259)

P. taeda V2. 56.2") (2-5-1227)
P. echinata 2 “17.1 “(3-8-30-5)
Quercus marilandica 3 42 (2.5-7.6)

472 PHYTO L.O.GHLA volume 75(6):463-480 December 1993

Table 4. Tree size.

Diameter class

dbh (cm)

No. of trees

two calcareous prairie remnants (Ratibida Prairie and K50H Prairie) and one
calcareous forest located near outcrops on the Kisatchie Ranger District. The
number of samples collected and analyzed from each area, zone, and site is
shown in parentheses in the table. The average is given where there is more
than one sample.

The area in which we located outcrops during our survey is a band several
miles wide that runs east-west across the entire Kisatchie District (a distance
of about 30 km). This band appears to correspond with the Lena Member
of the Fleming Formation, the chief characteristic of which is its “calcareous
clays” (Gorat & Roland 1984).

It was not surprising therefore to find that the soil samples confirmed what
the vegetation already told us. The soils were calcareous. In some places, we
found narrow strata consisting of nothing but calcareous concretions frequently
there were small calcium aggregations scattered on the surface and mixed
throughout the soils. This admixture may account for the low pH and high
calcium in the samples.

The Natchitoches Parish soil survey classifies the areas in which the out-
crops occur as Kisatchie soils; that is, “fine, montmorillonitic, thermic Typic
Hapludalfs” (Martin et al. 1990). With the exception of high calcium, they are
identical in acidity and mineral contents to the soils of the sandstone glades
we studied earlier (MacRoberts & MacRoberts 1992, 1993).

As the data in the table show, the soils in KG30-2 and KG30-3 are as
calcareous as the soils in the calcareous prairies. While some differences exist
between the outcrop soils and those described from the prairies, notably surface
pH and magnesium, the calcium content is approximately the same.

Tables 3 and 4 give information on tree distribution in transects in outcrop
communities.

A comparison of the data given in our previous papers (MacRoberts &
MacRoberts 1990, 1993) shows that outcrops and glades are very similar in

MacRoberts & MacRoberts: Sandstone outcrop flora 473

the distribution and abundance of trees, and that they differ in a number of
ways from bogs and pinewoods. In pinewoods there was one tree per 11 square
m, in glades there was one tree per 23.5 square m, and in bogs one tree per
35 square m. We found that in outcrops there was one tree per 24 square m.
In bogs, glades, and outcrops the trees are stunted and old growth. However,
bogs lack oaks, which are common in both glades and outcrops. In outcrops,
trees are almost entirely confined to zones 1, 4, and 5. Zones 2 and 3 lack
sufficient soil for trees to survive.

In our previous study, we found that the growth rate of pines differed
significantly among glades, bogs, and pinewoods. Trees in glades grew at the
slowest rate, averaging 11.5 rings per cm; bogs came next with 8.6 rings per
cm; and trees in upland pinewoods had 3.7 rings per cm. The growth rate
of pines from KG30-3 zone 4 was intermediate between bogs and glades, with
10.75 rings per cm (the four trees had 11, 11, 11, and 10 rings per cm). While
this sample is small, it is unlikely that a larger sample would significantly alter
the results since the trees in outcrops — as in bogs and glades ~ are clearly under
stress (stunted, gnarled, and with scanty foliage).

In the outcrop communities, pine seedlings were absent in zone 2 and scarce
in zone 3. In the ten one m square plots we examined in zone 3, there were
only two seedlings. In the 20 plots from zone 4, there were 13 seedlings. In
glades, pine seedlings fared better: 50 plots had 169 seedlings (MacRoberts
& MacRoberts 1993). But the end result is the same in these two habitats.
Irrespective of the number of pines that sprout and survive for a year or two,
the vast majority eventually die. By the end of summer, after a few July and
August droughts, very few pine seedlings remain.

Why do outcrop communities remain open? Several factors seem impor-
tant (MacRoberts & MacRoberts 1990, 1993). First, edaphic conditions may
be unfavorable. The soil itself appears to be nonabsorbant.. We have exca-
vated post holes in outcrops after two days of rain only to find that the soil
is dry 10-15 cm below the surface. Also in outcrops, as in glades, there is an
impermeable layer of rock. Further, even where soils are deep, the soil char-
acteristics themselves impede woody plant establishment. The soil is high in
calcium, which is known to deter growth in many plants including pines, and
is stiff and seasonally droughty with high shrink-swell properties (Martin &
Smith 1991: 64). Open areas are subject to very high summer temperatures
and short droughts put severe stress on pine seedlings. But the fact that trees
and midstory vegetation begin abruptly in zone 5 would indicate that there is
something different between the soils in zones 4 and 5. The soil samples did
not reveal what that might be.

During the course of this study we surveyed 36 outcrop communities in the
Kisatchie District. These ranged in size from 0.1-2.0 ha (average 0.8 ha). Most
contained all zones, but several consisted entirely or almost entirely of zone 4
(t.e., were prairie-like) but were on slopes, not hilltops. That we were dealing

474 Pi. YE OM O GAA volume 75(6):463-480 December 1993

Table 5. Statistics on Schoenoltrion wrighti populations.

Outcrop Size No. Plant Cattle
(ha) plants coverage grazing

(ha)

KG30-3 1.2
KG30-5 1.2 150 0.01 yes
KG30-8 0.4 75 0.005 yes
KG36-1 0.4 1000 0.2 no
SheardI 1.2

with an outcrop community at such sites was usually evident by the flora and
also by the presence, discovered with minimal searching, of a rock ledge down
slope, hidden by shrubby vegetation, that had not eroded out to pavement
dimensions. Since all of these outcrops occurred at approximately 75 m above
sea level, and since the rock layer was similar throughout, we assume that the
same geological strata are repeated wherever outcrops occur.

NOTEWORTHY SPECIES

During the course of this study we found a number of species that deserve
additional comment.

Schoenolirion wrightw. (MacRoberts & MacRoberts 1901 [VDB]). This
species is globally, federally, and state listed as rare (G3, C2, $1 Louisiana, S2
Texas) (see Nixon & Ward 1981; Orzell 1990; Grace 1993; for literature and
recent reviews).

Between April 17, 1993, when we first discovered Schoenohtrion wrighti on
the Kisatchie National Forest and May 7, 1993, when it had ceased blooming
and was becoming difficult to locate, we surveyed 19 outcrops in the western
part of the Kisatchie Ranger District and found it at five sites (26%), often
in large numbers (Table 5). The five populations are all located in T6N R8W
a few miles north of Kisatchie, Louisiana. Within this area, the closest two
populations are about 1 km apart; the most distant are 5 km apart.

In three outcrops Schoenolirion wrighti was confined to a small area. In
the other two, it was much more widespread. In these latter two sites, cattle
had been excluded for several years. In one of the outcrops where grazing
occurred, the small population of S. wright was entirely grazed down just
after it had set seed.

In an outcrop bisected by a road, a recent non-growing season prescribed
burn (February 13, 1993) had burned the southern half. Although Schoeno-

MacRoberts & MacRoberts: Sandstone outcrop flora 475

lirion wrighti bloomed and set seed in both burned and unburned portions,
plants appeared to be more prolific and larger in the unburned area.

We examined soils in the five outcrops in which Schoenoltrion wrightiw
occurred. These soils are the same as those reported for zone 3 in Table 2 and
can be as shallow as a few inches only.

Carex meadit. (MacRoberts & MacRoberts 1889 [VDB]). Prior to the
present study, this western species had been reported only once from Louisiana
(Williams 1977; MacRoberts 1989), and specimens from three other parishes
have recently been found in herbaria (Julia Larke, pers. comm.). It is currently
ranked as rare (S1) in Louisiana. Although we made no special attempt to
search for this species, we located three outcrops where it occurred in zone 4.
Two of these are within 1 km of each other; the other is about 6 km distant.

Selaginella arenicolassp. riddellit. (MacRoberts & MacRoberts 1809 |VDB)).
Riddell’s spikemoss is rare (S1) in Louisiana. It occurs in zone 3 and is often
associated with moss or lichens. We surveyed 36 outcrops in the Kisatchie
Ranger District and found it, often in large mats, in five. The plants are easily
dislodged by cattle trampling.

Talinum parviflorum. (MacRoberts & MacRoberts 1759, 1780 |VDB)).
This plant is rare in Louisiana (S1). At the beginning of this study, it was only
known from three closely adjacent outcrops on the Kisatchie Ranger District.
We found it in 24 of the 36 outcrops we surveyed, often in large numbers (more
than 1000 plants). It grows almost exclusively in thin soils in slight depres-
sions on the rock pavements in full sun (zone 3). It blooms in late afternoon.
We collected soils in which Talinum grew from eight outcrops. All appear to
be soils typical of zone 3.

Habranthus tubispathus (L’Herit.) Traub. (MacRoberts & MacRoberts 2093
[VDB]). While not considered rare, we found this West Gulf Coastal Plain
endemic at one outcrop, where it was abundant. It did not occur at KG30-2,
3, or 8.

It is found in barrens/glade/prairie habitat in southeast Texas (Orzell
1990).

DISCUSSION

During the course of this work, it became evident that we were sealing with
a community that consisted of a sandstone outcrop and upslope a calcareous
opening that would best fit the definition of prairie. But we have chosen not to
call these upslope openings prairies after examining “true” calcareous prairies,
which appear to be floristically somewhat different, are located on hilltops, not
side slopes, and which are alkaline and usually more calcareous. Nonetheless,
the upslope openings (zone 4) should be looked at as a type of calcareous
prairie since their soils are calcareous and their flora is calciphilous.

476 PHY FT OL,0 GIA volume 75(6):463-480 December 1993

The outcrop communities occurring in the Kisatchie National Forest ap-
pear to be very similar to the open (“prairie-like” or “barrens”) communities
described by Marietta & Nixon (1984), Bridges & Orzell (1989), Orzell (1990),
and Mohlenbrock (1993) for east Texas. They are similar also to a number
of barrens, glades, and prairies described for Arkansas and Missouri eastward.
Notably similar appear to be the various cedar glades of Tennessee and Ken-
tucky (Baskin & Baskin 1975, 1985, 1989; DeSelm 1986) and the “Black Belt”
flora of Alabama (Robert Kral, pers. comm.).

We did not divide out floristic surveys according to zones. But clearly if
we had done so, the different zones would have shown significant differences.
Many plants that grow on the thin soils overlaying rock outcrops do not grow
in the upslope zones, and vice versa. For example, Talinum and Selaginella are
found only on or near rock pavements (zone 3); whereas the many composites,
peas, and grasses typically occur in zone 4. It should be remembered, however,
that what does grow on these outcrops is in part determined by the soils that
erode down onto them from above and that, in the present case, these are
usually calcareous. Since we have not had the opportunity to study outcrops
with other soils upslope, we are not in a position to say how different they
might be.

KG30-2, 3, and 8 were grazed. While this was not an ideal condition under
which to make a floristic study, it did provide us with information on the ef-
fect of cattle on these communities. In a word, cattle have a disastrous effect.
Not only do they crop the plants (they ate all the newly seeded Schoenolirion
wrighti from one outcrop), but they crush and dislodge lichens and other
plants, especially in zones 2 and 3. Trampling also initiates erosion in zones
1 and 4, where soils are so unstable (Martin et al. 1990) that massive ero-
sion results, which not only sweeps away the soil above the rock shelf, but
undermines the rock itself. The devastation caused by cattle shows that these
communities require a good stable ground cover to keep them intact. Many of
the outcrops on the Kisatchie National Forest that were once grazed are now
free of cattle, and we are happy to report that the Forest Service has fenced
the outcrops where this study took place.

One thing that did surprise us was that, although the three outcrops were
grazed, we found no noticable difference in our floristic lists between them and
outcrops that had not been grazed for some years. The only difference was that
grasses in grazed areas were difficult to find and in short supply. Undoubtedly,
a study using plots would reveal many differences in composition and number
of species present between grazed and ungrazed sites.

Our work on open xeric rocky communities in the Kisatchie National Forest
has convinced us that there are at least two distinct types (MacRoberts &
MacRoberts 1993). This year’s field work establishes that sandstone outcrops
are clearly distinct from sandstone glades.

This finding clarifies some confusion that currently exists in the Louisiana

MacRoberts & MacRoberts: Sandstone outcrop flora 477

botanical literature. The community initially described by Smith (1988) was
a sandstone outcrop, but when Martin & Smith (1991) described the major
community types of the Kisatchie District of the Kisatchie National Forest,
they reiterated Smith’s 1988 description of the outcrop community, but exem-
plified it with a glade community. Hart & Lester (1993), without additional
research or reference to the growing literature, have perpetuated the confusion
by synonymizing glade and outcrop. Future community classifications should
distinguish between sandstone outcrops and sandstone glades.

ACKNOWLEDGMENTS

The continuing cooperation and assistance of the staff of the Kisatchie
National Forest have been instrumental in making this study, as in all our
botanical research, possible. Especially to be thanked are Tom Fair, Susan
Carr, and Viola Ritchie. Allan Tiarks, Southern Forest Experiment Station,
answered some questions we had about the soil data. Financial support was
provided, in part, by a Challenge Cost-Share Agreement with the Kisatchie
National Forest. Robert Kral vetted a number of the plants, notably our
Carer specimens. Julia Larke, Louisiana Natural Heritage Program, provided
information on Carer meadii and other rare plant species. Jerry Baskin and
Robert Kral made helpful comments on the paper.

LITERATURE CITED

Allen, C.A. 1992. Grasses of Lousiana. Cajun Prairie Habitat Preservation
Society, Eunice, Louisiana.

Bartgis, R.L. 1993. The limestone glades and barrens of West Virginia.
Castanea 58:69-89.

Baskin, C.C. & J.M. Baskin. 1975. The cedar glade flora of Bullitt County,
Kentucky. Castanea 40:184-190.

Baskin, J.M. & C.C. Baskin. 1985. A floristic study of a cedar glade in Blue
Licks Battlefield State Park, Kentucky. Castanea 50:19-25.

Baskin, J.M. & C.C. Baskin. 1989. Cedar glade endemics in Tennessee, and
a review of autecology. J. Tenn. Acad. Sci. 64:63-74.

Bridges, E.L. & S.L. Orzell. 1989. Longleaf pine communities of the West
Gulf Coastal plain. Natural Areas Journal 9:246-263.

478 PHY TOL:O GIA volume 75(6):463-480 December 1993

Caldwell, J. 1991. Kisatchie National Forest: Part of a 100-year heritage.
Forests & People 41(1):35-46.

DeSelm, H.R. 1986. Natural forest openings on uplands of the eastern United
States. Pp. 366-375. In D.L. Kulhavy & R.N. Conner (eds.), Wilderness
and Natural Areas in the Eastern United States: A Management Chal-
lenge. Center for Applied Studies, School of Forestry, Stephen F. Austin
State University, Nacogdoches, Texas.

DeSelm, H.R. 1990. Flora and vegetation of some barrens of the eastern
highland rim of Tennessee. Castanea 55:186-206.

Ebinger, J.E. 1979. Vascular flora of sandstone outcrops in Clark County,
Illinois. Castanea 44:38-44.

Gandhi, K.N. & R.D. Thomas. 1989. Asteraceae of Louisiana. Sida Bot.
Misc., No. 4:1-202.

Grace, S.L. 1993. Element stewardship abstract: Schoenolirion wrightit.
Unpublished report, The Nature Conservancy of Texas, Silsbee, Texas.

Greller, A.M. 1988. Deciduous forest. Pp. 287-316 In M.G. Barbour &
W.D. Billings (eds.). North American Terrestrial Vegetation. Cambridge
University Press, New York, New York.

Groat, C.G. & H.L. Roland. 1984. Louisiana Geological Survey, Geologic
Map of Louisiana. Louisiana Department of Natural Resources, Baton
Rouge, Louisiana.

Hart, B.L. & G.D. Lester. 1993. Natural community and sensitive species
assessment on Fort Polk Military Reservation, Louisiana. Louisiana De-
partment of Wildlife and Fisheries, Baton Rouge, Louisiana. Unpub-
lished report.

MacRoberts, D.T. 1984. The Vascular Plants of Loutsiana. Bull. Museum

of Life Sciences, Louisiana State University, Shreveport, Louisiana.

MacRoberts, D.T. 1989. A Documented Checklist and Atlas of the Vascu-
lar Flora of Louisiana. Bull. Museum of Life Sciences, Louisiana State
University, Shreveport, Louisiana.

MacRoberts, B.R. & M.H. MacRoberts. 1992. Floristics of four small bogs in
western Louisiana with observations on species/area relationships. Phy-
tologia 73:49-56.

MacRoberts & MacRoberts: Sandstone outcrop flora 479

MacRoberts, M.H. & B.R. MacRoberts. 1990. Size distribution of trees in
bogs and pine woodlands in west central Louisiana. Phytologia 68:428-
434.

MacRoberts, M.H. & B.R. MacRoberts. 1992. Floristics of a sandstone glade
in western Louisiana. Phytologia 72:130-138.

MacRoberts, M.H. & B.R. MacRoberts. 1993a. Why don’t west Louisiana
bogs and glades grow up into forests? Phytologia 74:26-34.

MacRoberts, M.H. & B.R. MacRoberts. 1993b. Floristics of two Louisiana
sandstone glades. Phytologia 74:431-437.

Marietta, K.L. & E.S. Nixon. 1984. Vegetation of an open, prairie-like com-
munity in eastern Texas. Texas J. Sci. 36:25-32.

Martin, D.L. & L.M. Smith. 1991. A survey and description of the natural
plant communities of the Kisatchie National Forest: Winn and Kisatchie
Districts. Unpublished report. Louisiana Department of Wildlife and
Fisheries, Baton Rouge, Louisiana.

Martin, P.G., C.L. Butler, E. Scott, J.E. Lyles, M. Mariano, J. Ragus, P.
Mason, & L. Schoelerman. 1990. Soil Survey of Natchitoches Parish,
Louisiana. United States Department of Agriculture, Soil Conservation
Service. Baton Rouge, Louisiana.

Mohlenbrock, R.H. 1993. Black Branch Barrens, Texas. Natural History
(March):30-32.

Nixon, E.S. & J.R. Ward. 1981. Distribution of Schoenoltrion wrightiz (Lili-

aceae) and Bartonta terana (Gentianaceae). Sida 9:64-69.

Orzell, S.L. 1990. Texas Natural Heritage Program Inventory of National
Forests and National Grasslands in Texas. Unpublished report. Texas
Parks and Wildlife Department, Austin, Texas, and U.S. Forest Service,
Lufkin, Texas.

Perkins, B.E. 1981. Vegetation of sandstone outcrops of the Cumberland
plateau. M.S. thesis. University of Tennessee, Knoxville, Tennessee.
140 pp.

Smith, L.M. 1988. The natural communities of Louisiana. Louisiana Natural
Heritage Program, Baton Rouge, Louisiana. Unpublished report.

Smith, L.M. 1991. Louisiana longleaf: An endangered legacy. Louisiana
Conservationist (May/June):24-27.

480 PHY TODOGIA volume 75(6):463-480 December 1993

Smith, L.M., N.M. Gilmore, R.P. Martin, & G.D. Lester. 1989. Keiffer cal-
careous prairie/forest complex: A research report and preliminary man-
agement plan. Unpublished report. Louisiana Department of Wildlife
and Fisheries, Baton Rouge, Louisiana.

Williams, J.R. 1977. Food plants of seven selected monocot families for
Louisiana wildlife. M.S. thesis, Louisiana Tech University, Ruston, Louisi-
ana.

Phytologia (December 1993) 75(6):481.

ARENARIA GYPSOSTRATA B.L. TURNER, A NEW NAME FOR A.
HINTONIORUM B.L. TURNER, NOT A. HINTONIORUM B.L. TURNER

Billie L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT

A new name (Arenaria gypsostrata) is required to substitute for
A. hintoniorum which is a later homonym of a previous A. hintontorum.

KEY WORDS: Caryophyllaceae, Arenarza, México, gypsophile

Arenaria gypsostrata B.L. Turner, nom. nov. Based upon Arenaria hinto-
niorum B.L. Turner, Phytologia 75:406. 1993 [Feb 1994]. Not Arenarza
hintoniorum B.L. Turner, Phytologia 7259. 1992.

In my overzealous effort to eponymise the remarkable Hinton family, a lapse

permitted the nomenclatural mistake corrected here, for which my apologies
to the Hinton family and the broader systematic community.

481

Phytologia (December 1993) 75(6):482.

CORRECTIONS AND ADDITIONS

Volume 74, issue 3, page 178, line 4 of abstract, “Fabanae” should not have
a hyphen.

Volume 75, issue 2, inside front cover, D.M. Sutherland was omitted as an
author of the paper entitled “ Chromosome numbers for Dalea species (

FabaceaeFabaceae) from southwestern New Mexico and southeastern Ari-
zona.

Volume 75, issue 3, inside front cover, the paper by MacRoberts & Mac-
Roberts begins on page 247, not page 248.

Volume 75, issue 4, the running heads on all even numbered pages should
read: bo YT OL .OcGAA 74(5):xxx-xxx October 1993”
rather than “PHY TOLOGIA 74(4):xxx-xxx October
1993”.

Volume 75, issue 4, page 277, line 1 should read “Phytologia (October 1993)
75(4):277-280.” rather than “Phytologia (October 1993) 74(4):277-280.”

Volume 75, issue 4, page 277, line 1 should read “Phytologia (October 1993)
75(4):281-324.” rather than “Phytologia (October 1993) 74(4):281-324.”

Volume 75, issue 4, page 277, line 1 should read “Phytologia (October 1993)
75(4):325-329.” rather than “Phytologia (October 1993) 74(4):325-329.”

Volume 75, issue 4, page 277, line 1 should read “Phytologia (October 1993)
75(4):330-332.” rather than “Phytologia (October 1993) 74(4):330-332.”

Volume 75, issue 4, page 277, line 1 should read “Phytologia (October 1993)
75(4):333-335.” rather than “Phytologia (October 1993) 74(4):333-335.”

Volume 75, issue 4, page 277, line 1 should read “Phytologia (October 1993) |
75(4):336-338.” rather than “Phytologia (October 1993) 74(4):336-338.” |

Volume 75, issue 4, page 277, line 1 should read “Phytologia (October 1993) |
75(4):339-340.” rather than “Phytologia (October 1993) 74(4):339-340.”

482

Phytologia (December 1993) 75(6):483.

INDEX TO AUTHORS, VOLUME 75

Allen, C.M. 336

Baird, G.I. 74

Brouillet, L. 224

Cereno, J.C. 192
Chrelashvili, L.G. 124
Cowan, C.C. 281
Cuatrecasas, J. 235
Garcia, A., A. 243

Grant, J.R. 170
Klackenberg, J. 199
Laferriere, J.E. 399

Lemke, D.E. 330
MacRoberts, B.R. 247, 463
MacRoberts, M.R. 247, 463
Mayfield, M.H. 178
McInnis, N.C. 159
McIntosh, L. 224

Miller, H.A. 185

Montes, L. 192

Nelson, C. 190

Nesom, G.L. 1, 45, 55, 74, 94, 113

7;

118, 163, 218, 341, 347, 358,
366, 369, 377, 382, 385, 391,

452
Nuciari, M.C. 192
Ochoa, C.M. 422
Pittman, A.B. 159
Rayner, T.G.J. 100, 149
Roberts, R. 330
Simpson, B.B. 341
Smith, L.M. 159
Spellenberg, R. 166, 224
Suh, Y. 341
Sutherland, D.M. 166
Thomas, R.D. 336

483

Turner, B.L. 121, 134, 136, 140, 143,

147, 176, 204, 221, 231, 239,
259, 277, 281, 325, 333, 400,
402, 404, 406, 409, 411, 417,
432, 481

Ward, D.E. 166

Phytologia (December 1993) 75(6):484-512.

INDEX TO TAXA, VOLUME 75

New taxa described in this volume are indicated in bold face type.

Acacia 228
constricta 228
greggtt 228
Acamptopappus 20, 24, 40
Acanthaceae 466
Acer 251
rubrum 251
Aceraceae 251
Acourtia 404, 405
hintoniorum 404, 405
tomentosa 404, 405
Actipsis 4
Adenosma 282
Adiantaceae 382
Aeginetza 399
saccharicola 399
Aesculus 470
pavia 470
Agalinis 252, 468
fasciculata 468
plukenettu 468
skinneriana 468
Agaloma 178
Agavaceae 466
Agave 233, 234, 326, 374, 386
lechugtlla 233, 234, 326
Ageratina 147, 148, 402, 403
acevedoana 402, 403
subg. Ageratina 147, 148, 402,
403
cardiophylla 148
gentryana 403
subg. Neogreenella 147, 148
parryana 402, 403
perezw 147, 148

484

Ageratina (cont.)
viejoana 147
warnocki 403

Agrostis 161, 466
elliotiana 161, 466

Aletris 250, 466
aurea 250, 466

Allardtia 171

Allieae 146

Allioniaceae 451

Allioniella 449
orybaphotdes 449

Alhionta 432, 433, 435, 439, 441, 444,

446, 449
albida 435
carletonit 441
ciliata 441
coahuilensis 435
comata 439
corymbosa 449

var. terensts 449
decumbens 444
diffusa 444
ezaltata 441
gausapordes 444
gigantea 439
glabra 439
grayana 435
hirsuta 441
latifolia 433, 435
linearis 444
nyctaginea 446
oblongifolia 435
pseudaggregata 435
rotata 435

Index to taxa, volume 75 485

Allioniza (cont.)
terensis 449
vaseyt 444
Allium 333-335, 466
canadense 466
glandulosum 333, 334
hintoniorum 333
Plummerae 334
stoloniferum 333
Alnus 251
serrulata 251
Aloysia 386
Amaryllidaceae 146, 250, 466
Amellus 61
Ammocodon 242
Amphiachyris 20, 24, 40
Amphipappus 17, 20, 21, 24, 25
Amphirhapis 4
Anacardiaceae 251
Anactis 5
Andigena 427
Andreana 427
Andropogon 466, 470
tenarius 466
Anemone 161
caroliniana 161
Antsophyllum 182
vellertflorum 182
Anoplophytum 171
Anthaenantia 250
rufa 250
Apiaceae 251, 466
Aplactis 3
Aplopappus 69
sect. Diplostephioides 69
Apocynum 470
cannabinum 470
Apodocephala 62
Apostates 61, 71
Aquifoliaceae 251, 466
Archibaccharis 55, 57, 62, 63, 71
Arenaria 400, 401, 481
hintoniorum 400, 401, 481

Arenarza (cont.)
gypsostrata 481
lanuginosa 400, 401
Aristida 161, 228, 250, 466, 470
longespica 466
oligantha 466
purpurascens 250, 466
var. virgata 250
Aronia 252
arbutifolia 252
Artemisia 69, 71, 91, 92, 386
Asclepiadaceae 199, 203, 251, 466
Ascleptas 251, 253, 466
lanceolata 253
longifolia 251, 466
rubra 251
viridiflora 466
Aster 17-19, 22-24, 27-31, 36-
39, 43, 45-48, 50-54, 61, 94-
99, 113-115, 117, 163-165,
251, 345, 452-461, 467
acuminatus 458
alpinus 114
ageratozdes 52
albus 28
amygdalinus 455
andohahelensts 98
subg. Aster 46, 47
subsect. Aster 47
asterordes 50
baccharordes 52
bakeranus 95
baronit 98
bifolatus 51, 54
sect. Brotra 47, 459
subg. Brotia 45
brickellioides 52
ciltatus 345
collinsu 51
conyzotdes 48, 50
cornifolius 455
curtus 51, 53
dimorphophyllus 457

486

PHY TODOGIA

Aster (cont.)

sect. Doellingeria 454

subg. Doellingerta 452-454, 461

dumosus 251, 467
dumosus 467
ericotdes 18
gracilis 46
grisebachi 163, 164
forma angustissima 164
harveyanus 95
hashimotoi 457
hemisphericus 17
infirmus 455, 461
sect. Integrifolit 47
japonicus 456
kingtt 53
komonoensis 457
lateriflorus 18
leucanthemus 50
sect. Leucoma 48
linaritfolius 467
lintfolius 51
longipetiolatus 457
lutescens 29
madagascariensis 98
mandrarensis 98
marchandi 456
marginatus 164
marilandicus 50
miqueltanus 456
nemoralis 47, 458
oolentangtensis 467
oregonensis 51
paludosus 18, 467
subsp. hemisphericus 467
panduratus 458
patens 467
paternus 50
plantagintfolius 50
sect. Ptarmicoidet 27, 28
ptarmicoides 27, 28, 30, 31
pubentior 455
sect. Radulini 47

volume 75(6):484-512

Aster (cont.)

reticulatus 47, 453, 458

rigtdus 51

rugulosus 456

saboureaut 98

scaber 456, 457

sekimotot 457

senectonotdes 98

sericeus 467

sericocarpotdes 251, 455

sect. Sericocarpus 48

subg. Sericocarpus 48

sect. Serratifolti 48

sohayaktensis 454, 456

series Sohayaktenses 454

solidagineus 50

subsect. Spectabiles 46, 47

spectabilis 46

striatus 458

subg. Symphyotrichum 17

sect. Teretiachaentum 456

tortifolius 48, 51, 54

trichanthus 457

sect. Triplopappus 454, 461

umbellatus 454, 455, 461
var. pubens 455

vials 52

Asteraceae 1, 36, 37, 39-41, 43-45,

December 1993

53, 55, 64, 69-73, 74, 79, 81,
91-94, 99, 113, 116, 118, 121,
123, 134-136, 140, 143, 146,
147, 163, 165, 176, 192, 204,
216-218, 220, 221, 224, 251,
257, 325, 341, 344, 345-347,
355, 357, 358, 365, 366, 368,
402, 404, 452, 460, 461, 467,
469, 478

x Asterago 30
Astereae 1, 7, 18, 19, 22, 25, 37, 39,

40, 42-45, 52-55, 57-59, 61,
62, 64, 70-74, 90, 92-95, 97-
99, 113, 117, 118, 120, 163-
165, 218, 220, 341-347, 348,

Index to taxa, volume 75 487

Astereae (cont.) 357, 358, 361, 364-
368
Asterinae 61, 62, 366
Astereae 452, 460, 461
Asteromoea 456
japonica 456
Atripler 71, 92
Aureolaria 468
pectinata 468
Aylacophora 59, 66, 358, 361, 362
deserticola 358, 362
Azonopus 466
affinis 466
Azorella 366
Aztecaster 41, 55, 60, 63, 64-68,
83, 93, 99, 347, 357, 360,
363, 365, 367, 368
matudae 55, 65, 68
pyramidatus 55, 64, 65, 68

Baccharidastrum 62
Baccharidinae 22, 41, 55, 57, 62-64,
73, 93, 95, 97, 99, 348, 357,
365, 368
Baccharidopsis 62
Baccharis 55-58, 62-64, 68, 71, 72,
348, 350, 351, 353, 359, 362
acaulis 348
sect. Discolores 56, 57
sect. Glandulocarpae 56
lucida 350
matudae 56-58, 64, 68
phillipw 362
phylictformis 350
pteronioides 56
pyramidata 56-58, 64, 68
quadrangularis 351, 353
spartioides 359
Baptisia 467
leucophaea 467
Bartonia 251, 479
paniculata 251
terana 479

Bellidinae 366
Berchemia 468, 470
scandens 468, 470
Betulaceae 251
Bidens 100, 101, 104-107, 109, 111,
149-152, 155-158, 192-198,
251
acuticaults 151, 152
aristosa 251
aurea 192-198
borianiana 100, 106, 109, 111
camporum 100, 101, 106, 107
cochlearis 150, 155
diversa 149-152, 155
subsp. diversa 149, 151, 155
var. diversa 150
subsp. filiformis 149, 151, 156
var. megaglossa 150, 151, 155
var. gutlembana 150, 155
var. typtca 155
filiformis 149, 151, 156
gledhillii 100, 104-107, 109, 111
laevis 192, 193
lineartloba 150
pilosa 192, 193, 196
steppta 156
subalternans 192
Bigelovia 20, 47, 56-58, 68, 76, 84,
86-88
sect. Chrysothamnopsis 76
glareosa 86
graveolens 86
var. glabrata 86
var. hololeuca 86
var. latisquameus 86
howardii 88
var. attenuata 88
juncea 86
letosperma 86
mohavensis 87
paniculata 84
pyramidata 56-58, 68
turbinata 88

488 PHY TOELEOQGIA volume 75(6):484-512 December 1993

Bigelowra 161, 467 Callisia (cont.)
nuttalliz 161, 467 rosea 277
Bignoniaceae 467 Callitriche 161
Biophytum 190 nuttalliz 161
dendroides 191 Calopogon 250, 253, 254
zunigae 190 barbatus 253, 254
Biotia 45, 456, 457 tuberosus 250
discolor 457 Calymenia 444
japonica 456 decumbens 444
Blakiella 59, 367 Campanulaceae 251, 467
Blechnaceae 250 Campsis 467
Boltonia 251, 456 radicans 467
diffusa 251 Capraria 287, 321
japonica 456 durantifolia 287
Boraginaceae 226 humilis 321
Brachychaeta 1, 10, 39,44 oppositifolia 287 —
Brachystegia 157 Caprifoliaceae 251
Brassicaceae 227, 231 Carez 250, 336-338, 463, 466, 475,
Brickellia 140-142, 224, 225 477
amplezicaulis 224 caroliniana 466
aramberrana 140-142 flaccosperma 466
coulteri 225 glaucescens 250
grandiflora 140, 142 hyalina 336-338
Brintonia 1, 5, 39 meadit 463, 466, 475, 477
Bromeliaceae 170, 175 Carya 468
Bromus 228 Caryophyllaceae 159, 227, 400, 481
rubens 228 Castilleja 228
Bumelia 470 ezserta 228
lyctoides 470 subsp. erserta 228
Burmannia 250 Castillejinae 229
capitata 250 Catopsis 171
Burmanniaceae 250 Caulanthus 227
lastophyllus 227
Cacaka 251 Cecropta 427
ovata 251 Cedrela 427
Calceolaria 428, 429 Celmisza 61, 66, 70, 96-99
Callicarpa 468 Centaurtum 259-275
americana 468 arizonicum 259, 260, 262, 263,
Callisia 277-280 266, 267, 269-27', 273
sect. Cuthbertia 277, 279 arizonicum x tezense 263
graminea 277 beyrichit 259-261, 263, 265, 268,
hintoniorum 277-279 271; 2%

ornata 277 var. beyrichi 260

Index to taxa, volume 75 489

Centaurium beyrichit (cont.) Chamaesyce crepitata (cont.)
var. glanduliferum 260, 265 var. longa 179, 181
breviflorum 259, 261, 263, 274 cumbrae 181
calycosum 259, 260, 262-266, 270, fendlerz 179, 181
271, 273, 274 var. triligulata 179
var. artzonicum 262-264, 271 fruticulosa 179, 181
var. calycosum 260, 263-265, var. hirtella 179, 181
273 fruticosa 179
var. breviflorum 260, 263, 265 geyert 179, 181
var. nanum 260, 263-265, 271, var. wheeleriana 179, 181
273, 274 glyptosperma 181
glanduliferum 259, 261, 265 golondrina 181
maryannum 259, 260, 262, 265, hirta 181
268, 269-271, 273 humistrata 181
multicaule 259-261, 271-273 hyperictfolia 181
nudicaule 259-261, 271-273 hyssoptfolia 181
parviflorum 268, 272 indivisa 181
pulchellum 259-261, 272-274 jejuna 181
terense 259-261, 263-265, 272- johnstonii 179, 180
274 laredana 181
var. glanduliferum 260 lastocarpa 180
var. breviflorum 260 lata 181
Centella 251 maculata 182
astatica 251 micromera 182
Centunculus 161 missurica 182
minima 161 nutans 182
Chaenactidinae 143 parryt 182
Chaetopappa 61, 113, 114, 344 perennans 182
Chamaesyce 178-182, 276 prostrata 182
acuta 181 revoluta 182
albomarginata 181 scopulorum 182
ammanniordes 181 var. inornata 182
angusta 181 var. nuda 182
arizonica 181 serpens 182
astyla 181 serpyllifolia 182
berterzana 180, 181 serrula 182
caprtellata 181 settloba 182
carunculata 181 simulans 179, 182
chaetocalyz 179, 181 stictospora 182
var. triligulata 179, 181 var. sublaevis 182
cunerascens 181 therzaca 180, 182
cordifolia 181 var. spurca 180, 182

crepitata 179, 181 velleriflora 180, 182

490 PHY TOLOGIA volume 75(6):484-512 December 1993

Chamaesyce (cont.) Chrysothamnus nauseosus (cont.) 363

villifera 182
Chaptalia 251
tomentosa 251
Chasmanthium 466
sesstliflorum 466
Chiliophyllum 60, 66, 83, 95, 363
Chiliotrichopsis 66, 363
Chiliotrichum 19, 55, 59, 61, 63, 66,
95, 347, 363
Chloris 336, 337
subdolichostachya 336, 337
Chodaphyton 282, 291, 292
ertctfolium 291
Chorizanthe 228
brevicornu 228
Chrysoma 2, 3, 20-22, 39, 46, 84
nauseosa 84
Chrysopsis 22, 28, 113-117
alba 28
gossypina 114
villosa 115
Chrysothamnus 17, 20, 22, 24, 25,
35, 41, 55-58, 68, 69, 71, 72,
74-93, 363, 365
affinis 88
albidus 77-79, 82
sect. Asiris 74
asper 88
bolanderz 90, 91
sect. Chrysothamnus 75, 76
consimulis 85
depressus 80
eremobius 75, 76, 91
sect. Graminez 75, 76
gramineus 75, 76
greener 80
linifolius 79, 80
sect. Macronema 74
monocephalus 89
sect. Nauseosi 56, 58, 75, 76, 80-
83
nauseosus 74-81, 84-88, 90-93,

subsp. albicaulis 79, 87
subsp. arenarius 85
subsp. bernardinus 85
var. bernardinus 85
subsp. bigelovit 85
subsp. constmilis 85
subsp. ceruminosus 85, 86
subsp. graveolens 86
subsp. glareosa 86
subsp. hololeucus 77-79, 86
subsp. iridis 86
subsp. junceus 86
subsp. latisquameus 86
subsp. letospermus 87
subsp. mohavensts 78, 87
subsp. nanus 87
var. nanus 87
subsp. nauseosus 87
subsp. nitidus 87
subsp. pstlocarpus 87
var. pstlocarpus 87
subsp. salicifoltus 87
subsp. tezensts 88
subsp. turbinatus 88
subsp. uzntahenszs 90, 91
subsp. viscosus 90
subsp. washoensis 88
oreophilus 85
var. artus 85
paniculatus 74-76, 81, 84
parryt 56, 74-77, 79-81, 88-90
subsp. affinis 88
subsp. asper 88
subsp. attenuatus 88
subsp. bolanderz 89
var. bolanderz 90
subsp. howardiz 88
subsp. zmulus 89
subsp. lator 89
subsp. monocephalus 89
subsp. montanus 89
subsp. nevadensis 89

Index to taxa, volume 75 491

Chrysothamnus parryi (cont.)
subsp. parryt 89
subsp. salmonensis 89
subsp. vulcanicus 89
sect. Pulchell: 75, 76, 81
pulchellus 75, 82
sect. Punctati 75-77, 79, 81-83,
83
pyramidatus 56, 57, 68
salictfolius 87
spectosus 87
var. albicaulis 87
var. gnaphalodes 86
var. spectosus 87
teretifolius 74-76, 80, 81, 84
viscidtiflorus 75, 78, 80, 91, 92
vulcanicus 89
Chusquea 427, 429
Cirsium 467
carolinianum 467
Cistaceae 467
Clusiaceae 251, 467
Cladonia 161
Colubrina 377
greggu 377
Columbiadorta 17, 20, 26, 41, 92
Columnea 318
trifoliata 318
violacea 318
Commelina 336, 338
benghalensts 336
Commelinaceae 277, 280, 336, 338,
406
Compositae 36, 38-40, 42-44, 52-54,
69-72, 90, 92, 93, 98-100, 111,
116, 117, 120, 139, 146, 149,
158, 165, 197, 198, 220, 230,
345, 346, 355, 357, 364, 365,
367, 368, 428, 429, 458, 460,
461
Connellia 171
Conobea 287

verticillaris 287

Convolvulaceae 467
Conyza 50, 51, 62, 164
asteroides 50
btfoltatus 51
lintfolia 50, 51
Conyzinae 62
Cordium 282
Coreopsis 100, 158, 161, 225, 230,
251, 467
californica 225
var. newberryt 225
camporum 100
sect. Fuleptosyne 230
lanceolata 467
linifolia 251
sect. Pugtopappus 230
tinctoria 161
tripterts 251
sect. Tuckermannia 230
Cornaceae 467
Cornus 467
florida 467
Coronopus 227
didymus 227
Corethrogyne 22, 114
Crataegus 468, 470
marshallit 468
spathulata 468
Crinitarta 460
Crotalaria 467
sagittalzs 467
Croton 467
capitatus 467
Crotonopsis 467
elliptica 467
Cruciferae 234
Cryptantha 226
nevadensis 226
Ctenium 106, 250
aromaticum 250
newtonit 106
Cupressaceae 466
Cuthbertia 277, 280

492 PHY POLO GEA volume 75(6):484-512 December 1993

Cyanotis 106 Desmodium (cont.)
longiflora 106 molliculum 389
Cynoctonum 252 macrostachyum 389
sessilifolium 252 neomezicanum 386, 389
Cyperaceae 250, 336, 338, 466 paniculatum 387
procumbens 390
Dalea 166-168, 467, 470, 482 pstlophyllum 389
albiflora 166 retinens 389, 390
alopecuroides 167 roset 385, 386
brachystachys 167 subrosum 385-387, 390
candida 467 tortuosum 389, 391
filiformis 167 Diaphoranthema 171
grayt 166, 167 Dicanthelium 250, 466
lachnostachys 167 aciculare 466
leporina 167 acuminatum 250, 466
nana 167 dichotomum 250
var. carnescens 167 enstfolium 250
neomezicana 168 sphaerocarpon 466
var. neomezicana 168 Dicerandra 185, 186
ordiae 167 christmani 185
pogonathera 168 cornutissima 185
var. pogonathera 168 frutescens 185
polygonoides 166, 168 immaculata 185
purpurea 467 thinicola 185, 186
versicolor 168 Diodia 468
var. sessilis 168 teres 468
Damnamenia 66, 70, 99 Dioon 136
Darcya 323 Diplopappus 28, 29, 113-116, 454
mutisit 323 albus 28, 29
Dastorima 12 var. lutescens 29
Dasyhirion 386 : delphinifolius 114
Delphinium 468 dubtus 114
carolinianum 468 intermedius 114
Dendropogon 171 lanatus 114, 116
Desmodium 385-390 lutescens 29
aparines 387 sect. Triplopappus 454
caripense 387 villosus 114-116
glutinosum 388 Diplostephioides 60
grahami 388 Diplostephium 55, 59-61, 63, 64, 66,
hartwegianum 388 69, 94-98, 114, 452
var. amans 388 madagascariense 95, 98
lindhermeri 388 schultzi 64

lineatum 388 Dissotis 104

Index to taxa, volume 75 493

Dissottis (cont.)
fruticosa 104
Doellingerra 28, 45, 113, 452-462
amygdalina 455
sect. Cordifoltum 452-454, 456,
459
series Cordtfoltum 453, 454, 456,
459
dimorphophylla 457, 459
sect. Doellingeria 452-454, 459
series Doellingerta 455
humilis 455
infirma 453, 455
japonica 456, 459
komonoensis 457
longtpetiolata 453, 457, 459
marchandi 453, 456, 459
obovata 458
series Papposae 457, 459
ptarmicoides 28, 458
pubens 455
reticulata 458
rugulosa 456, 457, 459
scabra 453, 457
sekimotoi 457
sohayakiensis 456, 459
sericocarpotdes 455
trichocarpa 458
umbellata 452, 453, 455
subsp. pubens 455
var. pubens 455
var. umbellata 455
Dolichogyne 349-351, 353
glabra 351
leprdophylla 349, 353
rigida 350, 351
rupestris 350, 351
sect. Tola 349
Donia 341, 344
ciliata 344
Drosera 251, 467
brevifolia 251, 467
capillaris 251

Droseraceae 251, 467
Dubautira 158

Eastwoodia 20, 24
Egerza 331
densa 331
Elaeophorbia 178
Eleocharis 250
tuberculosa 250
Engleria 22, 61
Eragrostis 250, 466
elltotti 466
spectabilis 250, 466
Erianthus 250
giganteus 250
Ericaceae 251, 467
Ericameria 22, 24, 41, 55-60, 63, 64,
66, 71, 72, 74-92, 361, 363,
365
sect. Asirts 58, 74, 75, 79, 80
bloomer: 80
<x bolanderi 89
cooper 80
cuneata 79, 90
discoidea 76, 79, 80, 90
sect. Ericameria 58, 74, 75, 79-
82, 363
ertcotdes 75, 79
lineartfolia 75, 82
sect. Macronema 58, 74-76, 79-
81, 363
nana 75
nauseosa 78, 84-88, 90
var. arenaria 85
var. arta 85
var. bernardina 85
var. bigelovii 85
subsp. consimilis 85
subsp. nauseosa 85
var. ceruminosa 85
var. glabrata 85, 86
var. glareosa 85, 86
var. hololeuca 85, 86, 90

494 PHYELOLO GEA

Ericameria nauseosa (cont.)
var. iridis 85, 86
var. jJuncea 85, 86
var. latisquamea 85, 86
var. leiosperma 85, 86
var. mohavensis 85, 87
var. nana 85, 87
var. nauseosa 85, 87
var. nitida 85, 87
var. psilocarpa 85, 87
var. salicifolia 85, 87
var. speciosa 85, 87, 90
var. texensis 85, 88
var. turbinata 85, 88
var. washoensis 85, 88
pachylepis 80
paniculata 84
parryi 88-90
var. affinis 88
var. aspra 88
var. attenuata 88, 90
var. howardii 88
var. imula 89
var. latior 89
var. monocephala 89
var. montana 89
var. nevadensis 89
var. parryi 89
var. salmonensis 89
var. vulcanica 89
sect. Stenotopsis 74, 75, 79
suffruticosa 75
teretifolia 84
x uintahensis 90
x viscosa 90
Erigeron 22, 40, 97, 113, 114, 117-
120, 164, 165, 218-220, 251,
253, 344, 467
annuactis 120
annuus 114
delphintfolius 114, 118-120
griseus 218-220
inoptatus 119

volume 75(6):484-512

December 1993

Erigeron (cont.)
jenkinsw 118-120, 220
mayoensis 218-220
neomezicanus 120
oreophilus 120
pappochroma 97
sect. Polyactts 117, 119, 120, 218,:
220
strigosus 467
vernus 251, 253
wislizent 218-220
Erinus 282, 321
verticillatus 321
Eriocaulaceae 250, 257
Eriocaulon 250, 253, 254
decangulare 250
tezense 250, 253, 254
Eriogonum 228, 229, 254
capillare 228
maculatum 228
vernus 254
wright 228
Eryngium 251, 466
integrifolium 251
yuccifolum 466
Erythraea 262-264, 273, 274
arizonica 262
beyrichi 263
calycosa 262, 264
var. arizonica 262
var. nana 264
nudicaults 273
tezensis 274
trichantha 263
var. angustifolia 263
Eucephalus 28, 45, 457
albus 28
scaber 457
Eupatorieae 140, 147, 402
Eupatorium 251
leucolepts 251
rotundtfolium 251
Euphorbia 106, 178-180, 182, 183,

Index to taxa, volume 75 495

Euphorbia (cont.) 467
subsect. Acutae 183
chaetocalyz 179

var. triligulata 179
corollata 467
crepitata 179

var. longa 179
depauperata 106
fendleri 179

var. triligulata 179
fruticulosa 179

var. hirtella 179
geyert 179

var. wheelertana 179
johnstonu 179, 183
lastocarpa 180
polycarpa 179

var. simulans 179
stmulans 179
theriaca 180

var. spurca 180
velleriflora 180

Euphorbiaceae 138, 178-180, 183, 467

Euphorbieae 178, 179, 183

Euphorbiinae 178

Eurybra 98

Euthamia 2, 20, 23, 31-33, 39, 41-

43, 47, 72, 93, 346
graminifolia 1, 31-33

Evolvulus 467

sericeus 467

Fabaceae 166, 251, 385, 409, 467,
469
Fabanae 482
Fagaceae 467
Felicia 22, 61, 113, 116
Fimbristylis 466
puberula 466
Floscaldasia 360, 365-367
hypsophila 366
Flosmutisia 59, 360, 366
Fouquteria 386

Fuchsia 428
Fuirena 250
squarrosa 250

Gaillardiinae 143
Galactia 467
volubilis 467
Galatella 47, 51, 460
obtustfolia 51
platylepis 51
Gaura 468
Gelsemium 468
sempervirens 468
Gentanthus 199-203
bicoronatus 200, 202
crasstfolius 199
hastatus 201-203
laurtfolius 199-203
siamicus 202, 203
Genttana 274
pulchella 274
Genttanaceae 251, 259, 275, 467, 479
Geocarpon 159-162
minimum 159-162
Gibasis 406-408
consobrina 408
gypsophila 406, 407
hintoniorum 406, 407, 408
karwinskyana 407, 408
pellucida 406, 407
Gladiolus 106
pstttacinus 106
Gleditsia 470
triacanthos 470
Gnaphalium 467
purpureum 467
Gramineae 429
Grangeinae 22, 366, 367
Gratiola 252, 302, 304
hookerz 304
pilosa 252
tetragona 302, 304
Greenella 40

496 PHY TOL OGdA

Grindelia 22, 341-346
adenodonta 344
buphthalmordes 343
ciliata 344
lanceolata 344
microcephala 344, 346
nuda 344
papposa 341, 344, 345
squarrosa 342

Guizotia 158

Gundlachia 20, 46, 47

Gutterrezia 17-23, 25, 40, 41, 43, 45,

46, 47, 53, 71, 226
tezana 18
Gymnosperma 20, 23, 40

Habranthus 475
tubispathus 475
Haloragidaceae 251
Hamamelidaceae 467
Haplopappus 22, 34, 35, 39, 52, 56,
57, 60, 68, 71, 74, 76, 77, 81,
83, 91-94, 96, 97, 99, 113,
163, 164, 341-343, 345, 365
sect. Asirts 71, 92, 365
ctliatus 345
sect. Ericameria 76
linearifolius 93
sect. Macronema 71, 92, 365
marginatus 163, 164
parry? 34
phyllocephalus 345
sect. Prionopsts 341
pyramidatus 68
Hazardia 343
Hedeoma 161, 468
hispidum 161, 468
Hedyotis 161, 252, 468
australis 161
boscu 252
crassifolia 161, 468
nigricans 468
rosea 161

volume 75(6):484-512

December 1993

Heleastrum 17, 28
album 28
Helenieae 143, 176
Helenium 251
drummondiu 251
Heliantheae 121, 123, 134-136, 139,
146, 192, 198, 204, 216, 217,
230, 257, 342
Helianthus 251, 467, 470
angustifolius 251, 467
hirsutus 470
Herpestris 321
diffusa 321
Hesperodoria 24, 41, 83, 92
Hesperonta 432
Heterothalamus 55, 57, 62, 359
spartiotdes 359
Heterotheca 23, 113-117, 467
graminifolia 467
sect. Phyllotheca 115
villosa 114, 115
Hinterhubera 55, 58-63, 66, 70, 347,
358-361, 363, 365, 367
columbica 360
scoparia 360, 361
Hinterhuberinae 22, 41, 55, 58-64,
66, 72, 83, 93, 94, 96, 97,
99, 347, 348, 357, 363, 365-
368
Humiriaceae 235, 238
Humirtastrum 235-237
colombianum 236
cuspidatum 237
dentatum 237
ezcelsum 237
glaztovit 236, 237
mussunungense 235, 237
ptraparanense 237
spiritu-sanctt 237
villosum 237
Hydrilla 330-332
verticillata 330, 331
Hydrocharitaceae 330, 331

Index to taxa, volume 75

Hyparrhenza 106
chrysargyrea 106
diplandra 106

Hypericum 251, 467
brachyphyllum 251
cruz-andreae 251
gentzanoides 467
hypericoides 467
setosum 251

Hypochaeris 225
radicata 225

Hypoxidaceae 377

Hypozis 250, 377-381, 466
decumbens 377-381

var. decumbens 379, 380

var. dolichocarpa 377, 378-

381
var. major 380, 381
hirsuta 377, 378, 466
mezicana 381
pulchella 377-379, 381
rigida 250
Hyptis 251
alata 251
Hysterionica 115, 163-165
dianthifolta 163, 164
filiformis 163, 164
marginata 163, 164
pinifolia 163, 164
pulvinata 164
villosa 164

Ilex 251, 466
cortacea 251
decidua 466
vomitoria 466
Inula 28
alba 28
Tonactts 113-115
sect. Chrysopsis 115
Iridaceae 250, 466
Isocoma 7, 41, 52, 56, 342, 344
pluriflora 52

Iva 161
angustifolia 161

Jatmehintonia 146, 279
gypsophila 279
Jalapa 444
longiflora 444

Juglandaceae 468

Juglans 427

Juncaceae 250, 466

Juncus 250, 466
debilis 250
marginatus 250, 466
scirpotdes 250
trigonocarpus 250

Juniperus 228, 466, 470
monosperma 228
virginiana 466, 470

Kalimeris 453, 456, 460
sect. Cordifolium 456
longipetiolata 457
marchandii 456
miqueliana 456

Kotschya 104
ochreata 104

Krigia 161, 467
occidentalis 161
virginica 467

Labiatae 158, 414

Lachnocaulon 250, 254, 257
anceps 250
digynum 250, 254
Laestadia 360, 366, 367
Lagenifera 97

Lasallea 54, 165, 461

Lauraceae 252, 378

Lechea 467

tenuifolia 467
Leguminosae 168, 169, 390
Letoligo 6, 7

subg. Breviligula 6

Lamiaceae 185, 251, 411, 414, 468

498 PHY TOLOGHIA

Letoligo (cont.)
subg. Doria 7
subg. Lininque 7
Lendneria 282, 285, 321, 323
agerattfolta 285
humilis 321
verticillata 321
Lentibulariaceae 252, 468
Leocus 106
lyratus 106
Leptactis 12

Lepidophyllum 59, 63, 66, 70, 347-

351, 353, 355, 363
abietinum 350
cupresstforme 347
cupressinum 353
lucidum 350
meyentt 353
phyliciforme 350

var. resinosum 350
quadrangulare 351
rigidum 350
terettusculum 355
tola 353
sect. Tola 349

Leucospora 282, 295
multifida 295
Iiatris 251, 467
acidota 251
aspera 467
earler 467
pycnostachya 251
squarrosa 467
Liliaceae 250, 333, 335, 466, 479
Linaceae 252, 468
Linosyris 47, 84, 85, 88, 89, 460
bigelovi 85
bolander: 89
ceruminosus 85
howardi 88, 89

var. nevadensts 89
parryt 88
teretifolius 84

volume 75(6):484-512

December 1993

Linum 252, 468
medium 252, 468
Llerasta 59, 60, 63, 66, 70, 81
Lobelia 251, 467
appendiculata 467
reverchoni 251
Loganiaceae 252, 468
Liquidambar 467
styraciflua 467
Lotus 409, 410
hintoniorum 409
sect. Hosackia 409, 410
oroboides 410
Loudetia 106
kagerensis 106
Ludwigia 252
hirtella 252
Lupinus 427
Luzula 161
bulbosa 161
Lycopodiaceae 250
Lycopodium 250, 254, 283
alopecurordes 250
appressum 250
carolinianum 250
cernuum 254
Lyonta 253
ligustrina 253

Machaeranthera 22, 40, 71, 92, 114,
342, 343, 345, 346
sect. Psilactis 71, 92
Macronema 90
bolanderz 90
Madagaster 61, 66, 72, 94, 97, 98
andohahelensis 98
madagascariensis 98
mandrarensis 97, 98
saboureaui 98
senecionoides 98
Magnolia 248, 252
virginica 248, 252
Magnoliaceae 252

Index to taxa, volume 75 499

Manfreda 466
virginica 466
Marshallia 251
graminifolia 251
ssp. tenutfolia 251
Mathiasella 420
bupleuroides 420
Matourea 282, 309, 311
pratensis 309, 311
Medicago 467
lupulina 467
Melampodium 136-139, 225, 230
americanum 136, 138
lineartlobum 138
mayfieldii 136-138
strigosum 225
Melanthium 250
virginicum 250
Melastomataceae 252, 427, 428
Mezobromelia 171
Microglossa 55, 62, 95
Microlecane 158
Mildella 382-384
fallax 382, 384
intramarginalis 382-384
var. serratifola 382-384
var. intramarginalis 382, 383
leonardii 382-384
Mimosa 228, 386
beuncifera 228
Minutifolia 427
Mirabilis 432-451
aggregata 449
albida 432, 433, 435-439, 441,
444, 446
var. lata 435

austrotexana 432-434, 436, 438-

440
carletont 441
ciliata 441
coahuilensis 435
coccinea 449
collina 446

Mirabilts (cont.)
comata 432, 434, 436, 437, 439
decumbens 444
diffusa 444
dumetorum 433, 435, 436, 439
entricha 435, 436
ezaltata 439, 441, 444
gausapoides 444
gigantea 432, 434, 438-440
glabra 432, 434, 436, 439-441
glabriflora 449
glabrifolta 442, 449
grayana 435
hirsuta 432, 434, 441, 443, 444
jalapa 432, 433, 443, 444
subsp. lindhetmert 444
var. lindhetmert 444
lindhermert 444
linearis 432, 434, 436, 441, 444,
445
longiflora 432, 433, 444, 447
var. longzflora 444, 447
var. wrighttana 444, 447
section Mirabilis 451
muellert 435
multiflora 432, 433, 446, 448
var. multiflora 446
nesomu 445
nyctaginea 432, 434-436, 439, 441,
446, 450
var. albida 435
var. hirsuta 441
oblongifolia 435, 439
orybaphoides 432, 433, 449, 450
pauctflora 435
pseudaggregata 435
subgenus Quamoclidion 451
rotata 435
texensis 432, 434, 442, 449
Monadenium 178
Monocotyledonae 332
Monoptilon 61
Morgania 282

500 PHYTOLOGTA

Muhlenbergia 250, 466
capillaris 466
ezpansa 250

Mutisieae 404

Myrica 252, 468
cerifera 252, 468
heterophylla 252

Myricaceae 252, 468

Myrtophyllum 251
aquaticum 251

Nannoglottis 460
Nardophyllum 59, 61, 66, 70, 355,
358-364

armatum 359, 361, 362
bracteolatum 359, 362
bryoides 359, 362
chiliotrichotdes 359-362
deserticola 358, 362
genistordes 360, 362
lanatum 359, 362
obtusifolium 359, 362
paniculatum 362
patagonicum 358, 362
scoparium 358-361, 362

Neptunia 470
lutea 470

Nerisyrenia 231-234, 326, 329
baconiana 231, 232, 234, 326
lineartfolia 231, 233, 234

var. lineartfolia 231, 233, 234
var. mezicana 231, 233, 234

mexicana 234

Nostoc 161

Nothoscordum 161, 198, 466
bivalve 161, 466
inodorum 198

Nyctaginaceae 239, 242, 432, 451

Nyssa 252
sylvatica 252

Nyssaceae 252

Oenothera 161, 468

volume 75(6):484-512

December 1993

Oenothera (cont.)
linifolia 161, 468
Olearza 55, 59-61, 66, 70, 94-98
argophylla 96
dentata 96
pannosa 96
tomentosa 96
Oligactis 48
Oligoneuron 1, 2, 7, 18, 20, 23-34,
39, 47, 53, 72, 93, 458, 459,
461
album 1, 25, 27, 28-34, 458
x bernardii 30
houghtonii 1, 25, 27-29
x krotkovii 30
xlutescens 29
x maheuxii 30
nitidum 25, 28
ohtoense 27, 29, 30
sect. Oligoneuron 26
sect. Ptarmicoidei 27
ser. Ptarmicoidei 28
riddelliz 28-30
rigidum 1, 25, 27, 30, 31
var. glabratum 1, 27
var. humilis 1, 27
var. rigidum 27
ser. Xanthactis 28
Olivaea 342-344
Onagraceae 252, 468
Opuntia 386
Orchidaceae 250, 258, 466
Oreochrysum 1, 2, 18, 20, 24, 25, 26,
34, 35, 460
parry? 34, 35
Oreostemma 60, 72
Oritrophium 59-61, 66, 72, 99, 361,
367
orizabense 72, 99
Orobanchaceae 399
Orthocarpus 229
purpurascens 229
Osmunda 250

Index to taxa, volume 75 501

Osmunda (cont.)
cinnamomea 250
regalts 250
Osmundaceae 250
Oxalidaceae 190, 468
Ozalis 428, 468
stricta 468
Orybaphus 432, 435, 439, 441, 444,
446, 449
albidus 435
coahutlensis 435
comatus 439
eraltatus 441
giganteus 439
glaber 439
hirsutus 441
linearzs 444
multiflorus 446
nyctagtneus 435, 436, 446
var. latifolius 435, 436
var. oblongifolius 435
pauciflorus 435
pseudaggregata 435
wrighti 449
Ozypolis 251
filiformis 251
rigidior 251
Oyedaea 204
ovaltfolia 204

Pachystegia 66
Paleaepappus 59, 66, 358, 361, 362
patagonicus 358, 362
Panicum 106, 250, 466
anceps 466
congoense 106
rigtdulum 250
tenerum 250
virgatum 250
Papaver 227
rhoeas 227
Papaveraceae 227
Parastrephia59, 63, 66, 347-354, 356,

Parastrephia (cont.) 360, 363, 365

ertcotdes 349-351
leprdophylla 347, 353
lucida 347-353
phylictformis 347, 350, 351
quadrangularis 347-351, 353, 354
teretiuscula 347, 349, 354, 356
Paspalum 250, 466
notatum 466
plicatulum 250
setaceum 250
Pediculareae 229
Pedilanthus 178
Pellaea 384
intramarginalis 384
var. serratifolia 384
Pentachaeta 61
Pentzia 226, 229
incana 226
Perityle 176, 177
feddemae 176
glaucescens 176
sect. Laphamia 176, 177
sect. Perityle 177
Persea 252
borbonia 252
Perymentopstis 204
Perymenium 121-123, 204-217
bishopit 205-207, 209
buphthalmoides 217
celendianum 204, 206-208
colombianum 204, 205, 209,
211
ecuadoricum 204, 214, 215
featherstonez 206, 208-211, 213,
216
grande 205
hintoni 123
hintoniorum 121, 123, 217
huascaranum 204, 205, 210,
PA erik
huentitanum 121-123
jelskit 204-206, 209, 210,

502 PY T Oo, OG: ieA

Perymenium jelskwu (cont.) 212-215
klatta 216
lineare 205, 212, 215
matthewsw 204, 213
oazacanum 217
rosmarinifoltum 215
serratum 204, 213
tamaulipense 121, 123
tehuacanum 217
Petradoria 2, 19, 24, 25, 35, 39, 74,
76, 79, 83
discoidea 76
pumila 79
Phaelypea 282, 287
erecta 287
Phlox 468
ptlosa 468
Phytophthora 424-426, 428, 430
infestans 424-426, 428, 430
Phytarrhiza 171
Pinaceae 243, 250, 466
Pinguicula 252, 466
pumila 252
Pinus 219, 243-246, 250, 377, 466,
471
echinata 466, 471
herrare 219
maziminor 244
oocarpa 243, 244
var. microphylla 243, 244
palustris 250, 466, 471
praetermissa 243-246
pseudostrobus 244
taeda 466, 471
Pitcairnia 171
Pityopsis 114
Plagiocheilus 368
Plantaginaceae 468
Plantago 161, 468
aristata 468
elongata 161
virginica 468
Plasmopara 124, 130

volume 75(6):484-512

December 1993

Plasmopara (cont.)

viticola 124, 132
Platanthera 250, 253, 254, 256, 258,

466

blephariglottis 253, 254, 256

ctlarts 250, 254

cristata 253

integra 253, 254, 258

nivea 253, 466
Platanus 377

rzedowskit 377
Pleurophyllum 66, 70, 98
Pluchea 251, 253

foetida 251

rosea 253
Poa 161

annua 161
Poaceae 250, 336, 466, 469
Poarium 321

veronicoides 321
Podocarpus 136
Pogonia 250

ophioglossotdes 250
Poinsettia 138, 178
Polemoniaceae 468 |
Polygala 252, 468

cructata 252

mariana 252

nana 468

ramosa 252

verticillata 468
Poltomintha 413
Polyclados 349, 350, 353

abietinus 350

cupressinus 349, 353
Polygalaceae 252, 468
Polygonaceae 228, 229
Populus 377

tremuloides 377
Portea 171
Portulacaceae 468
Prionopsis 341-345

ctlrata 341, 343, 345

Index to taxa, volume 75 503

Prunella 468
vulgaris 468
Prunus 468, 470
Psathyrotes 143, 146
Psathyrotopsis 143-145
hintoniorum 143-145
purpusi 143, 145
Pseudocatopsis 171
Pstadia 55, 62, 95
Psiadiella 55, 62, 95
Pteris 382, 384
fallar 382, 384
intramarginalis 384
Pteronia 22, 55, 61, 66, 83, 96, 97,
99, 363
Ptiimnium 251
capillaceum 251
Puya 171
Pyrrhopappus 467
carolinianus 467

Quamoclidion 432, 446, 449
multiflorum 446
orybaphordes 449

Quercus 219, 228, 244, 377, 378, 467,

471
arizonica 219
coccolobifolia 219
falcata 467
magnolifolia 244
marilandica 467, 471
pumila 468
rysophylla 378
stellata 467
turbinella 228

Ractnaea 171, 175

Ranunculaceae 391, 468

Ranunculus 391-397
carolinianus 397
cymbalarza 391, 392
fascicularts 395

fasciculatus 391, 393, 395, 396

Ranunculus (cont.)
geordes 395
hispidus 392, 395-397
var. caricetorum 396
var. hispidus 396, 397
var. nitidus 396, 397
macranthus 395
peruvianus 391-393
petrolarts 391, 392, 394-396
var. arsene2 391, 392, 394, 395
var. petrolaris 391, 392, 394,
395
var. sterrae-orientalis 391, 392,
394, 395
var. trahens 391, 394-396
pilosus 395
pringler 395
septentrionalis 397
var. pterocarpus 397
sierrae-orientalis 391, 393, 394,
396
trahens 391, 396
Rattbida 470
pinnata 470
Remya 61, 73
Rhamnaceae 369, 376, 468
Rhezia 252, 253
alifanus 253
lutea 252
mariana 252
petrolata 252
Rhododendron 253
canescens 253
oblongifolium 253
Rhus 374
Rhynchospora 250, 254, 256, 466
chalarocephala 250, 254
elliotti 250
globularis 250, 466
glomerata 250
gracilenta 250
inerzpansa 250, 466
latifolia 250

504 PHY TOLLtO GA

Rhynchospora (cont.)
macra 250, 254, 256
oligantha 250
plumosa 250
rariflora 250
Rhytachne 106
rottboellioides 106
Rigtopappus 61
Rochonia 55, 61, 64, 66, 94-98
senecionoides 98
Rosaceae 252, 468
Rubiaceae 252, 468
Rubus 429, 468
Rudbeckta 253, 254, 256, 257
scabrifolia 253, 254, 256
Ruellia 466
humilis 466

Sabatia 251, 253, 254, 256, 467
campestris 467
gentianordes 251
macrophylla 251, 253, 254, 256

Sageretia 369-376
elegans 369, 371-375
mexicana 369-373, 375
minutiflora 369, 372-375
thea 375
wrightw 369, 371-375

Salvia 386, 429, 468, 470
azurea 470
lyrata 468

Sarcanthemum 55, 62

Sarracenia 247, 252
alata 252

Sarraceniaceae 252

Satureja 411-414
sect. Gardoquta 411, 413, 414
hintoniorum 411-413
maderensis 411, 413
mezicana 413
seleriana 413

Schinus 429
molle 429

volume 75(6):484-512

December 1993

Schizachyrium 250, 466, 470
scopartum 250, 466
tenerum 250, 466

Schoenolrion 250, 253, 463, 466, 470,

474-476, 478, 479

croceum 250, 253

wrighti 463, 466, 474-476, 478,
479

Scirpus 161
kotlolepis 161

Scleria 250, 253, 466
ciliata 466
georgiana 253
oligantha 466
reticularis 250

Scrophularia 287
subhastata 287

Scrophulariaceae 228, 229, 252, 281,

324, 468

Schrankia 467, 470
microphylla 467, 470

Scutellaria 251, 468
integrifolia 251, 468
parvula 468

Secamone 199, 203

Secamoneae 199

Selaginella 463, 466, 475, 476
arentcola 463, 466, 475

var. riddelli 463, 466, 475

Selaginellaceae 466

Selinocarpus 239-242
lanceolatus 239-242

var. lanceolatus 241

var. megaphyllus 239, 242
maloneanus 239-242
megaphyllus 239-242

Senecio 221-223, 325-329

aureus 221
var. Balsamitae 221
var. borealis 221
claryae 325-328
douglasit 329
fendleri 221

Index to taxa, volume 75

Senecto (cont.)

flaccidus 325-329

var. douglas 327

var. flaccidus 327

var. monoensis 327, 328
multilobatus 221
neomezicanus 221, 222

var. metcalfer 222

var. mutabilis 222

var. neomezicanus 222

var. toumeyt 222
pattersoni 326
pinacatensis 327
plattensis 221
powellii 325, 326-328

Suffruticosa species-complex 325

Suffruticosi 328
thurbert 221, 222
tridenticulatus 222

Senecioneae 146
Sericocarpus 19, 20, 22-24, 41, 45-

52, 54
acutisquamosus 52
asteroides 46, 48-50

f. albopapposus 50
f. roseus 50
befoliatus 51, 52
var. acuttsquamosus 52
var. collinsu 51
californicus 51
collunstw 51
conyzotdes 46, 50
linifolius 48-50
oregonensts 45, 49-51
subsp. caltfornicus 51
var. californicus 45, 50, 51
var. oregonensis 50, 51
rigidus 48, 49, 51
var. californicus 51
var. laevicaulis 51
stper 52
solidagineus 48, 51
tomentellus 52

Sericocarpus (cont.)
tortifolius 48, 49, 51
var. collinsi 51
woodhousei 52
Setarra 466
geniculata 466
Silphium 467
lacintatum 467
Sisyrinchium 250, 466
atlanticum 250
sagttiferum 466
Smailaz 250, 466
laurtfolia 250
Solanaceae 422, 431
Solanum 422-431
acaule 423
subsp. albicans 423
var. albicans 423
series Acaulia 423
albicans 423
albornozt 423
andreanum 422
baezense 422
burtoni 427
calacalinum 423
caquetanum 424
chilliasense 429
chomatophilum 425
f. angustifolium 425
colombianum 424, 426
var. meridionale 424
f. quindiuense 424
var. trianae 424
f. ztpaquiranum 424

505

series Conicibaccata 422, 423

correll: 427
cyanophyllum 429
dolichocarpum 424
filamentum 424

flahaulti 425
huancabambense 428, 429
series Juglandifolra 422
juglandifolium 422

506 PEE YET OCL 36 p is volume 75(6):484-512 December 1993

Solanum (cont.) Solidago (cont.)
minuttfolrolum 428 bicolor 5, 6, 34
ochranthum 422 boottz 10
series Olmostana 426 subg. Brachyactis 8
olmosianum 426 ser. Brachychaetae 10
paucyugum 425 brachyphylla 10
sect. Petota 422, 431 buckleyt 7
series Prurana 429 caesta 6, 31, 32
subsect. Potatoe 422 calcicola 4, 14, 15
regulartfolium 428, 429 californica 9, 43
serratoris 422 canadensis 7, 8, 29, 31, 32, 34,
solisw 430 36, 37, 40, 41
suffrutescens 429 celtidifolra 9
series Tuberosa 422, 427 chapmanz 10, 11, 25
tundalomense 425, 426 chlorolepis 4
tuquerrense 430 sect. Chrysastrum 5

Solenstemnon 106 subg. Chrysastrum 5
monostachyus 106 chrysolepis 12

Solidagininae 1, 2, 10, 18-24, 26, 35, compacta 15

45-48, 59, 61, 83, 348, 459, confinis 11, 12

460 curtisiz 6

Solidago 1-20, 22-32, 34-47, 53, 72, sect. Corymbosae 26

(9282-093 s 2b lem2osec4o2. cutleri 4, 13

453, 459-461, 467 deamii 4
aestivalis 9 decemflora 8
alba 28 decumbens 4
subg. Albigula 5 delicatula 9
subsect. Albigula 5, 6 discoidea 5, 6
albopilosa 6, 36 drummondiu 9
alpestris 14 durangensis 8, 41
altiplanites 8 edisoniana 9
altisstma 8 ellrottu 9
amplezicaulis 29 elongata 8, 31
arguta 9, 10, 31, 40 erecta 4-6
ser. Argutae 10 sect. Erectae 4
sp.-group Argutae 9, 10 ser. Erectae 4
subsect. Argutae 9 ertcamertoides 11, 12
arizonica 8 fistulosa 9
aspera 9 flacctdtfolia 6
auriculata 10 flavovirens 12
austrina 12 flezicaulis 6, 31, 32, 36, 37
bellidtfolia 4 gattingerz 11

x bernardi 30 gigantea 8, 31, 36, 40

Index to taxa, volume 75

Solidago (cont.)

gullmani 4

glomerata 4

sect. Glomeruliflorae 6

subsect. Glomeruliflorae 6, 31

gluttnosa 4, 14, 15, 41, 42
var. monticola 14

gracillima 12

graminifolia 31

gutradonis 12

gypsophila 8

harperz 7

harrisiz 10

hintonitorum 7

hispida 5, 6, 32, 34

houghtoni 29, 40

humilis 42

ser. Integrifoliae 5

jejunifolia 7

juliae 8

juncea 11, 12, 32, 34, 40

ser. Junceae 11, 12

sp.-group Junceae 11

subsect. Junceae 11-13, 18

klughi 13

x krotkovi 30

lanctfolia 6

latisstmifolia 9

leavenworthiu 8

lepida 8, 19, 31, 32

ludoviciana 9, 10

x lutescens 29

macrophylla 5, 6, 14-16, 35

macvaughit 8

x maheuzi 30

sect. Maritimae 12

subsect. Maritimae 12, 13, 19

mezicana 12

microglossa 2, 8

microphylla 9

murabilis 9

missourtensis 11, 12, 19, 36, 42

mollis 8, 19

Solidago (cont.)

muellerz 11

multiradzata 4, 13, 15, 34
var. arctica 13, 15

ser. Multiradiatae 4

nana 4, 8

neglecta 12

sp.-group Nemorales 8

subsect. Nemorales 8, 9

nemoralts 8, 9, 11, 43

neomezicana 4

nittda 28, 467

odora 10, 11, 25

sp.-group Odorae 10

subsect. Odorae 10

ohioensis 27

subg. Oligoneuron 26

oreophila 4

orientalis 7

ouachitensis 6

paniculata 8

sect. Paniculatae 8

parryi 34

patula 10, 251

ser. Pauctradtatae 4

perlonga 12

petiolaris 7, 41

ptinetorum 11

subg. Plezactila 6

plumosa 4

porterz 5

pringler 11, 12

ptarmicoides 28, 31, 32, 36

puberula 5

pulchra 12

purshiz 12

racemosa 4

radula 9

randu 4

riddelliz 28

rigida 26, 27, 31, 39
var. glabrata 27
var. humilis 27

508

PRY OL O' GTA

Solidago (cont.)

roanensis 5

rugosa 9, 36, 253

rupestris 8, 36

salicina 10

sciaphila 4

sect. Secundiflorae 9

sempervirens 12, 13

ser. Serratae 4

shortiu 8

stmplez 4, 13, 15, 16, 41, 42

subsp. stmplez 15
subsp. rand 15

simulans 12

sect. Solidago 1-3, 5, 16, 17, 19,
25, 26, 35

subsect. Solidago 4-6, 18, 25, 31,
34

sparsiflora 9, 43

spathulata 4, 13, 15, 42

speciosa 7

ser. Spectabiles 11, 12, 13

spectabilis 12, 19

sphacelata 9, 10

spithamaea 4

squarrosa 4-6

sect. Squarrosae 5

subg. Stenactila 12

stricta 12, 13

strigosa 10

tarda 10

sect. Thyrstflorae 4, 7

ser. Thyrsiflorae 7

subsect. Thyrsiflorae 7, 18, 25,
35

tortifola 11

subg. Triactts 11

ser. Trinerves 8

sect. Triplinerviae 8

subsect. Triplinerviae 8

uliginosa 12, 37

ulmtfolia 9

subsect. Unicostatae 12

volume 75(6):484-512

Solidago (cont.)

sect. Unilaterales 1, 2, 7, 8, 19,

24-26, 31
ser. Unilaterales 7
uniligulata 12
velutina 8, 41
ser. Venosae 9
subsect. Venosae 9
verna 10
victorini 4
virgata 12
sect. Virgatae 12

virgaurea 3-6, 13-17, 31, 32, 39,

43
var. 3 13
var. y 13
var. alpestris 13, 14
var. alpina 13
subsp. aszatica 16
var. calcicola 14
var. Cambrica 13
var. ertcetum 13
subsp. gigantea 16
subsp. letocarpa 16
subsp. minuta 16
subsp. virgaurea 16
sect. Virgaurea 3, 4
subg. Virgaurea 3
wrightii 7
yadkinensis 10
x Solidaster 1, 26, 30-32, 37, 44
x hybridus 37
x luteus 30, 37
Sopubia 106
mannit 106
Spergularia 161
echinosperma 161
Sphagnum 248
Spiranthes 250, 466
lacera 466
longilabris 250
praecoz 466
vernalis 250

December 1993

Index to taxa, volume 75 509

Sporobolus 466

junceus +466

Stetractinia 216

klatti 216

Stellaria 227, 229

nitens 227

Stemodia 281-324

ageratifolra 285
angulata 281, 283-286
subsp. ageratzfolia 285
arenaria 321
arizonica 288
berteroana 287
bissez 288
chilensts 290
chodat 316
cructflora 318
damaziana 294, 302, 312
duranttfolia 281, 284, 285, 287-
291, 296, 314
var. angusttfolia 287
B angustifolia 288
var. chilensis 281, 285, 288-
290
var. duranttfolia 281, 287, 289
ehrenbergiana 287
erecta 287, 288
ericifolia 281, 283, 291-293
subsp. ertcifolia 292
subsp. vera 291, 292
foltosa 293, 311
fruticulosa 305, 307
gratioltfolta 308
harley: 281, 283, 292, 293
hasslerrana 281, 283, 293, 294
humilis 321
hyptordes 281, 284, 291, 295-297,
299, 304, 313, 314
var. aurtculata 295
var. platensts 295
var. stricta 313
jorullensis 285
subsp. reptans 285

Stemodia (cont.)

lanceolata 281, 284, 296, 298-
300, 304, 314
forma angustzfolza 298
var. angustifolia 298
forma latifolia 298
var. latifolia 298
forma laziflora 298
latefolia 281, 301
linearifolia 308
var. acutifolia 308
lobata 281, 283, 301-303
lobeltordes 281, 284, 296, 299,
302-304, 309, 314
macrotricha 321
maritima 281, 282, 284, 305, 306
var. rigtda 305
microphylla 281, 284, 303, 307,
319
mutisi 323
orbiculata 296
palustris 281, 284, 308, 310, 314
forma salicifolia 308
var. simpler 308
parviflora 321
pulcomayensis 295, 296
pwurensis 305
pratensis 281, 283, 309-311
scopartoides 298, 299
stellata 281, 283, 310, 312
stricta 281, 284, 291, 299, 304,
a13, s14, oui
subsp. glabriuscula 313
forma minor 313
var. multidentata 313
var. paucidentata 313
subhastata 287
suffruticosa 281, 283, 311, 315-
BIE
forma dentata 315, 316
var. villosa 315, 316
surinamensis 323
tetragona 302, 304

510 PAPY'IT OD OiG A

Stemodia (cont.)
trifoliata 281, 283, 318, 320
verontcotdes 281, 284, 294, 319,
320
verticillaris 287
verticillata 281, 283, 321, 322
Stemodtacra 282, 284, 287, 290, 291,
295, 298, 305, 308, 311, 313,
315, 318, 321
angulata 284
berteroana 287
chilensts 290
duranttfolia 287
errcifolia 291
folosa 311
gratiolifelia 308
hyptoides 295
lanceolata 298
lineartfolia 308
maritima 305
palustris 308
stricta 313
subhastata 287
suffruticosa 315
trifoliata 318
verticillata 321
Stenotus 20, 24, 83
Stephanodoria 342
Steyerbromelia 171
Stylisma 253
aquatica 253
Stylosanthes 467
biflora 467
Styraz 136
Synadenium 178
Synchytrium 426
endobtoticum 426
Syzygium 106
guineense 106

Talinum 161, 463, 468, 470, 475,
476
parviflorum 161, 463, 468, 475

volume 75(6):484-512 December 1993

Tephrosia 251, 467
onobrychoides 251
virginiana 467
Thestum 106
tenuisstmum 106
Thurovia 18, 20, 40, 47
Tillaea 161
aquatica 161
Tillandsia 170-175, 276
andreettae 170, 171
arpocalyz 171
barclayana 172
boeghii 170, 172
castaneo-bulbosa 172
ceretcola 172
curvispica 170, 172
cylindrica 170, 174
drewii 170, 172
harmstana 172
hitchcocktana 172
incurva 172
koideae 170, 174
limonensis 170, 173
olmosana 170, 173
var. pachamamae 170, 173
patula 173
pereziana 173
petraea 173
penduliscapa 170, 173
peruviana 170, 173
porphyrocraspeda 170, 174
subg. Pseudalcantarea 170, 171
rauhi 170, 174
robusta 174
sagasteguzt 170, 173
strobelii 170, 174
tequendamae 174
subg. Tillandsia 170, 171
tillandsioides 170, 174
werneriana 170, 174
yaconorensis 170, 174
Tillandsioideae 170, 175
Tithymalus 178

Index to taxa, volume 75 511

Tonestus 17, 20, 24, 35, 47, 53, 346 Verbesina 134, 135
Townsendia 344 aramberrana 134, 135
Tozicodendron 251 hintontorum 134, 135
verniz 251 zaragozana 134, 135
Tozocarpus 199 Verena 282, 294, 295
Tracyina 61 hassleriana 294

Tradescantia 161, 279, 407

occidentalis 161
Tragia 467
urtictfolia 467
Transaequatorialia 427
Tridens 250
ambiguus 250
Triodants 467
perfoliata 467
Tristachya 106
fulva 106
Tuberosa 427
Tubuliflorae 71, 357

Unamia 27, 28, 30
alba 28
lutescens 30
ptarmicoides 28

Unanvea 315
dentata 315
febrifuga 315

Ungnadia 378

Utricularia 252
cornuta 252
juncea 252
subulata 252

Vaccinium 251, 467
arboreum 467
corymbosum 251, 467

Valeria 282, 318, 319
trifohata 318

Vanclevea 24, 83

Vaviloviana 427

Verbena 468
haler 468

Verbenaceae 468

Vernonia 106, 350, 467
jaegerz 106
phylictformis 350

var. restnosa 350
tezrana 467

Vernontopsis 55, 62, 81

Viburnum 251, 470
dentatum 470
nudum 251

Viola 252, 253, 468
lanceolata 253
pedata 468
primulifolia 252

Violaceae 252, 468

Vriesea 170-174, 276
andreettae 171
arpocalyz 172
barclayana 172
boeght 172
castaneo-bulbosa 172
ceretcola 172
curvisptca 172
cylindrica 170, 174
drewtt 172
harmsiana 172
hitchcockiana 172
incurva 173
kotdeae 170, 174
limonensis 173
olmosana 173

- var. pachamamae 173

patula 173
penduliscapa 173
pereziana 173
petraea 173

rauhiu 170
sagasteguz 170, 173

512 PHY TOLOGIA volume 75(6):484-512

Vriesea (cont.)
rauhi 174
robusta 174
strobeli 174
tequendamae 174
tillandsioides 174

Vulpia 466
octoflora 466

Wedelia 213

jelskit 213
Westoniella 59, 360, 363, 366-368
Woodwardia 250

virginica 250

Xanthisma 342
Xanthocephalum 22, 40, 342-344, 346
gymnospermoides 343, 344
Xylorhiza 343
Xylothamia 20, 41, 72, 93, 346
Xyridaceae 251
Xyris 251, 254, 256
ambigua 251
baldwiniana 251
caroliniana 251
difformis 251
var. curttssit 251
drummondit 251, 254, 256
loutstanica 251
scabrifolta 251, 254, 256
torta 251

Yucca 326, 374, 386

Zigadenus 250, 254, 256
densus 250, 254, 256

December 1993

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