Journal of the Botanical Research Institute of Texas

J. Bot Res. Inst Texas ISSN 1934-5259

1971— William F. Mahler (right), professor of

botany at SMU and director emeritus of BRU,

inherited editorship and copyright.

1993— BRIT becomes publisher/copyright holder.

2007— First issue of 7. Bot. Res. Inst. Texas.

preparation, manufacture, and distribution

of botanical research and scientific discoveries

for the twenty-first century.

VOLUME 7 NUMBER 2 10 DEC 2013

Botanical Research Institute of Texas (BRIT)

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F6lix Llamas (Universidad of Le6n, Le6n, Spain)

Harold Keller (BRIT)

Robert J. O'Kennon (BRIT)

Richard Rabeler (MICH)

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science for plant conservation and education.

Direction and Coverage

The BRIT Press considers original research papers

concerned with classical and modern systematic

y, sensu lato, for publication in 1 Bot. Res. Inst. Texas.

e available

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n the BRIT Press website, httpy/www.britpress.org.

Bibliographical

Citation abbreviation for the

by the Botanical Research Institute ofTexa

Cover Illustration

Electronically tinted botanical illus

of Liatris aestivalis originally used o

BRIT'S anniversary poster 2001.

Jul-Aug(-Sep) and is endemic t<

Sida 19:768.2001.

Botanical illustration by

Press

Table of Contents

DEDICATION

In memory of BRIT co-founder William F. “Bill” Mahler (1930-2013) 615

SYSTEMATICS

Four new annual species of Euphorbia section Tithymalus (Euphorbiaceae) from North America

Mark H. Mayfield 533

Hackelia taylori (Boraginaceae), a new species from north central Washington state (U.S.A.)

Richy J. Harrod, Lauri A. Malmquist, and Robert L. Carr 649

Una nueva especie de Sobralia (Orchidaceae) de El Salvador

Jose L. Linares 659

a-MArquez, Oscar H. MarIn-GOmez, and John L. Clark

New records and notes on species from Parc National Pic Macaya, Massif de la Hotte, Haiti,

including a new species of Pilea (Urticaceae)

Lucas C. Majure, Gretchen M. Ionta,J. Dan Skean, Jr., and Walter S. Judd

Chusquea darkiae (Poaceae: Bambusoideae: Bambuseae: Chusqueinae): a new species in

section Longiprophyllae from Narifio, Colombia

Emmet J. Judziewicz and Ximena Londono

The spikelet callus of Eriochloa villosa (Poaceae)

Stephen J. Darbyshire, MARiE-JoseE Simard, and Robert E. Nurse

Changes to Potentilla rubricaulis s.L, P. hookeriana (Rosaceae), and erstwhile synonyms

in Flora of North America north of Mexico

Barbara Ertter, Reidar Elven, and David F. Murray

Previously unrecognized types of American Acacia species from the Torino Herbarium (TO)

David Seigler, Laura Guglielmone, and John Ebinger

s (Apocynaceae: Asclepiadoideae)

667

681

693

697

703

713

743

Micropetasos, a new genus of angiosperms from mid-Cretaceous Burmese amber

George O. Poinar, Jr., Kenton L. Chambers, and Joerg Wunderlich

BOTANICAL HISTORY

FLORISTICS, ECOLOGY, AND CONSERVATION

Alpine flora of Cerro Quiexobra, Oaxaca, Mexico

J. Andrew McDonald

Missouri botanical

DEC 2 7 2013

GARDEN LIBRARY

745

751

765

Flora and phytogeography of Cumbres de Monterrey National Park, Nuevo Leon, Mexico

Eduardo Estrada-CastillOn, JosE Angel Villarreal-Quintaniua,

Maria Magdalena Saunas-RodrIguez, Humberto RodrIguez-GonzAlez,

Javier JimEnez-PErez, and Mario Alberto GarcIa-Aranda

Flora and phytogeography of the Canon de Iturbide, Nuevo Leon, Mexico

MarIa Magdalena Salinas-RodrIguez, Eduardo Estrada-CastillOn, and

JosE Angel Villarreal-Quintanilla

Nuevos registros de Cactaceae y Solanaceae para el estado de Guanajuato, Mexico

Maricela GOmez-SAnchez, B.A. GonzAlez-HernAndez, Mahinda Martinez y

Luis G. HernAndez-Sandoval

para la flora de El Salvador

Frank Sullyvan Cardoza Ruiz y JosE L. Linares

Rediscovery of Callirhoe papaver (Malvaceae) in Alabama (U.S.A.)

Brian R. Keener and LJ. Davenport

Rediscovery of Persea borbonia var. borbonia (Lauraceae), Prosopis glandulosa var.

glandulosa (Fabaceae), and Pinus palustris (Pinaceae) in Arkansas, with three new

angiosperm species for Arkansas (U.S.A.)

Brett E. Serviss and James H. Peck

A floristic inventory of Phillips and Valley counties, Montana (U.SA)

Joseph L.M. Charboneau, B.E. Nelson, and Ronald L. Hartman

Taxonomic identity and historic

conservation concern in the Gre

James H. Locklear

Range expansion of Panicu

endangered species

Jeffrey T. Hutchinson and Robert B. Shaw

Annouud checklist of the vascular Dora of the Wind River Range, Wyoming (U.S.A.)

W F ' Fectig ' Ro,H,r T - M«sam, B.E. Neison, and Ronald L. Hatoian

Book Reviews, Notices, and Announccnn

820, 826, 840, 846, 900, 940

Reviewers for Volume 7 (2013)— 941

Index to Volume 7 (2013)— 942

Titles of Articles with Authors— 442

Authors— 944

Botanical Names and Subjects— 445

NewNan

'• 632 > 648, 658, 666, 680, 692, 712, 770, 8(

771

821

841

847

879

905

INDEX to new names and new combinations inj. Bot. Res. Inst Texas 7(2), 2013

Chusquea darkiae Londono & Judz., sp. nov. — 693

Columnea ceticeps J.L. Clark & J.F. Smith, sp. nov. — 668

Columnea ferruginea J .F. Smith & J.L. Clark, sp. nov.— 671

Columnea fractiflexa J.F. Smith &J.L. Clark, sp. nov— 673

Columnea laciniata J.L. Clark & M. Amaya, sp. nov. — 676

Euphorbia austrotexana M.H. Mayfield, sp. nov. — 634

Euphorbia austrotexana var. carrii M.H. Mayfield, var. nov— 636

Euphorbia georgiana M.H. Mayfield, sp. nov. — 639

Euphorbia nesomii M.H. Mayfield, sp. nov. — 639

Euphorbia ouachitana M.H. Mayfield, sp. nov. — 642

Hackelia taylori Harrod, Malmquist & Carr, sp. nov. — 652

Micropetasos G.O. Poinar, K.L. Chambers & J. Wunderlich, gen. nov— 746

j.O. Poinar, K.L. Chambers & J. Wunderlich, sp. i

Pilea vermicularis Majure, Skean & Judd, sp. nov.— 688

Sobralia paulancalmoi J. Linares, sp. nov. — 659

IN MEMORY OF BRIT CO-FOUNDER

WM. F. “BILL” MAHLER

1930-2013

Valued Director, Mentor, Visionary, Editor, Teacher, Botanist, Supporter, Friend

With indebtedness for your many contributions and service to our organization.

The Botanical Research Institute of Texas

WILLIAM FRED MAHLER

“BILL”

30 AUGUST 1930-2 JULY 2013

hometown ai^were married in 1955 years of age ' Bil1 in the back his brotber

In 1958 he went to Oklahoma Johnmthefront

State University (OSU) in Stillwater to pursue graduate work. Mahler re-

ceived his M.S. degree in Botany/Plant Taxonomy from OSU in 1960,

working under U.T. Waterfall. For the next six years he served as an assis-

teaching botany and establishing the HSU herbarium. Subsequently he

continued his graduate studies by attending the University of Tennessee at

Knoxville where he received a Ph.D. in Botany/Plant Taxonomy in 1968.

Upon graduation he joined the faculty of Southern Methodist University

(SMU) in Dallas, became editor and publisher of Sida, Contributions to

Botany in 1971 following the death of L.H. Shinners, and assumed leader-

ship of the SMU herbarium in 1973. Mahler was publisher of Sida, Botanical

js visible fromthe Miscellany after he and Barney Lipscomb founded the journal in 1987. Under

that copy after a buddy his guidance and own collecting, the SMU herbarium grew by 72,000

and ask for it.' specimens, eventually reaching about 400,000.

616

clarity

ande^7r b ™' da T S ’K M ^° /rttNMhC “^ T ““^a W ' W88 ).»eUl

. F. Mahler: In memory of BRIT co-founder

617

native flora of Texas. Other notable publications included the Keys to the Plants of Black Gap Wildlife Manage-

ment Area, Brewster County, Texas (1971), Flora of Taylor County, Texas (1973), and The Mosses of Texas (1980).

Mahler’s specialties include Fabaceae, Baccharis (Asteraceae), mosses, floristics, pollen morphology, and the

study of endangered plant species. In 1988, Mahler was the first recipient of the Harold Beaty Award from the

Texas Organization of Endangered Species for his work with endangered plant species in Texas. The Native

Plant Society of Texas again honored Mahler in 1995 with the Charles Leonard Weddle Memorial Award in

recognition of a lifetime of service and devotion to Texas native plants. Mahler also served on the Board of

Consultants for the North Texas Poison Center, Parkland Hospital, Dallas, Texas. He assisted the Poison Cen-

ter in identifying plants and mushrooms implicated mostly in human poisoning cases.

In 1987 SMU put its herbarium on permanent loan to a newly created organization. The Botanical Re-

search Institute of Texas (BRIT). Mahler received early retirement from SMU (Associate Professor Emeritus )

and served as the first Director of BRIT (1987-1992). Along with Andrea McFadden and long-time associate

Barney Lipscomb, they were instrumental in its establishment as a free-standing research institution.

In 1993, S.H. Sohmer assumed directorship of BRIT, and Bill served as Director Emeritus (1993-2013).

After retiring, he returned with his loving wife to his childhood home of Iowa Park, Texas, where he enjoyed

and shared life experiences with his many friends, family, and grandchildren. He was a native Iowa Parkan and

proud of his home town. There he kept his fingers in botany, attended many BRIT functions in Fort Worth

(often with Iowa Park friends, introducing them to BRIT), and worked tirelessly on the genealogy of the Mahler

family. The taxonomist in him never retired. On July 2, 2013, Bill retired to his final resting place adjacent to

his father, mother, and brother in Highland Cemetery, Iowa Park, Texas.

ournal of the Botanical Research Institute of Texas 7(2)

REFERENCES

Black, R.W. 1989. Rangers in Korea. Roster of 14th Co., P. 318. Ivy Books, Random House, New York, NY, U.SA

Diggs, G.M., Jr., B.L. Lipscomb, and R J. O'Kennon. 1 999. Shinners and Mahler's illustrated flora of North Central Texas. Sida,

Bot. Misc. 1 6. Botanical Research Institute of Texas, Fort Worth and Austin College, Sherman, Texas, U.S.A.

Taylor, T.H. n.d. Rangers lead the way. Roster, P. 1 53. Turner Publ. Co., Paducah, KY, U.S.A.

EDUCATION:

H.S., Wm. F. George High School, Iowa Park, Texas, May

1947

B.S. (Agriculture), Midwestern State University, Wichita

Falls, Texas, January 1955

M.S. (Botany, Plant Taxonomy), Oklahoma State Univer-

sity, Stillwater, August 1960

Ph.D. (Botany, Plant Taxonomy), University of Tennes-

see, Knoxville, August 1968

NSF SUMMER PROGRAMS:

1961 4th Summer Institute for College Teachers of

Botany, Washington State University, Pullman,

Washington.

1962 Marine Biology course. University of Oregon

(Biology Station, Charleston, Oregon).

19 63 Desert Biology course, Arizona State University,

of Bior | taJ ahler ' 1987 ' the SMU Her,,aliu,n • Associate Professor Tempe.

m Methodist University, Dallas, Texas. 1965, 1966 Research Participation Program for College

Teachers, Iowa State University, Ames.

GRADUATE ASSISTANTSHIPS:

Graduate Teaching Assistantship, Botany Department, Oklahoma State University, 1958-1960.

Knox^k^^^iges 131 Bl ° l0Sy (Research) ’ Department of Agricultural Biology, University of Tennessee,

ACADEMIC POSITIONS:

Hardin S.mmons University, Abilene, Texas; Assistant Professor, 1960-1969

Somhem MaWis, University, Dallas, Texas; Curator, SMU Herbarium 1971-1987; Assistant Professor,

1968-1974; Associate Professor, 1974-1987.

SUMMER TEACHING POSITIONS:

^ SMU “ d DaU “ Muscum °l Natural History,

Burgwin Research Center, Ranchos de Taos, New Mexico: SMU Departmental Courses, 1974-1977, 1979.

NON-ACADEMIC POSITIONS:

Botanical Research Institute of Texas, Fort Worth; Director 1987-1993; Director Emeritus, 1993-2013

MILITARY EXPERIENCE:

Germany^ 1950-1953 (14th Airbo ™ Ra "g« Infantry Company, 4th Infantry Division), U.S A. and

ORGANIZATIONAL MEMBERSHIPS

A#mc. Curator, Dallas Museum of NaturalHis, my

^o ^Natuml Area Preservation Assmiation

Botanical Research Institute of Texas

Wm.FJ

Chihuahuan Desert Institute

Consultant, North Texas Poison Center, Parkland Hospital, Dallas, Texas

Texas Plant Recovery Team, U.S. Fish and Wildlife Service

SOCIETIES:

American Association of Plant Taxonomists

American Bryological and Lichenological Society

Fort Worth Botanical Society

Heard Museum

International Association of Plant Taxonomists

Native Plant Society of Texas

North American Mycological Association

Southeastern Pecan Growers Association

Southwestern Association of Naturalists

Texas Committee on Natural Resources

Texas Mycological Society

Texas Nature Conservancy

Texas Organization of Endangered Species

HONORS:

Harold Beaty Award: for outstanding achievement in Endangered Plant Conservation, 1st recipient, Texas Or-

ganization of Endangered Species, 1988.

Donovan Stewart Correll Memorial Award: for scientific writing on the native flora of Texas, Native Plant Soci-

ety of Texas, Kerrville, TX, 1991.

Charles Leonard Weddle Memorial Award: in recognition of a lifetime of service and devotion to Texas native

plants, Native Plant Society of Texas, Waco, TX, 1995.

STUDY LEAVE (SABBATICAL):

Fall 1976, Mosses of Texas

Fall 1986, Mycological protocol for North Texas Poison Center, Parkland Hospital

PUBLICATIONS:

Books and Manuals:

1964 General Botany Laboratory Manual. By author. 51 pp.

1966 Keys to the Embryophyta of Taylor County, Texas. Hardin-Simmons University Bookstore, Abilene,

Texas, 86 pp.

1971 Keys to the Vascular Plants of the Black Gap Wildlife Management Area, Brewster County, Texas. SMU

Bookstore, Dallas , Texas. Third revision, 109 pp.

1972 Shinners’ Spring Flora of the Dallas-Fort Worth Area, Texas. Editor, 2nd edition. Prestige Press, Fort

Worth, Texas, 514 pp. (Class use: Tarleton State University, Tyler Junior CoUege).

1972 Keys to the Mosses of Texas. Biology Department, SMU (for class use), 43 pp.

1973 Flora of Taylor County, Texas: A Manual of the Vascular Plants with Selected Sketches. Published by the

author, SMU Bookstore, Dallas, Texas, 247 pp.

1975 Keys to the Bryophytes of Texas. SMU Herbarium, Dallas, Texas, 64 pp. (Class use: SMU, Texas A&M

University, Angelo State University).

1980 The Mosses of Texas: A manual of the moss flora with sketches. Published by the author, Dallas, Texas,

147 pp. (Class use: Texas A&M University, Angelo State University).

1984 Shinners’ Manual of the North Central Texas Flora, SMU Herbarium, Dallas, Texas, 360 pp. (Class

use: Austin College, Sherman; Baylor University; Fort Worth Botanical Garden; Fort Worth Nature

Center; Greenhills Environmental Center; Southeastern Oklahoma State University; University of

Texas-Arlington).

620

1988 Shinners’ Manual of the North Central Texas Flora. £

i, Bot. Misc. 3:1-313. Botanical Research Institute

of Payne County, Oklahoma. Bryologist

, and New Mexico. Southw.

arch Participation

, R.C. 1977. Dalea

Ikenberry, G.J., C.D. Bird, and W.F. Mahler.

63:112-113.

Mahler, W.F. and U.T. Waterfall. 1964. Baccharis (Compositae) in Oklahoma, T<

Naturalist 9:189-202.

Mahler, W.F. 1965. The pollen morphology of the tribe Psoraleae (Leguminosae). NSF Re

Report. 24 pp. Unpublished copy at New York Botanical Garden. Cited by Bamel

Imagines. Mem. New York Bota. Gard. 27:1-891.

Mahler, W.F. 1966. The pollen morphology of the subfamily Papilionoideae (Leguminosae). NSF Research

Participation Report. 21 pp. Unpublished copy at New York Botanical Garden.

Mahler, W.F. 1969. Pollen morphology of Desmodium (Leguminosae) in the United States. XI International

Mahler, W.F. 1970. Manual of the legumes of Tennessee. Tennessee Acad. Sci. 45(3):69-96.

Mahler, W.F. 1970. Pollen morphology of Dalea mollis - D. neomexicana complex (Leguminosae). Southw.

Naturalist 15:187-191.

Mahler, W.F. 1971. Lloyd Herbert Shinners 1918-1971. Sida 4:228-231.

Mahler, W.F. 1973. By any other name... Sida 5:180-181.

Mahler, W.F. 1973. Book review: The Agavaceae of Sonora by H.S. Gentry. Econ. Bot. 27:156.

Mahler, ^WF. 1974. Gnaphalium helleri Britton (Compositae - Inuleae) in the Texas flora. Southw. Naturalist

Mahler, W.F. 1975. The distribution of Pyropappus rothrockii Gray (Compositae - Cichorieae) in Texas. Southw.

Naturalist 20:139.

“ <1 diaribUti0n ° f ‘ he Varie,ieS ° f heUeri Britton (Compositae -

J976. Book review: The grasses of Texas by F.W. Gould. Southw. Rev. Spring: 220-222.

Mahl ?’ w F Q _^°| r en mor P hoI °gy Of Dalea section Theodora (Leguminosae - Psoraleae). Sida 6:328-331.

5(4)"41-42 Fe3tUred Institution ~ Southern Methodist University. Assoc. Syst. Coll. Newsl.

Mahler WF 1979 checklist ofNew Mexico mosses. Bryologist 81:593-599.

anler, W.F. 1979. Range extension of Brazoria pulchemma Lundell (Lamiaceae). Sida 8:211.

>. Rubus

r. duplaris (Shinners) Mahler,

/. (Rosaceae). Sida

Mahler, W.F. ]

Mahler,

8:211-212.

Mahler WF 1980 ' Checklist of mosses of Oklahoma. Bryologist 83:202-208.

Mahler W F 1981 ‘ P °u 16X35 3nd 0klahoma P lants - Sida 9:76-86.

M 1 1 ’ ' ' 8L Fledstudles on Texas endemics. Sida 9:176-186.

States Canada and r° mb l ^ A Synonymized checklist of the vascular flora of the Unite

Olwell M ^ T land - V ° l ^ The Bl0ta ° f North America. Sida 9:191-192.

etr in 1 rl' v 3 ^ l " 2 A populational stud y of the exomorphic variations in Vida minutiflora D

Mahler WfZZI r ™^* onii Wats - (Leguminosae). Sida 9:215-222.

Mahler’ W F 1983 ReZZ T * “ ° f ^ California > Mexico b Y EW. Gould and R. Moran. Sida 10:92-9:

Mahler] W mu “T and ” 0,es on tw0 other endemics. Sida 10:89-92

Beaty, H.E. and W.F. Mahler 1983 TOeTZ “ TT'™ ^ 10191

Organization ofEndlV Z endan 8 ered . threatened, and watch list of plants of Texas. Texa

Mahl w ot Endangered Species. Publ. 3, 1st rev. 7 pp.

Dallas, tS^ m3nUal N ° rth Central Texas flora - Southern Methodist Univ. Herbariun

r: In memory of BRIT c

Mahler, W.F. 1985. Review: Methods in plant virology by S. A. Hill. In: Methods in Plant Pathology 1, 1984. Sida

11:105.

Mahler, W.F. 1985. Review: Introduction to modern mycology by J.W. Deacon. In: Basic Microbiology Ser.

Vol.7, 1984. Sida 11:106.

Mahler, W.F. 1985. Additions to the moss flora of New Mexico. Evansia 2(2):30-31.

Coats, C. and W.F. Mahler. 1985. Barbula whitehouseae Crum, new to Oklahoma and distribution notes.

Southw. Naturalist 30:151-152.

Mahler, W.F. 1986. Review: A field guide to southwestern and Texas wildflowers. Quart. Rev. Biol.

61:107-108.

Mahler, W.F. 1986. Review: Plant diseases: Infection, damage and loss by R.K.S. Wood and G.J. Jellis eds. 1984.

Sida 11:353-354.

Mahler, W.F. 1986. Review: The vascular plants of South Dakota, second edition by T. VanBruggen, 1985. Sida

11:354-355.

Grauke, L.J., J.W. Pratt, W.F. Mahler, and A.D. Ajayi. 1986. Proposal to conserve the name of pecan as Carya

illinoensis (Wang.) K. Koch and reject the orthographic variant Carya illinoinensis (Wang.) K. Koch (Jug-

landaceae). Taxon 35:174-177.

Beaty, H.E. and W.F. Mahler. 1987. Endangered, threatened, and watch list of plants in Texas. Texas Organiza-

tion of Endangered Species. Publ. 3, 2nd rev. 11 pp.

Mahler, W.F. 1987. Leavenworthia texana (Brassicaceae), a new species from Texas. Sida 12:239-242.

Mahler, W.F. 1987. New combination and notes on the North Central Texas flora. Sida 12:250-251.

Mahler, W.F. 1988. Shinners’ manual of the North Central Texas flora. Sida, Bot.Misc. 3:1-313.

Mahler, W.F. 1988. Amorpha roemeriana Scheele (Fabaceae), an upland species. Sida 13:121.

Mahler, W.F. 1988. Dr. Russell Lee Kologiski. Sida 13:170.

Mahler, W.F. 1989. Agrimonia incisa (Rosaceae) new to Texas. Sida 13:383.

Bastien, J.W, W.F. Mahler, M.G. Reinecke, W.E. Robinson, Jr, J. Zalles, and Y. Shu. 1990. Testing of anti-HIV

compounds from Bolivian-Kallawaya medicinal plants. Paper presented at 2nd International Congress

of Ethnobiology, October 22-28, Kunming, China, by J.W. Bastien.

Bastien, J.W, W.F. Mahler, M.G. Reinecke, Y. Shu, W.E. Robinson, Jr, L. Horton, and J. Zalles-Asin. 1991. A

virgin pharmacopeia: Bolivian-Kallawayan medicinal plants as sources of anti-HIV compounds. Ab-

stracts of the International Congress of Natural Products 32, Abstract P-3.

Abdel-Malek, S, J.W. Bastien, W.F. Mahler, M.G. Reinecke, W.E. Robinson, Jr, andj. Zalles-Asin. 1994. Phyto-

chemical studies of Kallawaya medicinal plants potential anti-HIV Drugs. Abstracts Southwest Regional

Meeting of the American Chemical Society 50, Abstract No. 165.

Abdel-Malek, S, J.W. Bastien, W.F. Mahler, Qi Jia, M.G. Reinecke, W.E. Robinson, Jr, Yong-hua Shu, andj.

Zalles-Asin. 1996. Drug leads from the Kallawaya herbalists of Bolivia. 1. Background, rationale, proto-

col and anti-HIV activity. J. Ethnopharmacology 50:157-166.

NEW TAXA AND NEW COMBINATIONS

Asteraceae Gnaphalium helleri Britton var. micradenium (Weath.) Mahler— Sida 6(1):32. 1975.

Rosaceae Rubus trivialis Michx. var. duplaris (Shinners) Mahler— Sida 8(1): 211. 1979.

Brassicaceae Leavenworthia texana Mahler— Sida 12(1):239. 1987.

Fabaceae Pediomelum hypogaeum Rydb. var. scaposum (A. Gray) Mahler— Sida 12(1): 250. 1987 (IK)

Fabaceae Pediomelum latestipulatum (Shinners) Mahler— Sida 12(1): 250. 1987.

GRADUATE STUDENTS - M.S.

Olwell, Margaret. 1983. Geobotanical study of Penstemon cyanocaulis Payson in Lisbon Valley, Utah. Thesis.

December.

Ajayi, Anthonia. 1986. Phaseoleae (Leguminosae) of Oklahoma. Thesis. May.

PERSONAL TRIBUTES TO

WILLIAM F. MAHLER

Bill Mahler was a new faculty member when I arrived on the SMU campus in 1968. His appearance in faculty

meetings was always an occasion to visit with a valuable member of the faculty who was not located in Fondren

Science building but in the basement of the Science & Engineering Library. I recall a joint field trip to the Big

Bend region of Texas with Bill and several staff members of the Dallas Museum of Natural History, and we were

all taken back by Bill’s great knowledge of the plants there. We had no idea he was writing a monograph on

them. Bill was always a friendly person, eager to share his knowledge of plants.

—John Ubelaker, Biology Department , Southern Methodist University, Dallas

Remembrance for Dr. M. — Dr. Mahler was my graduate adviser at Southern Methodist University in Dallas,

Texas, over 35 years ago. I met Dr. Mahler in 1977 when I had just moved to Dallas after getting my under-

graduate degree in Botany at University of North Carolina, Chapel Hill. I wanted to talk with him about the

graduate program in Botany at SMU. He suggested that I wait one year before coming to graduate school be-

cause SMU was hiring 2 more professors with botanical/ecological expertise, and he thought there would be a

more well-rounded program for me. As usual, he always thought about what was best for the student. He was

absolutely correct, I waited one year and started my classes in 1978 with Dr. Mahler as my graduate adviser. He

taught me much about plants, mosses and lichens, pollen, etc., but the thing he taught best was humility. He

guided you gently, he listened intently (while squatting), and he never made you feel as though any question

you asked was dumb, even if it was! He opened many doors for me professionally, and 1 am forever grateful. I

am sure that the successes 1 have had as a botanist throughout my career are because of his unwavering belief

in my abilities and me.

Dr. Mahler loved plants, especially the flora of North Texas! He understood at a very deep level how im-

portant plants are for people, how important the SMU Herbarium was for botanical science, and how impor-

tant it was to teach people about plants and their significance for humanity. The Botanical Research Institute of

Texas is his legacy! I think Texas has lost a true Texan. He was, without a doubt, the best major professor a

student could have! He was a gifted scholar, a remarkable teacher, and most of all a truly wonderful human

being. It is with great fondness that I remember the venerable Dr. Mahler from Iowa Park. Dr. Mahler, you were

my favorite professor of all time and you will be sorely missed!

—Peggy Olwell, Washington, DC

William F. Mahler .— Dedicated educator, passionate, wonderfully sweet man. Dr. Mahler opened up a whole

botanical world to his students which we never considered. After taking an elective class in Botany, Dr. Mahler

introduced me to the world of mosses and fungi. We traveled all over north Texas and Oklahoma in search of

mosses. He worked tirelessly to preserve the SMU Herbarium collection for generations of students. I was one

of those many fortunate students blessed by his knowledge and passion for the plant life we studied.

The plant world knows Dr. Mahler as a taxonomist and botanist who was passionate about Texas flora,

the SMU Herbarium, and BRIT. But Dr. Mahler was more than an outstanding botanist. He was a good friend,

mentor, and surrogate dad. He always had time for his students, and unlike many professors who maintain an

academic aloofness from their students. Dr. Mahler was a warm and caring friend who was concerned about

his students’ welfare not only in the classroom but also outside of the classroom. Dr. Mahler was a man of tre-

mendous character, singularly committed to his work, his family, the Herbarium, and to his home town of

Iowa Park. I fondly remember Dr. Mahler quietly encouraging and always positive with pipe in hand. He was

the epitome of integrity, honesty, and curiosity.

Someone once wrote, “A friend should be one in whose understanding and virtue we can equally confide, and

whose opinion we can value at once for its justness and its sincerity.” Dr. Mahler was just such a friend. I greatly

valued his suggestions as he provided me invaluable advice particularly when I was making a career choice.

Although I did not pursue a flora-related career, he supported me as I chose a law career. I, like many of his

Wm. F. Mahler: In memory of BRIT co-founder

625

grabbed. We now knew that the plant did not grow so dose to the creek banks as did Amorpha fruticosa, but

high on the banks above the creeks. Knowing this, we eventually found two more sites.

1 worked with Bill and Barney Lipscomb at SMU until the new Botanical Research Institute of Texas was

formed in 1987. When we moved to the new location in Fort Worth in 1991, Bill and I supervised the move of

the herbarium cases and library books from the herbarium at SMU, and Barney and Andrea supervised the

unloading of the trucks at the new building in Fort Worth. Bill stayed on at BRIT for a number of years until

Bill Mahler was an incredible individual, very intelligent, focused, professional, the consummate gentle-

man, and admired friend to all. I will never forget the time I spent with him.

— Robert J. O’Kennon, Fort Worth

Bill Mahler: A Man to Remember! — One of the reasons 1 hate getting older is losing friends and people I have

known. Bill Mahler was one of those people 1 hated losing, although I had not seen much of him in the couple

or three years before he passed. My lasting impression of Bill will always be when 1 came to BRIT for the first

time in 1992 to interview for the director’s position that Bill was stepping down from. I found Bill, Barney

Lipscomb, and a couple of other people in a 10,000-square-foot warehouse in what was then “outer Siberia” of

downtown Fort Worth. He was a solid, tough guy, with a curmudgeonly and stubborn streak in him, but some-

one who knew who he was. There are not many people who can qualify as an Army Ranger, but he was one of

those. He always seemed to have a cigarette dangling from his Ups and used “F.Y.I." (“For Your Information”) a

lot. Since I also smoked in those days (but gave it up after accepting the director’s position of BRIT), he and I

would occasionally hang out on the second floor outside stairwell sharing a smoke. What was also startling

was that when I saw him at BRIT, I realized that this was the same person I had met when 1 started graduate

work at the University of Tennessee in Knoxville: he was leaving with a fresh Ph.D. and had been an older stu-

dent having gone back to school after doing his time in Korea. I am certain that the second floor outside stair-

well on Pecan Street in downtown Fort Worth, in BRIT’s original building, will always have BiU’s spirit there

that will last much longer than the cigarette butts left behind during his time in that building. Farewell to you,

good friend. I will always remember you fondly, and from all of us at BRIT, thank you!

— S.H. Sohmer, President, Botanical Research Institute of Texas, Fort Worth

Bill Mahler Leaves a Great Legacy. — Bill helped us create a truly significant and lasting institution. We could not

have done what we did with our other board colleagues if it weren’t for him. Just think of what a big move it was

for Bill to go from tenure at SMU to a warehouse in downtown Fort Worth under the auspices of a bunch of

neophytes! He leaves a great legacy [Botanical Research Institute of Texas].

—Ed Bass, Fort Worth

Dr. William F. Mahler: My Friend and Colleague —

1975-1981 (The Formative Years of My Career)

Mr. Robert Ziegler and Dr. Mickey Cooper were professors at Cameron University, in Lawton, Oklahoma, who

stimulated my interest in botany and, more specifically, taxonomy of aquatic plants, under Dr. Cooper. In 1973

I was accepted into graduate school in the botany department at the University of Arkansas in Fayetteville.

Once on campus, I met Dr. Edwin B. Smith, the taxonomist and discovered the herbarium. In the spring of

1975, 1 was finishing my master’s thesis and was on my way to becoming a botanist. After graduation, I won-

dered where I would get a job or what path I would follow. Under the direction of my adviser at the University

of Arkansas, Dr. Edwin B. Smith, I conducted a floristic survey of the aquatic plants of North Central Arkansas.

One of the last things to finish with my thesis was to visit the herbarium at Southern Methodist University in

Dallas. Since SMU had the Delzie Demaree Herbarium, an extensive collection of Arkansas plants, studying

the Demaree collections was essential for any serious floristic and distributional research on Arkansas plants.

Upon arrival at SMU, I met the herbarium botanist, Mr. Jerry Flook, and the director of the herbarium, Dr. Wil-

from the herbarium for another job. The position of herbarium botanist at SMU would soon be open for appli-

cations, and Dr. Mahler invited me to apply. 1 submitted an application and, lo and behold, on August 20, 1975,

1 launched my botanical and herbarium botanist career at SMU with some hesitation and trepidation. Dallas,

once surrounded by cotton fields, was a big city that seemed far away from the rolling red plains of southwest-

ern Oklahoma, where 1 was born and raised on a cotton farm near Temple, Oklahoma. I might have felt at home

in Dallas a century earlier when it was the world center for the cotton trade, but cotton was in the past in Dallas

in 1975. 1 needed support and guidance in a city of 850,000-plus people. Dr. Mahler was a new boss, but, more

important, he was a link to a familiar way of rural life. Dr. Mahler was born and raised in Iowa Park, Texas, a

small rural community some 35 miles southwest of Temple, Oklahoma, my birthplace. Bill owned a farm in

Oklahoma about 15 minutes as the crow flies from where 1 grew up — all comforting to a country boy in a big

city like Dallas. Dr. Mahler gave me confidence that Dallas would be good for me. So, I lifted my head high, and

a lifelong friendship and professional relationship began with Dr. Mahler. 1 always called him Dr. Mahler until

much later in life, when, years after retirement, I simply called him Bill. After all, working side by side in the

SMU herbarium and sometimes in the classroom, taking field trips together, attending meetings, enjoying

lunch and coffee almost every day, identifying plants together, and simply working with each other for almost

38 years, we were more than colleagues, we were close friends. His family became my family, and in more

senses than one, 1 was a sibling of the Mahler family. I enjoyed everything from dinners at the Mahler home to

tching Monday Night Football with them. Bill became my botanical mentor, field companion, and a confidant

:, and after I went through a divorce in 1976, Bill and his sweet

is death. The family

vife, L

along with my o

Fin 1968 a

ere my life support. My life turned out for the better

rter-time professor in the biology department, located in the

Fondren Science Building, and as a one-quarter-time employee of the herbarium, located in the basement of the

Science and Engineering Library. Lloyd H. Shinners (1918-1971) was director of the herbarium, and Bill as-

sumed the directorship after Lloyd’s death. Fondren Science and the science library were on the north side of

the campus, with only a parking lot separating them. Not long after my arrival at SMU, Bill sometimes invited

me to assist him with his taxonomy classes, in Fondren, and with labs, usually in the herbarium. The class-

room and laboratory interaction with Bill, which was over and beyond my herbarium duties, slowly gave me

confidence and a sense of place and purpose. After teaching class in Fondren Science, Bill would return to the

herbarium, which he used as his official office. When Bill was away from the herbarium, my life in the base-

ment was often quite isolated from the academic world above ground. Bill was my connection to the academic

life at SMU. My daily herbarium activities were regularly interrupted by science library staff looking for her-

barium books and journals being requested through interlibrary loan or by the occasional visiting scientist;

Ddzie Demaree in those early years was a regular visitor. Otherwise, my routine as a herbarium botanist in-

volved a diversity of duties that included processing loans and exchanges, filing Gray Herbarium Index cards,

processing and mounting plant specimens, filing specimens, identifying specimens, selecting botanical books

for purchase by the library, shelving new books and journals, and overseeing work-study students Every day

at least during the school year. Bill and I would sit at a table in the herbarium basement and have our lunch, ai

the very same place where Lloyd Shinners used to sit and sort plants. We discussed everything from Shinners

to botanical topics to SMU politics to local and national news. BUI was my advocate atSMU, always looking out

for me and trying to improve the quality of my life. My starting salary in 1975 was $7,500 and after six months

I got a hefty $500 raise.

1**7 12 m °7 hs 0r 50 doin S her banum work. Bill introduced me to the publishing world of botany. In

1976, Sida, Contributions to Botany, the journal Bill had inherited from Shinners, its founder and private pub-

lisher, was a 15-year-old taxonomic journal. After Shinners died, it was uncertain whether the journal would

sun™ hu. tndeed it Ih» today in prim and digital formats around the world because of several people, in-

cludmgjohn W. Thieret, but pnmanly because of the dedication and perseverance of Bill Mahler. In 1976, the

journal wasmits sixth volume, and in its early daysavolume had multiple issues, sometimes as many as seven.

Journal of the Botanical Research Institute of Texas 7(2)

ger, all 1 had to do was select a 17th-century herbal, Linnaeus’ Species Plantarum (1753), or an early issue of

Curtis Botanical Magazine (1787) from the shelves of Stunners’ library and examine or feel the letterpress print

on the pages. There is hardly anything quite like the feel of letterpress printing, which was the norm until the

mid-20th century, when offset printing was developed.

My herbarium duties were first and foremost, but at every opportunity I assisted Bill with the journal. In

1977, after one year of working with Bill on the journal, he appointed me as assistant editor; I was made full edi-

tor in 1982, but still with herbarium duties. In 1987, Bill and I cofounded the monographic book series, Sida,

Botanical Miscellany. Through it all, I received inspiration and guidance from my colleague and friend, William

F. Mahler. He was the light unto my path toward becoming even a semblance of an editor.

Early 1980s (The Struggle for Existence)

With the 1980s came social, economic, and broad change at SMU. Personal computers were introduced but,

more significantly, an economic change came, with the housing market crash, the savings-and-loan crisis, and

NCAA’s “death penalty,” canceling SMU’s 1987 football season. With all of this happening, there were serious

budget problems and space issues all over campus. As a result, the herbarium and library came under a watch-

ful administrative eye. All the while, Bill and I promoted the herbarium and library as an important resource to

the SMU community, to Dallas, and to the scientific community at large. We had consistent visitors and users

of the herbarium and library from on campus and off. I regularly processed loans of herbarium specimens to

researchers, assisted with interlibrary loans, and regularly edited manuscript submissions and published Sida.

The journal was an international journal of systematic botany and was being distributed internationally. Bill

and I struggled with annual budget cuts by the SMU administration, and this included my receiving an annual

dismissal letter every year for four or five years straight. SMU was looking at a variety of alternatives for closing

the herbarium and library to save money and gain space. Bill was an unflagging optimist and resourceful, and

he always found a way to keep me on the job and to keep the SMU Herbarium open for business. To suppor t the

annual budget, we took on botanical consulting jobs together, we wrote funding proposals, and took the her-

barium on the road, if you will, to generate awareness and support. Those were the days we started recording

in detail every phone call, every visitor, and every request on and off campus to provide at least some tangible

justification for keeping the herbarium and botany library open and functioning. Bill kept the closure of the

herbarium at bay for a few years through contract

work, but still that didn’t seem to be enough, and by

the mid-1980s, budgets were even tighter and space

on campus was at a premium. All the efforts to sup-

port the herbarium and library ultimately failed, or

at least they weren’t seen as enough, and SMU start-

ed again to seriously consider alternatives for the

herbarium and library. Ultimately, SMU placed its

herbarium and library on permanent loan first to

the Dallas Civic Garden Center and finally to a

group of citizens in Fort Worth who had the vision

and determination to set up a nonprofit called the

Botanical Research Institute of Texas that would ac-

cept and care for the Shinners collections.

Thank goodness for failures. Garrison Keillor

said to the Harvard chapter of Phi Beta Kappa, . . it

William F. Mahlef, )1 March 2012, atttie new BRIT facility, Fort Worth, Texas, j? f ° r y ° U “ thinkabout the ^fils

Failure is essential, a form of mortality. Without

failure, we have a poor sense of reality. In a nutshell,

629

my advice is Go, and have a crisis,” as if to say,

ane day that will stave off a crisis of greater

magnitude. Garrison was right. Those 1980s

:rises at SMU somehow, some way, prepared

Bill and me for reality, and ultimately, with as-

sistance from Bill’s student Andrea McFadden,

to establish BRIT in 1987 as a free-standing

1987-1992 (Brighter Days)

The first 17 years of my botanical caret

Bill Mahler at SMU are a source of en

memories, priceless photos, and roller-i

F. Mahler, 31 March 2012, atthenew BRIT facility, Fort Worth, Texas, i

s 25 years of age a

Edward P. Bass, and Lindsay Holland.

t SMU, and 1 w

as so impressionable in those early years; Bill’s presence was such a positive influence. 1 fondly

tersonal confidence, persistence, compassion, integrity, and loyalty, all of which touched and

ter botanist, colleague, editor, and individual for having known and worked with

Bill from 1975 to 1987 and even beyond. When BRIT was formed in 1987, Bill retired from SMU. Thus 1987 was

a new beginning, and I had the benefit and joy to work another five years with a dear friend and colleague.

After a happenstance meeting in June 1987, with an international real estate broker (Theodore McAlister)

and a client of his who had a large tract of Costa Rican timber to sell, the benefits of failure began to truly ma-

terialize. At this time, the SMU herbarium and its library had been placed on permanent loan to the Dallas

Civic Garden Center, but the arrangement wasn’t working out. After that early June meeting with Mr. McAli-

Texas. BRIT was established as nonprofit corporation in Texas on October 2, 1987. With a staff of three people

and a board of trustees, BRIT was on its way. Bill was director, I was curator/librarian/editor, and Andrea Mc-

Fadden was executive director. BRIT had a presence in Fort Worth, but the herbarium, library and staff were

still housed in the basement of the SMU science library. The search began for a new home in Fort Worth and,

through the support of its board, BRIT moved into a turn-of-the-century restored warehouse near downtown.

The basis of BRIT at this time was the Lloyd Shinners Collections in Systematic Botany and Sida, of which Bill

was still publisher and owner. Before the move to Fort Worth, the first two years of BRIT’s existence (1987-

1989) were formative and trying times for all of the staff and the new BRIT board. There were successes and

failures, but surely we would not face another? Our destiny was truly in our own hands. It is hard to see the

benefits of failure when you are experiencing crises. Bill Mahler always saw the glass as half full and never let

anything really get him down. At one point in the early life of BRIT, the payroll could not be met, and yes, none

other than Bill, lent money to the fledgling institution so work could continue. Years later Bill was repaid. He

always believed there were brighter days ahead, and it was so, and Bill got to enjoy and see the future of BRIT in

its new building, which opened in 2011. In February 1993, Dr. Sy Sohmer assumed directorship of BRIT and

Bill became director emeritus. Bill and Lorene retired to his hometown, Iowa Park. Iowa Park was on the route

I took to visit my family in Oklahoma, and that meant an occasional visit with Bill there.

My day-to-day interaction with Bill ceased with his second retirement, in 1992. Aside from my visits with

Bill in Iowa Park, I got to see him regularly in Fort Worth at BRIT events. In 1995, BRIT began an annual fund-

1995

630

through 2009 Bill and his wife were guests at the

gala. He and Lorene purchased tickets and often

brought friends to Fort Worth to join them; Bill and

his wife were always recognized at the event. The

years of hard work, along with the trials and tribula-

tions that came with keeping BRIT alive and going

forward, were truly recognized. Over the years, Bill

slowed down, and, in 2010 he was unable to make the

trip. I’ll never forget attending the 2010 banquet and

not seeing Bill and his wife there. I took home with

me one of the beautiful floral table arrangements and

made a visit the very next day to Iowa Park. I proudly

gave Lorene the floral arrangement and a copy of the

program, and enjoyed my visit with them. The 2009

for Bill, his family, the BRIT board and staff, ;

d. The new green building was more than anyone had dreamed

more than anyone could have envisioned on October 2, 1987, the day it was established as a nonprofit. Sine

t en, almost 27 years ago, it was a dream of the early founders, supporters, board members, and staff that on

day the BRIT collections would move to the dynamic Fort Worth Cultural District. For all of us, but especially

an ^ *”s fe m * ly ’ Ntoy 15. 2011, will be a day t° remember. H e was proud of BRIT, and everyone was proud

of Bill and his perseverance.

The last day Bill visited BRIT was March 31, 2012, wilh his family by his side. Bill was still of sound mind

and sptnt but needed a little help getting around. It was a very special day that will live forever with him, his

amily, and BRIT fnends. It was a reunion of the two early board members Lindsay Holland (from Midland),

t e first president of BRIT, and Ed Bass (from Fort Worth), vice president of BRIT, Andrea McFadden (from

Seattle), BRITs first executive director, and me (curator/editor/librarian). It was a homecoming that started

with a wonderful lunch in a new boardroom and concluded after two hours of touring and talking about the

Wm. F. Mahler: In memory of BRIT co-fi

new BRIT. We shared stories about BRIT’s early years at SMU and its first home in Fort Worth (Tindall Ware-

house, a restored tum-of-the-century building), where it was located from 1993 until 2011).

My memory of Bill lives on, and I’ll be forever grateful for what he gave me. His memory at BRIT is re-

flected in more ways than one. His portrait hangs in the BRIT library adjacent to that of Lloyd Shinners, whose

collections are the heart and soul of the Botanical Research Institute of Texas. It almost never fails that when I

walk into the BRIT library, 1 look over at Bill and Lloyd and smile, and in my heart, I say, “Well done, thou good

and faithful servants. Thank you.”

— Barney L. Lipscomb, Fort Worth

Journal of the Botanical Research Institute of Texas 7(2)

BOOK REVIEW

o Alaska. (ISBN-13: 978-1-

lbum, Washington 98001,

ng.com, 1-800-518-3541).

Andy MacKinnon and Jim Pojar. 2013. Alpine Plants of the Northwest: Wyoming

55105-892-4, pbk.). Lone Pine Publishing, 1808 B Street NW, Suite 140, A

U.S.A. (Orders: www.lonepinepublishing.com, order@lonepinepublish

$29.95, 528 pp., 5 W x 8 Vi".

Pojar and MacKinnon have created the format for field guides by which all others are measured. Alpine Plants

of the Northwest: Wyoming to Alaska follows in the footsteps of previous volumes in regard to the content, lay-

out, and presentation. The only shortcoming may be in the breadth of the material. From an ecological per-

spective, it makes sense to cover a region stretching from far northern Alaska southeast through the Cascade

Range and the continental divide to Wyoming. From the perspective of a hiker seeking to balance weight with

knowledge, the book may stretch half a continent too far. For an armchair ecologist, however, there is no better

guide to the cold-climate plants of the Pacific Northwest.

For a non-technical field guide, the Pojar and MacKinnon books offer a spectacular balance of usability

and detailed information on taxonomy. For the truly novice, the book offers a plethora of well-composed pho-

tographs. Each portion of a plant that is most distinctive is highlighted as needed, from flowers to leaf shapes

to canopy oudine for trees. Many guidebooks stop short at one or two photographs of a flower, leaving the

user out of luck should the trailside specimen be poor quality. For the more sophisticated user, the Pojar and

MacKinnon guides are organized by family with non-technical dichotomous keys emphasizing easily field-

discernible characteristics within each section.

The importance of a well-constructed key, organized by easily visible characteristics and supplemented

includes significant characteristics that distinguish a taxon from closely related or easily confused groups.

botanists more familiar with the flora of other ecoregions will find books by Pojar and MacKinnon invaluable.

From a simple physical perspective, the Pojar and MacKinnon guidebooks are constructed of high-quality,

water-resistant paper (a necessity in any PNW field guide!). The bindings are sewn, with the anticipation that

the guide will be well-thumbed and frequently used. Finally, each book strikes a good balance between page

dimensions (larger pages mean more room for photographs) and portability (smaller dimensions mean less

space). The weight of the paper adds some heft to the finished product, but this inconvenience is well worth the

additional life span of the tome.

Although not so useful for identification, short snippets of ethnobotanical information are included for

plants, whether edible or medicinal, contributes to the overall sense of each plant species as an element of an

Perhaps the most admirable element of Pojar and MacKinnon’s guides is their usefulness to the amateur

naturalist looking to learn more about a flora. The approachability and accessibility of the guide makes it very

useful to the inexperienced. The more detailed information about flower morphology and the characteristics

that distinguish common families in the Pacific Northwest provides an excellent opportunity for the enthusi-

ast to learn the basis by which plants are classified.

As stated initially, the only significant drawback of this guide is its heft for those hikers wishing to cut

every ounce, but the breadth of territory covered necessitates a substantial volume. This approach may be de-

sirable from an eco-regional perspective — and from the additional expense that would presumably follow

state-specific books. From the perspective of an interested naturalist who values knowledge over convenience,

however, there is really no substitute. Aspiring field guide authors from other regions of the US would do well

to emulate the Pojar and MacKinnon formula. — Brian Witte, PhD, Botanical Research Institute of Texas Research

Associate, Adjunct Professor of Biology at Collin College, and freelance writer.

FOUR NEW ANNUAL SPECIES OF EUPHORBIA SECTION

TITHYMALUS (EUPHORBIACEAE) FROM NORTH AMERICA

Mark H. Mayfield

Herbarium, Division of Biology

Kansas State University

Manhattan, Kansas 66506-4901, USA.

ABSTRACT

INTRODUCTION

North American species of Euphorbia L. include members of three of the four major clades discovered and cor-

roborated by numerous recent phylogenetic studies (e.g., Steinmann & Porter 2002; Bruyns et al. 2006; Horn

et al. 2012). Among those clades, the one circumscribed as Euphorbia subgenus Esula Pers. is highly diverse in

the Old World (>400 spp.), while in the New World (<50 spp.) it is primarily represented by Euphorbia section

Tithymalus (Gaertn.) Roep. The taxonomic limits of section Tithymalus were recently refined by Riina et al.

(2013): with the new circumscription this section is unique within subg. Esula in that most of its diversity oc-

curs in the New World, with 35-40 species of annual and perennial herbs. All of the members of section

Tithymalus share a lack of stipules, cyathial glands having either horns from the lateral margins, or having

marginal crenae (with or without long horns), seeds with conspicuous caruncles and smooth to variously pit-

ted or reticulate seed surfaces. The most recent complete taxonomic revision for North American species of

this group was by Norton (1899), who treated all taxa north of Mexico belonging to section Tithymalus (sensu

Boissier 11860], with nearly the same constitution as the contemporary subg. Esula sensu Riina et al. 2013).

Geltman et al. (2011) recently updated the nomenclature including a novel synonymy and lectotypifications

for essentially the same set of taxa (i.e., Euphorbia subgenus Esula sensu Riina et al. 2013) “. . . as a precursor to

the treatment of Euphorbia for the Flora of North America.” The latter made no taxonomic changes (other than

lectotypification) to the suite of short-lived taxa of the revised sect. Tithymalus relative to the same species in

i. Bot. Res. Inst Texas 7(2): 633 - 647. 2013

634

Journal of the Botanical Research Institute of Texas 7(2)

The annual North American members of section Tithymalus (not including E. commutata and E. crenulata,

which are usually biennials) are much less well known and more restricted in distribution in comparison to

the non-annual taxa, and they have been more stable taxonomically. From the time of Norton’s (1899) treat-

ment to now, these have included five species within the southern central U.S. states of Arkansas, Kansas,

Louisiana, Oklahoma, and Texas. Notably, all five species occur primarily or exclusively in Texas wherein

three are endemic and two are chiefly distributed (Correll and Johnston 1970; Turner et al. 2003). These species

have strong ecological affinities and clear separations in morphology and geography (Berry et al.. Flora of North

American North of Mexico, submitted). Most conspicuous among these is the relatively robust E. roemeriana

Scheele, endemic to the shaded canyonlands of the Balcones Escarpment of Central Texas. It is highly

branched, bearing conspicuous yellow-green, horizontal subcyathial bracts (“raylet leaves” in Riina, et al

2013), and having seeds with reticulate ridges. The remaining four species are more sparingly branched from

the base, and have raylet leaves that are greenish and are held at an oblique angle or perpendicular to the hori-

zontal (directed away from the axis of the plant). Euphorbia helleri Millsp. occurs on clayey soils near the Texas

coast from the lower Rio Grande Valley to the Coastal Bend area, and inland to Gonzales and Karnes Counties

(reports from Louisiana [e.g., Thomas & Allen 1996] were misidentified). Euphorbia longicruris Scheele occurs

sparsely across a broad swath of the central Texas Hill Country to the southern half of Oklahoma’s prairies on

thin soils over limestone and sandstone outcroppings, and in the Ouachita Mountains to Hot Springs, Arkan-

sas, on gladey shale outcrops. Euphorbia peplidion Engelm. is most prevalent in the Coastal Bend area, but

ranges widely from there north to Travis County, and west to eastern Pecos and Val Verde Counties (the latter

not indicated by Turner et al. 12003]). It grows in margins of eastern post oak, in Tamaulipan thomscrub, live

oak sandylands, and adjacent to coastal scrublands, where it seems to prefer silty or sandy soils over limestone.

Although it has yet to be collected from Mexico, from an ecological perspective, it is one of the more likely of

these species to have a natural distribution yet to be discovered there, as speculated by Johnston (1975). Lastly,

E. tetrapora Engelm. is here considered to be endemic to the western Gulf Coastal Plain from central Louisiana

to Wilson County, Texas, and north within the Cross Timbers to southern central and southeastern Oklahoma

(I have seen no specimens that can reliably be placed from east of the Mississippi River [e.g., “ Boykin , Georgia",

Engelmann 1858] and it has never been well-documented from there). It occurs in loose, sandy soils in open-

ings in mixed upland savannah woodlands with post oak (or sand post oak), often with loblolly pine.

This study presents four previously undescribed annual species and one variety within Euphorbia section

Tithymalus (sensu Riina et al. 2013). Field and herbarium work by the author on New World Euphorbia, includ-

ing access to collections not available to earlier authors, has enabled discernment of this diversity. This work

advances understanding of diversity in New World Euphorbia, and lays groundwork for broader, phylogenetic

study of section Tithymalus (Peirson et al.,

Glabrous annual herbs with taproots, 6-25(-28) cm tall, stems erect, 1 to a few stems spreading from the base.

Stem leaves alternate, sessile, strongly ascending to divergent (with age) above, blades 5-18 mm long, 0.8-2.5

mm wide at the widest point, narrowly oblanceolate to nearly linear, less often somewhat spatuliform, bases

linear to linear-attenuate, apices rounded to obtuse or acute. Ray leaves 3, sessile, 5-33 mm long, 1. 5-3.0 mm

wide, narrowly oblong to narrowly lanceolate. Primary inflorescence rays 3, usually with 1 to 3 internodes

from 1 .5-3.5 cm long, upper nodes becoming monochasial, secondary rays 0 to 3 in the upper mainstem leaf

axils. Raylet leaves free at the base, 3-8(-13) mm long, length/width ratio 0.7-1.5(-3.0), reniform-ovate to

subdeltate-ovate, rarely broadly lanceolate, bases obliquely truncate to rounded, apices broadly acuminate to

Mayfield, New species of Euphorbia sect. Tithymalus

635

Journal of the Botanical Research Institute of Texas 7(2)

attenuate, blades oriented vertically, adaxial surface directed away from the plant axis. Cyathia: involucres fun-

nelform, 0.8-1.0 mm high, 0.6-0.7 mm wide, on a stalk 0.1-0.2 mm long; cyathial glands lunate, 0.3-0.4 mm

wide, horns 0.5-0.7 mm long, relatively stout and blunt, entire to rarely slightly bifurcate at the apices, the

margins entire between the horns, involucral lobe apices rounded, short-ciliate on the margins; staminate

flowers 5 to 10. Pistil: styles 3, ca. 0.3 mm long, free nearly to the base, bifid at the apices for of the length,

the lobes divergent and spreading away, capitate. Capsules 2.0-2.2 mm long, 3.0-3.2 mm wide, columellas

1.8-2.0 mm long. Seeds 1.4-1.7 mm long, 1.0-1.3 mm wide, rotund-ovoid, rounded in cross-section, surface

ashen-gray to whitish at maturity, with deep, irregular to rounded, concave depressions crowded over the en-

tire surface; caruncles 0.7-0.8 mm wide, reniform-ovate, stipe present.

Etymology. — This plant is named for the region in which it occurs.

Chromosome number. — n = ca. 12-13 II (reported here, below).

M. Ph aneuflUO (KSC, LSU, MICH. TEX), also 20 Mar 2010, M.H. Mayfield el al 3843 (KSC, LSU, MICH, TEX); Exit 113 on southbound In-

terstate 37, E of onramp, S of Brite Cemetery Road, 460 ft, [corrected coordinates: N 29°03'06" W 98°25'59"1, 29 Mar 1992, M.H. Mayfield &

M. Phaneufll73 (LSU, MICH, TEX), also 26 Mar 1995, M.H. Mayfield&C.J. Ferguson 2160 (LSU, TEX). Bexar Co.: W side of IH-37, ca. 500 ft

N of mile marker 124, 1.3 roadmiles N of overpass at Priest Road-Mathis Road exit, 29°12'00"N, 98°25'12"W, 560-570 ft, 20 Mar 1993, W.R.

Carr & M. Mayfield 12504 (BRIT, KSC, LSU, MICH, TENN, TEX, UARK), also 13 Mar 1995 M.H. Mayfield 2128 (BRIT, LSU, TEX, UARK);

U.S. Hwy 281, S of San Antonio, 4.1 mi N of jet. with 536 (in Espey), 650 ft, 29°09'59"N, 98°28'56"W, 13 Mar 1994, M.H. Mayfield, C. Fergu-

son, & A.L. Hempel 1874 (LSU, TEX); 0.6-0.7 road mi ENE of 1-37 [at exit 122] on E side of Priest Road, S end of county, 620 ft, 29°iri5"N,

98°24'53"W, 4 Apr 1993, M.H. Mayfield,]. Mendenhall &J. P anew 1720 (BRIT, KSC, LSU, MICH, TEX). Wilson Co.: Gene Dodgen property,

EsideofFM [road] 1303. W.R. Carr etaL 13345 (TEX).

Populations of this species occurring on the South Texas sand sheet are recognizable as a distinct variety, with

the following key to distinguish them from the typical variety. Some of these plants were taken by Turner

(2011) to be evidence for the persistence of E. exigua L. in Texas; the latter taxon, an introduction from the Old

World, belongs to a separate section, E. sect. Exigiiae (Geltman) Riina & Molero, and can be readily distin-

guished by its seeds with tubercular prominences (Fig. 2).

Euphorbia austrotexana var. carrii M.H. Mayfield, var. nov. (Fig. 3). Type: UNITED STATES. Texas. Kenedy Co.: 50-200

dalgo, Kenedy, and Willacy counties, 50 ft, 16 Mar 2004, W.R. Carr & M. Pons 22784 (holotype: TEX).

» longer than wide), and seeds smaller, w

Euphorbia austrotexana occurs in stabilized sandy soiled habitats, with a range extending through a wide swath

of the south Texas plains. Although it is apparently locally abundant in some cases, only five documented lo-

calities for var. austrotexana are known. One of these is on Queen City Sand formation (both sides of 1-37

around Exit 113, Atascosa County), while the other four population areas are on a particularly massive repre-

and the San Antonio River (Wil-

t side of the San Antonio River in

suitable post oak/blackjack sandy savannah only yielded E. tetrapora in

tribution. The populations of E. austrotexana var. carrii are scattered acr<

the southwestern-most part of its dis-

iss the south Texas Sand Sheet, an area

Fk. 3. The hoiotypeofE austrotexana var. amir. Can & Pons 22784 (TEX).

Mayfield, New species of Euphorbia sect. Tithymalus

642

Journal of the Botanical Research Institute of Texas 7(2)

0.5-0.9, deltate to broadly deltate, the bases truncate, apices obtuse to abruptly acuminate, blades oriented

nearly horizontally, the adaxial facing upwards. Cyathia: involucres broadly cup-shaped, 0.8-1.3 mm high,

0.6-0.10 mm wide, on a stalk 0.3-0.5 mm long; cyathial glands lunate, 0.9-1.1 mm wide, horns 0.5-0.7 mm

long, attenuate-filiform, the margins entire between the horns, involucral lobe apices truncate and crenate on

the margins; staminate flowers 10 to 15. Pistil: styles 3, ca. 1.5 mm long, connate at the base ca. 0.2 mm, bifid at

the apices for of the length, the lobes divergent and ascending, capitate. Capsules 1.8-1.9 mm long, 2.3-

2.5 mm wide, columellas 1.7-1.9 mm. Seeds 13-1.5 mm long, 0.6-0.7 mm wide, oblong-ovoid, rounded dor-

sally, slightly flat ventrally in cross-section, surface white to tan, darker in the depressions at maturity, with

irregularly shaped broad concave depressions over the entire surface, ridges bordering pits rounded in relief;

caruncles 0.4-0.5 mm wide, reniform-ovate, base stipe absent or inconspicuous.

Euphorbia nesomii is the first annual species of the section to be described from Mexico, where it occurs on

relatively mesic limestone north-facing slopes in the mountains of northern Nuevo Leon in montane oak chap-

arral habitat under Brahea spp., Cheiropetalum schiedeanum, Fraxinus greggii, Osmanthus sp., and Ungnadia

speciosa. In addition to similarities in leaf and habit shared with E. roemeriana , it’s seeds resemble those of E.

peplidion in size and shape. Otherwise it has no clear shared features that might link it to any other of the other

annual species. Like the last species described above, it has been poorly collected and warrants additional

documentation and study.

Etymology. — This species is named for Dr. Guy L. Nesom, who first brought this plant to the authors at-

tention when he collected it in 1993. In addition to being invaluable as a mentor during the early phase of the

author’s career, he has had an indelible impact on the field of Plant Taxonomy. The epithet honors his many

contributions to botany and his generous spirit.

Glabrous annual herbs with taproots, from 12-28(-32) cm tall, stems erect-ascending, often basally decum-

bent, 1 to a few stems from the base. Stem leaves alternate, sessile or briefly petiolate, laxly ascending to hori-

zontal and spreading, petioles 0-3 mm long, blades 8-20 mm long, 3-9 mm wide at the widest point, broadly

oblanceolate to subspatulate, bases attenuate, apices rounded. Ray leaves 3, sessile, 10-25 mm long, 15-32 mm

wide, ovate-deltate to subrhombic-ovate. Primary inflorescence rays 3, usually with 3 or 4 intemodes from

2.5-6.5 cm long, upper nodes dichasial, secondary rays 1 to 5 in the upper mainstem leaf axils. Raylet leaves

connate to ca. 3 mm at the base, 6-18 mm long, length/width ratio 0.8-1.2, broadly deltate to subreniform,

bases truncate to broadly obtuse, apices obtuse to bluntly acuminate, blades oriented obliquely, adaxial surface

directed upwards and away from the plant axis. Cyathia: involucres funnelform, 1.3-1.6 mm high, 1.0-1.2 mm

wide, on a stalk 0.4-0.7 mm long; cyathial glands oblong, 0.7-1.0 mm wide, horns 0.2-0.4 mm long, attenuate-

filiform, the margins entire between the horns, involucral lobe apices rounded short-ciliate on the margins-

staminate flowers 15 to 20. Pistil: styles 3, ca. 0.8 mm long, free to the base, bifid at the apices for %-V4 of the

length, lobes ascending, capitate. Capsules 2.6-2.7 mm long, 2.5-2.7 mm wide, columellas 2.0-2.1 mm long.

Seeds 1.8-2.1 nun long, 1.1-1.2 mm wide, oblong-ovoid, rounded dorsally, slightly flat ventrally in cross-sec-

tion, surface dark-brown, lustrous at maturity, with deep, rounded, uniformly spaced pits in vertical rows (3-4

in a vertical row per each ventral facet, 14-18 in 4 rows on the dorsal facet); caruncles 0.6-0.7 mm wide, reni-

form-ovate, base stipe present, umbonate.

Mayfield, New species of Euphorbia sect. Tithymalus

Fk. 6. KSC isotype Of Euphorbia ouachitana : Mayfield 3551.

Etymology. — This plant is named for the region in which it is best represented.

Chromosome Number.— n = 13 II ( Urbatsch et al. 981 [LL], Urbatsch et al. 1975).

Euphorbia ouachitana occurs in semi-open forests and woodlands, often in soil*

also occurs in areas that are less sloped, and limited by thin soils on lii

it is associated with the herbs Packera obovata and Saxifraga virginiensi

The best-documented and expansive continuous part of the range for E. ouachitana is in the Ouachita

Mountains from southeastern Oklahoma to the area around Hot Springs County, Arkansas. The type is from

near the western margin of this range in Pushmataha County, Oklahoma. In Missouri, specimens are known

only from the area of Roaring River State Park in Barry County. In Tennessee, its range is very possibly under-

represented by collections. Five of the six known Tennessee populations are from north of the Cumberland

River (Trousdale and Smith Counties) in the low, isolated hills situated on the edge of the Nashville Basin and

eastern Highland Rim. A separate population further to the south in Rutherford County, Tennessee suggests

that this species may eventually be found across a wider area of the Cedar Barrens in middle Tennessee. In

Texas, the single population listed above was only recently discovered and reported as E. commutata by Sing-

hurst et al. (2013).

Most of the range of E. ouachitana occupies the region between the southwestern-most populations of

E. commutata and the northern-most populations of E. tetrapora in southeastern Oklahoma. Most previously

collected specimens have been identified as E. commutata, reflecting a resemblance in habit. Yatskievych and

Mayfield (2006) were the first to discuss its distinctiveness from E. commutata, referring to the populations in

Barry County, Missouri. Among the more prominent features that distinguish these two species are life span

and seed morphology (Fig. 2). Euphorbia ouachitana is a winter annual that completes most of its growth and

reproduction during a short span in the spring from mid March to early May, whereas E. commutata is most of-

ten a biennial that grows vegetatively during its first year, then bolts to flowering in spring about the same time

as E. ouachitana. At this time of year, the two species stand in stark morphological contrast: E. commutata is

much taller, and bears numerous, closely spaced, long-petiolate elliptic leaves at the base of the stems, whereas,

m E. ouachitana, the lowermost primary stem leaf blades are sessile and attenuate to the base. The lustrous

reddish-brown seeds of E. ouachitana (Fig. 2), with deep pits in rows, are also easily distinguished from those

of E. commutata and all similar species. Euphorbia commutata has darker brown seeds with more numerous

pits that are not distributed in obvious rows. Lastly, if existing chromosome counts (see below) are correct

and consistent across these taxa, E. ouachitana (n = 13) is cytologically distinct from E. commutata (n = 14).

Euphorbia ouacUtma is also somewhat similar and may be confused with the annual E tetrapora It is

disttmt from the latter in larger overall plan, size, and its substantially latger seeds with deeper, well-defined

pus (Fig. 2). These two species also grow near each other in southeastern Oklahoma but E tetrapora occurs on

flat togently ^sloping terrain with sandy soil in association with Post Oalt-Blackjack Oak mixed woodlands

The only species of section TUhymalus with which E omchUam is broadly sympatric is E loneicruris a

spec.es that also occurs within the Ouachita Mountains of Oklahoma and Arkansas in shale slopes or glades

ounchr t sl ® nl ^ cant tree ovetstory (T. Witsell, Arkansas Natural Heritage Commission, pers. comm!). Euphorbia

ouachitana occupies a more shaded woodland habitat with leaf litter and isdistinctinitsseeds (Fig. 2), and habit.

^^n^r“ e "“ n ' d:USA '“^^ G ^ 850 r, 9 Apr 1934 HR

dt^y S of 34 3167WV ' 200^ ^ mer ( MO, U AR K ); Ouachita M ’ ’ ^

34.66646N, 93.28357W, 6 May 2008, T. Witsell &]. Krystofik OsflO^ANHC.KS^)^^’

trusive area, rocky novaculite hills, 19 Mar 1938, D. Demaree 16713 (SMU); P.O. t

Apr 1939, D. Demaree 18874 (MO, SMU); P.O. Malvern 350 ft Lake rather™ c.„. ' D i

(MO, RSA,; near the boundary of Lake Catherine State Park, T4SR18W secAbW ^dApr R „

N side of Camp Road. 34.33751N, 93.05338W, 23 Apr 2008, T. Witsell

23 Apr 2008, R. Bledsoe RF-08-21 (,

R- Bledsoe 08-23 (ANHC). Howard Co.’: Baker Springs, 11 /

nity of 34.61820N, 93.49467W, 16 May

Journal of the Botanical Research Institute of Texas 7(2)

CHROMOSOME NUMBERS FOR NEW WORLD SECTION TITHYMALUS

Chromosome study was undertaken where possible for taxa of Euphorbia sect. Tithymalus. Developing buds

were field-collected in a solution of four parts chloroform, three parts 95% ethanol and one part glacial acetic

acid; with later transfer to 70% ethanol. Following the technique of B.L. Turner described in Jones & Luchs-

inger (1986), anthers were dissected out, stained with acetocarmine solution, squashed and examined for

meiotic figures. A new count of n = 1311 was made for E. longicruris ( Mayfield 2174 [TEX], Lampasas Co., Texas);

and imperfect meiotic figures were observed for E. austrotexana var. austotexana (n = ca. 12-1311, Mayfield &

Ferguson 2160 [TEX], Atascosa Co., Texas), E. peplidion (n = ca. 1411, Mayfield & Ferguson 2161 [TEX], Atascosa

Co., Texas), and E. roemeriana (n = ca. 1411, Mayfield 2158). One previously published count for “E. tetrapora ” of

n = 1311 corresponds to E. ouachitana (Urbatsch et al. 1975). A chromosome number of 2n = 28 was reported by

Perry (1943) for the common eastern annual/biennial E. commutata. Among North American perennials, a

chromosome number of n = 1411 was reported for E. brachycera Engelm. from Otero Co., New Mexico (Ur-

batsch et al. 1975), whereas Ward (1984) reported n = 13 for the perennial E. chamaesula Boiss. Numbers for

three additional perennial taxa from the western U.S.A. are documented by herbarium specimen annotations

as having n = 13 chromosomes: E. lurida Engelm. (Garfield Co., UT, Windham 96-035 [MO]), E. sp. nov. aff. lu-

rida (Clark County, Nevada, Windham 98-239 [MO]), and E. yaquiana Tidestr. (Gila Co., Arizona, Windham

94-24 [MO]). Euphorbia peplus, a species closely related to the North American section Tithymalus (Riina et al.

2013), has a haploid number of 8 (numerous counts in the literature; see Tropicos, http://www.tropicos.org/Na

me/12800171?tab=chromosomecounts). Together, these data suggest that the North American species may

have undergone significant chromosomal evolution prior to their diversification, and that aneuploidy may

have played a role in speciation. It is therefore likely that further investigations into the chromosome numbers

may provide insight into evolution of the New World members of E. section Tithymalus.

EY TO THE NON-PERENNIAL SPECIES OF EUPHORBIA SECTION TITHYMALUS

IN THE NEW WORLD (AND SIMILAR NON-NATIVE TAXA)

Mayfield, New species of Euphorbia sect. Tithymalus

647

ACKNOWLEDGMENTS

I thank many colleagues for valuable discussions and other assistance with this work, including Paul Berry, Bill

Carr, Carolyn Ferguson, Dmitry Geltman, Cleo Mayfield, Guy Nesom, Tom Wendt, and Ray White. Thanks to

Paul Berry and Jess Peirson for their comments on an earlier draft, to Carolyn Ferguson for editing the manu-

script, to Geltman and Eugene Wofford for helpful comments in review, and to Ziaming Zhao for help with in-

terpreting chromosomes counts. I gratefully acknowledge the following herbaria for loans and assistance dur-

ing visits: ANHC, ARIZ, BRIT, GA, GH, MO, LUTEX, KSP, LSU, NLU, NY, PH, OKL, OKLA, RSA, SBSC, SMS,

TENN, UARK, UC, US. This is contribution number 14-157-J of the Kansas Agricultural Experiment Station.

REFERENCES

Bruyns, P.V., RJ. Mapaya, andT. Hedderson. 2006. A new subgeneric classification for Euphorbia (Euphorbiaceae) in southern

Africa based on ITS and psbA-trnH sequence data. Taxon 55:397-420.

Correll, D.S. AND M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner.

Engelmann, G. 1858. Euphorbia. In: J.Torrey, W.H. Emory, Rep. U.S. Mex. Bound. 2(1 ):1 85-193.

Geltman, D.V., P.E. Berry, R. Riina, and J. Peirson. 2011. Typification and synonymy of the species of Euphorbia subgenus Esula

(Euphorbiaceae) native to the United States and Canada. J. Bot. Res. Inst. Texas 5(1):143-151 .

Horn, J.W., B.W. Van Ee, JJ. Morawetz, R. Riina, V.W. Steinmann, P.E. Berry, and KJ. Wurdack. 2012. Phylogenetics and the

evolution of major structural characters in the giant genus Euphorbia L. (Euphorbiaceae). Molec. Phylogen. Evol.

63:310-324.

Johnston, M.C. 1975. Studies of the Euphorbia species of the Chihuahuan Desert region and adjacent areas. Wrightia

5:120-143.

Jones, S.B. and A.E. Luchsinger. 1 986. Plant systematics, second edition. McGraw-Hill Book Company, New York.

Norton, J.B.S. 1 899. A revision of the North American species of Euphorbia of the section Tithymalus occurring north of

Mexico. Missouri Botanical Garden, St. Louis (Re-issued as Ann. Rep. Missouri Bot. Gard. 1 1 :85-144. 1900).

Peirson, JA, R. Riina, M.H. Mayfield, CJ. Ferguson, L.E. Urbatsch, and P.E. Berry, in prep. Evolution of New World leafy spurges:

phylogeny and taxonomy of Euphorbia sect. Tithymalus.

Riina, R., J.A. Peirson, D.V. Geltman, J. Molero, B. Frajman, A. Pahlevani, L. Barres, JJ. Morawetz, Y. Salmaki, S. Zarre, A. Kryukov,

P.V. Bruyns, and P.E. Berry. 2013. A worldwide molecular phylogeny and classification of the leafy spurges, Euphorbia

n ITS and ndhF

Singhurst, J.R., J.N. Mink, and W.C. Holmes. 2013. A short chronicle of Euphorbia commutata (Euphorbiaceae) in Texas.

Phytoneuron 2013-60:1-6.

Steinmann, V.W. and J.M. Porter. 2002. Phylogenetic relationships in Euphorbieae (Euphorbiaceae) basi

sequence data. Ann. Missouri Bot. Gard. 89:453-490.

Thomas, R.D. and C.M. Allen. 1996. Atlas of the vascular flora of Louisiana, Volume II: Dicotyledoi

Euphorbiaceae. Louisiana Department of Wildlife and Fisheries Natural Heritage Program, Baton Rouge.

Turner, B.L. 201 1 . Persistence of the weed Euphorbia exigua in Texas. Phytoneuron 201 1-20:1-3.

Turner, B.L, H. Nichols, G.C. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Volume 1— Dicots. Sida, Bot.

Misc. 24(1). BRIT Press, Fort Worth, Texas.

Urbatsch, L.E., J.D. Bacon, R.L Hartman, M.C. Johnston, TJ. Watson, Jr., and G.L Webster. 1975. Chromosome numbers for

North American Euphorbiaceae. Amer. J. Bot. 62:494-500.

Ward, D.E. 1984. Chromosome counts from New Mexico and Mexico. Phytologia 56:55-60.

Yatskievytch, G. and M.H. Mayfield. 2006. Euphorbiaceae. In: G. Yatskievytch. Steyermark's flora of Missouri: Vol. 2:1010-

1057. Missouri Department of Conservation, Jefferson City.

BOOK REVIEW

Werner Kunz. 2012. Do Species Exist? Principles of Taxonomic Classification. (ISBN-13: 978-3-527-33207-

6, hbk.). Wiley-Blackwell, 111 River Street, Hoboken, New Jersey 07030-5774, U.S.A. (Orders: www.

wiley.com, 1-877-762-2974). $99.95, 280 pp., 6 W x 9 fc".

Do Species Exist by Werner Kunz is in many ways Western science writ small; it is an attempt to impose an or-

derly understanding on an inherently unruly natural world. The order in which Werner Kunz devoutly be-

lieves is that not only do species exist, but there exists a rational means for distinguishing one from another.

Each system has had its difficulties, from Plato with his Forms that under-lie all of the observable world, to

Linnaeus with his system of nomenclature that, he believed, would reveal the mind of God at the moment of

creation. In the modern era, the problem of classification has been both illuminated and complicated by Dar-

win’s insights into the origin of species. No longer are groups of organisms perfect and unchanging. Rather, we

are seeing them at a moment in time — not at peace in some final form, but rather in the very midst of a con-

tinual process of change.

It is exactly this understanding of evolution and its impact on taxonomy that informs Kunz’ s manuscript

He rightly points out that the question is made all the more difficult by the simple fact that everyone knows

what a species is, even though no one has been able to offer a succinct and useful definition. Certain elements

may be widely agreed on, such as the consensus that any useful definition of a species must contain the stipula-

tion of common descent. That is, that all organisms bearing a particular taxonomic designation must be more

closely related to each other than to any outside group.

It is in just such endeavors that Kunz is at his strongest. The first portion of the book is a painstakingly

detailed examination of just what is agreed on in modern taxonomy, followed by a dissection of the various

systems that have been employed by taxonomists in classifying the natural world. If Kunz finds few novel in-

sights in this survey, he does at least offer a very well organized critique. This is, after all, well-tread territory.

Kunz even notes that Darwm, in his On the Origin of Species, had pointed out the difficulty of defining that

central term: species.

If Do Species Exist were merely a summary of definitions and the shortcomings of taxonomic systems, it

would better serve as a chapter in a textbook, albeit one laden with poorly chosen illustrations clumsily com-

posed in Photoshop. Kunz, however, goes one step further, and ventures into advocacy for phylogenetics as the

only practicable way of conducting taxonomy. This advocacy, however, appears only in the penultimate chap-

ter (?) and is given a curiously cursory treatment, considering the meticulous critiques of the preceding chap-

ters. It is almost as though Kunz, having shown the inadequacies of competing taxonomic systems, presents

the “phylocode” as the last option standing.

The shortfalls in Kunz argument are three-fold. The first and most obvious is that he presumes that cur-

rently extant systems of classifications are the set from which we must choose. To quote Sherlock Holmes, “. . .

[wjhen you have eliminated the impossible, whatever remains, however improbable, must be the truth.” While

other systems may have logical or practical inconsistencies, it does not follow that phylocode, devoid of “ar-

chaic Linnaean terminology”, is a more appropriate system of classification. It may be the most logically con-

sistent method we currently know of, but that does not mean that is the system which best reflects the

The second and more serious flaw in Kunz’s argument for the phylocode approach is that his proposal

simply does not pass the “smell test” for working scientists. In other words, phylogenetic methods may pro-

duce mathematically and logically satisfying results, but the results are frequently of little utility outside the

trees which they adorn. The species concept, after all, is based on the observations of field scientists cataloging

morphological differences among the specimens they observe in the natural world. Morphology is at best a dim

HACKELIA TAYLORI (BORAGINACEAE),

A NEW SPECIES FROM NORTH CENTRAL WASHINGTON STATE (U.S.A.)

Richy J. Harrod

Lauri A. Malmquist*

USDA Forest Service

Okanogan-Wenatchee National Forest

21 5 Melody Lane

Wenatchee, Washington 98801, U.S . A

rharrod@fs.fed.us

USDA Forest Service

Wenatchee River Ranger District

600Sherbourne

Leavenworth, Washington 98826, U.SA.

Imalmquist@fs.fed.us

Robert L. Carr

Department of Biology

Eastern Washington University

Cheney, Washington 99004, U.S.A.

ABSTRACT

RESUMEN

Hackelia Opiz. includes 45 species of perennial plants distributed within Northern Temperate region, Central

and South America combined (Mabberley 1987). In North America, 28 species are recognized, comprising 34

taxa, many of which are narrow endemics (Gentry & Carr 1976). Species of Hackelia can be found in a wide

range of habitats, including sagebrush steppe, steep talus slopes, moist rock crevices, open deciduous forests,

and Abies or Pinos forests; and the distribution of many species are narrowly restricted based on habitat, geog-

raphy, or elevation (Carr 1974; Gentry & Carr 1976). Gentry and Carr (1976) completed a comprehensive study

of Hackelia and clarified taxonomic relationships of the species and subspecies in North America. However, the

taxonomic status of H. venusta (Piper) St. John has remained in question (Gamon 1988) and recently has been

the focus of taxonomic research (Harrod et al. 1999; Hipkins et al. 2003).

Hackelia venusta, as originally described, includes a white-flowered form found at one low elevation site

(488 m) 9.6 km northwest of Leavenworth, Washington, and a blue-flowered form found at four currently

known high elevation (ca. 2050 m) alpine locations about 18 km northwest and southwest of Leavenworth,

Washington (Carr 1974; Gentry & Carr 1976; Hitchcock et al. 1959) (Fig. 1). The two forms were shown to be

distinct from each other based on morphological traits (Harrod et al. 1999). However, enzyme band pattern

analyses did not provide evidence for taxonomic separation of the two color forms (Hipkins et al. 2003). Al-

though at first this may seem to be a dilemma, taxonomy is often the result of synthesis across many lines of

evidence, some of which may fail to support the taxonomic hypothesis (Grant 1992; Winston 1999; Hipkins et

Journal of the Botanical Research Institute of Texas 7(2)

650

al. 2003). Taxonomic entities are defined on a combination of morphology, genotypic data (e.g., isozymes),

ecology, reproductive isolation, and geographic distribution. There are compelling reasons to consider the two

flower color forms to be separate species based on morphological and ecological distinctions

, He ™ £ Pr ™ dC 3 , techniCal descri P tio " for the »*** form at the rank of species, here named Hackelia

taylori. Although H. taylori was shown to be morphologically distinct from H. venusta in our previous work

va.ie.KS of H. iffusa (Doug ex Lehmann) Johnston which „e did no. study previously, but which were in-

eluded in the enzyme work of Hipkins et al. (2003).

MATERIALS AND METHODS

Study Sites

fr ° m 10 7 Ukti ° nS US£d by Harr ° d Ct 31 (1999) WUh 2 additi ° nal H Populations

the Rattlesnake HilL fRfn ^ V3r COtt(mii (Piper) Carr ’ located on Private land in

the Rattlesnake Hills (RH), 25 km north of Sunnyside, WA 150 m elevation; and 2) H. diffusa var diffusa lo-

ca e at Oneota Gorge (CXI) near Multnomah Falls, 50 km east of Portland, OR 75 m elevation.

Morphological Measurements and Statistical Analyses

We followed the same methods and used most of the same data (individual plants from original 10 populations

with missing morphological data were dropped from analysis) as described in Harrod et al. (1999). Nineteen

morphological characters from three categories (vegetative, floral, and fruit) were scored for statistical analysis

and an additional 11 descriptive characters (e.g., leaf shape, leaf surface, color) were recorded (Table 1). These

data were collected from 25 randomly selected individuals from each population except the Cashmere Moun-

tain population, which consisted of data from 14 individuals, and Crystal Cirque with only 10 individuals.

Both principal components and discriminant analyses were performed on the quantitative morphological data

using SPSS 16.0 (SPSS, Chicago, Illinois). Principal components analysis (PCA) was used to evaluate the natu-

ral groupings among each sampling unit or operational taxonomic unit (population). Discriminant analysis

was used to establish the non-arbitrariness of group assignments. This analysis places each case (plant) within

the group (population) with which it shares discriminating characters (Anderson and Taylor 1983). The analy-

sis is biased in that it positions cases within the ordination based on discriminating characters to achieve

maximum separation of the defined groups. A plot of the cases based on the first two discriminating functions

can assist in visualizing distinction among groups and species. The data for these analyses involved a 237 * 19

Descriptive characters were not subjected to statistical analyses but were used to further detail morpho-

logical characteristics of the new taxon.

The addition of H. diffusa var. cottonii and H. diffusa var. diffusa in both the principal components and discrimi-

nant analyses lead to tighter groupings of cases as compared to the results of Harrod et al. (1999). Of the 19 com-

ponents that accounted for all the variance in the PCA, the first three accounted for 65.1% (32.5%, 20.7%, and

11.9%, respectively). Characters highly correlated with the first component were lower cauline leaf width and

length, upper cauline leaf width, and radical leaf width (Table 2). Hackelia diffusa var. cottonii and H. diffusa var.

diffusa separated from other taxa along the first component (Fig. 2A). Plant height, radical leaf petiole length,

radical leaf length, and upper cauline leaf length were characters highly correlated with the second component

(Table 2). Along this second component, H. taylori (CC and CM) separated from the H. diffusa var. arida popu-

lations forming a distinct group that was somewhat overlapping with the H. venusta population (TC) (Fig. 2A).

The third component separated H. venusta from H. taylori and characters highly correlated with the third com-

ponent were upper cauline leaf length, limb width, fomice protuberance, and calyx length (Table 2, Fig. 2B).

The discriminant analysis showed H. taylori was more clearly distinct from H. venusta and almost the en-

tire H. diffusa complex (Fig. 3). The first two discriminant functions accounted for 81.8% of the ability to dis-

tinguish among groups (51.2% and 30.6%, respectively). Total predictability that a case from a certain popula-

tion was correctly classified to that population was 95.8%. Individual cases (plants) had high predicted group

membership with the sampled population from which they were sampled. Hackelia taylori plants from CM

were 100% correctly classified and those from CC were 95.8% correctly classified, with 4.2% classifying with

Harrod et aL, A new species of Hackelia from north central Washington

653

Factor 2

Fig. 2. A. Ordination of populations of Hackelia examined in this study based on scores of principal components 1 and 2. The first two components

accounted for 53.2% of the total variance, 32.5% and 20.7%, respectively. Hackelia taylori populations (CC and CM) are highlighted with heavy black

line. B. Ordination of populations of Hackelia venusta and H. taylori examined in this study based on scores of principal components 2 and 3. The third

component accounted for an additional 1 1 .9% of the variance for a total of 65.1% for the first three. H. diffusa var. arida : BM = Burch Mountain, TW =

Tumwater Canyon, SC = Swakane Canyon, PE = Ponderosa Estates, MC = Moses Coulee, DE = Derby Canyon, DC = Douglas Creek. H. diffusa var. diffusa-. OG

= Oneota Gorge. H. diffusa var. cottonir. RH = Rattlesnake Hills. H. venusta: TC = Tumwater Canyon. H. taylori: CM = Cashmere Mountain, CC = Crystal Creek.

Journal of the Botanical Research Institute of Texas 7(2)

Score 1

Tumwater Canyon. H. taylori: CM = Cashmere Mountain, CC = Crystal Creek.

Moderately short perennial, 1-2 dm tall; stems often many from a slender taproot, erect or spreading from

branched caudex. Leaves ciliate, antrorsely appressed strigose (Fig. 4A); radical leaves 3.7-10.4 cm long, 0.8-

2.9 cm wide, linear-elliptical, appressed strigose, petiolate for 1/3 to 1/2 their length; cauline leaves linear to

linear-lanceolate, sessile; lowermost cauline leaves 0.6-3.1 cm long, 0.4-1.2 cm wide; upper cauline leaves

1 .9-5.3 cm long, 0.5-1.3 cm wide, reducing upward to the inflorescence, becoming small bracts. Pedicel 2.3-

5.1 mm in flower (Fig. 3B) and 5.0-7.0 mm in fruit. Calyx length 2.4-3.4 mm, linear-lanceolate, strigose. Co-

rolla limb blue, 1.0-1.7 cm wide with five lobes, lobes 3.0-5.0 mm long (Fig. 4D, F). Fomices with appendages

showy, white, sometimes tinged pink, slightly emarginate, papillate-pubescent, rising 0.8-1.0 mm above the

throat; protuberances yellow, pandurate, 0.6-1.0 mm long (Fig. 4D, E). Anthers 0.8-1.0 mm long. Nutlets

1.8-3.6 mm long, lanceolate-ovate; dorsal surface verrucose-hispidulous, intramarginal prickles 7-13; mar-

ginal prickles connate for up to 1/2 their length, forming a flange 1.2-2.4 mm wide around the nutlet; distinct

prickle length 0.7-1.4 mm, a long prickle alternating with one or two shorter ones (Fig. 4C).

Etymology .— The epithet “ taylori ” honors Dr. Ronald J. Taylor, who taught botany at Western Washington

University, Bellingham, Washington. Dr. Taylor co-authored the first status report for H. venusta in 1979 and

was actively involved in native plant conservation in the Pacific Northwest for nearly 40 years.

Harrod et at, A new species of Hackelia from north central Washington

657

taylori flowers are always blue and H. venusta flowers are white but sometimes tinged blue suggesting perhaps

they share some genes for color.

Like H. venusta, this new species would benefit from well-developed conservation strategies. Populations

are at risk from loss due to stochastic events, such as rock slides, which were the cause of the loss of most of one

known population of H. taylori. Conservation strategies might include long-term seed banking so that popula-

tions could be re-established in the event of a stochastic loss.

ACKNOWLEDGMENTS

We would like to thank John Gamon, Loyal Mehrhoff, and Kali Robson for their work early-on suggesting a

new species description. Dottie Knecht, Mark Ellis, Cedar Drake, Ellen Kuhlmann, and Shelly Benson pro-

vided field assistance. Brandy Reed assisted with data analysis. We thank Pam Camp for help in locating popu-

lations of H. diffusa var. arida on BLM land. We thank Guy Nesom for providing the Latin description. This

project was cooperatively funded by the USDA Forest Service, USDI Fish and Wildlife Service, and the Wash-

ington Natural Heritage Program. Figures 1 and 2 were developed by Dan O’Connor, Okanogan-Wenatchee

National Forest. Illustrations of H. taylori in Figure 3 were drawn by Eve Ponder. We also thank two anony-

mous reviewers for detailed and constructive reviews.

REFERENCES

Anderson, A.V. and RJ. Taylor. 1983. Patterns of morphological variation in a population of mixed species of Castilleja

(Scrophulariaceae). Syst. Bot. 8:225-232.

Carr, R.L 1974. A taxonomic study in the genus Hackelia in western North America. Ph.D. Dissertation, Oregon State

il Heritage Program, Olym-

n of the genus Hackelia (Borac

Gamon, J. 1 988. Report on the status of Hackelia vt

pia, Washington.

Gentry, Jr., J.L. and R.L. Carr. 1976. A revisit

Mem. New York Bot. Gard. 26:1 21-227.

Grant, V. 1992. Systematics and phylogeny of the Ipomopsis aggregata group (Polemoniaceae): traditional and molecu-

lar approaches. Syst. Bot. 1 7:683-691 .

Harrod, RJ., LA. Malmquist, and R.L Carr. 1999. A review of the taxonomic status of Hackelia venusta (Boraginaceae).

Rhodora 101(905):16-27.

Hipkins, V.D., B.L. Wilson, RJ. Harrod, and C. Aubry. 2003. Isozyme variation in showy stickseed, a Washington endemic

plant, and relatives. N.W. Sci. 77:1 70-1 77.

Hitchcock, C.L., A. Cronquist, M. Ownbey, and J.W. Thompson. 1 959. Vascular plants of the Pacific Northwest Part 4: Ericaceae

through Campanulaceae. University of Washington Press, Seattle, WA.

Mabberley, DJ. 1987. The plant-book. Cambridge Press, Boston MA.

Winston J.E. 1999. Describing species. Columbia University Press, New York.

BOOK REVIEW

Werner Kunz. 2012. Do Species Exist? Principles of Taxonomic Classification. (ISBN-13: 978-3-527-33207-

6, hbk.). Wiley-Blackwell, 111 River Street, Hoboken, New Jersey 07030-5774, U.SA. (Orders: www.

wiley.com, 1-877-762-2974). $99.95, 280 pp., 6 W x 9

mirror in which to observe the fundamental differences among living things, resting as it does at the apex of

several shifting layers of DNA, epigenetics and environmentally- influenced gene expression patterns. And yet,

it is at the level of morphology, of phenotype, that influenced gene expression patterns. And yet, it is at the level

of morphology, of phenotype, that organisms meet the outside world in which their fitness is measured by na-

ture, red in tooth and claw. Morphology, in other words, is not an abstraction or obstruction to understanding

“true” relationships, but rather an essential part of classification.

The lure of the phylocode lies in the promise of certainty and objectivity. A DNA sequence is an objec-

tively observable truth, and the processes by which one sequence may mutate into another are reasonably well

described. The process of phylogenetics is not at issue, but rather the meaningfulness of the results of that

The meaningfulness criterion is simply that the genes and sequences currently used for phylogeny have at

best a tenuous connection to the fitness of an organism or even its habitat or ecological role. In other words,

leasure the distance between two organisms, but cannot speculate on the significance of

As a case in point from the world of microbiology, consider the humble Escherichia coli bacterium. Mea-

sured by the standards of molecular taxonomists, E. coli strain H7:OI57 and E. coli strain K12 are identical.

When mixed with hamburger meat, however, the H7:0157 strain can cause kidney failure and death while K12

will pass through the digestive tract without a trace. It is not the molecular taxonomists who have elucidated

these strains that have >99.9% identity in 16S rDNA sequence, but pathologists treating lethal food poisoning

outbreaks. In other words, those most concerned with life-as-it-is have identified the critical cell-surface anti-

gens overlooked by molecular taxonomists.

The issue with phylogenetics as presented by Kunz, in an admittedly brief treatment, is not that it fails at

mathematical consistency or clearly discernible taxa. It does. It is the relevance of those distinctions that is in

question and that he does not address.

The final substantive criticism of Kunz book lies with its questionable premise: that there exists a logi-

cally consistent means of classifying the elements of the natural world. In this quest one can hear the echoes of

Plato’s insistence on Forms. Rather than a perfect Form of an individual species, however, we have moved to

the perfect Form of taxonomy. We are not quite able to see it, but we know it exists. Each new generation of

mathematical algorithm refines the closest fit between our approximations and the perfect reality.

And yet, it may be that taxonomy is an inherently messy process. Life, after all, is inherently messy. There

are as many lineages stretching back to the Last Universal Common Ancestor (LUCA) as there are organisms

currently living. Each path is necessarily unique, and it may be that drawing lines around any subset is an ex-

minds does not necessitate tl

Rather it may be that t

nlydis

UNA NUEVA ESPECIE DE SOBRALIA (ORCHIDACEAE) DE EL SALVADOR

Jose L. Linares

Herbario CURLA

Orquldeas de El Salvador aparece un

taciones actuales de dicha entidad;

citada y llamada asi en todos los trat

pecies y una variedad de Sobralia Ruiz & Pav. (Hamer 1974, 1981). La

layores problemas, pues las especies son claramente diferentes entre

este genero para el pais fue el realizado por Hamer (1974, 1981) y ha

limiento del genero desde entonces. Entre las especies tratadas en las

ya description, distribution y ecologla no concuerdan con las delimi-

ita de la especie referida como Sobralia macro Schltr. en la obra pre-

ntos y trabajos subsiguientes relacionados con la flora de El Salvador.

era fue asignado sin objeciones a las plantas provenientes de la parte

suroccidental de El Salvador, que no encajaban ni en Sobralia macrantha Lindl. (y especies alines) ni en Sobralia

decora Bateman, este ultimo nombre asignado a las plantas de las zonas mis bajas. Sin embargo, los conoce-

dores y estudiosos del genero, como el Dr. Robert L. Dressier, identificaron los ejemplares de herbario depos-

itados en herbarios de Estados Unidos como pertenecientes a Sobralia leucoxantha Rchb.f., basandose en el

color mucho mas pilido de las flores, citado en las etiquetas como morado, violeta o rosado palido. A1 revisar el

material y la literatura referida a Sobralia macro, es notorio que esta especie tiene flores casi completamente

blancas y crece en lugares mucho mas humedos, ecolbgicamente distintos y se encuentra bastante alejado de

las localidades de El Salvador. Despues de una revision detallada de las posibles especies relacionadas presen-

tes en el pals o en palses cercanos, como Sobralia blancoi Dressier & Pupulin, Sobralia leucoxantha, Sobralia

macro, Sobralia macrantha, Sobralia pendula Dressier & Pupulin y Sobralia rogersiana Christenson y otras del

complejo Sobralia leucoxantha o cercanamente relacionadas (Christenson 2007; Dressier 2012; Dressier y Pu-

pulin 2008, 2012) se concluye que las plantas encontradas en El Salvador corresponden a una especie no de-

Sobralia paulancalmoi ] Linares, sp. nov. (Figs. 1-6). Tiro: ELS

Plantas cespitosas con 3-7(-9) tallos, las plantas adultas de aspecto desalinado, con 1-2 tallos florlferos. Tallos

delgados en relation con su longitud, por lo que se vuelven ligeramente arqueados (semierectos), cubiertos de

Linares, Sobraiia paulancalmoi, una nueva especie

ciaramente convexo; bas

mnade 4.7-4.8x0.6-07

curvada y con dos cornu

e dulces. Capsula no vi

661

rosada, la garganta rosado palido a casi

parte mas ancha y de 0.5 cm en su parte

omiculos falcados y triangulares en su porcibn distal,

tinte rosado a purpura en los comiculos, especialmente

te y algo desagradable, parecida a la del insecto del Or-

miiniscencias del olor de las bracteas de la inflorescen-

den Blattodea

E stado de Conservation. — de acuerdo con los criterios de la Lista Roja de Especies Amenazadas (UICN

2012) y segun las observaciones del autor de las unicas son subpoblaciones conocidas en la area de distribucion

de la especie, la categoria seria Vulnerable, VU Al(d).

Distribution, hdbitat yfenologla. — Conocida solo de la parte suroccidental de El Salvador (Fig. 1), en la

vertiente norte del macizo montanoso de El Imposible, en el ecosistema bosque tropical semideciduo latifolia-

do submontano, bien drenado, secundario y/o intervenido (WICE 2012) donde crece terrestre en riscos roco-

sos azotados por el viento, en lugares con acumulacion de humus. Entre la flora observada en la localidad ten-

emos arbustos pequenos de Dalbergia calycina Benth. (Leguminosae), Rapanea sp. (Myrsinaceae), Schoepfia

vacciniiflora Planch, ex Hemsl. (Olacaceae), Rondeletia lanijlora Benth. (Rubiaceae), Berberis johnstonii Standi.

& Steyerm. (Berberidaceae) y hierbas como Lasiacis sp. (Poaceae), Zeugites americanus var. mexicanus (Kunth)

McVaugh (Poaceae), Perezia sp. (Asteraceae). Entre las orquideas encontradas en esas localidades se encuen-

tran Corymborkis forcipigera (Rchb.f. & Warsz.) L.O. Williams, Epidendrum ciliare L., Epidendrum trianthum

Schltr., Maxillariella variabilis (Bateman ex Lindl.) M.A. Blanco & Camevali, Oncidium sotoanum subsp. pa-

palosmum R. Jimenez, Platystele ovalifolia (F. Focke) Garay & Dunst., Polystachia foliosa (Hook.) Rchb.f. y Stan-

hopea saccata Bateman

iero (en cultivo en San Salvador). Aparentemente no es de floracion gregaria, pues floi

l forma irregular, sin florecer todas las plantas de la especie al mismo tiempo. Fru<

Eponimia. — Es un honor dedicar esta especie a Paul Ancalmo (1948-), destacado y acucioso cultivador de

las orquideas de El Salvador, especialmente interesado en la conservacion y cultivo del genero Sobraiia, tan

descuidado y poco entendido en el pais. Gracias a los esfuerzos de Paul, ahora podemos decir que entendemos

mucho mejor las especies salvadorenas de este genero.

DiscusiOn. — Los rasgos mas caracteristicos de esta especie son su habito terrestre (o raramente litofitico),

el crecimiento vegetativo con tallos casi sin hojas en el habitat natural (Fig. 2), llegandose a observar ejemplares

casi completamente defoliados o con sblo dos hojas; la ausencia de crecimientos laterales (hijos o keikis); las

hojas usualmente de menos de 4 cm de ancho, versus hojas de mas de 4 cm de ancho en Sobraiia macro ; ademas,

las hojas de S. macra son eliptico-ovadas y acuminadas y la de S. paulancalmoi son angostasmente elipticas y

largamente acuminadas (Fig. 3); las partes florales de S. paulancalmoi son mas grandes que las de S. macra,

Uegando el sepalo dorsal a medir hasta 6.7 cm de largo contra solo 6.2 cm en S. macra; los sepalos laterales

pueden Uegar a medir 8 cm de largo mientras que en S. macra no pasan de 6.5 cm; por otra parte el labelo en S.

paulancalmoi puede Uegar a medir hasta 8.8 cm y en S. macra no pasa de 6.8 cm de largo; el habito de las flores

de abrir dos dias seguidos, el olor fuerte y la coloracion distinta de las flores, las cuales son rosado paUdo a algo

Las flores que duran, al menos en cultivo, mas de un dia, aunadas a la morfologia, la ubican ciaramente en

el complejo de Sobraiia leucoxantha (Dressier y Pupulin 2008, 2012), k que hizo que fuera confundida con esa

Journal of the Botanical Research Institute of Texas 7(2)

Fig. 6. Flordepe

REFERENCES

Christenson, E.A. 2007. Notes on the Sobralia macrantha complex. Orchideen J. 4:1 59-165.

Dressler, R.L. 2009. Can Sobralias be classified? Orchids 78:658-663.

Dressler, R.L. 2012. Sobralia decora. Orchids 81 :308-310.

Dressler, R.L. y F. Pupuun. 2008. La identidad de Sobralia leucoxantha, con tres especies m

lejana. Orquideologia 25(2):1 34-1 51 .

Dressler, R.L. y F. Pupulin. 2012. Una correccion al complejo Sobralia leucoxantha, con tres especies nuevas, dos muy

afines y una mas lejana. Orquideologia 29(1):28-30.

Hamer, F. 1974. Las Orquideas de El Salvador 2: 288, 289. Ministerio de Educacion, Direccion de Publicaciones, San Sal-

vador, El Salvador.

Hamer, F. 1 981 . Las Orquideas de El Salvador, Tomo III. Marie Selby Botanical Gardens, Sarasota, Florida, U.S.A„

UICN. 2012. Categories y Criterios de la Lista Roja de la UICN: Versibn 3.1 Segunda edicion. Gland, Suiza y Cambridge,

Reino Unido: UICN. Vi + 34 pp. Originalmente publicado como IUCN Red List Categories and Criteria: Version 3.1

Second edition. Gland, Switzerland and Cambridge, UK.

WICE (world iNsrmjn for conservation & environment). 201 2. Mapa de ecosistemas de El Salvador actualizacion 201 0 y mapa

tebrico de ecosistemas originales de El Salvador. Serie del Estudio de Racionalizacion y Priorizacibn del Sistema de

Areas Naturales Protegidas de la Republica de El Salvador, San Salvador.

journal of the Botanical Research Institute of Texas 7(2)

BOOK REVIEW

Richard Stephen Felger and Benjamin Theodore Wilder (in collaboration with Humberto Romero-Morales).

2012. Plant Life of a Desert Archipelago: Flora of the Sonoran Islands in the Gulf of California.

(ISBN-13: 978-0-8165-0243-1, hbk.). The University of Arizona Press, P.O. Box 210055, Tucson, Arizona

85721-0055, U.S.A. (Orders: uapress.arizona.edu, 1-800-621-2736). $65.00, 624 pp. (incl. 8-page color

inset), 388 species accounts with paired distribution maps, hundreds of line drawings and photos,

8 %" x 11", 3.7 pounds.

The natural history of this region is natural history brought to life and almost animated for you in this superb

compendium of information on the Gulf Islands. Anyone working in this region, in any rlicriplinp from begin-

ner to expert, will be dehghted by the wealth of detailed information written with heart, cheer, and energy,

lighting up the rich human and biological history of this very special region of the world.

This book is packed with rich tales of an island chain almost untouched by invasive plants, with a modern

history of many peoples sharing the rich biological resources. Almost every page has a picture, and albeit in

black and white, the images and the diverse objects throughout the book leave you yearning to give them color

and visit the islands for yourself.

Perhaps the most outstanding feature of the book is the quality of the science behind it, the precision in

every story and record and every dot on each map. The authors are meticulous scholars, and every detail has

been carefully researched, yet this in no way detracts from the readability. This book is a masterpiece in its

ability to be both a thoroughly enjoyable read and a scientific encyclopedia.

Felger took his first tripan 1954, and anyone familiar with Felger’s attention to detail will instantly know

the quality of this work. Wilder became involved a half-century later in 2005, and his focused studies in recent

years bring a contemporary energy to this mammoth volume.

The preface and foreword are both passionately written and convey the very personal connections of the

authors to the region. Part 1— The Islands and Their Vegetation is an introduction to the geology, biology,

ethnobotany, and natural history. It includes a section on invasive species and a section on administration and

conservation that boasts a photo of big-horn sheep being moved by helicopter in 2008! Part 2— Botanical

Explorations on the Sonoran Islands: Collectors, Associates, and Selected Personalities is a delightful

compendium of accounts documenting people who have lived and worked in the region. It is packed with tales

of adventure and travels in the gulf, giving rich insights into the lives of the big names in gulf biology. Part

3— The Flora contains the species accounts, the heart of the research. Each species has a distribution map, and

there are many great illustrations, packed with stellar detailed-yet-easily-digestible text. It includes keys to

species and the Seri plant names and uses in many cases, often provided by collaborator Humberto Romero-

Morales of the Seri nation. And anyone working in the region or researching its history will enjoy the detailed

place names in Part 4— The Gazetteer as well as the following useful appendices: Appendix A: Checklist of

the Flora of the Sonoran Islands (summarizing knowledge for each island); 1

Absent from Isla Tiburon and Mainland Sonora (an unusual and interesting addit

a watch-list for what might arrive or be found in future); an

the index and literature cited are exhaustive and excellent.

This book is a must-have for anyone working in the area, but it’s also a treasure for anyone that just wishes

they were there, a doorway into the magic of the region through the pages of a book.— Sula Vanderplank, PhD,

Biodiversity Explorer, Botanical Research Institute of Texas, Fort Worth, Texas, U.S.A.

J.Bot Res. Inst Texas 7(2):«

FOUR NEW SPECIES OF COLUMNEA (GESNERIACEAE)

WITH PRIMARY DISTRIBUTIONS IN COLOMBIA

James F. Smith Marisol Amaya-Marquez & Oscar H. Marm-Gomez

Department of Biological Sciences

Boise State University

1910 University Drive

Boise, Idaho 83725- 1515, U.S.A

jfsmith@boisestate.edu

Instituto de Ciencias Naturales

Universidad Nacional de Colombia

Apartado 7495

Bogotd, COLOMBIA

Department of Biological Sciences

The University of Alabama

Box 870345

ABSTRACT

RESUMEN

INTRODUCTION

Within Gesneriaceae, classification systems based on morphological characters have been notoriously chal-

lenging when trying to align them with evolutionary relationships (Wiehler 1983). Recent molecular work has

uncovered multiple cases of para- and polyphyletic genera (Moller & Cronk 1997; Smith et al. 1998, 2004;

Smith 2000; Clark & Zimmer 2003; Roalson et al. 2005a, 2005b, 2008; Clark et al. 2006, 2011, 2012; Moller et

al. 2009; Wang et al. 2010, 2011). Although the genus Columnea has been regularly recovered as monophyletic

in molecular systematics studies (Clark et al. 2012), its classification has not been entirely stable as Wiehler

(1973, 1983) split the genus into four genera (Columnea, Dalbergaria, Trichantha , and Pentadenia) and de-

scribed a fifth, Bucinellina (Wiehler 1981). Wiehler considered earlier classifications within Columnea si to

reflect convergent evolution by pollinator selection on corolla form rather than the evolution of the plants

themselves (Wiehler 1983). This split was not widely accepted and alternative subgeneric systems that treated

Columnea as a single genus were proposed (Morley 1974, 1976; Kvist & Skog 1993; Smith 1994). Currently the

genus is estimated to comprise 200 species (Kvist & Skog 1993; Weber 2004; Skog & Boggan 2007).

As part of the process of revising the subgeneric classification and placement of species in section Or-

tholoma (sensu Kvist & Skog 1993; Smith 1994), several specimens have been discovered that do not fit within

the range of variation for the previously described species in this section.

Colombia is home to tremendous species diversity in the genus Columnea with 90 species recorded there

(Clavijo et al. 2011) and numerous recent papers reflect the large number of species yet to be described from

this country (Amaya-Marquez 2010a, b; Amaya-Marquez & Marin-G6mez 2012; Amaya-Marquez & Smith

2012; Clark & Clavijo 2012; Clavijo & Clark 2013; Mora & Clark 2012; Amaya-Marquez & Smith submitted).

668

Here we describe four species

vhose primary distribution is in Colombia.

Columnea ceticeps J.L. Clark & J.F. Smith, sp. nov. (Figs. 1 & 2).

n, 5°29'18"N 75°54'20"W, 1'

Epiphytic herb; stems to 2.8 mm in diameter, red-brown, with zigzag appearance

tally appressed pilose with multicellular gold-colored trichomes; intemodes 1.3-4.8 cm long; leaf scars flush

with the stem. Leaves opposite, anisophyllous, larger lamina 2.5-8.8 cm long, 0.8-3.0 cm wide, ovate to el-

liptic, apex acuminate, base oblique, lateral veins 3-5 per side, adaxially green, appressed pilose with multicel-

lular trichomes with more or less pustulate bases, abaxially green, pilose with gold-colored multicellular tri-

chomes, denser on veins, margin crenulate to serrulate; petioles 0.1-0.15 cm long, pilose with multicellular

gold-colored trichomes; smaller lamina 1.7-2.5 cm long, 0.7-1.15 cm wide, lateral veins 1-3 per side, petiole

0-0.08 cm long, otherwise similar to larger lamina. Inflorescence of 1 flower per axil of leaf; bracts not seen,

presumably caducous. Pedicels 9.5-19.5 mm long, green, appressed pilose with multicellular trichomes. Calyx

loosely clasping, lobes 14.0-21.0 mm long, 0.8-3.5 mm wide, lanceolate, apex acute, exterior appressed pilose

with multicellular gold-colored trichomes and single-celled white trichomes or densely spreading pilose with

multicellular gold-colored trichomes (the latter true for most specimens from Narino), red, interior glabrous;

margin subentire to denticulate. Corolla 4.2-6.5 cm long, 0.55-1.4 cm at widest point which is the opening of

the throat, tubular, not ventricose, gibbous at base, 0.25-0.35 cm wide at narrowest point at the base, red to

orange, exterior pilose with multicellular red-colored trichomes, interior minutely puberulent; limb bilabiate,

upper lip formed by the two dorsal and two lateral lobes, lower lip formed by a ventral lobe; dorsal lobes con-

nate, rounded, 0.25-0.55 cm long, 0.5-0.6 cm wide, lateral lobes acute 0.1-0.2 cm long, 0.18-0.3 cm wide,

ventral lobe, oblong 1.05-1.8 cm long and 0.14 cm wide, galea 1.8-2.05 cm long. Filaments 1.5 cm long connate

0.25 cm and adnate to corolla another 0.15 cm, tomentose with glandular and non-glandular trichomes, an-

thers 2.5 mm long, 2.5 mm wide, quadrangular, included in corolla throat. Ovary 4.0 mm long, conical, gla-

brous, style 34 mm long yellow, minutely puberulent, stigma stomatomorphic, yellow, smooth. Nectary a

double dorsal gland. Fruit and seeds not seen.

Phenology.— Flowering specimens have been collected from December-February, April, May, and July,

presumably flowering continuously, no fruiting specimens are known.

Distribution.— This species is widespread mainly in the Cordillera Occidental of Colombia in the de-

partments of Antioquia, Chocb, Risaralda, Valle, Cauca, Narino, Putumayo, and Ecuador at elevations from

1900 to 2900 m. In Antioquia the species has been recorded both in Cordillera Occidental and Cordillera

Central.

Etymology .— The specific epithet is derived from the combining forms of whale (cetus) and headed (-ceps)

due to the similarity of the corolla in profile that looks like a sperm whale’s head with an open mouth (Fig. 2).

Cordillera Central, 12 May 2012 J.L. Clark et al.^^S)L); jardin, Sector alto de Ventanas,

San Jo^ del Palmar, Cerro del Torra, edge of the mountain, 21 Aug 1988, F.A. Silverstone-Sopkin et al 4641 (CUVC). Risaralda- Santuario

vereda las Colonias, 2 Feb 1983 ,J.H. Torres et al. 1446 (COL). Cauca: Cerro Munchique, F.C. Lehmann 7636 (US); El Tambo, PNN Munch-

ique, 20 Jun 2001, R. Bernal & P. Lopera 2830 (COL), 8 Feb 2000, C.E. Gonzdlez & M.M. Olives 2702 (COL), 30 Jul 1993, G. Lozano et al. 6587

(COL); El Tambito, Centro de Estudios Ambientales del Padfico Tambtto, trail Cerro del Perro, Rio Paloverde, 19 Dec 2000, C.E. Gonzdlez

3354 (COL); La Costa, 7 Sep 1936, K. von Sneidem 795 (COL), Sep 1936, K. von Sneidem 796 (COL); El Tambo, 4 Aug 1936 K. von Sne idem 990

(US), 19 Jul 1993, F. Gonzalez et al. 2756 (COL); 7 “La Gallera” El Micay 1 Jul 1922 E.P. Killip 7935 (US), Perez Arbelae z & Cuatrecasas 6266

(COL). A. Gentry et al. 60502 (MO), 30 Jul 1993, N. Ruiz et al. 204 (COL); El Cairo, vereda El Brillante, 30 May 2011 O -H Martn-GOmez &

D.A. Gdmez-Hoyos 196 (COL-2), 197 (COL), 198 (COL). Nariiio: Ricaurte, Reserva La Planada, 26 May 1985 O. de Benavides 5553 (US PSO),

29 Apr 1988 O. de Benavides 9598 (MO, US, PSO), 17 Jan 1990 O. de Benavides 11276 (US, PSO), 25 Sep 1989, 0. de Benavides 10809 (PSO), 1

Sep 1990, 0. de Benavides 11447 (PSO); Mpio. de Mallama, camino que conduce de la Planada a Pialapi, 15 Aug 1992, R. Giraldo 181 (PSO) 1 3

May 1992, R. Giraldo 121 (PSO), 3 Mar 1989J.F. Smith &M. Galeano 1522 (PSO); Barbacoas, Corregimiento de Altaquer, vereda El Barro, Mar

670

Journal of the Botanical Research Institute of Texas 7(2)

Smith et al., New species of Columnea in Colombia

Several herbarium specimens were annotated with unpublished names by Wiehler. It is curious that Wiehler

considered this species as a member of Trichantha, which is a genus that has few species with bilabiate corollas.

This species might more likely be placed in section Columnea , and corresponding to Wiehler’s genus Columnea

based on the bilabiate non-ventricose corolla with spreading corolla lobes. However, the corolla does not ex-

pand to a broadened galea (Fig. 2 A, B) as do all of the species in section Columnea and its sectional placement

has yet to be tested with molecular data.

The specimens from Narifio, Colombia are marginally different from other collections in that the exterior

vesture of the calyx lobes is densely pilose with the characteristic gold-colored trichomes found in other collec-

tions but much less dense. This species is similar to another undescribed species from Colombia and which

difference, pubescence on vegetative parts of species in Columnea tends to be consistent and glabrous leaves are

an unusual trait, therefore these specimens are considered a different species here.

Columnea ceticeps also is vegetatively similar to yet another undescribed species from Colombia, both of

which have been collected from Cerro Munchique. However the corollas are different between the two with the

undescribed species having corollas with a subradial limb and C. ceticeps with a bilabiate limb (Figs. 1, 2).

Columnea ferrugineaJ.F. Smith &J.L. Clark, sp. nov. (Fig. 3). Type: COLOMBIA. Valle: Municipio El Cairo, Corregimien-

Epiphytic herb; stems to 115 cm long, up to 5.5 mm diameter, brown, proximally pilose with multicellular

phyllous, larger lamina 6.2-10.0 cm long, 1. 8-3.1 cm wide, oblanceolate to narrowly oblong, slightly falcate,

apex acuminate, base oblique, lateral veins 9-12 per side, adaxially green, appressed pilose with multicellular

rust-colored trichom es , abaxially green, pilose with multicellular rust-colored trichomes, slightly denser ves-

ture on veins, margin serrulate and ciliate with multicellular red trichomes; petioles 0.2-03 cm long, densely

pilose with multicellular rust-colored trichomes, smaller lamina 1.2-2.1 cm long, 0.25-0.6 cm wide, lanceo-

late, apex acute to acuminate, base oblique, adaxially green, pilose with multicellular rust-colored trichomes,

abaxially green, densely pilose with multicellular rust-colored trichomes, denser on veins, margin serrulate,

petiole 0.08-0.1 cm long, pilose with multicellular rust-colored trichomes. Inflorescence of 1 flower per axil

of leaf; bracts 3.5 mm long, 0.2 mm wide, linear, apex acute, red-purple, pilose with multicellular rust-colored

trichomes. Pedicels 1.9-2.5 cm long, green or red, densely pilose with multicellular rust-colored trichomes.

Calyx loosely clasping, lobes 16.0-27.0 mm long, 1.5-1.8 mm wide, linear, apex acuminate, exterior densely

pilose with multicellular rust-colored trichomes, red or bright yellow-green; margin serrulate. Corolla 4.7-63

cm long, 0.95-1.4 cm wide at widest point, 0.9-1.25 cm before limb, 0.25-03 cm at gibbous base, tubular,

slightly ventricose, red, exterior sparsely pilose with multicellular rust-colored trichomes, interior with some

short trichomes and some stalked glandular trichomes; limb 0.95-33 cm in diameter (widest when lower lobe

is reflexed), lobes semi-orbicular to linear, 0.25-1.2 cm long, 0.25-035 cm wide, red. Filaments connate 3.5

mm and adnate to corolla 0.7 mm, with short stalked glandular trichomes, anthers 2.8 mm long, 2.8 mm wide,

quadrangular, included in corolla tube. Ovary 4.5 mm long, conical, pilose with multicellular transparent tri-

chomes, style pale yellow to white, pilose with multicellular transparent and short stalked glandular tri-

672

Journal of the Botanical Research Institute of Texas 7(2)

Smith et al., New species of Columnea in Colombia

673

chomes, stigma bilobed, papillate, included in corolla tube. Nectary a dorsal double gland. Fruit and seeds not

Phenology. — Flowering only known from January and May, fruits not seen.

Distribution. — Apparently a rare species found in Colombia between 2000-2320 m along the western

slopes of the Cordillera Occidental.

Etymology. — The specific epithet is derived from the rust-colored trichomes that cover most of the vegeta-

tive portions of this species.

Additional specimens examined: COLOMBIA. Valfc: Municipio El Cairo, Serranla de los Paraguas, Reserva Natural Cerro El Ingles, 1 Jan

1987, F.A. Siherstone-Sopkin 2850 (US); Corregimiento Boquerbn, Vereda La Amarillas, Serrania de los Paraguas, Cerro El Ingles, 14 May

1988 , J.L. Luteyn et al. 12346 (NY, US), 19 May 2013 J.F. Smith et al. 10772 (COL); Sector La Florida, camino Los Santicos a La Florida a Rio

Blanco, 20 May 2013, J.F. Smith et al. 10829 (COL), J.F. Smith et al. 10830 (COL).

Columneaferruginea is readily differentiated from all other congeners by the distinctive rust-colored vesture on

the abaxial leaf surface. It is similar to C. dictyophylla Donn. Sm. in that it also has a large, red, bilabiate corolla,

but differs in lacking the distinctive gold vesture of the abaxial leaf surface. Additionally, the tertiary leaf vena-

tion in C. dictyophylla is reticulate, abaxially prominent, and covered in hairs that contrast with the often red-

purple color of the lamina. The tertiary venation of C.ferruginea is suppressed and inconspicuous.

Columnea fractiflexa J.F. Smith & J.L. Clark, sp. nov. (Figs. 4 & 5). Type: Colombia. Antioquia: Municipio Urrao, Cor-

confluencia del rio Polo y antes del rfo San Pedro, sitio la Quiebra, 06°30’31”N, 76°14'W, 1600-1850 m, 31 Jan-2 Feb 2011, P. Pedra-

za-Penalosa,J. Betancur, M.F. Gonzdlez, G. Giraldo, F. Gdmez, A. Duque&J. Sema2137 (holotype: COL; isotypes: NY, UNA).

tween the corolla lobes.

Pendent to festooning epiphytic herb; stems to 1.5 mm diameter with a characteristic zigzag appearance, tan,

proximally sparsely appressed pilose with multicellular red-colored trichomes, distally denser; internodes

2.0-2.2 cm long; leaf scars slightly raised. Leaves opposite, anisophyllous, larger lamina 2.8-4.5 cm long,

0.85-2.8 cm wide, ovate, apex acute, base obtuse and slightly oblique, lateral veins 4-7 per side, adaxially red,

densely appressed pilose with multicellular red-colored trichomes, abaxially red, sparsely appressed to spread-

ing pilose with multicellular yellowish trichomes, denser on veins, margin serrulate; petioles 0.1-0.15 cm long,

pilose with multicellular red-colored trichomes, smaller lamina 0.6 cm long, 0.2 cm wide, lanceolate, sessile,

otherwise s imilar to larger lamina. Inflorescence of 1-2 flowers per axil of leaf; bracts 8.0 mm long, 1.5 mm

wide, lanceolate, apex acute, green, sparsely pilose with multicellular red-colored trichomes. Pedicels 8.0-11.0

mm long, green, densely appressed to spreading pilose with multicellular red-colored trichomes. Calyx clasp-

ing, lobes 7.0-8.0 mm long, 8.0-10.0 mm wide including lobes to the tips of the lobes, ovate, apex acuminate,

exterior spreading pilose with multicellular red-colored trichomes, green, interior glabrous, margin deeply

fimbriate. Corolla 4.4-5.1 cm long, 0.74-1.0 cm wide at widest point, 0.6-0.95 cm before limb, 0.3-0.35 cm at

base, tubular, slightly ventricose, somewhat falcate, gibbous at base, red, exterior sparsely spreading pilose

with multicellular trichomes, interior sparsely pilose inside throat dorsally, otherwise glabrous; limb slightly

bilabiate, 0.85-1.22 cm in diameter, lobes elliptic, 0.45 cm long, 0.2 cm wide, red. Filaments 47 mm long con-

nate for 8.0 mm and adnate to corolla an additional 0.5 mm, glabrous, anthers 1.7 mm long, 1.4 mm wide,

quadrangular, included in corolla tube. Ovary 3.0 mm long, conical, sparsely pilose with multicellular tri-

chomes, style 49 mm long, yellow, glabrous, stigma bilobed, papillate, included in corolla tube. Nectary two

dorsal double glands. Fruit and seeds not seen, but the label of the type collection indicates the fruits are

Phenology. — Flowering known from January, April, May, July, and November. Fruits not seen, but are re-

ferred to in the type collection made in January.

Distribution.— This species is known only from a narrow region in Antioquia Colombia where all speci-

mens have been collected from 1600—2050 m.

Etymology. — The specific epithet is derived from the zigzag appearance o

af the pendent stems (Fig. 5B).

Journal of the Botanical Research Institute of Texas 7(2)

0.3 cm

Smith et al.. New species of Columnea i

675

676

: of tubular subradial red corollas and pendent

s afractiflexa

Columneafractiflexa is similar to C. minor by the presenc

tooning habit (Figs. 4, 5). The leaves of Columned fractifL

5 cm). The presence of corolla appendages alternate between the corolla lobes

C.fractiflexa. Other species that have pendent habits are mostly confined to a section of Coll

Columned) that is characterized by strongly bilabiate corollas. The subradial corolla of Colun

(Fig. 5A) is different from the strongly bilabiate corollas in section Columned.

- Clark & M. Amaya, sp. nov. (Fig. 6). Type. COLOMBIA. Val

erva Natural Cerro El Ingles, 19 May 2013 J. F. Smith, O.

C.PSO.TULV, VALLE).

ate-ladniate cal^^r^ns.MfersfeKnLbhnnneajW^ia^a^ 15 ^ ^ *° ^°^ mnea ^ m ^ rica ^ x ^ ^ presence of fimbri-

EpipkyOc herb; stems to 50 cm, up to 3.5 mm diameter, brown, woody at base of stems, proximally nearly

glabrous to sparsely pilose with multicellular transparent trichomes, distally denser with multicellular trans-

parent and red-colored trichomes; intemodes 1.5-7.2 cm long; leaf scars raised. Leaves opposite anisophyl-

lous, appearing alternate as the smaller leaf is often lost on older portions of stems, larger lamina3.2-6.8 cm

long, 1.6 3.5 cm wide, ovate to elliptic, apex acuminate, base oblique, lateral veins 4-7 per side adaxially

green, sparsely pilose with multicellular transparent trichomes, abaxially red-purple, sparsely pilose with

mukiceUular transparent trichomes to nearly glabrous, slightly denser vesture on veins, margin entire; petioles

0 ^ whfe, <wale to lanceolate, apex acuminate, base oblhpieJatenilveins4-45 ^skk,^adarialtygreem

pilose with mult, cellular transparent trichomes to nearly glabrous, abaxially red-purple, sparsely pilose with

multicellular transparent trichomes to nearly glabrous, margin entire to crenulate, petiole 0.3-0 45 cm long,

apprised pilose with multicellular red-colored trichomes. inflorescence of 1-2 flowers per axil ofleaf; bracts

^ T, mm Wide ’ linMr - a[KX acute ' red -P u T*. with multicellular transparent tri-

ntTme r'l 5 1 °" 8 1° T ' 0ng ' :t PUrPl£ ‘ aPPreSS ' d PllOSe WUh mul,l “ llular ™sparen, or red-colored

reidimnes. Calyx loosely clasp, ng, lobes 14.0-20.0 mm long, 1.0-1.5 mm wide without laciniae, elliptic to

. ' , apeX aCUle ’ lnte "° r glabrous, exterior pilose with multicellular transparent or red-colored tri-

chomes, red-purple; margin laciniate. Corolla 2.3-3.6 cm long, 0.35-0.45 cm wide at widest point alone the

:^; 0 ' 5 T^ fore a Umb L a2cmatgibW

02-03 cm long oi *0 3 tnC ° m “ - ,r len0r g ' abrous; llmb 0.55-0.7 cm in diameter, lobes semi-orbicular.

7 Fib r ntS “ nna ' e5 5m m and adnate to cordU 0.5 mm, glabrous,

with muhLdluirnranspTrenrt^'ho^e^slyhTellow^Uib' 11 C ° r ^^ mm *° n ®’ con ' ca l> P'l° se

tube. Nec, ary nvodomaldtiuble glands. FmitJdsee^ no, se™ 181 ” 3 ^ bdobed-pap. lUte included in corolla

EWstributT Fl ° wer * ns °"/y ^"“w" from September, December, February to May, fruits not seen.

asmbuhon.-A rare spec.es from Colombia anjl bordering northern Ecuador between 800-2430 m

^he specific epithet is derived from the laciniate margins of the calyx lobes

P J dCl Palmar - Carmera Ah0 Gala P a « 0S a SanJos£ del Palmar, 22 May 2013, J.F. Smith et al 10875 (COL)

;b 1945J A - Ewan 16814 (MO). Valle: Municipio El Cairo, Serrania de Los Para^

S. Hoover ** 10768 (COL). ECUA-

lar 1991, A. Hirtz et al. 5253 (SEL).

Smith et ah, New species of Columnea in Colombia

677

Fie. 6. Columnea laciniata. A. Habit. B. Flower. C. Corolla opened to show androecium. D. Calyx and gyneocium with corolla removed. I. Enlarged view

of D showing the ovary and double dorsal nectary gland. All illustrations from J.A. Ewan 16814 (MO). Illustration by Sue Biackshear.

678

This species is most readily recognized by its corolla that is relatively elongate (to 3 cm) and densely covered

in trichomes (Fig. 6), but with deep serrations on the margins of the calyx (Fig. 6). This species is similar to

Columnea fimbricalyx L.P. Kvist & L.E. Skog in its laciniate calyx margins. However, the individual lobes are

ual calyx lobes difficult to distinguish in Columneafimbricalyx. It is also similar to other Columned species with

densely pubescent corollas but have leaves with crenate margins such as Columned purvifloru C.V. Morton, C.

lehmonii Mansf., and C. minutiflora L.P. Kvist & L.E. Skog. Columned Idcinidtd is distinguished from these by

the presence of laciniate calyx lobe margins (Fig. 6) in contrast to entire to shallowly serrate calyx margins. Ad-

ditionally this species is unique among species of Columned in that the leaf blades are nearly glabrous abaxially.

ACKNOWLEDGMENTS

We would like to thank the following herbaria for permitting loans and photography of their collections: COL,

MO, NY, SEL, and US. We are also grateful to Sue Blackshear for the illustrations. The authors thank the NSF

funded project: Flora of Us Orquideas National Park (DEB 1020623 to Pedraza), for supporting fieldwork and

specimen-based research that facilitated the discovery of Columned jractijlexa. A 2012 research expedition to

Colombia was a tremendous success because of logistical support from Alvaro Idarraga (HUA), Felipe Cardona

(HUA), Julio Betancur (COL), Alvaro Cogollo (JAUM), Laura Clavijo (UNA) and Diego Suescun (JAUM). We

also thank Gustavo Suarez and Luis Augusto Mazariegos-Urtado for facilitating an expedition to the Reserve

Mesenia-Paramillo and Uriel Rendon who was our guide and serves as the park guard for the Reserve Mesenia-

Paramillo. Fieldwork for the discovery of Columned ferrugined and C. lacinidtd was greatly facilitated by the

Organizacion Ambiental Comunitaria (SERRANIAGUA) and Johnier Arango Bermudez. Financial support for

this project was provided by NSF grants DEB-0949270 (to J.F. Smith and J.L. Clark) and DEB-0841958 (to J.L.

Clark) and the Marjorie Moore Davidson Foundation (to J.F. Smith). National University of Colombia sup-

ported MAM with time to do research. And finally, we thank Christian Feuillet (US) and Alain Chautems (G)

for their detailed and helpful reviews. The editorial staff at J. Bot. Res. Inst. Texas has been most supportive at

every step in this process and we thank them.

. . 3 (Gesneriaceae) de la Cordillera Oriental en los Andes Colom-

bianos. Caldasia 32:113-116.

W-Marquez, M. 2010b. Novedades taxonomicas en el genero Columned (Gesneriaceae). Rev Acad Colomb Cienc

34(132):301-307. t-Oiomb. Gene.

Vmaya-Mahchjez, m. O.H. Marin-gomez. 20,2. Column ea range* (Gestae), a new species from the Serrania de los

Paraguas in the Colombian Andes. Caldasia 34:69-74.

WMarquez IVUnd J.F. Smur 201 2. A rare new species of Columned (Gesneriaceae) from “Cordillera Occidental" in the

Colombian Andes. Rev. Acad. Colomb. Cienc 36:136-140.

r , * j CA , species of Gesneriaceae from Colombia. Caldasia.

Clark, J.L and EA Zimmer. 2003 A preliminary phylogeny of Alloplectus (Gesneriaceae): ir

flower resupination. Syst. Bot. 28:365-375.

Clark, J.L and L. Clavuo. 2012. Columned dntenniferd, a

v species of Gesneriaceae from tl

Colombian Andes. J. Bot. Res. Inst. Texas 6:385-390.

Clark, J.L, P.S. Herendeen, L.E. Skog, and EA. Zimmer. 2006. Phylogenetic relation:

Eoiscieae (Gesneriaceae) inferred from nuclear, chloroplast, and morphological data. Taxon 55:313-336.

J. Matos. 201 1 . Independent origin of radial floral sym-

oin Cuba.

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Syst Bot 36:757-767.

Clark, J.L., M.M. Funke, A.M. Duffy, and J.F. Smith. 201 2. Phylogeny of a

Clavuo, L and J.L Clark. 201 3. Drymonid crisp a (Gesneriaceae), a

10.1007/S12228-013-9310-4.

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„ Clark. 201 1 . Diversidad de Gesneriaceas de Colombia. Resum

/I Congreso Colombianc

Moller, M. and Q.C.B. Cronk. 1997. Phytogeny and disjunct distribution: evolution of Saintpaulia (Gesneriaceae). Proc. R

Moller, M., M. Pfosser, C.-G. Jang, V. Mayer, A. Clark, M.L. Hollingsworth, M.HJ. Barfuss, Y.-Z. Wang, M. Kiehn, and A. Weber.

ence with available tribal classifications. Amer. J. Bot. 96:989-1010.

Morley, B.D. 1 974. Notes on some critical characters in Columnea classification. Ann. Missouri Bot. Gard. 61:51 4-525.

Morley, B.D. 1 976. A key, typification and synonymy of the sections in the genus Columnea L. (Gesneriaceae). Contr. Nat.

Bot. Gard. Glasnevin 1:1-11.

Roalson, E.H., J.K. Boggan, and L.E. Skog. 2005a. Reorganization of tribal and generic boundaries in the Gloxinieae

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BOOK REVIEW

George Yatskievych. 2013. Steyennark’s Flora of Missouri , 1

deae) through Zygophyllaceae. (ISBN-13: 978-0-915279-13-5, hbk). Missouri Botanical Garden Press,

P.O. Box 299, Saint Louis, Missouri 63166-0299, U.S.A., in cooperation with the Missouri Department of

Conservation, P.O. Box 180, Jefferson City, Missouri 65201-0180, U.S.A. (Orders: www.mbgpress.info,

orders@mbgpress.org, 1-877-271-1930). $65.00, xvii + 1382 pp., 194 plates of b&w line drawings, 20

figures (incl. 27 b&w photographs), 798 distribution maps, 7" x 10" x 2.5".

At long last, the final treatment of the revision to Julian Steyermark’s (1963) Flora of Missouri has been completed by

the state’s premier botanist. Volume 3 is so well written, organized, and illustrated that it is hard to find fault. The late

Steyermark is someone the author has always looked up to, and a quote from Yatskievych’s acknowledgements is

worth repeating here. “Julian Steyermark is a model of what a botanist should be, and his high standards of scholar-

ship are something I continue to aspire to, but fear 1 will never reach.” In completing all three volumes of the revision

to Missouri’s flora, not only has Yatskievych reached Steyermark’s standards, he has exceeded them, and if Julian was

still with us I am sure he would concur. Volume 3 covers the Fabaceae (where Volume 2 left off) through the Zygo-

phyllaceae and includes treatments on 1,031 species, 65 infra-specific taxa, and 134 hybrids. All told, the entire

three-volume set is no less than 3,554 pages long and includes 582 plates and 2,726 county distribution maps involv-

ing 2,839 species and 3,166 taxa.

As in Volumes 1 and 2, one outstanding feature of Volume 3 is the thoroughness of the treatments, not only re-

value, chemical properties, medicinal value, and changes in distribution where applicable. There is only the need to

“• and poplar wood also was a popular material for stakes to strike through a vampire’s heart.” No one other than

George Yatskievych would digupsuch an interesting tidbit and include inaflora! The completeness of the treatments

is reflected in the fact that the author cited no less than 1,369 references!

Another highlight of the book is the contribution by 16 specialists and 10 separate illustrators. Despite the indi-

viduality of contributors, there is amazing continuity throughout the book. The excellent detail and enlargement of

flowers, fruits, and leaf vestiture, as well as illustrations of habit by all artists is certainly one of the noteworthy fea-

tures of the book. Another highlight is that some of the most difficult treatments were contributed by individuals

who are recognized experts on various taxa (e.g. Jay Reveill on various legumes, James B. Phipps on Crataegus, and

Mark P. Widrlechner on Rubus). Thus, it would be hard to argue taxonomy on some of the more controversial taxa.

Another positive point about the book is that the genus Rubus includes subgeneric and sectional keys using a combi-

nation of primocanes and floricanes as well as inflorescence characters. Similarly, keys to Lespedeza, Populus, and

Salix include both vegetative and reproductive chara ‘ ' •

his wit and sense of humor so the illustration of him

(plate 421) for scale is quite hilarious, and the illustrator John Myer should be c q _

of the drawing that includes Yatskievych in typical form with his beard, hat, bald head, glasses, hanging plant press,

and even hand clippers well depicted!

I suppose that all book reviewers are expected to find at least a few negative aspects for every book written es

pecially for one that is 1,382 pages in length, but I must confess that it was difficult to find anything to r ’l.f

about. Nonetheless, there are a few worth mentioning. The most obvious flaw is the fact that them is no family key

but simply the following statement on what would be page xviii; The key to dicot families will appear in a supple-

mentary publicanon." This is most unfortunate because no time table has been given when such a publication will

ever be completed. We can only hope that it is sooner rather than later, especially given so man taxonomic han es

in various dicot families. Another negative mark is that the key to legume subfamili Y aX ° n ° miC C angCS

Anyone needing to key out an unknown legume must potentially use both Vo

>t repeated in Volume 3.

NEW RECORDS AND NOTES ON SPECIES FROM PARC NATIONAL

PIC MACAYA, MASSIF DE LA HOTTE, HAITI, INCLUDING

A NEW SPECIES OF PILEA (URTICACEAE)

Department of Biology

University of Florida

Gainesville, Florida 32611 -8525, U.S.A

lmajure@ufl.edu

Department of Biology

Albion College

Albion, Michigan 49224, US.*

Gretchen M. lonta 1

Department of Biology

University of Florida

Gainesville, Florida 326 1 1-8525, USA.

Department of Biology

University of Florida

Gainesville, Florida 326 1 1-8525 , I

:ies nuevas de flora de la Reserva Bi6sfera Macaya (incluyendo Pan

s. La endtoica de La Espaiiola, Cyperus picardae, es reportada como

:eae), del Massif de la Hotte, se describe e ilustra.

Parc National Pic Macaya is located near Ville Formon in the Massif de la Hotte, ca. 36 km northwest of Les

Cayes, Haiti, encompassing ca. 2000 hectares (Judd 1987; Woods et al. 1992). The flora, plant communities,

land-use patterns, and history of biological exploration in the region are outlined in Judd (1987), Judd et al.

(1990, 1998), Woods and Ottenwalder (1992), Woods et al. (1992), and Sergile et al. (1992). The majority of

Haiti’s remaining undegraded natural forest lies within the boundaries of Parc National Pic Macaya and the

country’s other national park, Parc National Morne La Visite, in southeast Haiti. Parc National Pic Macaya

provides refuge for numerous endemic species (Woods et al. 1992, Rimmer et al. 2005, Huber et al. 2010), and

is the watershed for the agricultural region of the Plain of Les Cayes to the south (Woods et al. 1992). In 1987

the government of Haiti established the ca. 16,000 hectare Pic Macaya Biosphere Reserve, with the goal of

protecting and rehabilitating the biologically diverse forests of Pic Macaya National Park while fostering sus-

tainable development of the surrounding buffer zone (Sergile et al. 1992). For a map of Parc National Pic Ma-

caya, see Judd (1987).

Below we list species newly documented as occurring in Parc National Pic Macaya, along with previously

documented species for which we provide additional information based on recent observations. Most of these

records are based on collections made by the first three authors (L.C. Majure, G.M. Ionta, and J.D. Skean, Jr.)

during field work in the region in January, 2013. However, a few species are listed based on reconsideration of

earlier collections made by J.D. Skean or W.S. Judd. The taxa are listed in alphabetical order by family with

entries following the format of Judd et al. (1990, 1998). A complete set of voucher specimens is deposited at the

682

herbarium of the University of Florida, Gainesville (FLAS). A second set comprised of the collections made in

2013 is deposited with William Cinea at the American University of Les Cayes, and additional duplicates will

be distributed. Taxa endemic to Hispaniola (*) or the Massif de la Hotte (+) are indicated.

* Mikania polychaeta Urb.; vine, occasional; opening in forest dominated by Pinus occidentals Sw., associated

with Gleichenia sp., Miconia subcompressa Urb., M. umbellata (Mill.) Judd & Ionta, Piper sp., Renealmiajamai-

censis (Gaertn.) Horaninow, and Rhytidophyllum auriculatum Hook., 1466 m. This species is a new record for

Parc National Pic Macaya.

Begoniaceae

* Begonia cf. bolleana Urb. & Ekman; herb, uncommon; disturbed rak bwa (i.e., moist broadleaved forest over

karstic limestone) in mostly cleared forest, associated with Miconia navifolia Ionta, Judd, & Skean, ca. 900 m.

This species is a new record for the Macaya Biosphere Reserve.

* Cyperus picardae Boeck. var. brevinux Kuk.; herb, uncommon; growing out of heavy, disturbed, limestone

soils, in cleared forest, associated with Badris plumeriana Mart., Bunchosia haitiensis Urb. & Niedz., Calycogo-

nium hispidulum Cogn., Miconia curvipila (Urb. & Ekman) Ionta, Judd, & Skean, Miconia laevigata (L.) DC.,

M ecranium multiflorum (Desr.) Triana, Neea demissa Heimerl, Smilax havanensis Jacq., and Tabebuia berteri

(DC.) Britt., 728 m. Although C. picardae var. brevinux was collected outside of the Parc National Pic Macaya,

this species was previously only known from the Massif de la Selle in eastern Haiti. This collection represents

a significant range extension of the species to the west and the first collection of the species from the Massif de

la Hotte.

* Rh r hOSP ° ra d ° min8enSiS Urb ’ herb ’ uncommon ; moist, rich slopes in heavy soils of cloud forest domi-

nated by Pinus ocadentahs, Alsophyllus sp., Brunellia comocladifolia Humb. & Bonpl., and Cyathea sp., also as-

sociated with Mecranium microdyctium Urb. & Ekman, Miconia xenotricha Urb. & Ekman, and Rhynchospora

V ” ekm “ n " <Urb ' ) Mk -’ ^ 1878 ” ThiS SpeC ' eS ls a National Pic

a, S slopes of Mome Formon, 18.35160°N,

iii Urb.; small tree 2.5 m, rare, a single sterile indiv

xtremely disturbed area growing in patches of <

1; in rak bwa; 998 m. This spe-

forest with niicium hottense l

Guerrero, JuddSr A. B Morris, Meriania brevipcduncutoojudd Sr Skean, Miconia (Sagraea) polychaelc (Urb S

Ekman) Ionta, Judd, Sr Skean, M. pyramMis (Desr.) DC., and M. subcompressa This collection is

for Parc National Pic Macaya.

terfall, 18.32931°N, l^.OOigi^^Z.D. ^ S ** “

+ Calycogonium fonnonense Judd, Skean & Clase; shrub to 1.5

list of associates, 950-1200 m. Specimens initially identified and

m, rare; in rak bwa, see Judd e

reported by Judd (1978) as C.

cf. calycopteris

Majure et al.. New records from Parc National Pic Macaya 683

(Rich.) Urb. and later as C. hispidulum (Judd et al. 1998) were later recognized as representing a new species, C.

formonense, which is endemic to the Massif de la Hotte, and likely to the Macaya Biosphere Reserve (see Judd et

al. 2008). Calycogonium is polyphyletic (Michelangeli et al. 2008) and this species (along with its relatives) is

being transferred to Miconia (Judd et al., in press).

Voucher specimens: see Judd et al. (2008).

+ Henriettea hotteana (Urb. & Ekman) Alain; shrub ca. 2 m tall, rare, mixed pine/cloud forest with Arthrostyl-

idium haitiense (Pilger) Hitchc. & Chase, Calyptranthes hotteana Urb. & Ekman, Alsophila sp., Weinmannia

pinnata L., Phoradendron sp., 2219 m. This species is a new record for Parc National Pic Macaya.

74.02687°W, G.M. lonta 2031 (FLAS, NY).

+ Mecranium sp.; or small tree to 4.5 m tall in rak bwa, uncommon, 979-1188 m. This entity, previously

known from a single earlier collection, i.e., Skean 2093 (FLAS, IJ, S, US), was considered by Skean (1993) to be

a putative hybrid between Mecranium haitiense Urb. and M. revolutum Skean & Judd. Our additional collections

indicate that this entity is much more widespread in the Formon region than previously thought, and it may

1 in areas where M. haitiense and

Lon. The plants commonly have red, scurfy abrasions apparently caused by some type of

pathogen; the material is currently under study by J. Dan Skean, Jr.

74.003 17°W, 14Jan 2013, G.M. lonta 2046 (FLAS); Bwa Deron, W of Ville Formon, grow-

ing out of rak bwa, 18.32648°N, 74.020090°W, 14 Jan 2013, L.C. Majure 4312 (FLAS); Bwa Formon, S of Mome Formon, along road from Sou

Bwa to Ville Formon, karst hills, 18.31878°N, 74.00922°W, 8 Jan 2013 J.D. Skean, Jr. 5037 (FLAS, NY); near the crest along the Sou Bwa-Ville

Formon road, 18.31334°N, 74.01174°WJ.D. Skean, Jr. 5039 (FLAS, NY); S edge of Ravine Seche above region called Ravine Casco ca. 0.5 km

uphill from the waterfall, 18.33519°N, 74.01248°W, 9 Jan 2013, J.D. Skean, Jr. 5043 (FLAS, NY).

+ Meriania ekmanii Urb.; large shrub to 4.5 m tall, occasional; rak bwa in moist, broadleaf forest or cloud for-

est dominated by Pirns occidentals Sw., Gleichenia bifida (Willd.) Spreng., and G. revoluta H.B.K., associated

with Alsophila sp., Blechnum sp., Brunellia comocladifolia Bonpl., Didymopanax tremulum Krug & Urb., and

Weinmannia pinnata L. ; 1 170-1885 m. This species is a new record for Parc National Pic Macaya and was previ-

ously known only from the type specimen.

Pic Macaya and at the base of trail leading to Pic Macaya, 18.379223°N, 7

Le CM, 18.35512°N, 74.01967°W 12 Jan 2013 J.D. Skean, Jr. 5054 (FLAS); ra

Plain, 14 Nov 1989, W.S. Judd5852 (FLAS).

+ Miconia sp. This taxon was erroneously reported in Judd (1987) as “ Pachyanthus hotteana ,” an unpublished

name, based on the collections Judd 3939 and Skean 2080. It was recollected ( Ionta2023 , FLAS) on our 2013 trip

and is currently under study.

n ViUe Formon & Sou Bwa), 1 Jan 1987, J.D. Skean, Jr. 2080 (FLAS).

+ Miconia barkeri Urb. & Ekman. While preparing specimens of this species (of sect. Chaenopleura; see Judd

2007) during the January 2013 collecting trip, a distinct odor of cinnamon emanating from the leaves and in-

florescences was noted during the drying process. We subsequently detected a faint cinnamon odor on previ-

ously collected herbarium material of this species housed at FLAS. Thus we report this curious finding, which

has not been previously noted.

Majure et al., New records from Parc National Pic Macaya 685

+ Miconia cordieri Ionta & Judd; shrub to 1.5 m, uncommon; in disturbed rak bwa, 950-1200 m. This re-

cently described species (of sect. Sagraea; see Ionta et al. 2012) is endemic to the Macaya Biosphere Reserve

(Ionta & Judd 2012). It was initiaUy reported by Judd (1987) as Ossaea curvipila Urb. & Ekman (based on WJ.

Judd 3469).

m, 23 Jan 1984, W.S.Judd & D. Dod 3469 (FLAS, EHH, NY); between Ville Formon and “Experiment Station” on Deron Plain, 1170-1190 m,

14 Nov 1989, W.S.Judd 5859 (FLAS, 2 sheets).

+ Miconia curvipila (Urb. & Ekman) Ionta, Judd, & Skean; shrub to 1.5 m, common; in disturbed rak bwa,

915-1000 m. The report of this species was based upon the specimen Judd 3469, which actually represents M.

cordieri, although Skean 1320 (see specimen cited below), which actually represents this species, was initially

reported as Ossaea setulosa Urb., now Miconia rubrisetulosa Ionta, Judd & Skean (Ionta et al. 2012; Judd 1987).

and Ville Formon. 950 m, 9 Jun 1993, W.S.Judd 6892 (EHH, FLAS); Macaya Biosphere Reserve, Bwa Formon, disturbed “rak bwa” and fields

ca. 2 km SW of home of Robert and Tila Despagne, our base camp at Ville Formon, 915-945 m, 4Jan 1984 ,J.D. Skean 1320 (EHH, FLAS); Bwa

Formon, karst hills ca. 1 mi S of Ville Formon, 950-1000 m, J.D. Skean Jr. & C. McMullen 2465 (FLAS, MICH); Between Sou Bwa and Les

Platons, karst hills, 18°17'19.9"N, 73°59'28.1"W, 7 Jan 2013, G.M. I<mta20l2 (FLAS, NY); Soulet, Between Sou Bwa and Les Platons, karst

hills (disturbed rak bwa), 18.29623°N, 73.99812°W, 15Jan 2013, G.M. Ionta2050 (FLAS, NY); Macaya Biosphere Reserve, Soulet, karst hills

4- Miconia hottensis Ionta, Judd & Skean; shrub to 3 m, occasional; in disturbed rak bwa, 1100-1200 m. This

species (of sect. Sagraea), endemic to the Macaya Biosphere Reserve, was recently described by Ionta et al.

(2012). Until our recent collections, it was known from only two collections.

1100-1200 m, 13 Aug 1989 J.D. Skean, Jr. & C. McMullen 2557 (FLAS, JBSD, MICH, NY, S); Soulet, between Sou Bwa and Les Platons, karst

hills (disturbed rak bwa), 18.29623°N, 73.99812°W, 15 Jan 2013, G.M. Ionta 2052 (FLAS, NY).

+ Miconia navifolia Ionta, Judd & Skean; shrub to 2 m tall, occasional; in disturbed rak bwa, 1100-1200 m.

This species (of sect. Sagraea) is an endemic to the Macaya Biosphere Reserve and was only recently described

(Ionta et al. 2012). Until our recent collections, it was only known from two collections.

+ Miconia polychaete (Urb. & Ekman) Ionta, Judd & Skean (see Ionta et al. 2012); shrub to 1 m tall, uncom-

mon; in disturbed rak bwa, 1017-1100 m. This species (of sect. Sagraea) is a new record for the Macaya Bio-

sphere Reserve and previously was known only from the type specimen. Associated taxa include Andropogon

bicomis L., Cestrum bicolor Urb., Gleichenia sp., Lantana sp., Meriania brevipedunculata Judd & Skean, Miconia

pyramidalis, M. subcompressa, Ocotea sp., Paspalum sp., Smilax havanensis, Tabebuia berteri, and Vernonia

saepium Ekm.

Myrtaceae

+ Calyptrogenia ekmanii (Urb.) Burret; small tree to 3 m tall with purple-black fruits, uncommon, 1185 m.

This species is a new record for Parc National Pic Macaya.

+ Hottea torbeciana Urb. & Ekman; shrub, to 2 m

for Parc National Pic Macaya.

i, ca. 1200 m. This species is a new record

Poaceae

Dichanthelium aff. dichotomum (L.) Gould; herb, rare; forming small colonies in disturbed limestone soils

(from landslide) along the edge of forest dominated by Pirns occidental is, and associated with M iconic umbel-

lata, Pilea microphylla (L.) Liebm., 1466 m. Dichanthelium was not previously recorded for Parc National Pic

Macaya. This collection is not typical of D. dichotomum due to its densely pubescent leaf surfaces and sheaths,

small leaves (0.7-1.5 x 0.09-0.27 cm), lack of a basal rosette and spreading, colonial growth form.

Polygalaceae

* Badiera subrhombifolia Abbott; shrub to small tree to 7 m; in rak bwa, common, 950-1050 m. Previously

reported by Judd (1987) as Poly gala penaea L.; the populations of P. penaea - like plants occurring in the Macaya

Biosphere Reserve, the Massif de la Selle, and the Sierra de Bahoruco have been segregated and described as

Badiera subrhombifolia (Abbott & Judd 2011).

:ommon; in rak bwa

750-1560 m.Previ-

+ lUicium hottense A. Guerrero, Judd & A.B. Morris (Fig. 2); shrub to small tree to

and moist forests of Pinus occidentals, long unbumed, showing transition to cloud f<

ously reported by Judd (1987) as fflidum ekman „ A c. Smith, but ,h. populations of fllicium ekman,, iu the

Masstl de la Hotte have been segregated and described as 1. hottense (Guerrero et al. 2004) DNA sequence data

supports the sister-group mlaUonship of Mldum hottense and 1 eknuM AC. Smith, the latter a species occut-

nn g in the Massif du Nord, Haiti, and the Cordillera Central and Cordillera Septentrional of the Dominican

Republic (Guerrero etal 2004), fllicium has also been collected in the Massif de la Selle (Efemon H2230 (S), from

Morn 1 Hopttal te type of/.eWm subsp. sellemram Imkhanfiskaya); however, the specimen Ekman H2230

has no flowers (although fruits are present) and therefore cannot be identified with certainty. It is the most

rmm H22.3P 3 Hlsp aniola n fllicium - having papillate steins and young petioies and even fruiting pedicels. Elt-

munH2230 may represent an undescribed species related tttfllicium hottense, but flowering material is required

a, 18°19 , 43.2"N, 73°59'41.5

^s^ t1 Td a, * 0 h e r CC,1S (L } shrub to 2 m ta U, rare; rocky (limestone) soils along ephemeral stream

associated with, Gyrotaema myriocarpa Griseb., Lobelia robusta Graham, Senecio stenodonl Jrh Tibnurhna

longifoha (Vahl) Baill., and Thelypteris sp.; ca. 1400 m. Hunziker (1982) considered the genus Acnistus to be

15 P y ’ hOWeVer ’ Within the 8 enus Iochr °™> and they discussed

t0 SeParatC ^ tW ° §enera AUhOUgh the Ph ^ enetic of Smith

T T ! hr0ma “ ^ mono P h y^etic, they recovered two clades one of which con-

tamed the type species for the genus, I. cyanea. Acnistus arborescens is nested within rh , / T

type of the genus- however Ami^ ic th» u , ted Wlthin the clade containing the

and horticulturaliy important genus fodtromu is

S. Judd 5743 (FLAS); S slopes of Mome Fonr

Majure 4302 (FLAS, NY).

+ Ceslrum hotteanum Urb. & Ekman; shrub to 1 m tall, common, 933 m. Material originally identified as C.

hotteanum from the Pic Macaya Reserve (see Judd, 1987) is referable to Acnistus arborescens (i.e., W.S. Judd

5748). Thus C. hotteanum is reported here for the first time from Parc National Pic Macaya.

The genus Pika, the largest in the Urticaceae, comprises ca. 715 species (Monro 2004) of which approximately

235 occur in the Neotropics (Monro 2006). The Greater Antilles is one of the major centers of diversity of the

clade (Monro 2009). Pilea is especially diverse on Hispaniola, with nearly 100 species found there (Liogier

1996; Moscoso 1943). To date, 14 species of Pika have been recorded in Parc National Pic Macaya (Judd 1987),

including P. howardiana, a species described by Skean and Judd (1988) and the P. microphylla complex, which

forms a group of closely related species/entities, that is in need of further taxonomic work (see below). Addi-

tionally, previously unidentified specimens (listed as Pika sp. In Judd 1987) have been found to represent a

new species, which is described below.

Pilea aff. microphylla (L.) Liebm.— W.S. Judd 3624 (FLAS); J.D. Skean, Jr. 1530 (EHH). These collections were

reported under this name in Judd (1987). We note that they differ from the widespread and variable P. micro-

phylla in lacking small axillary branches along the major stems and in having elongate cystoliths on the adaxial

leaf surface that are arranged both longitudinally and transversely; typical material of P microphylla has cysto-

688

liths that are exclusively transversely oriented (i.e., oriented at right angles to the leaf axis). These

are extremely succulent, while typical material of R microphylla has leaves that vary from nearly hei

extremely succulent. We also note that the leaves of these specimens are not spathulate, as in the p

similar P. spathulifolia Groult of the Sierra de Bahoruco, in the Dominican Republic (Groult 1999).

Shrubs, apparently dioecious, woody, to 60 cm tall, highly branched; indumentum of simple, ± flattened, sil-

very, multicellular hairs; young stems rectangular in cross section with filiform cystoliths parallel to the stem

axis 0.1-0.3 mm long, pubescent with clear, multicellular, uniseriate, ascending hairs to 0.3 mm long; stems

becoming rounded in cross section and glabrous in age, 0.4-1.6 mm in diameter; leaves opposite, decussate,

moderately anisophyllous, i.e., the smaller leaf % to % the size of the larger leaf, to isophyllous, 0.45-1.7 x

0.25-0.9 cm, ovate to elliptic, the apex acute, the base acute to obtuse, serrate along distal % or more of the leaf

margin, mostly entire at base, each serration with a single vein, glabrous on both surfaces or with sparse, mul-

ticellular hairs to 0.5 mm long on the primary vein of the abaxial leaf surface, 3-veined, but with secondary

veins brochidodromous in distal portion of lamina, primary and secondary veins impressed adaxially and

raised abaxially, veins on the abaxial leaf surface with conspicuous filiform cystoliths produced parallel to the

vein axis, adaxial leaf surface covered in dense, conspicuous, oblong to filiform cystoliths 0.3-0.4 x 0.05-0.1

mm, produced in all directions (i.e., disorganized) and oftentimes overlapping, and becoming smaller toward

the center of the lamina, abaxial leaf lamina with punctiform to filiform, inconspicuous cystoliths, petioles

0.9-5.5 mm long, pubescent on the adaxial surface, hairs erect to ascending, glabrous on the abaxial surface

although with conspicuous punctate to elongate cystoliths; intrapetiolar stipules 1.3-2.8 x 1.2-1.7 mm, ob-

ovate to oblong, apices truncate, rounded or slightly 3-lobed, with conspicuous fil

parallel to the axis on the abaxial surface; carpellate inflorescences red, producing ±

along the main axis, 0.6-1.2 cm long, 0.15-0.8 cm across, bracts 0.3-0.6 x 0.05 mm, ovate to obovate with

apices acute, rounded or three-lobed as in the stipules; carpellate flowers with pedicels 0.3-0.4 mm long, tepals

3, dimorphic, the larger tepal saccate, 0.4-0.7 mm long, elliptic, forming a cup-shaped, fleshy structure enclos-

ing gynoecium, its margins incurved, clasping, and membranous, apex 3-lobed, the central lobe succulent and

exceeding the two membranous lateral lobes, the two lateral tepals 0.2-0.6 mm long, ovate to narrowly elliptic,

with acute apices; ovary appearing unicarpellate, stigma penicellate, with elongated papillae; staminate flow-

ers not seen; achenes elliptical, biconvex, brown, the surface moderately alveolate, 0.6-0.65 x ca. 0.4 mm.

Etymology. The specific epithet vermicularis refers to the wormlike appearance resulting from the disor-

ganized and oftentimes overlapping cystoliths on the adaxial leaf surface (Fig. 3D).

laris is morphologically similar to P. wullschlaegelii Urb. (Jamaica)

maica); all three species have “disorganized” cystoliths associated

wnn me aaaxiai teai sunace, glabrous leaf surfaces, and serrate leaves. Pilea vermicularis differs from these

sf*cies by to pubescent stems (Fig. 3B). Pilea vermicularis also differs from P. wullschlaegelii by the lack of

glands (hydathodes) on the lower leaf surface and the acute to obtuse vs. obtuse to rounded leaf bases (Fig. 3C,

E, F). Pilea vermicularis differs from P. radicans by its frutescent habit (vs. vine-like habit in the latter). Pilea

vmnruUnc .c oic« - i— — D . rufescens Fawc. & Rendle (Jamaica), although that species has densely pubes-

n the stem, instead of glabrous leaves and silvery stem hairs. Morphologically, P

to be closely related to Clade 2, Unit 5 of Monro (2006), sharing heteromorphic leaves

i P radicans (Sw.) Wedd. I

cent leaves and reddish hair

Majure et al. r New records from Parc National Pic Macaya 689

with incised margins and 3-merous carpellate flowers (Fig. 3C, E, G). However, those taxa are known from

Central America instead of the West Indies and are glabrous instead of pubescent. The West Indian subclade

(Clade 2, Unit 4) contains pubescent species with isomorphic leaves with entire margins, and 3-merous car-

pellate flowers (Monro 2006; see also “Clade H” in Jestrow et al. 2012). Piled vermicularis may belong to this

group if leaf margins are not consistent throughout the clade. Members of this clade occur in the Parc National

Pic Macaya (e.g., P. domingensis Urb.; Judd 1987). The phylogenetic analyses of Monro (2006) and Jestrow et al.

t this point is

(2012) are preliminary and include only a sm

phylogenetic relationships of P. vermicularis at

wn Pilea species, thus a

ACKNOWLEDGMENTS

We thank William Cinea for his invaluable assistance to the first three authors during their field work in the

Formon region. We also thank Florence E. Sergile for her help while these authors were in Port-au-Prince,

Haiti. The staff of the University of Florida Herbarium (FLAS) is gratefully acknowledged, especially Norris

Williams and Kent Perkins, for their help in processing herbarium material. Tom Zanoni and Jackeline Salazar

provided helpful comments on an earlier version of the manuscript. This research was supported, in part, by

the National Science Foundation Grant BSR-0818399. The trip (1984) on which Pilea vermicularis was collect-

ed was supported by Charles A. Woods, through his USAID Project No. 521-0191-A-00-7107-00.

tt, J.R. and W.S. Judd. 201 1 . Badiera subrhombifolia, a new species from Hispaniola. Brittonia 63:1

s Republique Dominicaine.

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Huber, B.A., N. Fischer, and JJ. Astrin. 201 0. High level of endemism in Haiti's last remaining forests: a revision of Modisimus

(Aranae: Pholcidae) on Hispaniola, using morphology and molecules. Zool. J. Linn. Soc 1 58:244-299.

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Ionta, G.M. and W.S. Judd. 2012. Miconia cordieri, a new species of Miconia sect. Sagraea (Melastomataceae) from the

J. Bot. Res. Inst. Texas 6:37-44.

, . 10 Clt McMullen- 201 2.Two new species of Miconia sect. Sagraea (Melastomataceae)

from the Macaya Biosphere Reserve, Haiti, and twelve relevant new species combinations. Brittonia 64-61 -72

Jestrow, B„ JJ. Valdes, FJ. Rodriguez, and J. Francsco-Ortega. 2012. Phylogenetic placement of the Dominican Republic

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Monogr. 81:1 -235.

Judd, W.S., J.D. Skean, Jr., and C.K. McMullen. 1 990. The flora

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Judd, W.S., J.D. Skean, Jr., and Dana G. Griffin, III. 1 998 . 1

and nomenclature! changes, II. Moscosoa 10:1 14-120.

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Judd, W.S., E.R. Becquer,ai

:i. Brittonia 60:265-270.

Michelangeu, F.A., W.S. Judo, D.S. Penneys, J.D. Skean, Jr., E.R. BEcquer, R. Goldenberg, and C.V

I in the Caribbean. Bot. Rev. 74:53-77.

Sergile, F.E., C.A. Woods, and P.E. Paryski. 1 992. Final report of the Macaya Biosphere Reserve Project. Prepared for USAID/

Haiti under contract #521 -01 91 -A-00-71 07. Gainesville, Florida, U.S.A.

Skean, J.D., Jr. 1993. Monograph of Mecranium (Melastomataceae-Miconieae). Syst. Bot. Monogr. 39:1-1 16.

Skean, J.D., Jr. and W.S. Judd. 1 988. A new Pilea (Urticaceae) from Hispaniola. Rhodora 90:21 5-222.

Smith, S.D. and D.A. Baum. 2006. Phylogenetics of the florally diverse Andean clade lochrominae (Solanaceae). Amer. J.

Bot. 93:1 140-1 153.

Woods, C.A. and JA Ottenwalder. 1 992. The natural history of southern Haiti. Prepared for USAID/Haiti under USAID con-

tract #521-01 91 -A-00-7107. Gainesville, Florida (available in FLAS library).

Woods, C.A., F.F. Sergile, and J.A. Ottenwalder. 1992. Stewardship plan for the national parks and natural areas of Haiti.

Prepared under USAID contract #521-01 91 -A-00-71 07. Gainesville, Florida (available in FLAS library), U.S.A.

BOOK REVIEW

George Yatskievych. 2013. Steyermark’s Flora of Missouri, Volume 3: Dicots, Fabaceae (subfamily Faboi-

deae) through Zygophyllaceae. (ISBN-13: 978-0-915279-13-5, hbk). Missouri Botanical Garden Press,

P.O. Box 299, Saint Louis, Missouri 63166-0299, U.S.A., in cooperation with the Missouri Department of

Conservation, P.O. Box 180, Jefferson City, Missouri 65201-0180, U.S.A. (Orders: www.mbgpress.info,

orders@mbgpress.org, 1-877-271-1930). $65.00, xvii + 1382 pp., 194 plates of b&w line drawings, 20

figures (incl. 27 b&w photographs), 798 distribution maps, 7" x 10" x 2.5".

mbers of the genus Acer that has bee

Volume 3. Anyone attempting to ki

st then use the key to the genus in \

e 3. That,

tit and di-

from the Aceraceae covered in Voluim

unknown Acer spp. after reaching thai

Obviously, the taxonomy of plants is in a constant state of flux so the unfortunate set of circumsu

genus Acer is no fault of the author, but it would have been helpful to repeat the key to the genus

however, was likely unachievable due to scheduled deadlines.

It would have been useful to include a short discussion in the introduction on some of the r

vergent changes in taxonomy such as Desmodium to Hylodesmum, some Lespedeza to Kummerowia, Psoralea to Or-

bexilum or Pediomelum, Coronilla to Securigera, Bumelia to Sideroxylon , Saxifraga to Micranthe s, Dodecatheon to Prim-

ula, Hybanthus to Cubelium, etc. The author does provide a short summary of some of the major familial changes in

the preface to the book, but it is difficult to comprehend the magnitude of such changes without the help of a table

that would list the old and new names for families, genera, and in some cases, species. There is considerable discus-

sion on the new families that have emerged from Scrophulariaceae and genera that are now merged into this family

on page 1106, but as noted above, the changes would have been best represented in a comparison table.

Some amateur botanists and naturalists are likely to have difficulty with many of the technical terms used in

family, genera, and species accounts, especially those associated with molecular genetics, but the author provides a

glossary in the back of the book with definitions for the overwhelming majority of sophisticated botanical language.

Due to scheduling deadlines, it is unfortunate that some species are not illustrated (e.g. Aeschynomene rudis, Cen-

trosema virginianum, Cotoneaster acutifolius, Dalea gattingeri, D. villosa, Lablab purpureus, Lathyru s tuberosus, Rhodo-

typos scandens, multiple species ofRubus spp., Spireajaponica, etc.). As with any botanical compilation, however, it is

impossible to keep up with new additions of taxa to state floras, and the author mentions in the preface that an aver-

age of nine species are added to the Missouri flora each year. Several species of Crataegus have been reduced to vari-

etal rank, but the lack of county distribution maps for the different variants prevents a visual evaluation of areas of

the state where such varieties may be found or a cursory examination of the conservation status of rare taxa based on

will complain about the $65 price tag, but the book is a bargain when compared to the information provided.

Overall, the negative comments should be considered nonconsequential compared to the overall quality of the

book. As with Volumes 1 and 2, Volume 3 of the Flora of Missouri should be on the book shelf of every botanist, natu-

ralist, and plant enthusiast in the Midwest. No longer should the entire treatment of Flora of Missouri Volumes 1 2

and 3 by George Yatskievych be compared to his mentor’s original work. With the completion of the revision it is

Yatskievych who has set the standard for similar state floras to follow!-Paul M. McKenzie, Ph.D., U.S. Fish and Wild-

life Service* 101 Park DeVille Drive; Suite A, Columbia, Missouri 65203, U.SA.

> of the

CHUSQUEA CLARKIAE (POACEAE: BAMBUSOIDEAE:

BAMBUSEAE: CHUSQUEINAE): A NEW SPECIES IN

SECTION LONGIPROPHYLLAE FROM NARINO, COLOMBIA

Emmet J.Judziewicz

Ximena Londono

Robert W. Freckmann Herbarium

Department of Biology and Museum of Natural Hist

University of Wisconsin-Stevens Point

Stevens Point, Wisconsin 54481, U.S.A.

emmet.judziewicz@uwsp.edu

Sociedad Colombiana del Bambu

Apartado Aereo 1 1574

Cali, COLOMBIA

ental of Narino, Colombia, is described and illu:

m (versus 1750-2200(-2700) m).

RESUMEN

la Cordillera Occidental de 1

A bamboo collected in Colombia along the Pasto to Tumaco road in 1990 by the second author, John Kress, and

Wilson Devia is, upon examination, a new species of Chusquea Kunth (Poaceae: Bambusoideae: Bambuseae:

Chusqueinae). Its combination of a scandent habit, infravaginal branching, tightly circular array of subsidiary

buds, narrow, erect, persistent, abaxially scabrous culm leaf blades, spikelets with well-developed glumes I and II,

and apiculate spikelet bracts all place it as a member of Chusquea Section Longiprophyllae L.G. Clark, a group that

now comprises seven species occurring at elevations of (950-)1130-2750min the Andes from Venezuela to central

Peru. The Section is most diverse in Colombia (Clark 1990).

clarkiae Londono & Judz., sp. n

Rhizomes unknown. Culms up to 10 m long, 1-1.5 cm in diameter, scandent, solid. Culm leaves at least 26 cm

long (base not seen), sheaths at least 20 cm long, abaxially strongly retrorsely scabrous, the midrib obscure;

blades 6. 5-8.5 cm long (in two examples available), folded width at base 3-5 mm, linear-attenuate, erect, per-

sistent, abaxially slightly retrorsely scabrous, the midrib inconspicuous, the margins glabrous; inner ligules

0.5-1 mm long. Nodes at mid-culm with a central bud subtended by 4 slightly smaller subsidiary buds; supra-

nodal ridge visible as a slightly raised line, not prominent; girdles 1.5-2.5 mm wide, brown to dark brown,

covered with fine, brown, retrorsely appressed, silky hairs 0.5-1 mm long. Branching infravaginal, with

emergent branches in available material diverging downwards up to 30° from the main culm; prophylls 2 — 4

cm long, narrow, glabrous; leafy subsidiary branches 4-5 per node, 4-8 cm long, ca. 1 mm in diameter, not

rebranching. Foliage leaves 4-7 per complement; sheaths glabrous, uniform in color, keeled toward the sum-

mit, the margins glabrous, summit extension absent to 2 mm long; blades 13-20 cm long, 1.5-2 cm wide, L:W

- 8-10, narrowly lanceolate, glabrous except puberulent on the adaxial surface of the pseudopetiole not tessel-

late, the midrib usually visible abaxially and prominent for ca. % the length of the blade, the base rounded-at-

tenuate, the apex acuminate-attenuate, the margins antrorsely scabrous to serrulate; pseudopetioles 3-5 mm

long; outer ligules 1-2 mm long, indurate, usually irregularly bilobed, each lobe in turn lacerate, glabrous; in-

6 %

Journal of the Botanical Research Institute of Texas 7(2)

Distribution —Known only from a recorded elevation of 1130 m along the Pasto-Tumaco highway in the Cor-

dillera Occidental in the Department of Narino, Colombia. Google Earth (accessed 18 June 2013) gives a likely

location of 1°19’N, 78°07'W and an elevation of ca. 1100 m.

Etymology .— We name this species in honor of our friend and colleague Dr. Lynn G. Clark, Director of the

Ada Hayden Herbarium at Iowa State University, and agrostologist specializing in bamboos, who has made an

immense contribution to our understanding of the genus Chusquea and the Chusqueinae (Clark, 1989, 1990,

1992, 1993, 1996, 1997, 2001).

We thank University of Wisconsin-Stevens Point undergraduate Rebecca A. Gregory for the illustrations, and

Robert J. Soreng and Christopher D. Tyrrell for helpful reviews.

Clark, LG.

Longifoliae (Poaceae-Bambusoideae). Syst. Bot. Monogr. 27:1-127.

Clark, L.G. 1990. Chusquea Section Longiprophyllae (Poaceae: Bambusoideaf

Syst. Bot. 15:617-634.

Verticillatae, Section Serpentes, and Section

Clark, L.G. 1992. Chusquea Sect. Swallenochloa (Poaceae: Bambusoideae) and allies in Brazil. Brittonia 44(4):387-422.

Clark, L.G. 1 993. Five new species of Chusquea (Poaceae: Bambusoideae) and a new combination. Novon 3(3):228-238.

Clark, LG. 1 996. Four new species of Chusquea (Poaceae: Bambuseae) from Brazil and Ecuador. Brittonia 48(2):250-262.

Clark, L.G. 1997. Diversity, biogeography and evolution of Chusquea (Poaceae: Bambusoideae). In: G.P. Chapman ed

The Bamboos. Academic Press, London, U.K. Pp. 33-44.

Clark, L.G. 2001. Diversification and endemism in Andean woody bamboos (Poaceae: Bambusoideae). Bamboo Sci.

THE SPIKELET CALLUS OF ERIOCHLOA VILLOSA (POACEAE)

Stephen J. Darbyshire

Marie-Josee Simard

Agriculture and Agri-Food Canada

960 Carling Ave., Saunders Building #49

Ottawa, Ontario, K1A 0C6, CANADA

Stephen.Darbyshire@AGR.GCCA

Agriculture and Agri-Food Canada

2560 Hochelaga

Quebec, Quebec, GIV2J3, CANADA

Agriculture and Agri-Food Canada

2585 County Rd. 20, RR#1

Harrow, Ontario, NOR 1G0, CANADA

ABSTRACT

The panicoid grass genus Eriochloa contains about 30-35, primarily tropical and subtropical, species which are

characterized by an unusual swelling and cup-like callus structure at the base of the spikelets (Clayton & Ren-

voise 1986). This unusual structure has been interpreted as a swelling of the lowest rachilla intemode sur-

mounted by the lower (proximal) glume which is reduced to a short flange of tissue (Hsu 1965; Shaw & Smeins

1979, 1983). However, in their anatomical examination of the callus of three Eriochloa species, Thompson et al.

(1990) found that the stele in the spikelet rachis remained unbranched until the upper glume traces, and inter-

preted the lack of stelar nodes within the callus region as an indication that no part of the callus is likely to

represent a vestigial lower glume. Although the callus structure has been investigated in a number of Eriochloa

species, it has not been studied in E. villosa (Thunb.) Kunth. This species is native to eastern Asia from the Rus-

sian Far East to northern Viet Nam, but has been introduced to other areas including North America where it

has been spreading as an agricultural weed (Darbyshire et al, 2003).

Shaw and Smeins (1979, 1983) examined epidermal characteristics of the spikelet callus in 20 species of

Eriochloa. They observed that the structure was divided into two parts, both with characteristics of the epider-

mis consistent with the interpretation that the callus is formed from an expansion of the lowest rachilla inter-

node and surmounted by the remnants of the first glume. Three types of calluses were described (Table 1). The

Type 1 callus was by far the most common in the genus, being seen in 16 of the species examined (E. aristata

Vasey, E. boxiana Hitchc., E. contracta Hitchc., E. eggersii Hitchc., E. ekmanii Hitchc., E. acuminata (J- Presl)

Kunth (= E. lemmonii var. gracilis (E. Foum.) Gould), E. lemmonii Vasey & Scribn., E. meyeriana (Nees) Pilger,

E. michauxii (Poir.) Hitchc., E. montevidensis Griseb., E. pacifica Mez, E. peruviana Mez, E. punctata (L.) Ham.,

E. sericea (Scheele) Vasey, E. setosa (A. Rich.) Hitchc., and E. weberbaueri Mez). Distinctive features of the Type

1 callus included a basal portion with a smooth epidermis beset with silica bodies and an apical portion of

plicate epidermal tissue. The Type 2 callus, seen in 3 species (E. distachya Kunth, E. grandiflora (Trin.) Benth.

bead macrohairs

Callus Type 1

Callus Type2

Callus Type J

and E. nelsonii Scribn. & J.G. Sm.), was similar to Type 1 but larger in size and lacking silica bodies. The Type 3

callus, seen in only one New World species (E. polystachya Kunth), had the basal portion without a smooth and

indurate epidermis and the apical portion was a membranous “reduced glume” encircling the base of the spike-

let. The Old World species E. rovumensis (Pilg.) Clayton (= E. biglumis Clayton) is somewhat anomalous in the

genus, possessing a swollen spikelet base and well-developed lower glume, although Shaw and Smeins (1983)

state that the lower glume is similar to that seen in E. polystachya.

The purpose of this study was to examine the characteristics of the spikelet callus in the Asian E. villosa

and compare it with the previously published observations on other species which are primarily from North

and South America and Africa.

MATERIALS AND METHODS

Spikelets originating from populations occurring in southern Quebec (Darbyshire et al. 2003) were used.

Thirty randomly selected spikelets from one population were measured under lOx magnification for spikelet

and callus size and the measurements were compared with those given by Shaw and Smeins (1979). Selected

spikelets were air-dried, coated with gold and examined with a Philips XL30 ESEM microscope at 5kV accel-

eration voltage. Spikelets stored in 70% ethanol were re-hydrated, hand sectioned and stained with methylene

blue or toluidine blue O. Air-dried spikelets were re-hydrated, hand sectioned and stained with Sudan IV.

Spikelets are broadly elliptical and dorsiventrally compressed (Fig. 1 A). The size is reported as 3.9-5.5(-6.5)

mm long, 2.0-2.8(-3.0) mm wide (Darbyshire et al. 2003; Shaw et al. 2003). Spikelets from the Quebec popula-

tion measured 5.3 (SD = 0.18) mm long and 2.7 (SD = 0.09) mm wide, with the callus 0.8 (SD = 0.09) mm long

and 1.5 (SD = 0.12) mm wide. The observed E. villosa callus size was larger than in any of the 19 species exam-

ined by Shaw and Smeins (1979). The callus length and width averaged 15% and 55% of the spikelet length and

width, respectively. Shaw and Smeins (1979) reported variation in callus sizes between species they measured,

with Type 2 calluses tending to be larger than Type 1, but no correlation between callus size (length/width)

and type was detected in this study (data not shown). Epidermal characteristics of the callus of E. villosa are

most similar to those described as Type 1 by Shaw and Smeins (1979).

Apart from the central bundles, no branching of vascular tissue was observed anywhere in the callus

structure (cf. Thompson et al. 1990). The callus was divided into two mart or less equal and strongly demar-

cated parts (Fig. 1). The basal part (bead) is formed of hardened tissues and the apical portion (cup) above is

°‘ ; the caltas usuall >' ™ f°nn a complete ring thereby incompletely clasping the spikelet

(Fig. 1 B), although sometimes the cup was almost completely encircling with only a small notch at the junction

699

fa- 1. Spikelets of Eriochloa villosa. A, left adaxial view, right abaxial view <

callus (SEM).

of the lateral edges (Fig. 1 A, left). The callus cup was directly opposite the upper glume or only slightly off

centre (i.e., not quite bilaterally symmetrical). An additional flap of tissue arising from the adaxial surface of

the callus cup is present in some species of Eriochloa and was referred to as an “extension” by Shaw and Smeins

(1979) and as a “ventral projection” by Thompson et al. (1990). The ventral projection in the callus of E. villosa

was observed to arise from the adaxial side of the cup at its base, where it formed a second inner, but incomplete

tion extended beyond the fleshy portion of the cu]

)t visible without dissection (Fig. 2 B, 4). The posi

membranous cup (Fig. 2 A, B). Sometimes the ve

(Fig. 1 A (right), 2 A), but usually this flange of ti

tion of the ventral projection is consistent with the position of the first glume, although there is no other evi

dence of homology.

No stomata or prickle hairs were observed on either part of the E. villosa callus. The bead epidermis wa<

hardened and smooth or minutely roughened (Fig. 3 A, B). Bicellular microhaiis were common apically (Fig. 2

W, but macrohairs were rarely seen. Rounded to broadly eliptic or shallowly lobed silica cells were commor

basally (Fig. 3 A). The abaxial epidermis of the cup was heavily plicate (Fig. 3 C) Silica bodies were not ob-

served on the cup. Bicellular microhairs were common throughout the cup and macrohairs were occasional!.

Darbyshire et al., Spikelet callus of Eriochloa villosa

seen (Fig. 3 C). Long cells with interlocking cell walls were seen in the epidermis of the cup and the ventral

projection.

Unlike the callus descriptions of most other Eriochloa species, occasional macrohairs were detected on

both the lower and apical portions of the E. villosa callus (Table 1; Fig. 3 C), but see also Shaw and Smeins

(1983). This is not unexpected as macrohairs are a common epidermal feature on other structures of E. villosa

(cf., Thompson et al. 1990) and their occasional detection simply a function of the large numbers of E. villosa

The callus of untreated spikelets has an oily or waxy appearance (Fig 1A). Parenchyma cells of the cup

portion and the ventral projection of the callus contained large vacuoles whose contents stained red with Su-

dan IV, indicating that this tissue is rich in lipids (Fig. 4). A lesser amount of staining occurred in the upper

portion of the harder bead tissues of the callus. The large amount of lipids present in the fleshy callus cup sug-

gests that this tissue may act as an elaiosome (myrmecochory) or animal attractant (Davidse 1987). No evi-

dence was observed of oil secretion or accumulation in concave cavity formed by the cup (or ventral projec-

tion), as was suggested by Arriaga (2000). It is uncertain what types of vectors might serve as effective dispersal

agents, but various types of insect and vertebrate (including birds and rodents) seed predators might be at-

tracted to the lipid food source. Optimal seedling emergence was observed from soil depths up to about 5 cm,

but occurred from depths > 9 cm (Liu & Owen 2003). This would suggest that diaspores can tolerate burial by

seed caching animals.

The specific gravity of plant oils typically range from 0.91 to 0.97 at 15°C (Lide 1990). Large quantities of

oils may affect buoyancy, movement and orientation (when unevenly distributed) of E. villosa diaspores under

aqueous conditions. While the callus cup is unlikely to have a major impact on hydrochory, the increased

buoyancy at the basal portion of the spikelet may provide some secondary functionality in diaspore transport

and placement when free water is present.

Eriochloa the lower glume i:

nsistent with those of Shaw and Smeins (1979, 1983) and

species in the genus, but provide no further evidence of the ontogeny of the

most similar to the Type 1 callus of Shaw and Smeins (1979), the most com-

e observed in the distribution of micro- and macrohairs. In most species of

tally described as absent or greatly reduced and fused with the callus bead,

ribed to the genus are said to have well-developed lower glumes and resemble

species of Brachiaria (Clayton 1975; Gibbs Russell 1981; Shaw & Smeins 1983). Detailed anatomical study of

these species (primarily African) is necessary for a b

the generic relationships of Eriochloa, Urochloa and Brachiaria.

ACKNOWLEDGMENTS

Clayton, W.D. 1 975. New species of Eriochloa from Africa. Kew Bull. 30:1 07-1 09.

Clayton, W.D. and SA Renvoize. 1986. Genera Graminum, grasses of the world. Kew Bull. Addit. Ser. 1 3:1 -389.

Darbyshire, SJ., C.E. Wilson, and K. Alison. 2003. The biology of invasive alien plants in Canada. 1 . Eriochloa villosa (Thunb.)

Kunth. Can. J. Plant Sci. 83:987-999.

Davidse, G. 1987. Fruit dispersal in the Poaceae. ln:T.R. Soderstrom, K.W. Hilu, C.S. Campbell and M.E. Barkworth, eds.

Grass systematics and evolution. Smithsonian Institution Press, Washington, D.C. Pp. 1 43-1 55.

Gibbs Russell, G.E. 1 981 . A new combination in Eriochloa. Bothalia 1 3:457.

Hsu, C-C. 1965. The classification of Panicum (Gramineae) and it

cule, style-base and lemma. J. Fac. Sci. Univ. Tokyo Sect. Ill Bot. 9:43-150.

Ude, D.R., ed. 1 990. CRC handbook of chemistry and physics. CRC Press, Boca Raton, L<

Liu, M-C. and M.D.K. Owen. 2003. Effect of seed ri

Weed Sci. 51:78-82.

Shaw, R.B. and F.E. Smeins. 1 979. Epidermal characteristics of the callus in Eriochloa (Poaceae). Amer. J. Bot. 66:907-91 3.

Shaw, R.B. and F.E. Smeins. 1983. Additional observations of the callus in Eriochloa (Poaceae). Iselya 2:15-19.

Shaw, R.B., R.D. Webster, and C.M. Bern. 2003. Eriochloa Kunth. In: Flora of North America Editorial Committee, eds. Flora of

North America north of Mexico. Oxford University Press, New York and Oxford. 25:507-51 6.

Thompson, RA, RJ. Tyrl, and J.R. Estes. 1 990. Comparative anatomy of the spikelet callus of Eriochloa, Brachiaria, and Uro-

chloa (Poaceae: Paniceae: Setariineae). Amer. J. Bot 77:1463-1 468.

CHANGES TO POTENTILLA RUBRICAULIS S.L., R HOOKERIANA (ROSACEAE),

AND ERSTWHILE SYNONYMS IN

FLORA OF NORTH AMERICA NORTH OF MEXICO

Barbara Ertter Reidar Elven

University of California

Berkeley, California 94720-2465, USA.

Snake River Plains Herbarium

Boise State University

Boise, Idaho 83725-1 51 5, U.S.A.

University of Oslo

P.O. Box 1172

Blindern, NO-03 18 Oslo, NORWAY

David F. Murray

University of Alaska Museum of the North

907 Yukon Drive

Fairbanks, Alaska 99775-6960, USA

Keywords: Potentilla rubricaulis, Potentate hookeriana, Potemilk nomenclature, Flora of North America North of Mexico

NOMENCLATURAL BACKGROUND OF THE POTENTILLA RUBRICAULIS COMPLEX

Preparation of the first continent-wide treatment of Potentilla (Rosaceae) for North America since that of Ryd-

berg (1908) required a full-scale revisionary effort to synthesize differing taxonomic concepts among regional

justify the resultant changes was well beyond the scope of a synoptic flora. This was particularly true for the P.

rubricaulis complex, which is accordingly published here instead.

L R«- Inst Teas 7(2): 703-711.2013

rubricaulis Lehm

r-papil-

> in the last century (e.g., Polunin 1940; Porsild 1957; Hulten 1968; Po:

:ber & Wittman 1996; Holmgren 1997) have generally used the na

licting manner, for plants with at least some palmate t<

(vs. strictly temate) leaves and short styles (less than 1.5 mm) that are usually thickened and glai

late basally. Alternatives have included the segregation of some elements as the ambiguous P. quinquefolia

Rydb. (e.g., Hitchcock & Cronquist 1961; Boivin 1967; Dorn 1977) or the inclusion of P. rubricaulis inP pulchel-

la R. Br. (Scoggan 1978). As further confusion, Sojak (1986, 1994) added P. hookeriana Lehm., a name previous-

ly used for a widespread species in the P. nivea L. complex, to the synonymy of P. rubricaulis s.l.

Although some of our early annotations and treatments (e.g., Elven & Aiken 2007) also reflect an inclu-

sive Potentilla rubricaulis, we now conclude that several reasonably distinct species can be parsed, based on our

combined studies of a broad range of material from the Arctic to the southern Rocky Mountains, increased at-

tention to vestiture and inflorescence architecture, and new analyses of types, including some lectotypifica-

tions (Ertter 2008; Sojak 1986). Most of these species collectively comprise Potentilla sect. Rubricaules (Rydb.)

originated as intersectional hybrids between members of temate-leaved sect. Niveae (Rydb.) A. Nelson and

pinnate- to subpalmate-leaved sect. Pensylvanicae Poeverl. Sojak (1986) considered at least 30 species, mostly

Eurasian, to be such intersectional hybrids. Jurtzev (1984) and Sojak (1986) have treated the Eurasian varia-

tion in this group in detail; Sojak’s work also dealt with Greenland and North American species. He initially

erican species in the complex, but later (1994) reverted to an inclu-

sive P rubricaulis for non-arctic members of the section.

Listed below are the species comprising sect. Rubricaules in the pending volume 9 of Flora of North Amer-

ica North of Mexico (FNANM), plus additional taxa that have been included in Potentilla rubricaulis s.l , with

full discussion and type paragraphs. Descriptions, keys, and distributions can be found in the pending FN-

ANM treatment. Interpretations of probable parentage, mostly by Sojak, are provided only for arctic and sub-

arctic species. Problematic or minor elements not given full treatment in FNANM are also addressed, as are

other erstwhile synonyms of P rubricaulis s.l. Additional information and discussion on arctic species is avail-

able at the Panarctic Flora website (http://nhm2.uio.no/paf/).

SUBARCTIC AND TEMPERATE SPECIES IN FNANM

Outside of the Arctic, seven species previously included in Potentilla rubricaulis s.l. are being treated as distinct

species in FNANM. These are only the best-defined elements morphologically and geographically, occurring

primarily in the Rocky Mountains and subarctic regions of Alaska and western Canada. Further investigation

is needed to resolve numerous poorly understood variants and transition zones, especially in western Canada.

For example, plants from the northern prairies of Saskatchewan, Alberta, and Montana evidently represent an

undescribed taxon, and collections from some other areas (e.g„ Hoback Canyon, Wyoming; Schell Peak, Ne-

vada) are also under investigation as possible novelties.

narrow circumscription of Potentilla rubricaulis is restricted to reli

ces occurring mainly in glaciated parts of subarctic northwesten

J6) initially interpreted P rubricaulis (including P furcata A.E. Poi

iP arenosa (Turcz.) Juz. (sect. Niveae) x P bimundorum Sojak (sect, f

rely large plants with open inflores-

lanada and southern Alaska. Sojak

i) as the hybrid species originating

nicae), but later (1994) c

t. Pensylvanicae parent for a broadly defined P. rubricaulis. The

distincuon between P rubricaulis and large forms of P arenosa with supernumerary leaflets is problematic,

though the latter lends to have more stiffly spreading petiole hairs and prominently peliolulate central leaflets.

705

ihared with P. bimundorum

;ins, and raised, partly reticulate vein:

rithin the glaciated area of overlap bet'

he unglaciated Beringian parts of Alas

md petiole hairs, sparse glands, strongly revolute leaf mar-

epicalyx bractlets and sepals. The range of P. rubricaulis is largely

1 P. arenosa and P. bimundorum, but the species is nearly absent from

“Vanamo" 2:25. 1947 [1949]. 1

The name Potentilla hookeriana, or its infraspecific equivalent under P. nivea, has traditionally been applied to

a widespread primarily ternate-leaved member of sect. Niveae. As noted by Sojak (1986), however, the type has

at least some 5-foliate leaves, as do most other populations outside of the Arctic. As a result, the ternate-leaved

arctic and subarctic material that was previously known as P. hookeriana has now taken P. arenosa (Turcz.) Juz.

as the next available name. This occurred after a brief period when it was called P. nivea s.s. (e.g., Sojak 1989;

Cody 1996) until that name was conserved with a conserved type (Eriksen et. al. 1999), thereby maintaining

P. nivea in its traditional sense for a separate species.

from the Rocky Mountains and eastern Great Basin. In addition to encompassing most of traditional P. hooker-

iana from this area, our new circumscription also includes many 5-foliate collections previously identified as P

Potentilla furcata A.E. Porsild, Bull. Natl. Mus. Canada 121:224. 1951. Potentilla hookeriana Lehm. var. furcata (A.E.

Although included within Potentilla rubricaulis by Sojak (1986, 1994), in our interpretation P. furcata differs in

several characters that suggest a hybrid origin from P. arenosa and a glandular member of the P. pensylvanica L.

complex. It is a characteristic species of the steppe bluffs of interior and south-central Alaska, Yukon Territory,

and northern British Columbia, mainly within the unglaciated Beringian region (i.e., largely allopatric to P.

Potentilla modesta Rydb., N. Amer. FI. 22:331. 1908. Potentilla concinna Richardson var. modesta (Rydb.) S.L. Welsh & B.C.

Potentilla modesta is the dominant component of P. rubricaulis s.l. in the Intermountain Region (e.g., Holmgren

1997, including illustration). Plants generally have more consistently 5-foliate leaves and more congested inflo-

rescences than P. hookeriana. Sojak (1994) and Holmgren (1997) considered P modesta to be a synonym of P.

rubricaulis s.l., while Hulten (1945) included P modesta in the synonymy of his primarily arctic concept of P.

nivea subsp. subquinata (Lange) Hulten. The epithet modesta is misapplied in Welsh et al. (1993), where the

combination P. concinna Richardson var. modesta (Rydb.) S.L. Welsh & B.C. Johnst. is used for long-styled

plants placed by us in P. concinna var. divisa Rydb. (sect. Concinnae (Rydb.) A. Nelson).

Potentilla pseudosericea Rydb., Mem. Dept. Bot. Columbia Coll. 2:98. 1898. 1

Texas 2:204. 2008: GH; isolectotypes: GH.JEPS, UC).

As discussed elsewhere (Ertter 2008), the traditional use of Potentilla pseudosericea for plants endemic to the

White Mountains on the border of California and Nevada was at odds with Rydberg’s (1908) citation of “Rocky

Mountains” as type locality. This citation resulted from the fact that two of the three syntypes were from the

Rocky Mountains, both now identified as P. bipinnatifida Douglas ex Hook, in sect. Pensylvanicae. The tradi-

tional application has been preserved by lectotypification on the third syntype, purportedly from Nevada but

actually from California (Ertter 2008). Whether the species occurs in Nevada remains to be determined. Sojak

(1994) included P. pseudosericea within P. rubricaulis si

mer. FL 22:3

U.S.A. Utah: La Sal Mts., Purpus 251 p.p. (holotwe: US; isc

Potentilla paucijuga, treated by us as endemic to the La Sal Mountains of Utah, was included in the synonymy

of P. rubricaulis si by Sojak (1994) and Holmgren (1997). Plants have subpalmate leaves, with somewhat larger

flowers and longer styles than sympatric members of sect. Rubricaules. In Colorado (e.g., Weber & Wittman

1996), the combination P pensylvanica var. paucijuga (Rydb.) S.L. Welsh & B.C. Johnston has been misapplied

to what is treated by us as P jepsonii Ertter (sect. Pensylvanicae).

(Rydb.) Th. Wolf,

Although often included in Potentilla rubricaulis s.1. (e.g., Holmgren 1997; Weber & Wittm

tana differs in having subpinnate leaves and smooth columnar styles. It is accordingly placed by us in sect.

Subjugae (Rydb.) A. Nelson rather than sect. Rubricaules, with many collections being transitional to P. subjuga

Rydb. The species is restricted to high elevations in the mountains of Colorado, Montana, Utah, and Wyoming.

Lectotypification of P. saximontana , provided above, is required only because a specific institution was not in-

dicated in the protologue. Otherwise the GH specimen could be accepted as holotype, as the only known du-

plicate annotated with this name by Rydberg.

The lectotype of Potentilla nivea var. dissecta S. Watson has been tentatively included in P saximontana

(Ertter 2008), but P. saximontana is not otherwise known from the Canadian Rockies. The name P pseudoseri-

cea var. grandiflora Th. Wolf, misapplied by Jepson (1936) to what is now called P. morefieldii Ertter, probably

represents the same entity, and possibly the same Drummond collection.

Although references to Hooker’s herbarium in the protologue might suggest th

pseudosericea var. grandiflora is at Kew, Wolf (1908) is clearly referring to the specimen in his own herbari

now at DR, which is annotated with this name and “NW Amerika/(ex herb. Hooker)/Lag mit echter P. diversi-

folia Lehm. zusammen”. The corresponding sheet in Hooker’s herbarium has a only a single specimen match-

ing the DR specimen of P. pseudosericea var. grandiflora (K000762566), mounted with two large plants of P.

glaucophylla Lehm. (formerly P diversifolia Lehm.). This sheet provides the additional collecting information of

J ” A specimen in the Bentham herbarium (K000762567) is probably a duplicate, in spite

“Rocky Mt/Druir

of Hooker being the only pers

tithe label.

ARCTIC SPECIES IN FNANM

Treatment of arctic variation in sect. Rubricaules is even more problematic than in temperate and subarctic ar-

eas, complicated by the challenge of applying the plethora of names based primarily on Eurasian types in a

region with limited access and a complex post-glacial biogeography. At present, we find only two arctic compo-

nents of sect. Rubricaules sufficiently uniform and widespread in North America to merit full treatment in

FNANM: Potentilla pedersenii (Rydb.) Rydb. and a species to which we provisionally apply the name P uschak-

ovii Jurtzev. These two species constitute the major portion of what has been called P. rubricaulis in arctic

Canada and Greenland (e.g., Porsild 1957; Hulten 1968; Bocher et al. 1978; Porsild & Cody 1980- Hulrtn &

Fnes 1986). The diagnostic morphological characters can be variable and overlapping, but the two species are

nevertheless treated separately in part because of differences in putative parental combinations

707

Potentilla uschakovii Jurtzev, Bot. Zhum. (Moscow & Leningrad) 73:1613. 1988. tvpe: RUSSIA: Wrangel island, “ad

Potentilla uschakovii was described by Jurtzev (1988) as endemic to Wrangel Island, the only place in Asia

where the putative parents (P. subvahliana Jurtzev [sect. Niveae] and P. pulchella [sect. Pensylvanicae]) are sym-

patric. The original description deviates in major features from the arctic Canadian and Greenland plants to

which we apply this name, not least in regularly temate leaves. Our expanded use of the name is based solely

on the interpretation of the same hybrid parentage, since P. subvahliana and P. pulchella are the only represent-

atives of their groups within the American range of plants that we treat as P. uschakovii in FNANM. Morpho-

logic features of North American P. uschakovii that are suggestive of P. pulchella as one parent include more

than three deeply dissected leaflets and leaflet teeth with soft well-developed apical hair tufts. Other features

point toward P. subvahliana as the second parent: caudex branches with persistent whole leaves, smooth petiole

hairs, and one- or few-flowered inflorescences with large flowers. There is much variation among the North

American plants, such that each island or population may have its own features. It is therefore probable that the

species has arisen from numerous hybridization events.

Sojak (1986) interpreted Potentilla pedersenii to have arisen from crosses between P. arenosa subsp. arenosa and

P. pulchella. Features of P. pedersenii that indicate P. arenosa as the sect. Niveae parent include no marcescent

whole leaves, verrucose petiole hairs, and inflorescences with more and mostly smaller flowers than P. uschak-

ovii. Like P. uschakovii, the species is polymorphic and probably the result of multiple hybridizations. It has,

however, a coherent range in arctic North American, including Greenland. Reports of P. pedersenii from north-

eastern European Russia may perhaps involve P. arenosa subsp. chamissonis (Hulten) Elven & D.F. Murray

rather than subsp. arenosa as one hybrid parent.

Jurtzev and Sojak (in Jurtzev 1984) described Potentilla tolmatchevii from northern Asia as a hybrid spe-

cies from P arenosa subsp. arenosa x P. pulchella. Material annotated by Jurtzev as P. tolmatchevii and numerous

collections from Ellesmere Island fit the American concept of P. pedersenii. The two species are therefore

merged by us under the priority name.

Since Potentilla pedersenii was introduced by Rydberg (1908) as a “sp. nov.,” the name is often treated as a

newly described species (e.g., Sojak 1986). However, Rydberg’s inclusion of P subquinata var. pedersenii in syn-

onymy establishes the varietal name as a basionym, making P pedersenii a new combination (K. Gandhi, pers.

comm. 2011).

TAXA NOT GIVEN FULL TREATMENT IN FNANM

In addition to the preceding species being treated in FNANM, there are numerous local or scattered popula-

tions of Potentilla in the Arctic that combine characters from sect. Niveae and sect. Pensylvanicae, or that have

otherwise been included in a broadly defined P. rubricaulis. Some of these may prove worthy of full taxonomic

recognition as species of hybrid origin, as they propagate independently of their putative parents (probably

mainly by agamospermy; cf. Eriksen 1996). Others are too different from place to place to deserve full species

treatment, i.e., they are not fully stabilized or sufficiently widespread. The following such entities— named and

adequately described as species— have been reported from arctic parts of North America and Greenland, but

the evidence is insufficient for full treatment in FNANM.

Journal of the Botanical Research Institute of Texas 7(2)

Sojdk (1985) interpreted Potentilla borealis as the hybrid of P. anachoretica Sojak (sect. Pensylvanicae ) x P. areno-

sa subsp. arenosa. Reports by B.A. Jurtzev (in Elven & Aiken 2007) of this species from the Seward Peninsula

(western Alaska) and Ogilvie Mountains (Yukon Territory) refer to plants with subpalmate to subpinnate

leaves, very slender and silky hairy leaflet lobes, and many glands on the epicalyx bractlets and sepals. The last

feature is not in accordance with the presumed parentage, so the identity of these North American plants as P.

:ev, Arktichesk. Fl. SSSR 9(1):318. 1984. Type: RUSSIA: East Chukotka: “F

tiae sinistrae fl. Czegitun),” 20 Jul 1972, .Jurtzev s.n. (holotypi

?Potentilla murrayi Jurtzev, Bot. Zhum. (Moscow & Leningrad) 78(11):80. 1993. Type: U.S.A.

Jurtzev (1993) interpreted Potentilla murrayi as a hybrid species from P. anachoretica and P. subvahliana. It is a

distinct local entity, forming significant and morphologically consistent populations in a small part of the

Brooks Range. The hybrid assumption is partly supported by morphology: leaf dissection and vestiture resem-

ble P. anachoretica, whereas the influence from P. subvahliana is very evident in its columnar tussocks, leaves,

and flowers. Reports of P. murrayi from outside the central Brooks Range are based on rather different plants,

not forming a morphologically consistent entity. It is debatable whether P. murrayi is distinct from P dezhnevii

of the Russian Far East, since Sojak (2004) suggested that both have the same parentage. If so, P. dezhnevii

would be the priority name.

Potentilla petrovskyi Sojak, Cas. Nar. Mus., Odd. Pdr. 153(2):102. 1984. Type: RUSSIA: South Chukotka: “Anadyr, r-n.

Potentilla tschaunensisjuz. ex Jurtzev, Arktichesk. Fl. SSSR9(1):317. 1

Both of these species, described from the Russian Far East, were interpreted by Sojak (2004) as Potentilla

anachoretica x P nivea si; he accordingly synonymized P. tschauensis under P. petrovskyi. Plants conforming to

Jurtzev’s description are present on the Seward Peninsula, western Alaska. However, in our evaluation these

collections do not constitute a coherent taxon, but rather a gathering of scattered hybrid biotypes. Other re-

ports from northwest North America are based on plants included by Sojak in P. psychrophila.

Potentilla psychrophila Sojak, Thaiszia 16:94. 2007 [2006], Type: U.S.A. Aiaska: ne Brooks Range Lake Peters area Coke

Creek drainage, 69°21'N 144°57W, 29 Jun 1973, Batten 2 50 (holotype: ALA).

The major portions of what had been annotated as Potentilla rubricaulis and P petrovskyi in Alaska and Yukon

Territory were transferred by Sojak (2007) to his new species P. psychrophila , which he assumed to be a hybrid

species fromP.IitomlisandP.nivea. The material annotatedby Sojak is polymorphic and may contain the prod-

ucts of several hybridizations, perhaps among different species. At least one part is morphologically consistent

and is known to represent fairly large populations in central and northern Alaska and in the Yukon Territory,

but whether this is an acceptable hybrid species remains in question.

Potentilla safronoviae Jurtzev & SojaMot. Zhurn. (Moscow & Leningrad) 73:1615. 1988. Type: RUSSIA. Siberia:

PotentiMalyngei Jurtzev & Sojak su bsp. spissa Sojak. Feddes Repert. 117:496. 2006. Type Greenland: Wollaston

r'Yenttlla subsp. spissa based on plants previously identified by Danish botanists

as either P. ntkncuuhs or P. ptdchella. This plant has a significant range and consistent morphology in northeast

Greenland. It is accordingly a candidate for recognition as an independent taxon, but in our understanding not

as a subspec.es of P. lyngei. We agree with Sojak s treatment of P. lyngei subsp. lyngei (sect. PensylvumcueJ as a

isunct taxon in north European Russia, with one close relative in the Russian Far East (P wrangelii V.V. Petro-

709

vsky, Wrangel Island). However, in our interpretation, plants annotated by Sojak as P. lyngei subsp. spissa (oran

unpublished combination as a subspecies of P. insularis ) are probably hybrids between P. pulchella and P

h yparctica Malte (sect. Aureae (Rydb.) Juz.). As such, P lyngei subsp. spissa is a synonym of P. safronoviae, de-

scribed from Siberia.

Potentilla insularis Sojak, Bot. Jahrb. Syst. 106:203. 1986. TVre: NORWAY: Svalbard, Spitsbergen, Hyperithatten, 28 Aug

Potentilla insularis, described from Svalbard and east Greenland, was interpreted by Sojak (1986) as P arenosa

subsp. chamissonis x P. lyngei s.l. Sojak believed P lyngei subsp. lyngei to be the sect. Pensylvanicae parent of the

Svalbard plants, but subsp. spissa (= P. safronoviae) to be the corresponding parent of the Greenland plants. The

hybrid origin of P insularis has been contested for morphological and molecular reasons (Hansen et al. 2000;

Hamre 2000). The Svalbard P insularis, however, has proved very close to Greenland P pedersenii in gross

morphology and to P arenosa subsp. chamissonis in RAPD multilocus phenotypes (Hansen et al. 2000).

Potentilla arctoalaskensis Jurtzev, Bot. Zhum. (Moscow & Leningrad) 78(11):79. 1993. Type: u s a Alaska:

Seward Peninsula, 18.5 miles SW of Deering near Utica Creek, 65°53'N 163°5'W, 23 Jun 1978, Wright 42 p.p. (holotype: ALA).

Potentilla arctoalaskensis is evidently known only from the type, which Jurtzev (1993) interpreted as P arenosa

x P. litoralis. The type sheet is a mixed collection with an unnamed variant of P litoralis that is the common

form in Alaska. Typical P litoralis barely enters Alaska from the southeast.

pentaphylla “Lehm.” and its substitute name P. quinquefolia Rydb. Sojak (1986), for example, included both

names in his synonymy of P hookeriana and designated a lectotype at PR, based on a specimen from the Hook-

er herbarium from “America septentr.” This interpretation results from the traditional attribution of P nivea

var. pentaphylla to Lehmann (Del. Sem. Hort. Hamburg. 1850:12. 1850), sometimes with reference to an earlier

publication (Lehmann in Fl. Bor-Amer. (Hooker) 1:195. 1832). The earlier publication includes a description

but lacks the actual combination, and by the time of the later publication the combination had already been

validly published by Turczaninow. Lehmann’s putative combination is therefore at best an isonym of P. nivea

var. pentaphylla Turcz., as accepted by Wolf (1908). By current taxonomy (e.g., Juzepczuk 1941; Sojak, pers.

comm., 2011), Turczaninow’s type is a variant of P altaica Bunge, endemic to central Asia. Neither the name P

nivea var. pentaphylla nor P quinquefolia therefore has any application in North America. As an alternative in-

terpretation, Porsild (1951) called P. quinquefolia a nomen confusum to be discarded.

Another commonly used name, Potentilla nivea var. subquinata Lange (= P subquinata (Lange) Rydb.), has

generally been considered a heterotypic synonym of the preceding names (e.g., Rydberg 1908, Hulten 1945).

However, we interpret the lectotype designated by Sojak (1986) as the casual hybrid of P. nivea and P. arenosa

that is common throughout the sympatric ranges of these species, for which P. prostrata Rottb. is the priority

name. Since both of the putative parents are in sect. Niveae, the epithet subquinata is not applicable to any mem-

ber of sect. Rubricaules. Normally trifoliolate species in sect. Niveae will occasionally produce supernumerary

leaflets, especially under favorable conditions (Eriksen & Nylehn 1999).

ACKNOWLEDGMENTS

We gratefully take this opportunity to dedicate this article and the pending treatment of Potentilla in FNANM

to the late Boris A. Yurtzev/Jurtzev (1932-2004; LE) and Jin Sojak (1936-2012; PR), whose joint expertise was

critical to synthesizing the previously independent taxonomic traditions in American and Eurasian Potentilla.

Without the resi

collaboratively \

treatment would be seriously deficient. This has been particularly true for

ixtensively revised as a result of recent trans-Beringian and trans-Atlantic

analyses.

We are likewise indebted to Bente Eriksen, Mats Topel, and Christoph Dobes for sharing both their

friendship and their latest research on the molecular phylogeny of Potentilla. A comparable debt for assistance

in resolving nomenclatural quandaries is owed to Kanchi Gandhi, James L. Reveal, John McNeill, and Dick

Brummitt. We are most grateful to Arnold Tiehm (RENO) and an anonymous reviewer for detailed reviews and

Support to the senior author from the Lawrence R. Heckard Endowment Fund of the Jepson Herbarium is

gratefully acknowledged, as is the University of California Botanical Garden for the maintenance of living col-

lections. The University and Jepson Herbaria at the University of California at Berkeley, the Natural History

Museums of the University of Oslo, the University of Alaska Museum of the North, the Snake River Plains

Herbarium at Boise State University, the Missouri Botanical Garden, and the National Museum in Prague Her-

barium are to be thanked for use of facilities and staff support, as are ALTA, ARIZ, BRY, CAN, COLO, DAO, DR,

GH, K, MONTU, NY, OSC, RM, US, UTC, and WTU for providing loans used in this research.

e & Sons Forlag, Copenhagen.

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BOOK REVIEW

Jane Goodall with Gail Hudson and forward by Michael Pollan. 2013. *

from the World of Plants. (ISBN-13: 9781455513222, hbk.). Grand Central Publishing, Hachette B

Group, 237 Park Avenue, New York, New York 10017, U.S.A. (Orders: www.hachettebookgroup.com/

publishers/grand-central-pubhshing). $22.36, 384 pp., 36 color photos, many b&w photos, 6" x 9".

In the early 1970s my father arrived home one evening with two tickets to attend a talk by Jane Goodall spon-

sored by the Appalachian Mountain Club of Boston. To a budding 12-year-old environmentalist, the opportu-

nity to see Jane Goodall, recently back from Gombe, and to hear her in person talk about her experience in Fast

Africa was a dream come true. Goodall was a personal heroine then and has been an inspiration since, and it

was with great joy that I jumped at the opportunity to review her recent book Seeds of Hope.

Known primarily for her work with chimpanzees, Jane Goodall has written a new book, Seeds of Hope:

Wisdom and Wonder from the World of Plants, that in many ways serves as a capstone to her life’s work as in it she

reflects upon her lengthy career and the impact that plants have had on her life and research. From a child

climbing the Beech tree in the family yard, to the forests of Gombe, and to her present work as a voice for the

environmental movement, Goodall reflects on the plant kingdom, weaving in stories from her own experi-

ences and from the many collaborations she has had over the years. In fact, she refers to her book as a gathering

Seeds of Hope is divided into four parts. It begins with a look at Goodall’s childhood home, “The Birches”

lumemouth, England, and how her “love for the natural world” developed. She then spends time discuss-

ing the plant kingdom, providing us with an overview of the structure and function of plants, and offers us a

glimpse of the great variety of plants around the world from the small, hardy tundra plants of Greenland and

ush forests of the Sumatran jungles, to those “unwanted weeds” that grow in our gardens. Goodall then

s to provide us with a history of plant collecting and its manifestations today. She begins with Carl Lin-

ts and his intrepid explorations through unmapped Lapland and brings us through to the 21st century to

:urrent “plant hunters” who work with botanical gardens and research institutes around the world to “col- J

ect, protect, cultivate, and propagate” plants, organizations such as the Kew Royal Botanical Gardens in Eng-

and and the Botanical Research Institute of Texas.

Part two of the book addresses the human uses of plants. In this section she discusses medicinal plants

md the importance of indigenous knowledge in identifying plants for medicinal purposes. She hig hlig hts two

ndividuals, Mark Plotkin in the Amazon and Vandana Shiva in India, both of whom have committed their

lives to working with indigenous groups to learn about medicinal plants and to ensure that indigenous knowl-

edge is not exploited by western pharmaceutical companies through what Shiva terms “biopiracy.”

In part three, Goodall turns to the history of plantation farming, focusing on the development of tobacco,

rice, cotton, wheat, potatoes, and com. Throughout she illustrates how the cultivation of these monocultures

“provided huge economic benefits to plantation owners while causing massive human suffering (including

health issues and child labor) and environmental degradation (including habitat and water loss and increased

pesticide, herbicide, and fertilizer use).” She then turns to a discussion of genetically modified organisms

(GMOs), addressing the genetic alteration of plant DNA and describing it as “a monstrous crime against

plants.” She highlights the development of the “super pests” and “super weeds,” the result of creating insect-

and herbicide-resistant plants. Goodall paints a dark picture of the multiple risks of GMOs to the health of

plants, animals, and humans.

Goodall, however, is an optimist She doesn't leave us in the dumps feeling there is nothing that can be

done. Instead she offers ushope. Inpart four, she provides examples of organically grown tea, coffee, and cacao.

demonstrating sustainable farming methods. She p

nity gardening.

PREVIOUSLY UNRECOGNIZED TYPES OF AMERICAN ACACIA SPECIES

FROM THE TORINO HERBARIUM (TO)

David Seigler

Department of Plant Biology

University of Illinois

Urbana, Illinois 61 801, U.SA.

seigler@life.illinois.edu

Laura Guglielmone

Department of Life Sciences and Systems Biology

Universitd degli Studi di Torino

V. le Mattioli 25, 10125 Torino, ITALY

John Ebinger

Emeritus Professor of Botany

Eastern Illinois University

Charleston, Illinois 61 920, U.SA.

jeebinger@eiu.edu

ABSTRACT

RESUMEN

The Herbarium Universitatis Taurinensis (TO) is a rich source of type materials for species that have been

placed in the genus Acacia. In the process of identifying type materials for species originally described as Aca-

cia sensu lato (s.l.) or at some time transferred to that genus, a search was made of herbarium materials of Colla,

de Spin, and Martius at TO. Previously unrecognized probable and potential type materials were discovered.

In order to put these findings into proper context, some historical background concerning the collectors, per-

sons associated with the collections at TO, and the history of the herbarium are needed.

History of Herbarium Universitatis Taurinensis (TO)

The Herbarium of the University of Turin (TO) was established in 1891, about 160 years after the foundation of

the Orto Botanico of the Universita di Torino (1729), and is one of the most important Italian herbaria with

about one million 1,000,000 specimens. In addition to two phanerogamic collections, Herbarium Pedemonta-

num (TO-HP, about 112,000 specimens) and Herbarium Generale (TO-HG, about 260,000 specimens), the

Herbarium possesses several important collections, such as Carlo Alhoni’s herbarium, from which materials

are not available for loan or exchange (Vignolo-Lutati 1951, 1952). These exsiccata or dried specimens docu-

ment more than 250 years of botanical studies including those of the first floristic research in Piedmont

(Piemonte), carried out by Carlo Allioni (1728-1804) and his collaborators (Allioni 1785), as well as several

important early expeditions including that of Vitaliano Donati (1717-1762) to Egypt (1759-1762) (Fomeris et

al. 2008), H.R.H. Luigi Amedeo di Savoia, Duke of Abruzzi, to Ruwenzori (1906) and the Wabi Scebele in

Ethiopia (1928-1929) (Mattirolo 1911; Fomeris & Montacchini 1984; Guglielmone 2004).

714

A number of American collections ofJ.J. Bemhardi, L.A. von Chamisso and A.R. Delile, (North America),

C. Bertero and F.W. Sieber (Antilles, South America), A.M.A. Bonpland (Central America), C.F.P. Martius and

G. Casaretto (Brazil), from the early 1800s, including those are conserved in the herbaria of Giovanni Battista

Balbis (18,000 specimens) and Luigi Colla (10,000 specimens) (Guglielmone et al. 2009; Baldini & Gugliel-

mone 2012).

Giovan Battista Balbis (1765-1831)

Giovan Battista Balbis, a member of prestigious Italian and European Academies, was an important botanist of

the 19th century; he was a correspondent with Augustin Pyramus DeCandolle, Jacques J. Labillardiere and

Kurt Sprengel, among others. Balbis was a student of Allioni; he graduated in medicine in 1785 and in 1794 he

was a physician in Napoleon’s Army in Italy. In 1801, Balbis was appointed Professor of Botany and Director of

the University Botanic Garden in Turin, posts he held until 1814 when, after the defeat of Napoleon and the

restoration of the Savoia monarchy, he was exiled. In 1819, Balbis served in the same role at the University of

Lyon, where he remained until 1830, when his poor health obliged him to leave France and return to Piedmont.

He died in early 1831 (DeCandolle 1831; Colla 1833; Stafleu & Cowan 1976:107-109; Fomeris & Pistarino

1990). Although Balbis’ studies involved the Italian and French flora [Flora Ticinensis (Nocca & Bal-

bisl816-1821)], written with the collaboration of Domenico Nocca (1758-1841), and Flore Lyonnaise (Balbis

1827-1828), between 1801 and 1814, he published 14 catalogs of the Botanic Garden of Turin and one of the

Garden of Lyon in 1826. In these publications, he added the descriptions of some new species that were culti-

vated in the Gardens. Under his direction, the number of species cultivated in the Botanic Garden of Turin in-

creased considerably. In 1801, there were more than 3,500 species, but in 1812 this number had grown to about

6,000 (Balbis 1801; 1812; 1813). Under his direction, there was an increased interest in exotic flora and the fo-

:xpanded from the geographic limits of the Piedmont region to a worldwide perspective,

e built, where many plants from Africa and America were cultivated; for some of these

species this represented the first introduction into Italy (Saccardo 1909; Maniero 2000). Many of these collec-

tions were obtained by personal exchanges with other Italian and European botanists and either University or

private Gardens. The richness of plants cultivated in the Botanic Garden is documented by the drawings col-

lected in Iconographia Taurinensis (Fomeris 2008). This collection, 7640 drawings in 64 volumes, represents

plants cultivated in the Garden between 1752 and 1868, the year when the last painter died (see Chiapusso Voli

1904). Many plants cited in the Balbis’ catalogs are included in volumes 35 to 47 of this series (Fomeris 2008).

Balbis made a remarkable herbarium, considered one of the most important of the time; the collection

includes the plants he collected as well as specimens obtained in exchange from other botanists. In 1831, after

Balbis’ death, his herbarium was bought by Giuseppe Giacinto Moris (1796-1869), director of the Botanic Gar-

den in Turin. Because the University did not hold a collection of dried specimens ( exsiccata ) at that time, Balbis’

materials were the first holdings of the current herbarium (Mattirolo 1929).

Carlo Giuseppe Luigi Bertero (1789-1831)

Carlo Giuseppe Luigi Bertero, a correspondent and close friend of Balbis, was one of the first Italian naturalists

to visit the New World, collecting a large amount of botanical material from little known areas. Bertero was

born on 14 October 1789 in Santa Vittoria d’Alba (Piedmont, near Turin). He studied Medicine at the University

of Turin, where he attended lessons of botany at the Botanic Garden by Balbis. After his graduation in 1811, he

practiced the medical profession for several years.

In 1816, Bertero moved to Paris where he met many important botanists such as Rene Louis Desfontaines,

lean LouisUiseleut-Deslongchamps and, especaily, Christiaan Hendrik Persoon; this botanist helped h,m tn

TL ng * r “ d “Sis'edhim in obtaining an appointment a, a ship’s doctor on the ship

Guadelupe, which sailed for Martinique in August 1816.

The details of this first expedition results were carefully recorded in Berteto’s manuscript, a field book in

about the localities visited and the plants observed, about his correspondence with Balbis and

it is collections. These documents trace the following itinerary: Guadeloupe (1816-1818),

Seigler etal., Types of American Acacia species

715

Saint Thomas (1818), Puerto Rico (1818-1819), Santo Domingo, Haiti (1819-1820), Colombia (Santa Marta,

Barranquilla, Mompos and part of Rio Magdalena) (1820-1821), and Jamaica (1821) (see also Urban 1902).

In 1821, Bertero returned to Paris, but later returned to Turin where he met with Balbis in order to study

his new collections. Balbis sent many of Bertero’s specimens to Kurt Sprengel in Berlin for identification, but

Bertero’s field book was sent to A.P. DeCandolle in Geneva; in this field book, 1746 species are cited; most of

these are described and the morphological details illustrated. DeCandolle (1825) included descriptions of the

new Caribbean taxa reported by Bertero in his Prodromus. In 1857, Alphonse DeCandolle returned Bertero’s

manuscript to Turin. At present, the largest part of Bertero’s collections from the West Indies and Colombia,

about 2,000 taxa, is conserved at TO in Balbis’ and Colla’s collections. Duplicates of Bertero’s material are also

found in other herbaria including B, FI, HAL, L, M, MEDEL, MO, MPU, NY, P, P-JU, S, WB.

In 1827, Bertero returned to Paris where he planned his next expedition to Chile, following the sugges-

tions made by A.P. DeCandolle and Benjamin Delessert to explore a land for which the flora was poorly known.

In October 1827, he embarked on a ship from Le Havre to Valparaiso, Chile, again as a ship’s doctor. Recon-

struction of this second journey is complex: data about the itinerary are based only on specimen labels and

Bertero’s correspondence with Balbis and Colla. The Chilean localities visited include: Valparaiso, Rancagua,

Quintero, M.te La Leona, M.te la Punta des Cortes, Quillota, Tagua Tagua lagoon, Concoa River, Rio Claro, and

Cachapoal.

In 1828-1829, Bertero published a list of several species he had observed in the newspaper Mercurio

Chileno. In this list, he proposed several new species, but unfortunately because he did not add descriptions,

these epithets are nomina nuda. In early 1830, Bertero went to the Juan Fernandez Islands (Isla Mas a Tierra)

with the English botanist A. Caldeleugh. On September 28, 1830, Bertero sailed from Chile to the Society Is-

lands with the General Consul of North America J. A. Morenhout and on November 4, 1830, he arrived in Ta-

hiti, where he collected plants actively for a few months. He departed on April 9, 1831, for Valparaiso, but died

in the shipwreck of his boat near Raiatea island (for detailed information about Bertero’s biography see: Matti-

rolo 1932; Vignolo-Lutati 1955, 1956; Delprete et al. 2002).

Before his final unlucky voyage, Bertero sent his Chilean collections to Baron Delessert in Paris who dis-

tributed duplicate sets of exsiccatae to Balbis, Colla, and A.P. DeCandolle, but kept back the rest of these collec-

tions (about 15,000 specimens) for Bertero. Several years after Bertero’s death, these were sold by Delessert’s

heirs to a Travel Company of Esslinger owned by E. Steudel and C.F. Hochstetter (1840); later these materials

were dismantled and dispersed. Bertero entrusted his Tahitian collection to Morenhout, who in 1834 sent

these materials to A. Dessalines d’Orbigny in Paris, although by that time, a portion of this material was miss-

ing (Guillemin 1836, 1837). Despite the fact that J.A.Guillemin reported that some duplicates were sent to the

Royal Academy of Turin, no Bertero specimens from Tahiti were conserved in Turin. At the present time,

Bertero’s Chilean material (about 300 specimens) are found in Colla’s herbarium,

iuigi Colla (1766-1848)

Luigi Colla was a lawyer and an expert botanist whose botanical knowledge was appreciated by the most im-

portant botanists of that time. A.P. DeCandolle, J. Lindley and K. Sprengel, named new genera after him. Colla

was a member of prestigious Italian and European academies; in 1822, he even became a member of the Acad-

emy of Natural History of Philadelphia. These affiliations permitted him to have many correspondents with

whom he made exchanges of plants and exsiccata. Colla published several works in the Memorie della Reale

Accademia delle Scienze di Torino between 1820 and 1848, including monographs of exotic genera such as the

genus Musa, which included the description of two species, Musa balbisiana and M. acuminata (Colla 1820).

Colla established a botanic garden in Rivoli (near Turin), the Hortus Ripulensis, for which he published catalogs

of plants cultivated between 1824 and 1831 (Colla 1824, 1827a, 1827b, 1829, 1831). In these catalogs, Colla also

reported descriptions of new species, often with the respective drawings made by his daughter Tecofila Colla

Billotti. In 1829, the number of species cultivated in the garden increased to more than 2000 species belonging

to approximately 700 genera (Colla 1831): the greatest part of these were exotic and included a number of

Plants obtained from seeds sent by Bertero from the Antilles.

716

al Research Institute of Texas 7(2)

Colla was a close friend of Bertero and, after Bertero’s death, Colla published descriptions of the new Chil-

ean genera and species based on Bertero’s specimens collected during the second expedition, including those

listed by Bertero in 1828-1829 in the Mercurio Chileno. In 1834, Colla (1834a) published a paper entitled “Plan-

tes rariores in regionibus Chilensibus a clarissimo M.D. Bertero nuper detectae”, in which he described several

montanum (Colla 1833-1837) (for more details about the priority of publications see Pichi-Sermolli 1951,

1952). In both works Colla enclosed drawings of new taxa made by his daughter. Two more species from

Bertero’s exsiccata were published by Moris (1834, 1835).

plants grown in his botanic garden, Bertero’s expeditions, and exchanges with several Italian and European

botanists including specimens from Giovanni Biroli (1772-1825) from Piedmont (Guglielmone, 2008), Jacob

Corinaldi (1782-1847), from Egypt (Fomeris et al. 2008), Carl Friedrich Philipp von Martius (1794-1868)

from Brazil (Fryxell 1976; Stafleu & Cowan 1981:325-339), and Maximilian A.P. von Wied-Neuwied (1782-

1867) (P.L.R. de Moraes, personal communication). On April 25, 1849, after Colla’s death, his son donated his

collection to the University of Turin; the catalog of this herbarium, handwritten by Colla himself , accompa-

nied the specimens (for more detailed information about Colla’s biography see: Parlatore 1850; Delponte 1852;

Mattirolo 1929:44).

Marquis Luigi Raimondo Novarina di Spigno (1760-1832)

Marquis Luigi Raimondo Novarina di Spigno (1760-1832), known as Marquis De Spin, was a passionate bota-

nist who established an important garden in San Sebastiano Po (near Turin) at the beginning of the 19 th cen-

garden in seven catalogs published between 1804 and 1823. He exchanged living plants and exsiccata with

several correspondents, among these were Balbis and Colla. The international relationships that he main-

tained allowed a constant and conspicuous increment of the species grown in the San Sebastiano garden;

moreover he acclimatized and introduced into cultivation several exotic species that were subsequently sent to

many Piedmont gardens.

De Spin acknowledged his gratitude to Bertero for the seeds, material and information obtained from him

through Balbis in his catalog of 1823. De Spin prepared specimens from several plants in his garden. Approxi-

mately 700 specimens based on Marquis’ living collections, now mostly included in Balbis’ herbarium, have

been identified in TO (for more detailed information about De Spin biography and collection see: Guglielmone

et al. 2006).

History of Acacia

As conceived by Willdenow (1806, 1809), Kunth (1823, 1825), DeCandolle (1825) and other early taxonomists,

the genus Acacia was quite diverse; many of the species now belong to other genera of mimosoid legumes,

among them Mimosa (Bameby 1991), Calliandra (Bameby 1998), Pithecellobium, Albizia, and Lysiloma (Barne-

by & Grimes 1996, 1997). Based largely on the concepts of Bentham (1842, 1875, 1876), the genus Acacia later

was limited to species with numerous stamens (20-200) and filaments free to the base. This concept remained

largely the same until the 20 th century when Vassal (1972) and Pedley (1978) refined the subgeneric treatment

of this rather large genus. Pedley (1986) suggested that the genus Acacia should be divided into three genera,

Acacia, Senegalia, and Racosperma. Nonetheless, most workers continued to accept Acacia s.l. until Maslin et al.

(2003) suggested that the time had come to depart from this viewpoint.

The taxonomy of Acacia became more contentious when in 2005, the International Botanical Congress in

Vienna approved a proposal to change the type of the genus Acacia Miller from an African species A scorpioi-

des (L.) W.F Wight [- A. nilotica (L.) Delile; Acacia subgen. Acacia) by recognizing an Australian species A.

penmnervis [Acacia subg. Phyllodineae] as a conserved type (Orchard & Maslin 2003; McNeill et al. 2005). Ac-

ceptance of this retypification remains controversial (Brummitt 2011; Linder & Crisp 201T Luckow et al.

2005; Moore & Cotterill 2011; Moore 2007, 2008; Moore et al. 2011a; Moore et al. 2011b; Rijckevorsel 2006;

Seigler et al.. Types of American Acacia species

717

Smith & Figueiredo 2011; Smith et al. 2010; Smith et al. 2006; Thiele et al. 2011). However, the retypification

was upheld recently at the XVIII International Botanical Congress in Melbourne (McNeill & Turland 2011;

Apart from these arguments, both morphological and molecular data for Acacia s.l. (Acacieae), Ingeae,

and Mimoseae, strongly support segregation of Acacia s.l. into at least five entities: the genera Vachellia (Seigler

& Ebinger 2005; Kodela & Wilson 2006), Senegalia (Seigler & Ebinger 2009, 2010; Seigler et al. 2006a), Acaci-

ella (Britton & Rose 1928; Rico-Arce & Bachman 2006), Mariosousa (Seigler et al. 2006b), and Acacia (primar-

ily Australian species of the former subgenus Phyllodineae ).

Acacia specimens in TO

A number of types or possible type material for American species of Acacia is found at TO, although many of

these are now recognized as members of other genera. Others were originally described in other genera, but

were later considered to be members of Acacia s.l. There are approximately 500 specimens of Acacia presently

assigned to 198 taxa in the “Herbarium Generale.” Several of these are included in the collections of herbaria of

Balbis and Colla.

Although Balbis listed 164 specimens of Acacia in the catalog of his herbarium, only 117 are presently

found in the collection; among these are 36 specimens sent by Bertero from Guadeloupe in 1819, Santo Do-

mingo and Puerto Rico in 1820, and Santa Marta (Colombia) and Jamaica in 1821. Dates on the labels refer to

the years when Balbis received the specimens (Vignolo-Lutati 1955). There are also eight specimens from the

Botanic Garden of Turin (between 1800 and 1813) and six from De Spin’s garden (between 1818 and 1825) (the

Marquis listed 47 species of Acacia in his catalogs).

The catalog of Colla’s herbarium includes 104 Acacia specimens; presently only 96 are found. Among

these, nine specimens from “Herbarium Martii,” probably represent duplicates of Martius’ collections; six of

these likely came from collections of von Wied-Neuwied (P.L.R. de Moraes, personal communication). There

are also 36 specimens cited in Hortus Ripulensis; Colla listed 50 specimens of Acacia in the catalogs of his gar-

den. Thirty-four more exsiccata are from the West Indies with labels handwritten by Bertero.

A careful search of materials at TO reveals specimens associated with many of the historically important

figures outlined above, that are either original materials examined by these botanists, or in a number of cases

type materials. Most of these specimens have not been considered in recent taxonomic studies involving mi-

mosoid legumes. The status of these exsiccata is discussed below:

anthera Schult. ex Colla, !

U.S OF ACACIA SENSU LATO AT TO

3. pedem. 2:266, n. 71. (Fig. 1). Nom. illeg. non Zeyh. ex Steud.

Neither flowers nor fruits are mentioned in the original description; the country of origin is not known. The

specimen lacks prickles and spines; the leaves are bipinnate with 2-3 opposite pairs of pinnae and 12 pairs of

leaflets per pinna. The rachillae of the pinnae are winged. The leaflets are opposite with more or less central

venation (Fig. 1). Colla (1834b 2:266, n. 71) notes that there is a gland at the base of the petiole. The gland ap-

gume. We cannot identify it to genus.

The collector was probably Joseph August Schultes (1804-1840), a botanical collector for Roemer or his

father Julius Hermann Schultes (1773-1831), an Austrian botanist in Brazil. Most of their collections were in

northeastern Brazil, especially Pernambuco.

it Catalogue 27. 1818. (Fig. 2). f

5 (Jacq.) 1

Seigler et al., Types of American Acacia species

Fk ‘ 1 0ri 9' nal materials of Acacia alba De Spin.

The species was listed and described in the catalog of S. Sebastiano garden (De Spin 1818, pp. 5, 27, note 1).

Moreover, it is described in the second appendix to the catalog of Hortus Ripulensis (Colla 1827b:339, note 1);

the plant was received from a garden sited in Buttigliera (on the hills near Turin) owned by Count Francesco

Lorenzo de Freylino (1758-1820) and at that time directed by M. Pangella. This garden was very important in

the 18 th century, particularly for the collections of exotic plants. In the three catalogs of the garden (Freylino

1785, 1808, 1810) this species was not listed. Colla (1834b 2:362, n. 33) also cited this species. This species was

early accepted (Steudel 1841, Bentham 1875; for complete descriptions and distribution see Bameby 1998;

Bameby & Grimes 1996; Hernandez 1986, 1989; Zuloaga 1999).

». pedem. 2:268. 1834. (Fig. 3). Senegaliaangico (I

gico;” TO-HG, herb. Colla ex herb. Martius.

Although Colla’s handwritten label confirms the earlier

the description (Colla 1834b) agrees more closely with the type of Acacia pirn

Colla did provide a Latin binomial and at least a generally a

was validly and effectively published. Further, the specimen at TO represents materials seen by Colla and we

designate that specimen as a lectotype for the species. Because the name of the species follows the type speci-

men, not the accuracy of the description, that name must be accepted.

Bentham (1876) considered Acacia angico Martius to be a synonym of Piptadenia rigida Bentham (1842)

[now recognized as Parapiptadenia rigida (Benth.) Brenan (1963)] suggesting that Bentham was influenced by

Colla’s descriptions, although Colla was not cited. A Martius specimen of authentic Acacia angico at Kew

(K264972), labeled as such in Martius’ handwriting and apparently identical to material seen by Colla at TO

was annotated by Bentham as Acacia plumosa Lowe. The descriptions of other authors are often based on er-

roneously interpreted materials of Acacia angico Martius in Colla (Boggan et al. 1997; Rico-Arce 2007; Bameby

This species was listed in the catalog of San Sebastiano garden (De Spin 1823) and described by Colla (1824:1,

note 2). In 1821, Bertero (1816-1821) recorded the description of the species in his fieldbook, in fascicle 13 of j

“Stirpes in Provincia S. Marthae Continent. Amer. Austral. Lectae 1821” (pp. 1042-1043, n. 1693). The four |

specimens can be considered as syntypes. Sample 2a reports the original collection number by Bertero and is

selected as the lectotype (Fig. 3). This species is discussed in more detail by subsequent authors (DeCandolle

1825; Mabberley 1981; Rico-Arce 2001).

s Colla, Mem. Reale Accad. Sci. Torino 33(1):135. 1829. (Fig. 5). Nom. illeg. non Sprengel (1826).

The specimen was from a plant cultivated in the garden owned by the Litta family in Lainate (near Milan) and

given to Colla by the gardener Giuseppe Tagliabue. The species was described in the third appendix of Hortus

Seigler et al., Types of American Acacia species

Seigler et al.. Types of American Acacia species

723

724

Ripulensis (Colla 1829:135, note 1) and listed in Herb. Ped. (Colla 1834b, 263, n. 47). Colla reported that the

plant has two glands on the petiole (Colla 1829:135, note 1), but we were unable to observe petiolar glands on

the specimen examined. A drawing of this species was found in Iconographia Taurinensis (Vol. 51, t. 16); the

volume is estimated to be dated between 1829 and 1831 (Fomeris 2008). These characters are compatible with

the genus Mimosa.

6. Acacia compta Mart, in Colla, Herb, pedem. 2:268. 1834. (Fig. 6). Type: brazil: “Acacia compta Man./s. Pedro d’Algoa

The flowers of the spicate inflorescences have 8-10 stamens. The leaflets have subcentral venation with con-

spicuous venation on the adaxial side. The prickles are paired at the nodes (Fig. 6). There is a small gland at the

base of the petiole (Colla 1834b). These characters are compatible with Adenopodia, Mimosa or most probably a

Piptadenia species. The specimen conserved in TO-HG is evidently part of the type material. There is no indi-

cation that the specimen in TO was the only material used by Martius, thus it is not considered as the holotype,

but designated as the lectotype (ICBN art. 9 note 1).

7. Acacia lasiopus Mart, in Colla, Herb, pedem. 2:267, n. 74. 1834. (Fig. 7). Type: brazil “Acacia lasiopu- Mart /AdFl.

The flowers of this specimen have approximately 50 stamens. According to Colla (1834b), the leaves have 4-6

pairs of elliptic leaflets and the species is eglandular. This specimen appears to be a mimosoid legume of tribe

Ingeae, but the paucity of characters does not permit further assignment of this taxon.

Although Colla (1834b) states that there are no petiolar glands, they are evident on the specimen itself. Cam-

pos Novos is in current-day Sta. Catarina. Based on Burkart (1979), this specimen, which is unarmed (Fig. 8),

may be Parapiptadenia rigida (Benth.) Brenan, which has been collected there in recent years. Perhaps other

genera of tribe Mimoseae should be considered as well.

myriophylla;” TO-HG, herb. Colla.

The specimen conserved in TO-HG is evidently part of the type material. There is no indication that the speci-

men in TO was the only material used by Martius, thus it is not considered as the holotype but designated as a

lectotype (ICBN art. 9 note 1). 6

Colla (1834b)

fig. 1). He noted that there

Seigler et al.. Types of American Acacia species

725

f* 6 - *■ Lectotype of Acacia c

Seigler et al., Types of American Acacia species

729

or possibly even 5) indicates that this species belongs to the Mimoseae, perhaps to the genera Parapiptadenia or

Pseudopiptadenia (Fig, 10).

10. Acacia plumosa Mart, ex Colla. Herb, pedem. 2:267, Jul 1834. (Fig. 11). Type: BRAZIL: “Acacia/ Villa Nova do Ver-

This specimen has spicate inflorescences and prickles, some of which appear to be paired at the nodes, but are

also scattered on the petioles. The glands are depressed at the base of the petioles. Leaflet venation is more or

less central. These characters are quite similar to those of Piptadenia trisperma (Veil.) Benth.

Although Colla’s handwritten label gives the name Acacia plumosa on the label written by Martius, the

description written on the reverse side of the Martius label (Colla 1834b) agrees more closely with the type of

Acacia angico Martius in Colla (see above). Because Colla did provide a Latin binomial and at least a generally

accurate description and diagnosis, Acacia plumosa was validly and effectively published. Further, the speci-

men at TO represents materials seen by Colla and we designate that specimen as a lectotype for the species.

Because the name of the species follows the type specimen, not the accuracy of the description, that name must

be accepted.

The fact that Bentham (1876) considered Acacia angico Martius to be a synonym of Piptadenia rigida Ben-

tham (1842) [now recognized as Parapiptadenia rigida (Benth.) Brenan (1963)] and a Martius specimen of au-

thentic Acacia angico at Kew (K264972), labeled as such in Martius’ handwriting and apparendy identical to

material seen by Colla at TO was annotated by Bentham as Acacia plumosa Lowe suggests that Bentham was

influenced by Colla’s descriptions, although Colla was not cited.

11 . Acacia pterocarpa Bertero in Colla, Herb, pedem. 2:265. 1834. (Fig. 12 ). Type: “Acacia pierocarpa?/es«n. Miss. A

12 . Acacia ramosissima Mart, in Colla, Herb, pedem. 2:268. 1834. (Fig. 13 ). Type: brazil: “in sylvis praemitivis Brasil

There are no petiolar glands. The number of stamens varies from 7-9. This species might possibly be Piptadenia

ramosissima (Bentham 1842; Lewis 1987), but alternatively could be a Mimosa species.

13- Acacia rubiginosa Martius in Colla, Herb, pedem. 2:268. 1834. (Fig. 14 ). Type: BRAZIL: “Mimosa/ Campos des

This specimen may have been collected at Campos de Goytacazes or Goitacazes (Goytacasen in German), a

ocalitym the northeastern part of present-day Rio de Janeiro, Brazil. Based on the image of the specimen and

3 later % ure (Colla 1837, tab. 62, fig. 2; Fig. 10)) this is clearly a Mimosa species and resembles Mimosa pellita

Humb. & Bonpl. ex Willd., but cannot be ascribed to that species with certainty. Other possible, but less likely,

s P € aes might be M. tarda Bameby or M. elliptica Benth. (Bameby 1991).

730

Journal of the Botanical Research Institute of Texas 7(2)

7, vol.8,tab.62.

Seigler et al., Types of American Acacia species

Journal of the Botanical Research Institute of Texas 7(2)

735

aspini Balbis ex De Spin (upper left).

736

Journal of the Botanical Research Institute of Texas 7(2)

738

14. Acacia spini Balb. ex De Spin, Supplement au Catalogue des Plantes du Jardin de St. Sebastien 8. 1823. (Fig.

In the original publication De Spin (1823) did not explicitly designate any specimen. Art. 9.2 of the code explic-

itly states that the “the original material” (from which a lectotype can be designated) “comprises: (a) those

specimens and illustrations (both unpublished and published either prior to or together with the protologue)

upon which it can be shown that the description or diagnosis validating the name was based...” However, all

specimens conserved at TO are dated after 1823. No material prior to that date was preserved, and possibly no

herbarium sheet existed at the moment of the description, as the name was apparently based on living speci-

mens. Therefore, one of the cited specimens can be selected as a neotype of Acacia spini. We select the specimen

labelled “S. Sebast. D. Spin 1826” from Balbis’ herbarium because it was taken from a plant grown in S. Sebas-

tiano Po, thus very likely the plant examined by De Spin for the description. Type locality: De Spin did not in-

dicate any locality. Colla (1829) re-described the species and indicated that the seeds had been collected in

Guadalupe near La Basse-Terre. Colla also included a drawing of this species made by his daughter T. Billotti

(t. 5). Another drawing of the species is in lconographia Taurinensis (vol. 51, tab. 10) (Fig. 16).

15. Acacia velutina Bertol., Syll. Pi. Hort. Bot. Bonon. 1827:3. 1827. (Fig. 17). Nom. illeg. nonDeCandolle (1825 ). Type

ACKNOWLEDGMENTS

The authors wish to thank several colleagues for advice concerning questions of nomenclature and general

taxonomic advice. Among these are: J. Leland Crane, Bruce R. Maslin, Fred Barrie, K.N. Gandhi, Victoria Hol-

lowell, Duane Kolterman, Mario Sousa, Charlotte Taylor, Gordon Tucker, Paul Wilson, and James Zarucchi.

We appreciate the input of Joseph T. Miller and an anonymous reviewer. We wish to acknowledge support by

the National Science Foundation (NSF BSR 82-15274 and NSF-PCM 82-17114, NSF 04-15803), United States

Department of Agriculture (OICD 58-319R-0-011), the University of Illinois Research Board (1994, 2001), a

Rupert Barneby Award by the New York Botanical Garden (1997), and the American Philosophical Society

(1992). We also acknowledge the assistance of Professor Pedro Luis Rodrigues de Moraes Universidade Esta-

dual Paulista “Julio de Mesquita Filho,” Instituto de Biociencias, Departamento de Botanica, Rio Claro, SP,

Brazil with regard to von Wied-Neuwied material.

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A NOMENCLATURAL NOTE ON

BIDAR1A INODORA AND B. TINGENS (APOCYNACEAE: ASCLEPIADOIDEAE)

Debjyoti Bhattacharyya and Lalawmkima Darlong

Department of Life

Assam Ui

Sikhar 78801 1,

e Flora of China, and o

B. inodora is the correct nami

Bidaria (Endl.) Decne

a Flora of China, y de B. (ingens (Roxb.) Dec-

In their treatment of Asclepiadaceae oflndia, Jagtap and Singh (1999) listed Bidaria tingens (Roxb.) Decne.

(based on Asclepias tingens Roxb.) as an accepted name with Cynanchum inodorum Lour, as a synonym. In con-

trast, the Asclepiadaceae treatment in Flora of China (Li et al. 1995) has Gymnema inodorum (Lour.) Decne.

(based on C. inodorum) as the accepted name with B. tingens as a synonym. Although the treatments of the

Chinese and Indian floras differ in their taxonomy for this species, their different usage of epithets for the ac-

cepted name posed a puzzle and led to the following study.

Decaisne (1844) elevated Gymnema [unranked] b. Bidaria Endl. to the rank of a genus as Bidaria (Endl.)

Decne. Within his new genus, Decaisne included five species [including B. inodora (Lour.) Decne. and B. tin-

gens (Roxb.) Decne.]. Although he did not cite a type species for his new genus, it is automatically typified by B.

tingens. This is because, for his infrageneric name Gymnema b. Bidaria, the basionym of Bidaria, Endlicher

(1838) included a single species, i.e., Asclepias tingens Roxb., which is the automatic type species of the preced-

ing infrageneric name.

The type species name Bidaria tingens is not the oldest within the genus. The priority of B. tingens, which

is based on Asclepias tingens starts from 1815. The priority of Cynanchum inodorum, however, starts from 1790.

Therefore, if A. tingens and C. inodorum are conspecific, then the correct name for this complex in the genus

Bidaria or Gymnema must employ the epithet inodora. It is evident that the Flora of China (Li et al. 1995) is cor-

rect in its usage of G. inodora, whereas Jagtap and Singh (1999) erred in their usage of B. tingens as the accepted

name. It is speculated here that Jagtap and Singh (1999) might have erroneously assumed that the type species

name has priority over the other names. Whatever may be the reason, their error must be corrected, and the

correct name within the genus Bidaria is B. inodora.

maria (Endl.) Decne. in A.R de Candolle, Prodr.

595. 1838. Type species: B. tingens (Roxb.) Decne. (Asclepi „

Bidaria inodora (Lour.) Decne. in A.P. de Candolle & A.L.P.P. de Candolle, Prodr. f

Journal of the Botanical Research Institute of Texas 7(2)

The genus Bidaria is closely related to Gymnema but differs from it in having bifarous pubescent internodes

and unpaired and non-bifid umbellate cymes. Additionally, the shape of the corolla and corona are different in

both genera. Although Hooker (1883) treated Bidaria as a synonym of Gymnema R. Br., Huber (1973) reinstated

the genus Bidaria which was further supported by Jagtap and Singh (1999).

As summarized above, besides making the new combination Bidaria inodora, Decaisne (1844) also made

the new combination Gymnema tingens. It is uncertain whether Decaisne made the two new combinations de-

liberately or inadvertently. Whatever may be the fact, both new combinations were validly made and are

treated as alternative names (see Melbourne Code Art. 36.2; McNeill et al. 2012).

VLEDGMENTS

and Kanchi N. Gandhi (GH) for his kind help during preparatio

:ript and reviewing the same before comm

Endlicher, S.F.L. 1836-41. Genera plantarum secundu

Hooker, J.D. 1883-85. The flora of British India. London, UK. 4:1-780.

Huber, H. 1973. Asdepiadaceae. In: B.A. Abeywickrama, ed. A revised handbook to the flora of Ceylon. Peradeniya, Sri

urales disposita. Vindobonae.

idaceae). Botanical Survey of India, Calcutta, India,

and H. Raven, eds. Flora of China. Beijing, China and

24:1-285.

Li, B, M.G. Gilbert, and W.D. Stevens. 1 995. Asdepiadaceae. In: Z.Y. Wu and H. Raven, eds.

St. Louis, Missouri. 16:189-270.

McNeill, J, F.R. Barrie, W.R. Buck, V. Demoulin, W. Greuter, D.L Hawksworth, P.S. Herendeen, S. Knapp, K. Marhold, J. Prado, W.F.

Pruo'homme van Reine, G.F. Smith, J.H. Wiersema, and NJ. Turland (eds. & compilers). 2012. International Code of Nomen-

clature for algae, fungi, and plants (Melbourne Code) adopted by the Eighteenth International Botanical Congress

Melbourne, Australia, July 201 1 . Regnum Vegetabile 1 54.

MICROPETASOS, A NEW GENUS OF ANGIOSPERMS FROM

MID-CRETACEOUS BURMESE AMBER

George 0. Poinar, Jr. Kenton L. Chambers

Joerg Wunderlich

Department of Zoology Department of Botany and Plant Pathology

Oregon State University Oregon State University

Corvallis, Oregon 9733 1, U.S.A. Corvallis, Oregon 9733 1, U.S.A.

Oberer Hauselbergweg 24

Hirschberg 69493

GERMANY

ABSTRACT

RESUMEN

INTRODUCTION

Burmese amber has been mined since AD 100, when an amber trade route was established with China. Around

1896, it was noted that the amber contained insect remains. In 2001, a new amber mine was opened in the

Hukawng Valley, southwest of Maingkhwan in the state of Kachin (Poinar et al. 2005). Amber-bearing strata in

this mine, known as the Noije Bum 2001 Summit Site, were initially assigned to the Upper Albian (97-110 Ma)

of the Early Cretaceous on the basis of paleontological (ammonite) and palynological evidence (Cruickshank

& Ko 2003). More recently, Shi et al. (2012) give an age of 98.79+/-0.62 Ma, based on U-Pb analysis of zircons

from the volcanoclastic matrix of the amber. This is slightly younger than the date of 100.5 Ma assigned to the

end of the Albian by the International Commission on Stratigraphy (2013, http://www.stratigraphy.org). How-

ever, Shi et al. (op. cit.) contradict themselves in claiming that the amber has not been re-deposited, yet stating

been re-deposited, because amber is not formed in such an environment. Just when it was first formed in a ter-

restrial forest is not known, leaving the true age of amber from Myanmar in question. With re-deposition, the

amber must be older than the zircon-based dates determined by Shi et al. (op. cit.).

The ancient age of amber from this site is supported by the presence of primitive insects in the deposit. For

example, a bee was discovered that possessed some characters of sphecid wasps, the taxon which, in tradi-

tional systematics, is considered to be ancestral to bees (Poinar & Danforth 2006). An elcanid grasshopper was

also found, representing a group (Elcanoidea) that first appeared in the Early Permian and continued only to

e m id-Cretaceous (Poinar et al. 2005). Thus, paleontological evidence, atomic dating, and the insect inclu-

in the amber favor an early mid-Cretaceous age for mines at the Noije Bum 2001 Summit Site. In their

°°k on early flowers of the Cretaceous, Friis et al. (2011, p. 34) propose that Myanmar amber is Late Creta-

Ceous or Early Cenozoic; however, they provide no new evidence in support of this revised date.

746

Nuclear magnetic resonance (NMR) spectra of samples taken from this locality indicate an araucarian

(possibly Agathis) source of the amber (Poinar et al. 2007b). While insect fossils dominate (Grimaldi et al.

2002), the deposits have revealed some interesting plants, including 2 early bambusoid grasses (Poinar 2004),

a staminate flower with affinities to the Monimiaceae (Poinar & Chambers 2005), an epigynous flower similar

to Comaceae (Poinar et al. 2007a), a pistillate, apetalous flower with possible connections to the rosid clade of

eudicots (Poinar et al. 2008b), and an epigynous flower with characters paralleling some modem Cunoniaceae

(Chambers et al. 2010, Grimaldi et al. 2002, fig. 13).

The present fossil consists of a basally bracteate fragment of a mixed cymose-paniculate inflorescence, 9

mm long (Fig. 1), whose branches bear 3-7 flowers on glabrous, relatively stout pedicels (Fig. 3). Eighteen flow-

ers are present, but only 10 are positioned well enough for study. The perianth lacks petals, consisting of a

single whorl of five basally connate sepals spreading laterally at anthesis (Fig. 2). The calyx lobes may be equal

or unequal in size. The numerous stamens are tightly clumped around the pistil, perhaps becoming aggregated

during preservation in the resin. On four flowers, 1 or 2 unusually long stamens are visible, separate from and

external to the others (Figs. 2, 3), and some of these appear to have a broadened, ribbon-like filament (Fig. 2).

The gynoecium consists of a single carpel bearing a short, curved, attenuate style, the ovary being mostly hid-

den by the surrounding mass of stamens (Fig. 3). Among eudicot flowers thus far described from this period

(Friis et al. 2006, 2011), Micropetasos is distinctive in its combination of five connate, spreading sepals, no pet-

als, numerous stamens with bisporangiate anthers, and a gynoecium comprising a single, superior carpel with

MATERIALS AND METHODS

The fossil inflorescence is preserved in a quadrilateral piece of amber with sides of 20, 15, 11, and 10 mm,

which initially was part of a larger piece containing a fossil scorpion. It is in the amber collection of J. Wun-

derlich, Oberer Hauselbergweg 24, Hirschberg, Germany 69493, and will eventually be deposited in the

Senckenberg Museum and Research Institute, Frankfurt-am-Main. The amber was obtained from the Noije

Bum 2001 Summit Site in the Hukawng Valley, Burma, as described above. Examination and photography

were made with a Nikon stereoscopic microscope SMZ-10R at 80X and a Nikon Optiphot microscope at 800X.

Flowers small, in groups of 3-7 on branches of a basally bracteate, glabrous, mixed cymose-paniculate inflo-

rescence (Fig. 1), pedicels relatively stout, glabrous, ebracteate (Fig. 3), flowers bisexual, hypogynous, calyx

5-merous, regularly or irregularly actinomorphic, sepals glabrous, basally connate, lobes triangular, obtuse or

acute, equal or unequal (Fig. 2), petals none, stamens numerous, bunched around the pistil (Fig. 3), filaments

linear, short on outer stamens, longer on those near the pistil, sometimes 1-2 outer filaments elongated (Fig?.

2, 3), anthers small, oblong-ovoid, bisporangiate (Figs. 2, 3), connective not prolonged, gynoecium of 1 carpel,

ovary conic or ovoid (hidden by the mass of appressed stamens), disc or nectaries, if any, not visible, style short,

arcuate, attenuate, stigmatic area terminal, papillate, not enlarged.

Micropetasos burmensis G O. Poinar, K.L. Chambers & j. Wunderlich, sp. nov. (Figs. 1-4). Tvph: MYANMAR j

unknown ambe^ BUnl ^ SUmmit ^ amber mine in the Hukawn g Valley, SW of Maingkhwan, 26°20'N, 96°36'E, 2012, I

eventually to be deposited in the Senckenberg Museum and Research Institute, Frat^^am-Ma^Gmllny)^ 11011

Flowers 0.8 mm in diameter at anthpgjs, conn

filament length variable, anthers 0.06-0.10 n

0.26-0.64 mm, pollen triaperturate (Fig. 4), diameter 10-14 p

E T^- < l e T name ' rom ‘ he Greek “ micro ' sma11 - >«**.• broad-brimmed hat, from the

imagined shape of the flowers • -

:alyx 0.13 mm, lobes 0.26-0.30 mr

a long, 0.38 mm wide, style 0.1

t, pedicels

-sis- Species name from the country of origin of the fossil.

748

Insome modem families withasingle superior carpel, such as Fabaceae, a curved style may be associated with

^ r „e Sid e M ^

*- «* «*-. and to mIw 2“ be

formless mass (Fig. 3). An irregular feature of the andrn ^ ^ P ° Ucn ^ ^ flattened ““ *

^vte“;:^ hads,am “ sof “"^

An irregularity in the gynoecium is present in one flower of Micropetasos, where two styles are present,

one short, straight, and appressed to the usual longer, curved one (Fig. 1). In is unclear whether this indicates

a fully bicarpellate gynoecium. Our ability to detect these kinds of developmental variations is due to the rare

example of a fossil in which several flowers of the species are connected in an inflorescence, as contrasted to the

usual case where flowers are detached and only one or two can be assigned to the same taxon (but see Poinar

et al. 2008a and Chambers et al. 2012, where six and four flowers, respectively, were available) In an example

cited by Friis et al. (2011, p. 32), two coalified fossil flowers of Lasistemon, male and female, could be associated

through the presence, in both, of pollen having a distinctive exine pattern. From the example of Micropetasos,

one might speculate that floral development was more flexible, i.e. less canalized, in the early evolution in some

angiosperm clades, but that this has gone undetected because the descriptions of fossil taxa have commonly

been based on one or a few flowers. In deposits of coalified or lignified flowers, where large numbers of speci-

mens are collected together (Friis et al. 2011), it is possible that this drawback can be overcome.

The consistent curvature of the style, together with the relatively stout floral pedicels of Micropetasos, may

have been associated with insect pollination, especially given the small size of the flowers. The presence of

po en grams on the style and calyx but not in the surrounding amber suggests that the grains may have been

adhesive. This feature would facilitate attachment to the body of visiting insects. Small insects would be the

most likely pollinators of minute flowers such as those of Micropetasos burmensis. Melittosphex burmensis, a

tiny bee just less than 3 mm in length that lived in the Burmese amber forest (Danforth & Poinar 2011) ii a

possible candidate.

A possible relationship of Micropetasos with a modem family in one of the redefined clades of eudicots

(APGIII, Stevens 200! onwanl) is problematic Features such as hypogyny, a counate calyx, numerous sta-

mens, a single carpel, and a curved style are suggestive of certain members of the Fabaceae. However the simi-

larity is only superficial, because the inflorescence differs from the racemose type found in that family, and

numerous stamens occur principally in the highly derived subfamily Mimosoideae. Furthermore, molecular

P ylogenetic studies date the origin of Fabaceae to the Early Tertiary (Lavin et al. 2005). In terms of phyloge-

netic systematics, Micropetasos appears to represent an early member of the Pentapetalae clade (Cantino et al

2007), also known as core eudicots. We prefer to leave the question of its exact familial relationships open at

ACKNOWLEDGMENTS

We thank James Doyle for his careful review and helpful suggestions.

Cant.no, P.D., J.A. Doyle, S.W. Graham, W.S. Judd, R.G. Oimstead, D.E. Solus, P.S. Soltis, and M J. Donoghue. 2007. Towards a phy-

logenetic nomenclature of Tracheophyta. Taxon 56:822-846.

nambers, K.L., G.O. Poinar, Jr., and R. Buckley. 2010. Tropidogyne, a new genus of Early Cretaceous eudicots (Anqiosper-

CH mae ) fr °m Burmese amber. Novon 20:23-29.

HAMBERS, K.L., G.O. Poinar, Jr., and A.E. Brown. 2012. A new fossil species of Colpothrinax (Arecaceae) from Mid-Tertiary

exican amber. J. Bot. Res. Inst. Texas 6:557-560.

~ T .C Nixon. 1998. Fossil Clusiaceae from the Late Cretaceous (Turanian) of New Jersey and implications

CRWckshank, R.D. and K. Ko. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. J. Asian Earth

w. 21:441-455.

Danforth, B.D. and G.O. Poinar, Jr. 2011. Morphology, classification, and antiquity of Melittosphex burmensis (Apoidea:

F «t5EM SPheCidae ) and implications for ear| V bee evolution. J. Paleontol. 85:882-892.

s ' ■ K.R. Pedersen, and P.R. Crane. 2006. Cretaceous angiosperm flowers: innovation and evolution in plant reproduc-

PrJ 0 "' Pala eogeog. Palaeodimat. Palaeoecol. 232:251-293.

M '' pR - Crane, and J.R. Pedersen. 2011. Early flowers and angiosperm evoli

Cambridge.

. Cambridge i

Journal of the Botanical Research Institute of Texas 7(2)

Grimaldi, D.A., M. Engel, and P.C. Nascimbene. 2002. Fossiliferous Cretaceous amber from Myanmar (Burma): its rediscovery,

Lavin, M., P.S. Herendeen, and M.F. Wojciechowski. 2005. Evolutionary rates analysis of Leguminosae implicates a rapid diver-

sification of lineages during the Tertiary. Syst. Biol. 54:575-594.

Pchnar, G.O., Jr. 2004. Programing burmitis gen et sp. nov., and P. laminatus sp. nov.. Early Cretaceous grass-like monocots

in Burmese amber. Austral. Syst. Bot. 17:1-9.

Poinar, G.O., Jr., R. Buckley, and A. Brown. 2005. The secrets of Burmese amber. Mid Amer. Paleontol. Soc. 29:20-29.

Poinar, G.O. Jr. and K.L Chambers. 2005. Paleoanthella huangii gen. and sp. nov., an Early Cretaceous flower (Angiosper-

mae) in Burmese amber. Sida 21:2087-2092.

Poinar, G.O., Jr. and B.N. Danforth. 2006. A fossil bee from Early Cretaceous Burmese amber. Science 314:614.

Poinar, G.O., Jr., K.L Chambers, and R. Buckley. 2007a. Eoepigynia burmensis gen and sp. nov., an Early Cretaceous eudicot

flower (Angiospermae) in Burmese amber. J. Bot. Res. Inst. Texas 1 :91-96.

Poinar, G.O., Jr., GJ.B. Lambert, and Y. Wu. 2007b. Araucarian source of fossiliferc

anatomical evidence. J. Bot. Res. Inst. Texas 1 -.449-455.

Poinar, G.O., Jr., K.L. Chambers, and A. Brown. 2008a. Lasiambix dominicensis gen a

can amber showing affinities with Fabaceae subfamily Caesalpinioideae. J. Res. Inst. Texas 2:463-471 .

Poinar, G.O., Jr., K.L Chambers, and R. Buckley. 2008b. An Early Cretaceous angiosperm fossil of possil

rosid floral diversification. J. Bot. Res. Inst. Texas 2:1 1 83-1 1 92.

Shi, G., DA Grimaldi, G.E. Harlow, J. Wang, J. Wang, M. Yang, W. Lei, Q. Lj, and X. Li. 201 2. Age constraint c

based on U-Pb dating of zircons. Cretac. Res. 37:155-163.

Stevens, P.F. 2001 onwards. Angiosperm Phytogeny Website, Version 12, July 2012 [and more or less

dated since]. <http//www.mobot.org/MOBOT/research/APweb> [Accessed on 20 August 2013].

. PUGH’S HERBARIUM

Alfred Traverse

752

Journal of the Botanical Research Institute of Texas 7(2)

Page from the Pennsylvania Agricultural College

Inventory of January , 1868.

One Articulated Skeleton, and otic Loose Skeleton, tor Instruction in Hu-

man Anatomy and Physiology,

Diagrams to Illustrate Human Anatomy and Physiology,

Dr. Pugh’s European Herbarium,

An Air Pump, Electrical Maohine, &c. ,

A very good assortment of Apparatus and other Appliances for Class-room

and Laboratory Instructions in Chemistry,

A Surveyor’s Compass, Chains and Pins,

A Boil Hood Transit Eail Road Level and Graduated Staff,

The use of Prof. James Clark’s private Zeological Collection,

A Mason & Hamm’s Organ,

It is dear from Pugh’s correspondence that he wanted the new college in rural Pennsylvania to become an

important center for agricultural research, and he regarded a herbarium as an essential asset for the aspects of

such research that depended on a breadth of botanical knowledge and information. His herbarium is also more

personal than the house he built but never lived in, since he obviously handled each and every plant specimen

that he collected, purchased, or otherwise obtained while in Heidelberg, Germany, in the spring and summer

of 1856. Among other things, the Pugh specimens demonstrate dramatically the durability of herbarium speci-

mens. Despite the many vicissitudes of the collection in Germany, the voyage to America, and the early days at

Penn State, most of the Pugh plants remain in excellent condition (Fig. 2).

Especially noteworthy in this connection are some specimens that Pugh obtained from the herbarium of

G.W. Bischoff, Professor of Botany at Heidelberg. They survived collection in various parts of the world, ship-

ping to Germany and then shipping again to the rather primitive environment of Centre County, Pennsylva-

nia, to a nascent institution in open farmland, fifteen miles from the nearest city, with no finished building to

receive the pressed plant collection.

Evan Pugh was not only the first president of what became Penn State University, he was also the youngest

and unquestionably the most talented president the institution has ever had (Fig. 3). Had he not died at the

early age of 36, it is certain that his impact on American education and science would eventually have rivaled

that of the legendary American college presidents of the 19th century, luminaries such as Charles W. Eliot of

Harvard. In science, his biochemical work in Germany, and especially in England, on the fixing of nitrogen by

plants, was widely acclaimed. His early correspondence as president of the nascent Pennsylvania college shows

that he was having an impact in the political/educational arena of America. For example, he persuaded others

of the importance of the developments leading to the Morrill Act of 1863, which set up a funding system for

what became the “Land Grant Colleges.” This was most dramatically displayed in Pugh’s actions in assuring

that the funds accruing to education in Pennsylvania from the Act would go to Penn State and only to Penn

State.

On one of the sheets of a herbarium specimen Pugh collected in Heidelberg, he noted that the specimen

was “collected for W. H. Brewer.” Brewer and Pugh were in Heidelberg at the same time— the summer of

1856— and both did extensive botanizing that summer in central Europe. Brewer later became head of the

Agricultural Department at the Sheffield Scientific School at Yale, but at the time Pugh arrived at the nascent

of 1856 and next to it (right), a specimen of the same species (Lathyrus

754

Penn State in 1859, Brewer was Profess

Washington, Pennsylvania. Brewer was

touch back home in Pennsylvania, for i

legislation.

Pugh was a chemist, but one who:

of Chemistry at what is now Washington and Jefferson College in

1 outstanding scientist. Presumably he and Pugh remained in close

mple, on such matters as the move toward the Morrill Land Grant

interests were broad, and whos

nected to plants. His early education, through what we would consider high school, was in various private

schools in eastern Pennsylvania and New York, with heavy emphasis on languages, mathematics, and the sci-

ences. He then taught in an elementary school and subsequently became principal of an academy, both in the

Philadelphia area. By 1853, at age 25, he had inherited a considerable fortune and was able to travel to Europe

and spend the next three years at the German universities of Leipzig and Gottingen.

After completion of his doctoral dissertation, which dealt mostly with mineralogical chemistry, at

Gottingen in early 1856, he accepted a research position in the renowned laboratories at Rothamsted Experi-

mental Station, England, where he worked for several years, beginning in the Fall of 1856. He made significant

discoveries there, showing that certain plants can take nitrogen from the atmosphere and incorporate it into

their chemical structure. This is a critical matter to Earth’s biochemistry, as it explains how plants can produce

proteins independently from those they can absorb from the soil. It has been suggested that if research of such

significance were published today, the author would likely get a Nobel prize or share one with co-authors. The

critical paper was Lawes, Gilbert and Pugh, 1862. Pugh was invited to lecture on this research by the British

Chemical Society and was made a fellow of the Society in reaction to the significance of his discoveries. Later,

the importance of his early scientific work was overshadowed by his accomplishment in turning “Farmers

High School” into the forerunner of the great American Land Grant universities. Over the years, the great sig-

nificance of his work with plant fixation of nitrogen has occasionally been noted in the literature (e.g., Browne

1930).

During the Spring and Summer of 1856, before Pugh went to Rothamsted, he spent several months in

Heidelberg. Why he went there is not certain. A letter home commented that his laboratory in Heidelberg was

the best he had ever had in Germany and that he loved the countryside around this storied University. The

chief chemist there at the time was Robert W. E. Bunsen of Bunsen-bumer fame, but I have found no evidence

that Pugh worked with him. One thing is certain about Pugh’s time in Heidelberg: he did much plant collecting

and was in contact with the botanical personalities of that time and place. It seems likely that the purpose of

the sojourn in Heidelberg was botanical. Pugh would certainly need to enrich his botanical knowledge to ac-

company his mastery of chemistry, to be prepared for work at an agricultural college in the mid-19th century.

plants for this purpose were collected by Pugh himself in parts of Baden-Wurttemberg near Heidelberg. Fur-

thermore, and very important, is that one of the leading botanists in Germany of the time, G.W. Bischoff

(1797-1854), was Professor of Botany at Heidelberg University and Director of the Heidelberg Botanical Gar-

den. Bischoff died not long before Pugh had finished his doctorate at Gottingen and headed for Heidelberg.

Bischoffs herbarium was sold at auction in Heidelberg on 21 July 1856, and Pugh bought a considerable part of

it- Those plant specimens, for the most part clearly marked with both Bischoffs and Pugh’s names, plus Pugh’s

own collections of 1856 from the Heidelberg area — including a number from the famous Heidelberg Botanical

arden— plus many specimens that he somehow obtained from other German collectors, make up “Dr. Pugh’s

European Herbarium” that he shipped to America (Figs. 2, 4-7).

One significant aspect of “Dr. Pugh’s Herbarium” is that Heidelberg’s Professor Bischoff was a prodigious

collector of plants, having made collecting forays from about 1820 to about 1850 to various parts of the world —

South Africa, South America, etc. On at least two of his trips he collected in or near Pennsylvania. There are a

number of sheets in the Pugh herbarium with labels indicating that the specimens were collected in the vicin-

,ty Reading, Pennsylvania, in 1831 and in New Jersey in 1849 (Figs. 5, 6 & 7). (Coincidentally, Evan Pugh

was born and raised in Oxford, a small rural town in southeastern Pennsylvania, less than 50 miles south of

W in 8 an d the same distance west of New Jersey.) The American Bischoff specimens would have crossed the

Atlantic with Bischoff to Europe in a sailing vessel, and then after some years in the botanical collections in

etdelberg, been bought by Pugh in 1856, and shipped back across the Adantic, this time probably by steam-

P, eventually ending up in the Penn State Herbarium,

n one of his letters, Pugh mentioned shipping a crate of rocks and minerals from Gottingen to relatives in

i area, presumably samples from his Ph.D. work. It seems probable that he arranged the same

^Philadelphia

756

Journal of the Botanical Research Institute of Texas 7(2)

Traverse, Evan Pugh's herbarium

<p , ftuf/,

SS-~ :

UcsnrttieA+ijfa J+4.

S3?'

p. p.

758

Journal of the Botanical Research Institute of Texas 7(2)

sort of shipment to Philadelphia of what would have been at least two or three large steamer trunks contaii

the infant Penn State herbarium. There, relatives would have been available to help him with further mo

chores. Such modern-sounding shipment was apparently quite ordinary in 1856. The journey of the cratt

trunks full of plant specimens from Heidelberg, probably to Rotterdam, then by sail or steamer to Philadelp

end eventually by train to Bellefonte, Pennsylvania, and by some sort of horse-drawn vehicle the last ten n

or so to State College, before there even was a State College or a significant road to the area, is fascinatin

EVAN PUGH AT THE PENNSYLVANIA AGRICULTURAL COLLEGE

Dr. Pugh arrived in Centre County in 1859 to head up the Farmers High School, which he soon renamed Agri-

cultural College of Pennsylvania. (It became Pennsylvania State College in 1874 and Pennsylvania State Uni-

versity in 1953.) Pugh’s first job was to rescue the then floundering agricultural college, beginning with secur-

ing funding for completion of the first, and for some time only, permanent building, the original Old Main. The

herbarium would have been kept in that building, which also provided classrooms, laboratories, dormitories,

refectory, and so on. It was constructed of Ordovician dolomite, quarried near the building. Students provided

much of the labor as part of their program. The ground plans for the structure show several small rooms la-

beled “Museum.” Presumably, the herbarium would have been kept in one of those rooms.

Evan Pugh died in 1864, some weeks after he had a disastrous buggy accident in the Bellefonte area, in

which one of his arms sustained multiple fractures, which were mismanaged locally. He then went to Philadel-

phia, where the bones were reset. Not long afterwards, back home, he died of a fever, frequently referred to as

typhoid; it seems more likely that sepsis from the mangled arm was responsible.

THE HERBARIUM SINCE PUGH’S DEATH

Soon after Pugh’s tragic death, there was a student in the college named William A. Buckhout, who had a keen

interest in plants and the science of botany. In 1869-1870 he studied at Harvard with Asa Gray, at that time the

most renowned American botanist. In the herbarium there isstill asmall collection of ancient photos of the great

man, certainly keepsakes that Buckhout brought back to Penn State from Cambridge, when he returned in 1871

to become head of the infant Department of Botany and Horticulture. He remained in charge of plant research

and teaching at Penn State until his death in 1911 (Fig. 8). Buckhout did not collect many plant specimens for

the herbarium, which was a unit of his department. However, one of the collections he did make was of a maple

(Fig. 9). That specimen has been important in tracing one aspect of the history of “Dr. Pugh’s Herbarium.”

For at least sixty years, everyone connected with the Penn State Herbarium has assumed that the hand-

written “Evan Pugh” on the label of every sheet of Pugh’s herbarium is in fact President Pugh’s signature. How-

ever, a trained herbarium curator from Guatemala, Ana Lu MacVean, at present a volunteer in the Penn State

Herbarium, has begun compiling a list of all Pugh/Bischoff specimens. She inadvertently happened on the

abovementioned sheet of Acer rubrum (red maple), collected by Dr. Buckhout in 1875, thirteen years after Dr.

Pugh’s death. The style of label, and the handwriting are identical to that on all of the labels in the Pugh her-

barium. More research in the Penn State Archives’ examples of Buckhout’s and Pugh’s handwriting confirms

that the labels in “Dr. Pugh’s Herbarium” are, in fact, the work of Buckhout. Apparently, about 1875, he decided

to improve the condition of the basic Penn State Herbarium by remounting, or in some cases only re-labeling,

the specimens from Dr. Pugh. In some instances, especially for the plants that Pugh had obtained from Bischoff

and other German botanists, Buckhout cut the labels off the original sheets and glued them on new sheets

along with the specimens; he also added a new label with the plant name and the inevitable “Evan Pugh, Hei-

delberg” (Fig. 6). This procedure seems to be true also for most of Pugh’s own collections, although some of the

specimens that he collected while in Heidelberg appear to be on the original paper, with a word or two in his

own handwriting, but a label prepared by Buckhout. It is interesting that Pugh put his name on only one or two

of the books from his library that Penn State still has. The lack of original signatures there and also on his bo-

tanical specimens possibly reflects his Quaker-influenced disapproval of emphasizing one’s person.

Traverse, Evan Pugh's herbarium

762

Journal of the Botanical Research Institute of Texas 7(2)

Traverse, Evan Pugh's herbarium

763

Still, it is curious and not really “correct” that Dr. Buckhout failed to indicate on the sheets or apparently

anywhere else, that he had so drastically altered the labeling of the specimens. He could have noted on the new

labels “information transcribed by WAB /date” or some words to that effect. It must be emphasized in his de-

fense, however, that his work greatly improved the physical condition of the collection. He used high-quality

paper, clearly acid-free, probably pure cellulose stock. In many instances one can tell from the transferred la-

bels that the original sheets were prepared using fair to poor paper. Buckhout also must be credited with rec-

ognizing the importance of the collection and assuring its permanence at Penn State. His emphasis on the

contribution of Pugh by putting the first president’s name on every sheet of “Dr. Pugh’s European Herbarium’’

was appropriate and scientifically correct. It should also be noted that at the time of Buckhout’s work, few col-

lectors provided much information on their labels.

The existence of Pugh’s herbarium, especially the Bischoff specimens, is of considerable interest to Ger-

man botanists, because many of Germany’s herbaria were destroyed or damaged during World War II. One

Pugh herbarium specimen has already accidentally turned out to be some sort of type specimen, because all

other specimens in Germany of that plant taxon have been lost. According to Professor Doctor Ulrich Kull,

distinguished retired botanist from Stuttgart University, many German botanists have recently expressed as-

tonishment that so much of BischofPs material ended up at Penn State. That was entirely new information to

It is somewhat surprising that the Penn State Archives have practically nothing about the history of “Dr.

Pugh’s Herbarium,” except for its mention in the inventory of the College’s meager possessions in 1868. One

can deduce that in Dr. Pugh’s time and at the beginning of Dr. Buckhout’s work, it will have been housed in the

onginal Old Main. When Botany moved to the newly constructed Botany Building (now “Old Botany”) in 1887,

Buckhout would have moved the herbarium there, but I have found no specific mention of that event. It is a

matter of living memory, that in the 20th century, when Buckhout Building was constructed to house the Biol-

ogy Department, the herbarium moved there, where it occupied several different locations. In the late 20th

century, it moved to various places in the Frear Building and then to its present location in Whitmore Lab.

Much of this moving can be attributed to the fact that, for some decades in the 20th century, natural his-

tory collections, including herbaria, were out of favor at universities. They were expensive to maintain and re-

garded as not for the most part connected to modem developments in science. However, with the realization

that herbaria provide banks of DNA, the tide turned in their favor. For example, voucher sheets of pollen- and

spore-bearing plants enable palynologists to check the identity of the plants from which slides of the pollen

were made, and to study the DNA of the same plant.

An example of such interplay between arboreta — where living, carefully documented plants grow — and

erbaria— where expertly dried and pressed, also carefully documented, plants are stored— is a living docu-

mented red maple tree (#1031) planted on the Mont Alto campus, when it was the location of the Penn State

Forestry School. We have in PAC a specimen of #1031, Acer rubrum, in flower, containing abundant pollen,

collected in 1948. It would be interesting to collect from tree #1031 again, in flower, to compare the pollen being

made now with that made 65 years ago. Have the years of exposure to atmospheric pollution and/or cosmic

radiation caused mutations in the tree’s DNA that are reflected in the pollen size and/or fine features of the pol-

Cn Wad mor phology? A scientist in the Biology Department at Penn State has shown the potential richness of

our herbarium records by germinating a seed from one of the Pugh plants, about 160 years old, and studying

e DNA of the resulting seedling. Surely, Pugh would be gratified to know that Penn State is now developing a

botanical garden/arboretum. Space will be created and designated to house the herbarium in one of the new

ec ^ addition to facilitating DNA experiments, the herbarium also provides a convenient venue for paleo-

18nn §1StS t0 StUdy Plant distribution for tar 8 et areas at times m the past. Our specimens range in age from near

oth *i 2 ° 13, and each s P ecimen contains in its dried plant tissues, samples of the carbon and oxygen and

* emental ls °topes from each of the represented dates and locations, a potentially rich source of informa-

n. Most specimens of herbaceous plants are collected with roots and therefore accidentally with some soil.

764

Journal of the Botanical Research Institute of Texas 7(2)

which soil can be analyzed in many different ways to get information about the woods of, say, Gettysburg,

Adams County, Pennsylvania, in the 1800s.

PAC is a function of the Penn State Biology Department, but has no budget or other funding specifically

for its work. Somehow it has survived despite this situation, because of volunteers who have recognized the

value of the collection, and help from persons in the university’s administration who have recognized both the

historic and scientific importance of the collection. As mentioned earlier, an expert, professional volunteer,

Ana Lu MacVean, is currently preparing an annotated list of all the sheets of the original, approximately 2,000

specimens of “Dr. Pugh’s Herbarium,” not an easy task, since they are interspersed with many thousands of

other specimens. The heart of PAC is clearly “Dr. Pugh’s Herbarium,” and that is turning out to have not only

importance as a relic of the work of Penn State’s scientifically important first president, but also potential bio-

logical significance. It incorporates much of the important early 19th century herbarium of Professor Bischoff

of Heidelberg University, purchased at auction by Pugh in July, 1856. Many other important collections, such

as the Harshberger Herbarium of Trees and Shrubs, which came to PAC from the former Penn State School of

Forestry at Mont Alto, have also, especially in recent years, found their way into the Penn State Herbarium.

It would be heartwarming if somehow PAC could be established as “The Dr. Evan Pugh Memorial Her-

barium.” There is nothing else at Penn State University that has so intimate a connection with the founding of

the institution as does this remarkable scientific collection.

ACKNOWLEDGMENTS

Professor Dr. Ulrich Kull, University of Stuttgart, Germany, assisted the author by getting much information at

the sources, about Evan Pugh’s years in Europe. Paul Karwacki and Jacqueline Esposito, of the Archives Divi-

sion of Penn State’s library system were of great help in assisting me in use of the archival material on Dr. Pugh

in the library. Ana Lu MacVean, who is preparing an annotated list of all specimens in PAC that came to Penn-

sylvania with Pugh in 1859, was very helpful in passing along information she encountered that was critical to

my study. I appreciate the careful reviews of Robert W. Kiger (CM) and an anonymous reviewer.

REFERENCES

Browne, C.A. 1930. European laboratory experiences of an early American agricultural chemist— Dr. Evan Pugh (1828-

1864). J. Chem. Edu. 7:499-51 7.

Lawes, J.B., Gilbert, J.H. and Pugh, E. 1862. On the sources of the nitrogen of vegetation; with special reference to the

question whether plants assimilate free or uncombined nitrogen. Phil. Trans. London, Part II for 1861. Pp. 433-577.

Thiers, B. 2008. Index herbariorum. A global directory of public herbaria and associated staff. New York Botanical Gar-

den's Virtual Herbarium, http://sweetgum.nybg.org/ih/.

ALPINE FLORA OF CERRO QUIEXOBRA, OAXACA, MEXICO

J. Andrew McDonald

Department of Biology

The University of Texas - Pan American

1201 W. University Dr.

Edinburg, Texas 78739, U.S.A.

RESUMEN

INTRODUCTION

%ne and subalpine vegetation is often difficult to delimit on summits or along ridges of mountains that ap-

proach timberline thresholds. Geophysical discontinuities in the angle and orientation of slopes, varying de-

^ ees °^ ex P°sure to wind and solar irradiation, sporadic fires and local edaphic heterogeneity create and affect

Q ontinu ities in the distribution of timberline species and their plant communities (Billings 1974; McDonald

o , UC ^ is l ^ e case f° r alpine plant communities on the poorly explored summit of Cerro Queixobra

6 1234 N; 96°11'51"W) in the Sierra Madre del Sur of southern Oaxaca (Fig. 1), where a species-rich associa-

te® of shade-intolerant species occurs exclusively on the mountain’s summit (ca. 3700 m).

^ rst comprehensive collections of Cerro Quiexobra were accomplished by the author in December of

^ A - McDonald 2900-2951 ) and October of 1990 (J.A. McDonald 2987-3040), followed by significant expe-

! Uons by Jaime Hinton and collaborators during October of 1995 (J. Hinton et al. 26133-26217) and August of

6 (J- Hinton et al. 26641-26843). These collectors encountered a rich herbaceous flora in moist glades of

fountain saddles (Alepidocline macdonaldana, Erigeron quiexobrensis, Hieracium abscissum, Luzula denticulata,

toa oenanthoides, Schiedeella hyemalis) and an array of herbs and short-statured (sl.O m), lignescent phanero-

£ytes> on exposed rock and cliff faces ( Ageratina pichinchensis, Aphanactis macdonaldii, Castilleja nivibractea,

onolaena macdonaldii, Verbesina macdonaldii, Valeriana pulchella). Several miniature, hemi-cryptophytes

chamaephytes ( Calandrinia micrantha, Lobelia macdonaldii, Oxalis hintoniorum, Sisyrinchium tenuifolium.

Journal of the Botanical Research Institute of Texas 7(2)

766

Tauschiafiliformis ) are loa

vith creeping, r

and a geophytic Peperomia (P. monticola). Most of the aforemen-

tioned genera occur frequently in disjunct alpine zones throughout Mexico. The timberline vegetation of Cerro

Quiexobra also harbors unique and highly specialized shrubs and short trees, such as the only known arbores-

cent bluebonnet, Lupinus jaimehintoniana, which occupies a shrub-dominated, subalpine ecotone between

surrounding pinelands (P. aff. hartwegii) and treeless zones.

The alpine flora of Cerro Quiexobra is extraordinarily unique in various regards, owing ostensibly to its

geographic isolation during the present age and glacial maxima of the Pleistocene (McDonald 1993). Compar-

ing closely in species richness to other alpine refugia of similar size in northeast and northwest Mexico (Mc-

Donald 1990, 2011), this highly restricted plant community comprises at least 80 species of herbs and short

shrubs. The vegetation is particularly noteworthy in terms of rare and endemic elements, supporting at least 19

species that are presendy known exclusively from this high point of the Sierra Madre del Sur, more than double

the number of single-peak endemics encountered in other alpine refugia of Mexico (Almeida-Lenero et al.

2007; Beaman 1962, 1966; Laur & Klaus 1975; McDonald 1993, 2011; Narave 1985). Unlike the closest known

alpine grasslands to the north and east, Pico de Orizaba of Veracruz and Volcan Tacana of Chiapas (respec-

tively), as well as alpine grasslands across central Mexico’s volcanic belt (19° Lat. N; Almeida-Lenero et al. 1994,

2004, 2007; Gimenez de Azcarate & Escamilla 1995; Rzedowski 1975), an array of forbs and perennial herbs of

Cerro Quiexobra, mosdy Asteraceae, are dominant over few grasses (5 spp.), caryophylls (2 spp.) and mustards

(2 spp.). Asteraceae are especially well represented in terms of species richness (27spp.) and plant cover, and

account for a substantial number of narrow endemics (11 spp.). The Apiaceae and Orobanchaceae rank second

767

in terms of species richness (4 spp. each), followed by the Ericaceae, Plantaginaceae, and Solanaceae, each with

three species. As presently known, only two non-native species occur among the natives, Poa annua and Sisym-

brium cf. irio, underscoring again the persistent insularity of this plant community.

The remarkable rates of narrow endemism at the summit of Cerro Quiexobra are apparently the result of

long-term isolation during both glacial and interglacial periods of the Pleistocene, as well as the distinctive

geological substrates of the Sierra Madre del Sur. The closest, aforementioned alpine zones to the north and east

of Oaxaca occupy loose basaltic soils and gravel, while Cerro Quiexobra’s flora inhabits solid, rocky substrates

of a mountain range that is formed by a complex mixture of Lower Cretaceous marine sediments, Proterozoic

metamorphic rock and extrusive Tertiary sedimentary volcanics (Ortega-Gutierrez 1992). With respect to evi-

dence that timberline plant communities of Mexico descend around 1000 m during glacial maxima, the alpine

vegetation of Cerro Quiexobra presumably expands in distribution during pluvials to cover around 170 km of

the east-west backbone of the Sierra Madre del Sur (McDonald 1993). The geographical limits of this relatively

broad distribution are still far-removed, however, from insular alpine zones to the north (Pico de Orizaba, Ve-

racruz) by at least 200 km and to the east (Volcan Tacana, Chiapas) by around 400 km (Fig. 1).

This epicenter of rare and potentially endangered species warrants serious attention in terms of research

and protection. While timber extractions and the establishment of logging roads during the last two decades

have altered surrounding pine woodlands considerably, as have widespread, recurrent and devastating fires

(fidej. Hinton; Turner 1995), the short- and long-term impacts of these disturbances are essentially unknown.

in recent years on account of over-grazing, fires, trail erosion, and human recreation (Gimenez de Azcarate &

Escamilla 1999; Almeida-Lenero et al. 2007), successful efforts to protect this biologically diverse refugium

would conserve valuable evidence to better elucidate Mexico’s deep and colorful, natural history.

Journal of the Botanical Research Institute of Texas 7(2)

Arbutus xalapensis Kunth {JAM 2939)

Arctostaphylospungens Kunth. {JAM 2941)

C. {JAM 2922; JH 26149)

Castilleja integrifolia Lf. (JAM 2937)

* Castilleja nivibractea G.L. Nesom ( JAM 2948, 3002)

*Castilleja quiexobrensis G.L. Nesom {JAM 2928)

esom IJH 26133, 26451,

in prep {JAM 301 7; JH 26799)

(JAM 2929)

Penstemon gentianoides (Kunth) Poir. (JAM 2925; JH 261 34, 26801)

Penstemon kunthii G. Don (JAM 2929, 3025; JH 26299)

903,3030)

pulchella M. Martens & Galeotti (JAM 3022)

ACKNOWLEDGMENTS

►r, Beryl Simpson, for years of assistance on this and re-

ess to the herbarium’s various collection databases and

ons. Lindsay Woodruff tracked down missing materials

Turner and Guy Nesom worked for years on specimen

I was adrift in other botanical endeavors. I thank Rich-

The author expresses gratitude to TEX and its Din

lated studies. Tom Wendt provided indispensable

assisted on numerous occasions in species determii

and facilitated my perennial visits to Austin. Billie

determinations and the publication of new taxa while I w

ard S. Felger and Remedios Aguilar Antelises for helpful reviews on an earlier version of the manuscript.

REFERENCES

Almeida, L, A.M. Cleef, A. Herrera, A. Velazquez, and I. Luna. 1994. El zacatonal alpino del Volcan Popocatepetl, Mexico, y so

posicion en las montanas tropicales de America. Phytoceonologia 22:391-436.

Almeida-Lenero, L., Gimenez de Azcarate, A.M. Cleef, and A. Gonzalez Trapaga. 2004. Las comunidades vegetales del zacatonal

alpino de los volcanes Popocatepetl y Nevado de Toluca, regidn central de Mexico. Phytoceonologia 34:91-132.

Almeida-Lenero, L., M. Escamilla, J. Gemenez de Azcarate, A Gonzalez Trapaga, and A.M. Cleef. 2007. Vegetacidn alpina de los

volcanes Popocatepetl, Iztaccihuatl y Nevado de Toluca. In: I. Luna, JJ. Morrone y D. Espinosa, eds. Biodiversidad de

la Faja Volcdnica Transmexicana. Universidad Nacional Autdnoma de Mexico, Mexico, D.F. Pp. 179-198.

Beaman, J. 1 962. The timberlines of Iztaccihuatl and Popocatepetl, Mexico. Ecology 42:377-385.

Beaman, J. and J.W. Andresen. 1 966. The vegetation, floristics and phytogeography of the summit of Cerro Potosf, Mexico.

769

Billings, W.D. 1 974. Adaptations and origins of alpine plants. Arctic Alpine Res. 6:1 29-1 42.

Gimenez de Azcarate, J. and M. Escamilla. 1 999. Las comunidades edafoxerifilas (enebrales y zacatonales) en las montanas

del centra de Mexico. Phytocoenologia 29(4):449-468.

Lauer, W. and D. Klaus 1 975. Geoecological investigations on the timberline of Pico de Orizaba, Mexico. Arctic Alpine Res.

7:315-330.

McDonald, J.A. 1990. The alpine-subalpine flora of northeastern Mexico. Sida 14:21-28.

McDonald, J.A. 1993. Phytogeography and history of the alpine-subalpine flora of northeastern Mexico. ln:T.P. Rama-

moorthy, J. Fa, R. Bey, and A. Lot, eds. Biological diversity of Mexico: origin and distribution. Oxford University Press,

New York, Pp. 681-703.

McDonald J.A., J. Martinez, and G.L. Nesom. 2011. Alpine flora of Cerro Mohinora, Chihuahua, Mexico. J. Bot. Res. Inst. Texas

5(2):701-705.

Narave, H. 1 985. La vegetacion del Cofre de Perote, Veracruz, Mexico. Biotica 1 :35-57.

Oktega-Gutierrez, F. 1992. Carta Geol6gica de la Republica Mexicana: escala 1:2,000,000. Institute de Geologia, UNAM.

Rzeoowski, J. 1975. An ecological and phytogeographical analysis of the grasslands of Mexico. Taxon 24:67-80.

Turner, B.L. 1 992. 1 995. A new species of Lupinus (Fabaceae) from Oaxaca, Mexico: A shrub or tree mostly three to eight

meters high. Phytologia 79:102-107.

770

BOOK REVIEW

Jane Goodall with Gail Hudson and forward by Michael Pollan. 2013. Seeds of Hope: Wisdom and Wonder

from the World of Plants. (ISBN-13: 9781455513222, hbk). Grand Central Publishing, Hachette Book

Group, 237 Park Avenue, New York, New York 10017, U.S.A. (Orders: www.hachettebookgroup.com/

publishers/grand-central-publishing). $22.36, 384 pp., 36 color photos, many b&w photos, 6" x 9".

She offers wonderful cases of children— the future of environmental protection— involved in educational ef-

forts from urban Dallas to the tip of India. She talks about grassroots movements where people rise up to pro-

tect their local resources, such as the Chipko movement in India where a group of women joined together to

protect their local forests. She highlights “forest warriors” like John Seed of Australia who took on the timber

industry, first in Australia and then across the globe; like Wangari Maathai, the founder of the Green Belt

Movement in Kenya, who fought to halt deforestation in her native country and led the planting of “hundreds

of thousands of trees around urban areas in many parts of Africa;” like Chief Almir of the Surui tribe in Brazil

who stood up to illegal loggers. These “warriors” inspire and challenge us to fight for the protection of our re-

sources. Goodall, a warrior in her own right, challenges us to never give up hope. She concludes the book with

a discussion on the resilience of our natural systems, providing us with several cases of plants and systems

that, even under the harshest conditions of deprivation, fight back with a strong will to survive. “Indeed,” says

Goodall, “nature is resilient. Therein lies our hope.”

Since publication, Goodall has been accused of plagiarizing parts of the book. Having recently read the

book, I can see some of the gray areas from which these accusations may come. However I question the fairness

of these accusations. Here is a woman whose lengthy career has spanned decades and continents, who has

been at the forefront of animal behavior research and conservation, and who has interacted with a multitude of

professionals and non-professionals on a whole host of environmental subjects. In Seeds of Hope, Goodall

weaves together a lifetime of conversations with groups and individuals. I imagine Goodall has a huge knowl-

edgebase formed from prior conversations and collaborations, so much so, that it would be hard to unravel

exact conversations in order to attribute a particular phrase or thought to a particular person when those

thoughts have intermingled with one’s own over decades of learning. Throughout the book she attributes ideas

and research to their appropriate authors, and in fact, includes 19 pages of what she terms “Gratitude” where

she thanks and gives credit to all those whose paths she has crossed in the writing of this book.

Seeds of Hope is a ramble through her long career, and I see no need in dragging an outstanding scientist

through the coals for something that has likely just become a part of who she is. At nearly 80 years old, Goodall

has been present for so many discussions and has taken environmental and conservation ideas to the world

arena that have benefitted humankind in ways we can only begin to see.

As Michael Pollan so eloquently puts it in his forward, “Seeds of Hope is not just a love letter to the plant

world, though it is certainly that. It’s also a call to arms, sounding the alarm about habitat destruction, the vio-

lence of industrial agriculture, and the risks of genetic engineering ... Goodall wants nothing less than to ex-

pand the circle of human affection once again, making it wide enough to take in the sunlight eaters.” Goodall

does so with humility and grace in this reflective and thoughtful book.— Gwen Michele Thomas, Chapter Coor-

dinator for the Society for Ecological Restoration’s Texas Chapter, Texas Master Naturalist, and Botanical Research

Institute of Texas Volunteer.

FLORA AND PHYTOGEOGRAPHY OF

CUMBRES DE MONTERREY NATIONAL PARK, NUEVO LEON, MEXICO

Eduardo Estrada-Castillon

Facultad de Ciencias Forestales

Universidad Autonoma de Nuevo Ledn

A.P. 41, 67700, Linares, N.L., MEXICO

aeduardoestradac@prodigy.net.mx

Marfa Magdalena Salinas-Rodrfguez

Facultad de Ciencias Forestales

Universidad Autdnoma de Nuevo Ledn

A.P. 41, 67700, Linares, N.L., MEXICO

madreselva_84@hotamil.com

Javier Jimenez-Perez

Facultad de Ciencias Forestales

Universidad Autdnoma de Nuevo Ledn

A.P.41, 67700, Linares, N.L, MEXICO

Jose Angel Villarreal-Quintanilla

Saltillo, Coahuila, MEXICO

javillarreal00@yahoo.com

Humberto Rodrfguez-Gonzalez

Universidad Autdnoma de Nuevo Ledn

A.P41, 67700, Linares, N.L., MEXICO

gonhumberto@gmail.com

Mario Alberto Garcfa-Aranda

Facultad de Ciencias Forestales

Universidad Autdnoma de Nuevo Ledn

A.P. 4 1, 67700, Linares, N.L., MEXICO

ABSTRACT

*7(2): 771 -801.2

tnii'

Journal of the Botanical Research Institute of Texas 7(2)

INTRODUCTION

The northeastern region of Mexico is characterized by climatic and landscape heterogeneity; its extensive

plains, high mountains, and scattered hills shelter an intricate and diverse mosaic of vegetation, characterized

by rich plant diversity and life forms. The heterogeneous physiography among different regions clearly differ-

entiates distinctive climatic zones, especially evident in the State of Nuevo Leon, where three physiographic

provinces are recognized: Gran Llanura de Norteamerica (North American High Plains), Llanura Costera del

Golfo Norte (North Coastal Gulf Plain), and Sierra Madre Oriental (INEG1 1986). These have contrasting par-

ticularities of soils, vegetation types, and plant diversity. The orthographic, edaphic, and climatic factors of the

physiographic provinces show'close relationships between the vegetation types, flora, and plant endemism

(Epling 1939; Woodson 1954; Bameby 1964; Johnston 1971, 1975; Powell & Turner 1974; Powell 1978; Zanoni

& Adams 1979; Nesom 1981; Turner 1994a, 1994b, 1994c, 1996, 1997, 2001; Valdes & Flores 1983, 1986; An-

derson 1987; McDonald 1990; Hinton & Hinton 1995; Allred & Valdes-Reyna 1997; Valdes-Reyna 1997; Es-

pejo & L6pez 1997; Henrickson & Johnston 1997; Valiente-Banuet et al. 1998; Estrada 1998; Valdes-Reyna &

Allred 2003; Mickel & Smith 2004; Estrada et al. 2007; Balleza & Villasenor 2011; Velazco-Maclas et al. 2011;

Estrada etal. 2012).

The vegetation is characterized by 11 plant communities: 1) Tamaulipan thornscrub, 2) piedmont scrub,

3) rosetophyllous scrublands, 4) microphyllous scrubland (Muller 1939; Rojas-Mendoza 1965; Rzedowski

1978; Estrada & Martinez 2003), 5) chaparral (Valiente-Banuet et al. 1998), 6) oak forest, 7) oak-pine forest

(Rzedowski 1978; Perry 1991), 8) conifer forest (Miranda & Hernandez 1963; Beaman & Andersen 1966; Rze-

dowski 1978; Perry 1991; Farjon et al. 1997; Graham 1999), 9) halophytic communities (Scott et al. 2004; Estra-

da et al. 2010), 10) alpine meadow (Beaman & Andersen 1966), and 11) aquatic vegetation (Rzedowski 1978).

The scrublands and forest types are present in the central part of Nuevo Leon and constitute the main

landscapes of the Cumbres de Monterrey National Park (CMNP), the largest National Park in Mexico, covering

an area of 177,367 ha (1773 km 2 ). The CMNP was established in 1939 by a presidential decree to preserve the

regional flora and fauna and is one of the most visited areas in Nuevo Leon for activities such as camping, hik-

ing, rappelling, and leisure. Small villages, ranches, private properties, and ejidos are widely distributed

throughout CMNP; the vibrant economy is based mainly on fruits such as apples, peaches, plums, and apri-

cots. The objective of the present work is to document the plant diversity and the phytogeography of the

CMNP.

Study area

The Cumbres de Monterrey National Park is located in the central-west portion of Nuevo Leon. It includes part

of seven municipalities: Ailende, Montemorelos, Monterrey, Rayones, Santa Catarina, San Pedro Garza Garcia,

and Santiago (25°4r-25°02'N, 100°45'-99°irw). The altitude ranges from 600 to 3400 m. The main urban

areas in CMNP are Puerto Genovevo, El Manzano, Cienega de Gonzalez, Laguna de Sanchez, El Tejocote, El

Hondable, La Camotera, La Trinidad, Potrero Redondo, El Pajonal, El Huajuco, La Huasteca, and San Antonio

de la Osamenta (Fig. 1).

Geology.— The most common outcrops are from sedimentary rock and clastic deposits of the Mezozoic

Era. Most outcrops in lowlands are constituted by lutites, conglomerates, and limestone. Higher in the Sierra

Madre Oriental are Mesozoic limestone and recent deposits of conglomerates and alluvial soils from the Qua-

ternary (INEG1 1986).

Soil- Predominant soils in lowlands are regosols and lithosols, developing into rendzinas and xerosols.

Intermountain valleys, as well as low plains, are mainly clay vertisols, frequently with high calcium carbonate

contents and poor drainage (INEGI 1986).

Climate .— The study areas have a seasonal climatic pattern. There is a dry season from November to May

and a humid season from June to October; however, some differences are evident for both the North Coastal

Gulf Plain (NCGP) and the Sierra Madre Oriental (SMO) physiographic provinces (Garda 1973; INEGI 1986).

Two main climates are dominant in the NCGP, warm and arid. The type A (C)w (semiwarm-subhumid) cli-

Estrada-C. et al.. Flora of Cumbres de Monterrey National Park, Mexico 773

mate, with 18°C as yearly mean temperature and annual rainfall averaging 1000 to 1200 mm, is distributed on

the windward slopes of the CMNP. The arid-warm (BS) and arid-semiwarm(Bw) climate types are found on the

leeward slopes south and west of the study area, averaging an annual mean temperature higher than 18°C and

annual mean rainfall 400 mm (Garcia 1973; INEGI 1986). Both types of climate are found at lower altitudes

(400-750 m) where scrublands with different physiognomy, structure, and plant composition are the main

plant vegetation. The temperate-subhumid (Cw) climate is common in the mesic and temperate areas above

750 m on windward slopes, averaging 12-18°C mean annual temperature and annual rainfall of 1500 mm.

This climate is found in the mountains and slopes over 750 m, where the main plant communities are oak and

Vegetation.— The main vegetation types in the CMNP, according to Muller (1939), Rojas-Mendoza (1965),

INEGI (1986), and Estrada et al. (2012a), are: Tamaulipan thomscrub, piedmont scrub, microphyUous scru-

bland, rosetophyllous scrubland, oak forest, oak-pine forest, pine forest, and Juniperus forest. The Tamaulipan

thomscrub is a plant community dominated by medium (1-2 m) to short (1 m) shrubs, present at lowest alti-

tudes in the CMNP.

Hie most common elements are: Agave fceheguilla, Bemardia myricifolia, Cemdittm macrum, Cordia bois-

»eri, Eysenhardtia texema. Hamrdia pattens .Jatropha dioica. Kanvmskia humMdtiana, LeucophyUumfrutescens,

Opuntia engelmannii, Schaefferia cuneifolia, and Acacia rigidula.

The piedmont scrub is composed of thorny and non-thorny evergreen or deciduous shrubs, reaching up

to 3 m tall, along the lower slopes of the mountains. The most common species are: Agave lecheguiUaAmyris

madrensis, A. texana, Bemardia myricifolia, Caesalpinia mexicana, Calia secundiflora, Celtis pallida, Cordia bois-

sieri, Eysenhardtia texana Forestiera angustifolia, Gochnatia hypoleuca, Hamrdia pallens, Helietta parx ifolia, Jat-

ropha dioica Karwinskia humboldtiana, Leucophyllum frutescens, Malpighia glabra, Neopringlea integrifolia,

Opuntia engelmannii Randia rhagocarpa, Schaefferia cuneifolia. Acacia berlandieri. Acacia greggh, Sideroxylon

774

Journal of the Botanical Research Institute of Texas 7(2)

lanuginosum. Acacia rigidula, and Zanthoxylumfagara. Higher in elevation, piedmont scrub is dominated by

Acaciella angustissima, Caesalpittia mexicana, Calia secundiflora, Juglans spp., Cercocarpus fothergilloides var.

mojadensis, Chiococca pachyphylla, Decatropis bicolor, Fraxinus greggii, Havardia pollens, Acacia coulteri, Neo-

pringlea integrifolia, Prunus serotina, Acacia roemeriana, Ptelea trifoliata, Randia rhagocarpa, Acacia berlandieri,

and Ungnadia speciosa.

The rosetophyllous scrubland is dominated by species with basal or apical leaves, arranged in a rosette,

and either caulescent or acaulescent. Most of these species are located in steep slopes and stony soils, areas with

low humidity and distributed mainly on the south and west faces of the CMNP. The most common species are

Agave americana, A. bracteosa, A. xictoriae-reginae, A. lecheguilla, A. lophantha, A. striata, Hechtia glomerata,

Hesperaloefunifera, Nolina spp., and Yucca spp.

The oak and oak-pine forest are quantitative and physiognomically the most important plant communi-

ties in the Sierra Madre Oriental, distributed from the 750 m to the summit of the higher peaks. The most

common species are Pinus pseudostrobus, P. greggii, P. strobiformis, P. teocote, Quercus canbyi, Q. coccolobifolia, Q.

fusiformis, Q. glaucoides, Q. hypoleucoides, Q. laceyi, Q. mexicana, and Q. polymorpha. The pine forest communi-

ties are located above 1600 m; Pirns cembroides and P. remota dominate on arid and almost flat slopes in low-

lands; while P. teocote, P. pseudostrobus, P. h artwegii, P greggii, and P strobiformis are distributed on moderate or

steep wet slopes. The juniper forest is distributed in temperate areas with low rainfall and is dominated mainly

by J. deppeana (Zanoni & Adams 1979).

The chaparral consists of shrubby species with coriaceous leaves and dense canopy cover, inherin g the

semiarid areas on mountains. Among the main species are several oaks of short stature (Valiente-Banuet et al.

1998) as well as Arctostaphylos pungens, Ceanothus fendleri, C. greggii, Garrya ovata, Lindleya mespilioides,

Malacomeles denticulata, and Purshia plicata.

METHODS

Field and lab work — Field work was carried out from 2005 to 2012, collecting plants in all plant communities

present in the CMNP. Plants were identified by authors and specialists for different groups. Plants are housed

m the CFNL herbarium, and duplicate sets of plants were sent to different herbaria as exchange (ANSM, BRIT,

MEXU, and TEX). Previous floristic studies for Mexico, as well as monographs of most of the genera, were used

to determine families and genera origin and distribution. The plant list follows -niome’s (1992 (monocots), 2000

(dicots)) classification system for Angiospermae, but for Scrophulariaceae (in part), we followed Olmstead et al.

(2001) and Anthericaceae (Kim et al. 2010). We followed Crabbe et al. (1975) for ferns and allies (Pteridophyta) and

Eckenwalder (2009) for Gymnospermae (conifers). The families, genera, and species of each major group are alpha-

betically arranged in Appendix 1.

Diversity

We recorded 137 families, 600 genera, 1300 species, and 173

(Table 1, Appendix 1). The families with the highest number

of species are shown in Table 2 and Table 3, respectively. Tv

66.5% of the genera and species, respectively, while the 13 n

species. The dicotyledoneae are by far the most common pla

sified taxa. The Asteraceae, Poaceae, and Fabaceae families ]

and Quercus, Euphorbia, Salvia, and Ageratina stand out as th(

four genera are distributed in all plant communities in the C

Endemism, precedence, and growth forms

From the total number of species for the CMNP, 34 of then

State of Nuevo Leon. Most of the endemic species belong to

(5). Astragalus is the genus with the highest number of ende

1 infraspecific taxa of vascular plants in the CMNP

of genera and the genera with the highest number

renty-four of the families (17.6%) include 65% and

aost diversified genera include 12.23% of the total

nts in the CMNP, whfle conifers are the less diver-

highlight as the most diversified groups of plants,

e most diversified genera. These three families and

(2.5%) are endemic (see Appendix 1 (*)) for the

wo families: Asteraceae (9 species) and Fabaceae

nic species (3). By far, the native species (1222 =

775

Table 3. Genera with the highest number of species in the CMNP, Nuevo Leon, Mexico.

776

94%) dominate over the introduced ones (75 = 6%). Most of species recorded are herbaceous (1040; 80%) and

shrubs (169; 13%), followed by trees (78; 6%), and plants with fleshy stems (26; 2%). The epiphytes, parasites,

and lianas are rare.

Phytogeography

The different plant associations found in the CMNP, along with their rich plant species diversity, allow recogni-

tion of different distribution patterns of the genera throughout its altitude gradient. Table 4 shows the origin

and genera number found by plant community.

The piedmont scrub (PMS) has the highest number of genera from neotropical (130) and warm origin

(122). Most of its floristic components are from warmer areas origin, highlighting those shrubs and trees which

are important components in this landscape. These include Acacia, Acaciella, Buddleja, Condalia, Cordia, Dio-

spyros, Esenbeckia, Gochnatia, Leucaena, Persea, and Smilax, as well as frequently abundant herbaceous ele-

ments such as Abutilon, Acalypha, Begonia, Commelina, Cyperus, Datura, Euphorbia, Hibiscus, Ipomoea, Jaco-

binia, Lantana, Mentzelia, Pavcmia, Phytolacca, Ruellia, Sechium, Serjania, Sida, Tagetes, Tetramerium, Tradescan-

tia, Tridax, Tripogandra, Tumera, Verbena, Verbesina, and Viguiera. Also, the PMS has the highest typically

Mexican genera such as Ageratina, Batesimalva, Carlowrightia, Chrysactinia, Ebenopsis, Eustoma, Fleischman-

nia, Havardia, Hechtia, Hesperaloejefea, Melampodium, Mirandea, Sanvitalia, Seymeria, Tagetes, Tigridia, Vige-

thia, and Zaluzania. Almost half of the warm Mexican and neotropical origin genera registered in PMS are

found also in the rosetophyllous scrublands (RS). Both plant communities (PMS and RS) encompass the high-

est number of Mexican genera, 34 and 27, respectively. Many of these genera are found in flat plains and moun-

tains of south and west Texas (Correll & Johnston 1970).

The oak forest (OF), oak-pine forest (OPF), and pine-forest (PF), by far, cover the largest amount of tem-

perate and North American origin genera. Most of these are from temperate and Nearctic origin, and almost

always are restricted to these forests. Among the most conspicuous elements in the landscape are Abelia, Acer,

Aquilegia, Arenaria, Arracacia, Ceanothus, Cercis, Comus, Crataegus, Geranium, Gibasis, Hexalectris, Lathyrus,

Lenophyllum, Malaxis, Monotropa, Omphalodes, Parthenocissus, Phacelia, Physocarpus, Pinguicola, Prunus, Ra-

nunculus, Securigera, Silene, Sisyrinchium, Stachys, Staphylea, Thalictrum, Tilia, Torilis, Toxicodendron, Trioda-

nus, Ulmus, Urtica, Vaccinum, and Veronica. The oak forest (OF) has the third highest number of Mexican gen-

era with (23).

The juniper forest harbors the highest number of genera (77) with typically North American distribution.

The most common genera ar e Ascyrum, Calylophus, Cercocarpus, Conopholis, Echeandia, Fendlera, Fendlerella,

Helenium, Hemichaena, Heuchera, Ipomopsis, Monarda, Nothoscordum, Onosmodium, Pediomelum, Phanero-

phlebia, Physaria, Pyrrhopappus, Schkhuria, Sisyrinchium, and Stephanomeria. Many of them are shared with

the other forest types but are rarely found in the scrublands.

DISCUSSION AND CONCLUSIONS

The Mexican Transition Zone (Darlington 1957; Halffter 2003) is a complex area where the neotropical and the

the Mesoamerican Mountain Province (Cabrera & Willink 1973), to the Mesoamerican Mountain Region

(Rzedowski 1978), in part to the Madrean sclerophyllous vegetation (Graham 1999), and to the Mexican Com-

ponent of Mountain (Morrone & Marquez 2003). According to Rzedowski (1998), the Mexican flora has three

basic geographic elements: meridional, boreal, and endemic (autochthonous). Endemism is high in plant com-

munities in the north (arid areas) (Rzedowski 1973, 1978, 1988; Medellin-Leal 1982) as well as in southern

Mexico (tropical and subtropical) (Graham 1998).

Most of the surface of the Cumbres de Monterrey National Park belongs to the Sierra Madre Oriental, lo-

cated into the Mexican Transition Zone, and consists of a series of folded strata mountains, forming deep can-

yons crossed by narrow intermountain valleys, reaching a significant altitudinal gradient from 700 to 3400 m.

This factor had favored the predominance of species of temperate affinity in the highest and rugged parts of the

mountains such as Quercus, Pinus, and other conifers.

778 Journal of the Botanical Research Institute of Texas 7(2)

merits of the Chihuahuan and Sonoran Desert, such as Larrea tridentata (Hunizker et al. 1972), and genera

such as Caesalpinia, Celtis, Cercidium, Condalia, Cryptantha, Demanthus, Evolvulus, Flourensia, Gaillardia,

Hedeoma, Hymenoxys, Gilia, Malvastrum, Mentzelia, Polygonum, Proboscidea, Salvia, Schkuhria, Sida, and

Ziziphus, among others (Raven 1963; Solbrig 1972). CMNP integrates patches of deciduous forest, very similar

in plant composition to those of northeastern North America recognized by Graham (1999), and the oak-

hickory (Marroquin 1968), oak-pine, and oak-hickory-Taxodium are the most common associations in the

wettest and warmer areas (mainly in the municipality of Santiago) between piedmont scrub and oak forest. At

higher elevations occur Acer, Cercis, Cornus, Fraxinus, Juglans, Magnolia, Ostrya, Prunus, Rhus, Tilia, and Ul-

mus. These genera and several more have been cited in previous works showing disjunctions between eastern

United States and eastern Mexico (McVaugh 1943; Miranda & Sharp 1950).

All plant communities are primarily Mexican genera. However, the highest number of Mexican genera is

distributed in the piedmont scrub and rosetophyllous scrubland, in the arid part of the CMNP. Most of the

endemic species belong to Asteraceae and Fabaceae.

Main families and representative genera in the CMNP

The 11 most diversified families include 50% of the total flora, and can be used as a parameter to establish some

nparisons of CMNP regarding the global diversity (Table 5). Eight of the flowering plant families have sub-

issicaceae, Euphorbiaceae, Lamiaceae, Fabaceae, Malvaceae, Poace-

e (Fagaceae) tropical-montane-temperate (Thome 1992,

>tribution (Aster

le, and Solanaceae), one (Rubiaceae

1000). Pinaceae shows a temperate distributic

). Two families, (Cactacea<

I. From the 2700 species (and 323

lemperate distribution and Pteridaceae, (mainly in tropical-subtropical areas; (Mickel & Smith 2004), are also

d Poaceae are the most representative family plants

1998); the same diversification pattern occurs also in the study area. The family Aste

number of species (32,000) (Hendrych 1985). This family is most diverse in mountai

tropical areas, and in the warmer temperate regions (Turner & Nesom 1991

genera) occurring in Mexico, 96% of them (2600) are believed to be endei

considerable diversity is undoubtedly due to the predominance of subtropical climate, and mountains adjacent

to desert regions (Turner & Nesom 1998). The study area has a similar pattern of climate (subtropical and arid)

and topography (warm and cool mountains and dry plains) which undoubtedly favors such high diversity,

covering 29% of the genera and 7.5% of the species occurring in Mexico. Ageratina, Verbesina, and Senecio are

the most diversified genera in Mexico (Turner & Nesom 1998) and also in CMNP.

After the Asteraceae, Fabaceae is the second family of plants most diversified in Mexico (135 genera and

1724 species), growing in all ecosystems, but most numerous in tropical areas (Sousa & Delgado 1998), and

66% (121) of the genera occurring in Mexico have been recorded for the Mexican northern States (Estrada &

Martinez 2003). The CMNP contains 37% and 7% of the genera and species, respectively, for Mexico. Several

genera deserve mention due to their importance in the landscape. Havardia is a Mexican genus with nine spe-

cies, six of them endemic to Mexico (Bameby & Grimes 1997); Havardia pollens is the only species found in the

CMNP. However, it is one of the most common species in plant associations of the Tamaulipan thornscrub and

piedmont scrub (Estrada et al. 2012a). Phaseolus, another Mexican genus, encompasses 37 species, highly di-

versified in Mexico (Freytag & Debouck 2002). Thirty-four of them occur in Mexico, of which 18 are endemic

(Sousa & Delgado 1998). Nineteen percent of the total species of Phaseolus are found in the PNCM, occurring

mainly in oak forest. Dalea, another typically Mexican genus, includes 161 species (Bameby 1977), 113 of them

distributed all over Mexico. Ten of them (10%) reach the CMNP, mostly found in semiarid areas. Desmodium is

a genus widely distributed in Mexico with almost 80 species (50 of them endemic) (Lewis et al. 2005). The

study area harbors 12.5% of the total species of Mexico, and most of the 10 species found in CMNP inhabit oak-

pine forest while a few of them (D. grahamii and D. lindheimeri) are distributed also in scrublands. Two highly

779

IwS. Plant families most diversified in CMNP compared to the diversity of the State of Nuevo Leon, Mexico, and worldwide (’Hendrych 1985; 2 Vald&& Cabral

1998; Hewis et al. 2005; 4 Walker et al. 2004; 5 Mabberley 1997; 6 FLN 2010; 7 Fryxell 1 988; 8 Rzedowski & Rzedowski 2005; 9 Martinez et al. 201 1 ; 10 Mabberley 1 997;

Johnston 1997; 17 Borhidi & Diego-Perez 2002; 18 Villarreal & Estrada 2008).

1000-32,000'

800-10, 000 2

in areas with anthropogenic influence. However, Mimosa (4 species) and Acacia (6) are not as diversified as

other genera of this family. Despite legumes being distributed all over the Park, the oak forest possesses the

highest diversity, mainly in the form of herbs and shrubs, but shrubs are quantitatively most abundant in

scrublands.

The Poaceae are the fourth most diversified family of plants in the world (Valdes & Cabral 1998). In

Mexico it is represented by 183 genera and 1151 species (Gould 1979; Beetle 1983, 1987a, 1987b; Beetle et al.

1991, 1995, 1999; Valdes & Cabral 1998). According to Cross (1980), the Poaceae are abundant in “open com-

munities.” This could be the reason that this family, while being the third most diversified in CMNP (42 genera,

93 species), does not constitute a physiognomically important part of the landscape, except for some weeds

such as Cynodon dactlyon, Cenchrus ciliaris, Rhynchelytrum repens, and Beusine indica, all occasionally found in

fragmented patches with anthropogenic disturbance. Paspalum, a New World genus (Renvoize 1995; Sanchez-

Ken 2010), with almost 87 species in Mexico (Guzman & Santana 1987), 20 of which are endemic, is one of the

most diversified genera in PNCM (8 species), inhabiting almost all plant communities.

Lamiaceae is the fourth most diversified family in CMNP. Four of its genera have five or more species

(Stachys, Hedeoma, Scutellaria, and Salvia). By far, Salvia dominates over the other genera: it represents almost

1,000 species around the world (Walker & Elisen 2001). Mexico is considered one of the areas with the highest

diversity of the genus in the world (Ramamoorthy 1984; Walker et al. 2004) with almost 300 species. It is con-

sidered the second most diversified in Mexico (Comejo-Tenorio & Ibarra-Manriquez 201 1). Approximately 6%

°f the species in Mexico are in the study area. Two of the endemic species recorded for the State of Nuevo Leon

are distributed in the CMNP: Scutellaria monterreyana and Stachys vulnerabilis (Villarreal & Estrada 2008).

^ee of the most common cultivated mints— Mentha pipertia, M. spicta, and M. rotmdifolia — are introduced

in America (Bailey 1951) and are used as ornamental and medicinal plants (Estrada et al. 2012b). They are com-

ply found in CMNP (Villarreal & Estrada 2008).

Malvaceae (with more than 100 genera and 2000 species) is mainly American and probably from South

^rica in origin. However, Mexico appears to represent a region of diversification of this family (Fryxell

198 8); the family is most richly developed in the lower elevations along both coasts (Pacific and Gulf). CMNP

“dudes 25% of the genera and 6.7% of the species of Malvaceae occurring in Mexico, most of them developing

® *e lower parts of the Park and commonly found in disturbed areas. The largest genera found in Mexico (as

Wel1 38 worldwide) are Abutilon, Hibiscus, Sida, and Pavonia (Fryxell 1988). Several species of them reach

especially those of Hibiscus (1 1% of the Mexican species) and Sida (11.5% of the Mexican species). Also,

^veral species of genera “almost endemic” to Mexico (Fryxell 1988), such as Anoda, Allowissadula, and M exi-

Journal of the Botanical Research Institute of Texas 7(2)

The subcosmopolitan Brassicaceae includes species especially distributed in temperate areas (Flora of

North America 2010). Most of them are distributed in the Northern Hemisphere (Rzedowski & Rzedowsld

2005), with 616 of them occurring in the USA (Flora of North America 2010). There is not an exhaustive study

of this family for Mexico. However, 27 genera and 52 species have been recorded for the Valle of Mexico (Rze-

dowski & Rzedowski 2005), and 30 and 123, respectively, for the Chihuahuan Desert Region (Henrickson &

Johnston 1997). Half of those same genera and almost half of those species are also found in CMNP. Some

genera such as Lepidium, Eruca, and Sisymbrium constitute an important part of the weeds found in abundance

at low disturbed areas, while others such as Cardamine, Diplotaxis, Lunaria, Physaria, and Thelypodium are

found most of the time in restricted higher altitudes in forest vegetation. Rorippa nasturtium-aquaticum grows

abundantly on the banks of rivers and streams after rainy seasons.

The Rosaceae from CMNP includes mainly subshrubs, shrubs, and trees. Most of the genera recorded are

almost the same (except for Petrophytum and Agrimonia ) as those recorded for the Chihuahuan Desert region

(CDR) (Henrickson & Johnston 1997). Most of the species inhabit cool areas, especially the wild ones. From

the 36 species recorded for the CDR and Mexican Valley (Rzedowski & Rzedowski 2005), 70% of them are

distributed in CMNP. Crataegus, Rubus, and Rosa are the most diversified genera.

Mexico is a center of diversity for Solanaceae (Cuevas-Arias et al. 2008). Most of its species are distributed

in warm and temperate areas (Rzedowski & Rzedowski 2005). CMNP contains 31.5% of the genera and 6.5%

of the species occurring in Mexico. Most of the species (64%) belong to four genera: Capsicum, Nicotiana, Phy-

salis, and Solarium.

Rubiaceae is the fourth most diversified family in the world (Mabberley 1997), distributed mainly in

tropical areas. This family is well represented in Mexico (Borhidi & Diego-Perez 2002), and most of the species

are distributed in the south region (Rzedowski 1978). In the study area 11.7% and 3.5% of the genera and spe-

cies occur, respectively. The most diversified genera are Galium and Hedyotis, representing small herbaceous

species, found in almost all plant communities. Randia is an important element of the piedmont scrub, some-

times as co-dominant species of this vegetation (Estrada et aL 2012a).

Mexico has one of the most highly diversified pteridophyte flora (ferns) in the world, inhabiting a broad

range of habitats, consisting of 1,008 species and 124 genera (Mickel & Smith 2004). The CMNP includes 15%

of the genera and 4% of the species for Mexico; most of the species inhabit shaded or partially shaded areas in

oak and oak-pine forest. Among the most diversified genera in Mexico are Asplenium (86 species), Selaginella

(30 species), Thelypteris (69), Cheilanthes (60), Polypodium (55), Nothoalena (24), and Adiantum (35) (Mickel &

Smith 2004). Several species of these genera reach CMNP, but only one, Chelianthes chipinquensis, is endemic

Euphorbiaceae is one of the most diverse families in the Angiosperms (Radcliffe-Smith 1987); 50 genera

and 826 species have been recorded for Mexico (Martinez-Gordillo et al. 2002). This family is the ninth most

diverse in CMNP; three of its genera, Euphorbia, Croton, and Acalypha, have most of the species (72%). CMNP

possesses 9.1% of the Euphorbia (sensu lato) species occurring in Mexico (241).

The Cactaceae are native from America (Hunt 1999); this family is characteristic of the landscape in arid

lands of Mexico (Bravo-Hollis 1978; Bravo-Hollis & Sanchez Mejorada 1991a, 1991b). Mexico has the richest

diversity of Cactaceae (Guzman et al. 2003). Surprisingly, CMNP is not as diverse in Cactaceae as other repre-

sentative families. Even though there are arid areas on the western and the most northern portion of CMNP,

this low diversity could be caused by the relative homogeneity of the landscape and altitude gradient where the

microphyllous (in plains) and rosetophyllous scrublands (hills and piedmont) occur; both of them are quite

homogeneous. CMNP contains 12% of the genera and 3.7% of the species registered for Mexico. Three typically

Mexican genera of this family are the most diversified in CMNP: Echinocereus, Mammillaria, and Opuntia. One

of the endemic cacti ( Echinocereus viereckn ssp. morricalii ) recorded for Nuevo Leon is found in the study area.

Much like the Cactaceae, the Agavaceae are not very diverse in the CMNP; they represent only 3.8% of the

261 species occurring in Mexico. However, 50% of its genera are present in CMNP. Within Agavaceae, Agave is

the genus most diversified: 136 (Gentry 1998)-159 (Garcla-Mendoza 2011) species, with almost 77% of these

781

endemic to Mexico (Garcla-Mendoza 2011). Agave albopilosa is the only species of Agavaceae endemic to

CMNP, restricted to tall cliffs, in lithosols.

Among the woody plants, Quercus is one of the most important, and the mountains of Mexico are the

center of diversity for the Western Hemisphere (Nixon 1998). The oak forest contains the main component in

terms of biomass, particularly in the oak-pine forest, chaparral (oak scrubland), and in the rain forest (Rze-

dowski 1978). The woody species exhibit several growth forms: rhizomatous shrubs, low trees, and tall trees.

It is estimated almost 200-225 species of oak exist for the west hemisphere; from those, 130-135 species are

distributed in Mexico, and 57 of them reach the eastern portion of Mexico (Nixon 1998). Oaks in eastern

Mexico are distributed mainly in the highlands of the Sierra Madre Oriental in the State of Nuevo Leon (Nixon

1998). Almost half of these species (24, 42%) are found in the CMNP and, by far, are the most important in

terms of canopy cover and density. The most common species of the CMNP are Q. emoryi and Q. polymorpha,

associated with conifer forest and piedmont scrub, constituting ecotones among shrublands and forest.

Economically and ecologically Pinus and Quercus, are two of the most important genera in Mexico, dis-

tributed in all areas where mountains are present. The genus Pinus is represented by 90-210 species (Silba

1984; Styles 1998) of which 49 (45.5%) of them are found in Mexico (Styles 1998). The genus inhabits mainly

temperate areas. The highest diversity centers, however, are located south of the Tropic of Cancer in the south

of Mexico, Central America, and the Caribbean Islands (Styles 1998). According to Eckenwalder (2009), in the

State of Nuevo Leon there are 10 genera and 34 species of conifers (Villarreal & Estrada 2008); of those, 15

species and 7 infraspecific categories belong to Pirns. Eight genera and 15 of these conifer species are found in

CMNP, and, by far, Pinus is the most diversified genus (9 species). Its species are widely distributed above 850

m throughout the area of the Park. The most common is P. pseudostrobus in wet and temperate areas, while P.

cembroides is the most common in the arid-temperate ones. Pinus remota, one of the pinyon group, occupies the

dry-temperate valleys in the western portion of the Park. Picea mexicana var. martinezi, Taxus globosa, Cupres-

sus wzonica var. arizonica, and Pseudotsuga menziesii have more restricted distributions. Picea and Taxus oc-

cur mainly in protected wet places; Taxus grows better along creeks with rocky soils (Garcia-Aranda et al.

2012), while Picea has even more restricted distribution in the Park, growing only in one place (El Butano, 1300

m). The latter is an amazing area where Abies, Juniperus, Picea, Pinus, Pseudotsuga, and Taxus share a small (2-3

ha) bowl-shaped area, protected by 150-200 m tall cliffs where dense fogs form; that constitutes a unique mi-

crohabitat capable of sheltering such a high diversity of conifer genera.

The CMNP holds almost a third of the flora known to occur in the State of Nuevo Leon. Most of its flora is

autochthonous. The dominant plant communities are scrublands and oak-pine forest. The highest diversity of

species was recorded in the oak forest and piedmont scrub. Most of the flora recorded is from subtropical affin-

ity. However, the highest parts accommodate many genera from temperate and cool regions. Two families,

Cactaceae and Agavaceae, dominate in the arid environments of the Park but are scarcely represented in the

orest and piedmont scrub. The most diversified plant families around the world, as in genera and as in species,

are also dominant in the CMNP.

Weeds and useful plants

A number of herbaceous weed species are present in the CMNP. They are dominant elements in the farms and

halted crops fields and are evident in plains and valleys. One of the most common is Rumex mexicanus, which

invades extensive areas after rains. Farmers and ranchers in the area mentioned the plant is used to feed cattle.

^ mon g the most conspicuous species introduced from Africa (Rzedowski & Rzedowski 1990) and present in

1 e study area are Cenchrus ciliaris, Cynodon dacytlon, Eleusine indica, Leonotis nepetifolia, Rhynchelytrum re-

Prns, Ricinus communis, and Sorghum halepense, found in small patches. Almost 240 species, 170 genera, and 69

unites of plants that grow in the CMNP are commonly used in the local subsistence culture, most as medici-

products while others are used for human consumption, fodder for livestock, firewood, construction mate-

Uve fences > etc. (Estrada et al. 2007). Vida villosa is used for fodder in U.S.A. (Gunn 1979) and also grows

m °Pen fields together with Rumex mexicanus. Local residents said this species is preferred by cattle over other

Journal of the Botanical Research Institute of Texas 7(2)

786

Estrada-C. et al., Flora of Cumbres de Monterrey National Park, Mexico

797

ACKNOWLEDGMENTS

We thank Tom Wendt and George S. Hinton for allowing us to check the plant specimens of the herbaria TEX/

Hand his private herbarium, respectively. We thank those colleagues who helped us in plant identification

and corroboration of plant speciemens: Jesiis Valdes Reyna and Ismael Cabreal Cordero (Poaceae), Alfonso

Delgado Salinas (Fabaceae), Juan Antonio Encina Dominguez (Fagaceae), Socorro Gonzalez Elizondo (Cy-

peraceae). Thanks to Mario Alberto Garcia Aranda for elaboration of the map. We thar

Allred, K.W. and J. ValdEs-Reyna. 1 997. The Aristida pansa complex and a key to the Divaricatae group of North Amei

(Gramineae: Aristideae). Brittonia 49:54-66.

Anderson, E.F. 1 987. A revision of the genus Thelocactus B. & R. (Cactaceae). Bradleya 5:49-76.

Andrade, G., G.C. De Rzedowski, S.L. Camargo-Ricalde, R. Grether, H.M. Hernandez, A. MartInez-Bernal, L Rico, J. Rzedowski,

M. Sousa. 2007. Familia Leguminosae, Subfamlia Mimosoideae. Flora del Bajio y regiones adyacentes. Fascfculo 1 :

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BOOK REVIEW

Daniel Chamovitz. 2013 (first paperback edition). What A Plant Knows: A Field Guide to the Senses. (ISBN-

13: 978-0-374-53388-5, pbk.). Scientific American / Farrar, Straus and Giroux, 18 West 18th Street, New

York, New York 10011 U.S.A. (Orders: us.macmillan.com). $14.00, 192 pp., 5 %" x 8 W.

In What A Plant Knows, Daniel Chamovitz does an exceptional job at presenting a brief introduction regarding

how plants experience the world through sensory perception; that is to say, what does a plant feel, hear, smell,

or even know? Attempting to enlighten the popular reader who may lack a degree in botany but simply wishes

to learn how or if plants are capable of gaining and interpreting knowledge through sensory perception,

Chamovitz does well when explaining these biological processes unique to plants as he abstains from isolating

the laymen through unexplained terminology or an absence of simplistic illustrations to convey his points and

enlighten his readers. Amazingly, as a writer Chamovitz has produced a decent way to give an intriguing yet

brief survey on this topic which not only abridges his own decades of research but the general findings of the

scientific community as well, all the while keeping his outline clear, his pace pleasantly brisk, and the presenta-

The book itself is divided into six chapters, each one respectively addressing what a plant sees, smells,

feels, hears, how a plant knows where it is, and what a plant remembers. Interestingly, while the reader may

have picked up this text with only plants in mind, they are bound to learn something about themselves in the

process: while addressing whether a plant has a certain level of sensory perception, Chamovitz is forced to

define these senses, especially beginning with how humans experience such phenomena, and then differenti-

ate between how plants utilize the same type of sense differently regarding various aspects such as their biol-

ogy, reaction, etc. Thus through an increased awareness of their own body, the reader is left with a better un-

derstanding of both similarities and differences in how plants interpret and react to the world around them.

Another layer of interest comes in Chamovitz’s decision to relate how mankind discovered that plants

have capabilities that resemble such actions as “seeing” and “smelling.” From Darwin playing his bassoon for

his plants to Thomas Knight strapping his seedlings onto a make-shift centrifuge in the form of the humble

water-wheel, curious stories arise in What A Plant Knows of scientists who failed or succeeded, sometimes aided

by pure accident, but eventually contributing to the knowledge mankind has been able to accumulate over the

years regarding how plants act and react. As active and intelligent as the science in What A Plant Knows portrays

plants, perhaps it is no wonder that Aristotle thought plants had souls!

there. Chamovitz’s lack of bias seems evident in his chapter addressing whether plants can hear like humans;

the chapter itself feels rather bare because in terms of research, Chamovitz admits, there is still far more work

to be done. Chamovitz seems quite comfortable in acknowledging this and is in no hurry to postulate wild

speculations in order to argue that plants have all the senses a human does. His goal is not to insist that plants

experience the world just as humans do. Rather he wishes to prompt new ways of thinking about the senses,

plants, and even humans themselves.

Ultimately, What A Plant Knows is a fascinating introduction for the popular reader to the world of sensory

perception in the realm of plants. For someone unacquainted with the vast world of biological processes in the

botanical field, this accessible text serves as an excellent way to both learn and appreciate more of the complex

facets of plants themselves, as well as the field of botany, those who studv it, and even the senses of humani-

ty— Alexander Petty, Historian and Volunteer at the Botanical Res

te of Texas, Fort Worth, Texas, U.SA-

FLORA AND PHYTOGEOGRAPHY OF THE

CANON DE ITURBIDE, NUEVO LEON, MEXICO

Maria Magdalena Salinas-Rodriguezand

Eduardo Estrada-Castillon

Jose Angel Villarreal-Quintanilla

Buenavista, Saltillo

Coahuila, MEXICO

Linares, Nuevo Le6n, MEXICO

ABSTRACT

y Terrestrial Region 82. A floristic survey w

Pausras Clave: Caft6n de Iturbide, flora, fitogeografta, Noreste de Mexico

The Cafion de Iturbide area is located at the Gran Sierra Plegada in the State of Nuevo Leon and ranges from 700

to 2,900 m in elevation. The area is covered by forests and shrublands, in relatively good condition, and whose

Doristic components have not been studied deeply. The National Commission for the Knowledge and Use of

Biodiversity (CONABIO) has designated it as the Priority Terrestrial Region 82. It represents an important area

in the corridor of the Sierra Madre Oriental, connecting the preserved forests for the transit of carnivorous spe-

cies such as black bear (Ursus americanus ), cougar ( Puma concolor ), jaguar ( Panthera onca ), and jaguarundi

(Pu mayagouaroundi ). It also serves as a migration corridor to the maroon-fronted parrot (R hynchopsitta terrisi )

which is endemic to the northern Sierra Madre Oriental. This study aims to contribute to the knowledge of the

regional flora and its origin.

J - »«*- Res. Inst Texas 7(2): 803-8

Study area

The study was carried out in the Priority Terrestrial Region 82 (Arriaga-Cabrera et al. 2000). It embraces a

surface area of 42,200 ha (Fig. 1) and is located at 24 o 40 , 19"-24°55 , 43"N and 99°45 , 36 M -99°59 , 50"W. Its geolo-

gy is constituted mainly of sedimentary calcareous, folded, anticlinal and synclinal rock layers from the upper

cretaceous (Rzedowski 1978), giving rise to steep sierras with narrow central valleys. The dissolution of the

rock by water has formed narrow canyons within the Cation de Iturbide study area such as El Potosi, Jaures,

Caribeflo, Venado, Canon Seco, Las Monedas, Pablillo, and La Escondida, as well as several important moun-

tains, such as Sierra La Muralla, Sierra Borrada, Sierra Cieneguita, Sierra El Gabacho, Sierra El Novillo, and

Sierra Santa Maria (INEGI 1986, 2010). The area is part of the San Femando-Soto la Marina hydrographic re-

gion, in the Rio San Fernando hydrographic subregion, and the Rio Conchos-Chorreras basin (CONAGUA

2007). The climate varies from warm (low parts) to temperate (higher parts), and from wet to dry, in this case

from East to West respectively. Climatic differences are caused by the Sierra Madre Oriental rising from the

Llanura Costera del Golfo Norte (Gulf Coastal Plain) toward the Altiplano Mexicano (Mexican Plateau), pro-

ducing a condensation effect, that generates rains in the eastern slope of the mountain chains where fog occurs

commonly, while the western slopes develop a “rain shadow” with dry or semi-dry climate, forming low forest

of oaks, known also as chaparral. The study area has three main climate types: (A)C(Wi) or semiwarm-subhu-

mid, C(w 0 ) or temperate subhumid, and BSlh(x’) or semiarid-temperate (Garcia 1998). The main soils of the

area are shallow lithosols, limited in depth to 10 cm above bedrock (FAO-UNESCO 1988). The main vegetation

types are mixed forest (Quercus-Pinus), conifer forest ( Pinus ), broad leaf forest, p

scrublands, and chaparral (Alanls 2004; Velazco 2009).

Field and lab work

Thirty sample sites representing all the plant communities previously recognized were randomly selected and

georeferenced. Plant specimens were collected from July 2011 to April 2012. All specimens were deposited in

herbarium in the Faculty of Forest Sciences, Linares, Nuevo Leon, Mexico (CFNL). Taxa were identified using

several floristic studies for the area or near it as well as monographs for most genera (Banda 1974; Briones 1991;

Estrada 2007; Estrada et al. 2012; Gonzalez 1972, 2003; Hernandez 1998; Luna et al. 2004; Martinez & Diaz

Salas 1995; Gonzalez-Medrano 1972; Mickel & Smith 2004; Puig 1991; Rojas 1965; Velazco 2009; Villarreal &

Estrada 2008). We also consulted and used the databases for the CFNL herbarium, Hinton Herbarium (Gale-

ana, N.L.), and TEX/LL, (Austin, Texas).

Sampled sites were compared according to their plant diversity. A species matrix based on presence

(l)-absence (0) data was used for comparison among sites using Sorensen Similarity Coefficient (Mueller-

Dumbois & Ellenberg 1974) by means of the polythetic agglomerative technique (Gauch 1982; Manly 1990)

and the UPGMA method was used for cluster analysis, using the MVSP Package (KCS 2005). Vegetation types

are according to Miranda & Hemindez-Xolocotzi (1963) and Rzedowski (1978). Floristic affinities are accord-

ing to (Rzedowski 1962, 1978), HemSndez-Xolocotzi (1953), Briones (1991), and Garcia (2009).

Floristic composition

A total of 113 families, 410 genera, 698 species, and 70 infraspecific taxa of vascular plants were recorded.

(Table 1, Appendix). Families with the highest number of genera, highest number of species, and genera with the

highest number of species are shown in Tables 2, 3, and 4, respectively. Seven of the families (6%) include 43% and

49% of the genera and species, respect ivelywhile the twelve most diversified genera include 23% of the total genera.

bile Coniferophyta and Cycadophyta are

it diversified families, and

The Magnoliophyta are the most common plants in Canon de Iturbide

the less diversified groups. The Asteraceae, Fabaceae, and Poaceae families are the n

Desmodium (13), Euphorbia (12), Dalea (9), and Quercus (10) stand out as then

rsified genera.

Salinas-R. etal.. Flora of Canon de Iturbide, I

Origin of the flora and phytogeography

e most c °mmon genera in the study area are from arid areas of the world and from Tropical America (Table

5 ' The P le dmont scrub community contains the highest number of genera. This type of vegetation is located

ln lower P a «s of the area, at the foot of the mountains, in transition -with Tamaulipan thorn scrub, and it is

common to find genera from warm areas such as Acacia, Bauhinia, Berberis, Zanthoxylum, Eheretia, and Phoebe,

^swell as those belonging to the American tropics such as Acaciella, Casimiroa, Caesalpinia, Amyris, Condalia,

0r ia, Decatropis, Esenbeckia, Eysenhardtia, Gochnatia, Helietta, Persea, and Schaefferia. Commonly found

er baceous species are Abutilon, Acalypha, Indigofera, Waltheria, Phyllanthus, Priva, Tragia, Capsicum, Course-

tia ' GlIia ’ Santana, Mimosa, Passijlora, and Verbesina.

Several genera with nearctic and temperate affinity are most commonly found in the oak-pine forests and

g P arr al communities, in the highest parts of the mountains and in the western flanks, where it is common to

P Pmus and Quercus as the dominant tree layer elements, associated with shrubby genera such as Juniperus,

nus ’ ^ erc is, Ceanothus, Prunus, Crataegus, Garrya, and Juglans, while the herbaceous layer is mainly rep-

ented by Aster, Conopholis, Hedeoma, Hymenoxys, Asclepias, Centaurium, Evolvulus, Hackelia, Packera, Par-

ocissus, Philadelphia, Tagetes, Taraxacum, Urtica, Vitis, Artemisia, and Eryngium, among others. Also, in the

Parral community, Quercus is the genus dominant in the shrub layer, along with several genera of Rosaceae

tem P erate and dry areas such as Amelanchier, Cercocarpus, Cowania, Crataegus, Vauquelinia, and

Th e pure oak forests found in the study area have an ir

c of neotropical elements followed by

Table 1.F

Table 2. Families with the highest number of genera in the study area.

Salinas-R. etal.. Flora of Canon de Iturbide, Mexico

nearctic origin elements. The former are much better represented in the herbaceous layer by the genera Agera-

tina. Begonia, Bouvardia, Castilleja, Cheilanthes, Cologania, Desmanthus, Desmodium, Mimosa, Phaseolus, Rivina,

Salvia, and Stevia, and several vines such as Gonolobus, Ibervillea, Matelea, Melothria, Serjania, and Smilax. Ele-

ments from temperate origin distributed in the study area are Arbutus, Carya, Celtis, Cercis, Pistacia, Prunus,

Quercus, and Ungnadia. It is noteworthy the presence of ferns such as Adiantum, Anemia, Asplenium, Cheilan-

thes, Uavea, Mildella, Pellaea, and Pleopeltis because many are epiphytes adapted to live on the bark of the oaks

and among the litter that keeps the moisture they need to thrive.

The southwestern portion of the study area is in the “rain shadow” of the sierra and shows evident semi-

arid conditions that favor the development of desert scrub plant communities, which include mostly nearctic

genera, highlighting species of Ambrosia, Baccharis, Bahia, Bouteloua, Dasylirion, Dyssodia, Lesquerella, Leuco-

phyllum, Machaeranthera, Mortonia, and Nissolia. Also, the xerophytic vegetation hosts the greater number of

typical Mexican genera such as Agave, Ferocactus, Glandulia

Riparian communities are found along seasonal and permanent rivers and also in some creeks with sea-

sonal streams that retain moisture. The most common genera from warm-areas found in these ecosystems are

Commetina, Cyperus, Digitaria, Oplismenus, and Paspalum, but also present are genera with cosmopolitan dis-

tributions such as Apium, Eleocharis, Lobelia, Rumex, and Samolus, and even genera from temperate origin such

as Arundo, Astranthium, Coriandrum, Equisetum, Geranium, Iris, Rorippa, Seymeria, and Talinum. The most evi-

dent and distinctive genera in the tree layer of the riparian areas are Platanus, Salix, and Sambucus.

According to the cluster analysis and the dendrogram (Fig. 2), based on presence-absence of species, five

ntain groups of plant associations are recognized: xeric scrublands, oak-pine forest (with different plant asso-

ciations), piedmont scrub, Tamaulipan thorn scrub and agricultural lands (Rzedowski 1978; Miranda &

Hernandez 1963).

Group I includes three sites located at xeric scrubland at an elevation of 1310 to 1500 m and is repre-

sented by the presence of the species Acacia berlandieri, A. roemeriana, Agave lecheguilla, A. striata, Berberis

Goch° lata Echinocereus P lat y<xanthus, Euphorbia antysiphilitica, Ferocactus hamatacanthus, Fraxinus greggii.

Group II is a heterogeneous complex of sites where oaks and pines are the dominant elements at an eleva-

tion of 600 to 2000 m; however, it shows floristic differences. It includes six subgroups of plant associations.

Subgroup IIA includes two sites dominated by 10-15 m tall pure oak-pine temperate forests at an elevation of

000 m. The dominates include Quercus affinis, Q. canbyi, Q. laceyi, Q. polymorpha, Pinus cembroides, P. pseu-

strobus, associated to Carya ovata, Cercis canadensis, Juglans mollis, Juniperus deppeana, Pistacia mexicana,

Verbesina olsenii. Subgroup IIB, made up of five sites (elevation of 1100 to 1500 m) and consists of a semi-

tfry oak scrubland in transition with pine forest. Subgroup lib is similar to the Subgroup IIA but with a greater

number of Rosaceae species. The dominant species in this subgroup are Amelanchier denticulata, A. paniculata,

myris m adrensis, Buddleja cordata, Ceanothusfendleri, C. greggii, Dalea capitata, D. melantha, D, luteajunipe-

^gosturanaj.flaccida, Undleya mespilioides, Pinus cembroides, P. pseudostrobus, Pistacia mexicana, Quercus

Miformis, Q galeanensis, Q. laeta, Q. microlepis, Q. sideroxyla, Q. striatula, and Rhus virens. These sites are

Salinas-R. et al.. Flora of Canon de

809

eguilla, Comarostaphylis polifolia, Croton ciliato-glandulifer, Dasylirion berlandieri,Juniperus deppeana, O. engel-

mannii, Painteria eleachistophylla, Pistacia mexicana, Tecoma stans, Vauquelinia corymbosa ssp. heterodon, and

Yucca treculeana. Subgroup IIF included two sites of pure oak forest located at 750 to 950 m and is dominated

by Dioon angustifolium in the lower strata and by Juglans mollis, Esenbeckia berlandiri, Quercus canbyi, and Q.

rizpphylla in the higher strata. Some epiphytes were also recorded such as Tillandsia bartramii, T. recurvata, and

T.usneoides.

Group III includes six sites reaching 600 to 1100 m along the Cabezones-Potosi canyon, showing a 4-5 m

tall piedmont scrub physiognomy with abundant thorny and non-thorny species. Several species are charac-

teristic of this plant community, highlighting species of the genera Acacia, Bauhinia, Buddleja, Celtis, Cercis,

Colubrina, Cordia, Decatropis, Diospyros, Ebenopsis, Ehretia, Erythrina, Esenbeckia, Havardia, Helietta, Phoebe,

Ungnadia, and Zanthoxylum, but also, when adjacent riparian habitats, they share several species such as Adi-

antum capillus-veneris, Cyperus minimae, C. odoratus, C. thyrsijlorus, Equisetum hyemale, Lobelia cardinalis,

Platanus rzedowskii, Rorippa nasturtium-aquaticum, Sambucus nigra ssp. canadensis, Salixjaliscana, and S. nigra,

Group IV includes four sites located at medium altitudes ranging from 1500 to 1530 m in agriculture

lands in Laguna de Santa Rosa, and adjacent to the semi-dry oak scrubland. It is dominated by weed species in

the herbaceous strata such as Abutilon hypoleucum, Altemanthera caracasana, Amaranthus hybridus. Ambrosia

psilostachya, Anagallis arvensis, Anoda cristata, Astragalus hypoleucus, Argemone mexicana, Bidens Jerulifolia,

Boerhavia anysophylla, Bouchetia erecta, Cirsium vulgare, Calyptocarpus vialis, Dyssodia papposa, Gaura coc-

dnea, Helianthus annus, Ipomoea coccinea, Kallstroemia parvifolia, Lactuca serriola, Meximalvafilipes, Nicotiana

trigonophylla, Oenothera rosea, Parthenium incanum, Ricinus communis, Solanum eleagnifolium, Taraxacum offici-

nale, Urticachamaedryoides, Verbena Carolina, and Zinnia peruviana.

Finally, Group V corresponds to a single site in the lowest part of the study area ranging from 600 to 650

m on the boundary with Tamaulipan thorny scrub. This group shows species not found in the other site's

samples such as Acacia amentacea, Acaciella angustissima, Bemardia myricifolia, Caesalpinia mexicana, Capsi-

cum annuum var. minus, Canavallia villosa, Celtis laevigata, C. pallida, Colubrina greggii, Croton cortesianus, C.

fruticulosus, Dyospyros texana, Ehretia anacua, Hibiscus martianus,Karwinskia humboldtiana, Randia rhagocar-

W and Schaefferia cuniefolia.

Endemism and protection status

The study area hosts at least five endemic species for Nuevo Le6n Mexican state: Notholaena leonina, living in

oak-pine forests, in the Canon Arroyo Seco; Verbesina olsenii, inhabiting mainly oak forest, adapted to live in

®oist ravines, near the Caracol Waterfall; Anoda leonensis, inhabiting piedmont scrub in transition to oak for-

La Palma and Canon El Caribeno; Thelocactus tulensis, recorded in the rosetophyllous desert scrub,

®°nt scrub near Canon El Caribeno. In addition Dioon angustifolium, is narrowly endemic to the mountains of

de and Linares counties in Nuevo Le6n, and to the Sierra de San Carlos, Tamaulipas (Astorga et al 2005).

’ 35 P art °f this work, a new Asteraceae species was discovered: Verbesina lanulosa (in press), which lives in

1 e transition between the piedmont forest and oak forests near the settlement of La Salitrera. The new species

recor ded in the piedmont scrub on rocky hillsides, with high sun e xposure. From the total species known

01 this area we recorded seven species protected by Mexican laws (NOM - 059 SEMARNAT 2010) (Table 6).

CONCLUSIONS

The heterogeneous topography and climates over all the study area allow for the development of rich plant

v ersity as well as contrasting plant communities and different life forms. The plains, mountains, creeks, and

classifi^ate different and variable ecosystems that are home to countless species of very different origins. The

cation of vegetation allowed us to recognize different patterns of plant association based on plant diver-

V* T he most diverse plant vegetation types were the oak and the oak-pine forest, although in this plant com-

unity were more sampling sites, our results agree with those of Estrada (2007), who found that of all sampled

810

Journal of the Botanical Research Institute of Texas 7(2)

Tuu 6. Species protected by Mexican law in the study a

vegetation types (which are close to our study area), mixed forests of oak and conifer always kept greater diver-

sity of species. Nearby areas with similar plant communities are found in the south of Nuevo Leon (Cerro Pena

Nevada) and Tamaulipas (Sierra de San Carlos), and both of them contain shrub and forest communities. In

Pefta Nevada, Moreno-Talamantes (2012) recorded 485 taxa, most of them of nearctic origin, while in the Si-

erra de San Carlos, Martinez (1995) recorded 676 taxa, most of them from neotropical origin: The Canon de

Iturbide shares strong similarities with both of them, but its flora is richer in diversity since it has all ecosys-

tems found in both areas, and it can be highlighted as an area for conservation of its plant diversity and its en-

demism. Five endemic species in the State of Nuevo Leon were recorded in the area: Anoda leonensis, Carda-

mine auriculata, Dioon angustifolium, Notholaena leonina, Thelocactus tulensis, and Verbesina olsenii. From the

768 taxa recorded, seven of them are protected by NOM 059-SEMARNAT-2010: Agave bracteosa, Braheaber-

landieri, Dioon angustifolium, Echinocactus platyacanthus, Litseaglauscecens, Pinus strobiformis, and Thelocactus

tulensis. The discovery of a new species, Verbesina lanulosa (in press), in this area shows that still more botani-

cal work is necessary to complete the floristic studies of northeastern Mexico. The plant diversity recorded in

the Canon de Iturbide includes genera and species from different origins, but the tropical elements are quanti-

tatively the most important ones, since they represent about 52% of families, 68% of genera, and 32% of the

species, and they are well represented in the different plant communities, especially piedmont scrub, oak for-

ft. et al.. Flora of Canon de Iturbide, f

818

Lantana camara L., E.E. 21044; M.S. 867.

1. Martens & Galeotti, M.S. 58.

Phyla incisa Small, M.S.2.

Verbena canescens Kunth var. canescens, E.E. 21 709 M.S. 1 1

Verbena Carolina L, M.S. 964.

Vitis berlandieri Planch., M.S. 1333.

Vitiscinerea Engelm., E.E. 21436, 21876.

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820

Journal of the Botanical Research Institute of Texas 7(2)

BOOK REVIEW

David Moore. 2013. Fungal Biology in the Origin and Emergence of Life. (ISBN-13: 978-1-107-65277-4, pbk.)

Cambridge University Press, 32 Avenue of the Americas, New York, New York 10013-2473, USA 1 1

(Orders: www.cambridge.org, 1-212-337-5000). $42.99, 236 pp., 28 b&w illustrations, 2 tables, 6" x 9"

Mycologists will welcome this book! Too often fungi are ignored in discussions about the origin of life as well

as their importance to world ecosystems. This book, as the title implies, considers the role of fungi in later eu-

karyote evolution. Heretofore emphasis on prebiological evolution, protocells, and biofilms led to prokaryotic

discussions of the earliest pathways of evolution and mostly ignored the role of fungi. Moreover, the central

thesis of this book “. . . is based on appreciation of the central role of the fungi grade of organization in the evo-

lution of higher organisms.”

Chapters 1 and 2 discuss the diversity of present-day fungi in form, function, and biology. Chapter 3 de-

scribes the formation of the solar system that gave birth to the Earth and Moon. Chapter 4 highlights the car-

bon-based molecules present that contributed to the early building blocks of organic compounds necessary for

life on Earth. The panspermia hypothesis (extraterrestrial origin of life) is explored in Chapter 5 which is not

a viable scientific option for the author. In Chapters 6 to 10 (pages 70-141) much attention is given to the first

formation and definition of life on young earth. The reader has to wait more than halfway through the book for

Chapter 11 “Toward Eukaryotes.” However, one of the more interesting observations is based on the theme

“are animals necessary?”, especially since humans asking the question are animals. The author argues fora

balanced life system: bacteria to begin, green plants as producers, and fungi as decomposers. The bottom line

is that Earth could exist without animals and still be balanced. I recommend Chapter 8 “It’s life, Jim . . .” for any

general biology student who wants to review the properties of a “living” system: LIFE.

Finally we arrive at the “Rise of the Fungi” in Chapter 12 and the discussion of Prototaxites, the largest

organism present during the Devonian period (up to 9 meters tall). Recent evidence strongly suggests that this

was a giant terrestrial saprotrophic fungus with hyphae and spores and was the dominant living form on this

ancient landscape. Imagine a fungus the size of a small tree that lasts for 40 million years about 420 million

years ago. Add to this the incredible diversity of fungal forms — chytrids, sexual reproductive stages, Ascomy-

cetes, and more, beautifully preserved, found in the Devonian Rhynie Chert of northern Scotland, mostly by

Tom Taylor and coworkers at the University of Kansas — and one appreciates that fungi have played a signifi-

cant role in the evolution of life on planet earth for a long time. Far longer than once thought! More discussion

here includes the importance of sclerotia (fungal mass of hyphae), free cell formation, filamentous growth, cell

fusion, and septum (cross wall) formation that are all found in fossil and extant fungi.

This is a must-read, fascinating book for anyone interested in evolution or fungi. Some chapters are highly

technical, but others are easily understood by a nonscientist. Table 1 gives a geological and biological chronicle

of events on Earth from 2500 million years ago to present day. References are cited and occupy pages 204-218.

many of more recent origin. There is no glossary which sometimes complicates some of the terminology, but

this is a minor distraction. The Index (pages 219-231) is a convenient shortcut to specific topics.

Every scientist interested in the broad topic of evolution of life on Earth should have this book on their

bookshelf. It is affordable, readable, understandable, and easily could be used in evolution-based seminars or

as selected reading for general biology courses. — Harold Keller, PhD, Research Associate, Botanical Research In-

stitute of Texas, Fort Worth, Texas, U.S.A.

NUEVOS REGISTROS DE CACTACEAE Y SOLANACEAE PARA

EL ESTADO DE GUANAJUATO, MEXICO

Maricela Gomez-Sanchez 1 , B.A. Gonzalez-Hernandez,

Mahinda Martinez & Luis G. Hernandez-Sandoval

Facultad de Ciencias Naturales

Avenida de Las Ciencias s.n.

76230 Quer&aro, Qro., MEXICO

is de distribution geografica de ti

En anos recientes, el conocimiento sobre la riqueza floristica del estado de Guanajuato ha mejorado notable-

ahora son parciales e incompletos. Las estimaciones mas recientes senalan que el estado de Guanajuato alberga

182 familias, 904 generos y 2774 especies de plantas vasculares (Zamudio & Galvan 2011). De esta riqueza solo

35 especies estan registradas en la Norma Oficial Mexicana, NOM-ECOL-059-2001 (Anonimo 2002) como

amenazadas o con protection especial y una en peligro de extincion, de las cuales 31 son endemicas de Mexico

y tan solo 26 son exclusivas de Guanajuato y la mayoria conocidas solamente de la localidad tipo. Conside-

rando el alto deterioro de la vegetation del estado y que muchas especies estsin sometidas a grandes presiones

de recolecta con fines omamentales, el bajo numero de especies protegidas resulta preocupante. La recolecta y

revision de material botanico como producto de estudios floristicos siguen aportando information notable que

incrementa el conocimiento de la flora mexicana. Asi, las cifras hasta ahora conocidas para el estado de Guana-

juato se siguen incrementando, pues investigaciones recientes siguen revelando la existencia de plantas que no

se habian registrado anteriormente para la entidad y no seria raro el descubrimiento de plantas que aun no han

sido descritas. Durante el desarrollo de estudios floristicos en el Area Natural Protegida “Sierra de Lobos” del

Estado de Guanajuato, en el Bajio mexicano, destaca el hallazgo de localidades no registradas previamente para

tres especies de Cactaceae y Solanaceae. Las recolectas recientes que provienen de bosque de encino en el Mu-

uicipio San Felipe, expande el Area de distribucidn conocida de Mammillaria mathildae Kraehenb. y Krainz

siendo este el primer registro para el estado de Guanajuato. Especlmenes de recolectas recientes en la ex-Haci-

enda San Juan de Otates, en el Municipio de Le6n, extienden el area de distribution de Physalis longiloba Var-

gas, M. Martinez & Davila (Cactaceae), taxon microendtinico conocido solamente de la Sierra de Manantlan

Bot Res. Inst. Texas 7(2): 8

822 Journal of the Botanical Research Institute of Texas 7(2)

en el estado de Jalisco y es un registro nuevo para Guanajuato. Asimismo, especimenes procedentes de El

Capulin, Municipio San Felipe extienden los llmites de distribucion de Physalis waterfallii Vargas, M. Martinez

y Davila, un taxon de distribucion mas amplia pero endemico de los estados de Jalisco y Michoacan, siendo este

el primer registro para el estado de Guanajuato.

Los especimenes de los taxa citados provienen de recolectas recientes que se hicieron en el Are

gida Sierra de Lobos, en el estado de Guanajuato, en la region del Bajio mexicano. Estos ejei

diaron detalladamente y la determinacion de la identidad de las especies se hizo mediante cto

nes de distintas obras floristicas (Bravo-Hollis & Sanchez-Mejorada 1991; Glass 1998; Scheim

et al. 1999, 2001, 2003). Los ejemplares botanicos estan depositados en el herbario QMEX.

n pocos los recolecto

Algunos especimenes de recolectas recientes se identifican

Physalis longiloba (Solanaceae) y Physalis waterfallii (Solanaceae) provenientes del bosque templado de Quercus

de los municipios San Felipe y Leon respectivamente, en el estado de Guanajuato. Las tres localidades amplian

la distribucion conocida de estas especies y representan el primer registro para el estado.

Mammillaria mathildae Kraehenb. y Krainz es una especie microendemica que habita en lugares secos del

bosque tropical caducifolio. Actualmente solo estan reconocidas y documentadas dos poblaciones de esta es-

pecie, la localidad tipo al sureste de la capital del estado de Queretaro, en La Canada, municipio El Marques

(Bravo-Hollis & Sanchez-Mejorada 1991; Glass 1998; Hernandez & Sanchez 2002) y la segunda al noroeste de

la capital queretana en Los Cajones, Provincia Juriquilla, municipio de Queretaro (Hernandez & Sanchez

2002). Esta especie habita areas perturbadas del bosque tropical caducifolio, entre 1830 y 2030 m de altitud,

donde destacan especies como:

Burserafagaroides (HBK) Engl. Myrtillocactus geometrizans (Mart.) Cons.

Bursera palmeri S. Wats. Opvmtia spp. Mill.

Calliandra eryophylla Benth. Senna poly anta (Collad.) Irwin yBameby

Ceiba aesculifolia Britt, y Baker Stenocereus dumortieri (Scheidw) Buxb.

Lysiloma microphylla Irwin y Bameby

En la localidad La Canada tambien estan presi

deterioro de la vegetacidn original (Zamudio e

Acacia famesiana (L.) Willd.

Karwinskia humboldtiana (Roem. y Schult.) Zucc.

Ipomoea murucoides Roem. y Schult.

Mammillaria mathildae Kraehenb. y Krainz es una especie catalogada como vulnerable por la IUCN (Walter &

Guillett 1998) y amenazada por la NOM-059-ECOL-2001 (Anonimo 2002). Sin embargo estudios detallados

de sus poblaciones (Hernandez & Sanchez 2002) la senalan como “en peligro de extincion” El habitat cono-

cido en la localidad tipo de esta especie se reduce a poco mas de una hectarea y los agentes de disturbio que

J poblacion son la ganaderia no planeada, la extraccion de lefta, la extraccion de

. 1992; Hernandez & Sanchez 2002), t

afectan considerablemente

i, la extraccion de plantas con fines o

landestinos (Cabrera-Luna & Gomez-Sanchez 2005).

La localidad del reciente especimen de Mammillari

1 estado de Guanajuato y amplia su distribucion y su

ntales y la presencia de tiraderos de basura y de escombro

823

presion de recolecta por su potential ornamental por lo que los datos precisos de esta localidad se omiten aqui

para evitar un posible saqueo de individuos. No obstante, cuando sea de interds, la information podra soliti-

tarsele al primer autor. Lo notable de esta nueva localidad es que alberga un bosque templado de Quercus L.,

con una altitud sobre los casi 2300 m, que sale de los lxmites conocidos hasta ahora, cerca de los paralelos 20°

Lat N, 100° Long W y 1850 msnm en los alrededores de la ciudad de Querdtaro. Esta nueva localidad es un

complejo de lomerios con bosque de encino compuesto principalmente por:

Quercus eduardii Trel. Quercus resinosa Liebm.

Quercus grisea Liebm. Quercus rugosaNee

En el estrato herbaceo, es <

iun la presencia de especies como:

Dasylirion sp. Zucc. Physalis patula Miller

Loeselia coerulea (Cav.) G. Don Tagetes sp. L.

Mimosa sp. L.

La presencia de Mammillaria mathildae Kraehenb. y Krainz es mas frecuente en pequenas areas abiertas por la

perturbation, dentro del encinar. Este hallazgo reciente tal vez representa el llmite de distribution de la espe-

cie, que va desde los alrededores de la ciudad de Queretaro Uegando hasta San Felipe, Guanajuato. Este taxon

sigue un patron de vegetation en el que se altema el bosque tropical caducifolio, el bosque espinoso y el mator-

ral xerofilo (como ocurre en la localidad tipo) Uegando al bosque de encino. Los registros de esta especie au-

mentan a medida que se hacen mas exploraciones y esto amplia su distribution. Asi entonces, una exploration

exhaustiva y una nueva evaluation de las poblaciones de Mammillaria mathildae, aunque pequenas, podrian

modificar su estatus de conservation hasta ahora conocido.

Physalis longiloba Vargas, M. Martinez y Davila es una especie endemica conocida, hasta ahora, solamente

de la Sierra de Manantlan, Reserva de la Biosfera, en el estado de Jalisco al occidente de Mexico. Crece de

manera abundante en lomerios, en areas abiertas o senderos del bosque mesofilo de montana y de bosque de

pino-encino muy humedo, entre las coordenadas 19°35’55” Lat N y 104°12’35” Long W, a 2100 msnm. Physalis

longiloba es superficialmente similar a P. lignescens Waterf. y se puede confundir con P. gracilis Miers (Vargas et

al. 2001, 2003). Sin embargo, el caliz de la Uor de 1.5 mm de longitud con largos lobulos acuminados de hasta 9

mm y el caliz cuando frutece con 5 angulos, en los ejemplares recientes, mostraron claramente su identidad.

Las recolectas recientes de Physalis longiloba (Mexico. Guanajuato, Ex Hacienda San Juan de Otates, mu-

824

Journal of the Botanical Research Institute of Texas 7(2)

nicipio de Leon, 21°12'58'' Lat N, 101°32’10" Long W, 2641 m de altitud, A. Gonzdlez et al 733-737, QMEX) son

nuevos registros para el estado de Guanajuato y amplian el area de distribution conocida para este taxon al

Bajio mexicano, region situada al noroeste de la distribution conocida hasta ahora. En esta localidad, PhysaHs

longiloba (Fig. 1) habita en un bosque de encino dominado por Quercus rugosa Nee, Q. eduardii Trel. y Q. poto-

sina Trel. En el estrato herbaceo destacan especies como:

Aegopogon cenchroides Humb. & Bonpl. ex WiUd.

Erodium cicutarium (L.) L’Her. ex Aiton

Eryngium carlinae F. Delaroche

Loeselia coerulea (Cav.) G. Don

Physalis sordida Fernald

Physalis chenopodifolia Lam.

Piptochaetium brevicalyx (E. Foum.) Ricker

Salvia spL.

Aunque esta nueva localidad amplia su distribution geografica, P. longiloba sigue representando un microen-

demismo, con presencia unicamente en la Sierra de Manantlan al sur del estado de Jalisco y la nueva localidad

en el muniripio de Le6n, Guanajuato.

Physalis waterfallii Vargas, M. Martinez y Davila es una especie de distribution mas amplia pero endemica

del estado de Jalisco y de una localidad al norte de Michoacan. Habita en bosques de pino-encino y mesofilo de

montana, tambien es comun localizarla en claros o pendientes, a veces asociada a cultivos y con frecuencia a la

sombra de encinos (Vargas et al 1999, 2003), a altitudes entre los 1700 y 2450 msnm. Esta especie se reconoce

facilmente en campo porque sus lobulos calicinos en la flor y en el fruto son largos, alcanzan hasta 1 cm de

,io TTT reClenteS de Physalis water f allii (Mexico. Guanajuato, El Capulin, municipio San Felipe,

1 22 26.1 Lat N, 101°18'54.66" Long W, 2385 msnm, A. Gonzdlez et al 159, QMEX) son registros nuevos para

el estado de Guanajuato. Esta localidad es una area comunal donde se permite el aprovechamiento de sus re-

cursos naturales y como Unidad de Manejo para la Conservation de la Vida Silvestre (UMA), esta registrada

ante la becretana de Medio Ambiente y Recursos Naturales (SEMARNAT, 2012) c

UMA-Ext-0019-Gto. En este sitio P. waterfallii (Fig. 1) habi

diiy Q. potosina. En el estrato herbaceo destacan especies o

el cOdigo SEMARNAT-

Aegopogon cenchroides Humb. & Bonpl. ex WiUd.

Bouteloua gracilis (Kunth) Lag. ex Griffiths

Bouteloua hirsuta T a g

Castilleja arvensis Schltdl. & Cham.

Dalea bicolor Humb. & Bonpl. ex Willd.

Desmodium aparines (Link) DC.

Desmodium grahamii A. Gray

Echinopepon milleflorus Naud.

Loeselia mexicana (Lam.) Brand

PeUaea cordifolia (Sesse & Moc.) A.R. Sm.

Pellaea temifolia (Cav.) finlf

Physalis hastatula Waterf,

Piptochaetiumfimbriatum (Kunth) Hitchc.

Salvia patens Cav.

Silene laciniata Cav.

Stachys coccinea Ortega

Tagetes lunulata Ortega

Con este nuevo hallazgo, se observa que Physalis waterfallii

encino del Occidente al Bajio mexicano. No obstante, esta espe

a la parte centro-sur del estado de Jalisco, una localidad en el f

i de los bosques humedos de

n una distribution restringida

al norte del estado de

1 noroeste del estado de Guanajuato,

agradecimientos

Lwmf I t ^ que atienden estos lugares y que amablemente hicieron posible la recolecta de

logia (CONACvT) aTrav^d 00 A * ^ ^ ^ aP ° y ° financiero del Consejo National de Ciencia y Tecno-

y a un icvisor anommo que amablemente mejoraron el escnto.

Gomez-Sanchez et al., Nuevos registros de Cactaceae y Solanaceae

825

de flora y fauna silvestres - 1

especies en riesgo. Secretary

Bravo-Hollis, H. y H. SAnchez-Mejc

para su inclusion, exdusibn o cai

lies. Diario Oficial de la Federacibi

1. 3. Universidad Nacional Autono

anada, Queretaro, Mexico. Bol. !

Cabrera-Luna JA y M. GOmez-SAnchez. 2005. Analisis floristico de La Canz

7735-50.

Glass, C. 1 998. Guia para la identification de CactSceas amenazadas de Mexico. Vol. 1 . Conabio-Cante. Mexico.

Hernandez M., M.M. y E. Sanchez M. 2002. Informe de una nueva localidad de Mammillaria mathildae, una propuesta para

modificar su categorfa legal de Conservacibn. Cact. Sue Mex. 47:4-1 0.

Scheinvar, L. 2004. Flora Cactolbgica del estado de Queretaro: diversidad y riqueza. Fondo de Cultura Econbmica. Mexico.

SecretarIa de Medio Ambiente y Recursos Naturales (SEM ARNAT). 201 2. http://www.semarnat.gob.mx/temas/gestionambien-

tal/vidasilvestre/Paginas/umas.aspx (Consulta: Mayo 2012).

Vargas, O., M. Martinez y P. Davila A. 1 999. Physalis waterfallii (Solanaceae), una especie nueva de los estados de Jalisco y

Michoatiin. Acta Bot. Mex. 48:21 -26.

Vargas, 0, M. Martinez y P. Davila A. 2001 .Two new species of Physalis (Solanaceae) endemic to Jalisco, Mexico. Brittonia

53505-510.

Vargas, 0., M. MartTnez y P. Davila A. 2003. La familia Solanaceae en Jalisco, El genero Physalis. Flora de Jalisco 16:1-126.

Walter, K.S. y H J. Guillett, eds. 1 998. 1 997 Red list of threatened plants. Compiled by the world conservation monitoring

centre. lUCN-The world Conservation Union, Gland, Switzerland and Cambridge, UK.

Zamudio, S., J. Rzedowski, E. Carranza y G. CalderOn de Rzedowski. 1992. La vegetacidn del estado de Queretaro. Concyteq-

Instituto de Ecologia A.C., Centro Regional del Bajio. PStzcuaro, Michoacan, Mexico.

Zamudio, S. y R. Galvan V. 201 1. La diversidad vegetal del estado de Guanajuato, Mexico. Flora del Bajio y de Regiones

Journal of the Botanical Research Institute of Texas 7(2)

BOOK REVIEW

Peter Roberts and Shelley Evans. 2011. The Book of Fungi: A Life-size Guide to Six Hundred Species from

Around the World. (ISBN-13: 978-0-226-721172-0, cloth, alk. paper). University of Chicago Press, 1427

E. 60th Street, Chicago, Illinois 60637, U.S.A. (Orders: Customer Service, Chicago Distribution Center,

11030 South Langley Avenue, Chicago, Ilinois 60628, 1-800-621-2736 (USA and Canada), 1-773-702-

7000 (International), orders@press.uchicago.edu). $55.00, 656 pp., 2000 color plates, 7.5" x 11" x 2".

This is a big, weighty book illustrated with life-size color images of fungal fruit bodies that appear in ones,

twos, or small groups but not in natural habitats. Each fungal species is given one full page that has the some

standardized information: for example Agaricus campestris (Field Mushroom) starts with a brief description

including habitats such as lawns, parks, pastures, and other areas of undisturbed short grass. Another section,

Similar Species, notes species that appear morphologically similar but with a brief description of the differ-

ences. At the top fourth of the page is a world geographical map with the family, distribution, habitat, associa-

tion, growth form, abundance, spore color, and edibility. The height and diameter of fruit bodies is given in an

The beginning sections include a Foreword; Introduction; What are Fungi?; Plant and Animal Partners;

Natural Recyclers; Pest and Parasites; Food, Folklore and Medicine; Distribution and Conservation; Collecting

and Identifying Fungi; and Guide to the Fungi. The authors state, “This book is not a field guide . . .” and there-

fore keys to families and species are lacking. A picture guide is provided that highlights groups— e.g., “Agarics”

are defined as having “fleshy fruit bodies, cap with or without stem, gills underneath cap,” and this is the first

group discussed, described, and illustrated beginning on page 31 and ending on page 323, representing 292

species or almost half of the 600 species. Next are the boletes, or fleshy pore fungi, on pages 325-361. The wood

rotters are represented by the brackets, crusts, and jelly fungi on pages 362-461. Tooth fungi, chanterelles,

clubs, and corals are a mixed bag of very different spore-bearing surfaces with little to argue for this grouping

on pages 462-505. The puffballs and earthstars, bird’s nests, and stinkhoms conclude the Basidiomycetes on

pages 507-547. The Ascomycetes begin with the cup fungi, morels, truffles, flask fungi, and lichens on pages

The text is written in nontechnical prose, readily understandable by the lay public, but a short 2-page

Glossary of terms also aids the reader. Supplementary information is included in the following sections: Re-

sources for further reading and general interest. The Classification of Fungi, Index by Common Name, and

Index by Scientific Name. Some of the common names applied here are clever andnew, for example, Beansprout

mte^olflahoi ^J eliqU * SCenS) ’ GreCn Skinhead ( Cortinarius austrovenetus ), and Collard Parachute (M aras-

This book is too bulky and heavy to carry into the field and therefore has limited value other than as a

coffee table display that highlights the diversity of fungi in color and form. Some species can be identified by

picture keying. Many mushroom books are similar in content especially in the introductory sections. The price

is a bargain for a book this size containing color illustrations.— Hamid W Keller, PhD, Research Associate, Bo-

tanical Research Institute of Texas, Fort Worth, Texas 76102, USA.

J. Batltes. Inst Texas 7(2): 826. 2013

SEDUM SALVADORENSE (CRASSULACEAE), UNA ESPECIE ENDEMICA Y RARA

REDESCUBIERTA PARA LA FLORA DE EL SALVADOR

Frank Sullyvan Cardoza Ruiz

Consultor independiente

Master of Science en ManejoyConservacidn

de Bosques Tropicalesy Biodiversidad (CATIE)

San Salvador ; EL SALVADOR

La Ceiba, HONDURAS

RESUMEN

ABSTRACT

INTRODUCClON

La familia Crassulaceae agrupa de 1,400 a 1,500 especiesy 34-35 generos. Son plantas perennes o rara vez anua-

•es o bianuales. Se caracterizan por tener generalmente hojas y tallos suculentos y flores hermafroditas, acti-

nomorfas, a menudo pentameras, con un ovario supero, carpelos libres y dehiscentes con una escama necta-

dfera en la base de cada uno. El metabolismo de muchas especies es del tipo del acido crasuldceo (Thiede &

Eggli 2007). Las especies de esta familia tienen una distribucion casi cosmopolita con prominentes centros de

diversidad en Mexico, Sudafrica, Este de Asia y la cuenca del Meditemineo; prosperando generalmente en zo-

tos montanosas sobre sustratos rocosos. Unos pocos representantes son epifitos o acuaticos (Mort & Mori

2004). En los tropicos, las especies estan confinadas a las regiones de montana (Standley & Steyermark 1946)

P°rsu follaje colorido y suculento, muchas son ampliamente cultivadas como plantas ornamentales.

Thome y Reveal (2007) propusieron el reconocimiento de unicamente dos subfamilias: Crassuloideae y

Sempervivoideae; esta ultima, contiene la mayor diversidad taxonomica, incluyendo aproximadamente 980

species y 28 generos y ha sido subdividida en cinco tribus por Thiede y Eggli (2007). Sedeae, la mas grande de

estas tribus, comprende aproximadamente 640 especies incluidas en dos grupos que no han sido aun nombra-

dos formalmente: el clado Leucosedum y el clado Acre (Carrillo 2009), con cerca de 530 especies (una tercera

Pane de la diversidad total de la familia), incluye a los representantes de Sedum (subgenero Sedum). El problema

** complicado que enfrenta la sistematica de la familia Crassulaceae es la parafilia de Sedum (Carrillo 2009),

iht >*tetT(HBj7(2):S27-«

828

Journal of the Botanical Research Institute of Texas 7(2)

fotografiadas ;

siendo un genero muy variable y sus limites taxonomicos no se han resuelto todavia (‘t Hart & Bleij 2003). El

genero Sedum L„ tiene distribucion cosmopolita; sin embargo, la mayoria de las especies crecen mejor en las

zonas templadas del Hemisferio Norte y comprende cerca de 420 especies. Alrededor de 170 especies crecen en

el Continente Americano ( t Hart & Bleij 2003) y en El Salvador, S. salvadorense es la linica especie del genero.

El redescubrimiento

Durante la expedition del botanico Paul C. Standley a El Salvador, en los meses de diciembre de 1921 y mayo

de 1922, especificamente en el occidente del pais (Ahuachapan) se encontro con una colonia de especies del

genero Sedum en la Finca Colima. Standley relata que estas plantas estaban “algo marchitas como resultado de

la larg3 estacion seca (17-19 enero de 1922).” Sin embargo, afirmo en aquel tiempo que esta especie se distin-

gma claramente de otras reportadas del centre de los Estados Unidos o de Mexico. Es asi como publico en el

Journal of the Washington Academy of Sciences en 1923 la especie Sedum salvadorense. El ejemplar tipo se en-

cuentra depositado en United States National Herbarium (US) con el numero 1,136,003.

Durante los recorridos realizados por medio del Proyecto “Mejor Manejo y Conservation de Cuencas

Hidrograficas Criticas” (MMCCHC), dentro del Parque Nacional El Imposible, al occidente del pais en los me-

ses de diciembre de 2007 y enero 2008, se encontraron plantas sobre rocas y materia organica que parecian ser

del genero Echevena, sin embargo despues de un analisis mas detallado se Uego a la conclusion que las plantas

olonias (individuos) que pertenecian a la rara y endemica es-

nlM . t .. :olectas, solo se conocia el ejemplar tipo por lo cual se conside-

d ’ ° r i a ^ e pres ^ ta en este articulo como un redescubrimiento colonias de especies en dos lugares

toThihifr l r e ,? CI ° n j ,mp0sible ' y tambien * ^ primeras fotograttas de la especie

(en su habitat y cultivadas) despues de 90 aflos de su publication.

Sedum

Desde su creation por Linnaeus, Sedum ha tenido limites difusos y ha s

caracteres morfologicos. Los analisis filogeneticos han demostrado que t

definido por atributos plesiomorficos. Dilucidar los limites y las relaciones de Sec„ ,

uno de los principales retos en la sistematica de las Crassulaceae. Por lo tanto, es probabrqL^rjfuture^-

,ez del material y al desconocimiento de sus caracteristi-

ise no ha sido analizada filogeneticamente, pero es probable que

mayor diversidad de especies.

n ser bastante estables en lo morfologico, otras, por el contrario, pueden

sus lurmas sean ditiriimpm .J° qUeSerefierealindumentt >y Porte entreotros que hace que algunade

Por ral ra zd„, se describeT™"ub a ZZ'ZT" “ raiSm0 T (CaS,I0Viej0 & ^

mando para ello medidas tamo de plantas en el cTmpo como IWmd^su^bilaTfcuhivadas)^ 3 ^^ 3 ^ l 'P' ca ' ^

DESCRIPCION DE LA ESPECIE

Sedum salvadorense StandLj. Washington Acad. Sci. 13:438. 1923. (Fig. 1). Tip<r El Salvador DepaktamentodeAhua-

chapA N: collected on a rock in forest, Finca Colima, Sierra de Apaneca, 17-19 Jan 1922, Standley 20143 (holoitpo: US-Imaged).

Son plantas perennes, con tallos sufructicosos, de unos 4-20 cm de ahum v m j .

^„th7ip^!r!i , ref e 7r sl 7 h T c T“ pa ' ulada " < ' bkn “ o1 ^

JOve” h T' ^ abaj ° ““n Rancho. piano, delgadoydacL. verde, las

'y , . P aios 3 •guales y camosos, verde-claro lustroso lisos de 1 rm de lareo

pecie Sedum salvadorense. Antes de r<

aum sea segregado en vanos generos. Debido

cas morfologicas y moleculares; S. salvadorer

pertenezca al subgenero Sedum donde

gunas especies parece

nte, especial]

s difusos y ha sido dificil de circunsc

s parafiletic

ir grandem

igualando los sepalos, cuspidado-agudo. Raiz tuberosa de 0.9-1.5 cm de largo y 04-0 6 mm de ancho cod

forma turbinada, esferica y fusiforme (Figs. 2 y 3).

Habitat. Crece sobre rocas verticales bas^lticas (acantilados) en colinas pedregosas, pequenos cauces de

agua o riachuelos con vegetacidn riparia en el bosque tropical latifoliado semideciduo submontano, bien dre-

nado entre los 720 hasta 1,130 msnm (Figs. 4 y 5). En la localidad tipo se observaron creciendo en el estrato

e, redescubierta de El Salvador

833

arboreo Garcinia intermedia (chaparron), Brosimum alicastrum (ujushte), Calophyllum brasiliense (varillo),

Swartzia simplex (naranjillo) y otras. En el estrato arbustivo existen Hirtella racemosa, varias especies de Psy-

chotria y algunas Nictaginaceas muy raras como Boldoa purpurascens, Pleuropetalum spruce i y Pisonia donnells-

mithii. En el estrato herbdceo son raras las grammeas y ciperaceas pero hay una gran diversidad de los generos

Justicia, Ruelliay Blechum. Abundan tambien Geophila repens y alguna piperaceas. Hay algunas palmas del ge-

nero Chamaedorea, algunas commelinaceas y orquideas terrestres de los generos Sacoila, Corymborkis, Tropi-

ca, Habenaria y M alaxis. La especie S. salvadorense crece espectficamente en los lugares mas secos, afloramien-

Pefia), algunos arbustos espinosos de mimosaceas, mientras que en los paredones y barrancos mas humedos se

pueden encontrar helechos pequenos y ocasionalmente algunos arborescentes como Cyathea costaricensis

(MARN 2012).

Fenologia. — Florece y fructifica de diciembre a febrero.

Etimologia. — Sedum es el nombre de diversas crasulaceas. El epiteto de la especie fue dedicado al pais de El

Salvador por el prolifico botanico estadounidense Paul C. Standley (1884-1963), maestro e investigador, quien

Visit6 y recorrio gran parte del pais por un periodo de 6 meses entre diciembre de 1921 y mayo de 1922, colec-

tando y describiendo especies nuevas para la ciencia.

Comentarios. — Muchas de las especies de esta familia se han adaptado a vivir en habitats extremada-

mente secos, por lo cual han desarrollado estructuras que permiten evitar la deshidratacion, como pruina,

Pelos> spinas; hojas y tallos camosos para acumular agua. Para sobrevivir bajo esas condiciones, se han espe-

834

a gemulas, renuevos esteriles, hojas enraizantes y tallos

ital por sus bellas flores blancas-amariUeii-

e esta especie considerandola extinta. Fue hasta 1997 que E.

Sandoval s.n. (LAGU) y E. Sandoval 1884 (LAGU, MO, B, BM, EAP) la colecto, es deck 75 anos despues de la

primera colecta de Paul C. Standley. Sin embargo, es hasta 2009 y 2012 que pudo ser colectada y documentada

fotograficamente por los autores, detallando de forma precisa la georeferenciacion de las localidades, pues las

coordenadas de las otras colectas no fueron asignadas en el campo sino en el herbario, tomando como punto de

CONCLUSIONES

En referencia al Estado de Conservation de esta especie; creemos que las plantas dentro del area del Parque

National El Imposible son poco frecuentes considerandose extintas en otras zonas debido al alto grado de de-

terioro y fragmentation de su habitat. De acuerdo con los criterios de la Lista Roja de Espeties Amenazadas

(UICN 2012) y segun las observaciones de los autores, estas plantas son las unicas poblaciones conocidas

dentro del Parque Nacional El Imposible, por lo tanto la categoria para esta especie seria Vulnerable, VU Al(d).

Nuestra gratitud a los dos revisores uno anonimo y para Tiana Rehman de BRIT por sus valiosos comentarios

y observaciones. Tambien a Barney Lipscomb por su valiosa revision, acotaciones, paciencia y apoyo en esta

publication que contribuira al conotimiento de la flora salvadorena.

REFEREN Cl AS

a Doctorado en Ciencias (Sistemitica).

ssulaceae). Trab. Dept. Bot. Fisiol. Veg.

Carrillo, P.R. 2009. Estudios sistem^ticos en el dado Acre (Crassulac

Instituto de Ecologla, A.C. Veracruz, Mexico.

Castroviejo, S. y R. Calvo. 1981. Notas citotaxonomicas sobre Sedu

Madrid 11:49-57.

Marn (Minister© de Medio Ambiente y Recursos Naturales). 2012. (Pendiente de publicar). Mapa de Ecosistemas de El Sal-

vador. Actualization 2012 y Detection de Cambios. World Institute for Conservation and Environment (WICE). San

Salvador, El Salvador.

Moran, R. 1942. Delimitation of genera and subfamilies in the Crassulaceae. Desert PI. Life 14:125-128.

Mort, M. and S. Mori. 2004. Crassulaceae. In: N. Smith, S.A. Mori, A. Henderson, D.W. Stevenson, y S.V. Heal, eds. Flowering

otevens, k. mart, m. van ham, M. Elema, M. Wildeboer, and J. Zwaving. 1995. Distribution of alkaloids and tannins in the

Crassulaceae. Biochem. Syst. Ecol. 23:157-165.

't Hart ' h - and b - Bleu - 2003 - Sedum - En: U. Eggli, ed. Illustrated handbook of succulent plants: Crassulaceae. Springer,

Berlin, Germany. Pp. 235-332.

Thiede, J. and U. Eggli. 2007. Crassulaceae. In: K. Kubitzki, ed. The families and genera of vascular plants Vol. 9. Springer,

Thorne, R. and J. Reveal. 2007. An updated classification of the class Magnoliophyta ("Angiospermae”). Bot. Rev. (Lan-

caster) 73:67-181.

Uhl, C. 1 956. The Crassulaceae and cytotaxonomy. Cact. Succ. J. (Los Angeles) 48:225-229.

Uhl, C 1992. Polyploidy, disploidy, and chromosomes pairing in Echeveria (Crassulaceae) and its hybrids. Amer. J. Bot

79556-566.

pja de la UICN:

REDISCOVERY OF CALLIRHOE PAPAVER (MALVACEAE) IN ALABAMA (U.S.A.)

Brian R. Keener

Dept, of Biological & Environmental Sciences

The University of West Alabama

Livingston, Alabama 35470, U.S.A.

LJ. Davenport

Dept, of Biological & Environmental Science

Samford University

Birmingham, Alabama 35229, U.SA

ABSTRACT

RESUMEN

The woodland poppy-mallow, Callirhoe papaver (Cav.) A. G

eastern Coastal Plain from Georgia and Florida, west to easl

southwestern Georgia, northern Florida, Alabama, and Mis

of the Mississippi River (Dorr 1990). In Mississippi and Alabama, it has been specifically attributed to the Pine

Hills or Lower Pine Region in the southern portions of both states (Mohr 1901; Dorr 1990).

In a recent treatment of the Alabama vascular flora (Krai et al. 2011), Callirhoe papaver was treated as

historic” or not collected in over 100 years and was mistakenly omitted from the latest inventory of rare,

threatened, and endangered species (ALNHP 2012). The 2012 rediscovery of C. papaver in Washington Coun-

ty, Alabama, reported here, is thus significant.

HISTORICAL SPECIMENS

Charles Mohr (1824-1901), in his monumental Plant Life of Alabama (Mohr 1901), listed Callirhoe papaver only

from Healing Springs in northern Washington County. His citation “Herb. Geol. Surv. Herb. Mohr” was based

on a single collection (UNA 10851; Fig. 1) that he made in July, 1873. “Herb. Geol. Surv.” refers to the collection

that Mohr made for the Geological Survey of Alabama upon which his book was based (Davenport 1978,

1979a, 1979b). That collection of over 4000 specimens, long maintained separately as the herbarium of the Ala-

bama Museum of Natural History (ALU), is now incorporated into the University of Alabama Herbarium

(UNA).

The ALU collection was started in late 1878, when Eugene Allen Smith (1841-1927), long-time Alabama

State Geologist (see Henderson 2011), asked Mohr for help with his plant identifications. (Mohr’s polite accep-

tence (Mohr 1878] of Smith’s request is housed in the University of Alabama Special Collections.) Their col-

•aboration soon led to the privately published Preliminary List of the Plants Growing Without Cultivation in Ala-

bama (Mohr 1880). In that list, Callirhoe papaver was noted as occurring only in Washington County, with

Mohr ( M”) as its collector.

h» contrast, “Herb. Mohr” refers to Mohr’s much larger personal herbarium, which he bequeathed to the

mithsonian Institution (US) (Anonymous 1901). Two Alabama Callirhoe papaver specimens are currently

Und at US. The first (US 774668; Fig. 2) has two labels. The first, original label is affixed to the lower-left

836

Journal of the Botanical Research Institute of Texas 7(2)

i «».«»*>«. |

urn y S<t/£ ■

Upr '|

Hb , UNAOOOtqKi r]

comer; it has a printed “Carl. Mohr” with the locality data and several sets of notes by Mohr, made at different

times. The second label, placed in the traditional lower-right comer, is a newer one, copied by Mohr, with

“Woodensprings, Washington Co” and “Aug 1872.”

The above locality, subsequently recorded by Dorr (1990) as “Wooden Springs,” is misspelled. Mohr, who

sought the healing powers of mineral baths and more healthful climates throughout his adult life (Davenport

1978, 1979a), was most likely one of the first customers at a resort founded by William Wooten in 1872 (Sulzby

1960; Foscue 1989). That resort, built around 17 small springs along abranch of Santa Bogue Creek in northern

Washington County, Alabama, was perhaps initially known as “Wooten Springs” after the name of its devel-

oper, then changed to “Healing Springs” in order to attract more customers.

The second US specimen (US 11818; Fig. 3) lacks important details. The label is a generic one used at US

during the late 1800s and was probably placed ex post facto. Thus, the species’ identification, locality, and col-

lector’s name (“Mobile Ala; Wm Harvey”) are clearly in Mohr’s handwriting and not original notes by the col-

kctor. A significant omission is that no collection date is recorded on the label.

Who was “Wm Harvey”? An 1871 Mobile city directory lists a William Harvey as a route agent for the

Mobile & Ohio Railroad (Ancestry.com 2011). Because no such person is listed in either the 1870 or 1880

United States Censuses, Harvey must have only lived briefly in Mobile. According to a U.S. Commissioner of

Agriculture Affairs report (Watts 1875), that department received a “package of plants collected near Mobile,

Alabama, by Mr. William Harvey” during 1874. Most likely, that package contained Harvey’s Callirhoe papaver

specimen — perhaps collected at a stop along the Mobile & Ohio Railroad line, which ran within 13 miles of

Healing Springs.

Mohr probably didn’t know about Harvey’s collection at US until the 1890s. It was during that decade that

he worked diligently on his flora of Alabama, and he visited US on several occasions (Davenport 1978, 1979a).

during one of those visits, he must have been given Harvey’s specimen to identify. On the label, Mohr wrote

“Mobile” to indicate the collector’s location and not the actual location of the specimen. Most importantly, he

Journal of the Botanical Research Institute of Texas 7(2)

did not include Mobile County— which lacks suitable calcareous habitat-as a Callirhoe papaver locality in

Plant Life of Alabama (Mohr 1901). In his later monograph of the genus. Dorr (1990) attributed C. papaver to

Mobile County by citing the Harvey specimen without knowledge of the specimen’s history outlined above.

REDISCOVERY 1

During a 2012 botanical survey of the -Limestone" or -Jackson Prairie* region of southwestern Alabam

collection of CtlflWioe papaver was made by the first author in Washington County, The locality is ca. 4.2 m

NNE of HealmgAVoodenAVooten Springs, the only ‘original" Alabama locality. This population contaii

approximately 40 individuals.

, . \ lhe j’° V n loca * lt ^ Was rev isited one year later. Callirhoe papaver and associated species seemei

be slightly delayed in flowering time horn the previous year. Further exploration in 2013 (Keene. ' “

resulted in the discovery of an additional population of 10 individual

(Keener etal. 7344).

:s ENE of the 2012 locality

Voucher specimens: ALABAMA. Washington Co.: 3.8 air mi N of MiUr

B.R. Keener 7344 with W.K. Webb (UWAL, SAMF, TROY, VDB); 4.3 air m

Co. Rd. 45 (Mt. Carmel Rd.), 31.67720°N, 88.26003°W, 21 jun 2013 Br

(UWAL).

ry, prairie W of Brier Creek, 31.69025°N, 88.31505°W, 24 Jun 2012,

»i NE of Millry, along primitive private road ca. 1.6 mi E of jet. with

TOe sods of both sites are clay with occasional thin areas of exposed limestone. The O

full sunor Jong the margin of prairie woods dominated by eastern red-cedar Ouniperus

Associated herbaceous species are Asclepias viridis Walter, Echinacea purpurea (L.) Moe

Nutt., P. violacea Aubl., Silphium laevigatum Pursh, and Tripsacum dactyloides (L.) L. Unfo

dominated by the invasive cogon grass, hnperata cylindrica (L.) P. Beauv.

allirhoe plants grow in

s virginiana L.) (Fig. 'fl-

ench, Polygala boykinii

irtunately, the sites are

In light of Callirhoe papaver being treated as “historic” and its “disappearance” from the Alabama flora

since the early 1870s, the above collections are deemed noteworthy. While C. papa\er is seemingly a globally

secure taxon, it remains as one of Alabama’s rarest vascular plant species.

ACKNOWLEDGMENTS

We thank Deborah Bell (US) and Steve Ginzbarg (UNA) for providing images of herbarium specimens. We

greatly acknowledge the field assistance of Wayne Webb and Alvin Diamond. We thank John Hall for image

editing assistance. We also thank John Clark (UNA) and Larry Dorr (US) for insightful comments that im-

proved the manuscript.

REFERENCES

Alabama Natural History Program [ALNHP]. 201 2. Alabama inventory list; the rare, threatened, and endangered plants and

animals of Alabama. Privately printed. Auburn, Alabama, U.SA

Ancestry.com. 201 1 . U.S. City Directories, 1821-1 989. http://search.ancestry.com/cgi-bin/sse.dll?h=92601 2876&db=USD

irectories&indiv=try. Accessed 02 Sep 2012.

Davenport, LJ. 1979a. Charles Mohr and Plant Life of Alabama. Sida 8:1-13.

Davenport, LJ. 1979b. Vascular plant type specimens in the Mohr Herbarium, University, Alabama. Taxon 28:567-571 .

Dorr, U. 1990. A revision of the North American genus Callirhoe (Malvaceae). Mem. New York Bot. Gard. 56:1-76.

Foscue,V.O. 1989. Place names in Alabama. University of Alabama Press, Tuscaloosa, Alabama, U.SA.

Henderson, A.K. 2011. Eugene Allen Smith’s Alabama: How a geologist shaped a state. New South Books, Montgomery,

ftiaDama, U.5.A.

Kral, R., A.R. Diamond, Jr., S.L. Ginzbarg, CJ. Hansen, R.R. Haynes, B.R. Keener, M.G. Lelong, D.D. Spaulding, and M. Woods. 201 1 .

Annotated checklist of the vascular plants of Alabama. Sida, Botanical Miscellany 36. Botanical Research Institute of

Texas Press, Fort Worth, Texas, U.S.A.

Mohr, C.T. 1878. Letter to E.A. Smith from Mobile, AL 18 Dec University of Alabama Special Collections, Tuscaloosa,

Alabama, U.S.A.

Mohr, C.T. 1880. Preliminary list of the plants growing without cultivation in Alabama, from the collections made by

Eugene A. Smith, Tuscaloosa, and Charles Mohr, Mobile, Ala. Privately published, Mobile, Alabama, U.S.A.

Mohr, C.T. 1 901 . Plant life of Alabama. Contr. U.S. Natl. Herb. 6. Government Printing Office, Washington, DC; also Ala-

bama Edition, Geol. Survey of Alabama Monogr. 5. Brown Printing Co., Montgomery, Alabama, U.SA

Suiav, J.F., Jr. I960. Historic Alabama hotels and resorts. University of Alabama Press, Tuscaloosa, Alabama, U.SA

Watts, F. 1 875. Report of the Commissioner. In: Report of the Commissioner of Agriculture for the year 1874. US. Gov-

ernment Printing Office, Washington, DC, U.SA Pp. 5-14.

BOOK NOTICE

Kenneth D. Heil, Steve L. O’Kane, Jr., Linda Mary Reeves, and Arnold Clifford. 2013. Flora of the Four Cor-

ners Region: Vascular Plants of the San Juan River Drainage. Arizona, Colorado, New Mexico, and

Utah. (ISBN-13: 978-1-930723-84-9, hbk). Missouri Botanical Garden Press, P.O. Box 299, Saint Louis,

Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.info, orders@mbgpress.org, 1-877-271-1930).

$72.00, 1098 pp„ 8 Vi" x 11 W.

From the Foreword by Peter Raven, President of the Missouri Botanical Garden :

“Certainly, readers of Tony Hillerman s well-appreciated detective stories will appreciate the natural mag-

nificence and geological diversity of the Four Comers region even if they have not had the pleasure of visiting

the area personally. The botanical diversity of this region, so well described in this attractive and useful flora,

is no less impressive. Clearly, the use of this book will inspire further studies and thus make possible an ever

greater depth of knowledge about these plants and their position in the habitats where they occur.”

From the publisher:

“The Flora of the Four Comers describes all of those species, subspecies, and varieties of vascular plants

that grow spontaneously in the drainage basin of the San Juan River, a major tributary of the Colorado River.

This region takes in major portions of Arizona, Colorado, New Mexico and Utah, and is centered on the Four

Comers, the only spot in the United States where four states meet at a common point. The entire area encom-

passes 65,382 square kilometers (25,244 square miles), an area the size of West Virginia or Connecticut and

about half the size of Alabama, Arkansas, New York, or North Carolina. The highest point is 4292 m (14,083

feet) at Mt. Eolus in the San Juan Mountains and the lowest point is 1130 m (3708 feet) where the San Juan River

flows into Lake Powell (the Colorado River). Because of this elevation gradient, and the varying climates pro-

duced by local topography, vegetation in the study area varies from alpine tundra to coniferous forests, moun-

tain shrublands, lowland sagebrush, blackbrush, and to the sparse communities seen on bare rocks and the

scorching sides of low-elevation canyons.”

J. Bot Res. Inst Texas 7(2): 840. 2013

REDISCOVERY OF PERSEA BORBONIA VAR. BORBONIA (LAURACEAE),

PROSOPIS GLANDULOSA VAR. GLANDULOSA (FABACEAE), AND

PINUS PALUSTR1S (PINACEAE) IN ARKANSAS,

WITH THREE NEW ANGIOSPERM SPECIES FOR ARKANSAS (U.S.A.)

Brett E. Serviss James H. Peck

Department of Biology P.O. Box 705

Henderson State University Cedar Key, Florida 32625, U.S.A.

Arkadelphia, Arkansas 71999-0001, U.S.A. jhpeck@uair.edu

ABSTRACT

INTRODUCTION

Though vascular plant floristics in Arkansas is widely considered to have commenced with the 1819 explor-

atory travels of Thomas Nuttall, one of the first trained botanists to visit the area as documented by his 1835

collections publication, much additional floristic investigation has been conducted on the Arkansas flora over

the past century, including the publication of over 1,200 articles by nearly 100 botanists documenting and

explaining the diversity, abundance, and distribution of the state’s flora and vegetation (Peck & Peck 1988;

Peck et al. 2001; Peck 2003). Continued floristic work in Arkansas led to the establishment of the Arkansas

Vascular Flora Committee (AVFC) in 1998, tasked with the creation of a checklist, atlas, and identification

manual for the nearly 3,000 species of native and naturalized vascular plants present in the flora. The AVFC

published the Checklist of the Vascular Plants of Arkansas in 2006, with the Atlas of the Vascular Plants of Arkan-

sas set for publication in late 2013. Over the past decade, numerous species have been added to the state’s flora,

a number of which were non-native to the U.S. (Peck & Serviss 2006; Serviss & Peck 2008; Serviss 2009; Peck

2011a, 2011b; Peck & Serviss 2011; Serviss et al. 2012), and although recent work by the authors has focused on

exotic species in urban and semi-urban environments, field work in 2012 and 2013 led to the rediscovery of

two rare (in Arkansas), native, woody species of angiosperms, Persea borbonia var. borbonia and Prosopis glan-

Mosa var. glandulosa that have not been encountered in the Arkansas flora for well over 50 years, and one

gymnosperm species, Pinus palustris, which has not been documented in the state for 30 years. These rediscov-

11 htR «-lnfl. Texas 7{2):8

842

eries vibrantly illustrate the fact that in-depth and comprehensive floristic investigation must continue to oc-

cur in Arkansas.

REDISCOVERIES IN THE ARKANSAS FLORA

Persea borbonia (L.) Spreng. var. borbonia (Lauraceae), Red Bay. Persea borbonia var. borbonia is a shrub or

small to medium-sized tree that is native to the southeastern U.S. from eastern Texas and Louisiana to Florida,

Georgia, and North Carolina (Wofford 1997). The only previous record of P. borbonia var. borbonia in Arkansas

is from Miller County (F.L. Harvey s.n„ 18 Aug, no year present on voucher, UARK). The specific location data

of the Harvey R borbonia var. borbonia specimen is ambiguous; however, Tucker (1974) elaborated somewhat

on the location, indicating that it was collected “in the vicinity of Texarkana in swampy habitat.” Persea borbo-

nia var. borbonia has not been collected in Arkansas since 1881 (Harvey 1883; Tucker 1974; Smith 1988).

Voucher specimen: ARKANSAS. Union Co.: a few trees and many additional shrubs growing in thicket along shore of backwater area in

mucky, sandy soil. Beryl Anthony Lower Ouachita WMA, 26 Sep 2012, Peck 2012189 (HEND).

Prosopis glandulosa Torr. var. gtandnlosa (Fabaceae), Mesquite. Prosopis glandulosa var. glandulosa is a shrub

Serviss and Peck, Arkan

843

or small tree that is native and widespread throughout much of the southwestern U.S., southern and central

Great Plains, and Mexico (McGregor 1986; Isely 1998). Tucker (1976) reported this species from Pulaski

County in Arkansas based on collections by D.M. Moore in 1954 and 1955, noting that it was a “true inventive”

of potentially long duration that was collected along the railroad tracks on the southern edge of Little Rock,

apparently brought in with livestock. Smith (1978, 1988, 1994) excluded this species from the Arkansas flora,

considering it to be only a waif, and not “part of the normal flora”, but Peck (2003) reinstated P. glandulosa var.

glandulosa as a component of the Arkansas flora based on the Moore vouchers (Moore 54343, 55517, UARK);

however, P. glandulosa var. glandulosa has not been collected in Arkansas since 1955 (Peck 2003).

Voucher specimen: ARKANSAS. Little River Co.: one tree with flowers and six nonflowering shrubs growing in riverine lowland woods

along the Red River, next to oxbow lake with sandy soil, off of state hwy. 41, Hudson Lake area, 6 Apr 2013, Peck 2013001 (HEND).

Pinus palustris Mill. (Pinaceae), Longleaf Pine. Pinus palustris is a large tree that is native to the southeastern

U.S. from eastern Texas and Louisiana to Florida, the Carolinas, and Virginia (Krai 1993). Shepherd and Ama-

son documented this species from Arkansas in Union County in 1983 ( Shepherd and Amason 187, UARK). This

specimen was cited by Smith (1988), but he considered it as a possible “long-lived waif” and not truly a compo-

nent of the state’s flora. The 1983 Union County specimen of P. palustris is the only previous record of this

species from Arkansas (our record is a distinct plant from that of the Shepherd and Amason record).

Voucher specimen: ARKANSAS. Union Co.: single tree, possibly adventive or persisting from arboricultur

regrowth pine tree farm from former hardwoods, but now mixed loblolly pine stand, mucky, sandy soil, aloi

Junction City, 26 Jun 2013, Peck 2 013011 (HEND).

ADDITIONS TO THE ARKANSAS FLORA

haceae). Globe Amaranth. Gomphrena globosa is an annual species native to

Itivated for its colorful, long-lasting flowers. Gomphrena globosa is sporadi-

cally naturalized in several southeastern states, including Louisiana and Texas, along with several states in the

northeastern U.S. from Ohio and Virginia eastward to Massachusetts and New York (USDA, NRCS 2013).

Gomphrena globosa is a prolific, self-seeding species and should be expected in other locations in Arkansas,

especially in the vicinity of areas where plants of G. globosa are cultivated.

Voucher specimen: ARKANSAS. Pulaski Co.: several clusters or colonies of plants growing in sandy soil of the Arkansas River flood plain,

kittle Rock, Twin Rivers County Park, 28 Jul 2007, Peck 07-1673 (HEND).

Oxalis debilis Kunth (Oxalidaceae), Pink Woodsorrel. Oxalis debilis is native to tropical America, but is natu-

ralized in several southeastern states from Texas eastward to Florida and South Carolina (Nesom 2009; USDA,

NRCS 2013; Figs. 1, 2). In Arkansas, O. debilis is weedy, occurring and spreading rapidly in areas with high

disturbance, including flower beds, shrub plantings, lawns, and roadsides. Oxalis debilis is easily confused

with another introduced species of Oxalis, O. articulata Savigny (Windowbox Woodsorrel; Figs. 1, 2). Oxalis

articulata is native to Brazil and Argentina, and is naturalized in Arkansas and other areas in the southern U.S.

°xaHs articulata is documented in Arkansas from several counties, but based on its overall morphological re-

semblance and similarity to O. debilis, some specimens of 0. articulata could be misidentified and may be, in

feet, O. debilis. Oxalis debilis can readily be distinguished from both O. articulata and the native O. violacea L.

(Violet Woodsorrel; Figs. 1, 2) by the following key (modified from Wunderlin 1998; Horne et al. 2013).

Journal of the Botanical Research Institute of Texas 7(2)

Ruellia nudiflora (Engelm. and A. Gray) Urb. (Acanthaceae), Violet Wild Petunia. Ruellia nudiflora is native to

the southwestern U.S. (Arizona and Texas) and Mexico (Bailey & Bailey 1976). It has also been documented

from Louisiana and Mississippi, but it is unclear whether or not it is native or introduced in those states (USDA,

NRCS 2013). Ruellia nudiflora is probably not native in Arkansas, but rather an introduction via seed or plant

contaminants in horticultural materials (soil, mulch, or potted ornamentals). Ruellia nudiflora is a robust pe-

rennial that prolifically self-seeds; thus, accidental introduction via horticultural endeavors seems plausible.

ACKNOWLEDGMENTS

We very sincerely thank Brent Baker (Arkansas Natural Heritage Commission— ANHC) and Jennifer Ogle

(University of Arkansas Herbarium— UARK) for their invaluable assistance with occurrence data and distri-

butional status of several species, Ann Willyard (Hendrix College) and Brent Baker for reviewing this paper,

and the Henderson State University Biology Department for supporting this work.

REFERENCES

is Third. A concise dictionary of plants cultivated in the United States and Car

Bailey, LH. and E Z. Bailey. 1976. H

Vol. 2. MacMillan, New York.

Harvey, F.L. 1 883. The forest trees of Arkansas. Am. J. Forestry. 1 :41 3-424, 45 1 -458.

Horne, H.E., T.W. Barger, and G.L. Nesom. 201 3. Two South American species of Oxalis (Oxalidaceae) naturalized in Alabama

and the USA, first report. Phytoneuron 54:1 -7.

e and naturalized Leguminosae (Fabaceae) of the United States (exclusive of Alaska and Hawaii).

l, R. 1 993. Pinus. In: Flora of North America I

Vols. Oxford University Press, New York and Oxford. 2:373-398.

^Uwrenc V 986 M ' mOSaCeae ' ln: Fl ° ra ° fthe Great P,ains - Great p|ains Flora Association. University Press of Kansas,

Nesqm, G.L. 2009. Taxonomic notes on acaulescent Oxalis (Oxalidaceae) in the United States. Phytologia 91:501-526.

^ u !!!' K r i a ' a: addltlons ' reinstatements, exclusions, and re-exclusions. Sida 20:1 737-1 757.

J* . u J ! a ’ u eW and noteworth y additions to the Arkansas fern flora. Phytoneuron 201 1 -30:1 -33.

^ e ul°on -3str S PKrld0phy,e S,UdlM wlth a " ew a ""°«ted checklist and ftoristic analysis. Phyto-

Pm J H Z BE sZs's 2006 ^ZdZteZrth 30 ' °| fA,kanSaS b ° tany - Proc Arkansas tod Sci. 41:5^73.

f " ' TT ° terdCed States, with new and

noteworthy records of several angiosperms in Arkansas. J. Bot. Res. Inst Texas 5-321-326

^s tod™!^Zm FO ' 1 2001 Ad<anSaS fidd botany (flora and »9«atlon) bibliography (1988-2000). J. Arkan-

Serviss, B.E., D.H. Mason, and T.L. Bray. 201 2. A first SDontanenuc nimrd a *■ • j- , ,, ...

In the United States Beta. J. Bot Res. Inst Texas d6,7-«0 »■ defcioso (Actrnidraceae)

^ £“ 3 ^ and n ° teW ° fth> reC ° rdS ° f SeVera ' n0n - na,lva va * ula ' P |a "< s P«ies in Arkansas. 1

Smith, E.B. 1978. An atlas and annotated and list ofthe vascular flora of Arkansas fr c. Artan „ s

SmiulERIMS. An atlas and annotated and !ist of the vascular flora of Arkansas, second edition. E.B. Smith. Fayetteville.

Smhh, LB. 1994. Keys to the flora of Arkansas. University of Arkansas Press Fayetteville Arkan™

Serviss and Peck, Arkansas floristic discoveries

845

USDA, NRCS. 201 3. The PLANTS Database (http://plants.usda.gov). National Plant Data Team, Greensboro, NC 27401-

4901 USA. Accessed on 20 June 201 3.

Wofford, B.E. 1 997. Persea. In: Flora of North America Editorial Cc

16+ Vols. Oxford University Press, New York and Oxford. 3:34-

Wunderlin, R.P. 1 998. Guide to the vascular plants of Florida. Univ

imittee, eds. Flora of North America north of Mexico,

sity Press of Florida, Gainesville.

BOOK REVIEWS

Jon Farrar. 2011. Field Guide to Wildflowers of Nebraska and the Great Plains. (ISBN-13: 978-1-60938-

071-7, pbk). University of Iowa Press, 119 W. Park Road, 100 Kuhl House, Iowa City, Iowa 52242-1000,

U.S.A. (Orders: uiowapress.org, 1-800-621-2736). $39.95, 384 pp., 279 color photos, 2 color maps, 83

drawings, 6" x 9 1/8".

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car seats or in backpacks as wild-flower enthusiasts explore prairies, wildlands, and wetlands in search of

those ever-changing splashes of color we call wildflowers. This second edition is much like the beloved first

edition, but better.”— Jon Farrar

Flower lovers are not all professionally trained botanists. Therefore, identification by color has become the

more common practice if the pictures are included in various publications. This book has sections for Green

Flowers; White Flowers; Yellow Flowers; Orange to Red Flowers; Pink to Red-Violet Flowers; and Blue-Violet to

Blue Flowers. And, of course, it includes an illustrated glossary, references, and also additional reading

The photography in this volume is outstanding. Colors ring true, and the pictures are large enough to be

able to see easily and enjoy immensely. Additional line drawings of various plants and/or plant parts also show

up occasionally, carefully identified or explained Helen Jeude, Volunteer, Botanical Research Institute of Texas,

Fort Worth, Texas, U.SA.

Timothy P. Spira. 2011. Wildflowers & Plant Communities of the Southern Appalachian Mountains &

Piedmont: A Naturalist’s Guide to the Carolinas, Virginia, Tennessee, & Georgia. (Southern

Gateway Guides). (ISBN-13: 978-0-8078-7172-0, pbk). University of North Carolina Press, 116 South

Boundary Street, Chapel Hill, North Carolina 27514-3808, U.S.A. (Orders: uncpress.unc.edu, 1-800-

848-6224). $26.00, 540 pp., 361 color photos, 760 color thumbnails, 22 line drawings, 2 maps, bibl.,

index, 6" x 9".

Not only is Tim Spira an excellent botanist, author, and presenter, he has an amazing ability to bring the flora

alive and keep the reader totally engrossed for all 500+ pages. The illustrations are absolutely beautiful; the

descriptions are precise; and the information keeps the reader happily reading and anxious to get on the road

and go visit the Southern Appalachian Mountains and Piedmont.

This hefty volume is also easy to use. The author thoughtfully designed the volume to indicate the specific

pages for each part by identifying it with a specific color along the edge of the page. (The closed book will show

a specific color for each section.) In addition, instead of lumping descriptions of plants by color, a system fre-

quently used, Spira explains: “The key idea at the center of the book is the organization of plants into 21 major

communities. Often, you’ll see wildflower books organized by flower color or family affinity, but the natural

community emphasis used here is an exciting, fresh approach that is connected to our growing understanding

of the environment and how all parts of the natural world are mutually dependent. ”

This is an incredibly beautiful and fascinating book, and the reader will find him- or herself going back to

it repeatedly. Moreover, the book is of a quality that it should not be easily damaged if you have taken it into the

field and are using it carefully. The author has presented us with an exceptionally beautiful and helpful vol-

ume.— Helen Jeude, Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas, USA.

J. Bat Res. Inst. Texas 7(2): 846. 2013

A FLORISTIC INVENTORY OF PHILLIPS AND VALLEY COUNTIES,

MONTANA (U.S.A.)

Joseph L.M. Charboneau 1 , B.E. Nelson, and Ronald L. Hartman

Rocky Mountain Herbarium

Department of Botany

University of Wyoming

1000E. University Ave.

Laramie, Wyoming 82071-3165, US. A

ABSTRACT

first floristic inventory of Phillips and Valley counties on the glacia

718 species, and 358 genera from 86 families. Among these are 1(

RESUMEN

Este estudio marca el primer inventario florfstico de los condados de Phillips y Valley de las Uanuras glaciadas del noreste de Montana. El

frea de 23.191 km2 (8.954 sq mi) fue estudiada en busca de todos los taxa de plantas vasculares en los espacios manejados por el Bureau of

INTRODUCTION

We report on a vascular plant inventory of public and private lands in Phillips and Valley counties in northeast-

ern Montana (Fig. 1). The area is bound by Canada to the north, the Missouri River to the south, Daniels

County and Fort Peck Indian Reservation to the east, and Blaine County and Fort Belknap Indian Reservation

to the west. Elevation ranges from 616 to 1,743 m (2,020 to 5,720 ft).

The area is located within the North American Prairies floristic province near the edge of the Rocky

Mountain province (Takhtajan 1986), although Lavin and Seibert (2011) have suggested that the area has a

greater floristic affinity to the Intermountain region than to the Great Plains. Botanical exploration of the area

began in 1805 and 1806 when the Lewis and Clark Expedition traveled along the Missouri River (Phillips

2003). Past treatments that have covered the area include Rydberg (1932; peripherally), Atlas of the Flora of the

Great Plains (GPFA 1977), and Flora of the Great Plains (GPFA 1986). State floras include Vascular Plants of Man-

ama (Dorn 1984) and the recently published Manual of Montana Vascular Plants (Lesica 2012). The area is one

of many on the western Great Plains for which basic floristic knowledge has been lacking (GPFA 1986). Indeed,

the area was not previously well collected: fewer than 1,200 collections from this area larger than the State of

New Jersey are vouchered at the Montana State University Herbarium (MONT; 2013) and the University of

Montana Herbarium (MONTU; 2013).

848

Journal of the Botanical Research institute of Texas 7(2)

This botanical inventory is part of the larger effort by the Rocky Mountain Herbarium (RM) to map in

relatively fine detail the geographic distributions of species based on vouchered specimens and to produce a

flora of the greater Rocky Mountain region (Hartman 1992; Hartman & Nelson 2008; Hartman et al. 2009).

Thus, flonstic inventories (49 as master’s degree projects) have been conducted during the past 34 years in Ari-

zona, Colorado, Idaho, Kansas, Montana, Nebraska, New Mexico, Oregon, South Dakota, Utah, Washington,

and Wyoming (e.g. Reif et al. 2009; Kesonie & Hartman 2011; Kuhn et al. 2011; Lukas et al. 2012). Over

650,000 new collections have been obtained by graduate students, staff, and research associates of RM. These

specimens form the core of the RM Plant Specimen Database (835,000 specimen records, 55,000 specimen im-

ages, and 4,000 field images; Hartman et al. 2009).

Study area. Various federal and state agencies manage lands in the area. In Phillips County, 4,374 sqkm

(1,689 sq mi) of Bureau of Land Management (BLM) lands are managed by the Malta BLM Field Office or in the

southwest comer of the county as part of the Upper Missouri River Breaks National Monument, which is ad-

ministered directly by the Montana/Dakotas BLM. The Glasgow BLM Field Office manages 4,095 sq km (1,581

sq mi) in Valley County. Also covered were 1,563 sqkm (603sqmi) ofU.S. Fish and Wildlife Service lands in-

cluding Charles M. Russell National Wildlife Refuge north of the Missouri River as well as Bowdoin National

Wildlife Refuge. The area also includes 1,635 sq km (631 sq mi) managed by the state, mostly as Montana State

Trust Lands or by Montana Fish, Wildlife, and Parks. Private lands visited include the American Prairie Re-

serve (133 sq km/51 sq mi) in southern Phillips County and the Matador Ranch (123 sq km/49 sq mi), owned

and operated by The Nature Conservancy in southwestern Phillips County. In total, 1 1 ,924 sq km (4,604 sq mi)

849

were accessible for collection within the 23,191 sq km (8,954 sq mi) area (the entirety of Phillips and Valley

counties exclusive of lands on the Fort Peck and Fort Belknap Indian Reservations). There are four wilderness

study areas (WSAs) managed by the BLM: Bitter Creek WSA (239 sq km/92 sq mi) in northern Valley County,

Burnt Lodge WSA (56 sq km/21 sq mi) in the Larb Hills (South), and Antelope Creek WSA (50 sq km/19 sq mi)

as well as part of Cow Creek WSA (138 sq km/53 sq mi in total) in the Upper Missouri River Breaks National

Monument. Grasslands National Park of Canada is located just north of the area in Saskatchewan.

Physiography.— The area is located on the Glaciated Missouri Plateau subregion of the northwestern por-

tion of the Great Plains physiographic region (Fenneman 1916). Figure 1 shows topographic features and bod-

ies of water in the area. The vast majority of the area was glaciated during the Pleistocene (Colton et al. 1961;

Fullerton & Colton 1986). Most of the area lies on broadly rolling hills with typically dry drainages, locally

called coulees. Grasses dominate these rolling hills with sagebrush ( Artemisia spp.) abundant in some areas as

well. Topographic relief is greater in the south on the Missouri River Breaks, where steep slopes can be covered

with ponderosa pine woodlands. The Little Rocky Mountains, one of several forested island mountain ranges

in central Montana, rise about 610 m (2,000 ft) above the surrounding plains in southwestern Phillips County

and southeastern Blaine County. The summit of Antoine Butte at 1,743 m (5,720 ft) is the highest point in the

Little Rockies and the area.

The entire area is located within the Missouri River watershed. Most of the area drains into the Milk River

except several drainages in the south that lead directly to the Missouri River and part of northeastern Valley

County, which is in the Poplar River watershed. The Milk River nearly bisects the area, entering in the west

near Dodson and reaching its confluence with the Missouri River in the east (Fig. 1). The Missouri River is

dammed near the town of Fort Peck by Fort Peck Dam, which was constructed by the U.S. Army Corps of En-

gineers during the 1930s (Bandy et al. 2004). Fort Peck Lake forms the shoreline of the Missouri River for much

of its length within the area.

Climate . — The region has a cold semi-arid climate (BSk in the Koppen-Geiger climate classification; Peel

et al. 2007), characterized by warm to hot summers and long cold winters (Bingham et aL 1984; Bandy et al.

2004; NCDC 2012). Average daily maximum temperatures range from 9.8 to 15.7°C (49.7 to 60.2°F), with the

north cooler than the south (PRISM 2004). Average daily minimum temperatures range from -3.3 to 2.3°C

(26.1 to 36.1°F), again generally lower in the north than in the south (PRISM 2004). Average annual precipita-

tion is relatively low, ranging from 26.7 to 55.1 cm (10.5 to 21.7 in) in the Little Rocky Mountains (PRISM

2004). Areas of locally high elevations tend to receive more precipitation, including the Little Rockies. About

half of the annual precipitation falls in the months of May, June, and July (NCDC 2012; WRCC 2012). Severe

thunderstorms throughout the summer can bring locally heavy precipitation as well as damaging winds and

hail (Bingham et al. 1984).

Precipitation was well above normal throughout most of the area in both field seasons of this inventory

(2010 and 2011). Annual precipitation in 2010 at Glasgow was 46.0 cm (18.1 in; 156 percent of average) and in

2011 was 58.4 cm (23.0 in; 198 percent of average), the highest ever recorded in Glasgow (NCDC 2012; NWS

2012). In addition, the 275.8 cm (108.6 in) of snow that fell in Glasgow during the winter of 2010 and 201 1 were

the most ever recorded, more than three times greater than the average of 91 cm (36 in; NWS 2012). This ab-

normally high level of precipitation created excellent conditions for conducting a floristic inventory but

brought extensive flooding as well.

Geology . — Three main events define the surficial geology of the area: the deposition of sedimentary rocks

m a shallow inland sea during the Late Cretaceous, the formation of the Little Rocky Mountains during the

early Paleogene, and the glaciation of nearly the entire area during the Pleistocene.

Throughout most of the area, the geologic layers exposed at the surface were deposited during the Late

Cretaceous when a large, shallow, inland sea known as the Western Interior Seaway covered the region (Mar-

shak 2005). Formations exposed from this time period are, from oldest to youngest, the Claggett shale, the

Judith River formation, the Bearpaw shale, the Fox Hills sandstone, and the Hell Creek formation (Collier 1918;

Vuke et al. 2007). The most commonly exposed of these Cretaceous-age materials is the Bearpaw shale (Vuke

850

Journal of the Botanical Research Institute of Texas 7(2)

et al. 2007). It consists of mostly dark-gray shale of marine origin and in some areas forms badlands and sticky

clay soils known locally as gumbo (Collier 1918; Jensen & Varnes 1964). Localized bentonite layers in the

Bearpaw shale, derived from volcanic ash deposits, have been mined in the area (Jensen & Varnes 1964; Bandy

et al. 2004).

A structure called the Bowdoin dome exists in the central and northern portion of the area, centered

about Nelson Reservoir and Lake Bowdoin (Bandy et al. 2004). Strata dip very slightly away from the center of

sures of two older formations, the Claggett shale and the Judith River formation (Collier 1918; Vuke et al.

2007). The older Claggett shale, which outcrops at the center of the dome, consists of a dark-gray marine shale

similar to the Bearpaw shale. The Judith River formation, which outcrops on the periphery of the dome, con-

sists of sandstones and shale of a freshwater depositional environment (Collier 1918; Jensen & Varnes 1964).

The Bowdoin dome has trapped natural gas in underlying Colorado Group sandstones (Bandy et al. 2004).

Natural gas production from this dome has occurred since the early part of the 20th century and continues

today (Bandy etal. 2004).

The Fox Hills sandstone and Hell Creek formation (famous for its dinosaur fossils; Jensen & Varnes 1964)

outcrop in the southern part of the area as well as parts of northeastern Valley County (Collier 1918; Vuke et al.

2007). These consist of mostly sandstones (Bandy et al. 2004). The sandstones of the Hell Creek formation are

more erosion resistant than the surrounding Bearpaw shale and often cap hills, particularly in the southern

part of the area (Jensen & Varnes 1964).

The Flaxville gravel, derived from alluvial terrace deposits from the late Neogene and early Quaternary, is

exposed in small parts of the north (Bandy et al. 2004). Resistant to erosion, it caps uplands and benches where

it is exposed (Collier 1918). Alluvium from the Quaternary is present in the Milk River Valley and lower parts

of larger creeks as well as on the Missouri River upstream of Fort Peck Lake (Bandy et al. 2004; Vuke et al.

2007).

The Little Rocky Mountains were formed during an early Paleogene orogeny in which intrusive igneous

rocks uplifted Precambrian basement rocks and overlying Paleozoic and Mesozoic sedimentary rocks around

the periphery of the range (Knechtel 1959). Precambrian metasedimentary and metavolcanic rocks outcrop

along with igneous rocks in the center of the Uttle Rockies (Knechtel 1959; Bandy et al. 2004; Vuke et al. 2007).

These igneous rocks at the core were intruded about 60 million years ago from alkaic magma (Wilson & Kyser

1988; Bandy et al. 2004). Gold and silver have been mined in the Litde Rockies since 1884 in a variety of opera-

tions (Wilson & Kyser 1988; Bandy et al. 2004).

The sedimentary rocks overlying the Little Rocky Mountains were uplifted during the orogeny and sub-

sequently have been eroded away over the core of the range, while remaining at the periphery (Knechtel 1959;

Vuke et al. 2007). The most prominent strata exposed at the surface are erosion-resistant calcareous rocks from

the Paleozoic, including dolomites of the Bighorn formation from the Ordovician, the Jefferson limestone of

the Devonian, and especially the Lodgepole and Mission Canyon limestones of the Mississippian (Knechtel

1959). Mesozoic rocks outcrop mostly in the foothills surrounding the Little Rockies and in small areas within

the range. These are mostly shales but also include some sandstones, conglomerates, and limestones (Knechtel

1959). Rocks from the Jurassic and Early Cretaceous are exposed in small areas around the periphery of the

range but once on the plains, strata from the Upper Cretaceous dominate at the surface (Knechtel 1959; Vuke

etal. 2007).

The Laurentide Ice Sheet covered the entire region during the late Illinoian glacial period (between

195,000 and 128,000 years ago) with the exception of the Little Rocky Mountains and an area east of Opheim

within the Poplar River drainage (Colton et al. 1961; Fullerton & Colton 1986). Following this glacial period,

extensive badlands formed subsequent to glaciation in the Wisconsinan (Fullerton & Colton 1986). Glaciers

returned between 21,000 to 16,000 years ago during the late Wisconsinan, although to a much smaller extent

than during the Illinoian (Fullerton & Colton 1986). During this time large areas remained ice-free in south-

i Phillips Coi

nuch of Valley Corn

Charboneau et al., Flora of Phillips and Valley counties, Montan

eluding the central portion (Colton et al. 1961; Fullerton & Colton 1986). Prior to these glacial episodes, the

Missouri River formed the broad valley that the Milk River now meanders through (Collier 1918; Bingham et

al. 1984; Bandy et al. 2004). Blocked by glacial ice, the Missouri River became entrenched in its current channel

during the Wisconsinan (Collier 1918; Alden 1932).

Paleovegetation . — Vegetational history following deglaciation is somewhat uncertain because of a pauci-

ty of fossil pollen data from northern Montana (Bamosky 1989; Strong & Hills 2005). However, it is likely that

after 12,000 years ago extensive grasslands similar to the present vegetation were established in the region,

unlike areas further east and north, which supported long-standing wide bands of boreal forest following de-

glaciation (Strong & Hills 2005). Fossil pollen data from Guardipee Lake, Montana indicates that by 12,200

years ago, temperate grasslands with shrubs in mesic habitats were present in northern Montana east of the

Rocky Mountains (Barnosky 1989). After 9,300 years ago these grasslands started to become more xeric as

they are today (Bamosky 1989).

Less clear is the nature of the vegetation following the maximum extent of the Laurentide Ice Sheet about

20,000 years ago (Fullerton & Colton 1986) but prior to 12,000 years ago. There is no direct evidence for for-

ests during this time, although the area may have been near the edges of both cordilleran and boreal forest

belts. A dry deciduous boreal forest or aspen parkland may have existed south of the boreal/cordilleran forest

zone in southern Saskatchewan (Klassen 1994), perhaps approaching northern Montana. The existence of a

belt of cordilleran forests during this time may explain the distribution of these tree species in theisland

mountain ranges of central Montana and the Cypress Hills in Canada (Thompson & Kuijt 1976; Strong&Htlls

2005). Presumably such a cordilleran forest belt stretched across the lowlands but was isolated after 14,000

years ago onto the discontinuous highlands of the region (Strong & Hills 2005), including the Little Rocky

Mountains. Thompson and Kuijt (1976) believed this a more plausible explr

dilleran conifers in the Cypress and Sweetgrass hills than long distal

Soils and Agriculture .— Substrates are important in detern

(Kruckeberg 2002), and in most of the area, soils rather than unw<

Many soils have developed from tills left following Illinoian and W

material is typically not far removed from its original source as the area was at the southern limit ot the c<

tal (Bandy et al. 2004). Therefore, these tills are derived pm

for the distribution of cor-

ispersal of seeds by wind or birds,

lg the distribution of plant species

red rocks are present at the surface,

nsinan glaciations. However, this till

nental ice sheet and scouring power was i

ily from Cretaceous shales. Tills are thickest in the

been removed completely by ere

erratics have been deposited fre

iart of the ai

places (Bingham etal. 1984; Bandy etal. 2004). A few large glacial

the Hudson Bay (Collier 1918; Bandy et al. 2004).

Is have also affected human settlemer

developed ft

scan be highly al

This alkalinity

Lh relatively low

^ ui ujc ^ i(j i i 0 f t he land unsuited for cultivation (Cooper et al. 2001). Many home-

st^err'wlwTlartedto'ar'rive following the establishment of the Great Northern Railway in 1887 (Bandy et al.

2004; now operated by the BNSF Railway), saw theit farms go bankrupt during the Great Depression <B,ng-

ham et al. 1984). The BLM now manages many of these lands that were repurchasedby the federal government

under the Bankhead-Jones Farm Tenant Act of 1937 (Mackie 1970; Cooper e, aL 2001k Today, most of the area

is utilized for cattle grazing, and toalesser extent, sheep grazing (Bandy etal. 2004).D^amifarmmgofsmaU

grains including spring wheat barley, and oats, as well as irrigated farming along the Milk River are suU im-

portant as welKBingham et al. 1984; Bandy et al. 2004). Today about 17 percent of the area is under culuvanon

(MTNHP 2010). The unsuitability of most of the area for cultivated agriculture and itsuse primarily as range-

land have left many of the grasslands and shrublands relatively intact (Cooper et aL 2001).

METHODS

The methodsusedfor this inventory largely follow those employed by other gmtotestudenri and srfatRM

for other floristic inventories in the greater Rocky Mountain region (Hartman 1992; Hartman & Nelson 2008,

Reif et al 2009- Kesonie Sr Hartman 2011; Kuhn et al. 2011; Lukas e. aL 2012). Our primary objective was to

852

document the diversity of vascular plants across the area throughout the growing season through the collec-

tion of voucher specimens. As such, we chose individual collecting sites in the field rather than visiting a set of

randomly distributed points. Collecting sites were selected for greatest potential diversity, often at the intersec-

tion of different vegetation types or on unique substrates, while spacing sites over the region during different

months of the field season. At each collection site, we used the “meander” search strategy (Goff et al. 1982;

Hartman 1992; Hartman & Nelson 2008). All species in flower or fruit or otherwise readily identifiable

through vegetative characters were vouchered at each site visited and relevant habitat and location data (in-

cluding GPS coordinates) were recorded. Specimens were collected within about 0.8 km (0.5 mi) of each re-

corded GPS point. Voucher specimens were collected, pressed, and dried in accordance with standard collect-

ing techniques described in Hartman (1992) and Hartman and Nelson (2008).

Joseph L.M. Charboneau and B.E. Nelson made collections in the field seasons of 2010 and 2011. In 2010,

we spent 53 person-days collecting between 8 June and 25 August and between 10 September and 21 Septem-

ber, generally alternating days collecting with days spent pressing. In 2011 , between 10 May and 15 August, we

spent 49 person-days collecting. In total, we made 12,768 collections from 308 sites at a density of 0.55 collec-

tions per sq km (1.43 per sq mi). Figure 2 contains a map of collection sites.

Specimens were identified using a number of floras including Dorn’s Vascular Plants of Montana (1984),

Flora of the Great Plains (GPFA 1986), Dorn’s Vascular Plants of Wyoming (2001), and Flora of North America

(1993+). All identifications were checked against specimens in RM verified by specialists. Nomenclature fol-

lows that of the RM Plant Specimen Database (Hartman et al. 2009). Specimen data have been entered into this

database and are available online (Hartman et al. 2009). All specimens are housed at RM, and duplicates have

been sent to MONT, MONTU, and other herbaria. We searched all databased records at MONT, MONTU, and

RM/USFS (USFS is the National Herbarium of the U.S. Forest Service, integrated with RM; Hartman et al.

2009; MONT 2013; MONTU 2013) from the area for taxa we did not collect as part of this study but were col-

lected by others and personally verified the identification of these specimens. These “historical” taxa are in-

cluded within the annotated checklist.

We described 19 vegetation types organized into six physiognomic categories based on the dominant

vegetation of each type, taking inspiration from the Montana Ecological Systems Field Guide (MTNHP 2012a).

These descriptions are based on our field notes, and the species listed in our vegetation type descriptions were

the most commonly collected within each type.

We performed two types of analyses to assess the adequacy of our collecting in documenting the actual

diversity of vascular plants. The first was a comparison of the environmental conditions and cover types sam-

pled by our collection sites and a set of randomly placed points based on the non-stratified environmental pa-

rameter analysis described by Neldner et al. (1995). Using ArcGIS v. 10.0 (ESRI 2011) we classified ranges of

three environmental variables across the area: elevation (USGS 2009), average annual precipitation, and aver-

age daily minimum temperature (PRISM 2004). We then created a raster file with combinations of these

classes and determined how many combinations were sampled by our collection sites and a set of random

points within the same accessible lands we collected. We also repeated this analysis using land cover type data

from MTNHP (2010) in place of the environmental data.

The second type of analysis used to evaluate our sampling adequacy was a comparison of the vascular

plant diversity we observed to estimates of the true diversity present. We used Estimates v. 9.1 (Colwell 2013)

to make taxon accumulation curves by collection days elapsed both chronologically and from 100 randomiza-

tions of collecting order using the default settings. For this purpose we used all collections that were defini-

tively identified even if they were eventually discarded for inadequate material. We estimated the total vascular

plant diversity using both the non-^arametric, asymptote-fitting Michaelis-Menten equation and parametric

richness estimators (i.e. based on the number of taxa collected only once or twice) such as the bootstrap, sec-

ond-order jackknife, and Chao 1 estimators (see Colwell & Coddington 1994 for a review of these methods).

We compared these estimates of actual taxon diversity to the number of observed taxa to estimate the percent-

age of actual taxon diversity documented.

RESULTS AND DISCUSSION

conservation concern, exotic taxa and noxious weeds, newly documented taxa, and sampling adequacy.

Summary of Taxa

We collected 762 unique taxa from 86 vascular plant families. The families with the highest diversity are

Asteraceae (134 taxa), Poaceae (111), Fabaceae (55), Brassicaceae (39), and Rosaceae (37). Genera with the

greatest number of taxa observed are Carex (Cyperaceae; 21 taxa), Astragalus (Fabaceae; 19), Elymus (Poaceae;

18), Poa (Poaceae; 11), and Potentilla (Rosaceae; 11). Below is a summary of the plants collected during the

study. Seventy “historical” taxa housed at MONT, MONTU, or RM/USFS were located from an additional four

families, 22 genera, and 68 species, bringing the total number of unique taxa to 832. Numbers in parentheses

below are totals including taxa collected by other workers.

c category:Taxa by special

108 (133)

14.2 (16.0)

(12)

(20)

854

Taxa by major plant group:

Fern Allies 7 (7)

Gymnosperms 7 (7)

Angiosperms 741 (810)

VEGETATION TYPES

Mackie (1970), Roberts (1980), Hansen et al. (1995), and Cooper et al. (2001) are among the few descriptions of

plant communities specific to the region. Using data taken from field notes, we describe 19 vegetation types

organized into six physiognomic categories based on the dominant vegetation. Delimitation of vegetation

types across the landscape is sometimes difficult as boundaries are often not clear-cut. The types we present

are not meant to be completely distinct and often blend into one another. Abbreviations for vegetation types

consist of an initial uppercase letter designating the physiognomic category followed by two lowercase letters

for the unique vegetation type. If only one infraspecific taxon was found for a species, only the species name is

listed in the vegetation type description.

Grasslands (G)

). — Mixedgrass prairie is the most common vegetation type, dominating over much

of the rolling plains. Although some sources classify the grasslands of eastern Montana as shortgrass prairie

(e.g. GPFA 1986), they are better classified as northern mixedgrass prairie (Coupland 1961; Singh et al. 1983).

Cool-season (C3) grasses dominate this mixedgrass prairie with a single short, warm-season (Q grass (Bou-

teloua gracilis ) present to varying degrees (Singh et al. 1983). Cool-season grasses dominant in mixedgrass

prairie are Elymus smithii, Hesperostipa comata, Koeleria macrantha, Nassella viridula, and Poa secunda subspe-

cies as well as the sedge Carex Jilijolia.

In addition to the dominance of grasses, Selaginella densa (spikemoss) can sometimes form a significant

component of these grasslands. Shrub cover can range from low to moderate as mixedgrass prairie blends into

sagebrush steppe. Shrubs commonly found are Artemisia cana, A. tridentata, Juniperus horizontalis, and Kra-

scheninnikovia lanata along with the cactus Opuntia polyacantha and the subshrub A.frigida. Forb diversity is

relatively high in mixedgrass prairie. Achillea millefolium, Allium textile, Antennaria spp., Astragalus spp., Boech-

era collinsii, Collomia linearis, Erigeronpumilus, Erysimum inconspicuum, Hedeoma hispidum. Heterotheca villosa,

Hymenoxys richardsonii, Lomatium foeniculaceum, Oenothera suffrutescens, Packera cana, Pediomelum argophyl-

lum, Penstemon spp.. Phlox hoodii, Plantago patagonica, Ratibida columnifera, Sphaeralcea coccinea, Vida ameri-

cana var. minor, and the exotic Tragopogon dubius are commonly found.

The area’s flora is more greatly influenced by regions to the west rather than by the eastern edge of the

Great Plains (Lavin & Seibert 2011). Grasses of the tallgrass or “true” prairie such as Andropogon gerardii Vit-

man, Hesperostipa spartea (Trin.) Barkworth, Panicum virgatum L., Sorghastrum nutans (L.) Nash, and Spo-

robolus heterolepis (A. Gray) A. Gray (Johnson & Larson 2007) indeed are entirely absent. But to say that the

area is little influenced by the Great Plains flora as indicated by Lavin and Seibert (2011) is dependent on how

one defines this flora. The Great Plains flora is in all parts recent and adventive, with species colonizing from

peripheral ecosystems (GPFA 1986).

A variant of mixedgrass prairie occurs in the north where mesic grasslands on soils derived from fine-

grained till are dominated by Hesperostipa curtiseta and Elymus lanceolatus varieties (Coupland 1961; Cooper

et al. 2001). This association will be discussed further with the moist coulee bottom and swale vegetation type.

Upland prairie (Gup).— Well-drained prairie uplands often have a distinctive suite of species in addition

to those common on typical mixedgrass prairie. Sandstone outcrops and sandstone-derived soils are often

present on uplands since sandstone erodes less easily than shale in this semiarid environment (Jensen & Var-

nes 1964). Thus many uplands often have sandier soil than surrounding areas. On these uplands, forbs such as

Astragalus gilviflorus, Comandra umbellata, Cryptantha celosioides, C. spiculifera, Dalea Candida, Eriogonum spp-.

Heterotheca villosa, Hymenopappus filifolius, Hymenoxys richardsonii, Lithospermum incisum, Lupinus pusilhs ,

Charboneau et al.. Flora of Phillips and Valley counties, Montana 855

Oenothera suffrutescens, Oxytropis sericea varieties. Paronychia sessiliflora, Penstemon nitidus, Physaria spatula-

ta, Stenotus armerioides, Tetraneuris acaulis, and Xanthisma grindelioides are common. Typical shrubs include

Juniperus horizontal is, Kraschenirmikovia lanata, Rhus trilobata. Yucca glauca, and the subshrub Artemisia camp-

estris var. pacifica. Graminoids often growing in this habitat are Achnatherum hymenoides , Bouteloua gracilis,

Calamovilfa longifolia, Carex filifolia, Elymus spicatus, Hesperostipa comata, and Schizachyrium scoparium.

Montane meadows (Gmm).— There are only a few montane meadows found on south exposures in the

Little Rocky Mountains. These often have many grassland species found at lower elevations but also have a

distinctive assemblage of forbs. Diagnostic forbs include Balsamorhiza sagittata. Delphinium bicolor, Drymocal-

lis glabrata, Lithospermum ruderale, Oxytropis splendens, and Solidago mollis. Some diagnostic graminoids are

Bromus porteri, Calamagrostis purpurascens, Carex hoodii, C. rossii, Festuca saximontana, and the exotic Poa

pratensis. The shrub Dasiphorajruticosa can also be found in these open meadows.

Shrublands (S)

Sagebrush steppe (Sss). — Sagebrush steppe intergrades extensively with mixedgrass prairie, sharing many of

the same graminoid and forb species. It is most prevalent in the southern part of the area. Sagebrush ( Artemisia

spp.) cover is dependent in part on climatic and edaphic factors, with areas receiving a greater proportion of

winter precipitation and greater soil moisture at depth likely to have higher sagebrush cover than pure grass-

lands (Knight 1994). Fires also greatly influence sagebrush cover. It may take more than 100 years for Wyo-

ming big sagebrush cover to return to pre-burn levels in eastern Montana sagebrush steppe (Cooper et al.

2011).

There are two primary sagebrush taxa forming sagebrush steppe: Artemisia tridentata var. w yomingensis

(Wyoming big sagebrush) and A. cana var. cana (silver sagebrush). Artemisia tridentata is typically is found on

never encountered it north of the Milk River. Artemisia cana is found throughout the area and is more tolerant

of higher soil moisture than A. tridentata (Knight 1994) and as such can often form sagebrush steppe in moist

coulees. Artemisia cana is also found in sandier soil than A. tridentata and is able to resprout after fires and

other disturbances unlike A. tridentata (Knight 1994).

shrubs Artemisiafrigida, Atriplex gardneri var. gardneri, Gutierreziasarothrae, and the cactus Opuntiapolyacan-

tha. Typical graminoids are Bouteloua gracilis, Elymus elymoides varieties, E. smithii, Hesperostipa comata,

Koeleria macrantha, Nassella viridula, Poa secunda subspecies, and the exotic grass Bromus japonicus. Forbs

commonly found in sagebrush steppe include Achillea millefolium. Allium textile, Antermaria parvifolia. Astraga-

lus missouriensis, Atriplex argentea, Dalea purpurea, Erigeron pumilus, Grindelia squarrosa, Heterotheca villosa,

M usineon divaricatum, Orobanche fasciculata, Pediomelum argophyllum, Plantago patagonica, Ratibida colum-

nifera, Senecio integerrimus var. scribneri, Vida americana var. minor, and the exotic Tragopogon dubius. As in

mixedgrass prairie, Selaginella densa can form significant ground cover as well.

Juniper steppe/woodland (Sjw).— This vegetation type is transitional between sagebrush steppe and pon-

derosa pine-juniper woodland, overlapping both considerably. It is found only in the south along the Missouri

River Breaks where Juniperus scopulorum (Rocky Mountain juniper) J. horizontalis (creeping juniper), and their

conspecific hybrid ,J. xfassettii, occur relatively sparsely on hillsides and coulees . Juniperusxfassettii (also

known as J. scopulorum Sarg. var. patens Fassett) is a decumbent shrub intermediate in stature between the

parental species that lacks the single-stemmed crown of J. scopulorum and the completely prostrate habit of J.

horizontalis (Adams 2011). Other common shrubs include Artemisia tridentata and Rhus trilobata.

Greasewood shrubland (Sgs).— Shrublands dominated by Sarcobatus vermiculatus (greasewood) are of-

ten found toward the bottom of coulees on soils derived from marine shales where there are saline soils and a

high water table (MTNHP 2012a). Artemisia tridentata is another common shrub in the fine-textured soils of

this vegetation type along with subshrubs Atriplex gardneri var. gardneri, Gutierrezia sarothrae, Suaeda calceo-

liformis, and the cactus Opuntia polyacantha. The forbs Atriplex suckleyi, Dieteria canescens, Grindelia squarro-

sa, Helianthus annuus ha axillaris, Musineon divaricatum, Plantago elongata, Sphaeralcea coccinea, Vida ameri-

856

Journal of the Botanical Research Institute of Texas 7(2)

cana var. minor are typically found along with exotics Melilotus officinalis. Polygonum aviculare, and Tragopogon

dubius. Common grasses include Bouteloua gracilis, Distichlis spicata, Elymus elymoides var. elymoides, E.

smithi i, Hordeumjubatum ssp. intermedium, and the exotic grass Bromusjaponicus. Sagebrush steppe and juni-

per steppe/woodland often intergrade into these greasewood shrublands from upslope.

Forests and Woodlands (F)

Thicket and woody draw (Ftw ). — In steep coulees there is enough moisture to support thickets primarily of

shrubs, especially Prunus virginiana, Rhus trilobata, and Shepherdia argentea but also Amelanchier alnifolia,

Comus sericea, Juniperus spp., Ribes spp., Rosa woodsii, Symphoricarpos occidentalis, and Toxicodendron

rydbergii. In the steepest, moistest coulees, trees such as Acer negundo var. interius, Fraxinus pensylvanica,

Juniperus scopulorum, and Populus deltoides can be found. Typical grasses in these thickets are Elymus ca-

nadensis, E. trachycaulus var. trachycaulus, Nassella viridula, Piptatherum micranthum, and exotics Bromus

inermis and Poa pratensis. Forbs such as Astragalus agrestis, Campanula rotundifolia, Geum triflorum, Glyc-

are often found along with exotics Camelina microcarpa and Fallopia convolvulus. Wooded draws with Fraxi-

nus pensylvanica are presently experiencing reduced seedhng recruitment and have been declining in quality

across eastern Montana due to the effects of overgrazing and the invasion of exotic grasses such as Bromus

inermis and Poa pratensis (Lesica & Marlow 2013).

ed by Populus deltoides (cottonwood) are found along the flood plains of the Milk and Missouri rivers and a few

larger creeks. Other trees sometimes found in these riparian forests are Acer negundo var. interius, Fraxinus

pensylvanica, and Salix amygdaloides, along with the exotic tree Elaeagnus angustifolia. Typical shrubs are

Prunus virginiana, Rosa woodsii, Salix eriocephala var. famelica, S. exigua ssp. interior, Symphoricarpos occidenta-

lis, and the subshrub Artemisia dracunculus. Fluctuating water levels and livestock disturb these forests so

weedy grasses such as exotics Bromus inermis, Eragrostis cilianensis, Setaria viridis, and natives Echinochloa

muricata and Panicum capillare are often found along with weedy forbs including exotics Euphorbia esula variet-

ies and Kochia scoparia. Also commonly found are Artemisia ludoviciana, Chamaesyce glyptosperma, Glycyr-

rhiza lepidota, and Solidago gigantea. In many of these forests, human alteration of hydrology has resulted in

highly altered, old cottonwood stands with limited regeneration since high water events are necessary for the

recruitment of new seedlings (Auble & Scott 1998). Flooding during 2011, however, resulted in the establish-

ment of many new cottonwood seedlings on the banks of the Milk and Missouri rivers.

on steep drainages. The upper canopy is typically fairly open and composed of Pinus ponderosa (ponderosa

pine), although Pseudotsuga menziesii (Douglas fir) may also be found on some of the steepest north exposures

in southern Phillips County. Typically there is also a thick understory of junipers, both Juniperus scopulorum

and J. x/assettii. Surrounding vegetation types like sagebrush steppe and juniper steppeAvoodland heavily in-

fluence ponderosa pine-juniper woodland vegetation. Artemisia tridentata, Juniperus communis, Ribes cereum,

Rhus trilobata, and Symphoricarpos occidentalis are common shrubs. Graminoids such as Achnatherum hymen-

oides, Carex inops, Elymus smithii, E. spicatus, Nassella viridula, Poa secunda subspecies, and the exotic grass

Bromusjaponicus are typically found. Achillea millefolium, Parietaria pensylvanica, Pediomelum argophyllum,

Phacelia linearis, Thermopsis rhombifolia var. rhombifolia, and the exotic Tragopogon dubius are common forbs.

Many of these woodlands and surrounding sagebrush steppe have a heavy cover of the exotic Melilotus offici-

nalis, which was often seeded by land managers in revegetation projects even though it can be highly invasive

on the Northern Great Plains (Lesica & DeLuca 2000). In addition to shading out native vegetation, M. offid-

nalis may allow other non-native plants to outcompete native ones by enriching soils with nitrogen (Lesica &

DeLuca 2000).

Montane ponderosa pine forest (Fpp).— These forests are found only in the Little Rocky Mountains in dry

areas at low elevations. Montane ponderosa pine forests occur from about 1,130 to 1,310 m (3,700 to 4,300 ft)

where they begin to transition into lodgepole pine forests. Above these elevations, ponderosa pine is more

857

scarce and usually only on sunny, south exposures. Ponderosa pine is at the northern edge of its range within

the area. In the Cypress Hills (in Canada) and the Sweetgrass Hills, only about 100 km (60 mi) further north

than the Little Rockies, ponderosa pine is absent, apparently because the climate is too cold (Breitung 1954;

Thompson & Kuijt 1976; USGS 1999).

Pinus ponderosa is the dominant tree in these forests with Juniperus scopulorum present in the understory.

The understory also includes such shrubs as Arctostaphylos uva-ursi, Berberis repens, and Juniperus communis

along with the subshrub Artemisia campestris var. pacifica. Representative grasses are Danthonia spicata, Ely-

mus albicans, E. trachycaulus var. trachycaulus, and the exotic Poa compressa. The suite of forbs found in these

montane forests is quite different from those found in the ponderosa pine-juniper woodlands of the Missouri

River Breaks. Anemone multifida, A. patens, Allium cemuum, Cerastium arvense, Cirsium undulatum, Fragaria

vi rginiana, Gaillardia aristata, Helianthus pauciflorus, Maianthemum stellatum, Monardafistulosa, Pterospora

andromedea, Solidago simplex, and Viola adunca are typical forbs.

Montane mixed conifer forest (Fmc).— This forest type is found in the Little Rocky Mountains on mesic

slopes at middle elevations. Tree canopy is made up of a mixture of the conifers Pinus contorta (lodgepole pine), P.

ponderosa, and Pseudotsuga menziesii along with the deciduous tree Populus tremuloides (quaking aspen). Com-

mon shrubs are Arctostaphylos uva-ursi, Berberis repens, Juniperus communis, and Shepherdia canadensis. Repre-

sentative grasses found in these forests are Danthonia spicata, Elymus spicatus, Poa interior, and exotics E. repens

and Phleum pratense. Typical forbs include Achillea millefolium, Campanula rotundifolia. Clematis occidentalis,

Gaillardia aristata, Galium boreale, Linnaea borealis, Maianthemum racemosum, Moehringia lateriflora, Monarda

Jistulosa, Osmorhiza chilensis, Prosartes trachycarpa, Pterospora andromedea, and the exotic Medicago lupulina.

Lodgepole pine forest (Ftp).— Lodgepole pine forests are found in the Little Rockies in dry areas at high

elevations. These forests typically have a closed canopy and an understory depauperate of species. Moderate

disturbance can add some diversity to these forests, but following fires, thick “doghair” stands of young trees

sprout from serotinous cones (Knight 1994). Such stands are common in the Little Rockies. Mountain pine

beetle infestations in these and other forests in the Little Rocky Mountains are minimal at this time. Shrubs

found in lodgepole pine forests are Ceanothus velutinus, Juniperus communis, Rosa nutkana, Salix scouleriana,

and Shepherdia canadensis. Other species commonly found include Galium boreale, Linnaea borealis, Orthilia

secunda, Pterospora andromedea. Spiraea betulifolia, and Thermopsis rhombifolia var. rhombifolia. There are no

subalpine forests found in the Little Rockies. Picea engelmannii Parry ex Engelm. (Engelmann spruce) has been

reported in the nearby Bearpaw Mountains (USGS 1999), which rise to a maximum elevation of 2,108 m (6,917

ft), nearly 365 m (1,200 ft) higher than the Little Rockies.

Montane riparian forest (Fmr).— This forest type is found along moist creek bottoms in the Little Rocky

Mountains, and we have included wetland species found in and along mountain creeks under this vegetation

type. Mixed conifers ( Pinus contorta, P. ponderosa, and Pseudotsuga menziesii) form the canopy with a thick

understory of the deciduous trees Betulapapyrifera (paper birch) and Populus tremuloides and the shrubs AmeL

onchier alnifolia, Comus sericea, Juniperus communis, Prunus virginiana, Ribes spp., Salix bebbiana, and Shepher-

dia canadensis. Typical grasses are Bromus richardsonii, Poa palustris, and exotics B. inermis, Phleum pratense,

and Poa pratensis. Common forbs include Achillea millefolium, Actaea rubra, Agrimonia striata. Clematis occi-

dentalis, Equisetum arvense, Galium boreale, G. triflorum. Geranium richardsonii, Heracleum maximum, Hieraci-

»m umbellatum, Linnaea borealis, Maianthemum racemosum, Mimulus guttatus, various orchids, Prosartes tra-

chycarpa Pyrola asarifolia Sanicula marilandica, Spiraea betulifolia, Symphyotrichum ciliolatum, Viola canaden-

sis, and the exotic Cirsium vulgare. The presence of paper birch in the Little Rockies suggests the presence of

boreal forests in the region following Pleistocene glaciations. Most of the flora of the Little Rockies, however, is

more indicative of a cordilleran influence as in the Sweetgrass Hills (Thompson & Kuijt 1976) and to a lesser

extent the Cypress Hills (Breitung 1954).

Wetlands (W)

Moist coulee bottom and swale (Web) —Some prairie species are most typically found in moist coulee bottoms

®d swales. This habilat also grades Into thickets and wooded conlees if there is enough moisture to support

858 Journal of the Botanical Research Institute of Texas 7 ( 2 )

more woody vegetation and into persistent wetlands if there is surface water. Common forbs in moist coulee

bottoms and swales include Achillea millefolium. Arnica fulgens, A. sororia, Artemisia ludoviciana, Cerastiwn

arvense, Geum triflorum, Glycyrrhiza lepidota, Grindelia squarrosa, Orthocarpus luteus, Potentilla spp., Thermop-

sis rhombifolia var. rhombifolia, Veronica peregrina, and Zigadenus venenosus along with exotics Draba nemorosa,

Thlaspi arvense, and Tragopogon dubius. Common graminoids are Carex brevier, C. praegracilis, Hordeumjuba-

tum subspecies Juncus arcticus, Nassella viridula, and the exotic Poa pratensis. The shrubs Artemisia canajuni-

perus horizontalis, Rosa woodsii, and Symphoricarpos occidentals can also be found.

Distinct from moist coulee bottoms and swales, vernal pools with seasonally standing water can be found

in otherwise flat topography. Eleocharis acicularis, E. palustris, Gnaphalium palustre, Myosurus minimus, Na-

varretia saximontana, Plagiobothrys leptocladus, P. scouleri, and Veronica peregrina are commonly found in ver-

nal pools. Several of the taxa of conservation concern we found grow in these vernal pools as well.

The coulee bottoms and mesic grasslands in the north, particularly in northeastern Valley County, seem

to be indicative of vegetation types more common to the north in Canada. In the Opheim Hills and to the east,

the shrubs Dasiphorafruticosa and Elaeagnus commutata can also be found in moist swales. Populus tremuloides,

rare on the plains of eastern Montana but more common further north in Canada (Coupland 1961; Cooper et

al. 2001), can be found in some of the coulees of the Opheim Hills and northeastern Phillips County as well A

few species found nowhere else were present in these moist habitats: Carex obtusata, Fragaria vesca , Geranium

viscosissimum. Primula pauciflora, Viola nephrophylla, and Zizia aptera. Many of these species are more com-

mon on the Canadian prairies further north (Budd 1979). Other species were only encountered elsewhere in

the Little Rockies including Carex bebbii, C. sprengelii. Delphinium bicolor, Heracleum maximum, Shepherdia ca-

nadensis, and Viola canadensis. The grasses Elymus lanceolatus varieties and Hesperostipa curtiseta were also

frequently found in these locations. Festuca hcdlii, the principal grass of the fescue prairies of Canada (Coup-

land 1961), was found only once in the study in northeastern Valley County just a few miles south of Canada.

This area receives slightly greater precipitation and is generally colder than the rest of the area (PRISM 2004).

The Hesperostipa curtiseta-Elymus lanceolatus grasslands found in northeastern Valley County are much

more common in Canada than in the U.S. However, they were once more prevalent in both countries before

such sites, which are well suited to grain production, were put under cultivation (Cooper et al. 2001). Indeed,

most of the lands east of Opheim are in cultivation and privately owned. A sizable expanse of this prairie as-

sociation in a large area of Montana State Trust Lands along Dry Fork Creek in northern Valley County repre-

sents one of, if not the best, remaining of its kind in the U.S. (Cooper et al. 2001).

Persistent wetland (Wpw). — Most persistent wetlands are located around small reservoirs although they

also occur along large creeks and small pools in creek beds where open water persists throughout the growing

season. Around the periphery of wetlands, which may be submerged in the spring and early summer but are

often dry by autumn, graminoids such as Beckmannia syzigachne, Carex spp., Echinochloa muricata, Eleocharis

acicularis, E. palustris, Hordeumjubatum subspecies Juncus arcticus, Poa palustris, and the exotic grass Polypo-

gon monspeliensis are common along with the forbs Conyza canadensis, Glycyrrhiza lepidota, Lycopus asper,

Mentha arvensis, Rumex spp., exotic Sonchus arvensis and the noxious weed Cirsium arvense. Common shrubs

on the periphery of wetlands are Rosa woodsii and Salix exigua ssp. interior. Occasionally the trees Populus deb

toides and Salix amygdaloides may occur as well. Emergent aquatic plants typically growing in standing water

throughout the growing season are Alisma gramineum, A. triviale, Bolboschoenusfluviatilis, B. maritimus, Limo-

sella aquatica, Persicaria amphibia, P. lapathifolia, Sagittaria cuneata, Schoenoplectus spp., Typha angustifoUa,

and T. latifolia. Common submerged aquatics are Ceratophyllum demersum, Potamogeton spp., Ranunculus

Alkaline wetland (Wal). — Many wetlands are alkaline at least to s

:a are derived from marine shales. Many species found in freshw;

:xtent because soils in most of the

rea are derived trom marine shales. Many species found in freshwater wetlands are also found in alkaline

wetlands but the most alkaline typically have a unique assemblage including Distichlis spicata, Glaux maritima,

Hordeumjubatum subspecies, Jva axillaris, Juncus arcticus, Puccinellia nuttalliana, Salicomia rubra, Spergularia

marina, Triglochin maritima, the subshrub Suaeda calceoliformis and the exotic Polygonum aviculare.

Charboneau et al., Flora of Phillips and Valley counties, Montana

Sparsely vegetated alkaline pan areas are also common. These pan areas are formed above high points on

the shale-till boundary beneath the soil surface. Salts from marine shales accumulate here and cause the forma-

tion of natric horizons in the subsoil, which greatly reduces infiltration of precipitation (Munn & Boehm

1983). Few plants can thrive in these water-stressed, alkaline conditions, so plant cover is very sparse with low

diversity. Atriplex suckleyi, Dieteria canescens, Distichlis spicata, Elymus smithii, Hordeum jubatum subspecies,

Iva axillaris, Monolepis nuttalliana, Oenothera cespitosa, Puccinellia nuttalliana, the exotic Polygonum aviculare,

and the subshrub Atriplex gardneri var. gardneri are among the few species typically encountered.

Sparsely Vegetated (V)

Badlands (Vbl ). — Badlands are common where marine shales are exposed. When wetted, these badlands form

slick, alkaline clay that cracks extensively upon drying and erodes so rapidly that little vegetation can be estab-

lished. The few species that can survive on badlands are often ruderal and tolerant of alkalinity. These include

Atriplex argentea, A. suckleyi, Eriogonum pauciflorum, Iva axillaris, Monolepis nuttalliana, Oenothera cespitosa,

Penstemon nitidus, and exotics Conringia orientalis and Polygonum aviculare occasionally with subshrubs Atri-

plex gardneri var. gardneri, Suaeda calceoliformis, and the shrub Sarcobatus vermiculatus.

Shale dunes, somewhat similar to badlands but less common, are found especially in the north in Bitter

Creek WSA and the Frenchman Creek valley. These dunes are formed by the wind when shale weathers into

sand-sized particles or small, thin flakes rather than clay minerals. Juniperus horizontalis typically stabilizes

these dunes. Other species commonly found are Artemisia longifolia, Eriogonum pauciflorum, Oenothera cespi-

tosa, Rosa spp., Stephanomeria runcinata, and Thermopsis rhombifolia var. rhombifolia.

Rock outcrops and talus (Vot ).— The Little Rocky Mountains have areas of both granitic and carbonate

rock outcrops. Chamerion angustifolium var. angustifolium, Cheilanthes feei, Draba cana, Erigeron composi-

tus, Eriogonum ovalifolium var. purpureum, Minuartia rubella, Poa glauca, Sedum lanceolatum, Townsendia

hookeri, and Woodsia oregana are among the herbaceous species found on these outcrops. The shrubs Da-

siphora fruticosa and Ribes cereum can be found as well.

There are also several large areas of sparsely vegetated talus fields in the Little Rockies. Ceanothus veluti-

nus, Chamerion angustifolium var. canescens, Prunus pensylvanica, Ribes cereum, R. oxyacanthoides var. oxyacan-

thoides, and Rubus idaeus are typically found on this talus.

Disturbed (D)

There are many disturbed habitats covered by ruderal forbs and grasses (many are invasive). These are primar-

ily found along roadsides but also in dry reservoir beds, on reservoir dams, and in reseeded fields. Areas dis-

turbed by natural action such as fires, flooding, and animal burrows have many of the same species. Typical

exotic forbs of these habitats include Alyssum desertorum, Camelina microcarpa, Descurainia sophia, Kochia

scoparia, Lactuca serriola, Lappula occidentalis, Medicago lupulina, M. sativa, Melilotus officinalis. Polygonum

aviculare, Salsola tragus, Thlaspi arvense, Tragopogon dubius, and the noxious weed Convolvulus arvensis. Na-

tives Chamaesyce spp., Chenopodium berlandieri, Grindelia squarrosa, Helianthus annuus, Lepidium densiflorum

varieties, Monolepis nuttalliana, Plantago patagonica. Polygonum achoreum, and Verbena bracteata are common

in disturbed habitats as well. Typical weedy grasses are the exotics Agropyron cristatum varieties, Bromus iner-

mis, B.japonicus, B. tectorum, Eragrostis cilianensis, Poapratensis, and natives Hordeum jubatum subspecies and

Munroa squarrosa.

A few species were only found planted and persisting at old homesteads and other such sites. These are

Caragana arborescens, Cotoneaster lucidus, Lonicera tatarica, Malus pumila, Ulmus americana, and U. pumila.

Syringa vulgaris L. was also present but never collected at such sites. Many of the “historical” taxa added to the

checklist were collected in disturbed areas including farm fields, gardens, and lawns. Over 35 percent of the

historical taxa added to the checklist are exotic to Montana while about 14 percent of the taxa we collected are

exotic (Mincemoyer 2012).

Taxa of Conservation Concern

Fifteen taxa of conservation concern were documented from 34 sites. These taxa are tracked by the Montana

Journal of the Botanical Research Institute of Texas 7(2)

Natural Heritage Program with state ranks of SI, S2, or S3 or are listed as sensitive by the Bureau of Land Man-

agement (MTNHP 2012b). These taxa are indicated by a diamond (♦) in the annotated checklist and listed al-

phabetically below.

Ammannia robusta (Lythraceae) was found in Valley County in a reservoir and adjacent mudflat. Voucher:

Nelson 81384.

Anagallis minima (Myrsinaceae) was found in Phillips and Valley counties in vernal pools. Vouchers:

Charboneau 2486, 7921.

Bacopa rotundifolia (Plantaginaceae) was found in Phillips County on the edge of a reservoir. Voucher:

Charboneau 9535.

Botrychium hesperium (Ophioglossaceae) was found in Phillips County in a rocky disturbed area in lodge-

pole pine forest. Voucher Charboneau 2120.

Carex scoparia var. scoparia (Cyperaceae) was found in Phillips County in a juniper thicket in the Missouri

River Breaks and in a montane meadow. Vouchers: Charboneau 2298, 7690.

Elodea bifoliata (Hydrocharitaceae) was found in Phillips County floating in reservoirs. Vouchers:

Charboneau 9431, 9516, 9541.

Phlox andicola (Polemoniaceae) was found in Phillips County in sagebrush steppe. Voucher: Charboneau

5069.

Physaria brassicoides (Brassicaceae) was found in Phillips County in a montane meadow. Voucher:

Charboneau 4812.

Physaria ludoviciana (Brassicaceae) was found in Valley County in mixedgrass prairie. Vouchers:

Charboneau 4862 ; Nelson 82012.

Plagiobothrys leptocladus (Boraginaceae) was found in Phillips and Valley counties in vernal areas. Vouch-

ers: Charboneau 1373b, 5791, 6144, 6870, 7209 ; Nelson 80119, 80180, 80542, 81590.

Psilocarphus brevissimus var. brevissimus (Asteraceae) was found in Phillips County in a vernal area.

Voucher: Charboneau 7286a.

Ranunculus hyperboreus (Ranunculaceae) was found in Valley County floating in a creek. Voucher:

Charboneau 2462.

Senecio eremophilus var. eremophilus (Asteraceae) was found in Phillips County in montane disturbed

areas. Vouchers: Charboneau 2141, 9167; Nelson 81011.

Sphenopholis intermedia (Poaceae) was found in Phillips County in mixed conifer forest. Voucher:

Charboneau 2199.

Suckleya suddeyana (Amaranthaceae) was found in Valley County in dried reservoir bottoms and shores.

Vouchers: Charboneau 2736, 3354, 3843, 3860; Nelson 81378.

nuda (Loasaceae), Penstemon grandiflorus (Plantaginaceae), Phacelia thermalis (Boraginaceae), Potentilla plat-

tensis (Rosaceae), and Schoenoplectus heterochaetus (Cyperaceae).

Exotic Taxa and Noxious Weeds

We collected 108 taxa exotic to Montana (Mincemoyer 2012), comprising 14.2 percent of the 762 taxa we col-

lected. These taxa are indicated in the annotated checklist by an asterisk (*). Nine species (10 taxa) of the 32

species recognized as noxious weeds in Montana (MNWP 2010) were documented. These were Acroptilon re-

pens, Centaurea diffusa, C. stoebe, Cirsium arvense. Convolvulus arvensis, Cynoglossum officinale, Euphorbia esula

varieties, Leucanthemum vulgare, and Tamarix chinensis. In the annotated checklist these taxa are indicated by

a circle (•). The most widespread and common of these noxious weeds are Euphorbia esula varieties and Cirsi-

um arvense. Two Montana regulated plants (priority three weeds; MNWP 2010) were also found: Bromus tecto-

rum and Elaeagnus angustifolia. Twenty-five of the 70 “historical” taxa added to the checklist (35.7 percent) are

exotic to Montana (Mincemoyer 2012). Among these additional taxa are two Montana noxious weeds (MNWP

a, 16.0 percent of the

u et al., Flora of Phillips and Valley counties, Montana

2010): Lepidium latifolium and Tanacetum vulgare. With the addition of the “historical” taxa

taxa included in the annotated checklist are exotic (Mincemoyer 2012).

Newly Documented Taxa

The area’s vascular flora was previously poorly documented. We collected 227 taxa and 201 species that had

previously been undocumented (GPFA 1977; Hartman et aL 2009; Lesica 2012; USDA 2012; Kartesz 2013;

MONT 2013; MONTU 2013). This accounts for 29.8 percent of the 762 taxa we collected. Only 8.8 percent of

these 227 taxa are exotic to Montana (Mincemoyer 2012) indicating that these newly documented taxa pre-

dominantly are not newly introduced to the area. Of the 12,768 specimens we collected, 446 or more than one

in every 29 collections are county records in either Phillips County or Valley County. On average we collected

over four county records per person-day in the field.

GIS analyses.— In our assessment of the sampling adequacy of environmental conditions by our collection

sites, we found that our sites did nearly as well as a set of random points. There were 66 combinations of eleva-

tion, average annual precipitation, and average daily minimum temperature classes within the lands accessible

for collecting. Our actual collection sites were located in 42 of these combinations while a random set of the

same number of points was located in 44 combinations. Our collection sites missed combinations comprising

2.2 percent of accessible lands, while the random points missed combinations totaling 1.1 percent.

lection sites outperformed random points in sampling land cover types. Thirty-nine land cover types are re-

ported within accessible lands (MTNHP 2010). These are the same types described in the Montana Ecological

Systems Field Guide (MTNHP 2012a). Our collection sites sampled 25 cover types, while the set of random

points was only in 15. Our collection sites missed cover types totaling 1.1 percent of accessible lands, while

random sites missed cover types making up 2.3 percent.

In both analyses, the frequency of collection sites and random points for the most part mirrored the fre-

quency of environmental class combinations and land cover types of accessible lands, with important excep-

tions. Our actual sites oversampled rare combinations and cover classes such as those found in the Little Rocky

Mountains while undersampling the most common combinations and classes. This allowed us to better docu-

ment all of the taxa found in rare habitats. Random points also have the disadvantage of often being further

in chronological order. The number of taxa collected levels ofF in the second year of the inventory as few new

taxa were encountered in May and June 2011, although almost 100 were encountered for the first time in July

and August 2011. In total 630 taxa (almost 83 percent) were encountered during the first field season, and an

additional 132 were collected for the first time during the second field season.

Figure 4 shows the taxon accumulation curve averaged from 100 randomizations of the order of collect-

ing days. The curve levels off fairly well with 90 percent of observed taxon richness encountered by about 60

of 102 collecting days. The asymptote of the species accumulation curve as predicted by the Michaelis-Menten

equation (see Colwell and Coddington 1994) reaches 797 taxa, only 35 more than we observed. Parametric

estimators gave higher estimates of diversity: the bootstrap estimator predicted 829 taxa, the Chao 1 estimator

885 taxa, and the second-order jackknife estimator 965 taxa. The addition of 70 “historical” taxa to the check-

list brings the total number of known taxa to 832, greater than predicted by the Michaelis-Menten equation

and the bootstrap estimator based on our collections. Many of the “historical” taxa added were collected in

habitats that we did not focus our efforts on such as lawns, gardens, and cultivated fields, which may explain

this discrepancy. Based on the addition of these “historical” taxa and our estimates of the taxon diversity pres-

ent in the area, we collected between 79 and 91.6 percent of the taxa growing in the area.

Our estimate of the actual diversity documented and our analyses of the environmental conditions and

862

Journal of the Botanical Research Institute of Texas 7(2)

land cover types sampled by collection sites show we performed adequately in documenting the diversity of

vascular pUnts^Because of the number of taxa documented for the first timein July and August of the second

fie!d season and the relatively short time spent collecting in September, the late summer and early fall likely

would be the most worthwhile part of the growing season for further collecting.

CONCLUSIONS

This inventory has greatly expanded the floristic knowledge of a 23,191 sq km (8,954 sq mi) area of northeast-

«n Momana. Approximately one in every 29 collections made (446 of 12,768) were county records in either

fionl'rtr ^ ^ley County, and about 3° percent of the taxa we documented were previously unknown

from the area. In total, we collected 762 vascular p!anttaxafrom86 families, an estimated 79-4>2 percent of the

actual vascular plant diversity present in the area. The addition of 70 “historical- taxa brings the total number

manv rarest r b r area *d 832. This study demonstrates there is still much to be learned about the flora of

many parts of the contiguous United States.

■ i

confusion Collection data are available ocbr^at'Ltpy^ww^huw^edu^Belo^is^key to'the a^bn

.ethefoiiowingform^

Menten estimator, Tobs = taxa observed. Data generated using Estimates (Colwell 2013).

County abbreviations:

PH Phillips

Riparian cottonwood forest

Thicket and wooded coulee

• Montana noxious weed

▼ Taxon of conservation c<

* Putative hybrid

VA Valley

Gup Upland prairie

$gs Greasewood shrubland

Sjw Juniper steppe/woodland

Sss Sagebrush steppe

Vbl Badlands

Vot Rock outcrops and talus

Wal Alkaline wetland

Web Moist coulee bottom and s

Wpw Persistent wetland

Charboneau et al.. Flora of Phillips and Valley counties, Montana

Verbena bracteata Lag. & Rodr. (14) PH, VA; 650-935 m; D

var .adunca (5) PH; 1 195-1440 m;Flp, Fmc, Fmr, Fpp

Viola vallicola A. Nelson ( 1 5) PH, VA; 71 0-1 440 m; Gmg, Gmm, Web

ACKNOWLEDGMENTS

Funding for this study was provided by the Montana/Dakotas BLM, Dr. Thomas Ford (Aven Nelson Fellow-

ship in Systematic Botany), and the University of Wyoming. We thank John Carlson and Wendy Velman of the

Montana/Dakotas BLM for facilitating the project. Lawrence Schmidt of UW Libraries was also instrumental

in the project’s creation.

The following individuals are greatly thanked for providing access for collecting, logistical support, or

on-the-ground knowledge of the area: Stephen Klessens and Raymond Neumiller of the Glasgow BLM Field

Office, Richard Adams of the Malta BLM Field Office, Robert Skinner and Aaron Johnson of Charles M. Russell

National Wildlife Refuge, Jessica Larson of Bowdoin National Wildlife Refuge, Shawn Cleveland of The Nature

Conservancy, and Damien Austin of the American Prairie Reserve. Damien Austin, Katie Butts, Michael

Dolan, Mary Frieze, Joshua Lamp, Mark Majerus, Matthew Ocko, Noorjahan Parwana, Alice Sawyer, and Ben-

jamin Wagner provided collecting help.

Matt Lavin and David Dyer are acknowledged for providing access to MONT and MONTU respectively to

verify “historical” collections. The first author thanks committee members that included Gregory Brown and

Larry Munn for their time and efforts improving this manuscript. Reviewers Mark Gabel and Peter Lesica are

recognized for their helpful comments on the manuscript.

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TAXONOMIC IDENTITY AND HISTORICAL ACCOUNTS OF

DALEA CYLINDRICEPS (FABACEAE), A SPECIES OF CONSERVATION CONCERN

IN THE GREAT PLAINS (U.S.A.)

James H. Locklear

Omaha, Nebraska 68 1 08, U.S.A.

Daka cylindriceps Barneby (Fabaceae) i:

1 herb (Figs. ]

INTRODUCTION

0 native to the western Great Plains of

Kansas, Nebraska, New Mexico, Okla-

homa, South Dakota, Texas, and Wyoming. Despite the large extent of its historical distribution, D. cylindriceps

has been collected infrequently and occurrences are scattered and local. As noted in the recent Flora of Ne-

braska (Kaul et al. 2011), “This distinctive species is rare almost throughout its wide geographic range.” Dalea

cylindriceps is tracked as a species of conservation concern in all but two of the states in which it has been docu-

mented, and is ranked by NatureServe (2013) as G3G4 (vulnerable).

Initial field survey by the author in the northern part of the range of D. cylindriceps indicates this species

may have experienced considerable population decline. In 2010, 1 searched the sites of 22 historical occur-

rences of D. cylindriceps (derived from herbarium specimen locality data) in Co orado, Nebraska, and Wyo-

ming and found existing populations at only 4 sites. It should be noted that D. cylindriceps is a lelatively large

plant (generally 3-6 [up to 8] dm tall) with distinctively elongate (up to 18 cm long) flowering spikes, making

it a noticeable object in the landscape.

Analysis of floristic and ecological literature informs conserv,

formation on distribution, abundance, ecological associations, etc. In the case ot j

sis is made difficult by the complex nomenclatural history of this taxon. C an cm

for this species will allow for more accurate assessment of its conservation sta

>sment by yielding historical in-

i-Bot Res. Inst. Texas 7(2): 879 - 890. 2013

Locklear, Dalea cyl indriceps, a species of conservation c

Fig. 2. Dalea cylindriceps, Sheridan County, Nebraska. Photo James Locklear.

The taxonomic history of this species was reviewed by Bameby (1977) in his monograph on the genu

L., in which he determined that a new binomial was required and proposed the currendy accepted n

cylindriceps Bameby.

The earliest published name for this taxon is Petalostemum macrostachyum Torr., <

material collected in 1820 by Edwinjames in present-day Nebraska (see below). In Flora of North America , Tor-

rey and Gray (1838, p. 309) altered the rendering of the generic name to Petalostemon (giving the full binomial

as Petalostemon macrostachyum) and listed P. omatum Douglas ex Hook, as a synonym. Gray later (1848) dis-

tinguished P omatum from P. macrostachyum, and the former is now recognized as Dalea omata (Douglas)

Eaton & Wright, a species of the Columbia Plateau and northern Great Basin. This ear

resulted in P macrostachyum being treated in certain 19th century botanical literature

Northwest as well as the Great Plains (e.g.. Coulter 1885).

Torrey and Gray later (1840, p. 690) added Dalea compacta Spreng. as a synonym under P macrostachyum

ta, as determined by Bameby 097^0 eompochr is a separate species .hat occurs in eastern .Texas and ac-

cent Arkansas and Oklahoma. Misinterprecation of D. ampacta, dong wkh misappUcatkm of (he speoffcepi-

thet, has been at the root of much of the nomenclatural confusion surrounding D. cylindriceps (Peterson 2000).

Torrey’s Petalostemum macrostachyum (cor

masculine gender of Petalostemon) appears in the early botanical lit.

wood 1893; Engelmann 1862; Gray 1848, 1863; Parry 1870; Porter & Coulter 1874), but subsided after pubhca-

ribed in 1827 from

a plant of the Pacific

the Great Plains region (East-

Journal of the Botanical Research Institute of Texas 7(2)

tion of Petalostemon compactus (Spreng.) Swezey in 1891 in Nebraska Flowering Plants (Doane College Natural

History Studies No. 1), a 17-page pamphlet written by Goodwin D. Swezey, professor at Doane College in Crete,

Nebraska. Swezey cited “P. macrostachyus Torr.” as a replaced synonym of his P. compactus. While proposed in

a relatively obscure publication, Swezey’s name apparently gained recognition and adoption thanks to the re-

view of his pamphlet by Nathaniel Lord Britton (1891) published in the widely-circulated Bulletin of the Torrey

Botanical Club.

Swezey’s P. compactus was a new combination based on D. compacta (Sprengel 1826), which Torrey and

Gray (1840, p. 690) included in the concept of Petalostemon macrostachyum (Torrey 1827) but which Swezey

elevated based on prior publication. It should be noted that Swezey applied the name P compactus to specimens

of what is in fact D. cylindriceps. Two such specimens are known to the author. One is in the herbarium of Do-

ane College in Crete, Nebraska, collected by E.E. Sprague (s.n.) on 25 July 1890 near Lewellen in Garden

County, Nebraska (annotated by S. Rolfsmeier [1987] as D. cylindriceps). The other is at NY (1259224) and ap-

pears to be a duplicate of the Sprague collection (with same collection date and locality) distributed under an

“Herb. ofG.D. Swezey” label (annotated by D. lsely [1958] andD. Wemple [1961-1965] as “P. compactum”).

Further clouding the nomenclature of this taxon was the reduction of Petalostemon Michx. and Dalea L.

to the genus Kuhnistera Lam. by Kuntze (1891), in which he proposed K. compacta (Spreng.) Kuntze, under

which he placed D. compacta and “ Petalostemon macrostachyum” as synonyms. Heller later (1896) added “Pet-

al ostemon compactus Swezey” to the list of synonyms under K. compacta. Use of the name K. compacta for what

is clearly D. cylindriceps occurred in a number of botanical and ecological publications in Great Plains states

(Hitchcock 1896; Pound & Clements 1900; Saunders 1899).

The publications of Per Axel Rydberg were foundational to state and regional treatments of the flora of the

Great Plains in the first part of the 20th century. Regarding D. cylindriceps, Rydberg initially (1894, 1895) took

up K. compacta for the taxon, citing Petalostemon macrostachyus and D. compacta as synonyms, but subse-

quently adopted Petalostemon compactus (Spreng.) Swezey, first in Flora of Colorado (1906), then in Flora of the

Rocky Mountains and Adjacent Plains (1917) and Flora of the Prairies and Plains of Central North America (1932).

During this same period, Britton and Brown (1897) treated this taxon as K. compacta but later (1913) adopted

Swezey’s name, rendering it “ Petalostemum compactum (Spreng.) Swezey.”

Subsequent treatments of this taxon in the flora of the Great Plains region followed Rydberg in using

Swezey’s Petalostemon compactus (often rendered P. compactum). These include floristic and ecological works

for the states of Colorado (Harrington 1954; Ramaley 1939), Kansas (Bare 1979; Barkley 1968; Gates 1940),

Nebraska (Petersen 1923; Webber 1892), New Mexico (Wooton & Standley 1915; Martin & Hutchins 1980),

Texas (Correll & Johnston 1970; Turner 1959), and Wyoming (Dorn 1977b), as well as other regional literature

(Barr 1983; Coulter & Nelson 1909; Dorn 1977a; Rogers 1953) including Atlas of the Flora of the Great Plains

(GPFA 1977).

The taxonomic identity and nomenclatural problems associated with P. compactus (Spreng.) Swezey re-

ceived further consideration and clarification in the 1970s. In his revision of the genus Petalstemon, Wemple

(1970) recognized “ Petalostemon compactum (Spreng.) Swezey,” treating both P macrostachyum and D. com-

pacta as synonyms. But, in his monograph on the genus Dalea, Barneby (1977) separated D. compacta from the

taxon originally described as P. macrostachyum, which excluded the type of D. compacta from his concept of P

macrostachyum, making the latter the oldest available name for the species. Since Barneby was reducing the

genus Petalostemon to Dalea, he needed to transfer P. macrostachyum to Dalea, but there was already a D. mac-

rostachya Moric., necessitating the “unwelcome new epithet” of cylindriceps. Dalea cylindriceps was adopted in

Flora of the Great Plains (GPFA 1986) and has since been used in most treatments of the flora of the region

(Dorn 2001; Kaul et al. 2011; Turner et al. 2003; Van Bruggen 1985; Weber & Wittmann 2012). Correct nomen-

clature and synonymy for this species is provided below.

Dalea cylindriceps Barneby, Mem. New York Bot. Gard. 27:227. 1977. Type: United States: “Long's 1st Expedition. Dr.

South Platte River in Lincoln County, Nebraska, 22-23 Jun 1820.

883

Discovery and Type Locality

The type specimen of what would eventually be recognized as D. cylindriceps was collected by Edwin James

while traveling with the Stephen H. Long Expedition of 1820. In his enumeration of the botany of the expedi-

tion, Torrey (1827) stated the locality of James’ collection of Petalostemum macrostachyum as “About the forks of

the Platte.” No date or locality information is provided on the holotype (the only known specimen) at NY (Fig.

3), only the statement, “Long’s 1st Expedition. Dr. James.” James did not mention this species in either his pub-

lished account of the expedition (James 1823) or in his personal diary (Goodman & Lawson 1995).

The North and South Platte rivers (“the forks of the Platte”) join to form the Platte River in Lincoln Coun-

ty, Nebraska, just east of the city of North Platte. From here westward to Ogallala, Nebraska (a distance of ca.

70 km), the main channels of the North and South Platte rivers run within a few (5-6) kilometers of each other

until they begin to diverge about 18 km east of Ogallala, the North Platte to the northwest and the South Platte

to the southwest. Thus, “About the forks of the Platte” could be a general description of the area between North

Platte and Ogallala, which the expedition traversed 22-25 June 1820 (Goodman & Lawson 1995).

But this phrase appears to refer to a more limited geographic area. After reaching the junction of the North

Platte and South Platte on 22 June 1820, the expedition continued west along the north side of the North Platte

River for a few miles then crossed the river heading southwest, making camp on the north side of the South

Platte River. In their reconstruction of the route and itinerary of the Long Expedition, Goodman and Lawson

(1995, p. 15) place the location of the camp of June 22 at 6-7 miles (10-12 km) west of the city of North Platte,

which would bVjust to the east of the present-day community of Hershey, Nebraska.

On the morning of June 23, the party traveled about two miles (3.3 km) upstream then crossed to the

south side of the South Platte. The crossing would have been in the vicinity of Hershey. James (1823, p. 468)

states that the party “had no sooner crossed the [South] Platte when our attention was arrested by the beautiful

white primrose (Oenothera pinnatijida. N .)” (« O. coronopifolia Torr. & A. Gray). In a footnote to his discussion

of the Oenothera, James lists several other species that were collected “about the forks of the Platte.”

Thus, it appears that the area referred to by James as “about the forks of the Platte” is tied to the locality of

this crossing of the South Platte River in present-day Lincoln County, Nebraska on 23 June 1820. The type of

D. cylindriceps was likely collected along the river somewhere between the towns of Hershey and Sutherland

(ca. 12 km to the west), the latter being the vicinity of the expedition’s camp of June 23 (Goodman & Lawson

1995, p 16) This conclusion is supported by a collection of D. cylindriceps made on 05 August 1989 (D. Suther-

land 6802 with S. Rolfsmeier at NEB, NY) along the South Platte River just south of Hershey.

Historical Accounts . ......

Armed with knowledge of the nomenclatural history of D. cylindriceps, the early flonstic and ecological htera-

tore of the Great Plains was searched for references to this specie* The following is a summary of notable

^ ndl Ihllea cylindriceps was encountered by many of the early botanical collectors traveling aooss the cc ”^

Great Plains in the MOs. The list includes Edwin James in 1820 (Toney 18CT Aug^ Fendfa m 1846

(Gray 1848), Ferdinand Hayden in 1857 (Engehnann 1862), and Elihu Hall and J.P. Harbour (with Charles C.

^^fretoatandance is indicated, D. cylindriceps was typically

ized, etc (Hitchcock 1896; Pound & Clements 1900; Ramaley 1939; Rogets 1953) Tins ts tn spue of fact that

the species is a noticeable object in the landscape when present, as reflected in the comments of Pound and

Clements (1900) in The Phytageogmphy of Nebraska, where they described D. cylindriceps (as K. ccmpacm) as

885

“the most remarkable species of this genus in our flora. Its stems, which are densely leafy, are often a meter high

and are very conspicuous on account of the long (a decimeter) nodding spikes of white or yellowish flowers .”

Dalea cylindriceps may have been more abundant prior to settlement and agricultural development within

its range. Eastwood (1893) included “ Petalostemon macrostachyus Torr” in A Popular Bora of Denver, Colorado ,

her book aimed at “students” and “beginners” and therefore focused on the showier or more common elements

of the flora. Eastwood stated the species occurred in “North Denver.” There are a number of D. cylindriceps

herbarium specimens collected in the late 1800s from what is today the Denver metropolitan area northward

toward Fort Collins, but the native vegetation of this area has been largely displaced by suburban and ex-urban

development.

The following references provide early accounts of the ecological associations of D. cylindriceps :

Colorado: “Sand Prairie” (Ramaley 1939, p. 14 as P. compactus). In his ecological study of the sandhill vegeta-

tion of northeastern Colorado, Ramaley recognized four plant communities occurring in upland sandy

soils (Loose Sand and Blow-out; Sand-hills-Mixed; Sand-Sage; Sand Prairie); he listed P compactus only

under Sand Prairie, where he placed it on the list of “Other species,” indicating relative infrequency.

Kansas: “Prairie” (Gray 1848, p. 33 as P macrostachyum ). Gray enumerated plants collected by Augustus

Fendler in the vicinity of Santa Fe, New Mexico, which he reached in 1846 after traveling from St. Louis

along the Santa Fe Trail. Gray reported the locality of the collection as “18 miles west of Lower Springs,

Cimarron [River].” Lower Spring (also called Wagon Bed Spring) was a site along the Cimarron Cutoff of

the Santa Fe Trail located on the Cimarron River in what is today Grant County, Kansas. A location along

the trail 18 miles west of Lower Spring would be in present-day Morton County, Kansas in the vicinity of

the Cimarron National Grasslands, where a number of recent collections of D. cylindriceps have been

Kansas: “Ulysses, Grant County... on sandy knolls along the South Fork of the Cimarron [River]. Rare.”

(Hitchcock 1896, p. 543 as K. compacta ). Hitchcock reported on the plants collected by C.H. Thompson

in southwestern Kansas in 1893.

Nebraska: “Sand hills along the Loup fork and Niobrara [rivers]” (Engelmann 1862, p. 189 as P. macrostachy-

ium [sic]). Engelmann enumerated plants collected by Ferdinand Hayden in the Elkhom, Loup, Platte,

and Niobrara river valleys of present-day Nebraska during an expedition led by Lieutenant G.K. Warren

of the U.S. Topographical Engineers in 1857.

Nebraska: “Localized in the sand-hills of Scotts Bluff county (sic)” (Pound & Clements 1900, p. 250 as K.

compacta). Pound and Clements likely based their account of this species on herbarium material col-

lected by PA. Rydberg ( Rydberg 61; NEB 177085, 177088; NY 1259216, 1259217) on 04 August 1891.

Rydberg identified his number 61 as “Petalostemon macrostachyus Torr.” in his journal (manuscript at

NEB), noting the locality as “On the sandhills north of the Platte River, Scotts Bluff Co.”

Such early references, as well as those found in more recent literature (Hazlett 2004; Kaul et al. 2011; Kuhn et

al. 2011; Neid et al. 2007; Reif et al. 2009; Rolfsmeier & Steinauer 2010; Sutherland & Rolfsmeier 1989), indi-

cate a strong association of D. cylindriceps with sandy habitat. Eolian sand sheets and sand dunes are common

landforms in the western Great Plains, notably in Nebraska, Colorado, Kansas, Oklahoma, New Mexico, and

Texas (Muhs & Holliday 1995). At present, these dune fields are mostly stabilized by vegetation.

Review of Great Plains floristic and ecological literature coupled with examination of herbarium speci-

mens and the author’s initial field survey, indicate that D. cylindriceps is frequently associated with plant com-

munities in which the shrub sand sagebrush (Artemisia filifolia) is a dominant or distinctive element. Occur-

rences of D. cylindriceps in southwest Nebraska, eastern Colorado (Fig. 4), and southwest Kansas are associated

with steppe communities comprised ofasparse to moderately dense layer oiA.filifolia interspersed with tailor

mid-height grasses, the component grass species varying with geography, precipitation, sod texture, etc.

(Kuchler 1974 Lauver et al 1999; Neid et al. 2007; Ramaley 1939; Rolfsmeier & Steinauer 2010). Frequently

referred to as “sandsage prairie,” these sometimes extensive communities occur on sands and loamy sands as-

sociated with eolian dune systems.

886

journal of the Botanical Research Institute of Texas 7(2)

Dalea cylindriceps has been collected (to a lesser exte

il deposits of streams and colluvium derived from sandstone outcrops and escarpments

jus plant communities that develop in such habitat occur as small patches or bands <

The association of D. cylindriceps with sandsage prairie could be a factor in the apparent decline of this

species. Throughout the Great Plains, extensive tracts of sandsage prairie have either been converted to irri-

Sexson 1983).

Future Research

University of Kansas botanist Ronald L. McGregor (1986) asserted nearly 30 years ago that D. cylindriceps

“needs more careful study ” and his statement holds true yet today. Additional research is needed to more fully

assess the conservation status of D. cylindriceps. Survey of historical occurrences throughout the range of the

species is needed to determine the number of existing occurrences and to develop a more detailed ecological

profile that includes habitat requirements, edaphic factors, disturbance factors, associated species, etc. As a

more precise understanding of the ecology of D. cylindriceps is gained, field workers can be more strategic in

arch for additional occurrences.

Research into the life history traits (phenology, reproductive ecology, etc.) of D. cylindriceps is needed t<

nine how these shape demography and population trends. This species appears to be a short-lived or evei

887

monocarpic herbaceous perennial (Bameby 1977; Kaul et al. 2011). McGregor (1986) noted that on dunes and

areas of loose sand, D. cylindriceps “sometimes flowers the first year and frequently expires at the end of the

second or third season,” yet is often a longer-lived perennial in more stable sandy areas.

Many plant species associated with sandy habitat are specialized to a particular ecological niche or stage

of recovery related to natural disturbance, and research is needed to determine if such is the case for D. cylin-

driceps. Sandsage prairie is a naturally dynamic plant community, with species composition, patterns of vege-

tation, and percent canopy cover changing over time in response to fluctuations in precipitation (Collins et al

1987; Farrar 1993b; Kelso et al. 2007; Ramaley 1939; Rondeau 2003, 2013). Such recurring natural disturbance

could result in fluctuations in the presence and abundance of D. cylindriceps over time.

Monitoring studies of existing populations of D. cylindriceps are needed to answer these and other ques-

tions of life history, demography, and population trends. Further study of D. cylindriceps would be of value not

only for the conservation and management of this species, but also holds promise for a better understanding of

the ecology and dynamics of sandsage prairie, a plant community that is of conservation concern throughout

most of its distribution in the western Great Plains (NatureServe 2013). Given its close association, D. cylindri-

ceps could serve as an indicator species of high quality occurrences of sandsage prairie and of habitat integrity

and health.

A Common Name for Dalea cylindriceps Bameby

The misleading common name “Andean prairie-clover” has been applied to D. cylindriceps and is currently in

use by NatureServe (2013) and other databases listing plants of conservation concern. Dalea cylindriceps is not

a plant of the Andes, and this common name may have had its origin in the binomial D. macrostachya Moric.,

which is similar to P. macrostachyum Tore, the first validly published name for what is now recognized as D.

cylindriceps. Dalea macrostachya is a replaced synonym for D. coerulea (L.f.) Shinz & Thell. var. longispicata

(Ulbr.) Bameby, which occurs in the Andean region of South America (Bameby 1977).

Several other common names have been adopted for D. cylindriceps that point to the dense, elongate spike-

like inflorescence which is a distinctive morphological feature of this species. These include “dense-flowered

prairie clover” (Britton & Brown 1897; Saunders 1899), “massive spike prairie-clover” (GPFA 1986), and

“large-spike prairie-clover” (Kaul et al. 2011; Stubbendieck & Conard 1989). The latter two names reflect Bar-

neby’s (1977) etymology of cylindriceps: “of the massive spike.”

The common name “sandsage prairie-clover” is proposed here based on the close association of D. cylin-

driceps with sandsage prairie and similar plant communities in which sand sagebrush (A. filifolia) is a domi-

nant or distinctive element.

acknowledgments

I would like to think several individuals who provided assistance with this research. David Sutherland, profes-

sor emeritus at the University of Nebraska at Omaha and author of the forthcoming treatment of the genus

Dalea for Flora of North America, reviewed the manuscript and brought to my attention pertinent information

in the Rydberg diary at NEB. Brad Elder and Rob Wikel, associate professor ami professor en.ert.us, respec-

tively, at Doane College (Crete, Nebraska), pnrvkfed information on die publications of Goodwm D. Swezey

and access to his botanical collections at Doane College. Robert Kaul, curator and research professor with the

Charles E. Bessey Hetbarium (NEB) of the Unhersity of Nebraska, provide . photocopy of ^1891

pamphlet, Nebraska Flowering Plants. Steven Rolfsmeier, director of the rihgh Hams i Herbanum «SOfl

Chadron State College (Chadron, Nebraska), shared his observations on die ecology of D. cylmdnccps and also

reviewed the manuscript.

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POTENTIAL DISTRIBUTION MODELING OF

PENSTEMON OKLAHOMENSIS (PLANTAGINACEAE)

J A Messick

Department of Geography and

100 E. Boyd St., SEC Suite 510

University of Oklahoma

Norman, Oklahoma 73019, U.SA.

B.W. Hoagland

University of Oklahoma

111 £ Chesapeake St.

Norman, Oklahoma 73019, U.SJL

bhoagland@ou.edu

ABSTRACT

INTRODUCTION

Advances in geospatial technologies and increased accessibility of species collection data have placed the map-

ping of species’ geographical ranges at the forefront of biogeographic research and conservation planning

(Guisan & Thuiller 2005; Franklin 2009; Peterson et al 2011). Location information from natural history col-

lections has become prevalent online in large databases (Elith et al. 2006; Newbold 2010; Peterson et aL 2011),

allowing for easier mapping of a species’ current and historic localities (Graham et al. 2004; Newbold 2010;

Peterson et al. 2011). Likewise, efforts to collect precise location data, with the assistance of GPS, also have

improved in recent years. Models of species distribution have provided insight into, generated hypotheses

about a species’ ecology, and aided in the location of new populations (Austin 2002; Hirzel et al. 2002; Guisan

& Thuiller 2005; Franklin 2009; Lobo et al. 2010; Newbold 2010; Naimi et al. 2011).

The objective of this study was to develop a predictive range map for the distribution of Penstemon oklaho-

mensis Pennell (Plantaginaceae). The genus Penstemon contains approximately 237 species and is one oi the

largest plant genera in North America (Freeman In prep; Lindgren & Wilde 2003; Nold 1999). Although P.

oklahomensis is one of 13 species of Penstemon that occur in Oklahoma, it is a unique regional endemic to the

southern plains region. It has been documented in 24 Oklahoma counties (Hoagland etal. ^ n ct ’

known populations were restricted to central Oklahoma until the recent discovery of a population in North

Texas (Mink et al. 2010). Penstemon oklahomensis is a native perennial that flowers from April to midjune and

■s °ie olonlv four species ol Penman with a closed .h, oat Horal morphology OI .hose four spcqK, ok laho-

mensis has the nJrestricted distribution (Clements et al. 1998; Pennell 1935). Penstemon oWohomensn most

892

frequently occurs in remnant Tallgrass prairie, but has also been found in other prairie types as well as open

woodlands (Hoagland et al. 2012). The Oklahoma Natural Heritage Inventory tracks P. oklahomensis as a state

rare species (SI). At the global level, it is ranked as a G3 (either very rare and local throughout its range or found

locally, even abundantly at some of its locations) and S3 (rare and local distributed within Oklahoma) (Okla-

homa Natural Heritage Inventory 2012).

This project is also intended to contribute to our understanding of the ecology of P. oklahomensis, for

which there are few published studies. A recent study of Penstemon oklahomensis habitat, indicated the soils

where populations occur ranged from sandy loam to loam with a pH range of 5.5-7.6 and relatively low nitro-

gen, phosphorous, and potassium levels. The same study also found P. oklahomensis populations to persist in

grassy roadside areas that are disturbed through various mowing regimes (Messick & Hoagland 2012).

Study Area

The study area encompasses the state of Oklahoma, although populations of P. oklahomensis have not been

documented in western parts of the state, congeners occur in all regions. We took this approach to determine

the regional extent of potential habitat. The long axis of Oklahoma has an east-west orientation that spans 6.5

degrees of longitude (from 94°30'W to 103°W) and 3.25 degrees of latitude (33°30’N to 37°N). Along this axis

northwest, at Black Mesa, to 110 m in the southeast, where the Little River exits the state into Arkansas. Aver-

age annual precipitation also follows a northwest to southeast gradient, with the lowest values in the northwest

(43 cm) and the highest in the southeast (142 cm). There is a weak south-north gradient in temperature. The

length of the growing season ranges from 225-230 days along the Red River and 175 days on the border with

Kansas. Average annual temperature increases roughly from 13.3°C in the northwest to 16.7°C in the southeast

(Johnson & Duchon 1995).

The successful analysis of species distribution relies upon the compilation of numerous layers of geospatial

data. The primary dataset for such analyses is location data, preferably in a geographic coordinates, derived

from specimen data. Location data for P oklahomensis

Oklahoma Vascular Plants Database (OVPD) (Hoagland et al. 2012), the Oklahoma Natural Heritage Inventory

(ONHI) (Oklahoma Natural Heritage Inventory 2012), and other sources (Freeman 1981). As noted earlier, a

population of P. oklahomensis has been reported from northeastern Texas, which is 51.5 km from the nearest

Oklahoma population (Mink etal. 2010), and was excluded from this analysis due to a lack of detailed informa-

tion on the population in question and access to geospatial data for Texas. We recognize the importance of this

population, however, and encourage the exploration of intervening areas between the Texas and Oklahoma

Once extracted, location data were compiled into a geodatabase and edited to remove duplicate records.

Duplicate records were found primarily in the OVPD and are a byproduct of specimen exchanges between in-

state institutions. Duplicate records also existed between the OVPD and the ONHI database. Next, geographic

precision of the records was assessed. Geographical coordinates were not provided on the majority of herbari-

um vouchers predating 2000, but either driving directions and/or land survey references (e.g. , township, range,

and section) were recorded. Thus it was necessary to manually assign geographical coordinates. Specimens

that listed only the county or equally vague geographic reference (e g., Indian Territory) were excluded from

analysis. The resulting dataset for analysis consisted of 142 location points (Fig. 1).

When mapping species distributions, it is important to examine both the extent of occurrence (EOO) and

area of occupancy (AOO). The EOO represents the entire area in which a species has been found, including

gaps between populations, and is bounded by the outermost occurrences of a species. The gaps between popu-

lations may simply represent inadequate sampling effort or are possibly areas of unsuitable habitat. The EOO

for a species can be mapped using the convex hull operation. The resulting map is a more accurate depiction of

a species distribution than one created using rectangles or circles encompassing all known locations of a spe-

893

cies (Podani 2009). The AOO is the area where the target species is actually found and is either equal to or

smaller than the EOO. The AOO does not include gaps between populations. Removing gaps or discontinuities

in an EOO results in the AOO for a species (Gaston & Fuller 2009).

The EOO map of P. oklahomensis was generated using the Convex Hull module of ArcGIS 10 using the

species occurrence data layer. Upon inspection, the resulting map exhibited a significant gap in the species

distribution between central and southwestern Oklahoma. Thus, we repeated the convex hull operation so

that the southwestern Oklahoma collection points (n=15) were aggregated into a polygon separate from a

larger central Oklahoma area polygon.

Maximum entropy (MaxEnt), the method most frequently used in species distribution modeling, was

chosen for modeling the distribution of P. oklahomensis. (Franklin 2009; Nairn! et al. 2011). MaxEnt was de-

signed specifically for use with presence-only data, such as the P oklahomensis dataset, and can analyze small

sample sizes (< 100 samples) and overcome sampling bias (Franklin 2009).

MaxEnt analyzes species occurrence data in conjunction with a suite of environmental data to calculate

an index of relative suitability for a species (Graham et al. 2008; Anderson & Gonzalez 201 1; Elith et al. 201 1).

Environmental factors are independent variables and are referred to as covariate or predictor variables. The

environmental variables used in this study were elevation, slope, aspect, land cover type, soU order, soil senes,

geology mean minimum annual temperature, mean maximum annual temperature, and mean annual precipi-

tation (Table 1). Slope and aspect were derived from a 30 m DEM using ArcGIS Spatial Analyst Toolbox and

then clipped to the political boundaries of Oklahoma. All of the remaining environmental variables were ac-

quired as vector data and were converted to raster format to match the extent and scale of the DEM.

MaxEnt attempts to derive a log-linear model that is dependent on the presence points and a set of se-

lected randomly from the environmental data layers, referred to as background points, to esthnate the proba-

bility of an occurrence or population in a locality. Sh<

background point, then the en

ence points because the presence poim

pecies presence point be selected as a

ire rescaled on a scale of 0-1, and an error boundary for

calculated from the environmental features rather than the pres-

iften biased (Elith et al. 2011).

it of MaxEnt is a probability of species occurrence b

ntropy .

TabuI.

Soil Series Associatioi

whether a pattern of occurrence is uniform across the landscape given the environmental variables used in the

model. A model is selected from replicates that have the highest test area under the curve (AUC) (Elith et al.

2006; Phillips et al. 2006; Franklin 2009; Elith et al. 2011; Warren & Seifert 2011).The area under the curve

(AUC) of a receiver-operating characteristic (ROC) plot is a threshold-independent metric (Franklin 2009;

Jimenez-Valverde 2012). A ROC plot graphs “the false-positive error rate on the x-axis (1 - Specificity) versus

the true positive rate on the y-axis (Sensitivity) based on each possible value of threshold probability” (Frank-

lin 2009). The AUC is calculated from the resulting curve and can range from 0.5 to 1.0. The value 0.5 repre-

sents random predictions while values above 0.5 represent “performance better than random” (Franklin 2009;

Jimenez-Valverde 2012). An AUC value between 0.5 and 0.7 indicates low or poor performance, between 0.7

and 0.9 indicates moderate performance, and values greater than 0.9 indicate high performance (Swets 1988;

Franklin 2009).

We used MaxEnt version 3.3.3e modeling software (www.cs.princeton.edu/~schapire/maxent) to model

the potential distribution of P. oklahomensis. The analysis was run with 0%, 10%, 20%, 30%, 40%, and 50% of

the P. oklahomensis point locations withheld for testing the model. Collectively these model runs were called

Model Set A. For each percentage category for which points were withheld, 15 replicates were generated. Re-

sponse curves, jackknife of variable importance, and maps of predicted distributions were also generated. The

jackknife of variable importance identifies the individual variable(s) that were most important in predicting

the species’ distribution (Elith et al. 2011). In order to evaluate the potential outlier affect of the P oklahomensis

occurrence in eastern Oklahoma, the analysis was conducted a second time and the resulting models were re-

ferred to as Model Set B.

MaxEnt created grids for the average, minimum, maximum, median, and standard deviation of the pre-

dictions for each percentage category withheld run based on the test AUC value. The average prediction grids

were converted to raster files and the resulting prediction AUC values were then compared. Based on the pre-

diction maps, the gap in the distribution between the southwestern populations and the central populations

was surveyed for P oklahomensis populations. This area included three counties; Grady County, Stephens

County, and Jefferson County. If a P oklahomensis population was discovered, voucher specimen was collected

and deposited at the Robert Bebb Herbarium (OKL) at the University of Oklahoma Norman OK

RESULTS AND DISCUSSION

:ounty-level map of the 142 Penstemm oklahomensis (Fig. 1) location points revealed that the majority of

:tton points were both in central Oklahoma and clustered near interstate or state highways. To verify this

m, we calculated Motaris I (I), which proved a significant pattern (1 . 0.371 z score = 3 385 p = 0.001)-

the AOO for this species was much smaller in area than the EOO and as noted earlier there were two

vonhy gaps in the EOO; the first in the southeast and a second in the southwest (Fig. 2).Since the gap in

juthwest was more pronounced geographically and was represented by a greater number of occurrences

(n=15) than the southeast (n=l), it became the focus of our analysis and groundtruthing. Our goal was to ascer-

tain whether this was a true gap in distribution or a sampling artifact.

The training and test AUC values for both model sets are listed in Table 2. Model 40 A had the highest

training AUC (0.954) while Model 10 A had the lowest training AUC (0.944). For the test data, Model 10 A had

the highest AUC (0.907) and Model 50 A had the lowest AUC (0.889). From the test data used in Model 10 A,

the jackknife of the environmental variables showed geology (25.6% contribution) and soil series association

(20.6% contribution) to be the most important (Table 3). Model 50 B had the highest training AUC (0.953) and

Model 0 B had the lowest training AUC (0.943). For the test data. Model 20 B had the highest AUC (0.900) and

Model 30 B had the lowest AUC (0.886). The jackknife of environmental variables from the test data used in

Model 20Balso showed geology (27.7% contribution) and soil series association (22.2% contribution) to be the

most important (Table4). Model 20B(Fig. 3) was selected as the best predictive map for the potential distribu-

tion of P. oklahomensis within Oklahoma because of its AUC value even though the value is the cut-off value

(0.900) between moderate and high performance according to Swets’ scale (1988).

MaxEnt predicted greater than 25% probability of occurrence of P. oklahomensis populations in extreme north-

ern Grady County and another in southern Grady County, a location within the southwesterndistnbution gap.

The two predicted northern locations were surveyed, but one proved to be a wheat field and the other a grazed

pasture. A new population (Fig. 4) of P oklahomensis was located, however, at the southern location where the

model redicted 25%-49% probability of occurrence. Another locality within the portion of Stephens County

in the gap with a greater than 25% probability of an occurrence was surveyed, but no populations were found.

A predicted location in northern Stephens County, with less than a 25% probability of occurrence, did produce

a new population (Fig. 4). Additional surveys of eounties in the southwestern gap dtd no. y«ld new popula-

^^T^poinllocafrons of die new populations were added to die overall point distributkm map and

■cted from the model. The Grady County population probability valuers 0.21 and the value

is 0.03. The two new location points were added to the same dataset used

with the same settings to produce Model Set C. The AUC

ability values e:

to produce Model Set B, and MaxEnt r

in importance values for Model 10 A.

values of Model Set C s

ues of all models i

the new Station points to the dataset lowered the AUC val-

era piv ~>ntrary to expectation.

CONCLUSION

“ E ‘T * b “ Kr “ nd 7'\ ndin « of f »°- controlling the d.s.nbuuon of oklaho-

mensis across its geographic range. Data for this effort was collected from Freeman (1981), QVP D (Hoagland et

898

Tabu 5. AUC values for Model Set C.

al. 2012), and ONHI. Our initial observation was that the highly clustered distribution pattern of P. oklahomen-

sis was a result of collector bias, which was correlated with ease of access due to roads. Further, collection loca-

tions were also clustered near cities with universities and in recreational areas.

We then analyzed the relationship of the distribution to environmental factors using MaxEnt. The result-

ing maps of potential probability of occurrence AUC values were deemed accurate, particularly those of the

Model 20 B (AUC = value of 0.90), the cut-off value between moderate and high performance (Swets 1988;

Franklin 2009). Groundtruthing of this model results lead us to two new populations within a “gap” in the

range of R oklahomensis. The low probability of occurrence values for the two new populations, however, sug-

gest that the predictor values used in the model may not be specific enough to locate additional populations P.

oklahomensis in southwest Oklahoma. Choosing the appropriate scale and type of predictor variables might be

confounded by the fact that P. oklahomensis exhibits relatively broad ecological tolerances. Messick and Hoa-

gland (2012), for example, documented that the greatest abundance of individuals of P oklahomensis was as

likely to occur in highway medians as in pastures dominated by native grasses.

Future surveys for P oklahomensis should be conducted to further evaluate the performance of the model.

It is important to note that during this study, the Southern Plains were experiencing a severe drought, which in

turn affected the number of stems present in known populations of P. oklahomensis. Thus we suggest the fail-

ure to find new populations within predicted areas could have been partially the result of drought.

ACKNOWLEDGMENTS

We greatly appreciate the helpful reviews of Stanley Rice and an anonymous reviewer.

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RANGE EXPANSION OF PANICUM REPENS (POACEAE)

INTO CENTRAL TEXAS (U.S.A.) MAY THREATEN ENDANGERED SPECIES

Jeffrey T. Hutchinson

U. S. Fish and Wildlife Service

<an Marcos Aquatic Resources Center

500 £ McCarthy Lane

San Marcos, Texas 78666, U.SA

jeffrey_hutchinson@fws.gov

Robert B. Shaw

Department of Ecosystem Science and

RESUMEN

La extension hacia el oestede la hierba acu4ticamuy invasiva, Parucum repens (torpedograss), enel Rto San Marcos en el Condado Hays, Texas

Panicum repens L., torpedograss, is a C 4 grass native to Europe, Asia, and Africa (Hossian et al. 1999). The

species is considered one of the world’s most aggressive grass weeds in agriculture (Holm et aL 1977) and natu-

ral areas (Langeland et al. 2008). It is widely naturalized throughout the New World tropics and subtropics

(Sutton 1996; Langeland et al. 2008). The species is currently distributed throughout the southeastern United

States, California, and Hawaii (Langeland et al. 2008). In Texas, the species is documented from herbarium

records in six counties (Cameron, Chambers, Galveston, Jefferson, Matagorda, and Trinity) in the eastern part

of the state (Shawetal. 2011; Shaw 2012). An additional voucher specimen ofR repens exists from Montgomery

County (Roger W. Sanders 6283, TEX 00207360) and un-vouchered observations are recorded for Calhoun

and Harris counties Qason Singhurst, unpubl. data). It is fisted as a noxious weed ^ Ae^Departmen^f

Agriculture (TDA 2013) and a prohibited exotic species by the Texas Parks and Wildlife Department (TPWD

2013).

perennial gras ilia, can grew io heighta of 1 merer. It Is mat-taring in water spreading

iron, an extensive network of rhizomes and stolons. The common name refers to •^a^pormed torpedo-

like tips of therhizomes. Panicum -epens is known to form extensive mars m water (16 to 1.2 meteis deep dui-

placing native aquatic plants (Tarver 1979). Rhizomes and stolons can grow to lengths of 6 meters (Langeland

1998) The svecies invades a wide variety of habitats and can be found growing in aquatic, riparian, wetland.

MmrLst^tata^T^theasmL United States, but thrives in wetland and riparian

establish in —

(Sartain 2003). Notwmtive grasses may have die ability to alter regional and even globalaspectsofeco^mteni

function (D Antonlo & Vitousek 1992). Being a C 4 grass, P. repens would have an advantage over na ,

902 Journal of the Botanical Research Institute of Texas 7 ( 2 )

grasses, such as the endangered Zizania texana Hitch., by its ability to sequester limited C0 2 during droughts,

low flows, and high water temperatures without photorespiration (Keeley & Rundel 2003). In Florida, P. repens

became established in over 5,665 ha (14,000 acres) in Lake Okeechobee and changed the structure and compo-

sition of the marsh forming a monoculture (Schardt 1994). Seed germination rates are highly variable, but

n, Extension and spread of Panicun

higher rates were reported for P. repens seeds under fluctuating temperatures (Martinez et al. 1992). Based on

the current literature, it is unlikely that P. repens seeds have high germination rates in the southeastern United

States (Wilcutetal. 1988).

The San Marcos River is spring-fed and supports a high diversity of threatened and endangered species

including Z. texana, Eurycea rathbuni Stejneger 1986 (Texas blind salamander), Eurycea nana Bishop 1941 (San

Marcos salamander), Etheostomafonticola Jordan & Gilbert 1886 (fountain darter), and Heterelmis comalensis

Bosse, Tuff & Brown 1988 (Comal Springs riffle beetle). An endemic species of fish Gambusia georgei Hubbs &

Peden 1969 (San Marcos gambusia) once found in the river is thought to be extinct. The upper 7.2 km of the San

Marcos River, from the headwaters at Spring Lake to the confluence with the Blanco River, is considered to be

one of the most diverse spring runs in Texas and is designated as critical habitat by the United States Fish and

Wildlife Service (USFWS 1996). Classification as critical habitat indicates a geographical area has all the at-

tributes needed for long-term success of endangered species’ but may require special management and protec-

tion measures to ensure species long-term survival. Threats to listed species in the upper San Marcos River

include dams, siltation, floods, decreased aquifer levels, low flows, recreation, and non-native species (Terrell

etal. 1978; USFWS 1996).

We believe that the similarity in habitat shared by P. repens and Z. texana is cause for concern. The most

upstream population of P. repens was observed less than 1 km from critical habitat and ca. 2.3 km from the

nearest population of Z. texana (Fig. 1). Based on our observations of P. repens just below Cumming’s Dam, the

species exhibits the ability to form monocultures along littoral and riparian habitat, as well as spreading into

uplands. This species has the ability to spread into critical habitat occupied by Z. texana from accidental or

natural movements of small stem fragments, rhizomes, and stolons upstream. In greenhouse studies, small

sections of P. repens stems with nodes produce roots in 1 day, and 79% of tiller segments and 93% of shoot seg-

ments produced new vegetative growth within four weeks (Sartain 2003). Panicum repens and Z. texana both

prefer open sunlight and reproduce vegetatively by rhizomes or tillers. Zizania texana is found at a mean water

depth of 0.75 m (Poole & Bowles 1999) which lies within the range where P repens is documented to form

The presence of P. repens just outside of critical habitat and the possibility that it could be introduced

further upstream poses a threat to native species of flora and fauna in the San Marcos River. The establishment

of P. repens within critical habitat, combined with the effects of other established non-native plants such as

Hydrilla verticillata (L.f.) Royle (hydrilla), Hygrophila polysperma (Roxb.) T. Anderson (East Indian hygrophi-

la), Cryptocoryne beckettii Thuill. ex R. Trim (Beckett’s water trumpet), and Colocasia escidenta (L.) Schott (wild

taro) could result in additional habitat loss for Z. texana , other native aquatic plants, threatened and endan-

gered species, and alteration of the habitat structure on which they are dependent for survival.

Voucher specimen: TEXAS. Hays Co.: San Marcos, San Marcos River (29°51'21"N,

ACKNOWLEDGMENTS

We thank employees at the USFWS’s San Marcos Aquatic Resources Center, Robert Doyle, Kenneth Unge-

land, and Jackie Poole for their review of this manuscript. Official journal reviews by Jason Singhurst and Da-

vid E Lemke are much appreciated. Jackie Fernandez and Javier Gomez generously provided the Spanish

translation of the abstract. The views expressed in this manuscript are the views of the authors and do not re-

flect the views of the U.S. Fish and Wildlife Service.

D'Antonio, C.M. AND P.M. VrrouSEK. 1

Ann. Rev. Ecot. Syst 23:63-87.

Holm, LG., D.L Plucicnett, J.V. Panch

’. Biological invasions by exotic grasses, the grass/fire cycle, and global change,

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Langeland, K.A. 1 998. Torpedograss— forage gone wild. Wildland Weeds 1 (3):4— 6.

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Florida's natural areas, 2nd edition. IFAS Communication Services, University of Florida, Gainesville.

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sowing often tropical dune species. Oecologia 92:343-353.

Poole, J. and D.E. Bowles. 1999. Habitat characterization of Texas wild-rice {Zizania texana Hitchcock), an endangered

aquatic macrophyte from the San Marcos River, TX, USA. Aquatic Conserv: Mar. Freshw. Ecosyst 9:291-302.

Sartain, R.T. 2003. Physiological factors affecting the management of torpedograss ( Panicum repens (L.) Beauv.). M.S.

Thesis. University of Florida, Gainesville.

Schardt, J.D. 1994. Florida aquatic plant survey 1 992. Technical Report 942-CGA. Florida Department of Environmental

Protection, Tallahassee.

Shaw, R.B. 2012. Guide to Texas grasses. Texas A&M University Press, College Station.

Shaw, R.B., B.S. Rector, and A.M. Dube. 201 1 . Distribution of grasses in Texas. Sida, Bot. Misc. 33. Botanical Research Insti-

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Sutton, D. 1996. Growth of torpedograss from rhizomes planted under flooded conditions. J. Aquatic PI. Managem.

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Tarver, D.P. 1 979. Torpedo grass ( Panicum repens L.). Aquatics 1 :5-6.

Terrell, R.E, W.H.P. Emery, and H.E. Beaty. 1978. Observations on Zizania texana (Texas wild-rice), an endangered species.

Bull.Torrey Bot. Club 10550-57.

TDA. 2013. Texas Department of Agriculture— noxious weed list, texasinvasives.org/plant_database/tda_results.

php?offset=20. Accessed 25 April 201 3.

TPWD. 2013. Texas Parks and Wildlife Department— Invasive, prohibited and exotic species, www.tpwd.state.tx.us/

huntwild/wild/species/exotic/prohibited_aquatic.phtml. Accessed 25 April 2013.

USFWS. 1996. United States Fish and Wildlife Service— San Marcos and Comal Springs and associated aquatic ecosys-

tems (revised) recovery plan. United States Fish and Wildlife Service, Albuquerque, New Mexico.

Wilcut, J.W., R.R. Dute, B.Truelove, and D.E. Davis. 1988. Factors limiting the distribution of cogongrass {Imperata cylindrica)

and torpedograss ( Panicum repens). Weed Sci. 36:577-582.

ANNOTATED CHECKLIST OF THE VASCULAR FLORA

OF THE WIND RIVER RANGE, WYOMING (U.S.A.)

Walter F. Fertig

Robert T. Massatti

Arizona State University Herbarium

School of Life Sciences, PO Box 874501

Tempe, Arizona 85287-4501, U.S.A.

wfertig@asu.edu

University of Michigan Herbarium

3600 Varsity Drive

Ann Arbor, Michigan 48108, U.SA.

B.E. Nelson

Ronald L. Hartman

Rocky Mountain Herbarium

Department of Botany, Dept 3165

University of Wyoming

1000 E University Ave.

Laramie, Wyoming 82071, U.S.A.

Rocky Mountain Herbarium

Department of Botany, Dept 3165

University of Wyoming

INTRODUCTION

The Wind River Range is located at the southern edge of the Central Rocky Mountains (Peet 2000) and Greater

Yellowstone ecosystem in northwestern Wyoming. These mountains, “named for flowing water named for

moving air” (Kelsey 1988) are the largest and highest range in the state (Blackstone 1993). More than 40 peaks

exceed 3800 m (12,500 ft). The tallest, Gannett Peak at 4208 m (13800 ft), is the highest point in the Rocky

Mountains between Canada and Colorado (Kelsey 1988). Scattered among these peaks is the second largest

concentration of glaciers within the contiguous United States and one of the most extensive areas of alpine

tundra (Scott 1995).

In June 1834, Thomas Nuttall collected the holotype of Eriogonum acaule at South Pass at the southern

edge of the Wind River Range, making him the first botanist to study the flora of the area (Dorn 1986). More

than 100 plant collectors have followed in Nuttall’s footsteps, including John C. Fremont, Aven Nelson, Elmer

D Merrill Edwin B Payson Cedric L. Porter, Frederick J. Hermann, Rupert C. Bameby, Reed C. Rollins, and

906 Journal of the Botanical Research Institute of Texas 7(2)

Erwin F. Evert. Since the mid-1960s, seven graduate students from the University of Wyoming’s Rocky Moun-

tain Herbarium (RM) have conducted floristic surveys of portions of the range. The purpose of this paper is to

compile an annotated checklist of the vascular flora of the Wind River Range based on a synthesis of the exist-

ing literature and specimen databases of the RM (Hartman et al. 2009) and other regional herbaria. The paper

will also highlight the biogeographic patterns and conservation significance of the area. The inventory of the

Wind River Range is part of a larger effort by RM to document the flora of the middle and southern Rocky

Mountains over the past 35 years (Hartman 1992) that has generated more than 50 master’s theses and over

650,000 herbarium collections.

METHODS

The species list (Table 1) was derived from a review of unpublished floristic theses, natural area inventory re-

ports, and rare species surveys conducted in the Wind River Range since the 1960s (Cramer 1997; Evert 2010;

Fertig 1992, 1995a, 1995b, 1997, 1998; Haines 1988; Kinter 2000; Massatti 2007; Mills & Fertig 1996; Newton

2008; Rosenthal 1999; Scott 1966, 1995). Additional reports were found in the collections of the RM and Cen-

tral Wyoming College (CWC) and records from the Wyoming Natural Diversity Database. At least one collec-

tion representing each species from the study area was re-verified at RM to corroborate its presence. Misidenti-

fied or falsely reported species from the Wind Rivers are listed in Table 2. Species nomenclature follows the RM

Plant Specimen Database (Hartman et al. 2009), which itself is based on Dorn (2001) and more recent literature

(including Barkworth 2003, 2007; Ertter 2007; Flora of North America Editorial Committee 1993+; Holmgren

et al. 2005, 2012; Mast & Reveal 2007; Nesom 2006; Wagner et al. 2007). Family names are based on the An-

giosperm Phytogeny Group (2009) classification. Each species entry in the checklist is annotated with addi-

tional information on the number of collections from the area, distribution within the Wind Rivers (by subre-

gions), elevation range, and general habitat types. Synonyms are provided for names that differ from Dorn

(2001), Fertig (1992), or Massatti (2007). Additional codes indicate species that are historical, introduced, or of

Setting. The Wind River Range straddles the Continental Divide from Togwotee Pass to South Pass, a distance

of 177 km (110 miles). At its northern end, the Wind Rivers are bordered by the Absaroka and Gros Ventre

ranges and the valley of the Buffalo Fork of the Snake River. To the east, south, and west, the range is bounded

by the Wind River, Great Divide, and Green River basins, respectively (Fig. 1). The entire range covers an area

of approximately 7800 square km (3010 square miles). For the purpose of this study, we excluded the Brooks

Lake area and Dunoir Valley northeast of U.S. Highway 26/287, which is included in the Wind Rivers by some

authors (Rosenthal 1999), because it is better considered part of the Absaroka Range.

The area is divided between Fremont, Sublette, and Teton counties. More than 80 percent of the Wind

River Range is managed by the U.S. Forest Service. The area west of the Continental Divide is located within

Bridger-Teton National Forest, while the lands to the east are part of Shoshone National Forest. More than half

of the Forest Service lands are within three Congressionally-designated Wilderness Areas: Bridger, Fitzpat-

rick, and Popo Agie. About one-quarter of the east slope of the range is in the Wind River Indian Reservation.

The foothills of the Wind Rivers include small parcels of private, state, and federal lands managed by the Bu-

reau of Land Management (BLM). Several small towns are located in the foothills of the range, including Pine-

dale on the west side. Lander off the southeast slope, and Dubois along the northeast boundary.

Climate. The Wind River Range lies in a partial rain shadow created by the Teton, Gros Ventre, Salt River,

and Wyoming ranges to the west and southwest. The higher elevations of the Wind Rivers receive 132-152 cm

of precipitation per year, with about 65 percent falling as snow (Manner 1986; Potkin 1991). Precipitation is

greatest at the northern end of the range and decreases southward (PRISM Climate Group 2007). Annual pre-

cipitation in the foothills averages 20-38 cm. Most precipitation occurs in April and May, with a secondary

peak in September and October (Western Region Climate Center 2007).

910 Journal of the Botanical Research Institute of Texas 7(2)

The modem terrain of the Wind River Range has been sculpted by Pleistocene glaciation. In the last

200,000 years there have been at least three major glacial advances (Lageson & Spearing 1988). The scouring

action of glaciers produced the large cirques, sharp-crested peaks, and U-shaped valleys common in the moun-

tains (Worl et al. 1986) as well as erratic boulders, moraines, and large glacial lakes in the foothills. Several

large glaciers still exist, mostly just east of the Continental Divide, and some of these show evidence of advanc-

ing in the past 100-2100 years (Mears 1997).

Vegetation. The vegetation of the Wind River Range and northwestern Wyoming has been described in

detail by Evert (2010), Gregory (1983), Habeck (1987), Knight (1994), Potkin (1991), Reed (1971, 1976), Tweit

and Houston (1980), Walford et al. (2001), Youngblood and Mueggler (1981), and Youngblood et al. (1985). We

recognize twenty-one general vegetation types for the Wind Rivers (Fertig 1992; Massatti 2007) based on dif-

ferences in physiognomy (forests/woodlands, shrublands, and forb/graminoid meadows), topography (slopes

or depressions), soil moisture, elevation zone (foothills, montane, subalpine, or alpine), and substrate (sand,

FORESTS AND WOODLANDS

Aspen forests.— Quaking aspen ( Populus tremuloides ) is common from the foothills to the upper subalpine

zones in the Wind River Range, especially in areas with a history of disturbance or on relatively calcareous or

mesophytic soils derived from sedimentary formations (Youngblood & Mueggler 1981). Common understory

species include Symphoricarpos oreophilus, Mahonia repens, Artemisia tridentata var. vaseyana, Arnica cordijolia,

Lupinus argenteus. Geranium viscosissimum, and Astragalus miser. Stands that are grazed by cattle or sheep often

have an understory dominated by Frasera speciosa, Hymenoxys hoopesii, Arctostaphylos uva-ursi, and Juniperus

communis (Reed 1971; Youngblood & Mueggler 1981). Aspen stands are usually serai to other coniferous forest

types, though climax aspen forests may occur in a band between low elevation sagebrush grasslands and

Douglas-fir or spruce-fir forests in the Wind Rivers (Reed 1971).

Douglas-fir forests. — Forests dominated by Douglas-fir ( Pseudotsuga menziesii var. glauca ) in the Wind

River Range occur primarily on steep, north exposures in the foothills and lower montane zones on limestone

or sedimentary soils (Reed 1976). Spruce-fir forests typically replace Douglas-fir on higher elevation acidic

soils derived from gneiss or granite. Common understory species include Juniperus communis, Mahonia repens,

Symphoricarpos oreophilus, Poa wheeleri, and Festuca idahoensis (Steele et al. 1983). On drier sites, Douglas-fir

forests intergrade with big sagebrush grasslands and limber pine woodlands.

Forest wetlands.— This broad category includes swamps, lakeshores, and streamsides dominated by En-

gelmann spruce ( Picea engelmannii), blue spruce (P. pungens), narrowleaf cottonwood (Populus angustifolia ),

quaking aspen, or other tree species (Fertig 1992). Forested wetlands differ from upland forest types by occur-

ring on wetter and more poorly-drained soils (usually in drainage bottoms) and by often having a richer herba-

ceous understory. Typical understory species include Carex microptera, Calamagrostis canadensis, Bromus cili-

atus, Senecio triangularis, Saxifraga odontoloma, Mertensia ciliata, and Equisetum arvense (Massatti 2007).

Juniper woodlands.— Relatively open canopy woodlands dominated by Utah juniper (Juniperus osteo-

sperma) or Rocky Mountain juniper (J. scopulorum) are restricted to the Red Canyon area along the southeast

flank of the Wind River Range (Massatti 2007). This vegetation type occurs on siltstone and sandstone-derived

soils. Common understory species include Artemisia tridentata, Leucopoa kingii, Purshia tridentata, and Rhus

trilobata (Massatti 2007).

Limber pine woodlands. — Limber pine ( Pinusjlexilis ) forms open woodlands on dry slopes and ridges in

the foothills of the Wind River Range (Steele et al. 1983). Stands usually occur on south exposures over well-

drained, rocky, sedimentary soils. Common associated species include Shepherdia canadensis, Juniperus com-

munis, Symphoricarpos oreophilus, Leucopoa kingii, Carex rossii, and Festuca idahoensis.

Lodgepole pine forests. — Forests dominated by lodgepole pine (Pinus contorta var. latifolia) cover extensive

areas of the foothill, montane, and subalpine zones of the Wind River Range. Lodgepole pine may form nearly

pure stands following severe disturbances, especially fire. Stands are usually serai to other coniferous forest

types, but climax lodgepole pine forests can persist on dry, cool, or relatively infertile sites (Steele et al. 1983).

Fertig etal.,l

Serai and climax lodgepole pine forests tend to have low species richness (Reed 1976). Typical species from the

understory include Juniperus communis, Vaccinium scoparium. Arnica cordifolia, Carex rossii, Poa wheeleri, Achil-

lea millefolium, Antennaria rosea, and Potentilla diversifolia. Calamagrostis rubescens and Carex geyeri, two spe-

cies that are often dominant in lodgepole pine communities elsewhere in western Wyoming, are absent or rare

in the Wind Rivers (Steele et al. 1983).

Spruce-fir forests. — Subalpine fir (Abies bifolia ) and Engelmann spruce (Picea engelmannii ) are the domi-

nant tree species in the montane and subalpine forests of the Wind Rivers. Abies bifolia is more likely to be

dominant (with R engelmannii being serai) on well drained, dry to mesic sites (Reed 1976; Steele et at 1983).

odic disturbance (Peet 2000; Steele et al. 1983). Potkin (1991) found the understories of different spruce-fir

habitat types to be floristically similar and relatively depauperate. Typical understory species include Vaccini-

um scoparium, Shepherdia canadensis, Juniperus communis. Arnica cordifolia, Antennaria microphylla, Carex ros-

sii, and Poa wheeleri. Calamagrostis rubescens, Carex geyeri, and Spiraea betulifolia are common understory

species in spruce-fir forests of the Teton and Wyoming ranges, but are rare to absent in the Wind Rivers (Steele

etal. 1983).

Whitebark pine forests.— Whitebark pine (Pirns albicaulis) occurs mostly from upper timberline to the up-

per montane elevation zone, where it is usually subdominant to subalpine fir and Engelmann spruce. Forests

dominated by whitebark pine are limited to ridges and slopes exposed to extreme cold, high winds, and

drought conditions unsuitable to spruce-fir forest (Reed 1976; Steele et al. 1983). Species commonly found in

these stands include Vaccinium scoparium. Arnica cordifolia, Carex rossii, Poa wheeleri, Juniperus communis, and

Ribes montigenum. These forests have been impacted by blister rust in the past decade.

SHRUBLANDS

Big sagebrush grasslands. — Shrub steppe dominated by mountain big sagebrush (Artemisia tridentata var.

vaseyana ) is the most widespread non-forested vegetation type in the Wind River Range. This type occurs on

upland sites from the foothills to the lower subalpine on well-drained, sandy, or coarse-textured soils (Knight

1994). Common associated species include Festuca idahoensis, Elymus spicatus, E. trachycaulus, Poafendleriana,

P. secunda, Achnatherum nelsonii, Koeleria macrantha, Purshia tridentata, and Ericameria nauseosa. Dry, rocky

slopes and talus in the sagebrush zone are often locally dominated by other shrub species, such as Acer gla-

brum, Amelanchier alnifolia, Cornus sericea, or Symphoricarpos oreophilus (Fertig 1992).

Riparian shrublands (non willow).— Riparian shrublands are a relatively minor vegetation type found

along streams, lakes, and moist, low-lying depressions, These communities are dominated by a mix of shrub

species that does not include willows or trees. The most common dominants are Betula occidentals, B. glandu-

losa, Cornus sericea, Lonicera involucrata, and Prunus virginiana. Typical understory species include Equisetum

arvense, Heracleum sphondylium, Poapratensis, Calamagrostis canadensis, and Carex microptera (Fertig 1992). In

some low elevation areas silver sagebrush (Artemisia cana var. viscidula ) may dominate streambank communi-

ties, with or without Dasiphora fruticosa (Youngblood et al. 1985).

Willow thickets — Willow (Solixspp.) thickets are the most widespread riparian shrub vegetation type in

the Wind River Range (Walford et al. 2001; Youngblood et al. 1985). Willow stands occur from the foothills to

above timberline and share a common physiognomy, but differ in the dominant species of Salix. At lower eleva-

tions, stands are dominated by Salix boothii, S. wolfii, or S. geyeriana with an understory of Deschampsia cespi-

tosa, Geum macrophyllum, Agrostis stolonifera, and Poapratensis. In the subalpine zone, Salix planifolia (or oc-

casionally S eastwoodiae) becomes predominant (Walford et al. 2001). Willow communities in the alpine tend

to be dominated by Salix glauca on well-drained mineral soils and S. planifolia on wetter sites (Potkin 1991).

r treehne on the highest peaks of the Wind

River Range. This vegetatii

Among the more comr

i type is similar to the dry meadows characteristic of subalpine and montane zones,

cushion-forming forte, dwarf shrubs, and low bunchgrasses (Scott 1966, 1995).

species are Geum rossii, Salix arctica, S. reticulata, Selagindla densa, Silene acaulis.

912

Journal of the Botanical Research Institute of Texas 7(2)

a woodlands of stunted Picea engelmannii,

(Fertig 1992).

Aquatic . — Aquatic vegetation consists of free-floating, submerged, or emergent forb and graminoid spe-

cies found within lakes, ponds, and slow-moving streams from the foothills to the subalpine of the Wind River

Range (Fertig 1992). This minor vegetation type is dominated by Potamogeton and Stuckenia spp., Callitriche

palustris. Ranunculus aquatilis, Carex utriculata, Sparganium angustifolium, and Alopecurus aequalis.

Badlands. — Sparsely vegetated badland communities are a minor type found sporadically in the northern

foothills of the Wind River Range on open, highly eroded, clay-rich slopes and drainage bottoms (Fertig 1992).

Common species include Pyrrocoma uniflora, Lomatium tritematum, Phlox multiflora, Oenothera cespitosa, As-

tragalus kentrophyta, and Antennariamicrophylla.

Disturbed sites. — Disturbed vegetation may be recognized by the dominance of non-native forb and

graminoid species (Fertig 1992). In the Wind River Range, this vegetation type occurs chiefly along roadsides,

trails, parking lots, or areas that have been logged, burned, or used heavily by livestock. Typical species include

Bromus inermis, B. tectorum, Phleum pratense, Poa pratensis, Taraxacum officinale, T. erythrospermum, Capsella

bursa-pastoris, Cirsium arvense, Thlaspi arvense, and Tragopogon dubius.

Dry meadows. — Dry meadow communities are park-like openings in upland sites from the foothills to

upper treeline dominated by perennial grasses and forbs with little to no mountain big sagebrush. These sites

often occur on relatively well-drained, fine-textured alluvial or colluvial soils with a dense sod that inhibits the

establishment of forests (Peet 2000). Common species include Festuca idahoensis, Elymus spicatus, E. trachy-

caulus, Lupinus argenteus, and Ligusticum filicinum. Dry meadows in the Range with a history of sheep grazing

tend to become dominated by Achnatherum lettermanii, Trisetum spicatum, Achillea millefolium, and Agoseris

glauca (Potkin 1991).

Granite or gneiss outcrops . — Exposed granite or gneiss bedrock, cliff faces, talus slopes, and boulder fields

are common along the Continental Divide and can extend into the foothill zone on the west side of the Wind

River Range. These sites are frequently disturbed by rock slides and have shallow, xeric soils due to exposure

to high winds (Gregory 1983). Vegetative cover is sparse and consists mostly of bunchgrasses or forbs (sites

with high sagebrush or montane shrub cover are classified as big sagebrush grasslands). At lower elevations,

granitic outcrops tend to be dominated by Balsamorhiza sagittata, Eriogonum umbellatum, and Elymus trachy-

caulus. At upper treeline and above, granite outcrops are commonly inhabited by Oxyria digyna, Erigeron

Limestone or calcareous outcrops. — Cliffs and talus fields derived from limestone sedimentary rocks occur

along the east flank of the Wind Rivers and on high peaks adjacent to the Green River Lakes on the west slope.

As with granitic outcrops, this community type is characterized by low cover of bunchgrasses and perennial

forbs and often lacks shrub or tree species. A suite of calceophilic plant taxa (including many rare species) are

restricted to limestone outcrops, such as Dryas octopetala, Saxifraga oppositifolia, Saussurea weberi, Parrya nu-

dicaulis, Antennaria aromatica, and Boykinia heucheriformis.

Marshes and bogs. — Marshlands dominated by sedges or grasses occur along waterways in valley bottoms

and adjacent to shallow lakes and ponds in the foothills and montane zones of the Wind River Range. Sites with

saturated or flooded soils are often dominated by Carex utriculata (Walford et al. 2001), while drier sites may be

dominated by C. aquatilis (Youngblood et al. 1985). Other common marsh species include Calamagrostis ca-

nadensis, Deschampsia cespitosa, Poa pratensis, and Geum macrophyllum. Heath bogs occur in similar environ-

ments with a high water table in the suhalpine zone, but are dominated by low shrubs or forbs (Potkin 1991).

Typical bog species are Kalmia microphylla, Vaccinium occidentale, Deschampsia cespitosa, Juncus drummondii,

Pedicularis groenlandica, Caltha leptosepala, Salix planifolia, and S. glauca.

Sand and gravel bars. — This vegetation type occurs on sandy beaches, gravel bars, and mud flats adjacent

to lakes, rivers, and streams throughout the Wind Rivers (Fertig 1992). Sites typically have only a modest cover

of graminoids and forbs. Common species include Equisetum arvense, Arnica chamissonis, Rorippa curvipes.

Ranunculus flammula, Alopecurus aequalis, Agrostis scabra, and Carex athrostachya.

913

Wet meadows.— Wet meadows are found along moist streamsides, lakeshores, and floodplains or in low-

lying areas that accumulate drifting snow. Similar in appearance to dry meadows, wet meadows are dominated

by a thick turf of perennial graminoid and forb species, but relatively few shrubs or trees. Species richness

tends to be high in these stands (Tweit & Houston 1980; Youngblood et al.1981). Dominant species may in-

clude Erigeron glacialis, Caltha leptosepala, M ertensia ciliata, Senecio triangularis, Deschampsia cespitosa, Poa

pratensis, P. palustris, Calamagrostis canadensis, Juncus arcticus, and Carexmicroptem. Wet meadows intergrade

with dry meadows and alpine tundra but tend to support a different suite of species.

Geographic Subregions. Based on differences in climate, geology, and elevation, the Wind River Range can be

divided into geographic subregions that reflect differences in local floristic composition. Fertig (1992) recog-

nized five subregions on the west side of the Continental Divide and Massatti (2007) identified six from the

east side. We have modified these subdivisions slightly and revised their names to avoid duplication to derive

the following ten geographic subregions for the range (Fig. 1).

Boulder Creek drainage— Formerly called the South-Central region by Fertig (1992), this subunit extends

from Mount Victor and Burnt Lake on the west side of the Continental Divide to Rennecker Peak and the Pros-

pect Mountains. The area includes the south half of the Bridger Wilderness and the Boulder Creek, East Fork,

Big Sandy, and upper Sweetwater River drainages. This region is similar to the Fremont Lake subregion to the

north in being predominantly gneiss and granite, but has a progressively drier climate and different grazing

history (Potkin 1991).

Dinwoody Creek drainage. — This area, located east of the Continental Divide, was previously named the

North-Central Region by Massatti (2007). It includes the granite/gneiss core of the northeastern Wind Rivers

from Union Peak to Milky Ridge, but excludes the calcareous and sandstone exposures along the east flank of

the range (separated out as the Torrey Lake subregion). Most of this region is included within the Fitzpatrick

Wilderness Area of Shoshone National Forest.

Fremont Lake— Previously called the North-Central region by Fertig (1992), this subregion occupies the

high elevation granite-gneiss core of the west slope of the Range from the Green River divide and Union Pass

south to the Boulder Creek divide between Half Moon and Burnt lakes. The Fremont Lake region includes the

north half of the Bridger Wilderness and has a wetter cli

Green River Lakes.— This region includes the calcareous mountains (Gypsum Peak, Big Sheep Mountain,

and White Rock) and adjacent valley surrounding the Green River Lakes west of the Continental Divide. This

is the only area on the west slope of the range that has retained its original mantle of limestone caprock, in

sharp contrast to the high elevation gneiss and granite bedrock of the surrounding Fremont Lake region. The

flora of the alpine areas of the Green River Lakes shares many rare and disjunct species with Arrow Mountain

and other calcareous peaks in the Torrey Lake subregion east of the Divide.

Limestone Mountain.— Located along the southeast flank of the range and formerly known as the South-

east Region (Massatti 2007), the Limestone Mountain area is characterized by sedimentary formations compa-

rable to those along the northeast slope of the Wind Rivers in the Torrey Lake region, but mostly at a lower el-

evation and with a drier climate. Major features include Red Canyon (the lowest point in the Wind River Range

at 1725 m), Fairfield Hill, Limestone Mountain, and the Freak Mountains.

Moccasin Basin. The Moccasin Basin region is located along the northwestern flank of the Range west of

the Continental Divide and encompasses the Cottonwood Creek and Fish Creek drainages between Togwotee

Pass and the Green River divide. Most of the basin is comprised of relatively low hills (mostly under 3050 m)

with volcanic soils or Tertiary-age claystones and sandstones (Devils Basin, Wind River, and Pinyon forma-

tions). Alpine vegetation is restricted to the summit of Two Ocean Mountain, just south of Togwotee Pass

(Fertig 1992). . . _

Popo Agie River drainage. — Known as the South-Central r<

region contains the granite-gneiss core of the southern Wind River Range o

Divide. The area is at lower elevation and has a drier climate than tl

Much of the region is within the Popo Agie Wilderness Area and includes Wind River Peak, the Deep Creek

lakes. Cirque of the Towers, and glacial deposits around Louis Lake.

South Pass. — Recognized by both Fertig (1992) and Massatti (2007), this region extends from the south

end of the Prospect Mountains and Miner’s Delight to South Pass City and Atlantic City east of Wyoming state

highway 28. The entire area is located south of the boundary of Bridger-Teton and Shoshone national forests.

The region consists of rolling hills with soils derived from sandstone, siltstone, granodiorite, and metasedi-

mentary rocks. South Pass is essentially a broad ecotone between the core of the Wind River Range to the north

Torrey Lake. — Originally named the Northeast Region (Massatti 2007), this area encompasses the sedi-

mentary lower flanks of the east slope of the Wind River Range from Warm Spring Mountain and the foothills

west of Dubois south to the Bull Lake area of the Wind River Indian Reservation. This region includes outcrops

of the Madison and Gallatin limestones. Bighorn dolomite, Flathead sandstone, and Gros Ventre Formation

and alpine summits of Whiskey Mountain, Arrow Mountain, and Dinwoody Peak. The Torrey Lake region

shares floristic affinities with the Green River Lakes region on the west slope of the range and the drier Lime-

stone Mountain region to the south.

Warm Spring Creek drainage.— This area (previously known as the North Region by Massatti 2007) is lo-

cated on the east side of the Continental Divide from Togwotee Pass south to Union Peak and east to Dunoir

and the north flank of Union Peak. Most of the subregion is drained by Warm Spring Creek and is underlain by

volcanic claystones and basalts of Quaternary gravel, rather than Mesozoic sediments and Precambrian gran-

ites and gneiss typical of the regions to the south. Alpine habitats are restricted to the summits of Lava Moun-

tain and Union Peak.

RESULTS AND DISCUSSION

Species Richness. Based on our field work and review of records from RM, CWC, and the literature, there are

1282 vascular plant taxa known from the Wind River Range (Table 1). This total includes 1190 full species and

92 separate varieties or subspecies. At least 32 of these taxa have not been relocated since 1970 and are consid-

ered historical. The flora of the Wind Rivers represents 44.6 percent of the vascular flora of Wyoming (2875

taxa; Fertig 2011). These species belong to 407 genera and 91 families (APG 2009). Angiosperms comprise

nearly 98 percent of the taxa known from the Range.

An additional 62 species reported for the Wind River Range or vicinity (Fertig 1992; Haines 1988; Mas-

satti 2007; Newton 2008; Rosenthal 1999; RM records) have been excluded from our annotated checklist.

Twenty of these taxa are now considered synonyms of other species already known from the flora (these are

listed in synonymy in the annotated checklist). Another nine taxa were rejected because the specimens cited

for the Wind Rivers were collected beyond our boundaries. The remaining 33 were misidentified (Table 2).

The Wind River flora includes 99 non-native species that account for just 7.7 percent of the entire flora

(Table 1). By comparison, introduced species make up 13.6 percent of the flora of Wyoming (Fertig 2011)-

Fourteen of the 26 officially designated state noxious weed species occur in the Wind River Range (Wyoming

Weed and Pest Council 2012).

Since 1987, twelve first records for the state of Wyoming have been documented from the Wind River

Range. Richard Scott discovered Lathyrus eucosmus in the southeastern foothills in 1987 and Potentilla hyparc-

tica near the Continental Divide in 1988. Fertig (1992) found three new species records in 1990: Coronilla varia

and Tanacetum parthenium (both introduced weeds) and Erigeron lanatus (a rare alpine endemic). Exploration

of the Arrow Mountain area on the east slope in 1996 generated first records of Arnica angustifolia var. tomen-

tosa and Braya humilis by Fertig and Helictotrichon mortonianum by Hartman. Massatti discovered Carex len-

ticularis var. dolia and Massatti and Wells (2008) found Festuca viviparoidea ssp. krajinae in 2005-06. Nelson

added Muscari botryoides from the vicinity of Atlantic City in 2005. Brasher and Enloe (2007) documented

Rorippa austriaca from two sites near Cora along the western boundary of the study area in 2006.

Rare Species. The Wyoming Natural Diversity Database currently lists 82 species from the Wind River

Fertig et al., Vascularflora of Wind River Range, Wyoming

915

Range as species of concern or potential species of concern (Heidel 2012). No plant species from the Wind Riv-

ers are listed as Threatened or Endangered under the US Endangered Species Act (ESA), although two species

were added to the Candidate list for potential listing in 2011. Whitebark pine (Pinus albicaulis ) is a Candidate

due to impacts from white pine blister rust, mountain pine beetles, and fire suppression (Ashe 2011). Small

rockcress ( Boechera pusilla ), a South Pass endemic, was added to the list because of a significant population

crash (Gould 2011). Of the 82 species of concern, 30 are designated Sensitive by the US Forest Service or BLM

Wyoming state office (Heidel 2012). Sensitive species are given special management attention to prevent their

continued downward population trends and potential listing as threatened or endangered. Most of the Sensi-

tive species are Wyoming or Rocky Mountain endemics or arctic/boreal disjuncts and tend to occur in calcare-

ous alpine or wetland habitats (Fertig 1997, 1998).

Richness by Vegetation Types. Of the 21 main vegetation types recognized for the Wind River Range,

dry meadows have the highest species richness, with 693 taxa or 54 percent of the total flora (Table 1). Nearly

as species-rich are wet meadows (572 taxa) and big sagebrush grasslands (607 taxa). Among forested vegeta-

tion types, forested wetlands have the highest species richness with 384 taxa (30 percent of the flora), followed

by aspen forests with 332 taxa. Lodgepole pine forests, often considered species-poor, support a higher number

of plant taxa in the Wind Rivers (324 taxa) than spruce-fir forests (263 taxa) or any other conifer-dominated

vegetation type. Granite and gneiss rock outcrops have relatively high overall species richness with 409 taxa

(32 percent) (Table 1).

Richness by Geographic Subregions. Fifty-four percent of the plant species of the Wind River Range

(694 taxa) occur in four or less of the ten geographic subregions, with 21 percent (278 taxa) restricted to a

single region (Table 1). By contrast, just 13 percent of the flora is found in nine or more subregions (168 taxa).

Each subregion has at least 12 species that are not found elsewhere in the range. The Limestone Mountain (77

taxa) and Fremont Lake (65 taxa) subregions have the greatest number of such “endemics”. Total species rich-

ness is highest in the Fremont Lake area with 877 taxa, representing 68 percent of the entire flora of the Wind

Rivers (Table 1). This is also the largest subregion and is relatively heterogeneous in elevation and vegetation

(though more uniform in geologic substrates). South Pass, one of the smallest subregions and lacking the eleva-

tional and vegetation diversity of other areas, has the lowest species richness with just 377 taxa, or 29.4 percent

of the total flora.

Based on Jaccard’s Index of Similarity, the average similarity between each pair of subregions is 0.428 (a

score of 1.0 indicates complete similarity, and 0.0 complete dissimilarity). With the exception of the Popo Agie

area, subregions at the far north and west slope of the range have higher average Jaccard’s similarity than re-

gions on the east slope or south end of the range (Table 3). Adjacent subregions tend to be more similar to each

other than to more distant subregions, suggesting that environmental changes are gradual. The Boulder Creek

and Fremont Lake subregions on the west slope are the most similar of any two pairs. Fertig (1992) noted the

primary difference between these regions is in management history, with Fremont Lake being traditionally

grazed by cattle and Boulder Creek by sheep. Massatti (2007) found that differences between the east slope and

west slope subregions were less pronounced when averaged together, with a Jaccard’s similarity of 0.645. The

similarity between east and west slopes may be enhanced by the presence of high elevation calceophilic plants

in the Green River Lakes region, which are otherwise found only on the east side.

The South Pass and Limestone Mountain subregions are the least similar to other areas of the range, with

average Jaccard’s similarity values of 0.318 and 0.375 respectively (Table 3). The distinctiveness of the South

Pass flora can be attributed in part to doristic similarities with the adjacent high desert of the Great Divide Ba-

sin and the absence of large lakes or subalpine and alpine plant communities found in other subregions. The

disparity in total species richness betv

(Fertig 1992). The relatively low elevatu

Wind River Basin and high local endem .

Significance of the Wind River Range. The Wind River Range has the second highest number of plant

taxa of any floristic region in western Wyoming (Table 4), trailing only Yellowstone National Parte This high

n South Pass and other regions can also dampen similarity scon

if the Limestone Mountain subregion, as well as its proximity to th

n are factors contributing to its distinctive flora (Massatti 2007).

Table 3. Jaccard's Index of Similarity for the flora of geographic subregions of the Wind River Range. Values across the top diagonal of the table are numbersofspeds

shared between subregions, while values across the bottom diagonal are the Jaccard's Index of Similarity scores. Jaccard's Index is determined by the formula j =

c/(N, + N r c) where c = number of taxa in common between two sites and N, and N 2 = the number of taxa in sites 1 and 2 respectively.

Popo Agie River drainage (P): 0.471

Table 4. Species richness of selected floras of western Wyoming.

species richness can be attributed to several factors. The Wind Rivers are one of the largest mountainous areas

in the state, and ecologists have long recognized the correlation between species richness and increasing area

(Rosenzweig 1995). The broad elevational range (1725 to 4208 m), diversity of geologic substrates (Precam-

brian granite and gneiss, Mesozoic calcareous formations, and volcanic outcrops), and wide variety of vegeta-

tion types also contribute to landscape heterogeneity and increased species richness. The intensity with which

the Wind River Range has been surveyed for over 180 years (resulting in over 28,600 voucher specimens) is

also a contributing factor to the richness of its flora.

Based on an analysis by Fertig (1992), the Absaroka Range has the most similar flora to that of the Wind

River Range. Both have similar vegetation and elevational range, but differ in geologic substrate (the Absarokas

are primarily volcanic). Geographic proximity may explain the high floristic similarity between adjacent

mountain ranges, but does not account for the disparity in species richness between the Wind River Range and

bordering desert basins. Floristic diversity in the Great Divide, Green River, and Wind River basins is lower

than the Wind River Range in part due to their greater aridity (which restricts the diversity of forest and wet-

917

land habitats) and limited elevational range which precludes alpine tundra and subalpine forest communities.

Likewise, the Wind Rivers have only a subset of the high desert or grassland species prevalent in basin areas

(Fertig 1992).

With nearly one-half of its area protected as Wilderness, the Wind River Range plays a significant role in

the conservation of Wyoming’s native flora. Nearly 76 percent of all plant species known from the Wind Rivers

(968 taxa) are found within the Bridger, Fitzpatrick, or Popo Agie wilderness areas, or other formally protected

lands (Kendall Warm Springs Special Interest Area, BLM Special Status plants ACEC, or The Nature Conser-

vancy’s Red Canyon Ranch Preserve) (Fertig 1995a, 1995b). Of the 265 unprotected native species, all but 20

are protected elsewhere in the state (Fertig 2011). The majority of unprotected species occur at low elevations

and are often uncommon within the Winds (including one-quarter of the 82 species of concern or potential

concern). Most unprotected species occur at the north end, the southeast flank, or the far southern portions of

The checklist is sorted by major phylogenetic groups (fern allies, ferns, gymnosperms, and angiosperms), with

families and species arranged alphabetically. Family taxonomy follows the classification of the Angiosperm

Phylogeny Group (2009). Species nomenclature follows the RM Plant Specimen Database (Hartman et al.

2009), which itself is based on Dorn (2001) and more recent literature. Each species entry includes the current

scientific name and authority, the number of specimens known from the study area (in parentheses), codes for

geographic subregions within the Wind River Range, elevation range in meters, codes for vegetation types, and

selected synonyms (limited to names used in earlier theses or in Dorn 2001 that differ from the currently ac-

cepted name). All vouchers are deposited at RM unless otherwise noted. Hybrid taxa are discussed under one

of the parent species. Individual codes are explained below:

* Species not native to Wyoming

• State of Wyoming noxious weed

♦ Species of conservation concern

# Historical (not relocated since 1970)

sa Sand and gravel bars

sff Spruce-fir forest (includes blue spruce forests)

wm Wet meadows (forb dominated)

wf Whitebark pine forest

wt Willow thickets

Vegetation type:

af Aspen forest

aq Aquatic (submerged or emergent)

at Alpine tundra (includes upper timber-line)

Big sagebrush grassland (includes shrub slopes)

Douglas-fir forest

Dry meadows

Disturbed sites

Granite or gneiss outcrops

Juniper woodland

Lodgepole pine forest

Limber pine woodlands

Marshes and bogs

sagebrush grassland)

Geographic regions within the study area

B Boulder Creek drainage (South Central/West)

D Dinwoody Creek Drainage (North Central/

East)

F Fremont Lake (North Central/West)

G Green River Lakes

L Limestone Mountain (Southeast)

M Moccasin Basin

P Popo Agie River drainage (South Central/East)

S South Pass

T Torrey Lake (Northeast)

W Warm Spring Creek Drainage (North)

Sensitive or Candidate plant species

BLM = Bureau of Land Management

BTNF = Bridger-Teton National Forest

USFS = US Forest Service,

R2 = Region 2 (Rocky Mountain Region)

USFWS: US Fish and Wildlife Service.

, F, G, L, M, P, S, W; 1

t. (94) B, D, F, G, L, M,P,T,W;

Kapoor (28} B, D, R G^P, T, \

, D, F, G, L, M, P, S, T, W;

937

In the summer of 2013, two new species in the Poaceae were documented for the South Pass subregion of the

Wind River Range: Bouteloua gracilis (H.B.K.) Lag. ex Griffiths (Heidel 3874, RM) and Muhlenbergia cuspidata

(Torr. ex Hook.) Rydb. (Heidel 3876, RM).

Funding for much of the fieldwork associated with this project was provided by Bridger-Teton and Shoshone

National Forests and the BLM Wyoming state office through the University of Wyoming and the Wyoming

Natural Diversity Database. Special credit is due to N. Duane Atwood, Kent Houston, Bill Noblitt, and Jeff Car-

roll for funding and logistic support. Joy Handley kindly tracked down some questionable specimens from the

RM for examination. Rob Thurston helped create the map of the study area. Richard Scott, formerly of Central

Wyoming College, shared many records from his nearly 50 years of exploring the Wind Rivers. A thank you is

given to Robert Dorn, Tim Hogan, and Bonnie Heidel for their review of the document and for providing ad-

ditional species records.

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ANNOUNCEMENTS

2013 Delzie Demaree Travel Award Recipients

The 25th Annual Delzie Demaree Travel Award was presented at the 50th Annual Systematics Symposium

(11-12 Oct 2013) at the Missouri Botanical Garden, St. Louis. Six students were presented the Travel Award:

Dakota Ahrendsen, University of Nebraska, Omaha; Shelly Aust, University of Nebraska, Omaha; Benjamin

Gahagen, Ohio University; Alison Scott, University of Wisconsin, Madison; Harlan Svoboda, Ohio University;

Eduardo Tomaz, Ohio University.

The 2013 Travel Awards were underwritten by 1) Contributors to the Delzie Demaree Travel Award En-

dowment, 2) Members of the Delzie Demaree Travel Award Committee, 3) Mary Isabelle Eggers, Williams-

burg, Virginia, and 4) John Clayton Chapter of the Virginia Native Plant Society.

Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the

travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research

Institute of Texas, to Barney Lipscomb, 1700 University Drive, Fort Worth, TX 76107-3400, U.S.A. 1-817-332-

7432; Email: barney@brit.org. Thank you.

The 2014 Applications for the Delzie Demaree Travel Award

Applications for the 2014 Delzie Demaree Travel Award should include a letter from the applicant telling how

symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major

professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology De-

partment, The College of William and Mary, Williamsburg, Virginia 23185-8795, U.S.A. 1-757-221-2799;

Email: ddmware@wm.edu. Applications may be sent to: Barney Lipscomb, 1700 University Drive, Fort Worth,

Texas 76107-3400, U.S.A. 1-817-332-7432; Email: bamey@brit.org. The period for receiving applications will

end three weeks prior to the date of the symposium if a sufficient number of applications are in hand at that

time. Anyone wishing to apply after that date should inquire whether applications are still being accepted be-

fore applying. The Systematics Symposium dates for 2014 are 10-11 October 2014 (dates tentative and subject

to change).

The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35

out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discov-

erer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He also

taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas A&M,

and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of the things he

enjoyed most as a botanist was assisting students with their field botany research.

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grass-like species, the book also ventures into territory often neglected by popu-

lar botanical texts. In short, the reader will become well-informed and enjoy the

I process! ” — Johnny Townsend, Botanist, Virginia Natural Heritage Program,

Department of Conservation and Recreation, Richmond, Virginia.

LTON AND Gustavus Hall. 2013. WiWJIoH*rs & Grasses of Virginia’s Coastal Plain. (ISBN: 978-1-

6, flexbound). Sida, Botanical Miscellany 40. Botanical Research Institute of Texas Press, 1700

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