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Table of Contents
SYSTEMATICS
them Appalachians
Viola calcicola (Violaceae), a new endemic violet from the Guadalupe Mountains
of New Mexico and Texas
Ross A. McCauley and Harvey E. Ballard, Jr.
Two new species of Eriastrum (Polemoniaceae) from California
Carex cyrtostachya (Cyperaceae), a new species of sedge endemic to the
Sierra Nevada of California
Peter F. Zika, Lawrence P. Janeway, Barbara L. Wilson, and Lowell Ahart
A re-evaluation of Carex specuicola and the Carex parryana complex (Cyperaceae)
A. A. Reznicek and D.F Murray i i r A I
New combinations in Carex kelloggii (Cyperaceae) MISSOURI BOT A
Barbara L. Wilson, Richard E. Brainerd, and Nick Otting
A revision of Cordia section Gerascanthus (Boraginales: Cordiaceae) OEC ^ ^ 2013
James S. Miller
Novelties in Serpocaulon (Polypodiaceae) GARDEN LIBRARY
Pedro B. Schwartsburd and Alan R. Smith
25
37
85
Two new species from the Brevanthemm clade of Solanum (Solanaceae) from eastern Brazil
Leandro L. Giacomin, Lynn Bohs, and JoAo R. Stehmann 95
New species of Aphelandra (Acanthaceae) from Peru and Ecuador
Dieter C . Wasshausen 1 09
Estudios en las Apocynaceae Neotropicales XLlIl; s
(Apocynaceae; Apocynoideae) en Colombia
J. Francisco Morales
-„mbuseae: Arthrostylidiinae): two new.
Emmet J.JuDziEwicz, EricJ. Geisthardt, Ch
1 (Poaceae: Bambusoideae:
vered species from Peru
Waas, and Sol Sepsenwol
i northern Andean species of Aulonemia (Poaceae: Bambusoideae:
wiuuuseae: Arthrostylidiinae) with veirucose culms
Emmet J.JuDziEwicz, Ximena Londono, and Lynn G. Clark
Dos nuevas especies de Guadua para el Peru (Poaceae: Bambusoideae: Bambi
Ximena Londono
Calathea croatii (Marantaceae), a new endemic species from Panama
Two new endemic species of Calathea (Marantaceae) from Ecuador
Helen Kennedy
El genero Polypogon (Poaceae: Agrostidinae) en Chile
Victor L. Finot, I fsi if Contreras, Wilson Ulloa, Alicia Marticorena,
Carlck M. Baeza & Eduardo Ruiz
145
155
169
203
(Asteraceae; Gnaphalieae) from South America
Claudia Monti, Nestor D. BayOn, Daniel A. Giuuano, and Susana E. Freire
Diferenciacion de especies Argentinas de Clinopodium (Lamiaceae; Nepetoideae)
a traves de caracteres morfologicos y anatomicos de sus frutos
Meuna Scandauaris and Gloria E. Barboza
Systematics, phylogeny, and biogeography of Chaetolepis (Melastomataceae)
Daniel Grimm and Frank Almeda
Miconia phrynosomaderma (Melastomataceae: Miconieae), a new species
from the Massif du Nord, Haiti, and sixteen new names and combinations
Lucas C. Majure and Walter S. Judd
Crataegus tenuior (Rosaceae) — an intriguing new species from the Okanagan
of British Columbia and Washington and a new variety of C. okanaganensis
J.B. Phipps
Identification, distribution, and habitat of Coreopsis section Eublepharis
(Asteraceae) and description of a new species
Bruce A. Sorrie, Richard J. LeBlond, and Alan S. Weakley
ArceuthobiumgilliiandA. nigrum (Viscaceae) revisited: distribution, morphology,
and rDNA-ITS analysis
Shawn C. Kenaley and Robert L. Mathiasen
The genus Platanthera (Orchidaceae) in Mississippi
Michael Wayne Morris
A new species of Ceropegia (Apocynaceae: Asclepiadoideae) from India with notes
on rare and threatened Ceropegia in Nilgiris of Western Ghats
P. Sujanapal, pm. Salim, N. Anil Kumar, and N. Sasidharan
A note on ‘ Eritrichum Schrad. 1819” and Eritricbium Schrad. ex Gaudin 1828 (Boraginaceae)
Kanchi N. Gandhi and David F. Murray
Notes on Eritricbium (Boraginaceae) in North America
David EMutiray
A review of the neotypification of Astrophytum capricome var. crassispinnm (H. Moller)
Okumura (Cactaceae)
Richard R. Montanucci and Heinz Hoock
paleobotany
A fossil flower of the genus Protinm (Burseraceae) in Mid-Tertiary amber
from the Dominican Republic
Kenton L. Chambers and George O. Poinar, Jr.
Hippocratea vohibUis (Celastraceae) in Cotui copal from the Dominican Republic
George O. Poinar, Jr., Kenton L. Chambers, and Alex E. Brown
CHROMOSOME STUDIES
- _ * (Poaceae: Chloridoideae) from north central Mexico
L Rosales Carrhio, Y. Herrera Arioeta, M.S. GonzAiez Euzondo, N. Almaraz Abarca,
L Mayek PErez, and A.L. Latorga Vanzela
217
265
275
299
311
323
341
347
351
355
367
375
381
DEVELOPMENT AND STRUCTURE
A survey of seed coat morphology in Oxytropis, sects. Arctobia, Baicalia, Glaeocephala,
Mesogaea, and Orobia (Fabaceae) from Alaska
Zachary J. Meyers, Stefanie M. Ickert-Bond, and Rose LaMesjerant
BOTANICAL HISTORY
The forgotten first checklist of the Iowa flora; John Henry Rauch’s 1851 “Report”
to the State Medical and Chirurgical Society
Thomas G. Lammers
ETHNOBOTANY
A preliminary report of mescaline concentrations in small regrowth crowns vs. mature crowns
of Lophophora williamsii (Cactaceae): cultural, economic, and conservation implications
M.Abul Kaiam, Molly T. Klein, Diana Hulsey, Keeper Trout, Paul Daley, and Martin Terry
FLORISTICS, ECOLOGY, AND CONSERVATION
Potential distribution of three native and one introduced grass species in semiarid highlands
of Mexico using GIS techniques
Armando Cortes Ortiz, Yolanda Herrera Arrieta, Jesus Herrera Corral, and
Daniel Hernandez Velazquez
A baseline vascular plant survey for Ocmulgee National Monument, Bibb County, Macon, Georgia
Wendy B. Zomlefer, David E. Giannasi, John B. Nelson, and L.L. Gaddy
Flora of calcareous upland glades in Gadsden and Jackson counties, Florida
A.F. Johnson, WW. Baker, L.C. Anderson, and A.K. Gholson, Jr.
History, dispersal, and distribution of Buddleja davidii (Scrophulariaceae) in Kentucky
Ralph L. Thompson and J. Richard Abbott
Astrolepis sinuata (Pteridaceae) new to the flora of Louisiana
Jeff McMiluan, Charles M. Allen, and Selena Dawn Allen
Notes on the identification and distribution of the species of the genus Galium (Rubiaceae)
Charles M. Allen
Synopsis of the genus Stylisma (Convolvulaceae) in Louisiana
Charles M. Allen
Confirmation of kariba-weed, Sahinia molesta (Salviniaceae) in the Calcasieu River
453
475
495
515
Ray Neyland and Jennifer Bushnell
A quantitative study of the vegetation surrounding a
(Ranunculaceae) population at Fort Polk in west central Louisiana
Charles Allen, Rachel Erwin, Jeff McMiluan, and Joe McMillian
Vascular flora and plant communities of Alleghany County, North Carolina
Derick B. Poindexter
517
519
529
Vascular plants of the Canyonlands Unit of the Big Thicket National Preserve, Tyler County, Texas
Kelly C. Hah f and Stephan L. Hatch 575
Heocharis microformis (Cyperaceae): rediscovered in North America from the Edwards Plateau
and Trans-Pecos regions of Texas
Robert J. O’Kennon and Kimberly Norton Taylor
587
vascular tlora of saline lakes in the Southern High Plains of Texas and eas
Davtd J. Rosen, Amber D. Caskey, Warren C. Conway, and David A. Haukos
Ecology and distribution of the n
exas endemic Dalea reverchonu (Fabaces
William F. “Bill” Mahler (1930-2013)
Book Reviews and Notices 8,36 52 54 84 94 108 144
346, 350. 3H 366, 374, 380, 434, 452,’ 474, 506, 308, 514
i 586,594
i4, 160, 168, 216, 264, 298, 340,
I Bot. Res. Inst. Texas 7(1), 2013
J.F. Morales, sp. nov.— 123
dra aucayacuensis Wasshausen, sp. nov-113
.K.«7.«..dradavidsoniiWasshausen sd nov— in
Aphelandra ^pinosae Wassshausen, sp. nov-li9
ASani^ plo^'
Aujonemia notata McClure ex J^r^GTlaiV/^^’n
A^onemiar«bralig„Iaujudz.&G;ist^^^^^^^
Calathea fredu H. Kenn sp. nov— 164
Calathea neillii H. Kenn. sp. nov— 161
^rex ^rtostachya Janeway & Zika, sp. nov— 25
&K.xbo^mgrc„io™n.Reznicek&D.F Murray nom r,« .
Carex utahensis Reznicek & D.F. Murray, sp. nov^^’^ Brainerd, comb, nov— 53
Chaetokpis phelpsiae var. cliiiiiant«i«i«
Ckmaiis vinacea Floden, sp. nov.— 2 ^ Almeda, stat. nov— 258
a^ulara LeBlond, Sorrie & Weakley, sp r».v_3oi
r. longispinaJ B. Phipps var nov—
s tenmorJ.B. Phipps, sp. nov. — ^281
" Gowen, sp. nov. — ^21
nse D. Gowen, sp. nov— 23
Guadua lynnclarkiae Londono, sp. nov— 146
Guadua takahashiae Londono, sp. nov— 149
Miconia aigentimuricata Majure & Judd, nom. nov— 268
Miconia asperifolia (Naudin) Majure &Judd, comb nov—
Miconia cubacinerea Majure &Judd, nom. nov— 268
Miconia cubana (Alain) Majure & Judd, comb nov— 268
Miconia granulata (Urb.) Majure &Judd, comb, nov— 268
Miconia hybophylla (Urb.) Majure &Judd, comb nov— 268
Miconia inaeqnipetiolata Majure & Judd, nom. nov— 268
Miconia jashaferi Majure & Judd, nom. nov— 268
Miconia kaiikmgu Majure & Judd, nom. nov — 268
Miconia kmgiana (Cogn.) Majure & Judd, comb, nov— 269
Miconia sect. Lima Majure & Judd, sect. nov. — ^266
595
603
611
613
A NEW LEATHERFLOWER (CLEMATIS: RANUNCULACEAE)
FROM THE SOUTHERN APPALACHIANS
Aaron Floden
TENN Herbarium
Department of Ecology and Evolutionary Biology
University of Tennessee
Knoxville, Tennessee 37996, U.S.A
INTRODUCTION
Several populations of a Clematis L. (Ranunculaceae) with affinities
shale talus slopes in the Ocoee River Gorge in Polk County, southeast Tennessee; an unusually dry habitat for
IS seen in C. crispa. Plants observed during anthesis in early spring
nee to the western North American C. hirsutissima Pursh. These
and single-flowered with pinkish-purple, sericeous sepals. The three to five pairs of opposite leaves were
highly divided and densely sericeous on both surfaces. Additional observations later in the season showed that
the plant can continue to grow from axillary buds at the base of the terminal inflorescence. When edaphic
conditions, i.e., abundant soil moisture, are present throughout the season the plant continues growing where-
up>on the habit becomes vine-like.
Examination of herbarium materials at TENN shows that Clematis vinacea has long been confused with
the Appalachian forms of C vioma L. which are glabrous in comparison to the pubescent forms encountered
in the Interior Low Plateau. Despite the similarities of the vestiture of C. vinacea to some C. vioma forms there
are differences in the narrower leaflets and lobes, the sericeous vestiture on the leaves, stem, and sepals, and
the distinctly abaxially and adaxially pinkish-purple colored sepals. It is Ukely that the distribution of this spe-
cies, which is outside the range of C. crispa, has partly contributed to its inclusion within the sympatric C
vioma. Nonetheless, the phenology of C. vinacea begins in early spring (see type collection date in late March)
and flowers throughout the growing season whereas C. vioma begins flowering in late May where the two are
sympatric. Both C. vinacea and C. vioma share plumose achene tails, however, the achene tails are gold plu-
mose in C. vinacea whereas they are typically brownish to whitish in C. ^
of this new species with C. vioma, the presence of tl
both surfaces, and growth habit suggests a close phenetic relationship to C. crispa. Despite their shared mor-
phology, the plumose achene tails, leaf indument that is usually lacking in C. crispa, and the only slightly
?. vinacea differ strongly from C. crispa.
the thick, leathery connivent sepals
which are urceolate and usually pendent (Pringle 1997). A common vernacular name for the group is “leather-
flower.” Clematis (Ranunculaceae) is a large genus comprising approximately 350 species (Wang 2005). Infra-
generic classifications have been based partly on floral morphology (Tamura 1967, 1995; Keener & Dennis
1982), floral and leaf morphology and plant growth habit (Erickson 1943), and a combination of the above and
eophyll orientation (Yang & Moore 1999; Wang 2005). Wang’s (2005) recent subclassification recognizes sub-
genera, subsections and series, of which our southeastern leather-flowers would fall under Clematis subg.
Vtoma A. Gray, sect. Vioma (Reichb.) Prantl, subsect. Crispae Prantl which is subdivided further into five se-
rial divisions. Pringle (1997) includes 25 species in Clematis subg. Vioma with 18 of these present in North
America. 1 follow this broader subclassification. As part of a revision of the North American Clematis subg.
Vioma (Estes, Floden, & Witsell, in prep.), examination of herbarium specimens and cultivation of nearly all
eastern North American Clematis subg. Vioma have provided data supporting the recognition of C. vinacea,
which appears to be endemic to a small region of the southern Appalachians.
horizontal ihizomes, some
1 , 1 1 n ^ midstem, spreading, to 1-3 dm long, longer than wide- petioles slender sub-
^al to bweroK^tleaneB, 1-6 cm long, sericeous pnbeseem;lealfeLynLlrtcal or
a^mmate, recurved, margins white sericeous;stamens linear filament u ’
oblong, 4-5 mm long, connectives Dubesrentrrtcaata j anthers oblong or narrowly
long, apex papillose and curved, plumose. ^yond anther; stigmas 10-14 mm
trichomes to 5 mm orbicular, brotvn, shortly sericeous; 4-6 mm, tails 3-5 cm!
louring; March-Sep.ember.fruitlngJune-Oc.ober.
Fefac. 6- B. Adam 11369 (TENN). ^ Gorge; W. Manek 88 (TENN); 17 Apr 1949. A.J. Sharp, C.J.
Living material of Clematis vinacea was first observed by the author in early June 2009 that was in anthesis
while also bearing nearly mature gold-plumose achenes. Notable features were the dense sericeous pubescence
of the pinkish-purple sepals, stem and leaves, and the leaves with more numerous and narrower leaflets than
C. \ioma (Fig. 1). Although annotations have consistently placed C. vinacea within the concept of C. vioma or
C. crispa, specimens of C. vinacea at TENN are easily segregated from the former by their terminal flowers
borne on ebracteate pedicels and narrower leaves and from the latter by the sericeous indument of the leaves
and sepals and plumose achene tails. Furthermore, the habit of C. vinacea due to continued growth in response
to sufficient moisture enables the plants to become vining later in the growing season whereas collections
early in the season appear distinctly erect with a single terminal flower. A number of collections state the plant
as “rather shrub-like” or “erect” which has been observed by the author both in and ex situ. Observations of C.
vioma in the field show that its inflorescences are strictly axillary and begin in late May on distinctly vining
plants. In contrast, early flowering does occur in C. crispa, but the plants are distinctly scandent and glabrous.
The late season collections of C. vinacea, despite their terminal ebracteate inflorescences, have contributed to
Based on similarities in morphology and phenology, the phylogenetic relationship of Clematis vinacea
seems to be with other terminally flowered Clematis of subg. Vioma; those from the American southwest and
southeast. On a phytogeographic basis a close comparison can be made with C. crispa rather than the morpho-
logically comparable western C. hirsutissima. It shares with C. crispa. C. baldwinii Torr. & A. Gray, and C. socia-
lis Krai several distinctive morphological and phenological characters: 1) ebracteate primary peduncles borne
3) early-spring flowering. In morphology, C. vinacea appears closest to its eastern counterpart C. crispa, but the
two are easily separated on account of their sepal morphology (narrowly or not crispate vs. broadly crispate)
and their achenes (plumose vs. non-plumose), respectively. Comparison of cultivated plants of C. hirsutissima
also shows similarities: a strongly clump forming habit from a short horizontal rhizome; finely divided lower
leaves; and terminal inflorescences, though the indument observed in C. hirsutissima is denser than that seen
in C. vinacea to nearly hoary in some parts of its distribution (Pringle 1997).
Observations of several genets of both Clematis vinacea and C. crispa cultivated in a common garden re-
veal that both exhibit similar growth patterns. Both display remontant behavior in response to ideal edaphic
conditions where the axillary buds subtending each terminal, single-flowered inflorescence enable continuous
'■*“*6 giuwiiig rupuii
le-like plants from early spring through September during 2
, 2010, and 2012. Field observa
revealed that during drought conditions C. vinacea remains a single stemmed, once flowering perennial. In
cultivation C. socialis and the western C. hirsutissima have not shown any remontant tendency, but have con-
tinued limited vegetative growth after their spring flowering. Both are erect perennials with short to long
branching rhizomes (Krai 1982). Rhizomes have been observed in several populations of C. vinacea where
single clones can have up to 12 or more stems separated by several centimeters each (Fig. 1), whereas rhizomes
are not present on C. crispa.
^ Perianth morphology of Clematis subg. Vioma differs little between taxa; all are 4-sepaled, urceolate, and
within a s^ies, although not usually within a population. Clematis crispa has broadly crispatllepal margin!
sibsectional status within the broader subg. Viontu (Grey-Wilson
^). Nonetheless, ^ml other taxa not included in this subsectional division are notable for their crispate
^1 matgttn as well: C. WdwitUi; C. suciulis, C. hirsutissima; and some forms of C. pitrheri Torr. & A Ly
^ the southwestern pan oflts range. Clematis vinacea differs front its phytogeog^phic neighbors by no'
havmptom,nent enspates^lmargitn.Punhetmore, the presence of pl„^^
e) achene tails on C. crispa. Plumose achene tails are shared with C. bald-
presence of the plumose achenes on C.
p r _ luse it also exhibits
from the typically
^.teepcirnf^composedofsiliceousphylhtesLpes^^^
m Tennessee are limited to the Walden-- ’ ' ^mcuitto
i between the Smoky h
s narrow band along the Ocoee River 0^100^0 is e:
All known populations
and west (Hardeman 1
n abundance of twailable habitat. It is known f.o„, .wo
rt '“outh of the gorge along Spring Creek; and further up-
1 ^rn^ecoHectionfromsoutheastemMurrayCo.,Georgia
reportedoccurrencesofCcrist«,andixitentM“i;c^^^^^^
^d C. crispa occur in distinctly different habi-
Floden, A new Clematis from the southern Appalachians
Journal of the Botanical Research Institute of Texas 7(1)
vinacea occurs. Most of the shrubby layer was killed to the ground and many herbaceous plants were killed
outright. Surprisingly, C. vinacea responded quickly to mid-summer rains and was in full flower by mid-Au-
gust as the predominant plant in several burned areas (Fig. 2). Excavation of several plants showed the root
crown was positioned 15-20 cm below the soil surface. The deeply positioned root crown is likely an adapta-
tion to the subxeric habitat that also provides protection from the occasional fire.
SUGGESTIONS FOR COLLECTING CLEMATIS SPECIMENS
A lack of material or too much plant material folded over itself on herbarium specimens can obscure the dis-
tinct characters of a collection of Clematis subg. Vioma. Morphology can also be misinterpreted when material
for preservation is taken from the proximal to middle part of the stem rather than a terminal section. Therefore,
collectors should note from which part of the plant the material has originated if it is not obvious, especially the
vining taxa, and also seek to press the whole stem in sections. The position of the hracts on the peduncle should
not be obscured by the leaves. Furthermore, the perianth color of Clematis herbarium material is usually
masked upon desiccation by turning brown (Keener 1975; Dennis 1976). Variation in color of the sepals can
differ between early season flowers and those produced later in the season. 1 have observed this in situ and ex
situ which has also been confirmed by D. Estes (pers. comm.). Noting the color, or providing a photograph of
the sepals, despite their variability, is useful in some groups, e.g. Glaucophylla-group, and Vioma-group.
ACKNOWLEDGMENTS
1 thank L.D. Estes for conversation regarding the genus while working towards a revision of subg. Viorna (of
North America) and the Dennis-Breedlove fund of the University of Tennessee for monetary support for field
work. I also thank the curators of GA and MO for facilitating visits to study Clematis. 1 also thank E. Schilling
for comments on the manuscript. Last I thank an anonymous reviewer and L.D. Estes for critical reviews of the
Tennl^^^' ^ Subsection Wornoe. Ph.D. d
OtNNis, W.M. 1976b. Chromosome morpi
1135-1139.
Ewckson. R.0. 1943. Taxonomy of Clematis
Grey-Wxson. C. 2000. Clematis the genus. 1
Hardeman, W.D. 1966. Geologic map of Tennessee. Tennes:
1:250,000.
xZr'rT; of ^oothern Appalachian endemics. Castanea 1 2:100-11 2
« ,9*7 A b»>,s«™,c sb,dy Of subsooboo ,«a„„„co,aceao). f. Elish, Mlfcholl Sb.
Keener, C.S. 1975. Studies
Keener, C. ano W.M. Dennis. 1 982. The :
north of Mexico. Taxon 3137-44.
Kral,R.1982.
Kral, R.1 983. A report
Isoetaceae through Euphorbiaceae and Voli
eds. 1997. Flora of h
ornae (Ranunculaceae). Canad. J. Bot. 54:
Card. 30:1 -62, platel.
of Geology Map, Nashville, TN, 4 sheets, scale
United States. III. Clematis L. Sida 6:33-47.
North America
Tamura, M. 1 987. A classification of
Tamuha, M. 1995. Clematis.
Humboldt Pp. 368-387.
/. 16:21-43.
a (4). Berlin: Ducker uni
Floden, A new Clematis from the southern Appalachians
Wang, W.T. 2005. A new system of classification of the genus Clematis (Ranunculaceae). Acta Phytotax. Sin. 43:431-488.
Wood, CE., Jr. 1970. Some floristic relationships between the southern Appalachians and western North America. In:
P.C. Holt, ed.The distributional history of the biota of the southern Appalachians. Part II: Flora. Virginia Polytechnic
Institute and State University, Blacksburg. Pp. 331-404.
Yang, T.Y. and D.M. Moore. 1999. A revision of the viorna group of species (Section Viorna sensu PrantI) in the genus
Clematis (Ranunculaceae). Syst. Geogr. PI. 68:281-303.
BOOK REVIEW
Michael Wojtech. 2011. Bark: A Field Guide to Trees of the Northeast. (ISBN; 978-1-58465-852-8, hbk).
University Press of New England, 1 Court Street, Lebanon, New Hampshire 03766, U.S.A. (Orders:
www.upne.com, 1-800-421-1561, 1-603-448-9429 fax). $25.95, 280 pp., 65 maps, 434 illus. (283 color),
keys,5Vi"x8y4".
Bark: A Field Guide to Trees of the Northeast is divided into six chapters: instructions on how to use this particu-
lar guide, bark structure, bark types, secondary identification keys, bark ecology, and species. This book con-
sists of numerous color images, line drawings, keys, and maps that provide the reader with further information
about the trees that one would encounter in the forests of the Northeastern U.S.A.
The opening chapter provides information on how a potential reader would begin to use this field guide.
The two chapters that follow contain detailed information regarding the structure and types of bark that can
n have a variety of appearances: smooth, peeling, curling, cracking, scales, plates,
er determine which species is in front of them. The author includes
- “S 1-2 images ofactual trees with these bark types. Also included is
a U.S.A. quarter in each photograph to establish a standard scale for each feature shown. Wojtech also included
a chapter on terk ecology to help the reader further understand the diversity and function of bark,
help ^u h^T T h ^ primary key is printed on the inside cover. This key will
f secondary key section to designate the age of the trTe:"yo Jn J(n mlium
(M), and old (O). This IS importantsince bark appearance can change as the tree ages.
that l^'ih T
thaUist the dominant common name, other possible common names, family name, and Latin binomid. He
parL^Xr account for the
Port Worth, Texas, U.S.A.
VIOLA CALCICOLA (VIOLACEAE), A NEW ENDEMIC VIOLET FROM THE
GUADALUPE MOUNTAINS OF NEW MEXICO AND TEXAS
Ross A. McCauley Harvey E. Ballard, Jr.
Fort Lewis College
Durango, Colorado 81 301, U.S./
mccauley_r@fortlewis.edu
Over the last half-century, three taxonomic treatments of North American (primarily U.S. and Canadian)
Viola (Violaceae) have been proposed by specialists (Russell 1965; McKinney 1992; Gil-ad 1997). These differ
substantially in the circumscription, rank, nomenclature and general evolutionary status of several taxa. In
these and earlier treatments, specialists have noted two broad groups of taxa with notched, lobed, or divided
leaves; homophyllous violets in which aU leaf blades are lobed or divided throughout the growing season, and
heterophyllous violets which produce undivided blades in the earliest and latest months of the growing season
and notched, lobed or dissected leaves during the height of anthesis and into chasmogamous fruit. All three
most recent treatments recognize some heterophyllous-leaved taxa, namely V. egglestonii Pollard, V septemloba
Leconte and V triloba Schwein., at some taxonomic rank. Russell maintained the hrst three as well as V. escu-
knta Elliott, V. lovdliana Brainerd and V viarum Pollard, as distinct species. McKinney synonymized the latter
three along with V triloba Schein. within a broadly delimited heterophyllous taxon under the name V palmata
L., based on his interpretation that the type of Linnaeus’ violet referred to a heterophyllous rather than a homo-
phyllous violet. Gil-ad used the presence of unique or extrem
cromorphological features in seed coat sculpturing and lateral beard trichomes (Gil-ad 1998), u
species status for V. egglestonii, V. septemloba and V triloba but relegated V. esculenta, V. lovelliana, and V. viarum
to hybrid derivatives based on intermediacy of various features. He provided a complete description and illus-
tration of eastern Mexican V. nuevo-leonensis W. Becker as a heterophyllous-leaved member of the subsection.
Non-specialists have run the gamut in their recognition of these violet taxa in North America. Gleason and
Cronquist (1991) took McKinney’s broad circumscriptions even further by merging nearly all heterophyllous-
leaved and homophyllous-leaved violets into a single taxon as well as synonymizing n '
Journal of the Botanical Research Institute of Texas 7(1)
leaved violets. The forthcoming treatment (co-authored by McKinney and R.J. Little) for the Flora of North
America series will remain largely faithful to McKinney’s original revision. The Biota of North America Pro-
gram website (www.bonap.org, Kartesz 2011) and the USDA Plants Database website (http://plants.usda.gov/
java/) follow Gil-ad’s taxonomy. Recently published regional and state floras, and checklists, follow either ex-
treme or a middle ground borrowing from both.
Specimens of a Viola clearly belonging to subsection Boreali-Americanae have been collected occasionally
throughout the Guadalupe Mountains, but their identity has engendered taxonomic confusion. Spellenberg et
al. (1986) ascribed a series of blue-flowered collections from three canyons in Eddy County, New Mexico by P.J.
Knight to V. viarum Pollard, noting the large (800 km) range extension eastward to Oklahoma, the westem-
U.S. species was proposed in light of other relict species also disjunct in the Guadalupes from their main south-
eastern hardwood forest ranges, although Spellenberg et al. acknowledged that further study might show the
acaulescent blue Guadalupe violet to be unique. Subsequent annotations of specimens by other taxonomists
and violet specialists have included V. lovelliana E. Brainerd, V. missouriensis Greene, and V. palmata L.
(Worthington 2002).
The Guadalupe violet’s unusual short vertical rhizome, sporadically heterophyllous nature, variable near-
white to moderately purple corolla, and restriction to limestone substrate in the region initially attracted the
attention of the authors. Examinations of specimens in herbaria with substantial Guadalupe Mountain angio-
sperm collections (NMC, SRSC, TEXA.L, and UNM), coupled with field studies of populations in New Mexico
and Texas, have revealed that all collected material with whitish to purplish flowers in the Guadalupe Moun-
tain region belongs to a single taxon, one that is distinct from previously described Viola guadalupensis with
yellow flowers (belonging to sect. Chamaemelanium) in the same region. Plants within a given site encompass
a broad range of leaf morphologies, with many plants bearing only unlobed leaves and others producing mid-
season leaves with marginal notches or shallowly to moderately lobed blades (rarely, the largest blades are as
deeply divided as typical V. egglcstonii). Furthermore, plants at the lowest elevations in a given site usually pos-
sess purple corollas, whereas plants at the top of the same canyon often bear pale violet to nearly white ones.
Scanning electron microscopy was conducted in order to document micromorphological characters
found to be taxonomically informative by Gil-ad (1998) in other acaulescent blue violets. Lateral petals and
mature seeds were removed from herbarium specimens of Viola calcicola. One lateral petal was removed from
e«h of two specimens, Knight & Fletcher 3419 (UNM) and Wamock & McVaugh 5418 (SRSC), and cropped to
t^ immediate area around the trichomes before mounting. Two mature seeds from each of two specimens
^ Johnston 16536 (SRSC), were prepared following the methods of Gil-ad
at 1/ mA tor 180 sec. Examinations were performed at The Ohio State University Microscopy and Imaging
measuremems of micromorphological
characteristics ofpetaltnchomes and seeds followed Gil-ad’s t- — ’ ' ’ - ^ ^
Table 1 details distinctions of V. calcicola ii
comparison with the four other h
s terminology and methods (1998).
1 macromorphological and micromorphological
1 by Gil-ad (1997). The new species
ofexTremef^an
nized heterophyllous-leaved orthospecies is far 1^6^11^ new violet and other currently recog-
phylloua-teared species. 1. unusual verrical Aizome. sporadlr^^^^^
McCauley and Ballard, A new endemic Viola from the Guadalupe Mountains
clavate or knob-shaped cucullate lateral petal trichomes set the new violet apart from all other lobed-leaved
violets. Its sparse lateral petal beards, broader cucullate lateral petal trichomes, seeds with paler ground color
and numerous darker patches, and restriction to limestone substrate are highly unusual features in the subsec-
mottled seeds with V. nuevo-leonensis, and its confinement to limestone with V. egglestonii. Following Gil-ad’s
(1997) key to flowering plants, unlobed-leaved specimens of Viola calcicola will key easily to V. cucullata,
whereas plants with lobed or divided blades will key to V triloba (if one ignores the sparse beards on the lat-
eral petals of the new species). In the Gil-ad key to cleistogamous plants, specimens of V. calcicola with unlobed
leaf blades will key easily to V nephrophylla if one ignores seed color (comparatively lighter than V nephro-
phylla in the new species), whereas plants with lobed or divided leaf blades will key generally to V. septemloba
if the seed color is ignored.
Lateral petal trichomes have the distinctly to strongly clavate shape of V cucullata although they are much
smaller than that species and scarcely overlap with V. nuevo-leonensis in width. The technical details of the
cuticular folds generally match those of V. septemloba, but the folds are not as densely packed, and the fine
structure of the folds (Fig. 4C-E) does not match any orthospecies or hybrid characterized by Gil-ad. Primary
and secondary structures on the seed coat of V calcicola share many details with V septemloba and few with
other species. Once again, subtle differences are obvious between the two species, with the ring-like secondary
structures in the former having essentially symmetrical and parallel outlines rather than “tear-drop” or pyri-
form asymmetrical outlines seen in V septemloba. Additionally, the central region within each of the ring-like
structures of V calcicola is generally uniformly depressed throughout rather than gradually diminishing in el-
evation from edge to center as in the structures of V septemloba. Overall, macromorphological traits and prefer-
ence for limestone ally V calcicola most closely to V egglestonii, whereas micromorphological features of petal
trichomes and seeds suggest a close relationship to V. septemloba in many details while also highlighting un-
usual features in the Guadalupe violet.
Short vertical rhizomes in the blue-flowered Guadalupe Mountain violet may be largely genetically fixed
but occasionally environmentally influenced, given that one plant growing mostly in deeper soils next to rock
exposures has been found with an ascending, elongate rhizome typical of other members of the subsection.
The habit of “sporadic heterophylly,” in which most Guadalupe Mountain plants within a given population are
completely unlobed whereas others produce mid-season leaves with notched or shallowly lobed (rarely deeply
dissected) blades, is novel among North American violets. Populations of other heterophyllous-leaved violet
species are typically comprised of plants that virtually all produce lobed or dissected blades during mid- or late
anthesis and into early fruit. Plants of the new species with variously unlobed or divided leaves are otherwise
identical in every other feature and are intermingled throughout a given site.
A de novo or recent hybrid origin for the violet under consideration is untenable, given that no other het-
erophyllous-leaved species inhabit the south-central U.S. within hundreds of kilometers of the region in ques-
tion. Two other, strictly unlobed-leaved members of the subsection, Viola missouriensis and V. nephrophylla,
have been recorded in nearby counties, although neither approaches the Guadalupe violet in any but a few
disparate traits. The available macromorphological and micromorphological evidence supports recognition of
the acaulescent blue violet in the Guadalupe Mountains as a divergent new orthospecies with an unusual re-
striction to limestone substrates. Characteristics and their measurements presented below follow Gil-ad
(1997), and are organized generally to allow direct comparison with his descriptions of accepted orthospecies.
TAXONOMY
a R.A. McCauley & H.E. Ballard, sp. nov. (Figs. 1-3). Type: U.SA. Texas. Culberson Co.: Si
It blue lines, 3 May 1947 (fl, fr), R. McVough 8149 (holotype: TEX!; iso
Acaulescent, heterophyllous, perennial herb in crevices of limestone, 3-9(-14) c
praemorse, rarely oblique and somewhat elongated, typically to 2.5 cm long, 5-7
i tall; rhizome vertical a
am diam., bearing stout
Journal of the Botanical Research Institute of Texas 7(1)
Journal of the Botanical Research Institute of Texas 7(1)
McCauley and Ballard, A r
colored when dry), sometimes finely spotted with red-purple, smooth, glabrous, (6.5-)7-9 mm long. Cleistoga-
mous peduncles erect or arching to declining, distinctly shorter than the leaves, slender, glabrous, green,
sometimes flushed or finely spotted with red-purple. Cleistogamous flower buds to 3 mm long; capsules
broadly obovoid to subglobose, green in life (straw-colored when dry), smooth, glabrous, 4.5-6 mm long.
Seeds from chasmogamous capsules yellow-brown (ISCC-NBS #74) with numerous slightly darkened patches,
broadly obovoid, ca. 1.75 mm long, 1.25 mm wide, with a small lateral caruncle 0.30-0.5 mm long extending
'^h-% the length of the seed from the funiculus; seeds from cleistogamous capsules slightly narrower and paler
than chasmogamous ones, yellowish olive-brown (ISCC-NBS #87) with somewhat more contrasting darker
patches giving a conspicuous mottled appearance, particularly toward the basal end, obovoid, 1.50-1.75 mm
long, 1.05-1.15 mm wide, the caruncle identical to that in chasmogamous seeds. Seeds of both types under
magnification displaying primary coat structure with cells superficially tetra-, penta- or hexagonal, isodiamet-
ric or elongated, the cell boundary furrowed, outer periclinal walls unevenly smooth; secondary coat structure
represented by symmetrically narrowly oblong, pentagonal or rounded, ring-like structures of spaced parallel
segments, the “basin” within the ring-like structures more or less uniform in elevation and mostly depressed
well below the top of the structures.
Distribution and Habitat. — Viola calcicola occurs in scattered locations principally along the eastern flank
of the Guadalupe Mountains and occasionally in the upper reaches of the larger mountain canyons. It is con-
fined to cracks of Permian Age limestone in sheltered canyons and springs. The species is commonly found
beneath an overstory association of Acer grandidentatum Nutt., Arbutus xalapensis Kunth, and Quercus
muehlenbergii Engelm. At the type locality V. calcicola, while common, is highly localized to rocks in close
proximity to water (e.g., rocky stream terraces or canyon walls flanking streams). The geographic distribution
of the species is hundreds of kilometers distant from the nearest population of other North American hetero-
Phenology. — Blooming April to May. Fruits present May to October.
Etymology. — ^Viola calcicola is named in recognition of its
Taxonomic distinctions and affinities. — Other currently recognized heterophyllous-leaved orthospecies in
subsection Boreali-Americanae are Viola egglestonii Pollard, found in limestone cedar glades of the east-central
U.S.; V septemloba Leconte, located in sandy open pine savannas in the southeastern U.S.; V nuevo-leonensis W.
Becker of sandy open oak-pine forests in northeastern Mexico; and V triloba Schwein., found in sandy loam or
clay soils in dry-mesic forests of the eastern U.S. and Canada (Gil-ad 1997). The new violet shares a restriction
to limestone substrate with V egglestonii, although the local floras of the two violets are dramatically different.
Viola calcicola resembles V egglestonii and V. septemloba in its glabrous foliage and eciliate sepals but diverges
in its peculiar sporadically heterophyllous nature. However, blades on lobed-leaved plants are similar to those
of V triloba var. triloba and V. nuevo-leonensis as they are usually shallowly divided, with the sinuses of the
lobes reaching less than halfway to the petiole summit, and the terminal lobe triangular or oblong-ovate. In V.
calcicola occasional plants with deeply divided leaf blades closely resemble typical V. egglestonii. Corolla color
in plants of certain sites often varies from nearly white to medium violet, and this appears to be at least partly
related to topographic position (for instance, plants with moderately purple or bluish-purple corollas are gen-
erally found at or near the bottom of a canyon or escarpment, while plants with nearly white corollas grow
principally in exposed microsites at or near the top of the canyon). Its sparse and localized lateral petal beard
is similar to that of V nuevo-leonensis and unlike other North American members of the subsection. Specimens
with unlobed leaves have been misidentified frequently as V missouriensis, whereas plants with conspicuously
lobed leaves have been misidentified most often as Viola (x) viarum. A misidentification of specimens assigned
to the latter may represent the basis for the disjunct county record of that taxon in the BONAP distribution
map. The Guadalupe violet shares most of its micromorphological features in petal trichomes and seed coat
with V septemloba.
li SW of Carlsbad, BLM Lonesome Rktge ACEC, perennial, growing in narrow cracks in
a, T26S R22E Sec 19 N % of SE Vi, 5040 ft, 29 Mar 1989 (fl), W
7 Apr 1937 (fl), L.N. Goodding2323 (NM
Journal of the Botanical Research Institute of Texas 7(1)
McCauley and Ballard, A r
n. (GUMO); Bear C:anyon, Pocket depres
t, 28 Apr 2002, (fl), Blaxland s.n. (GUMO);
trick Canyon, GMNP, fairly (
,uer,15Aprl988(n),A.M.Pc
i 1949 (fr), B.L. Turner 1249 (SRSC); frequent in i
30 April 1961 (n,fr),B.H.
Hunter’s Lodge, Guadalupe Moui
Canyon, Guadalupe Moui
18 Mav 1958 (fr), B.H. L M.C Johnston^6536 (SRSC); infrequent perennial, lavender flowers, in lower
i, 6000 ft., 18 May 1938 (fl), B.H. Wamocfe and M.C. Johnston 16556 (SRSC); infrequent pere
al Park, 26 July 1971 (fr), T. Weston 147 (SRSC).
Spring, Guadalupe Mt
DISCUSSION
Recosnition of V cdcicola adds a second endemic Viola to the Guadalupe Mountain flora, and the fimt hemro-
member ortheBomoli-Americanaegroup to the westernU.5.The^^^^^^^^
^is Powell and Wauer of sec. aamaemelanimn also occurs in 1 mestone along the ^
Ridge and is known from approximately 4 reported populations along the eastern escarpment ot the m
taii (Powell & Wauer 1990; Mullet etal. 2008; Marcussen et al. 2011; F. Armstrong, pers^comm.).
specimens of a third violet, referred to by others as Viola purpurea Kellogg, represent a ddferent yeUow-
emd species of Sect, Chamaemelauium that has been rarely collected in upland areas of the l^tns Mountaim
and in'^the Texas portionoft^G*^^
^“g^rnAe^tusonhCtodirthirisdisjunctby more than 50^
lations of the Purpureae to the west in Arizona. nf>w ^necies is
Theorigin and evolutionary relationshipsof Viola calcicalaare as yet unclear, although the new species^
clearly related to other heterophyllous-leaved eastern species. However, the ™
others in many vegetative and reproductive traits as well as the
chomes and seed coat. It shares fewer features with the other he* oovr*. an ecu
recognized species share amongst each other, suggesting long-t
erally, V. calcicola shares the most macromorphological its short vertical
mns'frTnd KMrX”tvreTkno?l^^ cuc!llale trichomes of ihe ktctal petals with eastern North
American V. cucullata. Its unusually broad range of corolla color pigmentation, even
correlate with elevation or ecologicalexposure(orboth),withplantsatthe bottom
producing deeply pigmented corollas and plants near or at the top pn^ucing
tionis that less pigmented (more reflective)corollas at higher topographic positions aidmreducing the delete
rious effects ofultraviolet radiation on reproductive structures in more exposed, sunny micro^^^^
Unlobed-leaved plants of V. calcicola could be mistaken morphologically for Viola missounensis or .
nephrophylla, both of which are reported from southeastern New rtoLTsp^^
ranee Uinit However the new species is easily distinguished from both by its short vertical rhizome, sparse
range hmiL im . ^ Generally clavate to knob-shaped, cucullate lateral petal trichomes; and yel-
lo^h town Ids'^rnumemus darker bmwn patches. Its
ates it further from V.misso«riensisa"'l’»‘=‘'l=’hroussDurred pet istinguis esit om
The inferred close relationship of Viola calcicola with V egglestonii and its broader affinities to other cat
tral U.S. and Mexican heterophyllous-leaved species are based on available morphological traits and ecologid
observations. Nevertheless, V. calcicola diverges in many features from other heterophyllous-leaved violets ami
possesses a few characteristics that are highly unusual or unique in the subsection. The proposed relationshf
hypotheses are testable using appropriate molecular markers. The authors are currently isolating and evahiat
ing the efficacy of potentially informative microsatellite loci as well as several nuclear and chloroplastgeneit
gions with sufficient polymorphisms to test such hypotheses. Future genetic studies on Viola calcicola and its
Boreali-Americanae kin will potentially illuminate phylogenetic relationships, biogeographic affinities, and
evolutionary origins of the new Guadalupe Mountain endemic and other North American species in this taxo-
nomically challenging but evolutionarily intriguing group of violets.
ACKNOWLEDGMENTS
The authors would like to thank the curators of the following herbaria for their loans of material or for provid
ing information regarding specimen data; MICH, NMC, NY, SRSC, TEX/LL, and UNM. We would also like B
thank A. Michael Powell and Rebecca A. Peters for their useful comments on the manuscript. RAM wouldlih
to thank Aurea Cortes-Palomec for assistance in the field and proofreading of the Spanish abstract, Fred Arm-
strong for providing information about the distribution of Viola in Guadalupe Mountains N.P., and Jonena
Hearst for providing access to the GUMO herbarium. HEB would like to thank Paul Knight, Tim Lowrey.Bob
Sivinsky, Larry Paul and Melissa Parker for guidance to the Big Canyon site, Paul Meloche for field assistance,
and Brian Kemmenoe at The Ohio State University Campus Microscopy and Imaging Facility for his expertise
with the SEM studies.
REFERENCES
Gil-Ad, N.L 1 997. Systematics of Vio/a subsection Boreali-Americanae. Boissiera 53:5- 1 30.
Gil-Ad, N.L 1 998. The micromorphologies of seed coats and petal trichomes of the taxa of Viola subsect. Boreali-omen-
canae (Violaceae) and their utility in discerning orthospecies from hybrids. Brittonia 40: 91 -1 21 .
GtfAsoN, H.A. AND A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada, 2 ed
New York Botanical Garden, Bronx, NY.
Kartesz, J.T. 201 1. The Biota of North America Program (BONAP). 2011. North American plant atlas (http://www.bo«d
org/MapSwitchboard.html). Chapel Hill, N.C [maps generated from Kartesz, J.T. 2010. Floristic Synthesis of Nottii
a Program (BONAP). (in press)].
F. Armstrong. 2011. Establishing tl
J.Bot. 98:1978-1988.
McKinney, LM. 1 992. A taxonomic revision of the acaulescent blue violets (Viola) of North America. Sida Bot. Misc7:1-d<l
Mullet, T.C., F. Armstrong, B. Zank, and C.M. Rrrzi. 2008. Predirting Viola guadalupensis (Violaceae) habitat in the Guadali4>«
Mountains using GIS: evidence of a new isolated population. J. Bot. Res. Inst. Texas 2:677-684.
Powell, A.M. and B. Wauer. 1990. A new species of Viola (Violaceae) from the Guadalupe Mountains, Trans-Pecos Texai
Sida 14:1-6.
Russeu, N.H. 1965. Violets (Viola) of the central and eastern United States: An introductory survey. Sida
WomuiNGTON, R.D. 2002. Inventory of the flora of the Guadalupe Mountains, New Mexico and Texas. Floristic Inventode^
of the Southwest Program, El Paso.TX.
TWO NEW SPECIES OF ERJASTRL7M (POLEMONIACEAE) FROM CALIFORNIA
David Gowen
111 Roble Road
Oakland, California 946 18, U.SA.
1davidgowen@gmail.com
RESUMEN
The genus Eriastrum, comprising 16 species from western North America, all of which occur in California, was
most recently treated by De Groot et al. (2012). The purpose of the current paper is to provide names for two of
the several unnamed taxa mentioned therein, both previously identified as E. hoovcri Qeps ) H. Mason.
Eriastrum hooveri, with corollas 5-6 mm in length, is one of the smallest flowered members of the genus.
The stamens are included and usually do not reach the corolla lobe sinus. Depending on flower size, E. hooveri
has stamens 1-1.5 mm long, inserted slightly more than that length below the sinus. The flowers are always
white. The only other species with corollas nearly as small is E. obramsii (Elmer) H. Mason. These two species
are easily distinguished from each other by a variety of characters, including leaf and bract lobe number, the
amount and location of woolly pubescence, flower color, and stamen length and position. Eriastrum obramsii
andbothofthetwonewspeciesproposedin this paperhavecorollasthatare pale blue,orhavebluestreaking.Al-
though many Eriastrum species can be recognized by vegetative or flower size and color characters, perhaps the
most important diagnostic characters involve stamen length and position (Mason 1945; Harrison 1959, 1968).
The seemingly minute stamen differences between these Eriastrum with small corollas, is more readily
apparent than the measurements would suggest. Splitting the corollas lengthwise and observing with a micro-
scope works well. 1 have also found that collecting corollas into a clear plastic sandwich bag, and placing this
in a plant press until dry, is useful. This allows one to simply hold the bag up to the light and observe the ar-
rangements of the flower parts with a hand lens.
TAXONOMIC TREATMENTS
e D. Gowen, sp. nov. (Fig. 1 A-E). Tm: U SA. Caufornia. C
Costa Co.: Lime Ridge Open Space, S
i; herbage lightly floc-
r of lateral lobes or entire, lobes 2-8
mm long. Inflorescences: heads 1 to few, floccose, 0.5-1.0 cm long excluding tips of bracts; bracts to 1.4 cm
long, exceeding heads, with 1 pair of lateral lobes. Calyx 4-7 mm long, densely woolly, lobes unequal, tips not
obscured by trichomes. CoroUa regular, sub-salverform, 5-7 mm long, throat plus tube 3-5 mm long, lobes
appearing almost white but with pale blue tint or blue streaking, elliptic-acute, 2 mm long, 1 mm wide. Sta-
Journal of the Botanical Research Institute of Texas 7(1)
mens 1.5-2.5 mm long; filaments 1 25-2 mm Inna cinnc
03-0.5 mm long, reaching the corolla lobe sint^’and usually e'^mdTt
stigmas0.25-0.5mmlong.Capsule4mmW ImmwiH ^ ^ ca. 2.5 mm long,
g ^.apsuie ^ mm long, 2 mm wide, 3-loculed, each locule 2-4 seeded.
saddle SE of microwave towers on summit, 18
Gowen, New species of Eriastrum
OcU«E„„r n»wera4,.„2003,G».„„aEPS),E,.d^
OEPS); power line area, 6 Jul 2005, Gowen 472 OEPS).
Dislrikution and lwbital.-Lime Ridge Open Space, slightly northwest of Mt. Diablo,
Clavton and Walnut Creek, is the only known location for E. ertterae. There are three sma si
plant grows, distant from each other by0.2-0.7kni.lt grows in finesandysoU in openingsor the
chaparral. Depending on weather patterns for any given year, each site might only have several u
son most likely with erroneous location data, that was mote likely collected in southern Califomta ®randegee
collMtons with erroneous locality information are, unfortunately, common^ It has of Moun
two decades thatE.al)mmsiiwasfound at two locations approximatelySandSkm east of the summttof
"efom a'^Ue in June 2003 to hud an Eriastrum at Lime Ridge, about 8 km northwest of
Mount Diablo while searching for a perplexing Navarretia I had seen there many years before. A review of the
supplementin' the just-published updated flora of Mount Diablo (Enter & Bowetman 2002) revealed that tor-
ham Enter and Tony Morosco had collected dried fruiting plants (not identifiable to
locahtyin 1998 Unfortunately, fresh floweringmaterialwas no easier to determme,asitdidntfitwellwithany
descritod species. Seveml years of additional fieldwork, morphological analysis, and common garden expen-
ments have confirmed that this extremely localized entity is a distinct taxon. , i
EriastrumertteraeismostsimilartoE.hooveri,buthasmorerobustcorollas(sizeandtexture),wrthapale
blue tint or streaking on the lobes. The stamens are longer, and the position of the anthers re ative to t e co-
rolla lobe sinus differs; E. ertterae has anthers placed at, or just beyond the sinus. The attachment
filament is almost at the mid distance of the total corolla length. By comparison, E. hooven has corolla lobes
that are proportionally longer to the total length of the corolla. Because the stamens are shorter and inc^de^
■he authersL placed not far beyond the mid length of the corolla. Geographically, E. ertterae ut
the closest population ofE.hotwerimthePanocheHilUof western Fresno County by apjrroxttnateVl^^^^^
BloomU time was found to differ between E. ertterae and E. hooveri, both m the field and wtth plants
grown under Lmon garden conditions by the authorinOakbnd,Cahfornia. The fimthowem^
E ertterae were almost a month later than for E. hooveri, which were grown using seeds collected from repre-
“X“^e;ri:™rhamEr^^^^
tion fo/the encouragement and guidance that she has given me over many years. For all the energy s e as
focused on the localBay Area flora, itisonlyfitting to haveavery local plant namedmher honor.
Eriastrum rosamondense D. Gowen, sp. nov. (Fig. 1 F-J).
SW quadrant of West Avenue D and 30th St. West, 12 May 2010, Gt
«ifcn„*l™E,h»„rtl.,h.v.agco™n.spakbl„.(».he,.tawhUe),.na~.^^
included). Differing from E. diifusum by having smaller flowers, and leaves entire (rather than leaves m y
nantscrect to spreadingannuals,2-8(-ll)cm; stem simple or branching, usuallyfrom^
to 1.5 cm; herbage and stems hghtly floccose to glabrescent. Leaves light green, to 1.5 cm ong and 1^ ,
subulate-awn tipped, entire. Inflorescences: heads 1 to few, floccose, 4-7
bracts4-14mm long, withlpairoflaterallobes 3-4 mm long, theseoftenrecunredC^^^^^
Iv woollv except toward the tips, lobes unequal, tips not obscured by tnchomes. Corolla regular sub sal e
fL5-6(-7)Llong throat plus tube ca.4mmlong,lobespalebluebutsometimesappearingalmost whim
b^'^hc-acute wi* ttps sLetimes slightly rounded, l.«(-2.)
mmlr=^nrf "“^or slS^ljTtoyrJthe mTIJIngrrf ^ coTl lobe. Style ca. 3 mm long, stigmas
ca. 0.3 mm long. Capsule ca. 4 mm long, 2 mm wide, 3-loculed, each locule 2-4 seeded.
Journal of the Botanical Research Institute of Texas 7(1)
part (RSA); Lancaster, May 1909, Brandegee s.n. (UC);
telope VaUey, W of Lancaster, N side of Avenue G, 27 Apr 1998, Porter
1 1834 (C:AS, RSA); Hwy. 138 (West Ave. D) Wof Lancaster, NE comer of intersection with 40th St. W., 11 May 2005, Gowen 309 OEPS); Hwy
138 (West Ave. D) W of Lancaster, areas on both sides of the road at the intersection with 30th St. West, 21 May 2003, Gowen 351 OEPS);
intersection of Hwy 138 (W. Ave. D) and 40th St. W. on the SW comer, 8 Jun 2005, Gowen 412 QEPS); 40th St. West and Avenue D, 12 May
Distribution.— The presently known distribution covers only a several mile area between Rosamond and Lan-
caster. It is centered near the intersection of Hwy 14 and Hwy 138 (West Avenue D), near the northern border
of Los Angeles County. Such a small area of occurrence for E. rosamondense makes this a rare plant deserving
of conservation.
Discussion. — Plants now proposed as E. rosamondense have been known from the Lancaster area from at
least 1892. A mixed collection of this plant and E. sapphirinum, by Davidson (RSA 469271) in that year, and a
Brandegee collection in 1909 (UC 130872), are known to the author. These collections have been referred to E.
dijfusum in the past. A collection by R. Hoffman, 29 Apr 1930 (CAS 178681) has been variously attributed to
Giliafilifolia var. diffusa [E. diffusum], annotated as “sp” by Harrison, and annotated by Hoover as E. hooveri.
More recently, plants from near Lancaster were attributed to E. hooveri (Boyd & Porter 1999). All of these col-
lections represent E. rosamondense. There is one known occurrence of E. hooveri from 7 km east of Isabella
Lake, a low valley near the southern end of the Sierra Nevada (Gowen 945), but E. hooveri is otherwise not
known from east of the San Joaquin Valley. It does not occur in the Mojave Desert.
The small flowers of E. rosamondense are easily confused with E. hooveri, but the pale blue flowers and
exserted stamens of E. rosamondense distinguish the two species. More problematic is the relationship with E.
dijfusum. Structurally, E. rosamondense is like a miniature of E. diffusum, and its placement as a subspecies of
that taxon was considered. However, E. diffusum does not occur closer than 100 km to the east, in San Ber-
nardino County. In addition, the leaves of E. rosamondense are entire, whereas E. diffusum most often has leaves
with a pair of lateral lobes.
Etymology.— The specific epithetrefers to the Rosamond Dry Lake area between Lancaster and Rosamond.
ACKNOWLEDGMENTS
I would like to thank the curators and staff at UC^EPS, CAS, RENO, RSA, and SBBG for the gracious use of
these facilities. 1 appreciate the insightful suggestions from Robert Patterson and Dieter Wilken, who reviewed
this paper. Barbara Ertter and Leigh Johnson provided many helpful comments on the manuscript, and over
many years, have greatly increased my knowledge of all things botanical. I'm grateful to Raiil Pozner (Darwin-
ton ^tamcal Institute, Argentina) for translation of the abstract. The illustration by Linda Vorobik was done
Press, Berkeley. Pp. t (Ml -1043.
Mason, H.L 1 945. The genus Eriastrum and the in
in Polemoniaceae. Madrono 8:65-91 .
CAREX CYRTOSTACHYA (CYPERACEAE). A NEW SPECIES OF SEDGE
ENDEMIC TO THE SIERRA NEVADA OF CALIFORNIA
Peter F. Zika
WTU Herbarium, Box 355325
University of Washington
Seattle, Washington 98195-5325, U.S.A.
Zikap@comcast.net
Lawrence R Janeway
The Chico State Herbarium
California State University, Chico
Chico, California 95929-0515, U.S.A.
Feather River Ranger District
Plumas National Forest
875 Mitchell Avenue, Oroville, California 95965, U.SA.
Uaneway@fs.fed.us
Barbara L. Wilson
Carex Working Group
1 377 NW Alta Vista Drive
Corvallis, Oregon 97330, U.S.A.
bwilson@peak.org
Lowell Ahart
9771 Ahart Road
Oroville, California 95966, U.S.A.
ahartbrothers@yahoo.com
•a Nevada of California. Detail
Several workeK in ihe llora of the northern Sierra Nevada of California have been puzzled by a sedge resem-
bling Curex ntendurinensis Olney ex W. Boon, but which differa from it on the basis of
and is disjunct from the coastal and coast range habitats of that species Oaneway 1992; Oswald & Atan im
Oswald 2002; CUfton 2003, 2005; Taylor 2010). After studying the Sierran populations, we describe them here
as a new species. It was first collected in 1968 by G. Ledyard Stebbins.
Carex cyrtostacliya Janeway & Zika, sp. nov. (Fig?. 1, 2A, 3A-B, 5)
Road 20N03 (Mooreville Ridge Road), 2 air mi SE of Sugar Pine Point, Pinkard Creek drt
Wilson, Ahart, Janeway (HOLorYre: WTU; isotypes; BRU, CAS, CHSC, DM
CarexortosrnchyadiffersfromCarexmendocinensisinitsmorenexuousdistalla
rnT'ce^I, with lort rhizomes; roots brown, not densely felted with root hairs Tertile cul^ (stems)
20-132 cm long, 0.4-2.3 mm wide, trigonous, smooth on the angles; basal sheaths scale-hke. Hapless, wtth
rale veins scabrouSHtillate, dark purple, grading above into red-purple, red-brown, and green sheaths the
upper of these bearing blades. Letwes (2-)3-5, mostly on lower half of culm; sheatte 5-80(-142) mm long
h^dy clasping culm, glabrous to scabrous, green, the sheath faces (opposite blade) green to whitish, often
m, 5 Aug 2011, Zika 25715 6-
f,OSC,RSA).
widely spaced perigynia, longer and
Journal of the Botanical Research Institute of Texas 7(1)
dotted with red or purple, glabrous or scabrous, the apex (mouth) U- or V-shaped, dilate, fragile; ' l-
“ long, longer "hau wide, apex acute or acuminate, occasionally rounded cliate; blades 0.5-42 cm
loTna S-4 5f-6')mmwide flat V- or W-shaped in cross-section, occasionally strongly scabrous on ad-
axWurfacenearligule,otherwi« smooth, with l-3paleprominentabaxialveir«,ttm,ginsscabrousd«^^^^^
Vegetative shoots similar to fertile shoots; pseudoculms (overlapping leaf sheaths of vegetative shoots, see
Remtcek 1986) 4-6 cm tall, with 3-6 leaves, the larger basal leaves 2.5-5 J mm wide. Ptoxtmal
bracts withsheaths(2.5-)6-42mm long, .be blades shorter than to exceeding mflorescence.5.5-^5cmong,
0.9-2.5 mm wide, gUbmus or slightly dilate, the distal bracts much reduced, ^rescimees 5-47 im long,
the lateral spikes pistillate, .be terminal spike staminate; proximal lateml spikes on peduncles 17-171 mm
Journal of the Botanical Research Institute of Texas 7(1)
long, the portion exserted from the sheath 10-1 14 mm long (rarely the proximal spike branching once from the
basal pistillate scale, Janeway 9080 CHSC, Rubtzoff 7088 & Howell CAS); lateral spikes solitary at the nodes,
arching, often drooping at maturity on flexible, smooth or slightly scabrous peduncles, the two proximal
spikes separated by 2-28 cm, the distal spikes overlapping, on peduncles 5-25 mm long, rarely sessile; termi-
nal spike erect on peduncle 5-26 mm long. Spikes 3-6, the lateral spikes pistillate (very rarely androgynous)
cylindrical, 18-93 mm long, 2-5.7 mm thick, with (10-)14-29 perigynia attached 0.7-3.5(-6) mm apart the
terminal spike staminate, 10-64 mm long, 1.1-2.3 mm thick, rarely gynecandrous (Rubtzoff 7088 & Howell
CAS)and the lower half staminate.PistiUate scales usuallyshorter than mature perigynia, obovate to oblong-
obovate, (2.2-)3-4.8(-6.8) mm long including glabrous or ciliate awn, acute, rounded or notched at apex
(apart from awn), glabrous with ciliate or erose margins distally, white-hyaline or faintly marked with red-
brown, uncommonly red-spotted; awn (0-)0.5-2.2(-3.1) mm long; midrib broad, green, aging brown with
1-3 vems, ramly with sparse bristles distally. Staminate scales densely overlapping, clasping at base, oblan-
ceolale to oHong-obUnceolate. 4-6.9(-8) mm bng, acme, roumled, or rrotched at apex, hyaline with green
midvetn, ^abrous, the proximal scales with prominent awn 0.5-3(-5) mm long. Anthem 3, 18-3 mm long
s tg t y ctlmte at a^x (40x1 Pengyma erect-ascending or oppressed when mature, (3.2-)3.8-5.8 mm long,
1. 1-1.8 mm wide, bases gradually narrowed to darker stipes 0.3-0.6 mm long; body * flattened-trigonous ±
sy_, mrop View, Cribbed and Bnely 12-15-vei„e4
p s ^ or purple dots, membranous, glabrous, gradually tapered to beak; beak (0.5-)0 7-1 4 mm Inno
(measured from inflection point); apical teeth obscure, 0.2-0.6 mm long often ciliate on nr h^t .u ^
Ache^ (nutlets) 1.9-2.7 mm long (not including style remnant orstipeU.1-1.6 mm wide trigoZl XT
7 f ‘ by perigynium pale brown naniZ
Cumx cvnZt aTr'" ® “ ^0“ fro" r.cessTons of
waxycuticleandtheouterpericlinXXlhhr^f^^^^^^^ """
Cure., mcndociitensis tends to have many papillae (unZT °7i Z“" "P“'“ Warent.
papillae often occupieda smaller percena^of*empZr''T7"''’'7'
.ended robe narrowandwithout':n.meS t“*^^^^^^
faces were simiUr for the two species (Fig 4C F 7l 77 “chene sur-
those of C. hirtissinta w. Boott (Waterway 1990a, Fig.V ' “..hene surfaces also resembled
exposure. Imnges Iro^fuM 7Zion 7 7 *P™*ng on elevation and
terraces on more mesic soils. "P"^" "“h samratedsoil, a! wel7' ZwTn
Cur« cyr,mmcl.ya commonly grows in the shade of wood
Torr. & A. Gray, Corylus cornuta Marshall, Lithocarpus densiflorus (Hook. & Am.) Rehder, Pinus contorta
Douglas ex Loudon, P. lambertiana Douglas, P. ponderosa Douglas ex Lawson & C. Lawson, P. sabiniana Doug-
las ex D. Don, Pseudotsuga menziesii (Mirb.) Franco, Quercus durata Jeps., Rhododendron occidentak (Torr. & A.
Gray) A. Gray, Rulms armeniacus Focke, R. laciniatus Willd., and R. leucodermis Douglas ex Torr. & A. Gray.
Herbaceous associates are diverse, corresponding to variation in soil moisture. Some typical associates are
Adenocaulon bicolor Hook., Agrostis palkns Trin., A. sccAra Willd., Aquikgia formosa Fisch. ex DC., Carex ampli-
folia Boott, C. bolanderi Olney, C.feta L.H. Bailey, C. hirtissima W. Boott, C laeviculmis Meinsh., C. lemmonii W.
Boon, C. pachystachya Cham, ex Steud., C. stipata Muhl. ex Willd., Circaea alpina L., Equisetum arvense L., E.
hyemale L., Fragaria vesca L., Galium bolanderi A. Gray, Geum macrophyllum Willd., Hieracium albiflorum
Hook., Juncus effusus L. subsp. pacificus (Feraald & Wiegand) Piper & Beattie, J. exiguus (Femald & Wiegand)
Lint ex Snogerup & Zika, J. laccatus Zika, J. trijormis Engelm., Linnaea borealis L., Luzula comosa E. Mey. var.
laxa Buchenau, L. parviflora (Ehrh.) Desv., Mimulus moschatus Douglas ex Lindl, Pteridium aquilinum (L.)
Kuhn, Rhynchospora capitellata (Michx.) Vahl, and Scirpus microcarpus J. Presl & C. Presl.
ving or pendulous (jcyrto-) pistillate
Etymology. — Carex cyrtostachya, or arching sedge, is named for its a
spikes (-stachya).
Distribution. — The range of Carex cyrtostachya is limited to the northern Sierra
northern high Sierra Nevada geographic subdivisions (Baldwin et al. 2012), from El Do
Yuba and Butte counties, California (Fig. 5). Populations of C. cyrtostachya are discontir
on the western slopes of the Sierra Nevada, with one cluster of populatio
adjacent Yuba County, and the other in El Dorado County. Only 13 populations have been documented, most
of them quite small, within an area of 120 x 30 km. No populations have been found in the intervening Placer,
Nevada, or Sierra counties. Arching sedge is thus a narrow endemic, restricted to just three counties in Califor-
nia. There are 12 other endemic Carex species in California (Mastrogiuseppe 1993, Zika 2012, Zika et al. 2012),
as well as additional endemic members of the Cyperaceae, such as the recently described Eleocharis torticulmis
S.G. Sm., restricted to a small area in the northern high Sierra Nevada of Plumas County (Smith 2001).
Conservation.— Most Carex cyrtostachya populations are small, difficult to locate, and potentially threat-
ened by logging and invasive Rubus. The species should receive some conservation attention. More inventory is
needed and systematic census data are lacking, but based on our initial field surveys and observations, and the
her^num records, there may be fewer than 20 populations and 2000 plants in total. A few populations are on
land managed by the US Forest Service, in the Plumas and El Dorado National Forests, but other locales are at
low elevations on private land and susceptible to development or hydrological changes.
an Creek bridge on Bean Creek Road, 609 m, 29 Jun 1988, Ahart 6108
J m, 2 Jul 1988 Janeway 2955 (CHSC, WS); small lake 1 mi S of Brush
'of Lost Creek Reservoir, 1036 m, 6Jul 1993, Ahart 7052 (CHSC); 0.5 mi
CHSC, DAV,HSC,JEPS,WTU); same site, 6Jul 2011, 2Sfea 25535 (CAS,
.2 road mi N of Lost Creek Reservoir dam, 6 Aug 2006 Janeway 8826 (CHSC WTU)- same
^ -ipper W branch of Know-Nothing Creek, Lumpkin
'aEPS,CHSC);sa '
miv 8868 (CHSC, WTU); S Fork Bacon S^onrarfelhicl™lRl^“^ ^ ^006 Jane-
same site, 19 Jul 1973, Rubtzpff 7187 & Holell ((^); Rlckc™yoTcreek2TO ^
Creek Special Interest Area, El Dorado National Fn«>« fisn ^ q i i “Pstream of confluence with Traverse Creek, Traverse
6-;a«cwa>(CHSC,OSC,WTl0. YubaCo.:HampshireCreekMealw^V“"r‘’pT’r‘^”^’.'^^^’ ^ay 2012, Zifea 25877
2007,Jan<way9116&Ahart, Hanson (WTU); same site, 7 Aug 2007 Ahart
Janeway (CHSC, jEPS,OSC, WTU). ’ ^ 5 Aug 201 1, 2iha25718& Wilson, Ahart.
CHSC. OSC, WTU); W side of Pinkard Crei
site, 6 jul 2011, Zika 25536 (CAS, CHSC, GH.JEPS, MICH, MO. OSC 1
CHSC, JEPS, NY
IZXx 'heirsy^ony^s (C. c.
establish,!, differences be, ween ,hema„dCc>nosk.41ZJZed.Z^^
.n synonymy might apply to C. cyrtostachya. The descriptions of several of rhe T
Whrch gathering was the type, bu, Wed to specify whiA duplicate wt IZ “ " “"P'”
type senes of c.mendociticnsis was a hybrid withe vv d ^ ^®*°‘ype. In addition, part of tl
and some lecmtypficariotrs were neeWThTZofZZ^
consecutively by Bolander and appear to be froZargl mkZZ W'e numbere
(1951) we belteve U is masonable to assume ) T Uke Howe
the ^pheates, a, DS, has the precise date of coUecion 1117^ urn ““ '
sts; hm original material at Harvard is the lectotype siktw b^f^' d T
Zika et al., A r
docinensis. The hybrid plants fail to exsert or dehisce their anthers and thus can be distinguished from the type
material of C mendocinensis on the same sheets, which, although often immature, has dehiscent anthers or
exserted filaments where the anthers were shed. Howell (1951) and Waterway (1988) reported isotypes of Bo-
lander 4701 at NY were also a mix with the hybrid, but we were unable examine them. We also inspected the
available types of C. cinnamomea Olney, a synonym of C. mendocinensis. Howell (1951) discussed the taxono-
my and cited type specimens at GH, NY, UC, and US, but did not select a lectotype. We designate Olney’s
original material at Brown University as the lectotype (Bolander 6477 BRU, ex herb. S. T. Olney). Carex gynody-
nama was also described by Olney (1868), citing only one gathering, Bolander 4700, but not specifying which
of the many duplicates was the holotype. Howell (1951) effectively lectotypified the name by citing Bolander
4700 (BRU) as the type. We were able to study the lectotype and nine isotypes of C. gynodynama; none ap-
peared to include hybrid elements.
Carex mendocinensis Olney ex W. Boott, Bot. California 2:249. 1880. Type: U.S.A. Caufornu. Mendocino Co.; swamps
DISCUSSION
Carex cyrtostachya is closely related to C. mendocinensis, and the two are compared in Table 1. Most specimens
arc easily segregated by inflorescence structure. The lateral pistillate spikes of Carex cyrtostachya arch or droop,
or dangle on flexuous peduncles, and the perigynia are more widely spaced (Figs. 1, 2A). Carex mendocinensis
usually displays erect or ascending distal lateral spikes that are densely fruited (Fig. 2B). Occasional plants of
C. mendocinensis show lax proximal spikes, but the distal pistillate spikes remain essentially erect. In addition
to the obvious inflorescence differences, several other characters are useful in identification Although none
are absolutely exclusive, in combination they allow reliable identification. The staminate scales of most plants
o C. mmdocmensis are awnless, although the lower scales rarely have an awn 0.3-0.5 mm (e.g.. Peck 8792 (GH),
Curry' Co., Oregon). Proximal staminate scales of C. cyrtostachya have prominent awns, often 1-5 mm long. In
^dition, the pistillate scales and their awns are longer in C. cyrtostachya, with some overlap in measurements,
he^ngyma and scales of C. mendocinensis are green, strongly marked with red or purple or red-brown;
T in comras, .he pis.lIU.e scale bodies ol
C. cyHosc^kya are usually wh.n.-hyaline, cmly rarely with a lain, red-brown wash, whereas .he perigynia are
the disunce bet^en .1^1.^^.^' placement within the perigynium differs slightly, so that
of C ntendochtemis ate often scabrous, while they tend m be smooth or fai«rscXusTn
Journal of the Botanical Research Institute of Texas 7(1)
ACKNOWLEDGMENTS
We thank the curators of BM, BRU, CAS, CHSC, DS, GH, HSC, ID, JEPS, MICH, NY, ORE, OSC, POM, RSA,
SOC, UBC, UC, WILLU, WS, anti WTU for access to their collections or loans. Some additional unaccessioned
materials were also examined at CAS, CHSC, and JEPS. For assistance in the field, we applaud Lowell Ahart
and Julie Nelson. We extend our gratitude to Joy Mastrogiuseppe, Tony Reznicek, Philippe Seguin, and Debra
Trock for their discussions and advice. Rebecca Peters at CAS was especially helpful in locating unmounted
collections by Peter Rubtzoff. The drawings were prepared by Krista Anandakuttan, the maps assembled by
Tom Ruehli, Fig. 3 photographed by Steve Matson, and the SEM images made by Wai Pang Chan at the Biol-
ogy Imaging FaciUty at the University of Washington; we are grateful to all of them. We are also indebted
to the three reviewers, Tony Reznicek, Leo Bruederle, and especially Ted Cochrane, whose comments and
suggestions resulted in a much improved manuscript. Comments from Barbara Castro also strengthened the
manuscript. Partial funding for herbarium study was provided by the Lawrence R. Heckard Fund of the Jepson
REFERENCES
Bacdvwn, B.G, D.H. Goldman, DJ. Keil, R. Patterson, TJ. Rosath, and D.H. Wilkin (eds.). 201 2.The .
of California, 2nd ed. University of California Press, Berkeley.
CuFTON, G. 2003. Plumas County and Plumas National Forest flora. Published by the auth
cy.CA.
s National Forest flora. Published by the author ai
s National Forest,
CuFTON, G. 2005. Plumas County ai
Quincy, CA.
Howell, J.T. 1951 . Studies in Carex— llhThe identity of Carex mendocinensis. Leaf!. W. Bot. 6:1 57-1 62.
Janeway, LP. 1992. Cyperaceae of Butte County, California. Part 1; Carex. Studies from the Herbarium California State
University, Chico, Publication No. 9.
Masthoouseppe, J. 1993. Carex, sedge. In: J.C. Hickman, ed.The Jepson manual: higher plants of California. Univ. of Cali-
fornia Press: Berkeley. Pp. 1 107-1 141 .
Olney, S.T. 1 868. Carices novae. Proc Amer. Acad. Arts 7-393-396.
Oswald, V.H. 2002. Selected plants of northern California and adjacent Nevada. Studies from the Herbarium California
State University, Chico, Publication No. 1 1 .
Oswald, V. and L Ahart. 1 994. Manual of the vascular plants of Butte County, California. California N
Sacramento, CA.
Hymenochlaenae (Cyperaceae) in Mexico and Central America. Syst.
SMriH, S.G. 2001. Taxonomic innovations in North American Eleocharis (Cyperaceae). Novon 11-241-257
Taylor, D.W. 2010. Rora of the Yosemite Sierra. Published by the author, Aptos, CA.
n Carex section Hymenochlaenae Drejer (Cyperaceae). Ph.D. dissertation,
e Plant ScKiety,
Cornell University, Ithaca, NY.
Vaterway, MJ. 1990a. Systematic ii
ceae). Canad. J. Bot 68:630-639.
racea^) in Callfomla. and C manddc/nensd
^ MJ. 2002. Carex Linnaeus sect Hymenochlaenae. In: f
Zika et al., A new species of Carex from the Sierra Nevada
of North America north of Mexico. Vol. 23, Magnoiiophyta: Commelinidae (in part), Cyperaceae. Oxford University
Press: New York. Pp. 461 -475.
Waterway, M J. and J.R. Starr. 2007. Phylogenetic relationships in tribe Cariceae (Cyperaceae) based on nested analyses of
four molecular data sets. Aliso 23:165-192.
Zika, P.F. 2012. Carex orestera (Cyperaceae), a new sedge from the mountains of California. Novon 22:1 18-124.
Zika PF, A.L. Hipp, and J. Mastrogiuseppe. 201 2. Carex. In: B.G. Baldwin, D.H. Goldman, DJ. Keil, R. Patterson, TJ. Rosatti, and
D.H. Wilkin, eds.The Jepson manual: vascular plants of California, 2nd ed. University of California Press: Berkeley. Pp.
1308-1339.
Journal of the Botanical Research Institute of Texas 7(1)
BOOK REVIEW
Amy Stewart. 2013. The Drunken Botanist: The Plants that Create the World’s Great Drinks. (ISBN: 978-1-
61620-046-6, paper overboard). Algonquin Books, P.O. Box 2225, Chapel Hill, North Carolina 27514-
2225, U.S.A. (Orders: www.amystewart.com). $19.95, 368 pp., 2-color illus., 6" x 8".
Amy Stewart got the idea for this book when she went to a liquor store and saw, in her wonderful imagination,
instead of bottles of liquor, the plants that make the drinks we know and love — or don’t know and don’t love as
the case may be. Her vision inspired a great little book for imbibers, botanists, gardeners, chemists, bacteriolo-
gists, and historians.
The book is handily sectioned into plants from which we create alcohol, the plants we use to flavor alco-
hol, and the plants we use to garnish our drinks. Each of these sections is in alphabetical order, making it easy
to find something specific. The index is quite detailed and is also helpful for anyone doing research. (The reci-
pes listed in the front of the book save time!)
In Part I, we are introduced to the plants used to make alcohol as well as the plants we use to age the brew.
This includes mostly grasses, fruit, and vegetables used to create alcohol. There are only a few oaks currently
used to make barrels in which rough spirits are tamed into the smooth scotch, whiskey, rye, rum, gin, & vod-
ka, with which we are familiar.
The history of various beverages is fascinating, and along the way Stewart debunks a myth or two. For
instance, vodka was made of grains long before the potato even arrived in Europe. Apparently Russia and Po-
land still quarrel over who invented the beverage. The history and importance of yeast in the whole process is
thoroughly and delightfully covered. ’Without the yeast, we would have no alcohol. My favorite category of side
articles in the book is “Bugs n’ Booze.” Earthworms, honeybees, and whatever falls off the rafters into the fer-
menting vat can bring yeast, flavor, and color to the mix.
Part II of the book covers the flavors we give to various alcoholic beverages. From allspice to wormwood,
angostura to sugar maple, apricot to yuzu, and almonds to walnuts, the reader is.led through a garden of fla-
vors. Given her directions, you could grow your own hops or sloes or citrus!
In addition, Ms. Stewart provides quite a few recipes and not a few horticultural tips on which species to
select and the prime conditions for growing various plants. There is a recipe for Capillaire syrup which uses
several stems of fresh maidenhair fern, water, sugar, and orange flower water. The resulting syrup comes in
handy if you wish to create Jerry Thomas Regent’s Punch.
Stewart winds up her book with garnishes for various drinks. Herbs, flowers, berries, vines, fruits, and
vegetables are touted here. A template for experimental garden cocktails will send you to your garden or the
local fanner’s market to try some or all of the recipes. You will be encouragerl to grow your own berries, fruit
and vegetables m a garden made exclusively for cocktails. She makes it sound easy.
Il''t>™"k™B«‘>"‘aisareallyfunread.Lotsoffactsatecrammedintothebook,bulMsStewartkeepsit
iiu luu latis lor cocKtail hour - " "
anyone who likes cocktails.— Kay Y,
Institute of Texas, Fort Worth, Texas, USA.
A RE-EVALUATION OF CAREX SPECUICOLA AND THE
CAREXPARRYANA COMPLEX (CYPERACEAE)
AAReznicek D.F. Murray
RESUMEN
INTRODUCTION
Plants referred to Carexparr^ana Dewey have always beenasource of taxonomic difficulty.Adding to this dif^
ficulty is that, even though some species are widespread, all are uncommon or rare. Carex parryana and its
close relatives differ from other members of section Racemosae in having ±elongated inflorescences that have
relatively narrow (2 4-6 8 mm wide), cylindrical, erect or ascending lateral spikes (if not unispicate), com-
bined with small perigynia 1.7-3.3 mm long (-3.9 in C. specuicola and C. utahensis). Mackenzie (1935) recog
nized four species in this complex, C. aborigii
ryana. Murray (1969) recognized C. parryana
E. Jones, C. halln Olney, C. idahoa L.H. Bailey, and C. par
; variable species consisting of di
nd treated them as subspecies; C. parryana subsp. halln (Olney) D.F. Murray, C. par
ryana subsp. idahoa (L.H. Bailey) D.F. Murray and C. parryana subsp. parryana. He did not treat C. obon^num,
realizing lhat ‘Carex olwriginum- as described in Mackenzie (1935) was close to C. parryana and a q-K
ent plane than C. nboriginum M E. Jones, which is a local Idaho endemic with much larger perig^m (4.W5-
6.6 mm long very similar to C. sermtodens W. Boon (Murray 2002). Hermann (1970) recognized C. hath,
idahoa, C. parryana, and C. aboriginum M.E. Jones as spe<
It noted that “Carex aboriginum” as described in
Maekenzie(19M)i; -completely different- from ME. Jones’s typ^Mype. He treatedCnborynum of
zie, not ME. Jones, as C. patrynnn var. brevisrparma FJ. Herm. Murray (2002) treated C. tdahoa. C linlln. C.
parryana. and C. ohoriginum as species, and also noted that Mackenzie s “C. aboriginum- was not C.
of M E. Jones, but did not recognize C. parryana var. brevisrfuoma. Finally, Goodnch (m Welsh et al. 2003)
concluded that C. specuicola, described by Howell (1949) as an endemic of hanging gardens m northern An.
zona, also belonged within C. parryana, and recognized only C. parryana in Utah. , ^ ,
in the last decade of floristic activity within the range of the Carex pa^ana compta,there has fortu-
nately been enough collecting to generate a great many new speamens. This w^ especially me lor C
specuLla, for which lew specimens existed when it was treated for flora of North America, Murray 2W2).
Botanists working with the Navajo Nation («e Roth 2004) discovered a number of new popuUuons, and we
38 Journal of the Botanical Research Institute of Texas 7(1)
now have an excellent series of specimens to study. We reviewed these new collections with several questions
in mind. Is C. specukola a good species? Is recognition of C. hallii, C. idahoa, and C. parryana as species sup-
ported by additional collections? Finally what is the identity of “Carex aborigimm” of Mackenzie (1935) not
M.E. Jones (= C. parryana var. brevisquama FJ. Herm.), and is it a good species? We also provide illustrations of
Plants referred to Carex parryana divide into two clear groups based on morphology and geography. One
group, including C. specukola, has strongly flattened perigynia much larger than and loosely fitting over the
achenes, an admixture of 2-styled and 3-styled flowers, and is essentially confined to Utah and Arizona. The
other group, including C. hallii, C. idahoa, C. parryana, and “C. aboriginum” of Mackenzie not M.E. Jones, with
the pengyma tightly enveloping the achene and entirely 3-styled flowers, is widespread, occurring from On-
tario to Alaska, but south only to Colorado, Utah, Nevada, and California.
The former group, with loosely fitting perigynia andamixof2-styledand3-styled flowers consists of two
species. One is the lax hanging garden endemic Carex specukola with pale perigynia. In addition, we recognize
anes^ntiallyalloF^tric and more widespread species fromadiversity of wetland habitats that isusually stiffly
upright, wnhclearlypurple-tintedperigynia, which wedescribeasCut^^^^^^ In the latter group, with pe-
ngyma tightly enveloping the achenes, the plants erroneously referred to Carex aboriginum by MackenL
i^wKlest 4-6.8 mm wuie. We recognize this taxonatspecies rank and dcscribeithemnasC.l.olmgrento™m
mm O 3 n.rt r “ ">“> “ 3.6
in Murrav Qonil ’ ' “ remaining species of the Cara parryana complex as recognized
I bestabuZhthe W^rl""”"^^^^^^^
eral spiL of C LlJii anH relatively broad, 3-5.5 mm wide Lat-
clrrhlfliUndr
and C. parryana show great variability in inflorescence size k .2 ,
ity. Nevertheless, most culms of Carex hallii iftyqftnf k k ^P^^e number, and spike sexual-
lon^tlatem, spike, in C.pnr,,nm.,ttZr::^~W^
X - “I”- ofient mfiTsTZo f 'rm “
heightrrZ^ofZlXXtmfcZ
Whi and C. parryana do not represent L emis otZmfZ ^^ows that C.
X parryana group
Spike number
2 3 4 5 6
Fk;. 1. Distribution of spike numbers in Carexhallii and C parryana.
Achenes 0.8-1 .2 nnm wide; lateral
5. Spikes (2-)3^-6), the 1
times as long as the longest
Loosely cespitose to slightly colonial from slender, ±elongate rhizomes 0.9-2.3 mm wide, rhizomes ca. 0.5-7
cm long between shoots, rhizome scales pale brown, rapidly disintegrating into fibers; culms (7.5-)19-70(-
diately below the inflorescence; phyllopodic, cataphylls and basal sheaths pale to dark brown, youngest some-
times tinged reddish-purple. Leaves 5-11, essentially basal; blades 1.8-38 cm long, 0.8-3.3(-4.5) mm wide.
Reznicek and Murray, Systematia of the Carex parryana group
stipe 0.2-0.45 mm long, lenticular or occasionally nattened-trigonous in cross section; the convex sides elliptic
to nearly circular in lenticular achenes, narrowly elliptic in trigonous achenes, brown, smooth; apex apicu ate
with an apiculus up to 0.15 mm formed by the peKislent style base; the style deciduous; stigm^ mostly 2,
sometimes up to 30% 3. Anthers 3, 1.2-2.7 mm long, with a conspicuous apiculus ca. 0.1-0.3 mm long.
Moist seeps in shallow caves or alcoves along sandstone cliffs— “hanging gardens.’ Care* spccuicota oc-
curs as local populations consisting of a few culms to as many as 400-300 plantssptead over many meters of the
cliff face. Elevation 1350-2323 m.Endemic.o northern Arizona andimmediatelyadjacentsouthemmos. Utah.
This is an endemic of Navajo sandstone “hanging gardens." Typical associated species of hanging gar ens
in ihe region include Adiantum cnpiUus-veneris L., Aniicica vuginula Rydb., Aquilcgia micrantlm Eastw., Cirsnim
rydbergii Petr., Epipoctis gigniitcn Hook., Mimulus caslwoodme Rydb., Plfltnnthera wtheam (Higgins & S.L.
4lsh*Kartesz & Lndht and a lew other Cnrex, especially C. hnssei L.H. Bailey. Though very local, this spe-
cies can form a dense turf and is the dominant cover in some sites. The lax, arching febit remains constant
even when the plants are growingonahorizontalsurface as shownin Figure 3, .hough this cantedifficu^o
discern in herbarium material. A sampling of perigynia, scales, and achenes are noted m Figure 4 an a yp
cal inflorescence from a herbarium specimen is shown in Figure 5B.
Representative specimens. ARIZONA. Apache Co.: Canyon del W
ASC); narrow canyonSWofIm
ie Chelly, 5 May 2001,
m 2276 (NAVA); Carson Mesa, 12Jun 2001, O’Kane, Jr. <
15 (NAVA); Walker Creek, 1 mi !
Mexican Water, 21 Aug 2003, Roth 1701 (NAVA); Tseyi-hatsosi Canyon N of Kayenta and SW of Boot Mesa,njul Wl^ev
N of Giant Canyon trail, 20 May 2002, Rinfe 1154 (ASC, BRY); Dancing Rocks NE of Rock Point, 21 Aug 2003. Roth 16W (
3, Roth & Holiday 841; small spring, W
; end of Keet Seel Canyon, 13 Sep 2000,
9,Hevron214b(NAVA);
h ins (NAVA). Navajo Co.: head of Far End Canyon N of Geshi Canyon. 30 Sep
watershed Dowozhiebito Canyon. 2 Oct 2000. Roth & Rink 923 (NAVA); Jackass Canyon. 1
sideofLongCanyon.SkeletonMesa,18May2004.Rinfe&Bungart3146(ASC.SJNM.NAVA). p ,„,„v4ave
n 1340 (NAVA);
2. Carex utahensis Reznicek & D.F. Murray, sp. r
/. Type; UNITED STATES. Utah. San Juan Co.; Angel Arch Canyot
inyon bottom and juniper-pinyon cc
1 slope, ca. 5.500 ft. 2Jun 1964, 5
Loosely cespilose to colonial from elongate rhizomes 1. 3-2.7 mm thick, rhizomes ca. 1-5 cm long between
shoots rhizLe scales brown to purplish-brown, disintegrating into coarse fibers; culms 22-55 cm, stiff,
.erect,' trigonous, papillose, 0.5-1.4 mm wide immediately below the inflomscence; camphylb
and basal sheaths brown, the youngest often reddish-purple. Leaves 6-12, esKutmlly ba« , blades 3^
long. 1.8-4.5 mm wide, much shorter than culms, .folded, margins and midnb smooth to Enely an ror^ly
scabrous distally, ±smooth adaxially, papillose abaxially; leaf sheaths 1.7-15 ^mlong, gla '
ish or pale brown, hyaline, thin, shallowly concave at the apex; hgules ca. 0.8-3.7 mm long .
shorter than wide, free portion whitish to pale brown, inflorescences 2^ cm long,
termlrmlgynaecandrous or occasionally starninatgpUtillm^^^^
t:X»ed.TLinalspikel2-31nimlongpistm^^^
widP ra f0-)L50-flowered staminate portion (0-)6-24.5 mm long, 1.8-3.1 mm wide, ca. (0-)2()-65 How
emd,'p^unchL8-lL5mmiong,Im.eralspikes 4-22.8 mmlong,3.5-6nrm,^e,cyhndnc,^^e^^
with ca 7-40 ascending perigynia. Staminate scales 2.3-3.7 mm long, 1. . mmwi , ’
brown to purple with narrow hyahne margins, especiaflydistally.apex acute to obtuse or rounded,
Lmerved center rarely prolongedmtoa±scabrous-ciliate apiculus or short awn up toO.3 mm long. Pistillate
s 1 9-3 3 mm long 1 3-2.2 mm wide, reddish-brown to reddish-purple, hyaline margins inconspicuous,
• ' ’ - 1 acute, the green 1-3-nerved center sometimes prolonged into a scabrous-cihate
apiculus or awn up to 0.5 mm long. Perigynia 2.6-3.9 mm long, 1.6-2.5 mm wide, ±strongly flattened to con-
cavo-convex m cross-section, ±obovate, not filled by the much smaller achene, herbaceous, papillose distally
green to reddish-purpledistally,greenish to brown proximally,2-nervedand frequently withafew faint nervei
Z ^ obscure to well defined cylindrical beak
teaks01-0.4tnm long, papillose and sometimes setulose. green to purplish-brown, apex *erose to obscurel,
ellinti r sometimes lenticular in cross section; the convex sides narrowly
cm gonousac enes, ™^dfy elliptic in lenticular achenes, brown, smooth apex usually apiculatewitl
%TATtto!3 'T'* ^ -"““y 3. “p “>
%2.A„te3.1.8-2.6mn.lo„g,„ftenwia.aco„spc„o„sapiculmca.0.1-0.3mmbnfr
Reznicek and Murray, Systematics of the Carex parryana group
Etymology.— FromthestateofUtahwherethetypecollectionisfrom. , ^
Seeps, wet slopes, alcoves and hanging gardens, wet meadows, and riparian bottomlands. Elevation
1400-3000 m. Endemic to Utah and adjacent westernmost Colorado. nORfO
PlantstreatedasCpanyana in Utahby Goodrich (in WeUhetal. 2003) and mapped by Albeeaal. (1988)
are this species and also C.holmgreniorum (see below).
Car« uMhcnsis appears to be endemic to the northern portion of the Colorado Plateau Jt ri similar to C.
speemeda in having achenes much smaller than the perigynia, the perigynia thus lonely fitting «ou^^
aeheneandnotfilledby it, Bute, utahensisisarnorestimyerectplant, with darkerpistilbm^
green and clearly obovate perigynia distinctly purple tinged apically. Cnrex specuicoln has a lax, “iri
Sder Distuiate scales and pale greenish, elliptic perigynia changing to whitish and iranslucent at maturity. A
Lphngotpengynia,scales,andachenesarenotedin Figure dCandatypicalinfloreseencefromaherbarium
specimen is shown in Figure 5C.
.rniOHDQ.Mo.ln,«C.iT47N,21)W.S13.L.S.ICi«kCy.l.5n.lEoiUul,-Ce...»a„K'iH„,Slanl994.
T.h,37M(COLO.UTC).OTAH.Ci«l«».Co.:TMSE17E.S17.3Ja„2«Xl,At,<,<»i25i72(Bm
8j™ 1979. WWn&Mooir S3 (BIt«lBigE«aMt.,8A«gl9r7,Le»ii3125(IlRY); San R.MSwdl.5Jun 1979,1
13 Jun 1989, Tuhy 3545; San Rafael Reef, Cottonwood Wash, 6 May 2001, Yeatfs 4573 &. Yeatt
1999, Atwood & Evenden 24445 (BRY). Juab Co.: Mt. Nebo peak trail from Pole Canyon, 17 .
Co ■ M 1 mi upstream from jet. San Juan R. with Slickhom Canyon, 1 jun 2005, Roth 1847 (NAVA
1983, Welsh & Chatterley 22024 (BRY); John’s Canyon bottom, 14 May 2000, ^16 (BRY). I
1900 (BRY). Uintah Co.: Firewater Canyon. Moonshine Canyon.3Jun2000.Atwood25765 (BRY)
niustration.— Mackenzie, K.K. 1940. N. Amer. Caric. 2, pi. 421, as C. abon^num.
Loosely cespitose from short, ascending rhizomes 2-3.5 mm thick, rhizomes ca. 0 5
shoots rhizome scales brown, disintegrating into coarse fibers; culms 20-90 cm tall,
trTlous very finely papillose, 0.6-1.1 mm wide immediately below the inflorescence;
phyllsandbasalsheathsbrown,theyounge3tusuanyreddlsh-purpk.Leavesca.M2^^m^l^^^^^^^^
3 3 48 cm long 1 5-4 mm wide, much shorter than culms, ^folded, margins and midnb mooth to finely an
™;!y^ltdis.aUy,Asm„othadaxial.y.papiUoseabaxiaUy,I^
trally whitish or pale brown, hyaline, thin, concave at the apex; hgules ca. 0.8-5.5(-7.5 mm long, acute to
rounded, slightly shorter than to longer than wide, the freeportionwhitishtopalebrownJn^^^^^^
cm lone with 3-5(-6) spikes, terminal staminate or sometimes gynaecandrous a era s p ,
e7ai4e3-eriappingVersou.e.in.essepa«e.lowe7.wospikesa^^^^
rriAtVi npHiinrle un to 14 2 mm long; lowest bracts ±bristle-like up to 7.5 cm long and 1.5 mm
raLss,up^rbractsabEupdy reduced. Terminal spte(6.5-)11.5-27 mm loug^^^^
10mmlong,4.5-7.2mmwide,ca.(0-)l-28-nowered,3tamina,eporuon4.5-2b3mmlo^ai^M^
ca. 10-65-flowered, peduncle 1.8-38 mm long. Lateral spikes 4 2 mm ong, ' ^ ^ ^ ^ 2 4
denselv flowered with ca 5-45 spreading-ascending perigynia. Staminate scales ' , . f’
mrjM"oL,e,reddI-brown\uhbroadbyaUne margins
"dcen.er.Pis.illa.esca.esl.^2.3mm,ong,1.5-2.2rn^
wwfr:2ied rrigonous ro plumply biconvex, broadly obovare^e— ^
beak; beaks 0 2^.5 mn. long, finely papillose and so.ne.imes .ru^, green ro br^n^x ™
Obscurely bidentulate with teeth up to 0.2 mm long. Achenes L7-2.4 mm long, L3-L6 mm wide, concavely
tapering to a thick stipe 0.1-0.2 mm long, flattened-trigonous in cross section; convex sides narrowly elliptic to
obovate, brown, smooth; apex usually apiculate with an apiculus up to 0.2 mm formed by the persistent style
base; style deciduous; stigmas 3. Anthers 3, 1.7-3 mm long, with an apiculus ca. 0.1 mm long.
Etymology.— Carex holmgreniorum was chosen to honor Arthur H. Holmgren and Noel and Patricia Hol-
mgren, who together have collected this species several times and whose scholarship has contributed so much
to our knowledge of the Intermountain flora.
Alkaline seeps, adjacent to springs, riparian zones, moist meadows, pasture land, calcareous and often
saline soils, with species such as Juncus balticus Willd., Eleocharis rosteUata (Torr.) Torr., and Schoenoplectus
pungens (Vahl) Palla. Elevation 1400-2000 m. UNITED STATES. Idaho, Nevada, Utah.
Some of the Utah localities of C. parryana mapped by Albee et al. (1988) presumably refer to this species.
Publication of Carex parryana var. brevisquama by Hermann (1968) lacked a citation of a type specimen
as was required by the Code. Valid publication did not occur until 1970 when the name was again published by
Hermann with Holmgren 8251 as the type for the name (the holotype, and apparently the only sheet seen by
Hermann based on his annotation on the sheet). The epithet brevisquama was preoccupied at the rank of
This is a distinctive species whose lack of recognition was due to its relative rarity and the confusion sur-
rounding the misapplication of the name C. aboriginum. It is easily recognized by its combination of large pis-
tillate lateral spikes, staminate or gynaecandrous terminal spike, broad achenes and perigynia, and pistillate
scales distinctly shorter than the perigynia. It is an uncommon and local endemic of the northern Great Basin
and western portion ol the Colorado Plateau. A sampling of perigynia, scales, and achenes are ,
4A and a typical inflorescence from an herbarium specimen is shown in Figure 5A.
Repre^ntauve specimens. IDAHO. Caribou Co.: Soda Springs, 9 Jun 1993, Moseley 2620 (BRY). NEVADA. Elko Co.: F
8745 (BRYM^ National Wildlife Headquarters, 20 Jun IS
1 in Figure
^ us,mnom.-Macltenzie. K.K. Im N. Amer. aric. 11: pt 412; CnmquUt, A, etal, 1977. iutermountain Flc
cm talUtiff, erect, trigonous, finely papillosewithltj::”^^^
mm wide immediately below the inflorescence- nhvllonorlir i, n . , ^ antrorsely scabrous, 0.5-1
usually reddish-purple.Uavesca. 6-12, ess^nttallylsaifcSS^'^^^^^^
shorter than culms, .folded, margins and midrib sLth to mXsI L'”
dislally, smooth adaxially, papillose abaxially leaf sheaths 2 n s i ' ‘'"''V umrotscly scabro
brown, hyaUne, thin, cotmave at theapexilJusItS^fl ^"”"*'®^'^^^^
lonpr than wide, the free portion whitish to pale brown iX *'
pistuiate, upper lateral spikes .overlapping bwer som,H™ *‘™"'«e, or rarely .mixed, lateta
wer sometimes separate, lower two spikes 0.3-5.2 cm distar
mm long, pistillate portion (0-)2.1-17.5
(0-)2-24.8 mm long, 1.5-2.8 mm wide.
.0.1-0.2mmlong.
T3S, R9W, Sec. 1 SW *A, 19 Aug 1997, Heidel 1612 (MICH). PowcU Co.: VI mi E of Kleinschmidt Uke, T14N RllW S18, 1 Jul 1993, Lesica 6027
(NY). Teton Co.: North Fork Willow Creek, ca. % mi SE of Duhr House, 25 mi W of Choteau, 23 Jul 1982, Lesica 2273 (NY). Wheatland Co.:
Galloway Creek, ca. 10 mi NE of Harlowton, 26 Jul 1988, Lesica 4693 (NY). WYOMING. Sublette Co.: Jackamore Creek, 21 Jun 1901 , Merrill
6. Wilcox 609 (GH, NY)
5. Carex haUii Olney in Hayden, Rep. U.S. Geol. Surv. Terr. 5:496. 1872 (non L.H. Bailey 1886). Type: UNITED
larbour 617 (holotype; BRUl; isotypes: DUKE fide Weber 1997, n.v., GH!, ISC fide Weber 1997, n.v., NY!,
n. New York Bot. Gard. 1:74. 1900 (nom. superfl. and illegit.); Carex panyana Dewey subsp. hallii (Olney) D.F. Murray,
Illustration.— Mackenzie, K.K. 1940. N. Amer. Caric. 2, pi. 411.
Colonial from ±elongate rhizomes 1.2-2.3 mm thick, rhizomes ca. 0.5-16 cm long between shoots, rhi-
zome scales brown, disintegrating into greyish fibers; culms (6-)9-38 cm, stiff, erect, trigonous, the angles
sometimes antrorsely scabrous, papillose, 0.4-1.1 mm wide immediately below the inflorescence; phyllopodic,
cataphylls and basal sheaths brown, the youngest often reddish-purple tinted. Leaves 4-10, essentially basal;
blades 1.8-18(-22) cm long, 1.6-4.3 mm wide, much shorter than culms, ±folded, margins and midrib smooth
to finely antrorsely scabrous distally, ±smooth adaxially, papillose abaxially; leaf sheaths 11-7.5 cm long, gla-
brous, ventrally whitish or pale brown, hyaline, thin, shallowly concave at the apex; ligules 0.5-2.7 mm long,
obtuse to rounded, shorter than wide, free portion whitish to pale brown. Inflorescences 1. 2-4.5 cm long, with
l-3(-5) spikes, terminal spike staminate (with sometimes l-3(-8) pistillate flowers at base or apex) or pistil-
late, rarely mixed, the laterals pistillate; lateral spikes overlapping or separate, the lower two spikes 0.3-0.7(-
1.8) cm distant, the lowermost sessile or on a papillose peduncle up to 8.2 mm long; lowest bracts ±bristle-like
up to 3.2(-6) cm long, 0.3-0.7(-1.8) mm wide, sheathless or nearly so, upper bracts abruptly reduced. Terminal
spike 12-33 mm long, 20-75-flowered and 3.2-5 mm wide when pistillate, 35-90-flowered and 2.2-3 8 mm
wide when wholly or largely staminate, the peduncle 2-12 mm long (if lateral spikes present). Lateral spikes, if
present, 3-14 mm long, 2.4-3.5 mm wide, cylindric, densely flowered with ca. 3-30 ascending perigynia. Sta-
minate scales 2.6-3.6 mm long, 1.4-2.2 mm wide, ovate, reddish-brown to pale purple with narrow hyaline
margins, especially distally, apex broadly acute to rounded, the green, 1-3-nerved center sometimes prolonged
into a ±short apiculus on the distal scales. Pistillate scales 1.8-3(-3.2) mm long, L4-1.9 mm wide, reddish-
brown to reddish-purple, with broad hyaline margins, ovate to ±oblong, broadly to rounded, tht green
1-3-nerved center sometimes prolonged into a scabrous-ciliate apiculus or broad awn up to 0.5 mm long. Pe-
ngyma L7-2.4 mm long, 0.9-1.8 mm wide, ±flattened-trigonous in cross-section, ±obovate, tightly wrapping
the achene, coriaceous, papillose distally, green to brown, 2-ribbed and frequently with a few faint nerves on
the faces, ta^nng to a short stipitate base, contracted into a short cylindrical beak; beaks 0.2-0.4 mm long,
^pillose and sometimes setulose, green to purplish-brown, apex bidentulate with teeth up to 0.1 mm long.
*■
T ■" “■"'ex sides obovate, brawn, smoolh; apex with a iiny apiculus up io 0.2 mm
formed by ihe persisieni style base; style deciduous; stigmas 3. Anthers 3, (1.5-12-3 6 mm long ^th a short
Moist rrequentlyalkaline,soilsoffens,mcadows,andptairies.Elevation200-3400m CANADA M-
representing a specimen also thought (by Porter?) to be C Lhi Th _ Pleasant Valley,
treatedasthetype,startingwithMatoJl9^^^^^ W specimen has been
also by Mnnay (1969) who followed Mackenzie and noted 2^5^^ “f
Junl821,«.yde„s,„.(Mva2bisshouldbethelecto.yP.i,nob::rra;^^^^^^
Reznicek and Murray, Systematics of the Carex parryana group
here that Olney’s C. hallii was clearly based on Hall & Harbour 617 in Olney’s herbarium (BRU!), which was
citedinside the quotes, and clearly fits the protologue, which calls foraspecimenthat^ resembk^^^^^^
dea when the latter hasasecond small spike.. ."This specimen was most likely collectedmSouth Park, Colo-
rado (Weber 1997). The Hayden specimen from Pleasant Valley is in conflict with the protologue, aving w
substantiallateral spikes, thus notatallhke^..C.sdrpoidea when thelatterhasasecondsma^^
is in fact C. idahoa. Pleasant Valley is located in the northeast corner of Yellowstone Na tonal Park in present
day Park Co., Wyoming, near the Montana Border at 44“55'43"N, U0°25'17"W 0. Revea pers. comm .
Carex hallii exhibits an unusual sex distribution in the innorescences. Termtnal sp*es am usually either
staminate or pistillate, with mmly one or a tew pistillate flowers at the base or apex of the ptedommantly sta-
minate spikes and occasionally a yery lew staminate flowers, often scattered, in the pistillate spikes. However,
lateralspikes,;f present, are alwayspistillate regardless of the sex of the.erminalspike.Th«t^^^^^
be distributed clonally; observations of sevetal populations in Colorado determined that clones
MW r^rtrfmrctlfan^^^^^^ porryanti van Wlii (e.g., Mastrogiuseppe 1993), were redeter-
mined as C. idahoa. See the discussion under C. idahoa.
n rANADA MANITOBA Brandon,24Junl951.Stevensim374(ALA);OakR.atLothair,21Junl906.Macoun&
7 ml Sot Kenned,. 13Jul IMl, Bmvin 6- Dere 7798 (DAO). UNtTED STATES. COt^RADO. «»;«“'■ “ ”
W of Salida. 22 Jun 1926, Erlanson 2020 (MICH). Cl
3332 (ALA, COLO,
132 (MICH, NY). El Paso Co.: F
E of Falconr22 Aug 1946, Livingston 1430 (COLO). Gunnison Co.: (
^ her&r Wittmmn 19051 (COLO). Lake Co.: I
IICH MO NY RM^UTC)' Twin Lakes 4Jul 1919,Cbfee)'3333(ALA,GH,MlCH,MO,NY,RM,US,UTC).
Honmsho'. Pa*. Rock; Ml. Nai. P«k, Itinn IRdl-Ko"- ^
'ollins Natural Area, 25 mi N of Fort Collins just S of Wyoming stat
1873 Wolfe s.n., (MICH, NY); South Park, 10 Aug 1927, Hanson 2687
l.ia3\cOLO);GenevaCreek,T6SR^W9Augl2Gi^^
- - Neely 2998 (COLO); Antero-Salt Creek Natural Area, ,ni SW of Fairplay. 16 Aug 2012, Reznicek
); South Park at Jefferson, 17 Aug 1960, Weber, Porsild & Hoi-
1, Wingate 9208 Regensberg 6- Clark (1
), MICH); E side of Fov
mi S of Fairplay, 23 i
Johnson Gulch tributary of Rock Creek, 18 Aug 2012, Reznicefe 12147 (KHD,
Creek ca 4 5 air mi NW of Grant (jet. US Hwy 285), opposite road from Bur
Summit Co.: Breckenridge, Aug 1901, Mackenzie 382 (NY). NEBRASKA. K
NY), 9 May 1913, (MO); 23 May 1927 (ALA, GH, NY, RM, US); Lexington, 16 J
Leeds, 10 & 19 Jun 1905, Lunell s.n. (MICH, NY). WYOMING: ~
y WTU); along County Road 62, paralleling Geneva
Campground, 18 Aug 2012, Reznicek 12149 (MICH).
>.: Minden, 26 May 1898, Hapeman s.n. (MICH, MO,
laveman s.n. (UTC). NORTH DAKOTA. Benson Co.:
Carbon Co.: Big Creek, 10 Jul
m 412 (MICH). Laramie Co.: Pin
nial, 26 Jul 1900, Nelson 7682 (NY). Platte Co.: Cold Springs, 13 Jul
6. Carex idahoa L.H. Bailey, Bot. Gaz. 21:5-6. 1896. l
niastration.-Mackcnzie, K.K. 1940. N. Amer. Caric. 2, pi. 410. 0 5-10 cm
mar Jitr!d'm1^rrZ^^'
Journal of the Botanical Research Institute of Texas 7(1)
shorter than wide, free portion whitish to pale brown. Culms apparently unisexual (and plants presumably
dioecious, but field observations unavailable); Pistillate inflorescences 1.6-5.3 cm long, with l-3(-4) spikes;
lateral spikes, if present, ±overlapping, the lower two spikes 0.6-1.7 cm distant, the lowermost ±sessile or on a
±papillose and scabrous-angled peduncle up to 8.5 mm long; lowermost bracts scale-like to bristle-like, blades
up to 1.8 cm long and 1.6 mm wide, sheathless, upper bracts abruptly reduced. Terminal spike 12.5-34 mm
long, (4.5-)5.5-9 mm wide, ca. 30-60-flowered. Lateral spikes 5.5-17.5 mm long, ca. 3-5.5 mm wide, cylin-
dric, ca. &-28-flowered. Staminate inflorescences poorly known, ca. 1.6-3.7 mm long, with 1-3 spikes; lateral
spikes, if present, ±overlapping, the lower two spikes ca. 0.4-1.3 cm distant, the lowermost ±sessile; lowermost
bracts scale-like to bristle-like, blades observed up to 1.1 cm long and 0.8 mm wide, sheathless, upper bracts
abruptly reduced. Terminal spike ca. 14-22.5 mm long, ca. 4-4.5 mm wide, ca. 40-60-flowered. Lateral spikes
ca. 4.5-15.5 mm long, ca. 1.5-3 mm wide, cylindric, ca. 8-25-flowered. Staminate scales 3.2-5.1 mm long,
center, the apex obtuse to acute. Pistillate scales (2.1-)2.4-4.6 mm long, 1.2-2 mm wide, dark reddish-brown
to purple-black, with at most narrow hyaline margins, broadly to narrowly ovate to elliptic, obtuse and cuspi-
date to acuminate, with a green to reddish purple 1-nerved center. Perigynia 2.1-2.7 mm long, 1.3-2 mm wide,
±strongly flattened-trigonous in cross-section, broadly elliptic to obovate, ±coriaceous, somewhat papillose
distally, green to brown, often reddish-purple tinged on and near the base of the beak, 2-ribbed, tapering to a
short stipitate base, ±abruptly contracted into a cylindrical beak; beaks 0.2-0.5 mm long, smooth to somewhat
papillose, sometimes slightly setulose, green to purplish-brown, apex ±erose to bidentulate with teeth up to 0.1
mm long. Achenes L5-1.8 mm long, 0.9-L2 mm wide, cuneately tapering to a short stipe ca. 0.1 mm long,
trigonous in cross section; the convex sides obovate, brown, smooth; apex apiculate with an apiculus up to 0. 1
mm formed by the persistent style base; the style deciduous; stigmas 3. Anthers 3, ca. 2 8-4.5 mm long, with a
short, triangular apiculus ca. 0.1 mm long.
Moist margins of seasonally moist calcareotis meadows where the surrounding vegetation is steppe. Ele-
vation 1400-3200 m. UNITED STATES. California, Idaho, Montana, Oregon, Utah, Wyoming.
We have not seen specimens of Carcx idahoa from Oregon, but the photographs in Wilson et al. (2008) are
convincing.
In order to be consistent with the International Code of Botanical Nomenclature, Boivin (1979, p. 87)
considered the correct spelling of the epithet to be idahoana, the adjectival form of Idaho. While idahoana may
be more correct linguistically, the spelling idahoa is not a correctable error covered by the Code, and we retain
Bailey’s original spelling.
Thedi«l„c,lvedarkcolor„f,hescalesinCa,ex,d^
darker than C. halht or C. panyana), and then it may resemble a robust C. haHii in having the lateral spikes
small, with the terminal spike (L2-)L5-3.3 times as long as the longest lateral or more, and also in ha^^^^
Afferent from C T II ^ ^ unispicate. This is
different from C. hallu where 87% of the inflorescences have only one lateral spike or are unispiLte Further-
though dark and longer than the perigynia are also shmter ' l ' ^ ^ P^suHate scale*
typfcal lor Rocky Mounuin plants. More coUmionsandsmdylZSd tetw^^
5, UNITED STATES. CALIFORNIA. Mono Co.: White Mts., Spring 75 m W of the dry bed of Poison Creek 0.25
T mi ENE of Sheet Mt., 7 Aug 2012, Zika et al 26045 (ALA, MICH);
R4WS10,Aug
N of Jackson, 8 Sep 1955,
12490 (MICH); B
nLakeT14SR12WS23, 29Jul 1989, Lesicafr Cooper 4979 (NY);
idoy Mtns, along Nicholia Creek just E of the confluence with
n 1995, Lesica 6762 (MICH); along a small spring 1 mi S of Basin
Hermann 12485 (MICH); W of Upper Red Rock Lake, 10 Sep 1955, Hem
1 miWofLowerHarknessLake,31Jul 1990, Lesica 5208 (MICH); EMoi
Tendoy Mtns, along Sourdough Creek, 9 Jul 1993, Lesica 6066 (MICH]
RockCreek,12Jull993,Lesica6088(MlCH);ca.lmiNofBoxSpring,; u 1 1 1997 Lesica 7484
SS- ^eid of CW^teek^^n^R^^^ Lesica 7495 (NY); along Coyote Creek, Coyote Uke, T15S RllW S7_W At^g 1^7,
lLA), Forks of the Madison, 26 Jul 1897, Rydberg 6- Bessey 3762 (NY),
te Highlands, Moose Creek, 22 Jul 1981, Lackschewitz 9728 (ALA, NY);
Shear 5462 (NY). UTAH. Wasatch Co.: T4S RlOW S6 EV^, 5E of Heber City, Strawberry Valley. Windy Ridge, 1 Jul
1983, Goodrich 18815 (NY). WYOMING. Park Co.: Pleasant Valley. Jun 1871, Hayden s.n. (NY). Teton Co.: Expedition of Capt. W.F. Rayn-
olds to the head waters of the Miss
NR9WS13.8Julll
s, Gros Ventre Pass, 6,500 ft, 18 Jur
ACKNOWLEDGMENTS
We thank the curatore at AlA. ASC, BRY, BRU, COLO, KHD, LCU, MICH, MO, NAVA, NY, RENO, US, UTC
and WIN for the loan of specimens or providing images; we are grateful to Steffi Ickert-Bond, Curator ALA, an
to Zachary Meyers for their assUtaneeinmany ways. Special thanks .oJohnSpen^National^ParkSe^ce, for
initiating this project and providing hnancial support for herbarium study and field work. Damela Roth w^
tremendously helpful showing one of us (AAR) sites for Curex specuicolu tn At^ona and Utah Leo Bnteder fe
Janet Wingate and Loraine Yeatts helped with field sites and specimens in Colorado. Peter Ztka sent matertal
of and commentary on Carex idahoa from California. We are grateful to Bruce Ford and Peter Zika for their very
helpful reviews.
REFERENCES
s of the vascular plants of Utah. Utah. Mus. Nat. Hist. Occ. Publ. f
/mces. Phytologia 43:1-106.
1872 Preliminary report of the United States Geological Survey of Montana and portions of adjacent tt
ogress. Washington, DC.
in Carices. Rhodora 70:419-421.
ritories; being
Hermann, FJ. 1968. Notes on Rocky
Hermann, FJ.
Mackenzie, K.K. 1935. Cyperaceae - Cariceae.
Mastrogiuseppe, J. 1993. Carex, sedge. In: Hie
Murray' 0^^969^3^x000^ of Carex sect Atratae (Cyperaceae) in the southern Rocky Mountains. Brittonia 21 :55-76.
M^rZ, DI. 2002. Corersert^i^n Racemosae. In: Flora of North America Editorial Committee, eds.. Flora of North America
Nnrth nf Mpiiiro vol 23 Oxford Univ. Press, New York. Pp. 401-414.
W.rm7. ofColorado botany: Charles Christopher 823-1 890. University Press of Colorado, N.wot.
Weish,S.L,N.D. Atwood, S. Goodrich, A
Wilson, B.L, R. Brainerd, D. Lytjen, a. newhouse, anu
Oregon State University Press, Corvallis, Oregon.
Zika, P.F., A.L Hipp, and J. Mastrogiuseppe. 2012. Cc
IS (eds.). 2003. A Utah flora. 3rd edition revised. Brigham Young L
Otbng. 2008. Field guide to the sedges o
B Pacific Northwest
n: Baldwin, B.a, D.H. Goldman, DJ. Keil, R. Patterson, TJ. Rosatti, and
scular plants of California, second edition. University of California Press,
Berkeley, CA.Pp. 1308-1339.
Journal of the Botanical Research Institute of Texas 7(1)
BOOK REVIEW
Carmine Stahl and Ria McElvaney. 2012. Trees of Texas: An Easy Guide to Leaf Identification. 2nd Edition.
(ISBN: 978-1-60344-515-3, flexbound). Texas A&M University Press, John H. Lindsey Building, Lewis
Street, 4354 TAMU, College Station, Texas 77845-4354, U.S.A. (Orders: www.tamupress.com, 1-800-
826-8911). $24.95, 338 pp., 18 color photos, 270 b/w photos, map, bib., index, 8 x 11".
detail is lost including fine distinctions of color in leaves, flowers, and bark. Presumably, this choice was made
to keep costs in line, but the lack of color does significantly lower the value of the illustrations for identification
purposes.
The single greatest lack, however, are the fine details that aid in distinguishing among trees with broadly
similar leaves (i.e., leaf margins, glands, and venation patterns, among others). Leaf surface characteristics
such as waxy cuticles, hairs, or scales are also missing from the illustrations. A simple close-up of key ancillary
characteristics would have significantly improved the certainty of identifications made with this guide. Like-
wise, floral and fruit characteristics are presented haphazardly. Some illustrations include fruits, some include
flowers, and some have neither. While these details may not be strictly necessary for the identification of a tree
species, they are an essential part of knowing and understanding what makes a species unique — and what
each species has in common with its relatives.
The best guidebooks intended for amateur audiences offer both user-friendly means of identification
along with a path to a deeper understanding of the philosophy underpinning how and why species are named
and classified as they are. There is a reason, after all, why the foliar characteristics used in Trees of Texas are not
used in determining plant families. That said. Trees of Texas does achieve its goak in making tree identification
much easier while providing interesting and useful tidbits of historical and ethnobotanical information.
— Brian Writte, Research Associate, Botanical Research Institute of Texas, Fort Worth, Texas, U.S.A.
LBrt. Res. tnst Taw 7(1): 54.2013
NEW COMBINATIONS IN CAREX KELLOGGII (CYPERACEAE)
Barbara L Wilson, Richard E. Brainerd, and NickOtting
Carex Working Group
1 377 NW Alta Vista Drive
Corvallis, Oregon USA. 97330
Author for correspondence: bwilson@peak.org
ABSTRACT
RESUMEN
INTRODUCTION
Recent molecular phylogenetic work on Carex lenticularis Michx., C. aquatilis L., and related species (Dragon
& Barrington 2009) resulted in the recommendation that western species of the Carex lenticularis complex are
sufficiently distinct from eastern C. lenticularis to warrant recognition at the species level, as C. kelloggii W.
Boott. Although the three varieties of this western taxon are not differentiated at the DNA sequences used dur-
ing this study, they can be distinguished morphologically (Standley 1983; Dragon & Barrington 2009). How-
ever, two of the taxa have not been published as varieties of C. kelloggii. In preparation for publishing a second
edition of our field guide to sedges of Oregon and Washington (Wilson et al. 2008), we are publishing those
Carex keUoggii W Boott var. impressa (L.H. Bailey) B.L. Wilson & N. Otting, comb, n
var. impressa L.H. Bailey. Mem. Tc
Syst. Bot. Monogr. 7:65. 1985. Type: U.S.A. C
1 (Holm) B.L. Wilson & R.E. Brainerd, c
9. Type: U.S.A. St. Paul Island: Bering Sea. Macoun 1661
ACKNOWLEDGMENTS
We thank Kenton L. Chambers for promptly and thoroughly reviewing this paper.
REFERENCES
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J.W. Thompson. 1 969. Vascular plants of the Pacific Northwest part 1 : Vascular
cryptogams, gymnosperms, and monocotyledons. Univ. Washington Publ. Biol. 17. University of Washington Press,
Dragon, J.A., and D.S. Barrington. 2009. Systematics of the Carex aquatilis and C. lenticularis lineages: geographically and
ecologically divergent sister clades of Carex section Phacocystis (Cyperaceae). Amer. J. Bot. 96:1 986-1 906.
Standley, L.A. 1985. Systematics of the Acutae group of Carex (Cyperaceae) in the Pacific Northwest. Syst. Bot. Monogr.
7:1-106.
Wilson, B.L., R. Brainerd, D. Lytjen, B. Newhouse, and N. Otting. 2008. Field guide to the sedges of the Pacific Northwest.
Oregon State University Press, Corvallis.
J. Bot. Res. Inst Texas 7(1): 53. 2013
Journal of the Botanical Research Institute of Texas 7(1)
BOOK REVIEW
Carmine Stahl and Ria McElvaney. 2012. Trees of Texas: An Easy Guide to Leaf Identification. 2nd Edition.
(ISBN: 978-1-60344-515-3, flexhound). Texas A&M University Press, John H. Lindsey Building, Lewis
Street, 4354 TAMU, College Station, Texas 77845-4354, U.S.A. (Orders: www.tamupress.com, 1-800-
826-8911). $24.95, 338 pp., 18 color photos, 270 b/wphotos, map, bib., index, 8 i/i" x 11".
(continued Jnmt p. 52)
detail is lost including fine distinctions of color in leaves, flowers, and bark. Presumably, this choice was made
to keep costs in line, but the lack of color does significantly lower the value of the illustrations for identification
purposes.
The single greatest lack, however, are the fine details that aid in distinguishing among trees with broadly
similar leaves (i.e., leaf margins, glands, and venation patterns, among others). Leaf surface characteristics
such as waxy cuticles, hairs, or scales are also missing from the illustrations. A simple close-up of key ancillary
characteristics would have significantly improved the certainty of identifications made with this guide. Like-
wise, floral and fruit characteristics are presented haphazardly. Some illustrations include fruits, some include
flowers, and some have neither. While these details may not be strictly necessary for the identification of a tree
species, they are an essential part of knowing and understanding what makes a species unique— and what
each species has in common with its relatives.
The best guidebooks intended for amateur audiences offer both user-friendly means of identification
along with a path to a deeper understanding of the philosophy underpinning how and why species are named
and classified as they are. There is a reason, after all, why the foliar characteristics used in Trees of Texas are not
used in determining plant families. That said. Trees of Texas does achieve its goals in making tree identification
much easier while providing interesting and useful tidbits of historical and ethnobotanical information.
—Brian Witte, Research Associate, Botanical Research Institute of Texas, Fort Worth, Texas, U.S.A.
J.Bot Res. Inst. Texas 7(1): 54.2013
A REVISION OF CORDIA SECTION GERASCANTHUS
(BORAGINALES; CORDIACEAE)
James S. Miller
The New York Botanical Garden
Bronx, New York 10458, U.S.A.
ABSTRACT
RESUMEN
INTRODUCTION
Cordia has historically been defined in a broad sense (e.g. Johnston 1930; Millet 2001) with estimates lot the
number of species ranging from 230 (Airy Shaw 1973) to 350 (Miller 2001). However, in the last two decade^
a number of advances in our understanding of its relationships with other related genera, relationships withm
the genus, and floristic studies of the constituent species have added considerably to what we unde«a.rf.
Several phylogenetic studies have shown that Boraginaceae, which have histoncaUy been considered to com-
prise four ^ distinct and morphologically ddferent subfamUies: Cordioideae, Ehrettotdeae, HeltorioptoA-
L, Boraginolae (lohnston 1931), were more complicated (Gottschling & Hil^r 2001; Gottschltng et al.
2005- Moore & Janse 2006; Nazaire Sr Hufford 2012; Weigend et al. submitted). Several other famthes, or
enigmatic genera, including Hydrophyllaceae, Lennoaceae, and the African genera Codon Hoplesligmu, and
WeLdtiawereallembeddedwithintheBoraginaceae,butalloftheoriginalfoursublamil«wereprovento
be monophyletic, altbough a few genera had historically been placed in the (Gottschltng et al.
2005). It is now clear that Sacccllitm and Coldcnin. both long considered allied with Ehreha, are more closely
related to Cordio, the former included within it and the later sister to Cordiu and Vorronm (Gmtschhng et ab
2005). This has lead to the recognhion of a broadly defined Boraginales (Gottschltng et al. 2W5), mcludmg
each of the four tmditional subfamilies elevated to familial status, Cordiaceae Ehretiaceae, Hehoriopuceae,
and Boraginaceae, and also inclusion of the Hydrophyllaceae, Lennoaceae, Hoplestigmataceae, Wellstedttace-
ae. and newly described Codonaceae (Weigend &Hilger 2010). . . w M
For the Cordiaceae, the studies of Gottschling et al. (2005) elucidated that the Old World prccumto
annual herb Coldeniu procumbens was sister to all of the other woody members of Cordiaceae and then CorAu
section Varronia was sister to all of the rest of Cordia and the three small genera that had been recognized be-
cause of unique, unusual fruits, which all have large accrescent calyces (Auxe^ Miers, Putag^ulu L , and
SuccelliuM Bonpl.), were all embedded in Coniia and allied with the species of Coirfio section Cordia. Based on
this evidence. Miller and Gottschling (2007) segregated Varronia and restricted Cordia to the membem of sec-
lions Cordia, Gerascanthus, Myxa, and Superbiflorae, though the molecular data clearly show that the Old
World and New World species of section Myxa are distinct clades, even though they appear morphologically
similar. Now that checklists are being compiled for the Online World Flora, there appear to be approximately
230 known species of Cordia and an additional 124 species of Varronia, though there are still clearly dozens of
undescribed species of both genera awaiting description from undetermined folders in major herbarium col-
lections.
Cordia section Gerascanthus has always been a difficult group (see quotes in Wheeler 1942; Johnston
1950). It is morphologically distinct and easily recognizable as a group of species, characterized primarily by
its unusual fruits, which are ellipsoidal, single-seeded, fibrous-walled, and most distinctively surrounded and
completely enclosed in the persistent calyx and marsescent corolla, with the corolla drying, turning brown
and apparently assisting like a small parachute with wind dispersal of the mature fruit. But while the section
may be easily recognized, the species have long been difficult to tell apart, prompting Chodat and Vischer
(1920) to write “We are, in fact, deahng with species which are but feebly defined morphologically and the
taxonomy of which will require revision from time to time as observations in the field increase in number.” The
species of section Gerascanthus are not distributed evenly, and though they occur throughout the Neotropics,
most of the South American species were known from few collections or were discovered relatively recently,
and the greatest concentration of species is in western Mexico, a group provided with a relatively solid revision
(Johnston 1950), though Johnston knew eight of his twelve species from fewer than 3 collections, four from the
type only.
Most species of Cordia sect. Gerascanthus are small to medium-sized trees. Only a few species are truly large.
Cordia megalontha is the largest, occasionally reaching 60 m tall, and C. alliodora and C. trichotoma often reach
30 m. Cordia insignis is a shrub of Brazilian cerrado that apparently grows from an underground xylopodium
and flowering individuals are generally only 1-5 m tall and at least in some situations, its above ground stems
appear to be annual.
Descriptions of bark are generally not included with most specimen label data, though bark is often a
characteristic feature. Most species have smooth pale gray bark. Cordia morelosana has dark gray, rough and
deeply fissured bark, which is one character that distinguishes it from C. sonorae, a species i
cally similar and difficult to distinguish from herbarium specimens but has smooth, pale gray bark.
Species of Cordia are not infrequently associated with ants but the association is generally casual, with
ants living under patches of bark or in hollowed twigs. There are, however, two true myrmecophilous species
of Cordia, which are unrelated and independently derived (Gottschling, 2005). Cordia nodosa of sect Myxa,
hasdomatia that are swollen, hollow petioles. Cordia alliodora, of sect. Gerascanthus, has domatia that are hol-
low, ell^td sweUings at the ends of branches and the bases of inflorescences that are generally inhabited by
small, bttmg ants of more than a dozen genera (Wheeler 1942). These swellings are filled with a loose pithy
material that is presumably easily removed by the ants. The domatia are characteristic of the plants and not
the .s„UdWesta.ion^theamsasdo™a.Ufon„nonnally on plants grownm, he gree^
absence of the ams.Anj5 have been associated wi, hall of the treesihave observed in Mexico, but they have
frequently been absent from trees in Central America.
Tnchomes of various types are often associated with E
in the genus
n vanous species of section Gerascanthus; the fifth type, echii
hairs, are known only in Cordia cymosa of section Myxa. The four types of hairs found in section Gerascanthus
include:
1) Simple hairs.— These are unicellular hairs that are found in the majority of the species and are variable in
2) SimplThl^lT/rom are thick-walled, unicellular hairs that arise from a swollen,
basal pedestal or cystolith. The shaft of these hairs is usually postulate and the hairs are mostly ap-
pressed giving rise to strigose, strigillose, and scabrous induments, depending on the length and
thicknes of the hairs. The pedestal or cystolith generally appears multicellular. In some cases, the shaft
of the hairs weathers and is lost with time, leaving the persistent cystoliths, which result in a rough sur-
face. Hairs from pedestals are common in the genus, but less common in section Gerascanthus.
3) Stellate hairs.-These are multiply-branched, star-shaped hairs that are characteristic of Cordia alliodora and
C. trichotoma in section Gerascanthus. Both of these species have stellate hairs with numerous branches
and the hairs are sessile. These differ from the stellate hairs in the related genus Varronia, which are
stalked and obviously have arisen independently.
4) Malpighiaceoushairs.-These are unicellular, T-shapedhairs with two arms andamedianattachment. They
are rare in Cordia but have arisen independently at least four times in the family, each time with a slight-
ly different appearance. They are known only from Cordia glabrata, among the species of section Ger-
ascanthus, where they give rise to a whitish cast to the undersurface of the leaves.
tL leaves of species of Cordia section Gerascanthus are simple, alternate, entire, and are presumed to be de-
ciduous, although this is not confirmed in all species. Many species flower without their leaves, so it is in some
cases difficuU to paiinoweringspedmens with sterile specimens with leaves. Leaves^ryconsiderablemsize
and form and leaf shape and texture are useful characters for distingnishingspecies-Allspecieshavepenolate
leaves,and the petioles are generally canaliculate on the adaxial surface, thoughafewspeciesare only flattened
and lack a nrominent groove.
While Boraginales are thoughtofascharacterized by helicoid or scorpioid inflorescences, that condtttonBap-
parent only occasionally in Conhu and then generally only in the ultimate branches of inflorescences, and tt ts
seldom apparent in species of section Gerascanthus. Inflorescences in the section vary from expanded cymes
and panicles to compressed and nearly umbellate. All of these are basically cymose-paniculate, but they vary
in the degree of elongation of the branches giving rise to a continuum of variatton between three different
imLulateinforescencesare expanded withadistmct central axisthatisapanicleof small cymes, suchasm
Cordia macrantha. . . r l j •
2) Cymose mnotescences are also expanded, but lack a central axis and broader tn form, such as ts found
CordiacolimensisandC.elaeagnoides. , • ■ ■ .
compressed with none of the branches elongating and giving rise to a
3) Umbellate inflorescences are very compress
nearly umbellatecluster of flowers, in Cordia glohuh/em these are not true umbeU but ratheramere con-
traction of the branches of a branched inflorescence, but in C. umheltea they appear to be truly umbel-
late with all flowers arising from a single point.
While inflorescence type is generally consistent lor each species, Cordia sonoroe has i^orescences that vary
S,^^olCordiasect.Getnscanthusaretypicallydistyh,usandmdividualpUntscanread%bese^ratedmm
long-style and short-style morphs, with respective stamenand style heights varyingsignificanUy between the
two. When species are known from adequate numbers of collections, floral measurements are provided sepa-
rately for the two morphs, though this unfortunately is not the case for the majority of species. In two species,
distyly has broken down. In Cordia alliodora, the long-style morph has been lost and all individuals have ex-
serted stamens and a short style. In Cordia trichotoma an unusual variation occurs, with both long-style and
short-style individuals being present in populations but with a third morph with equal stamen and style
heights (Gibbs & Taroda 1983).
The species of Cordia sect. Gerascanthus have a tubular calyx that is usually distinctly 10-ribbed. In a few
species, such as Cordia morelosana, the indument on the calyx is dense and long enough to obscure the ribs. In
other species, such as Cordia thaisiana, the calyx is merely striate and lacks the clearly defined ribs characteris-
tic of most species. The calyx persists in all species and encloses the mature fruit
Section Gerascanthus is characterized by corollas that are marcescent and persist, enclosing the mature
runs. As f^its mature, the corollas dry,tum brown, and act as small parachutes that aid in wind dispersal of
.he ta^Corolteofspedesof the section are ei, her tubuUrwith spreading lobes
^.al y 5 corolla lobes, tftough c^ollas with 4 or 6 lobes are not nncomnton in some spedes. Whether the
corolla l,^s are oblong, wtth parallel sides, or ovate to debate is a feature useful in distinguishing species The
a^ C ^ »>• ^nd only in Cordia ntegalantha, C. latiloba,
and C. umbellifera are the corolla lobes drawn to an acute, sharp apex
characurfordistinguislungspeciesandvar^bleonlyinCurdiagerascanthLandCm^^^^^
The gynoecturn of Cordw consists of a superior, bicaipellate ovary that U no. divided or lobed Earlv in
development tt IS btlocular, but it later becomes falsely 4.|ocular as a septum forms dividi t.
two separate chambers (Uunence 1937- Khaleel 1975 1987^1 fTT , *"*"*'"*
thotropus ovule but onlyonedevelopsmmaml aid fXonTa
species of the section have an annuL needle H- T ? A"
compared with the nectaries of species of mto le cto" 0^7 n ‘“^8'
grocilipesischaracterizedbyhavingadiscthatisciliatealoneth
and twice bifid with 4 separate stigma lobes that vary from fil f “PP*"^ ““Sin The style in Cor(ka is terminal
in section Cerascantims. ^ «‘l«oid but are always clavate
Fruits
The fruits in species of section Gerascanthus are thp a-
thin, fibrous wall, lacking a mesocarp, and are capped bv "
slightly during development and remains borne within the ners 7'™i
™|.dispe.sedwi.h.hemarcesce„.corona,aidinginwinr:;^^^^
PollenoftheBoiaginalesisknown.obediver!e(Avetissianl95fil m i,
thrm types of polfen grains in the genus Cmdia in a broad 7 <*’73) initm
and Gottschling Q007), Varronia and Cordia section Cordta
^mn C^o^ontlms shares pollen gmins that are ^0.^12' 7 7““' ’”'‘™
Tarodaand Gibbs(l986)groupedthese three section7nrr7 ^
taxonomic treatment
• Browne) G. Don, Gen. Hist.
380. 1837; A. DC, Prodr. 9:471 If
Geraxanthus Schlect & Cham., Linna
Miller, Revision of Cordia section Gerascanthus
Cordia section Cerdan
a{Ruiz&Pavon)Oken.
Small to large trees. Leaves deciduous, alternate, petiolate. Infloresce
sometimes with the axes much reduced and nearly umbellate. Flowers
dia alliodora: calyx tubular, usually distinctly 10-ribbed or occasionally merely striate; corolla whii
cent, tubular with spreading lobes or funnelform, usually 5-merous, the lobes usually spreading; si
same number as the corolla lobes, the filaments pubesc
an evident, usually annular, disc. Fruits borne within
soid, the walls thin and fibrous, capped by the discoid base of the style.
Cordia section Gerascanthus consists of 23 species, all restricted to the Neotropics. Thirteen species occur
in Mexico, and though the species are spread throughout Central America, the West Indies, and South Ameri-
ca, there is no other substantial center of diversity.
Journal of the Botanical Research Institute of Texas 7(1)
^^3-4):396. 1988. Type; PERU. HuAnuco: Pn^nr^ p.h, . . ,
.r«« ccraam Koem. & Schult., Syst. Veg. 4:467. 1819.
non. August, Hassler 7155 (holotype: G). '"*' in campis siccis, prope Concep-
S.TYPEiPERUrPovOns.)
Cordia alliodora var. glabra A. DC.. Prodr. 9:472. 11
Tree to 20(-25) m tall, slender cr b
f white, (8.5-)9.5-12(-141 mm i , attenuate teeth, these 0.5-K-l ^^(-12)-ftbbed,
(3.5-)4.5-6(-8 5) mm 1 ^ ^bese 5-7(_8 5) mm 1 *” marcescent,
Gerascmtims rang-
»tn.lAmencaa«l,heWes.lndies.,„B,BzU,
Miller, Revision of Cordia section Gerascanthus 61
Bolivia, and Argentina. It is replaced in Eastern Brazil by its closest relative Cordia trichotoma. It occurs across
a wide vareity of habitats, from dry to wet forests and ranges from sea level to 1,500 m or rarely as high as 1,900
lev. 30 ft, Whitfoord2866 (MO). Orange Walk: 1
5407 (BRH, MO, SEL). BOLIVIA. Beni: Prov. V
17557 (MO).
i: Cochabamba, Bang 1178 (MO, NY, US). La Paz. S. Y
samo 1224 (MO).
Krufeoiifs.n.(MO.NY).Pai
Hierba Buena, elev. 1,300
lo. W bank of Rio Madeira, 2 km N of Abuna, Prance et al. 6114 (MO, NY, I
, Steinbach 270 (MO). Tarija: 0.2-0.4 km NE of the bridge over Rio Pilcoi
road. 5.5 km (by road) W of ce
, Nee 53257 (MO. NY). BRAZIL. Acre: near mouth of Rio Embira (Tribuui
uth of Rio Embira (Tributary of Rio Tarauaca), Krukojf 4788 (MO, NY, U5
n 22 da Rodovia llheus^Ttabuna (BR 415), Jardim et al. 1102 (NY). Ceara: without localit
I, Heringer et al. 7183 (NY). Goias: Serra do Caiapd, ca. 5 km (straight line) S
a, 5 km de Imperatriz, Fires 16117 (MO). Mato Gross
iL 16668 (NY, US). Para: Upper Cupory River, plateau betwet
;a Valley, 15 km NE of Bolivar, Gentry & Renteria 23690 (MO). Bolivan Norosi-Tiquisio trail. Lands of Lova, elev. 1500-600 m.
Journal of the Botanical Research Institute of Texas 7{1)
n,Stehle603I(
600 ft, Proctor 18900 (US). St: Vincent: Smith 1249 (GH, NY). MEXICO. Campeche: A 18 km al S de Santa Maria Xcabak aproximadamente
road to Mai Palo, elev. 800 m, Breedlove & Thome 21019 (CHAPA, DS, F, LL, MEXU, MICH, MO NY, RSA). CoUma: 20-30 mi NE of Colima
iologia de Chamela, Perez 767 (ENCB, MEXU, ^
i. Rzedowski 17976 (ENCB, MEXU). Oaxaca: Mpio. of Chiltep
1. 11734 (ARIZ, F, GH, K, MO, N
Quinta
brera 1688 (MO). San Luis Potosi: Mpio. de Xilitla, El Jobo, elev. 200 m. Rzedowski 10151 (DS, ENCB, MEXU, SLPM, TEX, WIS). Sinaloa:
Imala, Gentry 4946 (ARIZ. F, MEXU, t
Mpio. Gomez Farias, 4-5 km al N W de Gomez Farias, «
MO). Yucatan: Lolon, Flores 4 (F). NICARAGUA. Atlantico Norte: E of Finca Waylawas, 2 km from base of Cerro Waylawas, elev. 90 m.
Pipoly 4346 (MO). Atlantico Sur: El Zapote, 40 km NE de Nueva Guinea, elev. 130-150 m, Sandino 4834 (MO). Boaco: Camino a Mom-
etepe, al lado N del Volcan Concepcidn, cafetales do las Angeles y sus alrededores, elev. 250-350 m, Sandino 536 (MO). Zelaya: vicinity of La
Luz-Siuna, elev. 150-200 m. Bunting & Ucht 657 (F, UC, WIS). PANAMA. Bocas del Toro: vicinity of Chiriqul Lagoon, Old Bank Island,
Wedel 1877 (MO, US). Canal Area: Barro Colorado Island, Croat 7694 (MO). Chiriqui: vicinity of San Felix, elev. 0-120 m. Pittier5279 (US).
i. Gentry 4349 (MO). Herrera: vi
Panama: San Jo
etal. 154 (AAU,
Island, Johnston 570 (GH). S
6 km SW of Aguadulce, elev. 0-50 m, Knapp et al. 3348 (MO). PARAGUAY. Alto Parana: Estancial Rio Bonito, Zardini &
i. Smith etal. 1682 (MO). P
Kayap 806 {MO). Cajam
a, Soukup 3839 (US). C
»: 1 km N of Pilcopata, edge of Rio Madre de Dios, e
Tlooking Rio Huallaga, elev. 350 m, Gentry et al. 37716 (MO). PUERTO RICO.
Adjnntas: Base of Pico Guilarte, 100 m, Liogier 10015 (NY, US). Agnada: On road from Rincon to Aguada, Vives 2999 (UPR). Aguadillas:
Aguadillas, Sargent 573 (US). Aibonita: Inter Aibonita et Cari-Blanco, Sintenis 2946 (US). Cabo Rojo: Cabo Rojo in Monte Grande, Sintenis
709 (MO. US). Caye; Caye, Hess & Stevens 2924 (NY). Ceiba: near Ceiba, Wagner 692 (A). Fajardo: near westernmost radio tower just S of the
Route3/Route 985 comer, elev. 10 m, Taylor 9133 (UPRRP). Isabela: Bosque Insular de Guajataca, Little 13516 (A, NY, UPR, US). Jayna: Ja-
)^a. Sargent 3050 (US). Juncos: Juncos, in Monte Goyo, sintenis 2038 (US). Manati: Km 26, Manati-Villalba highway. Little 13712 (US); Na-
V 465-720 m, Shafer 3476 (US). Penuelas: Penuelas, Sintenis 4883 (MO, U
; Coastal hill 2 mi. N of Mameyes (Palmar), elev. 50 m. Hartley 13376 (A, U
outes 162 and 717, Taylor 6432 (UPRRP). San Juan: I
e, Gregory 527 (UPR). Vega B
Journal of the Botanical Research Institute of Texas 7(1)
16342 (MO). Dis-
lidadde
il SW de los tanque de la Electricidad de Caracas (Cocuizal), elev. 900-1,100 m, Steyermark 91829 (US). Guarico:
15 km SW of Tamaco along main highway between Altagracia de Orituco and Chaguaramus, elev. 150 m, Davidse 4218 (MO). Maracaibo:
uatro Bocas, Aristeguieta et al. 6824 (US). Merida: 30 km SW of Mriida, along road to El
irais the most widespread species of section Gerascanthus, and it is very distinctive in its evident
Metepubescenceontostems, leaves, inBorescem^ebranches, and calyxsott
slenlellhaslhe smallest Dowers ot the seclion with corollas less than 14 mm long. Itis most closely related to
an most easi y confused with Coidlu trtchotmm, hut differs in having smaller Dowers, (8.5-)9 5-12(-14) mm
rT” i """I' >■“ 4nt domatia, which C. Iricholoma does not
Q^ndla^
■S^uuT:^ ?
7. (Fig. 2). Type: GUATEMALA, i
inn. Missouri Bot. Gard. 74(3):672-673, f. 2. 1
, Contreras 6835 (holotype: LL; isotypes: DS, F. LL US)
ersdlstylotm, on short spurs to L5 rmTongXTubuLT5‘!^t‘'T‘"T^r''^'^^^
evenly lobed and tearing upon dehiscence or dehMng circumscis^h sfriate o 7” 7“^'
Hremlent; corolla white, tubular with somewhat sprLing lobes 14 2 16 8 m T
oblong, 5.2-6.2 mm long, 4.8-5.9 mm wide the tube 4 7-5 1 mm I
long, the upper 7-7.6 mm free, glabrous, the anthers oblnna 9 t 9.8-11.5 mm
1.3- 2.5 mm broad; disc anular, 1.4 mm tall 1 3 mm hrni,4 k ^
1.4- 1.6 mm long, the stigma lobes clavate. Fruits unknown^ ^
occurrs^a7;d'::!:re:;™^=’^“"^"'-"»”'^'^^
ntlated to. and most easily" i^nhld
A 31:186. 1950. (F» 3),<
iving a calyx that is shorter
«(l.M.Johnst.)Borhidi.
nal, compressed cymose-panicu-
late, to 6 cm long, 8.5 cm broad, 11-30 or more flowered, the branches of the inflorescence tomentose, the hairs
wavy, short, dark brown. Flowers distylous, nearly sessile at the tips of inflorescence branches; calyx tubular,
12-15 mm long, 3-3.5 mm wide at the mouth, usually 5-lobed, sometimes with fewer lobes, the lobes un-
evenly shallowly triangular to deflate, 10-ribbed, each rib with a central furrow making the calyx appear
falsely 20-ribbed, glabrous; corolla marcescent, white, funnelform, 2.8-3 cm long, 5-lobed the lobes very
widely ovate to transversely widely oblong, 7-10 mm long, 9-11 mm wide, the tube 10-16 mm long, glabrous;
stamens 5, the filaments 21-24 mm long, the upper 10-15 mm free, pubescent at the point of insertion, the
anthers oblong, 2.5-3 mm long; ovary broadly obovoid, 1-2.5 mm long, 1-1.3 mm broad, glabrous; style 14-23
mm long, the stylar branches ca. 4 mm long, the stigma lobes clavate. Fruits unknown.
Distribution and habitat. — This species is known from only three collections from western Mexico where
it occurs in coastal parts of Colima and Jalisco from sea level to 550 m in dry forests.
Specimens examined MEXICO. Jalisco: ills between Bahia Navidad and La Manzanilla on Bahia Tenacaiita, E facing summits 3 mi. W of
the Autlan-Navidad highway, elev. 550 m, McVaugh 21017 (MICH); 1 km al N de La Manzanilla, sobre el camino a Tenacatita, Rzedowski
14840 (ENCB, MEXU, MICH, TEX).
L. DC., Prodr. 9:474. 1845. (Fig. 3). L
1891. Type: MEXICO. Oaxaca; Chjuitan inter Tehuantepec et Roca del Monte
rdia exsucca SessO & Mocino, H. Mexic, ed. 2, 45-46. 1894. Type: MEXICO: h
Journal of the Botanical Research Institute of Texas 7(1)
Tree to l5(-20) m tall, the bark furrowed, light brown, young twigs canescent ■ ■ u
3.3-.o(-a.3rr::™;htX"r^^^
and abn.p.ly decurrea. along .he pe.ii but
surfaceglabn.scen..ora.elyhmelIouMhehairsfromafU.pedel«r«rr;
•he budsapicutae; calyx urceolale to campanuUMS T'"""’
unmnly Ua-)13(-18) ribbed, unevenly 5-lobed the lobl!^ f ””"u *■
mm long, appressed; corolla marsescent white to cream fu ^ canescent, the hairs 0.2-0
^^';^*f73<-36)n.n.l„ng,.^esri^bte^™I“^“'”“““”«'>''et<hscato^^^^
■eugnunfcs u. a reasonably conuuon species in dry forest of wester
1 be found from sea level to 1,200 r
110 toward Manzanillo, Miller et al. 273 (MEXU, MO). Gueircro; along the road between Filo de Caballo and Milpillas, just W of Xochipala.
tween U Huerta and Km 231, elev. 500 m. Miller et all 388 (MEXU, MO). Mexico. Calera, dist. Temascaltepec, elev. 790 m, Hinton 5378 (G,
elev. 370 m, Miller & Tenorio 657 (MEXU, MO). Oaxaca: Km 12 de la carreteraXehuantepec al S de Salina Cruz, elev. 25 m. Cedilla Trigos &
Lorence 456 (MO). Quintana Roo: en el Palmar, a 7 km al sur de Ucum, sobre la carretera a U Unibn, cerca del Rip Hondo, Cabrera et al. 2636
(MO). Sinaloa: central plaza of Mocorito, Breedlove 16724 (CAS).
Cordia elaeagnoides is a common tree on the Pacific slope of western Mexico and is dominant in some of the
places where it occurs. Originally considered sectionally distinct and placed in section Rhabdocalyx by DeCan-
dolle (1845) because of its hard, bony-walled fruits, recent molecular studies show that it is definitely related to
the other species of section Gerascanthus (Gottschling et al. 2005). It is distinct in its cymose inflorescences and
hard, bony-walled fruits. Although this species seldom excedes 15 m in height, it is highly valued for its wood
in regions where it occurs. In western Mexico, it is one of the most commonly used species for the construction
of furniture. Common names include “Grisino” (Chiapas), “bocote,” “gueramo” (Michoacan, Guerrero), “oco-
tillo meco” (Oaxaca) and “barsino” (Guerrero, Michoacan) (Standley 1924).
i., Repert. Spec. Nov. Regni Veg. 12(322-324):240. 1913. Type: MEXICO. Gui
Tree to 15(-30) m tall, to 30 cm dbh, the bark light gray, smooth, the twigs glabrous. Leaves deciduous; petioles
(8-)12-30(-40) cm long, canaliculate on the adaxial surface, glabrous; blades elliptic oblong to lance-ovate,
(4.5-)6-15(-20) cm long, (2-)2.6-6(-^.5) cm wide, the apex acuminate or acute, the base acute to nearly ob-
tuse, the margin entire, the adaxial surface glabrous, the abaxial surface glabrous. Inflorescences terminal,
paniculate or condensed, often consisting of clusters of panicles, 2.5-12.5 cm long, 2.5-9 cm broad, the
branches densely pubescent to tomentose, the hairs usually dark. Flowers distylous, sessile or on short spurs
to 2 mm long; calyx tubular, 8-9(-9.8) mm long, 2.5-3.5 mm wide at the mouth, 10-ribbed, puberulent or
strigillose to velutinous or pubescent, usually 4-5 lobed, the lobes uneven, deltate-acuminate, (0.7-)l.l-1.7
mm long; corolla white, tubular with spreading lobes, 20-27 mm long, 5-lobed, the lobes oblong, 8-10(-13)
mm long, (5-)6-8 mm wide, the tube 6.2-8.5 mm long; stamens 5, the filaments in long-styled flowers 10.5-14
mm long, the upper 4.3-6.2 mm free, in short-styled flowers (12-)15-17(-21) mm long, the upper 6.7-11 mm
free, puberulent to sparsely pubescent at the point of insertion, rarely glabrous, the anthers oblong, (2-)4-5
mm long; ovary depressed ovoid to ovoid, 0.7-1.3(-1.8) mm long, l-1.6(-2) mm broad, glabrous; style in long-
styled flowers 10-17 mm long, in short-styled flowers 8.5-1 1(-12) mm long, the stylar branches 2-3 mm long,
the stigma lobes clavate. Fruits enclosed by the persistent calyx and corolla, brovm, ellipsoid to narriowly el-
lipsoid, 7-8 mm long, 3.5-4 mm broad, one-seeded, the wall fibrous.
Mia gerascanthus is one of the more widespread members of the section found throughout Mexico Central
nbbed calyx, and paniculate mhorescence. It is unusual in exhibiting geographic variation in seve .1 rh 7
giapratus Mart., Flora 21(2
IAZIL. Minas Gerais; high dry carr
2:977. mi. Cordia glabra
Tree to 5(-6) m tall, the twigs sericeous when young, often hollow and inhabited by ants, the hairs malpighia-
ceous, later glabrous and waxy. Leaves deciduous; petioles (6-)ll-27(-37) mm long, canaliculate to flattened
on the adaxial surface, glabrous to sericeous; blades ovate to widely ovate, (3.5-)6-14(-20.5) cm long, (3-)4-
9(-14) cm wide, the apex obtuse to acute or rounded and occasionally abruptly acuminate, the base rounded to
obtuse, occasionally approaching subcordate, the margin entire, the adaxial surface glabrous and lustrous,
sometimes with scattered patches of malpighiaceous hairs, the abaxial surface sericeous with malpighiaceous
hairs or these sometimes fused and reduced with the abaxial surface appearing essentially glabrous. Inflores-
cences paniculate, (2-)6-10(-18) cm long, (2-)5-12 cm broad, the branches sericeous to densely puberluent.
Flowers distylous, on short pedicels 1-2 mm long; calyx tubular, 8-10(-14) mm long, 3-4 mm wide at the
mouth, prominently 10-ribbed, densely white puberulent, 3-5 lobed, the lobes uneven, ovate to deflate, 1-2
mm long; corolla white 21-28(-34) mm long, 5-lobed, the lobes oblong, parallel-sided or nearly so, or nearly
rounded, 8-13 mm long, 7-10 mm wide, rounded at the apex, the tube 8-12 mm long; stamens the same num-
ber as the corolla lobes, the filaments 15-22 mm long, the upper 5-10 mm free, glabrous, the anthers oblong,
2-4 mm long; ovary ellipsoid, 2-3 mm long, the style 17-20 mm long, the stylar branches 2-4 mm long, the
stigmas filiform. Fruits enclosed by the persistent calyx and corolla, brown, ovoid, 8-10 mm long, 5-6 mm
wide, one-seeded, the wall fibrous.
Distribution and habitat. — Cordia glabrata occurs in SW Brazil and adjacent Bohvia and Paraguay, where
it is found in dry forest and cerrado vegetation from sea level to 500 m.
NY). BRAZIL. Bahia: Santa Maria da Vit6ria,
rapora de Monte Claros, Castellanos 24239 (F).
i3352(CTES,FCQ.
calk Artigas y Via Feirea, S
Cerro Leon, baldzone on top of Cerro Leon, Hahn 1559 (MO). Concepcii
, Arroyos Y Esteros, Rio Piribebuy, Zardini 6793 (MO, P
.Rojas 1327 (AS, MO).
Cordia glabrata is a common species in central Brazilan cerrado vegetation and it can be easily recognized by its
leaves that are pale white on the lower surface. It is the only species of the section wit
Type: MEXICO E.W. Nelson 6924 (w
Small tree to 6(-10) m tall, the bark smooth, light gray, the twigs sparsely strigillose, later glabrous and sparse-
ly lenticillate. Leaves deciduous; petioles 15-33 mm long, shallowly canaliculate on the adaxial surface,
a^lTinate ITnel (5-)8-14.5 cm long, (3-)5-7.5 cm wide, the apex
^akous, rough to .he touch, the abaxial surf “e ZTSl!
bellate, generally less than 1 cm long. Flowers distylous; calyx tubular, 8-12 mm long, 4-4.8(-5.6) mm wide at
the mouth, the ribs more or less obscured by the villous indument, the hairs translucent to silver-gray or
golden, usually 5-lobed, the lobes uneven, debate to shallowly triangular-acuminate 1 3-2 5 mm lone
lO-ribbed; corolla white, tubular with spreading lobes, (18-)21-25.5 mm long, 4-5-lobed’ the lobes oblone to
mm long, the upper 5.5-10.5 mm free, glabrous, the anthers oblong, 2-3 5 mm long- ovarv de-
.iT ■ ® “"'“'■■^terifonrr, 1-1.6 urm tall l-7mm
'T‘'“ TZ ”” ^-8 '
years,suu.,„es.l„..h.eas.:ha.l.ss.a.„slsa:tk“^^
son 2784 (MO); 4-22 km NW of Rio San Nicolas and 20-40 km ^ of Tomaof ^7 ^stacibn de Biologta Chamela, Loti & Atkin-
Nu«a. drv. 220 «2 * ™«i ■« N«™, U.,u,
•Wfc 6. ruart. 6S3 (MEXU. MOf .fesg ,|„ jj, | ^ f Iflu. « to W ,1 Haeun,
Miffer6.renono667(MEXU,MO);alongthedinroadbet H ^ ^ of Huetamo de Nunez, 16 km W of ()uet
mlk. efev. 450 m.Soto6- Boom 2062 (MEXU). S™,
Johnston (1950) described both Co,
Miston (1950) described both CnrAt
" ta tngrh of calyx and
MB CordiagWndi/em ban uncommon sn^Zf’C^Zr “'“cured by addition!
sn « is infrequently collected, largel
Miller, Revision of Cordia section Gerascanthus
because it flowers in the middle of the dry season when most other species are completely dormant. It can he
recognized by its parallel-sided corolla lobes and condensed, nearly umbellate inflorescences, and it flowers
8. Cordia goeldiana Huber, Bol. Mus. Goeldi Hist. Nat. Ethnogr. 6:89. 1910. (Fig. 4). c
andBraganca, 23 Sep 1907, Goeldi s.n. (MG8319) (hc
Tree to 30 m tall, the branches glabrous, often lustrous, the axillary buds densely puberulent, at least when
young. Leaves deciduous; petioles (13-)20-40(-48) mm long, narrowly canaliculate on the adaxial surface,
glabrous; blades narrowly elliptic or lanceolate to elliptic-oblong or obovate, (7-)8-16 cm long, 2.3-7 cm wide,
the apex narrowly acuminate to obtuse and abruptly acuminate, the base cuneate to decurrent or acute, the
margin entire, the adaxial surface glabrous and lustrous, the abaxial surface glabrous. Inflorescences terminal,
a panicle or cluster of small panicles, 7-12 cm long, 12-17 cm broad, the branches glabrous to sparsely puberu-
lent but becoming densely puberulent near the tips. Flowers distylous, sessile, borne with the leaves; calyx
tubular, 8.5-10.5 mm long, 5-6.3 mm wide at the mouth, the lobes 2-3, uneven, widely ovate, 2.8-3.8 mm
long, lightly striate, sometimes not very evidently so, ribs not present smooth in bud, glabrous to short, brown
tomentulose or granular puberulous; corolla marcescent, white, funnelform 22-30 mm long, 5-merous, the
lobes oblong, parallel-sided, 11.5-17 mm long, 6.5-10 mm wide, the apex acute and sometimes uneven, the
tube 5-7 mm long- stamens 5, the filaments 10.5-19 mm long, the upper 5.5-12 mm free, puberulent to villous
at and just above the point of insertion, the anthers oblong, 2.4-4.3 mm long; ovary obloid to ovoid-obloid,
1-2.6 mm long, 1.2-2 mm broad, glabrous; disc indistinct to anular and ca. 0.8 mm tall, 1.2-1.3 mm broad,
glabrous; style 7-18 mm long, the stigma lobes clavate. Fruits not seen.
Distribution and habitat.— Cordia goeldiana is apparently localized near the mouth of the Amazon along
the banks of its major tributaries and in adjacent French Guiana and Guyana. Little is known about its eco-
logical preferences, but it presumably grows on tierra firme well back from the river banks.
Specimens examined BRAZIL. Amapa: Rio Amapari, Serra do Navid, elev. 70-300 m, Cowan 38447 (F, NY, US); Rio falsino, approx. 10 km
upstream of conHuence of Rio Arauari. west bank. Rabelo 2378 (F. MO. NY) R„.„a Florestal Ducke. Manaus-ltacoatiara, km
Belizon, toJeen Llld Eaux Claires.’elev. 200-300 m, Mori 21614 (MO; NY). GUYANA: Station des Nouragues, bassin de I’Arataye. Sa-
batier & Prevost 2695 (MO).
Cordia goeldiana can be distinguished by its glabrous leaves, parallel-sided corolla lobes, and calyx that lacks
evident ribs. It is unusual in its occurence in wet forests near the mouth of the Amazon.
oles (12-)15-25’(-28) mm long, canaliculate to flattened on the adaxial surface, glabrous to villous; blades
narrowly elliptic to elliptic to obovate, (5.5-)6-9(-10.8) cm long, 2.3-4(-4.5) cm wide, the apex acuminate to
acute, the base acute, the margin entire, the upper surface glabrous, the
long, 6-ircrbroad^ the brashes of the inflorescence covered with a very short, dense, curly, dark-brown
tomentum. Flowers distylous, short pedicellate or nearly sessile; calyx tubular, 9-11 mm long, 4-5 mm wide at
the mouth (8-)10-ribbed, puberulent, the hairs dense, curly, very short, dark-brown, (4-)5-lobed, the lobes
shallowly triangular; corolla white, funnelform, 22-25 mm long, 5-lobed, the lobes widely depressed ovate,
7-8 mm long 10-1 1 mm wide, the tube 9-11 mm long; stamens 5, the filaments 15 mm long, the upper 9 mm
free, glabrous’, the anthers oblong, 2 mm long; ovary cylindrical, 1 mm long, 1 mm broad, glabrous; style 19
mm long, the stylar branches 2.5 mm long, the stigma lobes clavate. Fruits unknown.
Journal of the Botanical Research Institute of Texas 7(1)
e than 80
Distribution and habitat.— This species is still known from only the type collection made r
years ago in central Guerrero in the dry forests of the hills of western Mexico.
Cordia gracilipes is still known only from the type collection made in a poorly collected area of central
Guerrero. It is a distinctive species that can be recognized by its long-petiolate, glabrous leaves, calyx with
irs, and a ciliate nectariferous disc.
Small tree or shrub to 6 m tall, the twigs glabrous, with circular to elliptic lenticels. Leaves deciduous; petioles
(3-)5-10(-12) mm long, flattened to slightly canaliculate on the adaxial surface, short tomentose, the haiis
somewhat deciduous, yellow-brown; blades coriaceous, elliptic to obovate, (3.7-)5.6-9.4(-10.5) cm long,
(1.6-)2.8-5.6(-6.7) cm wide, the apex obtuse to rounded, the base acute or less commonly obtuse to rounded,
the margin entire, the adaxial surface glabrous, the abaxial surface tomentose to arachnoid, the hairs wavy,
thin, yellow-brown. Inflorescences terminal, paniculate, 6-15 cm long, 5-10 cm broad, the branches downy-
tomentose, the hairs short, yellow-brown. Howers distylous, short pedicellate to sessile; calyx tubular, 11-13
mm long, 3-4 mm wide at the mouth, pubescent, the hairs appressed, 10-ribbed; corolla white, funnelform
19-22 mm long, 5-merous. the lobes widely depressed ovate, 5-6 mm long, 7-9.5 mm wide, the tube9-13mn
upper 6-10.5 mm free, glabrous, the anthers oblong, 2-2.!
im long, ca. 1 mm broad, glabrous; disc depressed ovoid, ca. 1 mm tall
long, the stylar branches 3.5 mm long, the stigma lobes c
s 5, the filaments 15-18 mm k
.oiuiapan, Seler&Sder 1636 (K, NY).
6213 (HOLomr: GH; ,L^F. Jaen, 700-
canaliculate on JabZs Petioles (9-)12-22 mm 1
pubemlem; blades lanceoUte to ovate, (6.-l-)10-15 c.^ ' Z ''''T' '
panicuUte, often with a lew smaU leaL scattered am n ‘"'‘‘’'■'^"tnces term
brown tonwntnlose togUbrons.
‘“-bbbed, 3-5-l<*ed, the lobesnneven 10^1 .r? ""
long, funnelform, 5-lobed, the bbes broadly oJoid 5-8 mm
^me,« the same number as the corolU lobes, the fiUments 14 w , ‘
i, Cajamarca, and San Martin.
emative specimens examined: PERU, t
Cajamarca: San Jose de Lourdes,
i. Luya, Km 2
re Jazan y Bagua Grande, e
ruerin v^nucnunuaiii, e.cv. 550-650 m, Compos & Upez 4962 (NY); Prov. Jaen, ja6n, carretera hacia ^aranj
(NY); Prov. Jaen, in the forest, elev. 500 m, Woyffeowsfet 5599 (MO, US); Prov. Jaen, Jaen— San Ignacio road, N of Jae
S of Tamborapa elev 700 m. Gentry 61020 (NY); Prov. jaen, Shumba, desvio al oueste de la carretera e •
3934 (m); .tag ™d Bag., C.nde tai P«Iro Ruiz, zW. «0-1,000 ui, W,,# I«38 (NY) S
ron=«,,dilcp«ove,taki„gBU,Hu.il.g.,zW350»,Gta.Y«ul377(»(M^^^
Cordiaigud*R<maUanuncommonspeciesofnor,hemPeru.hatisdistmc.iveinit3glabrou3leav=sa„dcalyx
and its hard, bony-walled fruits.
12 Cordia icnalensis Bartlett Contr. Gray Herb. 36:632. 1909. (Fig. 5). Gerosconthus iguoknsis (Barden) Borhidi, Acta
Bot. h!SmJ<I7. 1988. Tm: MEXICO. GuBuiaD: lgu.1. Canyon. 28 D« 1906, C.G. Pringiz 13912 (Houmm; GHi noiro; ENCB.
I.MEXUUS).
Tree lo 15 m tall ihe twigs glabrous. Leaves deciduous; pettoles (8-)13-19(-28) uun long, canaliculate on the
adaxial surface, neariy glabrous withsparse,short,appressed hairs; blades narrowVollipdc to ovate or slightly
Ilf il i
Journal of the Botanical Research Institute of Texas 7(1)
oblanceolate, (5.7-)8.3-16(-17.5) cm long, (1.6-)4-6(-9) cm wide, the apex acute to acun
to obtuse, the margin entire to slightly undulate, the adaxial surface glabrous, the abaxia
nearly so with stiff, short hairs on major veins. Inflorescences terminal, cymose-paniculate, 5-13.5 cm broad,
Ae branches glabrous except near the ends which are densely covered with short, interwoven, black hairs.
Howers distylous, on pedicels to 2 mm long; calyx tubular, 6.5-8 mm long, ca. 3 mm wide at the mouth,
tmncme^wid^^^^^’fl^^ restricted to the grooves between the ribs, 10-ribbed, nearly
wWety depj^ ovate. 6.5-8 mm lon&
17 mm ong, ihe upper 7-10 mm free, glabrous, the ambers oblong, 2-3 mm long; ovary broadly obovoid to
hoadly oblong, 1-1.5 mm long, 1-1.7 mm broad. gUbrous; disc depressed obovoid, 0.5-1 mm Jl ca 1 mm
""" '™8- '“I** “ flattened
,r ,'i!^ZsT‘' “ ‘“•'O -a 1" Mexico
ir the intersection of the borders of the states of Guerrero, Mexico, and Michoacan.
pec. AcatitUn, Hinton 3176 (F, G. G
onset of the dry season, in November and December.
13. Cordia insignis Cham., Unnaea 8:122. v* -r,.
BRAZIL; Minas Gerais, Uiotzliy s.n. (hm.oty!>e; B). Litho
oLjahrb.Syst. 12:560.
Shmb.oX-4)m.all,(.omathickenedundergrou„d"lylopodtainsomeca b ,
twigs puberulent to tomentulose when vounp lat^r stems annual,
M) mm long canalicuUte to ilattened on ihtadaxral sZZbllTs’^ l^^ves deciduous; petioles (2-)12-2
sbghtly^,eorobovate,(7-)12-22(-30)cmlcmg,(4-)7-Tu-wZ H
c^rrnally acute, the base rounded to obtuse, the marain entire tb Z' ’■““"'‘'‘f "f
and Irmrous with only a few, widely-scattered hairs the abaxial nigose-bullate, glabr.
^pattern, coarse ^f^cent along the veZ
Distrifw tt>-t5 mm 1
to central Bolivia up to 500 m in elevadTr^ Cerrado and dry forests from the
: western part of Bahia
Y San Simon, elev. 250 m, Arroyo P. et al. 2604 (MO. USZ). Santa Cruz: Prov. Nuflo de Chavez, Consepcion, Krapovickas & Schinini 31914
(MO). BRAZIL. Bahia: Brumado, Baixa dos Flores, Sobrinhi 255 (US). Goias: 100 km S of Guara, along Belem— Brasilia highway, elev.
500-800 ft, Maguire et al 56118 (NY. US). Mato Gr
etal 16707 (MO, NY).
1, MT251, 30 km NNE of Cuiabi, Mori
Cordia insignis is a very distinctive species of the Brazilian cerrado that is unusual in being a shrub, apparently
growing from an enlarged root and at least in some cases the aerial stems are annual in duration, and also in its
large, distinctly bullate leaves.
Tree, the twigs glabrous. Leaves deciduous; petioles 15-45(-60) mm long, canaliculate on the adaxial surface,
glabrous; blades elliptic to elliptic-oblong, 4-15 cm long, 2.5-7 cm wide, the apex abruptly short acuminate or
acute, the base cuneate to obtuse, the margin entire, the adaxial surface glabrous, the abaxial surface glabrous.
Inflorescences terminal, paniculate, 4-15 cm long, the branches glabrous or minutely brown puberulent or
glandular. Flowers distylous; calyx tubular, 8-12 mm long, 2-3 mm wide at the mouth, 10-ribbed, glabrous or
sparsely glandular-puberulent, 2-3-lobed, the lobes uneven, debate or rounded to obtuse, 1.5-3 mm long; co-
rolla white, funnelform, 25-33 mm long, 5-lobed, the lobes debate to ovate, 5-1 1 mm long, 6-12 mm wide, the
apex acute, the tube 10-12 mm long; stamens the same number as the corolla lobes, the filaments 18-25 mm
long, the upper 8-14 mm free, with short stiff hairs at the point of insertion, the anthers oblong, 2-3 mm long;
ovary ovoid, 1-1.5 mm long, 0.8-1.2 mm wide, glabrous; style ca. 20 mm long, the stylar branches 3-5 mm
long, the stigma lobes clavate. Fruits enclosed in the persistent calyx and corolla, brown, ca. 12 mm long, 3.5
mm broad, one-seeded, the wall fibrous.
Distribution and habitat. — This species is reported only with imprecise localities in the area surrounding
Rio de Janeiro but has not been collected for more than one hundred years.
Specimens examined: BRAZIL. Rio De Janeiro: Rio Janeiro, Riedel s.n. (NY. S).
Cordia latiloba is a rare species of southern Brazil that is very distinctive in its debate corolla lobes with an acute
apex.
Tree to 10(-18) m tall, the twigs glabrous and waxy. Leaves deciduous; petioles (7-)14-31(-41) mm long,
broadly canaliculate on the adaxial surface, puberulent or sparsely strigillose to glabrous; blades widely elliptic
to elliptic-ovate, 10-30 cm long, 4.5-15.3 cm wide, the apex obtuse to rounded and often abruptly acuminate,
the base obtuse, the margin entire, the adaxial surface glabrous, the abaxial surface villous to sparsely pubes-
branches puberulent above, often glabrous near the base. Flowers distylous, sessile; calyx tubular, 18.5-20 mm
long, 6.4-6.9 mm wide at the mouth, prominantly 10-ribbed, densely puberulent, 3-lobed, the lobes debate-
acuminate, 2-2.2 mm long; corolla white, tubular with spreading lobes, 40-41.5 mm long, 5-merous, the lobes
very broadly ovate to depressed ovate, rounded at the apex, 10.3-14 mm long, 12.5-14.5 mm Avide, the tube
17-19 mm long; stamens 5, the filaments 23-28 mm long, the upper 11-13.5 mm free, pubescent at the point
of insertion, the anthers oblong, 3.3-3.5 mm long; ovary very broadly ovoid, 1.5 mm long, 1.8 mm broad, the
disc anular, 0.9 mm tall, 1.2 mm broad, glabrous; style 20-25 mm long, the stylar branches 3 mm long, the
stigma lobes clavate. Fruits enclosed by the persistent calyx and corolla, brovra, ellipsoid, 9-11 mm long, 4.4.5
mm broad, one-seeded, the wall fibrous.
Distribution and habitat.— This species is endemic to Ecuador where it occurs in dry forests on the coastal
plain up to 500 m in elevation.
Km 21. Guayaquil to Daule, eiev. zv-iqq Dods
13849 (MO); Capeira, Km 21, Guayaquil to t^uie eiev
Cevanos&NeiH 208 (MO, QCNE). ^ ^ Manabi.
m iw ^ ’ ^“ayaquil to Daule, eiev. 20-200 m, Dod:
». 22 „„ N of C„ay^.„ „„ ”
ili “uT ‘
Cordiam«ra«l„Uadis.tactivespcdesfo„„din,hedryforesBofw^^ p
^‘"""“'““’'’“''eleavesandaoweps.Py^ose-panJ^
S. Cordia macvangliHj 5
adaxW surface, pufK^mf '“"S’ “naliculate c
oUuse 10 ruuuded, .he base „b..«e „ acule. the marl eml ,hl'f T ‘T ® ™
the upper 5.5-11 mm fee, the lowest portion of the fee hlamenu thickenerl, pubescent at the poirit of inser-
tion, the anthers ohlong, 2-3 mm long; ovary eylindrical, ca. 1 mm long, 1 mm broad, glabrous; style 10.5-17
mm long, the stylar branches 1-2.5 mm long, the stigma lobes clavate. Fruits enclosed by the persistent calyx
and corolla, brown, wall thin, fibrous, mature fruits not seen.
Distribution and habitat.— This species is endemic to western Mexico, where it occurs in dry forests of
:en 600 and 1,300 m in elevation.
elev. 660 m, Miller & Tellez3082 (MEXU. MO, NY).
Cordiu mucvaughii is a well-marked species characterized by a large paniculate inflorescence with branches
coveredwithdownyvelulinous pubescence, corolla lobes thatarewider than long, the lack oflongpubescena
on the calyx, and rather large, broad leaves. It appears to flower while leafless, shortly after the leaves are shed.
17. Cordia megalantha S.F. Blake, Proc. Biol. Soc. Washington 36:200. 1923.
1988. Type; GUATEMALA: Izabal, Quebradas. 18 May 1919, S.F. Biafee 7498 (holotype: US; isotype; G).
Tree to 30(-60) m tall twigs glabrous, lenticellate. Leaves deciduous; petioles (8-)ll-33(-55) mm long, cana-
tauTe^n thlaxiai sull glabp
margS ^1^ the a^xihl MX^ghbrous, the abaxial surface glabrous. Inflorescences terminal, panicu-
late to 22 cm long 30 cm broad, the main branches gUbrous, puberulent at the tips. Flowers distylous, on
pedicels 2-5(-10) mm long; calyx tubular, (8.5-)9-10(-l 1) mm long, 4-5.5 mm wide at the mouth, stnate to
10-20-ribbed, glabrous to puberulent, the hairs dark brown, unevenly lobed, tearing upon dehiscence or de-
hiscing circumscissilly corolla white, funnellorm, 28-43(-50) mm long, 5(-6)-lobed, the lobes debate to
ovate acute at the apex, C8.5-)ll-13(-18) mm long, (8-)10-12.5 mm wide, the tube 7-9(-11.5) mm long; sta-
mens 5(-6), the filaments 14.5-19 mm long, the upper 5-10(-13) mm free, pubescent at the point of msertton
and frequently over the entire fee portion, the anthers oblong, 2-2.6(-4) mm long; ova^ ovotd to comcal,
l. 3-2.5M) mm long, 1-2 mm broad, glabrous; disc depressed obovoid, 0.5-1 mm util, 1-2 mm broad gb_
bums to ciliate; style (7.5-)16-19 mm long, glabrous or with a lew scattered hairs the stylar bmnch^ 3-4.5
mmlong, the stigma lobesclavate. Fruits enclosedhy the pemistentcalyx and corolla, brown, elhpsotd to nar-
rowlyellipsoid,8-12mmlong,4-6mmbroad, one-seeded, thewallthin,fibrous.
Distribution and habitat.-Cordia megalantha ranges from southern Mexico south through the Wland,
Atlantic wet forests to Costa Rica and also occurs in a disjunct population in wet forests of the Pacific slope on
theOsaPeninsulaofCostaRicaandadJacentBuricaPeninsulaofPanamauptohOOOminelevation.
Representative specimens examined: COSTA RICA. Alajucla: San Carlos, Llanura de San C
1505 (CR, NY). Goanacaste: Parque Nacional Rincon de la Vieja Quebrada Leiva a 1.5 km a^i
m. Rivera 1267 (MO, NY). Heredia: La Selva, Bawo 606 (MO). Limon: vicinity of Guapiles, elr
>a Peninsula, Liesner 1941 (MO). San Jose: F
1320 (CR, MO, NY). GUATEMALA. Izabal: lower Motagua,KuyIenG147(CFMR,F,
of La Ceiba. elev. 40 m. Stead SI42 (FHO, MEXU, MO). Yoro: Road to El Progreso at
I2k..rr™,BPr.g,.».kr.^lWMO).r~™------"---.r.o.d^^
et al. 32294 (MO). PANAMA. C
Cordia megutatha is the tallest of all Cordiu species, often exceeding M m, and ^casionally reachmgM m in
height. It also has the largest flowers of any of the species of section Geru^uuthu^wtth corollas 28-43(-M)
mr^ng It is easily recognized by its ovoid, apiculate buds and deltate com la lobes unth an acute apex. The
calyx is^nerally n« distinctly ribbed, but with 10-20 striations and the calyx opens by teartng tnto uneven
lobes or it dehisces circumscissilly.
18. Cordi. nK,„k«™a Standi, Contr. US. Natl Hetb. 2M220. 1924. (Fig. 7).
lung. 34.398. 1988. Type: MEXICO. Morelos: near Cuernavaca, elev. 5,000 ft, 17 Mar 1899, C.G. Pringle 8205 pro
)■■ rsoucroryns: ARIZ, ASU, BR, C, CAS, F, G, GH, IND, K, L, MEXU, MICH,
---ey designated Pringle 8205 at US (sheet #3544555) as the holotype, but the
-..t two gatnenngs and are therefore an invalid t ' undl^ ^ ‘collected on 29 Sep 1899, which repre-
Therefore,thenoweringbranchesareheredesignrd«thent^“”‘'"°^'‘‘'‘"‘""“
brous, the abaxial surface hirsute to less com ^ ^symetncal, the margin entire, the adaxial surface sca-
condensed, to 3.5 cm lone 3 5 cm y stngose or scabrous. Inflorescences terminal, paniculate or
buUr. 1 1-14.5(-19) mm long, t-Si-M) mm wtt auhl mou^Q B h "I ™
late-acuminaic, 1.4-2.7(-3.5) mm long 10 ribhfH ,b -i_ ‘h' >“b«i uneven, usually del-
halm ttanslucemor white to pale gta^mira •>-
ous, the lobes widely depressed ovate to depressldZT(^TmuT^\'""'
wide, the tube (8.6-) 9-13(-15.7) mm long stamens the ' ^ 'ung, (7.6-)9-12.5(-14) mm
long-styled flowers 11.5-16(-18) mm lone the imr^r-r a “ the corolla lobes, the filaments in
25) mm long, the upper (9.6-) 10.5-12 5(ll5) f ^^‘’’’t-^owered flowers (17.7-) 19-22(-
glabrous, the anthers oblong, 2.4-4 mm lone- ovarv of insertion of occasionally
brotm; dise annular-cratedform. O^U 2 oTsT r?fb ^
llowem (15.5-)17-20 mm long, in short-styW n 6Mm * '» W
■H-S) mm bug, the stigma lobes clavate. Fruits enclosed b ® branches (1.5-)2.3-
h* """ """
““'^"”'Morlta3''^?b!l‘! wCm hCrlmTT ““
800-1.650 m in elevation. tn dry forests and open areas from (340-)
elcv^m' MEXICO Goerrero: 19 km S of Iguala on M
(DS, SD). ^ Tepoz'lan, elev 1,900 ra, Rzedowski 22348 (ENCB ^ Morelos: Bar-
Cordiamorel ’
Erect tree to 5(-10) m tall, the bark light gray, smooth, the twigs sparsely strigillose or pubescent when young,
later glabrous and sparsely lenticellate. Leaves deciduous; petioles (3-)6-14(-19) mm long, flattened or shal-
lowly canaliculate on the adaxial surface, sparsely to evenly strigillose or pubescent with most of the hairs on
the adaxial surface; blades usually elliptic, occasionally narrowly elliptic or ovate, (3-)4-13(-18) cm long,
(1.4-)2.5-6.5(-8) cm wide, the apex usually acute, occasionally obtuse, the base obtuse to acute, the margin
entire, the adaxial surface glabrous or nearly so, usually with numerous papillae, the abaxial surface glabrous
or nearly so, often with scattered appressed hairs along the veins. Inflorescences terminal, sohtary or clusters
of sparsely branched panicles to 17 cm long, or condensed, often nearly umbellate, the branches villous to to-
mentose. Flowers distylous, sessile or on short pedicels to 3 mm long; calyx tubular, (10.6-)12.5-18(-20) mm
long, (3-)4-5 mm wide at the mouth, 10-ribbed, pubescent or velutinous to occasionally villous, unevenly
3-3-lobed, the lobes deltate-acuminate, (l-)2-3(-4) mm long; corolla white, funnelform, 24-36 mm long,
5(-6)-merous, the lobes depressed ovate to rarely widely depressed ovate, 10-12.5(-14.6) mm long, (9-)10-
1.5(-14.6) mm wide, the tube (8.4-)10.7-15.7 mm long; stamens 5(-6), the filaments in long styled flowers
16-23 mm long, the upper 5.7-9.5 mm free, in short-styled flowers (16-)19-27 mm long, the upper (9-)10-14.5
free, pubescent at the point of insertion or rarely glabrous, the anthers oblong, 2.8-4.1 mm long, ovary broadly
depressed ovoid, 0.9-1.5(-2) mm long, 1-2 mm broad, glabrous; disc transversely ellipsoid to obloid, 0.8-1
mm tall, 0.9-1.8 mm broad, glabrous or sparsely ciliate at the upper edge; style in long-styled flowers 20-30
mm long, in short-styled flowers (9.5-)ll-16 mm long, the stylar branches 2-4.6(-6.8) mm long, the stigma
lobes clavate. Fruits enclosed by the persistent calyx and corolla, brown, ellipsoid to narrowly ellipsoid, 7-10
mm long, 2-3 mm broad, one-seeded, the wall fibrous.
Distribution and habitat. — Cordia sonorae occurs in western Mexico in dry forests with a northern popula-
tion in Chihuahua and Sinaloa, a disjunct population on Maria Madre Island in Nayarit, and a more southern
population in Michoacan and Guerrero. It ranges from sea level to 600(-l,000) m in elevation.
Satnachique with Rio Batopilas, elev. 600 m Bye 3458 (INIF, MEXU). Guerrero: 50 km S of Iguala, 11 km S of Rio MeLala, elev. 600 in.
Miller et ol. 678 (MO). Michoacan: along Mexican Highway 200 between Playa Azul and Tecoman, Colima. 6 km W of Maruhuata, elev. 30
ra. Miller 6- Tellez 3088 (MEXU, MO). Nayarit: Maria Madre Island, Nelson 4207 (F. US). Sinaloa: between Rosario and Villa Union, Gentry
12598 (LL, MEXU). Sonora: about 5 mi. below Minas Nuevas, Rose et al. 12668 (NY. US).
Cordia sonorae is not uncommon in northwestern Mexico from Guerrero north to Sinaloa and Chihuahua. It is
most easily confused with C. morelosana (see remarks under that species).
20. Cordia thaisiana G. Agostini, Brittonia 25:174. 1973. (Fig. 6). Tyte: VENEZUELA. Zum: between Maracaibo and
Villa del Rosario, 15 Mar 1972, J. Steyermark, G.C. K. & E. Dunsterville 105,527 (holotype: VEN; isotypes: GH, NY, US).
Tree to 20 m tall, the twigs glabrous. Leaves deciduous; petioles 5-20 mm long, canaliculate on the adaxial
surface, glabrous or nearly so; blades elliptic-obovate, 4-12 cm long, 1.5-4 cm wide, the apex short acuminate
to acute, the base narrowly cuneate, the margin entire, the adaxial surface glabrous, sometimes pustulate, the
abaxial surface glabrous. Inflorescences terminal, cymose, 3.5-5 cm broad, 12-15-flowered, the branches
sparsely strigose. Flowers distylous, borne with the leaves; calyx tubular, 8-9 mm long, ca. 3.5 mm wide at th(
mouth, glabrous or nearly so, usually 3-4 lobed, the lobes uneven, ovate, 3-3.5 mm long, not evidently ribbee
butmerely striate; corolla white, 15-24 mm long, (4-)5-lobed, the lobes oblong, parallel-sided, 6-lOmm long
5-8 mm wide, the tube 9-10 mm long; stamens the same number as the corolla lobes, the filaments 1 3-1 5 mn
long the upper 7-8 mm free, pubescent at the point of insertion, the anthers oblong, 4-5 mm long; ovar
ovoid,l-1.2mmtall,lmmbroad,glabrous;style5-6mmlong,thestylarbranchesalmostnotevident,thefou-
lyx and corolla, brown, ellipsoid, ca. 5 mm long, one-seeded, the wall fibrous
Distribution and habitat.— Cordia thaisiana occurs in dry forests in wecn>rr, v i j j
Colombia. ^ Venezuela and adjacen
lo ampeche, Dugand553 (F); S«
w: along the rail
itheCerrejdnm
=che, Dugand 328 (F); in El Paraiso. Dugand 521 (F); road from
Y. 50-150 n,.Dugand573(US);nearMolinero,D„gand 575(F).
Forestal de T
Forestal de aparo, al None de los Rios Uribanic i ,
2894 (MO); Marcano-Bertt 2895 (MO). Falcon: A lo largo de la car
id,0.2-0.7
Journal of the Botanical Research Institute of Texas 7(1)
Tree 6-25 m tall, the twigs densely stellate-pubescent. Leaves deciduous; petioles 1-4 cm long, canaliculate on
the adaxial surface, densely stellate-pubescent; blades ovate to lanceolate, elliptic, or obovate, 6-15 cm long,
3-8 cm wide, the apex acute, the base acute, the margin entire, the adaxial surface glabrous or with sparse stel-
late hairs, the lower surface densely stellate-pubescent to stellate-tomentose. Inflorescences terminal, cymose-
paniculate, 8-30 cm broad, the branches densely stellate-pubescent. Flowers of three distinct morphs, two
distylous morphs and one homostylous, non-herkogamous morph, calyx tubular, 6.5-9 mm long, ca. 2.5-3.5
mm wide at the mouth, 10-ribbed, stellate-pubescent, (4-)5-lobed, the lobes acute, ca. 1 mm long, or nearly not
evident; corolla white, 12-17(-24) mm long, tubular with spreading lobes, 5-(6)-lobed, the lobes 6-9 mm long,
3-5 mm wide, the apex truncate, the tube 6-8 mm long; stamens the same number as the corolla lobes, the
filaments 11-15 mm long, the upper 5-7 mm free, pubescent at the point of insertion, the anthers oblong,
2.5-3 mm long; ovary ovoid to broadly ovoid, 1-1.5 mm tall, ca 1 mm broad, glabrous; style 6-10 mm long, the
style branches 2-3 mm long, the stigma lobes clavate. Fruits enclosed by the persistent calyx and corolla,
brown, subcylindrical, one-seeded, the wall fibrous.
Distribution and habitat. Cordia trichotoma ranges from Pernambuco in northeast Brazil through Bolivia
and Paraguay into northern Argentina from sea level to 1,000 m.
7. La Paz: Bautisu Saavedra, elev. 1,600 m, Zenteno 794 (M). Santa Crnx: Nuflo de Chave"
. 29.2 km S of Emborozu— Sidras road on road to Bermejo, (12.7 kn S of Naranjo Agrio), elev. 600 m. Sole
1. de Quebrangulo, Reserva Biologica de Pedra Ralhada, Cervi et al. 7192 (MO). Bahia; Mpio. of Iraq,
SSE of Iraquara. elev. 600 m, Mori et al. 14413 (MO, NY). Ceara: Iparana, mat
, — ™«--ade Rio deSao Bartolomeu, Herineeretal. 4535 (MO US) Esniritn i
J.5. 045/78 (MO). GoUs: ca. 5 km Sof CaiapSnia, elev. 850 m. Arulerson 9475 (US). Maranhao: Carolina, close to Rio UJes, Ratter 6767 (1
900-l,000m, miliams &Assis 6741 (F, MO,
JoJ«i-n,L dc. 150 m, Cair, ,| d. 59399 (MO). Ca„,d, Villa Ells. Pedtrs™ 31?3 njs) Co,*™,.. a 7
Belfo Horizonte, elev. 270 m, Zardim &■ Guerrero 41198 (AS MO) r™viilG a , ^ ^ Arroyo Tag^tiy^-Guazu at Esta
Ovetense. 12 km al NE de Chore, Zardim & Benitez3416 (MO, PY). ' ‘
leaf surfaces, lldmoslodily confused wi.hC.allW9rd,butdiL in
are vein, mens Wow.and larger ffo«rs.ItUvalnedasalimW tree
aKmipexAgoslini,AclaBot.Venez.9'292 1974 (Fi„ « ,
Tree to 8 m tall, the twigs glabrous. Leaves deciduous netiolP..3_Q ,
face, sparsely villous or glabrous; bladeseUiptic 3 5-m i ^ ® on the adaxial <
2-3 lobes the lobes uneven, ovate, 0.5-2 mm long, evidently 10-ribbed; corolla white, funnelform, 18-21 mm
long, (4-)5-lobed, the lobes 5.5-6 mm long, 4.5-4.7 mm wide, the apex acute, the tube 5-6 mm long; stamens
4-5 the filaments 12-17 mm long, the upper 7-9 mm free, pubescent at the point of insertion, the anthers ob-
long ca 2 mm long; ovary depressed ovoid, ca. 1 mm tall, 0.6-0.8 mm broad, the style 8-11 mm long, the
stylar branches 2-4 mm long, the stigma lobes clavate. Fruits enclosed by the persistent calyx and corolla,
brown narrowly ellipsoid, 5.5-6 mm long, ca. 2.5 mm broad, one-seeded, the wall fibrous.
Distribution and habitat.-Cordia umbellifera occurs in Eastern Venezuela and adjacent Guyana from sea
level to 450 m.
Specimens examined: VENEnJElA. Bolivar: Proc. Criollo, Ciudad Bolivar, entreei “Club NauticoYAngosturita,”elev. 20 m
(MO)- northern most slopes of Cerro Baraguan, 100-330 m. Wurdack & Monachino 41238 (MO, NY). Guanco: Entre Camatagua y m
breroAristeguieta5985(NY,VEN);7289(VEN);betweenOrtiz&GuaricoBridge.Pi«ierl2222(NY);Ortiz-GalerasdeElPao,20kmdeSan
FrandscodeTiznado,Aristeg«ietaetal.6269(VEN);PasoReal.roadtoCementerio,Plowman]9IO(US).
Cordia umbellifera is an uncommonly collected species that can be recognized by its corollas with lobes wider
than long, pubescent staminal filaments, and highly condensed, umbellate inflorescences.
ACKNOWLEDGMENTS
1 would like to thank Wayne Law for preparing the distribution maps. Thanks go to James 1. Cohen and an
anonymous reviewer for critieal and constructive reviews. Kanchi Gandhi (GH) kindly looked at the
Airy Shaw H K 1 973. A dictionary of the flowers plants and ferns. Cambridge University Press, U
Avetissian, E.M. 1956. Morphologie des microspores de Boraginaceae. Trudy Bot. Inst. Akad.
Armyansk. S.S.R
De Cahooue, A.P. 1 845. Prodromus systematis naturalis regni vegetabilis. Paris. B-^^aceae. 9^50E
Gibbs, P.E. and N.Taroda. 1983. Heterostyly in the Cordia alliodora-C trichotoma complex in BrazH. Rev. B as. Bot. 6 Ija
Gottschling M J S Miller M Weigend, and H.H. Hilger. 2005. Congrence of a phylogeny of Cordiaceae (Boraginales) in-
TrrfmlTSl“eTatawithmorpho
Johnson P.andR Mof^es 1972. A review of Cord/oa/Z/oc/oro (Ruiz &Pav.)Oken.Turrialba 22:21 0-220.
JoHNSToiii, I.M. 1930. Studies in the Boraginaceae 8. Observation on the species of Cordia and Tourr^efortia known from
Brazil’ Paraguay, Uruguay, and Argentina. Contr. Gray Herb. 92:3-89. 1
Johnston, 1.M. 1950. Studies in the Boraginaceae 19. B. Cordia section gerascanthus in Mexico and Central America. J.
Arnold. Arbor. 179-187. • * a- 1 Am^iH Arhnr
Johnston, I.M. Studies in the Boraginaceae 20. Representatives of three subfamilies in eastern Asia. J. Arnold
32:1-26; 99-122.
KHALEEL,T.F.1975.EmbryologyofCord/aBot.Gaz.136(4):380-387.
Khaleel,TF 1982 Embryology of Cord/flsebesfena (Boraginaceae). . yst. vo. .
Lawrence, J.R. 1937. A chelation of the taxonomy and the floral anatomy of certain of the Boraginaceae.
places
J.S. M.UO. 1990. Pollen morphology of .he Cordloideae (Boraginaceae)-. doxemmo. Cordio, and Po-
S™ou;p.C,g». Boraginaceae. ImTrees and shrubs o,Mexico.Conh.U.SNa,,^^^^
T^ooa. N. P.E. &..S. 1986. Smdies on dm genus (asrd/o L (Boraginaceae) ,n Braa,l. 1
and conspectus. Revista Brasil. Bot. 9:31-42.
ir. 3, 23:289-295.
id their ants. Bull. Mus. Comp. Zool., Harvard 90:1-2
Journal of the Botanical Research Institute of Texas 7(1)
JOURNAL REVIEW
Ashok Gadgil and Diana M. Liverman. 2011 (Nov). Annual Review of Environment and Resources Volume
36. (ISSN: 1543-5938; ISBN: 978-0-8243-2336-3, hbk). Annual Reviews, Inc., 4139 H (imi^Way RO
Box 10139, Palo Alto, California 94303, U.S.A. (Orders; www,AnnualReviews.org, service®annnalro
views.org, 1-800-523-8635, 1-650-493-4400). $80.00, 479 pp., 7 Vs" x 9 W.
This 36th volume of Annual Review of Environment and Resources focuses on emerging scientific and policy is-
es at the intertace ot environment, resource management, and development. This volume contains a series of
papers that review scientific, policy, technological, and methodological issues related to dobal life sunnort
tesearchandByhi^HBXr-"^^^^^
Br3.sectil7"banZ~^^
rdeoftheoceaninfuturecoi^mitmemsforl
andIn„^nchen.is.ry.AZ:“Zr^^^^^^^
become the most intensely altered of any ecosystems. freshwater ecosystems have
Thesecondsectionlooksatthepattems, drivers, and consequences nfhiiTnar,. r
and interactions with the environment. This section rom • ‘^ses of natural resources
technology for managing the imnacts of rorxi j several reviews related to energy (e.g., policy and
ergya,xfT„e.*yuJ.r.'aX.:X:Ta7^^^^^
mediate land degradation and the devplonmpr.t r i. ^ examine mechanisms to combat and re-
munication among various disciplines acf u ^ environment. The volume aims to foster com-
.hepublicwl„ryly„nsuchl„forma,i„„ .oaZZstaZ'"" 7 ”'7”*
S, U.SA.
NOVELTIES IN SERPOCAULON (POLYPODIACEAE)
This species has commonly been called Polypodium giganteum Desv. (Lellinger 1985, 1989; Moran 1995), or
Serpocaulon giganteum (Desv.) A.R. Sm. (Smith et al. 2006). However, the type of P. giganteum (Anonymous, ex
Herb. Desvaux [P*-00637550]) corresponds to a species of NiphidiumJ. Sm. (Polypodiaceae), probably N. cras-
sifolium (L.) Lellinger, widespread in the Neotropics. We here provide a better name (along with its new com-
bination) for the specimens long named “Serpocaulon giganteum” or “Polypodium giganteum”: Serpocaulon
The characterization of this species can be found in Lellinger (1989) and Moran (1995), as “Polypodium
giganteum.” Although Smith et al. (2006) cited this species (as Serpocaulon giganteum) from southern Brazil, in
Brazil it probably occurs only in the northern region, in the Amazon Forest (see Labiak & Prado 2008; Labiak
&Hirai2010).
Lellinger (1985) correctly attributed the combination Polypodium fraxinifolium subsp. articulatum to
Christ (1906), and not a new combination based on Desvaux’s illegitimate name. However, he mistakenly des-
ignated as lectotype a specimen cited by Christ: Pittier 9061 (US*-00065825). There is one sheet at P (P*-
00624694) that corresponds to Desvaux’s original material, and it is appropriate to consider that as the lecto-
type (ICBN Art. 9.17), redesignated here.
K Schwartsb. & A.R. Sm.,
lineous, copiously scaly, 5-10 mm diam.; scales lanceolate-
acuminate, 0.5-0.75 X 0.2-0.3 mm, peltate basally, clathrate, with margins entire, concolorous. Fronds erect,
monomorphic, 1.65-1.8 m long; petioles brown, glabrous, with two lateral lines of aerophores, 68-77 cm long,
4-6 mm diam.; laminae 1-piimate, linear-acuminate, slighdy broader at base, the apex conform, 100-110 x
50-72 cm, lateral pinnae 12-15 pairs; rachises glabrous; proximal and medial pirmae ascending (sometimes
strongly ascending), ca. 45-60° from rachis, linear, often falcate, 25-36 x 2.5-3 cm, with pinna bases inequi-
lateral, cuneate, basiscopically petiolulate, acroscopically sessile, the margins cartilaginous, crenate; costae
nearly glabrous, but abaxially and adaxially with scattered filiform scales 0.4-0.6 mm long and scattered
trichomidia 0.1-0.15 mm long, 1-2-celled; veins anastomosing in a goniophlebioid pattern, with 5-7 areoles
between the costae and the piima margins, nearly glabrous, but abaxially and adaxially with scattered tricho-
midia similar to those of the costae; laminar tissue between the veins abaxially and adaxially glabrous; sori in
4-6 uniseriate rows between main lateral veins running from costae to pinnae margins.
Distribution and ecology. — In the western region of the Brazilian States of Minas Gerais and Sao Paulo,
between the rivers Rio Grande and Tiete. It occurs terrestrially along the margins of streams of Seasonal semi-
Deciduous forest. It probably occurs also in the States of Mato Grosso do Sul and Goias.
Etymology. — The specific epithet is an allusion to the size of the fronds, which are among the largest in
AL. Barbosa & W. Forster 21 (SJRP); Nhandeara, ca. 20°43'S, 50°03'W, 18 Jun 1994, F.R. Nonato & MR. Pietrobom-Siha 24 (SJRP); Planalto,
ca. 21°02'S, 49°55'W, 15 Aug 1996, M.R Pietrobom-Silva& EM. Pedro 3423 (SJRP); Presidente Epitiicio, 22”05'S, 52“07'W. 250-300 ni, 23 Sep
1996, M.R. Pietrai>om-Siha & C.T. de Lucca 3476 (SJRP); Rosana, regiao do Pontal do Paranapanema, 22°33'S, 52'’44'W, 26 Jul 1997, M.R. Pit-
trobom-Silva 4086 (SJRP); Tanabi, 12 Dec 1990, L.M.S. Viona et al. 1 (SJRP); Teodoro Sampaio, Rio Paranapanema 22°38'S, 52°45'W, 24-26
Feb 1966, P.G. Windisch 4720 (^RP).
1, S. rex differs from S. triseriak
(Sw.) A.R. Sm. by stramineous rhizomes (vs. dark brown), concolorous rhizome scales with the margins entire
and clathrate (vs. scales bicolorous with margins subentire and hyahne),/ronds 1.65-1.8 m long (vs. 0.5-L2 m
Schwartsburd and Smith, Novelties in Serpocaulon
iirs ol pinnae (vs. 5-10 pairs), proximal and medial pinnae with bases basiscopi-
y sessile (vs. bases basiscopically sessile, acroscopically adnate), and with crenate
margins (vs. margins entire to slightly undulate), and 4-6 rows of son between costae and pinna margins (vs.
1-3 rows)— Figs. lA-H.
Serpocaulon rex differs from S. menisciifolium (Langsd. & Fisch.) A.R. Sm. and 5. hirsutulum (T. Moore)
Schwartsb. & A.R. Sm. by growing terrestriaUy (vs. epiphytically), /ronds erect (vs. arched) and much larger,
reaching 1.65-1.8 m long (vs. 0.5-1.4 and 0.45-0.55 m long, respectively), proximal and medial pinnae with
bases basiscopically petiolulate, acrocopically sessile (vs. bases basiscopically sessile, acroscopically adnate),
glabrous rachises (vs. rachises pUose or villous, with septate hairs and lanceolate scales), and 4-6 rows of sori
between costae and pinna margins (vs. 1-3 rows)— Figs. lA-H (confront with Figs. 2A-H of 5. hirsutulum).
Serpocaulon rex differs from 5. fraxinifolium Qacq.) A.R. Sm. by growing terrestrially (vs. epiphytically),
rhizomes thick, stramineous, and copiously scaly (vs. slender, black, and glabrescent or
overlapping, round scales), fronds much larger, reaching 1.65-1.8 m long (vs. 0.4-0.9 m long) a
arched), p
copically and acroscopically sessile), and 4-6 rows of sori between c
To verify that Serpocaulon rex had not been already described, we observed online the types of the syno-
nyms of Polypodium/roxini/olium Jacq., P. menisciifolium Langsd. & Fisch., and P. triseriale Sw. listed by Hensen
(1990) and ascribed to Brazil: Goniophlebium excelsior Fee (Glaziou 3334 [P*-6 sheets]; = S. mensiciifolium) G.
gauthieri Fee (Gauthier s.n. [RB*-31818], Glaziou2406 [P*]; =S. menisciifolium), P. albopunctatum Raddi (illustra-
tions in Raddi 1825!; and in Pichi Sermolli & Bizzarri 20051; = S. menisciifolium), P. distans Raddi (illustration
in Raddi 1825!; = S. fraxinifolium), P. elatius Schrad. (Wied-Neuwied s.n. [BR*-697061]; = S. menisciifolium) P.
lucens Schrad. (Wied-Neuwied s.n. [BR*-697025]; = S. triseriale), P. mosenii C. Chr. (Glaziou 4422 [P*-3 sheets],
Mosen 2208 [P*]; = S. mentsciifolium), and P. preslianum Spreng. (type?: Brackenridge s.n. [PH*-00021063]; = 5.
triseriale).
Besides the geographical distribution, Serpocaulon rex differs from 5. articulatum by laminae with 12-15
. ^ ^ . .. I wide (vs. elliptic, (3-)4-
largins 0
l-3(-4)
n (T. Moore) Schwartsb. & A.R. Sm., comb. nov. (Figs. 2A-H). ]
Plants epiphytic. Rhizomes creeping, chalky-white, copiously scaly, 5-6 mm diam.; scales debate, 15-3x1 5-2
!II!^ghLt!L^l^r^rena^^^^^ plantlete), 4-5 x^l-1.2 mm, peltate basally, clathrate, bicolorous, with
to light brown, glabrate, without aerophores, 15-20 cm long, 1.8^.8 mm diam.;
ovate, the apex conform, the distalmost pair of pinnae much reduced, 30-35 x 18-20 cm, lateral pinnae 9-13
pairs; rachises abaxially and adaxially densely villous, the hairs whitish, lax, 0.5-1.2 mm long, 5-12-celled,
abaxially also with sparse scales, the scales clathrate, lanceolate, 0.7-1 x 0.5-1 mm; proximal pinnae deflexed,
medial pmnae patent to slightly ascending, ca. 70-90° from rachis, linear-oblong with the apices round to ob-
tuse, 9 10 X 1.5-2.2 cm, with pinna bases inequilateral, basiscopically sessile, acroscopically adnate the mar-
gins cartilaginous, undulate, villous; costae abaxially and adaxially villous, abaxially sparsely scaly; veins
with 3(-4) areoles between the costae and the -y^rgins.
c Forests of the Brazilian Atlantic For-
Schwartsburd and Smith, Novelties in Serpocaulon
Journal of the Botanical Research Institute of Texas 7(1)
Schwartsburd et al. 2649 (UC!, V
This species has commonly been regarded as doubtful. After Raddi’s (1825) description, Fee (1869, 1873) re-
garded it as “not well known”; Brade (1951) considered it, with doubts, as a juvenile form of Polypodium menis-
ciifolium; and Hensen (1990) synonymized it under P. menisciifolium. Pichi Sermolli and Bizzarri (2005) found
a solely sterile type-specimen, and also regarded it as synonym of Goniophlebium menisciifolium (Langsd. &
Fisch.)J.Sm.
On the other hand, Moore (1862) recognized it a species, and included it in the genus Goniophlebium.
Rosenstock (1925) also recognized it as a species, citing one collection we have examined (Brade 8246), and
compared it to his Polypodium menisciifolium var. pubescens Rosenst. On Brade 8246 (UC), he wrote “Polypodi-
um raddianum, nom. nov., P. hirsutulum Raddi,” but this name was never published.
There is also an enigma regarding the proper type collection of this species at PI, the only material found
by Pichi-Sermolli and Bizzarri (2005) and consisting of a sterile frond, lacking the rhizome. Without doubt, it
was the plant illustrated in Raddi (1825: t. 29, fig. 2!). However, in the description of the species, Raddi (1825:
21) indicated “sons sparsis." Thus, it is possible that further type material will be found in other herbaria. Ser-
pocaulon hirsutulum appears to hybridize with S. menisciifolium (see below), thus blurring the interpretation of
Serpocaulon hirsutulum differs from S. menisciifolium s. str. by smaller /ronds, 45-55 cm long (vs. 50-140
cm), laminae with 9-13 pairs of lateral pinnae (vs. (20-)23-35 pairs), pinnae linear-oblong with apices round to
obtuse (vs. pinnae falcate, apices cuneate to acute) (Figs. 2A, B). In addition, S. hirsutulum has the laminae
markedly villous, with whitish lax 5-12-celled hairs, 0.5-1.2 mm long, on both sides of rachises, costae, veins,
laminar tissue between the veins, and laminar margins (Figs. 2C-F). On the other hand, S. menisciifolium has
stiff hyaline hairs with cross-walls reddish, 0.2-0.4 mm long, 3-5-celled, mainly restricted to rachises (espe-
cially adaxially), and scattered on costae; the other laminar regions of S. menisciifolium are totally glabrous.
The putative hybrids exhibit intermediate morphology, and since they have been formally described, we
combine the epithet in Serpocaulon.
’.nisciifoliun
4. Serpocaulon xpubescens (Rosenst.) Schwartsb. & A.R. Sm., comb, et stat. nov. Basionyi
Langsd. & Fisch. var. pubescens Rosenst., Repert. Spec. Nov. Regni Veg. 21:348. 1923. Type: BRAZIL. SAo Paulo: Moito Jaragua, uo
Dec 1912, A.C. Brade 5384 (iectotype, designated here: HBl-39800).
Distribution and ecology. — Epiphytically on the Coastal to Montane Forests of the Brazilian Atlantic Forest, in
the States of Rio de Janeiro, Minas Gerais, and Sao Paulo. Probably, it also occurs in the States of Espirito Santo
and Parana. Rosenstock (1925) listed it also to Rio Grande do Sul.
Additional specimens: BRAZIL. Rio de Jan
Sul?l, 04 Mar 1902, Gerdes 50 (S*-059227).
►: Regiao
3. Schwartsburd et al. 2650 (UC!, >
Serpocaulon xpubescens is a putative hybrid between S. menisciifolium and 5
everywhere both species are present. Although many sporangia and spores
some well-formed ones. Whether or not this putative hybrid can reproduce on its own is unknown.
Serpocaulon xpubescens presents an intermediate morphology between the two parents: it has the laminar
architecture (especially the ascending, falcate pinnae, with cuneate to acute apices) of 5. menisciifolium-, an inter-
mediate number of pinnae (9-20(-25) pairs); indument similar to S. hirsutulum on rachises and costae, but with
hairs much shorter and sparse on other laminar regions (hairs 0.2-0.4 mm long, 2-4-celled, sparse on both
sides of costae and veins, scattered on both sides of the laminar tissue between the veins and laminar margins).
I, lanceolate scales present on costae abaxially, and calcareous dots on the hydathodes
Schwartsburd and Smith, Novelties in Serpocaulon
L (C. Presl) Schwartsb. & A.R. Sm., comb. nov. Basionym: Mar^naria
num Raddi, Opusc. Sci. Bol. 3:287. 1819, nom. illeg. (non Salisb. 1
d 6:231. 1854 [1855?!, as “(Raddi) J. !
i) 1(2):523. 1870. GoniophleWum sehm. -
;. Senn.) Labiak & J. Prado, An
LIA: [in nemoribus aiiquamum uais auo> ...rellensium], [Nov 1817-May 1
by Pichi SermolU & Bizzarri 2005: 108: PI* [spec. 21; isoi£CTOTypes: BR*-697793; P*-0063:
we consider Presl’s (1836) Marpnarin la«a io be a nomm novam the illegitimaje
anH therefore the earliest basionym available in Serpocaulon for this species (ICBN Art. 58.1, Ex. 1). Charge
terization of this species can be found in Sehnem (1970). Pichi Sermolli and Bizzarri (2005), Labiak and Prado
;SoaI^ire.aU2010),respectively,asPo.,,»^^^^
mii, and S. sehnemil
THE DIVERSITY OF SERPOCAULON IN BRAZIL
Adding.heabovenoveUles.orecen,mforn,ation(Hense„1990;Smuheul.2m^biak&P«do^
&LabL 20)9; Labiak&Hirai2010), the Brazilian Alhntic Forest (soulhe,^»utheasternB«zilpteMu.h
em Bahia) contains 12 known species of Serpocaulon, plus a putative hybrid between ^
endemics to sub-formationsof the AtUntic Forest (iglondulosissimumlB^^^
(Copel.)A.R.Sm. from the highland elhn forests; andS-rexlromthe M^^^^
the percentage of the species richness (ca. 25%) and the num er o
latum, S. caceresu (Sodiro) A^R. Sm. (also ^ attenuatum (Willd.) A.R. Sm. Serpo-
(Klotzsch) A.R. Sm., S. sessih/ohum (Desv.) ' , ^ 2006; Zuquimet al. 2008; Labiak&Hirai
caulonjraxinifohum and S. msena e a so aanatum (Klotzsch) A.R. Sm. are better referred to S.
2010; Carvalho etal. 2012). Brazilian plants ascribed to i. a
richardii; S. adnatum occurs in western Venezuela, Co om la, an cua
.„totaf,iroriaspeciesa„dahyhridof^»^—
Of the total in the genus. Because of undescribed taxa. A new monographic revision is required
within the S. catharinae-latipes complex.
Journal of the Botanical Research Institute of Texas 7(1)
We thank Reinaldo A. Pinto for preparing the illustrations; George Cremers (CAY, P), Michel Boudrie (CAY),
and Germinal Rouhan (P) for confirming the identity of the type of Polypodium giganteum at P; Giovani Morales
(HB) for sending images and materials from HB; Leonardo Cocchi (PI) for sending images of the type of Serpo-
caulon hirsutulum; Markus Gastauer (VIC) for helping with German translation; the curators and staff of HB,
SJRP, and VIC; and the two reviewers for constructive comments (Fernando B. Matos and one anonymous).
Pedro B. Schwartsburd also thanks FUNARBEAJFV/Floresta-Escola, SECTES/HidroEx/FAPEMIG for
grants and sponsoring; Simone Ferreira da Silva, Marina Bond Schwartsburd, Markus Gastauer, and Lassi M.
Alves (in memoriam) for helping in the field expeditions; Joao A. Meira Neto (VIC) and Ana P. Santos-Gon-
galves (VIC) for the invitation to participate in the project ‘Floresta Escola’.
Assis, E.LM. AND P.H. Labak. 2009. Polypodiaceae da borda oeste do Pantanal sul-matogrossense, Brasil. Revista Bras. Bot
32(2):233-247.
Baker, J.G. 1 870. Cyatheaceae et Polypodiaceae. In: C.F.P. Martius and A.G. Eichler, eds. Flora Brasiliensis. Monachii, Lip-
siae: Fleischer in Comm. 1 {2):306-624.
Brade, A.C. 1951. Filices novae Brasilienses VII. Arq. Jard. Bot. (Rio de Janeiro) 1 1:21-36.
aRVALHo, F.A., A. Sauno, and C.E Zartman. 201 2. New country and regional records from the Brazilian side of Neblina Masif.
Amer. Fern. J. 1 02(3):228-232.
Christ, H. 1906. Primitiae Rorae Costaricensis, Rlices IV. Bull. Herb. Boissier, ser. 2, 6:45-58.
FtE, A.L.A. 1 869. Cryptogames vasculaires (fougeres, lycopodiacees, hydropteridees, equisetacees) du Bresil. J.B. Bailliere
et Fils Libraires, Paris.
FCe, A.LA. 1873. Cryptogames vasculaires (fougeres, lycopodiac^s, hydropterid^s, equis^tac^s) du Brasil, lie partie:
supplement et revision. J.B. Bailliere et Fils Ubraires, Paris.
V. 50(3-4):279-336.
JStor Punt Science websre. Available at http://plants.istor.org/. Accessed Aug-Sep 201 2.
Labiak, P.H. AND R.Y. Hirai. 2010. Polypodiaceae. In: Forzza, R.C. (and 30 others) (orgs.). Lista de esp4cies da flora do Brasil.
Jardim Botanico do Rio de Janeiro, Rio de Janeiro. Pp. 547-552.
Labiak, P.H. and J. Prado. 2008. New combinations in Serpocaulon and a provisional key for the Atlantic Forest species.
Amer. Fern J.98(3):1 39-1 59.
Lelunger, D.B. 1985. Nomenclatural and taxonomic notes on the pteridophytes of Costa Rica, Panama, and Colombia, It.
Proc Biol. Soc. Wash. 98(2):366-390.
Lelunger, D.B. 1989. The ferns and fern-allies of Costa Rica, Panama, and the Choc6 (Part 1: Psilotaceae through Dickso-
niaceae). Pteridologia 2A:1-364.
McNeilu j. (and 1 1 others) (eds.). 2006. International code of botanical nomenclature (Vienna Code). Regnum Veg. 146.
A.R.G. GantnerVerlag KG.
Moore, T. 1 86Z li^ex filicum: a synopsis, with characters, of the genera, and an enumeration of the species of ferns, with
Schwartsburd and Smith, Novelties in Serpocaulon
Moran, R.C. 1995. U. Polypodium L In: R.C. Moran and R. Riba, eds. Psilotaceae a Salviniaceae. In: G. Davidse, S.M. Souza
and S. Knapp. Flora Mesoamericana. Universidad Nacional Autonoma de Mexico. 1 :349-365.
PicHi Sermolli, R.E.G. and M.P. Bizzarre 2005. A revision of Raddi's pteridological collection from Brazil (1817-1818). Webbia
Prado, J., R.Y. Hirai, and P.B. Schwartsburd. 2010. Criptogamos do Parque Estadual Fontes do Ipiranga, Sao Paulo, SP. Pteri-
dophyta: 9. Grammitidaceae e 1 6. Polypodiaceae. Hoehnea 37(3):445-460.
Presl, C. 1836.Tentamen Pteridographia
nem exposita. Typis Filioru
Raddi, G. 1 825. Plantarum Brasilensium nova genera et species novae, vel minus cognita
Aloisii Pezzati, Florentiae.
;. Pars I (Filices). ExTypographia
Rosenstock, E. 1 925. Filices novae a Cl. A. C. Brade in Brasilia collectae. Repert. Spec. Nov. Regni Veg. 21 :343-349.
Sehnem, a. 1970. Polipodiaceas. In: R. Reitz, R., ed. Flora llustrada Catarinense. Herbario Barbosa Rodrigues, Itajai.
Smith, A.R., H.-P. Kreier, C.H. Haufler.T.A. Ranker, and H. Schneider. 2006. Serpocaulon (Polypodiaceae), a new genus segre-
gated from Polypodium. Taxon 55(4):91 9-930.
Swedish Museum of Natural History website. Available at: http-y/www.nrm.se/botany/krypto-s. Accessed on Aug-Sep 2012.
ZuQuiM, G., F.R.C. Costa, J. Prado, and H.Tuomisto. 2008. Guia de samambaias e licofitas da REBIO Uatuma - Amazonia
Central (Guide to the ferns and lycophytes of REBIO Uatuma - Central Amazonia). Attema Design Editorial, Manaus.
Journal of the Botanical Research Institute of Texas 7(1)
BOOK REVIEW
Adam Cross. 2012. Aldrovanda: The Waterwheel Plant. (ISBN: 978-1-908787-04-0, cloth). Redfern Natural
History Productions, 61 Lake Drive, Hamworthy, Poole, Dorset BH15 4LR, ENGLAND, U.K. (Orders:
www.redfemnaturalhistory.com, sales@redfemnaturalhistory.com, +44 1202 686585). $29.72, 249 pp.,
93 color photos, 6" x 9".
Most of us who have been around for a while probably grew up with the old adage, “You can’t judge a book by
its cover!” This reviewer says, “Oh, yes you can! — ^At least, a lot of the time!”
1 was totally fascinated by the cover of this beautiful volume when 1 first saw it. Even more exciting, as I
first thumbed through the pages, were the many color photographs and the careful, well-researched history of
this unusual, one-of-a-kind plant.
Best of all, however, the author has provided us with an incredibly fascinating and carefully researched
account of this monotypic genus and its only species, Aldrovanda vesiculosa, a Tertiary relict with an ancient
lineage dating back “tens of millions of years.” Cross has not only researched all the ancient information, he
has provided a thorough presentation of Paleo-History; Morphology, Physiology and Ecology; and Habitat and
Distribution, including all the continents in which Aldrovanda was and now is found. He also identifies areas
where populations of Aldrovanda are currently “introduced and artificial” and discusses population genetics;
conservation status, protection and national listings; integrated conservation and management guidelines;
and cultivation.
As the author points out, “Despite being widely regarded as one of the most difficult carnivorous plants to
maintain in horticulture, Aldrovanda is not as capricious as its reputation might suggest.” He also offers advice
on how to proceed with Aldrovanda, including the nurturing, protecting, and dealing with “pests, diseases,
and predators.”
This is a very careful, very well-researched account of Aldrovanda. The history and work of many promi-
nent botanists through the centuries is fascinating, and the volume is concise, to the point, thoroughly ex-
plained, beautifully illustrated, and very well documented. Most dedicated gardeners and curious, causal gar-
deners will enjoy “trying their hands” with this plant. (And the author does tell you specifically how to get
started at home!)— Helen Jeudc, Volunteer & Associate Editor, Botanical Research Institute of Texas Fort Worth,
Texas, U.SA.
i-lmt Teas 7{1): 94. 2813
TWO NEW SPECIES FROM THE BREVANTHERUM CLADE
OF SOLANUM (SOLANACEAE) FROM EASTERN BRAZIL
Leandro L. Giacomin
Lynn Bohs
Av. Antonio Carlos, 6627, 31270-901
Belo Horizonte, MG, BRAZIL
giacomin.leandro@gmail.com
Department of Biology
University of Utah
257 South 1400 East
Salt Lake City, Utah 841 12-0840, U.SA.
Joao R. Stehmann
ABSTRACT
RESUMO
INTRODUCTION
Solarium L. (Solanaceae) is among the largest genera of Howering plants and contains approximately 1500 spe-
cies with a worldwide distribution. Since 2004, it has been focus of a comprehensive species-level taxonomic
inventory aiming to provide online descriptions of all its species (the FBI Solanum project; Knapp et al. 2004;
httpy/www.solanaceaesource.org). This collaborative effort has led to the discovery of several new taxa from
different countries (e.g., Peralta et al. 2005; Granados-Tochoy & Orozco 2006; Knapp 2007; Knapp & Nee
2009; Nee et al 2006- Stem & Bohs 2009; Tepe & Bohs 2009; Knapp 2010a & b; Farmggia & Bohs 2011; Gia-
comin & Stehmann 2011- Vorontsova & Knapp 2012). The project was coincidentally developed together with
i 8ot Res. hKt Teas 7(1): 95 - 107. 2
Journal of the Botanical Research Institute of Texas 7(1)
a large-scale initiative in Brazil to achieve the Target 1 of the Global Strategy for Plant Conservation adopted by
the parties to the Convention on Biological Diversity (CBD) in 2002: the construction of a working list of the
plant species for the country (Forzza et al. 2010). The construction of the Brazilian list of Solanaceae (Stehm-
ann et al. 2010) revealed several poorly known or unknown taxa, and two of these are described here.
Previous molecular phylogenetic studies have identified the Brevantherum clade as one of the 12 to 15
main lineages of Solarium (Bohs 2005; Weese & Bohs 2007). This clade has been target of intensive studies by
the authors in order to better understand the evolution of the group and the relationships among its species. It
has a New World distribution, occurring from the southern United States through Argentina, with a center of
diversity in Brazil. It comprises unarmed herbs to shrubs with unbranched to variously branched and stellate
trichomes encompassing at least six sections treated in past classification systems. Molecular data have identi-
fied two subclades within the Brevantherum clade (see Weese & Bohs 2007 for more details). The first subclade
composes members of S. sect. Gonatotrichum Bitter, which are herbs to shrubs, mainly with an indument of
unbranched trichomes (except for S. Ugnescens Femald), and are easily characterized by the explosive dehis-
cence of their fruits (Stem & Bohs 2012; Stem et al. 2013). The other subclade contains species from five sec-
tions treated in past classifications (sects. Brevantherum Seithe, Cemuum Carvalho & Sheph., Extensum D’Arcy,
Lepidotum Seithe and Stellatigeminatum Child), and all share branched to stellate or peltate hairs as a main
character. Molecular phylogenies based on three gene regions (nuclear ITS and GBSSI and plastid tmT-F) indi-
cate that Solanum sect. Brevantherum is largely monophyletic (L. Giacomin et al., unpub. data), but the mono-
phyly of the other four sections has not been established and the relationships among them are uncertain. The
limits of these sections are not well demarcated and they are in dire need of a revision, as pointed out by other
authors (Child 1998; Nee 1999).
Although most recent classification systems place species with lepidote scales in Solanum sect. Lepidotum
and species with large leaves and usually bristly-pubescent or scaly inflorescence axes in sect. Cemuum, there
is no similar consensus about the characters defining S. sects. Extensum and Stellatigeminatum (Child 1998;
Nee 1999). Child (1998) proposed that the concept of S. sect. Extensum should be applied only to climbing or
sprawling species from Central America with somewhat foliaceous calyces and plurifoliate sympodia even
though the protologue for S. sect. Extensum did not list the sympodial stmcture as diagnostic (DArcy 1972).
The species from South America with foliaceous calyces (a diagnostic character cited by DArcy, 1972), but with
difohate-geminate sympodial units were placed in S. sect. Stellatigeminatum by Child (1998) Nee (1999) dis-
agreed with this proposition and, like DArcy (1972), included the species with foliaceous accrescent calyces
and fasciculate axillary inllorescences in sect. Extensum. The entire Brevantherum clade is currently under
investigation by the authors in order to better understand its infrageneric groupings
While sampling the species of the Brevanthemm clade from Brazil for phylogenetic studies an unde-
scnbed species was revealed from Espi'rito Santo state. Also, while reevaluating the species limits of Sofonum
megalochiton Mart, from the northeast, a new segregate species was discovered. The new species do not have
lepidote scales or scaly inflorescence axes, and are expected to belong to the undefined mixed clade to which
the type species of sections Extensum and Stellatigeminatum are assigned.
MaKrial fram , he following herbaria was reviewed (acronyms from Index Herbariorum. hupV/sweelgum.
HST R 1 PEUFR, RB and UFP, as well as the non-indexed
nes and Crilma (lUCN 2010) and for the extent of occurrence calculation a mtalmum convex^^lJ^n^CB
was b^d on Roe (1971) and Mentz et al. (2000). The delimitation of the new taxa was based on a morpW-
:t 1995).
Taxonomic Treatment
nov (Figs. 1, 2A-F, 3A-D). T
BHCB[BHCB0027301;ls
Shnibs to small trees up to 5 m, usually with a prominent trunk, branching in the upper stems, these spread-
ing. Young stems moderately to densely hirsute to hispid with few-rayed (4-6) porrect-stellate trichomes, these
sometimes multangulate, on multiseriate stalks 0.3-1. 2 mm long, the rays 0.2-0.8 mm, usually with a
2-3-celled central ray longer than the 1-celled lateral ones; old growth glabrescent to pubescent, usually with
broken trichomes comprising a hispid vestiture of multiseriate stalk remnants. Bark of older stems yellowish-
white, usually shiny; new growth whitish-green to dark purple near the insertion of the inflorescence, always
shiny. Sympodial units usually trifoliate, rarely di- or plurifoliate, often with solitary leaves; if geminate, leaves
paired only at branching points and differing mainly in size, the smaller up to the size of the larger ones.
Leaves simple, the blades 1.7-11.8(-13) cm long, 1.2-4.8 cm wide, narrowly elliptic to ovate-elliptic, membra-
naceous, weakly discolorous (dark shiny green above, pale green beneath when fresh), the abaxial surfaces
moderately to densely hirsute, the lamina always visible, with short- (more often on the lamina) to long-stalked
(more often on primary and secondary veins) porrect-stellate trichomes with 3-5 lateral rays supported by a
multiseriate stalk 0.3-0.9(-1.4) mm and a 2-3 celled central ray somewhat longer than the lateral ones (Fig.
2B), 0.8-1.2 mm long, the adaxial surfaces moderately to densely hirsute, the lamina always visible, with
mixed hyaline unbranched trichomes (Fig. 2D) up to 1.8 mm long, and sessile (more often on the lamina) to
short-stalked (more often on primary am
few (1-4; Fig. 2C) lateral rays and a 2-3-celled c
supported by a multiseriate stalk up to 0.4 mm long; venation camptodromous, impressed or not above (barely
visible in dried material), raised beneath, the secondary veins 5-9 pairs; base rounded to cuneate, asymmetric;
margins entire, slightly revolute; apex acute to long attenuate-caudate (only acute on the smaller geminate
leaves); petioles 1.8-5.5 mm long, densely hirsute, with the same trichomes as those of the abaxial midrib. In-
florescence an elongated unbranched cyme, terminal, lateral, intemodal, or subopposite the leaves, when in-
temodal, changing the orientation of the branch (see comments; Figs. 1, 3A-B), the axes (0.35-)0.9-4.3 cm
long, with (3-)6-15 flowers, subsessile to pedunculate, the peduncles 0.1-1.47 cm long, moderately hirsute
with porrect-stellate trichomes like those of the stems, the rachis pale green, densely pubescent with three
types of trichomes: hyaline, unbranched, 2-4-celled, eglandular or glandular-capitate trichomes, sessile to
short-stalked multangulate-stellate trichomes with few (1-4) lateral rays or only the 2-4-ceIled, sometimes
glandular-capitate central ray developed, this much longer than the^oAers, and supported by a inultismate
to 0.9 mm long; pedicek 0.3-L2 cm long in flower, 1.9-2.5 cm in fruit, pale green to dark pink to purplish at
base, articulated at base, 1 mm in diameter at base, densely pubescent with the same unbranched trichomes
found on the rachis, almost all with a glandular head, up to 2 mm long, mixed with sessile to short-stalked
multangulate trichomes with 1-4 lateral rays and a longer central ray. Buds slightly elongated with developed
calyx lobes, the corolla nearly included even when fully developed. Flowers perfect, 5-merous. Calyx tube 1-2
mm long, with lanceolate to narrow-ovate lobes 6-8 mm long, 3-4 mm wide in flower, the apices acuminate,
lobes broadly ovate to cordate, 8.6-12 mm long, 8-10 mm wide, the apices
ehairswith
acute, densely covered abaxi
= 5(»|im;E-J=100|ini.
Journal of the Botanical Research Institute of Texas 7(1)
Giacomin et al., New species of Solanum from Brazil
shorter stalks and the central ray always longer than the others and often glandular, adaxially with unbranched
2-4-celled (sometimes with a poorly developed secondary ray) glandular capitate trichomes up to 2 mm long
(Fig. 2E), the gland ellipsoidal, these commonly found also on the margin, associated with sessile to subsesstle
globose dark glandular trichomes (Fig. 2F), these concentrated on the midveins and near the base. Corolla
1.3-1.7 cm in diameter, white, stellate, lobed up to to the base, the lobes 4-6 mm long, 4.5-6.5 mm wide,
deltoid to ovate, with acute, slightly involute apices, spreading at anthesis, sparsely to moderately pubescent
abaxially with the same trichome types found adaxially on the calyx and with a tuft of small unbranched eg-
landular trichomes at apex, glabrous adaxially. Filament tube up to 1 mm long, with a minute free portion less
than 1 mm long, glabrous; anthers 2.8-4 mm long, 1.3-1.5 mm wide, sometimes slightly unequal, elliptic to
oblong, connivent, yellow, glabrous on both faces, poricidal at apex, with the pores directed introrsely, opening
into longitudinal slits with age. Ovary glabrous; style 6-7 mm long, cylindrical; stigma clavate, with a papillose
surface. Fruit a globose berry 1-1.4 cm in diameter, white to pale green when immature, purple to black when
mature, shiny, glabrous, drying brown to dark, completely covered by the accrescent calyx which spreads open
at maturity. Seeds 9-31(-35) per fruit, 2.3-3.4 mm long, 1.7-2.3 mm wide, flattened, reniform, golden to
brown, the surfaces with irregular pits.
9»13.53'S, 35“52.776'W, 550-600 m, 14 May 2001 (fl, fr), W.W nomas et al. 12441 (BHCB, NY. RB). Bahia: Mun. Ibirapitanga, Far. Paineiras.
17 Feb 1998 (fl, fr), A.S. ConceifOo 213 (MG). Pernambnco: Mun. Maraial^agoa dos Gatos, Serra do Urubu, 20 Apr 1994 (fi, fr), A.M. Miran-
da & L.P. Felix 1628 (EAC, HST).
Discussion— Solanum anisocladum is a species with a robust habit, forming small trees up to 5 m tall with
prominent trunks, with the upper leaf surface shiny, dark green, and drying dark. It is similar to Solanum meg-
aloAitm which nonnally fonns much branched shrubs up lo 3 m taU. with the surface matte, often covered
by a whitish-yellow indument, and normally pale green when dried. Both species share an inflated calyx that
is strongly accrescent at fruit maturity (Fig 3D). This feature was previously used to separate S. tnegulocliilOT
from similar species such as S. didymum Dunal. However, 5. utiisoclodum presents several other characters that
readily dislinguUh it from S. megalochiton. Its indument of very long stellate and unbranched hairs reaching
more than 2 mm long and drying ferruginous (Figs. 1, 2A-B) is easily visible to the naked eye. The old growth
becomes somewhat hirsute to hispid where only the multiseriate stalks of the stellate trichomes remain. The
indument in S mcgulocbiton is much softer, reaches up to 1.5 mm long, is normally transparem to yellowish
when dried, and the stems never become hispid. The inflorescence is also a distinctively robust unbranched
cyme (Fig. 3A-B), sometimes with an axis teaching more than 4 cm long whereas the inflorescences of S.
megulocliilon are normally shorter than 2.5 cm. Its adaxial leaf indument is composed of eglandular porrect-
stellate trichomes assocUted with unbranched hairs, the latter with muUicellular bases, apparently represem-
ing stellate hairs without developed lateral rays. The adaxialleafindumentofS. megulocliilon is composed only
of stelUte hairs these sometimes glandular. Solumim utiisoclodum also has a distinctive branching pattern not
observed in S. megulochiton (Fig. 3A). When arising from between the^s, Aejnflon
Strong change in growth direction (not always visible on small brar
in every specimen seen at the field). In contrast, the flowering brai
straight.
In the Brazilian plan, species list (Stehmannetal.2010),SoIunum megulochiton was cited as occurring tn
northeastern Brazil from the states of Alagoas and Bahia. Duni
1 herbarium sheets, I
s of S. megalochiton are
rbaria in Pernambuco and Bahia no
specimens matching it were seenco-occutnngwithS.umsocl4dum.AsheetfromAlagoas(71iomusl244I;BHCB.
NY, RB) determined as S. megulochiton by other Solunum experra acmally confo^ to our concept of S. umm^
cludum. Thus,innor.heas.ernBraz.l,S. megulochiton isknowntooccuronb-mafewoc^onsmthe^^^^^^
ofthesouthempartofBahiaandnotfurthernorthward.Solonummegooc ttoncana oun m
aKsoud,eas.ernandsouUremregionsofBrazil.wi.halewaddi.ionalcollecnonsm.hesrateofGo^andMa.o
Grosso. This suggests that Sunisocludum and S.megukxhit™ are not sympaurtc, butth.scanno.be
because not all important collections of the group
(e.g.,JPB).
tog pan
ingi
in unequal branches when the infloresce
Distribution and habitat. — Solanum anis
i from between the nodes.
>ccurs in forest edges of the Atlantic coastal rainforest of
northeastern Brazil in the states of Alagoas, Bahia, and Pernambuco, in partial to complete shade, at elevations
of 500-700 m (Fig. 4). Primary wet forest fragments at this elevation are quite rare in northeastern Brazil today,
and the type collection, despite the fact that it was found in a forest edge, was from a well preserved fragment
Phenology. — ^The species was collected with flowers and fruits from February through May, but a flower-
ing peak was observed in February and March, and fully developed fruits were found from March through
May.
only three localities and its calculated extent of occurrence using the MCP is 834 km^. Despite the fact that the
Bahia registry is far from the one in Pernambuco state, the MCP reflects the narrow range of coastal rainforeS
where the species is expected to occur. Although the type locality is now part of a private reserve owned by a
sugar company (RPPN Frei Caneca), it consists of a small group of forest fragments in a severely damaged
landscape mainly modified by extensive agriculture based on monoculture (plantations). Generally, the Atlan-
tic coastal rainforest has been severely damaged in northeastern Brazil in the past decades due to urban expan-
sion and extensive farming.
Shrubs, up to 3 m, sometimes scandent, much branched, the apical branches scandent, held downwards, arch-
ing. Young stems densely pubescent with sessile to short-stalked porrect-stellate trichomes normally with 8
lateral rays, rarely fewer, commonly of unequal length, up to 0.3 mm long, with a central ray much shorter than
the lateral ones (Fig. 2G); old growth sparsely pubescent to glabrescent. Bark of older stems bright white to
yellowish, shiny; new growth gray due to dense vestiture, with a white to dark purple lamina beneath. Sympo-
dial units 2-3 foliate, geminate, with paired leaves differing in size and shape, the smaller ones less than of
the size of the larger. Leaves simple, the blades 2.2-7 cm long, 1-3 cm wide, elliptic to broadly elliptic (the
smaller geminate leaves broadly elliptic to ovate, sometimes slightly obovate), membranaceous to chartaceous,
slightly discolorous, drying dark adaxially, the abaxial surfaces sparsely to moderately pubescent, the pubes-
cence denser on veins, with sessile to short-stalked porrect-stellate trichomes with 4-9 lateral rays up to 0.3
mm long, frequently unequal in length, and a minute central ray (Fig. 2H), always shorter than the laterals, up
to 0.1 mm long, the stalks multiseriate, reaching 0.1 mm long, the adaxial surfaces glabrescent, with hairs only
; frequently more rayed), up to 0.3 mm long, frequently unequal, with a slightly
the abaxial surface (Fig. 21), reaching 0.2 mm long, but still shorter than the lateral
m camptodromous, slightly to not raised above, raised beneath, the secondary veins 4-6 pairs;
base cuneate to obtuse (rounded on the smaller geminate leaves), asymmetric; margins entire, plane- apex
densely pubescent with the same trichomes as the stem and abaxial midrib. Inflorescence a congested un-
branched cyme, lateral, intemodal, or subopposite the leaves, rarely terminal, the axis sessile to subsessile up
nm or absent, green, moderately to densely covered with the same porrect
very closely spaced; pedicels
Journal of the Botanical Research Institute of Texas 7(1)
Fk. 5. botype of Sokmum aektolm Oiacomin & Stehmann. (K Dtmmer et aL 3273, BHCB [BHCB 002734]). <
105
3-8mmlonginflower,5-9mminfruit,greenatbase>comingpurplishdistally,ar^^^^^^
lmmindiameteratbase,moderatelytodenselypubescentwithsessiletosubsessileporrect-stellatet
with up to 8 lateral rays, the central ray much shorter than the others, and sometimes with 3-4 celled umsen-
ate glaLular hairs near the calyx. Buds globose, the corolla emergent from the calyx lobes, spreadmgjust^-
foreanthesis. Flowers perfect, 5-merous. Calyx tube up tolmm long, with narrow oblong to
1 6-2 5 mm lone 0 7-1 mm wide in flower, 2.5-3 mm long, 1-1.2 mm wide in fruit, with acute to ac
apices becomingovate, accrescent, but not covering the whole mature berry,coveredabaxially with the^^^
Lhon.es found on the pedicels, as^ciated wUh unbranchM 3-^ce..e
Lhon.es barely visible in dried material, adaxially with sparse porrec-stelkte mchomes, smaller than those
ta„na’xially with 4-7 lateral rays, often the same size as the central ray (Fig 2J . assoc.amd w. h un-
LLLglanlular capitate trichomes at thebase near the ova,y,andalsow^^^^^^
6 h H or ^metimes forked sometimes glandular hairs densely covering the margins, glabrous adaxially.
Foment tube up to 0 5 Inm with a free portion reaching 0.6 mm, sometimes slightly unequal. Y"'
ameter, white to pale green when immature, purple at m ^ , hserved in mature fruits) 2 5-3
.iallycoveredby,heaccrescentcalyxthatspreadsa.maturi,ySee^few(Mo^jyed— eta^^^ -
mmlong.2-2.4mm wide, na,tened,reniform. pale brown, the surface w.th.rregularp.ts.
Landislau. 19“18'39-S.
738 (BHCB.MBML).
Discussion.-Solanum caelicolum resembles S. ^ distinguished by its
leading to misidentifications of P dabrescent and usually drying dark adaxially (nor-
elliptic leaves, not ovate like those of S. hirte ^ J ^^,^hat gray to pale green due
mally moderately pubescent and pale green in S. „„bescent and pale green in S. hirtellum). The adaxial
to the conspicuous indument (normally inconspicuous y^ caelicolum has porrect-stellate trichomes
leaf trichomes are also distinct^
with a very reduced midpoint (central ray, g- ^ surfaces. Some variation in trichome
times multicellular and longer than the ^^j^xial leaf surfaces have a central ray
morphology occurs in S. hirtellum, but at leas distinctive sessile to subsessile congested and un-
longer than the lateral ones. Solanum caelicolum a s absent),
branched inflorescence (Fig ») with the few- to many-branched, and lax, with the
whereas the inflorescence m S. hirtellum is y ^ also be used to distinguish both species; they are
pedicels arising up to 0.5 cm from each other, e hirtellum. Another good character to distin-
upto8mmlonginS.caeIicoIumandfrequentlyexce ^^Hcolum the calyx lobes are larger than the
guish the two species is the size of the fruiting calyx lobes In 5. caenc y
106
Journal of the Botanical Research Institute of Texas 7(1)
Both species are known from Espirito Santo state, but they apparently do not co-occur. While S. coeii
colum seems to be restricted to north-central Espirito Santo, S. hirtellum has a much broader distribution, ex-
tending from northeast Argentina and Paraguay and reaching its northernmost limit in the south of Espirito
Etymology.— The epithet caelicolum derives from the Latin “dweller of heaven.” It references the regionof
Santa Teresa and surroundings, where the species was first collected and is most abundant. The region holdsa
considerable amount of the preserved forest fragments of the Espirito Santo state and plays an essential role in
the conservation of the plant species of the Brazilian Atlantic Forest, a severely damaged biome due to human
occupation. For Solanum specialists as well as other botanists, it is a heaven of plant diversity and new species
Distribution and habitat.— Solanum caelicolum inhabits the understory or shaded forest edges of well-pre-
served fragments of the Brazilian Atlantic coastal rainforest, and is known only from Espirito Santo state from
150 to 850 m (Fig. 4). Despite the fact that the type collection was from an outcrop, the species is believed to
inhabit only shaded environments and the reference might be to a granitic boulder in the forest understory,a
common situation in Espirito Santo forests.
Phenology.— The species can be found in flower from January through June. Fruits were found from Feb-
ruary through June, but fully developed ones were found from March through June.
Preliminary conservation status.— Endangered B1 a,b (i, ii, iii). Solanum caelicolum is known from four lo-
calities very close to each other in the state of Espirito Santo, and its calculated extent of occurrence using the
MCP is 31 km^. At the municipality of Santa Teresa, the locality of three of the collections, there still are sev-
eral well preserved forest remnants, some of them within protected reserves. One of the collections was found
at a city reserve (Estagao Biologica de Sao Lourengo) and therefore we are not treating this species as critically
endangered. The populations from further north in the Colatina region are more susceptible to changes in the
species area of occupancy due to urban expansion and farming.
ACKNOWLEDGMENTS
We would like to thank L. Kollmann, F. Saiter, and Y. Gouvea for assistance in the field at Espirito Santo and A
J.Silva ami A. SantiagoatPemambuco;UsimFreiCaneca, for the access permit at RPPNFmCaneca; thee™-
tors and institutions of the cited herbana; M.M. Duarte for preparing the trichome line drawings- G Wahlert
J. Lombardi, L. Assis, L. Mentz and T.E. Almeida for suggestions and critical review Reviews by two anony-
mous rt^ew™ am greatly appreciate. This work was supporte by CNPq thmugh Process 479921/2010-5
and by the SISBIOTA program through Process 563342/2010-2 toJRS and by NSF through grant DEB-0316614
to LB. A scholarship was provided to LLG by CNPq (Process 148363/2010-5).
Bohs, L 2005. Major clades in Solanum based on ndhF sequence data. In; R.C Keating, V.C Hollowell tr Crnat
eds. A iestmhtdtfor «„iam G. D'AtcyThe legacy of a taxonomist. Monegt. SysF Bot. 64,33000“; mi"™
CouvEutioii on BKHoacALDwasuYiCBDI. 2002. COP 5 Decision V/10; Global strategy for plant con<»o l a ./
declsion/cop/M=7152iaccessed16June20,2) »'“'P'4''*™nsenrat,on.(w»v.cbd.int/
Chbo, a 1998. Studies in Solanum and related genera (6). t
Feddes Repert. 109:407-427.
D'Afia, W.G. 1972. Solanaceae studies II; typification o
Farrugoa, F. and L Bohs. 201 1 . Two new South American species of Solanum s
1:67-77.
Forzza, FlC. AND 22 OIHERS. 2010. Catalogo das Plantas e Fungos do Brasil, 2 vols, Anc
Botanico do Rio de Janeiro, Rio de Janeiro.
Qacomin, L, and J.R. Stehmann. 201 1 . A new heterandrous species of Solanum section
from Bahia, Brazil. Phytokeys 7:1-9.
Granados-T ochov J.C and Cl. Orozco. 2006. Una nueva especie de Solanum seccion Ge
Caldasia 28:1-8.
' (Solanaceae) de Colombia.
107
8.0. Prepared by the Standards and Petitions Subcommittee in March 2010. http://intranet.iucn.org/webfiles/doc/
SSC/RedList/RedListGuidelines.pdf.
Knapp, S. 2007. Solanum coalitum (Solanaceae), a new enden
Knapp, S. 2010a. Four new vining species of Solanum (Duka
PLoS One 5:e1 0502.
Knapp, S. 2010b. New species of Solanum (Solanaceae) from Peru and Ecuador. Phytokeys 1:33-51.
Knapp, S., L. Bohs, M. Nee, and D.M. Spooner. 2004. Solanaceae: a model for linking genomics and biodiversity. Comp. Funct.
Gen. 5:285-291. ^ ^ ^ ^
unusual anther morphology. Novon 19:178-181.
Mallet, J. 1 995. A species definition for the modern synthesis. Trends Ecol. Evol. 1 0:294-299.
Mentz, LA., P.L Oliveira, AND M.V.DA Silva. 2000. Tipologia dos tricomas d
Regiao Sul do Brasil. Iheringia, Serie Bot. 54:75-106.
Nee, M. 1 999. Synopsis of Solanum in the New World. In: M. Nee, D.E. Symon, R.N. Lester, and J.R. Jessop, eds. Solanaceae
IV: Advances in Biology and Utilization, Royal Botanic Gardens, Kew. Pp. 285-333.
Nee, M., L. Bohs, and S. Knapp. 2006. New species of Solanum and Capsicum (Solanaceae) from Bolivia, with clarification of
nomenclature in some Bolivian Solanaceae. Brittonia 58:322-356.
Peralta, I.E., D.M. Spooner, and S. Knapp. 2005 New species of wild tomat(
from northern Peru. Syst. Bot. 30:424-434.
Roe, K.E. 1971. Terminology of hairs in the genus So/anum. Taxon 20:501-508.
Roe K E 1972 a revision of Solanum section Brevantherum (Solanaceae). Brittonia 24:239-278.
Stehmann, J.R., la. Mentz, M.F. Agra, M. Vignoli-Silva, and LL Giacomin. 201 0. Solanaceae. In: R.C. Forzza (and 22 others), eds.
Catalogo das Plantas e Fungos do Brasil, 2 vols, Jardim Botanicc
Stern, S.R. AND L. Bohs. 2009. 1
Pachyphylla (Solanaceae). J
Stern, S.R. and L. Bohs. 2012. /
s (Solanum section Lycopersicum: Solanaceae)
ode Janeiro, Rio de Janeiro. Pp. 1633-1649.
(Solanaceae). PhytoKeys 8:89-98.
Stern, S.R., L Bohs, LL. Giacomin, J.R. Stehmann,
38:471-496.
Tepe, EJ. and L. Bohs. 2009. Three new specie
L. Gonatotrichum. Syst. Bot.
eggplant. PhytoKeys 8:1-1
2013. A revision of Solanum S(
) section Herpystichum (Solanaceae). J. Bot. Res. Inst
201 2. A new species of Solanum (Solanaceae) from South Africa related to the cull
(Solanaceae). Syst. Bot. 32:445-463.
Journal of the Botanical Research Institute of Texas 7|1)
BOOK REVIEW
Timothy A. Block and Ann Fowler Rhoads. 2011. Aquatic Plants of PennsyKania: A Complete Reference
Guide. (ISBN; 978-0-8122-4306-2, cloth). University of Pennsylvania Press, Hopkins Fulfillment Ser-
vices, P.O. Box 50370, Hampden Station, Baltimore, Maryland 21211, U.S.A. (Orders: www.upenn.edu/
pennpress, 1-800-537-5487, 1-410-516-6998 fax). $59.95, 320 pp., 93 color illus., 508 b/w illus., 7" x 10'.
Most flower enthusiasts love the plants they know and spend hours tending to their personal gardens and
yards. Many spend time helping their local herbarium and surroundings. It’s a Labor of Love, and we are more
than willing to help in any ways we can. The authors of this volume carefully explain the “Purpose of the Book
in the opening paragraph of the Preface. In very short time, we are in Chapter 1. Evolution and Ecology— and
we are totally “hooked.” {Fascination is probably a more accurate — and scholarly! — designation.) The resultis
the same. We are definitely intrigued by this book. Your hubby or wife calls to you, asking when dinner will be
ready. The engrossed reader responds with “Uh, Honey, there’s a big frozen pizza in the ‘fridge’— put it in the
oven and we’ll have that for supper!” (I wonder how many of our readers will immediately identify with that
While many of us have probably seen a few water lilies and bulrushes and perhaps curly pondweed, most
of us are not at all familiar with the plants, their habits, and how they actually grow. In addition to the usual
descriptions of the various plants, the authors utilize line drawings, color photographs, and a drawing of the
state of Pennsylvania, with a red line indicating the route being examined and a “fill in” of dark blue dots to
indicate the presence of lakes. Such maps are also used to show the routes and places in Pennsylvania where
specific plants were located.
They also explain the types of ecosystems, the management of aquatic ecosystems (endangered, threat-
ened, rare, undetermined candidate), or watch list by the Pennsylvania Natural Heritage Program (at that
time). Table 1.2 lists the Endangered, Threatened, and Rare Aquatic Plants of Pennsylvania. In addition, the
authors created a table of Non-native, Invasive Aquatic Plants in Pennsylvania as well as a proliferation of “over-
abundant native species.”
In Chapter 2, the authors have created Identification Keys for the Aquatic Plants of Pennsylvania. They
explain the system they have devised and how to use the keys they have developed. In addition, they include
line drawings in areas that might be more difficult to visualize. The keys are designed to be as helpful as pos-
sible. The authors really did a great job. These keys may not appear to be the keys you’ve been accustomed to
using, but they do a terrific “either/or” approach which probably works a lot better for this type of identifica-
The balance of the volume describes the various types of plants, their family, the species, accompanying
line drawings of the plant described, and the map of Pennsylvania showing the route, the locations, and some-
times an accompanying comment.
This volume is fun to peruse, beautiful to see the color photos, easy to use the keys, easy to understand the
line drawings, and also creates an urgent desire in the reader to go immediately to Pennsylvania and travel
those map routes!— Helen Jeude, Volunteer & Associate Editor, Botanical Research Institute of Texas, Fort Worth,
Texas, USA.
NEW SPECIES OF APHELANDRA (ACANTHACEAE)
from PERU AND ECUADOR
Dieter CWasshausen
Department of Botany
f h ui230sDecleswmcnextendsfron.norlhwes.€mMexicosou.hcastwarf
Aphelandra, a neotropical genus o* a o f Catarina in southeastern Brazil and to the
through regions of 1x>th wet and
province of Salta in southwestern Argent . rpaches its zenith in the Andean mountains of
thenumber of species and inmorphological diversity described from Colombia, 41 species
western South America and in Brazil. Presently, 61 ^P““= “™^A,^^haceae, they are found to ^
from Ecuador, 38 species from Peru and 44 ^ ^ ^ widespread, in undisturbed forests and
extremely local distribution and range, only 15% are found
are rarely forests and are rarely collected in being devoid of cystoliths, the familiar
The genus is distinguished from ot er ^ ^ ^dh narrow, 1-thecous
character by which most acanthaceous p ^^ 3 longitudinal clefts, each of which often possess one
anthers and by having ellipsoidal pol en g ^^oluding the bracts and bractlets, and in certain species
pore. Its flowering spikes are often large a wy species, are the
variegated or colored leaves occur. Important ^ otherwise, on the margins of the leaf blades
presence or absence of spiny intepetiolar racts,o tee
orflowerbracts, and of ocelli on
The nearest relative is probably the Old g whereas in
rica and Asia. In Crossandra, the upper corolla lip is absent
Aphelandratheupperlipalwayspresent. ^ Maynas: Rio A^npiyacu. Pebas and
Aphelandra plowmanuWasshausen,spjiov(F^.J^^^
vicinity,3n0'S,7r49W,10Aprl977. 0 ^
Unbranched shrub or treelet 1.5-2 m tall stem short-petiolate, the petioles 5-40
densely hirsute, the trichomes appres^d, 1 _ J ^ acuminate at apex, gradually nar-
mmlong, hirsute, the blades dtcurrent atbase, the veins (12-14 pairs) puberulous,
rowed from the middle or slightly above the mid both sparingly and
prominent beneath, less so above, the upper nndnlate. Inflorescence a solitary, terminal, sessile
inconspicuously puberulous, the . (excluding corollas), moderately dense, the rachis spar-
m subsessile spike 12-19 cm long and ^5 cm
ingly pubescent with appressed “ 33_43 long, 17-18 mm wide, acute to rounded at
axis; bracts bright orange, recurving, broadly elliptic, 33-43
apex, attenuate basally, glabrous, rather firm, distinctly parallel veined, margin entire; bractlets elliptic to ob-
lanceolate, 20-22 mm long, 3-3.5 mm wide, glabrous, acute and reticulately veined apically; calyx deeply
lobed, 17-20 mm long, the lobes lanceolate, the posterior segment larger, 20 mm long and 4 mm wide, some-
what erose at apex, the anterior pair 19-20 mm long and 3.5 mm wide, the lateral pair 17 mm long and 3 mm
wide, all glabrous and subhyaline; corolla white tinged with magenta on lower lip or the tube carmine red, lip
yellow with two dark-red spots at throat, 80 mm long, tomentose, especially on the upper part of the tube,
throat and upper lip, the tube curved, 2.5 mm wide at base then gradually enlarged to 7 mm at throat, the upper
lip erect, ovate, 22-25 mm long and 3.5-4 mm wide, entire and cucullate at apex, lower lip spreading, 3-lobed,
the middle lobe obovate, 14 mm long and 1.75-2 mm wide, truncate at tip, the lateral lobes obovate 2 mm long
and wide; stamens slightly exserted beyond the mouth of the corolla tube; anthers unequal, posterior thecae
3.5 mm long, anterior thecae 6 mm long; ovary minutely punctate. Capsule not seen.
Aphelandra plowmanii is morphologically similar to A. crispata Leonard emend. Wasshausen from Ama-
zonian Colombia and Ecuador. The new species is distinguished by having leaf margins entire or slightly un-
dulate, bracts bright orange and recurving and corollas that are either white tinged with magenta on lower lip
or tube carmin red, lower lip yellow with two dark-red spots at throat. In contrast, in A. crispata the lateral
veins terminating in broad teeth 1-5 mm high and 2-2.5 cm apart, the bracts are red to purplish, crisped or
twisted but not recurved and corollas are pinkish-purple to violet.
Distribution. — ^Along trails and in mature upland forest on well-drained alluvial soil, 130-ca 600 m
Aphelandra knappii Wasshausen, sp. nov. (Fig. 2A-H). Type: p
Herb or shrub 1-4 m tall; stems terete to subquadrangular, glabrous, drying dark-purplish. Leaves long-petio-
late, the petioles 3-5 cm long, glabrous, drying dark-purplish, the blades broadly elliptic to ovate, 15-20 cm
long, 6-10 cm wide, acuminate at apex, narrowed and attenuate at base, glabrous, thin, the veins (10-12 pairs)
glabrous, conspicuous on both the upper and lower surface, the margin entire and slightly undulate. Inflores-
cence of 3-5 short, axillary and terminal spikes, these few-flowered and congested at the tips of the branches,
the peduncles 3-4 cm long, glabrous, drying dark-purplish; bracts obovate, somewhat asymmetrical, 12-15
mm long and 8-10 mm wide, rather thin, glabrous, obtuse or rounded at apex, vernation prominent, margin
entire; bractlets oblong to lanceolate, 4 mm long and 1 mm wide, minutely ciliolate at tip; calyx 4-4.5 mm long,
deeply lobed, lobes subequal, posterior lobe oblong, 4-4.5 mm long and 2 mm wide, truncate at apex, anterior
pair lanceolate, 5 mm long and 1.5 mm wide, the lateral pair 5 mm long and 1 mm wide; corollas yellow with
lobes apically green-tipped, 40 mm long, tube glabrous without and tomentose within at insertion of filaments
base, the tube suberect, 36 mm long, 1 mm wide at base, this to 5 mm above base, then gradually enlarged to
7-9 mm at mouth, the lips spreading, the upper lip 5-6 mm long, the 2 lobes 4 mm long and 3 mm wide,
rounded, the lower lip 3-lobed, the middle lobe 4 mm long, 3.5 mm wide, the lateral lobes 3.5 mm long and 3.5
mm wide; stamens exserted 8-10 mm beyond the mouth of corolla tube; filaments 40 mm long, tomentose at
base; anther thecae 5 mm long, dorsally tomentose; pollen white; ovary 2 mm high; style 55 mm long, exserted.
Capsule clavate, 15 mm long, 5 mm wide, blunt at tip, nitid, glabrous; seeds dark brown, flat, subcircular, 4 mm
in diameter, sparingly papillose.
Aphelandra knappii is morphologically similar to A.jacobinioides Lindau also found in Ecuador and Peru.
The new species is distinguished by having broadly elliptic to ovate, glabrous leaf blades, peduncles 3-4 cm
long, glabrous, these drying dark-purplish, bracts obovate, somewhat asymmetrical, 12-15 mm long, obtuse
or rounded at apex and corollas yellow, with lobes apicaUy green-tipped, 40 mm long, tube 36 mm long and
Journal of the Botanical Research Institute of Texas 7(1)
112
Fig. 2. A-F: Aphelmdra knappS {Knapp 8077). A. Habit B. Bract. C Bractlets and calyx lobes. D. Corolla :
I, New species of Aphelandra from Peru and Ecuador
113
late at apex and the corollas are orange to yellow, 48 mm long, the tube 42 mm long and
sand at elevations between 200-800 n
-9 mm wide at mouth. In contrast, in A. jacobinioides the leaf blades are oblong-elliptic, the pubescent pe-
mcles are 1-2 cm long,, not drying dark-purplish, the bracts a
uncateandsubmuci
3 mm wide at mouth.
Distribution.— Tropical w
aterial studied in addition to the type: PERU. San Martin: Lamas. Convento, trail to Tioyacu and Nuevo Lamas, km 68 of Tarapom-
5 Tarapoto-Yurimaguas, 6°16'S, 200-800 m, 22 Mar 1986, Knapp & Mallet 6896 (MO, US); Lamas, Convento trail to Tioyacu and
uevo Lamas (then to Rio Shanusi), km 68 of Tarapoto-Yurimaguas road, 6°16'S, 76-17^, ca. 200 m, 19 Apr 1986, Knapp & Mallet 7105 (MO,
S). Loreto: Balsapuerto, 220 m, Jan 1933, Klug 2847 (US); Balsapuerto (lower Rio Huallaga basin). 150-350 m, 28-0 Aug 1929, Killip &
nith 28617 (US).
wide, the lateral pair 0.5 mn
brous, the tube suberect, 25 n
a long and 2.5 n
Low shrub ca. 1.3 m tall; stem terete to subquadrate, glabrous, drying brownish. Leaves long-petiolate, the
petioles (unwinged portion) 1.5-2.5 cm long, glabrous to sparingly and inconspicuously puberulous, the
blades elliptic to oblong, 10.5-15 cm long, 3.5-5 cm wide, glabrous, acuminate at apex, narrowed from above
the middle and attenuate at base, firm, the veins (10-11 pairs) slightly raised and conspicuous both above and
below, glabrous, secondary veins conspicuous on lower surface, margin entire. Inflorescence of 4-5 short, axil-
lary and terminal spikes, these yellow, few-flowered and congested at the tip of branches, the peduncles 1.5-2
cm long canaliculated, glabrous; bracts yellow, narrowly obovate, somewhat asymmetrical, 10-12 mm long
and 7-8 mm wide, firm, glabrous, obscurely 2-lobed at apex, cuneate at base, margin entire; bractlets lanceo-
late to narrowly ovate, 3-3.5 mm long, 1-1.5 mm wide, striate-nerved, subhyaline, glabrous; calyx 5 mm long,
lobes lanceolate, striate-nerved, subhyaline, glabrous, the posterior lobe 2 mm wide, the anterior pair 1 mm
vide, both pairs mucronulate at apex; corolla yellow-green, 28 mm long, gla-
1 long, 2 mm wide at base, 4 mm wide at mouth, the upper hp 3-4 mm long, the
ide, rounded, the lower lip 3-lobed, the middle lobe 3 mm long and 3 mm wide,
the lateral lobes'3 mm loitg and 2 mm wide, all obtuse or rounded; stamens reaching the tip of the upper hp;
filaments tomentose at point of insertion; anther thecae 2.5 mm long; ovary 3 mm high; style 28 mm long, ex-
'‘""^hrrdtTnvT/sLi is also morphologically similar to A jncohinimties Lindau from Ecuador and Pem^
This new species is distinguished by beingalow shrub, ca.l.3mmll; stems glabrous, drying brovratsh; leaf
blades smaller, 10.5-15 cm long and 3.5-5 cm wide, firm; bracts yellow, narrowly obovate, somewhat asym-
metrical, obscurely 2-lobed at apex; bractlets glabrous at apex and corollas yellowish-green, 28 mm long, tube
25 mm long, 4 mm wide at mouth. In contrast, plants of A. jacobinioides are shrubs to 4 m tall; branches are
puberulous, not drying brownish; leaf blades are 16-20 cm long, 6-8 cm wide, membranous; bracts greenish,
broadly ovate, symmetrical, truncate and submucronate at apex; bractlets apically pilose and corollas orange to
Material studiedin addition to the type: PERU. Hnanuco: Divisoria, between Tingo Maria and Boqueron, 1500-1800m. Ferreyra 1675 (GH,
Herb orsImroTlsTi^Ihlml Lie in subqnndrangular, glabrous, drying dark reddish-brown Leaves
long-petiola.e, .he pe.ioles (unwinged portion) 1-3 cm long, glabraie to sparrngly puberulous. the blades
broadly eniplic to obovate, 11-20 cm long, 5-9 cm wide, acuminate a. apex, narroW arid attenuate at base,
glabrous, niembranous, the primary veins (9-11 pairs) glabrous, conspicuous on both the np,mr and lower
surface, the margin entire. Inflorescence of 3-« short, axillary and terminal spikes, these few-flowemd and
conges.eda.d.efipsorbranches,.hepedunclesl-3cmlong,darkreddish-browu;bracmgreen, lower broadly
115
ovate, 12-15 mm long and 12-13 mm wide, apically somewhat asymmetrically bilobed, becoming narrower to
ca. 10 mm wide toward apex, apically acute, all firm, glabrous, minutely gland-dotted, distinctly parallel
nerved, margin entire; bractlets lanceolate, somewhat shorter than the calyx lobes, 4 mm long, 1.5 mm wide,
striate-nerved, subhyaline, glabrous, minutely ciliolate at apex; calyx 4-5 mm long, the lobes lanceolate, stri-
ate-nerved, subhyaline, glabrous, the posterior lobe 1.5 mm wide, the anterior pair 1 mm wide, the lateral pair
0.5 mm wide, all minutely ciUolate at apex; corolla yellow, 30 mm long, glabrous, the tube suberect, 25 mm
long, 4 mm wide at base, 7-8 mm wide at mouth, the upper lip erect or spreading, 5 mm long, the lobes 1 mm
long and 2 mm wide, puberulous within, the lower lip 3-lobed, the middle lobe 4 mm wide, the lateral lobes 3
mm long and 3 mm wide, all obtuse or rounded; stamens reaching the tip of the upper lip; filaments tomentose
at point of insertion; anther thecae 3 mm long. Capsule green, clavate, 12-13 mm long, 4 mm wide, blunt at tip,
nitid, glabrous; seeds dark brown, flat, suborbicular, 4 mm in diameter, sparingly papillose.
Aphelandra ancayacensis is also morphologically similar to the more widespread species A. jacobinioides
from Ecuador and Peru. It is distinguished from the later species by plants being somewhat smaller herbs or
shrubs, 0.3-1. 3 m tall; stems dark reddish-brown; leaf blades elliptic to obovate; peduncles dark reddish-
brown, somewhat longer, 1-3 cm long; bracts green, 12-15 mm long and 10-13 mm wide, lower apically some-
what asymmetrically bilobed, upper apically acute, all minutely gland-dotted and corollas yellow, 30 mm long,
tube 25 mm long, 7-8 mm wide at mouth. In contrast, A. jacobinioides are plants of shrubs to 4 m tall; stems not
dark reddish-brown; leaf blades oblong-elliptic; peduncles not dark reddish-brown, shorter, 1-2 cm long;
bracts green, 10 mm long, 9 mm wide, all uniform in shape and size, truncate and submucronate at apex, lack-
ing glandular dots and corollas orange to yellow, 48 mm long, tube 42 mm long, 16 mm wide at mouth.
Distribution. — In well-drained soils, primary forest on the upper river terra
i: 23 km S of Nuevo San Martin
0 m, 5 Mar 1954. Woy tkowski 1146 (US). Pasco: I
a. 10“12’S, 325 m. 14-15 Apr 1983, D.N. Smith 3699 (MO, I
Aphelandra schunkei Wasshausen, sp. nov. (Fig. 5A-G). Type: Peru, h
Herbs or shrubs 0.4-1.5 m tall; stems subquadrangular, glabrous. Leaves petiolate, the petioles 1.5-3 cm long,
glabrate to puberulous, the trichomes sordid, the blades elliptic to broadly elliptic to oblong, 15-22(-26) cm
long, 4-7(-9) cm wide, acuminate at apex, narrowed and attenuate at base, rather firm, glabrous, shining in-
tensely olive-green, vernation prominent on lower surface, the primary veins (14-18 pair) glabrous or puberu-
lous, the trichomes appressed, the margin entire or slightly undulate. Inforescence of 2-10 short, axillary and
terminal spikes, these few-flowered and congested at the tips of the branches, the peduncles 2-2.5 cm long,
glabrous, canaliculated, bracts intensely green, narrowly obovate to ovate, 13-17(-24) mm long and 8-10(-l4)
tire; bractlets lanceolate, shorter than calyx lobes, 2.75-3 mm long, 1 mm wide, striate-nerved, subhyaline,
glabrous, ciliolate at apex; calyx 4-5 mm long, the lobes lanceolate, striate-nerved, subhyaline, glabrous, the
posterior lobe 1.5 mm wide, the anterior pair 1 mm wide, the lateral pair 0.5 mm wide, all minutely ciliolate at
apex; corolla yellow to greenish yellow, 35-40 mm long, glabrous, the tube suberect, 30-35 mm long, 4 mm
wide at base, 7-9 mm wide at mouth, the upper lip erect or spreading, 5-7 mm long, the lobes 3 mm long and
3.5 mm wide, puberulous within, the lower lip 3-lobed, the middle lobe 3.5-4 mm in diameter, the lateral lobes
3 mm long and 3.5 mm wide, all obtuse or rounded; stamens reaching the tip of the upper lip; filaments tomen-
tose at point of insertion; anther thecae 4 mm long, dorsally tomentose; ovary 3 mm high, glabrous, style 30-35
mm long, slightly exceeding the anther thecae. Capsule green, clavate, 13 mm long, 4 mm wide, blunt at tip,
nitid, glabrous; seeds dark brown, flat, suborbicular, 4 mm in diameter, sparingly papillose.
Aphelandra schunkei is morphologically similar to A. davidsonii Wasshausen also found in the Department
of Huanuco, Peru. This new species is distinguished by having larger leaf blades, 15-22(-26) cm long and
4-7(-9) cm wide; green spikes on longer peduncles 2-2.5 cm long; bracts intensely green, 13-17(-24) mm long
Wasshausen, New species of Aphelandra from Peru and Ecuador 1 1 7
Journal of the Botanical Research Institute of Texas 7(1)
a from Peru and Ecuador
119
and 8-10(-14) mm wide, acuminate at apex and corollas somewhat longer, 35-40 mm long, the tube 30-35
mm long and 7-9 mm wide at mouth. In contrast, in A. davidsonii the leaf blades are 10.5-15 cm long, 3.5-5 cm
wide; the spikes are yellow, the peduncles 1.5-2 cm long; bracts yellow, 10-12 mm long and 7-8 mm wide,
obscurely 2-lobed at apex and the corollas 28 mm long, the tube 25 mm long and 4 mm wide at mouth.
Distribution. — Primary and secondary wet forest at elevations between 350-900 m.
Herb to 0.5 m tall; stem subquadrangular, woody, glabrous, covered with warty protuberances. Leaves long-
petiolate, the petioles (unwinged portion) 2-2.5 cm long, canaliculated, glabrous, the blades oblong to ovate,
10-11.5 cm long, 4-5.5 cm wide, glabrous, acuminate at apex, narrowed and conspicuously long-attenuate at
base, membranous, the upper surface dark-olive, the lower surface pale-olive, the lateral veins rather incon-
spicuous, more prominent beneath than above, the margin entire or slightly undulate. Inflorescence of 1-3
short, axillary and terminal spikes, these few-flowered, 4.5 cm long and 2 cm wide (without spreading corol-
las), congested at tips of the branches, peduncles lacking; bracts densely imbricate, pale yellow with green ve-
nation and green apices, ovate, 15-16 mm long and 10-11 mm wide, somewhat asymmetrical, glabrous, acute
to acuminate at apex, the tip minutely apiculate, cuneate at base, 3-5-nerved, margin provided with 2 pairs of
minute dents, ciliolate; bractlets linear, subcarinate, 10 mm long, 1.5 mm vnde, aristate, striate nerved, sub-
hyaline, glabrous; calyx lobes linear, 12 mm long, aristate, striate-nerved, subhyaline, glabrous, the posterior
lobe 2.5 mm wide, the anterior pair 2 mm wide and the lateral pair 1.5 mm wide; corolla orange, 50-55 mm
long, sparingly puberulous, the tube erect, 2 mm wide at base, slightly narrowed at 5 mm above base, thence
gradually expanded to 8 mm wide at mouth, the upper lip erect, narrowly ovate, 16 mm long, 11 mm wide,
retuse at apex, the lower lip spreading, 3-lobed, the middle lobe ovate, 16 mm long, 11 mm wide, obtuse to
rounded and sparingly pilose at apex, the lateral lobes narrowly ovate, 12 mm long, 7 mm wide, obtuse or
rounded at apex; stamens exserted 10 mm beyond the mouth of the corolla tube; anther thecae 4 mm long;
ovary 4 mm long, 2 mm wide, glabrous. Capsules not seen.
Aphelandra espinosae is morphologically similar to A. guayasii Wasshausen also from Ecuador. The new
species is distinguished by being herbaceous, 0.5 m tall, having sessile spikes, glabrous bracts with margins
denticulate and corollas orange, 50-55 mm long, sparingly puberulous. In contrast, plants in A. guayasii are
suffrutescent, to 0.75 m tall, and have pedunculate spikes densely white-glandular pilose, and entire bracts and
corollas straw-colored, 55-65 mm long and sparingly pilose to glandular-pilose.
Distribution.— This species is known only from the type locality.
ACKNOWLEDGMENTS
My special thanks to Alice Tangerini for the skillfully prepared line drawings. The author further wishes to
thank Tom Daniel and Lucinda McDade for providing critical suggestions and corrections in their review of
this manuscript.
REFERENCES
Daniel, T.F. 1 991 . A revision of Aphelandra (Acanthaceae) in Mexico. Proc Calif. Acad. Sci. 47(8)253-274.
Journal of the Botanical Research Institute of Tex
!.The Acanthaceae of Colombia, II. Contr. U.S. Natl. Herb. 31(2);1 1 9-322.
McDade, la. 1'
complex (Acanthaceae). Ann. Missouri BotGard. 71:1 04-1 65.
McDade, LA.,T.F. Daniel, C.A. Kiel, and K.Vollesen. 2005. Phylogenetii
major lineages present contrasting patterns of molecular evolu
30:834-862.
Profice, S.R. 2004. Aphelandra R. Br. (Acanthaceae) novas combinacoi
Profice, S.R. 2005. Tres novos species de Aphelandra R. Br. (Acanthaceae) para o Brasil. Acta Bot. Bras. 1 9(4):769-774.
Wasshausen, D.C. 1 973. New species of Aphelandra (Acanthaceae). Phytologia 25:465-502.
Wasshausen, D.C. 1975. The genus Aphe/ondra (Acanthaceae). Smithsonian Contr. Bot. 18:1-157.
Wasshausen, D.C. 1996. New species and new combinations in Aphelandra (Acanthaceae) from Ecuador and adjacent
Peru. Nordic J. Bot. 16:389-407.
a 10:17-23.
ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES XLIII;
SINOPSIS DEL GENERO ALLOMARKGRAFIA (APOCYNACEAE;
APOCYNOIDEAE) EN COLOMBIA
J. Francisco Morales
AUomarkgrafia (Apocynaceae) es un genero de 10 especies, distribuidas desde el S de Honduras a Peru (Mo-
rales 1997, 2009), el cual pertenece a la subfamilia Apocynoideae, tribu Mesechiteae (Endress et al. 2007).
Dentrodeestegrupo,AnomarfegraiiaseencuentracercanamenterelacionadoconMesechites(e.g.,Siin6esetal
2004) y ambos generos se pueden separar del resto por sus inflorescencias cimosas y hojas sin domacios en la
superficie abaxial. AUomarkgrafia se separa de Mesechites por la presencia de cinco crestas infraestaminales que
se extienden desde la base de los aiamentos hasta la base del tubo de la corola. Adicionalmente, en AHomarfe-
medial del tubo (sobre las crestas), mientras que en Mesechites, el indumento esta restringido a los aiamentos
(Fig. D.Uncaractermorfologicoadicional que puedeayudarasepararespecimenesesterilesde ambos generos
es el hecho de que las venas terciarias en Allomarkgrcfia se encuentran dispuestas en forma mas o menos per-
pendicular al nervio central, mientras que en Mesechites la disposicion es irregular y sin un patron debnido.
Adiconalmente,enelprimer genero las venas son masconspicuasyestan dispuestas en forma mas aglomerada
que en Mesechites (Fig. 2). Otro caracter tradicionalmente utilizado ha sido la forma de la corola (infundibuli-
forme en AUomarkgrafia e hipocrateriforme en Mesechites); sin embargo, con la descripcion de A. ec^oriana
(Morales 1997) se demostro que corolas hipocrateriformes tambien estan presentes en Allomarkgraba. Algu-
nos estudios Blogen^icos recientes (e.g., Simoes et al. 2004), ban sugerido una cercana relacion entre amtes
alta variabilidad de la forma de la corola en generos de la tr—
rreducidoalasinoi
Basado en los registros de colecciones de m^ de 115 herbarios, se puede ahrmar que en general, Allomark-
un genero pobremente recolectado: de las 10 especies o
Ida extensamente (mas de 60 especimenes conocidos), mientras que
JBot Res. InsL Teas 7(1): 1
Journal of the Botanical Research Institute of Texas 7(1)
13 y 26 colecciones [A. brenesiana Woodson, A. campanulata (Markgr.) J.F. Morales, A. ovalis (Markgr.) Wood-
son) y de las cinco resiantes, 4 son endemicas y conocidas por menos de cualro colecciones (Cuadro 1). Las
especies son morfologicamente muy similares (en la forma y tamano de las hojas o foliculos y semillas) y son
necesanas flores para poder proceder con una identificacion apropiada.
Como parte del proceso de elaboracion de un tratamiento de Apocynaceae sensu stricio (subfamilias
Apocynoideae y Rauvolfioideae) para la serie Hora de Colombia, se presenta una sinopsis del genero en este
pais. Para facilitar el entendimiento de la clave, las partes florales de la corola se ilustran en la Figura 3
AVE PARA LAS ESPECIES DE ALLOMARKGRAFIA EN COLOMBIA
124
Journal of the Botanical Research Institute of Texas 7(1)
inferior del tube. b. Parte superior del tube.
5 18936, IMB). i,A.ptumeriifolki U.F. Morales 19000, INB). a. Parte
Etimologia.— El nombre de esia especie hace refei
Distribucidn. — Conocida por el momento unican
Allomarkgrafia antioquUim se puede confundir ct
porsusfloresmut
Allomarkgrafia
1 al Departamento de Antioquia en Colombia,
de la localidad tipo, a 1400 m de elevaciOn.
plumerii/lora, ya que ambas se c
grandes. El epfteto de esta especie esta dedicado al DepIrLTen to de Am^^i^a
torianaJ.F. Morales, Brittonia 49:340 1
1997. (Fig. 5). 1
'' '' emre 200-300 m
nurRgrrgia ecuatonana se puede reconocr
Morales, Sinopsis del genero Allomarkgrafia en Colombia
125
126
ffen Colombia.
Colombia (Morales 2006). ^ ^ reportada recientemente f
»rl979(n),ForerortaC
Distribuddn.— Colombia y Ecuador;
v“Ai-utfU€iai.2i7I6(COL,INB,K
1. 936 (PSO.QCA).
^hytologia 4M7. 1980. (Fi^ 5). Tao. COLOMBU. Ckoo, c.„
VHOLOTIPO: COL; isonpos: HUA, INB. MA. Mni
:HUA,1NB.MAJ
■ npucion.— Colombia y Ecuador en el nais pn 1
AUomarkgrafialaxifloraAH
“'•^'“"-CotatoyEcuado.enelpa.senOelOOvgoo
Por la forma de la corola, este taxa ^ IL r elevacion.
tienen corolas similares, pero A. laxiflora se ^ede semrar"^ tienen ambas espec
ispuestas y bracteas mas pequenas y escasas ^ orescencias con las flores laxame
127
Especimenes representativos exaininados. VaUe del <
abr 1993 (fl, fr), Devia et al. 3937 (COL, MO, TULV).
^**®n*ariigrafia plumeriiflora Woodson, Ann. Missouri Bot. Gard. 20:627. 933. (Fig. 6).
Tiro. COLOMBIA. BoyacA;
jsl.GH,K,MOlfotografla,
Distribucion. — S de Honduras a Costa Rica; en Colombia en elevaciones de 100-1200 m.
Allomarkgrafia plumeriiflora se puede distinguir por sus inflorescencias conspicuamente bracteadas, asi
como por el tamano de la corola. Se puede confundir con A. antioquiana, pero esta ultima especie tiene flores
mas grandes y de forma diferente.
Especimenes repre
MO,NY).Choc6-
staneda 2720 (COL). S
cur et al. 2799 (COL). Valle del Canca: Yatacue, Alto Anhicaya, v;
JAUM,MEXU,M0).
Mistrato, entre Puerto de Oro yjeguadas, 19 sep 1991 (fl), Betan-
iDagua, 17 jul 1984(0), Gentry 6- MonsoJve 48251 (COL,CUCV,
AGRADECIMIENTOS
Quiero agradecer a los siguientes herbarios por permitir el acceso y estudio de sus colecciones a traves de los
aflos: ARIZ, COAH, COL, CR, CUCV, F, G, GH, HUA, INB, JAUM, K, MA, MEDEL, MEXU, MO, NY, P, QCNE,
S. TULV, U, us. Asimismo, agradezco las facilidades logisticas brindadas por las siguientes personas, que fa-
cilitaron la estadia o el trabajo de campo en Colombia: Alvaro Idarraga (HUA), Dayron Cardenas (COAH), Fe-
Journal of the Botanical Research Institute ofTexas7(l)
Betancur (COL) y Ricardo Callejas (HUA). Mary Endress (Z) colaboro con la re-
Endress, M.E., S. Liede-Schumann & U. Meve. 2007. Advances in Apocynaceae: the enlightenment, an introduction. Ana
Missouri Bot. Gard. 94:259-267.
Morales, J.F. 1 997. A synopsis of the genus Allomarkgrafia (Apocynaceae). Brittonia 49:337-345.
Monies, J^F. 2006. Estudios en las Apocynaceae Neotropicales XXIII: una nueva especie de Mandevilla (Apocynoideae,
Mesechiteae) y nuevos reportes en las Apocynaceae (Apocynoideae, Rauvolfioideae) de Colombia. Anales Jard. Bot
Madrid 63:51-54.
Morales, J.F. 2009. Estudios en las Apocynaceae Neotropicales XXXIX: revision de las Apocynoideae y Rauvolfioideae de
Honduras. Anales Jard. Bot. Madrid 66:217-262.
SimOes, A.O., M.E. Endress, T. van der Niet, LS. Kinoshita & E. Conti. 2004. Tribal and intergeneric relationships of Mesechiteae
Apocynaceae): evidence from three noncoding plastid DNA regions and morphology. Amer. J.Bol
AULONEMIA DAVID-SMITHII AND A. RUBRALIGULATA
(POACEAE: BAMBUSOIDEAE: BAMBUSEAE: ARTHROSTYLIDIINAE):
TWO NEW SMALL-FLOWERED SPECIES FROM PERU
Emmet J. Judziewicz
Eric J. Geisthardt
Stevens Point, Wisconsin 54481, U.S.A
Christine M. Waas
Robert W. Freckmann Herbarium
Department of Biology and Museum of Natural History
University ofWisconsin-Stevens Point
Stevens Point, Wisconsin 54481, U.S.A.
ChristineMWaas@uwsp.edu
Stevens Point, Wisconsin 54481, U.SA.
Sol Sepsenwol
Stevens Point, Wisconsin 54481, U.S.A.
ABSTRACT
RESUMEN
INTRODUCTION
The woody bamboo genus Aulonemia Goudot (Poaceae;
its undescribed diversity throughout the Andes (Clark 1995; Judziewicz et al, 1999) in Venezuela Qudziewicz
etal. 1991; Judziewicz & Riina 2006), Colombia (Clark & Londono 1990; Clark et al. 1997, 2007; Judziewicz et
al- 2013), Peru Oudziewicz & TyrreU 2007) and Bolivia Oudziewicz & Clark 2011; Judziewicz et al. 2010,
2011).
Several Peruvian species are notable for their effuse panicles of small spikelets and presence of leaf sheath
niarginal hmbriae. Aulonemia parviflora 0- Presl) McClure (Huanuco, Cusco) has robust foliage leaf blades
(21-)35-52 cm long and (3-)3.7-5.8 cm wide that are strongly tesseUate abaxially, and extraordinarily long,
pale brownish inner ligules 40-80 mm long, and papery, confluent, non-sphtting hmbriae on its foliage leaf
sheath margins. On the other hand, A. humilUma (Pilg.) McClure (Loreto or San Martin) has much smaller
Journal of the Botanical Research Institute of Texas 7(1)
non-tessellate foliage leaf blades 8-17 cm long and 0.7-1 cm wide with sheath margins bearing regularly
spaced, discrete (non-confluent) marginal fimbriae, terminal sheath fimbriae 35-50 mm long, and inconspicu-
To this group of “small-spikeleted” taxa with marginal leaf sheath fimbriae may now be added two new
Peruvian species (Table 1). Aulonemia mbraligulata]udz. & Geisthardt (Amazonas, Cajamarca, and Junin) has
elongate, reddish inner ligules 20-45 mm long, foliage blades 15-25 cm long and 0.7-2.3 cm wide (that are not
at all tessellate), and pectinate-fimbriate sheath margins, while A. david-smithii Judz. & Waas (Pasco) has in-
conspicuous inner ligules, foliage leaf blades 25-38 cm long and 27-3.6 cm wide, sheath margins with densely
matted confluent fimbriae (these ultimately splitting) and terminal sheath fimbriae 10-15 mm long. Neither of
treatments of Peruvian grasses (Tovar 1993; Brako & Zarruchi 1994).
KEY TO THE ANDEAN SPECIES OF AULONEMIA
WITH MARGINAL FIMBRIAE AND EFFUSE PANICLES OF SMALL, SLENDER SPIKELETS
Plant parts were measured using a mm ruler, and a portion of the abaxial leaf blade epidermis (from the central,
non-marginal part of a well-developed blade) from one sample of both species was examined using air-dried,
uncoated, untreated material in a Hitachi S3400 scanning electron microscope; specimens were examined
from the following herbaria: F, K, MO, NY, P, US, UWSP, and WIS.
1; culms apparently less than 5 mm in diameter, glabrous. Midculm branching pattern and
culm leaves not seen. Foliage leaves glabrous and smooth; sheaths strongly keeled, the midnerve prominent
near the summit, the margins papery, chartaceous, and prominently cross-puckered on the lower leaves, on
the upper leaves the sheath margins breaking up into very dense, curling, orangish-brown, prominent fimbri-
ae 3-5 mm long, each fimbria attenuate into a mass of interwi^^^^ 3heath
summi ac mg auricles, but with erect, brownish, hispidulous fimbriae 10-15 mm long; outer ligule varying
from an indumte nm ca. 1 mm long to (one one side) an ovate, stramineous, shiny flange 2-4 mm long; inner
igule not evident, not large if present; pseudopetioles 2-4 mm long; blades 25-38 cm long, 2.7-3.6 cm wide,
TT ^°"g-»cuminate, the base obtuse and symmetrical, moderately strongly tes-
Leaf ■ I acute, paleas 1.8-3 mm long; flowers and fruits not seen.
Judziewiaetalj
Popinoe.— Abundant; globose to slightly compound; 4-6 pm in diameter.
Stomata.-Common; stomata alternating; stomatal rows 4-5, adjacent to costal
branched papillae present at each of the comers, giving
Interstomatal cells.—ca. 20-30 pm long, outline indeter
,htly compound, uniform in size.
Long Cells.— Dimensions difficult t
jparatus a plus sign (+) shape,
papillae present, abundant, globose U
133
Pricfeles.— Abundant, ca. 4/10,000 mostly in intercostal zones, but present in costal zones; 30-50 pi
long, 30-50 pm wide, tapered to a sharp tip, like a candy kiss.
Short Cells.— Difficult to discern due to dense papillae.
Microhairs— 2-celled, 55-60 pm long; basal cell 28-33 pm long, 7-10 pm wide; apical cell deflated, c
30 pm long.
Aulonetnia david-smUhii is distinctive in its densely matted orangish marginal fimbriae (densest in the
genus), as well as the Aulonemia queko Goudot -like flange at the summit of the leaf sheath, effuse panicle with
small spikelets, and somewhat abaxially tessellate leaf blades. The species is known from just a single collec-
tion made on the ridgetop cloud forest summit of the north-south trending Cordillera de San Matias in Oxa-
pampa, Pasco, Peru,at an elevationof910m.ltis named after DavidN.Smith(1945-1991),assiduous collector
of the Peruvian and Bolivian flora for the Missouri Botanical Garden. Smith gave the longitude as 75n2’W, but
from his field notes and label description, the collection was probably made at the summit of the cordillera at
75°06’W (according to Google Earth); 75°12’ would place the collection in lowland forest at 325 m elevation.
From Tropicos (accessed 24 April 2012), Smith’s collections from that site included many ferns and epiphytic
shrubs, withno vegetation greater than6mtall.Brako&Zarucchi(1994)citedSmith2045a5A.humtIlima, but
that species is known only from the type and lacks confluent marginal fimbriae, has shorter sheath summit
fimbriae, and has shorter, narrower leaf blades.
AdoMMnia rubraUgulat, Judz. & GeislharA. sp. nov. (Figs. 4-5). 7^^ PERU. 5.„jo«d.
Scrambling or climbing woody bamboo; cute ca.3nunmdiameter, reportedly up toSmlong, smooth. Culm
leaves appLntly not Lferenriated from foUage leaves. Midculm nodes bearing single^divergent branches
Foliage leaves wUh sheaths glabrous, strammeous to greenish, striate, the margins with abundant recurved
stramineous pectinate fimbriae 2-4 mm long spaced about 7-10 per cm, or when young confluent tnto a
chartaceous membrane; sheath summit lacking auricles and hmbriae; outer ligule ca. 1 mm lot^, tndurate
stramineous, prolongedon one side intoanovate stramineous, shiny flap 1.5-3.5t^ long; ,nnerhgule20^5
mm long, membranous, linear-lanceolate, acute, glabrous, reddish when young fadmg to «eous w^
a^;pseudopetioles2-3mmlong;blades.5-23cmlong0«^^^
rg\r:s:bt:s;s:c—
PedLle up toScm long mflorescence terminal, 8-20 cm tall, an open, ovotdpamcle; branches^
smooth,themainonesascendingata45°angle.Spikelets(12-)17-25mmlong,ca. mmwi e,
apex; sumens3,theimmatureanthersca.lmm long; pistil with2.mmaturest«mas,rru.t not seen,
doM for.,,. 14 J„„ 1978, in dammed Coalk. I>t9z&m.rdl» Te™ 7828 (MO. US, UWSP). Pm,.
'^teMl,P„ve9i,,t500-19<»m,3-4jmi929.dens.lo««,Kiiltp&S»*259®O<7.US).
Gentry et al. 22898 (MO) was also examined and exhibited a sim
Judziewicz et al.. Two new species of Aulonemia from Peru
Interstomatal cells.— Ca. 40-50 iim long, outline indeterminate; papillae present, abundant, globose to
slightly compound, uniform in size.
Long Cells.— Dimensions difficult to discern due to dense papillae.
Prickles.— Common, ca. 3-4/10,000 pm^; 35-45 pm long, 40-50 pm wide, tapered to a sharp tip, like a
Short Cells.— Difficult to discern due to dense papillae.
Microhairs.— 2-celled, 75-80 pm long; basal ceU 45-50 pm long, 7-l(
30 pm long.
Macrohairs. — None seen. i • r
Aulonemia rubraligulata is endemic to cloud forests in northern and centra Peru at e
1500-2000 m; it may eventually be found in neighboring Ecaudor, since one collection is only 8 km from t
border of that country. The prominent reddishfwhen fresh) inner ligule20-45mmlongis unique in the gen
and prompted the specific epithet. Tovar (1993) cited Killip & Smith 25959 as A. humilhma, but that species
known only from the type, is smaller of stature, has inconspicuous ligules, elongate leal sheath summit hi
briae and smaller spikelets with fewer florets.
ACKNOWLEDGMENTS
WethankUWSPsmdentEvaC.HathawayforthelinedrawingsandGerritDavidseCMCTfortheloanotspeci-
mens, assistance withobtaininginformation on David SmUh,andhU helpful rmew.Gem.Dav.dse (MO) and
Lynn Clark (ISC) provided valuable reviews.
references
e flowering plants and gymnosperr
N York Botanical Garden, Bronx. Pp.
Brako, L. and J.L. Zarucchi. 1994. Catalogue of tl
Monogr. Syst. Bot. 45:1 -1 328.
CuRK, LG. 1995. Diversity and distribution of tf
et al., eds.. Biodiversity and Conservation of Neotropical I
Cet” LG, JuDzinncr, .no CD. Tvmeil. 2007. Aulonemia xinmfiaetPox^^-. Bambusoldeae: Oambuseae), a new spe-
Clark, LG., X. LoNDofJo, and M. KoeAYASHi. 1 997. Aulonemia bogotensis (Poaceae: Bambusoideae), a new species from the
Cordillera Oriental of Colombia. Brittonia 49:503-507.
surface view. Bothalia 12:641-671.
JuoaEWKz, EJ. AND LG. Clark. 201 1 . Aulonemia cochabambensis (Poaceae: Bambusoideae: Bambuseae: Arthrostylidiinae),
an anomalous new species from Bolivia. Brittonia 63:375-378.
JuoziEwcz, EJ., LG., Clark, X. LoNoofto, and MJ. Stern. 1999. American Bamboos. Smithsonian Institution Press, Washing-
JuDZiEwo, EJ., G. Davidse, and LG. Clark. 1 991 . Six new bamboos from the Venezuelan Guayana. Novon 1 :76-87.
JuDziEwrcz, EJ., EJ. Geisthardt, LD. Gibbons, D.C. Ziegler, M J. Zueger, and S. Sepsenwol. 201 1 . Three new Andean species of
Aulonemia (Poaceae: Bambusoideae: Bambuseae) with sheath auricles. J. Bot. Res. Inst. Texas 5:485^98.
JUDZIEWICZ, EJ., X. LoNDOfio, AND LG. CuRK. 2013. Two new northern Andean species of Aulonemia (Poaceae: Bambusoi-
JuoziEWKZ, EJ. and R. Riina. 2006. Aulonemia dinirensis (Poaceae: Bambusoideae: Bambuseae), a new dwarf Venezuelan
species from the easternmost Andean paramos. Bamboo Sci. Cult. 19:11-15.
JuoziEwicz, EJ., E.L Shea, anoT.M. Wayda. 2010. Two new Bolivian species of Aulonemia (Poaceae: Bambusoideae: Bambu-
seae). J. Bot. Res. Inst. Texas 4:569-579.
D C.D. T
L 2007. y
Tovar, 0.1 993. 1
TWO NEW NORTHERN ANDEAN SPECIES OF AULONEMIA
(POACEAE: BAMBUSOIDEAE: BAMBUSEAE: ARTHROSTYLIDIINAE)
WITH VERRUCOSE CULMS
Emmet J.Judziewicz
Ximena Londono
Robert W. Freckmann Herbarium
Department of Biology ar)d Museum of Natural History
University of Wisconsin-Stevens Point
Stevens Point, Wisconsin 54481, USA.
Sociedad Colombiana del Bambu
Apartado Aereo 1 1574
Cali, COLOMBIA
ximelondo@gmail.com
Lynn G. Clark
Department of Ecology, Evolution and Organismal Biology
251 BesseyHall
Iowa State University
Ames, Iowa 5001 1,U.S.A.
Igclark@iastate.edu
ABSTRACT
INTRODUCTION
Exploration of the northern Andes continues to reveal new species in the woody hamhoo genus Aulo,
Goudot (Poaceae: Bambusoideae; Bambuseae: Aithrostylidiitiae). In the last two decades novelties havt
described from Venezuela Ondziewicz et al. 1991; Judziewiez 2004; Judziewicz & Riina 2005), and Colombia
(Clark & Londofto 1990; Clark et al. 1997, 2007). This study is based upon examination of material from these
herbaria; CAS, COL, F, K, MO. ISC, NY, P, UC, US, TULV, UWSP and WIS. Two distinctive new species may
BOW be added to the genus. Both are delicate, viniugbamboos with raspy culms that have the appearance and
■exture as if gritty bits of purple sand have been embeddedinthesurface (of evident utility when climbing) of
the culms, Verrucose, “raspy" culms ate otherwise knowuinAulunemiaonly in several southern Biaziltanand
ArgeutinUn species formerly ptaced in the segregate genus Cotathelia McClure Sr E.W. Smith Ondziewicz -
al. 1999; Tyrrell et al. 2012; Santos-Goncalves et al.. In prep ). The species of Auta™ -
*nay be distinguished by the following key:
h verrucose culms
2. Fertilp finrotc -yi i\ a ^ ®
2. Fertile florets 3-16, 6-103
3. Fertile florets 9-103 mm lon<
3.Fertileflorets7-9mml
4. Foliage leaf sheaths i
long, 0.3-1.
A. lanciflora McClure & LB. Smith
LB. Smith) McClure]
inibriae and sometimes also inconspicuous transferred into AuZemia]
-A. eingulata McClure & LB Smith rr ^ fertile florets 8-1 1 mm
•anyoftheBraziliancongenersofAu/onr>m- ' McClure & LB. Smith, McClur^^
«a (Hitchc.) McClure a much mnr. k . ^ only in th. r- Purple-spotted
’ species with ^ endemic A. vis-
P ^®™smooth,viscidculms.
Ian boundary. 700n_7snn r. tetania [7»27'N, 72'-26-W], along Rfo T;
....apaez (s,c. tor Villa Pdez. 7”29n, 72°27-W1 a.
a, bordering Colt
‘^ryPES; 1-3. US-2). (lowers purple; NV: pajiUa or corta caT
Vining,climbingbamboowithslenderculmsca 7r« . corta ca sOn. Steyermarfc
apically curling, easily rubbed off i ^1*”“’ long delicate "’^^S^ns smooth; sheath
mm l<mg, glabrous, smamiur ous bW^ H t '™- cUiaSmt a
asymmetrical at the base, long-acuminate T "" '”*• •anceoUteT''^’ ^"3
along one matgin beneath. Synflotescence usuab “"'“•omus or with °"f I'*""'"''
6-12 cm tall, 4-S cm wide. oLd det^f tb b T I- f ""P'
Flowers with lodicules not evident in spikelets ell a ^«iolate on Aete ^ nar-
yellow-brown; gynoecium not seen Fruit not se apparently 2, the anth ^ t bifid.
Known from one collection in Venezuela mTde alor, rh o
iamype made bythe lateZmll R ^ S ™ “'^^er spikelets 40-
sp«ipasnndescribedandsugges.edrXr-,I!IIn"d™We'^'“’',f®"-^^^^^^
andletdev^^^^arbe’::^^^^^^^^
‘^’^0-2.Uwsp).
Journal of the Botanical Research Institute of Texas 7(1)
JudziewiaetaLJ
Vining, scandent, delicate, intricately-branched bamboo. Culms 1.5^ mm in diameter, hollow with a lumen
ca. vs diameter of the culm, glabroua. simmineous, Ae larger, older culms det^ly verruco^ with ramed
Purple-bUck siUceous papillae about 130-180 pm in diameter and 4W00 pm tJl, the more sUnder, uppe
pontons of the culms merely purple-macuUte with papillae atenl. shght^ "h alll nten.^ ~
ginUe l-2mm wide producing (onlo«er,largea.lms)lsnonglyascendmgbranchwuhadtsultntemodeca
Icml h fi odu imordia that may develop into adventitious roots, the upper nodes pro
■fcicing™p,oLut^ualblcte"cCleaves (seen only in DepcMastklenacolto^^^
6 H jsuuequdiuid , orangish-browTi, the margins finely ciholate
atjunctureofsheath and girdle, the sheaths 3-5 cm long, stnateorang . f /mm
fcapexacme, truncate, Lughdyconcave,glabrous or (if concave) fnngedwtthafe«delicaie2-3mm gold
142
™ ^2 „.„, l„ng, 1.5-3 „.„ wi*
plish, adaxially pubescent; blades 9-17 cnflong 1 7-2 8 cm wid^’ T P^eudopetioles 2-5 mm long, pur-
somewha. asymmetrical to very slightly” ^rdlu auhTh “
glabrous to sparsely puberulem beneath, the margins cartitonou"* T”““' ” *'*'““*
With a slightly darker stripe along one margin beneath ^ i t ^^^rorsely scabrous, concolorous or
ovoid panicle 10-22 cm tall, 8-15 cm wide the few b i, ^ 5 cm long, glabrous. Synflorescence an
and smooth. Spikelets 40-70(-80) mm long slender^tram^ spikelei pedicels all glabrous
abundant purple spots, finely and densely appressed b strongly suffused with purple due to the
Plng such that the summit ofone note, extends abouZray“upTtaXf?H°“*^^^^
bract 0.3-1 mm long often present 0.5-1 mm below spikeler iL i ^ ^ ^ succeeding floret; scalelike
acute to apiculate, 5-7<-9)-nerved; upper glume 9-13 mmta r
late 7-9.„erved; fertile floreB 4-6. the lowermost floretsometo^rirf”'”
lie; fertile florets with lemmas 13-16 mm long, lanceolate fi uppermost 1 or 2 floret(s) ster-
trorsely scabrous awn 3-6 mm long; paleas 8-13 mm ^ ^~7(-9)-nerved, abruptly tipped with an an-
and concealed by the lemmas, bicarinate, clliolate on the ktlTm *' T*"’
1.2 mm long, rhombic, acute, transparent, obscurely 2-3-uerved tZ d Z' ° ®'
s.amens3,theamhers4mml„ng,h„ear;pishUwUh2hispZs,;gZ fZ
O.I»t,p..,meu.e»m,„rf:COlOM.U.CES«USl«;DaLIN4:Mp^ . f™>'notSeen.
Auhnemia verrucosa is a viuing species that has culms with distin f ' ’
etonga,e,slender,aw„ed, purple-maculate spilrelem.AColombiZXr'"""^^^^^
of An toquta and m the Serranla de Perija in Cesar/Magdalena about Z CoriUlcra Central
from 1700-2230 m (and perhaps up to 2620 m). I, is dZiy related to Tv”' « elevations
latter s^ciestesmallet.awnkss spilrelets 22-35 mm long With
e ferttle type specimen from the Depanment of AnUoquIa has eli^ b """
W2 cm long and 1-7-2.2 cm wide while the sterile Department MaZf with blades
a separate taxon, but we await fertile material before describing it as such. represent
ACKNOWLEDGMENTS
We thank UWSP student Eva C. Hathaway for the line drawings x
holotypeof24«fo„emmnotat«,GerritDavidse(MO)forthefoanof;pecimer^°^^^
and an anonymous reviewer for helpful comments. "" assistance, and Pedro Viana
REFERENi
CuvRK. LG., JuDaewiC4 EJ. and CD. Tyrrell 2
from Colombia and Venezuela. Bamboo Sci. Cult 20:1-6.
Judziewiaetal.,
143
Clark, LG., X. LoNooii)o, and M. Kobayashi. 1997. Aulonemia bogotensis (Poaceae: Bambusoideae), a new species from the
Cordillera Oriental of Colombia. Brittonia 49:503-507.
JuoziEWicz, EJ. 2004. Aulonemia. In: Steyermark, J.A., P.E. Berry, K. Yatskievych, and B.K. Holst, eds. Flora of the Venezuelan
Guayana, Vol. 8: Poaceae-Rubiaceae. Missouri Botanical Garden Press, St. Louis. Pp. 40-45.
JuDZiEwicz, EJ., LG. Clark, X. Londono, and M J. Stern. 1 999. American Bamboos. Smithsonian Institution Press, Washington,
DC.
species from the easternmost Andean paramos. Bamboo Sci. Cult. 19:1 1-15.
TvRREa, C.D., A.P. Santos-Goncalves, X. LondoNo, and L.G. Clark. 2012. Molecular phytogeny of the arthrostylidioid bamboos
(Poaceae: Bambusoideae: Bambuseae: Arthrostylidiinae) and new genus Didymogonyx. Molec. Phylogen. Evol.
144
BOOK REVIEW
Lytton John Musselman and Harouj J. Wiggins. 2013. The Quick Guide to Wild Edible Plants. (ISBN: 978-1-
4214-0871-2, cloth). The Johns Hopkins University Press, 2715 North Charles Street, Baltimore,
Maryland 21218-4363, U.S.A. (Orders: www.press.jhu.edu, 1-800-537-5487). $24.95, 144 pp., 116 color
photos, 5 Vi" X 8 Vi".
Growing up in New England and spending much time in the Mid-Atlantic States, I was interested to get my
hands on this little book to see what I knew about wild edible plants in my native habitat, and I was pleasantly
surprised. Musselman and Wiggins offer us an easy-to-use guide to common edible plants of the Eastern Sea-
board, most of which I am familiar with, some of which I have tried, and a number that sound, well . . . interest-
ing and worth an experimental taste test.
It is a self-described “quirky” guide which offers recipes for 31 wild edible plants, everything from
aperitifs to sweets. In fact, the guide is developed almost like a cookbook, starting with condiments, rang-
ing through greens and starches, and ending with cordials. The authors have a great sense of humor as they
describe their many plant collecting forays “crossing icy streams in winter and tenacious miasmic mud in
scorching summer heat . . . thorns and spines . . . clouds of insects . . . extracting plants from muck that seemed
to extend to Middle Earth . . . boiled, parched, ground, baked, and dehydrated only to find that what remained
was a taste that would gag a maggot.”
The authors don’t attempt to cover every possible edible plant out there, but they have picked some great
ones. It is a fast and easy read. Their recipes use few ingredients so as not to distract from the true flavor of the
plant and were tested over many years on their sporting families and students. The authors provide botanical
information on each of the plants and a cautionary overview of some of the key ones to avoid such as Poison Ivy
and Poison Oak.
Reading this guide, I was initially puzzled over who, exactly, was the target audience. It is not meant as
a survival guide for people dependent on foraging for existence as many of the recipes require patience on the
part of the gatherer for the precise moment when the plant is in bloom or ready to harvest. For example, in de-
scribing the recipe for Cane Crispies, the authors admit to finding fruiting cane only 3 times in their 75 years of
combined field work— though they claim it was worth the wait andsearch. It is hard to imagine someone lost in
the Appalachian Mountains for that long. Then it began to dawn on me, the book was intended for those with
enough background on plants to not make grave errors like the demise of Socrates, drinking Poison Hemlock.
Itistargetedfor those of us who aren’tdependent on foragingfor survival, those withaloveof the out-of-doors,
with a wilhngne^ to get a little dirty and to try new things. The book is intended for someone like ... me!
From Cattail Com Dogs to Pickled Orange Day Uly Buds (or Ditch Lilies as we used to call them) there
are electable treats to be had for the patient forager. How about some Black Locust Flower Fritters or Redbud
Floweis on one s salad or ice cream? Yum! I’ve tried Nettle Omelets, Field Garhc, and Sassafras Tea. The one
rnackfl^
IreXCa^tral^r
snenmaXsuX ^ussleman and Wiggins Wild Blueberry Cordial. Having
Musselman and Wiggins have spent years researching edible plants and refining their recines Their
expenence and knowledge extends way beyond this fun, little guide book. 1 appreciate their wit and humor
and their common-sense selections for easy-to-find Wild pHihUrrlc t d u ‘neir wit and humo
Edible Plants of Texas next?t_rw^iu “ they would like to take on Wild
DOS NUEVAS ESPECIES DE GUADUA PARA EL PERU
(POACEAE: BAMBUSOIDEAE: BAMBUSEAE: GUADUINAE)
Ximena Londono
Sociedad Colombiana del Bambu
A.A. 1 1574 Cali, COLOMBIA
Guadua Kunth (Poaceae: Bambusoideac. — - i n ni
genero tipo de la subtribu Guaduinae, la cual incluye ademas los generos. Apodad^ Eremocauon, tateay
meca (Soderstrom & Londono 1987; Judziewicz et al. 1999; Londono & Clark 2002; Ruiz-Sanchez et al 2008).
Estudios recientes basados en secuencias de ADN y en caracteres morfologicos (Ke chner
Phylogeny Group 2012) ratifican que la subtribu Guaduinae pertenece al dado e os am ties neotropica
lenosos y que conforma un linaje monofiletico en el arbol evolutivo de los bambiies. Esta subtnbu se caracten-
za por tener una micro morfologia de la lamina foliar inusual, en donde la lamina posee estomas en ambas su-
perbcies (amfiestomatica) rodeados por papilas, caracter que se observa prmcipa mente en a ^
adaxial(Soderstrom&Ellisl987;Ruiz-Sanchezetal. 2008); adem.s, la lamina
tipica nervadura central compleja de la mayona de los
■■ 1987).
29 especies de las cuales i/ taxas ban sido de^ritas en los ultimos 20 anos. La region
an«z<h.icade Brasil, Peru. Mivu, Ecuador, ColombiayV=nen.elasecon«Wcen«^^^^
E de esclerenquima intercostal ]
mil
bambiies realizado por Londono (2002) para America Latina y en particular para el Peru, se reportaron 37 es-
pecies y 8 generos. Los departamentos de Pasco y del Cuzco son los que albergan la mayor diversidad, mientras
que los departamentos de Madre de Dios y del Amazonas son los que presentan la mayor area cubierta por
bambiies (Takahashi & Ascencios 2004). Ancestralmente el bambu ha sido utilizado por comunidades nativas
y rurales del Peru, desde las tierras alias de los Andes hasta las tierras bajas de la region amazonica (Tovar
1993) y se utiliza en una cantidad proporcionalmente menor que olros materiales que usan para la construc-
ci6n, siendo los usos mas frecuentes techos, cercos, paredes, y como elemento decorativo de viviendas y locales
comerciales, sin cumplir realmente un papel estructural (Takahashi & Ascencios 2004).
Un viaje de coleccion por los departamentos de San Martin y Amazonas, en el noroccidente del Peru, ha
revelado dos nuevas especies de Guadua relacionadas con Guadua angustifolia Kunth, las cuales reiinen una
serie de caracteristicas como son el gran tamano, rectitud del culmo y espesor de pared mayor a 1.5 cm, lo que
las convierte en especies promisorias para el desarrollo local y regional en sectores como la construccion e
Se examinaron especimenes en herbarios nacionales e intemacionales (K, ISC, MOL, MO, USM, US) pero
bambiies tropicales varia en cada especie, con un rango de fluctuacion entre 3-60 anos. Ante la evidencia de
caracteres morfologicos vegetativos linicos en estas nuevas especies se toma la decision de describirlas sin flor.
La importancia economica de las especies afines a Guadua angustifolia Kunth obliga a utilizar caracteres
macromorfologicos, CSciles de visualizar por parte de los interesados con el fin de que se logre una pronta iden-
tificacibn, por eso se hace enfasis en estructuras morfologicas como rizoma, culmo, hoja caulinar, yema de la
region nodal y ramificacion. La estructura morfologica denominada hoja caulinar es una importante fuente de
caracteres para diferenciar las especies del genero Guadua y siempre debe ser recolectada; el tamano e indu-
mento de la hoja caulinar, la proporcion en tamano entre la vaina y la lamina caulinar, la forma y tamano de la
ligula, son de relevancia para la identificacion de las especies de este genero.
La siguiente clave diferencia las dos nuevas especies de Guadua de la especie tipo Guadua angustifolia
Kunth, que ocurren en el noroccidente del Peru.
uerpo del rizoma 20-40 cm longit
forma de tor tuga, L„
laao aei nzoma, con diametros 8-10 cm por raiz de sonorte- rafrc*; n . ^
diametro. Culmo (18-)20-27 m de altura, (6-)9-17 cm diametro erecto en la bai^^T
color verde con rayas verdes mas oscnra. .,.Mo ioven ^ ap^
joven, pubescente en el i/S sunerior h„Pros,
(15-)30-49 cm 1
n; nudo solitario, linea nodal horizontal, proi
Journal of the Botanical Research Institute of Texas 7(1)
ligeramente inclinada por debajo de la yema, con banda de pelos blancos, adpresos, arriba y abajo de la linea
nodal: banda superior 0.4-0.6 cm, banda inferior 1.2-1.5 cm; yema solitaria, ampliamente triangular, profilo
pubescente, obtuso hacia el apice, cubierto por pelos adpresos de color cafe, margenes ciliados. Hoja caulinar
triangular, 68-91 cm de longitud por 68-72 cm de ancho, coriacea, de color cafe, decidua, lamina 5 a 6 veces
1 que la vaina; vaina 57-77 cm de longitud por 68-72 cm de ancho, abaxialmente hispida, cubierta
pelos ondulados, aproximadamente 1 mm de longitud, hialinos, retrorsos,
adpresos, y b) pelos hispidos, rigidos, 2-2.5 mm, cafes, removibles, margenes conspicuamente ciliadas con
raya marginal oscura, cilios cafe pardo, aproximadamente 5 mm longitud, superficie adaxial glabra, brillante;
ligula interna 2-3 mm de longitud, truncada, arqueada en el centro y prolongandose ligeramente por un lado,
termina antes de la margen en ambos extremos, adaxialmente glabra, abaxialmente pubescente, margen den-
samente ciliolado, cilios hialinos, cortos, aproximadamente 0.2-0.3 mm de longitud, con o sin desarrollo de
setas orales en la parte media de la ligula en mimero de (4-)10-42; setas orales 1.5-2.7 cm longitud, basalmen-
te redondeadas, escabridas, oscuras, y rectas, luego ligeramente onduladas y pajizas; lamina 1 1-14 cm de lon-
gitud, 13-15 cm de ancho, triangular, erecta, persistente, abombada, apice mucronado, mucron fuerte, aproxi-
madamente 1 mm de longitud, cuando se rompe se divide en 2 o 3 partes, si son 3 una central mas ancha y dos
laterales mas delgadas, superficie abaxial glabrescente, superficie ada:
pelos cafe claro entre las nervaduras, margenes ciliados, cilios aproxir
cion intravaginal; rama primaria solitaria y con espinas, con 1-
sin desarrollo de complementos foliares en el extremo; sin desarrollo de ramificaciones en el VS medio del cul-
mo. Hojas del FoUaje 10-12 por complemento, dimorficas, las del % basal del culmo con laminas foliares an-
chas (hasta 4 cm) y con presencia de auriculas fimbriadas en la parte superior de la vaina, los del VS apical con
laminas foliares menos anchas (hasta 2.5 cm), con o sin presencia de auriculas fimbriadas sobre la vaina; vaina
adaxialmente glabrescente, con auriculas falcadas-lanceoladas y/o fimbrias en la parte superior, margenes ci-
hados, evidentemente mas oscuros, cilios 0.5-1 mm de longitud, usualmente hialinos; auriculas falcadas-lan-
ceoladas, 5-8 mm de longitud por 0.5 mm de ancho, verde o purpura, pubescentes, con las margenes fimbria-
color blanco-marfil, ligeramente onduladas, basalmente escabridas, 6-12 mm de longitud; h-
uncada, densamente pubescente, blancuzca, margen conspicua-
1 externa aproximadamente 0.5 mm de longitud, glabra,
brillante, hnamente ciliolada a lisa; pseudopeciolo 5-8 mm de longitud, adaxialmente glabro, abaxialmente
glabrescente, pulvinado, pulvino piloso, de color pajizo a cafe; lamina (15-)18-23 cm por (1.6-)2-4.2 cm de
ancho, L;W = 5-8:1, 17-20 nervaduras, lanceolada a linear-lanceolada, adaxialmente glabra, raramente con
tricomas estrigosos esparcidos, transparentes, tricomas cafe-rojizo en la base, con 3-4 nervios submarginales
escabndos, abaxialmente glabra, papilosa, teselada. nervadura central y nervios primarios amarillos y promi-
nentes en toda la longitud de la lamina, un margen escabrido, el otro escabridulos, apice acuminado, con mu-
cron retorcido, 3-4 mm longitud. Inflorescencia no vista.
Etimologia.—Esta especie se nombra en honor de Lynn G. Clark (Iowa State University) lid^
mahzanlt ^ Muabks " recolectarlos e
I^tribucmny habitat.-Se distribuye en el noroccidente del Peru, en el Departamento de San Martin,
enne »uu y yuo m a onlla de rios y quebradas. No tolera areas inundables. Quedan muy pocas poblaciones
debido a la severa destruccion de la selva con fines agricolas. Crece asociada en el sotobosque con heliconias,
piperaceas, arace^, convolvulaceas, Costus sp., ciperaceas, melastomataceas, ciclantaceas, helechos, iracasy
grammeas, y en el estrato superior con arboles del genero Ficus sp. (higueron u oiel
Nombre comiin.— “Marona,” y al rodal “Maronal.” ^
Usos.-Seobsej^pocousodees.ebambaendareado„dec,ecequWsdebidoalaatodanciadema<k-
r” mnstracdon de los techos con zinc, y para extracr
estenlla o guadua picada para paredes y corrales. Su gran tamano sumado al esnesor de su naJd O a 5 cm) y
asuspropiedadesfisico-mecanicashacenqueesta especie tengaungranpotencialparakm^^^^
a aproximadamente 0
a densidad de culmos por hectarea (7000 culmos/ha) y una aka lasa de creci-
a de 15 cm de diametro y 25 culmos por sitio en una
0 (5) anos de cultivo. Tuesta Vasquez (com. pers.), morador de Atumplaya, asegura que Guadua
lynnclarkiae florece y no se muere.
Afinidades. Guadua lynnclarkiae es similar a Guadua angustifolia Kunth. Esta dos especies comparten
los siguientes caracteres: a) culmos de gran porte, con alturas hasta de 27 metros y diametros superiores a 15
cm; b) region nodal con la banda inferior ligeramente mas ancha que la superior; c) Hoja caulinar coriacea, de
color cafe; d) ligula interna de la vaina foliar arqueada en el centro y Ugeramente prolongada por un lado, fina-
lizando antes de las margenes en ambos extremos; e) yema de la region nodal cubierta por un profilo pubescen-
te de color cafe; y f) rama primaria solitaria, con 1- 5 espinas por nudo. Sin embargo, Guadua lynnclarkiae se
diferencia de G. angustifolia por tener (a) rizomas paquimorfos con raices de soporte gruesas, en numero de
3-4 a cada lado, que le proveen al rizoma una apariencia de tortuga vs. raices de soporte delgadas, en numero
de 4-8 a cada lado, que le proveen al rizoma una apariencia de lagarto; (b) entrenudos de los culmos ligeramen-
te mas largos, 15-49 cm vs. 14-34 cm; (c) culmos con espesor de la pared de 2-5 cm vs. 1-3 cm; (d) presencia
0 no de setas orales en la parte media de la ligula caulinar vs. no presenta setas orales; (e) lamina foliar de mayor
tamano, 15-23 cm de longitud por 1.6-4.2 cm de ancho vs. 6-20 cm de longitud por 0.5-2 cm de ancho; (f)
vaina foliar con auriculas falcadas-lanceoladas y/o fimbrias en la parte superior de la vaina vs. solo fimbnas; (g)
pseudopeciolo glabrescente, con pulvino piloso vs. pseudopeciolo glabro, con pulvino glabro (Tabla 1).
La presencia de setas orales en la ligula de la hoja caulinar es frecuente en la subtribu Guaduinae, obser-
vandose en generos como Eremocaulon, Olmeca y Otatea (Londono & Clark 2002; Ruiz-Sanchez et al. 2011;
Judziewicz et al. 1999) sin embargo no se habia reportado para el genero Guadua como lo mencionan Ruiz-San-
chezetal. (2008; 2011). Guadua lynndarfeiae puede desarrollar setas orales en la parte media de la ligula cauh-
nar, de 1.5 a 2.7 cm longitud, de color pajizo y conspicuamente escabridas basalmente. Estas setas orales no se
ajustan exactamente a la definicion dada por Clark & Cortes (2004) quienes ™
a de la ligula, <
, en la boca de la vaina de la hoja c
n la bast
a 7 cm de longitud por 0.2 a 1.2 mm de diametro, y
adherWaalasuperficleabaxialdebligulaintema. Lasse, ason.lesenGu<«k.al>n„**iae no
en aoten y Eremocauton (Ruiz-Sanchez e, al. 2010; Soderstrom & Londono 1987): son redondeadas y muy ,
eibrldas basalmente, con una apariencia mas defimbria que de seta pero por suubicaconenUbocade lava,
caulinar yen la mitadde la ligula intemasedescriben como setas orales. u ,
Las plantulas (chusquines) de Guadua lynnclarkiae desarrollan ademas de lammas foliares anchas, au
culas fimbriadas en la parte superior de la vaina foliar, esios dos caraceres strven para d««enaar en f«
viveroG.ly„„clarfciaedeG.ungus,,/oliuKun.b(lamlnasfoliarasmasangos,as,pres™^^^^^^
culas). Las plantas en estado adulto de G. lynnclarkiae no siempre conservan e carac er
briadas en la parte superior de la vaina foliar: es posible observar e
y sin auriculas fimbriadas.
„OU; An,mpl.ya, ,3 M„ 1999, C. Rrng,-
ba. Comunidad Atumplaya, 5”50'48.9"S; 77°14'52.2''W, 883 m, Feb 2001, N. Ocean s.n. (CU ,
/o-Rui; 01 y 02 (CUVC, MOL).
Guad t k h h- d' (Pig 2) TlPO- PERU Amazonas: ProvinciaBagua, Distrito Aramango.aprox. 1km via
.An, r^,, S; reWZS-W.W 6 M.y 2010, X. ri-*. J. ^ «. ri,,,,™ ,043 ,Horo™: MOU;
aismo complementc
s especies de Guadua en Peru
151
Journal of the Botanical Research Institute of Texas 7(1)
apice mucronado, mucron fragil, aproximadamente 1 mm de longitud, cuando se rompe se divide usualmente
en 2 partes, superficie abaxial glabrescente, superficie adaxial pubescente, nervada, densamente cubierta por
pelos cafe claro entre las nervaduras, margenes ciliados, cilios facilmente deciduos, aproximadamente 0.5-1
mm de longitud. Ramificacion intravaginal; rama primaria solitaria, sin espinas o con espinas inconspicuas
en mimero de 1-3 por nudo. Hojas del Follaje 6-10 por complemento, linear-lanceoladas; vaina abaxialmente
glabra, estriada, brillante, con fimbrias en la parte superior de la vaina, margenes finamente ciliolados, cilios
hialinos, 0.05-0.1 mm de longitud; fimbrias basalmente escabridas, pajizas, apicalmente onduladas, hialinas,
3-7 (-14) mm de longitud; Ugula interna 0.2-0.3 mm, truncada, glabra, margen finamente ciliolado, cilios
hialinos; ligula externa 0.1-0.2 mm de longitud, glabra, brillante, finamente ciliolada; pseudopeciolo (2-)3-5
mm de longitud, adaxialmente puberuloso, abaxialmente glabro, pulvinado, pulvino glabrescente, brillante,
pajizo; lamina (8-)14-16{-18) cm de longitud por (0.8-)l-1.5 cm de ancho, L:W = 7-19:1, 9-13 nervaduras,
linear-lanceolada, adaxialmente con pelos estrigosos, esparcidos, hialinos, antrorsos, hasta 1mm de longitud,
con 3-4 nervios submarginales escabridos, abaxialmente con pelos estrigosos esparcidos, hialinos, antrorsos,
hasta 1 mm de longitud, densamente papilosa, teselada, nervadura central prominente, amarilla, un margen
escabrido, otro finamente aserrado, apice acuminado, termina en mucron recto, 1.5-3 mm de longitud. Inflo-
Etimologta. — Esta especie hace honor a Josefina Takahashi (Asociacion Peruana del Bambu), lider en el
conocimiento, promocion y desarrollo del bambii en el Peru.
Distribucidn y habitat. — Ocurre en el noroccidente del Peru, en el Departamento de Amazonas, entre 400
y 600 msnm, donde se observa a orilla de rios y quebradas o en zona de laderas.
Nombre comiin. — Guayaquil.
Usos.—Se utiliza en la construccion, para la elaboracion de entrepisos, paredes de bahareque, techos, vi-
raestructura agropecuaria como cercos, gallineros, y corrales para animales
Afinidades. — Guadua takahashiae comparte c
lua angustifolia y Guadua lynndarkiae la hoja cauli-
ton la Iiguia cauunar nnaiizando antes de las margenes, la yema de la region nodal
protegida por un profilo corto y el desarrollo de una rama por nudo. Guadua takahashiae es mas cercana a G.
ongusti/olm, con quien comparte la longitud de los entrenudos (10-34 cm), el espesor de la pared del culmo
(1.5-2.5 cm), la forma triangular y el tamano del profilo, la forma linear-lanceolada de la lamina foliar, la pre-
^ncia de tncoinas estrigosos, transparentes, antrorsos, esparcidos por ambas superficies de la lamina foliar y
la presenca de fimbnas en la parte superior de la vaina foliar. Guadua takahshiae difiere de G. angustijolia por
tener^(l) escasa presencia de espinas sobre el culmo y sobre el follaje vs. abundante presencia; (2) espinas
cmndo presemes en nnmero de 1-3 por nudo vs. 1-3 por nudo; (3) prolib de la yema nodal glabrescenre, pig-
— de r^o vs. pubescenre, de color cafe; (4) hoja caulirrar de color catt rosado vs. cL; (4) superficie
^al rle U hoja cauirnar con baja concentraciOn de pelos hispidos, largos, de color cafe-hlalino en U regidu
dorsal y basal vs. alra concentradrin de pelos de color cafe; (5) ligula interna de la hoja caulinar horizontal vs
arqueada en el centre (Tabla 1). ^
A pesardequeG.tafeahashiae comparte el color cafe rosadn dp la h^or-oaxt- ^ y .
volven.es.abrescen.es,LcX:r:r"r;r^^^^^
^sobmelcuIrnohacerrqueGrrrrdrrrrrrrfiuhasHarseaunaespecieLairariL^^^^^^
te, con un gran potencial para la industria de la construccion. ^ sosteniblente
>1042 (MOL, COL, CUVe, ISC).
>.5»24'56.3"S;78°27'16,5"V
153
AGRADECIMIENTOS
Esta investigacion se realize gracias al proyecto “Promocion de la Rehabilitacion, Manejo y Uso Sostenible de
los Bosques Tropicales de Bambu en la Region Noroccidental del Peru” PD428/06 Rev.2(F) financiado por la
Organizacion Internacional de Maderas Tropicales — ITTO. Especiales agradecimientos a Josefina Takabasbi,
coordinadora del proyecto, a John Leigbt quien ba apoyado siempre las investigaciones sobre el bambu en Peru,
a Hitler Panduro, German Guerrero y Carlos Godenzi por la colaboracion prestada, a las personas de las dife-
rentes comunidades visitadas en San Martin y Amazonas, a Mercedes Flores de la Universidad Agraria La
Molina por colaborar con el secado de los espedmenes de berbario y a Myriam Monsalve por las ilustraciones.
Agradezco a Emmet Judziewicz y al revisor anonimo por sus acertados comentarios y sugerencias.
REFERENCIAS
Bamboo Phylogeny Group. 201 2. An updated tribal and subtribal classification of the bamboos (Poaceae: Bambusoideae).
Bamboo Sci. Cult. 24:1 -10.
Clark, LG. and G. CortEs. 2004. A new species of Otatea from Chiapas, Mexico. Bamboo Sci. Cult. 1 8:1 -6.
Judziewicz, E., L.G. Clark, X. Londono, and M J. Stern. 1 999. American Bamboos. Smithsonian Insti
Kelchner, S.A., LG. Clark, and Bamboo Phylogeny Group. :
(Poaceae, Bambusoideae) based on five plastid nr
LondoNo, X. 2002. Evaluation of bamboo resources in U
Bamboo for sl
Bamboo Workshop. VSP & INBAR.
LondoNo, X. AND LG. CuRK. 2002. A rev
nae). Syst. Bot. 27:703-721 .
Londono, X. and P. Peterson. 1 992. Guadua chacoensis (Poe
dvon 2:41-47.
osA, AND M.T. Mejia-Saules. 2008. Phylogenetics of Otatea inferred from morphology and chloroplast
Jata and recircumscription of Guaduinae (Poaceae: Bambusoideae). Syst. Bot. 33:277-283.
. E.V. Sosa. 2010. Delimiting species boundaries within the Neotropical bamboo Otatea (Poaceae:
molecular, morphological and ecological data. Molec. Phylogen. Evol. 54:344-356.
Ruiz-Sanchez, E.V. Sosa, M.T. Mejia-Saules, X. LondoNo, and LG. Clark. 201 1 . A taxonomic revision of Otatea (Poaceae: Bam-
busoideae: Bambuseae) including four new species. Syst. Bot. 36:314-336.
Soderstrom, T.R. and X. LondoNo. 1987. Two n
deae). Amer. J. Bot. 74:27-39.
j of Brazilian bamboos n
luseae: Guadui-
ts taxonomic identity, morphology and
3 Guadua (Poaceae: Bambusc
ion of bamboo genera and allies in a system of grass classification. Ir
Sode^rom, K.W. Hllu, CS. Campbell, and M.E. Barkworth, eds. Grass systematics and evolution. Smithsonian In
tion Press, Washington, D.CPp. 225-238. „
T«ah»s», j. «d D. Aslaos. 2004 Inventarlo de Bambues en el Peni. GT7 Contrato 012459.44)01 .00/P14I30/03, L
Tovar, O. 1 993. Las grami'neas (Poaceae) del Peru. Ruizia 1 3:1 -480.
BOOK REVIEW
Patricia Fouey. 2011. The Guide to Oklahoma Wildflowers. (ISBN: 978-1-60938-046-5, pbk). University of
Iowa Press, 119 West Park Road, 100 Kuhl House, Iowa City, Iowa 52242-1000, U.S.A. (Orders: www.
uiowapress.org, 1-800-621-2736). $39.95, 312 pp., 415 color photos, 1 color map, 6" x 9".
With permission from Great Plains Research 23(l):81-82. 2013
Oklahoma is a beautiful state and home to some 2,500 species of flowering plants. Patricia Policy has captured
m full color some 200 striking and beautiful wildflowers. From the state wildflower (Gaillardia pukhella) to the
state grass (Sorghastrum nutans), this wildflower guide covers plants growing in the Rocky Mountain foothills
in the northwest to the cypress swamps of the southeast and a lot in between.
Each species is illustrated with at least two photos, showing the habit and a close-up of the flower. Text
wise, there is a short description, followed by the geographic area of the state where the species grows, flower-
ing time, and typical habitat. Common and scientific names are provided but not the family name. The organi-
zation of the book is taxonomic with monocots first then followed by dicots with families (not noted) and
genera in phylogenetic order. This arrangement is difficult to follow for the nonbotanist. It would seem an or-
ganization by flower color would have made using the book for identification num 1
index in the back of the book is organized into four color groups and then alphabetical by ^
un ess you know the common name, finding your plant by flower color can take time.
o not find the state tree of Oklahoma (redbud, Cercis ca-
>s of striking pink flowers— is more colorful and noticeable
). I looked hard but also failed to find the state floral emblem, the mistle-
But if you love wildflowers, and Oklahoma ones at tl
ers worthwhile to have in your library and perhaps a second copy in youi
tanical Research Institute of Texas, Fort Worth, Texas, U.SA.
you’ll find The Guide to Oklahoma Wildflow-
■ — Barney Lipscomb, Bo-
CALATHEA CROATII (MARANTACEAE),
A NEW ENDEMIC SPECIES FROM PANAMA
Helen Kennedy
UCR Herbarium, Department of Botany and Plant Science
University of California Riverside
Riverside, California 92521, U.S.A.
ABSTRACT
In preparation for the Flora Mesoamericana treatment, the species of Marantaceae from Panama has been a
special focus. With considerably more collecting since publication of the Woodson and Schery (1945) treat-
ment for Flora of Panama, listing 23 species, the total has more than doubled. Besides field work, recent her-
barium studies at Missouri Botanical Garden and University of Panama have uncovered addition new species.
Currently, a total of 67 species are recognized (ca. 191 per cent increase from the original Flora of Panama
treatment). Seventeen species are recognized as endemic, including the one described herein plus two as yet
undescribed taxa.
CalatheacroatiiH.Kenn
T.B. Croat 67776 (
n«ts rhizomatous caulescen. herbs, 1.5-2.05 m; cataphylU rarrowly ovate, red-purple; steua tok purple
jw below subtending leaf, the rest green, appressed tomentose, hairs 0.5-1 mm. Leaves 2-4 ^
gle cauline leaf, rarely 2, borne above an elongate stem intemode, ca. 0.9 m; leaf sheath often aur , g
(margins) dying back in age, often tearing away from central portion, wings apicaUy red-jmtple, appreK
mentose,hairs0.5-l mm, central back portiongreenin upper 10-15cm,red-purpleb^lly,appressed^
on Wings, hairs 0.5-1 mm, 15.7-^^ in subtending leaf, 58-73 cm in basal leaves; peuo e bean^ a
-“mow groove adaxially, green, subglabrous apically, haim more subglabrous apically, bans more prevalent
longer, to 0.5 mm, jusf above sheath, 9.1-41.3 cm in subtending Wf. 32-108 cm “^110^'™“
elbptic in cross-section, olive-green, minutely tomentose m narrow band adaxia y, e r g
156
Journal of the Botanical Research Institute of Texas 7(1)
1 provided by Missouri Botanical Garden (Croat 67776, MO).
Journal of the Botanical Research Institute of Texas 7(1)
Sepal length (mm)
d one new one, Calathea cmatii, in the"C. lanicaulis Group.”
?3-23
0.2-0.25 mm, 1.5-2.8 cm; leaf blade somewhat chartaceous, ovate, apex obtuse to rounded with acumen up to
1.9-2 cm, base rounded to subtruncate shortly abruptly attenuate, 34-58 x 8-21 cm in subtending leaf, 36-95
X 8-33 cm in basal leaves (length:width ratios 2.76-4.25[-4.43];l) lateral veins 30 to 33 per 3 cm (measured at
midpoint of each side of blade), adaxial surface matte green, glabrous, midrib yellow to greenish yellow, gla-
brous, abaxial surface light green, minutely tomentose along major veins, midrib yellow with caramel tinge,
densely appressed tomentose along sides, minutely so along center in basal ‘A, appressed tomentose through-
out apically, hairs colorless, 0.25-0.7 mm. Inflorescence terminal, 1 per shoot, imbricate when young, lax in
fruit, ovoid to broadly cylindrical, 14-22 x 7.5-15.4 cm; peduncle dark red-purple, densely appressed tomen-
tose, 1.8-2.8 cm. Bracts 31-45, spirally arranged, herbaceous apically, ovate to narrowly ovate, apical ones
proportionally longer and narrower, apex acute, very tip occasionally blunted, apex recurved, apical 0.5 cm of
margin smooth, subsequent 1-1.5 cm markedly undulate, 5-7.5 x 1.8-2.8 cm, abaxial surface of bracts dark
red-purple, appressed tomentose, hairs faint straw-colored, 0.4-0.7 mm, adaxial surface dark red-purple, ba-
salmost 1-1.5 cm pale pink to white, recurved apex dark, almost blackish, purple in younger, howering inflo-
rescences, sparsely pilose near apex, glabrous basally, bracts in age, in fruit, becoming deep red with apex and
apical margins dying; bicarinate prophyll membranous, ovate to elliptic, apex obtuse, translucent cream-
white, tinged red-purple at apex, 2.1-2.6 x 1.2-1.5 cm, 0.65-0.85 cm wide, carina to carina; secondary bracts
and bracteoles unknown. Flowers open spontaneously. Sepals membranous, narrowly elliptic, cream-col-
ored, very apex tinged pale purple, 21-23 x ca. 4 mm. CoroUa tube cream-colored to pale yellow apically, ca.
im; corolla lobes subequal, elliptic, apex obtuse, margins inrolled appearing acute, light purple abaxially,
I 11-. ^ 4_5 5 Staminodes 3; outer staminode broadly el-
% dark purple, basal portion yellow, ca. 13 x 9 mm; callose
minute acumen, yellow basally, apical % dark red-purple, ca.
stamen yellow with lateral pel
faint purple to n
liptic, apex rounded, clawed at base, apical V
staminode totally callose, apex rounded or w
15x6 mm; cucullate staminode yellow, ca.
smooth, cream-colored, 2.5 x 2 mm. Seeds 5.5-6 x 4.5-5 x 4-4.5 n
.Additional spedmens: PANAMA. Bugl.: beyond Hato Chanti on trail to Hacha, heavily disturbed humid forest, growing on steep
ri2"N, 8P46'48"W, 11 Apr 2008, C. Black &A. Rodriguezd? (PMA); Cocktbehind sawmill above El
too ft, 25 Aug 1983, WJ. Kress & B. Hammel 83-1592 (DUKE 293776).
to Panama. It is known from three localities in western
one m Clnnqu. Prov, the type locality, a second in the adjacent, Comarca Ngabe-Bugle and the third
. Jc Prov. It occurs in montane wet or cloud forest habitat. The first two collections were at 1200 m and
the third at ca. 735 m elevation.
Discussion.— Calathea crontii belongs to the infonnally treated “C. lanicnulis Groun" as outlined in Flora
of Ecuador (Kennedy 1988:47). These species are characterized by their habit of several basal leaves with a
Single jhne leaf, subKntog the innorescencefs) of spimlly arranged bracts, home above an elongate stem
mternode^Speaes m the C lunnoulis Group- are predominantly South American with a single sLies. C.
guzmn„.tvfesasfarnorthasCostaRica(Kennedy,2003),fivedescribedspeciesinPanama,siainCo!ombiaand
159
nine in Ecuador. Calathea croatii is most closely related to C guzmanioides sharing a similar habit, leaf venation
pattern with 7 minor veins between the major veins and inflorescence morphology. Calathea croatii differs
from C. guzmanioides in the glabrous vs. tomentose major veins of the adaxial leaf surface, the bracts ovate to
narrowly ovate with apex acute vs. broadly ovate to broadly elliptic vdth broadly obtuse to rounded apex (bract
length to width ratios 2.62-2.75:1 vs. 0.95-1.78:1). Calathea croatii differs from the related C. allenii Woodson
and C. confusa H. Kenn. by the purple to red vs. green to yellow-green bracts with an acute, recurved vs. emar-
ginate, inflexed apex.
Etymology.— The specific epithet, croatii, is in honor of Thomas Croat, P.A. Schulze Curator of Botany,
Missouri Botanical Garden. It is a pleasure to thus recognize his contribution to our knowledge of the Panama-
nian flora and thank him for the hundreds of Marantaceae specimens he has collected throughout Central and
South America.
ACKNOWLEDGMENTS
The Missouri Botanical Garden provided support for my accommodations while working in the MO herbari-
um (organized, thanks to Olga Martha Montiel). Fred Ganders provided the travel expenses for the trips to MO
and UCR. 1 am very grateful to Carla Black for the color photos of the plant in the field and the flower close-ups
with measurement taken in her garden. 1 sincerely thank Thomas Croat for collecting this uncommon species.
1 thank Gerrit Davidse, James Solomon and Teri Bilsborrow for their help in the MO herbarium and arranging
for the type scan.Ithank Charlotte Taylorand an anonymous reviewerforveryhelpfulsuggestions and correc-
tions and especially Barney Lipscomb for his help with the manuscript. 1 am grateful to Andrew Sanders for
providing facilities at the UCR herbarium and to Teresa Salvado for providing accommodations and transpor-
tation in Riverside. Layne Huiet provided a scan of the DUKE specimen. Missouri Botanical Garden provided
the type scan.
REFERENCES
W, H. 1988. Calathea. In: G. Marling and L Andersson, eds. Flora of Ecuador 32:1 1-191. Berlings. Arlov, Sweden.
2^3. Marantaceae. In: B. Hammel, M. Grayum, C. Herrera & N. Zamora, eds. Manual de Plantas de Costa Rica,
Vol. 2. Monogr.Syst.Bot. Missouri BoLGard. 92:629-665. . me
, In: Flora of Panama. Ann. Missouri Bot. Gard. 32:81-105.
Journal of the Botanical Research Institute of Texas 7(1)
ZSCXJ DeBRECZY and ISTVAN Racz, with Kath
BOOK REVIEW
2011. Conifers Around the World: Conifers of the
1 and 2. (ISBN: 978-963-219-063-1 [vol. 1], 978-963-
219-064-8 Ivol. 2], 978-963-219-061-7 [vols. 1-2], hbk). DendroPress, Ltd., Volgy u. 12, Budapest
H-1039, HUNGARY. (Orders: www.dendropress.com, interdendr.org, info@dendropress.com). $250.00,
2 volumes, 1089 pp., 15 lbs., 474 range maps, -1300 morphological iUus., >500 taxa represented by 3700
color photos, 9 Vi" x 12 Vi".
Beautiful and spectacular color pictures of conifer landscapes, habitats, morphological characters, along with
detailed drawings of vegetative and reproductive plant parts, all add something special to this book. This
makes these two volumes a publication that everyone interested in plants should own and display on a coffee
table or bookshelf. The high quality grade of paper is Hello Silk Paper (135/M2), chlorine-free with no bleed
through. This paper has the seal of approval from the Forest Stewardship Council as a certihed environment-
friendly product. The book spine has pages with Smyth Sewn binding and attractive head and tail bands. Ev-
erything about this book is the highest quality 1 have ever seen.
The three column format of text allows for color photographic images that either are full page, two col-
umns wide, or a vertical tier of one column. In every case the color images are sharply in focus with sufficient
detail to illustrate surface features as in the Bark Gallery for different taxa. Every attempt was made to take
photographs in the field, which required in some cases travel to pristine habitats in remote places. The scope of
this book involved more than 30 years of documentation and 340,000 photographs.
The two volumes are organized geographically by 11 regions and alphabetically by taxa. Volume 1 begins
with Europe, North Africa, Anatolia, Macronesia, and Asia, including China, Taiwan, and Japan. Volume 2
covers Western and Eastern North America, Mexico, Central America, West Indies, and the Southern Hemi-
sphere that includes ChUe, Argentina, and Australia (with particular focus on Tasmania and New Zealand).
There are no keys to the genera or species. Side bars of anecdotal stories enliven specific passages.
Features of Volume 1 include an 86-page introduction that includes a number of subsections. The Age <rf
Discoveries covers the history of exploration and discovery, especially in the conifer-rich regions of China and
Mexico where new discoveries continue. Conservation includes a discussion of threats faced by conifers in
natural habitats. Plant Classification and the ChaUenges of Conifer Identification gives an overview and
history of botanical nomenclature and classification systems. Morphology contains details about habits, bark,
’ i, foliage, cones, and seeds accompanied by illustrative photographs and drawings. Distribution and Cli-
, ., mifer geographic distribution superimposed by climatic
ther section titled Conifer Families and Genera, repre-
sented by beautiful line drawings of key morphological characters and concise descriptions that facilitate
identification of genera. This section has a separate list of references for information included in the genus
descriptions.
The geographic sections follow: Europe and Adjacent Regions, Continental Asia and Hainan, Japan
and Adjacent Islands, andTahvan. Here the conifers and the regions where they are native havespecies plates
with 3-5 detailed photographs that highlight the most characteristic traits and one image showing the trees
growing in natural habitats with essential descriptive and historical information
The bulk of Volume 2 is divided into seven geographic sections. Western North America (pp. 549-652),
with 70 range maps, 15 genera, and 79 species and varieties, includes the oldest living trees represented by
Finns langacvu(BristleconePine),datingback more than 4000 years, and the tallest hving trees represented^
\i History and Conifers describe c
i. Following this extensive introdui
s 7(1): 160. 2013
TWO NEW ENDEMIC SPECIES OF CALATHEA (MARANTACEAE)
FROM ECUADOR
Helen Kennedy
UCR Herbarium, Department of Botany and Plant Science
University of California Riverside
Riverside, California 92521, U.S.A.
ganders@mail.ubc.ca
ABSTRACT
RESUMEN
Since the publication of the treatment of Marantaceae for the Flora of Ecuador (Kennedy et al. 1988) there has
been a substantial increase in field work there. Consequently a number of new species have come to the atten-
tion of various specialists, including the two being described herein. Of the total of 96 species in the 1988 flora
publication, 64 were in the genus Calathea. Of these, 32 species were noted as endemic with 29 in the genus
Calathea. However, five of these 29 species were undescribed when collected in Carchi Prov., sufficiently near
the border with Colombia to reasonably be expected there as well, though not yet documented. Currently a
total of 69 species of Calathea are recognized for Ecuador: the two new species being described, t
undescribed species plus Calathea wallisn (Linden) Regel, known from Peru. Borchsenius et a
ed less than 15% of the species of Calathea in their molecular phylogeny, t
most species to the resurrected genus Goeppertia Nees. Both of these n'
horticultural potential as they have brilliant red bracts, a
o additional
[. (2012) includ-
e confident enough to remove
cies herein have considerable
icter rather uncommon in Calathea. 1 am less
species described herein seem most hkely to be re-
teed to either the “C Imicalis Group" (fennedy et al. 1988). which includes the red-hracted C. timothei H.
ton., or C. section Calathea. which has distichous bracts. Both of these groups remain in Catehea accordmg
to Borchsenius et al. (2012).
taxonomic treatment
Calathea neillii H. Kenn., sp. nov. (Fig«i. 1, 3). Tm: ECUADOR, z
region. Rto Machinaza watershed. E of Los Encuentros. Refugio de Vida S
i0m,0y50’37"S,78”3r48”W,
20 Jul 2009. D. Ndn. W. Quizhpe 6- D. V
163
from C. trianae in the narrowly ovate to narrowly ovate -elliptic vs. elliptic leaf blade {lengthrwidth ratios 5.87-6.93: 1
; closely spaced bracts (6 vs. 3 per 5 cm).
i, ca 0.6 m high; stem dark olive-green, densely tomentose at junction to
leaf sheath, bearing sparse, scattered hairs basally, hairs 0.2 mm, stem internodes between basal leaves seri-
ceous, hairs to 4 mm. Leaves 4-7 basal vshth a single cauline leaf borne above an elongate stem intemode, ca.
40 cm; leaf sheath not auriculate, sheath of subtending leaf ca. 7 cm with wings and base reddish purple, the
center adaxial portion deep olive-green in upper 3/4, tomentose at base, subglabrous to glabrous apically,
sheath of basal leaves 21-23 cm, tomentose apically on wings, margins densely appressed tomentose, hairs to
0.5 mm, hairs sparse on center back and wings basally; petiole green, glabrous, subglabrous just above sheath,
ca. 3.5 cm in subtending leaf, (6.5-)12-18.4 cm in others; pulvinus 1-1.4 cm, olive-green, minutely tomentose,
hairs to 0.5 mm, densely so adaxially, sparsely so abaxially and on sides, acuminately extended adaxially to 0.5
cm; leaf blade not ridged, the tissue between the raised major veins forming a smoothly curved shallow depres-
sion, narrowly ovate to narrowly ovate-elliptic, left side broader than right, apex acuminate attenuate, base
obtuse to 90°, shortly abruptly attenuate; 24.5-31.4 x 3.6-4.6 cm (length:width ratios 5.87-6.93:1), generally 7
minor veins between major veins, vein angle from midrib 40° to 50°, lateral veins ca. 13 per 2 cm, cross-veinlets
30 to 40 per 5 mm (veins measured at midpoint of each side of blade), adaxial surface semi-shiny deep green,
major veins minutely tomentose, in the basalmost 1 cm, the hairs are also present in the area between the veins,
the marginal 2 mm of right side of blade minutely tomentose with the acumen densely so, hairs colorless,
0.1-0.3 mm, midrib paler, whitish green, minutely tomentose, abaxial leaf surface light green, glabrous, midrib
glabrous except basal 1.5 cm tomentose where pulvinar cells extend onto midrib. Inflorescence terminal, 1
per shoot, ellipsoid-rectangular, ca. 5 x 1.5 cm; peduncle rose-red, densely appressed tomentose apically at
junction to bract, 1.1-1.4 cm. Bracts 7-8, distichous, subrotund, apex retuse, margins and apex straight, not
recurved, 2.1-2.3 x 1.9-2.2 cm, each subtending up to 3 or more flower pairs, abaxial surface rose-red, gla-
brous; bicarinate prophyll membranous, ovate, apex obtuse, translucent rose-red, glabrous except appressed
pUose abaxially in basal 1/3-1/2 adjacent to carinas, hairs 0.7 mm, 1.9-2.1 x 0.65-0.7 cm, 0.5-0.55 cm wide,
Carina to carina; secondary bracts absent; bracteoles 2 per flower pair, membranous, medial, 1 chanelled, 1
carinate, narrowly elliptic, glabrous, 1.3-1.4 (-2) x 0.3-0.35 cm. flowers opening spontaneously. Sepals
membranous, narrowly elliptic, broadly obtuse, glabrous, 14.5-15.5 x 3-3.5 mm. Corolla yellow, tube gla-
brous, 17-18 mm- corolla lobes subequal, elliptic, 6-8 x 2.5-3 mm, obtuse, margins inrolled appearing acute,
sparsely pilose in apical 1/4, hairs 0.2 mm. Staminodes 3, yellow; outer staminode obovate, emarginate; cal-
lose staminode petaloid apically, ca.6.5 mm; cucullate staminode 3.5-4.5 mm; stamen with lateral petaloid
appendage; anther 1.5-2 mm; ovary smooth, glabrous, 2-2.5 mm. Capsule unknown.
Distribution and habitat.-Calathea neillii is endemic to Ecuador, known only from the type locality m the
Cordillera del Condor region of Prov. Zamora-Chinchipe. It occurs at 1560 m elevation in the shady understory
ofwel forest habitat . The type was collected in nower in July. , . „ ,
Discussion.-Cnlntltea neillii, shares a similar general habit of several basal leaves and an inflorescence of
tiisUchous bright red bracts borne above an elongate stem intemode with both C/redii and C. t^nne. Calm ea
ndlliidiffersfrLc. trionaein the narrowly ovate tonarrowlyovate-elliptic vs. ellipticleafbfodelengA^^^^^
ratios 3.87-« 931 vs vs 3 35-4 38:1), minor veins usually 7 vs. 3 between major veins, the petiole glabrous vs^
pilose the shorter neduncle (1 1-1 4 vs 10-14.6 cm), and the relatively broader inflorescence (lengthwidth
rau« 2.4-2.91 vs. 6 0-7.33:1) with more closely spacedbracts (6 vs. 3 per 5 cm). It dillers from C./re«n *e
smooth rather than strongly cormgated leaf surface, the lower vein angle from nudnb (40 -50 vs. 73 -79 ),
the broader vein spacing (ca. 13 vs. 19-24 per 2 cm) and the yellow vs. purple petals
--Pe-fromLpicalwetforests —
»ricallycontrTlleLLacters in onler to preventsingkgenemutationsor morphs inapolymorphiccharacw
from being described as species.
Journal of the Botanical Research Institute of Texas 7(1)
Etymology.— Jht specific epithet, neiUii, is in honor of David Neill, Director of Investigacion, Universidead
Estatal Amzonica, Puyo, Ecuador, and principal investigator on the Ecuadorian “Cordillera del Condor proj-
ect.” As co-founder of the Estacion Biologica dejatun Sacha he has been and is a prolific collector of Ecuadorian
plants, including many undescribed species, of which this one, honoring him, is an especially attractive one.
Iconography.— Images of type scans of Calflthea trianae, to compare with those of C. neilii and Cfredii are
accessible through JSTOR at http:plants.jstor.org/search?plantName=”Calathea+trianae”&syn=l. A good im-
age is that at NY of an isotype.
i,00-=05'S,77“24'W,6
nzomatous, caulescent herbs, 0.9-1.2 m; roots stiff, v
glabrous below; cataphylls stiff, papyraceous, narrowly ova
rough to the touch, hairs, ca. 0.5 mm on i
a single cauline bladeless sheath above ai
ry; stem sericeous apically adjacent to cataphyll,
, apiculate, appressed tomentose to subhispid,
i. 26 cm. Leaves 3-8 basal and
t; leaf sheath not auriculate, wings
:r back slightly scabrid; bladeless sheath (cataphyll) subtending inflores-
cence 9.5-10.5 cm long, sheath of basal leaves 23-45.5 cm; petiole green, appressed tomentose adaxially in
apical 1 cm, minutely scabrid below to nearly smooth above sheath, (9-)16.5-62 cm; pulvinus round in cross-
section, 1.3-3.8 cm, olive-green, tomentose adaxially, the rest glabrous, hairs straw-colored 1-1.5 mm; leaf
blade plicate, markedly ridged with major veins on raised ridges, minor vein in depressions between, charta-
ceous, narrowly elliptic, apex acuminate-attenuate, slightly eccentric, base obtuse to 90°, 31.5-48 x 5.2-10 cm
(lengthiwidth ratios [4.8-] 5.24-7.45:1), lateral veins 19 to 24 per 2 cm, cross-veinlets ca. 50 per 5 mm (veins
measured at midpoint of each side of blade), vein angles from midrib 73°-79°, measured at midpoint of blade,
adaxial surface deep green, glabrous except sparsely pilose at base and along margin of wider side, midrib deep
green, glabrous basally, apical 10 cm pilose, abaxial leaf surface grey-green, scattered pilose, each hair on a
raised basal cushion of cells, hairs straw-colored, midrib yellow-green to caramel-colored, appressed pilose.
Inflorescences 1-3 per shoot, 1 terminal, subsequent ones borne in axil of bladeless sheath, imbricate, rectan-
gular, complanate, 5.5-8.3 x 3.1-4.6 cm; peduncle dark purple-brown, tomentose in apical 1-1.5 cm, glabrous
basally, 14-21.5 cm. Bracts 10-13, distichous, herbaceous, very broadly obovate to rotund, apex emarginate in
ermost the largest, each bract subtending up to 6 or more flower pairs, abaxial and adaxial surface of bracts red,
glabrous; bicannate prophyll membranous, ovate-triangular, apex obtuse to 90°, apiculate, translucent red,
1.8-2.2 x 1.1-1.2 cm, 0.45-0.6 cm wide, carina to carina; secondary bract membranous, ovate, apex acute,
red, glabrous, 1.8-2 x 0.8-0.9 cm; bracteoles 1-2 per flower pair, 1.2-1.4 x ca. 0.15 cm, membra-
cannate, or if 2, the second channeled, very narrowly elliptic to linear, glabrous. Flowers open-
membranous, very narrowly elliptic to sublinear, apex obtuse, 15-16 x 2 mm. Co-
rn, glabrous except for a few appressed hairs in a line in apical 1/2, hairs 0.7-1 mm;
iptic, 6-6.5 X ca. 2 mm, acute, apical 2/3 purple, cream-colored toward base, gla-
3; callose staminode, ca. 6 x 2.5 mm; cucullate staminode ca. 4 mm data unavailable on
node and stamen due to condition of specimen. Ovary smooth, pilose at apex, hairs colorless to 2
165
riBotanial Garden (Ato»7537, MO).
Journal of the Botanical Research Institute of Texas 7(1)
Kennedy, New species of Calathea from Ecuador
167
Distribution and habitat. — Calathea jredii is endemic to Ecuador. It occurs in eastern Amazonian Ecuador in
Napo and Sucumbios Provinces from 800-1450 m in premontane rain forest habitats. Flowering material was
collected in December and February as well as in August.
Discussion. — Calatheafredii shares the similar habit and inflorescence of distichous bright red bracts with
the related C. neillii and C. trianae. It would key out in Flora of Ecuador (Kennedy 1988: 47) under lead 31A
because of the distichous bracts. It differs from C. neillii in the strongly corrugated vs. smooth leaf surface, the
higher vein angle from midrib (73'’-79° vs. 40°-50°) the narrower vein spacing (19-24 vs. ca.l3 per 2 cm) and
the purple and white vs. yellow petals. Calathea jredii differs from C. trianae by the strongly corrugated vs.
smooth leaf surface, narrowly elliptic vs. elliptic leaf blade (lengthrwidth ratios [4.8-15.24-7.45:1 vs. 3.55-
4.58:1), and the purple and white vs. yellow petals. Also, it differs from C. neillii and C. trianae in lacking a
bladed leaf subtending the inflorescence. Here the leaf is bladeless, reduced to essentially to the leaf sheath.
Etymology.— The specific epithet, /redii, is in honor of Fred R. Ganders, professor emeritus and former
director of the UBC herbarium, in gratitude for his help writing the keys for the Flora of Ecuador treatment,
supporting my research trips, paying for needed equiptment to carry out such work, as well as for many other
trips to botanize and sample wine.
ACKNOWLEDGMENTS
The Missouri Botanical Garden provided support for my accommodations while working in the MO herbari-
um (organized, tbanks to Olga Martha Montiel). Fred Ganders provided the travel expenses for the trips to MO
and UCR. 1 thank David Neill for collecting both C. neillii and C jredii and for the use of his photographs of C.
neillii. I thank Robin Foster for the use of his color photographs of C jredii, his collections and scans of the Field
Museum specimens of it and John Kress for the color slide of C. trianae. 1 thank Mary Merello for help in the
MO herbariumandarrangingfor the type scans of C. neillii and C.^edu.ThankstoGordonMcPherson and an
anonymous reviewer for their helpful corrections and suggestions and to Barney Lipscomb for providing per-
tinent literature as well as helpful advice. I am grateful to Andrew Sanders for providing facilities at the UCR
herbarium and to Teresa Salvado for providing accommodations and transportation in Riverside. Missouri
Botanical Garden provided the type scans.
rantaceae). Syst. Bot. 37:620-635.
Judo, W.S., C.S. Campbell, E.A. Kellogg, P.F. Stevens, and
Third Edition. Sinauer Associates, Inc., Sunderland, MA.
references
Prince. 2012. Molecular phytogeny and redefined generic limits of Catathea (Ma-
Donoghue. 2008. Plant systematics: a phylogenetic approach.
BOOK REVIEW
<1 Musial, ED. 2011. Conifers Around the World: Conifers of the
12. (ISBN: 978-963-219-063-1 [vol. 1], 978-963-
219-064-8 [vol. 2], 978-963-219-061-7 [vols. 1-2], hbk). DendroPress, Ltd., Volgy u. 12, Budapest
H-1039, HUNGARY. (Orders: www.dendropress.coin, interdendr.org, info@dendropress.com). $250.00,
2 volumes, 1089 pp., 15 lbs., 474 range maps, -1300 morphological illus., >500 taxa represented by 3700
color photos, 9 W'x 12 ‘/i".
Scquoiadendwn giganteum (Sierra Redwood). Pinus edulis (Pinyon Pine), as the name implies, has edible seeds
important as a food item for Native Americans and wildlife. The section covering Eastern North America (pp.
653-701) has 32 range maps, 10 genera, and 34 species and varieties. Mexico and Central America (pp. 702-
830) has 80 range maps, 8 genera, and 95 species and varieties. Abies hidalgensis (Hilalgo Fir), Abies neoduran-
gensis (New Durango Fir), and Abies zapotekensis (Zapotek Fir) were described as new species from this area in
1995. These are distinct species but were isolated as small populations in more remote areas and were previ-
ously undiscovered. This emphasizes that even tree species that are conspicuous life forms because of their
larger size still represent the lure of the unknown as undiscovered new species. Another interesting story is
that of Pinus maximartinezU (Big Cone Pinyon Pine). Described as new in 1964, it was actually first discovered
at a food market in Juchipila, southern Zacatecas, Mexico, where Jerzy Rzedowski found its edible seeds being
sold as a delicacy. Additional sections of Volume 2 include the West Indies and Bermuda, Chile and Argen-
tina, Australia and Tasmania, and New Zealand.
There is also a unique Bark Gallery chapter containing 646 color images taken in natural settings; a
6-page graphic essay of the pollen cones of each genus; and 474 distribution maps that show a visual image of
the natural range of each species discussed. The Appendix in Volume 2 includes images and concise descrip-
tions of 28 additional taxa, a glossary, references, lists of the covered conifers by scientific and common names,
and a quick finder index.
This book is written in nontechnical narrative that will have broad appeal to a wide readership including
the lay public, general botanists, foresters, horticulturists, and dendrologists. College, university, and public
libraries will want to have this affordable book included in their biodiversity section. This book will be a valu-
able addition to botanical gardens, herbaria, nature centers, and national forest and national park visitor cen-
ters.—Harold W. Keller, Research Associate, Botanical Research Institute of Texas, Fort Worth, Texas, USA.
EL GENERO POLYPOGON (POACEAE: AGROSTIDINAE) EN CHILE
Victor L. Finot*,
Leslie Contreras
Universidad de Concepcion
Casilla 537, Chilian, CHILE
Wilson Ulloa, Alicia Marticorena,
Carlos M. Baeza & Eduardo Ruiz
Departamento de Botdnica
Facultad de Ciencias Naturales y Oceanogrdficas
Universidad de Concepcion
Casilla 160-C, Concepcidn, CHILE
INTRODUCClON
El gencro Pdypogon Desf. (gr., polys > muchos; pogon = bart«) (Walson & Dallwitz 1992) “ >>
rt™ Agrostidinae, Iribu Poeae y subfamilia Pooideae de la familia Poaceae (Soreng et al. 2003). Clayion &
Kmvoize (1986) lo sitOan en la Iribu Aveneae subtribu Alopecurinae que reune los generos con ^
unillonis previameute incluidos en la iribu Agroslideae. la subtribu Agrostidinae Soie"g “
«« 16 generos y cerca de 550 especies (Saarela et al. 2010), es el grupo mis diverse del clado con ADN plas-
Mal tipo AvenLe- de la iribu Poeae, que reune ademas a las subtribus Aveninae, Bruinae, Pbalandmae y
Torreyochloinae (Soreng etal. 2012). , y a i
AIgnnos de los generos incluidos en Agrostidinae son extremadamente diversos (v.gr. AgrMis L. ca. 220
W-; Calomagrostis Adans., ca. 270 spp.) y su taxonomia aun no esia completamente resue ^ ^
“pgriieticas ban stdo poco estudiadas y la circunscripciOn de los generos aparece como el pnncrpal problema
^onomicoen Agrostidinae (Saarela etal. 2010). u ■ r /x i
Polypogon con unas 26 especies de ireas templadas y templado-calidas de ambos hemisferios (Tzvelev
>«83;IWoiae 1998; Giialdo-cTftas 2004; Lu &Pbillips 2006), laet^rX
Ayosis L (Oayton & Renvoiae 1986), del cual difiere principalmenle por la ^ „
170
Journal of the Botanical Research Institute of Texas 7(1)
^stas en la inflorescencia cuando caen los frutos maduros (Clayton & Renvoize 1986). Ademas de estos carac-
teres, tienen algiin interes taxonomico la estructura de la palea, la micromorfologia de la lemma y la morfologia
de la cariopsis (Rugolo 1982; Finot et al. 2011b). El hlbrido intergenerico xAgropogon lutosus (Poir.) P. Foura
[=Agrostis stolonifera L. x Polypogon monspeliensis (L.) Desf.], con caracteres intermedios, fue citado para Chile,
bajo xAgropogon littoralis (Sm.) C. E. Hubb., por Rugolo & Molina (1997b), a partir de material recolectado en ,
la Region de Los Lagos (Osomo y Chilo^. I
Algunos autores reconocen a Chaetotropis Kunth como un genero diferente de Polypogon (v.gr. Kunth j
1835; Desvaux 1854; Pilger 1920; Skottsberg 1921; Bjorkman 1960; Nicora 1970, 1978, 1993; Marticorena & |
Quezada 1985; Nicora & Rugolo 1987; Morrone et al. 2005; Rugolo et al. 2008, Zuloaga et al. 2008, Rugolo '
2012a), del cual difiere fundamentalmente por la longitud de la palea con respecto al largo de la lemma (en i
Polypogon la palea es tan larga como la lemma; Chaetotropis tiene la palea Vi del largo de la lemma o menos) y
por la epidermis de la lemma provista de una malla de engrosamientos de las paredes periclinales de las cdulas
epidermicas denominada “trichodium net” que esta presente en Chaetotropis pero esta ausente en Polypogon s.s.
(Bjorkman 1960, Rugolo 2012b, Favret et al. 2007). Otros autores incluyen a Chaetotropis en Polypogon sect, i
Polypogonagrostis Asch. et Graebn. (v.gr. Trinius 1840; Steudel 1854; Martins & Eichler 1893; Stuckert 1904; |
Hitchcock 1931, 1951; Macbride 1936; Rosengurtt et al. 1970; Navas 1973; Gould & Shaw 1983; Tzvelev 1983; I
Muller 1985; Clayton & Renvoize 1986; Tovar 1993; Renvoize 1998; Soreng et al. 2003; Barkworth et al. 2007; |
Finotetal. 2009, 2011a, b).
Soreng et al. (2003) citan 11 especies de Polypogon para el Nuevo Mundo, una de las cuales (P. parvulus
Roseng., B.R. Arrill & Izag.), se encuentra solamente en Argentina y Uruguay (Rosengurt et al. 1970). Las otras !
10 especies han sido citadas para Chile, incluyendo P. maritimus Willd., cuya presencia en Chile se descarta !
aqui; una especie, P. linearis Trin., es endemica de Chile; 6 especies son nativas no endtoicas; P. australis
Brongn., P. chiicnsis (Kunth) Pilg., P. elongatus Kunth, P. exasperatus (Trin.) Renvoize, P. imberbis (Phil.) Johow,
P. interruptus Kunth y 2 son introducidas: P. monspeliensis (L.) Desf. y P. viridis (Gouan) Breistr.
Muller (1985) divide el genero en 2 secciones; Polypogon sect. Polypogon y P. sect. Polypogonagrostis Asch.
& Graebn. La seccion Polypogon, comprende plantas perennes o anuales, con glumas aristadas desde el apice o
con el apice bilobado y con la arista naciendo entre los lobulos, lemma de hasta 1,5 mm de largo, epidermis de j
la lemma sin trichodium net, y palea tan larga como la lemma. Esta seccion, tipificada por P. monspeliensis, in- |
cluye las especies usualmente tratadas bajo Polypogon s.s. (P. australis, P. interruptus, P. linearis y P. viridis). La !
seccion Polypogonagrostis comprende plantas perennes con glumas aristadas desde el apice, lemmas de ca. 1,5
mm de largo, palea de del largo de la lemma y epidermis de la lemma con trichodium net (Muller 1985).
Polypogon sect. Polypogonagrostis, tipificada por P. elongatus Kunth [= Ch. elongata (Kunth) Bjorkman], con-
tiene las especies tratadas por algunos autores bajo el genero Chaetotropis: P. chUensis (= Ch. chilensis Kunth), P-
elongatus [= Ch. elongata (Kunth) Bjorkman], P. exasperatus [= Ch. exosperata (Trin.) Bjorkman] y P. imberbis 1=
Ch. imberbis (Phil.) Bjorkman] (Muller, 1985; Finot et al. 201 lb).
En Chile continental, el genero se encuentra a lo largo de todo el pais y en Chile insular se le encuentra en
el Archipidago de Juan Fernandez (Rugolo & Molina 1997a; Baeza et al. 2007); crecen en lugares hiimedos, !
arenosos o salinos junto a cursos de agua (Glenn 1987), distribucion que a menudo se correlaciona con la dis- ^
persion hidrocora de las semillas (Li & Qiang 2009).
Trinius (1824) describio la primera especie chilena del genero, P. crinitus Trin basada en pi
por Eschhoh & Chamisso en CMe, sin indicar una tocalidad especlfica. El n^mbre propue'ste
unto.^oposreriordeP.crimrus(Scteb.)Nnt..,utUlzadoenl818porNntullpa,a.ransfcrirPblc»mcr«-
tumSchreb.algeneroPolpiwgoa.Kunlze(1898)lransfiri61aespeciedeTriiuusaPli,toralisSm coneltangode
vanedad [P littoralis var. crimlas (Trin.) Knnuel niiemras ,ne Hackel (1906) U uala como nna varicdad de P
m.errnplus|Pimer,riptmvar.cri„i,as(Trin,)Hack,l.Actnalmen,c,Pcr«mTrin.secor«ideraunsinOn^^^
tr^"*"'”' "“'''i’s especies para Chile; P a//inis Brongn. (. P. tiongalus) y P
Rnotetal, El genero Polypogon en Chile
Kunth (1830) describio Chaetotropis chilensis basandose en material recolectado en “La Conception”; esta
especie fue posteriormente transferida a Polypogon por Pilger en 1920.
Trinius (1836) describio P. linearis, basandose en material recolectado por Poeppig en 1827, en Concon.
Describio tambien P. monspeliensis var. interrupta (Trinius 1836), recolectada por Poeppig en “Los Andes de
Santa Rosa” (Los Andes). El tipo se encuentra en el herbario del Institute Komarov (LE) pero el estatus del
nombre aiin no ha podido ser esclarecido.
Trinius (1841) describio Agrostis exasperata basado en ejemplares recolectados por d’Urville en Concep-
cion. Esta especie fue posteriormente transferida por Renvoize (1998) al genero Polypogon [P. exasperatus
(Trin.) Renvoize]. Renvoize (1998) tambien transfirio Agrostis hackelii R.E. Fr.aPolypogon [P. hackelii (R.E. Fr.)
Renvoize], que se considera un sinonimo de P. exasperatus.
Joseph Dalton Hooker (1846) describio Polypogon chonoticus Hook, f., del Archipielago de los Chonos,
sobre la base de material recolectado por Charles Darsvin en diciembre de 1834; este nombre es un sinonimo
deP. australis Brongn.
Steudel (1854) describio Agrostis asperigluma sobre la base de material recolectado por Lechler en Huiti,
Region de Los Lagos, Chile. Esta especie fue posteriormente transferida a Chaetotropis por Nicora (1978) [Ch.
asperigluma (Steud.) Nicora = P. exasperatus (Trin.) Renvoize]. Steudel (1854) tambien describio P radicans
Steud. (= P australis), basado en material recolectado en Valdivia por Lechler. En el trabajo de Steudel (1854) se
encuentra tambien la descripcion hecha por Nees de P. longiflorus Nees ex Steud. (= P. linearis), basada en mate-
rial recolectado en Valparaiso por Cumming.
Emile Desvaux (1854) describio P. elongatu
en Santiago. Otras variedades han sido descrita
var. patagonica Speg.), lo que demuestra la vai
describio ademas dos variedades de P.
r. strictus, sobre la base de
erial recolectado por C. Gay
Hack, var. muticus Hack, y
riabilidad de los caracteres en esta especie. Desvaux (1854)
breviaristatus y var. longearistata).
variedades ha sido reconocida en los tratamientos modemos de Polypogon. Desvaux (1854) reconoce Polypogon
y Chaetotropis como gdneros separados; bajo Polypogon incluye 6 especies: P. crinitus (=P. australis), P. interrup-
tus, R chonoticus, P. monspeliensis, P. linearis y P elongatus y bajo Chaetotropis incluye solo a Ch. chilensis.
El mayor numero de especies fue descrito por R.A. Philippi. No obstante, la mayoria no ha sido recono-
cida por los botanicos modernos; [1859] Chaetotropis latifolia Phil. (=P chilensis); [1860] P. cachmalensis Phil.,
P microstachys Phil. (=P interruptus); [1891] P tarapacanus (=P. interruptus); [1896] Chaetotropis araucana (=P.
exasperatus), P brachyatherus (=P. interruptus), P breviaristatus (=P interruptus) yP. purpurascens (=P australis).
En este trabajo se revalida P cachinalensis Phil.
Pilger (1920) transfirio Ch. chilensis Kunth a Polypogon. Bjorkman (1960) transfirio a g^ero Chaetetr^s
tees especies: Agrostis exasperata Trin. [Ch. exasperata (Trin.) Bjorkman], Agrostis hackelii [Ch. hackehi (RE.
Fr.) Bjorkman] y Nowodworsfeia imberbis Phil. [Ch. imberbis (Phil.) Bjorkman]. Esta ultima especie habia sido
Previamente transferida a Polypogon por Johow (1896). .
Nicora (1978) transfirio Agrostis asperigluma Steud. al gCnero Chaetotropis (Ch, asperigluma (Steud.)
Martic'^M& Quezada (1985) catalogan 7 especies de Polypogon en Chile: R australis^P.
linearis, P lutosus (Poir.) Hitchc. (= x Agropogan lutosus), P. maritimus, monspe lensis y . ^miver ici
fforsk.) Hyl. (= P viridis). Reconoceu el gdneru Chaetotropis con 4 especKs: Ch. asperigluma (- P. exasperatus),
Oi chilensis (=P. chilensis), Ch. elongatu (-P. elongatus) yCh imheitisl-P.im^s).
Soreng etui. (2003) no recouoceu el genero Chaetotropiseiucluyenen Polygon s.l.imraClule:P.^stj
lh.P. chiteis, P. eLsperatus, P. imherfiis, P. interruptus, P. linearis, P. maritimus, P. monspehens.syP. .uidis. No
"zXgat3 Chaetotropis e incluyen para Chile las siguientes especies
^''.iXpan!,Pr™P.elon^
t’rnns,?. monspeliensis yP. viridis. rh
r-...- ./ a- ron las sieuientes especies para Chile- a<inermlumn Ch
Journal of the Botanical Research Institute of Texas 7(1)
chilensis, Ch. elongata var. eiongata y Ch. imberbis var. imberbis. Bajo Polypogon reconoce P. australis, P.
brachyatherus, P. interruptus, P. maritimus, P. monspeliensisy P. viridis.
En este trabajo reconocemos 11 especies y dos variedades de Polypogon para Chile. Las especies se
describen e ilustran y se entregan mapas de su distribucion, una clave para el reconocimiento de las especiesy
material estudiado de cada una de ellas.
MATERIALES Y METODOS
Se esludiaron las colecciones de Polypogon de los herbarios BA, CONC, SGO y SI (Holmgren et al. 1990), el
material tipo depositado en SGO y las imagenes de los tipos disponibles en JSTOR Plant Science (http://plants.
jstor.org/) de los herbarios B, BAA, BAB, C, CGE, F, G, HAL, K, LP, MO, P, SGO, TDC y W. Al mismo tiempo, se
consultaron las descripciones originales de las especies. Las observaciones se realizaron con un microscopic
estereoscbpico Zeiss SV 8 equipado con ocular micrometrico. Para diferenciar las especies se confecciono una
clave dicotomica. Las especies fueron descritas, ilustradas y se entrega la sinonimia de cada una de ellas, la
iconografia, distribucion geografica y el material examinado.
RESULTADOS
Morfologia
1. ado de vida.— Las plantas pueden ser anuales o perennes. Todas las especies asignadas a la seccion Polypo- j
gonagrostis son perennes con la excepcion de P. chilensis. De la seccion Polypogon son anuales P. linearis y P ,
ranaceas, de (0,5-) 1-6 mm de largo, truncadas. Las laminas son planas, glabras o escabrosas.
inflorescencia es una panicula subespiciforme o laxa (abierta o contraida), raramente
veces con tonaUdades purpureas. Las inflorescencias de mayor tamano
cm de largo, ca. 4 cm de ancho) y las mas pequenas en P. cachinalensis
(1 ,2-2,3 cm de largo, 0,4-0,5 cm de ancho). La compactacion de la inflorescencia ha sido considerado un
caracter con valor taxonomico en Polypogon (Keshavarzi et al. 2007). Paniculas laxas, abiertas (16-17 cm de
largo, 4-5 cm de ancho), se encuentran en P exasperatus y P. magellanicus. El raquis puede ser glabro o
fragmento de pedicelo de largo variable segiin la especie. En algunas especies el artejo que acompana a la es-
piguilla es muy breve (v.gr. P. monspeliensis, 0,l-0,2 nun), en otras corresponde a casi todo el pedicelo (v.gr. P
magellanicus, 1-4 mm). En la seccion Polypogon, el artejo alcanza 0,43 mm en promedio mientras en la seccion
Polypogonagrostis promedia 1,5 mm.
5. Glumas.— Las glumas son aproximadamente iguales en forma y tamano, ambas mayores que el antecio,
aristadas y usualmente escabrosas en el dorso. Las especies de la seccion Polypogonagrostis tienen glumas mas
largas (1, 6-5,0 mm, prom. 2,55 mm) que la seccion Polypogon (1,3-3,1 mm, prom. 1,90 mm). La gluma inferior
suele ser levemente mas larga que la superior en la seccion Polypogonagrostis. La especie con glumas mas cortas
es P cachinalensis (l,3-2,0 mm); las glumas m^ largas se encuentran en P. magellanicus (3,2-5,0 mm).
6. Arista de las ghnnas.— Las glumas pueden ser aristadas o mtiticas, a veces mucronadas. Polypogon chilen-
sis, P. exasperatus var. exasperatus, P. magellanicus y P. viridis carecen de aristas bien desarrolladas. Las aristas
mas largas (ca. 15 mm) se observaron en P australis y las mas cortas (0,3-0,5 mm) en P. cachinalensis. La arista
nace en el apice de la gluma o inmediaiamente por debajo del mismo. En P. elongatus, el cuerpo de la gluma se
extiende insensiblemente para formar una arista apical. Las caracteristicas de las glumas, especialmente de la
ansta (longitud y posicion) han sido utilizados como caracteres de valor taxonomico (Keshavarzi et al. 2007).
r que las glumas, hialina, glabra, 5-nervia. El dpice es truncado
e alia del apice como breves aristulas, todas del mismo largo o bien las exte-
s mutica o posee una arista inserta cerca del apice o en la zona media del dorso;
Finotetal., El genera Polypogon en Chile
ma (trichodium net). — El denominado ""trichodium net”
corresponde a engrosamientos en las paredes periclinales de las celulas epidermicas de la lemma. La presencia
de esta malla esta ausente en todas las especies pertenecientes a la seccion Polypogon y esta presente en la sec-
don Pofypogonagrostis. En P. magellanicus y P. exaspercUus var. kuntzei los engrosamientos periclinales estan
ausentes (Finotetal 2011b).
9. Palea. — La palea es hialina, 2-nervia y puede ser tan larga como la lemma (seccion Polypogon) o mas corta
que esta (seccion Polypogonagrostis), alcanzando aproximadamente % del largo de la lemma. La longitud de la
palea en relacion con la longitud de la lemma ha sido considerado un buen caracter para reconocer a Chaeto-
tropis como un genero independiente de Polypogon (v. gr. Nicora & Rugolo 1987).
:ariopsis puede ser ovoidal o elipsoidal, con surco ventral mas o menos marcado; escudete
1/4 de la longitud de la cariopsis; hilo punctiforme. Nicora & Rugolo (1987)
la morfologia de la cariopsis para separar Chaetotropis de Polypogon.
TRATAMIENTO TAXONOMICO
. 1:66. 1798. Especietipo: P. monspeliensts (L.) Desf.
;. Canas de 10 a 120 cm de alto. Llgulas membranaceas, glabras o
escabrosas; Uminas planas. Espiguillas unifloras, pediceladas, desprendiendose con el pedicelo o un frag-
mento del mismo de largo variable segiin la especie. Glumas mas largas que el antecio, subiguales, 1-nervias,
escabrosas, el apice agudo o bilobado, terminado en una arista dorsal o subapical de largo variable, a veces
ausente. Lemma hialina, glabra, 5-nervia, con el apice truncado, denticulado, mutico o con arista inserta cerca
del apice o en la zona media del dorso de la lemma, a veces los nervios laterales prolongados mas alia del apice
enaristulas breves. Palea hialina, 2-nervia tan larga como la lemma o menor que esta. Flor perfecta; lodiculas
glabras, agudas o redondeadas en el apice; estambres 3; ovario con estilos cortos y estigmas plumosos. Cariop-
sis ovoide o elipsoide; surco ventral mas o i
longitud de la cariopsis; hilo punctiforme.
Polypogon Desf., Fl. /
larcado; escudete embrional aproximadameni
Distribuci^n geogrdfica.-Genero cosmopolita, principalmente de regiones templadas o templado-calidas
aunque en las regiones tropicales puede encontrarse tambien en las montanas (Kabeer & Nair, 2007). En la
Argentina, Zuloaga et al (2008) reconocen 7 especies: P. australis, P. brachyatherus Phil, P. mterruptus, P
I^aris, P maritimus, P monspeliensts y P. nridis. En Bolivia, Renvoize (1998) cita 5 especies: P
^peratus, P. hackelii (= P. exasperatus), P interruptus, P monspeliensts y P vindis. En Peru, Tovar (1993) senala
4 especies: P. elongatus, P. interruptus, P monspeliensts y P. vindis (bajo P. semirerticiUatus) a las que debe
agregarseP.exasperat«scuyosin6nimo,ViI/aac«tigI«miaSteud.exLechler,fuedescrim
suelos arenosos y sobre todo hiimedos. En Chile se distribuye desde la Region de Anca y Pannacota, hasta la
Region de Magallanes y Antartica Chilena, incluyendo algunas especies en el Archipielago de Juan
Fernandez.
J. Plantas mayores (16-125 cm de alto); paniculas contraidas mayores de 3 cm; glumas agudas o
. Polypogon australis Brongn., Voy. Monde 2:21. 1829. Tipo
; de la ville de la Conception, au Chili. Jan 1825. D. U
Sci. Math.. Seconde Pt. Sci. Nat. 6.4(3-4):257. 1841
elago de los Chonos. Cape Tres Montes. Dec 1834. C
Perennes; canas de 35-125 cm; entrenudos de 7-17 cm de largo; nudos 2-3. Vainas de 6-14 cm de largo, mas
cortas o tan largas como los entrenudos, glabras o levemente escabrosas hacia el apice; llgulas de (0,5-)2-4 mm
de largo, levemente escabrosas en el dorso, el apice truncado y denticulado; laminas planas de 5-16 cm de largo
por 1-8 mm de ancho, escabrosas en ambas caras o, menos frecuentemente, glabras. Paniculas contraidas a
subespiciformes, de 6-18 cm de largo por 2-4 cm de ancho, verde-amarillentas a purpureas; raquis glabro.
Espiguillas se desprenden con un fragmento de pedicelo de 0,5-1 mm de largo; pedicelos de 0,8-2 mm de lar-
go, escabrosos; glumas subiguales de l,5-2,5 mm de largo por 0,4-0,6 mm de ancho, escabrosas en el dorso,
apice agudo o levemente lobado, provisto de una arista subterminal de 5-10(-14) mm de largo, usualmente
ondulada; lemmas hialinas, de 1-1,8 mm de largo, con el apice truncado, 5-nervias los nervios laterales e in-
termedios prolongados en cuatro aristulas de las cuales las dos extemas son mas largas que las internas; arista
de 3-8 mm de largo, inserta en el tercio superior del dorso de la lemma, a 0,2-0,35 mm del apice, ondulada
menos frecuentemente recta, exserta; callo glabro; palea de 0,9-l,4 mm de largo, tan larga como la lemma;
lodiculas de 0,5 mm de largo, hialinas, agudas; anteras de 0,4-0,8 mm de largo. Cariopsis elipsoidal, de 0,9-1.2
mm de largo por 0,25-0,4 mm de ancho, con marcada depresion ventral; endosperma blando.
konografia.-mchcock (1951: 365, fig. 513); Nicora (1978: 396, fig. 262a-e); Rugolo (2012b: 341).
Fenologia. — Florece y fructifica entre septiembre y marzo.
Distribucion geogrdficay habitat.— Habita en Argentina, Chile, Mexico y USA. Crece en zonas htimedas, a
orillas de canales o esteros, suelos arenosos y sitios pantanosos. En Chile se extiende desde la region de Arica
y Parinacota (Chapiquina, 18°12’S) hasta la Region de Aysen (Lago Bertrand, 47°02’S) y desde el nivel de mar
hasta los 4000 m de altitud.
Finotetal., El generoPolypogon en Chile 175
Caracteres distintivos. — Se caracteriza por sus paniculas amplias con espiguillas provistas de glumas con
aristas muy largas (4-6 veces mas largas que las glumas), onduladas. Especie muy cercana a P. interruptus, se
distingue de esta por las aristas de las glumas mas largas (hasta 14 mm en P. australis, hasta 4 mm en P inter-
ruptus), por la lemma con el apice 4-aristulado, las dos aristulas marginales mas largas que las aristulas inter-
nas (las cuatro aristulas de la lemma iguales en P interruptus).
Material adicional estudiado.— ARGENTINA. Ncnquen: Quila-Quina, Ugo Lacar, Sdnchez228 (BA). SahaiChicoana por Cachl, Valle En-
rantjHn Novorfl ct fll 1039 (SI). San Juan: jSchal, Quebrada Gualcamayo, entre Chepical y El Salto, Roig 2187 (BA). CHILE: Joseph 3316
(CONC); Salto, Philippi s.n. (SGO). Region de Arka y Parinacota: Arica, 20 mjaffuel 1109 (CONC); Arica, Chapiquifta, 4000 m, Zoellner
3926 (CONC); Arica, Valle de Azapa, 150 m, Schlegel 4866 (CONC); Parinacota, cerca de Putre, 3300 m, Arroyo 597 «
Tarapaca: Iquique, Corsa, 1300, Chong s.n. (CONC); Iqu
m,Ricardi 3426 (CONC); Iquique, Tarapaca, 1350 m. Fen
& Squeo 10532 (CONC); Antofagasta. Aguada Ojos del f
(CONC); Chiu
2184 (SGO). 1
(CONC);l
1. (CONC). I
00 m, Arando & Squeo 10316 (CONC); .
0 m. Baumann 47 (CONC); El Loa, Calan
2600 m, Villagrdn 2206 (CONC);
bradadeSocaire, 3840 m, Rodriguez deRui;
Loa, Rio San Pedro, San Pedro de Atacama, 11.,.
‘ ■ tn 107(SGO); Tilopozo, Salar de Atacama, 2300 m, Teillter3358 (SGO); T_-_
erillos, 3000 m, Arriagada s.n. (CONC); H
m 127 (SGO); 7
naetaI.303(CONC);Choapa,PangalLosVilos,25ra,Ji[cs2697(CONC);Elqui,
'ray Jorge, Munoz 4036 (SGO); lllapel, Landbeck s.n. (SGO); Limari, Cordillera de Ovalle, rio t
gua 10 km W de Rio Blanco, Canon del rio Aconcagua, 1250 m, Hutchisi
« Quebrada Puerto Frances. 100 m, Ldpez 11603 (CONC); Juan Femdndez. Ree.
inson Crusoe, La Vaqueria, 30 m, Stuessy & Crayeford 6439 (CONC); Isla Robinso
etd5308 (CONC); Isla Robinson Crusoe, Bahia del Padre, 10 m, Stuessy et al 11035 (CONC); Isla Robinson Cer
Robinson Crusoe, Sanjuan Bautista, Ldpez 6- Pertflilillo 11550 (SGO); Masafuera.CJuebrada . Quebrada Casas Cementerio
M. (CONC); Qulllou, Llm.ch.. Ulu Ul». 235 m. 2332. 2812 (CO ^ ^
SZ™OQu,m»,Co™deUC™.20„,^
1542 (CONC)- Quintero, Los Arrayanes, 20 m. Guncfee! s.n. (CONC); Quintero, Los Juanes. 20 m.
enda Rinconada Lo Cerda, Maipu, t^ueora
(S(K)), San Jose de Maipo, Caj6n del Rio N
Quebrada San Alfonso, 2000 m. Levi 1041 (CONC); Ceri
KONC);ElAlfalfal,RloO
m,Guncfeel 45970 (CONC)
i.n. (CONC); Cuesta Barriga, 500 m, Levi 2971
,tiago. 670 m, Aroya 59 (CONC); El Canelo, 900
Gunckel s.n. (CONC); Quebrada de Macul, 850 m. I
2250 m, Araya s.n. (CONC), Quebrada La P
Gabriel, 1500 m, Rojas s.n. (CONC); Vitacu
^nes, 700 m. Pfister s.n. (CONC); Termas de Cauquenes, 700 2(QHC0^^^^^ Negro, Rio Claro, ^-rv .... ...c.,..
ILONQ; Colchagua. Landbeck s.n. (SGO); Colchagua, La Rufma, 77U m, (qq^C)' San Fernando Termas del Flaco, 1700 m, Mon-
m L^i 1040 (CONC). Quebrada de Ramon,
a 740 m Schlegel 3205 (CONC); S.N. YerbaLoca.
omc 237 (CONC). Quebrada El Yeso,
Arroyo & Humana 855 (CONC); San
Colchagua. Undbecks.n. (SGO); Colchagu
C); Quebrada Camino Veeas del Flaco, Hu
Ptimilla Norte Uico, Villagnin & Tapia 111-90 (SGO); Camino Laguna d ^
(CONQ; Cerro Condell, 290 m, Barros 1627 (CONC); Curic6, 200 m, Barros 1
of Texas 7(1)
178
vista wtral. LCariopsis, vista lateral.
fie. 1. Polypogon cadiinalensis Phil. Uiks 483 1). A. Habito. B. Ugula. C Espiguilla. D. Lemma, v
Finot et al., El genero Polypogon en Chile
ceas, de 2,5-5 mm de largo, glabras o algo escabrosas en la cara abaxial, denticuladas en el apice; laminas de
ll-17{-25) cm de largo por 3-5 mm de ancho, planas, glabras o suavemente escabrosas en ambas caras, hordes
escabrosos. Pamculas espiciformes, apice lobulado, amarillentas o verde palido no violaceas, de (6-)9-16 cm
de largo por 1-3 cm de ancho, con ramas laterales de 2-4 cm de largo, contraidas o laxas; raquis glabro o esca-
briusculo. Espiguillas se desprenden con un fragmento de pedicelo de 0,5-2 mm de largo; pedicelos de 0,5-2,3
mm de largo, fuertemente escabrosos; glumas subiguales de 2-2,6 mm de largo por 0,3-0,6 mm de ancho, a
vecesla gluma inferior levemente mas larga que la superior, la quilla provista de escabrosidades de 0,2-0, 5 mm
de largo, el apice agudo, mucronado; lemmas de 1,2-1,7 mm de largo, glabras, hialinas, apice truncado, 4-£
tulado, las 2 aristulas externas mas largas que las internas; arista ausente o presente, recta o levemente gen
lada, de 0,5-l,2 mm de largo, subapical, inserta a 0,3 mm del apice, exserta o no entre las glumas; pale
0,4-0,8 mm de largo, menor que la mitad lemma, hialina; callo glabro; lodiculas de ca. 0,4 mm de largo; anteras
de 0,4-0, 6 mm de largo. Cariopsis de 0, 9-1,5 mm de largo por 0,2-0,4 mm de ancho.
Iconogra/i'a.— Nicora (1978: 400, fig. 266a-e bajo Chaetotropis chilensis); Riigolo (2012a:166 bajo Ch.
chilensis); Riigolo etal. (2008: 18: fig. 3a-c).
Fenologfa.— Florece entre octubre y febrero.
Dislribucte geogiafica > hMtat.-Especie americana, dislribuida en Argentina, Brasil, Chile, Paraguay y
Uruguay (Soreug et al. 2003; Rttgolo et al. 2008). En Chile se extiende desde la Region de Atacama hasta a
region de Los Lagos (Isla grande de Chiloe). Se encuentra tambien en el Archipielago de Juan Fernandez (Isla
Robinson Crusoe), donde fue probablemente introducida en el s. XVII (Skottsberg 1921). Habita principal-
mente zonas costeras o precordilleranas de baja altura, desde el nivel de mar hasta los 700 m de altitud en
suelosarenososyhumedos.Especiemuycomunenpraderas,consideradounforrajeapetecidoaunquedebajo
rendimiento (Rosengurtt et al. 1970). ^ ^ d lar o en el
Caracteres distintivos. — Se caracteriza por la presencia de escabrosidades e , - , mni e a g
dorso de las glumas (Vi del ancho de las glumas a tan largas como el ancho de las glumas); las glumas son agu-
a arista desarrollada; la lemma puede ser miitica o bien presentar una arista
ill
Rnotetal., El genera Polypogone
[. 7376 (SI). CHILE. Region Men
:); Rio Clarillo, 7
Finol & Solis 2288 (CONC-CH). MEXICO. C
CLAVE DE LAS VARIEDADES DE P. ELONGATUS PRESENCES EN CHILE
atus Kunth var. strictus E. Desv., Fl Chil. 6:302. 1854. (Fig. 2). l
(HOLOnpo: SCO 37180 imagen internet!).
Se distingue de la variedad tipica por tener paniculas compactas, de (2,3)3-8 cm de largo, 0,8-2 cm de ancho y
lemma miitica. Las plantas son generalmente pequenas, i
Fenologi'a.— Florece desde diciembre a febrero.
Distribudongeogrdficay hdhitat. — Se encuentra esp
la Region Los Lagos (Isla Grande de Cbiloe), entre 5
lenores de 35 cm de alto.
5urgidero, 30 m, Vinagrdn & A. Letva 7462 (CONC).
Iconografia. — Riigolo & Molina (1977a; 143, fig. 12a-e bajo Agrostis kuntzei).
Distribucidn geogrdjicay habitat.— Argentina y Chile (Soreng et al. 2003). En Chile presenta una distribu-
cion reducida a la Region de Coquimbo. Habita principalmente zonas de alta montana entre los 2000-2800 m
de altitud, generalmenie en suelos humedos.
Observaciones.—Se eligio como lectotipo el especimen NY-327669 que con
un dibujo interpretativo de la espiguilla, lemma, palea, ovario y antera.
s (Phil.)Johow, Est. Fl. Islas dejuan Fernandez 136. 1896. Nowodworskya imberbis Phil., Andes
s (Phil.) Bjorkman, Symb. Bot. Upsal. 17(1):14. 1960. Tipo: Halle esta grama en la isla
to del Ingles, Philippi s.n. (holotipo: SGO-PHIL no visto; isotipo: US-95294 fragm. ex
d; nudos /-J,
Hkrbas perennes, cespitosas, erectas; canas de 17-80 cm de alto; entrenudos de 5-9 cm de largo- 1
Uses, estriados, castaPos o marrones. Valnas de 3,5-9 cm de latgo, mas cortas o tan laigas como los emrena-
dos, glabras, estri^as, llgulas membranaceas, de l,5-3.5 mm de largo, glabras o levememe escabrosas en el
do.so;lamma5de4-10cmdeUrgopor2-6mmdeancho,plana5,glabras.Pamcula5e5plciformesosubespici-
tormes,de5-20mdeUrgoporO,8-2cmdeancho;raquisglabro.Espiguillassedesprendenconunfragmeiito
de pedtcelo de 1, 5-2,5 mm de largo; pedicelos de 1,8-3 mm de largo, escabrosos; glumas de 2-2,8 mm de largo
(excluidas las anstas), la inferior algo mayor que la superior, 0,5-0,8 mm de ancho, escabrosas en el dorso,
apice a^do, terminado en aristas de 0,8-l,7 mm de Urgo; lemma de 1,2-2 mm de largo, mntica, hiallna, apice
trnneado^ con 1m ner«M laterales tetminados en ttes pequenas aristulas de 0, 2-0,4 mm de largo; callo glabro;
Heade0,4-03mmdelargo,,gualaUmitaddelalemmaomenorqneesta,hialina;anterasde0,5-0,6nunde
largo. Unopsis elipsoide, de 0,9-1, 5 mm de largo por 0,25-0,4 mm de ancho
herlas"
Fenohgia. — Florece entre noviembre y junio
tnsnlm™Zntl7t‘^~fr^
ArchipieUgo dejnan Fernandez; en Chile continental se encuentra desde la Regiin
Journal of the Botanical Research Institute of Texas 7(1)
Caracteres distintivos.— Presenta una lemma con una arista recta levemente exserta entre las glumas,
paniculas no espiciformes, ligula glabra y cortas aristas en las glumas.
Obscrvacioncs.— Especie muy cercana a P. australis de la cual se distingue por el largo menor de las aristas
de las glumas; (en P. australis las aristas alcanzan hasta el doble de largo de la espiguilla). Sin embargo, la longi-
tud de las aristas es variable. Los tipos de P. breviaristatus difieren en el largo de las aristas: SGO-71535 tiene
aristas muy cortas mientras los tipos SGO-72936 y SGO-72945 tienen aristas largas (mas largas que las glu-
mas), como en P australis. |
La descripcion original de P. interruptus var. longearistata de Desvaux (1854) hace referenda a las aristas j
de las glumas (1 %-2 % lin. = 3,6-4,86 mm) que superan el largo de estas (1-1 V4 lin. = 2,16-2,7 mm), las que !
no difieren significativamente de las aristas mas conas observadas en P australis (5 mm). La var. longearistata |
fue descriia sobre la base de 4 sintipos: Santiago (Gqy), Valparaiso (Meyen), Rancagua (Bertero 586) y Quillota [
(Bertero 1256). Los tipos no fueron ubicados por lo que no fue posible tipificar y aceptamos la sinonimia estab- l
lecida por Soreng et al. (2003). '
El tipo de Polypogon interruptus var. breviaristatus E. Desv. no fue ubicado; seguimos a Soreng et al. (2003) i
Anuales, canas de 12-35 cm de alio; entrenudosde 4-8 cm de largo, glabros; nudos 2-3. Valnas de 3-7 cm de
largo, mas cortas o tan largas como los emrenudos, glabras; llgulas membranaceas, de (0,5-)2-3,5 mm de lar-
go, glabras, dentieuladas en el apice; laminas de 2,3-7 cm de Urgo, 2-3 mm de ancho, planas, escabrosas en los
nid^enes. Pantculas espiciformes a subespiciformes, de 3-6(-10) cm de largo, 0,5-l,5 cm de ancho; raqi*
gla to o escabroso. Espiguillas caen con un trozo de pedicelo mny reducido, de03-0,6mmde largo; pedicelos
deO,M mm de largo, escabrosos; glumas designales, la tnferiorde3a-3,8 mm de largo, la superior de2,7-3,l
mmdelar^lexcluidas las aristas),subignalesenancho(0,4-0,6mm), escabrosas en la parte inferior del dorse;
J.celobado,tttenosltecuentementealgoagudo,arritado;aristade5-7tnmdelargo,enalgunoscasostnasdel
dobledelargoquelasglunnri,rectas,escabrosas;lemmasdel-l,8mmdelargo,glLs,hlalitias4ptcem^^^^
do, con ^ ™l^, as extemas son levemente mds largas qne las intemas; arista recta o algo ondulada, caedi-
c:i,o«a;e7:pmrr^^^^
fconqgrq/la.— Gay (Hist. Fis.yPollt. Chile, Bot , Atlas Vol 1 t 77 18541
Fcnologm.-Floreceentrelosmesesdeoctnbieylehrero ' ’
liasJ^^dizti^r::;!^::"
costcras no superando los 500 m de altitud, en suelos humedos y
Rnotetal., El genero Polypogon en Chile
187
Caracteres distintivos. — Se distingue por las glumas lineares, algo desiguales, dorsalmente escabrosas en
la mitad inferior.
Observacion.— Polypogon linearis es morfologicamente cercana a P. monspeliensis, de la cual se diferencia
por sus espiguillas mas largas (3-3,8 mm en P linearis, l,4-2,2 mm en P monspeliensis). Tipicamente, en P
linearis la arista de la gluma es apical (en P monspeliensis inserta en la incision del apice bilobado).
Especfmenes adicionales examinados.— CHILE. Region de Coqnimbo; Choapa, Huentelauquen, 60 m Jiles 2841 (CONC). Region de Val-
paraiso: Casablanca, 250 m. Garmenta 5288 (CONC); Quintero, Los Juanes, 20 m, Navas s.n. (CONC); Campiche, 20 m, Gunckel 35725
(CONC); Quillota, Limache, Lliu-Lliu, 255 m, Garaventa 2254 (CONC). Region Metropolitana: Santiago, Chacabuco, Batuco, 480 m,
GuncM 26835 (CONC); Batuco, 480 m, Montero 563 (CONC); Batuco, 480 m, Gunckel 26719 (CONC); Batuco, 480 m. Looser 3631 (CONC);
Batuco, 480 m, Gunckel 26739 (CONC); Penaflor, 450 m, Montero 193 (CONC); Quebrada la plata, 740 m, Schlegel 3198 (CONC). Region de
a Sauzai, 125 m. Bliss & Lusk 808 (CONC). Region del Biobio: Concepcion, Hualpen, 20 m, Banos 159 (CONC); Nuble, entre San Nicolas y
Ninhue, Coipin, Finot 2304 (CONC-CH).
f«mn«,momatosas.de(lI-)25-90 cm de alto; en.renudosde8-17cmde largo; nud.^2-3glabros,Va
cm de largo, mas cortasquelose„trenudos.gkbras;llgulasde2-6mmdelargohrahna^^^
»g«lares,agudascnelapice,li5asoescabrosaseneldorso,decurrentesconlavama,aveceslac.raadas,lam.-
wplaaas o conduplicadas. de 5-15 cm de largo por l-« mm de anebo. glabr^ o escabrosas ^ “
yniegfnes, Panlculas larras o subespiciformes, de 5-16 cmde largo, l-6cmdeancho, vertex do^asopur^
Pteas eon ramas laterales contraidas o laxas; raquis escabroso-himuto. Esprgudte -
'tagmeiiiodepedicelodel-4mmdelargo;pedicelosdel-6ramdelargo,escabroM ns ^ ^
fc-k 3-5,5 mm de largo por0,4-0,8 mm deancho,verdescon ttn.es pnrpnreos,-nerv^^^^^
lisas en el dorso mttttcas o mucronadas, muerdn de 0,l-0,6 mm de largo; lemmas hralmas, de l,5-2,2
mm dp 1 , ’ , . „ p1 ^nire truncado o 3-dentado, 5-nervia, los nemos apenas
«im de largo, lisas o escabrosas en el dorso, con el apice truncaao o ^ del dnrso de
“"mdUos mas alia del apice; arista de 2-6 mm de largo, inserta en la mila o en e ^ escasos a cada
“'™nia,geniculadayre.orcida,escab.osa,exser.a;calbgUbrooc»nd»meAomsdep^se^^^^
>*fc.PebsdeO,2-0,4Lde largo; paleade0,4-0,8mmdetegoorednc.^;lodicnlasde0,4.^,5mn^*lm^^
;8n<lasobibbnladas;anterasdeO,5-0,7mmde largo. Cariopsisebpsoida^
'»^.0.4-0,6mmdeancho; macula embrionalde0,3-0,5mmdela^|h.o^^
fconograJin-Rngolo Sr Molina (1977a; 12f.jbajoAgmstts«gellmncn)
Fenologia. — Florece desde diciembre a febrero.
Distribucidn geogrdficay habitat.— Argentina y Chile (Soreng et al. 2003). En Chile se encuentra desde la
Region del Bio-Bio hasta la region de Magallanes y Antartica Chilena, entre 10 y 1200 m de altitud. Crecen en
suelos principalmente humedos, a veces arenosos.
Caracteres distintivos.— Se caracteriza por sus paniculas laxas amplias y antecios con arista larga (3-5,5
mm), inserta en la zona media de la lemma, geniculada y retorcida.
189
nas de (ll-)35-75 cm de alto; entrenudos de 5-15 cm de largo, glabros, es-
triados; nudos 2-4, glabros. Vainas de 4-11 cm de largo, mas cortas tan largas como los entrenudos, glabras o
escabrosas; ligulas hialinas, opacas, de 1-5 mm de largo, glabras o algo escabrosas en el dorso, denticuladas en
el apice; laminas de (3-)8-14 cm de largo, 2-6 mm de ancho, planas, escabrosas en ambas caras y margenes,
apice agudo. Paniculas espiciformes, amarillentas o verde palido, de 2-8 cm de largo, 1-2,2 cm de ancho;
raquis escabroso. Espiguillas se desprenden con un fragmento del pedicelo de 0,l-0,2 mm de largo; pedicelos
de 0,25-0,6 mm de largo, fuertemente escabrosos; glumas aristadas, subiguales, de l,4-2,2 mm de largo (ex-
cluidas las aristas), 0,4-0,5 mm de ancho, escabrosas en el dorso, con el apice bilobulado; aristas de 2-5 mm de
largo, mas largas que las glumas, rectas u onduladas, escabrosas; lemmas de 0,8-1, 3 mm de largo, glabras, hia-
linas, brillantes, apice truncado, 4-dentado; arista recta, de 0,8-2 mm de largo, inserta en el apice o en el tercio
superior de la lemma (a 0,2-0,3 mm del apice), exserta o no, escabrosa; callo glabro; palea de 0, 7-1,1 mm de
largo, poco menor que la lemma, hialina; anteras de 0,3-0,45 mm de largo. Cariopsis de 0,4-1 mm de largo,
0,25-0,35 mm de ancho, ovalado o ellptico, muy adherido a las glumas; macula embrional de 0,2-0,4 mm de
largo; hilo de 0,2-0,3 mm de largo.
fconogra/fa.— Doell & Hackel (1895: Tab. XII, fig. D; Hitchcock (1951: 364, fig. 510); Nicora (1978: 396, fig.
263a-e); Rugolo et al. (2008: 29, fig. 7a-c).
mmerocromos6mico.—2n = 14, 28 (Tucker 1996).
Fenologj'a.— Florece entre noviembrey febrero.
Distribucidn geogrdfica y hdbitat. — Especie de origen eurasiatico, introducida en America on e se ex
tiende desde Canada hasta Chile y Argentina. Se encuentra en Norteamerica (Canada, Estados Unidos, Mexi-
co), Centroamerica (Costa Rica Guatemala) y Sudamerica (Argentina, Bolivia, Brasil, Chile, Ecuador, Para-
S»y, Pert, Uruguay) (Hitchcock 1927; Soreng et al. 2003; Mgplo et al. 2008). En Chile se encuentra derfe la
ngionde Arica y Parinacota, hasta la region de Los Ugos. desde los 30 m de altitud en las zonas costeras asta
los 3000 n. de altitud en las zonas montanosas. Crece sobre lechos rocosos de embate, ori las de canuno,
tonas ribereflas junto a Cortuderia, Tessaria utemthWdes y Bucckaris. en chacras y orUlas del rl« (Vllupon
*17). en suelo arenoso (Cmckel 30442), suelo salobre (Gunchel 18719) y como maleza en cultivo de arroz (Ro-
^iuez&Baeza2345). . ,.
Caracteres distintivos.—Se caracteriza la panicula espiciforme, densa, con espigui as cortamente pe ice
ladas,lasglumasbilobuladas,provistasdelargasaristasquenacendesdelaincisi6ndelapiceb^
Wa^genemlmenteciliados,
Observacidn.— Especie afi'n a P. maritimus con la cual se puede conrun g
Icvemente bilobadas, los lobulos Ys-Ve del largo de las glumas; en P. mantimus as g u J
^(bslObulosalcanzany.-.Adellargodel.glumriyP«Ua™^^^^^
ausente en R maritimus). Los ejemplares de P. mantimus citaaos p s
OI™79)eorrespondenaP.monspeliensis.porloquesedescartalaprt*naadeP»^l.».nsenChde.Elt.pode
P "tonspeliensis f. nana no fue localizado; en la sinonimia seguimos a Rugolo (201 ).
190
ilNWde San Pedro deAtac
Brando 670 (CONC); Huasco, Freirina, 90 m, 1950, A. Pfister s.n. (CONC); Quebrada de
; Cavilol^n, 515 m, files 4040 (CONC); Illapel, cuesta El Espino, 1200 m,;iles 1615
:f al. 7579 (CONC); Termas de Soco, 100 m, Ricardi 5562 (CONC); Zorrilla, 350ni,Jilo
n, files 1657 (CONC). Region de Valparaiso: San Antonio, entre Mirasol y Algarrobo,
60 m, Schlegel 309 (CONC); Quillota, Tiinel de la Calavera, 320 m, Gunckel 46208
a 2250 (CONC); (Quintero, 20 m, Guncfeel 24386, 28055, 30442 (CONC); Ritoque, 10
i de Casablanca, 400 m. Looser 4419 (CONC); Petorca, Catapilco, 6,2 km Wde
1, Bliss 683, 706 (CONC); Marga-Marga, 150 m,Jaffuel & Piridn 1839 (CONO;
a 3281 (CONC); Archipielago Juan Fernandez, Isla Alejandro Selkirk, Quebrada Vacas, 40 m,
I N de quebrada Casas, 20 m, Stuessy & Ummers 8324-a (CONC pro-parte, mezclado conP.
tisla, Rondanelli & Humana 115J8 (CONC); Isla Robinson Crusoe, Quebrada Puerto Frances,
Ileo, 10 m. Mahu 166 (CONC); El Tabo, quebrada Cordova a 20 k
Looser 3437, 3632, 3634 (CONC); B
3952 (CONC); Marruecos, 430 m
(CONC); El Tagueral, cerca de Ui
San jos6 de Maipo, Cajbn Morale
100 m, Dpez 11603 (CONC); San Ai
1047 (CONC). Region Metropolitan
m, Levi 2210 (CONC); Batuco, 480 m, Montero2118 (CONC); laguna Batuco, 480 m. Looser 2038 (CONC); Batuco, .ucia uc .<o
:o,tranque Ortega, ScWegel3961 (CONC); Batuco, 480 m,Ja//ue!l869(CONC);Colina, 900 m, Schlegel
unchel 28749 (CONC); Polpaico, El Taco, 522 m, Soa 41182 (CONC); Melipilla, 175 m, GunM sx
, 470 m, Araya26 (CONC); Santa Luisa, cerca de Quilicura, 450 m, Matthei & Quezada 603 (CONO;
egas del Ruhillas, 2100 m, Teillier & Mdrquez 5181 (CONC); Santiago, cerro San Cristbbal, 800 m,
esta La Dormida, 1160 m, Guncfeel 29954 (CONC); Lampa, Taqueral 1985 Araya s n (CONC-CH).
L‘i'sr"(coNC)™’
erros frente a la playa, 20 m. Knot &■ Leppe 121 (CONC); Linares, a lo largo del rio Achibueno, desde
„ Ruiz & Lopez 1207, 1218, 1271 (CONC); Taka, predio Venecia, 1 kmalsurdeTalcaencultiw
C). Reeion del Bio-Bio: Concepcidn, 10m, Barros 1833 (CONC); Barrio Universitario,50
. . 4042 (CONC); vegasde
a 79 (CONC); camino entre Bulnes
t,P«rra&Rodrlguez77(CONC),ChilknEstaciqnExperiment-----''‘'”’“'"“‘’‘‘"‘"'’"^^^°''^'“^’®^^^^^^
O Nielol, 160 m, Guncfeel 12581 (CONC); Curarrehue, a
2299 (CONC-CH). R
E,CoipIn,Fino
,de 1,5-13 (
AGRADECIMIENTOS
A Robert J. Soreng (Smithsonian Institution, Washington, DC.), quien gentilmente nos envio material bibli-
ograhco y por la discusion de aspectos nomenclaturales. A Fernando Zuloaga y Norma Deginani (Instituto de
Botanica Darwinion, Buenos Aires), quienes amablemente nos enviaron especimenes para su estudio. A los
curadores de los herbarios BA, CONC y SCO el envio de material de herbario en prestamo. A Glenda Fuentes,
Gladys Munoz y Veronica Fuentealba, la ayuda tecnica. A un corrector anonimo y a Barney Lipscomb, editor de
Journal of Botanical Research Institute of Texas, por sus acertadas sugerencias para mejorar el texto de este
trabajo. Este estudio fue financiado por el proyecto DIUC 210.121.014-1.0.
REFERENCIAS
Baeza, C.M., C. MartkorenaJ. Stuessy, E. Ruiz, & M. Negritto. 2007. Poaceae en el Archipielago de Juan Fernandez (Robinson
Crusoe). Gayana Bot. 64:1 25-1 74.
Barkworth, M.E., K.M. Capels, S. Long, LK. Anderton, and M.B. Piep. 2007. Flora of North America north of Mexico, vol. 24.
Poaceae, part 1 . Oxford University Press, New York.
BjOrkman, S.0. 1 960. Studies in Agrostis and related genera. Symb. Bot. Upsal. 1 7(1 ):1 -1 1 2.
Brongniart, A.T. 1 829. Voyage autour du Monde, entrepris par Ordre du Roi, execute sur les Corvettes de S. M. Uranie et
la Physicienne. Botanique 2:19-21 .
Clayton, W.D and S.A Renvoize. 1 986. Genera graminum. Grasses of the world. Kew Bull. Addit. Ser. 1 3(1 0):1 -389.
Desvaux, E.E. 1854. Grami'neas. En: C. Gay. Historia fisica y politica de Chile, Botanica 6:233-469.
Doell, J.C. & E. Hackel. 1895. Gramineae II. In: C.F.P. von Martius, ed. Flora Brasiliensis. Enumeratio plantarum in Brasilia
hactenus detectarum... Vol. II Part 3. Monachii.
Favret, E.A., N.O. Fuentes, A.M. Molina, and LM. Setten. 2007. Description and interpretation of the bracts epidermis of
Gramineae (Poaceae) with rotated image with maximum average power spectrum (RIMAPS) technique. Micron
39:985-991.
Finot, V.L, C. MARTK:0REN^ AJ. Barrera, M.M. Schick, & MA. Negritto. 2009. Diversidad de la familia Poaceae (Gramineae) en
la Region del Bi'o-Bio, Chile, basada en colecciones de herbario. Gayana Bot. 66:1 34-1 57.
Finot, V.L, J.A. Barrera, C. Martkorena, and G. Rojas. 201 1 a. Systematic diversity of the family Poaceae (Gramineae) in Chile.
In: 0. Grillo and G. Venora, eds.The dynamical processes of biodiversity— Case studies of evolution and spatial distri-
bution. In-Tech Open Access, Croatia. Pp. 71-108.
Finot, V.L, W. Ulloa, CM. Baeza, A. Martkorena, & E. Ruiz. 201 1 b. Micromorfologia de la lemma de los generos Polypogon
Desf., xAgropogon P. Fourn. y Agrostis L. (Poaceae) en Chile. J. Bot. Res. Inst. Texas 5:237-253.
Giraldo-CaNas, D. 2004. El genero Polypogon (Poaceae: Pooideae) en Colombia. Caldasia 26:41 7-422.
Glenn, E.P. 1987. Relationship between c
Plant, Cell Environ. 10:205-212.
Gould, F.W. and R.B Shaw. 1 983. Grass systematics. Ed.Texas A & M University Press, College Station.
Hackel. E. 1 906. 1 79. Polypogon interruptus H.B.K. var. crinitus (Trin.) Hackel, nov. nomen in Stuckert, T. Segunda contribu-
ci6n al conocimiento de las Graminaceas argentinas. Anales Mus. Nac. Buenos Aires 1 3:409-475.
Hitchcock, A.S. 1927. The grasses of Ecuador, Peru, and Bolivia. Contr. U.S. Natl. Herb. 24:291-556.
Hitchcock, A.S. 1931 . Grasses of central Andes. Contr. U.S. Natl. Herb. 24:384-385.
Htchcock, A.S. 1 951 . Manual of the grasses of the United States (2“ ed). U.S.D.A. Burreau of Plant Industry, Miscellaneous
Publication N° 200. Washington, DC.
Holmgren, P.K., N.H. Holmgren, and LC. Barnett. 1990. Index herbariorum. Part I.The herbaria of the world, ed. 8. Regnum
Veg. 120:1-693.
Hooker, J.D. 1846. The botany. The Antarctic vovaae of H.M. discovery ships Erebus and Terror, Botany of Fuegia, tf
i Cervantes, Santiago de Chile. 289
Kabeer, K.A.A. AND VJ. Nair. 2007. Polypogon nilgiricus-a new species of Poaceae from Ir
** * " " ' " " Morphological varii
Science 3:1 18-1 28.
Kunth, CS. 1829-1835.
Bonpland; pr^ed^ d
Finot etal., El genero Polypogon en Chile
193
Kuntze,C.E.O. 1891-1898. Revisio generum plantarum. Pars 3. Leipzig.
L, R.H. AND S. Qiang. 2009. Composition of floating weed seeds in lowland rice fields in China and the effects of irrigation
frequency and previous crops. Weed Res. 49:417-427.
Lu, S. AND S. Phillips. 2006. Polypogon. Flora of China 22:361-363.
MacBride, J.F. 1936. Flora of Peru. Field Mus. Nat. Hist. 13:1-320.
Marhcorena, C. & M. Quezada. 1 985. Catalogo de la flora vascular de Chile. Gayana Bot. 42:5-1 57.
VON Martius, C.F.P. & A.G. Eichler. 1 893. FI. Brasiliensis 2:1 -342.
Morrone, 0., S.S. Aliscioni, & F.O. Zuloaga. 2005. Analisis de la diversidad y distribucion geografica de la familia Poaceae en
la provincia de Jujuy, Argentina. Ann. Missouri Bot. Gard. 92:595-639.
Muller, C. 1985. Zur verbreitung und taxonomie der gattung Polypogon (Poaceae) in Siidamerika. Wiss. Z. Karl-Marx-
Univ. Leipzig, Math.-Naturwiss. R. 34{4):437-449.
Navas, E. 1973. Flora de la cuenca de Santiago de Chile. Universidad de Chile. Vol. 1:1-299.
Ncora, E.G. 1 970. Gramineae. In: A.L. Cabrera, ed. FI. Prov. Buenos Aires 2:1 98-21 1 .
Nkora, E.G. 1978. Gramineae. In: M.N. Correa, ed. FI. Patagonica. 3(i-ix):1-563.
Nicora, E.G. 1 993. Notas sobre gramineas sudamericanas. Hickenia 2:85-92.
Nkora, E.G. and Z. Rogolo. 1 987. Los generos de Gramineas de America Austral: Argentina, Chile, Uruguay y ^reas limitro-
fes de Bolivia, Paraguay y Brasil. Ed. Hemisferio Sur S.A., Buenos Aires.
Nuttall,! 181 8. The genera of North American plants. Vol 1, Philadelphia.
PwuPPt, R.A. 1 859. Linnaea. Plantarum novarum chilensium. Centuriae sextae continuation 30(2):183-21 2.
PwLippi, R.A. 1860. Florula Atacamensis seu enumeriatio plantarum, quas in itinere perdesertum atacamense observavit
ta'lU.' 'Zl, Friedrich Philippi auf der Hocheberie de. Provinzen Antofagasta undTarapaca
gesammelten Pflanzen. Aufgestellt von Dr. R.A. Philippi. Leipzig, (viii). ^ ^ ^ VI de la obra de Gay Anales
Chile 94:5-34.
PiSER, R. 1920. Ueber einige gram
PoHL. R.W. and G. Davidse. 1971. Chromosome numbers of Costa Rican grasses. Brittonia 23:293 324.
j VII: Pooideae, agrostideae, Meliceae, Phala-
n Jardin botaniques de la Ville de Geneve.
iE., E. A.M. Mona., J.M, S«tes, ».o A. Z»«.c 2008. Gramineae VI
rirtzaces n_t=_^ . FH rnnsen/atoire ct J
194
SORENG, RJ., G. Davidse, P.M. Peterson, F.O. Zuloaga, E J. Judziewo, T.S. Filgueiras & O. Morrone. 2012. Catalogue of New Worid
grasses (Poaceae). httpy/www.tropicos.org/(first published 13 Jan 2010, accessed march 2013).
Steudei, E.G. 1854 [1855]. Synopsis plantarum glumacearum. Stuttgart 2 vols.- pars I. Gramineae. Synopsis plantarum
Stuckert, T. 1 904. Contribucion al conocimiento de las Graminaceas argentinas. Anales Mus. Nac. Hist. Nat. Buenos Aires
11:43-161.
Thellung, a. 1 91 2. La flore adventice de Montpellier. Imp. Emile Le Maout, Cherbourg.
Tovar, 0. 1 993. Las Grami'neas (Poaceae) del Peru. Ruizia 1 3:1 -480.
Trinius, C.B. 1 824. De Graminibus unifioris et sesquifloris. Imp. Acad. Imperiale Sci., Petropoli.
Trinius,C.B. 1836. Graminum in America calidiore ab E. Poepping lectorum pugillus primus. Linnaea 10:291-308.
Trinius, C.B. 1840. Genera Graminum, exposuit C.B. Trinius. V. Agrostidea. Mem. Acad. Imp. Sci. Saint-Petersbourg, Ser.6,
Sci. Math., Seconde Pt. Sci. Nat. 6(2):23-134. Reimpresion: Agrostidea. I, Vilfea. Petropoli.
Trinius, C.B. 1841. Gramina Agrostidea. II. Callus rotundus. Mem. Acad. Imp. Sci. Saint-Petersbourg, Ser. 6, Sci. Math.,
Seconde Pt. Sci. Nat. 4(3-4):352.
Tucker, G.C. 1 996. The genera of Pooideae (Gramineae) in the Southeastern United States. Harvard Papers Bot. 9:1 1-90.
Tzvelev, N.N. 1 983. Grasses of the Soviet Union. Amerind Publ. Co. Pvt. Ltd., New Delhi, India.
Watson, L. and M J. Dallwitz. 1 992. Grass genera of the worid. CAB International, Wallingford, UK.
Zuloaga, F.O., 0. Morrone, & M. Belgrano. 2008. Catalog© de las plantas vasculares del Cono Sur (Argentina, Sur de Brasil,
Chile, Paraguay y Uruguay). Pteridophyta, Gymnospermae, Monocotyledoneae. Monoqr. Syst. Bot. Missouri Bot.
Gard. 107(1); i-xcvi, 1-983.
NEW COMBINATIONS, NEW SYNONYMS, AND TYPIFICATIONS
IN PSEUDOGNAPHALIUM (ASTERACEAE: GNAPHALIEAE)
FROM SOUTH AMERICA
Claudia Monti
Nestor D. Bayon
Departamento de Ciencias Bioldgicas
Facultad de Ciencias Agrarias y Forestales
Universidad Nacional de La Plata
Avda.60entre116yl18
1900 La Plata, ARGENTINA
Area de Botdnica
Departamento de Ciencias Bioldgicas
Facultad de Ciencias Agrarias y Forestales
Universidad Nacional de La Plata
Avda.60entrell6yll8
1900 La Plata, ARGENTINA
Daniel A. Giuliano
Susana E. Freire
Facultad de Ciencias Agrarias y Forestales
Universidad Nacional de La Plata
Avda.60entren6yl18
1900 La Plata, ARGENTINA
a de Botdnica, Departamento de Ciencias Bioldgicas
Facultad de Ciencias Agrarias y Forestales
Universidad Nacional de La Plata
Avda.dOentre 116yll8
1 900 La Plata, ARGENTINA and
»«th American species of Pxudognaphatium Kirp. have been typically treated as Gnaphaliunt L. (Cabrera
>^3a97U, 74 1978- Dillon&Sa^tegui-Alval991a,b; Freire 1998, Bayon
>'« Achyrocbncid cudweeds groupCDrury 1970). However.studiesofdetatledtnorphologt^^^^^^
Deble & Marchiori 2006; Hind 2011; Freire et al. 2011, for South American species).
We found four combinations necessary for a consistent treatment of the genus during the course of the
revision of Pseudognaphalium (Monti, in prep.) and the treatment of Pseudognaphalium Kirp. in Chile and for
the Flora of Argentina (Freire, Bayon, & Monti, in prep.).
MATERIALS AND METHODS
lowing herbaria (abbrevi
The study is based on herbarium material froi
2011): CONC, CORD, LP, MCNS, MCNS, SCO, a
staining with 2% safranin. Characteristics of phyllaries were observed and recorded in cleared samples using
Dizeo de Strittmatter’s technique (1973). Observations were carried out using a light microscope (Gemalux),
equipped with a photographic camera PAL CCD. Scanning electron microscopy (SEM) imaging was carried
out on achene materials that were removed from herbarium sheets, rehydrated in water and fixed in FAA (70%
alcohol, formaldehyde and glacial acetic acid, 85 : 10 : 5), then dehydrated through an alcohol series of
90%:96%:100% and critical point dried using EMITECH K850. They were mounted on aluminum stubs with
double adhesive tape and coated with gold. Achenes surfaces were photographed using a FEI Quanta 200 scan-
ning electronic microscope at 20 kV.
TAXONOMY
s (J. Remy) C. Monti, N. Bayon & S.E. Freire, o
;o. Feb 1839, Gctys.
Observation 1
Gnaphalium phaeolepis Phil, and G. aldunateoides J. Remy are small plants from 5 to 10 cm
branched from base, and oblong to linear leaves. According to the protologue, Gnaphalium p
from G. aldunateoides by its brown tipped phyllaries (vs. whitish tipped in G. aldunateoides). ;
the type materials, we considered this character to be within the range of variation for G aldun
quently, Gnaphalium phaeolepis Phil, is now considered to be a synonym of G. aldunateoides
e name. The specimen
“Frumau 63, year 1^5’' kept at SGO(SGO 64452), which is in accordance with the protologue and has Phihp-
pis annotations, is here selected as the lectotype.
Observation 3
Eve„«A„dej^rg(1991)Uf.C„.pM«™aUu™,eaid«i„G»aph*^^
tams.„ce„hasdwKled.,ereon.e(Fig.lA)as,hisis.heprtaaryway,„differe„a^^^
la (Phil.) C. Monti, N. Bayon & S.E. Freire, comb, i
Condes, L. Nmarro s.n. (lectotype, designated here: SGO 64481 !).
Observation
Philippi (1895: 17) mentioned two syntypes in the protologue of G h I'd l S
ISSZ, SCO 64383. we s.lec.edas,ec.o.yp. SGO
the description in the protologue.
I (Griseb.) C. Monti, N. Bayon & S.E. Freire, comb.
197
The name Gnaphalium glanduliferum has an earlier author, i.e. G. glanduliferurr
T173. 1845, which is a synonym of Helichrysum glanduliferum Sch.Bip. ex Boi
ti (Phil.) C. Monti, N. Bayon & S.E Fr*
Jan 1837, Gay 739 (holotype. 5
G^halmm dimmutivum Phil, and G. perpustllum Phil. ^liffers from G. perpusillum by its Ic
^ lm«r leaves. According to .he prolologue. examining the type materials, we conside,
Phil. IS now considered to be a synonym of G. perpusdlum.
Monti etal., New combinations and t
A
C
B
D
etal. 700. SI). B. P. andkola {Burkart 9491. SI). C P glandulifenim {Hunziker
^9195. CORD). O.Pperpusillum {Teillier&Mdrquez5316. CONC). Scale bar A-0 = 0.1 m
Observation!
Evenifnoherbarii
i„„edfor,he.ypes,.dn«nintl«protologueofG„aph.Kumpe^^
“^ng to Stafleu and Cowan (1979), Philippi’s types are found in heibanum SCO.
turn “Cerro del Volcan ad originem
■Mippi (1864: 167) mentioned in the proiologue of CnapMiumdimi
Tinguiguiric.-, Because there are two sheets kept at SGO for type coUe«.on of G dnnn
^ the specimen SGO 64454 as lectotype, which presents the most complete pla ,
discussion
'"huc.asslfica.orys.udyofthegenusCnuphnli»mascnr.entlydefined,Dr^^
'fetnents: achyrocUnoid and anaphalioid cudweeds.
The former was characterized by having monochromous phyllaries with divided stereomes, yellowish
norets. pappus bristles subclavate at the tip with bases cohering by patent cilia, papillate achenes and capitula
nreTutrAfZn^e^Zs transferred
in his worldwide revision of Gnaphlliese, also recognizSIfe
70 sp^ies to II. Morphological comparison between Gnaphalium and Pseudogmphalmm is shown in Table 1.
In ihB paper, four species are transferred from Gnaphalium to Psendognaphalium owing to monochro-
mous phylanes with divided stereomes (Fig. 1); monomorphic (or slightly dimorphic) pappus briste
^ I ^ ?' of capitula arranged in corymbs; achenial epidermis glabrous and papillate (F
c -nduh/erum Fig. 3); or oblong duplex myxogenic hairs (P. aldmateoides, P. perpusillum. Fig-
considered to be diagnostic features for Pseudognaphalium (Anderberg 1991). However, the
(p aidunateoides, P. perpusillum), and pappus bristles smooth at
1991).
corollas with red purple teeth of n
the base of P glanduliferum (Fig. 2),
We thank Randall Bayer and an anonymous reviewer for useful comments on a first draft of the manuscript.
We are indebted to the curators of the Museo Nacional de Historia Natural de Santiago, Chile (SCO); Universi-
dad de Concepcion, Chile (CONC); Institute de Botanica Darwinion, San Isidro, Argentina (SI); Museo Bo-
tanico de Cordoba, Argentina (CORD); Museo de La Plata, Argentina (LP); and Museo de Ciencias Naturales
de Salta, Argentina (MCNS) for the loan of specimens. The authors are grateful to Maria Alejandra Migoya
(LPAG) for inking the illustration of pappus bristles and preparing the figures. Financial support was provided
by Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET), Argentina and Programa de Incen-
tivos, Decreto 2427/93, Secretarla de Polltica Universitaria del Ministerio de Educacion de la Nacion,
Argentina.
REFERENCES
Anoerberg, A.A. 1991. Taxonomy and phytogeny of the tribe Gnaphalieae (Asteraceae). Opera Bot. 1 04:1 -1 95.
Ballard, H.E., Jr., D.S. Feller, and G.L. Nesom. 2004. Cliff Cudweed at specific rank in Pseudognaphalium (Asteraceae:
Gnaphalieae). Sida 21:777-780.
Bayer, RJ., 1. Breitwieser, J. Ward, and C. Puttock [2006] 2007. Tribe-Gnaphalieae. In: The families and genera of vascular
plants, flowering plants - Eudicots: Asterales, vol. 8, eds J. W. Kadereit and C. Jeffrey (K. Kubitzki, ser. ed.). Berlin
Springer-Verlag. Pp. 246-283.
BavOn, N.D. 2009. Compositae: Gnaphalium. In: Flora Chaquena-Argentina (Formosa, Chaco y Santiago del Estero), S.E.
Freire and A.M. Molina, eds. Colecc. Cient. INTA 23:559-561 . Buenos A
Cabrera, A.L. 1963. Compositae: Gnaphalium. In: Flora de la Provincia d(
Buenos Aires.
Cabrera, A.L 1971. Compositae: Gnaphalium. In: Flora Patagonica, M.N. Correa, ed. Colecc. Cient. INTA 7:109-1 17. Bue-
nn<: Airac
a de Entre Rios (Argentina). A. Burkart, ed. Colecc. Cient.
,. In Flora de la Provincia de Jujuy, Republica Argentina, Colecc. Cient.
10-275-288. Buenos Aires.
Cerana, M.M. AND L. Awza Espinar. 2008. Familia Asteraceae: Tribu Inuleae {Achyrocime.
FI. Fanerog. Argentina Central 4:5-14, 26-52.
Deble.L.P.andJ.N.C. Marchiori.
9:13-16.
DiuxJn, M.O. AND A. SagAstegui-Alva.
Arnaldoa 1:5-91.
D^lon, M.O. AND A. SagAstegui-Alva. 1991b. Gnaphalium.
V. Fieldiana, Bot. n.s. 26 (1422):32-41
DE Strittmatter, C. 1 973. Ni
'^'w,D.G.1970.
present in New Zealand
Flora of Peru, Family Asteraceae: P
15:126-129.
. nn.. IV Inuleae in- A T Hunziker, ed. Flora Fanerogamica Argentina 1 4:3-60.
S.E., LP. Deble, AND L. Iharlegui 201 1 . Compostas: 5. tribo. Inuleae. in. riora
A. Reis, ed. Santa Catarina, HBR. Pp. 1 067-1 197. r n.nhaliinae Government Printer. Pretoria, South
O.M. 1983. Flora of Southern Africa, Part 7 Inuleae, Fasc. 2 Gnaphalimae. Gove
Africa. Ed. Balogh Scientific Books. 325 pp. Gnaohaliinae. Bot. J. Linn. Soc. 82:1 81 -232.
O.M. AND B.L. Burtt. 1 981 . Some generic concepts in Compositae Gnapha
DJ.N. 201 1. An annotated preliminary checklist of the Compositae of
.. 1960. Las especies de plantas d
Journal of the Botanical Research Institute of Texas 7(1)
Nesom, G.L 2001 . New records in Pseudognaphalium (Gnaphalieae: Asteraceae) for the United States. Sida 1 9:1 1 85-1 19a
Nesom, G.L 2004. Pseudognaphalium canescens (Asteraceae: Gnaphalieae) and putative relatives in western North
America. Sida 21:781 -790.
Nesom, G.L 2006. Pseudognaphalium. In: Flora of North America Editorial Committee, eds. Flora of North America north
of Mexico. Oxford University Press, New York and Oxford. 1 9:41 5.
Stafleu, fa. and R.S. Cowan. 1979. Taxonomic Literature, Vol. 2: H-Le. 2nd ed. Regnum Veg. 98.
Thiers, B. 201 1. Index Herbariorum: A global directory of public herbaria and associated staf. New York Botanical
Garden 's Virtual Herbarium. htp://sweetgum.nybg.org/ih/: 29.06.2012.
Ward, J., RJ. Bayer, I. BRErrwiESER, R. Smissen, M. Gaibany-Casals, and M. Unwin. 2009. Gnaphalieae. In: V.A. Funk, A. Susanna, TT.
Stuessy, and RJ. Bayer, eds. Systematics, evolution, and biogeography of Compositae. Vienna, Austria: International
Association for Plant Taxonomy (lAPT). Pp. 539-588.
DIFERENCIACION DE ESPECIES ARGENTINAS DE CLINOPODIUM
(LAMIACEAE: NEPETOIDEAE) A TRAVES DE CARACTERES MORFOLOGICOS
Y ANATOMICOS DE SUS FRUTOS
Melina Scandaliaris
BecariaSECyT
Cdtedra de Botdnica Taxonomica
Facultad de Ciencias Agropecuarias
Universidad Nacional de Cordoba
/. Valparaiso s/n, CC 509, Cordoba, ARGENTINA
mscan@agro.unc.edu.ar
Facultad de Ciencias Quimicas
Universidad Nacional de Cdrdoba
Cdrdoba, ARGENTINA
gbarboza@imbiv.unc.edu.ar
Clinopodium L. (tribu Mentheae, subtribu Menthinae) consta de ca. 100 especies circunscntas en su mayona a
America (desde Canada hasta Chile y Argentina) y a Eurasia templada. En la modema concepcion de este ge-
nero, se incluyen a las especies de Satureja L. del Nuevo Mundo, quedando este ultimo restringido a Eurasia y
Africa (Harley & Granda Paucar 2000; Harley et al. 2004). Acorde a Brauchler et al. (2010) tal circunscnpcion
hace de Clinopodium el genero taxonomicamente mas dificil y morfoldgicamente m^ diverse dentro de la sub
tribu Menthinae. Estos autores realizan la filogenia de esta subtribu, analizando 48 especies de Clinopodium y
demostrando que las especies de este genero no forman un grupo monofiletico; por lo tanto, su circunscnpcion
actual resultamuypoco natural. , , , .
En Argentina, habitan cinco especies nativas de Clinopodium: C. bolivianum (Benth.) Kuntze subsp. tan-
jense (Weddell ex Griseb.) J.R.I. Wood, C. gilliesii (Sw.) Kuntze y C odorum (Griseb.) Harley de la zona serrana
del centro y noroeste del pais, C. brownei (Sw.) Kuntze del Utoral y, C. darwinti (Benth.) Kuntze de la estepa
Patagonica (Harley 2008). , . ...
Diversos autores destacan la importancia taxonomica y filogenetica del pencarpo en Lamiaceae a distin-
204
Journal of the Botanical Research Institute of Texas 7(1)
tos nivelesjerarquicos, junto a otros caracteres floraks, palinologicos y seminales (Cantino et al. 1992; Ryding
1994, 1995, 2010; Moon et al. 2009). Segtin Ryding (1994), algunos caracteres de los frutos, particularmente
aquellos del esclerenquima, son algo conservativos a niveles supragenericos pero son consistentes dentro del
Harley et al. (2004) describen de manera general la exomorfologia de los frutos de Clinopodium en tanto
que otros autores ban estudiado la anatomi'a de un total de 8 especies, incluyendo a C. bolivianum (Wojcie-
chowska 1966; Ryding 1995, 2010; Duletic-Lausevic & Marin 1999; Martin Mosquero et al. 2004). For su
parte. Moon et al. (2009) proponen un analisis filogenetico de las Mentheae utilizando caracteres de los frutos
observados con microscopia electronica de barrido; en dicho estudio, C. vulgare L. y C. nepeta (L.) Kuntze son
los representantes de Clinopodium. Dates exomorfologicos de C. vulgare fueron tambien reportados por Wojcie-
chowska (1966) y Husain et al. (1990).
La mixocarpia, fenomeno debido a la presencia de celulas mucilaginosas en el epicarpo, es otro caracter
relevante en la subfamilia Nepetoideae, habiendose reportado que casi el 70% de las especies poseen frutos
mucilaginosos (Ryding 1994). Segtin Ryding (2001), se pueden presentar tres situaciones distintas: generos
exclusivamente no mixocarpicos, como ocurre en Alvesia Welw., Holostylon Robyns & Lebrun (Ryding 1992)
y Lepechinia Willd. (Scandaliaris & Barboza 2011); generos exclusivamente mixocarpicos como por ejemplo
Lavandula L. (Ryding 1992) y, generos con especies mixocarpicas y no mixocarpicas, caso de Hyptis Jacq. y
Ocimum L. (Ryding 1992). Cuando existe mixocarpia, las celulas que producen mucilago se pueden hinchar
sin liberar el mucilago o bien este queda totalmente expuesto. En Clinopodium, la mixocarpia no fue menciona-
da para las especies estudiadas por Ryding (2010) mientras que Duletic-Lausevic & Marin (1999) y Moon etal
(2009) comprueban la ausencia de este fenomeno en C. vulgare y C. nepeta.
Con respecto a las especies argentinas, la informacion sobre los caracteres morfologicos y anatomicos de
los frutos es mas que fragmentaria (Wood 2011). En cambio, cuatro de estas especies (C. odorum no incluida)
ban sido incorporadas en el analisis molecular realizado por Braucbler et al. (2010), apareciendo en dados
distintos y emparentadas con especies sudamericanas de otros generos.
En base a estos antecedentes y como parte de un trabajo integral sobre la carpologia de las especies de
Lamiaceae nativas de Argentina, los objetivos de este trabajo fueron: a) describir las especies de Clinopodium
nativas de Argentina a traves de los caracteres exomorfologicos y anatomicos de sus frutos; b) correlacionar la
estructura del pericarpo con el fenomeno de mixocarpia; c) valorar la incidencia de los caracteres carpologicos
para diferenciar las especies.
Material vegetal
Se trabajo con material de berbario (ACOR, CORD, CTES, LIL y SI) y
po. Frutos de estos ultimos se fijaron en solucion FAA (solucion de formol a
agua, en proporcion de 2:10:1.7) por 48 hs, y luego se guanlaron en alcohol enlico al 70% ha^
Los matenales berbonzados se depositaron en el berbario ACOR y CORD.
^ ck,j»
Lorenzo, junto al cauce del airoyo, 30 Ago I
1979, Schimni 19313 (CTES).
205
quinto puente colgante. 9 Feb 2010, Proyecto Coleccidn de Semillos (CS) 1417 (ACOR); Sierra Grande (falda E), entre El Cdndor y Copina, 5
Mar 1992, Zygodlo 146 (CORD).
nicroscopio electronico de barrido (MEB) para lo c
con oro y se observaron en un microscopio JEOL JSM 35 CF, en el Laboratorio de Microscopi'a y Microanalisis
de laUniversidad Nacional de San Luis (Argentina). Ademas, se efectuaron extendidos del epicarpo que se
colorearon con safranina para observar el ordenamiento de las celulas en vista superficial.
Los aspectos analizados fueron: 1) tamano, 2) forma en vista dorsal y ventral, 3) simetria, 4) color, 5) or-
denamiento de las cdulas en vista superficial, 6) ornamentacion de la superficie al MEB, 7) ubicacion y forma
delhilo carpico, 8) presencia y tipo de ceras.
Para estimar el tamano de los frutos, se midio el ancho y el largo en 3 muestras de 15 frutos por cada una
y por especie en tanto que para la forma se tomo como referenda a Steam (1983). El color del pericarpo se de-
terraino por la comparacion con la carta de colores de Munsell (2000). La descripcion de la superficie se realize
siguiendo los patrones establecidos por Budanstev & Lobova (1997) y Bonzani et al. (2011).
Para verificar la mixocarpia, se remojaron frutos en agua destilada por 72 boras. Posteriormente, la presencia
demucilagos se puso en evidencia utilizando azul brillante de cresilo al 1%. Se midio el ancho del halo produ-
cido y se tipified la reaccidn segtin Bonzani et al. (2011).
Se realizaron preparados permanentes con los frutos fijados en FAA y con los provenientes de herbario. Se
siguio la tecnica de Johansen (1940) y Gonzalez & Cristobal (1997). Los frutos extraidos de ejemplares de
herbario fueron previamenteremojados durante 10diasenalcoholetilico96%,glicerolyaguaen partes iguales
(Budanstev & Lobova 1997). Se efectuaron cortes transversales de 10-12 pm, con microtomo de acciona-
croscopio optico Axiophot Zeiss, con camara digital incluida.
Para las descripciones anatdmicas, se siguio el criterio de Wojciechowska (1966) quien considera a las
^lereidas como parte del mesocarpo.
^unopodiumbolivianumsubsp.tarijense ^ ^ ^
^orfologia (Figs. 1 A-F; 3 A).-Nuculas de 2-2,4 x 0,45-0,5 mm. Forma obclavado-subulada curvandose
Iwtael Spice, de base obtusa Asimetricas. De cotor manbn amarillemo oscuro. Epicarpo con grandes celulas
WcUaginosas rodeadas por pequenas celulas no mucilagiuosas (Fig. 3 A), Superficie al MEB espiuulosa eu la
"liUd superior, cou "espinas" (rricomas) estriado-acordouados. y superficie reticulado-areolado lisa a leve-
®ente estriada en la mitad inferior. Hilo carpico alargado y ubicado en la base hacia a cara ventra , zona pen-
hilar con ceras de tipo cristaloideesferico. _ , . .•
Mixocarpia (Fig. 2 G).— Reaccidn moderadamente fuerte, con un halo de muci ago con m
'^ Anntomta (Fig. 3 B, C).-Pericarpo de 25-32,8 pm de grosor. Epicarpo de 7,8-^
de celulas mudlaginosas mas o menos oblongas entremezcladas con cdulas no mucilagmosas isodiame-
^^asarectangulares;haciaelapicedeIfruto,cdulasnomucilaginosas^^
®ente intercaladas con tricomas bi- tricelulares no mucilaginosos, ^
^^rpo de 14,1-18 7 pm formado por tres regiones: la mas externa, de 1,6-3,1 pm, con una o dos capas de
aplanadas (alargLas tangencialmente) de Hmites difusos e intensamente ^
‘>«istrata.de4,7-6,2pm,concdulasparenquimaticasisodiamdricasincoloras;laregi^
Journal of the Botanical Research Institute of Texas 7(1)
unistrata, de 7,8-9,4 pm, con esclereidas alargadas radialmente o mas o menos isodiametricas, de paredes en-
grosadas y lumen redondeado pequeno. Endocarpo unistrato, de 3,1-4, 7 pm, con cdulas oblongas y paredes
con engrosamientos escalariformes.
Exomorfologla (Fig. 1 G-l).-NucuUs de 1-1,2 x 0,7-O,8 mm. Fornia elipsoidea, de base y apice redondeados,
de seccion mas o menos trigona. Levemente asimetricas. De color marriSn amarillento oscuro a marron rqjizo
207
oscuro. Epicarpo ^ „
ficie al MEB de tipo reticulado redondeado-radiado,
iibicado en la base hacia la cara ventral; zona perihil
Mixocarpia (Fig. 2 F). — Reaccion moderadamei
Testrla^ cuticulares radiales. Hilo carpico triangular y
r con ceras de tipo cristaloide esferico.
te fuerte, con un halo de mudlago continuo de 0,23-0,25
«lulas mucilaginosas en general alargadas tangencialmente e pxtpma de
*.^.sr^iaL«eTaLasaus.nLMesocanx.ae74,8-843p„.,c^^^^^^^^
6.i-7,8 pm, con una o dos capas de celulas aplanadas de limites difusos e i ^ ^
^ Wstiata, ac 3,1-4, 7 pm, con cClulas parenquimaticas ““ eng'rosaL y lumen reaonaeaao
la, de 63,5-71,8 pm, con esclereiaasalargaaasraaialmenleae pare , ®
P«l«no. Endocarpo unislralo, de ca. 1,6 pm, de celulas oblongas con paredes gruesas.
les, de base truncada y apice redondea-
ro. Epicarpo con grandes cdulas muci-
astmetricas. ue color marrun .»ia. ^ ^e tipo reticulado redondeado-
‘"^rodeadasporpequeflas cClulasno ^^eado en la base hacia la cara ventral;
“<>0 y estnas cuticulares concentricas. Hilo carpico triangular, ubica
‘aperihilar con ceras de tipo cristaloide esferico.
•"vrjoiogta
Levemenu
f}. — [Nucuias uc
s. De color marrbn amarillento a
208
Mixocarpia (Fig. 2 E). — Reaccion moderadamente fi
pos de celulas: las mucilaginosas, alargadas y mas o
mas pequenas e intensamente tenidas de rojo. Tricomas ausei
nes: la externa, de 4,7-7, 8 pm, c(
rojo; la interna, de 17,2-23,4 pm, con esclereidas en empalizada, alargadas radial
y lumen ledondeado. Endocaipo unistralo, de 3, 1-6,2 pm, con celnUs isodiamel,
des con engrosamientos escalariformes.
un halo de mucilago continue de 0,12-0,25
de 4,7-7, 8 pm, formado por dos ti-
s, entremezcladas con las no mucilaginosas,
Mesocarpo de 21,9-31,2 pm, con dos regie-
arectangularesdepaxe-
Sandaliaris & Barboza, Argentinean Clinopodium
Exomorfologia (Fig. 4 A-E). — Niiculas de 1,1-1, 3 x 0,5-0,6 mm. Forma obovoidea, apice
ybase turbinada (en forma de cono invertido), de seccion trigona. Asim^r
ron oscuro. Epicarpo con cdulas
conleves estriaciones concentric;
carpico eliptico ubicado en la base hacia la c
:eras de lipo cristaloide esferico.
Anatomia.— Pericarpo de 27,4-37,5 pm de grosor. Epicarpo de 7,8-9,4 pm, con una capa de celulas alarga-
das tangencialmente intercaladas con tricomas unicelulares disperses. Mesocarpo de 16,5-25 pm, con 3 regio-
nes unistratas cada una: la mas externa, de cdulas oblongas con paredes redondeadas; la media, de celulas
rectangulares mas aplanadas e incoloras; y la mas interna de esclereidas alargadas radialmente, de ca. 7,8 pm,
conparedes engrosadas y lumen redondeado. Endocarpo unistrato, de ca. 3,1 pm, con celulas rectangulares de
paredes con engrosamientos escalariformes.
Exmoifologia (Fig. 4 F-I).— Niiculas de 1,4-1,5 x 0,5-0,7 mm. Forma obclavado-subulada, curvandose hacia
el apice y de base turbinada. Asimetricas. De color marron oscuro. Epicarpo con celulas y tricomas unicelula-
res no mucilaginosos. Superficie al MEB espinulosa, las “espinas” (tricomas) estriado-acordonadas. Hilo carpi-
co alargado y ubicado en la base hacia la cara ventral; zona perihilar, con ceras de tipo cristaloide esferico.
0 produce mucilago.
Anatomia (Fig. 3 D).— Pericarpo de 59,2-79,9 pm de grosor. Epicarpo de 14 ^19 p
bias oblongas a redondeadas y tricomas unicelulares. Mesocarpo de 42,1-56,2 p
tema,de 23,4-31,2 pm, formadapordoscapasde celulas parenquimaticas mas c , , .
redes delgadas; la media unistrata, de 3,l-4,7 pm, con cdulas mas o menos rectangulares e incoloras; la mas
btema, de 15,6-20,3 pm, con esclereidas alargadas radialmente, de paredes engrosadas y lumen redondeado
a estrelbdo. Endocarpo unistrato, de 3,l-4,7 pm, con celulas parenquimaticas, aplanadas tangencialmente, de
En la Tabla 1 , se muestran los caracteres exomorfologicos diferenciales de los frutos, la reaccion uc’ g
nosa y b variacion en grosor de las capas del pericarpo, para las especies estudiadas.
Porotros tales como el tamano h omamentacion de la superficie y la forma del hrlo carpreo. Err cambro,
«i6n especlfica (Marrin Mosquero et al. 2004); asl, en general el apree es rerton
“*»• nrbsp. rarijense y C. odorum por presentar el apice alargado a muy alarga
“P'opia para cada especie salvo en C. odomm y C. giiltesii que companen la ct
S'gitalaclasificaciondeMoonelal. (2009). solo C.Wivimmn'5«‘»P “n
^ apice incluido), mientras que en el resto de las especres son ^
P^C.v,jg„„yC.nepeta(Husainelal.l990;Mar.lnMosqueroeral.2004,Moonei .
210
Journal of the Botanical Research Institute of Texas 7(1)
. . ^ ''"^a media eiirferior;I.Vistazona hilar. F-IC.«Joram.F.VistacaradotMl;6.V^
na hilar. Con fledia se muestra el hilo carpico. (A-E de CS 1418. F-l de CS 7417).
La presencia o ausencia de
posicion en los frutos y el numero de celulas aportan caracteres
espedfico. La variadon va desde fnitos glabros (C darwinii y C. brownei), apicalmente
bsp. tanjensey C gilliesii) a I'ntegramente cubiertos por tricomas (C. odorum). En
ionpluricelulares en C. boliviamm subsp. tarijense y unicelulares en las oirasdos;
1 los descritos por Budanstev & Lobo-
va (1997) para algunos generos de Nepetoideae y complementan a los mencionados por Moon et al. (2009) para
la tribu Menlheae. Asi, C. vuigare y C. nepeta, que poseen superficie reticulada con estructura secundaria es-
triada (Moon et al. 2009), comparten este caracter con C. bolivianum subsp. tarijense, C. brownei y C. darwinii.
Estas tres especies se distinguen a su vez por la forma de la superficie reticulada y la estructura secundaria (cfr.
Tabla 1), modelos que ya habian sido descritos para especies de Mentha (Bonzani et al, 2011). For su parte, C
gilliesii presenta superficie tuberculada y C. odorum espinulosa dada por la cobertura total de los tricomas.
El hilo carpico, ubicado en la base hacia la cara ventral, varia en su forma desde alargado (C. bolivianum
subsp. tarijense y C. odorum), eliptico (C. gilliesii) a triangular (C. brownei y C. darwinii). En cambio, la region
nado por Martin Mosquero et al. (2004) asi como a los de algunas especies de otros generos de Nepetoideae:
Mentha L. (Bonzani et al. 2011), Hedeoma Pers., Minthostachys (Benth.) Spach, Lepechinia Willd. (Scandaliaris
& Barboza 2011, 2012), Hyptis Jacq. (obs. pers.).
Mixocarpia
Segiin la literatura, los frutos de las especies de Clinopodium hasta ahora estudiadas no producen mucilagos
(Duletic-Lausevic & Marin 1999; Moon et al. 2009). Pudimos comprobar que este fenomeno esta presente en
el genero pero no es constante para todas las especies nativas de Argentina (cfr. Tabla 1). Este caracter, que re-
sulta novedoso para Clinopodium, se comporta de acuerdo a lo reportado para otros generos (Harley 1985; Ryd-
ing 1992, 2001), en lo que concieme a su presencia/ausencia y al tipo de reaccion producida; en este ultimo
caso, las especies mixocarpicas analizadas presentan mucilago expuesto con reaccion moderadamente fuerte.
Ryding (1992), para la tribu Ocimeae (Nepetoideae), concluye que tendencias evolutivas hacia la reduc-
cion de la mixocarpia se han observado en especies que crecen en ambientes humedos, o que tienen fmtos
grandes o en aquellas que poseen frutos con glandulas o con pelos (salvo que estos solo esten presentes en el
apice). Si relacionamos estas afirmaciones con las especies argentinas de Clinopodium (tribu Mentheae, Nepe-
toideae), se observaron distintas situaciones: en las especies de zonas hiimedas, C. browneii (litoral argentine)
y C. bolivianum subsp. tarijense (selva tucumano-boliviana), ambas con mixocarpia moderadamente fuerte, no
responden a lo supuesto por Ryding (1992). En cuanto a las especies de zonas semiaridas, Clinopodium gilliesii
(Argentina hasta Peru), no mixocarpica, se contrapone a lo esperado por este autor ya que segiin el habitat y la
presencia apical de pelos en el fruto tendrla que ser mixocarpica. En cambio C odorum (zona semiarida del
centra y none de Argentina) y C. darwinii (estepa patagonica), se ajustan a las afirmaciones de Ryding (1992)
siendo esta ultima una especie mixocarpica tipica de ese ambiente en tanto que la primera es no mixocarpica,
presentando sus frutos cubiertos completamente por pelos, los que, segiin Ryding (1992, 2001) cumplenla
funcion de anclaje reemplazando al mucilago.
A pesar de que los caracteres histoldgicos del pericarpo han demostrado ser de ii
a mvel de familia como en las categorias inferiores (Cantino etal. 1992; Ryding 1994, 1995), para ClinopodUm
la informacion sobre la histologia de los frutos es por demas escasa. Solo Martin Mosquero et al. (2004) han
desenpto en detalle la carpologia de C. vuigare subsp. arundanum (Boiss.) Nyman y C. nepeta (L.) Kuntze (sub
Wojciechowska (1966), Ryding (1995, 2010) y Dnleuc-
• ^'^"^ ,*"°^^''^"°""^"’'^^‘^‘'"™y''’™""t^‘:'^antitativossobrecadaregi6ndel pericarp
otras 6 especies, entre las que seincluye a C.b-’ - - ^ ^
evaluadon sobre su incidencia taxonomica.
Epicarpo.— La estructura del pericarpo de las especies a
dos patrones anatomicos en el epicarpo que se ;
mixocarpia, el epicarpo unistrato consta de cdulas n
213
tianscorte, que se entremezclan con otras no mucilaginosas alargadas radialmente, como ocurre en C. bolivi-
anum subsp. tarijense (cfr. Fig. 3 B), C. browneii y C. darwinii.
Al igual que en otros generos con especies mixocarpicas de las Menthinae, p.e. Conradina A. Gray,
Dicerandra Benth., Hedeoma Pers., Pycnanthemum Michx. y Rhabdocaulon Epling (Ryding 1995), las especies de
Clmopodium coinciden con el tipo 1 del ordenamiento de cdulas mucilaginosas y no mucilaginosas propuesto
por Ryding (1995), es decir celulas mucilaginosas solitarias o en pequenos grupos, separadas por pequenas
El segundo patron observado en Clinopodium (C. odorum. Fig. 3 D, y C. gillksii) corresponde a un epicarpo
con celulas que no producen mucilago las que, a diferencia de las no mucilaginosas del patron anterior, son
alargadas tangencialmente en transcorte. Estas caracteristicas celulares son similares a las observadas en C.
vulgare por Wojciechowska (1966). El engrosamiento de las paredes celulares en las especies aqui estudiadas
esuniformemente delgado, lo que difiere de lo reportado por Martin Mosquero et al. (2004) para C. vulgare
subsp. arundanum, quien describio e ilustro a las celulas no mucilagionosas con paredes radiales engrosadas.
Por su parte, C. nepeta esta reportada como especie no mucilaginosa (Moon et al. 2009) a la par que Martin
Mosquero et al. (2004) describe la estructura de su epicarpo acorde a la ausencia de mixocarpia. Por nuestra
parte, pndimos comprobar que esta especie, ademas de secretar mucilago, posee las caractensticas del epicar-
po del patron 1 senalado mas arriba.
La estructura del epicarpo en las especies no mucilaginosas de Nepetoideae es vana e. asi, e epicarpo
puedeestarintegrado por esclereidasexclusivamente(p.e.UpeAmu,,Rydmgl995,2010),s61o por celulas con
lodas sus paredes delgadas (p.e. Clmopodium. este Irabajo), o todas engrosadas (p.e. H^pBs, obs. pers.) o solo
por celulas con paredes langencialesdelgadasyradialesengrosadas(p.e.C. vulgare subsp^^nru^nnrMartm
Mosqneroetal.2004).opa.edestangencialesengrosadasyradUlesdelgadasCp.e.tnph^usAdans
«erFisch.&C.A.Mey.,Meebumu Britton ex Small&VaU,Budanstev&Loboval997);porulnmo,laseelulas
delepicarpopuedenaltemarcontricomasCCiinoHi“»'spp,ostetrabajo). toideaevarla
Mesocaipo.— Acorde a Ryding (1993, 2010), el niitnerodecapas del mesocarpo e as ejK
Jesde 2-15 (25), incinyendo la capa de escleteidas. En Clim>H‘“-". me^carpo consta de 3-4 “pas
dfendanclaratnenteen3reglones,porlalortnaytipodecelnlas(aplanad«o.sodnunetr.cas,^^^^^^^^
casoesclerenqulmaticas)ysucoloraeidn(tntensamentete«idasoincolo.^Estaorgan.za^^^^
esquematizado por Ryding (2010) para Mentheae. Sin embargo, en Clmopodium a region ex ema e
-gPu sc trate d! espec Jmixocarpicas o no; en las primetas, las celnbs son
■Musoseintensamente tenidas, a difemtcia de lo que observa Martin Mosquero et aL (20(M Pa™ ^
11 ^ c n muTinc rpdondeadas e incoloras tal como se indica
que en las no mixocarpicas las celulas son ">« “ “ de celulas iso-
corresponde a las esclereidas en empalizada, es el caracter mas variable
bolivianum subsp. tarijense) o alargadas radialr
y niimero de cristales en las esclereidas e
tudiados, pudiendo s
(C. broivnei, C. darwinii. C. gilUesii, C. odoruni). La pre^ncm^^^
"nr-goaserconsideradoAstCvulgurcsubspurundan^de^^^^^^^
2) mtentras que C. nepeta pt^e sol
s de Argentina aquI attaluadas que care^ ^
lumen (hast!
^cies nativas de Argentina aqui analizadas que
®iportancia taxonomica de este caracter al senalai
po es unistrato de cdulas parenquimaticas. Segiin
paredes celulares tiei
la presencia o ausencia de engrosamiento en las
, hemos observado que la mayoria d'*
eimportancia taxonomica. „-„j„iinoescalariformecoincidiendo con Martin
ee„paredescelnlaresconengrosan;^».»^
214
Brauchler et al. (2010) analizaron ca. 50% de las especies de Clinopodium mediante datos de ADN nuclear (ITS)
y cloroplastidial (tmK/tmL-F), definiendo numerosos subgrupos monofileticos dentro de este genero s. 1.
En el grupo Kurzamra, con buen soporte en el analisis cloroplastidial (cpDNA), estos autores ubicarona
Clinopodium danvinii junto a Kurzamra pulchella (Clos) Kuntze y otras dos especies: Clinopodium mukiflorm
(Ruiz & Pav.) Kuntze y Cuminia eriantha (Benth.) Benth., estas ultimas endemicas del centro-sur de Chile.
Desde el punto de vista carpologico, C. darwinii y K pulchella son mixocarpicas y comparten similar estructura
del pericarpo (obs. pers.); estos caracteres se suman al habito de matas en forma de cojin propio de zonas semi-
deserticas, si bien C. darwinii es de la estepa patagonica a diferencia de K. pulchella que vive a grandes altitudes
(3500-4100 m). En cuanto a las diferencias, estas dos entidades se distinguen no solamente por su caliz (osoi-
ramente biiabiado, tubuloso, lobulos netamente no subulados y no plumosos en C darwinii vs. actinomorfo,
tubo muy corto y cilindrico-infundibuliforme, lobulos largamente subulados y plumosos en K. pulchella) sino
tambien por la forma (piriforme vs. oblongo ovado) y tamano de los frutos (1,8-1, 9 mm vs. 1,5 mm). Un rasgo
destacable diferencial son los cristales en el interior de las esclereidas que estan presentes en K pulchella (obs.
pers.) y totalmente ausente en las especies de Clinopodium analizadas.
Clinopodium bolivianum aparece agrupado con especies de Minthostachys en el grupo Minthostachys,
tamo en el analisis cpDNA como en ITS (Brauchler et al. 2010); estos resultados agrupan a C. bolivianum junto
a C. gtlhesu, C. axillare (Rusby) Harley y C. vanum (Epling) Harley & A. Granda, los que, morfologicamente,
sonsimilarespor la forma del cahz (campanulado o cilindrico). Por su parte, Schmidt-Lebuhn (2008) agrupaa
estas imsmas especies junto a otras, bajo el grupo parafiletico Xenopoma en el arbol obtenido del anaUsis
c adistico morfologico y tambien en el logrado por analisis molecular. En cuanto a las especies argentinas, C.
M^umumy C. em,»re„,adas segUn estas propuestas, poseen frutos sitnilares por compartir tricomas
T hi ' por la forma, tamailo, reaccioii mudlaginosa y estructura del pericarpo (A
a a . be cjecen de dat^ carpologicos para las ottas especies de Clinopodium del grupo Minthostachys.
n.™ en , T ^ ol atiallsis molecular de Brauchler et al. (2010) donde apa-
rece en u.« postemn trresuelta demro de los Clinopmfium del Nuevo Mundo. Esta especie no presenta parheu-
carpologia siendo similar en la estructura del pericarpio y en la reaccion mucilagi-
laridades destacables i
nosa a C. bolivianum y C. dat
Por ultimo, Clinopodium odorum al
ha sido incluido en ningiin analisis filogenetico.
AGRADECIMIENTOS
herbarios ACOR, CORD, CTES LIE y s7por noner ^ V P^^^sonal tecnico de los
por la busqueda de bibliografia de diTicil accL a las d % ” material de estudio, a M. Cuaso b
ilustracionhnealdeestetrLjorawZ^’^^^^^
sugerenciasrecibidasparame^orLlmanuscriruT"'’'''^''^'''^''’^'^^^^^
concedida. ^ ^ pnmera autora agradece a Secyt-UNC por la beca doctoral
Bonzan, N.E., V.S. B.m. and G.E Barboza. 2011
(Lamiaceae) de Argentina. Caldasia 33(2):349-366
Brauchler, C.. H. Meimberg, and G. Heubl. 2010 Molecular
l^Gmera of Labiatae: status and classite
.. Royal Botanic Gardens, Kew. Pp. 51 1 -522.
215
DuletiC-LauSeviC, S. and P.D. Marin. 1999. Pericarp structure and myxocarpy in selected genera of Nepetoideae
(Lamiaceae). Nord. J. Bot. 19:435-446.
Gonzalez, A.M. and CL. Cristobal. 1 997. Anatomia y ontogenia de semillas de Helicteres Ihotzkyiana (Sterculiaceae). Bon-
plandia9(3^):287-294.
Harley, R.M. 1 985. Labiadas. I. Hyptis. In: R. Reitz, ed. FI. II. Caterinense, fasc. LABI: 1 -69.
Harley, R.M. 2008. Lamiaceae. En Zuloaga, F.O., O. Morrone and M J. Belgrano, eds. Catalogo de las Plantas Vasculares del
Cono Sur {Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). Monogr. Syst. Bot. Missouri Bot. Card. 1 07:2362-2392.
Harley, R.M. and A. Granda Paucar. 2000. List of species of Tropical American Clinopodium (Labiatae), with new combina-
tions. Kew Bull. 55:917-927.
Harley, R.M. (and 1 2 others). 2004. Labiatae. In J. W. Kadereit, ed.The families and genera of vascular plants, Vol VII, Flower-
ing plants: Dycotiledons (Lamiales except Acanthaceae including Avicenniaceae). Berlin and Heidelberg: Springer
Verlag.Pp. 167-275.
Husain, S2.. P.D. Marin, C. SiuC, M. Qaiser, and B. PetkoviO. 1 990. A micromorphological study of some representative genera
in the tribe Saturejeae (Lamiaceae). Bot. J. Linn. Soc. 1 03:59-80.
Johansen, D.A. 1940. Plant microtechnique. McGraw Hill Book Co, New York, New York.
Martin Mosquero, M.A., R. Juan, and J. Pastor. 2004. Estudio de las nuculas de Calamintha Mill, y Clinopodium L. (Lamiaceae)
en el suroeste de Espana. Lazaroa 25:1 35-1 41 .
Moon, H.K., S.R Hong, E. Smets, and S. Huysmans. 2009. Micromorphology and character evolution of nutlets in Tribe Men-
). Syst. Bot. 34:760-776.
Munsell, A.H. 2000. Munsell soil color charts. Munsell Color Company, I.N.C., Baltimore, Maryland, U.S.A.
Ryoing, 0. 1992. Pericarp structure and phylogeny within Lami
Bot. 12:273-298.
Ryding, 0. 1 994. The importance of pericarp structure in the classification of Labiates. Lamiales Newslett. 3:1-3.
Rywng, 0. 1995. Pericarp structure and phylogeny of the Lamiaceae - Verbenaceae - complex. PI. Syst. Evol. 198:
101-141.
Ryoing, 0. 2001. Myxocarpy in the Nepetoideae (Lamiaceae) with notes on myxodiaspory in general. Syst. & Geogr. PI.
71503-514.
Ryding, 0. 2010. Pericarp structure and phylogeny of tribe Mentheae (Lamiaceae). PI. Syst. Evol. 285:1 65-1 75.
Scandaliaris, M. and G.E. Barboza. 2011. Caracterizacion morfohistologica de frutos de especies argentinas de Lepechinia
(Lamiaceae). Bol. Soc. Argent. Bot.
Scanoauaris,M.,
(Griseb.) Epiing y"tomillo
Naturales del Litoral - III Reunion Argentina '
Schm'dt-Lebuhn, A.N. 2008. Monophyly and phylogenetic relations
amined using morphological and nrlTS data. PI. Syst. Evol. 270:25-38.
Stearn,W.T. 1983. Botanical Latin. Ed. David & Charles. Newton Abbot. Devon.
Wojoechowska, B. 1966. Morphology am
medicinal species. Monogr. Bot. 21:3-244.
''''ooo,J.r.|.20i
(Supl.):120.
lo" Hedeoma multiflorum Benth. (Lamiaceae, Nepetoideae). XI Jornadas de Ciencias
Bmily Labiatae with particular i
T Bull. 66:199-226.
216
Journal of the Botanical Research Institute of Texas 7(1)
Lorraine G. Bonney. 2011 . The Big Thicket Guidebook: Exploring the Backroads a
Texas. (ISBN: 978-1-5744-1318-2, cloth hbk). Big Thicket Association, University of North Texas Press,
1155 Union Circle #311336, Denton, Texas 76203-5017, U.S.A. (Orders: www.unt.edu/untpress, www,
tamupresscom, 1-800-826-8911). $29.95, 848 pp., 100 illus., 16 maps, bib., index, 6" x 9".
Whether you are the “Quick, Honey. Grab the kids and suitcases and let’s hit the open road!” or the “I’ll just curi
up in my favorite easy chair and read without actually having to go anywhere” type, you will undoubtedly en-
joy this large, carefully researched, informative, colorful, and delightfully humorous book. Canadian by birth,
Lorraine was married to the late Houston attorney Orrin H. Bonney. They were co-authors, and after his death
Lorraine finished two books. The Grand Controversy: History of Climbing in the Tetons in 1934 and Wyoming
This volume is actually Number 6 in the Temple Big Thicket Series, published by the Big Thicket Associa-
tion (University of Texas Press, Denton). It contains a wealth of background information, accompanied hy
current maps, very specific instructions, and delightful tales of the people who lived there — and still live
there— throughout the known history of the area. Be sure to read the General Introduction (pp. x-xiv) fora
quick assessment of what you are about t
to enjoy.
If you are planning to visit the Big Thicket, by all n
Texas and renting a car to visit the area— or for that mati
ms, take this volume with you! If you’re flying to
, if you live in Texas but have a passel of kids and
luggage— you might hesitate because of the book’s size. Nevertheless, you will benefit greatly by having it with
you. Part I, entitled “Some East Texas History,” provides a quick overview; Part 11, “Some Southeast Texas
Towns,” provides a great and often humorous account of the colorful towns and their equally colorful inhabit-
ants along the way; and Part III, “Roadside History of Southeast Texas, ” provides very, very specific-and cm-
rent!-road information about getting to all the colorful places. It identifies 15 tours, but it also includes spe-
cific sections in the larger areas. (How specific? Well, as an example. Tour 6 also has a Tour 6A, Tour 6B, and
Tour6CI)PartlIIs.artsonp. 199 and continues top. 787.Andii is allbothfunand fascinating, aswell as lelling
you specifically how to get to your intended destinations.
in the General Introduction, Professor Emeritus Pete A.Y. Gunter refers to the late comedian Johnny Cat-
sons fattwus chum that there was only one fruitcake in the entire world; it Just got passed from one family to
another. He continues:
^n one respect Lo™ne Bonney's The Big Thicket Guidehook is simple. It is a travelogue, guiding the
re^erthroughtheBtgThlcket.But,likethe-IhicketUtself,herwrttingisandhastolmaLgleolfolklou
Zret
indeed-the texture ofafruitcake:uneven,inpartsour,inpartsweet at times fun at times hard to digest.
and extremely rich (in details).” i sweet, at times tun, at times hard to a g
-Helen Jeude, VoluMcer Tf
, cai Kesearch Institute of Texas, Fort Worth, Texas, U.S.A.
SYSTEMATICS, PHYLOGENY, AND BIOGEOGRAPHY
OF CHAETOLEPIS (MELASTOMATACEAE)
Daniel Grimm and Frank Almeda
Department of Botany
California Academy of Sciences
55 Music Concourse Drive, Golden Gate Park
dgrimmO I @yahoo.com; falmeda@calacademy.org
RESUMEN
INTRODUCTION
Mi,. (Melastomalaceae) is a small mostly neotmpical genus of montane shmbs and subshrubs
fatelyannuaU). As interpreted here, it consismofll species thatare readily charac,enaed^4.merous,d.^^^
«emonous flowers, isomVic mostly inappendrcula.e stamens that lack welWeveloped pedoconnecuves^
MocuUr.aptcallyse, use ovaries, and cochleate seeds withatestathatconsists mostly or compre^flatened
i»h"ligi.adngcefc. The genus,asithasbeen circumscribed inthe past, hasindudedspecresof Andean South
America, the Guayana Highlands, southern Central America, and Cuba Recently the motrotypic genus
''-ophlr.NaudimLmweltropicalAfricawas.ramferredtoCIwtdephO-que^Felix
Study has attempted to evaluate Chaetolepis or Nerophila. Because the monophyly of the geographically dispa-
rate assemblage of^ taxa refel^ed to Chaetolepis had never been evaluated, it seemed ripe for testing.
218
netic relationships of the c
cladistic techniques. A ta:
mbersof^^
The primary goals of this study were to determine if Chaetolepis is monophyletic, assess the phyloge-
species based on morphology, and evaluate character evolution by using
L taxonomic revision is provided that includes detailed descriptions of all recognized
n microscope images of seeds for selected species, a summary of known chromosome
numoers, miormation on habitat and geographic distributions, flowering phenology, distribution maps, il-
lustrations of all recognized species, citations of specimens examined, and a list of excluded taxa. This was
supplemented by limited field study of four species in Costa Rica, Colombia, and Venezuela.
Tribal and Generic Affinities. — Chaetolepis belongs to the tribe Melastomeae, the core i
are characterized by cochleate seeds with a testa that is tuberculate or papillate, a crown oi inenomes ur ap-
pendages on the ovary apex, and the presence of staminal pedoconnectives (Renner and Meyer 2001; Freire-
Fierro 2002; Michelangeli et al. 2013). Based on overall morphological similarities, Wurdack (1973) hypoth-
esized that Chaetolepis is most closely related to Tibouchina Aubl. No cladistic analyses or molecular data,
however, had been used to assess the possible sister relationships of Chaetolepis. Other neotropical relatives
of Chaetolepis include Aciotis D. Don, Arthrostemma Pav. ex D. Don, Heterocentron Hook & Am., Microlicia D.
Don, Microlepts (DC.) Miq., Monochaetum (DC.) Naudin, Nepsera Naudin, Pterolepis (DC.) Miq., and Rhexia
L. based on a study using ndhF and rpll6 cpDNA sequences of 24 members of Melastomeae and outgroups
(Renner and Meyer 2001). Based on an expanded molecular sampling using DNA sequence data, Bucquetia
DC., Castratella Naudin, and Monochaetum are now considered the closest sister taxa of Chaetolepis (Peimeys
etal. 2010; Michelangehetal. 2013). Like Chaetolepis, the majority of species in thesegenera are also restricted
to higher elevation montane environments in the Andes of northwestern South America and all have tetramer-
ousn^ers, an apicallysetulose ovary thatUfourlocuUrandfree tarn the hypamhium, and capsularfrulB.
Morphology (Characters of Taxonomic Value).— The species of Chaetolepis are easily dis-
tinguished from each other by characters involving the indument, anther connective, seed surface, petal
coloration, and petal margin. The indument found on leaves, intemodes, nodes, and floral organs includes
eglandular tnchomes, glandular trichomes, barbellate/dendritic trichomes, and scales. The distribution of the
surfaces^IfTeeeLtS^lnTfl^^l structures vanes greatly. It ranges from absent to densely covering the
nrolnno^ft t 1^ 1 i organs. The anther connective prolongation ranges from absent to minutely
dwTrwtb T
imerdmatim'^ZttZ 1^^^^ prevailingly smooth with compressed/flattened
low to Lk or iTalel H t interdigitating cells. The petals vary in color fromyel-
low to pink or magenta and their margins are either glabrous or ciliate
ainoris.ics.udi«(Su„dCl9MTribJuW^™'’'lTo^^^
the soecies have been k a • ' Wurdack 1973; Almeda 2001). Over the past 80 years some of
.he placemen, of
Leon and Alain 1974; MamnsT989)
ciesa„d.h.eevade.ies. Because a,
identify symapomorphies and only cumorv aupmlon w, ^
vegetative and reproductive chamctem. llco^tunWiSr “ '
some of the chatacters used to characieri °f ” monograph of Melastoma.aceae, for example
varyingin color from rose .op„rple;4-Cw^t.ts“f"''“^^^^
connectives; and ovoid cochleateLds With smZr
ofasmallersamplingofspecielTstiWbuX^^^^^^^
and the seed surface appeam -smooth or fit^ly and dimta T
sculptural details of the seed testa because n SFM ,1 ^ ^uriculate.” No prior work mentioned any
detail relating to the seed and other structures mavlT'^ ^ of consistency and
iolepis, perhaps biased by the four-merous flowers and greltirreduc^
‘U greatly reduced pedoconnective. Furthermore, some
219
doubt about the monophyly of the genus has persisted because no cladistic analysis of the group has ever been
undertaken to evaluate character states and their evolution.
Ingroup. — ^All sixteen taxa thought to comprise Chaetolepis form the ingroup for this study. Chaetolepis alpina
var. alpina, C. alpina var. latijolia, C. anisandra, C. cubensis, C. ci^odontisii, C. gentianoides, C. lindeniana, C. lori-
carella, C. microphylla, C. perijensis var. perijensis, C. perijensis var. glandulosa, C. phelpsiae var. phelpsiae, C.
phelpsiaevar. chimantensis, C. santamartensis, C sessilis, andC. thymifolia.
Outgroup.— To assess the monophyly of Chaetolepis, seven species among the genera Tibouchina, Mi-
crolicia, and Monochaetum were used as the outgroup. The exemplars used were T. gleasoniana Wurdack, T.
narinoensis Wurdack, T. wurdackii Almeda and Todzia, Microlicia canastrensis Naudin, M. obtusifolia Cogn.
ex R. Romero, Monochaetum amabile Almeda, and M. compactum Almeda. Tibouchina was chosen as part of
the outgroup because Wurdack (1973) hypothesized that it likely shares a most recent common ancestor
with Chaetolepis based on many shared characteristics, such as perigynous flowers, campanulate or urceolate
hypanthia, superior ovaries that are apically pubescent, capsular fruits, and cochleate seeds. In addition, T.
^easoniana, T. narinoensis, and T. wurdackii were chosen as part of the outgroup because the tuberculate seed
testa and foliar scales (see Todzia and Almeda 1991) are identical to those found in C. cubensis. The species of
Microlicia and Monochaetum were chosen as other members of the outgroup because of their close phylogenetic
proximity to Tibouchina based on molecular studies by Renner and Meyer (2001).
Material Examined.— Over 400 herbarium specimens, including types, were used in the phylogenetic
analysis and taxonomic revision. Specimens from the following herbaria with significant holdings of Central
and South American Melastomataceae were borrowed or studied on site: BM, BR, CAS, COL, DS, DUKE, F,
FMB, G, GH, HECASA, K, M, MO, NY, P, US, and UTMC (acronyms according to Thiers 2012). Internet images
of type specimens OSTOR Plant Science) were examined for those taxa that were not readUy available on loan.
Characters.— Forty-one vegetative and floral characters were used in the analysis (Tables 1 and 2). The
type of indument covering both vegetative and floral structures was particularly informative in this study.
Data Analysis.— The software program PAUP* 4.0bl0 (Swofford 2002) was used to generate phylogenetic
estimates. Maximum parsimony with a branch and bound search was selected to evaluate the data matrix that
was generated in Maclade 4.03 (Maddison and Maddison 2000). Two analyses were conducted. Analysis 1
was performed to assess the monophyly of Chaetolepis and analysis 2 provided insight into ingroup relation-
ships after the exclusion of one of the species of Chaetolepis (C. cubensis) was prompted by the results of the
first analysis. The first analysis used 41 morphological characters and 23 taxa. Sixteen species formed the
presumed ingroup and seven species from the genera Tibouchina, Monochaetum, and Microlica formed the
outgroup. Default options were used to run the analysis. In analysis 2, a backbone constraint was appUed that
grouped all outgroup members plus C. cubensis together because of their unique floral and/or seed characters
grouped all ingroup taxa together. This was done by constructing a tree in MacClade (Maddison & Mad-
dison 2000) with the data matrix, opening the file in PAUP (Swofford 2002), derooting the tree, loading the
as a backbone constraint in the analysis, and finally enforcing the constraint in the search.
To evaluate the strength of the different clades in the various phylogenies generated, a bootstrap analysis
employing a heuristic search was utilized in PAUP. Stepwise-addition branch swapping was emp oyed; ot er-
icters among the lineages of Chaetolepis. The “l
. default options were used.
MacClade was used to infer the evolutic
ges,” and “all possible changes” options were chosen.
■O gain insight into the historical biogeography of Chartolepis, mo a™lys« were conducted^
erform^sri .. — ; j_ r^rioin and disoersal patterns of the diilerent taxa witnm
MacClade, a data mai
performed to provide insight into the origin and dispersal patl
^«nca. Central America, and Africa. This analysis used Fitch opir
'°"P«ratingallChaetolepi
5 {1); sessile glands (2).
Grimm and Almeda, Systematics of Chaetolepis
andalOOxoil
in which they are found (Table 3), and one of the 26 most parsimonious tree topologies generated in PAUP with
polytomies resolved arbitrarily. The second analysis was performed to infer how Chaetolepis taxa may have
migrated within the Andes. This analysis differed from the first analysis only in the number of area states (five)
used. In the analysis, all equally most parsimonious reconstructions (MPR’s) were examined.
H. Chromosome Cytology
Horal bud material was collected in the field, fixed in modified Camoy’s (Bradley 1948) solution (4 chloro-
form, 3 ethanol, 1 glacial aceitic acid, v/v/v) for 24 hours, transferred to 70% ethanol and stored under refrig-
eration until studied. The new chromosome count reported here for Chaetolepis cufodontisii v
acetocarmine smears of pollen mother cells using a Zeiss light microscope with phase contrast
immersion objective. Drawings of meiotic figures were made by camera lucida at a magnification of 2100x. The
voucher collection for this new count is marked with an asterisk among specimens cited.
:e all vegetative and reproductive characters based on dried her-
e taken from either one mature open flower in the center of an
were scored from matur^branches. To eLure that aberrant characteristics were r
specimens were examined and those with typical traits were used. . i ■
A scanning efecmn microscope (SEM) was used io study the seed morphobgy of “actoljs
Seeds were affixed to aluminum stubs with double-stick tape and sputter-coated t«th gold-palladium
^l»y.San,ples were photographed in the scanning electron microscope laboramryatthe^^^^
“(Sciences using a Leo 14M BP scanning electron microscope at 20kV, Voucher collectton data for the SEM
‘“'“ges are provided in the relevant figure legends.
RESULTS
ot being recorded, numerous
^fogeny.-in the first cladistic analysis, 22 n
was 0.66 and the reter
224
Cl, of which nine are unique, three are homoplasious and one is ambiguous. The unique characters include
appresse ’ ^te, flattened scales, basally attached with a raised thickening near the point where the scale is at-
tached to the leal surface and an erose margin on the hypanthium base, hypanthium apex, adaxial leaf surface,
abaxial leaf surface primacy veins, branch surface intemodes, branch surface nodes Lercalydne lobe sinus,
alyxvesiuure excluding the midrib, and calyx midrib (characters 3, 6, 11, 17, 20 23 25 28 31 respectively).
m homoplasious (char-
S^mes or b MUK/d ‘>"= ""■oo* egUndular trichomes, smooth glandular tri-
Grimm and Almeda, Systematics of Chaetolepis
ffi
226
n the intercalycine lobe sin
the abaxial leaf surface (betw
yellow petals. The homoplasious characters are smooth eglandular trichomes, smooth glandular trichomes, or
barbellate trichomes on the intercalycine lobe sinus. Clade D1 is supported by six characters, of which three
and one is homoplasious. The unique characters are roughened trichomes on
n primary veins), branch surface intemodes, and calyx midrib (characters 13,
and branch surface nodes (character 16 and 22). The homoplasious character is roughened trichomes on the
adaxial leaf surface (character 10). There are five ambiguous characters supporting clade D2 which include a
aenulate or serrulate leaf margin, punctate or sessile glands on the abaxial leaf surface, smooth eglandular tn-
chomes or smooth glandular trichomes on the abaxial leaf surface, branch surface nodes are glabrous or with
smooth eglandular trichomes, and yellow petals (characters 8, 12, 16, 22, 41). Clade El is supported by eight
characters, of which four are ambiguous and four are homoplasious. The four ambiguous characters include a
crenulate leaf margin, forked intercalycine lobe sinus, calyx vestiture that is absent or with smooth eglandular
s, and three to five leaf veins (characters 8, 24, 27, 38). The four homoplasious
im apex adaxial leaf surface, abaxial leaf surface (between primary veins), and
b smooth and elgandular trichomes (characters 5, 10, 13, 19). Clade E2 is sup-
. , characters of which two arc amhigttous and four ate homoplasious. The antbiguo.^ characters
are a serrulate leaf margin and punctate abaxial leaf surface (characters 8 and 12). The homoplasious char-
acters are sessile glands on the hypanthium base, hypanthium apex, adaxial leaf surface, and calyx vestiture
(characters 1, 4, 9, and 26). There are six characters supporting the clade consisting of only C. gentianotdes,
one of which is unique, four of which are ambiguous, and one of which is homoplasious. The unique character
is a seed testa with elongate, interdigitating cells. The ambiguous characters are abaxial leaf surface lacking
ghuds,stalked and variously hrauched trichomes at theintercalycmelohesinus,stnoothe^undulartrtchomes
™ the calyx lobes, and five elevated leaf veins (characters 12, 24, 27, 38). The homoplasious character ts
S~d. egUndular trichomes on the hypanthium base (character 2). There am three characters supporttng
the clade that includes C. phelpsiue var. phelpsioe, C. phelpsiae van chimmUnsis, C. thymi/oho and C. nnisnn
oneofwhichisan unclear derivedstate. This latter character stateUsmootheglandulattnchomesands^oth
Ihndnlar trichomes on the calyx lobes (character 30). The ambiguous char^ntrs are
Ckomes on the abaxial leaf surface and forked trichomes on the intercalycine lobe sinus (c arac _
ttreecharactets support the clade consisting of C. thymi/bliaandC. nrasnn ° ,
ahione of whichisan unclear derived state. The two ambiguous chametets are abaxial leaf surface withsess
*n*(character 12) and calyx vestiture devoid oteteCc^^m^^
ambiguous (not shown). However, a majority of the most parsimon
®alysis places Sierra Nevada de Santa Marta at the most basal node.
bCladistic Analysis
ofTraditionally CiKumscribed to the outgroup t
fflonophyletic as traditionally circumscribed because C. cubensis is m congeners (Fig. 1).
'^^china gkasoniana, Tibouchina narinoensis, and Ttbouchma ^ rtnberculate seed testa (Fig. 6 d-f)
^lepis cubensis and the species of Tibouchina in the ^ Vb to % of their
•nchomes modified into scales that are adnate to the fohar
'«'gtb. This species was probably transferred to Chaetolepis by appendage. Our results, however,
^markedly reduced pedoconnective and genus. Based on the phylogenetic results
^onsirate that the above charactersarenotsynapomo^hi^ortheg^^^^^^
*e character differences, we propose the return of C. cubensis to Tiboucnma
Character 1 - Andes Mountains (Eastern Cordillera,
Character 3 Talamanca Mountains, Costa Rica
Character 4 - Loma Mountains, West Africa
a branch and bound aarth of Onietolepis (analysis 2).
lillli
Likely sister outgroup oj Chaetolepis when excluding C. cubensis (Chaetolepis sensu strictoX— An analysis of
morphological data used in this study suggests that Monochaetum, not Tibouchina as proposed by Wurdack
(1973), is likely sister to Chaetolepis sensu stricto. The characters that support this relationship are a dorsal
orientation of the anther pore, four-merous flowers (Fig. 3). However, it cannot be ascertained from this study
whether these are synapomorphies for Chaetolepis and Monochaetum or for the remainder of the outgroup. Ad-
ditional similarities between Chaetolepis and Monochaetum include a four-locular ovary, cochleate seed shape,
compressed/flattened cells of the seed testa, a montane habitat in tropical America, and a chromosome number
based on X = 9 (Almeda and Chuang 1992; Solt and Wurdack 1980), although the latter two characters may
ultimately be demonstrated to be symplesiomorphies for these genera.
Infrageneric relationships and character evolution.— Despite the fact that the strict consensus tree exhibited
several areas of nonresolution, four major clades were resolved (clades D, E, F and Chaetolepis loricarella in Fig.
2). These clades are based largely on the kinds of trichomes found on various morphological organs, a charac-
ter that Wurdack (1986) surmised was of phylogenetic importance among Melastomataceae. Trichome diver-
sity in Chaetolepis includes scales, barbellate/dendritic trichomes, smooth eglandular trichomes or smooth
glandular trichomes, and sessile glands. Chaetolepis loricarella is the only species in the ingroup with an mdu-
ment of scales: thus this character is autapomorphic within Chaetolepis.
The clade comprising Chnelokpis micropkyU C. scntamartensis. and C. lindmam (Fig. 3. Clade Dl) m
the second cladistic analysis showing character state change is supported by four unambiguous c
whose character states all involve barbellate trichomes. The only other species in the analysis that has barbel-
late trichomes is Monochaetum compactum (a species within the outgroup). Thus this character state is likely to
be derived within Chaetolepis. Similarly, the clade comprising Chaetolepis gentianoides, C. phel^iae var. p e p-
siae, C. phdpsiae var. chimantensis, C thymifolia, and C anisandra (Fig.
biguous characters states that all involve smooth eglandular tricho " f
that has this character is Monochaetum amabile. Thus this charac
Chaetolepis.
-The clade comprising Chaetolepis cufodontisii, C. perijensis
e is also likely to be derived within
r. perijensis, C. perijensis var. glandulosa,
whose character states all involve the presence of sessile glands on the adaxial leaf snrface. This ts clearly a
derived character State within Chaetolepis for these characters.
-Clme.oleplsloricnrdla,C.llndemnnn,C.santnmariends,andaUtheontpoupt^haj^^^^^^^^^
petals (including C. cubensis), whereas all other species of Chaetolepis have yellow pe a . g y f
* am ambiguously placed in the character ^ cILe L“r^ins, Is do C. cubensis
Chaetolepis. The same three Chaetolepis spp. sensu stncto pe r-t, rrFntitmit acwpllasonce
andaUspecies of ribouchinn.Thischatacter is inferred to have evolved twicewulnnClinetoleps, as wel
Within the outgroup, Tibouchina. ^ elongate somewhat convex or domed
PtricUnal^nlTnfheKed ttsla (Fig. 6 g-i)
ihe second analysis. All other species of Chaetolepis have pendiM ce wa s o (oveolale or
^ttened and no. conspiculously elongate
PossiMe relationships of thisspecieswithotherrmm^
H“‘»™«'l!iogeography.-ThefirstF.tchoplim
aodVenezuelan Andes as the place of Origin fortheg ffg.^)^^^^^^^^^^^^^^^^^
g«ts an origin in the Sierra Nevada de ^^^^os. The Sierra Nevada de Sania Marta in far
constructions of area onto the topology result m d ^ ^ ^
■lotthem Colombia and its neatest montane neighbo , j Cgins (Todd & Carriker 1922; Irving
•heColombianAndeslatedtstinctmounuinsystemswtthindependentongt
230
Journal of the Botanical Research Institute of Texas 7(1)
massifwasu™lenva,erum,Uhe„.id-Te.;;^; ^"*''■0'- & Garza„-C. W971 ^
stantial block-lifting in the early Pleistocen7(Gansser continued with sul
Colombia and adjacent Venezuela (including I w H p
taoncluclmgtheSierradePenjaandMeridaAndesjbeganintheUpperCreu
231
'e® bu. substantial uplift abuve sunuunding terrain was only achieved in Teriiary time ^pnning in the
fdMcene with continued upward movement culminating in the Pliocene (van der Hammen 1961, Imng 1975,
Simpson 1975). • W t Af '
Kenner and Meyer (2001) postulated that the occurrence of Chaetolepis gentianoides in ^^est Afnca re
^ from a long^listance dispeisal event because of the -great morphological smularity
American “and West African taxa" of Melastomataceae,
support this hypothesis. In
study using molecular data, Givnish et al. (2004) concluded that long-distance dispersal was
232
sponsible for the present distribution pattern of Pitcaimia U Her. (Bromeliaceae), a genus centered in tropical
America with one endemic species in West Africa iPitcaimiafeliciana (A. Chev.) Harms & Mildbr.]. That study
postulated that many other South American-African disjuncts (see Thorne 1973; 1992) have resulted from
long-distance dispersal rather than vicariance events, because the split of these two continents is too old (ca.
80 million years) to account for many disjuncts, especially those that are highly similar morphologically, such
as Chaetolepis. The timing of dispersals to Africa from the Andes cannot be estimated because data from the
present study are based only on branching patterns, and not branch lengths. An independent test of whether
the amphi-Atlantic disjunction in Chaetolepis is due to ancient vicariance or more recent-long distance disper-
sal, as well as the timing of the other two dispersals inferred from the data (Venezuelan Guayana and Tala-
manca Range) will require clade divergence estimates based on molecular data, such as DNA sequences.
As for the distribution of Chaetolepis species within South America, long distance dispersal of different
taxa within Melastomataceae, Bromeliaceae, Cactaceae, and other families likely occurred intermittently
within South America because of climatic fluctuations during the Pleistocene, a period extending from 1.8
million to 1 1,550 years ago (Berry 1982). The movement of species within South America and beyond is specu-
lative but it may have been aided by bird migration. As bird species, like the Swainson’s Hawk (Buteo swainsoni
Bonaparte), traveled to breeding or wintering grounds along the Atlantic Coastal fly way or bird species like the
Arctic Tern (Sterna paradisaea Pontoppidan) migrated annually across the Atlantic from North America to
Europe, West Africa, and South America and back they may have unintentionally picked up Chaetolepis seeds
on their feet as they walked through mud in search of food (Lincoln et al. 1998). The Arctic Tern is the only
example of a bird with a migratory path passing through both South America and Africa, but it is not known to
frequent higher elevations in West Africa where Chaetolepis occurs. Further studies using molecular data are
needed to conhrm or reject the placement of C. gentianoides in the genus and to estimate divergence times that
could assess the likelihood of long distance dispersal to Africa.
Venezuelan Guayana taxa such as Chaetolepis phelpsiae var. phelpsiae and C. phelpsiae var. chimantensis in
clade E may have evolved as a result of vicariance. These species are closely related and have ranges that are ad-
jacent to one another. Probably through time, erosion of the sandstone landmass of the tepui landscape isolated
populations and led to speciation. Givnish et al. (2004) estimated that “chemical dissolution and erosion should
cause the margins of adjacent tepuis to retreat from each other at a rate of nearly 2.4 km/million years. Such low
rates would permit vicariance to play a role only on tepuis that are currently quite close to each other. Vicari-
ance seems unlikely to have played a role in speciation on tepuis that are now further apart than ca. 35 km.”
II. Chromosome Cytology
Meiotic chromosome numbers have been published for three species of Chaetolepis.
Favarger (1962) reported n = 9 for C. gentianoides (as Nerophila gentianoides Naudin) and Solt & Wurdack
(1980) reported n = 9 for C. microphylla and n = 18 for C. lindeniana. We report here the first count for C. cufodon-
tisii; n = 9. In this species meiosis was regular in all cells studied and all figures examined at diakinesis and telo-
phase II were clear and consistent (Fig. 7). Present data indicate that the base number for Chaetolepis isx = 9.
A base number of x = 12 has repeatedly been invoked as the original base number for the Melastomataceae and
the order Myrtales (Raven 1975; Johnson & Briggs 1984; Graham et al. 1993; Almeda 1997a; Almeda & Rob-
i^on 201 1). A base number of x = 9 could be derived through descending dysploidy. Thus, the three species of
aeto epts wit n = 9 have retained the base chromosome number in the genus whereas C. lindeniana, with
n- 8, is clearly tetraploid based on x = 9. Among the significant outgroups and sister genera of Chaetolepis,
BucquetiaDC.hasn = 18(Solt&Wurdack 1980); Monochaetum (DC.) Naudin hasx= 18 (Almeda 1997b); and
Tihouchma Aub . has x = 9 (Almeda 1997b). No chromosome numbers are known for the ditypic genus Costra-
tellu Naudim Of the five patterns of chromosomal evolution proposed to account for the origin and diversity
of numbers for neotropical Melastomataceae (Almeda 1997b; Almeda 2013; Almeda & Chuang 1992), Chae-
tole^s appears to fit the one in which intrageneric euploidy has played a role in speciation whereas the origin
of closely related genera such as Bucquetia and Monochaetum can be accounted f^ by paleopolyploid events.
Grimm and Almeda, Systematics of Chaetolepis
233
I
20 |jm
> indicated by DIAK (diakinesis), Til (Telophase II). a
=9,DIAK.b.Ccirfbdonl/s//,n = 9,TII.
Chaetolepis Miq., Comm. Phyt. 72. 1840.
(Bonpl.) Miq.
Erect or ascending, laxly or compactly brai
usually quadrate or subquadrangular, son
2. 1806-1808. Type: Chaetolepis mUrophylla
1 shrubs, subshrubs, or annual herbs. Distal cauline intemodes
les carinate; glabrous or covered with various kinds of tri-
— scales, or gianas; aistai cauime noaes somewhat thickened, often covered with trichomes, scales, or
glands like those found on the internodes but trichomes and scales when present tending to be longer. Older
intemodes terete. Principal leaves opposite, coriaceous or membranaceous, spreading or tightly appressed;
and abaxial leaf surfaces glabrous and punctate with sessile glands, or sparsely to densely covered with
Wchomes or scales; blades varying in shape from linear, oblong, elliptic, lanceolate, ovate, deltoid to obovate;
(l-)3-7 elevated, nearly parallel, longitudinal primary and secondary veins aU arising from the blade
'’ase or with secondary nerves arising from the midnerve above the blade base; margins ciliate or eciliate, en-
tire, crenulate, or serrulate, often revolute; blade petiolate or less commonly sessile or subsessile. Inflorescences
terminal (sometimes terminal on axillary branches);ilawers sohtary, paired, or in simple dich^ia (sometimes
ncompounddichasia). Bracts subtendingtheinflorescencesimilartopnncipalleavesmsize s pe,an in u
®ent, persistent or deciduous. Bracteoles subtending individual flowers ranging from anceo ate,
‘^c.oblanceolate, to narrowly ovate smallerin size thanbracts, persistent todeciduous.Hypanthia (at anthesis)
"rceolate to campanulate ovary free from hypanthium, calyx lobes, petals, and stamens inserted ^ the torus,
»yi«nthia (at maturity) simiJtohypanthiaatanthesisinbothsizeandshapebuU^^^^^
^“-triangular, apically acute sometimes rounded, margin fnngedw g rarelv erose
«theinargins,apexvaryingfromcuspidatetoobtuse,glabrous,withorwuno vellowor
ChMolepis alpim has been divided into two varieties, the nominate one and C aim
™niingtoCogniauxa89I) the nominate variety hasoblongleaveslO-15x3-VmmtdL
d^a mnivetu T m!"ho? to Cogniaux, we cou»
238
Journal of the Botanical Research Institute of Texas 7(1)
-.3,14:140. 1850 (Figs. 6 a-
is. Linn. Soc. London 28:50. 1871. Syn. nov. Type: COLOMBIA. Cundin/
V. 3400in, 1851-1857, Trianfl36J78 (holoiype: BM, photo of holotype, F
Yia 3:174. 1939. Type: VENEZUELA. E
O.TypeVENEZUEIA
y: Cordillera deBogoti,
It Auyan-tepui, elev. 2200 ni
A much branched subshrub 0.1-1 m tall. Older intemodes woody, terete and exfoliating with age. Distal cau-
line intemodes quadrate and carinate to narrowly winged, moderately to densely scabrous, the eglandular
and glandular trichomes 0.4-0.5 mm long, appressed to ascending. Distal cauline nodes similar to distal
cauline intemodes but with trichomes 0.7-1 mm long. Principal leaves coriaceous, adaxial surface moderately
covered with smooth, eglandular, evenly distributed spreading to erect trichomes; abaxial foliar surface mod-
erately covered with sessile glands and sparsely covered with appressed or sometimes spreading, eglandular
trichomes between the primary and secondary veins; moderately covered with sessile glands and moderately
to densely covered with erect, eglandular trichomes on the primary and secondary veins; blades 6-12 x 4-9
mm, ovate to nearly deltoid, acute to broadly acute apically, cuneate basally, 3-5-nerved, the outer pair arising
from the base and continuing one third or one half the length of the blade, margins ciliate-crenulate, petioles
2-4 X 0.23-0.5 mm, ciliate and/or sparsely beset with smooth trichomes but varying to glabrate. Inflorescence
terminal, flowers solitary or borne in a simple 3-flowered dichasium. Bracts and bracteoles 5-9 x 3-6 mm, oth-
erwise similar to the principal leaves, 3-nerved, sessile. Floral pedicels 0.5-1.0 mm long, or subsessile, moder-
ately to densely covered with minute, erect trichomes. Hypanthia (at anthesis) 3-4.5 x 2-3 mm, campanulate,
moderately covered with erect, smooth eglandular trichomes toward the apex, glabrous basally; hypanthia (at
maturity) with vascular ribs often conspicuous when drying. Calyx lobes 2-2.5 x 1-1.5 mm, erect, narrowly
tnangular, apically acute, with 1 or more smooth eglandular trichomes and/or smooth glandular trichomes
alongthe midrib, ciliate. Intercalycine lobe sinus with one or two forked trichomes or withasmoothegland^
ar tnchoine. Petals 7-9 x 4-5 mm, yellow, entire, glabrous, apically cuspidate, tipped with a single trichome
Filaments 3-9 mm long, yellow, anthers 2.5-4.5 x 0.5-1 mm. yellow, linear to arcuate, pore dorsally inclined
connective minutely prolonged 0.1-0.3 mm below the anther thecae. Ovary moderately covered apically with
smoot eglandulartnchomes.Style6-7mmlong,yelloworpalebrownwhendry.Seeds0.4-0.6mmlong,pale
Flowering.— September through January
cZ M ; ““ “ W50-2600 melcs. I. also «
” 2500-3400 melers.
Chaetolqris anisandra apnears to i i i
logical similarities as reflectTin the strict based on morpho
leaf surfaces, cauline nodes, intemodes and eglandular trichomes on ^
ciliate-crenulate leaf margins aleafbladethatisba 1 P««bermore, these species both hav<
nnectives, and a dorsally inclined anther nore
f«. 10. Oiaetolepis anisandra. A. Habit. B. Petal (adaxial surface). O. Stamen, dorsal view. Q. S
e view. D. Hypanthium. El. Seed, profile
Chaetolepis phelpsiae var. phelpsiae differs from C. anisandra in having smooth eglandular tnchomes that
distal region, an abaxial leaf surface that is punctate with eglandular
^ and secondary veins, an abaxial leaf surface that has eglandular
and glandular trichomes on the elevated veins, and cauline intemodes that are sparsely covered with eglandu-
lar and glandular trichomes (Fig. 22).
Chaetolepis thymifolia has been recognized as a distinct species based on 3-nerved leaves and ovate to
suborbicular petals (vs. 3-5-nerved leaves and obovate petals in C. anisandra). Because the petals of all species
in the genus are obovate and foliar venation is consistently 3-5-nerved in C. anisandra, we see no reason to
continue to recognize C. thymifolia. What has been recognized as C thymifolia occurs in central Colombia at
2500-3400 m whereas C. anisandra has traditionally been accorded a range extending from northern Colom-
bia to southeastern Venezuela at 1950-2800 m.
Representative specimens examined: COLOMBIA. Cesar Valledupar, paramo bajo entre Mamancana y Nabusimake, Carbono
(UTMC). Candtoamarca: El Portachuelo between UneandFosca. Cordillera Oriental vert. Oriental, en el mismo Boqueron, Cucureros
Jaramillo 28795 (US): Tacuate Valiev 3 km W of Gutierrez, 45 km S of Bogota, Grant 9651 (NY, US); Paramo de Chipaque, Karsten, s.n. (BR,
. Auyan-Tepui, Cardona 2720 (NY); Mt. Roraima, Quekh & McConnell 22 (K); Mt. Roraima, ascent of ledge along SW
: 1 157 (NY). Locality not given, Schombur^ 1013 (F, GH).
1. Field Mus. Nat. Hist., Bot. Ser. 18:792. 1938. (Figs. 7, 12). Type: COSTA
Erect, much-branched subshrub 10-40 cm tall. Distal cauline internodes quadrate, glabrous. Distal cauline
nodes sparsely to moderately covered with spreading or appressed, smooth eglandular trichomes. Older
branches woody, terete, and exfoliating with age. Principal leaves coriaceous, spreading, adaxial surface
sparsely covered with sessile glands; abaxial surface moderately punctate, sometimes sparsely covered with
antrorsely spreading trichomes between the primary and secondary veins, sparsely covered with eglanudar
trichomes on the primary nerve, sometimes glabrous; blades 6-10 x 1-2.5 mm, lanceolate to oblong-lanceo-
late, acute to rounded apically, acute basally, 3-nerved with an elevated median nerve and depressed secondary
nerves that extend the entire length of the blade; margin serrulate and eciliate; petioles 0.5-1.5 x 0.3-0.5 mm,
glabrous. Inflorescence a simple, terminal dichasium, sometimes flowers solitary or paired. Bracts similar to
principal leaves. Bracteoles 4-6 x 1-1.5 mm, also similar to principal leaves but subsessile or with petioles 1
mm long. Floral pedicels 0.5-2 mm long, glabrous. Hypanthia (at anthesis) 3.5-4.5 x 2.5-3.5 mm, urceolate to
subcampanulate, sparsely covered with sessile glands; hypanthia (at maturity) with conspicuous longitudinal
vascular ribs when dry. Calyx lobes 2.5-4 x 1-1.5 mm, erect to ascending, narrowly lance-deltoid, apically
acute, ciliate; calyx lobes sparsely covered with sessile glands and with antrorsely spreading trichomes on the
midrib, sometimes glabrous; intercalycine lobe sinus with one to three smooth flattened eglandular trichomes,
these sometimes bifurcate. Petals 5-7 x 4-5 mm, yellow, apex acute, margin entire, glabrous except for a single
terminal trichome. Filaments 4.5-5.5 mm long, yellow; anthers 2-3 x 0.4-0.6 mm, yellow, linear to arcuate;
pore dorsally inclined; connective minutely prolonged 0.2-0.3 mm below the thecae. Ovary apex sparsely to
:; style 9-10 mm long, yellow. Seeds 0.6-0.7 mm long, brown. Chromosome number: ti = 9.
Flowering. — ^All year.
Hc^itatand Distribution (Fig. 11).— Endemic to Costa Rica in San Jose and Cartago provinces on the Cor-
xposed slopes with shrubby vegetation at 2400-3600 m.
Discussion.-Chaetolepis cufodontisii is a much-branched subshrub readily recognized by the following
characters: adaxial leaf surface and hypanthium covered with sessile glands; abaxial leaf surface punctate be-
;en t^ secondary veins with a few eglandular trichomes on the elevated veins (Fig. 12); leaves
: intemodes; and dorsally inclined anther pores.
y resembles C. alpina of Colombia. Both species have eglandular tri-
eglandular trichomes be-
n addition these two species have hypanthia cov-
e intercalycine sinuses, and anthers with doisally
dilleradeTalam
becoUeci«i
metaL280
2349 (DUKE
(ianiC.cufodontisii.
242
to Cerro Chirripd, Luteyn & Chaveni 15419 (CAS, NY); Pai
SW of road to La Cima and 4.1 km NW of Cerro Asuncion
Vicinity of Cerro de la Muerte, Aflen 5410 (NY); direct line from Hotel La Georgina to Cerro Frio of the Ceno
with television and radio towers, Davidse 24987 (CAS, MO); Dos Burros, Cerro de la Mueite,
,]imenez2662 (F); enroute to Cerro Chirripd from Canaan and on Pico Sudeste Weston 3652 (CAS); Buena
deLosConejos,tiail
nca, Fosberg 27316 (NY); Cerro de la Mue
the Pan American Highway, Mori 6- Anders
t towards El Empalme, Wilbur & Abneda 17
i. Cordillera de Talainanca, Alme*
11 kmNEofVillaMilb
Burger & Bafeer95I8 (F, MO); neat
Cerro de la Muerte, Panamerican higlwa)'
itsner&Heithaus 241 (MO); Asuncidn (summit of Cerro de la Muerte) 83
' 207 (F); edge of the paramo ca. 2 mi N W from La Asuncidn on the Car-
! (DUKE,F);ca. 2kmSEofUGeorginaand8.3kmSEofUAs^c^
1993.*(FiJ. Nat., B, Adansonia, ser. 4,
Grimm and Almeda, Systematics of Chaetolepis
Erect, moderately to compactly brartched, annual herb 10-40 cm tall. Distal cauline internodes quadrate and
sparsely to moderately covered with appressed, smooth eglandular trichomes 1 mm long. D^tal
with spreading, smooth, eglandulartrichomes 1.5-2 mm long. Stems sparse ycovere asa yw
spreadlg,smLheglandLrtrichomes0.5-1.0mm long. Principal lea^^^
sparsely to moderately covered with minute, smooth eglandular tnchomes that are an rors y p §
»e^dismbu,edove..h=en.iresurface;abaxU.surface^-^^^^^
0).5.er^,j:”U«in.e;.wo,pal.oLcondaryveinsa^m^^^^
— „,wi.hpe«o,esln..lo„g,apart„gb,be..«^^^^^^
Bowers typically solitary but sometimes paired I”™ J 3.„e„ed, basally
lar tnchomes 0.5-0.8 mm long; hypanthia (at maturity) with ^
1-5-2.6 X 0.8-1.2 mm, erect (sometimes spreading). a stalked, bifurcate
«nootheglandulartrichomesnearthebase;mar^nscili^,
onrdurcatetrichome(stipitate-stelU^^^
Pped with a terminal trichome, glabrous, entire. Fda ^ ^-O.^ mm below the thecae.
K linear or somewhat arcuate; pore dorsally incline ’ 4-6 mm long, yellow, glabrous. Seeds
Ovary sparsely covered apically with smooth eglandular tnchome , ly
0-4~0.6 mm long, light brown. Chromosome number: n = 9.
Flowering.
-November through March.
n Senegal, Guinea-
3 wet, boggy areas
Discussion. — Jacques-Felix (1995) transferred Nerophila gentianoides to Chaetolepis based on vegetative,
floral, and seed characteristics. Morphological evidence presented here tentatively confirms his decision de-
spite differences in seeds, habit, and geography. Chaetolepis gentianoides shares many characters with the
neotropical species including 4-merous flowers, yellow petals, ventrally unappendaged isomorphic anthers,
trichome morphology, 4-locular ovaries, and a chromosome number of n = 9 (Favarger 1962; Bolkhovskikh et
al. 1969). Chaetolepis gentianoides also has a consistent nested position in all phylogenetic reconstructions of
the genus in the second analysis using morphological data. It has not been possible to study this species in the
field. DNA sequences will surely provide the decisive data set to evaluate the relationships and generic disposi-
tion of this species.
Chaetolepis gentianoides is distinguished from other congeners by its herbaceous annual habit, stalked
and branching trichomes at the intercalycine lobe sinuses (Fig. 13D), and the convex elongate periclinal ceU
walls of the seed testa (Fig. 6 g-i). All other taxa of Chaetolepis have more or less interdigitating compressed/
flattened periclinal cell walls (Figs. 5 & 6).
Among congeners, Chaetolepis phelpsiae seems most closely related to C. gentianoides based on the strict
consensus tree generated in this study and on their similar morphological characters. They both have dorsally
inclined anther pores, yellow petals, ovate leaves, ciliate-crenulate leaf margins, and smooth eglandular tri-
chomes on the hypanthia, both leaf surfaces, cauline nodes and intemodes.
Chaetolepis phelpsiae differs from C. gentianoides by its perennial woody habit, abaxial leaf surface that is
punctate with eglandular and glandular trichomes, forked trichomes between the calyx lobes- and calyx lobes
with sessile glands intermixed with eglandular and glandular trichomes.
near Dalaba, Adames 401 (li
given 51519 (K);Faoual-Kj
aJacques-Georges 176 (MO); si
28:51. 1871. (Figs. 5 d-f; 15). Hapbdes-
et image!).
Merida. Linden 402 (hootype: PI, photos: F!,
(bulla-based). Distal caultae ntTes *^trtrich”**'''l'u ''T "“ShcMd at base
basall, aud lunget. OUet caubne intentodes but ^ader
tic-ovate, obtuse apically and basaUy (often
and fringed with bulla-based trichomes; petioles W 5 a 0 7 ’ ’ conspicuously rev*
nodes and intemodes. Inflorescence termVnal ™ *“'> ‘rtchomes like those of tif
ered dichasia. Bracts identical to primarv leaves Rrl flowers solitary or in simple 3-flow-
Horal pedicels 2-5 mm long, strigose like distal cauF pnmary leaves but sraaUer.
uune mtemodes. Hypanthia (at anthesis) 2.5-3 x 2-2.5 mm;
<amj«nu ate to subcampanulate, magenta, demely covered with antrotsely, apptessed batbelUte trichomes
hke dMtal cauhne mt^odes. hypanthia (at maturity) with vascular ribs that are inconspicuous when dry; ca-
lyx o s (on tnmure ypanthta) 1-1.5 x 0.8-1 mm, ascending, spreading, or reflexed, lance-deltoid, acuteapi-
fttlM X ^5^ copiouslystrigose; intercalycine lobe sinus with one or several barhellate trichomcs
mao ^ fringed with trichomes. Filaments 3-4 mm long, datlc
the th “"A J * P°« docsally Inclined; connective not prolonged bek>«
maAeuta 5.^76^'“^ “ “-e apex; style 7-^ mm tag
magenta. Seeds 0.6-0.8 mm long, brown. Chromosome number: n = 18.
Flowering. — ^Throughout the year.
several paramos in the D ^ Santander from Paramo de Berlin to Paramo de Mogotocoro and m
mughetwd.richomeson.be\vatyape“;“^^^
axial leafsutfaces hyiA^tTta 7 ‘ntemodes and nodes, ah-
ers from C. Itmlenmtm in having slightly roughened trichomes (vs. densely
Grimm and Almeda, Systematics of Chaetolepis
247
tn El Picacho tc.. ...... .
, . . jyimKOL.
.cgas, ciiirc Dcriiii y Las vegas, Daraay &juajimay 10452 (P); El Picacho, on road between Bucata-
ez-Perez3726 (COL. FMB); Pamplona, La Baja, Punch & Schlim 1313 (P); Pamplona a Bucaramanp,
uunm-Dur figa cv jaiurrtillo 20000 (CAS, GH, P); Pamplona por Garcia, Garganta 990 (US); Pamplona. Kalbreyer 11930Q-,
d. Killip & Smith 8567 (GH, NY); edge of Paramo de Las Vegas, Kiliip &■ Smith 15698 (GH, NY); mountains E of Las Vegas,
1 r„. .. , near Vetas, Kiliip 6- Smitli
Killip&Sm
1), Fdbrega 990 (F); Cucutilla. Vereda
de Vetas v el Alto del Pirachn VHIamizar un tLUL;. «one ae ;>antander:
:e, Barriga 10034 (F, NY); Hoya de Musticua, vci uc
6- Cuatrecasas 10264 (F, NY); Municipio de Cacota, La Laguna, Contreras et al. 43 (HECASA); e
carretera, Cuatrecasas & Rodriguez 27912 (COL, F, G, K. NY, P); Pamplona, por Garcia (sur de la
Carrizal, Piramo El Romeral, Galvdn et al. SYG1025 (COL); E slope of Paramo de Santurban, to
NY); Municipio de Cucutilla, vereda El Carrizal, Paramo de El Romeral en limites con Santander, Sanchez et al 5088 (COL, HECASA); Her-
rtn, Tama, Se«or Orocue, 7‘‘25'31''N, 72”26’38'’W. Mendozaetal 7252 (FMB, HECASA); Municipio de Pamplona, Cerro del Rio Paraplonita,
Sdnchez 6- Solano 4862 (HECASA); Municipio de Chitaga, PSramo del Almorzadero, sector de Presidente, Sanchez et al 10383 (HECASA)
Provincia de Pamplona, paramos de la baja, Schlim 1739 (BR). Unknown location: Jeuxei 850800 (BR). VENEZUELA. Lara: Municipio Mo-
Tu R Los Rosas. Liesner et al 8030 (MO); between Buenos Aires and Paramo
e as Rosas, Steyermark 55483 (F, NY). Mcnda: 6.4 km SE of Laguna Mucubajl off the road to Barinas, King et al 10481 (CAS, F); Sierra Ne-
vada above Menda, Alston 6843 (BM, NY); near Uguna Negra, Paramo de Mucuchies. Alston 6937 (BM. NY); Laguna Mucubaji and Laguna
nstegiaefa2453 (F. NY); Paramo de Mucuchies, distrito Rangel, Aymard & Ortega 1349 (CAS); distrito Miranda, entre Laguna
^ Laguna de Mucubaji, Barclay &Juajibioy 9799
74(F);Laguna Negra. Sierra de Santo Domingo,Dennis2050(K); distrito Rangel. PAramode Santo
Ltorr & Barnett 5182 (CAS, NY); distrito Rangel, la cuenca del Rio Los
a. Dorr & Barnett 5577 (NY, P); distrito Ubertador,
Negra, Gines 1732m)-Tol2Zlonil^^^Z7l^"^"'^ ^aisay, Gehriger37 (GH. F. MO, NY) Lugana
Hanbury-Tracy 1 1 1 (NY); Laguna Negra. Huech!T(°F)- S^m Nev^^^'^"® ^ Loveless 27 (DUKE); Paramo de C^
Lateyn 6098 (CAS, NY); tril lead^rflt Las Escaleras y U Negrito
Mdgde/rau 650 (M); al norte de la carretera M^rBari M ^ Escalera. Luteyn 6167 (CAS, MO, NY); Pico Espq9
P).ElValle.OberwinhlerJ3420(M);Anden,OR '
Y); Paramo de Laguna Grande. Pittier 13245 (F,’g, K, b
3._, ,<.--*xro 127 (MO); distrito Libertador, Sierra Nevada d«. a ■ j ■ , j ■ * i
in2303(DUKE);SanRafael.emreloscaserfrKt pix. , de la Uguna Verde, al pie am
LI Molino y El Portachuelo, Distrito Chacdn, Ruiz-Terdn 3061 (P); al
es. Sierra Nevada de Santo Domingo, distrito Ranoel. Ruiz-Terdn 6356 1
ln7193(BR,F);p
1 Rio Chama, distrito Rar
jdeSanjosial
249
6. Chaetolepis loricarella Triana, Trans Linn. Soc. London 28:51.1871 (Fig. 17). Type: Colombia. Magdalena: Sierra
Nevada de Santa Marta, Purdie s.n. (holotype: BM?; isotype, K! [21).
Erect, much-braitched shrub to 1.2 m tall. Older intemodes woody, terete, exfoliating with age. Distal cauhne
intetiiodes terete and densely covered with appressed, narrowly lanceolate, flattened scales 0.5-0.75 x 0.25-
03 nun that are basally attached with a raised thickening near the point of attachment, margins etose. Distal
cauline nodes like intemodes except scales slightly longer (0.75-1 mm). Principal leaves cormceous, ap-
Ptessed-imbricate; adaxial surface glabrous apically, with a dense covenng of minute appressed scales at t e
base of the blade; abaxial surface covered with two kinds of scales: apical region with narrow, elongate sea es
13-2,5x0.5 mmadnate to the epidermis fortheir entire length, basal regionbeset with flattened erose scales
fcthoseofthenppercaulinein,emodes;blades3-5xl-2mm.ovatetolanceolate,concave,ttgh,lyappressed
wWi the abaxial side of the leaf facing outward and adaxial side facing inward toward the stem axis, apically
sente, basally obtuse with a single depressed vein visible only on the adaxial side; tnargijm seetnmgly enure
but minutely and obscurely ert^ anXor fringed with sessile or subsessile flattened scales with trreguUrly
emsemargins; sessile or with petiolesO.25-0.5 mm long covered withflattened erose sea esa axia y owem
scales like those of the distal intemodes; hypanthia (at matunty) wit n
1-1-5 X 0.3-0.5 mm, narrowly triangular, apically acute, margins
lobes, glabrous toward the apex, persistent. Intercalycine 1<
7-10 X 3-7 mm, magenta, apically obtuse, i
linear-oblong, bilobed ventro-basally and v
'lective prolonged 0.2-0.4 mm below the thecae; p
^les. Style 8-11 mm long, magenta. Seeds 0.7-0.
s usually with a single large scale. Pel
Jiate Filaments 5-6 mm long, magenta; anthers yell
Zgle deflexed lobe 1-2 x 0.25-0.5 mm dorso-basally; c
, pore dorsally inclined, ^ary
Rowering.-May through July. ...nr, is endemic to the Sierra Nevada de Santa M
Habitat and Distribution (Fig 16)— Chaetolepis Imcarellaisende
Colmnbua.»00^“wslvailablecdlec,ionsin^^^^^
250
Journal of the Botanical Research Institute of Texas 7(1|
B A
Discussion. Chaetolepis loricarella does not appear to have any close relatives. It has unusual trichoines
hat are modified into flattened scales (Fig. 17A, D). These are found on the cauline intemodes and nodes, both
kaf surfaces, and hypanthia. In addition, this species has roughened trichomes on the ovary apex, a single
Li r" T concave, appressed, imbricate coriaceous leaves with the abaxial surface
faungomwaniandexposed and .he adaxial surface facing inward toward the stemandco^^^
Representative specimens examined: COUJMRI A , _ .. pello, Village Nabest
IL). Magdalen*
US);NslopeofCuchillaCintura>-
Older branches woodyLrete olahmn. ' ‘ egianauiar inc.
leaves coriaceous somewh t t sparsely covered with moderately barbellate t.
spreading ^ “T" '“f "■“O'"''')' covered wiih antna^
L on anX^e^urT^^'t^'^'^" 01-05 otn. long; abaxjleaf surface densest 0
----- — pnm;
dendritic trichomes 0.1-0.5 .*x*xi ^
terete arms; blades2-6x2-4 mm.
vith short or well-developed axes and a moderate
te, rhombic, or rhombic-orbicular, acute (sometimt
apically, cuneate to attenuate basally, adaxial surface with three nerves that extend for the entire length of the
blade, sometimes 5-nerved with the outer pair arising from the base of the blade and extending one third to
one half the length of the blade before converging with the margin; margins entire, sometimes revolute, cili-
ate; petioles 1-1.5 x 0.5 mm, glabrous. Inflorescence terminal, flowers solitary or borne in a simple 3-flowered
dichasium. Bracteoles 2.5-5 x 2-3.5 mm, ovate, rhombic, elliptic, or obovate, indument of the adaxial surfaces
like that of the principal leaves except for a glabrous basal region, indument of the abaxial surface identical
to principal leaves, 3-nerved, petioles 0.5-1 mm long. Floral pedicels 0.5-1 mm long, moderately to densely
covered with antrorsely spreading, moderately barbellate trichomes. Hypanthia (at anthesis) 2-3 x 2-3 mm,
suburceolate to cylindric, glabrous or sometimes sparsely beset distally with moderately barbellate trichomes
on the vascular ribs; hypanthia (at maturity) with vascular ribs somewhat conspicuous on drying. Calyx lobes
1.5-2 X 1-1.5 mm, erect or ascending, lance-deltoid, apically acute, ciliate, beset with moderately roughened
trichomes adjacent to the midrib but not along the margins; intercalycine lobe sinus with 1-3 smooth eglan-
dular trichomes. Petals 2.5 x 2 mm, yellow, entire, apically obtuse with a single terminal barbellate trichome,
margins glabrous. Filaments 2.5-3.5 mm long, pale brown when dry; anthers 0.2 x 0.4-0.6 mm, yellow, linear;
pore ventrally inclined; connective prolonged 0.2-0.3 mm below the thecae. Ovary apex moderately to densely
covered with smooth eglandular trichomes; style 0.5-0.6 mm long. Seeds 0.4-0.6 mm long, brown. Chromo-
Flowering. — Throughout the year.
Habitat and Distribution (Fig. 19).— In Colombia this species is largely centered in the Cordillera Oriental
(Boyaca, Cundinamarca, Norte de Santander, and Santander) with outliers in the Cordillera Central (Antio-
quia) and Cordillera Occidental (Cauca). In Venezuela it ranges from the state of Merida west to T^chira. It
occurs between 2000-3700 m.
Discussion._Ch<.«olepismttroph>lfatereadllyrecognizedbyi>ssmaUfeafbladesandiuetonga.em^^
atly roughened trichomes on ihecaulineinleraodes. nodes, both foliar surfaces, andmidnbofthe calyx lobes.
Ouretolcpls microphylla is most similar to Chuctolepis suntamarttnsis based on morphologtcal s,m.lant.«
atidtheirpositionson the consensus trees (Fig. 2). These species share entire, ciliate ealmargr^a^^^^^^
ened trichomes on the cauline internodes and nodes, both foUar surfaces, and midnb of the calyx lobes (Fig. 18).
Chaetolepis santamartensis differs from C. microphylla in having slightly roughened tnchomes on the
cauline intemodes and nodes, both foliar surfaces, hypanthium base, intercalycine sinuses and ^lyx lobes,
five (vs. three) elevated veins on the abaxial leaf surface; magenta, ciliate petals (vs. yellow and ealiate); and a
dorsally inclined (vs. ventrally) anther pore.
il.2560(NY):SanPedi
:o, via Arcabuco-Villa dc Leyva, Alonso
il. 3459 (COL); Wramo de La Rusia. NW-N de Duiuma, Chef
,es.e.tu ere La Candelaria, cerros alrededor del Convento, Garcfa-Barriga 20247
hcy„„a«.oae,Os«,Cr-..^.*.COt,K,.2o,3.„d„o^^v^s^^
*I^.SA™i„p„,U„„„„n,amb,dd Hotel
amba de piscinas, Silverstone-Sopfein 5423 (NY); Municipio Samaca, Vereda
'*™talBecfemg&NcgretBN271 (COL). Cundinamarca: Boqueron de Bogota, Andre 736 ( .
& Castaneda 133J (NY); Bogota, Ariste-Joseph s.n. (GH, NY, US);
Cerrode Guadalupe, Barday 4()89 (COL);
Grimm and Almeda, Systematics of Chaetolepis
253
254
Erect, de^ly branched, spreading subshrub up to 40 cm tall. Distal cauline intemodes and nodes quadtatt,
winged beset with sessile glands and appearing minutely furfuraceous when dry. Older steins terete, eiifcliat
bl^r^T 'T' adaxlal and abaxial surfaces covered with sessile gWs
se^Ll ,h I
Flowed 7 " '“"8' ‘■“d often caducous; petioles 0,5-1.5 x 0.25 mm, glabmus
How^ term tml,ortenonUteralbtanches,soUtaty;bracts3.5^.5xO,Ll mm similar to principaltevesl-
«KikT d r f“'*'»':eous when dry. Hypanthla (at anthesis) 3-4 x 3-4 mm, nrceolate, beset wi*
conspicuous vl^„Urr\"'T‘’' when dry; hypanthia (at maturity)
covered with sessileglanili^'ammu^kLlf'
with one smooth, minute eglandular So ^7' ' ““ "®!
obtuse apically and lacking a terminal trichoLe F * r
Ffo>vermg.~November through February
depan“ “^<^3^::“ “ ■>'
and nodes lhaTa^l2rS^^”d7tS^7“ f hy its furfuraceous cauline interno^
abaxially punctate, and entire ^ glands. The leaf blades are oblong with one primary ve
The closest relative of Chaetolepis penjemis var nen •
F pe jemis var. penjensis appears to be C. alpina. These taxa
.both have
Grimm and Almeda, Systematics of Chaetolepis
se«ilp glands on the hypanthium, adaxial leaf surface, cauline internodes, and calyx lobes. Both taxa also
share a punctate abaxial leaf surface, a ciliate leaf margin, smooth eglandular trichomes on the intercalycine
sinuses and ovary apex, and anther pores that are dorsally inclined.
Chaetolepis alpina is unlike C. perijensis var perijensis in having ovate, 3-nerved leaf blades, crenulate leaf
margins, and the elevated primary vein on the abaxial leaf surface beset with sessile glands and smooth eglandu-
lar trichomes. In addition, the cauline nodes have smooth eglandular trichomes and lack glands altogether.
jul 1974, S.S. Tillett & K.W. Honig 747-946 (i
Erect, much branched subshrub to 30 c
minutely furfur,
branches woody.
iO cm tall. Distal cauline intemodes quadrangular, winged, glabrous or
r covered with stalked glandular trichomes and sessile glands. Older
with age. Principal leaves coriaceous and spreading; adaxial leaf surface
-.uuiciy .uriuraceous ana wiui sessde glands when dry; abaxial leaf surface glandular-punctate; blades 4-7.5
X 1-1.5 mm, oblong, obtuse to rounded apically, obtuse or broadly acute basally, the single primary nerve ex-
tending the entire length of the blade; margins obscurely ciliolate-serrulate, the cilia 0.1-0.2 mm long, ap-
pressed and often caducous; petioles 0.3-0.8 x 0.4-0.5 mm, glabrous. Flowers terminal and solitary, often on
lateral branches. Bracts 3-4 x 0.6-1 mm, similar to principal leaves but smaller, petioles 0.5-1 x 0.4-0.5 mm,
glabrous. Floral pedicels 0.6-1.0 mm long. Hypanthia (at anthesis) 3-4 x 2.5-3 mm, urceolate to narrowly
campanulate, moderately to densely covered with sessile glands, appeanng somewhat minute y u uraceous
when dry and moderately covered with glandular trichomes; hypanthia (at maturity) with vascular n s con-
spicuous when dry. Calyx lobes 0.8-1.2 x 0.9-1.1 mm. erect, lance-triangular to deltoid, acute apically, mar-
gins ciliate; calyx lobes glandular and appearing minutely furfuraceous when dry; intercalycine lobe sinus
With a Single eglandular trichome. Petals 4-5 x 2-3 mm, yellow, entire, glabrous, apex acute to obtuse and
lackinga terminal trichome. Filaments 3-4 mm long, yellow; anthers 1.5-2 x 0.25-0.5 mm, linear pore dor^
sally inclined; connective prolonged 0.1-0.2 mm below the thecae. Ovary apex sparely to moderately covered
with smooth eglandular trichomes; style 9-10 mm long, yellow to magenta. Seeds 0.7-1 mm long, brown.
Chromosome number: unknown.
Howering.— July. , in areas with
Habitat and Distribution (Fig. 21).— Known only from the type locality w ere
sparse, low lying vegetation on rocks and cliff bases at 3300-3650 m. , r -, ,
nomic status is possible. This variety differs from the nominate one oy iis yy
and geographic distribution. Chaetolepis perijensis var. glandulosa has ypan
will vancues occur on j ^ m an8 the nominate va
la. but Chaetolepis perijensis var. glandulosa
0-3350 m. Further study of these populatio
«oned differences are consistent and worthy of formal taxououn.
ELA- Amazonas: Ccrro Yavi, elev. 1400 m, 1-3 Mar 1947, K.D. Phelps &
257
»™,^'2brot;XxiaTsuZ
dulartrichomes between the primary and secondary veins, sparsely set wu eganu
*o.«„n.Hep.^.esa„a.conaaHes;Ha<.s,-.5x™
sparsely to moderately covered with eglandulartnchomes.Flowe n=L t v
dichasia.Bracts4.5Jxl3-33mm,liketheprincipalleavesbutsmal^^^^
pedicels 0.25-0.5 mm long, glabrous. Hypanthia (at anthesis) 3-4x2 n
moderately covered with somewhat appressed or spreading, s ascending, narrowly
snK«.h glandular .richomeMmercalycine lobe
f 5_5 5 mm long yellow anthers 4 — 3 x 0.3— 0.5
™..,W,„„ear.„arcua.e.,por=n;:!:::^^^^
mm. Ovary apex moderately covered with smooth eglan
04-0.6 mm long, pale brown. Chromosome number unknown.
lowering.— March. r^rrnYavi in Amazonas state, Venezuela, on rocks
Habitat and Distribution (Fig. 21).— Known only from Cerro ravi
and sandstone cUffs at 1400-1900 m.
Discussion. — Chaetolqris phelpsiae var phelpsiae is a
0 smooth eglandular unbranched trichom
distinguished by its dliate-crenulate leaf margin, punctate abaxial leaf surface, forked trichome on the interca-
lycine sinuses, and calyx lobes with sessile glands, smooth eglandular trichomes and smooth glandular
trichomes.
Chaetolqris phelpsiae var. phelpsiae most closely resembles C. anisandra. These two taxa have anther pores
that are dorsally inclined, and an ovary apex with smooth eglandular trichomes. These species also share ovate
leaves with cuneate bases; leaf margins that are cilate-crenulate; and smooth eglandular trichomes on the dis-
tal hypanthial area, both leaf surfaces, cauline nodes, and internodes.
Chaetolepis anisandra differs from C. phelpsiae var. phelpsiae in having a glabrous hypanthial base; cauline
internodes that are densely covered with a mixture of smooth eglandular and glandular trichomes; and calyx
lobes that are glabrous. It consistently occurs at higher elevations (2100-3400 m) than C. phelpsiae in northern
South America.
. Chaetolepis phelpsiae Gleason vi
Chaetolepis phelpsiae Gleason ssp. chime
Chimanti Massif, frequent along lower
A. Amazonas: Cerro Yavi, Phelps & Hitchcock 6 (NY).
r. chimantensis (Wurdack) Grimm & Ain
a, stat. nov.(Fig.22D2)
)4. Type: VENEZUELA. Boiyaj:
I m, 13 Jan 1953, J.J. Wurdaefe 34108 (houp
Much-branched subshrub ]
iiiiciiiuucs quaaranguiar, sngntiy wingeu.
ed with spreading glandular and eglandular trichomes 0.5-0.7 mm long. Distal cauline nodes 1
internodes quadrangular, slightly winged, sparsely
; 0.5-1 mm long. Older inter-
cept moderately to densely covered with smooth eglandular
nodes terete, woody, glabrous and exfoliating with age. Principal leaves coriaceous, somewhat unequal in size
m each pair, spreading; adaxially sparsely covered with spreading, smooth eglandular trichomes or glabrous;
a axia y punctate between the primary and secondary veins and with a sparse cover of glandular and eglan-
hi* r ^ primary and secondary veins with glandular and eglandular trichomes;
bdes7-16x4-ll mm,ovate,emptic-ovatetoorbicuW^^^
five elevated vemsabaxially (three that extend the entire length of the blade and two that arise at the baseand
extend for one tWrd to one half the length of the blade); margins ciliate-crenulate; petioles 0.5-2.5 mm long.
rimnkT with smooth eglandular or glandular trichomes. Flowers terminal, solitary or borne in a
^ " Hypanthium (at anthesis) 3.5-5 x 2.5-4 mm, urceolate, upper half
lower half elabrous- spreading, smooth eglandular (sometimes glandular) i
S vnaiYt ~7) with vascular ribs promi:
lobes 2-2 5"x 1 1 \ witn vascular ribs prominent and conspicuous when dry. Calyx
■ acute, erec.orspreadin/cita^ bes«wilh«»a<
ruZ«wuh^3' H
ear to arcuate, pore dorsallv i^^ V "" H yellow; anthers 2.5-3 x 0.4-0.5 mm, yeUow, lin*
to moderately covered with smoTth ^Td T ’’“I"”*®* ^ mtn below the thecae. Ovary apex spaM
long, pale brow.r.Chro™so.ne number: ””
Flowenng.— January through March.
Habitat and Distribution (Fig. 21). Chaetolt^^ u i ■
Venezuela on the Chimanta Massif at the upper faU^ of Rio tScT^ ^^*J«antensis is known only I
Sarven-tepui from 1900-2050 m ‘o Tinea and the lowei
while the apical region is sparsely beset w ‘^J^racters. A major portion of the hypanthium is glabrous
w. pbelpsiL has a hypanlum ^dlr T Chaetulepis pl^
var. chimantensis has ovate 5-nerved lp=,f eglandular trichomes. Chaetolepis phelpsi^
e, nerved leaf blades whereas C. phelpsiae var. phelpsiae has lanceolate leaf blades
onlyiromcoa^
t and talus forest of
259
that are 3-nerved. An evaluation of the differences between these varieties will be necessary when more ar
better material becomes available for study.
12. Chaetolepis santamartensis Wurdack, Phytologia 8:165. 1962. (Fig. 23). Type: COLOMBIA. Magdalena: Sierra
Nevada de Santa Marta, SE slopes, Hoya del Rio Donachul, near Col, 3070-3100 m, subparamos, 9 Oct 1959 (fl, fr),J. Cuatrecasas &
R.R. Castaneda 24669 (holotype: US!; botype: COL, internet image!).
Erect, compactly branched shrub up to 2 m tall. Older internodes woody, terete. Distal cauline internodes
quadrate and densely covered with spreading, elongate, slightly roughened trichomes 0. 5-1.0 mm long. Prin-
cipal leaves membranaceous and somewhat unequal in size in each pair; adaxial surface with antrorsely
spreading, minutely roughened trichomes ca. 0.5 mm long evenly distributed over the entire surface; abaxial
surface with erect, slightly roughened trichomes 0.4-0.8 mm long evenly distributed on and between the pri-
mary and secondary veins; blades 10-15 x 3-7 mm, ovate, elliptic to obovate, apex broadly acute to obtuse,
base rounded, 5(-7)-plinerved with the three innermost nerves extending the entire length of the blade and
the outermost pair extending one third to two thirds the length of the blade; margins entire to obscurely cili-
ate-crenulate; petioles 0.5-15 x 0.5 mm, densely covered with erect, slightly roughened trichomes. Flowers
terminal, solitary or in simple 3-flowered dichasia. Bracts and bracteoles 4-6 x 3-4 mm, like principal leaves
but smaller, petioles 0.4-0.6 x 0.2-0.3 mm. Flowers subsessile or on pedicels 1-2 mm long, densely covered
with spreading, slightly roughened trichomes. Hypanthia (at anthesis) 3-4 x 3.5-4 mm, campanulate, moder-
ately covered at the base with erect slightly roughened trichomes, upper two-thirds glabrous; hypanthia (at
maturity) with vascular ribs only faintly visible when dry. Calyx lobes 1.5-2.5 x 2.5-3 mm, erect to spreading
varying to reflexed, broadly deltoid, rounded apically, sparsely to moderately covered with erect slightly rough-
ened trichomes, margins ciliate. Intercalycine lobe sinus with one to three barbellate trichomes. Petals 6-9 x
3-5 mm, magenta, margins ciliate and erose, apically obtuse. Filaments 4-6 mm long, magenta; anthers 2-3 x
0.3-0.8 mm, yellow, hnear-oblong; pore slightly inclined dorsally. Connective not prolonged below thecae but
with a single deflexed thickening dorso-basally. Ovary apex densely covered with slightly roughened tri-
chomes; style 7-9 mm long, magenta, erect. Seeds 1 mm long, brown. Chromosome number: unknown.
Flowering.— October.
Habitat and Distribution (Fig. 21).— This species o
Santa Marta of Colombia in Andean forests and subparamos at 2500-3300 m. .
FHscussion. — Only five collections of Chaetolepis santamartensis are known ut it is stmctive an rea
% recognized. It is a sizable shrub or subshrub with prevailingly 5-plinerved leaf blades an a copious
nient of spreading roughened trichomes on uppermost internodes and on both leaf surfaces^^g.23b^^^^^^^^^
Chaetolepis santamartensis shares some similarities with C. mkrophylla. Bot species
leaf blades, compound dichasia, and roughened trichomes covermg the intern es, n es, „iahrous
Clue^lepJmic^phyiia differs from C.san.<™.-«n.is by its smallovatc
Wial base, and elongate moderately roughened to dendritic riiAomes covermg
leaves. Chuetolepis micmphylla has anthers with ventrally inclined (vs. ^
leaves that have thr« veins thatextendfmmthebaseolthe blade to its apex, and yellowCvs. magenta) echate
petals.
Vesenutive specimens examined: COLOMBIA. Cesar: Sierra Nevada de Santa Marta, Valledupa^ jwramo entre y
«l«,Cart»s3657(UTMC).M.(Ut.l™.:SiernN,™hdeSanoM.rU,.rikddR»^
^ Sanu Marta transecto dclAlto Rio Buritaca, Jaramillo et al. 5434 (COL); Sierra Nevada de Sanu M
^(COL).
»■ Chaetolepis sessilis Pittier.J. Wash. Acad. Sci. 13:385.
cagua, 3200 m, 31 Mar 1922, A. John 1037 (holotype: VEN, internet image..
the southeastern slopes of the Sien
Journal of the Botanical Research Institute of Tews 7(1)
with smooth, eglandular, antrorsplv
woody, terete and furrowed with agl PrinrinJr trichomes 1-1.5 mm long. Older inteniodes
brous; abaxial foliar surface glabrous on th ^ conaceous and spreading; adaxial foliar surface gla-
5-10 X 1.5-4.5 mm ellintir to nhlr. ii ^ and secondary veins and punctate between them; blades
2.5- 3 5 X 1-1 3 mm, narrowly elliptic to ’ I k "■"cl' like principal leaves. Bracleole
late between the primary and secondary vX lli ™' “"''^Pieally and hasally, glabrous abaxially.punc-
petioles 0.3-0.5 mm long Horal pedicTlsHr^ f T
4.5- 8 X 2.5-35 mm, nmeolate, m^ratelycL^d^ HyP“"'''ia
With appressed, eglandular trichomes- hvDamliia sparsely to moderately covered
lyx lobes 1.3-2.5x 0.8-1.2mm^Z-del^^^''^n"^^^^
ate; calyx lobes moderately covered with spreading or reflexed, cili-
Wlih bifnrcaie or iPfmcaie iPchomes. PeStfxT,^''
cuspidate with a terminal trichome. Filaments "^^^gins entire, glabrous, apically
arcuate; pore ventrally inclined; connective prolon T
apically with appressed smooth eglandular tri h ® tnm below the thecae. Ovary sparsely covered
brown. Chromosome number; unknown. "" Seeds 0.4-0.6 mm long,
Howering— March to June.
Habitat and Distribution (Fig. 21) Vn
paramos at 2800-3300 m. Mtrida, Venezuela, in Don Pedro and Aricagua
this species is readily d^istinguished°Ca°™!!^“'^h""“''“"“'™“*'''”a“‘iy Despite
axial leaf surface between the primary and secotL^Tv!!!^^' eglandular trichomes on the ab-
ends on the cauhne intemodes, calyx lobes, and hZanrh^’ hypanthia. It also has sessile
bifurcate or trifurcate trichome at each intercalvl’^ ’^'^^'^^^^b^^^^
The closest relative o( Chaetokpis sessilis appearl^L^c^ ^ '^^otrally inclined anther pore (Fig. 24).
tobeCperyensisvar.peryensis. Both species have ses-
Grimm and Almeda, Systematics of Chaetolepis
sile glands on the cauline internodes, calyx lobes, and hypanthla. They also share smooth egtodular tn-
chomes on the abaxial leaf surface between the primary and secondary veins and on the cauline nodes.
aa«nlepispen/ensisvar,pcrijensisisunlikeC.se»ilisinhavinglealbladeswithasinglepnmatyvem,.ts
leafmargins are Lte and enthe; the adaxUlleafsurfacesandcauUne nodes ate covered with ^ssile glands,
and the abaxial leaf surface is punctate. Furthermore, the hypanthium is glabrous, t e mterca ycme smuses
have smooth eglandular trichomes, and the anther pore is dorsally inclined,
active speci„.u.-.m,«.d:VWEZt,E.h.M«daiPa»«odeIXH.P«h..n™20-n
distrito Libenador, Ruiz-Terdn & Ldpez-Figueiras 8658a (F, US).
Chaetolepis cubensis (A. Rich.) Triana, Trans. Linn. Soc. Lo
to Sagra, Hist. Fis. Cuba, Bot. 10:550. 1845. Type: CUBA: de la Sc
cubensis (A. Rich.) Griseb., Cat. PI. Cub. 103. 1866. = Tiiwuchina ci
Chaetogastra cubensis var. brevistrigillosa Griseb., Cat. PI. '
JR!, G!, GH, MO!). Choetogistra
t: Savana del Ciego, Wright 2529
DC.,Monogr. Phan. 7:172. 1891.
Chaetogastra origanoides Griseb., Cat. Pi. Cub. 103. 186
Cte4;:;r:,q:LGriseb.Cai.Pl.Cub. 103.
BR!,G!, MO!, P!). Chaetolepis saturejoides(Gnseb.)Tnana,
Pleroma squamatum C. Wright, Anal. Acad. Ci. Habana 6:73. 1
(^ffoetolepis grS^cl^Cogn., in A.DC. & C.DC., Monogr. Phan
tl. 7:172. 1891. Type: «
We thank Peter W. Fritsch and Robert W. Patterson for early reviews of the manuscript and for patient help
with all the cladistic analyses. Two anonymous reviewers are thanked for critical and constructive reviews. We
are also grateful to Alan Chou for the line drawings, Charlotte Pfeiffer for technical assistance with many of the
figures, Scott Serata for assistance in the Scanning Electron Microscope Laboratory at CAS, Sabina King for
editorial assistance with many iterations of the manuscript, Carlos Parra and Eduino Carbono for facilitating
loans of important specimens from Colombia, and the curators and staffs of the following herbaria for loans or
assistance during study visits: BM, BR, CAS, COL, DS, DUKE, F, FMB, G, GH, HECASA, K, M, MO, NY, P,US,
and UTMC. We also thank the Andrew W. Mellon Foundation and JSTOR Plant Science for supporting and
facilitating the creation of a global online image resource for vascular plant type specimens. The facilities and
logistical support of San Francisco State University and the California Academy of Sciences is also gratefully
acknowledged.
REFERENCES
Almeda, F. 1997a. Chromosomal observations on the Alzateaceae (Myrtales). Ann. Missouri Bot. Card. 84:305-308.
Almeo^ F. 1997b. Chromosome numbers and their evolutionary significance in some neotropical and paleotropical
Melastomataceae. BioLlania Edicidn Esp. No. 6:167-190.
Almeda, F. 2001 . Chaetolepis. In: J.A. Steyermark, RE. Berry, K. Yatskievych, and B.K. Holst, eds. Flora of the Venezuelan
Guayana. Missouri Botanical Garden, St. Louis. 6:294-295.
Almeda, F. 2013. Systematic and phylogenetic significance of chromosome number diversity in some neotropical
Melastomataceae. Mem. New York Bot. Garden 1 08:1 55-1 77.
Melastomataceae. Syst. Bot. 17:583-593. ^ systematic significance in some Mexican
93':U1oT'^ ° '^ob'nson. 2011. Systematics and phytogeny of Siphanthera (Melastomataceae). Syst. Bot. Monogr.
^^rrinq Lnu VLr'''' 'Melastomataceae. In: A.A Federov, ed. Chromosome numbers
7.|_1 256. ° eds. Monographiae phanerogamarum. G. Masson, Pans.
and their systematic significance.Tmo^2 ”^1 ^naatia and Duabanga (Lythraceaei
MIjnbouwkd. 40:18t-188. Tertiary stratigraphy and tectogenests of the Colombian Andes. Ge
Irving, E. 1975. Structural evolution of the northernmost ah
United States Government Printing Office, Washington DC Professional Paper 846:H
Jaccjues-Feux, H. 1994 [1995]. Histoire des Melastom^r '
16{2-4):235-311. ® dAfrique. Bull. Mus. Hist. Nat., ser. 4, sect. B, Adansor
700-756. Myrtales and Myrtaceae - a phylogenetic analysis. Ann. Missouri Bot. Gard. i
LiNcoor, F.C, S.R. Pnirsort'^t
16:, -1,3. -"igradon of blrCs. United States Fish L Wildlife Ser^te,
lition, version 4.0PPC.
> Maddison. 2000. MacClade: a
ceae americaines d^rites par C. Naudin. J. Bot. Soc. Bot. France
Marcetia (Melastomataceae). Dissertagao de Doutorado, Universi-
37:3-111.
M/miNS, A.B. 1989. Revisao taxonomica d
dade Estadual de Campinas, Campinas, Brazil.
Micheiangeli, F.A., PJ. GuimarAes, D.S. Penneys, F. Almeda, and R. Kriebel. 2013. Phylogenetic relationships and distribution of
New World Melastomeae (Melastomataceae). Bot. J. Linn. Soc 171:38-60.
Miquel,F. 1840. Choefo/ep/s. In: Commentarii Phytographici. Leyden. P. 72.
classification of the Melastomataceae. Scientific Abstracts no. 509. Botany 2010, July 31-August 4, Providence,
Rhode Island.
RANca-CH., J.O. AND A. GarzOn-C. 1997. Sierra Nevada de Santa Marta, Colombia. In: Centres of plant diversity: A guide
and strategy for their conservation (Volume 3, The Americas), S.D. Davis, V.H. Heywood, 0. Herrera-MacBryde, J. Villa-
Lobos, and A
Publications Unit, Cambridge, U.K. Pp. 426-430.
Raven, P.H. 1975. The bases of angiosperm phylogeny: Cytology. A
Renner, S.S. and K. Meyer. 2001 . Melastomeae come full circle: E
Evolution 55:1315-1324.
Simpson, B.B. 1975. Pleistocene changes in the flora of the hiah tropical Andes. Paleobiology 1:273-294.
I.Wurdack.1980.C
ri Bot. Gard. 62:724-764.
Sou, M.L AND JJ. Wurdack. 1 980. Chromosome numbers in the Melastomataceae. Phytologia 47:1 99-220.
Standley, P.C. 1938. Melastomataceae. In: Flora of Costa Rica. Field Mus. Nat. Hist., Bot. Ser. 18:783-845.
SwoFFORD, D.L 2002. PAUP*:Phylogenetic analvcic usino oarsimonv (and other Methods). Ver. 4.0b10. Sini
httpV/sweetgum.nybg.org./ih/
J. Meggers, E.S. Ayensu, and
Thiers, B.M. 2012. [continuou:
New York Botanical Garden's Virtual Herbari
Thorne, R.F. 1973. Floristic
W.D. Duckworth, eds. Tropical forest ecosystems in Africa
5. Bot. Rev. 58:225-348.
rc« jR.'r92TT?;e b^rdronh^ Marta Region of Colombia: A study in altitudinal distribu-
tion. Ann. Carnegie Mus. 1 4:3-576.
Toozia, C. A. AND F. Almeda. 1991
Acad. Sci.47(6):1 75-206.
Tbana, J. 1871. Les M4lastomacees. Trans. Linn. Soc. London 28:1-11
U"iBE-U., L. 1972. Catalogo ilustra
Melastomataceae. Publ. Inst. Ciencias N
Wurdack, JJ. 1973. Melastomataceae. In: T. Lasser, ed. I
8:1-819.
isifloraceae, Begoniaceae,
racas, Venezuela.
BOOK REVIEW
Arthur Haines, with illus. by Euzabeth Farnsworth and Gordon Morrison. 2011. 1
Society’s FLORA NOVAE ANGLIAE: A Manual for the Identification of Native and Naturalized
Higher Vascular Plants of New England. (ISBN: 978-0-300-17154-9, cloth). Yale University Press, P.O.
Box 209040, New Haven, Connecticut 06520-9040, US. A. (Orders: www.yalebooks.com, 1-203-432-
0163, 1-203-432-8485 fax). $85.00, 1008 pp., 945 b/w illus., 7" x 9 ‘A".
The stales of Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont are usually
considered the New England area and they are the focus of this much needed, new and updated study of their
flora. In 2001 the New England Wild Flower Society hired Haines to visit every herbarium in the area and
check out every rare plant specimen to determine if it still existed in the area, if it had been properly identified,
and correctly named. His two-year study of the specimens indicated that one of every twelve specimens re-
1 to correct a misidentification or to identify a new species or infraspecific taxon that had
it had been collected.” Although a couple of new floras for the area had been published previ-
ously, they primarily relied on M.L. Femald’s 1950 edition (the 8th) of Gray’s Manual of Botany. This new vol-
ume is based on the outstanding work of current and recently deceased botanists, and it is a treasure-house of
excellent research, field studies, and dedication to botany.
Following a very helpful Introduction and an updated Glossary of Terminology, is the Key to the Fami-
lies, pp. 1-37. The heart of the book, though, is the whole set of taxonomic treatments (pp. 39-890). These aie
divided into the subgroups Lycophytes, Monilophytes, Gymnosperms Maenoliids Monocots, and
Tricolpates. r . &
most Mcated”!nd ^r^'^d ^ Cited section. This is an amazing citing of the works of some of the
this section. You will undoubtedly recognize name after name of botanists that you know or have known in the
recent past— or perhaps, you simply recognize through your reading— and whose research and contributions
have made a huge contribution to understanding better the flora of New England.
T contribution. The New England Wild Flower Society has provided a great
Texas USA Research Institute of
MICONIA PHRYNOSOMADERMA (MELASTOMATACEAE: MICONIEAE),
A NEW SPECIES FROM THE MASSIF DU NORD, HAITI,
AND SIXTEEN NEW NAMES AND COMBINATIONS
Lucas C. Majure and Walter S. Judd
Department of Biology
University of Florida
Gainesville, Florida 32611-8525, USA
lmajure@ufl.edu, vi/judd@botany.ufl.edu
RESUMEN
INTRODUCTION
St Erik L Ekman was a voracious collector and incredible taxonomist. He has
wild animal in the way he collected plants, as he made easy work of terrain quite
He was known for his hroad knowledge of the flora of Cuba and Hispaniola his
-tougnness, beautiful collections, and also his indifference to advetse climatic conditions while in the held
(Sandley 1931; Wolcott 1931). As an avid collector and well-trained hounist, he discovered hundreds rf new
snppipc P' U J n, 1 f ivfcvU fipsrribed either by German botanist, Ignaz Urban,
of die numerous collections made hy Ekman, many are still yielding species new to ^tence feg , Judd Sr
1988;Judde.al. 1988;Judd 1994;Guerremetal. 2004;Judd Sr Majure 2013). ^
oftspmsentedinthis paper is the resuhofacollectionofarareMiconin Ruiz SrPav.lrom northern Hammade
As currently circumscrihed, the genus Miconin of tribe Micomeae, is extremely poh^yletic, ®
■“oos other genera, e.g., Cniycogonium DC., Clidemia D. Don, Conostegin D. Don, ^
Inst Tew?
genera in the group to be arbitrary. It is quite clear that generic circumscription within this group is in need of
revision, however, the splitting of Miconia would lead to a proliferation of new genera, many of which would be
morphologically undiagnosable and of problematic circumscription (see lonta et al. 2012). In contrast, the
Miconieae can be diagnosed by several morphological synapomorphies: partly or entirely inferior ovaries, bac-
cate fruits, stamens without or merely with poorly developed connective appendages, and the absence of mega-
styloids (Michelangeli et al. 2008). Therefore, it is more useful to recognize a broadly circumscribed Miconia,
including all those species of Miconieae with berry fruits, while recognizing well-supported, less inclusive
clades with clear morphological synapomorphies as suhgenera or sections of Miconia (lonta et al. 2012; lonta
&Judd2012).
The Greater Antillean Lima clade within Miconia is a good example of the problem of arbitrarily circum-
scribed traditional genera within Miconieae, as it contains species that have been recogni:
Clidemia, Leandra, Miconia, Ossaea, Oxymeris DC., and Sagraea DC., although most species within tl
phyletic group are currently recognized within either Leandra or Ossaea (Alain 1957; Judd & Skean 1991; Uo-
gier 2000; Michelangeli & Becquer 2012). Leandra and Ossaea were traditionally recognized based on their
acute petal apices, a feature exhibited by most members of the Lima clade. Leandra is widely polyphyletic, al-
though, Leandra s.str. (including the type L. melastomoides Raddi; Martin et al. 2008) is a clade quite distantly
related to the Lima clade. The nomenclatural status of Ossaea is problematic, as no type has been designated.
However, of the species, which could be used as the type of the genus, none of those are closely related to the
included in Miconia s.l., neither Leandra nor
correct genera for members of the Lima clade. Thus, we herein describe this clade as a new
nia, i.e., Miconia sect. Lima. For the above stated reasons, we justify the description of the
Lima clade. So e
Members of Miconia sect. Lima represent a clade that is restricted to the Greater Antilles and consists of 17
knovm species, including Miconia phrynosomaderma, which is described herein. The monophyly of sect. Lima
IS well supported m molecular phylogenetic analyses (Goldenberg et al. 2008; Martin et al. 2008; MichelangeU
et al. 2008; MichelangeU et al, in prep.) and by morphological characters. Putative synapomorphies of this
clade are numerous, including the striking and well-developed bulla-based hairs on the adaxial leaf surface
(Fig. 1), as well as the ephemeral, long-stemmed, clavate-dendritic hairs on the adaxial leaf surface of develop-
mg leaves (which are produced from in between the large bulla-based hairs towards the leaf base along pri-
mary, secon^ry, and occasionaUy tertiary veins). These clavate-dendritic hairs are also produced at the *
mnfr ‘he bulla-based hairs. Bulla-based hairs are also produced on other plant
the ^ “^orescence axes, hypanthia, calyx teeth, and the upper surface (rf
Sessile T d * L 'TI'T' **»=* ‘" species, as compared to those of the adaxialleaf surfice.
taltrme^i "i'h ^cute to acumitute apices that bear large bulla-based hairs
apomorphic (HeT) pr II T*** p' *ese dUtinctive hairs likely is also sjn-
pore, both likelyapomor^hL ' ™ appendage and one dorsally oriented
dy re"«;mm »• distinctive, both are
scured by the dense bulla-based h
J typically slightly four- or five-lobed, although this if often ol
Evergreen shrubs; young stems terete, elliptic or slightly rectangular in cross section, lacking longitudinal
ridges, the indumentum of dense bulla-based hairs, these long appressed, spreading, or recurved, or short and
granulate. Leaves opposite, slightly anisophyllous; blade elliptical, ovate, or narrowly ovate, the margin crenu-
late to dentate, these crenulations/dentations obscured by large bulla-based hairs, which slightly fold over the
leaf margin, producing in some cases a moderately revolute margin, the indumentum of adaxial leaf surface
typically of broad bulla-based hairs ± filling areoles, although sometimes these hairs relatively narrow and
wide-spaced, not filling the areoles, with long-stemmed, clavate-dentritic hairs produced along the primary,
secondary and tertiary veins from between the bulla-based hairs, and also sessile to short-stalked glandular
hairs present on all parts of the lamina (between bulla-based hairs), the abaxial leaf surface variously covered
by narrow bulla-based hairs, these either long and well developed or short and granulate, these appressed,
spreading, or erect, the lamina with sparse, sessile glands, the venation acrodromous, with secondary veins
arching toward leaf apex, 1 to 3 pairs, basal to suprabasal, tertiary veins percurrent, ± perpendicular to the
midvein, sometimes mostly obscured by bulla-based hairs on the adaxial leaf surface, connected by quaternary
veins, the primary, secondary, tertiary and quaternary veins mostly impressed on the adaxial surface and
raised on the abaxial surface, domatia present or absent, occurring at the junctions of primary, secondary and
tertiary veins, forming a pocket-like structure in the axils of the primary and innermost secondary veins or
formed from a tuft of hairs in the vein axils. Inflorescences terminal, although often surpassed by the rapid
growth of axillary shoots, the flowers in 3-flowered dichasia, sessile, subsessile or pedicellate, thus forming
open cymes orsessileand nearly headhke clusters. Howers 4-5(6) merous, mostly actinomorphicor nearly so;
hypanthium 4-5 lobed, the lobes sometimes obscured by retrorse or antrorse bulla-based hairs, bu la-based
hairs long and well developed, or granulate, hypanthium also with sessile glands, calyx o s tnangu ar, acme
to acuminate, often covered by sessile glands throughout the adaxial surface or such glands restricte to t e
apex of the adaxial surface, ahaxial surface covered In hulla-based hairs and ± sessile glanrb; calyx teeth ±
eqmdo, longer thancalyxlohes, terete, mostly reilexed in fruit, coveredinlougandn-elld^eoped, or granu-
Ittt. bulla-hased hairs, sessile glands present or absent; calyx tube often with long stemmed^ clarmte jendrrue
hairs produced from apex along the margin, sessile glands ± present on adaxia sur ace, a axia su ace cov
ttediubulla-lased hairs; petals ovate to olxwateorslightly oblong, symrnetne or as^rnetnc,wh,te,red,rose
P^le, or white with pu^le tinge abaxially, apices acute to acuminate, with nrodenrtely bu la-tesed tar^
Ptoduced from the abaL7surlaces just below the petal apex and occasionally from the rmdral jmrtron o^e
M as well; stamens W0(-12), n^t geniculate, the filaments glabrous, the ambers ™h or
tWbasal appendage and a single, dorsally inclined pore; style straight to moderately curved, ^ '
iu ^middle, the stigma punctiform; ovary 2-5 locular, e inferior, wuh ^
intruded into each loculthLvary apex withoutacoUar but corrunonlywnthacrowno^m^^^^^^
P«arance to that of a file or rasp. Hence, we found it appropnate to use e n
^konia. , other
The following new combinations or new names for species . Although they are not part
Miconia maec{uipetiolata, M. karlkrugii and M. krugiam, are ^ ^ phylogenetic data
'rf‘heUmaclade(Michelangelietal.,unpubl.data,Majureetal.inprep.),morphologi P
^’^Sgest that they are closely related.
268
Journal of the Botanical Research Institute of Texas 7|1)
Miconia asperifolia (Naudin) Majure & Judd, comb. nov. Clidemia a
Note that Clidemia hirsuta Macfad., FI. Jamaica 2:45. 1850., also represents this species, but it is not considered
here to be effectively published, and thus is not validly published. The name Clidemia hirsuta (Sw.) Griseb. (Fl.
Brit. W. 1. 248. 1860.) represents a different species.
Miconia cubacinerea Majure & Judd, nom. nov. Clidemia cinereaGhsth, Cat. Pl. Cub. 97. 1866. Ojtymeris dnerea (Griseb.)
de Baracoa, 1 1 Jun 1860-1864, C. Wright 2483 (holotype: GOET! online image seen GOET007034; isotypes: BM! online image seen
BM000884493. BR! online image seen BR0000005185191, GH!, K! online image seen K000535607 MO! YU! online image seen
YU065014).
The specific epithet, cubacinerea, refers to the restricted distribution of this species, which is only known from
Miconia cubana (Alain) Majure &Judd, comb. nov. Ossaea,
1955. Type: CUBA: Isabel Maria, 16 Mar 1860-1864, C. Wright If
(UTb.)Alam,S.da 18:1026.1999. Type: HAm: Massif desCahos, Petite-Riviere delArtibonite,P^rc>din,atlngram,7Mar 1925, EL
'"HJ440 (holotype: S!;
1. Massif de la Selle, Croix-des-Bou-
;: K! online image seen 000329545,
Miconiajasbaferi Majure & ludd nom nnv l r „
The new epithet, like the original one, honors John Adolph Shafer (1863-1918).
Miconia karlkrugii Majure &r Judd, nom. nov. Col i,
Judd &Skean, Bull. Honda State MuL. Biol Sci 36-6/13"^”' Syst. 4:279. 1886. Leandra hrugii <
PUERTO RICO Maricao in decliv b ^ fCogn.) F.S. Axelrod, Sida, Bot. Misc 34:214. 201
The specific epithet kadkrugii honors the botanist Karl W
/ilhelm Leopold Krug (1833-1898) for which the spC'
I 111 I
image seen CORD00003622, FI, GOET! <
M0165768,MO!,NY!,US!).
Miconia limoides (Urb.) Majure &Judd, comb. nov. OssaealimoidesV^
mountain Laubwald, eruptives, 1300 m, 28 Jan 1926, E.L. Efeman H5462 («
_„_ _-ently recognized as 0. turquinensis, the specific epithet turquinens
a Oudd 2007). Therefore, the synonym 0. norlindii is transferred to Micoma.
Miconia ottoschmidtii (Urb.) Majure & Judd, (
■t.Spec.Nov.RegniVeg.2-
Cer«lZringo,LomalaCamranZrNovl929.E.L.EfemanHl]522(HOLOiin>E:S!;«HY^
The name Ossaea urbaniana was proposed to differentiate Ossuea polychaeta from O. polychaete Urb. & Ekman,
Ark.Bot.22A:60.1929.Leandraurbanmna(Alain)Alain,Sida20:1645.2003,nom.illeg.,isalaterh^^^^
urlmianaCogn.,inMart.,RBras.l4:148.1886.Anewnameisneededfor0.poIyc^^^^^^^^^
niana Cogn. (Bot. Jahrb. Syst. 42:139. 1908) and M. polychaeta Wurdack (Phytologia 23. a rea y exis
in this genus. The new epithetrefers to the condensed and distinctly pedunculate inflorescence ot this species.
a Majure & Judd, nom. nov. OssaeacapitataVrb., Rep.
The long calyx teeth of this species resemble
which also serves to mnemonically link the ne
-e & Judd, sp. nov. (F^. D- ^
^ section not ridged, the intemodes 1-3.2 cm long;
shrub (height unknown); steins roun m strongly dilated
^tn indumentum of bulla-based hairs 0.4-1. 2 mm long t ^ or slightly spread-
^ and others only narrowly dilated at the base, the P ^ ^ ^ 2 mm long. Leaves
-gwith apices recurved; nodal line present, made up of tnang«
^e.pa.,2.4^.3.«c.of.e„sU^^rf™
Majure and Judd, A new species of Miconia and new combinations 27 '
mm above the leaf base, positioned 2.5-53 mm in from margin at widest point of blade, the tertiary veins ±
1, 2.4-3.5 mm apart at mid-leaf; adaxial surface covered in bulla-based hairs, these
5 the lamina visible between the hairs, i.e., lamina areoles are not completely filled,
.n wide, apices of bulla-based hairs mostly erect to slightly spreading, the young
leaf adaxial surface with ephemeral, long-stemmed, clavate-dentritic hairs, these sometimes flattened at the
apex, arising from between the bases of bulla-based hairs along the primary and secondary veins toward the
base of the leaf, and with subsessile to short stalked glandular hairs along the lamina between bulla-based
hairs; abaxial leaf surface covered with bulla-based hairs, although the lamina clearly visible, also with bulla-
based hairs covering the primary, secondary, tertiary, and quaternary veins, the lamina covered in sessile
glands, also with depressions formed from the bulla-based hairs on the adaxial leaf surface; petiole 0.4-1.2
cm long, covered in bulla-based hairs, these spreading to retrorse and recurved on adaxial surface and mostly
appressed-retrorse on abaxial surface. Inflorescences terminal, well-developed to reduced cymes of 3-15
flowers, 1.7-3.9 cm long, 2.2-5.1 cm across, the peduncle 0.1-0.7 cm long, the proximal branches 0.7-1. 7 cm
long, the pseudopedicels 1.5-3.5 mm long, and pedicels 0.6-1 cm long; bracts narrowly ovate, 2-3 mm long
bmeoles narrowly ovale, persistenl, 2-2.2 mm long, ca. 0.2 mm wide; nodes of innorescence wnh mixed
bulla-based hairs and long-slemmed,denlrilic-clavale hairs, similar lolhose found allhe^ofyoungkaves.
Fluwersd-merous; hypanlhmm3.1-4 mm long, 5-5.2 mm wide,*spherical,shghlV4-loW,ahhough low
mostly obscured by bulla-based hairs 0.9-2.5 mm long, the free portion of hypant iuin . ® 8’ 8 Y
«helowlhelorns,holhhypanlhinmahaxUlsu^^^^^
ally andhnlla-lxised hairs abaxially; calyx lube0.4mmbng;pelals while, Imlpurpfeh on Aeataxialsn^,
o«e,5.1-5.2xca.3mm,lheapexacuminale,lhe margin membranous and enllre,cawedal base, wnh l^
slightly bulla-based hairs just below the apex on the abaxial surface, these 2 3 mm “
number of petals, the ftlaments 1.7-1.9 mm long, the anthers 1.4-1.5 mm long,
and asingle, dorsally inclined pore, the thecae 1.1 mm long. Style ca. 4.3 mm tag,
.-.estigma,suhtendedhyacrownoftag,multtalutahm.t^^^^
bulla-based hairs on the apex of the ovary; ovary ca. „ 6.5 mm wide,
placentation axile, placenta greatly intruded into the locules. bem g
blue-black at maturity. Seeds (immature) ca. 0.9 mm long, sickle-shaped.
Artibonite Province, Haiti (Fig. 2). It was
[ notes mentioned that Mome Belle Terre w
ommunity at the elevation where M. ph
sif du Nord, at Mome Belle Terre,
(Efeman H8204). Ekman in his fielc
but no information regarding plai
Phenology.— Miconia phrynosomaderrna was in flowering period is essentially
22nd), so it is likely that the species begins flowering earlier in the ye ,
Etymology.— Miconia phrynosomaderrna is named for the and
epidermis of Phrynosoma is covered in scales, which eve op m , cranium also produce
I’f Ac genus are commonly referred to “ ^h(^all^^^ "f M phiyosamad^rma
l^asrtatLImymAtaorrLd^^^^^^^
based on morphological characters, Miconia phrynosomade^ juc snarse indumentii
'bnoides and M. lima. Miconia phrynosomaderrna differs from - mo ^ ^
^lalleafsurfaceandthusclearlyvisibleabaxial epidermis, do not completely
bi® (Fig. ID-E), and by the spacing of the ad
®^eoles (Fig. IB-C). In M. limoides the bulla-l
er the leaf
abaxial lamina and completely
fill the areoles. Stem hairs ofM.hmoides are mostly ^ i
M. phrynosomaderma they are mostly appressedr«rL!^? ! tT°''"^^^
of theinflorescencesofM Laides tend tobe^lr T t ^
those of M. phrynosomaderma which are more o sessile to subsessik
needs to be investigated further (with moni collecttonsXhTt ‘“T"'
derma are longer (i.e., 3.5^.4 ntot) than those of r , a ’ '
exhibitsclawedpetalsandadorso-hasalaoDendaeeonT* «■ pliry"n»«“
Ucks clawed petals and has a reduced do^-lJLmte a * 1
aorso nasal anther appendage to 0.1 mm long or it is entirely al
MajureandJud(l,Anews
appressed-antrorse hairs (in M. lima), hypanthia 3.1-4 x 5 mm vs. 1.9-3 x 2.3-3.5 mm, and widely elliptical
(length to width quotient 0.92-1.95) vs. narrowly elliptical (length to width quotient 1.56-2.05) leaves. Mico-
niflphjynosomaderma differs from both M. lima and M. limoides by the purplish color of young leaves vs. the
lime-green color in M. lima and M. limoides, as well as the color of the petals, that is, white with purple abaxial
surfaces in M. phrynosomaderma vs. usually rose to red in M. lima and M. limoides. Miconia limoides may some-
times have white petals, but these are not purple abaxially. Also, both M. limoides and M. lima usually have an
androecial fringe on the apex of the free portion of the hypanthium, which is a continuation of the hairs pro-
duced on the free portion of the hypanthium, but this structure is absent in M. prhynosomaderma.
Miconiaphrynosomaderma satisfies the morphological-phenetic Qudd 2007) and diagnostic species con-
cepts (Wheeler & Platnick 2000), as it is easily distinguished, morphologically, from its putative closest rela-
tives, M. limoides and M. lima. Miconia phrynosomaderma also is allopatric from other members of the Lima
clade, and so is most likely reproductively isolated from other species, thus satisfying the biological species
concept (Mayr 1970, 2000). Finally, the clawed petals are likely autapomorphic, and thus M. phymosomaderma
is most probably a cladospecies (Donoghue 1985; Mishler 1985).
ACKNOWLEDGMENTS
This research was supported in part, by the National Science Foundation Grant BSR-0818399. We thank Nor-
ris Williams, Kent Perkins and Trudy Lindler, University of Florida Herbarium (FLAS) for their help in pro-
cessing specimen loans. We also thank the curators and staff of the herbaria (GH, S, and US) who generously
provided specimens on loan and L. Trujillo for introducing L.C. Majure to the genus Phrynosoma. We thank
Fabian Michelangeli and an anonymous reviewer for providing helpful comments on an earlier version of this
manuscript.
REFERENCES
Auin, Hno. 1 957. Flora de Cuba, Vol. 4. Contr. Ocas. Mus. Hist. Nat. Colegio “De La '
BfcQUEH Granados, E.R., K.M. Neubig, W.S. Judd, F. Michelangeli, J.R. Abbott, and D.S. Penneys. 2 . re ii
logenetic studies in Pachyanthus (Miconieae, Melastomataceae)^BoL Rev. 74:37-5 .
CoGNiAux, A. 1891. Melastoma -
Masson, Paris. Pp. 1-1256.
f^»«KHUE, MJ. 1985. A critique of the biological
Bryologist 88:1 72-1 81.
^««0N, HA 1932. A synopsis of the Melastomataceae of British Guiana
^^o^knberg, R., D.S. Penneys, F. Almeda, W.S. Judd, and F.A. Michelangeu. 2008
terns of stamen diversification in a megadiverse Neotropial ger’ '
^RRERo, A., W.S. Judd, and A.B. Morris. 200
de la Hotte, Haiti. Brittonia 56:346-352.
Ionta, G.M., W.S. Judd, J.D. Skean, Jr., and C.K. McMullen. 201 2.Two nev
from the Macaya Biosphere Reserve, Haiti, and twelve relevant
■owA, G.M. and W.S. Judd. 2012. Miconia cordieri,
Macaya Biosphere Reserve, Haiti. J. Bot. Res. Inst. Texas 6:37-44.
fr®o.W5. 1986. Taxonomic studies in the Miconi
38:150-161.
^.W.S. 1989. Taxonomic studies in th
Ann. Missouri Bot. Gard. 76:476-495.
W5. 1994. Miconia skeaniana (Melasi
A*®, W.S. 2007. Revision of Miconia
Monogr. 81:1-235.
"*t) R.S. Bea Brittonia 40:368-391 .
PI. Sd. 1 69:963-979.
rpecie7of ////c/um subsection Parviflora (llliciaceae) from the
Variation in inflorescence position. Brittonia
274
Judo, W.S., J.D. Skean, Jr., and R.S. Beaman. 1988. Miconia zanonii (Melastomataceae: Miconieae), a
paniola. Brittonia 40:208-21 3.
Judd, W.S. and D. Skean, Jr. 1991. Taxonomic studies in the Miconieae (Melastomataceae). IV.
among terminal-flowered taxa. Bull. Florida Mus. Nat. Hist., Biol. Sci. 36:25-84.
Judo, W.S. and LC. Majure. 201 3. Miconia becqueri, a
ovaries from the Sierra Maestra, Cuba. Brittonia 65(3):(ln Press). DO1 1 0.1 007/s1 2228-01 3-9312-2.
Liogier, A.H. 2000. La flora de la Espahola. Vol. 9. Instituto Tecnologico de Santo Domingo (INTEC), Santo Domingo.
Macbride, J.F. 1941. Melastomataceae. Flora of Peru. Fieldiana, Bot. 13:249-523.
Martin, C.V., D.P. Litue, R. Goldenberg, and fA. Michelangeli. 2008. A phylogenetic evaluation of Leandra (Miconieae
Melastomataceae): a polyphyletic genus where the seeds tell the story, not the petals. Cladistics 24:317-327.
Martin, C.V. and F.A. Michelangeli. 2009. Comparative seed morphology of Leandra (Miconieae, Melastomataceae).
Brittonia 61:175-188.
Mayr. E. 1 970. Populations, species, and evolution. Belknap Press of Harvard University Press, Cambridge, Massachusetts.
Mayr, E. 2000. The biological species concept. In: Q.D. Wheeler and R. Meier (eds.). Species concepts and phylogenetic
theory: a debate. Columbia University Press, New York. Pp. 1 7-29.
Michelangeu, F.A., D.S. Pennys, J. Giza, D.L Soltis, M.H. Hils, and J.D. Skean, Jr. 2004. A preliminary phylogeny of the tribe
Miconieae (Melastomataceae) based on ITS data and its implications on inflorescence position. Taxon 53:279-290.
Michelangeu, FA., W.S. Judo, D.S. Penneys, J.D. Skean, Jr., E.R. BEccjuer Granados, R. Goldenberg, and C.V. Martin. 2008. Multiple
events of dispersal and radiation of the tribe Miconieae (Melastomataceae) in the Caribbean. Bot. Rev. 74:53-77.
thV'" Melastomataceae. In: P. Acevedo R. and M.T. Strong, eds. Catalogue of seed
Mishler,B.D. 1985. Then t. 98.531 562.
ologist 88:207-214.
Presch,W.1
d phylogenetic basis of species (
xidentalis vol. 9, Berolini &
0 EL Ekman. I
a cl. EL Ekman 1 924-1928 lectae. Arkiv Bot. 22A:63-64.
a cl. EL Ekman 1924-1930 lectae. fi
a debate. Columbia University Press, New York. Pp. 55-69
Wu.ACK,JJ.l972.CertamenM.astom;t;i^^^^^
CRATAEGUS TENUIOR (ROSACEAE)— AN INTRIGUING NEW SPECIES
FROM THE OKANAGAN OF BRITISH COLUMBIA AND WASHINGTON
AND A NEW VARIETY OF C. OKANAGANENSIS
J.B. Phipps
London, Ontario, N6A 5B7, CANADA
ABSTRACT
RESUMEN
INTRODUCTION
of my accumulated duplicate collection prior to its recent dispersal revealed
■axmrnmyofPacificNorthwestCrnmcgusTteCrmaegmdescriWforthatregioncanbedmded™^^^^
orpurple-toblackJruitedgroupsbased on color of their fully ripe fruitandlxttharerepresent^^^^^^^
ThispaperdescribesCrataegustennior,spnov.(reddruited).andC.oknnag<memrivarJ<mg.sp,nn™^
l*-fruitcd),ensuringthattheveryrichvariationencounteredbytheauthormttegenem arra^ e n
lun*ia01m„aganisLlyaccount«ifor,atleastasfarasdistinctiveandrq«atedlyoccnrr.nglortnstsc^^^^^
^“d™fof the Pacific Northwest wererevisedhyPhi^^^^^
OhnaganheingintwLries.ser.Ro,„-nh/olineandser.M«r«m,.hne.The^^^^
MTMandCshcila-phippsiacandwasrevisedfmtherepo^^J^i'n
^ plane-sided nutlets and glandular petioles. Senes Macracmtlm, witn
P=Uoles,ishererepresenteLnlyhyC.-m.crncamhn.Crma,*»sch,,s«nrpaandC.marrncand.aarev^
“ngingspecies occurring from the Okanagan to the ^ mucracamhu
The new species, C. tenuior, is local and, at first g ance, v ry occidentalis with white anthers,
ex Uud. var. occUeulalis (Britt) Kruschke. Cmlaegus mmmcanAa var. occufeMom,
Journal of the Botanical Research Institute of Texas 7(1)
is the most usual form of C. macracmtha west of the Rockies and is the only variety of the species so far record-
ed from British Columbia. Crataegus tenuior was first collected by myself and R.J. O’Kennon of BRIT in Sep-
tember 1993 in the Shuswap area of British Columbia but, in spite of the similarities alluded to above, several
differences from C. macracantha var. occidentalis soon became apparent. These included: often more rounded
leaf apices, generally smaller and less coriaceous leaves, small pink anthers, generally more slender thorns, and
more delicate plant habit (Figs. 10, 11, 12). Moreover, C. tenuior, like C. chrysocarpa, but unlike C. macracantha,
has the sides of the nutlets ± plane or, occasionally, only irregularly and shallowly pitted, as well as glandular,
sometimes strongly so, petioles. In addition, C. tenuior shares with sympatric forms of both C. macracantha var.
occidentalis and C. chrysocarpa Ashe var. chrysocarpa, the most common form of the latter species, the trait of
ripening fruit passing through an apricot to salmon or orange phase on the way to ripening to a bright red (see
C. macracantha - Figs. 13a, 12b), different from most other forms of both species, particularly those to the east
of the Rockies, the fruit of which normally changes directly from green to red. However, in spite of sharing
important charactenstics with C. chrysocarpa, it is with a slender form of C. macracantha (Fig. 12b), sympatric
with C. tenuior, that C. tenuior is most easily confused.
Crataegus tenuior proves to have a quite limited distribution predominantly in the northern Okanaganof
southern Bntish Columbia, mainly from somewhat north of Armstrong to the vicinity of Enderby in the north
central Shuswap area where it is locally common. From this region it extends thinly into the generally drier
Okanogan of Washington where Crataegus as a whole peters out. Mainly collected in the mid 1990s and 2002,
C. tenuior remained undescribed until now because of the need to assess C. macracantha throughout its large
range, the greatest of any North American Crataegus, prior to deciding whether it should be included in that
specms. Although C. macracantha has proven to be very variable through much of its range, especially so in
Quebec and Montana, it has, except for a population in northwest Montana, strikingly eglandular petioles, a
trait usually thought of as species-specific. Consequently, a morphometric study of relevant Pacific Northwest
material was conducted to investigate these distinctions.
f study.— Fourteen flowering OTUs and 54 fruiting OTUs (Table 1), constituting all the
fomCT VWO (now TRT) specimens of pmative Crataegus lenulor and C. macracaMha from Brilish Columbia
Washington, were used. The flowering specimens were scored lor seven characteristics mean length of
three largest leaves, petiole glanduUrity (Figs. 9a, 9c), median thorn length, median thorn thickness, sepal
irargin serration (Figs. 9b, 9d), anther color (Fig. 11), and style number. The fruiting specimens used the same
c aracters except that nutlet number replaced style number (it is usually the same) and nutlet lateral pitting
placed anther coteMissing data, if in less than 10% of OTUs, was repUced by column (character) meansol
the b^est subset of OTUs considered, u pmteriuri, unequivocally the Le taxL if more than 10% missing
^N-TAX ZMO*™! cluster analyses were performed using Podanfs
5YN TAX 2000. The data sets were ranged before analysis. The results are as follows:
The ordtnattons PoCA (Fig 3) and MDS (Fig. 4) of the euclidean distance matrix siraibrly sepan
Cru,ueg„s,e„„.mandC.macr«cu„,hucleatdy.Thefirstaxisoftheprin7p:if^^^^^^^^^^^^^^
le group alv
for 62% of the variation f'"* ““
( - 4) m small, the concordance of very different methods emphasizes tf
comprising s
277
RZila,'WP'=Wm Phipps, and'TCB'=l
), 3 (6923), 4 (8370), 5 (8389), 6 (8400), 7 (8401 ), 8 (6902), 9 (6903), 10 (6963), 1 1 (8304), 1 2
4 (701 1), 5 (701 3), 6 (7171), 7 (8265), 8 (8274), 9 (8451), 10 (8452), 1 1 (8453), 12 (8454),
J), 18 (6792), 19 (8275), 20 (8473), 21 (6484), 22 (7103), 23 (7025), 24 (6783), 25 (8457),
4), 31 (6790), 32 (6781), 33 (6787), 34 (6793), 35 (81 76), 36 (6782), 37 (8275a), 38 (6976),
, 42 (TCB4041 ), 43 (8463), 44 (8464), 45 (8478), 46 (6789), 47 (6867), 48 (6871 ), 49 (Z1 7891 ), 50 (Z1 7907), 5 1
(Z20320), 52 (1 7904), 53 (Z22423), 54 (7265).
, , „Hr^tnflpnticulate sepal margins, 3 or 4 styles, short
TheCrataegus tenuior OTUs all had pink anthers, :.entxr occidentalis OTUs
Jwms (mean = 40.14 mm) and short leaves (mean = 41.66 mm) wh 1 _ ^
l>^whiteorivoryanthers,moredeeplyserratesepalmargms,2or3styles,shortto g
Aoms and short to longer leaves (mean = 47.00 mm).
•^Fnutiiigii
::;emeisolani(OTl]22)inFigure5whichis
uch larger nght-hand group. There is also an extr identifies the
•^edOTU of all and adds on after fusion of the two larger fairly laj-ge ei
‘blaster of OTUs
, ^ „Jf"STUs clustering in two fairly large groups
■‘'P‘'“l*“nd“’’;::;u^dLoT.?s22andM.SL,alsono<shown,isvery^^^^
278
UPGMA except that its left-hand group also includes OTU 39 and the large right-hand group lacks several
OTUs which are added only after the fusion of the two big groups. Here also OTU 22 is added last, further
emphasizing its isolated position. Finally, ISS (Fig. 6) divides all 54 OTUs into a left-hand group identical to
that in UPGMA and the remainder. In this, as in CL, OTU 22 is not as isolated as OTU 54.
Summarizing the cluster analyses, there is a consistently occurring group of OTUs 1-16 plus 19 and 26.
At least the great majority of these may be referred to Crataegus tenuior as they lack, or occasionally have a lovf
evd of nutlet pitting or scarring. Similarly, the remainder, comprising a large right-hand group, outliers, if any-
to t e left-hand group of C. tenuior, plus outliers, if any, to the combined main groups, may in large part be «-
terrri,„C.macrac»«haasd.evWh-.H„^„.„,„„...... 22 and 54. re nouble outliers™*
Ainong the ordinations, PCoA (Fig. 7) yields a left-hand group of OTUs 1-16 plus 19 and 22 and a tag!
group of the remainder. Here 41% of total
picked up by the first axis. MDS (Fig. 8) yields a top-ri^‘
mT “ “P “ bottom-left group of the remainder. In »
s 6 and 54 are peripheral. This strong isolation of OTU 22 in MDS reflects the UPGMA situation. As
the duster ^lyses, a discrete Crataegus tenuior group may be found that is characterized by a lack of, oral
^‘^^rri^g And as in the flowering analyses, all the OTUs with four nutlets
^’^cept for the isolated OTUs 22 and 54. Significantly, the two very differe^
‘he same main patterns of variation and also detect very similar groups to those
„ ^™‘^o’^istendesregardingOTUs22and26.Thefruitinganalysesarethussoin^
mumalfy ™ilar than the flowering analyses, an observation in line with the lower perceuug! of
vanance accountedforbyaxis 1 of the PCoA.
d-16 + 19.Inth^
nilar core grouping of OTUs 1-
lie in the putative u
ordination methods identify the &
the cluster analyses w
Phipps, Crataegus tenuiorsp. I
analyses, the core tenuior group contams those specimens wttn ^ ^
of 0-1 ona three-poin. scale), glanduUrpeHotejl«m^w^^^^
cbtactercouldstillberelUblyinlerpreted.nfru«,a^s^^^^^^ InU^ Hl5;andOTUslO,16, 19,
splits into three subgroups containing: OTUs 4, 5, 7, 9, 13,0 ’ ’ ’ ’ j tlir.iiah the significance
■l^taaad las. of LcUve higher petiole ^ularily than the seco«lsubgro„p.^^^^
of diisis not clear. Similar subgroups are found in Figure 6 (ISS) ^ they ^ ^
»"3(Figs.7,8).,ti^ofcourse,afeatureofglandstba.d.eyn«y^^
“tat^aybeaumreconsuntcharacteratanthesiscanbelessajelWeE^^^^^^^
S®t^g»d,h the tauiorcore ‘ht.thasa.mM.ng nntte^r _ fruitinganaly-
Cr<nnegnsmncrncan,ln.isrepresen.edbya ofdK«m g ^
. rhose specimens with P"** - "'1^,
18 70-54macracanthagroup,plusOTU
^except OTU 22, see below, interpretation. All the OTUs m tte 8^ 2^ ^
^ We nutlet sides significantly (score of 2) to suox^ bc ) ^
’^hich have middling nutlet side scores (respectively 1.25, UX i. )■ macracmtha
-t.in.esc.nster,oUwi.b.becote.em,iorgmuptbou^^^^^
Ponp but in the part closest to the tenuior group. Wit ^3 comprises the smaUer-
^-dchaster containing OTUs 17, 18, 20,21, 29,30,32,K^^^^^^
^v^specimens with dimensions more likeC. tenumrThe^^^^
thevanousanalyses,afactparticularlyseeninbolh ^^^^bly that the fimitingsam^^
CLdendrogram that is not shown. A major reason ^ diverse, this no doubt due to the
l^gerthanthe flowering one(540TUscompared to 14), andAe^^^
longer time-frame over which adequate specimens can be collect .
Journal of the Botanical Research Institute of Texas 7(1)
Thef
However differences as mentioned above-
xr ^ ruiting OTUs more variation is present and two mutually exclusive groups do
*^'*‘^’^g^;Neverthele^,agroupofl-16 + 19isconsistentlyfoundwhichisCrataegusfenMiorInfruit,thebest
interpretation seems to be to accept the presence of plane-sided nutlets as decisive for this species and to con-
oHowrnutlivemr'^'TT''1I'1^^^^^
haps Vbt^d betw^n chTiX '*■
up L test this and, minimal wtl^ldr" '"*r<tcanlha. However, the sntdy was not set
an oddity, perhaps cuItivatedXm the ^'XHtM^XT*'
form svith its eglandular petioles and pitted nutlet.; Okanagan. This is clearly a C. macracant
cnn,hnindividn*abintheXdy,as,.rt:Xh^‘:“““
more distinct features did nnt aet parts, again quite unlike the others studied. Some of its
tog thornless, it achieved thorns of meXenXs f“n''«inore>
toaanahtyntotedistinctthantheatalysessLandXtaXr^
g u-h!tn>mTonasket,Washmgton(Fig Mhlhisindwidualappearsthentost
V. and a new variety of C. okanaganensis
extreme of all in UPGMA and MDS, is second most extreme in ISS and is peripheral m PcoA. It is the only OTU
in the study to combine four nutlets, highly glandular petioles, and deeply pitted nutlet sides. Unlike C. tenuior,
ithasessentiallyglandlessleaf-teethatxlO.Also,it is the only specimen of the 68 studied tohaveadistmctively
non-macracanthoid leaf shape, one with clear resemblances to C. chrysocarpa. This, 1 suspect, is a very rare
^ple in the study area, perhaps unique in collections, of an hybrid or clone thereof, between C chryso-
‘^^^ndC.macracanthavar.occidentalis.
Confirmation that the calibration of fruiung and flowering groups to taxonomic names is consistent is
by two flower™ sDecimens attributed to Crataegus tenuior (JBP 6814 and JBP 8401) being from the
plants as JBP 7171 and JBP 8454 (fruiting) while for C macraeantlia var. occidentalis JBP 6903 (flowering)
comes from the same plant as JBP 6871 (fruiting).
282
Bus es, 2.5-4.0 m tall, multi-stemmed: l-year old twigs tan to reddish brown, older gray-brown; thorns 3-5
cmlong straight to slightly recurved, thinnish, glossy black-chestnutwhenyoungAeaves deciduous; peti^^^^
1.25 2.5 cm long, pubescent only in sulcus, variably gland-dotted; blades (2 5-)3-5(-7) cm long, broadly el-
liptic to ovate or rhombovate in general shape; apices subacute, bases broad-cuneate, shallowly 4-5(-^) lobed
persidMobesacute to obtuse;margins serrate, teeth with conspicuous black marginal glands (except±entire
m basal third), venation craspedodromous with 5(-6) pairs of lateral veins, veins adaxially not strikingly im-
pressed at maturity, adaxially densely short-appressed hairy, abaxially thinly pubescent on the veins, other-
wise glabrous, texture thin, autumnal color bright yellow. Inflorescences abt. 10-flowered; branches tomen-
mbranous, gland-margined bracteoles, glands subsessile. Flowers 15-17
ns ± glandular-dentic-
, , .^..lalKca. 0.6 mm); styles
t-, ± broad-ellipsoid, <
B most easdy distinguished in dower (Figs. 9, 11) when it diders from sympairic C
mnrmrandinbyavpety of characters asdisciissed above. In fruit. however.Clenui^
tmguish from smaller-leaved specimens ofC macroconthfl Without dissect fh Th ' because the
petiolar glandularity character has some overlap, the sepal margin cham^m'ayTrracnU m discern, d*
vithout dissection, and the nutlet numllr™'^' l-f^eoueml'^f™!,''' ‘''a
't irequently found, is not diagnostic. At that stage,
although autumnal
leaf color (yellowish ii
Phipps, Crataegus tenuior sp. nov. and a new variety of C. o
fit7.StandardizedPCoA of eudideandistance matrix of 54fruitingspedinensofs
group is C macmcantha; OTUs 22, 54 exduded. OTUs 22, 26 and 54 are starred.
i»y the smaller leaves, and greater delicacy of habit, twig, thorn, and leaf texture wtth the
otC. mncracnntha. With regmd to C. d,rysocarpa, in spite of sharing several chamctem m — J*
h«ior,seefcey,ithassttchadifferentfaciesthat it is easily separated nmcrosc^tcahy. The foUo™^^
■prizes critical distinctions between C tenuior, C. mneracn«.l.ovar.
Wclirysocarpainthecontextofallthered-fmitedCratacgiofthcOkatiagan. oe
it is best to check several fruits for nutlet pitting.
Journal of the Botanical Research Institute of Texas 7(1)
C. macracantha
uch'more'norifei^r mT'" ''*™**' to C macraamlha»
“'“'ifT 1®’ ‘i"™* n overlap'^
smaller-leaved and smaller innorescence tormsof C. maeracmrt..
suggested origin for the sporadic C. lotnemimm (OuleT «Tr “““,f "“"T*'' " '’'TTf ,m-
named form in Minnesota and perhans th u a ’’ ®
Great Plains (Phipps & O’Kcnnon 2007). Stffl Tis nombkT!'l '"“"'“'t
T'!^'!'r"‘‘^'^*7^“^”‘^^«reater01mnava„.^C
^ this paper). This being so, the distinctive Okanagan en-
Phipps, Crataegus tenuior sp. nov. a
Mmsttadbeconsideredacandidale parent. But CM ^ pitted nutlets. Thus, in
■Ple.latge-sepalled fruit of C. utrrutircns. Also, the cannot be discounted as a
k of precise intermediacy between any two sympatnc , passing, one may speculate
c^nism of origin and resolution of the matter ^ and in discussion of the morphometric
t the small-leaved form of C. macracantha, noted at th gi larger-leaved C. macracantha
'Jts.nughtrepresentintrogressionorhybridoriginbetweenC.tenm^ g
• occidentalis.
Phipps, Crataegus tenuior sp. nov. £
Phipps, Crataegus tenuiorsp.nov. a
289
drawing attention to the following additio
In the Mogollon Mts. of New Mexico a small and isolated population of a macracanthoid Crataegus with glan-
dular petioles is known. This is represented by O’Kennon, Huff & Snowden 13427 (TRT, flowering), Huff et al.
1380 (UNM 86261, fruiting), and Kirkeminde 897 (UNM 83115, fruiting) and has the further features of pink-
purple anthers and yellow autumnal color but clearly pitted nutlets. This population lies well to the south of the
southernmost populations of C. chrysocarpa in central Colorado but too little is known it about to warrant
further comment. Rather similar populations of C. macracantha in Arizona have pink-purple anthers and pit-
ted nutlets though typically eglandular petioles — such populations appear scarce west of the Rocky Mountain
. occidentalis and there is an unconfirmed report by Nadia Talent (pers.
ma there is a substantial population of a
pink-anthered, glandular-petioled, pitted-nutlet morphotype closely resembling C. macracantha.
CRATAEGUS OKANAGANENSIS VAR. LONGISPINA
Crataegus okanaganensis (ser. Purpureofructae) is a large hawthorn with stout thorns, readily distinguished
from C. douglasii and immediate allies not only by thorn measurements and fruit color but by the serial char-
acteristic of typically large, pointed, fruiting sepals in contrast to the much smaller, often blunt ones found m
burgundy in late August, purple-black in September .
•ita "'“k" T e tairy frui.. This ensemble of chamcierisilcs ^ no.
“«l^n,andfor,hisreasonanewvarie.yisrecognized. ^ southeastern
Crataegus okanaganensis var. longispina is a local taxon
Shuswap region, from Enderby to Salmon An
290
Phipps, Crataegus tenuior sp. nov. and a new variety of C. okanaganensis
^“shes, 3-8 m tall; extending twigs densely pubescent, at 1-yr. old ^
2%
Additional Cited Specimens [collectors abbreviated from following names: J.B. Phipps, RJ. O’Kennon, and R. Jackson (forJBP)]:
Aug 2002, Rj^BP 8452 (TRT, UBC); Back Enderby Road, W side, S of mailbox 5135, natural hedgerows, fruit orange-yellow to bright red, 25
Aug 2002, RJ/JBP 8453 (TRT, V); Back Enderby Road, W side, S of mailbox 5135, natural hedgerows, fruit yellow-orange, 25 Aug 2002, R]/]B?
8454 (DAO, TRT); Back Enderby Road, just N of Stepney Cross road, W side, S of mailbox 5135, natural hedgerows, fruit bright red-orange,
22 Sep 2002, RJ/JBP 8470 (TRT, UBC, V); Back Enderby Road, just N of Stepney Cross road, W side, S of mailbox 5135, natural hedgerows,
fruit bright red-orange, RJ/JBP 8471 (DAO, TRT, UBC), Back Enderby Road, just N of Stepney Cross road, W side, S of mailbox 5135, natural
hedgerows, fruit bright red-orange, RJ/JBP 8472 (TRT, V); north central Shi.swcp 2 km E of Enderby, bush 3 m, fruit scarlet, 25 Sep
1993,;BP& 0X6811 (DAO, TRT, UBC, V); 5 mi E of Enderby, bush 3 m, fruit scarlet, 25 Sep 1993, JBP& O’K 6814 (TRT); 5 mi E of Enderby,
bush 3 m, fruit scarlet, 25 Sep 1993, JBP & OX 6814a (CAN, DAO, TRT, UBC, V, WTU); Ca. 200 m E of Shuswap River bridge on Enderby
a^^UBC ^U)^ **^200^ of rivCT bank among scrub, slender bush, 4 m tall, fruit light salmon, type bush, 20 Aug 1994, JBP & OX 7011
hghtsalmon,20Augl994,JBP70i3&OX(DAO,TR^%alleyoftheShLwa’^t.TkmEofEnc2byaT«k^^
e, 7 Sep 2001 , JBP 6- O’K 8274 (TRT); Anderson Road, S side, at 1.5
fruit orange-red, 7 Sep 2001, JBP & O’K 8275 (TRT); RJ/JBP 8474
n, fruit scarlet, 4 Sep 2001, JBP & O’K 8265 (TRT).
I rlf , f British Columbia and adjacent Washington (e.g., Hitch
cock & Cron,u,« 1973; Taylor 1973; Brayshaw 1996) mported few species for the area intLive collectini
between 1993 and 20O2in the greater Okanaganhasyieldedafnrthere^t species and several varieties of tlii
gen^lPhipps&OKennon 1998, 2002, 2004). Thus, far frotnapositiot^^
Si"!" ' “ Bas sufficient open habUa! and the Lr^rature band!
relevant parts of the author's treatment for Crataegus in Flora of North Atttei
ACKNOWLEDGMENTS
1 wish to thank Bob O’Kennon of BRIT for many happy and fruitful hours of companionship in the field during
the discovery process for these new taxa and for several photographs as indicated on their captions. Antony
Uttlewood, Dept, of Classical Studies, The University of Western Ontario, is thanked for checking the Latin
diagnoses. Nadia Talent, University of Toronto, and Tim Dickinson, University of Toronto and Royal Ontario
Museum, are thanked for comments on an earlier version of the manuscript. Alan Noon, Dept, of Biology, The
University of Western Ontario, is thanked for photography of herbarium specimens and Ian Craig, Imaging
Unit, Dept, of Biology, The University of Western Ontario, for his assistance in preparing the figures for the
paper. Financial assistance for publication was provided by Barney Lipscomb and Bob O’Kennon.
BrayshawJ.C. 1996. Crataegus. In; Trees and shrubs of British Columbia. U.B.C. Press, Vancouver.
Hitchcock, CL and A. Cronquist. 1 973. Flora of the Pacific Northwest. U. Wash. Press, Seattle.
Pms, J.B. 1998.The red-fruited of western North America. Canad. J. Bot. 76:1863-1899.
Phiws, J.B. AND R J. O'Kennon 1 998. 1
C okanaganensis, and C phippsii. Sida 1 8:1 69-1 91 .
Phipps, J.B. and R.J. O'Kennon. 2002. New taxa of Crataegus (I
Shuswap diversity center. Sida 20:1 1 5-144.
Pms. J.B. AND RJ. O'Kennon. 2004. A review of Crataegus ser. Rotundifoliae in western Canada. S'da 21 :65-7 .
PW!,Ji.»DRJ.O'KE»o«.2007. The hawthomsoftheCypress Hills, Albertaand Saskatchewan. J.eot.R«.lnst. Texas.
1:1031-1090.
TATi£)fl,T.M.C. 1973. Crataegus. ln:The rose
/. Mus. Handb. No. 3
298
BOOK REVIEW
Geoff Williams and Paul Adam. 2010. The Flowering of Australia's Rainforests: A Plant and Pollination
Miscellany. (ISBN: 9780643097612, hbk). CSIRO PUBLISHING, P.O. Box 1139, Collingwood, Victoria
3066, AUSTRALIA. (Orders: WAvw.publish.csiro.au (Australia), wAvw.styluspub.com (U.S.A. & Cana-
da), Avww.eurospanbookstore.com (U.K., Europe, Middle East, & North Africa)). AU $99.95, US $93.00,
216 pp., 8" X lOi/i".
A word of caution before continuing: this is obviously a beautiful book, with attractive cover, excellent photo-
graphs, and appealing content and information. Be very alert to the second sentence in the Preface: “But read-
ers will find that from time to time we have taken a circuitous path that too easily voyages to the shores of other
t belong solely or even tenuously to the rainforest envii
related knowledge are given an airing in diverse quarters" (Italics mi__. .,
If you are planning to use the excellent and diverse information giver
pen or pencil close at hand. Note page numbers and subject content for key
ill find related
1 if you have
m find them easily. Comparisons
recorded locations of previous discussions of the same or related topics a
are not only helpful but occasionally unexpected.
After the explanatory Introduction, the first chapter, “Flowers and pollination in lore and legend” Avasa
bit unexpected but very informative. A nice addition, to be sure. The second chapter, “Categorising rainforest
plants,” is self-explanatory. Immediately following are 34 color photographs, beautifully providing an over-
view of plants and habitations.
Chapters 3 through 9 are quite diverse in subject content. They cover a variety of helpful (and sometimes
unexpected) topics. For example. Chapter 4 is entitled “Being a flower”— a relatively lengthy chapter; Chapter
5, “Introduction to breeding systems;” Chapter 6, “Spatial and temporal structure of rainforest: general media-
nisms that influence pollination and reproductive ecology;” Chapter 7, “Australian vegetation history audits
m luence on plant-polhnator relationships;” Chapter 8, “Pollination and the Australian flora;” and Chapter9,
Pollination ^dromes: who brings the ‘flower children’ in rainforest?” The titles definitely catch one’s atten-
E^rap^ndktmdtrbM'^T^'"^''^^^
rainforests: implications for the conservation of remnant communities,” echoes and confinns
Ancrraif fragmented parts of the Australian rainforests. Before the European settlements the
ta -dlir, ” on the eastern coastal fringe, existing commuaidrs
Ohvii. ? threatened by invasion of exotic weeds and animals and by human impacts.”
awaren -!!h / "" " ^ut of nearly Avorldwide
avvareness. e enjeude. Volunteer & Assistant Editor, Botanical Research Institute of Texas, Fort Worth. Texas,
I. Bot Res. Inst Texas W: 29*. 2013
IDENTIFICATION, DISTRIBUTION, AND HABITAT OF COREOPSIS
SECTION EUBLEPHARIS (ASTERACEAE) AND DESCRIPTION
OF A NEW SPECIES
Bruce A. Sorrie, Richard J. LeBlond, and Alan S. Weakley
CB. 3280, Chapel Hill, North Carolina 27599-3280, U.SA
RESUMEN
INTRODUCTION
C»^isisagenusof35 species, 28 in NorthAmericaand seven in .he neotropics
mSectionEuhlepharis Nutt, is strictly easleiarNorth American, rangingon the coastal plamto
Nova Scotia to southern Florida and eastern Texas. There has been hmite a^eeinent on e
thatpreriousauthorsrecognized (Table l),withdisagreementcenteringonC.gIad^
plant morphology (several characters), habitat fideltty, and “f™' from nuclear ITS and plastid
dons, presented below, are largely supported by tnolecularwor^^^ngseq ^^^^^^^^^^^^^^.^^
■oglomCCrawford&MarkTOOSXwhoreportawell-t^lved^^gwy ^
including monophyly of section Eublephans. The authors sample ^ ^ ^hat “additional field
C gladiata, C. integrifolia, C. linifolia, and C. rosea. However, Cm ^ provide
and labratory studies are needed to elucidate relationships m t pe^ ^ Boynton C jloridana E.B. Smith, C.
t^le for tecognizingnine species in sectionEublephnns|^CMau^-^J™^^^^^^^^,^^^^^^^^^
al other taxa not recognized here are synonymized in the treatment
Thecu d d d ■ 1 by the first author over a period of twenty years while gathering
specimen information on endemic species of the North American specimens accessioned
Smith served as a baseline, augmented by newly constructe for seven species; Godfrey and
«nce Smith’s work. Smith (1976) P'^ovided illustration
Wooten (1981) illustrated six species. These drawings we ygriation within each species. Measurements
^mples of the species; herbarium specimens provided data on ^ (jgyg ^978) cited
'^ere made of 20-23 mature achenes (1-2 per specimen) from spec
It is describe as follows:
1. Coreopsis aristulata LeBlond, Sorrie & Weakley, sp. nov. (Figs 1, 2).
Differs from C. palusiris in the following combination of character
S«rie (= C. heiianthoides Beadle misapplied), but differ markedly in
PlantsThizomatous; rhizomes ro 14+ cm, 2 mm wide, these and the ro«sy'>l^
giaied with roots or hardly distinguishable from lower stem. Stems g rous, ^ lowest 4-6
t-.lO-BmKiesbelowLflorei;ence.Canlme leaves gmduallyreducrf^"!^^^^^^^^
"“ies leaOess at anthesis. Petioles 2.0-3.5 cm long, leaf tissue ° ^
sheaths 1-2 mm long. Blades glabrous, linear ,o „ LVry) margins mi-
^Kxasbngas Wide, thick, s.iff,hri.ilewhendry,surfa«ssm^h.shno^^^^^^
Dutely scabrid. Inflorescence 13-50 cmlong, open, 0 8_1 ^ide at midpoint, 0.4-
me rays. Outer phyllaries narrowly lance-oblong, blunt, r Vt 5 mm long, 3-4.2 mm wide at
^aslongas the inner; innerphyllariesovate-oblong^pere^^^ .^^^^
''^estpoint.Achene bodies oblanceolate-oblong,(3.0-)3.2 •
Wings); awns 0.2-0.4 mm long; wings 0.1-0.3 mm wide,
slow Co.: Haws Run N
It, Thalictrum cooleyi, F
. -D (NCU). Pender Co.: S ^ j,.
ivirentumvar virgalum 23Oct2011.R.
, ’ 4 r aristulata are the only species in section Eublephans
s. — Coreopsis palustrts and ^ or two shriveled senes-
lous at anthesi:
‘^ent leaves. This is a striking character 4. .
to herbaria. The two species can be distinguished by the chatacte
NoKthatleafwidihandacheneawnlengthdoiiotoverlapbetweenthelwospecies;m{acl,awnlenglho
C. amlulata is only 38% of that of C. palusths. The narrow leaves of C. arislulala are simiUr to those of *
narrow-leaved extreme of C. glndtotn (described as C. Imgifolia Small), and specimens of the latter mayh
mistaken lorthefornrer.Howev^.leafbladesofCnrismlmaamevenlongerandmoren
of C ongi/olin , the lowest « leaves are absent at anthesis (vs. present), and the achene awns are sthkitigH
short (mean 0.3 mm vs. 1.5 mm).
Habitat and Dates^orecrpsis aristulata occurs in the Very Wet Loamy Pine Savanna natura
community (Schafale 2012), equivalent to the Pinus palustris-Pinus serotina/Magnolia virginiana/Sp
1^ Ti! V Woodland (CEGL004500) (NatureServe 2012). Community structure ii
^ th ^ 1 ^ dominated by Pinus serotina Michx., and usu
ally with someP.palitstnsP.MiUandTaxodinmnscendemBrongnTheshrnbUyer typically issparsetop^^^^^
M i. Characters ot Loreopsis amtulata vs. C palustns. N
IS of creeks and ri
e North Carolina tc
With Morelia caroliniensis (P. Mill.) Small, Cyrilla racemiflora L., Ilex glabra (L.) A. Gray, I. myrtifolia Wall, and
Vacciniumfuscatum Aiton prominent. Juvenile Acer rubrum L. var. trilobum Torrey & A. Gray and Nyssa biflora
Walter are often present. The herb layer is dense, and dominated by combinations of Ctenium aromaticum,
Sporobolus pinetorum, and several Rhynchospora taxa [e.g., R. pinetorum Small, R. latifolia (Baldwin ex Ell.)
Thomas, and R. thomei Krai]. Flowering dates: mid Sep-mid Oct.
Etymology.— The specific epithet, aristulata, means “with short awn" or “with short bristle.”
Range.— Coreopsis aristulata occurs only in Onslow and Pender counties. North Carolina (Fig. 3). Other
locally endemic species share tliis range and specialized habitat: Allium sp. nov. aff. cemuum Roth (Alliaceae),
Carex lutea LeBlond (Cyperaceae), Scleria sp. nov. aff. dliottii Chapman (Cyperaceae), and Thalictrum cooleyi
Ahles (Ranunculaceae); see LeBlond et al. (1994) and LeBlond (2001) for a discussion of this remarkable eco-
system. The three localities cited under Specimens Examined occur within eight km of each other and are part
of a complex of savannas that are remnants of a once larger ecosystem.
i F.E. Boynton, Biltmore Bot. Stud. 1(2):14L 1902.
This and C. nudata are the only spring/summer blooming species in the section and both have basally disposed
leaves^ C. nudata is easily distinguished by its pink to deep rosy ray ligules, terete leaves, and more southern
istnbution; the two species are allopatric. Seven out of 51 specimens at NCU lacked leaf lobes (or we could not
see them clearly); hence this species is keyed twice. It features very short achene awns, averaging 0.5 mm; only
those of C. anstulata are shorter. Furthermore, achenes of C.falcata are truncated at each end, unlike any other
p^ies. It inhabits seepage bogs, wet savannas, wet borrow pits, and seepy streamsides. C.falcata ranges from
ex reme southeastern Virginia [Fleming and Caljouw 10435 (GMUF, WILLI)] to southern South Carolina (Fig.
4). It was reported or Georgia in McIntosh and Bibb Counties by Jones and Code (1988) and there is a speci-
men at GH snnpty labeled “Georgia”; it is possibly more common in Georgia than these few records indicate.
s.n. (NCU)], which is far out of range but plausible if collected in a monta
May-late June(-earlyJul).
3. Coreopsis Horidana E.B. Smith, Sida 6:192. 1976.
^ and broader, resulting in an elliptical shape
“andZTb^'d ‘"e drawing in Lith (1976) represent
esemblmg C./alcotu. Uese two speetes ate amply distinguished by Dowering period, length of outer
s usua y present in C.falcata. Coreopsis jloridana inhabits wet savannas, flat'
Soriie et al.. Coreopsis sect. Eublepharis
305
Most plants have broadly elliptic basal and lower
leaves and are strikingly distinct from all other
taxa; only C palustris (=helianthoides) may have
similarly broad leaves, but those occur only in
the mid-cauline region because the basal and
lower leaves have senesced by anthesis. Core-
opsis gladiata and C. linifolia have the longest
respectively. Plants originally described as C.
longifolia Small represent the narrow extreme
of leaf width; achene morphology appears to
differences. This narrow leaved variant appears
more-or-less throughout the range, but is most
often encountered in peninsular Florida. Core-
opsis gladiata is most frequent in the East Gulf
Coastal Plain, where it inhabits wet pitcher-
plant bogs and streamhead seepages. Disjunct
populations occur in the mountains of the Car-
olinas and Georgia, where they inhabit mon-
tane seepage bogs. Coreopsis gladiata ranges
from eastern South Carolina to north Florida
and southeastern Mississippi; disjunct to mon-
tane Georgia, South Carolina, and North Caro-
lina (Fig. 6). It should be sought in southeastern
Louisiana. Howering dates; mid August-early
November.
Coreopsis integrifolia Poii
Suppl.2,352.1811.
s species is distinctive in its strictly opposite,
ite, and uniformly small-bladed leaves. Core-
is integrifolia is rare and local, known from
counties in northern Florida, five in south-
1 Georgia, and four in southeastern South
rolina (Fig. 7). We have not been able to verify
iport from North Carolina (Strother 2006). It
ts, and streamside seepages. This
species appears to prefer calcareous soils.
Flowering dates: mid August-early November.
Sonie et al.. Coreopsis sert. Eublepharis
307
Since its inception, C. hdianthoides Beadle has
been a misunderstood and often neglected
species. In part this is due to the general pau-
city of specimens in herbaria and in part due to
Beadle’s erroneous protolog. Beadle (1898)
designated a type [Aspalaga, Florida, October
1897, AW. Chapman s.n. (Biltmore Herb., holo-
type perhaps destroyed; isotypes MO!, US!)],
but the specimens actually are C. gladiata Wal-
ter, due to the presence of broad-bladed leaves
at all basal and lower culm nodes, and collect-
ed hundreds of kilometers outside of the prov-
en range of C. hdianthoides. We recommend
that the name C. gladiata Walter be conserved
for Walter’s plant; Smith has designated a neo-
type (see above). Therefore, the entity known
name: C. palustris Sorrie (Weakley et al. 2011).
The salient features of this highly distinctive
species have been overlooked by practically all
authors except Radford et al. (1968). Its leaf
phenology is matched only by C. aristulata,
with the lower 4-6 leaves senescent or decidu-
ous by anthesis, thus leaving the lower nodes
essentially naked. Coreopsis palustris is nota-
ble for the very broad achene wings (0.5 mm),
matched only by C. nudata. Coreopsis palustris
often grows prostrate with ascending distal
portions of stems, perhaps due to relatively
frequent inundation; of the species in section
Eublepharis, only C. palustris and C integrifolia
or stream overflow. Coreopsis palustris inhabits
swamp forests and openings, fresh tidal creek
margins, marshes, and borrow ponds. It rang-
es from southeastern North Carolina to Duval
County, Florida (last collected there in 1898)
(Fig. 10). Georgia specimens of C. palustris are
extremely rare; Camden County, 1 mi N of
Kingsland, 18 Oct 1950, Godfrey 50908 (FSU,
NCSC). Flowering dates: mid August-late
9. Coreopsis rosea Nutt., Gen. N. Amer. Pi.
The distribution of no other species in sec
Eublepharis has been as under-represented in
manuals and monographs as that of C. rosea. For example, Smith (1976) maps it in six
Rhode Island, New York, New Jersey, Delaware, and Maryland) plus Nova Scotia, Canada. Strother (2006) lists
six states (Massachusetts, Rhode Island, New Jersey, Delaware, and South Carolina) plus Nova Scotia. Howev-
er, there are specimens dating as far back as 1903 from Maryland (MO), 1871 from New York (GH), 1865 from
Pennsylvania (GH, MO), and 1900 from South Carolina (PH). Nuttall’s original discovery was made in Georgia
in October 1815: “about 20 mis. from Savannah on my way to Augusta I found . . . and a beautiful new purple
flowered species of Coreopsis!” (Graustein 1967, p. 101). This location would be in present-day Effingham
County. Nuttafl’s original description of C. rosea clearly stated that it occurred “from New Jersey to Georgia”
(Nuttall 1818, p. 179). Finally, there is a Leavenworth specimen collected in Georgia pre-1862 (Leavenworth
s.n. PH), as reported by Smith (1978). Figure 11 shows the distribution by county. Coreopsis rosea is unique in
the section in its very short and narrow achenes which lack wings and awns. The awnless achenes may suggest
affinity with some other section of the genus, as was done by Sherff (1936)— section Calliopsis— but molecular
data IS needed to guide current decision-making. Coreopsis rosea inhabits sandy to stony shores of ponds, lakes,
and other depressional wetlands, plus sandy margins of Waccamaw River, South Carolina. Flowering dates:
mid Jul-late Sep.
ACKNOWLEDGMENTS
WtgmeMlythank the cumorsand staff of .he followinghertaria for access to specimens:
GA,GH,MO,NCSC,NCU,NY,US,andUSCH.Twoanonymousrevtewerstmptoved.hema„uscnp.substa„.
tiaUy. The consulting firm of Yaupon, Dahoon, and Blolly provided logistical assistance.
tENCES
I the botany of the southeastern states. Bot. Gaz. 25:4
1.2005. Phylogenyo^ea^^^^^^^^^^
Cronquist, a. 1 980. Asteraceae. Flora of the southeastern United States. Carolina Press, C pe
Beadle, CD. 1898
Crawford, DJ.ani
from nuclear
bridge, MA.
Jo«s,S.B.,Jr.anc
^Bidno, RJ., A.S. Weakley, A.A. Reznicek, and WJ.
from North Carolina. Castanea 16:153-161.
■"■-"J.2001.Endernic plants of the Cape FearA.hr^ioa^^^^
E. 2012. NatureServe Explorer: an online encyclopeo
ihacaHlogue of the species throt.gh18t7.D,l
. Bttc 1968. Manual of .Ke .asculat flora of dte Carolinas. Univ. N
Guide to the Natural C Rasources, uareig...
1933. Manual of the southeastern flora. Univ. o ted States and Car ‘ ‘
E.B. 1976. A biosystematic survey of Coreopsis in eastern uni
h Carolina Natu-
I Resources, Raleigh.
, Bot. Ser. 11:277-^
la Press, Chapel Hin.
J States and Canada. Sida 6:123-21 5.
310 Journal of the Botanical Research Institute of Texas 7(1)
Smith, E.B. 1 978. Notes on Coreopsis. Sida 7:304-307.
Sttother, J.L 2006. Coreopsis. In: Flora of North America Editorial Committee. Flora of North America North of Mexico.
Vol. 21, Magnoliophyta: Asteridae (in part): Asteraceae. Oxford University Press, New York, NY. Pp 185-198.
Weakley, A.S., R J. LeBlond, B.A. Sorrie, C.T. Witsell, LD. Estes, K. Gandhi, K.G. Mathews, and A. Ebihara. 2011. New combinations,
rank changes, and nomenclatural and taxonomic comments in the vascular flora of the southeastern United States.
J. Bot. Res. Inst. Texas 5:439-440.
ARCEUTHOBIUM GILLII AND A. NIGRUM (VISCACEAE) REVISITED:
DISTRIBUTION, MORPHOLOGY, AND rDNA-ITS ANALYSIS
Shawn C. Kenaley
Department of Plant Pathology
and Plant-Microbe Biology
Cornell University
Ithaca, New York 14853, U.S.A.
sck26@cornell.edu
Robert L Mathiasen
School of Forestry
Northern Arizona University
Flagstaff, Arizona 8601 1, U.S.A.
Robert.Mathiasen@nau.edu
RESUMEN
le), was described in 1964
ujc w rsantalales ViscaceaeXwasdescnDeamiy
Gill’s dwarf mistletoe, Arceuthobium gilUi Hawksworth & ^lens ^ misdiaenosed as A. vagit
from southern Arizona (Hawksworth & Wiens 1964). Previous y, - gi (southwestern dwarf mist
^ (Will<fe„o„) Presl subsp, cryptopodum Arizona and northern Me
«)orA.divarica,nmEngeln«nn(pinyonp™dwrfm
«orA.driarica,n.Enge.n«nn(pinyonpinedwarf^^^^
• where these mistletoes are sympamc (Blumer W , physiologically from the latter species by its
giUii can be readily distinguished morpholog.«lly „f amhesis (Hawk-
skiwi color and sexual dimorphism, highly glaucous frui . P ^ ^ ^ Hawksworth & Wiens
Jl^th & Wiens 1964, 1996). Another similar dwarf ^ (Hawksworth & Wiens 1965)
(Wack dwarf mistletoe) which was initially described as a subspe , * , ^0,^ („ northern Mexi-
®l»thtaxaproducegUucou5fruits,sexuallydimorphicshoots.anas k
Journal of the Botanical Research Institute of Texas 7(1)
CO— Pinus leiophylla Schiede ex Schlechtendal & Chamisso, P. chihuahuana Engelmann, and P. lumholtzi Bl.
Robinson & Femald. However, A. nigrum was raised to the specific rank by Hawksworth and Wiens (1989)
based on morphological discontinuities with A. gillii; particularly, its larger and dark-green to black plants.
Moreover, A. nigrum reportedly has two flowering periods (March-April and September-October; Hawk-
sworth & Wiens 1989) while A. gillii flowers once annually (March-April; Hawksworth & Wiens 1996). Nev-
ertheless, these species remain difficult to distinguish from each other particularly in high-elevation pine for-
ests of central Durango, Mexico (Hawksworth & Wiens 1989, 1996; Mathiasen et al. 2008).
Because of the morphological similarities and host affinities of A. gillii and A. nigrum in northern Mexico,
the geographic distribution of these dwarf mistletoes has remained unclear (Mathiasen et al. 2003, 2008,
2010). We, therefore, have been gathering additional morphological measurements and phenological observa-
tions since 1999 from Mexico as well as southern Arizona. According to previous work by Nickrent et al. (1994,
2004), A. gillii and A. nigrum can be readily distinguished molecularly via DNA analysis of the nuclear ribo-
somal internal transcribed spacer (nrlTS) region. Therefore, we examined newly generated ITS sequences for
A. gillii and A. nigrum to better assess species boundaries and the geographic distribution of these mistletoes in
Mexico. Herein, we report the discriminatory morphological characters and phenology differences between A.
gillii and A. nigrum as well as our nrlTS analyses for both species across much of their geographic ranges.
To compare intra- and interspecific morphological characteristics of A. gillii and A. nigrum, we sampled 7
populations of A. gillii in southern Arizona and northern Mexico and 15 populations of A. nigrum from central
and northern Mexico (Fig. 1). Plants were also collected and measured from the type localities (populations)
for A. gillii and A. nigrum in Cochise County, Arizona, and Durango, Mexico, respectively (Hawksworth &
Wiens 1964, 1965) (Fig. 1; locations 3 and 9). Ten to twenty male and female plants were collected and the
dominant shoot from each plant was used for morphometric analyses. Plant characters measured were those
used previously by Hawksworth and Wiens (1996) for the taxonomic classification of Arceuthobium spp.: (D
height, basal diameter, third intemode length and width, and color of male and female plants; (2) mature fruit
length, width, and color; (3) seed length, width, and color; (4) length and width of staminate spikes; (5) stami-
nate flower diameters for 3- and 4-merous flowers as weU as the length and width of petals; and, (6) antherdi-
ameter and distance from the petal tip. Each plant was measured less than 24-h after collection using a digital
ca iper, a issectmg microscope with a micrometer, or with a Bausch and Lomb 7x hand lens equipped with a
m,crom«er. Stamtate spike and Dower measurements were made during peak anthesis, and fruit and seed
measurements were made during the height of seed dispersal. Because the seasonal occurrence of flowering
and st^d dlsper^l for A. gillii and A nigrum has received Uttle attention (Hawksworth & Wiens 1996), pheno-
ogical surveys for each taxon were conducted during the spring and fall of 1999, 2003, 2005, 2007, 2008, and
2010 as well as spring 2011.
A suite of multivariate and univariate sutistical tests were utilized to assess species differences, colkc-
tivelyartdseparately.among the 20 morphological characteristics measured forAgilliiandA.^
mulnvanate analysrs of variance (MANOVA) was performed to control for experiment error (lamily-«e
a^r ' d f of variance (ANOVA) was then used separately tf examine the «>•
ametn the mdrvtdual morphological characters of A giffiiandA. nigrum, and significant diHerences between
ired^ '“‘■>8 n posteriori contrast comparisons (u . 0.05). Morphological data were also ana-
lysed usmg dtscrtnnrtanr function analysis (DFA; Quinn & Keough 2006) to determine how well each olthe
d^ "fTTTd™ ** membetship-A, gillii or A rn-
gt^. in addttton, standaMtzed dircriminant function coefficients (DFC) were calculated to determine the
relative importance of each morphological characteristic as a discriminator between species. Univariate and
performed usingJMP 8 and JMP Pro 10 software (SAS Instiwl*
controls and blank reactions (i.e., minus genomic DNA) were also included in each PCR run as checks for
cross- and environmental contamination of template DNA, respectively.
Amplicon size for individual PCR products was checked via electrophoresis in 1.2% (wt/vol) agarose gels
followed by ethidium bromide staining (1.2 mg/L 0.5X Tris-acetate-EDTA [TAE]) and visualization under ul-
tra-violet huorescence. PCR productfs) of A. gillii and A. nigrum were purified using the Wizard® Preps PCR
DNA purification system (Promega) and the reagent ExoSAP-lT® (USB Inc.; 0.4 pL/pL of amplified DNA), re-
spectively Each amplicon was normalized (130 ng/reaction) and sequenced bidirectionally using an ABI 3730
DNA sequencer (Applied Biosystems), the referred primers (18s 1830 for and 26S 40rev), and a BigDye termina-
tors DNA sequencing kit (Applied Biosystems).
Phylogenetic Analyses and Variability of the ITS
ITS sequences were assembled and edited using CodonCode Aligner (CodonCode Corporation), and con-
firmed as belonging to the genus Arceuthobium by BLASTN and comparison (nucleotide identity) to authenti-
cated ITS sequences (Nickrent et al. 2004; Mathiasen et al. 2012). New DNA sequences generated for this study
were deposited in GenBank (accession no. KC543492-KC543496). For phylogenetic analyses, a dataset was
constructed consisting of the newly generated ITS sequences for A. gillii (KC543492-KC543494, KC543496)
and A. nigrum (KC543495) as well as previously published sequences of each taxon (A. gillii L25689; A. nigrum
AY288271, JQ723481-JQ723485, L23693) (Nickrent et al. 2004; Mathiasen et al. 2012). The dataset was then
complemented with ITS sequences (outgroups) of A. ooxacanum (AY288273) and A. yecorense (AY288288) -
sister taxa to A. gillii and A. nigrum (Nickrent et al. 2004). Sequences were aligned with the multiple sequence
alignment option implemented in ClustalXv2.1 (Larkin etal.2007),and the final alignment inspected in Co-
donCode Aligner. Maximum-likelihood (ML) analysis was performed using PAUP* 4.0bl0 (Swofford 2003)
and the best model of sequence evolution (Hasegawa-Kishino-Yano plus invariant sites [HKY-rll; Hasegawa et
al. 1985) selected by the Akaike Information Criterion (AIC; Akaike 1974) as implemented in jModeltest 0.1.1
(Posada 2008). Likelihood settings corresponding to the HKY+I model were as follows: Iset base = (0.3026
0T618 0.2074), nst = 2, tratio = 2.5765, rates = equal, and pinvar = 0.7820. All nucleotides were included in the
p y ogenetic analysis and gaps were treated as missing characters. The heuristic search was performed with
n bisection-reconnection (TBR) branch swapping, and
ULTREES in effect. Branch support was evaluated using 1000 bootstrap replicates and 100 RASper
pseudo-rephcate. ^
Bayesian inference of phylogeny via the Markov Chain Monte Cario (MCMC) method was also per-
formed usmg MrBayes 3.1.2 (Huelsenbeck & Ronquist 2001) as well as the best model and associated likeli-
o parametets executed in ML tree reconstruction. One cold and three heated Markov chainfs) were nm.
and samples were taken every 100 generations over 5.0 x 10« generations for a total of 30,000 sampled genera-
^ns. The potentral scale reduction factor (PSRF) for each ofthe model parameters was>1.0when the program
and l lTL d accessed by examining the avetage standard deviations of split frequetides
distamet^YJw““* ''‘‘"““‘'I' >TS for A. gillii and A. nigrum, mean pairwise genetic
ware IMP' Pro 10 M T “PUon 'n PAUP*and the statistical soft-
^ue^s wtthm and benveen spectes were also assessed in Getteiotts R6 (Biomattets Ltd.) and CodonCode
RESULTS AND DISCUSSION
Significant differences in morphology w
0.00579 (20, 179) = 1536.34; PeO.OOOl) with the f™ „
315
(Table 1). Discriminant function analysis indicated that the n
A. gillii and A. nigrum were the third internodal length of female plants (DFC = -7.97), basal diameter of male
(DFC = 5.53) and female plants (DFC = -5.03), and, the third internodal width of female (DFC = 3.95) and male
plants (DFC = 3.81). Other morphological characteristics such as the diameter of 4-merous flowers (DFC =
2.51), female plant height (DFC= -2.29), staminate spike length (DFC = -2.13), male plant height (DFC = -2.13),
seed length (DFC = -1.93), third internodal length of male plants (DFC = 1.68), and anther diameter (DFC = 1.0)
also significantly contributed to the discrimination of A. gillii and A. nigrum. The remaining flower, fruit, and
»ed charaaensrtcs contributed the least to the discriminant function CDFCe 11.01). Percent classifiottion to
the correct species using either a full-model (all 20 morphological characteristics) or reduced-model consist-
ingof the five, most discriminative characteristics described above was 100% (200/200).
Examining the morphometric results, male plants of A. gillii (mean = 12.3 cm) were significantly shorter
than those of A nigrum (mean = 24 3 cm). Similarly, female plant height of A. gillii (mean = 14.2) was also sig-
nificandy smaller than female plants of A. nigrum (mean . 19.6 cm)-apptoximately 5 cm shorter on average
The basal diameter and third internodal length and width of male and female plants, flower lameter an
1-meious), and the dimensions of fruit and seeds were also significantly smaller for A plitt when com[mred m
A nigrum (Table 1). Although mean staminate spike length was also significantly dtlferent between A gtllt.
ean =15.3 mm) and A. nigrum (:
= 20.6 ram), mean staminate spike width was 2.9 mm regardless of
edlyp™dr^™aXwo«hln?Wto(WMTm^^^^
centtalMelrirjScZTableVAlthoughthe^^^^^^
*rthatrfWlens(seeTablesland2forcompaj^nT^mt„„h ami Wiens. Yet, to date and to theb^^^
Ant^m, however, were shghtlysnuller ton those
out knowledge, the diametersofA-merous flowers, petal engthandw ^^^^^^^^^^^^
k»e not been reported. Similarly,Mathiasenetal. (2013^^
•oiu^esuchmeasurementotA-meromflmv^j^^uU^^^^^^^
Ourmeasurements of fruits were similar to the ^ ^ ^ measurements
otA. nigrum fruits, but our mean widths were larger (mea - _ ’ ^ 3 ^ ^han what they
Seated that A. gillii fruits are indeed larger (mean lengt = • > „ ,„ihose found by Hawksworth and
;;^rted;yet,seed"lengthandwtdthofAgi.-iiweresi«^^
(compare Tables 1 and 2). As for A. ^ '^3 j width = 1.5 mm).
^ths (1.3 mm) than our measurements (mean seed le g ^ diameter of 3-mer-
The only flower characteristic Hawksworth and Wiens „e mea-
-^ers (3.5 mm), which was slightly larger
e (mean - 32 mm). Our observations revra ^ averaged approximately 5 mm in itometer
well, thus, we sampled and measured these flowers ana ^ ^ significantly larger
(mean = 4.8; Table 1). The mean diameter of 3- and 4-merous ° ^ ^ nigrum were rela-
than those of A. gillii (3-merou5 mean = 2.8 mm, Wiens 1996; Mathiasen et al. 2012) as
‘^ely large in comparison to other dwarf mistletoes addition, examination of A. nigrum
^me lobes were >2 mm in length and nearly equally as larg phenotypic characteristic of this
flowers also revealed that the adaxial surface of petals were ar re
317
characteristic has gone unnoticed for nearly a half-century likely will
or-a phenotypic characteristic unaffected by plant age— should I
health specialist alike for differentiating A. nigrum and A. gillti.
lain unknown. However, petal col-
lized by field botanists and forest
According to our observatiom, flower phenology (period of anthesis)^^redJ»tob^add^^^
;^.icbet„eenA.gifliia„d..nigr„u..Insou.^r^^o^^^^^
arch, or as late as May m some years, and temiina y reported two flowering peri-
dually between September and January. 3,^„ding to our mcords and review of
“l^rch-April and September-October-for A. nigm .
^go, Mexico, in mid- to late-March 2003. 2005, and M7, and ,p.
owers were evident. Moreover, within the same loca ities an 19g9_2010, however, revealed
I»oaching maturity. Periodic fall surveys of the same A. nigrum November in Durango,
*^1 this dwarf mistletoe flowered starting m mid-Septem er sUghdy later (mid-Septem-
^exico,and peak flowering occurred in early-OctoberSumla^.-^^^^^
m central Mexico and continued into January; yet, t ^ indicated that A. nigrum began dispers-
*^ined uncertain. As with the advent of flowering, our ^ ^jd.November in Durango as well as
'«gseedinearly-Septemher,peaked in mid-October, and continued into
Journal of the Botanical Research Institute of Texas 7(1)
central Mexico. Likewise, seed dispersal by A. gillii initiated in e
These phenological observations for seed dispersal of A. gillii ar
ously reponed (Hawksworth & Wiens 1996); however, they i
PhytogenetKan3lysesof.helTSregio„fo,A.gimu„dA.„ig™msuppor,ed.hemorphologicaldau,eff^^^^
^imealmg secies « all planls diagnosed morphologicaUy as A. gillii resolved to a monophylelic group (Rg
4). T^e stze of the ITS region (ITS1-5.8S-ITS2) for each sequence of A. gillii and A. nigmn. was 600 bp whUe the
total amphcon per taxon consisted of 623 bp with a partial sequence of the 18S (4 bp) and 26S (19 bp) located
at the 5 - and 3 -end, respectively. The mean imerspecific genetic distance (HKYel) for both A. gfflii (n • W
pa.rw.se com,«nsons) ami A. nigntm (n . 21 pairwise comparisons) was 0.0018 substitutions/site. Similarly.
A. ttignim was 1.0 nucleotide. Although
related h loaned 11 T ■''anlts suggested that A. nigrum and A. gillii are more closely
Tnd LLu™ly“slLld!rir!j^ dentonstmted by Nkkrem et al. (2004). The alignment used lor ML
andA.mgrumu.ilLd in
L'n^ns~Cr”"'''TS’™“‘r P®‘“"y-W°™a.irThL°oL^^^^ MLnd Bay«iau
gly supported a single-clade (common ancestrv bootstran valnp 100% nnsterior probabil'
A A nigrum was 0.0083 substitutions/site
comparisons) and mean nucleotide divergence (n) between comparisons between A. gillii
4.9 nucleotides, respectively. Forty-eight of ^ and were separated by approxi-
A. nigrum shared 100% nucleotide identity (207/207 base ) consistent A/G nucleotide
"«y five or fewer substitutions across the 5.8S rRNA g® and ^
^"ges at positions 335. 421, 457, and 535. In „S sequences of A. gillii and A. nignm,
comparisons) and nucleotide divergence between our n ‘YE iUs,rnipsfFi2 2inNickrentetal.2004)was
^M41se,uence(GenBankL25693)utilizedbyNickrenta^jUra^ff^^
“1836 substitutions/site and 19.3 nucleotides, respecuvely. Tto ^ comparing the
«4 A. mgrum DLN 2041 were nearly equal to those cal^J^^se^
mterspecific genetic variability of outgroup taxa, A. yeco ^ potential hybrid — remains unclear and
“wcleotide difference). The identity of A. nigrum DLN 2041 - Therefore, the ITS sequences of A.
currently is being assessed (S.C. Kenaley & R-L. Mathiasen, p
nigrum presented herein, as well as in Mathiasen et al. (2012), should be utilized for future species-level identi-
fication in lieu of A. nigrum DLN 2041. Moreover, additional phylogenetically informative molecular markers
(e.g., single-copy nuclear genes; Duarte et al. 2010) and multigene analysis should be examined to further re-
solve the genetic relationships of A. gillii and A. nigrum.
Hosts
Our examination of host affinities for A. gillii and A. nigrum did not reveal any additional host-mistletoe com-
binations. Principal hosts of A. gillii included those already reported by Hawksworth and Wiens (1964, 1965,
1989, 1996) such as Pinus chihuahuana, P. leiophylla, and P. lumholtzii. Arceuthobium nigrum in Durango was
most often found parasitizing P. chihuahuana, P. leiophylla, and P. lumholtzii, as well as P. teocote Schiede ex
Schlechtendal & Chamisso; all pines were described previously as principal hosts (Hawksworth & Wiens
1996). Although P. arizonica Engelmann and P. cooperi Blanco are reported rare hosts of A. nigrum in northern
Mexico (Hawksworth & Wiens 1996), we did not observe either host-mistletoe combination during the course
of our work in that region of Mexico. However in central Mexico, A. nigrum was most frequently encountered
on P. teocote, and at several localities in Hildago and Puebla, we often found P. patula Schiede ex Schlechtendal
& Chamisso as a secondary host. Arceuthobium nigrum was also found on P. pseudostrobus Lindley - an occa-
sional host (Hawksworth & Wiens 1996); however, other occasional hosts of A. nigrum such as P. montezumae
A. B. Lambert, were not observed. This was disappointing as we purposefully examined forested sites in the
States of Hidalgo and Veracruz where Hawksworth and Wiens purportedly found A. nigrum infecting P. mont-
ezumae, finding only P. teocote severely infected with A. nigrum. The status of P. montezumae as a principal host
of A. nigrum, therefore, requires future examination and confirmation. Similarly, Hawksworth and Wiens
(1989, 1996) reported A. nigrum on P. lawsonii Roezl ex Gordon & Glendinning and P. oaxacana Mirovinthe
uthem Mexico
m^m is primarily distributed geographically along the Central Volcanic Cordillera of central
Mexico and north into Durango (Mathiasen et al. 2010, 2012). Although Hawksworth and Wiens (1989, 1996)
reported tha.Ag,iI.. andAmgrumoccurlnthe same mountainrange near Tepehuanes, Durango, our obsova-
w” l“:alions examined by Hawksworth in 1987 (Hawksworth S
Wtens 1996), did not support this report. All of the populations we examined in the Tepehuanes area repre-
antedAntgnim^Theclosesl, and thus far, only population ofAgillii we found to date in Durango wasnear la
Quebrada tmt^tately south of the border with the State of Cbihuahua (Fig 1 location 7). Ttwrefore, du
common dwarf mistletoe on P. leiophylla and P chihuoh^tna in Durango is A nigrtun while A giDii was the
it fmquent dwarf mistletoe on these pines in Chihuahua; we have L obser^d A uigrum in Chihuahw.
^ -'hi may be found to be sympatric with A. nigrum in Durango, we have not
3r general location.
The principal morphological and phenological differences dictinoi.i i, a ti r a ^imaresuoi-
Seed Dispersal
10.6(16.7)
13.8(223)
33(5.0)
4.2 (6.6)
243(533)
19.6(373)
6.9 (8.8)
a.samehos..i„nonhernMexico.However..hemeansizesofaU*emorphdogicalcha.^c.ersweex™med
txcep, suminale spike width, ate significantly smalkt for gtlW, which has green, not red tlow^ (compare
F«c2and3), and flowers in the spring and not the fall. Themfore, these ta«c» be dtanrgut^ro^^^^^^
other by plant size, flower size and color, fruit size, and flowering period (Table 3) and stanid be treated as
distinct species. Difficulties with identifying them in the field will be related pnman y to
habit and plant color, so it is necessary to examine many plants from a population, de erm p , ,
and fruit dimensions for at least a small sample of these parameters and observe J ° ,
-aleplantsare flowering, then the color oftheflowers(greenorred)andseasonTO
identify if the nonulaiinn renresents A eilUi or A. nigrum in northern Mexico where these species may co-
occuXd„nTcrXlledgeof.heirdis.ribnnon.^
^^Mexicosouibrofarnorrhern Durango, Mem»(Fi^a^^
Tepehuanes south as far as Puebla, Tlaxcala, and Veracruz (Mattiiasen herbaria records
he noted that most of the populations of A. gilUi reported by Mathiasen et a .
from southern and central Durango probably represent populations of A. nigrum.
acknowledgments
'‘'ewonldUkerorhankBrianP.ReifforperformingtoDNAe^^^^^^
^reaate the help of Gustavo Perez and Jeremy R. Thompson sincerely appreciated and helped
ReviewsofthemanuscriptbySocorroGonz^ezElizondoandGregFihparesincerely pp
in greatly improve the paper.
. Autom. Contr. 19:716-723.
322
; and Oryza and their phylogenetic utility across various
taxonomic levels. BMC Evol. Biol. 10:61.
Hawksworth, F.G. and D. Wiens. 1 964. A new species of Arceuthobium from Arizona. Brittonia 1 6:54-57.
Hawksworth, F.G. and D. Wiens. 1965. Arceuthobium in Mexico. Brittonia 17:213-238.
Hawksworth, F.G. and D. Wiens. 1 972. Biology and classification of dwarf mistletoes (Arceuthobium). Agriculture Handbook
401 , U5DA Forest Service, Washington, D.C.
Hawksworth, F.G. and D. Wiens. 1989. Two new species, nomenclatural changes, and range extensions in Mexican Arceu-
thobium (Viscaceae). Phytologia 66:5-1 1.
Hawksworth, F.G. and D. Wiens. 1996. Dwarf mistletoes: biology, pathology, and systematics. Agriculture Handbook 709,
USDA Forest Service, Washington, D.C.
Hasegawa, M., H. Kishino, and T. Yano. 1 985. Dating of the human-ape splitting by a molecular clock of mitochondrial DNA.
Molec. Evol. 22:160-174.
Huelsenbeck, J.P. and F. Ronquist. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinform. Appl. Note 17:
754-755.
Larkin, M.A., G. Blackshields, N.P. Brown, R. Chenna, PA. McGettigan, H. McWiluam, TJ. Gibson, and D.G. Higgins. 2007. Clustal W
and Clustal X version 2.0. Bioinform. 23:2947-2948.
Mathiasen, R.L., C.M. Daugherty, and V. Guerra De la Cruz. 201 0. Muerdagos enanos (Arceuthobium spp.) en el sur de Mexico:
Distribucion, hospederos y cambios en la nomenclatura. In: Proceedings of the XV Simposio Nacional de Parasitolo-
gia Forestal, November 18-20, 2009, Oaxaca, Mexico. Pp. 160-166.
Mathiasen, R.L., M.C. GonzAlez Euzondo, M. Gonzalez Elizondo, B.E Howell, I.L LOpez EnriqueZ, J. Scott, and J A Tena Flores. 2008.
Distribution of dwarf mistletoes (Arceuthobium spp., Viscaceae) in Durango, Mexico. Madrono 55:161-169.
Mathiasen, R.L, S.C. Kenaley, and B.P. Reif. 201 2. Distribution and morphological characteristics of Arceuthobium hondu-
rense and A. nigrum (Viscaceae) in Mexico. J. Bot. Res. Inst. Texas 6: 599-609.
Mathiasen, R.L, J. Meigar, J. Beatty, C. Parks, D.L Nickrent, S. Sesnie, C. Daugherty, B. Howell, and G. Garnett. 2003. New distri-
butions and hosts for mistletoes parasitizing pines in southern Mexico and Central America. Madrono 50:1 15-121.
Nickrent, D.L., M.A. GarcIa, M.P. Martin, and R.L Mathiasen. 2004. A phylogeny of all species of Arceuthobium (Viscaceae)
using nuclear and chloroplast DNA sequences. Amer. J. Bot. 91 :125-1 38.
A molecular phylogeny of Arceuthobium (Viscaceae) based on nudear
I.Bot.81:1149-1160.
c. Biol. Evol. 25:1253-1256.
C3ulNN,G.P.A^
r 1 3, 2009. MCMC Trace File Analyser, h
in. John Wiley an
SwoFFORD, D.L. 2003. PAUP*. Phylogenetic analysis using parsimony (*and other methods). Version 4. 5inauer Associates,
THE GENUS PLATANTHERA (ORCHIDACEAE) IN MISSISSIPPI
Michael Wayne Morris
Troy University
ABSTRACT
RESUMEN
1 1978), with approximately 20,000
s35,000 species (Dressier 1981J993). The family is moslspecioseinthe
topics, and the majority of taxa in this fam
The Orchidaceae is o
topics, and the majoi ilj' a.* y - -
Wi^sirialspeciesdisiributedprimarilyirithecir^mbomaU^^^^
^e.Species of P,a.a„.l.raco...rr.only lave either
late to oblanceolate leaves that become reduced to bracts art er p
«tep^au'^r“'”Tn“!^rSM™her™mmname fringed oichids, although they may^^
Nectarsputsareat flower bases. Among the speaes m the rilmris and P. inlcgrilaWo (Zetder et
® P clavellata (Luer 1975), to insect pollination involving u e ■ ^y.^y^ng sphinx moths in P.
"T 1996), butterflies, moths, and bees in P.Wephanglottis mi u„a«WpWs in P cristota and P Integra,
^WephariglottisfSmith&Snowl
m neman & Inoue 1997), bumble bees u
999) and P peramoena ( ape . ■ greatly influences phylogenetic rela
‘“‘Q even mosquitoes in P. ftava (Luer 1975). Reproduct ^
*^)s within Platanthera (Bateman et al. 2009; Hapeman & lnouel9y / J-
Sphagnum bogs, bay swamps and pitcher plant savannas, mesic field/woodland ecotones, and open, upland
pine woods with perched water tables. Disturbed habitats include highway medians, powerline rights-of-way,
and gas pipelines running through wetlands. Regarding the latter, the periodic mowing/bush hogging likely
promotes growth of fire-adapted Platanthera species needing high light intensities by removing encroaching
trees and shrubs. These practices benefit the orchids if done at the right time of year, if herbicides are not
sprayed, and if the ground is not mechanically disturbed. Several Platanthera species are of conservation con-
cern and are tracked by the Mississippi Natural Heritage Program (2006); they are as follows: P. conspicua (= P.
blephariglottis var. compicua) (S2), P. cristata (S3), P. Integra (S3), P integrilabia (SI), P lacera (S1S2), and P. per-
amoena (S2S3). Of these, two species have moderate to high priority global conservation rankings: P. Integra
(G3G4) and P integrilabia (G2G3). This study was conducted because a floristic manual for Mississippi does
not exist. Therefore, treatments of genera and families are needed as published journal articles at this time.
Dunngjuly, 2009 and July, 2010, when plants were at anthesis, 14 trips were made to locate extant populations
of Platanthera species, record habitat data, and visit the major herbaria in Mississippi. These excursions were
made to characterize specific habitat requirements of Platanthera species among the different physiographic
regions in Mississippi. Herbarium visits were conducted to accurately determine county-by-county distribu-
tion among the physiographic regions in Mississippi and to make morphological measurements. Platanthera
specimens m the following herbaria were examined: University of Mississippi (MISS), Mississippi State
University (MISSA), University of Southern Mississippi (USMS), Delta State University (DSC), Institute for
Botanical Exploration (IBE), Mississippi Museum of Natural Science (MMNS), The Crosby Arboretum, and my
per^nal collections. Representative specimens examined are listed at the end of each treatment for the spedes
in Missisippi. Matenal of P.Jlava deposited in MISS was on loan to Arkansas Tech at the time of this study. The
fdlowingso»rc«„l^„, for, he non. of the 50u.heastemUnUedSta.es were considered forsyn^^
^r. Clewell (1985), Godfrey and Wooten (1979), Luer ( 1972, 1975), Radford et al. (1968), Sheviak (2002).
The International Plant Names Index (2012), Weakley (2012), Wofford (1989), and Wunderlin (1998).
KtSULIS AND DISCUSSION
Ten species of Platanthera an. known to occur in Mississippi, a state with ten physionranhic regions (Lowe
yiava, P, tntegra, P. tntegrtlahia, P, lacera, P. nivea, and P. peran«na. Distribution of each taxon in Mississippi
widely distributed in the state Plat^ZZlrTu T
d hardwood forests at the easternmost edge of the latter. While
cilicris appears to be absent fmn. circum^nf f Coastal Pine Meadows, P
Ridve RlarW Rplf iinei r i d ^ and heavy clay soils characteristic of the Pontotoc
Ridge^lack Bek andJacksonPra.nes,L^ Bluffs, and Yazoo-Missiilppklm. Despite havingabrcu^
t inhabits shrub
flava is known from s'
nth a partially open canopy, and moist to we. ffatwoods.
P. peromoena. Both of the latter taxa snom^HuJl ““t*' ^ i"K«rilal.ia, P. lacera. and
meadows and woodland borders in CenTl^r documented from
River HUlsandLongleafPine Belt, The P”H«ions in the Tennessee
and adiacenr meadowc in in 7 ° P^^oena records are from bottomland hardwood forests
and adjacemmeadowsmiwer and creek ffoodplainsintheNorth Central Pla.eauandar.as transitional tothe
IS are also supported in the Ten-
c Ridge in the northeastern
i in the Jackson Prairie and sites
e Longleaf Pine Belt in central Missis-
mong the southernmost populations in
the United States. Platanthera integrilabia is the rarest and
most narrowly distributed of the fringed orchids in Mis-
sissippi and is only known from shaded wetland habitats
of the Tennessee River Hills.
Platanthera conspicua, P. Integra, and P. nivea are
all characteristic of open pitcher plant bogs and savannas
in the Longleaf Pine Belt and Coastal Pine Meadows of
southern Mississippi. However, P. nivea is the only taxon
not considered rare in the region; and it can even occur
at sites transitional to sandhiU habitats and infrequently
rights-of-way. All three orchids typically inhabit fire-
prone wetland sites with soils deficient in nitrogen in as-
sociation with species such as the following: Alctris lutea,
Aristida beyrichiana, Carphephorus pseudo-liatris, Erio-
j Uliumcatesbaei, Lophiolaaurea, Magnolia virginiana,
Nyssa biflora, Persea palustris, PinuselUomU P palusjris,
Ae genera Cyperus, Dicanthelium, Drosera. Hypericum,
I J ^ ■ Cl, rhn<nnrn Sobatia Sarracenio, Utricularia. and Xyris. Associated
^.pa, PotygM.^ c C t^osus. Clds,« h/a™, Pc^oniaopkio^-
praecox (Morris 1989).
GENUS DESCRIPTION AND KEY TO THE SPECIES IN MISSISSIPPI
Platanthera Rich. , FRINGED ORCHIDS
Glabrous erect perennial herbs with tuberous, fusiform, (
wetland habitats. Leaves simple, alternate, mostly ra«mo“si"Flowe/s white or
becoming reduced bracts toward the top of the plane " ^ ^he column; lateral
peenish toyellow, orange, or purple; Up lowermost (uppermost in P nivea), entire,
«Pals spreading or recurved; petals free, simple or p
divided, or fringed, base of labellum modified into saccate to ^ ^-celled; pollinia 2, attached
pedicel; column short; stigma with or without papillose process,
to partially to fully exposed viscid gland. Fruit a cylindric to ^ constructed based on ob-
An artificial dichotomous key to the ten Platanthera ^ presented below.
Nations of both herbarium specimens and living plants m the neiQ,
KEY TO PLATANTHERA SPECIES IN M
MISSISSIPPI
Journal of the Botanical Research Institute of Texas 7(1)
Plant erect, often robust, to 100 cm tall. Leaves mostly basal, lanceolate, keeled, reduced and becoming bracts
above, 15-30 cm long, 15-30 mm wide. Raceme densely or laxly flowered, bracteate, 5-17 cm long, 4.5-9 cm
wide. Flowers 15-25 mm long, yellow-orange to orange; sepals ovate, 7-8 mm long, 6-7 mm wide, dorsal con-
cave, lateral reflexed; petals linear, apex fringed, 5-6 mm long, 1-1.5 mm wide; lip oblong, 10-16 mm long,
mm wde, conspicuously ciliate-fringed; nectar spur slender, 2-3.5 cm long. Jul-mid-Aug(-early Sep).
Open to shaded bogs and seepage slopes, wet fields and savannas, seepy roadcuts and adjacent ditches, and
o^, upland pine woods. Coastal Pine Meadows, Longleaf Pine Belt, North Central Plateau, Interior Flat-
woods, and Tennessee River Hills. Infrequent to occasional (MA to Ont., Ml, and IL, s to FL w to e TX and OK).
Among Plotonthera species in Mississippi, P. ciliaris has the most varied habitat preferences. In southern
Mississippi, It frequently occurs m pitcher plant bogs and savannas with the same species listed above for R
conspicua, P Integra, and P nivea. In Grenada County, located in the north central part of the state, P dliaris is
ericountere in boggy springheads with much Sphagnum and in adjacent open, sandy seepage areas with the
o owing taxa: Amoglos^movatum, Carex atlantica, C. crinita, C. laevivaginata, C styloflexa, Chelone glabra,
Doellmgena sencocarpoides, Drosera brevifolia, Eryngium integrifolium, Gentiana saponaria Hex verticillata, Im-
LyccpoSclb appre^ Oromium a,pa,icum, Osm.Ma dn«a« 0. m
PhciMa pyn/ollo, Rhe^a nrginica. Rkododaulran cmescens.
^P'^‘‘gl<^"‘-‘‘‘.Sol:dagopatulaJphe^ispcnsyIyanica.SymphycRrichu^^
mun,un, ™dum Wood^arJR, arcolata. and Xyns ,„na (Morris 1988). Conrpanion orchid spec
Platantheraclavellata and Spiranthescemuua.
Gore Springs, (T22N, R
Moms), Hancock Ok Hillside Bog Natural Area of th. , , „
dens-Nyssa biflora, ^ Aug 1986, Morris 2423 (herbarium of the Oosby A
mi SE of intersection of hwy 35 and Natchez Trace Parkway,
I, Wieland 6550 (MMNS). Grenada Co.: ca. 2.5 miNNWof
;9Jul 1985, Morris 405 (mwm, pers. herb, of Michael Wayne
co tone of open bog and Magnolia virgini
oretum), Jackson Co.: ca. 2.0 mi N of Escatawpa oi
a, frequent, 8 Sept 1983, Norquist 1396 (MMNS). M«
m of Hwy, 25 and Hwy. 19, r<
inEsideof
. Platanthera clavellata (Michx.) Luer (Figs. 4, 5, ]
Plant erect, small, to 40 cm tall. Leaves mostly basal, usually one
becoming widely spaced bracts above, 8.5-14.5 cm long, 12-28 i
Journal of the Botanical Research Institute of Texas 7(1)
MofTis,Platanthera in Mississippi
a (Michx.) Lindl. (Fi^
•w « (M h)RB Orh stata Michx Platanthera pallida P.M. Brown.
^SSas pipelin^ p„^,U„e ri^.s-of-way. Coasu. Pine
^«»J«erto,Fla™oo<U,andTennesaeeRiverHills.fc.re.o.nfre,uen.(M^
Sute conservation status: S3.
Journal of the Botanical Research Institute of Texas 7(1)
Fk. 9. Known distribution of Platanthera cristata in Mississippi.
Habitats supporting P. cristata populations in Mis-
sissippi are mostly intermediate between more open
sites in which P. ciliaris is most often found and more
shaded sites in which P. clavellata occurs; however, P-
cristata may occur with both fringed orchid species
spring branches and more open grass/sedge meadows at
the edges of beaver ponds or along powerline rights-of-
way running through bay swamp communities. Here,
associated species include many of those listed for P. Cl-
aris, along with the following taxa: Aletris aurea, Barto-
nia paniculata, Carexglaucescens, Carex leptalea, Cypcrus
haspan, Drosera capillaris, Elephantopus nudatus, Eupato-
riadelphus fistulosus, Fuirena squarrosa, Hydrolea ovatu,
Hypericum crux-andreae, Hypericum gymnanthum, Ledi-
ea minor, Lespedeza capitata, Liatris spicata, Ludwigi^
hirtella, Lycopodiella alopecuroides, Lyonia ligustrina, Mi-
la, Rhynchospora inexpansa, and Sophronanthe pilo^
treola petiolata, Plucheafoetida, Polygala ,
Hexmyrtifolia, Rliciumjloridanum, Kalmia latifolia, Leuc(f
’ ■ 'soides, Rhododendron viscosum, Smilaxlaurifol^
arundinaceum, Hypericum galioides
23Jul 1981, Gordon 2567 (MMNS). Grenada Co.: ca. 4.5 mi SSE of Gore Springs, (T21N. R7 , ^ . i" e
W.M..rt.2«®E,™,.Ja.l».„C..:RcdC,..kMm*a.,onB.nkEotH»y57.m™2,«m^^^
uplands, 25 Jul 2006, Sullivan 06-1422 (MMNS). Monroe Co.: ca. 5 mi N of Aberdeen, (T14S,
lake,lAugl995,MacDonaId9023(MMNS).Ok,ibbehaCo.:ca.0.5miNofCedarGroveRd
age. fUtwoods, 19 Jul 1994, Liedolf 0526 (MMNS). Perry Co.: DeSoto National Forest, ea
^l.,».,.uck,k,„gs„».I9J„119nW,hck94-120MWa^
5. PUunthera fla't
indl. (Figs. 10-12). southern rein-orchid
HabenariaJIava (L.) R. Brown, Orchis flava U Perularia scutellata (Nutt.) Small
20-50 mm wide. Spike loosed, flowered, bmcteate, 7-19.5 cm
'®g. yellow-green; dorsal sepal ovale, concave, 2-4 mm o g, rounded, the base with
®«long,2-3mrnwide;petalsovate,concave.2-4mn.l™frM— ap^^
> itiangnlar lobe on each side. 5 mm long, 4 mm »>*, ^ bases of bluffs, and along braided
“h roX» CoaTal Pine Meadows, Longleal Pine Belt, North Cen-
Streams m dense cypress— hardwood swamp forests. C Flatwoods. Pontotoc Ridge, and
ttal Plateau, Loess Bluff/Yazoo-Mississippi ^ , TX and lA). Platantherafiava (L.) Lindl.
Tennessee River Hills. Infrequent (NS to Ont. and MN, s to c re,
'^r.jlavainMSnearcentralpartofrange. here this orchid grows. Plants tend to
Mosquitoes, the pollinating agents of P.Jlava (Luer 1975), ^ ^ channels filled with muck,
grow along the banks of braided streams in colluvial swamp o ’ . . ■ ^ p is not common. Asso-
^ttd at the bases of large bald cypress trees. proximity to the Yazoo-Mississippi Delta in
species in a swamp forest located in the Loess Blu Carpinus caroliniana, Cicuta maculata,
=«nada County .re: Acer ntbrum, Asimina trilobu, Itenvirginica, Undemben-
Cwns/oeminu. Dasistomtt mocrnpliyllum, Dtyoplerrs vtttutrttlci, Impanenr up
zoin. Lobelia cardinalis, Osmunda regalis, Platanthera
clavellata, Populus heterophylla, Rhynchospora miliacea,
Rudbeckiafulgida, Sabal minor, Saururus cemuus, Soti-
dago patula, Taxodium distichum, and Thelypteris
PUm erect to 65 cm taU. Leaves mostly basal, lanceolate, keeled, conduplicate, reduced and becoming btacts
above 5.5-19^1ong,^17mm unde. Raceme cylindrical, denselynowered,bracteate, 3-6 cm long, 2.5-3®
^e,Howers5-7mmlong,5-6mmwide.yellow-orange;sepalsobovatetoorbiculatdorealsepal4mmlong.
3tnm unde, con^ve, lateral sepals5mmlong,4mm Wide, oblique;petaUelliptic,4mm long, 1.5 mm «de;lip
^ ^ nectar spur 4-6 mm long. Jul-mid-
to n FL, » to e TX; TN; primarily coastaO. Global and ^'^™ion Mms'
333
Ptaheraimegraisfoundin pine savannas, pUcherpk„.bogs,and^
«g»winginassociationwi.hPin,egmani.hoselis«daboved.aiarealsotaowntogroww,,hP.ccm^^^
andP.nivea.
j s of Agricda School, 30.7458, -88.4986, large quaking bog
aide Bog Natural Area of the Crosby Aiboretum,
fiabenana blephariehttis (Willd.) Hook. var. infegriloma correu, . ..
PUn, , U 1 iHntir to lanceolate, keeled, reduced and becoming bracts
erect, to 60 cm tall. Leaves mostly basal, ' l'P'7‘’,'“““_(i^ered, braceate, 6.5-7.5 cm long,
^e, 20-23 cm long, 40-42 mm vvide. R-me lo^j broadly ovate, dorsal
5 cm wide. Flowers 10-12 mm long, 9-10 mm wide, wm , ^
^Pal 8 mm long, 6 mm wide, concave, lateral sepals 9 mm serrulate; nectar spur slen-
2®l«ng, 2.5 mm wide; lip spatulate-lanceolate^ ^^T.^n'^bogs in dense woods. Tennessee River Hills. Rare
«ate conservation status: G2G3, SI.
umal of the Botanical Research Institute of Texas 7(1)
336
spatulate to oblong-linear, denticulate, 4-8 mm long,
3-6 mm wide; lip tripartite, narrowed at base, cuneate,
middle lobe with central notch, 15 mm long, 12 mm
wide; nectar spur slender, club-shaped: curved, 22-25
mm long. Late June-mid-July. Bottomland hardwoods
and adjacent meadows in creek and river floodplains,
overgrown pastures, and grassy ditch/pine plantation
ecotones. Longleaf Pine Belt/Jackson Pr— =
areas, Jackson Prairie, North Central
Bluffs, Pontotoc Ridge. ’ "
(NJandPAtosILand
nARlPlatantherapen
in MS. State conservation status: S2S3.
A spectacular plant, P. peramoena sporadically
occurs in northern and central Mississippi. The habitat
® Attala County is a rich deciduous hardwood forest along a creek. superbum, Liquil
‘^‘r jhndanum. Arisaam, dracontium. A. , LoeK^uffs^onh Central Plateau
nyrociflun. Quercus utkhauxii, Q. pagoda, and ® ^^„^l,ttd hardwoods ecotone in the
transition area in Grenada County, P. peramoena is in a wet m ^ ^ ^
Y^busha River floodplain with Acer rubrum, Sites supporting populations of this rare
difficult to determine specific habitat requirements for P. peramoen . PP®
'P'tiesareoftennondescript. . , ,„,™r„,n»u.u™ck.<,nE«l.oIsh.HwyC,e.k. I7jd
L MISSISSIPPI- AtlaT
338
Journal of the Botanical Research Institute of Texas 7(1)
-urators ot the lollowing herbaria are grateiuiiy
Kiiowledged: DSC, IBE, MISS, MISSA, MMNS, USMS,
and The Crosby Arboretum. Data from the Mississippi
■ ,vided by Heather
images of
Natural Heritage Program i
neruage t'rogram were proviaea uy iivK— -
Sullivan. Additional input from Mac Alford is appreciated. Thanks to John Gwaltney for digital images of Plflt-
flaya, P. integnlabia, P. nivea, and P. peramoena. Additionally, I am grateful to Robert Stew-
MriTTlC ^ . 1
mthera conspicua, P. flava
■ C B ,r me on several field tripsin search 01
cies. Sam Faulkner is acknowledged for his assistance. Special thanks to Wayne and Lynda Edwards, and
Robert and She.^ Morris for their assistance with this project. I gratefully acknowledge Heather Sulli«n
Tl uTr Findewer for their helpful comments and suggestions. Appreciation a
extendi toJuheBatharee, Murray Hyde,and Glenn Cohen, Department of BiologicalandEnvironmentalSci
ences University, for technical assismnce. This research was partially supported by a Troy UnivcrriD
Faculty Development Summer Research Grant. / t't' y
339
REFERENCES
Bateman, R.M., K.E. Jones, Y.-B. Luo, R.K. Lauri, T. Fulcher, PJ. Cribb, and M.W. Chase. 2009. Molecular phylogenetics and mor-
phological reappraisal of the Platanthera clade (Orchidaceae; Orchidinae) prompts expansion of the generic limits of
Galearis and Platanthera. Ann. Bot. 104(3):43 1-445.
Clewell, A.F. 1 985. Guide to the vascular plants of the Florida panhandle. Florida State University Press, Tallahassee.
Correu, D.S. 1 978. Native orchids of North America north of Mexico. Stanford University Press, Stanford, CA.
Dressier, R.L. 1981 .The orchids: natural history and classification. Harvard University Press, Cambridge, MA.
Dressier, R.L. 1993. Phylogeny and Classification of the Orchid Family. Dioscorides Press, Portland, OR.
Godfrey, R.K. and J.W. Wooten. 1 979. Aquatic and wetland plants of southeastern United States. Monocotyledons. Vol. 1 .
The University of Georgia Press, Athens.
Hapeman, j.R. AND K. Inoue. 1 997. Plant-pollinator interactions and floral radiation in Platanthera (Orchidaceae). In: TJ.
Givnish and KJ. Sytsma, eds. Molecular evolution and adaptive radiation. Cambridge University Press, Cambridge.
LmvE, E.N. 1921. Plants of Mississippi, a list of flowering plants and ferns. Mississippi Geological Survey Bulletin 17. He-
derman Brothers, Jackson, MS.
Luer, CA. 1 972. The native orchids of Florida. The New York Botanical Garden, Bronx.
Luer, CA. 1975.The native orchids of the United States and Canada, Excluding Florida.The New York Botanical Garden,
Mississippi Automated Resource Information Systems (maris). 2013. Accessed at http-y/i«ww.maris.state.ms.us/.
Mississippi Natural Heritage Program. 2006. Special plants— tracking list. Accessed at http-y/
loads/science
Mor«s, M.W. 1988. Noteworthy vascular plants from Grenada County, Mississippi. Sida 13:177-186.
>hm. M.W. 1989. Spiramhes (Orchidaceae) in Mississippi. Selbyana 1 1:39-18.
»l«w, M.W. 1997. Contributions to the flora and ecoiogy of the northern longieaf P.ne Beit .n Rankin County, Missis
Iw!“h!e1^^,"1c.R.Beu.1968. Manual of die eascubrflora of the Carolinas.The University of North Ca™^^^^
Swrw, CJ. 2002. Platanthera Richard. In; Flora of North America Editorial CommitM, ^s. Flora of North Amenca north
). Pollination ecology o
d 10 May 201 3].
s. November 201 2 Version. University of North C
Smith, G.R. and G.E. Snow. 19:
Bot. Gaz. 137:133-140.
The International Plant Names Index. 201 2. Published on the Internet httpy/ipni.org [acces
USDA Plants Database. 201 3. Accessed at httpy/plants.usda.gov/
United States Geological Survey. 201 3. Accessed at httpy/pubs.usgs.gov/
Weakley, A.S. 201 2. Flora of the southern and mid-Adar
Wofford, B.E. 1 989. Guide to the vascular plants of the Blue Ridge. The Universi^
BOOK REVIEW
RT B. Shaw. 2012. Guide to Texas Grasses. (ISBN; 978-1603441865, flexbound). University of Texas A&M
Press, John H. Undsey Building, Lewis Street, 4354 TAMU, College Station, Texas 77843-4354, U.S.A.
(Orders: www.tamupress.com, 1-800-826-8911, 1-979-847-8752 fax). $45.00, 1096 pp., 1357colorpho-
tos, 34 b/w photos, 947 line drawings, 645 maps, 6.6 lbs, 7" x 10".
Table of Contents:
Preface
Why Grasses Are Important
The Grass Plant
Ecoregions of Texas
Classification of the Grasses of Texas
Generic Keys and Species Accounts
Appendix: Collection, Preparation, Handling, and Storage of Grass Specimens
Glossary
Index
Guide to Texas Grasses is a 7" x 10" flexbound text, consisting of 1096 very thick pages. The text contains nu-
merous color photographs and line drawings for about 670 grasses, both native and ornamental, found in
Texas. It is part of the ongoing Texas A&M Nature Guides series edited by Craig Nessler, director of Texas
AgriUfe Research, and Edward G. Smith, director of the Texas AgriLife Extension Service.
Chapter 1, Why Grasses Are Important, provides a brief summary of the family and discusses the eco-
nomic importance and uses of grasses such as food, forage, turf and ornamental uses, and biofu-'-
also touches on grasses often overlooked relative to their environmental importance for soil cc
their impact as invasive species.
Chapter 2, The Grass Mant, discusses the morphology of grasses and provides very clear drawings anu
imaapc nf r,l=,nt parts. Great drawings, many from Barkworth et al. (2003), are included detailing inflorescence
iictures, florets, spikelets, and disarticulation lines. The most detail in spikelets is shown for the
s. The chapter
ration and
Paniceae and Andropogoneae.
Chapter 3, Ecoregions ofTexas, briefly discusses the geography of the state anditsllecoregions, which
; defined as regions with homogenous natural features. The rhantPr »He complex geology ofTexas
j. The chapter also discusses
d vegetation types.
r , ui iciiy Ulscusses tne geography ot the «
are defined as regions with homogenous natural features. The chapter discus;
on a broad geologic timescale and how the distinct units were formed over l
how physiography, topogiaphy. and climate helped determine the different e^oregions'and vegetation types
^ik am discussedatthe level of soil order. Vegetation areas, naturalregions, and land uses are then diseased
based on the previously mentionedabioticfactors.Amom detailed disenssionthen begins withagooddetailed
summary of the 12 ecoregions, their location, dominant species, elevaUon, geology, soik, climate, and major
land use functions. Alist of uxa by grass subfamilies is also provided for eachecoregion.
e Grasses of Texas, discusses the different subfamilies of the grass fam-
, and primarily discusses the different taxonomic classifications and
Keys and Species Accounts, are where this book shines A key to groups is provided at
thestartofthesecnon,modelingtheIIIustratedHorao/NorthCentralTexas(Diggsetal 1999). Each group then
am mcluded to faedttate guidance. For each genus, a circumscription k provided as well as keys to species
J. B«t Res. Inn. Teas 7(1): 340. 20
A NEW SPECIES OF CEROPEGIA (APOCYNACEAE: ASCLEPIADOIDEAE)
FROM INDIA WITH NOTES ON RARE AND THREATENED
CEROPEGIA IN NILGIRIS OF WESTERN GHATS
P. Sujanapal
Kerala Forest Research Institute
Peechi, Thrissur - 680 653, INDIA
PM. Salim and N. Anil Kumar
MS Swaminathan Research Foundation
Puthoorvayal, Kalpetta - 673 111. INDIA
N. Sasidharan
Kerala Forest Research Institute
Peechi, Thrissur -680653. INDIA
uing studies in the genus Ceropegia in Nilgiri ph
manoharii Sujanapal, Salim, Anil & . . ju i nt hi
al study has shed light on the details of three edible Cempegia species, as hitherto undescribed host plants ol a b.
RESUMEN
INTRODUCTION
Nilgiris, or Blue Mountains, the meeting place of three mountain systems of a^d bks^d
evergreen-moist deciduous combinations in the west, aiu j southern
graphical oddity wrought the region a biodiversity hotspot and tne ce .^rAamarn eucalyptus
m aiKi Mk have resulted in the unprecedented destrucuon sub anthropogenic
"Cfflic vaUeys. Since the hill stations of Nilgiris are the j^mn ^ ^
engagements shattered the existing natural vegetation. Now, most ol th
Areas, are highly degraded. ,„„i„l01dWorldgenasofmorethan200species,exhib.
Ceropegia L, (Asclepiadoideae-Ceroj^ae), a trop adaptations (Dyer
'B tremendous diversity with respect to habit, hahut downwardly pointed
1983; Bniyns 1997; Kambale et al. 2012). Basely mllated tubu ^ „ap flowers' (Masinde 2004;
iBirs, torrnsaternporary trap for snmll flies and the group .stt^elyknot.m^ify Pf ^
«^onetal.20(^,.ALri%84),intherev..on«C^.^^^^
^rted to be endemic to India. Later six noveltnrs tave ^ ^ ^
'"dia by about 50 species, of which 38 are dismbuted m *e „„der different lUCN Red-List
Shendage 2010). Many species of Ceropegia are narrow endemics P
categories (Nayar & Sastry 1987-1989).
342
t tuberous, few, to 30 cm long, 1-1.5 cm diam.; stem to 0.5 cm diam., terete, gla-
brous, branched. Leaves simple, decussate, thickly coriaceous; petiole 0.8-1.5 cm long; lamina 5-10 x 2-3.5
cm, lanceolate, apex acuminate, base rounded or acute, dark green, shining above, pale green below, margin
smooth, laxly ciliate. Cymes axillary or extra axillary, mostly one- or two-flowered; peduncle 0.8-1.2 cm long,
ca. 1.5 mm diam., glabrous; pedicel 1.8-2.4 cm, ca. 1 mm diam., glabrous; bract solitary, 2-4 x 0.2-0.5 mm,
linear, acuminate, glabrous. Sepal 3-5 x 0.6-0.9 mm, linear, acuminate, glabrous. Corolla 3-4 cm long; tube
1.4-1.8 cm long, greenish yellow, dilated at base, conical vsdth sudden constriction above, striated with pur-
plish brown lines on the upper half, glabrous, mouth of the inflated part with inwardly directed hairs inside,
inner deeply purple; lobes 1.6-2.2 cm long, strictly and completely folded back along midrib, linear oblong
with rounded apex, lower half creamy white internally with an artistic roundish end, externally with brown
blotches, upper half dark greenish, margin with long brown adpressed hairs, lobes connate at tip forming an
ovoid head. Outer corona cupular, lobes 5, each lobe bifurcate into two divergent horn-like projections, each
projection ca. 1.5 mm long, ciliate, horns of adjacent lobes paired apically; inner corona of five linear, conni-
vent, purple lobes, ca. 2.5 mm long, apically rounded. Pollinia yellow, attached to the brown corpusculi by
short caudicles; pollinarium 0.3 x 0.25 mm in total. Follicle pairs unequal, larger 15-18 x 0.3-0.4 cm and the
smaller 10-13 X 0.3-0.4 cm, slightly curv ' ' ‘
1 long, white, silky.
1, glabrous; seeds c<
:-oblong, black; coma 0.8-15
Local status and Population.— Rare in the locality, a total of 44 matui
Montane Grasslands of Meppadi Forest Range. Most of the growing spots ;
and sharp rocky cuttings.
Habitat. — Grasslands between 1500-1850 m asl.
Geographical distribution.— So far known only from the montane grasslands of South Wayanad Forest
Division (Nilgiri, Western Ghats, Kerala, India).
Etymology — The specific epithet monoharii’ derived from a Malayalai
beautiful, denotes the elegant flowers of the new species.
Flowering and Fruiting.— August-February.
^ of Ceropcgia were collected from Nilgiris. Among the collections an unde-
scnbed, elegant species was included, named here as Ceropegia monoharii Sujanapal SaUm, Anil & Sasidh sp.
novshowsavery„an.owdMnbuaoni„*eMeppadiHUl,angesofWayajE^.C.W^^^
olher species are endemic lo die Pemnsular India - Sri Unba mega-diveisity center Ccropeein jmcea Roi* .
suc^i r T wT “ ■“*"'** ^Nilgiris.
IS foureln, the moist decidnonsforestsandsimiUr areas. Ceropegia decLconnWigh.^^
i.irsm<.Wigli.&Ani..andCdegi.ns Wall, are occurring in eveixreen forests and ^arhabitats above 500®
asl. Most of the speoes of Ceropcgin in dtis region are habitat specific witbanarr^idisiribii, ion ramteTbere-
Sujanapal et al., A new species of Ceropegia from India
343
344 Journal of the Botanical Research Institute of Texas 7(1)
Sujanapal etal., A new species of Ceropegiafrom India
345
Socio-ecology of Cervpegia of NUgiris
The genus Ceropegia is characterized by root tubers either as single large potato-like tuber or fascicled tuberous
roots. Tubers and leaves of many species of Ceropegia are used as food in various parts of the Old World (Mab-
berly 1987). It is also used in indigenous systems of medicines (Pullaiah 2006). Inquiry with the tribes, as well
as other local people of the study area, revealed that tubers of C. hirsuta and leaves of C. deccdsneana are con-
,. Though C. decaisneana doesn’t form a potato-like tuber, the leaves are used as
Amaranth) and are a delicious item during the fasting period such as ‘Eidh’.
a collected from the study area, seven are endemic to Western Ghats-Sri Lanka Biodiversity
HotspJt and three taxa are restricted to southern Western Ghats region. Among the three Western Ghats en-
demic species C. decaisneana is already listed in the Vulnerable category of lUCN; species such as C. ciliata and
C. manoharii are very rare and facing a higher threat than the red-listed species due to various anthropogenic
and environmental factors.
Ceropegia is distinctive with magnificent flowers and a few species have leafless succulent vines; because
of this peculiarity, many species of Ceropegia are widely cultivated as ornamental plants. Observation revealed
tat species such as C. juucett, C. mauohurii. C. elegms. C decaisncam, etc, are suitable for gardening. Stnce
many of these species are host plants of butterflies, they wilf attract butterflies to the gardens. It is found that
the caterpillar of the butterfly. Dunaus genutia, feeds on C. elcgnns, C. Ihwnitesii and the newly described C.
"Mtioborii. These speciesare hitherto undescribed host plants of Rtntntsg™^^^
ACKNOWLEDGMENTS
Anthois are thankfuf to S.R. Yadav, Kolhapur University, Maharastra; Peter Bruyns Bolus 5"“"'
AWca; David J. Goyder, Kew Herbarium; UK, Ulrich Meve. University of Bayreuth Genuany; SnWharthan
Surveswaran, University of Hong Kong and V.C. Balakrishnan, Kannur for relevant iteratures,
ments, expert opinion, and identification of the caterpillar. Thanks also to two anonymous reviewere or
careful review of an earlier draft. We ate thankfuf to Kerafa Forest Department for then “P d"™® Jh'
writ. First author is thankfuf to SERC, DST for fast track fellowship. We e
tribal communities and local people, \
Answh, M.Y. 1984. Asclepiadaceae: genus-Ceropegw. Fascicles of Flora of India, 1
k Calcutta; Botanical Survey of
. (Asclepiadaceae) of Silent Valley. Kerala, India. Rbeedea 7:1 07-1 1
PM, 1 997. A note on Ceropegia L .
Chaherjee, S. 1 995. Global 'hot spots' of biodiversity. Curr. Sci. 68:1 1 78-1
^ RA. 1 983. Ceropegia, Brachyste
'^ambaie, S.S, A.N. Chandore. and S.R.Y/
Western Ghats, India. Kew Bull. 67:843-848. On 1 14-1 1 5
^r.DJ.,987.The plant book. ambridgeUnl«ralvPre^-bnJ^^^^^
^nde, RS. 2004. Trap-flower fly pollination in East African C peg < Botanical Survey of India.
M.P. ™ A.R.K. s«nu. 1 987-1 989. Red Data Book of c^^pegio (Apocynaceae: Asclepia-
^^ttBnoN,J.,s.MAsiNDE, U. Meve, M. Picker, AND A.E. Whittington, z • r „.cni_i5i4.
doideae): biogeographic and phylogenetic perspectives. Ann. Bo • ^ _
'^H,T. 2006. Encyclopedia of world medicinal plants.
S.R. and M.Y. Kamble. 2008. Threatened Ceropegias <
Rawat, J. Chadha, and P. Kumar, eds. Special habitats ai
C?R:^s'M"8L^l...„10.Cempe9k.bb0d,.anewsp^^^
•^ew Bull. 65:107-1 10.
iceae: Ceropegieae) from
ts of India. Dehradun; Wildlife Inst
BOOK REVIEW
Robert B. Shaw. 2012. Guide to Texas Grasses. (ISBN: 978-1603441865, flexbound). University of Texas A&M
Press, John H. Lindsey Building, Lewis Street, 4354 TAMU, College Station, Texas 77843-4354, U.S.A
(Orders: www.tamupress.com, 1-800-826-8911, 1-979-847-8752 fax). $45.00, 1096 pp., 1357 colorpho-
tos, 34 h/w photos, 947 line drawings, 645 maps, 6.6 lbs, 7" x 10".
Somewhat frustratingly here, a number is given for each species to faciUtate direction, but page numbers would
likely be more helpful. Each species contains a brief circumscription and general habitat, an illustration of the
plant and florets, and a map of known Texas counties in which it occurs — though it would be helpful for the
circumscriptions to have mentioned complete distributions, beyond just Texas. Live images, growth form, and
inflorescence are often included as well. My only complaint is that the species circumscription does not include
any information about typical fruiting and flowering times. The circumscription could also be more detaUed
regarding morphology, making it a tad more difficult to obtain a positive identification from the key, but illus-
trations drawn to scale often make up for this. The illustrations are wonderful, though, and often take up half
of the page, so they are very easy to see.
The appendix provides a section on collection, preparation, handling, and storage for grass specimens
that should be very useful for new collectors. This section also discusses type designations and the importance
of vouchers and provides a how-to for collecting, pressing, and labeling herbarium specimens. Also provided
in this section is a list of herbaria in Texas, their addresses, specialties, specimen numbers, and herbarium
codes. While this is helpful, some of the information is incorrect or already out of date, and it would be wise to
verify this information with Index Herbariorum (www.nybg.org).
All in all this is a great resource for anyone, amateur or professional, looking to learn more about Texas
grasses. Guide to Texas Grasses is not a field guide by any means; it is bulky and heavy, even as a paperback, and
in a car or on a desk. It has unusually thick paper for a text of this sort, which makes the impeccable photo-
graphs all that more useful, but will also help maintain the life of the text, as pages will not tear easily. I would
recommend Guide to Texas Grasses to anyone wanting to delve into the magical world of grasses. — Rebecca K
Swadek, Field Biologist, New York City Natural Areas Conservancy, and Research Associate, Botanical Research In-
stitute of Texas, Fort Worth, Texas, USA.
arkworth, M.E K.M. Capels, S. Long, and M.B. Piep, eds. 2003. Flora of North America north of Mexico, VoL
25 Magnohophyta: Commelinidae (in part): Poaceae, Part 2. Oxford University Press, New York, U.S.A
iggs, G.M., Jr., BU. Lipscomb, and RJ. O’Kennon. 1999. Shinners & Mahler’s illustrated flora of North Cen-
T Miscellany series. Botanical Research Institute of Texas Press, Fort Worth,
Texas,U.S.A. 16:1-1626.
A NOTE ON ^TRITRICHUM SCHRAD. 1819” AND
ERITRICHIUM SCHRAD. EX GAUDIN 1828 (BORAGINACEAE)
Kanchi N. Gandhi
Harvard University Herbaria
22 Divinity Avenue
Cambridge, Massachusetts 02138-2094, U.S.A.
gandhi@oeb.harvard.edu
David F. Murray
University of Alaska Museum of the North
907 Yukon Drive
Fairbanks, Alaska 99775-6960, U.S.A.
t name for the genus.
RESUMEN
During a routine analysis of a draft
mendatural problem with a genus r
ncountered 2
It for the Flora of North America, Volun
i its type species.
wmie assembling material for an examination of the North American Eritrichium, we found that ^th
“Eritrichum Schrad. (1819: 186)” and Eritrichium Schrad. ex Gaudm (1828: 4, 57) have been use . ra ers
genusname,ifitweretobevalidlypublished,wouldhavepriorityoverGaudin’sEritrichium^
A sampling of monographs, revisions, and floras shows an inconsistency m spelling
(1901 407), Lechner-Pock (1953-1956: 100), and Weber (1990: 113) considered
mlidand used -Erirrichum- DeCandolle (1846:122), Popov (1953: 482),
757),Hulten (1968: 773), Scoggan (1979: 1284), Ovchinnikova (2009: 142) and Zhu “■( ; ^ *
™i, see below) accepied Schrader as theauthor of the gennsbutspeUed .he gen^nameErt^
apnlerencedrey can exercise in casesof orthographic variations. Abranw(1951: 537) andjoto^^^^^^
while using the spelling Erilricbtnm, gave the citation as Schrader in Gaudnr Etna y, a er
quist (1984: 222), and WeUh & al. (2003: 61) used Erilrichinm Schrader ex Gan in -Eritrichi-
The International Plan. Name index (IPNl 2012a) has the gentn name t^dtKbM^P^^
“I Schrad. in Comm. Gotting. iv. (1820) 186, mote. ^ I, j- 4 57. 1828." Additionally, the
mm (Plantarum) (ING 2012) listed “Eritrichium Schrader ex Gaudm, F .
Names in Current Use (NCU-3e; 2012) has essentially the same
Schrader (1819: 186) distinguished many taxa found m the gi freqnenlissima, cum discus
PhoK (. fruiting receptacule): Type ‘I. ImmtiMM s ptan Is. ^ ^ p.,
huou maturescente fomtam non mutat, ut m P ' ^imis Anchusae speciebus q) e. varus aliis;
ntatuntatem fructus magis vel minus elevatur, ut in Myoso , P on„s. Eritrichum (Myoso-
3. «cal. 11. = lice. - that is] hemisphoerico - conica, cuius sumths cum p _
-hna™),P„,„u„q,m,m^mcaetaff.uess^cm^
The question is whether the Type 3 character ^ (Pulmonarin vitginica and allied species),
or to two genera: Entrichum (Myosotis nann) and an unnameu g published by
tt|he key character does indeed apply to 3„a a„ unnamed genus, then the
Schrader. On the other hand, if the key character applies to both tn
genus name is not valid. both Eritrichum and an unnamed genus, and the
We have concluded that the key character does apply ns^ftWalidation of the name Eritrichium is
name was not validly published by Schrader. Therefore, Gaudms t
Journal of the Botanical Research Institute of Texas 7(1)
ear in any of Linnaeus’s later works and it
o Piante Rare Sicil. 1. 107. 1697) nottl
Eritrichium is typified by E. namm, which was based on Myosotis nana L. The validity of M. mm (pub-
lished in Amann, Fl. Alp. 13. 1756) has been questioned. The protologue is: Myosotis nana Bocc. T. 107 (1697).
The Linnean Typification Project (2012) does not list this name on its website. IPNI (2012b) t
nana L. as invalid and provided a remark: “The name doesn’t £
thought to be an abbreviated reference to Boccone’s phrase nan
publication of a new binomial (C. Jarvis, pers. comm.).”
After a discussion with IPNI’s editorial staff at Kew, the name Myosotis nana is deemed valid and the abot
remark is revised to: “The name is thought by some to be an abbreviated reference to Boccone’s phrase nan:
(Museo Piante Rare Sicil. t. 107. 1697) rather than publication of a new binomial (C. Jarvis, pers. comm.
(IPNI, 2012c).
Thus the correct citation of the generic name is as follows:
Eritrichium Schrad. ex Gaudin, Fl. Helv. 2:4, 57. June 1828. Type Species: E. nana (L.) Gaudin (Myosotis nana L. [correctauthc
ACKNOWLEDGMENTS
We thank Werner Greuter (B), Dan H. Nicolson (US), and James L. Reveal (CU) for a discussion, and Anthony
Brach (MO C/O: A, GH) for helpful suggestions. We also appreciate the review comments of Matt Guilliams
REFERENCES
Abiwms L. 1951 ‘Entrichiumr p. 557. Illustrated flora of the Pacific states: Washington, Oregon, and California. Vol. 3.
Stanford Univ. Press, Stanford.
DA Webb, eds. Ho, a europaea. Vol. 3. Cambridge Universlw Press, Cambridge. P. 1 1 7
Cronc!Uist,A. 1959. •fr/fr/ch/um.' In: Hitchcock CL A Cmnn.iict M . ■ * „fthp
'‘■‘^''''“‘"'^•'-'^O'^duistM.Owenby, and J.W. Thompson. Vascular plants of the
Pacific Northwest. Vol. 4. University of Washington Press, Seattle. Pp. 202-204
Tul*' T’ '““f T J.L Reveal, and P,K. Holmgren. Intar-
Pp S M3 “ ” ' ""''■"'ountaln West, U.SA Vol. 4. The New York Botanical Garden New Yort
Dt^o^A,846.-5,.«ch,„m.-ProdromosSys,ema,isNa,urallsRegnlVege,abllls.Vol.lO.^^^^^^^
Co, Ztirich.' > in homlnis anlmallumque uses vulgo cultamm continuata. Vol. 2. Orelli, Fuessll S
Hr»ra,E. 1968. Flora of Alaska and neighboring territories. Stanford Unlv Press Stanford
Tess^rA^rgoT ^ ” •’'<P://botany.sl.edu/ing/ingformcfm [ac-
|:=pNi“;r2o;2;rb;;::::r::;r::^
LiOwmXr. L ^955^956°^^!^.^™^““!“^^^
Lnnean Typification PROJEa. 2012. Published on the Internet at httoy/vZ!^ Verwandten. Phyton 6: 98-206.
ects/linnaean-typification/database/ [accessed on 22 April 2012] "^"^•^^“'^'•^earch-curation/researc
Names in current use for extant plant genera. Electronic version 1 .0. Published
ncu/genera/NCUGQuery.htm [accessed 22 April 201 2J
J. 1 965. Flora of Japan. Smithsonian Inst, Washington, D.C
Ovc„»wwAS.V.2009.0n,heposlt,onof,het,ibeEr,tncf.JedeintheBoraglnaceae
w, M.G. )953.“Eritrichiumr In: Shishkin, B.K. ed. Flora USSR. Vol. 1 9. Acad. Nauk, Moscow and Leningrad. Pp. 482-521 .
Schrader, H. A. 1819. De Asperifoliis Linnei Commentatio Recitata. Ii
ix..Commentat. Soc. Regiae Sci. Gott. Recent. 4(1 ):1 75-1%.
ScoGGAN, HJ. 1979. "fr/fr/ch/um."The flora of Canada. National Mus. Nat. Sci., Ottawa, Publ. Bot. N
Weber, W.A. 1990. Colorado flora: Eastern Slope. Univ. Press of Colorado, Niwot.
Welsh, S.L., N.D. Atwood, S. Goodrich, and LC. Higgins (eds.). 2003. A Utah flora. 3"* edition, revised. Brigham Young Univ.,
Wight, W.F. 1902. The genus Eritrichum in North America. Bull.Torrey Bot. Club 29: 407-414.
Zhu, G.-L, H. Riedu and R. Kamelin. 1 995. "Erifric/j/um." In: Z.Y. Wu & P.H. Raven, eds. Flora of China. Vol. 1 6. Science Press,
Beijing and Missouri Botanical Garden, St. Louis. Pp. 378-390.
Journal of the Botanical Research Institute of Texas 7(1)
BOOK REVIEW
Charles R. Hatch. 2007. Trees of the California Landscape; A Photographic Manual of Native and Orna-
mental Trees. (ISBN: 9780520251243, hbk). University of California Press, 2120 Berkeley Way, Berkeley,
California 94704-1012, U.S.A. (Orders: www.ucpress.edu, orders@cpfsinc.com, 1-800-777-4726, 1-800-
999-1958 fax). $63.00, 542 pp., >1000 color illus. (photos, line drawings, maps), 8 1/2" x 11".
The California Landscape
Topography
Geography
Native vegetation types and habitat
The interface: trai
ban landscapes
Trees in Urban Landscape Design
ween native and ur-
Creative applications and special effects
Special use and applications lists
Trees of the California Landscape (Compendium)
Taxonomy
Tree Structure
General Characteristics
Flower Types
Fruit Types
Glossary
Distribution of Trees by Classification
Trees by Family
Key to Tree Genera
lypes oj uroan landscapes Key to Tree Species
Arrangement of trees Index to Trees
Trees in various contexts Index to Common Names
Codes and ordinances affecting tree planting Arboretums
Design elements in tree selection Bibliography
Planting and irrigation
Trees o/lke California Undscope is an8V4-xll-hardcovcrbook of 542pages. It conlains numerous color pbo-
lographs, color drawings, and maps encompassing the entire state of California and its many vegetatioiBl
zones from the coast across the Central Valley, to the Cascades, Sierras, and the Mojave and Sonoran deserts,
to^t er, t ts text contatns tnorphologtcai descriptions, habitat information, photographs (leaves, bark.
and habtt), and drawings for 107 native andSll ornamental species.
Cli describing the relationships between vegetation, geography, topology, and
cltmate-tmthout excluding mans effects on vegetation and landscape. tL introLL gL a very brief
overvtew of the textand some of the major vegetatrotml zones. The nexVsection discusses thLffectsoft^
ogy, geography, and climate, with excellent maps. Also useful ate three (north central and south) cross sec-
tiotmMiagrarnsofCaliforniatopology,„hic^
tionoTr^ah
HiTfrh and give lists of typical vegetation from that particular ecosystem. HeK,
bto egtonal landscapes, whmh tncorpomte the features of the native environment inL parks and housing
and establtshes a de^r connectton to the natural processes" of rmtum. Examples include grading and
NOTES ON ERITRICHIUM (BORAGINACEAE) IN NORTH AMERICA
University of Alaska Museum of the North
907 Yukon Drive
Fairbanks Alaska 99775-6960, U.S.A.
and selection of a
nail cushion-forming plant of the Bering Strait region, which is how Wight
(1902) ami then Hnlten saw it (1968). Later, however, Hulten (1973) canne to believe that he had been led asttay
by Wight who applied E. chamissonis to the plants with flowers included within or barely exerted fmm the
cushion of stems and leaves. Hulterfs change in thinking was brought about by two errors. Fust, he decide
lhata cushion form should be named E. onrlioides (Cham.) DC, because PopoVs (1953) tllustratton of aretirn-
des- showed a densely pulvinate plant (Plate 25, fig. 1), and second, Hulten believed the type specin^o .
ctonissonis had long flowering stems, which is at odds with the original descnption of de Candolle (IWL
As to the first assumption, Jurizev and Petrovsky (1980) determined that Po^v had unknowmglytllus-
t«tedadistinet. densely pulvtnatespecies,not£ritrlchi»umtioides,butone that hadgonennrecognmeduntfl
tepubllcationofE.,schuhtschor„mJurtzsevSrW.Petrovsky,whichonleafmorphologyalotw«re^^^^
rable from both E. aretioides or E. chamissonis.
As to the second assumption, the descriptions by de CamioUe ofEritrichiun. uretiofc
■te brief, and the two species are not well differentiated; nevertheless, he made clear that E. ^ “
plant with the shorter stems of the two. In fact, the flowers of E chumissonn at anttois no « er
^rfaceofthetuft,whereasEuretioldeshasfloweimgsteins,mlUto«^
densely compact, but differ from E. chamissonis in the length ot the flowering
flowers. Hulten had no reason to doubt Wight’s account. n- awi i., ncnni chased on sneci-
De Candolle (Prodromus 10:125. 1846) the voyage of the Rurife 1816 or 1817. De
mens originating at LE, collected by Chamisso or Eschscholtz during <«nrti-Laurentii ad fre-
Candolle referred unequivocally and unambiguously to St. ^ localities and thus
t«mBeering(Cham.O”but then continued “inKamtchatkaa^
two specimens, or syntypes, were the basis for his new spe
had reported earlier (Chamisso & Schlechtendal, Linnaea 4.442. 18 ) y ^ ^ Chamisso in his
ously not all of the plants so named by Chamisso were regar e y reference to
account was clear that the locality for his M. viUosa was St. Lawre y-
Kamtchatka. with excellent digital images provided by G, then
AreviewofthetypesheetatG-DCwasundertaken,firs The^one from Kamchatka, collector
a visit to G. Mounted on the type sheet at G-DC ^ ^ j ^^^^ription). This speci-
unknown, provided to de Candolle by a colleague at li ( h. ac ^
®cn IS labeled Myosotis villosa Ledeb., and with its strong y exertis,” instead it clearly resem-
flescription of Eritrichium chamissonis: “. racemuhs pauciiio
'•««-lte.hBtT«as7(1):3
352
bles E. villosum (Ledeb.) Bunge. This specimen cannot be considered as E. chamissonis. Why de Candolle would
make a direct reference to the Kamchatka specimen is a curious contradiction of his own description.
The other specimen on the type sheet, also labeled Myosotis villosa, is a specimen that matches the de-
scription of Eritrichium chamissonis as the flowers barely exceed the cluster of sterile, leafy shoots, and it is
clearly attributed to Chamisso. However, this one is labeled “St. Lorenz insel” meaning St. Lawrence Island,.
This specimen, while a perfect match with the description of de Candolle, is in conflict as to location. However,
the Rurik visited both St. Lawrence Bay, Chukotka, and St. Lawrence Island, Alaska, during its voyages.
De Candolle took the locality Lawrence Bay directly from what Chamisso (in Chamisso and Schlechten-
dal 1829) had written under Myosotis villosa. Yet, neither of the two specimens on the type sheet is from St
Lawrence Bay. That de Candolle failed to add St. Lawrence Island to his description, despite the label on the
sheet before him, must have been a lapsus calami. It is otherwise impossible to reconcile the discordance.
An herbarium sheet at LE bearing Chamissio’s signature breve, labeled Myosotis villosa, and from St. Law-
rence Bay, would seem to be what is referred to in the description of Eritrichium chamissonis, but it is clearly not
consistent with it, inasmuch as the flowering stems are tall, leafy, rising well above the basal leaves, thus closer
in habit to E. villosum or E. aretioides than to E. chamissonis. That sheet cannot be considered as the basis forE.
chamissonis.
Another LE sheet has been labeled, incorrectly, as holotype by Ovchinnikova (2008). The specimens are
attributed to Eschscholtz. Ovchinnilova (2008) reported the locality as “Sin. St. Laur[enti]”; the handwriting
on the original labels attached to that sheet is not easy to decipher. Who wrote that note is not known (VY.
Petrovsky, pers. comm.). There is no evidence at LE or G-DC that this material was seen by De Candolle.
Not found at LE is a sheet with specimens of Eritrichium chamissonis from St. Lawrence Island from which
the^specimen on the type sheet at G-DC might have been taken. The source of that specimen at G-DC remains
Nevenheless, de Candolle did have before him and most certainly he based his description of Eritrichium
chomis^is on the Alaskan specimen. De Candolle did not designate a type, therefore, there can be no holo-
type. The specimen, a syntype, on the type sheet at G-DC from St. Lawrence Island, Alaska, is designated here
as lectotype for E. chamissonis DC.
Eritrichium chamissonis DC., Prod. 10:125. 184
olJes mixed with M. vIBostt, a. St, Lawrence Bay, Chukotka, andXhom St
two ItKalities were given by Chamisso, the nameM. ate, imdesmustbeb^ on one or more of the synw«
and a lectotypification is necessary.
Ovchinnikova (2008) cited an Eschscholtz specimen from St. Lawrence Island Clns. St. Lanrcntit
, t ' “ “ “miouks. Although her incorrect usage of the term ‘hololype'
notlec^^rrelT-trS^r^t^^^
.he name needs a fotn«i
unre a e to aretioides. Another is from the Academy in St Petersburg (LE) with a printed label
Myrnmuarntt^des, Sinus St. UurentiLbutacolfectot not tecord^l'Thf^^^^^
hB would be better as the type for M. orehoides. Wheteas a specimen at LE seen by Chamisso would ha«
353
greater standing, there is no Chamisso specimen at LE from which the one at G-DC might have been taken or
one that better represents Chamisso’s intentions. Since the specimen at G-DC was determined by Chamisso
and furthermore connects Chamisso to de Candolle, that specimen is designated here as lectotype.
ACKNOWLEDGMENTS
Thanks to my colleagues at the Komarov Botanical Institute (LE), St. Petersburg, V. Yu. Razzhivin, Andrei Sy-
tin, and especially to YN. Petrovsky for information on the Chamisso and Eschscholtz specimens there; to
Laurent Gautier, Curator, and Nicolas Fumeaux, Collections Manager at Geneva (G), for making it possible for
me to accomplish so much during my short visit to view types and associated specimens. 1 am grateful to re-
viewers who very kindly corrected my errors and generally improved the text of this note. James Reveal has
carefully read and commented on the manuscript. Reveal and Kanchi Gandhi clarified the authorship of the
Eritnchium names: A.P. de Candolle or DC, the father, the author of the Eritrichium names and descriptions,
notwithstanding the page heading that credits A. de Candolle or A. DC, the son.
. 1829. Dep
Chamisso, A.L von and D.F.L. von Schleci
rationem dicunt. Linnaea 4:435-496.
DeCanooue, A.P. 1846. Prodromus systematis naturalis regni vegetabilis sive enumerato coi
specierumque plantarum hucusque cognitarum, juxta methodi naturalis normas digest.
HultEn, E. 1968. Flora of Alaska and neighboring
HuLTfN, E. 1973. Supplement to flora of Alaska an
McNhu et al. 2012. International code of botanical nomenclature for
opted by the 18th International Botanical Congress ^
Ovchinnikova S.V. 2008. Conspectus of the genus Eritrichi
Russia 1:17-36. [in Russian]
Jurtzev,B.A. AND V.V. Petrovsky.
It world of Asiatic
iginaceae) species in North A
Eritrichium. ln:T0lma.he», A.I. .nd BA Jurtze., eds. Arctic Flora USSR. 8^38-245. tin
Popov, M.G. 1953. Eritichium. In: Shishkin, B.K., ed.. Flora
Wight, W.F. 1902. The genus Eritrichum in North Americi
USSR 1 9:482-521 . Akad. Nauk. Moscow & Leningad. [in Russian]
a. Bull. Torrey Bot. Club 29:407-414.
354
Journal of the Botanical Research Institute of Texas 7(1)
BOOK REVIEW
Charles R. Hatch. 2007. Trees of the California Landscape: A Photographic Manual of Native and Orna-
mental Trees. (ISBN: 9780520251243, hbk). University of California Press, 2120 Berkeley Way, Berkeley,
California 94704-1012, U.S. A. (Orders: www.ucpress.edu, orders@cpfsinc.com, 1-800-777-4726, 1-800-
999-1958 fax). $63.00, 542 pp., >1000 color illus. (photos, line drawings, maps), 8 Vz" x 11".
drought tolerant landscaping to preserve native oak forests in new developments and the use of floodplains for
open grassy park corridors instead of buildings.
The discussion on the use of trees in urban landscapes is very useful for landscapers and homeowners.
This section discusses different types of landscaping, purposes, tree arrangement, shape, spacing, and plant-
ing requirements of different trees. Lists are provided for different colored flowering trees, white barked trees,
fall colors, long-lived trees, desert accents, tropical accents, and more.
There are excellent sections at the end of the book showing leaf shape, cone and fruit shapes, and other
morphological characteristics of different species. Following these is a glossary of terms and a dichotomous
key to genus.
This book provides an excellent overview of CaUfomia ecology that would be useful to many different
disciplines, including landscape designers, teachers, naturalists, homeowners, etc. In fact, the only complaints
1 have about this text are map errors. In one error, Mariposa County is incorrectly labeled as Fresno County, so
Fresno County appears twice in maps containing county labels. Another error is for the species page of Joshua
Tree where the distribution map for California Bay on the previous page was duplicated, giving an incorrect
distribution for Joshua Tree. Something to consider for a revised or second edition!— Rebecca K. Swadek, Field
Biologist, New York Natural Areas Conservancy, and Research Associate, Botanical Research Institute of Texas, Fort
Worth, Texas, U.SA.
LT«b$7(1):354.2
A REVIEW OF THE NEOTYPIFICATION OF ASTROPHYTUM CAPRICORNE
VAR. CRASSISPINUM (H. MOLLER) OKUMURA (CACTACEAE)
Richard R. Montanucci Heinz Hoock
Dept, of Biological Sciences, Clemson University Weingartenweg 35
Clemson, South Carolina 29634-03 14, USA D-84036 Landshut, GERMANY
rrmnt@clemson.edu heinzhoock@kabelmail.de
ts wiOi Heinrich MoUer’s (1925) original description of this Uxon. Momberger’s assertion
tent. We designate ZSS-019963 as the neo-
,n Konflikt mit Heinrich Moller’s (1925) Original-Beschreibungdie:
n der Sierra Agua Chiquita, C
Nachden Bestimmungen des ICBN, Ariikel 9.17b, empfehlen wir, ULM-18572 aufzi
bezeichnen ZSS-019963 alsl
INTRODUCTION
to 1925, Heinrich Mailer described Ediinocartus capriconm var. crassispiaus Isle] bas^ on a sei^ of sp^i-
■«®colkaedbyhUbrother,ArthurMoller,whoresidedinSanPedn,deteCobn.as,C,«hu^Me^The
"rtglnal series coUectrfbyArlhnr Mailer was shipped to Heinrich Molkr for pnrpcs^ of study
«on.Ah„n.halfofd,eplan.ind.eoriginalser.sh^sparse,^^.a^^^^
the remaining plants were completely nude and green. The spines
™d numbered from 6 to 8 per areole. The flower was described as canary ye ow wit a su urou
IW The number of speci^et^ in the original series was not mentioneda.^»photo^pfe^^^^^^
0^ Original description. Furthermore, Heinrich MOller did not designate a hototype nor
specimens from the original series were deposited in a herbarium. The a nee o i ’
■kolaUum to find this plant at the type locality (see belowlset the sugeforadecades-long debate (M Mom
Her 2007) concerning theexistenceand identity of thistaxon^^^^^^^^^^__^^^^^^^^^^^^^^
Jbe type locaUty ofMOllefs var. Enough the efforts of many collector,
^planrimatchmgtheoriginaldescript.onmttau™n^nmg^^^^^^g^^^^^^^^^^__^^^^
occurrence of Astrophytum capricome var. seni e m P
documented (Hoock 2008, and references therein). It also occ the Sierra Mayran. Farther
*^rra de la Pena) and Cerro Bola, but not yet recorded trom tne sma
356
east, in the region near Parras de la Fuente, Riha and Busek (1986) reported the occur
capricome var. capricome (A. Dietr.) Britton et Rose.
In 1930, the eminent cactus collector, H.W. Viereck, discovered an Astrophytum in the vicinity of Cuatro
Cienegas, Coahuila, Mexico (Bernhard & Hoock 1986). The collected specimens belonged to the capricorn
group, but differed from the type species in having stout, flattened spines (usually 6 to 8 per areole), a large
stem size (a maximum of 50 cm in height), and an epidermis densely covered vsrith short, white hairs (tri-
chomes) having a felt-like texture. Viereck’s discovery was obviously morphologically similar to the previously
described var. crassispinum. Kayser (1933) described and named the population from the vicinity of Cuatro
Cienegas as Echinocactus capricomis var. niveus.
In 1933, plants identified as Astrophytum capricornus var. crassispinus [sic] were featured on postcards
created by Otto Stoye. The postcard photographs included both nude and sparsely flocked specimens from the
private collection of Kurt Kreuzinger (Hoock 2008:162). Our examination of the postcards led us to conclude
that the plants were very similar to those described by Heinrich Moller in 1925. The plants may have been
imported from Coahuila by A.V. Fric, who was a known associate of K. Kreuzinger, but the precise locality in-
formation was never published.
In 1985, a locality for “crossispinum-like” plants in “northern Coahuila” was discussed in an Astrophytum
“ring letter” which included a photograph of the habitat. Ulrich Bernhard and Heinz Hoock searched for, and
found the locality by matching the mountains in the photo with the actual mountain range on the landscape.
The locality was situated approximately 42 km west of Cuatro Cienegas; the habitat and its plants were de-
scribed by Bernhard and Hoock (1986). The authors reported that densely flocked, white plants (var. niveum)
and totally nude, green plants (which the authors called the “nude form” of Astrophytum capricome var. niveum)
occur together in the habitat in approximately equal numbers. But intermediate (sparsely flocked) plants were
encountered only very sporadically (Fig. 1). Similar observations concerning the relative abundance of flocked,
nude and sparsely flocked plants were also reported by Baumann (1992). Although no quantitative data were
presented, the sporadic occurrence of sparsely flocked plants was surprising considering the likelihood of
cross-pollination between the white and green forms.foha and Busek (1986) described variation in the spine
morphology and epidermal flocking of these Astrophytum from the locality west of Cuatro Cienegas. The au-
thors noted, aside from the typical var. niveum, several forms including nude plants with white, yellow, and
dark brown spines and yellow flowers.
Various authors, including Megata (1944), Haage and Sadovsky (1957 a,b), Sadovsky and Schutz (1979),
Ito (1981), and Busek (1981) regarded the nude and sparsely flocked plants with stout spines as ^'crassispinum^
It could not be determined whether the plants illustrated by these authors originated from a single locality «
from several different localities, or if any came from 42 km west of Cuatro Cienegas. foha and Busek (1«.
Bernhard and Hoock (1986), Baumann (1992), Hoock and Klesaewski (2006). and Hoock (2008) all indicatta
that the nude plants from west of Cuatro Cienegas were morphologically similar to Moller's crassispinunt
In 2007, Momberger reported the surprising discovery of the -missing” Astrophytum cupricomr vat.
crassispmum,cIaimmgthatthistaxonoccut5alongthcwestemslopesoftheSierraAguaChiquita.Thisinoun-
tain range is a northern extension of the Sierra de la Purisima which forms the eastern margin of the Bolson*
CtutroCimegas. The la, ter Unearly divided into eastern and western basim(bolsons) by the SiertaSanM^^
cos. Momberger accorded this Astrophytum population fonnal taxonomic recognition as Astrophytum cuF'
come snbsp ^ilc vat. crassispinum and designated a neolype (ULM-18572) which was deposited in UUI
(Herbanum Umversitat Ulm, Germany).
Pavlttek (2011) subsequently described the Sierra Agua Chiquita population of Astrophytum as a nr*
subspecies, Astrophytum capricome (A. men.) Brinone, Rose subsp sunyrntncnse. He designat«^
rib With spines as the holotype: PR 745389, Department of Bouny of the National Museum. Praha (PtagW*'
Czechoslovakia.
^acon^quence of MombergePsdiscovery.there are now rivo Astrophytum populations that poten.*
qualify as Moller’s var. crassispinum.
E morphological data p
358 Journal of the Botanical Research Institute of Texas 7(1)
bolson populations of Astrophytum and compare them with Heinrich Moller’s original description. Our pur-
pose is to determine if Momberger’s recognition of the Sierra Agua Chiquita population as var. crassispinwn
and his designation of a neotype for this taxon should be accepted or rejected under the provisions of ON
Article 9.17.
ORIGINAL DESCRIPTION
The original description of crassispinum by Heinrich Moller (1925) is quite explicit, presenting morphological
information on the vegetative and generative traits of this taxon. In our opinion, the potential of attaching this
name to a natural population has always existed owing to the completeness and detail of Moller’s description.
The original description reads as follows:
Our English translation of the original description reads as follows:
Body simple, spherical to conical in shape, 15 cm high, 8-9 cm in diameter, foliage-green. The body of half of
the obtained plants is completely naked; the body of the others is covered with sparse, irregularly scattered
wooly flecks. The apex is slightly depressed, covered with a little yellow felt. Ribs are always 8, sharp-edged.
Areoles are 20 mm apart, oblong, to 6 mm in size, with abundant yellowish hairs, which later turn gray and
dingy. Spines 6. maximally 8, flattened, very thick, black-brown in youth, later gray-white, chalky. The bottom
pair is strongest, bowed to the left and right and standing away from the plant, with the tips curved upwards.
The plant is surrounded by the loose tangle of spines which do not break off or prick each other. Flowers are
near the apex, about 6 cm tall. The ovary slender, covered with pointed scales. Hower tube with abundant
wool. Perianth funnel-shaped. Outer tepals acuminate, dirty yellow; inner tepals spatulate, denticulate, canary
yellow and satiny. The throat of the flower is not crimson, but sulfur-yellow, waxy. The stamens barely half as
long as the corolla tube. Filaments yellow, anthers slightly darker. The pistil has 8 bright yellow stigma lobes
and towers above the stamens. Fruit and seeds as in the type and other varieties.
MORPHOLOGICAL COMPARISONS
Morphological comparisons of the Astrophytim population from the western
bolson (42 km west of Cuatro
■ ‘ ■ *.luui^uu, with Moller’s (1925) original de-
below. Comparative data are drawn from the published literature, but the published
informationissupplemented by our direct observations of theseplants from the habitat and from horticultural
matenal. In comparing Moller’s original description with the Astrophytum from the Sierra Agua Chiquita, the
tollowmp Domts are TnaHp-
1) The plants 5, udKd by Moller consisted of complexly nude as well as sparsely flocked plants innearly«H
proportrons. ^e population front the Sierra Agua Ch^uita iscomprised of plants having sparse, irreg«-
wooUy spots but completely nude plants have not been found (Hoock Sr KlesrewsK
2006; Momberger 2007; Pavlfek 2011). Furthermore, as far as we can determine, none of the seve»l
rT-"t''w‘r® H'' ® Gottschlich, F. Jahn! K.P. Kleszewski, P.
Pavlicek, W. Sporbert) has ever reported finding completely nude plants
2) Moller (1925) described the new spines of rrnssispirtnm as having a black-brown cobr, turning gray-»l>*
with age. But, Momberger (2007) described the new spines as straw-colored in the plants from the Sierra
Agua Chiquita. Hoock and Kleszewski (2006) also described the plants from this mountain range as
having new spines with a golden-yellow hue, becoming dirty gray with age. PavliCek (2011) described
the new spines as gray-yellow, the older spines being predominantly gray in color and slightly frayed. In
culture, we have noted some plants with reddish brown spines. However, such plants are the exception;
the most common color is golden yellow or straw-colored. If Moller had observed any plants in his origi-
nal series with yellow spines at the stem apex he would have certainly mentioned it. Significantly, there
are no reports of plants from the Sierra Agua Chiquita with new spines of a black-brown color.
3) In Moller’s original description of crassispinum, the spines are described as “flattened” and “very thick, 6 to
8 per areole. According to Momberger (2007) the plants from the Sierra Agua Chiquita have 6 to 8 (-10)
spines. Pavlicek (2011) gave a slightly higher number, stating there are always more than 8, and usually
10 to 12 spines. According to Moller (1925) the bottom pair of spines is strongest, bowed to the left and
right and standing away from the plant, with the tips curved upwards. In plants from the Sierra Agua
Chiquita, this trait is only weakly expressed. Momberger (2007) stated that the bottommost pair is
mostly flattened and the upper spines mostly round. PavliSek (2011) described them as flexible, slightly
angular, and in some specimens almost triangular. He stated that they are only slightly stronger than in
the subspeciessenile. Direct comparisonsrevealthatthespinesoftheSierraAguaChiquitaplantsare not
as stout as those of the plants from the western BolsondeCuatroCienegas, but compare favorably with
the spines of var. aureum from the Sierra de la Paila (Fig. 2). . , , .
4) The blossoms otlheAstropMumfromSier™ Agua Ch^uiuarcyellowm.harcd.hn«,,W. the redcolor
usually reduced, and in exceptional specimens the throat is pure yellow, browntsh or honey-yellow
(Hoock & Kleszewski 2006; Momberger 2007). PavWek (2011) described .be Hower
brown, brown-yellow or pure yellow. Only the yellow-throated specimens agree with Moller s original
description of the flowers.
Momberger’s assertion that the plants from the Sierra Agua Chiquita are the
been influenced byphotospublUhed in Haage and Sadovsky (1957b) and Sadv^kya^Schu.z(1979^e^^^^
knowledge.ha.thepl.ossLwcloseag.eementwi.hd.eAstr„phy,„mhom*eS.^
foracomparison).However,acri.icallyimportantpointmust^^^^^^
onginal material collected by Arthur Moller. They represent p n - r, c^i,,-,rr 1Q7Q152I There-
Czechoslovakia, and credited to Konrad KayserandSadovskyhimselffSadovsky^^^^^
fore, Momberger’sstatement that thephotosdepimhe“re^^^^^n^^^^
g«y-green epidermis. Our taxonomic concept follows that sugp .,„3ChiQUita population
ever,wearepreparedtodefertoPavlftek%(2011)taxonom.cwnc,mmn^
tepre^ntsanewsubspeciesalthougho^catsymlmrf
A comparison of the plants horn the western ( information as well as direct
”aldescriptionispresentedbelow.Aspteviouslystated,wedrawupo p
obse,vationsolplantsinhabi.a.andculn.re.Thefollowingpoinma.emade;
I)Tlre original material collected by Arthur Moller inc^d»mpl«ely^wd^^^^^_^^^^^
populations west of Cua.ro CienegasrnctocompWy^d^.^^^^^^
plants. While the presence of nude and s^ y ^ problematic, and re-
prising the original series, the presence of densely flocked t
quires further discussion (see “7, ibkcklrnvn in color, becoming gray-white with
SNewspinesemergingfromtheapexofthestemareusualiyp
age, and hence agree with Heinrich Mollers ongina “flattened” and “very thick”, numbering 6
) The original description of crassispinum states that t ^ ^nd standing away from
to 8 per areole. The bottom pair of spines is stronge ,
362
Journal of the Botanical Research Institute of Texas 7(1)
the plant, with the tips curved upwards. The spines of the plants from the western holson are very stout
somewhat flattened, and usually number from 6 to 8. Typically, the bottom pair is strongest and charac-
teristically bowed to the left and right with the tips curved upwards. As mentioned previously, the spines
are considerably stouter than those of plants from the Sierra Agua Chiquita population, and better fit the
concept of the name “crassispimm.” Apparently, Heinrich Moller was quite impressed by the thickness
of the spines which he described. Also, he had in his possession at the same time var. aureim for the
purpose of publishing a taxonomic description in the same paper (Moller 1925). His published descrip-
tions are detailed, indicating that he had excellent knowledge of the comparative morphology of the two
taxa. Momberger (2007) suggested that the name “crassispimm” is a misnomer which resulted in a mis-
conception of the spine morphology of this taxon. However, Momberger’s opinion is not supported by
evidence. We note that the concept of the name is compatible with the detailed description of the spine
morphology given by Moller (1925).
4) Moller’s original description states that the throat color of the flower is sulfurous yellow. In plants from the
western bolson, the throat varies from peach to orange or orange-red, and in relatively few specimens the
throat color is pure yellow. Riha and Busek (1986) reported that among a series of 12 plants, four plants
(33.3%) had exclusively yeUow throats; the remaining eight had red color in the center. These plants
originated from seed material procured by A.B. Lau in 1977 from the western bolson. Heinrich MoUer
did not state how many blossoms he observed in the original plant material, and therefore it is not
known if the flowers displayed variation comparable to that observed among the plants from the west-
ern bolson. It is possible that H. Moller observed flowers from a small subset of the original series d
plants and thus was not aware of any color variation. Alternatively, the original plant material may have
been collected from a colony in which yellow throat color was locally fixed. In any case, we regard the
departure from the original description to be of minor significance considering that the variation of the
population includes yellow throat color.
CONCLUSIONS
ThepUnBfn,mtheSterraAguaChiqui,ade«n1bedasvar.c™ssKptambyMomberger(2007)differbyseve^
significant morphological characters from Mailer s original description of crassispinum. fn the Sierra Agw
ct 1 comprised of sparsely flocked plants; com-
sninpc lenegas) agrees quite closely with Moller’s original description. The plants have new
bottom n stout with a shghtly flattened cross-sectional shape; the
™^rn7dT"“T“'’ '"‘'“‘'''““"‘‘"8>>>»ndstatKilng away from the plant. ^
of heavtty nocked pUntsat the locationAZkmWnfCnatroCienegas. We oflerlplanat^for this discrep
■ts^'h H^nrich Momberger-s designated neotype (ULM-18572) seriously con
as the neotype is wan^nte^cTtl^^h taxonomic stability. The designation of another specimen
scription of 1925 Fimh^rmrsT- ^ ^ apparently no surviving plant material from Moller’s original de-
° T ■>='■>« cmssispinum should be attached to a herbarium specimen in onto o
richMoller'soriginaldescription. for its morphology agrees with Hetn-
Montanucd and Hoock, Astrophytum capricome var. crassispinum
redescription
1 (H. Moller) Okumura, Syaboten-no-
^ i. 1980s, HeinzSwoboda
:r2007:ULM-18572;™
K 1980s (see also PaWifek 2011).
A revised description of var, crassispin«. with additional morphological dau P'^
from the habitat, is herewith presented. Statistics of dispersion include t e samp e
followedbytheobservedlimitsinparentheses. r with age pale to dark leaf green; stem
Description.-Body single, young plants comprised of short hairs (tri-
«P to 35 cm in height; epidermis nude or only sparsely ^ ^ abundant
'Ptacs. Spines (-5)6-8(-ll) somewhat nattened, very stout, ^ ^ nght, directed
^.*own,hecom.nggtay-wh.te„ithage—
3way from Stem, with tips curved upward. Plants from ^ liotosinfoha&Bu^kl986).Flowerdiam-
tfrespinesalthoughthereisconsiderableindividualvariau^W^^^^^^^
«er is 77.8 mm . 3.5 mm (60-104 mm) in 44 flowers; ova^ inner tepals spatulate,
abundant wool; perianth vines from ’peach to orange to orange-red to red,
Circulate, canary yellow and satiny; flower throa y, filaments yellow, anthers
‘^rely sulfur-yellow or honey-brown; stamens bare y Pmit red or pale green with reddish
J;ghtlydarker; style with 7.5 .0.3(4-12)yeflowstigmaobesm ^
dehiscence basal in circular fashion; seeds usually conne
364
1 length; n = 30 seeds. Number of
niculus light pink, later drying. Seeds cap-shaped, 2.7 n
seeds per fruit: 146(96-280); n = 32 fruit.
Comparisons.— Variety crassispinum and var. niveur
to be closely related sister taxa. Var. niveum, however, is
giving the plant a snow white and felted appearance. Also the plants are slightly larger, reaching 50 cm in
height; maximum recorded height for crassispinum is 35 cm. As described above, var. crassispinum shows varia-
tion in the throat color of its flowers; in var. niveum the flower center is typically scarlet, but also rarely pure
yellow (Schatzle 1988). The flower of var. niveum is similar in diameter and number of stigmata to that of var,
crassispinum. Flower diameter is 74 mm ± 4.4 mm (60-80 mm) in 10 flowers; style with 8.4 ± 0.6 (6-12) stigma
lobes in 25 flowers. When the two varieties are compared at the same body size, var. crassispinum usually has
more areoles with spines. The greater density of spines may help protect the naked epidermis from solar radia-
sary protection from solar radiation. In both varieties, there are uncommon forms in their respective popula-
tions which have new spines with a whitish to yellowish color (Riha & Busek 1986).
Remarks on Type Locality.— The type locality for var. crassispinum, originally given as the Sierra de Parras,
has been a source of controversy for many years inasmuch as plants matching the type description have not
been found there. Some authors have speculated that Heinrich Moller may have deliberately misstated the type
locality as a diversionary tactic (Riha & Bu^k 1986). However, this seems improbable for a man of his profes-
sional standing; more likely, he would have simply withheld such information in the interest of plant conserva-
uon. Furthermore, in Moller’s description of var. aureum (in the same article with var. crassispinum) the origin
of the plants is correctly stated. His brother, Arthur Moller, would probably not have deliberately misled him
about the type locality, for he had provided accurate locality information for other new taxa sent previously. It
IS ^ssible that A. Moller inadvertently mislabeled the specimens or simply neglected to mention where they
had been collected, leading Heinrich Moller to assume the plants originated from the Sierra de Parras whichis
relatively close to San Pedro.
tentative restriction of the type locality deserves comment. Arthur Moller’s collection of plants in-
cluded both nude and sparsely flocked individuals in nearly equal proportions Assuming he collected plants
randomly from the habitat, the relative numbers of sparsely flocked and nude plants in his series would have
corresponded to their relative abundance in the habitat. At the present time, the populations at 42 km west of
Cuatro Cienegas are comprised primarily of nude plants (var. crassispinum) and densely flocked plants (var.
scarce. There are at least two possible explanations for the
at this locality, a natural change in relative abundance may
, , '‘•'“'ly Hocked plants increasing at the expense of the sparsely flockd
plants (iKrhaps taused by a strengthening of an incipient reproductive isolating mechanism between the two
vanettes), other explanation is that Arthur Moller collected his series from a different location, perhaps
farth„ west or southwestanda^y from the densely nockedpopulation. This would thenaccount for t^
u,^ T d h' ’’ T Po^Me explanation, we are aware of
^e undtsclosed, dtstant telity where nude and sparsely llocked plants of var. crassispinum have been oh
served, but no examples of var. niveum have yet been found.
is unknown and meni>
^ r -a cccs” (presumed hybrids?)
ruregreenphm.s.the;;^delnsTn.Cta^^T^^^^
even're™:^:, v^rs""^
365
under the type species, the classification system does not accurately reflect their relationships and evolutionary
history. Momberger (2011) has attempted to rectify this situation by the simultaneous use of the categories of
“subspecies” and “variety” for some taxa. Whether this system will be well received or rejected by taxonomists
remains to be seen.
In reviewing the descriptions of these taxa, we have determined that in some cases there are insufficient
morphological data for proper taxonomic diagnosis. It is evident that the “capricome group” is in need of care-
ful study in order to determine the number of valid taxa, their rank, interrelationships and evolutionary his-
tory. Future research should include not only detailed morphological analysis, but molecular genetic analysis
as well. In addition, the roles that physiographic features, \iz Desierto de Mayran, have played as vicariance
factors in the evolution of the “capricome group” members should be investigated.
ACKNOWLEDGMENTS
We thank Urs Eggli, Sukkulenten-Sammlung Zurich (ZSS), for providing herbarium records and digital pho-
tos of ZSS-019963. R. Russell, United States National Herbarium (US) made herbarium records of A. capricome
available to us. We also extend our gratitude to J. Busek and N. Turland for their assistance. Finally, we thank
A.M. Powell and an anonymous reviewer for their helpful, critical cor
1 earlier draft of the
REFERENCES
Baumann, H. 1992. Ungleiche Zwillinge von Astrophytum capricome (Dietrich) Britton & Rose var. niveum (
murafa, nudum. Kakteen And. Sukk.43(8):187-191.
Bernhard, U. and H. Hoock. 1986. Die Asfrophyfen von Cuatro Cienegas. Kakteen And. Sukk. 37(7)^141-147.^
BoSek, J. 1 981 . Astrophytum niveum und Astrophytum crassispinum. Die zwei schonsten Bisc o mutzen.
Sukk. 32(6)1138-140. . j c uu
Haage, W. and 0. SadovskV. 1 957a. Revision in der Gattung Astrophytum. Kakteen
Hoock, H. 2008. Astrophytum Lem. (Cactaceae). Schoendruck-media e. K., Landshut,
H. K.P. 2006. A«rophy,um capr^ome (Ca«ace«.) m,. -.,n gelbeo Bloten. K»k,ee
57(1):15-18.
ho, Y. 1981. The Cactaceae-classification and illustration of cacti. 'f
Kayser, K. 1933. Echinocactus (Astroph.) capricornis niveus var. n. Ka^nkunde ( )
Megata, M. 1 944. An account of the genus Astrophytum Lemaire. Mem. ColL g y ^ ^
Moeur, H. 1 925. Echinocactus capricornus Dietr. und seine Varietaten. Z Sukku en e ^ _
Momberger, P. 2007. Astrophytum capricome var. crassispinum (Cactaceae) K arung
Namens. Kakteen And. Sukk. 58(2):43-47.
P. 201 1 . Astrophytum capricome ss
e und Varietaten.
1,Oktober2011.
^P.20,,.N„v44a.oJansk6-as,roph,.um2poh^S^^^^^^
^ J. and j. Busek. 1 986. Astrophytum capricome var. niveum a va . P piora-Verlag, Titisee-Neustadt.
Wky, 0. AND B. SCHUTZ. 1 979. Die Gattung Astrophytum. Arten. Hybnd .
Federal Republic of Germany. Bliite. Kakteen And. Sukk. 39(3):
^Tzu, P 1 988. Astrophytum niveum (Kayser) W. Haage et Sadovsky
54-55.
Journal of the Botanical Research Institute of Texas 7(1)
BOOK REVIEW
Fred Dortort. 2011. The Timber Press Guide to Succulent Plants of the World. (ISBN: 9780881929959,
hbk). Timber Press, The Haseltine Building, 133 S.W. Second Avenue, Suite 450, Portland, Oregon
97204-3527, U.S.A. (Orders: www.timberpress.com, 1-800-327-5680, 1-503-227-3070 fax). $49.93, 344
pp., 755 color photos, 2 maps, 8 x 11 y^".
The author’s opening comment in his preface states, “The hrst time I visited a succulent plant nursery, I felt as
if I’d been handed the key to an odd but very interesting room. I saw plants there that resembled flowers, plants
that looked like rocks, and plants that fit no familiar description at all.” He was totally “hooked”! Or, as he
wrote, “. . . the owners themselves. . .were both extremely enthusiastic and extraordinarily knowledgeable
about their plants.” And, obviously, so was he.
Dortort was attracted to Berkeley’s University of California Botanical Garden, fascinated by their “scien-
tifically organized” plant collections and smitten with the desire to get involved. Soon he armed himself with a
paperback book and headed “to the great Southwest to look for the rarest, most localized plants in North
America.” This was the beginning of bis extensive and thorough work with succulents.
This is not only a beautifully illustrated book, it is carefully planned: descriptions are well written aid
thorough. While most of us think we’re fairly knowledgeable about succulents, the comprehensive inclusionof
both photographs and carefully written descriptions bring the reader into a far more extensive awareness of
the many diverse species. One can spend hours gazing at the incredible diversity of plants included in this
genre.
Chapter 1 explains and beautifully illustrates the diversity of succulents as they grow in nature. Chapter
2 gives an overall description and illustrations of succulents in cultivation. Chapters 3-29 deal with specific
FOSSIL FLOWER OF 1
MID-TERTIARY AMB
5 PROTIUM (BURSERACEAE) IN
:HE DOMINICAN REPUBLIC
George O. Poinar, Jr.
Journal of the Botanical Research Institute of Texas 7(1)
e to Lower Miocene Mam
y Group (Draper et al. 1994). The age of the amber is controversial, the oldest
i dates being 45-30 Ma, based on coccoliths (Cepek in Schlee 1999), and the youngest being 20-15 Ma,
n foraminifera (Iturralde-Vinent & MacPhee 1966).
e: HISPANIOLA. Dominican 1
It Oregon State University, Corvallis, Oregon 97331, U.S.A.).
n fully developed petals) 6.0 mm (Figs. 1, 2), a
lyx glabrous, cupular (Fig. 4), 1.4 mm long overall, sepals connate, the fused portion 1.0 mm, the lobes broadly
triangular, acute, 0.4 mm long, 0.7 mm wide at base; petals 5 (1 missing and 1 truncated near base), lanceolate
from a broad base, acute, spreading, distal portion recurved, length 2.4 mm including the recurved part, the tip
inflexed (Fig. 4), margin papillate (Fig. 4), abaxial surface glabrous, adaxial surface minutely papillate; stamens
10, strongly incurved, of 2 different lengths, the antepetalous ones up to 1.1 mm, the antesepalous ones up to
0.9 mm, anthers lance-linear, basifixed (Fig. 2), 0.5 mm long, laterally dehiscent; disc intrastaminal, glabrous,
annular, swollen, 1.7 mm in diameter, the central depression 0.7 mm in diameter, occupied by a reduced pistil-
lode (Fig. 2), ovary ovoid, glabrous, ca. 5-lobed, style fused, stout, 0.15 mm long, distally 10-lobed or sulcate
(Figs. 1, 2), floral pedicel papillate at base, otherwise glabrous, 2.8 mm long (Fig. 3).
Etymology.— From Greek “kallos,” beauty, and “anthos,” flower.
DISCUSSION
Petals of Protium may be either erect or reflexed at anthesis (Engler 1931), and the fossil is in the latter category.
Petals are vanously glabrous or pubescent in the genus, e.g., villous abaxially and adaxially in P. alvarevcmm
Daly & P. Fine and adaxially short-papillate in P. caroknse Daly. The fossil resembles the latter species in this
respect as well as in its widely recurved petals, but differs in other features (e.g., flowers 5-merous versus
4-merous). The form of calyx seen in P. callianthum is well represented in the genus, as described for many of its
species by Swart (1942). Anthers with basal rather than dorsal insertion are common in the genus and charac-
tenze P. callianthum (Figs. 2, 3). They are illustrated for P. altsonu Sandw., P. bahianum Daly, and others (Daly
1989 1992a). Stammate flowers possessing a thick intrastaminal disc whose central depression contains a te-
r“‘T Pfe'il. common In Prolinm (see ffluslrations of P. b<Mmm
IDaly 1992a| as well as P. mom Daly and other taxa [Daly I992bl). The ease with which the fossil keys to Par
hum onnoral traits (Swartl942; Daly etal.2011).as well as its Similarity to the various illustrated taxa men.
tioned above, strongly support its assignment to this genus
Several features of the fossil are worth noting. Papillate margins allow postgenital fusion hetweea the
valvate petals before they finally separate at anthesis. This occurs through imerd^tion of the papillae, m
xplained by Bachelier and Endress (2009). A si
1, which they also mention, is not
was notedforPrmiummorii by BacheherandEtnhess (2009) from noralbud'cr^s^s^cinrHowevert^
ers rom t e escnption of tbe species by Daly (1992), who gives 1.7-1.83 and 1.85-2.05 mm for the antepetal-
mtes,mlousones,respectively.Other species forwhichthedescriptions mention
,tenetal™,s p p dvareziamm (Daly & Fine 2011). The fm
ous than antepetalous
quency, in Protium, of the
Tribe Prot
inoe urotieae (the Protium Alliance per Daly et al 1201 1 D i« , ii j j • t mnlecular
.^s(Clarksonetal.2002;Weeksetal.‘’2005;{h:it::r]:;:r^^^^^
.bywhichtodistingnisham„ngthe7:rra'?™;tir^™
Chambers and Poinar,/
genus Crepidospermum by the lack of snaU-shaped trichomes (Solereder 1908)
usually differsLnxProtL and from thefossilinhaving the anth^
uients, the disc and pistillode fused into a conical “ovariodisc” m staminate owers, an
nate (Daly 1989). . r., . uv,
FoUowi„gh.descrtp«o„otPr..u™car„,e„«,D,V(~
Ulsect that feeds on the flowers. He found that m a majontyo ^ » f the remaininenerianth Other
whicheatstheinterioroftheflowerbutdoesnotpreventregularenlargement . ...yi^of amber con-
colleaiomof the species have developing fruits with no ,he stamens and top of
^tningthe type fossil ofP. ofsome kind. Whether the
•hedisc and pistilseen. to have been eaten, quite posstblybyanh*^^^ ^^^
‘usect/flower relationship described for P. carolense might also have charac
«nknown,butitdeservesn.entionforitspossiblebottm^
Protimn callianlltum is distinct from other « D„„i„ican Republic but differs from P, culli-
mo^tn species of Protium, P. gluucescens Urb., endemic m Cuba also have 4-merous
»*«tn its 4-merous, rather than 5-merous,flow^^The^
»»»ers, while P. attetmutum (Rose) Urb, of the Lesser Anulies, wttna op t-
a™ in its ovate, dotsifixedanthetsand 5-lobed (Becerra et al. 2012; De-Nova et al.
The two most recent molecular Pbylo8'«“”“^'““f®";:^'::“;es (Clarkson 2002; Weeks etal.
^912), although emphasizing the genus Bursera, go beyon p
372
2005; Thulin et al. 2008) by increasing the number of sampled species and proposing time-calibrated phylog-
enies of the family. According to the calculations of Becerra et al. (op. cit.), Burseraceae are over 92 Ma old, and
the evolutionary proliferation of many modem genera occurred in the period of 30±10 Ma (their Fig. 3, p. 339).
De-Nova et al. (op. cit.), however, give the age of the family’s crown group as 64.92 (60.33-69.67) Ma and of the
Bursera crown group as 49.43 (45.38-53.77) Ma. In a phylogenetic study of Protium by Fine et al. (under review),
the split between this genus and Bursera is dated as ca. 52 Ma, and the diversification of the Protium clade be-
gins ca. 20 Ma. The deposits of amber from the Dominican Republic are not precisely dated (see Materials and
Methods), the youngest estimate being 20-15 Ma. The large clade to which P. callianthum probably belongs,
containing, among others, the extant Caribbean species of Protium, diverged around 15 Ma, a figure that
clearly would favor the younger of the 2 estimates for the age of the amber. Protium callianthum may be still
older, on the stem of this clade, since it has no synapomorphies that match those in the larger, inclusive clade
(P.Fine,pers. comm.).
ACKNOWLEDGMENTS
We thank David Harris for his taxonomic advice and Peter Endress for pointing us in the direction of an ap-
propriate systematic placement for the fossil. Douglas Daly and Paul Fine provided valuable review comments
and suggestions, as well as information from unpublished research. Alex Brown kindly made available the
tise by Swart.
REFERENCES
Bacheuer, J.B. AND RK. Endress. 2009. Comparative floral morphology and anatomy of Anacardiaceae and B
(Sapindales), with a special focus on gynoecium structure and evolution. Bot. J. Linn. Soc. 159:499-571.
Becerra, JX, K. Noge, S. Ouvier, and D.L Venable. 2012. The
Burseraceae. Taxon 61:333-343.
Chambers, K. L. and G.O. Poinar, Jr. 2010. The Dominican amber fossil Lasiambix (Fabaceae: Caesalpinioideae?) is
(Chrysobalanaceae). J. Bot Res. Inst. Texas 4:217-218.
Chambers, K.L, G.O. Poinar, Jr., and A.E. Brown. 2
Bot. Res. Inst. Texas 5:457-462.
il flower of Perseo (Laurat
n Tertiary Dominic
i Protieae and Canarieae. Brittonia
„ . ^ '' Mid-Tertiary fossil flower of Swietenia (Meliaceae) in Dominican amber. J. Bot
Res. Inst Texas 6:1 23-1 27.
ChwK.UG.O. Poinar, Jr., AND A.S.CHANDERBAU. 2012. Trepfosfemon(Lauraceae),anew genus of fossil flower from Mid-
Tertiary Dominican amber. J. Bot. Res. Inst. Texas 6551-556
Clarkson, JJ., M.W. Chase, AND M.I
sequences. Kew Bull. 57:183-
Daly, D.C. 1989. Studies in neotropical Burseraceae. II Generic
41:17-27.
Daly, D.C. 1992a. Two new taxa of Protium from eastern Brazi
713-719.
Daly, D.C. 1992b. New taxa and combinations in Protium Bur
44:280-299.
Daly, D.C. 1998. Two new species of Protium from French Gui
50:517-523.
Daly, D.C., M.M. Harley, M.-C. MarUnez-Habibe, and A. Weeks 201 1
iring plar
Daly, D.C. AT
generali:
De-Nova, J.,
Studies in neotropical Burseraceae XVI. Syst. Bot. 36:939-949.
historical construction of species-rich Mesoai^^n ti^’llv dr^T*^' Tf
(Burseraceae, Sapindales). New Phytol. 193:276-287. ^
n neotropical Burseraceae Vlii. enuu...
In: K. Kubitzki, ed. Families and genera of
lialistand
Draper, G., P- Mann, and J.F. Lewis. 1 99^
tion.The University of the West Ir
Engler,A.1931.E
Leipzig. Pp. 405-456.
Fine, P.V.A., FZ Zapata, and D.C. Daly (In Review). The evolution ai
important lineage of tropical rain forest trees. Evolution.
Iturralde-Vinent, M.A. and R.D.E. MacPhee. 1966. Age and paleogeographic origin of C
1850-1852.
PoiNAR, G.O., Jr. ]99^.Hymenaea protera sp.n. (Leguminosae: Caesalpinioideae) from Dominican amber has j
finities. Experientia 47:1 075-1 082.
PoiNAR, G.O., Jr. and R. Poinar. 1 999. The amber forest. Princeton University Press, Princeton, NJ.
PoiNAR, G.O., Jr. 2002a. Fossil palm flowers in Dominican and Mexican amber. Bot. J. Linn. Soc. 1 38:57-61 .
Poinar, G.O., Jr. 2002b. Fossil palm flowers in Dominican and Baltic amber. Bot. J. Linn. Soc. 1 39:361 -367.
PfflNAR, G.O., Jr., K.L Chambers, and A.E. Brown. 2008a. Lasiambix dominicensis gen. and sp. nov., a eudicot flower i
can amber showing affinities with Fabaceae subfamily Caesalpinioideae. J. Bot. Res. Inst. Texas 2:463^ •
Poinar, G.O., Jr., K.L Chambers, and A.E. Brown.
cence in Dominican amber. J. Bot. Res. Inst. Texas 2:1 1 67-1 1 73.
ScHLEE, D. 1999. Das Bernstein-Kabinett. Stu1
SOEREDER, Ft. 1 908. Systematic ani
Stevens, P.F. 2001 o
Swart, JJ. 1942. A monograph of the genus Protium a
39:211-446.
Thuun, M., B-A. Beier, S.G. Razafinandimbison, and H.I. Banks. 2008. Ambilobea. a
of Aucoumea, and comments on the tribal classification of the frankince
J. Bot. 26:21 8-229.
Vol.l. Clarendon Press,
n Madagascar, the position
IS in Anacardiaceae. Blumea 51:165-195.
374
ir have you ever heard of Mr. Falkenberg?” (Fore-
BOOK REVIEW
Steve Manning. Foreword by Rudolf Jenny. 2010. Discovering New World Orchids. (ISBN: 978-0-9565594-0-
1, hbk). Steve Manning, 4 The Cedars, Nantwich, Cheshire, CW5 5GZ, U.K. (Orders: Steve Manning,!
The Cedars, Nantwich, Cheshire, CW5 5GZ, U.K.; orchidsmann@uwclub.net). £45 (approx. US $68.25),
669 pp., color throughout, 8 1/4" x 11
Rarely do we find such an exceptionally large, carefully researched and documented, beautifully illustrated
botanical volume. Where else do you find a chapter titled “POLITICS, INTRIGUE, and JEALOUSY— The story
of Masdevallia uniflora”?
Even the preliminary pages are fascinating:
• “Do you have any idea who Baron V
ward, Rudolf Jenny)
• “It is from South America that we obtain the best orchids we have,
might be related of the adventures experienced by collectors of Orchids.” (Introduction, F.W. Moore,
Orchid Review, June 1903:163)
By all means, read the Introduction first. In part: “The amateur enthusiasts who were willing to spend
fortunes on orchids and other tropical plants were often subject to that green-eyed monster Envy— as were the
sionals were incompetent, not all amateurs were self-centered and aln^LI^rere w^s much co-operation
between amateurs and professionals, between traders and customers.”
Each chapter is well written and contains excellent information. There are beautiful illustrations of
plants, photos, early illustrations of settlements and mines and flowers, and often maps, comments, bits of his-
tory, etc.
This is an excepiinnal volume, with exquisite illustrations, and carefully documented information. Ills
also a lascinating, “canT put it down" read Some readers might he a bit daunted hy a large, 669-pa|e vofomt.
but tt IS absotately beautifully presented, carefully researched, historically informational, and an astonishing
florrcultural history of the New World orchids.-HelenJeude, Volunteer und Assistant Editor, Botanical Rorart
Institute of Texas, Fort Worth, Texas, U.SA.
HIPPOCRATEA VOLUBILIS (CELASTRACEAE) IN COTUl COPAL
FROM THE DOMINICAN REPUBLIC
George O. Poinar, Jr.
Kenton L Chambers
376
Published estimates of the age of Cotui copal have ranged from Miocene to as young as 1000 ybp or less.
Our conclusion that the Hippocratea flowers are assignable to the extant species H. volubilis strongly supports
the latter, younger age, which was obtained from carbon-14 dating in 2 laboratories (“Beta Analytic” in the
United States and “Harz aus dem Tennengebirge” in Germany).
Hippocratea pollen is well represented in the fossil record (Muller 1981), hut to our knowledge these are
the first fossil flowers of the genus to be discovered.
MATERIALS AND METHODS
The specimens were collected from the upper 8 inches of soil at Zambrana Abajo near Los Ranchos in the vicinity
of Cotui, a village located in a valley between the Cordillera Central and Cordillera Septentrional, Dominican
Republic. The material is clear with a yellow tinge. Many samples are collected by children who follow theplows
during the planting of maize (personal correspondence from the late Jake Brodzinsky, 20 September 1987).
The age of fossilized resin from Cotui is unclear. Because of its light color and fairly soft texture, it has
been referred to as copal, a term used for semi-fossilized resin that possesses specific physical properties re-
lated to color, hardness, melting point, and solubility (Poinar 1992). Age estimates range from under 1000
copal, of several
1999).
nee spectroscopy (Lambert et al. 2012). In support of the older age was the discovery, in the
Comparing the maturity of fossilized resins with that of their enclosing bedrock can provide information
on the maturity, relative age, and biostratinomy of amber and copal. Such a study was conducted on amber and
Cotut copal from different mines in the Dominican Republic (Poinar & Mastalerz 2000). Maturity of the bed-
rock was determined by vitrinite reflection and that of the fossilized resin by FTIR (Fourier Transform Infrared
Analysis). The study found that Cotui copal showed a lower degree of maturity than that of the matrix rock,
indicating that it had been washedinto the older bedrock fairly recently and probably representsdeltaicdepos-
us. Further analysis of Cotui copal revealed that the plant source of the resin was Hymenaea courbaril (Mas-
talerz & Poinar, unpublished observations), a tree species that still exists on Hispaniola.
ola^Tm nTn ^ shallowly cupulate (Fig. 2), sepals 5,
lanceolate ellint’ 2 ^ blunt-tipped, abaxially papillate, petals 5, spreading,
lanceolate-elhptic, 2.1 mm long, 1.2 mm wide, 7-9-nerved, slightly overlapping at the base (Fig. 5), tip «-
cu^ed,abaxialsurfacetomentulose with short, curved trichomes,adaxm^
d d T r otherwise glabrous and dis-
mctlyvemeddownwardtothebase(Figs.l,4,5),marginscilia^^^^^^^^^ 1 2) disc 1.3 L high, 1.7 mtn
3) the^uScTdr^r "7 ’ above the mid-line (Fig^ 1.
biiH O^mm d recurved (Figs. 1, 3, 4), filaments 0.5 mm
mlrngplZ™"
For the purpose of comparison, we
South America and the Caribbean re
discussion
assumed that Hippocratea volubilis, which is widespread in Central and
:gion, is likely to be the extant species most closely related to the fossil-
377
elongated style, as illustrated by Matthews
and Endress (2005, figs. 49, 50). The adaxial
pubescence of the petals consists of a con-
spicuous band or arc of villous trichomes,
which is essentially similar to the pubes-
cence pattern in herbarium specimens of H.
volubilis. Such trichomes were, in fact, cited
by Smith (op. cit., p. 357) as a diagnostic
trait of the genus. The petals of the flowers
appear from photographs to be slightly con-
nate, but careful microscopic study shows
them to be basally imbricate. The floral disc
is divided into upper and lower segments,
thus differing from Smith’s illustration of H.
volubilis (op. cit., p. 357). However, similar
2-parted discs are often present in herbari-
um specimens of that species, including
many from the Caribbean region. In A. & P.
Liogier 30929, Puerto Rico (US2996523),
for example, the upper and lower disc seg-
ments are sharply differentiated, with both
parts being papillate, as in our fossils. In
both C.V. Morton 7839, Honduras
(US202361) and R.S. & E.S. Howard 9817,
Dominican Republic (US2228804), on the
other hand, the upper half of the disc is
lower half. The upper section of the disc is
sometimes reduced to a narrow papillate
band below the anthers (e.g., D.K. Christo-
pher et al. 69, NY s.n., Puerto Rico). A pecu-
liarity of petal pubescence in both the mod-
em and fossil flowers is that a zone of papil-
late trichomes may or may not be present
proximal to the adaxial band of villous tri-
chomes. This is illustrated in Fig. 1, al-
though
In the conservative tj
^ontothe
Rorida (Long & Lakela 1971). The pubescence ol penar
sils, as described above, falls within the variauon that ’
volubilis. With the exception of petal length and width,
^nges given for H. volubilis in Smith’s species descnptio
'’^S-'’0^)Se'UsH.v.««Us.HeUn.«ea.HU
extends north as far as the Everglades of
disc, androecium, and peduncle in the Cotui fos-
ve observed in available herbarium specimens of H.
the sizes of floral parts in the fossils are within the
1 (op. cit., p. 360). The petals are slightly shorter (2.1
379
M.A. AND R.D.E. MacPhee. 1 996. Age and paleogeographic origin of Dominican amber. Science 273:1 850-
Endress. 2005. Comparative floral structure and systematics in Celastrales (Celastraceae,
Parnassiaceae, Lepidobotryaceae). Bot. J. Linn. Soc. 149:129-194.
Muuer, J. 1 981 . Fossil pollen records of extant angiosperms. Bot. Rev. 47:1-1 42.
PoiNAR, G.O., Jr. 1 992. Life in amber. Stanford University Press, Stanford, CA.
PoiNAR, G.O., Jr. and M. Mastalerz. 2000. Taphonomy of fossilized resins: determining the biostratinomy of amber. Acta
GeoLHispanica 35:171-182.
Popov, YA. 1 987. A new species of the bug genus Empicoris Wolffe from Dominican copal, with the redescnptKjn of £
nudusMcAtee & Malloch. Stuttgarter Beitr. Naturk. Ser. B 134:1-9.
region. Molec. Phylogen. Evol. 7:145-147.
V. M.W. Ch«e, S.B. Hoor, CM. Mmton, D.E. Sans, C. Baw M.F. F.r, A. De B«j* S. Sous™, ano Y.-L Ou. 2000.
Phylogenetics of flowering plants based upon a combined analysis of plastid otpB and rtnrt gene sequences. Syst
Biol. 49:306-362.
ScHLEE, D. 1984. Bernstein-Neuigkeiten. Stuttgarter Beitr. Naturk. Ser. C 18:1-100.
SoftEE, D. 1990. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C 28:1-10
Simmons, M.P. 2004. Celastraceae. In: K.Kubitzki,ed. The families ai ’
er-Verlag, Berlin. Vol. VI. Pp. 29-64.
Simmons, M.P., C.C. Clevinger, V. Savoleinen, R.H. A
ferred from phytochrome B and morphology. Amer. J. Bot. 88:313-325.
nrDNA, phytochrome B, atpB, rbcL, and morphology. Molec. Phylogen. Evol. 19:353-366.
Smith, A.C. 1940. The American species of Hippocrateaceae.Brittonia 3:341 -555.
STtVENs, P.F. 2001 onwards. Angiosperm Phytogeny Website, Version 1 2, July 201 2 [updat sir
org/MOBOT/research/APweb/>, accessed 21 January 2013.
BOOK NOTICE
i. 2010. On Harper’s Trail; Roland McMillan 1
i. (ISBN: 978-0-8203-3100-3, cloth). The University of Georgia Press, 4435 At-
lanta Highway, Athens, Georgia 30602, U.S.A. (Orders; www.ugapress.org, 1-800-266-5842, 1-706-425-
3061 fax). $44.95 hbk, $24.95 pbk, 296 pp., 20b/w photos, 1 illus., 1 map, bib., index, 6" x 9".
From the publisher: “Roland McMillan Harper (1878-1966) had perhaps ‘the greatest store of field experience
of any living botanist of the Southeast,’ according to Bassett Maguire, the renowned plant scientist of the New
York Botanical Garden. However, Harper’s scientific contributions, including his pioneering work on the eco-
logical importance of wetlands and fire, were buried for decades in
1, Harper’s reputation as a scientist has often b
:. With this book, Elizabeth Findley Shores provides the first full-length biography of the ac
st, documentary photographer, and explorer of the southern coastal plain’s wilderness areas.
Incorporating a wealth of detail about Harper’s interests, accomplishments, and influences. Shores fol-
lows his entire scientific career, which was anchored by a thirty-five-year stint with the Alabama Geological
Survey. Shores looks at Harper’s collaboration with his brother Francis, as they traced William Bartram’s route
through Alabama and the Florida panhandle and as Francis edited the Naturalist Edition of The Travels of Wil-
liam Bartram. She reveals Roland’s acquaintance with some of the most important, and sometimes controver-
sial, scientists of his day, including Nathaniel Britton, Hugo de Vries, and Charles Davenport. Shores also ex-
plores Harpers personal relationships and the cluster of personality traits that sparked his interest in genetic
predestination and other concepts of the eugenics movement
Roland Harper described dozens of plant species and varieties, published hundreds of scientific papers,
an ma e notable contributions to geography and geology. In addition to explaining Harper’s eminence among
southeastern naturalists, this story spans fundamental shifts in the biological sciences-from an emphasison
field observation to a new focus on life at the molecular level, and from the dawn of evolutionary theory to the
modem synthesis to sociobiology.”
Elizabeth Findley Shores is a v
Boston University and the University of Arkansas, Uttle Rock.
. She holds degrees in history from
i. Bot Res. Inst Taas 7(1): 380. 2013
KARYOTYPES OF MUHLENBERGIA RIGIDA (POACEAE: CHLORIDOIDEAE)
FROM NORTH CENTRAL MEXICO
O. Rosales Carrillo
IPNCIIDIR Durango
Sigma 1 1 9 Fracc. 20 de Nov. II
Durango, Durango, 34220 MEXICO
M.S. Gonzalez Elizondo
IPNCIIDIR Durango
Sigma 1 1 9 Fracc. 20 de Nov II
Durango, Durango, 34220 MEXICO
N. Mayek Perez
IPNCBG
Blvd.del Maestro s/n esq. Elias Pina
Col. Narciso Mendoza
Reynosa, Tamaulipas, 88710 MEXICO
Y. Herrera Arrieta
IPNCIIDIR Durango
Sigma 1 19 Fracc 20 de Nov. II
Durango, Durango, 34220 MEXICO
N. Almaraz Abarca
IPNCIIDIR Durango
Sigma 1 1 9 Fracc. 20 de Nov II
Durango, Durango, 34220 MEXICO
Al. Laforga Vanzela
Laboratorio de Citogen^tica e Diversidade Vegetal
86051-970, Londrina, PR, BRAZIL
INTRODUCTION
.*s„„eof.he.arges.gene«cf.hePoaceaetaily,wi.happn«*
>»ose, 150 are distributed mainly in the temperate forests and ^ Peterson & AnnaWe
only six species are endemic to South Asia (Clayton & Renvotze 1986; f ^
Ml; Peterson & Orttz-Diaz 1998; Petenmn ”™j“^;,r!“merica“and Northern Mexico, some of
33species ofMuhlenbergia native to the Southern ^
them even reaching Central America (38) and South Amenca (25)^lh J
‘n Mexico (Beetle 1986; Espejo-Serna et al. 2000; Espejo ^ ^ j 2001; Peterson 2003; Peterson
neta&delaCerdaLemusl995;HerreraArrieta&Petersonl992^Petersone
&Valdes-Reyna 1999). Muhlenl^rgiaisagenusvdt^^^^^^
tive characters, ranging from very small plants (5 to ^ rieida(Kunth)Trin.haspopulations
ones (150-300 cm tall) in M.robusta(E.Foum.)Hitchc.Furthermo ’ ' ^
that differ in plant habit as well as in the shape and size of their inflorescence, which can be compact (dense) or
loose (open), large or small. This creates taxonomic confusion and problems with the specific delimitation.
Few chromosome studies have been carried out on the genus Muhlenbergia. Soderstrom (1967) stated
that Avdulov (1931) was the first who reported cy tological information as a basis for a new classification system
of Poaceae. Although Sodertstrom (op. cit.) did not carry out cytological studies, he does mention the results
from several studies which reported a base chromosome number for Muhlenbergia of x = 9, 10, or 21. These
studies included the 2n chromosome number for the species of sect. Epicampes. Peterson (1988) studied the
chromosome number of 25 annual Muhlenbergia species, reporting for the first time the chromosome number
of nine of them. Noteworthy among these are Muhlenbergia biloba Hitchc. and M. shepherdii (Vasey) Swallen
since they are n = 8. Herrera (1995) published the chromosome number of three species of the M. montm
(Nutt.) Hitchc. complex [(M. montana, M. quadridentata (Kunth) Trin., and M. y^irescens (Kunth) Trin.)], re-
cording for the first time the chromosome number of M. quadridentata, the disploid chromosome number ofM.
virescens and confirming the tetraploid condition of M. montana. Gould (1966) carried out karyotyping of 60
genera and 149 species of Mexican grasses, eight of which belong to Muhlenbergia. However, M. rigida was not
included in Gould’s study.
Muhlenbergia rigida is a grass with widespread distribution in the SW United States and ranges south to
Central and South America. In Mexico it is found from Baja California to Chiapas. The grass can be found
growing m pine-oak forests and grasslands, sporadically at the edge of roads, and at elevations between 1280-
2550 m (Herrera Arrieta & Peterson 2007).
This capacity to grow in such extensive areas with diverse habitats translates into a great morphological
diversity within the species (rigida). This diversity of traits is reflected mainly in the shape and size of leaves
and inflorescences, with vanation among the type of inflorescence (compact or loose) being particularly noto-
nous. However, there is no correlation between morphotypes and habitats or geographical distribution as k-
quently both^orphotypes^can be found growing in the same location. On the basis of the great morphological
study, the aim of the present study is to compare the karyotyproTt^moi^hm^^^^^ of ^
rigida located in north central Mexico. This comparison will reveal any relationship between the karyotype
and the morphological differences within the species as well as other relevant variations that allow the distinc-
tion of groups among the studied populations, or otherwise distinguish between the two morphotypes.
MATERIALS AND METHODS
I
d was collected from 27 locations (Fig. 1) and 30 populations of Muhlenbergia rigida, following
^ ^ Chihuahua, Durango, Zacatecas, Aguascalientes, Guanajuato, and Jalisco-
Mexico (Table 1). Live plants were collected (bunches) from natural populations and cultivated under —
and localities(mcludmggeographical coordinates and elevation) in which thestudiedplam^
Karyotyping
for 12 hours a.4C followed IvhydrolysUwUh three passes using dLuied water at 30-min„.eimc™^^
hen acid hydrolyias using O.IN HCI, and citrates buffer for 30 miLtes then a final digestion usinga 41tcd-
lulase enzyme mixture "Onozuka R-10 (Serxa)” and 1% pectin^ Y 23U h Ph rrcal) for 1 h-al
left to^ for^l^v removed after the samples were kept at -84°C. Preparations we«
CountinJatlLt H Chromosomes were stained using the acetocarmine reagent
gr y ,an I^ierptetationofthechromosomes(in7-10metaphaseslides)wascarriedoutusingan
383
software hup;//www.colostaie.edu/
Axioskop Zeiss microscope fitted with digital camera.
^tsmiology/MicroMeasure) was used for measuring the lengt o e the nomer
The ratio of the chromosome arms (r) (long arm: ^^ort arm) ^ ^
for describing chromosome morphology was that propt^ ^ ^ 00-6.99). Stebbins’ stanc
:00;l-69),sm = submetacentric(r=1.70-2.99),andst = ^
’on (1938) was used for classifying chromosomes accordi g
5-9 pm, medium-large; >9 pm, large.
In order to carry out the quantitative <
, of the karyotypes t
384
23°45'51.6”N y 104‘’25'29"W, 2014 m
Parte alta de la Canada el Caj6n, Santa Maria, El Oro, Durango, 25°24'42.8"N y 104'’57'28.4"W, 1914 m
Entronque Otinapa-Autopista Durango-Mazatlan, Durango, Dgo. 23°58'35.0"N y KM'Se'SOJ'W, 2446 m
Km 25 Autopista Durango-Torreon, Durango, Dgo. 24°1 1'12.0"N y104‘’29'31.7"W, 1864 m
Por la carretera 1 05, rumbo a la presa Bayacora, Durango, Dgo., 34°54'44.1 "N y 1 04°44'46.9"W, 21 85 m
Cerro de Los Gallos, Aguascalientes, Ags. 2r40'03.6"N y 102'’1 3'1 5.8"W, 21 91 m
Cerro de Los Gallos, Aguascalientes, Ags. 2r40'03.6"N y 102‘‘13'15.8"W, 2191 m
Entronque a Milpillas, Jesus Maria, Ags. 2r55’28.6"N y 102“33’57.7''W, 21 86 m
Cerro El Roble, Jesiis Maria, Ags., 2r47'30.7"N y 102°31 '26.3"W, 201 9 m
6 km antes de la caseta de cobro por la autopista Aguascalientes-Zacatecas, Guadalupe, Zac.,
22’’39'19.6"N y 102°26'45.5"W, 2305 m
jr la Ciudad de Guerrero, Guerrero, Chih. 28“30'1 5.8"N y 107°29'00.3"W, 2045 m
c, Chih. 28“24'09.4"N y 107“34'57.0"W, 2277 m
c, al norte deTemochi, Chih., 28‘’21 '20.7"N y107“49'26.0"W, 2075 m
calculated: (1) total chromosome length (LTC); (2) mean chromosome length (LMC); (3) centromere index
mean (short arm/total chromosome length x 100 [Cl]); (4) intra-chromosomal asymmetry index (Al) =
1- [I(b/B^), where b and B are the measures of the short and long arm of each homologous chromosome pair re-
spectively and n is the total number of homologues; (5) inter-chromosomal asymmetry index (A2) = s/x, where
s IS the standard deviation and x is the mean chromosome length; (6) Paszko Index Al = CVCL x CVCVlOO
where CVCL= (SC17XCL) x 100 is the relative variation of chromosome length, CVCI = (SCl/XCl) x 100 is
the relative vanation of the centromere length, respectively, XCL is the mean chromosome length and XCl
IS the Cl mean. Karyotype asymmetry was determined using Stebbins' categories (1971), Al and A2 indite
(Romero-Zarco 1986) and the Al index (Paszko 2006). The Al index is a quantification of Stebbins' asymme
try categories. It ranges between 0 and 1, and these are low when chromosomes tend to be metacentric. Basic
mtetpretanonofAlvaluesdeterminesthatthehigherthe value, the higheristheheterogeneityofchm^^^
length and/M the centromere index m a studied karyotype (Garcfa-Barriuso et al. 2010).
In the Ideograms, homologue chromosome pairs were ordered according to iheir length in decieasiC
size order. Four to seven metaphase cells were measured from various slides in order to obtain an average f«
constjmetion of the ideograms. Measurements were compared usingANOVA. The TCL, Cl, AlandAliodices
as well as chromosome number were considered
Rosales et al., Chromosome studies in Muhlenbergia
Clustering analysis of the karyotype data was carried out in order to examine karyotype similitude
among populations. A data matrix of 30 OTUs (operational taxonomic units) and five variables was construct-
ed. The following variables were used: LMC, Cl, Al , A2 and chromosome number. The first four variables were
used because they are not influenced by chromosome number. Nevertheless, ploidy level was also used since
different characteristics of various populations of Muhlenbergia rigida morphotypes are being compared. The
STATISTICA v.7.0 (StatSoft, 2004) software package was used to normalize the data matrix, calculate the aver-
age Euclidean distance, and generate an UPGMA dendrogram.
Also, in order to evaluate the contribution of each karyotype parameter to the population clustering, the
entities were subject to a principal component analysis (PCA) based on the 30 OTU data matrix and the afore-
The chromosome counts obtained from radicular cells of Muhlenbergia rigida in this study are consistent with
the numbers reported by Herrera and Peterson (2007) and Soderstrom (1967), both of which documented 2n =
40 and 44. This study recorded 2n = 30, 40, and 44; marking the first report of triploidy in this species. It is
noteworthy to mention that the 2n = 40 and 2n = 44 counts were found in the compact panicle morphotype
while the 2n = 30 count was present in the open panicle morphotype.
Our results support previous suggestions that the basic number of Muhlenbergia is x = 10, with 2n = 40
occurring in the majority of the species. Polyploidy is equally distributed in the morphotypes of M. rigida stud-
ied here: 43.33% of the plants are tetraploid (2n = 4x = 40), 43.33% are triploid (2n = 3x = 30) and only 13.33%
are disploid, derived from tetraploid (2n = 4x + 4).
We report for the first time the karyotype formula of 30 chromosomes in M.
the ideograms representing the three ploidy levels found (Fig. 2)
As a whole, the karyotypes of the species analyzed were composed of metac
trie (sm) chromosomes, with the former being predominant. The formulae among triploid populations were
30m (six populations), 29m + Ism (three populations), 27m + 3sm (two populations), and 25m + 5sm and 22m
+ 8sm in the remaining two populations.
Tetraploid populations were present in the following manner: 40m (five populations), 39m + Ism (three
populations), 38m + 2sm, 37m + 3sm, 35m + 5sm, 32m + 8sm, 34m + 6sm, 32m + 8sm m five popu ations.
Disploid populations had the following karyotype formulas: 44m (two populations) and 30m + 14sm, 35m +
9sm for the remaining two populations (Table 2). , ^ .
Chromosomes were sLu and medium-small in size (between 1.0 and 3.83 ^m), according to Stebbms
(1938). The mean chromosome length (LMC) ranged between 1.31 pm and 2.62 pm. The centromere tndex
variedfrom 12.94 to 21.76. . .. , fo KVninc
, , , nnH fall into categories lA and IB ot Stebbms
In general the karyotypes were moderately symmetrica
(1971).
The UPGMA dendrogram constructed with karyotype similitudes (Fig. 3) shows three
divided into subgroups. The first group is composed of three subgroups wuh gro^ “yo “3 “gs/
populations 4098, 4177, and 4016. Group 1-2 was formed by populations 4215 4170,
Group ..3wascomposedolpopulations4227 3982, 413942^^^^^^
Sit IT " i tTteTcroTelnSTcTraXw^^^ fimtroupTntains
to the morphological characteris v, c nnp disnloid and one tetraploid population, while
dtapbid and tetraploid populations, the second group has one dtsptad P
•Ito third group is completely composed of triploid populations that have open p
386 Journal of the Botanical Research Institute of Texas 7(1)
Measurements from triploid and tetraploid plants from the same 1
suiting in significant differences (P < 0.05).
itically compared re-
The results obtained from our study of Muhlenbergia rigida morphotypes partially confirms the previously re-
ported chromosome numbers for this species by several authors.
Of the 30 populations examined, 13 were found to be triploid, this being the first report for triploidy m
the species. Apart from the triploid cytotypes this study also found tetraploid and disploid plants, confirming
the results of previous studies. The triploid and tetraploid cytotypes are present in high frequency (86%) while
disploidy(2n = 44)hadlowfrequency,beingfoundinonlyfour^pu,^^^^^^^
ThisstudypresentsploidylevelsfoundinM.rigidapopulationsinnorthcentralMexico.Italsoshows,for
the first time, the karyotype and ideograms of the two Muhlenbergia rigida morphotypes (dense and loose
panicle) as well as the three cytotypes within the species.
Polyploidy has played an important role in the evolution of many eukaryotes (Solti
roiypioiuy tias played an important role in the evolution of many eukaryotes (Soltis et al. 1999), and the
inajontyofangiosperms(approximately7(m)have shown polypl^^^^
terson 1994). It is highly nrobahle that fhp w 1 . , , . i ;«nripin.
1 nnl f r : , have shown polyploidy during their evolutionary proce:
n 994). It is highly probable that the polyploidy of Muhlenbergia rigida is of autopolyploidy ir
very simuir^abk 2) chromosomes. General morphology of the studied plan^ —
Thcj=sultsof.hiss.udyrevealade.ail£ddescrip,i„n„t,hechromosomalMUsof.woMukk«i^^
da morphotypes, allowing the classification of the karyotype of M. rigida as symmetrical. The chromosoa®
bilsTwO ^‘'‘^“fding to the classification of Steb-
Stehto^ilqyT c' polyploids when compared to their disploid ancest*
Stehbins(1971)recognuesfivetnaturitystagesof polyploidy complexes: iniL, young, mature, declining.*-!
388
rehctual. According to him, the evidence of distribution patterns indicates that the majority of polyploid)’
comp exes that are currently mature originated in the Pliocene or Pleistocene. Our finding of 100% polyploid)
frequency mjhe^uhlenbe^iari^^ indicates a high evolutionary maturity.
Stebbins’ terminology (1971). All populations have predominamfy Mentrk cLonl^some” !"nd, to alesser
degree subrnetacentric chromosomes. The A1 index fiuctuated 0.1; for example, 50% of the populations had a
hi ^ I ^ ° the predominance of metacentric chromosomes,
asvmmer^ ngida karyotype constitutes the first karyotypic description of the genus. Karyotype
populations i h f ^ asymmetry is type A1 and Bl. This degree of asymmetry between
populations is high, reflecung low specialization (Stebbins 1971)
indicatingasimilarchroino-
The aggl„me„ttve clusuring analysis (UPGMA) revealed tha. the parameters which had greac« ih*'
Rosales et al., Chromosome studies in Muhlenbergia 389
ence in group and subgroup formation were the centromere index (Cl) and inter-chromosome asymmetry
(A2). The Cl values had an interval of 12.94 to 21.76 while the inter-chromosome asymmetry index values (A2)
range from 0.10 to 0.21, marking clear difference between morphotypes. Populations with the open panicle
morphotype had the highest A2 values (0.15 to 0.21) while the closed panicle morphotype populations had
lowerA2values (0.10 to 0.16).
The amount of metacentric chromosomes in the cytotypes studied, suggest that the karyotype of this spe-
des shows a tendency to be symmetrical, indicating a trend to become stable.
The ploidy levels of Muhlenbergia rigida are related to morphotypes. Plants that had compact indores-
cence were tetraploid and disploid (2n = 40 and 44) while those with loose inflorescence were triploid (2n = 30).
Therefore, cytological data provides a good complement to taxonomic studies. Knowledge on simple cytoge-
netic characteristics of a species such as chromosome number, behavior of the chromosomes during meiosis,
the mode of reproduction of individuals and their fertility can contribute to a better understanding of the pat-
terns of morphological variation and help to define taxonomic limits. The results obtained in this study con-
firm the differences between two morphotypes of M. rigida, which maintain their morphological and cytologi-
cal features even when growing at the same location, and may represent two taxonomic entities. However,
further evidence may be required to support their recognition at the species level.
The lack of cytogenetic information on species of Muhlenbergia makes the comparative study of karyo-
types and their quantitative characteristics difficult, limiting deeper discussion on the possible participation of
chromosome changes in the evolution of the genus, in its speciation processes, and the establishment of some
type of genome specialization in relation to the habitat. It is expected that a thorough cytogenetic study (with
banding, FISH, or GISH) could provide more elements to determine the evolutionary history of M. rig, da
morphotypes.
ACKNOWLEDGMENTS
We want to thank Instituto Politecnico Nacional (CIIDIR-DGO, SIP, and COFAA) for financial support, and the
«^e„.o.hero.,owi„gpr„iec.s:S,P—
ae) de Mexico,” SIP-20031025 “Estudio de los pastos mas comunes de Mexico,
:s de Mexico,” SIP-20110017 “Citologia de
c de Mexico,” and SIP-201 lObSlTitodiversidad y
y Ciperaceas ae imporianc.a ^ scholarship to the graduate
Ecosistemas de la Sierra Madre Occidental.” Many thanks to CONACyT by t „ revis-
student (CVU; 242097), first author of the present report. Finally we want to thank tew y
iched this report.
V.11L vv- V u. / j, nrst autnor oi i
ers and proof readers, whose observatu
ii.Bull.Appl.Bot.S
references
Avtxxov, N.P. 1931 . Karyo-systematische Untersuchungen der Familie Graminee
Sian with German summary. Citado en Soderstrom (1 967).
teu, AA. 1986. Noteworthy grasses from Mexico XII. Phytologia MMS ^ ^
'1«».W.D.,»SA Renvoize. 1986. G^neragramlnumigrassesof thetlWd few ^ ^ Lbpey.Ferrarl. eds. Las
drear;::;::™
tonoma Metropolitana-lztapalapa, y CONABIO,
0 en las Liliopsida Mediterranean species of Ophrys sect.
^5pex) Serna, A. 2u i z. ti enaemismo en las unuM^iuo ■ ■ Mo/4itPrrane
=««-B.».aso, M., S. 8E«a«,tx,s, .80 F. Amxh. 2010. Chromosomal
P^eudophrys (Orchidaceae): an analysis of karyotypes and ^ . got. 44:1683-1696.
F-W. 1966. Chromosome numbers of some Mexican grasses. ^ Durango. Phytologia 63:457-460.
KERiSRAARDin.. V nrso-, .. /a cspecie de Muh/enberg/o (Gramineae) de
some numbers report. Complex {Poaceae: Chloridoideae). Brit-
,n of the Muhlenbergia montana (Nutt.) Hitchc. P
'ia 50:23-50.
390
Journal of the Botanical Research Institute of Texas 7(1)
Terrera ARRtETA, Y. AND P.M. Peterson. 1 992. Muhlenbergia cualensis and M. michisensis (Poaceae: Eragrostoideae): two re
species from Mexico. Novon 2:1 14-1 1 8.
lERRERA Arrieta, Y. AND P.M. Peterson. 2007. Muhlcnbergia (Poaceae) de Chihuahua, Mexico. Sida, Bot. Misc. 29:1-109.
Masterson J. 1994. Stomatal size in fossil plants: evidence for polyploidy in majority of angiosperms. Science 264'
421-423.
Paszko, B. 2006. A critical review and a new proposal of karyotype asymmetry indices. PI. Syst. Evol. 258:39-48.
Peterson, P.M. and C.R. Annable. 1991. Systematics of the annual species of Muhlenbergia (Poaceae-Eragrostideae). Sysl
Bot. Monog. 31:1-109.
Peterson, P.M.1988. Chromosome numbers in the annual Muhlenbergia (Poaceae). Madrono 35:320-324.
Peterson, P.M. 2000. Systematics of the Muhlenbergiinae (Chloridoideae: Eragrostideae). In: S.W.L. Jacobs and J. Everett
eds. Grasses: systematics and evolution. CSIRO, Melbourne. Pp 195-212.
Peterson, P.M. 2003. Muhlenbergia. In: M.E. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds. Magnoliophyta: Ccm-
mehnidae, (in part): Poaceae, part 2. Flora of North America north of Mexico, vol. 25. Oxford Univ. Press, New York
and Oxford. Pp. 1-109.
Peterson, P.M. and Y. Herrera Arrieta. 2001 . A leaf blade anatomical survey of Muhlenbergia (Poaceae; Muhlenbergiinae).
Sida 19:469-506.
Peterson, P.M. and JJ. OrtIz-Diaz. 1998. Allelic variation in the amphitropical disjunct Muhlenbergia torreyi (Poaceae:
Muhlenbergiinae). Brittonia 50:381-391.
Peterson, P.M., RJ. Soreng, G. Dawdse, T.S. Filgueiras, F.O. Zuloaga, and EJ. Judziewicz. 2001 . Catalogue of New World grasses
(Poaceae): II. Subfamily Chloridoideae. Contr. U5. Natl. Herb. 41 :1-255.
Leon, Mexico. Sida 1 8:685-691 . ^ ^ ^ oaceae. Chloridoideae) a new species fr
^ ^ estimating karyotype asymmetry. Taxon 35:526-530.
SODERSTROM T.R. 1%7. Taxonomic study of subgenus Podosemus and section Epicampes of Muhlenbergia ((
Contr. U.S. Natl. Herb. 34:75-189.
Soms, P.S.. D.E. S^s, and M.W. Chase. 1 999. Angiosperm phylogeny inferred from multiple genes as a tool fc
tive biology. Natura 402:402-404.
Stebbins, G.L. 1938. Cytological
Bot. 25:1 89-1 98.
Ferent growth habits in the Dicotyledons. Ameci
Stebbins, G.L. 1971. Chromosomal evolution in
Watson, L. and MJ. DAawrrz. 1992. The grass ge
her plants. Addison Wesley, N
i of the World. CAB Internatio
il, Wallingford, Inglater
A SURVEY OF SEED COAT MORPHOLOGY IN OXYTROPIS, SECTS.
ARCTOBIA, BAICALIA, GLAEOCEPHALA, MESOGAEA,
AND OROBIA (FABACEAE) FROM ALASKA
Zachary J. Meyers
University of Alaska Museum of the North
907 Yukon Drive
Fairbanks, Alaska 99775, U.S.A.
zjmeyers@alaska.edu
Stefanie M. Ickert-Bond^
ABSTRACT
RESUMEN
INTRODUCTION .
Tfegenus Qvy,r„pi5 DC. (Fabaceae. subfamily PapUonoideae^&^ae^^rt.)^^^
P ex, comprising over 300 species worldwide (Yakovlev et a . , Northern Hemisphere, with its great-
^OOUewisetal. 2005). The genusis widely distributed
«t diversity found in the Tibetan plateau and I999. given & Murray 201 1). Qxytropis
/speaesarefoundintheArctic,22ofwhichoccurinAl ^^^^^^^^equently shares many morpho-
is thought to have derived from Astragalus L. approximate y ajsymmetrical leaflets, and an acaulescent
•s thought to have derived from Astragalus L. approximately - ^^etrical leaflets, and an acaulesc.
characteristics (Wojciechowski et al. 2005). A beake of the taxonomic relatu
‘^bitdistinguishQxytropisfromAstragalusmorphologica ^ (reviewed in We
V Within Qxytropis remain problematic, likely due to relatively recent
2001). ^
^®«-»«.hBLTe»s7(1):391-4
Several times in the Quaternary during glacial maxima an ice-free, continuous land bridge extended from
the Russian Far East to Alaska and Canada and played a significant role in the history of many northern taxa
(Hulten 1937, 1958; Elias et al. 1996; Ickert-Bond et al. 2009). In fact, a large ice-free refugium existed from tk
northwestern-most Arctic in Canada to the Lena River in arctic Russia, the area defined and named Beringiaby
Hulten (1937). Oxytropis present in Beringia, range in distribution from amphi-Beringian taxa to groups en-
demic to eastern (Alaskan) or western (Russian) Beringia to circumpolar elements, some with extensions
southward into the Rocky Mountains (Yurtsev 1997, 1999; Talbot et al. 1999; Ickert-Bond et al. 2009). Tk
suspected recent diversification of the genus coupled with specialized niches (i.e., O. kobukensis S.L. Welsh, 0.
kokrinensis A.E. Pors., and O. tananensis Jurtz.) has resulted in high levels of intraspecific morphological varia-
tion and phenotypic plasticity. In addition, the lack of definitive taxonomic characters in some complexes,
plethora of synonyms (reviewed in Welsh 2001).
Alexander Bunge’s worldwide treatment of Oxytropis (1874) established four subgenera based on the pres-
ence and variation of a septum in the legume fruit and the length of the calyx teeth: 1. Oxytropis (pod exerted
above the calyx, with pod always ventrally and sometimes also dorsally septate), II. Phacoxytropis (pod exerted
above torn calyx, without septum). III. Physoxytropis (small, somewhat inflated legume enclosed by intact a-
lyx), and IV. Ptiloxytropis (small pods enclosed by calyx with long villous calyx teeth). Bunge (1874) further-
more distinguished 17 sections within Oxytropis based on leaflet arrangement, raceme size and shape, legume
shape and anatomy, presence of glandular hairs, and presence of spines on the petiole. Many of Bunge’s infra-
genenc categories are still used in taxonomic works today (Barneby 1952; Yurtsev 1997, 1999; Zhu & Ohashi
2000; Polozhij & Malyschev 2006; Malyshev 2008a, 2008b; Ranjbar et al. 2009).
Revisionary work on members of Oxytropis was completed by Vasilchenko (1948), who added two sub-
genera exclusively of Old World taxa. In 1952 Barneby revised Oxytropis for North America, clarifying much of
the nomenclatural issues that had arisen. Yurtsev (1997, 1999) treated the genus from a Panarctic perspective
and noted the importance of substrate affinity to the distribution of certain taxa. Most recently, Welsh (2001)
revised Oxytropis m North America, largely agreeing with the species concepts proposed by Barneby. These
authors placed importance on characters such as stipuleshape, pubescence type, legume shape,andsubstmte
affanity to delineate taxa. Although the Panarctic Flora Checklist provides a unified view for all areas of tk
Arctic (http://nhm2.uio.no/paf), the high phenotypic plasticity observed in some taxa along with differencesin
species concepts between Russian and American taxonomists result in no widely accepted taxonomy for tk
The varied cWtes, lithologies, and landforms (many created and modified by cold climate geomorphic
processes) in Alaska generate a number of different habitats and ecological niches. Steep gradients of abiotic
factors such as disturbance, elevation, and moisture result in sharply contrasting adjacent habitats. Thus weak
geographic and ecological reproductive isolation among rapidly diversifying taxa might well have allowedin-
raspecific and interspecific hybridization in Oxytropis Oorgensen et al. 2003; Artyukova & Kozyrenko 2012)
These phenomena often caused by the influx of multiple rapid colonization e;ents have occurred in a number
including, Artemisia, Cassiope, and Rubus (Eidesen et al. 2007; Ehrich et al. 2008; Tkach etal.
eptedunlfiedtaxonomyforthefienus.loro.n..n...w.nn,^...__.,. ^ ^ „,arkers for populations
mnestrisando Jorgensen etal. (2003) using ITS and RAPD r
mpesms and O. orctica complexes in Alaska suggested that the taxa examined represent a recent racua
oeenetic study based on ITS se
..d r , 7 P ""'"^^^*^^Wstedthatthetaxaexaminedrepresentarecentraaid
non and are the result of multiple origins of polyploidy. Most recently a phylogenetic Ldy based on iTSse
20121. However, the 2012 study substantiates some infraa^n^d. namely th*
identification of section Mesogoea as the ferbZb ' relationship
tioneventsofOxytropisintheArct"^^^^^^^^
Within Fabaceaeseveralstudies have investigated the value of s^^^
namely tk
; multiple recent radia-
■omorphological c
as seen with SEM for identifying economically important taxa (Lersten 1981; Pandey &Jha 1988). Forty-eight
species of Turkish Astragalus were examined (Vural et al 2008) investigating seed micromorphology, the study
revealed two distinct seed coat types (i.e., rugulate and rugulate-reticulate, terms consistent with those used in
our study) and three distinct seed shapes (i.e., peroblate, suboblate, and oblate). Seed shapes were taxonomi-
cally useful only when combined with seed coat patterns in Turkish and Iranian Astragalus (Vural et al. 2008;
Ranjbar 2009). Seed coat patterns in Oxytropis have been documented as rugulate, lophate, or multi-reticulate.
However, only a few species have been examined (i.e., Oxytropis riparia Litv., Oxytropis campestris Hook., Oxy-
tropis lambertii Pursh), and none from Alaska (Lersten 1981; Pandey & Jha 1988; Solum & Lockerman 1991).
Recently Erkul and Aytac (2010) examined 13 Turkish species of Oxytropis and classified seed coats as striate-
reticulate, psilate-reticulate, reticulate, and striate-rugulate.
In the current paper we exended the SEM survey of seed coat micromorphology, shape and size into the
Alaskan oyxtropes. In order to test the value of seed coat micromorphology we examined representatives of
sections Arctobia, Baicalia, Glaeocephala, Mesogaea, and Orobia. Multmommal logistic regres
employed for several independent variables (tax(
tribution) to describe seed coat variability.
classification, seed din
and geographical dis
MATERIALS AND METHODS
Our survey of seed coal micromorphology of Oyxrmpis included 22 Alaskan uxa of the 67 arctic species that
have been described (Yurtsev 1999; Elven & Murray 2011; Table 1). Most of the remaining arctic species are
kom the Old-World (e.g., the Russian Far East, Siheria, and ScandinavU). Within the 22 Alaskan ^tropis we
mcluded nine species from sect. Arctobia (of 13 classified in this section), two from sect. Butcttlnt (of eight clas-
siSed in this section) three from sect. Gteocephalu (of 13 classihed in this section), one of sect Me^aea (of
:d in this section), and seven from sect Oruhir. (of 27 classified in 'h'*
:t. Arctobia (O. arctobia Bunge, 0. bryopWlu (Greene) Jurtz. subsp. lonchoporta
<7*«icaJurtz.,and0.merte,.sianaTnrcz.),weincludedadditionalsamplesthatwerecolfatedlmtn^
tagtegiorur on the Chukotka or Kamchatka peninsulas
*.nha:d::“::.:rarrbUd:Wana.om%asexam^^^
Wkion. For Scanning Electron Microscopy (SEM) whole “'^j^rTodel, and viewed with an
^ douhle.sided tape, sputter-coated with a palladium ta^e. (^
«-®-50 SEM at 1 5-20 kV at the Advanced Insttumen at the hilum
^hs. Following Lersten (1981), seed coat micromorphology was ex
at magnifications ranging from 500-2000x. micrometer on a stereoscopic
Measurements of seed length (L) and width (W) were made wuh^ ^^en are the mean
microscope as well as from the SEM image file with a scale bar ^ in millimeters
ength (measured transversely to the hilum) and width specimen three seeds were mea-
able 1). A minimum of three specimens per taxon were e Barthlott (1990).
mred. Terminology used for seed shapes and seed coats o o j ^at types (i.e., rugulate, lophate, foveo-
Statistical aualyscs.-We compared the relative fmq«n^
M With respect to taxonomic sections, species, seed leng , i„„i,.ncal regression was performed in R
^xonomic, physiological, and geographical & Ripley 2002) with the objective
'version 2.10.0 (R Development Core Team 2009) using the library ( species, latitude,
^ testing potential predictive variables of seed cMi as independent variables (IVs) in the
gitude, seed length, and seed length-to-width ratio ( information theoretical approach
with seed coat type as the dependent variable (DV). We used
!
§
il
!
Ill
III
laiiiisiiisiiiiii iiisiiiiilii
?3553?as2§?s5S3s? SSSsSsSSsSsS
S2!§s!22S22SSS??S 25?322?S2222
I
§
ll
ill III iplli 11 si i iiii Nil
ill ill Hills is ii 1 issi
.55 i55 ligiiJ55^ II 5 55J 5iis
III 3s! :§5§!? S3 S SS!S 3S53
l!l 333 :S! i23532 s3 5= ! §5 = ? —
397
to select the most parsimonious model ofall candidates by minimizingAkaike-sinformalioncrite^^^
detenttine which dependent variables explain variation of seed coat type aable 2. Akaike 1974). AlC^
implemettted as a relative measure for goodness of fit, which penalizes the number of parametetstn the mod.
to prevent overfitting (Akaike 1974).
Sted coal types and shapes.— The morphological chatacteristtes of the seeds ate J**' '' j)''
smlsexaminedappearsmoothnnderthedUsectingscopeandshowauniformdarktoltghtbrownp^menta-
Sufflgl). Typically the seedsarereniform,lmt when the lobes are utwqualmtogthsth^^^^^^^^
u'Lurl^fmilfFig.lC-ro.Fev.rseedsareovoidorglohose.dete^^^^
mitilomt settsu Murley (1951) (Fig. IC-D). Fewer seeds are ovoid or glotmse as uettueu o,
Seed anatomy— The the seed coat in all spec
at (Fig. 3
opidmnhUeoveredbyacnticleonLi^^r—
scleteids (Figs. IB-C), followed by a single row of osteoscleteids (hy^mus) an
pressed, ungentially elongate parenchyma cells of the foveoUte) were observed
Mietotitorphology.-Three primary seed coat types (i.e £ ^ P patterns among differ-
«hmanyspecies(i.e.,Qxytropisarcticavar.lmrnebyana)exhtbt.,^m*p^^
^htdivhluals (Fig. 5). The majority of seeds «■
charactenzed with irregularly roughened cuticular fold ( g ^ of interlacing ridges
s^coatswerecognizedamuhi-retic„lateform(MR),wh,ch^-._oU^^J^
and irregularly roughened cuticular folds and were found hate seed coat type, which consists of
Tablel).Approxiinately32%oftheseedsexaniineddemoi«trate t e op
tsUnct short ridges with irregular sides forming uplifted circuhro
2G'l, Table 1). We observed additional reticulation of the lopha pattern are rela-
tiwseaslophate-reticulate (LR, Fig. 2G, Table Xhe foveolate seed coat type character-
^ely thin when compared to the ridges in multi-reticu ate see ■ snrizee were only
^ by irregular circular folds with blunt edges and -hrek ceU ra^^ jp_f. Table 1)
"bse^ on 5% of seeds examined (F- |.™„tonis,andO. vtscida).
was found in only four species (i.e., O. tananensis, 0. jordahi, • y ^^^hc regression for the three major
Statistical Analysis.-Stepwise model selection of the “°®^^X:lassification. length of seed (size),
classes of seed coat types showed the strongest association descriptive model with an AIC score
a^L;W ratio (shape). These factors produced the classification, since the removal of
228.42 (Table 2). Seed coat type was associated with taxon ^he overall variation observed.
;hisfaaorincreasedtheAICscore(AAIC = .10.23),indicatmg^-^^^^^^^
2^*0 of the five sections differed in seed morphology; secuon M sog
398
Journal of the Botanical Research Institute of Texas 7(1)
il other sections examined (Table 2, Figi
•unt of variation in seed coat type (Table 2). Similarly, length
nformative of the independent variables (Tables 1, 2).
DISCUSSION
of*™ revealed ^veral distmct seed shapes, a uniform exotestal anatomy, and three main type
wMeTnfr^T"® of Alaskan oxy tropes. Seeds are typically reniform”
, w e in requent y ovoid and globose seeds were found. Anatomicallv all seeds examined have the
W T “ ““ “ ‘■istinctive combination of sclereid types: an epkknnal
lay^ol macrosclerreuls covenng an internal layer of osteosdereids. Based o
nized three seed coat types— ruguL ’ ' oasea o
some of these (ie., raulti-reticulate, a
Meyers etal. Seed coat mi(
Qxvfrn^i , ^ manifestation of the underlying epidermal cells (Fig.
^^opis may be entirely of cuticular origin and not a manitesta
*^iousstudiesinvestigatingAstragaIusandOig^tr^^d^^^™“^j“^^^
^^^0fUnriteduse.Astudythatexamined48speciesofTuricishA5tragd^ X
microphological types (rugulate and rugulate-reticulate) and three seed shapes (oblate, suboblate, and perob-
ate) (Vural et al. 2008). The authors concluded that seed coat patterns at both the species and infrageneric
evels were olhmited taxonomic utility in Astragalus unless supported by macromorphological characters.
Similarly, 13 Turkish species ofOxytropis were examined and classified as striate-reticulate, psilate-reticulate,
reticulate, and stnate-nigulate (Erkul and Ayta? 2010). Comparison of morphology of these Turkish species
with the current study is limited however as these seed coats types (striate-reticulate, psilate-reticulate, reticu-
ate. and stnate-rugulate) are not well defined in the study (Erkul and Ay tag 2010). The pioneering work of
testal topography examination by Lersten in 1981 revealed tribal associations in Leguminosae subfamily ?=>-
pihonoideae, specifically with Vicieae, Trifolieae, and Cicerae.
The multinomial logistic regression revealed species delimitation (independent variable) to describe tht
a y, species e mutation in Oxytropis has been controversial, resulting in competing taxonomies
agray sale (black = rugulate; gray = lophate; and white =foveolate).
■miaplethora of synonyn.s (reviewed in welsh 2001) andanumbcrofAlaskanOxy.i^uxaare^^^^^^^^
^IbleCseeB^rnelylUvurtsev 1997; WelshlOOhJorgenseneial. 2001^
fKognized, would be taxa of conservation concern (AKNHP 2013). We n has been de-
M.ba.sup,x.rtseparationof0.1u„unensisfro.avuri™.^^^
^ d as a distinct taxon by Yurtsev in 1993 base on synonymized
Sven and Murray (2011) S“-^‘ ^ -
‘jven and Murray (2011) sees It as part ot me naiuidiivaaa»vavr*.
0.tananensiswithhisO.campes£risvar.vanans.Sinularly,there^^
whi.e*we.d'pi;:;;fa;';;;catan^^^^^^^^^
Itself indicate a lack of taxonomic validity of these taxa. bserved some trends in frequency of seed
Despite the lack of taxonomic utility at the species leve we o rv sectional level for
coat types at higher infrageneric ranks. Seed coat type and Arctobia with 97% of the
^ArctoHa and Mcsogueu. Seed CMt “^'^rTed in the Alaskan members of sections
^ex^rned being strictly rugulate. Greater vanabr y Iniraspecihc vanabrl-
Dtoha, Baicalia, and Glaeocephala. Taxa examined from
"yin^dcoat types. m Alaskan Otylmpis. even within individual spe-
Despite the variability of seed shapes within sections , by having exclusively small globose
ctes, Qxytropis dejlexa of subgenus Phacoxytropis sect. Mesogaea is i ^ ^ compared to seeds
^cds, which range from 1 to 1.5 mm in length and 1.2 to 1.6 reniform (L = 0.92 to 2.45 mm, W
^*^®Mied from other subgenera and sections, which are ^
= to 2.72 mm, Table 1; Fig 1). Subgenus Phacoxytropis o imge
as Yurtsev (1997) as a distinct lineage within Oxytropis »
402
0. koyukensis (AF366334)
O. koyukensis (AF366332)
0. koyukensis (AF366330)
0. arctica var. barnebyana (AF36631 2)
0. arctico var. barnebyana (AF36631 0)
0. arctica var. barnebyana (AF 366^8)
Q maydelliana (HQ 1 76486) ~
a maydeiliana (HQ1 76485)
0. maydelliana (HQI 76484)
characters. Molecular evidence strongly support section Mesogaea as being an ancestral lineage of Oxytropi
(Archambaul,&Stn,mvik 2012). In addi, ion, matKsecfuence data for 6371egumetaxaincluding5evens,«i»
of Oxjrirop.s, o. deflexa was shown to be the earliest diverging branch (unpublished data, Martin F-
WojcrechowsM, Arizona State University, Tempe, Arizona), However, additional samples of Asiatic men*"
u. snbgenns Phncor^tntpis are needed to test whether small globose seeds represent the plesiomorphic seed
shapeandthatr^iforrnseedsrepresentthederivedCapontorphiclconditioninQ^^
nanemr 'ft ” “"T' observations on morphological diversification as they tev^
patterns of evolution tn the genus Qrytmpis, particularly since arctic members have been reported to be *
result of recent rapid radiaUons and hybridization involving polyploidy Oorgensen et al 2003, Archamba*
and Stromvrk 2012; Artyukova & Kozyrenko 2012). It is possible that the lack of consistent seed morphoW
wrthmspectes IS due to past introgression and more current hybridization Similar, ■ ■
have occurred in a number of other arctic genera including, Artemisia, Cassiope, and Ruhis (Eidesen et al. 2007
Ehrich et al. 2008; Tkach et al. 2008)
iiuuies investigating the importance of ecology in the speciation process might be particularly promising
lOxytropis, where sympatric species are documented to inhabit different ecological niches. For example, O.
nifcftfirfl is an acidonhilous snecies while O. <
ing that populations adapt to similar environments via different genetic pathways (Steiner et al. 2009) further
confounding the issue. Taxonomic resolution in a complex genus such as Qxytropis will only be achieved using
amulti-level approach that integrates phylogenetic, morphological, and ecological investigations.
ACKNOWLEDGMENTS
CAN, DAO, GB, GH, LD, LE, MO, NDG, NY, O, PH, S, UPS, and US (Thiers 2008). We would also like to thank
Ken Severin of the Advanced Instrumentation Laboratory (AIL), University of Alaska Fairbanks for access to
the scanning electron microscope, Dave Murray for his continued support and critical review of an earlier ver-
sion of this manuscript, Israel Loera for the Spanish translation of the abstract, Matt Carlson and an anony-
mous reviewer for their helpful comments. This study was funded in part by an Undergraduate Research Fel-
lowship to Rose LaMesjerant from Alaska EPSCoR, NSF award EPS-0701898, and the state of Alaska.
;w look at the si
references
._al model identification. lEEETrans. Automatic Control 6:71 6-723.
i). 201 3. Rare plants of Alaska tracking list. University of Alaska Anchorage, http://
y/co.any_tracking_page.htm. Accessed:! 6 Mar. 2013.
M.V. Stromvik. 2012. Evolutionary relationships in Oxytropis species, as estima e
ribosomal internal transcribed spacer (ITS) sequences point to multiple expansions in o e
770-779
EV. and M.M. Kozyrenko. 2012. Phylogenetic n
PhyWa (Pall.) DC. (Fabaceae) inferred from the data o.
operon and intergenic spacers of the chloroplast genome. ^^ad. Sci. 4:177-312.
rIT' revision of the North in plants. In D. Claugher, ed. Scanning elec-
BARTHunr, W. 1 990. Scanning electron microscopy of the eoidermai suna
C Re^ated,^ out of Baring,. «trogo,» embracas tha
Arctic J. Biogeogr. 34:1 559-1 574. , . g^jdag. Nature 382:60-63.
S-A. S.K. Short, C.H. Nelson, and H.H. Birks. 1 996. Life and times o e g ^ Aora (PAR. Vascular plants.
tLVw, R. and DM. Murray 201 1 . In R. Elven, ed. Panarctic 0^3
J|^^/nhm.uio.no/english/research/infrastructure/paf. (Fabaceae) in Turkey. XIII Optima Meet-
'"9- Anatalya, Turkey, March 2010. Abstract, P.1 64.
E. 1937. Flora of tb^ ^ .nd westernmost Alaska F
C^95B.Thaa,,,pb,.a.,an,icp,an,tandd,al-pM<^ecg«pNtaicon^^^^
"'■“ow, S.M., O.F. Mu«ba», Al® E. DiCham. 2009. Contrasting e^^aska Park Sd. 8:26-32
id 0. arctica (Fabaceae) comp
Lersten, N.R. 1981. Testa topography in Leguminoseae subfamily Papilionoideae. Proc. Iowa Acad. Sci. 88:180-191.
Lewis, G.P., B.D. Schrire, B.A. Mackinder, and M. Lock. 2005. Legumes of the World. Royal Botanic Garden, Kew.
Malyshev, LI. 20O8a. Diversity of the genus Oxytrop/s in Asian Russia. Turczaninowia 1 1:5-141.
Malyshev, LI. 20O8b. Phenetics of the subgenera and sections in the genus Oxytropis DC. (Fabaceae) bearing on ecology
and phylogeny. Sibirskii Ekolog. Z. 15:571-576.
Murley, M.R. 1951. Seeds of the Cruciferae of northeastern North America. Amer. Midi. Nat. 46:1 -81 .
Pandey, A.K. and S.S. Jha. 1 988. SEM studies on spermoderm of some Galegeae (Fabaceae). Curr. Sci. 57:1 008-9.
PoLOZHu, A.V. and LI. Malyschev. 2006. Flora of Siberia. Vol. 9: Fabaceae (Leguminosae). Science Publishers, Enfield, NH.
R Development Core Team. 2009. R: A Language and Environment for Statistical Computing. R Foundation for Statistical
Computing, Vienna, Austria. http://www.R-project.org
Ranjbar, M. 2009. Astragalus sect. Trachycercis (Fabaceae) in Iran. Nord. J. Bot. 27:328-335.
Ranjbar, M., R. Karamian, and S. Bayat. 2009. Notes on Oxytropis sect. Mesogaea (Fabaceae) in Ir
a new species. Ann. Bot. Fenn. 46:235-238.
Solum, D. and R.H. Lockerman. 1991 . Seed coat surface patterns and structures of Oxytropis ri
Medicago sativa, and Astragalus deer. Scan. Microscop. 5:779-786.
Steiner, C.C, H. Rompler, L. Boettger,!. Schoeneberg, and H.E. Hoekstra. 2009. The genetic basis of phenotypic convergence in
beach mice: similar pigment pattern but different genes. Molec. Biol. Evol. 26:35-45.
Talbot, S.S., B.A. Yurtsev, D.F. Murray, G.W. Argus, C. Bay, and A. Elvebakk. 1999. Atlas of rare and endemic vascular plants
of the Arctic. Conservation of the Arctic Flora and Fauna (CAFF) Technical Report 3, U.S. Fish and Wildlife Service,
Anchorage, Alaska.
Tkach, N.V., M.H. Hoffmann, M. Roser, and K.B. von Hagen. 2008. Temporal patterns of evolution in the Arctic explored in
Artemisia L (Astreraceae) lineages of different age. PI. Ecol. Diversity 2:167-169.
Thiers, B. 2008. Index Herbariorum. A global directory of public herbaria and associated staff. New York Botanical Gar-
den's Virtual Herbarium. httpy/sweetgum.nybg.org/ih/
Vasil'chenko, I.T. 1948. Oxytropis. In V.L. Komarov, ed. Flora of the U.S.S.R., vol. 13. Akademii Nauk SSSR, Moskow-Lenin-
grad. Pp. 1-229.
Venables, W.N. and B.D. Ripley. 2002. The Modern Applied Statistics with S. A**’ Edition. Springer Verlag, NewYork.
Vural, C, M. Ekiq, H. Akan, and Z. Ayta. 2008. Seed morphology and its systematic implications for the genus Astragalusl
sections Onobrychoidei DC., Uliginosi Gray and Ornithopodium Bunge (Fabaceae). PI. Syst. Evol. 274:255-263.
Welsh, S.L 2001 . Revision of North American species of Oxytropis de Candolle (Leguminosae). EPS Inc., Orem.
WoxiECHowsKi, M.F. 2005. Astragalus (Fabaceae): A molecular phylogenetic perspective. Brittonia 57:382-396.
Yakovlev, G.P., A. Sytin, and Y.R. Roskov. 1 996. Legumes of Northern Eurasia: A check-list. Royal Botanic Gardens, Kew.
Yurtsev, B.A. 1 993. Oxytropis tartanerisis, a new species of Section Baicalia (Fabaceae) from the interior of eastern Alaska.
Bot.Zhurn.78(9):59-65.
Yurtsev, B.A. 1 997. Analysis of evolutionary differentiation in key arctic-alpine taxa: Dryas, Oxytropis sect. Arctobia and
Taraxacum sect. Arctica. Opera Bot. 132:27-37.
B.A. 1999. Survey of arctic legumes with an emphasis on the species concept in Oxytropis. In I. Nordal andV.
in the High North - A Panarctic flora initiative. The Norwegian Academy of Sci-
ence and Letters, Oslo. Pp. 295-318.
Zhu, X. and H. Ohashi. 2000. Systematics of Chi
e Oxytropis DC. (Leguminosae). Cathaya 1 1 -1 2:1-218.
THE FORGOTTEN FIRST CHECKLIST OF THE IOWA FLORA:
JOHN HENRY RAUCH’S 1851 “REPORT” TO THE
STATE MEDICAL AND CHIRURGICAL SOCIETY
Thomas G. Lammers
Department of Biology
University of Wisconsin Oshkosh
Oshkosh, Wisconsin 54901, U.SA.
lammers@uwosh.edu
H primer gran compendiode la flora de Iowa no. . j jx j
deBuilington por mandato de la Sociedad Medica y Quinirgica del Estado de Iowa, fue publico dos ddcadas ai
la flora del estado por Charles Bessey. Rauch listo 516 especies er *
T^^ar«e,f„„.owardanoraof,owa-recogn.edbyH«^
>WS been viewed as such by the state’s botanists since Arthur ( “earliest significant re-
to the Flora of Iowa’ thus far made.” Prior mentions of plants from Iowa wer , scattered
cord” according to Eilers was Parry (1852). However, the Iowa reports in a^ of thf 727 snecies listed
^.ughoutawLdevotedprimaZ.MinnesotaandW^^^^^^
only 204 (most collected at Davenport m 1847) were explicitly Guldner 1960) and by a
we.co.,ec„dinthes.a.e,asshownbyF.rry-s„otebo„km.hePu.na™^^
bnef excerpt published in a local newspaper (Parry 1847) Areaderof that time could
(e|CthroughouttheNorthwest”or^banksoftheM„
®ly wonder if a “common” species such as Mimulus nngens L. or Songuma
"X°o!;iyearliermentlo„ofIowaplanBwhichEile.3(197»k™w^^^^^
nzed Karl Andreas Geyer’s collections from the upper Mississippi wae^ specimens from the state
(1852), Geyer’s
IS Geyer’s collections from the upper specimens from the state
, n Iowa was peripheral to a survey (shimek 1915; Bray & Bray 1976).
•*»»U.«ed mostly al Spirit Lake. 29
_ Recently, 1 had occasion to delve into the life and ^ ^ statement in several biographical
ri900; Kelly & Burrage 1920) that Rauch, as a
;o report on “the medical and economic
— juwa, Quring the 1850s. In my r
^hes (Anonymous 1894a, 1894b, 1894c; ’Wilson i
“lember of the fledgling state medical society, had been appointed l
‘™a>yo(lowa,"areport that was completed and published in 1851.
‘■'^wtetalninos-t
This came as a complete surprise to me. In 35 years of interest in the Iowa flora, I had seen no mention of
such a publication. Surely, I thought, if it were of even the slightest botanical merit, it would have been known
to Fitzpatrick (1899), Gilly (1947), Shimek (1948), Thome (1954), or others. I could only assume that it was
merely a few pages of quaint narrative about a handful of plants, identified largely by vernacular names of im-
precise application. But when I at last located a copy (Rauch 1851a), I was astonished to find that it was an ac-
tual checklist of the Iowa flora, in many ways comparable to the first floristic list for Illinois (Mead 1846). It
reported twice as many species as Parry (1852), following the nomenclature of leading contemporary bota-
nists. In the quality and quantity of information provided (observations on habitats, phenology, distribution
within the state, etc.), it was far superior to the hare-bones listings of Bessey (1871) and Arthur (1876). And it
had been completely forgotten!
Rauch’s (1851a) report offers a unique glimpse at Iowa’s flora just 18 years after settlement began. At that
time, the frontier lay a mere 50-75 miles west of the Mississippi River, and the state still contained millions of
untilled acres (Plumbe 1839; Galland 1840; Newhall 1841, 1846; Parker 1855). By the time Bessey’s (1871)
contribution was published, the state was essentially settled and largely in cultivation (Fite 1966). As such, this
forgotten first checklist provides valuable insights on the flora of Iowa prior to settlement, and enhances our
understanding of how it changed following the advent of widespread agriculture and the closing of the frontier
(Thompson 1980; Farrar 1981). It also casts light on a time when floristic study was largely the purview of dedi-
cated amateurs rather than paid professionals (Stuckey 1978, 1984; Rudolph 1988), and on early interest in the
therapeutic value of plants, an interest that continues today (Lewis & Elvin-Lewis 1995). The purpose here is
to assure for Rauch’s report the signal position it deserves in Iowa’s botanical history, by carefuUy analyzing
and interpreting the information that it contains.
John Henry Rauch (Fig. 1) was bom on 4 September 1828 in Lebanon, Pennsylvania, the first of ten children
bora to Bernhard Rauch and the former Jane Brown (Beatty 1991). He assembled his first herbarium as a stu-
dent at Lebanon Academy (Lee 1895). After graduating in 1846, he spent a year under the tutelage of local
physician and surgeon John Washington Gloninger (1798-1874), himself a student of the eminent botanist-
physician David Hosack (Kelly & Burrage 1920). Rauch then enrolled in the medical department of the Univer-
sity of Pennsylvania. Upon completing a thesis on the medicinal value oi Pohgonatum officinale L., he gradu-
ated an M.D. on 20 March 1849.
to Burlington, Iowa, to establish his medical practice. Adver-
1 local newspapers (Fig. 2) place his offices on Jefferson St. (Burlington Daily Telegraph, 9 June 1851);
on Mam St. (Burlington Daily Hawk Eye. 2 March 1853); and on Third St. (Daily Hawk Eye & Telegraph. 9 De-
cember 1856). Not only did he practice, but he conducted medical research as well, examining the efficacy of
topical applications of chloroform in alleviating various conditions (Rauch 1851b), the influence of ozone on
™ of ^
). His adv^acy for ihe men employed m navigation on the Mississippi resulted in his appointment to the
^mtssioncharpd with selecting sites forUS,Marine Hospital Service facUities, one of which was locatediti
Burlington (Kelly & Burrage 1920; Williams 1951).
s, . physicians who me. at Burlington to establish the lew
Sme Medttul and Chirurgical Society, which in 1856 shortened its name to the Iowa Medical Society (Wats®
1894; Fairchild 1927; Uwrence 2003). Ranch played an active role in this organization serving in ttim as U-
toanan (Ransom 1851), Recording Secretary (Anonymous 1854a), First Vice President (Beatty IWD, ^
F^stden.(An™ym^s 1858a, 18m He wasalsoafoundingmember of the DCS Moines County MedicalSoa
eht,estabhshedtnl852,servingasi.slirstCorrespondingSecretary(Bea.t^ 1991).
auc s mteresis in this period were not confined to medicine. As Lee (1895) commented, "He was a niaa
of t^ enlarged vrews, and possessed of roo mnch public spirit, «, be eonient with the ordinary drudgery of
medical practice, and the mere accumulation of wealth had no charms for him. The natural sciences, on tK
MU JOBS H. BAUCH*
&«eee m A. Cltrisc*» store, on Jeffersoastreet.
*igsstI5,5«
„ (Iowa) fti//rre<«?n#for9 June 1851.
, u .whocampaignedforstatefundingofagpological
'«herhand,strongly attracted him.” Rauch wasamongth^h^P^jS^^^
^fvey (Kelly & Burrage 1920), which came to fruition m the Abernathy 1907), serving
^HKster, he wasamember of the faculty at Burlington
^ Lecturer on Natural Sciences (Daily Hawk Eye & Telegraph F University as houseguest
«largezoologicalcollection,whichhetooktoLouisAgassiz(l^^^^^
and private pupil (Anonymous 1894d, Hamilton 1894; Beatty 1991). The celebrated zoologist described several
new species of teleost fishes from this collection (Agassiz 1854, 1855), including the cyprinids Chondrostom
pullum, Hypobsis dorsalis, and Rhinichthys meleaghs; and Ichthyobus rauchii, a catostomid named in his honor.
Numerous Rauch specimens were also cited in his monograph of North American turtles (Agassiz 1857). After
Agassiz’s death, Rauch was appointed to the celebrated zoologist’s memorial committee, serving among such
s Henry Cabot Lodge and Oliver Wendell Holmes, Jr. (Lawrence 1876).
ctive in at least three other
>. The Iowa Historical and Geological Institute (see below) co
(Wilson & Fiske 1900); he served at least one term as Recording Secretary (Parker 1855) and at the group's
age (Anonymous 1858b). He delivered the keynote address (Rauch 1853) at the annual meeting of the Southern
Iowa Horticultural Society on 20 October 1852. Likely it was through this connection that he “received a great
variety of Seeds from the Commissioner of Patents for distribution” to the public (Daily Hawk Eye & Telegraph,
20 November 1856). Rauch was also a Mason, a member of Des Moines Lodge, No. 1, in Burlington, and Grand
Orator of the Iowa Lodge for 1854 (Parvin 1858). At the national level in these years, he was a member of the
Academy of Science of St. Louis (Anonymous 1856), the Chicago Academy of Natural Sciences (Sperry 1904),
the Maryland Historical Society (Anonymous 1854b), and the American Association for the Advancement of
Science (Anonymous 1857a).
A local eulogist summed up well the high regard felt for Rauch during these years by his fellow Burlingto-
nians (Anonymous 1894a):
He was a leader in the society of his day and no event of importance was deemed complete without him.
He was always foremost, too, on all public occasions of a semi-social character, as the opening of the [Chi-
cago, Burlington, and (^incy Railroad], etc. He was a great admirer of ladies and was highly esteemed in
turn, yet died a bachelor. A street in Northern addition was named Rauch in his honor, but some supersa-
pient city council saw fit to change the name to Garnet, thus defeating the purpose of perpetuating the
Military service.— In response to the federal government’s call for volunteers to oppose the secession of
southern states, Rauch enlisted in the Army Medical Service in May 1861 (Lee 1895; Sperry 1904). He served
at the First Battle of Bull Run an ’
he became assistant medical director for the Army of the Potomac, in which capacity he participated in the
Battle of Cedar Mountain, the Second Battle of Bull Run, and the Battle of Antietam Early in 1863, he was
transferred to the Army of the Gulf, serving as special medical inspector and participating in the Siege of P«
Hudson and the Second Battle of Sabine Pass. Relieved from combat-zone dnty in 1864 he was appointed medi-
cal duector at Fon Wayne in Detroit. Michigan, and then at the army post in Madison, Indiana. At the latlet
posting, he was brought before a board of inquiry by a malicious subordinate on an array of charges, but all
were fonnd to be without merit (Beatty 1991). He was brevetted lieutenant colonel effective 13 Maich 1865 and
honorably discharged on 14 July 1865 (Anonymous 1887).
Chicugoyears.-ln 1857. Rauch became Professor of Materia Medina at Rush Medical College in Chicago,
Illinois, though he maintained his residence in Burlington until autumn 1858 (Wilson & Fiske 1900; Bead)
1991)i^Early the next year, he resigned this position. That September, he became Professor of Materia Medicaal
tte Chtrago College „ Pharmacy, the first such school west of the Alleghenies and one of the predecessors of
today s Univeisity of Illinois at Chicago (Hamilton 1894), Upon his return from military service, Rauch be-
erested in the relattonship between urban land use and public health (Wilson & Fiske 1900). Hecnh-
cized contem^raty burial practices as all-too-conducive to the contamination of groundwater and spread of
disease (Rauch 1866, 1M5), and advocated the planting of trees and the devebpment of public parklandsasi
fntemected in the closure of the decrephCh)'
* Six-year term on the city’s newly created Board of Health, where he
served as sanitary superintendent in the office of sanitary inspectors. His tenure with this agency was a con-
stant battle with powerful commercial interests polluting the Chicago River. “By simply shutting his eyes at
any time during that period, he could undoubtedly have secured an independent fortune. But his sturdy ‘Penn-
sylvania Dutch’ ancestry forbade the thought of any such venal dereliction” (Lee 1895). An even greater chal-
lenge came in the aftermath of the catastrophic conflagration of October
health of 112,000 homeless citizens (Hamilton
obliged everyone there to be v
t his sturdy Pen
i responsible for the
iamilton 189"!;. wnen smaiipox appeareu iii me iciugcc taiupa,
obliged everyone there to be vaccinated, thus preventing calamity (Lee 1895). In recognition of his preeminent
experience as a sanitarian, he was elected president of the American Public Health Association in 1876 (Rauch
1878; Davenport 1957).
(Davenport 1957).
Despite his professional focus on sanitation and public health during these years, Rauch maintained t
strong interest in natural history. In the winter of 1870-71, he traveled to Venezuela to study sanitaiy condi-
tions for the Orinoco Mining and Exploring Co. (Beatty 1991). Here “he made a large and valuable collection of
natural objects for the Chicago Academy of Natural Sciences” (Sperry 1904). Unfortunately no Rauch speci-
mens of any sort are to be found in the Academy’s collections today; likely all were destroyed in the firestorm
ofOctober 1871 (Dawn Roberts, pers.comm.).Rauch’spersonalherbarium,amanuscriptentitledSynopsisof
theFloraof the North West,” and allofhis notes fromhis South Americansojourn were also lostin that histone
conflagration (Beatty 1991). . . „ u n ^ decade of ser-
in 1877, the Illinois legislature created the State Board of Health. Rauch, after mo
*e„Chtagom.his field, wasanoWous candidate for membership. He was^cted .he
<knl, and then served as secretary the remainder of his tenure (Anonymous 1894b; ,
» 15 years, he crusaded against -untrained and unscrupulous pretenders to the medtcal arts (Le^^lW5),
•ihequak and charlatans who were preying upon the public, who could no. pass an enamtnanon tn the pro^
feil Which theytmXo be quaLi^ to pmctice; who imposed uj«n the*
•rang the last dollar from the victims olhopeless disease on pretense
ytnous 1891). To do this, he rigorously and verified the genu-
10 determine his qualifications to practice (Webster y in the stale (Anonymous
compelled almost 40% of Illinois' 3600 non-graduate physraans to ^ ^ declamation through
l»^).-Wm..hee„tireAmericanMedicalAssociationfaiW.o_*ct^^^^^
tomes ol decades, he accomplished in cupelled to conform" (Lee 1895).
®edical acquirement in the west to which the colleges in werful allies in the state legislature
These efforts earned Rauch many enemies - powerful enemies hostage to demands by
JWohster 1909). inhoth 1889and 1891, theBoard-shicnmal^Pjn^^^^^^
■^gislators for his resignation. Newspapers aligned with the lenient. ... his acts
«Uing Rauch “obnoxious . . . exceedingly arbitrary in or incompetent” and accusing him
generally have been such as to create the feeling that he w integrity (Anonymous 1892a). Re-
paying bribes to prevent revelation of damning facts a ^ r-anable physician was his enemy, no com-
PuHican newspapers rushed to his defense, pointing out that o p ^ opposed him, only those
““nity where he had checked disease, or prevented the 1891). The embattled Rauch
^hose empiricism was exposed by his efforts to protect the people (Anonym
My resigned, effective 30 June 1891. , „_^„ivania “wearied, worried, despondent
In November 1893, Rauch returned to his hometown m e youngest brother, Cyrus
'“d in ill health” (Guilford 1908). Here he took up residence
^^nymous 1894e; Beatty 1891). He died there of heart ai ure o
Cemetery (Lee 1895).
PREPARATION OF THE REPORT
At its inaugural meeting in June 1850, the Iowa State Medical and Chirurgical Society passed a resolution call-
ing for the preparation of a report on plants of medicinal and other economic value that were to be found in the
state (Matthews 1850). The committee appointed to carry out this task was chaired by Dr. John W. Brookbank
of Burlington (Anonymous 1850); its other members were Rauch and Dr. John F. Dillon of Farmington (Mat-
thews 1850, Sanford 1850). Regarding this committee and its charge, Sanford (1850) commented,
The floral riches of this beautiful and charming state seem almost inexhaustible, and no one doubts that,
amidst this profusion of nature’s eloquent and poetic beauties, mines of medicinal wealth exist, from
which the balm to many an ill, incident to our country, may be bountifully drawn. Numerous as are the
blessings which spring from the bosom of our mother earth, they may be greatly multiplied by the assidu-
ity of the medical botanist, and we sincerely trust a larger proportion of our physicians may be found de-
voting their attention to this subject. Our medical plants should be known and accurately classified in
order that we may resort to the great storehouse of nature — the fields and the forests — ^where no merce-
nary hand mingles with their life-giving principles, the seeds of death.
The finished report was read by Rauch at the Society’s second annual meeting, held on 7 May 1851 in
Fairfield. He stated that preparation of the report fell to him “in the absence of the chairman of the committee."
In fact, neither Brookank nor Dillon was practicing medicine during most of the year between the committee’s
formation and the presentation of the report, and all evidence points to Rauch as sole author.
John Forrest Dillon (1831-1914) was at that time but 19 years old. He was one of five graduates that spring
from the College of Physicians and Surgeons of the Upper Mississippi in Davenport, a forerunner of the Univer-
sity of Iowa School of Medicine (’Watson 1894), and was en route to Farmington to commence practice (StUes
1909). There he discovered that a medical condition (an inguinal hernia) made it very painful for him to ride
horseback, a requisite for a physician in that rough and thinly settled country. By that autumn, he had givenup
the practice of medicine, though not his membership in the Society (Anonymous 1854a) and not before pub-
lishing on one of his cases (Dillon 1850). He returned to Davenport, took up the study of law, and was admitted
to the Scott County bar two years later. His career as a jurist was long and illustrious. He founded and for many
years^edited the Central Law Journal, was elected to the state Supreme Court (Chief Justice 1868-69), and
Little is known about John W. Brookbank (Fairchild 1927). He was apparently the first physician to prac-
tice at Wapello in adjacent Louisa County, and represented that county at the 1844 state constitutional conven-
tion (Springer 1912). Some time later, he moved to Burlington and entered a partnership with Dr. H. Houghton;
that affiliation was dissolved “by mutual consent” two months before the founding of the Society (Burlington
Hawfe Eye. 11 April 1850). The next we hear of him isayear after the meeting, whenhe announced that he “has
resumed the Practice of Medicine” (Burlington Hawk Eye, 19 June 1851). Taken together, these notices suggest
that he, like Dillon, may not have been in a position to assist the committee during the year following his
appointment.
In his introduction, Rauch acknowledged that not every medi
eluded. This lack of completeness he attributed to two factors First
ration in Iowa prior to that time, which meant that he could not rely upon
anysigmficantextent. Second, because Rauch had been in thesiam,
gan,hi " • - -
in his repmrt from three sources.
) be found in Iowa w
ng literature or collections to
when the project
As a result, he assembled the information embodied
, , , , ‘ ^ meeting ended, “I endeavored to gain as much information as 1
fclycouldhy personal observation, at the same time collectingalltheindigenousmedicind
hemdo/tfes,usonlhad,„i.eafinecollection.uwasmyp„rposetohavepmsen.edd„plim^^^
LenX'ol o^rTtam 7"
1850 gtowingseason. Localities reported were Burhngton.KeSltt.'-NOTA^riofliZ-'Tai'esH
Lammeis, John Henry Rauch's Iowa flora checklist
State,” “S.W. of state,” and the Cedar, Wapsipinicon, and Missouri Rivers, which would imply that Rauch trav-
eled the length and breadth of the state while collecting. In light of the fact that Iowa had not a single mile of
operating railroad at that time, this would have been a considerable undertaking.
The enticing prospect that herbarium specimens exist to vouch for this forgotten checklist are dashed as
soon as they raised. In the very next sentence, Rauch stated, “unfortunately my specimens were all destroyed.”
How and when this calamity befell was not indicated, but it must have happened sometime after Rauch identi-
fied the specimens, but before he presented the report.
Curiously, it would seem that this was not the only herbarium in Burlington that was destroyed m tms
period. The Iowa Historical and Geological Institute was an amateur scientific society founded at Burlington in
1843 (Newhall 1846; Jewett 1850; Rhees 1859). According to Parker (1855), its holdings included “an herbari-
um containing the greater portion of the plants found in the State.” All of the Institute’s collections were de-
stroyed on 16 January 1853, when the building housing them burned to the ground (Anonymous 1853a,
1853b, 1857b). ..
The existence in the 1850s of two herbaria inBurhngton, both purporting to representwell the flora ol the
entirestate, seems odd. If the Institute’sherbarium was in existence when Rauchwaspreparing his report, why
didhenotrelyonitforinformation?Asamember(Parkerl855;Wilson&Fiskel900),hewou^^^^^^
about it. One possibility is that there has been some confusion and the herbarium mentioned by Par er { )
was actually Rauch’s, destroyed prior to May 1851. Alternatively, Rauch may have
stated in the introduction of his report, to rebuild the lost collection at some future peri , is a res
Southern Iowa Horticultural Society (Rauch 1853) does allude to botanical collecting during t e summer o
1852. If this had been done under the aegisof the lnstitute,he once again would have lost his herbarium When
Rauch also availed himself of such pubhcations as were available, specifically “tlm
various reports of the Geological and other explorations made of this state by the a852)
an allusion to Torrey (1843, 1845), but if so, Rauch did not take up all of the secies is e .
fullreport had not^t been pubhshed, though an excerpt hadbeen printed (Pa^^^^^^^
teown to Rauch (see below). But again, he did not take up all species listed t ere^ g
(e.g..Pike 1810; Allen 1814; Lea 1836; Allen 1846) did not contain any fornaal bot^^^^^^^
accounts of Plumbe (1839) and Galland (1840) ^ [LTand Gray, Bigelow, Rafinesque,
TTte literature mentioned by Rauch also included “wor^ ^LLryofWoodand
^ddel[sic],Eaton,Nuttall,James,Wood,Griffith’s^^^^^^^^^
^che. The last two (Griffith 1847; Wood&Bache 1836) no
'«Mostof,heremainder(e.g.,Nut.alll818;Toir£yl824;Eaton&Wn^. . y
« guides to identification and nomenclature, rather than as ^^ceus and
^ To the best of my knowledge, the only plants reined fam l^n
freiMndies alba by Wood (1847), and CaUbhae irumgulala by Gray . ^
Wta not the first. However, the reports ''T ^sh (1814), Nuttall (1818), or James
We been derived from the literature. Rauch seems to ha ^ Artemisia santonicum, Pursh 1814; Gcn-
d823, 1825) reported a species from “the plains of the Missoun explorers did not en-
fiana acuta, Nuttall 1818), it could be assumed to occur in wes
counter such plants until much farther upstream. formation obtained from three other men
OAerbotanists—Rauch’s third source for his info^^^^
®terested in botany. The first, whom he acknowledged for mu 1899; Wood 1955). In 1820, he
(1797-1861). James was a student of bothjohn Torrey and Amos Mountains, where he
as botanist and geologist on Maj. Upon completing his report of the ex-
Ae first recorded ascent of Pike’s Peak (Goodman & Lawso ^ assistant surgeon in the Army’s
Potion (James 1823, 1825; Torrey 1828), he secured a" ^3„ner in publishing his captivity
«^dical department (Benson 1968), during which time he assist
narrative (James 1830). In 1833 he retired from medicine and three ;
Burlington, where he spent the remainder of his days as an active advocate ol
1899; Pammel 1907-08). In the body of his report, Rauch related informatio
Dr. James” on the use of Eupatorium perfoliatum as malarial prophylaxis, as well as reporting tf
s Alfred Sanders (1819-1865), “who has done much as an amateur
botanist toward exploring and collecting the Flora of this State.” Sanders was owner, publisher, and editor ol
The Davenport Gazette, which he had founded upon his arrival in 1841 (Gue 1903); it was this newspaper that
had published Parry’s (1847) summary of his Iowa collecting. “In early life he made botany a specialty, and was
very successful in its prosecution. During the first year of his residence in Iowa he spent much time in the
Parry’s (1847) list, Sanders
cies not listed by Rauch
The final
as credited with collecting Penstemon grandijlorus Nutt, on the Cedar River, a spe-
i ne nnai man mentioned was David Sylvester Sheldon (1809-1886), who (along with Rauch and Sanders)
would “be happy to exchange [plant] specimens with persons of this or any other State.” Sheldon was Professor
of Natural Science at Iowa College in Davenport, the forerunner of Grinnell College. Here, he spent consider-
able time “scouring the adjacent country, securing specimens of plants, shells, fossils, etc., which by an exten-
sive system of exchanges, secured valuable returns from all parts of the world” (Parry 1893). During 1852-54,
Sheldon was assisted in these efforts by a tutor at the college, Sereno ’Watson (1826-1892), who later became
curator of the Gray Herbarium at Harvard University (Anonymous 1892b). Together they assembled “a very
complete series of local plants . . . most of which are now stored in the Davenport Academy [of Natural Sdenc-
es] Herbarium” (Parry 1893). Sheldon was the first president of that institution (Pratt 1882), which was the
forerunner of the Putnam Museum.
Publicatum.— The report was approved and ordered printed as part of the proceedings of the meeting
(Ransom 1851). For each of the 516 species recognized, Rauch provided its binomial (avowedly following Tor-
rey & Gray 1838-43), vernacular name(s), and pertinent synonymy, followed by succinct notes on habitat
(including locality in some cases), flower color, phenology, and uses. The species were arranged into 93 orders
(families in current usage) “according to the natural method, or that of Jussieu.” Sanford (1851) reviewed the
finished report favorably:
The report of our friend Dr. J. H. Rauch, also of the City of Burlington, likewise merits the highest com-
mendation. Dr. H. [sic] has the honor of exploring the medical Botany of Iowa, and an attentive examina-
tion of his report, will be sufficient to convince any one of the fidelity and zeal with which he has per-
formed his task. We hope his labors will ... constitute a nucleus around which a full collection and
classification of our indigenous and naturalized plants may be made.
Subsequent cwiits.— At the conclusion of the Society’s 1851 meeting, Rauch and Dr. Moses Cousins, Jr
(1826-1868), were appointed “a standing committee on Medical Botany” (Ransom 1851; Anonymous 1854a).
Cousins, a native of Vermont and an 1850 graduate of Cleveland Medical College had recently set up practice
,n Albia (AnonynK,us 1878, Hickenlooper 1896). Like Rauch, Cousins “was a lover of natural science. Botaa,
was a iavonte study with him; he deUghted in the examination and cultivation of plants, fruit and ornamental
trees, vines etc. ...” (Anonymous 1869). One might wonder why, given these interests, he had not been ap-
pomtedtotheoriginal ad hoc committeeinsteadofBrookbankorDiflon. This was almost certainly
was nm among those present at that first meeting (cf. Matthews 1850; Sanford 1850).
committee, its successor seems to have achieved little. At the Society’s 1852 meetmg.
held 5 May in Fairfield, “the Committee on Medical Botany was given further time to report” (Anonymous
1854a). In fact, it appears from the official record that neither Rauch nor Cousins was pre
terofco™iuecsrna<icrepor.3a.theSucic.yM853.„ee,iug.held8Ju.,ei„^^^
honuf*eCom.„.r,ee on MedicalBotarryiurhe official record
TlKd.acovcryo(Cousinsl«tanfcaUnlerestandacttvi.viu,kuJL.,.r„ .',..1.
3rd (Anonymous 1854a).
ty in this period may solve a long-standing my
involving species attributed to Iowa by Alphonso W. Wood. As noted above, two species were reported from
the state in the second edition of his original Class-book (Wood 1847); the collector of the vouchers was not
indicated. In his all-new Class-book (cf. Merrill 1948), however, Wood (1861) indicated that “Dr. Cousens gen-
erously supphed us with the plants of the state of Iowa. His name often appears in our pages.” Curiously, Wood
did not indicate the doctor’s first name or initials, or his city of residence, though these were routinely given for
Me n»her contributors. Even those species not explicitly credited to Cousens apparently originated with him.
l remains of Wood’s herbarium (cf. Arthur 1884, Rusby ''
In what remains of Wood’s herbarium (cf. Arthur 1884, Rusby 19361 is an undated Iowa specimen ui zurnu/.-
Aeraauriculata (Michx.) Raf. (NY-29001) collected by “Dr. Cousens.” Wood (1861) reported this species from
Iowa (as Gerardia auriculata Michx.) but did not mention Cousens.
I have seen no unequivocal evidence linking Moses Cousins, Jr., of Albia to Wood or his Class-book. Nev-
ertheless, 1 consider him the most likely candidate to have been the mysterious Iowa contributor. The differ-
ence in spelling is no real impediment to this assumption, as Wood was well known for his many typographi-
cal errors (Meehan 1881). At ten places in the text, including the protologue of Pyrus coronaria var. i^nsis A.
Wood (cf. hammers 1985, 1998), it was spelled as in the introduction: “Cousens.” However, under Marsilea
vestitaHook. & Grev. (cf. Arthur 1884), it was rendered “Couzens.” Veratrum woodii was one of
aedited to “Cousens,”butitwasattributed to “Cozzens” when the specimen was cited elsewhere (Wood im
In only one instance did Wood’s spelling conform to the Iowa physician’s name. A revised printing Wood
1869) was created by replacing a few of the stereotyped pages with newly composed printing plates (Merrill
1948). On one such page. Aster anomalus Engelm. was reported from Iowa, and credited to Cousins.
The last evidence we have of Rauch’s botanical activities was his speech to the Sout wn owa or i -
turalSociety (Rauch 1853).Amajorportion of this address wasdevoted to urging the memtership^^^^^^
moreplants native to Iowa, particularly those “whichgrowuponom
Aosewhoknewall the faslhonable exotics, “but take theminto our fieldsandfo^^^^^
distinguish one plant from another.” He encouraged the group to be Amencan m our o
govemment”asLtin.that “our lily moccasin flower, butter-fly weed, orchis, dogbane, and many others will
-iefcrbeau,y„,,hany.ha.arefoundin other pamof the world,-Heindic«edjha.dun^^^^
« (ie., 1852), he had exhibited attractive r^tive plants at the group's ~ ” h^ t^l^ d
Sieater interest in growing them, and intended to do sointhecorningseasonasweU. Whether heactualh^
deconstruction Of- iMt ht to
Before the botanical significance of Rauch’s report can purpose I have chosen
'^«continuetobemade(e.g.,Lewisl998;Norrisetal.2001;Cusick2()0^
offers the single best summary currently available. ^ literature often differ from those used to
SynoitynOi.— Names employed to denote plants J classifications are hypoth-
denote the same entities today. These differences come about methods
not facts (Donoghue 1987; LaDuke 1987). As such, they change ov species is moved to a
ar^lysisbecoL available (Stnessy 2009). When clas^ficanon^^^^^
«'«ntgenus,orasubspecies is segregated asadis.inc^^sX^l^^j;:;'^'^^^^
community today has a comprehensive and rh^bestowing of names prior to the Twentieth
Ctyfeo/Bohnthnl Nomenclature (McNeill et
Century was governed by little more than custom and capnee,
^ of current rules and must be brought into compUance. e„y,onvm of one in Eders and Roosa
J by Rauch is a heterotypic ^ ^ ^^3 ponced that the nai
«»4). Poe example. Paper Birch was called Betulap<W«“ oZJhld four years earlier. SUver Maple v
mnfera pertained to the very same species but had been p
t its correct name, A. saccharinum, was erroneously applied
to Sugar Maple (A. sacchanim). Heterotypic synonyms in the list that follows are denoted by the abbreviation
“incl.” for “including.” Each also bears a parenthetical reference to support the name change. In selecting these
references, 1 have given preference to the eight editions of Gray's Manual (Gray 1848, 1856, 1862, 1863, 1867;
Watson & Coulter 1889; Robinson & Femald 1908; Femald 1950), because this work has long been a primary
A name used by Rauch may also be a homotypic (nomenclatural) synonym of one used by Eilers and
Roosa (1994). For example, Aspidium acrostichoides (Michx.) Sw. is based on the same type (that of Nephrodim
acrostichoides Michx.) as Polystichum acrostichoides (Michx.) Schott. All three pertain to the same biological
entity; they merely differ in the genus to which it is assigned. Most such synonyms are obvious from the uni-
formity of the final epithet and its author, though sometimes this is not the case. Either way, homotypy is indi-
cated by simply citing the synonym alone, with neither “incl.” nor reference.
Some differences have resulted from the earlier misapplication of a name to a taxon that does not include
the type of that name. Case in point: the erroneous use of Acer saccharinum for Sugar Maple instead of Silver
Maple noted above. Such misapplied names are not, properly speaking, synonyms; synonyms are created by
differing opinions as to the circumscription, position, and/or rank of a taxon, or by ignorance of priority w
homonymy (McNeill et al. 2006). These misapplied names simply represent errors of identification that be-
came established and pervasive in the primary literature. They are denoted in the list below by the abbreviation
“misappl.” for “misapplied,” followed by a parenthetical documentary reference. The work of Jones and Fuller
(1955) was especially helpful in sorting out misapplied names, by providing insight into how certain names
were used by other Midwestern botanists of the day.
Finally, if a name used by Rauch but not by Eilers and Roosa (1994) is neither a synonym nor a misappli-
cation, it must simply represent an error in identification. Ordinarily, such errors are rectified through exami-
nation of voucher specimens. With the loss of Rauch’s herbarium, disposition of such names becomes prob-
lematic. I have attempted to divine what plant Rauch had in hand by considering what other members of the
genus or family are present in Iowa that he did not report, which might logically be confused with the unbkely
species. In doing so, I have home in mind the state of botanical knowledge in the 1850s, specifically the refer-
ence works used by Rauch. In most cases, I am able to suggest a plausible disposition (e.g., the report of Betuk
glanduIosalikelybasedonmaterialofB.pumilavar.glanduIz/era).Inothers,thedubiousreportmustbeallowed
to stand (e.g., Xanthorhiza simplicissimd).
Format.— In the following analysis of Rauch’s checklist, Rauch’s Jussieuean arrangement of taxa is i«-
placed by the sequence used in Eilers and Roosa’s (1994) list. Nomenclature also follows this standard, unless
the name they used is contrary to the Code (e.g., Quercus borealis, Salix rigida). Rauch’s name if different follows
m brackets; see the preceding section for details on notation. Most of the ancillary information provided by
Rauch (flower color, phenology, uses) is omitted; only his statements on habitat and distribution are retained.
^ information from his address to the Southern Iowa Horticultural Society (Rauch
1853) IS quoted for a few species.
Species stated by Eilers and Roosa (1994) to be native to Iowa are in Roman type, while non-nativcsa«in
, the year it was first documented in the state (Cratty 1929) is given. Species not listed by
p srriyi, asterisked name indicates that it is not native to
Uion of these supernumeraries are derived frc®
3CJ wi.
rfwMichx, which
laramers, John Henry Rauch's Iowa flora checklist
CONCLUSIONS
y of Rauch’s (1851a) r
what we know about the Iowa flora today?” In answering this question, native and non-native species will be
considered separately, in order to have a consistent standard for comparison. The number of species found in
Iowa today is far greater than in 1850, due to the naturalization of plants from other regions (Cratty 1929, Eilers
& Roosa 1994, Lewis 1998, Norris et al. 2001).
Native piants. — Of the 135 families with species native to Iowa (Eilers & Roosa 1994), 83 (61%) were
represented in Rauch’s report. The most glaring absences were Cyperaceae, with 160 native species, followed
by Potamogetonaceae (18 spp.) and Juncaceae (16 spp.). ’SVhen one adds to this the fact that the report had but
one of the 150 native species of Poaceae, no Amaranthaceae, and no r
suspect that Rauch felt some aversion for apetalous herbs.
Of Iowa’s 1516 native species (Eilers & Roosa 1994), 348 (23%) were listed by Rauch. Among these were
some of very rare occurrence or very limited distribution within the state, e.g., Abies balsamea, Asimina triloba,
Drosera rotundifolia, Filipendula rubra, Ranunculus gmelinii, and Sassafras albidum. Conversely, the three-quar-
ters of the native flora not listed did include several exceedingly common and conspicuous species, the sorts
of things any neophyte botanist in the Midwest learns in his first season, e.g.. Aster novae-angliae L, Calystegia
sepium (L.) R. Br, Phlox divaricata L., Rosa Carolina L., Rudbeckia hirta L., Sagittaria latifolia Willd., Solidag)
canadensis L., and Tradescantia ohiensis Raf. The absence of such common eye-catching plants is puzzling.
Of particular interest are the species Rauch reported that were not listed by Eilers and Roosa (1994), ie.,
the 86 denoted in the list above by an asterisk. The lack of voucher specimens means that these species cannot
be stated with certainty to have once been members of the Iowa flora. However, their presence in the report is
suggestive of possibilities to be explored, through further collecting in the field and searches of out-of-state or
e of these supernumeraries (e.g., Dentaria diphylla, Lindera benzoinl there
ure reports, which may increase the credibility afforded Rauch’s report.
Particularly credible are the 38 species whose current distributional limits lie immediately adjacent to
Iowa, m maoy cases, in border counties. Some of these very well may have grown in Iowa prior to settlemenl.
but been extirpated subsequently Populations at the periphery of a species' distribution may be particularly
prone to extirpation in the wake of habitat loss due to agriculturalization.
The ranges of 24 of these extend into southeastern Minnesota and/or southwestern Wisconsin from the
Moling ttepamm of might once have been native but rare in northeastern^^
n. t a-rrrr ^ .... Maionthetnum trifolia, Panax trifolius, Picea mariana, Pinus resinosa, P^'
1, Toxicodendron vemix, Tri
arpon, V. axycoccos. Viburnum acerifoliu^
neglected in-sta
Gaultheria procumbens, Geum r
The other 14 have ranges that extend into riuimcasici
and east. Thus it is plausible that some or all once may have oeen native out rare in souincasici n luwtt. lunuw-
ing the pattern of Diospyros \irginiana L., Froxinus quadrangulata, and Sassafras albidum (Davidson 1959).
These are Alnus serrulata, Carya glabra (recently discovered specimens may document its existence in the
state), Cunila origanoides, Erigenia bulbosa, Eupatorium fistulosum, Gillenia stipulata. Hydrangea arborescens,
Hypericum adpressum, Lindera benzoin, Medeola virginiana, Ranunculus laxicaulis, Sabatia angularis, Trifolium
stolonferum, and Trillium sessile.
Another 36 supernumeraries occur in adjoining states, 1
Gaultheriahispidula, Oxalis montana, Poly gala paucifolid) do n
particularly closely. Although s
{ credibility overmuch, others such as Tax
hm distichum and Oenothera macrocarpa seem very unlikely to have been native to Iowa due to their habitat
rquirements. Such reports likely are due to misidentification of specimens or (see above) the misinterpreta-
tion of distributions reported in the literature (e.g., Artemisia santonicum, Gentiana acuta).
are especially puzzling, as they are not native to aajaceni siaies, o. n.
within hundreds of miles of Iowa: Aristolochia macrophylla, Euphorbia ipecacuanhae, Ipomoeajalapa, Firms
palustris, Scutellaria integrifolia, Vitis labrusca, and Xanthorhiza simpbcissma. Most are so distinctive in appear
ance that it is difficult to imagine what native plant might have been mistaken for them. Given tl
isof some economic value, one wonders if they were in fact cultivated rather than naturally occui
'‘”ttrX.--Of.He44Upec.es..a.edasno„-na..eb,B.e.^^
(15%). Does this mean that the remaining 355 had yet to he introduced to the state? Clear y, t ™ ^ “
(ct,Pohl 1959; Pohl&Sylvester 1962; Lammeis 1997; Cnsick 2002). "’"“rtZ«rrs
species overlooked (see above), it does not seem wise to assume that the a ence o a non ^
i.W„„tbeenintroducedye..Consequently,ademiled temporal
PWicable. We can say, however, that these 66 species ^ ,he year of first
1® after the territory opened to settlement. In every case>^
*icumenlation reported by Cratly (1929). Many were umntenti . Brassica nigra, Polygonum
Ml). Many have gone on to become widespread, even ubiquitous rn the sla>e. c.&. * chdidonium
^tariaTaraxac„mo/>i„ale.Others,despitetheirearlyintroduction,arestdlqu.tes^^^^^^
Lidtospermum arvensc, Sencdo vulgaris (Origanum
Fifteen non-native species were listed by Rauch of North America and
^gare, Ranunculus bulbosus, Rumex sanguineus) are sporadica y as cultivated; Allium
^ir presence in Iowa would not be surprising. The remaining ^ jssa ojficirudis, Nicotiana rustica, N.
Cucurbita pepo, Euphorbia lathyris, Ipomoea batatas, ^inifera. Plants of this sort were
Ricinus communis, Solanum tuberosum, Symphytum officina g carota, Mentha
otpressly omitted by Filers and Roosa. It is worth noting that very early date, others that
^ato, Pastinaca sativa) had already established themselves outsi e gar officinalis. Cannabis saliva,
naturalized now were only known to Rauch from cultivation (e.g.,
^^birapomifera).
SUMMARY
r-wvement In a Single field season of four or
This forgotten first checklist of the Iowa flora was an amazing This represented almost
^ months, a 22-year-old amateur newly arrived in the state oc ^ ^^^^ve and included 87 species not current-
^-qnarter of the native species known today and 15% o t e non mprhanical means of land ti
•y accepted a;
ng in tne state' He did this at a time when there
in a trontier region lacking scholarly resources, and w
T>espite the lack of documentary specimens, the report p
I srarvcv relatively unmolested; *
lov was relatively unmoiestea; a uui*.
^ve had peripheral populations within the state;
h m easiei 11, and southern species may
itefrmanynon-nanve species had invaded Uu:
state and supplanted the indigenes. It also shows us what was possible for dedicated workers to achieve in a
time when floristic study was avocational, a hobby for educated men rather than the livelihood of professional
botanists. Further, it is a cautionary tale, reminding us that documentary herbaria held by individuals are far
more vulnerable and ephemeral than those in the custody of institutions.
Rauch’s report also provides interesting insights into the medical profession of the mid-Nineteenth Cen-
tury. Mainstream practitioners of that era often are characterized as practicing few therapies beyond venesec-
tion and the cathartic purging of the gastrointestinal tract, and as employing a limited pharmacopeia built
around toxic mineral substances such as mercurous chloride, mercuric chloride, and potassium antimonyl
tartrate (Watson 1894; Fairchild 1927; Berman 1957, 1958; Lawrence 2003). Great animosity was said to exist
sonian. Reformed Botanic, and Eclectic physicians, who eschewed such “heroic”
I instead embraced a diverse array of drugs derived from plants (Beach 1833; Stuckey 1978).
The attention afforded medicinal plants by the Iowa State Medical and Chirurgical Society in its first years
of existence, however, suggests that this animosity has been exaggerated. Medical societies such as this one
were explicitly founded to counter the influence of those viewed by mainstream physicians as quacks and
charlatans. But the review of Rauch’s report by Sanford (1851) makes clear that the use of medicinal plants per
se was not grounds for denigration. Rather, it was the uneducated peddlers of “Indian cures,” “snake oil,” and
other nostrums who were condemned.
... thedevelopement of the medical resources of the Vegetable Kingdom, n
Abominable tricksters have deceived the people in proclaiming themseh
tssert, and defy any of them to
ACKNOWLEDGMENTS
I extend my warmest appreciation to Donna Hirst (Hardin Library for the Health Sciences, University of Iowa)
for providing copies of the proceedings of the State Medical and Chirurgical Society’s meetings; Ronald H. Sims
(Gaiter Health Sciences Library, Northwestern University) for permission to use their portrait of Rauch; Chris-
tine L. Chandler (Putnam Museum) for providing a copy of C.C. Parry’s 1847-48 field notebook; Dawn Rob-
erts (Chicago Academy of Sciences) for checking their holdings for Rauch collections; and John Pearson (Con-
servation and Recreation Division, Iowa Department of Natural Resources) and Deborah Q. Lewis (Ada
Hayden Herbarium, Iowa State University) for their careful review of an earlier version of this manuscript.
Amer.J. Sci. (ser.2) 17:297-308,353-365.
nf of the ichthyological fauna of the Pacific slope of North America, chiefly from the collections
Allen, J. 1846. Ca
1 68, 29th Congress, 1 st session, b
Anonymous. 1850. State medical convention. The Davenport {\owa) Gazette {27 June):2.
Anonymous. 1853a. Fire! Burlington (Iowa) Daily Telegraph (17 January):!.
Anonymous. 1853b. Disastrous fire. (Burlington) /owo State Gozefte (19 January):!.
Anonymous. 1854a. The transactions of the Iowa State Medical and Chirurgical Society, third and fourth sessions. Held at
Fairfield, May, 1852 and Davenport, June, 1853. Telegraph Printing Co., Burlington.
Anonymous. 1854b. Annual report of the president of the Maryland Historical Society, and list of its members. John D.
1:29.
a State Medical Society. The Davenport (Iowa) Daily Gazette
Anonymous. 1 856. October 6, 1 856. Vice President, Dr. H.A. Prout, in the chair. Trans. Acad. Sci. St. Louis
Anonymous. 1 857a. Iowa physicians. The Davenport (Iowa) Daily Gazette (1 8 August):! .
Anonymous. 1857b. Commercial and industrial cities of the United States. Number XLVIl. Burlington,
chants’ Magazine and Commercial Review 36:687-691
Anonymous. 1858a. Abstract of the Proceedings of the lov
{18June):2.
Anonymous. 1 858b. Historical Society. The Weekly (Burlington, Iowa) Hawk Eye (21 December):
Anonymous. 1 859. State Medical Society. Davenport (Iowa) Daily Gazette (10 June):! .
Anonymous [“D.R.B— 1. 1 869. Moses Cousins, M.D. Journal of Materia Medica 8:31-32.
Anonymous. 1 878.The history of Monroe County,
cal directory of citizens, war record of its volunteers in the
Northwest, history of Iowa, map of Monroe County, Const
Western Historical Co., Chicago.
Springfield, Illinois. Pp. 99-101.
. c*,*aD.. hi rn Chicaao.
the Senate of the United States
Anonymous. 1 882b. History of Scott County, Iowa. Inter-State Publ. Co., Chicagc
Anonymous. 1 887. Tuesday, March 26, 1 867. In: Journal of the executive procei
of America, vol. 1 5. Government Printing Office, Washington.
Anonymous [“N.'^. 1891 . The war on the State Board of Health. / '
Anonymous. 1892a. State Board of Health. One of the Republi
(Decatur, Illinois) Daily Review (4 October):8.
Anonymous [“W.H.B.'I. 1 892b. Obituary. Sereno Watson.
r.J. Sci. (ser. 3) 43:441-444.
ical. The Burlington (Iowa) Hawk Eye (28
Anonymous. 1894c. Death of Dr.
Anonymous. 1894d. Obituary. Sanitarian 32:420-422.
1894e. A noted physician is dead. Dr. J(
Do//yA/ews (24 March):!.
■xplred suddenly last night, tehgnon (Pennsylvania)
20, 0.SalicaceaeMirbel. in: Flora of lyodh American
Oxford University Press, New York. Pp. 3-164.
A»^R, J.C 1876. Contributions to the flora of Iowa; a
Carles City, Iowa.
^R.J.C. 1884.Contributionstothefloraof lowa.-1^o.VI.Proc.i;
Bawoey, T.M., L. Brouillet, and J.L Strother (eds.). 2006. Asteraceae
Oxford University Press, New York. Vols. 1 9-21 . Nat. Hist. 19{5):1-‘
««UE.O.,n„p,H.Monson., 954. Marsh and aquadcangiosperms of l<«_^
W. ,833. The American practice of medicine, 3 J^j;ji„gs of the InsOtote of Medicine of Chicago
®^tty,w.K. 1991. John H. Rauch - p
^.M.F. 1968. Edwin James. Scientist, linguisthumanitarian.Un,
A. 1957. The botanic practitioners of 19th-Century Amenca.
A. 1 958.The heroic approach in 1 9th-Century therapeutics. University
Professional Pharmacist 23-.866-870,
Medical Bulletin 24:41 9-427.
Bessey, C.E. 1871. Contributions to the flora of Iowa. In; Fourth biennial report of the board of trustees of the Iowa State
Agricultural College and Farm to the governor of Iowa. G. W. Edwards, Des Moines. Pp. 90-1 27.
Bray, E.C. and M.C. Bray (eds.). 1 976. Joseph N. Nicollet on the plains and prairies. The expeditions of 1 838-39 with jour-
nals, letters, and notes on the Dakota Indians. Minnesota Historical Society Press, St. Paul.
Brooks, R.E. 1983. Trifolium stoloniferum, running buffalo clover: description, distribution, and current status. Rhodora
85343-354.
Clausen, J., R.B. Channell, and U. Nur. 1964. Viola rafinesquii, the only Melanium violet nativ
COOPERRIDER, M.K. 1957. Introgressive hybridization between Quercus marilandica and Q. i
44:804-810.
Cratty, R.1. 1 929. The immigrant flora of Iowa. Iowa State Coll. J. Sci. 3:247-269.
CusiCK, A.W. 2002. Six non-native species newly discovered in the Iowa vascular flora. Sida 20:405-407.
Davenport, F.G. 1 957. John Henry Rauch and public health in Illinois 1 877-1 891 . J. Illinois State Hist. Soc. 50:277-294.
Davidson, R.A. 1959. The vascular flora of southeastern Iowa. Univ. Iowa Stud. Nat. Hist. 20(2):1-102.
Dillon, J.F. 1 850. Rheumatic carditis. - Autopsical examination. The Westerm Medico-Chirurgical Journal 1:1-6.
Donoghue, MJ. 1987. Experiments and hypotheses in systematics. Taxon 36:584-587.
Eaton, A. and J. Wright. 1840. North American botany; comprising the native and common cultivated plants, north of
Mexico: genera arranged according to the artificial and natural methods, 8*^ ed. Elias Gates, Troy, New York.
Eilers, U. 1975. History of studies on the Iowa vascular flora. Proc. Iowa Acad. Sci. 82:59-64.
Eilers, LJ. and D.M. Roosa. 1994. The vascular plants of Iowa. An annotated checklist and natural history. University of
Iowa Press, Iowa City.
Ellsworth, W. 1 922. Parry's catalogue of Iowa plants of 1 848. Proc. Iowa Acad. Sci. 29:339-344.
EPLiNG,C.1942.TheA
Fairchild, D.S. 1 927. History of medicine in Ic
s of Scutellaria. Univ. California Publ. Bot. 20(1 );1--
Iowa State Medical Society, Des Moines.
ind fauna: a symposium. Proc. Iowa Acad. Sci. 88:1 .
n America. Rhodora 36:241-261, 285-305, 314-344, 353-371,
), M.L. 1 934. Draba in temperate
392^104.
Fernald, M.L. 1 950. Gray's new manual of botany, 8th ed. American Book Co., New York.
Fite, G.C. 1966. The farmers' frontier, 1865-1900. Holt, Rinehart & Winston, New York.
Pp. 507-538.
Galland, I.
Chillicothe, Ohio.
Guy, CL 1947. The flo
Goodman, GJ. AND C.A.L
Norman.
Gould, F.W. 1942. A SI
p, and general descriptions of Iowa Territory. Wm. C. Jones,
a progress report based on past contributions. Proc Iowa Acad. Sci. 54:99-106.
1. Bull. Torrey Bot Club 33:387-396.
's 1820 expedition. University of Oklahoma Press
r.Midl. Naturalist 28:71 2-742.
Gwy, A. 1862. School and college edition. Manudi ui me uma.-x u. u.e
Giay, a. 1 863. Manual of the botany of the northern United States, including Virginia, Kentucky, and all east of the Mis-
sissippi. Fourth revised edition. To w/hich is added Garden Botany, an introduction to a knowledge of the common
cultivated plants. Ivison, Phinney, & Co., New York.
Gray, A. 1867. Manual of the botany of the northern United States, including the district east of the Mississippi, and
& Co., New York. i, u • i,-
(kr™, R.L 1847. Medical botany: or descriptions of the more important plants used in medicine, with their history,
d mode of administration. Lea & Blanchard, Philadelphia.
Gue, B.F. 1 903. History of Iowa from the earliest times to the begini
The Century History Co., New York.
Guiford, W.M. 1908. The burial place of Dr. John H. Rauch. J. Amer. Med. Assoc. 51:2228.
Guidner, LF. 1960. The vascular olants of Scott and Muscatine Counties with some refen
ry.Vol. 4. Iowa biography.
port, Iowa.
Hau,J. AND j.D. Whitney. 1858. Report on the geological survey u.
IS of the years 1 855, 56 & 57. The Legislature of Iowa, Des A
. -)>47n-471.
B. 1894. The late Dr. John H. Rauch, M.D.
1, F. 1 896. An illustrated history of Monrr
Tom its earliest period of organization
m R.C. Barneby. 1 982. The American Cassiinae. A synoptical revision
. ^«;-1-918.
IsaY, D. 1981. Leguminosae of the unitec
Mem. New York Bot. Gard. 25(3):1 -264.
" P. 1823. Account of an expedition from Pittsburgh to the Rocl^
J.C Calhoun, Sec'y of War:
New York Bot. Gard. 35:1 -91 8.
States. 111. Subfamily Papilionoideae:
'20, by order of the F
Lea, Philadelphia.
James, E.P. 1825. Catalogue of plants collected during a journey to a
during thirty years residence among the Indians in the interior of North Arne
IwEn, CC 1 850. Appendix to the report of the Board of Regents 0
the public libn
s, 1st Session, 1850.
^-GJ. 1855. Budington University, Burlington, Urbar
^S.N ™oG.D.FuaB. 1955. Vascular American plant i
“«tesz, J.T. 201 1. The Biota of North America Program {BONAhi.
jn, Baltimore.
erican Holiness Movement. Unpub-
univc^iAx- North and cenira. niiierica. Brittonia 60:1 1-33.
GJ. AND P.D. Sorensen. 2008. A revision of Agrimonia (Rosaceae 36:60-64.
^Dwe, J.C 1987. The existence of hypotheses in plant ,owa Acad. Sci. 92:125-128.
T.G. 1 985. Vascular plant taxa originally described from ■ ^ _g^2.
■*^.T.G. 1997. Heterotheca latifolia (Asteraceae), new to the Carruth. Iowa Native PI. Soc. Newslett.
"**««-T.G.1998.lowa'sveryownlLPrairiecrabapple/>yrus/oens^^ • •
4(2):3. Museum of Comparative Zoology, a
1 876. The Agassiz memoria ' ^ e _ .i..ee...nr, ift7
•^^rvard College, in Cambridge: tog«
'^'^tight & Potter, Boston. Pp. 40-54.
itions, and the practice of medicine, part one: Establishing a pro-
fessional identity, 1833-1886. Annals of Iowa 62:151-200.
Lea, A.M. 1 836. Notes on the Wisconsin Territory; particularly with reference to the Iowa District, or Black Hawk Purchase.
Henry S. Tanner, Philadelphia.
Lee, B. 1895. In memoriam. Death of Dr. John H. Rauch. In: Tenth annual report of the State Board of Health and Vital
Statistics of the Commonwealth of Pennsylvania. Clarence M. Busch, Harrisburg, Pennsylvania. Pp. 391-394.
Lewis, D. Q. 1 998. A literature review and survey of the status of Iowa's terrestrial flora. J. Iowa Acad. Sci. 1 05:45-54.
Lewis, W.H. and M.P. Elvin-Lewis. 1 995. Medicinal plants as sources of new therapeutics. Ann. Missouri Bot. Card. 82:16-24.
Mack, R.N. 1 991 .The commercial seed trade: an early disperser of weeds in the United States. Econ. Bot. 45:257-273.
Matthews, H.M. 1850. Proceedings of the medical state convention. Burlington (Iowa) Hawk Eye (27 June):6.
McKavEY,S.D. 1955. Botanical exploration of the Trans-Mississippi West 1790-1850. Arnold Arboretum of Harvard Uni-
a Plain.
It blue violets (Viola) of North America. Sida, Bot. Mist
1-60.
I. Burdet, V. Demouun, D.L Hawkswortw, K. Marhold, D.H. Nicolson, J. Prado, P.C. Silva, J.E. Skog, J.
al Code of Botanical Nomenclature (Vienna Code) adopted by the
ein, Germany.
Mead, S.B. 1 846. A catalogue of plants growing spontaneously in the state of Illinois, the principal part near Augusta,
Hancock County. Prairie Farmer 6:35-36, 60, 93, 1 1 9-1 22.
Meehan, T. 1 881 . Prof. Alphonso Wood. Gard. Monthly & Hort. 23:92.
Meehan, T. 1 898. The plants of Lewis and Clark's expedition across the continent, 1 804-1 806. Proc. Acad. Nat. Sci. Phila-
delphia 50:12-49.
Melius, TO. 201 0. Department of the Interior. Fish and Wildlife Service. Endangered and threatened wildlife and plants;
5-year status reviews of seven Midwest species. Federal Register 75:55820-55823.
Merrill, ED. 1948. Unlisted new names in Alphonso Wood's botanical publications. Rhodora 50:101-130.
Meyer, A.W. 1959. George Berry, Iowa naturalist. Iowa Bird Life 29:74-78.
Myers, R.M. and R.D. Henry. 1979. Changes in the alien flora in two west-central Illinois counties during the past 140 years.
Amer.Midl. Naturalist 101:226-230.
Newhall, lb. 1841. Sketches of Iowa, or the emigrant's guide. J. H. Colton, New York.
). A glimpse of Iowa in 1846; or, the emigrant's guide, and state directory; with a description of the new
1. 3. Oxford University Press, New
Norris, W.R., D.Q. Lewis, M.P. Widrlechner, J.D. Thompson, and R.O. Pope. 2001 . Lessons from an inventory of the Ames. Iowa,
- i. 108:34-63.
s, and a catalogue of the species, to the year 181 7. D. Heartt, Phila-
survey of the Northwest, during the se
StltesTrlTuI^^^partm^^^^ ^ Nebraska Territory, m^e under instructions from the United
>ARRY, C.C. 1 893. Professor David S. Sheldon, LLD. Proc^ a^rl'^rt Acid ^Nafs^S^^TO 1 84
asons. at m annual communications, from A.L. 5854 to A.L. 5858, inclusive. The Gra
and Accepted Masons, at
Lodge, Muscat!
Percy, j.F. 1908. Dr
PlKE,ZJVl.
to the sources of the Arkan;
H. Rauch--A pioneer in the fight against quackery. J
softheMissi
r. Med. Assoc. 51:2074.
;r of the government of
Piumbe,J. 1839. Sketches of Iowa
ritories. Chambers, Harris & Knapp, St. Louis.
Pom, R.W. 1959. Introduced weedy grasses in lo
Poll, R.W. AND E.P. Sylvester. 1 962. Dipsacus
,tate Publishing Co., Chicago.
Rjsateri,W.P., D.M. Roosa, and D.R.
Acad. Sci. 100:29-53.
Ransom, ED. 1851. Proceedings of the Iowa State Medical and
field. May 7, 1 851 .The Whig Office, Keokuk, Iowa.
I and Chirurgical Society, second
k)wa.Pp. 11-52.
Rauch, J.H. 1851b. On the topical application
1993. Habitat and distribution of plants special tc
inhabitants of large
R«Jch,J.H. 1878. The sani
>^,1X1 885. In: Can cholen
f^KHiNGER, K.H. 1 937. The North American species
J-L, G.E. Moulton, and A.E. Schuyler. 1999. The Lewis and Clark
comments. Proc. Acad. Nat. Sci. Philadelphia 149:1-64.
R«s,WJ.l859.Manual of publiclibraries,institutions,and societies, mthe united
R-C. 1 993. The Cruciferae of continental North f ^"^°'^54.26_27.
E.D. 1988. Early botany in the Trans-Allegheny 37235-237.
J.F. 1850. Iowa State Medical and Chirurgical Socfety. Second Annual Meeting,
J-F. 1851. [Review of] Proceedings of the Iowa Sta ^
hdd in Fairfield, May 7th, 1851. The Western Medico-Chirurgi
1915. Early Iowa locality records. Proc. 'o^a Acad Scl 2 •
,a Stud. Nat. Hist. 1!
s, past and present, ol
^'cago. J. H. Beers & Co., Chicago. Pp. 1 1 7-1 20. ^ settlement to 191 2. SJ. Clarke Publ. Co, Chicago.
A. 1912. History of Louisa County Iowa from its io4-i33.
432
Stuckey, R.L 1 984. Early Ohio botanical collections and the development of the state herbarium. Ohio J. Sci. 84:148-1 74.
Stuessy, T.F. 2009. Plant taxonomy. The systematic evaluation of comparative data, 2nd ed. Columbia University Press,
New York.
SzczYGEL, B. AND R. Hewitt. 2000. Nineteenth-Century medical landscapes: John H. Rauch, Frederick Law Olmsted, and the
search for salubrity. Bull. Hist. Medicine 74:708-734.
Thompson, J.D. 201 0. The vascular flora of Boone County, Iowa (2005-2008). J. Iowa Acad. Sci. 1 1 7:9-46.
Thompson, S.A. 2000. Araceae Jussieu. In: Flora of North America north of Mexico, vol. 22. Oxford University Press, New
York. Pp. 128-142.
Thomson, G.W. 1980. Iowa's disappearing woodlands. Iowa State J. Res. 55:127-140.
Thorne, R.F. 1953. Notes on rare Iowa plants. Proc. Iowa Acad. Sci. 60:260-274.
Thorne, R.F. 1 954. Present status of our knowledge of the vascular plant flora of Iowa. Proc. Iowa Acad. Sci. 61 :1 77-183.
Torrey, j. 1824. A flora of the northern and middle sections of the United States: or, a systematic arrangement and de-
scription of all the plants hitherto discovered in the United States north of Virginia. T. & J. Swords, New York.
Torrey, J. 1828. Some account of a collection of plants made during a journey to and from the Rocky Mountains in the
summer of 1820, by Edwin P. James, M.D. assistant Surgeon U.S. Army. Ann. Lyceum Nat. Hist. New York 2:161-254.
Torrey, J. 1843. Catalogue of plants collected by Mr. Charles Geyer, under the direction of Mr. I. N. Nicollet, during his
exploration of the region between the Mississippi and Missouri rivers. In: Report intended to illustrate a map of the
hydrographical basin of the Upper Mississippi River, made by I. N. Nicollet, while in employ under the Bureau of the
Corps of Topographical Engineers, Senate Document No. 237, 26th Congress, 2nd Session. Blair & Rives, Washington.
DRREY, J. 1845. Catalogue of plants collected by Mr. Charles Geyer, under the direction of Mr. I. N. Nicollet, during his
exploration of the region between the Mississippi and Missouri rivers. In: Report intended to illustrate a map of the
hydrographical basin of the Upper Mississippi River, made by I. N. Nicollet, while in employ under the Bureau of the
Corps of Topographical Engineers, House of Representatives Document No. 52, 28th Congress, 2nd Session. Blair &
Rives, Washington. Pp. 143-165.
3RRE^ J. AND A. Gray. 1838-43. A flora of North America : containing abridged descriptions of all the known indigenous
New Yo?k ^ according to the natural system, 2 vols. Wiley & Putnam,
Historical Society, Iowa City.
W»m S. ™ m Co«m. 1 889. Manual of the botany of the notthem United States, including the district east of the
"“"n u?'* and Tennessee, sixth edition, revised and extended westward to the lOOth
meridian. American Book Co., New York.
mr' TW 1!^ ?' *“0^ 52:147-148.
Wnnwroi T.W. 1947. Amerxian origin of the cultivated cucurbits. Ann. Missouri Bot. Gard. 34:1 01 -1 1 1 .
Jussieu. In: Flora of North America north of Mexico, vol. 3.
Oxford University Press, New York. Pp. 85-271
Wiersema, J.H. and CB. Hellquist. 1 997. Nymp
University Press, New York. Pp. 66-77
Ei 1 908. John Bartram's cypress tree. In: Notable American t,„.. counrry u
Wilson, J.G. A1 A ■
' ■ r.D. Appleton & Co., New York.
;. In two parts. P
t: Flora of North America north of Mexico, vol. 3. Oxford
s. Country Life in America 14:590.
. 1 847. A class-book of botany, designed for colleges, a
C Crocker & Brewster, Boston.
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A PRELIMINARY REPORT OF MESCALINE CONCENTRATIONS
IN SMALL REGROWTH CROWNS VS. MATURE CROWNS
OF LOPHOPHORA WlLLlAMSll (CACTACEAE):
CULTURAL, ECONOMIC, AND CONSERVATION IMPLICATIONS
M.Abul Kalam
Sul Ross State University
Dept, of Earth & Phys. Sciences
Alpine, Texas 79832, U.S. A.
mkalam@sulross.edu
Keeper Trout
P.O.BOX561
Alpine, Texas 79831, U.S. A.
Molly T. Klein
Department of Biology
Sul Ross State University
Alpine, Texas 79832, U.S.A.
Paul Daley
AiexanderShulgin Research Inst
1483 Shulgin Road
Lafayette, California 94549, U.S.A.
Diana Hulsey
Department of Biology
Sul Ross State University
Alpine, Texas 79832, U.S.A
Martin Terry
Department ofBioiogy
Sul Ross State University
Aipine, Texas 79832, U.S.A.
mterry@sulross.edu
abstract
INTRODUCTION
Lophophora wiHiamsii (Lem. ex Salm-Dyck) J.M. Couk. (Cactaceae) — commonly known as peyote— is a small,
spineless, globular cactus of northeastern Mexico and adjacent Texas (Fig. 1). It is of cultural and economic
importance for its use as the religious sacrament of the Native American Church (NAC). The harvested crowns
of peyote plants, either fresh or dried, are ingested orally as the sacrament in NAC ceremonies. Although L.
williamsH contains over 50 alkaloidal substances (Anderson 1996), such spiritual use of the plant is evidently
based largely on the psychoactive properties of an adequate dosage of its principal alkaloid, mescaline (Huxley
1955). The supply of peyote to registered NAC members is available through regulated channels involving the
Texas Department of Public Safety and a very small number of licensed peyote distributors — currently there
are three — who purchase their supply from agents (peyoteros) who harvest peyote from wild populations in
South Texas. The peyoteros are paid by the piece, according to the number of buttons harvested and delivered
to the place of business of a peyote distributor.
Over the past few decades peyote has become scarce in many parts of its historical geographic range. The
largest part of the reduction in peyote population size is clearly habitat destruction associated with urban
sprawl and adverse agricultural practices, notably root-plowing, which uproots and kills peyote along with the
native brush, effectively exterminating the peyote along with the associated plants of its natural habitat, so that
the damage to peyote in a root-plowed tract is absolute and permanent. Another major cause of the decline of
peyote is overharvesting of the plant for ceremonial use by the NAC. Overharvesting of peyote has several dif-
ferent adverse effects on the wild populations:
(1) It reduces the harvestable population size, and selectively removes the largest crowns first, as these are
most valued in the peyote market.
(2) That reduces the quantitative reproductive output of the population, as a direct consequence of the removal
of the largest crowns that produce most of the seed in an unharvested population. In terms of population
genetics, while such selective harvesting of the largest plants may not have a marked short-term effect on
population size (assuming good harvesting practices, benign weather, and consequently a low mortaUty
rate in harvested individuals), it has the immediate effect of reducing the effective population size.
(3) Concomitantly, there is a qualitative genetic loss in the selective loss of seed production from the oldest in-
dividuak, which are ipso facto best adapted to local conditions. That loss may be temporary, if the har-
vested plants survive to produce regrowth buttons that are allowed to mature after a few years, or it may
be ^rmaiient, if mortality occurs in the old plants due to repeated harvesting of regrowth buttons (Terry
(4) The phenomenon of post-harvest regrowth of new crowns arising from areoles of the subterranean stem
(Terry & Mauseth 2006) temporarily increases the number of crowns in the population, but severely
201 total combined weights of crowns in the population (Terry et aL
(5) The decreased size of peyote buttons available to the NAC means that an individual in an NAC peyote cer-
emony must consume more buttons to equal the weight of the smaller number of buttons that would be
consumed if mature crowns were available. This leads to a vicious circle of more frequent harvesting to
supply the demand for greater numbers of buttons, which leads to the early harvesting of yet smaller
buttons-but now fewer, as the overharvested plants exhibit signs of decreased energy reserves for the
producuon of more new crowns Mowing repeated harvesting at two-year intervals (Terry et al. 2012)-
(6) Lookmgatthe quality of peyote for ceremonialuseinpurelypharmacological terms, there wouldappearto
be yet another disadvantage to regrowth buttons for ceremonial use apart from their small size, and that
is the ^bility that the dry-weight concentration of mescaline in the small regrowth buttons is sub-
stantiaUy lower than in mature peyote crowns, which now constitute a minor percentage of the total of-
fering in the regulated peyote market.
Accordingly, the purpose of the current study was to
• address the hypothesis that small regrowth
w atprials and methods
h Afresh weight) were collected from 14 in i-
m small regrowlh peyote crowns
plants in a South Texas population m March 201 •( :„ht from each individual) were field-collec
rthgroup^)Biopsysamplesofcrowntissue(ca.4g res ^
aOmatureCelght-ribbed) individual were s«
i to as the “Mature group'.) The cactus tissue from eac P P „f each group was ground
for a week on a drying rack at room temperature. 0"« <■'“ ^ pUntsamples in each group were
epowderwithamortarandpestle.A.tha.point^^^^^^^^pAsample^^
wed to constitute a single homogeneous sample t P of HPLC-^e m.
tissue fromeachgroupwas then placedina2M^^
Ided. The beaker for e-b samole was sealed with Pa ^ effect alkaloid extracti
El 111
438
Journal of the Botanical Research Institute of Texas 7(1)
the aqueous and dichloromethane layers were then allowed to separate overnight. The rationale was that pro-
tonated phenethylamines such as mescaline and protonated tetrahydroisoquinolines such as pellotine remain
in the acidified aqueous layer, while less polar substances dissolve preferentially in the dichloromethane layer.
The latter, containing fats and other nonpolar comp)ounds, was drained out of the separatory funnel and set
aside. Two additional defatting extractions of the remaining acidified aqueous phase in the separatory funnel
were done, each with 50 mL of dichloromethane, and in each case the aqueous and organic layers were allowed
to separate, whereupon the dichloromethane phase was drained from the separatory funnel and discarded.
After the third defatting extraction of the acidified aqueous phase, the aqueous phase was drained from
the separatory funnel into a beaker, and its pH was raised from 3 to 12 with sodium hydroxide (5 M), in order
to deprotonate close to 100% of the mescaline (pK^ = 9.5). The alkalinized aqueous extract was then poured
into a separatory funnel, and 50 mL of dichloromethane was added. The rationale was that alkalinizing the
aqueous phased in this classic acid-base cleanup procedure was that the deprotonated alkaloids would now be
more soluble in the nonpolar dichloromethane than in water. The organic and aqueous phases were mixed well
and left to separate. The dichloromethane layer was drained into a 200-mL beaker and saved. Another 50 mL
of dichloromethane was then added to the remaining alkaline aqueous phase in the separatory funnel. Upon
separation of the organic and aqueous phases, the organic phase was added to the first dichloromethane ex-
tract, and the combined extracts were left in a hood at room temperature to evaporate to dryness. The residue,
containing the mescaline and related alkaloids, was then redissolved in 10.0 mL of methanol and stored in the
freezer in a sealed vial. Samples of the two extracts (Small Regrowth and Mature) were run on an Agilent 1260
Infinity HPLC with a diode-array detector (G4212B) set at 205 nm, using 70% methanol (HPLC grade) as the
mobile phase, and a Phenomenex Gemini 5p CIS column as the stationary phase. Samples of 1.0 pL were in-
jected with a flow rate of 1.0 mL/min and run for 30 minutes. Each of the two samples of alkaloid extract, after
appropriate dilution to obtain on-scale HPLC peaks, was run three times, and the values of area under the
curve (AUC) of the three runs per sample were averaged. Appropriate dilutions were made of mescaline stan-
dard and run on the HPLC three times under the same conditions described above, to create a standard curve.
The mean AUC value for each sample of alkaloid extract was then interpolated on the standard curve to yield
the corresponding weight (in pg) of mescaline in the HPLC sample of alkaloid extract. From the latter value
(one such concentration value per sample group), we calculated the original concentration of mescaline in the
homogenized desiccated tissue sample from the Small Regrowth group and in that of the Mature group.
Confirmation of the identity of mescaline in these samples was achieved with GC/MS. A sample metha-
nol extract of L. wlliamsii from this study was evaporated to dryness under an stream at room temperature,
and the residue dissolved in dichloromethane for analysis. The instrumentation was an Agilent 6890 GC
equipped with a DB-5ms (0.25 mm l.D. x 30 m) column, splitless injection (250°) and an Agilent 5972 MSD
(transfer line at 275°), operated in full scan mode. Helium was used as the carrier, and the oven program was
70° (with a 1-min hold), then 20°/mm to 250°, with a final hold. The mescaline peak had an identical retention
time to that of an authentic reference sample, and the mass spectrum matched a spectrum in the NIST database
(Stein et al. 2005) with 93.5% probability of best match.
Group mescaline concentrations for the Small Regrowth and Mature groups sampled are presented in Table 1.
Thegroup values rqx«ed are physicalaveragesresuhingfron. the within-grouppoolingandhomorn^^^
of tissue Mrnples of the Individual plants sampled and are not statistical means. (A forthcoming manuscript
ftontour lab »ill report mescaUnecoucentralion data onasubstantially larger number of individuaU in differ-
enthfeslages and size iauges.withstalistical analysis of individualvalues of mescaline concentrationincrcwo
nsst^.) Mesealme concentrations »ere 3,80 g/lOOg tissue in the Mature group and 2.01 g/100 g tUsue in d*
Small R^rowth group, based ontissuesamples>2gfreshweighl from lOindividualsinthe Mature group and
14 individuals m the Small Regrowth group. The difference between these levels amounted to a 47% reduction
3 the Mature group. A gas chromatogwffl
439
t rr neak from peyote extract are presented in
f peyote extract and a mass spectrum of the most prommen
igs. 2 and 3, respectively. trations are reduced in small regrowth crowns
Ttiese daa support the hypothesis that mescaline conan p„vioosly harvested. This
f Wole in comparison to the concentrations in
anfinns the validity of the widespread opinion among another dimension to the damage
r” than larger, mature peyote crowns (T. Herrera, pers. o crowns have been largely replaced
ang done by the too frequent harvesting of peyote in Sout exas. ceremonial dose of the
y small regrowth crowns in the peyote market. In order to o number of small b
‘em,anNACparticipantinapeyotemeetingmustconsumeasubstam ..
would be the case with mature buttons. The finding in the current stud^^^^
»leed lower in
uttonsbyeach
appropriate spiri-
«n*byeachpar.icipan.inanNACmeeiinginorfer.ogf.-enna^----_^^^
State. Such compensatory increased consumption ^ , for the small size of the regrowth
^in addition to the increased consumption require leading to increased consumption
“ns—can only exacerbate the vicious circle of ^his vicious circle has been seen (MT
ing to increased harvesting. At the population level m ’ for l. mllumsii.
T.pers. obs.) to progress downward to the bottom of the extmcuo
Journal of the Botanical Research Institute of Texas 7(1)
ACKNOWLEDGMENTS
We are most grateful to the Alvin A. and Roberta T. Klein Foundation, Libbie and Jerald Mize, the Cactus Con-
servation Institute and its supporters, Sul Ross State University (Research Enhancement Grant), and the Welch
Foundation (Grant# AK-0023), for support of this work. We also greatly appreciate the generosity and under-
standing of the C.W Hellen Ranches, Ltd. - La Mota Division - Charles W. (Bill) Hellen, Managing Partner, for
access to the land where the field research was conducted. Norma Fowler provided helpful conceptual com-
ments on the project. Kelly Kindscher did a thought-provoking review of the manuscript, resulting in a num-
ber of needed improvements. Chris Ritzi and Karen Little were most helpful in providing lab facilities.
Andcrson, E.F. 1996. Peyote: The divine cactus. University of Arizona Press. Tucson.
Huxley, A. 1 955. Letter of 24 October 1 955 to Humphry Osmond. In: Horowitz, M., and C Palmer (eds.). Moksha: Aldous
Huxley's Classic Writings on Psychedelics and the Visionary Experience (1931-1963) Stonehill Publishing Company.
New York. P.81.
AND D.SPARKMANN.2005.The NIST
2.0 d. FairCom Corp., ChemSW,
(C^cT^^Ca^ R<»t-shoot anatomy and post-harvest vegetative clonal development in Lophophora
Terry. M.. K. Trout, B. Wiluams, T. Herrera, and N. Fowler. 201 1 . Limitations to natural production of Lophophora
(Cactaceae) I. Regrowth and Survivorship two years post harvest in a South Texas population. J. Bot. Res. Inst. Texas
s to natural production of Lophophora
1 a South Texas population. J. Bot. Res. Inst Texas
POTENTIAL DISTRIBUTION OF THREE NATIVE AND
ONE INTRODUCED GRASS SPECIES IN SEMIARID HIGHLANDS
OF MEXICO USING GIS TECHNIQUES
Armando Cortes Ortiz, Yolanda Herrera Arrieta,
Jesus Herrera Corral, and Daniel Hernandez Velazquez
ABSTRACT
introduction
, unUke any o
conditions. Different species require diverse conditions: suu*e plants will
while others require a different one (Ederra 1997). ,rv fFie L - i nmo
Horisuc studies of the north cenu.lregkm of
'‘“I'oaceK family (Herrera 2001;HerteraaPe>ersmM7^^^^^^^^.^^^„theevadab.hty of then
Cortes & Herrera 2011; Hen
cal modeling studie
nprep); their distributional
Fk. 1. Geographical location ofthe study area.
geographical distribution, ecological and ethnobotanical adaptation of a group of plants with geography,
ecology, climate and human settlements... ” (Guarino et al. 2002), can be used to analyze this distribution.
Ecological and geographical information on species is fundamental for conservation, planning, and prediction
(Ferner 2002; Funk & Richardson 2002; Rushton et al. 2004), as well as to reach an understanding of factors
which determine the patterns of spatial distribution of biodiversity (Elith et al. 2006; Ricklefs 2004). Ingen-
from ranXrsampli^^^ Ihaf^dT^""^^ of species are scarce and of presence-only since they generally come
^Xsluer^'t ""rwide "PP^ioations (Cortes & Herrera 2011). Spatial distribution
1 r ^ L ictions ol the distribution of species by connecting their presence with envi-
ronmen.al factors. Th^ have been applied in the study of telatiottships between environmental parametets
and s^c.« rtetaess (Herrera & CortCs 2010; MacNally & Fleishman 2004) and invasion by non-native spe-
aes(G^Uby20O4me present work attempts toanalyrealarge quantity ofdata compiled through noraand
faunastudtes.nordertotmprovecurrentknowledgeofspeciesdistribution.
Tothtsendl^ationdataCUtitudeandlongitudelforapproximatelySOOgrassspecieshavebeenobtained
nnrtrZir "™«=''““^Cortes&Herrer^
Tras^TitT Tb TT
grass spcctes. From these, asuto of 4 grasses from north central Mexico were chosen: 3 native and 1 in»-
ecosvstems^ w'’'T T c <>f 4 species indicate that they occupy dlvet«
CortK et al., Predicting plant distribution with GIS
porteri, and Melinis repens) collected in the study area were analyzed with Geographic Information Systems
(GIS) software, specifically the DIVA-GIS 7.1.7.2 version (http://www.diva-gis.org/), and GVSIG 1.9 for Vista
(http://www.gvsig.org/), using geo-statistics maps of the States of Chihuahua, Durango, and Zacatecas (INEGI
tabase (http://worldclim.org/current). The GIS software packages were developed for the analysis of species
diversity and distribution in order to clarify geographical, ecological, and genetic standards (Hijmans et al.
2004) and contain the necessary algorithms to generate the required data in a simple manner.
The study area encompasses the territory of the States of Chihuahua, Durango, and Zacatecas in the north
central Mexico, occupying approximately 446,000 km^ (Fig. 1) located between the extreme coordinates
21°02'24" to3r46'48"N and 109°04T2" to 100°44'24''W.
This area is also located within the physiographical provinces of the Sierra Madre Occidental and Sierras
del Norte, as well as partially within the Mesa del Centro (INEGI 2003b). As such, the topography of the study
area is highly varied and rugged with altitude variations between less than 500 m and up to 3200 m. Its latitude
position, the distance to and influence of the maritime zone, and the topographical characteristics of this zone
determine strong variation in climatic factors such as temperature and precipitation. Within this region are
low-altitude areas with an annual temperature ranging between 24 to 26°C and high-altitude areas with annual
temperatures ranging between 8 and 10°C. In terms of precipitation the study area has dry zones with little
annual precipitation (200 mm) as well as sub-humid zones that have an annual precipitation between 1000
and 1500 mm. Interrelations between these and other factors cause the study area to have varied plant commu-
nities and soil cover from zones with pine and oak forests to grasslands and xerophilus scrubs (microphyll
desert scrub, rosette desert scrub, crassicaule scrub), as well as small pockets dominated by halophilous scrub
communities or halophilous grasslands, due to salty soils (INEGI 2003a).
The potential distribution analysis was carried out using a subset of four (out of 500) species belonging to
diverse environments in order to determine the potential distribution and climatic characteristics of the distri-
bution area. A total of three native species were selected, Muhlenbergia peruviana from temperate semiarid
forests; Bouteloua gracilis from grasslands; Muhlenbergia porteri from the xeric region; as well as one nonnative
introduction, Melinis repens (Zizka rose Natal grass). The data used to carry out our analysis came from speci-
mens kept at several herbaria in Mexico (CHAPA, CIIDIR, ENCB, HUAA, lEB, MEXU, SLPM) as well as abroad
(TAES, US) that had been included in previous floristic inventories of the study area (Herrera 2001; Herrera &
Peterson 2007; Herrera & Cortes 2009, 2010; Herrera et al. 2010; Cortes & Herrera 2011).
Potential species distribution prediction was carried out using the Bioclim model included in the Di-
va-GlS program applied to Global Climate Data from the WorldClim bioclimatic database (http://www.
worldclim.org/current). In brief, the Bioclim model identifies patterns of geographical distribution of species
from 19 bioclimatic variables (contained in the database) and determines areas where species could thrive.
Unlike other models, this model uses presence-only data (Kenth & Carmel 2011; Rodriguez-Soto et al. 2011),
h are within the climate cover that have environmental characteristics similar to
under study. Bioclim can be used to describe the environment in which a certain
^ found in order to identify other areas where the species can live in the current
condition as well as to identify areas where the same species could be present if the climate landscape alters.
The WorldClim bioclimatic database is a set of global climate layers (raster format) with a spatial resolu-
tion of about one kilometer. It can be used to generate maps or create a spatial model in GIS or other computer
software. This database was generated by the interpolation of observed and representative data from the years
^^50 to 2000 (Hij-mans et al. 2005).
Creation of prediction maps of distribution.— Potential distribution maps were generated for the four grass
species with the Bioclim model initially classifying the area into the following categories of probability of oc-
currence: Unsuitable, Low (0-3^5 percentiles). Medium (3.5-5 percentiles). High (5-10 percentiles). Very High
U0'20 percentiles), and Excellent (20-34 percentiles). In order to determine the main climatic characteristics
relevant to the species distribution, the maps were remodelled so that only cells/areas with high, very high and
^^cellent probability were shown. ’
Creation of temperature and precipitation maps for each species. — Maps of the i
and annual precipitation of the study area were generated from the climate database
Afterwards, using overlapping map functions, annual precipitation and mean annu
generated for the high-probability distribution areas of each species.
maps were cross-mapped so that fi
precipitation and temperature val
perature values was obtained.
laps.— The annual precipitation and mean annual temperature
'try species, each high-probability distribution map cell also showed the
Furthermore, the cell frequency for the various precipitation and tem-
s (mode) were 500
Maps of the predicted distribution of four grass species were generated with the Diva-GIS Bioclim algorithm.
The selected species were Muhlenbergia peruviana, Muhlenbergia porteri, Bouteloua gracilis, and Melinis repem,
which were chosen because they have been collected in different ecosystems and our results show the variety
of climate conditions in which they can develop. Said maps show the level of probability of occurrence for each
of the species in the study area (Guarino et al. 2002; Hijmans et al. 2004).
Muhlenbergia peruviana is listed as a species commonly distributed in pine-oak forest communities and
the potential distribution map that was generated (Fig. 2) shows that areas of the Sierra Madre Occidental have
greater probability of occurrence for this species.
When the mean annual temperature and total precipitation maps of high, very high and excellent prob-
ability of distribution areas of M. peruviana were obtained, we found that this species is located in areas witha
relatively low mean annual temperature between 12 and 17°C, particularly at 14°C. Furthermore, the annual
precipitation ranged between 600 and 1000 mm, notwithstanding the fact that the largest area was comprised
in places with 600 mm (Tables 1 and 2). With this in mind, it is possible to establish that the distribution of M.
The potential distribution map of Bouteloua gracilis, which is known to be mainly distributed in natural
grasslands and within some xerophilus scrubs (Fig. 3), shows a large area with high to excellent values of po-
tential distribution.
The distribution area of B. gracilis, with greater likelihood values, had ann
400 to 800 mm, with an average of 560 mm although the most common precipitation vah
and 600 mm (these had the highest percentage of surface area). The r
our model showed a range between 13 and 19°C, with an average of 16.2°C and a mode located at 17°C.
The potential distribution map of Muhlenbergia porteri (Fig. 4),
rophilus scrubs (Fig. 4), placed this species within the driest regions of the study a
The area with the highest predicted distribution of M. porteri had a mean a
between 12 and 21°C, with the mean located at 17.8°C and the mode at 18°C, while the an
ranged between 300 and 900 mm with a mean centered at 482 mm and a mode of 400 mm.
Climatic requirements for each of the three native species are summarized in Table 3
expected, they belong to different ecosystems, j h t it
The potential distribution map of the nonnative introduced grass Mdinis repens (Fig. 5) showe t ^
had the smallest area of likely distribution. It is possible that these results are influenced by the fact t at
species was represented in low numbers with all collections used in this study. ^
Nevertheless, the distribution area with the highest likelihood of having Melinis repens showed
nual temperatures ranging between 14 and 18°C, with an average of 16°C and a mode also at 16“C.
nual precipitation reached an average of 609 mm, with a range between 500 to 800 mm and a mode o
3 (Herrera 2001; Herrera & Peterson 2007; Cortes & Herrera 2010; Herrera & Cortes 21
445
N. 2. Map of the potential distribution ofMuhlenbergiapenimna obtained with Biodim.
rera et al. 2010; Herrera et al. in prep) allowed us to construct probable distribution maps of close to 500 species
of which the actual areas of distribution are known; this work shows the results from a subset of four selected
species. The distribution of grass species in northern Mexico is mainly determined by climatic factors (Rze-
dowski 1978); therefore it can be assumed that this same analysis applied to the remainder of the grass species
within this area has a very high probability of increasing our knowledge on their potential distribution.
Journal of the Botanical Research Institute of Texas 7(1)
Tabu 1 . Proportion (%) of cells/areas with high probability of distribution ofMuhlenbergiaperuvianawiti\ regards to mean annual ti
Table 2. Annual precif
In this study we modeled the potential distribution of four grass species from the arid and semiaridre
gions in northern Mexico, allowing us to predict the area in which it is more likely to find each of them as w
as to determine the main climatic characteristics necessary for each species to be present.
The maps of potential distribution of Muhlenbergia peruviana, Bouteloua gracilis, Muhlenbergia porttn
and Melinis repens, generated with the Bioclim model using 19 bioclimatic variables, show the probab’ ity^^
distribution of these species within the study area in regard to their bioclimatic characteristics. These mapsa^
lowed us to determine the annual precipitation and mean annual temperature conditions in which eac spe
has been found. The possible distribution areas can be considered to be critical for further fieldwork as th^
show similarities in the main climatic characteristics with those areas where these species are
Our analysis clearly shows that the three native species that were selected grow under very
matic conditions, lending support to our initial decision of selecting species from different
Nevertheless, Melinis repens (the introduced species) was shown to share a large ecological niche wit
loua gracilis (the grasslands species) since the former has climatic requirements similar to those of the a
This fact could mean that this African species would not be able to invade other communities, such as ^
or xerophytic scrubs, only posing a problem for grasslands. With our model it is possible to apprecia e
tual distribution shared between a native species and an introduced species. hvooth'
The replacement of native species with less desirable introduced species affects biodiversity. The hyF^
esis that biodiversity has an influence on productivity and stability of the grassland ecosystem has been p^^^
ously tested in European grasslands by Hector et al. (1999) and in North American grasslands by
(1997). Both groups agree that the loss of biodiversity directly affects productivity and stability
One of the grasses that was studied in our work, Melinis repens (Zizka rose Natal grass), is an
species and the prediction of critical areas of future expansion is an important issue in order to preven
vasion and later displacement of native grassland species in Mexico. Melinis repens is an African grass that was
recently introduced into Mexico (less than 50 years ago), first found at the edge of roads, a fact that possibly
helped this species to spread, and it now invades and displaces native species in grasslands in northern Mexico
(Herrera Arrieta et al. 2011; Herrera Corral et al. 2011). This grass is highly aggressive and this characteristic
could be allowing it to increase its cover and dominate the distribution areas of Bouteloua gracilis (blue grass).
448
Fk. 4. Map of the potential distribution of Mt/Wenberff/oporferi obtained with Biodim.
However, Figure 5 shows that climatic conditions could be containing Zizka rose Natal grass from spreading
into every space that blue grass now occupies within these grasslands (see Fig. 4).
CONCLUSIONS
With this study we have shown that it is possible to carry out prediction modeling of the potentia
or ecological niche of grass species based upon presence-only data (Kent & Carmel 2011; Rodriguez
2011) that can be obtained from specimens collected with floristic inventory purposes.
Also, .his study demonstiauis tha. .he overUp in distribution bettueen naUve atrf invasive J*
shown, providing valuable inromtation that can be used for preventing the spread o mvastve speaes and the
displacement of native species. We also uncovered evidence that it is possible that climatic requireraen ar
preventing the expansion of the African-introduced species Melinis repens (Zizka rose Natal grassXalthoughn
lias spread into grasslands in northern Mexico during the last 40 years.
450
n on the potential distribi
s as well as for determining areas th
should provide support for decisic
n of grass species that could be useful in fu-
■elating to conservation and management of
ACKNOWLEDGMENTS
We want to thank Instituto Politecnico Nacional for financial support as well as the support granted to the fol-
lowing projects: “Floristica de Gramineas de Durango” DGPl-1998045 and CONABIO-R035, “Floristica de
Gramineas de Zacatecas” SIP-20070429 and CONABIO-EE014; and “Horistica de Gramineas de Chihuahua"
SIP-20100879 and CONABIO-GE003; in which the data analyzed in this work was obtained. We also want to
thank J.J. Ortiz and an anonymous reviewer for constructive reviews that improved this work. Helen Jeude
(BRIT) read and improved the English.
Zacatecas, Mexico. J. Bot. Res.
REFERENCES
2011. Distribucion y diversidad de la familia Poaceae en Chihuahua, Durango y
nst. Texas 5(2):689-700.
IS plantas y el equilibrio ecologico de nuestra Tierra, 2a Ed. Ediciones
Universidad de Navarra, S.A., Pamplona, Espana.
Elith, J., C.H. Graham, R.P. Anderson, M. Dudik, S. Perrier, A. Guisan, R J. Humans, F. Huettmann, J.R. Leathwick, A. Lehmann, J. L, L
Lohmann, B.A. Loiselle, J. Manion, K. Richardson, R. Scachetti-Pereira, R.E. Schapire, J. SoberOn, S. Williams, M.S. Wisz, and N.E.
ZiMMERMANN. 2006. Novel methods improve prediction of species' distributions from occurrence data. Ecography
29{2):129-151.
Ferris, S. 2002. Mapping spatial patterns in biodiversity for regional conservation planning: where to from here? Syst
Biol. 5 1(2):33 1-363.
Funk, V.A. and K.S. Richardson. 2002. Systematic data in biodiversity studies: use it or lose it. Syst. Biol. 51 (2):303-316.
Goolsby, J.A. 2004. Potential distribution of the invasive Old World climbing fern, Lygodium microphyllum in North and
South America. Nat Areas J. 24(4):351 -353.
Guarino, L, a. Jarvis, R J. Humans, and N. Maxted. 2002. Geographic Information Systems (GIS) and the conservation and use
of plant genetic resources. In: J.M.M. Engels, V. Ramanatha, A.H.D. Brown and M.T. Jackson, eds. Managing plant ge-
netic diversity. 12-1 6 June 20
y.Pp 387-400.
HEaoR, A., B. Schmid, C. Beierkuhniein, M. Diemar, and P.G. Dim^iakopoulos. 1 999. P
in European grasslands. Science 286:1 123-1 127.
IS de Durango. Instituto Politecnico Nacional-Comision Nat
nilia Poaceae para Chihuahua,
Herrera A., Y. & P.M. Peterson. 2007. ^
He«« & A. Corns Om,. 2009. Diversidad y distrlbuoidn dVia7GramIeard7za';
Herrera Arrieta, Y. & A. CoRrts Ortiz. 201 0. L
Durango y Zacatecas, Mexico. J. Bot. Res. Inst. Texas 4(2):7l'l-73’8. '
HiiMR. to,™ V, P.M. toBON & A. Coere Omz. 2010. Gramineas de Zacatecas, Mdxico. Sida, Bot. MIsc. 32:1-239.
I. Changes of vegetation
i Durango region, north Mexico.!
Herrera Arrieta Y, P.M. Peterson, and A. CortEs Ortiz. Gras
Herrera Arrieta, Y.,D.S.F
and diversity in grasslands, along 28 years of continuous grazing ir
Anim. Veterin. Advances 10(22):29 13-2920.
^^Cambto Tn t N. Almaraz Abarca, N. Naranjo JimEnez & FJ. GonzAlez GonzAlez. 2011-
F*. Mathur, I. Barrantes & E. Rojas. 2004. DIVA-GIS Versibn 4. Sistema de informaci^n
Cortes et al.. Predicting plant distribution with GIS
451
geografica para el analisis de datos de distribucion de especies. Manual, International Plant Genetic Resources Insti-
tute (IPGRI),y UC Berkeley Museum of Vertebrate Zoology, Berkeley, CA,USA.
iNEGi. 2003a. Diccionario de datos de uso del suelo y vegetacion, escala 1:250 000 (vectoriales). Institute Nacional de
Estadistica, Geografia e Informatica, Aguascalientes, Mexico.
INEGI. 2003b. Diccionario de datos fisiograficos, escala 1:1000 000 (vectoriales). Institute Nacional de Estadistica, Geo-
grafia e Informatica, Aguascalientes, Mexico.
Kent, R. andY. Carmel. 2001. Presence-only versus presence-absence data in species composition determinant analysis.
Diversity & Distrib. 1 7:474-479.
MacNally, R. and E. Fleishman. 2004. A successful predictive model of species richness based on indicator species. Con-
servation Biol.18(3):646-654.
Ricklefs, R.E. 2004. A comprehensive framework for global patterns in biodiversity. Ecol. Letters 7(1):1-1 5.
Rodr(guez-Soto, C., 0. Monroy-Vilchis, L. Majorano, L. Boitani, J.C. Faller, M.A. Briones, R. NOSiez, 0. Rosas-Rosas, G. Ceballos, and
A. Falucci. 2011. Predicting potential distribution of the jaguar (Panthera onca) in Mexico: identification of priority
areas for conservation. Diversity & Distrib. 1 7:350-361 .
Rushton, S.P., S.J. Ormerod, and G. Kerry. 2004. New paradigms for modeling species distribution. J. Appl. Eco. 41(2):
Rzedowski, J. 1 978. La Vegetacion de Mexico. Limusa.
Tilman, D., C.L. Lehman, and K.T. Thomson. 1997. Plant diversity and ecosystem productivity: Theoretical considerations.
Proc. Natl. Acad. Sci., USA 94:1 857-1 870.
452
Journal of the Botanical Research Institute of Texas 7(1)
JOURNAL NOTICE
Douglas J. Futuyma, H. Bradley Shaffer, and Daniel Simberloff, eds. 2012 (Dec). Annual Review of Ecology,
Evolution, and Systematics, Volume 43. (ISSN: 1543-592X; ISBN: 978-0-8243-1443-9, hbk). Annual
Reviews, Inc., 4139 El Camino Way, P.O. Box 10139, Palo Alto, California 94303, U.S.A. (Orders: www.
AnnualReviews.org, service@annualreviews.org, 1-800-523-8635, 1-650-493-4400). $89.00 indiv, 485
pp.,7%'-x9y8".
Contents of Volume 43;
1 . Scaling Up in Ecology: Mechanistic Approaches — Mark Denny and Usandro Benedetti-Cecchi
2. Adaptive Genetic Variation on the Landscape: Methods and Cases — Sean D. Schoville, Aurelie Bonin, Olivier Francois,
SUphane Lobreaux, Christelle Melodelima, and Stephanie Manel
Endogenous Plant Cell Wall Digestion: A Key Mechanism in Insect Evolution — Nancy Calderdn-Cortis, Mauricio Quc-
sada, Hirofumi Watanabe, Horacio Cano-Camacho, and Ken Oyama
New Insights into Pelagic Migrations: Implications for Ecology and Conservation— Daniel P. Costa, Greg A. Breed, and
PatricfeW. Robinson
The Biogeography of Marine Invertebrate Life Histories— Dustin J. Marshall, Patrick J. Krug, Elena K. Kupriyanova, Ma-
ria Byrne, and Richard B. Emlet
Mutation Load: The Fitness of Individuals in Populations Where Deleterious Alleles Are Abundant— Aneil F. Agrawal
and Michael C. Whitlock
From Animalcules to an Ecosystem: Application of Ecological Concepts to the Human Microbiome — Noah Fierer, Scott
Ferrenberg, Gilberto E. Flores, Antonio Gonzalez, Jordan Kucncman, Teresa Legg, Ryan C. Lynch, Daniel McDonald, Joseph
R. Mihaljevic, Sean P O’Neill, Matthew E. Rhodes, Sejin Song, and William A. Walters
Effects of Host Diversity on Infectious Disease— Richard S. Ostfeld and Felicia Keesing
Coextinction and Persistence of Dependent Species in a Changing World— Robert K. Colwell, Robert R. Dunn, and
:hes to Forecasting Species’ Responses to Cl
rough the Lens of Coexistence Theory— J. HilleRisLambers, P.B. Adler, W.S. Har-
NyeemaC. Harris
10. Functional and Phylogenetic Apprc
and Joel G. Kingsolver
1 1 . Rethinking Community Assembly i
pole,J.M. Levine, and M.M. Mayfield
12. The Role of Mountain Ranges in the Diversification of Birds-Jon FJeldsd, Rauri C.K. Bowie, and Carsten Rahbek
13. Evolutionary Inferences from Phylogenies: A Review of Methods— Brian C. O'Meara
14. A Guide to Sexual Selection Theory— Bram Kuijper, Ido Pen, and Franz J. Weissing
15. Ecoenzymatic Stoichiometry and Ecological Theory— Robert L. Sinsabaugh and Jennifer f Follstad Shah
16. Origins of New Genes and Evolution of Their Novel Functions-Yun Ding, Qi Zhou, and Wen Wang
17. Climate Change, Aboveground-Belowground Interactions, and Species’ Range Shifts— Wim H. Van derPutten
m — Noah T. Ashley, Zachary M. Weil, and Randy J Nelsoi
a Ecology Group (PEG) Model: Mechanisms Driving Plankton Succession— Ulri,
Adrian, Lisette De Senerpont Domis, James f Elser, Ursula Gaedke, Bos Ibelings, Erik levvesen A
Molinero, Wolf M. Mooij, Ellen van Donk, and Monika Winder
. Global Introductions of Crayfishes: Evaluating the Impact of Species Invasions on Ecosyster
Lodge, Andrew Deines. Francesca Gherardi, Darren CJ. Yeo, Tracy Arcella, Ashley K. Baldridge, 1
r r„rr.,.. r Crysta A. Gantz, Geoffrey W. Howard, Christopher L. Jerde, Brett W. Peters, Jody ^
- - - -ner, Marion E. Wittmann, and Yiwen Zeng
Cumulative ln<
n. Inst Texas 7(1): 452. 2013
A BASELINE VASCULAR PLANT SURVEY
FOR OCMULGEE NATIONAL MONUMENT,
BIBB COUNTY, MACON, GEORGIA
Wendy B. Zomlefer and David E. Giannasi
Department of Plant Biology
John B. Nelson
Moore Herbarium
Department of Biological Sciences
University of South Carolina
Columbia, South Carolina 29208-0001, U.S.A.
terra incognita
125 South Edisto Avenue
Columbia, South Carolina 29205-3301, U.S.A.
ABSTRACT
INTRODUCTION
Study Area
The -fall line" in Georgia is a pnrminent geological boundary ca. 32 km (20 mi) wide extending from Augusta
southwest to Columbus (Fig. lA) and represents the Mesozoic shoreline of the Atlantic Ocean. The ancient
beach demarcates the flat and sandy upper Coastal Plain to the south from the rolling rocky hills of the Pied-
mont to the north and separates significantly different plant and animal communities. As a consequence, the
diverse habitats within this narrow transition zone are characterized by a rich variety of flora and fauna
(Wharton 1978). Settlements developed where rivers descend along the relatively steep slope of the fall line
forming rapids or waterfalls that were used to generate power, and provided natural stopping points for travel
I- Bot. Res. Inst. Tenas 7(1): 453 - 473. 2013
454
Journal of the Botanical Research Institute of Texas 7(1)
ill line of the Ocmulgee River. The river,
),” provides the principal
and commerce. An example is the city of Macon, founded at
whose Native American name (Okmulgee) translates as -where the water boils i
watershed for much of the Piedmont and Coastal Plain of central Georgia (Fig 1 A)
Ocmnlgee National Monument (OCMU), administered by the National Park Service (NFS, US. Depan-
ment of the Imertor), is located along the Ocmulgee River in eastern Macon (Bibb County) at the fall line (NFS
2012a). The parkeompnses 283.9 ha (701.5 acres) in twoseparate land tracts (Figs. IBand 2): the much largn
ZhTh r'' •>’'= ^">^11 d«ached pared ca. 3.2 km S "■«
earth' ^ ^ acres). OCMU has signiBcant archeological sites including
earthen mounds associated with two Native American Mississippian cultures that settled along the river prior
to European contact (Hally 1994;see-BriefHistory of OCMU With Emphasison Land Use-below).Dueto the
TLIn™raT^;^e“(m::r2Cr“‘°^^
TOui'^rr^an m ** ^ '"■“W summers and mild winters (National Weather Service
ju“ h™r33
p sses through its southwestern border, parallel to the Ocmulgee Rivet. The ca. 9 km (5.5
of hiking trails traverse upland forests and fields and pass along the swamps and open wetlands associated
with the river and its tributary. Walnut Creek.
The most prominent landscape features at the Ocmulgee Mounds site are the seven rectangular earthen
mounds that rise up to 17 m (55 ft; Greater Temple Mound. Fig. 3) above the relatively flat topography of 85-
119 m (280-390 ff Marsh 1986). Other historic resources date from early European settlement to more mo -
em times, includingalateseventeenthcenturyBritishfortandtradingpostsite,aavilWargunemptont
andarestored antebellum residence (Dunlap House). The Visitor Centerincludesamuseum with exhtbitsand
a major archeological collection with emphasis on the Early Mississlppian culture that
park par^l from 900 to 1100 AD. The parks natumland cultural resourcesattractaboutm,000vus, tots per
year (NFS 2012b). ^ ^
Lamar Mounds unit.-This isolated part of OCMU (Lamar Mounds and Village site; coordinates
32.812562°, -83.591580°) is located between the Ocmulgee River and Interstate 16, ca. 5 km (3 mi) southeast o
456
Journal of the Botanical Research Institute of Texas 7(1)
downtown Macon (Fig. IB). The property is in swampland slightly elevated from the surrounding Hoodplam
(ca. 85 m, 280 ft) and an oxbow lake (Black Lake) nearby to the east (Fig. 2B; Wheeler 2007). An unfinished
levee, constructed in the late 1930s to protect the site from flooding, borders the northern, eastern, and part of
the southeastern sides of the property. The unit is the type site of the Lamar culture (Late Mississippia"’
1350-1600 AD; Williams 1999) and comprises a palisaded village area (8.7 ha, 21.5 acres) surrounding two
large earthen mounds: a rectangular (pyramidal) mound (Mound A in Figs. 2B and 4), 10.7 m (35 ft) tall, and
an unusual circular mound with a spiral ramp (Mound B, 6.1 m [20 ft] tall). Each mound is enclosed by a
locked chain-link fence. The Lamar Mounds unit has difficult access via a county road and an overgrown foot
trail and is open to the public via guided tours on a very limited basis.
Brief History of OCMU with Emphasis on Land Use
Based on archeological evidence, Ocmulgee National Monument was occupied continuously by a series of na-
tive North American tribes for ca. 1 2,000 years before European contact in the mid-seventeenth century (su»-
maries in Pope 1956; Marsh 1986; Hally 1998; NPS 2005; Wheeler 2007; Williams 2008). The mostsignific^
cultural period began when the early Mississippians (900-1100 AD) migrated to the area from the cent
Mississippi River Valley. This sophisticated and stratified society built a town (Ocmulgee Fields) supported
an agncultural economy managed by master farmers along the Ocmulgee River bottomlands. Their cerefflO'
nial complex included several structures that dominate the landscape today, including a circular earth lodge
and seven massive flat-topped pyramidal earthworks that served as temple platforms and hunal mounds (Figs.
2Aand3).
By 1350 AD, the Ocmulgee Fields declined as a ceremonial center, and a new culture, the Lamar or ^te
Mississippians, coalesced among people who lived in the swamps ca. 3.2 km (2 mi) downstream. One of then
major centers is now protected as the Lamar Mounds and Village site (Figs. IB, 2B, and 4). Represented by
vestiges of a stockade surrounding two temple mounds, this site includes a unique circular mound with a spira
ramp (Figs. 2B and 4). The spread of introduced European disease led to eventual decline in the^pulation,
and by 1650 the remnants of the culture had relocated westward to the Chattahoochee River (Williams &
Shapiro 1990). , , ,
Descendants of these Lamar (designated “Creek” or “Muscogee” by the European settlers) returned to
Ocmulgee Field, in 1690 to re-establi,h the town (“Okmulgee W) and to trade with the Bnttsh who had
built a trading post and fort near the sacred mounds (Fig. 2A). The Creeks were later (1717) expelled from the
area after losing a war with the British over land rights. After the Revolutionary War. the new state of Geo^m
obtained concessions to Creek tribal lands in 1826, following a series of contentious treaties (see Pope 1956;
Wheeler 2007). The acreage was incorporated into the new city of Macon, established in 18 ^
For the next century, agriculture and industry accelerated the progressive degradation of the namml and
cultural resources of the site (Froeschauer 1989; Wheeler 2007). The mam park area was a large
the mid-1850s. Grazing removed understory vegetation in the forested areas of Walnut Cree , ai
ricultural production around the mounds eroded topsoil that accumulated downriver at the Lamar site. Ik
Central of Georgia Railroad constructed two railway lines though “Ocmulgee Old Fields,” clearing vegetation
and removing much of Lesser Temple Mound and Funeral Mound in the process. After the Civil War, a brick
factory, fertilizer plant, and dairy farm were in operation at the site, and clay was mined from deep pits near
Great Temple Mound. By the 1920’s, the Ocmulgee mounds had become a popular recreation area for Macon
residents, and activities such as motorcycle racing (on the slopes) further eroded the earthworks. A large por-
tion of McDougal Mound was used as fill for constructing Emery Highway.
A group of concerned local citizens and politicians successfully secured New Deal funding for archeologi-
cal studies, organized by the Smithsonian Institution. The massive excavations, conducted from 1933-1942b)’
the Civilian Conservation Corps (CCC), stripped most of the vegetation at both sites (e.g., Fig. 4) to reveal
prehistoric landscapes and invaluable artifacts. The success of the federally sponsored study in addition to
by Presidential Proclamation in 1936 (Marsh 1986; Wheeler 2007). Under the jurisdiction of the National Park
Service, CCC workers constructed roads, trails, utilities, parking lots, and the Visitor Center for the new park
Loblolly pine (Pinus taeda) and Bermuda grass (Cynodon dactylon) were planted to control erosion.
More recently, natural resources have been heavily impacted by encroaching urbanization, particular!)
the construction of the Macon Levee (in 1950) on the western side of the river and of Interstate 16 (in the 1960s;
Fig. IB), which extensively changed the hydrology at both park units (Burkholder et al. 2010). One significant
consequence was a massive flood in 1994 that inundated hardwood forest and created the Walnut Creek wet-
lands, now a permanently flooded area (Figs. 2A, 3, and 5). Currently, the rest of the main park consists of
forested “natural” zones surrounding mowed lawns protecting the mound areas. The Lamar Mounds unit is
covered by dense floodplain forest that is managed on a minimal maintenance schedule with trail and mounds
occasionally cleared of vegetation. The NFS continues to negotiate for acquisition of up to 14,000 acres of Oc-
mulgee Old Fields eligible for the National Register as Traditional Cultural Property due to the significance to
the Creek people (David 2012).
Horistic Surveys of OCMU
A few unvouchered lists of plant species for the Ocmulgee National Monument and surrounding areas had
been produced for various park reports (e.g., Froeschauer 1989; Puckett 1997). In 1999, the National Park Ser-
vice inaugurated a long-term ecological inventory and monitoring program to establish baseline data onpaik
ecosystems for resource management decisions (see Fancy et al. 2009). As part of that initiative in the south-
eastern United States (DeVivo et al. 2008), a comprehensive and vouchered floristic survey of OCMU was
conducted by Gaddy and Nelson (2004). They collected 447 specimens from July 2002 to November 2004 and
also obtained 31 vouchers from informal 1986-1987 surveys by John D. Shepherd and his students at Mercer
University in Macon, Georgia. The most recent specimen verifications and supplemental collections m 20^
2009 by Zomlefer and associates enhance previous surveys with the goal of generating an accurate vouche
species list as a reference for on-going invasive species control and park-wide vegetation community mapping
and research projects (e.g., DeVivo et al. 2011).
The first author led 4 field trips for one year (27 May, 25 July, and 1 October 2008; 9 April 2009; ZontkJ^
2091-2136, 2249-2268, 2275-2292, 2359-2403) to collect plant specimen vouchers using standard fielda^
herbarium techniques (under collecting permit # OCMU-2008-SCI-0005). One follow-up trip was conduct
on 7 August 2012 to confirm vegetation community types. The GA Herbarium team also verified detennii^
tions of vouchers from previous surveys of the park (Nelson, Gaddy, and Shepherd collections),
unidentified specimens as well as a voucher of Salpichroa origanifolia sent to GA for identification m Nove
2010 by Theresa L. Hall, Interpretive Ranger at Ocmulgee National Monument. The complete set of®®""
vouchers for the park (610 specimens) is deposited at GA Herbarium. The floras of Radford et al (1968)^
Weakley (2011) were primary sources for plant identification, supplemented by Cronquist (1980),
Zomlefer et al, Floristic survey of Ocmulgee National I
459
RESULTS
Floristics
The “Annotated Checklist of Vascular Plant Tax
represented by vouchers in GA Herbarium. The (
I for Ocmulgee National Monument
ed 131 new numbered collections of
vascular plants representing 123 species; of these, 58 species are new vouchered taxa for the park (underlind
in the “Annotated List”). Added to the existing OCMU collections, the tally is 436 species (610 specimens) now
vouchered for this flora in three major plant groups: monilophy tes (1 1 spp.), gymnosperms (2 spp.), and angio-
sperms (423 spp.). Included in this total are three species in cultivation (indicated as “cult” in the annotated
list) or persisting from cultivation hut likely not naturalized. The most numerous families are: Poaceae (53
spp.), Asteraceae (36 spp.), Cyperaceae (33 spp.), Fabaceae (21 spp.), Rosaceae (18 spp.), Fagaceae (11 spp.),
Lamiaceae (10 spp.), Rubiaceae (9 spp.), Polygonaceae (8 spp.), Convolvulaceae (7 spp.), Onagraceae (7 spp.),
Plantaginaceae (7 spp.), and Sapindaceae (7 spp.).
Appendix 1 is a compilation of 189 additional unvouchered species cited for Ocmulgee National Monu-
ment in NPS databases and unpublished reports (Froeschauer 1989; Puckett 1997; Burkholder et al, 2010; NPS
2012c). Without preserved specimens, verification of these reports is not possible; therefore, this list should be
referenced with caution. Based on our previous national park survey work using unvouchered lists (e.g., Zom-
lefer et al. 2008, 2012), we predict that at least 20 percent of this list likely represents misidentihcations.
In general, the flora of Ocmulgee National Monument comprises a representation of the basic habitat
types of central Georgia (see “Plant Communities” section below) and also weedy plants common in disturbed
areas of the upper Coastal Plain and lower Piedmont (Zomlefer et al. 2008, 2012). The flora does not include
any Georgia endemics or species ranked as state or federally protected (threatened, endangered, or rare; GA
DNR 2010; USFWS 2012). However, Cayaponia quinqueloba is listed as a Special Concern species (at risk but
not formally protected) by the GA DNR (2011) with a state rank of S2 (imperiled; 6-20 occurrences) and a
global rank of G4 (apparently secure globally; no immediate conservation concern). This species was vouch-
ered from the western margin of Walnut Creek wetlands (Nelson 23064; see Fig. 2A) and from the apex of
Mound A at the Lamar Mounds unit (Zomlefer 2259; see Figs. 2B and 4).
One-hundred and six species at OCMU are non-native (indicated with an asterisk [*] in the “Annotat
List”). Excluding the three cultivated exotics, the remaining 103 introduced species represent 23.6 percent
the flora. Thirty-seven of the non-native species are invasive (Table 1) and ranked in four categories by Geoij^
Exotic Pest Plant Council (GA-EPPC 2006) depending on perceived threat to the native flora: Category 1 (13
spp.; serious invasive extensively invading plant communities, displacing native species); Category 2 (9spp.
moderate invasive, invading and displacing to a lesser degree than Category 1 species); Category 3 (8s^
minor invasive or potential threat not yet known, a threat in adjacent states); and Category 4 (8 spp.;
ized in Georgia, generally not posing a threat but additional data needed). Most of these species were v
ered from disturbed areas of the park (indicated as “da” in Table 1). ,
Currently, about 40 ha (100 acres) of OCMU have been successfully surveyed, treated, and manage ^
invasive plant species (G. LaChine, pers. comm.). For the past ten years, park personnel have been targe mg
most prevalent and critical of these exotics (all ranked as Category 1): Ailanthus altissima,
Melia azedarach, and Pueraria montana. A serious invasive of immediate concern and removal ef orts,
sebifera, has become widespread in the Walnut Creek wetlands area of the main park unit and is also pre^^^^
the Lamar Mounds unit. Other exotics, such as Lonicerajaponica and Hedera helix, have been
ever possible, especially from sensitive areas with cultural resources (NPS 2012d). In s
lannia keisak (1), and Myrii
„(2)arebe-
non-native grass species (Wheeler 2007; G. LaChine, p
3 (Table 1). In the main park unit, these exotics primarily occur intermixed with
Cynodon dactylon comprising the mowed lawns covering the mound
eight of these are ranked as
n^tivp grasses and pi^
Ocmulgee National Monument has a long history of disturbance that has drastically altered Q^jj,yigee
pecially over the last two centuries (see “Brief History of OCMU with Emphasis on Land Use )• ^ or
he vegetation.^
XheOcinnr
andfores-ea*
ip with some open disturbed areas h
Zomleferetal., Floristk survey of Ocmulgee National Monument
6A-EPPC (2006): Category 1 (serious invasive), Category 2 (moderate invasive). Category 3 (minor invasive or potential threat not yet known), and Category 4
the overgrown logging road andcleared mound areas (Fig. 2B). Below isasummary of the flora of these general
conrmunity .ypes based on Wharton (1978), Froeschauer (1989), specimen data, and field observations (W.B.
Zomlefer & B.L. Wichmann, pers. obs.): upland foresl (mixed hardwood and mixed hardwood-pine), swamp
btest, open wetland, and disturbed areas. These categories, which may tntergrade, are provided as a gutde (or
placing plant species within the context of a habitat.
UpUnd Forests-Ihe forested uplands at the Ocmulgee Mounds unit (Fig 5) have been mrrfrlied by
disturbance into secondary growth hardwood and mixed hardwood-pine (Wharton 1978; Burkholder et al.
2010). The dominants are oaks (e.g.. euercrts/ulcmu, 8 aigm. Q. phellos), hickories (e.g, Cuo'u gluhru, C. ovrrtu),
and loblolly pine (Pinus toedu); a few areas along the southeastern section of Ocmulgee National Moiiumenl
Road are dominated by pine. Other codominant trees include Acer spp. (e.g., A.iloridunum, A. ruhrrmr), Fugus
462
Journal of the Botanical Research Institute of Texas 7(1)
grandifolia, Liquidambar styraciflua, Liriodendron tulipifera, Magnolia grandiflora, and Ulmus Americana. The
composition of the understory shrubs and shrubby trees vary according to the overstory and location in the
park and commonly include Aesculus pavia, Asimina parviflora, Calycanthm floridus, Carpinus carolinim
Celtis laevigata, Cercis canadensis, Comusflorida, Crataegus spp. (e.g., C. spathulatd), Diospyros virginiona, Jk
spp. (e.g., I. decidua), Ligustrum sinense, Prunus serotina, and Vaccinium arhoreum. Gelsemium sempervirm,
Hedera helix, Lonicera japonica, Smilax spp. (e.g., S. bona-nox). Toxicodendron radicans, and Vitis spp. (e.g.,V
rotundifolia) comprise some frequent vine species intertwined amongst the shrubs and over the ground cover.
Understory vegetation is much more diverse in the mixed hardwood forests than in those areas where
pine predominates. The early spring flora species, inhabiting the forest floor before the hardwood canopy fully
closes, include: Erythronium umbilicatum, Hexastylis arifolia, Luzula echinata, Melica mutica, Myosotis macro-
sperma. Podophyllum peltatum, Polygonatum biflorum, and Viola sororia. Woodland species such as Aspleniim
platyneuron, Conoclinium coelestinum, Dichanthelium commutatum, Elephantopus carolinianus, Oplismenuskr-
tellus, Symphyotrichum spp. (e.g., S. pilosum), and Tipularia discolor appear later in the season, especially under
canopy openings such as trail sides.
Floodplain Forest.— Swamp or bottomland hardwood (floodplain, alluvial, or palustrine forest) consists
of seasonally inundated, mesic lowland that generally remains moist throughout the year (Wharton 1978). At
the Ocmulgee Mounds unit, this community type is associated with the Ocmulgee River and Walnut Creek in
the southwestern and eastern sections of the park, mainly bordering open marshland (Figs. 2A and 5). The
low-lying Lamar Mounds unit is almost entirely covered by dense floodplain forest (Williams 1999) since the
partial levee (Fig. 2B) provides little protection from periodic flooding by the nearby Ocmulgee River (Fig. IB).
Within the swamp forest, tall trees create a closed-canopy over an often impenetrable thicket of shrubby
understory. Dominant trees usually include Acer spp. (e.g., A. negundo, A. rubrum), Betula nigra, Fraxinuspenn-
sylvanica, Liquidambar styraciflua, Liriodendron tulipifera, Nyssa biflora, Platanus occidentalis, Quercus spp.
Q. nigra), and Ulmus americana. Swamp communities at OCMU can also be characterized by the following co-
dominants and/or understory woody species: Alnus serrulata, Asimina parviflora, Bignonia capreolata, Carpims
caroliniana, Crataegus viridis, Itea virginica. Hex spp. (e.g., I. vomitoria), Ligustrum sinense, Lonicera japonica,
Ostryavirginiana, Pinus taeda, Salix caroliniana, Smilax spp. (e.g., S. laurifolia), Triadica sebifera, Toxicodendron
radicans, and Vitis spp. (e.g., V rotundifolia). A relatively sparse understory of ferns, such as Onoclea sensiHis
and Woodwardia areolata, and angiosperm species, such as Arundinaria gigantea, Arisaema triphyllum, Com-
melina virginica, Juncus coriaceus,Justicia ovata, Leersia lenticularis. Lobelia cardinalis, Lycopus rubellus, and Pi-
lea pumila, may occur under openings in the bottomland canopy.
Open Wetlands. — Open wetland (or marsh) is a broad designation, here applied to aquatic areas at the
Ocmulgee Mounds unit where the canopy is lacking or sparse and standing water is often present for at least
part of the year. This habitat type is mainly associated with floodplain areas of the Ocmulgee River and its
tributary. Walnut Creek (Fig. 5). The l^rge marsh. Walnut Creek wetlands (Figs. 2A, 3, and 5), developed frow
hydrological changes caused by the construction of 1-16 (Wheeler 2007; Burkholder et al. 2010). Prior to 1994.
this open wetland was forested and seasonally flooded but the area now contains standing water all year.
In the marshy areas of this park unit, sedges and grasses predominate, including Carex spp. (e.g > ^
sianica, C. lupulina), Cyperus spp. (e.g., C. erythrorhizos, C. retrorsus), Leersia virginica, Panicum anceps, P- ngt^
lum, Phanopyrum gymnocarpon, Rhynchospora globularis, and Scirpus cyperinus. Floating aquatics indudeAf
temanthera philoxeroides. Myriophyllum aquaticum, and Najas minor. Other common herbaceous species
Erechtites hieraciifolius, Galium triflorum, Hydrocotyle verticillata. Iris hexagona, Juncus dichotomus, Ludvnpi^ j
spp. (e.g., L. decurrens), Mikania scandens, Mimulus alatus, Murdannia keisak, Packera glabella, Persicarid spP- i
(e.g.. P. hydropiperoides), Sagittaria latifolia, Saururus cernuus, and Typha latifolia. Berchemia scandens, Brun- ^
nichia ovata, Cephalanthus occidentalis, Cornus stricta, Forestiera acuminata, Hibiscus laevis, Rubus pensiha^ |
cus, Sabal minor, Sambucus canadensis, Smilax spp. (e.g., S. glauca), and Vitis spp. (e.g., V aestivalis) are |
of common woody vines and scattered shrubs that occur along the margins of these open wetlands. The ^
dering areas may also comprise some tree species (such as Acer floridanum) and seedlings of Triadica selnjera-
463
Disturbed or Ruderal Areas.— This general category refers to cleared sections around public access ar-
eas (parking lots, roadsides, trails, railroad right-of-way) and mowed fields surrounding earthworks and other
historic sites at the Ocmulgee Mounds unit (see Fig. 5). Disturbed areas in the Lamar Mounds swamp are re-
stricted to the most exposed sites (cleared mounds and trail margins) and are characterized by species prefer-
ring damp habitat such as Eupatorium sewtinum, Hypericum mutilum, Perillafrutescens, Persicaria longiseta,
and Smilax spp. (e.g., S. rotundifolia).
In the main park, ruderal areas are characterized by a much more variable and diverse flora that includes
many non-native and invasive species and a predominance of graminoids, including: Andropogon spp. (e.g., A.
glomeratus), Bromus racemosus, Cynodon dactylon, Dactylis glomerata, Danthonia spicata, Dichanthelium spp.
(e.g., D. dichotomum), Echinochloa crusgalli, Eragrostis spp. (e.g., E. curvula), Festuca subverticillata, Lolium pe-
renne, Panicum spp. (e.g., P. virgatum), Paspalum spp. (e.g., P. dilatatum), Setaha parvijlora, Sorghastrum nutans.
Sorghum halepense, and Vulpia myuros. Asteraceae also comprise a large component, with common species
such as Ambrosia artemisiifolia, Erigeron strigosus, Eupatorium hyssopifolium, Gamochaeta spp. (e.g., G. pensylva-
nica), Heknium amarum, Hypochaeris radicata, Krigia virginica, Pityopsis graminifolia, Solidago altissima, Son-
chus oleraceus, and Youngiajaponica. Allium canadense, Chaerophyllum tainturieri, Glandularia pulchella, Hous-
tonia pusilla, Ipomoea spp. (e.g., I. cordatotriloba), Lamium ampkxicauk, Lespedeza cuneata, Plantago spp. (e.g., P
lanceolata), Polypremum procumbens, Stellaria media, Trifolium arvense, Vida spp. (e.g., V. sativa), and Viola ar-
vensis are examples of other widespread ruderal herbs. The borders of these areas often include woody specif
rotundifolia).
OCMU Manage».^..^ r
Ocmulgee National Monument is a historically significant park that also supports diverse wiidlite as part oi a
greenway corridor to other natural areas to the south (DeVivo et al. 2008). The biological and cultural resourc-
es of the park have been heavily impacted by surrounding urbanization and associated degradation of the Oc-
mulgee River watershed. Threats affecting the flora and fauna include air and water pollution^CTosion, sedi-
a baseline for monitoring these habitats, thus providing a valuable reference for natural resource management.
Encroaching development threatens the native component of the OCMU flora while contributing to an in-
crease in non-native and weedy species that require documentation. Continued inventory efforts should also
concentrate on locating significant species-both native and exotic-previously reported in the park but not
yet verified by voucher specimens (see Appendix 1).
annotated checklist of vascular plant taxa
This list of 436 vascular plant species, representing 112 families, represents collections from Ocmulgee Na-
tional Monument now deposited at the GA herbarium. Genera, species, and infraspecific taxa are alphabetical
within each family under three major groups (monilophytes [“ferns and allies”], gymnosperms, and angio-
sperms). Family circumscriptions follow the following sources: Smith et al. (2006) for monilophytes; FNA
(1993) for gymnosperms; and APG HI (2009) for angiosperms. Scientific nomenclature follows ITIS (2012) and
Weakley (2011). Common names are from Weakley (2011) when available, or from Wunderhn and Hansen
(2011).ThefewsynonymsincludedinbracketsarealtematenamesusedinNPSreportsanddatabases.
Specimen notations: G = Gaddy (all s.n.) [GA accession number], H = Julie Howard (one specimen, Salpi-
chroa origanifolia), N = Nelson, S = John D. Shepherd, Z = Zomlefer; underlined ta.M = new vouchered taxa for
the park by Zomlefer * = exotic (ITIS 2012; Weakley 2011); invasive exotics (GA-EPPC 2006): [Cat 1] - Cate-
gory 1, [Cat 2] = Category 2, [Cat 3] = Category 3, [Cat 4] = Category 4; rare plants. [Con] = species of special
concern in Georgia (GA-DNR 2011); cult = cultivated (i.e., planted on park grounds); (s) = sterile (non-repro-
ductive) specimen. Locality/habitat data: LM = Lamar Mounds, da = disturbed areas, fp - floodplain swamp, pm
= pond/marsh (open water), uf = upland forest, hab? = habitat data not recorded on specimen voucher label.
Relative abundance: c = common (generally abundant throughout a particular habitat; species easily found), o
= occasional (locally common and/or several individuals distributed within a habitat; species not too difficult
to locate), i = infrequent (sporadic occurrence of a small number of individuals; species relatively scarce and
not easily found), r = rare (very few individuals encountered), ab? = abundance data not recorded on voucher
specimen label.
Zomleferetal.Jloristics
467
471
ACKNOWLEDGMENTS
The first author thanks Kristian D. Jones, Lisa Krueger, Patrick Lynch, and Jeremy Rentsch for their enthusias-
tic assistance in the field; Lisa Krueger also prepared specimen labels and identified 17 grass and sedge speci-
mens. Laura E. Lukas, (former) Collections Manager at GA, processed and databased the Gaddy, Nelson, and
Shepherd shipments, identified 70 of those specimens, and supervised the GA Herbarium team of plant mount-
ers. Brenda L. Wichmann, (former) Collections Manager at GA, facilitated verification of general vegetation
community composition in a follow-up field trip to OCMU (7 August 2012), and Kaitlin T. Brotman assisted
with preparation of Figure 1. We also greatly appreciate the invaluable support and cooperation of National
Park Service personnel: Guy LaChine, Chief of Operations at Ocmulgee National Monument; Joe DeVivo, Net-
work Coordinator; and Christina Wright, Data Manager. Biological Technicians Allen Huckabee, Stephen
Friedt, and Tel Vaughn provided transportation to the Lamar Mounds for WBZ and field associates. J. Richard
Carter, Curtis J. Hansen, Walter S. Judd, and Melanie Link-Perez provided constructive criticisms of the manu-
script. The fulfillment of this vouchered baseline survey was funded by National Park Service contract agree-
ments J2114-07-0026 andJ2114-09-0017 (WB. Zomlefer, PI, and D.E. Giannasi, coPl).
472 Journal of the Botanical Research Institute ofTexa57(l)
REFERENCES
Angiosperm PHYLCX3ENY Group (APG III). 2009. An update of the Angiosperm Phylogeny Group classification for the orders
and families of flowering plants: APG III. BotJ. Linn. Soc. 161:105-121.
Burkholder, J.M., E.H. Allen, and CE. Kinder. 201 0. Assessment of water resources and watershed conditions in Ocmulgee
National Monument, Georgia. Natural Resource Report NPS/SECN/NRR-201 0/276. National Park Service, Natural
Resources Program Center, Fort Collins, CO. http://nature.nps.gOv/publications/nrpm/nrr.cfm#2010. Accessed 15
August 201 2.
Cronquist, a. 1 980. Vascular flora of the southeastern United States, volume 1 . Asteraceae. University of North Carolina
Press, Chapel Hill, NC.
David, J. 2012. Ocmulgee National Monument. Old Fields boundary study & environmental assessment [brochure].
Ocmulgee National Monument, National Park Service, Macon, GA.
DeVivo, J.C., C. Wright, M. Byrne, E. DiDonato, and T. Curtis. 2008. Vital signs monitoring in the Southeast Coast Inven-
tory and Monitoring Network. Natural Resource Report NPS/SECN/NRR-2008/061. National Park Service, National
Resource Program Center, Fort Collins, CO. http://nature.nps.g0v/publications/nrpm/nrr.cfm#20O8. Accessed 15
August 201 2.
DeVivo, J.C., M.B. Gregory, T. Curtis, M.W. Byrne, and C. Wright. 2011. Southeast Coast Network climate science strategy.
Natural Resource Report NPS/SECN/NRR-201 1/436. National Park Service, National Resource Stewardship and Sci-
ence, Fort Collins, CO. http://nature.nps.g0v/publications/nrpm/nrr.cfm#2O1 1 . Accessed 1 5 August 201 2.
Fancy, S.G., J.E. Gross, and S.L Carter. 2009. Monitoring the condition of natural resources in US national parks. Environin.
Monit. Assessm. 151:161-174.
Flora of North America (FNA) Editorial Committee, eds. 1993-I-. Flora of North America north of Mexico. 16-F vols. Oxford
University Press, New York and Oxford.
Flora of North America (FNA) Editorial Committee, eds. 1993. Flora of North America north of Mexico, volume 2. Pterido-
phytes and gymnosperms. Oxford University Press, New York and Oxford.
Froeschauer, P. 1 989. A vegetation history of Ocmulgee National Monument, Macon, Georgia. CPSUTechnical Report No.
51 . National Park Service Cooperative Unit, Institute of Ecology, University of Georgia, Athens, GA.
Gaddy, LL and J.B. Nelson. 2004. Vascular plant inventory of the Ocmulgee National Monument, Bibb County, Georgia.
[Unpublished report.] National Park Service, Southeastern Regional Office, Athens, GA.
Georgia Department of Natural Resources (GA-DNR). 201 0. Protected plants of Georgia. Georgia Wildlife Resources Division.
Social Circle, GA. http://georgiawildlife.com/sites/default/files/uploads/wildlife/nongame/text/html/protected.
species/Plants.html. Accessed 15 August 2012.
Georgia Department of Natural Resources (GA-DNR). 201 1. Tracking list of special concern plants of Georgia. Nongame
Conservation Section, Social Circle, GA. http://www.georgiawildlife.com/sites/default/files/uploads/wildlife/nor'
game/pdf/rare_species_data/Tracking_List_of_SpeciaLConcern_Plants_of_Georgia.pdf. Accessed 1 5 August 2012.
Georgia Exonc Pest Plant Council (GA-EPPC). 2006. List of non-native invasive plants in Georgia. http://www.gaeppcorg/
list.cfm. Accessed 1 5 August 201 2.
Godfrey, R.K. 1988. Trees, shrubs and woody vines of northern Florida and adjacent Georgia and Alabama. University of
Georgia Press, Athens, GA.
Godfrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of the southeastern United States. Monocotyledons
University of Georgia Press, Athens, GA.
Godfrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of the southeastern United States. Dicotyledons. Univer-
sity of Georgia Press, Athens, GA.
Hally, DJ., eo. 1994. Ocmulgee archaeology 1936-1986. University of Georgia Press, Athens, GA.
Hally, DJ. 1998. Ocmulgee site. In: G. Gibbon, ed. Archaeology of prehistoric nativ
Publishing, New York, NY. Pp. 600-601 .
Isely, D. 1 990. Vascular flora of the southeastern United States, volume 3, part 2. Leguminosae (Fabaceae). Universrty
North Carolina Press, Chapel Hill, NC.
ms Steering Committee (ITIS). 2012. Integrated taxonomic information system, http://www.itis.gov/index.html. Ac-
cessed 15 August 2012.
Marsh, A. 1986. Ocmulgee National Monument: an administrative history. National Park Service,
the Interior, Washington, DC. http://www.nps.gov/history/history/online_bi
,: an encyclopedia. Garland
National Park Service (NPS). 2005. Ocmulgee National N
ment Printing Office, Washington, D.C.
National Park Service (NPS). 2012a. Ocmulgee National Monument, Georgia [official homepage], http://www.nps.gov/
ocmu/index.htm. Accessed 1 5 August 201 2.
National Park Service (NPS). 2012b. NPS stats. National Park Service Public Use Statistics Office, U.S. Department of the
interior, Washington, DC. httpy/nature.nps.gov/stats/index.cfm. Accessed 15 August 2012
National Park Service (NPS). 201 2c. NPSpecies, certified species lists. Certified species list for vascular plants in Ocmulgee
National Monument. National Park Service, Natural Resource Stewardship and Science, U.S. Department of the Inte-
rior, Washington, DC. https://irma.nps.gov/App/Species/Search. Accessed 15 August 2012.
National Park Service (NPS). 201 2d. Inventory & monitoring program status report, 201 2: Ocmulgee National Monument.
National Park Service, Southeast Coast Network, Athens, GA. http://www.nps.gov/ocmu/naturescience/upload/
OCMU_Program_Summary_2012-2.pdf. Accessed 15 August 2012.
National Weather Service. 2012. National Oceanic and Atmospheric Administration's Nationai Weather Service. U.S. Dept,
of Commerce, Silver Spring, MD. httpy/www.nws.noaa.gov/view/national.php?map-on. Accessed 15 August 2012.
Palmer, M.W. and J.C. Richardson. 201 2. Biodiversity data in the information age: Do 21st century floras make the grade?
Castanea 77:46-59.
Pope, G.D. 1956. Ocmulgee National Monument. Georgia. Nationai Park Service Historical Handbook Series No. 24. Na-
tional Park Service, U.S. Department of the Interior, Washington, DC. http://www.cr.nps.gov/history/online_books/
hh/24/hh24toc.htm. Accessed 15 August 2012.
Puckett, D.L 1997. Ocmulgee National Monument. A survey of natural resources. [Unpublished report.) Governor's
Intern Program, Ocmulgee National Monument, National Park Service, Macon, GA.
Radford, A.E., H.E. Ahles, and C.R. Bell. 1 968. Manual of the vascular flora of the Carolinas. University of North Carolina
Press, Chapel Hill, NC.
Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, and P.G. Wolf. 2006. A classification of extant ferns. Taxon
55:705-731.
United States Fish and Wildlife Service (USFWS). 2012. Species reports: listed plants. httpy/ecos.fws.gov/tess_public/pub/
listedPlants.jsp. Accessed 15 August 2012.
Weakley, A.S. 2011. Flora of the southern and mid-Atlantic states, working draft of 1 5 May 201 1 . httpi^/www.herbarium.
unc.edu/flora.htm. Accessed 1 5 August 201 2.
Wharton, C.H. 1978. The natural environments of Georgia. Bulletin 144. Georgia Department of Natural Resources,
Atlanta, GA.
Wheeler, BJ. 2007. Ocmulgee National Monument. Cultural landscape report. National Park Service, Southeast Regional
pdf. Accessed 1 5 August 201 2.
Williams, M. 1999. Lamar revisited. 1996 test excavations at the Lamar site. LAMAR Institute Publication 43, University
of Georgia, Athens, GA. http://shapiro.anthro.uga.edu/Lamar/images/PDFs/publication_43.pdf. Accessed 1 5 August
2012.
Williams, M. 2008. Ocmulgee mounds. The new Georgia encyclopedia (on-line), Georgia Humanities Council and
University of Georgia Press, Atlanta and Athens, GA. httpy/www.georgiaencyclopedia.org/nge/Article.jsp?id=h-
2866&hl=y. Accessed 1 5 August 2012.
Williams, M. and G. Shapiro, eds. 1990. Lamar archaeology: Mississippian chiefdoms in the deep south. University of Ala-
bama Press, Tuscaloosa, AL.
Wunderlin, R.P. and B.F. Hansen. 2000. Flora of Florida, volume 1. Pteridophytes and gymnosperms. University Press of
Florida, Gainesville, FL.
Wunderlin, R.P. and B.F. Hansen. 2011. Guide to the vascular plants of Florida, ed. 3. University Press of Florida, Gainesville,
FL.
ZOMLEFER, W.B., D.E. Giannasi, K.A. Bettinger, S.L. Echols, and LM. Kruse. 2008. Vascular plant survey of Cumberland Island
National Seashore, Camden County, Georgia. Castanea 73:251-282.
ZoMLEFER WB DE Giannasi A REYNOLDS,ANDK.HEiMAN.2012.VascularplantfloraofChattahoocheeRiverNationalRecre-
3. Rhodora 114:50-102.
JOURNAL NOTICE
FLORA OF CALCAREOUS UPLAND GLADES IN GADSDEN
AND JACKSON COUNTIES, FLORIDA^
A.F. Johnson
W.W. Baker
Florida Natural Areas Inventory 1 422 Qrestview Ave.
W18 Thomasville Road, Suite 200-C Tallahassee, Florida 32303, U.S.A
Tallahassee, Florida 32303, U.S.A.
ajohnson@fnai.org
Florida State University
Tallahassee, Florida 32306, U.S.A.
anderson@bio.fsu.edu
A.K. Gholson, Jr.
P.O.Box 385
Chattahoochee, Florida 32324, U.S.A
ABSTRACT
INTRODUCTION
Florida upland glades are small (0.03 to 0.8 ha), largely herbaceous openings in an otherwise forested land-
scape, occurring on thin soils over calcareous substrates in dissected terrain in Jackson and Gadsden counties
(plus one just north of the border in Decatur County, Georgia). They form the southeastern outpost of a series
of calcareous prairies and glades in the southeastern U.S., stretching from eastern Texas north to southern
Ohio and east to Georgia (Fig. 1). These isolated prairies and glades surrounded by forest are dominated on the
deeper soils by some of the same grasses as the Midwestern tallgrass prairies (Schizachyrium scoparium, Andro-
pogon gerardii, Sorghastrum nutans and Panicum virgatum; Sims 1988). Areas of shallower soil over calcareous
substrates are characterized by the annual grass, Sporobolus vaginiflorus (Ware 2002; Lawless et al. 2006).
While Florida upland glades share none of the endemic species of these calcareous openings to the north and
west, they do have many species in common with them, including some of the same dominant grasses
’We dedicate this paper to Steven W. Leonard, who in 1982 brought attention to n
JBot. Res. Inst. Texas 7(1): 475
(Schizachynum scoparium, Muhlenbergia capiUaris, and Sporobolus vaginiflorus\ Baskin and Baskin 2003). How-
ever, the Florida glades are also unique in having as one of their most characteristic dominants, the sedge
Schoenus nigricans, which is not found at any of the other sites shown in Figure 1.
These graminoid-dominated openings on calcareous outcrops have traditionally been referred to as
“glades” in Florida (or “upland glades” to distinguish them from the Florida Everglades). Baskin andBaskm
(2000) and Lawless et al. (2004) suggest confining the term ’’limestone cedar glade” to sites dominated by an-
nual grasses, cryptogams, and winter annuals, while using “xeric limestone prairies” and “barrens” to «'
note areas dominated by perennial grasses, in part because the two types have different endemics and diffei®
implications for management. Limestone cedar glades are naturally open due to shallow substrates, where^
prairies and barrens require anthropogenic fire to maintain them in open condition. However, the
eas have about equal proportions of the dominant annual and perennial grasses cited by the above authors
in addition, a dominant perennial sedge whose relationship to soil depth has not been determined. In 1#"
Johnson et al.. Flora of calcareous upland glades
477
their uniqueness, we will refer to them here as “Florida upland glades” as distinct from “limestone cedar
glades” sensu Lawless et al. (2004). Florida upland glades are denoted as “panhandle Florida limestone prairie"
Florida upland glades were first recognized as a distinct community by Gholson (Ward & Gholson 1987)
who began collecting specimens from River Junction and East Bank Campground glades in the 1970’s (includ-
ing the first examples from glades of species that are rare in Florida, e g., Delphinium carolinianum, Ratibida
pinnata, and Symphyotrichum pratense). The idea of Florida upland glades as a distinct community was bol-
stered in 1982, when, as part of a systematic survey of habitats and rare species along the east side of the Apala-
chicola River, Leonard and Baker (1982) discovered five more upland glades in Gadsden County with four
species new to Florida (Asckpias viridiflora, Echinacea purpurea, Stachys crenata, and Symphyotrichum shortii)
and one new to the Apalachicola region (Callirhoe papaver). Continued collecting by Gholson, J.B. Nelson, R.K.
Godfrey, and others on these upland glades, as well as on some in Jackson County, resulted in the finding of
more rare species, range extensions, and species new to Florida (including Primula meadia, Salvia urticifolia,
and Sporobolus vaginiflorus; Anderson [1984, 1988, 1989, 2007]). Other publications reported the finding of
individual species new to Florida, including Bouteloua curtipendula (Nelson 1985), Lepuropetalon spalhulatum
(Ward & Gholson 1987), and Carex microdonta (Bridges et al. 1989) and included short descriptions of the
glades vegetation, with the result that the current list of known rare species from the Florida upland glades was
fairly complete by the end of the 1980s. In 1987 Nelson, Anderson, and Gholson compiled a preliminary spe-
cies list for the Gadsden County glades which was updated by Gholson in 1990. The present paper builds on
the preliminary list for the Gadsden County glades, adds species from the Jackson County glades, and places
the flora of the Florida upland glades in the context of other well-studied calcareous glades and prairies to the
north and west. It is hoped this publication will spur recognition of the biogeographical significance of the
Florida upland glades and lead to protection of at least some of the better preserved examples.
PHYSICAL SETTING
Geology
Florida upland glades occur in two clusters about 40 km apart. The cluster of eleven Gadsden County glades
(plus a twelfth glade just over the border in Georgia, which will hereafter be included with the Gadsden
County glades) are scattered within a 47 km^ area north, south, and southeast of the town of Chattahoochee
along the uplands bordering the eastern side of the Apalachicola River (Fig. 2). They occur at an elevation of
27.4-30.5 m above mean sea level (90-100 ft contour line) and are underlain by the Chattahoochee Formation
of lower Miocene age, a silty, sandy dolomite with occasional occurrences of limestone, which dips steeply
southward from a high point in the vicinity of Chattahoochee (Rupert 1990). The cluster of nine Jackson
County glades are scattered within a 10 km^ area west and northwest of the town of Marianna (Fig. 3). They
occur between the 140- and 150-ft contour lines and are underlain by Marianna Limestone of lower Oligocene
age, a white to light gray limestone 7.6-12.2 m thick which dips southward from a maximum elevation above
mean sea level near Marianna of 45.7 m (150 ft; Moore 1955). The Marianna is a commercial grade limestone
that is being actively mined in the vicinity of the glades.
Soils
Coultas (1983) sampled soils along transects across three glades in Gadsden County, with a total of 13 samples
from open glades and 6 samples from forested edges of glades. Depth to hard limestone on the herb-dominated
open glades ranged from 10 to 36 cm; on the forested edge of the glades it ranged from 53 to >152 cm. Soil tex-
ture on the open glades was generally silt loam in the upper layers, followed by several inches of soft limestone
grading to hard limestone. On the forested edges, sandy loam in the upper layers gave way to clay and then to
soft limestone followed by hard limestone. The open glades soils were more alkaline (pH 7.5) than the soils of
the surrounding forests (pH ranging from 5.3 to 6.4). Coultas concludes from the lack of mottling in the subsoil
of the open glades and the position of one of the sampled glades (#5 on Fig. 2) at the crest of a hfll that they are
not seepage areas and that woody invasion is probably controlled by the shallowness of the soils.
479
5.Moranz;16.Bennett,17.
Brooks-4; 20. Brooks-2;
Climate
The area experiences mild winters and hot summers, with rainfall evenly distributed throughout the yean
21.4°C in July; mean annual rainfall is 140.6 cm (Weather-Warehouse 2012). The average length of the growing
season ranges from 260 to 280 days. Short drought periods occur most commonly in April and from September
through November (Fernald & Purdum 1992).
Twenty of the 21 glades were visited by the authors at least once in spring (March-june) and once in fall
(September-November) between 2005 and 2012, and most were visited multiple times throughout the grow-
ing season. The exception was Snake glade, which was destroyed by road widening sometime between 1985
and 2005. Records for this glade are from four visits by Baker and Gholson in the 1980s. At each visit, species
checklists were made and voucher specimens collected. Plants were listed from the open glade, from the edges,
including the shrub zone between the open glade and the forested area around it, or the forest edge within 10
m of the open glade, if no shrub zone was present. Common non-vascular plant taxa and ground lichens were
also noted, but no attempt was made to compile an exhaustive list of these taxa. Once a relatively complete
list of vascular plant species was developed, the Angus Gholson herbarium (formerly AKG, now part of FLAS)
and R.K. Godfrey herbarium (FSU) were searched for specimens from the glades to serve as further vouchers.
In 2005 one new glade was found and this spurred a systematic search of aerial photographs to identify
potential new glades in the vicinity of known glades. Criteria used to distinguish glades openings from clear-
ings were: (1) sites that remained open on all aerial dates consulted; i.e., 2004, 1999, 1994 for both counties.
plus 1954-55 for Gadsden County (Thomas et al. 1961) and 1969 for Jackson County (Duffee et al. 1979); (2) j
sites that were within or close to the elevation contour lines in which known glades occur in each county, i.e,
90-100 ft in Gadsden county and 140-150 ft in Jackson County; and (3) sites that had soils associated with
known glades and/ or rock outcrop symbols on the soil survey maps for the respective counties. In Gadsden
County these were Binnsville soils or Cuthbert, Boswell and Susquehanna soils on moderate to steep slopes ,
(Thomas et al. 1961). In Jackson County, soils were the Oktibbeha variant rock outcrop complex (Duffee et al. j
1979).
RESULTS AND DISCUSSION
New glades i
In the search for new glades using aerial photography, fifty-one openings were identified in Jackson County j
and seventeen of these were field checked, yielding one relatively intact glade. Old Car, which was added to this ,
study, and three possible former glades that are now highly disturbed. The latter three, in addition to one pos-
sible disturbed glade known prior to this study and one subsequently pointed out to us, brings to five the total
of possible former glades confirmed on the ground in Jackson County. In Gadsden County, twenty-two poten-
tial glades were identified from aerial photography and nine of these were field checked. No new intact glades
were found, but three possible former glades were identified, which, along with one possible former glade
subsequently pointed out to us, brings to a total of four the number of possible former glades identified on the
ground in Gadsden County. Possible former glades were identified by presence of characteristic glades domi-
nants, such as Muhlenbergia capillaris, Rhynchospora divergens, Sporobolus vaginijlorus, or Stenaria nigricans,
plus characteristic forbs such as Polygala boykinii. The non-native centipede grass (Eremochloa ophiuroides) is
the principal invading species on former glades, where it often forms a dense turf, apparently precluding fur-
ther colonization by native glades species.
In addition to the larger graminoid-dominated glades, small glade-like openings in forested areas with
some of the characteristic glades forbs are known from four areas; Three Rivers State Park in Jackson County,
the Angus Gholson Nature Park in Chattahoochee, an area along the Florida-Georgia line north of Chatta-
hoochee, and an area known as “Brooks-3” west of Marianna in Jackson County.
Vegetation
Vegetation of the twenty-one glades consists of an open graminoid-dominated portion bordered by a charac
teristic set of calciphile shrubs and small trees that grade into the surrounding forest matrix on deeper soils
(Fig. 4a-b). These shrubs and small trees may also form clumps or islands on the open glade. The surrounding
forests (where still intact) consist of mesic upland hardwood forests on the steeper lower slopes, and pine-oak-
hickory forests, or, in a few cases, remnants of longleaf pine/wiregrass pinelands, on the gentler upper slopes.
Jumperus virginiana is the predominant woody species found on the open glades and on the edges. Ot er
shrubs and small trees consistently present on the edges of most glades are Celtis laevigata, Cercis canadensis,
Comus asperifolia, Fraxinus americana. Ilex vomitoria, Myrica cerifera, Rhamnus caroliniana, Sideroxylon lanugi
nosum, S. reclinatum, and Viburnum rufidulum. There is some differentiation between the two counties in com
mon woody species found on the glades. Acer saccharum ssp.Jloridanum is more common in Gadsden County
and Quercus muhlenbergii is more common in Jackson County. Crataegus spathulata, C. pulcherrima, and
alata are found on glades only in Gadsden County. ^
Dominant species of the herbaceous portion vary from glade to glade and from place to place wit i
glade. Schoenus nigricans, a large, clump-forming sedge, forms dense, nearly monospecific stands at
the twenty-one glades studied (Fig. 4a). Other large graminoids dominating portions of glades are Mu en
gia capillaris, Sporobolus junceus, and Schizachyrium scoparium. A short turf composed of Sporobolus vagini)
rus and/or Rhynchospora divergens (Fig. 4b) is often found on other portions of the glades. Additiona s ^
grasses commonly found on the glades include Aristida longispica, A. oligantha, Sporobolus clandes ^
Schizachyrium tenerum, and Panicum flexile. Areas with much bare soil or broken rock at the surface
Stenaria nigricans, along with the moss Weissia jamaicense and a cyanobacterium, Nostoc sp. Characteris
Johnson et al., Flora of calcareous upland glades
herbs found at nearly all the glades include Allium ca-
nadense var. mobilense, Asclepias viridiflora, Carex
cherokeensis, Polygala boykinii, Nothoscordum bivalve.
Ratibida pinnata, Rhynchospora colorata, and Sabatia
angularis.
A number of characteristic herbaceous species of the
glades were found only in one county or the other, but
not both. Piriquetadstoidesssp.caroHniana,Selagineliaapoda var. Iudovidana,Canirhoepapavcr,andSymphyot-
richum pratense were present on the majority of Gadsden County glades, but were absent from those m Jackson
County. Coreopsis sp. (aff. lanceolata), Rudbeckia triloba, and Packera anonyma occurred on the majority of
Jackson County glades and were absent from those in Gadsden County. Dispersal or substrate may contnbute
to these differences. The glade clusters in the two counUesareatleast40kmapart,sepa.atedby.hefioodplam
of the Apalachicola River. In Gadsden County the glades substrate is dolomite, whereas in Jackson County it is
limestone.
^rnlTnumber of taxa noted either on, or around the edge of, the twentyone glades is 31 1 of which 302 are
vascular plant species (Appendix). Ninety-eight species were found only on the edge of the glades. Of the vas-
cular plants 280 are native, 21 non-native, and one advemive from the west (Oenothera speciosa). The largest
family represented is Asteraceae with 50 species, followed by Poaceae (39), Cyperaceae (13), and Umiaceae
(12), Fabaceae (11) is typically one of the three largest familiesin the Qorasof other calcareous glades and prat-
ties, but is here tied with Rosaceae for fifth place. , , , ^ „
Nine of the non-native species on Florida upland gUdes are considered invasive by the Florida Exotic Pest
Plant Council (FLEPPC 2011, Table 1). Of these, only the three species of Ligustrum were frequently noted on
the Florida upland glades. Two non-native species not listed as invasive by the FLEPPC EremtxWtnr ophmrai-
des and Pyracanthakoidjumii, were also frequently noted on disturbed gUdes and, together with the Ligustrum
species, tend to make up the majority of the cover of non-native species on these^des^
Fifteen glades species are tracked by the Florida Natural Areas Inventory (WAI 2012) as rare tn Hori^
(Table2).Fourteeu of these arealso listed as endangered by the stateofFloridaOTonda Department of Agncul-
and Consumer Ser " - sonar k,™. „r .base are northern caldnhiles near their southern range imit in
). Most of these are northern
482
Journal of the Botanical Research Institute of Texas 7(1)
FLEPPC Category I -species that are
disrupt native plant communities.
on Fiorida glades (n=11); * = species not listed by the Florida Exotic Pest Piant Councii but which are a threat to Florida giades,
? invading and disrupting native plant communities in Fiorida; FLEPPC Category il - species that have shown a potentid to
Species
Ligustrum japonicum
Ligustrum lucidum
Ligustrum sinense
Lonicerajaponica
FIEPPC-Categoryl FLEPPC-Category II
Table 2. Rare species on Fiorida upiand glades (n=15). FNAI rankings are at the global (6) and state (S) levels: 1- critically imperiled because of extreme rarity:
2-imperiled because of rarity; 3-very rare or local; 4-secure; 5-demonstrably secure; ?-rank provisional; state of Florida ranking: LE-listed endangered (native
species in imminent danger of extinction in state due to rarity).
Florida. Only one, Mateleajloridana, is globally rare, the rest being rare in Florida, but relatively secure over
their total range (NatureServe 2012).
Six species appear to be largely confined to the Jackson and/or Gadsden County glades and are not found
elsewhere in Horida (Wunderlin and Hansen 2011and personal observations of authors): Asdepias viricHflora,
Boutelouacurtipendula, Coreopsis sp. (aff. lanceolata), Ratibidapinnata, Sporobolus vaginiflorus, and Symphyotn-
chum pratense. The moss, Pleurochaete luteola, was only known from the Jackson county glades, but has just
recently been found on a limestone outcrop along the St Marks River in Wakulla County, Florida (first author,
pers. obs.). Four others are known in Florida only from the edges and vicinity of glades: Delphinium carohnui-
num, Echinacea purpurea. Primula meadia, and Symphyotrichum shortii.
Coreopsis sp. (aff. lanceolata) is characterized by infolded, narrow, glabrous leaves which distinguish «
from C. lanceolata (Fig. 4c). It is frequent on all nine Jackson County glades, as well as on the five possible for-
mer glades in that county. It is absent from Gadsden County glades where Coreopsis lanceolata is found on eig ‘
of the twelve glades. This narrow-leaved glabrous Coreopsis has not been observed by us in habitats other than
iL, Flora of calcareous upland glades
4«3
glades and maintains its distinctness from C. lanceolata when the two are grown together from seed in a com-
mon garden (first author, pers. observation).
Pleurochaete luteola has not previously been reported from Florida. In the New World it ranges on calcare-
ous substrates from Virginia west to New Mexico and south to South America (Flora of North America Edito-
rial Committee 2007 vol. 27). It can form the dominant ground cover in semi-shaded cedar glades in Tennessee
(Quarterman 1950), is known from the Ketona Dolomite glades in Bibb County, Alabama (Allison & Stevens
2001), and was noted by the first author in 2009 at a blackland prairie at Oaky Woods Wildlife Management
Area, Houston County, Georgia. In Florida we found it on two glades in Jackson County, in both cases as a
single isolated patch in partial shade of Juniperus virginiana.
Symphyotrichum pratense is a western disjunct whose main range is centered on prairies in eastern Texas,
Louisiana, and Arkansas Qones et al. 2008). Eastward from its central range, it has widely scattered disjunct
populations in open calcareous habitats in MS, AL, GA, TN, KY, and VA. The Florida population at the Gads-
den glades is an isolated outpost at the southeastern extreme of its range Qones et al. 2008, Fig. 2). Three other
western species, Callirhoe papaver, Hymenopappus scabiosaeus, and Linum medium var. texanum, are frequent
on Florida upland glades, but rare or absent on the Alabama blackbelt prairies, Ketona Dolomite glades, and
Oaky Woods sites.
Like Pleurochaete luteola, Bouteloua curtipendula is very rare on Florida upland glades where it is at the
southern limit of its range. It is common on glades and calcareous prairies to the north and west (Baskin &
Baskin 2003; Campbell & Seymour 2012; Morris et al. 1993).
Of the 311 species listed, the number found at individual glades ranged from 43 to 162 (Table 3) and the
average number per glade (excluding Snake which was not sampled as intensively as the others) was 105. So-
renson’s Index of Similarity, i.e., the percentage of the average number of species in two glades that are held in
common (Mueller-Dumbois & Ellenberg 1974), was calculated for six of the more intact glades, three in Gads-
den (Brickyard, EB, and Humphrey) and three in Jackson County (Brooks-1, Bumpnose, and Old Car). Simi-
larities ranged from 66-69% among the three Gadsden glades and 66 to 71% among the three Jackson glades.
Pairwise comparisons between glades from the two different counties showed somewhat lower similarities,
ranging from 53 to 63%. A value >50% is generally required for samples to be considered part of the same plant
community (Barbour et al. 1980).
Comparison with floras of other calcareous openings
The percentage of Florida upland glades species found on other calcareous glades and prairies in the Southeast
was determined from a literature review. Table 4 shows a comparison with sites where relatively comprehen-
sive species lists over the entire growing season were compiled. Perhaps not surprisingly, the Florida upland
glades have the highest number of species in common with those areas closest to them, namely, the blackbelt
prairies in Georgia, Alabama, and Mississippi, and the Ketona Dolomite glades of Alabama, with decreasing
numbers of species in common with sites farther to the north (TN, AL, GA glades) and west (isolated calcare-
ous prairies in LA and TX). Comparison with species lists combined from a number of sites, compiled from the
literature for the AL/ MS blackbelt prairies and for the Southeastern cedar glades, shows the same pattern.
Occurrence of some common Florida upland glades species at nearby calcareous sites (listed in Table 4)
are shown in Table 5. Species common on Florida glades that also occur on most other calcareous openings in
the Southeast include two of the most characteristic Florida upland glades species, found in few other habitats
in the state, Asclepias viridiflora and Sporobolus vaginijlorus. Common Florida upland glades species found only
on the closest sites (Ketona Dolomite glades, AL/MS blackbelt, and Oaky Woods) are generally southern
coastal plain species. An example is Evolvulus sericeus, which reaches its northern range limit in Georgia
(Bridges & Orzell 1989). This is also largely true of common Florida upland glades species that are absent from
other calcareous openings to the north and west. Among these is a dominant species on Florida upland glades,
Schoenus nigricans, which is spottily distributed from northern Europe to South Africa and North America. In
the U.S it is found in Florida, Texas, California, and Nevada. Over its entire range it spans a wide diversity of
habitats from acid bogs in northern Europe to hot springs in California (Munz & Keck 1959; Sparling 1968). In
IS visited 4 times prior to 2005.
Florida it occurs on moist to wet calcareous substrates in the panhandle, throughout south Florida, and in two
counties on the central west coast of the peninsula (Wunderlin & Hansen 2008). Of the twenty-seven spedes
frequently found on nearhy calcareous openings hut not found on Florida glades, six occur in Florida in other
habitats and the rest do not range south to Florida. Five of these have their centers of distribution in the West
or Midwest and are disjunct to the east on calcareous glades and prairies (Bridges & Orzell 1986; Brown 2003).
CONCLUDING REMARKS
In terms of dominant species, Florida upland glades differ from other calcareous glades and prairies to tte
north and west in the dominance of species in the Cyperaceae (Schoenus nigricans, Rhynchospora divergens) and
the relative paucity of species in the Fabaceae.
Questions outside the scope of this paper that remain for future studies to answer include the relati«
roles soil texture, soil depth, and fire frequency have in controlling woody colonization of the Florida up
glades, plus the relative roles of disturbance and soil depth in determining the distribution of herbaceous
dominants within any single glade.
In the context of the rest of the Southeast, the Florida upland glades can be considered a somewhat
anomalous southeastern outpost of a series of calcareous glades and prairies extending eastward from Texas
through Louisiana, Mississippi, Alabama, and Georgia. From an examination of aerial photographs of pot®
tial glade openings in the two Horida counties, it appears likely that glade openings were more numerous w
pre-settlement times, but probably did not extend much further geographically than at present. The Flon ^
upland glades community (denoted as “upland glade”) is ranked GlSl (critically imperiled) by the Florida a ^
ural Areas Inventory (FNAI 2010) and is the only highly ranked community in Florida without a promi^
site under protection. Currently one small Florida glade is protected on a state park (Moranz) and two ( ^
road and Pride) are on public land that is not actively managed for their protection. Of those on private lan^
two have been destroyed by mining (Brooks-4 and Brooks-2) and five (Bumpnose, Brooks-1, Brickyard,
492
66:154-205.
Anderson, LC. 1 984. N
Anderson, LC.1988.N
mtsfrom north Florida. Sida 10:295-297.
mtsfrom north Florida III. Sida 13:93-100.
Anderson, L.C. 1 989. Noteworthy plants from north Florida IV. Sida 1 3:497-504.
Anderson, L.C. 2007. Noteworthy plants from north Florida VIII. J. Bot. Res. Inst. Texas 1 :/
Barbour, M.G., J.H. Burk, and W.D. Pitts. 1 980. Terrestrial plant ecology. The Benjamin Cum
Barone, J.A. and J.G. Hill. 2007. Herbaceous flora of blackland prairie remnants in Mississippi and western Alabama.
Castanea 72:226-234.
Baskin, J.M. and C.C. Baskin. 2000. Vegetation of limestone and dolomite glades in the Ozarks and midwest regions ofthe
United States. Ann. Missouri Bot. Gard. 87:286-294.
Baskin, J.M. and C.C. Baskin. 2003. The vascular flora of cedar glades ofthe southeastern United States and its phytogeo-
graphical relationships. J.Torrey Bot. Soc. 130:101-1 18.
). Evolvulus sericeus (Convolvulaceae) in Georgia, v
look. (Cyperac
A.S.B. Bull. 33:155-166.
Bridges, E.L. and S.L. Orzell.
13:509-512.
Bridges, E.L.,S.L. Orzell, AND
Brodo, I. M., S.D. Sharnoff, and S. Sharnoff. 2001 . Lichens (
Brown, L.E., K. Hillhouse, B.R. MacRoberts, and M.H. MacRoberts. 2002. The vascular flora of Windham Prairie, Polk County,
east Texas. Texas J. Sci. 54:227-240.
Brown, R.L. 2003. Paleoenvironment and biogeography ofthe Mississippi Black Belt: Evidence from insects. In: E. Pea-
cock and T. Schauwecker, eds. Blackland prairies ofthe Gulf coastal plain. University of Alabama Press, Tuscaloosa.
Pp. 11-25.
s response patterns of re
3 Florida. Sida 13:378
Crum, H.A. and LE. Anderson. 1981 . Mosses of Eastern North America. (
Diamond, D.D. and F.E. Smeins. 1985. Composition, classification, and S|
ries in Texas. Amer. Midi. Naturalist 1 13:294-308.
Duffee, E.M., WJ. Allen, and H.C. Ammons. 1 979. Soil survey of Jackson County, Florida. U.S. Department of Agriculture Soil
Conservation Service in cooperation with the University of Florida Institute of Food and Agricultural Sciences Agri-
cultural Experiment Stations, Soil Science Department, Gainesville.
Echols, S.L. and W.B. Zomlefer. 201 0. Vascular plant flora of the remnant blackland prairies in Oaky Woods Wildlife Man-
agement Area, Houston County, Georgia. Castanea 75:78-1 00.
Erickson, R.O., LG. Brenner, and J. Wraight. 1942. Dolomitic glades of east-central Missouri. Ann. Missouri Bot. Card
29:89-101.
Fernald, E.A. and E.D. Purdum, eds.1 992. Atlas of Florida. University Press of Florida, Gainesville.
FLEPPC. 201 1 . www.fleppc.org. Accessed July 201 2.
FNA (Flora of North America Editorial Committee). 1993-201 1 (continuing). Flora of North America north of Mexico.
New York and Oxford. Efloras.org. Accessed January 201 3.
Florida Department of Agriculture and Consumer Services. 2004. Rule Chap. 5B-40, Florida Administrative Code, contains the
[."Available from: Florida Department of Agriculture and Consumer Services, Division of Pla"*
"Regulated Plani
Industry, Gaines
FNAI. 201 0. Guide to the natural communities of Florida: 201 0 edition, www.fnai.org. Accessed January 201 3.
FNAI. 201 2. www.fnai.org/trackinglist. Accessed July 201 2.
Fan, T.L 1989. Blackland pi
(Asteraceae: Astereae). J. Bot. Res. Inst. Texas 2:731-739.
Kucera, C.L. and S. Curk. 1 957. Vegetation and soil relatioi
Ecology 38:285-291.
UWLESS, PJ., J.M. Baskin, and C.C. Baskin. 2004. The floristic ecology of xeric limestone prairies in Kentucky, and a compari-
son to limestone cedar glades and deep-soil barrens. Sida 21:1 055-1 079.
Lawless, PJ., J.M. Baskin, and C.C. Baskin. 2006. Xeric limestone prairies of Eastern United States: review and synthesis. Bot.
Rev. 72:235-272.
Leonard, S.W. and W.W. Baker. 1 982. Biological survey of the Apalachicola Ravines biotic region of Florida. Report to the
Florida state office of The Nature Conservancy and to the Florida Natural Areas Inventory, Tallahassee.
MacRoberts, B.R. and M.H. MacRoberts. 1 995. Vascular flora of two calcareous prairie remnants on the Kisatchie National
Forest, Louisiana. Phytologia 78:18-27.
MacRoberts, B.R. and M.H. MacRoberts. 1996. The floristics of calcareous prairies on the Kisatchie National Forest, Louisi-
ana. Phytologia 81:35-43.
MacRoberts, M.H., B.R. MacRoberts, and L. Stacey. 2003. Louisiana prairies. In: E. Peacock andT. Schauwecker, eds. Blackland
prairies ofthe Gulf coastal plain. University of Alabama Press, Tuscaloosa. Pp. 80-93.
Moore, W.E. 1 955. Geology of Jackson County, Florida. Bulletin No. 37, Florida Geological Survey, Tallahassee.
Morris, M.W., C.T. Bryson, and R.C. Warren. 1993. Rare vascular plants and associate plant communities from the Sand
Creek chalk bluffs, Oktibbeha County, Mississippi. Castanea 58:250-259.
Mueller-Dumbois, D. and H. Ellenberg. 1974. Aims and methods of vegetation ecology. John Wiley and Sons, NewYork.
Munz, P.A. AND D.D. Keck. 1 959. A California flora. University of California Press, Berkeley.
NatureServe. 201 2. NatureServe .org. Accessed July 201 2.
Nelson, G. 201 1 .The trees of Florida: a reference and field guide. Second edition. Pineapple Press Inc., Sarasota, FL.
Nelson, J.B. 1 985. Bouteloua curtipendula in Florida. Castanea 40:58.
Nelson, P. and D. Ladd. 1 980. Preliminary report on the identification, distribution, and classification of Missouri glades. In:
C. Kucera, ed. Proceedings ofthe Seventh North American Prairie Conference, Southwest Missouri State University,
Springfield. Pp. 59-76.
Peacock, E. and T. Schauwecker. 2003. The nature, culture, and sustainability of blackland prairies. In: E. Peacock and T.
Schauwecker! eds. Blackland prairies ofthe Gulf coastal plain. University of Alabama Press, Tuscaloosa. Pp. 1-7.
Quarterman, E. 1 950. Major plant communities of Tennessee cedar glades. Ecology 31 :234-254.
Rupert, F.R. 1 990. Geology of Gadsden County, Florida. Bulletin No. 62, Florida Geological Survey, Tallahassee.
Schotz, a. and M. Barbour. 2009. Ecological assessment and terrestrial vertebrate surveys for Black Belt Prairies in Ala-
bama Alabama Natural Heritage Program, Environmental Institute, Auburn University, Auburn. Report submitted
to Alabama Department of Conservation and Natural Resources, Division of Wildlife and Freshwater Fisheries, State
Wildlife Grants Program, Montgomery.
Schotz, A. 201 1 . A checklist of the vascular flora in the Black Belt Prairies of Alabama. Alabama Heritage Program, Envi-
ronmental Institute, Auburn University, Auburn.
Sims, P.L. 1988. Grasslands. In: M.G. Barbour and W.D Billings, eds. North American terrestrial vegetation. Cambridge
University Press, Cambridge. Pp. 266-286.
Sparling, J.T. 1 968. Biological flora of the British Isles: Schoenus nigricans L. J. Ecol. 56:883-899.
Thomas, B.P., H.H. Weeks, and M.W. Hazen. 1961. Soil survey of Gadsden County, Florida. U.S. Department of Agriculture,
Soil Conservation Service in cooperation with the University of Florida Institute of Food and Agricultural Sciences,
Agricultural Experiment Stations, Soil Science Department, Gainesville.
Ward, D.B. and A.K. Gholson. 1987. The hidden abundance of Lepuropetalan spathulatum (Saxifragaceae) and its first
reported occurrence in Florida. Castanea 52:59-67.
Ware, R. 2008. Pers. Comm. 2 Idlewood Court NW, Rome, GA.
Ware, S. 2002. Rock outcrop plant communities (glades) in the Ozarks: a synthesis. Southw.
Weakley, A.S. 2012. Flora of southern and mid-Atlantic states. Working draft of 30 November 201 2. University of North
Carolina Herbarium (UNC), North Carolina Botanical Garden, University of North Carolina, Chapel Hill. Herbarium.
unc.edu/FloraArchives/WeakleyFlora_201 2-Nov.pdf. Accessed January 201 3.
White, M. and D. Arbour. 201 2. Symphyotrichum pratense (Asteraceae): new for the flora of Oklahoma. Phytoneuron 201 2-
WulKl^TrjrrZsriWw'AtSFMavascularplams. www.plantatlas.usf.eda
■ titute for Systematic Botany,
■rsity Press of Florida,
E. Peacock and T. Schauwecke
loosa. Pp. 110-145.
2003. A plant (
;r, eds. Blacklan
munity classification for Arkansas blackland prairie ecosystem. In:
airies of the Gulf coastal plain, University of Alabama Press, Tusca-
HISTORY, DISPERSAL, AND DISTRIBUTION OF BUDDLEJA DAVlDll
(SCROPHULARIACEAE) IN KENTUCKY
Ralph L Thompson
Berea College Herbarium
Berea College, Biology Program
Berea, Ker^tucky 40404-2121, U.S.A.
J. Richard Abbott
P.O.BOX299
Saint Louis, Missouri 63 166-0299, U.S.A.
richard.abbott@mobot.org
RESUMEN
INTRODUCE
The genus Buddleja was named by Linnaeus (1753, 1754) to commemorate the English botanist and clergyman
Adam Buddie (1660-1715). A New World specimen of B. americana from the Caribbean was brought to Lin-
naeus in the 1730s by Dr. William Houstoun (Stuart 2006; Dirr 2009). Buddleja is a cosmopolitan group of
some 90-100 species of which many of the Asiatic taxa have been grown as ornamentals in arboreta, botanical
gardens, landscape gardens, nurseries, parks, garden clubs, and yards (Stuart 2006; Dirr 2009). The centers of
diversity for Buddleja are South Africa and Chinese-Himalayan Asia in the Old World (Leeuwenberg 1979;
Stuart 2006; Tallent-Halsell & Watt 2009) and South America, Central America, and southwestern United
States in the New World (Norman 2000; Tallent-Halsell & Watt 2009).
Buddleja davidii Franch. (Scrophulariaceae), summer lilac or orange-eye butterfly bush (Bailey & Bailey
1976; Norman 2012), are unarmed, multi-stemmed Chinese deciduous to semideciduous shrubs up to 5.0 m
tall and a spread of 6.5 m. Buddleja davidii gained its specific epithet, davidii, by Franchet (1888) to honor Pere
Armand David (1826-1900), a French Jesuit missionary and naturalist in eastern Tibet, who introduced the
first B. davidii to Adrian Rene Franchet (1834-1900) at the Paris Museum of Natural History (Stuart 2006;
Tallent-Halsell & Watt 2009).
It was introduced from China by the English plant collector, E.H. Wilson into the Royal Botanic Garden,
Kew, England, in 1896, and then at the Arnold Arboretum, Harvard University, around 1900 (Stuart 2006;
Tallent-Halsell & Watt 2009). Orange-eye butterfly bush is native to 13 provinces of central and southwestern
China and has been introduced into Japan (Zheng-Yi & Raven 1996; Stuart 2006). The 13 Chinese provinces
are mapped by Zheng et al. (2006) and Tallent-Halsell & Watt (2009).
Buddleja davidii has been variously placed by recent workers in the Loganiaceae, Buddlejaceae, and most
e (IPNl 2005; Norman 2012; Tropicos 2012; USDA, ARS 2012a; Weakley 2012).
Journal of the Botanical Research Institute of Texas 7(1)
Buddleja certainly has strong affinities within the Scrophulariaceae in the order Lamiales based on current
molecular systematics. Nomenclature for all taxa in this article follows Weakley (2012).
Buddleja davidii is the most widely cultivated species grown especially for its attractive dark green and
grayish-white pubescent foliage, flowers which vary from colorful lilac-rose to lilac-lavender, lilac-blue, red-
violet, purple to white, pleasant fragrance, and a high nectar content for the attraction of butterflies, moths,
bees, wasps, and birds (Norman 2000; Stuart 2006; Dirr 2009; Tallent-Halsell & Watt 2009). Buddleja davidii
has over 150 cultivars and hybrids recognized to date (see: Leeuwenberg 1979; Stuart 2006; Dirr 2009). A vig-
orous, cold hardy ornamental shrub, it is well adapted to USDA hardiness zones 5-9 (Stuart 2006; Dirr 2009).
In Kentucky, B. davidii grows well in zone 6 (-10 to 0°F [-20.6 to 17.8°C]) within the United States Hardiness
Zone Map (USDA, ARS 2012b).
We report Buddleja davidii as unequivocally naturalized in Kentucky for the first time with documenta-
tion based on recent plant collections, examination of herbarium specimens, data from plant collector’s log
notebooks, and a Kentucky literature survey. Our research indicates B. davidii has non-introduced populations
established in five counties during the last 30 year period.
Buddleja davidii is currently naturalized (sensu laid), i.e., growing without cultivation, in Australia, Europe,
Fiji, and New Zealand in the Eastern Hemisphere and in Canada, Central America, Puerto Rico, South Ameri-
ca, and the United States in the Western Hemisphere (Tallent-Halsell & Watt 2009; Norman 2012). Tallent-
Halsell and Watt (2009) mapped 20 states of the USA and the Canadian Provinces of British Columbia and
Ontario. The USDA, NCRS (2012) mapped the same USA distributions, but deleted Alabama and added Illi-
nois. In the conterminous United States, Kartesz (2011) included the District of Columbia, Delaware, Indiana,
Kansas, and Rhode Island in addition to the above 21 states in his Biota of North America Program (BONAP).
The USDA, FS (2012) mapped the combined distributions in the United States of Kartesz (2011), Tallent-Hal-
sell and Webb (2009), and the USDA, NRCS (2012), but also included Arizona, Arkansas, Louisiana, Missis-
sippi, Nevada, New Mexico, Texas, and Utah. Tropicos (2012) included Missouri based on voucher specimens
(G. Yatskievych&K. Yatshievych 02-62-, T.E. Smith 3832) deposited at the Missouri Botanical Garden Herbarium
(MO). The overall distribution of B. davidii in the United States and Canada from these five distribution sources
totals 34 states (including Hawaii), as well as the District of Columbia, the territory of Puerto Rico, and the
Canadian Provinces of British Columbia and Ontario.
The above distribution sources do not specify whether the non-native status of Buddleja davidii in the
United States (e.g. in Kentucky) is cultivated, introduced, persisting, adventive, or naturalized according to the
definitions by Nesom (2000). In essence, these map distribution sources probably represent a combination of
all those non-native classification categories from literature reports and herbarium vouchers.
Common temperate habitats for Buddleja davidii growing without cultivation in the United States include
railroad lines, limestone quarries, coal surface-mined lands, abandoned cultivated areas, urban disturbed ar-
eas, successional woodland edges, roadsides, riparian corridors, streambeds, floodplains, and sandy lake
shores, among many others (Tallent-Halsell & Watt 2009; Norman 2012). Orange-eye butterfly bush plainly
has a broad ecological amplitude, which favors colonization, establishment, and naturalization in a diversity of
open, insolated and weedy disturbed habitats.
As with numerous woody plants, Buddleja davidii has also become invasive in several countries where it
has been naturalized, e.g., Australia, England, France, New Zealand, and Hawaii, Oregon, and Washington in
the USA (Tallent-Halsell & Watt 2009; USDA, ARS 2012a; Young-Mathews 2011; Norman 2012). Buddlejada-
vidii was listed as one of the invasive Asian plants established in the United States by Zheng et. al. (2006). In
Oregon, B. davidii is classified as a “B” designated noxious weed subject to quarantine, and in Washington it is
listed as a Class “C” noxious weed (Tallent-Halsell & Watt 2009; Young-Mathews 2011; USDA, NRCS 2012).
Tallent-Halsell and Watt (2009) have provided the most comprehensive discussion on the biology, distribu-
tion, ecology, and invasiveness of Buddleja davidii.
Fic. 1. Kentucky counties with naturalized Buddlejadavidir. 1=Pike (1960 & 1982), 2=Calloway (2003), 3=Greenup (2010), 4=Powell (2011), 5=Uurel
(2012).
Kentucky Literature and Buddkja davidii in Pike County
The ascription of this species to the flora of Kentucky warrants a discussion of previous Kentucky literature
accounts. Buddleja davidii was credited as established in Kentucky by Medley (1993) and state-mapped by the
USDA, NRCS (2012), and other workers (e.g., Kartesz 2011; USDA, FS 2012; USDA, ARS 2012a). Tallent-Halsell
and Webb (2009) used these distribution resources and further based naturalization of B. davidii in their North
America map on being “escaped from cultivation.” However, their reported Kentucky distribution was found-
ed in part on the misapplication of Buddleja altemifolia, which was planted at the Bernheim Forest and Arbore-
tum (see: Gunn 1959).
Browne and Athey (1992) in their Vascular Plants of Kentucky: an Annotated Checklist, did not list Buddleja
davidii. Likewise, Jones (2005) in the Plant Life of Kentucky did not include B. davidii (Buddlejaceae) as part of
the Kentucky flora, but noted it was “to be expected as an escape in Kentucky.” Clark and Weckman (2008) did
not discuss or map B. davidii in their Annotated Catalog and Atlas of Kentucky Woody Plants.
The first Buddleja davidii specimen in a Kentucky herbarium was a cultivated plant collected in Fayette
County in Lexington at the University of Kentucky Agricultural Experiment Station, 24 Aug 1923, H. Carman
s.n. (BEREA, UK). The first report of a potentially escaped, non-cultivated orange-eye butterfly bush for Ken-
tucky is a Pike County voucher (Fig. 1) from 1960 on file at the University of Kentucky Herbarium (UK); but,
this specimen was not referenced by Medley (1993). The label data is:
KENTUCKY. Pike Co.: Cedar Street near Chloe Creek, 15 Oct 1960, Mary Louise Miniard 42 (UK)!
Medley (1993) in his dissertation, An Annotated Catalog of the Known and Reported Vascular Flora of Kentucky,
reported Buddleja davidii Franch. (Buddlejaceae) as “Rarely established along railroads and in disturbed areas
in at least two localities in Pike County on the Allegheny Plateau.” His two specimens from Pike County, Med-
ley & Levy 5895-82 and 6364-82 (DHL) were eventually transferred to the Western Kentucky University Her-
barium (WKU). However, these unmounted specimens were not available for study by us since they have not
yet been incorporated into WKU from the Davies Herbarium (DHL) of the University of Louisville. Fortu-
nately, Medley’s original 1982 collection log notebook was available at WKU and the relevant data (pp. 175-
182) are:
KENTUCKY. Pike Co.: Pikevllle 7-5 minute Quadrangle, Pikeville, spontaneous along abandoned railroad track in SW part of town, associ-
ates: Chaenorhinum minus, Desmodium spp., Eupatorium sp.. Ipomoea lacunosa, Oenothera biennis, and Sonchus asper, 16 Aug 1982, Max E.
Medley & Foster Levy 6364-82 (DHL).
498
The 1982 collection label data from the two disturbed areas noted by Medley (1993) from Pike County are
noteworthy: In Pikeville, Medley (5895-82) collected Buddleja davidii at McDonald’s Restaurant, while Levy
s.n. collected a specimen at Wendy’s Restaurant, each with the same three botanists, but with different dates
and restaurants. Medley’s collection data were gathered from his collection log notebook, and Levy’s specimen
was examined at the University of North Carolina Herbarium (NCU) at Chapel Hill:
s, behind Wendy’s in a rock pile, 18 Aug 1<
Campbell and Medley (2012) in their Atlas of Vascular Plants in Kentucky on Buddleja davidii Franch. (Buddle-
jaceae) stated, “Although widely cultivated (as “butterfly bush”), there are few reports of this Chinese shrub
escaping in eastern North America.” Campbell and Medley noted that B. davidii may be increasing in Appala-
chian regions in the southeastern states and referred to the Pike County collection data from Medley (1993).
Campbell’s incidental collection of a seedling at a woods edge on a sandy ridge in Pulaski County, was not
found in any Kentucky herbarium. Regardless of their discussion, Campbell and Medley (2012) did not map B.
davidii in their Kentucky vascular plant atlas.
Buddlga davidii in CaUoway County
In Calloway County (Fig. 1), during the summer 2003, two flowering specimens of Buddleja davidii were col-
lected (Thompson 03-389) in Quaternary Ochlockonee alluvial gravelly loam soils along a roadside ditch
thicket near the southernmost boundary of Hancock Biological Station on Kentucky Lake. The vouchers were
deposited in the Berea College Herbarium (BEREA) and Murray State University Herbarium (MUR). The ma-
ture shrub was treated as a non-cultivated escape since there were no homesteads or building remains any-
where within its vicinity, nor was it close to any populated area. The nearby Pacer Point Marina, an asphalt boat
ramp landing, was not likely a source for the plant. Exact label data from this Calloway County site are:
KENTUCKY: Calloway Co.: Pacei Point Marina area. 0.8 mi E from jet. of Lancaster Road to Watersport Road, and 0.5 mi S on Wateispoti
Buddl^a davidii in Greenup County
Orange-eye butterfly bush was collected in Greenup County (Fig. 1) on a roadside along a creek near a residen-
tial area in 2010, but without a town or city listed on the label data. It is not known if the shrub was from a
cultivated or non-cultivated plant, although it may well have been an escaped plant. The label data from this
specimen at Morehead State University Herbarium (MDKY) are:
KENTUCKY. Greenup Co.: roadside at end of creek with much Quercus and Ambrosia, elev. 617 ft, near residential area. 38”26’24”N and
83°4r23’’W, 4 Oct 2010, Steffany Seagraves 31 (MDKY)!
Buddleja davidii in Powell County
David Taylor, USDAFS botanist, discovered a volunteering Buddleja davidii shrub in Powell County (Fig-
during the summer 2011 in the Daniel Boone National Forest within the Red River Gorge on Alticrest-Ramsey-
Rock Outcrop complex soils over Pennsylvanian-aged Lee Formation conglomeratic sandstone. The plant was
found during a reconnaissance of an area severely burned in the fall of 2010. The wildfire burned organic mat-
ter to mineral soil and killed most overstory, midstory and understory trees, and 50-60% of the shrubs. At the
time of collection, bare mineral soil accounted for 60-70% of the surface cover (D.D. Taylor pers. comm. 2012)
Data from his specimen, “On Permanent Loan From USDA Forest Service,” are as follows:
KENTUCKY. Powell Co.: Daniel Boone National Forest. Red River Gorge, lat. 37.825827, long. -83.678275, elev. 1220 ft asl. m moi«
TussUagofarfara. (BEREA)
Establishment and Dispersal of Buddlga davidii in Laurel County
The most recent Kentucky collections of Buddleja davidii were from th
sites of Laurel County (Fig- ’
500
Journal of the Botanical Research Institute of Texas 7(1)
during September and November 2012. While on a September 4th collecting trip towards a vascular flora of
Laurel County, we discovered a non-cultivated B. davidii established along the CSX Railway in the city of Lon-
don and voucher specimens were deposited in BEREA and the Missouri Botanical Garden Herbarium (MO). A
single established orange-eye butterfly bush 1.5 m high with a cymose panicle of lilac-colored flowers with
orange-eyed throats was present and appeared locally naturalized at this site. A second collecting trip to the
general site on September 10th revealed another flowering individual a short distance away from the first col-
lection site. On November 4th, a third trip was made to collect fruits from the two previous colonies at which
time three additional fruiting shrubs and four scattered seedlings were discovered on the east side of the CSX
railroad right-of-way, ca. 100 m south from either of the two previous plant sites. Representative voucher speci-
mens were taken from the populations of the second and third trips and placed on file in BEREA, MO, and
NCU. Duplicate specimens from the collecting trips have been distributed to other herbaria as designated by
herbarium acronyms from Thiers (2012).
Our collection label data of Buddleja davidii from these three reconnaissance trips to the CSX Railway in
London, Laurel County, are summarized below:
Characteristic associates growing in the granite, gneiss, and limestone railroad aggregate ballast were Ambro-
sia artemisiifolia, Cichorium intybus, Conyza canadensis, Digitaria sanguinalis, Euphorbia maculata, E. mtans,
Ipomoea coccinea, Kummerowia striata, Lathyrus latifolius, Oenothera biennis, Sonchus asper, Symphyotridm
pilosum, and Trifolium repens. Some associated plants at the ditch margin included Acer negundo, A. rubrum,A.
saccharinum, Ambrosia trifida, Catalpa speciosa, Echinochloa muricata, Eupatorium serotinum, and Paspalum
KENTUCKY. Laurel Co.: London Quadrangle, London, East 4th Street (KY 80) S of Flowers Bakery and 4.0 m S 2.0 m W of the CSX railroad
tracks adjacent to Railroad Street, elev. 359 m, lat. 37.116905, long. -84.066788, Powering shrub severely cut back, although not herbickk-
^prajed, 10 Sep 2012, R.L. Thompson 12-1097 (BEREA, MO); 4 Nov 2012, R.L. Thompson & K. Rivers Thompson 12-1138 (BEREA, BRIT, MO,
This orange-eye butterfly bush represents a second non-cultivated individual 16 m diagonally across from the
shrub at the concrete culvert, probably from a seed source spread by trains. Ruderal associated species ob-
served around Buddleja davidii in the railway aggregate between KY 80 and Railroad Street were Ambrosia otte-
misiifolia, Conyza canadensis, Daucus carota, Digitaria sanguinalis, Echinochloa crusgalli, Eleusine indica.
Eragrostis pectinacea, Eupatorium serotinum, Euphorbia dentata, E. maculata, E. nutans, Kummerowia striata.
Lespedeza cuneata. Polygonum aviculare, Setaria pumila, Solidago altissima, Symphyotrichum pilosum, and Ver-
bascum thapsus.
KENTUCKY. Laurel Co.: London Quadrangle, London, S of KY 354N, across East 4th Street (KY 80) S 120 m down to 216 McLemore Street
on E side of CSX Railroad right-of-way and grassy street margin, elev. 357 m, lat. 37.116880 long -84.066782, 04 Nov 2012, Rl-
& K. Rivers Thompson 12-1140 (BEREA. BRIT, EKY, KNK, MDKY, MO. MUR, NCU, TENN, UK).
This third collection corroborates further evidence for the naturalization of Buddleja davidii, as three addi-
tional fruiting shrubs and four seedlings were scattered for 35 m along the east side of the railroad right-of-way
A personal account from a local resident (C. House, pers. comm. 2012) noted several seedlings had become
established at the edge of his yard at 216 McLemore Street and the ballast of the railroad bed during the last
or 3 years. He remarked “this area hasn’t been [herbicide] sprayed and those three shrubs first appeared 3 or
years ago.”
Only 12 depauperate associates were identified at this grassy habitat due to recent heavy fall frosts. Exonc
and native herbaceous taxa in this habitat were Amaranthus hybridus, Andropogon virginicus, Conyza canaden-
sis, Croton monanthogynus, Cynodon dactylon, Digitaria sanguinalis. Euphorbia nutans, Lamium amplexicau e, ■
purpureum, Muhlenbergia schreberi, Schedonorus arundinaceus, and Setaria pumila.
501
Buddleja davidii Naturalized in Kentucky
We have classified Buddleja davidii as clearly naturalized for the first time in Kentucky based o
tion of the “naturalized” category of Nesom (2000): a non-native plant population introduc
from other introduced populations, which are spreading or dispersing from year to year wi
anthropogenic assistance.
The data from voucher specimens in Pike, Calloway, and Greenup
B. davidii in Kentucky over the last 30 years. Even if the species were orij
counties, their habitat locations would infer otherwise and they would still
into other non-cultivated areas. The 2011 Powell County collection of a volunteering, established non-intro-
duced B. davidii validates naturalization in Kentucky. In Laurel County, the five new non-introduced shrubs
and seedlings growing without cultivation within the railroad right-of-way habitats of London provide conclu-
sive evidence of naturalization of orange-eye butterfly bush in Kentucky. Nevertheless, the naturalization of B.
davidii in Kentucky at this time clearly does not imply classification of it as an “invasive species" based on clas-
sification standard criteria from the KY-EPPC (2012).
The most prudent explanation of plant propagule spread for this population of Buddleja davidii is the
phenomenon of “slipstreaming,” the low pressure drag and blowing of scattered diaspores caused by methods
of moving transportation (Eskridge & Hunt 1979; Gamier et al. 2008; von der Lippe et al. 2013). The numcr-
streaming wake of railway train locomotives and freight boxcars.
Although Buddleja davidii populations in North America are not restricted to active or abandoned railroad
habitats, we consider these three established colonies of B. davidii in Laurel County, Kentucky, to have been
naturalized through the dispersal of propagules by the CSX Railway.
Railways have played an important role in providing suitable habitats and dispersal means of vascular
plants. Slipstreaming along railroads has been verified as a means for the rapid dissemination of non-native
fruits and seeds in the United States from literature reports and herbarium vouchers, e.g., Chaenorhinum minus
(Plantaginaceae s.l). The seeds of this naturalized European annual have been effectively and rapidly dis-
persed by the action of railway trains and their linked boxcars (Arnold 1979, 1981, 1991; Widrlechner 1982;
Sauer 1988). Railroad rights-of-ways have been correlated with numerous exotic weedy invaders in Illinois,
Missouri, and Kentucky (Thompson & Heineke 1977; Thompson 1979; Cranfill & Thieret 1981; Medley et al.
1983; Thieret & Thompson 1984).
Four examples from the literature and herbarium specimens illustrate the occurrence of railroad trans-
portation of wind-dispersed seeds and establishment specifically for Buddleja davidii. Miller (1984) in his dis-
sertation reported that B. davidii had been originally spread throughout Great Britain along railroad tracks by
being carried on freight cars and through train slipstreaming.
Muhlenbach (1979) related the slipstream dragging effect by trains in St. Louis railroad yards and the ad-
dition of aggregate ballast as two means of dispersing propagules of herbaceous and woody plants, including
Buddleia davidii, as documented by:
MISSOURI. St. Louis Co.: collections from Railroad Sites: St. Louis Union Sution on track 9. one big specimen, about 5 ft apart from a
Fragaria sp. colony of huge size, not planted, determined by E. Norman, 18 Jul 1971, V. Muhlenbach 3622 (MO)!
Radford et al. (1968) in the Manual of the Vascular Flora of the Carolinas similarly reported Buddleja davidii as a
rare introduction along a railroad in North Carolina. Label data from NCU confirms that information:
north CAROLINA. Haywood Co.: cinder bed along railroad, 1.9 mi E of Canton, US 19-23, 15 Jul 1958, Harry E. Ahles &J.A. Duke 465 16
(NCU)!
In Knoxville, Knox County. Tennessee, an escaped population of Buddleja davidii was documented on Novem-
ber 21, 2012, along a railroad bed adjacent to Mead’s Quarry Lake, a part of tbe Ijams Nature Center. This popu-
lation was discovered from information given by Dr. Dwayne Estes, Austin Peay State University Herbarium
503
(APSC) curator, who discovered the B. davidii population in 2003 (D. Estes, pers. comm. 2012).
Many mature shrubs as well as smaller volunteering shrubs and seedlings have colonized a 75 m exten-
sion of a railroad bed to the SSE of the three demolished limestone kilns. Orange-eye butterfly bush is locally
naturalized at this Tennessee site, which provides evidence on the concept of wind-dispersed seeds being
spread and established shrub populations being created through the slipstreaming action of active trains. This
railway quarry site data are:
TENNESSEE. Knox Co.: Knoxville, Mead’s Quarry Lake of the Ijams Nature Center, elev. 269 m, lat. 35.950533, long. -83.867525, several
20 Nov 2012, R.L. Thompson & K. Rivers Thompson 12-1153 (APSC, BEREA, MO, NCU, TENN).
Representative herbaceous associates with Buddleja davidii included Ambrosia artemisiifolia, Andropogon vir-
ginicus, Conyza canadensis, Croton monanthogynus, Daucus carota, Lespedeza cuneata, Panicum flexile, Polymnia
canadensis, Schedonorus arundinaceus, Solidago altissima, Sorghum halepense, and Symphyotrichum pilosum,
among several others. Characteristic woody taxa between the railway tracks and kilns were Acer negundo, Ai-
lanthus atissima, Celtis occidentalis,Juniperus virginiana, Ligustrum sinense, Lonicerajaponica, L. maackii, Plata-
nus occidentalis, Rhus glabra, Robinia pseudoacacia, Rubus occidentalis, Pyrus calleryana, and Rosa multiflora.
The diagnostic description of Buddleja davidii here provides the first baseline taxonomic description directly
from Kentucky material. It is notably modeled after the excellent species descriptions of Norman (2000, 2012).
Measurements were made of 33 living branches from the three Laurel County habitats prior to prepara-
tion as herbarium specimens. Measurements of flowers, fruits, and seeds were made with the use of a 5 mm
Micro-ScaleTM and a 10 mm Micro-Scale™ Micro-Tools from Electron Microscopy Sciences, Hatfield, Pennsyl-
vania, in combination with a Bausch & Lomb StereoZoom 4 Microscope™.
Shrubs deciduous, unarmed, multi-stemmed, 1.2-2.2 m tall, 1.0-1.8 m spread, older branchlets greenish
brown to brown sparse stellate-tomentose to glabrescent; young branchlets tetragonous, densely grayish
white canescent to stellate-glandular tomentose (Fig. 2). Leaves opposite, simple, pinnate, 10-15 lateral veins,
blade narrowly to broadly elliptic-lanceolate to lanceolate-ovate 6.0-13.0 x 1.5-5.5 cm, dark forest green and
glabrous adaxially, white stellate-tomentose abaxially, apex sharply acute, base cuneate-attenuate, margin ser-
rate, new lateral leaf flushes white-canescent; petiole 5.0-10.0 mm, stipules, narrow auriculate, LO-2.0 mm
high, forming prominent stem lines adjoining leaf bases (Fig. 3). Inflorescences terminal, indeterminaf- -•
soid cymes tapered and secund, 10.0-36.0 x 3.0-5.5 cm, often with two lateral branches, pseudovciucc
cymes, 15-30 pairs each with 3-20 flowers, peduncles 5.0-15.0 mm (Fig. 3). Flowers numerous ^rfect
4-merous Calyx campanulate sessile to pedicillate 1.0 mm, persistent, greenish to red-purple, tube 2.4-3.1
1.0-1.6 mm stellate and glandular outside, glabrous inside; lobes acute, 0.8-L5 mm. Corolla salverform, lilac-
rose lilac-lavender lilac-blue to red-purple, tube subcylindrical 8.0-12.0 x L0-L2 mm, glabrous to sparsely
stellate outside pilose inside, lobes suborbicular to spatulate, 2.0-3.0 mm, glabrous, margin crenate-f—
limb 7.0-9 0 mm with a narrow white to yellow ring around an orange circle (eye) extending LO-1.5 mm into
throat. Stamens 4, sessile, anthers dorsifixed, LO-1.2 mm, inserted at or near the middle of the pilose corolla
tube. Ovary superior, ovoid, glabrous, syncarpous bicarpellate, 2.0-3.0 x 0.9-Ll mm, placentation axi
ma clavate 15-20x0 8-1 3 mm (Fig 4). Capsules septicidal, two-valved dehiscence, light olive green to
brown a, malnrity, narrowly linear-oblong to eUiptic, 9.0-1S.0 , 1.8-2,3 mm (Fig. 5). Seeds 30-40, linea
siform, long-winged, 4.0-5.3 x 0.4-0.5 mm. dark brown, dorsivcntrally compressed, wings translucent
brown, tapered at both ends, 1.7-2.1 mm (Fig. 5).
ACKNOWLEDGMENTS
We express our appreciation to Keith E. Clancy (USDA, APHIS, Jamaica, NY) ®
Derick B. Poindexter, University of North Carolina, Chapel Hill, and Ross C. Clark fm constmcttve manu-
script reviews. We convey thanks to David D. Taylor, USDAFS, Daniel Boone National Forest, for hts descnp-
Journal of the Botanical Research Institute ofTexas7(1)
tive site data and the Powell County specimen, and to Dwayne Estes, Austin Peay State University, for the loca-
tion of a Tennessee population. We thank the curators of APSC, EKY, KNK, KY, MO, MDKY, MUR, NCU
TENN, UK, and WKU for their courtesy during herbarium visits and/or for confirming their specimen
holdings.
REFERENCES
Arnold, R.M. 1979. Railroad lines as study sites in plant biology. Amer. Biol. Teacher 41:548-551.
Arnold, R.M. 1981. Population dynamics and seed dispersal of Chaenorrhinum minus on railroad cinder ballast. Amer.
Midi. Nat. 106:80-91.
Arnold, R.M. 1991. Dwarf snapdragon— the railroad weed. Weed Technol 5:467-468.
Bailey, LH. and E.Z. Bailey. 1976. Hortus Third: a concise dictionary of plants cultivated in the United States and Canada.
Macmillan, New York, New York.
Browne, E.T., Jr. and R. Athey. 1 992. Vascular plants of Kentucky: an annotated checklist. The University Press of Kentucky,
Campbell, J. and M. Medley. 201 2. The atlas of vascular plants in Kentucky. Draft of January 2012, with provisional listing of
authors. (httpsy/www.bluegrasswoodland.com). Six PDFs (https://www.onlinefilefolder.com, 20 Nov 2012).
Clark, R.C. and T.J. Weckman. 2008. Annotated catalog and atlas of Kentucky woody plants. Castanea, Occas. Pap. E. Bot.
3:1-113.
Cranfill, R. and J.W. Thieret. 1 981 . Thirty additions to the vascular flora of Kentucky. Sida 9:55-58.
Dirr, M.A. 2009. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propaga-
tion and uses, 6th Ed. Stipes Publishing L.L.C., Champaign, Illinois.
Eskridge, R.E. and J.C.R. Hunt. 1979. Highway modeling.1. Prediction of velocity and turbulence fields in the wake of
vehicles.! Appl. Meteorol. 18: 387-400.
Franchet, A.R. 1888. Nouv.Arch. Mus. Hist. Nat., Ser2, 10:103, or Pl.david. 10:103-104.
Garnier, a., S. Pivard, and J. Lecomte. 2008. Measuring and modelling anthropogenic secondary seed dispersal along
road verges for feral oilseed rape. Basic Appl. Ecol. 9: 533-541 .
Gunn, C.L 1 959. A flora of Bernheim Forest, Bullitt County, Kentucky. Castanea 24:61 -98.
IPNI. 2005. The international plant name index, (httpy/ipni.org/, 18 Nov 2012).
Jones, R.L 2005. Plant life of Kentucky: an illustrated guide to the vascular flora. The University Press of Kentucky, Lex-
ington.
Kartesz, J.T. 201 1 .The Biota of North America Program (BONAP). North America plant atlas. Chapel Hill, NC. (http//wvYW.
bonap.org/MapSwitchboard.html, 27 Nov 201 2).
Leeuwenberg, AJ.M. 1979. The Loganiaceae of Africa XVIII. Buddleja L II. Revision of the African and Asiatic species. Vee-
man. H. & Zonen, B.V. Meded. Landbouwhoogeschool Wageningen 79:1-163.
Linnaeus, C. [von LinnE, C.]. 1 753. Species plantarum: Ed. 1 :1 1 2. Tomus II. Holmiae.
Linnaeus, C. [von LinnE, C.]. 1 754. Genera plantarum: Ed. 5, 51 . Leyden, the Netherlands.
Medley, M.E. 1993. An annotated catalog of the known or reported vascular flora of Kentucky. Ph.D. dissertation. Univer-
sity of Louisville, Louisville, Kentucky.
Medley, M.E., R. Cranfill, and J.W. Thieret. 1983. Vascular flora of Kentucky: additions and other noteworthy collections.
Sida 10:114-122.
Miller, A. 1 984. The distribution and ecology of Buddleja davidii Franch. in Britain, with particular reference to conditions
supporting germination and the establishment of seedlings. Ph.D. dissertation, CNAA, Oxford Polytechnic, United
MOhlenbach, V. 1 979. Contributions to the synanthropic (adventive) flora of the railroads in St. Louis, Missouri, U.S.A. Ann.
Missouri Bot.Gard. 66:1-108.
Nesom, G.L. 2000. Which non-native plants are included infloristic accounts? Sida 19:189-193.
Norman, E.M. 2000. Buddlejaceae. Flora Neotropica Monograph 81 :l-225. New York Botanical Garden, New York.
Norman, E.M. 2012. Buddleja. Flora of North America, Provisional Publication, Vol. 1 7. Flora of North America Association.
9 March 201 2. (fna.huh.harvard.edu/files/Scrophulariaceae.pdf, 3 Dec 201 2).
Radford, A.E., H.E. Ahles, and C.R. Bell. 1 968. Manual of the vascular flora of the Carolinas.The University of North Carolina
Press, Chapel Hill, North Carolina.
sofgeograj
c patterning in seed p
Sauer, J.D. 1988. Plant migration: the
Press, Berkeley and Los Angeles.
Stuart, D.D. 2006. Royal Horticultural Society plant collector guide: Buddlejas. Timber Press, Inc., Portland, Oregon.
Tallent-Halsell, N.G. and M.S. Watt. 2009. The invasive Buddleja davidii (Butterfly Bush). Bot. Rev. 75:292-325.
Thieret, J.W. and R.L. Thompson. 1 984. Cleome ornithopodioides (Capparaceae): adventive and spreading in North America.
Bartonia 50:25-26.
Thiers, B. 2012 [continuously updated]. Index Herbariorum: a global directory of public herbaria and associated staff.
New York Botanical Garden's Virtual Herbarium, (httpy/sweetgum.nybg.org/in/, 22 Nov 2012).
Thompson, R.L. 1979. The vascular flora of Cedar Gap Lake and environs, Webster and Wright Counties, Missouri. Sida
8:71-89.
Thompson, R.L andT.E. Heineke. 1 977. Vascular flora of the Desoto-Hallidayboro railroad prairie strips, Jackson County, Il-
linois. Trans. Illinois State Acad. Sci. 70:1 14-127.
Tropicos. 201 2. Buddleja davidii Franch. Nomenclatural and specimen database of the Missouri Botanical Garden. (httpV/
www.tropicos.org/Name/1 9000534, 27 Nov 2012).
USDA, ARS. 201 2a. USDA Agriculture Research Service, National Genetic Resources Program. Germplasm Resources In-
formation Network (GRIN), Beltsville, MD. (httpy/www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?8081, 19 Nov 2012)
USDA, ARS. 2012b. USDA Agriculture Research Service. USDA 2012 Plant Hardiness Zone Map. (httpV/www3rs.usda.
gov/PHZMWeb/images, 20 Nov 2012)
USDA, FS. 2012. Weed of the week: Butterfly Bush (Buddleja davidii Franch.). USDA, Forest Service, Forest Health Staff,
Newtown Square, PA. Invasive Plants website: (httpy/na.fs.fed.us/fhp/invasive_plants/we€ds/butterfly_bush/pdf,
25 Nov 201 2)
USDA, NRCS. 2012. The PLANTS Database, (http://plants.usda.gov/java/profile7symbok BUDA2, 29 Nov. 2012). Na-
tional Plant Data Team, Greensboro, NC 27401-4901 USA.
Von Der Lippe, M., J.M. Bullock, 1. Kowarik, T. Knopp, and M. Wichmann. 201 3. Human-Mediated dispersal of seeds by the air-
flow of vehicles. PLOS ONE 8(1 ):e52733.doi:l 0.1 371 /journal, pone.0052733.
Weakley, A.S. 2012. Flora of the southern and mid-Atlantic states: working draft of 12 September 2012. University of
North Carolina Herbarium, Chapel Hill, North Carolina. (httpV/www.herbarium.unc.edu/flora.htm, 27 Nov. 2012).
WiDRLECHNER, M.P. 1982. Historical and phenological observations on the spread of Chaenorrhinum minus across North
America. Canadian J. Bot. 61:179-187.
Young-Mathews, a. 201 1 . Plant fact sheet for orange eye butterflybush (Buddleja davidii). USDA-Natural Resources Con-
servation Service, Corvallis Plant Materials Center, Corvallis, Oregon.
Zheng, H., Y. Wu, J. Ding, D. Binion, W. Fu, and R. Reardon. 2006. Invasive plants of Asian origin established in the United
States and their natural enemies. Vol. 1 . FHTET, 2nd Ed. Chinese Academy of Agricultural Sciences, Beijing, China, and
USDA Forest Service, Morgantown, West Virginia.
Zheng-Yi, W. and P.H. Raven, eds. 1996. Flora of China, Vol. 15, Myrsinaceae through Loganiaceae. Science Press, Beijing,
China, and Missouri Botanical Garden, St. Louis, Missouri.
506
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2. Arthur Kelman: Tribute and Remembrance — Luis Sequeira
3. Stagonospora nodorum: From Pathology to Genomics and Host Resistance — Richard P. Oliver, Timothy L
Friesen, Justin D. Paris, and Peter S. Solomon
4. Apple Replant Disease: Role of Microbial Ecology in Cause and Control — Mark Mazzola and Luisa M.
Pathogenomics of the Ralstonia solanacearum Species Complex — Stephane Genin and Timothy P. Denny
. The Genomics of Obligate (and Nonobligate) Biotrophs — Pietro D. Spanu
Genome-Enabled Perspectives on the Composition, Evolution, and Expression of Virulence Determinants
in Bacterial Plant Pathogens — Magdalen Lindeberg
Suppressive Composts: Microbial Ecology Links Between Abiotic Environments and Healthy Plants—
Yitzhak Hadar and Kalliope K. Papadopoulou
Plant Defense Compounds: Systems Approaches to Metabolic Analysis — Daniel J. Kliebenstein
L Role of Nematode Peptides and Other Small Molecules in Plant Parasitism — Melissa G. Mitchum, Xiaohong
Wang,Jianying Wang, and Eric L. Davis
. New Grower-Friendly Methods for Plant Pathogen Monitoring* — Solke H. De Boer and Maria M. Lopez
Somatic Hybridization in the Uredinales — Robert F. Park and Colin R. Wellings
.. Interrelationships of Food Safety and Plant Pathology: The Life Cycle of Human Pathogens on Plants-Jen
D. Barak and Brenda K. Schroeder
). Mechanisms and Evolution of Virulence in Oomycetes — Rays H.Y. Jiang and Brett M. Tyler
). Variation and Selection of Quantitative Traits in Plant Pathogens — Christian Lannou
r Gall Midges (Hessian Flies) as Plant Pathogens— Je//J. Stuart, Ming-Shun Chen, Richard Shukle, and Marion
O. Harris
i. Phytophthora Beyond Agriculture — Everett M. Hansen, Paul W Reeser, and Wendy Sutton
I Landscape Epidemiology of Emerging Infectious Diseases in Natural and Human-Altered Ecosystems^
Ross K. Meentemeyer, Sarah E. Haas, and Tomds Vdclavt'fe
3. Diversity and Natural Functions of Antibiotics Produced by Beneficial and Plant Pathogenic Bactena—
M. Raaijmakers and Mark Mazzola
I. The Role of Secretion Systems and Small Molecules in Soft-Rot Enterobacteriaceae Pathogenicity—^
Charkowski, Carlos Blanco, Guy Condemine, Dominique Expert, Thierry Franza, Christopher Hayes, Nico^
Hugouvieux-Cotte-Pattat, Emilia Lopez Solanilla, David Low, Lucy Moleleki, Minna Pirhonen, Andrew Pitman’
Nicole Perna, Sylvie Reverchon, Pablo Rodriguez Palenzuela, Michael San Francisco, Ian Toth, Shinji Tsuyttma,
Jacquievan der Waals,Jan van der WolJ, Frederique Van Gijsegem, Ching-Hong Yang, and Iris Yedidia
I Receptor Kinase Signaling Pathways in Plant-Microbe Interactions — Meritxell Antolin-Llovera, Martina
Ried, Andreas Binder, and Martin Pamiske
J. Fire Blight: Applied Genomic Insights of the Pathogen and Host — Michael Malnoy, Stejan Martens,Jo n
Norelli, Marie-Anne Bamy, George W. Sundin, Theo H.M. Smits, and Brion Duffy
LBot Res. InsLTexaj 7(1): 506.2013
ASTROLEPIS SINUATA (PTERIDACEAE) NEW
TO THE FLORA OF LOUISIANA
JeffMcMillian
Charles M. Allen Selena Dawn Allen
Almost Eden Plants
1240 Smith Rd
Merryville, Louisiana 70653, U.SA.
Colorado State Univ., Fort Polk Station
1697 23rd St
Fort Polk, Louisiana 71459, U.SA.
5072 Hwy 399
Pitkin, Louisiana 70656, U.SA.
ABSTRACT
RESUMEN
A recent collection of Astrolepis sinuata (Lag. ex Sw.) Benham and Windham from southwestern Louisiana is
apparently the first for the state. This species is not listed for Louisiana by Thieret (1980), Neyland (2011), Ben-
ham and Windham (1993), Thomas and Allen (1993), or the USDA NRCS (2012). It is reported from Arizona,
New Mexico, and western and central Texas. There the range extends eastward to Comal and Anderson coun-
ties (Diggs et al. 2006; Diggs & Lipscomb 2013), into Austin County (Diggs & Lipscomb 2013) and also to Ellis
County (USDA NRCS 2012). A disjunct population is reported from Meriwether County, Georgia (Benham &
Windham 1993). A duplicate specimen was verified by Dr. James Peck of the University of Arkansas at Little
Rock. This plant was the only one observed at the site but had multiple stems so our collecting did not eradicate
it. The plant is not known to be cultivated in the area so its mode of distribution to this site is unknown. No
additional plants of this species were located in a check of the other pillars on this bridge and those of nearby
bridges but additional populations of this fern might be found in similar habitats in the future in western Loui-
siana and eastern Texas.
ACKNOWLEDGMENTS
Michael MacRoberts (LSUS) and an anonymous reviewer provided helpful comments.
i,D.M.ai
1.1 993. In: Florae
e of Texas Press, Fort
Pteridophytes and Gymnosperms. Oxford Univ. Press, New York.
Diggs, G.M., Jr. and B.L Lipscomb. 2013. Ferns and lycophytes of Texas. Botanical Research Inst
Worth and Austin College, Sherman, TX.
Diggs, G.M., B.L. Lipscomb, M.D. Reed, and RJ. O'Kennon. 2006. Ill
Neyland, R. 201 1 . A field guide to the ferns and lycophytes
Mississippi. LSU Press, Baton Rouge.
Thieret, J.W. 1980. Louisiana ferns and fern allies. Lafayette Nat. Hist. Mus., Laf
Thomas, R.D. and C.M. Allen. 1993. Atlas of the vascular flora of Louisiana, V
L NRCS. 201 2.The PLANTS Database (httpy/plants.usda.gov, 1 5 June 201 2). National Plant Data Team, Greensboro,
NC 27401-4901 USA.
J.Bot.Res.li«t.Texas7(1):S07.3
508
Journal of the Botanical Research Institute of Texas 7(1)
JOURNAL NOTICE
Sabeeha S. Merchant, Winslow R. Briggs, and Donald Ort, eds. 2013 Gun). Annual Review of Plant Biology,
Volume 64. (ISSN: 1543-5008; ISBN; 978-0-8243-0664-9, hbk). Annual Reviews, Inc., 4139 El Camino
Way, RO. Box 10139, Palo Alto, California 94303, U.S.A. (Orders: www.AnnualReviews.org, service®
annualreviews.org, 1-800-523-8635, 1-650-493-4400). $97.00 indiv., 885 pp., 7 Vs" x 9 Ys".
l Chapman, Sandra Knapp, and Cathie Martin
,sforPlai
^typing — Fabio Fiorani and Ulrich Schurr
>me Engineering with Sequence-Specific 1
jster, Brighter: Advances in Optical Imaging of Living Plant Cells—
me Cytoplasmic Signaling— Jon Hughes
ptor Signaling Networks in Plant Responses to Shade— JorgeJ, Casal
ated Lipid Peroxidation and RES-Activated Signaling— Edward E. F
27. NetwoA Analysis of the MVA and MEP Pathways for Isoprenoid Synthesis— Eva Vranovd, Diana Coman, and Wilhelm Gruissem
28. Toward Cool C^ Crops— Stephen P. Long and Ashley K. Spence
29. The Spatial Organization of Metabolism Within the Plant Cell— Lee J. Sweedove and Alisdair R. Femie
Cumulative Index of Chapter Titles, Volumes 35-64
NOTES ON THE IDENTIFICATION AND DISTRIBUTION
OF THE SPECIES OF THE GENUS GALIUM (RUBIACEAE) IN LOUISIANA
Charles M. Allen
Colorado State University, Fort Polk Station 1697 23rd Sr.
Fort Polk, Louisiana 71459, USA
CharlesMAIIen l.ctr@mail.mil
RESUMEN
lave de las especies del genero Galiur
Twelve species of the genus Galium (Rubiaceae) are listed for Louisiana in the Plants Database (USDA, NRCS
2012) and by MacRoberts (1984) while 11 species are listed by Thomas and Allen (1998). Specimens of Galium
collected in Louisiana were examined and annotated from the following herbaria; Louisiana State University,
Baton Rouge (LSU); Louisiana State University in Shreveport (LSUS); Louisiana Tech University, Ruston
(LTU); University of Louisiana at Lafayette (LAP); and University of Louisiana at Monroe (NLU). The in-state
distributionisbasedon these data as wellastheworkofThomasandAllen(1998).Thekeys to species, descrip-
tions, and habitat information are based on field observations, herbarium specimens, and published works
including Correll and Johnston (1970), Diggs et al (1999) Lipscomb and Nesom (2007), Radford et al (1968),
and Wunderlin (1998). The current list of Galium species in Louisiana numbers 12 and includes three intro-
ducedannuaK ^o nativ^^^^^ Louisiana flora: Galium anglicum Huds., G. asprellum Michx.,
and G divaricatum Pourr. ex Lam. The following Galium taxa are excluded from the Louisiana Flora; G. obtu-
sum Bigelow, G. trifidum L., and G. orizabense Hemsl. Galium obtusum Bigelow was reported from a number of
parishes by Thomas and Allen (1998) but all herbarium specimens examined were G. tmctor^^^ three
petals only.Nospecimens with four petals were seen as reported forG.obtusum by Puff (1977). Thomas^^^^
Allen (1998) reported Galium trifidum L. from Orleans and West Feliciana
tified G. trifiorum Michx. Galium orizabense Hemsl. is reported for Louisiana by USDA NRCS °
Thomas and Allen (1984), but this pubhcatio ' ' "
G. orizabense from Louisiana were found.
n does not include the genus Galium. No herbarium specim
= GALIUM IN LOUISIANA
•Bot. Res. Inst. Texas 7(1): 509 -5
510
1. Galium anglicum Huds. (G. parisiense L. var. trichocarpum Tausch) is an introduced annual. It is known
from only one location in Louisiana. Bossier Parish: Median and roadbank of 1-20 at U.80 over-pass just W of
Rest Area, E of Filmore-HaughtonExitSec. 10, T18N RllW, Thomas 156334, 11 May 1998 (NLU). This collec-
tion is the first for Louisiana. Lipscomb and Nesom (2007) report this species from AL, AR, GA, MO, NC, OK,
SC,TN,TX,VA, andWV.
2. Galium aparine L. is a native annual that is common and widespread in the state in roadside ditches, lawns,
vacant lots and other disturbed areas and is reported from all 64 parishes (Fig. 1). It is reported from all 48
conterminous states and AK in the US and most of Canada (USDA NRCS 2012).
3. Galium asprellum Michx. is a native (but perhaps introduced in Louisiana) perennial. The collection data
for this species in Louisiana are; Caddo Parish: along railroad tracks near center of Kansas City Southern Rail-
road Yard W of La 173, E of Blanchard, sec 19 T18N R14W, Thomas 65094, 29 May 1979 (NLU). This is the 6rst
collection for Louisiana and possibly represents a one-time introduction that did not persist. This species is a
northern species and is reported from MO, TN, NC, and all states to the north of these except for KY (USDA,
NRCS 2012). It is also reported from New Brunswick, Newfoundland, Ontario, and Quebec in Canada.
4. Galium circaezens Michx. is a native perennial of well-drained forests mostly along streams in the pine
regions of the state (Fig. 2). It is reported from all the eastern states in the US west to TX, OK, KS, and NE (ab-
sent from ND and SD). It is also reported from Ontario and Quebec in Canada (USDA, NRCS 2012)
5. GaUum divaricatum Pourr. ex Lam. is an introduced annual. The collection data are Caddo Parish: along
railroad tracks near North Lakeshore Drive west of La 173 SWEof Blanchard Sec. 19, T18N R14W, Thomas
83085, 17 Apr 1983 (NLU); and at the same location; Thomas 99537, 26 Apr 1987 (NLU) and Thomas 88323, 28
Apr 1984 (NLU). Along railroad tracks in Kansas City Southern Yard west of La 169, south of Blanchard at
North Lakeshore Drive Sec. 19, T19N R14W, Thomas 76697, 21 May 1981 (NLU). These collections are the brsl
for Louisiana. Lipscomb and Nesom (2007) report this species from AL, IN, KY, and TX. It is reported from AT,
AR, CA, GA, IN, KY, LA, MO, MS, NC, OR, TN, VA, VT, and WV by USDA NRCS (2012).
6. GaUum hispidulum Michx. is a native perennial of sandy, well-drained soils. It is reported from three par-
ishes in SE and NW Louisiana (Fig. 3). It is an Atlantic and Gulf coastal plain species with reports also from AL,
7. Galium parisiense L. is an introduced annual. It is recorded from roadsides and other disturbed areas
mostly in western Louisiana (Fig. 4). Lipscomb and Nesom (2007) report this species from AL, IN, KY, M .
and TN. USDA NRCS (2012) reports it from CA, OR, and WA in the US and BC in Canada.
8. Galium pilosum Ait. is a native perennial that is widespread in Louisiana. Its habitats include pine for^'
hardwood forests, prairies, and the northern portion of the coastal marsh but is absent from the large fl
plain areas along the southern Mississippi River and lower coastal marsh (Fig. 3). It is also reported from
Allen, Galium in Louisiana
512
11. Galium uniflorum Michx. is a native perennial. It is a plant of hardwood forests along streams in fahty
well drained areas within the pine regions of the state (Fig. 8). In the US, it is restricted to the coastal plain area
with reports from AL, AR, FL, GA, LA, MS, NC, SC, TX, and VA (USDA NRCS 2012)
12. Galium virgatum Nutt, is a native annual. It is reported mostly from prairie areas in west central Louisiana
(Fig. 9). This species is considered to be rare in Louisiana (S2) (Louisiana Natural Heritage Program 2012). In
the United States, it is also reported from AL, AR, IL, KS, MO, MS, OK, SC, TN, and TX (USDA NRCS 2012).
ACKNOWLEDGMENTS
I greatly appreciate the detailed reviews by Michael MacRoberts and an anonymous reviewer.
REFERENCES
CORRELL, D. S. AND M. C. JoHNSTON. 1 970. Manual of the vascular plants of Texas. Texas Res. Found., Renner.
Allen, Galium in Louisiana
513
Diggs, G.M., Jr., B.L. Lipscomb, and RJ. O'Kennon. 1999. Shinners & Mahler's illustrated flora of north central Texas. Sida, Bot.
Misc. 16. Botanical Research Institute of Texas, Fort Worth.
Lipscomb, B.L. and G.L. Nesom. 2007. Galium anglicum (Rubiaceae) new for Texas and notes on the taxonomy of the G.
parisiense/divaricatum complex. J. Bot. Res. Inst. Texas 1 : 1 269-1 276.
Louisiana Natural Heritage Program. 2011. Louisiana Dept. Wildlife & Fisheries, Natural Heritage Program, Baton Rouge.
Available http://www.wlf.louisiana.gov/wildlife/rare-plant-species. (Accessed June 8, 201 2).
MacRobertS, D.T. 1 984. The vascular plants of Louisiana; an annotated checklist and bibliography of the vascular plants
reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci., Louisiana State University in Shreveport, Num-
PuFF, C. 1977. The Galium obtusum group {Galium sect. Aparinoides, Rubiaceae). Bull.Torrey Bot. Club 104:202-208.
Radford, A.E., H.E. Ahles, and C.R. Bell. 1968. Manual of the vascular flora of the Carolinas. Univ. North Carolina Press,
Chapel Hill.
Thomas, R.D. and C.M. Allen. 1984. A preliminary checklist of the pteridosperms, gymnosperms, and monocotyledons of
Louisiana. Contr. Herb. Northeast Louisiana University, No. 4
Thomas, R.D. and C.M. Allen. 1 998. Atlas of the vascular flora of Louisiana, Vol. 3: Dicotyledons Fabaceae-Zygophyllaceae.
Louisiana Department of Wildlife and Fisheries, Baton Rouge.
USDA, NRCS. 201 2. The PLANTS Database (http-y/plants.usda.gov, 4 June 2012). National Plant Data Team, Greensboro,
NC 27401 -4901 USA.
Wunderlin, R.P. 1 998. Guide to the vascular plants of Florida. University Press of Florida, Gainesville.
514
se Menges, Editor, maintained the photo
ichpage.
many plants included are native to the
from other countries. Included are non-
Many are weedy, with small flowers, not usually
NEW BOOK ANNOUNCEMENT
Helen Hamilton and Gustavus Hall. 2013. Wildflowers & Grasses of Virginia’s Coastal Plain. (ISBN: 978-1-
889878-41-6, flexbound). Sida, Botanical Miscellany 40. Botanical Research Institute of Texas Press
1700 University Drive, Fort Worth, Texas 76107-3400, U.S.A. (Orders: www.hritpress.org, 1-817-332-
4441). $24.95, 288 pp., 6" X 9".
From articles written and published by the first author in newspapers over the past 7 years, a book has been created about
Virginia’s Coastal Plain wildflowers and grasses. The second author Dr. Gustavus Hall, Professor Emeritus College of
William and Mary, rewrote portions of the text to ensure botanical accun
inventory, selected appropriate images for each page, and designed the laj
Sponsored by the John Clayton Chapter of the Virginia Native Plan
Coastal Plain of Virginia; some are introduced from other areas in the I
woody plants often seen along roadsides, in meadows, gard
seen in field guides. Also included are grasses commonly seen in Virginia’s Cos
The plants included here occur in most counties of the Coastal Plain of Virginia, and some may be found throughout
the Atlantic and Gulf coastal plains from Cape Cod to Mexico. Originally, the book was conceived to include only plants
native to the Coastal Plain. In decades of field work, the authors have observed very conspicuous non-native (introduced)
plants displacing natives in many locations. These familiar, introduced plants are in the book, to help users distinguish
desirable native plants from unwanted species.
The plants are arranged in the book by flower color (white, yellow, orange, red, pink, blue, violet, green, brown), in-
dicated by a colored rectangle on the upper edge of the page. The grasses and grass-like plants (tan rectangle) are in the last
section of the book. Within each color group the plants are arranged alphabetically by families. Photographs on each page
show the most prominent feature of each plant, usually the flower.
Text follows the photographs with user-friendly descriptions of the plant’s characteristics, habitat, range and growing
conditions, and interesting facts about uses in folk medicine, by the Native Americans, and origins of the plant’s names.
All photographs were contributed by members of local chapters of the Virginia Native Plant Society: Ellis Squires
from the Northern Neck Chapter; and from the local John Clayton Chapter, Teta Kain, Seig Kopinitz, Louise Menges,
Kathi Mestayer, Phillip Merritt, and Jan Newton; and Felice Bond from the Historic Rivers Chapter of the Virginia Master
Dr. Donna Ware, retired Herbarium Curator, College of William and Mary, has written the Preface; the book was
peer-reviewed by Dr. John Hayden, Botanist, University of Richmond; and Bland Crowder, Editor of the Flora of Virginia,
has done the copy editing.
Additional information about the plants can be found in the “Further Read-
ing” section of the book which follows the list of references the authors have used
within. Flora of Virginia (2012) is the definitive resource for technical botanical
e than a field guide
“Wildflowers and Grasses of Virginia’s Coastal Plain is
to the region. The scientific accuracy of the volume
and idiosyncratic text which is woven throughout. This style of writing allows
the reader to painlessly obtain a well-rounded education in taxonomy and species
relationships, the derivation of scientific names, uses of plants by humans, quirks
of plant habitats and distributions, and innumerable fascinating and useful facts
the book also ventures into territory often neglected by popular botanicaHexts!
:r will become well-informed and enjoy the process!”— Johnny
nist, Virginia Natural Heritage
1, Richmond, Virginia.
J. Bot. Res. Inst Texas 7(1); 514. 2013
SYNOPSIS OF THE GENUS STYLISMA (CONVOLVULACEAE) IN LOUISIANA
Charles M. Allen
Colorado State University
Fort Polk Station 1697 23rd St.
Fort Polk, Louisiana 71459, U.S.A.
Charles.M.AIIen l.ctr@mail.mil
ber of species includes three ni
ABSTRACT
e perennial vines: S. aquatica (Walt.) Raf., 5. humistrata (Walt.) Chapm., a
s, Stylisma patens (Desr.) Myint at ' " ”
RESUMEN
The genus Stylisma (Convolvulaceae) in Louisiana includes native perennial vine species with five reported by
the USDA, NRCS (2012), MacRoberts (1984) and Thomas and Allen (1998). The specimens of Stylisma from
Louisiana were examined and annotated from the following herbaria; Louisiana State University, Baton Rouge
(LSU), Louisiana State University in Shreveport (LSUS), Louisiana Tech University, Ruston (LTU), University
of Louisiana at Lafayette (LAF), and University of Louisiana at Monroe (NLU). The in-state distribution is
based on these data and the work of Thomas and Allen (1998). The report for S. patens was based on a misiden-
tified specimen of S. humistrata and the two reports of S. villosa were based on misidentified specimens of S.
aquatica. Consequently, these two species should be excluded from the Louisiana flora. Stylisma villosa has a
white corolla and villous filaments while S. aquatica exhibits a lavender, pink or purplish red corolla and gla-
brous filaments (Myint 1966). The range of S. villosa includes FL, GA, MS and TX with S. aquatica reported
from AL, AR, FL, GA, LA, MS, NC, SC, and TX (Myint 1966; USDA NRCS 2012). Stylisma patens has narrower
leaves (less than 1.2 cm wide) and mostly solitary flowers compared to the wider leaves (greater than 1.2 cm)
and flowers, mostly in cymes of three, of S. humistrata (Myint 1966). The range of S. patens includes Al, FL, GA,
MS, NC, and SC while S. humistrata is reported from AL, AR, FL, GA, LA, MS, NC, SC, TN, TX, and VA (Myint
1966; USDA NRCS 2012). The leaves of plants within the genus Stylisma are alternate, simple, entire, and with
pinnate major veins. The inflorescence is an axillary corymb. The perfect regular flowers have five imbricate
sepals, five fused petals with the corolla campanulate to funnelform, and five epipetalous stamens. The ovary
is superior and the fruit is a capsule with four seeds.
KEY TO THE SPECIES OF STYLISMA IN LOUISIANA
1- Stylisma aquatica (Walt.) Raf. Leaf blades oblong to elliptic, short densely pubescent, sepals pubescent,
corolla lavender, style deeply lobed to % or more of length, filaments glabrous. An infrequent plant of pine
flatwoods and savannahs in the southwest and southeast part of the state (Fig. 1). May-July (Bonamia aquatica
(M^afr.) A. Gray, Bonamia michauxii (Fern. & Schub.) K.A. Wilson, Breweria aquatica (Walt.) A. Gray, Breweria
'"ichauxii Fern. &Schub.).
516
Michael MacRoberts
2. Stylisma humistrata (Walt.) Chapm. Leaf blades
oblong to elliptic, sparsely pubescent, sepals gla-
brous, corolla white, style not lobed, filaments pubes-
cent. The most common and widespread of the Loui-
siana species of this genus in sandy pine forests in the
pine regions of the state (Fig. 2). May to October
(Bonamia humistrata (Walt.) A. Gray, Breweria hu-
mistrata (Walt.) A. Gray.
3. Stylisma pickeringii (Torr. ex M.A. Curtis) A.
Gray var. pattersonii (Fern. 61 Schub.) Myint. Leaf
blades linear, sparsely pubescent, sepals pubescent,
corolla white, style not lobed, filaments glabrous. An
infrequent species of very sandy areas in western and
northwestern Louisiana (Fig. 3). May-August (Brew-
eria pickeringii (Torr. ex M.A. Curtis) A. Gray var.
pattersonii Fern. & Schub., S. pattersonii (Fern. &
Schub.) G.N. Jones).
ACKNOWLEDGMENTS
nd Andrew McDonald provided helpful comi
Myint, T. 1966. Revision of the genus Sty/isma (Convolvulaceae). Brittonia 18:97-1 17.
Thomas, R.D. and C.M. Allen. 1 998. Atlas of the vascular flora of Louisiana, Vol. 3: Dicotyledons Fabaceae-Zygophyllaceae.
Louisiana Department of Wildlife and Fisheries, Baton Rouge.
USDA, NRCS. 201 2. The PLANTS Database (http://plants.usc
NC 27401 -4901 USA.
CONFIRMATION OF KARIBA-WEED, SALVINIA MOLESTA (SALVINIACEAE)
IN THE CALCASIEU RIVER BASIN, LOUISIANA
Ray Neyland and Jennifer Bushnell
Department of Biology and Health Sciences
McNeese State University
Lake Charles, Louisiana 70609, U.SA.
As described by Neyland (2011),
appears
andele-
n shore of Lake
2 molesta Mitchell, kariba-weed, is an introduced aquatic fern with
^uatic, and up to 4 cm long with rows of stiff hairs. Each hair has the
4 an eggbeater. The submerged leaves are dark brown with root-like fibers. Sporocarps are formed
in chains but rarely produce fertile spores. The species inhabits lakes, ponds, marshes, sluggish ri ^
streams along the southern tier of states from CA to NC. Plants are sensitive to freezing temperature
vated salinity.
Native to southeastern Brazil, Salvinia molesta has become a serious threat to aquatic systems in t e
warm-temperate regions of the United States Qacono et al. 2001). Exponential vegetative growth potential
(Mitchell & Tur 1975) and tolerance to environmental stress (Whitman and Room 1991) allows this species to
become a noxious weed in much of its range. On July 15, 2012, large numbers of individuals of Salvinia molesta
were observed floating down the Calcasieu River adjacent to Shell Beach Road along the s.
Charles in Calcasieu Parish, LA. This is the first recorded sighting of this species in the Parish.
On July 16, 2012, large numbers of S. molesta were obsen
along River Road in northern Lake Charles just south of the saltwater barrier dan
great abundance along the northern shore of Lake Charles on July 17, 2012. Specim
locations, including GPS coordinates, were collected and are housed in
urn (MCN). , , ^
Because this is the first recorded sighting of Salvinia molesta in Calcasieu J
prised at its great abundance. The stretch of the Calcasieu Riv
brackish. Sensitive to salinity and flowing \^
were being flushed from a location or locati , , . r n
Louisiana during the fi,s. two weeks of July is the probable cause for this llushing. For example ramfall re-
corded alUkc Charles from July 1-15 was 33.86 cm (13.33") (National Weather Service Website 2012L , . .
This event generates an important question: where is the source population
tnolesta? The Calcasieu River originate?
:rn side of the Calcasieu Riv
e observed i?
e not resident along this stretch of the riv(
r of Lake Charles. Unusually heavy rainfall in m
518 Journal of the Botanical Research Institute of Texas 7(1)
through Rapides, Allen, Jefferson Davis and Calcasieu parishes until it empties into the Gulf of Mexico in Cam-
eron Parish. Of those north of Calcasieu Parish, only in Jefferson Davis Parish, adjacent to Calcasieu Parish,
has S. molesta been documented (USDA, NRCS Website 2012). From information supplied by Mark Garland!
from USDA, NRCS, a population of S. molesta was observed by Vanessa Morgan, research assistant at Portland
State University, in /arm ponds near the vicinity of Fenton, Louisiana, on November 6, 2004. This was con-
firmed by Scott Schales, biologist at the Louisiana Wildlife and Fisheries Department. However, because no
specimen from this observation was collected, the exact location of the reported farm ponds is unknown. Un-
less these farm ponds drain into the Calcasieu River, either by Bayou Serpent or Little Bayou, these populations
of the recent occurrence in Calcasieu River.
of Allen Native Ventures, (pers. comm. 2012) reported that he has observed Salvinia
molesta in Fullerton Lake, north of Pitkin in Vernon Parish. Fullerton Lake drains into Sixmile Creek which
merges with the Whiskey Chitto, a tributary of the Calcasieu River. No specimen and, therefore, no specific
collection information exists for this sighting. Additionally, no specimens or records of S. molesta from Vernon,
Rapides, Allen, Jefferson Davis or Calcasieu parishes were found in a herbarium search from Louisiana State
University (LSU), University of Louisiana at Monroe (NLU), University of Louisiana Lafayette (LAF) or Mc-
Neese State University.
With the return of drier weather, observations on July 19, 2012 from the original collection sites revealed
that individuals of Salvinia molesta were no longer floating down the river and many individuals had died after
being washed up along the banks. However with the return of heavy rainfall from July 20-22, a new flush of S.
molesta occurred on the Calcasieu River. Additional specimens from this second flush were collected just up-
river from the saltwater barrier dam north of Lake Charles and under the I-IO Bridge near Westlake, LA.
The origin of the S. molesta observed in the Calcasieu River remains unclear. Numerous habitats upriver
from the saltwater barrier dam in Lake Charles could support populations of S. molesta. However, it is clear that
S. molesta is firmly established at some location or locations in the Calcasieu River basin. Because of the nega-
tive ecological and economic impacts that can accrue from this noxious species, we intend to closely monitor
the situation and will attempt to locate the original source of infestation.
ACKNOWLEDGMENTS
We are greatly to Michael MacRoberts (LSUS) and an anonymous reviewer for their helpful reviews.
Jacono, C.C., T.R. Davern, and D. Center. 2001 . The adventive status of Salvinia minima and S. molesta in the s
United States and the related distribution of the weevil Cyrtobagous salviniae. 2001 . Castanea f
Mitchell, D.S. and N.M. Tur. 1975. The rate of growth of Salvinia molesta (S. auriculata Auct.) in lal
conditions.! Appl. Ecol. 12:213-225.
National Weather Service Website. 201 2. http://www.srh.noaa.gov/
Neyland, R. 201 1 . a field guide to the ferns and lycophytes of Louisiana. Louisiana State University Press, Baton Rouge.
USDA, NRCS Website. 2012. The PLANTS Database {http://plants.usda.gov) 18 July 2012. National Plant Data Center,
Baton Rouge, Louisiana 70874-4490 USA.
Whitman J.B. and P.M. Room.1 991 . Temperatures lethal to Salvinia molesta Mitchell. Aquatic Bot. 40:27-35.
A QUANTITATIVE STUDY OF THE VEGETATION SURROUNDING
A XANTHORHIZA SIMPLICISSIMA (RANUNCULACEAE) POPULATION
AT FORT POLK IN WEST CENTRAL LOUISIANA
Charles Allen and Rachel Erwin
Colorado State University, Fort Polk
1645 23rd Street
Fort Polk, Louisiana 71459, U.S.A.
Jeff McMillian and Joe McMillian
ABSTRACT
RESUMEN
Yellow root (Xanthorhiza simplicissima Marsh.) is a small deciduous shrub in the Ranunculaceae family. It is
found throughout the eastern portion of the United States, ranging from Texas to Maine, with the exception of
Vermont and New Hampshire (USDA NRCS 2012). Yellow root has been found in two of the three states that
border Louisiana: Texas and Mississippi. The current county records for yellow root in East Texas include San
Jacinto, Newton, and Jasper. In Mississippi, yellow root is widely distributed and has been found in Tishom-
ingo and Itawamba counties in the northeast, Lafayette County in the north central, and Lauderdale, Jones,
and Forrest counties in the southeast (USDA NRCS 2012). This species is demonstrably secure globally, though
it may be quite rare in parts of its range, especially at the periphery, and has a global ranking of 5 (NatureServe
2012). It is considered imperiled in Texas (S2) and critically imperiled in Florida and Louisiana (SI).
Yellow root was first reported in Louisiana in 1987 (Allen et al. 1987) from Vernon Parish, the only loca-
tion in the state. In Vernon Parish, yellow root is located on US Forest Service/Fort Polk land in two locations
that are ca 6000m apart. The larger population is found on an unnamed creek that drains generally from east
to west and empties into the Ouiska Chitto Creek, just north of Lookout Road. At this location, the down-
stream distribution of yellow root apparently ends about 100m from where the tributary empties into the larger
Ouiska Chitto Creek. With the flooding that can occur along this small stream, yellow root is probably distrib-
uted downstream very readily. In theory, it should be found farther downstream and even along the banks of
the Ouiska Chitto Creek. Several searches downstream from this population on the banks of the tributary and
continuing along the Ouiska Chitto Creek have not yielded any populations of yellow root. The second and
much smaller population is south of Lookout Road on an unnamed creek that drains from east to west and
empties into Drake’s Creek.
The habitat reports for this species include shaded stream banks, moist woods, thickets, and rocky ledges
(Parfitt 1997). The Louisiana Natural Heritage Program reports that small stream forests are the preferred
520
Journal of the Botanical Research Institute of Texas 7(1j
habitat for this species (Louisiana Natural Heritage Program 2011). In the Carolinas, the plant has been re-
ported on shaded stream banks (Radford et al. 1968). Godfrey and Wooten (1981) described its habitat as river
and stream banks, moist thickets, and springy places, usually shaded. The objectives of this study were to
document the species that are associated with yellow root, describe the plant community where yellow root
grows, and compare and contrast the plant community and species association downstream in habitats where
yellow root has not been found.
Each Louisiana population is located in a baygall; the larger population within one of the baygalls studied
by Allen et al. (2004). The soil along the stream bank where yellow root grows is mainly the Guyton-luka com-
plex, frequently flooded (Soil Survey Division 2003).
METHODS
The larger population of yellow root was used for sampling. The distance (482 meters) along the creek from the
farthest upstream to the farthest downstream location was measured and recorded. A 482 meter long macro-
plot was created to include the known range of yellow root along the tributary. Using a random number genera-
tor, fifty plot centers out of the 482 points were selected for sampling. Beginning from the last downstream
yellow root location, a fifty meter macroplot was created downstream from that point and ten sampling loca-
tions were randomly selected out of the fifty potential points. All sixty (50 upstream and 10 downstream)
oriented perpendicular to the stream. The plant categories sampled included herbaceous plants, woody vines,
shrubs and saplings (woody non-vine species shorter than 6 ft = 1.83 m), and trees and shrubs (woody non-vine
species taller than 6 ft = L83m). During the sampling period, the number of stems in a sample for each species
were counted and recorded. For herbaceous plants, woody ^es, and shrubs and saplings, cover was deter-
mined by measuring the area occupied by the plant(s). The cover percent was calculated by multiplying the
area times the density and then dividing by the area of the sample (200,000 cm^). The cover was converted to a
percent by multiplying by 100. For the trees and shrubs taller than L83m, the dbh was measured at the stan-
dard L37m height using a diameter tape and recorded to the nearest 0.1 cm. The woody plant data in the sixty
samples were recorded in the fall only while the herbaceous plant data were recorded in the fall and again in the
spring. To facilitate relocation of plots, a metal pin was left in the center of the stream until the re-sampling was
completed.
All data were entered into a Microsoft Excel spread sheet for storage and calculation of variables. The
mean diversity (richness-species per sample) and mean density (stems per sample) were calculated for the
samples in each group (upstream or downstream) for all plant categories while the mean dbh was calculated
for trees and shrubs and mean cover percent was calculated for herbaceous plants, woody vines, and shrubs
and saplings. The frequency and mean density were calculated for each species in a sample group. The mean
dbh was calculated for tree and shrub species and the mean cover percent for all other species in a sample
group. The relative values for each of these variables (frequency, mean density, mean dbh, and mean cover
percent) were calculated by dividing the value for a species by the total for all species within the sample group
and plant category. Each value was converted to a percent by multiplying by 100 and the sum of these relative
values was used to calculate the importance value.
RESULTS
The number of species in the upstream area with yellow root totaled 122 and contained 24 trees and shrubs, 50
shrubs and saplings, 13 woody vines, and 57 herbaceous plants (Table 1). The number of species in the down-
stream area without yellow root totaled 69 with 15 trees and shrubs, 34 shrubs and saplings, 9 woody vines;
and 25 herbaceous plants. In the upstream area, the mean number of species per sample ranged from 3.44 for
trees and shrubs to 14.92 for shrubs and saplings while in the downstream study area the mean number of spe-
cies per sample ranged from 4.10 for trees and shrubs to 14.40 for shrubs and saplings. In the upstream area
where yellow root was present, the mean density ranged from 7.12 stems for trees and shrubs to 92.10 for
Allen et al., Vegetation study of Xanthorhiz.
523
524
Journal of the Botanical Research Institute of Texas 7(1)
Polk,LouisSna/ ty p , p y p j
Bignonia capreolata
:scending ii
group of species in the downstream area are in Table 10. The species are arra:
value in all eight tables.
The two herbaceous species with the highest importance value in both a
(L.) Yates and Mitchella repens L. but the importance value for Chasmanthium laxum (L.)Yates was much higher
in the downstream area (93.14) than in the upstream area(39.53) (Tables 3 and 4). The third species in the area
of yellow root was Dichanthelium commutatum (Schult.) Gould and in the area just downstream was Sderia
oligantha Michx. The top two species in importance value among the woody vines in both areas were Vitis ro-
tundifolia Michx. and Smilax glauca Walter but the importance value for Vitis rotundifolia in the upstream area
was much higher (93.28) compared to a value of 48.39 in the downstream area (Tables 5 and 6). The third spe-
cies in importance value in the area of yellow root was Gelsemium sempervirens (L.) W.T. Alton and in the area
just downstream from yellow root was Smilax rotundifolia L. In the upstream area, the shrub and sapling spe-
cies with the highest important value was Lyonia lucida (Lam.) K. Koch and was followed by Ilex coriacea
(Pursh) Chapm. and Vaccinium elliottn Chapm. (Table 7). In the downstream area, the shrub and sapling spe-
cies with the highest importance value was Vacdnium elliottii Chapm. and was followed by Halesia diptera Ellis
Hypericum hypericoides (L.) Crantz, and Symplocos tinctoria (L.) L’Her. (Table 8). The tree and shrub species
with the highest importance value in the area of yellow root was Ilex coriacea (Pursh) Chapm. , followed by
Magnolia virginiana L., Nyssa bifloraWalter, and Vaccinium elliottii Chapm. (Table 9). The tree and shrub species
with the highest importance value in the area downstream from yellow root was Vaccinium elliottii Chapm.,
followed by Acer rubrum L. and Nyssa biflora Walter (Table 10).
Thirty-three herbaceous species were found only upstream in the area containing yellow root (Table 3).
Two herbaceous species were found only downstream in the area without yellow root (Table 4). All other her-
baceous species were found in both upstream and downstream samples (Tables 3 and 4). Four woody vine
species were found only upstream with yellow root (Table 5). No vine species were unique to the downstream
samples. All other woody vine species were found in both upstream and downstream samples (Tables 5 and 6).
Twenty-two shrub and sapling species were found only upstream with yellow root (Table 7) and six shrub and
sapling species were found only downstream without yellow root (Table 8). All other shrub and sapling species
were found in both upstream and downstream samples (Tables 7 and 8). Twelve tree and shrub species were
found only upstream with yellow root (Table 9) and three trees and shrub species were found only downstream
(Table 10). All other tree and shrub species were found in both upstream and downstream samples (Tables 9
DISCUSSION
The yellow root habitat in Louisiana is best described as the bank and natural levee of a baygall stream with the
vegetation being typical of such a stream. We found the tree canopy vegetation associated with yellow root to
Journal of the Botanical Research Institute of Texas 7(1)
Diggs et al. 2006 and in central Louisiana by MacRoberts et al. 2004. The herbaceous flora associated with yel-
low root in our area is dominated by Chasmanthium laxum (L.) Yates, Mitchella repens L., and Dichanthelium
commutatum (Schult.) Gould; three species that have been noted by the senior author to be shade tolerant. Our
data seem to verify the qualitative reports of the shaded stream bank, small stream bank, and moist woods
habitat for yellow root. Yellow root was found in 19 of the 50 samples and had an importance value of 7.07 out
of 300 and was the 12‘*’ ranked shrub in importance value.
In the downstream samples, the vegetation contains species that are usually associated with less wet or
more mesic to dry sites. Red maple (Acer rubum L.) and southern red oak (Quercus falcata Michx.) are two tree
species with high importance values in the downstream samples and are usually found on more mesic sites. An
herbaceous species (Sderia oligantha Michx.) is usually associated with mesic sites as are the shrub species
Halesia diptera Ellis, Hypericum hypericoides (L.) Crantz, and Symplocos tinctoria (L.) L’Her. The dominance of
these species downstream and the absence of yellow root seem to indicate that yellow root cannot tolerate the
more mesic downstream sites. The big community difference in the downstream samples was the decrease in
herbaceous plants (richness, density, and cover percent). The mean density and cover percent also decreased
Allen et al., Vegetation study of Xanthorhiza simplicissima
527
alue for trees and shrubs in 50 samples in the area of yellow root at Fort Polk, Louisiana.
alue for trees and shrubs in 10 samples just downstream from yellow roof at Fort Polk,
downstream for shrubs and saplings while increasing for woody vines. The lack of yellow root and these
changes in the community seem to indicate a more mesic site created by a larger stream that produces a higher
528
; the first quantitative report on the vegetation surrounding Xanthorhisxi simplicissima Marsh,
round the other populations of Xanthorhiza simplicissima Marsh, throughout its range should
arison with our data so as to get a better idea of the variation, if any, of its habitat.
ACKNOWLEDGMENTS
We thank the reviewers, Michael MacRoberts and an unknown person for their thoughtful suggestions.
REFERENCES
Allen, C.M., H.D. Guillory, C.H. Stagg, S.D. Parris, and R.D. Thoma
Ranunculaceae New to Louisiana. Phytologia 62:5-6.
Allen, C.M., J. Pate, S. Thames, S. Trichell, and LEzell. 2004. Changes in baygall vegetat
west central Louisiana. Sida 21 :41 9-427.
Diggs, G.M., B.L. Lipscomb, M.D. Reed, and R.J. O'Kennon. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26:1-1594.
Godfrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of the southeastern United States. Univ. Georgia Press,
V Root [Xanthorhiza simplicissima Marshall)
n 1986 to 2001 at Fort Polk in
Louisiana Natural Heritage Program. 2011. Louisiana Dept. Wildlife & Fisheries, Natural Heritage Program, Baton Rouge.
Available http://www.wlf.louisiana.gov/wildlife/rare-plant-species. (Accessed June 8, 2012).
MacRoberts, B.R., M.H. MacRoberts, and L.S. Jackson. 2004. Floristics of baygalls in central Louisiana. Phytologia 86:1-22.
NatureServe. 2012. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe,
Arlington, Virginia. Available http://www.natureserve.org/explorer. (Accessed: June 4, 201 2).
Parfitt, B.D. 1 997. Xanthorhiza. In: Flora of North America Editorial Committee. Flora of North America: Volume 3; Mag-
noliophyta: Magnoliidae and Hamamelidae. Oxford Univ. Press, New York.
Radford, A.E., H.E. Ahles, and C.R. Bell. 1968. Manual of the vascular flora of the Carolinas. Univ. North Carolina Press,
Chapel Hill.
Soil Survey Division. 2003. Soil survey of Vernon Parish, Louisiana. United States Department of Agriculture, Natural Re-
USDA, NRCS. 201 2. The PLANTS Database (httpV/plants.usda.gov,4 June 2012). National Plant Data Team, Greensboro,
NC 27401 -4901 USA.
VASCULAR FLORA AND PLANT COMMUNITIES
OF ALLEGHANY COUNTY, NORTH CAROLINA
Derick B. Poindexter^
LW. Carpenter, Jr. Herbarium (BOON)
Appalachian State University, Biology Department
Boone, North Carolina 28608-2027, U.S.A.
RESUMEN
INTRODUCTION
Perhaps the most intense effort to document county floras in North Carolina and South Carolina was during
the production of the regionally renowned Manual oj the Vascular Flora of the Carolinas (Radford et al. 1968).
This manual was the product of many years of botanical exploration within these contiguous states and its
impact was far reaching. It served and continues to serve as a mainstay of taxonomic reference for the flora of
this area, and has been made extensible to neighboring states for various studies. However, there are limita-
tions to how much a single, large-scope work can cover regarding the mammoth task of documenting plant
distributions. Consequently, few if any counties within the range of Radford et al.’s (1968) original manual can
be considered “comprehensively inventoried”. Likewise, since its publication much time has passed. Taxo-
treatment (e.g., Weakley 2011). In light of global climate change, undiscovered new taxa,
edge of vegetation patterns and modern techniques for cataloguing plant diversity data, the
revived need for continued floristic research.
The objectives of this descriptive study were to: 1) document and georeference speci
vascular flora of Alleghany County, North Carolina; 2) describe general plant communities
mens of the known
; 3) analyze the flora
J- Bot. Res. Inst. Texas 7(1); 529 - 574. 201 3
Journal of the Botanical Research Institute of Texas 7(1)
1 context of species richness anti origin (i.e., native vs. multiple exotic categories) of taxa; and 4) make voucher
ata digitally available via a web interface to the botanical and general communities.
Physiography
Alleghany County, North Carolina is located in the northwest corner of the state, between 36.36° and 36.37°N
latitude, and 80.91° and 81.35°W longitude (Fig. 1). The county is the 4th smallest in North Carolina, with a
total area of 610 km^, comprised of 607.7 km^ of land and 2.3 km^ water (United States Census Bureau [USCB]
2011a). It is bordered by Surry County to the east, Wilkes County to the south, Ashe County to the west, and
Grayson County, Virginia to the north (Fig. 2). The southern and eastern boundary of the county is close to the
rim of the Blue Ridge Escarpment and comprises part of the Eastern Continental Divide. This county contains
two major river basins, the New River Basin and the Upper Yadkin River Basin. Most tributaries (e.g.. Little
River, Prathers Creek) in Alleghany County empty into the New River, which ultimately drains into the Gulf of
Mexico via the Ohio and Mississippi Rivers. The tributaries associated with the New River watershed drain ca.
93-95% of the county (Padgett 2011). The far southern and eastern portions of Alleghany are the only excep-
tion, with relatively few tributaries (e.g., Mitchell River headwaters) draining off of the escarpment into the
Upper Yadkin River Basin and eventually emptying into the Atlantic Ocean.
Alleghany County is principally montane. It is located within the Southern Section of the Blue Ridge
Physiographic Province (Fenneman 1938) of the Southern Appalachians (Braun 1950), although a small frac-
tion of the southeastern boundary is situated just below the Blue Ridge Escarpment (in the vicinity of Stone
Mountain State Park) in the Piedmont Upland Physiographic Province (Eenneman 1938). Topographic varia-
tion increases dramatically as the western and northern portions of the county grade into typical high to mid-
elevation mountain ridges that are more characteristic of this region (Fig. 2). This physiognomy of the land-
scape differs somewhat from the highest mountains in northwest North Carolina, which are exemplihed by
the Amphibolite Mountains Macrosite to the west of Alleghany County in adjacent Ashe and Watauga Coun-
ties. Many of the peaks in that area exceed 1500 m in elevation, and possess a prominent northern floristic
component typically associated with the inner, high mountains of the Blue Ridge. In contrast, the highest ele-
vation in Alleghany County is Catherine Knob (1272 m), which is part of a chain of peaks that run diagonally
northeast through the upper third of the county and contains several other notable peaks (Fender Mountain,
1213 m; Cheek Mountain, 1201 m; Twin Oaks Mountain, 1116 m; and Bald Knob, 1109 m). In addition, there
are several outliers to the south and southeast (e.g.. Bullhead Mountain, 1171 m; Bluff Mountain, 1139 m; Ma-
hogany Rock, 1103 m; and Green Mountain, 1018 m) that occur primarily adjacent to the edge of the Blue Ridge
Escarpment (Fig. 2). Though most of the lower elevation areas in the county occur within the New River drain-
age and headwaters of the Mitchell River, the minimum elevation occurs around the foot of Stone Mountain
(475 m) along the Wilkes County border. Ultimately, this difference in geography and topography, as compared
to the predominately higher elevations westward, results in a small reduction in the presence of northern spe-
cies, but in turn adds some lower elevation species with Piedmont affinities to the vegetation of the county.
Braun (1950) broadly classified the vegetation of Southern Appalachians as a part of the Oak-Chestnut
Forest Region, but this area is now more appropriately referred to as the Appalachian Oak Forest Region
(Kiichler 1964; Stephenson 1993) following the demise of the American chestnut. Despite these general re-
gional classifications, vegetation patterns in Alleghany County are largely a consequence of its highly variable
landscape and hydrology that contribute to a wide array of communities including both forested areas and
wetlands.
Climate
The continental climate of the southern Appalachians is temperate, humid mesothermal, lacking a distinct dry
season (precipitation shows only minor fluctuation), cool to warm summers, and mild to cold winters (Trew-
artha & Horn 1980). Alleghany County has a wide range of microclimate variation related to its mountainous
terrain. Due to incomplete data from the Sparta weather station, Alleghany County climate normals (1971-
532
tion consist of long horizontal bodies dominated by mica schist or phyllite (commonly graphitic) containing
garnet and magnetite, interlayered with lesser amounts of biotite-muscovite gneiss and amphibolite. Another
area dominated by amphibolite (some garnet based) and greenstone occupies a northeastern oriented sliver,
equidistant between Cherry Lane and Roaring Gap (Rankin et al. 1972; Epenshade et al. 1975; USGS 2011).
The Ashe Formation occupies a third to nearly half of the county adjacent to and west of the Alligator Back
Formation. It also runs horizontally in a northeast trajectory through the center of Alleghany. This formation
encompasses the Peach Bottom Mountain range, Doughton Mountain, the county seat of Sparta, and Stratford.
This late Precambrian (late Proterozoic) formation is dominated by rocks that are thinly layered and fine-
grained. As mapped by Rankin et al. (1972), Epenshade et al. (1975), and USGS (2011) primary bedrock mate-
rial IS comprised of biotite-muscovite gneiss, with varying amounts of mica schist, phyllite, quartz, feldspar,
amphibolite, and hornblende gneiss. Gneiss layers are most common and often very thick. Several long, nar-
row bands dominated by amphibolite and garnet amphibolite are scattered throughout the formation. These
mafic bedrocks are found in areas from Sparta to Ennice, south of Sparta, from Furches northeast to Stratford,
near Peden in the same trajectory through Amelia to the Virginia, and as another sliver from the northwest
section of the South Fork of the New River, south of Piney Creek and bisecting the northern “loop” of the New
River along the northern border of the county (Rankin et al. 1972; Epenshade et al. 1975; USGS 2011).
The Elk Park Plutonic Group or Suite is the second smallest lithologic assemblage in Alleghany County,
found in the northwest corner, surrounding Piney Creek and the confluence of the North and South Forks of
the New River, along with one section of the New River itself. It is comprised of intermediate Precambrian
(mid-Proterozoic), metamorphic and igneous rocks. The largest, most common body of rocks is referable to
Cranberry Gneiss (biotite granitic gneiss), containing rocks that range from diorite to granite, with quartz
monzonite that often bears biotite. Some hornblende (amphibolite), calc-silicate rock, and marble is present,
and sphene and epidote are common (Rankin et al. 1972; Epenshade et al. 1975; USGS 201 1)
Poindexter, Flora of Alleghany County, North Carolina
rery small area around Stoi
n State Park along
)rimarily of
muscovite, and quartz
1975; USGS 2011).
within the Ashe For-
The Spruce Pine Plutonic Group is res
the western side of the southeast leg of the county. This group ot
Paleozoic (Devonian) granitic substrates including quartz diorite
monzonite, with frequent epidote and local garnet (Rankin et al.
The ultramafic metamorphic rocks of this area occur in small bands or pockets wit
mation. Of these areas, Amelia, Edmonds, and Peden contain the largest consolidated loct
substrate. Likewise, the Edmonds portion extends northeastward into Virginia, constitutin]
bodies of ultramahc rock in the southern Appalachians. These ultramafic rocks are of an undetermined Pre-
cambrian and/or Paleozoic origin and are exceedingly rich in minerals. Areas containing this rock type consist
principally of chlorite-tremolite-magnetite schist with common deposits of either serpentine or talc, and local
occurrences of olivine (Rankin et al. 1972; Epenshade et al. 1975; Scotford & Williams 1983). Such bedrock re-
ring, and thus often produces habitats that harhor rare plants. Elements
^and potassium. Those of Alleghany County belong to two types, the Ed-
netasomatized, with the Edmonds-type rocks being less altered
the Todd-type in the western half (Scotford & Williams 1983).
Soils
The soils of the county are primarily classified as Ultisols, with a few examples of Inceptisols, and one series
with Entisols. Most soils in the county are derived from gneiss and schist, with occasional phylhte and rarely
granite. The soil texture is mainly loam to a fine sandy loam, containing varying amounts of mica. The soils of
Alleghany County are more specifically partitioned into five main units: the Watauga-Chandler-Fanntn asso-
ciation, Chester-Ashe association. Porters association, Clifton association, and Stony steep land assoctatton
(Brewer etal. 1973).
The Watauga-Chandler-Fannin association is widely distributed, occurring in large areas in the north-
534
Journal of the Botanical Research Institute of Texas 7(1)
least between Amelia and Twin Oaks, and continuing to the
Virginia border. Other areas dominated by this association occur in the southwest portion of the county sur-
rounding Laurel Springs and Citron, and then in a northeast trajectory from Whitehead up through Glade
Valley to the Blue Ridge Parkway. These soils are well-drained to excessively drained, occurring in areas that
are rolling to very steep in topography, particularly along upland side slopes and narrow ridge tops. They con-
tain a large micaceous component and occupy roughly 39% of the county. This association is near equally di-
vided into forested areas and farmland used for pasture and crop cultivation. Steep topography is the primary
disadvantage of this association, making these soils prone to erosion (Brewer et al. 1973).
The Chester-Ashe association is the second most prominent series of soils in the county. They occupy ar-
eas primarily in the eastern half of Alleghany County, but with one large region in the northwest corner of Al-
leghany County west of Amelia and encompassing most of the Piney Creek community. Other areas domi-
nated by this association occur from Whitehead through Sparta to the Virginia border, around the Eunice
community, and south surrounding Cherry Lane and Roaring Gap. These soils are generally classified as per-
meable to excessively permeable, and often have surficial bedrock. They occur on gentle to steep slopes, along
broad ridgetops and upland side slopes. This association occupies approximately 36% of the county, and is
mostly used for cultivation (only a small portion in forested areas). Like the previous association, soils of this
group are subject to erosion (Brewer et al. 1973).
The Porters association includes soils primarily in west-central section of the county, west of Whitehead
and south of Twin Oaks, beginning southwest from NC 113 and running northeast to the Virginia State line,
encompassing Fender Mountain and the Peach Bottom Mountain range. A second smaller area dominated by
this association occurs south of Glade Valley along the northern side of Bullhead Mountain. Soils of the Porters
association tend to be well-drained and occur on strong to very steep side slopes and narrow ridges of the
higher elevation areas in the county. This association occupies about 12% of the land in Alleghany County,
much of which is forested. The rugged topography dominated by this association makes agriculture difficult in
these areas, coupled with bedrock that is near the soil surface. Likewise, cultivation within this association is
not practical due to a high probability of erosion (Brewer et al. 1973).
The Clifton association is comprised of soils mainly in the north and northwestern portions of the coun-
ty. They occur in small bands, specifically around Peden northeast to Stratford, more or less parallel to US 221
and terminating just south of NC 93. This band continues along this trajectory on the north side of NC 93 until
reaching the Virginia border. The only other area where this association is found is around the Edwards Cross-
roads community and northeastward. Soils of this series are relatively well-drained. The Clifton association is
found in rolling to somewhat steep sites, along rather broad ridgelines and upland slopes. These soils are the
least common in the county, occupying about 6% of the land area with half in forest and half in cultivation.
Like the previous associations, these soils have a limited farming capacity due to steep topography and surfi-
cial bedrock (Brewer et al. 1973).
The Stony steep land association is found mainly along the rim of the Blue Ridge Escarpment, in the ex-
treme southern, southeastern, and northeastern portions of the county. These areas are adjacent to Wilkes and
Surry Counties. Soils of this association are very rocky with exposed bedrock, and generally occur on very
narrow ridgetops and steep side slopes, with especially narrow drainage ways. Most of this land belongs to the
Blue Ridge Parkway, and is thus relatively undisturbed. Likewise, the steep topography and unsuitable soils
make it non-conducive for agriculture. Approximately 7% of the county is occupied by this association, with
History and Special Features
The first known inhabitants of Alleghany County were Native Americans. Relictual evidence and other arti-
facts suggests that these cultures were present near the beginning of the Hypsithermal period, at the end of last
Pleistocene Glaciation, approximately 10,000 years ago (Alleghany Historical-Genealogical Society [AHGS]
1983). Three major divisions are recognized based on the progressively more advanced implements found in
the county, that ultimately resulted in a transition from a nomadic (seasonal hunting migration) lifestyle to
more sedentary (farming infused) society. These time frames correspond to the Paleo Period (ca. 10,000 B.C.),
the Archaic Period (8,000 B.C.), and the Woodland Period (ca. 0-1,700 A.D.). Interestingly, there was also a
shift in the areas normally inhabited by early peoples, with a transition from ridge top and uplands during the
Archaic Period to lowland areas (likely more conducive to agriculture) during the Woodland Period. The first
European settlers that migrated into what is now known as Alleghany County found few native Americans, but
this small group was comprised of Cherokee Indians (AHGS 1983).
The original Europeans (mostly of English, German, Scottish, and Irish descent) to inhabit this county
migrated south through the Shenandoah River Valley, as well as from other western portions of Virginia in the
mid to late 1700s. This county was initially part of a larger Ashe County before its separation from northeastern
Ashe and subsequent establishment as a new political entity by an act of the 1858-1859 session of the North
Carolina Legislature (Brewer et al. 1973; Alleghany County Historical Committee [ACHC] 1976). The location
of the county seat of Sparta was heavily debated for several years during the Civil War, and was not formally
recognized in its central locality until 1866, following the donation of 20.2 ha of land by James Parks, David
Landreth, and David Evans. County residents initially wanted to name this home for county government
“Parks” after the primary land donor, but instead he insisted on naming it “Sparta” after the ancient Greek city-
state. Likewise, the name “Alleghany” is purported to be derived from an alteration of the Delaware Indian
name for the Allegheny and Ohio rivers, and allegedly translates as “a fine stream” (ACHC 1976; AHGS 1983).
Two of the most important features of human interest in Alleghany County include the Federally owned
and maintained Blue Ridge Parkway (BRP), and the nationally significant New River. The BRP serves as a sce-
nic byway diagonally traversing part of the southern Appalachian Mountains. The lands preserved by the
parkway are of substantial importance due to the habitats they preserve. The first section of the BRP was built
at Cumberland Knob in Alleghany County between 1935 and 1939 (Penny 2010). Over the past couple of years,
the historic rock walls serving as roadside barriers and complimenting the natural aesthetics of this roadway
have been restored. Similarly, the New River is a natural physical element of the county of great interest. The
New River is part of the Ohio River watershed and is a tributary of the Kanawha River. It is believed by many to
be one of the oldest rivers in the world, with its origin predating the Appalachian Mountains. Due to its unique
nature, it is considered one of the nation’s American Heritage Rivers. This river not only provides a source of
recreation and beauty in the county, but also is rather pristine and serves as a sanctuary to many rare plants and
Land Use
Alleghany County is primarily a rural area with an economy driven by agriculture. According to the 2007 ag-
ricultural census (North Carolina Department of Agriculture and Consumer Services [NCDACS] 2012a), ap-
proximately 310.2 km2 (51.0% of the total land area) of this county is used for farmland. Some of the major
agricultural products for this county include Christmas trees (2nd leading producer in the state), hurley to-
bacco (5th in the state), corn for silage (3rd in the state), and milk cows (4th in the state) (NCDACS 2012a,
2012b). Boxwoods (Buxus spp.) are also planted in large quantity in the county, although this horticultural
crop is apparently not a major commodity and thus is not monitored by the North Carolina Department of
Agriculture (pers. obs.). This high level of agriculture-driven land use has led to a very fragmented and highly
altered landscape in the county.
Like many rural counties, Alleghany contains many small, local communities. Many of these communi-
e greatly dissipated over time and are hardly recognized currently, ht
e of these areas of human aggregation are important as they define c(
fluence. Many of the larger extant communities (including the county seat of Sparta) are located along
iclude Cherry Lane, Edmonds, Ennice, Glade Valley, Piney Creek, Roaring Gap,
Stratford, Twin Oaks, and Whitehead (Fig. 2).
In 1880, nearly 20 years after its formal establishment, Alleghany County had a population of 5,486 peo-
ple (Brewer et al. 1973). Based on the 2010 U.S. Census Bureau data (USCB 2012), Alleghany has an estimated
population size of 11,155 people, making it the 7th smallest in the state.
536
Journal of the Botanical Research Institute of Texas 7(1)
The first plant collections known to the author from Alleghany County, based on h.. wcic
in the early 1900s. Fieldwork in the county (as well as North Carolina in general) peaked during the 1950s and
1960s during the production of the Manual of the Vascular Hora of the Carolinas (Radford et al. 1968). Two M.S.
theses were conducted in Alleghany County, either as a site within the confines of the County’s political
boundaries (Bullhead Mountain, Michael 1969) or as a site that overlapped the boundary with a neighboring
county (Stone Mountain State Park, Taggart 1973, 1976). In the last decade, there have been only a few other
published works pertaining to newly documented plants in Alleghany County (Poindexter 2006 2008-
Denslow & Poindexter 2009; Poindexter 2010a, 2010b; Poindexter & Lance 2011; Poindexter & Nelson 2011;’
Poindexter et al. 2011). Botanists and avid collectors that have made significant contributions to the knowledge
ot flonstic diversity m this county within the last 50 years include A.E. Radford, J.B. Taggart, J.L. Mackay, J.L.
Michael, and county natives P.D. McMillan and myself.
sible. This documenta
e collected from the spring of 2008 through the summer of 2012. Despite the presence of preex-
attempt was made to recollect all known taxa to produce the most up-to-date records pos-
effort was augmented by herbarium searches for additional vouchers. The full col-
e University (BOON) was examined, while other herbaria were targeted for specific
records based on database and/or literature searches. These herbaria included: Catawba College, Clemson
University (CLEMS), Duke University (DUKE), Mecklenburg County Park and Recreation (UNCC), North
Carolina State Museum of Natural Sciences, University of Missouri (UMO), North Carolina State University
(NCSC), University of North Carolina-Chapel Hill (NCU), University of South Carolina- Columbia (USCH),
and Virginia Tech (VPI). All specimens were examined for accuracy and annotated
Weakley (2011) was the primary source for plant identification and nomenclature (exceptions are ad-
dressed separately). Other manuals consulted include: Bailey (1924), Bailey and Bailey (1976), ENA (1993+),
Fernald (1950), Gleason and Cronquist (1991), Radford et al. (1968), Rehder (1937), Small (1933), Stace (2010),
and Wofford (1989). Decisions regarding alien species inclusion followed a liberal philosophy (see Poindexter
et al. 2011; Weakley 2011), whereby all exotic plants that are either naturalized or demonstrate the ability to
migrate from an origin of cultivation or inadvertent seeding (e.g., adventives/waifs, escapes) are considered
1 acknowledges that establishment is not easily determined, and the
part of the flora. This ii
proach, taxa that are derived from cultivation but are commonly persistent (particularly around old homesites)
or planted with such regularity that they are perpetual agroeconomic elements of the county flora (e.g., Abies
/raserO were also documented. In general, all other cultivated taxa that were demonstrably maintained
through human interaction and not spreading were not vouchered.
The following terminology was applied to taxa not indigenous to the eastern United Stated and native
I Murrell (2008): exotic = any nonnative taxon considered
cultivated species as adapted from Poindexter ;
and regularly invade r
generally short-lived (i
se noted; invasive - naturalized exotics capable of becoming dominant in
il and disturbed habitats; adventive = unintentional and sporadi
tent from culth
= intentionally planted persis-
1 United States, but not necessarily to the study area; exotic persistent
from cultivation = planted and established (i.e., surviving for several years) but not spreading; and escaped =
generally perennial taxa (including both exotic and planted natives) that appear to weakly spread from cultiva-
tion and may possibly become established. Taxa were determined to be invasive in the southeast based on the
Southeast Exotic Plant Pest Council ([SE-EPPC] 2012). This list was followed closely, except in a few rare cases
where nativity is questionable (e.g., Solanum carolinense var. carolinense).
Putative assignments for county and state records were determined based on Radford et al (1968), Flora
of the Southeast atlas (FSE 2012), FNA (1993+), Kartesz (2012), and the PLANTS Database (United States De-
Poindexter, Flora of Alleghany County, North Carolina 537
partment of Agriculture, Natural Resources Conservation Service [USDA, NRCS] 2012). State records originat-
ing from Alleghany County in other recently published literature (e.g., Poindexter et al. 2011; Rothrock et al.
2011) were also noted. State and global rarity was accessed for each taxon based on the North Carolina Natural
Heritage Program (Buchanan & Finnegan 2010). “Significantly Rare” and “Watch List” species were reported
to the North Carolina Natural Heritage Program to promote conservation efforts. Taxa that were simply persis-
tent from cultivation were not considered as county or state records. Likewise, persistent or weakly escaped
cultivated natives were not assessed for rarity.
The full set of voucher specimens from this study was deposited in the Appalachian State University Her-
barium (BOON), with a limited set at the UNC-Chapel Hill Herbarium (NCU), and a partial duplicate set of
Carex at the New York Botanical Garden (NY). Additional select duplicates were sent to various institutions
and may be located in the customized online database described below.
All specimens were georeferenced with a handheld GarminT” GPSMAP 60Cx unit as they were collected,
using WGS 84 as the reference datum. Legacy specimens from other herbaria that lacked GPS coordinates were
assigned an estimated coordinate using Google Earth ™ and an existing knowledge of the geographical and
ecological attributes of the county. This heuristic method was employed, rather than utilizing less precise geo-
referencing software, to increase location accuracy. The flora was digitized and used to create an online search-
able database of specimens and their respective repositories, select field images, and associated label data.
Certain specimen locality data (but not images) were blocked due to land ownership or conservation concerns.
This tool was generated as a companion outlet for the extensive amount of floristic information that does not
traditionally occur in manuscripts. It was also created to help the general public, land managers, educators,
and researchers better understand the flora, and possibly add to our knowledge of the county’s vascular plant
diversity (via new additions and annotations/corrections) in the future. This website can be accessed at www.
vascularflora.appstate.edu (Poindexter 2012). Search filters are also provided in this database for specimens
associated with the Blue Ridge Parkway, as well as for vouchers corresponding to an ancillary biocontrol study
focusing on the vegetation dynamics and management of Persicaria perfoliata within the county. A download-
able copy of the annotated list (vouchered taxa only) in Microsoft Excel ® format is also provided on the web-
site to allow for data parsing and integration by researchers.
Species richness was evaluated for the Alleghany County flora using multiple power models. A conserva-
tive approach was taken, utilizing species numbers (rather than total taxa) to safely compare at the same level
of taxonomic resolution. These models are represented as 5 = cA^, where S = the number of expected species for
a given area (A), c = the y-intercept or constant, and ? = the slope or 2 coefficient. Area (A) units are in number
of hectares. This model is also accompanied by a coefficient of determination (r^), which ranges from 0 to 1 and
describes how well a regression line fits the data. ’Values closer to 1 indicate a better fit. Three unpublished
models, based on data from the FloraS of North America Project (the “S” symbolically distinguishes this proj-
ect from the similar-sounding “Flora” of North America Project; http://botany.okstate.edu/floras/index.
html), were supplied by M.W. Denslow (Appalachian State University, two North Carolina models) and M.W.
Palmer (Oklahoma State University, one continental United States model). The first model was broadly inclu-
sive for the North Carolina Mainland (including all physiographic provinces and excluding barrier islands; S =
ISO.SOA*^ r^ = 0.443), and the second model was based exclusively on floras from the mountains of North
Carolina (S = 76.10AO 210^ ^ o.537). The latter broad-scale model for the continental United States was based
on 3600 floras (S = 106.44A° = 0.557). In addition, published models from the Cumberland Plateau
(Huskins & Shaw 2010; S = 82.12A° ^613^ ^ = 0.780) and two models from the Mixed and 'Western Mesophytic
forest region (’Wade and Thompson 1991 [S = 272.10A"-113, r^ = 0.802]; Huskins and Shaw 2010, corrected
model [S = 260.82A° r^ = 0.769]) were assessed.
Plant communities were delineated through field reconnaissance and collections, as well as the evalua-
tion of several physical parameters, including general topography, aspect, moisture regimes, soil and geology,
dominant species, anthropogenic influence, and general vegetation composition. This study relies heavily on
these personal observations in conjunction with the North Carolina Natural Heritage Program Significant
Journal of the Botanical Research Institute of Texas 7(1)
Natural Heritage Inventory (Padgett 2011). Likewise, habitat affinities were derived for each taxon (where ap-
plicable) by using the PLANTS Database (USDA, NRCS 2012) to assign wetland indicator status (based on the
1988 list). This method was employed to qualitatively ascertain additional ecological patterns within the flora.
Modern techniques and current flora writing standards (e.g., Palmer et al. 1995; Palmer & Richardson 2011)
were followed as closely as possible to maximize the utility and accessibility of data within this study.
RESULTS AND DISCUSSION
Approximately 3754 specimens (including duplicates) were collected by the author. An additional series of
ca. 285 specimens were examined from various collectors and institutions (see Methods). A total of 1508
taxa, consisting of 1457 species in 642 genera and 161 families were documented, with only 36 taxon records
(mostly historical) attributed to other collectors. These taxa are represented by nine Lycopodiophyta, 39 Mo-
nilophyta, 23 Acrogymnospermae, and 1437 Angiospermae (Table 1). The latter clade can be further divided
into 412 Monocotyledoneae (Monocots) and 1025 “Dicots” in the traditional sense. This latter informal group-
ing is non-monophyletic, and to better reflect our current understanding of phylogenetics, it is best subdivided
into proper clades. As such, there are two Nymphaeales, 13 Magnoliidae, and 1010 Eudicotyledoneae. The
largest families are the Asteraceae (177 taxa), Poaceae (153 taxa), Cyperaceae (120 taxa), Rosaceae (74 taxa),
Fabaceae (61 taxa), and Lamiaceae (54 taxa). The most taxa-rich genera are Carex (83), Viola (24), Dichanthe-
lium (19), Solidago (16), Juncus (14) and Symphyotrichum (13). Four hundred and thirty-five taxa, constituting
28.8% of the total flora, are nonnative in the eastern United States, of which 141 (9.4% of total flora) are con-
sidered invasive in the southeastern United States (SF-FPPC 2012). This high exotic percentage is most likely
the consequence of high levels of disturbance associated with large-scale agricultural practices and residential
development.
To best discriminate the various origins of taxa reported in this flora, a quantitative summary of source
categories is provided (Table 2). As interpreted, 1408 taxa (1360 species) are naturally/sporadically occurring
exotic (including adventive) and native plants. The remaining 100 taxa (97 species) are derived from some
cultivated origin. Excluding adventives, the full flora consists of 1382 taxa (1335 species).
A total of 38 additional records are tentatively included in the list (for a total of 1546 taxa, 1495 species) as
they have been previously reported by reliable sources, but due to the lack of unequivocal physical evidence in
the form of voucher specimens or images, these taxa are not part of the formal taxonomic summary. Likewise
many other records have been excluded altogether due to either incorrect determinations or in case of sight
records, were highly implausible to occur in the county based on geographical affinities. Twenty-nine records
are from the FSE atlas (2012) database, which combines several data sources including Radford et al. (1968),
The Carolina Vegetation Survey (http://cvs.bio.unc.edu/), and others. Nine taxa are based on records from the
North Carolina Natural Heritage Program (NCNHP 2012). These collective sight records are uniquely denoted
within the annotated checklist to avoid confusion.
Rare Taxa, State and County Records, and Other Taxonomic Issues
Sixty-five taxa (Table 3) are currently considered “Significantly Rare” by the North Carolina Natural Heritage
Program (Buchanan & Finnegan 2010). Though several species from this county are listed as Federal Species
of Concern, no Federally Endangered or Threatened taxa have been documented. An additional category
monitored by the North Carolina Heritage Program contains “Watch List” species. This category accommo-
dates taxa that are rare or threatened and demonstrate serious population decline, but are not justifiably wor-
thy of major conservation efforts. Criteria for inclusion in this group range considerably. For instance, some
decline, or simply increasing in rarity as a consequence of commercial exploitation. The flora of Alleghany
County currently contains 100 taxa on this list.
Twenty-one state records have been previously published from plants in Alleghany County (Poindexter
2008, 2010a, 2010b; Poindexter & Lance 2011; Poindexter & Nelson 2011; Poindexter et al. 2011). An additional
541
As with any large scale study, many taxonomic issues were encountered. Some of these, for example.
1. Acer— collections referred to as Acer nigrum are atypical for this species and may represent a
tion within the A. saccharum species complex.
2. Calystegia — members of the “sepium complex”, as well as C. silvatica ssp. fratemiflora, are highly variable
and often difficult to segregate.
3. Elymus — some individuals appear intermediate between E. glabriflorus and E. macgregorii, while E. virgini-
cus is presumably absent from the flora.
4. Eutrochium — high levels of morphological variation in trichome structure for E. purpureum var. purpureum
are not well addressed in the literature.
5. Fallopio — many populations are intermediate between F. cristata and F. scandens with regard to fruit
morphology.
6. Lycopus — potential introgression between most L. uniflorus and L. virginicus (see Henderson 1962) in Al-
leghany County has produced a swarm of entities attributable to the hybrid L. xsherardii. Gene flow be-
tween these two species seems plausible, and few if any populations of either species appear “pure.”
7. Pycnanthemum — at least two distinctive and consistently separable entities are combined within the concept
of P. muticum. Likewise, many aberrant forms not assignable to any concept were encountered.
8. Tilifl— several populations, particularly in mafic sites, have abaxial leaf vestiture approaching var. americana
and clearly not densely stellate-tomentose as in the frequently encountered var. heterophylla- however,
these aff. var. americana populations do possess sparse stellate trichomes intermixed with acicular hairs
suggesting some local introgression between these taxa.
9. Vitis — like many other taxa with intergrading varieties (e.g., Eagus grandifolia), V. aestivalis var. aestivalis and
var. bicolor can only rarely be differentiated.
Species Richness
To date, only three floristic inventories that were explicitly considered comprehensive (i.e., collections made
over one or more full growing seasons) for a given county within North Carolina have been conducted (Horton
1957; Britt 1960; Blair 1967). However, other inventories have been published that consist of areas of compa-
542
s7(1)
rable size to a county, but were not within equally defined political boundaries (e.g., Sorrie et al. 2006; 734.8
km^ = 1206 taxa). Similarly, some floras have implied that they were county-wide in scope, but have either
admitted to the inadequate collection of certain plant groups (e.g., Memminger 1915) or were fractious and ill-
defined (Wood & McCarthy 1886; 1202 taxa; Peattie 1928, 1929a, 1929b, 1929c, 1930, 1931, 1937; 1090 taxa).
All three of the presumably complete county floras were M.S. theses that were not formally published. These
studies include Beaufort County (Blair 1967; 2142.4 km^, 951 taxa), Robeson County (Britt 1960; 2458.5 km^
931 taxa), and Rowan County (Horton 1957; 1324.4 km^,787 taxa). These floras represent areas within the
Tidewater/Coastal Plain, Coastal Plain, and Piedmont regions of North Carolina, respectively. This checklist
for Alleghany County constitutes the first full county flora primarily from the Mountain region of the state, as
well as the first such flora to be produced in the last 45 years. It also important to note that Memminger’s work
was within an entirely montane county (Henderson County). On a similar note, Peattie’s studies were chiefly
within a Piedmont county with some mountains to the far west (Polk County and adjacent South Carolina).
Likewise, Alleghany is a far smaller county, but contains more recorded taxa than previous county-wide com-
prehensive studies.
To assess the comprehensive nature of this study, several species area power models we
spite considerable variation between models (Table 4), all demonstrated a 40% or greater po
viation from the number of predicted species for this flora, except for the Cumberland Plateau model, which
indicated a slight negative deviation from the expected number of species. This divergence from the other
models is likely attributed to the source data used to develop this regression curve. No floras in excess of
10,300 ha were utilized hy Huskins and Shaw (2010), thus making an area the size of Alleghany County well
outside the predictive limitations of the derived model. Based on this analysis, Alleghany County exhibits a
species richness far above what would be expected for an area its size (Table 4). For example, when excluding
cultivated taxa, the total species predicted for the North Carolina Mainland is 711. The actual number of spe-
cies is nearly twice this prediction (1360 spp.) or 91.3% above the predicted number.
This high species richness is most likely the consequence of a broad array of community types and habi-
tats (see below), and the county’s small but notable ecotone-like transition from the Mountains to the Piedmont
(foothills). The geologic, edaphic, hydrologic, and climatic heterogeneity of Alleghany County plays a major
role in this pronounced species richness. Likewise, high levels of disturbance have added a prominent exotic
component to the flora, which undoubtedly contributes to this higher than predicted richness. Similarly, the
general paucity of large-scale (county wide or bigger) studies for model construction may also bias species area
predictions. In contrast, some recent data also suggest that newer floras generally exhibit more species than
older floras of comparable size areas (Denslow et al. 2010). A few additional variables that are likely responsible
for these results include the application of narrower taxonomic concepts, a liberal criterion for alien species
inclusion within the flora, time, effort, and prior floristic experience. Lastly, the fact that most models indicate
that Alleghany County is far more species rich than expected suggests that this study is comprehensive.
The landscape of this county is a mosaic of habitats, ranging from extremely dry to dry-mesic areas, particu-
larly along the edge of the Blue Ridge Escarpment and adjacent foothills, to mesic coves and inundated bogs
and wetlands. National Wetland Indicator Status 1988 list (USDA, NRCS 2012) provides a subjective measure
of a plant’s environmental preference via its hydrological amplitude. This status, though relative, adds to the
ecological attributes of a flora. “Regional Status” for the Southeast (since some plants demonstrate different
wetland preference in other geographical regions) was accessed for all taxa, with a separate analysis of taxa
from non-cultivated origins (i.e., naturalized + invasive + adventive + native) included in brackets. Of the total
flora, only 859 (57.0%) [835 59.3%] had a regional designation. These plants were divided near equally into
three groups. The first group was comprised of “upland” and “facultative upland” taxa (287, 33.4%) [279,
33.4%], which are most likely to occur in non-wetlands. A second group consisted of “facultative” taxa (260,
30.3%) [253, 30.3%] that are equally likely to occur in non-wetlands or wetlands. The third group contained
“facultative wetland” or “obligate wetland” taxa (312, 36.3%) [303, 36.3%], which were most likely to occur in
wetlands. These data confirm that Alleghany County has a wide range of habitat heterogeneity and thus help
to explain the high level of floristic diversity found here.
The plant communities of Alleghany County are initially divided into two categories: natural and disturbed.
As implied, natural communities are relatively unaltered areas containing native elements indicative of little
anthropogenic influence. Natural communities are organized into affiliated groups and modified (including
additions) from Padgett (2011), with subtypes and respective current ranks derived from the North Carolina
Natural Heritage Program database (NCNHP 2012). Entries for each provisional community include subtypes
listed in brackets with corresponding ranks, followed by a brief general description of the major community
as it idiosyncratically occurs in Alleghany County. These categories follow the definitions of Schafale and
Weakley (1990) and Schafale (2012). Community assessment was coarsely qualitative in nature and conse-
quently, several other communities and/or subtypes are likely present and may be recognized in the future
based on the updated and more finely divided classification scheme of Schafale (2012). Lastly, an additional
informal community type (Montane River Aquatic) is qualitatively expanded from previous concepts for the
purpose of this study. State Ranks (S) and Global Ranks (G) follow each community type in brackets. Exact
definitions for these ranks are enumerated by Padgett (2011), but in general, lower ranks indicate greater con-
servation concern, ranging from 1 (critically imperiled) to 5 (demonstrably secure). Disturbed communities
are generically defined.
natural communities
1) High Elevation Red Oak Forest [Heath Subtype S2S3 G4].— This community type is not very common as it
typically occurs at elevations in excess of 1067 m. Eorest structure is generally open and composed of a canopy
dominated by Quercus rubra var. rubra, a near absent subcanopy, a patchy to moderate shrub layer, and a
predominately thick orchard-like understory that exhibits less diversity than more mesic cove forests. Good
examples of this community occur at Bullhead Mountain and nearby along the Blue Ridge Parkway around
Mahogany Rock. In addition to Quercus rubra var. rubra, other rarely intercalated canopy species include Bet-
ula lenta var. lenta, Carya glabra, and even more rarely Fraxinus americana, Quercus coccinea and Q. montana.
Understory species include Acerpensylvanicum, Amelanchier arborea, Cornusjlorida, and Ilex montana. Charac-
teristic shrubs include Kalmia latifolia. Rhododendron calendulaceum, Vaccinium corymbosum, and 1/ pallidum.
Castanea dentata sprouts are often present as well within the shrub layer. The rather dense herb layer is often
composed of Ageratina altissima var. roanensis, Aralia nudicaulis, Carex brunnescens var. sphaerostachya, C. debi-
h C. laxijlora, C. virescens, Danthonia compressa, Dennstaedtiapunctilobula, Eurybia chlorolepis, Maianthemum
canadense, Solidago curtisii, and Thelypteris noveboracensis.
544
Low Elevation Mesic Forest
2) Acidic Cove Forest [Typic Subtype 55 G5].— This community is rather common in the mountains of North
Carolina. In Alleghany County it often occurs on midslopes at low to mid-elevations in narrow rocky gorges,
usually with some substantial rock outcrops. It differs from Rich Cove Forest due to nutrient-poor edaphic
accumulate. It often grades into oak-hickory dominated forests upslope, in drier ridge-top areas. The canopy
is primarily closed and often dominated by Acer rubrum var. rubrum, Liriodendron tulipifera var. tulipifera,
Quercus montana, Q. rubra var. rubra, and Tsuga canadensis, with infrequent Acer saccharum, Carya cordiformis,
and Fraxinus americana present. Subcanopy species are relatively few, with occasional Cornusflorida and Fa-
gus grandifolia var. caroliniana present. The shrub layer is dense, primarily comprised of evergreen ericaceous
shrubs such as Kalmia latifolia. Rhododendron catawbiense, and R. maximum, with rare occurrences of Leuco-
thoe fontanesiana and deciduous species such as Rhododendron periclymenoides. The herb layer is relatively
thin, with small patches of dense vegetation restricted to canopy gaps. Characteristic species include Asple-
nium platyneuron, Carex digitalis var. digitalis, C. nigromarginata, Chimaphila maculata, Galax urceolata, Maian-
themum racemosum ssp. racemosum, Medeola virginiana, Polystichum acrostichoides. Ranunculus allegheniensis,
3) Rich Cove Forest [Montane Intermediate Subtype S4 G4; Montane Rich Subtype S3 G3G4].— Rich Cove
Forest is generally common and well distributed throughout the southern Appalachian mountains. It is most
frequently associated with sheltered ravines along the Blue Ridge Escarpment, on upper slopes above the New
River (and other tributaries), and on north-facing slopes and associated colluvial fans of major peaks within
Alleghany County. These areas often include sheltered rock outcrops that are intermittent and too small to
be characterized as cliffs. The occurrence of this community type is frequently correlated with geology, with
most sites associated with mafic rock (amphibolite). The nutrient-rich conditions that characterize these forest
types are also responsible for high levels of species diversity and vigorous vegetation growth. Consequently,
the canopy layer of this forest type is generally closed, comprised of dense assortment of trees including, but
not limited to Aesculusjlava, Betula lenta var. lenta,Juglans nigra, Liriodendron tulipifera var. tulipifera. Magnolia
acuminata, M.fraseri, Prunus serotina var. serotina, and Tilia americana var. heterophylla. Subcanopy trees often
present are Cornus altemifolia, C.florida, Fagus grandifolia var. grandifolia, Halesia tetraptera, and Hamamelis
virginiana var. virginiana. The shrub layer is often sparse, comprised of Corylus americana. Hydrangea arbores-
cens var. arborescens, Lindera benzoin, and occasional Pyrulariapubera. The herb layer within this community
type is perhaps the most difficult to characterize. It is usually dense and diverse, with even small rock outcrops
dominated by a variety of species (e.g., Aquilegia canadensis, Micranthes virginiensis). A few of the characteristic
taxa include Adiantum pedatum. Allium tricoccum. Anemone acutiloba, Aruncus dioicus var. dioicus, Asarum ca-
nadense, Athyrium aspleniodes, Cardamine concatenata, Carex laxiculmis var. laxiculmis, C. laxiflora, C. plantag-
inea, C. woodii, Caulophyllum thalictroides, Collinsonia canadensis. Dicentra cucullaria, Diplazium pycnocarpon,
Dryopteris spp., Festuca subverticillata. Geranium maculatum, Huperzia lucidula, Hydrophyllum virginianum var.
atranthum, Luzula acuminata var. carolinae, Mitella diphylla, Osmorhiza claytonii, Panaxquinquefolius, Persicaria
virginiana, Poa cuspidata, Polygonatum biflorum var. biflorum, Prosartes lanuginosa, Sanguinaria canadensis,
Sanicula trifoliata, Scutellaria saxatilis, Tiarella cordifolia, Uvularia grandiflora, and Viola blanda.
Low Elevation Dry and Dry-Mesic Forests and Woodlands
4) Carolina Hemlock Forest [Typic Subtype 52 G2].— This community type is defined by the dominance of Tsuga
caroliniana, forming a closed or intermittently open canopy due to rocky substrate (Schafale & Weakley 1990).
This dominant species is a narrow Southern Appalachian endemic, and like its more widespread sister species,
Tsuga canadensis, populations are in rapid decline due to the herbivorous invasive Hemlock Woolly Adelgid
[Adelges tsugae (Annand)]. Both taxa are Federal Species of Concern, but the limited geographic range and
population size of Tsuga caroliniana makes it more susceptible to extinction. Consequently this community
is globally rare, found only in southern Virginia south to northern Georgia along the Southern Appalachians
over acidic soils, on steep slopes and bluffs. As noted by Padgett (201 1), this community may be fire dependent.
545
Carolina Hemlock Forest is also rare in Alleghany County, and due to their limited sizes, true well-developed
examples of this community do not exist. Most occurrences of this relictual community type are located along
the New River palisades near the Virginia border, within New River State Park. Other sites, such as the ridge
of Fodderstack Mountain (Doughton Park) along the Blue Ridge Parkway, are very small remnants. Additional
interspersed and generally rare canopy species include Betula lenta var. lenta, Carya spp., Liriodendron tulipifera
var. tulipifera, and Quercus spp. The subcanopy is almost absent, populated primarily by saplings, while the
sparse shrub layer contains species such as Kalmia latifolia and Vaccinium pallidum. The herb layer is scarce,
with rare occurrences of species such as Goodyera pubescens and Mitchella repens.
5) Chestnut Oak Forest [Dry Heath Subtype S5 G3; Herb Subtype S4 G4G5; Mesic Subtype S4? G4]. — ^This
is a frequently occurring community type that is most common at low to middle elevations, especially along
the Blue Ridge Escarpment and drier south and east-facing slopes within the county. The canopy tends to be
closed, but is not particularly dense, and gaps usually occur in very rocky areas. The primary canopy trees
include Quercus montana as the dominant species, fair amounts of Q. coccinea, and occasional Q. alba and Q. ru-
bra. Other intermittent species include Acer rubrum var. rubrum and Carya spp. Subcanopy trees often include
Amelanchier arborea, Cornus florida, Crataegus spp., and Oxydendrum arboreum. The shrub layer is variable
in density and is usually comprised of species such as Corylus comuta, Eubotrys recurva, Gaylussacia baccata,
Kalmia latifolia. Rhododendron calendulaceum, R. catawbiense, R. maximum, Vaccinium spp., and Viburnum ac-
erifolium. The herb layer is characteristically sparse and exhibits little overall diversity. Frequent components
include Carex appalachica, C. pensylvanica, C. swanii, Chimaphila maculata. Coreopsis major var. rigida, Dantho-
nia compressa, D. spicata, Dennstaedtia punctilobula, Epigaea repens, Galax urceolata, Galium pilosum, Gaultheria
procumbens, Lespedeza violacea, Pteridium aquilinum var. latiusci
6) Granitic Dome Basic Woodland [S2 G2].— The concept for this community ii
thin soils over granitic substrates and around the periphery of exfoliated rock outcrops. They are limited to the
upper Piedmont. In Alleghany County, this community type is confined to areas atop Stone Mountain’s larger
granitic domes and a few adjacent slopes, at the base of the Blue Ridge Escarpment. By definition, this commu-
material and there is a general lack of montane species. The canopy is closed to somewhat open, dominated by
stunted Quercus montana, with interspersed Carya glabra, C. tomentosa, Nyssa sylvatica, and rarely a few Pinus
virginiana. Per Schafale (2012), Carya spp. and Fraxinus americana are supposed to be abundant, with oaks
generally scarce, thus there is some deviation here from the principal community structure. The subcanopy
is essentially absent, while the very sparse shrub layer is occupied by scattered Gaylussacia baccata, Kalmia
latifolia, and Vaccinium pallidum. One of the characteristic and most prevalent elements of this community
type is its herb layer. This portion of the community lacks diversity and at Stoi
near monotypic dense stand of the indicator grass species, Piptochaetium avenac
include Carex glaucodea, Galax urceolata, Hexalectris spicata, and Tipularia discolor.
7) Montane Oak-Hickory Forest [Acidic Subtype S4S5 G4G5; Basic Subtype S3 G3].— Montane Oak-Hickory
Forest is widespread through Alleghany County and the mountains of North Carolina. ’Within the study site,
it is most often found along middle to upper slopes just off the Blue Ridge Escarpment, and along inner mon-
tane north and east-facing slopes. In contrast to Chestnut Oak Forest, this community occurs in more mesic
to sub-mesic, protected sites. As pointed out by Padgett (2011), the soils are variable and range from acidic to
somewhat basic, particularly in areas with slight mafic geology. The canopy layer is usually closed and domi-
nated by Carya cordiformis, C. glabra, C. ovalis, and infrequent C. ovata and C. tomentosa. Oaks also contribute
a major component to the canopy layer and include Quercus alba, Q. montana, and Q. rubra var. rubra. Fraxmus
and M. fraseri are also usually present in limited amounts. The subcanopy is
:e, comprised of species such as Amelanchier arborm. A. laevis, Cornus florida. Crataegus mac-
■ --yMenziesiapilosa,Ostryavirginiana,
ar. rubrum, Nyssa sylvatica, and Oxy-
a such Castanea dentata (sprouts), Kalmia
546
e of Texas 7(1)
latifolia. Rhododendron spp., Vaccinium spp. and occasional Viburnum acerifolium and V. prunifolium, with
woody vines such as Vitis spp. regularly encountered. The usually sparse to moderately dense herb layer can
be rather diverse, with some characteristic species including Agrostis perennans, Aplectrum hyemale, Asclepias
exakata, Aureolaria laevigata, Carex aestivalis, C. albicans, C. digitalis var. digitalis, C. pensylvanica, Danthonia
tatum, Dichanthelium latifolium, Dryopteris spp., Galium circaezans var. circaezans, Gentiana austromontana,
Hieracium paniculatum, Houstonia purpurea var. purpurea. Iris cristata, Prenanthes spp., Scutellaria elliptica var.
elliptica, Veratrum parviflorum, and Zizia trifoliata.
8) Pine-Oak/Heath [High Elevation Subtype S2 G2; Typic Subtype S3 G3]. — ^This community is most preva-
lent along steep, rocky ridgelines and crests of low to middle elevations, particularly along the edge of the Blue
Ridge Escarpment and south-facing portions of adjacent peaks (e.g.. Bullhead Mountain, Saddle Mountain).
Soils are characteristically dry and very acidic, generally as a consequence of topography, highly exposed
habitat, and dominant vegetation. The canopy is quite open and dominated by Nyssa sylvatica, Oxydendrum
arboreum, Pinus pungens, P. rigida, P. virginiana, Quercus alba, Q. coccinea, and infrequent Q. velutina. The sub-
canopy is essentially absent, but the shrub layer is usually well developed and characterized hy species such
as Castanea pumila, Comptonia peregrina, Eubotrys recurva, Gaylussacia baccata, Kalmia latifolia, Rhododendron
spp., and Vaccinium pallidum. The herb layer is sparse, but includes Aristida dichotoma, Epigaea repens, Galax
urceolata, Gaultheria procumbens, Melampyrum lineare var. americanum, Mitchella repens, Pteridium aquilinum
var. latiusculum, and the woody vines Smilaxglauca and S. rotundifolia.
Rock Outcrop Communities
9) Low Elevation Granitic Dome [S2 G2]. — ^This community is defined by large expanses of exfoliating granitic
rock with the absence of deep soil pockets and crevices (particularly along steeper sloping portions of the
domes) that are usually found on other more fractious rock types. The top of these outcrops are usually flat and
conducive to shallow soil accumulation. Where soil is most developed, this community becomes transitional.
The largest example of a Low Elevation Granitic Dome in Alleghany County occurs at Stone Mountain State
Park, where the presence of several large plutons makes this a site of national significance (Padgett 201 1). Other
smaller examples are scattered along the Blue Ridge Escarpment. Vegetation occurs in the shallow soil mats of
this community type, specifically at the margins of adjacent forested areas and gently sloping pockets. Because
of these constraints, no true canopy or subcanopy exists. Rarely shrubs and small trees from neighboring com-
munities (e.g.. Granitic Dome Basic Woodland) may become established, yet remain dwarfed and include
Gaylussacia baccata, Kalmia latifolia, Pinus spp.. Rhododendron maximum, and Vaccinium pallidum. The prima-
ry components of the community occur in the herb layer in thin soils and include species such as Bulbostylis
capillaris, Cyperus retrorsus, Dichanthelium meridionale.Juncus secundus, Linum medium var. texanum, Minuartia
glabra. Paronychia fastigiata var. paleacea, Phemeranthus teretifolius, Scleriapauciflora, and Selaginella rupestris.
10) Low Elevation Rocky Summit [Acidic Subtype S3 G3?]. — ^This community type occurs in scattered
localities throughout Alleghany County at middle to low elevations, generally below 1067 m. It is comprised
of exposed ridges of rugged rock outcrops with uneven vertical to sloped faces. This relatively uncommon
community type is located along escarpment ridges and along several of the larger mountains within the
county such as Bald Knob, Bluff Mountain, Doughton Mountain, and Twin Oaks Mountain, among others.
Vegetation in this community is restricted; very little, if any, canopy is present and most plant life is confined to
small islands of soil accumulation along the most horizontally oriented surfaces, in crevices of fractured rock,
ledges, and near cliff bases. Higher elevation species are generally lacking. This open canopied community
may occasionally support, in areas with deeper soils, shrubs and small trees (usually stunted) in areas with
deeper soils such as Chionanthus virginicus, Clethra acuminata, Eubotrys recurva, Pinus pungens, Salix humilis,
Sorbus americana, Chionanthus virginicus, Vaccinium erythrocarpum, and V. stamineum. The herbaceous layer is
localized and often contains species such as Andropogon virginicus var. virginicus, Avenellaflexuosa, Campanula
divaricata, Capnoides sempervirens, Carex rugosperma. Coreopsis spp., Crocanthemum canadense, Danthonia
spicata, Helianthus divaricatus, Heuchera villosa var. villosa, Hydatica petiolaris, Hylotelephium telephiodes, Hy-
pericum gentianoides, Schizachyrium scoparium var. scoparium, Selaginella rupestris, and Wbodsia spp.
11) Montane Cliff lAcidic Herb Subtype S3 G3G4].— The community type is defined by steep rock faces and
slopes that accumulate soil in small fissures, ledges, and talus. Ultimately, the sheer slope of this community
type eliminates the potential for canopy development, and much of the cliff face is barren except for bryophytes
and lichens. Most canopy species are found along the periphery of these cliffs, providing some indirect shad-
ing. These sites usually occur near the top of dry ridges and peaks and abruptly descend into mesic forests at
their bases. They occur throughout Alleghany County, usually as small isolated examples, but are most fre-
quent in watershed areas of the New River and adjacent tributaries, as well as north-facing slopes of some of the
higher peaks. Sparse vascular vegetation is limited to an herb layer and often consists of characteristic species
such as Asplenium montanum, A. trichomanes ssp. trichomanes, Heuchera villosa var. villosa, Micranthes carolin-
iana, and Polypodium appalachianum.
12) Low Elevation Acidic Glade [Grass Subtype S1S2 G1G21. — Occurrences in Alleghany of this communi-
ty type are few, and limited to small patchy areas surrounding and often associated with or transitional to Low
Elevation Rocky Summit communities. This community is characterized by gently to moderately sloping rock
outcrops with shallow soils and few crevices supporting a predominance of graminoids, scattered low shrubs,
and sparse small trees. As implied, the canopy is open, allowing for high light exposure. Some of the rarely
occurring small or stunted trees and shrubs in this community type include species such as Crataegus mac-
rosperma, Diospyros virginiana, Quercus montana, and Vaccinium spp. Vegetation mats are often accompanied
by lichens (Cladonia spp.) and consist of species such as Andropogon virginicus var. virginicus, Carex tonsa, C.
umbellata, Cyperus lupulinus var. lupulinus, Danthonia compressa, D. spicata, Festuca rubra var. rubra, F. trachy-
phylla, Schizachyrium scoparium var. scoparium, and Selaginella rupestris. Though dominated by graminoids,
this community type also harbors the rare occurrence of sexual diploids of Erigeron strigosus and the only
known locality for Polygonum tenue in the county. One of the best examples of this community type occurs at
Bluff Mountain, while additional smaller sites are scattered.
13) Montane River Aquatic [N/A]. — This community is described here to accommodate for fully aquatic vegeta-
tion within montane river systems. It is perhaps best treated in or at least affiliated with the Rocky Bar and
Shore community complex, as it shares affinities with the Rocky Bar and Shore (Riverweed Subtype). As stated
by Schafale (2012), the Riverweed Subtype “covers largely-submerged riffles where P
dominates, generally in nearly monospecific stands” and “this community is more ac
types, and may warrant a separate community type.” This community subtype currently lacks a state rank, but
is considered G3G5 globally. 1 concur that it seems distinctive, and here modify and expand the concept to in-
clude the presence of other aquatic vegetation in addition to Riverweed. As defined here, this community
illuvial soils or attached to rocky substrates within larger rivers, often away from
i. Plants are primarily submerged or rooted-floating herbaceous species in moder-
ately to swiftly-flowing waters, with a general lack of any emergent taxa. This community type notably occurs
within the Little River, South Fork of the New River, and the New River itself. Characteristic aquatic species
include: Elodea canadensis, Podostemum ceratophyllum, Potamogeton epihydrus, and Vallisneria amencana.
14) Rocky Bar and Shore [Alder-Yellowroot Subtype S3 G3G41.— This community is primarily found along
the South Fork of the New River and the New River proper, in the vicinity of river banks, but also as exposed
river islands. The best examples of this community type include areas that have either gravel or bolder depos-
its, or soil accumulations in rock outcrops. These riparian zones also include eroded cuts or channeling, but
characteristically include gravel and scour bars that are too regularly disturbed via periodic flooding to sup-
port a canopy or understory layer, but often leave demonstrable alluvial deposits. Trees are absent to rare and
include, Platanus occidentalis and Salix nigra. Shrubs and small trees characteristic of these areas include AIni«
woody vines such as Vitis labrusca and vulpina. Herb layer vegetation is highly variable and fleeting in this
community type, but often includes Boykinia aconitifolia, Carex spp., Cyperus flavescens, Eleocharis spp., Equi-
Journal of the Botanical Research Institute of Texas 7(1)
spp., Lobelia cardinalis, Ludwigia palustris, Myosotis scorpiodes, Persicaria pensylvanica, P. puncata, and Scutel-
laria lateriflora, among others. Some invasive exotics (e.g., Arthraxon hispidus var. hispidus and Microstegium
vimineum) can be found in this community type, presumably due to regular disturbance via river scouring. A
few small areas along the Little River and Prathers Creek approach the Twisted Sedge Subtype, however Carex
15) Montane Alluvial Forest {Small River Subtype SI G3].— This community is found in floodplains and
slopes along major tributaries within the county. Although it contains an amalgamation of cove and floodplain
species, it is the presence of this latter group of indicator taxa, coupled with a regular flooding regime that
distinguishes this community from Rich Cove Forests and Acidic Cove Forests. The best current examples
occur in areas adjacent to rivers that have steep slopes, below rocky cliffs that are for the most part agricultur-
ally inaccessible. It was formerly much more common, but the rich alluvium in riparian areas is prized by
farming, while the remainder of riverside landscape has fallen to other industries including construction and
timber production. Important mesophy tic canopy species that contribute a relatively dense cover include Acer
rubrum var. trilobum, Aesculus jlava, Betula alleghaniensis, Liriodendron tulipifera var. tulipifera, Platanus occi-
dentalis, Populus xjackii, and Tsuga canadensis. Characteristic subcanopy and shrub layer taxa are Acer negundo
var. negundo, Carpinus caroliniana var. virginiana, Prunus americana, Ptelea trifoliata var. trifoliata. Rhododen-
dron arborescens, R. maximum, Sambucus canadensis, Tilia americana var. heterophylla, Viburnum prunifolium,
and Xanthorhiza simplicissima. The herb layer is generally dense and comprised of many cove species, as well as
other taxa including Boehmeria cylindrica, Dichanthelium spp., Clyceria spp., Heracleum maximum, Impatiens
spp., Leersia spp., Packera aurea, and Viola spp.
Nonalluvial Wetlands
One of the most important natural aspects of Alleghany County is its possession of numerous nonalluvial
wetlands. As stated by Padgett (2011), this small county contains “some of the best examples of Southern Ap-
palachian Bog and Swamp Forest-Bog Complex natural communities in the state and the nation.” He continues
by enumerating a few exemplars including Brush Creek Bog, Laurel Branch Bog, Skunk Cabbage Bog, and
Sparta Bog. These wetlands are inherently fed by small spring seeps that are themselves uncommon and rather
unique communities. This hydrological and ecological community interdependence is a frail example of the
synergistic nature of our natural heritage. Consequently, the following communities are some of the most im-
periled in North Carolina, largely due to agriculture and residential development.
16) Low Elevation Seep {Montane Subtype S2S3 G2G3] . — Seeps are frequent scattered elements of the coun-
ty, often originating on mountain or hillsides and draining into low lying wetlands or other tributaries. Many
of the larger examples include very small woodland streams that spread out in lowlands creating seepage bogs.
This community type is transitional to a Rich Montane Seep, but generally lacks rich higher elevation indica-
tor species found in this community. Species composition is otherwise quite variable. Soils are often a mixture
of rocky intermittent stretches and mucky saturated areas. The canopies of Low Elevation Seep communities
are usually closed, with only the most inundated areas with gaps. Common mesic trees include Acer rubrum
var. rubrum, Aesculus Jlava, Betula lenta var. lenta, Liriodendron tulipifera var. tulipifera, Magnolia acuminata var.
acuminata, Quercus rubra var. rubra, and Tilia americana var. heterophylla. Subcanopy and shrub layer species
include sparse Clethra acuminata. Ilex montana, Kalmia latifolia, Oxydendrum arboreum. Rhododendron spp.,
and Vaccinium spp. Herbaceous vegetation is distinctive, occurring on mounds, in rock crevices, and in muddy
s Cardamine bulbosa, C. Jlagellifera var. Jlagellifera, C. pensyl-
itana, C. prasina, C. scabrata, C. stipata var. stipata, Chelone
glabra, Deparia acrostichoides, Glyceria melicaria, Hydrocotyle americana, Juncus spp. (one site withj. gymno-
sulcatum, Veratrum viride, rarely Veronica americana, common Viola cucullata, and V. macloskeyi ssp. pallens.
17) Southern Appalachian Bog {Low Elevation Subtype S1S2 G1G2; Skunk Cabbage Subtype SI Gl; Typic
Subtype S1S2 G1G2]. — ^According to Padgett (2011) this general community type is restricted to the mountains
of North Carolina, Tennessee, and Virginia. These communities occur throughout the county, particularly
549
along the Blue Ridge Parkway, NC 18 and US 21. Vegetation within these sites is very zonal with an absence of
canopy species, encroaching shrubs that are also interspersed throughout in lesser amounts, and distinctive
tussocks of graminoids and herbs that form an open inundated meadow. These communities serve as refugia
for many northern and coastal disjunct species, further adding to their uniqueness and varying in quality, with
the most altered examples containing few if any rare species and simply transitional to a degraded wet-mead-
ow. Substrate is variable, with many of Alleghany County’s bogs occurring over mafic rock, or in the case of
Savannah Church Bog, over ultramafic rock that imparts some fen-like qualities to the vegetation. Commonly
encountered shrubs and trailing woody species include Alms serrulata, Aronia spp., Hypericum densijlorum, H.
prolificum, Kalmia Carolina, Lindera benzoin, Lyonia ligustrina. Rhododendron viscosum, Rosa palustris, Salix
sericea. Spiraea alba, S. latifolia, Vaccinium fuscatum, V. macrocarpon, and Viburnum cassinoides. Herbs and
graminoids are especially diverse, with some consisting of Andropogon glomeratus var. glomeratus, Apios ameri-
cana, Bartonia virginica, Calamagrostis canadensis var. canadensis, C. coarctata, Calopogon tuberosus var. tu-
berosus, Carex atlantica, C. buxbaumii, C. echinata ssp. echinata, C. stricta, C. stylqflexa, Chelone cuthbertii, Ci-
cuta maculata var. maculata, Dichanthelium lucidum, Drosera rotundifolia var. rotundifolia, Eleocharis spp., Epilo-
bium leptophyllum, Eriocaulon decangulare var. decangulare, Eriophorum virginicum, Galium asprellum, Gentiana
saponaria, Glyceria laxa,Juncus brevicaudatus, J. longii,]. subcaudatus, Linum striatum, Lysimachia terrestris,
Osmunda spp., Osmundastrum cinnamomeum, Oxypolis rigidior, Panicum virgatum var. virgatum, Parnassia spp.,
Platanthera spp.. Polygala cruciata var. aquilonia, Pycnanthemum spp., Rhynchospora spp., Sanguisorba canaden-
sis, Scleria spp., Selaginella apoda, Stenanthium gramineum var. robustum, Thalictrum macrostylum, Thelypteris
palustris var. pubescens, and Xyris torta.
18) Swamp Forest-Bog Complex [Typic Subtype S2 G2].— This community type shares many affinities with
Southern Appalachian Bogs. The primary difference is in the physiognomy of these areas, which exhibits a
complex matrix of dense wooded thickets with intermittent small openings that correspondingly vary from
shade tolerant to shade intolerant species. As with bogs, they are mostly restricted to bottomlands. These com-
munities are considered to be drier than bogs, yet this varies considerably. The most common occurrences
of Swamp Forest-Bog Complex can be found along the Blue Ridge Parkway neighboring some of the streams
(e.g.. Brush Creek, Big Pine Creek) that parallel this road. Canopies often contain species such as Acer rubrum
var. rubrum, Pinus rigida, P. strobus, and Tsuga canadensis, with rare occurrences of Magnolia tripetala. The
subcanopy and shrub layers intergrade and include species such asAlnus serrulata, Hypericum densiflorum. Ilex
verticillata, Kalmia Carolina, K. latifolia. Rhododendron maximum, Sambucus canadensis. Toxicodendron vemix,
and Viburnum nudum. Species often encountered within the herb layer are Arisaema triphyllum ssp. stewardso-
nii, Carex bullata, C. folliculata, C. gynandra, C. intumescens var. intumescens, C. laevivaginata, C. longii, Cinna
arundinacea, Dryopteris cristata, Festuca subverticillata, Houstonia serpyllifolia, Osmunda claytoniana, Osmun-
dastrum cinnamomeum, Onoclea sensibilis var. sensibilis, Rubus dalibarda, R. hispidus, and Symplocarpusfoetidus.
DISTURBED COMMUNITIES
Culturally disturbed and ruderal communities are prevalent within the county. These areas are exemplified by
urbanization, roadsides and ecotones, residential lawns, annual crop lands, Christmas tree plantations, and
any other areas maintained by regular human activity. Due to the complex nature of these communities, larger
associations are not given.
19) Plantations.— This community category refers to areas of woody plant cultivation for agroeconomical
purposes. More specifically, these farms include regularly spaced plantings of Abies concolor, A.fraseri, Buxus
sempervirens, Pinus strobus, and other species that are harvested after several years of growth and maintenance.
Intermittent vegetation is usually comprised of exotic annual and perennial grasses such as Digitaria sanguina-
lis and Schedonorus arundinaceus, as well as an assortment of exotic herbs such as Arctium minus, Cerastium
spp. and Stellaria media, and native weedy species such as Ambrosia artemisiifolia and Chenopodium album.
20) Agricultural Fields and Farms.— In contrast to a plantation, this community category accommodates
both agricultural fields that experience an annual harvest of crops and regular tillage practices and less regu-
larly tilled fallow fields. In addition, this community refers to farm areas that harbor livestock (e.g., cattle and
550
y area that exhibits
hogs) in a localized and heavily disturbed environment. Many of the species encountered in this community
are exotic adventives that are introduced from the previous year’s crop rotation (e.g., Zea mays ssp. mays), are
weedy naturalized species that reseed themselves (e.g., Amaranthus spp.) or reemerge from perennial rootstock
(e.g.. Convolvulus arvensis). Likewise, weedy species are often inadvertently introduced as feed contaminants
for livestock (e.g., Marrubium vulgare) or seed and have ultimately become established elsewhere in the county
(e.g., Silenejlos-cuculi ssp. Jlos-cuculi).
21) Meadows, Pastures, and Ecotones. — Sites conforming tc
a predominance of graminoids (Cyperaceae, Juncacea*
ther maintained irregularly, with some succession allowed to occur (meadows), or are utilized for grazing cat-
tle and mowed one to several times annually for hay feed (pastures). In contrast to an agricultural field, soils
remain less disturbed. Reminiscent of plantations, they usually contain a predominance of annual and exotic
perennial grasses, but often also have native graminoids regularly interspersed within the site. In Alleghany,
pastures and meadows often abut tributaries and comprise the floodplains of these waterways. Such areas are
frequently wet, with partially inundated mid-fleld depressions. Sites like this are most common along the New
River and Little River and their associated tributaries. These wet meadows are sometimes transitional to South-
ern Appalachian Bogs, and in addition to exotics, they often contain native species such as Carexfrankii, C.
stylqflexa, Leersia oryzoides, Lilium grayii, Mimulus ringens var. ringens, Scirpus cyperinus, S. expansus, S. poly-
phyllus, and Spartina pectinata. In other areas, pastures are commonly surrounded by woodlands and are eco-
;. Upland taxa, particularly small native trees such as Crataegus spp. and Malus
22) Roadsides, Power Line Corridors, and Ecotones.— These areas are highly generalized and account for a
wide array of vegetation patterns. As with pastures bordering tributaries, roadsides are quite often periodically
flooded and wetland-like. Included within this community type are flat gravelly roadside shoulders, drainage
ditches, and woodland ecotones. As a consequence of this tremendous variability, these sites often contain
both exotic and native taxa, the latter of which are introduced from bordering natural communities. Drainage
ditches are particularly important as they can harbor hydrophilic native taxa such as Carex lurida, Persicana
hydropiper, Schoenoplectus tabernaemontani, and Scirpus hattorianus. In contrast, the physical instability of this
community type exposes it to the introduction (perhaps vehicular in some cases) of adventives (e.g.. Sorghum
bicolor var. bicolor and Tagetes spp.) and other naturalized exotics. Woodland borders and embankments are
especially important as they provide open habitat for native taxa such as Desmodium spp., Helianthus atroru-
bens, Lespedeza spp.. Phlox spp., Physostegia virginiana ssp. praemorsa, Spiranthes cemua, and Veronicastrum vir-
ginicum. Power line corridors are often similar to meadows, yet contain considerable small shrubs and stump
sprouts from periodic clearing procedures. Most taxa within these sites are native successional species.
23) Old Homesteads.— Like any area that has been inhabited for long periods of time, Alleghany County
s and lots that have been abandoned and allowed to dilapidate. In many cases, the
chimney of a house may be visible. In other situations, only the level area and sur-
rounding vegetation provide evidence of a possible human-derived structure. In any case, these sites often
contain cultivated species that have either remained persistent (e.g., Chaenomeles speciosa, Cunninghamia lan-
ceolata. Thuja occidentalis, and Viburnum opulus var. opulus) or appear to have spread, mostly vegetatively, to
surrounding areas (e.g.. Aster tataricus and Lycium chinense). One aspect of this community category that is
particularly interesting is that it provides some insight into the historical preference of cultivated species in the
24) Residential and Urban Areas- This community differs from well maintained pastures and old home-
steads in regard to the close anthropogenic activity associated with it. In essence, residential areas are com-
prised of lawns, urban areas, and disturbed areas around gardens that provide habitat for the infiltration of
exotics, particularly those cultivated for aesthetic and/or consumption purposes. These introduced taxa may
readily, but sparingly become naturalized in small exposed areas adjacent to their point of origin. In some
cases, homes and yards are developed around streams, often allc
otic wetland
Poindexter, Flora of Alleghany County, North Carolina
:akly escaping or natui
im majus, Chionodoxa
ilized species in-
species (e.g., Glyceria declinata, Nasturtium officinale). Ej
elude herbaceous exotic taxa like Aegopodium podagra}
woody species like the native tree Crataegus phaenopyrum or exotic shrub Prunus tomentosa. Similarly, many
rural residences have vegetable gardens and waste heaps that occasionally provide a source of inoculation lor
species such as Solanum lycopersicum and S. tuberosum. Mulch beds also occasionally harbor adventives intro-
duced from intercalated seeds that sporadically germinate. • v, K a
25) Ponds and Reservoirs.— This community includes all manmade bodies of water within t e county, an
is treated here mainly because of their anthropogenic origin. These areas include smaUpondsandh^^^^^^^
of various sizes. Lake Louise, located at Roaring Gap Club, is the largest reservoir in the county (Ft^. & 3). It
was constructed in 1927, and other than general maintenance and recreational use, it has received httle altera-
tion It was built in close proximity to several known bogs and harbors many taxa around its margins with
bogdike affinities. Consequently, this lake and the vegetation that surrounds it ate likely relicts of a once natu-
ral community. Little Glade Mill Pond and Hare Mill Pond along the Blue Ridge Parkway are other examples of
such communities. The semi-natural aspect of these sites is problematic for ciassification purposes, and they
annear to be closely associated with Piedmont/Mountain Semipermanent Impoundment communities (&ha-
lale & Weakley 1990; Schafale 2012). Though many taxa found in these areas are exotta that are capitahmng
on the onen environment most are native. Serial wetland strata in this community include open water, free-
floating herbaceous species such as Lemmi minor, rooted noating herb species exemphWhy^^^^^^
phyllavar.hemropMlaandPotamogetondhersi/oliurandsubmer^^^^^^
These portions are the most bog-like and often contain many infrequent to rare taxa sue a
var. cuLens, Heodinris pnlustrls, and jnneus brevicnndams. Other notable rion-grammoid herbs me ude H-
poris loeselii, Lysimncliin terreslris, Trindenum virginienm, and Spimntlies lucida. Woody species are less com-
mon but usually include Alniis sernilata, Hypericum densiflorum, and Spiraea tomenlosn.
Z~Padge.t (2011), Alleghany county contalnsalarge number of Signifi^^^^
(SNHA) tracts for such a small county. Forty-eight SNHAs have been identified, four of which are Nationally
Lhcant,21thatare State Significant, 15 of Regional Sign, ficance,andseven that are
nificant. This high number ofSNHAsstands in stark contrast to the larger ptetureofoveraheavy^
the county, which would suggest that this area would have far fewer areas of significance than currently
'”""Xugh this area is not dominated by urbanization, the demands of agriculture have ultimately contnb-
u.edtoagfeat,ydistnrbed„^^^^^^^^^^^
ticularly evident in the numerous bog-hke remnants that nave Dcenc „cn.rP^ndromnlexvee-
draining practices throughout the area. To
etationofth^.^^^^^^^^
ciesiXuntds“nd
to thiirrarity and the factlthese rather disparatelyb^^^^^
onlyuncommoningeneral,butareoftenfoundinno^
habitats has reduced the natural area quality in Alleghany County, weax y
Poindexter, Flora of Alleghany County, North Carolina
553
Cursory examination of type
Bog, and Skunk Cabbage Bog) while remnants of natural \
ow bog communities. Although these disturbed areas stil
Schafale 1994), they are a bleak reminder of the enormous
e apparent by numerous disturbed mead-
:ew rare plants (as noted by Weakley and
)act on the natural heritage of this county.
NOTATED CHECKLIST
Nomenclature and plant origin (exotic vs. native) follow Weakley (2011) except where taxa were recently de-
scribed, have new nomenclatural combinations, follow a different taxonomic concept, or are not currently
recognized as occurring in the southeast or mid-Atlantic states (see Table 5). Plants of questionable nativity in
Weakley (201 1) are assigned status based on the PLANTS Database (USDA NRCS 2012). Along with traditional
infraspecihc ranks of variety and subspecies, three forms, five cultivars, and four potentially novel taxa are
geographic affinities that imply need for further study. Taxa are arranged alphabetically within each major
clade by family, genus, and species. Authorities are abbreviated according to the Brummitt and Powell (1992)
scheme, which is continuously updated and available online (Harvard University Herbaria 2012). Major clade
organization follows the PhyloCode as derived from Cantino et al. (2007) rather than a linear system (e.g.. Re-
veal 2012) to better reflect phylogenetic relationships and includes four primary groups: LYCOPODIOPHYTA,
MONILOPHYTA, ACROGYMNOSPERMAE, and ANGIOSPERMAE (here consisting of the Monocotyledon-
eae, Nymphaeales, Magnoliidae, and Eudicotyledoneae).
The scientific name of each taxon is preceded by a symbol denoting origin and invasive status as: natural-
from cultivation (A), taxa that appear to have escaped or are weakly spreading from cultivation (po = exotic, □ =
invasive, ^ = native), or a lack of notation for naturally occurring native taxa. The scientific name is then fol-
lowed by putative record status including: previously published state record (+), new state record (++), and
county record (o), where applicable. “Significantly Rare” taxa (see Table 2) are in bold type, and “Watch List”
taxa are underlined. A primary community of occurrence, a relative abundance value, representative voucher
specimen number(s), and respective repository conclude each taxon entry. An italicized voucher specimen(s)
number by the author is in a year-number (e.g., 08-274) format and corresponds to the primary collection
554
‘ Botanical Research Institute of Texas 7(1)
Symbols
t = Exotic Persistent From Cultivation
= Rich Cove Forest
= Rocky Bar and Shore
houseci at the Appalachian State University Herbarium (BOON). Taxa represented by collections from other
individuals are identified by the collector’s name and number, along with respective repository and date of col-
lection in brackets (e.g.J.L. Michael 792 [NCU, 25 June 1968]). Herbarium acronyms follow Index Herbario-
rum (Thiers, continuously updated).
Unvouchered records derived solely from the Flora of the Southeast Atlas (2012) are indicated with a
source reference: 1) “FSE-CVS” = Carolina Vegetation Survey data, and 2) “FSE-RAB” = Radford et al. (1968).
Similar records from the North Carolina Natural Heritage Program, though also partially integrated into the
Flora of the Southeast Atlas (2012) database, are represented separately as “NCNHP” due to rarity status. These
collective literature and sight reports are unverified and should be acknowledged with caution. Taxa that are
derived from these three sources are not included in the taxonomic summary.
Relative abundance is assigned here as inclusive for the entire study site and is adapted from Murrell and
Wofford (1987) and Estes (2005): Very Rare (V) = found in a single locale, usually in a small population; Rare
(R) = known from one to two localities, in small to moderate populations; Scarce (S) = several small or one to
two moderate to large populations; Infrequent (I) = scattered throughout in many small populations, or several
564
570
Despite severe anthropogenic influence and a heavily modified landscape, Alleghany County has a wealth of
flonstic diversity. This is also quite remarkable in light of the very small size of this study area. Part of this high
taxon diversity can be attributed to the rugged topography (inducing variable microclimates), disturbance
(high exotic richness), and the geographic positioning of the county, which adds a physiographic “ecotone”
effect (coupling of Mountain and Piedmont habitats). Ultimately, this flora is the most comprehensive survey
for Alleghany County at present, but can in no way be considered complete due to human factors (missed taxa)
and the dynamic nature of vegetation patterns (loss and gain of taxa over time). The usage of contemporary
techniques, particularly the digital documentation and georeferencing aspect of this project, will hopefully
serve as a model for making additional floristic projects more readily available, dynamic, and useful.
ACKNOWLEDGMENTS
First and foremost, this study would not have been possible without the gracious support of various donors at
or affiliated with Roaring Gap Club in Alleghany County. This work is a direct result of the private grant they
provided. It was a rarity to find so many willing individuals who desired to contribute to the preservation and
understanding of the flora of this area. I would like to thank James Padgett (North Carolina Natural Heritage
Program) for companionship and expertise while conducting fieldwork in Alleghany County, as well as the
numerous colleagues who accompanied me to various site visits. Likewise, I am indebted to the North Caro-
lina Natural Heritage Program for helping me to procure specific site information and locality data. I would
also like to extend my gratitude to the following specialists for their assistance with identifications and/or veri-
fications of various taxa reported in this publication: Wesley M. Knapp (Maryland Natural Heritage Program,
Juncus), David j. Keil (OBI, Centaurca), John C. Semple (WAT, miscellaneous Asteraceae), Paul M. Peterson and
Robert J. Soreng (US, Glyceria), Richard J. LeBlond (Dichanthelium, Panicum, and Paspalum), Richard D. Noyes
(UCAC, Erigewn strigosus diploids), and Robert F. C. Naczi (NYBG, Carex). In addition, I am grateful for the
curators at the following institutions for their assistance with loans and/or allowing me to peruse their collec-
tions (digitally and/or physically): Catawba College, CLEMS, DUKE, NCSC, North Carolina State Museum
of Natural Sciences, NCU, UMO, UNCC, USCH, and VPl. I would like to thank Alan S. Weakley, J. Richard
Abbott, Michael W. Palmer, and Ralph L. Thompson for their early reviews of this work. I would also like to
express my appreciation to Doug Goldman and John Taggart for their critical reviews and insightful sugges-
tions that improved the quality of this manuscript.
ITT, J.R. A
I. 2011
REFERENCES
ions in Phoradendron leucarpum (Viscaceae).
t. Res. Inst. Texas
5:139-141.
Abbott, Jr., R.N. and LA. Raymond. 1984. The Ashe Metamorphic Suite, northwest Nc
observations on geologic history. Amer. J. Sci. 284:350-375.
Alleghany County Historical Committee (ACHC). 1976. History of Alleghany County, 1
Carolina.
Alleghany Historical-Genealogical Society (AHGS). 1 1
Bailey, LH. 1924. Manual of cultivated plants. Macmillan Publishing Company, New York, New York.
Poindexter, Flora of Alleghany County, North Carolina
lary of plants cultivated in the United States and Canada.
I mountains. Univ. of North Carolina Press, Chapel F
Bailey, LH. and E.Z. Bailey. 1976. Hortus third. A concise d
MacMillan Publishing Co., New York, New York.
Bentley, S.L 2000. Native orchids of the southern Appala...,„..
Blair, A.E. 1 967. Vascular flora of Beaufort County, North Carolina. M.S. thesis. North Carolina State Universiti Raleigh
Braun, E.L 1950. Deciduous forests of eastern North America. Macmillan Publishing Company, New York New York
Brewer, E.O., R.M. Brown, and J.H. McIntyre. 1973. Soil survey of Alleghany County, North Carolina. United States Depart
ment of Agriculture, Soil Conservation Service, Washington, D.C
Britt, R.F. 1 960. The vascular flora of Robeson County, North Carolin
\. thesis, University of North Carolina, Chapel
5. Royal Botanic Gardei
e plants of North Carolina. North Caro-
Brummitt, R.K. AND C.E. Powell. 1 992. Authors of plant r
Buchanan, M.F. and J.T. Finnegan (eds.). 2010. Natural He
lina Natural Heritage Program, Raleigh.
Burnham, S.H. 1 906. A new species of Monotropsis. Torreya 6:234-235.
Canting, P.D., J.A. Doyle, S.W. Graham, W.S. Judd, R.G. Olmstead, D.E. Soltis, P.S. Soltis, and M J. Donoghue. 2007. Towards a phy-
logenetic nomenclature of Tracheophyta. Taxon 56:822-846.
Denslow, M.W. and D.B. Poindexter. 2009. Mentha suaveolens and M. xrotundifolia in North Carolina: a clarification of distri-
bution and taxonomic identity. J. Bot. Res. Inst. Texas 3:383-389.
Denslow, M.W., M.W. Palmer, and Z. E. Murrell. 2010. Patterns of native and exotic vascular plant richness along an eleva-
tional gradient from sea level to the summit of the Appalachian Mountains, U.S.A. J. Torrey Bot. Sci. 1 37:67-80.
Espenshade, G.H., D.W. Rankin, K.W. Shaw, and R.B. Neuman. 1975. Geologic map of the east half of the Winston-Salem quad-
rangle, North Carolina and Virginia: U.S. Geol. Survey Misc. Inv. Map 1-709-B, scale 1 .250000. United States Geological
Survey, Washington, D.C.
Estes, D. 2005. The vascular flora of Giles County, Tennessee. Sida 21:2343-2388.
Fenneman, N.M. 1 938. Physiography of eastern United States. McGraw-Hill Book Co., New York, New York.
Fernald, M.L 1950. Gray's manual of botany. Eighth edition. American Book Company, New York, New York.
Flo^ of North America Editorial Commttee, eds (FNA). 1 993-t-. Flora of North America north of Mexico. 1 4-f voIs. New York,
Flora of the Southeast (FSE). 2012. US Southeast flora atlas. University of North Carolina, Chapel Hill, North Carolina.
[http://www.herbarium.unc.edu/seflora/firstviewer.htm, accessed 1 3 Jan 201 2]
Gleason, H.A. and A. Cronquist. 1991 . Manual of vascular plants of northeastern United States and adjacent Canada. Sec-
ond edition. The New York Botanical Garden, Bronx.
Grant, E., and C. Epling. 1 943. A study of Pycnanthemum (Labiatae). Univ. Calif. Publ. Bot. 20:1 95-240.
Harvard University Herbaria (HUH). 2011. Index of botanists. [httpWasaweb.huh.harvard.edu:8080/databases/botanist_
index.html, accessed 2 January 2012].
Henderson, N.C. 1962. A taxonomic revision of the genus Lycopus (Labiatae). Amer. Midi. Naturalist 68:95-1 38.
Hitchcock, A.S. and A. Chase. 1950. Manual of the grasses of the United States, second edition. U.S. Dept, of Agriculture
Miscellaneous Publication No. 200 (reprinted in 1 971 by Dover Publications, New York).
Horton, J.H. 1 957. A vascular flora of Rowan County, North Carolina. M.A. thesis. University of North Carolina, Chapel Hill.
Huskins, S.D. and j. Shaw. 2010. The vascular flora of the North Chickamauga Creek Gorge State Natural Area, Tennessee.
Castanea 75:101-125.
Kartesz, J.T. 201 2. North American Plant Atlas. The Biota of North America Program (BONAP), Chapel Hill, North Carolina.
[http://www.bonap.org/MapSwitchboard.html, accessed 28 Jan 2012]
KOchler, A.W. 1964. Potential natural vegetation of the conterminous United States (map and accompanying manual).
American Geographical Society, Special Publication No. 36. New York, New York.
Memminger, E.R. 1915. A list of plants growing spontaneously in Henderson County, NC. J. Elisha Mitchell Sci. Soc.
31:126-149.
Michael, J.L 1 969. The vascular flora of Bullhead Mountain, Alleghany County, North Carolina. M.S. thesis. University of
North Carolina, Chapel Hill.
Murdock, N.A. 1994. Rare and endangered plants and animals of the southern Appalachian wetlands. Water, Air and Soil
Pollution 77:385-405.
Murrell, Z.E. and B.E. Wofford. 1 987. Floristics and phytogeography of Big Frog Mountain, Polk County, Tennessee. Cas-
tanea 52:262-290.
I. Agricultrual Statistics — Summary of Com-
modities by County; Alleghany County, [http://www.ncagr.gov/stats/codata/alleghany.pdf, accessed 2 Feb 201 2]
North Carolina Department of Agriculture and Consumer Services (NCDACS). 201 1 b. Agricultrual Statistics— County Estimates:
Christmas Trees. [http://www.ncagr.gov/stats/facts/ChristmasTrees.pdf, accessed 2 Feb 2012]
North Carolina Natural Heritage Program (NCNHP). 2012. Heritage Data Search, [http://portal.ncdenr.org/web/nhp/
database-search, accessed 1 Jul 2012]
Padgett, J.E. 201 1 . An inventory of the significant natural areas of Alleghany County, North Carolina. North Carolina
Natural Heritage Program, Raleigh.
Palmer, M.W. and J.C. Richardson. 2011. Biodiversity data in the information age: do 21st century floras make the grade?
Castanea 77:46-59.
Palmer, M.W., G.L Wade, and P.R. Neal. 1995. Standards for the writing of floras. BioSci. 45:339-45.
Patrick, T.S. 1984. Trillium sulcatum (Liliaceae), a new species of the southern Appalachians. Brittonia 36:26-36.
Peattie, D.C. 1928. Flora oftheTryon region of North and South Carolina: An annotated list of the plants growing spon-
taneously in Polk County, North Carolina, and adjacent parts of South Carolina, in Greenville and Spartanburg coun-
ties. Part 1 . introduction: soils, climate, etc, ferns and conifers (Pteridophyta, Gymnospermae). J. Elisha Mitchell Sci.
Soc. 44:95-1 25.
Peattie, D.C. 1 929a. Flora of the Tryon region of North and South Carolina: An annotated list of the plants growing
spontaneously in Polk County, North Carolina, and adjacent parts of South Carolina, in Greenville and Spartanburg
counties. Part 3. Willow family to Rose family (Salicaceae-Rosaceae). J. Elisha Mitchell Sci. Soc. 44:1 80-229.
Peattie, D.C. 1929b. Flora oftheTryon region of North and South Carolina: An annotated list of the plants growing
spontaneously in Polk County, North Carolina, and adjacent parts of South Carolina, in Greenville and Spartanburg
counties. Part 2. Cat-tail family to Orchid family (Typhaceae to Orchidaceae). J. Elisha Mitchell Sci. Soc. 44:141-180.
Peattie, D.C. 1929c. Flora oftheTryon region of North and South Carolina: An annotated list of the plants growing
spontaneously in Polk County, North Carolina, and adjacent parts of South Carolina, in Greenville and Spartanburg
counties. Part 4. Mimosa family to Dogwood family (Mimosaceae to Cornaceae). J. Elisha Mitchell Sci. Soc. 45:59-1 00.
Peattie, D.C. 1930. Flora oftheTryon region of North and South Carolina: An annotated list of the plants growing spon-
taneously in Polk County, North Carolina, and adjacent parts of South Carolina, in Greenville and Spartanburg coun-
ties. Part 5. Wintergreen family to Lobelia family (Pyrolaceae to Lobeliaceae). J. Elisha Mitchell Sci. Soc. 45:245-290.
Peattie, D.C. 1931. Flora of the Tryon region of North and South Carolina: An annotated list of the plants growing
spontaneously in Polk County, North Carolina, and adjacent parts of South Carolina, in Greenville and Spartanburg
counties. Part 6. Daisy family (Compositae). List of new names published. Errata. Summary. J. Elisha Mitchell Sci. Soc.
46:129-160.
Peattie, D.C. 1937. Additions, corrections, and deletions for the flora of the Tryon region. J. Elisha Mitchell Sci. Soc.
53:311-323.
Penny, M. 2010. We Drivers': Alleghany County's Relationship with the Automobiie.
University Library, The University of North Carolina at Chapel Hill. [http://do(
overlooks/we_drivers/, accessed 1 1 February 201 2]
Perry, K.B. 1998a. Average growing season for selected North Carolina locations. NC State University, Horticulture Leaf-
e Extension Service, [http://www.ces.ncsu.edu/depts/hort/hil/hil-709.html, accessed
Perry, K.B. 1998b. Average last spring frost dates for selected North Carolina locations. NC State University, Horticulture
Leaflets. North Carolina Cooperative Extension Service, [http://www.ces.ncsu.edu/depts/hort/hil/hil-707.html, ac-
cessed 11 Jan 2012]
Poindexter, D.B. 2006. Eight new plant distributional records to Alleghany County, North Carolina. J. North Carolina Acad.
Sci. 122:101-105.
North Carolina. J. Bot. Res. Inst. Texas 2:649-650.
Poindexter, D.B. 2010a.Ab/es//rm£? (Pinaceae) naturalized in North America. Phytoneuron 2010-41:1-7.
Poindexter, D.B. 2010b. Persicaria perfoliata (Polygonaceae) reaches North Carolina. Phytoneuron 2010-30:1-9.
Poindexter, D.B. 2012. Vascular flora of Alleghany County, North Carolina database. Appalachian State University, I.W.
Carpenter, Jr. Herbarium, Boone, North Carolina 28608. [www.vascularflora.appstate.edu, 14 January 2012]
Poindexter, D.B. and R.L. Unce. 201 1 . Crataegus chrysocarpa var. dodgei (Rosaceae) new to North Carolina. J. Bot. Res. Inst.
Texas 5:349-350.
573
Poindexter, D.B. and Z.E. Murrell. 2008. Vascular flora of Mount Jefferson Sti
North Carolina. Castanea 73:283-237.
Poindexter, D.B. and J.B. Nelson. 201 1. A new hedge-nettle (Stachys: Lamiaci
tains. J. Bot. Res. Inst. Texas 5:405-414.
Poindexter, D.B., A.S. Weakley, and M.W. Denslow. 201 1 . New exotic additions a
North Carolina. Phytoneuron 201 1-42:1-14.
PySek, P., D. M. Richardson, M. RejmAnek, G.L. Webster, M. Williamson and J. Kirschn
towards better communication between taxonomists and ecologists. Tc
Rankin, D.W., G.H. Espenshade, and R.I
North Carolina, Virginia, and Ten
logical Survey, Washington, D.C.
Radford, A.E., H.E. Ahles, and C.R. Bei
Press, Chapel Hill.
Rehder, a. 1937. Manual of cultivated trees and shrubs. The Macmillan Company, New York, New York.
Reveal, J.L. 2012. An outlineof a classification scheme for extant flowering plants. Phytoneuron 2012-37:1-221.
Rothrock, P.E., A.A. Reznicek, and C.T. Bryson. 201 1. Geographic range and morphological and chromosomal variability
Carex molestiformis (Cyperaceae) east of the Mississippi River. Castanea 76:1 78-1 82.
Schafale, M.P. 2012. Guide to the nal
ither noteworthy records for the flora of
304. Alien plants in checklists and floras:
53:131-143.
IN. 1972. Geologic map of the west half of the Winston-Salem quadrangle,
U.S. Geol. Survey Misc. Inv. Map 1-709-A. Scale 1 :250,000. United States Geo-
. Manual of the vascular flora of the Carolinas. University of North Carolina
mmunities of North Carolina, fourth approximation. N.C. Natural Heriti
s. httpV/www.ir
e Summary. [httpV/www.sercc.
V. Press, Cambridge.
s. In: Martin, W.H., S.G. Boyce, and A.C. Echl
nities. John Wiley and Sons, N
n State Park in North Carolina. M.S. thes
Schafale, M.P. and A.S. Weakley. 1990. Classification of the natural
N.C. Natural Heritage Program, Div. of Parks and Recreation, Raleigh.
SCOTFORD, D.M. and J.R. Williams. 1 983. Petrology and geochemistry of metamorphosed ultramafic bodies in a portion of
the Blue Ridge of North Carolina and Virginia. Amer. Mineralogist 68:78-94.
Small, J.K. 1933. Manual of the southeastern flora. The University of North Carolina Press, Chapel Hill, North Carolina.
SoRRiE, B.A., J.B. Gray, and PJ. Crutchfield. 2006. The vascular flora of the longleaf pine ecosystem of Fort Bragg and Wey-
mouth Woods, North Carolina. Castanea 71:129-161.
Southeast Exotic Pest Plant Council (SE-EPPC). 2012. Invasive plants of the thirteen southern st
org/seweeds.cfm.
Southeast Regional Climate Center (SRCC). 201 1.Transou, North Carolina (318694)-
com/cgi-bin/sercc/cliMAIN.pl?nc8694, accessed 13 Jun 2012]
Stage, C. 201 0. New flora of the British Isles, third edition. Cambridge
Stephenson, S.L, A.N. Ash, and D.F. Stauffer. 1993. Appalachian Oak Forests. In: Martin,
nacht, eds. Biodiversity of the southeastern United States: Upland terrestrial comm
York, New York. P. 255-303.
Taggart, J.B. 1 973. Floristic survey and vegetational analysis of Stone Mountain
North Carolina State University, Raleigh.
Taggart, J.B. 1 976. Additions to the flora of Alleghany and Wilkes Counties, North Carolina. Castanea 41 :342-346.
Thiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New
York Botanical Garden's Virtual Herbarium. [httpV/sweetgum.nybg.org/ih/, accessed 1 2 May 201 2]
Trewartha, G.T. AND L.H. Horn. 1 980. An introduction to climate, 5th ed. McGraw Hill Book Company, New York, New York.
United States Census Bureau (USCB). 201 2. Annual estimates of the resident population for counties: April 1 , 201 0 to July
1 201 1 [http//www.census.gov/popest/data/counties/totals/201 1/CO-EST201 1 -01 .html, accessed 20 Jun 201 2]
Un^d States Department of Agriculture, Natural Resources Conserva.on Service (USDA, NRCS). 2012. PLANTS Database. Na-
tional Plant Data Team, Greensboro, North Carolina 27401-4901, USA. [http://plants.usda.gov, accessed 13 Jan 2012]
United States Geological Survey (USGS). 2012. North Carolina geology, [http://mrdata.usgs.gov/sgmc/nc.html, accessed
Wade, G.L. and R.L. Thompson. 1991. The species-area curve and regional floras. Trans. Kentucky Acad. Sci. 52:21-25.
Weakley, A.S. 2011. Flora of the southern and mid-Atlantic states. Working draft of 15 May 2011. University of North
Carolina Herbarium, North Carolina Botanical Garden, Chapel Hill.
Weakley, A.S., RJ. LeBlond, B.A. Sorrie, C.T. Witsell, L.D. Estes, K. Gandhi, K.G. Mathews, a
rank changes, and nomenclatural and taxonomic comments in the vascular fl
J.Bot. Res. Inst. Texas 5:349-350.
574
Journal of the Botanical Research Institute of Texas 7(1)
Weakley, A.S. and M.P. Schafale. 1994. Non-alluvial wetlands of the southern Blue Ridge— dive
tern. Water, Air Soil Pollut. 77:359-383.
WicHMANN, B.L 2009. Vegetation of Geographically Isolated Montane Nonalluvial Wetlands c
of North Carolina. M.S. Thesis, North Carolina State University, Raleigh.
Williams, J.R. 1979. A geochemical investigation of the metasomatized ultramafic bodies in
rangle, northwestern North Carolina— southwestern Virginia. M.S. thesis, Miami Universit
Wofford, B.E. 1989. Guide to the vascular plants of the Blue Ridge. University of Georgia Pres
Wood, T.F. and G. McCarthy. 1 886. Wilmington flora. A list of plants growing about Wilmington
of flowering. With a map of New Hanover county. Edwards, Broughton & Co., power print
of the Southern Blue Ridge
1 the Winston-Salem Quad-
ity, Oxford, Ohio,
ss, Athens.
n. North Carolina, with date
ters and binders, Raleigh.
VASCULAR PLANTS OF THE CANYONLANDS UNIT OF THE
BIG THICKET NATIONAL PRESERVE, TYLER COUNTY, TEXAS
Kelly C Haile
Stephan L Hatch
Department of Ecosystem Science and Management
Texas A&M University
Texas A&M University
2138TAMU
College Station, Texas 77843, U.S.A.
2138TAMU
College Station, Texas 77843, U.S.A.
kelly.haile@hotmail.com
INTRODUCTION
This paper provides the first checklist of the vascular plants in the Canyonlands Unit of the Big Thicket Na-
tional Preserve. The checklist will be utilized by ecologists and botanists interested in the flora of the Big
Thicket National Preserve and southeastern Texas. It provides a list of plants that are currently found within
this area. Wildlife biologists and wildlife ecologists can benefit from this checklist by using the data for imple-
menting management practices within the unit. This information could also be helpful in determining the
different wildlife species within the area. The checklist indicates the invasive/noxious plant species in the area
and can serve as a starting point for tracking and managing these invasive species.
SITE DESCRIPTION
The Big Thicket National Preserve is located in the Pineywoods vegetation area (Hatch et al. 1990) of south-
eastern Texas. Established in 1974, it became America’s first National Preserve with approximately 34,803
hectares (Peacock 1994). In 1981 the Preserve was designated a UNESCO Biosphere Reserve by the United
Nations, and in 2001 the American Bird Conservancy designated the preserve as a Globally Important Bird
Area. Today the preserve consists of just over 40,468.6 hectares, comprising 15 units that spread over parts of
seven different counties in eastern Texas (Watson 2006). This project focuses on the Canyonlands Unit of the
Big Thicket National Preserve (Fig. 1). The Big Thicket National Preserve acquired the Canyonlands Unit in
1993. It consists of 597.3 hectares, located on the eastern border of Tyler County along 5.63 km of the Neches
River, on the east side of the unit.
Within these hectares are a variety of environments, including upland habitat (Fig. 2), wetland and riparian
areas (Fig. 3), cypress swamps and pineywoods (Fig. 3). The Canyonlands Unit is made up of various canyons
and topography changes with elevations ranging from 18.3-61 m above sea level. A floristic study was con-
ducted on this unit to determine the number of the vascular plant species within the Canyonlands Unit. This
floristic study occurred over parts of four years, November 2008-November 2011.
The climate for this area is described as subtropical (MacRoberts 2008). The average temperature for Tyler
J.Bot. Res. Inst. Texas 7(1): 575 -5
576
Journal of the Botanical Research Institute of Texas 7(1)
County is 18.8-20°C with an average annual rainfall of 127-137 cm (CoCoRaHS 2012). This region has the
highest average annual rainfall in Texas (MacRoberts 2008). However, during the study period this area of
Texas received below average rainfall for both 2010 and 2011, 93 cm and 79 cm respectively.
The Canyonlands Unit is made up of a variety of 20 different soil types: sand, silt, clay or a combination of
all three. The most abundant soil is the Estes Angelina Complex, which makes up 36% of the unit. It is a clay-
dominated soil and is found in the bottomlands and low-lying areas. The second most abundant soil is the
Woodville Fine Sandy Loam, which occurs on the slopes of the main canyons within the unit. For more specif-
ics on soils, see Haile (2012).
METHODS
Eleven collection trips were made over the three year period. Trips were made during the different growing
seasons in order to collect plant species while they were in flower and/or fruit. Each collection site within the
unit was visited several times throughout the project to make a more complete checklist of the vascular plants
within the Canyonlands Unit (Haile 2012). A voucher specimen and/or a photo were taken to document the
species within the unit. The specimens of each species included in this checklist are housed in the S.M. Tracy
Herbarium (Texas A&M University) including other holdings of the National Park Service.
Collection sites were determined based on soils, topography, and vegetation communities in an area. Dur-
ing collection trips up to four collectors were present in order to better cover each location. At each site the
collectors would collect all the flowering vascular plant species that had not previously been collected. Once
collections were made, each specimen received collection numbers, site descriptions, an associated species list,
soil description and GPS locations. Before leaving the site, each specimen was pressed in a plant press and set
to dry, later keyed, identified and verified.
Identifications for the species were made by using the following books, including: Illustrated Flora of North
Central Texas (Diggs et al. 1999), Illustrated Flora of East Texas, Volume 1 (Diggs et al. 2006), Aquatic and Wetland
Plants of Southeastern United States: Dicotyledons (Godfrey & Wooten 1981), Aquatic and Wetland Plants of
Southeastern United States: Monocotyledons (Godfrey & Wooten 1979), and Gould's Grasses of Texas (Hatch
2010), Trees, Shrubs, & Woody Vines of East Texas (Nixon 1985). Hatch (2010) was used for the current scientific
names of all the Poaceae. The current scientific names for all other families and species were found at USDA
Plants, and Diggs et al. (1999, 2006). Angiosperm Phylogeny Group (APG) classification system was used for
placement of genera within families (Angiosperm Phylogeny website).
RESULTS AND DISCUSSION
The Canyonlands Unit has 103 families, 246 genera, and 388 species and 16 infraspecific taxa (Appendix 1).
Figure 4 and Figure 5 are two species collected and documented for the unit. Of these 388 species, 29 are intro-
duced and 3 are considered invasive/noxious. An invasive species is defined as a plant whose introduction
causes or is likely to cause economic harm, environmental harm, or harm to human health (www.invasive.
org). Invasive species are weedy species; they can be either native or introduced. The plants listed as invasive
species are listed on the invasive and noxious weed list for Texas (USDA Plants).The four largest families in this
unit were Poaceae (68 species), Asteraceae (43 species), Cyperaceae (23 species), and Fabaceae (18 species).
Floristic studies and the resulting species checklist of an area are never complete. It is estimated that this
list includes 90 percent of the plant species within the unit. Due to the dry weather conditions that occurred
during this study, some of the cool season annuals may not have been observed and collected. The next goal is
to make future collecting trips to this unit in order to continue to add to this species checklist.
Appendix 1 is the annotated checklist of the vascular plant species observed and collected within the
Canyonlands Unit of the Big Thicket National Preserve. The voucher specimens are housed at the S.M. Tracy
Herbarium (TAES). The checklist is arranged first by phylum (Pteridophyta, Pinophyta, Magnoliophyta).
Families are arranged alphabetically within phylum, genera are arranged alphabetically within family, and
species are arranged alphabetically within each genus. The scientific name and authority is given for each of
the species (Hatch et al. 1990; Diggs et al. 1999; Diggs et al. 2006; Hatch 2010), along with the longevity, origin.
577
579
Fig. 3. Bonomland area within the Canyonlands Unit of the Big Thicket National Preserve, Tyier County, Texas (Photo by S.L. Hatch).
580
584
Styraxgrandifolius Ait, KH; N, P, C
U/mus o/ato Michx., KH 863; N, P, C
Ulmus americana L, AG 514; N, R C
Viola primulifolia L, KH 678; N, P, C
This projected was funded by the Big Thicket Association and by Mrs. Lucile Gould Bridges (Frank W. Gould
Research Award). We also thank Aminda Gallardo, Steven Goertz, Katherine Haile, Robert Haile, Tyler Hatch,
Dale Kruse, George Umphres, and Erin Wied for their contributions. Thanks to the staff at the Big Thicket
Field Research Station in Saratoga, Texas for allowing us to use their facilities during field work. Also, we offer
many thanks to two anonymous reviewers for improving this publication. Thanks to Katherine E. Winsett for
help with Figure 1.
Botanical Research Institute of Texas. 2011-2012. http://www.brit.org/herbarium. Accessed March 2012.
Brown, LE., B.R. MacRoberts, W.W. Pruess, I.S. Elsik, and S.B. Walker. 2008a. Annotated checklist of the vascular flora of the
Loblolly Unit of the Big Thicket National Preserve, Liberty County, Texas. J. Bot. Res. Inst. Texas 2:1 481-1 489.
Brown, L.E., B.R. MacRoberts, M.H. MacRoberts, W.W. Pruess, I.S. Elsik, and S.B. Walker. 2008b. Annotated checklist of the
vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. J. Bot. Res. Inst. Texas
2:651-660.
Brown, L.E., B.R. MacRoberts, M.H. MacRoberts, P.A. Harcombe, W.W. Pruess, I.S. Elsik, and S.D. Jones. 2006. Annotated check-
list of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County, Texas. Sida
22:1175-1189.
Brown, L.E., B.R. MacRoberts, M.H. MacRoberts, P. Harcomb, W.W. Pruess, S. Elsik, and D. Johnson. 2005. Annotated checklist of
the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin County, Texas. Sida
21:1807-1827.
Community Collaborative Rain, Hail Snow Network (CoCoRaHS). 2012. http://www.cocorahs.org/. Accessed February 2012.
Diggs, G.M., Jr., B.L. Lipscomb, and RJ. O'Kennon. 1 999. Shinners & Mahler's illustrated flora of North Central Texas. Botanical
Research Institute of Texas, Fort Worth.
Diggs, G.M., Jr., B.L. Lipscomb, RJ. O'Kennon, and M.D. Reed. 2006. Illustrated flora of East Texas, volume one. Introduction,
pteridophytes, gynosperms, monocotyledons. Botanical Research Institute of Texas, Fort Worth.
Godfrey, R.K. and J.W. Wooten. 1 979. Aquatic and wetland plants of the southeastern United States: Monocotyledons. The
University of Georgia Press, Athens.
Godfrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of the southeastern United States: Dicotyledons. The
University of Georgia Press, Athens.
Haile, K.C. 2012. Vascular plant survey of the Canyonlands Unit of the Big Thicket National Preserve, Tyler County, Texas.
Thesis, Texas A&M University, College Station.
Hatch, S.L 1990, K.N. Gandhi, and LE. Brown. 1990. Checklist of the vascular plants ofTexas. MP-1655. Texas Agric. Exp.
Sta., College Station.
Hatch, S.L. 2010. Gould's grasses ofTexas. Copy Corner, College Station, Texas.
Haile and Hatch, Flora of the Canyonlands Unit, Big Thicket
585
e Big Thicket National Preserve, Tyler County, T
MacRoberts, M.H. and B.R. MacRoberts. 2
2(1):665-671.
MacRoberts, M.H. and B.R. MacRoberts. 2008. The Big Thicket as floristically u
665-671.
Nixon, E.S. 1985.Trees, shrubs, and woody vines of East Texas. Bruce Lyndon Cunningham, Nacogdoches, Texas.
Peacock, H. 1 994. Nature lover's guide to the Big Thicket. Texas A&M University Press, College Station.
Texas Water Development Board. 201 2. http://www.twdb.state.tx.us/. Accessed February 2012.
Thunder Snow Interactive. 201 1. http://www.thundersnow.com/pdfdocs/weedlistv4sci.pdf. Accessed November 2012.
University of Texas Herbaria. 2011-2012. (TEX-LL) httpy/www.biosci.utexas.edu/prc/types.html. Accessed October 2011.
USDA, NRCS. 2009-201 2. The PLANTS Database http://plants.usda.gov. National Plant Data Center, Baton Rouge, Loui-
siana 70874-4490, USA. Accessed September 2011.
USDA, NRCS. 2012. Web Soil Survey http://websoilsurvey.nrcs.usda.gov. Soil Survey Staff, Las Cruces, New Mexico
88003-0003, USA.
USFWS. May 1988. National list of plant species that occur in wetlands. St. Petersburg, Florida 33702, USA. Accessed
October 2011.
Watson, G.E. 2006. Big Thicket plant ecology - an introduction 3rd edition. University of North Texas Press, Denton.
586
ANNOUNCEMENT
Report from a New Botanical Nomenclature Course
Kanchi N. Gandhi
Harvard University Herbaria (HUH)
22 Divinity Avenue
Cambridge, Massachusetts 02138, U.S.A.
A Botanical Nomenclature Course workshop was jointly organized by the Botanical Survey of India (BSI)
and ENVIS Centre on Floral Diversity-BSI in Kolkata from January 11-13, 2013 at the Hotel Presidency Inn
Pvt. Ltd., Kolkata. This workshop was the first of its kind organized in India. Dr. Kanchi N. Gandhi (Senior
Nomenclatural Registrar, Harvard University Herbaria, USA) was the Course Director; Dr. Paramjit Singh
(Director, BSI) was the Course Convener; and Dr. P. Lakshminarasimhan (Scientist ‘D’, BSI) was the Course
Coordinator. The BSI botanist late Dr, Mithilesh K. Pathak was helpful in organizing the workshop. There were
96 total participants, 51 from BSI and 45 from outside BSI. The participants included Research Fellows/Associ-
ates from BSI, faculties from universities/colleges, scientists from research institutes, one Project officer from
Foundation for Ecological Security (Gujarat), and a Deputy Director of Public Instruction (UGC), Education
Directorate, Govt, of West Bengal, Kolkata.
On the inaugural day. Dr. Lakshminarasimhan welcomed the gathering and outlined the course program.
His talk was followed by two lectures; 1) The importance of Botanical Nomenclature by Prof. P.K. Mukherjee
(Calcutta University) and 2) Taxonomy and Global Strategy for Plant Conservation by Dr. D.K. Singh (Scientist
‘F’, BSD. Two lectures were given by Mr. R.L. Mitra (Scientist (Retd.), BSI): 1) Scientific Names of Plants — Some
Facts and Misconceptions (on January II) and 2) Type Concept and Nomenclatural Problems (on January 12).
Dr. Gandhi’s lectures were spread over the three days, and he covered the following topics:
1) An historical overview of the botanical nomenclature from Linnaeus to the Melbourne Code
2) Review of the physical structure of the Melbourne Code
3) Preamble, ranks and names of taxa, and effective publication (Articles 1-28)
4) Validity of names. Part I (Articles 32-45)
5) Validity of names. Part II
6) Authorship citation (Articles 46-50)
7) Rejection of names (Articles 51-60)
8) Anamorphic fungal and hybrid names
In the concluding function on January 13, there was a lecture by Dr. Paramjit Singh on “Floristic Diversity in
India: An overview.” After his talk, certificates were distributed to all the participants. Valedictory remarks
were delivered by: Dr. R.K. Chakraverty (Additional Director (Retd.), BSI), Dr. Kanchi N. Gandhi, Dr. P. Laksh-
minarasimhan, Prof. G.G. Maity (Retd; Kalyani University), Prof. N.D. Paria (Calcutta University), Dr. D.K.
Singh, and Dr. Paramjit Singh.
ELEOCHARIS MICROFORMIS (CYPERACEAE):
REDISCOVERED IN NORTH AMERICA FROM THE EDWARDS PLATEAU
AND TRANS-PECOS REGIONS OF TEXAS
Robert J. O'Kennon
Botanical Research Institute of Texas
1700 University Drive
Fort Worth, Texas 76107, USA.
Kimberly Norton Taylor
1700 University Drive
Fort Worth, Texas 76107, U.S.A.
ktaylor@brit.org
ABSTRACT
(L.) Roem. & Schult. and E. atropurpuea (Retz.) J. Presl & C. Presl. Henry K. Svenson placed E. microformis in synonymy under E. geniculata
INTRODUCTION
Eleocharis microformis Buckley, in subgenus Eleocharis sect. Eleogenus, subser. Rigidae (Gonzalez-Elizondo &
Peterson 1997), was described by Samuel Bostford Buckley in 1862 (Buckley 1862) from a specimen collected
in “northern Texas” (Buckley s.n., PH). Later that year Asa Gray remarked that E. microformis “[a hybrid name]
is near E. intermedia, Schultes” (Gray 1862). Buckley responded by saying E. microformis “is probably a good
species” (Buckley 1870). Svenson (1939) reduced E. microformis to a synonym under E. geniculata (L.) Roemer
& J.A. Schultes and the name has been overlooked in taxonomic treatments for the last 74 years. Although
described from Texas, E. microformis has never been mentioned either as a species or as a synonym in any
Texas or North American flora or checklist (Gould 1962; Correll & Johnston 1970; Correll & Correll 1972;
Godfrey & Wooten 1979; Hatch et al. 1990; Kartesz 1994; Jones et al. 1997, Diggs et al. 1999; Kartesz &
Meacham 1999; Smith et al. 2003). Hooker and Jackson (1895), in the first volume of Index Kewensis, following
Gray’s comment gave Eleocharis intermedia Schult. as a synonym of E. microformis. Eleocharis intermedia is a
species in section Eleocharis, known from the northeastern United States, with golden-brown, compressed-
trigonous achenes with narrowly pyramidal tubercles. In contrast, E. microformis is in section Eleogenus with
The epilaphs Eleocharis gemculala. E. caribaca (Rottb.) S.F.BIake, E. microformis. and E elcgans R^m. & Schult.
have been confused for decades (Svenson 1929, 1937, 1939, 1957). Henry Knute Svenson, who for decades
worked on the genus Eleocharis, recognized E. microformis as a distinct species in the first part of his Mono-
graphic studies in the genus Eleocharis'^ (Svenson 1929). Of the collections from six different lo-t,nn. th.i
Svenson examined, he annotated each one as E. microformis at least once a
1937. From 1948 to 1967 he re-annotated most of these specimens as E. car
Journal of the Botanical Research Institute of Texas 7(1)
treated as a synonym of E. geniculata. He stated, “The dwarf round-headed E. microformis from Texas with
achenes 0.7-0.9 mm long, often confused with E. atropururea, I now believe to be a small extreme of E. genku-
lata (E. caribaea), similar to specimens from Nicaragua (Maxon, Harvey, & Valentine 7291, NY) and from Hon-
duras (Schipp No. 913 & Standley No. 56671, NY)” (Svenson 1939). Upon recent examination by O’Kennon, the
collections from Honduras appeared to be smallish but otherwise typical E. geniculata. The collection from
Nicaragua was a short E. geniculata with smaller than usual achenes (0.7-0.8 mm long), but had none of the
characteristics of E. microformis. Svenson re-annotated a Reverchon collection (1672, TEX) of E. microformis
from Blanco County, Texas as E. caribaea with the comment that it was a “form with small heads and small
achenes . .
Of the five taxa within subser. Ridigae, only E. atropurpurea (Retz.) J. Presl & C. Presl and E. geniculata
are widespread in North America (Gonzalez-Elizondo & Peterson 1997,). Eleocharis bahamensis Boeckeler and
E. sintenisii Boeckeler are endemic to the Bahaman islands and Puerto Rico, respectively, and E. microformis is
endemic to Texas. Upon cursory examination, Eleocharis microformis resembles both E. atropurpurea and E.
geniculata. Herbarium specimens of E. geniculata and E. atropurpurea were examined from the following her-
baria: BRIT, SMU, TEX-LL, GH, MO, US, NY, and TAES. Twenty-one specimens (including duplicates) were
identified matching Buckley’s description of E. microformis. Upon examination of specimens of E. geniculata,
E. atropurpurea, and E. microformis, it became immediately obvious to us that E. microformis was distinct and
supported the resurrection of Buckley’s original name.
Eleocharis atropurpurea is easily distinguished from the other species by its small (0.4-0.5 mm) black
achenes with a small subconic tubercle usually less than Va, as wide as the achenal body, generally with bright
white bristles shorter than to equaling the achenal body (Table 1). Eleocharis microformis is distinguished from
E. geniculata by its smaller achenes (less than 0.65 mm vs. greater than 0.7 mm) with a larger tubercle (Vi as
broad as the achene vs. Vi as broad), fewer scales per spikelet (generally fewer than 20 vs. more than 28), shorter
scales (1 mm vs. 1.5-2 mm), shorter bristles (shorter than or equaling achene length vs. longer than achene),
thinner culms (0.2-0.3 mm vs. 0.5-1.0 mm), and arching to ascending habit (vs. erect to ascending) (Table 1,
Fig. 1).
Eleocharis microformis and E. geniculata are sympatric on the Edwards Plateau, and we have observed
them growing side by side on four sites. Although there are minor variations in size within each species and
occasional overlaps in size between these species, no recognizable intermediates have been observed. El-
eocharis microformis also occurs on the same sites with E. atropupurea, a diminutive species with which it is
often confused in the field but is distinguished by its smaller tubercle and achenes (see key and Figures 1 & 2
The first known collections of what Buckley later called Eleocharis microformis were made in July of 1852 by
Charles Wright (1930, 1932, 1961 GH) while he “was on the Pecos on his way back to San Antonio (Shaw
1987).” Label information for no. 1930 indicates “Howard Creek, Texas” on 4 July. Howard Creek meets the
Pecos River in northwestern Val Verde County, about 40 km (25 mi) downriver from Live Oak Creek where
Wright made collections two days prior (Gray 1852-1853). This is likely the “Howard Creek” cited on the
specimen. Numbers 1932 and 1961 lack locality information but are probably in the vicinity (Svenson 1929).
Eleocharis microformis was next found by Buckley in June 1861 (s.n., PH), and described by him in 1862.
In the protologue of his collection, Buckley did not include an exact location for that 1861 collection other than
“northern Texas.” According to Dorr and Nixon (1985), Buckley worked as a geologist and naturalist for the
Geological Survey of Texas in June 1861, and he traveled from Austin to work on the survey of Navarro County
about 250 km north of Austin. This apparently was his northern-most excursion in Texas and the specimen
was likely collected somewhere along his route.
Reverchon collected E. microformis in 1885 from “swamps” in Blanco Co. (1672, GH, LL, MO, NY, US;
3594, MO, NY, US), and from “Texas” (s.n. GH). Reverchon’s collection number 1673 of E. geniculata (Reverchon
1673, MO) was from the banks of the Pedernales River in Blanco Co. It is likely that his “swamps” are also
590
Fw. 2. Photographs of achenes of three closely related taxa. From left to right: Eleocharisatmpurpurea {O'Kennon 10663). Eleocharismicrofomis (O'Kennon
10171). Eleocharis geniculata (O'Kennon 10710).
Estes, Proctor, and Benesh found an additional site of E. microformis in 1995 in Bexar Co. (BUL0856, BRIT).
This makes a total of approximately 18 sites in 14 counties, since one of O’Kennon’s sites is likely the same as
Heller’s 1894 site.
A field survey was made by O’Kennon in July 2009 to relocate all recent and historic sites of E. microfor-
mis. Plants were found at only five of the 14 locations known at the time (the site in Bexar Co. was not searched
due to restricted access, the sites in Hudspeth and El Paso counties and Wright’s “Howard Creek” were un-
known to us at the time and have not been searched) with a total of only 37 plants observed. The only historical
site relocated was south of Kerrville along Turkey Creek near Hwy 16 which could have been Heller’s 1894
location “about Kerrville”. The other four locations were at sites located by O’Kennon in 1991 and 1992 (Fig. 3).
An extensive search for the plants was made by O’Kennon in Blanco County, but none were found. Places that
could have been Reverchon’s “Swamps” were not found. Plants were never found near Lake Austin, south of Big
Lake, or in Edwards Co. southeast of Rocksprings.
Conservation Status
There are only 18 known locations for Eleocharis microformis in the world in 14 Texas counties. As of 2009, only
5 of the 14 sites surveyed had extant populations. Seven of the historic collection locations have not been seen
in over 70 years and may be extirpated. With so few sites, this Texas endemic is rare and perhaps warrants
conservation status.
KEY TO NORTH AMERICAN ELEOCHARIS (sUBGENUS ELEOCHARIS SECT. ELEOGENUS SUBSERIES RIGIDAe)
NOTE, IN THIS KEY, THE ACHENE IS THE ACHENAL BODY NOT INCLUDING THE TUBERCULE
O'Kennon and Taylor, Eleocharis microformis rediscovered in Texas 591
Eleocharis microformis Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:5-10. 1862. type: U.S.A. Texas: northern Texas Jun
Plants annual, densely tufted, without creeping rhizomes. Culms arching to ascending, setaceous, lightly
sulcate (l-)3-6(-9) cm long, 0.2-0.3 mm thick. Leaves: distal leaf sheaths firm, distally tightly sheathing,
apex acute Spikelets ellipsoid to ovoid, acute, 10-20(-26)-flowered, 2-3(-4) mm long; floral scales obtuse,
often rufescent, with a greenish-yellow midrib and light brown hyaline margins. Howers: perianth bristles
4-6, light-brown to whitish, generally shorter than the achene, retrorsely barbed (120x); styles bifid. Achenes
broadly obovoid-pyriform, shiny black when ripe, biconvex, 0.5-0.65 x 0.4-0.5 mm, not constricted proximal
to tubercle, surface minutely pitted at 40x. Tubercles white, depressed-umbonate, at least Vi as broad as the
acnene. i i •
Fruiting Jun-Aug. Wet areas including pond margins, creeks, riverbanks, and ephemeral swales m a va-
riety of soil substrates including sand, clay, and granite. Specimens examined with known site locations are
from central and west Texas. The exact location of the type collection protologue from “northern Texas is
unknown; elevation 258-1455 m.
Associated plants include; Ammannia coccinea Rottb., Ammannia robusta Heer & Regel, Lysmachia mini-
ma (L.) U.Manns & Anderb., Centella erecta (L.f.) Fernald, Cyperus acuminatus Torr. & Hook., Cyperus haspan
L., Cyperus squarrosus L., Eleocharis atropurpurea (Retz.) J. Presl & C. Presl, Eleocharis coloradoensis (Britton)
Gilly, Eleocharis engelmannii Steud., Eleocharis geniculata (L.) Roem. & Schult., Eleocharis obtusa (Willd.)
Schult., Fimbristylis autumnalis Roem. & Schult., Fimbristylis vahlii (Lam.) Link, Fuirena simplex Vahl, Heteran-
thera limosa Willd., Isoetes lithophila N. Pfeiff., Juncus texanus (Engelm.) Coville in Small, Limnosciadium pin-
natum (DC.) Mathias & Constance, Lindernia dubia (L.) Pennell var. anagallidea (Michx.) Cooperr., Hemicar-
pha micrantha (Vahl) Pax, Ludwigia glandulosa Walter, Marsilea vestita Hook. & Grev., Pilularia americana A.
Braun, Rotala ramosior Koehne, Schoenoplectus saximontanus (Fernald) J.Raynal.
W103°00'45", 4200 ft, 18 Sep 1988, Worthington 17456 (UTEP). Kerr Co.: about Kerrville, 12-19 Jun 1894, Heller 1851 (GH, MO, SMU, US);
6 plants along margin of Turkey Creek by crossing of Fall Creek Road, N29“57'40.7", W099°10’28.8'', elev. 499 m, 30 Jun 1993, O’Kennon
11696 (BRIT). Lampasas Co.: 21 Sep 1892, Plank s.n. (NY). Llano Co.: Enchanted Rock State Natural Area, ca. 5 plants along sandy margin
of Moss Pond, N30‘’30'33'’, W098°49'3r', elev. 458 m, 10 Oct 1991, 0’Kennon 101 71 (BRIT). Mason Co.: ca. 5 plants along sandy banks of Hay
Branch on FM 152, 8.7 km W of Castell, N30°39'22", W099°01T3", elev. 384 m, 25 Jun 1992, 0’Kennon 10677 (BRIT). Reagan Co.: 2 mi S of
Big Lake in mud, 24 Aug 1939, Cory s.n. (GH); 2.5 mi S of Big Lake, 24 Aug 1939, Cory 32765 (TAES). Travis Co.: silt of Colorado River, Lake
Austin, 12 Aug 1922, Tharp 2128 (TEX, US); 2129 (US). Uvalde Co.: margin of pond, Winston-Mize Ranch, N29°13'16'', W099°13'36'', elev
258 m, 7 May 2005, 0’Kennon 19994 (BRIT). Val Verde Co.: Howard Creek, 4Jul 1852, Wright 1930 (GH); Wright 1932 (GH, US); Wright 1961
ACKNOWLEDGMENTS
Thanks to The Academy of Natural Sciences, Philadelphia (PH), Harvard (GH), Smithsonian (US), Missouri
Botanical Garden (MO), University of Texas (TEX, LL), Texas A&M (TAES, TAMU), and N.Y. Botanical Garden
(NY) for loans of plant material for research for this paper. Thanks also to Amanda Neill, Director of the Her-
barium of the Botanical Research Institute of Texas (BRIT), for the excellent drawings of the achenes. We thank
Galen Smith, Jackie Poole, and an anonymous reviewer for constructive reviews of our manuscript.
REFERENCES
Buckley, S.B. 1862 (1863). Descriptions of new plants from Texas. No. 2. Proc. Acad. Nat. Sci. Philadelphia 14:5-10.
Buckley, S.B. 1870. Remarks on Dr. Asa Gray's notes on Buckley's new plants of Texas. Proc. Acad. Nat. Sci. Philadelphia
CORRELL, D.S. AND M.C. JoHNSTON. 1 970. Manual of the vas
CoRRELL, D.S. AND H.B. CoRRELL. 1972. Aquatic and wetlar
tection Agency. U.S. Government Printing Office, W
Diggs, G.M. Jr., B.L Lipscomb, and RJ. O'Kennon. 1999. Shii
s Research Foundation, Renner.
United States. U.S. Environmen
ngton, D.C.
s & Mahler's illustrated flora of North Central Texas. Sida, Bot.
Dorr, LJ. and K.C. Nixon. 1985. Typification of the oak (Quercus) taxa described by S.B. Buckley (1809-1884). Taxon
34:211-228.
Godfrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United States, monocotyledons. Uni-
versity of Georgia Press, Athens.
Gonzalez-Elizondo, M.S. and P.M. Peterson. 1 997. A classification and key to the supraspecific taxa in Eleocharis (Cypera-
593
il and Mechanical College of Texas,
Gould, F.W. 1962. Texas plants, a checklist and ecological si
College Station.
Gray, A. 1852-1853. Plantae Wrightianae te
5; vol. V, art. 6 Smithsonian contributions to knowledge; v. 3, art. 5; v. 5, art. 6.
Gray, A. 1862. Notes upon the "description of new plants from Texas, by S.B. Buckley," published in the Proceedings of the
Academy of Natural Sciences of Philadephia, Dec. 1 861 and Jan 1 862. Proc. Acad. Nat. Sci. Philadelphia 1 4:1 61 -1 68.
Hatch, S.L, K.N. Gandhi, and LE. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Misc. Publ.
No. 1655.
Hooker, J.D. and B.D. Jackson. 1 895. Index Kewensis. 1 :830.
Jones S.D., J.K. Wipff, and P.M. Montgomery 1 997. Vascular plants of Texas: a comprehensive checklist including synonymy,
bibliography, and index. Univ. of Texas Press, Austin.
Kartesz, J.T. and C.A. Meacham. 1 999. Synthesis of the North American flora. Version 1 . North Carolina Botanical Garden.
Chapel Hill.
Kartesz, J.T. 1 994. A synonomized checklist of the vascular flora of the United States, Can;
Volumes 1 and 2. Timber Press, Portland, Oregon.
Shaw, E.A. 1987. Charles Wright on the boundary 1849-1852 or Plantae Wrightianae revi
Westport, Conn.
Smith, G.S., JJ. Bruhi, M.S. Gonzalez-Elizondo, and F.J. Menapace. 2003. Eleocharis. In: Flora of North America Editorial Com-
mittee, eds. 1 993-t-. Flora of North America North of Mexico. 1 2-f voIs. New York and Oxford. Vol. 23.
Svenson, H.K. 1 929. Monographic studies in the genus Eleocharis. Rhodora 31 :224-242.
Svenson, H.K. 1937. Monographic studies in the genus Eleocharis-WI. Rhodora 39:259-262.
Svenson, H.K. 1 939. Monographic studies in the genus Eleocharis-^. Rhodora 41 :43-77, 90-1 1 0.
Svenson, H.K. 1 957. Poales: Cyperaceae: Scirpeae (Continuatio) [Fuirer^a, Hemicarpha, Eleocharis]. N. Amer. FI. 1 8:520-521 ,
532-533.
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ANNOUNCEMENT
The 2013 Applications for the Delzie Demaree Travel Award
Applications for the 2013 Delzie Demaree Travel Award should include a letter from the applicant telling how
symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major pro-
fessor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology Depart-
ment, The College of William and Mary, Williamsburg, Virginia 23185-8795, U.S. A. 1-757-221-2799; Email;
ddmware@wm.edu. Applications may be sent to: Barney Lipscomb, 1700 University Drive, Fort Worth, Texas
76107-3400, U.S. A. 1-817-332-7432; Email: barney@brit.org. The period for receiving applications will end
three weeks prior to the date of the symposium if a sufficient number of applications are in hand at that time.
Anyone wishing to apply after that date should inquire whether applications are still being accepted before
applying. The Systematics Symposium dates for 2013 are 11-12 October 2013.
The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35
out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, dis-
coverer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He also
taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas A&M,
and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of the things
he enjoyed most as a botanist was assisting students with their field botany research.
J.Bot. Res. Inst. Texas 7(1): 594. 2013
VASCULAR FLORA OF SALINE LAKES IN THE
SOUTHERN HIGH PLAINS OF TEXAS AND EASTERN NEW MEXICO
David J. Rosen
Amber D. Caskey
Department of Biology
Lee College
Baytown, Texas 77522-0818, U.S.A.
Department of Biology
Lee College
Baytown, Texas 77522-0818, U.S.A.
Warren C. Conway
Arthur Temple College of Forestry and Agriculture
Stephen F. Austin State University
Nacogdoches, Texas 75962-6109, U.S.A.
David A. Haukos
U.S. Geological Survey
Kansas Cooperative Fish and Wildlife Research Unit
Kansas State University
Manhattan, Kansas 66506, U.S.A.
dhaukos@ksu.edu
In the High Plains of the westei
hydrological surface features.
management focus, primarily
functional ecology and structc
son et al. 2012). Numbering ne
alized regions in North Ameri
INTRODUCTION
rn Southern Great Plains, playa wetlands and saline lakes form the predominant
. This is particularly evident in the Southern High Plains (SHP) of northwest
CO Playas of the SHP have received a tremendous amount of conservation and
stemming from concerns regarding natural and anthropogenic threats to playa
are (Bolen et al. 1989; Haukos & Smith 1994; Luo et al. 1997; Smith 2003; John-
>arly 20,000, playas are shallow, isolated, freshwater, precipitation filled recharge
scribed as islands of biological diversity in one of the most intensively agricultur-
ica (Bolen et al. 1989; Haukos & Smith 1994; Johnson et al. 2012). Playas are dy-
596
Journal of the Botanical Research Institute of Texas 7(1)
namic and biologically productive wetlands that provide critical habitat for migrating, breeding, and wintering
waterfowl (Ray et al. 2003; Haukos et al. 2006; Moon & Haukos 2006); migrating (Davis & Smith 1998) and
breeding shorebirds (Conway et al. 2005a, b); and wintering sandhill cranes (Grus canadensis; Iverson et al.
1985). The playa flora has also been the focus of numerous studies, including checklists (Rowell 1971), field
guides (Haukos & Smith 1997), and more detailed analyses of plant community structure as related to soils,
playa area, and physical location on the landscape (Smith & Haukos 2002; Haukos & Smith 2004; O’Connell
etal. 2012).
In contrast, saline lakes of the SHP are poorly studied, and have received little or no conservation atten-
tion. Despite the conservation focus upon playas, saline lakes are arguably the most imperiled wetland ecosys-
tem in the region and have been identified as a High Priority Community within the High Plains Ecoregion
(Texas Parks and Wildlife Department 2005). Approximately 1 saline lake exists for every 500 historical playas
regionally (Reeves & Temple 1986; Haukos & Smith 1994; Smith 2003), but their numerical rarity makes
them, and their associated biota, particularly susceptible to both natural and anthropogenic perturbations.
Saline lakes differ from playas ecologically, geologically, and hydrologically (Reeves & Temple 1986; Os-
terkamp & Wood 1987; Hall 2001). Saline lakes are generally larger than playas, have saline water chemistry,
(>200 g/L of dissolved solids [Osterkamp & Wood 1987; Dockery 1989]), are often underlain by areas of deep
Permian salt dissolution (Dockery 1989), and usually are bordered by dunes on their leeward side from defla-
tion (Osterkamp & Wood 1987). Geologically older than playas (-300,000 ybp [Wood 2002]), most regional
saline lakes have evidence of human occupation 10,000-12,000 ybp (E. Johnson, unpublished data).These
settlements were likely due to the permanence of one or more freshwater springs connected to the Ogallala
Aquifer in an environment with otherwise rare surface water. Still considered discharge wetlands, saline lakes
currently receive most surface water via direct precipitation and from overland flow, as natural artesian fresh-
water springs associated with most saline lakes have ceased (or have dramatically reduced) flowing during the
last 50 years (Brune 1981). Some evidence suggests that ground water elevation in saline lakes is at least 7-8 m
lower than 25-55,000 ybp (Wood 2002). As such, surface water presence (and residence time) operates as an
ecological driver of floral and faunal (e.g., bird and insect) community attributes in saline lakes (Conway et al.
2005 a, b; Andrei et al 2008; Conway unpublished data; Saalfeld et al. 2012).
Conservation concerns surrounding saline lakes are multifaceted, but cessation of spring flow causing a
reduction in surface water residence time and an increase in salt concentrations are arguably the most pressing.
Saline lake hydrology is impacted by interactions among variable hydroperiods, high evapotranspiration rates,
unpredictable precipitation patterns, and anthropogenic disturbances (i.e., groundwater pumping; Conway et
al. 2005a, b; Saalfeld et al. 2011, 2012). For example, when annual irrigation practices of groundwater pumping
coincide with extended drought and surface water evaporation, surface water is very ephemeral. Predictions of
increasing evaporation rates in models of future climate change will further reduce hydroperiods in saline
lakes, which are considered to be at “regionally high risk” of alteration under current climate change models
(Matthews 2008).
In saline lakes, rooted vascular plants are typically restricted to close proximity to remnant artesian
spring discharge zones, leaving most of the basin floor without vegetation. Due to their inhospitable environ-
ments, saline lakes possess a fraction of the floral diversity of playas (Rowell 1971; Haukos & Smith 1997) and
provide habitat for more specialized avian fauna than playas (Iverson et al. 1985; Conway et al. 2005 a, b; An-
drei et al. 2008; Saalfeld et al. 2011, 2012). For saline lakes, much of the basic biological inventory and natural
history data are lacking. The purpose of the research reported here is to provide an annotated checklist of
vascular plants collected from accessible saline lakes in the SHP of Texas and eastern New Mexico.
The SHP is an approximately 80,000 km^ region occurring from the panhandle of Texas (south of the Cana-
dian River), into New Mexico (east of the Pecos River), and south to Midland, Texas, at a regional elevation of
approximately 1000 m (Osterkamp & Wood 1987). Level topography is broken by shallow freshwater playa
Rosen etal.. Flora (
le lakes in Texas and New Mexico
wetlands; draws; and large, deep, saline lakes. Although playas numerically dominate, approximately 40 saline
lakes also occur; and combined, these features comprise the regionally dominant wetland systems. Regional
annual precipitation averages approximately 45 cm/yr, with annual evaporation exceeding 200 cm/yr, as sum-
mer temperatures may reach 40°C. Within saline lakes, temperatures huctuate diurnally during summer,
ranging from 7-54“C (Saalfeld et al. 2012). The growing season in the region ranges from 193-287 days (NRCS
We acquired access to seven individual SHP saline lakes (Fig. 1). Three on private lands (Mound Lake,
Rich Lake, and Tahoka Lake) are considered to be the best remaining saline lakes (Saalfeld et al. 2011, 2012)
as they maintain some surface water with seasonally functional artesian springs. An additional three lakes
(Goose Lake, Paul’s Lake, and White Lake) are located on the Muleshoe National Wildlife Refuge and possess
less regular spring flow. Grulla Lake, on the Grulla National Wildlife Refuge, no longer contains flowing arte-
sian springs. Processes by which these lakes have formed have been widely debated, but recent work, via core
sampling of lake basins, indicates some basin sediments are more than 300,000 years old (Wood 2002). Most
saline lakes contain aeolian deposited lunettes on the lee side of the basin, consisting of fine lacustrine sedi-
ments, deposited when saline lakes are dry (Wood 2002). As the basin floor of saline lakes were historically
connected to the top of the Ogallala Aquifer, soils of the basin are typically hydric loamy lacustine deposits
(NRCS 2000) in form, and in some instance may be greater than 30 m deep, reflecting some aeolian deopo-
sition on basin floors during periods of extended dry conditions (Wood 2002). Soils surrounding artesian
springs, in which we focused our floristic surveys, tend to be sandier and overlay the hydric loamy lacustrine
soil mentioned above.
Collecting trips were made to these saline lakes (Table 1) in 2009 from May 25th-28th and again from
September 18th-19th. For reasons mentioned, collecting was focused near artesian spring discharge zones
within the lake basins that we were confident experienced current or historical seasonal flooding. A complete
set of voucher specimens is housed at the University of Texas at Austin Plant Resources Center Herbarium
(TEX).
RESULTS AND DISCUSSION
Thisresearchresultedinacollectionof49speciesofvascularplantsrepresentingl6familiesand40genera (Table
2).ThefourfamilieswiththemostspecieswereAsteraceae(12),Amaranthaceae(8),Cyperaceae(5),andPoaceae
(12). Non-native species (Bromus catharticus, Poa compressa, Polypogon monspeliensis, Sonchus oleraceus, Kochia
scoparia, and Tamarix ramosissima) accounted for 10% of the total species. Overall species richness within each
lake excluding Tahoka Lake, was poor, only ranging from 3-14 species (Table 1). Tahoka Lake contained 43 of
the 49 species identified during the surveys, and 21 species that were not collected from any other saline lake
(Table 2). Mound, Goose, and Grulla Lakes contained one unique species each, while Rich Lake contained three
(Table 2). The only species common to all saline lakes was the exotic invasive Tamarix ramosissima (Table 2).
Reasons for disparities in species richness among saline lakes may be related to historical anthropogenic
alterations and surrounding land use practices. Each saline lake is bordered by some combination of upland
grasslands, row crop agriculture, or former cropland planted to exotic perennial grasses under the US Depart-
ment of Agriculture, Conservation Reserve Program. Row crop agriculture impacts local hydrology via
groundwater pumping for irrigation. To date, no studies have been performed to estmate water use of these
areas surrounding each basin Nonetheless, Tahoka Lake remains the least impacted saline lake because no
anthropogenic alterations nor any petrochemical exploration has occnrred on the lake basin bottom (unlike
MonnriandRichlakes). Tahoka Lake still contains threefnnctionalanesian springs, located at the north-west,
.uthwestern boundaries of the lake basin. Although spring flow is more intermittent now than in ttie past
ay et al 2005 a b' Saalfeld et al. 2011, 2012), it maintains the plant richness reported herein. On its
astern border. Mound Lake contains several small seeps and one dominant spring surrounded by a
hoenus maritimus subsp. paludosus dominated marsh (Fig. 2). A long history of petrochemical develop-
n the north and south sides of the lake have probably impacted other springs. Rich Lake contains on y
im
598
Journal of the Botanical P
Rosen et al. Flora of saline lakes in Texas and New Mexico
one primary spring on the northwestern margin of the basin, now dominated by Tamarix ramosissima, and
extensive alteration (i.e., dikes and levees) on its southern side. Grulla Lake no longer has any functional
springs, and had the poorest richness found in this study. Two Muleshoe NWR Lakes (Goose and White) also
have very intermittent springs, where discharge occurs only during the early growing season. Conversely,
Paul’s Lake has remained flooded for an extended period of time, as removal of T. ramosissima in this basin has
apparently allowed spring flow to recover. However, all three Muleshoe saline lakes contain an elevated levee
partitioning each lake. Constructed as Work Projects Administration (WPA) projects to improve water reten-
tion in a portion of each basin, these levees have dramatically altered water flow in each lake, effectively creat-
ing two different systems within each basin; one with some spring flow, the other without. As no rooted plants
reflects the importance of maintaining springs and minimizing impacts to individual lake basin hydrology.
Saline lakes contain slightly more than 10% of the species richness of playas (approximately 350 species
[Rowell 1971; Haukos & Smith 1997; Haukos & Smith 2004]). Nineteen species collected from saline lakes
have not been reported from playas (Table 2). Dominant species of the saline lakes we surveyed are similar to
those described for temperate desert wetlands of the Intermountain Region of Western North America and
occur in well defined zones in response to topographic gradients (West & Young 2000). We observed Distichhs
spicata to form a “meadow” in seasonally saturated to very shallowly flooded soils. Whereas an emergent
marsh of Bolboschoenus maritimus subsp. paludosus or submerged “beds" of Ruppia maritima occur where sur-
face water is deeper and persists longer (Fig. 2). These species (and a few interstitial species in our checklist) are
also known to be important members of saltmarsh communities of the temperate and tropical Gulf Coastal
Plain. Two other species we recorded, Suaeda nigra and Sporobolus airoides, are listed as important members of
lowland salt-desert shrub communities of all four North American deserts (West & Young 2000). More work
is needed to better understand the floristic relatedness of saline lakes of the SHP to other halophytic plant com-
munities of North America.
This survey represents the first attempt to characterize saline lake flora of the SHP of northwest Texas and
eastern New Mexico. Since the survey in 2009, the region has been experiencing an extended drought, which
has at least temporarily altered the norainsonreof these lakes,TahokaLakemparticular.Reductionsinspnng
now even in these saline lakes that maintain some artesian How, combined with little or no precipiution, may
permanently alter thU distinctive regional Hora. These wetlands are unique systems and provide virtually the
only nesting habitat for snowy plovers (Churadrius nivosus) in the region (Conway el al. 2005 a, b; Saalfeld et al.
2011, 2012). This survey provides further evidence of the importance of freshwater spring conservation in SHP
saline lakes.
ACKNOWLEDGMENTS
Financial and logistical support was provided in part by die Arthur Temple College of Forestry and Agricul-
ture Stephen F Austin Slate University, the U.S. Fish and Wildlife Service, Division of Migratory Birds, Region
2 The Rumsey Research and Development Fund, and Texas Parks and Wildlife Department. We thank the
staff at the Muleshoe National Wildlife Refuge for logistical support, and private land^ners for ^ce^ to their
properties. Weare grateful loD.Sullinsfor assistance with preparationofFigurel.andKennethD.Hedandan
anonymous reviewer for their helpful comments.
Ano„,,A.E.,LM.Sm™,D.A.H.u«os,»dJ.G.Su«s.2008. Habitat use by migrant shoteblrdslnsallnelakesofthesoumern
Great Plains. J. Wildlife Managem. 72:246-253.
Bolen, E.G., L.M. Smeth, a.nd H.L. Schramm, Jr. 1989. PI
39:615-623.
Brune, G.M. 1981. Springs of Texas: Volum
Conway, W.C., L.M. Smith, and J.D. Ray. 2005a. E
of Texas. Waterbirds 28:1 29-1 37.
., Fort Worth, Texas.
j in wetlands of the Playa Lakes Regio
602
Journal of the Botanical Research Institute of Texas 7(1)
Conway, W.C., LM. Smith, and J.D. Ray. 2005b. Habitat use and nest site selection of shorebirds in the Playa Lakes Region.
J. Wildlife Managem. 69:1 74-1 84.
CORRELL, D.S. and M.C Johnston. 1 970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner.
Davis, C.A, and LM. Smith. 1998. Ecology and management of migrant shorebirds in the Playa Lakes Region of Texas.
Wildlife Monogr. 140:1-45.
Dockery, R.M. 1989. Origin of Shafter, Whalen and Lazy X Ranch lake basins, Andrews County, Texas. MS Thesis. Texas
Tech University, Lubbock.
Hall, S.A. 2001 . Geochronology and paleoenvironments of the glacial-age Tahoka formation, Texas and New Mexico
High Plains. New Mexico Geology 2001:71-77.
Haukos, D.A. AND LM. Smith. 1994. The importance of playa wetlands to biodiversity of the Southern High Plains. Land-
scape Urban Planning 28:83-98.
Haukos, D.A. and LM. Smith. 1997. Common flora of the playa lakes. Texas Tech University Press, Lubbock.
Haukos, D.A. and LM. Smith. 2004. Plant communities of playa wetlands in the Southern Great Plains. Spec. Publ. Mus.
Haukos, D.A., M.R. Miller, D. Orthmeyer, J.Y. Takekawa, J.P. Fleskes, M.L. Cassazza, W.M. Perry, and J.A. Moon. 2006. Spring migra-
tion of Northern pintails from Texas and New Mexico, USA. Waterbirds 29:1 27-1 36.
Iverson, G.C., P.A. Vohs, and T.C. Tacha. 1985. Habitat use by sandhill cranes wintering in western Texas. J. Wildlife Mana-
gem. 49:1 074-1 083.
Johnson, LA., D.A. Haukos, L.M. Smith, and S.T. McMurry. 2012. Loss and modification of Southern Great Plains playas. J.
Environm. Managem. 1 12:275-283.
Luo, H., L.M. Smith, B.L. Allen, and D.A. Haukos. 1997. Effects of sedimentation on playa wetland volume. Ecol. Applic.
Matthews, JJ. 2008. Anthropogenic climate change in the Playa Lakes Joint Venture region: understanding impacts,
discerning trends, and developing responses. Report to the Playa Lakes Joint Venture.
Moon, J.A. and D.A. Haukos. 2006. Survival of female Northern pintails wintering in the Playa Lakes Region of northwest-
Natural Resources Conservation Service. 2000. Soil survey, Lynn County, Texas. httpV/soildatamart.nrcs.usda.gov/manu-
scripts/TX305/0/Lynn%20County.pdf
O'Connell, J., L. Johnson, L. Smith, S. McMurry, and D. Haukos. 2012. Influence of land-use and conservation programs on
wetland plant communities of the semi-arid United States Great Plains. Biol. Conservation 146:108-1 15.
Osterkamp, W.R. and W.W. Wood. 1987. Playa-lake basins on the Southern High Plains of Texas and New Mexico: Part I.
Hydrologic, geomorphic, and geologic evidence for their development. Bull. Geol. Soc. Amer. 99:21 5-223.
Ray, J.D., B.D. Sullivan, and H.W. Miller. 2003. Breeding ducks and their habitats in the High Plains of Texas. Southw. Natu-
ralist 48:241 -248.
Reeves, C.C., Jr. and J.M. Temple. 1 986. Permian salt dissolution, alkaline lake basins, and nuclear-waste storage. Southern
High Plains, Texas and New Mexico. Geology 14:939-942.
Rowell, C.M., Jr. 1971 .Vascular plants of the playa lakes of the Texas Panhandle and South Plains. Southw. Naturalist 15:
407-417.
Saalfeld, S.T., W.C. Conway, D.A. Haukos, and W.P. Johnson. 201 1 . Nest success of snowy plovers {Charadrius nivosus) in the
Southern High Plains of Texas. Waterbirds 34:389-399.
Saalfeld, S.T., W.C. Conway, D.A. Haukos, and W.P. Johnson. 2012. Snowy plover nest site selection, spatial patterning, and
temperatures in the Southern High Plains of Texas. J. Wildlife Managem. 76:1 703-1 711.
Smith, LM. and D.A. Haukos. 2002. Floral diversity in relation to playa wetland area and watershed disturbance. Conserva-
tion Biol. 16:964-974.
Smith, LM. 2003. Playas of the Great Plains. University of Texas Press, Austin.
Texas Parks and Wildlife Department (TPWD). 2005. Texas Comprehensive Wildlife Conservation Strategy 2005-201 0. Texas
Parks and Wildlife Department, Austin.
West, N.E. and J.A. Young. 2000. Intermountain valleys and lower mountain slopes. In: M.G. Barbour and W.D. Billings, eds.
North American Terrestrial Vegetation, 2nd ed. Cambridge University Press, NY. Pp. 255-284.
Wood, W.W. 2002. Role of ground water in geomorphology, geology, and paleoclimate of the Southern High Plains,
USA. Ground Water 40:438-447.
ECOLOGY AND DISTRIBUTION OF THE NORTH CENTRAL TEXAS ENDEMIC
DALEA REVERCHONII (FABACEAE)
Kimberly Norton Taylor and Robert J. O'Kennon
Botanical Research Institute of Texas
1700 University Drive
Fort Worth, Texas 76 107-3400. U.S.A.
ktaylor@brit.org; okennon@brit.org
RESUMEN
INTRODUCTION
Dalea reverchonii (S. Watson) Shinners was described in 1886 by Sereno Watson from an 1882 collection made
by lulien Reverchon (1273, GH, NY, SMU, US) “on the rocky top of Comanche Peak” (Watson 1886), Hood
County, Texas. Eggert recollected the species from Comanche Peak in 1900 (Mahler 1984). Seve^^^^
have attempted to relocate the type locality on Comanche Peak but were unable to find any plants (Orzell
1987). In 1964 Barneby collected D. reverchonii in “limestone breaks” near Springtown in Parker County (Bar-
neby 13529, NY). This collection represented the first time the species had been seen in 64 years. Mahler relo-
cated this population in 1984 (Mahler 9594, SMU) and found several new locations in Parker and Wise counties
(Turner 1959- O’Kennon 2010). Mahler (1984) recorded eight populations of D. reverchonii m 1984 but noted
that “the actual observed populations are thought to reflect only a small percentage of the total number possi-
ble within the Grand Prairie.” Barneby (1977) agreed with this sentiment saying “the species must be expected
to turn up elsewhere around the northern fringes of Edwards Plateau. , j - ii
Several surveys have been completed documenting the range of Dalea reverchonii, including Orzel
(1987), Singhurst and Horner (1997). and McLemore and O’Kennon (2003). As of 2003 there °
wUh only 23 plants remaining in 2003 (McLemore Sr O'Kennon 2003). In 2003, McLemore and O ennon
relocated the "ype locality onL top of Comanche Peak for the Brst time in 103 years, where they found 11
plants(0'Kennou&McIemorel8»3,BRin
Dalea reverchonii ts a calctphile, known to grow y ' . j ^ ,^^,5
Mahler (1084, described
of Texas 7(1)
noting that the species is restricted to Walnut Limestone glades. Much of the Walnut (Kwa) and Goodland
(Kgl) limestones are mapped as undivided units (i.e. Kgw), possibly leading to the uncertainty in geology. Co-
manche Peak, the type locality for the species, is topped with Edwards Limestone (Ked). Dalea reverchonii has
only been collected in limestone substrate on what have been termed limestone glades by Swadek and Burgess
(2012). These glades consist of little to no soil overtopping limestone bedrock.
As of 2003, this rare Texas endemic was only known from Hood, Parker, and Wise counties in Texas.
high risk of extinction “due to very restricted range, very few populations, steep declines, or other factors”
(NatureServe 2013). The recent discovery of additional sites for several associate species suggests the possibil-
ity of finding additional D. reverchonii populations (Swadek & Burgess 2012; Taylor et al. 2012). Mahler (1984)
believed “continued field research will undoubtedly produce more populations.” The purpose of this study was
to update the status of D. reverchonii, by surveying for additional populations and providing a thorough de-
scription of D. reverchonii habitat, including underlying geology and associated species.
In 2011 the authors began an updated status survey of Dalea reverchonii. All known sites were revisited from
spring 2011 through fall 2012 and population numbers were estimated. Associated species, geology, and habi-
tat preferences were recorded. Satellite imagery and geologic maps were used to locate limestone outcrops
representing habitat similar to that seen on known sites across nine north central Texas counties. All potential
suitable habitat identified was searched for additional populations.
RESULTS AND DISCUSSION
Dalea reverchonii was found at 33 additional sites across eight north central Texas counties (Fig. 1). This brings
the total number of sites to 69 and represents new county records for Bosque, Erath, Johnson, Somervell, and
Tarrant counties. Population sizes ranged from one to several thousand plants, though most sites had fewer
than 100 plants. The Aurora sites in Wise County were the largest population, consisting of an extensive sys-
tem of linked glades. Each of the approximately 10 glade groups in the Aurora complex was estimated to have
over 1000 plants. Population numbers are largest in Wise and Parker counties, with the populations in Bosque,
Erath, Hood, Johnson, Somervell, and Tarrant counties typically consisting of fewer than 10 plants. Associated
species include Minuartia michauxii (Fenzl) Farw. var. texana (B.L. Rob. ex Britton) Mattf., Phemeranthus ca-
lycinus (Engelm.) Kiger, Sporobolus vaginiflorus (Torr. ex Gray) Wood, Hedyotis nigricans (Lara.) Fosberg, He-
liotropium tenellum Torn, Hedeoma drummondii Benth., Plantago helleri Small, Erioneuron pilosum (Buckley)
Nash, Opuntia phaeacantha Engelm. var. major Engelm., Tetraneuris scaposa (DC.) Greene, and Croton monan-
thogynus Michx. Less common associates include Pediomelum reverchonii (S. Watson) Rydb., Coryphantha sul-
cata (Engelm.) Britton & Rose, Dalea frutescens A. Gray, Bouteloua pectinata Feath., Paronychia virginica
Spreng., and Yucca pallida McKelvey. From Hood County southward, the following species were typically as-
sociated with D. reverchonii: Erigeron modestus A. Gray, Linum rupestre (A. Gray) Engelm. ex A. Gray, Thele-
sperma simplicijolium (A. Gray) A. Gray, Verbena canescens Kunth var. roemeriana L.M. Perry, Vemonia lind-
heimeri A. Gray & Engelm., and Melampodium leucanthum Torr. & A. Gray.
Dalea reverchonii appears restricted to crevices in exposed limestone bedrock, very shallow soils over
bedrock, or Walnut marl where the shell hash has been removed (Fig. 2). The roots are embedded in the Wal-
nut marl directly below the indurate Texigryphaea mucronata shell hash. The plant was rarely seen where veg-
etative cover is taller and denser, which seems to be associated with deeper soils overlying the limestone. The
closely related limestone cedar glade endemic Dalea gattingeri (A. Heller) Barneby has high light requirements
and poor competitive ability, suggesting the species is endemic to limestone glades of Tennessee, Alabama,
Georgia, Missouri, and Arkansas not because it prefers the habitat but because it is not shaded and outcom-
peted by other taxa (Breeden 1968). It is likely that D. reverchonii is restricted to limestone glades for similar
reasons. Gankin and Major (1964) concluded that rare plants often occur in areas where light competition
Taylor and O'Kennon, Ecology and distribution of the endemic Dalea reverchonii
Fig. 2. Top - Dalea reverchonii plants showing prostrate habit of older plants (left) and more erect habit of younger growth (right). Photo taken 7 May
2011, New Highland Rd. site, Parker County. Bottom - Dalea reverchonii plant showing woody taproot and seeds (inset). Seeds have a hard coat and
are 1.7 x 2.7 mm. Photo taken 11 Nov 2011, Summit Ridge glade site, Parker County.
ciated vegetation is decreased “by ;
xtraordinary soil parent material . . . (Baskin & Baskir
1988).
All Dalea reverchonii populations, with the exception of the Comanche Peak site, were found on texigry-
phaeid oysterbeds of the Walnut Limestone Formation (Fig. 3). In the northern and western parts of its range
D. reverchonii sites tend to be on higher parts of the landscape, appearing as erosional remnants capping the
softer Paluxy Sandstone below. In the southern and eastern parts of its range D. reverchonii tends to be at lower
parts of the landscape where the softer Goodland/Comanche Peak Limestone or clays in the Fredericksburg
group erode away exposing the harder Walnut caprock below. Therefore, D. reverchonii is typically found at the
edges of the Walnut Formation near the contact with either the underlying or overlying strata. Thicker soil
development above the hard Texigryphaea oyster beds, and the associated taller vegetation, rarely support the
plant. Therefore populations will only be stable in areas where the bedrock remains exposed forming an
edaphic climax community. This explains why the plant is not found further to the south on the Edwards
Plateau where the Walnut Formation is quite expansive but is typically located in valleys and supports more
developed soils. A few D. reverchonii sites were found where soil development was more advanced, and the
plants were overtopped by annual and perennial grasses. These sites have likely progressed from open glades
to deeper-soiled barrens as described by Quarterman (1950) for the cedar glades of central Tennessee. Dalea
reverchonii could potentially persist for some time in glades where it was well established that have become
overgrown, but population numbers would be expected to decline, ultimately resulting in extirpation from the
The Dalea reverchonii population from the top of Comanche Peak (Fig. 4) is the only known location for
the species not in Walnut Limestone. The type locality is actually in thin soil on Edwards Limestone near the
contact with Comanche Peak Limestone (very similar to Goodland Limestone to the North of Hood County)
and far from (60 meters above) the Walnut Limestone. The peak base is surrounded by Walnut Limestone with
an abundance of exposed bedrock similar to that at other D. reverchonii sites. This area is currently grazed by
cattle which may explain the plant’s absence from this otherwise ideal habitat. We hypothesize that the Walnut
Limestone surrounding Comanche Peak historically supported a large population of D. reverchonii. The plants
on the top of Comanche Peak were likely opportunistic and only survived due to the difficulty for cattle to
reach them. The Comanche Peak population suggests that it is not the chemical makeup of the Walnut Lime-
stone that determines the plants viability, but the edaphic conditions that the hard erosion-resistant caprock
produces. These mesas capped with Edwards Limestone, which are similar to the cuestas of the Grand Prairie
capped with Walnut Limestone, are abundant throughout the Lampasas Cut Plain south of Hood County (Hill
1901). These areas are mostly privately held ranch land and may represent an unexplored habitat for the
species.
On surfaces stabilized by a caprock glade of Walnut Limestone, the vegetation tends to be lower, more
open, and less grassy than typical mixed grass prairies. This vegetation may be part of the “Walnut Prairie-
described by Hill (1901), which he recognized as distinct from the Fort Worth Prairie. This community should
not be confused with mixed grass prairies, especially since it has at least one endemic plant
these landscape units were distinguished by some label other than “prairie.” The processes s‘"
Swadek and Burgess (2012) proposed the open rocky areas where D. reverchonii tnrives ne lermea vvai-
nut Limestone glades.” Proposing a name for the unique landscape and community that this species inhabits
would aid stewards of environmental heritage in promoting the appreciation and conservation of this distinc-
tive regional ecosystem. Swadek and Burgess (2012) define a Walnut Limestone glade as “open areas of ex-
posed limestone outcrops and rocky areas with soils around 0 to 5 cm deep. Texigryphaea fossil shell fragments
TLse Walnut Limestone glades are both structurally and floristically “reminiscent of those found in the
ssee” (Taylor et al. 2012). Dalea gattingeri fills the same ecological niche in the Tennessee
Central Basin of Tennes
LxiiL:.
ing woody encroachment into the glades will be essential in maintaining viable populations. Dalea reverchonii
appears to thrive in sites where the Walnut caprock has been removed, leaving the softer underlying marl ex-
posed. These quarried sites appear to be well suited for D. reverchonii as scraping removes competing vegeta-
tion and the hard shell hash, allowing roots easy access to the Walnut marl. It is unclear if this success will
persist as soils may develop and competing vegetation invade with the absence of the hard rock shell hash.
In 2005, Dalea reverchonii plants were transplanted to three locations in the Lyndon B. Johnson National
Grasslands in Wise County. The plants that were introduced are thriving, but no new plants have come up
from seed. This lack of recruitment is likely due to the hard seed coat (Fig. 2), but could have implications for
establishment of new viable populations. An additional population was planted in the Fort Worth Nature Cen-
ter and Refuge in Tarrant County in 2012. All reintroduced populations were planted on Walnut Limestone
glades. It is likely that D. reverchonii plants would have historically occurred at both the LBJ National Grass-
lands and Fort Worth Nature Center and Refuge. A history of livestock grazing at both locations could explain
their current absence from these locations. These transplanted populations represent the only populations on
city, state, or federally protected land. Dalea reverchonii currently has a conservation rank of G2S2, but with
this recent range expansion this rank may warrant modification.
Representative voucher specimens; USA. TEXAS. Bosque Co.: CR 2650 at Cr 2640, ca 1.5 mi W of Walnut Springs, h
fence line, 2 plants. 32.057893, -97.775105, elev. 284 m, 30 May 2011, OKennon 23497 (BRIT). Erath Co.: Rough Cre
ca 4.2 mi SE of hwy 67, 10.4 mi SW of Glen 1
OKennon 24574 (BRIT). R
of Comanche Peak, plants prostrate to the ground, 11 plants observed, collected from type locality. 3
Aug 2003, 0’Kennon & McLemore 18793 (BRIT); 7 plants observed, 4 May 2011, ?
610
Journal of the Botanical Research Institute of Texas 7(1)
and New Highland Rd, ca 3 mi SW of
I m, 6 May 2011, Norton & O’Kermon 790
li SE of hwy 67, 10.4 mi SW of Glen Rose, growing on Walnut Limestone
24 m, 1 Dec 2011, Norton & OKennon 1286 (BRIT). Tarrant C^.; FM 718 ca 1.5 mi SE of Newark on Sside
me glade, 4 plants, 32.988185, -97.462911, elev. 224 m, 25 May 2013, OTCennon 25829 (BRIT). Wise Co.:
1, Walnut Umestone glade, abundant, 33.070957, -97.495660, elev. 252 m, 8 May 2012, Norton 6- OTiamon
n United States:
ACKNOWLEDGMENTS
We thank Caren Marcom (McLemore), Tony Burgess, Becca Swadek, and Sam Kieschnick for their help in the
field and their valuable insight; Austin Sewell for his assistance in locating populations; the Lyndon B. Johnson
National Grasslands and the Fort Worth Nature Center and Refuge for allowing us to transplant plants; and all
of the landowners who allowed us to collect on their property. We also thank Jason Singhurst, Allan Nelson,
and Becca Swadek for their thoughtful reviews of the manuscript.
REFERENCES
Barneby, R.C. 1977. Daleae Imagines. Mem. New York Bot. Gard. 27:1 -892.
Baskin J.M. and C.C. Baskin. 1 988. Endemism in rock outcrop plant com
an evaluation of the roles of the edaphic, genetic and light factors. J. Biogeogr. 1 5:829-840.
Breeden, J.L 1968. Ecological tolerances in the seed and seedling sages of two species of Petalostemon (Leguminosae). Ph.D.
thesis, Vanderbilt University, Nashville, Tennessee.
Gankin, R. and J. Maxw. 1 964. Arctostaphylos myritifolia, its biology and relationship to the problem of endemism. Ecology
45:792-808.
Hill, R.T. 1901. Geography and geology of the Black and Grand Prairies, Texas. U. S. Geological Survey. Annual Report
21 .United States Geological Survey, Washington, D.C.
Mahler, W.F. 1984. Status report [on Dalea reverchoniil Report prepared for U.S. Fish & Wildlife Service, Albuquerque.
Unpublished. Botanical Research Institute of Texas, Fort Worth.
Mclemore, C. and R j. O’Kennon. 2003. Dalea reverchonii (S. Watson) Shinners Status Survey. Report prepared for The
Nature Conservancy's Texas Conservation Data Center. Unpublished data. Botanical Research Institute of Texas, Fort
Natureserve. 2013. Comprehensive Report Species— Dalea reverchonii. http://wviw.natureserve.org (accessed January
3,2013).
O'Kennon, RJ. 201 0. Dalea reverchonii (Fabaceae).The Reverchon Naturalist, Natural Resources Conservation Service 2:1 .
Orzell, S. 1 987. Reid survey of north-central Texas, 1 8 June-2 July 1 987. Report prepared for The Nature Conservancy
(TNC) of Texas Conservation Data Center (CDC), San Antonio.
Quarterman, E. 1 950. Major plant communities of Tennessee cedar glades. Ecology 31 :234-254.
Singhurst, J.R. and P. Horner. 1 997. A field survey of Dalea reverchonii in North-Central Texas of 1 -2 August 1 997. Report
prepared for the TNC Texas CDC, San Antonio.
Swadek, R.K. and T.L. Burgess. 201 2. The vascular flora of the north central Texas Walnut Formation. J. Bot. Res. Inst. Texas
6:725-752.
Taylor, K.N., R J. O'Kennon, and T.F. Rehman. 2012. Expanded distribution of /soetes butleri (Isoetaceae) in Texas. J. Bot. Res.
Inst Texas 6:753-757.
Tuwjer, B.L1959.The legumes of Texas. University of Texas Press, Austin.
Watson, S. 1 886. Contributions to American botany. Proc. Amer. Acad. Arts 21 :449.
IN MEMORIAM
REMEMBERING RUTH SCHOTT O’BRIEN
1921-2012
Roy L Lehman
Department of Life Sciences
Texas A&M University-Corpus Christi
6300 Ocean Dr., Unit 5802
Corpus Christi, Texas 78412-5802, U.S.A.
Ruth O’Brien, native plant specialist and curator of the Ruth O’Brien Herbarium located on the campus of
Texas A&M University-Corpus Christi, passed away on December 26, 2012. Ruth was bom October 13, 1921,
in Sioux City, Iowa. She received her Bachelor of Science degree in Bacteriology in 1943 from the University of
Minnesota. Her family relocated to Corpus Christi, Texas in 1949 where she became interested in the native
flora of Texas. It was this deep-rooted interest in native plants and taxonomy that led her to complete a Master’s
of Science degree in Biology from Corpus Christi State University (now TAMU-CC) in 1980. A year after gradu-
ating, she founded the university herbarium and became its curator in 1983.
She was interested in the mesquite bmsh community of plants because they were so adaptive to the area’s
many environments. Ruth had a thirst for knowledge, and said that her life mission was to pay back her debt to
society through teaching and devoting her time and expertise to “her” herbarium. Recently, the Native Plant
Society of Texas presented Ruth, Dr. Roy L. Lehman and Tammy White with the Donovan Stewart Correll
Memorial Award for scientific writing in the field of native flora of Texas for their book Plants of the Texas
Coastal Bend. She was also co-author of Plants of Webb County and wrote recovery plans for two local endangered
i. Bot Res. Inst Teas 7(1):6n -612.2013
612
s spent at the herbarium identifying
plant species for the U. S. Fish and Wildlife Service. Most of her ti
plants, answering questions - - - -
dergraduate students in plant projects.
She was a member of numerous societies and volunteered many years of service. Most notably were the
Botanical Research Institute of Texas, Native Plant Society of Texas, Corpus Christi Museum of Natural His-
tory, Corpus Christi Botanical Gardens, Daughters of the American Revolution and American Association of
University Women. Through the years, Ruth and her husband, Joe, donated funds to improve the herbarium
and, in 2008, started the Joseph A. and Ruth S. O’Brien Endowed Herbarium Internship. The couple also
served on the TAMU-CC University President’s Council for 18 years and was inducted into the Endowed
Scholarship Council in October 2009.
Ruth completed 50 years of volunteer service for the Corpus Christi Museum of Natural History. In rec-
ognition, the museum named a wing after her which houses an extensive botanical and shell collection. Dur-
ing her residence at the museum she donated her time as a docent, auxiliary officer, board member and honor-
ary curator of the Museum’s herbarium. In addition, she volunteered for over 30 years at the herbarium on the
scientist. Just late last year, I stopped by the university herbarium and there Ruth was, at the age of 91 , working
with graduate students on plant identifications. She was amazing!
I consider myself fortunate to have worked alongside Ruth for these past 22 years. She was a talented tax-
onomist that was able to explain to students the difficult details that are necessary to be successful in plant
identifications. She had an eager intellect, a kind heart, and a wonderful sense of humor. She
her students, colleagues and friends.
It is with much affection, admiration and honor that I write these few words for Ruth.
IN MEMORIAM
WILLIAM F. “BILL” MAHLER
1930-2013
The following obituary is reprinted with the family’s permission. As Dr. Mahler was especially important to this jour-
nal and its parent organization, a special feature by the editors will appear in the next issue. If you would like to con-
tribute a story, a remembrance, an experience, or a fond memory about Wm. F. Mahler, please send to bamey@bnt.
orgby 1 October 2013.
William F. “Bill” Mahler, of Iowa Park, Texas, passed away Tuesday, July 2, 2013, at his home. Bill was bom Au-
gust 30, 1930, to Herman & Magdalena (Maier) Mahler on Aldine Street in Iowa Park. He graduated from W.F.
George High School in 1947, and attended Hardin College in Wichita Falls, Texas. He served in the U.S. Army
from Sept. 1950 to Sept. 1953 with the 14th Ranger Infantry Company (Airborne) at Fort Bennmg, Georgia,
Fort Carson, Colorado; and Friedberg, Germany. In 1954, he returned to college and received his Bachelor’s
degree in 1955 in Agriculture from Midwestern State University. During this time, he met Wanda Lorene Lin-
desmith, of Addington, Oklahoma, and they married in January of 1955 in Burkbumett, Texas.
Bill earned his Master’s degree in Botany/Plant Taxonomy in 1960 from Oklahoma State University m
Stillwater, Oklahoma. For the next six years, he served as an assistant professor at Hardm-Simmons University
in Abilene, Texas, where he established the HSU Herbarium. He went on to earn his Ph.D. from the University
of Tennessee in Knoxville, Tennessee, in 1968. After graduation, he moved his family to
where he joined the faculty of Southern Methodist University in C
Herbarium. He became editor and publisher of Sida, Contributio
Botanical Miscellany. He authored and co-authored many plant taxonomy t> . , , n
being the Shinners’ Manual of the North Central Texas Flora (1984, 1988). For his work. Bill received the Dono-
van Stewart Correll Memorial Award for scientific writing on the native flora of Texas in 1991, from the Native
Plant Society of Texas. The Native Plant Society of Texas again honored him in 1995 with the Charles Leonard
Weddle Memorial Award in recognition of a lifetime of service and devotion to Texas native plants. Bill was a
member of many conservation organizations, including the Texas Organization for Endangered Species. He
also assisted the National Poison Control Center in identifying mushrooms involved in poisonings.
In his dedication to preserve the SMU Herbarium collection, one of the largest herbarium collections in
tbe world, he worked with Ed Bass [and others] of Fort Worth, Texas, to establish the Botanical Research Insti-
tute of Texas (BRIT). After an early retirement from SMU, he served as the first director of BRIT from 1987-
1992. In 1992, Bill & Lorene officially retired to his home town of Iowa Park, Texas, where he was an active
ogy and producing family albums with the computer, renewing friendships from childhood and his military
days, and being with his family.
Bill is preceded in death by his parents and his brothers, Herman, Jr. and John Edward. He is survived by
his wife, Lorene, of 58 years; his daughter, Brenda Cornitius and husband, Tim, of Dodge City, Kansas; his son,
Shawn and wife, Carol, of Iowa Park; and five grandchildren: Matthew Cornitius and wife, Victoria, of Dodge
City, Kansas; Rebecca Reed and husband, James, of Ft. Riley, Kansas, and Hayley, Jacob, and Nicholas Mahler
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