PHYTOLOGIA

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

Vol. 78 Apnl 1995 No. 4

CONTENTS

DEBRECZY, Z. & I. RACZ, New species and varieties of conifers from

Mexico 217"

WIPFF, J.K. & S.D. JONES, Nomenclalural combination in Poaceae 244 -

ANDERSON, L.C., E.L. BRIDGES, & S.L. ORZELL, New data on

distribution and morphology for the rare Hasleola robertiorum (Asteraceac)..

246 '

SHAW, R.B. & P.P. DOUGLAS, Carex inversa R. Br. (Cypcraccae), new to

Hawaii and the United States , 249 -^

EGGER, M., New records and notes concerning Caslilleja spirantfioides

(Scrophulariaceae) 256 ^

RAO, B.H. & K.V. KUMAR, Lithocysts as taxonomic markers of the species

of Cordia L. (Boraginaceae) 260 ""

HAYS, J., A floristic suney of Falls Hollow sandstone glades, Pulaski County,

Missouri 264 ''

HE, PING & HONG LI, Studies on the isozyme \anation pattern among

individuals and populations of the endangered species Deulzia imdliradiata

(Hydrangeaceae) on Mt. Jinfo of Nanchuan 277

HE, PING, A new variety of Deulzia (Hydrangeaceae) in China 282

TURNER, B.L., A new species of Potentilla (Rosaceae) from Ccrro Quiexobra,

Oaxaca 285

LANDRY, P., A revised synopsis of the pines 6: Supplement to the subgenera,

and concl usion 287

MACROBERTS, M.H. &. B.R. MACROBERTS, Noteworthy \ascular plant

collections on the Kisatchie National Forest, Louisiana 291

Books received 314

Back issues available 315

Phytologia Memoirs available 316

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Phytologia (Apnl 1995) 78(4) 217-243

NEW SPECIES AND VARIETIES OF CONIFERS FROM MEXICO

Dr. Zsolt Debreczy & Dr. Istvdn R^z

International Dendrological Research Institute, Inc. (I. D.R.I.) (5 North Meadow

Road. Medfield. Massachusetts 02052 U.S.A.)

Botanical Department of the Hunganan Natural History Museum (H-1476 Budapest,

P.O. Box 222, HUNGARY)

ABSTRACT

Four new species and six new vaneties are described here; three species
and two vaneties of Abies, one species and three vaneties of Pinus, and one
new vanety of Pseudolsuga. Within the genus Abies only one Mexican
species was known with consistently more than two resin canals, and five
having cones with enclosed bracts. The authors here report two additional
(new) species, one from the state of Hidalgo and one from Oaxaca, with more
than two resin canals and, including these, altogether three spjecies with
enclosed bracts. A new vanety of Abies guaiemalensLs Rehd. was found in
Guerrero. Abies lowiana {Gord.) A. Murr., often unreasonably neglected as
a separate species and included in A. concolor (Gord. & Glend.) Lind. ex
Hild. as its vanety, is herein reported to be represented by a new vanety in the
state of Chihuahua. A distinct species of Pinus, discovered in Sonora, with
five needles in a fascicle, stout branches, and strong scales such as in P.
michoacana Mart., but with cones clearly placing this species in
"Pseudostrobus" (like P. estevezii [Mart.] Perry or P. nubicola Perry). This
new pine may play a key role in understanding the process of divergence of
"Montezumae" and "Pseudostrobus" pines. A vanety of the new species was
also found by the authors in the state of Sinaloa. The southernmost localities
of the genus Pseudolsuga so far were known from the states of "Hidalgo and
Puebla. The authors report on the species, as a new vanety, further south, in
Oaxaca A new variety of Pinus lawsonii Roezl ex Gord. and one of P.
oaxacana (Mart.) Mirov is also reported from the latter area

KEY WORDS: New taxa, Coniferae, Mexico, Abies, Pinus, Pseudolsuga,
Hidalgo, Guerrero, Oaxaca, Sonora, Sinaloa

NOTE: the species and vaneties descnbed here have been mentioned, along
with a concise descnption and photographs, in the No. 6 issue of the
NewsBnef of the International Dendrological Research Institute, Inc.
(I.D.R.I.), Sep. 1994. The taxa ''Abies A-E" are those listed in this article
under numbers 1-5, ''Pinus A-D" are those under 6-9, and "Pseudolsuga A"
is under 10 in the list below (discussion). For information contact I.D.R.I.,
as given above.

217

218 PHYTOLOGIA volume 78(4): 2 17-243 Apnl 1995

INTRODUCTION

While documenting the Gymnosperms and their plant communities m Mexico for
the Dendrological Atlas project we encountered several conifers which we consider
new taxa.

1: Abies hidalgensis spec. nov. (Hidalgo); 2: Abies neodurangensis spec,
nov. (Durango); 3: Abies zapotekensis spec. nov. (Oaxaca); 4: Abies
guatemalensis Rehd. var. longibracteata var. nov. (Guerrero); 5: Abies
lowiana A. Murr. var. viridula var. nov. (Chihuahua); 6: Pinus yecorensis
spec. nov. (Sonora-Sinaloa); 7: Pinus yecorensis Debreczy & Ricz var.
sinaloensis var. nov. (Sinaloa); 8: Pinus oaxacana (Mart.) Mirov var.
diversiformis var. nov. (Oaxaca); 9: Pinus lawsonii Roezl var. gracilis var.
nov. (Oaxaca); 10: Pseudotsuga menziesii (Mirb.) Franco var. oaxacana var.
nov. (Oaxaca).

For a long time (since 1932), only Abies hickelii Rous & Gaussen, a species of
southern Mexico, was known to have more than two (typically 8-12) resin canals in its
leaves (A. oaxacana Martfnez is treated here as a synonymous with A. hickelii.). A
new species was found in the montane "cloud forest" region of Hidalgo {Abies
hidalgensis Debreczy, Ricz, & Gufzar, see below) with (typically 4-5) up to 7 median
resin canals in the leaves, and soon after, surprisingly, another species with 4 resin
canals was located in the similarly mild and humid habitat of a hidden valley in the
state of Oaxaca (A. zapotekensis Debreczy, Ricz, & Ramfrez, see below). The
taxonomic value of the number and position of the resin canals is well discussed in the
literature (Orr 1937; Liu 1971; e/c). Gaussen (1971) thinks that marginal position of
the resin canals is a younger character than median, supported by the observation that
leaves on sterile shoots usually have marginal resin canals, while fertile ones of the
same specimen are typically median (Ferrd 1941). The phylogeneUc importance of the
resin canal number is little understood. Except for A. hickelii in Mexico, only two
species: A.Jirma Sieb. & Zucc. of Japan and A. bracteata (D. Don) A. Poit. of Pacific
North America were known so far occasionally to have more than two, namely 4,
resin canals. We suspect that this is either an ancient, "pnmitive" character or the
consequence of rapid changes in climatic circumstances during the history of the
species. In the ancient group of Gymnosperms, Podocarpus in a broad sense, there
are several species with more than two resin canals-e.^., up to 10 in sect.
Eupodocarpus such as in P. rostratus Laurent-and their position is typically median.
Further study for the better understanding of the systematic position of these newly
discovered relict endemic species, should involve the invesugauon of questions
relating to the number of the resin canals.

There were only a few fir species known in Mexico having enclosed bracts:
excluding the southernmost types oi Abies concolor (Gord. & Glend.) Lindl. ex Hild.
known from northern Mexico together with Abies magnifica A. Murr. in Baja
California, only Abies guatemalensis Rehd. [not including A. g. var. tacanensLs
(Lund.) Mart, and the variety described here (var. longibracteata Debreczy & Racz)],
Abies durangensis Mart., A. d. var. coahuilensis (l.M. Johnst.) Mart, and Abies
vejarii Mart. var. tnexicana (Mart.) Liu. The first species was descnbed by Alfred
Rehder in 1939, the others by Maximino Martfnez in 1942. More recently (1992,
Biol. Sergio Zaimudo Ruiz of the Instituto de Ecol6gia, C.R. Del Bajio, Patzcuaro,
Michoac^ reported (pers. comm.) a species from the state of Queretaro with two resin

Debreczv & Racz: New conilers Ircim Mexitx) 219

canals and short, enclosed bracts. Our recent exploration extended this list by three
more species having this character Abies hidiilgensis, A. neodurangensis Debreczy,
Rdcz, & Sala7iir, and A. zapotekensis (see below),

Abies guatetnalensLs, a species with enclosed bracts has an extended population in
Guerrero w iih large cones and exserled bracts. So far only a southern form with Ime
needles, hairy branchlets, and smaller, narrow cones were known to have bracts
exceeding the cone scales (var. tacanensis) from the high region of the Volcdn Tacana,
Chiapas. The variety, seems to be much more extended in Mexico than the species
itself, is descnbed here as A. guaiemaletisis \ar. longibracteata.

Abies lowiana (Gord.) A. Murr., a species not consistently recognized by
botanists and mistakenly, often included in A. concolor {A. c. var. lowiana [Gord.]
Lemm.), is reported here from "south of the Rjo Grande", north of the town of
Madera, Chihuahua. With its A. grcmdis (Dougl. ex D. Don) Lindl. -like pectinate
arrangement of the leaves on shade branches and young trees, a character which never
appears in A. concolor, A. lowiana is more related to the previous species than to A.
concolor in a strict sense. It is important to distinguish this species from the latter
taxon, fairly frequent in the northwestern region of Mexico. Found here with a
surpnsingly lustrous green foliage (always pruinose gray in the north), it is considered
a new vanety of A. lowiana.

Whether a pine species belongs to the "Montezumae" or "Pseudostrobus" group in
classification is often difficult to establish. Pinus yecorensis Debreczy & Ricz has
characters of both sections or subsections: the strong, thick branches with large,
prominent pnmary scales suggest that it belongs to "Montezumae", while the cone is
that of "Pseudostrobus". At this time, only the discovery of the species and a
comparative morphology of this and some of the reb^ed species will be discussed.
Chemical composition analysis and a more thorough siudy are still to come.

Pinus oaxacana (Mart.) Mirov is the most common species in Oaxaca between an
altitudinal range of 1900-2500 m. With its long pendent, thin/fine shiny green foliage
and the fairly large cones close to the apex of the strong branches for most of the year,
this species is the most typical element of the "encinar y pinar" belt of the region.
During our expeditions the variability of the species was thoroughly studied and a
large number of samples collected. These proved a wide range of vanability of the
species, particularly in their cone morphology. A form with yellow (not purplish red)
developing male strobiles is also worth mentioning here. While making this study,
distant small populations with P. montezumae Lamb. -type elongated, just barely
asymmetnc cones of much less protuberant scales draw our special attention,
particularly when noticing that this type consistently had brown branchlets instead of
pruinose, typical for the species. If P. montezumae were present in the area, one
would easily suspect that this pine is a hybnd between P. oaxacana and that species.
Here we describe it as a vanety of the Oaxacan pine, knowing that this solution may
just open this taxonomic problem for further study and consideration.

Pinus lawsonii Roezl is the most common species of the relatively low altitude
( 1400-2500 m) coniferous forests of southern Mexico. Over one hundred collections
were made to observe its natural vanability. Several types, which include trees with
different growth pattern, color, or other characters of the needles, along with a w ide
range of the vanability in the cones, were observed'. We felt that all of them belong to
the natural vanability of the species and were not worth distinguishing with formal
taxonomic epithets e.xcept for a type with smaller cones and conspicuously thin
needles. This type, growing consistently in the lower limits of the distnbution of the
species, is treated here as P. lawsonii \ds. gracilis Debreczy & Ricz see below .

220 PHYTOLOGIA volume 78(4): 2 17-243 Apnl 1995

The distnbution of the genus Pseudolsuga in North Amenca has been known as
far south as the stales of Hidalgo and Puebla only. Recent discovery of the genus in

the slate of Oaxaca has extended the known distnbution of the genus to 16° N lat., and
the new location with those being discovered since the first report, represent the
southernmost location of the genus worldwide. The Me.Xican Sierras, including the
area of the Sistema Norte del Oaxaca, are known to be within an earlier wider range of
the distnbution of northern corufers including the genus Picea, known recently only as
far south as the norlhem states of Mexico. The (by now) isolated distnbution of the
northern taxa here, is practically just the continuation of their distnbution southward
{e.g., in Picea, that of Picea pungens Engelm. and Picea engelmanii Parry) with great
morphological similanty to those of the northern ones, differing from them only in
some characters and typically showing a wide range of infraspecific vanability anyway
(e.g., the length and thickness of the needles, number of the stomatophorus lines,
shape of the cone scales, etc.). The Oaxacan Douglas fir is also just a southern
representative of the variable northern Pseudotsuga menziesii (Mirb.) Franco
segregated to isolated populations during its repeated migrations and survived in a
wide range of different habitats representing a wide ecological spectrum. These
ecotypes were separated on species rank {Pseudotsuga flahaultii Rous in Chihuahua-
Durango-Zacatecas in the west, Coahuila-Nuevo Le6n in the east, Pseudotsuga
guinieri Flous in Chihuahua-Durango, Pseudolsuga macrolepis Rous in Chihuahua-
Durango and Coahuila-Nuevo Lebn-Hidalgo-Puebla, Pseudolsuga rehderi Rous in
Chihuahua-Durango and Nuevo \jo6n). They ait practically distnbuted throughout the
range of the genus with geographic ranges that are close to or even overlapping one
another. This "over-splitting" of the major species resulted in such "ndiculous" cases,
that (for example) from just in the vicinity of Galeana, Nuevo Le6n, three species of
Douglas-fir were reported: Pseudotsuga flahaultii and Pseudotsuga macrolepis from
Cerro Potosf, NW of Galeana, and Pseudotsuga rehderi from the Canon de los
Charcos, SE of the town. The extreme variability of the main species (Pacific and
continental. Rocky Mountain forms) and the companson of cultivated trees with wild
specimens show the difficulty and weak basis of evaluating them as different species,
or even as varieties. Dropping the idea of placing the Oaxacan Douglas fir under a
"questionable" and ill-defined taxon such as, most likely, Pseudotsuga macrolepis,
and considering the distant isolation of our taxon, we propose that it be separated on
the level of variety within the main species. We are, however, aware of the fact that
cultivation and further investigation which should include cold hardiness tests, may
prove that this is not even a variety. Further data on the distnbution of the species in
Oaxaca (see description #10), reported soon after the first publication of our discovery
in a local newspaper, indicate how widely the species was distnbuted in this region in
earlier climatic periods.

1. Abies hidalgensis Debreczy, Rdcz, & Gui'zar^, spec. nov. Figure 1: cl-c3;
Plate 1:1; "Abies A" in NewsBnef No. 6 of 1. D.R.I. (Sep. 1994). TYPUS:
MEXICO, eastern Hidalgo: 4-5 km north of Metepec, 2300 m, June 12, 1994,
Debreczy. Rdcz. Bird et al. M0323 (HOLOTYPUS: BP; Isotypi: A,CHAP,E,
MEXU,NA).

Arbor ad 18 m alta, truncosimplici; habitus columnan-conicus; cortex
rasilis, in arbonbusjuvenilibus pallide cinereus, postea in laminas irregulares
magnas ruptus, in arbonbus vetustis valde crassus et suberosus, pars intenor

Debreczy & Rdcz: New conifers from Mexico

corticis sanguinea; rami fusci, vel cinereo-fusci, usque ad 5-6 annos pilosi;
ramuli \alidi, rasiles, pili.s 0.2-0.3 mm longis, dcnsis, llaNo-brunncolis
diaphanis ccxiperti; gemmae (partim foliis brevibus, incurvalis, termmalibus
occultae) rolundalo-descclae, lenuitcr resinosac, squamis usque ad 15
tnangulanbus vel saltem apicibus squamorum tectae; folia pecUnatim
dislnbuta, ( l-)3-5<-6) cm longa, abrupte abbreviata (usque ad 1 cm longa) m
ramulis exlemis leviicr deorsum ct/vel versus apicem ramulorum mclinata,
aciebus parallelis, apicibus roiundata el breviier (0.2-0.4 mm), sed conspicue
emargmata vel subtililer bifida, in positione umbrosa mollia fle.xibilia vel in
acumme arbons aliquanlum quasi succulenta; pagina dorsalis eorum
atrovindis vel glauco-vindis, lineis stomatum usque ad 14, in ramulis ad
lucem expositis deorsum medium folii attingentibus, sed in positjone umbrosa
plerumque 6 lineis brevibus, ad apicem folii litteram "V" formantibus; pagina
ventralis argenieo-cinerea cum margine angusto (0.2 mm), vindi et costa
media 0.2-0.4 mm lata, saepe cum lineis stomatum infractis; limitibus
lateralibus bilaleraliter singulis argenteis, lineis stomatum typice 14-15 (in
positione umbrosa in foliis angustis 5-6) instructis; lineis stomatum cum
stomatibus 9 pro 1 mm; stomata 0.06 mm longa, elongato-ovaia, intense
pruinosa; folia ramulorum infemorum in secuone transversali plano
tnangulana, sed ramulorum fertilium tnangulana cum canalibus resiniferis
medianis usque ad 7(!); strobili feminei juveniles 2.5-3.0 cm longi,
bracteae eorum irutio adpressae postea reflexae (4-5 mm), rotundato-
tnangulares, irregulariter serrato-incisae, pallide vindes; strobili masculini
12 mm longi, pedunculo 3 mm longo, squamis pilosis ciiica 100 (20 x 5),
1.8 mm latis; strobili maturi 6.5-8.0 cm longi. 3.5-4.0 cm lati, pedunculo
0.4-1.0 cm longo, cylindracei, apice rotundati, leviter umbilicati; squamae
130-170(25-34 x 5) pro strobilis, flabellatae, virides, glauco-vmdes, dense
velutinosae, pili 0.2 mm longi, albidi, adpressi; bracteae 0.5-0.8 cm longae,
sessiles, leviter spathulatae, cum ala acute inciso-serrata, rotundata, sine apice
protrudenti; axis anguste-conicus; semina anguste-tnangulana, resino-
vesiculana cum ala cinereo-straminea.

DESCRIPTION: tree with straight, typically single trunk, eolumnar-conical
crown, first ascending, later descending branches, shiny- (or in sunny and dryer
places) dull grayish-green, pruinose foliage; bark: smooth, light gray in young trees,
later ruptured to irregular large plates, very thick and suberous on old trees with blood-
red inner bark; branches: light brown, later grayish brown, hair>' (up to 5-6 years);
branchlets: rather strong, smooth, slightly prominent below the leaves on strong
shoots, densely pubescent with 0.2-0.3 mm long yellowish brown, semi-clear hairs;
buds: (partially hidden by short, incurving terminal leaves) flat, covered by up to 15
triangular scales or their tips, thinly resinous; leaves: pecunately arranged,
"distichous", abruptly shortened (to 1 cm) on the upper (dorsal) side of the branchlets,
slightly downward and/or forward curving, (l-)3-5(-6) cm, parallel sided, rounded
and shortly (0.2-0.4 mm), but markedly, emarginate or finely bifid, soft flexible
(shade) or rather fleshy (upper crown), shiny dark- or dull grayish-green with up to 14
stomatophorus lines on the upper (dorsal) side reaching below the middle of their
length on stronger exposed stems, usuzilly 6 short lines form a (0.6-1.0 mm long) "V"
below the apex on shade leaves; lower (ventral) side is silvery gray with narrow green
margins (0.2 mm) and midnb (0.2-0.4 mm, often with broken lines of stomata), two
broad silvery stnpes typically with !4-15 (in narrower shade leaves 5-6)
stomatophorus lines with 0.06 mm long, elongated ovate, intensively pruinose.

222 PHYTOLOGIA volume 78(4):2 17-243 Apnl 1995

densely set (9/mm) stomata; leaves in cross section flat tnangular on lower, tnangular
on fertile branchlets wilh up to 7(!) median resin canab, strobiles: female- 2.5-3.0
cm long with short, first appressed later reflexed (4-5 mm) rounded-tnangular,
irregularly serrate-incised pale green bracts; male- strobile 12 mm long on 3 mm
peduncle, consists of about 100 (20 x 5) 1.8 mm wide, hairy ferule scales; cones:
6.5-8.0 X 3.5-4.0 cm, short (0.4-1.0 cm) stemmed (peduncled), cylindncal with
about 130-170 (25-34 x 5) cone scales; rounded, slightly impressed (umbilicate) at
apex, pruinose green at maturity; cone scales: flabellale (wide cyathiform), green,
grayish green, densely velutinous with short (0.2 mm), white, appressed hairs; bracts
(0.5-0.8 cm) short, enclosed (sessile), slightly spathulate with sharply incised-serrate,
rounded wings and without a protruding tip; axis narrow corneal; seeds: narrowly
triangular with resin blisters and narrow, light yellowish gray wings (not fully
developed at the time of the descnption).

NAMED: after the central Mexican state of HidaJgo, where the species was found.
The name also recalls Miquel Hidalgo y Costilla (1762-181 1), a prominent personality
of the post-Columbian Mexican independence movement, after whom the state itself
was named.

DISCUSSION: this rare and distinct tree with 5-7 resin canals in the needles and
green cones with short, enclosed bracts was located in the state of Hidalgo in an area
where (in the wider region) only Abies religiosa (H. B. K.) Schltdl. & Cham, was
known. The latter is completely different with its strongly curved, pointed (not bifid)
leaves with two resin canals (not up to 7) and 2-3 times larger cones with long
exserted, recurved bracts of prominent tip. Abies guatemalensis, a southern sp)ecies
with (typically) "bractless" cones not known as far north as Hidalgo, has twice as
many leaves on an equal length of (shade) branchlet with straight needles curving to
almost a right angle from the stem (not curved, directed forward), with two resin
canals (not up to 7). In A. guatemalensis the cones typically pruinose blue (not
pruinose green changing to light brown), have wide cyathiform "wing shaped" (not
flabellate) cone scales with elongated (not short) basal section and slightly enclosed or
finely exserted (not short, sessile) bracts with finely protruding (not absent) tip. Other
species such as A. vejarii Mart of northeastern Mexico (with short, curved, succulent
needles with two resin canals, close to or exserted bracts) are too distinct even to be
compared with the new species.

DISCOVERY: the species was found in a deep canyon running eastward, 3 miles
above the small village of Metepec. The first tree was seen on Apsnl 3, 1994 on the
upper nm of the canyon just where the road makes a sharp turn downward toward
Poza Rica and Tampico at an elevation of 2350 m. The tree, old and struggling with
only a few upper branches alive, had no cones or cone axils, but the distinct foliage
and bark immediately suggested that it does not belong to any species known. In this
extremely cloudy area we had no opportumty to continue the exploration at that time.
Stenle branches were transferred to the Division de Ciencias Forestales, Universidad
Aut6noma Chapingo (DCF/UACH) to preserve the (possibly sole) tree under i38672.

We returned to the cirea on June 12. This time we saw more trees deep in the
canyon along with old cypress trees (Cupressus lusitanica Mill.). In spite of an early
start, we had only a few minutes before the clouds filtered up and hid the valley.
Realizing the difficulty of reaching the trees from our position and having no evidence
of cones, we went to the section of the canyon where the lonely tree was found a few
months earlier. We had good fortune for a few seconds and located a tree on a nearly
vertical slope, 20 m above the right side of the nver. The tree has had many cones
w hich were not npe yet but seemed to be fully developed in size. Soon we found it to
be identical to our #38672, but it was much younger with different bark and less

Debreczy & Racz: New conifers from Mexico 223

glaucous needles {DRB ^40323; type). Conifers associated with this fir are Pinus
palula Schltdl. & Cham., Pinus apulcensis Lindl. {P. pseudosirohus Lindl. var.
apulcensis [Lindl] Shaw), and Pinus leocole Schltdl, & Cham, in iLs upper
distnbuUon, and (a few) P. palula in the evergreen cloud forest in the canyon. At the
site of the specimen we found 36 accompanying species among w hich the dominaUng
were Quercus laurina H. & B., Ternslroemia pringlei Rose, and Cestrum fascicuUilum
(Schltdl.) Miers. Ipurpureum {L\nd\.) Standi, with Alnus firmifolia Femald, Buddleia
cordata H. B. K., Plelea trifoUaia L., Sambucus mexicana Presl., and Syngonium
podophyllum auct.

PRESERVATION: Declaring the entire canyon as conservation land was
proposed. Further study and conservation was planned in collaboration with Dr.
Enrique Guizar Nolazco, DCF/UACH.

PARTICIPANTS IN HELD WORK: Dr. Gyongyvdr Bir6, Dr. Zsolt Debreczy,
Dr. Istvdn Rdcz, (I. DR. I. and Hung. Nat. Hist. Museum, Budapest, Hungary), Eva
Kert6sz (Botanical Collection, Munkicsy Mihdly Miizeum, B^k^scsaba, Hungary),
Vince Zsigmond (University of Horticulture, Budapest, Hungary), Yinghao Zhao
(Botanical Garden of the Chinese Academy of Sciences, Beijing, China).

2. Abies neodurangensis Debreczy, Ricz, & Salazar*, spec. nov. Figure 1: al, a2
Plate 1:2; "Abies B" in NewsBnef 6 of I. D.R.I. (Sep. 1994). TYPUS:
MEXICO, southwestern Durango: near Los Bancos (Ejido la Victoria Paraje, San
Antonio, Ejido El Brillante Paraje, Santa Barbara, Arrovo del Infiemo), 2500 m.
May 13, 1994, Debreczy. Rdcz. Bird el al. »39936 (HOLOTYPUS: BP; Isotvpi:
A,CHAP.E.MEXU,NA).

Arbor ad 35-40 m alta, trunco simplici; habitus initio laxe conicus,
postea densior, vertice rotundata; cortex rasilis, in arbonbus juvenilibus
pallide cinereus, postea in arbonbus vetustis in laminas irregulares magnas
ruptus, pars intenor corticis sanguinea; rami ferruginei, usque ad 5-6 annos
sporadice pilosi, pilis subtiliter pellucidis; ramuli pallide vindes ad cinereo-
brunnei, dense vel in positione umbrosa parce pilosi, pilis pellucidis 0.10-
0.15 mm longis; gemmae in ramulis vegetativis typice subglobosae, 2-3 mm
longae, non-resinosae, gemmae in ramulis terminaJibus planae (6-7 x 3 mm)
cum 12-18 squamis tnangulanbus, 3 mm longis, nitidis, brunneis costa
prominenli et margine eroso instruclis; folia dichotoma, late distanlia, leviter
ascendentia, in latere supenon ramulorum prorsus inclinata, aciebus
parallelis, apicibus rotundata et leviter (0.1 mm) emarginata; flexibilia, in
pagina dorsali vindia, lineis stomatum usque ad 7 paene apicem attingentibus
et lineis aliquot prope basin foliorum praeter costam impressam; in pagina
ventrali argenteo-alba, cum margine angusto vindi (0.1 mm) et costa (0.25
mm lata), et limitibus lateralibus bilateraliter singulis argenieis, lineis (6-)9(-
10) stomatum instructa; stomata 0.03 mm, ovata; folia in ramulis fertilibus
saepe asymmetnce acuta, aliquantum succulenta cum limite pruinoso 0.2 mm
lato supeme lineis stomatum usque 8; costa crassa usque 0.7 mm lata, praeter
costam lineis 6-8 stomatum angustis 0.35 mm laUs; folia ramulorum
infemorum in sectione transversali eiliplico-aviculana, obovata, folia
ramulorum fertilium in secuone transversali obovata (ellipUca), canalibus
resinifens marginalibus 2, latero-ventraiibus; strobili breves, columnares,
6.5-8.5 cm longi. 3.0-3.6 cm lati, initio vmdes, postea vmdo-brunnei,
umbone prominenti; strobili velutini, pilis 0.10-0.13 mm longis. pellucidis.

224 PHYTOLOGIA volume 78(4):2 17-243 Apni 1995

erectis; squamae 150-200 (30-40 x 5), flabellatae; bracteae sessiles, 0.5-
0.8 cm longae, ala angulan, protrudenti, apice 3-4 mm longo; axis anguste-
conicus; semina 0.8-1.0 x 0.8 cm, ala 1.0-1.4 cm longa, obiusato-
thangulan instructa.

DESCRIPTION: tree up to 35-40 m tall with single trunk, loose conical, later
dense, round-topped crown with first ascending, later descendmg branches; bark:
smooth light gray first, later ruptured, in old trees very thick, suberous with blood-red
inner bark; branches: reddish brown, sporadically hairy with Tine clear hairs up to
5-6 years; buds: vegetative buds typically subglobose, 2-3 mm long, terminal one
flat (6-7 X 3 mm) on strong fertile shoots with 12-18 triangular, 3 mm long, wide,
shiny brown scales with prominent midnb and eroded margin, not resinous;
branchlets: light green to grayish brown, densely or sparsely (shade) hairy with
0.10-0.15 mm long clear hairs; leaves: on stem, "dichotomous", widely spreading,
slightly ascending, forward curving on the upper side, 3-5 cm long, 1.5-2.2 mm
wide, parallel sided, rounded and slightly (0. 1 mm) emarginate at apex, flexible, shiny
dark green above with up to 7 stomatophorus lines near the apex and a few lines down
near the base of the leaves along the impressed midrib, the ventral side is silvery white
with narrow green margins (0.1 mm) and midnb (0.25 mm) and two silvery stnpes
with (6-)9(-10) stomatophorus lines, stomata 0.03 mm, ovate, leaves on fertile shoots
often asymmetrically (on one side) acute, more succulent with wide (0.2 mm) pruinose
stripe above, with up to 8 stomatophorus lines, midrib strong, up to 0.7 mm wide, the
stomatophorus stripes are narrower (0.35 mm) with 6-8 lines of stomata. In cross
section the leaves are ellipto-aviculate to obovate (elliptical) on lower, obovale on
fertile branchlets with 2 marginal latero-ventral resin canals; cones: short, columnar
(6.5-8.5 X 3.0-3.6 cm) with prominent umbo, consist of 150-200 (30-40 x 5)
conescales, bright light green first, greemsh brown later, velutinous with 0.10-0.13
mm long upright clear hairs, conescales are flabellate with short (0.5-0.8 cm),
enclosed (sessile), bract with protruding angular wings exceeding the 3-4 mm long
apex; axis narrow conical; seeds: 0.8-1.0 x 0.8 cm with 1.0-1.4 cm long, rounded-
triangular wing; fully developed but not mature when studied.

NAME: after the state where the species was found with neo- (Lai. prefix = new-)
as antenor is to distinguish it from Abies durangensis, the well known and widely
distributed (although not common) species of the region.

DISCUSSION: this fir with ''Abies grandis (Dougl. ex D. Don) Lindl.-like"
appearance is characteristically with two resin canals in its needles, the early
appearance of the strobiles, and green cones with enclosed, sessile bracts. It was
found in a canyon on the Pacific side of the sierra close to the Durango-Sinaloa border.
Although enclosed bracts and two resin canals exist in the leaves of both, it can readily
be distinguished from A. durangensis by its densely hairy, dark purplish brown (not
subglabrous-glabrous yellow or pruinose) branchlets, glabrous (not thickly resinous),
straight (not curved), green (not gray pruinose), rounded-emarginate (not fwinted)
leaves with median (not marginal) resin canals, cones with very short, enclosed
(sessile) bracts with prominent, laminal wings forming a "V" exceeding the shortly
protruding tip (not subsessile), nearly reaching the "exposure line" (the inner limit of
the exposed part, "scale head" of the conescale) and tnangular-acute at end. It has
cones almost fully developed in size by the time A. durangensis "blooms". It is very
distinct from and not even comparable to A. concolor or A. lowiana.

DISCOVERY: On May 13, 1994, while heading from Capilla Textla, Sinaloa to
El Salto, Durango, just 3 miles before reaching the plateau, that is on the warmer,
western side of the sierra (identified later as Ejido la Victona Paraje, San Antonio and

Debreczy & R^cz: New conifers from Mexico 225

Ejido El Bnllante Paraje, Santa Barbara. Arroyo del Infiemo), suddenly a fir with fully
developed, although not mature, green cones came into sight. Traveling to dtxument
the cone-mllorescences (strobiles) of Abies durangerisis, we were surprised to see a
species in cone. Having the samples in hand, we realized that it was a aimpletely
different species, at first sight most similar to A. grandis, a species of the Pacific
North. At an altitude of 2500 m, the trees are growing in the margin and upper part of
deep canyons and associated with the dominating Hydrangea seernannii Riley and
twelve other species such as Alnus firmifolia, Cornus discijlora DC, Garrya laurifolia
Hartw., Litsea glaucescens H. B. K., and Quercus magnoliifolia Nee.

CONSERVATION: We found only a few mature trees. Although the difficult site
seems to secure the species, the preservation of the habitat(s) is immediately
necessary. Young plants are present, but not many. We reported to Dir. Tec. Felipe
Norberto Cona Quinone and Biol. Rosalva Miranda Salazar at Unidad Conservaci6n y
Desarollo (UCODEFO) No-6, El Salto, and proposed at least a few hectares of
conservation land in the canyon. The protection seems to be secured and further
studies are ongoing.

PARTICIPANTS IN HELD WORK: Dr. Gyongyv^r Bir6, Dr. Zsolt Debreczy.
Dr. Istvin Rdcz, (I. D.R.I, and Hung. Nat. Hist. Museum, Budapest, Hungary). Eva
Kert^sz (Botanical Collection, Munkicsy Mih^y Miizeum. B^k^scsaba. Hungary),
Vince Zsigmond (University of Horticulture, Budapest, Hungary).

3. Abies zapotekensis Debreczy, Rdcz, & Ramirez'*, spec. nov. Figure 1: d; Plate
I:3;"A/?/e.sC"inNewsBnefNo. 6of I.D.R.I. (Sep. 1994). TYPUS: MEXICO.
Oaxaca; Sierra de Juirez. near Porlillo, 2700 m, June 30, 1994, Debreczy. Rdcz,
Bird, etal. #40675a (HOLOTYPUS: BP; Isotypi: A,CHAP,E,MEXU,NA).

Arbor ad 20 m alta, trunco simplici, habitus laxe-conicus; cortex
rasilis, arboris juvenilis pallide cinereus, postea in laminas irregulares ruptus;
rami castaneo-brunnei. usque ad 5-6 annos sporadice pilosi; ramuli vindes-
flavovirides, tenuiter pubescentes, pilis 0.10-0.13 mm longis. leviter
infuscato-subpellucidis. basin versus leviter contractis; gemmae (foliis
brevibus, incurvatis, tenninalibus partim occultae) rotundato-desectae.
squamis usque ad 15 tnangulanbus tectae; exteme non-resinosae; folia
subdisticha. ascendentia, ad apicem leviter inclinata, litteram "V" patentem
formantia. in ramulis latens dorsalis aliquantum breviora et conspicue reflexa.
3-5 cm longa. 2 mm lata, aciebus parallelis, apice rotundata et breviter (0. 14-
0.20 mm) emarginata; flexibilia. aliquantum conacea nitide atrovmdia; folia
cum/sine lineis stomatum usque ad 5. 2-3 mm longis. prope apicem folii. vel
in ramis crassis lineis 2-3 deorsum infra medium folii; pagina ventralis
argenteo-cinereus cum margine angusto (0.15-0.20 mm) vindi et costa media
0.2-0.4 mm lata, saepe cum lineis stomatum infractis vel a stomatibus omnino
tecta, limitibus bilateraliter singulis argenteis. lineis stomatum typice 14-18 (in
positione umbrosa in foliis angustis 10-12) instructis; stomata 12-13 pro 1
mm. 0.03-0.04 mm longa, anguste elongata; folia in sectione transversali
plana, aviculana usque epsilon-formia cum canalibus resinifens medianis 2-
4{!); strobili feminei juveniles 3-4 cm longi, bracteis brevibus, iniUo
adpressis postea reflexis (4-5 mm), rotundatis, irregulanter serratis. pallide
viridibus, sine costa protrudenti marginem excendenU; strobili masculini
1.2 cm longi, squamis pilosis circa 100 (20 x 5) 1.8 mm latis; strobili
maturi 8-9 cm longi, 3.6-4.2 cm lati, cyiindracei, rotundati, vmdes, glauco-

226 PHYTOLOGIA volume 78<4): 2 17-243 Apnl 1995

virides. pedunculus 0.4-0.8 cm longus, squamae 150-190 (25-38 x 5) pro
strobilis, alifoimes, subtiliter velutmosae, pilis 0.03-0.06 mm, erectis;
bracteae 0.8-1.0 cm longae, sessiles, apice leviier litteram "V"-formantes
cum alls 2 inciso-serratis, fimbnatis costam non-prommentem e.xcendentibus;
axis anguste-conicus; semina 1 1 mm longa, 3 mm lata, ala 21 mm longa, 12
mm lata, cucullo 7 mm longo.

DESCRIPTION: tree with straight, usually single trunk, loose conical crown,
ascending and honzontally spreading, later slightly descending branches, with shiny
dark green foliage; bark: thin, smooth, light gray in young age, later ruptured to
irregular plates; branches: chestnut brown, sporadically hairy up to 5-6 years;
bud: (partially) hidden by short, incurving terminal leaves, flat, covered by a few, up
to 15 triangular scales, not resinous outside; branchlets: green, yellowish green,
thinly pubescent with fine (0.10-0.13 mm long), faintly brownish subclear hairs,
slightly contracted toward their base; leaves: on stem "subdistichous", ascending,
and slightly forward directed, forming an open "V" on the upper (dorsal) side of the
branchlets, somewhat shorter and conspicuously reflexed when (in helical order) they
reach or leave their uppsermost position on the branchlets, 3-5 cm long, 2 mm wide,
parallel sided, rounded and shortly (0.14-0.20 mm) emarginate, flexible and rather
leathery, shiny dark green with or without up to 5 (2-3 mm long) stomatophorus lines
on the tip of the leaves or on stronger branches with 2-3 lines reaching below the
middle of the length of the leaves, their ventral side silvery gray with narrow green
margins (0.15-0.20 mm) and midrib (0.2-0.4 mm), often with broken lines of, or
completely covered by stomas and two broad silvery stripes, typically with 14-18 (in
narrower shade leaves 10-12) stomatophorus lines with densely set (12-13/mm), fine
(0.03-0.04 mm) narrow elongated stomata; leaves in cross section flat, "avicular" to
"epsilon" shaped with 2-4(!) median resin canals; strobiles: female- 3-4 cm long,
with short (4-5 mm), first appressed, later reflexed, rounded, irregularly serrate, pale
green bracts with sessile midrib exceeded by the margins; male- 1.2 cm long on 3 mm
peduncle, consist of about 100 (20 x 5), 1.8 mm wide, hairy fertile scales; cones: 8-
9 X 3.6-4.2 cm, short (0.4-0.8 cm) stemmed "peduncled", cylindncal, rather smooth
in outline with 150-190 (25-38 x 5) cone scales; rounded at apex, green, grayish
green before fully ripened, cone scales are "wing-shaped", finely velutinous with very
short (0.03-0.06 mm), upright hairs; bract short (0.8-1.0 cm), enclosed (sessile),
slightly "V" shajjed at end with incised serrate, fimbriate wings exceeding the non-
prominent midnb; axis narrow conical; seeds: 11x3 mm, wing 21 x 12 rrun with
7 mm long hood.

NAMED: after the indigenous people living in the area where the species was
found. Zeipotecs (here Zapotek to avoid the sound modification in Latin of "c" before
"e") are the largest native nation of Oaxaca, with a macro-otomangue language and old
culture (Monte Alban). They are by nature fnendly and excellent in forest management
and conservation.

DISCUSSION: this rare species with two to four resin canals in the leaves, green
cones with short (sessile), inserted bracts, was found in a hidden valley of the cloud
forest region northeast of the town of Ixtldn de Judrez, Oaxaca, in an area where only
Abies hickelii and (very locally in the wider region but not near the site) A.
guatemalensis was known. Although crown and foliage is similar at first sight, Abies
hickelii is completely different with its slightly emarginate (not bifid) and very dark
green leaves with up to 13 resin canals (not 2-4) and larger (10-12 cm long), blue-
pruinose (not pruinose green) cones with long exerted, upright, pointed (not enclosed,
sessile) bracts with prominent tip. Abies guatemalensis has similarly "bractless"

Debreczy & Racz: New conifers from Mexico 227

cones, but it has a denser foliage (close to twice as many leaves as it is in this species
on equal length of comparable shade branchlets), straight and vertical (not
subdistichous, partially ascending on the stem) and has two (not 2-4) resin canals.
The cones of that species are typically pruinose blue (not green) with wide cyathiform
"wing shaped" (not flabellate) cone scales with elongated (not short) base section, its
bracts are slightly enclosed or occasionzilly finely exserted, (not short, "sessile"), with
finely protruding (not absent) tip.

DISCOVERY: The species is icnown so far only from a wide valley about 10 km
directly northeast of Ixtldn de Judrez, toward the place called Portillo, at an alutude of
about 2700 m. We first visited this site in December 1991 and later on February 4,
1992. Faced with logging roads, often muddy and with many divisions, we asked the
help of Mr. Gustavo Santiago Ramfrez, a native of Ixtl^ (then a biology student at the
Institute Polit&nico Nacional, Ciudad de Mexico) to help us in exploring the area.
During our second trip with the local commissanat and with the guidance of Sr.
Ignacio Crisoho, we made collections of a fir we considered to be Abies hickelii from
a distance, butcunous because of its green cone-inflorescences (DRB #32957a). On
the basis of the different shape of the bracts and green color of the strobiles, we
suspected that it was a new species and decided to make a new collection later. On
June 30, 1994, we made a third trip to the area and we obtained full size cones which
proved it to be a new species to science. Finally, in October, Mr. Gustavo Santiago
Ramirez (DRB ^f 41 300) collected fully developed cones for further study. The cones
this time of the year seemed to be mature, but still green. This new fir of the humid
cloud forest ("bosque mes(3filo de montana") region grows on steep slopes in rich,
mixed evergreen forests, associating with 65 (or more) species, including Alnus
glabrata Femald, Arbutus xalapensis H. B. K., Cleihra mexicana (Lindl.) DC,
Comarostaphylis conzattii Small largula Zucc., Cornus disciflora, Crusea coccinea
DC, Gaultheria hirtiflora Benth., G. acuminata Schltdl. & Cham., Quercus casianea
Nee, Q. crassifolia H. B. K., Q. taurina, Senecio andrieuxii DC. Completion of a
vegetation profile of the habitat is in progress. More study is necessary to establish the
full distribution of the species.

CONSERVATION: the habitat of this distinct, rare species is very important to
preserve. Mr. Ramirez, in collaboration with Dr. Enrfque Martinez y Obeja (Institute
Tecnol6gico de Oaxaca), works to secure conservation land around the habitat of the
species and conduct further investigation of the distnbution and habitat of the species.
Seeds will be distnbuted from Ixtlln later.

PARTICIPANTS IN FIELD WORK: Sr. Ignacio Crisoho (Commissanat, Ixtldn
de Julrez), Dr. Gyongyv^r Bir6, Dr. Zsolt Debreczy, Dr. Istvln Rlcz, (I. D.R.I, and
Hung. Nat. Hist. Museum, Budapest, Hungary), Vince Zsigmond (University of
Horticulture, Budapest, Hungary), Mr. Gustavo Santiago Ramirez (SERBO Inc.,
Oaxaca).

Abies guatemalensis Rehd. var. longibracteata Debreczy & Rdcz, var. nov.
Plate 1:4; ''Abies D" in NewsBnef No. 6 of I. D.R.I. (Sep. 1994). TYPUS:
MEXICO. Guerrero: Sierra Madre del Sur, near Yextla, 2400 m, January 10,
1994, Debreczy, Bird, Rdcz, el al. M34763 (HOLOTYPUS: BP; Isotvpi: A,
CHAP,E,MEXU,NA).

A specie typica strobilis matuns 10-12 cm longis, 4 cm latis et bracteis
2.0-2.2 cm longis, excertis differt.

228

PHYTOLOGIA volume 78(4): 2 17-243 Apnl 1995

1 mm

Fig i . Analyiical sketches of leaf ctoss sections of the new Abies species, partly in comparison

with other species,
al. a2: Abies neoduran'jensis sp. nov.
bl,b2: Abies dunuiijensis Mart,
cl. c2, c.'^: Abies hi(lal};ensis sp. nov.
d: Abies /.apotekensis sp. nov.

Debreczy & Ricz:
Plate I

New conifers from M^xicx)

229

Plate I: cones of the newly descnbed Abies species and vaneties. 1: A. hidalgensis
(resin canals 5, up to 7); 2: A. neodurangensis {res\n canals 2); 3: A. zapotekensis
(resin canals 4); 4: A. guatetnalensis var. longibracieaia (with its long exserted
bracts).

230
Plate II

PHYTOLOGIA volume 78(4): 2 17- 243

Apnl 1995

Plate II: newly descnbed Pinus species and vaneties. 1-2: cone and details of the
crown of P. yecorensis (Sonora). 3: cone of P. oaxacana var. diversiformis
(Oaxaca). 4: cones of P. /aw^onn var. graci'/ts (Oaxaca).

Debreczy & Ricz: New conifers from Mexico 231

NAME: longus (Lat. adj.) = long; bngibracleala = long bracted; refers to the
differentiating character of the variety in a:)mpanng the species.

DESCRIPTION: tree, identical with the species except for the larger cone ( 10-12
cm), the wider and longer cone scales (3.0-3.2 x 2 cm) with longer (2.0-2.2 cm) and
prominently (4 mm) exserted, acuminate, upcurving bracts, and the wider, more
angular, llabellate seed scale, wing.

DISCUSSION: comparing the vanety with the typical Abies gualemalensis, it
differs from the typje in the size of the cones (10-12 x 4 cm; not 8-11), in the wider,
longer (3.0-3.2 x 2 cm) cone scales with 2.0-2.2 cm (not 1.5-1.7 cm) long,
acuminate bracts with long (4 mm) exserted, upcurving tip (not short, enclosed,
abruptly acute or rounded at apex or with lateral wings, exceeding the up) and by the
more angular and flabellate (not rounded) wings. Other characlenstics are equal with
those of the species. Abies gualemalensis var. lacanensis (Lund.) Mart., reported
from Volcjin Tacand from the stale of Chiapas from between 3500-3800 m is similar,
but the branchlets of var. bngibracleala are glabrous or faintly hairy (not densely
hairy), its leaves are longer, 3.5-5.5 cm (not 1.2-3.6 cm) long, the cones larger (not
10 X 4 cm), the cone scales larger (3.0-3.2 x 2 cm, not 2.7 x 1.5 cm) with
consistently long exserted (not subequal to slightly exserted bract). The tree is fairly
common in the Yextla area associating with over 65 higher plant species. Its plant
communities are dominated by Cupressus lusiianica, Quercus spp. (4),
Chiranthodendron pentadactylon Larreategui; among shrubs. Dahlia excelsa auct.,
Mahonia lanceolata Fedde, Oreopanax xalapense (H. B. K.) Decne. & Planch.,
Philadelphus mexicanus Schltdl., and Solanum cervantesii Lag. are frequent.

CONSERVATION: The Convention on International Trade in Endangered
Species of Wild Fauna and Flora (CITES) lists Abies gualemalensis among the most
protected corufers. This will not save the species itself: for example, logging in most
areas where this conifer grows is common. Taking further steps toward full protection
are needed, including preservation of the beautiful conifer stands with var.
longibracieata. It is doubtful whether the new vanety will be recognized as "CITIES
1" and distinguished from Abies religiosa (H. B. K.) Schltdl. & Cham., frequent in
the area.

PARTICIPANTS IN HELD WORK: Dr. Gyongyv^r Bir6, Dr. Zsolt Debreczy,
Dr. Istvln Rlcz, (I. D.R.I, and Hung. Nat. Hist. Museum, Budapest, Hungary), Eva
Kert&z (Botanical Collection, Munkicsy Mih^y Miizeum, Bek^scsaba, Hungary),
Yipghao Zhao (Botanical Garden of the Chinese Academy of Sciences, Beijing,
China).

5. Abies lowiana (Gord.) A. Murr. var. viridula Debreczy & Rdcz, var. nov.
''Abies E' in NewsBnef No. 6 of I. D.R.I. (Sep. 1994). TYPUS: MEXICO.
Chihuahua; Sierra Madre Occidental, near Ciudad Madera, 2700 m, Mav 6, 1994,
Debreczy, Bird, Rdcz. elal. $39711 (HOLOTYPUS: BP; Isotypi: A, CHAP, E.
MEXU.NA).

A specie typica foliis ramulorum stenlium conspicue regulariter
dichotomice ordinatis, in latere dorsali lucide vindibus; lUis ramulorum
fertilium prorsum directis et parum declinatis differt.

DESCRIPTION: tree with straight, usually single trunk, loose conical crown,
ascending and horizontally spreading, later .slightly descending branches, with shiny
light green foliage; bark: thin, smooth, light gray in young age, later ruptured to

232 PHYTOLOGIA volume 78(4):2 17-243 Apnl 1995

irregular thick plates; branches: brownish yellow, sporadically hairy up lo 5-6
years; bud: ovate, thickly covered by resin; branchiets: yellowish green when
mature, thinly pubescent with fine (0.08-0.13 mm long) brownish, faintly brownish
or subclear hairs, often long pointed widened at their base; leaves: on stem
"subdistichous", spreading, slightly forward directing (shade) or upcurving (exposed)
at nght angle, forming an open "U" on the upper (dorsal) side of the branchiets,
slightly shorter when (in helical order) they reach or leave their uppermost position on
the branchiets, 3-5 cm long, 2.2 mm wide, parallel sided, rounded (fertile) pointed or
very finely (0.04 mm) emarginate on shade leaves; leathery, shiny dark green above
with up to 10 stomatophorus lines all along on middle of leaves, their ventral side
greenish gray with neurow (0.2-0.3 mm) green margins and wider midnb (0.5 mm),
often with few broken lines of stomas and two broad silvery stnpes, typically with 9
(in narrower shade leaves 4-6) stomatophorus lines with moderately densely set
(9/mm), fine (0.03-0.04 mm) oval stomata; leaves in cross section elliptical, with 2
median resin canals; strobiles: female- 5-6 x 0.8 cm with first appressed, later
reflexed, short (4x5 mm), rounded triangular, irregularly serrate bracts with slightly
prominent tip of the bract-needle (midvein), pale green or less often reddish green;
cones: not seen, cone scales regular, as in the species.

NAME: viridulus, viridula (Lat. adj.) = greenish; refers to the color of the foliage,
markedly distinguishes this geographically isolated variety from the northern type with
gray, pruinose needles.

DISCUSSION: Abies lowiana was not distinguished and recorded in Mexico so
far. The species, mistakenly considered as a variety of A. concolor (A. c. var.
lowiana [Gord.] Lemm.), should be considered as the "continental vicanant" of the
Pacific species A. grandis. It is easy to separate it from A. concolor in the strict sense
even by some prominent, easily available outer morphological characters: in A.
lowiana the shade branches and branchiets are rough and hairy (not smooth and
glabrous), the winter branchiets are brownish yellow (not greenish gray or gray); the
leaves, positioned in regular helical order are spreading evenly out to horizontal
position appearing "dichotomous" or "pectinate" on sterile, or pectinately upcurving in
right angle to both sides of the branchlet on fertile shoots (not diverging in all
directions according to the helical order and often curving back- and forward when in
upper position). On primary terminal shoots, "leaders", the leaves are abruptly
upcurving to parallel the shoot such as in A. grandis (not spreading or ascending up lo
45°). They are 3-5 cm long, rounded at apex on regular shoots (not up to 10 cm or
more, acute and abruptly pointed); the stomata arranged in two silvery stnpes below,
with up to 9 stomatophorus lines in each and in a defined central stomatophorus band
near the central longitudinal groove above the midvein, with up to 11 lines (not evenly
spread in both sides); in cross section the shade leaves are elliptical (not rectilinear);
the cones of A. lowiana are typically conspicuously uneven in outline, light reddish
brown, rarely green (not more or less even in outline, purplish, less often green). The
variety (var. viridula) differs from the northern type by having less pruinose leaves
above and shiny green leaves below. This could be a result of the ecological
differences in the habitat of the two plants: such as the canyon environment with
shorter dormancy period in the southern type, while exposed high mountain slopes
with long winter and hot, dry summer in the northern one.

DISCOVERY: the species, represented here by the vanety, is found in a canyon
north of Madera, Chihuahua. We saw several specimens in the shady north-
northeastern side of the wide valley about 10 km northeast of Madera, Chihuahua at an
altitude of about 2700 m. We visited this site on May 4, 1994 and found the tree with

Debreczy & Ricz: New conilers Irom Mdxico 233

well developed female and male cone-inflorescences (strobiles). We asked the help of
Mr. Leonel Iglesias Quitierrez, Director Forestal NC Chihuahua, Chihuahua, and
M.C. Raul Narvaez Rores, Jefe del Campo Expenmental Madera, associates of
INIFAP (InstJtuto Nacional de InvesUgaciones Forestales y Agropecuanas), CIR
(Centros de Investigaci6n Regional del INIFAP) Noroeste to make this exploration
possible. The trees formed a mixed stand with PseudoLsuga rnenziesii (Mirb.) Franco
S.I., with p»articularly big (some 120 cm DBH) specimens of the latter along the litlle
stream in a side valley, all cut down in the near past. Other associated species include
Pinus durangensis Mart., Pinus ayacahuite Ehrenb. var. brachyptera Shaw, Acer
glabrum Torrey, Arbuius glandulosa Mart. & Gal., Cestrum nocturnum L., Cornus
sericea Michx., Juniperus deppeana Steud. var. robusta Mart., Lonicera involucrata
(Rich.) Banks, Lonicera pilosa (H. B. K.) Willd., Mahonia repens (Lindl.) G. Don
var. pumila (Greene) Fedde (or a form of M. eutriphylla Fedde), Populus tremuloides
Michx., Quercus candicans Nee, Ribes madrensis Coville & Rose, Salix gooddingii
Ball., Salix sp., Symphoricarpos roiundifolius A. Gray (the other 35 herbaceous or
semiwoody species include Erigeron, Euphorbia, Fragaria, Galium, Gnaphalium,
Lupinus, Luzula, Poa, Polypodium, Plantago, Potentilla, Ranunculus, Rubus,
Senecio, Tluilictrum, Valeriana, Veratrum, and Viola).

CONSERVATION: the habitat of this distinct fir, probably rare in its southern
range, would be very important to preserve. Sr. Leonel Iglesias Quitierrez, Director
Forestal Noroeste Centre, Chihuahua, Chihuahua, and M.C. Raul Narvaez Flores,
Jefe del Campo Experimental Madera had been asked to collaborate in secunng the
species and to work on the possibilities of the conservation of the land around the
habitat, also to conduct further investigation of the distribution and habitat of the
species and this variety.

PARTICIPANTS IN HELD WORK: Dr. Gyongyv^r Bir6, Dr. Zsolt Debreczy,
Dr. Istvdn R^cz, (I. D.R.I, and Hung. Nat. Hist. Museum, Budapest, Hungary), Eva
Kert^sz (Botanical Collection, Munkicsy Mihdly Miizeum, B^kdscsaba, Hungary),
Vince Zsigmond (University of Horticulture, Budapest, Hungary).

6. Pinus yecorensis Debreczy & Ricz, spec. nov. Rate 11:1-2 '"Pinus A" in
NewsBnef No. 6 of I. D.R.I. (Sep. 1994). TYPUS: MEXICO, southeast
Sonora: near Y6cora, 1800 m. May 9, 1994, Debreczy, Rdcz, Bird, el al. #39798
(HOLOTYPUS: BP; Isotypi: A,CHAP,E,MEXU,NA).

Arbor 18 m alta, trunco simplici, recto; habitus late conicus
("araucarioideus"); truncus juvenilis diamelro 10 cm et plus cortice exciratus,
dense squamosus cum squamulis faale separabilibus panmculos pallide
ferrugineos relinquentibus, postea fuscus usque atrocinereus; in truncis 40 cm
diametro 7 costae discontinuae laminarum corticis (25-40 x 5- 10 cm), laminis

2 cm crassis, planis, rotundato-angulanbus; rami crassi (in arbonbus vetustis

3 annorum 3 cm diametro), foliis persistentibus 4 annorum; rami conspicue
squamati, squamis magnis, aurantiaco-brunneis, ultra 5 annum squamae
pnmordiales cinereae; squamae corticis ultra 8 annum plane evolutae; ramuli
crassi, dense squamulosi, glabn; brachyblasti cum squamis non-deciduis.
squamae basales valde cannatae (4 mm), squamae supenores 5, lenues,
rugosae, et superius filiferae; in arbonbus adultis squamae pnmordiales
primum longe decurrentes, |x>stea basi breve (6 mm), cordata, lucide
aurantiaco-brunnea cum apice filifero, longe acuminate; folia 5 pro
fasciculis, ad basin crassa cum squamis basalis validis, 4.0-4.5 mm longa;

234 PHYTOLOGIA volume 78(4): 2 17-243 Apnl 1995

28-35 cm longa, in sectione transversali paene equilalero-tnangularia, in latere
dorsali 1.2-1.4 mm lata, dense serrata, lucide vindia, m latere dorsali lineis 3-
4 stomatum, m latenbus ventralibus 3 lineis, margme tenuissimo (O.CX^ mm)
instructa; dentes m aciebus foliorum densae (4-5 pro 1 mm), ellusae, tenues
(0.03 mm), pellucidae; strobili feminei juveniles ovajes, 2.0-2.5 cm
longi, 1.5-2.0 cm lati, ex 200 squamis compositi; squamae 3-4 mm longae,
ad 2.0-2.5 mm ascendentes, flavo-brunneae, pruinosae (hmc subroseae), m
apices 1 mm longos, rectos, obtusos, verticaJiter expansos terminatae;
pedunculo 1.2-1.4 cm longo; strobili maturi 8-12 cm longi, 6-9 cm lati
(inclusi), 7.0-9.5 cm lati (aperti), ovati, circa medium latissimi, leviter
asymmetrici, quasi sessiles, firmiter affixi, non-serotmi, pallide brunnei;
strobili ex 130-150 (26-30 x 5) squamis compositi; apophysibus plarus \el
protuberantibus, retrorse-rotundatis, umbone 3.5 mm longo, 2.2-3.0 mm
lato, cum aculeo inconspicuo, obtuso, 0.6 mm longo terminate; semina
ovata, 6.5-4.0 mm longa, in latere ventrali levia, ordinatione mosaica pallide
brunnea, pallido- et atrocinerea, latere dorsali aspero, subuliter sulcato et
vanole notato, supenus pallide flavo-brunneo; ala 3 cm longa, infra medium
latissima, latere intenore recta, semen ad basin circumdans.

DESCRIPTION: tree, 18 m tall, with straight, typically single trunk, broad
conical ("araucaroid") crown, with upcurving, ascending, horizontally spreading and
upcurving, finally descending or declining thick pnmary branches with relatively few
(usually 25-30) lateral branches with large clumps of heavily pendent, shiny bnght
green foliage; trunk: straight, usually clean, up to 4-10 m; bark: nbbed from 10 cm
upward, densely scaly with easily attached plates leaving light reddish-brown patches,
changing to grayish brown to dark gray (at 40 cm DBH: 7 discontinuous nbs of 25-
40 cm X 5-10 cm and 2 cm high flat topped rounded-angular plates); branches:
thick (mature tree, 3rd year fertile: 3 cm across), leafy for 4 years, prominently scaly
with large orange-brown, later, (from year 5), gray, eroded pnmordial (pnmary)
scales; cortical scales appear from year 8; branchlets: strong, densely scaly,
glabrous; brachyblasts: with non-deciduous scales (sheaths), basal scale pair
strongly keeled (4 mm), upper scales 5 in number, thin, wrinkled and filiferous at
margin above; leaves: 5 in a fascicle thick at base (4.0-4.5 mm at basal scale pair),
28-35 cm long, 1.2-1.4 mm wide on dorsal side, triangular in cross section, densely
serrate, shiny bnght green with 3-4 stomatophorxis lines on dorsal and 3 on the two
inner (ventral) sides, margin very narrow (0.06 mm), teeth densely set (4-5/mm),
spreading, fine (0.03 mm), clear; pnmordial leaves of young plants (euphyllum) not
studied; on adult plants the scales are first long decurrent, later the base is short (6
mm), cordate, shiny orange brown with filiferous, long acuminate apex; conelets:
large to the genus (2.0-2.5 x 1.5-2.0 cm), oval, on 1.2-1.4 cm long stem, subtended
with recurved or revolute entire budscales, consist of about 200, 3-4 mm wide 2.0-2.5
mm high, elevated, yellowish brown, pruinose (hence pinkish) scales, terminated in 1
mm long vertically spreading straight blunt tips; cones: medium sized, 8-12 x 6-9
cm when closed, 8-12 x 7.0-9.5 cm when open, ovate, widest in the middle of its
height, slightly asymmetrical, neariy sessile, firmly attached, non-serotinous, pale
brown, the 130-150 (26-30 x 5 developed) scales have fiat or protuberant, slightly
retrorse, rounded apophysis with 3.5-5.0 x 2.2-3.0 mm umbo terminating in
inconspicuous (0.6 mm) blunt pnckles; seeds: small, ovate, 6.5 x 4 mm, smooth
with pale brown, light- and dark-gray mosaic pattern on lower/ventral side, uneven,
finely grooved and warty, |>ale yellowish brown above. The wing is 3 cm long.

Debreczy & Ricz: New conifers from Mexico 235

broadest below the middle, straight on inner side, light yellowish brown with some
dark shade and lines toward the edge.

NAME alter the mountain village Yd<x)ra in western Sonora, above which the
species was found.

DISCUSSION: This very distinct new pine from the high mountains of Sonora
has five needles in a fascicle and, in general appearance, it is ver\' much like Pinus
tnichoacana Mart., but with conspicuous, shiny light green (not dull grayish green)
needles and very different cones, somewhat similar to the northeastern species Pinus
estevezii (Mart.) Perry or to its close ally, Pinus aputcensis of central Mexico. The
cone is also comparable to Pinus nubicola Perr\ of the very south. It differs from P.
estevezii with much thicker branches (3 cm; not 1.4 cm in third year), green (not
pruinose) branchlets, larger primordial (scale) leaves (1.4 x 0.4 cm, not 0.8 x 0.2
cm), longer and thicker (28 x 6 mm, not 16.0 x 3.8 mm) enclosed (basal) section of
the stronger (1.3 mm, not 0.8 mm) and less densely serrate (40 teeth, not 50 teeth/cm)
needles. The cones of f. yecorenju are somewhat smaller (10-12 x 7-9 cm, not 12-
14 X 8- 10 cm), widest in the middle (not toward base), with ngid (not flexible) cone
scales, light brown (not reddish-chestnut brown) around the seed pnnt with light (not
reddish brown) attachment zone below the more rounded- protuberant apophysis on the
dorsal (abaxial) side, with larger umbo ended in short (0.7 mm) and blunt (not 1 mm,
pointed) arista. The seeds have a brown (not black) pattern on the ventral side, with
wings more gradually (not abruptly) narrowing toward the tip. Easy to distinguish P.
yecorensis from P. engelmannii Can", common in the lower region of the area with its
bark (flat, shallow nbs, thin plates change light brown to dark gray, not high, dark
gray nbs with conspicuous orange fissures), 5 needles/fascicle with 3 stomatophorus
lines on dorsal side and three resin canals in each (not 3 needles, with 7-8 lines, 5-8
resin canals) and shorter, thicker (28 x 6 mm, not 30 x 3 mm) enclosed section of the
needles. The tips of the conelet scales in P. yecorensis are honzontally spreading and
pruinose (not directing downward, not shiny brown), the cones are egg shaped with
rounded somewhat flat triangular tip of the scales with short blunt umbo (not long
cylindrical with triangular, abruptly elevated scales with prominent, prickly umbo).
Pinus michoacana with its sparse, thick branches is very similar in habit, but P.
yecorensis has shiny green needles of thick (7-8 mm) base, 3-4 stomatophorus lines
and narrow margin with spreading teeth on dorsal side (not 4.5 mm' wide at base,
pruinose grayish green with 6-8 lines, margin [0.15 mm] and teeth directing somewhat
inward). The cones are verv' different in P. michoacana, they are long (20-30 cm),
cylindrical, often curved. Compared to P. yecorensis, P. durangensis Mart, and P.
maximinoi H.E. Moore, present in the region, have slender stems, finer needles and
much smaller cones with thin, flexible scales, not comparable with our sjaecies.

DISCOVERY: On the way from Barranca del Cobra on May 8, 1994, we turned
on the road leading to the town of Obregon. Just 12 miles before reaching the
mountain village of Y&ora, we saw a few strange, long needled, young pines in the
dark along the roadside. They were neither Pinus durangensis nor Pinus maximinoi.
Next morning, we returned to the area and found adult trees forming scattered stands
in a dry canyon, associated with Pinus herrerai Mart., Arbutus arizonica (A. Gray)
Sarg., A. glandulosa, Quercus candicans Nee, Q. coccolobaefolia Trel., Q.
magnoliifolia, Quercus spp., Prunus serotina Ehrh. subsp. capuli (Ca\.) McVaugh.
and Rhamnus betulaefolia Greene. The altitudinal range of the species, around 1800
m, seemed to be very narrow, a hundred meters below P. leiophylla Schiede ex
Schltdl. & Cham. var. chihuahuana (Engelm ) Shaw and P. lumliolizii B.L. Rob. &
Femald to just above Y6cora, where P. engelnumnii Carr. lakes over the area.
Leaving Y6cora, a few miles toward Obregon, we again came across scattered stands

236 PHYTOLOGIA volume 78(4):2 17-243 Apnl 1995

of this species, but for only a few miles along the road. Interestmgly, it was found
again lOO's of miles south of there, in the mountains of SinaJoa, along the road from
Mazatldn to El Salto, Durango. With its distinct characters, we considered the latter to
be a new geographical vanety (see below).

CONSERVATION: It is located far from cities, so there was no opportunity to
make steps in this matter. In the same area, other conifers such as Pinus durangensis,
P. engelmannii, P. herrerai, P. leiophylla, P. I. var. chihuahuana, P. lumholtzii, P.
maximinoi, and P. oocarpa Schiede ex Schltdl. are common, but do not appear in
distribution maps, suggesting that the area is not yet well explored. The tree seems to
be restncted to the localized sites and protection of some populations should be
considered.

PARTICIPANTS IN FIELD WORK: Dr. Gyongyv^r Bir6, Dr. Zsolt Debreczy,
Dr. Istv^ Rdcz, (I. D.R.I, and Hung. Nat. Hist. Museum, Budapest, Hungary), Eva
Kert6sz (Botanical Collection, Munkdcsy Mih^y Miizeum, B^k^scsaba, Hungary),
Vince Zsigmond (University of Horticulture, Budapest, Hungary).

7. Pinus yecorensis Debreczy & Ricz var. sinaloensis Debreczv & Rdcz, var.
nov. "Pinus B" in NewsBnef No. 6 of I. D.R.I. (Sep. 1994). TYPUS:
MEXICO. Central Sinaloa, Capilla Textia, 1200 m. May 9, 1994, Debreczy,
Rdcz, Bird, et al. #39896 (HOLOTY PUS: BP; Isotypi: A,CHAP,E,MEXU,NA).

A specie typica strobilis maturis (apertis) 4.5-6.5 cm longis, 4.5-6.0 an
latis et peduncuiis 0.5-1.2 cm longis differt.

DESCRIPTION: general character is idenucal to that of Pinus yecorensis (above),
but the trees are usually smaller (12-16 m tall); needles: narrower (1 mm; not 1.2-
1.4 mm wide) with more dense serration (6-7/mm; not 4-5/mm) and with 3
stomatophorus lines on the dorsal sides, 2-3 lines on the two ventral sides (not 3-4
slomatophorus lines on the dorsal and 3 on the two ventral sides); conelets: about
half in size (1.2 X 1.1 cm; not 2.0-2.5 x 1.5-2.0 cm) with smaller scales (1.8 x 1.2
mm; not 3.5-5.0 x 2.2-3.0 mm) and shorter (0.5-0.7 mm; not 1 mm) but often
sharper prickles; cones: smaller (4.5-6.5 cm long, 4.5-6.0 cm wide when open; not
8-12 X 6-9 cm, or 8-12 x 7.0-9.5 cm when open), consisting of 80-110 (not 130-
150) developed scales with elevated, usually rounded, occasionally conical and
pointed apophyses (up to 7 mm) on one side, flat and rhombic on the side, facing to
the branches, along with a peduncle of 5 to 12 mm; seeds: smaller (with wing, 1 .4
X 0.4 cm), densely shaded with dark lines (not 3 x 0.9 cm and light brownish
yellow).

NAME: after the Mexican state of Sinaloa in which the species was first found.

DISCOVERY: While traveling from Mazatldn, Sinaloa to H Salto, Durango we
were surprised to see a similar (unknown) pine we found on the slopes around Yecora
hundreds of miles north. It is fairly common on hot, dry, rocky slopes between 1 , 100
and 1300 m altitudes for just a few miles above and below Capilla Textia, Sinaloa, 74
miles east of Mazatl^. It associates with Pinus durangensis and P. maximinoi, and as
in around Capilla Textia, with P. oocarpa and Quercus magnoliifolia. The cones often
remain on the branches for several years and, from a distance, the trees can easily be
mistaken for P. oocarpa.

CONSERVATION: Steps will be taken in collaboration with the UCODEFO in H
Salto.

PARTICIPANTS IN FIELD WORK: as with the above species.

Debreczy & Ricz: New conifers from M^xico 237

8. Pinus oaxacana (Mart.) Mirov-* var. diversiformis Debreczy & R^cz, var.
nov. Figure: Plate \\.3\"Pinus C" m NewsBnef No. 6 of I.D.R.l' (Sep. 1994).
TYPUS: MEXICO. Oaxaca; south of the village of I.xlldn de Judrcz, 2000 m.
Feb. 26, 1994, Debreczy, Bird, Rdcz. el at. #37897 (HOLOTYPUS: BP, Isotvpi:
A,CHAP.E,MEXU.NA).

A specie typica ramuiis olivaceo-brunneis, strobilis maturis angustionbus
( 12 cm longis, 6 cm latis), apophysi busque potius aequaliter protuberantibus
differt.

NAME: (Lat. adj.: diverse = different; diversiformis = different form) to express
its divergent characteristics, comparing it with the species, Pinus oaxacana (Mart.)
Mi rev.

DESCRIPTION: large tree, 20 m tall or probably more, with broad, rounded
crown, very similar if not identical with that of Pinus oaxacana. The needles, 5 in a
fascicle, long (25-30 cm) and fine, just like in the species but with a slightly different
shade with less shiny bright green color. The branchlets are green first, soon
yellowish green and brown, not bluish white pruinose as is typical in the species. The
cones (just like those of Pinus anenuata Lemm. from a distance) are elongated ovate to
cylindrical, typically twice as long as wide (12x6 cm) with elevated, but not strongly
protuberant, spiny umbos (as it is in the spjecies) and with slightly elevated scales with
strong transverse keel toward the apex.

DISCUSSION: this pine, occumng sporadically above the Rio Grande valley in
Central Oaxaca, typically intermingled with the species and with other pines, is distinct
with its narrower, more symmetncal cones (not strongly asymmetncaJ ovate) with
short, evenly protuberant apophyses toward the base of the cone (not with 1.5-2.5 cm
long, strong projection of the apophysis particularly toward the base on the dorsal
[abaxial] side). The young branchlets are shiny green, later yellowish-greenish brown
and finally brown, not pruinose bluish white.

DISCOVERY: while studying the variability of Pinus oaxacana, we encountered
single trees and small groves of this vanety, intermingled with the species and other
trees between and around Ixtldn and Guelatao, Oaxaca, at 1,800-2,(XX) m. The namral
association in the region of this conifer had been destroyed, and its occurrence by now
is likely restricted to secondary grasslands and scrubs with scattered stands of Acacia.
Quercus, woody Verbesina, and Vemonia sp)ecies. Among conifers, besides P.
oaxacana, only P. leiophylla, P. lawsonii, and P. michoaca/ia was found to grow
nearby.

CONSERVATION: as descnbed above, this rare vanety, found on both pnvate
and community lands, is an important pine from a phylogenetic aspect. Its scattered
specimens and groups, intermingled with other species, can be preserved only after
careful survey and mapping the specimens in collaboration with the local commissariat
and conservancy pjersonnel.

PARTICIPANTS IN HELD WORK: Dr. Gyongyv^r Bir6, Dr. Zsolt Debreczy,
Dr. Istvln Rdcz, (I.D.R.l. and Hung. Nat. Hist. Museum, Budapest, Hungary), Eva
Kert^sz (Botanical Collection, Munkicsy Mihaly Miizeum, Bek^scsaba, Hungary),
Vince Zsigmond (University of Horticulture, Budapest, Hungary), Yinghao Zhao
(Botamcal Garden of the Chinese Academy of Sciences, Beijing, China) and
Earthwatch volunteers.

238 PHYTOLOGIA volume 78(4):2 17-243 Apnl 1995

9. Pinus lawsonii Roezl var. gracilis Debreczy & Rdcz, var. nov. Plate 11:4;
'T//iwjD"in NewsBnef No. 6of I.D.R.I. (Sep. 1994). TYPUS: MEXICO.
Oaxaca; above the town of Oaxaca, 4 km below EJ Estudiantes on dr\ rocky
outcrops, northwest facing slope, close to route #175, 1650 m, March 24, 1994,
Debreczy, Bird. Rdcz. el al. #38542a (HOLOTYPUS: BP; Isotypi: A,
CHAF.E,MEXU.NA).

A specie typica foliis tenuionbus, multo elasticioribus, 1.0-1.2 mm latis,
strobilis matuns lucidionbus, pedunculis eorum usque 1.5 cm longis,
squamis strobilorum angustionbus (6-11 mm latis) magis flexibilibus,
umboneque prommenti, aculeo brevi instructo differL

NAME: (Lat. adj. gracilis = thin, slender) was chosen to express its fine foliage
and relatively small cones.

DESCRIPTION: small to medium sized tree up to 15 m with typically rounded
crown; bark: separated to wide, flat, densely scaly nbs, some easily detached plates
and others, firmly attached and leaving chestnut brown patches, changing brown to
light brown to ash gray with age, fissures orange red, conspicuous on rapidly growing
trees when whole sections of the trunk show orange red; branches: upward curving,
densely scaly, orange to reddish-brown with easily detached scales, the pnmary scales
detach readily, replaced by thin bark scales; buds: small (1 cm long), pointed,
reddish brow'n with a few'white threads, bud scales (8x3 mm) narrow, triangular
with wide hyaline margin at base; branchlets: thin (6-8 mm in lateral fertile shoots),
densely covered with white bloom and narrow, nearly parallel sided scales; leaves: 3
in fascicles, 14-16 cm long, 1.2 mm wide on the dorsal side, 0.7 mm on the ventral
side, the basal section covered up to 1.4 cm, partially eroded by the following year
(0.4 cm) and remaining for 2 years, the initial scale pairs are psapery thin with 9
stomatophorus lines on the dorsal side and 4 lines each on the ventral sides, densely
serrate (60-68/cm) with small teeth (up to 0.16 mm long, with 0.03 mm clear Up);
conelets: small (1.4 cm long), oval to ovate, abruptly contracted toward the base,
consist of about 180 (56 on the contracted section), small (25 x 1.5 mm) scales, each
with 0.5 mm long blunt pnckle, directed (axilwise) upward on conspicuous (1.4 cm
long), slender, recurved peduncle with easily detached scales; cones: developing on
recurved, slender peduncle (1.3-1.5 cm long), firmly attached, non-serotinuous, small
3-6 X 2.3-4.0 cm), ovate, widest above base or at the middle of their length, slightly
asymmetrical, consist of 80-110 (6-22 x 5 developed) scales, irregularly rounded at
upper margin, shiny brown, dark chestnut brown on dorsal side with a gray seed scale
pnnt and nanxjw blackish zone on ventral (adaxial) side around the seed scale,
apophysis (5-9 x 2-4 mm at mid-cone) fiat or slightly elevated with 3-5 clearly visible
keels, umbo (3-4 x 2 mm) terminating in inconspicuous (0.4-10.0 mm long), blunt
pnckles directing forward (as to continuation of the scale); seeds: small (4.5 x 2.5
mm), ovate or rounded quadrangular, evenly brown, black and gray dotted, w ith small
(1.2 x 4.5 mm), shiny purplish gray (light yellowish gray, delineated purplish gray)
wing.

DISCUSSION: this interesting rare vanety of Pinus lawsonii, with typically
rounded crown, fine, flexible, gray to grayish green needles in threes and relauvely
small, shiny, ovate cones with prominent pnckles and up to 1.5 cm long peduncle,
grows in the lower limits of the disUibution of the species. It differs from that w ith its
slender needles (1.0-1.2 mm across), with narrow, 1.5 mm wide, enclosed basal
section, at most 2.4 mm across, measured at the papery thin basal scales (not 2.4 mm
wide and with conspicuous basal scale pair, 3.5 mm across); the needle is 1.2 mm

■■■■■■iBaaBitovwr ttv<:

240 PHYTOLOGIA volume 78(4):2 17-243 Apnl 1995

DESCRIPTION: Narrow columnar tree up to 30 m tall with first upcurving, later
ascending-upcurving, short main branches forming a columnar pyramidal habit, with
trunk diameter up to 80 cm; bark: thick with high nbs, suberous like in the smaller
trees of the species; branches: densely scaly with pnmary scales or remnants up to
8-10 years, shiny gray, later replaced by thin, gray to dark gray cortical scales; buds:
(4x2 mm) cylindncal reddish-brown, not resinous, the lower scales recurved in dry
weather; branchlets: typically short, relatively thick (1st year 3 mm across),
sparsely pubescent, first light green, later purplish brown; needles: typically short
and relatively wide (1.0-1.7 cm x 1.0-1.2 mm), parallel sided, abruptly acute,
mucronulate at the end, deeply grooved above the midnb, pruinose wi'hout
stomatophorus lines above, with two stomatophorus bands below on the upper,
adaxial surface, with 4 glaucous lines in each and conspicuous but narrow (0. 16 mm)
margin and strong (0.2-0.3 mm) midrib; cone inflorescences (strobiiies):
appears in early March, female- (1.5-2.0 x 0.6-1.0 cm) pink or green, typically
pruinose; male- (0.8 x 0.5 cm): light yellow to pinkish yellow; cones: 2-4 x 1.5
cm, rarely up to 6.5 cm long, with often less than 10, otherwise 25, rounded,
relatively small cone scales wider than long (1.5 x 2.2 cm), with dense, appressed,
short (0.25 mm) hairs; the cone scales are prominently veined on the abavial (dorsal)
side; there are shorter hairs inside around the two, smooth seed pnnts (9x4 mm);
bracts exserted, slightly longer than the midscales, upcurving toward the base (before
the cone matures or when wet), 2 cm long at the widest scales, narrow (3 mm wide)
and serrate below the middle, usually tricuspidate with short points on the outer edge
of the wings forming a short stemmed, often "closed", "V" from 5 mm below the
shortly protruding tip, becoming needle-like or nearly so; lateral lobes of the bract are
inconspicuous or absent toward the base; seeds: (old when observed) 4 x 3.0-3.5
mm, light yellowish gray-pale brown with 9x4 mm wing, light grayish yellcw in
color with or without a few short brownish lines.

NAME: after the state of Oaxaca in southern Mexico, where it was found. The
locality is just 15 km as the crow flies from the capital of the state, with the same
name, Oaxaca de Judrez.

DISCUSSION: this narrow columnar Douglas-fir (from a distance appearing like
cultivated Cupressus arizonka Greene) with grayish foliage and small cones, was
found around a huge limestone rock, Pena Pneta (Pena Piedra), on the northeaiitem
slope of a valley in the San Felipe Mountains at 2700 m altitude, a few miles north of
the town of Oaxaca. Following this, on June 28, 1994, Prof. Boone Hallberg cl the
Instituto Tecnol6gico de Oaxzica reported that "Sr. Evodio, consejo de Vigilaiicia,
Ixtepeji and Sr. Alfredo Yescas" located the tree in Rosa Blanca, a few kiloireters
north of the first location. Since September, the species has also been reported from

the (newly identified) highest mountain 3751 m, 12, 303' [not Cerro Zampoaltcpetl,
3395 m(!) known previously as the highest] of Oaxaca (according to Boone Hallberg,
"from lat. 15°22' N, long. 96°06' W; must be 16°22' N, 96°06' W, Cerro QuiexDbra,
Sierra Madre del Sur( !) 1 10 km SE of the town of Oaxaca, 35 km SE of Miahuatlan de
Porfino Diaz). All these data indicate how widely the species had been distnbuted in a
cooler, relatively more humid penod. Applying Pseudotsuga menziesii in a broad
sense, this location represents the southernmost distnbution of the species and,
simultaneously, that of the entire genus worldwide (P. wilsoniana Hay. and P.
brevifolia W.C. Cheng & L.K. Fu grow near the 22nd latitude in Taiwan and in south
China, respectively). To understand the systematic position of our variety, it is
necessary to evaluate the relationships of all species descnbed from Mexico with those
in the north and evaluate this new variety within that relationship. As discussed in the

Debreczy & Ricz: New conifers from Mexico 241

introduction, species such as the north Mexican P. flahaultii, P. rehderi, P. guinieri
and the one, considered the southernmost member of the "menziesii group", P.
macrolepis, are not consistent m their morphological characters and better to be
considered a synonym of P. menziesii. This is why the Oaxacan Douglas fir is better
treated as the southernmost and isolated variety of the main species (see introducuon).

DISCOVERY: we found this vanety in the San Felipe Mountains at 2700 m
altitude, north of the city of Oaxaca. It associates with Pinus oaxacana, Pinus
pseudostrobus Lindl. (including var. megacarpa Loock), and Pinus teocote, Arbutus
xalapensis, Litsea glaucescens, Prunus serotina subsp. capuli, Quercus candicans
Iccdophylla Cham. & Schltdl., Q. crassifolia H. B. K., Q. Umrina, and Q. castanea in
the upper level and Comarostaphylis conzattii largula, Senecio angularis auct., S.
barba-johannis DC. in the scrub, Gaultheria hxrtiflora, Pemettya cUiala Small,
Vaccinium conferium H. B. K., with a rich (Thuidium, Polytrichum) moss level with
Sedum, Crassula, F*teridophytes {Elaphoglossum, Adiantum) and a few herbaceous
plants such as Heuchera, in the lower scrub level. High stemmed (2-4 m tall)
Dasylirion lucidum aucL with nodding terminal is common in the shade of the trees.
The small forest in the deeper soil under the big rock is rather poor in the number of
species. The associations of the spjecies in this location are distinct from any other
location of the Douglas firs.

CONSERVATION: the area was visited by professors from the Instituto
Tecnol6gico de Oaxaca and re[M"esentatives of the Oaxacan and local governments.
With the new discovery of Douglas fir in the nearby Pena Rosa, two patches, each not
more than 2-3 hectares has been known. Even if the most recent discovery of
Douglas-fir (evidently the same vanety) in Cerro Quiexobra, raises the number of the
known locations to three, the tree should still be considered rare and very local,
necessitating designation of all known locations for strict conservation policy. The
protection of two locations of this rare tree, together with other rare laxa found in the
larger area, is proceeding as suggested, within the framework of a proposed "San
Felipe National Park".

PARTICIPANTS IN FIELD WORK: Dr. Gyongyvdr Biro, Dr. Zsolt Debreczy,
Dr. Istvdn Rdcz, (I. D.R.I, and Hung. Nat. Hist. Museum, Budapest, Hungary), Eva
Kertfez (Botanical Collection, Munkdcsy Mih^y Miizeum, B^k^scsaba, Hungary),
Yinghao Zhao (Botanical Garden of the Chinese Academy of Sciences, Beijing,
China) and Earthwatch volunteers.

ACKNOWLEDGMENTS

Along with the support of I. D.R.I, members and Earthwatch volunteers, we
express our sincere thanks for the generous contnbutions of Dr. Carol Dolinskas, Mr.
Walter Hunnewell and Mr. B^Ia Kalman of the Board of Directors of I.D.R.I., the
Rare Conifer Foundation, the Cncket Foundation, the William P. Wharton Trust and
the Hungarian National Science Foundation (OTKA Grant #3168). The research, of
which part of the result is descnbed here, has been partially supported by Earthwatch
and Earthwatch volunteers in 1991/92 and 1994. Our field work in Mexico has been
supported by the Secretana Desarollo Social, SEDESOL, and continuously assisted by
the Instituto Tecnol6gico de Oaxaca (ITO), where we are particularly indebted to Prof.
Boone Hallberg and Dr. Enrique Martfnez. Here we zilso give thanks for the
continuous attention of Dr. Francisco Becena Luna of Instituto Nacional de
Investigaciones Forestales y Agropecuarias, INIFAP, and his colleagues at the
Secretaria de Agricultura y Recursos Hidraulicos. SARH. For field assistance, many

242 PHYTOLOGIA volume 78(4):2 17-243 Apnl 1995

thanks are due lo Ing. Roman Perez Aranjo, Subdelegado Forestal de Guerrero, and
Ing. Rafael Norberto Jaimes, Supervisor Forestal Federal. In Coahuila, Biol. Antonio
Cano Pineda and while in Chihuahua, Dr. Enrique J. Sanchez Granillo, Mr. Leonel
Iglesias and their colleagues assisted our field trips. We also express our gratitude to
Biol. Santiago G. SaJazar Hem^dez of the Instituto Tecnologico Forestal, ITF, No.
1 and Biol. Rosalva Miranda Salazar of Unidad de Conservaci6n y Desarollo Forestal,
UCODEFO, for their help in Durango. In the Division de Ciencias Forestales,
Universidad Aut6noma Chapingo, we are particularly obliged to Dir. Saul B. Monreal
Rangel for his support all over the country and for opening the research facilities and
the Expenmental Station at Aculco, in P.N. Zoquiapan on the Ixtaccihuatl, for our
team of up to twelve at one time. We are also grateful to Biol. Emma Estrada
Martinez and Prof. Inv. Enrfque Gui'zar Nolazco of the herbarium of the above
institution for their cooperation. Finally, our thanks to Mr. Istvdn Gerendds,
Honorary Consul of the Hungarian government in Guatemala, for his support.

LITERATURE

Farjon, A. 1993. Names in current use in the Pinaceae (Gymnospermae) in the ranks

of genus to variety. NCU-2, Regnum Vegetabile vol. 128.
Ferrd, Y. 1941. La place des canaux rdsinif^res dans les feuilles des Abielin^es.

Trav. Lab. For. Toulouse. T. I, III, art. XII. Toulouse, France.
Gaussen, H. 1971,1973. Les Gymnospermes Actuelles et Fossiles. Fasc. 4andl2.

Toulouse, France.
Hallberg, B. 1994. Hallazgo de un nuevo gdnero de conifera. Noticias, Domingo

27. de Febrero de 1994., p. 9a. Oaxaca de Julrez, Mexico.
Hallberg, B. & E. Martinez y Ojeda. 1994. Coniferas en peligro de extincion.

Noticias, Viemes 27 de Majo de 1994., p. 9a. Oaxaca de Judrez. Mexico.
Liu, Tang-Shui. 1971. A Monograph of the Genus Abies, pp. 607. Taipei, Taiwan
Martfnez, M. 1948. Los Pinos Mexicanos. pp. 361. Ciudad de Mexico, Mexico.
Martfnez, M. 1942, 1963. Las Pinaceas Mexicanos; pp. 399. Ciudad de Mexico,

Mdxico.
Orr, M.Y. 1937. On the value for diagnostic purposes of certain of the anatomical

features of Conifer leaves. Not. Roy. Bot. Gard. Edinb. vol. XIX. Edinburgh,

Great Britain.
Perry, J., Jr. 1989. The Pines of Mexico and Central America. Timber Press,

Portland, Oregon.

'A compact tree (6 x 5 m) with very dense foliage was located in the land of Ixtldn de

Ju^ez, which would be valuable for gardens in zone VIII-IX; for availability and

further reference, contact Mr. Gustavo Santiago Ramirez, Ixtldn de Judrez, Oaxaca.

Gray-blue forms of Pinus lawsonii are p>articularly sf)ectacular in Guerrero, in the

Chilpancingo region, e.g., along the road from Xochipala to Yextla.

^M.C. Gufzar N. Enrique, Div. de Ciencias Forestales, Universidad Aut6noma

Chapingo, Mexico.

'Rosalva Miranda Salazar, UCODEFO No. 6, El Salto, Durango, Mexico.

'^161. Gustavo Ramfrez Santiago, of Ixtldn de Judrez, Oaxaca, Mexico.

Debreczy & Ricz: New conifers from Mdxjco 243

^ote: We believe that Pinus oaxacam (Mart.) Mirov should be the properly
authorized name. No better description with precise and clear illustrations could be
given, than what was provided by Maximino Martinez with his P. pseudosirobus
Lindl. var. oaxacam Mart. Unfortunately, in most literature Martinez's name has been
improperly dropped after Mirov elevated the variety to species rank.

Phylologia (Apnl 1995) 78(4) 244-245

NOMENCLATURAL COMBINATION IN POACEAE

Joseph K. Wipff & Stanley D. Jones

Herbarium (BRCH), Botanical Research Center, P.O. Box 6717, Bryan, Texas
77805-6717 U.S.A.

ABSTRACT

The following nomenclatural change in Poaceae is proposal:
Sporobolus compositus var. clandestinus (J. Biehler) comb. nov.

KEY WORDS: Sporobolus, nomenclature, Poaceae

Sporobolus compositus (J. Poiret) E. Merrill var. clandestinus (J. Biehler) J.
Wipff & S.D. Jones, comb. nov. BASIONYM: Agrostis clandestina J. Biehler,
Pkmtarum Novarum ex Herbaria Sprengelii Centuriam 8. 1807. Muhlenbergia
clandestina (J. Biehler) K. von Trinius, De Graminibus Unifloris et Sesqmfloris
190. 1824. Vilfa clandestina (J. Biehler) Nees von Esenbeck ex E. von Stcudel,
Nomenclator Botanicus, ed. 2, 2:767. 1841. Sporobolus clandestinus (J.
Biehler) A. Hitchcock, Contributions of the U.S. National Herbarium 12:150.
1908. Sporobolus asper (P. de Beauvois) K. Kunth var. clandestinus (J. Biehler)
L. Shinners, Rhodora 56:30. 1954. TYPE UNITED STATES. Pennsylvania:
Muhlenberg 1 15 (J^PE: PH; Type Fragment US).

Sporobolus canovirens G. Nash in N. Britton, Man. 1042. 1901. Sporobolus
asper (P. de Beauvois) K. Kunth var. canovirens (G. Nash) L. Shinners,
Rhodora 56:30. 1954. Sporobolus clandestinus (J. Biehler) A. Hitcicock
var. canovirens (G. Nash) J. Steyermark & J. Kucera, Rhodora 63:24. 1961.
TYPE UNITED STATES. Kansas: St. George, 3 September 1890.
Kellerman {TYPE: NY).

Kartesz & Gandhi (1995) reported that the basionym of Sporobolus asper
[Agrostis aspera A. Michaux (1803)] is a later homonym and illegitimate, aiuJ that
Agrostis composita J. Poiret is the earliest available name for this taxon. Sporobolus
clandestinus (J. Biehler) A. Hitchcock and S. compositus (J. Poiret) E. Merrill are
very similar morphologically. Riggins ( 1977) separated the two taxa by the following
characters: Sporobolus clandestinus has spjarsely, appressed, pubescent lemmas and a
pencarp loose when moist, whereas, S. compositus has glabrous lemmas cjid a
pericarp gelatinous when moist. We believe that this kind of morphological vanation
is best recognized at the variety level; thus making the above combination necessary.

244

245 PHYTOLOGIA volume 78(4): 244-245 Apnl 1995

ACKNOWLEDGMENTS

We are grateful to W.E. Fox (TAES) and Gretchen D. Jones (USDA, AWPMRU)
for reviewing this manuscnpt.

LITERATURE CITED

Kartesz, J.T. & K.N. Gandhi. 1995. Nomenclatural notes for the North Amencan

nora. XIV. Phytologia78:l-17.
Riggins, R. 1977. A biosystematic study of the Sporobolus asper complex

(Gramineae). Iowa State Journal of Research 51:287-321.

Phytologia (Apnl 1995) 78(4) 246-248.

NEW DATA ON DISTRIBUTION AND MORPHOLOGY FOR THE RARE
HASTEOLA ROBERTIORUM (ASTERACEAE)

Loran C. Anderson

Department of Biological Science, Florida State University, Tallahassee, Ronda
32306-3043 U.S.A.

Edwin L. Bridges

Fairchild Tropical Garden, 1 1935 Old CuUer Road, Miami, Honda 33156-4299

U.S.A.

&

Steve L. Orzell

Florida Natural Areas Inventory, 1018 Thomasville Road, Suite 200C, Tallahassee,
Honda 32303 U.S.A.

and

Fairchild Tropical Garden, 1 1935 Old CuUer Road, Miami, Ronda 33156-4299

U.S.A.

ABSTRACT

Newly discovered localities and their somewhat different habitats are
described for the rare Florida endemic Hasleola robertiorum. The basic species
description is modified to include minor variations found in these new
populations.

KEY WORDS: Hasteola robertiorum, Asteraceae, Rorida, range extension,
morphological variation

DISCUSSION

The recently described //ojteo/a robertiorum L.C. Anderson is a very rare Ronda
endemic related to the more widespread H. suaveolens (L.) Pojarkova, formerly
known as Cacalia suaveolens L. (Anderson 1994). Hasteola robertiorum was
descnbed from only a few populations in Levy County, Ronda. New populauons

246

247 PHYTOLOGIA volume 78(4): 246-248 Apnl 1995

were discovered in 1994 from somewhat different habitats in Lake County, Ronda, m
an area disjunct 120 km from the Levy County sites. Additionally, the species
descnpuon must be emended slightly to accommodate certain features found on plants
from these new pxjpulauons.

The abbreviated collection data of the Lake County populations are: soggy black
muck of seepage-saturated hydnc hammock along spnng-fed blackwater stream
(Sulphur Run) in Seminole Slate Forest, ca. 1 air km SE of Lake Jordan, ca 25 air km
ENEof Eustis, 18 Oct 1994, S.L. Orzell. E.L. Bridges, & G. Reese 23332 (FLAS,
FSU,FTG,NY,TEX,USF); soggy, often quaking, deep black muck of hydnc
hammock along Sulphur Run, Seminole State Forest, ca. 0.8 air km SE of Lake
Jordan, ca. 24 air km ENE of Eustis, 18 Oct 1994, S.L. Orzell, E.L. Bridges. & G.
Reese 23336 (FSU,FTG,TEX,USF).

Many thousand plants of Hasteola robertiorum are found over a distance of at least
one km in the Sulphur Run hydnc hammock. The two collection sites are firm to
quaking muck microhabitats that are scattered within a blackwater creek valley hydnc
hammock. There are some isolated areas of saline seepage; the surface soil pH ranges
from generally 6.8 to 7.2 (near saline seeps). The canopy of the quaking muck site is
dominated by Magnolia virginiana L., SatxU palmetto (Walt.) Lodd. ex Schult. &
Schult., and Acer rubrum L., with a tall shrub layer of Cornus foemina Mill., Myrica
cerifera L., and Leucothoe racemosa (L.) A. Gray. The canopy dominants of the
firmer muck site are Sabai palmetto, Quercus laurifolia Michx., and Tilia caroliniana
Mill., with a subcanopy-shrub layer of Carpinus caroliniana Walt., Agarista
populifolia (Lam.) Judd, and Myrica cerifera. Herbaceous taxa found at both sites
include: Boehmeria cylindrica (L.) Sw., Carex leptalea Wahl., Dryopteris ludoviciana
(Kunze) Small, Mikania cordifolia (L./) Willd., Oplismenus setarius Lam., Osmunda
cinnamomea L., Panicum commutatum Schult., Rhynchospora miliacea (Lam.) A.
Gray, and Thelypteris palustris Schott. An additional 63 vascular plant species were
associated m\h Hasteola in at least one microhabitat.

Rare species found at one of both of the Lake County sites include Carex
chapmanii Steud., Cirsium muticum Michx., Rhapidophyllum hystrix (Pursh) Wendl.
& Drude, and Salix floridana Chapm.; of these, only Carex chapmanii is found at the
Levy County sites. The relatively drier Levy County sites also differed noticeably in
that they had very few Sabal palmetto, which was abundant in the Lake County sites.

Lake County plants (especially those of Orzell et al 23332) are generally more
robust than those from Levy County. Plants are up to 15 dm tall with basal (radical)
leaver- up to 54 cm long with petioles up to 32 cm long. Lower cauline leaves are up
to 38.5 cm long and 16.5 cm wide. These measurements surpass those of any wild-
collected Levy County plants (Anderson 1994), but the progeny of Levy County
plants thai were garden-grown by Anderson in Tallahassee were even larger. Lake
County plants also have more purple pigmentation on lower stems and peuoles, and
the prominent basal auricles on some cauline leaves measure up to 9 mm long.

Involucres on Lake County plants have average lengths for the species, but the
phyllaries number only 7-8, and the 4-7 subtending bracteoles are only 4-6 mm long.
Flowers are fewer (8-10 per head). Corollas are average in length, but the tubes are
generally longer (5.0-5.4 mm long), the throats concomitantly shorter (1.7-2.4 mm
long), and the lobes 2.0-2.8 mm long. Anthers are somewhat shorter (2.0-2.5 mm
long). The balusterform anther collars and enlarged stylopodia are typical for the
species.

Achenes are shorter (6-7 mm long), and the pappus somewhat longer (5-6 mm
long) on Lake County plants compared to 7-9 mm long and 4-5 mm long,
respectively, for Levy County plants. The carpopodium on Lake County plants has 4-

Andersons/ a/.:

Distnbution of Hasteola 248

6 rows of disuncuve square to procumbent cells; those of Uvy County plants

'°"rnTene^''LTnvolucr:U character of the Lake County plants, f^cularly
phyllar^ number and number of Rowers per head, expand the vanauon of HasteoUi
robertihrum away from the related species. H. suaveolens, prov.dmg further evidence
of its distinctness. However, Horal characters generally indicate more overlap in
noral measurements between H. robertiorum and H. suaveolens^ had previously
been known. This is not surpnsing considering that H robertiorum had been
descnbed from only a few populations in a very narrow local area, ^^ the Lake
County populations may have been isolated from those in Levy County or a
siRmficam penod. Nevertheless, the consistent trend in reducUon of phyllary number
aifd number of flowers per head in Flonda Hasteola suggests that they are remnants of
a once more continuous range. . ,

The Lake County plants, though slightly different in some aspects of morphology
from those of Levy County, are not sufficiently distinctive to warrant special
tSonon^c rm^gmtion as a'vanety or subspecies^ The discovery of Hj^^o^
robertiorum in Uke County has greaUy enlarged the known range for this species (the
species being more abundant here than in Levy County), but it still remains a very- rare
endemic. The Ronda Endangered Plant Advisory Council has recommended that it be
listed as "endangered" in Florida

ACKNOWLEDGMENTS

We wish to acknowledge Gary Reese, who discovered the hydnc hammock sites
in Lake County as part of a natural areas inventory of Uke County conducted wth
fun£ providL byTe Lake County Water Authonty and the St. John's River Water
fZiglment District He also provided assistance in the field with data collo^t.on
andin momtonng the Hasteola phenology. T.M. Barkley and M.O. Moore kindly

reviewed the manuscript.

LITERATURE CITED

Anderson, L.C. 1994. A revision of //a5teo/a(Asteraceae) m tiie New World. Syst.
Bot. 19:211-219.

Phytologia (Apnl 1995) 78(4) 249-255.

CAREX INVERSA R. BR. (CYPERACEAE), NEW TO HAWAII AND THE
UNITED STATES

Robert B. Shaw & Patricia P. Douglas

Center for the Ecological Management of Military Lands, Department of Forest
Sciences, Colorado State University, Fort Collins, Colorado 80523 U.S.A.

ABSTRACT

Carex inversa R. Br., a rhizomatous perennial species endemic to .Australia
and New Zealand, is reported for the first time from the Hawaiian Archipelago.
The taxon was discovered in an area known as the Kipuka Alala near the center
of the island of Hawaii at an elevation of approximately 1675 m (5500 ft). The
species occurs in Styphelia Mixed Shrubland, Dense Dodonaea Shrubland, and
Sparse Metrosideros Montane Treeland plant communities. Two other
introduced Australian species [Lepidium hyssopifolium Desv. and Crassula
sieberiana (Schult.) Druce] are restricted to the same general area as Carex
inversa. The occurrence of the taxon probably represents an inadvertent
introduction by domestic livestock or in grass seed mixtures. Long distance
transport by wind or birds is another possible but highly improbable mode of
introduction for the species. Rather large clones, some up to 4 m in diameter,
and occurrence of the taxon in several diverse habitats suggests that the original
introduction may not be recent.

KEY WORDS: Carex, Hawaii, Australia, New Zealand, introduction

The first taxonomic revision of the Carex of the Hawaiian Islands listed nine
species which were all thought to be native (Krauss 1950). The most recent revision
of the carices recognizes eight species from the Islands (Koyama 1990); however,
only four species are listed as endemic and four are indigenous. The geographic
affinities for the indigenous species are: one with Eurasia, one with Europe and
Pacific North America, one with the Caroline Islands, and one possibly with eastern
Asia.

Carex inversa R. Br. (Figures 1, 2), commonly known as knob sedge, is being
reported for the first time from the Hawaiian Islands. This taxon is a member of
subgenus Vignea and section Inversa. It is a perennial herb (0.25-0.75 m high) that
occurs frequently in clonal swards because of its spreading rhizomes. Culms are
slender and leaves are basal as well as cauline. The leaf blades are linear (1-4 mm
wide) and involute at the tip. The leaf sheaths tightly clasp the stem and project
beyond the juncUire with the blade producing a membranous appendage (conira-
ligule). The gynecandrous inflorescence is composed of 2-3(-4) short, ovoid terminal
spikes (5-15 mm long). The lowermost spike is subtended by leaf-like involucral

249

Shaw & Douglas: Carex inversa in Hawaii 250

bracts, at least one of which surpasses the entire intlorescence. Each pengymum is
subtended by an ovate, short-awned scale (2.0-3.0 mm long). The midnbs of the
scale are prominent and pale green, and margins are broad, translucent and entire.
Male flowers have 3 stamens (1.9-2.1 mm long), and the anthers are 2-celled ( 1. 1-1.6
mm long, including the anther hairs) with numerous basally-fused hairs present at the
anther's ape.x. Female flowers are composed of a single pisul with 2 stigmas. The
pistil IS contained within a beaked pengymum (~ 3 mm long) and the branches extrude
from a small bifid onfice at the apex. The pengymum is slightly winged above the
middle, and the margin is finely serrate. Abaxial and adaxial faces are strongly veined
(9-11 and 6-8, respectively). The achene is flattened, lenticular and reaches - 2 mm in
length at matunty. Our collections were venfied by companson with specimens from
Australia (US 31117, 1349954, 2072356, 2072359, 2072373, 2126591, 3165854)
and New Zealand (US 1239957, 3165853, 3165855, 3203467) at the US National
Herbanum and by Drs. Anton A. Reznicek (University of Michigan) and Karen
Wilson (New South Wales Herbanum). Bentham (1878), Moore & Edgar (1970),
and Black (1978) all listed vaneties for C inversa; however, most recent usage
suggests that separation into intraspecific categones is unwarranted without further
study.

Carex inversa grows in the Australian states of Queensland, New South Wales,
Victoria, Tasmania, South Australia, and West Australia. The taxon also is reported
from both the North and South Islands of New Zealand. Burbidge & Gray (1970)
described the plant as a weedy sedge which forms dense patches or swards in poorly
sown or modified nattiral grasslands. In New Zealand tussock grassland
communities, C. inversa grows in association with Poa caespitosa Spreng., Festuca
novae -zeaUmdiae (Hack.) Cockayne, Danthonia gracilis Hook., Danthonia unarede
Raoul, and Dichelachne crinila (L.) Hook. (Cockayne 1967). Burbidge & Gray
( 1970) and Beadle et al. ( 1972) reported the species from Eucalyptus woodlands and
swampy areas or creek banks. Within its range, the taxon is relatively common and
widespread in mesic sites of localized communities. Healy & Edgar ( 1980) r£|X)rted
that C. inversa is a troublesome weed in lawns, paths, and rockeries. Because of its
rhizomatous habit, C. inversa may be invasive and difficult to eradicate (Willis 1962).
It appears to be tolerant of grazing and may be of value in providing a limited tinount
of fair quality forage when it occurs in quantity (Leigh & Malham 1965). The plant
also appears to be resistant to trampling and mowing.

Carex inversa was discoveraJ during fionstic inventones on the Ponikuloa
Training Area (PTA), Hawaii (Figure 3). PTA is the largest U.S. Army installation in
Hawaii, over 43,700 ha ( 108,000 ac) in size, and is situated in the saddle betwet.n two
volcanoes (Mauna Kea and Mauna Loa) which are over 4200 m (13,800 ft) in
elevation. Approximate average annual precipitation for the saddle area is 39 cm (15
in) [data from Bradshaw Army Airfield which is 14 km (9 mi) northeast and 200 m
(650 ft) higher in elevation than the collection sites for C. inversa]. Substrate on the
installation is composed of numerous younger (< 10,000 year old) Mauna Lea lava
flows and some older (> 10,000 year old) Mauna Kea flows. The diversity in lava
substrates combined with the fairly wide altitudinal gradient gives nse to a complex
mosaic of plant communities. Castillo el al. (1994) have delineated 24 plant
communities on the installation. Shaw & Douglas (1995) have documented ten
endangered or threatened species, five Category 2 candidate species, and one
presumed extinct species on the installation. Carex wahuensis C.A. Mey. subsp.
wahuensis dixxd subsp. rubiginosa (R. Krauss) T. Koyama are the only other members
of the genus found on the installation.

251

PHYTOLOGIA volume 78(4):249-255 Apnl 1995

FIGURE 1. Carex inversa R. Br. (Douglas et al. 4618) a. general habit illustrating extensive
rhizome development, b. inflorescence composed of three spikes, and c. single qynecandrous
spike

Shaw & Douglas:

Care.x inversa in Hawaii

O^T

FIGURE 2. Carex inversa R. Br, (Douglas et al. 4618) a. Male flower with insert showing
terminal anther hairs, b, perigynium with bract, c. abaxial face of the perigynium, and
d. leticular achene

253

PHYTOLOGIA volume 78(4):249-:55

Apnl 1995

U.

.^'

h

^•i

i

i

h

I

V

Shaw & Douglas: Carex inversa in Hawaii 254

Carex inversa was first collected near and m Kipuka Alala, on the southwestern
portion of the instaJlaUon (Figure 2) (Douglas el al. 3507 [MICH], 3508 [BISH],
4612 (CS], 4618 [CS], 5045 [RM], 5048 [CS)). A kipuka is an older and more
heavily vegetated lava flow that is surrounded and isolated by younger Hows with
speirser vegetation. Kipuka Alala is approximately 1750 ha (4325 ac) in size and the
substrates are Mauna Loa lavas ranging in age from 1500 to > 4000 years old
(Lx>ckwood el al. 1988). Vegetation within Kipuka Alala consists of Dodonaea,
Styphelia, and Myoporum dominated shrublands (Castillo el al. 1994). Adjoining
younger flows (< 1500 years old) support open Metrosideros treelands.

Carex inversa is rare to scattered in the following plant commumties: Siyplielia
Mixed Shrubland, Dense Dodonaea Shrubland, and Sparse Metrosideros Treeland
(Castilloes a/. 1994). Plants occur in soil and/or ash accumulations which are slightly
more mesic than the surrounding landscape. The taxon generally grows in full sun.
Plants were observed flowering dunng the wet period of the year (NovemlDer to
February), which differs from the flowering penod for Australian and New Zealand
populations (October to April) (Marchant el al. 1987). Plants grazed by feral sheep
and/or goats were seen; however, no adverse effects of grazing were noticed.

A vascular plant survey of the installation documented 249 species growing on
PTA (Shaw & Douglas 1995). Approximately 61% (157 species) of these taxa are
naturalized. Nineteen taxa are introduced species from Australia and/or New Zealand.
Three Australian introduced grass species [Danthonia pilosa R. Br., Ehrharla slipoides
Labill., and Eragroslis brownei (Kunth) Nees ex Steud.] are common within the area.

There are two other examples of Australian or New Zealand species introduced
only into the saddle area and PTA (Shaw & Douglas 1995; Wagner el al. 1990).
Lepidium hyssopifolium Desv. (Brassicaceae) was first found in 1975 in the .saddle
area and was known from only two collections unnl recently venfied from PTA
(Wagner et al. 1990). Crassula sieberiana (Schult.) Druce (Crassulaceae i is a
widespread interstitial species throughout the installation. Wagner el al. (!990)
reported that this taxon was first collected in 1978 and apparently was a recent
introduction.

Several explanations exist for the occurrence of these Australian species only in the
saddle region of the island of Hawaii. First, a single natural event may have biought
the species to the saddle area {i.e., storm or birds). Second, the species were
introduced by humans in forage seed mixtures, by herds of grazing animals, or by
military vehicles. Third, the introduction of these taxa may be completely unrelated
and represent separate events. FYior to the mid 1970's, relatively few introduced taxa
were reported from the saddle region which suggests the region was either still fairiy
pristine at this time or pooriy collected. If the former is true, then the Ausiralian
species may indeed be rather recent introductions. Conversely, the large clones
indicate that Carex inversa may have occurred at PTA before this date. Studies are
ongoing to examine the genetic vanability throughout the species and to determine
similarities between Hawaiian and Australian/New Zealand matenal. These data
should help determine the ongin of the Hawaiian populations as well as the
approximate time of introduction.

ACKNOWLEDGMENTS

This research was funded by the Department of Defense, Legacy Resource
Management Program; U.S. Army Support Command Hawaii (USASCH); and U.S.

255 PHYTOLOGIA volume 78(4): 249-255 Apnl 1995

Army Corps of Engineers, Pacific Ocean Division. The field assistance of J M
Castillo, C. Hindes, M. McFadden, E. O'Regan, B. Painter, C. Popolizio and t'
Tiemey is appreciated. Special thanks to Tracy Wager for the excellent line drawing of
the species. We are very grateful to Drs. Anton A. Reznicek (University of Michigan)
and Karen Wilson (New South Wales Herbanum) for venf.caUon of thTsSs
.dentificauon. Also, review of the manuscnpt by Drs. Reznicek, TimotLy I^^ey
(University of New Mexico), Richard D. Laven (Colorado State University^ I^d
Derral Herbst (U.S. Army Corps of Engmeers) is appreciated.

LITERATURE CITED

Beadle N.C.W.. O D. Evans, & R.C. Carolin. 1972. Flora of the Sidney Region
Sydney. Australia. A.H. and A.W. Reed Dty Ltd t^egwn.

Bentfiam.G. 1878 (repnnt 1967). Flora Australiensis: A Description of the Plants of

Black"TM ,?7?'^'?, ^°'- 7i'- \ ^'''' ^'^ ^°- ^"^«"' Great Bntain. ^

;^ni^Q.^ A ^^r"" i.^?"'} ^^^<^^^ P^ I. Third Edition. Government
Printer, South Australia, Adelaide, AusU^ia

Burbidge, N.T^ & M. Gray. 1970. Flora of the Australian Capital Territory
Australian National University Press. Canberra, Australia armory.

PaHk^^L' J-^- '^'T^^' S ^-^^ ^^^- PJant Commumues of the

Pohakuloa Training Area. Hawaii, Hawaii. Map. Center for the Ecoloacal

aTrldo""' ""^ ^'''^ ^"^"^ ^^'^^'^^ ^'^^ University' Fort CduS

^'^i£r^:i^l^^n'^^^ ^•^- ^-- G-er^nent

Healy. A.J. & E. Edgar. 1980. Fbra of New Zealand. Vol. III. Adventive

Cyperaceous Petalous and Spathaceous Monocotyledons. PD Kasselbere

Government Pnnter, Wellington. New Zealand ' '^**^^^"^'^8'

^°^i^'^J^/ ^/^"f,^ ^"- W^8"er. W.L., D.R. Herbst. & S.H. Sohmer

i^n u^ of the Flowering Plants of Hawai'i. Umversity of Hawaii pVess

and Bishop Museum Press, Honolulu Hawaii

"^"a^x L,nc1!;i 4^^282™' """°" °' "'^ "^"""" ^^'" °^ ^' ^^"-
^tcalild^^s^B^STAi^a ^^^ ^^^ ^ '^ ^^^^rine Pla.n. The

^cZr^v •'h^- ^-^c- i'P"^^' ^•'^- P^^^'-^^"' & FR- Warshauer. 1988
Generalized ages of surface lava flows of Mauna Loa Volcano. Hawaii. USDI '
US. Geological Survey, Misc. Investigations Senes, Map 1-1908

Marchant. N.G., ^ Wheeler, B.L. Rye. E.M. Bennett. N.S. Lander & TD

HeTtSur-Denfof a'"''", '^ t ^"'^ ^^^'^''- ^^^^ "" ^^^^^ ^llliSi
Herbanum Dept. of Agriculture. Western Australia. Australia

Moore, L.B. & E. Edgar. 1970. Flora of New ZeaUmd. Volume II Indigenous
Shaw R S'P^.^P p^-,?;^*^.^"^' Government Pnnter. Wellington. New ZeS

P^h^ I V-P-^"^'^- ^^^- ^^^^ Pl^t Inventory of the U S Arniv

Pohakuloa Traimng Area. Island of Hawaii. Hawaii IN PREP ^

Wagner W.L D.R Herbst, & S.H. Sohmer. 1990. Manual of the Flowering

^1"^: ^"'^^^^'^^ ^^ "^-^' ^-^ -'^ Bishop Lseum Pr^^^
W.Ihs James H. 1962. A Handbook to Plants in Victoria. Volume I. Ferns Conifers

and Monocotyledons. Melbourne Umversity Press. Canbena, Australia

Phytologia (Apnl 1995) 78(4)256-259

NEW RECORDS AND NOTES CONCERNING CASTILLEJA SPIRANTHOIDES
(SCROPHULARIACEAE)

Mark Egger

C/O Herbarium (WTU), Department of Botany, KB- 15, University of Washington,
Seattle. Washington 98195 U.S.A.

ABSTRACT

Newly identified collections of the little-known Castilleja spiranthoides are
documented, and a summary of its morphology, ecology, distribution and
possible relationships is provided. This species appears to be an anomalous
member of sect. Castilleja, with its nearest relative probably being Castilleja
ctenodonta.

KEY WORDS: Castilleja, Castilleja spiranthoides, Castilleja ctenodonta,
Scrophulariaceae, chromosome numbers, Sinaloa, Mexico

Castilleja spiranthoides Standley is a distinctive but little known taxon previously
reported only from the type collection (Ortega 6896 [F!]), made in the vicinity of Los
Gusanos, San Ignacio in south-central Sinaloa. Its description by Standley (1936)
contains only the Latin description, the collection number, the location cited above and
the sparse collection notes, "in dry soil, March 1931". Since that time references to C.
spiranthoides appear to be absent from the botanical literature.

Recently, a review of undetermined collections of Mexican Castilleja from the
herbaria of ASU and ARIZ revealed three additional collections of Castilleja
spiranthoides, extending its known range at least 70 km to the southeast. More
importantly, they provide additional insight into the morphology and possible
relationships of this rather unusual species within Castilleja.

The three newly identified collections, all from Sinaloa, Mexico are as follows:

1. Near El Batel," along highway from Mazatldn to Durango, mixed oak and pine
forest with steep southeriy slopes, elevation 5,000 to 6,000 ft., 6 February 1952,
Gentry 11562 (ARIZ!). The specimen label also contains the notauon, "annual with
red Oowers". This sheet holds four complete plants. Unpublished collection records 11
in Gentry's hand list the collection date as 5 February 1952 and include the locality '
notation of "Palmito-Batel, Sinaloa" (P. Jenkins, ARIZ, pers. comm.). A duplicate of
this collection is deposited at MICH and was independently identified as Castilleja j,
spiranthoides by G.L. Nesom in 1992 (Nesom, pers. comm.). I

256

Egger Notes on Casiilleja spiranthoides 257

2. Just west of El Palmito, Rancho El Liebrd, pine forest hillside and deep
barranca, elevation ca. 7,000 ft., 13 March 1980, Uhlo 24384 (ASU!). This sheet
holds four complete plants and two partial stems.

3. About I mi. north of Mexican Highway 40, between Villa Union and Ciudad
Durango, about 5 miles west of Durango state border, elevation ca. 7,600 ft., 14
March 1985, Darnel 4028 (ASU!). This sheet holds two complete plants. The
specimen label also provides the notation, "Bracts reddish; occasional."

In addition, a fourth new collection of Casiilleja spiranthoides is deposited at
UCR, according to notes and photographs of the collection on file at UC(!). This
sheet was annotated as C. spiranthoides by L.R. Heckard in 1992. The onginal
citation is as follows: Rancho Libre Barranca, 2 air miles northwest of H Palmito,
north of Hwy. 40 £ind very near the Durango state line, 27 March 1984, Sanders
4903. Additional notes by the collector indicate that the plants occurred in cloud forest
on north slopes in pine/oak vegetation on exposed ridges and that the plants were
"fairly common in duff under pines". The annotation by Heckard also notes that the
calyx is divided more deeply in front than behind and that the lobes of the lower lip of
the corolla are infolded.

While the Latin description by Standley appears to accurately represent the plants
in the newly identified collections, a few additional and potentially diagnostic
characters are apparent in these plants. The plants represented in the collections of
Lehto and Daniel are vigorous specimens in prime flowering condition, and the
measurements from these plants for all vegetative and flowering parts are consistently
and proportionately larger than are those found in the holotype or in the Gentry
collection, which appear to possess somewhat defjauperate and/or desiccated
inflorescences. Measurements from the descnption by Standley (1936) of the type
collection more closely resemble those of the Gentry collection. Whether these
differences are due to ecological conditions or to natural variation among healthy plants
is unknown. However, it is clear that all five of these collections are of a single
species. The following notes are based upon Standley's description combined with
data from the newly identified collections and are intended to supplement Standley's
type descnption of Castilleja spiranthoides:

Plants annual; stems 25-35 cm tall, mostly simple, erect, slender and densely
villosulous with gland-tipped hairs; leaves numerous, crowded and ascending below,
becoming shorter, widely spaced, erect, and with a finely sinuate-margined apex
above, 15-50 mm, entire, linear and narrowly acuminate, sessile to subclasping but
not at all auriculate, villosulous with gland-tipped hairs; infiorescence at first a
subcapitate spike, the intemodes elongating with age; bracts (15-)20-24 mm long,
usually entire (occasionally with several short, acuminate, apiculate teeth) greenish
basally, becoming pink to light red-purple above with prominent dark red-purple
veins, blundy ensiform to narrowly pandurate with a rounded, sinuate-margined,
petaloid tip, moderately to densely villosulous with gland-tipped hairs; calyx (10-)18-
27 mm long, entirely pink to red-purple with prominent dark red-purple veins,
sfjarsely stipitate-glandular, tube 2-3 mm wide for most of its length, then widening to
3-5 mm near the base of the lobes, abaxial cleft 12-16 mm long, adaxial cleft 6-8 mm
long, with secondary clefts 2-4 mm long, narrowly acuminate, slightly unequal, often
falcately upturned; corolla (l2-)20-30 mm long with tube and beak of abJout equal
length, only slightly exceeding the calyx in length but often at least partially exserted

258 PHYTOLOGIA volume 78(4):256-259 Apnl 1995

outward from the abaxial calyx cleft, mostly glabrate, tube pale, beak becoming
entirely reddish except for a pale green and shortly bearded dorsal surface, lower lip of
three dark greenish, incurved teeth, 1.0-1.5 mm in length.

Castilleja spiranthoides appears to flower dunng most of February and March,
although the full duration of the flowering penod has yet to be determmed. Little is
known of its ecology, although it is now known to occur in pine-oak forests on sleep
slopes between 1,500-2,300 m. It appears to be a rare endemic to the western slopes
of the Sierra Madre Occidental in southeastern Sinaloa, but it likely occurs in adjacent
western Durangoas well. The locations of all four of the newly idenufied collections
cited above are within a few kilometers of each other in the vicimty of the botanically
well-explored highway from Mazatldn to the Durango border. Also, it seems likely
that the range of C. spiranthoides may be more or less continuous from the vicimty of
the type collection to the area of the new collections, as the intervening mountainous
terrain provides similar habitat but is relatively inaccessible and poorly known
botanically.

The relationships of Castilleja spiranthoides within the genus remain somewhat
unclear. Based on coloration and morphology of the inflorescence, this species is
surely a member of subg. Castilleja, as defined by Chuang & Heckard ( 1991). More
equivocally, its irregularly cleft and conspicuously colored calyx, somewhat
pendulously exserted corolla, and relatively long corolla beak appear to place C.
spiranthoides within sect. Castilleja, as defined by Holmgren (1976) and by Nesom
(1992). However, neither of these authors includes C. spiranthoides among the sect.
Castilleja found within their area of treatment. This apparent oversight may be due to
the paucity of specimens of this species previously available for comparative study.
Nesom (pers. comm.) now considers C. spiranthoides to be a member of sect.
Castilleja, based on calyx and corolla morphology.

While it does appear to belong to sect Castilleja, Castilleja spiranthoides is a rather
anomalous member of that group, with the only other annual species apparently being
Castilleja fili/lora Nesom, a limited endemic from Chiapas, Mexico (Nesom 1992) that
is unlike C. spiranthoides in a number of morphological features. The species to
which C. spiranthoides is perhaps most closely related and which it most resembles in
form, leaf arrangement, vestiture, coloration, and numerous features of the
infiorescence is the rarely collected C ctenodonta Eastwood. Apparently, C.
ctenodonta is known only from a few collections made in the vicinity of the type
locality in the Sierra de Clavellinas in Oaxaca, although, as Nesom ( 1992) pointed out,
C ctenodonta is very closely related also to the Guatemalan endemic species, C.
altorum Standley & Steyermark. Eastwood's (1909) descnption of the type of C.
ctenodonta indicates that this is a perennial species, although its stems are very slender
and apparently rhizomatous in the specimens I have seen, including an isotype of
Pringle 4986 (UC!) and two sheets of Smith 5i9 (NY !,UC!). CastUleja ctenodonta
differs from C. spiranthoides most strongly in that both the leaves and bracts of C.
ctenodonta have short, narrow, nearly pectinate divisions and that the bracts are not at
all apically pandurate.

Some aspects of the morphology of Castilleja spiranthoides are atypical of sect.
Castilleja. In such features as its annual duration, basally clumped but then more
widely spaced, upright to erect cauline leaf arrangement, often sinuate-margined upper
leaves and bracts, and often pandurate bract shape, C. spiranthoides more closely
resembles some species of sect. Euchroma (Nutt.) Benth., such as C. rnacrostigrna
Robinson and C. ornata Eastwood. However, the calyx and corolla morphology of C.
spiranthoides appear to preclude its placement in that group. Both additional field

Egger Notes on Castilleja spiranihoides 259

work, and chromosomal and biochemical analyses of this species would be desirable
in sorting out its evolutionary relationships.

ACKNOWLEDGMENTS

I wish to thank Guy Nesom (TEX) and Margriet Wetherwax (JEPS) for helpful
reviews of the manuscript, Phil Jenkins (ARIZ) for providing unpublished field notes
of H.S. Gentry, and the staffs of ASU and ARIZ for the loan of the specimens upon
which this note is pnmarily based.

UTERATURE CITED

Chuang, T.I. & L.R Heckard. 1991. Generic realignment and synopsis of subtnbe

Castillejinae (Scrophulariaceae -- Tribe Pediculareae). Syst. BoL 16(4):644-666.
Eastwood, A. 1909. Synopsis of the Mexican and Central American species of

Castilleja. Proc. Amer. Acad. Arts 44:565-591.
Holmgren, N.H. 1976. Four new species of Mexican Castilleja (subgenus Castilleja,

Scrophulariaceae) and their relatives. Brittonia 28: 195-208.
Nesom, G.L. 1992. Taxonomy of the Castilleja tenuiflora group (Scrophulariaceae)

in Mexico, with an overview of sect Castilleja. Phytologia 73(5):389-415.
Standley, P.C. 1936. Studies of Amencan plants ~ VI. Field Mus. Nat. Hist. Bot.

Ser. 11(5): 145-276.

Phytologia (Apnl 1995) 78(4)260-263

LITHOCYSTS AS TAXONOMiC MARKERS OF THE SPECIES OF CORDIA L.

(BORAGINACEAE)

B. Hanumanlha Rao & K. Vjaya Kumar
Department of Botany, Andhra University, Visakhapatnam - 530 003, A. P., INDIA

ABSTRACT

The structure and distribution of lithocysts in the foliar epidermis of eight
species of the genus Cordia of Boraginaceae are studied. These are mostly
distributed on the adaxial surface of the leaf and are restricted to the epidermis
only. The qualitative and quantitative characteristics of these cystolith containing
cells are found to be useful as taxonomic markers in the identification of the
various species of Cordia.

KEY WORDS: Cordia, Boraginaceae, foliar epidermis, lithocysts

INTRODUCTION

Lithocysts are the cystolith containing cells. These are situated in the epidermis of
leaf, usually on the adaxial surface and occasionally on both the surfaces. A perusal
of the so far available literature (Solereder 1908; Metcalfe & Chalk 1950, 1979, 1983)
clearly reveals that the information available on this aspect in the family Boraginaceae
is highly limited. Therefore, in the present investigation, eight species of Cordia are
studied with emphasis on the structure and distribution of lithocysts.

MATERIALS AND METHODS

Fresh leaf material of Cordia alba L; C. dichotoma Forst.; C. monoica Roxb.; C.
sebestena L., and C. waltichii G. Don were collected and fixed in Formalin- Aceto-
Alcohol, whereas herbarium specimens were secured for C. domestica Roth., C.
evolutior Gamble, and C. macleodii Hook. /. & Thomps. For the latter, the material
was initially rehydrated by boiling in water. Whole mounts, epidermal peels and
transverse sections of the leaf were prepared using traditional methods and microtomy.
The frequency, were prepared using traditional methods and microtomy. The
frequency, distribution, and size (length and width in surface view and depth m
sectional view) were recorded. The presence of calcium carbonate in the cystoliths,
was confirmed adopting methods of Jane ( 1970).

260

Rao & Kumar:

Lithocysts in Cordia

261

OBSERVATIONS AND DISCUSSION

Lithocysts are observed in all these species of Cordia (Figures 1-8). The
quantitative features of them are presented in Table I. These are distributed in the
costal and intercostal regions of both the epidermal layers in C. dicholoma and C.
sebestena, whereas these are restricted to the adaxial surface only in the remainder of
the species. The lithocysts are usually larger than the adjacent epidermal cells and
protrude into the mesophyil region in the form of cell cavities (Figures 5-8). In
surface view a nng of radiating epidermal cells encircles them, giving the total
structures the appearance of trichome bases (Figures 1-4). But the secUonal view
confirmed the presence of cystoliths inside their cell cavities (Figures 5-8). Sometimes
the cystoliths protrude above the general surface giving a papillate appearance as in C.
monoica, C. dichotoma, and C. macleodii. A limited number of epidermal cells
encircles the lithocysts in C. evolutior and C. sebestena, (Figures 2,4) whereas
numerous small epidermal cells surround them in C. domestica (Figure 1). In C.
wallichii, lithocysts form a characteristic feature of the surface morphology with
deeply stained encircling cells giving a rosette appearance to them. The maximum
frequency of the lithocysts is recorded in C. wallichii (28 per mm^). The largest and
smallest lithocysts are observed in C. monoica and C. evolutior (Figure 7) to
ellipsoidal as in C. wallichii (Figure 8). Their surface is uniformly verrucose ( Figures
5-8).

Table 1. Quantitative characteristics of lithocysts in different species of Cordia in the
present investigation.

Species
Number

Species Name

Frequency
(per mm^)

Length
(urn)

Width
(^m)

Depth
(Jim)

1.

Cordia alba

6

95.40

76.32

43.29

2.

C. dichotoma

27

81.40

71.86

58.17

3.

C. domestica

12

78.86

71.23

64.00

4.

C. evolutior

14

70.40

64.05

56.00

5.

C. macleodii

7

208.29

195.57

73.26

6.

C. monoica

20

77.59

66.48

7.

C. sebestena

3

61.05

61.05

52.16

8.

C. wallichii

28

77.59

70.73

69.25

The lithocysts in Cordia alba are less specialized with isolated groups of cells in the
upper epidermis having thicker outer walls from which knob-like processes
impregnated with calcium carbonate project into the cell cavity. In addition to the
original lithocyst, adjacent epidermal cells around it may also contain cystolith-like
bodies as in C. macleodii (Figure 3). Such groups of lithocysts were earlier reported
in the family Opiliaceae (see Mauseth 1988).

The lithocysts containing the cystoliths are generally treated as the excretory bodies
with reference to their function. Haberlandt (1914) stated that these excretory
structures become transformed into rejxsitories of reserve matenals and the lime is

262

PHYTOLOGI A

volume 78(4):26()-263

Apnl 1995

■

ECO

E
E
^^

Et^

£
E

o

6 -

o

OnT

o

.

Figures 1 -8. Li thocysts of Corrf/a. 1-4. Surface view. 1. C. domestica\ 2. C,
evolution 3. C. macleodii; 4. C. sebestena. 5-8. T.S. of adaxiaJ epidermis. 5. C.
alba\ 6. C. dichotoma; 7. C. monoica, 8. C. wallichii.

Rao & Kumar: Lithocysts in Cordia 263

reintroduced into the metabolic cycle. Bider ( 1935), on the other hand, considered that
the species which possess these lithocysts and related bodies do not flounsh unless
lime is present in the soil. Therefore, species of Cordia may be useful as indicators of
lime in the soil.

On the basis of a few available characteristics of lithocysts of the various species of
Cordia studied here, a tentative key is presented.

1 . Lithocysts distributed on both surfaces of the leaf 2

2. Lithocysts small and less frequent C. sebesletia

2. Lithocysts large and more frequent C. dichotoma

1. Lithocysts distnbuted only on the adaxial surface of the leaf 3

3. Lithocysts not protruding into the mesophyll C. alba

3. Lithocysts protruding into the mesophyll in the form of deep cavities 4

4. Lithocysts in groups C. macleodii

4. Lithocysts solitary 5

5. Lithocysts surrounded by numerous small epidermal cells

C. domestica

5. Lithocysts surrounded by limited number of large epidermal cells 6

6. Cystoliths ellipsoidal C. Wallichii

6. Cystoliths spherical 7

7. Stalk of the cystolith is wide C.monoica

7. Stalk of the cystolith is narrow C. evolutior

Thus the variability expressed in different qualitative and quantitative features by
the lithocysts and the cystoliths in the foliar epidermal cells of the species of Cordia
provide useful taxonomic markers in the infrageneric classification of Cordia.

UTERATURE CITED

Bider, J. 1935. Beifrage Zur pharmakognosie der Boraginaceen und Verbenaceen

Vergleichende Anatomia de Laubblatem. Thesis, Basel, Switzerland. 124 pp.
Haberlandt, G. 1974. Physiologiccd Plant Anatomy, (translated from the Fourth

German edition by Montagu Drummond). Macmillian, London, United Kingdom.
Jane. F.W. 1970. The Structure of Wood. Adam and Charles Black, London,

L'nited Kingdom.
Mauseth, J.D. 1988. Plant Anatomy. Benjamin Publishing, Inc. California, U.S.A.
Metcalfe, C.R. & L. Chalk. 1950. Anatomy of the Dicotyledons. Vol. I, Clarendon

Press, Oxford, United Kingdom.
Metcalfe, C.R. & L. Chalk. 1979. Anatomy of the Dicotyledons. 2nd ed.. Vol. I.

Clarendon Press, Oxford, Umted Kingdom.
Metcalfe, C.R. & L. Chalk. 1983. Anatomy of the Dicotyledons. 2nd ed., Vol. II.

Clarendon Press, Oxford, United Kingdom.
Solereder, H. 1908. Systematic Anatomy of Dicotyledons. Vols. I and II.

Clarendon Press, Oxford, United Kingdom.

Phytologia (Apnl 1995) 78(4) 264-276.

A FLORISTIC SURVEY OF FALLS HOLLOW SANDSTONE GLADES,
PULASKI COUNTY, MISSOURI

John Hays

Missouri Department of Conservation, Natural History Division, 2901 W. Truman
Blvd., Jefferson City, Missouri 65109 U.S.A.

ABSTRACT

The vascular flora of Falls Hollow sandstone glades, Pulasid County,
Missouri, is reported. A total of 137 sf)ecies is listed. Twenty-three new taxa
are added to the known vascular flora of Pulaski County. Three species
collected during this study are on the Missouri Department of Conservation's
list of Rare and Endangered Species.

KEY WORDS: Sandstone glade, flora, Missouri

INTRODUCTION

Missouri glades are open expanses of bare rock in non-prairie areas charactenzed
by a herbaceous flora, a lack or scattered occurrence of woody plants, and droughty
soils that are often seasonally saturated. The original objective of this study was to
survey the vascular flora of three sandstone glades at Falls Hollow on Fort Leonard
Wood Army Base, Pulaski County, Missouri, to determine their suitability for
nomination as a natural area by the Missouri Department of Conservation. These three
glades were located by Skirmer (1991), and I located a fourth glade during the present
study. The purpose of this paper is to present the results of a floristic survey of these
previously unbotanized sandstone glades in Missouri.

DESCRIPTION OF STUDY AREA

Falls Hollow glades are located approximately 17.7 km south-southeast of St.
Roberts, Pulaski County, Missouri, on Fort Leonard Wood Military Installation
(Bloodland Quadrangle, T34N, Rl IW. Sec. 22 NWl/4). Pulaski County lies within
the Upper Ozark Section of the Ozark Natural Division (Thorn & Wilson 1980). Falls
Hollow consists of four glades totaling 2.0 ha. The largest of these glades is 0.8 ha.,
whereas the three smaller glades are approximately 0.4 ha each. The aspect of the
glades at Falls Hollow is neutral and the slope is mostly flat to gentle. Each glade is
separated from the others by dry to mesic sandstone forest dominated by Quercus spp.
In the case of the three smaller glades a narrow ecotone exists between the forest and

264

Hays: Flonstic survey of Falls Hollow 265

the glades, with Vaccinium arboreum Marshall and Quercus mariUuidica Muenchh.
most common in this zone. The substrate of the glades at Falls Hollow is Roubidoux
sandstone (Ryan 1992; Wolf 1989), a medium to fine-grained, sparsely fossiliferous
sandstone that ongmated dunng the Ordovician Period (Koenig 1961). Although
sandstone occurs widely in the Missoun Ozarks, exposed areas of sandstone bedrock
in non-praine regions large enough to warrant the name glade are now uncommon
(Nelson 1987); this is particularly so with scmdstone glades of the Roubidoux
formation. Known glades on this formation are rare (< 10 ha) and restncted to four
counties in south-central Missouri (Nelson 1987; Currier 1991; Ryan 1992; Rvan &
Smith 1991).

The glades at Falls Hollow are particularly interesting in that they are exclusively
associated with intermittent streams. During heavy rziins, which are frequent in the
spring, these streams flood the glades. The impact of such flooding is evident in the
obvious patterns of erosion present (Figure 1). Frequent flooding has been a
significant factor in keeping these glades open, whereas fire has probably played a
secondary role in maintaining these sites historically. Although Hooding has kept
substantial areas of bedrock open and largely free of vegetation, it has also created
habitats suitable for certain plants. Because sections of the sandstone are more
resistant to erosion and weathering (Beveridge 1990), the glades have developed
ledges, depressions, and other structures where water pools and soil accumulates.
These processes have added floristic richness to the glades by creating areas where
plant species not ordinarily associated with glades, such as Alopecurus carolinianus
Walter, Cyperus acuminatus Torrey & Hook., Gratiola neglecta Torrey. Leersia
oryzoides (L.) Swartz, Lythrum alatum Pursh, and Rolala ramosior (L.) Koehne, can
thrive.

There is no evidence of prior botanical work at Falls Hollow. Prior to 1940 this
area was utilized for farming and grazing, and in 1940 the land was purchased by the
Department of the Army for the construction of Fort Leonard Wood. Given the small
size of each glade and their location on a military reservation, it is likely that previous
workers either overlooked or were unable to access these glades (Steyermark 1963;
Johnson et al. 1990). Skinner (1991) located the glades at Falls Hollow and Ryan
(1992) assessed them as part of a natural features inventory, but neither botanized the
site intensively.

METHODS

Forty-one trips were made to Falls Hollow on a weekly basis from 1 Apnl to 23
October 1994, and all glades were surveyed during each visit; during late spring and
early fall, the glades were often visited twice weekly. Vouchers were deposited at
UMO, with the exception of specimens representing county records and rare and
endangered species, which were deposited at MO. Verification of county records and
difficult taxa was made by Dr. George Yatskievych of the Missoun Department of
Conservation. Grasses were venfied by Dr. Michel Leiong of the University of South
Alabama, and Dr. Robert Krai of Vanderbilt University verified the sedges.

Determinations were made principally using Steyermark (1963). Plants were
occasionally identified with Gleason & Cronquist ( 1991). Identification of Aster spp.
was made with Jones (1989). Nomenclature follows Yatskievych & Turner ( 1990),
with the exception of Panicum, which follows Leiong (1986), and Heucfiera x
hirsuticaulis (Wheelock) Rydb. which follows Gleason & Cronquist ( 1991).

266

PHYTOLOGI A

volume 78(4): 264-276

Apnl 1995

Figure 1. Glade at Falls Hollow, as seen from
south (top photo) , and west (bottom photo) . Top
photo shows course of intermittent stream; arrow
indicates the point where the stream enters the
glade .

Hays: Flonstic survey of Fails Hollow 267

RESULTS/DISCUSSION

Only the flora of the three smaller glades is reported here. My decision to exclude
the largest (0.8 ha) glade is based upon the severe impact that has occurred due to the
construction of a military finng range adjacent to the glade. Large amounts of soil (in
the form of numerous dirt mounds), gravel, and other debns (expended nfle shells,
glass and metal containers and plastic items) were deposited on the glade dunng the
construction of the range; frequent maintenance of the range has produced a constant
supply of fresh debris. It is, in fact, often hard to detenmine which parts of the glade
are natural and which ones are the result of intense human activity. By comparison,
the three smaller glades show no signs of recent disturbance and are comparable lo
other Roubidoux sandstone glades judged to be high quality communiues (Ryan
1992). It was felt that the inclusion of the largest glade, with its large number of non-
native and weedy taxa, would misrepresent the true nature of the Falls Hollow glade
community.

Due to the rarity of sandstone glades in Missouri, I would encourage those
involved with land management at Fort Leonard Wood to conserve Falls Hollow
glades. The greatest potential for conservation lies with the three smaller, high quality
glades. These glades are not directly impacted by the finng range, as is the largest
glade, and management efforts would be minimal. In fact, as long as there is no
human disturbance in the form of logging, construction, or vehicular traffic, these
three glades would require only slight management in the form of a prescnbed bum
plan. Although they are maintained principally by flooding, rather than by fire, a fire
management plain would nonetheless be beneficial. By reducing woody invasion
along the borders of the glades - the areas least affected by flooding - fire would
diversify the habitat around the glades by maintaining or expanding (or in some cases
creating) the ecotone between the forest and each glade.

A total of 137 taxa, representing 48 families and 104 genera, was collected. A list
of the plant taxa collected at Falls Hollow glades follows. Poaceae (26) and
Asteraceae ( 13) are the two families with the largest number of representatives. Three
species identified during this study, Silene regia Sims, Sporobolus ozarkanus Femald,
and Trifolium reflexum L. var. reflexum, are currently on the Missoun Department of
Conservation's list of Rare and Endangered Species (1992). Due to the ranty of
Silene regia at Falls Hollow, a photo voucher was made in lieu of an actual collection.

Plants charactenstic of Falls Hollow glades include Crotonopsis elliptica Willd.,
Diodia teres Walter, Hypericum gentianoides (L.) Bntton, Juniperus virginiana L.,
Schizachyrium scoparium (Michaux) Nash. Sporobolus spp., Vaccinium arboreum,
and Vulpia octoflora (Walter) Rydb. Each of these species, with the exception of
Juniperus virginiana and Sporobolus spp., is listed as charactenstic of sandstone
glades in Missouri by Nelson & Ladd (1983). Studies indicate that the species
composition of sandstone glades vary, however, based upon the age and ongin of the
substrate, slope and aspect, and recent vegetational history. Based on the list of taxa
reported from previous work, only Quercus marilandica, Schizachyrium scoparium,
and Vaccinium arboreum are icnown to occur on all sandstone glades (Bacone et al.
1983; Jeffenes 1985, 1987; MacRoberts & MacRoberts 1992, 1993). The literature
also indicates that dominants vary. Bacone et al. (1983), Jeffenes (1987), and
MacRoberts & MacRoberts (1992, 1993), list Schizachyrium scoparium as the
dominant or co-dominant plant species on sandstone glades. Jeffenes (1985) found
Coreopsis grandijiora (Hogg) ex Sweet and Crotonopsis elliptica to be dominant on
calico sandstone in northern Arkansas. At Falls Hollow, Schizachyrium scoparium
and Sporobolus spp. were apparently co-dominants at one glade, whereas Sporobolus

268 PHYTOLOGIA volume 78(4):264-276 Apnl 1995

spp. and Aristida spp. were apparently co-dominants on the other two glades. This is
interesting, because glades where Sporobolus spp. were previously noted as dominant
or abundant had substrates other than sandstone (Quarterman 1950; Kucera & Manin
1957; Baskin & Basicin 1973, 1977, 1978; Nelson 1987). But as MacRoberts &
MacRoberts (1993) and Jeffenes (1985) have noted, sandstone glades are less studied
than glades of other substrates. Future studies should reveal more about the
similarities and differences among sandstone glades.

Scientific names of new taxa for Pulaski County are preceded by an asterisk.
Within each group, families, genera, and species are arranged alphabetically.

PTERIDOPHYTA

Adiantaceae

Cheilanthes lanosa (Michaux) D. Eaton

Aspleniaceae

Asplenium platyneuron (L.) Bntton, Stems, &. Pogg. vax. platyneuron

Dryopteridaceae

*Cystopteris tennesseensis Shaver
Dryopteris marginalis (L.) A. Gray

PINOPHYTA

Cupressaceae

Juniperus virginiana L. var. virginicma

MAGNOLIOPHYTA

MAGNOLIOPSIDA

Acanthaceae
Ruellia hwnilis Nutt.

Hays: Ronstic survey of Falls Hollow 269

Anacardiaceae

Rhus copallina L.
R. glabra L.

Asteraceae

Ambrosia artemisiifolia L.

Ambrosia bidentata Michaux

Antennaria planlaginifolia (L.) Hook.

Aster linariifolius L. var. linariifolius forma linariifolius

Aster pilosus Willd.

Aster sericeus Vent, forma sericeus

*Heliopsis helianthoides (L.) Sweet var. occidentalis (T. Fisher) Steyerm.

Hieracium gronovii L.

Krigia dandelion (L.) Nutt.

K. virginica (L.) Willd.

Rudbeckia missouriensis Pursh

Solidago nemoralis Dryander

S. ulmifolia Muhlenb. ex Willd.

Brassicaceae

Cardamine concaienata (Michaux) O. Schwarz

C. parviflora L. var. arenicola (Britton) O. Schwarz

Draba brachycarpa Nutt. ex Torrey & A. Gray

Cactaceae

*Opuntia humifusa (Raf.) Raf. var. humifusa

Caesalpmaceae
Cercis canadensis L.

Callitrichaceae

CalUtriche heterophylla Pursh var. heterophylla

Capri foliaceae

Lonicera flava Sims
Symphoricarpos orbiculatus Moench
Viburnum rufidulum Raf.

270 PHYTOLOGIA volume 78(4): 264- 276 Apnl 1995

Caryophyllaceae

Arenaria serpyllifolia L.

Cerastium brachypetalum Pers.

* Paronychia fastigiata (Raf.) Femald var. paleacea Femald

Silene regia Sims

Clusiaceae

Hypericum gentianoides (L.) Bntton
H. punc latum Lam.

Comaceae
Cornus florida L

Ebenaceae

*Diospyros virginianaL. var. platycarpa Sarg. (orma. platycarpa

Ericaceae

Vaccinium arboreum Marshall

Euphorbiaceae

Croton capitatus Michaux var. capitatus
Crotonopsis elliptica Willd.
Euphorbia coroUata L
Tragia betonicifolia Nutt

Fabaceae

Baptisia alba (L) Vent

Stylosanthes biflora (L.) Britton, Steams, & Pogg

Tephrosia virginiana (L.) Pers.

Trifolium reflexum L. var. reflexum

Fagaceae

Quercus alba L

Q. marilandica Muenchh.

Q. stellata Wangenh. var. stellata

Hays: Floristic survey of Falls Hollow 271

Juglandaceae

Carya texana Buckley

Linaceae

Linum medium (Planchon) Bntton var. texanum

Lythraceae

Cuphea viscosissima Jacq.
Lythrum alatum Pursh var. alatum
Rotala ramosior (L.) Koehne

Mimosaceae

Schrankia nultallii (DC. ex Bntton & Rose) Standley

deaceae

Fraxinus americana L

Onagraceae

Ludwigia altemifolia L.
Oenothera linifolia NutL

Oxalidaceae
Oxalis violacea L.

Plantaginaceae

PUmtago aristala Michaux
P. pusilla NutL var. pusilla
P. virginica L.

Polemoruaceae

* Phlox pilosa L. subsp. ozarlcana (Wherry) Wherr\

272 PHYTOLOGIA volume 78(4):264-276 Apnl 1995

Polygalaceae

Poly gala sanguinea L. iovms. sanguinea

Polygonaceae

Polygonum tenue Michaux
Rumex acetosella L.

Pbrtulacaceae

Pomdaca oleracea L
Talinum calycinum Engelm.

Ranunculaceae

Ranunculus fascicularis Muhlenb. ex Bigelow
R. harveyi (A. Gray) Britton forma harveyi

Rhamnaceae

Rhamnus caroliniana Walter

Rosaceae

*Prunus hortulana L.

P. mexicana S. Wats.

Rosa Carolina L.

Rosa setigera Michaux var. setigera forma setigera

Rosa setigera Michaux var. tomentosa Torrey

Rubus flagellaris Willd.

*Rubus invisus (L. Bailey) Britton

Rubiaceae

Cephalanthus occidentalis L.
Diodia teres Walter
Hedyotis crassifolia Raf .

Sapotaceae

Bumelia lanuginosa (Michaux) Pers.

Hays: Roristic survey of Falls Hollow 273

Saxifragaceae

Heuchera x hirsuticaulis (Wheelock) Rydb.

Scfophulariaceae

Agalinis tenuifolia (M. Vahl) Raf.
*Gratiola neglectalow^y
Leucospora multifida (Michaux) NutL
*Nunallanthus canadensis (L.) D. Sutton
Penstemon pallidas Small

Ulmaceae

Celtis tenuifolia Nutt. var. tenuifolia
Ulmus rubra Muhlenb.

Violaceae

Viola pedata L. forma pedam
V. rafinesquii Greene

UUOPSIDA

Commelinaceae
Tradescantia ohiensis Raf.

Cyperaceae

*Bulbostylis capillaris (L.) C.B. Clarke

Carex bushii Mackenzie

Carex flaccosperma Dewey var. glaucodea (Tuckerman) Kiik.

Cyperus acuminatus Torrey & Hook.

Cyperus aristatus Rottb.

Fimbristylis autumnalis (L) Roemer & Schultes

Upocarpha micrantha (M. Vahl) G. Tucker

* Rhynchospora globularis (Chapman) Small var. recognita Gale

Iridaceae

Sisyrinchium campestre E. Bickn. forma campestre

274 PHYTOLOGIA volume 78(4): 264-276 Apnl 1995

Juncaceae

Luzula bulbosa (Alph. Wood) Rydb.
Juncus interior Wieg.

Liliaceae

Allium canadense L. var. canadense
Allium canadense L. var. mobilense (Regal) F. Ownbey
Camassia scilloides (Raf.) Cory forma scilloides
Hypoxis hirsuta (L.) Cov. forma vilosissima
Nothoscordum bivalve (L.) Bntton

Orchidaceae
*Spiranthes tuberosa Raf.

Poaceae

Agrostis elliottiana Schultes

Agroslis hyemalis (Walter) Britton, Stems, & Pogg var. hyemalis

Agrostis perennans (Walter) Tuckerman

*Alopecurus carolinianus Walter

Andropogon gerardii Vitman var. gerardii

Aristida dichotoma Michaux var. dichotoma

*Aristida longespica Poiret var. longespica

* Aristida purpurascens Poiret

Daruhorda spicata (L.) P. Beauv. ex Roemer & Schultes var. spicata
*Digitaria cognata (Schultes) Pilger var. cognata
*Leersia oryzoides (L.) Swartz
L. virginica Willd.

* Muhlenbergia mexicana (L.) Trin.
Panicum acuminatum Swartz var. acuminatum
P. depauperatum Muhlenb.

P. flexile (Gattinger) Scribner

*P. philadelphicumThn. var. philadelphicum

P. virgatum L.

Schizachyrium scoparium (Michaux) Nash

Sphenopholis obtusata (Michaux) Scnbner var. obtusaia

*Sporobolus asper (Michaux) Kunth var. asper

Sporobolus clandestinus (Biehler) A. Hitchc.

*Sporobolus ozarkanus Femald

*Sporobolus vaginiflorus (Torrey) Alph. Wood

Tridens flavus (L.) A. Hitchc. var.flavus

Vulpia octoflora (Walter) Rydb. var. glauca (Nutt.) Femald

Hays: Ronslic survey of Falls Hollow 275

ACKNOWLEDGMENTS

I would like to thank the following individuals for their assistance during this
study: Dr. Robert Krai of Vanderbilt University, Timothy Smith and Janet Siemburg
of the Missoun Department of Conservation, and Dr. George Yatskje\ ych of the Rora
of Missoun project at the Missoun Department of Conservation. A special thanks
goes out to Dr. Michel Leiong of the University of South Alabama. His teaching and
continual encouragement are a constant source of inspiration. And finally to my w ife,
whose tolerance of a house continuously cluttered with specimens and manuals is
lovingly appreciated.

LITERATURE CITED

Bacone, J. A., L.A. Casabere, & M.D. Hutchison. 1983. Glades and barrens of

Crawford and Perry Counties, Indiana. Proc. Indiana Acad. Sci. 93:291-301.
Baskin, J.M. & C.C. Baskin. 1973. Observations on the ecology of Sporobolus

vaginiflorus in cedar glades. Castanea 38:25-35.
Baskin, J.M. & C.C. Baskin. 1977. An undescnbed cedar glade community in

Middle Tennessee. Castanea 42: 140- 145.
Baskin, J.M. & C.C. Baskin. 1978. Plant ecology of cedar glades in the Big Barren

Region of Kentucky. Rhodora 8a 545-557.
Beveridge, T.R. 1990. Geologic wonders and curiosities of Missoun. Missoun

Department of Natural Resources, Rolla, Missouri.
Currier, M. P. 1991. Missouri Namral Features Inventory: Camden Co., Cole Co.,

Cooper Co.. Gasconade Co., Maries Co., Miller Co., Moniteau Co., Morgan

Co., Osage Co. Missouri Department of Conservation, Jefferson City, Missoun.
Gleason, H.A. & A. Cronquist. 1 99 1. Manual of the Vascular Plants of the

Northeastern United States and Adjacent Canada, 2nd ed. New York Botanical

Gardens, New York, New York.
Jefferies, D.L. 1985. Analysis of the vegetation and soils on calico rock sandstone in

northern Arkansas. Bull. Ton-ey Bot. Club 1 12:70-73.
Jefferies, D.L. 1987. Vegetation analysis of sandstone glades in Devil's Den State

Park, Arkansas. Castanea 52:9-15.
Johnson, F.L., R.A. Thompson, CM. Sladewski, J.R. Estes, & G.D. Schnell.

1990. Floral Inventory of Fort Leonard Wood, Missoun. Oklahoma State

Biological Survey, Norman, Oklahoma.
Jones, A. G. 1989. Aster and flrac/jyacrw in Illinois. Bull. Illinois Nat. Hist. Surv.

34:139-194.
Koenig, J., ed. 1961. The Stratigraphic Succession in Missouri, 2nd ser., vol. 40.

State of Missouri, Division of Geological Survey & Water Resources, Rolla,

Missouri.
Kucera, C.L. & S.C. Martin 1957. Vegetation and soil relationships in the Glade

Region of the southwest Missoun Ozarks. Ecology 38:285-291.
Leiong, M.G. 1986. A taxonomic treatment of the genus Panicum (Poaceae) in

Mississippi. Phytologia 61:251-269.
MacRoberts, M.H. & B.R. MacRoberts. 1992. Flonstics of a sandstone glade in

western Louisiana. Phytologia 72: 130-138.
MacRoberts, M.H. & BR. MacRoberts. 1993. Flonstics of two Louisiana

sandstone glades. Phytologia 74:43 1-437.

276 PHYTOLOGIA volume 78(4):264-276 Apnl 1995

Missouri Department of Conservation. 1992. Rare and Endangered Species of

Missouri Checklist. Jefferson City, Missoun.
Nelson, P. 1987. The Terrestrial Natural Communities of Missouri, rev. ed.

Missoun Department of Natural Resources, Jefferson City, Missouri.
Nelson, P. & D. Ladd. 1983. Preliminary report on the identification, distnbution

and classification of Missoun glades. Pages 59-76 in C. Kucera (ed.).

Proceedings of the Seventh North Amencan F*raine Conference, August 4-6,

1980. Southwest Missouri State University, Spnngfield, Missoun.
Quarterman, E. 1950. Major plant communities of Tennessee cedar glades. Ecology

31:234-254.
Ryan, J. 1992. Missouri Natural Features Inventory: Phelps Co., Pulaski Co..

Laclede Co. Missouri Department of Conservation, Jefferson City, Missoun.
Ryan, J. & T. Smith. 1991. Missouri NaturcU Features Inventory: Howell Co.,

Texas Co., Wright Co. Missouri Department of Conservation, Jefferson City,

Missouri.
Skinner, M. 1991. Rare and Endangered Plant Survey of Fort Leonard Wood

Military Reservation. Missouri Department of Conservation, Nanjral History

Division, Jefferson City, Missouri.
Steyermark, J. A. 1963. Flora of Missouri. Iowa State University Press, Ames,

Iowa.
Thorn, R.H. & J.H. Wilson. 1980. The natural divisions of Missoun. Trans.

Missouri Acad. Sci. 14:9-23.
Wolf, D.W. 1989. Soil Survey of Pulaski County, Missouri. U.S. Department of

Agriculture, Washington, D.C.
Yatskievych, G. & J. Turner. 1990. Catalogue of the Flora of Missouri.

Monographs in Systematic Botany, vol. 37. Missoun Botanical Garden, St.

Louis, Missouri.

Phyiologia (April 1995) 78(4); 277-281

STUDIES ON THE ISOZYME VARIATION PATTERN AMONG INDIVIDUALS

AND POPULATIONS OF THE ENDANGERED SPECIES DEUTZIA

MULTIRADIATA (HYDRANGEACEAE) ON MT. JINFO OF NANCHUAN

Ping He & Hong Li

Department of Life Science, Southwest China Normal University, Beibei, Chongqing

630715 CHINA

ABSTRACT

With the electrophoretic technique, we studied the peroxidase (PER), catalase
(CAT), esterase (ES), amylase (AA), citric acid dehydrogenase (CDH), and
glutamate dehydrogenase (GDH) in leaves of 41 plants from five small
populations with different altitudes and habitats of the endangered species
Deutzia multiradiata which is endemic to Mt. Jinfo of Nanchuan, China. The
band number variation of those isozymes is studied by clustering analyses on
Manhattan metric by UPGMA. The result indicates that though individuals
within the same population show certain similarities in the isozyme band number,
the electrophoretic difference in the band number and the genetic divergence
within the same populations are notable, while most individuals from different
populations show higher resemblance in the band number of those isozymes,
which reveals that there are few relations between the band number of those
isozymes and the collecting habitats of those plants.

KEY WORDS: multivariate analyses, isozyme variation, Deutzia multiradiata,
Hydrangeaceae, China

Deutzia miMradiata W.T. Wang is a species endemic to Mt. Jinfo (29° 05' N,

107° 10' E) of Nanchuan County in Sichuan Province and belongs to subsect.
Cymosae Rehder of sect. Deutzia in the genus Deutzia Thunberg of Hydrangeaceae
(He 1989, 1990). It was published as a new senes named ser. Multiradiaiae P. He
because of its unique morphological characteristics in subsect. Cymosae Rehder (He &
Hu 1989; He & Pan 1994). Further studies show it also possesses some unique
biologicjil features. It is scattered as small populations in a very resUicted area smaller
than 10 km^ from 600-1200 m altitude on Mt. Jinfo of Nanchuan. Based on our
detailed observations and statistical works in the field over many years, we found there
are no more than 800 living individuals of the species within its entire geographic
distribution and that it is indeed an endangered species. It competes poorly when
growing with other species such as Comjxisitae and Poaceae with strong competitive
abilities, but because of its highly developed root system, it mostly favors those
exposed habitats with water-exuded limestone crevices where other plant sf)ecies

277

Ping He & Hong Li:

Isozymes in Deutzia muUiradiala

278

cannot survive. Whether it is an adaptive strategy for a species such as Deiuzia
multiradiata to occupy the exposed micro- habitats where other species cannot survive,
or this distnbulion is a retrogressive phenomenon, is not known. It is known that the
survivorship and development of a given species is not only aifected by the
environmental conditions, but also regulated by its own genetic features. It is
necessary to understand the genetic variation of such a restncted endemic and
endangered species as Deutzia multiradiata.

MATERIAL AND METHODS

After multiple field observations and collections, the authors dug 78 living
individuals from five populations of Deutzia multiradiaia in Apnl 1994. and
transplanted them in native soil to plastic pots. These pots were then moved to the
Botanical Garden of Southwest China Normal University. The collecting locations
and their micro-habitats are shown in Table 1.

Table 1. The collecting localities and micro-habitats for the living material used in this
study.

Population

Elevation (m)

Locality

Micro- habitat

A

600

Shanquan

limestone crevice

B

720

Banhe

damp scrub

C

800

Lower Daheba

cliff crevice

D

760

Yihaoqiao

slightly dry scrub

E

1200

Upper Daheba

talus slope

Sample preparation follows Wu (1979). The vertical plate polyacrylamide gel
electrophoresis was conducted following Wu (1979) and Hu & Wan ( 1985).

Based on the schematic figures of the enzyme bands, we obtained the total band
number for each individual and then inserted them into the original data matrix which
was processed on a 386DX40 computer using BASIC programs for clustenng
analyses using a Manhattan metric under UPGMA (Zhong et al. 1990).

RESULTS AND DISCUSSION

In the isozymic dendrogram produced as a result of this study (Figure 1), 41
individuals were clmtered into four Manhattan metnc-0.82 isozymic phenons. The
"A" Manhattan metnc-0.82 phenon is composed of fifteen individuals from among
populations A, B. C, D, and E, while the "B" Manhattan metnc-0.82 phenon is
composed of five individuals ongmating from populations A and C. The "C"
Manhattan metric-0.82 phenon consists of thirteen individuals collected from
populations A, B, C, and E. The "D" Manhattan metnc-0.82 phenon is formed of a
mixture of eight individuals from populations A, B, and D. Generally speaking,
though some groups of individuals from particular populauons cluster very closely
(such as numbers 1-2-8, 12-17, 15-19, 3-4, and 21-22) and show high resemblance in

279

PHYTOLOGIA volume 78(4): 277-281 Apnl 1995

1.0 0. 9 0. 8 0. 7 0. 6 0. 5 0. 4 0. 3 0. 2 0. 1

1. 0 0. 9 0. 8 0. 7 0. 6 0. 5 0. 4 0. 3 0. 2 0. 1
Manhattan Metric

Fig.l Dendrogram by the Manhattan metric with WPGMA, showing
the variation in band number of six isozymes in leaves of 41
individuals from 5 populations of Deutzia multiradiata .

Ping He & Hong Li: Isozymes in Deitlzia multiradiala 2K0

isozyme band number, most clusters are composed of mdividuais from several
populations. On the other hand, some individuals from separate populations (e.%.,
numbers 31-41, 40-24, 9-16, 29-37, 32-14-5, 18-34, and 36-25) have great similarity
in band number. Therefore, isozyme band numbers of a plant do not show good
correlation with the habitat from which it was collected.

Though there are similanties in band numbers for PER, CAT, ES, AA, CDH, and
GDH in leaves of some individuals from the same population, certain genetic vanation
does occur among individuals within the same populauon. Conversely, some
individuals from different populations show notable convergence in the isozyme band
numbers. This indicates that isozyme band numbers of given plants are not correlated
with the micro-habitat from which the plant was collected. Deutzia multiradiala was
supposedly widely distnbuted in the past, with the current distribution restncte«l to a
small area around Mt. Jinfo. During the long evolutionary history of the species, the
isolated distnbution of different populations limits gene exchange between individuals
of different populations, which explains why some individuals from different
populations show higher similarities in the isozyme band numbers, while the
remarkable hybridizations among individuals of the same population lead to greater
gene exchange and stronger divergence in the isozyme band numbers among
individuals within the same papulation. Because the species can flower and pollinate
easily, but most of its seeds are abortive, which makes the transfemng of those nch
genetic variations within the same population to their offspring nearly impossible so
that it is more difficult for the fixation of those genetic vanations as a genetic resource
for the development of such a narrowly distnbuted endangered species iis D.
multiradiata. Though a few scattered habitats such as moist limestone crevices in a
restricted area from 600 to 1200 m altitude on Mt. Jinfo can be favorable tor the
survival of D. multiradiata presently, owing to its paucity of genetic vanation, it does
not have a bright future if these habitats are changed.

ACKNOWLEDGMENTS

This f>aper is one part of the project "Studies on the interspecific alfinitits and
microevolution of Deutzia in China" granted to the first author by the "Natural Science
Foundation of China" with grant no. 39270058 and also included in the project
"Biosystematics of Deutzia in China" granted to the first author by the "Sichuan ^'outh
Science and Technology Foundation" (SYSTF). We are indebted to Prof. Dr. F. Tan
and Mrs. J.F. Yan for their assistance with field work. Thanks are extended to Prof.
L.C. Hu and Prof. J.H. Xiong for their review of the paper.

UTERATURE CITED

He, P. 1989. A taxonomic study on the genus Deutzia of Sichuan (I) (in Chinese).

J. Southw. Chin. Norm. Univ. (Sci. Nat.) 14<l):67-74.
He, P. 1990. Taxonomy of Deutzia (Hydrangeaceae) from Sichuan, China.

Phytologia 69(5):332-339.
He, P. & L.C. Hu. 1989. A taxonomic study on the genus Deutzia of Sichuan (II)

(in Chinese). J. Southw. Chin. Nonn. Umv. (Sci. Nat.) 14(4): 1 17-127.

281 PHYTOLOGIA volume 78(4): 277-281 Apnl 1995

He, P., & T.C. Pan. 1994. Multivanate analyses of varation pattern in the Deulzia

A«M///>rt<^/a/a group (Hydrangeaceae) in Ml. Jinfo of Nanchuan. J. Southw. Chin.

Norm. Univ. (Sci. Nat.) 19(4):409-418.
Wu, S.B. 1979. The polyacrylamide gel disc-electrophoresis of proteins and

isozymes in plant tissues (in Chinese). Plant Physiology Letters 1:30-33.
Hu, N.S. & X.G. Wan. 1985. Isozyme Technology and its Applications (in

Chinese). Changsha, China: Hunan Science and Technology Publishing House.

pp. 112-117.
Zhong. Y., J.K. Chen, & D.S. Huang. 1990. The Methods and Progratns for

Numerical Taxonomy (in Chinese). Wuhan, China: Wuhan Universitv Press, pp.

203-212.

Phyiologia (Apnl 1995) 78(4) 282-284

A NEW VARIETY OF DEUTZIA (HYDRANGEACEAE) IN CHINA

Ping He

Department of Life Science, Southwest China Normal University, Beibei, Chongqing

630715 P.R. CHINA

ABSTRACT

A new variety named Deutzia rfwco/or Hemsl var bicruristyli P He collected
from Hubei Province in China is described.

KEY WORDS: Deutzia, Hydrangeaceae, new taxon, China

Deutzia discolor Hems!, var. bicruristyli P. He, var. nov. Plate 1. TYPE;
CHINA. Hubei: Enshi, Damuxiang, 1380 m alt., frutex 2 mm altus, Ibliie
cilbidis, 25 June 1958 (fr). Fang Minyuan 24492 (Typus var. in SZ conserv.);
eodem loco, 25 June 1958 (fr). Fang Minyuan 24192 (SZ).

Frutex 1-2 m altus; ramuli purpureo-brunnei vel fumeus, glabn, cortia>
non delapso. Folia petiolata; lamina chartacea, ovato-lanceolata vel oblongc-
lanceolata, 3-7 cm longa, 1-2 cm lata, apice acuminata, basi cuneata, margine
denticulata (dentis 1-2 mm longis), supra vindis, pilis stellatis 5-8-radiati5
sparse conspersa, subtus albida, pilis 10-15-radiatus dense obtecta ad costam
nervosque sine simpliciter puberula, nervis lateralibus utnnsecus 4-5; petiole 3-
6 mm longi. Cyma multiflora, 3-7 cm diam., pedicelli purpureo-brunnei, 7-15
mm longi, pilis 10-14-radiatis sparse conspersis; lobis calyx lanceolat's,
persistentibus, ca. 3 mm longis; styli 3-4, 5-6 mm longi, apice bicruns.
Capsula subglobosa, 4-5 mm diam.

A Deutzia discolor Hemsl. typica differt plerumque cortice ramulis non
delajDso; foliis supra 5-8-radiatis stellato-pilosis pilis adpressis conspersis;
subtus sine simpliciter puberulis; pedicellis 7-15 mm longis; stylus bicruns.

This variety differs from Deutzia discolor var. discolor mainly because its twigs
are not fissured, upper side of leaf blades are sparsely covered with 5-8-rayed
appressed stellate hairs, lower side of leaf blades lacking simple hairs, pedicels 7-15
mm long, and most importantly because its style apex is two-lobed.

282

Ping He:

New Deutzia from China

Fig.l Morphology of Deutzia discolor var. bicruristyli P. He

a. stellate hairs on upper side of lamina, X 40;

b. stellate hairs on lower side of lamina, X 40;

c. stellate hairs on fruiting pedicel, X 50;

d. fruit;

e. fruiting inflorescence.

284 PHYTOLOGIA volume 78(4): 282-284 Apnl 1995

ACKNOWLEDGMENTS

This study was supported by the "National Science Foundation of China" (NSFC)
with grant no. 39270058 and partly supported by "Sichuan Youth Science and
Technology Foundation" (SYSTF)- Much thanks are due to Prof. Dr. L.C. Hu of
Sichuan University and J.H. Xiong in Southwest China Agricultural University for
their critical review of the paper.

Phytologia (April 1995) 78(4) 285-286

A NEW SPECIES OF POTENTILLA (ROSACEAE) FROM CERRO
QUIEXOBRA, OAXACA

B.L. Turner
Departmentof Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

A new species of Potentilla, P. macdonaldii B.L Turner, is described from
the subalpine areas of Cerro Quiexobra, Oaxaca It is perhaps most closely
related to P. richardii Lehm. of the sect. Multijugae but differs in having
thinner more dentate leaves, flowers 2-5 arranged in open cymes, and smooth
achenes

KEY WORDS: Rosaceae, Potentilla, Mexico

Routine identification of Mexican plants has revealed the following novelty.

POTENTILLA MACDONALDII B.L. Turner, spec. nov. TYPE MEXICO.
Oaxaca; Mpio. Miahuatldn, 35 km ESE of Miahuatl^, 5 km NE of Santo
Domingo Ozolotepec, Cerro Quiexobra, "subalpine glades surrounded by pine
forests on ndgetops and in mountain saddles", 3500-3700 m, 3 Oct 1990, Andrew
McDonald 2995 (HOLCfTYPE: TEX!; Isotype: MEXU).

PotentiUae richardii Lehm. similis sed differt foliolis tenuioribus dentibus
numerosionbus (dentes marginales 5-9 per foliolum vs. dentes. 3-5 per
foliolum), flonbus in cymis laxis dispositis in pedunculis 1-4 cm longis (vs.
plerumque solitariis axillanbus in pedunculis 1 cm longis vel minus), et
acheniis laevibus (vs. omatis nervatura prominenti).

Prostrate or subprostrate perenmal herbs 5-18 cm high, arising from woody
taproots. Stems moderately villous-hirsute, the hairs mostly 0.8-1.5 mm long.
Leaves mostly (3-)5-7 pinnately parted, the lower mostly 3-6 cm long, 1.5-2.5 cm
wide; stipules umted forca. 1/2 their length to the lower petiole proper; peuoles 1.5-
2.0 cm long, pubescent with silky-villous spreading hairs 1-2 mm long; divisions of
the leaf flabellate, irregularly dentate with 5-9 teeth, the latter 0.6-1.4 mm long.
Inflorescences cymose, 2-5 flowers per primary stems; peduncles mostly 1-4 cm long,
pubescent like the stems. Receptacle corneal, pubescent. Bracts of the calyx 3.5-4.5
mm long, broadly ovate in outline, clearly 3-lobed. Sepals lanceolate, 5-6 mm long,
ca. 2 mm wide, uruted below, sparsely villous without, glabrous within. Petals
yellow, spreading, 6-7 mm long, 5-6 mm wide, the apices emarginate, the cleft ca. 1

285

Turner. New Poieniilla from Mexico 286

mm deep. Stamens 18-21 in 2-3 senes. Styles lerete-ctinical, oulcurved, ca. 0.5 mm
long, glabrous. Seeds ovoid, brown, ca. 0.75 mm long, glabrous, smooth.

ADDITIONAL SPECIMEN EXAMINED: MEXICO. Oaxaca; about same
locauon as the type, 3650-3800 m, 10 Dec 1989. McDonald 2919 (TEX).

This taxon, because of its pinnately compound leaves, is obviously related to
Potentilla richardii but can be readily reaignized by characters indicated in the
following coupleL

1. Flowers 2-5, arranged in open cymose panicles, the ultimate peduncles 1-4 cm
long; petals spreading, mostly 6-7 mm long; bracts at base of sepals mostly 3-cIeft,
rarely not; achenes smooth P.macdonaldii

1. Rowers mostly solitary, axillary, the peduncles 1 cTn long or less; petals erect,
mostly 4-5(-6) mm long; bracts at base of sepals mostly elliptical, sometimes 2-
cleft; achenes nervate P- richardii

While compared with Potentilla richardii of the sect. Multijugae {sensu Rydberg
1902) in the above diagnosis and key, P. macdonaldii also possesses characters
suggestive of P. heterosepala Fritsch., namely flowers arranged in leafy cymes and
bractlets 3-lobed. In Johnston's (1985) key to sections, P. tnacdonaldii, largely
because of its pinnately compound leaves and small styles, will key to the sect.
Multifidae. In truth, P. macdonaldii appears to combine characters of both P.
heterosepala (a poorly studied complex of Guatemala and southernmost Mexico) and
P. richardii (a better known complex of the trans-volcanic belt of southcentral
Mexico), standing somewhat between these in both morphology and geography.

It is a pleasure to name this species for its only known collector. Dr. Andrew
McDonald, currently at Harvard University and first botanist to ascend and collect on
Cerro Quiexobra, from which numerous novelties have been named, many in his
honor (c/. Nesom 1995).

ACKNOWLEDGMENTS

I am grateful to Guy Nesom for the Latin diagnosis, and to him and Piero Delprete
for reviewing the manuscript

UTERATURE CITED

Johnston, B.C. 1985. Studies in Potentilla. I. Key to North American sections.

Phytologia 57:292-302.
Nesom, G.L. 1995. Two new species of Cas////e/a (Scrophulanaceae) from southern

Oaxaca, Mexico. -Phytologia 78<2): 131-133.
Rydberg, P.A. 1908. Potentilla, in N. Amer. R. 22:293-352.

Phyiologia (Apnl 1995) 78(4) 287-290

A REVISED SYNOPSIS OF THE PINES 6: SUPPLEMENT TO THE
SUBGENERA

Pierre Landry
50 Dussault, suite 306, Hull, Quebec CANADA J8X 4G2

ABSTRACT

This chapter presents an alternate key to the subgenera, eliminating the
complex morphology of the twigs, but keepmg in accordance with Linnean
philosophy. Basic particulars of the seven subgenera are given. The conclusion
is made that plant behavior is an important part of plant systematics. The
proposed division of Pinus into seven subgenera is presented as natural and
clear-cut. Such a treatment eliminates artificial groups such as sect. Parrya
Mayr emend., by splitting them.

KEY WORDS: Pinus, Pinaceae, systematics

A NEW MORPHOLOGICAL KEY TO THE SUBGENERA

In the previous chapter, (Landry 1994, pp. 74-75) a traditional (Linnean) key to the
subgenera was given. This key was influenced partly with the characters of the twigs.
These characters are now known to be very complex. In order to simplify the key
structure, we hereunder present a new key without twig characters. We have replaced
features of the twigs by leaf characters.

This alternate key is more detailed, having in mind the promotion of the relations
of morphology with behavior.

A. Cones spend three growth seasons to mature, as witnessed by their double
concentric umbos (for Pinus torreyana Parry ex Cam^re, a hand lens is useful to
see them) 1 . Pinus subgenus Pinea

A. Cones spend two growth seasons to mature, as witnessed by their simple umbos. .

B

B. Seeds bodies three times longer than broad 2. Pinus subgenus Gerardia

B. Seeds bodies less than two times longer than broad C

C. The persistent stalks (peduncles) of the cones (and conelets) are thick (4-8

mm diam.) (very long, turning at least 60°. The conelets continue growth
for over three months. Fresh seeds yellow with a faint reddish area at the

apex. Leaves connate, but easily separated

3. Pinus subgenus Tamaulipasa

287

288 PHYTOLOGIA volume 78(4): 287-290 Apnl 1995

The stalks not persistent, or when persistent do not show the same
combination of thickness, length and degree of curvature. The conelets
grow for at most one month. Fresh seeds colored otherwise. Leaves not

connate D

D. Umbos are dorsal (central) at least on the mtcnor (inner) face of the

middle part of the cones E

E. Leaves grouped in such a way that they form dense, long, brush-
like masses. Leaves persist at least ten years

4. Pinus subgenus Balfouria

E. Leaves grouped into rather sparse masses, w hen examined closely.
Leaves persist less than ten years 5. Pinus subgenus Pinus

D. Umbos are terminal or simili-ienminal on all faces of the middle part of
the cones. All the subtending apophyses long taper to a point, or are
thin F

F. Umbos stout, massive, curved slightly or much, simili-terminal. . . .

6. Pinus subgenus Sabinia

F. Umbos not massive, not curved, truly terminal

7. Pinus subgenus Strobus

PARTICULARS

\. Pmiis subgenus Pinea (Endlicher) Landry compnses three species:
Pinus pine a Linn^, Type;
Pinus leiophylla Schiede & Deppe;
Pinus torreyana Parry ex Cam^re.

2. Pinus subgenus Gerardia E Murray is monotypic:

Pinus gerardiana Wallich ex D. Don.

3. Pinus subgenus Tamaulipasa Landry is also monotypic:
Pinus nehonii Shaw.

4. Pinus subgenus Balfouria E Murray comprises only the Foxtail Pines:
Pinus balfowiana Greville & Balfour is their type species.

5. Pinus Linn^ subgenus Pinus comprises a large number of species.

Its type speaes is Pinus sylvestris Linn^.

6. Pinus subgenus Sabinia E Murray is constituted by two species:

Pinus sabiniana Douglas ex D. Don, type, and

Pinus coulteri D. Don

Note: See photo (Figure 1) of a closed cone of Pinus sabiniana, herewith

reproduced from Chaumeton & Durand (1990, species 44) (with their

authorization), showing that the degree of terminality of the apophyses is similar to

that of Pinus albicaulis Engelmann, of subgenus Strobus Lemmon.

7. Pirms subgenus Strobus Lemmon compnses a large number of species.

Its type is Pinus strobus Linn^.

CONCLUSION

This chapter concludes "A Revised Synopsis of the Pines".
This revision enhances the importance of plant behavior for dividing and
subdividing a genus. Four sections:

Landry:

Synopsis of the pines 6

289

Figure 1 Pinus sabiniana an example of a closed cone pine.

290 PHYTOLOGIA volume 78(4): 287-290 Apnl 1995

Pinus sect. Qiiinquefoliis Duhamel Du Monceau emended Landr>' (Landry 1989a

[chapter 1]),

Pinus sect. Cembra Spach (Landry 1989b [chapter 2]),

Pinus sect. Pinea Endlicher emended Landr>' (Landry 1989c [chapter 3]),

Pinus sect. Leiophylla Van der Burgh (Landry 1992 [chapter 4]),

were used to do so.

A new division of Pinus into seven subgenera is proposed. This treatment is very

natural and clear-cuL It moreover eliminates artificial groupings of species (such as

Pinus sect. Parrya Mayr emended), by splitting them.

LITERATURE CITED

Chaumeton, H. (director) & R. Durand (author). 1990. Les Arbres. Guide vert.

F*aris. 298 numbered species, 384 pp.
Landry, P. 1989a. A revised synopsis of the white pines: The subgenera of (Pinus,

sect. Quinquefotiis). Phytologia 65:467-474.
Landry, P. 19i89b. A revised synopsis of the pines 2: The arolla pines (Pinus, sect.

Cembra). Phytologia 65:475-481.
Landry, P. 1989c. A revised synopsis of the pines 3: The Parasol Pine (Pinus, sect.

Pinea). Phytologia 66:477-48 1 .
Landry, P. 1992. A revised synopsis of the pines 4: The Chihuahua Pine (Pinus,

section Leiophylla). Phytologia 72:373-377.
Landry, P. 1994. A revised synopsis of the pines 5: The subgenera of Pinu.;, and

their morphology and behavior. Phytologia 76:73-79.

Phytologia (Apnl 1995) 78(4)291-313.

NOTEWORTHY VASCULAR PLANT COLLECTIONS ON THE KISATCHIE
NATIONAL FOREST, LOUISIANA

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

ABSTRACT

Eighty plant species that are recognized as sensitive, threatened, or
endangered by the Kisatchie National Forest or by the Lx)uisiana Natural
Heritage Program, or that are on the Louisiana Natural Heritage "watch list,"
are known to occur on the Kisatchie National Forest, Louisiana. Over the past
few years, surveys have been undertaken by various botanists and ecologists to
locate and census many of these species. The results add to the growing
botanical knowledge of the West Gulf Coastal Plain.

KEY WORDS; Rare, threatened, endangered plants, Kisatchie National Forest;
Louisiana; West Gulf Coastal Plain

INTRODUCTION

Many authors have pointed out the relatively poor state of knowledge of the
Louisiana and east Texas vascular flora (Thieret 1967b, 1968; MacRoberts 1984;
Bridges & Orzell 1989a, 1989b). In 1967, Thieret stated that although there had been

resident botanists in Louisiana for the past two hundred years "... the flora of
Louisiana is probably the least known of any state in the union" (Thieret 1967b: v).
Two decades later. Bridges & Orzell (1989a- 12) maintained that, although the general

flora of the West Gulf Coastal Rain is fairly well known, "... more exploration of rare
localized and specialized habitats is still needed in order to fully document the
occurrence of plants which are very rare or restncted in their habitat preference . . .."

The Kisatchie National Forest is located in western Louisiana in the center of the
West Gulf Coastal Plain. Over the past few years, a number of surveys have been
conducted of localized and specialized habitats on Forest Service land, one aim of
which was to gather data on rare and specialized plant species. A large body of data
on these species and their micro-distribution now exists, which should be made
acces,sible in order to further the understanding of plant ecology in the West Gulf
Coastal Plain.

Therefore, like Bridges & Orzell (1989a, 1989b) before us, we turn from the
macro-distnbution of species as reported in atlases or floras (MacRoberts 1989;
Thomas & Allen 1993) to the micro-distnbution in the existing landscape. Knowledge
of locally rare species aids in pinpointing rare and vanishing habitat as well as

291

MacRoberts & MacRoberis: Rare plants of Kisatchie National Forest 292

assessing the conservation value (ranty, diversity, and representativeness) of an area
(Bourgeron et al. 1995).

This paper is based on our own surveys, on the rare species files of the Kisatchie
National Forest and the Louisiana Natural Heritage Program, herbarium specimens,
published papers, and unpublished reports. We include onlv data collected before
1995.

Since te.xts can be read in many ways, and since one of our reviewers mistook this
to be an historical account, let us at the outset explain that our purpose is not histoncal
but biogeographical. Our aim is not to provide a chronology of discovery and
document collectors' pnonty on the Kisatchie NaUonal Forest, but to provide
documentation for the occurrence of the taxa discussed. Consequently, we document
occurrences emphasizing our own collections since we can vouch for them: we have
had many vetted by experts and we know their provenance. Additionally, we
emphasize the collections in the Kisatchie National Forest herbanum (SFRP) because
we are most familiar with them. The history of botany and plant collecting in
Louisiana and on the Kisatchie National Forest has yet to be wntten (MacRoberts
1984, 1989; Ewan 1967; Sundell 1979).

Unfortunately, only some plant communities on the Kisatchie National Forest have
been systematically inventorial and censused, notably, bogs, glades, prairies, sandy
woodlands, and sandstone outcrops. Consequently, information on rare species is
biased in the direction of these communities. It is hoped that, in time, other
communities will receive equal attention.

As in any endeavor like this, we keep turning up new records and finding mistakes
in existing ones. But if we were to wait until all inconsistencies and mistakes are
corrected and each report triple checked, nothing would get published. We provide the
following account in the hope that future researchers will use it as a baseline to make
emendations.

The Kisatchie National Forest consists of six districts that comprise about 242,000
hectares, or about two percent of Louisiana (Figure 1) (Caldwell 1991). The major
community types in the Forest are upland longleaf pine forest, nparian forest, pine
flatwoods/savannah, shortJeaf pine/oak-hickory forest, and mixed hardwood-loblolly
forest (Martin & Smith 1991, 1993). There are also many inclusion communities such
as bogs, prairies, glades, outcrops, baygalls, and sandylands (Smith 1988; Smith et
al. 1989; MacRoberts & MacRoberts 1993a, 1993b, 1993c, 1994, 1995; Marun &
Smith 1991, 1993; Guillory el al. 1990). The majority of the plant survey work has
been done on the Vernon, Kisatchie, and Winn districts. The Caney, Evangeline, and
Catahoula districts have been relatively neglected except for brief surveys undertaken
in 1993 and 1994 to examine some of their more promising habitats.

About half of the endangered, threatened, and sensitive plant species (ETS) on the
Kisatchie National Forest are also on the threatened and endangered list for the Texas
National Forests and Grasslands (Anon. 1994; Orzell 1990; Natural Hentage 1994).

There are two classes of plants reported on in this paper. The first are plants that
are on both the Kisatchie National Forest ETS list and the Louisiana Natural Hentage
"Rare Plants Species" list The second are the Louisiana Natural Hentage "watch list"
species. These are species that are of interest either because their distribution is
imperfectly known or because they may be decreasing in numbers. For this work, we
use the last 1994 Natural Hentage list dated "Nov. 1994." Because records on watch
list species are not kept by the Forest Service, in the accounts that follow, information
on them is more anecdotal and scanty than that on ETS species. We use Kartesz
(1994) for nomenclature, adding synonyms where recent name changes could be
confusing.

293

PHYTOLOGIA volume 78(3):291-313 Apni 1995

Kisatchie

UouUonal

RANGER DISTRICTS

Figure 1 . Locations of Ranger Districts in the
Kisatchie National Forest.

MacRoberts & MacRoberts: Rare plants of Kisatchie National Forest

294

Figure 2. Louisiana parishes.

295 PHYTOLOGIA volume 78(3) : 29 1-3 13 Apnl 1995

We follow recent authors (e.g.. Bridges & Orzell 1989a) in giving detailed
distnbutional information in order to make our data useful to present and future
tolanists interested in the plants and habitats; they need not borrow herbarium
specimens, or more important in this case, try to locale the many site records that are
presently scattered among many agencies, organizations, and individuals and which
have no voucher specimens. However, we give only Township/Range/Section since
this will suffice for most botanical/ecological needs and is general enough to
discourage unscrupulous collectors.

In addition to Township/Range/Section, plant locations are given by Ranger
District and Parish. For the sake of completeness, using the Louisiana Natural
Heritage rare species list and other sources (e.g., Larke & Smith 1994) we give the
other panshes in Louisiana where the species is reported (Figure 2).

The following abbreviations are used:
KNF = Kisatchie National Forest

WGCP = West Gulf Coastal Plain (west Louisiana and east central Texas)
CatD = Catahoula Distnct (Grant Parish)
CnyD = Caney Distnct (Webster and Claiborne parishes)
ED = Evangeline Distnct (Rapides Parish)
KD = Kisatchie Distnct (Natchitoches Parish)
VD = Vernon District (Vernon Pansh)

WD = Winn District (Natchitoches, Winn, and Grant parishes)
LAF = Herbarium, Department of Biology, University of Southwestern Louisiana,

Lafayette, Louisiana
LSU = Herbarium, Department of Botany, Louisiana State University, Baton Rouge,

Louisiana
LSUS = Herbanum, Biology Department, Louisiana State University, Shreveport,

Louisiana
LTU = Herbarium, Biology Defjartment, Louisiana Tech University, Ruston,

Louisiana
NATC = Herbanum, Biology Department, Northwestern State University,

Natchitoches, Louisiana
NLU = Herbarium, Biology Department, Northeast Louisiana University, Monroe,

Louisicma
SFRP = Herbanum, Range N/Ianagement Research, Southern Forest Experiment

Station, U.S. Forest Service, Pineville, Louisiana
VDB = Herbarium, Botany Department, Vanderbilt University, Nashville, Tennessee

SPECIES ACCOUNTS

Amsonia ludoviciana Vail. {Martin 364 [SFRP]; MacRoberts & MacRoberts 1578
[VDB], 1629, 2209 [SFRP]). Louisiana blue star is a near Louisiana endemic with its
main populations centering on the KNF. It has been found at 21 sites m three districts:
KD (Natchitoches Pansh, nine sites, T5N R6W Sec. 29; T5N R7W Sees. 2, 14, 20,
21, 23. 37; T5N R8W Sec. 24), WD (Winn. Grant, and Natchitoches panshes. mne
sites, T13N R5W Sees. 2, 29. 30; T12N R6W Sec. 1; T9N R3W Sec. 36). and CatD
(Grant Parish, three sites, T8N R2W Sees. 16, 29; T8N R3W Sec. 1). It grows m
wet areas such as roadside ditches and npanan forest (Lemke 1987; Larke & Smith
1994). The habitat in which this species occurs has not been the focus of any specific

MacRoberts & MacRoberts: Rare plants of Kisatchie National Forest 296

survey on the KNF. Lx)uisiana blue star is also reported from Allen, Bienville,
Calcasieu, Red River, and Vernon parishes (Natural Heritage 1994).

Asclepias stenophylla A. Gray. (Urbatsch 4468[LS\J]). Narrow -leaved milkweed, a
western species that reaches its eastern limit in Lx)Uisiana, is known from a single
calcareous praine on the WD (Winn Parish, TUN R4W Sec. 7). It was onginally
reported from Louisiana by Riddell in 1852 under the synonym Aceretes angusiifolia
(Nutt.) Dene. (MacRoberts 1984, 1989) but not reported again until recently (S.
Lynch, pers. comm.; Natural Heritage 1994).

Asclepias viridiflora Raf. {MacRoberts & MacRoberts 1730 [VDB], 2032. 2439
[SFRP]). This species is on the Louisiana Natural Heritage watch list. While we have
not systematically kept records on its occurrence and the Forest Service does not
monitor it, we have noted it regularly in calcareous praines and calcareous soils in
sandstone outcrops on the KD and WD (Natchitoches and Winn parishes)
(MacRoberts & MacRoberts 1993c). It has been reported from scattered locations
statewide (MacRoberts 1989).

Astragalus crassicarpus Nutt. var. trichocalyx (Nutt.) Bameby. {Brown & Lenz
7620, 7617 [LSU]; Martin s.n. [LSU]; Thomas et al. 92314, 95456 [NLU]; Gilmore
& Smith 3561 [LSU]). Ground-plum, a mid-central U.S. species, is known from
four calcareous prairies in the WD (Winn Parish, TUN R4W Sec. 18; TUN R5W
Sees. 13, 24, 26). It was first collected on the WD in 1939 by C.A. Brown and W.
Lenz but was not correctly identified and was not relocated there until the mid-1980's
(Smith et al. 1989; Lasseigne 1973; MacRoberts 1989). It has also been reported from
Natchitoches tmd Cameron parishes (Natural Heritage 1994).

Burmannia biflora L. {MacRoberts & MacRoberts 2484 [SFRP]; Thomas 33333,
74027 [NLU]). Two locations for Northern Burmannia are known on the KNF. One
is on the CatD (Grant Parish, T8N Rl W Sec. 1 1) in a hillside wooded seep, the other
is on the VD (Vernon Parish, TIN R6W Sec. 31) in riparian woodland. This is a
southeastern and coastal plain species that is near the penphery of its range in the
WGCP (Thomas 1970). It has also been reported from Bienville, DeSoto, Ouachita,
and Webster jjarishes (Natural Heritage 1994).

Calopogon barbatus (Walt.) Ames. {Holmes 3007 [NATC]; MacRoberts &
MacRoberts 304 [LSUS], 660 [VDB], 1917 [SFRP], 1610 [NLU]). We know of
five locations for Bearded grass-pink on the KNF. Four are in the VD (Vernon
Parish, TIS R8W Sec. 26; TIN R7W Sec. 29; and TIN R5W Sec. 20) and one is in
the KD (Natchitoches Parish, T6N R7W Sec. 22). The KD site has been known since
the 1970's (Hdmes & Mathies 1980; MacRoberts & MacRoberts 1988). Four of the
five populations occur in bogs and one is in the upslope mesic area above a bog
(MacRoberts & MacRoberts 1993b; Hart & Lester 1993). This is a coastal plain
species that reaches its western limits in the WGCP. It has also been reported from
Allen, Jefferson Davis, and St. Tammany panshes (Natural Heritage 1994), but since
it is likely that C. barbatus and C. pallidas Chapman could be confused, the published
distributions of these species need verification (MacRoberts 1989; Thomas & Allen
1993a).

Camassia scilloides (Raf.) Cory. {Allen 17637 [NLU]; Mclnnis & Smith s.n.
[LSU]). Wild hyacinth, an eastern species, has been found at two locations in the WD

297 PHYTOLOGIA volume 78(3): 29 1-3 13 Apni 1995

(Winn Parish, TUN R5W Sees. 24, 35). It is a calcareous forest species (Larke &
Smith 1994). It has also been reported from Bossier, Caddo, Natchitoches, and
Webster panshes (Natural Heritage 1994).

Carex hyalina Boott. {Allen 17890 [NLU]). This species has been found at one
location on the ED (Rapides Parish, T3N R2W Sees. 23, 25). It occurs in wet
bottomland hardwoods (Thomas & Allen 1993b). It has only been recorded for
Rapides Pansh in Louisiana (Natural Heritage 1994).

Carex meadii Dewey. {MacRoberts & MacRoberts 1889, 2216 [VDB], 1903, 1924,
2186 [SFRPl, 1889, 2289 [NLU], 7970 [LSU]). Mead's sedge, a widespread
northern species, is known from five locations on the KD (Natchitoches Parish, T5N
R6W sec. 4; T5N R7W Sec. 2; T6N R7W Sec. 35; T6N R8W Sees. 28, 29, 32; T5N
R8W Sec. I) and two locations on the WD, (Winn and Grant panshes T9N R2W Sec.
6; T12N R4W Sec. 34). It grows in sandstone outcrop communities and in calcareous
prairies (Williams 1977; MacRoberts & MacRoberts 1993c). It is also reported from
Acadia, Allen, and Jefferson Davis panshes (Natural Heritage 1994).

Carex microdontaToney & Hook. {MacRoberts & MacRoberts 1945, 2220 [VDB],
2215, 2220, 2284 [SFRP], 2272 [NLU], 2285, 2286 [LSU]). Small-toothed sedge,
a species of the south central U.S., has been found at 20 sites on the WD (Winn and
Grant panshes. TUN R4W Sees. 7, 8, 18; TUN R5W Sees. 13, 24, 25, 26, 35;
T12N R4W Sees. 33, 34, 35; T9N R2W Sees. 5, 6). It is abundant in calcareous
prairies. It has also been reported from Acadia, LaSalle, and Vernon parishes (Natural
Heritage 1994).

Carex tenax Chapman. {MacRoberts & MacRoberts 2282 [VDB], 2295, 2296, 2339
[SFRP], 2294 [NLU], 2334 [LSU]). Strong sedge has been found at eighteen sites in
KD (Natchitoches Parish, T7N R6W Sec. 64; T6N R6W Sec. 6; T6N R7W Sees. 1 ,
2, 4, 5, 9. 15, 16, 17, 18, 20, 21; T7N R7W Sees. 33, 34, 36) and three in WD
(Natchitoches Pansh, T13N R6W Sees. 7, 18) (MacRoberts & MacRoberts 1994). It
occurs in upland xenc sandhill woodlands. It was for a long time known from only
one site in Louisiana (Williams 1977; MacRoberts 1989). This species is also
recorded from Vernon Parish (Thomas & Allen 1993a).

Ceimothus herbaceous Raf. {MacRoberts & MacRoberts 2018 [VDB]). Prairie
redroot has been found in one calcareous prairie on the WD (Winn Pansh, Tl IN R5W
Sec. 13; TUN R4W Sec. 18). The only other Louisiana location is one calcareous
prairie remnant in Caddo Parish (Natural Heritage 1994).

Chamaelirium luieum (L.) A. Gray. Fairy wand, an eastern species, has been found at
one wooded seep on the KNF (WD, Natchitoches Parish, T13N R6W Sec. 8). The
species was not collected because of its rarity and because there were only seven stems
in the population. It has also been recorded from Lincoln, Ouachita, St. Helena, St.
Tammany, Tangipahoa, Washington, and W. Feliciana panshes (Natural Hentage
1994).

Cheilamhes Umosa (Michx.) D.C. Eat. {MacRoberts & MacRoberts 2401 [VDB];
Thomas <&. Carroll 73889 & 2383 [SFRP]). Hairy lip-fem was first found on the KD,
Natchitoches Parish, in the late 1930's (Brown & Correll 1942). It has been
momtored over the years and still grows in the same place in crevices of sandstone

MacRoberts & MacRoberts: Rare plants of Kisatchie National Forest 298

boulders in shaded upland longleaf pine forest (T6N R8W Sec. 27). While we have
searched many boulders in the KNF for this species, it is sull known only from the
onginal locality. From earlier accounts, it apparently has become more restncted at the
onginal locauon (Brown & Correll 1942; Thieret 1980). It is widespread in the
eastern U.S.

Cyperus grayioides Mohlenbrock. (MacRoberts & MacRoberts 1801. 2072 [VDB],
1798, 2071 tSFRP], 2070 [NLU], 1803 [LSU]). Mohlenbrock's umbrella sedge has
been collected many times in eastern Texas and western Louisiana. There are seven
known sites for it on the KD (Natchitoches Parish, T6N R6W Sec. 6; T6N R7W
Sees. 1,2. 18, 21, 34; T6N R8W Sec. 21; T7N R7W Sec. 33), and four sites on the
WD (Winn and Natchitoches panshes, T13N R7W Sees. 2, 11; T13N R6W Sec. 7;
T13N R5W Sec. 32; T12N R5W Sec. 28). It grows in open areas in upland and
stream terrace xenc sandhill woodlands (MacRoberts & MacRoberts 1994). It seems
to be associated with deep sands especially in Illinois, Missoun, Texas, and Louisiana
(Badges & Orzell 1989a; Orzell 1990; Carter & Bryson 1991). In Louisiana it is
recorded from Allen, Bienville, and Vernon panshes (Natural Hentage 1994).

Cypripedium kentuckiense C.F. Reed. Kentucky yellow lady's slipper occurs at
scattered localities in the south central U.S. extending south into the WGCP. It has
been reported from many northern parishes in Louisiana, but most records are
relatively old (MacRoberts 1989; Thomas & Allen 1993a; Natural Hentage 1994). Its
only known occurrences in the KNF are on the CatD (Grant Parish, one site, T9N
RIW Sec. 32) and the ED (Rapides Pansh, three sites, T2N R3W Sees. 1 1, 12). It
occurs in mixed hardwood-loblolly forest and hardwood slope forest. Because of its
extreme rarity, we photographed it rather than collect it.

Dodecatheon meadia L. {MacRoberts & MacRoberts 2240 [VDB,SFRP]). Shooting
star has been found at one location on the KD (Natchitoches Parish, T5N R7W Sec.
14) in mixed hardwood-loblolly forest. It is a widespread northern species that
reaches its southern limit in the WGCP. It is also reported from Bossier juid Caddo
parishes (Natural Heritage 1994).

Draba cunei/olia Nutt. ex Torrey & A. Gray. {Brown 5184 [LSU]). Wedge-leaf
whitlow-grass is widely scattered through the south and central U.S. Brown collected
It in 1934 in the vicinity of Goldonna, Louisiana The herbanum label says "Goldiana
[sic]. La., Winn Parish," but Brown's field notebook says only "Goldiana" [sic],
which is in Natchitoches Parish. And his notebook shows that on March 27 he had
moved around a good bit collecting in both panshes. The exact habitat is also unclear
on the herbarium label it says "mixed hardwoods," but his notebook reads "roadside,
mixed pine and hardwoods." The Natural Hentage data files record it as T12N R5W
Sec. 28 assuming it was collected in Winn Parish. With this confusion, it is probably
safe to say only that it was collected in the vianity of Goldonna on a private inholding
in the Winn District. This species is also known from Caddo Parish (Nauiral Heritage
1994).

Eriocaulon texense Kbm. {MacRoberts <& MacRoberts 1589 [VDB], 1579 [SFRP]).
This species is on the Louisicina Natural Hentage watch list. While we have not kept
precise records on its occurrence, we have found it only in VD (Vernon Pansh) bogs
where it can be plentiful (MacRoberts & MacRoberts IS^b). It is also reported from
a few other west central panshes (MacRoberts 1989; Thomas & Allen 1993a).

299 PHYTOLOGIA volume 78(3): 29 1-3 13 Apnl 1995

Eriogonum longifolium Nutt. (MacRoberls & MacRoberls 1800 [VDB], 1797
[SFRP]). Long-leaved wild buckwheat, a Great Plains species, is known from one
stream terrace sandhill woodland in an inholding on the WD (Wmn Parish, T12N
R5W Sees. 28, 29). It is also reported from Caddo and Vernon panshes (Natural
Hentage 1994).

Eriogonum multiflorum Benth. (MacRoberls & MacRoberls 1808 [VDB], 1850, 2084
[SFRP], 1820 [NLU]). Many-flowered wild buckwheat, a species that ranges from
northeast Mexico to Oklahoma and western Lxiuisiana, has been located at ten sites in
two districts (WD, Winn and Natchitoches panshes, five sites, T13N R5W Sees. 7,
33; T13N R6W Sees. 2, 7; T13N R7W Sees. 2, 11; KD, Natchitoches Parish, five
sites, T6N R6W Sec. 6; T6N R7W Sees. 5, 20; T7N R6W Sec. 64; T7N R7W Sec.
36). It grows in xenc sandhill woodlands. It also is reported from Bienville, Caddo,
and Webster parishes (Natural Hentage 1994).

Erythronium rostratum W. Wolf. {MacRoberls & MacRoberls 2169 [VDB], 2169,
2204 [S¥RP], 2169 [LSU]). Yellow trout-lily, a species of the south central U.S., is
on the Louisiana Natural Heritage watch list. We found it at one location in the ED
(Rapides Pansh) and two locations on the WD (Natchitoches Pansh). It prefers
npanan forest (Krai 1966). It has been found in a number of north Louisiana panshes
(Thomas & Allen 1993a; MacRoberts 1989).

Euphorbia bicolor Engelm. & A. Gray. {MacRoberls & MacRoberls 2415 [NLU],
2435 [SFRP]). This western praine species, called snow-on-the-praine, while
common in Texas is rare in Louisiana and is on the Louisiana Natural Hentage watch
list It is known to occur in one WD (Winn Parish, TIN R4W Sees. 7, 8) caicareous
praine. It is also known from Vernon Pansh (MacRoberts 1989).

Evolvulus sericeus Sw. {MacRoberts & MacRoberls 1948 [VDB]). Creeping
morning glory is on the Louisiana Natural Hentage watch list. While we have not
regularly searched for it, we found it in sandstone outcrops on the KD (Natchitoches
Pansh, MacRoberts & MacRoberts 1993c). It has also been reported from a number
of parishes in northeastern and southwestern Louisiana (MacRoberts 1989).

Heiiotropium lenellum (Nutt.) Torrey. {MacRoberls & MacRoberts 2019 [VDB],
2019, 2417 [SFRP]). Slender heliotrope, a calcareous praine species of the south
central U.S., has been found at fourteen sites on the WD (Winn and Grant panshes,
TUN R4W Sees. 7, 18; TUN R5W Sees. 13, 24, 26; T9N R2W Sees. 5, 6, 7, 8)
(Smith el al. 1989). It is also known from Vernon Pansh (Natural Hentage 1994).

Hexalectris spicata (Walt.) Bamh. {Rhodes 16606 [LTU]). Crested coral-root has
been found once on the CnyD (Claiborne Parish, T23N R4W Sec. 34). Recent
attempts to relocate these populations have failed. It is an uncommon but wide-
ranging species extending across the southern United States to New Mexico and
Mexico. In our area it is found in mixed hardwood loblolly and hardwood slope forest
(Larke & Smith 1994). It has also been reported from a number of central and north
Louisiana parishes (Natural Hentage 1994; Thomas & Allen 1993a; MacRoberts
1989).

MacRoberts & MacRoberts: Rare plants of Kjsatchie National Forest 300

Houstonia purpurea L. var. calycosa A. Gray. [SY = Hedyolis purpurea (L.) Torrey
& A. Grav var. calycosa (A. Grav) Fosberg]. (MacRoberts & MacRoberts 2206
[VDB], 2205, 227i 2273 [SFRP]' 2207 [LSU]). Purple bluet, a calcareous praine
sf)ecies has been found at nineteen sites in the WD (Winn and Grant parishes, T9N
R2W Sees. 5, 6; TUN R4W Sees. 7, 8, 18; TUN R5W Sees. 13, 23, 24, 26, 35;
T12N R4W Sees. 33, 34. 35) (Smith et al. 1989; Martin & Smith 1991). It has also
been reported from Bossier, Caddo, LaSalle, Caldwell, and Webster parishes (Natural
Heritage 1994).

Koeleria macrantha (Ledeb.) J. A. Sehultes. (MacRoberts & MacRoberts 2021a
[VDB], 2440 [SFRP]). June grass, a widely distributed grass in North Amenea, is
known from three calcareous praines on the WD (Winn Parish, TUN R4W Sec. 18;
TUN R5W Sees. 13, 26). It is reported from no other pansh (Smith et al. 1989;
Allen 1992; Natural Hentage 1994).

Lachnocaulon digynum Kbm. (MacRoberts & MacRoberts 531, 1776 [VDB],
1777B. 2471 [SFRP], 1470 [LSU]). Pineland bog buttons, a coastal plain species,
was not reported for the WGCP unul the late 1980's (Bndges & Orzell 1989a;
MacRoberts 1989; Thomas & Allen 1993a). It has been found in one bog on the WD
(Natchitoches Parish, T12N R6W Sec. 12) and in just over 50 bogs on the VD
(Vernon Pansh; there are so many widely scattered sites that we do not give locations
here) (Hart & Lester 1993; MacRoberts & MacRoberts 1993b). It is also recorded
from Beauregard, Sabine, St. Tammany, and Washington parishes (Natural Heritage
1994; MacRoberts 1989; Julia Larke, pers. comm.).

Liatris tenuis Shinners. (Grelen 1821 [NLU]). Slender gay-feather, a WGCP
endemic, has been found at a single location on the VD (Vernon Parish, TIS R6W
Sec. 6) in upland longleaf pine forest (Gandhi & Thomas 1989). It is also reported
from Caddo, Calcasieu, DeSoto, Rapides, and Sabine parishes (Natural Heritage
1994).

Lilium michauxii Poir. (MacRoberts & MacRoberts 2457 [SFRP]). Carolina lily is
on the Louisiana Natural Heritage watch list It has not yet been carefully monitored.
We have found it in a couple of beech-magnolia riparian forest areas on the KD
(Natchitoches Parish). It is probably locally common but generally rare on the forest.
It has been reported from a number of parishes in western and southeastern Louisiana
(MacRoberts 1989; Thomas & Allen 1993a).

Uthospermum caroliniense (Walt, ex J.F. Gmel.) MacM. (MacRoberts <&
MacRoberts 2223 [SPT^^]). Puccoon is on the Louisiana Natural Hentage watch list.
It appears to be locally common in upland xenc sandhill woodlands and upland
longleaf pine forest, notably on the KD (Natchitoches Pansh) and WD (Natchitoches
and Winn panshes) «md in central and northwestern Louisiana (MacRoberts 1989).

Lycopodiella cernua (L.) Pichi Sermolli [SY = Lycopodium cernuum L.]. (Holmes
3360 [NATC]; Thomas 74133 [NLU]). Nodding clubmoss, a coastal plain species,
has been found at only one site in the KNF. In the 1970's it was found in a bog on
the KD (Natchitoches Parish, T6N R7W Sec. 22) (Thomas et al. 1978). While it
apparently persisted at this location for a number of years, it has not been found there
for more than a decade (MacRoberts & MacRoberts 1988, 1993b). It is also reported
from Ouachita and St. Tammany parishes (Natural Hentage 1994).

301 PHYTOLOGIA volume 78{3):29l-3 13 Apnl 1995

Lyonia mariana (L.) D. Don. (Latimore Smith s.n. [LSU]). Staggerbush lyonia, an
eastern species that reaches its western limit in east Texas, is known from one site on
the CnyD (Claiborne Parish. T2N R4W Sec. 13). It has also been found in Caddo
Parish (Natural Heritage 1994).

Maianthemum racemosum (L.) Link [SY = Smilacina racemosa (L.) Desf.]. {Thierei
24686 [LAF]). There is only one known occurrence of False solomon seal, a northern
species, on the KNF (Thieret 1967a). This species has been repeatedly searched for at
the onginal site (CnyD, Claiborne Parish, T22N R4W Sec. 10) but without success.
It occurs in mixed hardwood slope forest. This is the only parish for which this
species has been reported (Natural Heritage 1994).

Malaxis unifolia Michx. {MacRoberts & MacRoberts 341 [VDB]). We have found
Green adder's mouth, a Lx)uisiana Natural Heritage watch list species, at several
scattered locations on the KD and WD, Natchitoches and Winn panshes. There has
been no systematic survey for it. It is never common where found and apjsears to
prefer shaded areas in mixed hardwood-loblolly forest, often under dogwood (Cornus
florida L.). Its secretive habit and short stature may account for its apparent rareness.
It has been found in many parishes in northern and southeastern Louisiana
(MacRoberts 1989; Thomas & Allen 1993a).

Mayaca fluviatilis Aubl. [SY = Mayaca aubletii Michx.]. {Hampsien el al. s.n.
[SFRP]; MacRoberts & MacRoberts 2160, 2466 [VDB]). Bog moss, a coastal plain
species, is known only from the ED (Rapides Parish). A survey for this species in
1993 revealed six ED locations (T3N R2W Sees. 17, 18, 20; T3N R3W Sees. 24,
25). It occurs in shallow water in bayhead swamps. It is also reported from
southeastern Louisiana in St. Tammany, Tangipahoa, and Washington panshes
(Natural Heritage 1994).

Monotropa hypopithys L. {Rhodes 14856 [LTU]; Can & Bruser s.n. [SFRP]).
Amencan pinesap, a widespread northern species, has been found in several locations
in northern Louisiana, but only three are on the KNF. Two are on the CnyD (T22N
R4W Sec. 10; T23N R4W Sec. 34) in Claiborne Parish, and one is on the WD in
Winn Pansh (TUN R5W Sec. 13). One of the CnyD populations has not been
relocated in recent years. It grows in shortleaf pine/oak-hickor>' forest and mixed
hardwood loblolly forest in filtered shade (Larke & Smith 1994). It is found in a
number of parishes on the northern tier of Louisiana (Natural Hentage 1994).

Obolaria virginica L. (Carr, MacRoberts, & MacRoberts 2168 [VDB]; MacRoberts &
MacRoberts 2254 [SFRP]). Pennywort is on the Louisiana Natural Heritage watch
list. We have encountered it only twice on the Kisatchie National Forest in deeply
shaded hardwood slope forest (ED, Rapides Pansh, T2N R3W Sees. 11, 12). How
common it may be is not known since very little surveying has been done in mi.xed
hardwood slope forest. It is known from a number of north Louisiana panshes
(MacRoberts 1989).

Orobanche uniflora L. {Mclnnis 3990 [personal collection]). Broomrape, a
widespread northern species that also occurs in Texas and Arkansas, was first reported
for Louisiana in 1852 (MacRoberts 1984, 1989). But it was not reported again until
1990 when it was found in upland longleaf pine forest on the VD (Vernon Pansh,

MacRoberts & MacRoberts: Rare plants of Kisatchie Nauonal Forest 302

TIN R8W Sec. 36) (Hart & Lester 1993). It has also been reported for Rapides
Parish (Natural Heritage 1994).

Panicum flexile (GdXi.) Scribn. {Brown & Lenz 8236 [LSU]; Smith & GiUnore 3713
[LSU]; Thomas & Allen 94224 & 13430 [NLU]). Wiry witchgrass, a widespread
eastern species that is uncommon at most localities where it occurs, is recorded from
four calcareous prairies in the WD (Winn and Grant parishes, T9N R2W Sec. 5; T 1 IN
R4W Sec. 18; Tl IN R5W Sees. 13, 26) (Smith el al. 1989; Allen 1992). It has also
been reported for Caddo, Ouachita, and Vernon parishes (Natural Hentage 1994).

Paronychia drummondii Torrey & A. Gray. {MacRoberts & MacRoberts 2083
[VDB], 2281, 2344 [SFRP], 7805 [LSU]). Drummond's nailwort, a Texas near
endemic that reaches its eastern limit in western Louisiana, is known from eight
locations on the WD (Natchitoches Pansh, T12N R5W Sec. 8; T31N R6W Sees. 7,
12, 18; T13N R7W Sees. 2. 11; Winn Parish, T12N R5W Sec. 28; T13N R5W Sec.
17). It occurs in upland and stream terrace xeric sandhill woodlands. It has also been
recorded from Bienville and Caddo parishes (Natural Heritage 1994).

Pediomelum hypogaeum (NutL ear Torrey & A. Gray) Rydb. [SY = Psoralea subulata
Bush]. {MacRoberts & MacRoberts 2236 [LSU,VDB,SFRP,NLU]). Awl-shaped
scurf-pea occurs in east Texas, south Oklahoma, southwest Arkansas, and western
Louisiana (Lasseigne 1973). It has been found at one location on the KD
(Natchitoches Parish, T7N R7W Sec. 36) in upland xenc sandhill woodlands. It also
has been found in Caddo Parish (Natural Heritage 1994).

Penstemon murrayanus Hook. {MacRoberts & MacRoberts 1612 [VDB]; Grelen 2052
[SFRP]). The only known KNF site of cupleaf beardtongue is in a stream terrace
sandhill woodland in an inholding on the WD (Winn Parish, T12N R5W Sees. 28,
29). This species is found in Arkansas, Oklahoma, Texas, and in northwestern
Louisiana (Vincent 1982) where it has been found in Caddo Parish (Natural Hentage
1994).

Phacelia strictiflora (Engelm. & A. Gray) A. Gray. {MacRoberts & MacRoberts 2211
[VDB], 2532 [SFRP]; Thomas & Kessler 75762 <& 1334 [NLU]). Phacelia, a species
of the south central U.S., is known from two WD stream terrace sandhill woodlands
(Winn Parish, T13N R5W Sees. 32,33;T12N R5W See. 28) (Thieret 1967a). It has
also been recorded for Caddo, Natchitoches, and Bienville parishes (Natural Heritage
1994).

Platanthera blephariglottis (Willd.) Lindl. White-fringed orchid, an eastern species,
was first found as a single stem on the VD (Vernon Parish) in August 1990 (Parker
1990). In September 1992, the site was revisited and three stems were found at two
locations. It occurs in TIN R6W Sec. 14 in bog habitat. The species has not been
collected (although it has been photographed) because there were too few individuals
(MacRoberts & MacRoberts 993b; Hart & Lester 1993). It is also reported from St.
Tammany Parish (Natural Hentage 1994).

Platanthera Integra (Nutt.) A. Gray ex Beck. {MacRoberts & MacRoberts 1146
[VDB], 886 [SFRP]). Yellow fnngeless orchid, a coastal plain bog species, was
unknown on the KNF until 1980 (Holmes 1983). It was known from only one site
until systematic bog surveys revealed it at nine KD (Natchitoches Parish, T6N R7W

303 PHYTOLOGIA volume 78(3): 29 1-3 13 Apnl 1995

Sees. 11, 14, 22. 24, 25, 35, 36; T5N R8W Sec. 36) and twelve VD bogs (Vernon
Parish, TIN R5W Sec. 33; TIN R6W Sees. 14. 15, 17, 23, 26, 35; TIN R7W Sees.
21, 28; TIS R8W Sees. 14, 15, 17, 20, 21) (MaeRoberts & MacRoberts 1988,
1990a, 1990b, 1991, 1993b; Hart & Lester 1993). Its range is New Jersey to Honda
and along the Gulf Coastal Plain to Texas, where it is very rare (Bndges & Orzell
1989a; Orzell 1990; Grace 1993). In Louisiana it is also reported from Beauregard
and St. Tammany parishes (Natural Rentage 1994).

Polanisia erosa (Nutt.) litis. {MacRoberts & MacRoberts 1802 [VDB], 2340, 2431
[SFRP], 1821 [LSU]). Clammy weed, a species apparently confined to Texas,
Oklahoma, Arkansas, and western Louisiana, has been found at four sites in upland
and stream terrace xeric sandhill woodlands in the KD (Natchitoches Parish, T6N
R7W Sees. 5, 12; T7N R7W Sees. 29, 36), and three sites on the WD (Winn and
Natchitoches panshes, T12N R5W Sec. 28; T13N R6W Sec. 7) (MacRoberts &
MacRoberts 1994). It has also been reported from Bienville and Sabine panshes
(Natural Hentage 1994).

Polygala verticillata L. {MacRoberts & MacRoberts 1975 [VDB], 1968. 2363
[SFRP], 1666 [LSU]). Prairie milkwort is on the Louisiana Natural Heritage watch
list It has not been monitored on the KNF. While often difficult to find because of its
small size, we have found it regulariy in sandstone outcrop and calcareous prairie
communities in Natchitoches and Winn parishes (KD and WD) (MacRoberts &
MacRoberts 1993c). It appears to be scattered widely around the state (MacRoberts
1989).

Polygonella arnericana (Fisch. & C.A. Mey.) Small. {MacRoberts & MacRoberts
1799 [VDB], 2428 [SFRP]). Southern jointweed, a spjecies found in xenc sandylands
from South Carolina to New Mexico, occurs at one xeric stream terrace sandhill
woodland in an inholding on the WD (Winn Parish, T12N R5W Sec. 28). It has also
been recorded from Bienville, Caddo, and Sabine parishes (Natural Hentage 1994).

Polygonella polygama (Vent) Engelm. & A. Gray. {MacRoberts & MacRoberts 2128
[VDB], 920, 2147 [SFRP]; Gilmore. Smith, & Johnson 3729 [NLU]). October
flower, an upland xeric sandhill woodland plant, was first found in Louisiana in 1988
on the WD (Natchitoches Parish, T13N R6W Sees. 7, 18) where it is abundant
(Johnson & Johnson 1990). However, it is not known from any other locality in
Louisiana. It ranges across the southern U.S. from Virginia to Texas.

Polytaenia nuttalUi DC. {MacRoberts & MacRoberts 2377 [SFRP]; Thomas et al.
92238, 104893 [NLU]). Prairie parsley is on the Louisiana Natural Heritage watch
list It has not been specifically monitored so records are not available, but it occurs in
several calcareous prairies on the WD (Winn Pansh) (Smith et al. 1989). It occurs m
several scattered locations in western and central Louisiana (MacRoberts 1989).

Prenanthes barbata (Torrey & A. Gray) Milstead. {MacRoberts & MacRoberts 2082
[VDB]; Thomas 78298 [SFRP]). Barbed rattlesnake-root, a southeastern species, is
found in mixed hardwood-loblolly and riparian forest. It has been found at rune
locations on the WD (Winn Pansh, TON R5W Sees. 2, 3, 4. 9, 10, 14, 15, 22, 24;
Natchitoches Pansh, T13N R6W Sec. 1), and one location on the ED (Rapides Pansh,
T3N R2W Sec. 29) (Gandhi & Thomas 1989). It is also reported from Vernon Pansh
(Natural Hentage 1994).

MacRoberts & MacRoberts: Rare plants of Kisaichie National Forest 304

Pteroglossaspis ecristata (Femald) Rolfe. [SY = Eulophia ecristala (Femald) Ames].
{Thomas et al. 12265 [NLU]). Wild coco, a southeastern species, is known from
only one site in the KNF. This is a wooded seep in the CatD (Grant Parish, T8N
RIW Sec. 11) (Thomas 1972). Although found in the late 1960's, it has not been
regularly monitored. It was apparently found at this location again in 1978 but not
relocatal in a 1986 survey. We did not relocate it in our survey of the site in 1994. It
has cilso been recorded for Allen, Beaureg2U"d, Jefferson Davis, St. Tammany,
Tangipahoa, and Washington parishes (Natural Heritage 1994).

Rhynchospora, macra (C.B. Clarke) Small. (MacRoberts & MacRoberts 1432, 1469
[VDB], 1430, 1467 [SFRP], 1464, 1494 [NLU], 1434, 1787 [LSU]). Large
beakrush, a coastal plain species that reaches the penphery of its range in the WGCP,
was first found in Louisiana in the eariy 1970's in a bog on the KD. This remained
the only known WGCP site until extensive bog surveys on the VD and KD in the late
1980's and early 1990's located it in eight KD (Natchitoches Parish, T5N R8W Sec.
3; T6N R6W Sec. 34; T6N R6W Sees. 11. 17, 22) and 22 VD (Vernon Parish, TIS
R6W Sec. 6; TIS R7W Sees. 2, 3; TIS R8W Sees. 1, 13, 20, 21; TIN R5W Sec. 28;
TIN R6W Sees. 13, 14, 15, 17, 20, 21, 22, 23, 24) bogs (MacRoberts &
MacRoberts 1988, 1992, 1993b; Buras 1993; Hart & Lester 1993). It has not been
reported from any other parishes in Louisiana.

Rhynchospora oligaruha A. Gray. {MacRoberts & MacRoberts 956 [LSU], 707. 769,
924 [VDB], 934, 1259 [SFRP]). This bog fidel is on the Natural Heritage watch list.
We have found it to be abundant in most bogs in the WD. KD, and VD (Vernon a.id
Natchitoches parishes). Thomas & Allen (1993a) also report it from Beauregard and
Allen parishes.

Rudbeckia laciniata L. {MacRoberts & MacRoberts 2468 [SFRP]). Goldenglow, a
watch list species, is known from one riparian woodland on the ED (Rapides Parish,
T3N R2W Sees. 28, 29). It is also reported from Caddo. Evangeline, West Feliciana,
and Washington parishes (MacRoberts 1989; Gandhi & Thomas 1989). Gandhi dc
Thomas ( 1989) state that R. laciniata is cultivated and can escape. Whether or not the
population in Rapides Parish is natural is not known.

Rudbeckia scabrifolia L.E. Brown. {MacRoberts & MacRoberts 859, 1686, 1747
[VDB], 1468, 1748, 2331 [SFRP]). Sabine coneflower, a recenUy recognized
WGCP endemic (Brown 1986). has been found in about 50 VD (Vernon Pansh) bogs
and baygalls (there are so many sites that we do not give locations except to point out
that the species appears to be confined to the western half of tlie VD). It also occurs in
two WD (Winn and Natchitoches panshes, T13N R5W Sec. 9; T13N R7W Sec. 2)
sites; one is a bog, the other a wooded seep (Bridges & Orzell 1989a; MacRoberts &
MacRoberts 1993b; Hart & Lester 1993; Orzell 1990). It has also been recorded from
Sabine Parish (Natural Heritage 1994).

Sabatia macrophylla Hook. {MacRoberts & MacRoberts 1722 [VDB], 1704 [SFRP];
Grelen & Thomas 1780 [SFRP]). Large-leaved rose gentian, a coastal plain species
that reaches the limit of its range in the WGCP. has been found in about 50 VD
(Vernon Pansh) bogs. Since these are scattered over the entire distnct. we do not give
precise locations (MacRoberts & MacRoberts 1993b; Hart & Lester 1993). It has also

305 PHYTOLOGIA volume 78(3): 29 1-3 13 Apnl 1995

been reported for Beauregard, St. Tammany, and Washington parishes (Natural
Hcntage 1994).

Schoenolirion crocewn (Michx.) Wood. (MacRoberts & MacRoberts 1586 [VDB],
1920 [SFRP]). Yellow sunny-bells is on the Louisiana Natural Hentage watch list. It
has not been monitored on the KNF. It is abundant in some, but by no means all,
bogs on the VD (Vernon Pansh). It has been reported from Beauregard,
Natchitoches, and Sabine panshes (MacRoberts & MacRoberts 1993b; Thomas &
Allen 1993a).

Schoenolirion wrightii Sherman. {MacRoberts & MacRoberts 1901 [VDB], 1914
[LSU.NLU], 1914, 1921 [SFRP]). Texas sunny-bells was first found on the KNF m
1993 on sandstone outcrops. A survey of nineteen outcrops located it in five. At one
site up to 1000 plants were found. All sites are on the KD (Natchitoches Parish, T6N
R8W Sees. 27, 28, 29, 32; T6N R7W Sec. 35) where the plant grows on unshaded
thin soils on open rock pavement. In the KNF, Texas sunny-bell is always associated
with Talinum parviflorum NutL (MacRoberts & MacRoberts 1993c). The species has
an unusual distribution of disjunct populations from Alabama, Arkansas, Louisiana,
and Texas (Nixon & Ward 1981; Oraell 1990; Sherman & Becking 1991). These
KNF populations may be some of the most protectable in the world. It has also been
found in Red River Parish (Natural Heritage 1994).

Selaginella arenicola Underw. subsp. riddellii {Man EselL) R. Tryon. (MacRoberts <&
MacRoberts 1809 [VDB], 1779 [SFRP]; Thomas & Kessler 75765 & 1337 [SreP]).
Riddell's spikemoss has been known since the middle of the last century from areas
now probably part of the KNF (Brown & Correll 1942; Thieret 1980). On the KD. it
is known from nine sites (Natchitoches Parish, T6N R6W Sec. 21; T6N R7W Sec. 5;
T6N R8W Sees. 27, 29, 32, 35, 36; and T7N R7W See. 36) (MacRoberts &
MacRoberts 1994). It occurs in two communities: sandstone outcrops, and upland
and stream terrace xerie sandhill woodland. In the former, it grows on sandstone
pavements (MacRoberts & MacRoberts 1993c), in the latter in deep xenc sands. On
the WD (Natchitoches and Winn parishes) it is known from three sites in sandy
woodlands(T13NR6WSec. 7;T13NR5WSee. 32;T12NR5WSee. 28). This is a
coastal plain species that is rare in the WGCP. It has also been reported from
Bienville, Caddo, Sabine, and Vernon parishes (Natural Heritage 1994).

Solidago auriculata Shuttlw. ex S.F. Blake. {MacRoberts & MacRoberts 2498
[SFRP], 2495, 2496 [NLU], 2497 [LSU]). Clasping goldenrod is a Louisiana
Natural Heritage watch list species that is sometimes abundant in mi.xed hardwood
slope forest. We have found it at several locations on the KD (Natchitoches Pansh).
It has also been reported from several northwestern parishes (Gandhi & Thomas
1989).

Sporobolus ozarkanus Femald. {Gilmore & Smith s.n. [NLU]; Mclnnis & Smith
3695 [LSU]). Ozark dropseed has been found at one calcareous praine in the WD
(Winn Pansh, TUN R4W Sec. 18; TUN R5W Sec. 13) (Smith er a/. 1989). It also
occurs in Missouri, Arkansas, and Texas. It has yet to be reported from any other
peirish in Louisiana (Natural Heritage 1994; Allen 1992).

Stenanthium graimneum (Ker-Gawl.) Morong. {MacRoberts & MacRoberts 737
[VDB], 811, 1724 [SFRP]). Featherbells is a Louisiana Natural Hentage watch list

MacRoberts & MacRoberls: Rare plants of Kisatchie National Forest 306

species. We have found it in several communities on the KD (Natchitoches Pansh),
often in large numbers, specifically in damper areas of upland longleaf pine forest and
in nparian forest. It has also been reported from Beauregard, Caddo, Red River,
Sabine, St. Tammany, Washington, and Winn panshes (Thomas & Allen 1993a).

Streptanthus hyacinthoides Hook. {MacRoberts & MacRoberts 2298 [VDB], 2332,
2347 [SFRP]). Smooth twistflower occurs in upland and stream terrace xenc sandhill
woodlands at two locations on the WD (Winn Parish, T13N R5W Sec. 32; T12N
R5W Sec. 28) and one location on the KD (Natchitoches Parish, T7N R7W Sec. 36).
It is also found in Bienville and Caddo panshes (Natural Heritage 1994), and in
Arkansas, Texas, and Oklahoma.

Sty lismaaqmtica(W^ ail.) Raf. (MacRoberts & MacRoberts 1696 [SFRP]). Stylisma
is on the Louisiana Natural Heritage watch list. We know it from only one site on the
VD (Vernon Parish). It was found in an open mesic area just above a bog in
conjunction with Viola kmceolata L. and Calopogon barbatus. It appears to be
scattered around the state (MacRoberts 1989).

Taerudia integerrima (L.) Drude. (MacRoberts & MacRoberts 2180 [VDB], 2372
[SFRP]). Yellow pimpernel is an eastern species that is rare in the WGCP. It has
been found at three locations on the WD (Natchitoches and Winn panshes, T13N
R6W Sec. 32; TUN R4W Sec. 18) where it occurs in rip>arian calcareous forest
(Larke & Smith 1994). It is also reported from Bienville, Caddo, and Caldwell
parishes (Natural Heritage 1994).

Talinum parvijlorum Null. (MacRoberts <& MacRoberts 1759, 1780 [VDB], 2056
[SFRP], 1493, 1758 [LSU]. Small-flowered flame flower, a Great Plains species,
was known from only a few sites on the KD (Natchitoches Pansh) since the 193()'s .
A survey of sandstone outcrop communities in 1992 and 1993 located it at 24 sites
scattered over the southern half of the KD (T5N R6W Sees. 4. 5, 25, 33; T5N R7W
Sec. 6;T5N R8W Sec. r,T6N R8W Sees. 26, 28, 29, 32. 35, 36). It grows, often
in large number (> 1000), in full sun almost exclusively in thin soils in slight
depressions on flatrock pavements (MacRoberts & MacRoberts 1993c; Thieret 1959).
It also is recorded from Caddo, Red River, Sabine, and Winn panshes (Nai.ural
Heritage 1994).

Tetragonotheca ludoviciana (Torrey & A. Gray) A. Gray ex Hall. (MacRoberts &
MacRoberts 1988 [VDB], 1691 [SFRP], 2371 [LSU]). There are six known
locations for Louisiana square-head, a WGCP endemic, on the KD (Natchitoches
Pansh, T6N R7W Sees. 1, 2. 5, 9, 17), and one on the VD (Vernon Parish, TIN
R7W Sec. 22) (Hart & Lester 1993). It grows in upland xeric sandhill woodlands
(Grace 1993; Gandhi & Thomas 1989; Turner & Dawson 1980). It has also been
reported from Allen, Caddo, and Sabine panshes (Natural Heritage 1994 ).

Tradescantia reverchonii Bush. (MacRoberts & MacRoberts 1565 [VDB], 2276, 2407
[SFRP]). This species is on the Louisiana Natural Hentage watch list. It is found
almost exclusively in upland and stream terrace xenc sandhill woodlands of western
Louisiana and eastern Texas south to the Rio Grande Valley. It is common in all
sandylands in both the KD and WD (Natchitoches and Winn panshes). It appears to
especially like disturbed areas (MacRoberts 1980). It has been reported from Caddo,
Beauregard, Bienville, Sabine, and Vernon parishes (MacRoberts 1989).

307 PHYTOLOGIA volume 78<3):29 1-3 13 April 1995

Triphora trianthophora (Sw.) Rydb. {MacRoberts <Sc MacRoberis 2509 [SFRP]).
Three-birds orchid, a widespread eastern species, has been found only twice on the
KNF, KD (Natchitoches Parish, T7N R7W Sec. 20) in "damp, shady woods" in
September 1977 (Mathies el al. 1983; Holmes & Mathies 1980) and ED (Rapides
Parish, T2N R3W Sec. 11) in hardwood slope forest m 1994. It has also been
recorded for Caddo and West Feliciana parishes (Natural Hentage 1994).

UtriculariajunceaWahl (MacRoberts & MacRoberis 825, 1179 [VDB], 872 [SFRP],
1442 [LSU]). This bladderwort is on the Louisiana Natural Hentage watch list. It is
locally common in bogs on the KD and VD (Natchitoches and Vernon parishes)
(MacRoberts & MacRoberts 1988). It has also been reported from Beauregard, St.
Tammany, and Washington parishes (MacRoberts 1989).

Utricularia purpurea Walt F*urple bladderwort is on the Louisiana Natural Heritage
watch list. It has been reported from only one bog on the VD (Vernon Parish, TIN
R6W Sec. 22), but there is no ioiown voucher specimen (Parker 1990). It has been
reported from Evangeline, Cameron, Tangipahoa, and St. Tammany panshes
(MacRoberts 1989).

Uvularia sessilifolia L. (Mclnnis 4003, s.n. [personal collection]). Sessile-leaved
bell wort, a northeastern species, known from the northern tier of Louisiana panshes,
has been found at two locations on the VD (TIN R7W Sees. 13, 33) (Hart & Lester
1993) and one location on the WD (Natchitoches Parish, T13N R6W Sec. 8). It
occurs in wooded seeps and bayhead swamps. It is also recorded from Caddo,
Claiborne, Lincoln, Morehouse, Union, and Webster panshes (Natural Hentage
1994).

XarUhorhiza simplicissima Marsh. {Allen 14892; Thomas & Allen 98050 [NLU]).
Yellowroot, a southeastern species that is rare in the WGCP, was first found in
Louisiana on the VD (Vernon Parish) in 1986 and has subsequently been found at
another location about three miles away (TIN R7W Sec. 34 and T IS R8W Sec. 1)
(Allen el al. 1987; Buras 1993; Hart & Lester 1993). It favors shaded npanan forest
sites. These are the only locations reported in Louisiana (Natural Hentage 1994).

Xyris drummondii Malme. {MacRoberts «fe MacRoberts 868, 885 [VDB], 1158,
1775A [SFRP], 841, 1775B [NLU], 1781, 1841 [LSU]). Dmmmond's yellow-eyed
grass, a coastal plain species, was first reported for the WGCP in the mid-1970s from
a bog in the KD (Natchitoches Parish, Krai 1983). Surveys of bogs on the KNF
since the late 1980's have located it in about 75 bogs on the KD and VD (Natchitoches
and Vernon parishes, Hart & Lester 1993, MacRoberts & MacRoberts 1988, 1990a,
1991, 1992, 1993b) It is so widespread in these two distncts that we will not give
loGitions. It is also known from many other bogs in western Louisiana and eastern
Texas (Bndges & Orzell 1989a; Orzell 1990). It is also found in Beauregard Pansh
(Natural Heritage 1994).

Xyris scabrifolia Harper. {MacRoberts & MacRoberis 1491, 1543 [VDB], 1448,
1793 [SP-RP], 1754, 1782 [NLU]). Rough-leaved yellow -eyed grass, an uncommon
coastal plain species, was first reported for the WGCP in the early 1980s. It has been
found in sixteen bogs scattered widely over both the KD (Natchitoches Parish, T5N
R8W Sec. 3; T6N R7W Sees. 12, 21, 22; T6N R8W Sec. 35) and VD (Vernon

MacRoberts & MacRoberts: Rare plants of Kjsatchie National Forest 308

Pansh, TIS RSWSecs. 12. 14; TIN R5W Sec. 28; TIN R6W Sees. 14, 20, 21, 23,
25; TIN R7W Sec. 29; TIN R8W Sees. 1, 34) (MacRoberts & MacRoberts 1992,
1993b; Bndges & Orzell 1989a; Hart & Lester 1993). It also occurs in Beauregard
Parish (Natural Hentage 1994).

Zigadenus densus (Desr.) Femald. (MacRoberts & MacRoberts 1646 [VDB], 1966
[SFRP], 1646 [LSU]; Hampsten s.n. [SFRP]). While black snakeroot and other
Zigadenus species have been reported from Louisiana for a long ume, there has been
confusion over taxonomic status and distnbution (Thieret 1972). Zigadenus densus is
a coastal plain species. Dunng surveys in 1992 and 1993, it was found in fifteen VD
bogs (Vernon Pansh. TIN R5W Sees. 28, 30, 31, 32; TIN R6W Sees. 21, 23; TIN
R7W Sees. 24, 25, 27; TIN R8W Sees. 15, 22) (MacRoberts & MacRoberts 1993b;
Hart & Lester 1993). It is also reported from St. Tammany and Washington panshes
(Natural Hentage 1994).

Zomia bracteata J.F. Gmel. (MacRoberts & MacRoberts 1897 [VDB], 1807. 2413
[SFRP]). Vipenna, a southeastern species, occurs in a stream terrace sandhill
woodland at one inholding on the WD (Winn Parish, T12N R5W Sees. 28. 29)
(Lasseigne 1973). It has also been reported for Caddo, Natchitoches, Vernon, and
Tangipahoa panshes (Natural Hentage 1994).

EXCLUDED SPECIES

Two species that have been on the KNF rare species list have not been venfied.
One is Xyris stricta Chapman reported from several localities on the VD (Vernon
Parish, and from southern Natchitoches Pansh) (Hart & Lester 1993). The voucher
specimens for these reports turn out to be other Xyris species (Linda Watson and
Bruce Hoagland, pers. comm.). The other is Asclepias hirtella (Pennell) Woodson
reported for the KD (Natchitoches Pansh). After rep)eated searches, the voucher
specimen for this has been located and it turns out to be A. longifolia Michx. (S.
Lynch, pers. comm.).

DISCUSSION

The Kisatchie National Forest comprises about two percent of the Louisiana land
surface. The 1994 Louisiana Natural Heritage rare species and watch lists contain 407
species. Eighty, or 20%, of these have been found on the KNF. The Natural
Heritage has 2071 rare plant speaes site records for the entire state (Julia Larke, pers.
comm.). Add to this many records from the KNF not yet in their data base, and the
figure probably approaches 2500. There are about 570 rare species site records for the
KNF. The ICNF therefore accounts for about 23 percent of the Louisiana total.
Relatively speaking, therefore, the KNF is either very neh in rare species sites or has
been exceptionally well surveyed. The latter, of course, is the case largely because it
is open to investigation, because it has relatively vast tracts of undisturbed habitat, and
because several community types within it have been the subject of special surveys.

Companng these records with what was known about these species in the WGCP
ten years ago makes it obvious that while much work has been done, much remains to
be done in documenting the fiora of the WGCP (Bndges & Oizell 1989a; 63). For

309 PHYTOLOGIA volume 78(3): 29 1-3 13 Apnl 1995

example until two decades ago, Xyris drummondii was not known to occur in the
WGCP, and for years after its discover>' it was known from only one bog. Recent
surveys of bogs in west Louisiana and east Texas have found it in over 100 locations
(Bndges & Orzell 1989a; Orzell 1990; MacRoberts & MacRoberts 1993b; Han &
Lester 1993), its highest known concentration. Rudbeckia scabrifoUa was not
recognized as a separate species until 1986 (Brown 1986). Due to intensive bog
surveys, it too has now been documented from over 100 WGCP bogs and baygalls
(Bridges & Orzell 1989a; Orzell 1990; MacRoberts & MacRoberts 1993b; Han &
Lester 1993; Parker 1990).

The same applies to community types. A concerted effort is currently under way
to negotiate a taxonomy of community types for the WGCP (Smith 1988, in prep.;
Diamond et al. 1987; Bndges & Oraell 1989b; Orzell 1990; Martin & Smith 1991,
1993), but as more research is undertaken, it is clear that these taxonomies are
preliminary.

At the same time, we are only beginning to map the extent and location of different
communities. Surveys of major community types, as well as those of inclusion
communities, are beginning to uncover the distnbution, extent, and condition of these
rare, localized, and speci^ized habitats where rare plants often occur in surprising
abundance. Unfortunately, progress is slow since, in the entire WGCP, only a
handful of botanists and ecologists are working.

ACKNOWLEDGMENTS

The continuing cooperation and assistance of the staff of the Kisatchie National
Forest was instrumental in making this work possible. Especially to be thanked are
Susan Carr, Ken Dancak. Karen Belanger, Tom Fair, and John Baswell. Part of our
research was sponsored by volunteer and cost-share agreements. Part of our survey
work on the Vernon District was conducted while on conu^act with The Nature
Conservancy. We wish to thank the Louisiana Natural Heritage Program, Louisiana
Department of Wildlife and Fishenes, and The Nature Conservancy, for sharing with
us their information about rare WGCP plants. In 1994 some of the data were gathered
while the authors were on contract with the Forest Service ("Sensitive Plant Survey",
#RFQ R8-6-94-417). We wish to thank the Forest Service for allowing us to publish
that data. Robert Krai, Will McDearman, Steve Orzell, Steve Lynch, and Julia Larke
vetted a number of difficult species. Julia Larke, Latimore Smith, and Nelwyn
Mclnnis supplied much information concerning rare species and Louisiana ecology.
D.T. MacRoberts, Julia Larke, Latimore Smith, Phil Hyatt, and Rhonda Stewart made
useful comments on the manuscnpt. Many others aided in vanous ways, notably Dale
Thomas, Dcm Rhodes, Steve Lynch, Rhonda Stewart, and Charles Allen, Robert
Krai, Tom Wendt, Steve Lynch, Linda Watson, D.T. MacRoberts, David Moore, and
Bnxe Hoagland were especially helpful in clearing up some problematic items. Since
people, not institutions, fourxi and recorded these plant locations, thanks are due the
individuals who searched for these plants and who were sedulous enough to make
collections and/or see that their findings get into a centralized data base - either of the
Louisiana Natural Hentage FYogram or the Kisatchie National Forest - or into a
herbarium or into pnnL

MacRoberts & MacRoberts: Rare plants of Kisatchie National Forest 310

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Phylologia (Apnl 1995) 78(4)314

BOOKS RECEIVED

Aquatic Weeds, The Ecology and Management of Nuisance Aquatic Vegetation.
Arnold H. Pieterse & Kevin J. Murphy (eds.). Oxford Science Pubhcations,
Oxford University Press, 200 Madison Avenue, New York, New York 10016.
1994. xviii. 593 pp. $49.95 ISBN 0-19-854840-0 (paper).

This book was designed as a textbook on aquatic weeds. Authors (30)
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