Journal of the Botanical Research Institute of Texas

J. Bot. Res. Inst. Texas ISSN 1934-5259

Fort Worth, Texas 76107-3400, USA

817-332-7432; 817-332-4112 fax

Electronic mail: barney@brit.org; jbrit@brit.org

History and Dedication

1962— Lloyd H.Shinners

(left), a member of the

Southern Methodist University

(SMU) faculty and a prolific

researcher and writer, published the first issues of S/do,

Contributions to Botany (now J. Bot. Res. Inst. Texas)

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

botany at SMU and director emeritus of BRIT,

inherited editorship and copyright.

1993— BRIT becomes publisher/copyright holder.

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

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of botanical research and scientific discoveries

for the twenty-first century.

The BRIT Press— bringing out the best in botanical

science for plant conservation and education.

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The BRIT Press considers original research papers

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botany, sensu lato, for publication in J. Bot Res. Inst. Texas.

Guidelines for submissions are available

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Bibliographical

Citation abbreviation for the

Journal of the Botanical Research Institute of Texas is

J. Bot. Res. Inst. Texas following the principles

of B.P.H. (informally JBRIT).

International Standard Serial No. (ISSN 1934-5259)

Frequency of Publication

J. Bot. Res. Inst. Texas is published semiannually

(summer/fall) as one volume

by the Botanical Research Institute of Texas.

Felix Llamas (Universidad of Le6r

Editorial Board

Harold Keller (BRIT)

Robert J.O'Kennon (BRIT)

Richard Rabeler (MICH)

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Composition

rhorngraphics, Plano, Texas; rlrcf

Cover Illustration

Electronically tinted botanical ill

of Liatris aestivalis originally user

BRITs anniversary poster 2001.

Summer gayfeather flowers mic

Jul-Aug(-Sep) and is endemic t

Oklahoma and Texas.

Sida 19:768. 2001.

Press

Table of Contents

SYSTEMATICS

A new Sisyrinchium (Iridaceae) from cedar glades in northern Alabama

Bruce A. Sorrie, Wesley M. Knapp, L. Dwayne Estes, and Daniel D. Spaulding

Polemonium elusum (Polemoniaceae), a new species from east central Ida

Joshua J. Irwin, Rebecca Stubbs, and Ronald L. Hartman M ' S

ind James L. Reveal

Systematics of Lindleya (Rosaceae: Maloideae)

James Henrickson

Illustrations and studies in neotropical Orchidaceae. 5. The I

(Pleurothallidinae) with three new species from Costa Rica

Diego BogarIn, Christina M. Smith, and Daniel JimEnez

SSOURI BOTANICAL mi

DFK 0 4 7ni2 “9

GARDEN LIBRARY 341

Helen Kennedy

of the Colombian A

a new species of Gesneriaceae from the Cordillera Central

[ohn L. Clark and Laura Clavijo

\ new variety of Dedieuxia cacui

Joseph H. Kirkbride, Jr. and Piero G. Delprete

A previously unrecognized species of Senegalia (Fabaceae) from northeastern Brazil

David S. Seigler, John E. Ebinger, and Petala Gomes Ribeiro

391

397

403

Cytotypic variation in Phlox pilosa ssp. pilosa (Polemoniaceae) at the western edge

of its range in the central United States

Lindsey Worcester, Mark H. Mayfield, and Carolyn J. Ferguson

A new combination and a new species in Combretum (Combretaceae) from India

K.A. Sujana, M.K. Ratheesh Narayanan, and N. Anil Kumar

i, Mexico. I. Convolvulus, Cressa

443

453

Richard S. Felger, Daniel F. Austin, Thomas R. Van Devender, J. JesOs Sanchez-Escalante,

and Mihai Costea

id J. Jesus SAnchez-Escalante

~. Austin, Thomas R. Van Devender,

PALEOBOTANY

459

529

551

557

A new fossil species of Colpothrinax (Arecaceae) from Mid-Tertiary Mexican amber

Kenton L. Chambers, George 0. Poinar, Jr., and Alex E. Brown

Additional fossils in Dominican amber give evidence of anther abortion in Mid-Tertiary

Trichilia (Meliaceae)

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

DEVELOPMENT AND STRUCTURE

Limitations to natural production of Lophophora williamsii (Cactaceae) 11.

Effects of repeated harvesting at two-year intervals in a South Texas population

Martin Terry, Keeper Trout, Bennie Williams, Teodoso Herrera, and Norma Fowler

HERBARIUM METHODS AND TECHNIQUES

A cost-effective method for constructing magnetic fumicels for herbarium cabinets

FLORISTICS, ECOLOGY, AND CONSERVATION

Nuevos registros de Poaceas para el norte de Mexico

Y. Herrera Arrieta, C.A. Silva Salas, L. Ruacho Gonzalez y O. Rosales Carrillo

Distribution actual y potencial de Taxus globosa (Taxaceae) en Mexico

Mario A. GarcIa-Aranda, CEsar CantO-Ayala, Eduardo Estrada-CastillOn,

Marisela Pando-Moreno y Antonio Moreno-Talamantes

and A. nigrum (Viscaceae) in Mexico

Robert L. Mathiasen, Shawn C. Kenaley, and Brian P. Reif

Taxonomic history, rediscovery, and assessment of threat status of Streblus ilicifolius

(Moraceae) from India

Bikarma Singh, Arun Chettri, Dibyendu Adhikari, and Saroj K. Barik

Three noteworthy additions to the Alabama flora

Alvin R. Diamond and Brian R. Keener

A first spontaneous record of Actinidia chinensis var. deliciosa (Actinidiaceae)

in the United States flora

Brett E. Serves, David H. Mason, and Troy L. Bray

Pallenis maritima (Asteraceae)

of salt-tolerant ornamental plants

Richard E. Riefner, Jr. and Werner Greuter

Vascular flora and plant communities of Dead Horse Knob (Rucker’s Knob),

Madison County, Kentucky

Ralph L. Thompson, Derick B. Poindexter, and J. Richard Abbott

Vascular plants of the Ya 2

in Grenada County, Mississippi

Michael Wayne Morre and John R. MacDonald

ity, Pennsylvania

New Pennsylvania county occurrences for Beilis perennis. Hibiscus moscl

Lamiutn maculatum, and Robinia hispida

Jerky G. Chmielewski and David Krayesky

561

567

579

583

599

615

617

621

653

681

707

A NEW SISYRINCHIUM (IRIDACEAE) FROM CEDAR GLADES

IN NORTHERN ALABAMA

Bruce A. Sorrie

NC Natural Heritage Program and

University of North Carolina Herbarium

North Carolina Botanical Garden

Chapel Hill, North Carolina 27599-3280, U.S. A.

nruce.sorrie@ncdenr.gov

L. Dwayne Estes

Department of Biology &

Center for Field Biology

Austin Peay State University

Clarksville, Tennessee 37044, USA.

Wesley M. Knapp

Maryland Department of Natural Resources

Wildlife and Heritage Service

P.O. Box 68, Wye Mills, Maryland 21679, U.S.A

wknapp@dnr.state.md.us

Daniel D. Spaulding

Anniston Museum of Natural History

P.O. Box 1587

Anniston, Alabama 36202, U.S.A.

RESUMEN

Sisyrinchium L. is a taxonomically difficult genus with 37 species in North America north of Mexico. Of these,

only Sisyrinchium albidum Raf. and S. capillare Bicknell have paired inflorescences (Cholewa & Henderson

2002). Each of the inflorescences is subtended by two bracts or spathes, which terminate the unbranched

stems. Although this combination of characters is rare in North American Sisyrinchium, a third entity shares

these character states and is the subject of this paper. We propose Sisyrinchium calciphilum Sorrie for this spe-

cies, which is endemic to limestone glades in northwestern Alabama.

Plants: perennial, cespitose, 20-42 cm tall. Stems simple, clearly but narrowly winged, 0.7-1.3(-1.5) mm wide,

each wing wider than stem core, glabrous, margins entire, stem base purple to pinkish brown. Leaves: gla-

brous, bases not persistent as fibrous tufts. Inflorescences: paired one sided cymes, i.e., rhiphidia, each sub-

tended by a pair of spathes, these subtended by a bractlike leaf 41-82 mm long that often obscures inner inflo-

rescence; bractlike leaf and spathes purple tinged, spiculate, spathe keels denticulate, outer spathe of outer pair

12-18 mm, spathes of outer inflorescences averaging 4.4 mm longer than spathes of inner inflorescences,

spathe margins hyaline, translucent to purple. Flowers: tepals blue with yellow bases, 7-9 mm long, apex

weakly emarginate, aristate; filaments connate most of length, egandular; ovary green and glandular-hairy.

Capsules: fully mature capsules not seen; immature capsules more-or-less globose, 4-5 mm long and wide,

pale green, pedicels spreading to ascending, glabrous or with very sparse glandular hairs. Seeds: mature seeds

324

Journal of the Botanical Research Institute of Texas 6(2)

Sorrie et al., A new Sisyrinchium from Alabama

325

27 Apr 1993, R. Whetstone, D. Spaulding, J. Ballard, & T. Dobson 16436 (JSU); N of Courtland, just S of Wheeler Station Sporting Clays, recent-

' 4 May 1996, R. Whetstone &S.Hruska 17457 (JSU); Prairie Grove Glades Preserve, abundant on thin clay

e, 19 Apr 2003, R. Krai 93898 (AMAL, APSC); 2 mi SE of Speake on Alabama 157, clearing in limestone woods,

1968, R. Krai 30489 (GA); 1.5 mi SW of Flat Rock, limestone cedar glade, tepals dark purple, 17 Apr 2006, B. Keener & D. Spaulding

2817 (AMAL, UNA); Bankhead National Forest, off FSR-264, limestone woods, 17 Apr 2005, D. Spaulding & B. Keener 12252 (AUA.UNA).

CR-55 near Emmanuel Road, disturbed limestone glade, 13 Apr 2003, D. Spaulding 11723

«d E of Massey, 16 Apr 2011, B. Sorrie, D. Estes, & W. Knapp 12746 (DUKE, GA, NCSC,

n slope in dryish woods, 16 Apr 2011, B. Sorrie, D. Estes, & W. Knapp 12747 (NCU, NY).

Sisyrinchium albidum was the first of the blue-eyed-grasses with paired inflorescences to be named (Rafinesque

1832). Much later, Bicknell split out S. capillare with its wiry, barely winged stems (Bicknell 1899). Since there

appeared to be overlapping characters between these two species, some authors declined to recognize S. capil-

lare (e.g., Radford et al. 1968) or confounded their habitats and ranges (Cholewa & Henderson 2002). Recent

herbarium and field work by the authors show that S. capillare occurs strictly in the Atlantic Coastal Plain

Physiographic Province whereas S. albidum occurs primarily in the Piedmont, Interior Low Plateau, and Inte-

rior Highlands Physiographic Provinces, northward to the Great Lakes states; and with populations southward

to the Gulf Coastal Plain (Fig. 6). In addition, there are strong morphological differences, notably the slender

and nearly wingless stem of S. capillare (vs. clearly winged in S. albidum), stem with dense fibrous remains of

leaves (vs glabrous to only moderately dense), nearly equal spathes (vs. very unequal), and short outermost

spathe (vs. long) (Table 1). Note that Table 1 divides S. albidum into two groups, east and west of the Appala-

chian Mountains. These groups appear to differ from each other, notably in stem width and the difference be-

tween inner and outer spathe length, but there is much overlap. At this time we are reluctant to recognize any

infraspecific taxa and treat S. albidum as a widespread, variable species.

Specimens of S. calciphilum are quite different from S. capillare, where they had been placed due to their

slender stems. Stems of S. calciphilum are definitely (albeit narrowly) winged, the fibrous remains of leaf bases

are absent, the two outer spathes exceed the two inner by a mean of 4.4 mm, spathes and leaflike bracts are

strongly purple tinged and spiculate. Corolla color of S. calciphilum is medium blue, unlike the pale blue to

whitish color of S. capillare (Figs. 2 & 3). Moreover, S. calciphilum is restricted to high pH limestone glades,

whereas S. capillare to acidic, fire-maintained longleaf pine savannas (Table 1).

calciphilum is less strikingly distinct. While a single morphological charac-

em populations of S. albidum, the suite of characters distinguishes 5. calci-

philum (Table 1). Critical differences are: stem width of S. calciphilum is similar to that of many plants from east

of the Appalachians, but only half as wide as sympatric plants from west of the Appalachians. Stem margins of

S. calciphilum are smooth, but denticulate (often strongly so) in S. albidum. The difference between the two in-

um blue, unlike the pale blue to whitish color of S. albidum (Figs. 2 & 4). Coupled with the restricted range and

habitat preference, these morphological differences are significant at the species level.

DISTRIBUTION AND HABITAT

: counties in northwestern Alabama (Fig. 5), a region underlair

.. Heller) Bameby, Forestiera ligustri

Journal of the Botanical Research Institute of Texas 6(2)

j Sisyrinchium from Alabama

327

Journal of the Botanical Research Institute of Texas 6(2)

Table 1. Co

U 9 2-)1.5-2.5(-2.8)

330 Journal of the Botanical Research Institute of Texas 6 ( 2 )

BOOK REVIEW

Noel H. Holmgren, Patricia K. Holmgren, James L. Reveal, and Collaborators. 2012. Intermountain Flora,

Vascular Plants of the Intermountain West, U.S.A. Volume Two, Part A: Subclasses Magnoliidae-

CaryophylUdae. (ISBN: 978-0-89327-520-4, hbk.). New York Botanical Garden Press, 2900 Southern

Blvd., Bronx, New York 10458-5126, U.S.A. (Orders: http://www.nybgpress.org, 718-817-8721, fax 718-

817-8842). $150.00, 731 pp, color frontispiece, grayscale epilogue, line drawings throughout, Th n x

Volume 2A, the Intermountain Flora Team brings to a successful close a decades-long, j

multigenerational effort to document the plant life of the colder drylands occupying the large region in the

western United States more or less bounded by the Sierra Nevada to the west, the Rocky Mountains to the east, j

and stretching from central Idaho south to northwestern Arizona. Originally conceived by Bassett Maguire j

around 1940, the ensuing research became a driving force in the careers of Arthur Holmgren and Arthur Cron- |

quist, as well as the botanists who authored this final volume. Publication of the six volumes in eight big books

(most floristicians have difficulty estimating space requirements for their works) took forty years, but time

alone is a poor estimate of the scope of the exploration and research that went into the project. The full series j

treats 3,847 species in 898 genera. For those who do not own these wonderful books, the publisher is cur- ;

rently offering the whole set for $520 (a $640 value). There also are rumors of a planned ninth volume with a

Volume 2A of the work, which includes the beginning of the dicots in the Cronquistian classification -j

system, treats 147 genera, 611 species, and 301 additional infrataxa in 31 families, including such regionally j

diverse important groups as the Papaveraceae, Ranunculaceae, Polygonaceae, Caryophyllaceae, Chenopodia-

ceae, Amaranthaceae, Nyctaginaceae, Montiaceae, and Cactaceae. The presentation is identical to that in ear-

lier volumes, with indented keys and detailed descriptions followed by range/ecology statements and very j

useful critical notes on taxonomic and other issues (along with copious literature citations). The treatments j

also include extensive synonyms, with complete citations of types. The numerous plates of line drawings (by ;

several artists), which cover varying numbers of taxa per genus, are uniformly excellent in composition and ;

detail. The drawings of Cactaceae are particularly beautiful. A useful addendum summaries the 6 new combi- J

nations, 34 new typifications, and 1 new cytological report included in the volume. Throughout, the contents

are encyclopedic and the treatment of any family or genus provides a marvelous introduction to the taxonomy,

nomenclature, ecology, and uses for that group.

There is little to criticize in this enormous work. Perhaps some of the more recent volumes are a bit pricey,

but sadly that has become the nature of the printed page. The lack of a comprehensive key to dicot families

(such keys exist for the remaining major plant groups in other volumes) might be addressed in the proposed

follow-up volume, as might a more detailed taxonomic summary of the plants in the series or a discussion of

plants of conservation concern in the region (a 70-year study surely can offer a unique perspective on regional

patterns of plant endangerment). However, such potential additions in no way diminish the present utility of

the volumes. Between the burnt-orange covers of these volumes lies a treasure-trove of information and in-

sights on the vascular plants unequaled for most other regions of the country. The authors, past and present,

are to be congratulated on their persistence in seeing this project through to successful completion . — George

Yatskievych, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, USA.

POLEMONIUM ELUSUM (POLEMONIACEAE),

A NEW SPECIES FROM EAST CENTRAL IDAHO, U.S.A.

Ronald L. Hartman

mm

Perennial herbs, caespitose, glabrous to pubescent with trichomes sparse to moderately dense, minute, flaccid

to erect, often glandular trichomes, not odiferous; branches subterranean, 3-15. Stems 20-50 cm tall. Leaves

mostly basal and the bases sheathing to scattered along stem, 5-20, 10-30 cm long proximally to 2 cm distally,

1.5-3(-4) wide; petioles 2— 7(— 11) cm long; pinnae 12-24, mostly 2-parted, alternate or paired and thus appear-

ing whorled, narrowly to broadly elliptic or spatulate, occasionally obovate, 4-28 mm long, 1-3.5 mm wide.

Inflorescences a thyrse, often diffuse; branches 2-5, 5-11 cm long, strict; flowers 15-60, congested distally;

pedicels 5-6 mm long. Flowers 5-merous; perianth regular; calyx fused at base, campanulate; lobes lanceo-

late, green to purplish in part, 4.4-8 mm long, diverging at a 35-40 degree angle; connective membrane in bud,

flower, and fruits translucent, whitish, oblong to spatulate, 0.4-0.7 mm wide; sinuses rounded; corolla Ught

blue to white, campanulate, 10-13 mm long, tube 4-5.2 mm long; lobes 5-7.8 mm long, spatulate to obovate;

apex rounded, minutely apiculate; stamens usually exserted; filaments attached 2-3 mm above base, at which

point minutely and densely tomentose for 0.8-1 mm; anthers basifixed, narrowly to broadly sagittate, broadly

elliptic to orbicular following dehiscence, 1.4-1.8 mm long; ovary greenish, broadly ovoid, 2 mm long, with 5

raised nerves at least in part; placentation axile, chambers 3; styles 6.5-8 mm long; branches flattened, 2.5-3.2

mm long; stigma glandular over adaxial surface. Fruit globose, tricarpellate, 4.5-5 mm long, split for 60-70

percent of length. Seeds brown, trigonous, the abaxial surface rounded, the edges acute, 24—2.8 mm long;

Etymology.— The epithet was chosen as this species has eluded discovery for well over a century, this despite

sporadic botanical forays to the region.

Distribution and ecology. — Polemonium elusum is known from the mountains and foothills surrounding

the town of Ellis, in east central Idaho (Fig. 2). Populations occur from 1440 to 2560 m in elevation, where

vegetation transitions from sagebrush and mountain mahogany to Douglas-fir woodland, depending on slope

and aspect. Within this range of vegetation, it was found in a variety of microhabitats, thus the species appears

to be an ecological generalist. Microhabitats in which plants are found include the margins of talus fields, dry

Douglas-fir woodland, outer riparian margins, and shaded rock outcrops. Plants occur in stable but loose,

coarse textured colluvial soils. All populations were found on various geologic units of the Challis volcanics

group (Zientek et al. 2005) and were most abundant from lower Cow Creek southward across the adjacent un-

The initial discovery was made by

potential habitat for Cryptantha salmon

(Rydb.) Blank, var. salmonensis P £

found growing under more mesic conditi

1-June 2010. This locality was surveyed as

Payson, Astragalus amblytropis Barneby, and Oxytropis besseyi

demies was encountered, but Polemonium elusum was

Id be expected for the aforementioned taxa. Further

ivn that populations of P. pulcherrimum var. pulcherrimum surround the distribution of P.

cies have several similarities, including overlapping elevation ranges, dry habitats, and a

e of the corolla and the inflorescence. Additionally, P viscosum has been observed as low as

p canyons above the Salmon River to the north and south of P. elusum. However, these con-

Journal of the Botanical Research Institute of Texas 6(2)

Phenology . — Flowering begins in late-May at more xeric, low elevation sites, continuing through mid-July

at higher, cooler sites; fruiting from mid-June through August.

Conservation .— Perhaps the greatest threat to the existence of Polemonium elusion is the small total popu-

>tal about 1,300 plants, occupying eight km .

Other probable threats include herbivory by ungulates and habitat invasion by exotic weeds. Interestingly, the

population at Gerry Gulch occurs at a location where the vegetation is significantly impacted by cattle. Her-

bivory by white-tailed deer has been shown to be a significant factor for population survival in Polemonium

vanbruntiae Britton (Bermingham 2010). If herbivory by cattle and wildlife affects the survival of P. elusum,

then the maintenance of ungulate populations below certain levels may be required.

Other aspects of the biology of this taxon are unknown. However, Polemonium vanbruntiae is self compat-

i species of Polemo

ible (Hill et al. 2008). Because reproductive and life history traits influence the survivorship of plant popula-

tions, further research on P. elusum is necessary in order to develop a more effective conservation strategy.

Molecular analysis. — Sequence data from 28 taxa were analyzed. The 46 vouchers used in this phyloge-

netic study are listed in Appendix 1. Twenty-six of these were accessions from GenBank. Leptosiphon, Linanthus,

and Phlox were used as outgroups because they have been proposed as genera closely related to Polemonium

(Porter 1997; Johnson et al. 2008; Hankamp 2011).

Field collections of fresh plant material were preserved in silica gel and voucher specimens are deposited

in the herbarium at San Francisco State University (Table 1). DNA extracted from herbarium sheets follow the

same process. Extraction protocols are described in DNeasy (2004). The 1TS1, 5.8S, and ITS2 regions were

amplified using primers 1TS4 and 1TSLEU (White et al. 1990). Both regions were duplicated using standard

Polymerase Chain Reaction (PCR) in 25 pi volume reactions. Excess nucleotides and primers remaining in the

samples from the PCR were removed using ExoSAP-it kit.

An Applied Biosystems 3100 Genetic Analyzer was used for the capillary electrophoresis of all samples.

Fragments were sequenced using BigDye following the manufacturer’s protocols. Precipitation was with

EDTA/Ethanol/Sodium acetate, and the remaining cycle sequence products were resuspended in Hi-Di before

being denatured. Fragments were visualized using an Applied Biosystems PRISM 3100 Genetic Analyzer. Run

modules were conducted using liquid polymer POP-6 or POP-7. Sequence files were base-called using Sequenc-

ing Analysis 5.1 (Applied Biosystems 2003), and the forward and reverse reads were formed into a consensus

sequence using Sequencher 4.8 (Gene Codes Corporation 2007). The consensus sequence contig was loaded

into MacClade 4.08X (Maddison & Maddison 2005) to visually confirm the coherence of the bases, and into

ClustalX Version 2 (Larkin etal. 2007) for a complete alignment. The nexus file was analyzed in PAUP4.0al 12

(Swofford 2002), Mr. Bayes 3.1.2 (Ronquist & Huelsenbeck 2003), and GARLI 0.951-GUI (Zwickl 2006).

This ITS data set included 46 samples with a total of 706 characters, 163 of these were variable. Of the variable

characters, 61 were parsimony informative. Bootstrap analyses were performed in PAUP 4.0all2 (Swofford

2002) using a heurist search with 100 repetitions and number of trees increasing by 100. Trees were also ana-

lyzed using maximum likelihood and Bayesian analysis.

For statistical selection of the best fit model, jModelTest (Posada 2008) was used. The -InL using jModelT-

est was 2412.83. The results indicated GTR+I+G, and these parameters were employed. Bootstrapping was

tested with GARLI and Bayesian analysis provided numbers for branch support.

DISCUSSION

Molecular analysis of the ITS region of Polemonium weakly suggests P. elusum is sister to R mexicanum, P. pauci-

florum, P. grandiflorum, and P cameum (Fig. 3). This is the best inference from the maximum likelihood tree

but it is not statistically supported by either bootstrap (56%) or Bayesian (0.84) values. A few conclusions can

be drawn from these results. First, the four species that are most closely related to P. elusum are all included in

Grant’s (1959) sect. Polemonium and P. elusum also fits into this section. Worley et al. (2009) further divided

Grant’s sect. Polemonium into three species complexes, and except for P. mexicanum, the other three species are

all included in the P. pauciflorum complex. This species complex includes plants that are erect to decumbent.

The inflorescences are panicle-like cymes or the flowers are arranged in groups of one to three. Polemonium

elusum meets these criteria.

Second, under Wherry’s (1942) classification, P. pauciflorum , P. grandiflorum, and P cameum all fall under

the large flowered section Eupolemonium. Once more, P. elusui

These species all have overlapping elevation ranges and low ir

North America. Although there is not much statistical support for this grouping, morphometric features help

validate the placement of Polemonium elusum with this group.

Polemonium elusum is congruent with the rest of the genus in its campanulate corolla, habit, compound

leaves, stamen attachment, pubescent filaments, and brown seeds. This species notably differs in the calyx

The leaf-

Journal of the Botanical Research Institute of Te

ACKNOWLEDGMENTS

We would like to thank Teresa Prendusi and Bruce Smith for their support of the floristic inventory, thus mak-

ing this discovery possible. Lucinda Haggas coordinated our logistical needs during summer field work. Bob

Patterson and Dieter Wilken provided helpful suggestions regarding the manuscript. Wendy Irwin graciously

provided Figure 1.

Applied Biosystems. 2003. Sequencing Analysis* version 5.1 . httpy/www.appliedbiosystems.com.

Bermingham, L.H. 2010. Deer herbivory and habitat type influence long-term population dynamics of a rare wetland

plant. PI. Ecol. 210:359-378.

DNeasy, Q. 2004. Plant DNA extraction handbook, http://www.qiagen.com.

Gene Codes Corporation. 2007. Sequencher* version 4.8 sequence analysis software. Ann Arbor, Ml. http://www.gene-

codes.com.

Grant, V. 1959. Natural history of the Phlox family, vol. 1, systematic botany. The Hague: Martinus Nijhoff.

Hankamp, P.Z. 201 1 . Molecular systematics of Leptosiphon (Polemoniaceae). M.S. thesis. San Francisco State University, CA.

Hartman, R.L. 1992. The Rocky Mountain Herbarium, associated floristics inventory, and the Flora of the Rocky Moun-

tains project. J. Idaho Acad. Sci. 28:22-43.

Hartman, R.L. and B.E. Nelson. 201 1 . General information for floristics proposals. [The Boiler Plate]. http:/www.rmh.uwyo.

edu.

Hartman, R.H., B.E. Nelson, and B. Legler. 2009. Rocky Mountain H<

Hill, LM., A.K. Brody, and C.L Tedesco. 2008. Mating strategies <

Polemonium vanbruntiae. Acta Oecol. 33:314-323

Johnson, LA., L.M. Chan, T.L. Weese, L.D. Busby, and S. McMurry. 2

strong inference of higher phylogenetic relationships in th

Kesonie, D. (Scott) and R.L. Hartman. 201 1. A floristic inventory of Grand Teton National Park, Pinyon Peak Highlands, and

vicinity, Wyoming, U.S.A. J. Bot. Res. Inst. Texas 5:357-388.

Kuhn, B, B.E. Nelson, and R.L. Hartman. 2011. A floristic inventory of the Cimarron National Grassland (Kansas) and the

Comanche National Grassland (Colorado). J. Bot. Res. Inst. Texas 5:753-772.

Larkin, M A, G. Blackshields, N.P Brown, R. Chenna, P.A. McGettigan, H. R

Thompson, TJ. Gibson, and D.G. Higgins. 2007. Clustal W and Clustal X version 2.0. B

Maddison, D.R. and W.P. Maddison. 2005. MacClade 4: Analysis of phylogeny and char

Sinauer Associates, Sunderland, MA.

:r DNA sequences. Aliso

Posada, D. 2008. jModelTest: phylogenetic model averaging. Molec. Biol. Evol. 25:1253-1256.

Reie, B, J. Larson, B. Jacobs, B.E. Nelson, and R.L. Hartman. 2009. Floristic studies in north central New Mexico, U.SATheTusas

Mountains and the Jemez Mountains. J. Bot Res. Inst. Texas 3:921-961 .

RONQU.ST, F. AND J.P. Huelsenbeck. 2003. MRBAYES 3: Bayesian phylogenetic inference under mixed models

19:1572-1574.

Swofpord, D.L. 2002. PAUP*. Phylogenetic analysis using parsimony (*and Other Methods). Version 4. Sin;

d direct sequencing of fungal ribosomal RNA genes

White, TJ.,T.D. Bruns, S.B. L

for phylogenetics. In: Innis, M A, D.H. Gelfand, JJ. Sininsky, and TJ.White.'eds. PCR ^oio^ols^’guide tom* hods'and

applications. Academic Press Inc., San Diego, CA. Pp. 31 5-322.

Wherry, E.T. 1942. The genus Polerru

Worley, A.C., H. Ghazvini, and D.W. Schemske. 2009. /

length polymorphism (AFLP) markers. Syst. Bot. 34:149-161.

Zientek, M.L, P.D. Derkey, a

i datasets

NEOTYPIFICATION OF AMORPHA ROEMERIANA (FABACEAE: AMORPHEAE)

Shannon C.K. Straub 1 and James L. Reveal

L.H. Bailey Hortorium, Department of Plant Biology, 4 12 Mann Library Building

Cornell University, Ithaca, New York 14853, USA.

ss463@cornell.edu; jlr326@cornell.edu

Se designa un neotipo (WR Carr et cd 27f

RESUMEN

In the process of preparing a treatment of Amorpha L. for a forthcoming volume of Flora of North America, we

attempted to locate original material of Amorpha roemeriana Scheele, a relatively rare species found primarily

on the Edwards Plateau of Bandera, Bexar, Blanco, Comal, Gillespie, Hays, Kendall, Kerr, Kinney, Medina,

Travis, and Uvalde counties of central Texas (Turner et al. 2003: 306), with disjunct populations in Coahuila,

Mexico, that scarcely differ from the Texas populations. The type was collected by Carl Ferdinand von Roemer

(1818-1891) “In margine rivulorum prope, Austin” during his 1845-1846 visit to Texas where he concentrated

on geology (Simonds 1902). Upon his return to Europe, Roemer wrote several books and articles on his visit

parleying these activities eventually into a professorship in geology at the University of Breslau. Roemer’s plant

specimens were given to George Heinrich Adolf Scheele (1808-1864) who accounted for several new Texas

species (Scheele 1848) gathered by Roemer and, independently, by Ferdinand Jacob Lindheimer (1801-1879),

including A. roemeriana which was reported to have been collected, in flower, in April of 1846. Roemer (1849:

429) himself stated that the plant was found “Bei Austin am Bachrande,” and the “Bluthen sehr wohlriechend,

violet.”

Scheele’s types were housed at Berlin, but as noted by Wilbur (1975), no one has reported, in the literature

at least, the existence of any original material of Amorpha roemeriana. We attempted to locate a specimen at the

Roemer-und Pelizaeus-Museum in Hildesheim, Germany, which at one time contained specimens gathered by

Ferdinand Roemer (the Museum is named for his brother, Hermann Roemer [1816-1894]), only to be in-

formed that the plant collection was sent to Berlin in the early 1900s. Nonetheless, there is no record at the

Museum that there was a specimen of Amorpha from Austin, Texas, gathered in 1846.

The detailed description written by Scheele provides the characters necessary to distinguish Amorpha

roemeriana from A.fruticosa, the most widespread species of the genus and the only one co-occurring with A.

roemeriana. The inclusion of the swollen mucro, glandular punctuate leaflets, petiolules bearing purplish

glands, and glandular vexillum in the description clearly point to A. roemeriana as that name is applied cur-

rently (Schneider 1907; Rydberg 1919; Palmer 1931; Wilbur 1975; Isley 1998), and contrast with the long, slen-

der mucro, commonly eglandular leaflets, eglandular or inconspicuously glandular petiolules, and generally

eglandular vexillum of A. fruticosa. Confirming the assumption that the name as currently applied is correct

could be only accomplished by examining type specimen material. Being unable to locate any original mate-

rial, we are here designating a neotype to maintain the current concept of A. roemeriana, which we believe to

be consistent with the original description. With the able assistance of William R. Carr of the Nature Conser-

vancy of Texas and Dr. Thomas Wendt of the University of Texas, to whom we are grateful, a suitable collection

was obtained.

340

irpha roemeriana Scheele, Linnaea 21:461. 1848. Type UNITED STATES. Texas. Hays Co.: Ashe juniper woodland on

extremely shallow stony clay loam on top of N- to WY-facing bluff of Fredericksburg Limestone ca. 40-50 ft above the S bank of

ir mi W of the junction of State Route 150 and Ranch Road 2770 near M

e 150 and Ranch Road 3237 at Hays City, at N30°00'37.5", W097°58'01.r

Kf.WR. Carr, B.Johnso

ACKNOWLEDGMENTS

We wish to thank the curator of TEX for loan of material that was essential to completing this work and our

reviewers, Billie L. Turner and Stanley L. Welsh.

REFERENCES

Isley, D. 1 998. Native and naturalized Leguminosae (Fabaceae) of the United States. Monte L. Bean Life Science Museum,

Brigham Young University, Provo, Utah.

Palmer, E J. 1 931 . Conspectus of the genus Amorpha. J. Arnold Arbor. 1 2:1 57-1 97.

Roemer, C.F. von. 1 849. Texas. Mit besonderer rucksicht auf deutsche auswanderung und die physischen verha'ltnisse des

landes nach eigener beobachtung geschildert von Ferdinand Roemer. Mit einem naturwissenschaftlichen anhange

und einertopographisch-geognostischen karte von Texas. Adolph Marcus, Bonn.

Rydberg, P.A. 1919. Amorpha. N. Amer. FI. 24(1):26-34.

Scheele, A. 1 848. Beitrage zur Flor von Texas. Linnaea 21 :453-472, 576-602, 747-768.

Schneider, C.K. 1 907. Conspectus generis Amorphae. Bot. Gaz. 43:297-307.

Simonds, F.W. 1902. Dr. Ferdinand von Roemer, the father of the geology of Texas; his life and work. Amer. Geol. 29:

131-140.

Turner, B.L., H. Nichols, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, vol. 1 : Dicots, vol. 2: Ferns, gym-

nosperms, monocots. Sida, Bot. Misc. 24.

Wilbur, R.L. 1 975. A revision of the North American

i genus Amorpha (Leguminosae-Psoraleae). Rhodora 77:337-409.

SYSTEMATICS OF LINDLEYA (ROSACEAE: MALOIDEAE)

James Henrickson

Plant Resources Center

University of Texas, Austin, Texas 78712, U.SA.

ABSTRACT

Chihuahua, south to central San Luis Potosi and southwestern Tamaulipas; in Hidalgo-Queritaro; and Puebla-Oaxaca with outlying collec-

is discussed along with a call for further modifications of the classification of the Rosaceae.

RESUMEN

INTRODUCTION

The monotypic genus Lindleya (Rosaceae) consists of sclerophyllous, evergreen shrubs with conspicuous

white flowers and distinctive, woody, 5-carpelled capsular fruit. It is native to the arid and semi-arid chaparral

or matorral of Mexico, with populations known from Puebla-Oaxaca, Hidalgo-Queretaro and the mountains

of the Sierra Madre Oriental and the Chihuahuan Desert from San Luis Potosf, Tamaulipas, Nuevo Le6n, Coa-

huila and eastern parts of Zacatecas, Durango and Chihuahua.

The genus has a complex but interesting taxonomic history that is reviewed below. Data on vegetative and

reproductive features are presented and illustrated along with a formal taxonomic treatment and distribution

maps. The most interesting aspect of Lindleya is its place in the phylogeny of the Rosaceae. With its five-car-

peled capsular fruit, its placement within the family has been problematic.

MATERIALS AND METHODS

The study is based on empirical observations from field-collected and herbarium material as well as extensive

field studies. Herbarium specimens were borrowed from A, GH, MO, MICH, NY, TEX-LL and US. Additional

data was obtained from collections from ARIZ, ASU, CAS, DS, ENCB, MEXU, RSA, POM during visits to those

herbaria. Anatomical studies incorporated standard paraffin techniques (Johansen 1940) for production of

serial sections of leaves and flowers. All plant measurements were made from dried material unless otherwise

342

TAXONOMIC HISTORY

Taxonomic History.— The genus Lindleya was named by Kunth (1824) in the sixth volume of Humboldt and

Bonpland’s Nova Genera et Species Plantarum [5 Jan 1824-pertinent dates from International Plant Name Index

(wwwipnixHg) and Taxonomic Literature II (Stafleu & Cowan 1976-1988)] with a complete description. An

illustration of the sole species, Lindleya mespiloides Kunth, was published in Apr 1824 (fig. 562). The genus was

named for the then young John Lindley (1799-1865), British botanist, student of the Rosaceae and Orchida-

ceae, who went on to be professor of botany at the University College London (1829-1860).

However, prior to this, the name Lindleya had been used as a generic name several times. The first use of

Lindleya as a genus name was by Nees von Essenbeck (1821), in a paper (21 May 1821) determining specimens

from the Brazilian collections of Spix and Martinus. His Lindleya was a genus in the Theaceae, which Index

Kewensis states is based on a species Lindleyafruticosa Nees ( =Laplacea semiserrata Cambess.). Nees’ Lindleya,

however, is a homotypic synonym of a slightly older Wilkstroemia Schrader (5 May 1821) and thus is a superflu-

ous name. Kunth in Humboldt, Bonpland loc. cit. (25 Feb. 1822) recognized this genus as Laplacea Kunth.

Kunth himself had previously used the generic name Lindleya twice. His first use was on plates of two

species that were published in volume five of Nova Genera et Species Plantarum. According to Taxonomic Lit-

erature II, the fascicle of plates bearing the name Lindleya (plates 479-480) were published on 25 Feb. 1822,

while the text, (vol. 5: 361-367) was not published until 24 March 1823, however, in the text, the species were

placed in the genus Casearia (Flacortiaceae). The use of generic name Lindleya on the plates has been consid-

ered as a nomen nudum by Index Kewensis and Index Nominum Genericorum database and l.P.N.I. But the

International Code of Botanical Nomenclature (McNeil et al. 2006) Art. 42.3 considers an illustration with

analysis (i.e., with separate figures showing details to aid with identification) as being acceptable in place of a

written description or diagnosis for a species description. But Art. 42.1, 42.2 notes that this can apply only if the

genus is monotypic. As there were two species illustrated, the use of Lindleya here can be considered as a nomen

Kunth (1822) again u

le and Tiliaceae (20 Apr, J

m. 46D) between Theo- j

broma Linn, and Guazuma Plum., in Sectio [subfamily] II “Buttneriaceae verae” of Buttneriaceae (once Stercu- ,

liaceae, now Malvaceae s.l.). This is purely a listing of a known or proposed name and again is a nomen inval, j

without description given, implied or cited.

Rydberg (1908a) in his Notes on Rosaceae preceding his North America Flora treatment (Rydberg 1908b)

listed the previous use of Lindleya by Kunth and Nees (as noted in Index Kewensis) and offered a new generic

name Lindleyella for the Rosaceaous genus, and described a second species L. s chiedeana Rydb., characterized

by more obovate leaf blades and longer, more obovate petals. In 1940 Fedde, for some reason, rejected the ear-

lier Lindleyella of Rydberg (1908), in favor of a later described Lindleyella Schlechter (1914), a genus of Orchida-

ceae with five species that were named from 1914 to 1924, and substituted a new name, Neolindleyella Fedde for

Rydberg’s genus. As to why Fedde would choose a younger name ( Lindleyella of Schlechter 1914) against an

older name ( Lindleyella of Rydberg 1908) is unknown as it goes against the rules of priority, which he so

strongly espoused. Fedde’s work on botanical nomenclature had become such a disruption on nomenclatural

stability that the provision for conservation of generic names was enacted during the Vienna Congress in 1905

(Briquet 1906).

In 1930, at the fifth International Botanical Congress (Briquet 1935), the rule concerning homonyms was

altered. Prior to that time, a later homonym could be accepted as a valid name if the earlier name had become a

synonym and was not being used. The fifth congress maintained strict nomenclatural priority and rejected

later homonyms with the understanding that all well known generic homonyms, as far as possible, should be

To this end, a systematic search was made to validate later homonyms via generic conservation (Rehder et

al. 1935). The task was assigned to various botanists each responsible for genera based on alphabetical group-

ings. Genera beginning with the letters L through P were assigned to Dr. Rudolf Mansfield of the Botanical

Henrickson, Systematic of Lindleya

Museum of Berlin. Through his work, Lindleya Nees was rejected in favor of Laplacea H.B.K. (nom. cons.).

Kunth’s use of Lindleya as a replacement name for Casearia was considered a nomen nudum, and Lindleya of

Kunth in Humboldt and Bonpland (1824) was conserved against Lindleyella Rydb. (Green 1940). Thus the cor-

rect name for the genus, under the present rules, following that round of conservation, would be Lindleya

H.B.K., or following those who object to the H.B.K. designation, Lindleya Kunth in Humboldt & Bonpland

(nom. cons.) or just Lindleya Kunth following ICBN 2006, Art. 46, ex. 9, note 1 (McNeill et al. 2006).

Of interest, in 1858, J. Agardh, in his Theoria Systematis Plantarum, placed Lindleya in its own family

Lindleyaceae J. Agardh. In the same paper, he also recognized Cercocarpaceae J. Agardh, Coleogynaceae J.

Agardh, and many other new families of dicotyledoneae.

>RPHOLOGY AND ANATOMY

» few-stemmed shrub (Fig. 1A). In more mesic habitats, the main

s an obovoid-spreading shrub to 2-4(-5) m tall with the uppermost

branches either ascending or arching outward. In dry habitats the plants form more rounded, densely and di-

varicately branched shrubs to 1.3 m tall and wide. These arid-land shrubs may be misshapen due to browsing

by goats, or somewhat thorny due to the placement of dried lateral stems.

Stems . — Stems are heteroblastic. Young long-shoot stems are initially maroon, glabrous, with internodes

7-15 mm long. They develop a single band of vascular tissue and dense strands of primary phloem fibers. The

cortex and pith cells both develop thickened walls. Secondary growth initiates soon after stem development.

Short-shoots develop from the nodes. As they mature they may develop into variously branched short-shoot

systems to 1— 5(— T) cm long with intemodes 0.4-1.5(-5) mm long. The epidermis of young stems is soon re-

placed by a light gray periderm. Older branches have a smooth gray periderm marked by distinctive horizontal

lenticles (Fig. ID).

Stipules . — Paired yellow-green stipules occur at each long- and short-shoot node at the margins of the

canaliculate petioles. They are typically acicular, 0.4-0.9(-2.2) mm long and are bordered on two sides by well-

spaced or crowded, distinct, reddish-maroon multicellular glands to 0.2-0.3 mm long (Fig. 3A). When shorter,

which decrease in size acropetally, make the stipules appear more deltate. When longer,

icular with well spaced marginal glands. The stipules mark the separation of the

broadened leaf base from the canaliculate petiole and occur just below the zone of leaf abscission. The petioles

mm long 2.5 mm wide to mesophylls 55 mm long and 17 mm

owly oblanceolate, narrowly obovate to obovate. The apices

range from narrowly acute, obtuse, rounded, to retuse often with an acuminate to mucronate tip. Any one plant

may have some leaves with acute tips, other leaves with rounded tips. They are cuneate at the base with the

margins forming decurrent narrow wings along the short, canaliculate, yellowish petioles. The margins are

crenate; the crenations are gland tipped, with rounded distal and proximal margins. The crenations are usu-

ally uniform in distribution, but are more widely separated along the lower blade margins. The leaf blades are

glabrous throughout. The marginal dark red-maroon glands are compressed ovoid, sessile, the marginal ones

ascending towards the tip, the distal ones erect, 210-290(-350) pm long, to 100-120(-160) pm wide at the

broad base. The gland’s marginal cells are anticlinally elongated. The glands fall from older leaves.

The dorsi-ventral leaf blades are generally ascending, with the adaxial surface slightly concave (Fig. IB).

The blades range from 220-360(-430) pm in trans-section thickness and are 350-400(-430) pm thick at the

midvein. The midvein is raised on both the adaxial and abaxial surface proximally, but only on the abaxial

surface in the mid section and distally, except in the thickest blades where the midvein is not raised (Fig. 2A).

narrow external cell with rather dense cytoplasm and a much larger internal water-storage cell. The outermost

wall of the upper epidermal cell is about 5.5-7 pm thick, with the cuticle 4-5.6 pm in thickness. The abaxial

Journal of the Botanical Research Institute of Texas 6(2)

epidermal and guard cells and the structure is difficult to sort out. The epidermal cells are larger and have a

smaller external cell, but the guard cells do not. The adaxial leaf mesophyll consists of 2(-3) layers of crowded

palisade cells. A palisade-type cell orientation may or may not occur in the abaxial mesophyll but cells are not

as dense. The primary, secondary, and usually tertiary veins have vascular cambia. A mass of support cells

develops adaxial to the phloem. In some leaves these consist entirely of collenchyma cells, in others collen-

chyma mixed with fibers, and in the primary and secondary veins consist of masses of lignified fibers. Usually

:curs both above and below the primary-secondary-tertiary veins

rntinue to the fourth and fifth order veins (seen throughout the

a collenchymous

times bundle sheath e

345

Fig. 2. Leaf structure of Lindleya

specimens from Coahuila-Nuevo Leon (B-H), Hidalgo (l-L), and Puebla-Oaxaa (M-Q). B. Henridaon 22056 (TEX)-NL, C Henrickson 6225 (TEX)-Coah.

D. Johnston etaL 1 1465 (LL)-Dgo. L Steward 294 (OH)-Coah. F. Steward 378 (GH)-Coah. €. Henridaon 22062 (Tex)-LL. Stanford etai 626 (GH)-Tamp. I.

Gonzalez 21443 (F)-Qto. J. Gonzalez 2401 (LL)-Hgo. K. Gold 324 (TEX)-Hgo. L Moore 2481 (GH)-Hgo. M. Salinas 6 Solis 3238 (TEX)-0ax. N. Tenorio 6882

(TEX)-Pue. 0. Dorado 8, Salinas sj). (Tex.)-Pue. P. Tenorio 18341 (TEX)-0ax. Q. Tenorio Gonzdlez 18341 (TEX)-0ax. B-S. Leaf clearings. R. McVaugh 10346

(US). S. Gonzalez 1443. (MEXU). Sale in A = 0.3 mm, in B-Q = 20 mm, in R-S = 1mm.

Journal of the Botanical Research Institute of Texas 6(2)

states of Queretaro, Hidalgo, Puebla, and Oaxaca). The mesophyll has scattered cuboidal crystals that are more

common in bundle-sheath extensions.

Venation, as seen in cleared leaves is pinnate, weakly brochidodromous to semicraspedodromus (Ellis et

al. 2009). The primary vein is straight, symmetrical, and moderately thickened (Fig. 2R-S). Secondary veins

are irregularly alternate, extending from the primary vein at angles of 20-40 degrees. In narrow leaves they

tend to arch slightly towards the tip hut do not extend to the leaf tip. The secondary veins exhibit moderate

branching towards the margins, often forming through connections with adjacent secondary veins one or

more series of arches below the margins, with the crenations served by tertiary or smaller veinlets. Intersec-

ondary veins are composite and muched branched, connecting to the lower portions of adjacent secondary

veins. Tertiary veins form random reticulations, mostly at right angles to the secondaries; they are percurrent

and forked. There are 5 orders of vein branching; the areoles are well developed, random, 4-5 angled, the

smallest veinlets are 1-3 times branched. Venation of small leaves differs in that the primary and secondary

veins are much thicker than the tertiary and smaller veinlets, and secondary veins may not connect with super-

adjacent veins.

Stomata are restricted to the lower (abaxial) leaf surface and range in number from 96-175 per mm 2 in

small leaves ca. 13 mm long, and 140-175 per mm 2 in larger leaves to 27 mm long. Stomata do not occur below

the veinlets with bundle sheath extensions. The stomata are usually subtended by four subsidiary cells. Subsid-

Inflorescences . — Flowers are usually solitary terminating new growth of the season (Fig. 1C), but occa-

sionally, with vigorous new growth, more than one flower will develop, with the subtending flower(s) develop-

ing from lateral shoot(s) from a subterminal node in the new growth of the same season. In such conditions the

flowers will appear to form simple corymbs or three-flowered cymes (Fig. IE). The uppermost, reduced leaves,

that form with the season’s growth are typically crowded below the flower and extend onto the pedicel. The

uppermost cauline leaves are typically only 8-18 mm long, petiolate with slender to deltate, gland-margined

stipules. They have canaliculate petioles and gland-margined, glabrous blades (Fig. 3B). The uppermost leaves

are reduced to linear to lanceolate bracts to 3-5 mm long, 0.5-1.2 mm wide, without stipules but with glands

continuing from the petioles to the blade margins. True pedicels, when present, may be up to 2 mm long. The

terminal flowers disrupt terminal growth of a shoot, continued growth occurs through development of sub-

:rminate in thick-walled, coriaceous, obconic, hypanthia topped with 5

sepals, 5 petals and about 20 stamens (Figs. 3C, 5A). The ovaries are not attached to the hypanthia except at

their sessile bases. The free portion of the hypanthia is about 0.7-1 mm in thickness. The outer hypanthial

surface is light green, glabrous and shiny. The inner surface is yellowish-green in color, nectariferous, some-

times vertically ribbed (the ribs reflecting the position of stamen traces), with a somewhat expanded inner rim

subtending the filaments.

Sepals . — The hypanthia terminate in five coriaceous, light greenish, broad-based, ovate to oblong-ovate,

lance-ovate, usually acuminate, imbricate sepals (Figs. IE, 3B). Where the sepal margins overlap, underlying

sepal margins are membranous to 0.5(-0.9) mm wide and are sometimes cordate at the broad sepal base. The

external or overtopping sepal margins are usually not membranous but usually have distinctive reddish glands

similar to those found on leaf margins. The sepal tips usually terminate with a reddish gland. The sepals may

be glabrous throughout the abaxial surface or variously villous distally. The inner surfaces typically are gla-

brous in the lower half but distinctly villous distally and along the distal margins, with slender, crinkly, white

hairs to 0.6(-0.9) mm long. There are some differences in sepal size throughout the range of the species; plants

from Oaxaca-Puebla tend to have shorter sepals, but this is not consistent as similar short sepals occur in small

flowers throughout its northern range. Sepal size usually reflects overall flower size, and flower size can vary

with environmental conditions. The sepals usually persist on the rim of the hypanthium as ascending or re-

flexed structures (Fig. 3G).

Petals .— The five broadly obovate, broadly clawed petals are borne equally along the hypanthium rim al-

ternate to the sepals. The petals are spirally arranged in bud and are oblique distally with the portion of the

347

by dry hypanthium and sepals. Note the style bases have been split apart. H. Post-dehiscent fruit with some sepals removed showing dehisced carpels.

I. Longitudinal view of mature fruit with carpel interior exposed showing seed. J. Seed. K. Embryo. Magnifications as indicated. A-E ( Henridaon 22 1 12,

TEX); G-K [Henridaon 22222, TEX).

petal that develops under adjacent petals being much larger than the portion exposed in the bud (Figs. IF,

In most petals the right half of the petal is interior in bud and largest. But this will vary from flower to flow<

one plant, with occasional flowers having the left half of the petal interior and larger than the right half,

petals are white, waxy appearing, well veined, thickened at the base, and thinning towards the somet

crinkled margins. They spread outward at anthesis (Fig. IF) and tardily abscise after anthesis. The mid

often terminates in a single gland When clearing the petals, the thick basal portion of petal often stains dark—

Atubvecia . — Flowers typically have 20 stamens borne at the inner rim of the hypanthia in one series or

occasionally in two weak series with occasional smaller stamens born inside larger outer series of stamens

(Fig. 3D). The inner and outer stamens usually develop opposite the petals with the interior stamen having a

shorter incurved filament. The white filaments are acicular, very broad at the base, tapering distally. The fila-

ments are attached to the mid-portion of the versatile anthers (Figs. 3D). Anthers are introrse in bud, 1.5-4 mm

long, oblong, apiculate at the tip, with V

anther sacs, and the medial septa betwe<

The gyn.

icated basal lobes. Anthers are light yellow, longicidal, with 4

inther sacs appears to be secretory (glandular) as it is dull red-

dish in color (in dried anthers) and stains strongly in microscope slides. Within the buds, the outer, larger

stamens are erect with straight anthers, while the inner stamens have inflexed filaments as their anthers de-

velop further within the crowded hypanthial cup. These inner anthers are typically bent below the attachment

to the filaments (Figs. 3D). They usually retain this shape at anthesis.

ound ovary is 5 carpeled, ovoid, glabrous, slightly 5-angled in transverse section,

the receptacle apex (Figs. 3C, 5A) and the carpels are united ventrally up to the

level of ovule insertion forming a thick-walled compound ovary with axile placentation (Fig. 6D). The carpels

are laterally united from the dorsal edge to near the ventral margin. The carpels, however, are not connate

centrally where a five-lobed opening extends to the base of the ovary (Fig. 4A). Sterling (1966) noted that this

is a carpel fusion pattern similar to that found in the Maloideae. He also noted that occasionally the carpels will

not be fused ventrally. Each carpel contains two apical, pendent, collateral ovules, about 0.7-1 mm long at-

tached at the inner tip of the locule and receiving a downward-oriented vein from the adjacent ventral trace

(Fig. 5A). The short, thick funiculus appears to function as an obtruator. The ovules have a broad outer integu-

ment, a thinner inner integument, and the embryo sac is quite large with a crassinucleate nucellus.

The carpels are free from the inner hypanthium walls except at the very base. The five styles are separate,

terminal, slender, cylindrical and are obliquely expanded at the stigmatic tip (Fig. 3E). A split develops across

the oblique tip, exposing papillate interior tissue. Dried styles may persist on the fruit, or they may break off

above the bases (Fig. 3F-I).

Floral vascularization. The pedicel contains a single cylinder of vascular bundles (Fig. 5E-1). At the

base of the hypanthium, 10 (occasionally more) traces separate from the central traces and extend up the hy-

panthium (Fig. 5E-3-4). These produce an irregular series of lateral traces that extend horizontally into the

hypanthial tissue. The hypanthial traces branch in the mid toupper hypanthium to produce additional stamen

traces (Fig. 5D). The traces opposite the sepals typically divide to form 2 stamen traces that then pass parallel

branch near the mid-hypanthium to form additional stamen traces, or one or moresumen tr^may^eparatt

from the trace near the top of the hypanthium. In addition the trace opposite the petal divides near the top of

the hypanthium to produce two lateral sepal traces, one to each of the two adjacent sepals (Fig. 5B, D). At the

run oi the hypanthium, the 5 sepals then each receive one medial trace and two lateral traces derived from

adjacent petal traces. The lateral traces each divide int.

ree separate traces at the base of the sepal,

ore parallel traces that extend up through the sepal base and branch above (Fig. 5B).

.single basal trace that quickly divides into five to seven traces. These continue to

3 the petal in a pattern as shown in Fig. 5C. The 20 stamens each receive

separate from the original 10 hypanthial traces at some point in the mid to

The petals each receive

branch and anastomose further

single traces that, as noted abov

upper hypanthium.

.p . ^' e ^ entra ^ tratesret ^ a ' n | n S af . er the initia! hypanthia! traces diverge, form into five central packets (Fig.

cf, r i maSS ’ c Ve dor5al ‘ races div “8 e opposite the sepal traces receiving tissue from two adjacent

vascu ar tissue (Fig 5E 6). These leave behind five dense masses of vascular tissue that becomes the

W ™er ventral carpel traces (Fig 5E-6). The dorsal paces continue upwatd akmg the ovary periphery. They

grve off senes of branch traces to the ovary wall, but disposition of these late J traces PcLLd by the

demtely staining, tannin-containing cells of the developing ovary wall. The ventral vascular tissue forms ten

As noted by Sterling (1966), no wing traces diverge from the ventral traces at this time

one .SToTon^nh b gl ° bOSe ' 5CarpeUed ' W °° d * loculiddal -P-Ksubtended in the lower

I of upper portion of two ovules from B. All from Henrickson 22 1 18, TEX (Galeana area, Nuevo Leon, Mexico). Bar

At maturity, the carpels dehisce loculicidally directly through the ventral traces, splitting the ventral traces

and the base of the style through a suture that is visible even in the developing ovary wall (Fig. 6A). The dehis-

cence splits the style bases in half, and continues onto the distal portion of the abaxial fruit surface (Fig. 3G-

H). The inner lateral walls of the carpels are smooth and cartilaginous (as in an apple), lined with a single layer

of macrosclereids 100-120 pm thick and subtended by a thick layer of brachysclereids each 35-65 pm in diam-

eter with lignified walls 11-22 pm thick (Fig. 6E). The highly lignified tissue extends 1.5 mm in radial thick-

ness in the triangular segments between the locules. Only the outermost 0.5 mm of the abaxial-most portion

of the triangular carpel segments is not lignified.

Seeds.— Two compressed, half-ovoid, dull-brown seeds are produced in each locule. The seed shape con-

forms to the locule space, being straight along the ventral edge and rounded on the dorsal edge, with one flat-

tened surface (where contacting the adjacent seed) with the outer surface convex (Fig. 3J, 6G). The seed coat is

crystals (Fig. 6F). The embryo is oriented with the hypocotyl superior to (i.e., above) the cotyledons. The em-

bryo, occupies about 70 percent of the total seed length, leaving a thin wing, 0.2-0.5 mm wide, at the dorsal

margins. The embryo consists of two compressed, oblong cotyledons and a smaller, obovoid hypocotyl (Fig.

3K). Endosperm is absent at maturity. The seeds are wind dispersed. Upon germination the cotyledons form

the first seed leaves of the seedling.

TAXONOMIC TREATMENT

LINDLEYA KunthNov. Gen. Sp. [H.B.K.] 6:240 (ed. qto.); 188 (ed. foL). 1824 (nom. cons.), nonUmfleyaKunihNov.

Evergreen, multistemmed shrubs; periderm gray, smooth. Stems heteroblastic, tending to form shortened axil-

lary spurs in arid conditions. Leaves simple, alternate; leaf bases short, the stipules acicular to debate, maroon,

sometimes with marginal glands; leaf blades narrow to broadly oblanceolate to obovate, ovate, acute to round-

ed, sometimes retuse at tip, cuneate with the margins decurrent on short petiole, the margins closely crenate,

the marginal teeth terminating in distinct glands, the blade coriaceous, shiny green, glabrous on both surfaces.

3S0 Journal of the Botanical Research Institute of Texas 6(2)

Henrickson, Systematics of Lindleya

351

major veins with bundle sheath extensions, venation brochidodromous to semicrospedodromus. Flowers

complete, perigynous, usually solitary and terminal on long and short shoots, these sometimes forming termi-

nal corymbs when the shoots aggregated or on long-shoot stems; bracts reduced, linear, gland-margined,

borne on upper pedicel; hypanthia obconic, hemispherical in fruit, green, glabrous outside, nectariferous and

yellow-green inside, coriaceous; sepals 5, imbricate, debate to ovate, acute to acuminate, glabrous outside, vil-

lous near tip inside, somewhat coriaceous except along overtopped margins, spreading, persisting on fruit;

petals 5, borne at the rim of the hypanthium, obliquely obovate, asymmetrical, white, spreading at anthesis,

tardily deciduous, aromatic; stamens usually 20; filaments subulate, broadened at the base, borne at the inner

rim of the hypanthium in a single (rarely two) series; anthers lanceolate, large, versatile, longicidal, yellowish;

ovary; styles terminal, distinct, the stigmas terminal, oblique; ovules 2 per carpel, collateral, apically attached,

pendent, the micropyles superior, the funiculi thickened, the tissue serving as an obtruator. Fruits spheroidal,

Journal of the Botanical Research Institute of Texas 6(2)

woody, loculicidally dehiscent capsules, the lower third surrounded by the coriaceous, persisting hypanthi-

um, the carpels dehiscent distally along ventral and dorsal sutures, with thick, inwardly cartilaginous valves;

seeds 2 per locule, compressed, narrowly winged abaxially, brown, the seed coat thin, the embryo with oblong-

ovate, flattened cotyledons, the hypocotyl small, superior; endosperm absent. With one species.

Lindleya mespiloides Kunth, No

22(3):259. 1908. Type: MEXICO. Hn

ENE of Actopan, Hgo.), May (holoti

lotype: NY!). Distinguished by Rydberg as

ien. Sp. [H.B.K.] 6:240. 182!

;o: Inter La Puente de la Madre de D

specimen of L. mespiloides with very large features. Other specimens from

smaller leaves. As noted al

N W of that of L. mespiloides. As it is distinguished only by quant

Erect, much-branched, evergreen shrubs to small trees l-3(-5) m tall, in dry habitats forming small, tightly,

divaricately branched shrubs with many short shoots, in mesic habitats forming erect-ascending, moderately

branched, tall shrubs; stems heteroblastic; long-shoot branches with intemodes 7-15 mm long, glabrous, ini-

tially maroon, developing a close smooth, gray periderm; short-shoot branches l-5(-7) cm long, variously

branched, with intemodes 0.4-1.5(-5) mm long. Leaves with petioles l-2(-4) mm long; stipules 0.4-2.2 mm

long, acicular to deltate, maroon, when longer more attenuate and bearing marginal glands; leaf blades nar-

rowly oblanceolate, oblanceolate, spatulate, obovate, sometimes elliptical-oblanceolate, rarely somewhat

ovate, (3.5-)ll-32(-55) mm long, (1.6-)4-13(-21) mm wide, acute, acute-acuminate, rounded to emarginate

at the tip, narrowly cuneate with the margins forming wings above the short petiole at base, the margins

closely crenate to crenulate with 5-10(-15) teeth per cm of margin, the teeth each terminating in a maroon,

conical gland 0.1-0.3 mm long, the blades coriaceous, glabrous throughout, shiny, dark green, slightly concave

above, more yellow-green beneath, the midvein yellowish and raised on both surfaces. Flowers terminal, soli-

tary on short, leafy shoots, rarely in terminal 3-flowered racemes, the subtending leaves reduced, the upper-

i to linear, gland-margined bract(s) to 5 mm long; pedicels 1-2 mm long, glabrous,

4-5 mm wide (to 7 mm wide pressed), green, glabrous outside, yellowish-green,

„ , m : the ovar y- the distal rim ^ghtly expanded; sepals 5, ± imbricate, deltate to ovate, !

ally^Uhr^t^^no^s mm W ' de ’ *“'* “* COriaCeOUS ^ 8rKn med ''

glabrous at the base but densely villous n^r t^e tipinsitk^ind ^tenTiUous^dlL^hrou^ouu^ffwrgins* tfw ]

s With conical glands along the margins, the thin inner margins somewhat cordate at the

urity; petals broadly obovate to ob-

bowers W-)22-40 mm tot,! diameter], obliquely rounded to emarginate at the dp, broadly cuneate above a

very short and broad claw at the base, firm, thickish, waxy, white, spreading, emitting a street aroma remain-

t™””" -tfter anthesis, eventually deciduous, stamens usually 20; filaments linear-subulate, dis- I

n long, to 0.5-0.8 mm broad at the expanded bases, borne in the inner rim of the hypan-

u e with the shortest laments occurring inside a larger filament opposite the

a ^ 2 ^ 3 5( - 4> r m “ ^ p - co " I

itue, reddish to usually dark yellow; ovary two-thirds +

most leaf(s) often ri

ectariferous i

base, the sepals spreading in flower and persisting around the fru

--orbicular, obliquely asymmetrical distally, contorted in bud wit!

:t, (2-)3.5-7(-8) n

n, of variable length and s

date, the basal lobes often spreading or folded -

TXT Z " a, h “ Rnded t,ps - Fruils of ' TOod * ovoid-spheroidal, long-persistent capsules, 7-10

adlZienUn^dis, Tft T° Unded by ““ I”*** ‘W—W™. the carpelsLh locuB-

y ’ 8 ’ 8 ark red to maro °n, drying dark brown, glabrous; ovules 2 per locule, pendent;

Henrickson, Systematics of Lindleya 353

seeds two per locule, oblong-ovate in outline, flattened, 4.3-6 mm long, 1.8-2.6 mm wide, narrowly winged on

the curved abaxial margin, the adaxial margin straight; cotyledons ovate, endosperm absent, radical superior.

The species is characterized by its shrub to small tree growth habit with smaller plants of more xeric

habitats often developing short-shoot branches and a somewhat thorny aspect, by its smooth gray bark, by its

small to moderately large shiny green, mostly oblanceolate, acute to round-tipped leaves with gland-tipped

finely crenulate margins, by its large terminal, sweetly aromatic flowers with a thickened obconic hypanthium

that bears 5 glabrous sepals with thin villous-ciliate margins, 5 large, white, obliquely obovate to somewhat

orbicular petals, and ± 20 stamens with subulate filaments and large versatile anthers, and by its superior,

5-carpelled ovary with 2 suspended ovules per locule, and 5 separate styles. The fruit are globose, woody, locu-

licidally dehiscent capsules with each locule producing two flattened seeds, each with a short wing along its

outer margin. The flowers are conspicuous and remain on the plant after the anthers have shed their pollen

creating a conspicuous floral display. The fruits are often long persistent, allowing recognition of the species in

the field.

The species has three regions of distribution (Fig. 7): northern Oaxaca and adjacent Puebla, where it oc-

curs oak-pine woodlands, chaparral and thorn scrub in association with species of Malacomeles, Vauquelinia,

Comarostaphylos, Rhus, Quercus, Juniperus, Pinus, Acacia, Leucanea, Beaucamea, and Yucca etc., from 1900-

2400 meters elevation; in Hidalgo and adjacent northern Queretaro, where it again occurs from pinyon-oak-

juniper woodland, chaparral association with many of the same genera. Its largest distribution is in the Sierra

Madre Oriental and Chihuahuan Desert region from southwestern Tamaulipas, southern Nuevo Leon, north-

ern San Luis Potosi, northern Zacatecas, northeastern Durango, the southern half of Coahuila and adjacent

southeastern Chihuahua where it occurs in pinyon-juniper woodland, chaparral and canyons in desert scrub

often in limestone, rarely gypseous or sandstone substrates in association with species of Rhus, Garrya, Ceano-

thus, Fraxinus, Cercocarpus, Gochnatia, Vauquelinia, Berberis, Arctostaphylos, Quercus, Arbutus, Juniperus, Agave,

Yucca, Dasylirion, Ungnadia, Leucophyllum, Condalia, Mimosa, Mortonia, Foresteria, Acacia, and Pinus from

1100-2700 m elevation. Flowering typically occurs in May but may occur from March to September usually

following rains, with occasional flowering occurring throughout the year.

Throughout the wide range of the species some notable variation occurs. Most apparent is the variation in

leaf size. Most specimens from Hidalgo and Oaxaca have moderate large leaves 18-55 mm long, 8-18 mm

wide. In contrast, most specimens from Coahuila and Nuevo Le6n have smaller leaves, 10-26 mm long, 3.5-8

mm wide. The separation is by no means complete, as some specimens from both Hidalgo and Oaxaca also

have small leaves. Leaf size corresponds to habitat, as plants in dry exposed areas have very small leaves that

contrast with larger leaves of plants of nearby less-arid, shaded sites. As noted above, plants from Hidalgo,

Queretaro, Puebla and Oaxaca have bundle-sheath extensions extending to the 4th and 5th order of vein

branching in contrast to only the 1st, 2nd, and in part to the 3rd order of branching in more northern range.

Specimens observed from Puebla and Oaxaca also tend to have shorter sepals measuring 3.0-5.2 mm long. In

contrast, sepals from Hidalgo and northward specimens tend to be larger, (3.0-)4.5-7(-9) mm long, but there

is considerable overlap, with sepal length corresponding to flower size and being variable even on a single

specimen. Flowers range in total diameter (petal tip to petal tip) from 17 to 40 mm. Fruits also vary in size, with

occasional specimens having fruits much smaller or larger than average.

PHYLOGENETIC RELATIONSHIPS

The placement of Lindleya within the Rosaceae has been problematic. Numerous subfamilial and tribal classi-

fications of the Rosaceae have been proposed (Focke 1888; Hutchinson 1964; Schulze-Menz 1964; Cronquist

1981; Takhtajan 1987 1997, 2009; Kalkman 1988, 2004; Thome 1983, 1992) etc. and most all retain four sub-

families, (some older treatments have five subfamilies with the inclusion of the Chrysobalanoideae, or six with

the Neuradoideae) that corresponded well to fruit types: the Spiraeoideae, with follicles, (rarely achenes — Ho-

lodiscus, or capsules — Vauquelinia, Lindleya)- Rosoideae with achenes (rarely druplets — Rubus); the Amygda-

loideae with drupes ( Prunus etc.), and the Maloideae (Pomoideae) with pomes ( Malus etc.). Within these sub-

families, the treatments differed in the placement of genera within tribes. See summary in Potter et al. (2007).

Journal of the Botanical Research Institute of Texas 6(2)

There has always existed a problem group of six genera with woody fruit and winged seeds some with

follicles, others with capsules, consisting of: Quillaja, (plant polygamo-dioecious, fruit of five radiating woody

follicles each with many (10-16) seeds in two series per carpel— northern South America); Kageneckia (dioe-

cious, fruit with five separate follicles with many seeds in two series per carpel— western South America);

Vauquelinia and Lindleya, (dry capsules with two-winged seeds per carpel— Mexico); Exochorda (plant polyg'

amo-dioecious; fruit with five, rather compressed woody follicles— Eurasian) and Lyonothamnus (two follicles

ing x=9, the Amygdaloideae x=8, the Rosoideae

x=7, 9 (rarely 8) and the Maloideae x=17. The high chromosome number in Maloideae caused Sax (1931, 1932,

1933) and later Stebbins (1950, 1958) and many others (see Phipps et al. 1991; Rohrer et al. 1991, Cronquist

1981) to promote the view that the Maloids arose via paleo-allopolyploidy from x=9 Spiraeoideae and x=8

Amygdaloideae ancestors or from within the Spiraeoideae (Gladkova 1972).

In 1976 Goldblatt contributed chromosome numbers of several of the problem genera noted above. Of

these Exochorda was found to be x=8, Kageneckia and Lindleya x=17; Vauquelinia x=15; and Lyonothamnus x=27

and Quillaja x=14. The high numbers in these taxa again gave clues that their relationship may lie with the

Maloideae, and Lindleya and Vauquelinia were subsequently transferred to the Maloideae (Pyroideae) by

Thorne (1983) and Takhtajan (1987). The 5-carpelled, dry-fruited Exochorda (x=8) was considered by Goldb-

latt (1976) to belonging to the x=8 Prunoideae, which has been supported by molecular data (Morgan et al.

1994). Kageneckia and Quillaja, which have similar appearing fruits, remained in subfamily Quillajeoideae

(Thorne 1983; Takhtajan 1987).

Floral morphology Data . — Sterling (1966), on the basis of ovary morphology, considered the Quillajeae

(containing Exochorda, Kageneckia, Lindleya, Quillaja, and Vauquelinia) to have sharp differences in the gynoe-

cium structure and number and orientation of ovules. He noted that Lindleya had ovaries with complete lateral

intercarpellary fusion of a type characteristic of the Maloideae, but with minimal hypanthium fusion. He also

noted that the carpels of Vauquelinia were also maloid in nature though fused only basally and ventrally while

being separate laterally.

Data from Rusts . — Savile (1979) summarized the use of fungus-host relationships in plant phylogeny. He

notes that cedar-apple rusts Gymnosporangium occur throughout Pomoideae (Maloideae) with species also

known from Myricaceae, Hydrangeaceae and from two genera of Spiraeoideae ( Vauquelinia and Porteranthus —

now Gillenia ). He cites Gymnosporangium vauqueliniae Long and Goodding (1939) on Vauquelinia califomica

from Arizona and interestingly, Gymnosporangium externum Arth. & F. Kern in Arth., on Gillenia in eastern

United States, which had been known since 1903. The on-line “Fungal Database” also gives references of Gym-

ditional collections of G. externum on both species of Gillenia in the eastern United States (http://nt.ars.grin.

gov/fungaldatabases). Gymnosporangium has not been reported on Lindleya.

r Data . — If we stop here and look at the data accumulated by the 1980s, we

number as members of the then recognized Maloideae, and has

some floral features in common with the Maloideae. We also have data from Cedar-apple rusts linking the re-

lated Vauquelinia with the Maloids. But confusingly, the rusts also occurred in another Spiraeoid, namely Gil-

lenia. But as there was evidence that indicated that Vauquelinia and Lindleya were related to Maloids, there was

no data indicating whether they were derived from the fleshy-fruited Maloids, or if they were basal to the fleshy-

fruited Maloids. When we monographed Vauquelinia (Hess & Henrickson 1978), we had no evidence pertain-

ing to the relative placement of Vauquelinia to the fleshy-fruited Maloids, so in that paper we concentrated just

on species relationships. In the 1991 symposium on the Evolution in the Maloideae (Rosaceae) published in

Systematic Botany, Vauquelina and Lindleya was not mentioned nor included in data sets (Phipps et al. 1991;

Robertson et al. 1991). But by the mid 1990s, molecular data began to provide answers and laid the ground-

work for a new phylogeny of the Rosaceae.

ar data presented by Morgan et al. (1994) from chloroplast rbcL sequences

vidence for a needed subfamilial rearrangement of the Rosaceae. Their paper

divided the family along the same lines as cytological data. Their data, however, showed that the old Rosoideae

was polyphyletic, that thex=7 genera ( Filipendula, Fallugia, Geum, Waldsteinia, Potentilla, Fragaria, Agrimonia,

Rosa, Rubus and the x=8 Alchemilla) formed the core of the Rosoideae and the x=9 genera formed four distinct

Journal of the Botanical Research Institute of Texas 6(2)

groups: the Neviusia, Rhodotypos group; the Cercocarpus, Purshia, Lyonothamnus group; and the core Spiraeoi-

deae with basal Spiraea-Aruncus-Holodiscus group, separate from a derived Physocarpus-Neillia group, and an-

other Sorbaria group associated with Chamaebatiaria including the achene-bearing Adenostoma. Exochorda fell

into the x=8 Amygdaloideae. Their data excluded the South American Quillaja from the Rosaceae.

Of significance to this paper, their data showed Kageneckia, Undleya and Vauquelinia were basal to the

remainder of the x=17 Maloideae, i.e., their data showed Vauquelinia and Lindleya, with capsular fruits, and

Kageneckia with follicle-like fruits, as remnants of a clade that have given rise to the core Maloideae. That is,

they were basal to the Maloids, not dry-fruited derivatives of fleshy-fruited Maloids.

Campbell et al. (1995), using the internal transcribed spacers (ITS) of the nuclear ribosomal DNA region,

studied phylogenetic relationships within the Maloideae. Their study, however, concluded that the Maloideae

was not monophyletic. They also noted that Vauquelinia forms a well supported clade with fleshy-fruited Erio-

botrya and Raphiolepis that is the sister group to the remainder of the Maloideae. These data, however, were

badly skewed, as their “ Vauquelinia ” sample was actually Raphiolepis collected at the University of Arizona

campus by a graduate student, vouchered by a specimen collected twenty years earlier in 1975. Their data set

did not include material of Lindleya, Kageneckia, and as it turned out, also did not contain Vauquelinia.

Data presented by Evans et al. (2000) on the “Granule-Bound Starch Synthase 1” gene (GBSSI) provided

further insights into the phylogeny of Rosaceae. The portion of this nuclear gene used (near the 5’ end) consists

of seven complete, short exons, and parts of two other exons alternating with non-coding introns. While all

other diploid families in which this gene has been used, have only one GBSSI sequence, all diploid Rosaceae

have two distinct sequences (designated as GBSSI-1 and GBSSI-2) that differ in the length of, or presence or

absence of, the introns between particular exons providing evidence of Rosaceae monophyly. Species of Maloi-

deae, with their higher chromosome number, have two copies or loci of each sequence, each of which have

differences in their base-pair sequences that are designated GBSSI-1A and IB, and GBSSI-2A and 2B. So there

are six different sequences or loci, the GBSSI-1 and -2 in the diploid non-maloids studied, and GBSSI-1A, -IB,

-2A, and -2B occurring the Maloideae. The sequence data, using only exon base pairs, showed that various

sampled collections of Kageneckia had GBSSI-1A, -2A and -2B loci; and Vauquelinia had GBSSI-1A, -IB, -2A and

-2B loci as in members of the Maloideae. Their results again showed that Kageneckia and Vauquelinia were

A later report by Evans and Campbell (2002) used CBSSI gene to investigate the origin of the x=17 Maloi-

deae. Their phylogenetic analysis of some 42 genera showed that GBSSI-1 and -2 alleles of Prunus (Amygdaloi-

deae) were not closely associated with the Maloideae, but rather the sequences from the genus Gillenia (a her-

baceous, x=9 Spiraeoid, with compound leaves and 5 separate ovaries each with 4-6 ovules that form folicular

fruits with non-winged seeds) were strongly associated with, and basal to, sequences of Vauquelinia, Lindleya

and Kageneckia at the base of the Maloideae clade. Gillenia has a GBSSI-1 locus that shares distinct intron dele-

tions and additions with the GBSSI-1B loci of the Maloideae and the GBSSI-2 intron shared a distinct base pair

substitution with the GBSSI-2B loci of Vauquelinia, Kageneckia and Lindleya and core Maloideae placing the

diploid Gillenia at the base of the Maloideae.

Thus Gillenia would appear to be an extant survivor of a lineage ancestral to the Maloids. But what is the

other parent? Their survey of the Amygdaloideae has found no potential parent. The other parent has either not

been sampled or is long extinct. They conclude that the other parent could have been another x=9 Spiraeoid

that, in forming a hybrid via amphiploidy, could double the chromosomes to x=18, (2n=36) and this could be

reduced to x=17 (2n=34) via aneuploidy (Evans & Campbell 2002). But whatever the other parent would be,

there is no reason to expect its lineage to be extant today. But definitely one of the parental lineages has a sur-

viving member, that being Gillenia, a genus of two species native to the eastern United States.

The most recent molecular study of relationships within Rosaceae was presented by Potter et al. (2007)

that investigated the relationships of 88 genera using nucleotide sequence data from six nuclear and four chlo-

roplast regions. Their paper resulted in a complete infrafamilial rearrangement recognizing three clades as

subfamilies. The basal subfamily Rosoideae consists of herbs, shrubs rarely trees, that lack cyanogenic glyco-

sides and sorbitol. They have alternate, usually compound, stipulate leaves; ovaries are usually numerous

(rarely 1), separate, free from the hypanthium, some borne in a spiral arrangement on expanded receptacles,

and the fruit are indehiscent [x=7(-8) — including Agrimonia, Filipendula , Fragaria, Geum, Potentilla, Rosa,

Rubus etc.]. The subfamily Dryadoideae clade consists of shrubs, subshrubs, with cyanogenic glycosides and

traces of sorbitol and a tendency to have nitrogen fixing symbionts. The leaves are simple or compound, stipu-

late; ovaries are 1 or many; fruits are achenes, (x = 9) including Cercocarpus, Chamaebatia, Dryas and Purshia

(including Cowania). The Dryadoideae is sister to a highly diverse Spiraeoideae 1 clade consisting of mostly

shrubs, small trees with some cyanogenic glycosides and strong sorbitol presence. Leaves are usually simple,

alternate, the stipules persistent (deciduous in Prunus); ovaries number 1-5, mostly separate, radially oriented,

usually free from the hypanthium, and fruits ranging from achenes, drupes, to pomes (x= 8, 9, 15, 17). It in-

cludes what was in the subfamilies Spiraeoideae ( Spiraea , Holodiscus, Petrophyton etc.), Amygdaloideae

(Prunus), and Maloideae (Amelanchier, Crataegus, Malus, Pyrus etc.) of previous classifications and several

Within the subfamily Spiraeoideae, the Maloid clade was designated in Potter et al. (2007) as the super-

tribe Pyrodae [the subfamily name Pyroideae Burnett (1835) named for Pyrus, the pear, has priority over name

Maloideae Weber (1964)]. Basal in the supertribe is the genus Gillenia (x=9), that Evans and Campbell (2002)

showed is a surviving genus of a lineage that gave rise to the Maloids. The supertribe has a single tribe, Pyreae,

containing the three genera Lindleya, Kageneckia (x=17), and Vauquelinia (n=15) — its chromosome number the

product of further aneuploidy and the tribe Pyreae has one subtribe, Pyrinae, that includes all the core maloids

with apple-like fruits (pomes). But I see a problem with this classification. There remains no rank to distin-

guish the variation in the core Pyrinae (the maloids) as investigated by Lo and Donoghue (2012). And that is

the subject of another paper.

Lindleya vs. Vauquelinia . — Both Lindleya and Vauquelinia (Hess & Henrickson 1987) are moderately

large, evergreen shrubs, distributed from northern, east-central to southern Mexico, with Lindleya ranging

from northern Oaxaca to western Coahuila and the three species of Vauquelinia occurring from central Oaxaca

to northern Baja California Norte, south-central Arizona and trans-Pecos Texas. Both genera occur in arid to

semiarid scrublands, often limited to more mesic niches on north-facing slopes and along drainages or on

rocky habitats where their roots can reach deeper moisture. Both have coriaceous leaves with well developed,

fibrous bundle-sheath extensions with leaves of Vauquelinia being much larger and usually

than those of Lindleya.

The smaller flowers of Vauquelinia are arranged in distinct well-branched compound corymbs (but see

Evans & Dickinson 1999, who consider the inflorescences to be determinate, alternately branched dichasia

with lateral pleiocasia similar to those found in some Crataegus, sensu Weberling 1989), not mostly solitary as

in Lindleya. As in Lindleya, their leathery hypanthia bear five, ± thick, persistent sepals, five, white, ovate pet-

als, (18-)20 stamens with tapered filaments, yellowish, introrse anthers and the sessile 5-carpelled ovary is

mostly free from the hypanthium and topped with five separate styles, each somewhat compressed distally

with broad stigmas. But unlike Lindleya, sepal margins of Vauquelinia do not contain the multicellular glands;

anthers and petals are much smaller; ovaries are villous; and carpels are connate only along the inner (adaxial

or ventral) margins — the outer (lateral and abaxial) margins are free (see Hess and Henrickson 1987). In Vau-

quelinia, each carpel has two basal-attached ovules (not apically attached as in Lindleya); the ovules, as in

Lindleya, have two integuments, and the upper portion of the ovule develops into a wing. In Vauquelinia, ma-

ture fruits are more ovoid to oblong-ovoid (not ovoid-globose) in shape, and the fruit body is distinctly five

lobed in cross section with five radial incisions, villous to strigose (not glabrous), but as in Lindleya, each carpel

Journal of the Botanical Research Institute of Texas 6(2)

is loculicidally dehiscent all across the ventral and distal portion of the dorsal sutures, splitting the persisting

style base to shed the winged seeds. The embryos in Vauquelinia are about half the total seed length, are ascend- j

ing, with basal hypocotyls (not two thirds the seed length with apical hypocotyls as in Lindleya). In both genera

the seed coat is thin, brown, and endosperm is lacking. They also differ in chromosome number, with Vau- j

quelinia n=15 and Lindleya n= 17.

The South American (Chile, Peru, Brazil) Kageneckia (n= 17) is quite distinct. The 3(-4) species are dioe- j

cious, ± large shrubs, vegetatively quite similar to some species of Vauquelinia in having thick, coarse leaves

and occurring in dry scrublands. They have flowers ± 20-40 mm in diameter with attenuate sepals, moder-

ately large, oblong-ovate white petals, 15-20 stamens along the hypanthial rim in male flowers but with re-

duced staminodia, and 5 separate vertical ovaries in female flowers, with terminal short styles and 2 rows of j

ovules (10-12 total) along the ventral traces. In fruit, the separate carpels expand abaxially and radiate out-

ward to enclose the seed wings and the fruiting ovaries open along both the ventral and dorsal sutures to dis-

perse the 10-12 winged seeds.

Spjut (1994) refers to the fruit of Vauquelinia as a coccetum “a multiple fruit with dehiscent fruitlets” j

with each carpel having a separate style-stigma. But the distinction between the fruits of Vauquelinia and Lind-

leya is only in the amount of lateral connation of the 5 carpels, being restricted to near the interior (ventral or

adaxial) region in Vauquelinia but throughout the lateral surfaces in Lindleya. In overall structure and function,

they are both loculicidal capsules. In contrast in Kageneckia. the carpels are completely separate, and Spjut

(1994) would designate the fruit as a follicetum (i.e., a cluster of follicles) but while dehiscence occurs primar-

ily along the dorsal suture, it continues onto the ventral suture, and his definition removes it from the follicle

category, making it fit Spjut’s definition of a coccetum as in Vauquelinia.

In molecular phytogenies that include Vauquelinia, Lindleya and Kageneckia [Potter et al. (2007); Camp-

bell et al. (2007); Lo & Donoghue (2012)], Lindleya and Kageneckia are most often associated and sister to

Vauquelinia and the rest of the pome-bearing Pyrinae. But also see Campbell 2007 for analysis of separate

GBSSI genes.

APPENDIX 1

rra de Pampas, W of Hacienda El Berrendo, 27°20’N, I04°43'W, 25 Aug 1972, Chians J

mmit Sierra de Chupaderos, 27°12’N, 104°43'W, 5300 ft, 2 Oct 1973, Henrickson 13776 J

(TEX). Coahuila: E slope Sierra Almargre, 4800 ft, 5 May 1973, Gentry 6- Engard 23219 (CAS, US); Vicinity of Santa Elena Mines, E foothills j

of Sierra de las Cruces, 30 May 1941, Stewart 378 (F, GH, LL); Sierra Mojada, S of Esmeralda, 27°16'N, 103°4rW, 1 Sep 1972, Chiang etal. j

9086n (LL); 23 (air) mi NW of Las Delicias, Valley N of Sierra de las Delicias, 26°23‘N, 102°52’W, 4800 ft, Henrickson 6124 (TEX); ± 29 (air) :

mi WNW of Cuatro Cienegas, N slope of Sierra de la Madera, 7.8 (rd) mi W of Rancho Cerro de la Madera, Cafidn Desiderio, 27°08'N,

102°30'W, 12 Aug 1976, Henrickson & Prigge 15310 (TEX); Cerro San Pedro, near N.L. line, 2 mi E of Ejido Presa de San Javier, 24°44’N,

100°46'W, 2200 m, 21 Aug 1974, Wendt & Lott 608 (TEX); Sierra dejimulco, 8 km NE Estacion Otto, 27 Sep 1972, Chiang et al. 9552h (LL); ?

Sien-a de Parras, 5500 ft, Shreve & Tinkham 9859 (GH); 10 mi W Saltillo, near Las Barrancas, 3 May 1959, Correll & Johnston 21400 (GH, NY, J

, 4 Sep 1938, Shreve 8581 (US); Concepcion del Oro, 2500-2700 m, 18-19 Jul |

atecas-Coahuila state line, 24°44'N, 101°10'W, 1990 m, 29 Mar 1973 Johnston

et al. 10491a (LL). San Luis PotosE 15.1 id. mi N of Zac-SLP, hwy 49 on rd to Charcas, near Cerro Tecalote, 22°30'N, 101°09'W, 2100 m, 23

Sep 1978, Henrickson & Lee 17553 (TEX); Lajoya, 4 km NW de Ventura, Mcpio Villa Hidalgo, 1900 m, 11 Jan 1955, Rzedowski 5705 (TEX); 20 |

ss & Byrne 4710 (F); E de Nunez, km 84

t, Rzedowski 5549 (ENCB). Nuevo Leon: Mts near Monterrey, Jul 1933; Mud- j

E of San Rafael, 25°03'N, 100°25 r W, 22 >

jn 1934, j

«,US);13kmalESan

22 Sep 1974,

ordetal 626 (DS, GH, NY); 34 km

80 km NE of Querttaro, above

1. 7425b (GH).

d to Pinal de A moles, 2700 m, 24 Apr

1, Pennell 17419 (GH, h

Henrickson, Systematic of Lindleya

2850 (DS, TEX); Cerro San Miguel, 14 km NNE de Actopan, 2300 m, 6 May 1965, C

lan, road to Zacualtipan, 1600-1800 m, 24 Mar 1947, Moore 2481 (GH). Puebla: Los Naranjos, May 1908, Purpus 3234 (F, GH, NY); Mcpio

Caltepec, Cerro El Mirador al SW de Coatepec, 19 Apr 1985, Tenorio8822 (TEX); 8kmalNE Acatepec, 16 May 1981, Chiongrtol. 1959 (TEX).

Oaxaca: 3 km al SW de Tamazulapan sobre carr. Tamazulapin-Chilapa de Diaz, 15 May 1982, Rico et al. 332 (F); Cerro sobre el camino de

Teposcolula a San Andres Lagunas, 10 May 1981, Cedillo et al. 770 (CAL, F); 3 km al S de Santiago Teotongo por la Terraceria a San Pedro

Nopala, 17°45'N, 97°33W, 12 May 1986, Salinas &■ Solis F3238 (TEX); 3 km SW de Magdalena Jicotlin a Santiago Teotongo, 97°29'N,

ACKNOWLEDGMENTS

I thank the curators for the herbaria mentioned for loans and Thomas Wendt for reading over an earlier version

of this paper, the reviewers of the article, which has been awaiting publication for more than a decade. Figures

3 and 5 were drawn by Bobbi Angell. Timothy A. Dickson (TRT) and Joseph R. Rohrer (UWEC) gave the

manuscript a critical review.

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ILLUSTRATIONS AND STUDIES IN NEOTROPICAL ORCHIDACEAE. 5.

THE LEPANTHES OVALIS GROUP (PLEUROTHALLIDINAE)

WITH THREE NEW SPECIES FROM COSTA RICA

1 Diego Bogarfn Christina M. Smith

Daniel Jimenez

Journal of the Botanical Research Institute of Texas 6(2)

Bogarin 2010); L. schizocardia Luer (Pupulin et al. 2010). Two other groups are under review: L. guatemalensis

Schltr. (Pupulin & Bogarin in prep.) and L. minutilabia Ames & C. Schweinf. (Smith et al. in prep.).

In this paper, we studied a group of species related to the Jamaican endemic Lepanthes ovalis (Sw.) Fawc.

& Rendle, the first species of Lepanthes to be described and which was originally placed in Epidendrum L.

(Swartz 1788). The other species of this group are L. erinacea Rchb.f., one of the first species described in Costa

Rica (Reichenbach 1855), L. pan Luer & Dalstrom from Ecuador (Luer 1996), L. seegeri Luer from Panama

(Luer 1987), L. trichocaulis Luer & R. Escobar (Luer 1988) and L. viridis Pupulin & Bogarin, recently published

from Costa Rica (Pupulin & Bogarin 2011). All members share similar plant morphology, having ramicauls

with long-ciliate sheaths with markedly dilated, long-ciliate ostia. The leaves are subcoriaceous, always light

green without purple stripes or spots. The congested racemes arise under or above the leaves and are shorter

than the leaves (rarely larger in some specimens of L. erinacea). The sepals are ovate, acute to obtuse, pubescent

or glabrous. The upper lobes of the petals are wider and longer than the lower lobes. The lip is triangular, cor-

date or subcordate, with the sides strongly folded and embracing the column at sides (rarely expanded), the

blades are flattened with cuneate connectives, the apex is cleft leaving a narrow space between the apices with

a pubescent appendix (Luer 1987; Luer 1996; Pupulin & Bogarin 2011). At the moment the group range in- ^

eludes Costa Rica, Panama, Colombia, Ecuador and Jamaica. We add three species proposed here as new to

science. A key to the species of the group is provided. Additional material is provided online at www.epiden-

E COSTA RICA. CArrAGO.Jin

Plant epiphytic, caespitose, pendent herb, up to 15 cm long. Roots slender, flexuous, up to 1 mm in diameter.

Ramicaul more or less descending, up to 10.5 cm, enclosed by 11-21 ciliate, lepanthiform sheaths, lightly cili-

ate especially on new growth; the ostia markedly dilated, ciliate. Leaves coriaceous, green, elliptic, subob- ;

ovate, acuminate with a short apiculus, 3.0-4.0 x 1.0-1.5 cm; cuneate base narrowing into a petiole ca. 3.5 mm

long. Inflorescence racemose, distichous, successively flowered, borne above the leaf, shorter than the leaves,

up to 3 cm, peduncle 1.5-2 cm long, rachis 0.5-1.0 cm. Floral bracts 1 mm long, ciliate. Pedicel 1.5 mm long,

persistent. Ovary up to 1 mm long. Flowers light orange, yellow, red-orange and scarlet; sepals light orange;

petals yellow with red-orange and scarlet; lip scarlet; column red. Dorsal sepal ovate, acute, connate to the

lateral sepal for about 1 mm, 3.2 x 2.9 mm. Lateral sepals ovate to elliptic, acute, connate for about 1 mm, 2.5

x 2.5 mm. Petals essentially glabrous, transversely bilobed, entire, 0.8 x 2.4 mm; the upper lobes broadly un-

cinate, rounded, overlapping, the apex broadly rounded; the lower lobes ovate, falcate, with apex broadly

rounded, slightly smaller than the upper lobes. Lip bilaminate, adnate to the column, 1.8 x 1.3 mm expanded;

blades narrowly oblong, pubescent, with narrowly rounded ends, falcate; connectives broadly cuneate, up to 1

mm long, perpendicular to the column and not leaving it exposed; body broadly oblong, connate to the base of

the column; appendix narrowly oblong, pubescent. Column cylindric, 1 mm long; anther apical; stigma ven-

tral. Pollinia two, ovoid. Anther cap cucullate.

Distribution.— Only known from Costa Rica.

Habitat and ecology. -Epiphyte in secondary premontane wet forest on the Atlantic watershed of Cordik

lera de Talamanca, between 1000 and 1200 m of elevation.

Bogarin et al.. New species of Lepanthes from Costa Rica

363

Drawn by CM. Smith from £ Pupulin 7477 (JBL-Spfrit).

368

Journal of the Botanical Research Institute of Texas 6(2)

i. E) Up, spread. F) Pollinarium

Bogarin et al„ New species of Lepanthes from Costa Rica

apex. Dorsal sepal broadly ovate, obtuse or subacute, connate to the lateral sepal for about 1.3 mm, 3.3 mm x

2.8 mm. Lateral sepals ovate to elliptic, obtuse, connate for about 1.4 mm, 2.9 mm x 2.7 mm. Petals minutely

pubescent, transversely bilobed, entire, 1.2 x 3.2 mm; the upper lobes oblong-obovate, truncate, not overlap-

ping, the apex broadly rounded; the lower lobes oblong, obtuse, subfalcate, the apex broadly rounded, smaller

than the upper lobes, lip bilobate, spreading, adnate to the column, 1.2 mm x 1.5 mm, expanded; the blades

ovate, with rounded ends, falcate; the connectives cuneate, up to 0.5 mm long, perpendicular to the column

leaving the appenndix exposed; the body oblong, connate to the base of the column; the appendix oblong,

pubescent. Column cylindric, 1 mm long; the anther apical; the stigma ventral. Pollinia two, ovoid. Anther

Distribution. — Only known from the southern Pacific region of Costa Rica.

Habitat and ecology. — Epiphytic in tropical wet forest, premontane belt transition at 800 m of elevation

along the Pacific watershed of Cordillera de Talamanca in southern Costa Rica.

Etymology. — From the Latin expansus “spread” and labium “lip,” in reference to the spreading lip with the

lobes not touching each other, leaving the appendix exposed.

Phenology. — Plants have been recorded in flower in April.

Discussion. — It is similar to L. erinacea but differs mainly by the pendent plants (vs. erect) with ramicauls

with sparsely ciliate sheaths (vs. heavily ciliate), the petals with the lobes oblong, subsimilar (vs. the upper

flabellate, the lower minute, oblique), the lower lobe yellow (vs. red), the lip spreading with the lobes separated

and not touching each other, leaving the appendix exposed (vs. the apices of the blades approximate, hiding

the appendix) (Fig.l). Other differences are summarized in Table 1.

k Lepanthes ovalis (Sw.) Fawc. & Rendle, FI. Jamaica 1:71. 1910 . 1

As noted by Pupulin (2009), there is no way to assume that any of the specimens at BM is the holotype. The

species should be lectotypified after a more detailed study of the available material; drawing of type, W). For a

complete description and illustration see Luer (2009).

Distribution. — Only known from Jamaica.

Habitat and ecology. — Epiphytic in cloud forest along the Blue Mountains.

Discussion. — Plants are recognized by the erect to suberect arching habit, the inflorescences developed

above the leaf, the upper lobe of petals suborbicular, rounded, yellow with red-purple base, the lower lobe yel-

low, pubescent and the inconspicuous appendix.

5. Lepanthes pan Luer & Dalstrom, Monogr. Syst. Bot. Mis

UADOR. Imbabura: Los Cedros Reserve, wet forest near Rfo Los Cedi

Ingram-FerrcU 2200 (holotype: MO). For a complete description and

i Bot. Gard. 61:133-134, f. 178. 1996. Type ec-

1200-1300 m, 24 Mar 1996, 5. Dal strOm, S. Ingram & K.

■ation see Luer (1996).

Distribution. — Only known from Ecuador.

Habitat and ecology. — Epiphytic in wet forest at around 1200-1300 m of elevation.

Discussion. — It is distinguished by the erect plants with inflorescences developed behind the leaf, the

lateral sepals acute, pubescent, the upper lobe of petals with a lobule at the inner apical comer, the lower lobe

long-ciliate, pubescent, the apex of the lip is shallowly cleft with apiculate apices in apposition with a small

triangular appendix.

6. Lepanthes seegeri Luer, Orchidee (Hamburg) 38:60. 1987. Type: PANAMA. CHnuqut: epiphytic in forest above Boquete,

alt. 1600-1700 m, Apr 1985, collected by H. Seeger 678A, flowered in cultivation at Heidelberg University 16 Oct. 1985, C. Luer

11424 (holotype: MO). For a complete description and illustration see Luer (1987).

Distribution. — Only known from Panama.

370

Journal of the Botanical Research Institute of Texas 6(2)

Habitat and ecology.— Epiphytic in cloud forest at around 1600-1700 m of elevation along the Cordillera

de Talamanca in western Panama.

Discussion .— It is recognized by the erect plants with inflorescences developed behind the leaf, the lateral

sepals obtuse, glabrous the upper lobe of petals with a lobule at the inner apical corner, the lower lobe long-

ciliate, pubescent and the lip with a long, slender, descending, pubescent appendix that protrudes from a shal-

nsis Bogarin & D. Jimenez, sp. nov. (Figs. ID, 5). Type: COSTARICA. SanJose.- Dota, Sa

i and Naranjillo de Tarrazu, 9°35'33.35"N 83°58'26.74"W, 1615 m, epiphytic, legit Daniel JimEnez, f

Plant epiphytic, caespitose, erect herb, up to 14 cm tall. Roots slender, flexuous, up to 1 mm in diameter.

Ramicauls erect to suberect, 3.5-9.0 cm long, enclosed by 7-10 ciliate, lepanthiform sheaths, ciliate especially ;

on new growth; ostia markedly dilated, ciliate. Leaves coriaceous, green, elliptic to oblong, acute to acuminate

Inflorescence racemose, distichous, successively flowered, beneath (rarely above) the leaf, shorter than the

leaves, up to 2.5 cm long; peduncle 1.3 cm long; rachis 1.2 cm. Floral bracts 1 mm long, ciliate. Pedicel 2 mm

long, persistent. Ovary up to 1 mm long. Flowers sepals light orange, petals and lip with the bases scarlet-red

and the apices yellowish-orange, the column pinkish-purplish. Dorsal sepal broadly ovate, acute, glabrous,

connate to the lateral sepals for about 1.2 mm, 3.7 x 3.6 mm. Lateral sepals broadly ovate, acute, glabrous, con- \

nate for about 1 mm, 3.9 x 2.5 mm. Petals transversely bilobed, 1.8 x 4.2 mm, slightly ciliate along the margins; 1

upper lobes oblong-trapeziform, overlapping, apex broadly rounded or truncate; lower lobes smaller than the

upper lobes, oblong to narrowly triangular, oblique, apex rounded. Up bilobate, adnate to the column, 1.5 x

2.2 mm expanded; blades ovate, glabrous with ciliate, rounded apices, subfalcate; connectives cuneate, up toi

0.7 mm long, perpendicular to the column and not leaving it exposed; body oblong, connate to the base of the

column; the appendix oblong, shortly pubescent. Column cylindric, 1.2 cm long; anther apical; stigma subapH

Distribution .— Only known from the central-southern Pacific region of Costa Rica.

Habitat and ecology.— Epiphytic in premontane and lower montane rain forest along the Pacific watershed

of Cordillera de Talamanca.

Etymology.— From the locality of Tarrazu in San Jose province. Tarrazu comes from an indigenous Hu-

etar word of unknown meaning.

Phenology.— Plants were recorded in flower from February to March.

Discussion. It is similar to L. seegeri but differs mainly by the glabrous petals and the lip glabrous with

ciliate apices (vs. long ciliate-pubescent), the upper lobe of petals truncate, without lobules at the inner apical

corner (vs. oblong, rounded, with lobule at the inner apical comer), the petals and lip basally red, tinged with

yellow-orange at apex (vs. completely red without stains), the blades of the lip touching each other above the

column (vs. slightly expanded and not touching each other), the appendix shorter, a small triangular lobule,

not extending far from the apex of the column (vs. appendix longer, descending, extending far from the apex

of the column) (Fig.l). Other differences are summarized in Table 1.

8. Lepanthes trichocaulis Luer & R. Escobar, Orquideologia 17:224-226. ]

& J. Posada at Colomborquldeas 26 Mar 1984, R. Escobar 3245 (holotyfe: MO). For a cc

1. Type: COLOMBIA. A

Distribution . — Only known from Colombia.

Habitat and ecology.— Epiphytic around 1820 m of elevation (Luer 1988).

Discussion.— It is recognized by the erect plants, the inflorescences developed behind the leaf, the upper j

372

Journal of the Botanical Research Institute of Texas 6(2)

lobe of petals oblong, rounded and not overlapping each other, the lower lobe longer or as long as the blades of

the lip a couple of longer hairs in place of an appendix.

9 Lepanthes viridis Pupulin & Bogarin, Lindleyana in Orchids, Mag. Amer. Orch. 5

COSTA RICA. Turk, alba: Tuis, Cien Manzanas, 9°51'25"N 83=3347 AN ca. 3,1 17 ft (950 m), along a mi.

premontane wet forest, secondary vegetation and remnants of primary, 26 May 2003, F. Pupulin 4801,

G. Gerlach, P Kindlmann, H. Ledn-Patz, S. Pugh -Jones & E. Serrano (holxjtype: JBL). For a complete des

Distribution . — Only known from Costa Rica.

Habitat and ecology.— Epiphytic in premontane wet forest on the Caribbean slopes of Cordillera de Tala-

Discussion.— It is superficially similar to L. pan, from which it mainly differs by the distinctly obovate

leaves (vs. orbicular and shortly acuminate), the yellow flowers with bright green petals (vs. peach and red), the

lateral sepals shorter than the dorsal sepal (vs. equal in length), the densely villose petals (vs. upper lobe pubes-

cent, lower lobe pubescent on the outer half), with the upper lobe rounded (vs. truncate), and the apices of the

lateral lobes of the lip acute (vs. obtuse).

KEY TO THE SPECIES OF THE LEPANTHES OVALIS GROUP

;rthan the lobes of the lip, apex orange-yellow, lip red

L. dikoensis Bogarin & C.M. Sm. (Costa Rica)

is Pupulin & Bogarin (Costa Rica) "

ACKNOWLEDGMENTS

We acknowledge the scientific services of Ministerio del Ambiente, Energia y Telecomunicaciones de Costa |

Rica (MINAET) and Sistema Nacional de Areas de Conservation (SINAC) for issuing the Scientific Passports

under which wild specimens treated in this study were collected. The curators and staff at CR, 1NB and US|

herbaria for granting access to the collections and for the facilities provided. Franco Pupulin, Jose F. Morales,

Lisa Thoerle, and one anonymous reviewer, provided useful discussions and comments that improved the

manuscript. We thank Adam Karremans for helping with fieldwork and literature. William Chacon of

Orquideario Happy Garden, Coto Brus, Puntarenas kindly provided plant material and data. The present pa-

per is part of the Project 814-A0-052, “Flora Costaricensis: taxonomia y filogenia de la subtribu Pleurothallidi-

nae (Orchidaceae) en Costa Rica,” sponsored by the Vice-Presidency of Research, University of Costa Rica. J

Bogarin et al.. New species of Lepanthes from Costa Rica

373

REFERENCES

Barringer, K. 1 986. Typification of Schlechter's Costa Rican Orchidaceae. I. Types collected by A Brenes. Fieldiana, Bot.

Bogarin, D. and M. Fernandez. 2010. Lepanthes arenasiana (Pleurothallidinae: Orchidaceae), a new species from Costa Rica.

Lankesteriana 9:487-489.

Luer, C.A. 1987. Vier neue Arten im Subtribus Pleurothallidinae: Lepanthes hubeinii, seegeri, rauhii und Platystele rauhii.

Orchidee 38:58-62.

Luer, CA 1988. Species of the genus Lepanthes from Colombia. Series 2-3. Orquideologla 1 7:145-230.

Luer, C.A. 1992. New species in Lepanthes Sw. (Orchidaceae). Lindleyana 7:100-1 18.

Luer, CA 1996. leones Pleurothallidinarum XIV. Systematics of Draconanthes, Lepanthes subgenus Marsipanthes, and

subgenus Lepanthes of Ecuador (Orchidaceae). Part Three: The genus Lepanthes subgenus Lepanthes in Ecuador.

Monogr. Syst. Bot. Missouri Bot. Gard. 61 :1-255.

Luer, C.A. 2009. leones Pleurothallidinarum XXX. Lepanthes of Jamaica: Systematics of Stelis: Stelis of Ecuador. Part four

Addenda, systematics of Masdevallia: new species of Lepanthes from Ecuador, and miscellaneous new combinations.

Monogr. Syst. Bot. Missouri Bot. Gard. 1 15:1-265.

Pupuun, F. 2009. Pupulin, F.Typi Swartziani Orchidacearum Indiae Occidentalis in Herbario Vindobonense conservandi.

Ann. Naturhist. Mus. Wien 1 10B:21 3-247.

Pupuun, F. 2010. Orchidaceae werckleanae: typification of Costa Rican orchid species described from collections by K.

Werckle. Bot. J. Linnean Soc. 163:111-154.

Pupuun, F. and D. Bogarin. 2010. Illustrations and studies in Neotropical Orchidaceae— The Lepanthes jimenezii group

(Pleurothallidinae). Harvard Pap. Bot. 15:11 1-1 21.

Pupuun, F. and D. Bogarin. 2011. Two new Lepanthes from Costa Rica. Lindleyana in Orchids, Mag. Amer. Orch. Soc.

80(3):1 78—1 81 .

Pupuun, F., D. Bogarin, and C.M. Smto. 2010. Two new species of Lepanthes from Costa Rica close to L schizocardia (Orchi-

daceae: Pleurothallidinae). Lankesteriana 9:423-430.

Reichenbach, H.G. 1855. Symbolae Orchidaceae. Bonplandia (Hanover) 3(15-16)212-227.

Reichenbach, H.G. 1 858. Lepanthes Sw. Xenia Orch. 1:151—1 52, 1 54. Tab. 49.

Schlechter, R. 1923. Beitrage zur Orchideenkunde von Zentralamerika, II. Additamenta ad Orchideologiam Costaricen-

sem. Repert. Sp. Nov. Reg. Veg. Beih. 19:3-307.

Swartz, 0. 1788. Nova genera et species plantarum; seu, Prodromus descriptionum vegetabilium, maximam partem in-

cognitorum quae sub itinere in Indiam Occidentalem annis 1 783-87 / digessit, Olof Swartz, M.D. - Stockholm, Upsala

and Aboa: Bibliopolid Acad. M. Swederi.

374

BOOK REVIEWS

Christian Ziegler. Introduction by Michael Pollan. 2011. Deceptive Beauties: The World of Wild Orchids.

(ISBN-13: 978-0-226-98297-7, hbk.). The University of Chicago Press, Chicago, Illinois 60637, U.S.Af |

(Orders: www.sinauer.com). $45.00, 183 pp., color photos throughout, 9 3 A" x 9 3 /4".

The book jacket describes the author as “a biologist turned photographer specializing in tropical natural his- ]

tory .” That is obviously a true statement— but it doesn’t go far enough. This is a person who greatly enjoys what I

he’s doing and it is clearly reflected in the beauty, the layout, and the genuine “feel” — and love — of orchids. |

Michael Pollan’s introduction, “Sex Among the Orchids,” definitely catches the reader’s attention with his

opening statement: “We animals don’t give plants nearly enough credit. “ His following discourse is not only |

fascinating and treated with fact, history, and humor, the reader is easily fully engaged — and can’t put the book

The photography is outstanding. Ziegler’s accompanying chapters are well written, explanatory, and pro-

vide exceptional background, diversity, descriptions of various habitats, atmospheric changes, uses, concerns,

biological changes that influence the habitat environments, and, of course, effects of human behavior as well—

Expect to take time to browse and enjoy this volume. It is really well done, and it makes one very aware of

potential challenges in the upcoming years. — Helen Jeude, Volunteer and Assistant Editor, Botanical Research

Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, U.SA.

Carol Grace with a foreword by Eric Lamont. 2012. Spring Wildflowers of the Northeast: A Natural History.

2012. (ISBN: 978-0-691-14466-5, hbk.). Princeton University Press, 41 William Street, Princeton, New

Jersey 08540, U.S.A. (Orders: http://press.princeton.edu/titles/9668.html, 609-258-4884, 609-258-

6305 fax). $29.95, 290 pp., 512 color illus., T/i” x 10".

What a welcome addition to wildflower literature! This is the ideal book for readers of all levels of expertise,

from the professional botanist to the armchair naturalist. Princeton University Press affirms the value of Carol

Gracie’s passion and attention to detail, her botanical research and travel, her years of teaching, her insightful

writing, and 500 revealing photographs. The book first appears to be a gorgeous coffee table book, but then, I

opened those first pages, and found myself dropping down the rabbit hole into Carol Gracie’s extraordinary

botanical world rarely experienced by most people. Her teaching years at the New York Botanical Gardens, her

intelligence and curiosity come alive to show the identification and natural history of a select group of wood-

land wildflowers from baneberries to violets. She thinks of the question before you ask it. She touches on com-

plicated topics of climate change, ethnobotany, horticulture, medicine and more, without being tedious. In

addition, she refers to Indian lore, botanical literature and art. Carol Grade explains the reason for taxonomic

name changes and adds an impressive list of references, which is testament to the extensive research to prepare

this work. I have heard that some times when we look, we don’t see. Carol Gracie teaches us to see. I salute this

gifted plantswoman for focusing on the beauty and intricacies of northeastern wildflowers, adding a valuable

contribution to our knowledge of the flora of the United States.— Jane Austin Bruckner, graduate of Rutgers Uni-

versity, New Jersey, a Registered Nurse, a Certified Master Gardener and BRIT Volunteer, 1700 University Dr., Fort

Worth Texas 76107-3400, USA.

CALATHEA BASIFWRA (MARANTACEAE), ANEW SPECIES ENDEMIC TO PANAMA

Helen Kennedy

Department of Botany and Plant Science

University of California Riverside

Riverside, California 92521, U.S.A

ganders@mail.ubc.ca

RESUMEN

In the Flora of Panama (Woodson & Schery 1945), a total of 23 species of Marantaceae were listed with 14 in

the genus Calathea. By 1972, Dressier (1972: 184) reported 35 species. Later, Kennedy (1976: 312-313) noted a

total of 49 species and predicted an eventural increase to 60 or 70 because of the species richness found along

the Rio Guanche(Col6nProv.) and the El Llano-Carti road (Panama Prov. and San Bias Prov.) and general inac-

cessibility of those areas at the time. More recently, Kennedy (2012: 49) reported 63 species of Marantaceae

from Panama. Currently 67 species (ca. 191% increase from the 1945 Flora of Panama treatment) are recog-

nized with 49 in the genus Calathea. Eighteen species are recognized as endemic, including the one described

herein plus two as yet undescribed taxa. Additional collecting in Costa Rica, but especially in adjacent Colom-

bia, will reduce this number.

TAXONOMIC TREATMENT

Calathea basiflora H. Kenn., sp. nov. (Fig. 1) Type: PANAMA. San Bias. Rio PlayOn Chico, Campamento Nebba Dummat, Filo

del Sureste 200 120 m 09°14'N 78°15'W, lOJul 1994, H. Herrera 1661 (holotype: PMA; isotypes: MO 6315278, PMA 75534).

Calathea basiflora differs from C vmcunda in the wider leaf blades, 17.5-21.5 vs. 4-8 cm, the obtuse vs. acute to acuminate bract apex; from

both C. verecunda and C. cleisthantha by the presence of 1-2 claviculate bracteoles vs. all bracteoles membranous; and from C. rhizanthouks

by the larger leaf blades, 42-46 x 17.5-21.5 vs. 17-30 x 9.1-16.5 cm. two inflorescences per shoot vs. a single inflorescence, and the ratto of

peduncle length to inflorescence length less than 1.2 vs. 3 or more.

Plants rhizomatous, perennial, herbs, 40-60 cm; cataphylls herbaceous, narrowly ovate, minutely pilose,

hairs ca. 0.2 mm, 3.5-5.5 on shoots bearing an inflorescence. Leaves all basal, 1-2 per shoot; leaf sheath not

auriculate, appressed pilose, hairs more dense along margins of wings and toward base, base and intemode

below, sericeous, ca. 40 cm; petiole with a distinct groove adaxially, scattered minutely pilose, hairs ca. 0.1 mm,

2.5-3 cm; pulvinus minutely tomentose adaxially, sides and abaxial portion glabrous, 1-1.5 cm; leaf blade soft,

J78 Journal of the Botanical Research Institute of Texas 6(2)

Etymology . — The specific epithet, basiflora, refers to the inflorescences borne directly from the rhizome at

the base of the plant, well below the leaves.

ACKNOWLEDGMENTS

The Missouri Botanical Garden provided support for my accommodations while working in the MO herbari-

um (organized, thanks to Olga Martha Montiel). The travel expenses for the trip to MO were provided by Fred

Ganders. I am very grateful to Gerrit Davidse, Jim Solomon and Mary Merello for their help in the MO her-

barium, to Barry Hammel, Isabel Perez and Mireya Correa for help with the Spanish resumen and to Heraclio

Herrera for collecting this species. I thank Mireya Correa and an anonymous reviewer for helpful suggestions

and corrections. The type scan was provided by the Missouri Botanical Garden.

Dressler, R.L 1972. Terrestrial plants of Panama. Bull. Biol. Soc. Wash. 2:179-186.

Kennedy, H. 1976. Notes on Central American Marantaceae II. New species from Panama and Costa Rica. Bot. Not

128312-322.

Kennedy, H. 2012. Calathea rhizanthoides and C. peregrina (Marantaceae), new species endemic to Panama. J. Bot. Res.

Inst. Texas 6:49-54.

Woodson, R.E. Jr. and R.W. Shery. 1 945. Marantaceae. In: Flora of Panama. Ann. Missouri Bot. Gard. 32:81 -1 05.

TWO NEW DISTICHOUS-BRACTED CALATHEA (MARANTACEAE)

SPECIES FROM CENTRAL AMERICA

Journal of the Botanical Research Institute of Texas 6(2)

2.2-2.36:1 in cauline leaves, ca. 17 lateral veins per 3 cm and 15-19 cross-veinlets per 5 mm (both measured at

midpoint of sides of blade), adaxial surface green, glabrous, midrib minutely pubescent in basal half, hairs 0.1

mm, abaxial surface nearly glabrous, minute hairs to 0.2 mm along some lateral veins close to midrib, midrib

below minutely tomentose along sides, hairs 0.1-0.2 mm. Inflorescences 1-2 per shoot, first one terminal,

peduncle glabrous, 22-36 cm. Bracts 14-16, herbaceous, distichous, overlapping at base when live, rachis usu-

ally not visible, broadly ovate in basalmost, transverse broadly elliptic in subsequent ones to broadly elliptic in

uppermost, apex rounded with acumen in basalmost one, rounded to broadly obtuse in upper ones, margin

straight (not recurved), 4-5.3 x 3.6-5.3 cm, each bract subtending 3 or more flower pairs, abaxial surface of

bracts green, glabrous, adaxial surface glabrous; bicarinate prophyll membranous, rectangular-elliptic, apex

rounded to subtruncate, glabrous, 3-3.2 x 1.0-1.5 cm, 0.9-1. 1 cm wide, carina to carina; secondary bract mem-

branous, elliptic, apex rounded to minutely tridentate, glabrous, 2.6-3 x (0.85-) 1.25-1.5 cm; bracteole 1 per

flower pair, medial, membranous, 1.1-1.35 x 0.15-0.2 cm. Flowers open spontaneously, pale yellow (fide Croat

16835). Sepals membranous, narrowly obovate, obtuse, apical margin incurved, glabrous, 18-20 x 3-4 mm.

Corolla tube glabrous, 26-32 mm; corolla lobes subequal, elliptic, obtuse to 90°, glabrous, 12-14 x 3.5-5 mm.

Staminodes 3; outer staminode spathulate, 9-10 x 6-8 mm; callose staminode rounded apically, ca. 11.5 mm;

cucullate staminode 6-8 mm; stamen with lateral, petaloid appendage, 1 mm wide, anther 2.5 mm; ovary

smooth, glabrous, ca. 2.5 x 1.5 mm. Capsule unknown.

Distribution and habitat. — Calathea ravenii is known only from the type collection near Puerto Obaldia in

wet tropical forest habitat. Considering the proximity of Puerto Obaldia to the border with Colombia, it is

highly likely it will be eventually found there as well. It is apparently uncommon, as in two days collecting in

July, 2005, in the vicinity of Puerto Olbalda, we failed to find any plants of it, even sterile ones, though another,

previously unknown, undescribed species was found.

Discussion . — Calathea ravenii belongs to Calathea G. Meyer section Calathea, having the characteristic

habit of several basal leaves and a cauline leaf subtending the 1-several distichous-bracted inflorescences. It is

distinguished from other Panamanian species of Calathea with distichous, complanate, bracts by the elliptic

leaf blades, length 2.2-2.36 x width, the greenish, glabrous bracts with non-recurved apical margins, 4-5.3 x

3.6-53 cm, the single membranous bracteole per flower pair, and pale yellow flowers. A comparison of five of

the distichous-bracted Panamanian species is given in Table 1 (Kennedy 2011: 202). It is readily distinguished

from the other three distichous-bracted species not mentioned in the table, C. lasiostachya, C. caesariata and C.

similis H. Kenn. by its glabrous, green bracts and the glabrous adaxial surface of the leaf blade.

Etymology .— The specific epithet ravenii, is in honor of Dr. Peter Raven, director emeritus of the Missouri

Botanical Garden. It is with deep gratitude that I have the privilege of naming a species in his honor as he was

instrumental in my choosing the Marantaceae initially as an OTS field project and encouraging me to continue

this for my PhD research, besides indirectly introducing me to my partner. I am glad to finally be able to thus

acknowledge the debt of gratitude I owe for his help throughout my career, and that of numerous other botan-

ists, in this mai

Plants perennial, rhizomatous, caulescent, herbs, 2 or more m high. Leaves several, basal, and 1 cauline above

an elongate stem intemode; leaf sheath not auriculate, green, appressed tomentose, hairs to 1 mm, very dense-

ly tomentose to sericeous at very base in cauline leaves, 16-25 cm in cauline leaf; petiole green, appressed to-

mentose in basal %, subglabrous to glabrous apically 47-60 cm in cauline leaf; pulvinus glabrous except for

row of minutely tomentose hairs adaxially, hairs ca. 0.2 mm, glabrous near junction with petiole, 5-7 cm in

382

Journal of the Botanical Research Institute of Texas 6(2)

Fig. 2. Calatheaoscariana H. Kenn. Holotype (A Vetiz&M. Veliz 94.3926, BIGU). I

384 Journal of the Botanical Research Institute of Texas 6(2)

oscariana (Vdiz 99.7115, MEXU) and C. sclerobracteata ( Vdiz 99.7115B, BIGU), growing together under cultiva-

tion, are the notes: “Planta cultivada dendro de cafetales o areas con banano, sus hojas son empleadas para

envolver la masa de los tamales.” Likewise, such cultivation is noted on the type specimen, “Planta cultivada

utilizada para la elaboration de tamales (Envoltorio)” and similar on Vdiz & Vdiz 94.3983B (BIGU). Interest-

ingly, such use is not restricted to the campo, as I was served a tamale wrapped in the leaves of C. oscariana at

the buffet in my hotel in Guatemala City. The portion of leaf was saved and pressed for later comparison.

Etymology. — The specific epithet, oscariana, was recommended by Mario Veliz (BIGU), who collected the

type and provided photos and scans of the new species, as a dedication to his friend Senor Oscar Archila Euler

(+), “ la persona que la colectb y la tiene actualmente cultivada en su casa. . . .”

ACKNOWLEDGMENTS

I am especially grateful to Andrew Sanders of the UCR herbarium for allowing me the space to store and study

all the Mesoamericana loan material and for taxonomic discussions. I thank the following for help and use of

the herbarium facilities: Gerrit and Jeany Davidse (MO), Mario Veliz and Jorge Vargas (BIGU), Marie S. Cerma-

kova (USCG), Mario Sousa, Gloria Andrade and Gerardo Salazar (MEXU), Christine Niezgoda (F) and Car-

men Galdames (SCZ). I gratefully acknowledge the considerable help of Mario Veliz in providing color photos

of the inflorescences of both C. oscariana and C. sclerobractea as well as photographs of specimens for com-

parison. I am most grateful to Tom Croat for making the one and only (known) collection of C. ravenii. Teresa

Salvato provided accommodations and transport for my work at UCR. I am deeply indebted to Fred Ganders for

personally funding the cost of the herbarium visits and to the Missouri Botanical Garden for providing support

for my accommodations while working in the MO herbarium (organized, thanks to Olga Martha Montiel). I

thank Barry Hammel and an anonymous reviewer for help with the Spanish resumen and Gordon McPherson

and the anonymous reviewer for their helpful suggestions and corrections. Thanks to the curators of F, MEXU,

MO, PMA, and SCZ for the loan of their specimens. The type scan of C. ravenii was provided by the Missouri

Botanical Garden and the scan of C. oscariana, by Mario Veliz (BIGU).

REFERENCES

Kennedy, H. 201 1 . Three new distichous-bracted species of Calathea (Marantaceae) from Panama. Novon 21 :201 -21 1 .

Standley, P.C. and JA. Steyermark. 1952. Marantaceae. In: P.C. Standley and J.A. Steyermark, eds. Flora of Guatemala. Field

Mus. Nat. Hist. Bot. Ser. 24:207-221.

COLUMNEA ANTENNIFERA , A NEW SPECIES OF GESNERIACEAE

FROM THE CORDILLERA CENTRAL OF THE COLOMBIAN ANDES

Department of Biological Sciences

Box 870345

The University of Alabama

Tuscaloosa, Alabama 35487, U.S.A.

Laura Clavijo

Box 870345

The University of Alabama

Tuscaloosa, Alabama 35487, U.S.

lauriclav@gmail.com

Key Words: Columnea, Colombia, Episcieae, G

INTRODUCTION

The genus Columnea L. is primarily epiphytic and belongs to the New World subfamily Gesnerioideae and tribe

Episcieae. Columnea ranges from Mexico south to Bolivia and is most diverse in the northern Andes of Colom-

bia and Ecuador. With over 200 species, Columnea is the largest genus in the subfamily Gesnerioideae (Burtt &

Wiehler 1995; Weber 2004; Skog & Boggan 2006). The genus is distinguished from other closely related genera

by an indehiscent berry instead of a fleshy bivalved capsule.

Columnea antennifera J.L. Clark & Clavijo was discovered during a 2012 research expedition to the Co-

lombian department of Antioquia in the Cordillera Central of the northern Andes. A remarkable character of

Columnea antennifera is the presence of five elongate appendages near the corolla sinuses. The presence of co-

rolla appendages and where they appear has been discussed in numerous artificial classifications of groups

now recognized as Columnea, such as the section Ortholoma Benth. and the genus Trichantha Hook. (Morton

1963, 1971; Morley 1976; Smith 1994). Corolla appendages have not been thoroughly evaluated in a phyloge-

netic context and most likely this character is convergent within Columnea.

Columnea antennifera J.L. Clark & Clavijo, sp. nov. (Fig. 1) Tyre: COLOMBIA. Antioquia: Municipio Valdivia, Cordillera

Central, road Ventanas to Briceilo, before the quebrada El Oro, 07°05'20*N, 75°29’20"W, 1802 m, 19 May 2012 01), JX. Clark, J.

Anderson, L. Clavijo, M. Mazo & D. Suescun 13036 (holotype: COL; isoiytcs: BRIT, HUA, MO, NY, UNA, US).

Obligate scandent epiphytic climber; stems elongate and horizontal, 2-3 m long, suffrutescent, glabrescent

below, sparsely pilose above. Leaves opposite, strongly anisophyllous in a pair; larger leaf with petioles terete,

4-10 mm long, blade coriaceous when dry, elliptic to oblong, 3-12 x 1.3-3.4 cm, base rounded to oblique.

387

sometimes asymmetrical, apex acute, margin entire, adaxially shiny green, abaxially light green, sparsely pi-

lose on upper surface and densely pilose on lower surface; smaller leaf greatly reduced relal

nearly sessile, orbicular to ovate, 1-2.7 x 0.5-1.5 cm, base rounded to cordate, apex acute, m

faces and vestiture similar to larger leaf. Flowers solitary and erect; pedicels 1.3-2 cm long, red, pilose; calyx

2-3.5 cm long, uniformly bright red, inside and outside sparsely pilose, inside lanate at the base, lobes 5, erect

at anthesis, each lobe tightly appressed to adjacent lobe and folded lengthwise, ovate, apex broadly acuminate

margin entire, 4 lobes nearly equal 0.7-1 x 1.2-1.5 cm, fused at the base for 1.5-2.6 cm, 5 th lobe (dorsal) slight-

ly smaller, fused at the base for 1.1-1.5 cm; corolla tubular, posture erect in calyx, 3.0-4.5 x 0.6-1.0 cm; outside

uniformly bright red and tomentose, internally glabrate, lobes bright yellow, appressed, 5-6 x 2-2.5

ovate; corolla appendages present in immature and mature flowers, located in each sinus alt*

lobes, 14-20 mm long, pilose, bright yellow with or without a dark spot at the base; stamens 4, didynamous

included; filaments 12-15 mm long, coiled after anthesis, connate and adnate to the base of the corolla tube,

glabrous; anthers connate, longer than broad, 2-2.5 x 1-1.5 mm, dehiscing by longitudinal slits; staminode not

seen; nectary a dorsal gland, glabrous; ovary superior, lanate, ca. 5 x 4 mm, style ca. 30 mm long, glabrous,

stigma included and capitate. Fruits not seen.

Columned antennifera is morphologically similar to C. dissimilis C.V. Morton (Fig. 2, A-D). These two spe-

cies are easily differentiated by the elongate corolla appendages in Columned antennifera (Fig. 1) in contrast to

the relatively short corolla appendages in C. dissimilis (Fig. 2A). The corolla appendages are developed in im-

mature flowers of Columned dntenniferd (Fig. 1C). In contrast, the corolla appendages in Columned dissimilis are

either absent or significantly reduced when the flowers are immature (Fig. 2B). Another species that has co-

rolla appendages and is morphologically similar to C. dntenniferd is C.filomentosd (Figs. 2E, F). These two spe-

cies are readily differentiated by the uniformly red corolla in Columnedfildmentosd (Fig. 2E) in contrast to the

red corolla with bright yellow lobes in C. dntenniferd (Fig. 1). Vegetatively these two species are differentiated

by the isophyllous leaf arrangement in Columnedfildmentosd in contrast to the strongly anisophyllous C. dnten-

niferd. A third species that has corolla appendages is Columned rosed (C.V. Morton) C.V. Morton (Fig. 3E). How-

ever, the calyx lobes of Columned rosed are deeply serrate to fimbriate (Fig. 3E) in contrast to the entire calyx

margins of C. dntenniferd (Fig. 1).

Distribution and habitat. — Columned antennifera is known from the northern Cordillera Central of the

Colombian Andes in the department of Antioquia from montane forests (1800 m). Herbarium collections of

Columned antennifera were not seen during recent visits to the National University of Colombia (COL) or the

University of Antioquia (HUA), but an additional population was observed and photographed near the type

locality between Yarumal and Ventanas during a 1996 field expedition by Gunter Gerlach from the Munich

Botanical Garden (Botanischer Garten Munchen-Nymphenburg).

Etymology. — The specific epithet, antennifera, refers to the resemblance of the elongate appendages at the

apex of the corolla tube to insect antennae.

Classification. — Columned antennifera appears to belong to section Ortholoma Benth. because of the pres-

ence of corolla appendages and an obligate epiphytic habit. However, the traditional sectional classification of

Columned is artificial and arbitrary. As an example, the section Ortholoma has been recognized at the generic

level as T richantha Hook, by previous authors (Morton 1963; Wiehler 1973, 1975). A monographic revision of

Trichantha by Morton (1963) was followed by another paper by the same author (Morton 1971) with a reduc-

tion of all species recognized as Trichantha to Columned. The type species for Trichantha is Columned minor

(Hook.) Hanst. and is characterized by the presence of appendages at the sinuses of the corolla (Fig. 3C, D). It

is important to note that corolla appendages are not a unifying character for section Ortholoma or genus

Trichantha. The type species for section Ortholoma is Columnea anisophylla DC., which lacks corolla append-

ages as do many other species that have been assigned to this section. The traditional sectional classification of

Columnea has been shown to be artificial because many sections do not represent monophyletic lineages

(Smith 1994; Smith & Sytsma 1994; Clark et al. 2006). A revised sectional classification system based on mo-

lecular sequence data is currently a collaborative research focus by numerous authors (e.g., James Smith, John

L. Clark, Lacie Schulte and others).

Journal of the Botanical Research Institute of Texas 6(2)

389

Fw - 3 Variation in corolla appendages present in Columnea. Aft B. (oiumnea coronata Amaya, LE. Skog ft LP. Kvist C ft 0. Columnea minor (Hook.)

Hanst. E. Columnea rosea (CM. Morton) CV. Morton, f, Columnea mem (Wiehler) L.E. Skog & LP. Kvist (Photos A & B from/L Clark etal. 12990; C from

J .L Clark etaL 10870; D from J.L Clark etal. 9647; E from J. Betancur 12394; F from J.L Clark etal. 7140).

Corolla appendages vs. corolla lobes. — The presence of corolla appendages is widespread in Columnea and

this character is often not accurately distinguished from corolla lobes. Various taxa from different sections

have corolla appendages and this character is probably convergent within Columnea. For example, Columnea fi-

lifera (Fig. 3F) has been assigned to section Collandra Lem. (Kvist & Skog 1993) and genus Dalbergaria Tussac.

(Wiehler 1992; Kvist & Skog 2004) because of subsessile leaves, dorsiventral shoots, and a facultative epiphyt-

ic habit. A recently described species, Columnea coronata Amaya, L.E. Skog & L. P. Kvist, was assigned to sec-

tion Collandra (Amaya et al. 2004), where the specific epithet refers to a “corona” at the apex of the corolla tube.

More accurately, the “corona” in Columnea coronata is homologous to reduced corolla lobes like those found

in Columnea antennifera. Thus, what Amaya et al. (2004) referred to as “petals” in the description of Columnea

coronata are actually appendages that appear petaloid (Fig. 3A) and what was described in Amaya et al. (2004)

as the “corona” is homologous to reduced corolla lobes. Another species that has reduced corolla lobes and pet-

aloid appendages is Columnea filifera (Fig. 3F). The petaloid appendages in Columnea filifera (Fig. 3F) are visible

in the field and in photographs, but the corolla lobes are only visible with a hand lens or microscope. Phyloge-

netic studies on the evolution of corolla appendages and their presence in numerous lineages of Columnea will

play an important role in understanding their function, homology, and role in plant-pollinator interactions.

ACKNOWLEDGMENTS

This study was supported by funds from the National Science Foundation (DEB-841958 and DEB-0949169).

We thank Christian FeuiUet (US) and William R. Anderson (MICH) for help in selecting the specific epithet;

Alain Chautems (G), Laurence E. Skog (US), and Jeremy Keene (BHO) for providing helpful reviews of the

manuscript. Our 2012 research expedition to Colombia was a tremendous success because of logistical sup-

port from Alvaro Idarraga (HUA), Felipe Cardona (HUA), Julio Betancur (COL), Alvaro Cogollo (JAUM), and

Diego Suescun (JAUM). We thank Gunter Gerlach from the Munich Botanical Garden (Botanischer Garten

Munchen-Nymphenburg) for sharing his observations and images of Columnea antennifera. We gratefully ac-

knowledge Norris H. Williams (University of Florida) for his carefully curated slide collection that included

field images of voucher specimens (e g., Columnea filamentosa- Fig. 2 E, F) from a 1972 research expedition to

Colombia with Hans Wiehler. We thank Julio Betancur from the Universidad Nacional de Colombia (COL) for

providing images of Columnea rosea (Fig. 3E).

w species of Columnea (Gesm

Amaya, M., LE. Skog, and L.P. Kvist. 2004. Novae Gesneriaceae Neotropicarum XII: four n

ceae) section Collandra from Colombia. Edinburgh J. Bot. 60:41 5-424.

Burtt, B.L. and H. Wiehler. 1 995. Classification of the family Gesneriaceae. Gesneriana 1 *1-4

Cl* J.L.PS. Herendeen, L.E. Sk*. » EA 2-mer. 2006. Phytogene, ic relationships and generic boundaries in the Eph

^'^aasn^n’^n' 10 " ^ “"** SK,k>m ** 9 *" us Contr. Natt

Morton, C.V. 1963. A revision of Trichantha (Gesneriaceae). Contr. U.S. Natl. Herb. 38:1-27.

Morton, C.V. 1 971 . A reduction of Trichantha to Columnea (Gesneriaceae). Phytologia 22:223-224

Skog, LE. andJ.K. Boggan. 2006. A new classification of the Western Hemisphere Gesneriaceae. Gesneriads 56:12-17.

^4:1^9 yS,ema,ICS of Columnea SMion and section Srygnomhe (Gesneriaceae). Syst. Bot. Monogr.

,M4 ' MOleCUlK and m ° rPh0l09y: COn9nK "“ <* d “ (Gesneriaceae). PI. Syst

familiM ^ r era ° f vascuia ' pian,s - f,mem

7:63 1 58 H mdC<ide Inciua| ng Avicenmaceae). Springer-Verlag, Berlin & Heidelberg.

Wiehler, H. 1992. New species of Gesneriaceae from the neotropics. Phytologia 73:220-241.

Ill

A NEW VARIETY OF DECLIEUXIA CACUMINIS (RUBIACEAE)

FROM THE STATE OF TOCANTINS, BRAZIL, AND A

REVIEW OF THE VARIETIES RECOGNIZED IN THE SPECIES

Joseph H. Kirkbride, Jr.

Piero G. Delprete

USDA-ARS, U.S. National Arboretum

Floral & Nursery Plants Research Unit

350 1 New York Avenue NE

Washington, DC, 20002-1958, U.S.A.

joseph.kirkbride@ars.usda.gov

Herbier de Guyane, IRD - UMRAMAP

Boite Postale 165, 97323 Cayenne Cedex

Guyane Fran^aise (French Guiana), FRANCE

piero.delprete@ird.fr

ABSTRACT

RESUMO

INTRODUCTION

Kirkbride (1976) revised the genus Declieuxia Kunth, and recognized 27 species and seven varieties in three

species: D. deltoidea Mull. Arg. with two varieties, D. cacuminis Mull. Arg. with two varieties, and D. cordigera

Mart. & Zucc. ex Schult. & Schult. f. with three varieties. A few years later (Kirkbride 1983), he added a third

variety for D. cacuminis, D. cacuminis var. decumbens].H. Kirkbr. With this, the known range of D. cacuminis

was expanded from central Minas Gerais to south central Bahia. In 1997, another species was added to the ge-

nus, D. decumbens J.H. Kirkbr. (Kirkbride 1997), increasing the number of species to 28.

The Rubiaceae were treated for the Flora dos E stados de Goids e Tocantins by Delprete (2010). The Distrito

Federal, which is surrounded by the state of Goias and only touchs the state of Minas Gerais on its southeastern

comer, was also covered by the treatment. Delprete found nine species of Declieuxia in the state of Goias, six in

the Distrito Federal, and just one, D.fruticosa (Willd. ex Roem. & Schult.) Kuntze (the most widespread of the

genus), in the state of Tocantins. Early in 2012, while finalizing a treatment of the Rubiaceae for the Flora do

Distrito Federal, Brasil, Delprete encountered a specimen of Declieuxia from the state of Tocantins in the herba-

rium of the Reserva Ecologica do Instituto Brasileiro de Geografia e Estatistica (1BGE) in Brasilia, that attracted

his attention. Since he had reported just a single species of Declieuxia from the state of Tocantins, he immedia-

tely knew that the specimen represented a new taxon for this state, and possibly a new taxon for science. He

consulted Kirkbride about the identity of the specimen, who in turn compared it with the Declieuxia speci-

mens in the herbaria of the Smithsonian Institution (US) and New York Botanical Garden (NY). They conclu-

ded that the specimen is a member of D. cacuminis and is a fourth, new variety of the species, which is described

and illustrated below. In addition, a key to the four varieties of D. cacuminis, short descriptions, and selected

specimens for each variety, are also presented.

J- Bot Res. Inst Texas 6(2): 391 -396.2012

393

395

Journal of the Botanical Research Institute of Texas 6(2)

of Tocantins, west of the Rio Sao Francisco, and at lower elevation in cerrado vegetation (a type of savannah),

on sandy, reddish soils poor in nutrients and rich in aluminum (Fig. 1).

This new variety was collected in the Jalapao region, which has large areas of white-sand soils. The eleva-

tion is lower with respect to the elevations of the other three varieties; however, the type locality is not on the

Brazilian Planalto. It is on a lower plateau, at the northernmost end of the Serra Geral de Goias, which is the

divide between the basins of the Rios Sao Francisco and Tocantins at the border between the states of Tocan-

tins and Bahia. Relative to the nearby river valleys, this area is 50-100 m higher, so the new variety occurs at :

the highest elevations in its area of distribution as do the other three varieties in their areas of distribution.

ACKNOWLEDGMENTS

We thank Laurence J. Dorr, Department of Botany, Smithsonian Institution (US) for his helpful review of the

manuscript and the directors and curators of the following herbaria for allowing us access to their collections

and making loans of material: IBGE, NY, UB, UFG, and US.

REFERENCES

Delprete, P.G. 201 0. Rubiaceae, parte 1 , introdu^ao, generos A-H. In: J.A. Rizzo, ed. Flora dos Estados de Gioas e Tocantins.

IRD & UFG, GoiSnia, GO, Brazil. Vol. 40(1 ):1 -580.

Kirkbride, J.H., Jr. 1976. A revision of the genus Declieuxia (Rubiaceae). Mem. New York Bot. Gard. 28(4):1-87.

Kirkbride, J.H., Jr. 1983. A new variety of Declieuxia cacuminis (Rubiaceae) from Bahia. Ann. Missouri Bot. Gard. 70:

204-205.

Kirkbride, J.H, Jr. 1997. Manipulus rubiacearum— VI. Brittonia 49:354-379.

A PREVIOUSLY UNRECOGNIZED SPECIES OF SENEGALIA (FABACEAE)

FROM NORTHEASTERN BRAZIL

David S. Seigler

Department of Plant Biology

University of Illinois

Urbana, Illinois 61801 U.S.A.

seigler@life.illinois.edu

John E. Ebinger

Department of Biology

Eastern Illinois University

Charleston, Illinois 61920 USA

Petala Gomes Ribeiro

Departamento de Botdnica

Universidade Estadual de Feira de Santana

Av. Transnordestina, s/n, Novo Horizonte

44036-900, Feira de Santana, Bahia, BRASIL

RESUMEN

During the course of our work on the genus Senegalia Raf. of Brazil, we encountered a collection [M. Blanchet

2772 (F, GH, K)] that Bentham (1842) cited first as Acacia velutina DC. and subsequently (1875, 1876) as Acacia

monacantha Willd. This material had mostly have been accepted under the latter name, but as more collections

have become available, it is clear that they represent a previously undescribed species distinct from both Sen-

egalia monacantha (Willd.) Seigler & Ebinger (syn. Acacia monacantha) and Senegalia velutina (DC.) Seigler &

Ebinger (syn. Acacia velutina ).

Senegalia paganuccii Seigler, Ebinger, & Ribeiro, sp. nov. (Fig. 1). Type: BRAZIL. Bahia: Municlpio Rio de Contas, 10 km

do Rio de Comas na estrada para Marcolino Moura, Caatinga (13°36S, 41°43W, elev. 500-600 m), 15 Nov 1988, gemmae (buds,

gem.) & flowers (fl.), R.M. Harley, D.J.N. Hind & T.B. Cavalcanti 26439 (holotype: HUEFS; isotypes: CEPEC, CTES, F, K, NY, SP, SPF).

Shrub or small tree to 6 m tall, ramified from the base; bark grayish; twigs reddish brown to dark purple, not

flexuous, terete, usually puberulent; short shoots absent; prickles brown to purple-brown, apex usually darker,

flattened, recurved, woody, 1-3 mm long, 1-3 mm at the base, glabrous, scattered along the twig, petiole and

rachis, commonly paired at some nodes; perulate buds commonly presently at leaf axil, ovate or elliptic in

profile, 5-7 x 2-4 mm. Leaves alternate, commonly paired at some nodes, 30-100 mm long; stipules light to

dark brown, lanceolate, symmetrical, flattened, straight, herbaceous, 4-10 x 1.5-5.5 mm, pubescent and cili-

ate, early deciduous; petiole slightly grooved adaxially, 6-19 mm long, pubescent; petiolar gland solitary, usu-

ally located just below the lowest pinna pair, sessile, attached near the middle, the margins of the petiolar gland

turning upward, oval to orbicular, 0.6-1.5 mm across, cup-shaped, glabrous; rachis adaxially grooved, 25-80

mm long, pubescent, an oval gland 0.4-0.9 mm across between the upper and sometimes other pinna pairs,

rachis glands 0.4-0.8 mm across, oval, cup-shaped, glabrous; pinnae 4 to 11 pairs per leaf, 25-45 mm long,

4-10 mm apart; paraphyllidia 0.3-0.7 mm long, mostly absent; petiolule 0.8-1.5 mm long; leaflets 25 to 40

pairs per pinna, opposite, 0.7-1.1 mm apart, linear, 4-6 x 0.9-1.4 mm, glabrous, lateral veins not obvious, 1

>■ Bot Res. Inst Texas 6(2): 397 - 401 . 2

398

Fk. 1 . Senegaliapaganuccii Seigler, Ebinger, & Ribeiro. k. Flowering branch. B

F. Flower at anthesis. G. Gynoecium showing detail of ovary. H. Prickles. I. Vegetative bud detail. J. Fruit. K. Detail of innt

H-l from RM. Harley etal. 26439 (HUEFS); J-L from >1.4. Conceipoetal. 1925 (HUEFS).

vein from the base, base oblique and truncate on one side, margins ciliate, apex obtuse, midvein submarginal

Inflorescence a densely 40-75-flowered cylindrical spike, 9-15 x 20-45 mm, 1 to 3 in the leaf axils; peduncles

6-22 long, 0.4-0.6 mm thick, pubescent; receptacle not enlarged; involucre absent; floral bracts spatulate.

8-15i

AACCM-MCGCGCGTCCMCTGGACG. .

. . . . GCGCCCGAGGCCT-

SCCMCM? ? ? TGCACCGCACGGAA

Fk. 2. ITS sequence data for Senegalia paganuccii.

Paraphyllidia

" 1X0 1 ' Jlna estraaa Serrana e Mocambmho, 5 Sep 1974, M. Magalhaes & M.B. Ferreira 5021 (IPA). HAUL Sao Raimundo INonare-

entre Sao Raimundo Nonato e Anisio de Abreu, caatinga, 17 Nov 1981 (fl.), M.R. DeVArco & E. Nunes 2214 (IPA, TEFB).

Bentham (1842) in his discussion accompanying the description of Acacia velutina DC., first mentioned A

monacantha Willd. and concluded that if it were a true Acacieae, it would belong to this tribe. Under A. velutina,

Bentham (1842) cited Blanchet 2772 from Bahia and Pohl s.n. from Brazil, as possibly belonging to this species.

However, our studies show that the type of Acacia velutina DC. (BRAZIL. RIO DE JANEIRO: [holotype: G,

(photos F, MO, SI); isotype: G]), is distinct both from Blanchet 2772 (“habitat ad Utinga in deserto fluminis S.

Francisco provinciae Bahiensis”) and the Pohl collection (“inter Praia et Bom Jardim Provinciae Minas

Gerais”). Later, Bentham (1876) considered Blanchet 2772 (and another collection, apparently from the same

location (Blanchet 3772), the Pohl collection and a Lindberg coUection (“ad Santos in provinceae Sao Paulo”) to

represent Acacia monacantha Willd. Additionally, he considered Acacia velutina Benth. (1842) to be a synonym

of A. monacantha Willd. Today these species are regarded as distinct, namely, Senegalia monacantha (syn. Aca-

tina ), and the Blanchet and Pohl collections i

da monacantha ) and Sengalia velutina (syn. Acatia velut

to our new species, 5. paganuctii.

The descriptions of Bentham (1875, 1876) and Lewis (1987) for Acada monacantha and the description

and illustrations (p. 201, HI, H2, and H3) of Senegalia monacantha of de Queiroz (2009) all appear to be based

on Blanchet 2772 and similar collections and not on the original type material of Acacia monacantha Willd. We

The authors wish to thank several colleagues for advice concerning questions of nomenclature and general

taxonomic advice, in particular, K.N. Gandhi. We wish to acknowledge support by the National Science Foun-

dation (NSF DEB 04-15803), by the American Philosophical Society (1992) and by the CNPq for financial sup-

port to PGR, on her master’s degree in the Postgraduate Program in Botany at the Universidade Estadual de

Feira de Santana, Bahia, Brazil; also to the curators of the herbaria that were visited or provided loans of speci-

mens for our study (ALCB, BHCB, CEN, CEPE C, F, G, H, HRB, HST, HUEFS, IBGE, IPA, K, MBM, MO, NY, RB,

R, SP, SPF, TEX, UB). We thank Bruce R. Maslin and an anonymous reviewer for helpful comments.

REFERENCES

Bentham, G. 1842. Notes on Mimoseae, with a synopsis of species, London J. Bot. 1:318-392, 494-528.

Bentham, G. 1875. Revision of the suborder Mimoseae. Trans. Linnaean Soc London 30:335-664.

Bentham, G. 1876. Leguminosae. In: C.F.P. von Martius, Flora Brasiliensis 15(1 &2). Munich and Leipzig. 1-527 pp., 1-138

Queiroz, LP., de, 2009. Leguminosas da Caatinga. Universidade de Feira de Santana, Feira de Santana.

Lewis, G.P. 1 987. Legumes of Bahia. Royal Botanical Gardens, Kew, England.

appears to be most closely related to S, tenuifolia (L.) Britton & Rose. The characters used to distinguish Senega-

402

Journal of the Botanical Research Institute of Texas 6{2)

BOOK REVIEW

WelbyR. Smith. Illustrations by Vera Ming Wong and Bobbi Angell. 2012. Native Orchids of Minnesota. (ISBN-

13: 978-0-8166-7823-5, hbk.). University of Minnesota Press, 111 Third Ave. South, Suite 290, Minne-

apolis, Minnesota 55041, U.S.A. (Orders: www.upress.umn.edu; 612-627-1980 fax; 612-627-1970

phone). $34.95, 288 pp„ 93 b/w illustrations, 174 color plates, 53 maps, 7" x 10".

Nineteen years after publishing Orchids of Minnesota, the author has published an updated, comprehensive,

and beautifully illustrated new volume. Native Orchids of Minnesota. His Table of Contents, map of the counties

in Minnesota where orchids are most likely to be found, and Preface provide an excellent “starting platform"

for the Introduction and his gentle guidance into the descriptions and basic understanding of Minnesota’s or-

chids.

After providing straightforward answers to the usual questions flower lovers ask, he provides a thorough

and easily useful “Key to the Genera of Orchids Found in Minnesota.” An unusual but very helpful addition to

the keys is the author’s careful inclusion of two or three simple line drawings of the described plant part at the

end of each leg of the key. The reader has an immediate and accurate vision of where and what to look for as (s)

he examines the plant.

Each genus is introduced on the left page and accompanied by a photo on the right page. The following

pages provide thorough descriptions, explanations, photos, and illustrations of the individual species in each

genus. Most orchid fanciers will find themselves totally enchanted by the “reading and admiring” process.

Obviously, this book would be incredibly useful to have with you in the field — not only for identifications but

also to have use of the accompanying maps of Minnesota counties, showing where each species is most likely

to be found.

(This is such a beautiful book, I think I would need to buy two copies: one to take into the field for quick

and accurate identification and one to keep safely at home in an easily available, beautiful “pristine condi-

tion!”).— Helen Jeude, Volunteer and Assistant Editor, Botanical Research Institute of Texas, 1700 University Dr.,

Fort Worth, Texas 76107-3400, U.SA.

TAXONOMY OF LANTANA SECT. LANTANA (VERBENACEAE):

II. TAXONOMIC REVISION

Roger W. Sanders

Bryan College It 7802

721 Bryan Drive

Dayton, Tennessee 37321, U.S.A.

RESUMEN

Due to a long history of cultivation, hybridization, and invasiveness, the taxonomy of Lantana L. sect. Lantana

resists partitioning into easily identified species (see Sanders 2006 for review). While some workers might

prefer the convenience of recognizing a single highly variable species, Lantana camara L., previous biosyste-

matic studies (Sanders 1987a, 1987b, 1987c, 1989) have shown the presence of morphologically discrete dip-

loid taxa having coherent ecological and geographic ranges where they appear to have speciated in situ. These

studies have been corroborated by a recent molecular analysis of the taxa in Florida (Maschinski et al. 2010).

The present study attempts to delineate the indigenous taxa of sect. Lantana, even in the face of rampant hy-

bridization due to human-induced ecological disturbance and the failure of odd polyploidy as a breeding bar-

rier in this group. This second paper in the series builds on the first (Sanders 2006), which detailed the typifica-

tion of species of sect. Lantana. As suggested by Sanders (2006), the identity of individual specimens constitut-

ing the hybrid plexus found growing outside cultivation today cannot be unraveled by morphology alone, and

it may be recalcitrant even to molecular genome analysis. Thus, variation encompassing the indigenous species

now connected by hybrids may appear to be constituted more of adaptive peaks rather than bell curves sur-

rounded by discontinuity. Furthermore, no phylogenetic analysis has been attempted here because the out-

group relationships of sect. Lantana are not understood, and the significant trichome and inflorescence char-

acters are homoplastic with regard to potential outgroups.

The rank of series is established only for grouping species of presumed origin by divergence from the an-

cestor of sect. Lantana. Species of interseries hybrid origin are not placed into series and are listed separately.

These species are presumed to have originated by natural selection acting on the variable pool of original hy-

brids resulting in one or a few closely similar phenotypes and, thus, may not be strictly intermediate to the

Journal of the Botanical Research Institute of Texas 6(2)

parental species. The surviving phenotype has become self-propagating and has attained a geographic range

exceeding the original area of sympatry. Spontaneous and cultivated hybrids that have received Latin names

but do not behave biologically as species are in a third separate list.

1 view varietas as the least inclusive taxonomic rank composed of a minimum of one breeding population |

(as inferred from available ecological data) having geographic coherence in a limited part of the species range

and imperfect discontinuity from similar, geographically adjacent taxa within the species. Subspecies is used

either to group varieties or to recognize a taxon within a species with geographic coherence over an extensive

geographic range (e.g., usually several islands or subcontinental areas) and having imperfect discontinuity orl

minor differences from similar, geographically adjacent subspecies. One impetus to employ subspecies in Lon - 1

tana has been to avoid instability in infraspecific names that could be caused by the subsequent discovery of i

poorly known varietal names, of which there are many.

MORPHOLOGICAL TAXONOMIC CRITERIA AND ANALYTICAL CAVEATS

Caveats for identification and descriptions are given in italics.

Prickles . — Whereas a majority of species either lack prickles or bear only small weak straight or recurved |

prickles, pronouncedly stout recurved prickles are inconsistently present (varying among herbarium collec-

tions and field populations) in the remaining species, notably Lantana camara subsp. aculeata, L. hirsuta subsp. 1

amazonica, L. horrida, L. nivea, L. planaltensis, L strigocamara, L. urticoides, and L. viscosa. The tendency to 1

produce prickles appears more pronounced in hybrids than in most indigenous species.

Trichames . — The form of trichomes on the abaxial leaf surfaces (filiform vs. setiform vs. strigiform) and

their length are highly correlated with ecological and geographic coherence of indigenous taxa and provides |

can be nearly twice as long as those on the remaining tissue. Therefore measurements in the key and descriptions for I

adaxial hairs are taken between the secondary veins. Likewise, hairs on the nodal lines of the stems are often about

twice as long as other hairs along the stem and are excluded from measurements in the key and descriptions.

Filiform hairs and setae are both erect from the base with the setae differing primarily by greater length

and stouter, and a more conical proximal portion. Both types may be somewhat flexuous, arching or curly 1

Strigae are stiff conical hairs that are geniculately bent in the proximal quarter or third with the remain-

ing distal portion directed antrorsely. On the adaxial leaf surfaces, the antrorse portion is more or less ascend- 1

ing and arching. The broadened base emerges from a buttressing ring of epidermal cells that form a pustulate

base. Especially in Ser. Strigosae or taxa of hybrid origin with genes of its species, the strigae often are decidu- 1

ous leaving the pustulate bases as rough points. In some species the bases enlarge with age and become vitre- J

ous (clear or white). In taxa and hybrids with strigae on the abaxial surface, the strigae lack the buttress base,

arising directly from the epidermis and the antrorse portion is held more or less parallel to the epidermis. In 1

some Lantana nivea, they are so short that the antrorse portion is not well developed, appearing as a short point 1

angling upwards. In recognizing the abaxial strigae, one must also be aware that filiform or setiform hairs that are

crushed against the surface during pressing can be mistaken for strigae, which occur consistently over the pertinent j

The co-occurrence on the abaxial surfaces of filiform hairs or setae with stn

brid nature or heritage of the specimen at hand.

The presence of stipitate glands on the twigs, peduncles, petioles, and ei

several taxa, noteably Lantana camara subsp. portoricensis, L. horrida subsp. 2

crantha, L. paraensis, and L. planaltensis. While glands are consistent in L. le

indicat

1 of the hy-

■af-blades is variable within

ii and subsp. tiliifolia , L. mi-

1 the basis of glands. Therefor

it presence of glandular ha

iara is simply variable in tl

nages, with the widest poi

alf, making it difficul

shape. Bases of the blades in almost all cases abruptly taper to a narrow wing onto the petiole distally. Leaf-

blades are considered to be triplinerved (as opposed to pinninerved) if the basal pair (or two pairs) of secondary veins

among them. Nigrescence refers to a distinct blackening of mature leaves occurring during drying for preservation,

not normal senescence. While such blackening is diagnostic for certain species, newly emerging leaves can blacken in

most species. Leaves atypical for size, shape, bases, apices, and venation are present on most plants. Leaf shape, size,

and vestiture traits are measured only on fully developed, non-senescent leaves. Measurements for marginal teeth are

taken mid-margin, avoiding the reduced teeth near the base and apices of the blades.

Inflorescences.— The basic structure of the inflorescences has been discussed in detail (Sanders 2001).

Peduncle length in the key and descriptions are given for fully opened inflorescences and infructescences.

Bracts generally decrease gradually in length and width from the proximal to the distal series that seem-

ingly spiral up the receptacle. The distal (inner) bracts are about 2-4 mm long and about 0.3-1 mm wide in

most species and, thus, are not detailed in the descriptions. The exceptions appear to be diagnostic in Ser. Spi-

catae and a few taxa in Ser. Lantana in which almost all the bracts are the same dimensions. In some taxa there

is an abrupt diminution from the proximal (basal or outer) two or three series of bracts to the more distal series.

Shape and size of the proximal bracts appear to be consistent and diagnostic, with the exception of one or

rarely two subfoliaceous bracts that develop sporatically in almost any taxon; hence, these atypical bracts are exclud-

ingfrom the measurements.

Flowers . — Corolla color has been discussed by Sanders (2001, 2006). It often changes from bud to early

opened flowers to late flowers to fading flowers, especially in plants that produce both yellow to carmin pig-

ments and purplish pigments. The throat is often not only different but changes during flower maturation.

This developmental variation is often further complicated by intraspecific variability. Unfortunately, detailed

information is usually lacking from collection labels. Corolla shape is nearly uniform in the group, but size

appears to be consistent within taxa when measured from fresh material. However, dried corollas are often

shrivelled and difficult to measure; in the descriptions, “fresh” size has been extrapolated from dried specimens.

Cytology. — Chromosome numbers are not given in the descriptions because those for only a few taxa are

known (See Sanders 1987a, 1987b, 1989).

Phenology . — Flowering time is not given in the descriptions because any species can flower anytime dur-

ing the year whenever moisture is available. This is true even of species l

seasons when the species are grown in frost-free areas.

TAXONOMIC TREATMENT

See Sanders (2001) and Sanders (2006) for further characterization of Lantana and comparison of sect. Lantana

with other sections. Also, see Sanders (2006) for details of species typifications, which are supplemented here

only as needed. Please note that, in the type and other specimen citations, the abbreviation “di” refers to a

digital image made available online or as a courtesy by the cited herbarium. Many thousands of specimens

representing this group are in herbaria awaiting identification. For this study only a small sample, primarily

from major U.S. institutions, has been selected for annotation and citation here as these specimens will be most

easily available for consultation by other professionals. Even so, annotations made during quick visits to her-

baria may differ than those cited herein as a result of reflection and more careful study of digital images that I

made or were sent to me. Selection of specimens was to establish only distribution limits of the species, as well

as document as many hybrid combinations as possible, thus, resulting in a falsely apparent predominance of

hybrids in some cases. To assist those attempting to identify specimens of sect. Lantana, a richly illustrated

interactive key (in which vernacular names are also discussed and provided) has been made available online

(Offutt & Sanders, 2012).

Lantana L. sect. Lantana

Shrubs or rarely treelets, erect to decumbent or subscandent, height (or length) (0.1-)0.5-3 m (to 4 or even 6 m

m subscandent, especially aggressively naturalized forms); the internodes usually less than to almost twice as

Journal of the Botanical Research Institute of Texas 6(2)

long as leaves (mostly twice to thrice as long in L. splendens), with or without weak to stout, conical to recurved

prickles; vestiture antrorsely strigose to puberulent, pilose, setose, glabrescent, or stipitate-glandular and thus

markedly viscid, the trichomes of twigs, peduncles, and petioles often noticeably longer and stiffer than those

on remaining herbage. Leaves opposite or sporadically ternate, petiolate, simple; blades usually ovate or lan-

ceolate to elliptic, usually hardly to moderately rugose, i.e., puckered between tertiary veins (strongly so in

some L. horrida and usually bullate in L. leonardiorum, which is puckered between the secondary veins), usu-

ally longitudinally flat or somewhat undulate (incurved in L. depressa ); base attenuate to cordate; apex attenu-

ate, acuminate, acute, obtuse, or occasionally rounded; margin usually finely serrate-crenate but coarsely so in

L. urticoides and some L. hirsuta and L. kingii or subentire in some L. cujabensis, flat to revolute, usually green

(often purple-tinged in L. kingii); adaxial surface strigose (strigae typically ascending distally to antrorsely

bent, ± appressed in L. kingii, flaccid and strongly appressed in L. hodgei), strigose-villous, setose-villous, or j

nearly glabrous; abaxial surface strigose, pilose, puberulent, setose, or glabrescent, with the veins green to pale

brown or sometimes nigrescent or occasionally tinged with purple (frequently purplish in L. kingii). Inflores-

cences pedunculate, capituliform spikes, one (or sporadically two in several species) per subtending leaf; pe-

duncles about a third to twice the length of leaves (up to four times in some L. horrida); axis (common recepta-

cle) ellipsoid or fusiform, spongy; bracts nearly always subtending a flower, linear triangular or linear lanceolate

to oblong, elliptic, or spatulate, ± appressed to spreading or reflexed. Flowers in several series, two to three

series in anthesis at a time, zygomorphic; corolla salverform with inflexed tube and four unequal lobes, pig-

ments either yellow to reddish or pink to purple or admixtures of both (in hybrids or taxa of hybrid origin) or

lacking. Drupes usually blue-black (but sporadically described by collectors as dark violet-purple), usually

with a metallic iridescence; pulp watery-mealy; endocarp turbinate-obpyriform with an inflated commissure and

external circumferential ridge below the seed chambers. x=ll.

407

409

broadly ovate or oblong-deltate to elliptic-lanceolate, (l-)3-8(-16) cm long, the length (0.9-)1.5-2.5 x width,

usually not nigrescent, papery, pinninerved; base subcordate, truncate, rounded or broadly cuneate, usually

very briefly, narrowly cuneate onto petiole at very base; apex acute to acuminate, occasionally attenuate or

rounded; marginal teeth 6-35(-50) per side, rounded to acute, spreading to appressed, sometimes with tips

recurved, with sinuses 0.2-2 mm deep; adaxial surface dull, antrorsely strigillose to strigose-pilose or with

stipitate glands mixed in, the hairs occurring on veins and intervening tissue, thin canopy of hairs only 0.2-0.5

mm (occasional hairs 0.7 mm in subsp. aculeata ) with understory of shorter hairs not well developed, 10-90(-

strigae ca. 0.1-0.2(-0.3) mm in diam.; abaxial surface duller green than adaxial surface, moderately densely

(occasionally sparsely) pilose, the hairs on all veins and intervening tissue, 0.2-0.5 mm, all about same length,

(10-)40-250/sq. mm. Inflorescences remaining hemispheric; peduncles 0.3-2 x leaf length. Proximal bracts

above middle, with 3 veins from the base, appressed or spreading, deciduous after flowering; apex attenuate to

rounded; indument pilose to strigillose, sometimes stipitate-glandular, somewhat or not ciliate, the longest

hairs £ 0.5 mm. Corolla yellow to or aging reddish orange (infused with pink or purple in subsp. aculeata),

rarely white; corolla tube 4-12 mm.

Distribution and habitat. — Mexico, Central America, West Indies, and northern South America; cultivat-

ed and escaped pantropically, especially in Australia; disturbance openings in tropical evergreen and decidu-

ous forest, open pine forest, thorn shrubland, savanna; 0-2000 m.

Lantana urticifolia Mill., Card. Diet. ed. 8, Lantana 5. 1768. Lanu

(misapplied to L. camara subsp. aculeata). Type: MEXICO. >

un s.n.. Herb. Sloan 6:84 (lectotype: BM-

id Virgin Is

412

Journal of the Botanical Research Institute of Texas 6(2)

Stems usually without prickles or with few weak, straight ones; twigs, peduncles and often petioles moder-

ately to densely covered with usually ascending, soft to somewhat stiff, curled or straight hairs, the hairs 0.1- ;

0.5(-0.7) mm, mostly ca. 0.3 mm. Leaf-blades broadly ovate to oblong-deltate to elliptic lanceolate, widest

usually in or near proximal third, sometimes near middle, (1.5-)3-7(-9) cm long, the length (1.1— )1.3— 2 x|

width; marginal teeth 20-35(-50) per side, rounded, obtuse, or acute, often appressed, with sinuses 0.3-0.8(-|

1) mm deep; adaxial surface antrorsely strigillose to strigose-pilose, the hairs mostly about 0.3 mm or less.

Peduncles 0.5-1.2 x leaf length. Bract series gradually reduced in size; proximal bracts linear-oblong, oblan-

ceolate-oblong, linear-lanceolate, or linear-triangular, 4-8 mm long, 0.5-1.5 mm wide, widest near the base or

the outermost series sometimes widest above middle (if 4 mm or less long, then widest near the base); apex

acute to attenuate. Corolla yellow to or aging reddish orange; corolla tube 7-12 mm; corolla limb 6-9 mm in

a subsp. glandulosissima (Hayek) R.W. Sanders, comb. & stat. i

epert. Spec. Nov. Regm Ytg. 2 IM l'H'o Tm: MEXICO. Jalisi .. Tequila 2 Jul

ilO!, NY!, P[2,di!]).

Stems usually without prickles or with few weak, straight to recurved ones; twigs, peduncles and often peti-

oles densely covered with stipitate glands or also with eglandular filiform hairs mixed in, the hairs (and

glands) 0.1-0.5 mm, mostly 0.2-0.3 mm. Leaf-blades broadly ovate or broadly elliptic to oblong-lanceoate or

elliptic-lanceolate, widest usually near proximal third or middle, (l-)4-10(-16) cm long, the length (1.2-)l-jB

2.1 x width; marginal teeth 10-30(-45) per side, usually rounded or obtuse, usually spreading, with sinuses

(°.4-)0,7-1.5(-2) mm deep; adaxial surface mixed antrorsely strigillose to strigose-pilose and stipitate-glandu-

lar, the hairs 0.1-0.5 mm. Peduncles 0.5-1.8 x leaf length (often almost doubling in length in fruit). Bract se-

nes gradually reduced in size; proximal bracts oblanceolate-oblong (rarely obovate) to triangular-oblong or !

Sanders, Taxonomy of Lantana sect. Lantana

413

linear-lanceolate, (2.5-)4-8 mm long, 0.8-1.7(-2) mm wide, widest above or near the middle or near the base;

apex acute to attenuate, often rounded at very tip. Corolla yellow to or aging reddish orange, rarely white; co-

rolla tube (5-)7-12 mm; corolla limb 6-9 mm in diam.

Distribution and habitat . — Mexico (northwestern, central, and southern) and Central America to north-

ern Colombia and Venezuela; open pine-oak forest, thorn and tropical deciduous shrubland and woodland,

and savanna, especially in disturbance openings; 0-2000 m.

Lantana camara subsp. glandulosissima differs from subsp. camara only in the strong development of

stipitate glands in place of filiform hairs on twigs, peduncles, petioles, and leaf surfaces and in the longer bracts

and corollas. Because the development of glandular hairs is variable within several other taxa in sect. Lantana,

parapatric or narrowly sympatric in the vicinity of Veracruz (as evidenced by the intermediate or hybrid

specimen, Gilly et al. 75, MSC), perhaps due to human activity. Although no specimens of subsp. glandulosis-

sima from Veracruz came to my attention, interspecific hybrids (see section below) further evidence its pres-

ence there. Furthermore, at least in Bocas del Toro Prov., Panama, subsp. glandulosissima intergrades with

subsp. moritziana ( Peterson &Annable 868 , MO).

Lantana camara subsp. glandulosissima is broadly sympatric with L. horrida. The two “pass the test of

sympatry” (Stebbins 1966, p. 95-96) despite occasional hybrids that are probably limited to disturbed areas. I

take this as evidence that L. camara and L. horrida are distinct. On the other hand, if one considered the differ-

ences in length of the adaxial leaf-surface trichomes an inadequate species criterion and submerged L. horrida

414

Journal of the Botanical Research Institute of Texas 6(2)

If. Lantana camara subsp. aculeata (L.) R.W. Sanders, Sida 22:394. 2006. Bask

Lantana sanguined Medik., Hist. & Commentat. Acad. Elect. Sci. Theod.-Palat. 3. Phys. 229. 1775. Camara aculeata (L.) Kuntze [var.

subinermis Kuntze] f. sanguined (Medik.) Kuntze, Revis. Gen. PI. 2:503. 1891. nom. illeg. (see synonyms below). Lantana aculeata L

r. Stirp. Chap. Allerton. 1C

Ecole Bot., ed. 3 (Cat. PI. I

n. illeg. Type: None selected.

Stems usually with stout, recurved prickles, often abundant; twigs, peduncles and often petioles moderately

covered with antrorse to ascending or retrorse, curled or straight hairs or also stipitate glands, the hairs 0.1-0 7 f

mm. Leaf-blades broadly ovate or oblong-deltate to elliptic lanceolate, widest usually in or near proximal

third, sometimes near middle, 3-9 cm long, the length (l.l-)1.3-2 x width; marginal teeth 10-30(-45) per

side, usually acute or obtuse, sometimes rounded, usually spreading, with sinuses 0.5-2 mm deep; adaxial

surface antrorsely strigillose to strigose-pilose, the hairs 0.1 -0.5 mm (occasional ones to 0.7 mm). Peduncles

0.5-1.2 x leaf length. Bract series gradually reduced in size; proximal bracts linear-oblong, oblanceolate-ob-

long, linear-lanceolate, or linar-triangular, 4-8(-10) mm long, 0.5-1.5 mm wide, widest near the base or

proximal third, sometimes the outermost one or two slightly broader above middle; apex usually attenuate.

Corolla yellow to or aging red-orange and usually infused with purple or opening pink aging to deep reddish

purple; corolla tube (5-)7-12 mm; corolla limb 6-10 mm in diam.

Distribution and habitat . — Historically cultivated worldwide and escaped pantropically, especially com-

mon in Africa and Australia; disturbance openings in tropical evegreen, deciduous, and thorn forest and sa-

vanna; 0-2000 m.

Selected specimens examined: AUSTRALIA. Queensland: Day 8 (BRIT). KENYA. Taita Taveta: Wakanene et al. 383 (MO). ZAIRE. Haut-

Katanga: Fabri 60415 (MO).

Presumed hybrids with: 9i-cvx20. L. Callowiana Hybrid Group cultivars (L. depressa-tetraploid x strigocamara). AUSTRALIA

Queensland: Day 64 (BRIT); Riding 76 (BRIT); Robazza & McAndrew 17 (BRIT). 12b. L. nivea subsp. mutabilis. AUSTRALIA. New South

Wales: Day 38 (BRIT); Day 42 (BRIT); Day 71 (BRIT). Queensland: Hannan-Jones 29 (BRIT); McAndrew 48 (BRIT); McAndrew 81 (BRIT);

McAndrew 84 (BRIT). CHINA. Guangdong: Deng Lang 10459 (BRITldi]). RHODESIA. Guruve: Nyariri 167 (MO). TANZANIA. Kiliman-

jaro: Mlongwa et al. 459 (BRIT). Tanga: Mwongofea & Kayombo 113 (MO). U.S.A. Hawai i. Oahu: Degener 11467, identification uncertain

(SMU). North Carolina. Forsyth Co.: cult,. Schallert 1352 (SMU). 20. L strigocamara. AUSTRALIA. Queensland: Day 69 (BRIT). TANZA-

Shrubs erect, rounded, lax, or trailing, dense to open; stems 0.5-3 m; branches ascending and normally sev-

eral to decumbent and few; twigs, peduncles and often petioles puberulent, setose, glabrescent, or stipita te '

glandular, the hairs 0.1-1.5 mm. Leaf-blades broadly ovate to rotund, elliptic or lanceolate-deltate, (0.5-)l-9(" ;

Sanders, Taxonomy of Lantana sect. Lantana

12) cm long, the length 1-2.2 x width, not nigrescent, papery, pinninerved; base usually rounded to truncate,

sometimes broadly cuneate or cordate, briefly narrowly cuneate onto petiole at very base; apex acuminate,

acute, obtuse, or rounded; marginal teeth (4-)6-25(-45) per side, acute, obtuse, or rounded, spreading or ap-

pressed, then sometimes with tips recurved, with sinuses 0.2-2.5 mm deep; adaxial surface dull, antrorsely

dense canopy of hairs 0.6-0.8 (-1) mm with understory of hairs 0.2-0.5 mm, (5-)10-50(-150)/sq. mm, not

noticeably vitreous-pustulate, the circular bases of the strigae ca. 0.1-0.2 mm in diam.; abaxial surface dull

green, moderatly densely (occasionally sparsely) pilose, the hairs on all veins and intervening tissue, 0.3-0.5

mm, all about same length except for a few scattered arching hairs 0.7-1 mm on the midrib or secondary veins,

10-200/sq. mm. Inflorescences remaining hemispheric; peduncles (0.5-)0.8-4 x leaf length (usually about

equalling to almost twice when mature). Proximal bracts lanceolate-triangular, lanceolate-linear or narrowly

elliptic, narrowly oblanceolate, narrowly oblong to oblanceolate-spatulate, 2-12 mm long, 0.5-3 mm wide,

widest in proximal, middle, or distal third, with 3(-5) veins from the base, appressed or spreading, persisting

or not; apex acute, attenuate or obtuse to rounded; indument strigose-pilose or setose, often sessile- or stipitate-

glandular, ciliate or not, the longest hairs mostly 0.3-1 mm. Corolla yellow to or aging reddish orange; corolla

tube 4-12 mm.

Distribution and habitat .— Mexico and West Indies to subtropical South America; tropical savanna with

gallery forest, montane humid, pine, or dry forest, and disturbed successional woodland, shrubland and grass-

land; 0-2500 m.

Journal of the Botanical Research Institute of Texas 6(2)

45:296. 1980. Type: CUBA. Isle of Pines: Siguanea^ 21 May 1910 Jennings 458 (holotype: NY!).

Shrubs erect, rounded, or lax, dense to open, the central axis ± developed, branches ascending or clambering

and several; twigs, peduncles and often petioles moderately setose, rarely with stipitate glands mixed among

the eglandular hairs. Leaf-blades ovate to broadly ovate or broadly elliptic, (l-)3-9 cm long, moderately to

weakly rugose, puckered between tertiary veins; apex acute to acuminate, occasionally obtuse or rounded;

marginal teeth (4-)10-25(-35) per side, with sinuses 0.5-2 mm deep; adaxial surface antrorsely strigose-velu-

tinous, the hairs 10-50/sq. mm. Peduncles (0.5-)0.8-2 x leaf length (usually about equalling to almost twice

when mature). Bract series gradually reduced in size and width; proximal bracts lanceolate, lanceolate-linear,

narrowly elliptic or rarely narrowly oblanceolate, 5-12 mm long, 0.5-1. 5 (rare outermost one subfoliar to 2.5)

mm wide, widest in proximal third (often near base), sometimes near middle or distal third; apex attenuate;

indument setose or pilose, ciliate or not; distal bracts 3-5(-8) mm long. Corolla yellow aging yellowish or red-

dish orange; corolla tube 7-12 mm; corolla limb 6-10 mm in diam.

Distribution and habitat . — Mexico (northwest, central, southern), Central America (Guatamala to central

Panama), Cuba; cultivated and escaped in Old World tropics; littoral and thorn shrubland, open pine-oak and

deciduous montane forest and woodland; disturbance openings in tropical evergreen, sclerophyll, and decidu-

ous forest and woodland; tropical savanna; 0-2500 m.

e Cooper 5892 (US). San Jose: Sidney 42, also x L. nivea

subsp. mutabilis (F); Tonduz 3377 (US); Tonduz 7 035 (LL[di]). GUATEMALA. Aha Vera Paz: Turckheim 39 (US). MEXICO. Chiapas: Ton

7048 (TEXIdi)). Jalisco: Gregory & Eiten 209 (SMU). Nayarit: Waterfall 16328, also x L. kingii (SMU[di]). Veracruz: Nee et al. 25132

subsp. glandulosissimal (MO). 5. L. insularis. CUBA. Santiago de Cuba: Britton et al 12624 (NY); Ekman 7 972 (NY); Havard 125 (NY). 6.%J

scabrida. COSTA RICA. Gnanacaste: Tonduz 13630 (LL[di]). Limon: fiminez 1903 (NY). CUBA. Havana: Ledn 1744 (NY). Matanzas: Brit-

ton et al 2 34 (NY). Santiago de Cuba: Britton et al. 12898 (NY). HONDURAS. Comayagua: Wilson 478 (NY). PANAMA. Bocas del Toro:

Peterson & Annable 7269 (MO). Codfc Gonz0kz23 (MO). Coldn: Blum & Dwyer 2119 (MO); Mill er & Miller 908 (MO). Panama: Ebinger29

camara). AUSTRALIA. New South Wales: Riding 77 (BRIT). Queensland: Hannan-Jones 73 (BRIT). 10. L. kingii. MEXICO. Hidalgo:

Carney 31 (BRlT[di]). Oaxaca; King 1178 (NY). Sinaloa: Gentry 7133 (NY). Sonora: Frye & Frye 2308 (NY). 12b. L. nivea subsp. mutabilis

(L. horrida subsp. uncertain, could also be L. horrida subsp. tMfolia). AUSTRALIA. New South Wales: Day 70 (BRIT); Day 72 (BRIT).

RWANDA. Butare: D’Arcy 8700 (MO). 20. L. strigocamara. AUSTRALIA. Queensland: Hannan-Jones 35 (BRIT). See also taxa la, Id, le

2b. Lantana horrida subsp. zanonii (R.W. Sanders) R.W. Sanders, comb. nov. Basionym: Lantana urticifolia Mill, subsp.

Shrubs erect, rounded, lax, or trailing, dense to open, the central axis well-developed to abortive, branches

ascending to arching and several or decumbent and few; twigs, peduncles and/or petioles densely setose and

stipitate-glandular. Leaf-blades ovate, ovate-elliptic, trullate, ovate-deltate, or broadly ovate, or smallest ones

subrotund, (0.5-)l-4(-6) cm long, moderately to prominently rugose, puckered between tertiary and/or sec-

ondary veins, apex acute to rounded; marginal teeth 6-20 per side, with sinuses 0.2-2.5 mm deep; adaxial

surface antrorsely strigillose to strigose-velutinous, viscidly stipitate-glandular or not, the hairs 10— 150/sq- ;

mm. Peduncles 1-4 x leaf length (usually about 2 when mature). Bract series gradually reduced in size and

width or all similar; proximal bracts lanceolate, lanceolate-linear, narrowly oblong or oblong-obovate to spatu-

late, 2.5-10 mm long, 0.5-1.5 mm wide, widest in proximal, middle, or distal third; apex rounded, obtuse or

acute; indument strigose-pilose or setose, often sessile- or stipitate-glandular, ciliate or not; distal bracts 2-4

mm long. Corolla yellow and aging yellow, orange or orange-red; corolla tube 4-10 mm; corolla limb 4-8 mm

Sanders, Taxonomy of Lantana sect. Lantana

Distribution and habitat . — Eastern Cuba, Jamaica, Hispaniola, Puerto Rico, Virgin Islands and northern

Lesser Antilles; brushland and open tropical deciduous to semi-evergreen woodland or open pine woodland

on rocky (often calcareous) slopes; 0-600 m.

When I originally described L. urticifolia subsp. zanonii, it initially appeared to differ from L. arida var.

s argentii in leaf and bract shape and some vestiture traits (Sanders 1989). However, careful examination for the

present study failed to produce consistent distinctions or geographic correlations. If L. subcordata had not

proved to be partially continuous with L. horrida var. sargentii, there would have been no need to recognize

varieties within the subspecies. However, there is overlap in bract shapes, and one gathering (Dominican Re-

public. Santiago: Liogier 13272, LL, NY), otherwise identical to L. subcordata, is an erect shrub as in var. sargen-

tii. Recognition of only two varieties within Lantana horrida subsp. zanonii has resulted in the lack of a nomi-

nate variety because autonyms exist only for infraspecific taxa that include the type of the species (ICBN Art.

26, Note 1, McNeill et al. 2007). Because, u sargentii” has priority at the varietal level, I am not free to publish the

name L. horrida var. zanonii for the variety that includes the type of L. horrida subsp. zanonii (ICBN Recom-

mendation 26A, Examplel).

2b.i. Lantana horrida subsp. zanonii var. sargentii (Moldenke) R.W. Sanders, comb. & stat. nov. Basionym: Lan-

Shrubs erect or rounded and open to dense, the central axis well-developed, branches ascending to arching

and several; twigs, peduncles and/or petioles densely setose and stipitate-glandular. Leaf-blades ovate, ovate-

elliptic, trullate, ovate-deltate, or broadly ovate, or smallest ones subrotund, (0.5-)2-4(-6) cm long, moder-

ately to strongly rugose, puckered between tertiary and/or secondary veins; apex acute to rounded; marginal

teeth 6-20 per side, with sinuses (0.5-)l-2.5 mm deep; adaxial surface with hairs 10-50/sq. mm. Peduncles

1-3 x leaf length (usually about 2 when mature). Bract series gradually reduced in size and width; proximal

bracts lanceolate, lanceolate-linear, elliptic-lanceolate, narrowly oblong, oblong-oblanceolate, or spatulate,

5-10 mm long, widest in proximal, middle, or distal third; apex rounded, obtuse or acute; distal bracts 2-4 mm

long. Corolla yellow aging orange or orange-red; corolla tube 5-10 mm.

Distribution and habitat .— Eastern Cuba, Jamaica, Hispaniola, Puerto Rico, Virgin Islands and northern

Lesser Antilles; brushland and open tropical deciduous to semi-evergreen woodland or open pine woodland

on rocky (often calcareous) slopes; 0-600 m.

See description and illustration in Sanders (1989).

Selected specimens examined: CUBA. Guantanamo: Britton 2168 (NY). Santiago de Cuba: Britton et al. 12622 (NY). DOMINICAN RE-

PUBLIC . Azua: M. /.ii a ,1/ I ,sl ,X\ II \ M 1 1 IVdernalcs: !. ' I >•%<• > > ' Ul«> Peravia: / ,o<ui » r - ' .

22025 (NY); Pelaez 202 (NY). Santiago: Jimenez 8607 (LL[dil). Santiago Rodriguez: Garcia & Pimentel 2284 (NY). JAMAICA. St. Cathe-

rine: Proctor 32628 (NY); Yunker 17476 (NY). St. Thomas: Arague-Molina & Barkley 22J4 (LLtdi)); Boars Lc 5 (BRIT). PUERTO RICO.

Guanica: Britton et al. 55 08 (NY). Lajas: Smith PR. 18 (LLldi]). VIRGIN ISLANDS. St. Thomas: Woodbury s.n. (NY). Tortola: Fishlock 45

(NY); Proctor 44899 (NY).

Presumed hybrids with: 6. L. scabrida. JAMAICA. St. Andrew: Baars Lc7 (BRIT). St. Elizabeth: Baars Lc8 (BRIT). LESSER ANTIL-

LES- St. Maarten: Krauss 1670 (LLtdi]). 7. L. splendens. CUBA. Gua nt a n a m o: Britton 2216, identification uncertain (NY). 20. L. strigoca-

mara. DOMINICAN REPUBLIC: Santana: Zanoni 17698 (NY). See also taxa la and lc.

2b.ii. Lantana horrida subsp. zanonii var. subcordata (Urb.) R.W. Sanders, comb. & stat. nov. Basionym: Lantana

subccmlcii.Hrl, Mmh Am, II |l. rh 1 : V,1 1»12 T.n LH>MI\|, AN Kl PI Bl K near \imi.i>y \t„<mbwgh id (Otyfe: K. bar-

code K000470761Idi!]; iscrrrrc: P[di!]).

Shrubs trailing or sprawling, the central axis abortive or weakly developed, branches decumbent, few; twigs,

peduncles and often petioles moderately puberulent, setose, or glabrescent, at least peduncles usually stipitate-

glandular. Leaf-blades ovate, deltate, or ovate-oblong, 0.5-2 cm long, strongly rugose, puckered between veins

and veinlets; apex abruptly rounded or acute; marginal teeth 8-15 per side, with sinuses 0.2-0.8 mm deep;

adaxial surface with hairs 50-150/sq. mm. Peduncles 2-4 x leaf length. Bract series similar in size or distal

series only partially, gradually reduced; proximal bracts oblong or oblong-obovate to -oblanceolate, 2.5-7 mm

long, widest just above middle to distal third; apex rounded or obtuse, often reflexed; distal bracts 2-3 mm

long. Corolla yellow aging to dark yellow or orange-yellow; corolla tube 4-8 mm.

Distribution and habitat . — San Jose de las Matas region of Cordillera Central, Dominican Republic, His-

paniola; open pine and deciduous montane forest; 100-600 m.

See further discussion and illustration in Sanders (1989).

2c. Lantana horrida subsp. tiliifolia (Cham.) R.W. Sanders, c

(WILLD 11502) (lectotype: B-WlLLD[di!]). Remaining syntypes: BRAZIL. 1

: PERU: Sep 1829, Poeppig 1375 (i Ft iotype: G[di!|!; .

Lantana tiliifolia Cham. var. glandulosa Schauer, FI. Bras. [Martius] 9:257.1851. Lantana tiliifolia Cham. f. glandulosa (Schauer) R. Fern, ;

Blanchet20 (G-DC[di!]); BRAZIL. Bahia: 1

j-DC[di!], MPU[di!], P[di!]).

VENEZUELA. N

a: Campo Ella, 14 Aug 1938, Han-

bury-Tracy 31 (holotype: K[di!];

Shrubs erect or rounded and open, occasionally lax and subscandent, the central axis well-developed, promi-

nent, branches ascending and several, occasionaly clambering and few; twigs, peduncles and often petioles

moderately to densely setose, or mixed setose and stipitate-glandular, or predominantly stipitate-glandular.

Leaf-blades ovate to broadly ovate or broadly elliptic, (l-)3-9(-12) cm long, moderately to weakly rugose,

puckered between tertiary veins, apex usually abruptly, briefly acuminate (triangular tip ca. 5 mm) to acute,

occasionally obtuse or rounded; marginal teeth 15-35(-45) per side, with sinuses 0.3-1.2(-1.5) mm deep; ad-

axial surface antrorsely strigillose to strigose-velutinous and often viscidly stipitate-glandular, the hairs (5-

)10-30(-50)/sq. mm. Peduncles (0.5-)0.8-2 x leaf length (usually about equalling to almost twice as long

when mature). Bract series all small or proximal 2 or 3 series longer with distal series abruptly shortened;

proximal bracts elliptic, narrowly ovate, oblong, or oblong-obovate, 2-5(-10) mm long, (0.8-)1.2-3 (rare out-

middle; apex acute to obtuse, sometimes briefly, abruptly acuminate or, if bract is over 5 mm long, attenuate;

indument thinly pilose, stipitate-glandular or not, ± ciliate; distal bracts 2-4 mm long. Corolla yellow to orf|

aging reddish orange or dark red; corolla tube 7-12 mm; corolla limb 6-10 mm in diam.

Distribution and habitat .— The Guianas, Venezuela, Colombia, Ecuador, Peru, Chile, Bolivia, Paraguay,

northern Argentina, and southeastern to eastern Brazil; cultivated and escaped in Old World tropics; tropical

savanna with gallery forest, montane humid forest, and disturbed successional woodland, shrubland and

grassland; 150-2400 m.

Andean plants are predominantly stipitate-glandular. Otherwise, I have been unable to discern any geo-

graphic or ecological patterns separating the eglandular, mixed eglandular-glandular, and predominantly

stipitate-glandular plants, especially in the Brazilian Planalto. Extensive field work is needed

Selected specimens examined: ARGENTINA. Buenos Aires: Torres et al. 2154 (MO). Cordoba: Botta & Miami 3J4 (MO). BRAZIL. Bahia:

Harley 18478 (LLfdil); Mori et al. 10635 (LL[di]); Smith Braz. 33 (LL). COLOMBIA. Boyac* Cuatrecasas 1134 (F); Cuatrecasas 1920 (F).

Cauca: Cuatrecasas 19542 (F). Choco: LeOn 312 (MO). Norte de Santander: Cuatrecasas & Rodriguez 27954 (F). Santander Cuatrecasas &

Garcia 9855 (F); Luteyn et al. 7619 (F). Tolima: Cuatrecasas 10515 (F). PERU. Cusco: Galiano et al. 6275 (MO); Valenzuela et al. 4620 (MO).

Journal of the Botanical Research Institute of Texas 6(2)

on veins but usually not on non-innervated t

ining hemispheric, prolate-globose in fruit.

, setiform, 0.7-2.0 mm. Inflorescences ar-

4. Lantana hirsuta M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11:326. 1844. Type: MEXICO. Va

Shrubs erect, rounded, or subscandent; stems 0.7-3 m; branches ascending and several to clambering and few;

twigs, peduncles and often petioles moderately to densely setose, the hairs (0.8-)l-2.5 mm, mostly all the same

length. Leaf-blades broadly ovate, ovate, or ovate-elliptic, rarely lanceolate, (2-)4-12 cm long, the length 1-2J

x width, not nigrescent, membranous to papery, pinninerved; base usually rounded to truncate, sometimes

marginal teeth 10-35(-40) per side, acute to rounded, spreading to appressed, then sometimes with tips re-

curved, with sinuses 0.3-3 mm deep; adaxial surface dull, setose to villous, the hairs occurring on veins and

intervening tissue, longer ones 1-1.5 mm or more, 1-40/sq. mm, not noticeably vitreous-pustulate, the circular

bases of the setae ca. 0.1-0.2 mm in diam.; abaxial surface slightly lighter or duller green than adaxial surface, ■

setose, with the setae restricted to midrib and veins, (0.5-)0.7-1.5 mm or more, usually without shorter hairs,

0.5-80/sq. mm. Inflorescences occasionally 2 per leaf axil, remaining hemispheric; peduncles 0.3-K-2) x leaf

othermost series often dilated in distal third and widest there or equalling broadest proximal portion, with 3

veins from the base, appressed or spreading, persisting or not; apex attenuate or acute or rarely rounded at very

tip; indument setose or pilose, ciliate or not, the longest hairs 0.3-1.5 mm. Corolla yellow to or aging reddish

orange; corolla tube 8-12

Distribution and habi

tropics; openings in tropi<

cal semi-evergreen fore:

na with gallery forest, a:

id northern Argentina; cultivated and escaped in Old World |

t and montane evergreen forest on poor soils, open pine-oak

eas of dense woodland, shrubland, and grassland; 0-2000(- 1

4a. Lantana hirsuta subsp. hirsuta.

Lantana scorta Moldenke, Publ. Carnegie Inst. Wash. No. 522 (Bot. Maya Area): 161. 1940. Type: MEXICO. QuerEtaro: San Juan del Rfe

17 Aug 1905, Roseetal. 9520 (holotype: NY!; isotype: MEXUldi!]).

Leaf-blades broadly ovate, ovate, or rarely lanceolate or ovate-elliptic; apex acuminate (abruptly contracted to

triangular tip 3-8(-15) mm long), acute or rarely obtuse or rounded (triangular tip lacking); marginal teeth

10-25(-40) per side, with sinuses (0.7-)l-3 mm deep; adaxial surface with the setae l-7(-15)/sq. mm; abaxial

surface with the setae erect and usually rigidly straight, 0.5-7(-I5)/sq. mm. Peduncles (3-)6-14 cm, OJ^H

x leaf length (usually about equalling when mature). Proximal bracts with longest hairs 0.5-1.5 mm.

Distribution and habitat . — Mountains and coastal plains of Mexico (frequent in eastern Mexico but collec-

tions are known from a few disjunct localities in western Mexico from Guerrero to Baja California), Central

America, and extreme northwest Colombia; cultivated and escaped in Old World tropics; open pine-oak wood-

land, openings in semi-evergreen tropical forest and brushland, thickets, and grasslands; (0-)1000-1600 &.

The few collections from western Mexico may represent another infraspecific taxon, as they tend to have

smaller leaf-blades and denser trichomes. Further work on this variation is needed.

Selected specimens examined: GUATEMALA. Pettn. Contreras 42 02421); Contreras 439 (LL[2]>. Quezaltenanjjo: Skutch 1347 (NY,

Sanders, Taxonomy of Lantana sect. Lantana

& Hernandez 596 (NY); Ortega et al. 19650 (NY); Sharp 44197 (TENN); Vasquez 488 (NY). U.S.A. Alabama. Mobile Co.: cult., Mohr s.n. (US).

i. GUATEMALA. Retalhuleu: M axon et al. 3523 (US). MEXICO. Chiapas: Martinez 8949

x L. kingii (BRIT[dil). Quintana Roo: Cabrera & Cabrera 4046 (TEX[dil). PANAMA. Bocas

il. 898 (MO). Chiriqui: Churchill & Churchill 6136 (MO); Lewis et al 616 (MO); Tyson 897 (MO). Kuna Yala (San Bias):

McDonagh etal. 167 (MO). Panama: Knapp et al. 3298 (MO). 9i-cvx20. L. CaUowiana Hybrid Group cultivars (L. depressa-tetraploid x

siri B oiamara) TANZANIA. Kilimanjaro: Ml, , UJ I'l. l I 10. I . kingii. MEXK O. Oaxaca: c : n-iiXalicrnaiively x / scabrida

or L. CaUowiana Hybrid Group cv. (TEX[dil). Puebla: Arsine s.n. 3 Nov 1908 (US). San Luis Potosfc Davis 24 4 (NY); Krai 2 4812 (VDB); Parry

L. hirsuta subsp. amazcmica). AUSTRALIA. Queensland: Day 56 (BRIT); Day 87

)UTH AFRICA. Limpopo: Day 6 (BRIT). 18. L. urticoides. MEXICO. Nuevo

subsp. uncertain, could also be L. hirsuta subsp. amazonica). AUSTRALIA.

)Sp. nov. (Fig. 1). Type: BR/

IBGE.NY!).

Leaf-blades broadly ovate-elliptic to ovate, oblong-ovate, or elliptic; apex acuminate (abruptly contracted to

triangular tip (5-)10-15 mm long), acute or rarely obtuse or rounded (triangular tip lacking); marginal teeth

(20-)25-35 per side, with sinuses 0.3-1.5 mm deep; adaxial surface with the setae (3-)10-40/sq. mm; abaxial

surface with the setae erect and usually arching or sinuate, 10-80/sq. mm. Peduncles 2-5(-8) cm, 0.3-0.5(-l)

x leaf length. Proximal bracts with longest hairs 0.3-1 mm.

Distribution and habitat . — Brazil (eastern Amazon Basin and Planalto), Paraguay, northern Argentina,

Bolivia, Peru, Ecuador, and Colombia to western Venezuela; cultivated and escaped in Old World tropics;

openings in tropical semi-evergreen forest and montane evergreen forest on poor soils, tropical savanna with

gallery forest and areas of dense woodland, shrubland, and grassland; 0-2000(-3000) m.

Schauer (1847, 1851 [t. 42]) recognized Lantana hirsuta subsp. amazonica as a species but misapplied the

name L. mista L. to it (see hybrid synonymy lfx4).

ularis Moldenke, Caribl

>;16. 1940. Type: JAMAICA: [1

Shrubs erect, apparently pyramidal and open; stems 1-3 m; branches ascending and several; twigs, peduncles

and often petioles spasely setose but viscid with conspicuous, dense stalked glands, the setae 1.2-2 mm, stipi-

late glands, 0.3-0.7 mm. Leaf-blades ovate-triangular to ovate-elliptic or lanceolate, 4-10 cm long, the length

Journal of the Botanical Research Institute of Texas 6(2)

423

a and Cuba; openings or disturbed areas in cloud c

Presumed hybrids with: 6. L. scabiida. CUBA. Sancti Spiritus: Alain/

C. Lantana sect. Lantana series Strigosae R.W. Sanders, ser.

Adaxial leaf-surfaces strigose, strigillose, or scabrid, the hail

abaxial leaf-surfaces strigose, usually sparsely so, the hairs o

innervated tissue, strigiform, 0.03-1 mm. Inflorescences an

bose in fruit (continuing to initiate flowers somewhat in L. n

ciduous in lower portion leaving it more or less bare w

>tent bases) about ]

of flowers/fruits active).

c Sagua de Tanamo, 8 Apr 1987, Mendez

3373 (holotype: HIPC[di!]).

Shrubs rounded and open, lax or subscandent; stems 0.5-6 m; branches ascending to clambering, often few;

twigs, peduncles and often petioles thinly to moderately strigose or setose, the hairs 0.2-1 mm. Leaf-blades

ovate to elliptic-lanceolate or lance-oblong, (2-)5-15 cm long, the length (1.5-)1.7-2.5(-3) x width, nigrescent,

papery to subcoriaceous, triplinerved; base attenuately to abruptly tapering onto petiole from middle or just

below middle; apex acuminate, acute, obtuse, occasionally rounded; marginal teeth (12-)17-25(-35) per side,

obtuse, rounded, or acute, spreading to appressed, then sometimes with tips recurved, with sinuses 0.5-1.5

mm deep; adaxial surface lustrous, thinly strigose or scabrous, the hairs occurring on veins and center of are-

oles, 0.1-0.6(-1.2) mm (longest usually 0.2-0.4 mm, except in “scandens” morph where 0.6-1.2 mm), (4-)6-

12/sq. mm, sometimes with conspicuous vitreous or whitened pustulate bases 0.1-0.3 mm in diam.; abaxial

surface whitish or pale green but not glaucous, antrorsely strigose-scabrous, with the strigae scattered on veins

and veinlets, (0.03-)0.1-0.5(-1) mm (longest mostly 0.3-0.5 mm, except in “scandens” morph where 0.5-1

mm), (4-)6-12(-25)/mm sq. (“scandens” morph 0-5)/sq. mm). Inflorescences remaining hemispheric; pe-

duncles 0.5-1 x leaf length. Proximal bracts narrowly lanceolate, lance-elliptic or -oblong (including those

with slight constriction in proximal third; occasionally 1 or 2 outermost bracts subfoliar or narrowly spatu-

late), 4-8 mm long, 1—2 mm wide, widest at very base, in proximal third, or in middle third (then often equally

wide at base), with 3 veins from the base, appressed or spreading, usually deciduous after flowering; apex at-

tenuate or acuminate; indument sparsely strigose, somewhat or not ciliate, the longest hairs 0.3— 0.5(— 1) mm.

Corolla yellow to or aging reddish orange; corolla tube 7-12 mm.

Distribution and habitat .— West Indies (Cuba, Jamaica, possibly Hispaniola, Puerto Rico, possibly the

Virgin Islands, and the northern Lesser Antilles), eastern coastal Mexico (Tamaulipas southward) with local-

ized disjunction in southwestern Michoacan (long-haired, scandent morph). Central America, and Caribbean

coast and slopes of Colombia, Venezuela, and the Guianas; disturbance openings, savanna and man-made

grassland in tropical dry to humid forest; 0-1600 m.

Lantana scabrida is replaced in upland and western Mexico by L. kingii, but the two are apparently sympat-

ric in Tamaulipas. Field work is needed to determine the status of the isolated scandent morph in Michoacan.

Sanders (1987b) reported the chromosome number (2n = 44) of Lantana scabrida in the Luquillo Mountains

of Puerto Rico ( Sanders 1510 ) as L. camara due to misapplication of the name and confusion with L. strigocamara.

Sanders, Taxonomy of Lantana sect. Lantana

tensely lustrous, strigose-glabrescent but smooth, the hairs occurring on veins (scattered) and center of are-

oies, 0.2-0.4(-0.8) mm, 0-2/sq. mm, flaccid and strongly appressed to surface (unique in this species), often

deciduous not pustulate based; abaxial surface whitish or gray-green (but not glaucous), weakly strigose and

nearly glabrous, with the strigae scattered on veins and veinlets, 0.1-0.6 mm, 0-4/sq. mm. Inflorescences re-

maining hemispheric; peduncles Vfe-VS x leaf length. Proximal bracts oblanceolate to narrowly oblong to nar-

rowly lanceolate, (4-)6-10 mm long, 1-1.7 mm wide, widest in proximal third to distal third, with 3 veins from

the base, appressed or spreading, deciduous after flowering; apex attenuate; indument glabrescent, not cihate,

the longest hairs 5 0.5 mm. Corolla yellow to or aging reddish orange; corolla tube 7-10 mm.

Distribution and habitat.-Ce ntral Lesser Antilles (Dominica, Martinique, probably Saint Lucia); sunny

slopes in borders and openings of montane rainforest; 400-1000 m.

See discussion in Sanders (1987c).

Sheeted specimens examined: LESSER ANTILLES. Dominica: Lee 9 (NY); Lloyd 201 (NY); Smith 10216 (SMU). Martinique: Bailey & Bailey

ANTILLES. Dominica: Hill 23959, altera

9. Lantana depressa Small, Bui

v York Bot. Gard. 3:436. 1905. 1

Shrubs trailing to erect, dense to ± open; stems 0.1-3 m; branche

several to numerous; twigs, peduncles and often petioles thinly setose or strigose-s

Leaf-blades ovate-elliptic to elliptic, l-6(-8) cm long, the length 1.7-2.3(-3) x width, induplicately curved at

maturity (unique to this species), nigrescent, papery, triplinerved; base obtuse or acute, tapering onto petiole

from middle or just below middle; apex abruptly tapered, obtuse or acute; marginal teeth 3-15 per side,

rounded to acute, often appressed, with sinuses 0.5-1.5 mm deep; adaxial surface lustrous, antrorsely strigil-

lose to strigose the hairs occurring on veins and center of areoles, 0.1-0.7(-l) mm, 2-8/sq.mm, not noticeably

vitreous-pustulate, the circular bases of the strigae ca. 0.1-03 mm in diam.; abaxial surface slightly lighter or

duller green, antrorsely strigose-scabrous, with the strigae scattered on veins and veinlets, longest ones 0.5-1

mm, 0.5-8/sq. mm. Inflorescences remaining hemispheric; peduncles 0.7-2 x leaf length. Proximal bracts

elliptic-lanceolate, 4-7 mm long, 0.5-1.5 mm wide, widest at middle or just below, with3veins from the base,

appressed or spreading, persisting and reflexed from base in fruit; apex acute to attenuate, indument strigose,

ciliate or not, the longest hairs 0.1-1 mm. Corolla yellow aging to a dark yellow or dull, pale orange; corolla

tube 5-11 mm.

Distribution and habitat .— Peninsular Florida; limestone pinelands, wet prairies, and dunes; 0-50 m.

See more complete discussion in Sanders (1987a) and Maschinski et al. (2010) and illustrations in Sandeis

(1987a).

i. Lantana depressa var. depressa

ttoon. Plant Buyer’s Guide, ed. 6, 1

ssa Small var. depressa (see Sanders 2001, [

V. Long, Rhodora 72:34. 1970. Type U.S.A. Florida. E

426

Shrubs low mounded, dense, 0. 1-0.3 (spreading to 1) m, the central axis abortive or hardly developed; branch-

es prostrate or decumbent, twigs, peduncles and often petioles with hairs 0.5-1 .5 mm. Leaf-blades 1-3 cm

long (to 4 or 5 cm in some cultivars). Corolla with tube 5-9 mm; corolla limb 5-8 mm in diam.

Distribution and habitat.— Peninsular Florida (Miami Ridge); cultivated and escaped in tropics and sub-

tropics worldwide; limestone pinelands; 0-25 m.

Because of its drought tolerance, compact habit, and profuse flowering, Lantana depressa var. depressa has

been cultivated widely since the 1950s. A tetrploid cultivar and L. strigocamara apparently are the parents of

the currently popular Callowiana Hybrid Group cultivars, which have the floral colors of L. strigocamara and

are cultivated worldwide and escaped pantropically (see Sanders 2001, specimen citations below, and 9i-cvx20

in the section on hybrid synonymy).

Selected specimens examined: AUSTRALIA. Queensland: cult., McAndrew MJH-647 (BRIT). U.S.A. Florida. Dade Co.: Deam 60894 (NY);

Demaree 10208 (SMU); Krai 53943 (VDB); Krai 53964 (VDB); Krai 66236 (NY, VDB); Krai 70742 (VDB); Small et at. 3482 (NY); Traverse 646

e: Moldenke 797 (NY),

i: Moldenke 19862 (SMU). GUATEMA1

(BRIT). MEXICO. Jalisco: Jimenez 357 (NY). Sinaloa: Hutchison 2544 (NY). Veracrnz: Day 15 (BRIT). TANZANIA. Kilimanjaro: Mlangwa

352, possibly with genes of L. hirsute (BRIT[di]). Tanga Sallu 248 (BRIT). TRINIDAD & TOB,

U.S.A. Florida. De Soto Co.: Krai 5 7309 (VDB). Lee Co.: Krai 11998 , alternatively a backcross in

Co.: Carter & Carter 13218 (FTG). Louisiana. Iberia Par.: Webb 4764 (VDB). Massachusetts: ci

Co.: cult., Dunn 13828 (BRIT). South Carolina. Beaufort Co.: Leonard & Radford 2743 p.p. (VDB). Tennessee. Davidson Co.: cult., Hackney

s.n. 10 Oct 1990 (VDB). Texas. Blanco Co.: cult., Sanders 5470 (BRIT). Dallas Co.: cult., Shinners 8526 (SMU); cult., Skinners 29844 , probably

‘Cream Carpet’ (SMU); cult., Wansbrough 251 (SMU). Galveston Co.: cult.. Traverse 2515 (BRIT). Harris Co.: c

Traverse 2401 (BRIT). Hidalgo Co.: cult., Green/ield5(BRIT). Montg

Presumed further hybrids of I9i-cvx20] with: 10. L. kingii. MEXICO. Guerrero: Guerra 36 (LL[di]). I

AUSTRALIA. Queensland: Day 86 (BRIT); Horrocks 55 (BRIT). TRINIDAD & TOBAGO. 5

planaltensis. BRAZIL. Bahia: Ddbereiner & Tokarma 1490 (LL[di]). PARAGUAY. Central: Perez 890 (MO). 1

Anderson Co.: cult., Couch 73 (SMU). Cameron Co.: Traverse 1030 (BRIT). 20. 1

(SMUjdi]). Leon Co.: Bratcher 57, alternatively x L. depressa var. depressa (BRIT);

(VDB). Seminole Co.: SchaUert378 (BRITldi]). See also taxa la, lb, Id, le, If, 2a, 4i

a and 20 and section on hybrid synonymy: 9i-cvx20. |

9ii. Lantana depressa var. floridana (Moldenke) R.W. Sanders, Syst. Bot. 12:55. 1987. Basionym: Lantai

Shrubs rounded, open, 0.4-1 m, the central axis ± developed, but not prominent; branches arching or ascend-

ing; twigs, peduncles and often petioles with hairs 0.5-1 mm. Leaf-blades 3-6(-8) cm long. Corolla with tube

7-11 mm, corolla limb 8-10 mm in diam.

Distribution and habitat. — Peninsular Florida; Atlantic barrier dunes and interior sand ridges, stabilized

and relictual dunes; 0-50 m.

Selected specimens examined: U.S.A. Florida. Brevard Co.: Moldenke 2 18 (NY). Martin Co.: Krai 15385 (VDB); Krai 57137 (NY, VDB). N*

Beach Co.: Small 2134 (NY). St. Johns Co.: Morton 4508 (SMU); Small 2313 (NY). Volusia Co.: Ray et al. 10817 (SMU). Geogia: cult., Bol**

s.n. (NY).

Presumed hybrids with: 20. L. strigocamara. U.S.A. Florida. Duval Co.: Curtiss 1968 (NY[2]). Lake Co.: Small 8666 (NY[21). Vd*

ria Co.: Krai 18406 (VDB); Whetstone 145 18 (VDB). See also section on hybrid synonymy: 9iix20.

9iii. Lantana depressa var. sanibelensis R.W. Sanders, Syst. Bot. 12:55. 1987. Type: u.SA. Florida. Lee Co.: Sanibd*

land, 11 May 1954, Cooley 2674 (holoiype. GH!; isotypes: USF[2]!).

Shrubs erect and pyramidal, proximally dense, distally open,

nent; proximal branches decumbent, distal ones ascending; i

1-1.5 mm. Leaf-blades 3-6 cm long. Corolla with tube 7-10 v

Distribution and habitat.— Peninsular Florida; wet limesto

barrier islands; 0-25 m.

-3 m, the central axis well-developed, pro®*-

vigs, peduncles and often petioles with hairs

427

ock 268 (NY).

Presumed hybrids with: 20. L. strigc

Brumbach 8182 (NY); Brumbach 8283 (NY, U

; Brumbach 9058 (VDB).

f 7737 (FTG, NY); Sheehan s n. 7 Mar 1919 (NY). I

10. Lantana kingii Moldenke, Phytologia 8:161. 1962. Type: MEXICO. Oaxaca: Isthmus of Tehuantepec, Niltepec, I7Jul 1959,

IQng 1775 (holotwe: TEX!; isotype: US!).

Shrubs erect or rounded, open; stems 0.5-2(-3) m; branches ascending and several; twigs, peduncles and often

petioles glabrescent with scattered antrorse hairs, the hairs ca. 03-0.7 mm. Leaf-blades ovate, ovate-elliptic or

ovate-triangular (rarely, especially if less than 2 cm long, obovate to rotund), 1-8 cm long, the length (0.8-)1.2-

attenuately tapering onto petiole from middle or just below middle to abruptly contracted and broadly cuneate,

sometimes forming a short narrow petiolar wing; apex acute or acuminate (or abruptly to broadly rounded);

marginal teeth (3-)6-15(-25) per side, obtuse or rounded, spreading to appressed, then sometimes with tips

recurved, with sinuses 1-2.5 mm deep; adaxial surface lustrous, scabrous, the hairs occurring on veins and

intervening tissue (sometimes just center of areoles), 0.1-0.5 mm, 3-7(-12)/sq. mm, mostly deciduous leaving

the noticeably vitreous-pustulate circular bases, these 0.2-0.5 mm in diam.; abaxial surface whitish or pale

green but not glaucous, antrorsely strigose-scabrous to nearly glabrous, with the strigae scattered on veins and

veinlets, 0.1-0.7 (longest ones usually 0.3-0.4) mm, l-8(-12)/sq. mm. Inflorescences remaining hemispheric;

peduncles about 0.5 to 1.5 x leaf length. Proximal bracts spatulate, oblanceolate, or oblong-oblanceolate, oc-

casionally oblong-elliptic to broadly elliptic, (3.5-)5-10 mm long, 1.2-3 mm wide, widest in distal half (often

just above middle), occasionally at or just below middle, with 3 veins from the base, appressed or spreading,

persisting and reflexed from base in fruit; apex obtuse to acute; indument strigose, hardly ciliate, the longest

hairs 5 0.5 mm. Corolla yellow to orange aging orange to orange-red; corolla tube 6-10 mm.

Distribution and habitat. — Mexico (central highlands to eastern slopes of the Sierra Madre Oriental, Pa-

cific slope, and Isthmus of Tehuantepec) to northern Central America (only to central Guatemala?); thorn for-

est and schrubland; 0-2000 m.

Selected specimens examined: MEXICO. Coahuila: Johnston 9325 (LL[di]); Henrickson 18926 (NY); Waterfall 16661 (SMU). Colima: Grego-

ry & Eiten 334 (BRIT). Guerttero: Flyr 623 (SMU). Guerrero: Mocktord & Rowell 2790 (SMU). Michoacan: Turner 2024 (BRIT, SMU). Neuvo

Le6tt: Garcia 10 (SMU); Pennell 16860 (NY); Waterfall 13187 (SMU); Waterfall 15312 (SMU[di]). Oaxaca: King 1328 (NY); King 1464 (NY);

King 1598 (NY); Purpus 7306 (NY). Puebla: Chiang et al. F-2610 (TEX); Davis 211 (NY); Martinez 21705 (TEXldi]). Sinaloa: Gentry 11454

(LL); Palmer 1511 (NY); Rose et al. 13366 (NY). Sonora: Wiggins & Rollins 138 (NY). Tamaulipas: Smith Mex. 94 (LLldil).

Presumed hybrids with: 2a/4a. L. horrida subsp. horrida or L. hirsuta subsp. hirsuta. MEXICO. Hidalgo: Cawrey3 (BRIT). 18. L.

urticoides. MEXICO. Coahuila: Henrickson 11352 (LL[dil); Muller 3069 (LL(dil); Reveal et al. 2604 (NY): Wehbe 052 ( TEX[di] ): Wy nd &

Mueller 88 (NY). Neuvo Le6n. Frye & Frye 2447 (NY); Meyer & Rogers 2686 (NY); Pringle 11670 (SMU). Tamaulipas: Meyer & Rogers 2499

(NY); Stanford ctal. 2302A (NY). 20. L. strigocamara. MEXICO. Tamaulipas: Krai 24799, alternatively x L. scabrida (VDB). Seealso taxa le,

2a, 4a, and 9i (as cv x 20) and section on hybrid synonymy: lexlO, Iex2axl0, lexl0/20, 2axl0, 4axl0, 10xla/le?, and 10x2a/4a?

11. Lantana ovatifolia Britton, Bull. New York Bot. Gard. 4:123. 1905. Type: BAHAMA ARCHIPELAGO. Grand Bahama:

Feb 1905, Britton & Millspaugh 2450 (holotype: NY!; botype: F!).

Shrubs lax and trailing, sparse; stems 0.3-1 m; branches prostrate, few; twigs, peduncles and often petioles

moderately strigose, the hairs 0.2-1.5 mm. Leaf-blades ovate to ovate-elliptic, 2-6 cm long, the length 1.2-1.7

* width, not nigrescent, papery to subcoriaceous, pinninerved; base rounded to tapering onto petiole mostly

from proximal third; apex acute to obtuse or rounded; marginal teeth 8-18 per side, rounded to acute, often

appressed, with sinuses 0.7-1.5 mm deep; adaxial surface lustrous, antrorsely strigose to scabrous (due to loss

of deciduous longer hairs), the hairs occurring on veins and intervening tissue (sometimes just center of are-

oles), 0.2-1 mm, 2-7/sq. mm, noticeably vitreous-pustulate, the circular bases 0.3-0.5 mm in diam.; abaxial

surface slightly lighter or duller green than adaxial surface, antrorsely strigose-scabrous to nearly glabrous,

with the strigae scattered on veins and veinlets, 0.2-1 (longest ones usually 0.4-0.8) mm, 2-7/sq. mm. Inflo-

rescences remaining hemispheric; peduncles about 1—1.5 x leaf length. Proximal bracts spatulate or oblan-

ceolate, 6-12 mm long, 1-3 mm wide, widest in distal half, with 3 veins from the base, appressed or spreading,

428

Journal of the Botanical Research Institute of Texas 6(2)

persisting and reflexed from base in fruit; apex obtuse to acute; indument strigose, hardly ciliate, the longest

hairs £ 0.6 mm. Corolla yellow aging to a dark yellow or yellow-orange; corolla tube 6-10 mm.

Distribution and habitat.— Northern Bahama Archipelago; limestone pinelands with open or low shrubby

understory; 0-25 m.

In certain respects Lantana ovatifolia is rather similar to L. kingii. However, as a narrow endemic with a

narrow range of variation, it is best kept as a distinct species. This is further supported by its distribution,

which is oddly disjunct from that of L. kingii. See Sanders (1987a) for detailed discussion and illustration.

Selected specimens examined: BAHAMA ARCHIPELAGO. Grand Bahama: Brace 3686 (NY); Correll & Popcnoe 45402 (FTG, NY, SMU);

Cone II &■ Krai 42892 (FTG, VDB); Correll & Krai 42946 (FTG, VDB).

Shrubs lax, rounded and open, sometimes forming treelets; stems 0.5-4 m; branches ascending or divaricate,

several to numerous; twigs, peduncles and often petioles thinly to moderately strigillose, setulose, or scabridu-

lous, the hairs 0.03-0.4(-0.7) mm. Leaf-blades ovate, lanceolate or elliptic, 3-12 cm long, the length some-

times of those subtending inflorescences distinctly reduced (unique to this species), (1.5-)1.7-3(-3.6) x width,

nigrescent, membranous to papery, triplinerved; base attenuately tapering onto petiole from widest point or

acute; marginal teeth (13-)18-40 per side, obtuse, rounded, or acute, usually appressed, sometimes spreading

at tip, with sinuses 0.2-1 mm deep; adaxial surface lustrous, antrorsely strigose to scabrous or scabridulous,

the hairs occurring on veins and intervening tissue, 0.03-0.5 mm, 10-60/sq. mm, often with conspicuous

vitreous or whitened pustulate bases 0.1-0.3 mm diam.; abaxial surface slightly lighter or duller green than

adaxial surface, antrorsely strigillose to strigose-scabrous, with the strigae scattered on veins and veinlets and

sometimes on intervening tissue, 0.03-0.4(-0.7) mm, 10-60/sq. mm. Inflorescences sometimes 2 per leaf

axil, hemispheric but receptacle often elongating by slight separation or prolonged initiation of nodes and be-

coming naked below the hemispheric flower cluster at apex; peduncles 0.5-1.2 x leaf length. Proximal bracts

narrowly triangular, lanceolate or linear-lanceolate, occasionally oblanceolate, 2.5-7(-10) mm long, 0.6-13

mm wide, widest at very base, proximal third, or occasionally distal third, with 3 veins from the base, ap-

pressed or spreading, usually deciduous after flowering; apex acute, attenuate, or subulate; indument strigil-

lose-scabridulous, hardly ciliate, the longest hairs 5 0.5 mm. Corolla white aging white, pale pink or light blue,

or opening pink, cream or yellow aging cream, yellow or orange infused with purple, pale yellow throat usu-

ally developed and fading with age; corolla tube 7-12 mm.

Distribution and habitat. — Eastern and southeastern Brazil; cultivated world-wide, escaped pantropically

understory and disturbance openings and man-made grasslands in tropical humid forest, occasionally in dry

forest; 0-1500 m.

Sanders, Taxonomy of Lantana sect. Lantana

Leaf-blades with the indument of the adaxial surface composed of strigae and rough points, 0.03-0.3 mm;

indument of the abaxial surface composed only of strigae 0.03-0.3 mm, never with short, straight fililform

prolonged initiation of nodes and becoming naked below the hemispheric flower cluster at apex. Corolla white

aging white, bluish or pale pink, or pink aging light purple.

Distribution and habitat . — Eastern and southeastern Brazil; cultivated world-wide, sometimes escaped

pantropically; understory and disturbance openings and man-made grasslands in tropical humid forest, oc-

casionally in dry forest; 0-1500 m.

Many of the native collections have narrowly elliptic leaf-blades. However, other native collections vary

toward having the more ovate or lanceolate blades typically seen in cultivated plants of the species (e.g., the

type specimens of Lantana nivea and L. triplinervia). There is a tendency, especially in the collections from Ba-

hia, for a marked reduction in size of leaves subtending the inflorescences, resulting in a corymb-like arrange-

Selected specimens examined: AUSTRALIA. Queensland: McAndrew 32 (BRIT). BRAZIL. Bahia: Mori et al. 10266 (LL); Sieber s.n. 1826

(BR[di]); Silva58360 (NY, US). Distrito Federal: Duarte & Pereira 4740 (US). Minas Gerais: Hatschbach 31331 (VS); Irwin 2112 (NY, US). Rio

de Janeiro: Araujo 3034 (NY); Brack 24153 (NY); Carauta et al. (Herb. 18132) (LL); Nee 3389 (US); Pereira et al. 4383 (NY). MEXICO. Distrito

Federal: cult,. Bonpland s.n. (P[2di]). SRI LANKA. North Central: cult., Moldenke et al 28233 (US).

(NY); Harley 21658 (NY); Harley 21927 (NY); Hatschbach 46372 (LL[dil); Irwin et al. 27777 (LL[di]). 17. L. planaltensis. BRAZIL. Bahia:

Harley et al. 21617 (US). Minas Gerais: Anderson 9142 (F, US); Mexia 5436 (F, NY). Rio de Janeiro: Segadas-Vianna et al. 593 (SMU). 20. L.

12b. Lantana nivea subsp. mutabilis (Hook.) R.W. Sanders, Sida 22:395. 2006. Bason™: Lantana nivea Vent. var.

Leaf-blades with the indument of the adaxial surface composed of strigae, 0.05-0.5 mm; indument of the ab-

axial surface composed of strigae, 0.05-0.7 (longest ones mostly 0.4-0.5) mm, often with short, straight filil-

form hairs to 0.3 mm mixed in but not dominant or codominant. Inflorescences remaining hemispheric; re-

ceptacle rarely elongating and becoming naked below. Corolla opening creamy white or pink, aging cream,

yellow or orange infused with purple or opening yellow aging purple.

Distribution and habitat. — Probably of cultivated hybrid origin, cultivated world-wide and escaped pan-

tropically (especially in Australia), but also collections from southeastern Brazil apparently of natural hybrid

origin; understory and disturbance openings and man-made grasslands in tropical humid forest, occasionally

in dry forest; 0-1500 m.

Selected specimens examined: AUSTRALIA. Queensland: Day 53 (BRIT). BRAZIL. Goias: Irwin et aL 25204 (NY). Rio de Janeiro: Carauta

& Araujo 2244 (LL[di]); G6es 64 (NY); Sarahyba 80 (NY); Segadas-Vianna 4053 (SMU). BURUNDI. Bujumbura: Lewalla 5717 (MO). CO-

LOMBIA. Caldas: cult., Ldpez-Palacios 4023 (LL). DOMINICAN REPUBLIC. La Vega: Ososki & Rodriguez 249 (BRIT). GOLD COAST.

Knmari: D arko 598 (MO). TRINIDAD & TOBAGO. Smith Tr. 2 (LL[dil); Smith Tr. 24 (LL[diJ). MEXICO. Tabasco: Taylor & Taylor 12574

(BRIT). PARAGUAY. Canendiyu: Zardini 48550 (MO). SIERRA LEONE. Freetown: Johnston s.n. 3J Jan 1882 (MO); Morton SL185 (MO).

y 19 (BRIT). SRI LANKA. Central Moldenke et al 28159 (US); A

i. Dade Co.: cult., Avery 1867 (NY).

tania: Crosby &Doore 162 (SMU). 13. L. cuja

i: Huge 36 5 (LL[diJ). 17. L. planaltensis. BR

,n hybrid synonymy; ldxl2b, 2cxl2b, and l:

13. LantanacujabensissJu.au Prodr [A. P. dc Candolle! 11 1847 i i u M (

ing syntypes: BRAZIL. ;Rio Negro, Martius s.n. (M[di!l). PERU: Poeppig 148 5 (G[di!]).

(Urban 1906)], Riedel s.n. (holotype: LE, n.v.).

Lantana tenuifolia Rusby, Phytologia 1:74. 1934. Type: BOLIVIA. [La Paz:) Bopi River, 11 Sep 1921, Rusby 653 (holotype: NY!).

Lantana cujabens is Schauer var. parvijolia Moldenke, Phytologia 9:186. 1963. Type: PERU. La Libertad: Otuzco, 1 Jul 1951, Angulo &

Rusby 715 (holotype: NY!).

Shrubs rounded and ± open, lax, or subscandent; branches ascending or clambering, several to few, occasion-

ally herbaceous; twigs, peduncles and often petioles glabrescent to thinly setose or scabrous, the hairs (0.1- J

)0.3-0.5(-l) mm. Leaf-blades ovate, lanceolate, elliptic, or narrowly oblong, (3-)4-12(-16) cm long, the

length 1.3-2.5(-4) x width, nigrescent or not, papery to subcoriaceous, triplinerved to pinninerved; base

rounded or truncate and abruptly tapered onto petiole or cuneate and often forming an attenuate wing; apex

acute, abruptly acuminate, or attenutate; margin with the teeth (15-)25-40 per side, rounded to acute, often

appressed to strongly so and barely discemable, the sinuses 0.1-1.5 mm deep; adaxial surface dull to occasion-

ally lustrous, antrorsely strigillose to strigose-pubescent, the hairs occurring on veins and intervening tissue,

0.1-0.4(-0.7) mm, 3-20/sq mm, not noticeably vitreous-pustulate, the circular bases of the strigae ca. 0.1-02 J

mm in diam; abaxial surface slightly lighter or duller green than adaxial surface, sometimes whitish green but

not glaucous, antrorsely strigose-scabrous to nearly glabrous, with the strigae scattered on veins and veinlets,

0.1-0.6 mm, 3-KVsq mm. Inflorescences with peduncles 0.5- 2 x leaf length. Proximal bracts lanceolate,

oblong, or ovate-elliptic, often subfoliaceous, (4-)6-20 mm long, 2-8 mm wide, widest in proximal to middle

third, occasionally distal third, with 5-7 veins from the base, appressed to spreading, persisting and recurved

or reflexed in fruit; apex acute, briefly acuminate, or obtuse, often rounded at very tip; indument thinly stri-

gose, usually not distinedy ciliate, with longest hairs 0.1-0.6(-l) mm. Corolla yellow, orange, or light red aging

reddish orange to bright red (or occasionally intense reddish purple); corolla tube 7-12 mm.

Distribution and habitat. — Brazil (westernmost Amazonia, southern and central Planalto), the Guianas,

Venezuela, Colombia, Ecuador, Peru, Bolivia and Paraguay; understory and disturbance openings in tropical

humid forest, occasionally in dry forest or hard-pan savannas; 100-3000 m.

Lantana cujabensis is variable with respect to leaf-blade shape and width (broadly ovate to narrowly ellip-

tic-oblong) and marginal serration, outer bract length and width, and altitude preference. Apparently the type

specimens of L. cujabensis and L. riedeliana represent the broad-leaved, toothed vs. narrow-leaved, subentire

extremes, respectively; that of L. tenuifolia is intermediate. Different specimens exhibit all possible combina-

tions, which do not correlate with geography, therefore, no infraspecific taxa are justifiable based on the sam-

ple studied.

t eetdl&

Selected specimens examined: BOLIVIA. Cochabamba: Jaramillo etal. 1212 (MO); Ritter 1644 (MO); Steinbach 644 (F, N

1 1927 (MO). Santa Cruz: Guillen &Roca 3334 (F); Steinbach347 (NY, SMU); Steinbach 794 (SMU). BRAZIL. Acre: A I

i: Maas &• Maas 273 (MO). Bahia: dos Santos & Barreto 65 (LL[di]). Teixeim et al. 331 (M w

k Diaz 3478 (MO). Meta: Betancur 1336 (M0*

!. 2877 (F); Holm-Nielsen etaL 3016 (ft

Napo: Abbott 15637 (MO); Campos 135 (F); Cento 278 (MO); Croat & Hannon 93505 (MO); Ponce & Ght a 320 (MO). PERU. Amazonas: Casin'

1. 18983 (MO); Lewis et al. 18100 (MO). Cusco: Huamantupa et al. 4024 (MO). Macurdy 1007 (F, M0*

(MO); Santos et

il. 18843 (MO>, Castro et

2(F).L

k Fosberg 29017 (F); Fosberg 29104 (F); Gentry et al. 15599 (F); R

Sanders, Taxonomy of Lantana sect. Lantan

431

Z 745 (MO); Gudnchez 2132

(MO); Uesner 4021 (MO); Liesner 7454 (MO). Guarico: Ramirez 2027 (NY). Merida: Ldpez-Palados 2695 (LL).

Presumed hybrids with: 15. L. micrantha. BOLIVIA. Chuquisaca: Jimenez & Flores Til (MO). 17. L. planaltensis. BRAZIL.

Minas Gerais: Gentry et al. 49588 (MO), identification uncertain. PARAGUAY. Itapua: Perez 179 (MO). 19. L. paraensis. SURINAME: Si-

paliwini: Miller etal. 9367 (MO). VENEZUELA. Bolivar. Sanoja 2343 (MO). See also taxa Id, 2c, 4b, 9i (as cv.) and 12b and section on hy-

brid synonymy: 2cxl3, 4bxl3, and 15x13/20?

D. Lantana sect. Lantana series Spicatae R.W. Sanders, ser. nov. Type: Lantana viscosa Pohl ex Schauer

Adaxial leaf surfaces strigose-villosulous to setose-villous, the hairs up to 2.5 mm; abaxial leaf surfaces setose

or pilose, often densely so, the hairs occurring on veins and non-innervated tissue, setiform or filiform, 0.2-2.0

mm. Inflorescences initially hemispheric becoming short-cylindric by prolonged initiation of flowers or elon-

gation of internodes.

14. Lantana viscosa P

Shrubs erect or rounded and open to lax and subscandent; stems 1-3 m; branches ascending and numerous to

clambering and few; twigs, peduncles and often petioles sparsely setose but viscid with dense understory of

conspicuous stalked glands mixed with short hairs, the setae 1-2 mm, the stipitate glands and short hairs, ca.

0.5 mm. Leaf-blades broadly ovate, ovate, or ovate-elliptic, 2-8 cm long, the length 1.4-1.8(-2.2) x width, not

nigrescent, papery, pinninerved; base usually rounded to truncate, sometimes broadly cuneate or subcordate,

briefly narrowly cuneate onto petiole at very base; apex usually abruptly acuminate, sometimes acute; mar-

ginal teeth (10-)20-35 per side, acute to rounded, spreading to ascending, rarely appressed with tips recurved,

with sinuses (0.3-)0.6-1.5 mm deep; adaxial surface dull, setose to villous, the hairs occurring on veins and

intervening tissue, longer ones 1-1.5 mm or more (shorter ones ±. 0.5 mm, often mixed glandular and eglandu-

lar), 30-70/sq. mm, not noticeably vitreous-pustulate, the circular bases of the hairs ca. 0.1-0.2 mm in diam.;

abaxial surface slightly lighter or duller green than adaxial surface, setose to villous, the hairs on all veins and

intervening tissue, longer ones 1-1.5 mm or more (understory hairs 0.3-0.8 mm, these often mixed glandular

and eglandular), 50-150/sq. mm. Inflorescences occasionally 2 per leaf axil, becoming short-cylindric by

prolonged initiation of flowers; peduncles 0.5-1.3 x leaf length. Proximal bracts lanceolate or elliptic to ovate-

elliptic, (2.5-)4-7 mm long, 1.5-3 mm wide, widest in proximal or middle third, with 3 veins from the base, ±

spreading, persisting (proximally ± cupped around enlarging fruit) and becoming reflexed (± distally only) in

fruit; apex abruptly acuminate with prolonged tip to caudate; indument setose-pilose, usually ciliate, often also

stipitate-glandular, the longest hairs 0.7-1.5 mm. Corolla reddish purple to pale pink, often with white or yel-

low throat, occasionally white; corolla tube 5-10 mm.

Distribution and habitat.— Venezuela, Brazil (northern, eastern, and southern), Paraguay, Bolivia, and

Peru, possibly also Ecuador, Colombia, and the Guianas; widely distributed but infrequent; openings in tropi-

cal evergreen forest, tropical savanna with gallery forest, and areas of dense woodland, shrubland, and grass-

land; 100-1200 m.

Selected specimens examined: BRAZIL. Distrito Federal: Heringer et cd 7087 (NY). Pari: Strudwick & Sobel 4236 (LL[di], NY). Pernam-

buco: Figueiredo et al 18 (US). PARAGUAY. Amambay: Zardini & Guerrero (NY). VENEZUELA. Bolivar: Steyermarh cl al. 115523 (NY).

Presumed hybrids with: 15. L. micrantha. PARAGUAY. Cordillera: Merries & Degen 5522 (MO). See also taxa 2c, 4b, and 12a and

section on hybrid synonymy: 12axl4.

15. Lantana micrantha Briq., Annuaire Conserv. Jard. Bot. Geneve 7-8:299. 1904. Type PARAGUAY: AscunriOn, Apr

1874, Bdlansa 1039 (holotype: G(dU]; isotypes: Kldi!], P[2,di!]).

Shrubs erect or rounded, open; stems 0.5-2 m; branches ascending and several; twigs, peduncles and often

petioles moderately to densely setulose, setose, pilose, or also stipitate-glandular, the hairs usually 0.5-1.2 mm

(these sometimes lacking), mixed with shorter glandular and eglandular hairs about 0.2-0.3 mm. Leaf-blades

Sanders, Taxonomy of Lantana sect. Lantana

tween the more or less sympatric taxa L. camara subsp. camara and L. s plendens, probably prior to European

colonization or perhaps human habitation. Moreover, the range of variation is narrower than expected for a

hybrid swarm, and the distribution extends beyond the area of sympatry.

Selected specimens examined: BAHAMA ARCHIPELAGO. Cat Island: Britton & Millspaugh 5763 (NY); Correll 46083 (FTG, NY); Correll

46098 (FTG, SMU). Eleuthera: Lewis 7232 pp. (FTG). Grand Caicos: Gillis 12317 (LL). Great Exuma: Correll & Correll 42298 (FTG, NY);

oles moderately to densely puberulent or setulose, sometimes with stipitate glands intermixed, the hairs

mostly 0.1-0.4 mm, occasionally up to 0.8, rarely to 1 mm. Leaf-blades ovate, lanceolate or elliptic, 2-10 cm

long, the length (1.4-)l,7-2.7 x width, ± nigrescent, papery, triplinerved; base attenuate onto petiole from wid-

est point or rounded and abruptly narrowed to an often attenuate or cuneate petiolar wing; apex usually acu-

minate, sometimes acute; marginal teeth (14-)20-35(-50) per side, rounded, obtuse, or acute, spreading to

appressed, then sometimes tip recurved, with sinuses 0.3-1.2 mm deep; adaxial surface dull to somewhat

lustrous, antrorsely strigillose to strigose-pilose, the hairs occurring on veins and intervening tissue, forming

a thin canopy of hairs only 0.3-0.5(-0.8) mm with understory of shorter hairs often well developed, (2-)20-

80(-200)/sq. mm, sometimes vitreous-pustulate, the circular bases of the strigae ca. 0.1-0.3 m in diam.; abax-

ial surface slightly lighter or duller green than adaxial surface, moderately densely to sparsely pilose, if some

strigiform hairs mixed in, then filiform hairs dominating, the hairs on all veins and intervening tissue, 0.05-

0.5 mm, all about same length (or those on areoles evenly much shorter), occasionally a few along midrib to 0.8

mm, 20-150/sq. mm. Inflorescences occasionally 2 per leaf axil, remaining hemispheric; peduncles 0.2-0.7(-

1.3) x leaf length. Proximal bracts linear-, narrowly elliptic-, or lanceolate-oblong (occasionally 1 or 2 outer-

most bracts subfoliar or narrowly spatulate and distinctly longer), 4-7 mm long, 0.5-1. 5 mm wide, widest in

proximal or middle third, with 3 veins from the base, appressed or spreading, usually deciduous after flower-

ing; apex acute to attenuate, often rounded at very tip; indument pilose to strigillose, hardly ciliate, the longest

hairs 0.2-0.3(-0.5) mm. Corolla opening yellow or white with yellow throat, aging to dark yellow, orange, or

red, or opening white becoming infused with pink, blue, or purple, or opening pink, aging pink, purple, or

white with yellow throat (those opening with yellowish pigments becoming infused with purple also expect-

ed); corolla tube 7-12 mm.

Distribution and habitat — The Planalto of eastern and southern Brazil, northeastern Argentina, and east-

ern Paraguay; openings in tropical semi-evergreen forest, tropical savanna with gallery forest or areas of dense

woodland, shrubland, and grassland; 0-1000 m.

Lantana planaltensis to have arisen by hybridization between L. horrida subsp. tiliifolia and L. nivea subsp.

nivea. Furthermore its geographic distribution far exceeds the zone of contact of the probable parental species,

verifying its status as an independent species. The new name is required because L. hispida Kunth (= L. horri-

da) already exists. Most of the plants annotated by H.N. Moldenke and me as “ Lantana triplinervia " are in-

cluded here.

Selected specimens examined: ARGENTINA. Bnenos Aires: Cabrera 7020 (SMU, VBD); Krapovickas 2 891 (SMU) Misiones: Ekman 1985

(F); Schwarz 3635 (SMU); Schwarz 4074 (F); Zuloaga et al. 6633 (MO). BRAZIL. Distrito Federal: Heringer 13834 (NY); Heringer et al. 4253

(NY). Golds: Anderson 9479 (NY). Mato Grosso do Sol: Hatschbach 49116 (LLjdi]); Salvador 3094 (US). Minas Gerais: Hatschbach 25966

(US), Hatschbach 46673 (NY); Mexia 5436 p.p. (US). Parana Hatschbach 11224 (F); Hatschbach 16038 (VBD); Hatschbach 241 54 (US); Hatsch-

bach 41549 (NY); Wasum 2498 (BRIT); Winder 001 (BRIT). Peranamlmco: Silva etal.82 (MO, US). Rio de Janeiro: Carauta 3430 (LL[di]). Rio

Grande do Sol: M acedo 5507 (NY); Sehnen 9562b (US); Wasum et al. 1425 (US); Winder 006 (BRIT). Santa Catarina: Reitz & Klein 1778 (F);

ir rounded, open; st<

often petioles thinly to densely si

ovate or ovate-deltate to rotund, (

so, membranous to papery, pinnii

marginal teeth 5-15 per side, acut

1.5 (longest o

lairs 0.1-1.8 mm, the longest 0.8-1.8 mm. Leaf-blades broadly

long, the length 1-1.5 x width, not nigrescent or only somewhat

e rounded, truncate, or cordate; apex rounded to abruptly acute;

, spreading, with sinuses (l-)1.5-5 mm deep; adaxial surface dull

/illous, the hairs occurring on veins and intervening tissue, 0.1- f

usually 0.7-1.5) mm, (2-)5-20/sq. mm, noticeably vitreous-pustulate or not, the circular

bases of the hairs ca. 0.1-0.5 mm in diam.; abaxial surface slightly lighter or duller green than adaxial surface,

thinly to moderately densely setose or setulose to pilose, the hairs on most veins and some intervening tissue,

longest ones 1.5-2 mm on proximal portions of major veins, those increasingly distal gradually reduced (near

margin ca. 0.7 mm), those on intervening tissue mostly 0.2-0.5 mm, 5-20/sq. mm. Inflorescences remaining

hemispheric; peduncles 0.8-2.3 x leaf length (usually nearly twice when mature). Proximal bracts narrowly

oblanceolate or spatulate to elliptic-oblong, (5-)7-12 mm long, (l-)1.5-3 (rare outermost one subfoliar to 5)

mm wide, widest in distal half or near middle, with 3 veins from the base, appressed or spreading, persisting

and reflexed from base in fruit; apex obtuse or acute; indument strigose, ciliate or not, the longest hairs 0.3-0.7 |

mm. Corolla opening yellow, aging to red-orange; corolla tube 7-12 mm mm.

Distribution and habitat . — Central and southern Texas, Mexico (Coahuila, Nuevo Leon, Tamaulipas), ap-

parently cultivated and naturalized across the southwestern and southeastern United States from northern

Texas to California and to Florida and North Carolina; open woodlands, brushland, thickets, and grasslands

on calcareous clays or sandy soils; 0-1000 m.

Lantana urticoides likely originated by hybridization between L. hirsuta subsp. hirsuta and L. kingii, hav-

ing developed greater frost tolerance and a more northerly distribution than either parental species.

Selected specimens examined: MEXICO. Tamaulipas: Dominguez & McCart 8231 (SMU).U.S.A. Texas. Aransas Co.: Uzzell 51 (US). Comal

(SMU); San Patricio Co.: Jones 83 (SMU). Somervell Co.: Helm s.n. 9 May 1948 (SMU). Starr Co.: Garza et al. 8470 (SMU). Travis Co.: Han-

sen 26 (VDB); Lundell & Lundell 8928 (SMU). Uvalde Co.: Dickey 70 (SMU). Willacy Co.: Lundell & Lundell 8751 (SMU).

Presumed hybrids with: 20. L. strigocamara. MEXICO. Coahuila: Hazard s.n. May 1883 (US). Nuevo Leon: Rodriguez 62 (SMU);

Dodge 100 (NY). Tamaulipas: Berland ier s.n. 1836 (NY). U S A. Aiabama. Baldwin Co.: Krai 39530 (NY, VDB). Crenshaw Co.: Diamond

11455 (BRIT[di]). Florida. Citrus Co.: Krai 4542 (SMU). Marion Co.: Slaughter 13954 (BRIT[di], SMU). Monroe Co.: Krai 53896 (VDB).

RIT(di], NY). South Carolina. Orangeburg Co.: Leonard et al. 5001 (VDB). Texas. Blanco Co.: Sanders 5143 (SMU). C

» 6244 (US). Dallas Co.: cult. Niblack 50 (SMU). Fayette Co.: Krai 68519 (VDB). Galveston Co.: Waller 2579 (US). Sai

MU). Tarrant Co.: Krai 91937 (VDB); Whitehouse 16027 (SMU). Val Verde Ca: Spjut & Marin 15152 (BRIT). See also

Sanders, Taxonomy of Lantana sect. Lantana

435

e FRENCH GUIANA: Grand-Santi, 26 Aug 1961, Schndl

;: BRAZIL. ParA: 0° 55’S, 54° 26’W, 23 Jul 1981, Strudwick

Shrubs rounded and ± open, lax, or subscandent; stems 0.5-5 m; branches ascending and several or clamber-

ing and few, occasionally herbaceous; twigs, peduncles and often petioles moderately setose or pilose, often

with stipitate glands mixed in, or occasionally glabrescent, the hairs (0.1-)0.3-0.6(-l) mm. Leaf-blades

broadly to narrowly ovate or ovate-elliptic, 3-9 cm long, the length 1.4-2.2 x width, nigrescent or not, papery,

usually pinninerved; base rounded to subcordate and abruptly tapered onto petiole or cuneate; apex acumi-

nate, sometimes with a prolonged narrow tip, or acute; marginal teeth 15-35 per side, rounded or acute, often

appressed, then sometimes with tips recurved, with sinuses 0.5-1.5 mm deep; adaxial surface dull to occasion-

ally lustrous, antrorsely strigose or strigose-setose, the hairs occurring on veins and intervening tissue, 0.1-

0.6(-1.2) mm, 3-20/sq. mm, usually not noticeably vitreous-pustulate, the circular bases of the hairs ca. 0.1-

0.2 mm in diam.; abaxial surface slightly lighter or duller green than adaxial surface, sometimes whitish green

but not glaucous, antrorsely strigose to spreading-setose, with the hairs frequent on veins and veinlets, 0.1-0.7

mm, 3-10/sq. mm. Inflorescences often 2 per leaf axil, becoming short-cylindric (resembling spikes of Carex

lupulina ) by prolonged initiation of flowers; peduncles 0.5-l(-2) x leaf length. Proximal bracts lanceolate to

lanceolate-elliptic, (4-)6-8(-10) mm long, (1.5-)2-3 mm wide, widest in proximal third to just below middle,

with 5-7 veins from the base, spreading, persisting and recurved or reflexed (± distally) in fruit; apex acumi-

nate with prolonged tip; indument setose or strigose to glabrescent, usually distinctly ciliate, the longest hairs

(0.3-)0.5-l mm. Corolla rose-pink or white with pale yellow throat and aging pinkish purple, rarely yellow to

red-oranage; corolla tube 7-12 mm.

Distribution and habitat . — Brazil (central and eastern Amazonian), the Guianas, Venezuela (Amazonian)

Colombia (Amazonian),and Bolivia (Amazonian); disturbance openings, savannas and man-made grasslands

in tropical humid forest; 50-300 m.

Lantana paraensis exhibits characters of both L. cujabensis and L. viscosa , suggesting that it arose from

natural hybrids of the two. Its species status is suggested by greater consistency in the expression of its traits

than expected of a hybrid swarm and the much wider distribution than the area of sympatry of the proposed

Selected specimens examined: BOLIVIA. Santa Crnz: Guillen &Roca 3523 1

Croat 62252 (MO); Tsugaru. & Sano B-598 (MO). Para: Ginzberger 822 (F); P

et al. 664 (F, NY). COLOMBIA. Vichada: Davidse5196 (MO). FRENCH G

(MO). SURINAME. Paramaribo: Florschutz&Florschutz 1693 (SMU). Sipal

*6>Black27459 (LLldi]). Amazonas:

20. Lantana strigocamara R.W. Sanders, Sida 22:392. 2006. Type: U.S.A. Florida. Dade Co.: Coral Gables, 23 Sep

1981, Sanders 1450 (holotype: FTG!; isotype: NY!).

Shrubs erect or rounded, open; stems 0.3-3 m; branches ascending, several to numerous; twigs, peduncles and

often petioles thinly to moderately strigose, setose, or pilose, the hairs 0.1-1.2(-1.5) mm, the longest mostly

0.5-1 mm. Leaf-blades ovate to broadly ovate, (2-)5-10 cm long, the length 1-1.7 x width, usually not nigres-

cent, papery, pinninerved; base rounded, truncate, or cordate, shortly and narrowly cuneate onto petiole at

very base; apex usually acuminate; marginal teeth 15-40 per side, rounded to acute, often appressed, some-

times spreading at tip, with sinuses 0.5-1.5 mm deep; adaxial surface usually dull, antrorsely strigose or stri-

gose-setose, the hairs occurring on veins and intervening tissue (sometimes just center of areoles), 0.2-1.2

(longest mostly 0.5-0.8) mm, 1-12/sq. mm, not noticeably vitreous-pustulate, the circular bases of the strigae

ca- 0.1-0.2 mm in diam.; abaxial surface slightly lighter or duller green than adaxial surface, antrorsely stri-

gose-scabrous, with the strigae scattered to moderately dense on veins and veinlets, 0.1-0.6 (longest ones usu-

ally 0.4-0.6) mm (sometimes accompanied by scattered short [mosdy <: 0.3 mm] erect filiform hairs along

Sanders, Taxonomy of Lantana sect. Lantana

. horrida subsp. horrida

160. 1978. Type: MEXICO. Durango: 1

Iex2axl0. Lantana camara subsp. glandulosissima x L. horrida subsp. horrida x L. kingii

Lantana urticoides Hayek f. aculeata Moldenke, Phytologia 49:182. 1981. Type: MEXICO. Puebla: TehuacSn, 4 Aug 1966

Mex-28 (holotype: MEXU[di!]).

Lantana mdu ma ioldei 2:2 1982.1 PAN \M \ I on Sherman, 14 ]un 1923, M axon&

xlO. Lantana camara subsp. glandulosissima x L. kingii

Lantana glandulosissima Hayek f. parvifolia Moldenke, Phytologia 49:182. 1<

7933 (holotype: MEXU[di!]).

e: MEXICO. San!

lexl0/20. Lantana camara subsp. glandulosissima x L. kingii or L. strigocam

los: Yantepec, 14 Aug 1950, Wyatt 45 (holotype: MEXU[di!]).

Ifx2. Lantana camara subsp. aculeata x L. horrida (probably subsp. horrida)

/ iimwii.i niiKnlulivL I W'cieel. I'lijsioer. SaUk. I l.tndl I 4(> r’l. I \i i > nil < ireilsuald llr.i ( i.iril . >

lfx4. Lantana camara subsp. aculeata x L. hirsuta

Lantana mista L., Syst. Nat., ed. 12. 2:417. 1767. Camara aculeata (L.) Kuntze [var. subinermis Kuntze

Lantana albopurpurea Desf., Tabl. Ecole Bot., ed. 3 (Cat. PI. Horti Paris) 392. 1829. Type: cult., Hort. Paris, s. coll, (le

lfxC. Lantana camara subsp. aculeata x L. sp. Ser. Strigosae

2xC. Lantana horrida (probably subsp. horrida) x L. sp. Ser. Strigosae

2ax4a. Lantana horrida subsp. horrida x L. hirsuta subsp. hirsuta

Lantana poly acantha Schauer, Prodr. [A.P. de Candolle] 11:597. 18'

17481, F!]; lectotype, here designated: P, barcode P00713484[di!])

Lantana horrida Kunth f. inermis Moldenke, Phytologia 52:130.1982. Type MEXICO: Yucatan, Gamier 808 (holotype US!).

iaxlO. Lantana horrida subsp. horrida x L. kingii

dez424 (holotype: MEXU[di!]).

2ax20. Lantana horrida subsp. horrida x L. strigocamara

V 1948, Moldenke & Moldenke 19861 (holotype: N

L. SAo Paulo: Muji-Gaugu, 31 Jul 1!

2cxl2b. Lantana horrida subsp. tiliifolia x L. nivea subsp. mutabilis

^Wona camara L. f. glandulosa R. Fern., Bol Soc. Brot. sir. 2, 61:132. 1988. Type: ANGOLA, t

(holotype COl, n.v.; isotypes: BM|dil], K(dil], USU p.p.).

438

Journal of the Botanical Research I

? of Texas 6(1)

2cxl3. Lant

2cxl7. Lantana horrida subsp. tiliifolia x L. planaltensis

2185 (holotype: NY!). Alternatively nivea subsp. mutabilis could possibly be the second pare

2/4x? Complex hybrid involving Lantana horrida or L. hir

2. Type: MEXICO. Nuevo Le6n: Cafion Dient

Lantana horrida xinth f. bracteosa Moldenke, PhytologiT 52:231. 1982. Type: MEXKX). Puebla: Puebla, 15 Sep 1910, Arsdne 5426 (how-

4bxl3. Lantana hirsuta subsp. amazonica x L. cujabensis

Lantana cujabensis Schauer var. hispida Moldenke, Phytologia 46:58.1980. '

4bx 15. Lantana hirsuta subsp. amazonica x I ... micrantha

9i-cvx20. L. Callowiana Hybrid Group cultivars (derived from tetraploid cv. L. depressa var. depressa x

L. strigocamara)

viana Monrovia Nursery, Monrovia Nursery Catalog 1952-1953:44. 1952. nom. illeg. Type: none.

ra L. var. nana Moldenke, Phytologia 28:402. 1974. Lantana camara L. f. nana (Moldenke) Moldenke, Phytologia 45:296

e: U.S.A. New Yokx: cult.. New York. Bot. Gard., 14 Oct 1941, Moldenke & Moldenke 11903 (holotype: NY!). This is either an

r an independent cultivation of a wild-collected hybrid betw

Moldenke & Moldenke 29885 (holotype: LL, n.v.; isotype:

!). Lantana Callowiana Hybrid Group ‘Cre

n L. depressant.

id, 20 May 197V

Sanders (2001) argued that the parents of the Callowiana Hybrid Group were Lantana strigocamara and L. de-

pressa var. depressa rather than L. strigocamara and L. montevidensis (as claimed by Monrovia Nursery, see How-

ard 1969) based on character intermediacy and chromosome number incompatability of the latter combina-

tion. However, one likely hybrid of L. montevidensis with L. strigocamara was seen for this study (see taxon 20),

but it is very different in character details from the Callowiana Hybrid Group cvs, as well as appears to be sterik-

9iix20. Lantana depressa var. floridana x L. strigocamara

lOxla/le? Lantana kingii hybrid (x L. c

antana kingii hybrid (x L. horrida subsp. horrida or L. hirsuta subsp. hirsuta^

10x2a/4a? I

439

12x6/7? Lantana nivea x L. scabrida o

12axl4. Lantana nivea subsp. nivea x L. viscosa

Lantana pohliana Schauer, Prodr. [A.P. de Candolle] 11:601. IE

BRAZIL. GoiAs: “Inter Pirapora et Jenipapa,” Pohl 3088 (I

strayed [MacbrideNeg. 17480 at BRIT!, F!, GH!], F [fragm

The collections known to me combine the reduced leaves subtending inflorescences typical of some Lantana

nivea subsp. nivea (see comments, taxon 12a) and the stipitate glands, bracts, and elongating receptacles of L.

viscosa. They are geographically restricted near the type locality of L. pohliana, which is the area of sympatry of

the two species. For a narrowly endemic taxon, they exhibit a pronounced inconsistency in structure, length,

and density of trichomes compared to other natural taxa. These data suggest that the collections represent in-

dependent spontaneous hybrids, or at most, an unstable hybrid swarm. Thus, L. pohliana is not recognized as

12bxl7. L. nivea subsp. mutabili

;e) Moldenke, Phytologia 28:403. 1974. Type: BRAZIL. Bahia: Cocos

I? L. micrantha x L. cujabensis or L. strigocamara

2:468.1948. Type: ARGENTINA. Chaco: Colonu

18x20. L. urticoides x L. strigocama

Journey Mex., t.5 (top, facing p. 410). IS

EXCLUDED AND DUBIOUS NAMES

Lantana asperata Hort. ex Vis., Orto Bot. Padova 142. 1842. nom. nud. Type: unknown.

Lantana hahiensis Turcz., Bull. Soc. Imp. Naturalistes Moscou 36:206. 1863. Type: BRAZIL. Bahia: Salzrnann s.n.

(holotype: KW[di!]). Species of Lantana sect. Callioreas. (Misapplied to L. planaltensis by Moldenke in

sched.) c

Lantana Camara L. var. rosea Mattoon, Plant Buyer’s Guide, ed. 6. 167. 1958. nom. nud. Lanlam Camara L. f.

r«eu(Mattoon) Moldenke, Phytologia 45:296. 1980. nom. illeg. Type: none.

Lantana camara L. var. rubra Mattoon, Plant Buyer’s Guide, ed. 6. 167. 1958. nom. nud. Lantana camara

rubra (Mattoon) Moldenke, Phytologia 45:296. 1980. nom. illeg. Type: none.

Lantana cujabensis Schauer var. punctata Moldenke, Phytologia 2:411. 1948. - Lantana lopez-palacn Moldenke,

Phytologia 27:359. 1973. Type: COLOMBIA. Antioquia: Ceja, 1 Nov 1947, Barkley et al. 1536 (houttype:

MEDEL,n.v.). Species of Lantana sect. Callioreas.

Lantana hispida Kunth f. alba Moldenke, Phytologia 9:99. 1963. Type: GUATEMALA. PetEn: Ttka rums, jun

I960, Contreras 1056 (holotype: LL!). Aff. L. hirta Grah. of sect. Callioreas.

«— hispida Kunth f. parvt/dia Moldenke, Phytologia 52:130.1982. Tvm: HONDURAS. M^ Ct^ad

Univereitaria, 26 May 1978, Romero 71 (uoutrYm: MOD. Aff. L. velutina M. Martens & Galeom of sect

Lwa^m !ZL Briq. var. beck,, Moldenke, Phytologia 50:13. 1981. Tvrr: BOLIVIA. Bern: BallivMn, 12 Apr

1981, Beck 5339 (holotype: LL!). Aff. Lfucata Lindl. of sect. Callioreas.

Lantana multicolor Lem., FI. Serresjard. Eur. 3:239. 1847. nom. dub. Type: unknown.

Lantana notha Moldenke, Phytologia 1:422. 1940. Type: MEXICO. StNAUts: Fuerte, 27 Mar 1910, Rose et al. 13573

(holotype: NY!). Aff. L. hirta Grah. of sect. Callioreas.

bantam pulchra Larrahaga, Escritos DSntaso Antonio Larranaga 1:406. 1922 [Pub. Inst. Geog Uragl. nom.

dub. Type: Destroyed.

Lantana purpurea (Jacq.) Benth. & Hook!., Gen. Pi. [Benth. & Hooker f.] 2(2):1142. 1876. nom. illeg. (non Hor-

nem.) Species of Lippia or Lantana sect. Callioreas.

Lantana riedeliana Schauer var. pubescens Moldenke, Phytologia 19:435. 1970. Type: BRAZIL: Rio de Janeiro,

Pabst 9310 (holotype: LL!). Aff. L.fucata Lindl. of sect. Callioreas.

Lantana rosea Raf., Sylva Tellur. 83. 1838. Type: unknown. Probably a species of Lantana sect. Callioreas.

EPITHET INDEX (USING TAXON AND HYBRID SYNONYMY CODES’,

EDN=EXCLUDED AND DUBIOUS NAMES)

aculeata:la, If, 12a, 12b, Iex2ax6, lfx4, 2ax20, 2/4x?, 10xla/le?. aculeatissima:lex2a. aculeifera:ldx2c.

alba:12a, EDN. albiflora:le, 19, lexl0/20, 2cxl7, 9i-cvx20. albopurpurea:lfx4. amazonica:4b

amethystina: 1 2b. antidotalis:lx2xl2. antillana:2. arida:lb, 2b, 2bi. armatarld, 12b, 19, ldx6, 2cx4b,

4bxl5. asperata:EDN. aurea:9i. bahamensis:9ii, 16, laxl6, 9i-cvx20. bahiensis:EDN. bartramii:9iix20.

beckii:EDN. bracteosa:4axl0. brittonii:6. caffertyi:lax6. callowiana:9i-cvx20. camara:l, la-le, 2a, 2b,

7, 12a, 12b, 14, lax6, ldx2c, ldxl2b, lexl0/20, lfx4, 2xC, 2ax20, 2cxl2b, 9i-cvx20, 10xla/le?, 12x6/7?,

12axl4, EDN. canescens:laxl6. coccinea:lf. crenulata:6. crocea:la, 6. cujabensis:13, 19, 4bxl3, EDN.

cummingiana:2c. depressa:9, 9i-9iii. eitenorum:2cx4b. flava:le, 6, 2xC. floridana:9ii, 9iix20. foetida:2c

formosa:la. g!andulosa:2c, 2cxl2b. glandulosissima:le, 2bi, lex2a, lex4a, lexlO. glutinosa:2c,

2cxl3. grandiflora:2a. grandis:lex4a. guatemalensis:6. guianensis:19. hirsuta:4, 4a, 4b. hirta:4axl0.

hispida:2a, 17, 4bxl3, 10x2a/4a?, EDN. hispidula:18. hodgei:8. horrida:2, 2a-2c, 18, lex2a, 2ax4a,

4axl0. hybrida:2/4x? incarnata:12b. inermis:2ax4a. insularis:5. kingii:10. latibracteata:18. leonardio-

rum:3. longibracteata:4bxl2a. lopez-palacii:EDN. macrantha:2a. macrophylla:2axl0 micrantha:15

2cx4b, 4bxl5, 15x13/20?, EDN. microphylla:lex2a. minasensis:12a, 17, 4bxl2a, 12bxl7. mista:lf, If* 4

moldenkei:lc. montevidensis:9i. morii:12a. moritziana:ld, le, ldx2c, lexl0/20. multicolor:EDN. mul-

tiflora: 12x6/7?. mutabilis:lf, 12b, 20, lfx2. nana:9i-cvx20. niveailf, 12, 12a, 12b. normalis:lf, 12b.

notha:EDN. obtusifolia:la orientalis:2c. ovatifolia:9i, 11. paraensis:19. parviflora:2a. parvifolia:le

9i, 13, lexlO, 10xla/le?, EDN. planaltensis:17. pohliana:12axl4. polyacantha:2ax4a. portoricensis:lb

puberulenta: 1 2bx 17. pubescens:EDN. pu!chra:EDN. punctata:EDN. purpurea:lfxC, EDN. redinata:9i

riedeliana: 13, EDN. robusta:4bxl2a. rosea:EDN. rubella:2ax20. rubello-flavescens:ldxl2b. rubra:18x20,

EDN. ragosa:2cxl3. sandersii:6 sanguinea:lf. sanibelensis:9iii sargentii:2bi. scabra:ldx2c. scabrida:6

scabrifolia:13. scandens:6. scorta:4a. splendens:7. strigocamara:20. suaveolens:lf. subcordata:2bii

subinermis:la, If, 12a, 12b, lfx4. tenuifolia:13. temata:2a, 4axl0, 10x2a/4a? temifolia:ld. tiliifolia:2c,

ldx2c, 2cxl7. triplinervia:12a, 12b, 17, 4bxl2a, 12bxl7. urticifolia:la, lb, lc, 2b. urticoides:18, Iex2ax6,

2axl0. varia:lf. variagatarlf. velutina:ldx6 violacea:15xl3/20? viscosa:14. vulgaris:la. weberbaueri:2cx4b

zanonii:2b, 2bi, 2bii.

ACKNOWLEDGMENTS

I thank the following herbaria for searches, loans, digitization of specimens, and access to collections: A, AAU

BH, BM, BM-SL, BR, BRIT, B, B-WILLD, C, DWC, E, F, FI, FTG, G, G-DC, GH, GOET, HIPC, HOH, JE, K, KW,

LASCA, LINN, LIV, LL, M, MANCH, MEXU, MO, MPU, MSC, MVFQ, NY, OXF, P, P-HBK, PAD, PH, PL RB, &

SMU, STU, TENN; TEX, UC, UPRRP, UPS, US, VDB, W, WECO, WIS, WLU, WS, WTU, WU. Special thanks

go to the curators of BRIT and TEX for extensive digitization and of TENN for hosting loans. Financial support

was provided by the Appalachian College Association and Queensland Department of Primary Industries-

Constructive comments by Michael Nee and an anonymous reviewer are appreciated.

REFERENCES

Howard, R.A. 1969. A checklist of cultivar names used in the genus Lantana. Arnoldia 29:73-109.

Maschinski, J E. Sirkin, AND J. Fant. 201 0. Using genetic and morophological analysis to distinguish endangered taxa fro*"

their hybrids with the cultivated exotic pest plant Lantana strigocamara (syn: Lantana camara). Conservation Gen*

11:1607-1621.

146).

Offutt, ICE. and R.W. Sanders. 201 2. Identification guide to the Lantana camara complex: an interactive, multi-access key.

Bryan College, Dayton, TN. Accessed November 23, 2012 at http://www.bryancore.org/sliks/

Rotman, A.D. and M.E. Mulgura de Romero. 2010. Novedades nomenclaturales en los generos Lippia y Lantana

Sanders, R.W. 1 987a. Identity of Lantana depressa and L ovatifolia in Florida and the Bahamas. Syst. Bot. 1 2:44-60.

Sanders, R.W. 1987b. Taxonomic significance of chromosome observations of Caribbean species of Lantana

(Verbenaceae). Amer. J. Bot. 74:914-920.

Sanders, R.W. 1987c. A new species of Lantana (Verbenaceae) from Dominica, Lesser Antilles. J. Arnold Arbor. 68:

343-348.

Sanders, R.W. 1989. Lantana sect. Camara (Verbenaceae) in Hispaniola: novelties and notes. Moscosoa 5:202-215.

Sanders, R.W. 2001 . The genera of Verbenaceae in the southeastern United States. Havard Pap. Bot. 5:303-358.

Sanders, R.W. 2006. Taxonomy of Lantana sect. Lantana (Verbenaceae): I. Correct application of Lantana camara and

Santos Silva, T.R. 2001. Lectotypifications and neotypifications in Lantana and Lippia (Verbenaceae). Taxon 50:

1115-1118.

Schauer, J. C. 1 847. Verbenaceae. Prodr. [A.P. de Candolle] 1 1 : 522-700.

Schauer, J. C. 1851 . Lantana. FI. Bras. [Martius] 9: 251-266.

Stebbins, G.L 1966. Processes of organic evolution. Prentice-Hall, Englewood Cliffs, NJ.

Urban, 1. 1906.Collectores: Riedel, Ludwig. FI. Bras. [Martius] 1(1):89-91.

Journal of the Botanical Research Institute ofTexas6(2)

BOOK REVIEW

Craig Pittman. 2012. The Scent of Scandal: Greed, Betrayal, and the World’s Most Beautiful Orchid. (ISBN:

978-0-8130-3974-9, hbk.). University Press of Florida, 15 Northwest 15th St., Gainesville, Florida 32611-

2079, U.S.A. (Orders: http://www.upf.com). $24.95, 299 pp., b/w photos, 6" x 9 M .

After carefully reading this book, I turned to the back cover, and read the small heading: “TRUE CRIME/GAR-

DEN1NG.” The first review stated: “FANTASTIC. If I did not know most of the main players I would have

thought the author had a vivid and twisted imagination.” — Paul Martin Brown, author of Wild Orchids of

Florida ” — (and also a number of other orchid books and papers.) 1 was delighted. AND I had had about the

same reaction as he did. I had worked with Paul when 1 was Sr. Technical Editor for the Flora of North America j

North of Mexico project and had edited some of his contributions to the Orchidaceae. I also edited some of his j

other orchid manuscripts separately. I have tremendous respect for his contributions — AND his knowledge of

orchids.

Long, long ago, I had married the son of a well-known African Violet (and companion plants) family. After ;

college and a stint in the Air Force, we moved back to Illinois and setded down in a nearby small town. We im-

mediately got involved with the plants (Gesneriaceae and Orchidaceae, especially) — AND my mother and I

were quickly introduced to the monthly African Violet Club. (My father lovingly considered us all “a bunch of

delightful screwballs”)

People who REALLY love orchids, African violets, and companion plants really do tend to get very in-

volved, and often competition can get pretty nasty. Of course, there are local shows, state shows, regional, and

national shows. Ribbons are nice to earn, but tempers can really skyrocket at times. The more involved one

is — and the more competition gets heated— well, it can wreak havoc, even at the local level. When you get in-

volved enough to go to the state shows and the national shows, most people are really committed to doing their

best. However, while competition for new varieties is always a big challenge, when it comes to finding beautiful

and truly new species in native soils in other countries, it can become truly illegal, involved, and potentially

unlawful to bring in plants collected in other countries and illegally slipped through customs.

The Scent of Scandal is an actual, very carefully documented account of a rare and unusually beautiful or-

chid that got into the country, was recognized as an extremely new and beautiful specimen, caused all kinds of

interest, all kinds of trouble, was illegally named, and becomes an absolutely true, carefully documented case

history. Craig Pittman has done an incredible job of researching all strands of information, documentation,

actions (good or bad), timing, and ultimately providing 242 pages of page-turning fascination, 43 pages of fine

print, and a bibliography, tracing the path of one extremely beautiful, very rare, illegally named orchid but Stitt

bound to the name given it according to legal practice.

This is a page turner— with real people, real emotions, good intentions, devious actions, careless deci-

sions, and a very beautiful plant— legally or illegally “officially named.” It prompts numerous concerns all the

way: legalities, illegal actions, some knowingly, others perhaps not so knowingly. And every bit is carefully

researched and documented by the very capable author . — Helen Jeude, Volunteer and Assistant Editor, Botanic d

Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, U.S.A.

CYTOTYPIC VARIATION IN PHLOX P1LOSA SSP. PILOSA (POLEMONIACEAE)

AT THE WESTERN EDGE OF ITS RANGE IN THE CENTRAL UNITED STATES

Lindsey Worcester, Mark H. Mayfield, and Carolyn J. Ferguson

Herbarium and Division of Biology

Kansas State University

Manhattan, Kansas 66506-4901, U.SA

ferg@ksu.edu

RESUMEN

Polyploidy is frequent in plants and is considered an important factor in plant evolution (Otto & Whitton

2000; Adams & Wendel 2005; Soltis et al. 2009; Soltis et al. 2010). Autopolyploidy, or genome doubling within

a species, can lead to intraspecific ploidy level (cytotypic) variation (although some autopolyploids alterna-

tively meet criteria for species recognition; see Soltis et al. 2007). Such cytotypic variation within species may

be underappreciated, as systematists have sometimes made assumptions about ploidy levels for entire taxa

based on one or few chromosome counts. Advances in flow cytometry methods enable rapid assessment of

genome size and inference of ploidy level (Dole&l & BartoS 2005), and recent studies have demonstrated in-

triguing cytotypic variation in some species (e.g., Balao et al. 2009; Cires et al. 2009; Whittemore & Olsen

2011). Documentation of cytotypic variation is a critical first step toward understanding ecological, genetic,

and taxonomic consequences of autopolyploidy in particular groups.

Phlox pilosa L. (Polemoniaceae) is a showy, upright, perennial species ranging throughout most of the

eastern United States, from the east coast to the edge of the Great Plains and Central Texas (Wherry 1955; Great

Plains Flora Association 1986; Gleason & Cronquist 1991). Phlox pilosa exhibits noteworthy morphological

variation across its range, and several subspecies are usually recognized (Wherry 1955; Levin & Smith 1965;

Levin 1966; Locklear 2011; see also Ferguson 1998). Phlox pilosa ssp. pilosa occurs in prairies and woodland

openings across most of the general range of P. pilosa as a whole, with westernmost populations occurring in

north central Texas and south central Oklahoma. Most published chromosome counts for P. pilosa are diploid,

and represent the wide-ranging P. pilosa ssp. pilosa (e.g., Kelly & Wahl 1928; Flory 1931 [P argillacea Clute &

Ferris, a taxonomic synonym], 1934; Meyer 1944; Levin & Smith 1965; Levin 1966, 1968; Smith & Levin 1967;

see also Levin & Schaal 1970; Levy & Levin 1974). However, there has long been a suggestion of polyploidy

near the western edge of the range of P. pilosa ssp. pilosa. Smith and Levin (1967) reported a tetraploid chromo-

some count for material of P. pilosa ssp. pilosa from a mile south of Sachse, Texas (Dallas County, in north

central Texas) and one tetraploid count for material of P. pilosa ssp. pilosa from the state of Arkansas (precise

locality unknown). This has led to some question regarding ploidy level and possible cytotypic variation in P.

Phlox pilosa is one of the most thoroughly studied species of the genus Phlox (due particularly to the extenr

sive systematic and ecological research conducted by D.A. Levin and colleagues in the 1960s and 1970s; e.g.,

citations listed above), and further work on P. pilosa advances the utility of Phlox as a study system. Phlox is a

genus of ca. 60 species of perennial herbs occurring mostly in North America (Wherry 1955; Locklear 2011;

Ferguson et al. in prep. [FNA vol. 15]). The base chromosome number for Phlox is x=7 (Flory 1934), and diploid,

tetraploid and hexaploid chromosome counts have been reported (see Kelly & Wahl 1928; Flory 1931, 1934,

1937, 1948; Meyer 1944; Levin 1964, 1966, 1968; Levin & Smith 1965; Eater 1967; Smith & Levin 1967; Love

1971; Strakosh 2004; Fehlberg & Ferguson 2012 and in press). Furthermore, polyploidy has been implicated

as a factor contributing to complicated patterns of phylogeny, including incongruence between nrDNA and

cpDNA phylogenies for samples of P. pilosa and close relatives (Ferguson & Jansen 2002; see also Ferguson et

al. 1999). Study of cytotypic variation within P. pilosa ssp. pilosa can thus be placed within an extensive broad-

In this study, patterns of cytotypic variation in P pilosa ssp. pilosa were explored at the western edge of its

range, from north central Texas to eastern Oklahoma, western Arkansas, and southeastern Kansas. Flow cy-

tometry was conducted chiefly on fresh field-collected leaf material, as well as previously preserved material

for some populations (silica gel-dried, frozen, or herbarium material). Meiotic chromosome counts were per-

formed on bud material from several populations to enable ploidy level inferences based on genome size

(Dole&l et al. 2007).

MATERIALS AND METHODS

Field collection. — Material was collected from over 50 populations of P. pilosa ssp. pilosa ranging from the gen-

eral area of the earlier reported tetraploid count in Texas (Smith & Levin 1967) and north along the western

edge of the taxon range as far as southeastern Kansas. Fresh leaf material from an individual plant from each

population was reserved for flow cytometry. For some populations, leaves from multiple individuals were col-

lected in silica gel to enable later testing for cytotypic variation within populations. For populations at an early

stage of flowering, bud material for chromosome counts was collected (and preserved in a solution of 3 parts

95% ethanol: 1 part glacial acetic acid, with subsequent transfer to 70% ethanol). Voucher specimens fromaB

populations were prepared and deposited in the Kansas State University Herbarium (KSC). This study was

further augmented with material from several populations previously collected (material that had been col-

lected in silica gel or fresh-frozen and stored at -70°C, and one recent herbarium specimen).

Flow cytometry. — DNA content per cell was assessed for individuals using flow cytometry. At least one

individual was sampled per population, with additional within-population sampling for 11 populations. Leaf

material (0.04-0.10 grams) was chopped with a fresh razor blade in 2 mL of chopping buffer specified by Da-

vison et al. (2007; modified from Bino et al. 1993). The resulting slurry was filtered through 30 pm nylon mesh

(Small Parts, Inc.), followed by vortexing and centrifugation (500 RCF for 7 minutes). The supernatant was

decanted off of the pellet, or if a pellet did not form, approximately half of the liquid was decanted. Propidiu®

iodide staining solution (BioSure; 700 pL) was added to each sample to fluorescently stain the DNA; and 2 pf

of chicken erythrocyte nuclei (CEN) singlets (BioSure) were added to each sample as an internal standard

Samples were vortexed and stored on ice for 1-2 hours prior to processing on a Becton Dickinson FACS-Cafr

bur flow cytometer at the Flow Cytometry Lab of the College of Veterinary Medicine, Kansas State University

The flow cytometer quantifies the amount of DNA by measuring the fluorescence per nucleus. The result'

Worcester et al., Cytotypic variation in Phlox pilosa ssp. pilosa

ing histogram for each sample was analyzed using CellQuest software (Becton Dickinson), and a coefficient of

variation (CV) was obtained. To ensure highest quality data, we aimed to retain samples with measurements

based on ca. 10,000 nuclei per sample and having CV values <5% (see Galbraith et al. 1997; Dole£el & Bartos

2005). Picogram (pg) amounts were then calculated using the equation from Dolezel and BartoS (2005).

Chromosome counts. — Chromosome counts were conducted using a modified version of B.L. Turner’s

squash technique for meiotic cells (Jones & Luchsinger 1986). These counts enabled linking of picogram DNA

amounts to chromosome numbers for P. pilosa ssp. pilosa, and thus inference of ploidy levels based on flow

cytometry data.

Flow cytometry data were retained for 46 populations: these samples had 10,005-11,205 nuclei scored and CV

values from 1.91-4.98% (Table 1). Flow cytometry worked well for the Phlox material; most runs yielded high

quality data, although some samples did not meet criteria for inclusion (see Materials and Methods; due to

lower numbers of events, high CV values, or, occasionally, poorly defined peaks). Based on these data (Table 1),

cytotypic variation in P. pilosa ssp. pilosa was inferred, with 13 diploid populations, 30 tetraploid populations,

and three unusual populations with higher DNA content (discussed below). DNA content per nucleus for the

diploid and tetraploid populations averaged 10.66 pg (range 9.22-13.36 pg) and 21.90 pg (range 18.27-24.81

pg), respectively (based on measurements of all individuals; Table 1). The three remaining populations had an

average DNA content per nucleus of 26.55 pg (range 23.59-29.46 pg; Table 1).

For 11 populations, additional individuals (2-9) were analyzed. No cases of intrapopulational cytotypic

variation were inferred (Table 1). However, measured within population variation in picogram values ranged

from 0.26 pg (in diploid population 16) to 4.13 pg (in tetraploid population 17; Table 1).

Chromosome counts were obtained, including a diploid count (n= 7) and a tetraploid count (n=14; Fig. la,

lb; Table 1). Cells undergoing meiosis were also observed for two of the three populations that exhibited high

genome size values (populations 12 and 13). Cells from some buds of population 12 showed clear tetraploidy

(n=14), while those from other buds suggested additional chromosomes or chromosomal fragments (although

clear counts could not be obtained). Cells from buds of population 13 appeared to have more chromosomes

than the tetraploid material (n=17+), although the chromosomes could not be sufficiently spread apart. While

it is possible that higher level polyploids are present, these results may also be due to the presence of B chromo-

somes (supemumary chromosomes; see Smith & Levin 1967 for reports of B chromosomes in P. pilosa; see also

Meyer 1944 for notation of “fragments” in meiotic figures of some Phlox taxa).

Mapping the cytotypic data for sampled populations revealed a generally east-west pattern (Fig. 2), with

diploid populations occurring in the eastern part of the sampling range, tetraploid populations in the western

part, and the three populations with unusually high DNA content in southeastern Oklahoma (Fig. 2).

DISCUSSION

This study demonstrates that P. pilosa ssp. pilosa occurs as diploid and tetraploid populations at the western

e dge of its range from the southeastern Great Plains (sensu Great Plains Flora Association 1986) and south into

northern Texas. This variation does not correspond to any previously recognized infraspecific taxa in P pilosa,

and no morphological differences were noticed among cytotypes (future work will investigate micromorphol-

ogy of these populations). Interestingly, two subspecies of P. pilosa occurring further west in Texas (onto the

Edwards Plateau) are known to be tetraploid (P. pilosa ssp. latisepala Wherry and P. pilosa ssp. riparia Wherry;

see Levin 1966, 1968; Smith & Levin 1967). Some workers have recognized these Central Texas taxa at the

specific level, partly due to the polyploid condition: P aspera E.E. Nelson and P. villosissima (A. Gray) White-

house (Levin 1968; Levin & Schaal 1970; Levy 1973); or, as a single species under P villosissima (Turner 1998;

Locklear 2011). Our documentation of tetraploid populations of P. pilosa ssp. pilosa in northern Texas (and

northward) indicates that there are no ploidy level differences precluding intergradation with the Central

Texas taxa. Investigation of population level morphological, genetic, and cytotypic variation are warranted

is (in P pilosa ssp. latisepala and P.

within P pilosa thus occurs in the region sampled in the present study 0

northern Texas; in P. pilosa ssp. pilosa), west and south through Central Te

pilosa ssp. riparia).

Ploid, three populations in southeastern Oklahoma had unusually high DNA content per nucleus, and com-

pound light microscopy of meiosing cells suggested presence of chromosomes (or chromosome fragments)

beyond the tetraploid number. Further cytological study of these populations will be necessary to definitively

document ploidy. Accessory, B chromosomes have been documented in P pilosa ssp. pilosa (Smith & Levin

1967 > »n populations from central and northern Alabama, northeastern Arkansas, and northwestern Indiana),

and may well explain the variation detected here. Although perhaps less likely, higher ploidy levels are also

Journal of the Botanical Research Institute of Texas 6(2)

Worcester et al., Cytotypic variation in

>. pilosa 449

known in Phlox. For example, Fehlberg & Ferguson (2012; in press) have found diploid, tetraploid and hexa-

ploid cytotypes in R amabilis Brand and P woodhousei (A. Gray) E.E. Nelson (with evidence from chromosome

counts, flow cytometry, and population genetic data); Eater (1967) obtained diploid, tetraploid and hexaploid

chromosome counts for P nana Nutt, (following current taxon recognition; e.g., Wilken & Porter 2005); and a

hexaploid count is known for P. andicola E.E. Nelson (Love 1971).

Tetraploid individuals of P. pilosa ssp. pilosa have a genome size approximately double that of diploids

(averaging 21.90 pg [n=58] vs. 10.66 pg [n=19]; populations with unusually high genome size values averaged

26.55 pg [n=6]). Ranges in genome size within a ploidy level (e.g., 9.22-13.36 pg for individual diploids) were

somewhat greater than expected. For example, Fehlberg and Ferguson (2012) found picogram ranges within a

ploidy level of less than three picograms for R amabilis and P. woodhousei. However, their sampling on a per

cytotype basis within a taxon was more limited (with a maximum of eight individuals for a single cytotype

P. woodhousei; Fehlberg & Ferguson 2012). In the present study, there were no clear geographical patterns to

genome size variation within cytotypes: in some cases, geographically proximate populations of the same

ploidy level differed greatly (e.g., DNA content for two Wagoner County, Oklahoma, tetraploid populations was

21.60 vs. 24.15 pg), and measurements from individuals within populations differed (with the extreme being a

tetraploid population with a range in values slightly over four picograms). Variation in genome size observed

within a ploidy level for P pilosa ssp. pilosa may reflect, at least in part, an error level for flow cytometry mea-

surements in this study; it may reflect true variation in genome size within cytotypes, and this could be due in

part to differential presence of B chromosomes (discussed above; see Smith & Levin 1967); or, cytotypic varia-

tion may actually be more complex than inferred here (i.e, some additional cytotypes such as triploids and

hexaploids may be present).

Spatial patterns of cytotypic variation in P. pilosa ssp. pilosa show that tetraploids generally occur west of

diploid populations, potentially under more xeric conditions. While diploid and tetraploid populations were

detected in close proximity (within eight miles), we did not detect mixed cytotype populations (though, with-

in-population sampling was limited). Cytotypes are somewhat separated by geographic features, particularly

river systems: for example, the Neosho River in southeastern Kansas separates sampled diploids to the east and

tetraploids to the west. Into central Oklahoma (where most of our sampling was conducted), tetraploid popula-

tions tend to occur at slightly lower altitudes relative to diploids and west of major river systems, but detailed

study of potential ecological correlates of cytotypic variation in the area will require additional sampling.

Documentation of intraspecfic cytotypic variation in P. pilosa ssp. pilosa at the western edge of its range is

intriguing, and sets the stage for further study exploring ecological and cryptic morphological correlates of

this variation. This work advances study of systematics of Phlox by adding to our understanding of cytotypic

variation. Current work in our lab seeks to synthesize chromosome count and flow cytometry data for Phlox in

a revised taxonomic context. In a broader context, cytotypic variation may in general be underappreciated m

plants: for certain, systematists should not assume inference of ploidy level for an entire taxon based ononeor

few chromosome counts alone. Broader surveys (made easier through techniques such as flow cytoi

detailed studies of cytotypic patterns promise to improve our gen

sity in plants.

tnding of this aspect of biodiver-

ACKNOWLEDGMENTS

We gratefully acknowledge support from the following programs for LWs undergraduate research: NSF Un-

dergraduate Research Mentoring Grant URM-1041199, the KSU McNair Scholars Program, NSF-ESPCoR

(EPS-0903806), and the State of Kansas through Kansas Technology Enterprise Corporation; and from the

KSU Division of Biology. We thank Richard Noyes and Alan Prather for valuable reviewer comments, and

Shannon Fehlberg and the Ferguson lab group for valuable comments on an earlier version of the manuscript.

We thank Theresa Melhem Susan Rolfsmeier, Wanda Worcester, and Bethany Wright for assistance with held

work; Mike Herman for use of his camera-equipped microscope; and Molly Bernstein, Bernard Friebe, Theresa

450

Journal of the Botanical Research Institute of Texas 6(2)

Melhem, Susan Rolfsmeier, Spencer Tomb, Bethany Wright, and the KSU Flow Cytometry Facility (School of

Veterinary Medicine) for technical assistance. Online specimen database resources are gratefully acknowl-

edged: R.L. McGregor Herbarium (KANU), Oklahoma Vascular Plants Database (particularly Oklahoma State

University Herbarium [OKLA] and Robert Bebb Herbarium [OKL]), and Flora of Texas database (LLTEXk

This is publication 12-452-J of the Kansas Agricultural Experiment Station.

l.Bot. 95:99-1 10.

ersity of California, Santa

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Journal of the Botanical Research Institute of Texas 6(2)

BOOK REVIEW

David Bramwell and Juu CaujapE-Castells (eds.). 2011. The Biology of Island Floras. (ISBN 978-0-531-11808- j

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lbs., 72 b/w illus., 36 tables, 9 3 A" x 6%".

It is estimated that the biodiversity of 53 archipelagos includes between 50,000 and 52,000 plant endemic species woHd- |

wide that are highly threatened, and 20,000 of these species are in critical danger of extinction as part of fragile ecosystems. I

This suggests the urgency and importance of continued study of island biodiversity and conservation highlighted in this I

book. The 21 separate chapters and 48 authors discuss the evolution, diversity, and conservation of island vascular plants. I

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el B. Crespo, Michael F. Fay, Mark W. Chase, Alain Vanderpoorten, Ben Laenen, Rosalina Gabriel, Juana M. Gonzalez-

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Jorge Rodriguez, Sara Oldfield, and Vernon H. Heywood.

The Socotra archipelago in the Indian Ocean off the coast of Yemen currently counts 835 as endemic vascular plant

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I.BotRes. Inst Texas 6(2): 452. 2

A NEW COMBINATION AND A NEW SPECIES IN

COMBRETUM (COMBRETACEAE) FROM INDIA

K.A. Sujana, M.K. Ratheesh Narayanan 1 , and N. Anil Kumar 1

Central Botanical Laboratory, Botanical Survey of India

AJCB Indian Botanic Garden, Botanical Garden

P. 0., Howrah, West Bengal - 71 1 103, INDIA

Author for correspondence: sujanacabc@yahoo.com

lalabaricum (Bedd.) Sujana

RESUMEN

INTRODUCTION

The genus Quisqualis L. (Combretaceae) comprises ca. 16 species of lianas from the Old World tropics (Exell &

Stace 1966). Recent research, however, has shown that the supposed differences with Combretum Loefl. cannot

be maintained on the basis of molecular evidence (Jongkind 1991; Stace 2007; Maurin et al. 2010). Quisqualis

dates from 1762, Combretum from 1758, and the latter name therefore has priority. Gangopadhyay and

Chakrabarty (1997) recognized two species of Quisqualis from India, viz., Q. indica L. and Q. malabarica Bedd.

The latter is here transferred to Combretum and a new combination, viz., Combretum malabaricum (Bedd.) Su-

jana, Ratheesh, & Anil is proposed. Both species have been recorded for Kerala (Nayar et al. 2006; Sasidharan

2011).

During botanical exploration as part of a project on “Investigation of rare, endemic and threatened woody

climbing plants of the Western Ghats,” one of us (Sujana) collected specimens with recurved petals from the

Aralam Wildlife Sanctuary in 2008 which did not match with any known species of Combretum or Quisqualis

and is therefore described as a new species here. The species, although morphologically similar to the Combre-

tum malabaricum (Bedd.) Sujana, Ratheesh, & Anil, comb, nov., differs from it in some characters; these data

are given in Table 1.

Combretum malabaricum (Bedd.) Sujana, Ratheesh, & Anil, comb. nov. (Fig. 1). Quisqualis malabarica Bedd., icon.

PL Ind. Or. 1:33, 1. 155. 1874. Type: INDIA. Kerala: Carcoor ghat Wayanad, dev. ca. 450 m, Beddome s.n. (lectotype: MH - Acc. No.

20405; isolectotype: BM).

Combretum recurvatum Sujana, Ratheesh, & Anil, sp. nov. (Figs. 2, 3). Type: India. Kerala. Kannur district: Aralam

Wildlife Sanctuary, ca. 800 m, 28 Feb 2008, Su jana 0509 (holotype: CAL; isotypes: CAU).

A liana, 15-20 m tall. Stems 15-25 cm in diameter; bark light brown with vertical striations, spines 1.5-2.5 cm

lon g, at the base of the branchlets; young branchlets puberulous, purplish, terete. Leaves opposite, ovate to

oblon g, symmetric, 4-15 x 4-7 cm, membranous, glabrous above, lead-coloured when dry, brown beneath,

domatia as tufts of brown hairs in the axils of the secondary nerves, base rounded, margins entire to undulate

**• Res. Inst Texas 6(2): 453 - 458. 2012

Sujana et al., A new s

455

Flt - 2 ‘ ^return recurvatum Sujana, Ratheesh, & Anil. A. Habit, B. Inflorescence, C Flowers - dose up, D. Fruit

Journal of the Botanical Research Institute of Texas 6(2)

457

458

with tawny hairs, apex shortly acuminate or acute, lateral veins 5-8 pairs, slender, adaxially faint, abaxially

prominent, arcuate, tertiary nerves adaxially faint, sometimes abaxially prominent, scalariform, marginal ul-

timate veins recurved to form loops; petioles 1-1.2 cm long, sulcate, densely pubescent. Inflorescence termi-

nal, spicate, 18-26-flowered; peduncles slender, purplish, puberulous, 7-11 cm long; bracteoles linear-lanceo-

late or triangular, ca. 5 x 1 mm, tomentose, deciduous. Flowers 5-merous, 1.5-1.7 x 1.5-1.6 cm; pedicels 0-1

mm long, puberulous. Calyx tube pale green, narrowly infundibuliform, ridged, 0.9-1 cm long, puberulous;

teeth triangular, ca. 3 x 2 mm, ciliolate, apex recurved. Petals white, turning pink with age, ovate or oblong,

13-14 x 6.8-7 mm, recurved with 3 distinct nerves, glabrous, apex obtuse. Stamens 5 + 5, inserted in the throat

of the calyx tube; filaments ca. 1 mm long, white; anthers yellow, basifixed, ca. 1 x 0.7 mm. Ovary 4-5 x 1-1.2

mm, tawny puberulous, ridged, 1-celled with 3 ovules; upper free portion of style 1-2 mm long, yellowish

white, slender, flattened; stigma capitate, creamy white. Fruits chocolate brown, chartaceous, 2-2.3 x 1.8-2.2

cm, villous when young, glabrous when mature, horizontally faintly striate, widely ellipsoid; stalk ca. 1 mm

long, wings 5, thin, 10-12 mm broad. Seed solitary, fusiform, ca. 13 x 0.5 cm. Funicle ca. 6 mm long, curved;

testa brown, membranous.

Distribution. — Combretum recurvatum is so far known only from Aralam Wildlife Sanctuary, Kannur

district of Kerala, India (Fig. 4).

Flowering. — -January-March; Fruiting: February-May.

Habitat. — Rarely seen in the semi-evergreen forests of the Aralam Wildlife Sanctuary, Kerala (Fig. 3), at

an altitude of 800 m above sea level and is found associated with Anamirta cocculus (L.) Wight & Am., Baccrn-

rea courtallensis (Wight) Mull.-Arg., Combretum latifolium Blume, C. razianum K.G. Bhat, Sterculia foetida L.,

and Vateria indica L.

Additional specimens examined: INDIA. KERALA. Kannur district: Aralam Wildlife Sanctuary, alt. 800 m above sea

level, 26 Mar 2009, Sujana 0714; 6 May 2009, Sujana 0786; 2 April 2010, Sujana 0833 and 6 Apr 2011, Sujana 0913 (Herbari-

um of M.S. Swaminathan Research Foundation, Wayanad).

The research grant provided by Sir Dorabji Tata Trust, Mumbai, is gratefully acknowledged. Sincere thanks are

expressed to J. F. Veldkamp (L) for valuable remarks and comments. We very much thank Clive Stace (LTR) for

confirming the novelty; Carel Jongkind (WAG) and Brent Berger (WIS) for their advice on the Combretum/

Quisqualis relationships; and P. Lakhshminarasimhan (BSI) for reviewing the paper. Thanks to K.M. Manudev,

former Research Fellow of MSSRF for the drawings.

REFERENCES

Exell, A.W. AND CA Stace. 1 966. Revision of the Combretaceae. Bol. Soc. Brot. II 40:5-25.

Gangopadhyay, M. and T. Chakrabarty. 1997. The family Combretaceae of Indian subcontinent. J. Econ. Taxon. Bot. 21: 1

Jongkind, C.C.H. 1991. Novitates Gabonenses, 6. Some critical observations on Combretum versus Quisqualis (Combre-

taceae) and description of two new species of Combretum. Bull. Mus. Natl. Hist. Nat. Sect. B, Adansonia 12:275-280.

hase, M. Jordaan, and M. van der Bank. 2010. Phylogenetic relationships of Combretaceae inferred from

jstid DNA sequence data: implications for generic classification. Bot. J. Linn. Soc. 1 62:453-476.

egam, N. Mohanan, and G. Rajkumar. 2006. Flowering plants of Kerala— a handbook. Tropical Botanic

Sasidharan, N. 2011. Flowering plants of Kerala - Ver 2.0 (CD-ROM). Kerala Forest Research Institute, Peechi, Thrissur,

Kerala, India.

Stace, C.A. 2007. Combretaceae. In K. Kubitzki, ed. The families and genera of vascular plants. Springer-Verlag, Berlin,

Heidelberg. 9:67-82.

CONVOLVULACEAE OF SONORA, MEXICO. 1.

CONVOLVULUS , CRESSA, DICHONDRA, EVOLVULUS, 1POMOEA,

JACQUEMONTIA, MERREMIA, AND OPERCULINA

Richard S. Felger

Herbarium, University of Arizona

P.O. Box 210036, Tucson, Arizona 85721

and Sky Island Alliance, Tucson

rfelger@ag.arizona.edu

Thomas R. Van Devender

Daniel F. Austin

Arizona-Sonora Desert Museum

fi 021 N. Kinney Road, Tucson, Arizona 85743, U.S. A.

and Herbarium, University of Arizona, Tucson

J. Jesus Sanchez-Escalante

Universidad de Sonora

Dept, de Investigaciones CientificasyTecnoldgicas

Rosales y Ninos Heroes, Centro

Hermosillo, Son, 83000, MEXICO

jsanchez@guayacan.uson.mx

Mihai Costea

Dept, of Biology

Wilfrid Laurier University

75 University Avenue W

Waterloo, ON, N2L 3C5, CANADA

mcostea@wlu.ca

RESUMEN

INTRODUCTION

The Convolvulaceae constitute a large and diverse assemblage of 58 genera and about 1800 species worldwide

(Staples 2011). This monophyletic family has greatest diversity in tropical and subtropical regions worldwide

and does not generally occur in higher latitudes. The sweet potato ( Ipomoea batatas) and water spinach (I.

aquatica) are the only major crop plant in the family. In addition to species with horticultural value (e.g., vari-

ous morning glories), there are significant crop weeds (e.g., certain species of Convolvulus, Cuscuta, and

Ipomoea) and many medicinal uses among Sonoran people and worldwide. This publication is the first taxo-

nomic account of all the known Convolvulaceae (convolvs) native or naturalized in the state of Sonora, Mexico

(Fig. 1). We include approximately 84 species and 2 infraspecific taxa in 9 genera for the state. Due to the large

size of the article, we divided it into two parts: the first part includes all the Convolvulaceae genera minus Cus-

cuta, while the second part provides a floristic/taxonomic treatment for the latter genus (Costea et al. 2012a).

This introduction includes the 9 genera.

Although a pohtical border is not necessarily an ecological or biological boundary, the borders of Sonora

are to varying degrees biologically and logistically significant. The western boundary (except the extreme

northwestern comer) is the Gulf of California. The east boundary mostly coincides with the continental di-

vide. The north boundary marks a division between the better-known continental flora of Arizona and the

relatively less-known flora of northern Sonora. The southern border with the state of Sinaloa border separates

a continuous flora but is far enough south to include the northern climatological limits of the New World trop-

ics (in east-central Sonora at 28-29°N).

Sonora encompasses 185,934 km 2 (Molina-Freaner & Van Devender 2010) and is the second largest state

in Mexico after Chihuahua. The region is topographically and biologically diverse (Molina-Freaner & Van

Devender 2010). Three major river systems, the Rio Colorado, Rio Yaqui, and Rio Mayo, and several minor riv-

ers systems course through the state and empty into the Gulf of California. The eastern margin of the state is

comprised of the Sierra Madre Occidental and numerous north-south trending Sky Island ranges forming the

Madrean Archipelago. The highest elevation is 2625 m in the Sierra de los Ajos in northeastern Sonora. The

Sierra San Luis in the extreme northeastern comer is nearly as high, and another high peak, further south on

the Chihuahua border, near Mesa Tres Rios may be equally high. Numerous other ranges, generally decreasing

in peak elevations westward, spread across the rest of the state, interspersed with broad valleys and expansive

plains.

Sonora includes the northern limits of tropical and subtropical biota as well as some of the most arid des-

ert regions of North America. Tropical species follows the lowland tributaries in the Rio Yaqui drainage system

northward in northeastern Sonora.

Total annual precipitation decreases from south to north and east to west, and increases with elevation.

Rainfall is largely bi-seasonal with summer and winter-spring r ainy seasons. Precipitation increases from west

to east: for example from the Rio Colorado River to northeastern Sonora/New Mexico, and with elevation and

from north to south. Total annual precipitation varies from less than 40 mm in the extreme northwest of the

tr the delta of the Rio Colorado to about 1000 mm in southeast and east-central Sonora (e.g., Y£cora at

and probably considerably more farther north, such as the Tres Rios-Sierra Huachinera region (Brito-

et al. 2010; Felger et al. 2001; Martinez-Yrizar et al. 2010).

man populations in Sonora remained sparse and major roads were relatively few until the mid-twenti-

1500 ir

Felger etal., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

eth century (e.g., Stoleson et al. 2005). Much diversity and richness of natural habitat remain, but like else-

where, assault* on the environment are escalating (e.g., Castellanos-Villegas et al. 2010). Since the latter part of

the twentieth century large areas of the state have been established for conservation purposes (e.g., Burquez

and Martinez -Yrizar 2007, Felger and Broyles 2007).

Major works including ethnobotanical information on convolvs in Sonora include Felger and Moser

(1985), Gentry (1942, 1963), Yetman and Felger (2002), and Yetman and Van Devender (2002). For summary

information see species accounts for Ipomoea ancisa, I. arborescens, I. bracteata , and I. pedicellaris.

While soil moisture is the principal limiting factor in this generally arid region, freezing temperatures

limit the northern distributions of the more tropical or subtropical species (e.g., Brito-Castillo et al. 2010;

Shieve 1951; Turner et al. 1995; also see the species accounts for Ipomoea arborescens ). Freezing weather is in-

frequent across most of Sonora, especially in the southern part of the state and at low to moderate elevations,

and many habitats are essentially frost-free, but freezing temperatures become more frequent and severe north-

ward and at higher elevations (Brito et al. 2010).

The total flora of the state includes about 3700 taxa (Van Devender et al. 2010 and new records). Within

this rich flora, the Convolvulaceae is the seventh largest family, following the Asteraceae, Poaceae, Fabaceae,

Euphorbiaceae (Steinman & Felger 1997), Malvaceae (s.l.), Cactaceae (Paredes et al. 2000), and Cyperaceae

(Van Devender et al. 2010). With 42 taxa, Ipomoea is the fourth largest vascular plant genus in the state and the

largest genus in the family worldwide. Among the documented convolv species in Sonora, we include species

that occur in Arizona and New Mexico close to the northern boundary and are expected to occur in Sonora.

Growth forms. — Most convolvs throughout the world are scandent annuals or herbaceous and woody

perennials. Three native Sonoran Ipomoea are trees or large shrubs: I. arborescens, I. chilopsidis, and I. seaania.

The non-native I. cornea subsp. fistulosa is a shrub while I. bracteata and some Jacquemontia may become some-

what shrubby. Ipomoea bracteata and I. longiflora form large tuberous roots, as do I. capillacea and I. plummerae,

while Merremia palmeri, Operculina pinnatifida , and O. pteripes probably have large tuberous roots because

their closest relatives have them.

The majority of Sonoran convolvs are vining or have twining stems. The liana growth form, characteristic

of tropical regions, occurs among Ipomoea bracteata, I. pedicellaris, several Jacquemontia, Merremia quinquefo-

lia, and Operculina pteripes. All occur in tropical deciduous forest in Sonora and some also range into thom-

scrub and oak woodland. Cuscuta spp. are always annual when parasitic on annual hosts; however, when para-

sitizing on perennial woody hosts, some species such as C. americana, C. corymbosa var. grandiflora, and C.

tinctoria are often perennial, regenerating every year from haustorial tissues left inside the stems of the host

(Costea & Tardif 2006).

Endemism and Rare/Endangered Taxa.— Although many species may be locally rare and known from

few collections in Sonora, most of these are common elsewhere. Only a small number of Sonoran convolvs are

rare and/or endangered throughout their range.

Ipomoea seaania, the only convolv endemic to the state of Sonora, is known only from the vicinity ot the

type locality and seems to be globally restricted to a relatively small area. Among Cuscuta, C. dentatasquamata

is known in Sonora only from the type collection; C. solina is known from one locality in Sonora but is common

elsewhere, and the only known Sonoran C. tinctoria specimen is a parasite on a cultivated tree and is not native.

Tkere are two records for Dichondra brachypoda in Sonora, both in mountains in the northeastern part of the

state. Evolvulus prostratus is known from two collections in Sonora but is widespread elsewhere in Mexico.

Ipomoea alba, with two records, is common elsewhere and may or may not be native in Sonora. Ipomoea ancisa

is a relatively narrow endemic in mountains in eastern Sonora and western Chihuahua but is locally common.

Jacquemontia abutiloides is widespread in Baja California (none) and Gulf of California islands and approaches

Sonora on Isla Tiburon, although a thorough taxonomic investigation may render it a synonym of a mainland

s P ec ' es - Merremia cissoides, with a single Sonora collection, is a cosmopolitan species.

Non-natives. — About eight convolvs found in Sonora are not native to the state. Convolvulus Owens ts and

1 xl-wunthu are widespread weeds. Ipomoea cornea subsp. fistulosa, I. triloba, and Merremia disjecta are prob-

463

ably of Caribbean origin and I. batatas, the sweet pototo, is of tropical American origin. The origin of I. hep-

taphylla is not known although it is probably native to the Old World. Ipomoea batatas and I. camea subsp.fis-

tulosa are cultivated and are sometimes encountered outside of cultivation. Cuscuta campestris and C. indecora

are weedy and probably have been introduced, and C. tinctoria probably is also not native to the flora.

Diversity. — In comparison with neighboring areas to the north and west (Arizona, the two Baja Califor-

nia states, California, and New Mexico), the convolvs are quite diverse in Sonora. This is due to the more tropi-

cal affinities of much of the Sonoran flora as well as habitat diversity. As one moves further south in Mexico and

Central America, the convolv diversity continues to increase. A comparison of the convolv diversity in Sonora

with adjacent and comparable areas in Mexico and the southwestern United States is shown in Table 1.

Vegetation of Sonora and the Convolvs

The major habitats or vegetation types in Sonora include mangroves and coastal vegetation, tropical deciduous

forest, thomscrub, desertscrub (both Sonoran and Chihuahuan), grassland, oak woodland, pine-oak forest,

and mixed conifer forest (Fig. 2, Table 2). The Sonoran convolvs are distributed as follows: Chihuahuan desert

(11), Sonoran desert (33), coastal thomscrub (30), foothills thomscrub (31), tropical deciduous forest (41),

grassland (13), oak woodland (34), pine-oak forest, 27), mixed conifer forest (1), salt scrub (1), mangroves (1).

These vegetation regions are briefly mentioned below, with some examples of characteristic convolvs. For dis-

cussions of the vegetation of Sonora the reader is referred to Brown (1982), Felger et al. (2001), Gentry (1942),

Martin et al. (1998), Martinez-Yrizar et al. (2010), Rzedowski (1978), and Shreve (1951).

Coastal vegetation. — Mangroves occur sporadically along the coastal fringe of the southern two-thirds

of the state, bordering salt scrub. Cressa truxillensis occurs along beaches and extends into tidal marshes (lo-

cally called esteros ) of salt scrub and sometimes at the inland border of mangroves, as well as inland in some

agricultural areas. Ipomoea imperati and I. pes-caprae are found along beaches.

northeastern Sonora and adjacent southeastern Arizona at elevations below about 1430 m. This landlocked

desert covers much of north-central Mexico between the Sierra Madre Occidental and Sierra Madre Oriental

extends into adjacent inland areas of the southwestern United States. Hard freezes may occur in the Chi-

diich accounts for the absence of columnar cacti and reduced convolv diversity. Rainfall

mostly occurs during the summer. The Sonoran portion of the Chihuahuan desert is bordered by grassland

and oak woodland. As with the Sonoran desert, there is open ground, and shrubs predominate. The substrate

often consists of limestone and alkaline soils. A total of 12 convolv species are recorded for the relatively small

area of the Chihuahuan desert in Sonora (one occurs adjacent in Arizona) and all except Ipomoea cardiophylla

also occur in adjacent vegetation.

Sonoran desert.— The Sonoran desert, defined and described elegantly by Forrest Shreve (1951), covers

roughly the northwestern two-thirds of Sonora. Shreve divided the Sonoran desert into seven geographic veg-

etation zones, five of which occur in Sonora, although Shreve’s Foothills of Sonora is reclassified as thomscrub

(Felger & Lowe 1976), leaving the Plains of Sonora, portions of the Arizona Upland, the Sonora portions of the

Crural Gulf Coast, and the Lower Colorado Valley. Within the span of environments in the Sonora portion of

the Sonoran desert there is great variation in vegetation cover and structure and regional plant diversity. Thir-

ty-two convolv species are documented for the Sonora portion of the Sonoran desert.

Ray6n 33, Rosario

, San Luis Ri'o Colorado 1, San Miguel de Horcasitas 42, San Pedro de

>, Tubutama 13, Ures 43, Villa Hidalgo 30, Villa

a !.

1 ss»ss

] !h!i!

3 2 i

i.ciflt

,-Jll

toslilil

.»*Ss!if

I»

Felger et al., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

^Unlike thornscrub there is much open ground. Most of the perennial Sonoran desert plants have evolved

° m ,ro P ical or subtropical relatives that are found today in thornscrub and tropical deciduous forest. Many of

1 Perennials are sensitive to winter freezing, their northern limits fixed by an invisible line of freezing tem-

peratures and drought. The desert is essentially frost-free at its southern limits, such as near Guaymas, and the

***** and Oration of freezing increases northward. Similarly, drought is more severe northward and west-

(Felger 2000; Turner et al. 1995).

show a strong seasonality linked with monsoonal rains and for the most part are highly frost-sensitive. Unlike

TDF, natural thornscrub generally does not form 100% perennial coverage. The stature of the vegetation is

generally lower and the dominant species tend to have smaller leaves than those of the TDF. Thornscrub in

Sonora was described as Sinaloan Thornscrub by Brown (1982) and Thom Forest by Gentry (1942).

Two thornscrub formations can be discerned in Sonora: the coastal thornscrub (CTS) of southwestern

Sonora (Felger and Lowe 1976; Friedman 1996; Martin et al. 1998) and the interior, foothills thornscrub (FTS)

along the east side of the Sonoran desert and at higher elevations within the Sonoran desert. We document 30

convolv species in CTS in Sonora and 31 in FTS.

Coastal thornscrub is on the coastal plain from Empalme-Guaymas southward. Foothills thornscrub is

on inland, often rocky slopes. In southern Sonora, FTS is below tropical deciduous forest in elevation. In cen-

tral Sonora, it is the transitional vegetation between the Plains of Sonora subdivision of the Sonoran desert on

the west and oak woodland in Sky Island mountain ranges and the Sierra Madre Occidental to the east. In the

north FTS is replaced by desert grassland as winters become colder and periodic fires become ecological pro-

cesses. The northern limits of FTS in Sonora are at about 30°11'N in the Rio Sonora Valley and 30°26'N on the

Rio Bavispe at the southern end of the Sierra El Tigre. FTS does not reach Arizona, but the distributions of a

number of FTS species extend into southern Arizona in desert grassland or oak woodland.

Coastal thornscrub extends southward on the coastal plain from the southern margin of the Sonoran

desert in the vicinity of Guaymas into coastal northwestern Sinaloa. Southward and eastward this vegetation

type merges with TDF. Much of the coastal thornscrub has been converted to large-scale modem agriculture.

Foothills thornscrub. — This vegetation type is essentially synonymous with Shreve’s (1951) Foothills of

Sonora subdivision of the Sonoran desert (Felger and Lowe 1976; Felger et al. 2001). FTS is shrubby or semi'

arborescent vegetation with a nearly closed canopy of small trees and large shrubs. FTS extends northward

along the eastern side of the Sonoran desert, becoming narrower in geographic and elevational range towards

its northern limits. Northward it gives out at about the vicinity of Arizpe, where rise in elevations and winter

freezing as well as drier conditions become pronounced.

Tropical deciduous forest. — The northern arm of TDF, sweeping northward from the tropics, ends in

the mountains of eastern Sonora and southwestern Chihuahua. Sonoran TDF is sandwiched between thorn-

scrub to the west at lower elevations and oak woodland eastward at higher elevations. Northward, along the

east side of the Sonoran desert, TDF merges into foothills thornscrub (Felger et al. 2001; Martlnez-Yrizaretd.

2010).

There is a long dry season interrupted by a short but intense rainy season generally from mid-June to

October. Summers are long and hot and winters short and mild. Freezing weather is rare and most Sonoran

TDF species are highly frost-sensitive. Awesome and seemingly sudden transformation to luxuriant tropical

green occurs with the onset of the summer monsoon. TDF, with 44 documented species, supports a greater

convolv species richness than any other vegetation in the state. The tree morning glory, lpomoea arborescens is

a common and conspicuous component, as are others such as lpomoea bracteata with its spectacular pink in-

florescences. lpomoea muricata, I. quamoclit , Merremia cissoides (with one record), M. dissecta, and M. quinqut

folia are more or less restricted to TDF.

Grasslands.— The southwestern extension of the Great Plains grassland biome in the mid-continent is in

the southwestern United States and northwestern Mexico. The best developed regional grasslands with mo*

rainfall and colder winter temperatures termed Plains Grassland are restricted to the Animas and San Rafael

Valleys along the Arizona-Sonora border. Most other areas are termed desert grassland (McClaran & Van

Devender 1995), with warmer, drier climates. Dominant or common species in the regional desert grassland

have fluctuated four times during the last 4,000 years— between bunch grasses during wetter periods and

shrubs such as mesquites (P rosopis glandulosa, P. velutina ) and others during drier periods (Van Devender

1995). Today desert grassland controlled by human disturbance related to cattle grazing is widespread in val-

ley lowlands below oak woodland from northeastern Sonora west to the Sdsabe area southeast of the Baboqui-

van Mountains and south to the Cananea-Fronteras area within 100 km of the Arizona border.

In southeastern Arizona and northeastern Sonora, valley bottom desert grassland forms a mosaic with

Chihuahuan Desert on rocky limestone slopes. To the west, Sonoran desert borders desert grassland at its

lower elevations. To the south, desert grassland is replaced by foothills thomscruh below oak woodland, al-

though there are local areas of grassland still present. In addition, there are open grassy areas within oak

woodland and pine-oak forest in the higher Sky Island mountain ranges and the Sierra Madre Occidental. As

freezes decrease southward, foothills thornscrub replaces desert grassland. Only 13 convolvs are documented

from grasslands in Sonora. Characteristic species include Ipomoea longifolia, a large herbaceous perennial often

conspicuous sprawling across expanses of short grasses, and Evolvulus seriteus, a small, silvery-leaved herba-

ceous perennial.

Oak woodland. — Oak woodland is widely distributed at elevations above desert, grassland, thornscrub,

and tropical deciduous forest, but below pine-oak woodland. In Sonora, 34 convolv species occur in OW.

Across Sonora, and northern Mexico, the species composition and density of oak woodland changes both with

elevation and from south to north. Although these oak zones have been called Madrean Evergreen Woodland

(Brown 1982), many Sonoran oaks and associated species are deciduous during the late spring drought, and

their biggest flush of new foliage occurs with the renewal of summer rains. In these regions "autumn color”

occurs in late spring as the air and ground desiccate and temperatures soar.

Extensive areas in northeast and north-central Sonora are dominated by open woodland of Quercus emo-

ryi. This species, with Q. oblongifolia and Q. arizonica, are among the most common low-elevation oaks in the

northern part of the state. At lower elevations the oaks border desert grassland, foothills thornscrub, and des-

ertscrub. There is sometimes a broad ecotone between oak woodland and grassland where the oak trees be-

come widely spaced and grasses predominate. Oak woodland sometimes occurs on acidic, hydrothermally

altered soils within tropical deciduous forest where the ecotone between the two plant communities is often

only a few meters wide.

Oak woodland in southeastern Sonora, called Oak Forest by Gentry (1942), shows considerable tropical

affinity. The lower limits border tropical deciduous forest, and the boundaries are often remarkably well de-

fined, apparently maintained by fire. Fire, however, is not an ecological process in desertscrub, thornscrub, or

tropical deciduous forest. Across mountains in the Rio Mayo and Rio Fuerte drainages, low fires creeping al-

most harmlessly through dry grasses and forbs among the leafless oaks used to be a common sight in May and

June. These fires destroy small TDF trees and shrubs but not the perennial grasses, forbs, and oaks. Many of the

oaks in east-central and southeastern Sonora and nearby southwestern Chihuahua are tropical montane oaks.

Ipomoea chilopsidts is the most spectacular and unique convolv among the diverse 34 convolv species in oak

woodland.

Pine-oak forest.— There are numerous montane islands of pine-oak forest in the mountains of eastern

Sonora. However, pine-oak forest is more extensive east of Sonora in Chihuahua along the east side of the con-

tinental divide. In comparison, on the western slope of the Sierra Madre Occidental the climate is generally

somewhat wetter, with presumably milder winter temperatures, resulting in a more diverse flora with more

tropical-derived pines such as Pirns engelmannii, P. h errerae, P. oocarpa, and a number of tropical-montane

oaks such as Q. tarahumam. The pine-oak forest has been included within the concept of Madrean Evergreen

Woodland and Madrean Montane Conifer Forest (Brown 1982; Martin et al. 1998). Towards southeastern

Sonora the pine-oak woodland is floristically and structurally more like Mexican pine-oak woodland than the

temperate pine-oak woodland to the north. Pine-oak forest, where the pines form the overstory while the oaks

generally form an understory, is continuous with oak woodland at lower elevations. Among the 27 convolv

species in POF, Cuscuta dentatasquama (the type collection is the only Sonora record), Ipomoea ampullacea (one

ttC0rd > see the species accounts), and Ipomoea decasperma (one record) appear to be restricted to this habitat.

Mixed conifer forest.— This zone is restricted to limited areas on the several highest mountain tops in

northeastern Sonora and very limited areas near Yecora, in the upper Rio Mayo drainage. Three conifers, Abies,

Pinus ’ especially p strobiformis, and Pseudotsuga , define this vegetation. Evolvulus wtundifolius is the only con-

volv recorded in mixed conifer forest in Sonora.

470

Convolvulaceae Jussieu - Morning glory Family

is, subshrubs, shrubs, or trees, some species with milky sap. Rootsto<

simple, entire to pinnately lobed or pecti

Twining herbs, ltanj

tuberous, otherwise fibrous. Leaves a]

cies palmately compound; stipules absent. Inflorescences solitary in leaf axils or in racemose or paniculate

cymes, some dichasial basally and monochasial above. Flowers small and inconspicuous to large and showy,

but usually wilting quickly after opening (mostly within 4-5 hours) except Cuscuta and often excepting plants

flowering during cooler weather, bisexual (or unisexual in som<

irregular. Sepals 5, distinct, imbricate, equal or unequal, persistei

alous, tubular, funnelform, campanulate, urceolate, or salverform, 5-lobed, 5-toothed, or ± entire, with plicae

(areas folded in bud) and interplicae (unfolded in bud), usually induplicate and often also convolute in bud.

Nectary disc annular or cup-shaped, sometimes 5-lobed, occasionally absent. Stamens 5, distinct; filaments

inserted on the corolla tube base alternate with corolla lobes; anthers dithecal, usually linear or oblong, ex-

trorse or introrse. Ovary superior, 2-4(-6)-carpellate, usually with as many cells, placentation basal or basal-

axile, ovules 2 (4-6) per cell, or ovary 1-celled and ovules 4, these erect, anatropous; style 1, filiform, simple or

bifid, or sometimes with 2 distinct styles; stigmas capitate or bilobed, or, when stigmas 2, then linear, ellipsoid,

or globose. Fruits capsular, dehiscent by valves, transversely or irregularly, or indehiscent and baccate or nut-

like. Seeds l-4(-6; to 10 in Ipomoea decaspema), often fewer than ovules, glabrous or pubescent, endosperm

mated 1880 (Staples 2011), cosmopolit;

a 9, species about 84 in Sonora.

SPECIES ACCOUNTS

infraspecific taxa. The months or seasons noted refer to the recorded times of reproduction. Flowering and

fruiting usually overlap broadly, and therefore we generally do not distinguish separate flowering and fruiting

times. Many species that are reproductive at various seasons do so facultatively, mostly depending on soil

moisture and temperature. There is, however, a marked tendency for members of the family to have flowering

initiated by short day length and thus they usually flower in the autumn.

Representative specimens are cited in the last paragraph of each species or infraspecific taxon account

We include the Sonora municipios (mpio.), as of 2011, to help place the collection localities (Fig. 1). We also in-

clude records from Gulf of California islands nearest the Sonora coast (Islas Tibur6n, Alcatraz in Bahia Kino,

Datil, San Esteban, and San Pedro Nolasco; see Felger and Wilder 2012, Felger et al. 2011). We have seen all

specimens (except those specified as “not seen”) and use “!” for all specimens or images that we have seen, ex-

cept in Cuscuta where “!” is used for the types seen. All Cuscuta specimens cited have been seen by Costea. All

other convolvs have been seen by Austin and/or Felger.

Unless otherwise indicated, specimens cited are deposited at the University of Arizona Herbarium

(AR1Z); specimens in other herbaria are indicated by the abbreviations given in Thiers (2011). Most of our col-

lections are duplicated in the herbaria of USON, MEXU, and SD and other regional and international colfcc-

len is at AR1Z, we generally do not cite duplicates at other herbaria. When a specimen lacks*

collection n

r, it is identified by the date if

nple: Ezeurra 9 Nov 1982. We gei

abridge label information, but provide enough that one can find the specimen at a herbarium or search addi-

tional information in a data base, especially SEINet (Southwest Environmental Information Network 2012) j

and MABA (Madrean Archipelago Biodiversity Assessment 2012). Usually only the first collector’s name's

listed. Elevations and reproductive times (flowers and fruiting) are mostly from herbarium label data and I

specific only for Sonora. Coordinates for specimens cited are often available in SEINet (Southwest Environ-

mental Information Network 2012), however these might not have come from the collectors’ labels. Many co-

ordinates were added/determined by students entering information by looking on Google Earth, or other maj* |

and some may be inaccurate (especially for specimens from Mexico). Coordinates for many specimens cited ;

are not repeated here to save space. North America is defined here as Mexico northward, excluding Cen0»

Felger et al., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

America. Hundreds of photographs that illustrate diagnostic details of morphology for the majority of species,

are provided on a companion website — Convolvulaceae (morning glories) of Sonora, Mexico, which is hosted

at ARIZ and WLU (Costea et al. 2012b). Plants not native to flora area are marked with an asterisk (*).

TYPIFICATIONS

Special attention has been given to study of type specimens for the taxa included, not only to assure the correct

names but also for proper identification. These data are included because there have been errors in citing types

in the literature. Types have been checked by consultation of protologues and corrected where necessary. Type

sheets have been studied at various herbaria or as loans, and also from online databases at various herbaria,

some through JSTOR Plants (2011). Moreover, because of the information now available, it is possible to note

many more duplicates of type collections than previously known. The information provided here gives a more

complete picture of the available resources for each taxon than formerly obtainable.

Problems and errors in typification are discussed in several cases. Some taxa have never had lectotypes

selected. In several cases, we designate lectotypes to establish proper use of names and concepts. In other cas-

es, various problems with former selections of types have been located and these are discussed and resolved

except in the case of E, alsinoides var. angustifolia. Lectotypes are selected for nine species and new information

and clarifications are provided for others: Convolvulus palmatus (in Merremia dissecta). Convolvulus pennatus

(in Ipomoea quamoclit), Cuscuta corymbosa var. grandiflora, lpomoea ancisa, Ipomoea alata (in Opcrculina

pteripes), I. alatipes (in Operculina pteripes), I. decasperma, l. pedicellaris, Merremia dissecta, Operculina pinnati-

fida, 0. pteripes, and O. roseana (in Operculii

sBenth., Pi. Hartweg. 16. 1839. Type: MEXICO. Nuevo Leon: 1837. Hart weg 98 (m

des A. Gray. Syn. Fl N. Amer. 2(1):216. 1878. Type: U.S.A. T

Convolvulus inconus sensu Kearney & Peebles, not Vahl (see also Staples et al. 2006).

Perennial herbs with branched, prostrate or decumbent steins arising from a taproot; densely hairy. Leaves

variable, ovate-elliptic to triangular-lanceolate or narrowly oblong with projecting basal lobes, blades most

often deeply indented basally, 1-7 cm long, 0.2-4 cm wide, densely hairy on both surfaces with loosely ap-

pressed indumentum, margins toothed or lobed or both, rarely entire; petioles 0.25-5 ci

usually 1-flowered, less often 2 or 3 and cymose, on peduncles 0.5-10.5 c

late, 1.5-3 mm long or sometimes scale-like, hairy like the leaves. Flowers often solitary, on short pedicels

5-24 mm long. Sepals oblong to ovate, 6-12 mm long, 3-6 mm wide, obtuse to weakly retuse apically, ap-

pressed sericeous, the margins membranaceous, subcordate with age. Corollas (1.5—) 2.5-3 cm long, cam-

panulate, white or pink to pale lavender, at times with a reddish center, sericeous on the petal lobes. Capsules

± globose, 7-8 mm wide, glabrous. Seeds 1-4, 4-4.5 mm long, black, granulate, glabrous, n = 12.

Sonora . — Northeastern part of the state in Chihuahuan Desert, grassland, and oak-mesquite scrub; often

in disturbed sites; 1200-1600 m. Flowering April-December.

General distribution. — California, Arizona, New Mexico, Texas; Chihuahua, Coahuila, Guanajuato, Hi-

dalgo, Edo. Mexico, Michoacan, Nuevo Leon, Oaxaca, Puebla, QuerCtaro, San L

Mna2009-1747\ Mpio Cans

CRESSA L., Sp. PI, 223. 1753. [Greek, based on feris or kriti , “from Crete,” a Cretan woman; apparently not ety-

mologically related to cress or cressa (Cruciferae) of Germanic derivation].

Alkali weed

Perennial herbs or subshrubs, gray appressed pilose to sericeous, usually much-branched, the stems erect to

decumbent. Leaves sessile or short-petiolate, entire, small or scale-like. Inflorescences axillary, 1-flowered.

Flowers appearing sessile or on short peduncles (at least some, e.g„ C. truxillensis), bracteate, in spicate to

head-like clusters at tips of branchlets, bracteoles unequal in length. Sepals ovate to obovate, imbricate. Corol-

las salverform, the limb 5-lobed, the lobes mostly ovate, imbricate, spreading to reflexed. Stamens and styles

exserted; filaments filiform; pollen 3-colpate. Ovary 2-locular, 4-ovulate; styles 2, distinct to the base; stigmas

capitate. Fruit capsular, ovoid, unilocular, 2-4-valved, usually 1-seeded. Seeds glabrous, smooth and shining

to reticulate, dark brown.

Species 4, two in the Americas, one in Asia, and one in Australia.

Selected reference .— Austin (2000b).

Prodr. 9:440. 1845

ed.) 3:119. 1818 [1819]; also folio ed. 3:93. 1819. Type: PERU:

ioto F!, isotype: F!). Cressa cretica var. truxillensis (Kunth) Choisy in DC.,

7-July3,l

474

s gray, appressed pilose to silvery-sericeous, not twining, usually much-branched, at

first erect, becoming decumbent to spreading, often 8-25 cm long in open, sunny habitats, or often with weak

slender stems to 75 cm long when growing through and branching over the tops of other salt-marsh halo-

phytes; stems dying back during adverse times to thickened rhizomes and/or rootstocks often 8-15+ cm below

the surface; lower stems often semiwoody. Leaves on main branches often larger than those on branchlets,

with pubescence like the stems, subsessile or with petioles 0.5-2 mm long; blades mostly 3-10 mm long, 1-4

older leaves sometimes thick and succulent in hypersaline habitats. Inflorescences of solitary flowers, axil-

lary, usually concentrated in the upper leaf axils and appearing almost spicate. Flowers on stalks (peduncles

and pedicels) 2-6 mm long, the pedicel much reduced and more slender than the peduncle, the bracteoles

ovate to ovate-lanceolate, acute, unequal, 2-3 mm long, mostly 1 mm wide. Sepals ± equal or the inner slightly

longer; outer sepals obovate, elliptic, 3-4 mm long, 2.5-3 mm wide, obtuse or acute, pubescent; inner sepals

obovate, 3-4.5 mm long, 2-3 mm wide, acute, with scarious margins, appressed sericeous only at the apex.

Corollas white, becoming scarious when dry and somewhat persistent, salverform, 5-6.5 mm long, the tube

3-3.5 mm long, the limb 5-lobed. Corolla lobes obtuse to ± acute, about as long as the tube, somewhat pubes-

cent on the outer surface, becoming reflexed with age. Stamens exserted, 4-6 mm long, usually equal, the fila-

ments basally pubescent with glandular indumentum; anthers red and becoming purple with age, 1-1.5 mm

long, oblong with the base cordate to ± bilobed. Ovary ovoid, unilocular to ± bilocular, apically somewhat hir-

sute; styles pure white, unequal, 3-5 mm long; stigmas pure white, capitate, smooth. Capsules 5-6 mm long,

ovoid, shiny brown, surrounded at least basally by the calyx, apically hirsute, unilocular. Seeds usually 1,

ovoid, 3-4 mm long, brown, glabrous. 2n = 28.

Sonora . — Often common in tidally wet saline mud and sandy soils among saltscrub and margins of man-

groves, esteros, bays, and low-lying coastal soils from the Rio Colorado delta to the Sinaloa border. In north-

western Sonora sometimes in small inland playas but near the coast, and as an agricultural weed south of San

Luis, especially in fine-textured silty-clay alkaline soils. Sonoran desert and coastal thornscrub: 0-20 ffl.

Flowering March-December.

In contrast to populations in northwestern Sonora, this species has apparently become relatively rare, at

least in recent years, in nearby southwestern Arizona (Austin 1992). In the early 1990s Cressa was a common

weed in sandy soil of seawater-irrigated experimental plots at the Environmental Research Laboratory adja-

cent to Estero Morua at Puerto Penasco. Cressa had been evaluated earlier as a potential halophytic seed crop

in these plots. Substantial seed crops were obtained, but no economic value was found and further evaluation

was terminated. This is probably the first report of a weed in seawater-irrigated agriculture (Felger 2000).

General distribution.— California, Arizona, Nevada, New Mexico, Texas, Utah; Baja California (norte) and

Sur, Chiapas, Chihuahua, Coahuila, Colima, Jalisco, Nuevo Le6n, Oaxaca, San Luis Potosi, Sinaloa, Revilla-

gigedo Islands; Ecuador, Peru, Chile, Argentina.

San Luis R.C.: Colorado River, opposite mouth of Hardy River, 29

6 Oct 1985, Felger 92-989. SONORAN ISLANDS: Alcatraz, Bahia

Miguel, E side of island, 23 Nov 2006, Wilder 06-366! (See Felger a

>r 1990, some older leaves succulent, Felger 91-40. Mpio

ns 2842 (CAS!); Ci£nega de Santa Clara, 5 km S of Rilhto,

17, Felger 07-167!; Tiburdn, base and N side of Punta San

DICHONDRAJ.R. <Sr G. Forster, Char. Gen. PI. 40, t.

r, PENNYWORT, FALSE PENNYWORT

[Greek 6ixov6pa, di “double” and chondra “a

Herbs with slender glabrous or pubescent and mostly repent stems, from perennial taproots; stems sometimes

rooting at nodes. Leaves petiolate; blades cordate-orbicular to reniform, small, the margins entire. Inflores-

cences of inconspicuous, axillary, solitary (rarely paired) flowers on short to long peduncles. Flowers usually

less than 1 cm wide, greenish-yellow or white (purple in D. occidentals, but that one not in the flora area). Se-

pals ± equal, barely united basally, often spathulate. Corollas broadly campanulate to subrotate when living,

appearing funnelform at times when dried, usually deeply 5-lobed, the lobes induplicate. Stamens typically

included, more or less equal. Pollen 3-colpate. Ovary 2-lobate to emarginate, the lobes distinct or usually ba-

sally united, 2-locular. Styles 2, attached between the lobes and appearing almost gynobasic, filiform, the

stigmas capitate. Fruits capsular or utriculate, 2-lobed to almost entire, membranaceous, usually 2-seeded,

irregularly bivalvate or indehiscent. Seeds obovoid, smooth, the cotyledons linear, 2-plicate.

Species ca. 15; tropics and temperate regions: 8 species in North America, ±7 in South America, the others

in Australia and New Zealand.

Selected references . — Austin (1998a), Johnston (1963), Tharp and Johnston (1961).

Dichondra argentea Humb. & Bonpl. ex Willd., Enum. Pi. 297. 1809. Type: COLOMBIA: "Habitat in America merid.ona-

li prope Hundam” (from protologue), no date, Humboldt & Bonpland s.n. (B-W, presumably barcode B -W 05468-01-0 and the re-

verse, 05468-02-0, having “ Dichondra argentea (W)" in what seems to be Willdenow’s handwriting).

Perennial herbs, whole pi ant except roots, stamens, corolla and styles silvery-canescent with a dense pubes-

cence of long, silky, usually appressed, flattened hairs; taproots perennial, dark brown, 1.8-4 mm thick; sto-

lons annual, 10-35 from the crown, ca. 0.7-1.1 mm thick, rarely branched; intemodes l-2.5(-3) cm long;

nodes often bearing adventitious roots and occasionally short-shoots with crowded nodes. Leaves reniform,

5~13 mm long, 12-20 mm broad, apically often shallowly emarginate, basally truncate or broadly and shal-

lowly cordate, with a narrow cuneate base where the blade joins the petiole; petioles (l-)2-3(-5) cm long,

0 5-1 mm thick, erect, straight. Inflorescences on pedicels 4-6 mm long, 0.5-1 mm thick, basally sharply re-

CUrved - Flowers 3.4-4 mm wide. Calyx broadly campanulate, 2-2.6 mm long, accrescent to 2.4-3 mm in fruit,

5-lobed two-thirds to three-fourths the length, the lobes linear-oblong, apically blunt and often somewhat re-

curved - Corollas nearly cylindrical, cream-colored, 5-lobed about half the length or more, the lobes subulate.

on 'y slightly bilobed. Fruits capsules 2.2-2.S mm long, and 2-2.1 mm wide. Seeds 1.9-2.4 mm long,

°* Did t0 Pynform, dark brown to black when fully mature.

This species is known from southeastern Arizona close to Sonora as well as southwestern New Mexico

and Chihuahua and is likely to be in the Chihuahuan desert region of northeastern Sonora (Austin 1998a).

Ak ° m ch iapas, Coahuila, Durango, Edo Mexico, Nuevo Leon, San Luis Potosi,Tamaulipas, Zacatecas.

u AWZ0n A: Cochise Co.: Bisbee, 26 Sep 1931 , 1

; of Texas 6(2]

Felgeretal.,

a, Mexico

477

108°45'W, 1210 m, 7 Sep 1995, Fishbein 2526!; La Pila, 0.6 km E of El Trigo Moreno, riparian forest in pine-oak forest, 28°17'48"N,

108°47'irW, 1450 m, 30 Aug 2008, Rei na-G. 2009-429'.

EVOLVULUS L., Sp. PI., ed. 2, 1:391. 1762. [Latin, evolvulare, to enroll, an allusion to their non-twining habit.]

Herbs or small suffrutescent shrubs, annual or perennial, not twining but sometimes creeping. Leaves usually

small; blades ovate to almost linear, entire. Inflorescences of 1-several-flowered axillary cymes, pedicellate,

pedunculate, or sessile. Flowers on pedicels about as long as calyx or pedicels apparently absent. Sepals equal

or subequal. Corolla conspicuous, blue, or inconspicuous, faded pale bluish-white, rotate, funnelform or sal-

verform, the Umb plicate, mostly subentire, the lobes pilose externally. Stamens with filiform filaments, the

anthers ovate to oblong or linear; pollen 3-colpate, 12-rugate. Ovary 2-locular, each locule 2-ovulate, some-

times 1-locular and 4-ovulate. Styles 2, free or partially united at the base, each style deeply bifid for at least half

its length into long, terete, filiform to subclavate stigmas. Fruits of capsules, globose to ovoid, 4-valved. Seeds

1-4, small, smooth or minutely verrucose.

Species ca. 100; all native to the New World. Two species have become widespread in the Old World.

There are 7 or 8 species in Sonora and 17 in North America.

Selected references .— Austin (1990b), van Ooststroom (1934), Ward (1968).

m UT6 ^ babocillo (fide Rea 1208): mouse ears; huhodam sah’i (Pima Bajo, “laughing bush/herb, ” fide Rea

Herbs ’ Penial; stems prostrate or ascending, 6-50 cm long, loosely appressed pilose and with some hairs

spreading. Leaves ovate, oblong or elliptic to lanceolate, 8-25 mm (exceptionally to 44 mm in extreme SE

^nora mountains) long, 3.5-11 mm wide, the apex obtuse and mucronulate, the base acute to rounded,

sparsely to densely pilose on both surfaces, with strongly and loosely appressed, soft, short, grayish trichomes.

Inflorescences 1 or 2 flowers on filiform peduncles, shorter or longer than the leaves. Flowers on pedicels 2-4

mm lon & short pilose; bracteoles linear-subulate. Sepals lanceolate, 2-2.5 mm long, acuminate, short pilose.

P^ ollas Pale blue or white, rotate, (5-) 7-10 mm wide, filaments 2-3 times as long as the anthers. Ovary glo-

Z 05 * t0 ov «id, glabrous. Capsules 3-4 mm wide, globose, 4-valved, reflexed, glabrous. Seeds 1-4, ovoid, tan to

br own, glabrous.

478

Sonora. — Sonoran desert, desert grassland, coastal and foothills thomscrub, tropical deciduous forest,

oak woodland, and pine-oak forest. Natural and disturbed sites, often somewhat xeric and rocky habitats.

Nearly statewide and one of the most widely distributed members of the family in Sonora. Near sea level-1650

m. Flowering much of the year, depending on soil moisture and temperature.

General distribution. — Arizona, New Mexico, Texas, Florida; Aguascalientes, Baja California Sur, Chi-

apas, Chihuahua, Coahuila, Nuevo Leon, Tamaulipas; Mesoamerica and South America.

Van Ooststroom (1934) recognized fifteen varieties around the world. Sonora, Arizona, New Mexico, and

Texas plants belong to var. angustifolia Torrey. This variety typically has shorter sepals, narrower leaves, less

densely pubescent stems, and tends to have shorter stems than other varieties.

2004-111 01 Mpio Alamos: San Bernardo, G<

on SON 89, foothills thomscrub, 30°19'23"

Ft, 15 Sep-4 Oct, White

ntry winter 1934-35!; Gu

i blue, open 12:00 a.m„ 16 Sep 2000, Reina-G . :

Sierra de los Ajos, 30°44'N, 109°59'W, 1150 m, 22 Apr 1995, Fishbein 2280! Mpio Bavispe: Santa Rosa Canyon, near Bavispe, 3850 ft, 20Jnl

1938, White 611!; Rio Bavispe, Candn de la Petaquilla, 15 Aug 1940, White 3332! Mpio Carbo: 14.6 tni by MEX 15 N of Pesqueira Junction,

1800 ft, 19 Aug 1960, Felger 38451, El Tecolote road (1.2 mi N of El Oasis) W of Mex 15, 27 Aug 1983, R eichenbacher 14711 Mpio Cucuipc

Palm Canyon, 17 mi SE of Magdalena, Van Devender 16 Jul 1977\ Mpio Gen. Plutarco Elias: 16 mi S[E] of Sonoyta on MEX 2, 14 Apr 1963.

Felger 7537! Mpio Guaymas: Nacapule, 25 Feb 1985, Felger 85-248! Mpio HennosiUo: Rocky lower slopes of Sierra de Calena S of Villa dt

Sens. 13 Nov 193 9. Mron.-l c- KiJi.mK >i(m ' Ml >' ■ Ho., kv hillside llcrmo,ilk> 220m.2^W'N. I Id >4 W. 23 Nov |UW & Kuh.uJ;

3754 (MO!); Sierra Seri, 550 m, 2 Feb 1969, Felger 181301 Mpio Hutabampo: Camahuiroa, 23 Nov 1993, Friedman 330-93 ! Mpio La Cob-

b ocher 1891 Mpio Onavas: Onavas, 1 1 Oct 1986, Rea 1208! Mpio San Javier: Cerro Verde to San Javier, Martin & Ferguson 10 Mar 1 990! Mp*>

foothills thomscrub, 1 Sep 2009, Van Devender 2009-649 (USON!).

ep 1996, Van Devender 96-357! Mpio Ures: 6 mi N of Ures, 20 Sep 1!

a, 23 Sep 1977, Goldberg 77-170!; Agua Amarilla, 1000 m, Martin 14 *

Suffrutescent herbs, densely appressed pilose to almost woolly tomentose throughout. Stems few to many,

arising from a woody base, erect to ascending or decumbent, 10-30(-45) cm long. Leaves lanceolate to linear- j

lanceolate, 10-25(-35) mm long, 2.5-6(-14) mm wide, gradually decreasing in size toward apex, the upper I

leaves linear, acute or obtuse apically, attenuate basally; petioles absent or short, sparsely to densely pilose on |

both surfaces, with strongly and loosely appressed, soft, short, grayish trichomes. Inflorescences cymose, 1-3

flowered on slender peduncles usually as long as or longer than the leaves. Flowers on pedicels 3-4(-8) n®>

long, reflexed in fruit; bracteoles linear-subulate, 1.5-3 mm long. Sepals equal, lanceolate, acuminate, 3-33 |

mm long. Corollas rotate to broadly campanulate, blue or blue with white stripes, (10-)12-22 mm wide. Fib- |

ments inserted near the corolla base, 1.5-2 times as long as the linear anthers. Ovary globular, glabrous. Cap-

sules globular, 3.5-4 mm long, reflexed, glabrous. Seeds 2-4, 1-1.25 mm long, tan to brown, glabrous.

Sonora.— Widespread in eastern and central Sonora; Sonoran desert (Plains of Sonora), grassland includ-

ing mesquite grassland, foothills thomscrub, tropical deciduous forest, oak woodland, and pine-oak M*

natural and disturbed sites, often in rocky habitats; 150-ca. 1200 m. Flowering mostly August-December. |

General distribution. — Arizona, New Mexico; Chihuahua; disjunct in Argentina.

This species is easier to recognize in living material than on some herbarium specimens. Some have#'

ognized two varieties that are not distinct.

Selected reference.— Austin (1990b).

Mpio Agua Prieta: Rancho Nuevo, Caj6n Bonito, 2May 1976, Mason 3213b! Mpio Alamos: Mocuzari, La Cruz, 17°13'N, 109 °° 5 . 5 ’W.^^ (

Guayabo (upper) crossing of the Rio Cuchujaqui 3 km NE of Sabinito Sur and 15 km (airltaT) ESE of Alamos, near 27°00'N, 108°4^ I

Felgeretal., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

479

Evolvulus filipes Mart., Flora 24 (2, Beibl.):100. 1841.

tofthes

Annual herbs, the stems erect to ascending, delicate, generally sparsely pubescent. Leaves linear or narrowly

lanceolate, l-2.5(-3) cm long, usually 2-5 mm wide, sessile or ± sessile or sometimes short-petioled on larger

leaves, cuneate basally, acute apically, sparsely pilose on the upper surface, slightly more pubescent on the

lower surface. Inflorescences cymose to solitary, on peduncles usually 1-2.5 cm long or slightly longer. Flow-

ers l(-3) on pedicels to 2.5 mm long, short-pilose; bracteoles lanceolate to subulate, 1-2 mm long. Sepals lan-

ceolate 2-2.5 mm long, glabrous or pubescent and ciliate. Corollas pale blue or white, 3-5 mm long, rotate,

with 5 sericeous bands outside, filaments ca. 2 mm long. Ovary globose, glabrous. Capsules globose to ovoid,

3-4 mm long, reflexed, glabrous, brown. Seeds 1-4, smooth, dark brown to b

Sonora.— Several specimens identified to this species are from the eastei

ciduous forest and oak woodland.

General distribution.— C oahuila, Durango, Michoacan, Sinaloa, Veracruz; Mesoamerica; South America.

This species is weakly differentiated from E. alsinoides. In South America they appear to be distinctive,

while in Mexico they are not always separable. The Sonora specimens have the appearance of first season

plants of E. alsinoides. Perhaps those called E. filipes in part (or all) of the range are nothing more than depau-

perate or first season E. alsinoides, but there are insufficient data to confirm or deny that suspicion and we re-

luctantly retain them as separate until more detailed studies are made.

n E of Alamos, 2r0'20"N, 108°4710"W, 250 m, Martin 16 Mar 1989 (FTG-FAU!). Mpio Cucurpe:

), Toolin 453-C (2 sheets, AR1Z 220685!, ARIZ 221490)!

Schult., Syst. Veg. 6:198. 1820, New name

E.-USA Missouri: Rapid River, “On the banks of the Missour., v F .<

tly destroyed during the Second World War, although van Ooststro

fwlwdus oeophlu Greene, Leafl. Hot. Observ. Crit. 1(U):151. 1905. TnK USA New Mexico: Sima [Co.l: S e^oftheBladt Rang®. *

miW of Hillsboro, dry hills, 5500 ft, 16 Aug 1904, Metcalfe 1228 (u-cronre designated here: US!; isotypes: MO. NMC. in

Suffrutescent herbs, the stems several, erect to ascending, 10-15 cm long, densely spreading-pilose with an

indumentum of ferrugineous, brown, fulvous or gray color. Leaves linear-oblong, narrow-1;

row-oblanceolate or rarely oblong, 8-20 mm long, 1.5-5 mm wide, attenuate basally, acute tc

densely pilose on both surfaces; petioles short or absent. Inflorescences solitary, m axils

fength of stem; peduncles short or absent, flowers on pedicels 3-4 mm long, becoming reflexed in fruit; brac-

teoles subulate, 1-4 mm long. Sepals lanceolate to narrowly lanceolate, long-acuminate, 4-5 mm long, sprea -

ing villose. Corollas rotate to broadly campanulate, 8-12 mm wide, subentire, purple o

Journal of the Botanical Research Institute of Texas 6(2}

long, oblong, basally auriculate; filaments twice as long as the anthers. Ovary subglobose, glabrous. Capsules

ovoid, about as long as sepals, reflexed, glabrous. Seeds (1) 2, brown, smooth.

This species grows in the mountains of southeastern Arizona near the Mexican border (Austin 1991,

1998b) and probably will be found in northeastern Sonora in oak woodland and pine-oak forest; 820-2450 m.

Flowering April-September.

General distribution. — Arizona, New Mexico, and Texas to Montana, North Dakota, Illinois, Arkansas,

disjunct in Tennessee; Chihuahua, Coahuila.

i B.L. Rob., Proc. Amer. Arts 29:320. 1894. i

Y!; isolectotypes: GOET!, K!, PH!). MEXICO, t

:n). MEXICO: Valley of Mexico at Santa Ffi, 5 Jul 1865-66, Bourgeau 323 (synty

Perennial herbs; stems few or several from a woody perpendicular root, prostrate, 10-20 cm long, sericeo-vil-

lose, with fulvous, grayish or whitish trichomes, glabrescent. Leaves distichous, more or less imbricate, at

right angles to the stems or somewhat reflexed, broadly-ovate to orbicular or sometimes broader than long,

9-16 mm long, 7-16 mm wide, apically rounded to somewhat emarginate, basally truncate to cordate or

rounded, covered with appressed silky-villous, light-brown to grayish trichomes below, green and glabrous

long. Flowers 1(2) on pedicels 2-3 mm long, appressed-villose; bracteoles oblong to linear-oblong, to 3.5 mm

long. Sepals ovate-oblong to ovate, 3-4.5 mm long, acute, appressed-villose. Corollas white, outer surfaces of

corollas in bud and midpetaline conspicuously tawny sericeous-hirsute, rotate to broadly funnelform, tube

short, the limb 10-12 mm wide. Anthers 1-2 mm long, filaments 3-4 times as long as the anthers. Capsules

globose, 4-valved, 3-5 mm long, glabrous. Seeds 1-4, brown, glabrous.

Sonora. — Pine-oak forest in east-central Sonora; 1550 m. Flowering at least in September.

General distribution. — Chihuahua, Distrito Federal, Durango, Hidalgo, Jalisco, Michoacan, Nayarit,

Nuevo Leon, Oaxaca, Pubela, San Luis Potosl, Veracruz.

Mpio Yecora: Ydcora, 28°23%'N, 108°54Vi'W, 1550 m, open pine forest, occasional, 7 Sep 1995, Fishbein 2481 !; Yecora, near baseball fieldand

cemetery, 28°22'25"N, 108°56’W, 1540 m, common on clay flats, 5 Sep 1996, Reina-G. 96-377 (FTG-FAU!).

Evolvulus rotundifolius (S. Watson) Hallier f., Bot. Jahrb. Syst. 16:530. 1893. Type: MEXICO. Chihuahua: damp places,

Perennial herbs; stems several from a woody root, prostrate, 10-15 cm long, sericeo-villose, glabrescent to-

ward the base. Leaves distichous but not imbricate, at right angles to the stem, ovate to elliptic, sometimes

broad-ovate, 13-22 mm long, 8-12 mm wide, apically obtuse to somewhat emarginate, basally rounded to

cordate, appressed sericeo-villose below, glabrous above, the petioles 1-2 mm long. Inflorescences axillary,

sessile, and 1-flowered. Flowers sessile; bracteoles linear-oblong, to 6 mm long. Sepals ovate-oblong to oblong,

4-5 mm long, acute, appressed-villose. Corollas blue with a white throat and midpetaline stripes, rotate to

broadly funnelform, the tube short, the limb 10-13 mm wide. Filaments 2-4 mm long; anthers 1-2 mm long

Capsules globose, 4-5 mm long, 4-valved, glabrous. Seeds 1-4, brown, glabrous.

Sonora.— East-central and southeastern Sonora near the Chihuahua border; arroyos and open areas in

oak woodland, pine-oak forest, and mixed conifer forest (with Abies durangensis, etc.); 1600-2100 m. Flower-

ing August-September;

General distribution. — Chihuahua, and Durango; reported but not verified for Aguascalientes and

Queretaro.

!I Aug 1991 ! Mpio Yecora: Mesa El Campaner

-FTG!); El Kipor (Quipur), Arroyo El KIpor, p

sahuaribo), 27°19'N, 108°40'W, 1600 m, 21 Aug 1991, i

el Salto, 2040 m, arroyo margin in pine-oak forest will

!; N end of Mesa Campanero at head of Bart

e-oak forest, 28°24'25"N, 108°35'40"W, 1

i in tropical deciduous forest at about 500

central Sonora in oak woodland, ca. 940 m, and southe;

m. Flowering September-November.

General distribution . — Sinaloa southward and perhaps in all Mexican states; southeastern United States;

Mesoamerica; Colombia, Venezuela, Guyanas, Ecuador, Peru, Brasil, Argentina; Caribbean. Often cultivated;

native to tropical America. Flowering September-May.

Apparently first recorded in the New World by Oviedo (1526). This plant was probably first carried

around the world for its medicinal seeds. Later it was spread for the nocturnal, fragrant flowers. Determining

the region of nativity of this species within the New World is particularly difficult since it was carried from at

least Cuba around the world in the early 1500s by the Spanish and probably the Portuguese. However, the

center of diversity is tropical North America (McDonald 1993a). Typically, the plants are associated with wet-

lands, the seeds being distributed by water. Seeds arrive on the coasts of the British Isles with regularity, and

surprisingly a few of them are viable and germinate. The occurrence in southern Sonora may be part of the

native range, or may result from cultivated plants that have escaped.

Perennial herbs from a tuberous root, the stems woody below, retrorsely hispid. Leaves 8-10 cm long, almost

as wide, cordate to broadly ovate, entire or 3-lobed, remotely appressed sericeous. Inflorescences dichasial,

axillary. Flowers 1-4 on peduncles 10-12 mm long. Sepals 2.5-4 cm long, ovate, apically acuminate to obtuse,

and becoming spatulate-attenuate with age, the outer ones appressed sericeous, the inner ones glabrous on the

margins, sericeous on the dorsal regions. Corollas funnelform, white, 3-4 cm long, pubescent on the outer

surface. Capsules 8-10 mm long and wide, almost globose, brown, glabrous. Seeds 1-4.

Sonora . — Known in the state from a single record in pine-oak forest, 1220 m. Flowering in September.

General distribution.— Sinaloa, Guerrero, Edo. Mexico, especially tropical deciduous forests. The occur-

rence of this species in pine-oak woodland in Sonora seems to be an anomaly as compared with its general

distribution.

Y«ora: Rio Maycoba at MEX 16 (20.5 km W of Maycoba, 28.6 km E of Yecora), 28°22'15"N, 108°45'30” W, common, to 2 m in shrubs.

Ipomoea ancisa House. Ann. N.Y. Acad. Sci. 18:187. 1908. im MEXIC O Chihuahua: 22-24 Aug I8<w. tor..,. •

Rom ®IA DEM SIERRA

Ere «, suffrutescent shrubby perennials, 1-1.5 (2) m tall, the stems erect or ascending, glabrous. (John Palting,

Personal communication 28 September 201 1 , found the roots of a few plants to be deep and relatively thick but

not ^berous.) Leaves ovate in outline, 3.5-11 cm long, 2-5 cm wide, irregularly pinnately divided into 6-9,

obtuse-tipped divisions, the lobes linear to filiform, entire to irregularly toothed or lobed, 1-7.5 (8.7) cm long,

1-2 10111 wide, glabrous, the petioles 7-15 mm long. Inflorescences of solitary (rarely 2) flowers. Flowers on

Muncies 3-12 cm long, the pedicels 1-1.5 cm long, accrescent and recurved in fruit; fruiting peduncles and

Pedicels becoming somewhat woody; peduncles and pedicels with squamose, caducous bracts. Sepals un-

T 31 ’ ^ outer slightly shorter than inner, broadly elliptic to ovate, 6-10 mm long, 4-8 mm wide, smooth,

obtUsc or truncate, rarely acute, the margins scarious. Corollas funnelform, 5-12 cm long, white to pale laven-

der ’ the tube whitish, glabrous, the limb 4-7 cm wide. Capsules pear-shaped to ovoid, 1.5-1.8 cm long, 1.1-1.7

001 wide. Seeds 4, 7-10 mm long, ellilpsoid, gray-brown, puberulent.

Sonora . — A narrow endemic in mountains in eastern Sonora and western Chib

a, 1220-1500 mir

Sonora and to ca. 2000+ m in adjacent Chihuahua. Juniper-oak woodland and pine-oak forest. Flowers and

fruits July-September. This large morning glory, sometimes locally common, can be a spectacular sight to-

wards the end of the summer rainy season with its many, large flowers open in early morning among the bright

green, feathery foliage. John Pairing (personal communication 30 Sep 2011) and Van Devender found it abun-

dant in the Sierra de Bacadehuachi in early Sep 2011. Pairing wrote, “The stout plants in the sun were spectacu-

lar in bloom (100 flowers or more). Upon blooming the largest plants are a round mound about 3 feet high.

They become more decumbent in the shade and bloom less. This is definitely a plant with high ornamental

value.” The flowers produce jasmine-like fragrance.

Gentry (1942: 213) reported the herbage of Ipomoea ancisa in the upper Rio Mayo region of Sonora and

adjacent Chihuahua “is decocted and drunk for stomach ailments.”

Both I. ancisa and I. sescossiana are erect, shrubby perennials with pinnately compound leaves bearing

very slender lobes, and are xenogamous with large bee-pollinated flowers (McDonald 2001). I. sescossiana oc-

curs in Chihuahua and might be found in easternmost Sonora. Some specimens of I. ancisa from Sonora have

been incorrectly attributed to I. sescossiana. The much smaller leaves and darker flowers readily distinguish!

sing, 28°22.5'

i. White 3672!; C

-oad to Moris (Chihuahua), 28°19'40"N, 108°39’20"W, 1500 m, 26 Sep 1997, Van Dozen-

VIEX 16, grassland on rocky slope in oak woodland, 28°22'52"N, 108°49'33"N, 1410m,

Mpiojanc

e: MEXICO. Guerrero: Inter

Palo santd, palo blanco; tree morning glory; jOtuguo (Mayo)

Trees 3-15 m tall, the trunk to 50 cm diameter, the bark gray, whitish or yellowish, stems with abundant latex,

tomentose when young with trichomes 0.1-2.5 mm long, becoming glabrescent. Leaves: blades often 9-19 cm

long, 6-9 cm wide, ovate to lanceolate, often glabrescent or pubescent below (especially among var. pachyleuta)

(velvety below particularly in the southern end of the range near Mexico City), the apex acuminate, the base

cordate, with trichomes longer that those on the branches; petioles of larger leaves often 6-8 cm long. Inflow

cences terminal or axillary, monochasial, racemose, forming compound-cymose clusters. Flowers in 1 (2) p®

cyme. Sepals 6-14 mm long, ovate to rarely orbicular, more or less equal, tomentose, the apex obtuse to obtuse-

mucronate. Corollas 4-6 cm long, funnelform, tomentose at least on the lobes, white, with green, yellowish, or

puplish within the tube. Capsules 17-25 mm long, 4-valvate, brown, glabrous. Seeds 1-4, 10-16 mm long-

brown, pilose on the margins with trichomes 10-15 mm long.

Howard Scott Gentry (1942: 213) wrote, “A spectacular tree 7-10 m high, with smooth, white-gray bark

like the hide of a hippopotamus. It flowers in winter when leafless, holding a high, thin spread of white corollas

like stars against the morning sky. These stars soon fall upon the ground, where the deer eat them. With the

summer rains, the tree forms dense foliage, which on the characteristically recurved branches is somewhat

plumelike, especially from a distance.” The flowers an important early spring food source for migrating hunt'

mingbirds (Martin et al. 1998).

Sonora . — Sonoran desert, thornscrub, tropical deciduous forest, and lower oak and pine zones. These

trees generally leaf out with the first summer-monsoon rains and fall away as the rains cease in fall, usually

around September and October. Flowering during cooler months, when the trees are essentially leaflet

mostly November-April and sometimes with a few flowers into May; near sea level-1100 m. The flowers ope°

487

in the late afternoon or early evening and may remain open much or all of the day during cooler weather. Many

flowers, however, tend to fall before sunrise.

General distribution . — Mexico at least in Chiapas, Chihuahua, Guerrero, Jalisco, Michoacan, Morelos,

Oaxaca, Puebla, Queretaro, and Sinaloa.

lpomoea arborescens can be confused with 1. pauciflora M. Martens & Galeotti of Mesoamerica. Farther

south in its range, one may distinguish 1. arborescens by the pubescent sepal and leaves that are silky pubescent

below. However, in Sonora I. arborescens does not have those silky leaves. That paucity of indument led Gentry

Sinaloa, which may warrant recognition as species. In Sonora the varieties apparently are allopatric, although

in the vicinity of Aduana, near Alamos, they were observed growing intermixed in a disturbed tropical decidu-

ous forest habitat.

lpomoea arborescens has been used medicinally by the Guarijios and Mayos including a remedy for snake-

bite and to alleviate the pain of a scorpion sting, and also to treat toothache. The wood is burned to produce

smoke to keep away mosquitoes. The soft, spongy, and moisture-rich wood is used as emergency fodder for

cattle. It is chopped up for cattle feed (Gentry 1942, 1963; Yetman and Felger 2002; Yetman and Van Devender

2002). The Tepehuan used the wood in bows for violins (Pennington 1969). They also used the light-weight

wood for foreshafts in the composite arrow.

Selected references.— Austin et al. (2005), Felger et al. (2001), Gentry (1942), Martin et al. (1998), Turner et

Gray (1887). Type: MEXICO. Sonora: Alamos, N

RUd SANTO, PALO BLANCO', TREE MORNING GLORY; JOTUGUO (Mayo)

Sonora. — This variant is widespread through the range of the species (except where var. pachyleuta o

the Sonoran desert, coastal and foothills thomscrub, tropical deciduous forest, and sometimes in 1<

zones; near sea level to ca. 1090 m. The flowers in Sonora are visited by bees, hawkmoths, and humm

and south of Sonora by bats.

General distribution.— Sonora southward to Chiapas including Sinaloa, Guerrero, Michoacan,

°axaca, and Puebla.

r — „ J 9 vuy M 6 — 5! M P io Cok,rada: 1:

ms of Sonora desertscnib, 28°3717"N, 110°05'30"W, 5

"«-G. -2008-147! Mpio Cncurpe: 14.2 km NW of Sinoquipe on road to Cucurpe, 30.19222°N, 1

[ 5 - Mpio Gnaymas; Microondas Avispas, Sierra Baviso, S end of Sierra Libre, 28°29'N, lll o 02'V

^t-.CenoHVigfc

l5 ’ Plains of Sonora d,

mscnib, 29°29’09"N, 110°59'55"W, 5

it, open all day, 27 Dec 2000, R(

liN of Hermosillo, 900 ft, 23 Feb IS

r) of Melchor Ocampo, 26.57500°N,

109.3000°W, n

c 1993, Friedman 343-93 (ASU!). Mpio M

s, 5 Jul 1971, Hastings 7.

m, very common 3-10 m tall tree, flowers white with a little magenta in throat, 17 Feb 1993, Rrina-G. 97-118 (FAU-FTG!). Mpio Opodept:

Publ. 527:213. 1942.1

r. pachylutea Gentry, Publ. Carnegie Inst. Wash.

Palo santo amarillo; ToamO, tochiguO (Guarijfo), jOtuguo (Mayo)

Tree 7-8 m tall. Petioles, 3-7 cm long; blades often 8-18 cm long. This variety is distinct from the tautonymic

1. arborescens farther south in its range, and may warrant taxonomic revision. The wood is harder than the

lowland I. arborescens. See Austin et al. (2005) for details.

Tropical deciduous forest, oak woodland, pine-oak forest; 400-1200 m.

Ipomoea aristolochiifolia G. Don, Gen. Hist. 4:277. 1838. Type VENEZUELA: Humboldt &Bonpland 679 (isotype: B!, bar-

code: B-W-03661-01-0). Convolvulus aristolochiifolius Kunth, Nov. Gen. Sp. (quarto ed.) 3:102. 1819, not Mill. (1768).

Ipomoea tuerckheimii Vatke ex Donn.-Sm., Bot. Gaz. 40:8. 1905. Type: GUATEMALA: Alta Verapaz, von Turckheim 386 (isotype: US!).

Ipomoea austin-smithii Standi., Publ. Field Mus. Nat. Hist, Bot. Ser. 18:566. 1938. Type: COSTA RICA:. Brenes 16899 (holotype: F!).

Annual herbs, the stems twining, delicate, 1-3 m long, glabrescent to puberulent. Leaves 4-8(-10) cm long,

cordate to ovate-cordate, entire, more or less glabrous. Inflorescences cymose, with flowers on long peduncles

that pass between the basal lobes of the leaves. Flowers 1-6. Sepals 3-5 mm long, ovate, more or less equal-

rugose on outer surfaces. Corollas 20-25 mm long, campanulate, the throat pale purple, the tube cream with-

out, glabrous. Capsules 9-10 mm long, ovoid, brown, glabrous. Seeds 1-4, 4-5 mm long, black to brown, p»'

Recognized by the small pale flowers on peduncles that pass between the basal lobes of the leaves. Also, «

is distinctive by having rugose sepals and small, lavender corollas.

Sonora . — Eastern Sonora, in Chihuahuan desert, oak woodland, and tropical deciduous forest; 180-129®

m. Flowering September-October.

General distribution. — Arizona; Baja California Sur, (probably in Chihuahua but no records found), Coa-

huila, Guerrero, Jalisco, Edo. Mexico, Michoacan, Morelos, Oaxaca, Sinaloa, Veracruz; Mesoamerica; Colom-

bia, Venezuela, Ecuador, Peru, Bolivia, Brasil.

Felger et al., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

Annual herbs, the stems low-climbing, glabrous. Leaves orbicular-ovate in outline, 3-8 cm long, 1.5-8.5 cm

wide, glabrous, with or without glandular dots, basally cordate, deeply 5-7 lobed, the lobes lanceolate and nar-

rowed toward base, acute to acuminate; petioles 1-5.5 cm long. Inflorescences cymose, axillary. Flowers 1-3

on peduncles 2-6 cm long, glabrous or remotely appressed hairy, the pedicels 4-5 mm long, erect in fruit;

bracts foliaceous, elliptic to linear, 5-6 mm long. Sepals ± equal, 10-12 mm long, 1-2 mm wide, lanceolate,

with elongate narrowly linear tips typically erect to spreading in flower, reflexed in fruit, the bases slightly di-

lated, hispid-pilose. Corollas funnelform, 1.6-2(-2.5) cm long, glabrous, blue to light rosy-purple or white, the

limb 1.8-2 cm wide, the throat yellow, glabrous. Capsules glabrous, 8-9 mm long, rounded, apiculate,

2-3-locular. Seeds 1-6, 4-5 mm long, pyriform, dark brown to black, appressed pubescent.

Sonora . — Desert grassland, thornscrub, tropical deciduous forest, and oak woodland; 200-1300+ m.

Flowering at least August-September.

General distribution.— Arizona, New Mexico, Texas; Baja California Sur, Sinaloa, disjunct to Oaxaca.

Floweringjuly-December.

In many ways this species resembles L hederacea, which may grow with it, but I. barbatisepala is more

delicate, more of the leaves are lobed, and the corollas are typically smaller. The two have sepals that are almost

identical, but the pubescence is mostly near the base in I. hederacea and extends all the way to the tip in I. bar-

batisepala.

^ Arn^o MpUo, near Colonia Oaxaca, 30 JuU938, mite 746! Mpio inmris: 10 tar NE of Imunson MEX 2, 1300 m, desert

white, sepals drying purple, open 7:20a.m., 29 Aug 2000, Van Devender 2000-463 (USON!). Mpio Yecora: 1.9 km W of Tepoca, 675 m, 22 Sep

Ipomoea batatas (L.) Lam., Tabl. EncycL 1:465. 1791. Type: INDIA: herb. Linnaeus (LINN-77 .5 S'). Convolvulus batatas l

H. 154. 1753.

Batatas edulis (Thunb.) Choisy, Mem. Soc. Phys. Genfcve 6:435. 1834. Type: JAPAN: Thunberg (holotype: UPS!).

Perennial herbs, the stems twining or prostrate, to 5 m or more, pubescent with appressed to erect trichomes,

less often glabrous. Leaves 5-10 cm long, broadly ovate to cordate, entire to dentate or 5-7-lobed, the apex

acuminate, glabrous to pubescent. Inflorescences, axillary, in monochasia or dichasia. Flowers (l-)3-°o. Se-

Pak unequal, 8-15 mm long, oblong, the two outer shorter than inner and abruptly acuminate, glabrous or

Pubescent. Corollas (3-H-7 cm long, funnelform, lavender or sometimes white, with a tube darker within.

Pubescent within near the base. Capsules infrequent, 4-5 mm long, rounded, brown, pubescent or glabrous,

^eds 1-4, 3-4 mm ] on g tan to brown, glabrous. 2 n = 60, 84, 90.

Sonora. — -Cultivated in gardens and occasional found in the southern margin of the Sonoran desert,

C ° asta l an d foothills thornscrub, and tropical deciduous forest, especially in disturbed areas in densely vege-

tated ar eas such as washes. Flowering at least March, April, September, and October.

General distribution.— Mexico; Central America; Argentina, Bolivia, Brasil, Chile, Colombia, Ecuador,

S of Tepache, 560 m, 25 Apr 1971, Turner 7 i-58! Mpio Yecora: Arroyo San Nicolas, E

Nicolis sobre el camino a Yecora (Sonora 16), 29.452°N, 109.15806°W, hierba trepador

95-266 (USONOl Mpio Ures: 10 km W-SW ofMazocahui on SON 89, foothills thomscr

11Q°10'20"W, 715 m, 29 Nov 2000, Reina-G. 2000-879!

a, perenne, bracteas y flor lila, 7 May 1995, Reina-G.

■ub/tropical deciduous forest transition, 29 S 30'02 ,, N;

*IpomOea cairica (L.) Sweet, Hort. Brit, 2:287- 1826. Convolvulus cairicus L., Syst. Nat. ed. 10, 922. 1759. “Convolvulus Aegyp-

tius" Vesling en Alpinio, Pi. Aegy pti 73 & 74 (1640) (uscTomd).

Perennial vines, the stems to 5 m or longer, twining but also lying on the ground, glabrous. Leaves 3-10 cm

tong, 3-10 cm wide, glabrous, ovate to orbicular, palmately divided to the base into 5-lobes, these lanceolate or

ovate-lanceolate to ovate or elliptic, the basal segments acuminate, the two basal segments generally lobulate

or otherwise parted, mostly with pseudostipules (small leaves from the axillary buds at the base of the petiole),

acute to obtuse at the apex. Inflorescences cymose. Flowers 1 to few. Sepals 4-6.5 mm long, more or less

equal, or the outer sepals slightly shorter, ovate, obtuse to acute, the inner sepals wider, obtuse, glabrous but

frequently somewhat tuberculate. Corollas 4.5-6 cm long, rarely shorter, funnelform, usually blue-purple or

white, with a reddish-purple throat, glabrous. Capsules 1-1 .2 cm long, more or less globose, straw-colored or

brown, glabrous. Seeds 1-4, globose, 5-6 mm long, brown, densely short tomentose or at times also with long

silky trichomes on the margins.

Sonora.— Known from one collection from 1939. Due to wide naturalization in other parts of the world

the species is included. Future searching may reveal additional records for Sonora.

General distribution.— Naturalized in Alabama, Florida, Louisiana; cultivated in Arkansas, California,

Texas?; Mexico (at least in Oaxaca); South America; West Indies; Africa; Asia; Australia. Cultivated in the trop-

ics and subtropics around the world; nativity uncertain, perhaps Africa.

Ipomoea capillacea (Kunth) G. Don, Gen. Syst. 4:267. 1837. Type: COLOMBIA: between San Miguel and Rio Pure*, Nov

1801 Hum bold i Hon r h„n I s n. <n .lOTYFE P. microfiche seen, photos I !. Gil! .•T,n • P<2>!) ivnu.lvnlio u, r ,IUcus Kunth. Nov

Gen. Sp. PI. 3:97.1819.

Ipomoea murieata Cav., Icon. 5:52. 1799, not Jacquin (1798). Type: MEXICO. Guanajuato: Oct, Cavan Hies i.n. (holotwe: MA!; isotype:

MA!). Ipomoea muricatisepala Matuda, Ann. Inst. Biol. Mex. 34:124. 1964. New name for I. murieata Cav.

Perennial herbs with erect to ascending glabrous stems, from an underground elongate tuber. Leaves with

blades sessile, incised and appearing compound, the segments usually 5, filiform, 5-15 mm long. Inflores-

cences axillary, solitary. Flowers on peduncles 5-7 mm long, the pedicels 3-6 mm long, reflexed in fruit. 5e-

Pak ± equal, 5-6 mm long, the outer one 2 mm wide, the inner ones 3 mm wide, acute to obtuse and mucronate

apically, the outer sepals elliptic to oblong, the inner sepals ovate, muricate-tuberculate at least on the midvem.

Corolla, 3-4 cm long, funnelform, lavender .0 reddish purple, .he limb 2-2,5 cm wide, glabrous Xapsules *

globose, glabrous, 4-5 mm wide, apiculum 1 mm long. Seeds 1-4, 3 mm long, ovoid, black to dark brown,

shortly erect hispid.

Sonora. — Oak woodland and pine-oak forest in eastern Sonora; 1100-2100 (to 2500 m in nearby SW

Chihuahua). Flowering August-October.

General distribution.— Arizona, New Mexico, Texas; Baja California (norte) and Sur, Chihuahua, Coa-

buila, and southward to Puebla; Central America; South America.

See comments under I. murieata and also I. plummerae for potentially confusing species.

V. AUmos; Sierra . A r 7n All „ 1Q o 4 steinmonn 94-741; Vallecito, 3 km SW of Santa Barbara, 1100 m, Martin 4 Oct 1990 .

i, 1372-1585 m, south

6 Aug 2003. K amp 445 (COCHISE!). Mpio

*“ anJos * fountains, 3 Aug 1893, Mearm 1628 (US, not seen, cited by McDonald 1995). M P .o Nacoxar. de

' eras ' 20 Aug 1941, While 4059 (Gil specimen not relocated, cited by McDonald 1995). Mpio Yecora: Arroyo Otro

"e. Van Devender 95-840!; Mesa de los Corona

' *'20 m , 28°23'49”N, 108°54’48"W, 7 S

^upanero, 28°19’30"N. 109°01'40'W, 2100 n

^ ^Barranca El Salto, pine-oak forest, 2

j 1995, flowers ]

flowers pink, 1 Sep 2000, Van Devende

00 m, 6 Sep 1996, Van Devender 96-396!

Felgeretal., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

493

Ipomoca chilopsidis Standi, Publ. Field. Mus.

purple throat, singularly of the high and arid crags, 1

er. 17:206- 1937. Type: MEXICO. Cmh

n high, few and irregularly branched, flo'

2391 (hOlotyfe: F!; isotypes: AR1Z 76041!, 2

Multiple-stem small trees and large shrubs 2-5 m tall, the stems broadly ridged on drying, glabrous. Leaves

mostly l0-17.5(-20) cm long, 0.5-1.3 cm wide, with 18-30 pairs of secondary veins, entire, linear, apically

acute, basally cuneate, glabrous; petiole 0.5-1.2 cm long. Inflorescences monochasial, terminal on reduced

branches. Flowers l(-3) on peduncles 0.4-2.2 cm long, glabrous except inner surfaces of sepals. Sepals 12-16

mm long, 7-9 mm wide, ovate, about equal or the outer ones slightly shorter than the inner ones, apically ob-

tuse to acute, the inner surfaces short-pubescent. Corollas 8-9.5 cm long, 8-9 cm wide, funnelform, white

with a purple throat. Capsules 18-22 mm long, conic, brown, glabrous. Seeds often 4, 10-15 mm long, oblong,

brown, woolly with long trichomes on the ventral margins.

Sonora .— High and arid rock ridges and on indurated ash in woodland, often with Acacia pennatula,

Dodonaeaviscosa, and Quercus chihuahuensis ; 1000-1300 m. Flowering documented May-November.

General distribution . — Oak woodland in southeastern Sonora and southwestern Chihuahua; 1000-1800 m.

The leaf shape of this shrub makes it unique among the Ipomoea in the Americas. As the species name

indicates, the leaves resemble those of Chilopsis linearis (Bignoniaceae).

loom, 24 Aug 1992, SU

tensis, 1100 m, large wl

t, 27 D 58 , 20"N, 109°06'30"W,

ridge, ca. 5000 ft, Qjun 1963, Friger 8078b!; Barranca de de Batopilas, 5 mi S of Quirare, rocky

:n CF39 (UCR!); La Bufa, canyon Rio Batopilas, short tree forest, barancan oak forest, 3300-

d new settlement, 10 Aug 1971, Bye 1977 (UCR!); Sierra Madre Occidental between Creel and

ior near mouth of creek called Arroyo San Fernando, 3330-4180 ft. 27 Jun 1982, Siplivinsfey

Bound. 149. 1859. type: l

Herbs, annual, from a slender taproot; stems erect at first, in age trailing or twining at the tips, 2-3 m long,

glabrous. Leaves with blades ± sessile or on petioles 1-3 cm long, deeply palmately divided with the lateral

divisions two-cleft (pedatisect), the segments 5-9, linear or linear-lanceolate, 7-25 mm long, glabrous. Inflo-

rescences mostly solitary, axillary. Flowers on peduncles l-3(-7) cm long, the pedicels 15-25 mm long, erect

in fruit. Sepals slightly unequal, the outer ones 3-5 mm long, 1-2 mm wide, the inner ones 4-6 mm long, 2-3

™n wide, oblong-lanceolate, acute, mucronulate, scarious margined, at least the inner slightly rugose along

the veins. Corollas 10-12 mm long 5-10(14) mm wide, pale lavender to pink, usually with a white throat.

Capsule* ± globose to ellipsoid-globose, 4-5 mm wide, with a 1-2 mm caducous apiculum, tan, glabrous.

Seeds 3 or 4, ■

i long, ovoid, black, glabrous,

land, coastal

Sonora.— Southern and eastern margins of the Sonoran desert, Chihuahuan desert, gra

and foothi U thomscrub, tropical deciduous forest, oak woodland, and pine-oak forest; 50-1550+ m. Flowering

August-November.

Cncrd distribution. — Arizona, New Mexico, Texas, Baja California Sur, Chihuahua, Coahuila, Nuevo

U6n ’ Tar nauli pas south to Chiapas; introduced into South America.

This widespread species is self-fertilizing (McDonald et al. 2011); indicators are the annual habit and the

inconspicuous small flowers.

495

Felger et al., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

mi S of Esqueda, 22 Jul I960, Felger 4045! Mpio Guaymas: Canon Nacapule, 18 Nov 1985, Felger 85-1310! Mpio Hennosillo: Cafldn Las

Chivas, Sierra Libre, al Sur de la ciudad de Hermosillo, por la Carr. Fed. 15, 28.583°N, 110.96528°W, 8 Oct 1997, Burquez 97-501 (USON!).

Sep 2006, Boyle 7840! Mpio Yecora: Near Los Vallecitos, 14.7 km E ofYecora on MEX 16, pine-oak forest, 28°22'38"N, 108°50'03"W, 1470 m,

10 Sep 1996, Van Devender 96-521 !

i. Type: MEXICO. Hidalgo: rocky hills. Lens Static

1905. Pringle 10034 U

Perennials from a large, thickened root, the steins 1-1.5 m long, twining, finely white-pubescent, becoming

glabrous with age. Leaves 3-5 cm long, 3-4 cm wide, entire or trilobate, or almost 5-lobed, white-pilose, some-

what sagittate to cordate, the apex acute, mucronulate; petioles 2-3.5 cm long, white-pilose. Inflorescences

with axillary peduncles 3-20 cm long, white-pilose. Flowers solitary on slender pedicles 8-10 mm long, pi-

lose; bracts 2-4 mm long, linear, pilose. Sepals 10-15 mm long, 7-8 mm wide at the base, herbaceous, more or

less equal, pubescent, triangular-ovate, acute (not attenuate), white-pilose. Corollas blue to bluish, 5 cm long,

4-5 cm wide, the tube white. Capsules about as long as the calyx, glabrous, 5-valved, 5-locular, often 10-seed-

ed, the 5 valves often bifid; pericarp chartaceo-ligneus, brownish. Seeds 1-potentially 10, 4 mm long, discoid,

black, with short erect trichomes.

Sonora.— East-central part of the state in pine-oak forest, 1300 m. Flowering August-September.

General distribution.— Hidalgo, Michoacan, Edo. Mexico, Sinaloa, Zacatecas; reported from Chiapas (Nel-

W49, NY and GH, neither seen).

This perennial species is similar to and might be confused with annual I. purpurea. The sepals of I. deca-

sperma lack the setose trichomes of I. purpurea ; leaves of I purpurea, while variable, are not sagittate as in I.

decasperma.

Mpio Yecora: Tributary of Arroyo de Pilar

magenta ’ 2 2000, Reina-G. 2 000-622, 2000-627 (USON!).

Ipomoea hederacea Jacq., Collect. Bot. 1:124, pi. 36, 1787 [title page 1786 1^' “^ IERICAS ” (ac (

“g to the Code) since Jacquin did not mention the Dillenius plate.

Ipotnoea desertorum.House, Ann. New York Acad. Sci. 18:203. 1906. Type: U.S.A. Arizona: Thurber 29 (holotype: NY!,

Ipomoea hederacea var. integriuscula A. Gray, Syn. FI. N. Amer. ed. 2, 2:433. 1886. Type: U.S.A: East Florida, Curtiss (h

T *°MPiUO, TRQMPILLO AZUL, TMM1LLO MORADO ; IVY-LEAF MORNING-GLORY; HEHE QUVJAM ‘PLANT THAT-CURLS-AROUND-.T,’

mm] W K spun (like a top)’ (Seri)

Annual herbs, the stems twining, often 2-3+ m long, densely to sparsely pubescent throughout. Leaves with

^des ovate to * orbicular, (2-) 3.5-13 cm long and about as wide, entire to 3- or 5-lobed, basally cordate, the

a P ica % acute to acuminate, pubescent; petioles to 12 cm long, rarely longer. Inflorescences cymose.

Flowers l-3(-6), on peduncles 5-10 cm long, the pedicels 3-7 mm long, erect in fruit; bracts foliaceous elhp-

lc to lanceolate, 5-8 mm long. Sepals ± equal, 12-25 mm long, 4-5 mm wide, herbaceous, lanceolate, abrupt-

497

Hobs, annual, the stems twining or prostrate, 2-8 m long, glabrous. Leaves petiolate, 1-4 cm long and wide,

5-foliolate, the lobules lanceolate to linear-lanceolate, sessile, apically acute, glabrous. Inflorescences axillary

and terminal, flowers usually solitary, with filiform peduncles as long as or longer than the petioles, often

spirally twirled. Sepals ± equal, 5-7 mm long, ovate to oval, apically obtuse to rounded, glabrous. Corollas

1.8-3 cm long, funnelform, glabrous, rosy (or sometimes white) with the interior of the tube red-violet. Cap-

sules 8-10 mm long, globose, brown, glabrous. Seeds 1-4, 4-6 mm long, ovoid, brown, finely pubescent.

Sonora .— Coastal thomscrub of the coastal plain in southwest part of the state, apparently not common.

Margins of temporarily flooded or swampy places including roadside; below ca. 50 m. flowering September-

October.

General distribution.— Texas, SE USA; Mesoamerica; Argentina, Brasil, Ecuador, Paraguay, Peru;

pantropical.

Although no one knows where this species is native, it shares no obvious relatives in the New World. It

may well be an Old World species and perhaps is related to I. cairica, which is probably native to Africa.

Ipomoea heptaphylla has been confused at least with I cairica and is morphologically more similar to that spe-

cies and others like it in Africa than those elsewhere. Roxburgh (1824) pointed out that the plant appeared

unexpectedly in the nursery at the botanical garden in Calcutta, but he did not think it native, at least in his

region.

Verdcourt (1961: 11) thought that the plant climbed with the spiraled peduncles. Although Austin has

this plant few times, no note was made of the peduncles behaving like tendrils. Perhaps someone examin-

ing living plants in the field will be able to clarify the role of the spiraled structures.

^om^ C b l,,: 17 011 MCX HWy ^ ^ °^ unct * on ° f road t0 (= edge of

P°ttUi Oct 1985, Mger 85-12641 Mpio Navojoa: Navojoa, water edge of river back-water, limb and tube white, throat purple, 27 Oct 1939,

a imperati (Vahl) Griseb., Cat. Pi. Cub. 203. 1866. Type: imperato, Hist.

>) (lectotyfe!). Convolvulus imperati Vahl, Symb. Bot. 1:17-18. 1

J24. 1763. Type: based on Convolvulus Joliis obtusis, Plu

Convolvulus stoloniferus Cyrillo, PL Rar. Neap. 1:14. 1 5, 1788. Type: based on

‘iw Plate (lectotype!).

Herbs > Perennial, the stems prostrate, rooting at the nodes and often underground, reaching 5 m or more in

‘ en gth. Leaves 1.5-8 cm long sometimes linear to ovate or oblong, lobed or not lobed, the size and shape van-

l-T — a- in th<« morning. Se-

** Ul5 ^ long, unequal, oblong, the outer two sepals shorter, acute to obtuse, glabroi

Felgeretal.,

499

loba, with larger flowers, is usually allogamous (outcrossing; McDonald et al. 2011). The honeybee, introduced

from the Old World, carries pollen from one species to the other and is the only insect known to move between

the two. However, certain other pollinators, when present, will visit I. lacunosa and the hybrid. Presumably the

hybrid is moved, as it has been around the world, as a contaminant in seeds of cultivated plants.

, 1200 ft, 28 Oct 1939, Gentry 4767!; Rancho San Pedro, E entry to El Caj6n

-26!; Alamos, in streambed, 390 m, flowers pink, 2 Feb 1992, Van Devender

imin Hill: 24 km S of Rancho El Seri, 8.2 km S of Rancho El Carrizo (16.9

le, 29°58'58"N, 111°15 , 24"W, 735 m, 22 Aug 2007, Van Devender 2007-890!

1, Friedman 438-94 ! Mpio Navojoa: Navojoa, w

river back-water, 27 Oct 1939, Gentry 4747! Mpio Onavas: Onava

1986, Rea 1210! Mpio San Lois R.C.: 5 km S of San Luis, roadside

morning, 6 Oct 1985, Felger 85-1032 ! Mpio Yecora: Santa Ana 2

28°22'40" N, 109°08' W, 950 m, Otts, Martin, et al. 10 Oct 1988.

Ipornoea longifolia Benth., PI. Hartw. 16. 1839. Type: MEXICO. Zacatecas: Hartwcg 97 (iiolotype: K!; isotypes: NY!, P!>; other

sheets with species #97 (species numbers?) are from Chihuahua (K!) and Leon (BR!, LD!).

Large perennial herbs forming large tuberous roots, somewhat fleshy, rhizomatous, glabrous, the stems long,

trailing, decumbent, or ascending. Leaves with blades linear to oblong-lanceolate, often 10-12(-20) cm long,

2-4 cm wide, entire, glabrous; petiole 0.5-1.7 cm long. Inflorescences mostly solitary. Flowers on peduncles

3.5-4(-ll) cm long, the pedicels 2-3.5 cm long, mostly erect in fruit. Sepals ovate, the outer ones 12— 14(— 17)

mm long, 6-7 mm wide, the inner ones 15-20 mm long, 7-8 mm wide, coriaceous, glabrous. Corollas funnel-

form, 7-10 cm long, the limb whitish to cream-white, 7-8 cm wide, the throat purple or deep reddish purple.

Capsules ovoid, 14-16 mm wide, with an apiculum 2-3 mm long. Seeds 1-3(4), 10-11 mm long, ovoid, brown,

with long pilose indumentum on the margins and near the apex.

Sonora. — North-central and northeast part of the state in grassland, oak woodland, foothills thomscrub,

and tropical deciduous forest; 975-1850 m. Flowering April-September.

General distribution . — Southeastern Arizona and northern Mexico in Aguascalientes, Chihuahua, Du-

The flowers are open in the evening and are pollinated by moths, although there have been reports of bees

also visiting (Austin 1986).

Perennial herbs, roots tuberous, the stems erect, scandent, or less often twining, 4-50 cm long, mostly

branched from the base, glabrous. Leaves variable, the first ones usually elliptic, rhombic, or narrowly lanceo-

kte, 1.5-5 cm long, 3-20 mm wide, occasionally lobed, the lobes to 6 mm long, 5 mm wide, margins entire or

Regularly dentate, apex acute mucronulate the base attenuate, the distal leaves sometimes grading into pal-

•^tisect laminas with 3-5 segments, about equal or unequal, filiform, linear, or lanceolate, 7-42 mm long, 1-6

mm Wide - the outer segments often shorter than the inner ones, margins entire, apex obtuse or acute, the base

attenuate, glabrous; petioles almost absent or 2-10 mm long. Inflorescences monochasial axillary cymes.

,1-3 ml

nn

502

Sonora .— Central and mostly southeastern part of the state in tropical deciduous forest, ca. 150-790 a.

Flowering at least August-November.

General distribution . — Texas and Arkansas to Florida; Mexico including Baja California Sur, Chihuahua,

uila, Guerrero, Jalisco, Edo. Mexico, Sinaloa; Argentina, Ecuador, Peru, Venezuela. Native to Mexico and

ttive in the southern United States and South America, and also naturalized in many parts of the Old

d. This species has been widely spread through the southeastern United States as a contaminant in soy-

seeds (Gunn 1969a, 1972). It continues to be dispersed and established in new areas.

Confusion between names for I. capillacea and I. muricata began in the 1800s and a new layer of confusion

was added by Gunn (1969a, 1969b, 1972). What these authors failed to realize, or at least point out, in addition

was clarified by Staples et al. (2005). Ipomoea capillacea is a moming-flowering herb with corollas 3-4 cm long

while I. muricata is an evening- and night-flowering vine with corollas 3-7.5 cm long. The leaves are also nota-

bly different: I. capillacea has nearly sessile highly dissected leaf blades with linear segments and I. muricata has

petiolate entire to 3- or 5-lobed blades. The current residual confusion involves Gunn’s conclusions so the

earlier problems will not be addressed. The underlying problem is that Gunn (1969b) used the wrong date for

the names involved, rejecting I. muricata (L.) Jacq. (1798) and using I. turbinata Lag. (1816). Staples et al. (2005)

Ipomoea nil (L.) Roth, Catal.Bot. 1:36. I/ 1 -)/. Type: Dillenius, Hort. Eltham. 1:96 i 80 1 91 1732(lec totype!). PhariritisnilO-

Herbs, annual, herbaceous, the stems twining or prostrate, to 3-4 m long, hispid, the trichomes yellow. Leave*

ovate to ± orbicular, 5-15 cm long, 2-14 cm wide, entire to 3- or 5-lobed, the apices acuminate, hirsute. Inflo-

rescences cymose, dichasial. Flowers 1-4. Sepals 1.5-2.5(-3) cm long, ± equal, linear-lanceolate, basally nar-

rowly ovate, the base densely hispid with yellow trichomes, hispid to strigose on the upper parts, or distally

glabrous. Corollas (2-)3-6 cm long, funnelform, purplish to blue, at times white or red, the tube white or yel-

low within, glabrous. Capsules 8-12 mm long, rounded-depressed, 3-locular, brown, glabrous. Seeds l-4(-®-

4-9 mm long, dark brown to black, densely short-pubescent. 2 n = 30.

Sonora .— Near the margins of the Sonoran desert, coastal and foothill thomscrub, and tropical deciduous

forest, in disturbed as well as natural habitats; 50-ca. 1400 m. Growing and flowering mostly following war®

weather rains, especially August-October.

General distribution .— Mexico (probably most or all states); Mesoamerica; Argentina, Bolivia, Brasil, Co-

lombia, Guyanas, Ecuador, Venezuela; Caribbean; cultivated and introduced into the Old World.

The seeds have been utilized as a laxative (Austin 2000c). There are three species of somewhat simi#'

Felgeret al, Convolvulaceae (excluding Cuscuta) of Sonora, Mexico S03

appearing morning glories that are often confused. One is I. nil, characterized by long, straight, subulate sepals,

in tropical regions worldwide. The second is I. hederacea, also characterized by long sepals, but curving and

with an ovate base and abruptly attenuate apex, in temperate regions by latitude or higher elevations in tropical

latitudes. The third member is I. purpurea, which has short fat sepals and is worldwide.

Ipomoea nil and I. hederacea can be hybridized with difficulty (Yoneda and Takenaka 1981). Historically

they were geographically isolated. Superficially I. nil seems similar to I. hederacea, but detailed morphological

and molecular genetics show that they are not as close as they may seem. Furthermore, I. nil seems closer to I.

eriocalyx Mart, of South America than to I. hederacea (Austin et al. 2001).

Selected references.— Austin (2000c), Austin et al. (2001), McDonald et al. (2011).

nth) G. Don, Gen. Hist. 4:275. 1838. Tm: VENEZUELA: Humbol

is parasiticus Kunth, Nov. Gen. Sp. (quarto ed.) 3:103. 1818 [1819].

Proc. Amer. Acad. Arts 21:319-320. 1894. Type: MEXICO. Jalisco: Tequila, 1

Herbs, annual, the stems twining, herbaceous, to 2-7 m long, with fleshy tubercles on the older parts, or

smooth, glabrous or glabrate. Leaves 5-10 cm long, 6-9 cm wide, entire, cordate to broadly cordate, apically

acuminate to rounded and cuspidate, lightly pubescent or glabrous above, mostly glabrous below. Inflores-

cences usually simple cymes, less often compound-cymose. Flowers 2-10. Sepals 4-6 mm long, more or less

e qual, the outer ones ovate to ovate-lanceolate, more or less acute and cuspidate, with small appressed-puber-

u, ent indumentum. Corollas 2.5-3 cm long, funnelform, purple, sericeous on the interplicae. Capsules 10-12

mm lon g> ovoid to ovoid-globose, reflexed, glabrous. Seeds 1-4, 6-7 mm long, narrowly ellipsoidal, brown,

Wlth sma11 Puberulent indumentum or glabrous.

Sonora.— Eastern Sonora in Chihuahuan desert, foothills thomscrub, tropical deciduous forest, and ri-

P-Jrtan in oak woodland; 260-1290 m. Flowering November to February. Although not known from the Unit-

e States, its occurrence at Agua Prieta suggests that it might be found in nearby Arizona.

General distribution .— Baja California Sur, Chiapas, Chihuahua, Guanajuato, Guerrero, Jalisco, Edo. Mexi-

c °. Michoacan, Morelos, Nayarit, Oaxaca, Sinaloa, Veracruz; Mesoamerica; Brazil, Colombia, Peru, Venezuela.

These Pknts contain an ecdysone that was considered at one time as a potential insecticide. Subsequently,

or similar compounds were found in I. nil and became commercially known as “kaladana” (Austin

504

Journal of the Botanical Research Institute of Texas 6(2)

. Type: MEXICO. Chiapas: wooded slope 9 km N of Tuxda GutiCrrez along

MANTELA DE MARIA, TROMPILLO; JtCURE ‘TWISTED YARN’ (Mayo)

Lianas, perennial, the stems 2-10 m long, twining with a woody base, often winged, the younger parts herba-

ceous, smooth to somewhat winged, glabrous or pubescent. Leaves 3.5-14 cm long, 3-12 cm wide, ovate, often

broadly ovate, entire or 3- or 5-lobed, chartaceous, basally cordate, apically acuminate, glabrous or pubescent

on both surfaces at least near the base. Inflorescences cymose, axillary. Flowers (2-)5-15(-35). Sepals un-

equal; the 2 outer sepals 4-7 mm long, ovate-lanceolate, broadly elliptic to oblong, acute, muricate or more of-

ten with wings on the lower portion; inner sepals 5-8.5 mm long, obtuse to obtuse-mucronate, coriaceous, the

margins scarious, glabrous or pubescent only toward the apex. Corollas 5-9 cm long, funnelform, purple,

slightly pubescent at the apices of the petals. Capsules 10-19 mm long, ovoid, brown, apiculate, glabrous.

Seeds 1-4, 6-7 mm long, ellipsoid, brown, minutely puberulent or glabrescent.

Sonora . — Central and southern parts of the state in coastal and foothills thomscrub and tropical decidu-

ous forest; ca. 50-550 m. Flowering September to November. The seeds, ground, roasted, and boiled, were

used by the Guarijlos as a purgative (Gentry 1942). The Guarijlos also used a morning glory, possibly this spe-

cies, as follows: “If a woman does not wish to have a child, she will sometimes eat the seeds of trompillo, grind-

ing up the seeds, mixing the gruel into water, and drinking it” (Yetman & Felger 2002:193). The Mayos con-

sider the herbage of this fast-growing summer vine to be valuable forage for cattle and goats (Yetman and Van

Devender 2002).

General distribution. — Chiapas, Chihuahua, Colima, Guerrero, Hidalgo, Jalisco, Michoacan, Morelos,

Nayarit, Oaxaca, Puebla, San Luis Potosl, Sinaloa, Tamaulipas, Veracruz; Mesoamerica.

Selected reference .— Austin (1997).

Mpk> Alamos: San Bernardo, 24 Aug 1935, Gentry 1616 ; Rio Mayo Raft Trip, confluence of Rio Mayo with San Ignacio, 27°55'N, 108°4TW,

Jenkins & Rondeau 26 Sep 1991!; 23.3 mi by road W of Alamos, Soule & Krizman 2 8 Aug 1964!; Arroyo el Mentidero at El Chinal road, 113 b»

S of Alamos, 26°54 I 45"N, 108°55'05" W, 240 m, 20 Sep 1993, Van Devender 93-855!; Alamos, 28 Sep 1991, Van Devender 91-755 !; Gueybatnpo.

E edge of Arroyo Tojibampo, 26°42'30"N, 109°16’W, 50 m, 22 Sep 1994, Van Devender 94-713 !; 0.3 km E of Tqjibampo (S side of the Sierra*

Alamos), 26°48'50"N, 108°58'W, 240 m, 21 Sep 1994, Von Devender 94-659!; Arroyo Huirotal, Rancho Las Uvalamas, E slopes of Sierra*

Alamos, 550 m, vine 6-8 m in trees, flowers purple, 13 Sep 1994, Van Devender 94-564 (AR1Z!, ASU!). Mpio Huatabampo: 1 km SEofCer-

rillos, 9.5 km SE Melchor Ocampo, 40 m, flowers purple, 21 Sep 1994, Friedman 347-94 (ASU!). Mpio La Colorada: 7 mi NE of Colorado,

between Colorado and Mazatan, climbing vine to 8 or 10 m, corolla rich purple, 6 Sep 1941, Wiggins & Rollins 323 (CAS!): 3.5 km SE * '

from Navajoa-Alamos, sandy bottomlands, thorn forest, seeds used as purgative, M antela de Maria, 5 Nov 1939, Gentry 4880 !

Felger et al., Convolvulaceae (excluding Cuscuta) of Sonora, K

Ipomoea pes-caprae (L.) R. Br. in Tuckey subsp. I

isiliensis (L.) Ooststr., Blumea 3:533. 1940. i

Batatilla, cmmisTATE de playa; bejuco de playa; beach morning glory, railroad vine, g

Herbs, perennial, the stems mostly prostrate on beaches, rarely twining, fleshy, to 10 m long (usually much

shorter in Sonora), glabrous, and with milky sap. Leaves 3-10 cm long, 5-10 cm wide, ovate to reniform, ba-

sally rounded, truncate to cordate, apically normally emarginate, the blade with 2 glands near the base. Inflo-

rescences axillary, monochasial and/or dichasial. Flowers 1-5. Sepals 5-11 mm long, equal or unequal, ellip-

tic, ovate-elongate to orbicular, glabrous. Corollas 5-6 cm long, funnelform, pinkish or lavender, the throat

darker within, glabrous. Capsules 1.5-2.2 cm long, rounded, straw-colored or brown with purplish patches,

glabrous. Seeds 1-4, 8-9 mm long, rounded, densely brown-tomentose. 2 n » 30, 60.

Sonora . — Cultivated on beaches from the vicinity of Bahia Kino southward and sometimes weakly estab-

lished, and perhaps native or at least established on beaches along coastal thomscrub in extreme southwestern

Sonora; near sea level. Flowering at least during the warmer months.

General distribution . — This species occurs on beaches and coastal dunes worldwide; subsp. brasiliensis is

widespread in the New World tropical shores. Texas; Baja California (norte) and Sur, Chiapas, Guerrero, Jalis-

co, Michoacan, Oaxaca, Sinaloa, Tamaulipas, Veracruz; Mesoamerica; Brasil, Colombia, Ecuador, Guyanas,

Pern, Venezuela; Caribbean. Subspecies pes-caprae occurs in a narrow range in the Indian Ocean. A close rela-

tive, I. asarifolia (Desrousseaux) Roemer & Schultes, has been introduced into wetlands in the American trop-

ics (Austin 2005).

Selected reference.— St. John (1970).

it, 26.6917°N, 109.587°W, o

a (by air) S of Us Bocas, 26.65°N, I09.331°W, s<

Perennial herbs with globose, tuberous roots, the stems erect to procumbent, ascending, not twining or only

slightly at tips, glabrous. Leaves 1-3 cm long, orbicular in outline, palmately and pedately lobed (rarely cune-

ate-obovate in which case the apex is laciniate-dentate), basally cordate, the lobes acute, glabrous; petioles 2-5

mm ^ng Inflorescences solitary. Flowers on peduncles 1.5-2.5 cm long, erect or reflexed in fruit; bracts ca-

duceus. Sepals unequal; outer sepals 5-8 mm long, 2-3 mm wide, oblong, obtuse to acute, mucronate, muri-

ate at least along the midrib; inner sepals 7-9(-10) mm long, 3-4 mm wide, broadly ovate, acute to acuminate,

muricate on the midrib or glabrous. Corollas funnelform, 2.5-3.1 cm long, purple, glabrous, the limb 1.8-2.2

cm wide. Capsules ± globose, 5-6 mm wide, with an apiculum to 5 mm long. Seeds 1-4, 2-2.5 mm long, ovoid,

black to dark brown, finely tomentose.

Sonora.— At least in southern Sonora, coastal thomscrub and in pine-oak forest in southeastern Sonora

near the Chihuahua border. Also southeastern Arizona near the border and expected in nearby Sonora and

Perhaps elsewhere in mountains in eastern Sonora. Open rocky slopes. 20-above 1250 m. Flowering April-

October;

General distribution. — Arizona, New Mexico, Texas; Chihuahua, Coahuila, Distrito Federal, Durango,

^go, Jalisco, Edo. Mexico, Michoacan, Puebla, Veracruz, Zacatecas; Argentina, Bolivia, Peru. McDonald

" 5) rec °gnized several widespread varieties, with the Sonoran population being var. plummerae.

Journal of the Botanical Research Institute of Texas 6(2)

Although McDonald (1995) recognized var. cmeifolia as distinct, DFA has seen individual populations in

southern Arizona with both “var. cuneifolia” and “var. plummerae ” growing intermixed. We suspect that the

two “varieties” are nothing more than variants in leaf shapes.

This species is easily confused with I. capillacea. Leaf segments are <1 mm wide in I. capillacea, and >1

mm wide in 1. plummerae. Also, I. capillacea is an erect herb, while I. plummerae is a prostrate or rarely twining

Selected reference. — McDonald (1995).

Mpio Alamos: Cliffs 5 km W of Chiribo, Sierra Saguaribo, 1400 m, pine-oak woods, Martin 24 Aug 1993. Mpio Yecora: Yecora, 28°22'25"N,

108°56'W, 1540 m, 6 Sep 1995, Rema-G. 96-373. CHIHUAHUA. Mpio Buenaventura: Rio Santa Maria, 10 km SW of Buena Ventura [Buena-

ventura], 2100 m, 14 Aug 1989, Jenkins et al. 89-311.

Ipotnoea pubescens Lam., Tabl. Encycl. 1:4

5. 1791 [1793]. Type: AMERICA, c

1707. Type MEXICO: horto Regio, 17

York Acad. Sci. 18:196. 1908. Type: U

7. Ortega (lectotype MA! ,

Perennial herbs from a large, oblong root; stems twining, hirsute with retrorse trichomes. Leaves with blades

2-8 cm long, 2-9 cm wide, ovate, nearly entire with the margins sinuate, or 3- or 5-lobed and palmate, basally

cordate, the lobes elliptic to ovate, hirsute with antrorse trichomes, at times sericeous, apically acute to obtuse,

mucronate; petioles 2-5 cm long. Inflorescences 1- or 2-flowered. Flowers on peduncles 15-18 mm long;

pedicels 3-10 mm long, erect in fruit; bracts 5-12 mm long, subulate, acuminate. Sepals unequal; outer sepals,

9-21 mm long, 5-11 mm wide, ovate, basally truncate, acuminate; middle sepals 9-19 mm long, 3-8 mm wide,

asymmetrical, ovate, acuminate; inner sepals 9-20 mm long, 2-4 mm wide, ovate-lanceolate, antrorsely his-

pid, at times sericeous. Corollas 5.5-8 cm long, funnelform, blue to violet with a white throat, glabrous, the

limb 6-7 cm wide. Capsules 10-12 mm wide, ± globose, surrounded by the sepals. Seeds (1-) 3-6, 5-6 mm

long, ovoid, brown to black, densely hairy with velvety trichomes.

Sonora.— Mountains in eastern Sonora. It grows in south-central and southeastern Arizona near the bor-

der and is expected in nearby Sonora. Oak woodland and pine-oak forests, especially rocky areas and near

streambeds; ca. 1200-2000 m. Flowering August-September.

General distribution. — Arizona, New Mexico, Texas; Chihuahua, Durango, Hidalgo, Michoacan, Quere-

taro, San Luis Potosi; disjunct to South America.

The Arizona plants have larger flower than those listed from South America by O’Donell (1959a) and

there is perhaps more than one taxon involved. McDonald et al. (2011) have placed all the large-flowered plants

of northern Mexico and the SW United States in I. lindheimeri. We cannot agree because the sepals are distinct

within the populations that we have studied in Sonora and Arizona. McDonald (pers. comm., June 2012) says

that he has found what appear to be hybrid swarms between the typical I. lindheimeri with narrow lanceolate-

linear, cuneate sepals and plants of I. pubescens with broadly ovate, basally truncate sepals. We have not seen

such populations and maintain them as separate species until future studies can be made of their relationships

Selected reference.— Austin (1991).

s: Sierra Saguaribo, Chir

d, rare, one patch along s

1 24 Aug 19931; Sierra Sahuaribo, ca. 2 km (by air) NE of La Vinateria, 27°17'30 , N,

Ipio San Felipe de Jesus: Cajbn Infiemo, northern Sierra Aconchi, 4480-5120 ft

•row part of canyon, 18 Sep 1982, Reichenbacher 11391 Mpio Yecora: ReS "‘^

;. 2000-561 (USONt).

Ipomoea purpurea (L.) Roth, Bot. Abh. Beobs

Ipomoea hirsutula Jacq. f.

U.84,f.97,1732(tECTorrf*

ype. Pharbitis htspida Cho^f’

r. diversifolia (Lindl.)

Journal of the Botanical Research Institute of Texas 6(2)

Ipomoea scopulorum Brandegee, 1

(holotype: UC!, photo FTG-FAU!).

: 5:169. 1903. ^MEXICO. B*a California [Sur]: Cape F

Perennial twining herbs, with tuberous roots?, 1-2 m long, sparsely to densely pubescent throughout with

simple and stellate trichomes. Leaves 6-8 cm long, 4-6 cm wide, cordate-ovate, entire, stellate-pubescent

above and below; petioles 2-5 cm long. Inflorescences cymose or solitary, axillary. Flowers on peduncles 2-9

cm long, the pedicels 2-3 cm long, erect in fruit; bracts 2-3 mm long, subulate, caducous. Sepals 7-10 mm long,

4-6 mm wide, unequal, broadly oblong to broadly ovate, smooth, rounded to obtuse, the upper margins some-

what scarious, glabrous or less often sericeous on the outside of the outer two sepals. Corollas 6-8 cm long, 6-7

cm wide, funnelform, white with a purple throat or less often purplish throughout, glabrous. Capsules ovoid,

10-14 mm long, 8-10 mm wide. Seeds 1-4, 6-8 mm long, ovoid, brown, silky, with white or tawny trichomes

8-10 mm long on the margins, otherwise glabrous.

Sonora . — Southern to north-central parts of the state; Sonoran desert, coastal and foothills thornscrub,

and tropical deciduous forest; 150-725 m. Flowering August-September.

General distribution . — Baja California Sur, Sinaloa.

Ipomoea seaania Felger & D.F. Austin, Sida 21:1296. 2005. Type: MEXICO. Sonora: Mui

1 km north of Bahia San Carlos on old road to Bahia Algodones, 27 Feb 1985, Felger & Devini

MEXU!, MO!, NY! See label information below).

Openly-branched shrubs 1-4 m. tall, with multiple woody stems from the base, the upper twigs sometime 5

sinuous or moderately spiraling, sometimes becoming extremely slender. Herbage largely glabrous or glabrate

except newest growth densely to sparsely short-pilose, the trichomes mostly spreading. Leaves drought de

ciduous, lanceolate to ovate, long shoot leaves often 2-8 cm long, 1.5-2 cm wide, the blades lanceolate to ovate,

with 6— 8(— 10) lateral pairs of primary veins, the base obtuse to subtruncate, the apex obtuse to emarginate, tk

midrib often ending in a short mucrone, glabrous; petioles 8-15(-20) mm long, with a pair of glands, usua^

conspicuous, at the junction of petiole and blade, seen on the lower leaf surface. Spur branch leaves often 4^

cm long, to 4-11 mm wide, linear to linear lanceolate with 6-10 lateral pairs of primary veins, the base obtu#

to subtruncate, the apex obtuse or blunt, or sometimes shallowly emarginate, the midrid often ending io 1

Felgeretal., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

509

short mucrone; petioles to 2-9(-13.5) mm long. Inflorescences of 1 or 2 (3) flowers, appearing solitary but

cymose on short-shoots 2-5 mm long, these sometimes with a few small leaves; bracts 5-8 mm long, quickly

deciduous, broadly oblong with an obtuse tip; peduncles very short, to 5 mm long, the pedicels 8-22 mm long.

Sepals 12-17 mm long, 6-8 mm wide, broadly lanceolate to mostly ovate, puberulous to villous, the inner

surfaces generally more densely hairy than the outer surfaces, the trichomes white, appressed to mostly as-

cending, and curly to straight. Inner (adaxial) sepals obtuse, the surfaces with trichomes 0.15-0.6 mm long;

outer (abaxial) 2 sepals acute, slightly narrower and more sparsely pubescent than the inner 3, the trichomes

0.1-0.5 mm long, the sepal margins scarious and glabrous or glabrate. Corollas 4-6 cm long and 7-8 cm wide,

showy, funnelform, glabrous, white with yellowish interplicae and a maroon band at inside base of the tube.

Stamens 5, with 4 filaments 25-26 mm. long, the fifth stamen 23-24 mm long, filaments pubescent on the

basal 4 mm; anthers 6-7 mm, long oblong, sagittitate; pollen spheroidal, spinulose. Ovary 3 mm long, gla-

brous; style 37-38 mm long, glabrous; stigma 2-globose.

Sonora . — Near the southern margin of the Sonoran desert where it is endemic to hills and mountains west

of Bahia San Carlos, generally on rocky ridges, steep colluvium and rhyolite slopes of canyon sides and cliff

base; near sea level-100 m. Although locally common, the San Carlos population is threatened by tourist de-

velopment (Felger 1999). There are no other records for this unusual shrub. Flowering January-April.

Ipomoea sescossiana Baill., Bull. Mens. Soc. Linn. Par. 1:385. 1883. Type MEXICO. San Luis Potosc Sescosse s.n. (holo-

Perennial erect herbs, with tuberous roots, the stems shrubby, spreading, glabrous. Leaves 2-4 cm long, 0.5-

1-5 cm wide, orbicular in outline, irregularly pinnately divided into 5-9 obtuse-tipped divisions, the lobes

linear to filiform, entire or rarely toothed, glabrous; petioles 3-15 mm long. Inflorescences mostly of solitary

flowers, at times 2 or 3 flowers. Rowers on peduncles 0.8-3.5(-8) cm long, the pedicels 0.6-1.5 cm long, ac-

crescent and erect in fruit; bracts 2-3 mm long, lanceolate. Sepals 6-ll(-13) mm long, 3-6(-8) mm wide,

subequal, oblong-ovate, smooth, obtuse-rounded to emarginate or cuspidate, the margins scarious. Corollas

6-10 cm long, funnelform, rose-purple to purple, the tube whitish, glabrous, the limb 6-8 cm wide. Capsules

14-16 mm long, 10-14 mm wide, ovoid to almost globose. Seeds 1-4, 8-9 mm long, compressed ovoid, black,

finely appressed tomentose.

Although not documented for Sonora, it occurs in Chihuahua near high elevations in east-central Sonora

and * likel Y to turn up in the highlands of easternmost Sonora.

General distribution.— Chihuahua, Coahuila, Durango, San Luis Potosi, Zacatecas; 1000-2400 m. Flow-

erin gJune-October.

This species is an erect herb with bee-flowers. This xenogamous species is sister to I. ancisa and I. stans

(McDonald 2001; Ana Rita Simoes, pers. comm. 23 Nov 2011).

CHIHUAHUA. Mpio Moris: La Cieneguita, Rfo Mayo Upper Sonoran, 10 Sep 36, Gentry 26481 [ca. 35 km E of Sonora]; Mpio Madera:

510 Journal of the Botanical Research Institute of Texas 6 ( 2 )

Chuichupa, Aug 1936, Horde LeSueur 850\ [ca. 25 km E of Sonora] . Mpio Temosachi: N W of Yepdmera on hwy 180, 2100 m, flowers mostly

pale lavender, shrub Jenkins, Martin, & Moore 21 Jul 1986\

Ipomoea tenuiloba Torn, Rep. U.S. Mex. Bound., Bot. 148-149. 1859. Type: U.S.A. Texas: hills and rocky places near

pedately 5-9 lobed, the lobes linear to lanceolate, 0.5-6.5 mm wid

Perennial herbs, with tuberou

long, 3-10 cm wide, orbicular :

entire, glabrous; petioles 2-38 1

mm long, sessile or the pedicels to 8 mm long,

ate. Sepals unequal; outer sepals 5-11.5 mm lor

rved in fruit; bracts 1-3 mm long, 1

-3 mm wide, oblong-lanceolate, muricate along the midrib

nate, smooth, the margins scarious, glabrous. Corollas 4.5-10 cm long, funnelform or salverform, completely

white or white with pale rose to purple limb, glabrous, the limb 3-3.6 cm wide. Capsules 4-8 mm wide, ±

globose to broadly ovoid, with an apiculum 4-5 mm long. Seeds 1-4, 3.5-5 mm long, ovoid, black to dark

brown, finely appressed tomentose. This delicate twiner has moth flowers that McDonald (2011) considered

xenogamous.

Sonora . — Documented in Sonora with four specimens from mountains in the eastern part of the state, in

hills and rocky sites, oak woodland and pine-oak forest; ca. 1280-1950 m. Flowering August-September.

General distribution. — Arizona, New Mexico, Texas; Chihuahua.

Two varieties are recognizable: Variety tenuiloba with 5-7 leaf divisions to 1.2 mm wide, a mostly white

salverform corolla 6.5-10 cm long and the free portions of the filaments 8-11 mm long. Variety lemmoni (A.

Gray) Yatsk. & Mason with 7-9 leaf divisions to 6.5 mm wide, a funnelform corolla 3.5-6.5 cm long, with a

white tube and rose to purple limb and the free portion of the filaments 14-19 mm long. In some areas these

varieties intergrade, and a specimen from northeast Sonora (White 3474) was considered an intermediate by

Yatskievych and Mason (1984). The species is nocturnal and those studied in southern Arizona opened about

1:00 a.m. and often closed before dawn (Austin 1991).

Mpio Alamos: Sahuaribo and vicinity, 27°19'N, 108°34'W, 1550 m, Martin 23 Aug 19921 Mpio Bavispe: Sierra el Tigre, Las Tierritas deB

28.38°N, 108.8258°W, 1410 m, very common herbaceous perennial in dense

open at 9:30 a.m., 2 Sep 2000, Van Devender 2000-640 (USON!); Yficora, 28.3636°N, 108.9228°W, 1540 m, grassland, locally conn

.,5 Sep 2001, V

fectively selected the Thurber 977

Annual herbs, with a fibrous root system; stems slender and procumbent, twining at tips, glabrous. L*® 1 **

with blades pedately 5 or 7 parted, 1.5-3 cm long and wide, basally cordate, the segments linear, apically acute,

glabrous to remotely setose; petioles to 2 cm long. Inflorescences of solitary flowers or cymose. Flowers 1 ° r

2, the peduncles slender, equaling or exceeding the leaves, the pedicels (10-)15-18(-20) mm long, erect in

fruit; bracteoles 2-2.5 mm long, subulate. Sepals unequal; outer sepals 8-9 mm long, 2-3 mm wide, lanceolate;

a wide, attenuate-acuminate, glabrous or hirsute at least on the three main

ns, otherwise membranaceous. Corollas 2.5-3.2 cm long, funnelform, purple (oc-

casionally white), glabrous, the limb 3.2-4.6 cm wide. Capsules 3-5 mm wide, ellipsoid to ovoid depending on

the number of seeds, with an apiculum 5 mm long or longer. Seeds 1-4, 5-6 mm long, ovoid, black and gray

mottled, minutely appressed tomentose.

Sonora . — Statewide except the northwest; Sonoran and Chihuahuan deserts, foothills thomscrub, tropi-

cal deciduous forest, mesquite-grassland, and oak woodland; near sea level-ca. 2000 m. Flowering January

andJune-November.

General distribution .— The species, with 3 allopatric varieties, is widespread in deserts and tropical de-

dduous forests of southwestern United States to El Salvador. Variety leptotoma , the northernmost one, occurs

in southeastern Arizona, southwestern Chihuahua, and most of Sonora and Sinaloa, and Baja California Sur.

This species is self-incompatible and bee pollinated (McDonald 1995; McDonald et al. 2011).

Selected reference .— McDonald (1995).

MpioAgoaPrieta: Colonia Morelos, 2600 ft, 5 Oct 1941, White 4619! Mpio Alamos: Alamos, 16 Oct 1936, Gentry 2928!; 3.7 km N of Guiro-

coba, 450m, abundant annual infield, flowers white, 23 Sep 1994, Van Devender 94-759!; El Guayabo (upper) crossing of the Rio Cuchujaqui

3kmNE of Sabinito Sur and 15 km (airline) ESE of Alamos, near 27°00'N, 108°47’W, 350 m, tropical deciduous forest, 12 Oct 1992, Sanders

white flowers among abundant purple-flowered plants on disturbed roadside, flowers open 3:45 p.m., foothills thomscrub, 16 Sep 2000

1934, Shreve 6794!; Rancho el Carrizo, 2400 ft, Tomelson 7 Aug 1968\\ 2 km S of Los Chinos, 23 km N of El Oasis, foothills thomscrub, 665

m, flowers lavender, drying magenta, open 2:00 p.m., 6 Jan 2001, Van Devender 2001-14! Mpio La Colorada: 3 mi E of La Colorada, 1300 ft,

12 Jan 1981, Fischer 6869!; 4 km W of Cobachi (dirt road between paved roads to Tecoripa

Cnrarpe: Rancho Agua Fria, flowers white, Van Devender 6 Sep 197 6!; 4 mi SW of Agua Ft

Gunpas: 5 km ESE ofjecori, road to Moctezuma, 26 Oct 1984, Felger 84-380! Mpio Fronteras: 2

nal purple flowers, 4200 ft, 7 Sep 1960, Felger 4048! Mpio Guaymas: Arroyo Las Pirinolas, roat

Joside Robinson, 28.1112°N, 111.036667°W, 100 m, Felger 01-7421 Mpio Hermosillo: La Puerca,

10 a.m, 4 Sep 2000, Van Devender 2 000-671! Mpio Moctezu

ith white throat, open 8:00 am, 14 Aug 2006, R eina-G. 2006-468! Mp!

DV-NW of San Jose de Masiaca, 26.78170°N, 109.295°W, 200 m, 19 Jan 1995, Friedman

.an N of Teachive, SW end of Cerro 1 nscmb. 26°48.1'N, 109°14AV7, 100 m, 28 Sep 1'

96-582! Mpio Opodepe: 5.6 mi on Mex Hwy 15 S of Be

154-95 (ASU!); 2

51 3206°N, 110.367°W, 1555 m, grassland with isolated

, 30°23'38"N, 110°3r43"W, Meling I Oct 1986 (USON!). Mpio

:s, 29°25'30"N, 110°23'21"W, Meling 19 Sep

19 &(US0N!).

Ipomoea thurberi A. Gray, Syn. Fl. N. Amer. 2:212. 1878. Type: U.S.A. Arizona: Thurber 966 (ho^typh: GH!).

Ipomoea gentry! Sundl., Field Mus. Nat. Hist. 22:46. 1940. Type: MEXICO. Chihuahua: Gentry 2497 (holotype: F!; isotype: MO!).

Glabrous perennial twining herbs from an elongate, tuberous root, the stems trailing. Leaves with blades 1-5

on long, 2-6.5 cm wide, sagittate to pedately 5 or 7 lobed, the lobes divergent, lanceolate, linear to oblong

^tate, sparsely strigose; petioles 0.9-2 cm long. Inflorescences solitary. Flowers on peduncles 3

1,1111 lon g, the pedicels 7-8 mm long, erect in fruit; bracts ca. 1 mm long, scale-like. Sepals 12-15 mm long, 3

mmurirb, . . ,, -u obscurelv warty at base or not.

wide, ± equal, lanceolate, « HI |

-orollas 5-8 cm long, funnelform-salverform, white with a

ous-caudate, obscurely warl 7

mb and green tube (drying purple), opening

512

in the evening, glabrous, the limb 5-6.5 cm wide. Capsules 6-7 mm wide, ± globose to ovoid, with an apieu-

lum 4-5 mm long. Seeds 3-6, 3.5-4 mm long, ovoid, black to dark brown, finely tomentose.

Sonora . — Mountains of eastern Sonora in grassland, oak woodland, and pine-oak forest; at least 1150-

1600 m. Flowering Juiy-September. The flowers open at dusk when they are visited by hawkmoths, perhaps

Hyles lineata and probably others. Also near the Sonora border in Cochise and Santa Cruz Counties in Arizona

Herbs, annual, the stems prostrate and twining, 1-3 m long, glabrous or occasionally sparsely pubescent, the

indument concentrated on the nodes. Leaves 2-8 cm long, 2-7 cm wide, broadly ovate to orbicular, entire,

coarsely dentate to deeply 3-7-lobed, basally cordate, the basal lobes rounded to angular or lobed, apically

acute to obtuse, both surfaces glabrous or sparsely pilose. Inflorescences axillary, cymose. Flowers 1— 3(-12)

with peduncles variable in length from shorter to longer than the petioles, glabrous, angular, minutely ver-

ruculose at least toward the apex. Sepals 6-7 mm long, ± equal, the outer sepals oblong to narrowly elliptic-

oblong, obtuse to acute, mucronulate-caudate, glabrous or sparsely pubescent without, conspicuously cihate,

mostly glabrous otherwise. Corollas 1-2 cm long, funnelform, lavender, glabrous. Capsules 5-6 mm wide, ±

globose, brown, bristly pubescent. Seeds 1-4, 2.5-3.2 mm long, globose, dark brown, glabrous. 2 n = 30, 60.

Sonora .— A weed carried, at least partly, as a contaminant in rice and others seeds. Sonoran desert,

coastal thomscrub and tropical deciduous forest, often in disturbed sites; 0-950 m. Flowering all year.

General distribution. — California; Campeche, Chiapas, Guerrero, Jalisco, Edo. Mexico, Nayarit, Nuevo

Le6n, Oaxaca, Sinaloa, Tabasco, Tamaulipas, Veracruz, Yucatan; Mesoamerica; Argentina, Bolivia, Brasil, Co-

lombia, Ecuador, Paraguay, Peru, Venzuela; Caribbean.

Felgeretal., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

Vines, herbs or suffrutescent shrubs; stems herbaceous toward tips, procumbent to twining (except some-

times in ]. agrestis ), perennials or occasionally annuals, glabrous or hairy. Leaves chartaceous to herbaceous,

mosdy cordate, glabrous or hairy usually with stellate trichomes, entire or variously repand, dentate, or lobate,

petiolate. Inflorescences in scorpioid cymes, head-like cymes, umbelliform, or flowers solitary. Flowers on

pedicels 5-30 mm long, the bracts small and linear or lanceolate or large and foliose. Sepals equal or unequal,

variable in shape, hairy or glabrous. Corollas blue, lilac, or white (red in one West Indian species), subrotate,

campanulate, or funnelform, deeply lobed, dentate or almost entire, glabrous or hairy. Stamens and styles in-

cluded (in our species). Pollen 3-colpate, 12- or 15-rugate. Ovary 2-locular, 4-ovulate, glabrous or hairy; styles

1, filiform; 2 stigmas ellipsoid or oblong and flattened. Fruits capsular, 2-celled, with 4 or 8 valves, globose to

subglobose. Seeds 1-4, glabrous or pilose, or tuberculate, or winged.

About 80-100 species in the Americas and several in Australia.

Selected references .— Austin (2006), Robertson (1971).

Jacquemontia abutiloides Benth., Bot. Voy. Sulphur 34-35. 1844. Type: MEXICO. Baja Cauforn* [Sur]: Bay of Magdalena,

Hinds s.n. (holotype: K!; photo MO!).

Scrambling and twining perennial vines, the stems generally woody near the base and slender above growing

into and overtopping shrubs reaching 2 (3) m long; younger stems and herbage with 3-branched stellate hairs,

mostly crowded but varying from densly overlapping to sometimes moderately dispersed when vigorously

growing following ample rain, the hairs nearly white when young and golden-brown with age. Leaves ovate to

broadly ovate, to 8 cm long and 4.5 cm wide, usually Vi to% that size, apically obtuse, acute or rarely retuse,

mucronate, acuminate or cuspidate, basally cordate with a deep and wide sinus, pubescent; petiolate. Inflores-

cences axillary, cymose, rarely solitary, the dichasia 1-2 times compound, the peduncles 2.5-8(-13.5) cm

lon g. Flowers on pedicels 1-6 mm long, the bracts linear, to 12 mm long. Sepals of different sizes, the outer 2

ovate to narrowly ovate with attenuate apices, 7-11 mm long, 3-6 mm wide, the middle sepal narrowly ovate,

attenuate apically, sometimes slightly falcate, the inner 2 sepals ovate or broadly ovate, 5-7 mm long, 2.5-3 mm

^de, apically attenuate, the bases narrowed with a stipe ca. 1 mm long; all 5 sepals pubescent on the outer

surfaces and to a lesser extent on the inner surfaces, enlarging slightly in fruit. Corollas (1.8-) 2-3.5 cm broad

when open, broadly campanulate to rotate, blue. Stamens ± equal or unequal, the anthers 1.5-2.5 mm long,

^ary ovoid, 1.5-2 mm long, the styles 6-7 mm long, longer than the stamens. Capsules 5-6 mm ong

broadly ovoid, usually opening by 8 segments, partly enclosed by sepals. Seeds 1-4, 2.7-3.5 mm long, l.B-2.3

““a wide, trigonous, minutely areolate and ruminate. j J . .

, ^ region. — Sonoran desen on Ida Tiburdn from near sea level-a. least 490 m; widespread m the

b 'en mountain mass (Sierra Kunkaak) and its eastern bajada to the south shore of the island, especially a o g

Was ^ les and canyons, andalsoon desert plains and rocky slopes (Felger et al. 201 2). Flowering with sufficient

-tat a. various seasons, flowers recorded October-April Not known from mainland Sonora, whteh

Seems unusual since seemingly similar habitat to that on the island occurs on die adjacent Sonora mainland

* heic Fel s er et al. (2012) have searched for it.

515

Jacqnemontia pentanthos (Jacq.) G. Don

This species has not been verified for Sonora. While it is common in eastern Mexico, it is either absent or rare

in western Mexico. According to Robertson (1971) J. pentanthos is at the center of a group of species that in-

cludes J. abutiloides.J. albidaj. eastwoodianaj. polyanthaj. pringlei, and five others. This group of taxa ranges

from Arizona to Central America. We tentatively accept the distinctions made by Robertson (1971) in these

segregates (except for J. albida), although the relationships of these taxa generally remains confused. These

taxa are often misidentified or difficult to identify on herbarium specimens, while living plants are often con-

siderably different. Within this alliance J. polyantha and J. pentanthos are closely related (see J. polyantha ).

Jacquemontia albida Wiggins & Rollins, Contr. Dudley Herb. 3:277. 1943. Type: MEXICO. Sonora: [Mpio Hermosillo] Along an arroyo 1

mile NE of El Zapo, between Hermosillo and Tastiota, 4Sep 1941, Wiggins & Rollins 273 (holotype: DS at CAS!; isotypes: ARIZ!, GH!,

LL!, MICH!, MO!, NY!, RSA!,US!).

Paneya (Guarijio, Gentry 1081)

Perennial vines, sparsely to densely pubescent with 3-armed trichomes. Stems twining to several meters long

or procumbent, herbaceous or becoming woody near the base. Leaves: Blades 2.5-11.5 cm long, 1.5-6.5 cm

wide, ovate, broadly ovate, to almost circular; basally cordate to less often truncate; apically highly variable—

pand to undulate. Petioles 0.2-6.5 cm long. Inflorescences axillary, loosely cymose, of simple or often com-

pound dichasia; peduncles 5-13 cm long and often longer than the leaves. Flowers on pedicels 0.5-2 cm long,

the bracts linear, often 5-10 mm long. Sepals unequal to more or less equal; the outer two sepals 6-6.5 mm

long, 3.5 mm wide, elliptic, broadly elliptic, or ovate to broadly ovate, the bases narrowed, the the apices acute

or acuminate, or shortly attenuate; the inner sepals smaller (narrower and shorter), glabrous or sparsely to

densely stellate. Corollas 1-2.5 cm long, funnelform to campanulate, white, glabrous. Stamens unequal, 5-12

mm long; anthers 1-2 mm long. Ovary 1.5 mm long, ovoid to cylindrical, glabrous; styles 6-11 mm long. Cap-

sules 4-5 mm long, ovoid, partly enclosed by the sepals. Seeds 1-4, 2.7-3 mm long, rotund, brown, minutely

Sonora.— From near Hermosillo and the east-central part of the state southward and e;

desert, coastal thomscrub, and tropical deciduous forest; near sea level-770 m. Flowering a

March, September-December.

General distribution— Widespread in Mexico including the Sierra Madre Occidental i

Oriental, and Sierra Madre del Sur, and at least in Chihuahua, Guerrero, Edo. M<

Tamaulipas, Veracruz; from desert margins to oak and pine-oak zones

Robertson (1971: 133) reported that J. albida is closely related to J. polyantha, “which has outer sepals that

are broadly ovate or ovate and pubescent instead of elliptic and glabrous.” He knewj. albida only from the vi-

cinity of Hermosillo. Many additional specimens available to us from a wide range indicate that differences

between J. albida and J. polyantha are too few and do not distinguish them. As Robertson suspected, we con-

clude that they are best treated as a single species.

Robertson (1971: 168-169) also pointed out that, “Both J. polyantha and J. pentanthos are very closely re-

lated.” Primary differences between J. polyantha and J. pentanthos are in inflorescences, sepals, and flower color.

Jacquemontia pentanthos has compact cymes and usually rhomboidal outer sepals with long attenuate apices; J.

Polyantha tends to have open cymes and elliptic to ovate outer sepals and acute or acuminate to short-attenuate

apices. Moreover, J. pentanthos has blue flowers and those of J. polyantha are usually white.

516

Perennial vines, the stems twining, with 4-6-armed stellate trichomes, rarely glabrate. Leaves broadly ovate

to ovate, 2-6.5 cm long, 1.5-4.8 cm wide, basally shallowly cordate to truncate, apically acute or less often re-

tuse to obtuse, sometimes mucronate. Inflorescences axillary, loosely cymose or solitary. Flowers 1-7 on pe-

duncles 1-1 1 cm long, the pedicels 2-10 mm long, erect to nodding in fruit; bracts linear, to 6 mm long. Sepals

subequal, the outer ovate to broadly ovate, the middle sepal falcate, the inner narrowly ovate, 5.5-9 mm long,

the apices acute, pubescent without and to a lesser degree within. Corollas 1.4-2.7 cm long, funnelform, laven-

der to white, glabrous. Stamens 5-10 mm long, included, anthers 1.5-2 mm long. Ovary 1-1.5 mm long, ovoid,

2-locular, glabrous; styles 7-9 mm long. Capsules 5-6 mm long broadly ovoid, enclosed by accrescent sepals.

Seeds 1-4, 2-2.5(-3) mm long, trigonous, brown, minutely areolate and ruminate.

Sonora .— Widespread except the northwestern corner of the state; Chihuahuan and Sonoran deserts,

coastal and foothills thomscrub, and tropical deciduous forest. Near sea level-1000+ m. Flowering (March)

August-December.

General distribution. — Pima, Yuma and Cochise Cos, Arizona; southwestern Chihuahua to northwestern

Sinaloa.

Felger et al., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

517

Lianas or herbaceous annual or perennial vines (also flowering in first season or perhaps annuals), the stems

twining or prostrate. Leaves simple or palmately compound, ovate, hastate, sagittate, variably palmately or

pinnately divided, the segments 3-9, ovate-linear, the margins entire or sinuate-serrulate, sessile or if present

the petiole slender, cylindrical, occasionally sulcate, or rarely scale-like. Inflorescences axillary, in dichasia or

monochasia, sometime umbellate. Flowers 1-40, mostly diurnal. Peduncles usually similar to the stems and

petioles or reduced or absent, the pedicels usually shorter than the peduncle, smooth, striate, or notably five-

angled, slender, stout to distinctly clavate. Bracts glabrous or pilose, usually two, prominent and foliaceous to

reduced or scale-like or absent, usually caducous or fugaceous, rarely persisting in fruit, the bracteoles, when

present, similar to the bracts. Sepals 5, persistent, imbricate, ovate-lanceolate, herbaceous, membranaceous-

coriaceous, glabrous, pilose-appressed, or hirsute, the margins entire, the apex acute to obtuse, attenuate, or

rarely emarginate. Corollas campanulate to funnelform, the tube widening gradually or abruptly, the limb

more or less entire, the lobes 5-10, white, yellow, or rose-purple, usually glabrous. Stamens 5, included, in-

serted at the base of the corolla tube, erect, glabrous or pubescent with glandular trichomes; anthers helicoid-

contorted; pollen 3-colpate, or polycoplate (Austin et al. 2008; Ferguson et al. 1977). Style 1, white, glabrous,

the stigma 2-globose or superficially 4-globose. Fruits capsular, globose to more or less conical, brown to

straw-colored or gray or transparent when dry, entire or 4-lobed, the locules 2-4, the valves usually 4, glabrous,

dehiscence valvular or irregular. Seeds 1-4, brown or black (straw-colored in M. aegyptia ), rounded or 1-3-an-

gled, glabrous, puberulent, or tomentose.

About 100 species, largely in tropical zones; 27 species known from the Americas.

Selected references — Austin (1979, 1995), Gunn (1977), O’Donell (1941). Ana Rita Simdes (atBM, working

on the Merremieae, pers. comm. 2011).

11862],

Stems to 6 m long, herbaceous at least above, with appressed trichomes. Leaves palmately compound, the

leaflets 5, 1.5-6 cm long, 0.7-1.4 cm wide, ovate, the base decurrent, the margins entire, undulate, dentate or

dentate-sinuate, the apex acuminate, membranaceous, glabrous or appressed-pubescent, strigose or hirsute on

both surfaces. Petioles 3-5 cm long, hirsute-pilose to glabrous and/or glandular. Inflorescences of monocasia

519

Felgeretal., Convolvulaceae (excluding Cuscuta) of Sonora, Mexico

(USONt). Mpio Baviaccora: Suaqui along SON 89, 29 0 41'46''N, 1 10°09'18"W, 680 m, solitary vine to 3 m in tree away from houses, flowers

Merremia palmeri (S. Watson) Hallier f., Jahrb. Hamburg. Wiss. Anst. Beih. 3:38. 1899. Type: Mexico. Sonora:

Trompiuo, huirote (these are general names; trompillo for any morning glory and huirote in Mexico for vine)

Large, robust perennial vines, glabrous, with milky sap, the stems to 8 m long; growing and flowering during

hot weather, often covering trees and shrubs, and leafless and dying back severely in dry seasons. Leaves

broadly ovate to circular in outline, palmately compound; leaflets 5, lanceolate 5-14 cm long, 0.6-2.5 mm wide

(the median one largest), basally attenuate, apically truncate and mucronate, the margins entire or nearly so;

petioles 1.5-2.5 cm long. Inflorescences solitary, axillary. Flowers nocturnal (opening around dusk and

quickly wilting with the early-morning heat), on peduncles greatly exceeding the petioles, the pedicles 2.2-4

cm long, the bracts deltoid-ovate and apically caudate, 4-5 mm long, 1-1.5 mm wide. Sepals 2.6-3.5 cm long,

in fruit and becoming pock-marked with glands on the inner surfaces. Corollas 5.5-7.3 cm long, 4-6(-8) cm

wide, white, salverform to funnelform-salverform, glabrous. Capsules brown, globose to ovoid, 1.5 cm long

and wide, subtended and partly enclosed by the accrescent sepals. Seeds 2-4, 9-11 mm long, 6-9 mm wide,

black, wedge-shaped, puberulent when fresh, glabrescent.

Endemic to Sonora and Sinaloa: Sonoran desert, coastal and foothills thomscrub, and tropical deciduous

forest. Northern Sonora from the vicinity of Trincheras, Benjamin Hill, and Presa Angostura southward to

Sinaloa; 10-800 m. Flowering recorded (January) March, May-October.

This species is self-incompatible. The nocturnal flowers are pollinated by hawk moths, but excess pollina-

tor visits can lead to a decline in female reproductive success as a result of flower damage or pollen interference

(Willmott & Burquez 1996). Alberto Burquez (personal communication 6 October 2001) writes that “the flow-

ers emit a faint, sweet fragrance after opening that lasts through the night. Hawkmonths use it as a long-dis-

tance cue. When close, they use eyesight and proboscis.” In contrast, pollination ecologist Robert A. Raguso

described the fragrance of flowers remaining open in the morning as being disagreeable (specimen label for

Raguso RAR 98-162).

The closely related species M. platyphylla (Femald) O’Doneli appears to have bat-pollinated flowers. M.

palmeri is also related to the Mexican M. tuberosa (L.) Rendle and M. discoidesperma (Donn. Sm.) O’Donell

(Austin 1998c).

m, 6 Nov 1982, Starr 192'. Mpio Sahuaripa: Mountains 6.7 mi Wof Sahuaripa, Gates 9 Sep 1959'. Mplo Soyopa: NE side of Rio Yaqui bridge

Perennial or sometimes annual herbs, twining or prostrate, glabrous or sparsely hirsute, the stems herbaceous

toward the apex, woody toward the base, to 5 m long. Leaves 1.5-5 cm long, 2.5-7 cm wide, palmately com-

pound, circular in outline, the leaflets 5, elliptic or lanceolate to oblanceolate, the margins sinuate-serrulate,

the segments more or less sessile, apically and basally acute to acuminate, glabrous. Petioles 2-9 cm long, gla-

brous or with a few scattered patent trichomes. Inflorescences of monochasia or dichasia. Flowers 1-10, diur-

nal. Sepals unequal, the outer ones 3-5 mm long, the inner ones 4-7 mm long, oblong, ovate to elliptic, coria-

ceous to chartaceous, the margins scarious or not, obtuse, mucronulate, glabrous. Corollas 1. 5-2.5 cm long,

campanulate, cream to white, glabrous. Capsules 5-8 mm long, more or less globose, straw-colored to brown,

glabrous, the sepals partly surrounding and covering the fruit. Seeds 1-4, 3-5 mm long, straw-colored or

black, pubescent with short, brown trichomes.

Sonora. — Southeast and east-central part of the state in tropical deciduous forest , often in disturbed sitesand

riparian habitats near rivers and along arroyos; 120-460 m. Flowering March-May and September-October.

General distribution. — Florida; Baja California Sur, Chiapas, SW Chihuahua, Colima, Guerrero, Jalisco,

Edo. Mexico, Michoacin, Nayarit, Oaxaca, San Luis Potosi, Sinaloa, Tamaulipas, Veracruz; Mesoamerica; Co-

lombia, Venezuela, Ecuador, Peru, Guayanas, Brasil; Antilles.

OPERCULINA Silva Manso, Enum. Subst. Braz. 16. 1836. [From Latin operculum, a lid or cover, and -inus,

pertaining to, referring to the covering of the capsule.]

Lianas or small twining herbs, the stems prostrate or climbing, to 6 m long, smooth or striate, glabrous.

Leaves ovate, broadly ovate, pinnatly or palmately lobed or compound, the segments 5 or 7 or entire, glabrous;

petioles and pedicels sometimes winged, mostly glabrous. Inflorescences in axillary monocasia. Flowers few

or solitary, often with foliose bracts. Sepals equal or unequal, enlarging in the fruit and becoming coriaceous,

sometimes irregularly dentate on the margins, glabrous. Corollas broadly campanulate, funnelform or salver-

form, white, yellow, or reddish to salmon, the interplicae pilose, the plicae glabrous. Stamens included (ex-

serted in O. pteripes); anthers twisted when fully mature; pollen 3-colpate. Ovary glabrous, bilocular, each

locule 2-lobed; style included (exserted in O. pteripes), filiform; stigma of 2 globose lobes. Fruits dehiscent, the

upper part separating by a circumscissile epicarp, the upper part more or less fleshy and separating from the

lower segment and from the endocarp, 2-locular. Seeds 1-4, ovoid to ovate, glabrous or pubescent.

Species about 15; 10 known from only the Old World.

Selected reference. — Staples and Austin (1981).

Gaujma, pata be GALLO (Friedman 1996); tansy-leaf lid-pod (USDA).

Perennial herbs, the stems twining or prostrate, to 6 m long, sometimes angular, glabrous. Leaves 2-12 cm

long, 2-11 cm wide, ovate in outline, pinnatisect to palmately compound, the segments 5-9, linear, lanceolate

toelliptic-obovate, apically obtuse-acuminate, basally truncate or auriculate, with few trichomes on upper and

lower surfaces, the margins entire. Inflorescences of monocasia. Flowers 1-3, on winged peduncles (at least in

the upper part), glabrous. Sepals 1.1— 1.6 cm long, equal, straw-colored to slightly rosy during anthesis, ovate to

obovate, accrescent in fruit, membranaceous, glabrous. Corollas 3.4-5.3 cm long, campanulate, white, the limb

more or less entire, widening gradually, the interplicae sericeous. Capsules 1.2-2 cm long, transparent,

brown, glabrous. Seeds 1-4, 5-7 mm long, ellipsoidal, black, glabrous.

Sonora .— Arroyos in coastal thomscrub in the far southwestern part of the state and foothills thomscrub

in the central part of the state; 10-730 m. Probably flowering May-October

General distribution— Texas; Guerrero, Edo. Mexico, Michoacan, Morelos, Nuevo Leon, Oaxaca, San Luis

Potosi, Sinaloa, Tamaulipas, Veracruz; Mesoamerica.

Through much of its range this species is known as queibra-platos and is considered such a drastic laxative

that even handling the plants will cause dishes to break at home (Alcorn 1984). In some areas it is an important

medicinal plant.

°perculina pteripes (G. Don) O’Donell, Lilloa 23:435, t. 6, 1950. Type: ECUADOR: Guayaquil, Ruiz & Pawn mil

1778-88 (printed label), 1800 (typed label) (holotype: MA!; isotype: F!). Cahmyction pteripes G. Don, Gen. Hist 4:264. 1838.

522 Journal of the Botanical Research Institute of Texas 6 ( 2 )

Campanula chocolate

Perennial herbs or lianas, glabrous, the stems twining, pendulous, or prostrate, reaching 5 m long, becoming

woody toward the base, striate to angular, glabrous. Leaves 3-17 cm long, 2-8 cm wide, simple, entire, ovate

to broadly-ovate, apically acute to acuminate, attenuate, mucronate, basally cordate to almost truncate; petioles

to about half as long as the blades. Inflorescences monocasial to dicasial. Rowers (1)2-12, on peduncles with

3 wings in the central part 0.6-3.5 mm wide, becoming attenuate toward both ends. Sepals more or less equal,

the outer ones 2.2-2.6 mm long, ovate to ovate-elliptic, the apex acute to obutse, the inner ones 2.3-2.5 cm

long, ovate, obtuse, glabrous or more often pubescent near the base, surrounding the base of the corolla. Corol-

las 4-7 cm long, salverform, the limb more or less entire, broadening abruptly, reddish or red-orange to salm-

on, tomentulose on the tube and the interpliae. Capsules 1.5-2 cm long, transparent, brown, glabrous. Seeds

1-4, 7-9 mm long, ovoid to ellipsoid, black, glabrous.

Sonora — Foothills thornscrub, tropical deciduous forest, and oak woodland in the southeastern and

east-central part of the state; 240-1250 m. Flowering July-October.

General distribution. — Chiapas, southwestern Chihuahua, Colima, Distrito Federal, Edo. Mexico, Guer-

rero, Jalisco, Michoacan, Morelos, Nayarit, Oaxaca, Sinaloa; Mesoamerica; Colombia, Venezuela, Ecuador,

Peru.

The flowers of this vine have the shape and color of bird flowers and are visited by hummingbirds. There

is an incredible variation in colors of the flowers even within the limited range available. No study of pollina-

tion or color variation has been made beyond casual observations. However, the various names given to it, and

the many illustrations from the 1800s onward show the fascination that Europeans had with the flowers.

Mpio Alamos: Arroyo el Mentidero at El Chinal road, 11.3 km S of Alamos, 26°54'45”N, 108 o 55'05"W, 240 m, 20 Sep 1993, flowers red. Van

Devender 93-843!; Rio Mayo Raft trip, 2 km beyond Palmarito on bank of Rio Mayo, 27°53'N, 108°48'W, 26 Sep 1991, Rondeau & Jenkins 91-

180 (FTG-FAU!); 1.3 km S of Guirocoba Road, 3.3 km S of Alamos on road to El CKinal, Sinaloa, 26°59'30"N, 108°55'25"W, 340 m, 11 Oct

ACKNOWLEDGMENTS

Over the years many friends and colleagues have provided information and assistance, and/or accompanied us

in the field. In this regard, we thank Thomas Bowen, Kathy Bunnell, Alberto Burquez-Montijo, Mark A. Dim-

mitt, Exequiel Ezcurra, Ana Luisa Rosa Figueroa-Carranza, Lloyd Findley, Mark Fishbein, Francisco Molina-

Freaner, Juan Pablo Gallo-Reynoso, Pedro Garcillan, Edward Erik Gilbert, Powell B. “Gill” Gillenwater, lllji®

Henrickson, Cathy Moser Marlett, the late Paul S. Martin, Angelina Martinez-Yrizar, J. Andrew McDonald,

William (Bill) Risner, the late Alexander Russell, Jean Russell, Andrew C. Sanders, Silke Schneider, Barbara

Straub, Raymond Marriner Turner, Benjamin T. Wilder, and Michael F. Wilson.

The staff at several herbaria have been especially helpful of our multiple requests and have provided sig-

nificant assistance: Bradley (Brad) Lome Boyle, Benjamin Daniel Brandt, W. Eugene Hall, Sarah Hunkins,

Philip D. Jenkins, and Michelle (Shelley) McMahon (ARIZ); Comision Nacional de Areas Naturales Protegidas

(CONANP); Prescott College Kino Bay field station; Jon P. Rebman, John F. Sanborn, and especially Judy Ann

Gibson who provided innumerable search results and other data (SD); Herbarium of the University of Sonora

(USON); and Herbarium of Centro de Investigaciones Biologicas del Noroeste (HCIB).

People at many herbaria provided generous assistance. In this regard we especial thank: Anne Barber an

Elizabeth Makings (ASU), Ana Rita Simoes (BM), Ria DAversa and Debra Track (CAS), Wendy Caye Hodgson

523

and Andrew Michael Salywon (DES), Jacqueline Kallunki (NY), Layne Huiet (DUKE), Brett Jestrow (FTG),

Laurent Gautie (G), Brian Franzone, Melinda Peters, Julie Shapiro, and Emily Wood (Harvard University Her-

baria), Kenneth R. Robertson (ILLS), Anita F. Cholewa (MIN), James C. Solomon, Cynthia Strickland (MO);

Richard Spellenberg (NMC), Tom Zanoni (NY), Ronald L. Hartman and Burrell E. Nelson (RM), Sula E. Van-

derplank (RSA); George Staples (SING); J. Andrew McDonald (PAUH), Andrew S. Doran (UC); Andrew C.

Sanders (UCR); Maria Teresa Buril (UFP), and Larry Hufford and Mare Nazaire (WS).

Ana Lilia Reina-Guerrero translated the abstract for the resumen and Pedro Garcillan drafted the map.

Two reviewers, J. Andrew McDonald and Javier Ortega provided constructive comments improving the manu-

From Tom Van Devender: I thank my wife Ana Lilia Reina-Guerrero for 16 years of botanical adventures

in her native Sonora. I thank Father William Trauba, Capuchin Franciscan Missionary, for sharing our field

work in the Yecora area. The late Paul S. Martin inspired us and a talented group of botanists in the Rio Mayo

area of southern Sonora.

The following herbaria have provided Cuscuta plant material: AAU, ALTA, ARIZ, ASU, B, BAB, BOL,

BRIT, CANB, CAS, CEN, CHR, CHSC, CIIDIR, CIM1, CTES, DAO, F, G, GH, H, HUFU, IAC, IEB, IND, J, JEPS,

LL, LP, LPB, LPS, K, MEL, MERL, MEXU, MICH, MO, NMC, NY, OAC, OKLA, OSC, OXF, PACA, PRE, QCNE,

QFA, P, PACA, RB, RSA, SAM, S, SD, SGO, SI, SPF, TEX, TRT, TRTE, UA, UB, UBC, UCR, UCT, UNB, UNM,

UPRRP, UPS, US, USAS, WTU and XAL. Cuscuta research was supported by a Natural Sciences and Engineer-

ing Research Council of Canada Discovery grant to Costea (327013-06 and 327013-12). Felger acknowledges

support from the Wallace Research Foundation and the David and Lucile Packard Foundation. Botanical col-

lections were made under Mexican Federal collecting permits including NOM-126-SEMARNAT-2000 with

the generous assistance of Exequiel Ezcurra.

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528

BOOK REVIEW

Emily Monosson. 2012. Evolution in a Toxic World: How Life Responds to Chemical Threats. (ISBN-13: 978-

1-59726-976-6, hbk.). Island Press, 1718 Connecticut Ave., NW, Suite 300, Washington, DC 20009,

U.S.A. (Orders: http://islandpress.org/index.html). $35.00 hbk., $29.95 pbk., 240 pp., 10 figures, notes,

“The best way to envisage the situation is as follows: the environment presents challenges to living species, to

which the latter may respond by adaptive genetic changes.” — Theodosius Dobzhansky.

. . . And this is exactly what Emily Monosson discusses throughout the ten chapters of Evolution in a Toxic

World: How Life Responds to Chemiccd Threats. This 222-page book is a thought-provoking summary of an im-

portant but often ignored subject matter, environmental toxicology. The introduction (Chapter 1) is a fascinat-

ing overview of the subject matter. Monosson opens her introduction with two powerful sentences: “All of life

is chemical. But not all chemicals are compatible with life.” Think about that contrast! The author gives an

overview of the four sections of the book: 1) Nature’s Toxicants (everything is a poison in the right dose), 2)

Evolutionary History of Toxicology, 3) Toxic Evolution in Action, and 4) Looking Forward by Looking Back.

Chapter 2 — Shining a Light on Earth’s Oldest Toxic Threat, looks at UVR, a highly energetic and destruc-

tive force to be reckoned with. We all know metals like arsenic and the secondary plant metabolite strychnine

are poisonous, but Monosson throws at the reader chemical toxicants that most of us don’t think about, like

oxygen. Monosoon points out, “we cannot live without it, yet every day we struggle to coexist with this highly

reactive and potentially toxic chemical.” Oxygen is discussed in detail in Chapter 3 — When Life Gives you

Oxygen, Respire. Chapter 5 looks at the many toxic metals that life deals with. Chapter 6 discusses chemical

warfare ... the battle to protect and to survive. The combatants? Plants and animals. Sounds like a sci-fi movie,

and it very well could be. Maybe people would watch and then take notice of the thousands of chemicals in our

environment. If you like war, then Chapter 6 is a captivating account of this historical battle and challenge to

stay alive; it’s my favorite chapter in the book. This is not to say that the rest of the book is not interesting. Quite

the contrary. Chapter 7 — Sensing Chemicals is enticing, Chapter 8 — Coordinated Defense is inviting, Chap-

ter 9— Toxic Evolution is captivating, and Chapter 10 is all about toxic overload: “How will life’s toxic defense

mechanisms respond to industrial age chemicals?” And yes, there are many. “In 2009, the Chemical Abstracts

Service, which catalogs and tracks all known chemicals, announced the registration of its fifty-millionth

‘novel’ chemical — the last ten million chemicals having been registered over the preceding nine months. There

are plenty more novel chemicals to be found.” In the conclusion, Monosson writes that this toxic world we live

in is challenging to each and every human. We do live in a sea of toxic chemicals. “Life on Earth is now subject

to a virtual onslaught of chemicals associated in one way or another with human activity. We are a society built

on chemicals, and there is no turning back ” Monosson says that we must strive to better understand how

chemicals affect wildlife and human health. “We have to do so. There is no higher ground, no corner on earth

where life can escape the influence of toxic chemicals. The choice must not be to ‘evolve or die.’”— Barney

Lipscomb, Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, USA

). Bot. Res. Inst. Texas 6(2): 528. 2012

CONVOLVULACEAE OF SONORA, MEXICO. II: CUSCUTA

Mihai Costea Richard S. Felger

Department of Biology

Wilfrid Laurier University

75 University Avenue W

Waterloo, ON, N2L 3C5, CANADA

Daniel F. Austin

Arizona-Sonora Desert Museum

2021 N. Kinney Road, Tucson, Arizona 85743, U.SA.

and Herbarium, University of Arizona

P.O. Box 21 0036, Tucson, Arizona 8572 1, U.SA.

Herbarium, University of Arizona

P.O. Box 210036, Tucson, Arizona 85721, U.S.. ,

and Sky Island Alliance, P.O. Box41165

Tucson, Arizona 85717, U.S.A.

Thomas R. Van Devender

Sky Island Alliance, P.O. Box 41 165

Tucson, Arizona 85717 and Herbarium

University of Arizona

P.O. Box 21 0036, Tucson, Arizona 85721, U.SA

VanDevender@skyislandalliance.org

J. Jesus Sanchez-Escalante

Dept, de Investigaciones Cientificas y Tecnoldgicas

Rosales y Ninos Heroes, Centro

Hermosillo, Son, 83000, MEXICO

RESUMEN

INTRODUCTION

^ is the second part of a comprehensive study regarding the Convolvulaceae of Sonora (Felger et aL 2012).

® the firs t Part, we included a general introduction, information about the growth forms, cases of endemism/

f s P«te, species diversity, and types of vegetation for all the genera/species. The first part also provided

Wentihcation keys, and extensive taxonomic and floristic data for all the Convolvulaceae genera except Cus-

530

Journal of the Botanical Research Institute of Texas 6(2)

cuta (dodder). Since the diversity of dodder species and their association with certain vegetation types were

presented together with the other Convolvulaceae (Felger et al. 2012), this article concludes the monographic

study of this botanical family in Sonora with a taxonomic and floristic account of the genus Cuscuta.

Cuscuta is nearly cosmopolitan and comprises over 200 species (Costea 2007-onwards). Although some

dodders are agricultural pests (Costea & Tardif 2006), more numerous Cuscuta species require conservation

measures (Costea & Stefanovic 2009). Roughly 75% of species are native to the Americas and about 30% to

Mexico and southern USA (Stefanovic et al. 2007). We have documented 21 species of Cuscuta in the flora of

Sonora, a number that is higher or comparable to that of the neighboring areas to the north and west (Arizona,

the two Baja California states, California, and New Mexico; see table 1, Felger et al. 2012). Interestingly, while

the species diversity of the other Convolvulaceae genera increases towards southern Mexico and Central

America, dodder species richness declines sharply in these geographical areas (Felger et al. 2012). The 21 spe-

cies that occur in Sonora belong to nine of the 15 clades of subgenus Grammica, the most complex infrageneric

taxon of Cuscuta (Stefanovic et al. 2007). The best represented in Sonora is the C. umbellata clade (clade “L”)

with seven of the nine North American species growing in Sonora (C. desmouliniana, C. legitima, C. leptantha,

C. odontolepis, C. polyanthemos, C. tuberculata, and C. umbellata). This strongly suggests that Sonora is part of

the genetic center of origin for this clade, which has a complicated evolutionary history shaped by reticulate

evolution (Costea & Stefanovic 2010). The other eight clades of subg. Grammica are represented in Sonora by

one species (C. salina, clade “A”; C. tinctoria, clade “G”; C. corymbosa var. grandiflora, clade “J”, C. indecora, clade

“M”; C. campestris, clade “B”; C. vandevenderi, clade “N”), two species (C. azteca and C. chinensis var. applanata,

Clade “H”; C. americana and C. macrocephala, clade “D, or three species (C. boldinghii, C. costaricensis, andC.

erosa, clade “K”).

CUSCUTA L.,Sp. PI. 124. 1753. [Based on the Aramaic and Hebrew triradical root of the verb K-S-Y PH Kaph, ® Shin, ’ Yodh), XT.

Common names.— Dodder; fideo

Herbaceous vines. Stems filiform, yellow or orange, trailing or dextrorsely twining and attached to the host by

numerous small haustoria, glabrous. Leaves reduced to minute, alternate scales. Inflorescences monochazial

clusters further grouped in cymose inflorescences that are often confluent. Flowers 4-5-merous, small, always

± fleshy when fresh, thick or membranous-thin when dry, white, white-cream, sometimes yellowish or red-

dish. Conic-cylindrical papillae present or absent on the pedicels, perianth and ovary/capsule; laticifers visible

or not in the calyx, corolla, ovary/capsules, translucent, white, yellow or orange, isolated or arranged in rows

especially in the midveins of the calyx and corolla lobes, round, ovoid or elongated. Calyx gamosepalous; lobes

basally overlapping or not, sometimes with multicellular projections (C. chinensis var. applanata, C. boldinghii)-

Corolla gamopetalous with lobes imbricate in bud, sometimes with a subapical cusp or horn-like multicellular

appendage (C. boldinghii, C. costaricensis, C. erosa). Stamens alternating with the corolla lobes. Pollen 3-colpate

(sometimes 4- or 5-colpate in the same anther), tectum imperforate to reticulate. Infrastaminal scales com-

monly present, scale-like appendages dentate or fringed, bridged and adnate to the corolla tube base, forming

a corona alternating with the corolla lobes. Ovary superior, 2-locular, each locule with 2 anatropous ovules.

Styles 2, terminal, distinct or united, equal or unequal. Stigmas spherical to linear (only distinct unequal styles

with spherical stigma in the species from Sonora). Fruits capsules, indehiscent (sometimes opening irregu-

larly between the styles) or circumscissile by a ± regular line near the base. Seeds 1-4 per capsule, 3-angled or

dorsoventrally compressed; endosperm nuclear; embryo uniformly slender, 1-3-coiled, without cotyledons,

consisting mostly from the hypocotyl; seed coat alveolate when dry and papillate when hydrated (rarely not

alveolate/papillate with cells ± rectangular, puzzle-like arranged).

Cuscuta campestris and C. indecora are weeds, the former subcosmopolitan, the latter widespread in North

and South America. Both species have been likely introduced to Sonora with contaminated legume seeds (e g -

alfalfa; Costea & Tardif 2006). Another species, C. tinctoria, may also have been introduced to Sonora on the

cultivated tree Schinus terebinthifolia. Although none of the Cuscuta spp. are endemic to Sonora, many occur

only here and in adjacent geographical areas (Table 1, Felger et al. 2012). The diversity of Cuscuta species in

Sonora is most similar to that of the Baja California Peninsula and Arizona (12 and 11, respectively common

species), followed by New Mexico (9 common species), while only three of the Sonoran species occur in

California (Table 1, Felger et al. 2012).

Selected references . — Stefanovid et al. (2007), Welsh et al. (2010), Wright et al. (2011, 2012), Yuncker (1932,

1965).

Identification of most Cuscuta spp. is a lengthy process because rehydration of flowers, dissection, and

examination under a microscope are usually necessary. Measurements of floral parts were done on rehydrated

herbarium material. Length of flowers was measured from the base of calyx to the tip of straightened corolla

lobes. The texture of flowers and the color of calyx were noted on dry herbarium material. Observation of pa-

pillae and laticifers requires magnifications of at least 100 x. Examination of seed surface requires magnifica-

tions of at least 150 x. In describing the stem, the following categories based on stem diameter were used

(Yuncker 1921): “slender” with the diameter of 0.35-0.4 mm, “medium” with the diameter of 0.4-0.6 mm, and

“coarse” when diameter is greater than 0.6 mm. The geographical distribution, both in Sonora and Mexico, is

The host range is also based on herbarium labels as well as observations by Richard Felger; hosts ob-

served in other geographical areas are included when they are present in Sonora.

SEM images of the flowers for some species are provided to help identification. The vouchers are indicated

(“SEM”) in the lists of typical collections examined. Pictures were taken with the scanning electron micro-

scopes Hitachi S-570 and LEO 1530 FE-SEM at 15 KV. Samples were coated with 30 nm gold using an Emitech

K 550 sputter coater. Numerous images for all the species, including the types, with details of dissected flowers

are available from Digital Atlas of Cuscuta (Costea 2007-onwards). All specimens cited have been seen by

Costea and are at ARIZ unless otherwise indicated. For citation of herbarium specimens see Felger et al. (2012).

Plants not native to flora area are marked with an asterisk (*).

532

-0.5 x 0.2-0.3 1

. nil

537

desert grassland, 31°11'23"N, 110°36’22"W, 1340 m, on Ambrosia confertiflora, 17 Aug 2001, Von Devender 2001-710 (ARIZ, USON, WLU)

[SEMI.

Cuscuta corymbosa Ruiz & Pav. var. grandiflora Engelm., Trans. Acad. Sci. St. Louis 1:483. 1859; Syst. Ar-

rang. Sp. Cuscuta 33. 1859. (Engelmann published the same name in both publications, the same year).

Type: COLOMBIA, [from Popaydn?]: Humboldt 2002, ex herb Witldenow 3157 (lectoiyfe here designated: B!; fragment at MO!). En-

. Sulphur 35. 1844. Type MEXICO. Baja California Sur: Magdalena Bay, Bentham s.n. (K!).

ay. Sulphur 138. 1844. Type: MEXICO. Guerrero: Acapulco, Bentham s.n. (K!).

Stems medium, orange. Inflorescences dense, corymbiform or umbellate, often confluent. Pedicels 2-6 mm

long. Bracts 1 at the base of clusters, absent at the base of pedicels, 0.5-1 mm long, ovate lanceolate, margins

entire, apex acute. Flowers 5-merous (Fig. le), 4.5-6.5(-7) mm long, membranous, white when fresh, creamy

brownish when dried, papillae absent. Laticifers barely visible in the calyx and corolla lobes, isolated, ovoid to

elongate. Calyx 2-2.5 mm long, straw-yellow to brownish, membranous, finely reticulate, not shiny, cylindric

campanulate, V2- 3 /4 as long as the corolla tube, divided ca. l A its length; tube 1.1-2 mm long; lobes 0.4-0.75 mm

tong, not overlapping or only slightly so, rounded, not carinate, margins entire. Corolla 4-6 mm long; tube 3-5

nun long, cylindric, becoming dilated in the lower W; lobes 1-1.5 mm long, initially erect, later spreading, VS-Va

of the corolla length, ovate, mar gins entire, apex obtuse to rounded, ± incurved. Stamens included, shorter

than the corolla lobes; filaments 0.1-0.3 mm long; anthers 0.5-0.6 x 0.3-0.4 mm, subround to elliptic. Infra-

staminal scales 1.5-2 mm long, Vi-% as long as the corolla tube, bridged at 0.2-0.35 mm, forming oblong

ridges with fringed margins, fimbriae 0.4-0.15 mm long. Styles 2.4-4.2 mm long, much longer than the ovary,

evenly filiform. Capsules circumscissile, 2-2.9 x 2.2-2.6 mm, globose to slightly depressed, not thickened or

risen around the small interstylar aperture, translucent, surrounded by the withered corolla. Seeds 2-4 per

capsule, 1.1— 1.3 x 0.7-0.9 mm, slightly angled, broadly elliptic, sometimes with a longitudinal groove on the

ventral face; seed coat cells alveolate/papillate.

Sonora.— Locally in Gulf Coast of the Sonoran Desert in a large, ecologically pristine canyon in the Guay-

mas region opposite Isla San Pedro Nolasco and in foothills thornscrub in east-central Sonora; also on Islas San

Pedro Nolasco and San Esteban; ca. 5-450 m. Flowering December-April. Probably more widespread in the

state. Parasitic especially on Colubrina viridis and Vaseyanthus insularis (Felger et al. 2011); alsc

urtfornicum, Acalypha califomica, Cottsia linearis (Janusia linearis ),

n ka, and Poaceae.

General distribution.— One of the most common species in Mexico, where it is sometimes weedy. Baja

California Sur, Chiapas, Colima, Durango, Edo. Mexico, Guerrero, Guanajuato, Hidalgo, Jalisco, Michoacan,

Morelos, Nayarit, Sinaloa, Tamaulipas, Veracruz; also in Central and South America.

Selected references — Felger and Wilder (2011, 2012), Yuncker (1932).

M P» Guaymas: Caft6n las Barajitas Sierra el Aguaje, 28°03'27"N, lll o 09’27"W, 90 m, 19 Feb 1995, Felger 95-208. Mpio Soyopa: 0.6 km N

16 on road to T6nichi just E of Rfo Yaqui, 28.5758°N, 109.5505°W, 270 m, foothills thornscrub, on Janusia linearis, 7 Jan 2001. Van

I96« ,der2001 ' 16 SONORAN ELANDS. San Esteban: N side of island, o

S ?' Fdger 1755 °- St eep N slope of NE peak, 28°42'N, 112 0 35'W, 450 m, o

® Nolasco: NE side of Island, 18 Jan 1965, Felger 12082 [S

5, Felger 12082; l'

i. TorreyBot. Club 18:227. 1932. i

e: MEXICO. Durango: Santiago Papasqui;

Journal of the Botanical Research Institute of Texas 6(2)

The type locality is in pine-oak forest and the host plant is Bouvardia temifolia. In addition to the type collec-

tion, C. dentatasquamata is recorded from a canyon in southern Arizona mountains in oak woodland and

should be sought elsewhere with Bouvardia in southern Arizona and northern Sonora mountains.

General distribution. — Arizona; Chihuahua, and Sonora; apparently very rare.

Selected reference. — Yuncker (1932).

S Biol. Monogr. 6(2-3):40-41. 1931. Type: MEXICO. SONORA: hills near Altar,

UZ!, ASU1, Gl, GH!, IND, MEXU!, MO!, NY!, US!).

iol. Monogr. 6(2-3):41. 1921. Regarding Pringle 105, Yuncker (1921, p 41) raen-

Following two distinguishable varieties (“typical” and “attenuiloba”).

rr. 6(2-3):43. 1921. Type: MEXICO. Sonora: Guaymas, 22 Feb 1904, Palmer 1209

Stems slender, yellow-orange. Inflorescences loose, umbellate, often confluent. Pedicels 1-5 mm long. Bracts

1 at the base of clusters and 0-1 at the base of pedicels, 0.6-1 mm long, ovate-lanceolate, margins entire, apex

acute. Flowers 5-merous (Fig. 3d), 2-3 mm long, membranous, white when fresh, creamy-white when dried.

Papillae usually present on the pedicels, calyx, abaxial and adaxial epidermis of corolla lobes, and sometimes

on the ovary/capsule. Laticifers not visible or hardly so in the midveins of corolla lobes, elongate. Calyx 0.6-1.2

mm long, brownish-yellow, ± reticulate or shiny, campanulate, equaling or somewhat longer than the corolla

tube, divided VS-Vi the length; tube 0.25-0.5 mm long; lobes 0.5-0.76 mm long, not overlapping, triangular-

ovate to lanceolate, weakly to distinctly carinate, with small dome-like multicellular projections on the mid-

veins, margins irregular, ± revolute at the base and forming angled sinuses (especially when lobes are triangu-

lar ovate), apex acute to acuminate. Corolla 1.5-2.9 mm long; tube narrow-campanulate, 0.8-1.5 mm long

lobes 1-1.5 mm long, initially erect, later spreading or reflexed, slightly longer than the tube, lanceolate, mar-

gins entire sometimes involute upon drying and appearing very narrow, apex acute, ± incurved. Stamens

short-exserted, shorter than corolla lobes; filaments 0.4-0.7 mm long; anthers 0.4-0.6 x 0.2-0.3 mm, ovate to

oblong. Infrastaminal scales 0.6-1 mm long, ca. % of the corolla tube, bridged at 0.1-0.2 mm, oblong to spath-

ulate, short-fringed, fimbriae 0.05-0.15 mm long. Styles 1.2-2.1 mm long, longer than the ovary, evenly fili-

form. Capsules circumscissile, 1.5-2 x 0.9-1.7 mm, globose, to globose-depressed, slightly thickened and

risen, or with a few protuberances around the inconspicuous interstylar aperture, translucent, capped by the

withered corolla (Fig. 3e). Seeds 2-4 per capsule, 0.75-0.9 x 0.7-0.8 mm, angled, subrotund to broadly elliptic;

seed coat cells alveolate/papillate.

Sonora. — Common in the Sonoran Desert, especially on sandy flats, valley floors, and bajadas, and in

coastal and foothills thomscrub; near sea level-300 m. Recorded on herbaceous hosts including Amaranthus

watsonii, Boerhavia (including B. coccinea, B. triquetra). Euphorbia subgenus Chamaesyce (especially E. poly-

carpa), Pectis (including P. coulteri , P. papposa), and occasionally on Tumamoca macdougalii.

General distribution. — Baja California (norte) and Sur, Sinaloa.

Selected reference. — Costea and Stefanovic (2010).

Mpio Caborca: 35.2 km W of Caborca on road to Desemboque, desertscrub on sandy flats with Lama and Olneya, 30°44'35"N, 112°26'32’W,

63 m, 16 Jan 2002, Van Devender 2002-23 (WLU). Mpio Guaymas: 6 mi NW of Guaymas, 28 Feb 1933, Shreve 6134; Old road to Algodone.

San Carlos, 27°57'42"N, 111«03’43"W, ca. 35 m, 26 Dec 2000, Reina-G. 2000-917 (CAS, US, WLU) Mpio Hermosillo: 5 mi by road E of Bahia

Kino, 19 Oct 1963, Felger 9046. Mpio Huatabampo: 6.25 km E of Camahuiroa, 1.9 km W of Tierra y Libertad, 26°33'00"N, 109°12’45’W, 23

tn 20 |.,n f 2H-*.s Mpio Pitiquilo: at coast on N side of headland ca 10 ,h „f PcsomhocpK |sm hyuM-l 22 Man

Spellenberg 4943 (NMC). Mpio Soyopa: Arroyo Los Garambullos, 1.5 km E of T6nichi, 28°34'10"N, 109°33'00"W, 180 m, 15 Sep 1998> V*

Devender 98-1 120 (ARIZ, WLU); 4 Sepl996, Von Devender 96-360 (ARIZ, NMC, WLU) [SEM]. SONORAN ISLANDS. DAtik NW side of

land, 20 Dec 1966, Felger 15313A. Tiburon: SE side of Agua Dulce Valley, ca. 12 mi S from Tecomate, 28°57'20"N, 112°24.5 , W, ca. 280®^

26 Sep 2008, Felger 08-120. poy rpa, qu

Cuscuta erosa Yunck., Illinois Biol. Monogr. 6(2-3):26. 1921. Type: MEXICO. Sonora: 1869, Palmer s.n. (holotype: US!)

Cap-

543

ment bases, bridged at 0.5-0.7 mm, obovate to spathulate, rounded, uniformly dense, long fringed, fimbriae

0.2-0.4 mm long. Styles 1-2.5 mm long, ca. as long as the ovary, evenly filiform. Capsules indehiscent, 1.9-4(-

5) x 2-3.5 mm, globose to subglobose, thickened and risen around the medium large interstylar aperture,

semi-translucent, surrounded or capped by the withered corolla. Seeds 2-4 per capsule, 1.42-1.86 x 1.25-1.6

mm, shape heterogeneous on the same plant: dorsoventrally compressed to weakly angled, broadly elliptic to

transversely oblique, seed coat cells variable: alveolate/papillate, polygonal (not alveolate/papillose), and puz-

zle-like arranged, or both kinds present on the same seed. In * 30.

Sonora.— Sonoran Desert and coastal thornscrub; ca. 10-100 m. Flowering August-November(-March).

Apparently spreading as a weed of alfalfa (with contaminated seeds); other hosts include Baccharis, Chenopo-

dium, Helianthus, Heterotheca, Ipomoea, Pluchea, Polygonum, Rhynchosia, and Tephrosia.

General distribution. — throughout most of the USA; Aguascalientes, Chihuahua, Coahuila, Edo. Mexico,

Jalisco, Michoacan, Nuevo Leon, Puebla, Queretaro, Sinaloa, Tabasco, Tamaulipas, Veracruz, Zacatecas, Yuca-

tan; West Indies; Central America; South America. As in the case of C. campestris, C. indecora may be more

widely distributed in Mexico than current herbarium data suggest.

Selected reference. — Costea et al. (2006b).

ir Garcia 9 Aug 2010 Cm-

a 2010-01574,” WLU). Mpio Hus

Van Devender 92-1 101. and 1 '> M.ir log \ p, u „ J, , ■. t-,522 ■ \RIZ U.UR ARI7 l ( >.2 km Wot lurr.i v l.ik-i Lid mi \ ro.ul (<’ c .tm.i-

hmroa, 26°33'50”N > 109°12'50"W. 24 Nov 1093, Van Devender 93-1283 (ARIZ, ASU, CAS, TEX, UC.UCR, USON). Mpio Etchojoa: Etchojoa,

Febrero, weed in alfalfa, Ley 21 Aug 2010 (“muestra 2010-01980,” WLU). Mpio San Ingnacio: Ejido San Ignacio Rio Muerto, weed in alfalfa,

Sdanr-Garda 9 Aug 2010 (“muestra 2010-01573,” WLU).

Cuscuta legitima Costea & Stefanov., Taxon 59:1795. 2010. Type: MEXICO. Sonora: NW side of Rio Yaqui at MEX 15 near

r. reflexa (J M. Coult.)

, 1889, Nealley 338 (holotype: US!; iso

Stans slender, yellow-orange. Inflorescences dense to loose, umbelliform, confluent. Pedicels 2-10 mm long,

tacts 1 at the base of clusters and 0-1 at the base of pedicels, 2.0-3.6 mm long, broadly triangular-ovate, mar-

gins entire, apex acuminate. Flowers 5-merous (Fig. 3f), 4.0-5.5(-6.0) mm long, membranous, white when

fresh, creamy-white when dried. Papillae absent. Laticifers evident in the bracts, calyx, corolla, tips of infras-

taminal scale fimbriae, and ovary, isolated, ovoid. Calyx 2.5-3.2 mm long, straw-yellow, finely reticulate,

slightly shiny, campanulate, longer than corolla tube, divided ca. % the length, tube 0.6-1.0 mm long, lobes

l5 ~ 22 mm long, not basally overlapping, ovate-lanceolate, not carinate, margins entire, apex acuminate.

Corolla 3.8-5.2(-5.6) mm long, tube 1.6-2.1 mm long, campanulate, lobes 1.8-3.0 mm long, initially erect,

fcr reflexed, longer than the tube, linear-lanceolate, margins entire, apex acuminate, straight. Stamens ex-

j^ ned ’ starter than the lobes, anthers 0.50-0.70 x 0.24-0.36 mm, elliptic to oblong, filaments 0.6-1.0 mm

0ng hrfrastaminal scales 1.8-2.2 mm long, equaling or slightly longer than the tube, bridged at 0.2-0.4 mm,

^thulate to obovate, uniformly dense-fringed, fimbriae 0.2-0.5 mm long. Styles 0.9-2.5 mm, longer than the

° Var y- eve nly filiform. Capsules circumscissile, 2-3 x 1-2 mm, depressed, irregularly thickened and slightly

'lj Cn arou nd the inconspicuous interstylar aperture, translucent, surrounded or capped by the withered cor-

Seeds 2-4 per capsule, 0.9-1.2 x 0.8-0.9 mm, broadly elliptic to subround.

Sonora. Chihuahuan and Sonoran Deserts, coastal and foothills thornscrub, tropical deciduous forest;

200 m - lowering August-November. Common but not weedy; hosts herbaceous, including Allionia in-

, Boerhavia, Chamaesaracha, Evolvulus, Kallstroemia, Salsola, Solanum, Tidestromia la-

„.n, and Tribulus terrestris.

General distribution. Arizona Kansas New Mexico, Texas; Baja California (norte). Chihuahua, Coa-

tala, Tamaulinas

544

Selected reference . — Costea and Stefanovic (2010).

Mpio Agna Prieta: S edge of Agua Prieta on Mex 17, Chihuahuan desertscrub, 31°18’21"N, 109°34'55"W, 1204 m, 13 Sep 2006, Van Devender

h, ii, m. • II 'AH Mpio Manios: t .> pi i ,i hu.t -..i 2oBT W N I0S V. W In) m i=> Vp Van Dcvntdei ••’.-J/.MiARIZ

UC, UCR). Mpio Cajeme: Ciudad Obregdn, 29 Sep 1933, Gentry 272 (ARIZ, MICH); Cerro La Antena, 1 km N of Microondas La Cabana,

27°27'45"N, 109°46'20"W, 200 m, Sinaloan thomscrub, 19 Sep 1994, Van Devender 94-603 (ARIZ, ASU, MEXU, UC, USON); NWsideof Rio

Yaqui at MEX 15 near Esperanza, ca. 9 km N of Ciudad Obregdn, 27°35'45"N, 109°56’W, ca. 40 m, 10 Sep 1994, Van Devender 94-458 (ARIZ,

ASU, MEXU, UC, UCR) [SEM], Mpio Hermosillo: New Year’s Mine, 20 mi S of Hermosillo, Jones 28 Oct 1926 (MO); 27 mi W of Hermosillo,

on road to Kino Bay, 28 Aug 1941, Wiggins & Rollins 1 33 (ARIZ, CAS, DS). Mpio Navojoa: Sanjose de Masiaca; 26°45'N. 109 B 50"W 70 m

22 Sep 1994, Van Devender 94-710 , 94-711 (ARIZ, MEXU, UC). Mpio Puerto Penasco: Pinacate Region, MacDougal Crater, 8 Sep 1964, Fel-

Pinacate Region, 155 m, 8 Dec 1970, Felger 20035 - 1 km SWW Papago Tanks, 28 Sep 1964, Felger 10608 ; Rancho Grijalva (Rancho Guadalupe

Victoria), 32°00'35"N, 113°34'25"W, 225 m, Ezcurra9Nov 1982. Mpio Soyopa: Tdnichi, 28°35'55"N, 109°33'50"W, 200 m, 17 Aug 2006, Van

Devender 2006-627 (ARIZ, NMC, WLU); Arroyo Las Tinajas below ruins of Toledo smelter, near Loma Maderista, 3.5 km S of TOnichi, W

(MEXU, WLU), Reina-G. 2006-606 (WLU), Reina-G. 2006-612 (WLU).

a leptantha Engelm., Tra

:r544 (holotype: NY!; ts

Stems slender, yellow-orange. Inflorescences loose, umbellate, confluent. Pedicels (l-)2-7 mm long. Bracts 1

at the base of clusters and 0-1 at the base of pedicels, 0.75-1 mm long, triangular ovate, margins entire, apex

acute. Flowers 4-merous (Fig. 3h), 3.5-4.5(-5) mm long, membranous, white when fresh, creamy-white when

dried, papillae usually present on the pedicels and perianth. Laticifers not visible. Calyx 1.5-1.8 mm long,

straw-yellow, not reticulate or shiny, campanulate, V1-V2 of the corolla tube, divided ca. Vi the length, the tube

0.5-0.8 mm long, lobes 0.8-1 mm long, not basally overlapping, triangular-ovate, not carinate, margins entire,

apex acute. Corolla 3-4 mm long, tube cylindric, 1.5-2.5 mm long, lobes 1.5-2 mm long, initially erect, later

spreading or reflexed, as long as the tube, lanceolate, margins entire often involute upon drying and corolla

lobes appearing narrow, apex acute ± cucullate. Stamens short-exserted, shorter than corolla lobes, anthers

0.4-0.6 x 0.35-0.45 mm, subround to broadly elliptic, filaments 0.3-0.6 mm long. Infrastaminal scales 1.3-2J

mm long, ca. Vi of the corolla tube, bridged at 0.4-0.8 mm, oblong, uniformly short-fringed, fimbriae 0.05-0.15

mm long. Styles 1.2-2. 1 mm long, longer than the ovary, evenly filiform. Capsules circumscissile, 1-5-2 *

1.6-1.9 mm, globose, slightly thicken and risen or with a few protuberances around the inconspicuous mter-

stylar aperture, translucent, capped by the withered corolla. Seeds 2-4 per capsule, 0.75-0.9 x 0.7-0.8 mm,

angled, subrotund to broadly elliptic, seed coat cells alveolate/papillate.

Sonora. — Sonoran Desert; 5-150 m. Flowering December-May. Parasitic on Euphorbia subgenus Charnae-

syce, especially E. polycarpa.

General distribution. — Texas, New Mexico; Baja California (norte) and Sur, and Sinaloa.

Selected reference. — Costea and Stefanovtf (2010).

Mpio Hermosillo: 1 .5 mi E of Santa Rosa, 15 Feb 1965, Felger 12575; 4 mi by road NW of Rancho Noche Buena at ca. 0.5 mi E of crest of “Sm

Pass,” Sierra Seri, 14 May 1966, Felger 14035 ; Roadside 3.7 mi S of Punta Chueca, 13 Apr 1980, Bowers 1966; Playa Esthela, just N of Bahia

Kino, 28°52’28"N, 112°0r20"W, 50 m, 31 Dec 2000, Von Devender 2000-933 (WLU) [SEM] . Mpio Pitiquito: 5.9 mi S of Desemboque RioS»

Ignacio, 14 Apr 1968, Felger 17762; 19.8 mi S of Desemboque Rio San Ignacio, 14 May 1966, Felger 14080; ca. 1 mi E of 19 mi by roadSd

Desemboque, vie. 29°20'N, 11214^, 18 Feb 1968, Felger 17205. SONORAN ISLANDS. Tiburon: SW Central Valley, Felger 17342 ; 1

inland at Zozni Cmiipla, at base and N side of Punta San Miguel, 23 Nov 2006, Wilder 06-368; Canyon at base of CapxOlim, 24 Nov

Wilder 06-381.

Cuscuta macrocephala W. Schaffn. ex Yunck., Illinois Biol. Monogr. 6(2-3):36. 1921. Type: MEXICO. Sinau* !®>

date], Schaffners.n. (uoi.otypi : NY!).

Stems orange, coarse. Inflorescences dense, paniculiform-glomerulate. Pedicels 0.3-3.2 mm long. Bracts 1 i[

the base of clusters and 0-1 at the base of pedicels, 1.5-3 mm long, ovate, margins entire, apex obtuse. Flo** 1 *

5-merous (Fig. lb), 5-6.5 mm 1

ca. % of its length, tube 1.3-2

1 per capsule, 1.4-1.9 x 1-1.3 mm, a

(osteaetal..

547

x 1.25-1.43 mm, ± visible through the pericarp, dorsoventraily compressed, broadly elliptic to subround; sur-

face of seed coat epidermis alveolate when dried and papillate when hydrated.

Sonora. — The species is apparently localized at Sonoyta (400 m) and Quitobaquito (at the Arizona-

Sonora border, 335 m). Flowering April-May. Growing on Suaeda moquinii.

General distribution. — Arizona, California, Nevada, New Mexico, Utah, Texas; Baja California (norte) on

herbaceoushosts (e.g. , species ofFrankenia, Salsola, Suaeda, Wislizenia) from inland salt flats, marshes, and ponds.

Selected references. — Costea et al. (2006c, 2009), Felger (2000).

I Type: MEXICO. Oaxaca; 1827,

Pedicels 0.5-2.6 mm long. Bracts 1 at the base of clusters, usually absent at the base of pedicels or flowers,

1.5-3 mm long, oblong to oblong lanceolate, acute to obtuse, margins entire. Flowers 5-merous, 4-5.2 mm

long, thick, white when fresh, reddish-brownish when dried. Papillae absent. Laticifers visible in the calyx,

corolla, isolated or in rows, ovoid to elongated. Calyx 2-3 mm long, reddish-brownish, more or less reticulate,

± glossy, campanulate, equaling corolla tube, divided VS-*A the length, tube 0.5-1 mm long, lobes 1.6-2.2 mm

long, broadly overlapping, round to broader than long or occasionally broadly elliptic, not carinate or with

multicellular protuberances on the midveins, margins entire, apex rounded. Corolla 3.5-5 mm long, tube

2.3-3 mm long, campanulate, lobes 1.5-2.5 mm long, initially erect, later reflexed, equaling or shorter than the

lube, oblong-ovate, overlapping, margin entire, apex rounded, straight. Stamens exerted, shorter than corolla

lobes, anthers 0.7— 1.1 mm long, oblong-elliptic, filaments 0.8-1.2 mm long. Infrastaminal scales 2.5—3 mm

long, equaling corolla tube, bridged at 0.8-1.2 mm, oblong to ovate, uniformly dense-fringed, fimbriae 0.2-0.5

mm long. Styles 1.2-2.1(-3) mm long, longer than the ovary, thick, but uniform. Capsules circumscissile,

1-5-3 x 1.8-2.5 mm, globose to depressed-globose, not thickened and/or risen around the small interstylar

aperture, translucent, capped by the withered corolla. Seeds (2-)4 per capsule, 1.5-2 x 12-1.9 mm, angled or

slightly dorsoventraily compressed, elliptic-oblong to subround, seed coat cells alveolate/papillate or wrin-

Sonora.— This species may be a sporadic introduction. It was collected in northern Sonora in 1994 from a

cultivated Schinus terebinthifolia tree. On many subsequent visits, however, it was not found again. Ornamental

trees and shrubs grown in Sonora are often brought from nurseries in Guadalajara, Jalisco, which points to a

Potential source for new introductions. Sonoran Desert. Flowering December-January.

General distribution.— Juncker (1932, 1965) mentioned that C. tinctoria is common throughout Mexico to

Guatemala. This species is part of the largest and most complicated taxonomically clade in Mexico (clade “G”;

Stefanovic et al. 2007) and the delimitation of species and their distribution require more study.

Selected reference.— Yuncker (1932).

10 °8(ariz, mexu, uson, WLU) ISEM], 5153 gd

Guscuta tuberculata Brandegee, Univ. Calif. Publ. Bot 3 38Q 1909 Type MEXICO. Lower California [Baja California

S ‘ ems fiIif °rm, yellow-orange. Inflorescences loose, umbelliform or racemiform, confluent. Pedicels 2-3(-5)

!" m lon & Bracts 1 at the base of clusters, usually absent at the base of peduncles, 0.5-0.75 mm long, ovate-

nce °late, margins entire, apex acute. Flowers 5-merous (Fig. 3i), 2.5-4 mm long, membranous, white-

when fresh, creamy when dried. Papillae present especially at the base of the corolla tube. Laticifers

^visible in the corolla, isolated, ovoid to elongated. Calyx 0.5-1.5 mm long, yellow, not or finely reticulate,

^Pulate-angular, Vi-Vi as long as the corolla tube, divided almost to the base, tube 0.2-0.5 mm long,

fr-1-3 mm long, not basally overlapping, triangular to lanceolate, carinate and/or with multicellular pro-

Costea et al., Convolvulaceae of Sonora, Mexico: Cuscuta

General distribution. — Arizona, Colorado, Ne'

Federal, Guanajuato, Guerrero, Edo. Mexico, Hidalgo, Jalis

rftaro, San Luis Potosi, Tamaulipas, Veracruz; West Ind

Selected reference . — Costea and Stefanovic (2010).

i, Durango, Distrito

on, Oaxaca, Puebla, Que-

and South America.

Cuscuta vandevenderi Costea & St<

:r. Pedicels 0.7-6 ir

*s corymbiform cymes arranged in dense globose inflorescences, 1-3 cm in diame-

lg. Bracts 1 at the base of clusters, 0.5-0.75 mm long, triangular-ovate, margins entire

. Flowers (4-) 5-merous, 2-2.6 mm long, membranous, white turning cream yellow-

ish when fresh, creamy-light brown when dried. Papillae absent. Laticifers prominent in the calyx and corolla

articulated or isolated, rectangular, ovoid to elongated. Calyx 0.9-1.6 mm long, yellow-gray, not shiny, cam

panulate, divided y h- l h mm, tube 0.4-0.8 mm long, lobes 0.5-1.2 mm long, not overlapping to overlapping

triangular to triangular-lanceolate, carinate, margins ± entire to serrulate, apex acute, acuminate to obtuse

Corolla 1.5— 2.1 mm long, tube campanulate, 0.9-1.3 mm long, lobes 0.6-1.3 mm long, erect to slightly spread

ing, triangular, margins entire to irregular, apex obtuse to acute. Stamens equaling to longer than corolla lobes

anthers 0.3-0.5 x 0.3-0.4 mm, subround to broadly elliptic, filaments 0.6-1.5 mm long. Infrastaminal scales

1-1.2 mm long, equaling corolla tube, bridged at 0.4-0.5 mm, oblong-ovate to truncate, densely fringed, fim-

briae 01-0.2 mm long. Styles 0.8-1.2 mm long, longer than the ovary, uniformly filiform. Capsules indehis-

cent , 1-8-2.4 x 0.8— 1.2 mm globose— depressed to globose— obovoid, slightly thickened but not risen around

the relatively large interstylar aperture; persistent corolla surrounding the base of capsules. Seeds 3-4 per

tapsule, 0.9-1.1 x 1-1.2 mm, subrotund, seed coat cells alveolate/papillate.

Cuscuta vandevenderi resembles C. gracillima from which it differs through the smaller flowers and inde-

hiscent capsules surrounded by persistent corollas, and the often serrate calyx and corolla lobes. From C. del-

tottko, which is the closest related species, it can be separated by the indehiscent capsules and denser inflores-

cences (Costea et al. 2008).

Sonora.— Southeast and central part of the state in tropical deciduous forest, oak woodland, and pine-oak

forests (sometimes in openings); 350-1550 m. Flowering September-December. Parasitizing various herbs,

e &,Ayenia, Chamaecrista, Cosmos, Euphorbia, Sida, and Evolvulus.

General distribution .— Sonora and Baja California Sur.

Selected reference .— Costea et al. (2008).

^ Ahmos: Sierra Tecurahui, 1200-1500 m, 26-28 Oct 1961, Gentry 19423 (US); 3.9 km above Rancho El Palmarito, 23.9 km, E-NE of

AUmos ’ 27 °03'04"N, 108°45'51"W, 516 m, 1 Oct 2006, Van Devender 2006-983 (WLU); El Guayabo Crossing of Rio Cuchujaqui, 14 Km (by

^E-SEof Alamos, 27°00’05"N, 108°47'08"'W, 370 m, 2

3, Steinmann 93-349 (ASU). Mpio Mazatan: Sierra de Mazatfn,

E-SEofYecora, 2

^ see Felger et al. (2012) for full acknowledgments.The following herbaria have provided Cuscuta plant

^ nal: AA U ALTA, AR1Z, ASU B BAB BOL BRIT, CANB, CAS, CEN, CHR, CHSC, CIIDIR, CICY, CIMI,

rf- DA 0. F, G, GH, H, HUFU, IAC, IEb] IND, j, JEPS, LL, LP, LPB, LPS, K, MEL, MERL, MEXU, MICH, MO,

C ’ ***• ° A Q OKLA, OSC, OXF, PACA, PRE, QCNE, QFA, P, PACA, RB, RSA, SAM, S, SD, SGO, SI, SPF,

550

TEX, TRT, TRTE, UA, UB, UBC, UCR, UCT, UNB, UNM, UPRRP, UPS, US, USAS, WTU and XAL. Cuscuta

research was supported by a Natural Sciences and Engineering Research Council of Canada Discovery grant

to Costea (327013-06 and 327013-12). We thank J. Andrew McDonald and Javier Ortega for providing helpful

Austin, D.F. 1982. 165.Convolvulaceae. In: G.W. Harling and B.B. Sparre, eds. Flora of Ecuador, vol. 15. University of Gote-

borgand Swedish Museum of Natural history, Goteborg and Stockholm. Pp. 1-98.

Costea, M. 2007-onwards. Digital atlas of Cuscuta (Convolvulaceae). Wilfrid Laurier University, Ontario, Canada, https://

www.wlu.ca/page.php?grp_id=2147&p=8968&pv=1 (viewed 16 June 2011).

Costea, M. and FJ. Tardif. 2004. Cuscuta (Convolvulaceae) — the strength of weakness: a history of its name, uses and

parasitism concept during ancient and medieval times. Sida 21:369-378.

Costea, M. and FJ. Tardif. 2006. The biology of Canadian weeds. Cuscuta campestris, C. gronovii, C. umbrosa, C. epithymum

and C. epilinum. Canad. J. PI. Sci. 86:293-31 6.

Costea, M., and S. StefanoviC. 2009. Cuscuta jepsonii (Convolvulaceae), an invasive weed or an extinct endemic? Amer. J.

Costea, M., and S. StefanoviC. 2010. Evolul

evidence of extensive hybridization from discordant nuclear and plastid phylogenies. Taxon 59:1 783-1800.

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TREPTOSTEMON (LAURACEAE), A NEW GENUS OF FOSSIL FLOWER

FROM MID-TERTIARY DOMINICAN AMBER

Kenton L. Chambers George 0. Poinar, Jr. Andre S. Chanderbali

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 9733 1, U.S.A.

chamberk@science.oregonstate.edu

Department of Zoology

Oregon State University

Corvallis, Oregon 97331, U.SJ

Department of Biology

University of Florida

Gainesville, Florida 3261 1, U.S.A

ABSTRACT

be assigned to any modem gem

RESUMEN

INTRODUCTION

Amber deposits of the Cordillera Septentrional, Dominican Republic, continue to yield interesting fossil angio-

sperm flowers from the low-elevation tropical forests characteristic of Mid-Tertiary Caribbean vegetation

(Poinar & Poinar 1999). We have recently described fossils assignable to Licania (Chrysobalanaceae) (Poinar

etal. 2008a, revised by Chambers & Poinar 2010), Persea (Lauraceae) (Chambers et al. 2011a), Thchilia

(Meliaceae) (Chambers et al. 2011b), Swietenia (Meliaceae) (Chambers & Poinar 2012), and Trochanthera (pos-

sibly Moraceae) (Poinar et al. 2008b). A second flower of Lauraceae is described in the present paper. It has

several well-marked features, including 6 fertile stamens in whorls 1 and 2, whose anthers have 2 pairs of pores

tlwt °P en extrorsely. The stamens of whorl 3 display 2 upright valves but are possibly staminodial, since their

terminal portion is dissimilar in size and shape from the fertile anthers. The anthers are closely adjacent and

are on short filaments. The epidermis of the anthers and staminodes is densely covered with minute trichomes.

No PisfiHode or whorl 4 staminodes are visible. Because of the extrorse dehiscence of the 6 anthers of whorls 1

3nd 2 ’ the fossil cannot easily be accommodated in any modem genus of Lauraceae. We therefore propose to

Separate 11 as the new genus and species Treptostemon domingensis.

MATERIALS AND METHODS

AH the amber fossils referred to above came from mines in the Cordillera Septentrional, between Puerto Plata

®nd Santiago, Dominican Republic. Dating methods applied to the deposits are thus far equivocal. An age of

30 ®ybp was assigned by Cepek in Schlee (1999) based on fossil coccoliths, and one of 20-15 mybp was

ermined by Iturralde-Vinent and MacPhee (1996) based on foraminifera. The amber is found in turbiditic

552

sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994). Animals and plants of

the forest vegetation present at the time of amber deposition were described by Poinar and Poinar (1999), the

original resin having been a product of the arborescent genus Hymenaea (Fabaceae). In Poinar and Poinar (op.

ciL), the present flower was inadvertently assigned to the genus Nectandra (Pg. 21, Fig. 10).

DESCRIPTION

Treptostemon K.L. Chambers, Poinar, & A.S. Chanderbali, gen. nov. Type Species: Treptostemon domingensis K.L. Chan-

Diagnosis.— Flower staminate, radially symmetrical (Fig. 1), receptacle flat, pedicel remnant strigose, tepals6,

in 2 whorls of 3, separate, approximately equal, spreading, lightly strigose on both surfaces, margins involute,

glabrous (Fig. 2), stamens 9, in whorls of 3, at least the outer 2 whorls fertile, anthers minutely puberulent,

those of whorls 1 and 2 with 4 pores arranged as 2 pairs, one above the other, all extrorse, opening by apical

valves, stamens of whorl 3 functional or staminodial, their distal portion dissimilar in size and shape from the

fertile anthers (Fig. 2), 2 or 4 small pores probably present, the distal pair opening extrorsely by upturned

valves, glands of whorl 3 stamens small, whorl 4 staminodes and pistillode not evident. Pistillate flower un-

Etymology. — From Greek “treptos,” turned, and “stemon,” stamen.

Description.— Tepals lanceolate, acute, laterally spreading, 3.1 to 4.0 mm long, 1.1 to 1.6 mm wide (Fig. 1), an-

thers of whorls 1 and 2 0.9 to 1.0 mm wide, ca. 1,0 mm long, with short filaments, pores extrorse, circular to

oblong, ca. 0.3 mm in diameter, valve remnants visible in a few cases (Fig. 2), connective forming an adaxial

ridge, stamens of whorl 3 club-shaped, distal portion cylindrical, 0.7 mm in diameter, pores extrorse, the distal

pair defined by small, upturned valves (Fig. 2), otherwise not observable, glands of whorl 3 stamens 0.19 mm

wide (only 1 observed).

Etymology. — From source of amber in Domin

DISCUSSION

Examination of the fossil is best done from above, as in Figures 1 and 2. In a lateral view, obtained with diffi-

culty, the outer anthers display all 4 extrorse pores, although in apical view, the lower pair may be barely visible

(Fig. 2). The filaments of the outer stamens are quite short and curve at the tip, so that the anthers face dorso-

laterally. The positioning of the stamens does not allow observation of pores on whorl 3, and their number is

uncertain. The upraised valves of the distal pores (Fig. 2) are in an extrorse position. Despite the reduced size

and cylindrical shape of these anthers, it is uncertain whether the whorl 3 stamens are fertile or staminodial. A

staminodial condition is possible, considering the report by Kubitzki and Kurz (1984) that in staminate flow-

ers of 3 dioecious species of Ocotea they studied, “[tjhe staminodes often possess valves that open after the

wilting of the stigma” (i.e., stigma of the pistillode). Only one gland of the whorl 3 stamens is in view (Fig. 2)

On some anthers, small, black air bubbles that have settled among the surface trichomes may be confused for

pores, but these are artifacts. The extrorse dehiscence in all androecial whorls of Treptostemon is unique in

Lauraceae, although in Pleurothyrium there are 9 tetrasporangiate, latrorse anthers which may look almost

extrorse (J. Rohwer, pers. comm.).

The shortness of the stamens of Treptostemon and their close positioning in the flower are features similar

to Aniba and Aiouea (Kubitzki and Renner 1982), among other genera, although a close relationship to these 2

taxa, which are hermaphrodite, with upright tepals and only 2 pores per anther, is not likely. Furthermore, the

usual laurad introrse dehiscence of whorls 1 and 2 anthers occurs in the 2 genera. Stamens with short filaments

also occur in the New World dioecious genera Rhodostemonodaphne (MadriMn 2004) and Ocotea (the

Chambers et al., Treptostemon, a new f

553

v of complete flower. Scale bar = 1 .4 mm.

with bisexual or unisexual flowers). Both have 4-pored, introrse anthers, but in Khodostemonodaphne the pores

form a more or less horizontal row (Rohwer 1993; van der Werff 1991 , Fig. 2E), unlike the pores of Treptostemon

anthers. The latter two genera are similar in lacking staminodes of whorl 4 but differ in their receptacle, which

15 narrow ly tubular in Rhodostemonodaphne (Rohwer op. cit.). The closest relative among genera with 4-pored

*«heis is probably Ocotea, a species-rich and variable taxon characterized by Rohwer (1993, p. 382) as “the

dustbin of the Perseae.” Dioecious species such as O. pyramidata (Allen 1945) may have anthers 1.0 mm long,

Wuh the filament only 0.5 mm. (As an aside, although pollen is shed inwards by these anthers, it is also released

outwards from the 2.1 5 mm-long stamens of whorl 3). According to Rohwer (op. cit.) the whorl 4 staminodes

of Ocot ea are absent in unisexual flowers, the receptacle varies from flat to deeply tubular, and the pistillode in

flower s may be present or absent. The principal difference between Ocotea and Treptostemon, therefore, is

1 at the former has anthers of whorls 1 and 2 dehiscing introrsely.

Pollen had recently been discharged from the Treptostemon flower before it became imme

555

in, as shown by a cloud of tiny pollen grains belc

would have had to release pollen from the stami

visiting insects for transfer to pistillate flowers on another tree. It is likely that dioecy, nectar ]

extrorse anther dehiscence were adaptive features of the species’ pollination syndrome, but w

moved in time to reconstruct other details of its floral biology. A brief review of reports on poll

present-day Lauraceae may be of interest, nonetheless, in conjunction with a discussion of avail

in the insect fauna already known from Dominican amber (Michener & Poinar 1997; Poinar & Poinar 1999).

A report on pollinators identified from modern tropical rain forests in Costa Rice (Bawa et al. 1985) lists

only one member of Lauraceae, an unidentified species of Ocotea. It is cited as hermaphroditic and a member

of the subcanopy, its major pollinators being beetles. Whether the flowers offer both nectar and pollen as food

rewards is not mentioned. However, in her generic description of Ocotea, Allen (1945) states that stamens of

the inner series always bear two sessile or stipitate glands, presupposing the presence of a nectar reward for

visitors. In a study of dichogamy and dioecy in Neotropical Lauraceae, Kubitzki and Kurz (1984) included ob-

servations of pollinators for 3 dioecious species of Ocotea. Two species were seen to be visited by brown bees,

about 1 cm in length, while small diptera, wasps and moths sucking nectar were occasionally encountered.

Larger flies were observed visiting flowers of O. guianensis and O. opifera but could not be caught for identifica-

tion. A more detailed study of pollination in a single dioecious species, Laurus azorica of Macaronesia, was

published by Forfang and Olesen (1998). In a sample of over 200 trees, flower visitors included at least 11 dif-

ferent insect species. These were: Hymenoptera-2 species of Halictidae, 1 of Apidae, and 1 of Ichneumonidae;

Diptera-1 species of Musicidae, 1 of Syrphidae, 1 of Tachinidae, and 1 of Bibionidae; Lepidoptera-1 species of

Nymphalidae; Coleoptera-1 species of Nitidulidae; Hemiptera-1 species of Pentatomidae. Only Halictidae and

Tachinidae were common, comprising 97% of the visits. Bees are also considered to be the principle pollinators

of the avocado, Persea americana, even though flies and other insects also visit the flowers (Free 1993).

Generalized, radially symmetrical flowers of Lauraceae, with spreading perianth and exposed stamens

and pistil, are open to visits by many different groups of anthophilous insects. A potential pollinator might

develop a specific attraction to the odor of pollen or nectar in such a flower and become faithful to one or a few

species (Faegri & van der Pijl 1979). However, it is unlikely that an unspecialized flower like that of Treptoste-

mon - even with its modified form of pollen presentation atypical of the family, would have been closely adapted

to a single kind of pollinator. Since their origination in the Early Cretaceous (Danforth & Poinar 2011), bees,

which rear their young on pollen, have been among the most dependable pollinators of angiosperms. Based on

the above reports indicating that bees are frequent visitors of present-day Lauraceae, it is likely that this group

of insects also visited and pollinated T. domingensis in the Tertiary forests of Hispaniola. The bee fauna of Do-

toinican amber includes representatives of families Apidae, Andrenidae, Colletidae, and Halictidae (Michener

& Poinar 1997). Of these, the most common bee in Dominican amber is the small, stingless Proplebia domini-

““ (Apidae, tribe Meliponini), which could well have been significant in the pollination system of Treptoste-

mon - D espite their presence in the fossil record, stingless bees do not occur in Hispaniola today. Their disap-

pearance as keystone species could have led to the elimination of many plant species, including Treptostemon.

ACKNOWLEDGMENTS

We thank Jens G. Rohwer and Santiago Madrinan for their helpful review comments.

REFERENCES

^ CA 1945. Studies of the Lauraceae, VI. Preliminary survey of the Mexican and Central American species. J. Arnold

Arbor. 26:280-434.

Bawa ' K - s - s -H. Bullock, D.R. Perry, R.E. Coville, and M.H. Grayum. 1 985. Reproductive biology of tropical lowland rain forest

bees- "• Pollination systems. Amer. J. Bot. 72:346-356.

T*®*’ K.L and G.O. Poinar, Jr. 201 0. The Dominican amber fossil Lasiambix (Fabaceae: Caesalpimoideae?) is a Ucama

hrysobalanaceae). J. Bot. Res. Inst. Texas 4:21 7-21 8.

556

K.L., G.O. Poinar, Jr., and A.E. Brown. 201 la. A fossil flower oiPersea (Laura

tes. Inst. Texas 5:457-462.

», K.L, G.O. Poinar, Jr., and A.E. Brown. 201 1 b. Two fossil flowei

nst. Texas 5:463-468.

», K.L. AND G.O. Poinar, Jr. 2012. A Mid-Tertiary fossil flower of Swietenia i

Danforth, B.N. and G.O. Poinar, Jr. 201 1 . Morphology, classification, and antiquity of Mellittosphex burmensis (Apoidea:

Melittosphecidae) and implications for early bee evolution. J. Paleontol. 85:882-891 .

and J.F. Lewis. 1 994. Hispaniola. In: S. Donovan and T.A. Jackson, eds. Caribbean geology: an introduc-

sity of the West Indies Publishers' Association, Kingston, Jamaica. Pp. 129-150.

der Pul. 1979. The principles of pollination ecology. Ed. 3. Pergamon Press, Oxford.

Forfang, A-S. and J.M. Olesen. 1 998. Male-biased sex ratio and promiscuous pollination in the dioecious island tree Laurns

azorica (Lauraceae). PI. Syst. Evol. 21 2:1 43-1 57.

Free, J.B. 1993. Insect pollination of crops. Ed. 2. Academic Press, New York.

1850-1852.

Kubitzki, K. and H. Kurz. 1984. Synchronized dichogamy and dioecy in Neotropical Laurai

Kubitzki, K. and S. Renner. 1982. Lauraceae I (Aniba and Aiouea). FI. Neotropica 31:1-124.

MadriNAn, S. 2004. R

’aceae). FI. Neotropica 92:1-102.

Michener, C.D. and G.O. Poinar, Jr. 1997. The known bee fauna of Dominican amber. J.

1. Syst. Evol. 147:253-266

s Entomol. Soc 69, suppL

1996:353-361.

Poinar, G.O. Jr. and R. Poinar. 1 999. The amber forest. Princeton University Press, Princeton, NJ.

Poinar, G.O. Jr., K.L. Chambers, and A.E. Brown. 2008a. Lasiambix dominicensis gen. and sp. nov., a eudicot flower in Domini-

can amber showing affinities with Fabaceae subfamily Caesalpinioideae. J. Bot. Res. Inst. Texas 2:463-471.

Poinar, G.O. Jr., K.L Chambers, and A.E. Brown. 2008b. Trochanthera lepidota gen. and sp. nov., a fossil angiosperm inflores-

cence in Dominican amber. J. Bot. Res. Inst. Texas 2:1 167-1173.

Rohwer, J.G. 1 993. Lauraceae. In: K. Kubitzki, J.G. Rohwer, and V. Bittrich, eds. The families and genera of vascular plants

II. Springer-Verlag, Berlin, Germany. Pp. 366-391 .

Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C, 28.

VAN DER Werff, H. 1 991 . A key to the genera of Lauraceae in the New World. Ann. Missouri Bot. Gard. 78:377-387.

A NEW FOSSIL SPECIES OF COLPOTHRINAX (ARECACEAE) FROM

MID-TERTIARY MEXICAN AMBER

Kenton L. Chambers George 0. Poinar, Jr.

Alex E. Brown

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

:e.oregonstate.edu

Department of Zoology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

629 Euclid Ave.

Berkeley, California 94708, U.S.A.

RESUMEN

INTRODUCTION

A new fossil species of palms, described here as Colpothrinax chiapensis, is represented by 4 flowers embedded

m Mid-Tertiary amber from a mine in the state of Chiapas, Mexico. The floral morphology of the species is

similar in general features to the modern genus Colpothrinax, whose three species are found in Cuba and Cen-

tral America, from Guatemala to Panama (Evans 2001). The pistil, in particular, is like that of this genus, with

3 separate but apposite carpels and an elongate style, which appears to be fused and grooved rather than com-

posed of 3 separate units. The style tip is tapering and lacks an enlarged stigma. The basally connate petals,

forming a short tube surrounding the ovary, and the almost completely united sepals with short, triangular

lobes, provide further similarities. Principal differences are in the androecium of C. chiapensis, in which the

longer, linear-subulate filaments are widened only near the base and slightly united into a staminal tube. The

anthers are large, dorsifixed, and latrorse in dehiscence, as described for Colpothrinax (Dransfield et al. 2008).

The probable age of the fossils, 22.5-26 Ma, is commensurate with an estimated mean stem age for the genus of

28-56 Ma (Bacon et al 2012). The species’ Mexican origin gives evidence for an evolutionary radiation of Col-

pothrinax in the Caribbean region, as suggested by Bacon et al. (op. cit.) for Copemicia and certain other genera

s . Specimen originated from an amber mine in the

unojovel area in Chiapas, Mexico. Maps of the area i

Was P rod «ced by Hymenaea mexicana (Poinar & Brc

Lorthem mountain ranges (Chiapas Highlands) of the

e given in Poinar (1992). Amber from Chiapas, which

m 2002), occurs in lignitic beds among sequences of

Journal of the Botanical Research Institute of Texas 6(2)

primarily marine calcareous sandstones and silts. The particular amber under study is associated with the

Balumtun Sandstone of Early Miocene and the La Quinta Formation of Late Oligocene, with radiometric ages

from 22.5-26 Ma (Berggren & Van Couvering 1974). The amber is secondarily deposited in these marine for-

mations and may be somewhat older than the above dates. The original vegetation was probably a wet to moist

tropical to subtropical forest.

Colpothrinax chiapensis K.L. Chambers, Poinar, & A.E. Bi

deposited in the Poinar amber collection maintained at Oregon Stat

2, catalogue number Sd-9-181B, Figs. 3-4, c

nov. (Figs. 1-4). Type: MEXICO. Chiapas: amber 1

y, Corvallis, Oregon 97331, U.S.A.; paratypes: see Fig

Flowers hermaphrodite, ca. 6 mm in diameter when petals spread, calyx cupulate, glabrous, 0.4 mm (Fig. 4),

sepals united, with 3 free, triangular, 0.4-1.1 mm lobes, petals 3, glabrous, ovate-lanceolate, ca. 3.3 mm,

spreading or reflexed, with numerous parallel veins and no raised mid-nerve adaxially (Fig. 1), shallowly

grooved abaxially (Fig. 2), tip boat-shaped (Fig. 1), stamens 6, exserted, glabrous, filaments subulate, 1.0-1.2

mm, widened at base, the connate portion short (Fig. 3), attached at the mouth of the corolla tube, anthers ca.

2.3 mm, dorsifixed, locules 4, dehiscence latrorse (Fig. 2), gynoecium glabrous, 3-carpellate, ovary loosely

enclosed in corolla tube (Fig. 1), carpels free, apposite, style appearing united, ca. 1.2 mm, 3-grooved, tip acute,

stigma not enlarged (Fig. 2).

Etymology. — from the fossils’ origin in Chiapas, Mexico.

r, the species’ pres-

DISCUSSION

The fossils differ from other genera of Trachycarpeae in such features as the enlarged anthers, persistent petals

(cf. Pritchardia), cupulate calyx (cf. Brahea, Acoelorrhaphe), long filaments (cf. Copemicia), and presence of a

staminal tube (cf. Serenod) (see Dransfield et al. 2008, pp. 272—285, for illustrations). The orientation of the

flowers does not allow measurement of the depth of the corolla tube. In other respects, the fossils can be

matched with excellent descriptions and floral illustrations of Colpothrinax in Uhl and Dransfield (1987),

Dransfield and Uhl (1998), Dransfield et al. (2008), and Evans (2001). Whether the proposed new species falls

within the full morphological variation of any of the extant taxa is uncertain, but we believe that the androe-

cium, in particular, sets it apart from Colpothrinax wrightii (Bailey 1940) and C. cookii (Read 1969), and

large petals and long, subulate, scarcely united filaments are adequately distinct from C. aphanopetala (Evans

2001). The published treatments of the 3 modern species include much information about stem and leaf mor-

phology, inflorescence, pollen (esp. Dransfield et al. 1990), habitats, and associated vegetation. Vegetativi

inflorescence morphology is, of course, unknown for C. chiapensis, as is the pollen; howevr

ervation in resin from Hymenaea mexicana is evidence of moist tropical or subtropical forest vegetation as u*

preferred habitat. .

The phylogenetic relationships of the genera of tribe Trachycarpeae (Dransfield et al. 2008 [formerly

istoneae, Dransfield et al. 2005]) have begun to be clarified by recent molecular studies utilizing nuclear

plastid DNA (Asmussen et al. 2006; Roncal et al. 2008; Baker et al. 2009; Bacon et al. 2012). In the first 3 pape*

cited, Colpothrinax is in a clade that includes several other genera with which it had previously been assoCiat ^

including Pritchardia, Washingtonia, Serenoa, Acoelorraphe, Brahea, Lmstona, and Chamaerops. However, in ^

report by Bacon et al. (2012), involving matK, ndhF, tmD-tmT and 3 nuclear loci, Colpothrinax is alone m

highly supported clade that is sister to 2 large clades containing, inter alia, Brahea, Chamaerops, Serenoa,

lorraphe, and Lmstona, and is well separated from Washingtonia and Pritchardia. The subtribal classification

Trachycarpeae is presently in flux, it appears, as perhaps is true also in other large families now under int

molecular phylogenetic study.

The new intrafamilial relationships of palms revealed by molecular research have been used to in J eSt ^

additional questions beyond taxonomy alone. These include the origin and global diversificatic

rain forests (Couvreur et al. 2011) and the effects of Miocene dispersal on island radiations ii

559

560

Journal of the Botanical Research Institute of Texas 6(2)

tailed attention by Bacon et al. (2012). Of interest here is the authors’ estimation of the age of crown and stem

nodes for the clades that they resolve within this tribe. The clade comprising Colpothrinax alone, mentioned

above, is assigned a mean stem age of 28.56 Ma (dates for upper and lower age estimates were not well sup-

ported in posterior probabilities analysis). The mean crown age for the genus is 7.87 Ma, with upper and lower

estimates of 14.44 and 2.74 Ma. The age range assigned to the Mexican amber (Berggren & Van Couvering

1974) of 26-22.5 Ma is therefore in good agreement with a stem position for C. chiapensis. It is reasonable that

crown species radiation would have occurred later, in the Central American and Caribbean area. This is one of

the regions of interest to Bacon et al. (op. cit.), which they specifically mention as characteristic of the evolution

of this genus, as well as of Rhapidophyllum, Brahea, Washingtonia, and Caribbean Copemida (their Fig. 5).

ACKNOWLEDGMENTS

Careful review comments by John Dransfield and Scott Zona are much appreciated. We thank Deborah Car-

roll, Valley Library, Oregon State University, for help with bibliographic material.

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Dransfield, J. and N.W. Uhl. 1 998. Palmae. In: K. Kubitzki, ed.The families and genera of vascular plants. IV. Springer Verlag,

Berlin. Pp. 306-389.

Dransfield, J., N.W. Uhl, C.B. Asmussen, WJ. Baker, M.M. Harley, and C.E. Lewis. 2005. A new phylogenetic classification ottne

palm family, Arecaceae. Kew Bull. 60:559-569.

Dransfield, J., N.W. Uhl, C.B. Asmussen, WJ. Baker, M.M. Harley, and C.E. Lewis. 2008. Genera palmarum— the evolution and

classification of palms. Royal Botanic Gardens, Kew, Richmond, UK.

Evans, RJ. 2001 . Monograph of Colpothrinax. Palms 45:1 77-1 95.

Poinar, G.O. Jr. 1992. Life in amber. Stanford Univ. Press, Stanford, CA.

Poinar, G.O. Jr. 2002. Fossil palm flowers in Dominican and Mexican amber. Bot. J. Linn. Soc. 1 38:57-61 .

Poinar, G.O. Jr. and A.E. Brown. 2002. Hymenaea mexicana sp. nov. (Leguminosae: Caesalpinioideae) from Mexican am

indicates Old World connections. Bot. J. Linn. Soc. 1 39:1 25-1 32.

Read, R.W. 1 969. Colpothrinax cookii — a new species from Central America. Principes 1 3:1 3-22.

Roncal, J., S. Zona, and C.E. Lewis. 2008. Molecular phylogenetic studies of Caribbean palms (Arecaceae) and their relat

ships to biogeography and conservation. Bot. Rev. 74:78-102.

ADDITIONAL FOSSILS IN DOMINICAN AMBER GIVE EVIDENCE

OF ANTHER ABORTION IN MID-TERTIARY TRICHILIA (MELIACEAE)

Kenton L. Chambers George O. Poinar, Jr.

562

Journal of the Botanical Research Institute of Texas 6(2)

MATERIALS AND METHODS

The fossils originated from mines in the Cordillera Septentrional of the Dominican Republic, between the cit-

ies of Puerto Plata and Santiago, island of Hispaniola. Two differing ages have been proposed for this amber

deposit the older being 45-30 Ma, based on fossil coccoliths (Cepek in Schlee 1999) and the younger being

20-14 Ma, based on foraminifera (Iturralde-Vinent & McPhee 1996). Most of the amber is secondarily depos-

ited in turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994). The

specimens described here were found during a search of the Poinar amber collection for additional flowers of

Meliaceae that might relate to those already described. They are designated as numbers D-9-27C, D-9-27D and

D-9-27E in this collection. A newly described species of Swietenia, which incidentally had been illustrated

earlier (Poinar & Poinar 1999), is also representative of fossil Meliaceae flowers in these amber deposits from

the Dominican Mid-Tertiary tropical forest (Chambers & Poinar 2012).

RESULTS

The first illustrated flower, designated A (Figs. 1, 2) resembles Trichilia glaesaria in the shape and pubescence

of its calyx and corolla (cf. Fig. 2 in Chambers et al. 2011). Its lobed, strigillose calyx differs from that of T. an-

tique which was described as glabrous and rotate. In apical view (Fig. 1), the ovary of flower A is well devel-

oped, 1 mm wide, strigillose, and situated at the base of the staminal tube. Four anthers can be seen, showing

different degrees of extreme abortion. Three have knob-like terminal structures, which resemble the one much

reduced anther seen in A. antiqua (op. cit.. Fig. 5). A short filament is present on one of these (upper left arrow,

Fig. 1), while the fourth remnant is a tiny filament with a mere suggestion of an anther (upper right arrow).

Anthers are absent at other potential sites on the irregularly lobed, partly obscured staminal tube. On the evi-

dence from this flower, the filament stubs of T. antiqua shown in Fig. 3 of Chambers et al. (op. cit.) might be

reinterpreted as lacking anthers from the beginning, rather than having them removed earlier by an herbivo-

In the flower designated B (Figs. 3, 4), the adaxial and abaxial sides of the petals are densely covered with

shiny droplets of a glassy deposit, which obscures the natural pubescence of these surfaces. The calyx, barely

in view, also has this deposit. In lateral view (Fig. 3), anthers with much reduced locules are visible in alterMt-

ing higher and lower notches between acute lobes on the rim of the staminal tube. The apical view of is

flower (Fig. 4) shows approximately 10 anthers, which are also covered with the glassy deposit. The stigma ol

the pistil is visible, but the ovary is too deeply placed to be characterized as to its pubescence or size.

The third flower, C, is illustrated in lateral view in Figure 5. Its calyx is not in view, and its petals a«

mostly too darkly stained to show the necessary details for a complete description. However, on the petal at

left (arrow), the abaxial surface can be seen to be strigillose, as was the abaxial petal surface in T. glaesaria

(Chambers et al. 2011, Figs. 1, 2), and the petal at the right appears to have the papillate adaxial surface charac-

teristic of flower A. At least 8 much reduced anthers are present on the rim of the staminal tube, in alternating

higher and lower positions between acute lobes as in flower B, above, and in T. glaesaria (op. cit.. Fig. D-

locules of these anthers appear to be more reduced than in flower B. In an apical view of flower C (not shown,

the ovary is glabrous, unlike that of flower A. It is interesting that in two extant Trichilia species, the ovary «s

glabrous before fertilization but develops pubescence afterward, as does the fruit (Pennington et al. 198k P-

11). The intense staining of flower C does not allow further details to be observed. Flowers A and C i e

greatly in anther development; however, as noted by an anonymous reviewer, “(t)he range of anther reduction

between the 3 flowers described here could easily be found in a single modem day species.”

DISCUSSION

The flowers under consideration here were not discovered until our earlier paper had been published.

fossils are linked through their having been excavated from amber mines in the same ocean-deposite

Tertiary strata, now uplifted in the Cordillera Central of Hispaniola. As discussed by Poinar and Poinar (1^ ^ ^

Chambers and Poinar, Fossil (

563

possible that the Trichilia fossils described here do not represent contemporaneous species. What they illus-

trate is that Trichilia taxa in these forests were diverse in features such as perianth pubescence, pistil pubes-

cence, and especially the nature of anther reduction and loss in pistillate flowers. However, lacking evidence

relating to staminate flowers or to differences in the vegetative parts and inflorescence structure of the sampled

taxa, we are limited as to any broad taxonomic conclusions that might be drawn from the observed variability

in pistillate flowers. It is important, nevertheless, that these fossils be placed on record as part of the history of

floral evolution in the genus, to be considered in possible future discussions of Meliaceae phylogeny.

The loss and extreme reduction of anthers in flower A, and probably also in Trichilia antiqua , goes beyond

the description by Pennington et al. (1981) of androecia in functionally female flowers. These authors simply

^te (p. 26) that in Trichilia the “antherodes [are] narrower than anthers, not dehiscing, without pollen ” In

view of the review comments quoted above, this may be a generalization that omits a complete description of

anther abortion and loss in modern species. That two of the Dominican fossils display such androecial reduc-

tion is evidence of its occurrence earlier in Trichilia floral evolution than might have been expected.

As discussed by Pennington et al. (1981), Trichilia is today the largest genus of Meliaceae in the New

World, with ca. 85 species in lowland tropical America as well as ca. 14 species in Africa and 2 in the Indo-

Malesian region. Its floral diversity, particularly in the androecium, well exceeds our sample of amber fossils

(Pennington et al. op. cit., illustrations on pages 36-226). Respecting the age of Trichilia, Muellner et al. (2006)

^ evidence from known Meliaceae fossils, combined with chloroplast rbcL data, to estimate divergence times

» the family. The DNA cladogram presented by these authors includes a monophyletic clade of Trichilia and 1 1

other genera, which is assigned an origin in the Oligocene. Apart from Trichilia, members of this clade are to-

limited to the Old World tropics, from Africa to Madagascar, India, lnto-China, Malesia, and Austroasia.

Muellner et al. (2006) propose a West Gondwanan Cretaceous origin for the family, followed by dispersal

**°ss Eurasia and between Eurasia and North America over the Beringian and North Atlantic land bridges.

Eurther movement from North to South America occurred via island hopping and/or direct land connections

uunng the Tertiary ( see also Muellner et al. 2010, with a dense sampling of Cedreleae). As noted previously

^Chambers et al. 2011), Pennington et al. (1981) recognized only two sections in Trichilia, Sect. Trichilia and

T* Moschoxyhim C. DC. The fossils discussed here are best placed in the emended Sect. Moschoxylum, the

defimn g trai ts of which are the valvate petals and completely united filaments.

565

ACKNOWLEDGMENTS

We thank Andrea Muellner and 2 anonymous reviewers for their helpful comments, which improved the con-

tent and clarity of our presentation.

REFERENCES

Chambe «. K.L, G.O. Poinar, Jr., and A.E. Brown. 201 1 . Two fossil flowers of Trichilia (Meliaceae) in Dominican amber. J. Bot

Res. Inst. Texas 5:463-468.

K.L. and G.O. Poinar, Jr. 2012. A Mid-Tertiary fossil flower of Swietenia (Meliaceae) in Dominican amber. J. Bot.

Res. Inst. Texas 6:1 23-1 27.

Drapb '. G, P. Mann, and J.F. Lewis. 1 994. Hispaniola. In: S. Donovan and T.A. Jackson, eds. Caribbean geology: an introduc-

tion. The University of the West Indies Publishers’ Association, Kingston, Jamaica. Pp. 129-150.

itURRAiDE-ViNENT, M.A. and R.D.E. Macphee. 1966. Age and paleogeographic origin of Dominican amber. Science 273:

1850-1852.

Mueuner ' A.N., V. Savolainen, R. Samuel, and M.W. Chase. 2006. The mahogany family 'out-of-Africa": divergence

estimation, global biogeographic patterns inferred from plastid rbcL sequences, extant and fossil distributi

diversity. Molec. Phylogen. Evol. 40:236-250.

AJU.D. Pennington, A.B. Koecke, and S.S. Renner. 2010. Biogeography of Cedrela (Meliaceae, Sapindales) in Ce

tral and South America. Amer. J. Bot. 97:51 1-518.

p INGT0N ' ID - B.T. Styles, and D.A.H. Taylor. 1981. Meliaceae. FI. Neotropica 28:1 -470.

G -°” jR - R- Poinar. 1 999. The amber forest. Princeton University Press, Princeton, NJ.

*** D - 1999- Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C, 28.

566

BOOK REVIEW

Scott Calhoun. 2012. The Gardener’s Guide to Cactus: The 100 Best Paddles, Barrels, Columns, and

Globes. (ISBN-13: 978-1-609469-200-6, pbk.). Timber Press, Inc., The Haseltine Building, 133 S.W.

Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: www.timberpress.com, 800-

327-5680). $24.95, 227 pp„ color throughout, 8” x 9”.

Thinking of going to the Desert Botanical Garden in Phoenix, Arizona? If so, now is the time to read Scott

Calhoun’s The Gardner’s Guide to Cactus: The 100 Best Paddles, Barrels, Columns, and Globes. The juxtaposition

of Dale Chihuly’s innovative glass sculptures with cactus and succulents was a stroke of genius. Likewise is Mr.

Calhoun’s new cactus guidebook.

Succulent plant expert Scott Calhoun has selected 100 of the best cactus available and shows how they

can make striking additions to desert gardens or serve as dazzling container specimens. His writing style is

one which offers a clean, helpful, well-presented text with reliable design suggestions.

Mr. Calhoun’s stunning photograph on the book cover clearly demonstrates to the book publishing world

and its gardening readership how easily cactus appreciation can be for everyone— not just specialists. He is the

quintessential model of cactus authorship. It is a great book for beginners and expert gardeners.

Paddles, barrels, columns, and globes add new meaning to the cactus gardening neophyte! One look at

the Tuna Colorado cactus in its purple high-winter coloration photograph at the Desert Botanical Garden sur-

rounded by a purple Chihuly glass sculpture suggests Mr. Calhoun’s new book is set apart from normal cactus

guidebooks. First of all, it is a total delight to look at!

As an effort to simplify the issues related to plurals, Mr. Calhoun has chosen to call the plants cactus. He

describes cactus conservation and defines Crassulacean Acid Metabolism (CAM). A glance at the Introduction

and Table of Contents describes planting and care, cactus planted in the ground and containers, tools of the

prickly trade, and the art of showing cactus. Mr. Calhoun not only describes how one plants and cares for

cactus; he gives tips for staging cactus at shows sponsored by the Cactus and Succulent Society of America

(CSSA). The 100 plants are grouped as follows: low and mounding, barrels and globes, paddles and rods, and

columns. Each entry has a colorful photograph, scientific and common names, habitat, mature size, hardiness,

flowering season, cultivation, design suggestions, and, yes, culinary value!

In addition, the book has extensive cross-referenced index, bibliography, plant hardiness tables, cactus

pests, cactus for special purposes, and selected cactus nurseries. One can tell after reading The Gardner’s Guide

to Cactus: The 100 Best Paddles, Barrels, Columns, and Globes, cactus sagacity can be great fun and rewarding.

When is your first CSSA entry to demonstrate what you have learned from Mr. Calhoun ?— Kay M.

Stansbery, Ph.D., Library Volunteer, Botanical Res

76107-3400, U.SA.

e of Texas, 1700 1

y Dr., Fort V

I. Bot. R«. Inst Texas 6(2): 566. 2012

LIMITATIONS TO NATURAL PRODUCTION

OF LOPHOPHORA WILLIAMSU (CACTACEAE)

II. EFFECTS OF REPEATED HARVESTING AT TWO-YEAR INTERVALS

IN A SOUTH TEXAS POPULATION

Martin Terry

KeeperTrout Bennie Williams

Sul Ross State University

Alpine, Texas 79832, U.SA.

Cactus Conservation Institute

P.O.Box 561

Alpine, Texas 79831, U.S.A.

Cactus Conservation Institute

P.O.Box561

Alpine, Texas 79831, U.S.A.

Teodoso Herrera

Norma Fowler

P.O. Box 460346

San Antonio, Texas 78246, U.S.A

The University of Texas at Austin

Department of Integrative Biology C0930

1 University Station

Austin, Texas 78712, U.SA.

RESUMEN

568

Journal of the Botanical Research Institute of Texas 6(2)

Lophophora williamsii (Lem. ex Salm-Dyck) J.M. Coult. (Cactaceae), known as peyote both in Spanish and in

English, is a small cactus (rarely exceeding 10 cm in diameter) of northeastern Mexico and adjacent border

areas of Texas. The aerial crowns of plants are approximately hemispherical in shape. Some plants are caespi-

tose; i.e., they have multiple crowns arising from a single rootstock. The literature on the biology of this plant

up to the mid-1990s is summarized by Anderson (1996), who first suggested that the species might be endan-

gered by overharvesting (Anderson 1995).

There is active commercial trade in the harvested crowns of peyote, which are collected and sold by li-

censed distributors to the Native American Church (NAC) for religious use as protected by U.S. law. There is

substantial concern that the rate of harvest of peyote from wild populations is not sustainable. Anecdotal re-

ports by members of the NAC include descriptions of the decline or decimation of natural populations and a

decrease in both the availability and the quality of peyote being offered for sale in the regulated peyote market

(TH, pers. obs.). A number of papers in the scientific literature have described the decline of peyote in its native

habitat, apparently due to overharvesting (Anderson 1995; Trout 1999; Terry & Mauseth 2006; Powell etal.

2008; Terry 2008a, b,c; Terry et al. 2011). Despite such reports involving both Texas and Mexican populations,

the species is not (yet) considered in danger of extinction (NatureServe 2012; Fitz Maurice and Fitz Maurice

2009), except in Texas, where NatureServe determined it to be in the S4 (imperiled) category. The work of

Terry et al. (2011) was the first experimental investigation of the effects of harvesting on peyote plants in situ.

In that paper we reported the effects that were detectable two years after the initial harvest. The present report

focuses on effects detectable four years after the initial harvest.

MATERIALS AND METHODS

The study site was described in Terry et al. (2011). Because of the multi-year duration of the ongoing study and

the complexity of the study design, it is appropriate to provide a clear, detailed description of what was done to

which plants, and when.

At the start of the study, in March 2008, 100 L. williamsii plants that appeared not to have been previously

harvested were individually numbered and tagged along a transect through the population. The number of

crowns on each plant was counted and the horizontal diameter of each crown was measured. Fifty of these

plants that were single-crowned were then harvested (i.e., the crown of each plant was cut off transversely at

ground level and removed), and the other 50 plants (most but not all of which were single-crowned) were left

unharvested as controls. The harvested crown of each plant in the harvested group was weighed, to determine

the harvested fresh biomass obtained from each of these “virgin” plants.

At the end of the second year of the study, in March 2010, all surviving plants from the original groups of

50 harvested and 50 control plants were located, the number of crowns on each plant was counted, and the

diameter of each crown was measured. Then the 43 surviving plants in the harvested group were divided into

two subgroups: 20 multiple-harvest plants and 23 single-harvest plants. All regrowth crowns were harvested

from the 20 multiple-harvest plants, leaving these now twice-harvested plants without crowns (and thus with-

out photosynthetic tissue) for the second time in two years. Reharvest at two-year intervals is typical in current

commercial harvest (MT, pers. obs.). The harvested crown(s) of each multiple-harvest plant were weighed to

obtain harvested fresh biomass at a second harvest. A comparison of harvested biomass between the 2008 and

2010 harvests was reported by Terry et al. (2011). The single-harvest plants were not reharvested, and the sur-

viving plants of the 50 original control plants continued to serve as unharvested controls.

At the end of the fourth year of the study, in March 2012, all surviving plants were again located, counted,

and measured. In addition, all new regrowth crowns were again harvested and weighed from the 16 surviving

plants in the multiple-harvest treatment. In summary, control plants have never been harvested, single-harvest

All stalls

lalyses were done with SAS 9.1 (

:, Cary, NC, USA).

Teny et al., Harvesting effects on wild populations of Lophophora in Texas

569

RESULTS

Survival. — Of the 100 plants of the initial (2008) census, 4 (2 control, 2 harvested) were dug up by feral hogs

and were therefore dropped from all further analyses, leaving 96 plants. Of these 96 plants, 6 (1 control, 5

harvested) died before the second (2010) census. Ninety plants were still alive in 2010 (census 2): 47 control

plants and 43 plants that had been harvested in 2008. Of these 43 surviving plants that had experienced one

harvest, 23 were assigned to the single-harvest treatment and 20 were assigned to the multiple-harvest treat-

Of the 47 control plants alive in 2010, 45 were still alive at census 3 in 2012. Nineteen of the 23 single-

harvest plants (83%) and 16 of the 20 multiple-harvest plants (80%) were still alive in 2012. By 2012 the sur-

vival rate of control plants from census 1 through census 3 was significantly higher than the survival rate of

harvested plants over the same interval (94% [45/48] versus 73% [35/48], x 2 = 8.65, P = .0033; Fig. 1).

Harvested mass .— Weights of all the crowns of a plant were summed to calculate harvested fresh mass per

plant. Mass per crown was calculated for each plant by dividing its total mass by its crown number; these val-

ues were then averaged for statistical analysis and for Figure 2. Average harvested mass per plant decreased

from census to census: 44% between the first and second censuses and 32% between the second and third

censuses (Fig. 2, solid line). The differences between harvests were significantly different from zero (paired t-

tests: harvest 1 vs harvest 2: 14.0 g average difference, N = 20, t = 6.73, P < 0.0001; harvest 2 vs harvest 3: 6.9 g

average difference, N = 16, t = 4.24, P = 0.0007).

Plants initially responded to harvesting by increasing the average number of crowns per plant (Fig. 2,

dashed line), although this increase was not sufficient to counterbalance the decrease in mass per crown (Fig.2,

dashed and dotted line). After the second harvest, both the average number of crowns per plant and the average

mass per crown decreased.

Volume.— The above-ground volume of each plant was estimated by first estimating the volume of each

trown as a hemisphere from its measured diameter: estimated volume = % n (diameter/2) 3 .

The estimated volumes of all the crowns on the plant were then summed to estimate total plant above-

ground volume. The estimated volume of each plant in 2012 was very closely correlated with its harvested

fresh mass in 2012 (Fig. 3).

Volumes were log-transformed before analysis of covariance (ANCOVA) to improve normality of the re-

siduals. Volume at census 2 (2010) was used as a covariate. Treatments did not differ significantly in their

sfopes: the slope of the relationship between log-transformed volume in 2010 (x-axis) and log-transformed

volume in 2012 (y-axis) was the same for each treatment. Therefore the final ANCOVA model assumed equal

Note that equality of slopes in a model fitted to log-transformed data does not imply that slopes will be

near w hen untransformed data are graphed on a linear scale (e.g., Fig. 4).

Estimated plant volume at census 3 (2012) was closely related to estimated plant volume at census 2; 74%

the variation in the former was explained by variation in the latter amount. Treatment accounted for an ad-

r°nal 10% of the variation among plants at census 3. The effects of the single-harvest treatment did not differ

,llose of the control (Scheffe contrast, Fj 75 = 1.33, P * 0.25), but each of these treatments differed signifi-

cantly fro m the multiple-harvest treatment (Scheffe contrasts; control versus multiple-harvest: F 1J5 = 29.20, P

< 0001; single-harvest versus multiple-harvest, F, 75 = 41.46, P < 0.0001; Fig. 4). In other words, the surviving

^-harvest plants were growing (on a logarithmic scale) about as fast as the surviving control plants be-

, een 2010 and 2012, but they began the interval with much smaller sizes than the control plants. In contrast,

mu ltiple-harvest plants were decreasing in size. For example, the final ANCOVA model predicts that a

COntro1 P lant with a volume of 10.0 cm 3 , the average size of all 90 surviving plants at census 2, would have

to 21.9 cm 3 and a single-harvest plant of the same size would have grown to 26.3 cm 3 (not significantly

1 erent from 21.9 cm 3 ), but a multiple-harvest plant with a volume of 10.0 cm 3 would have decreased slightly

nsi ze, to 9.66 cm 3 .

Regional harvesting trends in South Texas .— Annual peyote sales data covering the years 1986-2011 (Texas

Anient of Public Safety, unpublished data) are presented in Figure 5. Although these figures do not in-

570

Journal of the Botanical Research Institute of Texas 6(2)

census

s. Solid line: control plants; dotted line: harvested plants.

elude all sales of peyote (Terry et al. 2011), it is reasonable to assume that the number of buttons sold in the

regulated trade is positively correlated with the total number of buttons harvested in the region of South Texas

known as “the Peyote Gardens.” In 2011, the DPS-regulated peyote sales totaled slightly over 1.4 million but-

tons, continuing the generally downward trend which such sales have followed since 1997. It is noteworthy

that prior to the current decline there was a decrease in numbers of buttons sold during the late 1980s that

appears to have corresponded to the historical decline in the available harvest of mature plants, followed in the

early to mid-1990s by a marked increase in numbers of buttons sold when the proliferation of small regro

buttons began to be harvested to meet the needs of the NAC. Anecdotal accounts from NAC meetings during

the period of temporary increase in numbers of buttons noted the prevalence of fresh buttons as small as dira®

(TH & KT, pers. obs.). The number of buttons sold in 2011 was the lowest for any year in the last quarter o a

century. As the annual number of buttons sold has declined steadily since 1997, the price has shown a ma

increase; the price per button is roughly equal to total sales (in U.S. dollars) divided by the number of buttons

sold.

DISCUSSION

Effects of harvesting on plant survival and growth

The negative effects of harvesting on survival may be delayed. The initial harvest did not significantly reduce

survival during the first two years after the harvest (2008 to 2010; Terry et al. 2011), but its effects were higWy

significant by 2012 (73% survival to the four-year time point in 2012 among plants harvested in 2008, vers®

94% among control plants). Any delayed effects of the second (2010) harvest on survival were not yet evide»

in 2012. sub-

file effects of precipitation may also be delayed. The six months preceding the 2010 census received su

era 9 e fresh above-ground mass per harvested plant, number of crowns per plant, and average fresh mass pc

wrage fresh mass per crown, an average was calculated for all cr r _

Ptots were used to calculate the values in this graph. Vertical bars: 1 standard error.

n at each date. To calculate

stantially more rain (32.4 cm October-March precipitation) than the six months periods preceding the other

censuses (6.0 cm and 15.4 cm October-March precipitation preceding the 2008 and 2012 censuses, re-

U.S. Department of Agriculture 2012), but the average size of control plants declined in the first in-

terval and incre ased in the second (Fig. 4). However, it may be that in wetter years peyote experiences more

“^Petition from other plants that have responded rapidly to the increased soil moisture. Harvesting also sig-

m Cantl >’’ and strongly, affected plant growth rate and therefore plant size (Fig. 4). Each harvest reduced plant

tom ratCS ThC 2008 harvest reduced the average growth rate of all harvested plants (Terry et al. 2011). The

harvest of the multiple-harvest plants significantly reduced their growth rate below that of the single-

est plants (harvested only in 2008) as well as below the growth rate of the never-harvested control plants

/ & 4). While the single-harvest plants and the controls had about the same growth rate between 2010 and

-the single-harvest plants were so much smaller in 2010 (due to the 2008 harvest) that they remained

. smaller than the control plants in 2012. Meanwhile, the multiple-harvest plants continued to decline in

tween 2010 and 2012. Plant size and plant survival are usually highly correlated (Harper 1977), so we

1*^0 ^ continuing excess mortality of the multiple-harvest plants,

infec . re are P r °bably several reasons why harvesting reduces growth rate, size and survival rate. Microbial

out ft, 0 " ° f Ae ° PCn Wound crated by the act of harvesting the crown of a plant, for example, cannot be ruled

Mon mectlanism that appears to be an inevitable consequence of harvesting is that of exhaustion due to

n ged deprivation of solar energy. The crown, being the only aerial organ of the peyote plant, is the plant’s

Journal of the Botanical Research Institute of Texas 6(2)

only site of photosynthesis. Without photosynthesis, the plant cannot use solar energy to create and store car-

bohydrates, and nutrient, carbon, and water uptake are greatly reduced. The crown is also the part of the plant

that is always harvested for ceremonial use. When the crown is harvested— thereby becoming a button in the

peyote trade — the plant’s ability to photosynthesize is ipso facto reduced to zero. The harvested plant then uses

stored energy, nutrients, and water to regrow its above-ground biomass. If reharvesting occurs before the p

has had time to rebuild its stored reserves from photosynthesis in its regrown above-ground tissue, it w

come successively smaller at each harvest and eventually die. The reduced size and growth rate, and increase

mortality, of harvested plants strongly support the hypothesis that a two-year cycle of harvesting of this spe-

cies is too frequent for plant recovery. A sustainable frequency of harvesting would be low enough to allow a

plant to fully regrow and to fully rebuild its supply of stored resources between harvests. We hope eventua }

to be able to determine the maximum sustainable harvesting frequency. Whatever that frequency may be, our

As is true of most plant species, the removal of the apical meristem (part of the harvested crown of pep

te), in addition to stimulating regrowth, probably also de-represses axillary meristems, resulting in the forma-

tion of multiple crowns. In the absence of the continual secretion by the apical meristem of the hormone t

normally suppresses lateral branching (presumably auxin, based on Mauseth and Halperin 1975), one <*

more) of the axillary meristems in the areoles on the subterranean portion of the stem is de-repressed and

gins to form a new crown at the apex of a lateral branch which emerges from the subterranean stem and g roWS

toward the surface of the ground. This phenomenon accounts for harvested plants of this species having ®

crowns than unharvested plants (Fig. 2). It is a temporary phenomenon, however, because eventually a P la ^ s

stored resources are exhausted by too frequent harvesting, causing the number of crowns per plant to e

573

census

single-harvest

multiple-harvest plants

► average - control plants

* average - single-harvest plants

* average - multiple-harvest plants

buttons sold (millions)

574

annual numbers of buttons sold (in millions)

•••&•• total sales (in thousands of US $)

Fig. 5. Annual peyote sales by licensed distributors in South Texas from 1986 through 2011.

crease. This stage appeared to be reached at the second harvest: the number of crowns per plant increased after

the first harvest but declined after the second.

The negative impact of harvesting will be greater if portions of the subterranean stem are also removed

To avoid such damage to the harvested plants, in this study we used only best harvesting practices, viz., cutting

the crown at its base, parallel to the surface of the ground. However, commercial harvesting practices may re-

move a substantial portion of the subterranean stem along with the crown (Terry & Mauseth 2006). Such re-

moval of subterranean stem tissue reduces the number of areoles available to initiate lateral branching, reduc-

ing the number of new crowns that can be formed. In addition it removes even more resources from the plant,

reducing the amount available for regrowth.

Changes in harvest yield over time

As a result of the effects of harvesting on plant size, the yield per plant of harvested biomass decreased after

each biennial harvest, first by 44% and then by 32% (Fig. 2, solid line). If one includes mortality in these calcu-

lations, the decrease in harvest yield is even more marked: the third harvest produced only 25% of the bioma*

that the first harvest did. For example, if we had begun with 100 plants, the first harvest would have yield* 1

3125 g (i.e., 100 plants x 31.25 g/plant), the second harvest would have yielded 1547 g (100 plants x 0.895 sur-

vival rate of harvested plants 2008-2010 x 17.27 g/plant), and the third harvest would have yielded 769 g (10°

plants x 0.895 x 0.800 survival rate 2010-2012 x 10.73 g/plant), declines of about 50% per harvest. The absolute

(as opposed to relative) decline was smaller in the second two-year period, but only because it began fro® 4

Terry etal., Harvesting effects on wild populations of Lophophora in Texas

575

lower baseline. These are exactly the effects on harvest yield to be expected if harvesting is occurring too fre-

quently for plants to regrow and to rebuild their stored resources.

The fact that harvested mass was even more closely correlated with estimated volume at census 3 (Fig. 3)

than at census 2 (Fig. 3 in Terry et al. 201 1) may seem surprising in that the crowns of most adult peyote plants

do not appear to have a true hemispherical shape, but rather the shape of half of an oblate sphere. However,

many of the younger plants — and especially young regrowth crowns — do indeed have vertically extended

crowns, and all peyote plants tend to expand vertically in response to rain (MT, pers. obs.), which would tend

to balance out the more flattened shape of the adults and the flattening effect of drought, over time.

Peyote shows many of the hallmarks of a classic case of unsustainable harvesting of a wild resource. First, the

decline in total harvest combined with an increase in price/unit is characteristic of overharvested wild species

(cf. Fig. 1 in Schippman et al. 2002). A declining number of wild plants is a likely explanation for the failure of

the harvest to increase in response to the increase in unit price (because a declining population causes de-

there are still individuals to be harvested (Hilbom & Walters 1992; Thurstan et al. 2010). Second, there are

anecdotal reports of declining unit (button) size (TH, pers. obs.). Declining body size is another classic indica-

tor of overharvesting (Stergiou 2002; Berkeley et al. 2004; Genner et al. 2010). Third, there are anecdotal re-

ports of declining quality of the harvested buttons (TH, pers. obs.). Fourth, the harvesting frequency (every

other year) shown to be unsustainable by the present study is typical. Finally, our results may underestimate

impacts of harvesting, as our harvests may have been less damaging to individual plants than a commercial

harvest, due to the care taken in the harvests of this study.

As far as we are aware, this study is the first well documented case of overharvesting of a cactus species

(but see Jimenez-Sierra and Eguiarte 2010, in which browsing was also involved). It is also one of a limited

number of well documented cases of overharvesting of non-timber plant species in general. Most well docu-

mented cases of overharvesting of wild resources involve marine and freshwater animal species (Jackson et al.

2001; Allen et al. 2005; Genner et al. 2010). There are detailed reports of overharvesting of many tree species

(e g., Schwartz et al. 2002; Schulze et al. 2008). There are some detailed reports of overharvesting of herbaceous

plant species, of which ginseng ( Patiax quinquefolius) is perhaps the best documented (Nantel et al. 1996; Mc-

Graw 2001; Case et al. 2007; McGraw et al. 2010). However, many hundreds (at least) of other plant species are

threatened by overharvesting, especially plant species harvested for medicinal uses (Schippman et al. 2002),

for lumber (Oldfield et al. 1998), or for collectors (Oldfield 1997).

The regulatory panorama

At the moment there are only two major interested parties with any standing in the discussion about the fate of

P^ote in its natural habitat: (1) the Native American Church (NAC), whose right to consume peyote for reli-

cs purposes is protected by legislation such as the American Indian Religious Freedom Act (AIRFA), and (2)

the Drug Enforcement Administration (DEA), which is obligated by the Controlled Substances Act to regulate

^ use and distribution of peyote by and for the NAC, and to prevent the diversion of peyote to non-authorized

Persons. Neither of these parties is speaking very audibly about regulatory solutions to mitigate the deteriorat-

ln g state of the wild peyote populations. This is unfortunate, as the problem has a feasible solution, namely the

regulated cultivation of peyote by and for the NAC, which would reduce the harvesting pressure on the wild

Populations (as in, e.g., Kay et al. 2011). Furthermore, this solution is technically within reach (Chandra et al.

311(1 oulturally acceptable (TH, pers. obs.). The barrier to bringing this solution to fruition is essentially

a regulatory one. Cultivation of L. williamsii is anticipated in the American Indian Religious Freedom Act (as

tended 1994), which “. . .does not prohibit such reasonable regulation and registration by the Drug Enforce-

Administration of those persons who cultivate. . .peyote. . ..” But to date no interested party (e.g., the NAC

° orth America) has petitioned the DEA to promulgate any such “reasonable regulation” spelling out the

QClaik for SUc h registration. Pending such action, cultivation of peyote, though not illegal, lacks the needed

te ** ulator y framework to provide legal certainty and protection for NAC members who would prefer to produce

populations of peyote continue to produce steadily decreasing yields, as demonstrated in this study and in the

regulated peyote market. Under the current system— which can accurately be described as “management by

extirpation” — at some point the conservation crisis will become so critical that the U.S. Fish and Wildlife Ser-

vice will be obligated by the terms of the Endangered Species Act to evaluate the conservation status of the

species Lophophora williamsii. At that point the regulatory situation will become substantially more complex.

If a regulatory stalemate then ensues, the NAC’s options may broaden (or narrow) to include the Supreme

Court and/or Congress as sources of relief.

ACKNOWLEDGMENTS

We are most grateful to C.W. Hellen Ranches, Ltd. - La Mota Division - Charles W. (Bill) Hellen, Managing

Partner, for providing access to his ranch and good company. We also thank the younger generations of the

Herrera and Terry families for their help with the hard labor involved in the logistics of conducting the study.

Garry Stephens kindly provided USDA Field Office climate data for Hebbronville, Texas. Essential funding for

the study was generously provided by Libbie and Jerald Mize, the Alvin A. and Roberta T. Klein Foundation,

Sul Ross State University (in the form of a Research Enhancement grant), and all the donors supporting the

scientific work of the Cactus Conservation Institute, Inc. We appreciate the helpful comments of Michael Pow-

Allen, J.D., Ft Abell, Z.E.B. Hogan, C. Revenga, B.W. Taylor, R.L. Welcomme, and K. Winemiller. 2005. Overfishing of inland waters.

Bioscience 55:1041-1051.

Anderson, E.F. 1995. The "Peyote Gardens' of South Texas: A conservation crisis? Cact. Succ J. (U.S.) 67:67-73.

Anderson, E.F. 1996. Peyote: The divine cactus. University of Arizona Press, Tucson.

Berkeley, SA., MA Hixon, R J. Larson, and M.S. Love. 2004. Fisheries sustainability via protection of age structure and spatial

distribution offish populations. Fisheries 29:23-32.

Case, M.A., K.M. Funn, J. Jancaitis, A. Alley, and A. Paxton. 2007. Declining abundance of American ginseng ( Panaxquinque-

folius) documented by herbarium specimens. Biol. Conservation 134:22-30.

Chandra, B„ L Palni, and S. Nandi. 2006. Propagation and conservation of Picrorhiza kurrooa Royle ex Benth.: an endan-

gered Himalayan medicinal herb of high commercial value. Biodivers. & Conservation 1 5:2325-2338.

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Version 201 2.1 . <www.iucnredlist.org>. Accessed: 28 June 201 2.

Genner, M J., D.W. Sims, AJ. Southward, G.C. Budd, P. Masterson, M. McHugh, P. Rendle, EJ. Southall, VJ. Wearmouth, and SJ.

Hawkins. 2010. Body size-dependent responses of a marine fish assemblage to climate change and fishing over a

century-long scale. Global Change Biol. 16:51 7-527.

Harper, J. 1977. Population biology of plants. Academic Press, London.

e, dynamics, and uncertainty. Chapman

Jackson, J.B.C., M.X. Kirby, W.H, Berger, K A Bjorndal, L.W. Botsforo, B J. Bourque, R.H. Bradbury, R. Cooke, J. Erlandson, J A Es®>

T.P. Hughes, S. Kidwell, C.B. Lange, H.S. Unihan, J.M. Pandolfi, C.H. Peterson, R.S. Steneck, M J. Tegner, and R-R. Warner- 2001-

Historical overfishing and the recent collapse of coastal ecosystems. Science 293:629-637.

JimEnez-Sierra, C. and L Egularte. 2010. Candy barrel cactus ( Echinocactus platyacanthus Link & Otto): a traditional P^

Kay, J. r A. Strader, V. Murphy, L. Nghiem-Phu, M. Calonje, and M.P. Griffith. 201 1 . Palma Corcho: a case study in botanic garde*’

conservation horticulture and economics. HortTechnol. 21:474-481.

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of American ginseng (Panax quinquefolius L.). Nat. Areas J. 30:202-210.

Nantel, P, D. Gagnon, and A. Nault. 1996. Population viability analysis of American ginseng and wild l<

stochastic environments. Conservation Biol. 10:608-621.

NatureServe. 2012. NatureServe Explorer: An online encyclopedia of life [web application]. Version 1

Arlington, Virginia. Available at http://www.natureserve.org/explorer (Accessed: July 3, 201 2.)

Outfield, S. (ed). 1997. Cactus and succulent plants— Status survey and conservation action plan. IUCN/SSC Cactus and

Succulent Specialist Group. IUCN, Gland, Switzerland and Cambridge, UK.

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Powell, A.M., J.F. Weedin, and S.A. Powell. 2008 . Cacti of Texas: a field guide. Texas Tech University Press, Lubbock.

Schippmann, U„ DJ. Leaman, and A.B. Cunningham. 2002. Impact of cultivation and gathering of medicinal plants on biodiver-

sity: global trends and issues. In: Departmental Working Group on Biological Diversity for Food and Agriculture, FAO

(ed). Biodiversity and the Ecosystem Approach in Agriculture, Forestry and Fisheries. Satellite event on the occasion

of the Ninth Regular Session of the Commission on Genetic Resources for Food and Agriculture. Rome, 1 2-1 3 Octo-

ber 2002. FAO, Rome. Available at http://www.fao.org/DOCREP/005/AA010E/AA010E00.HTM

Schulze, M, J. Grogan, C. Uhle, M. Lenttni, and E. Vidal 2008. Evaluating ip# (Tabebuio, Bignoniaceae) logging in Amazonia:

sustainable management or catalyst for forest degradation? Biol. Conservation 141 :2071 -2085.

Schwaru, M.W., T.M. Caro, and T. Banda-Sakala. 2002. Assessing the sustainability of harvest of Pterocarpus angolensis in

Rukwa Region, Tanzania. Forest Ecol. Managem. 170:259-269.

Stehgiou, K.1. 2002. Overfishing, tropicalization of fish stocks, uncertainty and ecosystem management: resharpening

Ockham's razor. Fish. Res. 55:1-9.

Terry, M. 2008a. Stalking the wild Lophophora. Part 1 . Chihuahua and Coahuila. Caet. Succ. J. (US) 80:1 81 -1 86.

Terry, M. 2008b. Stalking the wild Lophophora. Part 2. Zacatecas, San Luis Potosi, Nuevo Leon, and Tamaulipas. Cact

Succ. J.(U.S.) 80:222-228.

Terry, M. 2008c. Stalking the Wild Lophophora. Part 3. San Luis Potosf (central), Quer#taro, and Mexico City. Cact. Succ.

J. (US.) 80:310-31 7.

Terry, M. and J.D. Mauseth. 2006. Root-shoot anatomy and post-harvest vegetative clonal developnr

williamsii (Cactaceae: Cacteae): implications for conservation. Sida 22:565-592.

Terry, M, K. Trout, B. Wiluams, T. Herrera, and N. Fowl£r. 201 1 . Limitations to natural production of Lophophora willi

(Cactaceae) I. Regrowth and survivorship two years post harvest in a South Texas population. J. Bot. Res. Inst

5:661-675.

Texas Department of Public Safety. Peyote sales data. Unpublished.

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578

BOOK REVIEW

Ellen Sousa, Foreword by William Culuna. 2011. The Green Garden: A New England Guide to Planning,

Planting, and Maintaining the Eco-Friendly Habitat Garden. (ISBN-13: 978-1-59373-091-8, pbk).

Bunker Hill Publishing, 285 River Road, Piermont, New Hampshire 03779, U.S.A. (Orders: www.bun-

kerhillpublishing.com). $34.95, 224 pp., color throughout, 7" x 10".

A growing number of people are becoming more agricurious. They are interested in how things are grown,

land stewardship, animal husbandry, and soundscape ecology.

Thus, planning an eco-friendly habitat garden and landscape is like becoming a creative gourmet cook

orchestrating a perfect dinner party with tried and tested recipes. Ellen Sousa’s The Green Garden: A New Eng-

land Guide to Planning, Planting, and Maintaining the Eco-Friendly Habitat Garden is the ideal manual/reference

book to use if one wants to establish a notable green garden, particularly for those wanting to stop the deteriora-

tion of the natural world around us. Ms. Sousa presents new concepts for gardening that allow the gardener to

work with nature to develop an area’s natural ability to maintain its own health and sustainability. Her cogent

writing style is a true testimonial that she is a cognoscente of natural habitat gardening. She and her husband

live on a farm landscaped as a Certified Wildlife Habitat and Monarch Waystation. Renowned native plant au-

thor William Cullina’s Foreword in the book is a validation that Ms. Sousa is the real deal!

Even though the title suggests it is only about the New England area of the country, it is not. Remember,

the fight for establishing eco-friendly habitats is a universal challenge.

Fifty-plus contributing photographers fill this book with amazing wildlife photos and native plants. It is

not only a wonderful primer for those just learning about natural habitat gardening but also a valuable educa-

tional resource for seasoned gardeners seeking to increase their eco-friendly habitat gardening knowledge,

A quick glance at the Table of Contents denotes the principles Ms. Sousa cites are applicable from forests

to seasides, from lots to farmlands and, yes, to what Victorian gardeners called a “moon garden” of fragrant

tese are for gardeners who are only able to enjoy their garden during the evening

In addition, the book includes a guide of the best plants for the region. The plants are grouped by plant

type in alphabetical order by genera. Common names are listed after each plant genus name. A key of numer-

ous icons is included.

Thoreau wrote “the world is but a canvas for your imagination” — when are you going to begin developing

your own Eden?— Kay M. Stansbery, Ph.D., Library Volunteer, Botanical Research Institute of Texas, 1700 Univer-

sity Dr, Fort Worth, Texas 76107-3400, U.S.A.

J. Bot Res. Inst. Texas 6(2): 57S. 2012

A COST-EFFECTIVE METHOD FOR CONSTRUCTING

MAGNETIC FUMICELS FOR HERBARIUM CABINETS

Richard Carter

Director, Herbarium (VSC), Biology Department

Valdosta State University

Valdosta, Georgia 31698-0015, U.S.A.

ABSTRACT

RESUMEN

INTRODUCTION

Substantial literature exists on the importance and efficacy of various pest repellants in herbaria and their

health risks (e.g.. Hall 1988; Strang 1999). However, methods for containing and securing repellants such as

naphthalene and paradichlorobenzene (PDB) in herbarium cabinets have received little attention. A variety of

methods are used for containment in herbaria, e.g., cloth bags with drawstrings, paper envelopes, small paper

bags, small cardboard boxes, or small compartments in cabinet doors, and some entomologists use fumicels

constructed from small cardboard boxes with wire mesh tops (Anonymous 2012). Because naphthalene and

PDB vapors are heavier than air and tend to settle at the bottom of the cabinet, the repellant is normally placed

011 t0 P uppermost folder of specimens in the uppermost pigeonhole of each column of the herbarium

^net. However, there are several annoying aspects of this widely used method. When specimens are re-

trieved from the uppermost pigeonholes, the packets of repellant fall out, often striking the individual remov-

mg the specimens, or the packets are inadvertently pushed toward the back of the pigeonhole where they slip

°w of sight. Retrieving the “lost” packets usually requires use of a ladder or step-stool, which is time-consum-

wg and potentially hazardous.

I devised, tested, and recommend placement of solid repellant in fumicels constructed from acid-free

8“Ssine envelopes suspended from the metal ceilings of herbarium cabinets by small neodymium magnets

ed to small rectangles cut from herbarium paper. This method uses archival materials, is cost effective,

a «d— except for magnets and glassine envelopes — employs materials commonly available in the herbarium.

MATERIALS AND METHODS

United States, neodymium magnets and glassine envelopes are normally sold by English units; thus,

x 8 9 UnitS are P rilT tarily employed here. Paper magnet-holders were made from 2 Vi inch x 3V4 inch [6.4 cm

Thec Cm ] P a P er rectangles cut from 0.12 caliper archival herbarium paper using a guillotine type paper cutter.

‘^terofeach rectangle was marked usinga wooden pencil, and a V4 inch x Vie inch [6.4 mm x 1.6 mm] N42

^ 1Um (NdFeB) disc magnet was affixed at the center with a small dot of water-base herbarium glue. The

^ ve was allowed to dry, and the paper magnet-holder was placed in a 2)4 inch x 3)4 inch [7 cm x 9.5 cm]

matehT’ glassirie envelope with a single opening along one 2)4 inch end and a foldable end flap. Approxi-

Og of naphthalene (three Enoz® moth-balls) was placed in the packet against the surface of the paper

Vnolder opposite the magnet. The envelope was closed, and a 1V+ inch stainless steel paper-clip was

3.75 inches

placed across one corner to secure the flap. See Figs. 1, 2 and 3. Alternatively, the loose flaps can be taped

closed, but this would most likely prevent reuse of envelopes. Glassine paper is relatively non-porous and ini'

pervious. Therefore, 1 punctured the lower surface of each gWw envelope about 20 times with a steel sewing

needle to increase its permeability to naphthalene.

Magnets.— % inch x Vis inch [6.4 mm x 1.6 mm] Grade N42 Neodymium (NdFeB) Disc Magnet, 3x nickel

plated, pull force 2.5 lbs: MAGNETman® via amazon.com®. http://www.amazon.com/shops/A2WOM254L'

CFKNF

Glassine envelopes.— Vh inch x Vh inch [7 cm x 9.5 cm] acid-free, glassine envelope with foldable end Ihp

and single opening along 2Y4 inch end: Uline®, 12575 Uline Drive, Pleasant Prairie, W1 53158, USA. h«F

RESULTS AND DISCUSSION

The major expense was the cost of the neodymium magnets ($0.24 each, including shipping; pack of 1°°**

$12.99, excluding shipping), the acid-free, glassine envelopes ($0.07 each, including shipping; box of lOOOfor

jft-OO, excluding shipping), and stainless steel paper-clips ($0.07 each, including shipping; box of 500 for

9.95, excluding shipping). The cost of the other, commonly used materials was negligible. Thus, I equipped

rtwrium cabinets, each with two fumicels, at a cost of less than $0.40 per fumicel. Although 1 recommend

siting the fumicels to ceilings of cabinets, they also seem to work well on cabinet doors, which might be a

consideration when the uppermost pigeonholes are packed with specimens.

j. ^ ve t0 l hcir size, neodymium magnets are very strong, and they are readily available from several on-

vendors. Marketed as permanent, they are subject to chipping if allowed to snap together. Although I used

^ 1 10g of repellant per fumicel, these small magnets are sufficiently strong to suspend at least 20g of

k Kpellant (Le., six moth-balls and the maximum capacity of the fumicel envelope) securely, and they can

^easily detached from the herbarium cabinet. In addition to being dangerous, larger, stronger neodymium

C al^ CtS 310 m ° re ex P ens * ve and would require considerable effort to detach from the metal surface of the

Caching them would most likely result in tearing both glassine envelope and paper mag-

«ver magnet £ rade and size recommended herein seem ideal for this particular application. How-

iUm ma S nets c ome in a variety of sizes and shapes, and heavier containers could easily be at-

e Wlth larger, stronger magnets.

This article should not be construed to be a recommendation by either the author or this journal

th at ^ g h n ^ hthalene ’ PDB » or other pest repellants in herbaria. Rather, its purpose is to provide information

* beneficial t0 th °se who find it necessary to use repellants or other solid materials such as desic-

3 Patt ofthe overall program for herbarium pest management.

ACKNOWLEDGMENTS

^ assistant Zachary J. Buning is gratefully acknowledged. The constructive criticism of Wendy B. Zorn-

5g2 Journal of the Botanical Research Institute of Texas 6(2)

lefer (GA), Kent D. Perkins (FLAS), and Melanie Link-Perez (AASU) improved this paper. This work was sup-

ported in part by NSF DBI-1054366 (J R- Carter, PI).

Anonymous. 2012. Australian Entomological Supplies Pty. Ltd. http://www.entosupplies.com.au (18 January 2012).

Hail, A.V. 1988. Pest control in herbaria. Taxon 37:885-907.

Stoang, TJ.K. 1999. A healthy dose of the past: a future direction in herbarium pest control? In: DA. Metsger and S.C.

Byers, eds. Managing the modern herbarium, an interdisciplinary approach. Society for the Preservation of Natural

History Collections (SPNHC). Washington, D.C. Chapter 3:59-80.

NUEVOS REGISTROS DE POACEAS PARA EL NORTE DE MEXICO

Y. Herrera Arrieta 1 , C.A. Silva Salas, L. Ruacho Gonzalez y 0. Rosales Carrillo

Instituto Politicnico Nacional, CIIDIR Unidad Durango-COFAA

Sigma 1 1 9, Fracc. 20 de Noviembre II, Durango, Dgo. 34220, MEXICO

ABSTRACT

& Columbus, is ex

Durante el desarrollo de estudios de biodiversidad (floristica y citologica), cuyo objetivo principal fue realizar

lesisde Maestrfa en el IPN CIIDIR Durango, sobresale el hallazgo de localidades no registradas con anteriori-

para dos especies de gramineas (familia Poaceae), familia reconocida por su importancia ecologica y gran

calidad alimenticia de muchas de sus especies (ejemplo cereales, forrajes, cana de azucar). Se encontro que la

•ocalidad de colectas recientes en la vegetation de cimas del cerro Huehuento, San Dimas, Durango expande el

area de distribucion conocida de Muhlenbergiafiliculmis Vasey (taxon de distribucion boreal conocido de las

nontafias del W de Estados Unidos), resultando ser este e 1 primer registro para Mexico. As! tambien, colectas

recientes en la Laguna de Salitrillo, Zacatecas expanden el area de distribucion de Distichlis eludens (Soderstr.

&H.F. Decker) H.L. Bell & Columbus, taxon escaso y endemico de suelos halofilos en terrenos inundables del

“°ne-centro de Mexico, cuya distribucion conocida se restringia a dos localidades disyuntas: Humedal de

ga, Durango y Salinas de Hidalgo, San Luis Potosi, siendo la nueva localidad un punto intermedio entre las

Baades an, ertormeme conocidal.

METODOS

esludia r°n ejemplares de los taxa incluidos, provenientes de colectas recientes, realizadas en las areas de

n^ribucion registradas en el reporte presente. La determination de la identidad de las especies se llevb a cabo

rzando claves y descripciones de diversas obras floristicas para Norteamerica (Hitchcock & Chase 1951) y

^“nas regiones de Mexico (McVaugh 1983; Herrera 1998; Herrera 2001; Herrera y Pamanes 2006; Herrera y

RESULTADOS

^identifiearon ejemplares con el nombre de Muhlenbergia filiculmis de la vegetacibn de alta montana en el W

range y de Distichlis eludens proveniente de la laguna del Saladillo, Zacatecas. Ambas localidades am-

n a distribucion conocida de estas especies.

** 3ra ^ x * co y P ara e * f stado de Durango

n r g»a filiculmis Vasey es una especie que habita en lugares templados frios de los

1 ** s - Inst Teas 6,2): 583 _ 5g$ j 01 2

bosques de

584

Journal of the Botanical Research Institute of Texas 6(2)

conlferas de las montanas del NW de Estados Unidos, entre los 2500 y 3300 m de altitud, con distribution

conocida en Arizona, California, Colorado, Nuevo Mexico, Utah, Washington y Wyoming (Herrera, 1998). Es

una especie perenne pero de porte pequeno, de 10 a 35 cm de alto, por lo que pudo haber pasado desapercibida

a los ojos de colectores anteriores, sumado a ello la caracteristica de que es apetecida por los herbivorosyse

encuentra generalmente ramoneada. Es una especie del complejo aparentemente monofiletico de Muhlenberg

montana (Nutt.) Hitchc., propuesto por Herrera (1998), cuya caracteristica morfologica principal es tenerla

segunda gluma 3-nervada; siendo ahora reconocido como un linaje natural (subgenero Clomena), con base a

el andlisis filogenetico del ADN (Peterson y col. 2010).

Las colectas de material botanico en la vegetation de alta montana de la Sierra Madre Occidental son es-

casas, especialmente en el municipio de San Dimas, Durango, donde lo inaccesible del terreno hace quesean

pocos los colectores que se internan en sus caminos para realizar estudios floristicos.

La localidad del ejemplar de M.filiculmis [Mexico. Durango. Municipio San Dimas: Cima del cerro Hue-

huento, 24°04’3rN, 105°44’24”W, 3262 m, M. Gonzdlez 4288A (CIIDIR)] es nueva para Mexico y se constituye

como el extremo septentrional de distribution para esta especie, a un limite altitudinal sobre los 3200 m,

misma que sale del limite sur conocido hasta ahora, cerca del paralelo 32° N en Arizona y Nuevo Mexico hasta

cerca de 24° N correspondientes a la nueva localidad en San Dimas, Dgo. Podria sospecharse la existenciade

otras localidades en que posiblemente se encuentre creciendo esta especie, sin embargo dichas localidades

debieran ser en las altas montanas de la Sierra Madre Occidental, a fin de compensar las diferencias de latitud

con altitud que permite se presenten los requerimientos climaticos del habitat de la especie.

Las localidades de alta montana en Mexico son escasas. Challenger (1998) marca 18 picos altos con vegetation

alpina o subalpina, 10 de ellos en el Eje Volcanico Central, 7 en la Sierra Madre Oriental y 1 en la Sierra Madre

Occidental (el cerro Mohinora). Sin embargo en la Sierra Madre Occidental, se encuentran otros picos que

quizes no Uegan a tener vegetacidn alpina tipica, pero que presentan elementos de vegetacion alpina por la alti-

tud que alcanzan, como es el caso de los cerros siguientes: Cerro Mohinora (3,293 m), mpio. Guadalupe y

Calvo, Chih.; Cerro Barajas (3,280 m), mpio. Guanacevi, Dgo.; Cerro Huehuento (3,233 m), mpio. San Dimas,

Dgo., donde se encontrd M. filiculmis para este registro; Cerro Gordo (3,328 m), mpio. Pueblo Nuevo, Dgo.;

Cerro de las Antenas (3,221 m), mpio. Canatlan, Dgo. Ruacho (2011), inicio muestreos y dejo marcas enlas

localidades de algunas de estas montanas (en aquellas donde la seguridad actual lo permitio), con el fin de re-

alizar monitoreo para ver cambios posteriores en la vegetacion de la Sierra Madre Occidental.

II. Nuevo registro para el Estado de Zacatecas

El gdnero Distichlis, fue propuesto por Rafinesque en 1819 (L6pez y col. 2009). Se le conoce vulgarmente como

“pasto salado” ya que habita en suelos naturalmente salitrosos o salinizados artificialmente, desde elevadas

montaiias hasta la orilla del mar, se destaca en la clasificacion sintaxonomica, debido a la relation tan estrecha

que tiene con los ambientes en los que habita (Pelliza y col. 2005). Generalmente, estas plantas permanecenen

estado verde durante el periodo de sequia, al tiempo que el resto de los pastos forrajeros en el sitio no estan

disponibles, por esta razon el ganado lo consume (Estrada-Hemandez y col. 2005).

Un nivel elevado de salinidad en el suelo aumenta el punto de marchitamiento de las plantas como conse-

cuencia del incremento de la presion osmotica y de los efectos toxicos de las sales (Proyecto SoCo 2009). U

salinidad puede llegar a ser un factor importante en la reproduction de las especies de Distichlis, lo que hace

que su estudio sea de singular importancia (Proyecto SoCo 2009).

Distichlis esta formado por un pequeno grupo de plantas herbaceas, dioicas, perennes y rizomatosas, con

espiguillas multifloras y hojas conspicuamente disticas, pertenecen a la familia Poaceae, subfamilia Chloridoi-

deae, tribu Cynodonteae y subtribu Monanthochloineae (Lopez y col. 2009). La taxonomia de este grupo, fue

definida recientemente por Bell y Columbus (2008) con base en caracteres de la topologia nuclear y del cloro-

plasto, adicionales a los caracteres morfolbgicos y anatomicos del grupo. Concluyendo que los generos MoruW-

thochloc y Rcederochloa estan anidados dentro de un Distichlis parafiletico, conformado por unas siete especi*

distribuidas en sitios salinos del Hemisferio Occidental y Australia (Bell y Columbus 2008, Barkworth y BeU

2011), cinco de esas especies se encuentran en Mexico.

Distichlis eludens (Soderstr. & H.F. Decker) H.L. Bell & Columbus (antes Reederochloa eludens Soderstr.

& H.F. Decker) (Fig. 1) es una planta perenne, dioica, estolonifera que se-desarrolla en zonas de alta concen-

tration de sales en pastizales halofilos o en claros de matorral xerofilo y requiere de altos niveles de agua para

su crecimiento. Se encuentra en cualquier epoca del ano aunque este marchita y es por ello que el ganado la

consume. Es una planta que en ocasiones es dificil de encontrar por el tamaiio tan pequeno que presenta y por

elgrado de ramoneo que el ganado ejerce sobre ella, asi tambien en otras ocasiones, se puede Uegar a observar

facilmente ya que la densidad de individuos que forman una colonia puede ser hasta de 10 m continuos. La

especie se consideraba endemica de las dos localidades disyuntas, en los Estados de Durango y San Luis Potosi,

d ejemplar recientemente colectado [Mexico. Zacatecas: orilla SE de la Laguna el Saladillo, 22°38’38.7 , 'N,

102°02'24.9"W, Silva, Carrillo y Herrera 29 (CIID1R)], amplia el area de distribution conocida para este taxbn

alEstado de Zacatecas, region intermedia a las de distribution conocida.

El estudio citologico en poblaciones de D. eludens realizado por Silva (2011), proportion un ideograma

del cariotipo de esta especie, confirma el numero cromosomico (x=9) y la aneuploidla (2n=4x+2=38) reporta-

dos anteriormente, con resultados novedosos adicionales que se reportaran en un artlculo en proceso.

CONCLUSION

U revi si6n y colecta de material botanico con fines de estudios de biodiversidad floristica, citologica, ecologica

0 de otra indole, siguen aportando information novedosa y relevante para el conocimiento de nuestra flora

®exicana. Se dan a conocer a la comunidad botanica localidades nuevas de dos importantes gramlneas que se

rep °rtan por primera vez: Muhlenbergiafiliculmis nuevo registro para Mexico y Distichlis eludens nuevo registro

P® 3 el Estado de Zacatecas.

AGRADEC1MIENTOS

%adecemos a la Secretaria de Investigation y Posgrado del 1PN por el financiamiento recibido en los proyec-

^ S ' ^E-20110017 “Citologia de Gramineas y Ciperaceas de importancia economica del n-c de Mexico y

S1P2 °1 10681 “Fitodiversidad y Ecosistemas de la Sierra Madre Occidental”. Asi tambien agradecemos al

CONACyT por las becas de estudios de posgrado a la 2 a (VU: 331664) y 4° coautores. Por ultimo agradecemos

a J J. Ortiz y un revisor anonimo por las observations y sugerencias al manuscrito del presente reporte.

586

Barkworth, M.E. and H.L. Bell. 201 1 . 1 7.04 Distichlis Raf. http://herbarium.usu.edu/treatments/Distichlis-1

Bell, H.L. and J.T. Columbus. 2008. Proposal for an expanded Distichlis (Poaceae: Chloridoideae): support

morphological, and anatomical characters. Syst. Bot. 33:536-551.

Challenger, A. 1998. Utilizacton y Conservacton de los Ecosistemas Terrestres de Mexico, pasado, presente y futuro.

CONABIO (Comiston Nacional para el Conocimiento y Uso de la Biodiversidad)-UNAM Universidad Autonoma de

M6xico)-Agrupaci6n Sierra Madre, S.C., Mexico.

Estrada-HernAndez, A., E. Troyo-Dieguez., J.L GarcIa-HernAndez, H. HernAndez-Contreras, B. Murillo-Amador, y R. LOpez-Aguiuul

2005. Potencial forrajero del pasto salado Distichlis spicata (L.) Greene en ecosistemas costeros de Baja California Sur,

Mexico porel m&odo de"componentes principales.'Tec. Pecuaria Mexico 43(1):1 3—25.

Herrera A., Y. 1 998. A reviston of the Muhlenbergia montana (Nutt.) Hitchc. complex (Poaceae: Chloridoideae). Brittonia

50:23-50.

Herrera A., Y. 2001 . Las Gramfneas de Durango. Institute Politecnico Nacional (IPN)-CONABIO, Mexico.

Herrera A., Y y D.S. PSmanes G. 2006. Guia de pastes para el ganadero del Estado de Durango. IPN-COCyTED-Fundac6n

Produce, Durango, A.C.

Hitchcock, A.S. y A. Chase. 1951. Manual of the grasses of the United States. 2nd ed. USDA Misc. Publ. 200.

Herrera A., Y„ P.M. Peterson, y A. Cort£s 0. 201 0. Gramfneas de Zacatecas, Mexico. Sida, Bot. Misc. 32:1 -239.

Lopez S., Ma.M., S.D. Koch, M. Flores-Cruz y E.M. Engleman. 2009. Anatomfa comparada de la lamina foliar del genera Disti-

chlis (Poaceae). Acta Bot. Mexicana 89:1-23.

McVaugh, R. 1983. Gramineae. In: Anderson, W.R., ed. Flora Novo-Galiciana Vol 14. University of Michigan Press, Ann

Peluza, A., L. Borrelu y G. Bonvasuto. 2005. El pasto Salado ( Distichlis spp.) en la Patagonia: una forrajera adaptada a la

aridezy a la salinidad. Facultad de Ciencias Agropecuarias. Rev. Ci. Agropecuaria 9(2):1 19-131.

Peterson, P.M., K. Romaschenko, and G. Johnson. 2010. A phytogeny and classification of the Muhlenbergiinae (Poaceae:

Chloridoideae: Cynodonteae) based on plastid and nuclear DNA sequences. Amer. J . Bot. 97:1532-1554.

Proyecto SoCo. 2009. Agricultura sostenible y conservation de los suelos, procesos de degradation del suelo. Ficha

informativa n°4: Salinidad y Sodificacton.

Ruacho G., L 201 1. El elemento alpino en la vegetation de cimas de la Sierra Madre Occidental. Tesis de Maestrfa, IPN

CIIDIR Dgo., Mexico.

&lva S., C.A. 201 1 . Citologfa de especies de gramfneas en el norte-centro de Mexico. Tesis de Maestrfa, IPN CIIDIR Dgo,

distribucion actual y potencial

DE TAXUS GLOBOSA (TAXACEAE) EN MEXICO

Mario A. Garda-Aranda, Cesar Cantu-Ayala, Eduardo Estrada-Castillon,

Marisela Pando-Moreno y Antonio Moreno-Talamantes

Facultadde Ciencias Forestales

Universidad Autdnoma de Nuevo Ledn

Km 145 Carr. Nacional, Apartado Postal 41

C.P. 67700, Unares, N. L, MEXICO

RESUMEN

Clave Taxus globosa, Modelacion, Habitat, MaxEnt

ABSTRACT

NOM-059-SEM ARNAT-20 1 0 , a modeling distribution procedure of maximum entropy or MaxEnt, based on 40 occurrences of field records

REM1B-CONABIO records. Two different scenarios of geographical and environmental coverages— climatic and topographic— were

Proved statistically. The BIOCL1M variables scenario with topography data show a more accurate model in the range of 0.9 to 1.0 probability

roth a 53% coincidence; it was statistically significant at (AUC=0.985). Slope variable indicates an environmental variable with more influ-

““ in potential modeling, followed by Biol4 (Precipitation of Driest Month). In agreement with the MaxEnt analysis with 19B10CLIM and

3 topographic variables, the main plant communities in the modeling area (where Taxus globosa is distributed in Mexico) are pine-oak for-

^osk-pine forest, pine forest, and cloud mesic forest. Taxus globosa is potentially distributed in Hidalgo, Nuevo Leon, Puebla, Tamaulipas,

aodVm

k prediction de la distribution potencial de especies raras o en riesgo de extintion es relevante para llevar a

un se guimiento y conservation, ya que proporciona un punto de partida en practicas de recuperacion de

suscondiciones ambientales o del establecimiento de areas de repoblacion artificial (Leal 2009). Taxus globosa

« una conifera incluida en la NOM-059-SEMARNAT-20I0 en la categorla de sujeta a protection especial

(SEMARNAT 2010). La information de registros de especies raras como Taxus globosa, en bases de datos o

Publicaciones, es escasa, aun lo es mas sobre las areas de distribucion local o regional (Zamudio 1992; Garcia y

^Uo 2000; Contreras y Luna 2001; Zavala 2002; Zavala et al. 2001; Bonilla y Canchola 2008). Existen diver-

** metodos para el modelado de las areas de distribucion potencial (Phillips et al. 2006; Ward 2007; Phillips y

Dud* 2008; Hernandez et al. 2008; Garcia 2008), algunos de los cuales se han aplicado en estudios en plantas

en categoria de riesgo de extintion o de distribucion restringida (Engler et al. 2004), ademas de un estudio

sobre el modelado del nicho de Taxus globosa con analisis de los efectos del cambio en uso de la tierra y

conservation en Mtirico (Contreras etal. 2010).

H proposito del estudio fue modelar el area de distribucion potencial de Taxus globosa y conocer la m-

de enables bioclimaticas y topograficas en la precision del modelado a traves de una comparacion.

i el objetiv

aportar information para planes c

MATERIALES Y MfiTODOS

Se visitaron 40 sitios donde crece Taxus globosa en recorridos de campo ei

Nuevo Leon y Tamaulipas, en el noreste de Mexico, en estos sitios se tom:

cuales fueron incorporadas a una base de datos.

En estos sitios se colectaron ejemplares botanicos Estrada et al, con numeros de colecta 11804, 15993,

16129, 16299, 16373, 16383, 16507, 16545, 16820, 16828, 19155, 19436, 2000 los cuales fueron depositadosen

el herbario CFNL. Una serie de 39 registros (coordenadas geograficas) de Taxus globosa se obtuvieron de la

consulta a la base de datos de la Red Mundial de Information sobre Biodiversidad REMIB, CONABIO, http#

www.conabio.gob.mx/remib/doctos/remib_esp.html. Los 40 registros de campo se sumaron a los 39 registros

de REMIB integrando una base de 79 registros totales utilizados en la modelacion de la distribution potenciaL

Posteriormente del uso de esta base de datos, se utilizaron 42 registros de colecta adicionales obtenidos de visi-

tas de campo posteriores con el propbsito de realizar pruebas de validation de los modelos por porcentaje de

Del sitio web de WorldClim (Hijmans et al. 2005; http://www.worldclim.org/bioclim) se obtuvieron cu-

biertas geograficas de variables ambientales de 1960-1990 en formato raster de un km 2 de resolution. La base

de datos BIOCL1M utilizada en el modelado de distribution consta de 19 cubiertas (Cuadro 1), la cual fue

combinada en una segunda modelacion con un grupo de tres variables topograficas altitud, exposition y pen-

diente (Cuadro 1) que fueron calculadas a partir de la cubierta ‘altitude’ de WorldClim (Hijmans et al. 2005;

http://www.worldclim.org/bioclim), todas estas fueron recortadas en ArcGIS version 9.2 a los limites de

118°20'3.42 n a 86°25'3.42" Oeste y 34° 01'0.08" a 12° 5530.76" Norte, correspondiente al territorio de Mexico;

las capas se transformaron en ArcGIS version 9.2 a formato ASCII Grid.

Se utilizb el modelado Maximum Entropy Distribution o MaxEnt, el cual es el mejor metodo (Kumar y

Stohlgren 2009), es de los que procesan bases de datos con pocos registros (Hernandez et al. 2006; Pearson et

al. 2007). Es un mtiodo basado en un protocolo que estima la probabilidad de ocurrencia de las especies basa-

do en requerimientos ambientales que genera una estimation de probabilidad de presencia de la especie am

valores de 0 y 1, donde 0 se considera como la minima y 1 para la maxima probabilidad (Phillips et al. 2006),

para realizar el analisis solo se requiere de datos de presencia de la especie (no ausencia) y cubiertas geograficas

de variables ambientales de las areas de analisis (continuas o categoricas). El software MaxEnt versibn 3.3.2 es

de uso libre y esti disponible enhttp://www.cs.princeton.edu/~schapire/maxent/.

La base de datos compilada de 79 registros de Taxus globosa y los cuatro grupos de variables ambientales

fueron utilizadas para la modelacion de un escenario climatico actual (periodo de 1950 a 2000). El programs

generb intigenes logisticas acumulativas y analizo a un maximo de 500 aproximaciones sucesivas (iteraciones).

Los dos modelos generados se sometieron a las pruebas curva de respuesta (analisis de omision/comision

y sensitividad ROC AUC=Area Under Curve) y una prueba Jackknife para medir el efecto de cada variable u0-

lizada en la generacibn de los modelos. Para realizar esto se dividio la base de datos de 79 registros en un

subgrupo de 40 registros para el modelado de habitat potential y otro de 39 para pruebas estadisticas de vaK-

Con una base de 42 registros adicionales de Taxus globosa, a traves del modulo Point Analyst 10 pan

Arcview, se registrb el valor de los dos modelos de distribution potential generados en formato raster los cualti

presentan valores de probabilidad de habitat de 0.1 a 1.0, de manera que solo se contabilizaron los registroscon

valores entre 0.8 y 1 .0 para expresarlos en porcentaje de ocurrencias >80 % de probabilidad de tener un habitat

potential para Taxus globosa.

Se realizb un analisis de distributibn de la superficie actual y potential (entre los pixeles de 0.8 a 1.0 pR>*

babilidad) generado en el mejor modelo de acuerdo al uso actual de la tierra y vegetation (INEGI 2005). A*

mismo, se determino su area de distributibn actual y potential en Mexico.

Garcia et al.. Distribution de Taxus globosa en Mexico

0*«»1.Ustadodelasvariablesambientales BIOCLIM 2 y topografi

589

RESULTADOS Y DISCUSlON

de distribution potencial para Taxus globosa en Mexico se muestra en la Figura 1 el cual considera

^ 19 cubiertas geograficas BIOCLIM sin la de topografia. Este mapa tiene un alto nivel de prediccion. El

del area bajo la curva ROC (AUC = 0.987) (Cuadro 2). En contraste el mapa del modelo de habitat poten-

Pwa T. globosa considerando las 19 cubiertas geograficas BIOCLIM y las tres cubiertas geograficas de topo-

2 X genero una tasa de exito mayor en su prediccion con un valor de area bajo la curva ROC (AUC -

M , ) (Cuadr ° 2), estos valores son menores al obtenido por el modelo (AUC = 0.997) generado por Contreras

tt *l (2010).

ilidacion de MaxEnt (Cuadro 2), los dos modelos generados tuv

Rirv-T Vaiiables 9 ue tuvieron mayor ponderacion en la generacion de los modelos, fueron: para el modelo

tem Peratura maxima del mes mas calido (Bio05) 39.4%, isotermalidad [P2A>7] [*100] (Bio03)

’ Pecipitacion del mes mas seco (Biol4) 15.3% y temperatura minima del mes mas frio (Bio06) 10.3%;

590

Journal of the Botanical Research Institute of Texas 6(2)

las de mayor probabilldad.

para el modelo BiOCLIM con topografla: Pendiente (Topo03) 39.8%, precipitacion del mes mis seco (Biol4)

25.5%, elevacibn (TopoOl) 14.0% y temperatura minima del mes mas frio (Bio06) 7,5%, muestra que la pen*

diente del terreno tiene relevancia en uno de los dos modelados, seguida de la precipitacion en el mes mas seco

y elevacibn. Esto significa que el segundo modelo produjo areas mas especificas que discriminan zonas p®

topografla (Fig. 3, Cuadro 1),

Los patrones de distribucion biogeografica estan cambiando en respuesta a recientes alteraciones del cfr

ma, como lo establece un indice que mide la velocidad de cambio de la temperatura (km afio) y la velocidad

cambio varia en los ecosistemas (Loarie et aL 2009). En relacibn a la influencia de la topografla, dicho tadis

establece que el efecto topografko, influye de manera importante en la velocidad de cambio de la temper®" 1 *’

siendo menor (0.11 km ano) en el ecosistema con bosques de coniferas tropicales y subtropicales, dondest

distribuyen las comunidades vegetales con presencia de Taxus globosa. Ademas, el tiempo de residencia Ok®"

po que permanece un ecosistema en una regibn) de este es mayor (63.6 anos) respecto a otros ecosistemas

analizados (Loraie et al. 2009).

La Figura 4 muestra la distribucion de los registros usados en el modelado de habitat potencial con la '^

liable Topo03 (pendiente), donde se observa que los datos se concentran entre los 89.99 y los 90.00 grades

pendiente, lo cual indica que la especie prefiere crecer en los canones. En la Figura 5 se muestra la distribuc*®

de los registros en la variable Biol4 (precipitacion del mes mas seco), los datos se concentran entre 8.0 y 32®*®

de precipitacibn (mes mSs seco del ano), lo que indica zonas con periodos de sequla. La Figura 6, muesttf

distribucibn de los registros para la variable Topo02 (elevacibn), en la que se observa una preferencia por

aones etitre 1,200 y 3,000 m, ubicando la condition del habitat en zonas elevadas. En la Figura 7, se muestra la

nbucidn de los registros en la variable Bio06 (temperatura minima del mes mds frlo), los datos se ubican

entre 24 y 33 °C, lo que denota una preferencia por sitios templados,

En el analisis de distribucion por deciles de probabilidad (entre 0.1 y 1.0) de presencia de distribution

P^ial de Toxus globosa en Mexico para lo dos modelos, se observo que los uitimos dos deciles (0.8-0.9 y

1 °) re P re sentan las areas donde sc advierte mayor probabilidad de encontrar a la especie y coincide con la

20113 nticleo del area potential de distribucion de la especie (Figs. 1 y 2).

Earaelpcnultimodecilde probabilidad (0.8-0.9) la superficie calculada para la distribution potencial fue

en el modelo BlOCLIM con 9,299.38 km 2 que representa un 0.47% del territorio de Mexico, mientras

qUCenel mod elo Biocilm+Topografia genero una superficie de 8,581 km 2 que representan el 0.43% del territo-

Porelc.

**** nna suj

^CLIM+Topografia genero una superficie de 1,804 km 2 (0.09% del territorio mexicano). Es probable que la

cam CnCia Cn tendencia Sca resultado del efecto de la pendiente y elevaciOn ya que los registros tornados en

Po estan ubicados en zonas de elevation mayores a 1,200-3,800 m y con pendiente pronunciada (Fig. 4).

1 grupo de 42 registros adicionales de Taxus globosa se realizO una prueba de validation de

s generados, contabilizando los registros que coinridieran geograficamente con las zonas de los

e marcaron valores de probabilidad entre los deciles 0.8-0.9 y 0.9-1.0, estos se expresaron en valo-

Journal of the Botanical Research Institute of Texas 6(2)

ft. s. Mapa de la distnbucidn de 79 registros de Taxus globosa en reladdn con la variable ambiental Bio14 (precipitation promedio del mes mis seco).

res porcentuales de coincidencia con los modelos. Para el modelo BIOCLIM se obtuvo una precision general de

52% en el deed 0.8-0.9 y 2% en el deed 0.9-1 .0) mientras que para el modelo BIOCLIM+Topografia se

obtuvo una precision de 55% (36% en el decil 0.8-0.9 y 19% en el deed 0.9-1.0), Considerando el total de valo-

probabilidad de los pixeles, el modelo BIOCLIM tiene mayor precision con 83%, mientras que si consi-

^ ran,os Flores del decil de mayor probabilidad de ocurrencia de habitat potencial para Taxus globosa, el mo-

*° B I0CLIM+ Topografia tiene la mayor precision con 55% (Cuadro 4).

U P meba muestra que el modelo de BIOCLIM+Topografia tiene una mayor eficiencia en el Ultimo percen-

" de ma r or probabilidad de ubicar Taxus globosa de 0.9-1.0, mientras que el modelo BIOCLIM es mA s preciso

* wpercentil de menor probabilidad de ubicacion de Taxus globosa 0.8-0.9 (Fig. 8), en contraparte Contreras

C,a ' ^ 01 °) obtuvieron un 94 5% de eficacia en la prediction para su modelo; esta diferencia se establece ya que

^subdividio el modelo en deciles y fue considerada la totalidad de la superficie del modelo, la eficiencia de

“todelos considerando las coincidencias de los registros de la prueba de eficiencia son del 90% para el mo-

un ° y 81 % p ara m0{ j e l 0 2 ( a partir de una probabilidad del 0.6 al 1.0.

t ^ fealizo un analisis de la superficie del habitat potencial (0.8-1.0) para T. globosa, de acuerdo al uso de la

y Ve g e taci6n (INEGI 2005), al cruzarla information se analizO la distribution de los 10,385 km 2 en rela-

° n a 'os tipos de vegetation y uso actual de la tierra. De este analisis se obtuvo que la comunidad con mas

P^dicie del area de distribution potencial es el bosque de pino-encino con 2,230 km 2 , (21.48%), el bosque de

"O-Pino eon 1,638.5 km 2 (15.78%), el bosque de pino con 1,244.75 km 2 (11.99%). Existe un Area abierta a la

594

agriculture de temporal de 1,031.53 km 2 (9,93%), el bosque mesofilo de montana con 1,027.12 km 2 (9.89%), d

bosque de encino con 10,016 km 2 (9.78%), ver Cuadro 5.

En un analisis de distribucion posterior de la superficie generada de habitat potencial (0.8-1.0) para I

globosa de acuerdo al modelo BIOCLIM+Topografia, los estados con mayor superficie potencial son: Hidalgo,

Nuevo Leon, Veracruz, Puebla y Tamaulipas. En el percentil de mayor probabilidad (0.9-1.0) los estados de

Veracruz (474,5 km 2 ), Puebla (401.3 km 2 ), Hidalgo (329.9 km 2 ) y Nuevo Leon (310.1 km 2 ) son los que tienen

mayor superficie. En el percentil medio (0.8-0.9) los estados de Hidalgo (1,935.95 km 2 ), Nuevo LeOn (1,775

km 2 ), Veracruz (1,342 km 2 ), Puebla (1,258 km 2 ) y Tamaulipas (1,173 km 2 ) son los que tienen mas superficie

(Cuadro 6).

Los modelos desarrollados en MaxEnt para obtener distribucion potencial para Taxus globosa en Mexico, advi-

erten un area muy reducida, con una superficie entre los 9,650 km 2 (BIOCLIM) y los 10,385 km

(BIOCLIM+Topografia) que representan entre el 0.49% al 0.52% del territorio nacional.

El modelo BIOCLIM (19 variables climaticas) resulta tener el mejor porcentaje de precision global enel

percentil de probabilidad (0.8-0.9) de presencia de habitat potencial para Taxus globosa en Mexico.

Para el percentil de mayor probabilidad de presencia de la especie (0.9-1 .0), el modelo BIOCLIM+Topogr 3 " 2

resulto ser mas preciso con el 55% en la prueba de validacion. La variable de pendiente es de mayor influence

en el modelado de habitat potencial, seguida por Biol4 (precipitation del mes mas seco).

II /IIIIIIIII! I

Garcia et al.. Distribution deTaxus globosa en Mexico

595

1,843,243.20

56,857.47

23.614.17

13.427.17

10,439.67

8,558.69

8,093.88

1,828,028.46

62,255.25

27,856.23

16,507.72

It, 496.47

9,701.42

La ubicacion del habitat potencial de Taxus globosa en zonas de montana correspondientes al bosque de

atferas " ’ • 1 im bio de temperatura por efecto global, asf como una

[iempo de residencia de este habitat

L °S tipos de vegetation que

el habitat potencial de los ultimos dos percentiles de probabilidad

de encino-pino, bosque de pino y bosque mesofilo de montafla,

Garda etal., Distribution de Taxus globosa en Mexico

597

Gmm» 5. Distribution de la superfide generada de habitat potential para Taxus globosa (0.8-1 .0) con base en ei mapa uso del suelo y vegetation (INEGI, 2005).

Gmdro6. Superficies (km 2 ) de habitat potential para Taxus globosa (km 2 ) de los principles estados en Mexico en los percentiles 0.8-0.9 y 0.9-1 .0 de probabilidad

scgun el modelado BIOCLIM+Topografia.

odstiendo para el 2005 una area abierta a la agricultura de 1,027 km 2 que representa casi un 10% de este habi-

tat potencial en Mexico.

De acuerdo al analisis MaxEnt utilizando las 19 cubiertas geograficas BIOCLIM y tres topograficas, los

estados de Hidalgo, Nuevo Leon, Veracruz, Puebla y Tamaulipas poseen mayor superficie donde se puede dis-

tnbuir Taxus globosa en Mexico.

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DISTRIBUTION AND MORPHOLOGICAL CHARACTERISTICS OF

ARCEUTHOBIUM HONDURENSE AND A. NIGRUM (VISCACEAE) IN MEXICO

Robert L. Mathiasen

School of Forestry

Northern Arizona University

Flagstaff, Arizona 86011 U.S.A.

Robert.Mathiasen@nau.edu

Shawn C. Kenaley

Department of Plant Pathology

and Plant-Microbe Biology

Cornell University

Ithaca, New York 14853 U.S.A.

sck26@cornell.edu

Brian P. Reif

School of Forestry

Northern Arizona University

Flagstaff, Arizona 860 1 1 U.S.A.

Brian.Reif@nau.edu

ABSTRACT

RESUMEN

Tfe genus Arceuthobium (Santalales: Viscaceae) consists of 42 species (Hawksworth & Wiens 1996) that are

aenal parasites of Pinaceae or Cupressaceae. Many of the species, commonly known as dwarf mistletoes, are

cognized as serious forest pathogens (Hawksworth & Wiens 1996; Mathiasen et al. 2008). Morphological

characters consistent throughout the genus include small flowers produced on male and female plants, leaves

reduced to squamate scales, and morphologically similar bi-colored fruits (Hawksworth & Wiens 1996). The

8enus ^ long been considered a taxonomically difficult group because of the extreme morphological reduc-

’'° n dialed with the parasitic habit and the morphological similarities between species (Hawksworth &

lens 1996 )- Factors that complicate classification and identification include a large amount of variation in

morphology and geographic distributions as well as flowering periods that occasionally overlap. Two species

' 1 exem Phfy the problems associated with field identification of morphologically similar dwarf mistletoes

^ in southern and central Mexico: Arceuthobium hondurense Hawksw. & Wiens (Honduran dwarf mistle-

106 3ndA - "igrum Hawksw. & Wiens (black dwarf mistletoe).

Journal of the Botanical Research Institute of Texas 6(2)

Arceuihobium hondurense was originally described from central Honduras (Hawksworth & Wiens 1970)

and thought to be extremely rare, even on the verge of extinction due to rapid and extensive harvesting of its

pine hosts (Hawksworth & Wiens 1972). However, Honduran dwarf mistletoe was later found to occur from

northern Nicaragua, through much of Honduras into Chiapas, Mexico and as far north as central Oaxaca (Ma-

thiasen et al. 2001; Mathiasen et al. 2002a; Mathiasen et al. 2003; Mathiasen et al. 2006; Mathiasen & Melgar

Wiens 1977; Mathiasen et al. 2003). Another dwarf mistletoe, originally described as A. hawfesworthii Wiens

and C.G. Shaw (Hawksworth’s dwarf mistletoe), was recombined as a subspecies of A. hondurense (Mathiasen

2007). Therefore, A. hondurense presently consists of two subspecies: A. hondurense subsp. hondurense which

occurs from Nicaragua north to Oaxaca, Mexico (Mathiasen et al. 2010) and A. hondurense subsp. hawksworthu

(Wiens & C.G. Shaw) Mathiasen which is primarily distributed in the Mountain Pine Ridge area of Belize

(Mathiasen 2007), but also has been reported from central Honduras (Mathiasen et al. 2002b).

Because plants of Arceuthobium hondurense subsp. hondurense (hereafter referred to as A. hondurense) art

similar in size and color to those of A. nigrum, determining the geographic distribution of these species in

southern Mexico has been difficult (Hawksworth & Wiens 1989, 1996; Mathiasen et al. 2001, 2002a, 2003,

2010). Although A. nigrum was thought to be distributed from northern Durango, Mexico into southern Mexi-

co (Hawksworth & Wiens 1996), it is now thought to be distributed only as far south as central Mexico (Ma-

thiasen et al. 2010). Further, both A. hondurense and A. nigrum produce red flowers that bloom in the fall. We

have collected additional morphological data for A. hondurense and A. nigrum since 1998. Here we report our

findings and discuss the distribution of these dwarf mistletoes in Mexico based on our field observations and

morphological measurements. Because prior studies (Mathiasen et al. 2003; Nickrent et al. 2004) have success-

fully used ribosomal DNA (rDNA) sequence information to discriminate between A. hondurense and A ni-

grum, we conducted additional analyses of the internal transcribed spacer (ITS) region for several populations

of both species, particularly populations in central Mexico where these species may be sympatric (Nickrent et

al. 2004). In addition, because A. vaginatum (Willd.) Presl subsp. vaginatum (Mexican dwarf mistletoe) is also

morphologically similar to A. hondurense as well as A. nigrum, and often confused with them, we have provided

information on how to distinguish these species from A. vaginatum. The primary objective of this study, how-

ever, was to provide additional data on how to discriminate A. hondurense from A. nigrum — and vice versa-

and in so doing, better determine their geographic and host ranges. Morphological data and ITS sequences for

A. vaginatum were taken from Hawksworth and Wiens (1996) and obtained from GenBank, respectively.

MATERIALS AND METHODS

Morphology and Phenology

To compare morphological characters we sampled 16 populations of Arceuthobium hondurense (two fro®

Oaxaca, Mexico, one from Nicaragua, and 13 from previous work by Mathiasen (2007)) and 14 populations

A. nigrum from throughout its geographic range (Fig. 1). Plants were measured from the type locality for both

mistletoe species (Hawksworth & Wiens 1965, 1970, 1977) (Fig. 1; locations 6 and 23). From each population,

10-20 male and 10-20 female plants were collected and the dominant shoot from each infection was used for

morphological measurements. Characters measured were those used by Hawksworth and Wiens (1996) far

taxonomic classification of Arceuthobium: height, basal diameter, third internode length and width, and color

of male and female plants; mature fruit length, width, and color; seed length, width and color; length and width

of staminate spikes; staminate flower diameters for 3- and 4-merous flowers; length and width of stamina*

flower petals; and, anther diameter and anther distance from the petal tip. Plants were measured within 2

hours after collection using a digital caliper and a Bausch and Lomb 7X hand lens equipped with a micromeW

Staminate spike and flower measurements were made during the peak of anthesis and fruit and seed measure-

ments were made during the peak of seed dispersal. One-way analysis of variance (ANOVA) was used to exam-

ine the variance in the above characters for A. hondurense and A. nigrum and significant differences between

means were determined using a Tukey’s honestly significant difference (HSD) post-hoc test (a = 0.05). All s»

tistical analyses were performed using JMP 8.0.2 software (SAS Institute, Cary, NC).

601

Because the times of flowering and seed dispersal for Arceuthobium hondurense and A. nigrum are poorly

known (Hawksworth & Wiens 1996), additional observations of the phenology of these taxa were made dur-

ing the spring and fall of 1999, 2003, 2005, 2007, 2008, and 2010 as well as during the early spring of 2011.

DNA Extraction and ITS Sequencing

^nples of DNA were obtained from five and six specimens, each representing a geographically separate popu-

!ati °n, of Arceuthobium hondurense and A. nigrum , respectively. Locality and voucher number for each speci-

"•en (bold print) are presented in Fig. 1. For each specimen, total DNA was extracted using the DNeasy™ Plant

MlruKit (Qiagen, Valencia, CA) according to the manufacturer’s instructions. DNA purity and concentration

Were quantified for each sample using a NanoDrop ND-1000 (Thermo Fischer Scientific, Wilmington, DE).

^'kngth, ITS sequences (comprising ITS1, 5.8S rDNA gene, and ITS2) were PCR-amplified using the primer

P® 1 IBS 1830for and 26S 40rev (Nickrent et al. 2004). PCR amplifications were carried out in 25 pL reaction

"jhuures containing 12.5 pL of 2X AmpliTaq Gold® Master Mix (Applied Biosystems, Foster City, CA), 0.5 pL

ofea ch20pM primer, 11.25 pL nuclease-free water, and -2-18 ng (0.25 pL of 8-78 ng/pL) of genomic DNA.

602

PCRs were performed in an Eppendorf Mastercycler® pro thermal cycler (Eppendorf, Westbury, NY) with the

following cycling parameters: initial hold for 6 min. at 95°C; 5 cycles at 94°C for 30s, 55°C for 30s, and 72°C for

1 min.; 33 cycles at 94°C for 30s, 48°C for 30s, and 72°C for 1 min.; and, a final extension step of 72°C for 10 min.

Blank reactions (i.e., minus genomic DNA) were run concomitantly to check for contamination of the reagents.

The size of each PCR product (bp) was checked separately by ultraviolet fluorescence after 1.2% agarose

gel electrophoresis in 0.5x TAE buffer and staining with GelRed™ (Phenix Research Products, Candler, NQ.

Amplification products were purified directly from reactions using ExoSAP-lT (0.4 pL per pL of reaction prod-

uct; USB Inc., Cleveland, OH) and normalized to 130 ng per sequencing reaction. Sequencing was carried out

using a BigDye terminators DNA sequencing kit (Applied Biosystems), ABI 3730 DNA sequencer, and the

above forward and reverse primers. PCR products were sequenced in both directions. Sequences were proof-

read and assembled in CodonCode Aligner (CodonCode Corporation, Dedham, MA). Boundaries to the 5-

and 3'-region of ITS1 and ITS2, respectively, were previously identified by Nickrent et al. (1994). ITS sequences

for A. hondurense (n=5) and A. nigrum (n=6) produced in this study were deposited in GenBank.

Phylogenetic Analysis

ITS sequences for Arceuthobium hondurense and A. nigrum obtained in this study and from GenBank (A. hondu-

rense AY2888263 and A. nigrum AY288271) as well as A. vaginatum subsp. vaginatum (AY288286 and

AY288287) and A. douglasii Engelmann (L25687; outgroup) were included in the dataset. Sequences were

aligned using ClustalX ver. 2 (Larkin et al. 2007) and visually edited as necessary in CodonCodon Aligner.

Maximum Likelihood (ML) trees were constructed using PAUP* 4.0bl0 (Swofford 2003). The DNA substitu-

tion model TIM2 and the parameter estimates for tree reconstruction were determined using the Akaike Infor-

mation Criterion (AIC; Akaike 1974) as implemented in jModelTest 0.1.1 (Posada 2008). All nucleotides were

included in the phylogenetic analysis; gaps were treated as missing characters. Heuristic searches were per-

formed with 200 replicates of random sequence addition and tree bisection-reconnection (TBR) branch swap-

ping. Branch support was evaluated using 1000 bootstrap replicates and 10 random additions of sequences per

pseudo-replicate. Inter- and intraspecific genetic distances were also examined using Kimura’s two-parameter

model (K2P; Kimura 1980) for base substitution as implemented in PAUP*.

Bayesian analysis was also performed using MrBayes 3.1.2 (Huelsenbeck & Ronquist 2001). The best-fit

model for DNA substitution was determined as described previously; however, Hasegawa-Kishino-Yano (HKY;

Hasegawa et al. 1985) model and parameter estimates were determined using the Bayesian Information Crite-

rion (BIC; Schwarz 1978). One cold and three heated Markov chain(s) were run, and samples were taken every

100 generations over 5.0 x 10 6 generations. The potential scale reduction factor (PSRF) for each of the model

parameters was > 1.0 when the program was terminated. Stationarity was accessed by examining the average

standard deviations of split frequencies and likelihood values. Bum-in value (10%) was determined using

Tracer vl.5 (Rambaut & Drummond 2009). The r emaining trees were used to calculate a 50% majority rule

consensus tree and to determine the posterior probabilities.

RESULTS AND DISCUSSION

Our measurements of Arceuthobium hondurense indicate it forms larger plants than previously reported by

Hawksworth and Wiens (1970, 1996); they reported plant heights averaged approximately 14 cm with a ma»-

mum height of 21 cm, but we measured plants (male and female combined) that averaged 22 cm. We found

some male plants in Chiapas, Mexico that were over 65 cm in height (Table 1). The discrepancy in maximum

heights is probably related to Hawksworth and Wiens only measuring specimens from central Honduras

where plants were generally smaller than in southern Mexico (Mathiasen et al. 1999; Mathiasen 2007). A*

mean basal diameter of dominant shoots was the same as reported by Hawksworth and Wiens (1996); ap

proximately 5 mm. However, we measured some shoots with basal diameters of nearly 13 mm, while Hawk'

sworth and Wiens (1996) only reported a maximum of 9 mm for this character. The means and ranges for m^

of the remaining morphological characters were similar to those previously reported (Hawksworth & W*® 5

Previously, Mathiasen et al. (2003) listed the pines parasitized by Arceuthobium hondurense in Central

America and Chiapas, Mexico. Our observations support their findings, except that A. hondurense also parasit-

ized Pirns teocote Schiede ex Schlechtendal & Chamisso in central Oaxaca. Although we have only observed

A. hondurense infecting this pine in one location north of Ixtlan (Fig. 1, location 13), the level of infection (>

90%) indicated Pinus teocote was a principal host. This population was misidentified as A. vaginatum subsp.

vaginatum by Hawksworth and Wiens (1996), as our morphological and ITS analyses indicated this population

was indeed A. hondurense. Mistletoe plants on P. teocote at this location were dark brown to black, similar in

color to A. vaginatum, but male plants flowered in the fall producing dark red flowers. Large male plants at this

locality also had swollen nodes and parasitized P tecunumanii Eguiluz et J. P. Perry, a principal host of A. hon-

durense elsewhere in southern Mexico (Mathiasen et ai. 2003). In addition, A. hondurense was found parasitiz-

ing P. lawsonii Roezl ex Gordon & Glendining in central and northern Oaxaca. Although Hawksworth and

Wiens (1977, 1996) reported A. hondurense (but identified as A. nigrum ) parasitizing Pinus oaxacana Mirovand

P. patula Schiede ex Schlechtendal & Chamisso in Chiapas, we did not observe these host-mistletoe combina-

tions during our field work in southern Mexico.

Arceuthobium nigrum is morphologically very similar to A. hondurense. Both species produced relatively large,

dark brown to black plants on their pine hosts (Hawksworth & Wiens 1996). Male and female plants of A ni-

grum averaged nearly 25 and 18 cm in height, respectively, but were not significantly larger than A. hondurense

(Table 1). It is difficult to compare our results for plant heights with those of Hawksworth and Wiens (1996)

because they only provided a range of heights for A. nigrum (15-35 cm, maximum = 45 cm). However, the larg-

est plant we measured for A. nigrum was a male over 53 cm tall from Puebla, Mexico. We also found that the

basal diameter of shoots averaged two-mm larger (7.6 mm) than that previously reported (5 mm, Hawksworth

& Wiens 1996) with a maximum basal diameter nearly twice that described by Hawksworth and Wiens. Mea-

surements of the third intemode widths also indicated that A. nigrum produced thicker plants (about 5 mm)

than what Hawksworth and Wiens reported (about 4 mm). The means for the basal diameters of male (7 mm)

and female plants (7.8 mm) and third intemode widths of A. nigrum were significantly greater than those for A

hondurense (Table 1). Furthermore, the mean length of the third internode of female plants of A. nigrum (16.5

mm) was significantly longer than those of A. hondurense (13.8 mm) but not the mean length of the third inter-

node of male plants.

The only flower characteristics Hawksworth and Wiens reported for Arceuthobium nigrum was the diam-

eter of 3-merous flowers (3.5 mm), which was sUghtly larger than the mean diameter for the 3-merous flowers

we measured (3.2 mm). Our observations indicated that A. nigrum commonly produced 4-merous flowers also,

thus, we sampled these flowers and found they averaged nearly 5 mm in diameter (Table 1). Collectively, the

mean diameters of 3 and 4-merous flowers of A. nigrum were significantly larger than those of A. hondurense.

Petal sizes were also relatively large for A. nigrum when compared to other dwarf mistletoes (Hawksworth &

Wiens 1996). We found petals longer than 2 mm and nearly as wide, both significantly larger than those for A

hondurense (Table 1). Another key characteristic of A. nigrum flowers that was consistent throughout its geo-

graphic range, was that the adaxial surface of its petals was dark red. While this characteristic was easily ob-

served, and is similar to flowers for A. hondurense, a review of the literature (Hawksworth & Wiens 1989, 1996)

on A. nigrum revealed that the petal color, as a diagnostic character, had never been mentioned before,

The mean f

fruits 8.8 rr

of fruits we

Arceuthobium nigrum were remarkably glaucous and large compared to other dwarf mistletoes

it length was nearly 7 mm, which is what Hawksworth and Wiens reported. However, we found

in length compared to 9.0 mm by Hawksworth and Wiens (1996). In contrast, the average width

amined was larger (4.1 mm) than that reported by Hawksworth and Wiens (3.5 mm). We

lat were shorter, but wider on average than those examined by Hawksworth and Wiens (1996)

• fruit length and width of A. nigrum were significantly larger than those of A. hondurense, ho*

i length and width of seeds were similar for both species (Table 1).

605

The phenology of A. nigrum requires additional observations as we were unable to confirm the incidence

of two flowering periods — one in March-April and one in September-October— as reported previously by

Hawksworth and Wiens (1989, 1996). We examined male plants of A. nigrum at several localities in mid- to

late-March during 2003, 2005, and 2007 as well as in early April 2011 and never observed open flowers. Fur-

thermore, staminate flowers did not appear to be approaching anthesis in late March or early April. Our field

observations, however, indicated that it flowers beginning in mid-September and continued into November in

Durango, Mexico, while in central Mexico, A. nigrum flowers in late-September and continued into at least

January. Peak flowering was in early October in Durango; however, the peak flowering period in central Mex-

ico is still poorly understood. Seed dispersal initiated in early September and peaked in mid-October continu-

ing to mid-November in Durango and elsewhere in central Mexico.

Our field observations of pines infected by Arceuthobium nigrum in Mexico did not reveal any additional

hosts. The principal hosts of A. nigrum in Durango were clearly Pinus leiophylla Schiede ex Schlechtendal &

Chamisso, P. lumholtzii B. L. Robinson & Femald, P. teocote, and P chihuahuana Engelmann as reported by

Hawksworth and Wiens (1996). This mistletoe also been reported to rarely infect P. arizonica Engelmann and

P. cooperi Blanco in northern Mexico (Hawksworth & Wiens 1996), but we have not observed it on these infre-

quent hosts. In central Mexico its principal host was P teocote. Pinus patula Schlechtendal & Chamisso was a

secondary host at several locations in Hidalgo and Puebla. While we agree with the classification of P. pseu-

dostrobus Lindley as an occasional host of A. nigrum (Hawksworth & Wiens 1996), we were unable to verify

whether P montezumae A. B. Lambert was also an occasional host. Moreover, we did not observe any P mont-

ezumae at the location where Hawksworth and Wiens reported an infestation of A. nigrum in Hidalgo nor in

Veracruz where we found large, mistletoe-free P. montezumae growing near P teocote severely-infected with A.

nigrum. Therefore, the susceptibility of P. montezumae to A. nigrum needs further study. Although Hawksworth

and Wiens (1989, 1996) reported that both P. lawsonn and P. oaxacana Mirov were principal hosts of A. nigrum ,

this host susceptibility classification was based on infection of these pines by A. hondurense in Oaxaca and

Chiapas, respectively (Mathiasen et al. 2003).

DNA Analyses

DNA sequence analysis demonstrated that Arceuthobium hondurense, A. nigrum, and A. vaginatum occurred in

three well supported clades (Fig. 2). All samples identified morphologically as A. hondurense and A. nigrum

yielded a 627 and 623 bp fragment, respectively, consisting of the 3'end of the 18S (4 bp), complete ITS1-5.8S-

HS2 sequence (604 and 600 bp), and the 5'end of the 26S (19 bp). Four of five sequences for A. hondurense were

identical (mean K2P value= 0.0017); however, RLM 98107 differed by an A/G nucleotide change at positions 22

and 40 in 1TS1. Similarly, ITS sequences for A. nigrum were nearly identical (mean K2P value= 0.0018); except

for A/T nucleotide changes at position 508 in ITS2. The alignment for phylogenetic analyses consisted of 649

characters including those of A. douglasii and A. vaginatum. Of these characters, 574 were constant, 50 were

tree supported three distinct clades (Fig. 2) with bootstrap values >98% and posterior probability values equal

to 1.00, respectively. Each plant identified according to morphometric data as A. hondurense or A. nigrum

formed a distinct clade with either A. hondurense (RLM 0136, Nickrent et al. 2004) or A. nigrum (DLN 2019,

Nic krent et al. 2004). Sequences of A. nigrum differed from those of A. hondurense and A. vaginatum by ap-

proximately 43 nucleotides (mean nucleotide difference = 43.07, mean K2P value= 0.0775). Likewise, the mean

number of nucleotide changes between A. hondurense and A. vaginatum was 13.0 (mean K2P value= 0.0224).

Nickrent etal. (2004) reported thatA. hondurensemay occur in Veracruz based on molecular data (GenBank

accession no. L25693; voucher DLN 2018), but our results did not support this. Plants collected south of Sierra

de A A Ua (RLM 1083), the approximate location where Nickrent collected DLN 2018 (D. Nickrent, pers. comm.),

Were morphologically similar to those of A. nigrum. As suspected, the ITS sequences generated from RLM 1083

identical to L25693 (data not shown). However, in a separate phylogenetic analysis (data not shown),

the se collections/sequences were unrelated to A. nigrum (AY288271) as well as GenBank accessions of A. du-

r ®«go«eHawksw. & Wiens, A. gUlu Hawksw. & Wiens, A. hondurense, and A. vaginatum. Nickrent previously

A. nigrum in Mexico

607

identified DLN 2018 (L25693) as either A. vaginatum (Nickrent et al. 1994) or A. hondurense (Nickrent et al.

2004). The species identity of this mistletoe (RLM 1083 and DLN 2018), therefore, remains unresolved and

requires further study.

SUMMARY

Although Arceuthobium hondurense, A. nigrum, and A. vaginatum are morphologically similar and often diffi-

cult to distinguish from each other in situ, there are several diagnostic characteristics that can be used to iden-

tify t h e m in central Mexico where they may be sympatric. Our results support the classification of these taxa

as distinct species and the principal morphological and physiological characters that can be used to distin-

guish these species are summarized in Table 2. While the overall height of male and female plants and their

color cannot be used to easily separate these species, A. hondurense is a more slender plant than both A. nigrum

and A. vaginatum. It also has swollen, rounded nodes, particularly near the base of older plants. This character-

istic is most evident on older, male plants. In contrast, A. nigrum and A. vaginatum lack swollen, rounded nodes

near the base of plants.

Another key characteristic of A. hondurense that separates it from the other dwarf mistletoes is the width

of its staminate spikes. While the length of staminate spikes often is too variable to be of any diagnostic value,

the width of the staminate spikes of A. hondurese are thinner (mean 1.7 mm) compared to those of A. nigrum

(2.9 mm) andA. vaginatum (2.0 mm). Furthermore, the staminate spikes of A. nigrum and A. hondurense gener-

ally do not form secondary branches, while those of A. vaginatum typically do.

Arceuthobium hondurense primarily forms 3-merous flowers, and occasionally 4-merous flowers, but A.

nigrum andA. vaginatum commonly form both 3- and 4-merous flowers. Although the adaxial surface of petals

of male flowers for A. hondurense andA. nigrum is distinctively dark red, the 3-merous flowers of A. hondurense

are smaller (2.5 mm) on average than those of A. nigrum (3.5 mm). The color of male flower petals of A. vagina-

turn, however, is dark brown to green. While the fruits of both A. hondurense and A. nigrum are usually mark-

edly glaucous, the fruits of A. nigrum are larger than those of A. hondurense as well as A. vaginatum. Addition-

ally, A. hondurense and A. nigrum primarily flower from late August through September and October, but A.

vaginatum flowers from March through April (Hawksworth & Wiens 1965, 1996). Additional observations of

A. nigrum are still necessary to determine if it flowers in the spring as reported by Hawksworth and Wiens

(1989, 1996). Our observations of A. nigrum over multiple seasons and years in Durango, Mexico, do not sup-

port a spring flowering period for A. nigrum. Furthermore, our analyses confirm that these species can be

readily distinguished using ITS-rDNA sequences as previously demonstrated by Nickrent et al. (1994, 2004).

The host specificity of these mistletoes may help separate them, depending on the locality in Mexico. In

Arango, Arceuthobium nigrum and A. vaginatum both parasitize P. teocote, but P. teocote is less susceptible to A.

vaginatum (a secondary host) (Hawksworth & Wiens 1996). Moreover, A. vaginatum does not parasitize P

Iftophylla, P lumholtzii, nor P. chihuahuana, which are all highly susceptible to A. nigrum. In central Mexico, the

Principal host of A. nigrum is P. teocote, but P. patula is also infected by A. vaginatum there. Now that A. hondu-

rense h as been discovered severely infecting P teocote in Oaxaca, Mexico, infection of this pine cannot be used

10 distinguish A. nigrum from A. hondurense, since these mistletoes both flower in the fall, have red flowers, and

are similar in size and color. The width of internodes and staminate spikes, therefore, are likely the best char-

acters for distinguishing between them. The size of 3-merous flowers, petals, and fruits will also assist in dis-

tln guishing A. nigrum (larger flowers and fruits) from A. hondurense (Table 2).

Based on our field observations and measurements of plant characteristics of the dwarf mistletoes in

**thern Mexico, we do not agree that Arceuthobium nigrum or A. vaginatum occur in Oaxaca or Chiapas,

hese species are primarily distributed along the Central Volcanic Cordillera of central Mexico and north into

Durango. Arceuthobium vaginatum extends as far north as central Chihuahua in the Sierra Madre Occidental

an d as far north as southern Coahuila in the Sierra Madre Oriental (Hawksworth & Wiens 1996). However, the

graphic distribution of A. nigrum is centered on the eastern side of the Central Cordillera and extends north

rato Durango (Fig. 1). Arceuthobium vaginatum is sympatric with A. nigrum in central Mexico (Hawksworth &

608 Journal of the Botanical Research Institute of Texas6(2)

Wiens 1996) and since it also extends as far north as Chihuahua, it is probably sympatric with A. nigrum in

Durango.