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JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS J. Bot. Res. Inst. Texas ISSN 1934-5259
VOLUME 3 NUMBER 1 15 JULY 2009
COPYRIGHT 2009
Botanical Research Institute of Texas (BRIT) 500 East 4th Street
Fort Worth, Texas 76102-4025, USA
© Hugh H. Itis
EDITOR: Barney Lipscomb
ASSISTANT EDITOR: Brooke Byerley
Botanical Research Institute of Texas
500 East 4th Street
Fort Worth, Texas 76102-4025, USA 817-332-7432; 817-332-4112 fax
Electronic mail: barney@brit.org; jbritebrit.org
HISTORY AND DEDICATION
1962— Lloyd H. Shinners (left), à member of the Southern Methodist University (SMU) faculty and a prolific researcher and writer, published the first issues of Sida, Contributions to Botany (now J. Bot. Res. Inst. Texas)
CONTRIBUTING SPANISH EDITOR
1971—William F. Mahler (right), professor of botany at SMU and director emeritus of BRIT, inherited editorship and copyright.
1993—BRIT becomes publisher/copyright holder. 2007—First issue of J. Bot. Res. Inst. Texas.
MISSION AND GOALS
The BRIT Press seeks innovation and excellence in preparation, manufacture, and distribution
of botanical research and scientific discoveries for the twenty-first century.
The BRIT Press—bringing out the best in botanical Science for plant conservation and education.
DIRECTION AND COVERAGE
The BRIT Press considers original research papers concerned with classical and modern systematic botany, sensu lato, for publication in J. Bot. Res. Inst. Texas. All submissions are peer-reviewed.
Guidelines for submissions are available
from the BRIT Press website, http://www.britpress.org.
BIBLIOGRAPHICAL
Citation abbreviation for the
Journal of the Botanical Research Institute of Texas is J. Dot. Res. Inst. Texas following the principles
of B.P.H. (informally JBRIT).
International Standard Serial No. (ISSN 1934-5259)
FREQUENCY OF PUBLICATION
J. bot. Res. Inst. Texas is published semiannually (summer/fall) as one volume
by the Botanical Research Institute of Texas.
Félix Llamas
Dpto. de Botánica, Facultad de Biologia Universidad de Léon
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COVER ILLUSTRATION
Electronically tinted botanical illustration of Liatris aestivalis originally used on BRIT's anniversary poster 2001.
Summer gayfeather flowers mid Jul-Aug(-Sep) and is endemic to Oklahoma and Texas.
Sida 19:768. 2001.
Botanical illustration by
Linny Heagy €2001.
BRIT PRESS
TABLE OF CONTENTS COMMENTARY
Learning about other species: an updated component of a liberal arts education Guy L. NESOM AND ÁLAN S. WEAKLEY
SYSTEMATICS
Orobanche riparia (Orobanchaceae), a new species from the American Midwest
L. TURNER COLLINS, ALISON E.L. COLWELL, AND GEORGE YATSKIEVYCH
Bristlecone pine dwarf mistletoe: Arceuthobium microcarpum subsp. aristatae (Viscaceae), a new subspecies of western spruce dwarf mistletoe from northern Arizona
JARED M. SCOTT AND ROBERT L. MATHIASEN
A new cliff-dwelling species of Zamia (Zamiaceae) from Belize
MICHAEL CALONJE
A new species of Zamia (Zamiaceae) from the Maya Mountains of Belize
MICHAEL CALONJE, JAN MEERMAN, PATRICK GRIFFITH, AND GEOFFREY HOESE
What is Zamia prasina (Zamiaceae: Cycadales)?
MICHAEL CALONJE AND JAN MEERMAN
Festuca aloha and F. molokaiensis (Poaceae: Loliinae), two new species from Hawai'i PILAR CATALÁN, ROBERT J. SORENG, AND PAUL M. PETERSON
A new genus, Mexotis, for five Mexican species of Hedyotideae (Rubiaceae)
EDWARD E. TERRELL AND HAROLD ROBINSON
Capsicophysalis: a new genus of Solanaceae (Physaleae) from Mexico and Central America
JOHN E. AvERETT AND MAHINDA MARTÍNEZ
Synopsis of Gonolobus s.s. (Apocynaceae: Asclepiadoideae) in Trinidad and Tobago ALEXANDER KRINGS
An overview of the Diospyros campechiana complex (Ebenaceae) and description of three new species
MITCHELL C. PROVANCE AND ÁNDREW C. SANDERS
Novedades y notas misceláneas en las Bromeliaceae de Mesoamérica J. FRANCISCO MORALES
Una nueva especie y notas misceláneas en el género Oreopanax (Araliaceae) en Centro América
J. FRANCISCO MORALES AND ALVARO IDARRAGA
Five new species of Myrtaceae from Ecuador MARIA Lúcia KAWASAKI AND BRUCE K. HOLst
Folia taxonomica 12. Paradrymonia (Gesneriaceae: Episcieae) from the Guiana Shield: P. magu- irei, a new species from Amazonas, and distribution and floral morphology of P. maculata
CHRISTIAN FEUILLET
New species of Baccharis (Asteraceae: Astereae) from Rio de Janeiro state, southeastern Brazil G. HEIDEN, J.F.A. BAUMGRATZ, AND R.L. ESTEVES
Diplycosia indica (Ericaceae): a new species and a new generic record for India M.R. DEBTA AND H.J. CHOWDHERY
Amorpha confusa, a new name for an old Amorpha (Fabaceae: Amorpheae) SHANNON C.K. STRAUB, BRUCE A. SORRIE, AND ALAN S. WEAKLEY
A new combination in North American Tephrosia (Fabaceae)
Guy L. NESOM AND JAMES L. ZARUCCHI
23
31
43
51
59
71
77
85
113
117
123
133
139
147
151
157
A recombination for varieties of Anticlea elegans (Melanthiaceae) WENDy B. ZOMLEFER AND WALTER S. JUDD
Taxonomic overview of Eurybia sect. Herrickia (Asteraceae: Astereae) Guy L. Nesom
The phylogeny of Selenia (Brassicaceae) inferred from chloroplast and nuclear sequence data JAMES B. BECK :
Preliminary insights into the phylogeny and speciation of Scalesia (Asteraceae), Galápagos Islands JEREMY D. BLASCHKE AND RoGER W. SANDERS
Introduced lesser celandine (Ranunculus ficaria, Ranunculaceae) and its putative subspecies in the United States: a morphometric analysis
ANGELA R. Post, ALEXANDER KRINGS, WADE A. WALL, AND JOSEPH C. NEAL
The types of Astragalus section Diphysi (Fabaceae), a complex endemic to Western North America, Part I: lectotypifications, epitypifications, and new combinations of several taxa
J. ANDREW ÁLEXANDER
The typification of Crotalaria rotundifolia and Crotalaria maritima (Fabaceae)
DaNiEL B. WARD
Lectotypification of Cardamine flexuosa (Brassicaceae)
ANGELA R. Post, ALEXANDER KRINGS, QUI-YUN (JENNY) XIANG, BRYON R. SOSINSKI, AND JOSEPH C. NEAL
An investigation of morphological evidence supports the resurrection of Pyrrocoma scaberula (Asteraceae: Astereae)
Curtis R. BJORK AND MARK DARRACH
Miscellaneous typifications, one new series, and one new varietal combination in Crataegus (Rosaceae) J.B. Pipers
What is Ranunculus gelidus (Ranunculaceae)? ALAN T. WHITTEMORE
Infrageneric taxonomy of Astrophytum (Cactaceae), with remarks on the status of Digitostigma RICHARD R. MONTANUCCI
Mycological literature on Texas fleshy Basidiomycota, two new combinations, and new fungal records for Texas
Davip P. Lewis AND CLARK L. OVREBO
Twenty-eight new three-letter family acronyms for vascular plants (with comprehensive listings on-line)
NEIL SNOW
CHROMOSOME NUMBERS
Chromosome numbers of miscellaneous angiosperm taxa Marc BAKER, Jon REBMAN, BRUCE PARFITT, DONALD PINKAVA, CHARLOTTE CHRISTY, ANDREW SALYWON, RAUL PUENTE-MARTINEZ, ALLAN ZIMMERMAN, AND J. Huso COTA
ANATOMY AND MORPHOLOGY
Anatomia de la epidermis foliar en las especies Mexicanas del género Muhlenbergia (Poaceae) OCTAVIO ROSALES CARRILLO AND YOLANDA HERRERA ARRIFTA
FLORISTICS, ECOLOGY, AND CONSERVATION
Primer reporte de Cystopteris reevesiana (Aspleniaceae) para México José A. VILLARREAL-Q. AND CELESTINO FLORES-L.
159
161
169
177
193
211
219
227
231
239
245
251
257
273
279
285
307
Floristic composition, abundance, and distribution pattern of woody plants in a tropical savanna in northern Ghana ALEX ASASE, PATRICK K. EKPE, AND JOHN Y. AMPONSAH
New state records of vascular plants for the Southern Rocky Mountain region PAMELA F. SMITH, DONALD L. HAZLETT, AND NEIL SNOW
Pyracantha koidzumii (Rosaceae) new to the Arkansas flora
BRETT E. SERVISS
Medicago rigidula and M. truncatula (Fabaceae): new to the California flora RICHARD E. RIEFNER, JR. AND ERNEST SMALL
New and noteworthy plants from Florida
Jonn M. KUNZER, RICHARD P. WUNDERLIN, LORAN C. ANDERSON, AND JAMES R. BURKHALTER
Vascular flora of Churchill Ranch, Sarasota County, Florida ALAN R. FRANCK AND RICHARD P. WUNDERLIN
Vascular flora of a saline prairie in Winn Parish, Louisiana BARBARA R. MACROBERTS, MICHAEL H. MACROBERTS, CHRISTOPHER S. REID, AND PATRICIA L. FAULKNER
Vascular flora of Morse Clay prairies in northwestern Louisiana BARBARA R. MACROBERTS, MICHAEL H. MACROBERTS, CHRISTOPHER S. REID, AND PATRICIA L. FAULKNER
Floristics of upland shortleaf pine/oak-hickory forest in northwestern Louisiana BARBARA R. MACROBERTS AND MICHAEL H. MACROBERTS
A revision to the vascular flora of Calcasieu Parish, Louisiana Ray NEYLAND
Additions and emendations to the vascular flora of Caddo Parish, Louisiana
BARBARA R. MacRosrnrs, MICHAEL H. MACROBERTS, CHRISTOPHER S. REID, AND PATRICIA L. FAULKNER Mentha suaveolens and M. xrotundifolia in North Carolina: a clarification of distribution and taxonomic identity
MICHAEL W. DENSLOW AND DERICK B. POINDEXTER
Castilleja angustifolia var. dubia (Scrophulariaceae), a new record for South Dakota
GRACE KOSTEL AND LYNN HETLET
Glyceria declinata (Poaceae) new to the flora of Texas
CHARLES M. ALLEN, PATRICIA LEWIS, AND Davip P. Lewis
Vegetation areas of Texas: concept and commentary
Ray C. TELFAIR II
Vascular plant species richness of the Big Thicket, East Texas
MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS
A classification of geographic elements and analysis of the flora of Big Bend region of Texas ANNA SAGHATELYAN
Annotated checklist of the vascular flora of the Menard Creek Corridor Unit of the Big Thicket National Preserve, Polk, Liberty, and Hardin counties, Texas Larry E. Brown, BARBARA R. MACROBERTS, MICHAEL H. MACROBERTS, AND WARREN W. PRUESS
Cyperus pilosus (Cyperaceae) new to the flora of Texas
RICHARD CARTER, CHARLES M. ALLEN, PATRICIA LEWIS, AND Davip P. Lewis
Cryptostegia grandiflora (Apocynaceae: Asclepiadoideae), a new non-native weed for Texas THOMAS F. PATTERSON AND Guy L. NESOM
Tournefortia hirsutissima (Boraginaceae) new to the flora of Texas
ALFRED RICHARDSON AND W. KEN KING
309
317
319
323
331
339
349
355
367
375
379
383
391
393
395
443
457
461
Book Reviews and Notices 12, 22, 30, 42, 50, 76, 84, 122, 146, 156, 168, 192, 210, 226, 244, 272, 278, 284,308, 322,330, 0239197804390, 400, 406, 442, 456, +00: 404
Announcement 468
INDEX to new names and new combinations in J. Bot. Res. Inst. Texas 3(1), 2009
Amorpha confusa (Wilbur) S.C.K. Straub, Sorrie @ Weakley, comb. et stat. nov.—154 Anticlea elegans var. glaucus (Nutt.) Zomlefer € Judd, comb. nov.—159 Arceuthobium microcarpum subsp. aristatae J.M. Scott & Mathiasen, subsp. nov.—14 Astragalus lentiginosus var. bryantii (Barneby) J.A. Alexander, comb. nov.—215 Astragalus lentiginosus var. iodanthus (S. Watson) J.A. Alexander, comb. nov.—215 Astragalus lentiginosus var. pseudiodanthus (Barneby) J.A. Alexander, comb. nov.—216 Baccharis altimontana Heiden, Baumgratz & Esteves, sp. nov.—140
Baccharis friburgensis Heiden, Baumgratz & Esteves, sp. nov.—143
Calyptranthes compactiflora M.L. Kawasaki & B.K. Holst, sp. nov.—123 Calyptranthes fusca M.L. Kawasaki & B.K. Holst, sp. nov.—125
Calyptranthes sparsiflora M.L. Kawasaki & B.K. Holst, sp. nov.—125 Capsicophysalis (Bitter) Averett & M. Martínez, stat. nov.—72
Capsicophysalis potosina (B.L. Rob. & Greenm.) Averett & M. Martínez, comb. nov.—72 Crataegus series Altaicae J.B. Phipps, ser. nov.—243
Crataegus chrysocarpa var. subrotundifolia (Sarg.) J.B. Phipps, comb. et stat. nov.—240 Diospyros camposii M.C. Provance & A.C. Sanders, sp. nov.—93
Diospyros crotalaria M.C. Provance & A.C. Sanders, sp. nov.—95
Diospyros haberi M.C. Provance & A.C. Sanders, sp. nov.—98
Diplycosia indica M.R. Debta @ H.J. Chowdhery, sp. nov. —147
Eurybia aberrans (A. Nels.) G.L. Nesom, comb. nov.—166
Eurybia kingii (D.C. Eaton) G.L. Nesom, comb. nov.—166
Eurybia kingii var. barnebyana (Welsh & Goodrich) G.L. Nesom, comb. nov.—166 Festuca aloha Catalán, Soreng & PM. Peterson, sp. nov.—51
Festuca molokaiensis Soreng, PM. Peterson & Catalán, sp. nov.—54
Hygrocybe chamaeleon (Cibula) Lewis & Ovrebo, comb. nov.—264
Hygrocybe mississippiensis (Cibula) Lewis & Ovrebo, comb. nov.—264
Mexotis Terrell & H. Rob., gen. nov.—60
Mexotis galeottii (M. Martens) Terrell & H. Rob., comb. nov.—61
Mexotis kingii (Terrell) Terrell & H. Rob., comb. nov.— 64
Mexotis latifolia (M. Martens & Galeotti) Terrell & H. Rob., comb. nov.—61
Mexotis lorencei Terrell & H. Rob., sp. nov.—67
Mexotis terrellii (Lorence) Terrell & H. Rob., comb. nov.—64
Myrcia aequatoriensis M.L. Kawasaki € B.K. Holst, sp. nov. —128
Myrcia verticillata M.L. Kawasaki € B.K. Holst, sp. nov. —130
Oreopanax paramicolus J.F Morales & A. Idárraga, sp. nov.—118
Orobanche riparia L.T. Collins, sp. nov. —7
Paradrymonia maguirei Feuillet, sp. nov.—134
Paradrymonia subg. Pagothyra (Leeuwenb.) Feuillet, comb. et stat. nov. —134 Tephrosia xintermedia (Small) G.L. Nesom & Zarucchi, comb. nov.—157 Werauhia luctuosa J.F Morales, sp. nov. —114
Zamia decumbens Calonje, Meerman, M.P Griff. & Hoese, sp. nov —31
Zamia meermanii Calonje, sp. nov.—23
COMMENTARY
LEARNING ABOUT OTHER SPECIES: AN UPDATED COMPONENT OF A LIBERAL ARTS EDUCATION
Guy L. Nesom Alan S. Mid 2925 Hartwood Drive Herbarium/NC B Fort Worth, Texas 76109, U.S.A. Univ North Carolina, Coker Hall, E 2 guynesom@sbcglobal.net Chapel Hill, North Carolina 27599-3280, U.S.A. weakley@unc.edu
Some aspects of a liberal arts education—critical thinking, respect for truth and clarity of expression—are unchanging in value. Knowledge of humans and human culture remains central. But if the goal of the cur- riculum is to prepare students for broad engagement in the world, an understanding of the Earth and all of life calls for equal time.
Humanity is, for now at least, bound to Earth. Here, unbridled growth of the human population and profligate use of natural resources threaten our own existence and put at risk the well-being and continuance of all life and the environment. An appreciation of the planet and preparation toward committed stewardship should be among the primary goals of ae E arts education can not afford to be self-centered, in the sense of study of human cult l context and without attention to the thousands
of species that are our evolutionary associates. We propose a course that guides students towards an appreciation of the Earth as home to many spe- cies. At the end of the essay, the course is integrated within an updated liberal arts curriculum.
Learn to know other species. Perhaps the most effective path toward an appreciation of life on Earth is through first-hand knowledge of species other than our own. The primary goal of the proposed course is to learn the identities of commonly encountered species. The course need not be heavily laden with deep academic underpinning—the value lies in learning to recognize species by sight, as they are encountered in everyday life, and to identify them by name.
A know-the-species course provides foundation for one of the most basic lessons in humility—humans are one species among hundreds of thousands. So many of us are hardly aware that others exist, except in an abstract way. Daily acquaintance with other species on a "first-name" basis greatly broadens one's view of the natural world and engenders a closeness, a sense of unity, with it. Even if names of species eventually are forgotten, at least the memory remains that each is unique, distinguished by features that most have
previously been unaware of.
One can quickly learn to identify a half-dozen species (hollies, oaks, maples, etc.) and to see that some of the species are more similar among themselves while others are more unique. Such an appreciation of variation and diversity, even in a simple system, is the quickest way possible toward real insight into the process of evolution— patterns of similarity among species result from common ancestry. Each species on Earth is the product of long evolutionary lineage, interrelated among the others, each unique and beauti- ful.
“Lam strongly induced to beli hat, as in music, tl ho understands every note will, if he also possesses a proper taste, more thoroughly enjoy the whole, so ene que examines a part of a aoe view, n c puo iia the full d be a botanist, for in all I "Charles
Darwin, 1836, Voyage i the E.
Nature of the course. Plants are the most conspicuous and abundant expression of non-human life, even in urban settings, and we believe they are the best focus for the proposed course. It's joyful to walk along a
J. Bot. Res. Inst. Texas 3(1): 1 — 2. 2009
sidewalk, through a park, or through the woods and recognize species of oaks, elms, and maples by name. Even in a cityscape, dozens of species are commonly planted and usually many more are scrabbling out their existence, unassisted, seeding themselves in unlikely places.
Awareness of other species also could be gained from an introduction to birds, beetles, fish, or fungi, or practically any other kind of organism, but none of these is as easily accessible as plants. Do other courses in biology (e.g., physiology, genetics, molecular biology, cell biology) offer the same potential insights? Simply, no—most are based on study of a single species or of no species in particular. Some courses in ecology re- quire a basic knowledge of species diversity, but the focus is different. Studies of geology, meteorology, and astronomy may be doorways to recognition of the diversity of everyday extra-human experience and could be analogous to identification of plants, but diversity, or at least every-day accessibility, is much lower.
Residents of the United States and other highly urbanized areas are so removed from other species that it is easy to be unaware that plants underpin our existence. Do undergraduates recognize that our diet is mostly of roots, stems, leaves, fruits, and seeds? Do typical students know that most of the energy expended by human individuals comes directly from starch in corn, rice, cassava, wheat, and potatoes? Beans, ba- nanas, and sorghum are the world's dietary staples. Chocolate, coffee, sugar—the plant foods that tempt us. Cotton, flax, hemp, mulberry—the plants of fashion. Even the meat in our diet is but one step away from plant nutrition. Plants are responsible for nearly all our oxygen, our clothes (directly or indirectly), and most of our non-food energy (fossil fuels —oil, natural gas, coal). These topics deserve a place in a liberal arts
education.
Since the suggested course is not an abstruse undertaking, basic knowledge of other species should be taught not only as part of a liberal arts collegiate education but in elementary through high school as well. Of course, this is one among many basic topics, but lack of knowledgeable teachers is a constraint and the responsibility usually comes to rest at college level.
Suggested course outline. (A) identification of 100 species, concentrating on first-hand recognition of those most easily at hand and abundant; (B) a brief overview of plant vegetative features and reproduction (flower and fruit structure usually are critical in identification) and common families encountered among the species; (C) perspectives on evolution, the interrelatedness of life, and global patterns of diversity; and (D) a brief introduction to ecological and economic values of plants tered in everyday life (e.g., urban forests, invasive species, food, clothes, construction, fuel). Simply learning to know the species is the first
priority.
Goals of a liberal arts education. In the broad view, we believe that a liberal arts curriculum should be aimed (I) toward continued existence of our own species and all others and (2) toward maximizing the potential for freedom and happiness (including creativity, appreciation of beauty, productivity, service, and increasing understanding of the world).
Overview of an updated liberal arts curriculum. We divide a suggested curriculum into four major, overlap-
ping categories.
1. The Earth and all things non-human. Life and life processes (humans as animals, other species, ecology, evolution), geology, weather and climatology. Our proposed course would figure as a central focus in this category.
2. Intellectual outlook. Reasoning and critical thinking (including scientifi hod), respect for truth, clarity of expression, philosophy and ethics.
3. Humans and human culture. Beauty (visual art, music, perception), cultural diversity and history (history, language, philosophy, ethics, and religion).
4. The Universe and universal principles. Astronomy and cosmology, physics and chemistry, mathematics.
OROBANCHE RIPARIA (OROBANCHACEAB), A NEW SPECIES FROM THE AMERICAN MIDWEST
L. Turner Collins Alison E.L. Colwell Science and Technology Department Resources Management and Science vangel University Yosemite National Par
111 N Glenstone PO. Box 700, 5083 Foresta Road Springfield, Missouri 65802, U.S.A. El Portal, California 95318, U.S.A. collinst@evangel.edu alison_colwell@nps.gov George Yatskievych
Missouri Botanical Garden PO. Box
St. Louis, Missouri 63166, U.S.A. george.yatskievychemobot.org
ABSTRACT Orobanche dedi mE oe in the n Plains, is tl idely distributed species i t. Nothaphyllon. It was first described by T in 181 ted at Ft. Mandan, North Dakota. During the foll and a half, most botanists had HQ field Ee "ih living plants of the g ] very few herbarium peci ith e to a ke newly collected eae! bl fusion al he ci ion of thi i lits g N hyllon ( M hiza) by B (1890, 1930) aud 1930) 1 led | i l species
name too broadly to par whose mn was s discordant with Nuttall's original protologue, a the dinde ofa an Bn. broad led hi O. ludoviciana
DO
pn (Collins 1973) 1d his problem, including f l gnition of logical itia O. ludoviciana sensu stricto with different h inhabi land d tl I fined to I land iated with major river systems. Recent additional investigations have revealed fu further evid pporting gnition of tl listi ic entiti : oe logical distincti i 1 : p i f the upland f O. ludoviciana ] gnition the | land f distinct species, O. riparia, sp. nov. RESUMEN
Orobanche ludoviciana, cuya extensión abarca mayormente la zona de Great Plains, es la especie de mayor distribución en la Sección Nothaphyllon. El o que n una ee fue pas Nuttall en 1818, que se K en una muestra NAME en Ft. Mandan, Dakota del N D te el g Į i p g y con ejemplares de a con las due comparar los e obtenidos igui hubo! fusión en torno a la circun- scripción de esta especie y su distribución geográfica. Los estudios ROM con la Sección Nothaphyllon (sec. Myzorrhiza) hechos por i ie pid y Munz Coa hacen que los botánicos apliquen de modo demasiado amplio los nombres de las plantas cuya uttall. El problema que permanece es el de un concepto demasiado amplio de
E 8 la E Estudios doctoral ducid por el autor pip durante los primeros años de la década de los setenta resultaron en TRE d 1 I O. ludoviciana (Collins 1973), incluso un reconocimiento formal de dos tro de O. ludovici j distribución algo dif éspedes. Una de las plantas era s tierras
altas y es s retenida aquí como O. ludoviciana y l finada a ti baj iadas principal i Jecerincion: Reci ] lido a 1a] asd dich t a. En r
trabajo se comentan las diferencias morea entre estos dos ecotipos y se propone reconocer les adams de las tierras bajas
ta O. riparia, sp. nov.
COLL p
Key Wonps: Orobanche, Orobanchaceae, parasitic plants, broomrape, Flora of North America
Orobanche L. sect. Nothaphyllon (A. Gray) Heckard (sect. Myzorrhiza Phil.) comprises of an array of ca. 13 species that account for most of the taxonomic diversity within New World holoparasitic Orobanchaceae. It
has been considered a monophyletic group by most botanists based on morphological and molecular data (Olmstead et al. 2001; Schneeweiss et al 2004; Bennett & Matthews 2006). Within sect. Nothaphyllon, two
J. Bot. Res. Inst. Texas 3(1): 3 — 11. 2009
major clades exist whose relationships have not yet been fully resolved: the O. californica Cham. & Schltdl. complex (Heckard 1973), which has diversified mainly in the far-western United States; and the O. ludoviciana Nutt. complex (Collins 1973), which is widespread in the remainder of the United States and Canada, and southward into Mexico (also disjunct in southern South America).
Orobanche ludoviciana Nutt., with a range primarily in the Great Plains, is the most widely distributed species in sect. Nothaphyllon. It was first acude. by Thomas Nuttall in 1818 from a specimen collected at Ft. Mandan, North Dakota. B ists during the following century had limited field experience with living plants of the genus and had very few herbarium specimens with which to compare newly collected materials, the species name was applied too broadly to include plants whose morphology is discordant with Nuttall's original protologue. The result has been considerable confusion about the circumscription of this species and its geographic distribution. All those who have treated Section Nothaphyllon (sect. Myzorrhiza) (Beck 1890, 1930; Munz 1930; Collins 1973; Heckard 1973), have at times recognized various infraspecific taxa, some of which are no longer accepted and therefore excluded from this discussion. Nevertheless, the lingering problem has been that of an overly broad species concept. Even Philip Munz (1930), who provided good insights into taxonomic variation within O. ludoviciana in his otherwise excellent taxonomic revision of sect. Myzorrhiza, overlooked significant morphological variation. Subsequent authors of floristic works who relied on his revision to determine specimens continued to some degree to overlook the taxonomic heterogeneity of plants treated by him as O. ludoviciana var. genuina Beck.
Doctoral studies by the senior author led him to suggest a number of ludoviciana complex (Collins 1973), one of which was formal recognition of two ecological races within O. ludoviciana sensu stricto with differing host preferences: one inhabiting only uplands and the other confined to bottomlands associated with major river systems. Recent discoveries of new populations and examina- tion of additional existing herbarium specimens of the bottomland plants have added evidence in support of this proposal. The present paper therefore discusses the morphological distinctions between these two ecotypes, and we propose to retain the upland form as O. ludoviciana and to recognize the bottomland form as a distinct species, O. riparia. It should be noted that, based on Collins's dissertation, the epithet O. riparia was discussed far in advance of the present publication by Musselman (1982, on Orobanchaceae in Virginia) and by Kartesz (1994, in a North American plant checklist), but neither of these usages affects the validity of the present description.
Bringing this new taxon to the botanical literature requires a brief review of the existing nomenclature and morphology of O. ludoviciana sensu stricto (O. ludoviciana var. genuina, sensu Beck). Several species of Orobanche in North America are commonly misidentified as O. ludoviciana, but are currently recognized as distinct species. They include O. multiflora Nutt., O. cooperi (A. Gray) A. Heller, O. multicaulis Brandegee, and O. valida Jepson. Of these, the two with the most morphological similarity and name association are O. ludoviciana and O. cooperi, and must therefore be contrasted with O. riparia.
Collins (1973) found that Munz's 1930 description and illustration of the corolla lobes of O. ludoviciana did not match the morphology of Nuttall's type specimen. The corolla lobes of the type specimen are rounded on both the upper and lower lips, but Munz's illustration, based on a specimen from White County, Illinois, shows a corolla with triangular-pointed lobes. Because Munz used material that matched his illustration in assembling his description of O. ludoviciana, subsequent regional and state floras continued to overlook the mu gical eub pee Mp i to i euer
ated
1
within the O.
range has revealed that the character of founded e ibe is consistent throughout for UN PR (Fig. 1). In contrast, the corolla lobes of bottomland plants, hereafter referred to as O. riparia, are consistently triangular and pointed (Fig. 1). The populations of O. riparia share several additional characteristics (Table 1): D They are found almost exclusively in river bank and sandbar habitats in floodplains; 2) The native hosts are annual members of Ast tribe Heliantheae, subtribe Ambrosiinae, primarily Ambrosia trifida L. and Xanthium strumarium L., and occasionally Ambrosia artemisiifolia L. and Dicoria canescens A. Gray 3)
Collins et al., Orobanche riparia, a new species
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Fic. 1. Comp ludoviciana.
6 Journal of tt IR h Institute of Texas 3(1) Tape 1. C | id f Orot | ludoviciana, and cooperi Character Orobanche riparia Orobanche ludoviciana Orobanche cooperi Calyx len 7-11 mm 8-14 mm 7-12 mm Corolla length 15-22 mm 14-20 mm 15-32 mm ape and size Triangular-acute, 4-5mm, Rounded-obtuse, 4-8mm, Triangular-acute, 5-8 mm, of corolla erect or slightly reflexed erect or ascending apiculate apex, ascending lips/lobes or reflexed Corolla tube Strongly curved, tube Slightly curved, tube ascending, Strongly curved, tube
Corolla color
horizontal slightly flared distally Pallid, or upper lobe apex purple/lavender
slightly flared distally Purple, lavender, rose, or pallid, tube exterior whitish
ae nalig: (alee ay Dar | half with T eeii and throat
Pubescence pubescent with glandularand Pubescent with "uad Pubescent with glandular eglandular trichomes or trichomes except in trichomes on tube, eglandular glabrate lobe surface on inner surface of lobes
Flowering August-October April-August December-April
Inflorescence Open raceme, flowers in Compact raceme, flowers in Open or compact raceme with
arrangemen loose, regular spiral on dense, often irregular spiral on flowers in loose, regular spiral
Primary hosts
Habitat and
floral axis
Annuals; Ambrosia, Xanthium, icoria
River banks, sandbars eastern
floral axis
Perennials; Grindelia, Artemisia, Heterotheca
Prairie, desert, dunes; central and western U.S.
on floral axis
Paranniale Amhrocin +
Sonoran Desert, ravines, dry stream beds, dunes;
main range U.S. and prairie provinces
southwestern U.S,
The flowering period is from mid-August to early October; 4) Inflorescence color is typically pallid with only the interior tips of the corolla lobes tinted purple; 5) The corolla tube is strongly curved with the distal portion of the tube horizontal, slightly flared distally, and palatal folds that are either glabrous or densely pubescent.
Compared with O. riparia, O. ludoviciana has slightly shorter corollas with a flared throat and rounded corolla lobes (Table 1). It occurs mainly through the Great Plains (and eastward along the Prairie Peninsula) in various habitats, including sandy prairies, sand dunes, dry washes, calcareous (caliche) soils, and lower slopes of the Rocky Mountains. Its hosts are almost exclusively perennial members of Asteraceae tribes As- tereae and Anthemideae, including Grindelia squarrosa (Pursh) Dunal, Heterotheca spp., Artemisia spp., and occasionally other genera. The principal flowering period is from late April in the southern plains through mid-August in the northern plains, although sporadic flowering has been observed rarely at other times in marginal habitats. The corolla tube is slightly curved at the constriction, ascending, and flared distally, with the palatal folds sparsely pubescent. Floral pigmentation varies from purple to pale rose and occasionally yellow, usually with dark purple upper lobes (Table 1).
Additionally, populations of O. riparia are ecologically and somewhat geographically distinct, and the habitat distinction is maintained in the western populations, where the two ecological races are geographi- cally somewhat sympatric. Populations east of the Mississippi River (but including ell are allopatric with O. ludoviciana. West of the Mississippi River, sympatry of m eastern Nebraska and Kansas to eastern Colorado, southward into New Mexico and the Texas Panhandle. A small overlap in flowering times in early August could provide an opportunity for hybridization between the two species. Although no suspected cases dix uen have been addas based on morphology or field observa- tic st ded to answer this question. Molecular
+} + ors
tions of populations, data from g
studies by Colwell are currently underway. Munz (1930) treated O. cooperi as a variety of O. ludoviciana, but subsequent authors mostly have ac-
cepted it as a separate, but closely related species (Shreve & Wiggins 1964; Munz 1974). Orobanche cooperi is
Collins et al., Orobanche riparia, a new species 7
distinguished by its dark purple color, larger, di lla tube, and erect or strongly reflexed, triangular lobes with apiculate apices. The corolla tube has a whie exterior and is slightly curved and flared distally, with palatal folds pubescent at the mouth and becoming glabrous in the throat. The habitat is primarily Sonoran Desert scrub, often associated with washes that remain dry except in monsoon season. Its hosts are shrubs in various tribes of Asteraceae, principally species of Ambrosia and Encelia (all Heliantheae, Am- brosiinae), but also reported on Gutierrezia (Astereae). The flowering period is mainly December through April (Table 1). Collins (1973) found that the cross-sectional stem anatomy of O. cooperi and O. riparia are very similar and that O. ludoviciana differs significantly from both of the other taxa.
Several characters suggest that O. riparia is more closely related to O. cooperi than to O. ludoviciana. Orobanche riparia and O. cooperi share an affinity for riparian or water-eroded habitats and hosts in the sub- tribe Ambrosiinae. They differ in that O. cooperi occurs only in seasonally arid habitats, whereas O. riparia occurs in both mesic and semi-arid habitats, and that the former parasitizes shrubby hosts, whereas the host taxa of the latter are annuals. The pubescence of the corolla lobes in both species is eglandular on the inner surface with vestiture considerably longer than the short glandular trichomes typical of the exterior surface. Likewise, the palatal folds are invested with eglandular trichomes, differing between the species only in their distribution on the folds. This shared pattern of trichomes is more evident in the western populations of O. riparia than in the eastern populations. Similarly, the eastern populations have reduced indumentum as compared with the western populations. One can speculate that O. riparia and O. cooperi share a recent common ancestor. A possible scenario is that a progenitor’s range became bisected, with populations that became O. riparia adapting and dispersing into new habitats eastward across the American Midwest and populations now referable to O. cooperi attaining a distribution in the southwestern U.S. and adjacent Mexico. If this is the case, O. riparia adapted quickly to annual hosts and a late summer to early fall flowering period, in contrast with the shrubby hosts and late winter to early spring flowering period of O. cooperi. Of this character set, O. ludoviciana shares only the semi-arid habitat and perennial hosts with O. cooperi, although the host taxa of O. ludoviciana often are herbaceous.
Orobanche riparia L.T. Collins, sp. nov. (Fig. 1). ter: UNITED STATES: Inpiana: Gibson Co.: common on bank of slough .5 mi N of Griffin, 16 Aug 1931, Deam 50941 (noLorvee: IND; isorvees: A, E GH, IND, MINN, WIS).
Ab O. ludoviciana differt lobis corollae apice acutis, puberulis vel cs eure horizontali, tricto flexoq per ovarium,
inflorescentia laxe racemosa. Ab O. cooperi differt corolla 15-22 mm longa, lo! I piculati flexis, inflorescentia
pallide purpure
Annual root parasite, lacking chlorophyll. Coralloid roots few, rudimentary, stout, fleshy, usually forming a single host attachment. Stem stout or slender, fleshy, 5—35 cm long, mostly subterranean, solitary or clus- tered, Aoi or more ee ae from near base with a few to rarely numerous branches,
each beari larly enlarged or not, 5-20 mm in diameter; glabrous,
creamy siue: yellow, or tan, drying Piraeus: dark bone. or black. Leaves reduced to scales, 7-10 mm long, cauline, spirally alternate, appressed, broadly ovate to ovate triangular, becoming narrower distally. Inflorescence an open spike-like raceme, glandular pubescent; bracts mostly ca. 1 cm long, the distal ones shortened, narrowly oblong-lanceolate, acute at the tip, pale lavender to dark purple. Flowers sessile or the proximal ones on short pedicels, evenly and spirally inserted on the axis, subtended by 2 small, linear-subulate bracteoles; calyces 7-11-13) mm long, purple, deeply 5-lobed, the lobes lance-linear to linear-subulate, about equal in length, densely glandular pubescent; corollas (13—)15-22 mm long, constricted above the ovary, strongly curved at the constriction, the tube nearly horizontal, exterior surface glandular puberulent or pubescent, persistent, the tube narrow or slightly flared distally, cream-white sometimes tinted with purple, the throat with yellow palatal folds, glabrous or pubescent on folds and in grooves with eglandular trichomes; upper lip ca. 5 mm long, erect, divided about half its length, the 2 lobes triangular-acute, lavender or purple, puberulent or densely pubescent with eglandular trichomes; lower lip 3-4 mm long, divided to its base into 3 linear lobes with an acute apex, ca. 1 mm wide, pubescent with eglandular trichomes, pale, often with 1-3 purple veins. Stamens 4, inserted on the corolla tube at its constriction, generally glabrous
or with a few scattered hairs, anthers ca. 1 mm long, white, glabrous or sparsely pubescent, equalling or shorter than style. Ovary somewhat asymmetrically ovoid, 2-carpellate, nectary not evident; placentation parietal, with 4 lateral placentae; style filiform, equaling the corolla tube or shorter, stigma peltate, discoid- crateriform or rarely somewhat 2-parted Fruits 2-valved capsules, 0.7-1 cm long; seeds numerous, 0.3-0.5 mm long, ovoid to broadly ellipsoid, reticulate-pitted, light tan to dark brown. 2n = 48 (Collins 1973). Flowering mid-August to early October.
The species ranges (Fig. 2) from the Potomac and Shenandoah River valleys of Virginia to southern Ohio and Indiana along the Ohio River to Central Illinois along the Wabash and Sangamon Rivers, to the Meramec River in eastern Missouri (near St. Louis), southward to a tributary of the Mississippi River in western Ten- nessee, and westward along the Platte River in Nebraska and Eastern Colorado. Some apparently isolated populations occur in the Kansas, Oklahoma, Texas, and New Mexico along the Canadian, Cimarron, and Rio Grande rivers and their tributaries. Only historical collections are currently known from the District of Columbia, Virginia, and West Virginia. The species is found entirely on mixed sand and silt deposits on stream banks of eastern rivers or on sandbars or sandy banks of western rivers. Hosts are Ambrosia trifida and Xanthium strumarium, and occasionally Ambrosia artemisiifolia and Dicoria canescens. Rarely reported alternate hosts are Nicotiana tabacum L. and N. glauca Graham.
Representative specimens: UNITED STATES. COLORADO: Crowley Co.: cultiv: m ae near nee 22 Je 1948, ld: s.n. (CS). Jackson Co.: N sandhills E of Cow uA 1 Oct Mine Wi pd 13922 is ).I ] near Ster- == anonymous, 8 Sep 1965 (CS). P ,6mi E of Two Buttes Peak, 24 Aug 1964, Martin s.n. (COLO, CS). Weld Co.: sandhills 3 mi NE of Roggen, 1 Sep 197, pud s.n. He CS). DISTRICT OF COLUMBIA: E end of New Long Bridge, 2 Sep 1905, Ward s.n. (US). ILLINOIS: nim: ~ s of Embarras River, 2 mi E of Greenup, 18 Sep 1980, Collins 1618 (MO). Mason Co.: bank of Salt Creek and edg field, 5 mi N per cati - nis in os 1615 (MO). Menard Co.: bottoms, Athens, 1860, 1861, 1866, 1867, E. Hall s.n. (F, GH, IL, MO, POM, WIS). V mel along banks of Wabash River and Greathouse Creek, 18 Sep 1980, Collins 1619 (MO). INDIANA: Harrison Co.: W of ig at ER 25 Aug 1945, Deam 63566 (IND). Jefferson Co.: tob field 0.5 mi E of Brooksburg, 15 Sep 1935, Banta s.n. (IND). Knox Co.: along ditch 2 mi N x ud 9 Oct 1938, Deam 59255 Ends vieni Co.: E bank of Wabash River at New Harmony, 28 AE 1971, Collins 1555 (MO, WIS). k, 6.5 mi SW of Carlisle, 1 Sep 1939, Deam 59424 (IND). Vandenburgh Co: river slough, 17 Oct 1941 Zeiner s.n. (IND), Warrick Co.: banks of Ohio River at Newburg, 28 Aug 1971, Collins 1543 (UWM). KANSAS: Allen Co.: Iola, Sep 1925, anonymous (KSC). Greenwood Co.: potato patch near Eureka, 24 Sep 1935, Farmer s.n. (KS). Morton Co.: Cimarron River bed N of Wilburton, Aug 1929, Gates 16205 (F, KSC, MO). KENTUCKY: Pendleton Co.: Ohio River bank at Ivor, 27 Nov 1942, Chicoine 4745 (MO, US). MISSOURI: St. Louis Co.: a pes a bar on S bank of Meramec River, 2 mi N of Crescent, 1 Sep 1991, Yatskievych & Colwell 91-195 (MO). NEBRASKA: B Platte River, 2 mi S of Odessa, 8 Oct 1999, Collins 1620 (MO, s Co.: sandbar in Platte River, 2 mi NE of Louisville, 15 Sep D. Morrison 1342 (NEB). Dawson Co.: sandbar in Platte River, S of Lexington, 8 Oct 1999, Collins 1621 (MO). Kearney Co.: sandbars of Platte River, Minden, 15 Sep 1922, Hapeman s.n. (PH, POM/RSA, WTU). Lincoln Co.: sandbar in Platte River, Sutherland, 13 in 1936, anonymous (NEB). Merrick Co.: sandbar in Platte River, near Central City, 22 Aug 1938, I. Mueller s.n. (NEB). NEW MEXICO: Doña Ana Co.: sand, od e 25 s 1906, Wooten & Stanley s.n. (US). Sandoval Co.: sandy soil, Arroyo Chico, 22 Sep 1973, ee 1758 (ARIZ). San River, 30 m S of Farmington, pipeline ee 7 on BIA Road 2, 4 sep 2007, Collins & Heil 2032 (MO). Socorro Co.: sand along Rio Cani San Marcial, 1 Aug 1897, H k 82 ). Union Co.: idle field, 17.5 mi SW of Clayton, 1 Sep 1936, Cogdell 125 (GH). OHIO: 5 mi E of Chilo, 20 Oct 1949, Bardy s.n. (OS, US). Hamilton Co.: moni a ae Miami River, 25 pace wee Haller s.n. (US). Scioto Co.: Nile Bae a Ohio River, 29 Aug 1949, zn s.n. An f Canadian River, 10 mi NW of N , 8 Sep 1946, Goodman 4078 (MO). T dale C hofLower Forked Deer Creek, ca.2 m NE of in 16 Oct 1972, Piehl s.n. (TENN). pois Brewster Co.: »: Boquillas Given. Chisos Mts., sand near Rio Grande River bank, 5 Aug 1937, Warnock 838 (ARIZ, TEX, US). D dy soil, Rita Blanca Creek N of Dalhart, Aug 1949, York & Rogers 233 (OSU, TTC). El Paso Co.: vicinity of El Paso, 1911, Stearns 40 (MO, US). Hartley Co.: deep sand on creek bank, Punta del Agua Creek 30 mi SW of Dalhart, 26 Sep 1965, Collins 821 (OSU, TTC). Wheeler Co.: 5 mi 5 & 7 mi E of Miami, Jul 1967, Baggerman s.n. (TTC, WIS, UWM). VIRGINIA: Campbell Co.: 8 mi from Lynchburg, 13 Oct 1933, Freer s.n. (US). Page Co.: banks of Shenandoah River, Luray, 22 Aug 1912, Holtzman s.n. (US). Warren Co.: sandy field, Front Royal, 18 Sep 1897, Miller s.n. (US). WEST RGINIA: Summers Co: New River, above Shanklin's Ferry, 23 Aug 1946, McNeill s.n. (WVA)
Eastern and western populations have an apparent disjunction in distribution between eastern Missouri and east central Nebraska, a distance of ca. 650 km. A full list of specimens examined is available upon request to the authors. It is uncertain whether this is a true disjunction or simply the lack of intensive field work in this region to locate obscure populations (many botanists avoid botanizing in the midst of dense
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g allergy season). The eastern and western populations of O. riparia
stands of ragweed and cockleburs d
exhibit slightly different morphologies. Plants of the eastern populations (including the Missouri plants) are generally pallid with very pale lavender or purple pigmentation in the corollas. Pubescence is reduced and sometimes almost absent, especially on the corolla lobes. The primary host is Ambrosia trifida. The habitat of these populations is floodplains with mixed sand and heavy silty loam soils of the midwestern
rivers cited above.
The western populations (west of eastern Missouri) have more deeply purple-tinted corolla lobes, calyx lobes, and floral bracts. The upper stems, where exposed to sunlight, may be infused with a rosy purple. Pubescence is rather dense on the inflorescence, corolla lobes, and palatal folds. The primary host is Xanthium strumarium. Populations of this variant are concentrated along sand bars of the Platte River in Nebraska, with a few records of populations widely scattered in Kansas, Oklahoma, Texas, Colorado, and New Mexico. Specimens of this species collected along the Rio Grande River from central New Mexico to
the Big Bend of Texas have proven problematic for identification throughout the study of Section Nothaphyl- lon. Only recently, when they were compared with the Nebraska plants, did it become apparent that they
represent populations of O. riparia. It is tempting to name these two variants formally as subspecies within O. riparia. However, the life
history and morphological distinctions are not sufficient to warrant taxonomic status at this time. This problem requires more intensive study of both morphological and genetic variation within and between populations before a conclusion can be reached as to the merits of segregating infrataxa within O. riparia. Host information for Orobanche is generally difficult to assess from herbarium labels. Collectors sometimes list as a host any adjacent plant without confirming a physical connection or list no host at all. However, host data for O. riparia reported here have been observed by all three authors. Only two non-Asteraceous
MO Joumalof tani Insti Texas 3(
host reports have been confirmed. Some populations in southern Ohio occasionally parasitize cultivated tobacco (Nicotiana tabacum). Likewise, some populations along the Rio Grande in western Texas have been reported on the weedy, introduced N. glauca (the only perennial host with which the species has been associ- ated). It is not possible at this time to judge He a of ak exceptions, if any. Anecdotal evidence suggests that other species of Orobanche lternate hosts at the margins of their ranges.
The populations east of the Appalachian Mountains are of particular interest because they represent the extreme eastern portion of the range and may possibly be extirpated. Virginia records are represented by four herbarium specimens from four counties (on the James, Shenandoah and Potomac rivers). The most recent collection we have seen was made in 1933. A single specimen was seen from the New River, Summers County, West Virginia. The present study could not confirm extant populations in the District of Columbia, Virginia, or West Virginia. The species seems reasonably secure elsewhere in its range, although current herbarium vouchers document a highly fragmented range. For example, the single Tennessee record rep- resents a southward disjunction of ca. 325 km from the St. Louis County, Missouri station. However, field experience of the authors suggests that this species is probably more common than the herbarium records indicate. The senior author discovered several previously unreported populations in Nebraska, Illinois, and Indiana. Additionally, a few populations were observed in Illinois, Ohio, and Texas that were located on minor tributaries (creeks) of the major river systems. Some of the historical populations were determined to be extant. Thus, it is likely that concentrated field work will reveal additional populations of O. riparia. The status of the historical populations in the District of Columbia, Virginia, and West Virginia especially needs to be updated by further field work.
The habitat of O. riparia is disturbed annually by levents and h tivity. Although it is widely distributed, populations seem to be locally isolated and therefore fragile. The senior author has observed the obliteration of a large population in Posey County, Indiana, as a result of riverbank development. Because of its unique distribution of widely separated populations in regularly disturbed habitats, the species is likely to be threatened on a local or state basis.
ACKNOWLEDGMENTS
The authors wish to express their appreciation to the staff and curators of the following herbaria for the loan of specimens and/or for providing images and label data from specimens in the Orobanche riparia alli- ance: A, ARIZ, COLO, CS, F, GH, ILL, IND, JEPS, KS, MICH, MIN, MO, NEB, NY, OSC, OKLA, PH, POM, RSA, TENN, TTC, UC, US, UWM, WIS, WTU, and WVA. Collins is grateful to Ken Heil (San Juan College) for accompanying him during field work in the southwestern U.S. Financial support for some of the field work of Collins was provided by a grant from the Evangel University Faculty Summer Research Program. Colwell's initial contributions to this project during her doctoral studies in St. Louis were supported by the Department of Biology at Washington University. The Flora of North America Project additionally provided funding for one of Colwell's trips to the Missouri Botanical Garden.
REFERENCES
Beck von MANNAGETTA, G. 1890. Monographie der Gattung Orobanche. Biblio. Bot. 4(19):78-85.
Beck von MANNAGETTA, G. 1930, Orobanchaceae. In: A. Engler, ed. Das Pflanzenreich: Regni vegetabilis conspectus im Auftrage der Kóngl. preuss. Akademie der Wissenschaften vol. IV (261), issue 96. Wilhelm Engelmann, Leipzig. Pp. 1-348.
BENNETT, J.R., AND S. MarraEws. 2006. Phylogeny of the parasitic family Orobanchaceae inferred from phytochrome A. Amer. J. Bot. 93:1039-1051.
Cotuns, LT. 1973. Systematics of Orobanche section Myzorrhiza. Ph.D. dissertation, University of Wisconsin, Milwaukee.
HeckarD, L.R. 1973. A taxonomic reinterpretation of the Orobanche californica complex. Madroño 22:41-104.
Collins et al., Orobanche riparia, a new species 11
KARTESZ, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland, 2 vols. Timber Press, Portland, Oregon.
Munz, PA. 1930. The North American species of Orobanche, section Myzorrhiza. Bull. Torrey Bot. Club 57: 611-624.
Munz, PA. 1974. A flora of southern California. Univ. of California Press, Berkeley.
MussLeman, L.J. 1982. The Orobanchaceae of Virginia. Castanea 47: 266-275.
NurraLL, T. 1818. The genera of North America plants, and a catalogue of the species, to the year 1817, 2 vols. Published by the author (printed by D. Heartt), Philadelphia, Pennsylvania.
OLMSTEAD, R.G., C.W. DePampnitis, A. Wotre, N.D. Young, W.J. ELison, AND P. Reeves. 2001. Disintegration of the Scrophu- lariaceae. Amer. J. Bot. 88:348-361.
ScHNEEWEISS, G.M., A. CouweLL, J.-M. Park, C.-G. JANG, AND T.F. Steussy, 2004. Phylogeny of holoparasitic Orobanche (Orobanchaceae) inferred from nuclear ITS sequences. Molec. Phylogen. Evol. 30:465-478.
Shreve, F. AND ILL. Wicains. 1964. Vegetation and flora of the Sonoran Desert, 2 vols. Stanford University Press, Stanford, California.
SrRAUsBAUGH, P.D. AND E.L. Core. 1978. Flora of West Virginia, ed. 2. Seneca Books, Grantsville, West Virginia.
I Al S£ Das anli D L De if
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o r T: rr Pul stars for surpassing the highest standards.
This book is divided into five chapters: (1) Evolution, Classification, and Botanical Characteristics; (2) Ecology and Habitats of Cacti, Agaves, and Yuccas; (3) HK Profiles; (4) Conservation and d and (5) Exploring the Cactus Country of California and Nevada. The eight appendices a lly informative and increase the value of the book: Cactus, Agave, and Yucca Species of California and Nevada; Men Vegetation Types and ppt for Cacti, Dee dy Yuccas; senec Names ie Non- ates Species; Cholla Species C C
y Species List for ME Public tended in Califonia and Nev ada The species profiles for each entry includ name, the scientific name, a phonetic pronunciation guide, recent syn- elie oth common dé a commentary al he plant, a landscape, habit, and flower photograph pue a range map. Ad ing the species. The chapt
ipti f the habit, stems, spines or leaves, flowers, and f pprot on oo ation 2 Cultivation”: pecially y ] ion i Siena aves cultivation
A E 3111 1 1 1 1 1
g ? li ] ific si 1 lected i be found on
bs section on on "Exploring n Cactus Country of calina ae Ney ada ate parks, national parks, nati I sildeenéss areas, wildlife ee and recreation areas. c : 1 wa. : 1 ] id i y common, in igenous rare I new species are
th llent | th dagl hat defi ] inol lin the book. The 15 genera of Cactaceae due cli
that occur in California. Ru Nevada are a by d ios ao paintings by Eugene O. Murman and th M dd Susan text. For cad “Packrat Middens me Vegetation Histor *Yuccas and Yucca Moths" in both dd to tl iosi iated plants, especially the yucca moth-flower o biology often featured in biology textbooks as obligate un. tho push the Pus are not M in ue text, the pees has more n 200 citati ly published in the last ten
Bazell. T
*t*r.l
ears. J ing ted afr a : 1 decr ] A px
by g genus and species li
s book must ls on Hee E a did WB. botanist, a: on o landscaper, conservationist,
dde]
E E E 1 L JE 1 1 L th
and national parks E in Western states. Cachuma Press, the editors will be cherished by all bibliophilesi—Harold W. Keller, Ph.D., Research Associate Botanical Research Institute of dois Fort Worth, Texas
76102-4025, U.S.A
J. Bot. Res. Inst. Texas 3(1): 12. 2009
BRISTLECONE PINE DWARF MISTLETOE: ARCEUTHOBIUM MICROCARPUM SUBSP. ARISTATAE (VISCACEAE), A NEW SUBSPECIES OF WESTERN SPRUCE DWARF MISTLETOE
FROM NORTHERN ARIZONA Jared M. Scott Robert L. Mathiasen School of Forestry School of Forestry Northern Arizona University Northern Arizona University Flagstaff Arizona 86011, U.S.A. Flagstaff, Arizona 8601 1, U.S.A. Jared Scott@nau.edu Robert. Mathiasen@nau.edu ABSTRACT
We describe Arceuthobium microcarpum subsp. aristatae (Viscaceae), a parasite of bristlecone pine (Pinus aristata Engelm.) and Engelmann spruce (Picea engelmannii Parry ex Engelm.) in northern Arizona. oe and aer Me data were used to com- pare Á. microcarpum populations parasitizing bristlecone pine on the San Francisco Peaks, A a, with populations on Engelmann ME and a e (Picea pungens a from nto Pid in a and New Mexico. diit ieu e phenological, and
Francisco Peaks and Kendrick
rr r
Peak in northern veneer as a new als
RESUMEN Se describe A thobi i p subsp. aristatae (Vi ,un e de Pinus aristata NEN y de Picea engeimannii Parry e Engelm en el norte de Arizona. S 1 fol 5gi y g I p de A. microcarpum que D. 1
r 1 F Shee [PES f cee wees Aéxic Dis fala f, J hili Jad Anal
parasitan a Pi istata de San F i , Arizona, blaci i Pi l ti APEA EN Pod 1
mentan la clasificación de las SNAM de A A. microcarpum de San Fand Peaks y de eig Peak en el norte de Arizona como una subespecie nueva
Key Wonps: Arceuthobium, Picea engelmannii, Picea pungens, Pinus aristata, Arizona
Western spruce dwarf mistletoe (Arceuthobium microcarpum (Engelm.) Hawksw. & Wiens) is an important parasite of blue spruce (Picea pungens Engelm.) and Engelmann spruce (Picea engelmannii Parry ex Engelm.) in the southwestern United States (Mathiasen et al. 1986; Hawksworth & Wiens 1996). It occurs as far south as the Pinaleño Mountains, Arizona and Sacramento Mountains, New Mexico and as far north as the Kaibab Plateau, Arizona. However, western spruce dwarf mistletoe is most common in the White Moun- tains, Arizona and Mogollon Mountains, New Mexico (Mathiasen et al. 1986; Hawksworth & Wiens 1996). On the San Francisco Peaks, Arizona, it also parasitizes Rocky Mountain bristlecone pine (Pinus aristata Engelm.) (Mathiasen & Hawksworth 1980). Initially, the dwarf mistletoe infecting bristlecone pine on the San Francisco Peaks was classified as limber pine dwarf mistletoe (Arceuthobium cyanocarpum (A. Nelson ex Rydb.) Coulter & Nelson) by Hawksworth and Wiens (1972). It was later classified as A. microcarpum based on its host affinities and chemical characters (flavonols) (Crawford & Hawksworth 1979; Mathiasen & Hawksworth 1980; Hawksworth & Wiens 1996). Although Mathiasen and Hawksworth (1980) reported morphological, phenological, and host susceptibility differences between the A. microcarpum populations on the San Francisco Peaks and other A. microcarpum populations in the Southwest, they did not recom- mend giving the San Francisco Peaks populations taxonomic status at that time. In 2006, we began a more detailed analysis of the morphological characteristics of the A. microcarpum populations in Arizona. Our data further substantiated the differences between the A. microcarpum populations on the San Francisco Peaks and those in other mountain ranges of Arizona first reported by Mathiasen and Hawksworth (1980). Because of the differences in plant size, plant color, phenology, host affinities, and geographic isolation of
J. Bot. Res. Inst. Texas 3(1): 13 — 21. 2009
14 J t ical Insti Texas 3(
the dwarf mistletoe populations on the San Francisco Peaks, and nearby Kendrick Peak, there is sufficient evidence to describe these populations as a new subspecies of A. microcarpum.
Arceuthobium microcarpum (Engelm.) Hawksw. & Wiens subsp. aristatae J.M. Scott & Mathiasen, subsp. nov. (Figs. 1-2). Tre: U.S.A. Arizona. Coconino Co.: near summit of Schultz Peak, a southeast subsidiary ridge of the San Francisco Peaks, Coconino National Forest, elev. 3,060 m, Lat. 35? 18' 43" N, Long. 111? 37' 52" W, parasitic on Pinus aristata, 8 Aug 2006, J.M. Scott 2006-6 ( : ASC; IsoTYPES: JEPS, UNM, US).
Plantae (1237) lt li principales basi (121.8(—3) mm diam; fructus maturi 3.3 x 2.1 mm; anthesis mense Julio-Augusto;
antde i=;
; F F fructus maturitas Augusto—Septembri; in Pinus aristata et Picea engelmannii parasiticae.
Male plants (0.8-)2.7(-7.0) cm in height, basal diameter of dominant plants (1.0-)1.8(3.0) mm; female plants (1.4—)3.6(-7.0) cm in height, basal diameter of dominant plants (0.6-)1.8(3.0) mm; male and female plant plants primarily light green, but some brown-green or purple; male flower diameter 2.5 mm, perianth lobe dimensions 1.2 x 1.0 mm, anther diameter 0.5 mm, distance from anther to perianth tip 0.5 mm; fruit dimensions 3.3 x 2.1 mm; seed dimensions 2.4 x 1.1 mm.
Common name.—bristlecone pine dwarf mistletoe.
Etymology.—We selected aristatae, a name that indicates the principal host of this dwarf mistletoe is Pinus aristata.
Phenology.—Anthesis from early July to late-August with peaks in late-July to early-August; seed dis- persal from mid-August to late-September with peaks in mid- to late-August to early-September.
Habit.—Parasitic principally on Pinus aristata and occasionally on Picea engelmannii on Schultz Peak, parasitic principally on Picea engelmannii in the Inner Basin of the San Francisco Peaks and on nearby Ken- drick Peak. Not known on Picea pungens. Rarely infects Pinus flexilis James and Abies lasiocarpa (Hook.) Nutt. on Schultz Peak.
Distribution.—Bristlecone pine dwarf mistletoe only occurs in three small populations on the San Francisco Peaks (Schultz Peak, Weatherford Trail, and Inner Basin, Fig. 3). It also occurs near the summit of Kendrick Peak (Fig. 4). It ranges in elevation from 2890 to 3130 m on Schultz Peak.
METHODS
Collection Locations.—1n 2006 and 2007, plants of A. microcarpum were collected from 18 populations in Arizona (Appendix A; Appendices A—F available online at http://www.for.nau.edu/SOFArchive/MSF- Students/JMScott/20090325.pdf). Four populations were sampled near Flagstaff, Arizona (host plants): Schultz Peak (bristlecone pine and Engelmann spruce), Weatherford Trail (bristlecone pine), Inner Basin (bristlecone pine and Engelmann spruce) (Fig. 3), and Kendrick Peak (Engelmann spruce) (Fig. 4). A total of 12 populations of A. microcarpum were sampled in the White Mountains (host plants): Cache Cienega (blue and Engelmann spruce), Coleman Creek (blue spruce), Hannagan Creek (blue spruce), Johns Canyon (blue spruce), Lee Valley (blue spruce), Williams Valley (blue spruce), Forest Road 249 (blue spruce), For- est Road 402 (blue and Engelmann spruce), and Forest Road 72 (blue and Engelmann spruce) (Fig. 4). In addition, two populations were sampled from the North Rim of the Grand Canyon: Point Royal Road (blue spruce) and Route 67 (blue spruce).
In 1975 and 1976, plants of A. microcarpum were collected by the junior author from the same loca- tions and hosts on the San Francisco Peaks that were sampled in 2006-07. Plants were also collected from the North Rim of the Grand Canyon (blue spruce), the White Mountains (blue and Engelmann spruce), the Mogollon Mountains (blue spruce), and the Pinaleño Mountains (Engelmann spruce) (Fig. 4).
Morphological Measurements.—The following morphological characters of male plants were measured: dominant plant height, dominant plant basal diameter, flower diameter, anther diameter, perianth lobe length, perianth lobe width, and distance from the outer edge of the anther to the tip of the perianth lobe. The following morphological characters were measured for female plants: dominant plant height, dominant plant basal diameter, length and width of both fruits and seeds. The color of plants, fruits, and seeds were recorded. Plant heights and basal diameters were measured with digital calipers to the nearest 0.1 cm. All
Cents his ; A thah: : H aricfataa 15
Plants of Arizona VISAS
tatae
Arceuthobium microcarpam sap. aris da M; "ems Mathiassen
rona Tesoro C6 Cocenmo NF Servite Eevu tOnoze
Habitat: On birstiecone pine Jared Scot 2006.8 23 August 2005
Northern Arizona University (NAU)
Fic. 1. Holotype of Arceuthobium microcarpum subsp. aristatae J.M. Scott & Mathiasen, subsp. nov., J.M. Scott 2006-6, 8 Aug 2006 (ASC). Photo by R.L. Mathiasen.
; Arizona. Photo
F f 7,714 2008 hv 1 M Scott (Rott 1 Mala A
taken 30
May 2008 by J.M. Scott.
Schultz Peak, Arizona. Photo
Scott and Mathiasen, Arceuthobium microcarpum subsp. aristatae
San Francisco Peaks m
Arizona
Fic. 3. The four infestations of western spruce dwarf mistletoe on the San Francisco Peaks, AZ. Dark circles indicate an infestation on bristlecone pine, white circles indicate an infestation on Engelmann spruce, half-dark and white circles indicate an infestation on both bristlecone pine and Engelmann Spruce.
A LL ÉEc North im » e San D . Francisco Kendrick Peaks Peak @ a New . White Mexico Arizona „Mountains ss” e Mogollon Mountains inaleño Mountains PI te locations for plant collections of western spruce dwarf mistletoe in 1975 and 1976 (gray squares) and in 2006 and 2007 (closed circles). Plant lj 1f, : ER nara dd Sec a ap liqus 2007 (dosi of the Grand Canyon.
other morphological characters were measured using a 10x hand lenses with a micrometer to the nearest 0.1 mm. Male plants were collected during peak anthesis and female plants were uda NE fruits were
1
mature. Over E male or female ps were collected for each population and
1
were comp characters that were measured in 2006—07 were measured lou the 1970s ae Characters EXER for both datasets included male and female plant height and basal diameter, flower diameter, perianth lobe length and width, fruit length and width, seed length and width, and plant color.
Phenology Observations.—Anthesis and seed dispersal data were recorded when plants were collected for morphological measurements in 1975, 2006, and 2007. Data for the last two years were primarily from the San Francisco Peaks and Kendrick Peak due to frequent visits to these areas. During these years, the White Mountains were only visited twice each year: once during anthesis and again during seed dispersal. Therefore, the precise times of peak anthesis and seed dispersal were estimated for the White Mountains in 2006-07. In 1975, phenology data was collected from the San Francisco Peaks, Kendrick Peak, White Mountains, and Pinaleño Mountains, Arizona.
Host Susceptibility.—In 2008, temporary 0.012 ha (radius 6 m) plots tablished around 26 severely infected bristlecone pines on Schultz Peak and 13 severely infected Engelmann spruces in the Inner Basin. In each plot, all trees > 1.4 m in height were examined for dwarf incas dud For each tree, species, diameter at breast height (dbh; to the nearest cm), and a dwarf mistl g R, Hawksworth 1977) were
] qos 1 ] using ten randomly selecrea 1 plants f IOT each led Not el of
c 4 I Mash f A hah’ ` | | I aristatae 19
recorded. These data provided information on the relative susceptibility of bristlecone pine and Engelmann spruce to dwarf mistletoe infection on Schultz Peak and for Engelmann spruce in the Inner Basin.
Data Analysis. Morphological data were combined from hosts on the San Francisco Peaks (bristlecone pine and Engelmann spruce) and Kendrick Peak (Engelmann spruce) and also for the White Mountains and North Rim (blue and Engelmann spruce) for analyses. Morphological data from 1975 and 1976 were included within these two data sets, but the 1970s data also included morphological measurements from the Mogollon and Pinaleño Mountains. A one-way analysis of variance (ANOVA, P-value < 0.05) was used to test for significant differences between the means of each morphological character measured.
RESULTS
Morphological Data.—On average, male and female plants of western spruce dwarf mistletoe (WSDM, subsp. microcarpum) were larger than those of bristlecone pine dwarf mistletoe (BPDM, subsp. aristatae) and the differences in size were significantly different (Table 1, Appendices B-E, online). Although the largest plants we measured in 2006 and 2007 were collected from blue spruce in the White Mountains (Appendix B), the largest plants measured in the 1970s were from Engelmann spruce in the Pinaleño Mountains (Appendix E). Plant heights for the 1970s data were slightly larger than those for the 2006-07 data for BPDM, but both sets of data demonstrated that plants of WSDM were larger than BPDM. The mean basal diameter of male plants was similar, but the mean basal diameter of female WSDM plants was larger on average than female plants of BPDM and significantly different. The means for flower diameters, fruit lengths, and seed widths were significantly different, even though the differences were only about 0.1 mm or less (Table 1).
The color of male and female plants of BPDM was commonly light green or green-brown. However, many plants on the San Francisco Peaks appeared purple (Fig. 2). The color of male and female plants of WSDM was similar to BPDM, except that some plants in the White Mountains were green-blue.
Phenology.—Peak anthesis for BPDM occurred one to two weeks earlier on the San Francisco Peaks than for WSDM in the White Mountains in 2006 and 2007. Seed dispersal of BPDM also starts and ends one to two weeks earlier on the San Francisco Peaks than seed dispersal of WSDM in the White Mountains. The 1970s observations also found that BPDM flowers and disperses seed earlier than WSDM (Mathiasen & Hawksworth 1980)
Host Susceptibility.—Infection of bristlecone pine on Schultz Peak was 9696 (n = 111) indicating it is a principal host of bristlecone pine dwarf mistletoe there. However, infection of Engelmann spruce was only 3496 (n = 224) on Schultz Peak placing it in the occasional host class of Hawksworth and Wiens (1996). Engelmann spruce is clearly much less susceptible than bristlecone pine on Schultz Peak. However, infec- tion of Engelmann spruce in the Inner Basin, just 5 km north of Schultz Peak, was 94% (n = 101). All of the bristlecone pines growing near severely infected Engelmann spruces in the Inner Basin were infected, but there are only about 10 bristlecone pines in the area. We also observed rare infection of limber pine (2 trees) and subalpine fir (3 trees) by BPDM on Schultz Peak.
Table 2 summarizes the principal morphological and physiological differences between BPDM and WSDM used to distinguish the two subspecies. The shoot heights, colors, and phenology, of BPDM on the San Francisco Peaks and Kendrick Peak were similar enough to group them as one population. Morpho- logical data from Mathiasen and Hawksworth (1980) and our data demonstrated that BPDM plant heights were smaller than for WSDM populations in southern Arizona and central New Mexico. The differences in mean plant height between BPDM and WSDM were statistically significant. Additionally, color differences were found between BPDM (purple plants) and WSDM (blue-green plants). Furthermore, the periods of flowering and seed dispersal for BPDM on the San Francisco Peaks occurred one to two weeks earlier than for populations of WSDM in the White Mountains in 1975-76 and 2006-07.
Another important set of A ecc characteristics that separates BPDM from WSDM are their host affinities. Bristlecone pine pal host of BPDM on Schultz Peak (9696 infection), but we found that only 3496 of the Engelmann Soria were infected there. Mathiasen and Hawksworth (1980) also
20
TABLE 1. por : morphological characters of Arceuthobium microcarpum subsp. aristatae and A. microcarpum subsp. microcarpum. Data combine measurements from and 2006—07 for all hosts and are presented as means (ranges) [n]. Characters ul an asterisk had significantly different ica Using ANOVA (P < 0.05). The means for seed width were
rounded to the nearest 0.1 mm, but actual values g different. Character A. microcarpum subsp. aristatae A. microcarpum subsp. microcarpum Plant Height (cm)
Male* 2.7(0.8-7.0) [152] 5.6(1.8-14.9) [283]
Female*
Basal Diameter (mm) Male Female*
Flower Diameter SUBE
3.6(1.4-7.0) [177]
1.8(1.0-3.0) [121] 1.8(0.6-3.0) [167] 2.5(1.8-4.0) [287] 3:3(3.5-5.1) [281] 2.1(1.7-2.9) [281]
6.4(2.0-15.7) [353]
1.9(0.8-3.4) [157] 2.0(0.8-3.8) [313] 2.4(1.6-3.1) [266] 34(3.4-5.2) [440] 2.2(1.9-3.1) [440]
Seed Length (mm) 24(1.5-34) [107] 24(1.3-3.4) [224] Seed Width (mm)* 1.1(0.8-1.4) [107] 1.1(0.7-1.5) [244] Tagle 2. Summary of the principal E and physiological differences between Arceuthobi j 1 ubsp.
aristatae and A. microcarpum subsp. microcarp
Character A. microcarpum subsp. aristatae A. microcarpum subsp. microcarpum Mean Plant Height (cm)
Male Plants 27 5.6
Female Piants 36 64 Plant Color Light green; green-brown; purple Light green; green-brown; blue-green Peak Flowering Period One to two weeks earlier than One to two weeks later than
subsp. microcarpum subsp. aristatae
Host ad
Bristlec ne Principal Unknown
ey spruce Principal/Occasional? Principal
Blue spruce nknown Principal
Limber pine Rare Unknown
Subalpine fir Rare Rare
Southwestern white pine Unknown Immune 'Host susceptibility classificati j described in Hawl th and Wiens (1996). The host susceptibility designa-
tions used for A, nin un micorcarpum are based on information in Hawksworth and Wiens (1996) also.
25 | | fiar
| host for Subsp: aristatae in the Inner Basin of the San Francisco Peaks and on Kendrick Peak, Arizona, but it was an sal host of | n Schultz Peak.
reported that bristlecone pine was the principal host of BPDM on Schultz Peak, and they reported a similar level of infection for Engelmann spruce (32%), but they used a different method of collecting infection data; a systematic sampling design using 27 rectangular plots. Although they classified Engelmann spruce as a secondary host of BPDM on Schultz Peak, the low incidence of infection we found indicates Engelmann spruce should be classified as an occasional host there. Based on our observations and those of Lynch (2004), blue spruce is more susceptible to infection by WSDM than Engelmann spruce, but both are principal hosts of WSDM. Peas blue spruce has not been reported on Kendrick Peak or the San Francisco Peaks, we were unable to infection data for BPDM on this host. There is also a report of dwarf mistletoe on Chihuahua spruce (Picea chihuahuana Martinez) in northern Mexico by Ledig et al. (2000) which we assumed would be WSDM. However, our examination of several Chihuahua spruce populations in Chihuahua and Durango
Scott and Mathi , A thobi i | bsp. aristatae 21 reported to be infested with dwarf mistletoe, indicated that the spruces were infected with spruce broom rust (Chrysomyxa arctostaphyli Dietel) and not dwarf mistletoe. Spruce broom rust induces the formation of witches' brooms on Chihuahua spruce that are similar to those caused by dwarf mistletoes (Cibrián et al. 2007), which was undoubtedly the reason for the report of dwarf mistletoe on Chihuahua spruce in Mexico. Therefore, none of the Mexican species of Picea have been reported as the host of a dwarf mistletoe thus far (Hawksworth et al. 2002).
Hawksworth and Wiens (1972, 1996) defined a subspecies as geographically restricted populations having small, but consistent variations. Variations in dwarf mistletoe plant size, color, host range, and phe- nology have been used as the primary characteristics to identify subspecies of dwarf mistletoe (Hawksworth & Wiens 1965; Hawksworth & Wiens 1972, 1977, 1996; Hawksworth et » T Wass & Mathiasen 2003; Mathiasen 2007; Mathiasen & Daugherty 2007). Furthermore, the g lation of the San Francisco Peak/Kendrick Peak BPDM populations is also consistent with Hw and Wiens (1972, 1996) defini- tion that a subspecies should be a "geographically restricted population."
ACKNOWLEDGMENTS
We thank M. Socorro González Elizondo for the Spanish translation for the Resumen. We also appreciate the field assistance of James White and Laine Smith in northern Arizona and Brian Howell in northern Mexico. Gregory M. Filip and Del Wiens provided helpful suggestions to an earlier version of the manuscript.
REFERENCES
CIBRIAN, T.D., D. ALVARADO, AND S.E. GARCIA. (EDS.). 2007. Forest diseases in Mexico. Universidad Autonoma Chapingo, Mexico.
CRAWFORD, D.J. AND F.G. HawksworTH. 1979. Flavonoid chemistry of Arceuthobium (Viscaceae). Brittonia 31:212-216.
HawkswoRTH, F.G. 1977. The 6-class dwarf mistletoe rating system. Gen. Techn. Rep. R.M. U.S. Forest Serv. 48:1-7.
Hawksworth, F.G. AND D. Wiens. 1965. Arceuthobium in Mexico. Brittonia 17:213-238.
HawksworTH, F.G. AND D. Wiens. 1972. Biology and classification of dwarf mistletoes (Arceuthobium). Agriculture Handbook 401, USDA Forest Service, Washington, D.C.
Hawksworth, F.G. AND D. Wiens. 1977. Arceuthobium in Mexico: Additions and range extensions. Brittonia 29: 411-418
HAWKSWORTH, F.G. AND D. Wiens. 1996. Dwarf mist! biology, pathology, and systematics. Agriculture Handbook 709, USDA Forest Service, Washington, D.C.
HAwKSWORTH, F.G., D. Wiens, AND B.W. Gas. 2002. Arceuthobium in North America. In: Mistletoes of North American conifers. BW. Giels and T.D. Cibrian, tech. coords. Gen. Techn. Rep. R.M.RS. U.S. Forest Serv. 98:29-56,
HAWKSWORTH, F.G., D. Wiens, AND D.L. NickRENT. 1992, New western North American taxa of Arceuthobium (Viscaceae). Novon 2:204-211.
LEDIG, Eis M. ebay el B. diia V. HERNANDEZ Reyes, C. FLORES LOPEZ, AND me bin ARTEAGA. 2000.
Mexico and the d | f Pi Madroño
Locations of g | por graphy of! 47:71-88.
LyncH, A.M. 2004. Fate and characteristics of Picea damaged by Elatobium abietinum (Walker) (Homoptera: Aphididae) in the White Mountains of Arizona. W. N. Amer. Naturalist 64:7-17.
Mathiasen, R.L. 2007. A new combination for Hawksworth's dwarf mistletoe (Viscaceae). Novon 17:217-221.
MATHIASEN, R.L. AND C.M. DaucHerry. 2007. Arceuthobium tsugense subsp. amabilae, a new subspecies of hemlock dwarf mistletoe (Viscaceae) from Oregon. Novon 17:222-227.
MATHIASEN, R.L. AND F.G. HAwkswonrH. 1980. Taxonomy and effects of dwarf mistletoe on bristlecone pine on the San Francisco Peaks, Arizona. Res. Pap. R.M. 224:1-10
MATHIASEN, R.L., FG. HAw«swonrH, AND C.B. EDMINSTER. 1986. Effects of dwarf mistletoe on spruce in the White Moun- tains, Arizona. Great Basin Naturalist 46:685-689.
Wass, E.F. AND R.L. MATHIASEN. 2003. A new subspecies of A g (Viscaceae) from British Columbia and Washington. Novon 13:268-276.
7
BOOK REVIEW
Arnos Farjon. 2008. A Natural History of Conifers. (ISBN 978-0-88192-869-3, hbk.). Timber Press, Inc., The Haseltine Building, 133 S.W. Second Avenue, Suite 450, Portland, Oregon 97204, U.S.A. (Orders: www.timberpress.com, 1-800-827-5622). $34.95, 304 pp., 157 color photos, 39 line drawings, 7 3/8"
x 10 3/8".
1 A 3M» 3115 E ‘A J E J 11 x x 1 A x zt plant group y g y This is what we have in this volume. Mr. Farjon, retired head of tł ti f the Kew Herbarium, is an obviously dedicated botanist and always had a working knowledge of the conifers. | i d decidi ite a book on th j d porn) them all over again. io we are dus an extensive T of the conifers - enlightening naper on corrected classificati y methods
t y o DNA studies, g ore accurate ob of the group. T} 1 fé if, JL 1 es D 1 AR th lah
their viia is Hong and involved. DH course w are e also a pano of the pope Since some are fast-growing and quite dense. their
t connecti is very important b 2 2 E
ff, ] ¡lod inf, ; hath th
1 ae ne |
TI iting should be of i I ienti ientist alike. The photographs by tl I ; some of which depict trees you would have to travel long, difficult ways to see for yourself, make this a wonderful book.—Elaine Bell,
Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 3(1): 22. 2009
A NEW CLIFF-DWELLING SPECIES OF ZAMIA (ZAMIACEAE) FROM BELIZE
Michael Calonje Montgomery Botanical Center 11901 Old Cutler Road
iami, Florida 33156, U.S.A michaelcamontgomerybotanical.org
ABSTRACT Zamia meermanii (Zamiaceae), a new cliff-dwelli f l Belize is described. It is distinguished by having 1 to 3 pendent iid EE coriaceous ue that are entire or r erenulately nee on e uis un MM on A gda surface, and ich it most closely resembles, as well as to
E d
M : Jiff_q 1 š 7 ; hha} Gach Cchut & Deh or r£ O
RESUMEN
z> do Roli Cantal Pee € PD po : Se describe Zamia meermanii oe j tral q precip p guir porque tiene de l a 3 f T lad ] ee ee eee 2m
t T Y en la superficie adii. y persi la superficie abaxial. Esta especi I Zamia furfi L.f., la especie más his 1 T : des "e E Z3 semejante, yt Y A 1 4 A t Vovides, Schut & Dehgan y Z lovalii Nelson INTRODUCTION
Zamia is a New World genus ranging from the southeastern USA to Bolivia. It currently comprises 57 species (Hill et al. 2007) and is considered to be the most ecologically and morphologically diverse cycad genus (Norstog & Nicholls 1997). It includes species growing in areas with extremely high rainfall, such as Z. roezlii Linden from the Colombian Chocó region, and adjacent coastal Ecuador to species growing in full sun in extremely dry conditions, such as Z. encephalartoides Stevenson from the Santander region of Colombia. The genus also includes the only known obligately epiphytic cycad, Z. pseudoparasitica Yates, as well as two cliff-dwelling species: Z. cremnophila Vovides, Schutzman & Dehgan, and Z. sandovalii Nelson.
Jan Meerman, a Dutch ecologist who resides in Belize, li a third cun dwelling species of Zamia in February of 1999 together with Martin Meadows of the Belize Botanic G during a Rapid Ecological Assessment of a private protected area. Since discovering the plant, Meerman has peen collecting ecologi- cal and distribution data for this species, some of which is used in this description. He was instrumental in bringing a team from Montgomery Botanical Center to Belize to study this taxon and other Belizean Zamiaceae in August and September of 2008.
DESCRIPTION Zamia meermanii Calonje, sp. nov. (Figs. 1-2). Tr: BELIZE. Beuze D : 60 m, 28 Aug 2008, M. Calonje, J. Meerman & P Griffith BZ08-152 (moLoryre: BRH; isotypes: FTG, MO, NY, XAL). Species insignis habitui pulicol dex brevis, ape pendulis y litus; foliola coriacea ginil integris vel crenulatis triente apicali, is distinctis adaxialil pagini 3 : pro]
Stems globose to cylindrical, to 27 cm long and 18 cm wide, MM or occasionally branching on older plants. Cataphylls chartaceous, stipulate, triangular with lanceolate apex. Leaves 1—3 per stem, pendent,
2-146.5 cm long. Petiole 26-60 cm long with abruptly swollen base 2-3 cm wide, variously unarmed or carrying numerous prickles (50+) up to 3 mm tall. Rachis 50 to 109 cm long, unarmed or bearing a few prickles (« 20). Leaflets 7—23 pairs per leaf on adult plants, jam 3- > cm c from each other with the point of attachment to the rachis 8.5-13.3 mm wide, obovate t narrowly oblong, coriaceous, 12.5-32 cm long, 3.4-7.5 cm wide, with 31-56 veins prominently raised on adaxial surface, visible but not
J. Bot. Res. Inst. Texas 3(1): 23 — 29, 2009
Fic.1 (A) Habit of plant 1 M Inge ME fal . ight (Di E ey ee | : gl £4 le, E, " L hs y (0 Detail
of trunk and petioles. (D) Variation in leaflet sh (E) à f daxial surface of leaflet showi ised vei d late t r
flat L £: " lnc el : ETT RAT fev ct £ al. :4l £. laaflat L
duit leaf. A and C rep t Calonje et al. BZ08-152; B and F represent Calonje et al. BZ08-156; D, E and G represent Calonje
et al. BZ08-125.
Calonje, Zamia meermanii, a new cliff-dwelling species from Belize
[M 4 : : vn hl IDVA Emi Ave ile 2 IL 1 f^ id lu £ mat vll Fig. 2 f J t F 3 pi F t ve) b LÁ (PI Ad 1 £ hall /EY RA : tun? a PA (E\ Ra H trnhil $ 1 dehicri (G) Mature V F PM SAT ort J f 3 > 3 1 J uir nne "m el hull Dhatanranhe A E£ m 1 : r RUD + al fall E ire 2 rJ q“ t d 2" kil hab: lew RA f RA J u tc Calanie et al. R708- y j 152.
26 J t ical i Texas 3(
protuberant on abaxial surface; margins entire or crenulately notched in the distal third; surface on emergent leaves reddish-brown, but densely white-tomentose, turning cream-colored with saffron-orange tomentum, and gradually maturing to green and glabrous adaxially, with some persistent brown tomentum abaxially along leaflet margins and near the po of Su to the rachis. Eophylls typically carrying a single pair of ovate leaflets. } te strobilus conica l-cylindrical, erect, at pollen release 13-16 cm long, 2.2—2.5 cm diam. pu individually or in groups of 2—5 or more, cream to brown colored; peduncle 9-10 cm long, 0.9-1 cm diam., covered by ferrugineous pubescence. Microsporophylls arranged in 12-18 columns, 17-30 rows; obtrullate, distal face hexagonal to oblong-hexagonal, 5.5-6 mm wide, 2.5-3 mm tall, and 1-2 mm thick ; microsporangia spheroidal, 1-1.6 mm diam., present on abaxial surface, absent on adaxial surface, 22-28 per microsporophyll arranged in two separate groups along margins. Megaspo- rangiate strobilus cylindrical, solitary, erect at maturity, 12-20 cm long, 6-7 cm diam., tan to light-brown tomentose from emergence to maturity; peduncle 8-12 cm long, 1.5-3 cm wide, tan to brown tomentose. Megasporophylls arranged in 6-10 columns, 7-15 or more rows; distal face oblong-hexagonal, 1.5-2.2 cm wide, 1.0—1.5 cm tall, 1-1.2 cm thick, extruded to a narrow, depressed terminal facet. Seeds with sarcotesta ovoid-pyramidal, red at maturity, 2.0-2.5 cm long, 1-1.2 cm wide; sclerotesta ovoid to ovate, 1.5-1.8 cm long, 0.9-1.1 cm wide.
ined: BELIZE. Belize District: 40-70m, 25-26 Aug 2008, M. Calonje & J. Meerman BZ08-120 (FTG), BZ08-122 (FTG), BZ08-124 (FTG), BZ08-125 (BRH, FTG, MO, NY, XAL), BZ08-126 (FTG), BZ08-140 (FTG), BZ08-146 (FTG); 60 m, 28 Aug 2008, M. Calonje, J. Meerman & P. Griffith B208-154 (FTG); 70m, 27 Aug 2008, M. Calonje, J. Meerman & P. Griffith BZ08-156 (BRH). Etymology.—The specific epithet honors Jan Meerman, who discovered the species.
Distribution and habitat Endemic to Cayo and Belize Districts in Belize, it occurs in seasonally dry tropical evergreen broad-leaved lowland forests on steep karstic hills as defined by Meerman and Sabido (2001), reaching elevations of up to 200 m. The arboreal vegetation is characterized by species such as Co- mocladia guatemalensis Donn. Sm., Metopium brownei (Jacq.) Urb., Plumeria rubra f. acutifolia (Poir.) Woodson, Bursera simaruba (L.) Sarg., Forchhammeria trifoliata Radlk. var. trifoliata , Coussapoa oligocephala Donn.Sm., Caesalpinia gaumeri Greenm., Erythrina standleyana Krukoff, Pseudobombax ellipticoideum A. Robyns, and Thouinia paucidentata Radlk. Disturbed open spots are characterized by the endemic Louteridium donnell- smithii S. Watson. These forests are semi-deciduous, with more than half of the trees being leafless for at least one month during the dry season, and with several species being leafless for four months per year. This semi-deciduous cl implies that the amount of light reaching the forest floor (and under-canopy
cliffs) is dramatically different between the dry and rainy seasons.
Scattered throughout these hills are vertical limestone cliffs where this species is found. It appears to be an obligate cliff-dwelling species, as it is typically found growing in cracks and crevices on sheer vertical walls and absent from th ling forest floor. The cliff faces are largely bare but depending on the level of hering of the rock and the amount of shading they receive, the accompanying cliff vegetation consists largely of herbaceous plants, hemi-epiphytes and vines such as Adiantum tenerum Sw., Agave angustifolia Haw., Anthurium schlechtendalii Kunth subsp. schlechtendalii, Anthurium verapazense Engl., Philodendron radiatum Schott var. radiatum, Clusia sp., Begonia sericoneura Liebm., Pitcairnia recurvata (Scheidw.) K. Koch, Trades- cantia spathacea Sw., Passiflora cobanensis Killip, and Passiflora xiikzodz J.M. MacDougal subsp. xiikzodz.
Climate.—The average annual precipitation within this species’ range is estimated at 2000-2500 mm, with a distinct dry season occurring from February through May. The wettest month is July with an average monthly range of 300-440 mm, and the driest month is April with a range of 50-70 mm. The temperature ranges from 18°C to 31°C, with an annual mean temperature of 25°C. The coldest month is January and the warmest month is May. (Data derived from GIS analysis using Worldclim 1.4 climate layers as described by Hijmans et al. (2005)).
Conservation status. —The extent of for thi is estimated to be 750 sq. kmas delimited by the eee range occupied by suitable karst hills in Belize and Cayo Unas The area of occupancy,
1
which incl only these karst formations, is 300 sq. km. Based on an g population density
Calonje, Zamia meermanii, a new cliff-dwelling species from Belize 27
of 30 adult plants per km?, the estimated total population size for this species is 9,000 plants (Meerman, unpub. data). The karst hills where this species occurs are often isolated and separated by unsuitable habi- tats such as lowland forest, lowland savannas, and agricultural areas. In addition, this species occurs in discrete populations of only a few individuals, and does not appear to occur on all suitable habitats within its geographic range. As a result of the isolation between karst mountains and the clumped distribution of this species, populations are considerably fragmented. Visits to 12 different localities where this species Occurs appear to indicate that reproduction is occurring and seedling regeneration is healthy.
The primary threat to this species appears to be fire, utilized in adjoining areas for slash and burn agriculture. These fires have been observed spreading up surrounding karst hills and decimating native vegetation. Another important threat to this species is mining for construction aggregates and dolomite, with several active and planned quarries in the region. Recent fieldwork in Belize suggests that illegal wild collection of cycad plants and seeds has occurred in the past and may present an additional threat to this species. Hopefully, the difficult terrain and inaccessibility of this cliff dwelling plant, combined with the fact that approximately 5096 of its area of occupancy lies within protected areas will help minimize the risk of illegal harvesting. Specific locality information has been purposefully withheld in order to further
minimize this risk.
Due to its limited extent of occurrence and area of occupancy, the fragmented nature of its populations, and the threats caused by fire, mining, and illegal harvesting, this species should be listed as Endangered (EN) based on IUCN Red List criteria ab(i-iv) and 2ab(i-iv) (IUCN, 2001).
Reproductive phenology.—Elongating microsporangiate strobili have been observed in August and September in habitat, and pollen releasing strobili have been observed in od and OVID: under cultivation at Green Hills Botanical Collections near San Ignacio. A si with about a third of the seeds already dehisced, was collected in September of 2006, ud o en tócally dehisced with most seeds cleaned of sarcotesta, was collected in January 19 of 2006 by Martin Meadows of Belize Botanic Gardens. During the course of fieldwork with this species in August of 2008 our team found several distinct groupings of seedlings with newly-emerged eophylls, indicating that strobili probably disintegrated two to three months earlier. Further work is required to peier A the reproductive phenology of this species.
Ecology.—A large number of young seedlings observed during the course of fieldwork indicate that reproduction is healthy and the pollinating agent is active, although it has not yet been observed. Seedlings germinating on the forest floor suffer 100% mortality within a year or two, suggesting the forest floor habitat is unsuitable for this species (J. Meerman, pers. comm.). In August of 2008 our team observed a female plant with its peduncle attached and several seedlings growing directly underneath on the forest floor. Several of these seedlings from this same strobilus had also germinated in a fissure on the cliff directly above the mother plant, indicating that some unknown dispersal agent had moved individual seeds or a piece of the strobilus to this location. Larvae of Eumaeus toxea Godart butterflies were observed feeding on emergent
leaves. DISCUSSION
Zamia meermanii is most similar in appearance to Zamia furfuracea L.f. from southeastern Veracruz, Mexico. Both species have obovate to oblanceloate or narrowly oblong coriaceous leaflets with persistent tomentum on the abaxial side of leaflets, distinct veins on the adaxial side of leaflets, and a similar number of maxi- mum leaflet pairs per leaf (ca. 25). Zamia meermanii differs from Zamia furfuracea in having usually solitary stems as opposed to freely branching stems; holding 1-3 pendent leaves per crown, rather than 5-7 erect or slightly arching leaves per crown; eophylls with a single pair of leaflets rather than eophylls with two pairs of leaflets, leaves cream-colored and covered with saffron-orange tomentum just prior to maturing as opposed to light green with predominantly white tomentum prior to maturing; coriaceous leaflets to 0.8 mm thick compared to coriaceous or extremely coriaceous leaflets to 1 mm thick; leaflet length to 28
28 1 intel ID t + f Texas 3(1) Taste 1. D lucti | for Za ia furfi (d cliff-dwelli g Zamia species Zamia Zamia Zamía Zamia meermanii cremnophila sandovalii furfuracea Microsporangia per microsporophyll 22-28 14-18 14-18 40-42 Microsporangia diameter (mm) 1-1.6 0.65-0.78 0.6 0.8-1.2 Microsporangiate strobilus 9-10 2.5-3 6.7 8-10 peduncle length (cm) Megasporangiate strobilus length (cm) 12-20 8.5-14 9-17 10-24 Megasporophyll rows 7-15 4-8 4-7 4-15 Megasporophyll width (cm) 1.5-22 1.6-2.6 2.5-3.5 1.5-1.8 Megasporantiate strobilus 8-12 4—5 3.6-6 10-20 eduncle length (cm) Sclerotesta length (mm) 1.5-1.8 1.5-1.7 1.8-2.2 1.1-1.5
Data derived from measurements by author, Whitelock 2002, Nelson 2006, and Schutzman et. al. 1988.
cm compared to 18 cm; protuberantly raised veins on adaxial surface to 1.5 mm wide rather than slightly raised, narrower veins to 0.8 mm wide; and 22-28 microsporangia on abaxial side of microsporophylls as opposed to 40 or more on Z. furfuracea Zamia meermanii shares this unusual cliff-dwelling habit with two other Mesoamerican Zamia species: Zamia cremnophila from Mexico and Zamia sandovalii from Honduras. All three species typically carry 1-3 pendent leaves. Zamia meermanii is easily differentiated from the two other species by leaflet features alone (see key below). 1. Leaflets with distincly raised veins on adaxial surface and persistent tomentum on abaxial surface Z. meermanii
1. A | | I 1 : jore QE | f. lue . 1 "sl £.
Z. cremnophila Z. sandovalii
2. Adaxial surface of petiole with d 2. Adaxial surface of petiole tod ded without d qd groove
It has distinctly raised veins on the adaxial surface, persistent tomentum on the abaxial surface, and mar- gins that are entire or crenulately notched in the upper third, whereas Z. sandovalii and Z. cremnophila both have plane leaflets with no distinct raised veins on the adaxial surface, are glabrous on the adaxial surface, and have margins that are distinctly toothed on the distal half. Zamia sandovalii median leaflets are strongly falcate as opposed to those of Z. meermanii and Z. cremnophila, which are typically straight. Zamia meermanii and Z. cremnophila eophylls typically possess two leaflets, whereas Z. sandovalii eophylls typically possess four leaflets.
In addition to the vegetative characters that easily distinguish these three species, reproductive char- acters are also useful (Table 1). Zamia meermanii microstrobili have longer peduncles and possess more and hyll than Z. cremnophila and Z. sandovalii. Megastrobili of Z. meermanii
larger i I gi pet
can be longer and with more sporophyll rows and have longer peduncles than either Z. cremnophila or Z. sandovalii. Seeds of Z. sandovalii are the largest of the group, and those of Z. meermanii and Z. cremnophila are of a similar smaller size. Until a well-resolved phylogeny of Zamia is available, we have no way to determine if the cliff-dwelling habit evolved more than once in the genus.
ACKNOWLEDGMENTS
Funding in support of the 2008 Belize expedition which resulted in the description of this species was generously provided by the Association of Zoological Horticulture (AZH), Tim Gregory, and Montgomery Botanical Center (MBC). Jan Meerman of Biodiversity & Environmental Resource Data Systems of Belize introduced me to this species and participated in all fieldwork. Patrick Griffith (MBC) assisted in data and
Calonje, Zamia meermanii, a new cliff-dwelling species from Belize 29
specimen collection, and editing the manuscript. Green Hills Botanical Collections donated significant in- kind support by providing lodging, equipment and work infrastructure in Belize. Belize Botanic Gardens provided megasporangiate strobilus images and phenological data. Alan Meerow and Bart Schutzman pro- vided assistance with the Latin diagnosis and comments on the manuscript. Alberto Taylor (PMA), Anders Lindstrom of Nong Nooch Tropical Botanic Garden, Miguel Angel Pérez-Farrera (HEM), and Chad Husby (MBC) provided comments on the manuscript. The Belize Forest Department granted the scientific research and collecting permit (CD/60/3/08(45)) supporting research on Belizean Zamiaceae, as well as herbarium infrastructure for preservation of voucher specimens.
REFERENCES
HUMANS, R.J., S.E. CAMERON, J.L. PARRA, P.G. JONES, AND A. Jarvis, 2005. Very high resolution interpolated climate surfaces for global land areas. International J. Climatology 25:1965-1978.
Hitt, K.D., D.W. STEVENSON, and R. OsgorNE, 2007. The world list of cycads. Proc. 7th Int. Conf. on Cycad Biol. (CYCAD 2005), Xalapa, Mexico, January 2005. Mem. New York Bot. Gard. 97:454—483.
IUCN. 2001. IUCN Categories & Criteria, ver. 3.1. IUCN Species Survival Commission, Gland, Switzerland. Website: <http://www.rediist.org/info/categories_ criteria2001.html>. Accessed 16 October 2008.
Meerman, J. AND W. Sagipo. 2001. Central American ecosystems: Belize. Programme for Belize, Belize City. 2 volumes.
Netson, C.H. 2006. Dos plantas del genero Zamia (Gimnosperma) nuevas de Honduras. Ceiba 46:41-44.
Norstoc K.J AND T.J. NicHoLLs. 1997. The biology of cycads. Cornell University Press, Ithaca, New York.
ScHuTZMAN, B., A.P VoviDEs, AND B. DEHGAN. 1988. Two new species of Zamia (Zamiaceae, Cycadales) from southern Mexico. Bot. Gaz. 149:347-360.
Wuitetock, L.M. 2002. The cycads. Timber Press, Portland, Oregon.
BOOK REVIEW
Lynne CHAPMAN, NOELENE DRAGE, Di DursTON, JENNY JONES, HILLARY MERRIFIELD, AND BILLY West. 2008. Tea Roses: Old Roses for a Warm Climate. (ISBN 9781877058677, hbk.). Rosenberg Publishing Pty Ltd, PO. Box 6125, Dural Delivery Centre, New South Wales 2158, Australia. (Orders: www.rosenbergpub.com. au, 61-2-9654-1502, 61-2-9654-1338 fax). $59.95, 240 pp., color photographs, glossary, appendices, bibliography, index, 9 1/4" x 11 1/4”.
This beautiful book is presented by six ladies who love roses with a passi ] realized the lack of hi £d tralia. The tea rose or P Scented China rose hidd China f. ies. They were BUB to Europe in about 1820 and later to Australia. They | d, studied, and hybridized tt hth losing an g over and over again.
bia over two hundred pages covering every De of the tea rose, this book is a sioe work and just fascinating for a rose lover. T d in detail, from the horticultural and botanical facts to bs po history. ue teas are presented in all, each with a list of distinguishing features, cultivation background, location, The pepe is quite fantastic as is rud polo by the AE as as ems um close- NUS T aen MS views.
and other p Of additio note is the authors’ f di listinguisl ] ] f family ions. Tea rose “imposters” are discussed and a “What not to grow” list from 1922 is given. ook for the beauty of the rose and a book for the factual study of the rose, Tea Roses should strike a chord with a variety of
readers.— Elaine Bell, Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst, Texas 3(1): 30. 2009
A NEW SPECIES OF ZAMIA (ZAMIACEAE) FROM THE MAYA MOUNTAINS OF BELIZE
Michael Calonje Jan Meerman Montgomery Botanical Center Green Hills Butterfly Ranch 11901 Old Cutler Road and Botanical Collections
Miami Florida 33156, U.S.A Cayo District, BELIZE michaelcamontgomerybotanical.org Patrick Griffith and Geoffrey Hoese Montgomery Botanical Center 11901 Old Cutler Road Miami, Florida 33156, U.S.A
ABSTRACT Zamia decumbens (Zamiaceae), a new species from the Maya Mountains of Belize, is described and illustrated. It has affinity with
Zamia tuerckheimii, Zamia bussellii/onan-reyesii, and Z. monticola, to which it is compared.
RESUMEN
mu acce = - q 1 (7 : A le] nac M de Belize M t finidad Zamia ES
tuerckheimii, Zamia bussellii/onan-reyesii, y Z. monticola, con las que se compara.
INTRODUCTION
In December of 1997, while lucting research on wild nutmeg trees, John Janovec and Amanda Neill were led to the bottom of a large sinkhole in the Maya Mountains of Belize by naturalist guide Valentino Tzub to look at an interesting group of plants he called “corn palms.” At the bottom of the sinkhole, they found an old Mayan kiln and many pottery shards, and a dense population of Zamia (Janovec & Neill 2003). The plants did not match the description of any Zamia known from Belize at the time, but shortly after their return, Dr. Dennis Stevenson of the New York Botanical Garden informed them that the holotype of Zamia prasina Bull, a species described in 1881 (Bull 1881), had recently been re-discovered at the Kew herbarium. Janovec and Neill (2003) wrote that they connected these specimens to the sinkhole plants, and the name Zamia prasina has since been applied to this plant in the horticultural trade and several publications (see Balick 2000; Whitelock 2002).
In August, 2008, Montgomery Botanical Center (MBC) sponsored an expedition to Belize to shed some light on this poorly-known species and other Belizean Zamiaceae. During the course of the expedition, it became clear that the leaflets of these sinkhole-dwelling plants, which were chartaceous, gradually acumi- nate, with a distinctly raised longitudinal fold, and margins that are entire or with a few minute teeth at the apex, were very different from the leaflets of the Z. prasina holotype, which were coriaceous, abruptly acuminate, strongly serrulate and lacking a longitudinal crease. We determined that Z. prasina was in fact a prior valid name for Z. polymorpha DW. Stev., A. Moretti & Vazq. Torres (see Calonje et al., pages 43 — 49, this volume), and that the cycads found in the sinkhole belonged to an undescribed species.
During the cycad research expedition, the authors studied four separate populations of this species in Belize, gathering enough data to formally describe it.
SPECIES DESCRIPTION
Zamia d I Calonje, Meerman, M.P Griff. & Hoese, sp. nov. (Figs. 1-3). Tree: BELIZE. Torno District: bottom of sinkhole, 350—400 m, 2 Sep 2008, M. Calonje, J. Meerman & P Griffith BZ08-201 (HoLorYrE: BRH; isotypes: FTG, MO, NY, XAL).
Species Z. tuerckheimii Donn.Sm. affinis sed caule d bente et pedunculo pollinis longissimo.
J. Bot. Res. Inst, Texas 3(1): 31 — 41. 2009
Journal of the Botanical Research Institute of Texas 3(1)
Fic. 1. Habit and habitat of Z. decumbens. A. Adult female plant with naturalist guide Valentino Tzub. B. View from sinkhole bottom at type locality of Z. decumbens.
tains, Belize
Calonie etal.. Z. J} ,
16 mm
K
Fic. 2. Rey j f £7 d b All pl g pl j jfi yl | lity | I lation. A. C f I g bilus, adaxial side. B. C cti f | g bilus, abaxial side. C Rhopalotria sp., put i I ll fZ decumbens. D. Pharaxonotha SP., putative pollinator of Z. decumbens. E Microsporophyll abaxial side. F. Mi I iate strobilus. G. M g gi ti bil | Il | ivi y stag H M gas gi trobilus. |. Seed witt test 1.J. M I l yll f KM poro Í yll [ 1, abaxial side
34 i i Texas 3(
Stems globose to cylindrical, to 80 cm long and 6.7-11 cm wide, decumbent and occasionally branch- ing on older plants, in habitat often rotting at base and rooting adventitiously from along the horizontal stems. Vegetative cataphylls tan-tomentose, chartaceous, stipulate, shallowly triangular to triangular with short, brown apex; strobilar cataphylls tan-tomentose, chartaceous, narrowly triangular with elongated light-tan lanceolate apex. Leaves on adult plants 5-17 per crown, erect or arching, 90—175 cm long, light green when emerging, turning dark green when mature. Petiole 30—55 cm long with abruptly swollen base 2.2-3.2 cm wide, moderately to heavily armed with prickles up to 5.3 mm long. Rachis 60-120 cm long, unarmed or sparsely armed in the proximal third. Leaflets to 28 pairs per leaf, 2.8-4.1 cm wide, 17.5-29 cm long, spaced 2—5.5 cm apart from each other with the point of attachment to the rachis 4.5-6 mm wide, chartaceous, lanceolate with gradually tapering apex and distinct longitudinal furrow, straight or slightly falcate, margins entire or with a few teeth restricted to apex. Eophylls typically with a single pair of ovate leaflets. Microsporangiate strobilus conical-cylindrical, emerging erect with peduncles progressively lean- ing outward horizontally, at pollen release 10—16.5 cm long, 2.1—2.5 cm diam., occurring individually or in groups of 2-6 or more, cream to brown; peduncle 12-23 cm long, 0.9-1 cm diam., green with brown pubescence. Microsporophylls arranged in 16-18 columns, 26-36 rows; obtrullate, distal face hexagonal to oblong-hexagonal, 3.0-3.5 mm wide, 4.0—4.5 mm tall; microsporangia spheroidal, 1-1.2 mm diam., present on abaxial nae absent on ue surface, 12-18 per microsporophyll arranged in two separate groups
along margins. Meg bilus cylindrical, solitary, erect at maturity, 12-20 cm long, 5.7-6.4 cm diam. , emerging cream- color tomentose and maturing to glabrous light or dark green with patches of tan tomentum ing on inner facet and margins of megasporophylls; peduncle 7-13 cm long, 1.5-2 cm wide,
green to brown tomentose. Megasporophylls arranged in 8-12 columns, 4—16 rows; distal face hexagonal to oblong-hexagonal, 2.25-2.9 cm wide, 1.6-1.8 cm tall, 2.5-4 mm thick, with a depressed terminal facet 9.5-12mm wide, 3.5—4.55 tall. Seeds with sarcotesta ovoid to ovoid-pyramidal, red at maturity, 1.7-2.0 cm long, 1.1-1.45 cm wide; sclerotesta ovoid to ovate, 1.45-1.65 cm long, 1.05-1.43 cm wide.
Other vouchers examined: BELIZE. Cayo District: 170—200 m, 19 Aug 2008, M. edd J. Meerman, M.A. Perez-Farrera, B. Arevalo BZ08-040 (FTG), BZ08-041 eg BZ08-053 (BRH). Toledo District: 100—200 m, 9,11,12 Mar 1987, Davidse & Brant 32232 (MO); bottom of sinkhole, 700 m, 3 Oct 1999, Janovec & Neill 1185, 1186 (FTG). bottom of sinkhole, 300—320m, 1 Sep 2008, M. Calonje & P. Griffith BZ08-180 (FTG), BZ08-189 (FTG), BZ08-194 (FTG); bottom of sinkhole, Toledo District, 700 m, 2 Sep 2008, M. Calonje, P. Griffith, J. Meerman, & V. Tzub BZ08-222 (FTG), BZ08-224 (FTG), BZ08-225 (FTG), BZ08-227 (FTG), BZ08-231(BRH), BZ08-232 (FTG); 350-400m, 2 Sep 2008, M. Calonje, J. Meerman & P. Griffith BZ08-180 (FTG), BZ08-189 (FTG), BZ08-194 (FTG), BZ08-202 (FTG).
Etymology.—The specific epithet refers to the decumbent habit of the stems.
Distribution and habitat. —Known from several locations in the Maya Mountains of Belize in Toledo, Cayo, and Stann Creek districts at elevations of 150-700 m. Currently considered a Belizean endemic, but some populations occur near the border with Guatemala, and since the Maya Mountains extend into South- eastern Petén Province, it is likely to occur here as well. These locations were within Tropical Evergreen Broadleaf Lowland Forest variants as defined by Meerman and Sabido (2001) and over a number of geologi- cal formations, most notably late Cretaceous limestones, but also older Triassic metamorphic bedrock to a
lesser extent (Cornec d The authors h rved seven different f thi i f 1 throughout the Maya Moun-
tains of Belize, encompassing an area of approximately 2600 km". OR each of these populations was extremely small and restricted to rocky mountaintops and ridges or on the bottom of sinkholes. The largest populations found have been inside two steep-walled sinkholes in Toledo district, each approximately 50-80 m wide and 30-60 m in depth, with approximately 150 large plants of reproductive age. The mountaintop and ridge populations observed to date have been much smaller, with a maximum of 20 plants observed, all much smaller in size than those observed in the sinkholes. The factor limiting the distribution of this species appears to be reduced soil moisture. In the sinkholes, the plants are found growing near the sinkhole walls within the sinkhole overhang drip line, where there is less light and they are protected from direct rainfall. Few other plant species are present in these areas, in low densities, with large areas of unoccupied soil (Table 1). The mountaintop populations may benefit from the quick-draining substrate provided by
Calonie et al., Z J ,
Tage 1. À g | wit! t locality for Za d b Taxon Family Justicia sp. Acanthaceae Louteridium donnell-smithii Acanthaceae Adiantum macrophyllum Adiantaceae Syngonium podophyllum Araceae Chamaedorea geonomiformis Arecaceae visa veracruzana Aristolochiaceae Forchhammeria trifoliata var. trifoliata Capparidaceae lis spathacea Commelinaceae Dracaena americana Dracaenaceae ponen Ho URN Fuphorbiaceae Episcia pun Gesneriaceae Calatola d Icacinaceae Dorstenia lindeniana Moraceae
icus Sp. oraceae Passiflora lancetillensis Passifloraceae Peperomia sp. Piperaceae
Piper spp. Piperaceae Deherainia smaragdina var. smaragdina oo Myriocarpa heterostachya Urticac
Myriocarpa longipes i Urera Ho Urticaceae
the rocky environment. In Toledo district, other lithophytic species such as Chamaedorea adscendens and Chamaedorea schippii were also observed growing in the same environment.
Climate—The average annual precipitation within this species’ range is estimated at 1800-2800 mm, with a distinct dry season occurring from February through May. The wettest month is July, with an aver- age monthly range of 205-580 mm; the driest month is April, with a range of 50-70 mm. The temperature ranges from 16-31°C, with an annual mean temperature of 22-25?C. The coldest month is January and the warmest month is May (data derived from GIS analysis using Worldclim 1.4 climate layers as described by Hijmans et al. (2005).
Reproductive phenology.—The reproductive phenology of this little-studied species is not well known and no distinct periodicity is evident based on the authors' visits in September of 2008, as well as the examina- tion of photographic evidence archived at Montgomery Botanical Center from John Janovec and Amanda Neill's visits to the sinkhole populations in August of 1999, and June of 2001. Immature, pollen-releasing, and old microsporangiate strobili were observed during all three visits. Receptive megasporangiate strobili were observed in August and September, and near-mature strobili were observed in June and September.
Ecology.—Several microsporangiate strobili at pollen-release stage observed at both sinkhole localities harbored numerous snout weevils of an unknown Rhopalotria species, as well as clavicorn beetles of an unknown Pharaxonotha species (Fig. 2c, d). Both genera are believed to be pollinators of Zamia and have previously been found in mi giate strobili of Zamia furfuracea L.f. (Vovides 1991) and Z. pumila L. (Tang 1987), as well as entering nt exiting both microsporangiate and megasporangiate strobili in many natural Zamia populations in Panama (Alberto Taylor, pers. comm.). A preliminary examination of the Rhopalotria insects suggests that they are either conspecific or close relatives to Rhopalotria mollis, one of the known pollinators of Zamia furfuracea L.f. (William Tang, pers. comm.).
Ceratozamia robusta Miq. occurs throughout the range of Z. decumbens and both species were observed growing sympatrically at one location in Northern Cayo district. Zamia decumbens was also observed grow- ing together with Zamia variegata Warsz. at one location in southern Toledo district, with no evidence of hybridization. These two species are unlikely to overlap much geographically, as most observed populations of Z. decumbens occur above 300 m, which is the maximum of the altitudinal range of Z. variegata.
—
JOU OF
Notice
f Z. decumbens.
c. 3. Di
Fi
D. Eophylls
a
Calonie et al., Zamia d b ies from the Maya Mountains, Belize 37 J 7 7 f P 7
DISCUSSION
Zamia decumbens appears to be most closely related to Zamia tuerckheimii Donn.Sm. from Guatemala as well as a new species described from Honduras. The taxonomic priority for this Honduran species, described separately as Zamia bussellii Schutzman, R.S. Adams, J.L. Haynes & Whitelock and Z. onan-reyesii C. Nelson & Sandoval is currently in dispute and will poo be aed here as Z. bussellii/onan-reyesii.
Zamia decumbens differs from both species in 1 tems to 80 cm long that often rot at the base and re-root horizontally. Zamia tuerckheimii has erect or leaning stems to 3 m tall, and Z. bussellii/onan- reyesii has erect or leaning stems to 2 meters tall. All three species have leaflets with a marked longitudinal crease most prominent near the point of attachment to the rachis, but the leaflets of Z. bussellii/onan-reyesii are characteristic enough to distinguish it vegetatively from the two other species (Fig. 4). Zamia bussellii/ onan-reyesii leaflets are papyraceous and with prominent teeth on the lower margin compared to those of the other two species, which are chartaceous with margins entire or minutely toothed at the apex. Leaflets of Z. decumbens, measuring 2.8—4.1 cm in width, are Mia s narrower than those of Z. tuerckheimii, which
typically measure 4—9 cm in width. Another useful veg tinction between the two species is that Z. decumbens has moderately to heavily armed petioles wl | f Z. tuerckheimii are unarmed or sparsely armed.
Mature microsporangiate strobili of Zamia decumbens are easily distinguished from those of the two other species as they have long peduncles 11-23 cm in length causing them to lean outward, whereas Z. tuerckheimii has erect to slightly leaning strobili with peduncles to 6.5 cm long and Z. bussellii/onan-reyesii has erect to slightly leaning strobili with peduncles to 8.5 cm long. Zamia bussellii/onan-reyesii has the larg- est microsporangiate strobili, measuring up to 27.5 cm long and 3.8 cm in diameter, whereas those of Z. tuerckheimii measure 14-17 cm in length and 2.5 to 4 cm in diameter, and those of Z. decumbens measure 10-16.5 cm. Zamia bussellii/onan-reyesii is further differentiated from the other two species because it has more than 40 microsporangia arranged in a single group on the abaxial surface of the microsporophyll whereas 2. decumbens has 12-18 ón arranged in two separate groups and Z. tuerckheimii has 18-26 I gia arranged int The microsporophylls of Z. tuerckheimii are elongate triangular, approximately 1. 8-2. 5 times as one as they are wide, whereas those of Z. decumbens and Z. bus- sellii/onan-reyesii are roughly triangular and n a 1.3 to 1.5 times as long as they are wide.
TI bili of Z. bussellii/o the largest of the group, measuring 22.4—25.2
cm tall and 10- 11. 2 cm mi wide with sporophylls faces 4. 5-5 cm tall "m 4.3-4.6 cm wide, compared to those of Z. tuerckheimii which are 16.3—22 cm tall and 8.3-10 cm wide with sporophyll faces 1.8-2.2 cm tall and 2.3-3.8 cm wide, and to those of Z. decumbens which are 12-20 cm tall and 5.7-6.4 cm wide with sporophyll faces 1.6-1.8 cm tall and 2.25-2.9 cm wide. See the following dichotomous key and Table 2 for summary of diagnostic characters. The known geographical ranges of the three species do not overlap (Fig. 5).
KEY TO Z. BUSSELLII/ONAN-REYESII, Z. DECUMBENS, Z. TUERCKHEIMII
. Leaflets papyraceous with prominent teeth on lower margin Z. bussellii/onan-reyesii . Leaflets chartaceous with entire margins Or us toothed at the apex.
2. Stem decumbent | g 11-23 cm long Z. decumbens 2. Stem erect or leaning, ¡ gi bili peduncle to 6.5 cm long Z. tuerckheimii
One other species, Zamia monticola Chamb. appears to share some similarities with Z. decumbens in leaf length, leaflet shape and size, and microsporangiate strobilus size. It was described by Charles J. Cham- berlain based on a single male plant cultivated from seed reportedly collected near Naolinco Crater in the vicinity of Xalapa, Mexico. Chamberlain (1926) contends that the single plant sprouted from a batch of seeds thought to be Ceratozamia mexicana. The species has never been found again in the area where it was reportedly collected, and it is thought that it is either extinct, or that Chamberlain's seeds became mixed in his glasshouse and this species was acquired elsewhere (Hill 2004). Material collected in southern Alta Verapaz in Guatemala is currently being identified as this species, but this Guatemalan material is poorly
understood at this time. Considering the fact that the species was described from a single cultivated indi- vidual obtained under unusual circumstances, that its present geographic distribution remains unclear, and that its megasporangiate strobili are unknown, a detailed comparison cannot currently be made between this species and Z. decumbens. However, based on Chamberlain's description, this species appears to have an erect stem 14 cm in diameter compared to the stems of Z. decumbens, which are decumbent and up to 11 cm across. In addition, Z. monticola is described as having 20-32 microsporangia per microsporophyll, whereas Z. decumbens has 12-18.
Conservation status.—Although we observed seven different populations in the Maya Mountains add- ing up to an extent of occurrence of 2600 km”, the populations were all small, occupying a small area, and limited to the very specialized mountain top and sinkhole habitats. Despite visiting seven populations, only the two sinkhole populations had numerous large plants of reproductive size (about 150 each). The rest of the populations, located on rocky mountaintops, all had less than 15 individual plants of a much smaller size than those observed in the sinkholes. In total, less than 350 plants were observed within a combined area of occupancy of less than a square kilometer. Discussion with local residents throughout the range of Z. decumbens confirms that plants of this species have been extracted commercially in the past and seeds continue to be extracted for commercial purposes. During our fieldwork in the sinkholes, we found evi- dence of past mechanical removal of megasporangiate strobili, and a seemingly low number of seedling and young plants, perhaps suggesting that continued harvesting of seeds may be affecting the health of these populations. Specific locality information has been purposefully withheld from this paper in order to further
minimize the risk of illegal harvesting.
Considering how fragmented the distribution of this species is, most likely resulting from its specific habitat requirements, the small number of healthy, reproductively active populations observed, the small combined area of occupancy for this species, and the evidence of past and present commercial exploitation,
Calonje etal. Z
Belize
TABLE 2. $ y of di icd
Enr 7
tuerckheimii, Z. bussellii/onan-reyesii, and Z. decumbens.
Z. tuerckheimii
Z. bussellii/onan-reyesii
Z. decumbens
Microstrobilus length Microstrobilus width Microstrobilar peduncle
eng Microsporophyll shape Microsporangia
Megastrobilus length Megastrobilus width
Petiole armature
Leaflet length
Leaflet texture Leaflet margins
Eophyll leaflet pairs Stem height Stem habit
14-17 cm 2.5-4 cm to 6.5 cm
Elongate triangular
18-26 Arranged in two rows
Sparse or unarmed
19-30 cm to 14
cm Chartaceous
Entire or with a few
27.5 cm 38cm 8.5 cm
Broadly triangular ed in single row
22.4-25.2 cm
Light to moderate 15.5-36 cm to 23
Papyraceous rominent teeth on
minute teeth at distal end the lower margin
1
to3m Erect or leaning
4 to2m
Erect or leaning rotting at base and
11-16.5 cm 2.1-2.5 cm 11-23 cm
Broadly triangular 12-18 arranged in two rows
Moderate to heavy 17.5-29 cm to 28
Chartaceous
Entire or with a few minute teeth at distal end ]
to 80 cm long.
Decumbent, often
rooting horizontally
Zamia bussellii/onan-reyesii measurements derived from Schutzman et al., 2008. Zamia tuerckheimii measurements derived from Donnell Smith (1903), Standley & Steyermark (1958), Vannini 2008, and measurements taken by primary author. Zamia decumbens measurements taken in-situ by authors. All measurements based on mature plants.
we consider this species critically endangered and therefore recommend a Red List Category of CR for this species based on criteria B2ab (i, ii, iii, iv, v) (IUCN 2001). It ue be noted that although the criteria have changed somewhat based on new information, ntinuation of the Red List category currently listed for this species under the misapplied name “Zamia prisa
ACKNOWLEDGMENTS
The Association of Zoological Horticulture (AZH) funded our research and conservation work with this species through the grant "Zamia prasina in Belize." Tim Gregory and Montgomery Botanical Center sup- ported additional work in Belize.
John Janovec and Amanda Neill were the first to scientifically document and collect the sinkhole popu- lations of Z. decumbens, providing the inspiration for our return to Belize on a research expedition. Green Hills Butterfly Ranch and Botanical Collections donated significant in-kind support by providing lodging, equipment and work infrastructure in Belize. The Belize Forest Department granted the scientific research and collecting permit (CD/60/3/08(45)) supporting research on Belizean Zamiaceae, as well as herbarium infrastructure for preservation of voucher specimens. Valentino Tzub and Boris Arevalo provided field as- sistance. Willie Tang offered to identify the possible pollinators of Z. decumbens and provided the images of them. Jay Vannini pru information on Z. tuerckheimii and comments on the manuscript. Bart Schutzman helped in the Latin diag 1 provided comments on the manuscript. Anders Lindstromm, Chad Husby, and Alberto Sidney Taylor provided comments on the manuscript. Nancy Korber provided assistance in locating hard-to-find references. Dennis Stevenson provided herbarium specimen images and references.
E E di a § m. 4 BE vd e tn gett Tos Mexico 9o d - j 9 j i a S y d is f d f A iai S E A P ; ë | i y ah aw | Co | Belize a ns í + ES ] ] a ;o. #8 x AS ee z i Fa , = x a X \ | E ^m, ( pos i å "d Sy i y ff E i de i EOM Í » E $ i a? d E = č } a uel NEU cu Som Guatemala i p > | a H i Tr. — ooo «SN mos . h. Mi X we S zi e i SW ° "d 2 e. ^ a en e i p AA (Species | E Honduras | p IE Z busselli/onan-reyesii, | 4 Z decumbens | 4 ; | ates A 100 e Z. tuerckheimii : D km Á
Fic. 5. Geographical distribution of Z. bussellii/onan-reyesii, Z. decumbens, and Z. tuerckheimii.
REFERENCES
BaLick, MJ, M.H. Nee, AND D.E. Atha. 2000. Checklist of the vascular plants of Belize, with common names and uses
Mem. New York Bot. Gard. 85:246. Butt, W. 1881. Retail List 20.
CHAMBERLAIN, C.J. 1926. Two new species of Zamia. Bot. Gaz. 81:218-227
Cornec, J. 2003. Geology Map of Belize. Private Publication. Donnett SMITH, J. 1903. Undescribed plants from Guatemala and other Central American republics XXIV. (Zamia
tuerckheimii). Bot. Gaz. 35:8-10. IUCN 2001. IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN, Gland
Switzerland and Cambridge, U.K. HUMANS, R.J., S.E. CAMERON, J.L. PARRA, PG. JONES, AND A. Jarvis, 2005.Very high resolution interpolated climate surfaces
for global land areas V. Int. J. Climat. 25:1965- : Hitt, K. 2004. The Cycad Pages. Zamia monticola. Sydney: Royal Botanic Gardens. http//plantnet.rbgsyd.nsw 8].
gov.au/cgi-bin/cycadpg?taxname=Zamiat+monticola [Accessed 27 December 200 Janovec, J.P. ANO A.K. Neti. 2003. Exploring the palms and cycads of the Maya Mountains of Belize: reflections on
MBC-sponsored expeditions to Belize, 1999-2001. The Montgomery News 11(1):5-6.
MEERMAN, J. AND W. Sasipo. 2001. Central American ecosystems: Belize. Programme d Belize) Belize City. 7 ZU A A L aceae) aer
NELSON-SUTHERLAND, C.H. AND G.G. SANDOVAL-GONZALEZ. 2008. Una especie nueva de
Ceiba 49:135-136.
aadlil
] lU
a
Calonie et al., Z J 7
la decumbens, pecies from the Maya Mountains, Belize "
SCHUTZMAN, B., R. ADAMS, J.L. HAYNES, AND L.M. WurreLock. 2008. A new endemic Zamia from Honduras (Cycadales: Zamiaceae). Cycad Newsl. 31(2/3):22-26.
STANDLEY, P.C. AND J.A. STEYERMARK. 1958. Zamiaceae. In: Flora of Guatemala, Part |. Field Mus. Nat. Hist., Bot, Ser. 24:19-20.
TANG, W. 1987a. Insect pollination in the cycad Zamia pumila (Zamiaceae). Amer. J. Bot. 74:90-99.
VANINI, J.P. 2008. Notes on the Guatemalan cycad Zamia tuerckheimii Donn. Sm. Cycad Newsl. 30(3):4-7.
Vovipes, A.P. 1991. Insect symbionts of some Mexican cycads in their natural habitat. Biotropica 23:102-104.
WhrreLock, L.W. 2002. The cycads. Timber Press, Portland, Oregon.
BOOK REVIEW
Bonnie J. GiseL, with images by STEPHEN J. Joseph. 2008. Nature's Beloved Son: Rediscovering John Muir's Botanical Legacy. (ISBN 978-1-59714-106-2, hbk.). Heyday Books, PO.Box 9145, Berkeley, California, 94709 U.S.A. (Orders: www.heydaybooks.com, 1-510-549-3564, 1-510-549-1889 fax). $45.00, 247 pp., 9 1/2" x 12 1/4".
A 1 1 13.21 14 1 ža mi } des fTIAl € ees - C TON, | Es J Lt. P NEA O J 2 I GE and yes, his Eu as s explored and found p f t ifers, all of which he considered friends as well as expressions of divinity. The auth 1 photographs follows Muir's ventures into wilderness with assages from the explorer's own works, drawings from hisi journals and EUM d i d i ross the United p P Į p States. In a way the book is a botanical Bormo of Muir—not definitive, as that would take vol , but llent sampling of his botanical endeavors. Tl 1 1 1 +1 f. iH 1 4l ] ce | fil $4] f, 1 in th « p y I I I Plant Gallery Citations," with luced pi , a herbarium legend, and a quotation from Muir on his discovery of each plant. Di 1 1 T Jis 1 1 lod farh: : 1 i ; [1 1 : ; 1 e r d I I cC adding contrast, and eli 1 f he label d d 1 ] 1 d t Lo e a r Y Y "T 1 iE 1 ben bear es 1| N34 11 3111 e 1 11 1 J CNA. 4 i=) Joann Karges, Texas Christian University Library (retired), Box 298400, Fort Worth, Texas 76129, U.S.A.
J. Bot. Res. Inst. Texas 3(1): 42, 2009
WHAT IS ZAMIA PRASINA (ZAMIACEAE: CYCADALES)?
Michael Calonje Jan Meerman Montgomery Botanical Center Green Hills Butterfly Ranch 11901 Old Cutler Road and Botanical Collections Miami, Florida 33156, U.S.A. Cayo District, BELIZE ABSTRACT Maya Mountains: in Belize. An examination of the 1 ical ial of Zami ina, herbarium specimens, and wild Zamia popula- tions in Pipe ] ] d t l y to o this taxon but instead is a valid prior name for Zamia
] l 1 A lectotype for Z 4L ra
I» Is E
7 : 1 1
Key Worps: Nomenclature, Priority, Belize, Zamia prasina 1 I RESUMEN El nombre Zamia prasina se está aplicando en horticultura y literatura sistemática a un taxó ido de las Montañas Maya de Belize. U del material histórico de Zamia prasina, pecí de herbario, y poblaciones e de Zamia en imd nos 1] 1 1 A 1 1 1 z X: r L
yporl i prioridad 1 1. Se designa un lectotipo [ Z 1
INTRODUCTION
Zamia prasina Bull was described in 1881 in English nurseryman William Bull's horticultural catalog. Historically, this species has received little mention in the literature, but in recent years the name has been applied in the horticultural industry and systematic literature to a relatively unknown taxon from the Maya Mountains of Belize. We re-examined the historical material related to Zamia prasina as well as herbarium specimens and wild Zamia populations in Belize in order to uncover the true identity of this species, which is presented in the following analysis.
HISTORICAL MATERIAL AND EARLY USAGE OF THE NAME ZAMIA PRASINA
William Bull's 1881 horticultural catalog pa the ne ias for Zamia prasina:
“A hand d distinct-looking Cycad, i luced from H f g h Honduras, now Belize]. The stems in the young plants roundish or roundish-ovate, clothed with the imbri les left by the falling of the 1 The leaves are equally pinnate, the dark Eigen ae terete, with an indistinct PME in genu and Enea with afew small Men prickles. The leaf-blade is oblong-ovate, pinnate, I I entire, and tar base, where they are set on by a oy swollen articulation, denticul 1s tł he upp f f a bright grass-green colour, whence the name. 1 Y? guinea.”
Bull also provided a leaf of Zamia prasina, which was subsequently mounted as an herbarium specimen at Kew. In a letter attached to this specimen, dated March 22, 1881, he invites William Turner Thiselton-Dyer, at the time Assistant Director at Kew, to stop by his nursery and collect a leaf of Z. prasina. Although there was no notation on the specimen indicating when it was collected, it was probably shortly after Thiselton- Dyer received the invitation.
The specimen at Kew tated ible hol fZ. prasina by Dennis Stevenson of the New
York Botanical Garden on September of 1999. However, it has not been designated as a type for the species in any of the printed matter we consulted. This specimen, presumably provided by Bull the same year as he described the species in the horticultural catalog, undoubtedly represents the original material on which the species was based. However, because no reference was made to this specimen in the protologue, it cannot be considered a holotype under article 9.1 of the 2005 International Code of Botanical Nomenclature ("Vienna Code”). We remedy this situation by here designating this specimen as the lectotype for Z. prasina.
J. Bot. Res. Inst. Texas 3(1): 43 — 49, 2009
4 tani i Texas 3(
Zamia prasina Bull, Hort. Cat. 176:20. 1881. Ter: [BRITISH] HONDURAS [BELIZE]: cultivated from Belize, William Bull s.n., ar 1881 (Lecrorvre, here designated: K!). Zamia polymorpha D.W. Stev., A Moretti & Vázq.Torres, Delpinoa 37—38:3-8. 1998. Tyre: BELIZE. Caro: 22 Jan 1989, D. Stevenson et al. 1119 (uoLorvpe: NY; isotypes: BRH, FTG, MO, NY, U.
In addition to this specimen, there is an illustration of a leaf and caudex of Zamia prasina at Kew (dated 1881). A letter attached to the illustration (dated June 15, 1881) provides some background on Bull’s original importation of Zamia prasina. The letter, addressed to “J. Smith Esq.” (presumably Kew curator John Smith), mentions that a “Mr. Watson” (presumably Kew assistant curator William Watson) “talked him out of two plants that he did not want to give up.” He also mentions receipt of two separate shipments of Zamia from Belize and that each of the plants taken by Mr. Watson came from a separate shipment. The first plant, described as having “widely separate” leaflets was the only surviving plant from one of the shipments. The second plant, which Bull called Z. prasina, was one of the larger plants from the second shipment of which only a few plants survived. Mr. Bull clearly did not want to let go of the two plants and was asking Mr. Smith to return them. In return, he promised to provide “a couple” of the smaller surviving plants of Z. prasina for the collection at Kew. The illustration appears to be traced from the original Z. prasina plant provided by Bull before it was returned, as it is on a herbarium-specimen-sized sheet and bears the hand-written words “returned Zamia prasina.” Another inscription on the illustration reads “cf. Z. latifolia Lodd.,” and an inscription on the herbarium specimen reads “Z. latifolia Lodd.” These notes, in the same handwriting, were most likely added by Thiselton-Dyer, as he included Z. prasina in his treatment of Mexican and Central American cycads (Thiselton-Dyer 1884) as a synonym of Z. latifolia Lodd. ex A.DC (see de Candolle 1868). Thiselton-Dyer added a question mark next to his listing of Zamia prasina indicating he was uncertain about its synonymy with Z. latifolia. In fact, today it is not entirely clear what Z. latifolia is, as it was a horticultural name from Loddiges’ catalog for which no type specimen or illustration was provided. Miquel (1843) considered Zamia latifolia a synonym of Z. muricata var. obtusifolia, later bringing it to species rank (1849). In 1847, when he still considered Z. latifolia a variety of Z. muricata, he published an illustration of a leaflet labeled Zamia latifoliae (Linnaea 19(4): Tab. VII, fig. a. 1847), which Stevenson and Sabato (1986) selected as the neotype for Z. muricata var. obtusifolia (Fig. 1a). Since so little is known about the true identity of what was originally called Z. latifolia in Loddiges’s catalog, it must be considered a nomen dubium as well as its synonym, Z. muricata var. obtusifolia. Furthermore, the leaflet illustrated in Miquel's publication is unlikely to have been derived from a Belizean species, as none of the known species have leaflets with strongly serrated rounded leatlet tips. The leaflet shape and serrations in Miquel's illustration match closely those of Z. furfuracea plants at Montgomery Botanical Center grown from seed eed near Alvarado, Veracruz, Mexico (Fig. 1b).
Thiselton-Dyer clearly considered Bull's specimen to be di than Z. muricata Willd., as both species are compared on the Zamia prasina illustration in the same handwriting as the "Z. latifolia" annotations. The comparison indicates that Z. prasina has leaflets that are coriaceous, abruptly acuminate, and serrulate at the apex, whereas Z. muricata has leaflets that are chartaceous, gradually acuminate, and spinulose-serrate towards apex. Zamia muricata was originally described from Venezuela, and as currently circumscribed, is a species occurring only in Venezuela and Colombia.
In his 1932 cycad treatment, Schuster listed Z. muricata var. obtusifolia as well as Zamia prasina as synonyms of Z. loddigesii var. latifolia (Lodd.) J. Schust., incorrectly citing the publication date for Bull's horticultural catalog as 1822 rather than 1881 (Schuster 1932).
MODERN USAGE OF THE NAME ZAMIA PRASINA
Since Schuster's synonymization of Zamia prasina, the species name received little use until recent years, as apparently the original description and Bull's specimen were misplaced and not re-discovered until 1998 (Whitelock 2002). Since then, the name has been misapplied to a rare and little known taxon from isolated localities in the Maya Mountains of Belize (see Balick et al. 2000; Whitelock 2002; Janovec & Neill 2005; Hill 2004). This species, formally described in this issue (Calonje et al., page 31), differs from Zamia prasina in having an epigeous stem, leaflets that are chartaceous, gradually acuminate, with a raised longitudinal
Lenta I Jin RAS 1(184N RI flat nf 7, DEF AP £ I 4 Marta]
Fic. 1. A. Leaflet of Zamia muricata var. obtusi ifolia (= Ze Ji ] ) I near Alvarado, Veracruz, Mexico (MBC Accession # 20011305*A).
crease, and margins that are entire or with few minute teeth at the apex, rather than a hypogeous stem and leaflets that are coriaceous, abruptly acuminate, without a raised longitudinal crease, and strongly serrulate on the upper half (Fig. 2).
Since it is clear that the plant currently being called Z. prasina in horticult d recent literat not match the original description, lectotype or illustration of this species at Kew, then the logical ensuing question to ask is: What is Zamia prasina?
The only other described Zamia species currently known from Belize are Z. variegata Warsz. and Z. polymorpha DW. Stev., A. Moretti & Vázq.Torres. While Zamia variegata has strongly serrulate leaflets as described in the protologue for Z. aiio. ee are pola rather than coriaceous, and are variegated with yellow flecks, a unique and horti that Bull undoubtedly would have men- tioned in his horticultural catalog. However, e coriaceous, strongly serrulate leaflets of Zamia polymorpha exactly match the description of Zamia prasina.
A
SYNONYMIZATION OF ZAMIA POLYMORPHA
Zamia pol } tl ibed ranges from the Yucatan Peninsula in Mexico to Belize (Steven- son et al 1998). It was arene considered to be a form of Z. loddigesii Miq. with a highly variable karyo- Vee (see Mog & Olivares 1996). Stevenson et al. (1998) noted the karyotype variability as an important f this species in addition to its high level of pol; phism in leaf and leaflet morphology. This variability was ascribed to phenotypic plasticity due to dii censes in sun exposure, with plants exposed to full sun having shorter and narrower leaves with lanceolate leaflets, those growing in Mdb shade nas longer, broader leaves with ellit ] late leaflets, and those g gini
displaying leaf morphologies that are intermediate between the two extremes.
During a cycad research expedition to Belize in August of 2008 sponsored by Montgomery Botanical Center, an extensive survey of populations matching Stevenson et als description of Z. polymorpha with strongly serrulate leaflets uncovered some P ao with remarkably consistent vegetative morphology, and others that were highly variable. Tl ] 11
y consistent populations included 1 narrow-leafleted
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plants growing in savannas closely resembling Zamia loddigesii, and also other populations growing in tropical evergreen seasonal broad-leaved forests with larger leaves and wider leaflets. In addition to these morphologically consistent populations, others were found to be highly variable, containing the two forms mentioned above as well as intermediate plants.
The variability in these mixed populations appeared to be at least partially genetically determined, as plants with wide leaflets were sometimes found growing in full sun, and some narrow-leaflet plants were found growing in more shaded areas. Plants with narrow leaflets collected in an open savanna by the second author have retained their morphological characteristics despite years of growing in a shaded environment at Green Hills Botanical Collections.
A survey of one highly variable population halfway between Belmopan and Belize City uncovered in- dividual plants closely matching Bull's holotype (Fig. 3) of Zamia prasina and the drawing of this species at Kew (Fig. 4), as well as the holotype for Z. polymorpha (Fig. 5), indicating that Z. prasina and Z. polymorpha are the same species, and therefore Z. prasina takes nomenclatural precedence over Z. polymorpha.
In addition, there is also some circumstantial evidence suggesting that Z. prasina is a prior valid name for the species now considered Z. polymorpha. William Bull did not provide a locality or specific habitat information for where in Belize Z. prasina may have been collected, but two years after its description, Morris (1883) identified plants common on ridges and banks near Point Ycacos as belonging to this species. Morris did not provide a description or illustration of the plants he saw, but the cycad that is common in this area, visited by the second author, matches the description of Zamia polymorpha. Furthermore, this species is the most common cycad throughout Belize and would very likely be the first species encountered by collectors for William Bull.
Zamia prasina (now including Z. polymorpha) appears to belong to a species complex that includes Z. loddigesii and other related species such as Z. paucijuga Wieland, and Z. spartea A. DC (Schutzman 1987). Members of this a a display high levels of morphological variation (Gonzalez-Astorga et al. 2006) and may easily hybridize (Schut 1987). Although Schuster’s (1932) taxonomic work with cycads is not highly regarded (see Stevenson & Sabato 1986; De Luca 1990, Taylor et al. 2008), his inclusion of Z. prasina as a synonym of Z. loddigesii suggests that the name at the time may have been applied to plants within this P ne oe
l variability observed within populations of Zamia prasina, coupled with the high Ec RI vatisbility between p lations and the la ce a by Stevenson
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populations i d coriaceous,
and Sabato (1986), indicate that while it i serrulate leaflets represent one highly poyini species, further research including genetic sampling
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Fic. 3. Same-scal al. BZ08-086).
across a wide range of populations will be needed to better clarify the relationship between the different forms and their placement within the iia piod d iil complex. Until species nn are PEN we
cognition of Z I ith Z. polymorpha included within it ACKNOWLEDGMENTS Funding in support of the 2008 Belize expedition was g ly provided by the Association of Zoological
Horticulture (AZH), Tim Gregory, and Montgomery Botanical Center (MBC). Green Hills Botanical Col- lections donated significant in-kind support by providing lodging, equipment and work infrastructure in Belize. Patrick Griffith, Chad Husby, Anders Lindstrom, Walter S. Judd, and Bart Schutzman provided comments on draft versions of the manuscript. The Belize Forest Department granted the scientific research and collecting permit (CD/60/3/08(45)) supporting research on Belizean Zamiaceae, as well as herbarium infrastructure for preservation of voucher specimens. Dennis Stevenson and Andrew Vovides provided im- ages of historical material of Zamia prasina. Nancy Korber of Fairchild Tropical Botanical Garden graciously assisted me in the search of references.
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REFERENCES
BaLick, M.J., M.H. Nee, AND D.E. ATH. 2000. Checklist of the vascular plants of Belize, with common names and uses. Mem. New York Bot. Gard. 85:246. The New York Botanical Garden Press, Bronx.
Butt, W. 1881. Zamia prasina. Bull horticultural catalog. ÉS E
De Luca, P. 1990. A historical perspective on cycads fi tl t. Mem. New York Bot. Gard. 57:1—7.
DE CANDOLLE, A.P. 1868. Zamia latifolia. In: Prodromus systernati naturalis regni vegetabilis sive enumeratio con- tracta ordinum, generum specierumque plantarum huc usque cognitarum, juxta methodi naturalis normas digesta. Paris. P. 533.
GONZÁLES-ASTORGA, J., A.P. VOVIDES, P. OCTAVIO-AGUILAR, D. AGUIRRE-FEY, F. NICOLALDE-MOREJÓN, AND C. IGLESIAS. 2006. Genetic diversity and structure of the cycad Zamia loddigesii Miq. (Zamiaceae): implications for evolution and con- servation. Bot. J. Linnaean Soc. 152:533-544.
Hitt, K. 2004. The ean pages Zamia pana Sydney: Royal Botanic Gardens. http://plantnet.rbgsyd.nsw.gov. au/cgi-bin/cycadpg Zamia+prasina [Accessed 27 December 2008].
Janovec, J.P. AND A.K. Nei. 2003. Exploring the palms and cycads of the Maya Mountains of Belize: Reflections on MBC-sponsored Expeditions to Belize, 1999-2001. The Montgomery News 11(1):5-6.
McNett, J, ER. Barrie, H.M. Bunper, V. Demoutin, D.L. Hawksworth, K. MARHOLD, D.H. NicoLson, J., PRADO, P.C. Siva, J.E. Skoc, J.H. WIERSEMA, AND N.J. TuRLAND, eds. 2006. International code of botanical nomenclature (Vienna Code)
AE YORE PRA TAR d
Fi. 5. $ | pari f Zamia polymorpha holotype (right) | bet Imo; d Beli y (left, Michael Calonje et. al BZ08-085). Image of Zamia polymorpha holotyt | with permission from The C.V. Starr Virtual Herbarium of the New York Botanical
adopted by the Seventeenth International Botanical Congress Vienna, Austria, July 2005. Gantner Verlag, Ruggell, Liechtenstein.
Miquel, FAW. 1843. De cycadeis loddigesianis epistola. Tijdschrift Nat. Ges. Phys. 10(1):68-74.
Moue, FAW. 1847. Collectanea nova ad Cycadearum cognitionem. Linnaea 19:411—430.
Miquel, FA.W. 1849. Epicrisis systematis cycadearum. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 2:280-302.
Morass, D. 1883. The colony of British Honduras, its resources and prospects; with particular reference to its indigenous plants and economic productions. Edward Stanford, 55, Charing Cross, London.
SCHUSTER, J. 1932. Cycadaceae. In A. Engler, ed. Das Pflanzenreich 99(4):1-168.
SCHUTZMAN, B. 1987. Mesoamerican Zamias. Fairchild Trop. Gard. Bull. 42:16-19.
STEVENSON, D.W, A. Moretti, AND L. Gaupio. 1998. A new species of Zamia (Zamiaceae) from Belize and the Yucatan Peninsula of Mexico. Delpinoa 37-38:3-8.
STEVENSON, D.W. AND S. SABATO. 1986. Typification of names in Zamia L. and Aulacophyllum Regel (Zamiaceae). Taxon 35:134-144.
TAYLOR, A.S. J.L. HAYNES, AND G. HoLzMAN. 2008. Taxonomical, nomenclatural and biogeographical revelations in the Zamia skinneri complex of Central America (Cycadales: Zamiaceae). Bot. J. Linnaean Soc. 158:399-429.
THIsELTON-Dyer, WT. 1884. Cycadaceae. In: Helmsley, W.B. Biologia Centrali-Americana, Botany 3(16):190-195.
Vovipes, A.P. AND M. Otivares. 1996. Karyotype polymorphism in the cycad Zamia loddigesii (Zamiaceae) of the Yucatan Peninsula, Mexico. Bot. J. Linnaean Soc. 120:77-83.
WhrreLock, L.W. 2002. The cycads. Timber Press, Portland, Oregon.
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Botante Research institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 3(1): 50. 2009
FESTUCA ALOHA AND F. MOLOKAIENSIS (POACEAE: LOLIINAE), TWO NEW SPECIES FROM HAWATT
Pilar Catalán Robert J. Soreng and Paul M. Peterson Departamento de Agricultura (Botanica) Department of Bota T Satie Superior de el esca National Museum of Naural “a ersidad de Zaragoz mithsonian Instit Ctra. ree um 1, 22071 freu SPAIN Washington, DC oa USA, pcatalangunizar.es sorengr@si.edu, peterson@si.edu ABSTRACT
Festuca aloha, sp. nov. of Kaua'i and F. molokaiensis, sp. nov. from Molokz'i are described and illustrated. The two species are en- demic to the Hawaiian archipelago and grow on steep mountain slopes and volcanic cliffs with other native mesic flora. A key to the
species of Festuca found in the Hawaiian Islands is given.
Key Words: Festuca, new species, Hawaii, Loliinae, key
RESUMEN Se describen e ilustran Festuca aloha, sp. nov. de Kaua'i y F. molokaiensis, sp. nov. de Moloka'i. Las d peci démicas del sataj Torr ti 31,1 o lad = J; a f1 A R + E © 4 E mésicos € H 1 da 3 if L da] i Jo Eu Lakit 1 icl Ap] heel r L K e
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While revising specimens of Festuca deposited at the United States National Herbarium (US) the first author found two specimens of fine-leaved individuals from Kaua'i that were identified as F. rubra L. However, these individuals exhibit several characters that separate them from holarctic red fescues of the F. rubra complex, including; leaf blades flat with involute margins, sheaths open, relatively long ligules, and ovaries with densely hairy apices. After reviewing specimens of Festuca from Molokai'i deposited at the Bishop Museum (BISH) we noticed an additional taxon. Previously, the only species of Festuca and relatives re- corded from Hawai'i and the Pacific include the endemic, F. hawaiiensis Hitchc. (Hitchcock 1922), a robust broad-leaved species placed in F. subg. Drymanthele V.1. Krecz. & Bobrov sect. Banksia E.B. Alexeev (Alexeev 1980), and three Eurasian species: the fine-leaved F. rubra L. (F. subg. Festuca sect. Aulaxyper Dumort.); and the broad-leaved F. arundinacea Schreb. and F. pratensis Huds. [F. subg. Schedonorus (P. Beauv.) Peterm. sect. Schedonorus (P. Beauv.) Endl]. The taxonomy of Festuca s.l., the largest genus of monophyletic subtribe Loliinae Dumort., is i tate of flux. Much additional research is needed to properly align the taxa, althoug preliminary data from DNA sequencing (Catalán et al. 2007; Inda et al. 2008) suggest that two major clades may exist. As elements of the "broad-leaved" clade, the latter two species have been classified within the separate genus Schedonorus P. Beauv., as Schedonorus arundinaceus (Schreb.) Dumort and S. pratensis (Huds.) P. Beauv. (Soreng et al. 2003; Snow 2008). In addition, Festuca subg. Drymanthele is sometimes recognized as the genus Drymochloa Holub. The “fine-leaved” clade of fescues, including F. rubra, are placed in Festuca s.s. (Soreng et al. 2003). However, results are still equivocal concerning the resolution of the division at the base of the two major clades.
Specimens ined from the Bishop Museum (BISH), National Tropical Botanic Garden (PTBG), Missouri Botanical Garden (MO), and the United States National Herbarium (US).
Festuca aloha Catalán, Soreng & PM. Peterson, sp. nov. (Figs. 1A-J, 2). Tye: U.S.A. Hawar: Kava‘, Hanalei district, Kalalau Rim, Kalala, side bendi and W iun "i d neo. lookout, isolated hanging side-valley of iind diverse mesic forest
Peucedanum, Poa mannii, Hedyotis, 1 hia el Melicope
surrounded by F pallida, NA dtm. Dubautia. Dodd: POSER Li ta, Lobelia niil is, Lipidium, Neti Hibiscus kokio, Eragrostis, a Myrsine, Acacia, and Psychotris, 790 n m, 13 Mar 1992, K.R. Wood 1701 & S. Verba (HoLOTYPE: PTBG-17679; Iso TYPE: US-32522
J. Bot. Res. Inst. Texas 3(1): 51 — 58. 2009
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A Festuca hawaiiensis Hitchc. paniculis 8-11 cm longis, angustibus, ramis erectis lentibus; pedicellis (11.5—3.5(-5) mm longis;
lemmatibus 5.8-8 mm longis; laminis 1.5-2.4 mm longis, recedit.
Plants perennial, cespitose, with extravaginal innovations; cataphylls conspicuous, brownish. Culms 45-70 cm tall, erect, slender (ca. 1 mm diam.), nodes 23, smooth and glabrous. Leaf sheath margins fused for 1-2 mm at base, overlapping more than !^ the length below, sparsely villose abaxially when young, becoming glabrous at maturity, purple-brownish at the base, greenish above, becoming fibrous in age at base; auricles absent; collars glabrous; ligules 1-1.5 mm long, scarious, glabrous, brownish, apex obtuse, dentate-erose; leaf blades of vegetative shoots 22-33 cm x (12)1.5-1.7 mm, erect to pendant, flat with involute margins, glabrous and smooth abaxially, hirsute along protruding ribs adaxially, the hairs up to 0.1 mm long, cross section (Fig. 2) about 1 mm bearing 13-16(-20) larger vascular traces or veins, each with prominent ribs adaxially alternating with smaller ribs on the secondary and tertiary veins, scleren- chyma strands forming trabeculae or girders on most veins; leaf blades of culms about 22 cm x 2.4 mm, erect to pendant. Panicles 8-11 x 2.5—4 cm, erect, axis and peduncle smooth; loosely contracted with 34-66 spikelets, spikelets mostly at branch tips; panicle branches 2(3) per node, basal branches 4-6 cm and ca. 1/2 the length of panicle, angled, erect or ascending, smooth proximally to scabrous, angled dis- tally. Spikelets 8.2-13 mm x 3.5-6.0 mm, 4-6-flowered, broadly lanceolate, pale green, imes tinged with purple; pedicels (1-21.5-3.5(-5) mm long; rachilla internodes 0.8-1.2 mm long, scabrous; glumes lanceolate, green, margins very narrowly scarious («0.5 mm), apex acute; lower glumes (3.6-)3.9-4.2(-4.5) mm long, 1-veined; upper glumes 5.2—5.5(-6.0) mm long, 3-veined; lemmas 5.8-8.0 mm long, 5-veined, glabrous below, scabrous near the apex, green or tinged with purple at the apex, mucronate to awned, the mucro or awn 0.5-1.1 mm long, scabrous; callus rounded, obliquely angled (to 45°), smooth to dorsally minutely scabrous at the apex; paleas longer or shorter than lemma, scabrous on and between keels; stamens 3, anthers 3.1-4.2 mm long; ovary densely hairy on upper 1/3. Caryopsis not seen.
Additional material examined. U.S.A. Hawaiʻi. Kaua‘i: Hanalei district, Kalalau Rim, N of Kahuamaa flat, 990-1020 m, 3 Mar 1991, K.R. Wood 631, M. Query, S. Montgomery (PTBG-9498); north aspect, 1060—1190 m, 5 Apr 1991, K.R. Wood 901 & M. Query (PTBG-13876); Kalalau side below and W of the first una lookout, 13 Mar a K.R. Wood 1704 & S. Perlman (PTBG-17678, topotype?; Honopu Rim, undisturbed cliffs, r ith old i seen with Panicum lineale, 2800 ft, 6 Nov 1993, K.R. Wood 2832 (PTBG-29690; MO- pened Kalalau valley, Gumping on N facing al cliffs, seen with Poa mannii, Panicum lineale, Eragrostis vari- abilis, Carex Mn ca. 3000 ft, 17 Jun 1994, K.R. Wood 3245 (PTBG-20438); Pohakuao, hanging valley between Kalalau and Hanakoa,
and Kaaalahine Ridge, 2000 ft, 4 Jan 1992, K.R. Wood 1761, S. Perlman & J. Lau (BISH-621133, PTBG-12875, US-3250261); Waimea D. with Panicum lineale pn P. FPE 2000 ft, 30 Mar 1993, K.R. Wood 2470, & S. Perlman (PTBG-18788), K.R. Wood 2471
& S. Perlman (PTBG-18779); A alley, N facing slopes above stream, 0.5 mi along trail, growing with Wilkesia gymoxiphium and Lipidium serra, 3300—3500 ft, 18 May 1994, K.R. Wood 3193, S. Perlman & M. Hartley (PTBG-15270); Upper Kawaiiki: Kaluahaulu Ridge PP dh W of ] i iiis into Hager Acacia n Metrosideros iss morpha mixed mesic for- est with D 11 1 Poa ie eee S nd Cl
1149 m, 14 Nov 1996, K.R. Wood 7605 (PTBG-42661).
Comments.—Festuca aloha has panicles 8-11 cm long, narrow, branches erect and ascending; lemmas 5.8-8 mm long; and leaf blades 1.5-2.4 mm wide; in contrast F. hawaiiensis has panicles 30-40 cm long, widely open, branches patent or patent-erect; lemmas 9 mm long; and leaf blades 2-3 mm wide.
The illustrator Alice Tangerini noticed that all florets with mature anthers of F. aloha contained small, undeveloped ovaries (Fig. 1H). Mature ovaries with well-developed styles were found only in florets that had already shed their anthers (filaments were still present) [Fig. 11]. Therefore, F. aloha is apparently protandrous.
Etymology.—The specific epithet F. aloha name derives from the Kalalau mountains of Kauai.
Conservation status —Festuca aloha has been found in at least six different localities of Kauaii, covering a distribution area of approximately 700 km?. The number of individuals varies among populations but usually there are less than 1000 individuals per population. Based on these preliminary data, the new spe- cies clearly falls within Vulnerable (VU) category as defined by the IUCN (2001). The major threats to in- dividuals of F. aloha are the allochthonous plants, such as: Cyperus meyenianus Kunth, Erigeron sp., Kalan- choe pinnata (Lam.) Pers., Lantana camara L., Melia sp., Passiflora mollissima (Kunth) L.H. Bailey, Rubus
Ec? l £hlad na £ tati L £ Fact Inha Catalán Sorena & PM DP 1110 2353320 Y ). Scale bar = 0.1 mm Li 3 A
rosifolius Sm. ex Baker, Setaria gracilis Kunth, Triumfetta sp.; and goats and pigs (K.R. Wood, on herbarium label). Festuca molokaiensis Soreng, PM. Peterson & Catalán, sp. nov. (Fig. 1K). tw U.S.A. Hawari. Moroka‘: Kupaia Gulch, on steep slopes in mesic forest, occasional, associated vegetation: Metrosideros, Coprosma, Leptecophylla, Dodonaea, Alyxia, Viola, Melicope, Myrsine, Diospyros, Dryopteris, Phyllanthus, Luzula, Selaginella, A Carex, 21.119564 N 156.936999 W, 880 m, 3 Apr 2007, H.L. Oppenheimer H40704, S. Perlman & N. Tangalin (noLorvre: BISH-7287
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A Festuca aloha Catalán, Soreng & P.M. Peterson, p giori 1.5-2.8 mm, antheris brevioribus 2-3 mm, shins inferis (s. 0-)5.2- 5. sc 5.7) mm, EET superis (6.0-)6.2-6.8(-7.0) mm, oe Plants perennial, cespitose, with extravaginal innovations; cataphylls conspicuous, brownish. Culms up to 60-70 cm tall, scabrous below panicle. Leaf sheaths with margins fused for 1-2 mm at base, overlapping more than Y the length below, purple-brownish at the base, greenish above, becoming fibrous at base in age; auricles absent; collars glabrous; ligules 1.5—2.5 mm long, scarious, glabrous, brownish, apex obtuse, dentate-erose; leaf blades of vegetative shoots approximately 34 cm x 2 mm, more or less pendant, flat with involute margins, glabrous and smooth abaxially, hirsute along protruding ribs adaxially and on mar- gins and apex, dull green, cross section luu to that of F. aloha, about 2 mm wide bearing 25 larger vas- cular traces or veins, each with prominent ril ially alternating with smaller ribs on the secondary and tertiary veins, bulliform cells between adaxial ribs, sclerenchyma strands present forming trabeculae or girders on most veins, sclerenchyma strands with thin cell walls (adaxially) and thick cell walls (abaxially); leaf blades of culms about 21 cm x 2 mm, flat with involute margins, densely scabrous adaxially. Panicles 8.5-13 x 4 cm, loosely contracted with approximately 75 spikelets, spikelets located at the ends of the branches, 3(4) branches on basal node, 2(3) branches in other nodes, axis and branches strongly scabrous; basal branches as long as 7 cm, more than 1/2 the length of the panicle, angled, suberect to ascending. Spikelets 7-9 x 3-4 mm, 5-6-flowered, broadly lanceolate, pale green; pedicels Q— 4(-9) mm long; ra- chilla internodes scabrous; glumes lanceolate, green, margins very narrow, scarious, apex acute; lower glumes (5.025.2—5.5(5.7) mm long, 1-veined; upper glumes (6.0—)6.2-6.8(-7.0) mm long, 3-veined; lemmas (5.0-)5.5—6.0 mm long, 5-veined, scabrous dorsally and near apex, green, awn 1.5-2.5 mm long, scabrous; callus rounded, obliquely angled (to 459), scabrous; paleas slightly shorter than lemma, bidentate at apex, keels scabrous, between keels scabrous; stamens 3, anthers 2.8-3.0 mm long; ovary densely hairy on upper 1/3. Caryopsis not seen.
Comments.—Festuca molokaiensis differs from F. aloha by having more delicate and thinner, more nu- merous and lax branches; scabrous panicles and spikelets; lemmas scabrous throughout, awns longer 1.5-2.8 mm; anthers shorter 2-3 mm; lower glumes (5.0—)5.2—5.5(-5.7) mm; and upper glumes (6.0—6.2— 6.87.0) mm. In comparison to F. aloha, culms are scabrous on the nodes; panicles 8.5-13 cm long, more delicate, axis and peduncles scabrous, branches densely scabrous, angled; spikelets more delicate; rachilla more slender; lower glumes (5.0-)5.2—5.5(-5.7) mm long; upper glumes (6.0—)6.2—6.8(-7.0) mm long; lem- mas 5-6 mm long, evenly scabrous across the back from the base to the apex, 3-veined or infrequently faintly 5-veined (lateral veins obsolete or obscure), awned, the awn 1.5-2.8 mm long; and anthers 2-3 mm
long.
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Etymology.—The specific epithet of F. molokaiensis derives from the island of Moloka'i.
Conservation status.—Festuca molokaiensis is only known from the type locality where it has been re- corded as occasional (Oppenheimer, on herbarium label). Due to the scarcity of available data on the po- tential distribution, demography, and population status of F. molokaiensis, we consider to be a Data Deficient (DD) taxon (IUCN 2001).
TAXONOMY AND PHYLOGENY
Festuca aloha and F. molokaiensi logically similar to the Hawaiian endemic, F. hawaiiensis in ligule size and shape, ovary hairiness, and leal hee y (Hitchcock 1922). However, F. hawaiiensis is a more robust plant up to 150 cm tall with longer (30-40 cm) and wider panicles that are open with 3—5 spreading and drooping branches at the nodes, and larger lemmas up to 9 mm long. Festuca hawaiiensis grows at higher elevations, at ca. 2000 m, in rich soil on moist wooded hills on the island of Hawaii.
Festuca aloha and F. molokaiensis resemble the western North American, F. californica ssp. hitchcockiana (E.B. Alexeev) Darbysh. [Darbyshire & Pavlick 2007] in leaf blade cross section, panicle features, and hairy ovary apices. However, F. californica ssp. hitchcockiana differs from the new species by having a more robust and cespitose habit, possession of a continuous sclerenchyma layer on the abaxial side of the leaf blade, a partially hairy collar, and longer ligules with ciliate apices.
The new Festuca taxa also show similarities with Macaronesian volcanic cliff dwellers: F. agustinii Lind- ing., F. jubata Lowe, and F. francoi Fern.Prieto, e Aguiar, E. Días & M.I. Gut. (Saint-Yves 1922; Fernández Prieto et al. 2008) because all sh d flat to inrolled (or conduplicate) leaf-blades with some complete sclerenchyma cele However, the latter species all differ from the new species by having glabrous ovary apices and short (« 1.5 mm) ligules.
We analyzed the ITS and trnL-F sequences of two samples of Festuca aloha from different populations from Kauai [F. aloha 1: Kalalau, US-3252239 (isotype), GenBank GQ162205 (ITS) and GQ162208 (trnLF) accessions; F. aloha 2: Pohakuao, US-3250261, GenBank GQ162206 (ITS) and GQ162209 (trnLF) accessions] and one sample of F. molokaiensis from the only known population from Molokai [F. molokaiensis 1: Kupai Gulch, BISH-728771 (holotype), GenBank GQ162207 (ITS) and GQ162210 (trnLF) accessions]. We then conducted heuristic parsimony analyses for the combined ITS and trnL-F dataset (10,000 random entry trees, TBR, mulpars off, saving no more than 10 trees of length «10 per replicate; 1000 bootstrap replicates with the same parameters as in the original search) using these three samples and combined them with our previous data for Festuca on a worldwide level (Catalán et al. 2007; Inda et al. 2008).
All three samples grouped together within the broad-leaved Festuca clade (Fig.3), nested within the well supported clade Subulatae + Leucopoa p.p. [92% bootstrap(BS)], which were separated from F. califor- nica, F. rubra, and the M ian Festuca sect. Aulaxyper s.l. species. Festuca aloha was monophyletic (9996 BS) and sister to F. molokaiensis (98% BS).
The two F. aloha specimens from dn ode on Kauai (F. aloha 1 & F. aloha 2) had similar sequences (9996 BS; 1 trnL-F nucleotid titution difference); however, the sample of F. molokaiensis from Molokai showed several nucleotide diff in both the ITS (five substitutions) and trnL-F (two substitu- tions) regions with respect to F. aloha. These molecular differences also support the taxonomic separation of the two species. Despite the limited infraspecific sampling, the ITS variation found between F. aloha and F. molokaiensis, from the same Hawaiian archipelago, is remarkable as these species show more nucleotide differentiation than that found between other closely related species [e.g. the Pyrenean F. eskia Ramond ex DC., and F. gautieri (Hack.) K. Richt. with only two ITS nucleotide substitutions], and between species from different Macaronesian archipelagos (e.g., Madeiran F. jubata and Azorean F. petraea Guthn. ex Seub. with three ITS nucleotide substitutions). Unfortunately, F. hawaiiensis could not be included in the molecular analysis because it is extremely rare and we have no material other than the type.
The closest relative of F. aloha and F. molokaiensis in our analyses (Fig. 3) was the eastern Asian F. parvigluma Steud. (8896 BS), followed by the ] North American F. subulata Trin. (7796 BS), both belonging to F. subg. Subulatae, and then by the Siberian-North American F. altaica Trin. (9296 BS), placed
Festuca luciarum Festuca multinodis
à Vulpia membranacea Vulpia unilateralis Cienopsis mE Vulpia ci
u
Fesucrm Festuca violac Vulpía octo Hora
gautieri
Festuca quadriflora
Festuca rieulmis estuca thurbert
Festuca arundinacea Festuca atlantigena 1 Festuca leo urnexiana
Festuca gincescens
Festuca mai
Festuca fen
Festuca bačtica
foe einig pad
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Festuca aloha2
Festuca moloksiensis s parvigrum
[| l af pl JUuiriidi OF
Drymanthele s.s. gr.
Exaratae p.p./ American Vulpia
Leucopoa p.p. +
pee S.S. +
Aulaxyper s.l./ Vulpia 2x
Festuca/ Wangenheimia
Micropyrum Narduroides
Loretia gr. + Ctenopsis
Psilurus/ Vulpia (4x-6x)
Exaratae p.p.
American II
American ]
Eskia- Dimorpha gr. Leucopoa p.p. Neozeylandic I./ American
Schedonorus gr. Subulbosae Subulatae +
Asian-American
Leucopoa s.s.
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Fine-leaved Loliinae
Broad-leaved Loliinae
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within F. subg. Leucopoa sect. Breviaristatae. In a separate ITS strict consensus tree the F. aloha/F. molohai- ensis clade fell within the broad-leaved clade, and was closely related to the Asian F. parvigluma and F. modesta Steud. [sometimes treated as Drymochloa modesta (Nees ex Steud.) Holub]. In contrast to the ITS data, a trnL-F strict consensus tree placed F. aloha/F. molokaiensis in a polytomy at the base of the fine-leaved clade, among fine-leaved taxa and taxa intermediate between fine- and broad-leaved forms of Festuca.
Given the different resolutions between separate nuclear and plastid DNA sequence topologies for our species, it is possible that F. aloha and F. molokaiensis are of allopolyploid origin. On morphological grounds F. aloha, F. molohaiensis, and F. hawaiiensis are all part of the same complex, representing a common pattern of morphological speciation in upland habitats of different islands along the Hawaiian archipelago.
KEY TO THE SPECIES OF FESTUCA IN HAWAI
1. Lower leaf blades with auricles; leaf blades usually 5-12 mm wide, flat. 2. Auricles ciliate on margin; lemmas awned (awn 2—3 mm), more or less scabrous distally Festuca arundinacea 2. Auricles not ciliate on margin, lemmas muticous, smooth or only sparsely scabrous distally — baba pratensis 1. All le af bl f blades 0.3- 5m m wide, i involute to flat but when flat 3. Ligules 0.1-0.5 mm long; c ovary apices glabrous; leaf blades conduplicate; leaf sheaths usually closed below (open « Y their length); leaf blades of vegetative shoots in cross section without sclerenchyma girders. estuca rubra . Ligules 1-2.5 mm long; ovary apices densely hairy; leaf blades uu d Tel margina; m Aa generally open Y their length [they are fused only for 1-2 mm att fi in cross d with M dps Sues
w
4. Panicles 30-40 cm long, open, t | ling or drooping; leaf blades 3-5 mm wide ; known only dd M. wall Festuca hawaiiensis 4. Panicles 3.5-25 cm long, usually narrow, the branches patent, erect or ascending; leaf blades 0.3-2.4 mm wide. 5, qui scabrous only in the upper third, i liate veins distinct, awns 0.5-1 mm long; anthers 3.1-42 mm long; ligules 1-1.5 mm long; lower glumes (3.6-)3.9—4.2(-4.5) mm long; upper glumes eh 5.5(- is mm long; aca only from pale! i Festuca aloha
veins absent or obscure, awns 1.5-2.8 mm; ee 2-3 mm long: ligules 1.5— 25m mm long; lower glumes (5.0-)5.2—5.5(-5.7) mm long; RES gium (6.0-)6.2-6.8(-7.0) mm long; known only from Moloka'i Festuca end:
ACKNOWLEDGMENTS
This work has been subsidized by the Spanish Ministry of Science and Technology Grant Project CGL2006- 00319/BOS and Sabbatical Research Stay Grant PR2008-0020 to Pilar Catalán. We thank Stephen Darbyshire and Neil Snow for their critical review of an earlier version of the manuscript; Napua Harbottle, Tim Flynn, and Gerrit Davidse, for facilitating for us the study of Hawaiian Festuca materials deposited at BISH, PTBG and MO, respectively; Hank Oppenheimer, Neil Snow, and Cliff Morden for valuable information on the ecology and distribution of F. aloha and F. molokaiensis in Kauai'i and Molokai'i; Sasha Savytskyy, Dai Tsuchiya, and Juan Viruel for helping us with figure preparation; Alain Touwaide for correcting the Latin diagnoses; and Alice R. Tangerini for providing the illustration.
REFERENCES
ALEXEEV, E.B. 1980. Novye podrody i sekzii ovsjaniz (Festuca L.) severnoj ameriki | meksiki. (Festuca L. subgenera et sectiones novae ex America Boreali et Mexica). Novosti Sist. Vyssh. Rast. 17:42—53.[in Russian].
CATALÁN, P, P. TORRECILLA, J.A. Lopez-Ropricuez, J. MULLER, AND C.A. Stace. 2007. A systematic approach to subtribe Loliinae (Poaceae: Pooideae) based on phylogenetic evidence. Aliso 23:380—405.
DaRBYSHIRE, S.J. AND L.E. PAvuck. 2007. 14.01 Festuca L. In: Barkworth, M.E., K.M. Capels, S. Long, L.K. Anderton, and M.B. Piep, eds. Magnoliophyta: Commelinidae (in part): Poaceae, part 1. Flora of North America north of Mexico, volume 24:389—444. Oxford University Press, New York.
FERNÁNDEZ PRIETO, J.A., C. AGUIAR, E. Días, AND M.I. Gutiérrez ViLLARÍAS. 2008. On the identity of Festuca jubata Lowe
L D H In l.l PA fT,
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(Poaceae) and the description of a new Festuca species in the Azores Islands. Bot. J. Linnean Soc. 157:493—499.
HitcHcock, A.S. 1922. The grasses of Hawaii. Mem. Bernice Pauahi Bishop Mus. 8:100-230.
INDA, L.A., J.G. SEGARRA-MORAGUES, J. MULLER, PM. Mine AND P. CATALÁN. e Pains iS biogeography of the temperate Loliinae (Poaceae, Pooideae) grasses in t tl h Molec. Phylogen. Evol. 46:932—957.
IUCN. 2001. IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK.
Saint-Yves, A. 1922. Les Festuca (subgen. Eu-Festuca) de l'Afrique du Nord et de les Isles Atlantiques. Candollea
Snow, N. 2008. Notes on grasses (Poaceae) in Hawaii. Bishop Mus. Occas. Pap. 100:38-43. Soreng, RJ, PM. Peterson, G. Davipse, E.J. Jubziewicz, FO. ZULOAGA, AND O. Morrone 2003. Catalogue of the New World grasses (Poaceae): IV Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730.
A NEW GENUS, MEXOTIS, FOR FIVE MEXICAN SPECIES OF HEDYOTIDEAE (RUBIACEAE)
Edward E. Terrell Harold Robinson Research diy Department of Botany, MRC 166 Department of Botany, MRC 166, National Museum of Natural History National Museum of Natural History 20. PO. Box 37012 Smithsonian Institution Smithsonian Institution Washington, DC 20013-7012, U.S.A. Washington, DC 20013-7012, U.S.A. terr60@ msn.com robinsoh@si.edu ABSTRACT In furtl icti f th t of the g Hedyotis in Mexico, four | pecies, H. galeottii, H. kingii, H. latifolia, and H. terrel- lii, are m inanew eee Mexotis. A new species, M. lorencei, i gni d described. Morphological and distributional dat RESUMEN Enl icción adicional 1 pto del gé Hedyotis en México, lel peci idas, H. galeottii, H. kingii, H. latifolia
9 al +
^
ie. M. lorencei r i i
y H. terrellii,
y de distribución y se citan las PA representativas.
In early 20* century literature on the tribe Hedyotideae of the Rubiaceae the name Hedyotis L. was applied to many members of the tribe in both hemispheres. The main recent proponent of the name Hedyotis was Fosberg (1943) in his treatment of 24 Polynesian species of Hedyotis. He stated that he was adopting Hedyo- tis in the broadest sense because “the criteria to segregate the species involve not very fundamental differ- ences in structure" (Terrell 1996:2—5). Under Hedyotis he recognized five subgenera and commented that many botanists would consider these as genera. Despite his broad view of genera, he favored extreme split- ting for subdivisions of species, e.g., he described 39 M forms of Dou acuminata a & Schltdl.) Steud.(now Kadua acuminata Cham. & Schltdl.). He als idered seed cl because
“they vary so much that no two are alike in a single capsule". In subsequent papers rockers continued to use the name Hedyotis.
A recent study of Asian Hedyotis shows that many species related to the type, Hedyotis fruticosa L., of Sri Lanka and India, have unique capsule and seed morphology (Terrell & Robinson 2003). These species, in the genus Hedyotis, so far as known, are restricted to Asia and the western Pacific. Excluded from this concept are the mid-Pacific elements more recently treated as Kadua Cham. & Schltdl. (Terrell et al. 2005) and the New World species that have currently been placed in Hedyotis.
Many of the American Hedyotideae that were once placed in Hedyotis are now recognized as separate genera on the basis of structural features, particularly seeds and habit. Some of the segregate genera were described as new, including Carterella, with one Mexican species (Terrell 1987), Stenaria with five U.S. and Mexican species (Terrell 2001a), and Stenotis with seven Baja California and one Arizona species (Terrell 2001b). In addition, Houstonia and North American species of Oldenlandia each have five Mexican or Mexican-U.S. species (Terrell 1996; Terrell & Robinson 2006). In recent years further study has included about 12 additional hedyotoid genera. None of these genera were found to have seeds and other characters similar to those of the five species studied here.
We provide a further needed reclassification for Mexican Hedyotis species. A new genus, Mexotis, is described, including Hedyotis galeottii, H. kingii, H. latifolia, H. terrellii, and a newly described species, M. lorencei.
The removal of Western Hemisphere Hedyotideae from Hedyotis is supported by two recent studies
J. Bot. Res. Inst. Texas 3(1): 59 — 70. 2009
60 Journal of tani i Texas 3(
using DNA sequencing. In the DNA study of Andersson and Rova (1999), such species once placed in Hedyotis are not closely related to the Asian and eastern Pacific Hedyotis, the latter represented by H. mac- rostegia Stapf or the Central Pacific H. hillebrandii (Fosberg) Wagner & Herbst. (now Kadua axillaris (Wahra) Wagner & Lorence). In a more complete survey, Kárehed et al. (2008) show the division more clearly, and they call for the narrower interpretation of Hedyotis. Thus, the morphological and DNA evidence agree in the need to exclude American species of Hedyotideae from the genus Hedyotis. The Kárehed et al. study subdivides the Hedyotidae (as Spermacoceae) into many discrete clades that are not only distinct from Hedyotis, but distinct from each other. Two such groups are the Arcytophyllum/Houstonia/Stenaria Clade, to which many of the previously studied American Hedyotideae listed above belong and the Manettia/Bou- vardia branch, the latter being close to the Spermacoce Clade. No DNA sequences are available for any of the species placed here in Mexotis, and conclusions regarding which group is closest to Mexotis are based on structural features.
A paper by Church (2003) contributed an important molecular study on Houstonia and related genera, but did not include any of the five Mexican species that we investigate here.
Mexotís includes four species of large herbs or subshrubs with woody stems, long internodes, large elliptic or ovate leaves, long petioles, filiform pedicels, and generally flattened seeds with central punctiform hila. These species are native to Oaxaca and Veracruz. A fifth smaller species, M. latifolia, has herbaceous or woody-based stems and smaller leaves. It appears closely related to the four more robust species and ranges from Veracruz to Guatemala. This species is the only one of the five with a known chromosome number, n - 17, a number not known to occur in any other species of Hedyotideae.
The five species (Table 1; Figs. 1—5) have significant reproductive and vegetative characters. The seed types fall into two groups, the kingii group and the galeottii group. The kingii group is represented by M. kingii, the type species, which has somewhat thicker seeds, sometimes polygonal in outline and being often obtusely angulate or obtusely ridged. The compression is moderate or slight. These exact characters have not appeared in other Hedyotideae species. Mexotis terrellii is also in this group because of its apparent similarity to M. kingii; however, unfortunately the collections have immature seeds. The second seed type group includes M. galeottii, M. latifolia, and M. lorencei. These species have thin, flat or strongly compressed, concavo-convex seeds. In M. galeottii and M. latifolia the seeds are slightly to moderately concavo-convex. In M. lorencei the seeds become strongly concavo-convex and may appear cupulate. In M. galeottii and M. latifolia the seed areoles (cells) are radially elongated, and M. galeottii is unique in having narrow, usually partial, sometimes complete, wings. These same two characters also occur in the genus Bouvardia, which, however, carries the winged character to extremes, as many Bouvardia species have broad, conspicuous wings (and large capsules) (Blackwell (1968).
Mexotis Terrell & H. Rob., gen. nov. Tee species: Mexotis kingii (Terrell) Terrell & H. Rob.
Plantae UNS saepe Luis b i a herbaceae vel suffrutescentes. Caules ad 1.5 m alti; laminae foliorum ad 11 cm longae vel ellipticae; planatae aut leniter compressae interdum alatae, hilis centralis punctiformibus,
ovatae
He d PA Dal 1 1: 1
Perennial herbs or subshrubs typically with thick, woody, erect to decumbent, stems ca. 0.4-1.5 m long, in M. latifolia stems 15—55 cm tall, woody only at base. Leaves opposite, blades ovate to elliptic, acute to slightly acuminate, glabrous to sparsely or minutely puberulent or pubescent. Stipules interpetiolate. Inflo- rescences with slender or filiform pedicels, branching alternate or partially opposite. Calyx lobes lancolate. Corollas white or purplish to rarely rose, 3-13 mm long, funnelform, valvate, 4-lobed, not appendaged, glabrous to densely pubescent inside; heterostylous, with pin and thrum forms. Capsules subglobose to broader than long, where stages are first loculicidally then septicidally dehiscent. Seeds numerous, concavo- convex or obtusely ridged, one species with narrow wing, hilum central, punctiform, areoles isodiametric or radially elongated.
The genus name, Mexotis, is derived from Mex- as in Mexico and —otis as in Hedyotis, referring to an
, Mexotis, g f Rubi 61
KEY TO SPECIES OF MEXOTIS
1. Plants decumbent or erect, slender and only slightly woody at base, height 15-55 cm; leaf blades 4-32(-45)
x 3-24 mm; calyx lobes 0.4-1.5 mm long 2. M. latifolia 1. Plants erect, with woody stems and stout woody bases, height 45-150 cm; leaf blades 25-110 x 8-40 mm;
calyx lobes 1-6 mm lon
2. Stems and infl i ly hirtell g ls oft bvi ly wil iged 1. M. galeottii 2. Stems and infl lab t winged. 3. Stipules to 10 mm long; calyx lobes 5s 6 mm long; capsules 5.0-5.5 x 6-7 mm 5. M. lorencei
3. Stipules 2-4 mm long; calyx lobes 1-4.7 mm long; capsules 1-4.5 mm in diam. 4. Inflorescences 3-15 x 6-18 cm; pedicels 7-32 long; calyx lobes 1-2 mm long; corollas 5-6.5 mm
long; stipule appendages 1-2 mm long 4. M. terrellii 4. Inflorescences to 8 x 11 cm; pedicels 3-13 mm long; calyx lobes 1.5-4.7 mm long; corollas 7-13 mm long; stipule appendages only 0.3-0.5 mm long 3. M. kingii
1. a galcote s ban. Terrell & H. Rob., comb. nov. (Fig. 1A—D). Declieuxia galeottii M. Martens, in d. Roy: Sci. Brux. 11:231. 1844. Hedyotis Minas (M. Martens) Terrell & Lorence, Phytologia 66:1—4. 1989. Tyre: MEXICO. Oaxaca: Llano Verde, 3-7000 ft, Apr 1840, H. Galeotti 2603 BR!; isorvres: BR!, US!)
Manettia liebmannii Standley, J. Washington Acad. Sci. 17:337—338. 1927. Tere: MEXICO. Oaxaca: Pelado, Aug 1842, Liebmann 11485 (HOLOTYPE: C; ISOTYPE fragment: US 13157721). Paratype: MEXICO. Oaxaca: Cuesta de San Juan del Estado, Liebmann 11487 (C; frag- ment US 1315771!)
Perennial herbs or small shrubs. Stems 60—150 cm tall, to ca. 2.5 mm thick and woody toward base, stout, subterete, erect, glabrous or minutely hirtellous on upper stem and in inflorescence, few-several branched. Leaves with slender petioles 2-7 mm long, blades 40-85 x12—38 mm, ovate or ovate-lanceolate, acuminate glabrous above, glabrous or hirtellous on midrib beneath. Stipules 1-3 mm, broadly deltate, cuspidate with tip 1-2 mm long, glabrous or with few thick marginal teeth. Inflorescence with terminal, few-flowered cymes, branches densely and minutely hirtellous, flowers heterostylous, pedicels 2-7 mm long, very slender or filiform. Hypanthium glabrous; calyx lobes 1-2 x 0,3-0.7 mm, lanceolate, acute or submucronate, gla- brous. Corollas 4.5-9.7 mm long, funnelform or broadly funnelform, white, glabrous externally; tubes 2.8-4 mm long, 1-2.5 mm wide at base, 2.2-4.5 mm wide at throat, densely puberulent within; lobes 2.2—5.2 x1-2.5 mm, ovate, densely downy on adaxial face with white gland-tipped hairs. Pin flowers with stigma lobes 0.6-1.5 mm long, oblong or linear, exserted 1-2 mm beyond throat, anthers located at 3/4-point from base of corolla tube. Thrum flowers with anthers 1.0—1.5 mm long, linear or narrowly oblong, sessile or on filaments 0.5 mm long, attached at sinuses, stigmas located at midpoint of corolla tube. Capsules 2.2—4.0 x 2.2-4.5 mm, subglobose or somewhat compressed, 3/4—4/5 inferior, rather thick-walled, glabrous, loculicidally dehiscent. Seeds 0.8—1.2 x 0.8—1.2 mm, black, strongly compressed, very thin, flat, slightly to moderately concavo-convex, in outline orbicular or suborbicular, margins thin, wings present or absent, complete or partial, very narrow, 0.1-0.2 mm wide, fragile, hilum punctiform, testa finely reticulate, areoles radially elongated (Fig. 1). The pollen was found by Joan Nowicke (Terrell et al. 1986) to be similar to that of Hedyotis kingii. Chromosome number unknown. Distribution and Habitats.—Forests and rocky slopes. México: Oaxaca. Llano Verde area, Sierra de Juarez, north of the city of Oaxaca; Veracruz, in Sierra Madre Oriental northeast and northwest of Xalapa. Notes.—Additional nomenclatural and morphological notes were included in the Terrell and Lorence (1989) description of Hedyotis galeottii. The minute pubescence on the stems and inflorescence branches a the species. Addi i Labelled Hedyotis liebmannii ( l bove) and cited ! don data from D. H. Lorence. MEXICO Veracruz: Mpio. Alto Lucero, El Cerro la Cima, entre Plan de las Hayas y Tierra Blanca, 1600m, Castillo C. & Narave 2158 (XAL); Mpio. Atzalan, Cerro del Aguila, 850 m, 10 Sep 1982, Ventura A. 19740 (ENCB); Mpio. Atzalan, Alseseca, 950 m, 24 Mar 1975, Ventura A. 11118 (ENCB); Mpio. ii La Calavera, 1000m, 27 Apr 1978, Ventura A. 15232 (ENCB); Mpio. Naolinco, Naranjillo, 1250 m, 13 Nov 1976, Ventura A. 13605 (ENCB); Mpio. Yecuatla, Santa Rita, 1450 m, 12 pu 1973, Ventura A. 7824 (ENCB); Mpio. Yecuatla, Loma Santa Rita, 1480 m, 12 Jan hn Ventura A. 4764 (ENCB). Oaxaca: Yolox R de, al oesta de Cuasimulca, Miranda 1094 (MEXU).
2. Mexotis latifolia (M. Martens & Galeotti) Terrell & H. Rob., comb. nov. (Fig. 1E, F). Oldenlandia latifolia M.
£ 41 D A ID Li PS £T
62 Journal of Texas 3(
£-4- Ot hue
latifolia. Purpus 2951 ( NY), Veracruz. A, B, E. Seeds, ventral views, flat, gs. C D, F. Areoles enlarged.
Fic. 1. Seeds examined by SEM. A-D. Mexotis galeottii. A. Castillo C. & M 2158 (XAL) V B-D. Ventura A. 11118 (ENCB), Veracruz. E, F. Mexotis E A Il he I Init I tad ch .
owina hilum y
T Hu JI DAR: , Mexotis, g £ DAR: 63
Martens & Galeotti, Bull. Acad. Roy. Sci. Brux. 11:235. 1844. Hedyotis latifolia (M. Martens & Galeotti) Walp., Rep. 6:55. 1846.
Type: MEXICO. Veracruz: Mirador, at 3000 ft, near Jalapa, May 1840, Galeotti 2556 (holotype: BR; isotypes: F! GH! K!).
tune:
Hedyotis dichotoma Sessé & Moc., Fl. Mexic. ed.2. 20.1893, non H. dichotoma Cav., Icon. 6:53. 1801. Type: Sessé, Mociño, Castillo, <= Mala 2 aang y
Houstonia gracilis Brandegee, Zoe 5:238. 1907, non Hedyotis gracilis DC., Prodr. 4:419. 1830. Type: MÉXICO. Veracruz: on rocks near Orizaba, May 1905, C.A. Purpus 1251 (lectotype: UC! isolectotypes: F! MON.
Hedyotis exigula WH. Lewis, Rhodora 63:221. 1961. New name. Perennial herbs with roots sometimes woody. Stems 15—55 cm tall, occasionally rather woody at base, tet- ragonal or rounded-tetragonal, slender, erect, decumbent or spreading, glabrous or rarely pubescent, sparsely to much branched. Leaves with petioles to ca.10 mm long, blades 4—32(-45) x 3-14(-24) mm, median blades ovate to broadly elliptic or ovate-lanceolate, upper blades becoming lanceolate or linear- lanceolate, glabrous or puberulent above, glabrous beneath, base rounded to cuneate, apex obtuse or rather acute. Stipules to 2 mm long and wide, ovate or deltate, glabrous or short-pubescent, margin with several glandular teeth to ca. 1.5 mm long, apices truncate to lacerate. Inflorescence with terminal, few- flowered cymes, flowers heterostylous, closed at night, pedicels to 5(-10) mm long, very slender or filiform. Hypanthium glabrous or puberulent; calyx lobes 0.4—1.5 x 0.2-0.7 mm, scarcely to somewhat longer than capsules, erect, deltate to lanceolate, apices acute or obtuse, margins and sinuses glabrous or with white hairs to ca. 0.1 mm long. Corollas 3-7.3 mm long, funnelform or broadly funnelform, white, purplish, or rarely rose, glabrous externally; tube (222.3-3.3(-4) mm long, usually 1-2 times longer than lobes, some- what widened distally, 0.6-1.4 mm wide at base, 1.8-3 mm wide at throat, glabrous or puberulent within; lobes 13.3 x 0.6-2 mm, ovate, glabrous or puberulent within. Pin flowers with stigma lobes 0.4-1.3 mm long, linear, exserted 0.5—2 mm beyond corolla throat, style 2.4-4.3 mm long, filiform, anthers 0.5-0.8 mm long, 0.2 mm wide, narrowly oblong or elliptic, yellow or whitish, included near midpoint of corolla tube, sessile or subsessile. Thrum flowers with anthers 0.7—1.2 x 0.2-0.3 mm, narrowly elliptic or narrowly oblong, yellow or whitish, exserted at throat on filaments 0.2-0.8 mm long, stigma lobes 0.4-0.6 mm long, in- cluded near midpoint of corolla tube on style 1-1.6 mm long, filiform. Capsules 1-2.7 x 1-3.5 mm, usually slightly wider than long, or subequal, usually 3/4 to 9/10 inferior, moderately compressed, thin-walled, glabrous or with few hairs on calyx margins, cuneate or rounded toward base, apex retuse, dehiscing widely loculicidally and then septicidally. Seeds 8-24 per capsule, 0.5-1 x 0.4-0.9 mm, black, strongly compressed, very thin, slightly to moderately concavo-convex, in outline broadly elliptic to suborbicular, margins thin or slightly rounded, hilum punctiform, testa finely reticulate, areoles radially elongated (Fig. 1). Chromosome number: n = 17 (Lewis in Terrell et al. 1986, as Houstonia gracilis).
Phenology.—Flowering throughout year.
Distribution and Habitats.—Moist soil, shaded places, banks, slopes, among rocks, sides and bases of cliffs, montane rain forest; altitudes recorded for a few collections vary from 330-1430 m (1000-4300 ft). Mexico: Veracruz (mainly from Cofre de Perote and Jalapa south to Orizaba), northern Oaxaca, Chiapas, and west and central Guatemala.
Notes.—Stephen Koch reported (pers. comm. 1980) that this species was for sale in Coyoacan, Mexico City, as a house plant. It grows rapidly and continually produces numerous small flowers.
Standley (1918) listed Oldenlandia latifolia as a synonym of O. microtheca DC., but the type of O. latifolia (see above) has flat seeds and reticulate testas, as opposed to O. microtheca (Schltdl. & Cham.) DC. which has angular seeds with the areole walls coalescent. The basionym O. latifolia thus supplants the previously accepted names, Hedyotis exigula and Houstonia gracilis.
A paper on Oldenlandia (Terrell & Robinson 2006) provided a key to Oldenlandia microtheca and Hedyotis latifolia (Mexotis latifolia), two superficially similar species differing by seed characters and chro- mosome numbers (the former n = 11. latter n = 17).
Mexiotis latifolia is a variable species. Its relationships to the other four species in Mexiotis remain to be determined. It resembles Houstonia purpurea L. of eastern U.S. in aspect and leaf shape and size, but differs in seeds and reproductive characters.
Das A ID h l titt. ET
1 I £ al 5f 64 Journal of Texas 3(
Representative specimens: MEXICO. Chiapas: Lagos de Montebello, 42 km NE of La Triniteria, 23 Oct 1971, Breedlove & Thorne 21112 (DS, MICH, MO). Veracruz: mpio. Ixtaczoquitlan, Parque de Cerveceria Moctezuma, 1 km SE of Fortin, 970 m, 9 Nov 1978, Koch & Fryxell 78185 (CHAPA, ENCB, XAL); Cerro de Chicahuaxtla-Cuauhtlapan, 22 Jan 1968, Marino Rosas R. 1034 (CAS, MO); Mt. Orizaba, Seaton 101 (F, GH, K, NY, US); near Rio Blanco and Orizaba, 4300 ft, 15 Sep 1944, Sharp 44889 (GH, MEXU, TENN, US); side road between Orizaba and Fortin, 3200 ft, 23 May 1973 Terrell & King 4450, (US); Tepejilotla, mpio. Chocaman, 29 Jun 1979, Ventura A. 16282 (ARIZ, CHAPA, ENCB, MO). GUATEMALA. Huehuetenango: 5 mi SE of Barillas, Sierra de los Cuchumatanes, 1150 m, Steyermark 49537 (GH)
3. Mexotis kingii (Terrell) Terrell & H. Rob., comb. nov. (Figs. 2, 3). Houstonia kingii Terrell, Brittonia 32:491. 1980. Hedyotis kingii (Terrell) G.L. Nesom, Syst. Bot. 13:434. 1988. Tre: MEXICO. Oaxaca: E-facing slopes, mountains along rt. 175, 28 km by road S of Valle Nacional on way to Ixtlan de Juarez; rain forest abounding in ferns, bromeliads, orchids, and aroids; elev. ca. 2000 m; not common; full sun; gravelly soil, 30 Jul 1959, R.M. King 2107 (HoLorvre: MICHI, 2 sheets, one flowering, one fruiting).
Perennial herbs with thick, woody roots. Stems to 45 cm tall, to ca. 5 mm thick and somewhat woody toward base, tan or straw-colored, subterete, rather stout, erect, glabrous, branched. Leaves with petioles to 10 mm long, blades 40-85 x 8-30 mm, elliptic, ovate, or lanceolate-ovate, acuminate, glabrous, slightly paler be- neath. Stipules to 2 mm long, broadly rounded, lobed, the central lobe with recurved glandular hairs. In- florescence in terminal, few-flowered cymes, widely branching, to 8 x 11 cm, flowers heterostylous, pedicels to 13 mm long, very slender or filiform. Hypanthium glabrous; calyx lobes 1.5-4.7 x 0.3-0.8 mm, 1/2-3/4 as long as corolla tube, linear or narrowly lanceolate, acute or acicular, sometimes divaricately spreading. Corollas 7-13 mm long, broadly funnelform, white or light purple, glabrous externally; tube 3-9 mm, 3-7 mm wide at throat, glabrous or puberulent within; lobes 2.5-7.0 x 1.5-3.3 mm wide, shorter than to slightly longer than the tube, ovate, glabrous or puberulent within. Pin flowers with stigma lobes 0.5—1.3 mm long, narrowly oblong or linear, near corolla throat or slightly exserted, anthers located at 1/3-1/4-point from base of corolla tube. Thrum flowers with anthers 1.3-2.0 mm long, narrowly oblong, purple, attached at corolla throat, subsessile or filaments to 1.5 mm long, stigmas located at 1/4—point from base to midpoint of corolla tube. Capsules (only previous year's dehisced capsules seen) 3-4 x 3-4 mm, subglobose, thin- walled, glabrous, 3/4—7/8 inferior. Seeds 0.6—1.1 x 0.5-0.9 mm, dark brown, moderately to slightly com- pressed, often polygonal in outline, obtusely angulate or obtusely ridged, margins rounded, hilum puncti- form, testa finely reticulate, areoles small, isodiametric. Pollen illustrated in Terrell et al. 1986, figs 21, 22. Plant and seeds illustrated here in Figures 2, 3. Chromosome number unknown.
Phenology.—Flowering December to July.
Distribution.—México: Oaxaca. Original collection in a cloud forest of the Sierra Juarez range along or near highway 175 from Ixtlán de Juarez to Valle Nacional, 28 km south of Valle Nacional. Additional occur- rences in Oaxaca cited below are mostly along hwy. 175 between Tuxtepec and Oaxaca city; the location of Tarabunde is unknown. On 22 July 1987 Stephen Koch and Terrell, somewhat hampered by a torrential rain, searched unsuccessfully for the species along Highway 175.
Additional specimens examined. MEXICO. Oaxaca: Distr. de Ixtlan, roadside, trop. v itl ferns, 24 mi S of Valle Nacio- nal, Hwy. 175, 22 Mar 1978, Poole et al. 1283 (LL); bosque mesofilo perturbado, 1750 m, 38 km S of Valle Nacional, 11 Apr 1976, Rze- dowshi 34092 (ENCB); km 128 entre Llano de Las lores y Tuxtepec, 26 Dec 1965, Delgadillo 121 (MEXU), reported by Lorence; Mpio. de Comaltepec, Hwy. 175, “Vista hermosa,” selva secundaria, km 137, 1500 m, 11 May 1966, Martinez Calderon 801 (ENCB, MEXU, XAL); Tarabunde, 7 Feb 1966, MacDougal 17 (US); 22-26 mi S from Valle Nacional on Hwy. 175, cloud forest, 7 Jan 1989, Todzia et al. 2839 (TEX)
4. Mexotis terrellii (Lorence) Terrell & H. Rob., comb. nov. Hedyotis terrellii Lorence, Novon 7:51.1997. Tyre: MÉXICO. Oaxaca: Distrito de Mixe, Municipio de Totontepec, 3 km al SO de Totontepec, carretera a Mitla, 1910 m, bosque mesófilo, 6 Sep 1986, R. Torres C. & C. Martinez 9067 (HoLorvre: PTBG 12314; isotype: MEXU)
The following description is modified from that of David Lorence (1997). Perennial herbs. Stems 45-80 cm
tall, branching from the base with several erect stems 2.0-3.5 mm diam. and somewhat woody toward base,
terete, pee adi with petioles 3. on mm long, blades (25—)45-100 x 10-40 mm, elliptic to ovate-
elliptic, the b decurrent, te, glabrous, chartaceous, margins sometimes with few small
teeth. Stipule en to 4 mm ine: MR or deltate, with apical and lateral linear gland-tipped teeth 1-3
, Mexotis, g f Rubi 65
Fe? € de afhalat £ M. n ae A Joy CEA) A R Seeds polygonal, flat, IE ele PIPER] I CD A E el I l,
not radially elongated.
mm long. Inflorescences terminal, compound cymes 3-15 x 6-18 cm, subtended by reduced leaves, branch- ing dichasially to 2° or 3°, the primary branches 2-5 cm long, the axes slender, glabrous, subtended by stipule-like bracts, flowers on filiform pedicels 7-32 mm long. Hypanthium broadly obconic, glabrous. Calyx lobes 1-2 x 1-2 mm, deltate, glabrous. Corolla in bud quadrangular, rounded at apex, at anthesis 5-6.5 mm long; tube 2-2.5 mm long glabrous, lobes 3-4 mm long, erect or slightly spreading, elliptic, acute at apex, externally glabrous, internally villosulous with crinkled, white trichomes. Stamens attached near middle of tube below sinuses, the filaments 0.5 mm long, anthers 1 mm long, linear-elliptic, style 2-2.5 mm. Stigma lobes 1 mm long, linear. Capsules 2-3 x 3-4 mm, slightly wider than long, 1/2 inferior, thin- walled. Seeds immature. Plant illustrated in Lorence 1997.
Phenology.—Collected in flower in March, and in flower and fruit in June and September.
Distribution. —Known only from the area around the type locality in cloud forest at ca. 1900 m. near Totontepec in the mountainous Mixe District of Oaxaca.
Notes.—Lorence (1997) compared certain characters of M. terrellii with those of M. galeottii and M. kingii. Mexotis terrellii mede resembles M. hingii, as summarized in Table 1
d. P MEXICO. Oaxaca: Distrito Ixtlán, Llano Verde, collector unknown, Herb. Reichenbach fils
III-396(W); Dio Mixe, mpio. ded COE Totontepec, bosque mesófilo de montaña, 1900 m, 17?15'N, 9602"W, 11 Mar 1990, Rivera R. &
66 Journal of tl
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Fic. 3. Mexotis kingii, holotype. A. Habit. B. Flower. C. Corolla with st style, and stigma D Ovary sectioned, ith calyx lol