Journal of the Botanical Research Institute of Texas

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

Copyright 2012

Botanical Research

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Southern Methodist University

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researcher and writer, published the first issues of Sida,

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

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inherited editorship and copyright.

1^93 — BRIT becomes publisher/copyright holder.

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

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Journal of the Botanical Research Institute of Texas is

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Table of Contents

SYSTEMATICS

Penstemon kralii (Plantaginaceae), a new specie:

updated key to the southeastern U.S. taxa

A new species of Fragaria (Rosaceae) from Oregon

Kim E. Hummer

Ranunculus glacialis subsp. alaskensis subsp. no^

otherwise Atlantic species

Boris A. YuRTZEvt, David F. Murray, and Reidar Elven

S from AiAMSSOURirBQeTANIOAL

AUG 0 6 2012

w | garden library

New combinations for the Montana flora

17

Lomatium brunsfeldianum: a new species of Lomatium (Umbelliferae) from northern Idaho

Richard P. McNeill

Miconia cordieri, a new species of Miconia sect. Sagraea (Melastomataceae) from the Macaya

Biosphere Reserve, Haiti

Gretchen M. Ionta and Walter S. Judd

A new species of Vachellia (Fabaceae: Mimosoideae) from Haiti

David S. Seigler, Ricardo GarcIa, MilcIades MejIa, and John E. Emnger

Calathea rhizanthoides and C. peregrina (Marantaceae), new species endemic to Panama

Helen Kennedy

Calathea carolineae (Marantaceae), a new species endemic to Honduras

Helen Kennedy

Two new endemic species of Calathea (Marantaceae) from Panama

Helen Kennedy

Paradrymonia apicaudata (Gesneriaceae), a new species from western Colombia

M. Marcela Mora, John L. Clark, and Laurence M. Skog

Drymonia atropurpurea (Gesneriaceae), a new species from northwestern South America

Laura Clavijo and John L. Clark

A taxonomic study of the Galapagos endemic Vammia (Cordiaceae) species with

Julia K. Stutzman, Edgar B. Lickey, Andrea Weeks, and Conley K. McMullen

A new species of Tapeinostemon (Gentianaceae) from the Chocd of northern Ecuador

Jason R. Grant

Lectotypification of Bauhinia nervosa (Leguminosae: Caesalpinioideae)

S. Bandyopadhyay

Impatiens johnsiana (Balsaminaceae), a new scapigerous balsam from Western Ghats, India

M.K. Ratheesh Narayanan, N. Anil Kumar, Jayesh P. Joseph, C.N. Sunil, and T. Shaju

Commelina clavatoides (Commelinaceae), a new species from Kerala, India

Sheba M. Joseph, A. Anna Ancy Antony, and Santhosh Nampy

A Mid-Tertiary fossil flower of Swietenia (Meliaceae) in Dominican amber

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

Corrigenda: A new species of Erigeron (Asteraceae) from southwestern Oregon

Kenton L. Chambers

29

37

45

49

55

59

65

75

101

113

119

123

129

FLORISTICS, ECOLOGY, AND CONSERVATION

Woody species richness and abundance in a tropical savanna of Northern Ghana

Damian Tom-Dery, Patrick Boakye, and William J. Asante

latorral submontano de Rosaceas de la Sierra de

Zapaliname, Coahuila, Mexico

Juan A. Encina Dominguez, Sergio G. Gomez P£rez y Jesus Valdes Reyna

Flora of Tinajas Altas, Arizona — a century of botanical forays and forty thousand years

of Neotoma chronicles

A floristic inventory of Dismal Key and Fakahatchee Island — two shell mounds situated

within the Ten Thousand Islands region in the Gulf of Mexico (Collier County, Florida)

GJ. Wilder and MJ. Barry

Noteworthy plant records from Louisiana

Christopher S. Reid and Lowell Urbatsch

Further additions and emendations to the vascular flora of Caddo Parish, Louisiana

Barbara R. MacRoberts, Michael H. MacRoberts, Christopher S. Red, and Rosanna Ohlsson-Salmon

Discovery of Hydrocotyle bowlesioides (Araliaceae) in Louisiana

Robert W. Thornhill and Alexander Krings

An annotated checklist of the myxomycetes of the Big Thicket National Preserve, Texas

Katherine E. Winsett and Steven L. Stephenson

Cyperus albostr iatus (Cyperaceae) new to North America and Fimbristylis cymosa

(Cyperaceae) new to Texas

David J. Rosen, Richard Carter, Alfred Richardson, and Ken King

Phemeranthus calcaricus (Montiaceae) new to Texas

Rebecca K. Swadek

259

273

283

287

303

Osvaldo Morrone: 1957-2011

Liliana M. Giussani 309

Edward Groesbeck Voss: 1929-2012

Richard K. Rabeler and Anton A. Reznicek 311

Geraldine Ellis Watson: 1925-2012

Barney L. Lipscomb and George M. Diggs, Jr. 313

Book Reviews and Notices 16, 28, 48, 58, 64, 70, 100, 108, 112, 128, 130, 142, 258, 282, 286, 308, 315-321

Announcements 322

PENSTEMON KRALII (PLANTAGINACEAE),

A NEW SPECIES FROM ALABAMA AND TENNESSEE,

WITH AN UPDATED KEY TO THE SOUTHEASTERN U.S. TAXA

Dwayne Estes

Austin Peay State University

Department of Biology & Center of Excellence for Field Biology

P.O.Box4718

Clarksville, Tennessee 37044, USA

estesL@apsu.edu

ABSTRACT

RESUMEN

is y P. smallii. Se aporta ui

Penstemon Schmidel is the largest vascular plant genus endemic to North America. More than 250 of the 270+

species of this taxonomically challenging genus occur in western North America with the center of diversity in

the Intermountain Region (Wolfe et al. 2006). About 20 species occur naturally in the southeastern United

States. Many new species have been described over the past few decades from western North America but none

have been described from eastern North America since Pennell’s (1935) monograph, The Scrophulariaceae of

eastern temperate North America. During a recent examination of Penstemon specimens for the Flora of Tennes-

see project, specimens of an unusual Penstemon from just across the state line in northeastern Alabama were

encountered that did not fit any known species. These specimens bore a similarity to P. smallii A. Heller of the

Southern Appalachians and P. tenuis Small of the southern Mississippi River Valley. Further assessment led to

the determination that these populations represent an undescribed species. This species joins Blephilia sub-

nuda Simmers & Krai (1992), Clematis morefieldii Krai (Krai 1987), Polymniajohnbeckii D. Estes (Estes & Beck

2011), Silphium brachiatum Gatt., and Vemoniaflaccidifolia Small as the latest addition to the endemic flora of

rocky limestone woods of the southwestern escarpment of the Cumberland Plateau.

Penstemon kralii D. Estes, sp. nov. (Figs. 1-2). Type: UNITED STATES. Alabama. Jackson Co.: 3.8 km Sof Paint Rock, eastern

base of Keel Mountain at the western edge of the Paint Rock River Valley, along NW side of County Road 5, 1.6 km by road W from

jet. with US Hwy 72, edge of dry deciduous forest over Mississippian limestone, 34.626176° N, -86.329163° W, 4 May 2011, D. Estes

Plant a perennial herb from a caudex. Stems 4-7 dm tall, erect to ascending, solitary or clustered, unbranched

below inflorescences, terete, 4-5 mm in diameter at midstem, with 6-9 leafy nodes below inflorescence, pale

green, densely to moderately puberulent with spreading or retrorse, straight, eglandular, sharp-tipped white

hairs 0.1-03 mm, the eglandular pubescence above the distalmost non-bracteal leaf becoming more sparse

and replaced by mostly spreading, glandular-capitate hairs 0.5 mm. Leaves simple, oppositely-decussate,

spreading, often with apices recurved in life (not often evident in specimens), proximal oblanceolate and taper-

Journal of the Botanical Research Institute of Texas 6(1)

mg to a narrowly attenuate petiole-like base, medial lanceolate or oblong, those just proximal to the inflores-

cence ovate-lanceolate, medial blades 12-15 cm x 3-4 cm, usually larger than proximal and distal cauline

leaves, apex acute to acuminate, margins sharply and coarsely serrate with mostly 4-7 acuminate teeth per cm

(fewer on proximal leaves), base subcordate to rounded and clasping (proximal leaves tapered to a narrowly

clasping base); blades dark green, abaxial surface glabrate or those toward inflorescence moderately pubes-

cent; adaxial surface with scattered, eglandular, straight, sharp-tipped white hairs 0.1-0.3 mm, especially near

midvein, otherwise glabrate. Inflorescence a terminal thyrse, 5-20 cm x 3-12 cm, bracts at base of inflores-

imetimes equaling the distalmost non-bracteal stem leaves, primary

with dilated bases, secondary inflorescence bracts narrowly lanceo-

1 at base; inflorescence glandular-capitate. Flowers slightly nodding;

pedicels 1.5-4 mm, pubescent with glandular-capitate hairs. Calyx campanulate with 5 subequal sepals; these

distinct nearly to base, lance-attenuate and becoming strongly recurved, green with thin hyaline margins

proximal to middle, 4-5.5 mm x 1 mm, abaxial surface and margins moderately covered with gland-tipped

Journal of the Botanical Research Institute of Texas 6(1)

hairs, adaxial surface glabrous. Corolla bilabiate, tubular-funnelform, 10-16 mm, prominently 2-ridged

within on abaxial surface, pale lavender (adaxial surface of lobes nearly white) or sometimes nearly white with

prominent (sometimes faint) purple guide lines, especially on the abaxial surface of the corolla throat and on

the adaxial surface of corolla lobes, pubescent externally with glandular-capitate hairs; tube 6-7 mm, throat

5-7 mm long, open, abruptly expanded from the tube, lobes 5, 4-5 mm, abaxial lobes 3, spreading and de-

clined, projecting beyond the adaxial lobes, adaxial lobes 2, slightly shorter than the abaxial ones, upwardly

curved to erect. Stamens 4, included in throat, filaments strongly curving, 10-12 mm, anthers very finely

papillose, the 2 locules strongly divergent, each 0.8 mm x 0.5 mm. Staminode 7-8 mm, reaching the orifice

and slightly exserted, densely bearded in distal 4-5 mm, beard golden-brown, beard hairs 1-1.2 mm. Cap-

sules broadly ovoid, narrowly rounded at apex, 5-6 mm x 3.5-4 mm. Seeds ca. 1 mm (not seen at full matu-

rity). Chromosome number unknown.

Additional specimens examined: UNITED STATES. ALABAMA. Blotrnt Co.: banks of small calcareous branch near NW base of Wamock

[sic, recte Womock) Mountain, 4 May 1938, R. Harper 3658 (MO); 2.25 air mi. NW of Brooksville, area just N of Curtis Williams Lake, E of

US Hwy 231, 30 Apr 2011, B.R. Keener 6430 with T. Keener & W.K. Webb (UWAL). Jackson Co.: 0.4 of a mi E on US 72 from its jet. with Jack-

son County Road 63, 19 May 1990, S. & G. Jones 4880 (VDB). Madison Co.: mountain region, Huntsville, 29 May 1896, L.M. Underwood s.n.

Etymology.— The epithet “ kralii ” was chosen to honor Dr. Robert Krai (1926-), one of the top 20th-Century

American plant taxonomists. Dr. Krai is Professor Emeritus of Vanderbilt University, curator of the VDB her-

barium and research associate at the Botanical Research Institute of Texas in Ft. Worth. He has arguably been

one of the most significant contributors to the flora of the eastern United States and is widely renowned for his

knowledge of the flora of the southeastern U.S. Dr. Krai’s professional accomplishments are extraordinary.

During his 52-year career he has collected nearly 100,000 specimen numbers, not to mention hundreds of

thousands of duplicates that have been widely diwminatpH to dozens of herbaria. Additionally, he has named

more than 100 new species, including new taxa from both North and South America. He has tackled many

difficult taxonomic problems in eastern North American botany and his work has clarified such difficult gen-

era as Carex, Eriocaulon, Fimbristylis, R hynchospora, and Xyris. Although not widely circulated, Dr. Krai’s two-

volume, 1,305-paged U.S. Forest Service report, “A Report on Some Rare, Threatened, or Endangered Forest-Relat-

ed Vascular Plants of the South ” remains perhaps the most important single contribution to our understanding

of rare and narrowly endemic plants of the southeastern U.S. since J.K. Small published his Manual of the

Southeastern Flora in 1933. Dr. Krai continues to publish and work on taxonomic treatments, especially of New

World Xyridaceae, and The Flora of Alabama and Middle Tennessee. Given Dr. Krai’s exceptional taxonomic eye

and the fact that he has collected plants all over the Southeast, it is no surprise that he collected this new Pen-

stemon from Morgan County, Alabama in 1970. 1 have always admired and been inspired by Dr. Krai’s accom-

plishments and quality of work and it is my pleasure to name this new species in his honor.

Common Name. — Krai’s Beardtongue

Distribution and Ecology. — Penstemon kralii is endemic to the lower slopes of the southwestern Cum-

berland Plateau escarpment of northeastern Alabama and southeastern middle Tennessee (Fig. 3). It is

found at the edges of calcareous juniper-oak-hickory woodlands over Mississippian-aged limestone at

elevations ranging from 198-335 m (650-1,100 ft) above sea level. Tree species characteristically found

with P. kralii include Quercus muehlenbergii Engelm., Q. shumardii Buckley, Carya carolinae-septentrionalis

(Ashe) Engl. & Graebn., Fraxinus americana L .Juniperus virginiana L., Ulmus alata Michx., and Cotinus

obovatus Raf. Small tree and shrub species include Cercis canadensis L., Viburnum rufidulum Raf., Sideroxy-

ton lycioides L„ Hypericum frondosum Michx., and Forestiera ligustrina (Michx.) Poir. Herbaceous species at

Estes, A new species of Penstemon from the southeastern U.S.

these sites include Camassia scilloides (Raf.) Cory, Clematis glaucophylla Small, Helianthus microcephalus

Torr. & A. Gray, Polymnia canadensis L., Silphium brachiatum Gatt., Sisyrinchium sp. nov., Solidago auricu-

lata Shuttlew. ex S.F. Blake, Spigelia marilandica (L.) L., Verbesina virginica L., and Vemonia flaccidijolia Small.

Tennessee. The paucity of collections in herbaria gives some indication as to its rarity. Judging from the ab-

sence of herbarium specimens, P. kralii apparently has not been collected from Tennessee prior to this study.

This is intriguing given the abundance of fieldwork that has been conducted in southern Franklin County.

Although more extensive surveys will almost certainly result in the discovery of additional populations it is

likely that this species is truly rare and deserving of protection at the state level.

Collection History . — The earliest collection found for P. kralii dates to 1896 from near Huntsville, Alabama.

In 1938, Roland Harper collected it in Blount County, Alabama. His typed label originally did not give a spe-

cific epithet for the collection. Sometime after the original label was made, the inscription “laevigatus?” was

added in pencil, though it is not clear whether this was by Harper or someone else. In 1970, Robert Krai col-

lected this species from Morgan County, Alabama, and identified his specimen as “P. laevigatus ?”. From Krai’s

inclusion of the question mark, it is clear that he was not satisfied with his determination. Between 1964 and

1997, the species was collected several times by various botanists, adding additic

Jackson and Madison counties in Alabama. The first collections of this species fr<

May 2011 during this study. A review of specimens at TENN, UOS, UCHT, APSC,

Comparison with Similar Species . — In the herbarium Penstemon kralii resembles P. laevigatus Ait., P. tenuis,

and P smallii. Penstemon laevigatus is considered to be closely allied to P. calycosus Small and P. digitalis Nutt, ex

Sims (Pennell 1935). Penstemon laevigatus shares the following characteristics with P. digitalis and P. calycosus:

(1) leaves generally entire to remotely toothed; (2) stems, below the inflorescences, eglandular-puberulent with

very short trichomes either arranged in lines or essentially lacking, usually not consistently covering the

stems; (3) corolla tubes conspicuously and greatly dilated into the mi

surfaces of the corolla throats flat and not distinctly 2-ridged. In coi

that are consistently closely and sharply serrate; (2) stems that are closely and consistently eglandular-puberu-

lent throughout; (3) corolla tubes that are slightly expanded and lack conspicuously dilated throats; and (4)

interior abaxial surfaces of the corolla throats distinctly and prominently 2-ridged. It should be noted that co-

rolla tube dilation is relative. Penstemon kralii has tubes that are dilated compared to other Penstemon species

(e.g., P. tenuiflorus Pennell), but they are not nearly as dilated as in P. calycosus, P. digitalis, and P. laevigatus.

The combination of features described above for P. kralii is also found in P. smallii and P tenuis. None of

these species overlaps in distribution with P. kralii. From P. tenuis, P. kralii differs in its wider leaves, paler corol-

las (lavender to whitish with conspicuous dark purple veins vs. darker lavender, purple, or pink and inconspic-

uously-lined in P tenuis), and less-dilated corolla throats. Penstemon kralii also grows in a very different habitat

compared to P. tenuis. It is an upland plant of dry to submesic or rarely rich soil in edge habitats or along

hardwood forests, seasonally wet flatwoods, wet ditches, and wet prairies. Penstemon kralii and P. tenuis also

occur in two different regions. Penstemon kralii is endemic to the southern Cumberland Plateau escarpment of

southeastern Missouri where it is found mostly in the floodplain of the Mississippi River and its major tributar-

ies. Penstemon kralii is separated 300 km from the nearest populations of P. tenuis in the Mississippi Alluvial

Plain of west-central Mississippi. Small’s (1903) report of P. tenuis from Alabama may be based on P. kralii.

Based on morphology, P. kralii and P. tenuis seem to form a natural group with P. smallii. Penstemon smal-

lii is readily differentiated from smaller-flowered P. kralii and P. tenuis (corollas of both 10-16 mm) by its much

larger corollas (28-35 mm long). The corollas of P. smallii are dark pinkish or purplish and more closely re-

semble the pinkish or purplish corollas of P. tenuis than the very pale lavender to nearly white corollas of R

kralii. It also has generally larger leaves. In terms of habitat P. smallii and P. kralii are somewhat similar. Both

occur in upland, dissected, and generally mountainous terrain, and both often grow in edge habitats. Penste-

mon smallii typically occurs in moister habitats than P. kralii and is rarely associated with limestone-derived

Journal of the Botanical Research Institute of Texas 6(1)

Fig. 3. Distribution of Penstemon kralii (circles). The triangle represents a fruiting specimen (Gunn & Bailey 2008, VDB) that is either P. kralii or P. smallii; its

specific identity cannot be determined atthis time withoutflowers-Physiographicdesignations primarily follow Griffith etal. (2001). CuP= Cumberland

Plateau; eCuP = Cumberland Plateau Escarpment; DP = Dissected Plateau; EHR = Eastern Highland Rim; LM = Little Mountain; MV = Moulton Valley;

ONB = Outer Nashville Basin; SH = Shale Hills; STP = Southern Table Plateaus; SV = Sequatchie Valley; WHR = Western Highland Rim.

soils. Penstemon smallii is a Southern Appalachian endemic found in the Blue Ridge Mountains and adjacent

Ridge and Valley province. It is known from western North Carolina, eastern Tennessee, northern Georgia, and

northwestern South Carolina. The southwestern-most occurrence of P. smallii is from banks of the Tennessee

River near Chattanooga, Hamilton County, Tennessee, ca. 95 km NE of the nearest P. kralii population in Jack-

son County, Alabama. In the course of this study, all records of P. smallii from Tennessee and Alabama were

carefully checked for accuracy. Previous Alabama reports from Clay and Pike counties proved to be based on

misidentifications of Penstemon australis Small (C. Hansen, Auburn University, pers. comm. & A.R. Diamond,

Troy University, pers. comm.). Two additional specimens from northeastern Alabama were found that were

identified as P. smallii. One of these, a collection from Madison County (Threlkeld & Duke 1044, JSU), is actually

P. kralii. The other is a fruiting collection from bluffs above the Tennessee River in eastern Jackson County (Gunn

& Bailey 2008, VDB). This population is located 44 km E of the nearest P. kralii population in extreme western

Jackson County and 55 km SW of the nearest occurrence of P smallii in Hamilton County, Tennessee (also on a

bluff of the Tennessee River). Because this specimen is in fruiting condition, it cannot be positively determined

whether it is P kralii or P smallii. A visit to this population during flowering will be necessary to make a correct

identification. If it is P. kralii, then it would be the easternmost population. If it is P. smallii, then it would be the

southwestemmost occurrence for what is primarily a Blue Ridge species and Alabama’s only population.

UPDATED i

) PENSTEMON OF THE SOUTHEASTERN* UNITED STATES

Estes, A new species of Penstemon from the southeastern U.S.

■y obscurely lined (except P. at

ding-ascending, diverging fi

17. Corollas 20-32 mm.

22. Corollas 28-35 mm, lavender, violet, or purple; staminodes 1 5-1 8 mm, densely bearded in the

distal 13-15 mm; largest leaf blades mostly 30-60 mm wide R s

22. Corollas 10-16 mm; staminodes 7-9 mm, densely bearded in the distal 4-5 mm; largest leaf

23. Leaves mostly 30-40 mm wide; corolla pale-lavender to whitish, usually strongly lined with

dark violet lines, sepals linear-attenuate; plants of dry or mesic calcareous uplands

_P.kraliiD. Estes

*Penstemon fendleri Torr. & A. Gray, P. grandijlorus Nutt., and P. triflorus A. Heller occur along the extreme

western portion of what some consider the Southeastern U.S. in central Texas. These species have ranges that

occur primarily west of the Eastern Deciduous Forest region and are excluded from the above key. Measure-

ments in the key and text refer to length unless specified otherwise.

ACKNOWLEDGMENTS

1 thank the Austin Peay State University Center of Excellence for Field Biology and the Department of Biology

for travel support and release time for research. My graduate student, Clea Klagstad, kindly prepared the dis-

tribution map. Aaron Floden (TENN) and Wes Knapp (Maryland Department of Natural Resources) and my

graduate students, Dawn York and Courtney Gorman (APSC), assisted with fieldwork. I gready appreciate the

feedback provided by Wayne Chester (APSC), Craig Freeman (KANU), Noel Holmgren (NY), and Theo Witsell

(ANHC) who reviewed the manuscript prior to submission. Lastly, I especially thank Brian R. Keener (UWAL)

and Andrea D. Wolfe (OS) for helpful review comments.

Estes, D. andJ. Beck. 2011 .Anew species oiPolymnia (Asteraceae: Tribe Polymnieae) from Tennessee. Syst. Bot. 36:48 1-486.

Griffith, C.E., J.M. Omernik, J.A. Comstock, G. Martin, A. Goddard, and VJ. Hulcher. 2001. Ecoregions of Alabama. U.S.

Environmental Protection Agency, National Health and Environmental Effects Research Laboratory, Corvallis, OR

Kral, R. 1987. A new “Vioma” Clematis from northern Alabama. Ann. Missouri Bot. Gard. 74:665-669.

Pennell, EW. 1935. The Scrophulariaceae of eastern temperate North America. Acad. Nat. Sci. Philadelphia Monogr.

No. 1.

Simmers, R.W. and R. Kral. 1992. A new species of Blephilia (Lamiaceae) from northern Alabama. Rhodora 94:1-14.

Small, J.K. 1903. Flora of the southeastern United States. New Era Printing Company, Lancaster, PA.

Wolfe, A.D., C.P. Randle, S.L. Datwyler, J.J. Morawetz, N. Arguedas, and J. Diaz. 2006. Phylogeny, taxonomic affini-

ties, and biogeography of Penstemon (Plantaginaceae) based on ITS and cpDNA sequence data. Amer. J. Bot. 93:

1699-1713.

A NEW SPECIES OF FRAGARIA (ROSACEAE) FROM OREGON

Kim E. Hummer

USDA ARS National Clonal Germplasm Repository

33447 Peoria Road

Corvallis, Oregon 97333-2521 U.S.A.

Kim.Hummer@ars.usda.gov

ABSTRACT

RESUMEN

•: U.S.A. Oregon. Lane Co.: US NFDR 5897, southern exposure, dis-

n, with Pseudotsuga menziesii, 8 Aug 2011, K.E. Hummer CFRA 2111

Plants perennial, acaulescent, crown-forming herb with stolons and adventitious roots; petiole, pedicel, run-

ner and calyx with dense, spreading hairs. Stolons red, sympodial. Leaves generally trifoliate, rarely quadri-

or pentifoliate; slightly bluish green; adaxial surface with scattered appressed white hairs about 1 mm; not

thick or coriaceous, and not reticulate-veiny beneath. Shape of leaflets variable, elongated to obovate, cunei-

form or broadly round, truncate; margin variably serrate, entire to only upper third of leaflet; teeth small and

pointed, rarely crenate, ciliate; distal tooth of the terminal leaflet always narrower and shorter than adjacent

ones; accessory leaflets. Inflorescence compound dichasium with aborted epipodium including the terminal

flower, of variable length, sub-to super foliar, 4 to 15 cm. Flowers subdiecious with plants growing in separate,

or interspersed male, female, or hermaphrodite colonies; pistillate flowers 16.3(13.0-20.0) mm wide; stami-

nate flowers 19.7(15.0-24.0). Calyx green rarely regular with 5 lanceolate sepals and 5 bractlets, frequently ir-

regular, with 11 or 12 sub-equal parts. Petals white, usually 5 rarely 4 or 6, depressed obovate to very widely

obovate, entire, emarginate, or unequally-bilobed, partially overlapping or not; Stamens about 0.5 mm long,

protruding above sub-reflexed corolla; hermaphrodite flowers 23.1(17.0-30.0.) mm wide. Pollen grains about

34 pm air dried with longitudinal ridges (Type 1, Staudt 2009). Pistils light yellow, numerous, 1mm long, the

group centrally located within the flower as attached to receptacle about 3(3-4) mm. Receptacle in Fruit

shape variable from globose, subglobose, to ellipsoidal, and irregular; fruit set may be irregular, poor, or some-

times full, 1.0(0.5-1.2)cm wide x 1.5(1.0-1.75) cm long; fruit skin ripens red; fruit flesh white, juicy and sweet-

ish, with flat to sour flavor. Achenes sometimes set in pits; length - 1.56 mm; ovary vasculature prominent on

surface; lateral stylar edge frequently concave, forming a comma-shape. Chromosome number: 2 n = lOx = 70

(Nathewet et al. 2010; Hummer et al. 2011).

Differs from F. chiloensis and F. xananassa nothosp. cuneifolia by leaves not thick and coriaceous, not re-

ticulate-veiny beneath. Differs from F. vesca subsp. bracteata by leaves usually slightly bluish green, and distal

tooth of terminal leaflet always shorter than adjacent teeth as compared with leaves Kelly green and distal

'■ Bot Res. Inst Texas 6(1):9-1

Journal of the Botanical Research Institute of Texas 6(1)

tooth of terminal leaflet usually longer than adjacent teeth. Differs from F. virginiana subsp. platypetala by

leaves with adaxial, scattered, appressed white hairs (~1 mm) and achenes frequently comma-shaped, lateral

stylar edge concave as compared with adaxial leaf surface glaucous and achenes dome-shaped with straight

lateral stylar edge. The distribution range is in the western Oregon Cascades at elevations from 1,000 to 3,800

m. Differs from all sympatric congeners by the decaploid chromosome number (2 n = lOx = 70), whereas F.

vesca subsp. bracteata is (2n = 2x = 14) and F. virginiana subsp. platypetala is (2n = 8x = 56).

L. Wynd 202I(OSC). Hood R

t. OREGON. Deschutes Co.: Carpenter IV

e, Crater Lake

n 16829 (OSC). Jefferson Co.:

). Lane Co.: US NFDR 5897 leading to Waldo Lake, K. Hummer 2102 (OSC) 2104 (OSC) 2105

(OSC); US NFDR 5897 leading to Waldo Lake, K Hummer 2102 (OSC); US NFDR 5897 losing to Waldo Lake, K. Hummer 2104 (OSO; US ;

NFDR 5897 leading to Waldo Lake, K. Hummer 2105 (OSC); 2 mi above Alder Springs Campground, McKenzie Hwy, O R. Ireland592 (OSC).

Linn Co.: Hoodoo Butte, M.M. Thompson 95003 (OSC); Hoodoo Butte, M.M. Thompson 95025 (OSC); H

' >S< Sand) Spring, McKenzie Pass Summit, I “ ~ '

Big Lake, 44.37065, -121.86562, PI 551527; E of Big Lake, Pacific Crest Trailhead, 44.37933, -122.08560, PI 658460 ; W of Big Lake, 44.37766,

-121.87058, PI 657862; W of Big Lake, 45.37766, -121.87058, PI 657865; W of Big Lake. 44.37442, -121.87445, PI 657863; W of Big Lake,

45.41302, -122.087546, PI 657867.

Etymology. — This species is named for the Oregon Cascade Mountains.

Common name. — I suggest Cascade strawberry.

Distribution and habitat. — Populations of Fragaria cascadensis occur at elevations of 1,000 m to 3,800 m,

to the west of the divide in the Oregon Cascade Mountains; from the Columbia River in the north, to the south-

ern and western vicinity of Crater Lake in the south. This region is referred to as the “high peaks” (Alt &

Hyndman 2009). Habitat: open alpine meadows, or on forest path edges, where direct sunlight breaks through

the canopy; along stream banks or in roadside drainage ditches; growing in sandy-clay loam of volcanic origin.

At these locations the dominant vegetation is usually Douglas fir [Pseudotsuga menziesii (Mirbel) Franco] or

silver fir [Abies amabilis (Douglas ex Louden) Douglas ex Forbes]. Associated plants include: Gaultheria humi-

fusa (Graham) Rydb., Epilobium ciliatum Raf., Lupinus latifolius J. Agardh., Montia parvi/olia (Mocifio ex de

Candolle) Greene, Vida americana Muhl. ex Willd., Hieracium albiflorum Hook., Artemisia ludovidana Nutt.,

and Agoseris grandiflora (Nutt.) E. Greene. The soils have udic moisture and frigid soil temperature regimes.

The mean annual precipitation in the northern part of the decaploid distribution range is 200 to 250 cm

(Franklin & Dymess 1973), but it is 100 cm or lower in the southern area.

Phenology. — Fragaria cascadensis begins growing after snowmelt in late May or early June, flowering in

early July, about 2-3 weeks later than F. virginiana subsp. platypetala (Rydb.) Staudt at lower elevation below

1,000 m. Runner production begins after flowering. Fruit is ripe during August for about 2 weeks with plants

at ^ 1,500 m elevation ripening 1 to 2 weeks later than those at 1,000 m.

Journal of the Botanical Research Institute of Texas 6(1)

<■>

& % y 4

Fig. 3. Achenes of a) Fragaria vesca subsp. bracteata (PI 551 81 3, CFRA 464, near Lost Lake, Oregon); b) F. cascadensis Hummer. (PI 664461 , CFRA 21 12, Big

Lake Road, Oregon), and c) F. virginiana subsp. platypetala. (PI 551553, CFRA 176, near Prineville, Oregon). Bars represent 1 mm.

DISCUSSION

Throughout much of Oregon, F. virginiana subsp. platypetala and F. vesca subsp. bracteata (A. Heller) Staudt are

sympatric (Staudt 1999; Cook & Sundberg 2011). Sample collection for this study confirmed this frequent

sympatry as well as that of F. vesca subsp. bracteata with F. cascadensis at locations near Mt. Hood, Hoodoo

Butte, Iron Mountain, and Echo Mountain trail (Hummer et al. 2011).

From physical examination of native living plants at 17 locations from Mt. Hood through Crater Lake

where F. cascadensis was present, no sympatric octoploids, or other plants of intermediate ploidy, as deter-

mined by flow cytometry, were observed (Hummer et al. 2012). However, three plants collected by L. Wynd

(numbers 2020 for two plants and 2021 for one plant) in 1928, near the south entrance of Crater Lake, which

were mounted together on a single herbarium sheet (OSC 46316), comprise two congeners: two plants with

adaxial leaf surfaces that were glabrous and third plant with scattered leaf hairs (with hand written annotation

by the collector on the hairs of that specimen). Although the collector originally determined that the pressed

specimens on the sheet were one species despite the differential occurrence of the hairs, these specimens can

now be re-interpreted as two F. virginiana subsp. platypetala plants and one of F. cascadensis, growing in sym-

patry near Crater Lake. The occurrence of diploid F. vesca subsp. bracteata, and octoploid F. virginiana subsp

platypetala in sympatry with each other and separately with decaploid F. cascadensis supports the idea that

these taxa may have progenitor-descendant relationship.

Combination of morphological traits

Fragaria cascadensis has some morphological traits similar to those of F. virginiana subsp. platypetala and oth-

ers more like those of Fragaria vesca subsp. bracteata. From several meters away, leaf and inflorescence mor-

phology of the F. cascadensis is similar to that of F. virginiana subsp. platypetala, which is how this species es-

caped notice until recently. A defining difference is the presence of scattered appressed white hairs about 1 mm

long on the adaxial leaf surface of F. cascadensis. This trait differs from the smooth, glabrous adaxial leaf sur-

face of F. virginiana subsp. platypetala, but is similar to that ofF. vesca subsp. bracteata, which has scattered to

dense appressed, adaxial leaf surface hairs.

Another morphological difference separating Fragaria cascadensis from F. virginiana subsp. platypetala is

achene shape. Some of the achenes of the Cascade strawberry are comma-shaped, having a concave lateral

stylar edge, while those of F. virginiana subsp. platypetala are tear-drop shaped. The lengths of the achenes of F.

cascadensis and F. virginiana subsp. platypetala are similar, 1.56-1.85 mm, and achenes of both species tend to

onger than those of F. vesca subsp. bracteata, which are about 1.37-1.56 mm long (Staudt 1999). Achenes of

• vesca subsp. bracteata frequently have a persistent style, while those of the other two taxa do not.

Inflorescence lengths of Fragaria virginiana subsp. platypetala are often short, usually shorter than the

Hummer, A new species of Fragaria from Oregon

IB

foliage. The inflorescence length in F. cascadensis is variable within a population, ranging from 4 to 15 cm and

from shorter to taller than the foliage. That of F. vesca subsp. bracteata tends to be taller than the foliage.

1) Ploidy and Genetic Features

Decaploidy in Fragaria cascadensis was confirmed by chromosome counts and flow cytometry (Nathewet et al.

2010; Hummer et al. 2011, 2012). The Cascade strawberry has more alleles and more unique alleles (Njuguna

2010) than do octoploid strawberry species F. virginiana or F. chiloensis. A previous report of high genetic

variation in F. viriginiana subsp. platypetala (Hokanson et al. 2006) partly resulted from the unknowing inclu-

sion of three decaploid F. cascadensis samples (CFRA 101, CFRA 110, and CFRA 440) in that study.

Potential Progenitors

With a significant combination of morphological traits, F. cascadensis seems strongly linked to both F. vesca

subsp. bracteata and F. virginiana subsp. platypetala. The facultative outcrossing of F. vesca subsp. bracteata and

the findings of Dermen and Darrow (1938) and Njuguna et al. (2010) suggest that the direction of a hybridiza-

tion event would likely be F. vesca subsp. bracteata (or a polyploid derivative) as the mother, and F. virginiana

subsp. platypetala (or a derivative) as the pollen parent, if there was a direct cross. Reports summarized by

Dermen and Darrow (1938) and Darrow (1966) indicated that autotetraploids arising from a diploid as mother,

with the pollen of higher ploidy cytotypes, could produce a successful cross. The reciprocal cross (with pollen

from autotetraploid F. vesca) was unsuccessful in the laboratory (Dermen & Darrow 1938). This is, however,

counter to Bringhurst’s (1990) discussion of the origin of the California F. xbringhurstii Staudt pentaploids.

These genotypes have arisen from natural crosses of F. chiloensis with autotetraploid F. vesca subsp. califomica

(Cham. & Schltdl.) Staudt (Bringhurst 1990; Staudt 1999). Future work comparing plastid single nucleotide

polymorphisms between the putative parents and F. cascadensis could determine the maternal parent.

2) Previously unrecognized cytotype

This unusual cytotype was overlooked by many studying Fragaria taxonomy. Other than the decaploid chro-

mosome number, F. cascadensis appears similar to the octoploid F. virginiana subsp. platypetala as described by

Staudt (1999). Close inspection is needed to see the morphological differences of hairy adaxial leaf surfaces

and comma-shaped achenes. Therefore, F. cascadensis has gone unnoticed. Darrow (1966) who collected

strawberries and performed many ploidy experiments including synthetic development of multiple levels of

polyploidy was unaware of wild Fragaria species with more than 56 chromosomes. Hancock et al. (2001) and

Harrison et al. (1997) extensively studied octoploid F. virginiana Mill, and F. chiloensis Losinsk. from North

America, including F. v. subsp. platypetala from the Western Cascade Range of Washington, where all indi-

viduals are octoploid. Hokanson et al. (2006) unknowingly included three decaploid cytotypes (CFRA 101,

CFRA 110, and CFRA 440) in their analysis, but assumed them to be octoploid.

3) For the present study, 23 OSC herbarium specimens were determined to be F. cascadensis, judging from

adaxial leaf surface hairs and western High Cascades localities of origin. An additional 16 were determined as

F. vesca subsp. bracteata, judging from adaxial leaf surface hairs, inflorescence length, leaf dentation (size, loca-

tion, and number per leaf) and collection locality. Strawberry specimens at the OSC herbarium were reviewed

and annotated by Staudt. His (Staudt 1999) described the upper leaf surface of F. virginiana subsp. platypetala

as “smooth” based on his studies of strawberries from the Pacific North Western North America. With the

definition of F. cascadensis sp. nov., I suggest that “smooth and glabrous” is a more appropriate description for

the adaxial leaf surface of F. virginiana subsp. platypetala.

IDENTIFICATION KEY FOR WILD FRAGARIA IN OREGON

!. Leaves somewhat th

133-1.75 mm long); plants o

>t thick and coriacious, n<

CONCLUSIONS

Fragaria cascadensis, sp. nov., the Cascade strawberry, occurs in the western Oregon High Cascade Mountains.

Fragaria cascadensis is visually similar in plant, inflorescence, and most traits of leaf morphology to F. virgin-

iana subsp. platypetala octoploids as described by Staudt (1999). However, the decaploids also share two key

characters with F. vesca subsp. bracteata, including adaxial leaf hairs and predominantly comma-shaped

achenes.

We provide a cautionary note to breeders who wish to enhance the octoploid strawberry gene pool from

wild North American material. If decaploids are inadvertently used in breeding programs in lieu of octoploids,

crosses are possible but the resulting enneaploid progeny would have reduced fertility. As an example, a lower

fecundity was observed in second generation crosses from Staudt et al. (2009) of F. iturupensis (a species since

shown to be decaploid) with F. x ananassa cultivars (octoploid). From additional scattered sampling of wild F.

virginiana subspecies across North America (Hummer et al. 2011), decaploid zones, other than the one in the

Oregon High Cascade Mountains, have not been detected. However, detailed collection and examination of

ploidy is warranted where native diploids and octoploids are sympatric congeners.

Fragaria cascadensis presents the possibility for developing and breeding a new class of cultivated straw-

berries at the decaploid level. The wild Oregon decaploids, the Kurile decaploid, F. iturupensis (Hummer et al.

2009), and cultivars of the artificial species F. xvescana (Bauer 1993) may be inter-fertile, because of their

equivalent ploidy. Recently Japanese breeders reported on new synthetic decaploids that included genes from

the aromatic diploid F. nilgerrensis Schltdl. ex J. Gay (M. Morishita, pers. comm. 2012).Crossing within the

decaploids should produce fertile offspring. Perhaps new flavors or resistant genes could be available to con-

This newly described taxon enforces the view of post-glacial polyploid evolution (Harrison et al. 1997;

Hancock et al. 2004; Hokanson et al. 2006) in the North American strawberries — not only with continued

gene flow between sympatric octoploids, but with the additional dimension of hybrid combinations between

multiple ploidy levels, such as diploids with octoploids.

ACKNOWLEDGMENTS

The support for this project from USDA ARS CRIS 5358-21000-038-00D is gratefully acknowledged. The au-

thor thanks Kenton L. Chambers for comments on the diagnosis; Nahla Bassil and Wambui Njuguna for ge-

nomic information; Tom Davis, Aaron Liston, Jim Oliphant, for frank discussions of data interpretation; Sugae

Wada for photographic assistance; and Tom Davis and Daniel Porter for helpful reviews.

Hummer, A new species of Fragaria from Oregon

REFERENCES

15

Alt, D.D and D.W. Hyndman. 2009. Roadside geology of Oregon. Mountain Press Publishing Co., Missoula, Montana.

Bauer, A. 1993. Progress in breeding decaploid Fragaria xvescana. Acta Hort. 348:60-64.

Cook,T. and S. Sundberg (eds.). 201 1. Oregon vascular plant checklist. [Rosaceae]. http://www.oregonflora.org/checklist.

php. Accessed [2011-10-31].

Dermen, H. and G.M. Darrow. 1938. Colchicine-induced tetraploid and 16-ploid strawberries. Proc. Amer. Soc. Hort.

36:300-301.

Darrow, G. 1 966. The strawberry. Holt, Rinehart and Winston, New York, New York, USA.

Franklin, J.F. and C.T. Dyrness. 1973. Natural vegetation of Oregon and Washington. USDA Forest Ser

Technical Report PNW-8. Seattle, Washington.

Hancock, J.F., P.W. Callow, A. Dale, JJ. Luby, C.E. Finn, S.C. Hokanson, and K.E. Hummer. 2001 . From the Andes ti

native strawberry collection and utilization. HortSci. 36:221-225.

Hancock, J.F., S. Serce, C.M. Portman, P.W. Callow, and JJ. Luby. 2004. Taxonomic variation among North and South American

subspecies of Fragaria virginiana Miller and Fragaria chiloensis (L.) Miller. Canad. J. Bot. 82:1632-1644.

Hokanson, K.E., J.M. Smith, A.M. Connor, JJ. Luby, and J.F. Hancock. 2006. Relationships among subspecies of New World

octoploid strawberry species, Fragaria virginiana and Fragaria chiloensis, based on simple sequence repeat marker

analysis. Canad. J. Bot. 84:1 829-1 841 .

Harrison, R.E. JJ. Luby, G.R. Furnier, and J.F. Hancock. 1 997. Morphological and molecular variation among populations of

octoploid Fragaria virginiana and Fragaria chiloensis (Rosaceae) from North America. Amer. J. Bot. 84:61 2-620.

Hummer, K.E., N. Bassil, and W. Njuguna. 201 1 . Fragaria. Chapter 2. In: C. Kole, ed. Wild crop relatives: genomics and breed-

ing resources, temperate fruits. Springer WCR Series Vol. 6:17-44.

Hummer, K., N. Bassil, J. Postman, and P. Nathewet. 201 2. Chromosome numbers and flow cytometry of strawberry wild rela-

tives. Acta Hort. (accepted; in press).

Hummer, K., P. Nathewet, and T. Yanagi. 2009. Decaploidy in Fragaria iturupensis (Rosaceae). Amer. J. Bot. 96:71 3-71 6.

Manage 201 0. Karyotype analysis in wild diploii

strawberries, Fragaria (Rosaceae). Cytologia 75:277-288.

Njuguna, W. 2010. Development and use of molecular tools in Fragaria. Oregon State University, Corvallis. (Thesis).

Njuguna, W., A. Liston, R. Cronn, and N. Bassil 2010. Multiplexed Fragaria chloroplast genome sequencing. Acta H

859-315-321.

Staudt, G. 1 989. The species of Fragaria, their taxonomy and geographical distribution. Acta Hort. 265:23-24.

Staudt, G. 1999. Systematics and geographic distribution of the American strawberry species: taxonomic studies in the

genus Fragaria (Rosaceae: Potentilleae). Univ. Calif. Publ. Bot. 81:1-162.

Staudt, G. 2009. Strawberry biogeography, genetics and systematics. Acta Hort. 862:71-83.

Staudt, G., S. Schneider, P. Scheewe, D. Ulrich, and K. Olbricht. 2009. Fragaria iturupensis : a new source for strawberry improve-

ment? Acta Hort 842:479-482.

nstitute of Texas 6(1)

BOOK REVIEW

Nan K. Chase. 2010. Eat Your Yard: Edible Trees, Shrubs, Vines, Herbs and Flowers for Your Landscape.

(ISBN 13: 978-1-4236-0384-9; ISBN 10: 1-4236-0384-2, pbk.). Gibbs Smith, P.O. Box 667, Layton, Utah

84041, U.S.A. (Orders: www.gibbs-smith.com, 800-835-4993). $19.99, color throughout, 160 pp„ 7.5" x

Growing up during WWII meant most of our vegetables came from my father’s Victory Garden. Endive, mint

and parsley were grown in our yard near the fig tree. Stalking the Wild Asparagus , by Euell Gibbons, was one of

my favorite books back in the early 1970s. Later, when I had grandchildren, they learned what plants they could

eat rather than the ones they should not eat. They tasted the thyme, rosemary, garlic chives, etc. Every August

was kept free for “putting up” the peaches. The pantry and freezer were full of preserves, chutney and pies from

our delicious freestone peach tree. When the peach tree died, it was replaced with a pomegranate, which has

flowered but not yet produced fruit.

Eat Your Yard is right up my alley . . . planting for food and beauty. Ms. Chase has chapters on edible trees,

fruits, nuts, herbs and flowers. She includes histories of various plants; growing tips (locations and soil and sun

conditions); recipes for using the harvests; and information for freezing, canning, dehydrating, pressing and

Beautiful photographs illustrate how the trees and plants can be used to beautify the landscape around

your home. Ms. Chase’s choice of trees and plants takes into consideration their year round attractiveness in

your landscape.

— Ann Schrader, Volunteer,

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, U.SA

RANUNCULUS GLACIALIS SUBSP. ALASKENS1S SUBSP. NOV. (RANUNCULACEAE),

A BERING1AN RACE OF AN OTHERWISE ATLANTIC SPECIES

David F. Murray

Boris A.Yurtzev

Journal of the Botanical Research Institute of Texas 6(1)

A. Gray, also based on R. andersonii. The morphological and molecular analyses of Horandl et al. (2005), Pawn

et al. (2005), and Emadzade et al. (2010), using sequences of the malK/tmK and psbJ-petA regions, support

Beckwithia as a genus apart from Ranunculus, but monotypic, consisting only of B. andersonii.

The two northern species of Ranunculus — glacialis and camissonis — have been allied with European spe-

cies of sect. Aconitifolii Tutin containing the tail-grown and white-flowered R. aconitifolius L. and R. platanifolius

L., which are confined to montane and subalpine tail-herb communities, and with R. seguieri Vill. and R. kuepfcri

Greuter & Burdet, alpine species of scree slopes and margins of snowbeds, respectively. The northern taxa, R-

glacialis and R. camissonis, have a contrasting history, ecology, and distribution, but studies by Paun et al.

(2005) show that the species of section Aconitifolii are phylogenetically coherent.

Ranunculus glacialis and R. camissonis have the following characteristics in common: Perennial plants

with most leaves basal, long-petiolate, with blades deeply trisected or temately divided (with more or less peti-

olulate leaflets in R. glacialis), main segments mostly dissected anew; flowering stems one to several with 2-4

leaves, similar to the basal ones or less divided, and with 1-3 (4) flowers; lower parts of the plant subglabrous

or glabrous, upper parts with peduncles and sepals more or less densely pubescent with dark golden-brown

hairs; petals 5-8, broadly obovate, white, becoming red-tinged and then purple after pollination. Both species

are diploids with 2n = 16. We recognize three taxa: two species, R. camissonis and R. glacialis, the latter witn

two infraspecific taxa subsp. glacialis and subsp. alaskensis. Subspecies alaskensis is new to science.

Yurtzev et al. Ranunculus glacialis subsp. ;

lus glacialis L.

In our concept, this is a European, amphi-Atlantic, and American Beringian species with two subspecies, the

ranges of which are reported below. It is found in wet to damp gravelly or stony habitats, in open vegetation

directly on mineral substrates without peat or other soil profiles, often in high alpine and arctic snowbank en-

vironments, on gravel bars along brooks and rivers, in periglacial environments including fresh morainal

gravel and nunataks, and in wet or north-facing humid screes.

Ranunculus glacialis subsp. glacialis

The range is amphi-Atlantic. It is found in eastern Greenland, Jan Mayen, the southernmost part of Svalbard,

Iceland, Fennoscandia eastwards to the Khibiny Mountains on the central Kola Peninsula (Russia), and in the

major central European mountain ranges from the Pyrenees eastwards to the Carpathians. It reaches the alti-

tudinal limit among vascular plants in Scandinavia (2370 m.s.m. Lid and Lid 2005) and is among those plants

reaching the highest elevations in the Alps.

Ranunculus glacialis subsp. alaskensis Yurtzev, subsp. nov. (Fig. 2). Type: U.S.A. ALASKA: Seward Peninsula, Kigluaik

Habitat in peninsula Sewardensi Alaskae Occidentalis in montibus kigluaikensibus in decliviis et summatibus

lapidosis glareosisque (non calcareis).

Subspecies alaskensis differs from subsp. glacialis by having in the upper parts of stem and peduncles

dark brownish-red pubescence (not glabrous or subglabrous as in subsp glacialis ), sepals with hairs up to 4 mm

long, patent, erect to descending (vs hairs subappressed, shorter and flexuous). It differs from R. camissonis in

its blades reniform in outline (not cuneate), temately divided with petiolulate main segments, more numerous

and more obtuse segments, the lateral ones shorter (not narrowly lanceolate and subacute), stems ascending or

rarely prostrate (not suberect or erect), and sepals obovate (not ovate). See Figure 2.

This subspecies is known only from western Alaska: Seward Peninsula, four sites in the Kigluaik

Mountains (Fig. 1) on rocky slopes of platy scree and on ridges and summits on non-calcareous substrates.

Ranunculus camissonis Schltdl.

Ranunculus glacialis subsp. camissonis (Schltdl.) Hulttn; Ranunculus glacialis var. camissonis (Schltdl.) L.D. Benson; Beckwithia

camissonis (Schltdl.) Total.; Beckwithia glacialis subsp. camissonis (SchltdL) A. Ldve & D. Lftve

ii-Beringian species reaches from the eastern Anyui mountains of northeastern Siberia and

ssian Far East eastwards to western and northern Alaska with outliers in the alpine tundra of

; in the Interior (cf„ Hulten 1968). It is found in moist to wet tundras, often on peat but mostly

Fig. 2. Ranunculus glacialis L. subsp. alaskensis Jurtzev, subsp. nov. Habit, flower, and sepal, based on ALA V1 14478. Alaska: Solomon Quad.: Seward

Peninsula, Big Creek Valley, 23 Jul 1993, D.F. Murray, B.A. Yurtuv& T Kelso 1 1763. Illustration: Anne Elven.

with underlying calcareous bedrock or morainal material, with a vegetation of forbs, sedges and mosses, often

in seepages, in saddles (interfluves), along streams and temporary watercourses, and on moist, non-sorted soil

circles (“frost boils”). The species has a preference for circumneutral substrates and avoids both dry limestone

and very acidic sites.

Ranunculus camissonis differs from R. glacialis in several characters. No transitions have been observed in

the material studied. Whether this is due to reproductive isolation or to their radically different habitats, mak-

ing co-occurrence nearly impossible, is not known.

The range of Ranunculus glacialis subsp. alaskensis is within that of R. camissonis. Although the two are in

a broad sense sympatric, due to the very different site requirements R. glacialis subsp. alaskensis is locally al-

lopatric and has not yet been found at or even very close to a site of R. camissonis. The two plants are visually

quite different, especially in growth shape and leaf shape, and are unlikely, in our view, to be mistaken for each

other. Regrettably the illustration provided by Schlechtendal (1819) does not accurately portray the basal

leaves of R. camissonis. See instead Figure 3 prepared for this paper.

The habitat of subsp. alaskensis is very similar to that of Atlantic R. glacialis s. str. and quite dissimilar to

that of R. camissonis. The features it shares with R. camissonis are quantitative ones in the amount and length of

pubescence and in petal proportions (see Figs. 2 and 3 for sepals). The constellation of characters separating

subsp. alaskensis from R. camissonis and the lack of any transitions is sufficient to justify subsp. alaskensis as a

taxon separate from R. camissonis and belonging to R. glacialis.

Fk. 3. Ranunculus camissonis Sdiltdl. Habit, flower, and sepal, based on ALA VT14333. Alaska: Bendeleben Quad.: Seward Peninsula, Minnie Creek

headwaters, 16 Jul 1993, D.F. Murray, B.A. Yurtzev& T. Kelso 1 1597. Illustration: Anne Elven.

Differences between Atlantic and Beringian races of R. glacialis are to be expected from the large disjunc-

tion, either across the Polar Sea (more than 3000 km) or across nearly the entire continents with gaps of more

than 130° longitude from eastern Greenland west to Alaska and more than 150° longitude from Alaska west to

the Murman area in European Russia.

22 Journal of the Botanical Research Institute of Texas6(1)

Schonswetter et al. (2003, 2004) postulated the persistence of R. glacialis in peripheral refugia of the Alps I

with post-glacial expansion to North Europe by long distance dispersal. However, the disjunctions and the

discontinuous morphological variation in R. glacialis subsp. alaskensis and R. camissonis suggest origins well f

prior to the postglacial period of 10,000-15,000 years ago, although not nearly as early (Miocene) as proposed j

by Hoffmann et al. (2010). Whereas we agree with an origin in European mountains for R. glacialis subsp. gla-

cialis or its precursor, we are still unable to offer an explanation for the Beringian R. glacialis subsp. alaskensis ■

or for R. camissonis, which are wholly arctic in distribution.

The Beringian taxa suggest migrations or dispersal events from a European center across to Beringia d- 1

ther by an easterly or westerly direction. Dispersal across Eurasia or North America, without leaving remnant

populations is not probable, although that is precisely the explanation offered for similar disjunctions in 1

Androsace : dispersal of ancestors or of species from this genus during a warmer interval (interglacial) and sub- J

sequent persistence in the Beringian refugium throughout the last glacial maximum with the extirpation of the |

intervening populations during the full-glacial expansion of glaciers and ice-sheets (Schneeweiss et al. 2004). |

Fragmentation and significant loss of populations from a formerly more continuous, widely Eurasian or cir- :

cumpolar range of R. glacialis seems unlikely to account for its present limits inasmuch as iL has site require-

intervals. How does one account for the remarkable geographic gaps?

A similar case of subspecies with gaps of distribution is Saxifraga rivularis subsp. rivularis and subsp.

arctolitoralis, the former European, the latter Beringian with disjunct extensions to Greenland. The two sub-

species are explained in this case as two separate, post-glacial, long-distance dispersal events from refugial

centers established prior to the last major glaciation (Westergaard et al. 2010).

Any hypothesis as to how R. glacialis attained its disjunct range, and subsp. alaskensis its characters, is |

pure guesswork, and the former question may be unanswerable.

A phylogeographic study including R. camissonis , R. glacialis subsp. alaskensis and subsp. glacialis, em-

ploying plastid DNA and ITS data (Ronikier et al. in prep.), shows that the genetic divergence between R. gla-

cialis subsp. glacialis on the one hand and R. camissonis and R. glacialis subsp. alaskensis on the other is much

more shallow than the genetic split between populations of R. glacialis subsp. glacialis from the western and

eastern Alps, indicating that dispersal to Beringia happened relatively late in the history of the complex. As the

Beringian populations are phylogenetically deeply nested in R, glacialis subsp. glacialis and subsp. alaskensis

and R. camissonis are not readily distinguishable genetically, they prefer to treat R. camissonis as subspecies of

R. glacialis and are reluctant to recognise R. glacialis subsp. alaskensis as new taxon before better resolution

with genetic data becomes available (Schonswetter pers. comm.).

We acknowledge the lack of agreement. Incongruence between phylogenetic and morphological studies

of the same taxa are not uncommon. However, we see and by the key demonstrate that there are morphological

discontinuities that distinguish the taxa. The fact remains that R. glacialis subsp. alaskensis has morphological

features not found in subsp. glacialis. It is possible to discriminate consistently the two subspecies. It follows

that the development of these features is under genetic control. It is obvious, too, that subsp. alaskensis is dis-

tinct from R. camissonis and more closely aligned with R, glacialis, albeit geographically distant from subsp.

glacialis. We see no taxonomic value in making the three taxa equivalent. That R. glacialis and R. camissonis

share genetic material is not surprising, but it is not inconsistent to present a taxonomy that emphasizes the

sharply contrasting phenotypes and distinct ecological separation between R. glacialis subsp. alaskensis and R.

camissonis.

(ALA); Glacial Lake, Sinuk River valley, 64*52™, 165°45'W, 702 11 JuLM 93, D.F. Murray, BA. Lrtzev, T. Kelso 11398. holotype (ALA);

Big Creek valley, 64°59'N, 164*50^ 23Jul 1993, D.F. Murray, BA Yuriev, T. Kelso 11763 (ALA); Upper Fox Creek, 64°59'N, 165°03'W,914

m.s.m., 5 Jul 1996, D.F. Murray, R. Upkin 1218 3 (ALA).

Boris Alexandrovich Yurtzev presented us with the original manuscript prior to his death, which we have edited

and expanded, but his views with respect to the new subspecies have been faithfully retained, and he alone is

the author of the new subspecies. Sylvia Kelso was one of the field team with Yurtzev and Murray when the

discovery of the new taxon was made, and we thank her for numerous helpful comments while preparing this

manuscript. Our thanks to Anne Elven for the line drawings, to Zachary Meyers for the map, to Rob Lipkin for

his work with us in the field 1996 and 1997. Dale Taylor, then with National Park Service, Anchorage, a strong

supporter of Beringian studies, made possible the two summers of joint U.S.-Russia expeditions to the Seward

Peninsula. Without his enthusiastic, hands-on support, these would not have been so successful. We are grate-

ful to Peter Schdnswetter for not only allowing us to see unpublished data on the molecular genetics of

Ranunculus glacialis and R. camissonis, but also for carefully reviewing the manuscript, for summarizing the

molecular results, and for suggesting important changes. Finally, our thanks to Alan R. Batten for his meticu-

lous review of the manuscript for this journal.

Al-Shehbaz, I., J.R. Grant, R. Lipkin, D.F. Murray, and C. Parker. 2007. Parrya nauruaq (Brassicaceae), a new species from Alaska.

Novon 17:275-278.

Emadzade, K., C. Lehnebach, P. Lockhart, and E. Horandl. 2010. A molecular phylogeny, morphology and classification of

genera of Ranunculeae (Ranunculaceae). Taxon 59:809-828.

Hoffmann, M.H., K.B. von Hagen, E. Horandl, M. Roser, and N.V. Tkach. 2010. Sources of the arctic flora: origins of arctic species

in Ranunculus and related genera. Int. J. PI. Sci. 1 71 50-1 06

HOrandl, E., 0. Paun, J.T. Johansson, C. Lehnebach, T. Armstrong, L. Chen, and P. Lockhart. 2005. Phylogenetic relationships and

evolutionary traits in Ranunculus s.l. (Ranunculaceae) inferred from ITS sequence analysis. Molec. Phylogen. Evol.

36:305-327.

HultEn, E. 1 937. Outline of the history of arctic and boreal biota during the Quaternary Period. Bokforlags Aktiebolaget

Thule, Stockholm.

HultEn, E. 1968. Flora of Alaska and Neighboring Territories. Stanford Univ. Press, Stanford, CA.

Ickert-Bond, S., D.F. Murray, and E. DeChaine. 2009. Contrasting patterns of plant distribution in Beringia. Alaska Park Sci.

8(2):26-32.

Kelso, S. 1 983. Range extensions of vascular plants from the Seward Peninsula, northwest Alaska. Rhodora 85:371-379.

Kelso, S. 1 987. Primula anvilensis (Primulaceae): a new species from northwestern Alaska. Syst. Bot. 1 2:9-1 3.

Kelso, S. 1989. Vascular flora and phytogeography of Cape Prince of Wales, Seward Peninsula, Alaska. Canad. J. Bot.

67:3248-3259.

Kelso, S., BA Yurtzev, and D.F. Murray. 1994. Douglasia beringensis (Primulaceae): a new species from northwestern

Alaska. Novon 4:381-385.

Lid, J. and D.T. Lid. 2005. Norsk flora, 7th edition by R. Elven. Det norsk samlaget. Oslo.

Love, A. and D. Love. 1 956. Cytotaxonomical conspectus of the Icelandic flora. Acta Hort. Gotob. 20(4):65-291 .

Murray, D.F., S. Kelso, and B.A. Yurtzev. 1994. Floristic novelties in Beringia: patterns and questions of their origins. In:

Meehan, R. H„ V. Sergienko, and G. Weller, eds. Bridges of science between North America and the Russian Far East.

Proc 45th Arctic Sci. Conf. Anchorage and Vladivostok. Pp. 1 86-192.

Paun, O, C. Uhnebach, J.T. Johansson, P. Lockhart, and E. Horandl. 2005. Phylogenetic relationships and biogeography of

Ranunculus and allied genera (Ranunculaceae) in the Mediterranean region and in the European Alpine System.

Taxon 54:91 1-930.

Porsild, AE. 1 939. Contributions to the flora of Alaska. Rhodora 41:141-183; 1 99-254 + pi 55 1 -553; 262-301 + pi 554.

Roniwer, M„ G.M. Schneeweiss, and P. Schonswetter. In prep. The extreme disjunction between Beringia and Europe in the

arctic-alpine Ranunculus glacialis does not coincide with the deepest genetic spilt - a story of the importance of

temperate mountain ranges in arctic-alpine phylogeography.

Schlechtendal, D.F.L 1819. Animadversiones Botanicae in Ranunculeas Candollii. Sectio prior. Berlin.

Schneeweiss, G.M., P. Schonswetter, S. Kelso, and H. Niklfeld. 2004. Complex biogeographic patterns in Androsace

(Primulaceae) and related genera: evidence from phylogenetic analyses of nuclear internal transcribed spacer and

plastid trnL-F sequences. Syst. Biol. 53:856-876.

60)1

SchOnswetter, P., O. Paun, A. Tribsch, and H, Niklfeld. 2003. Out of the Alps: colonization of northern Europe by east Alpine

populations of the Glacier buttercup Ranunculus glacialis L. (Ranunculaceae). Molec. Ecol. 1 2:3373-3381 .

SchOnswetter, P., A. Tribsch, I. Stehlik, and H. Niklfeld. 2004. Glacial history of high alpine Ranunculus glacialis (Ranunculaceae) 5

in the European Alps in a comparative phylogeographic context. Biol. J. Linn. Soc. 81 :1 83-1 95.

Westergaard, K.B., M.H. Jorgensen, T.M. Gabrielsen, I.G. Alsos, and C. Brochmann. 2010. The extreme Beringian/Atlantic dis-

junction in Saxifraga rivuiaris (Saxifragaceae) has formed at least twice. J. Biogeogr. 37:1 262-1 276.

Whittemore, A.L 1 997. Ranunculus Linnaeus. In: Flora of North America Editorial Committee, eds. Flora of North America

north of Mexico. 3. Magnoliophyta: Magnoliidae and Flamamelidae. Pp. 88-1 35.

Yurtzev, B.A. 1 972. Phytogeography of Northeastern Asia and the problem of transberingian floristic interrelations. In: A.

Graham, ed. Floristics and Paleofloristics of Asia and Eastern North America. Elsevier, Amsterdam. Pp. 1 9-54. ; j

NEW COMBINATIONS FOR THE MONTANA FLORA

Peter Lesica

Division of Biological Sciences

University of Montana

Missoula , Montana 59812 USA

peter.lesica@mso.umt.edu

Examination of herbarium material in preparation for a new floristic manual for Montana has convinced me

that several new combinations are warranted.

is lackschewitzii Douglass M. Hend. & R.K. Moseley, Syst. Bot. 15:462^65. 1990.

ies: var. aurantiaca has lanc<

trrowed to the beak; var. purpure

Cronquist has ovate-attenuate, outer phyllaries, ciliate but glabrous on the outer surface and a

ally tapered to the beak (Cronquist 1994; Baird 2006). Agoseris carnea Rydb. has tradit

under A. aurantiaca var. aurantiaca because of its similar involucre (Hitchcock et al.

Baird 2006) although it has pink rather than orange rays in fresh material (Rydberg 1900). 1

(1990) described this pink-flowered Agoseris as A. lackschewitzii from material collect*

and adjacent Montana, U.S.A. and were unaware of the plant occurring in British Columbia, Cana*

ing been previously described as A. carnea by Rydberg. However, they correctly pointed out tha

flowered form has narrow, villous phyllaries as in A. aurantiaca var. aurantiaca but achenes gradu

to the beak as in var. purpurea, and that it occurred in moist to wet meadows, an un

variety of A, aurantiaca. Though Henderson et al. (1990) believed these plants represented a distin

the close relationship to A. aurantiaca cannot be denied, and I believe it is better placed as a third v;

J Bot. Res. Inst. Texas 6 ( 1 ): 25- 27.2012

Journal of the Botanical Research Institute of Texas 6(1)

Columbia and Alberta south to Wyoming and Idaho. Rydberg (1900) gives the type locality as Mt. Queest, h

“Mt. Avert” is given on the holotype.

Representative specimens examined: CANADA. British Columbia: Mt.

National Park. Sunshine Meadows, 8 Sep 1991, P. Achuff6038 (MONTU). I

Artemisia lindleyana has been treated as a distinct species (Hitchcock & Cronquist 1973; Dorn 1984) or a sub- i

species of the European A. vulgaris L. (ssp. lindleyana H.M. Hall & Clements). The most recent treatment of

Artemisia for North America (Shultz 2006) considers A. lindleyana conspecific with A. ludoviciana Nutt. Shultz

(2006) recognized six subspecies within A. ludoviciana, and A. lindleyana was reduced to synonymy under ssp.

incompta (Nutt.) Keck. Similar suffrutescent habit, flowers and involucre indicate a close relationship between 1

A. lindleyana and A. ludoviciana. Within this complex both A. ludoviciana ssp. incompta and A. lindleyana have

leaves that are glabrate above. However, the former has glabrate phyllaries, a paniculate inflorescence and

deeply lobed leaves, while the latter has racemes of heads with tomentose involucres and leaves that are entire |

or nearly so. I agree with Shultz that A. lindleyana should be placed within A. ludoviciana, but believe that dif- 1

ferences between A. lindleyana and A. ludoviciana ssp. incompta preclude subsuming the former in the latter I

Shultz (2006) suggested that A. lindleyana may warrant infraspecific status under A. ludoviciana, and Cronquist j

stated that he had observed A. lindleyana growing adjacent to A. ludoviciana sensu stricto without intermedi- j

ates (Hitchcock et al. 1955). For these reasons I propose lindleyana as a seventh subspecies of A. ludoviciana. |

Artemisia ludoviciana ssp. lindleyana is found on sandy, gravelly or rocky banks of rivers from southern .

British Columbia to Oregon and east to Idaho and Montana west of the Continental Divide (Hitchcock et al .

1955). The other three subspecies of A. ludoviciana in Montana generally occur in different habitats than ssp. j

lindleyana; ssp. ludoviciana occurs in grasslands, sagebrush steppe and meadows; ssp. candicans (Rydb.) Keck

is found in grasslands, streambanks and roadsides; ssp. incompta (Nutt.) Keck occurs in stony soil of talus

slopes, rock outcrops and sagebrush steppe. Montana’s four subspecies have different combinations of a few

variable characters and can be differentiated with the following key:

1 . Leaves glabrate and greenish above.

mens examined: U.S.A. Montana. Flathead Co.: Middle Fork Flathead River, 13 Aug 2004, P. Lest ca 89 36 (MONTU)

Lake Co.: Flathead Indian Reservation, Lower Flathead River, 22 Aug 1984, S. Gregory 2894 (MONTU). Mineral Co.: Clark Fork River, 26

Oct 1978, K. Lackschewitz 8799 (MONTU). Missoula Co.: Blackfoot River, Johnsrud Park, 19 Sep 1967, M. M ooar5559 (MONTU). Sanders

Co.: Thompson Falls, 6 Aug 1901 ,J. Blankenship 409 (MONT); sandy riverbanks, Thompson Falls, 2 Aug 1957, W. Booth 571198 (MONT). |

Symphyotrichum foliaceum (Lindl. ex DC.) G.L. Nesom va

» Gray) P. Lesica, o

i. Gray) Cronq., /

29:429-468. 1943. Symphyotrichum cusickii (A. Gray) G.L. Nesom, Phytologia 77:141-297. 1!

Aster cusicku was first described by Asa Gray based on a Cusick collection from northeastern Oregon (Grayj

1880). Sixty years later Cronquist argued that the plant graded into other forms of A foliaceus and was best

treated as a variety of that species (Cronquist 1943). Nesom moved the taxon into the genus Symphyotrichum

and recognized it at the species level (Nesom 1994). In the most recent treatment of the group, Brouillet et al.

(2006) also recognized the taxon at the species rather than at the infraspecific level. My review of mainly

Montana material suggests that low-elevation segregates of Symphyotrichum foliaceum sensu lato from the

Northern Rocky Mountains often cannot always be confidently distinguished from each other based on the

plastic, continuous, vegetative characters purported to separate them (Cronquist 1943). Thus, I prefer to recog-

nize this plant as a variety of Symphyotrichum foliaceum rather than at the species level; however, the desired

combination had not been previously made. Symphyotrichum foliaceum var. cusickii can be distinguished from

other varieties of S. foliaceum usi n g keys presented by Cronquist (1943) and Hitchcock and Cronquist (1973).

ACKNOWLEDGMENTS

The review comments of John C. Semple (WAT) and one anonymous reviewer are gratefully acknowledged.

Kanchi Gandhi (GH) and Nicholas Hind (MO) helped with nomenclatural advice.

Pp. 323-335.

Brouillet, L., J.C. Semple, G.A. Allen, K.L Chambers, ano S.D. Sundberg. 2006. Symphyotrichum. In: FNA Editorial Committee,

Flora of North America, Vol. 20, Oxford University Press, New York. Pp. 465-539.

Cronquist, A. 1943. Revision of the Western North American species of Aster centering about Aster foliaceus Lindl. Amer.

Midi. Naturalist 29:429-468.

Cronquist, A. 1994. Intermountain flora. Volume 5: Asterales.The New York Botanical Garden, Bronx.

Dorn, R.D. 1 984. Vascular plants of Montana. Mountain West Publishing, Cheyenne, WY.

Gray, A. 1 880. Botanical contributions 1 880. 1 . Notes on some Compositae. Proc. Amer. Acad. Arts 1 6:78-1 02.

Henderson, D.M., R.K. Moseley, and A.F. Cholewa. 1990. A new Agoseris (Asteraceae) from Idaho and Montana. Syst. Bot.

15:462-465.

Hitchcock, C.L., A. Cronquist, M. Owenby, and J.W. Thompson. 1955. Vascular plants of the Pacific Northwest. Part 5:

Compositae. University of Washington Press, Seattle.

Hitchcock, C.L. and A. Conquist. 1 973. Flora of the Pacific Northwest. University of Washington Press, Seattle.

Keck, D.D. 1 946. A revision of the Artemisia vulgaris complex in North America. Proc. Calif. Acad. Sci. Ser. 4, 25:421-468.

Nesom, G.L. 1994. Review of taxonomy of Aster sensu lato (Asteraceae: Asterae) emphasizing the New World species.

Phytologia 77:141-297.

Rydberg, P.A. 1 900. Catalogue of the flora of Montana and the Yellowstone National Park. Mem. New York Bot. Gard. Vol. 1 .

Shultz, L.M. 2006. Artemisia. In: FNA Editorial Committee, Flora of North America, Vol. 20, Oxford University Press, New

York. Pp. 503-534.

Journal of the Botanical Research Institute of Texas 6(1)

BOOK REVIEWS

Lynn M. Steiner. 2010. Prairie-Style Gardens: Capturing the Essence of the American Prairie Wherever

You Live. (ISBN: 978-1-60469-003-3, hbk.). 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). $34.95, 304 pp„ 182 color photos, 1 map, 7" x 9".

Prairie-Style Gardens: Capturing the Essence of the American Prairie Wherever You Live is a motivating, beautiful

book with many photographs of wildflowers and grasses found in American prairies. The book is divided into

four chapters and includes profiles of flowers and grasses and sedges. It begins with a chapter about the inspira-

tion for the prairie and by the time I finished that chapter I was hooked. Steiner clearly is enamored with the

prairie plants and convinces the reader anyone can help re-create a bit of the prairie. The author conveys her

passion for the prairie through her writing and photographs.

This book encourages the reader to add prairie plantings into the garden no matter how big or small the

available space. Steiner makes a good case for restoring prairies and biodiversity. She shows great respect for

the plants that survive through high winds, drought conditions, and repetitive disturbances. There are helpful

tips throughout the book on everything from preparing the soil to winning the neighbors over with your natu-

The Flowers Index is quite extensive with a colored photograph next to every detailed description. This is

incredibly helpful to the novice prairie gardener. The information covers wildlife attraction, landscape uses, -

knew which flower she was currently referencing. Perhaps covering the flowers first in the book would allevi-

ate this problem. The Grasses and Sedges section was a disappointment with only nine different species repre-

sented. Many of the photographs were captioned with both the scientific and common name while others only

included the common name. It would be helpful to use both names throughout. Overall this is an inspiring

book for any gardener, not just those already intrigued with native plants.

—Julie Donovan, Coordinator of Volunteers, BRIT, 1700 University Drive, Fort Worth, Texas 76107-3400, USJjM

Marty Wingate. 2011. Landscaping for Privacy: Innovative Ways to Turn Your Outdoor Space into a

Peaceful Retreat. (ISBN: 978-1-60469-123-8, pbk.). Timber Press, Inc., The Haseltine Building, 133 S.W.

Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: wwwtimberpress.com, 800-

327-5680). $19.95, 155 pp„ color photos, 8" x 9".

Andrew Buchanan’s book cover photograph serves as the subliminal concierge to Marty Wingate’s Landscaping

for Privacy: Innovative Ways to Turn Your Outdoor Space into a Peaceful Retreat.

Ms. Wingate’s book is a clearly visible written example of her landscape gardening/design prowess. I® 1

addition, it demonstrates she is a savvy author who has garnered an extraordinarily talented team of profes- 1

sional photographers, book designers, and a well known publisher of books on gardening, horticulture, etc. ;

The reader’s sojourn into the Table of Contents will ensure the reader will turn the next page. While the

writing is for a general readership, it is cerebral fodder for soundscape ecologists. It is notable for the striking

fidelity of its illustrations, photographs, and the ease with which they provide identification of buffers, barriers, J

screens, and plant lists. Plant lists offer ideas for readers to consider for optimum landscape privacy design-

Scientific and common plant names are included in each category of the listings. Also, an extensive index cross

indexes scientific plant names and common names. In addition, conversion tables, plant hardiness zones and

resource names/websites are included at the end of the text.

Why wait to develop innovative ways to turn your outdoor space into a peaceful retreat?

— Kqy M. Stansbery, Ph.D„ Library volunteer,

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, U.sM

LOMATIUM BRUNSFELDIANUM:

A NEW SPECIES OF LOMATIUM (UMBELLIFERAE) FROM NORTHERN IDAHO

Richard P. McNeill

Luna Community College

P.O. Box 2843

Las Vegas, New Mexico 87701, U.S.A.

ABSTRACT

h Fork del Canon del rio Clearwater, C

a lo largo del rio Coeur d’Aler

Lomatium brunsfeldianum Kemper & R.P. McNeill, sp. nov. (Figs. 1 - 5 ). Type: UNITED STATES. Imho. Idaho Co.:

Plants caulescent perennial, 37-86 cm tall at maturity; root cylindric, fleshy, branched, simple to branched

caudex. Stems 1-several, deliquescent, shallowly corrugated, glabrous to densely papillate, 10-30 cm long.

Leaves 1-several per stem, alternate; petiole sheathing stem; blades trullate, glabrous to densely papillate,

ternate-pinnately dissected, 2-6 times divided, 13-33 cm long, 17-48 cm wide; ultimate segments 30-71 per

1 cm 2 , filiform or linear, with mucronulate apices, 0.73-4.12 mm long, 0.14-0.62 mm wide. Scapes 1-2, 4.4-

40-1 cm long. Inflorescence compound umbel, sparsely to densely papillate; involucre absent; rays 5-16,

Journal of the Botanical Research Institute of Texas 6(1)

Journal of the Botanical Research Institute of Texas 6(1)

papillate, divaricate, 1.2-12.8 cm long; pedicels 4-19, caespitose, 2.9-10.1 mm long; involucel bractlets ±

present, free, linear or filiform, up to 4 mm long, up to 0.2 mm wide. Flowers sepals 5, connate, inconspicU* j

ous; petals 5, yellow obcordate; anthers 2, yellow, exserted; filaments flattened, yellow; superior ovary, car-

pel 2, styles 2. Mericarps narrowly elliptic, deplanate, 10.75-15.63 mm long, 3.06-4.61 mm wide, lateral

wings 0.29-0.99 mm wide; vittae interval 1-2, commissural 2-6, may be obscure, or incomplete; stylopodi*

yellow, smooth to paleaceous; carpophore yellow, 9-14 mm long. Flowering March through June. Occurring j

on wet rocky outcrops and talus slopes.

Additional specimens examined: U.S.A. IDAHO. Co.: Blackrock Canyon, Lochsa River, ca. 0.4 mi E of Split Cr. Along HWY 12, on

rock out crop, ca. 1800 ft (046 o 121'16.33"N, 115°24'10.85" W); 30 May 2003, Tyson Kemper 93 (ID); North Fk. Clearwater River, SSW facing

cliff complex adjacent to road, some seepy spots (046°50.422’N, 115°33.317'W; 046°39.080'N, 115°31.88rW); 16Jun 2004, TysonKemper266

OD); Clearwater Range Just S of mouth of Tumble Creek W of Lochsa River, E aspect (T34N R8ESWAof the SE V4 Sec. 25), 24 May 1993,

Karen Gray 27 (ID) Kootenai Co • growing on mossy rock out-crops NW side of Rose LK (N 047°33.586', W 116 0 27.64T); 23 June 2004,

Tyson Kemper 287 (ID)- rock cliff complex on Rd. 108 ca. 7 mi E of Couer d’Alene SSE aspect (N 047°42.957\ W 116“40.082* 23 Jun 2004,

Tyson Kemper 288 (ID); W margin of Rose Lake Bog, growing on a dry mossy cliff immediately above West Shore Rose Lake Road, 17 Jul

2002, Curtis R. Bjork 6481 (ID). Shoshone Co.: Couer d’Alene River, !

loos « rocky soil (N 047°37.225*. W 116°12.833'); 24Ju

Consequently, it is home to many rare and endemic species, and many disjunct populations of species that are

normally distributed either west of the Cascade Mountains or east of the Rocky Mountains (Brunsfeld et al.

2001). In 2003, an anomalous population of Lomatium was discovered in the Lochsa River Canyon by Tyson

Kemper. Additional populations were later found on the North Fork of the Clearwater, St. Joe and Couer

d’Alene rivers. It was not possible to key specimens from these populations reliably in the Flora of the Pacific

Northwest or the Intermountain Flora (Cronquist et al. 1994; Hitchcock et al. 1961). Morphologically speci-

mens from these populations were found to be somewhat similar to both Lomatium bicolor (S. Watson) J.M.

Coult. & Rose var. bicolor and L. grayi (J.M. Coult. & Rose) J.M. Coult. & Rose var. grayi (Fertig 2000). It was

concluded, however, that differences in morphology, ecology, and/or geography were significant enough that

these populations should be recognized as a new species, L. brunsfeldianum. The characters distinguishing L.

brunsfeldianum from L. bicolor var. bicolor and L. grayi var. grayi are discussed below.

The morphological characters of Lomatium brunsfeldianum easily distinguish it from other Lomatium spe-

cies, with the exceptions of L. bicolor var. bicolor, and L. grayi var. grayi. Lomatium bicolor var. bicolor is similar

in leaf structure, and mericarp size and shape. It also has overlap in all of the size measurements with L.

brunsfeldianum, but with a much smaller statistical mean. There are four main characters that allow the two

species to be separated. The most obvious is the large separation in geographical range. There is approximately

400 km separation between the known occurrences of the two taxa. The root structure is also very different; L.

bicolor var. bicolor has moniliform, globose, or tuberous roots that are un-branched and somewhat woody,

while L. brunsfeldianum has cylindric, fleshy roots that may have multiple branchings (Harris & Woolf Harris

2003). Lomatium brunsfeldianum occurs on rocky outcrops and talus slopes, which is very different from the

wet meadows and high clay content soils of L. bicolor var. bicolor habitat. Lomatium brunsfeldianum has a very

distinctive umbel structure with numerous (5-16), long (12-127 mm), divaricate rays, while L. bicolor var. bi-

color has fewer (2-9), shorter (7-81 mm), caespitose rays. When Schlessman (1984) described L. bicolor var.

bicolor, he included some samples of L. brunsfeldianum as L. bicolor var. bicolor. There were few specimen of L.

brunsfeldianum collected at that time, which made it difficult to separate from L. bicolor var. bicolor. Lomatium

grayi var. grayi is distinguished from L. brunsfeldianum by the structure and size of the mericarps, a stout tap-

root, lack of papillae in the umbel, and it is essentially acaulescent, with the leaves apparently basal, while L.

brunsfeldianum is caulescent. Lomatium grayi occurs on sites that are much dryer than sites where L. brunsfeld-

ianum is found and generally at lower elevations in north Idaho. Lomatium bicolor var. leptocarpum is very dif-

ferent from L. brunsfeldianum, with the only similarity being the mericarps. It has been considered in this paper

because it is the only variety of L. bicolor that occurs in north Idaho.

Lomatium brunsfeldianum has only been found in four deep river canyons of northern Idaho: the Lochsa

River Canyon, the North Fork of the Clearwater River Canyon, the St. Joe River Canyon, and the Couer d’Alene

River Canyon (see Fig. 1). Lomatium brunsfeldianum occurs only on moist rocky outcrops, talus slopes, and soil

at the base of cliffs in the river valleys and canyons in the mesic cedar/hemlock forest of northern Idaho be-

tween 480-1800 m in elevation. The soil it occurs on in the St. Joe River canyon was classified as an Udorthent,

which are young well-mixed soils with no horizon development, a high portion of rock fragments, a low clay

fraction, and an udic moisture regime (Soil Survey Staff 2003). The sites where it occurs have a south aspect and

are possible ground water discharge areas. Lomatium brunsfeldianum flowers from March to early June and the

fruit matures in late June through August with timing largely dependent on elevation and exposure. Lomatium

brunsfeldianum occurs in association with the following species: Alnus rubra Bong., Amelanchier alnifolia

(Nutt.) Nutt. ex. M. Roem., Centaurea stoebe L. ssp. micranthos (Gugler) Hayek, Claytonia cordifolia S. Watson,

Collinsia parviflora Lindl., Fragaria vesca L., Holodiscus discolor (Pursh) Maxim., L. ambiguum (Nutt.) J.M.

Coult. & Rose, L. dissectum (Nutt.) Mathias & Constance var. multifidum (Nutt.) Mathias & Constance,

Mimulus clivicola Greenm., M. guttatus DC., Orobanche fasciculata Nutt., Penstemon w ilcoxii Rydb., Philadelphus

lewisii Pursh, Pinus ponderosa C. Lawson, Poa bulbosa L., Pseudotsuga menziesii (Mirb.) Franco var. menziesii,

and Sedum stenopetalum Pursh.

tidemic to the Northern Rocky Mountains of Idaho and should be consid-

ered for lif tin g as a sensitive or threatened species due to its limited geographic range. It occurs on the north

side of river drainages, which is also where most of the major roads are located. This puts L. brunsfeldianum at

risk in a number of ways: encroachment by invasive species introduced by the initial road building and spread

by current road maintenance and traffic, habitat destruction caused by additional road building, and eradica-

tion through herbicide application. Future efforts should focus on a more thorough mapping of this and other

plant species in Idaho and integration of this data into management plans.

ACKNOWLEDGMENTS

I would like to thank Pam Brunsfeld, Ronald L. Hartman, Mark Schlessman, Curtis Bjork, and Matt Parks for

their help with this research. The following herbaria kindly assisted with loans: Stillinger Herbarium (ID)

University of Idaho, Moscow; Rocky Mountain Herbarium (RM) University of Wyoming, Laramie; University

of Montana (MONTU) Missoula; Missouri Botanical Gardens (MO) St. Louis; Montana State University

(MONT) Bozeman; Colorado State University (CS) Fort Collins. I would also like to thank Mary Ann Feist and

Ronald L. Hartman for helpful reviews and Barney Lipscomb for his help and patience in getting this paper

d P.S. Soltis. 2001. Comparative phylogeography of northwestern North America:

Integrating ecological and evolutionary processes in a spatial

context. Oxford: Blackwell Science. Pp. 319-339.

Cronquist, A, N.H. Holmgren, and P.K. Holmgren. 1994. Intermountain flora: vascular plants of the intermountain west,

U.SA, Vol. 3: part A subclass Rosidae (except Fabales). New York: New York Botanical Garden Press, Bronx.

Fertig, W. 2000. Lomatium bicolor var. bicolor — Wasatch Biscuitroot. State species abstract, Wyoming Natural Diversity

Database.

Harris, J.G. and M. Woolf Harris. 2003. Plant identification terminology

Payson, Utah.

Hitchcock, C.L., A. Cronquist, M. Ownbey, and J.W. Thompson. 1961. Vascular plants of the Pacific Northwest, |

Saxifragaceae to Ericaceae. University of Washington Press, Seattle.

“ V 1 984 - Systematics of tuberous Lomatiums (Umbelliferae). Syst. Bot. Monogr. 4:1-55.

f. 2003. Keys to soil taxonomy. United States Department of Agriculture, Natural Resources

d glossary. Spring Lake Publishing,

» Conservation

MICONIA CORDIERI, A NEW SPECIES OF MICONIA SECT. SAGRAEA

(MELASTOMATACEAE) FROM THE MACAYA BIOSPHERE RESERVE, HAITI

Gretchen M. lonta

Walters Judd

Department of Biology

University of Florida

Gainesville, Florida 3261 1-8525, U.S.A

Department of Biology

University of Florida

Gainesville, Florida 3261 1-8525, U.S.A.

wjudd@botany.ufl.edu

RESUMEN

While conducting a systematic revision of the Caribbean species of Miconia Ruiz & Pavon section Sagraea

(DC.) Ionta, Judd & Skean, two specimens collected in 1984 and 1989 in the Parc National Pic Macaya, Massif

de la Hotte, Haiti and identified as Ossaea curvipila Urb. & Ekman ( Miconia curvipila (Urb. & Ekman) Ionta,

Judd & Skean) were examined and determined to represent an undescribed species in the section. Species of

Miconia sect. Sagraea are easily differentiated from other Miconia species by their axillary, four-merous flowers

with well-developed calyx teeth (Judd 1986a, 1989), and the presence of minute, subsessile to short-stalked,

furrowed gland-headed hairs on vegetative surfaces (Ionta et al. 2012). Until recently Sagraea was recognized

as a genus within Miconieae (Judd 1989; Liogier 2000) or treated as a section within Clidemia (Cogniaux 1891).

A number of recent phylogenetic analyses (Michelangeli et al. 2004, 2008; Goldenberg et al. 2008; Martin et al.

2008) confirmed the monophyly of Sagraea, but at the same time revealed that nearly all of the genera of

Miconieae, including Miconia, itself, are polyphyletic (Michelangeli et al. 2004, 2008; Goldenberg et al. 2008;

Becquer-Granados et al. 2008; Martin et al. 2008). The most plausible solution to this classificatory problem (as

clearly evident in Figs. 1-3 of Goldenberg et al. 2008, or Fig. 1 of Michelangeli et al. 2008) is to place most spe-

cies of Miconieae (including Sagraea species) within a greatly expanded Miconia, comprising the molecular-

supported clade within Miconieae that can be diagnosed by the synapomorphy of berry fruits (Ionta et al.

2012). Thus we describe this new species as Miconia cordieri.

Miconia cordieri is morphologically most similar to the la Hotte endemic M. woodsii (Judd & Skean) Ionta,

Judd & Skean, and to a complex of species that includes the la Hotte endemics M. curvipila, M. lanceifolia (Urb.)

Ionta, Judd & Skean, M. rubisetulosa Ionta, Judd & Skean, and the Hispaniolan endemic M. ellipsoidea (Urb. &

Ekman) Ionta, Judd & Skean; species delimitations and relationships within this “M. ellipsoidear species com-

plex are being investigated by the first author, and a consideration of these delimitations is beyond the scope of

this paper. Miconia cordieri, M. woodsii, and the members of the M. ellipsoidea complex have a number of fea-

tures in common, most notably the presence of minute, subsessile to short-stalked pseudopeltate hairs with

flattened branches on abaxial leaf surfaces, which are likely homologous to the short-stalked, furrowed gland-

Journal of the Botanical Research Institute of Texas 6(1)

headed hairs typical of (and putatively synapomorphic for) most other species of Miconia sect. Sagraea. These

distinctive, flattened hairs are also present in a group of related species (differentiated from the previously

mentioned species by their square, winged stems) that includes M. h ottensis lonta, Judd & Skean, M. navifolia

Ionta, Judd & Skean, M. capillaris (Sw.) M. Gomez (the hairs peltate in M. capillaris), and M. tetraptera (Cogn.) |

lonta, Judd & Skean (Ionta et al. 2012), and may prove to be synapomorphic for this entire group of species, j

Miconia cordieri, exhibiting a distinctive combination of morphological characters, satisfies the defini- ■

tions of the morphological-phenetic (Judd 2007) and diagnostic (Wheeler & Platnick 2000) species concepts, j

The most distinctive feature differentiating Miconia cordieri from M. woodsii and from species in the M. ellipsoi-

dea complex is the presence of simple, erect, multiseriate hairs (0.1-03 mm long) scattered sparsely on the J

veins and on the abaxial leaf lamina (arising from veinlets). Simple hairs are absent from the abaxial leaf sur-

faces of M. woodsii, and in the species of the M. ellipsoidea complex they are present only on quaternary veins

and higher, and the hairs tend to be longer (e.g., from 0.3-0.7 mm long in M. ellipsoidea, and to 1.3 mm long in

M. rubisetulosa) and more densely distributed. The pattern of simple hairs on the stems of these plants mirrors

that of the abaxial leaves, i.e., simple stem hairs are absent in M. woodsii, short erect hairs are scattered on the

stems of M. cordieri, and more densely distributed, longer hairs are typical of species in the M. ellipsoidea com-

plex. Dendritic hairs, which are present on the stems of many species of Miconia sect. Sagraea, can be diagnos-

tically important. The stems of M. cordieri and M. woodsii are densely covered with sessile to stalked globular

dendritic hairs (to 0.2 mm long in M. cordieri, to 0.1 mm long in M. woodsii); these hairs are also scattered on

the major abaxial veins, particularly along the midvein adjacent to the petiole. Dendritic hairs on the steins of

species in the M. ellipsoidea complex are more sparsely arranged than in M. cordieri and M. woodsii, and tend to

be sessile to very short-stalked (occasionally to 0.1 mm long in M. rubisetulosa ).

The leaves of Miconia cordieri are morphologically similar to those of M. woodsii and species in the M. el-

lipsoidea complex. In these species, the leaves are more or less elliptic (often elongate to narrowly ovate in some

species), bullate (to varying degrees within and among species), with scattered, stiff setae on the adaxial sur-

face, serrulate margins, and adaxial surfaces that tend to turn dark maroon when dry. In M. cordieri and spe- :

cies in the M. ellipsoidea complex, abaxially raised percurrent intertertiary veins are common. In M. woodsii,

percurrent intertertiary veins are uncommon, and when they occur they are indistinct and not raised abaxially.

Inflorescence morphology of Miconia cordieri is similar to that of M. woodsii and most species in the M.

ellipsoidea complex from the la Hotte region, i.e., few-flowered, compact inflorescences (e.g., up to 5 flowers in

inflorescences up to 4.5 mm long in M. cordieri, and 7 flowers in inflorescences up to 8 mm long in M. woodsii),

the exception being M. rubisetulosa, which has up to 12 flowers in longer, less compact inflorescences to 24 mm

long. Floral traits that differentiate M. cordieri and M. woodsii from species in the M. ellipsoidea complex include

hypanthium shape and size (± cylindrical, flaring distally, and to 1.6 mm long in M. cordieri and M. woodsii, vs. •

urceolate and to 2.5 mm long in members of M. ellipsoidea complex), indumentum (low warty projections pres- j

ent on proximal half of hypanthium in M. ellipsiodea complex are absent in M. cordieri and M. woodsii), and

ornamentation of the inner hypanthium surface and the lower rim of hypanthium tube (the lower rim in M j

cordieri and M. woodsii appearing scalloped, with 8 low triangular projections corresponding to the insertion

points of the stamens and extending down the inner hypanthium and forming low ridges, and the lower rim of j

the hypanthium tube ringed with a fringe of inwardly-oriented hairs, vs. inner hypanthium surface and rim

glabrous, smooth, and un-omamented in species of the M. ellipsoidea complex).

Other floral features distinguishing Miconia cordieri from M. woodsii include the presence of stout, coni-

cal, somewhat curved hairs to 0.3 mm long on the hypanthium (absent in M. woodsii, and multiseriate hairs of

species of M. ellipsoidea complex longer and thinner, and not conical, when present), elongate, gland-tipp^

hypanthium hairs (absent in M. woodsii, but common in species in the M. ellipsoidea complex), and ovary posi-

tion (Vi inferior in M. cordieri, Vi inferior in M. woodsii, and mostly inferior, except for the collar, in other spe-

cies, but we note that data are based on only ten collections, and ovary position tends to vary infraspecificalfy j

the Gran Ravine du Sud <

: the Macaya ;

Biosphere Reserve (Sergile et al. 1992; Woods & Ottenwalder 1992), an extraordinary area of species ende-

mism in the Massif de la Hotte (Haiti) that continues to yield new angiosperm species (see Ackerman &

Whitten 2010; Judd et al. 2008). Miconia cordieri is known only from a few isolated fragments of original vegeta-

tion in cut-over moist broadleaved forests (i.e., rak bwa) on rough “dog-tooth” limestone at the southern base of

Mome Formon, from 950 toll90 m on the ridge just below (south) of Ville Formon and between Ville Formon

and the “Experiment Station” on the Deron Plain (west-northwest of Formon). For a detailed map showing the

locations of Morne Formon, the Deron Plain, and the karstic ridge just S of Ville Formon see Woods & Harris

(1986, Map 4). The vegetation of the Formon region (sometimes spelled Formond) is described in Ekman

(1928), Judd (1987) and Judd et al. (1990, 1998).

The Massif de la Hotte, the westernmost mountain range on the Tiburon peninsula of Haiti, harbors

many interesting species of Melastomataceae (see Judd 2007; Skean 1993). Other melastomes occurring in the

vicinity include Calycogonium apiculatum Urb. & Ekman, C.formonense Judd, Skean & Clase, C. torbecianum

Urb. & Ekman, Clidemia umbellata (Mill.) L. O. Williams, Mecranium birimosum (Naudin) Triana, M. haitiense

Urb., M. revolutum Skean & Judd, Mericmia brevipedunculata]udd & Skean, Miconia pyramidalis (Desr.) DC., M.

rubrisetulosa, M. subcompressa Urb., M. tetrastoma Naud., and Tibouchina longifolia (Vahl.) Baill. ex Cogn.

The discovery of Miconia cordieri brings to 15 the number of described species of Miconia sect. Sagraea

known from the Mome Formon-Pic Macaya region of the Massif de la Hotte (see Ionta et al. 2012). Of these

species, 10 are endemic to Hispaniola, and nine are endemic to the Massif de la Hotte.

Shrub to 1.5 m tall. Indumentum of minute, multicellular, subsessile pseudopeltate globular-dendritic hairs

with flattened and coalesced branches, multicellular, sessile to stalked elongate-dendritic hairs to 0.6 mm long,

simple, erect, multiseriate hairs to 0.4 mm long, and broad based, curved, simple, multiseriate hairs from 0.1-

0.2C-0.3) mm long. Young twigs to 2.2 mm wide, rounded-quadrangular in cross section, becoming terete with

age, the young stems densely pubescent, with sessile to stalked, globular-dendritic hairs to 0.2mm long (oc-

casionally to 0.3 mm long in interpetiolar regions), and scattered, simple, erect, multiseriate hairs to 0.4 mm

long, the stems becoming glabrous with age; intemodes 0.6-6.5 cm long. Leaves with petiole 1.5-9.3 mm long,

± rounded-triangulate, with hairs similar to those of the stem, elongate-dendritic hairs to 0.6 mm long often

present adaxiaUy, the petiole hairs persistent; blade 1.3-5.7 cm long, 0.7-2.7 cm wide, leaf pairs occasionally

slightly to moderately anisophyllous, the blades elliptic, falcate, with discrete areas bounded by adjacent ter-

tiary and quaternary veins adaxially convex, thus the leaves often bullate, the blade coriaceous, the apex acute,

± straight to slightly acuminate, culminating with a small, blunt mucro, the base ± symmetric, obtuse to

rounded to slightly cordate, the margin plane, serrulate; largest teeth to ca. 0.2 mm, hairs on teeth incurved, to

ca. 0.5 mm long; venation acrodromous, basal, with prominent midvein and 4 secondary veins, 2 conspicuous

secondary veins positioned 1.1-4 mm from margin, and 2 inconspicuous secondary veins 0.1-0.4 mm from

margin, numerous percurrent tertiary veins oriented sub-perpendicular to midvein and 0.9-3 mm apart, per-

current quaternary veins common, the higher order veins orthogonal-reticulate; adaxial surface glossy green

(dark maroon when dried), initially with scattered minute, subsessile, pseudopeltate globular-dendritic hairs

with flattened and coalesced branches, but these soon shed, elongate dendritic hairs (to 0.6 mm long) common

at midvein particularly towards the leaf base and scattered on secondary and tertiary veins, and with broad-

based, curved, simple, multiseriate hairs from 0.1-0.2 (-03) mm long, clustered in groups of one to six in the

center of convex areas bounded by tertiary and percurrent quaternary veins, the midvein, major secondary and

tertiary veins strongly to only slightly impressed, the percurrent quaternary veins moderately to slightly im-

Journal of the Botanical Research Institute of Texas 6(1)

pressed, the surface smooth; the abaxial surface pale green (brownish-green when dry), with scattered, simple, |

erect, multiseriate hairs to 0.3 mm long on midvein, secondary to quaternary veins, and even on lamina, where

they originate from veinlets, and minute, scattered subsessile, pseudopeltate hairs with fused, flattened

branches, and sessile to short-stalked dendritic hairs to 0.1 mm long restricted to quaternary and higher order

veins, the midvein strongly raised, the major secondary and tertiary veins moderately raised, and the percur-

rent quaternary veins slightly raised, the higher order veins ± flat. Inflorescences axillary, in one or both axils

of leafy nodes, occasionally more than one inflorescence per axil; glomerate cymes with up to 5 flowers, lack-

ing major branch-pairs, ca. 4.5 mm long, 3 mm wide, inflorescence branch segments to 0.9 mm, the ultimate

Journal of the Botanical Research Institute of Texas 6(1)

branches (pseudopedicels) to 0.5 mm long, inflorescence branches sparsely pubescen

dendritic hairs and occasional, simple, erect, multiseriate hairs to 0.4 mm long; pedum

similar indumentum, occasionally with an additional pair of bracts mid-peduncle, tl

early deciduous, narrowly triangular, to 1.5 mm long, 0.5 mm wide, 1

peduncle segment densely pubescent, with globular dendritic hairs similar to stem hairs; each inflorescence I

branch associated with a pair of persistent, narrowly triangular, linear bracts, 0.6-1 mm long, 0.3-0.5 mm

wide, the bracts with globular-dendritic hairs similar to those of inflorescence branches abaxially and glabrous I

adaxially, and with sparse elongate-dendritic hairs to 0.3 mm long along nodal region between bract-pairs; i

each flower subtended by 2 persistent, narrowly triangular to linear bracteoles to 1 mm long, ca. 0.3 mm wide, I

with sparse hairs similar to those of inflorescence branches abaxially, glabrous adaxially, the apices acute.

Flowers in dichasia, pedicels absent to 0.1 mm long. Hypanthium ± cylindrical, slightly flaring distally, 1-5-1.61

mm long, free portion ca. 1 mm long, the outer surface with scattered sessile to sub-sessile, globular-stellate

hairs, scattered, minute globular to elliptic dark brown glands, simple, erect, multiseriate gland-headed hairs

to 0.6 mm long, and simple, erect, stout conical hairs to 0.3 mm long, the inner surface glabrous. Calyx lobes 4,

at anthesis 0.3-0.4 mm long, 1.1— 1.3 mm wide, broadly triangular to rounded, the outer surface with hairs

similar to hypanthium, the inner surface glabrous, the apex rounded to slightly acuminate, the margin entire

to slightly erase; calyx teeth 4, 0.9-1.2 mm long, 0.15-0.2 mm wide, narrowly triangular with a ± terete distal

portion, with an acute apex, hairs similar to those of hypanthium; the calyx tube ca. 0.1 mm long, inner surface *

glabrous, the lower rim appearing scalloped, with 8 low triangular projections corresponding to the insertion |

points of the stamens, and ringed with a fringe of inwardly-oriented hairs to 0.1 mm long. Petals 4, ca. 2 mm

long, 0.4-0.5 mm wide, narrowly triangular, glabrous, the apex acute, slightly cupped, with or without a min-

ute abaxial projection, the margin entire. Stamens 8, not seen. Ovary 4-loculate, Vi inferior, ovoid to narrowly

ovoid, 1.2-1.4 mm long, 1.1-1.2 mm wide, with cylindrical apical collar encircling style, 8-lobed and distinctly |

ridged, with a crown of fine hairs to 0.1 mm long at the lobe apices; style and stigma not seen. Placentation

axile; ovules numerous. Fruits not seen, presumably globose, blue berries.

Distribution and Ecology. — Miconia cordieri is endemic to Bwa Formon and Bwa Deron, on the high pla- ,

teau with karstic hills south of Mome Formon in the Massif de la Hotte, growing at 950-1200 m. The two

collections of this rare species were found growing in disturbed moist broadleaved forests on limestone. I

Phenology.— The flowering period is poorly known and has been documented only in November.

Etymology .— The specific epithet refers to Dan and Tia Cordier, in acknowledgment of their important

logistical assistance during W. S. Judd’s 1984 fieldwork in the Massif de la Hotte, which resulted in the first |

collection of this species. Without their critical support, this fieldtrip and the subsequent trip to the same re-

gion byJ.D. Skean, Jr. (then a graduate student working under the direction of W. S. Judd) would not have been

ence (see Guerrero et al. 2004; Judd 1986b, c; Judd & Skean 1987a, b; 1993; Judd et al. 2008; Skean 1993, 2000;

Skean & Judd 1986, 1988a, b) and facilitated the documentation of the diverse flora of this region (Judd 1987). |

Formon, S of Mome Formon, 950-1040 m, 23 Jan 1984, W.Judd & D. Dod3469 (FLAS, EHH, NY).

ACKNOWLEDGMENTS

This research was supported, in part, by National Science Foundation Grant BSR-0818399. The staff of the

University of Florida Herbarium (FLAS) is gratefully acknowledged, especially Norris Williams, Kent Perkins

and Trudy Lindler, for their help in processing specimen loans. Thanks also to the staff of the Electron

Microscopy & Bioimaging Laboratory at the Interdisciplinary Center for Biotechnology Research, University

of Florida. We thank Charles A. Woods, who organized (and supported, through his grants) the fieldtrips to

the Massif de la Hotte in 1984 and 1989 during which the specimens of Miconia cordieri were collected. In ad-

dition, Paul Paryski, Sonny Parafina, Roy Voss, and Genness McBride provided field assistance on the 1989

trip, and Donald Dod assisted in fieldwork in 1984. Finally, we thank Darin Penneys and Taylor Suita® j

Quedensley for their helpful comments.

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logenetic studies in Pachyanthus (Miconieae, Melastomataceae). Bot Rev. (Lancaster) 74:37-52.

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Goldenberg, R., D.S. Penneys, F. Almeda, W.S. Judd, and F.A. Michelangeu. 2008. Phylogeny of Miconia (Melastomataceae): pat-

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38:150-161.

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Skean, J.D., Jr. and W.S. Judd. 1 988b. A new Salvia (Labiatae) from Hispaniola. Brittonia 40:1 6-2 1 .

Skean, J.D., Jr. and W.S. Judd. 1 993. A note on S

). Moscosoa 7:199-200.

Wheeler, Q.D. and N.I. Platnick. 2000. The phylogenetic species concept (se

R. Meier, eds. Species concepts and phylogenetic theory: a debate. Columbia University Press, New York. Pp. 55-69. I

Woods, C. A. and L. Harris. 1986. Stewardship plan for the national parks of Haiti. Prepared for USAID/Haiti under con-

tract 521-01 69-C-00-3083-00. Florida Museum of Natural History, and School of Forest Resources and Conservation, ;

University of Florida, Gainesville.

Woods, C.A. and JA Ottenwalder. 1992. The natural history of southern Haiti. Prepared for USAID/Haiti under contract

521-0191-A-00-7107. Florida Museum of Natural History, Gainesville.

A NEW SPECIES OF VACHELLIA (FABACEAE: MIMOSOIDEAE) FROM HAITI

David S. Seigler

Department of Plant Biology

Ricardo Garcia

Jardin Botdnico Nacional

Urbana, Illinois 61801, US. A

Milciades Mejia

Jardin Botdnico Nacional

“Dr. Rafael Ma. Moscoso “

Santo Domingo, DOMINICAN REPUBLIC

Santo Domingo, DOMINICAN REPUBLIC

John E. Ebinger

Emeritus Professor of Botany

Eastern Illinois University

Charleston, Illinois 61 920, U.S.A.

ABSTRACT

liti, de donde ha sido descrii

RESUMEN

I. Seigler, es una especie que se puede encontr;

i especies afines de Haiti y de la Republica Doi

During the course of field work in Departement Ouest, Haiti, shrubs of a previously undescribed species were

collected. Most closely related to four other species of the genus Vachellia from the Dominican Republic (Clarke

et al. 2009; Seigler et al. submitted), V. barahonensis (Urban & Ekman in Urban) Seigler & Ebinger, V. cucuyo

(Bameby & Zanoni) Seigler & Ebinger, V. oviedoensis (R. Garcia & M. Mejia) Seigler & Ebinger, and V. azuana

R. Garcia, T. Clase, Ebinger, & Seigler, ined., and one from Haiti, V. caurina (Bameby & Zanoni) Seigler &

Ebinger, this new species occurs in the central part of Haiti. This taxon seems most closely related to V. baraho-

nensis, but it is clearly distinctive and is proposed as a new species.

Vachellia koltermanii R. Garcia, M. Mejia, Ebinger, & Seigler, sp. nov. (Fig. 1). Type: HAITI. Quest: 13 km al none de

Shrub to 4 m tall; bark unknown; twigs dark gray to dark reddish-brown, terete, not flexuous, glabrous to

minutely puberulent; short shoots, 0.5-1.3 mm long, present above the stipular spines; prickles absent, leaves

alternate, also clustered on short shoots, 2.5-15.0 mm long; stipular spines light reddish-brown, becoming

gray with age, symmetrical, terete, straight, not inflated, 1-5 x 0.3-0.7 mm near the base, glabrous to minutely

puberulent; petiole adaxially grooved, 1.0-2.3 mm long, glabrous to minutely puberulent; petiolar gland soli-

tary, located just below the pinna pair, columnar, 0.2-0.7 mm long, apex circular, 0.2-0.6 mm across, de-

pressed, glabrous; rachis adaxially grooved, 0-13.5 mm long, minutely puberulent, lacking glands between

the pinna pairs; pinnae (commonly 1 pair of pinnae on leaves from the short shoots) 2 to 6 pairs per leaf, 6-15

mm long, 2-6 mm between pinna pairs; paraphyllidia absent; petiolules 0.2-0.6 mm long; leaflets 12 to 32

pairs per pinna, opposite, 0.2-0.5 mm between leaflets, linear, 1.8-3.5 x 0.5-0.8 mm, lightly puberulent and

with scattered minute glands, lateral veins not obvious, 1 vein from the base, base oblique, margins entire, apex

obtuse; midvein central. Inflorescence a densely-flowered globose head, 6-9 mm across, probably solitary

from the short shoots; peduncles 3-6 x 0.2-0.5 mm, puberulent; receptacle slightly enlarged; involucre 4 to

5-lobed, located near the base of the head, persistent; floral bracts spatulate, 0.3-0.6 mm long, sparsely pu-

berulent, persistent. Flowers sessile, yellow; calyx 5-lobed, 0.4-0.7 mm long, glabrous; corolla 5-lobed, 1.6-2.3

e, leaflets, b. petiolar gland, c

mm long, lightly puberulent, lobes one-quarter the length of the corolla; stamens 17-26; stamen filaments

3.0-4.0 mm long, distinct; ovary lightly pubescent, stipe absent to 0.3 mm long. Fruits oblong, 23-35 x 7-10

mm, slightly curved, nearly terete in cross section, not constricted between the seeds, coriaceous, not striate

but wrinkled, glabrous, eglandular, indehiscent, dark reddish-brown to black; stipe 0.3-0.7 mm long; apex

obtuse, not beaked. Seeds unknown.

Some of these data were obtained from a mass of spider-webs attached to one of the specimens (ILL) i°

which detritus including a few leaflets and peduncles with an involucre and floral bracts attached to the recep-

tacle, and a few Vachellia flowers were imbedded.

Flowers. — May-July.

Chromosome number . — Not determined.

Distribution . — Dry thorn-scrub vegetation in Departement Ouest, Haiti. This species occurs at low and

intermediate elevations (to 400 m), where there is a definite dry season.

Etymology . — Named after Duane Kolterman, Herbario, Universidad de Puerto Rico, Mayaguez, one of the col-

lectors and a student of rare and endangered species of the Greater Antilles.

SeigleretalJ

KEY TO SPECIES OF VACHELL1A FROM THE DOMINICAN REPUBLIC AND HAITI

c. Legumes inflated, nearly terete in cross section, seeds in two rows (biseriate) or irregularly arranged Vachellia farnesiana

c. Legumes not inflated, mostly flat in cross section, seeds in a single row.

d. Legumes less than 60 mm long, not constricted between seeds.

e. Mostly two pairs of pinnae per leaf; leaflets 0.6-2.0 mm wide, glabrous, lateral veins obvious Vachellia <

e. Pinnae 2 to 6 pairs; leaflets 0.5-0.8 mm, lightly puberulent, lateral veins not obvious Vachellia kolte

ACKNOWLEDGMENTS

The authors wish to thank Kanchi Gandhi for advice concerning questions of nomenclature and Scott Zona

and an anonymous reviewer for helpful comments. Nonetheless, the views (and errors) in this manuscript are

our own and do not necessarily reflect their judgments. We wish to acknowledge support by the National

Science Foundation (NSF 04-15803) and the University of Illinois Research Board (1994, 2001). We wish to

acknowledge support by the National Science Foundation (NSF DEB 04-15803) and by the American

Philosophical Society (1992). We wish to thank Alexa E. Musgrove for the preparation of drawings.

REFERENCES

Clarke, H.D., D.S. Seksler, and J.E. Ebinger. 2009. Taxonomic revision of the Vachellia acuifera species group (Fabaceae:

Mimosoideae) in the Caribbean. Syst. Bot. 34:84-101 .

Seksler, D.S., R. Garcia, T. Clase, and J.E. Ebinger. Submitted. A new species of Vachellia (Fabaceae: Mimosoideae) from the

Dominican Republic. Novon.

BOOK REVIEW

David Small and Ella May T. Wulff. 2008. Gardening with Hardy Heathers. (ISBN 13: 9780881927825, hbk.)J

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). $39.95, 296 pp„ 1 line drawing, 214 color

photos, 17 maps, 2 charts, 7"x 10".

The authors ask the question, “Why Grow Heathers?” in the opening chapter. The answer is convincingly sup-|

plied in following chapters. The reader is drawn into the fascinating and diverse world of the large group of

plants collectively called heathers. Heathers occupy several genera, including Calluna, Erica, Daboecia, and

This book offers extensive information on care and cultivation of heathers, as well as propagation meth-

ods, guidance on designing a heather garden, and uses of companion plants. The authors share their knowl-

edge of the popularity of heather breeding while clarifying the muddy waters of heather nomenclature. 1

Many of the heathers originated in the Northern Hemisphere. Their preferred climate is temperate with

annual rainfall over 40 inches. Many types are found at altitudes over 1,000 feet in elevation and in alpine en-

vironments. Because heathers thrive in areas with poor soils, near coastal regions, in windy locations, and in

boggy soils they are one of the most adaptable groups of plants, useful in a wide variety of garden applications. :

Heathers are most readily associated with two types of ecosystems: the heaths and moorlands in Western Eu-

rope. Here they grow mixed with grasses, forbs, other shrubs, and occasionally trees to create essent ia l habitat

for many animals, especially birds.

Humans have also found many uses for heathers: as material for making rope, baskets, brooms, bedding,

fences, animal fodder, as well as ingredients in traditional medicines and dyes. One of the first uses of heather

was as a fuel source. These plants have also been used for building material, primarily as the source of thatch

for roofing. However, sod houses (called turves in Scandinavia) were widely constructed from heathers. Our

modem word “turf is derived from “turve.”

Care and cultivation of heathers are discussed in detail. While heathers are found in harsh and marginal

environments, the perception that they require no special care when cultivated is erroneous. Heathers can be

successfully grown in hot summer climates if afternoon shade is provided. Likewise, they require winter pro-

tection when grown in zones colder than their rated tolerance. Proper soil conditions are also essential for

growing heathers. A soil that is acidic, high in organic matter, well draining yet moisture retentive is desirable, j

Heathers cannot survive if planted in clay.

The authors describe pruning as the least understood aspect of heather culture. These plants benefit from

annual pruning that removes spent flower stems. Pests that harm heathers are also discussed. Rabbits, hares,

and other rodents can cause damage, though heathers are not preferred by deer. Other pests include the

heather beetle, red spider mite, vine weevil, and parasitic nematodes.

In the design discussion, careful planning for site preparation is emphasized. Many heather-themed gar-

dens strive to express a natural or informal effect, but heathers are well suited to frequent pruning, so they arc

also used in formal knot gardens. While heathers are generally thought of as low-growing shrubs or ground C0*|

ers, the authors provide information on taller varieties (tree heaths) which can add height to garden plantings.

The book includes photographs and maps of species ranges. Methods for production of cut flowers and

techniques for drying heathers are described. The last chapter, “Heathers for Special Uses” is an excellent refer-

ence for growers, providing lists of heathers with similar traits and care needs.

—Jackie Peel, Volunteer,

Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, Texas 76107-3400, VSM

CALATHEA RHIZANTHOIDES AND C. PEREGRINA (MARANTACEAE),

NEW SPECIES ENDEMIC TO PANAMA

Helen Kennedy

UCR Herbarium

Department of Botany and Plant Science

ABSTRACT

a. Calathea rhiztmthoides i

ier leaf blades (9.1-16.5 vs. 4-8 cm), the

2-4(-9) leaves per shoot. Calathea pereg-

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 and recently Kennedy (2011: 201) re-

ported a total of 59 species of Marantaceae from Panama. Currently 65 species of Marantaceae (ca. 182% in-

crease from the 1945 Flora of Panama treatment) are recognized from Panama of which 47 are in the genus

Calathea. Seventeen species are e n de mic, including the two described herein. Calathea, which is restricted to

the Neotropics, is the largest genus within the Marantaceae with approximately 286 species, comprising

roughly 57% of the Neotropical Marantaceae species. The majority of Calathea species are South American

with Brazil (at least 95) and Ecuador (at least 66) being the most diverse. For the whole of Central America,

including Mexico, there are but 75 currently recognized species of Calathea.

TAXONOMIC TREATMENT

sH. Kenn .sp. nov. (Fig. 1). Tm: PANAMA. Panama. Kii. lt> on (lie road lo Carti (El Llano-Carti road).

Plants rhizomatous, perennial herbs, 36-55 cm high; the inflorescence may terminate a single-leaved shoot

but more commonly is borne on a leafless shoot directly from the rhizome; cataphylls herbaceous, narrowly

ovate, obtuse, apiculate, minutely puberulent along margins, innermost cataphyll 6-15 cm. Leaves 2-5 from a

single rhizome, each leaf borne separately from the rhizome, or a leaf may subsequetly develope in the axil of

the cataphyll subtending an inflorescence; leaf sheath not auriculate (apical margins of sheath not extended

above junction of sheath with petiole), green, subglabrous except at very base and onto node sericeous, (1-)

ca. 1.3 x 0.25 c

, ca. 13 x 3 mm.

basal leaves present, details unknown; leaf sheath not auriculate when petiole present, auriculate (apical mar-

gins of sheath extended above junction of sheath with petiole), if petiole absent, green, densely tomentose api-

cally, especially the auriculate portion and along margin in upper 1-2 cm, sides and back subglabrous to gla-

brous, basalmost 7-8 mm sericeous, 8.5-18 cm in cauline leaves; petiole green, when present, bearing a groove

adaxially, sparsely tomentose just below pulvinus, glabrous basally, (0-)0.6-1.8 cm in cauline leaves; pulvinus

elliptic in cross-section, densely tomentose in a band adaxially, the rest glabrous, hairs to 0.6 mm, 0.25-0.4 cm

in cauline leaves; leaf blade herbaceous, elliptic, apex acuminate, base obtuse, shortly abruptly attenuate,

15.5-18.5 x 4.8-6.5 cm in cauline leaves (length:width ratios 2.45-3.3:1), lateral veins 11-14 per 3 cm, cross

veinlets 33-40 per 5 mm (veins measured at midpoint of each side of blade, ruler at right angles to veins), vein

angles from midrib 38°-47°, measured at midpoint of blade, adaxial surface green, glabrous except apical 2 cm

less 0.3-0.5 mm, abaxial surface light grey-green, minutely pilose along veins, hairs 0.1-0.25 mm, additionally

apical two thirds, midrib pilose throughout, hairs to 0.4 mm in acumen. Inflorescence terminal, 1 per shoot (a

new shoot may develop in the axil of the leaf subtending the inflorescence), imbricate, ellipsoid, ca. 5 x 2.3 cm;

peduncle green, glabrous, ca. 5 cm. Bracts ca. 8, spirally arranged, herbaceous, lowermost bract broadly trans-

verse ovate, apex obtuse with acumen, upper ones broadly ovate, apex obtuse, margin not recurved, 1.3-1 .8 x

1.2-2 cm, each bract subtending 6 or more flower pairs, abaxial surface green, very minutely pilose near mar-

gins and apex (30x magnification), hairs ca. 0.1 mm, central and basal portions glabrous, adaxial surface

densely pilose apically; bicarinate prophyll membranaceous, elliptic, apex obtuse to rounded, glabrous or very

sparsely pilose on sides at apex, 1.5-1.6 x 0.8-1 cm, 0.6-0.7 cm wide, carina to carina; secondary bract mem-

branaceous, elliptic, apex obtuse to rounded, glabrous, ca. 1.6 x 0.8 cm; bracteoles 1 per flower pair, claviculate,

medial, 1.7-1.9 cm. Flowers open spontaneously, white. Sepals membranous, elliptic, obtuse to 90°, glabrous,

1.6— 1.8 x 4—5 mm. Corolla tube glabrous, ca. 22 mm; corolla lobes subequal, elliptic, glabrous, ca. 11—12 x

3 5-4.5 mm. Staminodes 3; outer staminode subrotund, apex rounded, ca. 9 x 8 mm; callose staminode

spathulate, petaloid apically, apex rounded, ca. 9 mm; cucullate staminode ca. 5 mm; stamen with lateral pet-

aloid appendage, ca. 1 mm wide, anther 2.5 mm; ovary glabrous, ca. 2.5 x 1.5 mm. Capsule unknown.

Distribution. — Calatheaperegrina is known only from the type, collected near Masargandi, Darien Prov.,

DISCUSSION

Calathea peregrina belongs to Calathea section Breviscapus Benth. It is most closely related to C. portobelensis,

both having basal and cauline leaves, the inflorescence borne on the leafy shoot, spirally arranged green bracts,

claviculate bracteoles and white flowers. It differs from C. portobelensis in the narrower angle of divergence of

the lateral veins from the midrib, 38°-47° vs. 46°-62°, the more widely spaced lateral veins, 11-14 vs. 17-22 per

vs. 2 lateral, membranous bracteoles in addition to a medial, claviculate bracteole per flower pair.

Etymology. — The specific epithet peregrina, meaning foreign, exotic or strange is in reference to the spe-

cies turning up as the joker in the deck, something foreign or strange. It was only in going through a pile of

Calathea portobelensis specimens to compare with another similar species, that the specimen of C. peregrina

struck one as clearly the odd one out.

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

CALATHEA CAROLINEAE (MARANTACEAE),

A NEW SPECIES ENDEMIC TO HONDURAS

Helen Kennedy

UCR Herbarium

Department of Botany and Plant Science

University of California Riverside

Riverside, California 92521, U.S.A

ganders@maii.ubc.ca

RESUMEN

Currently 18 species of Marantaceae are recognized for Honduras, 10 in the genus Calathea, including the new

species described herein. The related, C. matudae (Kennedy & Ganders 2011: 59), had been previously identi-

fied as C. atropurpurea, a species now recognized as endemic to western Mexico. Thus, the total number of

species has only increased by one, but that species, C. carolineae, is the only Marantaceae species endemic to

Honduras.

TAXONOMIC TREATMENT

Calathea carolineae H. Kenn., sp. nov. (Fig. 1 ). Type: HONDURAS. Cortes: W of San Pedro Sula, Sierra de Merenddn, Parque

National Cusuco, El Cortecito Camp-site, valley bottom in montane rain forest, 1305 m, Grid ref. 0361758 1716496, 17 Jul 2008, D.

vatis, a C. micans (L. Mathieu) Kent, foliis adaxialiter minute pilosis atque habitu deciduo distinguitur.

Hants deciduous, rhizomatous, herbs, 40-60 cm. Leaves 3-5, basal; leaf sheath 11-18 cm, not auriculate, pi-

lose; petiole 1.5-14 cm, bearing a groove adaxially, pilose along groove, becoming subglabrous basally but

hairs more dense just above sheath; pulvinus 0.6-1.2 cm, densely tomentose in a narrow band adaxially, the

rest glabrous; leaf blade 10-21 x 4.5-9 cm, thin, herbaceous, elliptic, asymmetric, ratio of larger to smaller side

of blade 1.31-1.54:1, apex obtuse with acumen, acumen to 1 cm, slightly eccentric, base obtuse, shortly abrupt-

ly attenuate, lowermost the smallest, lenghtrwidth ratios 1.96-2.74:1, 10-15 lateral veins per 3 cm, 17-19 cross

veinlets per 5 mm (veins measured at midpoint of each side of blade); adaxial surface of blade green with scat-

tered minute hairs, each borne on a slightly raised basal cluster of cells, hairs 0.1-0.2 mm, more dense in acu-

men, midrib lighter green, appressed tomentose, hairs 0.3-0.5 mm; abaxial surface of blade light green, gla-

brous except acumen sparsely tomentose, hairs 0.3-0.5 mm, midrib light yellow-green, glabrous except to-

mentose along sides in basal 1-2 cm. Inflorescence terminal, 1 per shoot, usually borne above the leaves, 2-2.2

x 1 -5-1.7 cm, imbricate, turbinate to broadly ovoid; peduncle 40-56 cm, glabrous. Bracts 8-10, 1.2-1.4 x ca.0.8

cm, spirally arranged, herbaceous, broadly ovate to ovate, apex acuminate attenuate in lowermost, upper ones

acute, apex not recurved; abaxial surface of bracts cream basally, green apically (fide Kelly 12033), densely

J -*«t Res. Inst Texas 6(1): 55 - 37.2012

pilose, hairs 0.2-0.3 mm, less densely so centrally near base; adaxial surface sparsely minutely pilose apically;

bicarinate prophyll 0.7-0.9 x 0.45-0.6 cm, ca. 0.35 cm wide, carina to carina, membranaceous, elliptic, apex

obtuse, translucent cream-colored, glabrous; secondary bract ca. 0.75 x 0.4 cm, membranaceous, elliptic, apex

obtuse, glabrous. Flowers pale orange (fide Kelly 12033). Sepals ca. 10-11 x 3 mm, membranous, narrowly

elliptic, acute, glabrous. Corolla tube glabrous; corolla lobes ca. 7 x 3 mm, subequal, elliptic, apex obtuse,

margins inrolled, appearing acute, light orange, glabrous. Ovary ca. 1.5 x 1.5 mm, smooth, glabrous. Capsule

obpyramidal, glabrous, crowned by a persistent calyx.

1: HONDURAS. C

Elections near El Cortecito Camp in Parque Nacional

Cusuco.

Discussion. — Calathea carolineae belongs to Calathea section Breviscapus. It is most closely related to C.

atropurpurea Matuda and C. matudae H. Kenn. but is readily distinguished from those in lacking claviculate

bracteoles. It is further distinguished from C. matudae by the glabrous vs. tomentose abaxial leaf surface. In

dried specimens, the bract tips usually retain their green color, a feature shared with C. atropurpurea. However,

in C. atropurpurea the bract margin is distinctly recurved vs. not recurved and has 23-30 vs. 15-19 cross vein-

lets per 5 mm. All three species share the deciduous habit and presence of hairs on the upper surface of the leaf.

Small individuals of C. carolineae might be mistaken for an especially large, robust individual of C. micans.

However, C. carolineae is clearly distinguished from C. micans by the minutely pilose vs. glabrous adaxial leaf

blade (excluding midrib), the glabrous vs. pilose and generally longer, > 40 cm vs. < 32 cm peduncle.

Etymology. — The specific epithet, carolineae, is in honor of Caroline Whitefoord (BM) in gratitude for her

help at the BM herbarium during my visits over the past 40 years, her many valuable collections of Marantaceae

for Flora Mesoamericana and moreover for showing me this important Honduran material.

ACKNOWLEDGMENTS

I am grateful to Caroline Whitefoord for making the recently collected Honduran specimens available for me

to study at BM. I thank the curators of BM and MO for arranging the loan of this material to MO. The Missouri

Botanical Garden provided support for my accommodations while working in the MO herbarium (organized,

thanks to Olga Martha Montiel). The travel expenses for the trips to BM and MO were provided by Fred

Ganders. Thanks to Roy Gereau (MO) for the Latin diagnosis, to Zoila Avila for digital photos of the isotype at

HEH and to Cyril Nelson and George Pilz for their reviews and critical corrections. The holotype scan was

provided by the Missouri Botanical Garden. 1 greatly appreciate the nomenclatural input of Kanchi Gandhi

(GH).

REFERENCES

Journal of the Botanical Research Institute of Texas 6(1)

BOOK REVIEW

David W. Hall, WiluamJ. Weber, edited by Jason H. Byrd. 2011. Wildflowers of Florida and the Southeast.

(ISBN: 978-0-615-39302-9, hbk.). D.W. Hall, Wildflower Book 8135 Meadowlark Court Melrose, FL j

32666, U.S.A. (Orders: D.W. Hall, Wildflower Book 8135 Meadowlark Court Melrose, FL 32666; Ama-

zon at http://www.amazon.com/Wiklflowers-Floricla-Southeast-David-Hall/dp/0615395023). $40 (plus

s&h: $8 USPS Media Mail, $18.95 USPS Priority Mail), 819 pp.

Florida and the southeast U.S. possess a rich and abundant flora highlighted by their wildflowers. Florida

ranks behind only Texas and California as the U.S. state with the most plant species. With this tome, the au-

thors provide both amateur and professional botanists with an essential guide for identifying the local wiU»|

flowers of the region. The authors emphasize Florida, yet many of the featured plants occur through the Gulf :

and Eastern Coastal Plains, especially North Carolina and west into east Texas.

This book covers more than 750 species with descriptions and color photographs. The area of plant oc-

currence is listed along with habitat type, the frequency of occurrence, and flowering season for each species. 5

Frequently, more than one common name is listed for each species; the first name listed being the one most

commonly recognized. The treatment is organized by flower color and these colors were determined using the

authors “best judgment.” Some subjectivity was used in grouping the plants, for example purple is listed in the

“blue” category and flowers that exhibit more than one color were grouped by the most common color for the

species. In some cases, the authors have added photographic inserts displaying color variation. The flower

color categories include red, pink, blue, orange, yellow, white, and green.

The book begins with a description of the treatment and an introduction to ecological communities that

provides a brief description of each (coastal, salt flats, coastal scrub, etc) with select pictures. Within the treat-

ment, the plant descriptions appear on one page, vis-a-vis the plant photos on the opposite page. The plant

photos are of various sizes and some species have more than one photo. These photographs by William Weber

are excellent quality and many point out key botanical characteristics useful for identification. The book closes

with a glossary of botanical terms and an index of scientific and common names.

I recommend this book to those among us who are eager to learn these regional wildflowers. The breadth

of the book makes it far more comprehensive than most wildflower guide books and that fact alone should see

it widely utilized by botanists of the southeast. The high quality photographs and botanical and ecological in-

formation make this a valuable book for learning the wildflower flora of Florida and the southeast U.S. ,J|

— Kevin Janni, 5WC A Environmental Consultants, Arlington, TX, kjanni@swca.com, and Research Associate,

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, USA j

TWO NEW ENDEMIC SPECIES OF CALATHEA (MARANTACEAE)

FROM PANAMA

Helen Kennedy

UCR Herbarium, Department of Botany and Plant Scie

University of California Riverside

Riverside, California 92521, US. A

ganders@in terchange. ubc.ca

rate leaf blade (leaflength

TAXONOMIC TREATMENT

Calathea lanibracteata H. Kenn., sp. nov. (Fig. 1). Type: PANAMA. Bocas del Toro: along old pipeline road from continental

divide, forest, 900 m, 08°48'04"N, 82°15'04"W, 27 Dec 1986, G. McPherson & J. Aranda 10170 (holotype: MO; isotype: PMA).

Plants rhizomatous, caulescent, herbs, ca 1.5 m high; stem villous, hairs pale straw-colored, 2-3 mm. Leaves

3-4 (or more) basal with a single cauline leaf borne above an elongate stem intemode, ca. 1.25 m; leaf sheath

auriculate, reddish, villous, sheath of subtending leaf 11-12 cm, others at least 50 cm; petiole reddish (fide

Hammel et al. 14721), villous, hairs 3-4 mm, 3.8-S.5 cm in subtending leaf, 32-44 cm in others; pulvinus el-

liptic in cross-section, tomentose adaxially, the rest glabrous but tomentose throughout just above junction to

petiole, 3.9-4.5 cm; leaf blade stiff, chartaceous, narrowly ovate-elliptic, apex acuminate attenuate, base acute

to 90°; 44-52 x 7.4-9.5 cm in subtending leaf, 55-71 x 8.5-10 cm in others, (lengthiwidth ratios 5.42-7.17:1)

lateral veins 25 to 31 per 3 cm (measured at mid-point of each side of blade), cross-veinlets 36 to 40 per 5 mm,

adaxial surface green, glabrous, midrib glabrous, abaxial surface light green, tomentose, hairs pale straw-col-

ored, most dense along veins, more sparse on acumen, up to 0.5 mm, midrib tomentose along sides in basal Vs,

tomentose throughout apically. Inflorescence terminal, 1 per shoot, imbricate, fusiform, 8.3-14 x 3-3.7 cm;

peduncle reddish, villous, 1.2-10.3 cm. Bracts 17-28, spirally arranged, central portion stiff, hardened, mar-

gins and apex soft herbaceous, elliptic, apical 7-10 mm of bract surface concave, forming a shallow channel

centrally, apex truncate to rounded, apex slightly spreading, margin in apical half undulate, undulations ca.

8-10 per cm, 4.5-5.8 x (1.3-)1.8-3 cm, each bract subtending up to 3 or more flower pairs, abaxial surface of

bracts reddish, villous throughout or with hairs sparser centrally and basally, adaxial surface villous at apical

abrous in basal third; bicarinate prophyll membranaceous, ovate, apex acute, few

sparse hairs to 1.5 mm on sides near junction with carina, the rest glabrous, 2-2.3 x ca, 1.1 cm, 0.5-0.8 cm

wide, carina to carina; secondary bracts and bracteoles absent. Flowers open spontaneously, yellow through-

out ( Hammel et al. 14721) or yellow and red ( McPherson & Aranda 101 70). Sepals membranous, narrowly linear-

elliptic, apex obtuse, margins inrolled appearing acute, glabrous, 18-20 x ca. 4 mm. Corolla tube with few

sparse hairs 1.5 to 2 mm in apical half, less sparse toward lobes, 29-34 mm long; corolla lobes subequal, ellip-

tic, obtuse, with scattered pilose hairs, hairs 0.1 to 0.25 mm, 16-18 x 4-5 mm. Staminodes 3; outer staminode

obovate, emarginate, 10-12 x 9-10 mm; callose staminode totally callose, apex obtuse with minute acumen,

ca. 15 mm; cucullate staminode ca. 6 mm; stamen with lateral petaloid appendage to 1 mm wide, extending to

basal quarter of the anther; anther 3 mm; ovary sericeous apically, hairs straw-colored, 1.5-3 mm, glabrous

basally. Capsule unknown.

Additional specimens examined: PANAMA. Bocas del Toro: Hill just S or Chiriqui Grande, at end of pipeline access road 2 mi N of 2nd large

Sanders 14721 (MO).

Distribution and habitat. — Calathea lanibracteata is

Prov. Bocas del Toro, both from along the pipeline a

s road, from 350-900 n

Discussion. — Calathea lanibracteata belongs to the informally treated “ Calathea lanicaulis Group” as out-

lined in Flora of Ecuador (Kennedy 1988:47). These species are perennials characterized by their habit of sev-

eral basal leaves with a single cauline leaf, subtending the inflorescence(-s) of spirally arranged bracts, borne

above an elongate stem internode. Calathea lanibracteata is most closely related to C. allenii Woodson and C.

confusa, sharing a similar habit, similar inflorescence and bract morphology, and the absence of both second-

ary bracts and bracteoles. Calathea lanibracteata differs from C. allenii and C. confusa by the narrowly ovate-

elliptic vs. ovate leaf blade (leaf length to width ratio 5.42-7.17:1 vs. 1.57-2.83:1), narrower leaf blades (7.4-10

vs. 12-18 cm wide), apex acuminate attenuate vs. obtuse with acumen, base acute to 90° vs. obtuse, the villous

(hairs 3-4 mm) rather than glabrous or minutely tomentose petioles, and the villous rather than glabrous to

pilose bracts.

Etymology.— The specific epithet, lanibracteata , is in reference to the distinctly hairy bracts.

Haec species a Calathea alien

8-14 vs. 2.5-4.5 mm) atque v

:nis tessellatis laxiorobis (27 ad 34 v

. 42 ad 47 per 5 mm) distinguitur.

Plants rhizomatous, caulescent herbs, 1-1.3 m. Leaves 3-5 basal with a single cauline leaf borne above an

elongate stem internode, 30-55 cm; leaf sheath green, appressed tomentose at apex and along margin, the rest

glabrous except very base sericeous, 8-15.5 cm in subtending leaf, 18-26 cm in basal leaves; petiole green,

glabrous but occasionally minutely appressed tomentose, absent in subtending leaf, 16-41 cm in basal leaves;

pulvinus yellow-green, sparsely tomentose adaxially with the rest glabrous to minutely tomentose throughout,

1-8-3 cm in subtending leaf, 3.3^47 in basal leaves; leaf blade coriaceous, major veins raised, felt as ridges,

ovate, apex obtuse with acumen to slightly acuminate, acumen 1.5-3 cm, usually curved to the left, base ob-

tuse, shortly attenuate, 22-32 x 12-16 cm in subtending leaf, 40-49 x 16-18 cm in basal leaves, (length:width

ratios 1.57-2.83:1) lateral veins 20 to 27 (to 29) per 3 cm (measured at midpoint of each side of blade), (6-) 8-14

mm between major veins and commonly 7 minor veins between the major veins, cross-veinlets 27 to 34 per 5

mm, adaxial surface green, tomentose along major veins and throughout near right (inner coiled portion of

blade) margin and acumen to nearly glabrous with sparse hairs along major veins only at base and on acumen,

midrib tomentose to glabrous, abaxial surface grey-green, tomentose throughout to glabrous except sparsely

tomentose near margin, midrib yellow-green, glabrous to densely tomentose, hairs 0.8-2 mm. Inflorescence

terminal, 1 per shoot, imbricate, fusiform, 12-17.5 x 3.5-5.1 cm in flower, to 11 cm wide in fruit; peduncle

Journal of the Botanical Research Institute of Texas 6(1)

green, 1-2.3 cm. Bracts 18-30, spirally arranged, central portion firm, margins and apex thinner, herbaceous,

elliptic, apex emarginate, the two lobes of the apex overlapping, margin undulate apically, 6-8.5 x 2-3.4 cm,

abaxial surface of bracts yellow-green, glabrous centrally, right margin appressed pilose, left margin only

sparsely pilose, adaxial surface densely appressed pilose apically, just below the lobes, lobes sparsely pilose

3.3 x 0.9-1.1 cm, ca. 0.5 cm wide, carina to carina; secondary bracts and bracteoles absent. Flowers open

spontaneously; pedicel of flower pair sericeous, hairs 2 mm. Sepals membranous, narrowly elliptic to sublin-

ear, apex acute, translucent whitish to transparent, glabrous, 30-36 x 3-4 mm. Corolla tube faint cream-or-

ange, pilose, hairs to 1.5 mm, 38-40 mm; corolla lobes subequal, elliptic, acute to 90°, pale yellow, pilose, more

densely so than on tube, 14—15 x 3-4 mm. Staminodes 3; outer staminode bright yellow, ca. 11 mm; callose

staminode yellow, ca. 13 mm; cucullate staminode 8-9 mm; anther ca. 3 mm. Ovary villous, 4-4.5 x 1.5-2

mm. Capsule unknown; seed black, aril white (fide Nee 7270).

Additional specimens examined: PANAMA. Colon: Rio Guanche, above the bridge on Portobelo Road, ca. 3-5 km above bridge, 50-200 m,

8Jul 1976, T. B. Croat 36967 (MO); Rio Guanche, ca. 1.5 mi up-stream from the bridge, lowland wet forest, c. 10 m, 23 Aug 1972, H. Kennedy

Distribution and habitat. — Calathea confusa is endemic to Panama. It occurs along the Atlantic slope in

Comarca San Bias (now Comarca Cuna Yala) west to Prov. Bocas del Toro from sea-level to near 500 m in moist

to wet forest habitats. It is generally found in the shaded understory of the forest.

Discussion . — Calathea confusa also belongs to the informally treated “Calathea lanicaulis Group” as out-

lined in Flora of Ecuador (Kennedy 1988: 47). Calathea confusa is most closely related to C. allenii Woodson

sharing a similar habit, inflorescence and bract morphology, and also the absence of the secondary bracts and

bracteoles. These two sister species are so similar in general aspect, that C. confusa has been consistently mis-

taken for C. allenii by myself and others since the 1970’s. When comparing a stack of C. allenii specimens with

C. lanibracteata for differences, the difference of venation among the supposed C. allenii material became ap-

parent. Calathea confusa differs from C. allenii in the more numerous minor veins between the major, raised,

lateral veins (7 vs. 1 to 3), the greater distance between major veins ([6—] 8-14 vs. 2.S-4.5 mm) and the more

widely spaced cross-veinlets (27 to 34 vs. 42 to 47 per 5 mm). The two tend to separate out ecologically as well.

Calathea confusa occurs from sea-level to nearly 500 m in moist or wet forest habitats while C. allenii is found

from 500-1000 (-1200) m in montane or cloud forest habitats.

Etymology.— The specific epithet, confusa, is in reference to its being confused with C. allenii and also the

unusual (somewhat confusing) variability of leaf pubescence between populations.

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 and Jeany Davidse for their help in the MO herbarium, to Roy Gereau

(MO) for the Latin diagnosis, Barry Hammel for the Spanish resumen and to Gordon McPherson for helpful

suggestions, corrections, and his collections. The type scans were provided by the Missouri Botanical Garden.

I- 1988. Calathea. In: G. Harling and L

REFERENCES

Andersson, eds. Flora of Ecuador 32:1 1-191. Berlings, Arlov, Sweden.

Journal of the Botanical Research Institute of Texas 6(1)

BOOK REVIEW

Stewart McPherson and Donald Schnell. 2011. Sarraceniaceae of North America. (ISBN: 978-0-9558918-6-1,

hbk.). Redfem Natural History Productions, 61 Lake Drive, Hamworthy, Poole, Dorset BH15 4LR, ENG-

LAND, UK. (Orders: inside UK 01202 686585, outside UK +44 1202 686585, wwwredfemnaturalhisrl

tory.com, sales@redfemnaturalhistory.com). £34.99, 808 pp., 571 images.

This beautifully illustrated hefty tome (800 pages) is the second volume in the two-volume set of Sarracenia-|

ceae. The authors chose to place Sarraceniaceae of South America as the first volume, because it describes HeU-

anthphora, the largest genus of the family. It also includes introduction chapters. This one is the second volume

of the two-volume set.) Each volume can stand independently but the authors heartily recommend using them

together.

Obviously, when the reader — at first glance — leafs through this volume, he/she will be totally fascinated

by the number of exquisite color photographs accompanying the described species — and recognize instantly

that this volume is not only extremely heavy, it is too beautiful to risk taking into the field! Take your trusty

(and presumably small) camera instead, keep careful notes and identifications, and compare and identify when

you’re comfortably back at home or office!

The authors not only describe the plants, they carefully observed and recorded data re: identification of

location, nature of climate, and when possible, how the plants grow, react to natural conditions, and — espe-

cially— the relationships between plants and insects and/or small rodents.

According to the carefully observed and recorded data of the authors, some 730 species representing 17 |

genera included carnivorous species — two identified as “snap drops” and seven as “pitcher plants.”

This is an extremely well done and very valuable contribution. The authors and artists have obviously

dedicated themselves to intensive research. They have also included maps, indicating specific areas where

various species grow; they have identified historical backgrounds for many of the species; and they have in* j

eluded helpful current information about Societies, Recommended Suppliers, various specialty places, e.g,

Hampshire Carnivorous Plants; New Taxa, and a Glossary.

— Helen Jeude, Volunteer,

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, U.SM

BOOK NOTICES/REVIEW FORTHCOMING

Stewart McPherson, Andreas Wistuba, Andreas Fleischmann, and Joachim Nerz. 2011. Sarraceniaceae of South

America. (ISBN: 978-0-9558918-7-8, hbk.). Redfem Natural History Productions, 61 Lake Drive, Ham-

worthy, Poole, Dorset BH15 4LR, ENGLAND, UK. (Orders: inside UK 01202 686585, outside UK +44

1202 686585, www.redfemnaturalhistory.com, sales@redfemnaturalhistory.com). $58.00, 562 pp.,488

images.

An elegant and beautiful volume! Hiding in the appendix are four newly described species and one new com-

bination, each with a beautifully executed line drawing: Heliamphora arenicola sp. nov., Heliamphora ceracea

sp. nov., Heliamphora collina sp. nov., Heliamphora parva comb, et stat. nov., Heliamphora purpurascens sp. nov.

( ilRM I VN z " u 1 K - lNTRODU 1 ION BY Mic;hael ™-LAN- 2011. Deceptive Beauties: The World of Wild Orchids.

(ISBN: 978-0-226-98297-7. hbk ). The Universiiy of Chicago Press, Chicago, Illinois 60637, US A.

( ers: www.press.uchicago.edu). $45, 183 pp., color photos throughout, 9.75" x 9.75".

Deceivingly beautiful!

I. BotR«.lret Tens 6<1): 64. 2012

PARADRYMONIA APICAUDATA (GESNERIACEAE),

A NEW SPECIES FROM WESTERN COLOMBIA

M. Marcela Mora

John L. Clark

Department of Biological Sciences

The University of Alabama

Tuscaloosa, Alabama 35487-0345, U.S.A.

mmmorapinto@ua.edu

Department of Biological Sciences

The University of Alabama

Tuscaloosa, Alabama 35487-0345, U.S.A.

jlc@ua.edu

Laurence E. Skog

Smithsonian Institution

National Museum of Natural History

PO Box 37012

Washington, DC 2001 3-7012, U.S.A. skogl@si.edu

in the genus by the combination of spathulate leaf

RESUMEN

INTRODUCTION

The plant family Gesneriaceae is mostly tropical or subtropical with over 3500 species distributed in 150-160

genera (Weber 2004; Weber & Skog 2007). The family is divided into four subfamilies with the subfamily

Gesnerioideae found almost exclusively in the Neotropics (Burtt & Wiehler 1995; Smith & Carrol 1997).

Although different tribal arrangements are recognized in the literature based on morphological data (e.g.,

Hanstein 1854; Fritsch 1893-1894; Wiehler 1983), more recent studies based on molecular evidence divide the

subfamily Gesnerioideae into eight tribes (Weber 2004; Skog & Boggan 2007; Weber & Skog 2007). Of the

eight tribes, Episcieae is the largest and most diverse with 22 genera and an estimated 784 species (Clark et al.

2006; Clark 2009).

Paradrymonia Hanst. with 38 currently recognized species is the fourth largest genus in tribe Episcieae

after Columnea L. (200+ spp.), Drymonia Mart. (100+ spp.) and Nautilocalyx Linden ex Hanst. (ca. 60 spp.)

(Clark 2009). The genus occurs throughout the Neotropics except southeast Brazil and the Caribbean, with

centers of diversity in Colombia and Ecuador. Ongoing phylogenetic studies support that traditionally recog-

nized Paradrymonia, Nautilocalyx, and Chrysothemis are not monophyletic. The new species described here

shares the following morphological features with currently recognized members of an informal clade com-

prised of Paradrymonia: facultative epiphytic herbs; leaves clustered in a basal rosette; petioles U-shaped in

cross section; corollas trumpet-shaped to salverform; anther dehiscence longitudinal; leaf pairs anis

i-fleshy bivalved d

a by the first a

a Colombia that is described here

J. Bot Res. Inst Texas 6(1): 65 - 69 . 2

Journal of the Botanical Research Institute ofTexas6(1)

Facultative epiphyte. Stems subwoody, subquadrate, elongate and creeping with erect shoots, intemodes 3-6

cm long near base, becoming apically clustered, rhytidome glossy and tan, adventitious roots present. Leaves

opposite, unequal in a pair, the larger leaf with petioles l-2(-4) cm long, cross section U-shaped, wine-red,

densely sericeous; the blade 8.5-24 x 3.5-6.5 cm wide, spathulate to oblanceolate with denticulate margins,

base decurrent on petiole, apex abruptly acuminate to caudate, dark green above, light green below, sometimes

tinged with purple, matte (non-glossy), sericeous on both surfaces, young leaves densely sericeous; the lateral

pairs of veins 10-14(-16), departing the midrib at 40-50° angle. The smaller leaf strongly reduced, to 2 cm

long, linear-lanceolate with denticulate margins. Inflorescence a reduced pair-flowered cyme, of 1-2 flowers in

axillary clusters, the prophylls linear-lanceolate, denticulate, less than 2 cm long, reddish, sericeous; the pedi-

cels 0.8-1.5 cm long, rose-colored, sericeous. Calyx lobes subequal 15-30 x 1-2 mm, linear, sericeous, pale

green to yellowish green, with one or two subovoid teeth on the margin, the teeth sometimes in pairs. Corolla

oblique relative to the calyx, ca. 3.5 cm long, salverform with a flattened limb and spreading lobes, spurred at

base, pilose, proximal half a narrow tube, white, distal half gradually expanding with yellowish hues inside the|

tube, pilose outside, with short glandular hairs inside, the lobes 5, subequal, 0.5-0.6 cm, slightly broader than

long. Androecium with 4 stamens, included, didynamous, 1.8-2.3 cm long, the filaments adnate for up to 3

mm from the base of the corolla tube, white, glabrous, each anther apically coherent in a pair, each thecae ca. 1

mm long, the thecae dehiscent by longitudinal slits. Gynoecium with the ovary ovoid, 7 mm long, densely se-

riceous, style up to 2.5 cm long, densely glandular-pilose, stigma capitate, nectary a bilobed dorsal gland, ca. 2

mm long, entire, glabrous. Fruit a bivalved semi-fleshy (not succulent) capsule, green, and sericeous. Seeds

oblong-ovoid, tapered at both ends but more acutely at one end, 1.2-1.4 x 0.3-0.4 mm, light reddish brown

Distribution and habitat. — Paradrymonia apicaudata is endemic to Colombia and is known from the west-

ern foothills of the Cordillera Occidental in the departments of Choco and Valle. Its habitat ranges from

Tropical wet forest (Twf) to Tropical rain forest (Trf) to Premontane (P) life zones (Holdridge et al. 1971X |

Collections range from sea level to 700 m in primary forest, fragmented tracts of primary forest with few large

trees and many epiphytes, or growing on steep rocky roadside embankments.

Phenology. — Paradrymonia apicaudata has been collected in flower in October to May and in fruit in |

February to May.

Paradrymonia apicaudata is distinguished from other species in the genus by the combination of sericeous :

pubescence on the leaf blades, pedicels, and calyces; spathulate leaf blades with conspicuous caudate apices

and denticulate margins; Unear calyx lobes with subovoid teeth; and white salverform corollas. Paradrymonia

apicaudata is similar to P. ulei, an endemic species from the northeastern Andes of Peru. Both species have sal-

verform corollas with sericeous pedicels and calyces. However, Paradrymonia apicaudata differs from P. ulei by

the presence of white corollas (vs. red-orange), linear calyces (vs. lanceolate) and conspicuously caudate leaf

apices (vs. acuminate).

Etymology.— The specific epithet, apicaudata, is in reference to the apical portion of the leaf blade. The leaf

apex is elongate and “tail-like” as in the order of Amphibians, Caudata (salamanders and newts), which is de-

fined by the presence of tails as adults.

Conservation and IUCN Red List category. — Most of the specimens of Paradrymonia apicaudata are from

the Bajo Calima region in the Department of Valle on the Pacific coast of Colombia near Buenaventura. From

the 1950s until 1995, the Bajo Calima site was a timber concession to Carton de Colombia. The site was logged

on a 30-year rotational basis and during the late 1980s and 1990s the same timber company encouraged the

Journal of the Botanical Research Institute of Texas 6(1)

Paradrymonia apicaudata M.M. Mora & J.L. Clark, sp. i

Facultative epiphyte. Stems subwoody, subquadrate, elongate and creeping with erect shoots, intemodes 3-6

cm long near base, becoming apically clustered, rhytidome glossy and tan, adventitious roots present. Leaves

opposite, unequal in a pair, the larger leaf with petioles 1— 2(— 4) cm long, cross section U-shaped, wine-red, ‘

densely sericeous; the blade 8.5-24 x 3.5-6.5 cm wide, spathulate to oblanceolate with denticulate margins,

base decurrent on petiole, apex abruptly acuminate to caudate, dark green above, light green below, sometimes •

tinged with purple, matte (non-glossy), sericeous on both surfaces, young leaves densely sericeous; the lateral

pairs of veins 10-14(-16), departing the midrib at 40-50° angle. The smaller leaf strongly reduced, to 2 cm

long, linear-lanceolate with denticulate margins. Inflorescence a reduced pair-flowered cyme, of 1-2 flowers in \

axillary clusters, the prophylls linear-lanceolate, denticulate, less than 2 cm long, reddish, sericeous; the pedi-

cels 0.8-1.5 cm long, rose-colored, sericeous. Calyx lobes subequal 15-30 x 1-2 mm, linear, sericeous, pale

green to yellowish green, with one or two subovoid teeth on the margin, the teeth sometimes in pairs. Corolla

oblique relative to the calyx, ca. 3.5 cm long, salverform with a flattened limb and spreading lobes, spurred at ■

base, pilose, proximal half a narrow tube, white, distal half gradually expanding with yellowish hues inside the

tube, pilose outside, with short glandular hairs inside, the lobes 5, subequal, 0.5-0.6 cm, slightly broader than

long. Androecium with 4 stamens, included, didynamous, 1.8-2.3 cm long, the filaments adnate for up to 3

e base of the a

-, glabrous, each anther apically coherent in a pair

mm long, the thecae dehiscent by longitudinal slits. Gynoecium with the ovary ovoid, 7 mm long, densely se- 1

riceous, style up to 2.5 cm long, densely glandular-pilose, stigma capitate, nectary a bilobed dorsal gland, ca. 2 |

mm long, entire, glabrous. Fruit a bivalved semi-fleshy (not succulent) capsule, green, and sericeous. Seeds

obfong-ovoid, taPCred 3t b ° th CndS bUt m ° re acutely at one end ’ 1 - 2-1 - 4 x 0 3-0.4 mm, light reddish brown

Distribution and habitat.— Paradrymonia apicaudata is endemic to Colombia and is known from the west- i

ern foothills of the Cordillera Occidental in the departments of Choco and Valle. Its habitat ranges from

Tropical wet forest (Twf) to Tropical rain forest (Trf) to Premontane (P) life zones (Holdridge et al. 1971), 1

Collections range from sea level to 700 m in primary forest, fragmented tracts of primary forest with few large

trees and many epiphytes, or growing on steep rocky roadside embankments.

Phenology. Paradrymonia apicaudata has been collected in flower in October to May and in fruit in

February to May.

Paradrymonia apicaudata is distinguished from other species in the genus by the combination of sericeous 5

pubescence on the leaf blades, pedicels, and calyces; spathulate leaf blades with conspicuous caudate apices

and denticulate margins; linear calyx lobes with subovoid teeth; and white salverform corollas. Paradrymonia

apicaudata is similar to P ulei, an endemic species from the northeastern Andes of Peru. Both species have sal- ;

verform corollas with sericeous pedicels and calyces. However, Paradrymonia apicaudata differs from P. ulei by 1

the presence of white corollas (vs. red-orange). Unear calyces (vs. lanceolate) and conspicuously caudate leaf

apices (vs. acuminate). 7

Etymology. The specific epithet, apicaudata, is in reference to the apical portion of the leaf blade. The leaf

r^rr* 7, asm the order of A m ph ibians, Cauda, a (salamanders and newts,, whtch is de-

fined by the presence of tails as adults.

Conservation and IUCN Red List category. Most of the specimens of Paradrymonia apicaudata are front

the Bajo Calima region in the Department of Valle on the Pacific coast of Colombia near Buenaventura Front

, he 1950sun,il 1995, theBajoCaltoasi.ewasatimber concession toCandnde Colombia. The she was logged

on a 30-year rotational basts and during the late 1980s and 1990s the same timber comnanv enrnuratted the

Weber, A. 2004. Gesneriaceae. In: K. Kubitzki and J.W. Kadereit, eds. The families and genera of vascular plants. Vol. 7.

Flowering plants, dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae). Springer-Verlag, Berlin and

Heidelberg, Germany. Pp.63-1 58.

Weber, A. and L.E. Skog. 2007 (onward): The genera of Gesneriaceae. Basic information with illustration of selected spe-

cies. Ed. 2. http://www.genera-gesneriaceae.at

Wiehler, H. 1983. A synopsis of the neotropical Gesneriaceae. Selbyana6:1-219.

Journal of the Botanical Research Institute of Texas 6(1)

BOOK REVIEW

Bernardo Gut. 2008. Trees in Patagonia. (ISBN: 978-3-7643-8837-9, hbk.). Birkhauser Verlag AG, P.O. Box

133, CH-4010 Basel, SWITZERLAND. (Orders: 'www.springer.com). $89.95, 283 pp., 760 illustrations, -1

600 color photos, 9" x 12.25".

When I first started to examine this book I thought that 1 was going to be looking at something for the Patago-

nian area that would be comparable to those written by Texas authors Neil Sperry, Howard Garrett, or Sally ;

Wasowski. Rather it is something between the works by those authors and is intended for use as a field guide.

From the back cover: “This book is a guide to the native trees and approximately 95% of the introduced

arboreal species of Argentine and Chilean Patagonia. Introductory chapters convey an overview on what is j

termed ‘Patagonia’, as well as on the geology, climate, soils, and vegetation of Southern South America. Keys

based on vegetative characters and richly illustrated descriptions of more than 170 species form the core of the

manual. These chapters are followed by concise entries on afforestations, urban trees, and plantations of fruit |

trees in climatic enclaves. Renowned experts describe the most important National Parks of both Argentine

and Chilean Patagonia. The last chapter is devoted to Carl Skottsberg and his remarkable expedition through

Patagonia a hundred years ago.”

Following the chapter on Skottsberg are a number of appendices including a glossary, abbreviations, an

extensive bibliography, and the English, Spanish, and scientific plant names.

Each species is provided with a botanical description, followed by some or all of the following: status and

distribution, hardiness, uses, habitat, elevation, and remark(s). Each is illustrated by color photographs or

drawings, sometimes of the tree, its flowers, fruits or leaves.

This was intended to be a ‘field guide’ but its size weighs against that usage. It is an excellent resource and

contains a considerable amount of useful secondary information such as a good botanical key and descriptions

icapes.

r purchase by major academic libr

searchers with an interest in the region. It will be useful t

i a landscape or garden which includes trees.

Botanical Research Institute of Texas, 1700 University Dri

rticultural

BOOKS RECEIVED/REVIEWS FORTHCOMING

William A. Weber and Ronald C. Wittmann. 2012. Colorado Flora: Western Slope. A Field Guide to the Vas-

cular Plants, Fourth Edition. (ISBN: 978-1-60732-142-2, pbk.). University Press of Colorado, 5589

Arapahoe Ave., Suite 206C, Boulder, CO 80303, U.S.A. (Orders: http://www.upcolorado.eom/content/_ j

welcome, 800-627-7377). $27.95, 608 pp., 108 line drawings, 1 map, 4 tables. 5.5" x 8.5".

William A. Weber and Ronald C. Wittmann. 2012. Colorado Flora: Eastern Slope. A Field Guide to the Vas-

cular Plants, Fourth Edition. (ISBN: 978-1-60732-140-8, pbk.). University Press of Colorado, 5589

Arapahoe Ave., Suite 206C, Boulder, CO 80303, U.S.A. (Orders: http://www.upcolorado.com/content/_l

welcome, 800-627-7377). $27.95, 608 pp., 104 fine drawings, 1 map, 4 tables. 5.5" x 8.5".

Elray Nixon, Illustrated by Bruce Lyndon Cunningham. 2012. Trees, Shrubs, and Woody Vines of East Texas,

Third Edition. (ISBN: 978-0-934115-10-0, pbk.). Bruce Lyndon Cunningham (Orders: brucelc@sud-

denlink.net or call 936.569.6965). $59.95, 256 pp., 345+ b/w illustrations, 8" x 10".

DRYMONIA ATROPURPUREA (GESNERIACEAE),

A NEW SPECIES FROM NORTHWESTERN SOUTH AMERICA

Laura Clavijo

John L. Clark

Department of Biological Sciences, Box 870345

The University of Alabama

Tuscaloosa, Alabama 35487, U.SA.

lvclavijoromero@crimson.ua.edu

Department of Biological Sciences, Box 870345

The University of Alabama

Tuscaloosa, Alabama 35487, U.SA

jic@ua.edu

RESUMEN

INTRODUCTION

The flowering plant family Gesneriaceae is a member of the order Lamiales (APG III 2009) and is primarily

pantropical with extensions into the subtropics and temperate regions (Weber 2004; Skog & Boggan 2006).

The family contains ca. 150 genera and ca. 3500 species and is classified into four major groups (Weber 2004).

In the Neotropics the highest concentration of species diversity for the family is found in Colombia with 32

genera and more than 400 species (Kvist et al. 1998), followed by Ecuador with 29 genera and 240 species (Skog

& Kvist 1997), Brazil with 28 genera and 207 species (Forzza etal. 2010), and Peru with 28 genera and 150 spe-

cies (Kvist et al. 2005).

(272 species) and Besleria (200 species) (Weber 2004). It ranges from Mexico to Bolivia, including Brazil,

French Guiana, and the Caribbean. The highest species richness is found in Colombia with 31 species (Clavijo

& Clark 2008) and Ecuador with 30 species (Clark et al. 2006). Drymonia is a genus of terrestrial subshrubs,

vines or herbaceous epiphytes with campanulate, tubular or hypocyrotid flowers. Drymonia is especially di-

verse in the tropical wet forests along the western Andean slopes of southern Colombia and northern Ecuador

where there are over 35 species.

Drymonia has a wide range of morphological variation as a result of different pollinators and dispersal

mechanisms (Roalson et al. 2005; Clark et al. 2006). The morphological variation has made Drymonia a diffi-

cult genus to circumscribe. Ongoing studies on pollination biology have facilitated a better understanding of

the morphological variation that pertains to pouched flowers (bird pollinated) and campanulate flowers (eu-

glossine bee pollinated).

Drymonia atropmpurea ( l.ivijo | L t lark, sp tu>\ 'Fig. 1' I , hi \ix« i ■ .... ram-quia Aim

78°33'39" W, 695 m, 29 May 2008 (fl, fr) J.L. Clark 10443 (holotype: US; isotypes: BRIT, K, MO, NY, QCNE, UNA).

Differs from other congeners by the subshrub habit, relatively large leaves to 46 cm long, dark purple inflorescence bracts, bullale purple

dark «lyx, and angulate corolla tube.

Journal of the Botanical Research Institute of Texas6(1)

73

Terrestrial subshrub, 1-1.5 m tall. Stem erect, unbranched, quadrangular in cross-section, shallowly sulcate,

subwoody, green with red spots, glabrate basally, strigose apically, trichomes ca. 1 mm long, unbranched, yel-

low when dry; internodes (2.4-)5-15 cm long. Leaves opposite, decussate, equal in a pair; petioles 2.8-5.2 cm

long, terete, green with red spotting, base swollen with several pinkish gland-like enations, glabrate to strigil-

lose, strigose in the immature leaves, trichomes less than 1 mm long, yellow when dry; blade elliptic to oblong,

20-46 cm long, 6.3-21 cm wide, coriaceous, upper and lower surface green, sometimes lower surface green

suffused with light red, apex acuminate, base cuneate or sometimes slightly oblique, margin serrulate, becom-

ing revolute when dry, upper surface glabrous, lower surface glabrate; 6-8 lateral pairs of veins, venation raised

below, green when live, red-brown when dry, main vein strigose, secondary veins strigillose, higher order ve-

nation only evident abaxially, green when live, red-brown when dry. Inflorescence a reduced pair-flowered

cyme, 1 inflorescence per axil with 2-6 flowers; peduncle reduced to less than 1 mm long; inflorescence bracts

ca. 23 mm long, ca. 12 mm wide, dark purple, strigillose, elliptic, apex acute, margin entire; floral bract one,

10-23 mm long, 1-8 mm wide, dark purple, linear to spatulate, base decurrent, apex rounded, margin entire,

strigose to strigillose towards the apex; pedicel 16-32 mm long, red-orange, strigillose to strigose, glands scat-

tered along the pedicel. Calyx dark purple, coriaceous, bullate, persistent in fruit, apex rounded, margin en-

tire, base truncate to cordate, glabrate to strigose at the base, venation evident, the main vein strongly raised

abaxially, strigose; calyx lobes 5, 4 nearly equal, 5th lobe (dorsal) slightly smaller, lobes fused at the base for

2-4 mm, ventral lobes 18-25 mm long, 14-21 mm wide, rotund, margin occasionally involute, lateral lobes

19-28 mm long, 13-17 mm wide, ovate, margin involute apically, dorsal lobe 14-19 mm long, 8-13 mm wide,

ovate. Corolla zygomorphic, funnelform; corolla tube strongly angulate (bent) at the base, posture perpen-

dicular relative to the calyx, ca. 29 mm long, outer surface beige ventrally with some darker longitudinal lines,

red-brown dorsally, glabrous; corolla base ca. 7 mm wide, spur ca. 11 mm long, ca. 8 mm wide, white; throat

ca. 14 mm wide, light yellow, glandular trichomes on the inner surface; corolla lobes 5, subequal, red-brown,

apex obtuse, margin fimbriate, glabrous, ventral lobe longer than the other four lobes, spreading, ca. 13 mm

long, ca. 15 mm wide, orbicular, upper lobes reflexed, 6 mm long, 6-7 mm wide, orbicular, lateral lobes re-

flexed, 9-10 mm long, 8-9 mm wide, rotund. Androecium of 4 stamens, didynamous, 24-28 mm long, adnate

to the corolla tube for 9-10 mm, glabrous, staminode absent; anthers oblong, dehiscence by basal pores that

develop into longitudinal slits, 6-7 mm long, 1.5-2 mm wide. Gynoecium with a single dorsal nectary gland,

ovate, emarginate, 2.0-2.6 mm long; ovary superior, ca. 6 mm long, ca. 4 mm wide, glabrous, oblong, laterally

compressed; style ca. 13 mm long, strigillose; stigma stomatomorphic, ca. 3 mm diameter. Fruit a bivalved

laterally compressed fleshy capsule, ca. 18 mm long, ca. 14 mm wide, rounded, externally red-brown with yel-

low patches towards the apex and the base, glabrous, seeds numerous, immersed in a mass of fleshy funicular

tissue, 0.4-0.6 mm long, 0.2-0.3 mm wide, brown when dry, rhombic, covered by a transparent aril.

Distribution and habitat.— Drymonia atropurpurea is known from wet forests on the western slopes of the

Andes in northern Ecuador and southern Colombia, between 350 and 1400 meters. In Colombia it has been

found in the understory of protected cloud forests that are part of the Rio Nambi Natural Reserve in the

Department of Narino above 1000 meters. In northern Ecuador D. atropurpurea has been collected in remnant

patches of wet forest between 350 and 695 meters in the Esmeraldas Province along the San Lorenzo-Ibarra

highway.

Drymonia atropurpurea is distinguished from other congeners by the subshrub habit; large elliptic to ob-

long leaves to 46 cm long; dark purple bracts (inflorescence and floral) and calyces (Fig. 1A); bullate calyx

lobes; corolla tube strongly angulate (bent) at the base (Fig. ID); corolla posture perpendicular relative to the

calyx; and dark reddish-brown corolla lobes. Drymonia atropurpurea is similar to Drymonia turrialvae because

of their large leaves, relatively large funnelform corollas, and similar inflorescence. The two species are differ-

entiated by the subshrub habit in D. atropurpurea in contrast to the herbaceous habit in D. turrialvae; the non-

bullate leaves with the abaxial surface green in D. atropurpurea in contrast to the bullate leaves with the abaxi-

al surface wine-red in D. turrialvae; and a laterally compressed fleshy capsule in D. atropurpurea (Fig. IB) in

contrast to a globose indehiscent berry in D. turrialvae.

Journal of the Botanical Research Institute of Tei

Etymology. — -The specific epithet is Latin for dark purple: atro (=black), purpurea (=purple), in reference to

the dark purple color of the bracts and calyx.

Conservation and IUCN Red List category. — Drymonia atropurpurea is known from recently documented

populations in Ecuador and Colombia. The population in Colombia is from the well-established protected i

area, Rio Nambi Natural Reserve (Narino). The population from Ecuador is from an area that is almost com- ]

pletely deforested along the San Lorenzo-Ibarra highway. According to the IUCN Red List criteria for esti-

mated range, area of occupancy and population size (IUCN 2001), and considering the uncertain future of

habitat conservation in Ecuador, Drymonia atropurpurea should be listed in the category NT (Nearly

Threatened).

to, M. Flores & A. Vdsquez 1689 (COL, PSO). ECUADOR. Esmeraldas: ParroquiaSan

7 N, 78°35’50" W, 1 2 Feb 2003 (fl, fir), J.L. Clark 71 45 (QCNE, SEL, UNA, US).

ACKNOWLEDGMENTS

Support for this study was provided by the National Science Foundation (DEB-0841958 & DEB-0949169 to |

JLC) and the Nellie Sleeth Scholarship from The Gesneriad Society, Inc. (to LC). We thank Alain Chautems,

Christian Feuillet, Alejandro Zuluaga and Steve Ginzbarg for providing helpful reviews; the Herbario National

Colombiano (COL) for access to their collections; and the Fundacion Ecologica los Colibries de Altaquer

(FELCA), especially Alex Vasquez and Mauricio Flores, for logistical support for the 201 1 expedition to the Rio

Nambi Natural Reserve in Narino (Colombia).

REFERENCES

sification for the orders

Clark, J.L, P.S. Herendeen, L. E. Skog, and E.A. Zimmer. 2006. Phylogenetic relationships and generic boundaries in the

Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and morphological data. Taxon 55:313-336.

Clavuo, L and J.L Clark. 2008. El g4nero Drymonia (Gesneriaceae). Rapid Color Guide #244 (www.fmnh.org/plantguides). 7

Chicago Field Museum.

Forzza, R.C. (and 29 others), eds. 2010. CatSlogo de plantas e fungos do Brasil. Andrea Jakobsson Estudio/Jardim

Botanico do Rio de Janeiro.

Kvist, L.P., L.E. Skog, and M. Amaya-Marquez. 1 998. Los generos de Gesneriaceas de Colombia. Caldasia 20:1 2-28. ij

Kvist, LP., LE. Skog, M. Amaya-Marquez, and I. Saunas. 2005. Las Gesneriaceas de Peru. Arnaldoa 12:16-40.

IUCN. 2001. IUCN Red List Categories and Criteria, Version 3.1. Prepared by the IUCN Species Survival Commission.

International Union for Conservation of Nature and Natural Resources, Gland, Switzerland and Cambridge.

Roalson, E.H., J.K. Boggan, LE. Skog, and LA. Zimmer. 2005. Untangling Gloxinieae (Gesneriaceae). I. Phylogenetic patterns

and generic boundaries inferred from nuclear, chloroplast, and morphological cladistic dataset. Taxon 54:389-410.

Skog, LE. and LP. Kvist. 1 997. The Gesneriaceae of Ecuador. In: Valencia, R. & H. Balslev, eds. Estudios sobre Diversidad y

Ecologia de Plantas, Memorias del II Congreso Ecuadoriano de Botanica. Pontificia Universidad Catolica del Ecuador,

Quito, Ecuador. Pp. 13-23.

Skog, L.E. and J.K. Boggan. 2006. A new classification of the Western Hemisphere Gesneriaceae. Gesneriads 56(3):1 2-17.

Weber, A. 2004. Gesneriaceae. In: Kubitzki, L. and J.W. Kadereit, eds. The families and genera of vascular plants. Vol. 7.

Flowering plants, dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae). Springer-Verlag, Berlin#

Heidelberg, Germany. Pp. 63-1 58.

A TAXONOMIC STUDY OF THE GALAPAGOS ENDEMIC VARRONIA

(CORDIACEAE) SPECIES WITH NOMENCLATURAL NOTES

Julia K. Stutzman

James Madison University

Harrisonburg, Virginia 22807, U.S.A.

Edgar B. Lickey

Andrea Weeks

George Mason University

Fairfax, Virginia 22030, U.S.A.

Conley K. McMullen

Harrisonburg, Virginia 22807, U.S.A.

ABSTRACT

the life-history of the species.

RESUMEN

INTRODUCTION

Background

Within the past several decades, the Galapagos Islands have become increasingly exposed to threats from in-

vasive species and rapidly expanding human populations (Atkinson et al. 2010; Tapia et al. 2010). Invasive

nonnative species pose the greatest threat to the native and endemic species on the Islands (Trueman et al.

2010), and the Galapagos are particularly vulnerable to invasion by alien species (Denslow et al. 2009; Reaser

et al. 2007). Accurate identification and documentation of the flora of the Galapagos is essential for conserva-

tion efforts, particularly for those species that suffer from taxonomic uncertainty. The purpose of this study

was to review and update the characterization of the endemic species of genus Varronia and to create a more

complete identification key for them based on foliar morphology. Scientists have been unable to accurately

evaluate these species for conservation status because morphological distinctions between species do not hold

up in practice. This uncertainty may have discouraged research on these species of Varronia.

The Galapagos archipelago is composed of volcanic islands located in the Pacific Ocean with a total land area

of 7880 km 2 within a geographical area of 45,000 km 2 (Neall & Trewick 2008). The archipelago is approxi-

mately 1000 kilometers west of Ecuador and is the product of a stationary mantle hotspot (Morgan 1971); the

t Res. Inst Texas 6(1): 75 - 99.2012

crease eastward due to the movement of the Naz

From the velocity of the plate motion, scientists <

c data predict ;

Journal of the Botanical Research Institute ofTexa$6(1)

ental land mass. The ages of the individual islands in* 1

: on which the Galapagos are located (Rassmann 1997).

e that none of the islands above ocean level is older than

; recent origin of the present archipelago (White etaL

1993). The older islands are estimated to be between three and four million years old, while the younger islands

may be one to two million years old (Hickman & Lipps 1985). The flora and fauna that dispersed to the Gala-

pagos originated in various regions around the Pacific basin, including North, Central, and South America, the

Caribbean, and Asia/Australasia (Grehan 2001; Jackson 1993). The climate of the Galapagos is extremely arid |

compared to most tropical archipelagos. The Islands experience a warm season from January to May, during

which time lowland vegetation reaches its peak, and a cool season from June to December, where lowlands

remain dry while highlands remain constantly wet from a mixture of light rain and mist (McMullen 1999). f

Today, the Islands have a well-documented flora of 614 indigenous angiosperm species, of which 238 are

endemic, 314 are native and 62 putatively so. The Islands also host 825 introduced species of which 324 are

considered to be naturalized (McMullen 2011). The Galapagos Islands consist of three ecological zones that

include the Littoral (Coastal) Zone, the Arid Lowlands, and the Moist Uplands (Johnson & Raven 1973). The

Littoral Zone occupies the shoreline and is comprised of salt-tolerant vegetative species; the Arid Lowlands are

characterized by Opuntia cacti and shrubby, herbaceous species that tolerate dry living conditions much of the

year; the Moist Uplands are characterized by ferns, sedges, and evergreen tree species, including Scalesia (Mc-

Mullen 1999). Varronia species from this study are primarily found in the Arid Lowlands, but may occasion-

ally be found in the Moist Uplands.

The angiosperm family Cordiaceae is represented in the archipelago by two genera, Cordia L. and Varronia P.

Br., and seven species: C. lutea Lam., C. alliodora (Ruiz & Pav.) Oken, V. polycephala Lam., V. canescens Anderss,

V leucophlyctis (Hook, f.) Anderss., V revoluta (Hook, f.) Anderss., and V. scouleri (Hook, f.) Anderss. (Jaramillo

Diaz & Guezou 2011; Wiggins & Porter 1971). Of these seven, four are endemic (V canescens, V. leucophlyctis,

V. revoluta, and V scouleri), with the remainder either native (V polycephala) or introduced (C. lutea, C. aflio-

dora). Until recently, all species were placed in the genus Cordia, but Miller and Gottschling (2007) resurrected

the genus Varronia to circumscribe the monophyletic group of shrubby species limited to the Americas.

Varronia species are multi-stemmed shrubs with regularly serrate leaf margins and have inflorescences that are |

condensed, capitate, spicate, or if cymose, then small and few-flowered; their pollen grains are porate. By con-

trast, Cordia species are single or few-stemmed trees with leaf margins that are entire or irregularly sharply

dentate on the apical half and have inflorescences that are cymose; their pollen grains are colporate.

Varronia is an entirely New World genus with approximately 100 species r ang ing from Arizona to Argen-

tina, with the predominance of the species being limited to the frost-free sub-tropics (Miller and Gottschling

2007). The presence of Varronia in the Galapagos is likely due to long-distance seed dispersal by frugivorous

birds (McMullen 2009; Itow 2003) from western South America. Porter (1983) hypothesized that three of the

endemic species of the “ leucophlyctis-complex ” (V. canescens, V. leucophlyctis, and V scouleri) are most closely

related to V macrocephala Desv. (syn.= V polyantha Benth.), a native of Columbia and Peru, and the findings of

Weeks et al. (2010) are consistent with this hypothesis.

Of the four endemic species, V revoluta is the most easily distinguished based on its linear, revolute leaves ?

and tubular corolla. The three remaining endemic species are morphologically similar; these members of the

so-called “leucophlyctis-complex” are quite difficult to distinguish reliably. The taxonomic key produced by

Wiggins and Porter (1971) provides diagnoses based on leaf and calyx pubescence:

i and do not pro-

In practice, however, their diagnoses do not cover the full range of Varronia species variation

vide the necessary separation needed for accurate and reliable identification.

Molecular phylogenetic data have not been able to fully resolve questions regarding the origin and evolu-

tion of the endemic Varronia species. Sequence data from the nuclear and chloroplast genomes indicate a pos-

sibly reticulate evolutionary history for the endemic species, with nuclear ribosomal DNA internal transcribed

spacer (ITS) data supporting separate origins of V revoluta and the species of the leucophlyctis-complex and

chloroplast ndhF genic data supporting a single radiation event. Moreover, sequence data were not able to

distinguish separate lineages within the complex. Fossil-based calibrations of the phylogeny indicate lineages

are younger than some of the extant islands in the archipelago (< 4.5 Mya) and that the complex is likely less

than 2 Mya. This finding suggests species boundaries may be difficult to distinguish because of the relative

youth of this species complex (Weeks et al. 2010).

The difficulty in distinguishing these endemic species may have conservation implications. Currently, the one

endemic species easily identifiable, V. revoluta, is considered “Near Threatened” by the International Union for

the Conservation of Nature (IUCN), whereas V. scouleri is considered “Vulnerable” (Tye 2000c, d). A Vulnera-

ble species is one that has been categorized by the IUCN as likely to become endangered unless the circum-

stances threatening its survival and reproduction improve. A status of Near Threatened is assigned to a species

that may be considered threatened with extinction in the near future, although it does not currently qualify for

the Threatened status. Both V canescens and V. leucophlyctis are classified as “Data Deficient” due to taxonomic

uncertainty (Tye 2000a, b). Data Deficient is a category applied to a species when the available information, in-

cluding abundance and distribution, is not sufficient for a proper assessment of conservation status to be made.

Introduced species have become increasingly represented in the flora and fauna of the Galapagos Islands (At-

kinson et al. 2010). The effects of these introduced species on the indigenous plants and animals of the archi-

pelago must be monitored because so many of the species (39%) are endemic, such as the Varronia in this study

(McMullen 2011). Because of the relatively unknown impact of introduced species on many endemic species,

it is important to establish and maintain an accurately documented floral baseline for reference in future stud-

ies. Documentation of this sort, which includes updated nomenclature, taxonomic keys, and distribution

maps, will provide useful information to Galapagos researchers, park personnel, and visiting researchers as

they strive to develop sensible and effective conservation programs.

Objectives

The major objectives of this study were: 1) revise the nomenclature of the Varronia Galapagos endemics, 2)

revise the leaf, trichome, and calyx lobe morphological descriptions for each Varronia species, and 3) produce

a more reliable key for the endemic Varronia using characters that can be used in the field.

MATERIALS AND METHODS

A total of 348 herbarium specimens were examined: 133 specimens collected by CKM during trips to the Ga-

lapagos in 2005 and 2007 (JMUH) and an additional 215 loaned specimens collected between the years 1825

and 1983. A total of three specimens were obtained from the Brooklyn Botanic Garden (BKL), 49 from the

California Academy of Sciences (CAS), eight from the New York Botanical Garden (NY), 56 from Harvard Uni-

versity (GH), 26 from the University of Copenhagen (C), 55 from the Swedish Museum of Natural History (S),

and 18 from the Royal Botanic Gardens at Kew (K). Eight type specimens from K and one type specimen from

S, as well as photographs of three type specimens from the University of Cambridge Herbarium (CGE) were

studied for a total of 11 type specimens included in this study. A total of 81 specimens of Varronia canescens, 95

specimens of V leucophlyctis, 89 specimens of V. scouleri, and 83 specimens of V. revoluta were examined.

Journal of the Botanical Research Institute of Texas 6(1)

Specimens were first sorted into piles based on relative similarities in leaf shape, regardless of the identi-

fication on the label. The specimens were then examined with a dissecting microscope and the piles were read-

justed based on both leaf shape and morphological similarities in leaf pubescence. During this process, four

distinct sets of morphological leaf shape and leaf pubescence characteristics were observed, as described in the

following species narratives. In addition, measurements of leaf length and width on two leaves per specimen

were taken, and observations of calyx length and pubescence were also observed. This information is also in-

cluded in the species descriptions.

Endocarp structure of fruits from 17 different individuals was observed. Only JMUH specimens were

utilized. Few specimens from other herbaria had fruits available, and most of these specimens were consider-

ably older and in more fragile condition. Preparation of the endocarp included boiling fruits in distilled water

for 15 minutes, and soaking them in a room temperature water bath for a 24-hour period to help loosen the

fleshy material surrounding the endocarp. Any remaining material was removed with tweezers before observ-

ing the endocarp under a dissecting scope. Endocarp structure varied minimally between individuals ob-

served, and was not useful in distinguishing species. This information is therefore not included in the new

To obtain detailed images of the plants’ leaf surface morphologies, a LEO 1340 VP Scanning Electron

Microscope was used at the James Madison University SEM/EDS Regional Facility. The SEM operated with an

acceleration voltage of 25 kV, a spot size of 300 (equivalent to a probe current of 102 picoamps), and a working

distance of 25 mm. Samples of dried herbarium material were coated with gold from a sputter coater prior to

imaging, and the images obtained are secondary electron images. The specimens selected to obtain SEM im-

ages were chosen as representative samples of the species based on observations with a dissecting microscope.

In addition to work at James Madison University, a trip to the Galapagos Islands in February 2011 allowed di-

rect observation of live specimens in the field, observation of herbarium specimens at the Charles Darwin Re-

search Station (CDS), and field testing of the new taxonomic keys (Tables 1 and 2).

MORPHOLOGICAL CHARACTERIZATIONS

In the following species characterizations, all descriptions of leaf shape and size, leaf pubescence and color,

and calyx pubescence and size at anthesis are the result of this morphological study. All other descriptions are

from Wiggins and Porter (1971), and ar

1. Varronia revoluta (Hook, f.) Anders

h within each description.

revoluta (Hook f.) Borhidi,

Species description.— Varronia revoluta is a slenderly branched shrub 2-4 m in height, with dark gray-brown,

shallowly fissured bark marked by transversely elliptic lenticels; young twigs densely fine-pubescent with ap-

pressed simple hairs (Wiggins & Porter 1971). Leaves linear with strongly revolute margins, 2.5-110 cm long,

0.2-1.0 cm wide; adaxial surfaces dark green and densely pubescent, hairs white, simple, and appressed (Fig-

la); abaxial surfaces lighter green and densely villous pubescent, with more robust appressed hairs found on

the veins; midrib is markedly elevated (Fig. 2a). Inflorescence globose-capitate on a slender, terminal peduncle

1.0-5.0 cm long with appressed-pubescence (Wiggins & Porter 1971). Calyx broadly cup-shaped to nearly

globose, 2.5-3.5 mm long, about 3.0 mm wide, slightly contracted at apex, shallowly 5-lobed, only slightly

Journal of the Botanical Research Institute ofTexas6(1)

la. lb. lc. Id.

Distribution and habitat. — Endemic to the Galapagos Islands. This species is found among other shrubs

and in forests as an understory tree. It is also found on lava flows and rocky soil. It can be found at elevations

from near sea level to 1420 m. Specimens have been collected from Femandina, Floreana, Isabela, Santa Cruz,

and Santiago Islands (Fig. 4).

Specimens examined. ECUADOR. GALAPAGOS ISLANDS. Femandina Island: Apr 1899, R.E. Snodgrass and E. Heller 327 (GH 244027);

Apr 1906, Alban Stewart 3177 (CAS 27260, GH 244032); Feb 1964 John R. Hendrickson 56 (CAS 619244); Feb 1964, F.R. Fosberg 45099

749292); Feb 1965, D. Wiens 3825 (CAS 516647); Mar 1967, Inga Eliasson 1700 (S s.n.). Floreana Island: Jul 2005, Conley K. McMullen 83?

•IM1 II I *707 >. |ul K M, Mullen 8 is ( |M( H I1705<. |ul 2005. ( „„| 0 K M, M„II, * i7 ( |ML II I 170V.. |un 1S«I G &««

(GH 233528); May 1932 John Thomas Howell 9372 (CAS 468109); Feb 1964, Syu® Itow 189 (CAS 579308); Oct 1905, Alban Stewart 664 (C AS

Stutzman et al., Taxonomy of Varronia

«*440U) Kb NnT lr.il \\ ,«,«s ,mj Pun ,' an M IV, in Vo<X As 52o3n5>.Oci I- il II 1 1 '• .UK l M '>• M - n . IV. I -no

Eliasson 785 (K H2007/01159/13 S s n )• May 1967, Jnga E Hasson 2099 (S s.n.); Oct 1932, H.J.F. Schimpf/215 (S s.n.). Isabda Island: Aug 2005,

Conley K. McMullen 850 (JMUH 13729); Aug 2005, Conley K. McMullen 851 (JMUH 13732); Aug 2005, Conley K. McV

13734); Aug 2005, Conley K. McMullen 853 (JMUH 13736); Aug 2005, Conley K. McMullen 860 (JMUH 13750); Aug 2005, C

861 (JMUH 13752)- Aug 2005 Conley K McMullen 862 (JMUH 13754); Aug 2005, Conley K. McMullen 863 (JMUH 13756):

K.McMullen8490MUH849);Aug2005 1 Conl^K.McMullen^^l3^^1 8 ^G^l^M3^A^l^

MS Heat's (G^oS; M^S^n 2 and E. Heller S (GH 244023); Feb 1899, R.E. Snodg

Heller 272 (GH 244028); Aug 1905, Alban Stewart 3169 (GH 244025); Mar 1906, Alban Stewart 3173 (GH 244031); Jul 1891, G. Bai

rt 3172 (GH 244033); Jur

E. Heller 897 (GH244C

Journal of the Botanical Research Institute of Texas 6(1)

E. Heller 28 (GH 244019); Aug 1905, Alban Stewart 3170 ( CAS 27261); Apr 1974, H.H. van der Werf/1025 (CAS 606128); Sep 1975, H.H. 1

Werff 2332 (CAS 6061 29); Jun 1963, David Snow 289 (CAS 579303); Jan 1967, Ira L. Wiggins and Duncan M. Porter 207 (CAS 525322); Ja®

Ira L. Wiggins and Duncan M. Porter 231 (CAS 525339); May 1967, Inga Eliasson 2196 (S s.n.); May 1967, Inga Eliasson 2173 (S s.n.); Apt

Inga Eliasson 2013 (S s.n.); Mar 1967, Inga Eliasson 1711 (S s.n.); Mar 1967, Inga Eliasson 1548 (S s.n.); Feb 1967, Inga Eliasson 1305 (S s.r

1959, Gunnar Marling 5357 (S s.n.); Jun 1959, Gunnar Harling 5385 (S s.n.); 1825, Janies Macrae s.n. (K 449167); 1825, James Macrae s.n. if

449166); Sep 1975, H.H. van der Werff 2332, (K H2007/01159/14). Santa Cruz Island: May 1932, John Thomas Howell 9465 (NY s.n.). Santiago

Island: Jul 1905, Alban Stewart 3176 (GH 244026); Dec 1905, Alban Stewart 3175 (GH 244024)- Dec 1905 Alban Stewart 3175 (CAS 27257)'

Oct 1835, Charles Darwin s.n. (K 449161); Oct 1835, Charles Darwin s.n. (K 449162).

Nomenclatural history.— Varroma revoluta is the most distinct of the four endemic Varronia species found on

Stutzman et a!., Taxonomy of Varronia

83

the Galapagos Islands. Hooker first described this species in 1847 as Cordia revoluta Hook. f. In the same pub-

lication, Hooker also described Cordia revoluta Hook. f. var. nigricans Hook. f. and Cordia linearis Hook. f. Be-

cause Cordia linearis A.DC. was named in 1845, two years before Hooker, Hooker's C. linearis is not a validly

published name (see Art. 45.3 and Art. 53.1, McNeill et al. 2006). Porter (1980) reviewed old specimens from

Hooker, and though he did not discuss the taxonomy, he named Macrae s.n. (K 449166) as the lectotype of

Confia revoluta Hook f var. nigricans Hook. f. In observing this specimen, no obvious differences could be

found between it and the other types, and it is therefore treated as a synonym in this study. Andersson (1855)

transferred C. revoluta Hook. f. and C. linearis Hook. f. into Varronia. In 1891, Kuntze transferred V. revoluta

(Hook, f.) Anderss. into Lithocardium, and in the same publication named a new species, Lithocardium hooke-

rianum Kuntze. The type specimen for L. hoofeerianum Kuntze, which was housed in Leipzig (LZ), cannot be

located and is presumed to have been destroyed with the herbarium in 1943 during a bombing raid in World

War II (Thiers 2012). Gurke (1893) later transferred L. hookerianum into Cordia. In 1933, Fnesen transferred V.

linearis (Hook, f.) Anderss. and C. revoluta Hook. f. into Sebestena, without regard to Anderssoris earlier renam-

ing of C. revoluta Hook. f. as V revoluta (Hook, f.) Anderss. and Kuntze’s later reclassification to Lithocardium

revolutum (Hook, f.) Kuntze. More recently Borhidi (1988) moved Cordia revoluta Hook. f. back into Varronia as

Journal of the Botanical Research Institute ofTexa$6(1)

Specimen: Aug 2005, Conley K. McMullen, 848 (JMUH 1 3725).

V. revoluta (Hook, f.) Borhidi without regard to Andersson’s (1855) origin;

Therefore, Borhidi’s change is invalid (see Art. 45.3 and Art. 53.1, McNeill et

resurrected by Miller and Gottschling (2007) and because of this, the name

derss. has priority as the species name.

Contribution of this study. — After examining the types and other specimens, this species has distinctive >

linear leaves with revolute margins (Fig. 5), separating it from the other three Galapagos Varronia. Varronia

revoluta also has very consistent leaf hairs that do not vary among the specimens examined. Adaxial leaf sur-

faces are densely pubescent with distinctive white, simple hairs that are appressed with tips oriented towards

the leaf apex and/or outer margins (Fig. 6). Abaxial leaf surfaces also have simple, dense hairs. Margins are

villous pubescent with simple hairs. More robust, appressed white hairs are found on the veins, which also

bend towards the leaf apex/outer margins (Fig. 7). Calyxes are densely villous with simple hairs which are a

uniform length throughout the entirety of the calyx. Calyx lobes are much shorter than the calyx tube at anthe-

sis. The characters observed in this study agree with the description reported by Wiggins and Porter (1971> 1

However, this study provides a more complete description of the leaf hairs, which are deemed to be the distin-

guishing features among the other three endemic Galapagos Varronia species.

2. Varronia leucophlyctis (Hook, f.) Andersson, Kongl. Vetensk.-Akad. Handl. 1853:203. 1855 Cordial

cophlyctis Hook. f„ Trans. Linn. Soc. London 20:199. 1847. Type: ECUADOR. GalApagos Islands. Albemarle Island (Isabela Island*

1835, Charles Darwin s.n. (uectotype, designated by Porter 1980: CGE 00285). Paralectotypes: ECUADOR. C

): 1825, Macro

nomenclature for the species.

L 2006). Recently, Varronia was

Varronia revoluta (Hook, f.) An-

Stutzman et al.. Taxonomy of Varronia

appressed simple hairs. Stale represents 400 pm. Spedmen: Jul 2005, Conley K.

Species descriptioi

ing stems; bark d

oles, and pedunc

2.0-11.0 cm long

pubescent with n

-Varronia leucophlyctis is an open shrub 1.0-2.5 m tall, with one to several erect to ascend-

•k brownish gray, with scattered, reddish tan, almost circular lenticels; young twigs, peti-

, densely pubescent with simple hairs (Wiggins & Porter 1971). Leaves broadly lanceolate,

3 8-4 0 cm wide with finely serrate to crenate margins; adaxial leaf surfaces dark green,

stly erectandsimple hairs (Fig. lb);abaxial leaf surfaces lighter green, often denselypubes-

jstly simple hairs; simple hairs on veins are commonly robust (Fig. 2b). Inflorescence capi-

e on terminal peduncles 1.0-5.0 cm long (Wiggins & Porter 1971). Calyx broadly cup-

f campanula, 6.0 mm long, 4.0 mm wide, 4- or 5-lobed (Wiggins & Porter 1971); calyx

lobes about as long as calyx tube at anthesis (Fig. 3b); calyx pubescent with erect and simple hairs that are no-

ticeably longer on calyx lobes than calyx tube (Fig. 3b). Corolla white, usually 5-lobed, funnelform, tube 7.0-

8.0 mm long, 7.0-8.0 mm wide; lobes rotate-spreading to slightly reflexed (Wiggins & Porter 1971). See Wig-

gins and Porter (1971) for a more thorough description of the reproductive parts and fruits of this species.

Distribution and habitat.— Endemic to the Galapagos Islands. This species is found among lava boulders

and in rocky soil among other shrubs. It is also sometimes found as an understory shrub in forested areas.

Found from near sea level to 1750 m. Specimens have been collected from Espaftola, Femandina, Floreana,

Isabela, San Cristobal, Santa Cruz, and Santa Fe Islands (Fig. 4).

ECUADOR . GALAPAGOSISl^NDS-Espaaola Island: Feb 1906,^5^^

• Mar 1899 R.E. Snodgrass and E. Heller 342 (CAS 9251, GH 244008); Apr 1899, R E. Snodgrass

Stewart 3168 (CAS 27273). Fa

Hamaroi 222 (C 24/2007/22).

7, Inga Eliasson 1701 (S s.n.); Sep IS

Island: Jul 2i '

P. Schmidt 2535 (C 2

IcMullen 840 (JMUH

I); Jul 2005, Conley K. \

m 844 (JMUH

5, Conley K. McMullen, 838 (JMUH 13705).

I >7 In 1 Pei Mhn Micisv.M.Si-Kssn > P o, Nun. 81 .5 (S s n l Dec l')M«.%itli l isv.n^ l Sv„ LVc I'-’nr. Iny,; I

877 (Ss.n ). Isabela Island: Aug 2005, Conley K. McMullen 859 (JMUH 13748); Aug 2005, Conley K. McMullen 857 (JMUH 13745); Aug 2005,

13739); Mar 2007, Conley K McMullen and Michael Shane Woolf 881 (JMUH 13797); Mar 2007, Conley K. McMullen and Michael Shane W<4

877 (JMUH 13788); Mar 2007, Conley K. McMullen and Michael Shane Woolf 878 (JMUH 13790); Mar 2007, Conley K. McMullen and Mich‘S

Shane Woolf 879 (JMUH 13792); Mar 2007, Conley K. McMullen and Michael Shane Woolf 880 QMUH 13794); Mar 1906, Alban Stewart 3159

(GH 244014); Mar 1906, Alban Stewart 3164 (GH 233526); Mar 1899, R.E. Snodgrass and E. Heller 195 (GH 244007); Feb 1899, R E. Snodgrass

and E. Heller 291 (GH 233534); Jun 1899, HE. Snodgrass andE. Heller 893 (CAS 9260, GH 244011);Jun 1899, R.E. Snodgrass and E. Helkr85J

(GH 244013);Jun 1899, R E. Snodgrass andE. Heller 881 (GH 244012); Aug 1891, G. hour 212 (GH 233532);Jul 1891, G. Baur210 (GH 23353®

no date, R.E. Snodgrass and E. Heller 136 (GH 244009); Dec 1898, R.E. Snodgrass and E. Heller 75 (GH 244010); Apr 1932 John Thomas Howell

8976 (CAS 468107); Mar 1906, Alban Stewart 3161 (CAS 27270); Jan 1967, Ira L. Wiggins and Duncan M. Porter 232 (K H2007/01 159/9, CAS

525340); Mar 1906, Alban Stewart 3160 (CAS 27272); May 1932 John Thomas Howell s.n. (CAS 468103); May 1932 John Thomas Howell 9431

(CAS 468111); Apr 1974, H.H. van der Werff 1037 (CAS 606063); Apr 1974, H.H. van derWerff 1033 (CAS 606062); Jun 1959, C,unnarHarM

5622 (S s.n.); Jun 1959, Gunnar Harling 5360 (S s.n.); Jun 1959, Gunnar Harling 5312 (S s.n.); Apr 1967, Inga Eliasson 2034 (S s.n.); Mar 1967,

Inga Eliasson 1713 (S s.n.); May 1967, Inga Eliasson 2219 (S s.n.); May 1967, Inga Eliasson 2208 (S s.n.); Feb 1967, Inga Eliasson 1285 (S s.n.)i F*

1967, Inga Eliasson 1303 (S s.n.); Sep 1972, Michelle and Ole Hamann 2243 (C 24/2007/24); Jul 1972, Michelle and Ole Hamann 1780 (C

24/2007/20); Jul 1972, Michelle and Ole Hamann 1771 (C 24/2007/19); Jul 1972, Michelle and Ole Hamann 1709 (C 24/2007/18); Jul 1972, Mi-

chelle and Ole Hamann 1615 (C 24/2007/17); Sep 1972, Michelle and Ole Hamann 2320 (C 24/2007/12); 1825 James Macrae s.n. (K 449163);

1825 James Macrae s.n. (K 449164); Apr 1932 John Thomas Howell 8976 (KH2007/D1159/15). San Cristobal Island: Feb 1967, Ira L. Wigg&

and Duncan M. Porter 379 (CAS 526675, GH 233549). Gw. Cruz Apr 1930, H.K. Svenson 7 (GH 233525-1)- Feb 1964, Ira L. Wi0 ni

(BKX8B51 nT 26 ’) ^ ^IdyW m ^ 233531)1 ^ 1938 ’ ^ ^ ^ 262726) ’ 1933 ’ JP Chapi " ^

1966, Inga Eliasson 514 (S s.n.); Apr 1972, P. Pritchard 1197 (C 24/2007/13); Mar 1981, O. Hamann andO. Seberg 1772b (C 24/2007/16); MS’

lO.Sebergl]

A (CAS 27266, GH 1

Nomenclatural history.— Hooker (1847) first described this species as Cordia leucophlyctis Hook. f., moved b>

Andersson (1855) into Varronia, then transferred into Lithocardium by Kuntze (1891). In 1855, Andersson de-

scribed a new species, Varronia scaberrima Anderss.; however, as Varronia scaberrima Bert, ex Spreng. was al-

ready in existance, V. scaberrima Anderss. is not valid (see Art. 45.3 and Art. 53.1, McNeill et al. 2006). In 1891

Kuntze moved both Varronia P. Br. and Cordia L. into Lithocardium Kuntze; however, Kuntze renamed V. scab

errima as Lithocardium galapagosenum Kuntze, because the epithet “ scaberrima ” was already occupied by L

scaberrima, which had priority. Lithocardium galapagosenum was later moved by Gurke (1893) into Cordia, anc

was considered taxonomically synonymoui

Gottschling’s (2007) resurrection of the gei

correctly recognized name for this species.

Contribution of this study. — The

cens or V. scouleri. The leaf shape ten

but this is not entirely consist*

icophlyctis u

a be broadly lanceolate with finely se

be distinct from those of either V. canes-

ite margins (Fig. 8),

luta, V. leucophlyctis

ypes across all of the examined specimens. Adaxial leaf surface hairs are mostly

erect and simple (Fig. 9). Abaxial leaf surfaces are often densely pubescent, with mostly simple, erect hairs;

simple, larger hairs can be found on the veins (Fig. 10). Very rarely, forked or stellate hairs may be found on

either leaf surface of V leucophlyctis. Calyx hairs are also erect and simple, and noticeably longer on the calyx

lobes that on the calyx tube. The calyx lobes themselves are about as long as the calyx tube at anthesis.

Though leaf morphology of this species is often distinct, V. leucophlyctis c

canescens and V scouleri In this situation, the hairs on the adaxial and abaxi;

ifused with both V

Journal of the Botanical Research Institute of Texas 6(1)

acter to use for identification. Neither V. canescens nor V. scouleri possesses predominately simple, erect hairs

on both leaf surfaces, and in the same abundance as found on V. leucophlyctis.

3. Varronia canescens Andersson, Kongl. Vetensk.-Akad. Handl. 1853:203. 1855. Type: ECUADOR. GaUpacos Is-

lands. Charles Island (Floreana Island): 1853, Andersson s.n. (holotype: S 04-1959!).

Species description.— Varronia canescens is an erect, moderately branched shrub 1.0-3.0 m tall, with one to

several stems from the base; bark dark brown and thin (Wiggins & Porter 1971). Young twigs densely pubes-

cent with appressed stellate, forked, and simple hairs. Leaves lanceolate, 1.3-11.8 cm long, 0.5-4.3 cm wide,

with finely serrate to crenate margins; adaxial leaf surfaces dark green, glabrous to sparsely pubescent with

hairs typically appressed or sometimes weakly erect, stellate and forked, sometimes simple (Fig. lc); abaxial

leaf surfaces lighter green, densely pubescent with appressed stellate, forked, and simple hairs (Fig. 2c). Inflo-

rescence subcapitate to short-spicate, on slender peduncles, 1.5-10.0 cm long (Wiggins & Porter 1971). Calyx

narrowly campanulate, 3.0-5.0 mm long, 4- or 5-lobed (Wiggins & Porter 1971); calyx lobes as long as or

shorter than calyx tube at anthesis (Fig. 3c); calyx appressed pubescent with stellate, forked, and simple hairs,

generally uniform length throughout (Fig. 3c). Corolla white, 5-lobed, funnelform, tube 5.0-8.0 mm at anthe-

sis, lobes rotate-spreading to reflexed (Wiggins & Porter 1971). See Wiggins and Porter (1971) for a more

thorough description of the reproductive parts and fruits of this species.

Stutzman et al.. Taxonomy of Varronia

Fk. 10. Varronia leucophlyctis abaxial leaf surface displaying robust, erect simple hairs on veins, and simple hairs in margins. Scale represents 300 pm.

Distribution and habitat .— Endemic to the Galapagos Islands. This species is found among lava boulders

and in rocky soil among other shrubs. It is also sometimes found as an understory shrub in forested areas.

Found from near sea level to 950 m. Specimens have been collected from Espaftola, Floreana, Isabela, Pinta,

Pinzon, San Cristobal, Santa Cruz, and Santiago Islands (Fig. 4). This species is the most widely distributed of

the endemic Galapagos Varronia.

Journal of the Botanical Research Institute of Texas 6(1)

:e margins. Photograph courtesy of Conley K. McMullen. Specimen: N

Nomenclatural history. — Andersson first described this species in 1855 as Varronia canescens Anderss. In 1891,

Kuntze moved both Varronia P. Br. and Cordia L. into Lithocardium Kuntze. However, the epithet “ canescens ”

was occupied by Cordia canescens Kunth which had priority, so Kuntze (1891) renamed V. canescens Anderss.

as Lithocardium anderssonii Kuntze. Subsequently, Gurke (1893) later reclassified L. anderssonii Kuntze as Cor -

dia. More recently Borhidi (1988) moved C. anderssonii (Kuntze) Gurke back into Varronia as V. anderssonii

(Kuntze) Borhidi without regard to Andersson’s (1855) original nomenclature for the species. Therefore, Bo-

rhidi’s change is invalid (see Art. 45.3 and Art. 53.1, McNeill et al. 2006). As such, V canescens Andersson

should be considered the correct name for this species based upon Miller and Gottschling’s (2007) reinstitu-

Contribution of this study. — Based on characters observed on the type specimen, this species has lanceo-

late leaves with finely serrate to crenate margins (Fig. 11), but these characters were variable when observing

all herbarium specimens. Varronia canescens has a propensity to look most similar to V. scouleri, though the

leaves of V canescens do not tend to be as long as those found on V. scouleri, nor are the leaf margins on V. cane-

scens as coarsely serrate as those generally found on V. scouleri. The pubescence on the adaxial surface of the

leaves of V canescens, when present, is typically appressed (Fig. 12) or weakly erect (Fig. 13) and composed

mostly of stellate and forked, occasionally simple hairs. The abaxial leaf surfaces are densely pubescent, with

i. Specimen: Jul 2005, Conley K.

McMullen, 824 (JMUH 13675).

appressed stellate, forked, and simple hairs (Fig. 14). The abaxial pubescence of V. canescens is the densest of

any of the four endemic species and the most obviously appressed. In addition, the hairs on the veins do not

tend to be any larger than those found in the margins, providing another character that distinguishes V canes-

cens from the other three endemic species. The pubescence on the calyx is composed of appressed stellate,

forked, and simple hairs, which are a uniform length through the length of the calyx. In contrast to the other

species, the hairs are generally very short. The calyx lobes are as long as or shorter than the calyx tube at anthesis.

4. Varronia scouleri (Hook, f.) Andersson, Kongl. Vetensk.-Akad. Handl. 1853:204. 1855. = Cordu, scouleri Hook. f„

Trans. Linn. Soc. London 20:200. 1847. Type (Porter 1980): ECUADOR. Galapagos Islands. James Island (Santiago Island): no date.

Species description.— Varronia scouleri is an ascending branched shrub, with dullish dark gray bark with circu-

lar or transversely lenticular lenticels (Wiggins & Porter 1971). Young twigs sparsely pubescent with generally

appressed stellate and forked, or sometimes simple hairs. Some noticeably larger, more robust hairs may also

be observed. Leaves lanceolate, 2.3-17.5 cm long, 0.5-5.0 cm wide, with coarsely serrate margins; adaxial leaf

surfaces dark green and pubescent with erect simple, forked, and sometimes stellate hairs (Fig. Id); abaxial leaf

surfaces lighter green and sparsely pubescent with weakly appressed stellate and forked, sometimes simple

bairs which are noticeably more robust on elevated veins (Fig. 2d). Inflorescence globose-capitate, on pedun-

cles 3.0-5.0 mm long (Wiggins & Porter 1971). Calyx globose in bud, campanulate after anthesis, 5-lobed;

lobes are 4.0 mm long, excluding elongated slender lobe tips, and 3.0-4.0 mm wide (Wiggins & Porter 1971);

Journal of the Botanical Research Institute ofTe:

elongated lobe tips may be as long as, or longer than the lobe, 4.0-8.0 med mer; calyx lobes usually longer than

calyx tube at anthesis (Fig. 3d). Pubescence on the calyx composed of erect and simple, sometimes forked or

stellate hairs. Simple hairs noticeably longer on the calyx lobes than on the calyx tube (Fig. 3d). Corolla white,

5-lobed, tubular-funnelform, with a tube 6.0-8.0 mm long; lobes short, crispate (Wiggins & Porter 1971). See

Wiggins and Porter (1971) for a more thorough description of the reproductive parts and fruits of this species.

Distribution and habitat. — Endemic to the Galapagos Islands. This species is found among lava boulders

and in rocky soil among other shrubs. It is also sometimes found as an understory shrub in forested areas. Var-

ronia scouleri is found from near sea level to 400 m. Varronia scouleri on Santa Cruz Island near the Charles

Darwin Research Station and town of Puerto Ayora are known to be found among Bursera graveolens, Opuntia

echios var. gigantea, Jasminocereus thouarsii var. delicatus, Parkinsonia aculeata, Croton scouleri var. scouleri,

Walfheria ovata, Castela galapageia, Cordia lutea, Clerodendrum molle, Scutia spicata var. pauciflora, Passijlof a

foetida, and Acacia rorudiana. Specimens have been collected only from Floreana, Santa Cruz, and Santiago

Islands (Fig. 4). This species has the most limited distribution of the endemic Galapagos Varronia species.

Specimens examined. ECUADOR. GALAPAGOS ISLANDS. Floreana Island: Dec 1966, IngaEliasson 836 (S s.n.); Feb 1928, BorghiWR^

33 (Ss.n.). Santa Cruz Island: Mar 2007, Conley K. McMullen and Michael Shane Woolf 876 (JMUH 13787); Mar 2007, Conley K. McMullen and

Michael Shane Woolf 875 (JMUH 13784); Mar 2007, Conley K.

McMullen and Michael Shane Woolf 873 (JMUH 13782); Mar 20(

Conley K. McMullen 836 (JMUH 13701); Jnl 2005, Conley K. Me

Jul 2005, Conley K. McMullen 833 (JMUH 13693); Jul 2005, C

m and Michael Shane Woolf 874 (JMUH i;

113775); Jul 2005,

832 (JMUH 13692);

7 (GH 233525); F*

Stutzman et al., Taxonomy of Varronia

as found in margins. Scale represents 150 pm. Specimen: Jul 2005, Conley K. McMullen, 824 (JMUH 13675).

Thomas Howell 9075 (CAS 468104, GH 233550, NY s.n.); Jan 1964, Syuzo Item 1 (CAS 579305); Mar 1970, Syuzo how 30800-3 (CAS 623522);

May 1970, Syuzo Itow 52200-1 (CAS 623519); Feb 1964, fra L. Wiggins 18472 (CAS 619424); Jun 1970, Syuzo how 62600-1 (CAS 623520); Feb

1964, Luis A. Fournier 157 (CAS 705256); Feb 1967, Ira L. Wiggins and Duncan M. Porter 710 (CAS 528135); Feb 1964, Syuzo Itow 45 (CAS

579313); Jim 1932, H.J.F. Schimpff40 (NY s.n.); 1939, T.WJ. Taylor s.n. (NYs.n.);Jun 1932, H.J.F. Schimpff40 (BKL 81350, S s.n.); May 1959,

Gunnar Harling 5194 (S s.n.); Dec 1966, Inga Eliasson 986 (S s.n.); Jan 1967, IngaEliasson 1192 (Ss.n.); De 1966, Inga Eliasson 985 (S s.n.); Dec

Nov 1966, Inga Eliasson 515 (S s.n.); Nov 1966, Inga Eliasson 513 (S s.n.); Jun 1972, Michelle and Ole Hamann 1598 (C 24/2007/11); Jan 1972,

Michelle and Ole Hamann 155 (C 24/2007/9); Jan 1972, Michelle and Ole Hamann 154 (C 24/2007/8); Mar 1981 , 0. Seberg 1747 (C 24/2007/7);

Apr 1972, Michelle and Ole Hamann 1177 (C 24/2007/4); Mar 1972, Michelle and Ole Hamann 726 (C 24/2007/2); Apr 1983, Phyllis S. Bentley 318

OK H2007/01159/12); Apr 1930, H.K. Svenson 7 (K H2007/01159/11). Santiago Island: Aug 2005, Conley K. McMullen 864 (JMUH 13758);

Aug 2005, Conley K. McMullen 865 (JMUH 13761); Aug 2005, Conley K. McMullen 866 (JMUH 13763); Aug 2005, Conley K. McMullen 867

(JMUH 13764); Aug 2005, Conley K. McMullen 868 (JMUH 13766); Aug 2005, Conky K. McMullen 869 (JMUH 1 37869); Mar 2007, Conley K.

McMullen and Michael Shane Woolf 887 (JMUH 1381 1); Mar 2007, Conley K. McMullen and Michael Shane Woolf 888 (JMUH 1 381 3); Mar 2007,

Conley K. McMullen and Michael Shane Woolf 889 (JMUH 13815); Mar 2007, Conley K. McMullen and Michael Shane Woolf 890 (JMUH 13817);

Aug 1891, G. Bern 209 (GH 233524); Aug 1972, Michelle and Ole Hamann 2140 (C 24/2007/3); 1825, John Scouler s.n. (K 449165).

Nomendatural history.— Hooker first described this species in 1847 as Cordia scouleri Hook, f., transferred by

Andersson (1855) into Varronia, and later Lithocardium by Kuntze (1891). Borhidi (1988) renamed the species

Varronia scouleri (Hook, f.) Borhidi, without regard to Andersson’s (1853) designation rendering this combina-

tion illegitimate (see Art. 45.3 and Art. 53.1, McNeill et al. 2006). The currently accepted name for this species

is Varronia scouleri Anderss., based upon Miller and Gottschling’s (2007) reinstitution of genus Varronia.

Contribution of this study.— Based on characters observed in the type specimen, this species typically has

lanceolate leaves with coarsely serrate margins (Fig. 15), but this was variable when observing other herbari-

um specimens. As previously mentioned, V: scouleri tends to have longer leaves and more coarsely serrate

margins than V. canescens, the most similar species. The adaxial leaf surface of V scouleri is pubescent with

simple or forked (Fig. 16) or sometimes stellate (Fig. 17) hairs. Abaxial leaf surfaces are sparsely pu-

and forked, sometimes simple, hairs (Fig. 18). Much longer and more noticeably

:an be found on the veins. These large hairs on the veins are very distinctive of this species. Pubes-

:alyx is composed of erect and simple, and sometimes forked or stellate hairs. Simple hairs on the

calyx lobes are noticeably longer than the hairs on the calyx tube. The calyx lobes themselves are usually lon-

ger than the calyx tube at anthesis. Varronia scouleri is distinct in that its calyx lobes can be exceptionally long

and outwardly splayed compared to the other three species.

DISCUSSION

All four species of Varronia endemic to the Galapagos Islands deserve recognition as distinct species. Based

upon this morphological study, each species can be distinguished by a combination of characters. In addition,

these characters appear to have remained constant over the 185 years these species have been collected. While

this is not a substantially long period of time in terms of evolution, it does indicate that the characters used in

this study remain relatively stable during the time period in which these specimens were collected and studied.

No obvious hybrids (plants with characteristics of more than one species) were observed during the morpho-

logical study, suggesting that there may be little if any hybridization between species. Based on personal obser-

vations, the individual species of Varronia do not generally appear to be found within close proximity to one

another, further reducing the possibility of hybridization through shared pollinators. Additionally, these Var-

ronia species are distylous, and initial research on V. scouleri indicates that almost all fruit set is the result of pin

and Michael Shane Woolf, 876 (JMUH 13787).

> and forked hairs. Scale represents 300 pm. Specimen: Mar 2007, Conley K. h

flowers that have been pollinated with pollen from thrum plants of the same species (CKM and JKS, unpub-

lished data). Thrum plants produce few, if any, fruits, indicating that only about 50% of a local plant population

is capable of producing fruit. This initial research also indicates that only about 14% of flowers produce fruits.

The limited ability of Varrania to effectively produce fruit further limits the possibility of established mature

hybrid individuals. Future research on these species will include controlled pollination crosses between spe-

cies to determination if hybridization is possible, and if so, the viability of the resulting hybrid individuals.

Due to the limited fruiting ability of these species, future recruitment may be cause for concern if the

habitat of these species is disturbed. In 1959, the Ecuadorian government created the Galapagos National Park,

which comprises about 97% of the total landmass of the archipelago (Camancho 2005). Because such a signifi-

likely to be in these protected areas. This existing protection decreases the risk of habitat destruction that

could potentially reduce the survival of these species. Based on February 2011 observations on Santa Cruz Is-

land near the Charles Darwin Research Station and the town of Puerto Ayora, few seedlings are visible among

established populations, though mature adult plants appear to be relatively common and vigorous.

At this time, little is known about the abundance or complete distribution of Varronia in the Galapagos,

and future field studies of these species would benefit from assessments of these populations. Data from her-

barium collection labels has provided enough information to determine on which islands each species can be

found (Fig. 4), but specific collection locations were rarely, or only partially provided, apart from those speci-

mens collected by CKM. Based on current information, it is evident that each species retains its foliar morphol-

ogy and can be identified across multiple islands suggesting that the characters are not geographically deter-

mined. Several islands have all four species of Varronia present (Fig. 4). It is possible that this is the result of

collection intensity, and future studies may find that the least distributed species may in fact be present on

other islands that have not previously undergone intensive collecting. Detailed GPS coordinates for Varronia

populations in the Islands would provide the information needed to construct more complete distribution

maps for each species. These species do not currently appear to be under any increased threat, and their con-

servation needs are unlikely to change in the near future; however, the IUCN would benefit from updated in-

formation and distributions of these species in order to reevaluate, or evaluate for the first time, the status of

each species.

This study supports the existence of four morphologically distinct endemic Varronia species in the Gala-

pagos Islands. While Weeks et al. (2010), did not find V. leucophlyctis, V. canescens, and V scouleri to be geneti-

cally distinct from one another, this may have been due to the ITS regions that were used in the study. These

regions may not be variable if these species have diverged relatively recently. Weeks et al. (2010) suggest that

fingerprinting techniques that record nucleotide polymorphisms, such as microsatellites or amplified frag-

ment length polymorphisms, would be a better method for use in the future. Future molecular studies may M

that genetic variation exists in now-unexamined regions, but until that time, these populations deserve recog-

nition as morphologically different species based on this study.

Morphological characterization indicates the existence of four endemic species of Varronia in the Galapagos

Islands that are distinct. The results from this study will be helpful in evaluating these species for their conser-

vation statuses. Over 100 misidentified specimens are now correctly identified through the course of this

study, and the diagnostic information provided here in the new taxonomic keys will help provide the informa-

tion necessary for other specimens to be reevaluated, or new collections to be correctly identified. In addition,

an effort must be made to assess the size and stability of the populations of each endemic Varronia species in

order to evaluate how these species should be managed. Protecting the endemic species of the Galapagos is

important both for the survival of the individual species as well as the continuation of the Galapagos Islands as

a unique example of island evolution and diversity. Effective conservation programs to protect this exceptional

diversity can only result from research that strives to document the flora and fauna of these islands. Accurate

descriptions of the distinctive species of the Galapagos, such as those accomplished in this study, are the first

step towards protecting and ensuring their future on the islands.

ACKNOWLEDGMENTS

JKS and CKM thank the Charles Darwin Foundation, the Charles Darwin Research Station, and the Galapagos

National Park Service for logistical support during fieldwork in the Galapagos Islands. CKM thanks Lenin

Cruz, Michael S. Woolf and Beattie G. Sturgill for field assistance. Thanks are extended to Michael H. Renfroe

and Heather Peckham Griscom for reviewing an earlier version of the manuscript and providing constructive

urnal of the Botanical Research Institute of Texas 6(1)

suggestions. James L. Reveal is gratefully acknowledged for nomenclatural advice. A special thanks also to

Lance Kearns for his help in obtaining the SEM images used in this paper that provide an excellent example of

many of the characters used in the new taxonomic key. Many thanks to Susana Chamorro for translation as-

sistance, and JKS would also like to thank Reuben and Catarina Heleno for graciously providing lodging in

their home in Galapagos during her field work for this study. Marc Gottschling and an anonymous reviewer

provided helpful suggestions.

The following funding sources have allowed travel to the Galapagos Islands and the completion of this

research: The Eari Core Student Award (Southern Appalachian Botanical Society), the Norlyn L. Bodkin Schol-

arship for Arboretum and Botanical Field Studies (James Madison University), the Peter T. Nielsen Annual

Award for Botanical Studies (James Madison University), and the James Madison University Department of

Biology. Appreciation is extended to the following herbaria for use of their Varronia specimens: BKL, C, CAS,

CGE, GH, JMUH, K, NY, and S.

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galapagospark.org/documentos/DPNG_plan_de_manejo.pdf

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Biotropica 41 :1 62-1 70.

Friesen, C. 1 933. Liste des Varroniae exduendae et conservandae. Bull. Soc. Bot. Geneve 2(24):1 80-1 83.

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Johnson, M.P. and P.H. Raven. 1973. Species number and endemism: the Galapagos archipelago revisited. Science

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Journal of the Botanical Research Institute of Texas 6<1)

BOOK REVIEW

Sylvan T. Runkel and Dean M. Roosa. 2010. Wildflowers of the Tallgrass Prairie: The Upper Midwest. (ISBN:

978-1-58729-796-0, pbk.). University of Iowa Press, 119 West ParkRoad, 100 Kuhl House, Iowa City, Iowa

52242, U.S.A. (Orders: www.uiowapress.org, 319-335-2000). $29.95,308 pp., 131 color photos, 5.5" x 8.5".

Growing up in America, it is hard not to be exposed to the wonderful tales of Laura Ingles Wilder and her fam-

ily’s efforts to survive the blessings and hardships of life on the prairie. As a girl, I envisioned myself as Laura,

romping through the tallgrass prairie with my pup at my side, experiencing the many adventures from eking

out a living from the hard black earth, to harsh winters, to encounters with Native Americans, to one-room

schoolhouses, and to lying by a stream watching the butterflies flit back and forth.

Having recently moved from Southern Wisconsin, the land of the Tallgrass Prairie, to Northern Texas’

Blackland Prairie, it was meant to be that I review this wonderful guidebook, “Wildflowers of the Tallgrass

Prairie, The Upper Midwest” by Sylvan T. Runkel and Dean M. Roosa. The prairie is in my blood. I would even

go so far as to describe myself as a “prairie rambler” as the authors are described in the book’s forward, al-

though in no way can 1 claim the expertise of these two esteemed prairie experts.

Once covering approximately 221,436 square miles from Canada to Texas, tallgrass prairie was subject to

the natural conditions of grazing buffalo, harsh winters, and prairie fires. Humans, however, have interfered

with its natural cycles. With the invention of the steel plow, the rich and tough prairie lands soon became

farmland. Today, the tallgrass prairie is estimated to cover only 1% of its original expanse. Remnants of the

tallgrass prairie remain dispersed among the Upper Midwest states. In Wisconsin alone, only about 4,000

Runkel and Roosa offer us a wonderful guidebook with stunning full page, color photographs by botanist

Thomas Rosburg, that make one’s heart long for the tallgrass prairie. The book is organized by season begin-

ning with the early blooming pasque flower, followed with summer beauties such as Maximilian sunflower

and prairie coreopsis, and ending with the lovely fall blazing star and closed gentian. Prairie grasses such as big

bluestem, Indian grass, and switchgrass, among others, are an integral part of the book. The authors provide

readers with both common and Latin names, as well as definitions which help the amateur wildflower enthu-

siast remember them. They provide detailed plant and habitat descriptions as well as historical and medicinal

uses of the plant by Native Americans and pioneers.

While primarily a guidebook, the authors make an important link to the need for conservation of these

vital and yet sensitive prairie lands. As they say, “If the prairie goes, an entire community is lost forever— a

community that was the foundation of our Midwestern society.” Find yourself a copy of this book, a pair of

boots, and go ramble on the prairie. You too will be convinced of its beauty, and the need to protect it.

— Gwen Michele Thomas, Texas Master Naturalist & Volunteer,

e of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, 1/.SA1

lical Research Inst

BOOKS RECEIVED/REVIEWS FORTHCOMING

Patricia Fouey. 2011. The Guide to Oklahoma Wildflowers. (ISBN: 978-1-60938-046-5, pbk.). University of

Iowa Press, 119 W. Park Road, 100 Kuhl House, Iowa City, LA 52242-1000, U.S.A. (Orders: http://www.

uiowapress.org/books/2011-fall/guide-oklahoma-wildflowers.htm, Phone: 319-335-2000, Fax: 319-335-

2055, Phone orders: 800-621-2736, Fax orders: 800-621-8476, E-mail: uipress@uiowa.edu). $39.95, 312

pp., 415 color photos, 1 color map, 6" x 9".

Craig Pittman. 2012 The Scent of Scandal: Greed, Betrayal, and the World’s Most Beai

978-0-8130-3974-9, hbk.). University Press of Florida, 15 Northwest 15th St., Gaine

2079, U.S.A. (Orders: http://www.upf.com). $24.95, 299 pp., b&w photos, 6" x 9".

rtiful Orchid. (ISBN:

sville, Florida 32611'

A NEW SPECIES OF TAPEINOSTEMON (GENTIANACEAE)

FROM THE CHOCO OF NORTHERN ECUADOR

Jason R. Grant

Laboratoire de Botanique Evolutive, Institut de biologie, University de Neuchdtei

Faculty de Sciences, Unimail, rue Emile-Argand 1 1

2000 Neuchdtei, SWITZERLAND

RESUMEN

While examining specimens of neotropical Gentianaceae at the Missouri Botanical Garden for work largely on

Macrocarpaea (Grant 2011; Grant & Trunz 2011), a distinctive specimen came to light. With John Pruski and

Ron Liesner we determined that this was a member of the small genus Tapeinostemon. Species of this genus

range from annual to perennial herbs that are woody at the base. The pentamerous flowers are positioned in lax

axillary to terminal cymes, and depending on species are distinctly nodding or not (Struwe et al. 1999; Struwe

et al. 2002). The seeds are minute with a reticulate testa with numerous important characters for species dif-

Tapeinostemon occurs primarily on tepui formations of the Guayana Highlands with six species: T. brew-

eri Steyerm. & Maguire, T. jauaensis Steyerm. & Maguire, T. longiflorum Maguire & Steyerm., T. rugosum

Maguire & Steyerm., T. sessiliflorum (Humb. & Bonpl. ex Schult.) Pruski & S.F. Sm. and T. spenneroide s Benth.

(Struwe et al. 1999). Two species occur in the Andes: the first Tapeinostemon zamoranum Steyerm., which has

the broadest distribution of any species in the genus, ranges across several biogeographic zones from southern

Colombia to the Amotape-Huancabamba region of southern Ecuador and northern Peru, to the Cordillera

Central of Peru and northern Bolivia, always on Amazon-facing slopes; the second is a new species, T. adulans

J R. Grant described here, which occurs on Pacific-facing slopes of the Chocd region northern Ecuador. The

distribution of Tapeinostemon, a Guayana Highland-centered genus with several species also occurring in the

Andes, is similar to that of Bonyunia (Loganiaceae), Potalia (Gentianaceae), and Tachia (Gentianaceae) (Grant

2009); yet in Tapeinostemon there is an absence of a widespread species in the Amazon basin as in the other

three.

J - 8ot Res. Inst. Texas 6<1): 101-1

h. ( Liesneri Camevali 22666 [MO]). Photo Neil V.llard.

Journal of the Botanical Research Institute of Texas 6(1)

r 1258 [MO]). Photo Neil Villard.

105

Unbranched herb 0.5 m tall, glabrous throughout. Steins rounded below to quadrangular above. Leaves (40-)

130-140 mm long, long-petiolate; petioles (3-)20-25 mm long, blades lanceolate to ovate, (37-)110-120 x

(16-)37-43 mm, entire, thin; leaf base attenuate and decurrent on the petiole; leaf apex acuminate. Inflores-

cences: lax axillary cymes, 30-78 mm long; branches spreading laterally, 5-20 mm long, 4-10 laxly flowered

Per branch. Lower bracts lanceolate to ovate, decreasing in size and becoming obovate toward the apex of the

106

Golden translucent

irregular polygonal cells

(pentagonal and hexagonal,

some lengthened with the

Reticulate with irregular polygonal

Straight to curving, thin, strongly

raised vertically; inner walls with

band-like thickenings

inflorescence; lower bracts petiolate, upper bracts sessile; bract petioles 0-3 mm; bracts 2-14 x 0.5-32 mm;

bract base cuneate to attenuate and decurrent on bract petiole; bract apex obtuse to acuminate.. Flowers pedi-

cellate, pedicels 1-8 mm. Sepals separate to the base, glabrous, ecarinate, narrowly triangular, 1.0-1.5 x

0.25-0.33 mm, apex acuminate to acute. Corolla funnel-shaped, 1.5-2.5 x 0.33-0.5 mm, greenish (.Croat

99788), corolla lobes less than 0.5 mm long, corolla lobe apex acute. Androecium and gynoecium unknown

(too few flowers to risk damaging the specimen). Capsules ellipsoid to obovoid, 3-5 x 1-2 mm, rugose, bilocu-

lar, each locule with around 80 seeds (ca. 160 seeds per fruit). Seed characters in Table 1.

Morphology and similarities. — Tapeinostemon adulans (Fig. 1) is distinct in being an unbranched 50 cm tall

herb with short axillary inflorescences and globose-shaped seeds. It is similar to T. spenneroides (Fig. 2) in its

overall form, inflorescence architecture, leaf shape, non-undulating leaf margins, and seed morphology, but

differs in its short stature, smaller calyx 1.0-1.5 vs. 1.0-2.0 mm long, smaller corolla 1.5-2.5 vs. 4-6 mm, and

seed characters listed below. Tapeinostemon zamoranum (Fig. 3) has crenate undulating leaf margins, flowers

that are distinctly nodding, and an unique seed morphology. In the key to species of Tapeinostemon by Struwe

et al. (1999), both T. adulans and T. zamoranum will key to T. spenneroides.

Seed morphology .— The seeds of Tapeinostemon spenneroides have been characterized by Bouman et al.

(2002). Here, the seeds of two additional species T. adulans (Fig. 4 E-F) and T. zamoranum (Fig. 4 A-B) as well

as T. spenneroides (Fig. 4 C-D) are described and illustrated. The calyx and corolla which are usually good

characters in differentiating species, are so small in Tapeinostemon that they seem somewhat unreliable when

dried as herbarium specimens for clear species differentiation. Therefore, seed anatomy is a useful character to

distinguish these three species. The seeds of Tapeinostemon range in color from solid reddish brown to golden

translucent and are minute (200-300 x 155-250 pm), globose, elliptic to ovoid, with a reticulate testa. The

seeds of T. adulans and T. spenneroides are similar to that of two other neotropical genera, Coutoubea and Schul-

tesia, while the seeds of T. zamoranum appear similar to Cicendia (see photos in Bouman et al. 2002). The char-

acteristics of these three species can be seen in Table 1.

Distribution and habitat.— Tapeinostemon adulans occurs in the diverse Choco region on lower elevation

Pacific-facing slopes of the Andes of northern Ecuador.

107

Etymology . — From the Latin, adulans, adulatory, for the collector of the type, Thomas B. Croat of the Mis-

souri Botanical Garden.

I thank Jennifer Kuhl at the Missouri Botanical Garden for her help in finding and sending information on

various specimens. I also acknowledge assistance from several other colleagues at MO during my visit there in

summer 2011, especially Ron Liesner and John Pruski. Scanning electron microscope photos were prepared

with Mireille Leboeuf at the CSEM (Centre Suisse d’Electronique et de Microtechnique) in NeuchStel, Switzer-

land. Marion J. Jansen-Jacobs and James S. Pringle provided helpful review comments, thus improving the

Bouman, R, L Cobb, N. Devente, V. Goethals, PJ.M. Maas, and E. Smets. 2002. The seeds of Gentianaceae. In: L. Struwe and V.A.

Albert. Gentianaceae, systematics and natural history. Cambridge University Press. Pp. 498-572.

Grant, J.R. 2009. A revision of neotropical Bonyunia (Loganiaceae: Antonieae). Ann. Missouri Bot. Gard. 96:541 -563.

Grant, J.R. 201 1 . De Macrocarpaeae Grisebach (ex Gentianaceis) speciebus novis IX: a synopsis of the genus in Bolivia.

Harvard Pap. Bot. 16:389-397.

Grant, J.R. and V. Trunz. 201 1 . De Macrocarpaeae Grisebach (ex Gentianaceis) speciebus novis X: a synopsis of the genus

in montane Atlantic forests of Brazil. Harvard Pap. Bot. 16:399-420.

Struwe, L, J.W. Kadereit, J. Klackenberg, S. Nilsson, M. Thiv, K.B. von Hagen, and V.A. Albert. 2002. Systematics, character evolu-

tion, and biogeography of Gentianaceae, including a new tribal and subtribal classification, in: L. Struwe and V.A.

Albert. Gentianaceae, systematics and natural history. Cambridge University Press.

Struwe, L., PJ.M. Maas, O. Pihlar, and V.A. Albert. 1 999. Gentianaceae. In: P.E. Berry, K. Yatskievych, and B.K. Holst, eds. Flora

of the Venezuelan Guayana. Vol. 5. Missouri Botanical Garden, St. Louis. Pp. 474-542.

BOOK REVIEW

Candace Savage, Principal Photograph by James R. Page. Illustrations by Joan A. Williams. 2011, Prairie: A

Natural History. (ISBN: 978-1-55365-588-6, pbk.). Greystone Books, 2323 Quebec Street, Suite 201,

Vancouver BC V5T 4S7, CANADA. (Orders: www.dmpibooks.com). $29.95, 320 pp., color throughout,

hAv illustrations, 7.5" x 10".

Prairie: A Natural History provides both a historical and present-day story of the North American prairie; a

story that eloquently weaves together threads of botany, geology, biology, archaeology, chemistry, ranch man-

agement, and anthropology to produce a tapestry of the prairie. Savage’s writing style will transport readers to

the sites of her research. “Climb up there on a blue day in early September,” she writes about the Frenchman

Valley in southern Saskatchewan, “out and onto the bald steppe at the top, and the wind will slam against you

as if it had a grudge against anything that dared to raise its head above the grasses.” Corresponding pictures,

maps, and botanical illustrations provide greater detail to this piece of art— augmenting readers’ understand-

ing and appreciation for this ecological wonder. The final details that truly make this book a piece of art are the

side stories that compliment the main threads of investigation, e.g., Cheyenne folk lore, excerpts from Lewis

and Clark’s diaries, agricultural practices, and animal behaviors.

Savage has written a literary piece that belongs both on a coffee table and in the classroom. The chapters,

which focus on individual elements of the prairie (e.g., soil, water, woodlands, animals, and land manage-

ment), are written so they can stand alone, yet together they provide a holistic representation of a vastly com-

plicated system. Both prairie novices and experts will find this book illuminating due to its diversity of topics

and its inclusion of both basic and intermediate levels of subject matter. From simple definitions to more com-

plex connections, this book will find its way into the hearts of those who read it.

Prairie: A Natural History argues for the conservation of one of the most endangered ecological systems.

Savage provides a historical view of how the nature of the prairie has changed due to both natural and human-

made conditions, and she delicately balances the severity of the issue of the prairie’s survival with hope for the

future. Individuals who ask the question, “Why save the North American prairie?,” will be illuminated with

each turn of the page.

— Amanda Stone Norton, Ph D-.

Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, Texas 76107-3400, USA

BOOKS RECEIVED/REVIEWS FORTHCOMING

Robert B. Shaw. Photographs by Paul Montgomery and Robert B. Shaw. 2012. Guide to Texas Grasses. (ISBN:

978-1-60344-186-5, pbk. flexbound). Texas A&M University Press John H. Lindsey Building, Lewis

*Street, 4354 TAMU, College Station, Texas 77843-4354, U.S. A. (Orders: www.tamupress.com, 800-826-

8911). $45, 1080 pp., line drawings, distribution maps, color throughout, 7" x 10".

Fred Dortort. 2011. The Timber Press Guide to Succulent Plants of World: A Comprehensive Reference to

More Than 2000 Species. (ISBN: 978-0-88192-995-9, hbk,). 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). $49.95, 344 pp., color throughout, 8.5" x 11".

Scorr Calhoun. 2012. The Gardener s Guide to Cactus: The 100 Best Paddles, Barrels, Columns, and

Globes. (ISBN: 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".

LECTOTYPIFICATION OF

BAUHINI A NERVOSA (LEGUMINOSAE: CAESALPINIOIDEAE)

S. Bandyopadhyay

Central National Herbarium

Botanical Survey of India, P.O. Botanic Garden

Howrah - 71 1 103, West Bengal, INDIA

subirbandyopadhyay@yahoo.com

ABSTRACT

tauhirrid nervosa (Benth.) Wall, ex Baker.

In the course of my taxonomic studies on the Indian Bauhinias I have sometimes faced a great difficulty in lo-

cating a particular specimen cited as type by the author(s). This paper deals with one of such problems encoun-

tered by me.

Larsen and Larsen (1984: 35) cited India ‘(Sylhet (type), Assam, Khasya], China (?), Upper Burma’ under

the distribution of Bauhinia nervosa (Benth.) Wall, ex Baker [basionym Phanera nervosa Benth.].

Zhang and Zhu (2006) cited Type: Bangladesh. Sylhet: Without precise locality, Wallich 5777 (holotype,

K!)’. They have, however, neither given a clue in their publication so that one can find out the holotype exam-

ined by them at K, nor any specimen at K has been annotated by them accordingly.

In the protologue of Phanera nervosa, Bentham (1852: 262) cited ‘Bauhinia nervosa, WALL. Cat. n. 5777. In

mont. Sillet’ after the morphological description.

The specimen at K-WALL1CH bears a piece of paper on the top left hand side on which the name Bauhinia

nervosa Wall is inscribed and written as: Ching akahoo (?) this was found in the Katchar Hill climber yellow

flower stamens reddish like style yellow this flower has good smell this was found in the month of October

1822. Pinned to it there is anot her strip of paper on which it is inscribed as: 5777 Bauhinia nervosa Mont. Sillet

LD (?) in an exactly same pattern of handwriting as in Wallichian Catalogue. In another small piece of paper

affixed on the top comer of the right hand side there is the name Bauhinia nervosa Wall, 5777, Cachar Montes

1823. This specimen is unlikely to have been studied by Bentham because neither any of the morphological

characters stated on the sheet has been included in the protologue nor the place of collection has been stated to

he Katchar Hill/Cachar Montes and so also the date of collection.

At K (000760781) there is a relevant herbarium sheet bearing the stamp ‘HERBARIUM BENTHAMIA-

NUM 1854’. This stamp suggests that the specimen was donated by Bentham in 1854 from his own herbarium

to the Kew herbarium when he found that the maintenance of his own herbarium had become too expensive

for him (http://en.wikipedia.org/wiki/George_Bentham; http://www.kew.org/heritage/people/bentham.html;

http://www. anbg.gov.au/biography/bentham.george.html) [also see under Bentham’s HERBARIUM and

TYPES in Stafleu and Cowan, TL2 1:173. 1976]. This sheet has 3 separate leaves (afl in detached condition) and

an inflorescence. One of the leaves is folded, slightly damaged, and its lobe tips are not evident. The data in-

scribed on the sheet in pen are as follows: 1) Bauhinia nervosa, Sillet Mont, H.1. 5777 (seen on the left bottom of

the inflorescence); 2) Phanera nervosa Bth, Pi. Jungh. 1. 262 (seen on the left bottom of the folded leaf); and 3)

almost in the middle of the sheet, it has an inscription in pencil as follows: 5777, B nervosa, Sillet Mont, cad As-

sam Griff in [illegible] (not associated with any of the 4 specimens). One of the remaining two leaves is on the

top left and the other is at the bottom left. The left top leaf lobes are distinctly acute, whereas the left bottom leaf

Fk. 1. Photograph of the leetotype of Bauhinian

Bandyopadhyay, Lectotypification of Bauhinia nervosa

Within the protologue, the “leaflets” (an error for leaf lobes) were described therein as ‘acutiusculis v(el).

acute.” Furthermore, the scandent habit that has been stated in the protologue must have been taken from

some other source because the specimen neither has any tendril nor there is any such inscription on the sheet.

As no other collection of “Wall. Cat. num. list no, 5777” used by Bentham could be located, 1 am lecto-

typifying here the name with the aforementioned Kew specimen (000760781), which was originally part of

‘HERBARIUM BENTHAMIANUM:

i (Benlh.) Wall, ex Raker in J.D Hooker, I I But India 2:283. 1878. im same jsof hasmnym rim-

Any specimen at K having the locality spelled as ‘Sylhet’ could not be located but it may be so that Zhang and

Zhu (2006) cited the same specimen studied by me and changed the spelling of ‘Sillet’ to ‘Sylhet’ following the

current usage. Although Art. 9.8 of the ICBN (McNeill et al. 2006) allows “The use of a term defined in the Code

(Art. 9.1-9.7) as denoting a type, in a sense other than that in which it is so defined, is treated as an error to be

corrected (for example, the use of the term lectotype to denote what is in fact a neotype),” Zhang and Zhu’s

(2006) citation ‘holotype, K!’ cannot be construed as an inadvertent lectotypification, even if the same speci-

men has been studied by us i.e, D.-X. Zhang and G.-H. Zhu and myself. This is because, Art. 7.11 of the ICBN

(McNeill et al. 2006) requires that on or after 1 January 2001, the typification statement must include the

phrase “designated here” (hie designatus) or an equivalent, which is not the case in Zhang and Zhu’s (2006)

type citation.

ACKNOWLEDGMENTS

I am thankful to the Director, Botanical Survey of India and Additional Director, Central National Herbarium,

BS1 for their help, to the Director, Royal Botanic Gardens, Kew for providing literature and cibachrome photo-

graph of the Kew specimen, V.P. Prasad, IBLO, RBG, Kew for sending the image of the specimens at K-W and

BM, K.N. Gandhi for refining the manuscript and to Dave Boufford and Emily W. Wood, Harvard University,

USA for their comments.

REFERENCES

Baker, J.G. 1878. Bauhinia. In: J.D. Hooker, Flora of British India 2(5);275-285. L. Reeve & Co., London.

Bentham, G. 1852. Leguminosae. In: F.A.W. Miquel, ed. Plantae Junghuhnianae. Leiden. Pp. 205-269.

Ursen, K. and S.S. Larsen. 1 984. Bauhinia. In: T. Smitinand and K. Larsen, eds. Flora of Thailand 4(1 ):4-45. The Forest Her-

barium, Royal Forest Department, Bangkok.

McNeill, J„ F.R. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, K. Marhold, D.H. Nicolson, J. Prado, P.C. Silva, J.E. Skog, j.H.

Wiersema, and NJ. Turland (eds.). 2006. International code of botanical nomenclature (Vienna Code). Adopted by the

Seventeenth International Botanical Congress Vienna, Austria, July 2005. A.R.G. Gantner Verlag, Ruggell, Liechten-

stein. (Regnum Veg.146].

Zhang, P.-X. and G.-H. Zhu. 2006. The identity of Bauhinia claviflora and of B. dioscoreifolia (Leguminosae)

Sin. 44:651-653.

0. Acta Phytotax.

112

Journal of the Botanical Research Institute of Texas 6(1)

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— Grace Bascom, Research Associate

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, USA

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ture, and art. The photos capture the beauty of these plants and also illustrate the concepts discussed in the text

IMPATIENS JOHNSIANA (BALSAMINACEAE),

A NEW SCAPIGEROUS BALSAM FROM WESTERN GHATS, INDIA

^.K. Ratheesh Narayanan, N. Anil Kumar, and Jayesh P, Joseph

C.N. Sunil

Emakulam-683516, Kerala , INDIA

T.Shaju

ABSTRACT

INTRODUCTION

The genus Impatient L. (Balsaminaceae) is a genus of over 1,000 species widely distributed throughout the

Northern Hemisphere and tropics (Mabberley 2008). In India, the concentration of Impatiens species is re-

markably local and occurs in two well-defined regions, viz., the Himalayas in the North and the Western Ghats

in the South (Hooker 1908; Gamble 1915; Vivekananthan et al 1997). There is no doubt that the Western Ghats

is the second richest area in the Indian subcontinent and perhaps in the world (Bhaskar 1981; Viswanathan &

Manikandan 2003). Out of the 203 species known to occur in India, about 86 species are found in the Western

Ghats (Nair 1991). In India, scapigerous Impatiens form a peculiar Section ‘Scapigerae’ and are restricted to the

Western Ghats-Sn Lanka Hotspot of Biodiversity (Grey-Wilson 1980; Vivekananthan et al. 1997). Except for I.

acaulis Arn., which is distributed throughout the wet tropical forests of the Western Ghats and Sri Lanka, all

the other species have a very narrow distribution and are endemic to small microcenters in the Western Ghats,

especially in the southern part (Kumar & Sequiera 1996; Bhaskar 2006; Jyosna et al. 2009; Anil Kumar et al.

2011).

The Wayanad district of Kerala forms a biodiversity rich area in the Nilgiri phytogeographical region of

the Western Ghats. During an exploration of the flowering plants there, 21 species of Impatiens were collected

in various habitats (Ratheesh Narayanan 2009). One, collected in a high altitude evergreen forest, was quite

distinct from the hitherto known scapigerous species and it is described and illustrated here as new.

Journal of the Botanical Research Institute of Texas 6(1)

Narayanan et al., A new species of Impatiens from India

115

H. Leaf, J. Fruit.

116 Journal of the Botanical Research Institute of Texas6(1)

Scapigerous perennial herbs, epiphytic, 15-22 cm tall; rootstock tuberous, yellowish. Leaves 1, rarely 2, radi-

cal, petiolate, pendulous, distantly serrate, broadly elliptic to oblanceolate, acute, mucronate at tip, base

broadly cordate, unequal, 14-22 x 4-5 cm, fleshy, hairy above, glabrous below, 5-7-nerved, adaxially green,

abaxially pale pink, deep pink along midrib and lateral nerves; petiole 5-8 cm long, glabrous, green above,

light pink below. Inflorescence scapose, unbranched, semi-pendulous, peduncle with a bend above the middle,

shorter than the leaves, 12-20 cm long, fleshy, glabrous, pink, terete, less than 10-flowered; flowers in racemes,

distal, ca. 1.5 cm across, deep pink with white center; bract fleshy, light green with pink streaks, broadly ovate

to oblong, 4-5 mm long, acute, glabrous; pedicels shorter than the flower, 1.6-1.8 cm long, terete, slender,

white, elongating in fruit, glabrous; lateral sepal 1.7-2 x 2-2.5 mm, broadly ovate to oblong, obtuse-acute at tip,

white with pink tinge, glabrous; lower sepal (lip) ca. 8 x 5 mm, white, glabrous, spurred; spur 1.5-1.7 cm long,

funnel-shaped, coiled with broad tip, white, glabrous; dorsal petal (standard) broader than long, 0.5-1 cm

across, hooded, glabrous, pink, margins wavy; lateral united petals 3- lobed, 1.9-2 x 1.2-13 cm, lobes unequal,

with a tuft of yellow papillae little below the basal lobes; basal lobes larger than distal lobes, ca. 1.3 x 0.6 cm,

broadly obovate, bent upwards, midlobes ca. 0.5 x 0.5 cm, straight, distal lobes ca. 0.5 x 0.5 cm, oblong, tip

obtuse, straight; stamens 5, connate, ca. 2 x 1 mm, filaments pink, connate above middle, anthers pale white;

ovary pale yellow, broadly elliptic, acute at apex, glabrous. Capsules broadly ellipsoid, acute, ca. 1 cm long,

glabrous; seeds many, ca. 2 mm long, tubercled, with a dense tuft of hairs at each end.

Paratype: INDIA. KERALA. Wayanad District: Kattimattom hills, Nll°30.716' E76°06.239’ ca.1700 m, 20 Sep, 2011, M.K. Ratheesh Naray-

anan, C.N. Sunil & T. Shaju MSSH 4455 (Herb. MSSRF, Kalpetta, Wayanad).

Flowering and fruiting.— Flowering is from July with a peak in August. Fruit matured during September-October.

Etymology.— The specific epithet is in dedication to the late Prof. John C. Jacob (who was popularly

known as ‘Johnci’), an ardent naturalist who devoted his life to conserve the biodiversity of the Western Ghats,

India.

Distribution and conservation status.— Endemic to the Kattimattom Hills of the Chembra-Vellarimalahill

ranges, Wayanad District, Kerala, India. Small population of this species is distributed on densely clothed tree

trunk in evergreen forests at an altitude of 1500-1700 MSL. The species is usually seen as an epiphyte on the

moss-covered trunk of evergreen tree species. Our observations showed that there were less than five hundred

mature individuals restricted to a 10 km 2 Vested Forest area. By following IUCN criteria (IUCN 2001) for as-

sessing the status of Rare and Threatened plants, Impatiens johnsiana is assessed as belonging to Critically

Endangered (CR) category. Its range (extent of occurrence) is less than 100 km 2 , and the quality of habitat is

declining continuously.

Taxonomic note.—lmpatiensjohnsiana resembles I. dendricola Fisch. in its pendulous habit, 3-lobed lateral

united petals with a tuft of papillae, and seeds with a dense tuft of hairs at each end. The shape and size of

leaves, color of flowers, shape and size of the lateral sepal, dorsal petal, spur and lateral united petals, and tex-

ture of seeds differentiates this species from the other. Impatiens johnsiana has 14-22 cm long, elliptic-oblan-

ceolate leaves, pink flowers, broadly ovate lateral sepals, and a dorsal petal with wavy margins in contrast to the

less than 10 cm long, ovate to oblong-lanceolate leaves, white flowers, falcate-ovate lateral sepals deeply lobed

at base and entire dorsal petal of I. dendricola. The new species has 1.5-1.7 cm long, coiled spur with broad

rounded apex in contrast to 1-1.5 cm long, straight, clavate spur of I. dendricola. The lateral united petals of the

new species are not auricled, curved basal lobes larger than the distal lobes, and it has tubercled seeds. But in I

dendricola, the lateral united petals have an auricle, straight strap-shaped basal lobes of the lateral united petal 5

smaller than the distal lobes, and seeds muriculate. The new species also resembles I. scapiflora in some of its

characters, e.g., large pink flowers, lateral united petals without an auricle, and recurved basal lobes larger than

the distal lobes, but differs by the pendulous elliptic-oblanceolate leaves, up to 2 cm long pedicels, less than 2

cm long, coiled spur with broad rounded apex, tubercled seeds, with a dense tuft of hairs at ends in contrast to

an ovate to obovate leaf, up to 5 cm long pedicels, up to 7 cm long, tubular spur with narrow apex, and seeds

hairy all over. Diagnostic morphological characters of Impatiens johnsiana and related species are provided in

Table 1.

Narayanan et al..

Table 1. Diagnostic morphological characters ot Impatiens johnsiana an

Basal lobes of lateral ui

than the distal lobes.

Capsules ellipsoid o

ACKNOWLEDGMENTS

The authors are grateful to the Chairman Prof. M.S. Swaminathan and Executive Director of the M.S. Swami-

nathan Research Foundation, Chennai, India, for providing facilities and support. The logistics provided by

the Kerala Forest Department for the fieldwork and the help rendered by K.T. Satheesh, V.R. Volga, V. Mini,

Jithin M.M., and S. Mithunlal, Community Agrobiodiversity Centre, M.S. Swaminathan Research Foundation,

Kalpetta are thankfully acknowledged. Sincere thanks are expressed to J.F. Veldkamp (L) for critical remarks

and the Latin diagnosis. Anil Zakharia and Sethu Madhavan for the field support and encouragement. Two

anonymous reviewers provided constructive and helpful reviews.

3, Adlard & Son, London, reprinted by

REFERENCES

Anil Kuw* n, M.K. Rntneesh IWw*,. P. Suiwwmt. R. Mw»l KA Suw* and M™»lal 2011. Imports veeropazhnii

(Balsaminaceae), a new scapigerous balsam from Wayanad. Western Ghats. India. J. Bob Res. Inst Texas 5:1 53-158.

Bhaskar, V. 1 981 . The genus Impatiens in South In

Bhaskar, V. 2006. Impatiens clavata Bhaskar sp. nov. A new scapigerous

Ghats, South India. Cur. Sci. 91 (9):1 1 38-1 1 40.

Gamble, J.S. 1 91 5. Flora of the Presidency of Madras 1 :1 38-145. West N

Bishen Singh Mahendra Pal Singh, Dehra Dun. . ......

Gbey-Wiison, C. 1 980. Impatiens of Africa. Morphology, pollination and pollinators, ecology phytogeography, hybridiasa-

tioit keys and a systematic treatment of all African species, with a note on collection and conservation. AABalltema,

Rotterdam.

Hooker, J.D. 1 908. An epitome of the British Indian species of Impatiens. F

Jyosna, R.N. Dessai, Laly Joseph, and M.K. Janartha

Western Ghats, India. Taiwania 54:149-151. ^ ...

Mabberley, DJ. 2008. Mabberley's plant-book, ed. 3, 89, 420, 429.

Kerala, India. Rheedea 6:51-54.

Nair, N.C. 1 991 . Endemism on the Western Ghats with special reference to Impatiens L. In: C. Karunakaran,

ings of the Symposium on rare, endangered and endemic plants of the Western Ghats. Kerala Forest I

Kerala. Sp. Publ. No. III. Pp. 93-102.

118

Journal of the Botanical Research Institute of Texas 6(1)

Ratheesh Narayanan, M.K. 2010. Floristic study of Wayanad district with special emphasis on conservation of rare and

threatened flowering plants, (unpublished) Ph.D. Thesis submitted to University of Calicut, Kerala, India.

Viswanathan, M.B and U. Manikandan. 2003. A new species of Balsaminaceae, Impatiens tirunelvelica, from Peninsular India,

Bull. BotSurv. India. 45:1 -4.

Vivekananthan, K., N.C. Rathakrishnan, M.S. Swaminathan, and L.K. Ghara. 1997. Balsaminaceae. In: P.K. Hajra, VJ. Nair and P.

Daniel, eds. Flora of India (Malpighiaceae-Dichapetalaceae) Vol. 4. Botanical Survey of India, Calcutta. Pp. 95-229.

COMMELINA CLAVATOIDES (COMMELINACEAE),

A NEW SPECIES FROM KERALA, INDIA

Sheba M. Joseph, A. Anna Ancy Antony, and Santhosh Nampy*

Plant Systematics & Floristic Laboratory

Department of Botany, St Joseph's College

-

120

A MID-TERTIARY FOSSIL FLOWER OF SWIETENIA (MELIACEAE)

IN DOMINICAN AMBER

George 0. Poinar, Jr.

124

Journal of the Botanical Research Institute of Texas 6(1)

has proven to be controversial, the oldest proposal being 45-30 mybp, based on coccoliths (C6pek in Schlee

1999) and the youngest being 20-15 mybp, based on foraminifera (Iturralde-Vinent & MacPhee 1996). The

amber is principally deposited in turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey Group

(Draper etal. 1994).

Swietenia dominicensis K.L. Chambers & Poinar, sp. nov. (Figjs. 1-3). Type: HISPANIOLA. Dominican Republic: amber

mine in the northern mountain range (Cordillera Septentrional), 1995, unknown amber miner s.n. (holotype: catalogue number Sd-

9-7, deposited in the Poinar amber collection maintained at Oregon State University, Corvallis, Oregon 97331, U.S.A.).

Flower putatively staminate, total length (pedicel apex to tip of filament tube appendages) 3.85 mm, calyx shal-

lowly cupulate (Fig. 2), ca. 1.0 mm long, fused portion 0.54 mm long, lobes 0.45 mm long, broadly rounded,

spreading, margins ciliolate, petals 5, free, ca. 4.5 mm long, 2.2 mm at widest part, spreading or reflexed, some-

what contorted, elliptic to oblanceolate, obtuse, glabrous (including margins), filament tube urceolate (Fig. 1),

glabrous, 2.9 mm long excluding appendages, 2.9 mm wide, appendages 10, deltoid-acuminate, ca. 0.8 mm

long, slighdy spreading, anthers 10, sessile, alternating with appendages, ca. 0.6 mm long (only a few visible for

measurement), dehisced, hypothesized to be fertile, pistil present, ovary not visible, style exserted from fila-

ment tube, barely exceeding appendages, stigma enlarged, disc-shaped, 1.3 mm in diameter, densely minutely

pitted, obscurely marked into 5-6 fused segments (Fig. 3).

Etymology. — From the amber’s source in the Dominican Republic.

We hypothesize that the flower is staminate in function, although its anthers are dehisced and are not well

enough displayed to be sure that pollen was produced. The flower’s overall shape, including the position and

morphology of the stigma, matches well the excellent drawings of staminate flowers of Swietenia mahagoni pub-

lished by Miller (1990, p. 478). We also compared it with illustrations of the other modem species, S. humilis and

S. macrophylla, in Styles (1981) and with the description and photographs of the Mexican fossil species, S. mio-

cenica, in Castaiieda-Posadas and Cevallos-Ferriz (2007). Together with the examination of herbarium material

of S. mahagoni, these references support our judgment that the Dominican fossil is best placed in Swietenia.

As noted by Styles (1981) and other authors (e.g. Helgason et al. 1996), the 3 modem species of the genus

are “poorly defined biologically,” are largely allopatric at the present time, and are “reasonably distinct mor-

phologically” (Styles 1981). Putative hybrids, from the natural zones of contact in Costa Rica and Guatemala

involving S. humilis and S. macrophylla, are intermediate for both ecological tolerance and morphological char-

acters such as rachis length and leaflet size (Whitmore & Hinojosa 1977; Helgason et al. 1996). An extensive

polyploid series of chromosome numbers is reported for the genus (Khosla & Styles 1975; Helgason et al.

1996). A further complication is that S. mahagoni, in particular, has been widely planted outside its original

range in southern Florida and the larger Caribbean islands, thereby increasing the occurrences of species sym-

patry. The key distinction that we relied on to separate the present fossil from all other species is its combina-

tion of a ciliolate calyx with glabrous-margined petals. Swietenia mahagoni is described as having glabrous ca-

lyx lobes and petals, while the other 2 species under discussion have ciliolate petals and calyx (Styles 1981)-

Miller’s remark (1990, p. 479) that “this distinction was not apparent in numerous specimens . . . that I studied’

is offset by his statement, perhaps in error, that the calyx and corolla are ciliolate in S. mahagoni and smooth-

margined in the other 2 species. Nonetheless, the differences in perianth pubescence described by Styles (of*

cit.) have held true in practice (Pennington, pers. obs.).

Castafleda-Posadas and Cevallos-Ferriz (2007) differentiated Swietenia miocenica from this Dominican

fossil, which they saw illustrated, but not described, in Poinar and Poinar (1999), by differences in the shape of

the filament tube and petals. In addition, they described S. miocenica as having “a free calyx composed of five

. . rounded lobes . ..,’’ which would differentiate it from all other known species of the genus. To their species

characterization we have added the lack of petal ciliolation in S. dominicensis versus its presence in S. mioceni-

ca. The age of S. miocenica given by these authors (Late OUgocene-Early Miocene, 26.0-22.3 mybp ) appn*i'

Chambers and Poinar, /

iir bubbles. Scale bar = 1.5 m

126

Journal of the Botanical Research Institute of Texas 6(1)

mates that of S. dominicensis. Nonetheless, we conclude that S. dominicensis is best described as a separate

species, especially given the different provenance and the morphological details of calyx, filament tube, and j

perianth pubescence.

Swietenia is placed in the monophyletic subfamily Cedreloideae (formerly Swietenioideae) by Muellner et j

al. (2003, 2006). However, compared with the tribe Cedreleae, composed of Cedrela and T dona (Muellner et al

2009), the remaining tribes Swietenieae and Xylocarpeae, as described by Pennington and Styles (1975),

not monophyletic (Muellner et al. 2003, 2006). Capuronianthus had previously been classified as a separate

subfamily by Pennington and Styles (1975), but was later shown to be nested within Cedreloideae (Muellner et

al. 2003). According to Muellner et al. (2003), “some genera and most tribes can only be diagnosed by usings j

combination of several characters.” A well resolved and sufficiently supported molecular phylogenetic recon-

struction including all genera of Cedreloideae (compare Muellner et al. 2011 for a recent test of more DNA

markers in Cedrela, Swietenia, and other genera), along with a detailed morphological re-investigation, will ul-

timately lead to a new and robust tribal classification within Cedreloideae.

Muellner et al. (2006) discussed the fossil history of Meliaceae as a whole, with later reviews, in

detail, for tribe Cedreleae (Muellner et al. 2009, 2010; Pennington & Muellner 2010). Fossils of Cedreleae ha#

been reported from the Eocene and later periods, at both northern and mid-latitude sites in the Old and New

Worlds. In proposing a West Gondwanan origin for family Meliaceae (Muellner et al. 2006), the authors diS'

cussed an “out-of-Africa” scenario, “with dispersal. . ..[via] important routes across Eurasia and between Eur-

asia and North America provided by Beringia and the North Atlantic land bridge, and between North AmerK*

and South America through island chains and/or direct land connections.” A crown group age for Swietenia**

Chambers and Poinar, /

not yet been proposed. Based on fossil evidence (Europe, North America) and relaxed molecular clock dating,

crown Cedreleae are believed to date back at least to the Early Eocene (Muellner et al. 2010; Pennington &

Muellner 2010). The Late Oligocene-Early Miocene age of our fossil and that of Castaneda-Posadas and Ceval-

los-Ferriz (2007) verifies the Mid-Tertiary occurrence of Swietenia in Central and South America.

ACKNOWLEDGMENTS

We thank the herbaria of Harvard University and The New York Botanical for their loan of specimens used in

this study. The valuable review comments of Terence D. Pennington and Alexandra N. Muellner are gratefully

acknowledged.

REFERENCES

Castaneda-Posadas, C. and S.R.S. Cevellos-Ferriz. 2007. Swietenia (Meliaceae) flower in Late Oligocene— Early Miocene

amber from Simojovel de Allende, Chiapas, Mexico. Amer. J. Bot. 94:1 821-1 827.

Chambers, K.L and G.O. Poinar, Jr. 2010. The Dominican amber fossil Lasiambix (Fabaceae: Caesalpinioideae?) is a Licania

(Chrysobalanaceae). J. Bot. Res. Inst. Texas 4:217-218.

Chambers, K.L, G.O. Poinar, Jr., and A.E. Brown. 201 1 a. A fossil flower of Persea (Lauraceae) in Tertiary Dominican amber. J.

Bot. Res. Inst. Texas 5:457-462.

Chambers, K.L., G.O. Poinar, Jr., and A.E. Brown. 2011b. Two fossil flowers of Trichilia (Meliaceae) in Dominican amber. J. Bot.

Draper, G., P. Mann, and J.F. Lewis. 1 994. Hispaniola. In: S. Donovan and TA. Jackson, eds. Caribbean geology: an introduc-

tion. The University of the West Indies Publishers' Association, Kingston, Jamaica. Pp. 1 29-1 50.

Helgason, T., SJ. Russell, A.K. Monro, and J.C. Vogel. 1 996. What is mahogany? The importance of a taxonomic framework

for conservation. Bot J. Linn. Soc 1 22:47-59.

lTURRALDE-ViNENT,M.A.ANDR.D.E.MACPHEE.1966.AgeandpaleogeographicoriginofDominicanamber.Science273:1850-1852.

Khosla, P.K. and B.T. Styles. 1 975. Karyological studies and chromosome evolution in Meliaceae. Silvae Genet. 24:73-83.

Miller, N.G. 1 990. The genera of Meliaceae in the southeastern United States. J. Arnold Arbor. 71 :453-486.

Muellner, A.N., R. Samuel, S.A. Johnson, M. Cheek, T.D. Pennington, and M.W. Chase. 2003. Molecular phylogenetics of

Meliaceae (Sapindales) based on nuclear and plastid DNA sequences. Amer. J. Bot. 90:471-480.

Muellner, A.N., V. Savolainen, R. Samuel, and M.W. Chase. 2006. The mahogany family 'out-of-Africa'’: divergence time esti-

mation, global biogeographic patterns inferred from plastid rbcL DNA sequences, extant, and fossil distribution of

diversity. Molec. Phylogen. Evol. 40:236-250.

Muellner, A.N., T.D. Pennington, and M.W. Chase. 2009. Molecular phylogenetics of neotropical Cedreleae (mahogany

family, Meliaceae) based on nuclear and plastid DNA sequences reveal multiple origins of ‘Cedrela odorata." Molec.

Phylogen. Evol. 52:461-469.

Muellner, A.N.,T.D. Pennington, A.V. Koecke, and S.S. Renner. 2010. Biogeography of Cedrela (Meliaceae, Sapindales) in

Central and South America. Amer. J. Bot 97:51 1-518.

Muellner, A.N., H. Schaefer, and R. LaHaye. 201 1 . Evaluation of candidate DNA barcoding loci for economically important

timber species of the mahogany family (Meliaceae). Molec. Ecol. Res. 1 1 :450-460.

Pennington, T.D. and B.T. Styles. 1975. A generic monograph of the Meliaceae. Blumea 22:419-540.

Pennington, T.D. and A.N. Muellner. 201 0. A monograph of Cedrela. DH Books, Sherborne, UK. P. 1 1 2.

Poinar, G.O., Jr. 1 991 . Hymenaea protera sp.n. (Leguminosae: Caesalpinioideae) from Dominican amber has African af-

finities. Experientia 47:1 075-1 082.

Poinar, G.O., Jr. and R. Poinar. 1 999. The amber forest. Princeton University Press, Princeton, N J.

Poinar, G.O., Jr. 2002a. Fossil palm flowers in Dominican and Mexican amber. Bot. J. Linn. Soc. 1 38:57-61 .

Poinar, G.O. Jr. 2002b. Fossil palm flowers in Dominican and Baltic amber. Bot. J. Linn. Soc 1 39:361 -367.

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 .

cence in Dominican amber. J. Bot. Res. Inst. Texas 2:1 167-1173.

Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C, 28.

Styles, B.T. 1981 . Swietenia. ln:T.S. Pennington, B.T. Styles, and D.A.H. Taylor. Meliaceae. FI. Neotropica 28:390-406.

W htmore, J.L. and G. Hinojosa. 1977. Mahogany ( Swietenia ) hybrids. Forest Service Research Paper ITF-23. Rio Piedras,

Puerto Rico: USDA Forest Service Institute of Tropical Forestry.

128

Journal of the Botanical Research Institute of Texas 6(1)

BOOK REVIEW

Charles M. Allen, Kenneth A. Wilson, and Harry H. Winters. 2011. Louisiana Wildflower Guide. (ISBN: 978-

0-9718625-3-1, pbk.). Allen’s Native Ventures, LLC, 5070 Hwy 399, Pitkin, Louisiana, 70656, U.S.A.

(Orders: www.nativeventures.net, 337-328-2252). $30, 246 pp., 500 + color photos.

Louisiana Wildflower Guide is a welcome addition to the libraries of Louisiana botanists. Previously, profes-

sional botanists and plant lovers in Louisiana relied on the broader regional field guides for identifying local

plants. With this publication, Louisiana finally has a dedicated field guide to its abundant wildflowers.

The book is divided into sections for monocots and dicots and then alphabetically by family and genus.

Each plant gets a paragraph that includes a technical description with notes on habitat, parish records, and

synonymy. Each species also has a high-quality color photograph. Those trying to identify and learn plants for

the first time will appreciate the many charts provided to help narrow down your identification. For example,

some of the charts included, list monocot genera that are annuals, list of monocot genera that are scapose, list

of dicot genera with lavender flowers, and several other lists. This will lead the identifier to the index to find the

page with photo and description. Because the authors chose not to arrange the book by flower color, like so

many field guides commonly do, this is a nice format. Indeed, flower color can vary by species and among

genera. Further, field guides organized by flower color can disassociate related plants. With the format of this

guide, the user is more likely to narrow the plant down to the generic level, thus leading them to the section of

the book where all the species of that genus are discussed. The book also contains color photos of various

habitat types (marsh, prairie, swamp, pine forest), a glossary of botanical terms, and line drawings of many of

the botanical terms.

As stated in the book’s introduction, the authors have aimed to provide a guide to Louisiana’s wildfloweis

that “... split(s) the difference between the technical and too simple.” The book succeeds in doing so and I ex-

pect it will gain wide readership considering that All About Louisiana Wildflowers by Jan Midgley (2003) is ;j

geared toward gardeners and Wildflowers of Louisiana and Adjoining States by Clair Alan Brown (1980) is long

out of print. I would recommend this book to anyone working in Louisiana and East Texas. It is small enough

to fit in your backpack, but covers enough species to take it beyond a simple roadside plants guide.

— Kevin Janni, Botanist, SWCA Environmental Consultants,

3901 Arlington Highlands Blvd, Suite 200, Arlington, TX 76018, kjanni@swca.com and Research Associate,

Botanical Research Institute of Texas, 1 700 University Drive, Fort Worth, TX 76107-3400, U.SA

BOOKS RECEIVED/REVIEWS FORTHCOMING

Zsolt Debreczy and IstvAn RAcz. Edited by Kathy Musial. 2011 . Conifers Around the World. Volumes 1 and 2.

(ISBN: 978-963-219-063-1 [vol. 1], 978-963-219-064-8 [vol. 2], 978-963-219-061-7 [vols. 1-2], hbU

DendroPress, Ltd., Budapest, Hungary. (Orders: www.dendropress.com). $250.00, 2 volumes, 1089 pp

474 range maps, nearly 1300 illustrations, more than 3700 color photos, 9.25" x 10.25".

Stunningly beautiful!

Michael Wojtech. Forword by Tom Wessels. 2011. Bark: A Field Guide to Trees of the Northeast. (ISBN: 978-

1-58465-852-8, pbk.). University Press of New England, One Court St., Suite 250, Lebanon, New Hump-

shire 03766, U.S.A. (Orders: www.upne.com). $24.95, 264 pp., color throughout, 5.5" x 8.75".

CORRIGENDA:

A NEW SPECIES OF ERIGERON (ASTERACEAE)

FROM SOUTHWESTERN OREGON

Kenton L. Chambers

Department of Botany and Plant Pathology

Oregon State University

2082 Cordley Hall

Corvallis, Oregon 97331-2902, U.SA

Corrigenda

In my paper A new species of Erigeron (Asteraceae) from southwestern Oregon, J. Bot. Res. Inst. Texas 5:415-419,

2011, the species epithet in the description, p. 415, was misspelled as “stansellii” rather than the correct

“stanselliae.”

REFERENCE

Chambers, K.L. 201 1 . A new species of Erigeron (Asteraceae) from southwestern Oregon. J. Bot. Res. Inst. Texas 5:41 5-41 9 .

J-tatlte. Inst Texas 6(1): 129.2012

130

Journal of the Botanical Research Institute of Texas 6(1)

BOOK REVIEW

John W. Tunnell Jr., Jean Andrews, Noe C. Barrera, and Fabio Moretzsohn. 2010. Encyclopedia of Texas

Seashells: Identification, Ecology, Distribution, and History. (ISBN: 978-1-60344-141-4, hbk.). Texas

A&M University Press, John H. Lindsey Bldg,, Lewis St. 4354 TAMU, College Station, Texas 77843-

4354, U.S.A. (Orders: www.taraupress.com, 800-826-8911). $50.00, 512 pp., 8.5" x 11".

Contents: 1) Shells in Texas Coastal History. 2) Chronology of Marine Malcology in Texas. 3) Molluscan Ecology

and Habitats. 4) Collecting Seashells. 5) General Features of Mollusks. 6) Texas Seashells. 7) Class Aplacophora;

a-Class Polyplacophora (Chitons); b-Class Gastropoda (Snails); c-Class Cephalopoda (Squid, Octopus);

d-Class Bivalvia (Bivalves); e-Class Scaphopoda (Tusk Shells). 8) Appendix: Outline of Classification and

Checklist of Texas Seashells. 9) Glossary. 10) References. 11) Index.

Encyclopedia of Texas Seashells is just that: an encyclopedia providing everything an amateur, naturalist, or

researcher would need regarding Texas seashells. While perhaps written with a scientific audience in mind, this

text is accessible to a wide range of audiences. This book is 8.5 x llinches, hardbound, and the size and weight

of a textbook; it is not a field guide by any means. The book is full of 987 color photographs, 12 black and white

photos, 15 maps, 18 line drawings, several figures and tables, and a complete glossary, bibliography, and index.

Chapter 1, Shells in Texas Coastal History, provides a brief summary of geologic history of coastlines and

how they formed, as well as a summary of the uses (construction, tools, ornaments, food, etc.) of seashells dat-

ing back to the early Archaic era (over 4,500 years ago).

Chapter 2, Chronology of Marine Malacology in Texas, provides a list of texts on malacology in chrono* 1

logical order, and then discusses some of the scientists instrumental to the field.

Chapter 3, Molluscan Ecology and Habitats, provides a physical description of the Texas coast with black

and white maps and a few color photographs. This section describes the bays and estuaries and the rivers that

flow into them. The next section discusses the factors that affect mollusk distribution, e.g. temperature, seasons,

salinity, turbidity, etc. There are ecological descriptions of habitats (coastal marshes, bay bottoms, reefs, sea

meadows, flats, jetties, beaches, banks, etc.) where mollusks can be found. These descriptions include species of

mollusk common in that habitat, typical vegetation type, temperature, salinity, and predator-prey interactions

within these habitats. Also includedarecolorplates with imagesofthespecies assemblages typicalofeachhabitat

Chapter 4, Collecting Seashells, contains very brief descriptions of how to collect, maintain, buy, sell, and

trade shells. Lacking from this section is a suggested listing of necessary data if you are collecting shells for

scientific purposes.

Chapter 5, General Features of Mollusks, describes the different feature

plete with detailed, labeled images (including posterior, anterior, dorsal, an

clear, and easily understood.

Chapter 6, Texas Seashells, makes up the bulk of the text (295 pp.), son

class, family, genus, and species. The taxonomic hierarchy is at the top of each page, making it easy t<

information while flipping through the text. A brief paragraph discusses each class and family. Each

description includes a photograph, the scientific name of the organism and its authorities, common

distribution, size, brief physical description, habitat, remarks, and synonymous s<

This information is given in sections with bold headings for quick reference. Fc

phenotypic variation and juvenile identification are provided.

Overall this appears to be a very complete overview of Texas Seashells and “must have” reference mate-

rial for anyone interested in them. Again, this book is textbook size and is not meant to be a small field guide.

It contains clear photographs of every species with a black background and is organized in a manner that

makes it good quick reference material.

—Rebecca K. Swadek, Texas Christian University and,

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, U.SA |

ic names if applicable.

e species, sections on

J. Sot Res. Inst Texas 6(1):

WOODY SPECIES RICHNESS AND ABUNDANCE IN A TROPICAL SAVANNA

OF NORTHERN GHANA

Damian Tom-Dery

University for Development Studies

Faculty of Renewable Natural Resources

Nyankpala Campus, Box TL 1882, Tamale, GHANA

tom_dery@yahoo.co.uk

Patrick Boakye

University for Development Studies

Faculty of Renewable Natural Resources

Nyankpala Campus, Box TL 1882

Tamale, GHANA

William J. Asante

University for Development Studies

Faculty of Renewable Natural Resources

Nyankpala Campus, BoxTL 1882, Tamale, GHANA

In Ghana the savanna vegetation serves as a source of livelihood for about 30% of the population through the

provision of economic resources such as Vitellaria paradoxon (Shea), Parkia biglobosa (Dawadawa), forage and

fuelwood (Yaro 2008). Because of intensive farming, fuel wood harvesting and shortened fallow periods be-

tween cropping periods, savannas, especially those in Africa, are undergoing rapid changes in vegetation pro-

ductivity, structure and composition (Jansen 1988; Lewis & Berry 1988). The guinea savanna zone of Ghana

is estimated to cover about 60.77% of the country’s total land mass and is the most dominant vegetation type

(Anonymous 2002). Previous research carried out on the guinea savanna vegetation type^Ghana included

Vigne (1936), Taylor (1952), Lawson et al. (1969), Hopkins (1979), Houssain and Hall (1996), Oteng-Yeboah

(1996), Asase and Oteng-Yeboah (2007). All lacked an extensive documentation of diversity and abundance of

plant species in the different savanna zones. However, arecentstudy by Asase et al. (2009) mcluded abundance

and distribution data of savanna woody species at the Sinsablingbini Forest Reserve, located 20 km from

Tamale, the Northern Regional capital.

Journal of the Botanical Research Institute of Texas 6(1)

The Damongo Scarp, unlike Mole National Park and Kenikeni forest, is the smallest reserve in the West

Gonja District of Northern Region, Ghana. Some research has been carried out on the major reserves in the

district, especially Mole National Park with monitoring programs initiated since the 1960s to the 1970s (Hall

&Jenik 1968; Pegg 1969; Jamieson 1972).

Botanical assessments such as floristic composition, diversity and structure studies are vital in the frame-

work of understanding the scope of plant diversity in various ecosystems (WCMC 1992; Addo-Fordjour et al

2009). They are helpful for the estimation of sustainability of ecosystems since they play a major role in the

conservation of plant species and ecosystem management (Tilman 1988; Ssegawa & Nkuutu 2006). Ecological

data obtained in this regard are not only valuable for the application of sound management practices, they are

also useful in identifying important elements of plant diversity, protecting threatened and economic species,

and monitoring the state of vegetations (Tilman 1988; Ssegawa & Nkuutu 2006; Addo-Fordjour et al. 2009).

It is in this light that this research was carried out at the Damongo Scarp to ascertain its woody species’

composition and density. Such information is urgently needed to serve as baseline data for management plans

that aim at ensuring conservation and sustainable use of savanna vegetation. Specific objectives of this study

were: (i) to determine the tree species richness and abundance of the scarp; (ii) to determine the shrub species

richness and abundance; and (iii) to determine the regeneration status of trees at the scarp.

The study was carried out at the Damongo Scarp (39.36 km 2 ) in the West Gonja District of Northern Region. It

is located north of Damongo (Fig. 1) and lies between longitude 1°3' and 2°58W and latitude 8°32' and 10°2'N.

It shares boundaries in the south with Central Gonja District, Bole and Sawla-Tuna-Kalba Districts in the West,

Wa East District in the North West, West Mamprusi in the North, and Tolon-Kumbungu District in the East

(WGDA 2008). The vegetation is categorized as guinea savanna (Dickson & Benneh 1988). The scarp has high

temperatures with the maximum occurring in the dry season. The mean monthly temperature is 27° C.

Rainfall is unimodal with an average annual precipitation of about 1144 mm.

METHODS

A total of 10 nested square sampling plots were laid. The first square plot of area 225 m 2 was laid at random and

the subsequent plots laid 50 m away from each other. Trees with diameter at breast height (DBH) of 10 cm and

above were recorded within the 225 m 2 plots. Smaller squared plots of 100 m 2 were located in the larger plots

for the survey of young trees above a height of 1.30 m and below a DBH of 10 cm. Smaller squared plots of 25

m 2 were further located within the 100 m 2 plots for regenerating plants below 30 cm diameter. All individuals

in these regeneration plots and subplots were counted by species.

All species of woody plants found rooted within each plot were identified and their individual plants

counted. The identification of tree and shrub species was based on their physiognomic characteristics and later

confirmed using relevant literature (Hutchinson & Dalziel 1957-1972; Arbonnier 2004; Hawthorne &

Jongkind 2006), and by comparison with already identified specimens at the University for Development

Studies Herbarium. All botanical nomenclature in this paper follows IPNI (2008).

RESULTS

General Findings

A total of 61 woody species (trees and shrubs) belonging to 51 genera and 21 families were identified in the

Damongo Scarp. Fifty (50) tree species belonging to 17 families were identified (Table 1) while 11 shrub species

belonging to 8 families were recorded. About 36% of woody species were of the family Leguminosae.

The woody species accumulation, which is the cumulative number of species recorded as a measure of the

sampling effort, shows that the majority of woody plants in the scarp were sampled during the study as seen on

the curve (Fig. 2) as it reaches the asymptote.

Tree species richness and abundance . — A total of 693 individual trees belonging to 20 plant families, 41

Tom-Dery et al.. Woody species richness in a tropical savanna

133

I I Regions

Fk- 1. Map of Ghana indicating the Damongo Scarp.

genera and 50 species were recorded. About 85% (Table 1) of tree species genera consisted of only one species.

Genera with two or more species were Acacia (2 species), Combretum (3 species), Lannea (2 species), Lophira (2

s pecies), Strychnos (2 species) and Terminalia (2 species). The most species rich families were Leguminosae

(40.8%), Combretaceae (12.1%), Meliaceae (9.9%) and Sapotaceae (7.9%), as illustrated in Table 2.

Tom-Dery etal.,V

Fb. 2. Woody species accumulation ci

Tiliaceae

3. Families of shrubs identified with their respective species richness and relative abundances in the Damongo Scarp.

The five most abundant trt

Vitellaria paradoxa, Burkea afrit

decreasing order of abundance. ]

ber of individual

species in the scarp, contributing about 32% of total individual trees, were

ria, Detarium microcarpum, Terminalia avicennioides, and Daniella oliveri, in

contrast, the least abundant were Mitragyna inermis, Hexalobus monopelalus,

ina, Monotes kerstingii, and Vitex doniana, representing 2% of the total num-

r 100 m 2 .

:corded. The density of woody tree species was determined to be 69.3 ti

Shrub species richness and abundance. — A total of 185 individual shrubs belonging to 8 families, 10 genera

and 11 species was recorded. The shrub family Rubiaceae recorded the highest number with 4 species, repre-

senting 36% (Table 3). The remaining plant families— Annonaceae, Celastraceae, Combretaceae,

Euphorbiaceae, Leguminosae, Olacaceae, and Rhamnaceae — all recorded just one species each. The most

abundant shrub species in the reserve were Annona senegalensis, Gardenia aqualla, and Nauclea latifolia con-

tributing about 60% of the total number of shrubs identified. In contrast, the least abundant species were

Ximenia americana, Ziziphus mauritiana, Nauclea diderrichii, and M acaranga barteri, representing about 10% of

the total species recorded. The density of woody shrub species calculated per 100 m 2 was 18.5.

Regeneration status of trees in Damongo Scarp.— The regeneration study revealed a total of 39 tree species in

all sampling sites, representing 18 families. The diameter class 0-4.99 cm (56.6%) was highest with 392 indi-

viduals, followed by the diameter class 5-9.99 cm (22.9%) with 159 trees, and lastly, big trees 10 cm and above

(20.50%) had 142 trees. This indicates that the majority of individual trees were regenerating trees (56.6%) as

illustrated in Figure 3.

Trees and shrubs which are widely distributed in the area constitute an integral component of the Damongo

Scarp (Fig. 4). The species identified at the Damongo Scarp are similar to those reported in other guinea sa-

vanna areas of Ghana (Lawson et al. 1969; Hall 1976; Schitt & Adu-Nsiah 1993; Houssain & Hall 1996; Oteng-

Yeboah 1996; Asase & Oteng Yeboah 2007; Asase et al. 2009). However, Belanities aegyptiaca, a common

woody species recorded in surrounding areas of the guinea savanna, was not recorded in this study. Some

species like Rourea coccinea, Dalbergia afzeliana, and Hoslundia opposita, recorded by Asase et al. (2009) but not

in other savanna areas, were also not found in this study.

In a study of woody plant composition in a tropical savanna in northern Ghana, Asase et al. (2009) noted

43 species belonging to 13 families. Bright (2007), in a study of gradation in density of woody species around

Lake Taakor in Tolon-Kumbungu in northern Ghana, recorded 47 species belonging to 22 families. Also

Yahaya (2007) recorded 46 tree species belonging to 20 families. Schitt and Adu-Nsiah (1993) recorded a very

high number (148) of tree species at the Mole National Park, Damongo (MNP), while Houssain and Hall (1996)

reported a lower number (90) of tree species in the same MNP. Though most of the species recorded in this

current study were recorded in all the above studies, a few different species (22%) were also recorded. This in-

dicates that floristic composition of savanna is variable, even over relatively homogenous areas (Hopkins 1979,

Lawson 1985).

Tom-Dery et al.. Woody species richness in a tropical savanna

137

The high species richness and abundance of tree families such as Leguminosae (formerly Fabaceae) and

Combretaceae in the guinea savanna have been noted elsewhere (Hopkins 1979; Asase & Oteng-Yeboah 2007;

Asase et al. 2009). In a study of three traditional groves in northern Ghana, Oteng-Yeboah (1996) also reported

the predominance of the family Leguminosae (Caesalpinaceae, Papilionaceae, and Mimosaceae) and

Combretaceae in the different groves which is similar to the results of this study. The famiUes Leguminosae

(Mimosaceae and Caesalpinaceae) and Combretaceae, found to be dominant in this study, were also common

among the dominant families reported by Bright (2007) and Yahaya (2007). However, Meliaceae, though re-

ported as dominant in this study, was not a dominant family in the other studies. Hence, the same families may

nm through the various guinea Savanna zones of northern Ghana but some may be dominant whereas others

may not. Interestingly, some families (Sapotaceae and Bombacaceae) with few species recorded very high

abundances; it follows that not all the families with many species are abundant (Asase et al. 2009) in the scarp.

It was observed that important economic trees such as Vitellaria paradoxa (Shea) and Parkia biglobosa

(Dawadawa) were among the dominant species in the reserve, which conforms with the suggestion of Yaro

(2008) that savanna vegetation of Ghana provides ecological conditions for economic trees such as Shea and

Dawadawa. Apart from Azadirachta indica and Acacia spp., which were exotic savanna species, all theother

species observed were native savanna species. The savanna vegetation is generally under threat from bushhres

(Abatania &Albert 1993; Gordon & Ametekpor 1999) and grazing animals (Winter 1991; Smith & Franks

2000) which promotes the spread of weeds. Four woody species ( AfzeliaAfricana , Lophiraalata, Naucleadider-

Khii, and Vitellaria paradoxa ) are stated as vulnerable in the IUCN red list of threatened species (IUCN 2012).

The DBH class 0-4.99 cm recorded the highest percentage (56.6%) of the total individual tree species,

indicating that most of the plant species were in the regenerating stage. Although Asase et al. (2009) reported

138

Journal of the Botanical Research Institute of Texas 6(1)

Tom-Dery et al., Woody species richness in a tropical savann*

dominant in their study, this

l Dejong (2001) as forests regener

Chokkalingam and Dejong (2001) as forests regenerating largely through natural processes after significant

human and/or natural disturbance of the original forest vegetation.

The shrub species identified on the scarp are among the species that have been reported in guinea sa-

vanna areas of Ghana (Lawson et al 1969; Hall 1976; Schitt & Adu-Nsiah 1993; Oteng-Yeboah 1996; Asase &

Oteng Yeboah 2007; Yahaya 2007; Asase et al. 2009). A total of 11 shrub species belonging to 8 different families

were recorded in the scarp. Schitt and Adu-Nsiah (1993) reported 61 shrub species while Asase et al. (2009)

recorded 19 species belonging to 8 different plant families elsewhere in guinea savanna of Northern Ghana and

Yahaya (2007) recorded 15 species belonging to 11 families.

The results suggest that generally guinea savanna zones of Ghana are less rich in savanna shrubs. Five

families— Euphorbiaceae, Annonaceae, Celastraceae, Rubiaceae, and Fabaceae (now Leguminosae), reported

by Asase et al. (2009)— were also found in this study, with Rubiaceae being the species rich family. Yahaya

(2007) recorded 15 shrub species belonging to 11 families, with Rubiaceae being the species rich family.

Similarly Bright (2007) identified 8 shrub species belonging to 5 plant families, with Rubiaceae the dominant

family. Comparatively the shrub species recorded in this study was either higher or lower than the above men-

tioned studies, however, Rubiaceae was commonly the dominant family throughout the studies. This indicates

that Rubiaceae may be the family with the largest number of shrub species in the guinea savanna zone of

Northern Ghana. The abundance of two species recorded, Annona senegalensis and Gardenia aqualla in the

reserve, confirms their frequent occurrence in the study area, as also reported previously by Schitt and

Adu-Nsiah (1993).

Densities of 69.3 trees per 100 m 2 and 18.5 shrubs per 100 m 2 were recorded in the scarp. Tom-Dery and

Schroeder (2011) stated this method of estimating density as being systematically incorrect, but used it to com-

pare survey results with those of other authors in Ghana, and it is herein used for such comparative purposes.

Yahaya (2007) recorded a density of 39 trees per 100 m 2 and 8 shrubs per 100 m 2 similar to Asase et al. (2009)

who also recorded a density of 39 trees per 100 m 2 and 15.4 shrubs per 100 m 2 . The higher density of trees than

shrubs recorded in this study compared to the two previous studies, suggests that densities of trees are far

higher than shrubs in guinea savanna areas which implies that more tree species are widely distributed in the

savanna areas than shrubs. This further implies that savanna areas are richer in trees than shrubs. This finding

confirms the report of Cole (1986) and Stott (1991) that in savannas, tree densities are higher.

To conclude, the study is the first work on the woody species abundance and species richness in the

Damongo Scarp. The assessment has revealed that the characteristics of the vegetation in the scarp were gener-

ally similar to those of other guinea savanna areas. It has also revealed that the floristic composition (species

richness) and abundance of woody species in different areas of the guinea savanna zone are unique. It is there-

fore vital to study the diversity of plants found in the different areas of the guinea savanna zone of Ghana to

effectively conserve the botanical resources of the different areas, especially unique land masses such as the

ACKNOWLEDGMENTS

We wish to sincerely thank James Braimah of AROCHA GH for all the support during the field work. We are

also grateful to Mrs. Elena Hietsch for translating the abstract and not forgetting B.N. Baatuuwei for proof read-

ing this script. And we thank the journal reviewers Michael MacRoberts and an anonymous colleague for their

detailed and critical reviews.

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istraints of legume production in the fa

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Nyankpala Agricultural Experimental Station, NAES, Tamale, Ghana. Pp. 1 70-1 81

^Forojour, P.S. Obeng A.K. Anning and M.G. Adoo. 2009. Floristic composition, stru

Ring systems of Northern

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Anonymous. 2002. National biodiversity strategy for Ghana. Ministry of Environment and Science, Accra.

Arbonnier, M. 2004. Trees, shrubs and lianas of West African dry zones. CIRAD, Margraf Publishers.

Asase, A. and A.A. Oteng-Yebeoah. 2007. Assessment of plant biodiversity in the Wechiau community Hippopotamus

Sanctuary in Ghana. J. Bot. Res. Inst. Texas 1 :549-556.

Asase, A., K.E. Patrick, andYA John. 2009. Floristic composition, abundance and distribution pattern of woody plants in a

tropical savanna in Northern Ghana. J. Bot. Res. Inst. Texas 3:309-316.

Bright, Y.B. 2007. Gradation in density of woody species around Lake Taakor in Nawuni. Unpublished BSc. Thesis,

University for Development Studies, Tamale, Ghana. Pp. 26-30.

Chokkaungam, U. and W. Dejong. 2001 . Secondary forest: a working definition and typology. Int. Forestry Rev. 3(1):19-26.

Cole, M.M. 1986. The savannas: biogeography and geobotany. Academic Press, London.

Dickson, K.B. and G. Benneh. 1988. A new geography of Ghana. Longman Group UK Limited. Longman House, Burnt Mill,

Harlow, Essex, England.

Gordon, C. and J.K. Amatekpor. 1999. The sustainable integrated development of the Volta Basin in Ghana. Volta Basin

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Hall, J.B. and J. Jenik. 1 968. Contribution towards the classification of Savanna in Ghana. Bull. J' I.F.A.N 30:84-99.

Hawthorne, W. and C. Jongkind. 2006. Woody plants of Western African forests: a guide to the forest trees, shrubs and lianes

from Senegal to Ghana. Royal Botanic Gardens, Kew Richmond Surrey, UK.

Hopkins, B. 1979. Forest and savanna, 2 nd ed. Heimemann Educational Books Ltd., London, UK.

Houssain, M. and J.B. Hall. 1 996. The trees of Mole National Park, Damongo, Ghana. Revised by C.C.H. Jongking. 2nd ed.

GIJ Press, Accra, Ghana.

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IPNI. 2008. international plant nomenclature index. Published on the internet http://www.ipni.org [Accessed 4

IUCN. 2012. IUCN Red List of threatened species. Published on the internet http://www.redlist.org [Accessed 6 May,

Jamieson, B. 1 972. Final report-wildlife resource survey team 1 970-72. Department of Game and Wildlife, Accra, Ghana.

Jansen, D.H. 1988. Tropical dry forest; the most endangered major tropical ecosystem. In: E.C. Wilson, ed. Biodiversity.

National Academic Press, Washington, DC. Pp. 130-137.

Lawson, G.W. 1 985. Plant life in West Africa. Ghana Universities Press, Accra.

Lawson, G.W., J. Jenik, and K.O. Armstrong-Mensah. 1969. A study of a vegetation catena in guinea savanna at Mole Game

Reserve (Ghana). J. Ecol. 56:505-522.

Lewis, L.A. and L. Berry. 1988. African environments and resources. Boston, Unwin Hyman.

Oteng-Yeboah, AA 1 996. Biodiversity in three traditional groves in the Guinea Savanna, Ghana. In: L J.G. Vander Maesen

et al., eds. The biodiversity of Africa plants. Kluwer Academic Publishers, Dordrecht, London. Pp. 1 88-1 97.

Pegg, PJ. 1 969. Wildlife Management in Mole Game Reserve. FAO Report TA 2623, Rome. 29.

ScHrrr, K. and M. Adu-Nsiah. 1 993. The vegetation of Mole National Park-FRMP. GWD/IUNC Project 9786, Accra. P. 48.

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vegetation management in Queensland. Brisbane, Department of Natural Resources.

Ssegawa, P. AND D.N. Nkuutu. 2006. Diversity of vascular plants on Ssese Islands in Lake Victoria, Central Uganda. African

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Stott, P. 1991. Recent trends in the ecology and management of the of the World's Savanna formations. Progress in

Physical Geography 15:18-28.

Taylor, CJ. 1962. Synecology and siviculture in Ghana. Thomas Nelson and son, Edinburgh. P. 418.

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Princeton, New Jersey.

Tom-Dery, D. and J-M. Schroeder. 201 1 .Tree species abundance and regeneration pot

ments of the Ashanti Region of Ghana. J. Bot. Res. Inst. Texas 5:733-742.

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asation and the peasant-environment debate. West

Journal of the Botanical Research Institute of Texas 6(1)

BOOK REVIEWS

Jim Kimmel Photographs by Jerky Touchstone Kimmel. Foreword by Andrew Sansom. 2011. Exploring the Brazos

River From Beginning to End. (ISBN: 978-1-60344-432-3, pbk.). Texas A&M University Press, John H.

Lindsey Bldg., Lewis St., 4354 TAMU, College Station, Texas 77843-4354, U.S.A. (Orders: www.tamu-

press.com, 800-826-8911). $24.95, 192 pp„ 72 color photos, 7 maps, 8.5" x 10".

The Brazos River crosses Texas from the northwest to the Gulf of Mexico. Along the way the longest river in

Texas travels through six of the ten major ecoregions regions of the state. The author does a marvelous job of

guiding us along the entire route of the Brazos. In words and photographs he introduces us to the plants, ani-

mals, landforms, rivers, and geology of Texas.

Starting in the dry High Plains in New Mexico the river and the author follow a winding pathway across

the High Plains, Rolling Plains, Cross Timbers, and Prairies. Both then move on to the Blackland Prairie, Post

Oak Savannah, and Gulf Prairie and Marshes regions. In each section, the author points out key features of the

river and the landscape. He thoughtfully lists state parks, historical landmarks, and places to visit.

Texas is a large and diverse state and it is difficult to grasp this diversity easily. By using this single river

the Kimmels provide the reader with a cross section of the state. Sections of the book deal with the science of

rivers and water flow, plant and animal ecology, and key agricultural activities. I especially appreciated the sec-

tions titled “Where to Experience the Land and River.” Within each region the author notes these points of

interest and invites the reader to travel to these sites.

This book is a guide for travelers, an introduction to a variety of natural sciences, and a history of the state

as it expanded from coastal settlements to the high plains. The book is a fun read and is a welcome companion

for anyone exploring the various sections of the Brazos Basin.

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas

Gil Nelson. 2011. Botanical Keys to Florida’s Trees, Shrubs, and Woody Vines: A Guide to Field Identifica-

tion. (ISBN: 978-1-56164-499-5, pbk.). Pineapple Press, Inc., P.O. Box 3889, Sarasota, Florida 34230,

U.S.A. (Orders: www.pineapplepress.com). $19.95, 208 pp., 2 b/w plates, 6" x 9".

Morphological characteristics used to identify species are defined in a 9-page glossary along with 2 pages of

line drawings for leaves. There are no photographs included, which limits the use of this book as a standalone

publication for the identification of trees, shrubs, and woody vines in Florida. Indeed, the flyer suggests that

this book should be a companion to The Trees of Florida and The Shrubs and Woody Vines of Florida. The book is

designed primarily for field use and the handheld size and light weight will facilitate carrying the book in the

field. The dichotomous keys are constructed as numbered couplets, usually less than 14 leads and based

mostly on leaf morphology. Most of the key characteristics are based on easily seen features with emphasis on

traits observable throughout the year. There are two sets of keys: one a master key to major groups of woody

plants and another to 128 families. The families are arranged alphabetically by scientific name, followed by the

author’s name but without regard to phylogeny or higher ranks of classification. Choice of family names is

based on the Angiosperm Phylogeny Group classification.

Two paginated indices, one for genus and another for common names, facilitate location of names within

the text. Native and non-native species are included and endangered, threatened, or invasive species are desig-

nated by symbols. Taxa listed include habitat descriptions and county distribution patterns. Persons with a

special need to identify Florida plants will find this book useful.

—Harold W. Keller, Research Associate,

Botanical Research Institute of Texas, 1 700 University Drive, Fort Worth, Texas 76102-4025, V.SJc

J. Bot Res. Inst Texas 6(1): 142. 2012

COMPOSICION FLORISTICA Y ECOLOGIA DEL MATORRAL SUBMONTANO

DE ROSACEAS DE LA SIERRA DE ZAPALINAME, COAHUILA, MEXICO

Juan A. Encina Dominguez, Sergio G. Gomez Perez y Jesus Valdes Reyna

En Mexico el matorral xerofilo ocupa el 40% del territorio, se distribuye en regiones con clima arido y semiari-

do (Rzedowski 1978) y, junto con los pastizales, tiene una riqueza de 6,000 especies de plantas vasculares, lo

cual representa el 20% de la flora del pais (Rzedowski 1991). Una variante de esta vegetation es el Matorral

submontano (Rzedowski 1966), el cual tambien ha sido nombrado como ' piedmont scrub " (Muller 1939), o

chaparral (Pinkava 1980; Henrickson y Johnston 1983; Valdez y Aguilar 1983) y matorral esclerofilo perenni-

Coahuila hasta el centro-sur de Mexico (Muller 1947; Rzedowski 1978; Valiente et al. 1998), en climas semia-

1 Res. Inst Texas 6(1): 143 - 156. 2012

Journal of the Botanical Research Institute of Texas 6(1)

ridos y altitudes inferiores a los 2,000 m. Se trata de un matorral denso de hasta 3 m de alto, integrado por ar-

bustos microfilos e inermes (Rzedowski 1978). En el estado de Coahuila el matorral submontano representa el

5.14% de la superficie estatal (Anonimo 2001), es frecuente en altitudes superiores a los 1,800 m, en laderas

bajas y canones de los sistemas montanosos, en sitios con suelos someros, aunque su distnbucion se extiende

hasta comunidades boscosas y en sus limites inferiores se mezcla con el matorral rosetofilo (Henricksony Jo-

hnston 1983). Las especies dominantes pertenecen a los generos: Quercus, Rhus, Diospyros, Acacia y Calia,

ademas de especies de la familia Rosaceae.

Una variante de esta vegetacion es el matorral denso inerme parvifolio de rosaceas (sensu Marroquin

1976; Arce y Marroquin 1985) o matorral submontano de rosaceas, el cual se distribuye en la Sierra de Zapali-

name entre 1,800 y 2,480 m de altitud en la ecotonia con el matorral desertico chihuahuense y los bosques de

encino (Villarreal y Valdes 1992-93; Encina et al. 2007), donde representa el 2.86% (cerca de 1,820 ha) del

macizo montanoso (Anonimo 1998). La Sierra de Zapaliname se localiza en el sureste de Coahuila, es una es-

tribacibn de la subprovincia fisiografica de la Gran Sierra Plegada en el noreste de Mexico, se ubica en la zona

de transition con el Desierto Chihuahuense (Anonimo 1998) y es un area natural protegida, decretada en 1996

como Zona Sujeta a Conservation Ecologica por el gobiemo del estado de Coahuila (Anonimo 1996). El mato-

rral submontano de rosaceas se distribuye adyacente a la ciudad de Saltillo, por lo cual los incendios forestales,

el cambio de uso del suelo y la recreation mal planificada han ocasionado que sea la comunidad vegetal mas

afectada, lo que ha contribuido a reducir su superficie y modificar la composici6n de especies. Arce y Marro-

quin (1985) consideran que la intensidad de disturbio esta en relation directa con la cercanla a la zona urbana.

El objetivo del presente estudio es determinar la composition, estructura y aspectos ecologicos del matorral

submontano de rosAceas de la Sierra de Zapaliname.

Area de estudio

La Sierra de Zapaliname forma parte de los municipios de Saltillo y Arteaga y se ubica aledana a la ciudad de

Saltillo, entre los 25°15'00" - 25°25'58.35" latitud norte y los 100°47'14.5" - 101°05'3.8 n longitud oeste (Fig. ti-

Al norte colinda con la ciudad de Saltillo, al este con la carretera 57 (Mexico - Piedras Negras), al oeste con la

carretera 54 (Saltillo - Zacatecas) y al sur esta limitada por la coordenada de latitud 25°15'. La region pertenece

a la subprovincia fisiografica de la Gran Sierra Plegada; el macizo incluye valles, planicies y montaflas. La orien-

tation de los pliegues transversales es de este a oeste, las altitudes van desde 1,590 m en el pie de monte hasta

los 3,140 m en el cerro El Penitente, su mayor elevation, y con valles intermontanos a 2,200 m. La sierra esta

disectada por canones, con pendientes abruptas y topografla accidentada. Las rocas que afloran en el area son

sedimentarias marinas del Jurasico y Cretacico, las calizas cubren 43% del area, las areniscas y los conglome-

rados 17%. Los suelos aluviales ocupan casi 30% del area, son de profundidad variable y constituyen planicies

con abanicos al pie del macizo, en los valles son profundos y con buen drenaje. Abundan los suelos de tipo lito-

sol y rendzina, ambos constituyen casi 80% de la superficie del area; los litosoles son superficiales y sobreyacen

superficial de humus, sobre roca caliza o material rico en cal en el pie de monte y valles y representan 29%. En

menor proportion se localizan los xerosoles calcicos y feozem calcarico (Anonimo 1983, 1998). ^

C(Wo) (Anonimo 1998). La caseta meteorologica mas cercana se ubica en Buenavista, Saltillo, Coah. (23°38'N,

103°38 r W, a 1588 m s.n.m.), donde se registra una temperatura media anual de 16.9° C, la precipitation media

anual de 498 mm, valores que se consideran proximos a los correspondientes a las partes bajas de la Sierra de

Zapaliname; las lluvias son de tipo convectivo, coinciden con los meses calientes del ano. Arce y Marroquin

(1985) describen 11 comunidades vegetales para el macizo montanoso, mencionan que la cubierta vegetal en

areas con exposition sur presenta matorral rosetofilo, mientras que en las partes altas se presentan bosques de

pino y oyamel, en los canones se localizan bosques de encino y en las laderas bajas de exposition norte y oeste

se presenta el matorral denso inerme parvifolio de rosaceas. Los tipos de vegetacion mejor representados son

el bosque de pino que ocupa 14.09% de la superficie total del area protegida, el bosque de pinonero 12.54%y ®

bosque de pinonero con matorral xerofilo 9.55% (An6nimo 1998).

145

’■ Ubka d«>n del area de estudio: Sierra de Zapaliname y localization de los sitios de muestreo.

Journal of the Botanical Research Institute of Texas 6(1)

MATERIALES Y MfiTODOS

Para la medicion de la vegetacion se utilizo el metodo de parcela (Mueller-Dombois y Ellenberg 1974). Se levan-

taron 23 sitios de muestreo, su establecimiento se realizd mediante la compensation por pendiente propuesto

por Medina (1983). Las especies arbustivas se evaluaron en parcelas de 75 m 2 mientras que las especies del es-

trato herbaceo se midieron en tres parcelas rectangulares de 2 m 2 por sitio de muestreo, en ambos estratos se

tomo la altura y cobertura de las especies. En cada sitio se registro la altitud y coordenadas geograficas con un

receptor de GPS, profundidad de suelo con barrena de 1.5 m, exposition topografica (con una bnijula) y pen-

diente (con pistola Haga). La textura del suelo se determino con el metodo del hidrometro y el pH se miditi

mediante la concentration de iones de hidrogeno en una suspension de suelo, agua o solution de sal a traves de

un electrodo de vidrio. Los valores de precipitation, temperatura y tipo de suelo se tomaron del Sistema de In-

formation Geografica para el Manejo y Planeacion por Microcuencas (SIGMAPLAN-Coahuila sureste) (Ano-

nimo 2003). Se recolectaron muestras botanicas las cuales se herborizaron, identificaron e incluyeron en la

coleccion del herbario ANSM de la Universidad Autonoma Agraria Antonio Narro (Holmgren y Holmgren

1990). Para todas las especies se calcularon los atributos basicos de la vegetacion y con la suma de los valores

relativos se obtuvo el valor de importancia relativa por especie (Krebs 1999). La diversidad se cuantifico me-

diante el indice de Shannon-Wiener (Ludwig y Reynolds 1988; Magurran 1988) con la ecuacion: H’ = - Epi (In

pi), donde pi es la densidad relativa de las especies en cada sitio de muestreo. Para el calculo de la diversidad se

utilizo el logaritmo natural (Magurran 1988), por lo que el indice se expresa en “nats” (Pielou 1969). La rique-

za de especies es medida y analizada en terminos del numero de especies. Se calculo la equitatividad de espe-

cies (E), la cual es una medida de la uniformidad en la abundancia relativa de estas en cada parcela, calculada

con base a Pielou (1966): E = H’/H’ max = H’/lnR, donde H’ es la diversidad actual de la comunidad, H’ max es la

diversidad maxima potencial y R es riqueza o numero de especies. Se realizo un analisis de regresidn lineal

simple para determinar la riqueza y diversidad de especies a traves del gradiente de altitud, para lo cual se uti-

lizaron los indices de diversidad y el numero de especies por sitio asi como los valores de altitud.

La flora vascular se integra por 49 familias, 144 generos y 223 especies, incluyendo taxa infraespecificos, de los

cuales el 95.96% (214 taxa) pertenecen a la division Magnoliophyta, el 3.14% (7) a Pteridophyta y el resto

(0.90%) a Pinophyta (Tabla 1); las dicotiledoneas aportan el 73.09% (163 taxa) a la riqueza floristica del mato-

rral, mientras que las monocotiledoneas el 22.86% (51). La distribution de los taxa por familia presenta dife-

rencias marcadas, ya que las 10 familias mas ricas (con seis o mas taxa) agrupan en conjunto el 67.26% de la

flora registrada y las 39 familias restantes (79.59%) contribuyen con el 32.73% de la riqueza. Asteraceae esla

principal familia con 48 taxa, le siguen Poaceae (36), Fabaceae (15), Lamiaceae y Cactaceae (9), Euphorbiaceae

y Rosaceae (7). Los generos con mayor numero de taxa son: Salvia (6), Ageratina (6), Aristida, Bouteloua, Dalea,

M uhlenbergia y Quercus (c/u con 5) (ver listado floristico en Anexo 1). Las especies arbustivas mas frecuentes

son: Lindleya mespiloides, Malacomeles denticulata. Mimosa biuncifera y Purshia plicata.

En el Tabla 2 se compara la composition (especies mas frecuentes) y la distribution altitudinal del mato-

rral submontano de rosaceas de la Sierra de Zapaliname con la del chaparral y la vegetacion esclerofila peren-

nifolia en otras localidades en Mexico. La semejanza con estas comunidades se hace evidente, ya que todas sc

ubican por arriba de los 1,600 m de altitud y se componen principalmente por especies de las familias Rosaceae

y Fagaceae; en todas ellas prevalecen: Lindleya mespiloides, Malacomeles denticulata, Rhus virens, Garrya ovaM

y Arbutus xalapensis, asi como especies de los generos Quercus y Cercocarpus. La mayor similitud se presents

entre localidades mas cercanas al area estudiada (Coahuila y Nuevo Le6n) con las cuales comparte el 75% de

las especies, mientras que con Durango y Puebla se comparten el 62% y el 50% respectivamente.

La comunidad esta dominada por elementos arbustivos, inermes, microfilos y esclerofilos, aunque tain-

bien son frecuentes plantas espinosas. Las formas de vida arboreas estan poco representadas y se distribuyen

en las partes mas altas, asi como en canones y arroyos intermitentes, por su parte, las herbaceas son abundan-

tes, sobre todo en verano, y crecen con frecuencia bajo el dosel de los arbustos y otras en espacios abiertos.

Encina etal., Rosaceae de la Sierra de Zapaliname

149

1800 1900 2000 2100 2200

Altitud (m)

Fifi. 2. Valores de importancia relai

I Mimosa biuncifera

a traves de un gradiente altitudinal.

mere de especies arbustivas y el indice de diversidad de dicho estrato (Fig. 3) se relacionan de manera positiva-

con la altitud a un promedio de incremento de 2.17 especies y 0.3 nats por sitio de muestreo cada 100 m de al-

tttud, en condiciones homogeneas de exposicidn topografica (noroeste).

La mayoria de los sitios de muestreo se ubicaron al pie de monte de la sierra, en laderas bajas y medias

(valor de pendiente de 25 y 45%), en altitudes desde 1,778 hasta 2,220 m, con exposition topografica hacia el

noroeste. El sustrato geologico esta formado a partir de rocas sedimentarias de tipo conglomerado, los suelos

^ pedregosos, moderadamente alcalinos, con valores medios en el contenido de materia organica y con una

Profundidad inferior a los 21 cm (Anexo 2).

Journal of the Botanical Research Institute of Texas 6(1)

150

| '■

discusiOn

La riqueza floristica del matorral submontano de rosaceas equivale al 6.95% de la flora de Coahuila (Villarreal

2001) y representa el 53.79% de los 409 taxa registrados por Arce y Marroquln (1985) para la Sierra de Zapali-

name. Esta comunidad ocupa el segundo lugar en riqueza de especies dentro del area protegida, superada por

el bosque de encino, el cual presenta un total de 259 especies (Encina et al. 2007); la alta riqueza espedfica es

atribuible a que se ubica en transicibn con otras comunidades como el matorral micrbfilo, rosetofilo, bosque de

pino y/o encino, ademas de que algunas especies son ruderales y su presencia se favorece por el impacto an-

tropico derivado de la cercanla con la zona urbana de la ciudad de Saltillo. Esta riqueza floristica es superior a

la del "Chaparral" de la Sierra de la Madera, Coahuila reportada por Pinkava (1980) de donde se registran 138

especies; un comparativo con la vegetacion esclerdfila perennifolia del valle de Tehuacan, Puebla, muestra altas

similitudes en numero, pues en dicha localidad se reportan 214 especies (Flores et al. 1999), la similitud flo-

ristica a nivel de familia entre estas comunidades tambien se repite, ya que en ambos Asteraceae, Fabaceae,

Lamiaceae, Euphorbiaceae, Poaceae y Rosaceae, tienen una mayor riqueza de especies; sin embargo, tambien

se observan algunas diferencias como la mayor abundancia de Poaceae y Asteraceae en Zapaliname, lo cuales

atribuible a una mayor aridez y un clima mas templado (Rzedowski 1978), ademas de que el area estudiadase

ubica en la region con mayor diversidad de Asteraceas en Coahuila (Villarreal et al. 1996). En general la com-

donde son mayoria las especies de afinidad holartica (Valiente et al. 1998). Los valores de diversidad calculados

para el estrato arbustivo (2.76 nats) son superiores a los reportados por Estrada (1998) y Huerta y Garcia (2004)

para matoiral submontano (1.03 y 1.88 nats); estas diferencias, aunadas a un moderada equitatividad (68.58%)

del estrato, asi como a una riqueza y diversidad alta de herbaceas son evidencia de la actividad antropiea en esta

comunidad (Margalef 1991; Aronson y Schmida 1992). No obstante, la alta diversidad del matorral tambien

puede ser debido a la transicibn con otras comunidades de la sierra, tal como lo mencionan Canizales etaL

(2009), quienes concluyen que la alta diversidad del matorral submontano de Nuevo Leon se debe a la for-

macion de ecotonos con otros tipos de vegetacion. Encina et al. (2007) senalan una mayor diversidad de los

bosques de encino en las partes bajas del macizo montanoso, a una altitud en la cual se da la transicibn con el

matorral submontano de rosaceas, lo cual corrobora los resultados encontrados en el presente trabajo. El incK-

mento de la diversidad en relacibn a la altitud, muestra que esta variable ecolbgica es la de mayor influencia en

la diversidad del matorral, ya que en el area la mayor altitud se traduce en un incremento de la precipitate

(obs. pers.), factor comunmente asociado a una mayor diversidad en comunidades de zonas aridas y semiaridas

(Aronson y Schmida 1992; Montaftay Valiente 1998).

El matorral de rosaceas de la Sierra de Zapaliname forma parte del matorral submontano de Rzedowski

(1966) y de manera general es equiparable a los chaparrales de Durango (Gonzalez et al. 2007), Coahuila

(Pinkava 1980), Desierto Chihuahuense (Henrickson y Johnston 1983) y Nuevo Leon (Muller 1939; Rojas

1965; Valdez y Aguilar 1983), al matorral submontano de Coahuila (Villarreal y Valdes 1992-93) y a la vegeta-

tion esclerofila perennifolia (Valiente et al. 1998; Flores et al. 1999). Si bien el comparative floristico mostro

mayor similitud entre las localidades mas cercanas, todas se caracterizan por la dominancia estructural de ar-

bustivas menores a 2 m de altura (principalmente de Fagaceae y Rosaceae), inermes y esclerofilas, con una co-

bertura del 100%, ademas de una elevada riqueza de herbaceas; no obstante, contrario a lo observado en dichas

comunidades, en el area estudiada la estructura del matorral esta dominada por especies de rosaceas, lo que ha

llevado a considerarlo como una vegetacion secundaria que se origino por la tala de bosques de pino y encino

(Arcey Marroquin 1985). Sin embargo, no hay evidencias historicas que confirmen esta aseveracion, ademas

de que los bosques de pino se localizan a mayores elevaciones y en otros sectores de la sierra; aunque dos de las

rosaceas dominantes ( Malacomeles denticulata y Purshiaplicata) creemos que su abundancia en el area se vin-

cula directamente al ciima y suelo, ya que dichas especies crecen junto con Lindleya mespiloides en climas

templado-seco y en sustratos de origen calizo (Rzedowski y Calderon 2005). La mayor densidad de estas espe-

cies se presenta en las porciones mas aridas (menor altitud o exposicion sur) y en sustratos calizos; otros indi-

cadores de mayor aridez en este matorral son la presencia de especies de afinidad xerica de las familias Cacta-

ceae y Agavaceae. Los VIR de las especies de rosaceas presentaron variaciones con respecto a la altitud, lo cual

refleja tambien la influencia de aspectos climaticos en la distribucion de la comunidad y de las especies domi-

nantes, resultados que concuerdan con Flores et al. (1999), quienes encontraron que Malacomeles denticulata

es dominante en los pisos altitudinales inferiores, mientras que Valdez y Aguilar (1983) senalan que Lindleya

mespiloides y Purshia plicata son abundantes en chaparrales ubicados en laderas altas con exposicion sur, pa-

tron altitudinal similar al que describen Rzedowski y Calderon (2005) para estas especies en el centra de Me-

xico. Aunque el estrato arbustivo inferior presento la mayor densidad y el mayor numero de especies, mientras

que el superior es el que tiene mas importancia en la comunidad dado que contribuye al incremento de la di-

versidad, esto se observo en los sitos dominados por especies de este estrato (principalmente rosaceas) donde

* present a mayor riqueza de herbaceas, entre ellas de gramineas; esto se debe a que las copas de los arbustos

modifican la disponibilidad de luz y humedad, lo cual permite el establecimiento de las especies, tal como lo

senalan Madrigal et al. (2007) quienes enfatizan la importancia del nodricismo en los matorrales mediterra-

neos de Espana. Desde una perspectiva general el matorral estudiado se encuentra dentro del rango ecologico

descrito para el matorral esclerdfilo perennifolio y los chaparrales del norte del pais, los cuales se distribuyen

altitudes superiores a los 1,5

n un horizonte petre

e y Aguilar 1983); sin embargo, diferencias como una menor precipitacion n

icion con climas templados y secos, s

o bien desarrollado (

en zonas montanosas, e_

suelos poco profundos c

$07; Pinkava 1980; Val

^ual (450 a 550 mm) hacen de esta comunidad una variante seca de dichos matorrales, donde la precipita<

P r °medio varfa entre 600 a 800 mm anuales; este deficit hidrico se traduce en una cantidad importante de

elementos xerofilos en el matorral y una baja participation estructural de generos de zonas mas templadas y

u ®edas como Quercus y Pirns, asi como en una menor altura de la vegetacion.

El matorral submontano de rosaceas del area protegida de Zapaliname es una de las comunidades mas

a 'j tadas Por actividades antropicas, debido a que se ubica en los lhnites con la zona urbana de la ciudad de

j iUo - ^ cercania ha propiciado la perdida de 74% del matorral en solo 20 anos debido a la urbanization y

el( ambio de uso de suelo (Portes 2001), actividades que tambien son consideradas como las principales causan-

* es de Perdida y diminution de la diversidad en otras areas protegidas de Mexico (Figueroa et al. 2009), asi

r°° en ec osistemas cercanos a zonas urbanas (Pisanty et al. 2009). Otros factores como el pastoreo excesivo,

tocendios, l a extraction de materiales y la recreation mal planificada han modifkado la estructura y com-

mon del matorral (Arce y Marroquin 1985), efecto que se aprecia con mayor claridad en los sitios proximos

* C ' Udad donde Mimosa biuncifera es dominante; la presencia de esta arbustiva en sitios impactados concuer-

COtl ^ observaciones de Melgoza (1977) para el matorral submontano de Nuevo Leon, quien atribuye su

154

Journal of the Botanical Research Institute of Texas 6(1)

Exposition Pendiente Tipodesuelo Clase textural

19 25.36723 100.96449

21 2537496 100.97194

22 25.38092 100.95592

23 25.37773 100.95572

Agradecemos al herbario ANSM de la Universidad Autonoma Agraria Antonio Narro las facilidades brindadas

para la realization del presente trabajo, a Jose A. Villarreal por su ayuda en la determination de las especies

colectadas. Se agradece a Oscar L. Briones, Socorro Gonzalez E., asi como a los revisores anonimos por las

valxosas sugerencias para mejorar el manuscrito. De igual forma a Felipe N. Hernandez S. por la elaboration dd

mapa del area de estudio, a Gualberto J. Perez por elaborar la Figura 2, asi como al personal del area protegida

Sierra de Zapaliname por su apoyo durante la toma de datos de campo.

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FLORA OF TINAJAS ALTAS, ARIZONA— A CENTURY OF BOTANICAL FORAYS

AND FORTY THOUSAND YEARS OF NEOTOMA CHRONICLES

Richard Stephen Felger

Herbarium, University of Arizona

Tucson, Arizona 85721, U.S. A and

Sky Island Alliance, P.O. Box41 165

Tucson, Arizona 85717, USA.

Thomas R. Van Devender

Sky Island Alliance, P.O. Box41165

Tucson, Arizona 8571 7, U.S.A. and

Herbarium, University of Arizona

Tucson, Arizona 8572 1, US. A

School of Natural Resources and the Environment

University of Arizona

Tucson, Arizona 85721, U.SA.

ABSTRACT

This flora of the vascular plants of the Tinajas Altas region, within the Lower Colorado Valley subdivision of the Sonoran

vegetation and flora are dynamic, changing even now, and have changed dramatically during the past millennia, along with

spans more than 43,000 years of plants inadvertently collected and curated by packrats ( Neotoma spp.) and more recently

by botanists. We document a present-day flora of 227 species in 175 genera and 46 families. We also document at least 119

species in 96 genera and 36 families from the fossil record and among these fossils at least 28 species in 17 genera and 6

Boraginaceae, Cactaceae, Brassicaceae^Fabaceae, Solanaceae, Euphorbiaceae, Polygonaceae, and Nyctaginaceae. The most

diverse genera are Cryptantha, Ambrosia, and Eriogonum. There are 12 non-native species in the flora, representing only

53% of the modem flora, but only Sahara mustard ( Brassica toumefortii) and Arabian grass (Schismus arabicus) are likely to

negatively impact the native plants.

The famous waterholes, the Tinajas Altas, were critical for desert travelers and prehistoric people. The Tinajas Aims

Mountains, in one of the most arid parts of North America, have one of the richest fossil records for Ice Age plants in the

Wor ^The radiocarbon-dated plant assemblages provide a detailed record of dramatic changes in geographic ranges of spe-

w isconsin. Ice Age woodlands with single-leaf pinyon ( Pirns monophylla), California juniper (Juniperus califomica), Utah

^ (Juniperus osteosperma), Sonoran scrub oak ( Quercus turbinella ), and Joshua tree (Yucca brevifolia) were at Tinajas

Abas and elsewhere in Sonoran Desert lowlands. The earliest known creosotebush (Larrea divaricata) in North America,

l8 '°°0 Years before present, from a Tinajas Altas midden, was already the modem tetraploid Sonoran Desert race.

The Tinajas Altas region encompasses 80,000 acres (32,375 hectares) adjoining the western margin of the Cabeza

£*ta National Wildlife Refuge and is within the Barry M. Goldwater Range. Scientific, cultural, and aesthetic values dictate

158

Journal of the Botanical Research Institute of Texas 6(1)

como fosiles, de los cuales al menos 28 especies, 17 generos y 6 familias ya no existen en el area. Las familias mas diversas,

fortii) y el zacate arabe (Schismus arabicus ) probablemente tendran un impacto negativo en las plantas nativas. Estas famosas

tinajas eran una fuente de agua indispensable para los viajeros del desierto y los habitantes prehistoricos. La Sierra de las

torral des6rtico moderno. Los bosques de la Edad de Hielo con Pinus monophylla, Juniperus californicaj. osteosperma,

Quercus turbinella y Yucca brevifolia se encontraban en las Tinajas Altas y en otras regiones bajas del Desierto Sonorense hace

divaricata) en Norteam6rica, 18,000 aflos antes del presente, de un deposito de rata de campo, representa la raza tetraploide

The Tinajas Altas, or High Tanks, are famous through history and were utilized for millennia by Native Amer-

icans (Broyles et al. 2012). If you were traveling across the Camino del Diablo, part of the old Yuma-Caborca

trail, you would head for the Tinajas Altas (Fig. 1). Sometimes the lower tanks would be dry and getting to the

became a focus for desert research by the likes of WJ McGee and Edgar Mearns. Meams (1907: 122) wrote,

“This important station is at the east base of the [mountains], beside the lowest of a chain of natural rock tanks,

in a steep ravine, containing an unfailing and almost inexhaustable supply of good water. The upper tanks are

easily overlooked and difficult of access, which facts afford the most plausible explanation of the loss of lives of

The Tinajas Altas Mountains are a rugged range, rising 500 meters from the desert floor of southwestern

Arizona. There are no foothills, only slabs of pale Eocene granite rising abruptly at an unlikely angle of 45 de-

grees (Fig. 4). Along a skyline fretted with spires and windows there are no trees or saguaros to provide scale,

no sense of near or far. If you camp in a comfortable arroyo near the foot of the mountains and look to the peaks

on a full-moon evening the Tinajas Altas are as brilliant as snow and as eye-popping as any great mountain

(Fig. 5). The extremely arid Tinajas Altas region is situated along the Mexican border at a remote edge of the

Barry M. Goldwater Range, about 70 km southeast of Yuma, Arizona (Fig. 6). The Tinajas Altas Mountains are

35 km long, trending southeast-northwest from the international border to the Gila Mountains, where the

granite is rather dramatically replaced by dark schist. The Tinajas Altas site (the waterholes) is about 6.5 km

north of the Mexican border and is reached by the Camino del Diablo, a graded dirt road running south from

Interstate 8 and then eastward across the Lechuguilla Valley and the Cabeza Prieta National Wildlife Refuge

In spite of many visitors, the mountains and much of the adjacent valleys remain in near pristine condition,

although the upper bajadas and various other areas continue to be severely impacted by illegal off-road driving

and other recreational activities, and more recently by the massive construction activity of the U.S. border

fence in 2008 (see Department of Homeland Security 2008).

We selected the flora area because of the rich fossil and modem botanical record and because it embraces

a variety of desert habitats, is a convenient biological and cultural zone (delineated by U. S. Fish and Wildlife

Service) and is an area of major conservation concern. The core flora area is the 80,000 acres (32,375 hectares)

of the Tinajas Altas region encompassing the Tinajas Altas Mountains from the Cipriano Pass to the Mexico

border at Frontera Canyon and the south end of the Lechuguilla Valley drained northward by Coyote Wash

and southward by La Jolla Wash (Fig. 7). We also include some plant records from the nearby Butler Mountains

tanto actuales como fosiles, son Asteraceae, Poaceae, Boraginaceae, Cactaceae, Brassicaceae, Fabaceae, Solanaceae, Euphor-

11,000 afios durante el periodo Wisconsin Medio y el Wisconsin Tardio. El registro mas antiguo de gobemadora ( Lama

moderna del Desierto Sonorense. La region de las Tinajas Altas formo parte del refugio de vida silvestre Cabeza Prieta Na-

tional Wildlife Refuge y actualmente del Barry M. Goldwater Range. Los valores cientificos, culturales y esteticos sugieren que

las Tinajas Altas deben recibir mayor atencidn y protection.

INTRODUCTION

upper tanKs couia De aaunung, ana some travelers aia not Know aoout tne upper pools or were noi auie

reach them. Missing the tinajas and their water could spell death to desert travelers. The importance of Tinajas

Altas grew during the California gold rush of the mid-nineteenth century and by the end of that century they

many persons whose bones and graves were thickly scattered about our camp” (Figs. 2 & 3).

Felgeretal., Flora ofTinajas Altas, Arizona

159

*° provide perspective for the Ice Age record. A few records from immediately adjacent areas are included when

We expect them to occur in the core area. Specific geographic information is given in the regional gazetteer by

Hfes et al. (1997, 2007) and abridged in the list of place names below.

The Tinajas Altas region is nested among five major, contiguous protected areas and a sixth de facto area

^makeup the heart of the Sonoran Desert: Organ Pipe Cactus National Monument, Cabeza Prieta National

Wi ldlife Refuge, and Sonoran Desert National Monument in Arizona, and Reserva de la Biosfera El Pinacate y

Gran D esierto de Altar in Sonora and Reserva de la Biosfera Alto Golfo de California y Delta del Rio Colorado

® Sonora and Baja California. The Barry M. Goldwater Range in Arizona is not a protected area per se but gen-

^'ly « managed as one by the Air Force and the Marines. Spanning 210 miles (338 km) from San Felipe, Baja

^niia, to just southwest of Phoenix, Arizona, these reserves cover 7,515,221 acres (3,041,410 ha), or 11.7

P^em of the Sonoran Desert, making them the largest zone of contiguous protected desert anywhere in the

ricas (Felger et al. 2007a). Felger et al. (2007b) documented a flora for this bio-network that included 845

^ Ular P lant <axa (species, subspecies, and varieties) in 439 genera and 104 families, including 85 non-native

T; Cles (Fel ger et al. 2007b). Some additional records have been discovered since then (e.g., Felger et al. 2012).

QlO(vJT° re than one - third of total flora of the entire Sonoran Desert, a region covering 100,000 mi>

,000 V) in five states in Mexico and the United States (Shreve 1951).

Journal of the Botanical Research Institute of Texas 6(1)

Plants of the Tinajas Altas region have been collected inadvertently by packrats ( Neotoma spp.) for more

than 43,000 years in fossil amberat and purposefully collected by botanists for more than a century. Together

these two lines of collecting document a unique record revealing both the deep ecological history of the So-

noran Desert and its vegetation, as explained in a section by Van Devender, and the broad, fascinating range of

modern-day plants and their species relationships, as compiled by Felger from his own field studies and those

of his colleges and predecessors. In a section on botanizing and a list of collectors, we pay tribute to the role of

curious collectors and the process of collecting, reminding readers of the fulfilling joy of discovery. Other sec-

tions on geology, climate, human history, land management, and place-names expand our knowledge and ap-

preciation of this special place and provide a baseline as we look from the deep past to the future. Sites like

Tinajas Altas will be key stations for studies of global climate change and its effects on particular biological

communities. Meanwhile, perseverant neotomas continue to gather floral data.

GEOLOGY

The Tinajas Altas Morn

ing 400-500 meters at

5 ,and higher peaks stand-

e is block-faulted,

:ar and narrow with steep slopes, rugged c

:y floor. Typical of Basin-and-Range tecto , 0 .

trends northwest, and is separated from neighboring ranges by broad alluvial valleys or plains. The geology

Tinajas Altas has been strongly influenced by movement of the North American, Pacific, and Farallon conti-

nental plates. The Tinajas Altas range batholith is mainly composed of light-colored Gunnery Range gra nite

exposed following the Laramide Orogeny (Figs. 2-4). Basaltic lava dating from the mid-Tertiary Orogeny

(16-10 mya) crowns the adjacent Raven Butte (Figs. 8 & 9) as well as Tordillo Butte visible east of Tinajas Alta*

the *•* andesite and latite cap of Cabeza Prieta Peak dates to this time, too (Kresan 2007; Shafiqullah et al.

198 °)- Within the past 5.5 to 4.5 million years rifting and subsidence created the proto-Gulf of California and

Salton Basin as well as an inland sea that reached as far as modern Phoenix, Arizona, and Las Vegas, Nevada,

and sur rounded Thuyas Altas before retreating (Ledesma-Vasquez & Carreno 2010; see Lucchitta et al. 2001,

and s P en cer & Pearthree 2001). At that time the Colorado River had a much different course than today

( ^°ung & Spamer 2001). Over the past 1.6 million years strike-slip faulting opened several off-set passes

jurough the main range (Tucker 1980), but the range shows little recent uplift or lateral movement (Biggs &

“ emse y 2000). Granite bedrock has been the source for virtually all of the sediment in the Tinajas Altas area,

fusion occurs primarily by infrequent bursts of rainfall runoff. Tributary channels are dendritic and the

woad valley floors are characterized by large, ephemeral washes that flow roughly along the central axis of the

valle ys (Biggs & Demsey 2000). Striking geologic features include tafoni, inselbergs, and embayments.

CUMate

Not and dry! Nothing defines a desert like the Sonoran more than high temperatures and the paired factors of

rainf aH and unpredictability— or variability, as shown so elegantly by Shreve (1951) and then by Ezcurra

^Rodrigues (1986) and Comrie and Broyles (2002). Climate variability is the norm, as rainfall fluctuates on

® ne scales ranging from seasons and years to millennia. Plant distributions and growth in general are delim-

Fig. 4. Upper elevations along the east side of Tinajas Altas Mountains. 1 6 March 2011. Photo by JM.

ited in the Tinajas Altas region by drought and heat, especially the seasonal drought and hot weather of late

spring and early summer. Wintertime freezing also may play a role for some plants. This brief climate sum-

mary incorporates information from Adams and Comrie (1997), Comrie and Broyles (2002), McGee (1906),

Sellers et al. (1985), and Sheppard et al. (2002), as well as our own observations. The nearest long-term weather

data, from Wellton (40 km to the north) and Yuma (about 70 km to the west), are summarized in Tables 1 and 2.

Summers are long and hot, and winters mild and relatively warm. Rainfall, however, varies wildly— with

perhaps 7.6-10.2 cm (3-4 in) average at Tinajas Altas. Rainfall is bimodal, with summer rains and winter-

spring rains, although summer rainfall is often nonexistent, rare, or substantially reduced. Summer mon-

soons, generally occurring July to September, are a northern extension of a tropical phenomenon, and greatly

reduced from neighboring regions at somewhat higher elevations to the east, such as most of the Cabeza Priew

Refuge and Organ Pipe Cactus National Monument. Summer rains average about one-third as much as the

cool-season rains. Often highly localized and violent, summer thunderstorms can bring heavy rainfall of brief

duration, but such rains tend to be sporadic, spotty, and undependable, often drenching one place and leaving

adjacent places bone dry. Occasional downpours produce flashfloods in the Tinajas Altas Canyon (Fig- 1°)>

scouring the pools and removing or preventing vegetation in or adjacent to the pools.

Fall can be dry, or the occasional hurricane-fringe or tropical depression storms in late summer and fall

( backdoor monsoons”) can dump substantial amounts of rain over great expanses, turning the desert greeir

These rams can result in spectacular development of the major perennials because the plants are often already

in an active state of growth from the summer monsoon— the soil is still relatively moist and the weather still

warm. Monsoonal thunderstorms or tropical Gulf of California hurricane-fringes on rare occasions can surpass

Feigeretal., Flora of Tinajas Altas, Arizona

T«i I.Yuma Valley climate summary. November 1, 1930 to December 31, 1992, maximum average temperature (F), averai

average total rainfall (inches). From www.wrcc.dri.edu/cgi-bin/cliMAIN.pl7azyuva (Verfied 14 July 2008).

T«i 2. Wellton, Arizona, climate summary. March 18, 1922 to December 31, 1980, maximum average temperature (F), average minimum temperature (F), and

wage total rainfall (inches). From www.wrcc.dri.edu/cgi-bin/cliMAIN.pl7azwell). (Verified 14 July 2008).

Aug Sep Oct

bridge the summer monsoon and winter rains. This happened in the flora area in late September 2003 as a re-

sult of Tropical Depression Marty, when summer a nnuals continued growing and flowering intermixed with

early germination and flowering of winter-spring (cool-season) annuals. A similar situation occurred in fall

Cool-season (winter-spring) rains, frontal storms originating in the Pacific Ocean, may begin as cooler

weather approaches in November and can variously occur off and on through March or early April. Typically

these rains are gentler than the summer rains and can dehver widespread precipitation. Sporadic El Nino years

sometimes bring exceptional amounts of winter-spring rains, which can result in spectacular displays of

spring wildflowers. This being an extremely arid region, the cool-season rains are often scanty or insignificant.

such as dui

a La Nina year, e.g., 2011 (CL1MAS 2011).

April and especially May and June become increasingly hot and dry— desiccating is „ x

^ hot weather and seasonal drought of late spring and early summer, the fore-summer of Shreve (1931), se-

vere, y limit the survival and distribution of Sonoran Desert plants. Spring annuals dry up and die and many

tees and shrubs lose their leaves and some may even perish during the driest years. High temperatures rival

^ords set elsewhere in North America and frequent winds intensify the aridity. Maximum daily tempera-

>Ures c °mmonly exceed 38-45°C (100-113°F) from late April to early October. Summer temperatures in the

region are known to sometimes exceed 48.9°C (120°F)— on 28 July 1995, Yuma reached its all-time high at

5I °C (124°F). Average July maximum temperatures are 40.6°C (105°F).

Average winter daytime temperatures are about 24°C (75°F) and average monthly minimum winter tem-

peratures are about 4.4°C (40°F). Temperatures commonly dip several degrees below freezing on a few nights

ring each of the colder months. Many species in the region are frost sensitive, e.g., Bursera microphylla and

^°pha cuneata. Certain places or microhabitats, however, can be nearly or entirely frost-free, permitting a

shelf ° f Species with subtropical affinities to thrive. Such microh

and cliff bases, and solid rock slopes and aprons that channel v

as radiant heat in winter.

anitic rock

and often provide shade in si

!t * “repressive to find the vegetation and flora so rich and varied in this extremely arid place.

HISTORY

“ terafly Tinajas Altas is in the Western Papagueria region near the intersection of three major groups: Ho-

Trincheras, and Patayan (McGuire & Schiffer 1982). Because water is a limiting factor for human ac-

ties ’ the tinajas likely have been a focus of interest since people first arrived in the region during the Paleo-

k 6 - Tinajas Altas Region, Cabeza Prieta National Wildlife Refuge, and adjacent areas, showing U.S. Interstate Highway 8 and Mexico Highway 2.

Drafted by Cathy Moser Marlett.

«*an Period (Malpais Period) that extends from 11,000 BCE or earlier to 8000 BCE. An Archaic Period ex-

fended from 8000 BCE to 800 CE. A major archaeological survey conducted at Tinajas Altas in 1998 found that

“The archeological evidence ranges in age from the Middle Archaic period (5000-2000/1000 BCE) to the

19 40s. ... There is no evidence from the Paleoindian period” (Hartmann & Thurtle 2000: xxviii). However,

archaeological evidence from the surrounding region dates from earlier times, and people likely utilized the

Slte during the Early Archaic and Paleoindian periods (Adrianne G. Rankin, personal communication 2008;

Altschul & Rankin 2008; Carpenter et al. 2008). At Tinajas Altas “Prehistoric evidence points to dominant use

® the Ceramic period (ca. 800-1850 CE) by the Patayan, those enigmatic gatherers/hunters/farmers who oc-

orpied a vast area centered on southwest Arizona and southeast California” (Hartmann & Thurtle 2001: 512).

Evidence from the historic period points to use by Hia Ced O’odham (Hartmann & Thurtle 2001; Hartmann

etal - 2007). A Hia Ced O’odham camp, known as O’ovak, was once located on the Mesa del Muerto, the terrace

t0 the east of the tanks (Ahlstrom & Lascaux 2000: 6). A trail used by Cocopahs traveling between the Colo-

^ River delta and the lower Gila River valley intersected Tinajas Altas (Broyles et al. 2012), and an extensive

"Sworkof foot trails crisscrossed the region allowing trade and travel at least as far as the Pacific Ocean, Colo-

^ Plateau, and interior of Mexico (Becker & Altschul 2008; Broyles et al. 2012). Modem Native American

a *®»l affiliations with Tinajas Altas involve as many as two dozen native nations (Fortier & Schaefer 2010).

Historically, the first Europeans known to have drunk from Tinajas Altas were Jacob Sedelmayr and his

^ m 1750. Althoughithas been speculated thatSpanish explorer Melchior Diaz went there in 1540, evidence

V*** Expeditions by Padre Eusebio Kino (1699, 1700, 1701, 1702) utilized other tinajas such as Heart,

Ube « Prieta, and Baker tanks In 1774 and 1776 Juan Bautista de Anza did camp at Coyote Water, which he

^ Pozo de En Medio but not at Tinajas Altas (Broyles et al. 2012). With discovery of gold in California in

Journal of the Botanical Research Institute of Texas 6(1)

;s north of Tinajas Alias Pass. Drafted by

Felger etal.. Flora of Tinajas Altas, Arizona

167

k *• *»» Butte, east-facing slope. Brittlebush (Encelia farinosa), desert lavender ( Hyptis albida ), and saguaro (Carnegiea gigantea). 27 April 2010.

1848 > a number of argonauts utilized the El Camino del Diablo route and sought water at Tinajas Altas. Follow-

ing the Gadsden Purchase, which added Tinajas Altas to the United States in 1854, Tinajas Altas was visited by

boundary surveyors (Emory 1987; Gaillard 1896; International Boundary Commission 1898), scientists (Mc-

Gee 1901, 1905, 1906; Meams 1907), speculators and geologists (Bailey 1963; Bryan 1925), hunters (Roosevelt

192 °; Sheldon 1993), and naturalists and travelers (Lumholtz 1912; Pumpelly 1870, 1918). The area has long

***** unpopulated and unexploited commercially but popular among campers, tourists, and desert devotees.

Management and status

The boundaries for the current flora area coincide with those of the Tinajas Altas Addition to Cabeza Prieta

National Wildlife Refuge as proposed by U.S. Fish and Wildlife Service (Master Plan Brochure 1970: map; U.S.

Fl *h and Wildlife Service 1970a) and as incorporated within the refuge for a fewmonths in 1975 by the Depart-

men t of the Interior (Tunnicliff et al. 1986: 3.14). The boundaries extended the northern refuge boundary

M^tward to a point about one mile west of Cipriano Pass and then southward to the US-Mexico border. The

Altas addition encompassed land sections in townships 12, 13, and 14S in range 17W and townships

7** 13S in range 18W, totaling approximately 80,000 acres (Master Plan, Backup Volume 1970: 62; U.S. Fish

^Wiidlife Service 1970b) and including Raven Butte, the Tinajas Altas Mountains, and portions of the Lechu-

jf^rt and Davis Plain, butthe land was not congressional^ included within the refuge. In turn because

01 lts special biological, historical, archaeological, geological, and scenic values, the Bureau of Land Manage-

ment in coordination with the Department of Defense subsequently designated much of the former addition as

**** Critical Environmental Concern (ACEC) of irregular configuration comprising about 60,500 acres

background is part of the Tinajas Altas Range. 27 April 2010. Photo by JM.

(Departments of the Air Force, Navy, and Interior 2006: 4.11), and later as a Special Natural Interest Area of

comparable though not congruent boundaries (Range Management Department 2007).

Since 1941 Tinajas Altas has been managed as part of the expansive Barry M. Goldwater Range (BMGR,

formerly known by several names including Luke Air Force Range). The U.S Marine Corps manages BMGR-

West (which includes Tinajas Altas) and the U.S. Air Force manages BMGR-East. About 90% of the Range, in-

cluding the Tinajas Altas region, remains largely unscathed by military pilot training and other national de-

fense-related purposes. The Tinajas Altas region was managed by the Yuma Field Office of the Bureau of Land

Management (BLM) in collaboration with the military until 2001. The intent of the Tinajas Altas Mountains

Area of Critical Environmental Concern (ACEC) continues to be honored by the Marine Corps and the Tinajas

Altas region has been designated as a Special Interest/Natural Area (see Departments of the Air Force, Navy,

and Interior 2006). Other initiatives have been proposed to give the area special status. “In 1974 the Depart

ment of the Interior proposed adding to CPNWR [Cabeza Prieta National Wildlife Refuge] the 79,000 acres

immediately adjacent to the Refuge’s western boundary. . . . This area, was actually added to the Refuge in a

1975 public land order. . .but was revoked several months later due to some technical flaws in the order” (T uB '

nicliff et al. 1986). Since 1998 citizens have variously proposed incorporating Tinajas Altas within a Sonoran

Desert National Park and Preserve or a biosphere reserve, incorporating Tinajas Altas within the CPNWR , <*

designating it as a Traditional Cultural Property. A major management plan was conducted under the direction

of the U.S. Marine Corps (Villarreal et al. 2011). Special designation, stricter regulations, increased staffing, a# 1

heightened management attention may be required to surmount expanding tourism, recreation, military &&

FelgeretaL, Flora of Tinajas Altas, Arizona

rises, and illegal activities such as smuggling and ensuing law enforcement. Conservation of plants and wild-

life and protection of cultural resources at Tinajas Altas are not only merited but necessary under existing

stewardship laws.

PUCES

Information for the place names is based in part on the regional gazetteer by Broyles et al. (1997, 2007), which

was based on maps and other sources using NAD 27 datum.

Borrego Canyon. A major canyon in the southern part of the Tinajas Altas Mountains. The canyon mouth is at

32°17'58"N, 114°02'25"W, 1240 ft. Borrego Tank is in the canyon at 32°17’07"N, 114°03 , 03"W, 1590 ft.

Butler Mountains. This small, extremely arid, granitic range is 6 km (4 mi) west of the Tinajas Altas Moun-

tains and adjacent to our core flora area. Its base is partially buried by drifting sands. It is a significant packrat

fossil-midden study site (Van Devender 1990) and is named for Gurdon Montague Butler, dean of College of

Mines at University of Arizona (1915-1940) and director of Arizona Bureau of Mines (1918-1940). Also

known as Las Cuevitas on some older maps in recognition of the many eroded caves (tafoni) in the granitic

rock. Summit is at 32°22'34”N, 114°12'23"W, 1169 ft.

El Camino del Diablo. This is the infamous, ancient route from Sonoyta to Yuma. It crosses the northern part

of the Pinacate region, runs through Cabeza Prieta National Wildlife Refuge to Tinajas Altas, and ends at the

Gila River. Also known as the Old Yuma-Caborca Trail, this was the route taken by some early Spanish explor-

ers, by settlers from Mexico going to the San Diego, Los Angeles, and San Francisco areas, and by many forty-

niners to the California goldfields (Fig. 1). The first motorcar trip across the Camino was in 1915 by Raphael

Pumpelly (1918) and segments of it remain a primitive road.

The U.S. Fish & Wildlife Service (2008) provides the following summary:

tarting at Caborca, Sonora, Mexico, the 250-mile trip featured rest stops at Quitovac, Sonoyta, and Quitovaquito, be-

fore plunging headlong into the expanse of desolation that lent the trail its name. This stretch between flowing wa-

ter on the Sonoyta River at Agua Dulce, and the rumor of stagnant water at Tinajas Altas, proved arduous to all, and

le lj °-tnile stretch from Sonoyta, N

taking 3 to 10 days. Water might 1

i the cooler months, the trip might be only a slow plod,

if fortune really smiled, rain could deliver pools at Las

F and people require two gallons of water a day just to

way with the single-minded thought of reaching Tinajas

:s of that leg; by one count 65 graves near Tinajas Altas

c, priano Pass. A pass that separates Tinajas Altas Mountains from Gila Moi

**** Butte. Apparently named for Cipriano Ortega, a nineteenth century reformed bandit who may have

“men cattle through this gap It was also used by smugglers and called Smuggler Pass. 32°25'35"N,

ca. 1000 ft. Another Cipriano Pass is a gap in the southern Cipriano Hills, 3 mi N of Quitobaqui-

j! ' n 0r 8 an Pipe Cactus National Monument (Broyles et al. 2007).

Wote Wash. The major axial arroyo draining the LechuguiUa Valley northward to the Gila River, though

^ it is blocked by the Mohawk Canal. It used to meet the Gila River at 32°41'32 n N, 114 0 11'12"W.

Wote Water. A waterhole on Coyote Wash 6 km northeast from Tinajas Altas, 32°19'59"N, 1 13°59'23" W , 960

lL ^orically important and known as Pozos en Medio (Broyles et al. 2012). Bryan (1925; 422) writes that

WoteWater

* 0ut 3 feet high. The sand lies in great waves, which indicates the way in which th

IIoods - The water remains in a cavity or scour depression in the hardpan that under

^inedby digging about 4 ft. A wooden signboard and the marks of coyote or huma

170

Fig. 10. A flash flood atTinajas Altas, 13 Sep 201 1 . Photo by Scott Fischer, Arizona Game and Fish Department

finding the exact spot. The water is small in amount and not very permanent and has a bac

due to decaying plant roots.

Davis Plain. Sandy plain between the Butler and Tinajas Altas mountains. Named for geologist W. M. ^

(probably William Morris Davis, 1850-1934). The plain slopes gently to the southwest and supports the great-

est number of ocotillos ( Fouquieria splendens ) in the study area. Ironwood ( Olneya tesota) and blue palo verde

( Parkinsoniajlorida ) are widespread along the washes and runnels, although many of these trees died during

the severe drought of 2003-2004. The approximate center of the plain is 32°26'15"N, 114°13’27 M W, 900 ft-

Frontera Canyon. A large canyon draining the southern part of the Tinajas Altas Mountains. The canyon was

trends northwest from the vicinity of El Sahuaro (along Highway 2 in Sonora, Mexico) and becomes a canyon

entering the mountains as it courses towards the high point in the range. The canyon wash crosses the interna

171

tional boundary, hence the name “frontera ,” coined in March 1998 by a group of hikers including Gayle Hart-

mann (personal communication, 1998). Some Nolina bigelovii in the canyon are unusually tall (see the species

accounts). The canyon mouth is at 32°15'02"N, 114 o 0r31"W, 1100 ft.

High Tanks Gate. A seldom used name for a former border “crossing” along a dirt track in a low, broad pass at

the international boundary between the Tinajas Altas Mountains on the west and the Sierra Lechuguilla on the

east, 7.2 km (4.5 mi) south-southeast of Tinajas Altas and west of Border Marker 190. The name is a misnomer,

since there was neither a gate nor a fence until 2008, when the site was obliterated by the tall, metal border bar-

rier fence (Fig. 11). 32°15’08"N, 114°0r04"W, 1100 ft.

Lamb Tank. Named for young bighorn sheep, it is a subterranean artificial water tank built for wildlife in 1995

atthe mouth of Surveyors Canyon, 32°16’26"N, 114°0r36"W, 1200 ft.

Lechuguilla Valley, Lechuguilla Desert. This is the arid valley-plain between the western margin of Cabeza

Prieta Mountains and the eastern flank of the Tinajas Altas Mountains. This piedmont has minimal topo-

graphic relief as the broad plain slopes gently from the mountains to the axial wash, which drains the valley to

the north by Coyote Wash and to the south by La Jolla Wash. The Camino del Diablo logically crosses the ap-

proximate divide for the northward and southward drainages since there is no arroyo cutting here. Lechuguilla

is a common name in Mexico for agave, here Agave deserti.

Mesa del Muerto, Mesa of the Dead. This small mesa is named for the graves, once prevalent at the site, of

travelers who perished on the trail. The mesa is a few hundred meters east of the Tinajas Altas. Access to the

Tinajas and Tinajas Altas Canyon is from the parking area on the mesa. A wide trail leads from the mesa to the

areas near the lower tinajas. McGee (1905) called it Mesita de los Muertos. 32 0 18'45"N, 114°02’53"W, 1160 ft.

Raven Butte. Distinctive basaltic butte abutting the east side of the Tinajas Altas Mountains (Figs. 8 and 9).

Tbe butte is possibly named for its raven-black color, but the Board of Geographic Names (U.S. Geological

Survey 1987) implausibly states that it was named for the large number of relatively tame ravens found there in

lhe ear ly 1900s. Summit at 32°24'15"N, 114 o 06'33"W, 1773 ft.

Raven Butte Tank. A tinaja above huge granitic rocks in a canyon south of Raven Butte, at about 1220 ft.

Surveyors Canyon. Canyon in Tinajas Altas Mountains, 1.9 mi NNW of international boundary monument

190. Mouth at 32°16'30"N, 114'01'38"W, 1160 ft. Surveyors Tank in Surveyors Canyon is an intermittent natu-

ral tinaja used in 1890s by surveyors of the international boundary (International Boundary Commission

lg 98), 3.5 mi SSE of Tinajas Altas. It may equal Engineers Tank, shown on USGS 7.5 map at 32°16'20"N,

1H °02'00"W, 1360 ft. A tank can be found there, but this site does not completely fulfill historic descriptions

(see Sheldon 1993 : 25 , 91 ).

Tinajas Altas. These are the most significant tinajas or bedrock waterholes in the region. Dependable water

raade these large tinajas “renowned in the pioneer history of the district,” according to Lumholtz (1912: 239,

3%) ' though he was displeased in January 1910 to find “pieces of cast-off clothing, rusty tin cans, and other

^wless marks of human occupancy.”

u There are nine sets of intermittent and perennial pools that hold at least 77,664 liters (22,000 gallons)

When htK (Broyles 1996; Broyles et al. 2007). An arroyo from a hanging valley (Tinajas Altas Canyon) in these

Sratmic mountains downcuts steeply through joint fractures to scour and pluck a staircase of pools. Prehis-

^ P eo Ple were camping and variously using the area for millennia (Hartmann & Thurtle 2001). Although

w aterholes are generally regarded as permanent, at times they have gone dry or nearly so and Broyles

^informs us that“even the stalwart Tinajas Altas were virtually dry in the summer of 1905 (McGee

19 06), June 1946 and winter 1957 (CPNWR files).” McGee (1906: 722) wrote, “The supply has been

***** ^failing; though singularly enough the water was practically exhausted in the summer following the

^*Phonally wet winter and spring of 1905, seemingly for the reason that the unusually prolonged and gentle

Potation served rather to fill the tinajas with sand than to sweep them clean and leave them brimming, as

£ the ordi nary rains of the region.” In fact, the lack of wetland plants (except Typha domingensis) at these tina-

*W*bly isdue to the scouring effect of flash floods and the lack of soU at the margins of the waterholes.

"kny Wayfarers arrived to find the lower tinajas dry. . . . Perilously, they clambered up steep cliffs to retrieve mere buck-

ttfuU for themselves and their struggling animals. Some victims, finding the bottom tanks dry, and unable to feebly

Ilf!

Felgeretal., Flora of Tinajas Altas, Arizona

the Tinajas Altas Mountains in June 1992 sparked my interest. I was privileged to accompany Bill Broyles for

six-day Held trip in Cabeza Prieta and the Tinajas Altas Mountains. It was the second week of June and I ex

good hikes, and of course botanical exploration. We saw n

first day when a van with Cabeza Prieta workers passed u

fiadbeen

white fruit.

generous that spring. New growth of c

had plenty of water and time for

i else the whole time, except the morning of the

the road at the east end of the refuge. The rains

:s encroaching narrow roads was full of fuzzy

I was surprised to si

egetation so lush and dense, being that the annual rainfall is so scanty

i day under 37.8°C (100°F) is rare.

% first really significant botanical reconnaissance of the Tinajas Altas region was in March 1998. In just

3 few days, Luke Evans and I covered much of the region. Luke was verifying data for a revision of the Dry

^ders gazetteer (Broyles et al. 2007). I had heard of giant tree nolinas (Vatina bigelovii, see species accounts)

^wingin the place Gayle Hartmann and friends named Frontera Canyon and Luke knew how to get there.

18 March 1998. We leave Tinajas Altas early in the morning and head for the border on the old Surveyors

^d along the east side of the Tinajas Altas Mountains. The road is a rough dirt track to the border at a

f Uce ^times called “High Tanks Gate,” although there is not a gate or even a fence. The road continues

r nora > Mexico, where there has been considerable vehicle traffic that does not continue northward— th

rf7*cks are from drug or people smugglers not driving into the U.S. We lock up the vehicle on the U.S.

Q ‘ hope « Win be here when we get back) and walk several hundred meters into Sonora, going south and

Toward, skirting the

i. border patrol helicopter 1

the sand with landing skids on the dirt road on the Mexico side. They come back later t<

INI

174

prints, indicating illegal border crossers. Mexico Highway 2, the only road in Mexico linking mainland Mexico

with the Baja California Peninsula, is only a few kilometers to the south. After walking around the southern toe

of the mountain we head northward into the large, main canyon draining southward from the Tinajas Altas

peaks, soon crossing back into the U.S. There are no border markers or a fence here and we are soon in Frontera

Canyon heading northward. This canyon is a “dead end” leading into steep, near impassible steep, rock slopes.

We come across recent trash and “El Yori 1998,” the name of a popular Mexican rock band, freshly carved

into an overhanging ironwood branch. We hike up the canyon, up and over huge boulders making up the can-

yon floor, and I record the plant species encountered. We soon find the Nolina bigelovii giants — the biggest ones

I have ever seen (see the species accounts). We record GPS readings, measure the nolinas, and 1 collect speci-

mens of the more interesting plants, putting them in plastic bags to prevent them from wilting, to be pressed

later. Luke carries out yesterday’s trash. Was it left by border crossers or smugglers waiting for nightfall?

Later in the day we are back at Tinajas Altas and climb up above the tanks. From the ridge along an indis-

tinct trail I look down onto ironwood ( Olneya tesota ) and palo verde trees ( Parkinsonia microphylla) sparsely

strung along the bottom of a high hidden canyon, which we are calling Tinajas Altas Canyon (Fig. 12). The

place looks eerily pristine, without human influence. The steep slopes across the canyon are marked with

switchback animal trails, like the kind you see where there are too many cattle. No cows here, these are from

desert bighorns (Ovis canadensis mexicana ). I remind myself that with all this tinaja water, people have been

here since Ice Age times and must have had impacts on the place. Surprisingly, I find no more than several

mesquites (Prosopis glandulosa) along the canyon bottom. We were lucky that it is a “good” spring and the

vegetation is luxurious. By the end of the day my backpack is crammed full of plant specimens in Ziploc plastk

bags, each marked with a piece of paper keyed to notes in my pocket notebook. That night and early the next

morning I fill the plant presses.

On other occasions it is so dry that even creosotebush leaves are shriveled, such as a November 2002

trip with Curtis McCasland, then ecologist at Cabeza Prieta National Wildlife Refuge and later manager of the

Refuge. Returning with Curt on October 24, 2004, the margins of Coyote Wash, at Coyote Water along the bot-

tom of Lechuguilla Valley, are like a jungle. The hard damp vertical banks, about 1 m high, are shaded in green-

ery and the soil held together with a carpet-like biological crust of blue-green algae, liverworts, soil lichens,

green mosses and the like (see Belnap 2007; Rosentreter et al. 2007). I find “semi-wetland” plants such as Ama-

FelgeretaU

176

ranthus crassipes and Cyperus squarrosus that I would not have expected, as well as robust annuals, some bigger

than 1 have ever seen. Recent tropical depression storms soaked the place while it was still warm enough for the

summer annuals to keep going or germinate and late enough in the year for the winter-spring annuals to get

started. There is a full complement of summer annuals (ephemerals) such as Amaranthus, Pectis, and Tidestro-

mia, as well as the first cool-season annuals such as Oenothera and Plantago. Green gallery walls on both sides

of the wide wash are a tangle of honey mesquite ( Prosopis glandulosa ) to about 8 m tall with heavy trunks, cat-

claw acacia ( Senegalia greggii), desert wolfberries ( Lyciumfremontii , L. macrodon, and L. parishii), brittlebush

(Enceliafarinosa), canyon ragweed ( Ambrosia ambrosioides), and numerous other species. But ironwood is

missing from this lowest part of the valley floor, the nearest ones a few kilometers away on the middle to upper

bajada closer to the Tinajas Alas Mountains, and 1 don’t have a good explanation for its absence.

Late afternoon we go to where the Camino del Diablo crosses the divide of the Lechuguilla Valley almost

at the Mexican border. There is sheet flow here but no wash has formed, for it is here that the valley drains

northward to form Coyote Wash, coursing towards the Gila River, and southward to form La Jolla Wash, cours-

ing into the sands of the Gran Desierto (Broyles et at 2007). It is humid and warm and an aromatic yellow

carpet of desert chinchweed (Pectis papposd) and other summer annuals almost completely covers the ground

between the creosotebushes, mesquites, lycium shrubs ( Lycium macrodon and L. parishii), desert saltbushes

( Atriplex polycarpa), white bursages ( Ambrosia dumosa ), canyon ragweeds, brittlebushes, and button encelias

( Enceliafrutescens ). Half-grown sphinx-moth caterpillars graze on Boerhavia plants and there is a constant

buzz of insects. The soil is glistening mud made solid by biological soil crust. Some of the annual desert daturas

(Datura discolor) are shrub-like about one meter tall.

We turn around to go to Tinajas Altas to camp for the night. It is 5:55 and we have not passed another ve-

hicle since entering the Goldwater Range south of 1-8, although a border patrol vehicle is in front of us. They are

on a jeep trail roaming around looking for illegal border-crossers on foot and in vehicles. Between the illegals

and border patrol the desert floor is being crushed and eroded.

Soon after dark a 4-wheel-drive SUV shows up with four “herpers.” They ask Curt if he knows where they

might find rosy boas ( Lichanura trivirgata) and head off with headlamps. They come back after an hour or so

and report young red-spotted toads everywhere and one subadult female chuckwalla. I ask if they are collecting

and they say, “we hardly do anymore.” Later Curt says it’s not against the law to collect most herps. They leave

and the night is peaceful except for noisy mosquitoes. Bright lights to the northeast are from aerial flares in

military maneuvers on the Goldwater Range.

The next morning, October 25, we climb nearly to the top of the Tinajas Altas Mountains, going way up

past where Luke and I went in March 1998. Again I find only a very few mesquite trees along the high canyon

above the tinajas. Barrel cacti are also scarce— we see only a few large ones on inaccessible cliff faces where

bighorn sheep cannot reach them (see species accounts for Ferocactus cylindraceus).

During the day I see remains of about half a dozen rusty military ordinances (rocket motors and other

remnants of military aerial gunnery training). A golden eagle tops a high ridge. About three-quarters of the

way to the top we see a still-standing dead nolina and estimate the trunk to be 6 m tall. On north- and east-

facing slopes the vegetation has become denser in places where there is enough soil. There are several more

very large, downed and dead nolinas. Apparently the soil eroded away and they fell over. Perhaps they are the

same age class as the Frontera Canyon giants. A bighorn sleeping-place in an eroded (tafone) cave is f«d of

fresh, smelly bighorn droppings. Farther up I find a small, localized patch of desert spike-moss (Selaginefa

arizpnica). We have not seen it elsewhere on the climb, although I would expect it since this spike-moss is com-

mon in similar habitats elsewhere in these mountains. Has it been eliminated by bighorn sheep? I am some-

what surprised to find, apart from subtle increases in vegetation density, only minor elevational differences in

the plant life from low- to high-elevation.

Before climbing down steep rough granitic rocks, I stop to look out across a vastness of desert silence-

Dark xeroriparian ribbons writhe like double helices across the distant pale brown valley floor scratched only

by the Camino del Diablo.

177

By 2008 the Tinajas Altas portion of the desert had started to make a gradual recovery due to reduced il-

legal border-crossing traffic, a result of multiple factors, including increased border patrol presence and an

economic downturn in the U.S. By mid-year the U.S. Department of Homeland Security began building a

14-foot-high metal fence along the border (Fig. 11).

On 20 November 2008, Ben Wilder and 1 drive from Tucson to Tinajas Altas, arriving at 1:40 pm. Driving

south from 1-8 on the wide graded road we pass numerous huge trucks engaged in the border fence construc-

tion They have cleared what appears to be hundreds of acres on the valley floor northeast of Tinajas Altas as a

headquarters and staging area. Looks like a moonscape.

The lowermost Tinajas Altas tank has water swarming with mosquito larvae and multitudes of tiny,

young red-spotted toads are all about. A sprawling Boerhavia coccinea is same plant among rocks in front of the

tinaja that I have seen on every visit. Bighorn scats everywhere, plus a small quantity of larger scats that appear

to be from a burro (donkey). The vegetation is somewhat dried up — it has not rained for quite a while. Ben

climbs to a lower-middle tinaja, tank #4, and finds numerous tadpoles and the elusive cattail ( Typha domingen-

sis, see species accounts). I investigate the lower slopes and adjacent areas for worthwhile records. The large

blue palo verde ( Parkinsoniaflorida ) in the canyon wash east of tinajas is gone: it was here in 2004. We camp on

the Mesa cjel Muerto east of the tinajas and the roar from border-fence construction trucks goes on all night.

The next morning we climb the steep hanging canyon just north of the tinajas. From the ridge/saddle

looking down at Tinajas Altas Canyon we see the numerous bighorn switchback trails on the lower slope

across the canyon, leading into the upper tinajas (Fig. 12). In the canyon bottom wash leading to the uppermost

tinajas we find the sand pocked with bighorn tracks and their scats are everywhere. Some saguaros ( Camegiea

gig anted) about 1 to 1.5 m tall have been bashed in by bighorn seeking moisture (Fig. 14).

On the morning of the 22nd we drive south from the Tinajas Altas area along a wide, new graded road

approximately along the old Surveyors Tank Road to the new border fence at the High Tanks Gate area (Fig. 11).

The desert along the border is scraped bare. The fence stops at the steep granitic mountain, providing a place to

walk to and from Mexico.

Returning northward, about halfway to Tinajas Aims, the road cross two large washes coursing eastward

from mountain canyons. These washes have their margins jammed full of dense green vegetation and a blaze

flowers, apparently the result of a recent flash flood, including yellow of brittlebush ( Enceliafarinosa ) and red

of hummingbird bush (Justicia calif arnica), as well as Ambrosia dumosa, A. ilicifolia, Bebbiajuncea, Brandegea

fydovii, Datura discolor, Funastrum hartwegii, Horsfordia alata, Lycium andersonii, L. brevipes, Lfremontii, L.

m acrodon, Hyptis albida, Larrea divaricata, Olneya tesota, Parkinsonia florida, and Senegalia greggii (Fig. 15). The

adjacent desert vegetation is dry and drought-stressed.

We turn east on a wide new road past a new mountain of sand being worked by bulldozers for making

c «nent for border fence posts. The border fence construction site is off-limits. We turn back and head for Tucson.

On March 25 to 29, 2010, 1 accompany Bill Broyles and Joan Scott on a field trip across the Camino del

^Wo, from Organ Pipe Cactus National Monument through Cabeza Prieta National Wildlife Refuge to the

T,ni *jas Altas region. At the Refuge offices in Ajo we see Curt McCasland, the Refuge manager. He tells us the

alon g the Camino del Diablo in the Refuge is not really wilderness anymore: emergency stations (with

^tvival water for border crossers) are every 5 miles and there is a heavy presence of Border Patrol. In camp that

n, ght we meet up with the rest of our party. There are six of us: Bill Broyles and his wife Joan Scott, a wildlife

expert retired from the Arizona Department of Game and Fish, Luke Evans, an environmental consultant and

re gional scholar, David Taylor, a Guggenheim fellow from New Mexico State University who is re-photograph-

ic the border monuments from Texas to California, and Richard Laugham, a fine art photographer from

‘J 0enix who has been tracking specific desert plants year after year. The last two days are spent in the Tinajas

* ltas region.

On the 28th, Richard Laugharn and I stop at the Camino crossing of the Lechuguilla Valley bottom and

2" ore the mini-jungle of mesquites at the beginning (southern limit) of Coyote Wash. The mesquites, mostly

^'5) m tall, are nearly impenetrable. 1 am looking for Teucrium glandulosum, which I had previously found

178

Journal of the Botanical Research Institute of Texas 6(1)

gn of it. The ground is mostly covered by the invasive bana*

e variation. The soil is still damp from last rains and where the

itinuous blackish surface of microphytic crust with blue greens

al. 2008). Milkweed vines ( Funastrum hartwegii) interlace the

but only as a dry skeleton, but there was no

mustard (Brassica toumefortii) of incredible <

ground is not covered by ephemerals 1 see a c

and bacteria (see Belnap 2007; Rosentreter ,

jungle, reaching into the tops of the mesquites. Inside the jungle the air is hot, humid, and still. Lactucaserrio-

ia and Sphaeralcea emoryi are new records for the Tinajas Alias region. Later in the day 1 am botanizing around

Tmajas Altas. Near full moon and using a headlamp I press the day’s specimens, all in good shape in Zip**

plastic bags in the frig in the back of Luke’s pickup.

On the March 29, after a quick breakfast, I go with Luke Evans in his Toyota pickup. He has a laptop floor-

mounted in the cab with topo maps on the ready and other techno conveniences. We drive south and then

westward into Surveyors Tank Canyon, to a small parking area near the mouth of canyon at Lamb Tank, an

elaborate artificial buried water catchment leading into a small cement water tank for bighorn sheep. Bigh° rn

scats are abundant as well as Sahara mustard. The tank is full and swarming with honeybees, which we hado*

seen elsewhere on the trip. Surveyors Tank is further up the canyon and is entirely natural. There are bedrock

grinding holes near the tinaja. We do not see barrel cacti and agaves are surprisingly few, probably due to big-

horn sheep—their scats are numerous.

In the late morning we drive back north to our cs

a dirt track west from the Camino into a large canyon

and hike up a canyon with huge granitic rocks. Like n

dozen conspicuous major perennials and more than

barrel cactus ( Ferocactus cylindraceus).

Leaving for Tucson in the early afternoon, 1 am i

road along the sandy Lechuguilla Valley to Wellton I

but no buffelgrass. We pass spectacular ocotillo “fort

full flower. We stop for cold drinks at a convenience

daily from San Luis, Sonora, are spending money on

all the way from Wellton to Tucson.

np near Tinajas Altas, pack up, and head north. We

n the granitic mountains. We park at the end

wo-dozen winter-spring annuals. I saw only one

ding with Bill and Joan. Going north on the

ee large plants of Sahara mustard all along tl

its” along the sandy valley floor, the ste

store in Wellton. A long line of farm workers DU ^-

ank food and pop. In the back seat I am pressing P 1 ^

sredtipP^

Felger et al.. Flora of Tinajas Altas, Arizona

179

DEEP HISTORY AT TINAJAS ALTAS, TOM VAN DEVENDER

Des erts are notoriously bad places for preservation of fossils. In wetter climes plant materials and dead animals

fall into water, are quickly buried, and minerals may replace their organic fabric. Yet the Tinajas Altas Moun-

tains ’ in °ne of the most arid parts of North America, have one of the richest fossil records for Ice Age plants in

lhe World - The humble packrat has collected and curated hundreds of species in the southwestern United

*** for more Aan 43,000 years. In 1962, Phil Wells, a plant ecologist, and Clive Jorgenson, a mammalogist,

c imbed the isolated, desolate Spotted Range on the Nevada Atomic Test Site. On the way down they entered a

l0dt shelter to eat lunch, where they found a dark, organic mass full of twigs and seeds that Wells recognized

** juniper and fecal pellets that Jorgenson knew were from packrats. It was the rich dark reddish-gold color of

^ rat ’ the Tver’s name for the shiny crystalline veneers of packrat urine on cave walls. Wells later radiocar-

-dated a piece of the deposit to more than 10,000 years ago. Wells named the deposits ‘middens,’ a term

°*r°wed horn archeologists referring to human garbage piles. The 1980s were a time of discovery— the Am-

r Rus h had begun (Van Devender 2007) These studies are summarized in Packrat Middens: The Last 40,000

^Biotic Change (Betancourt etal. 1990).

. Packrals (ako called woodrats) are medium-sized rodents that carry plant and animal materials and

""objects back to their houses or dens. In dry rock shelters some of this material on ledges or in crevices

f0m the ma in den deposit may become cemented with their urine into hard, dark organic middens (Be-

w J° Un et al 1990). These deposits reflect many years of gradual accumulation of plant material collected

1 ° nl y ab out 30 m of the rock shelters, and are excellent for reconstructing past vegetation and climate.

n February of 1979, botanists Art Phillips and Barbara Phillips from the Museum of Northern Arizona

U* ' explorin g the Tinajas Altas Mountains when they found a packrat midden in a canyon on the east side of

ltl0re ange ’ f° uth -southeast of the famous ‘high tanks’ area. In March of 1980, 1 returned to the area in search of

middens and the promise of reconstructing the history of vegetation in a new area in the Sonoran Desert.

On the map, the most direct route was from Interstate 8 east of Yuma down the west side of the Gila Mountains

to the old Fortuna Mine and southward through Tinajas Altas Pass. This was a real adventure with stark, beau-

tiful landscapes and wondrous spring wildflowers. But it was a slow trip crossing the myriad of arroyos drain-

ing the mountains and a strain on my Volkswagen Beetle and kidneys. Eventually the Tinajas Altas were

reached on the east side of the mountains where broad grussy arroyos (from grus, sand-like erosive material

from granite) adorned with blooming chuparrosas (Justicia califomica ) drain white granite slopes. As we

worked our way above the tanks, we spotted a coiled speckled rattlesnake ( Crotalus mitchelli), a medium-sized

rock dweller. This one was nearly white with black speckling — a perfect match for the pale granite. Looking

eastward, the thin ribbon of the Camino del Diablo stretched to the horizon across the broad Lechuguilla Val-

ley. Here the desertscrub is so sparse that the small desert trees are restricted to xeroriparian strips along

washes in a sea of widely spaced diminutive creosotebush ( Larrea divaricata) and white bursage (Ambrosia

Ancient packrat middens proved to be common, resulting in a remarkable fossil record (Van Devendei

1990). A total of 24 middens with 31 different stratigraphic units were found in 12 rock shelters (Table 3). Each

discrete unit in a midden was numbered and analyzed separately. A total of 30 radiocarbon dates obtained on

materials from 21 of the samples from 11 of the middens ranged in age from 245 to more than 43,000 ybp (ra-

diocarbon years before 1950). An additional 12 dates obtained from nine samples from six middens from the

nearby Butler Mountains ranged in age from 740 to 11,250 ybp. The Tinajas Altas and Butler Mountain samples

also yielded abundant animal remains, including invertebrates (Hall et al. 1988, 1989, 1990) and small verte-

brates (Van Devender & Mead 1978; Van Devender et al. 1983, 1991).

The radiocarbon-dated plant assemblages provide a detailed record of dramatic changes in geographic

ranges of species and the succession from Ice Age woodlands to modern desertscrub. Prior to 11,000 years ago,

in the middle and late Wisconsin, woodlands with single-leaf pinyon ( Pinus monophylla), California juniper

(Juniperus califomica), Joshua tree ( Yucca brevifolia), Mohave sage (Salvia mohavensis), and desert tree-bear-

grass (Nolina bigelovii) were present on the slopes near the tanks. The lowest fossil record for single-leaf pinyon

was at 460 m in the Tinajas Altas Mountains. The lowest elevation record for California juniper was 240 m ina

creosotebush desertscrub assemblage in the Butler Mountains.

The midden fossils mostly record woodland plants at low elevations, reflecting dramatic changes in veg-

etation with Sonoran Desert dominants mostly displaced to refugia, probably in central Sonora. The trough of

the Colorado River Valley likely was desertscrub throughout the Pleistocene, with much simpler vegetation

changes than at higher elevations. Middens from Picacho Peak, California (just north of Yuma) recorded con-

tinuous desertscrub for the last 12,730 years, where Mohave desertscrub with Joshua tree-blackbrush (Ck*

gyne ramosissima)-creosotebush changed to brittlebush ( Enceliafarinosd)-pygmy cedar (Peucephyllum schot-

tiO-creosotebush and was later replaced by a simpler Sonoran desertscrub (Cole 1981). The creosotebush-

white bursage desertscrub in the flats below the Tinajas Altas Mountains was probably more stable, with less

change or impact from the Ice Age climates than the vegetation on rocky slopes above at higher elevations.

Ice Age climates with greater winter rainfall from the Pacific and reduced summer monsoonal rainfej

from the tropical oceans likely favored woody cool-season shrubs with northern affinities (Neilson 1^

rather than the summer-rainfall trees, shrubs, and cacti of dry tropical origin and subtropical deserts. Pal

matic reconstructions based on the modem distributions of these cool-season species inferred greater win^

rainfall, greatly reduced summer monsoon rains, and cooler summers (Van Devender 1990). The Tinajas Al

middens showed a dramatic shift from predominantly winter annuals in Ice Age woodland assemblage*

mixed winter-summer annuals as the summer monsoon developed in the Holocene and Sonoran desertscn®

formed.

Creosotebush is one of the most important plants in the warm deserts of North America. Unlike

other Sonoran Desert dominants, it evolved in South America and immigrated to the Chihuahuan Desert .

oldest record for creosotebush in North America is a midden sample from the west base of the Tinajas Al

Mountains. A C-14 date of 18,700 ybp on its twigs documents its presence in the area during the last full glac** 1 ’

Felgeretal., Flora of Tinajas Altas, Arizona

Tinajas Alias #18B

Tinajas Alias #15A

Tinajas Altas # 18 A

Tinajas Altas # 20 A

TinajasAltas # 2 B

Tinajas Altas #3B

TinajasAltas # 3 B

Tinajas Altas # 1 B

Tmajas Altas #3A

Tmajas Altas #ia

Tm a)as Altas #16B

Tma ias Altas #16A

Tinajas Altas # 1 6A

r "»tas Altas # 13 B

r "»jasAltas#i3A

>AHas#i3B

T^jas Altas #20C

T*’ajasAftas#2A

>43,200

>37,000

18,700±1050

18,530±1070

16,150±400

1 5,680±720

15,050±350

11,040±270

10,600±420

10,300±110

10,070±110

9700±100

9230±140

8970±75

8650±430

5860±60

5820±310

5080+80

4490±230

4010±70

1 770±300

1230±100

245±250

AA-1720

USGS-959

A-3730

A-3570

USGS-957

av. AA-779 & A-8255

AA-777

A-3738

AA-525

Larrea divaricata

°f maximum development of continental and montane glaciers at high latitudes and elevations in

2oo n :* erica ’ though it surely emigrated from South America much earlier (Hunter et al. 2001; Lia et al.

Cleric eSC ICC AgC creosotebus hes were growing in a California juniper-Joshua tree woodland. In North

chr ^evolution in creosotebush resulted in the widespread Larrea divaricata subsp. tridentata with three

de^ 050 ™ 6 races roughly found in the Chihuahuan (diploid), Sonoran (tetraploid), and Mohave (hexaploid)

tj* Gran and the unusual willowy dune creosotebush (var. arenarid) in the Algodones Dunes of California and

Desiert0 of northwestern Sonora (Felger 2000). In South America there are four species of Larrea, all

**an j)_ Accordin g to Hunter et al. (2001), the full-glacial creosotebush in the Tinajas Altas was the same So-

^ Sert tetraploid chromosomal race that occurs in the area today. Surprisingly, they also reported all

182

three chromosomal races in middens from Picacho Peak, California, to the northeast across the Colorado

River from Yuma. All of the late Wisconsin creosotebush midden records were diploids or hexaploids from

below 310 m in the Lower Colorado River Valley. The first Mohave Desert hexaploids appeared in the Picacho

Peak midden record at 8420 ybp (Hunter et al. 2001). This is interesting because Vasek (1980) inferred that the

King Clone creosotebush (Sternberg 1976) in the Mohave Desert of California, with an average diameter of

about 14 m, was 11,700 years old, even though creosotebush was not found in packrat middens in the nearby

Lucerne Valley before 5880 ybp (King 1976). The King Clone may be younger than popular myth would like.

The packrat midden assemblages are rich with 25 to 60 plant species per sample. The fossil records of the

majority of the plants, which are not structural dominants, provide different insights. Holly-leaf bursage (Am-

brosia ilicifolid), as well as desert agave (Agave deserti), desert tree-beargrass, many-headed barrel-cactus (Edo-

nocactus polycephalus ), mountain barrel-cactus ( Ferocactus cylindraceus ), and Spanish bayonet ( Hesperoyucca

whipplei) were present in Ice Age woodlands in Tinajas Altas, unlike Sonoran Desert saguaros and palo verdes,

Another interesting pattern is that short-lived plants and animals were not affected as much by Pleisto-

cene climates as larger trees, shrubs, and succulents. The midden assemblages have many records of anoma-

lous associations in Ice Age woodland, including holly-leaf bursage with single-leaf pinyon, California junker.

Joshua tree, Mohave sage, and skunk bush (Rhus aromatica) in the Tinajas Altas Mountains (Van Devender

1990). The composition of plant communities has changed continuously because each individual species re-

sponds differently to climate events, and climate fluctuations range from thousands of years on continental*

global scales to months or days for regional or local droughts or freezes. Even in pristine-appearing habitats,

some species are increasing, decreasing, absent, or just arrived in response to some drought, rainfall, freeze,*

other climate event (see Amaranthus crassipes and Cyperus squarrosus in the species accounts).

Middens have provided remarkable insight into evolutionary processes. Over 400 species of plants and

small animals have been identified from Sonoran Desert packrat middens. These fossils are indistinguishable

from their living descendents, no matter what their age. Midden fossils tell us that species are well adapted®

fluctuating climates, and simply expand or shrink their ranges.

Considering that t

> 15 t(

/interglacial cycles in the Pleistocene, species d

unity compositions have been very dynamic in the last 2.4 million years. Similar successional stages

from widespread woodlands to expanded deserts likely occurred during each interglacial. Although the late

Holocene desertscrub communities likely resembled the original late Miocene Sonoran Desert, relatively®^

era desertscrub communities were developed for about 5-10% of the 2.4 million years of the Pleistocene- W

Age climates with woodlands in the desert lowlands typical of about 12,000 ybp persisted for about 80-90%

this period (Porter 1989; Winograd et al. 1997).

Middens in the Tinajas Altas Mountains yielded at least 119 plant taxa, although not all could be identi-

fied to species (Van Devender 1990, and this publication, Table 4). Of these, about 28 taxa no longer occur®

the flora, including the community dominants single-leaf pinyon, California juniper, and Joshua tree. Other

woodland and Mohave plants no longer found in the area are white sage ( Artemisia ludoviciana ), cliff golden-

bush ( Ericameria cuneata ), turpentine bush (E. laricifolia), green rabbitbrush (E. teretifolia), Spanish bayonet

mint ( Monardella arizonica), pancake prickly-pear ( Opuntia chlorotica), skunk bush, Mohave sage, andeV *f*

few winter annuals ( Calycoseris parryi, Cryptantha utahensis, Daucus pusillus, Festuca microstachys, and W

sanocarpus curvipes). Among the fossil specimens, 16 species of trees, shrubs, and succulents are shown®*

diagram in Van Devender et al. (1990). The rest are published here for the first time.

The inference from the packrat chronicles is that you look at the modem communities, and most ot

larger trees and shrubs you see have immigrated into the Tinajas Altas Mountains during the Holocene (9-

thousand years ago). Other species such as desert agave and holly-leaf bursage. Lower Colorado Valley

endemics, and many short-lived plants and small animals, probably have been here for most of the Pkist

Unlike saguaros and palo verdes, they did not retreat and then return. This scenario shows that although ®"“

era and fossil species are virtually identical, the vegetation and community composition of the region"* 5

changed dramatically and continuously through time.

Glandularia spVspp.tt

ZYGOPHYLLACEAE

; COLLECTORS

Botanists have documented the flora of Tinajas Altas for more than a century. The collectors of specimens cited

in this flora are listed here. The main repository of their collections is at ARIZ or as indicated. There are, how-

ever, more specimens from other collectors that we do not cite to avoid duplication of information (also see

Southwest Environmental Information Network 2011).

Marc A. Baker. 24 January 1999, with Ami Pate (ASU).

R.J. Blackwell. 15 April 1932, with A. B. Akens.

Rodney G. Engard.18 April 1976 (ASU), with Russell A. Haughey.

Richard Felger. 3 and 4 February 1990, with Bill Broyles, Stephen Bell, Gayle Hartmann, Paul Huddy, and Cad

Wachtmeister. 16 June 1992, with Bill Broyles. 18 and 19 March 1998, with Luke Taylor Evans. 29 No-

vember 2002, with Curtis McCasland. 10 January 2002, with Curtis McCasland. 25 and 26 October

2004, with Curtis McCasland. 18 February 2005, with Karen Louise Reichhardt. 21 February 2005,

with James L. Heard and Karen Reichhardt. 30 December 2005, with Bill Broyles and Stephen Oertle. 20

to 22 November 2008, with Benjamin Theodore Wilder. 28 and 29 March 2010, with Bill Broyles, Luke

Howard Scott Gentry. 28 October 1937. The 1<

probably from or near the actual Tinajas Altas waterholes. There are collections at ARIZ and RSA by

Gentry from 1 October 1939 that are labeled “Telegraph Pass, Tinajas Altas Mountains” and some of the

same date merely labeled “Tinajas Altas” that are presumably also from Telegraph Pass (as evidenced by

the specimens collected) and therefore not part of our flora. In 1939 the Gila and Tinajas Altas Moun-

tains were often considered to be a single range.

Edward Alphonso Goldman. 20 November 1913 (US).

Leslie Newton Goodding. 22 November 1934. 5 and 6 December 1935 with Edward W. Hardies. 17 November

1936, with Edward H. Morris. 1 December 1938. 7 March 1940, with Arthur L. Hinkley. The numbers on

his herbarium labels often represent various accession numbers of the U.S. Soil Conservation Service in

Tucson, where he worked. These numbers are not consecutive, but are sufficient to identify the Tinajas

Altas specimens.

Stephen Ferris Hale. 28 March 1981, with Frank W. Reichenbacher. 5 March 1983, with Thomas R. Van Dev-

ender and Timothy Louis Van Devender. 26 Mar 1983, with Thomas R. Van Devender.

Richard Ray Halse. 31 March 1973.

Charles F. Harbison. 6 and 9 March 1937 (ARIZ, SD). The numbers on his herbarium labels represent acces-

sion numbers at SD; he did not use collection numbers for his herbarium specimens.

George J. Harrison. 5 March 1927, with G. O. Belden. 28 and 29 March 1930, with Thomas Henry Kearney.

Wendy Caye Hodgson. 18 April 1983 and 8 March 1984 (DES), with Rodney Engard. 11 Feb 1993, (DES), with

Lynda P. Kozak, Liz Ecker, and Ted Anderson.

Thomas Henry Kearney. 29 March 1930, and 23 and 25 March 1935. On many labels it is not apparent if the

collection number is that of Kearney, George J. Harrison, or Robert H. Peebles.

Edwin Bernard Kurtz. 17 and 18 April 1948, with William Frank McCaughey and Horace S. Haskell.

Cynthia A. Lindquist. 25 and 26 March 1983. See Thomas R. Van Devender.

Steven Paul McLaughlin. 19 and 20 February 1979, with Janice E. Bowers, Martin Karpiscak, Arthur M. Phil-

lips III, Barbara G. Phillips, E. M. Peterson, and J. M. Downs.

Roger E. McManus. 29 February 1976, with Paul Fugate and Steven P. McLaughlin.

Edgar Alexander Meams. Wand 21 February 1894 (CAS/DS.US). The first set of his specimens is at US, with

duplicates at numerous other herbaria. See Meams (1907).

Gale W. Monson. 25 September 1955 (herbarium at Cabeza Prieta National Wildlife Refuge headquarters, Ajo)

J.R. Morrison. 16 April 1993 and 10 November 1993, with PJ. Walter (ASU).

Robert Hibbs Peebles. 1935 and 1940. See George J. Harrison and Thomas Kearney.

Felgeretal., Flora of Tinajas Altas, Arizona

185

Timothy Reeves.12 February 1977 with Donald J. Pinkava andJ.B. Rodriguez (ASU). “My collection numbers

are 5352-5452, so there were 101 collections that day” (Tim Reeves, personal communication, 31 July

2008). Unfortunately not all 101 seem to be at ASU, but at least 81 were accounted for in Southwest Envi-

ronmental Information Network (2011). A few Reeves collections at ASU are entered as Pinkava collec-

tions, but the numbers are Reeves’, e.g., Dudleya arizonica, Pinkava R5370.

Frank W. Reichenbacher. 9 and 10 March 1980, with Thomas R. Van Devender, Sandra Hunt Stein, etc. (see

Thomas Van Devender). Also 28 March 1981.

Forrest Shreve. 26 and 27 March 1932. 21 March 1933. Edward Tattnal “Tad" Nichols IV (2007: 89, Fig. 9.5)

shows a photo of himself, Thomas Dwight Mallery, Shreve, and Ira Wiggins at “Tinajas Altas, April

1933” (reproduced in Fig. 16). He provides a fine narrative of the trip and tells us, “The trip was about five

or six nights, with two nights at Tinajas Altas” (Nichols 2007: 88). The date, however, must have been in

March and not April.

Norman Montgomery Simmons. 21 May 1965. He was assistant refuge manager, Cabeza Prieta National

Wildlife Refuge, from 1961 to 1966, during which time he collected plants on the Refuge and adjacent

areas (Felger & Broyles 2007; Simmons 1966, 2007).

Peter Christian Sundt, Jr. 2 April 2011, with Mara MacKinnon and Jim Malusa ( Psorothamnusfremontii ; see

OIneya tesota in the species accounts).

Thomas Roger Van Devender. 9 and 10 March 1980, with Scott Mills, Gary Paul Nabhan, Barry Spicer, Frank

W. Reichenbacher, and Sandra Hunt Stein. 5 and 6 March 1983, with Stephen Ferris Hale and Timothy

Louis Van Devender. 25-27 March 1983, with Owen K. Davis, Stephen Ferris Hale, Cynthia Lindquist,

and Brent E. Martin. 9 March 1986, with Rebecca Kay Van Devender.

Charles T. Vorhies. 16 April 1924, with Walter P. Taylor.

Grady Under Webster. 17 March 1980. The first set of his collections is at DAV.

ha Loren Wiggins. 21 March 1933, with Forrest Shreve and others (see entry for Forrest Shreve; Fig. 16).

Benjamin Theodore Wilder. 20 November 2008, with Richard Felger.

Loraine Yeatts. 21 March 1995. In March 1995 she collected in Cabeza Prieta National Wildlife Refuge and the

Tinajas Altas region. Her collections are at ARIZ, CAB, CS and KHD and the herbarium at Cabeza Prieta

NWR headquarters, Ajo).

THE FLORA

We present a listing of all the vascular plants known to us from the Tinajas Altas region, from the present-day

Plants as well as fossils recovered from packrat middens. The vegetation and flora are dynamic, changing even

n °w, and have changed dramatically during the past millennia, along with shifting climate and human pres-

ses. This is the first publication of a comprehensive temporal flora for any region and spans more than 43,000

r* rs of P laru s inadvertently collected and curated by humble packrats ( Neotoma spp.) and more recently by

botanists. We document a total of 255 species in 192 genera and 52 families, of which at least 119 species in 96

®**r* and 36 families are known from the fossil record and among these fossils at least 28 species in 17 genera

^ 6 families are no longer present (Tables 4 and 5). A summary of the flora and comparison with adjacent and

Felger and Van Devender (2012) provide an abridged checklist of the present-day flora in a comprehensive

W ° rk on the Tinajas Altas region by Broyles et al. (2012), and an updated listing of the modem flora is included

ln a regional checklist by Felger et al. (2012). We include some plants from immediately adjacent localities that

316 expected to occur in the core area.

The most diverse families, present-day and fossil taxa (Table 7), are the Asteraceae (49 species and 2 hy-

Poaceae (21 species); Boraginaceae (16 species); Cactaceae (14 species); Brassicaceae, Euphorbiaceae,

^ «ae, and Solanaceae (10 species each); Polygonaceae (9 species); and Nyctaginaceae (8 species). This flo-

tlc spectrum represents somewhat of a departure from the usual floristic makeup of the present-day Sonoran

^ Sen 11013 including that of the Gran Desierto of northwestern Sonora) in which the most diverse families.

m decreasing order, are the Asteraceae, Poaceae, Fabaceae, and Euphorbiaceae (e.g., Felger 2000; Van Devender

et al. 2010). The most diverse genera in the Tinajas Altas region are Cryptantha (7 species, including one fossil),

Ambrosia (6 species and 1 hybrid), Eriogonum (6 species). Euphorbia (5 species), and Boerhavia, Cylindropuntia,

and Lycium each with 4 species.

There are 12 non-native species in the flora, which is only 5.3% of the total modem flora. Across the So-

noran Desert region this extremely low percentage of non-natives is matched only in the Gran Desierto dune

fields of northwestern Sonora (Felger 2000) and some Gulf of California islands (Felger & Wilder 2012; Felger

et al. 2011; Van Devender et al. 2009; Wilder & Felger 2010; Wilder et al. 2007). Sahara mustard (Brassicd

toumefortii) and Arabian grass (Schismus arabicus) are the only invasive species that are likely impacting native

plants (e.g., Felger 2000; Felger et al. 2003; Felger et al. 2012). Red brome ( Bromus madritensis subsp. rubens)

and buffelgrass (Cenchrus ciliaris) are serious invasives elsewhere in the Sonoran Desert, but have not become

a problem in the Tinajas Altas region, although Cenchrus ciliaris poses a potential threat. Boerhavia coccineu

(native to North America but not to southwestern Arizona) and netleaf goosefoot ( Chenopodium murale) are

ocahzed at the Tinajas Altas visitor area, while the two Sonchus species are widely scattered but not common.

Lactuca s erriola is highly localized and Erodium cicutarium and Eragrostis cilianensis are well established but

scarcely seem invasive in the flora area. Mollugo cerviana, reported to be non-native in the New World, is an

innocuous small an nual

Species Accounts

The plants in this flora are grouped as ferns and lycophytes, gymnosperms, dicotyledons (eudicots), and mono-

cotyledons. Within these categories the plants are listed alphabetically by family, genus, and species. Plan'

ami les iollow the APG 111 format (Angiosperm Phytogeny Group; e.g., Stevens 2008). Selected, pertinent

felgeretal., Flora of Tinajas Altas, Arizona

de Altar, Dry Borders = the six contiguous protected areas of Organ Pipe, Cabeza Prieta, Sonoran Desert National Monument, Barry M. Goldwater Range, Pinacate

Reserve, and the Reserva de la Biosfera Alto Golfo de California y Delta del Rio Colorado (Felger & Broyles 2007).

7.The ten most diverse plant families of the Tinaias Altas reoion.

synonyms are in square brackets following the accepted scientific name. Vernacular (common) names follow

| e Sc >entific names and when known are given in English, Spanish, and O’odham, respectively. The Spanish-

“Wguage names are italicized. The O’odham names are from Felger et al. (2007b). Plants not native to the flora

are marked with an asterisk (*). Fossil taxa and/or specimens are indicated with a dagger symbol (t).

nobotanical uses are from Felger (2007). Years before present (ybp) are the approximate years before 1950,

! * on radiocarbon dating— therefore some summarized dates in the species accounts are about one half

than thC y b P' radiocarbon date - Fl° wer color refers to the dominant color and not necessarily to

bS? fl ° Wer Part ° r structure - 11 be assumed that the species are common unless otherwise stated. The

| e descriptions and identification keys for the various taxa are based on plants from the flora area and are not

cessanly applicable to other regions. More extensive descriptions can be found in works by Felger (2000), Fel-

J/* 1 aL (2007b ). Wiggins (1964), the Vascular Plants of Arizona Project (Vascular Plants of Arizona Editorial

j ^«»ee 1992), and the Jepson Flora Project (2012). Nomenclature follows the regional checklist by Felger et

Ads floristic account is a specimen-based except for some records verified by an observation, photo docu-

taUon , 0r Published report. Extensive collections from the Tinajas Altas region are at ARIZ, ASU, and DES.

There are often multiple collections of a given species or taxon and in si

for each species or taxon. Extensive references to herbarium specimen

mental Information Network (2011; also known as SEINet) and other herbarium databases.

Specimens cited are at the University of Arizona Herbarium (ARIZ), unless otherwise indicated by the

standardized abbreviations for herbaria (Index Herbariorum 2009), or the herbarium at Cabeza Prieta Na-

tional Wildlife Refuge headquarters in Ajo, Arizona, identified as CAB. All herbarium specimens cited have

been seen by Felger, although some have been seen as images, unless otherwise noted. The name of the collec-

tor and collection number is given. In cases where more than one collector is listed on a label, generally only the

first collector’s name is stated. Locality descriptions and other information on herbarium labels generally are

abridged. If no collection number is provided on the herbarium label, then the specimen is identified by the

date of collection, for example, Harbison 6 Mar 1937. When the date of collection is significant, such as collec-

tions of historic interest, especially non-native species, both the collection number and date are given. It should

be noted that the labels on many of the early herbarium specimens merely state “Tinajas Altas Mountains” or

“Tinajas Altas.” Before the mid-twentieth century it was customary to regard the Gila and Tinajas Altas Moun-

tains as a single range and thus some of these specimens might be from the Gila Mountains, which currently

are regarded as the range to the north of the Tinajas Altas.

Although we provide extensive documentation of plants from this region, intrepid botanists should be

able to discover additional species.

FERNS AND LYCOPHYTES

Astrolepis cochisensis (Goodd.) D.M. Benham & Windham subsp. cochisensis [Notholaena cochisensis

Goodding] Scaly cloak-fern, jimmy fern

Above Tinajas Altas, 1500 ft. Van Devender 10 Mar 1980.

Cheilanthes parryi (D.C. Eaton) Domin [Notholaena parryi D.C. Eaton] Parky’s up-fern

Small desert ferns with short rhizomes and gray-hairy leaves.

Sheltered among rocks throughout the mountains. This fem extends into drier habitats than any other Sonoran Desert fern-

Tinajas Altas, Wiggins 21 Mar 1933. Tinajas Altas, above the tinaja containing Typha, shady crevices, in vertical granite

faults, 18 Apr 1976, Engard 920 (ASU). Frontera Canyon, 18 Mar 1998, Felger (observation).

Notholaena californica D.C. Eaton subsp. califomica [Cheilanthes deserti Mickel] California cloak-fern

Small tufted desert ferns with bead-like leaf divisions.

Felgeretal., Flora of Tinajas Altas, Arizona

; with scale leaves; the plants curl up when dry, and quickly expand with rain to reveal green foliage.

ingly scarce in the vicinity of the Tinajas Altas waterholes, which is likely a result of grazing by desert bighorn. Tim

Reeves, an expert on desert ferns, made the following entry in his field notebook for 12 February 1977, at Tinajas

Altas: “All ferns [the three species] green and healthy. Suggests fairly recent rain. Area searched for Selaginella ere-

mophila, none seen, various good habitat [for it].”

SE side of Tinajas Altas Mts, ca. 2 mi SE of Tinajas Altas, dense, locally extensive mats, not seen elsewhere, Felger 92-619.

0.8 mi SE of Tinajas Altas camping area, steep N-facing slope, a patch, 10 x 10 m, Reichenbacher & Hale 28 Mar 1981 .

Tinajas Altas Canyon, among rocks, ca. 0.7 km SW of upper tinaja, 1800 ft, shaded niche at canyon bottom, localized

(not seen elsewhere in vicinity), Felger 98-134.

GYMNOSPERMS-CONE-BEARING PLANTS

CUPRESSACEAE — REDWOOD OR CYPRESS FAMILY

tjuniperus califomica Carriere. Califc

junct from the primarily distribution along the Pacific side of California and mountains in northern Baja California.

fButler Mts, twigs, seeds, 10,360-11,250 ybp (3 samples). tTinajas Altas, twigs, seeds, 8970-18,700 ybp (14 samples), &

>43,000 ybp.

EPHEDRACEAE JOINT-FIR FAMILY

Ephedra aspera Engelm. ex S. Watson [E. nevadensis S. Watson var. aspera (Engelm. ex S. Watson) L.D. Ben-

son] Boundary ephedra; cautillo; ku’ukpalk

Woody shrubs; twigs green with paired, short, brown scale-leaves.

sils were originally identified as E. nevadensis, but are more likely to be E. aspera. We are unable to distinguish one

from the other on the basis of fossil specimens, and they are indeed closely related and often impossible to distin-

guish. It was an important medicinal plant (Felger 2007).

^ % mi S of Tinajas Altas, Hodgson 2100 (DES). Tinajas Altas Canyon, Felger 08-176. Borrego Canyon, 16 Jun 1992,

Mger (observation). fButler Mts, twigs, bracts, seeds, 740-11,250 ybp (7 samples). tTinajas Altas, twigs, seeds,

1230-18,700 (19 samples), & >43,000 ybp.

PINACEAE PINE FAMILY

tPi ® as mon °phylla Torr. & Frem. Single-leaf finyon; pinOn

Small trees. Unlike most pinyon taxa, the fascicles bear only a single leaf. Cones relatively small with wingless, edible

4 lee Age woodlands in sub-Mogollon Arizona. Tinajas

the lowest-elevation regional record for a pine in the Pleistocene. This pinyon presently occurs in northwest

ti Baja California, California, Idaho, Nevada, New Mexico, and Utah.

) with P. edulis Engelm. var./ollox Little, which today oi

le Mogollon Rim in central Arizona and in southern New Mexico. “Pinus ed

r — immures of P. edulis and P. monophylla. More study is needed” (Krai 1993: 383).

“ajas Altas, 460 m, 11,040-15,680 ybp, & >43,000 ybp.

190

Journal of the Botanical Research Institute of Texas 6(1)

ANGIOSPERMS— FLOWERING PLANTS

DICOTYLEDONS (EUDICOTS)

ir disk florets often si

3. Inflorescence and flo

and often with greer

Plants Leafless or Essentially So, or with Scale Leaves

(note: excluded are most seasonally leafless plants and annuals that have shed their leaves)

3. item noaes witn semi-persistent and opposite s

3. Leaves few, quickly deciduous, usually not scale

key).

h lens-shaped glands, the plants noticeably aromatic (citrus-li

Trees and Shrubs with Compound Leaves

stipules none; leafstalks (petiole and rachis) without glands; fruits globose or nearly so, 1-seeded Burseraceae

2 ' Plants armed or unarmed, not aromatic; trunks and limbs not unusually thick or semi-succulent; stipules generally

present (except Parkinsonia microphylla ); leafstalks sometimes with a prominent gland; fruits of pods, multiple-

seeded ( Olneya pods sometimes 1 -seeded) Fabaceae ( Olneya , Parkinsonia, Prosopis, Psorothamnus, Senegalia)

Trees and Shrubs with Simple Opposite Leaves (sometimes with some alternate leaves, or whorled)

192

Non-woody Succulents (stems and/or leaves succulent)

>r red-pink, collapsing with daytime heat; fruits of capsules opening

4. Stipules usually present; flowers bilateral, some or all petals separate; stamens 1 0; styles 1, the stigma unbranched;

fruits of pods, not globose ; Fabaceae {Acmipon, Doiea, LuptoRl

194

9. Stems yellow-green, dotted with lens-like glands, the plants aromatic, often citrus-like; flowers to 1 cm long,

Coyote Water, Felger 04-24.

>r short petioled, the blade narrowly lanceolate. Infloresc

Common and widespread at low elevations, especially washes, bajadas, and canyon bottoms.

Coyote Water, Felger 04-25. Tinajas Altas, 26 Oct 2003, Felger (observation). tButler Mts, calyx, 8160 ybp.

Unidentified amaranths, likely A. fimbriatus and/or A. pabneri, are recorded at Tinajas Altas dating to about 1

ago (see Doubtful, enigmatic and potential records).

tTinajas Altas, seeds, 9900 & 10,600 ybp.

Atriplex polycarpa (Torr.) S. Watson. Desert saltbush; chamizo cenizo; ’onk ’i:vakI

Woody shrubs. Fruiting bracts bearing 7 to many small, finger-like blunt teeth. Flowering in various seasons, espe-

cially with hot weather following rains.

One of the more common and widespread shrubs in the region. Washes, sandy valley plains, bajadas, pediments, and

lower slopes. It has been in the region for more than 10,400 years ago.

Camino del Diablo, E of Raven Butte, Felger 02-04. SW side of Tinajas Altas Mts, flats and granitic hills, 10 Jan 2002,

Felger (observation). tButler Mts, leaves, 10,360 ybp.

Chenopodium murale L. Netleaf goosefoot; chual, choal ; ’onk’iivakI, kaupdam

This species has been collected twice in the flora area, 18 years apart, between the lowermost Tinaja and the parking

area — the sites most often frequented by visitors. Our observations indicate that it is rather scarce at Tinajas Altas

but apparently well established. It is common in nearby northwestern Sonora at waterholes in the Sierra El Pinacate

and as a weed in agricultural fields (Felger 2000). The young herbage and especially the seeds are edible. This spe-

cies, widespread in the New World, is reported as native to the Old World.

Tinajas Altas, disturbed bank, Van Devender 10 Mar 1980. Parking area E of Tinajas Altas [Mesa del Muerto], Felger 98-

(Sanchez-del Pino & Flores Olvei

iassoniana Sanchez-del Pino & Floi

1 Pino] Woolly honeysweet; hierba cm.

and spreading, the herbage often scurfy-whitish; flowers mil

s Olvera [T. eliasso

Coyote Water, Felger 04-64. Tinajas Altas, 19 Mar 1998, Felger (observation; dry plants from previous year).

ANACARDIACEAE— CASHEW FAMILY

NN»ns aromatica Alton cf. var. trilobata (Nutt.) S. Watson [R. trilobata Nutt.] Skunk bush; umtta

Shrubs with winter-deciduous leaves. Ice Age Rhus seeds seem to be from a skunk bush, likely this one. The nearest

present-day population occurs on the north side of Pinacate Peak (Felger 2000).

*■ ar0matica cf - trilobata, Tinajas Altas, seeds, 9230-11,040 ybp (5 samples).

WlUS kearne y» F A. Barkley subsp. keameyi {Schmaltzia keameyi (F.A. Barkley) F.A. Barkley] Desert sumac; l/mita

^ldesierto. Fig. 17.

Hardwood shrubs, 2-4 m tall, the leaves tough, green, and generally evergreen, or sometimes partially or fuUy leafless

during extreme fore-summer drought. Flowers white or pink; fruits red, about 1 cm long, November-March. The

surfaces of freshly ripe fruits are wet with gooey but not sticky exudate; this liquid has a pleasantly tart (acidic) taste.

A ^freshing, tart drink is made from the fruits. The growth form and habit of this shrub is unique in the region. The

t ) r P e collection is from Tinajas Altas.

Altas Mountains, mosdy in steep north-facing canyons and cliff bases, from near the lower tinajas to near peak

elevations.

■Hfc subspecies occurs in the Gila and Tinajas Altas Mountains and in the nearby north-facing, steep granitic moun-

tains in northwestern Sonora west of the Pinacate region (Felger 2000). Disjunct populations of this subspecies oc-

t^r in the Sierra del Viejo southwest of Caborca, Sonora, and Sierra San Pedro Martir in Baja California. Two other

Journal of the Botanical Research Institute of Texas 6(1)

Fig. 1 7. Desert sumac ( Rhus kearneyi). North-facing slope at Tinajas Altas. 20 November 2008. Photo by Benjamin T. Wilder.

subspecies occur in mountains of Baja California and Baja California Sur. Rhus kearneyi appears to be a relict of a

more widespread Pleistocene distribution. This species seems most closely related to R. integrifolia (Nutt.) Benth. &

Hook. f. ex Rothr. of the Pacific Coast of the Califomias; the two species differ in part by leaf shape and pubescence.

Tinajas Altas Mts, [probably at Tinajas Altas], 29 Mar 1930, Harrison & Kearney 6573 (isotype). Tinajas Altas, Goldman

2311 (US). NE Tinajas Altas Mts, ca. 500 m SSW of Raven Butte, granitic mountainside, steep NE-facing ravine,

shrub to 4 m tall, 488 m, Morrison 590 (ASU). Borrego Canyon, vicinity of Borrego Tank, Felger 92-612.

APIACEAE— CARROT OR PARSLEY FAMILY

Recorded for Tinajas Altas more than nine millennia ago. The nearest present-day records are from the nearby Pints

Sands and Las Playas in Cabeza Prieta Refuge.

tTinajas Altas, fruits, 8970 & 9790 ybp.

APOCYNACEAE— DOGBANE FAMILY (includes Asclepiadaceae)

Asclepias albicans S. Watson. White-stem milkweedji/mate, candbulla . ,

Shrubs to 3 m tall, the stems reed- or wand-like, generally few, white-waxy, semi-succulent, and often leafless or with

Fairly common on arid, exposed rock slopes, and occasional on the bajada-plain of the Lechuguilla Valley. This bigM?

xerophytic milkweed has been in the region for at least five millennia. The fruits are edible after being roasted in coals-

Tinajas Altas, Shreve 5941. Lechuguilla Valley, Felger 08-216. tTinajas Altas, 1230-5080 ybp.

Asdepias erosa Torr. Giant sand-milkweed, desert milkweed; hierba del cuervo

Robust perennials, often 1-1.8 m tall, leaves large and rather thick. Flowers greenish '

ering in late spring and early summer in amazing heat when nothing else seems i

Scattered on sandy soils of Lechuguilla Valley.

Lechuguilla Desert, Simmons 21 May 1965.

Asclepias subulata Decne. Desert reed-stem milkweed; jumete, mata candeulla

Flowers waxy, cream- and yellow-white,

ccasional in the Lechuguilla Valley, recorded near

amino del Diablo, NE of Raven Butte, Felger 04-3.

tall. Lea.

:, few and quickly deciduc

Funastrum hartwegii (Vail) Schltr. [F. cynanchoides (Decne.) Schltr. var. hartwegii (Vail) Krings. Sarcostemma cynan-

choides Decne. subsp. hartwegii (Vail) R.W. Holm] Climbing milkweed; cuirote

Mosdy along washes and canyon bottoms. The fresh flowers can be eaten as snacks, and chewing gum was made from

the milky sap.

Coyote Water, Felger 04-46. Camino del Diablo at Coyote Wash, locally common in mesquites, Felger 10-163. Tinajas

Altas, Van Devender 9-10 Mar 1980.

ASTERACEAE— ASTER OR DAISY FAMILY

This is the largest family in the flora, the Sonoran Desert, and globally. The 43 present-day species represent

19% of the flora — comparable to the 15% of adjacent northwestern Sonora (Felger 2000). Additionally, 8 spe-

cies are known from the flora area only by fossils. Approximately 22 species, or 58% of the composite species

» the flora are annuals or ephemerals, comparable to 52% of adjacent northwestern Sonora; in contrast, world-

wide about 25% are annuals. Most of the res t of the local composites are herbaceous perennials or small shrubs,

andBaccharis sarothroides and Peucephyllum schottii are woody shrubs reaching more than 1.5 m in height. Two

small composite shrubs form important and widespread components of the local desert vegetation: Ambrosia

<kmosa and Encelia farinosa. Lactuca serriola and the two Sonchus species are the only non-native composites

» the flora area.

Most composites in the flora are insect-pollinated. More than 16 species have yellows flowers, e.g., Baileya

pbniradiata, Encelia farinosa, Gymnosperma glutinosum, and Pectis papposa. Stephanomeria pauciflora has noc-

tornal or crepuscular white or pinkish flowers; whitish flowers are found on about 10 species. A few species

we lavender rays but yellow disk flowers, e.g., Erigeron lobatus and M onoptilon bellioides. There are no red-

wered composites in the region. The Ambrosiinae, e.g., Ambrosia, are wind-pollinated, with corollas absent

female flowers and reduced on male flowers. Others, such as the filaginoids Logfia and Stylocline, have

mi " Ute ’ educed flowers and are undoubtedly selfing.

■ «ants often with milky sap; all florets conspicuous, bisexual, similar in shape (inner florets often smaller) with strap-like

w ray-like (ligulate — the ligules 5-lobed) corollas.

1 7 ?*“ ***** thread-like (capillary i. n

3. Florets 3 <

5 medium t0 lar 9 e - the larger phyllaries (13-)17-22 mm long; ligules of larger (

5 Heads" 9 ’ a jj henes taperin 9 int0 a slender beak; pappus bristles (6-)9.5-K — 1

); pappus bristles 2.2-8 nr

198

Journal of the Botanical Research Institute of Texas 6(1)

Journal of the Botanical Research Institute of Texas 6(1)

medicinal plant (Felger 2007).

Tinajas Alias, Van Devender 9-10 Mar 1980. Coyote Wash, 10 Jan 2002, Felger (observation).

tAmbrosia confertiflora DC. Sumleaf bur-sage; estafiate; mo’ostauc

Herbaceous perennials, winter and spring dormant.

Not known from the Tinajas Altas area today, but it was in the Butler Mountains more than 10,000 years ago. The nearest

present-day records are from nearby areas in Cabeza Prieta Refuge in poorly drained, clayish soils of the large playas.

Ambrosia deltoidea (Torr.) W.W. Payn,

Bushy subshrubs; tardily drought-deciduc

small burs.

■; CHAMIZO FORRA/EXO;

:r dormant. Flowers green or )

Valley plai

e it is one of the most widespread and abundant perenni:

in the more xeric Tinajas Altas region it is replaced by A. dumosa. A single Tinajas Altas c

esting anomaly, although tellingly, it is noted as being “rare.” In nearby regions, such as nc

toidea does not extend into such extremely xeric areas as does A. dumosa (Felger 2000).

ins, Tinajas Altas, 1200 ft, rare on granite, 0.5 m shrub. Van Devender 26 Mar 1983.

. Gray) W.W. Payne [Franseria dumosa A. Gray] White bur-sage; chamizo; tadsad

bage often whitish. Summer dormant, flowering and fruiting fall to spring; seeds in smal

our shoes and lower pant legs were coated yellow with white bursage pollen.

, and rocky slopes to higher elevations, and extending into the harshe

across much of the Sonoran and

5 , Reeves R5398(ASU).tTinajas

Mojave deserts. It has been in the flora area for more than 15,700 yeai

SE of Raven Butte, bajada, Felger 04-7. Tinajas Altas, Felger 10-177. Tina

Altas, leaves, burs, 1230-15,680 ybp (11 samples).

Ambrosia dumosa x A. ilicifolia

of the presumed hybrid was seen in the flora area.

Tinajas Altas, steep slope N of the tinajas, two branches collected from the one plant seen, Felger 98-136.

Ambrosia ilicifolia (A. Gray) WW. Payne [Franseria ilicifolia A. Gray] Holly-leaf bur-sage

Broad, spreading shrubs, often 1-2+ m wide. Leaves firm and holly-like, the dead leaves persistent. Summer dormant;

flowering and fruiting winter and spring. Seeds in burs.

Washes, canyon bottoms, and sometimes on rocky slopes. The dry, dead leaves rustling in the wind may sound star-

tlingly like a rattlesnake. Its presence at Tinajas Altas dates from more than 15,700 years ago.

Tinajas Altas, Vorhies 16 Apr 1924. 1.7 km WNW of Tinajas Altas Peak, 340 m, Baker 13308 (ASU). Frontera Canyon, 18

Mar 1998, Felger (observation). tTinajas Altas, leaves, burs, 1230-15,680 ybp (17 samples).

Ambrosia salsola (Torr. & A. Gray) Strother & B.G. Baldwin var. pentalepis (Rydb.) Strother & B.G. Baldwin

[Hymenoclea salsola Torr. & A. Gray var. pentalepis (Rydb.) L.D. Benson] White burrobush, cheesebush; ’lwadHOD

Shrubs, usually about as wide as tall, with many spreading and interlacing branches. Flowers straw-colored, the femak

flowers and seeds in small, winged burs. Flowering March and April; the wet plants smell like a dead animal.

Widespread across much of the Lechuguilla Valley and canyons in the mountains, and present in the region at least

9000 years ago.

Camino del Diablo at Coyote Wash, Felger 10-166. Coyote Water, 18 Mar 1998, Felger (observation). Tinajas Altas: 1200

ft, Lindquist 26 Mar 1983; Webster 24245. tButler Mts, bur, 8570 ybp. tTinajas Altas, burs, 8970 ybp.

'Artemisia ludoviciana Nutt, subsp. albula (Wooton) D.D. Keck. Western mugweed, white sage, silver worm-

wood; ESTAFIATE

Herbaceous perennials, flowering in late spring and probably with summer rains.

Western mugweed was at Tinajas Altas from at least 11,000 to more than 18,700 years ago. The nearest present-day*"

cords are from mountains in Organ Pipe Monument and higher elevations in the Sierra Pinacate.

tTinajas Altas, leaf fragments, 11,040-18,700 ybp (4 samples).

Felgeretal., Flora of Tinajas Altas, Arizona

as; mostly late spring to early fall.

Mostly along canyon bottom and at higher elevations in the Tinajas Altas Mountains; localized and not common.

Tinajas Altas, Goodding 1186. Tinajas Altas Canyon, Felger 04-79. Frontera Canyon, canyon bottom among rocks, Felger

Baccharis sarothroides A. Gray. Desert broom; escobaamakga, romemllo; susk kuagig

Broom-like shrubs; branches green, mostly leafless or sparsely leaved. Flowers whitish, the fruits become airborne with

copious white pappus; flowering and fruiting mostly October and November.

Localized along Coyote Wash; not common.

Coyote Water, Felger 04-26, 04-27.

Bahiopsis parishii (Greene) E.E. Schill. & Panero [Viguierapariskii Greene. V. deltoideaA. Gray var. parishii (Greene)

Vasey & Rose] Parish’s goldeneye; ariosa

Small shrubs; stems slender and brittle, the leaves rough-surfaced (scabrous). Flower heads with bright yellow ray and

disk florets.

Canyons and slopes to the summit.

Tinajas Altas, 990 m, Felger 04-84. Borrego Canyon, 3 Feb 1990, Felger (observation).

Baile y a Pleniradiata Harv. & A. Gray. Woolly desert-marigold

Non-seasonal annuals, but mostly in spring. Flowers yellow and showy.

Wside of Tinajas Altas Mts, Felger 05-48. FrontenJcanyon, Me

Mts, Van Devender 27 Mar 1983.

^bbiajuncea (Benth.) Greene var. aspera Greene. Sweetbush, chuckwalla deugh

Globose, bushy perennials or shrubs with slender stems and rough (scabrous) herbage; 1

drought deciduous. Flowers heads yellow and fragrant, with disk florets only, attra

“any other insects; flowering response non-seasonal. Karen Reichhardt (persons

has observed chuckwallas (Sauromalus ater) at Tinajas Altas eating chuckwalla delight.

mmon an d widespread; bajadas, washes (Fig. 15), canyons, and rocky slopes. Present in the region

8 Mar 1998, Felger (observ

jno del Diablo, SE of Raven Butte, Felger 04-6. Tinajas Altas Pass, Reeves R-5411 (AS

ybp. tTinajas Altas, achenes, 4010-10,750 ybp (8 samples).

atractyloides A. Gray var. atractyloides. Spiny-leaf brickellbush

s rubs or small bushy perennials. Leaves firm with spinescent marginal teeth. Floi

rse, the leaves quickly

mication 21 February

at least 10,800

achenes, 8160

s m 7r C0UIteriA G rayv

S ^ l *™bs.I.» asa(T J j>l

*>PartofCabeza Prieta Refuge and the Sierra pin .

t JaSAUas , involucres, 4490 & 10,950 ybp.

< ^* Sens P arr yi A. Gray. Yellow tack-stem

annuals. Flowers pale yellow.

above 550 m.

tTinajas Alias, achenes, 9230 & 9900 ybp.

Calycoseris wrightii A. Gray. White tack-stem

Cool-season annuals with milky sap, branched at or near the base, ;

above, otherwise glabrous or glabrate. Rays of flower heads w

v Surveyors Tank, 29 Mar 2010, Fi

CHAENACHS

Spring annuals; flower heads of disk florets, white and often suffused with pink.

Chaenactis carphodinia A. Gray var. carphodinia. Pebble pincushion

Foliage not cobwebby, the phyllary tips needle-like. Flowers white.

Sandy flats, washes, bajadas, and rocky slopes.

TinajasAltas, Van Devender 26 Mar 1983.

Chaenactis stevioides Hook. & Am. Desert pincushion

Plants generally slightly more robust than those of C. carphodinia ; stem base often with at least some cobwebby hairs;

phyllary tips relatively blunt. Flowers white or pinkish white.

Sandy soils of valley bottoms and rocky slopes. It has been in the region for at least 10,000 years.

Coyote Water, 18 Mar 1998, Felger (observation). W side of Tinajas Altas Pass, Lindquist 26 Mar 1983. tTinajas Alias,

ENCELIA

Small shrubs, the leaves alternate, and with yellow flowers, and sometimes with brownish disk florets.

Encelia farinosa A. Gray ex Torr. var. farinosa. Britelebush; inoenso, herba del bazo rama blw ca; tohaves. Fig?- 8

and 9.

Shrubs, not long-lived. Leaves highly variable with soil moisture. Flowering branches usually raised well above the foli-

age; massive displays of daisy-like flowers with bright yellow rays and yellow or brownish disk florets, mostly in

spring but also with summer-fall rains. At maturity the flower heads turn downward, dumping out the seeds.

One of the most common desert perennials in the region; widespread including all slope exposures to summits, dry

washes, canyons, and bajadas; generally not in open creosotebush flats. The yellowish resin was used as chewing

gum and an all-purpose adhesive and sealant. Brittlebush has been in the region for at least 43,000 years.

Tinajas Altas, 5 Dec 1935, Goodding 2204. El Camino del Diablo, E of Raven Butte, Felger 01-586. tTinajas Altas, leaf

fragments, achenes, 1230-18,700 (19 samples), & >4300 ybp.

Encelia farinosa var. farinosa x E. frutescens

bright yellow rays and disk florets

Scattered, isolated plants in the Lechuguilla Valley.

Camino del Diablo, E of Raven Butte, Felger 01-585.

Encelia frutescens A. Gray. Button encelia

Short-lived shrubs, densely branched and often mound-shaped to about 1 m tall. Flowers heads yellow, of disk flot**

Felger etal., Flora ofTinajas Altas, Arizona

203

Locally common on sandy soils of Lechuguilla Valley.

Coyote Water, Felger 04-65. Camino del Diablo at W boundary of Cabeza Prieta, Felger 10-159.

tEricameria cuneata (A. Gray) McClatchie var. spathulata (A. Gray) H.M. Hall [ Haplopappus cuneatus A. Gray

unmanked spathulata (A. Gray) S.F. Blake] Wedge-leaf goldenbush

Perennial herbs or subshrubs.

The nearest present-day population is in the Ajo Mountains, mostly at higher elevations. It was widespread in the region

during the Ice Age.

tTinajas Altas, leaves, 8970-15,680 (4 samples), & >43,000 ybp.

tEricameria laricifolia (A. Gray) Shinners [ Haplopappus laridfoliusA. Gray] Turpentine bush

Shrubs with bright green, resinous foliage.

The nearest present-day population is in the Ajo Mountains, mostly at higher elevations. It seems to have been very com-

mon and ranged across the region during the late Wisconsin Ice Age.

tButler Mts, twigs, leaves, involucres, 10,360 ybp (common in this sample). tTinajas Altas, leaves, 8970-18,700 ybp (10

samples).

tEricameria teretifolia (Durand & Hilg.) Jeps. [Chrysothamnus teretifolius (Durand & Hilg.) H.M. Hall] Green rab-

tTinajas Altas, leaves, invoh

ijas Altas from 15 to at least 43 millennia ago. The nearest present-day populations are in

tlso in the Mojave Desert in southern California at the western edge of the desert into south-

estem Utah at the upper limits of desert and extending into pinyon-juniper vegetation,

ucres, 15,050-18,700 & >43,000 ybp (4 samples).

Erigeron lobatus A. Nelson. Deseri

r. Disk florets yellow, tl

I r slender and pale lavender.

Geraea canescens Torr. & A. Gray. Desert sunflc

Coarse spring annuals; showy suni

Surprisingly, it has not been found

gravelly and rocky soils. It has been in the

Buder Mts, Van Devender 27 Mar 1983. TButlei

DESERT GOLD

ike heads with yellow-orange disk florets and large, bright yellow rays

imediate Tinajas Altas area, although it is often abundant nearby on sa

the region for at least 9 millennia.

i, 3820 & 8570 ybp.

1 " nos P enna glutinosum (Spreng.) L

^mall shrubs or subshrub perennials wit!

ray florets; flowering during the warmer months.

The plants are amazingly drought resistant and often retain bright green foliage and may ever

during extended drought when nearly all of the surrounding plants are dormant and leafless

area for more than 15,700 years.

Tinajas Altas, McLaughlin 1969. Surveyors Canyon, canyon bottom, Felger 10-208. tTinajas Alt;

1230-15,680 ybp (10 samples).

t has been in the flora

is H. sessiliflora (Nutt.) Shinners >

em Sonora (Felger 2000). The Tir

010 to >43,000 ybp (13 samples).

Hymen <Kka salsola, see Ambrosia salsola

ibout 4000 years ago. This is the only record for this genus

is presently occur in the region. The nearest population c

var. thiniicola (Rzed. & C. Ezcurra) G.L. Nesom, a rare dr

tTinajas Ai

204

*Lactuca serriola L. Prickly lettuce, compass pi

Annuals, often germinating in late winter or sprii

turned basally to hold the leaf edgewise and u

ering late spring and summer; flowers pale yellow.

Locally at the south end of Coyote Wash in the Lechuguilla Valley. Usually a weedy species, native

Camino del Diablo at Coyote Wash, locally abundant in mesquite “forest,” some dry dead stalks fr

m in height, Felger 10-162.

> the Old World

a last year reach 2+

Laennecia coulteri (A. Gray) G.L. Nesom [ConyzacoulteriA. Gray] Coulter’s horseweed

Warm-weather annuals; flowers inconspicuous. Infrequent along Coyote Wash.

Coyote Wash at Camino del Diablo, Felger 02-14.

LOGFIA. Fluffweed

Diminutive or small spring annuals, often white-woolly. Flowers minute. Logfia is a genus segregated from Filagp.

logfia arizonica (A. Gray) Holub [Filago arizonica A. Gray] Arizona fluffweed

Small cool-season annuals, often white-woolly; branching pattern usually symmetric (2 equal side-bran

at each

Sandy, gravelly or clayish-silt soils; washes, canyon bottoms, floodplains, andbajadas and perhaps soil pockets on rocky

slopes. Often where rainwater may briefly accumulate. Commonly growing intermixed with L.filaginoides.

Camino del Diablo at Coyote Wash, Felger 02-9. Coyote Water, Felger 98-114. Surveyors Canyon, canyon bottom, Felger

10-212.

Logfia filaginoides (Hook. & Am.) Morefteld (L. califomica (Nutt.) Holub. Filago califomica Nutt] California fluffweed

mostly sparsely papillate, with a pappus of 17-23 bristles falling away in complete or partial rings; lobes of disk co-

rollas mostly 4, usually red-tipped.

Washes and canyons, bajadas, and among rocks on slopes.

Tinajas Altas, Felger 98-144. Surveyors Canyon, canyon bottom, Felger 10-213. Canyon below Raven Butte Tank, Felger

10-234.

Machaeranthera pinnatifida, see Xanthisma spinulosus

Monoptilon bellioides (A. Gray) H.M. Hall. Mojave desert star

Low growing cool-season annuals. Flower heads showy, the rays white or lavend

disk yellow; pappus bristles of shorter, outer white bristly scales, and more ni

Seasonally common, sometimes producing showy displays on otherwise nearly

of Tinajas Altas Pass, Halse 31 Mar 1973. Tinaj

Felger 10-215. Tinajas Altas Canyon, 19 Mar 1998, Felger (obser

foxia arida B.L. Turner & M.I. Morris var. arida. Spanish n

Reeves 5399 (.

» linear. Flower heads of disk florets, dull white or pale pin

en gravelly-rocky flats; sandy »

ver slopes.

Surveyors Canyon, canyon bottom.

r 1983.

Felger et al., Flora of Tinajas Altas, Arizona

Pectis papposa Harv. & A. Gray var. papposa. Desert cmnchweed; manzanilla del am*

One of the most abundant summer-fall annual wildflowers in the region. Plants pungently ai

oil glands. Flowers showy and bright yellow with ray and disk florets.

Sandy to rocky soils, washes, plains, and less common on slopes.

Coyote Wash at Camino del Diablo, Felger 04-66. Coyote Water, Felger 04-56.

r. Desert rock daisy

M MANZANKYA

11,000 years ago.

Coyote Water, Felger 04-57. Canyon b

1998, Felger (observation). Canyc

9230 & 10,950 ybp.

Y-CEDAR; ROMERO DEL DESIERTO

mbling a small conifer, with twisted trunks and branches, and shreddii

t, of yellow disk florets.

Rocky and often exposed mountain slopes and in the region for more than 15,700 years.

S of Tinajas Altas, N-facing slope, 8 ft tall, 7-8 ft wide, Hodgson 2096 (DES). Tinajas Altas, Van Devender 5 M

Frontera Canyon, 18 Mar 1998, Felger (observation). tButler Mts, achenes, 740-11,060 ybp (5 samples). 1

Altas, leaves, involucres, achenes, 1230-15,680 ybp (6 samples).

Pleurocoronis pluriseta (A. Gray) R.M. King & H. Rob. [Hofnu

Small shrubs or subshrubs. Leaf blades very narrowly arrow-shap

often scarcely wider than the petiole.

Often growing from crevices on rock faces, canyons, cliffs, and slopes.

Tinajas Altas Mtn, on cliffs and rock slopes, 4 Mar 1927, Belden 3604. Tinajas Altas Tanks, on rocky hills above tanks,

Pleurocoronis sp./spp.

The younger specimens are probably P. pluriseta. The older ones may be P. laphamioides (Rose) R.M. King & H. Rob.,

which nowadays occurs in the Ajo Mountains and farther south in northwestern Mexico.

tButler Mts, involucres, achenes, 740-11,250 ybp (7 samples). tTinajas Altas, involucres, achenes, 4010-15,680 ybp (9

samples).

Porophyfluni gracile Benth. Slender poreleaf; odora, merba del vena do

Herbaceous perennials, pungently aromatic. Stems and leaves bluish green, the foliage sparse and quickly deciduous,

the leaves slender. Flower heads with pinkish-white disk florets. Growing and flowering at various seasons.

Widely scattered; rocky slopes, canyons, arroyos, and upper bajadas. It has been in the region since at least 8600 years ago.

Tinajas Altas, Van Devender 5 Mar 1983. Tinajas Altas Canyon, 19 Mar 1998, Felger (observation). tButler Mts, achenes,

involucres, 740-8570 ybp (4 samples). tTinajas Altas, involucres, achenes, 1230-8255 ybp (3 samples).

Pltn « 1 thella exigua (A. Gray) Rydb. Brightwhite

Diminutive spring annuals. Flower heads 5 mm long, with 3 or 4 ray florets, the rays white with violet tips.

Higher elevations in the mountains.

Tinajas Altas, above the tinajas, Felger 98-132.

A. Gray. Desert velvet, turtleback

lly, and strongly scented. Leaves velvety gray-green with deeply incised veins. Flower

urs and pappus bristles bright, iridescent copper-colored,

nd of the Tinajas Altas region and the desert plains west of the Tinajas Altas

range, observed on a steep granitic slope where it was uncommon and on a cobble-rock bajada.

Camino del Diablo, E edge of Davis Plains, Halse31 Mar 1973 (probably slightly north of the flora area). Along the “old”

Tinajas Altas Pass road (a bit north of the present Tinajas Altas Pass road), 7 72 843 E, 35 81 704 N, Zone 11, WGS

H 1050 ft, common, photo, Malusa 18 April 2011.

s Pring annuals,

heads of yellow disk florets. Ache

Apparently highly localized at the no

206

Cool-season annuals; plants glabrous (as opposed to Calycoseris wrightii), the flower heads showy, of ray-like florets,

white tinged with pale rose-purple and yellow. Plants often browsed and growing in the protection of small shrubs,

the flowering steins overtopping the nurse shrub such as Ambrosia dumosa or, especially in drier years, spiny shrubs.

Sandy to rocky soils; washes, plains, and slopes, lowlands to higher elevations. Present at Tinajas Altas more than

10,000 years ago.

Coyote Water, Felger 05-151. Tinajas Altas Pass, along wash, McLaughlin 1974. tTinajas Altas, achenes, 10,070 ybp.

Senecio mohaven

Winter-spring ai

A. Gray. Mojave groundsel

als, often small and delicate. Stems and lower leaves glabrous and often purple-g

*SONCHUS. Sow THISTLE

Winter-spring annuals, the flower heads pale yellow with ray-like (ligulate) florets.

*Sonchus asper (L.) Hill subsp. asper. Prickly sow thistle; chinita; ho’idkam, Y.vakI

Winter-spring annuals; larger plants conspicuously spiny-prickly and sometimes robust.

Localized and seldom common; documented at two localities near the Mexican border— in a wash and a canyon bot-

Monument (Felger 2000). Y Y Q

Coyote Wash at Camino del Diablo, Felger 02-10. Frontera Canyon, 18 Mar 1998, Felger (observation).

*Sonchus oleraceus L. Common sow thistle; chinita; hauvI, hehewo

Winter-spring annuals, mostly less than 80 cm tall.

Widely scattered but not common; washes and canyons, sometimes on slopes. This sow thisde is well established in the

a Canyon, 18 Mar 1998, Felger (observatit

Stephanomeria pauciflora (Torr.) A. Nelson. Desert-straw, desert wire-lettuce

Globose or mound-shaped bushy perennials with sparse foliage. Flowers pale pink, closing by mid-day or earlier in hot

weather; flowering non-seasonally.

Camino del Diablo at Coyote Wash, Felger 10-168. 1 mi E of Tinajas Altas, Van Devender 86-144.

Stylocline micropoides A. Gray. Desert neststraw

Diminutive, woolly, cool-season annuals; flowers minute and inconspicuous.

Locally common in the Tinajas Altas Mountains on upper bajadas, benches, and washes, and sometimes in soil pockets

on rocky slopes and in canyons.

Coyote Water, 18 Mar 1998, Felger (observation). Tinajas Altas Canyon, Felger 98-122. Vicinity of Tinajas Altas, Van

Devender 05 Mar 1983.

Trichoptilium incisum (A. Gray) A. Gray. Yellow-head

Annuals with woolly leaves; mostly growing and flowering during cooler seasons, especially in spring, and soi

flowering into early summer. Flower heads of yellow disk florets.

Widespread in the region, mostly on rocky or gravelly soils; washes, bajadas, and rocky slopes.

Tinajas Altas, Van Devender 10 Mar 1980. W end of Tinajas Altas Pass, Reeves R 5425 (ASU).

Trixis califomica Kellogg var. califonuca. California threefold

Small shrubs or subshrubs with brittle stems and thin, upright leaves. Flowers yellow and ray-like (bilabiate).

Felgeretal., Flora ofTinajas Altas, Arizona

207

Canyons, bajadas, and rocky slopes including extremely arid hills to higher elevations in the mountains. It has been in

the region for more than 11,000 years.

Tinajas Altas, Vorhies 16 Apr 1924. Tinajas Altas Pass, 4 mi W ofTinajas Altas, Webster 24257. Granitic hills at SW side of

Tinajas Altas range, 10 Jan 2002, Felger (observation). TButler Mts, leaf fragments, 740-8160 ybp (3 samples). tTina-

jas Altas, leaf fragments, 4010-10,950 ybp (7 samples).

Vig uieraparishii, see Bahiopsis parishii

Xanthisma spinulosum (Pursh) D.R. Morgan & R.L. Hartm. var. gooddingii (A. Nelson) D.R. Morgan & R.L.

Hartm. [Machaeranthera pinnatifida (Hook.) Shinners var. gooddingii (A. Nelson) B.L. Turner & R.L. Hartm. Haplo-

pappus spinulosus (Pursh) DC. subsp. gooddingii (A. Nelson) H.M. Hall] Spiny goldenweed

Short-lived herbaceous perennials and also flowering in the first season. Ray and disk florets bright yellow, mostly in

spring but also with summer rains.

Mostly in canyons and on slopes to the summit, and sometimes along washes and on the desert floor. This or a closely

related Xanthisma has been in the flora region for more than 11 millennia.

Borrego Canyon, Felger 92-617. 1 mi N ofTinajas Altas, Kurtz 1169. tX. cf. spinulosum: Butler Mts, involucres, achenes,

740-11,250 ybp (7 samples). tTinajas Altas, achenes, 4010 & 9900 ybp.

BORAGINACEAE — Borage Family (includes Hydrophyllaceae)

There are 17 species in this family recorded for the flora area and all are annuals or semi-herbaceous perennials

and all except Tiquilia grow only during the cooler seasons (see Felger 2000).

^SINCKIA — Fiddleneck

Winter-spring annuals, mosdy erect, the herbage with conspicuously coarse hairs. Flowers bright yellow-orange.

1 the nutlets rough with ragged-

edged ornamentations. Flowers yellow-orange,

idespread in the region, from lowest to high elevations. Documented for the region from more than 11

1,250 yea

208

amino del Diablo at Coyote Wash, Felger 10-169. Tinajas Altas, Van Devender

ybp. tTinajas Altas, nudet, 9230 ybp.

inckia tessellata A. Gray var. tessellata. Desert fiddleneck; cetkom

Sandy soils at least in the soi

in the region for at least 15,000 years.

Camino del Diablo at W boundary of Cabeza Pi

lets, 9230-15,050 ybp (3 samples).

iedia, but often more rob

>n of lobes), the nutlets w

he Lechuguilla Valley. G

Felger 10-16U

beza Prieta Refuge. It has been

D, 360 ybp. tTinajas Altas, mtt-

CRYPTANTHA

Winter-spring annuals and one species of long-lived annuals or short-lived perennials; growing and flowering

only during the cooler seasons, the plants beset with harsh, glassy hairs. Flowers small and white. Fruits with

1 or 4 nutlets enclosed in a calyx covered with harsh, glassy hairs, the whole structure resembling a miniature

bur. Nutlets all similar or dissimilar either in size and/or ornamentation and how readily they fall out of the

calyx. Cryptanthas are among the most diverse and widespread plants in the region.

1. Nutlet margins knife-edged or winged.

2. Calyx broad (lobes ovate to lance-ovate); nutlet wing usually as wide as the nutlet body and fringed with finger-like

2. Calyx n

/v (lobes lanceolate); nutlet wing n;

I . Nutlet margins rounded or angled but not knife-edged

s in the region. Flowers white. This is one of ti

Cryptantha angustifolia (Torr.) Greene. Narrow-

plants that gets in your socks and sleeping bag, re

Valley bottoms and plains, canyons, and rocky slopes.

Tinajas Altas, Van Devender 5 Mar 1983. Tinajas Altas Pass, Reeves R 5405 (ASU).

Cryptantha barbigera (A. Gray) Greene. Bearded cryptantha

Cool-season annuals; flowers white. The plants are someth

Widespread across the flora area; sandy soils of washes and plains, and on rocky slopes to higher elevations. It was inti*

region more than 5100 years ago

Tinajas Altas Mts, above the tinajas, Felger 98-137. tTinajas Altas, nutlets, 5080 ybp.

Cryptantha maritima (Greene) Greene [C. maritima var. pilosa I.M.Johnst.] White-haired cryptantha

Cool-season annuals; flowers minute. As with C. barbigera, the plants are sometimes deformed by mites and do not

produce flowers. The varieties do not seem worthy of recognition; both occur intermixed (Felger 2000).

Common and widespread; washes and sand flats, and bajadas to rocky slopes at higher elevations. It has been in tbe

region for more than 11,000 years.

Tinajas Altas, Kearney 10905. Canyon below Raven Butte Tank, Felger 10-220. Tinajas Altas Pass, Reeves R5401 (A Stt

annotated by Jeff Brasher, 2001, “mixed collection, var. maritima & var. pilosa”). tButler Mts, calyx, nutlets, ^

10,360 ybp (3 samples). tTinajas Altas, fruits, nutlets, 4010-11,040 ybp (8 samples).

Cryptantha micrantha (Torr.) I.M. Johnst. subsp. micrantha. Dwarf cryptantha

Cool-season annuals; plants diminutive with very slender stems, the roots and stems staining red-purple, the fl° werS

Felgeretal., Flora ofTinajas Altas, Arizona

209

minute and white. This is the smallest cryptantha in the region. It is common in immediately adjacent regions and

Cabeza Prieta Refuge: Pinta Sands, Felger 92-17.

Cryptantha pterocarya (Torr.) Greene var. cydoptera (Greene) J.F. Macbr. Wing-nut cryptantha

Cool-season annuals. Readily recognized by the broad sepals, green when fresh and only moderately hairy, relatively

large fruits, and relatively large nutlets with broad, ornamented wings.

and present in the region more than 11,000 years ago.

Tinajas Altas, Van Devender 10 Mar 1980. Canyon below Raven Butte Tank, Felger

ybp. tTinajas Altas, nutlets, 11,040 ybp.

Cryptantha racemosa (S. Watson ex A. Gray) Greene. Bushy cryptantha

Annuals, highly variable in size, or short-lived perennial herbs or subshrubs; growing and flowering only during c<

seasons; flowers white. This species and C. holoptera, known from nearby northwestern Sonora, are the

cryptanthas in the region that survive more than one season.

Rocky si.

sandy gravelly washes in the Tinajas Altas Mountains to higher elevati.

Vicinity of Tinajas Altas, 1700 to 1900 ft, on cliffs, Van Devender 5 Mar 1983. W end Tinajas Altas Pass, Reeves 85423

(ASU; originally identified as C. holoptera (A. Gray) Macbr., the fruits are not mature). tButlerMts, calyx, nutlets,

10,360 ybp. tTinajas Altas, calyces, nutlets, 8970-11,040 ybp (3 samples).

tCryptantha utahensis (A. Gray) Greene. Scented cryptantha

Annuals. This species was at Tinajas Altas 11,000 years ago and is presently in the Mojave Desert and pinyon-juniper

regions of Mohave County, Arizona, above about 950 m, and also in east-central California to southwestern Utah.

tTinajas Altas, calyx, nutlets, 11,040 ybp.

Eucrypta micrantha (Torr.) A. Heller. Desert hideseed

Delicate cool-season annuals, aromatic and viscid-glandular; leaves pinnately divided; flowers small, the corollas pale

violet or lavender with a pale yellow throat.

Often in protected niches beneath shrubs or sheltered among rocks. Widespread; washes, bajadas, canyons, and rocky

slopes.

Coyote Water, Felger 05-134. Tinajas Altas, Van Devender 5 Mar 1983. Canyon below Raven Butte Tank, Felger 10-219.

NAMA — Purple mat

1 Plants semi-prostrate, matted; longer s

ma dei >»issum A. Gray var. demissum. Purple mat

Cool-season annuals, mostly semi-prostrate, low and often matted. Flowers bright lavender-pink.

Mostly along washes and gravelly soils of bajadas and plains; often in open, otherwise barren areas such as desert pave-

ment. The plants are more compact, closer to the ground, generally smaller and with brighter and darker-colored

r . Hitchc. [N. coulter! A. Grayl Bristly r

Cool-season annuals, erect with ascending branches, or with age sometimes spreading-prostrate but noi maueu. r.uw-

ers P®le lavender to purple.

Widespread and common across the flora area, especially washes, sandy flats, bajadas, and canyons.

Coy °te Water, Felger 04-52. Tinajas Altas, Felger 10-191. Tinajas Altas Pass, Reeves 5413 (ASU). Surveyors Canyon, can-

yon bottom, Felger 10-214. Frontera Canyon, 18 Mar 1998, Felger (observation).

^EOCARYA — Comb-bur, comb-seed

s P rin g annuals. Leaves narrow, lower ones opposite, appearing as a basal rosette, the stem leaves alter-

210

nate. Inflorescences not helicoid or only moderately so. Flowers minute, at least soi

petals white. Nutlets 4, spreading open like tiny jaws, the margins variously toothed.

Pectocarya heterocarpa (I.M. Johnst.) I.M. Johnst. Mixed-nut comb-bur

Cool-season annuals; flowers minute, white with a yellow throat.

Widespread; sandy soils of washes and valley floor to the mountains. It was in the Butler Mi

; than 10,000

Coyote Water, Felger 05-137. Tinajas Alias, Felger 93-204. tButler Mts, nutlets, 10,360 ybp.

Pectocarya platycarpa (Munz & I.M. Johnst.) Munz & I.M. Johnst. Broad-wing comb-bur

Cool-season annuals. Similar to P. heterocarpa but the nutlets are noticeably larger with broader and usually yellowish

Widespread including washes, desert flats, and bajadas to rocky slopes. It was at Tinajas Altas more than 10,600 years

ago.

Coyote Water, Felger 05-137A. Tinajas Altas, Van Devender 5 Mar 1983. tTinajas Altas, nutlets, 10,600 ybp.

PHACEL1A— Scorpion-weed

The phacelias in the flora area are cool-season annuals, glandular pubescent and can cause unpleasant derma-

titis. Inflorescence branches are generally helicoid (curled at the tip like a scorpion tail) except in P. neglecta,

and the flowers lavender, lavender blue, or white.

igua (M.E. Jones) J.F. Macbr. [P. ambigua M.E. Jon

t scorpion-weed

Cool-season annuals, mostly erect, stinky, glandular pubescent, and can cause unpleasant dermatiti

branches curved (helicoid or ‘scorpioid’). Flowers lavender-purple. Seeds 4 per capsule.

This is the most abundant and widespread phacelia in the region; in many habitats including washes, a

plains, and rocky slopes to their summits.

Coyote Water, Felger 05-128. Tinajas Altas, Van Devender 5 Mar 1983.

Phacelia neglecta M.E. Jones. Alkali phacelia

Short, stubby cool-season annuals, often less than 8 cm tall, semi-succulent with the main stem and ro<

Flowers white. Seeds many per capsule. Unlike most phacelias, the plants are not stinky and the in

short and not coiled (helicoid).

General!)

pediments near mountain bases; not common in the flora area.

E margin of Davis Plain, W branch of Camino del Diablo, decomposed granite pediment directly downslope fromd#*?

hill, Felger 05-62.

Phacelia pedicellata A. Gray. Pedicellate scorpion-weed

Robust plants, conspicuously glandular-pubescent with an offensive odor, the herbage often pale green. Flowering

Canyons bottoms and among rocks, especially on north- and east-facing mountain slopes, from base to higher elevation

Tinajas Altas, canyon bottom E of tinajas, Felger 98-145. Tinajas Altas Canyon, Felger 98-127.

tPhaceliasp.

This is this only fossil record for a hydrophyll (Hydrophyllaceae, sensu stricto) in the flora area. The seeds are excavated

on one side like those of many other phacelias with four seeds per capsule.

tTinajas Alias, seeds, 1230 ybp.

Tiquilia palmeri (A. Gray) A.T. Richardson. Palmer’s crinklemat

Low-growing perennials forming small semi-prostrate mats, the areal portions drying back during drought and re-

sprouting from very slender roots issuing from deep, very thick, long, black roots. Stems with a forked branching

pattern (pseudo-dichotomous). Leaves with 2 or 3 (4) pairs of shallowly impressed veins; the blades 3-8 mm long.

Growing and flowering non-seasonally following spring and summer-fall rains; flowers essentially sessile and axil-

Butler Mts, Von Devender 27 Mar 1983.

BRASSICACEAE — Mustard Family

The nine species in the flora area are cool-season annuals except Lyrocarpa.

1. Fruits of 2 disk-shaped halves (spectacle-shaped) joined along less than 25% of their margins; flo

2. Leaves with broad, coarse teeth or lobes; petals (fresh) 10-’ 9P

1. Fruits not of disk-shaped halves; flowers various colors, fragra

long; fruits usually widest well above

ss, ca tournefortii Gouan. Sahara mustard; mostaza del Sahara

^-season annuals, extremely variable in size, less than 10 cm tall with k

with leaves to 80 cm long. When dry and dead the larger plants may bre E

Flowers pale yellow and apparently selfing (autogamous),

idespread and seasonally common in lowland disturbed and natural site;

soils; washes and valley plains, and sometimes extending onto rocky si

t the root and become tumbleweeds,

s the flora area, especially on sandy

e flora region. The broad, spreading basal-rosette leaves can

mnuals. Densely crowded or drought-stressed plants may be

soil moisture may grow to more than 1 meter across and nearly 1 m tall, making this species the largest herbaceous

On u** Plant in tHe regi ° n (see Felger 2000) -

tch 28, 2010, Richard Laugham and Felger poked around in the miniature mesquite thicket in l

pmmd Where thC Camino del Diabl ° crosses Coyote Wash and Felger made the following observatio

e Lechuguilla

212

Journal of the Botanical Research Institute of Texas 6(1)

ing with only a single fruit with 3-7 seeds, the larger leaves 1.0 or 1.4 to several cm long, and basal rosette leaves of-

ten only 1-3 or sometimes not developing, the stems extremely slender, unbranched and the plants 8.7- 11.4 cm and

more in height. Other, less crowded, robust plants have rosette leaves more than 30 cm long. Obviously B. toumefor-

Native to the Old World, B. tournefortii has spread explosively across the Sonoran Desert, especially in sandy-soilhabi-

It was first reported from southeastern California by Jepson (1923-1925) and established in agricultural areas in

southeastern California by 1938, recorded from Yuma in 1957, northwestern Sonora in 1966, and widespread across

the Sonoran Desert region since at least the 1970s (Dimmitt & Van Devender 2009; Felger 2000; Malusa et al. 2003;

Van Devender etal. 1997).

Coyote Water, Felger 04-33. Camino del Diablo at Coyote Wash, 28 Mar 2010, Felger 10-172. Surveyors Canyon, Lamb

Tank, Felger 10-197. Canyon below Raven Butte Tank, Felger 10-242. Camino del Diablo, E of Raven Butte, 29 Nov

2001, Felger (observation). Tinajas Altas Mts, above the tinajas, 19 Mar 1998, Felger (observation).

Caulanthus lasiophyllus (Hook. & Am.) Payson [ Guillenia lasiophylla (Hook. & Am.) Greene. Thelypodium lasio-

phyllus (Hook. & Am.) Greene) California mustard

ward as they mature.

Widespread; washes, valley plains, and mountains. Often in shaded niches beneath spinescent shrubs.

Coyote Water, Felger 05-131. Tinajas Altas Pass, Reeves 5404 (ASU). Tinajas Altas, Felger 10-181. Tinajas Altas Mts, above

the tinajas, 19 Mar 1998, Felger (observation).

Descurainia pinnata (Walter) Britton subsp. ochroleuca (Wooton) Detling. Western tansy-mustard; pamtm;

SU’UVAD

Spring annuals; leaves highly dissected, the flowers minute, pale yellow.

Widespread across the flora area, sandy to rocky soils, washes, plains, bajadas, hills, and mountains. Often in shaded

niches, mostly beneath spinescent shrubs. This tansy mustard was an important medicinal plant. It was present in

the region more than 10,000 years ago.

Vicinity of Tinajas Altas, Van Devender 05 Mar 1983. Coyote Water, Felger 05-152. Frontera Canyon, 18 Mar 1998, Felger

(observation). tButler Mts, fruit, 10,360 ybp.

Ditnorphocarpa pinnatifida Rollins. Dune spectacle-pod

Spring annuals, often becoming large and sprawling. Flowers white and fragrant; fruits of two disc-shaped segments,

A single, large and many-stemmed plant was encountered in the Tinajas Altas Mountains. It was probably a waif and

apparently not reproducing, presumably due to lack of cross-pollination since they are outcrossers (Andrew Saly-

won, personal communication 2007). Endemic to dunes in nearby northwestern Sonora and southwestern Arizona.

Tinajas Altas, canyon bottom above uppermost tinaja, one large plant seen, no fruit setting, Felger 98-138.

Dithyrea californica Harv. California spectacle-pod

Spring annuals. Flowers cream white and fragrant.

Characteristically on sandy soils. Recorded east of the Butler Mountains and in the Pinta Sands in Cabeza Prieta Refug'-

Butler Mts, Van Devender 27 Mar 1983.

Draba cuneifolia Nutt, ex Torr. & A. Gray [D. cuneifolia var. integrifolia S. Watson. D. cimeifolia var. sonorae (Greed

Parish] Wedge-leaf draba

Small winter-spring annuals; leaves broad and in a basal rosette. Bowers small and white.

Seasonally common; washes, desert plains, sand flats, canyons, and rocky slopes especially in sheltered niches.

species or a similar one was at Tinajas Altas 11,000 years ago.

Camino del Diablo, SSE of Raven Butte, Felger 05-36. Tinajas Altas, 1200 ft, onslopes and in washes. Reeves R5382 (ASU

Frontera Canyon, 18 Mar 1998, Felger (observation). tD. cf. cuneifolia, Tinajas Altas, fruits, 10,950 ybp.

Lepidium lasiocarpum Nutt, subsp. lasiocarpum. Sand pepper-weed; lentejilla

Felgeretal., Flora of Tinajas Altas, Ariz

213

Common and widespread; washes and flats, canyons, and less common on rocky slopes. It has benn in the region for at

least 10,400 years.

Coyote Water, Felger 05-130. Tinajas Altas, wash floor above tanks, Van Devender 10 Mar 1980. Canyon below Raven

Butte Tank, Felger 10-233. tButler Mts, fruits with flattened pedicels, seeds, 8160 & 10,360 ybp.

tLepidium sp./spp. Pepper-weed

Pepper-weed has been in the flora area for more than 18,700 years.

tTinajas Altas, fruits, seeds, 8700-18,700 ybp (5 samples). Butler Mts, fruits, 11,250 ybp.

Lyrocarpa coulteri Hook. & Harv. Lyre-pod; ban censanig

Weak-stemmed and sprawling herbaceous perennials, sometimes to 1+ m tall, and sometimes flowering in the first

season; plants with stellate hairs. Flowers especially fragrant at night, yellow to purple-brown, the petals slender

and often twisted, often 1.5-2+ cm long. Growing and flowering any time of year with sufficient soil moisture.

Widespread in the mo untains , often growing in the protection of large rocks and beneath shrubs and trees. Washes,

canyons, gravelly bajadas, and rocky slopes to higher elevations. This is the only perennial member of the mustard

family in the flora area.

Tinajas Altas, 29 Mar 1930, Harrison 6563.

hysaria tenella (A. Nelson) O’Kane & Al-Shehbaz [Lesquerella tenellaA. Nelson] Desert bladderpod

Cool-season annuals; flowers bright yellow; fruits globose, hollow and bladder-like.

Widespread, especially at lower elevations; washes, sand flats, valley plains, and mountains, often growing in the pri

tection of small shrubs.

Coyote Water, Felger 04-58. Tinajas Altas, Van Devender 5 Mar 1983. Tinajas Altas Canyon, 19 Mar 1998, Felger (observ

tThysanocarpus curvipes Hook. Lacepod

Cool-season annuals. Lacepod was well established in the flora region from at least 9000 to 15,000 years ago. The near-

est present-day population is in the Agua Dulce Mountains on the east side of Cabeza Prieta Refuge.

tButler Mts, 10,360 ybp. Thysanocarpus cf. curvipes, Tinajas Altas, 8970-15,050 ybp (8 samples).

BURSERACEAE— Torchwood Family

Bursera microphylla A. Gray. Elephant tree; toroie; ’usabkam.

large shrubs or small trees, with thick, semi-succulcni limbs and pungcnily aromatic dr.mghi-dcLiduous leaves. Flow-

ers minute, cream-color, produced with the emerging leaves in the first summer rains.

B°cky slopes to peak elevations and sometimes on upper bajadas. The plants often show signs of repeated freeze dam-

age. A large elephant tree in Frontera Canyon measured 4 m tall and 6 m across, with a trunk 1 10 cm tall and 130 cm

in circumference. This species was at Tinajas Altas 6000 years ago, indicating a nearly frost-free habitat.

Tinajas Altas, Vorhies 16 Apr 1924. Tinajas Altas Mts, Van Devender 86-12. tTinajas Altas, seeds, 5940 ybp.

CACTACEAE— Cactus Family

j* me of the 13 cactus species in the present-day flora are defining aspects of the landscape, while others can

** * en onl y by diligent searching. Two species, the many-headed barrel cactus (Echinocactus polycephalus )

3nd beavertail cactus ( Opuntia basilaris), do not range eastward from the arid, general region of the flora area.

214

CYLINDROPUNT1A — Cholla; cholla

Chollas are a conspicuous feature of the landscape and the most diverse cactus genus in the region.

Cylindropuntia acanthocarpa (Engelm. & J.M. Bigelow) F.M. Knuth var. coloradensis (L.D. Benson) Pink*

( *d aCanth0Carpa Engelrn ' & J M Bigelow var - coloradensis L.D. Benson] Buckhorn cholla; CHOW, a° m

Shrub-sized plants; flowers yellow-orange to dull golden yellow, March and April. Fruits become dry upon ripening 18

Common and widespread on upper bajadas, canyons, and rocky slopes to higher elevations. The flower buds were pre-

pared as a vegetable. This species has grown in the region for more than 10,000 years.

E end of Tinajas Alias Pass, Reeves R5395 (ASU). Mesa just E of Tinajas Alias, Felger 08-190. SW side of Tinajas Alias

range, flats adjacent to granitic hills, 10 Jan 2002, Felger (observation). tTinajas Alias, seeds (actually the aril-O*'

ered seeds), 1230-10,070 ybp (6 samples; variety unknown).

Cylindropuntia bigelovii (Engelm.) F.M. Knuth [ Opuntia bigelovii Engelm.l Teddybear cholla; choya gDERa; had-

s adkam (also called hanamI, the general term for cholla). Fig. 3.

215

pale yellow-green; March and April. Frui

cally from readily rooting fallen joints. Packrats have been collecting teddybear cholla for more than 15.7 millennia.

Tinajas Altas, Meams 348 (US). 0.5 km N of Tinajas Alias, Reeves 5393 (ASU). Hats at SW side of Tinajas Altas range, 10

Jan 2002, Felger (observation). tTinajas Altas, seeds (actually the aril-covered seeds), 1230-15,680 ybp (10 samples).

tBuder Mts, spines, seeds, 11,060 ybp.

ylindropuntia echinocarpa (Engelm. &J.M. Bigelow) F.M. Knuth [Opuntia echinocarpa Engelm. &J.M. Bigelow.

0. wigginsii L.D. Benson] Silver cholla

Small chollas (the plants generally smaller than those of C. acanthocarpa). Howers silvery white with green filaments;

Upper bajadas at base of the mountains, onto sandy flats, and generally downslope of the somewhat simillar C.

acanthocarpa.

SE side of Tinajas Altas Mts, 3 Feb 1990, Felger (observation),

ylindropuntia ramosissima (Engelm.) F.M. Knuth [Opuntia ramosissima Engelm.] Diamond CHOLLA

name derives from the diamond-like pattern formed by the low-relief tubercles. Flowers small, pale yellowish to

greenish brown or cream-color with reddish purple outer tepals; opening mid- to late-afternoon, April and May.

Especially common on bajadas along the western side of the Lechuguilla Valley in the Goldwater Range and the Tinajas

Altas region. This xerophy tic cholla is documented from the Butler Mountains more than 10,400 years ago.

E end of Tinajas Altas Pass, Reeves 5394 (ASU). Camino del Diablo SE of Raven Butte, Felger 04-08. SW side of Tinajas

Altas range, nats adjacent to granitic hills, 10 Jan 2002, Felger (observation). tButler Mts, stem, 10,360 ybp.

r. polycephalus. Many-headed barrel cactus, cotton-

TOP cactus, cannonball cactus; biznaga

Multiple-headed, mound-forming barrel cacti with a dense cover of stout spines; young areoles densely white woolly.

The spine surfaces are red but covered by felt-like, overlapping, short, white hairs imparting a dull pink-gray color.

This covering suddenly becomes translucent when wet, revealing the blood-red spine surfaces — the spines turn red

in the rain but soon dry and revert to their usual dull color. Howers yellow and confined (protected?) by the dense

spine cover; flowering late May and June. Fruits densely woolly, drying soon after maturity, tenaciously held among

the closely set spines and may remain in place for one year or more, the seeds falling as the fruits disintegrate. Scat-

tered, mostly on upper bajadas and sandy valley flats.

People in Yuma call it the “cannonball cactus” by because of “the small rounded stems piled up.” A giant mound-shaped

plant on a sandy plain near the eastern boundary of the flora area measured 190 x 201 cm wide, 81 cm tall, and had

more than 152 stems (“heads”). We have not seen seedlings or small juvenile plants, and recruitment presumably is

TTtts cactus has been in the Tinajas Altas region for more than 43,000 years. Evidence from the fossils indicates it was

more common on rocky slopes than today, which is more like the habitat of the northern, or Grand Canyon popula-

tion, var. xeranthemoidesJ.M. Coult., than that of var. polycephalus. In fact, the fossils may be more like the northern

95 kmEof Camino del Diablo on road to Borrego Canyon, Felger 90-20. TButler Mts, seeds, 3820-11,060 ybp (5 sam-

ples). tTinajas Altas, spines, seeds, 9700-15,680 (4 samples), & >43,000 ybp.

-. chrysocentrus (Engelm. & J.M. Bigelow) ROmpler.

j *kawberr y hedgehog cactus; htayita; ’isvigI

Stems several to many, branching mostly from near the base. Howers very showy, 7.5-9 cm long, bright purple-magen-

ta; flowering March and early April. Fruits sweet and edible, ripe late May and early June. Upon ripening, the fruit

P«lp and seeds are quickly consumed by a variety of animals, especially birds that poke a hole in the side of the fruit.

Ants quickly finish off what the birds leave and hollowed-out fruits are common,

tdespread in the Tinajas Altas Mountains, upper bajadas and rocky slopes.

r,I *jas Altas, Blackwell SF-709. Bajada at SW side of Tinajas Altas Mts, 10 Jan 2002, Felger (observation). Ridge above

Tinajas Altas Canyon, 19 Mar 1998, Felger (observation).

216

Journal of the Botanical Research Institute of Texas 6(1)

f Echinocereus sp./spp. (probably E. engelmannii and/or other specie

Hedgehog cacti have grown in the Tinajas Altas Mountains for mon

of present-day Echinocereus are known from Organ Pipe Cactr

Wildlife Refuge (Felger et al. 2007b; 2012).

tTinajas Altas, seeds, 1030-10,600 (5 samples), & >43,000 ybp.

Ferocactus cylindraceus (Engelm.) Orcutt [F. acanthodes (Lem.) Britton & Rose. F. acanthodes var. eastwoodiaeU).

Benson] Mountain barrel cactus

Stem nearly always solitary, reaching 0.5-1+ m tall, straight (erect), becoming considerably taller than wide. Spines red-

dish pink; flowers yellow-green.

Widely scattered on upper bajadas and steep rocky slopes and cliffs. Barrel cacti are fairly scarce in the vicinity of the

waterholes such as at Tinajas Altas and Surveyors Tank and on up to the peaks. Most of the larger ones grow on inac-

cessible cliffs and rock faces. Their scarcity near the tinajas is probably due to bighorn sheep, which dislodge the

plants with their horns and eat the fleshy stem (Warrick & Krausman 1989).

This barrel cactus was at least reasonably common from about 11,000-3800 years ago, but its absence in younger mid-

perhaps bighorn sheep, which knock over the plants and eagerly eat

species characteristically occurs on rock slopes (e.g., Felger 2000).

Borrego Canyon, Felger 90-5. Lower slopes adjacent to Tinajas Altas, Fe

Coyote Water, 24 Oct 2004, Felger (observation). tButler Mts, spine

Altas, seeds, 4010-10,070 ybp (10 samples).

s this change due to climate change or

across much of its geographic range this

82. Near Camino del Diablo on road to

3820-11,250 ybp (5 samples). tTinajas

Grusonia kunzei (Rose) Pinkava [Opuntia kunzei Rose. O. stanlyi Engelm. ex B.D. Jackson var. kunzei (Rose) L.D. Ben-

son. Corynopuntia kunzei (Rose) M.P. Griff.] Desert club-cholla, devil cholla

Thick-stemmed club chollas, commonly forming sprawling colonies, and well armed with rather large and very sharp

spines. Flowers pale yellow or cream-color; May and June, the fruits ripening late August through the following

Scattered on the mid-bajada and upper bajada of the Lechuguilla Valley.

Tinajas Altas, Harbison 9 Mar 1937 (2 specimens, SD 16999 & 17000). Camino del Diablo, E of Raven Butte, 29 Nov 2001,

Felger (observation).

MAMMILLARIA

D. Benson. M. microcarpa Engelm. M. milleri (Britton & Rose) Bodel

cactus; cabeza de viejo; ran ’isvig, ban cekida

Small globose to cylindroid cacti with hooked spines. Flowers pink and rather showy, p

generally following rainfall from April to September. Fruits red and fleshy, s

seeds small and black.

Widely scattered across the region; sandy and rocky soils; upper bajadas and rocky slopes i

cactus, probably this species, has been at Tinajas Altas for more than 18,700 years.

Tinajas Altas, Reeves 5391 (ASU). Granitic mountain at NW side of Raven Butte, Felger 05-384. SE side of Tinajas Alias

Mts, 3 Feb 1990, Felger (observation). tM. cf. grahamii, Tinajas Altas, spines, seeds, 1230-18,700 ybp 01 samples)-

Mammillaria tetrancistra Engelm. Cork-seed fishhook cactus; cabeza de vie jo

Small globose to cylindroid cacti with hooked spines. Flowers pink, spring and summer, fruits red, fleshy, and spine-

less. Basal one third of each seed seated in a corky cup. Although the fleshy part of the fruit is edible, the coiky see*

n Dese

ctushasacorkf

Felgeretal., Flora of Tinajas Altas, Arizona

217

tButler Mts, hooked spines, 740 & 3820 ybp (probably one or both of the present-day species).

Opnntia basilaris Engelm. & J.M. Bigelow var. basilaris. Beavertail cactus. Fig. 18A, B.

Dwarf prickly-pears with a very thick caudex and very thick, mostly upright and short pads. Areoles bear glochids but

no spines. Flowers showy, rose-pink, the stamens with yellow filaments and yellow anthers; flowering early April;

» Refuge and the Tinajas Altas Mountain

ostly on nearly barren desert pavements

han 43,000 years.

In drought the pads shrivel and curl inward, and some or even most of the pads drop off in severe drought almost like

deciduous leaves. Many pads have a dead, dried area below the old flowers or fruits — the pad tips seem to desiccate.

In cultivation, such as in Tucson, these “dwarf” plants can become substantially larger than those in the wild.

Observations: Borrego Canyon, 3 Feb 1990, Felger, Tinajas Altas, 29 Nov 2001 , Felger. tTinajas Altas, seeds (actually the

aril-covered seeds), 1230-11,040 (8 samples), & >43,000 ybp.

iflpuntia chlorotica Engelm. & J.M. Bigelow. Pancake prickly-pear; nopal rastrero

Shrub-sized prickly-pears. The nearest present-day populations are in mountains in the eastern and central parts of the

Cabeza Prieta Refuge and it is more common in the higher mountains of Organ Pipe Monument. This species gener-

ally occurs at the upper elevations of the desert and into oak-grassland. It grew at Tinajas Altas more than 11,000

years ago and has been in the Ajo Mountains for at least 32,000 years. Unlike most other cacti in the region, this

species seems to have expanded its range during the wetter rainfall climates of the Wisconsin glacial environments,

a time of greater winter rainfall and drastically reduced summer rains.

tTinajas Altas, seed (actually the aril-covered seed), 11,040 ybp.

var. transmontanus Engelm.l Desert night-blo

Stems slender, to ca. 1.5 m tall, 4-angled, grayish to grayish purple, mo:

root. Flowers large and white with a long tube; spectacularly fragi

quickly after sunrise. Fruits bright red throughout, the pulp juicy a

one or a few nights between late May and July, the

a nurse shrub. The plants

characteristically grows at low densities

** are edible as a fresh snack food. '

e been utilized medicinally. Packra

iems llke a popsicle. An old, forlorn plant at the Borrego Springs Tank was repeatedly eaten back to a stump

bighorn sheep (Broyles et al. 2007: 597).

■najas Alias- Harbison 6 Mar 1937 (SD 16828); Meams 2811 (DS, US). E end Tinajas Altas Pass, Reeves 5396 (ASU).

CAMPANULACEAE— Bellflower Family

' is orientals (McVaugh) Morin [N. glanduliferus Jeps. var. orientate McVaugh] Redtip thread plant

Co yoteWz

lthough probably n

e widespread.

Felger 05-147. Canyonb

CARYOPHYLLACEAE— Pink Family

s Pecies in the flora area are small, spring annuals.

Journal of the Botanical Research Institute of Texas 6{1)

Felger et al., Flora of Tinajas Altas, Arizona

219

Is, quickly forming a slender t

2.5cmta

partially obscured by needle-lil

Locally common on partially barren pediments of decomposed granite or

Tinajas Altas, adjacent to parking area, a large but highly localized populat

areas without other plants, Felger 93-202.

: side of Tinajas Altas tv

mg white gra

CHENOPODIACEAE, see AMARANTHACEAE

CRASSULACEAE— Stonecrop Family

>r perennial herbs.

ir. eremica (Jeps.) M. Bywater & Wickens.

1. Thick-stemmed perennials; leaves alterm

Crassula connata (Ruiz & Pav.) A. Berger [C. c

&Am.] Pygmy stonecrop

Diminutive winter-spring annuals, the plants s

white, minute.

Sandy and gravelly soils of bajada slopes and car

Camino del Diablo, SSE of Raven Butte, Felger 05

nubs, Felger 10-184.

iarney & Peeblesl Ariz<

?e towards tips; April ai

1 early May. Growing during cooler »

i dry up and the remaining leaves shri

CROSSOSOMATACEAE— Crossosoma Family

Oossosoma bigelovii S. Watson. Ragged rock-flower

Unarmed shrubs to 1.5 m tall, evergreen or tardily drought deciduous. Bowers white and fra

^ early spring.

flora area for more than 43,000 years.

Borre g° Canyon, vicinity of Borrego Tank, Felger 93-194. Tinajas Altas, Webster 24254. tTinaj;

4010-15,680 ybp (9 samples), & >43,000 ybp.

CUCURBITACEAE— Gourd Famil

s with tendrils, the flowers unisext

l. The aerial parts of members of

B^ndegea bigelovii (S. Watson) Cogn. Desert star-vine

nnuals growing with cool-season rains, fall to spring, perishing in late spring; sometimes growing as <

f* 1 - Slender, herbaceous vines from a stout, carrot-shaped white root. Leaves thin and highly v;

sma11 - white, and delicately fragrant.

Felger et al., Flora of Tinajas Altas, Ariz

nd flats and dunes D. serrata

litaxis lanceolata (Benth.) Pax & K. Hoffm. [Argythamnialanceolata (Benth.) Mull. Arg.] Narrowleaf silverbush

Sparsely to densely branched perennial herbs to subshrubs. Herbage silvery pubescent. Rowers inconspicuous, white

and green, flowering during the warmer months.

Common and widespread; washes, canyons, bajadas, and rocky slopes to summit elevations. Often heavily browsed,

probably by bighorn. It was in the Butler Mountains about 8200 years ago.

Tinajas Altas, Felger 04-72. Borrego Canyon, 16 Jun 1992, Felger (observation). tButler Mts, leaves, fruits, 8160 ybp.

litaxis neomexicana (Mull. Arg.) A. Heller [Argylhamnianeomexicam Mull. Arg.] New Mexico silverbush

Non-seasonal annuals or small, short-lived perennials. Rowers green and white, small and inconspicuous.

Common and widespread; washes, valley plains, bajadas, and rocky slopes. It has been in the flora area since more than

Camino del Diablo, SE of Raven Butte, Felger 04-10. tButler Mt

Ditaxis serrata (Torr.) A. Heller var. serrata lArgythamnia s<

Is or perhaps short-lived herbaceous pereii

>uth of Frontera Canyon, 18 Ma

i, 5860 to 8255 ybp (3 samples)

its, seeds, 3820 & 8570 ybp.

) Mull. Arg.] Sand silverbush

:s gray-green. Rowers green and white, s

list and the leaf tips generally truncate ra

uth end of the Tinajas Altas tv

EUPHORBIA— Spurge

Annuals and perennial herbs with milky sap. The individual flowers are generally minute and borne in a cup-

like structure, the cyathium, which looks somewhat like a single flower. The petal-like (petaloid) appendages

of the cyathia of some species can be attractive and the whole structure is indicated here as the “flower.” In

Mexico the Chamaesyce are generally known as golondrina, the term for a swan, apparently in reference to the

female flower on a curved, neck-like stalk, and the general Hia C’ed O’odham name is v’ibam (see E. polycarpa).

p t ate, leaves alternate below and oppos y Subgenus Agaloma: E. eriantha

GOLONDRINA

Small annuals, mostly prostrate, growing with hot-season rains and probably also with cool-season rain. He.

brownish or reddish. The minute cyathia, reduced petaloid appendages, and glabrous capsules are distinctr

Saad y and gravelly soils; washes and sandy flats ol

^ote Water, 25 Oct 2004, Felger 04-44. Coyote \

iablo, 25 Oct 2004, Felger 04-68.

Euphorbia eriantha Benth. Beetle spurge

Non-seasonal annuals mostly encountered in spring,

branched main axis and sparsely branched above, and sparse fo

Sandy, gravelly soils of washes and bajadas of the Lechuguilla Valley

occasionally short-lived perennia

Euphorbia platysperma Engelm. ex S. Watson [Chamaesyce platysperma (Engelm. ex S. Watson) Shinnersl Dune

ing to the glandular-sticky buried portions of the stems forming a sand jacket. Leaves elliptic to oblong or obovatt,

relatively thin, the midrib prominent. “Flowers” yellowish. Seeds smooth, flattened, markedly longer than wide (14

x 1.4 mm). The seeds are larger than most Chamaesyce species and not mucilaginous, unlike those of the other

Chamaesyce in the flora area that become moderately or conspicuously mucilaginous when wet and adhere tena-

ciously to a substrate upon drying.

This species is rare in Arizona and California. It is a dune and sand soil endemic, common in the Gran Desierto dunes

of northwestern Sonora, and also known from extreme northeastern Baja California, and Arizona on dunes and

sand soil at the southwestern margin of the flora area west of Tinajas Altas Mountains near the Mexican border, and

occasional waifs are recorded from southeastern California dunes (Felger 2000; Felger et al. 2007b).

NW corner of Yuma Dunes, 7.4 km N of Mexico border, Douglas 876 (ASU).

Euphorbia polycarpa Benth. [E. polycarpa var. hirtella Boiss. Chamaesyce polycarpa (Benth.) Millsp.] Desert spurge;

golondrina; vi’ibam

also occurring as small perennials. “Flowers” maroon and white, or drought-stressed plants may fail to develop the

white, petal-like appendages.

This is the most common and widespread euphorbia in the region. Sandy soils of washes and bajada-plains, and rocky

Camino del Diablo, SH of Raven Butte, Felger 04-13.

side of Tinajas Altas Mts, 10 Jan 2002, Felger (ol

"Euphorbia cf. polycarpa &/or E. micromera Boiss. ex Engelm

One or both of these species were in the region more than 10,000 ye

taxa are distinguished by subtle differences in the cyathia.

tButler Mts, capsules, seeds, 8160 & 10,360 ybp.

Euphorbia setiloba Engelm. ex Torr. [Chamaesyce setiloba (Engelm. ex Torr.) Millsp.] Fringed spurge; gowndrina

Non-seasonal annuals, especially with summer-fall rains. Plants glandular-pubescent, the herbage yellow-green during

hotter months, otherwise reddish. Cyathia with seven tooth-like white or pink appendages (hence the specift

name) giving the Rower cluster a unique star-like look. The plants are frost-sensitive.

Widely scattered, probably mostly at low elevations; often in broad, sandy-gravelly washes, canyon bottoms, and bajadas.

Camino del Diablo, SE of Raven Butte, Felger 04-14. Wash, 2 mi SE of Tinajas Altas, Felger 08-205.

tEuphorbia sp./spp. (Chamaesyce)

One or more unidentified Chamaesyce were at Tinajas Altas at least 5,000 to more than 11,000 years ago.

tTinajas Altas, capsules, 5940-11,040 ybp (4 samples).

Jatropha cuneata Wiggins & Rollins. Desert lime

Shrubs often 1-1.5 m tall, oozing blood-like sap v

drought deciduous, appearing with rains at var

small and white, with rains from July to Octobei

widespread on rocky slopes and upper bajath

2 mi SE of Tinajas Altas, Felger 08-205. Granitic hills, SW

seeds appear identical and the two

north. This w

the region is about 1250 years old.

t— hence the name sangrengado (dragon’s blood). Leaves

sons except during the coldest weeks or months. Flow® 5

sometimes freeze-damaged during severe win

h of Tinajas Altas Pass, uncoi

n plant for basket-making by the Hia C’ed O’odhai

,r absent farther

ie prehistoric record in

Felger etal., Flora of Tinajas Altas, Arizona

Tinajas Altas, near base, Harrison 6567. Borrego Canyon, 3 Feb 1990, Felger (observation), t Tinajas Altas, 550 m, leaves,

seeds, 1230 ybp.

FABACEAE — Legume Family

Annual or perennial herbs, shrubs, trees, or vines, many with nitrogen-fixing bacteria in root nodules. Leaves

alternate and once- or twice-pinnately or palmately compound. The trio of Olneya, Parkinsonia, and Prosopis

are the most important desert trees of the region and are significant nurse plants for an array of herbs and

larger perennials including saguaros. These trees featured importantly in the economies of the local peoples

for food, fuel, shelter, fiber, medicine, and provided some of the rare shade (e.g., Felger 2007; Felger & Moser

1985; Felger et al. 1992; Hodgson 2001; Rea 1997). The Sonoran Desert flora includes 281 legume species, or

about 11% of the total flora, while legume species make up only 4% of our flora.

Acacia, see Senegalia

acmispon

Annuals or herbaceous perennials. Leaves pinnate with 3-7 leaflets. Bowers yellow to red-orange, pea-like.

Pods multiple seeded.

Perennial subshra

guide soon de

.) Greene. Hosackia rigida Benth.l Desert

n long, the corollas at first bright yellow, a

:the Tinajas Altas Mountains, arroyos and canyons among boulders and rocky slopes, often east-a

north-facing. Documented at Tinajas Altas from about 4000-10,800 years ago.

^jas Altas, Harrison 3610. Tinajas Altas, crevices in granite, 1400 ft, Engard 925 (ASU). Surveyors Canyon, cany

bottom just below Surveyors Tank, Felger 10-203. Frontera Canyon, 18 Mar 1998, Felger (observation). tTina

Altas, fruits, seeds, 4010-10,750 ybp (7 samples).

cmis P«n strigosus (Nutt.) Brouillet [Hosackia s trigosa Nutt. Lotus strigosus (Nutt.) Greene var. tomentellus (Greei

kely. Ottleya strigosa (Nutt.) D.D. Sokoloff] Hairy LOTUS

W inter-spring annuals, mostly low growing or prostrate; leaves succulent on well-watered plants. Flowers 7.5 9 n

iong, the petals bright yellow with red nectar guidelines on the banner.

Wadies and canyon bottoms, and soil pockets on rocky slopes.

Tinajas Altas, Van Devender 9 Mar 1980. Tinajas Altas Canyon, 19 Mar 1998, Felger (observation).

Journal of the Botanical Research Institute of Texas 6(1)

Dalea mollis Benth. Silky dalea

occurs in adjacent areas. It was at Tinajas Altas about 5900-6000 years ago.

margin of Davis Plain at base of Gila Mountain, W branch of Camino del Diablo,

sandy soil, Felger 05-67. tTinajas Altas, fruits, 5860 & 5940 ybp.

; semi-succulent. Flowers pale lavender-pink.

Tinajas Altas, Van Devender 25 Mar 1983. Tinajas Altas Pass, Reeves R5400 (ASU). Surveyors Canyon, canyon bottom,

Felger 10-212.

tLupinus sp./spp.

Lupines are documented for the region for nearly 10,000 years.

tButler Mts, fruits, calyx, seeds 740 & 3820 ybp. tTinajas Altas, seeds, 1230-9900 ybp (9 samples).

i and limbs often

e drought £

nd population might not flower ei

er stature on rocky slopes. A number of mighty ironwoods are scattered about the arroyo-wasl

Muerto and the lower tinajas, and the lai

a massive trunk (Fig. 19). This species is

s of desert bighorn and n

s. Olneya is one c

trly 10,000 years, which is the oldest known record for this species anywhere,

tortant source of wood and fuel, and the seeds were variously prepared for food (Felger 2007).

vood stumps can be seen throughout the region. For example, numerous large ironwood stumps and

era Canyon appear to be axe- or saw-cut, and many such stumps were noted on the desert flats about

i of the Tinajas Altas site. The ironwood population has largely recovered from the extensive wood

few palo verde or ironwood trees are still standing and may furnish horse feed, but the traveler should bring his own

1.5 miles:

Tinajas Altas, wash just E of the tinajas, Felger 08-171. Tinajas Altas Canyon, 19 Mar 1998, Felger (observation). Fronton

Canyon, 18 Mar 1998, Felger (observation). tButler Mts, spines, leaves, fruits, 740-8570 ybp (3 samples). tTmaj*

Altas, twigs with spines, leaflets, 1230-9900 ybp (4 samples).

PARKINSONIA— Palo verde

Trees and large, heavy-branched shrubs with green bark and relatively soft wood. Leaflets small, the leaflet 5

and leaves quickly drought deciduous, the leaflets often falling independently; leaves twice pinnate or appear-

ing once pinnate. Flowers yellow, caesalpinioid, produced in prodigious quantities in spring. Pods ripening®

early summer, indehiscent to tardily partially dehiscent, 1-several-seeded.

Fdgeretal., Flora of Tinajas Altas, 1

Parkinsonia Honda (Benth. ex A. Gray) S. Watson [Cercidium floridum Benth. ex A. Gray] Blue palo verde; palo

verde; ko’okomadk, kausp. Fig. 15.

One of the several larger tree species in the region, usually with a well-developed trunk. Flowers bright yellow; mass

flowering in spring and sometimes sporadically in fall.

Blue palo verde trees are fairly common in canyon bottoms and the larger washes along the eastern side of the Tinajas

Altas Mountains, such as the broad wash leading out of Borrego Canyon, and many of the small washes on the upper

bajada. Only two or three blue palo verdes, however, were found between 2000 and 2006 in the wash below Tinajas

Altas the absence of more of the trees perhaps due to earlier woodcutting, but by 2008 even these trees were gone,

apparently due to drought and firewood collecting by visitors. Although the soft wood is of poor quality for fuel, it

was utilized because little else was available. The seeds served as food for people. The trees were common on the

Davis Plain west of the Gila Mountains, although many of them perished during extreme drought of 2004 and 2005.

Palo verdes and ironwoods were not seen along Coyote Wash in the vicinity of Coyote Water, yet palo verde is com-

mon and ironwood present only eight km south in La Jolla Wash. At the northern end of the study area, blue palo

verde replaces foothill palo verde in certain unexpected circumstances, such among the basalt boulders of Raven

Butte— hardly typical habitat, but the basalt is a veneer atop course granitic aUuvium, and perhaps well drained and

not so different from an arroyo. This species has been in the region for at least 8300 years.

Tinajas Altas, wash. Van Devender 86-11. Arroyo 2 mi SE of Tinajas Altas, Felger 08-195. Borrego Canyon, 16 Jun 1992,

Felger (observation). Camino del Diablo, E of Raven Butte, 25 Oct 2004, Felger (observation). Canyon wash at NW

side of Raven Butte, 30 Dec 2005, Felger (observation). tButler Mts, twigs, spines, fruits, 740 & 3820 ybp. TTinajas

Altas, twigs, spines, 4010 & 8255 ybp.

Parkinsonia microphylla Torr. [Cercidium microphyllum (Torr.) Rose & I.M. Johnst] Foothill palo verde, little-

leaf palo verde; palo verde; kek cehedagI

Small trees or large shrubs. Flowers pale yeUow with a white banner petal; mass flowering in April and early May.

Upper bajadas on the east side of Tinajas Altas Mountains; canyons, washes, and rocky slopes from the base to higher

elevations on all slope exposures in the mountains but scarce north of Tinajas Altas Pass. The seeds were a signifi-

cant food resource for people.

Tinajas Altas Canyon, Felger 08-177. Frontera Canyon, 18 Mar 1998, Felger (observation).

Prosopis glandulosa Torr. var. ton

Benson. P. odorata Torr. & Fr6m.,

Trees and shrubs. Leaves appearing :

a (L.D. Benson) M.C. Johnst. [P. juliflora (Sw.) DC.

R. Palacios 2006| WESTERN HONEY MESQUITE; MEZQl JITE:

aneously in spring after the last freezing weather, the ]

one pair of pinnae. Flowers dull ;

sporadically through early fall; pods mostly ripening in early summer ar

squites are concentrated along the larger drainageways

:ur, especially along Coyote V

a primary seasonal food for

Mesquite was the single most useful plant in the Sonoran Desert; providing wood for const™

weapons, and was the preferred cooking fuel. The mesocarp of the pods w

fashioned into cordage. The whitish gum is edible and was used medicinal

bark. This black pitch was also used as dye (Felger 2007; Felger & Moser 1985).

Mesquites in the flora area have leaves with one pair of pinnae and the typically elongated and rather widely spaced

leaflets characteristic of P. glandulosa, but some specimens are conspicuously pubescent like P. velutina rather than

being glabrate or glabrous as in “typical” P glandulosa. The Tinajas Altas population is within a broad area oi mes-

quites of intermediate appearance (Felger 2000). „ „ ,

Water, Felger 04-62. Base of Tinajas Alias, Harrison 6581. 1 mi N of Tinajas Altas, 17 Apr 1948, Kurtz ll»lherb-

pubescent!. Surveyors Canyon, canyon bottom below Surveyors Tank, Felger 10-216. Frontera Canyon, 18 Mar

!998, Felger (observation).

Proso PB glandulosa &/or P. velutina Wooton. Mesquite

Mesquite has been well established in the region for more than 18,750 years

determine which of the two species were gathered by Ice Age packrats.

tButler Mts, leaflets, 740 & 8160 ybp. TTinajas Altas, spines, leaflets, endocar

> the black pitc!

The fossil specimens do not allow

«, 1230-18,700 ybp (14 samples).

Journal of the Botanic

Felgeretal., Flora ofTinajas Altas, A

Lfrhng iiilla Valley in larger washes and less ofte

r drainageways of the bajada, and Tinajas Altas Moun-

»cky slopes, especially east- and north-facing, from low to high elevations. Catclaw

is one of the favorite foods of desert bighorn (Russo 1956). Packrat collections show it has been in the Tinajas Altas

Mountains for more than 43,000 years.

Coyote Water, Felger 04-21. Wash just E of Tinajas Altas, Felger 08-188. Canyon above Tinajas Altas, 26 Oct 2004, Felger

(observation). tButler Mts, stem prickles, leaflets, fruit fragments, 740 to 8160 ybp (3 samples). TTinajas Altas, twigs

(prickles), leaves, 1230 to 11,040, & >43,000 (14 samples).

FOUQUIERIACEAE — Ocotillo Family

Fouquieria splendens Engelm. subsp. splendens. Ocotillo; ocotilw, melhog. Figs. 9 and 11.

Unique, long-lived shrubs with wand-like spiny branches arising from a very reduced trunk. Short-shot

at almost any time of year following a ground-soaking rain except during freezing weather, and are

the soil dries or with freezing weather. Long-shoots are produced during times of warm or hot weath

moisture. Flowers red-orange, generally in spring.

Widespread; sandy soils of upper bajadas and rocky slopes t<

traditional Hia C’ed brush house. The flowers provide s

smut elevations. Ocotillo served as a framework for the

presence of ocotillo sticks, identified as ‘spiritual sticks’ in a rockshelter near Tinajas Altas” (Rankin 2000

The oldest Sonoran Desert records are from the late Holocene, about 4500 years ago in the Homaday Mounts

Pinacate region of nearby northwestern Sonora (Van Devender et al. 1990a). Considering the ecological e

( range of habitats) and wide geographic range of this species, its apparent late arrival and rarity in packrs

assemblages in the flora area is somewhat surprising.

Camino del Diablo, SE of Raven Butte, Felger 04-15. Tinajas Altas Canyon, Felger 08-181. Tinajas Altas Mts, neai

mit, 26 Oct 2004, Felger (observation). TButler Mts, stem fragment with spines, 740 ybp.

GERANIACEAE— Geranium Family

ERODIUM— Storks bill

Winter-spring annuals with small, lavender-pink flowers. The long, corkscrew beak on each fruit segment

uncoils when moistened and screws the sharp-pointed and heavier seed-bearing end into the ground.

Not widespread in the flora area although common in nearby areas, especially in disturbed sites. Documented in the

vicinity of Coyote Water and rare along the canyon above the Tinajas Altas. Native to the Mediterranean region and

widely naturalized in the New World.

Coyote Water, Felger 05-143. Tinajas Altas Mts, above the tinajas, among rocks, rare, grazed, 19 Mar 1998, Felger

Erodhun texanum A. Gray. False filaree, desert stork’s bill

Widespread and common; washes, bajadas, plains, and sandy flats. It has been in the flora area for at least ten millennia.

Gumno del Diablo, SSE of Raven Butte, Felger 05-34. TButler Mts, fruit, 10,360 ybp. TTinajas Altas, fruit, 8970 ybp.

HYDROPHYLLACEAE, see BORAGINACEAE

KOEBERLINIACEAE— Allthorn Family

tK «*ertinia spinosa Zucc. Crucifixion thorn, allthorn; corona de cristo

Wscent woody shrubs, essentially leafless.

■Hte nearest population occurs along the northeastern border of Organ Pipe Monument, except for a single known shrub

from the Pinacate Region (Felger 2000). There is one fossil record for it in the Butler Mountains.

tButfer Mts, twig, 10,360 ybp.

Journal of the Botanical Research Institute of Texas 6(1)

KRAMERIACEAE— Ratany Family

KRAMER1A

Small shrubs with small drought-deciduous leaves; reported as root parasites on other shrubs. Flowers bilater-

ally symmetrical, purple and yellow, and attractive; the 5 sepals are petal-like and the 5 petals modified into a

landing-pad lip and two slab-like, thick elaiphores. Female Centris bees are the pollinators, gathering saturated

fatty acids from the elaiphores (Simpson & Neff 1977).

1 . Branches mostly straight and without knotty spur-branches; the 3 upper petals distinct (free), the blades near orbicular;

; claws of the 3 upper petals fused toward base, the blades

r, 1906] White ratany; cOsahw; edho, he:d

various seasons following sufficient rainfall, especially in

: the region, especially in sandy to gravelly soils of washes,

Krameria bicolor S. Watson, 1886 [K. grayi Rose & Pail

Sprawling shrubs. Flowers magenta-purple (deep rose),

spring.

Common and widespread through most of the lowlands

plains, and bajadas, and sometimes on rocky slopes.

Camino del Diablo, SE of Raven Butte, Felger 04-16. Tinajas Altas, canyon below tanks and on rocky slopes, 19 Mai

Felger (observation).

Krameria erecta Willd. ex Schult. [K. parvifolia Benth.] Little-leaf ratany, range ratany; cOsahui

Dwarf woody shrubs with knotty short-shoots. Flowers bright magenta-purple, at various seasons following suf

Scattered on sparsely vegetated granitic-gravi

tains for more than 8200 years.

0.3 mi SE of Tinajas Altas, Felger 08-191. Butle

and spines from fruits, 740 & 8160 ybp.

lissected bajadas, often with K. gray i

5, 27 Mar 1983, Van Devender (obsen

LAMIACEAE— Mint Family

:s opposite. Flowers bilaterally symmetrical, violate, blu<

Hyptis albida Kunth [H. emoryi Torr.J Desert

Widespread, especially in washes, also in canyon bottoms and on rocky slopes to the mountain summit. The fossil re-

cord for the flora area extends to 8700 years.

Hyptis emoryi seems best regarded as a synonym of H. albida (see Felger 2007; Martin et al. 1998; Felger & Wilder 2012).

The Sonoran Desert populations tend to be more densely white-pubescent than those of H. albida from non-desert

regions farther south in Mexico, but the variation is continuous. The leaves of well-watered desert plants, even in the

Tinajas Altas region, especially following favorable, warm-season rains, can be much larger, greener (sparser pubes-

Tumer et al. 1995). Several varieties of H. emoryi have been described and across the large geographic range there is

1 mi N of Tinajas Altas, Kurtz 1161. Tinajas Altas Pass, Reeves R-5421 (ASU). Frontera Canyon, 18 Mar 1998, Felger (ob-

servation). Camino de Diablo, E of Raven Butte, 26 Nov 2001, Felger (observation). Tinajas Altas Canyon, Fel S eT

183. TButler Mts, twigs, leaves, fruits, 740-8570 ybp (3 samples). TTinajas Altas, leaves, calyces, seeds, 1230-87W

ybp (6 samples).

tMonardella arizonica Epling. Arizona monardella

Bushy, herbaceous or subshrub perennials.

229

This mint is documented from the Tinajas Altas Mountains from about 9000 to more than 43,000 years ago. The nearest

present-day population occurs in canyons in the Ajo Mountains in Organ Pipe Monument. It occurs elsewhere in

southern and western Arizona, but generally above the desert.

tTinajas Altas, involucres, fruits, 8979-18,700 (9 samples), & >43,000 ybp.

tSalvia mohavensis Greene. Mojave sage

Broad, spreading shrubs with woody stems.

The fossil record in the flora area extends from about 9300 to more than 43,000 years. It was widespread in the Sonoran

Desert region through the late Pleistocene. The nearest present-day occurrences are relictual populations at higher

and Mohawk Mountains in southwestern Arizona. Mojave and Sonoran deserts in southeastern California, south-

3,360 & 11,060 ybp. tTinajas j

fragments, calyces, 9230-18,700, & > 43,000 ybp

Scutellaria mexicana (Torr.) A.J. Patoi

•xicana Torr.] Bladder-sage

y with sparse foliage, the stems interlacing and branching at right angles, twigs often rigid a

t. Flowers white and blue; mass flowering at various seasons following rains. The calyx enlai

Shaded canyon bottoms in at least several mo u ntains i

>llas) relatively large (1.8-2.5 cm long) and attractive, white with

Cabeza Prieta Refuge, one record in Organ Pipe Monument, and

ut probably more widespread in the region. The Coyote Wash

:ona and California where it is rare.

' species in this large genus confined

Coyote Wash at Camino del Diablo, clay-like soil in dense mesquite brush, Felger 02-14.

LOASACEAE— Stickleaf Family

Annuals or perennials with silicified or calcified, barbed hairs (the barbs usually in whorls), these hairs result-

®g in the leaves and fruits sticking like Velcro.

1 ^ or subshrub perenr

^OTKELIA Stickleaf; pegapka

or summer annuals, and sometimes herbaceous perennia

albicaulis (Douglas ex Hook.) Douglas ex Torr. & A. Gray [This species complex includes M. deserto-

r (Davidson) H.J. Thomps. & J.E. Roberts] WhiteSTEM blazing STAR

“‘-season annuals. Flowers bright vellow, sometimes with a dark orange or reddish center.

230

Sandy and gravelly soils of the bajadas, arroyos, and washes. M. desertorum is distinguished on technical features.

Butler Mts, dunes, Van Devender 27 Mar 1983. W end of pass just N of Tinajas Altas, Van Devender 6 Mar 1983.

Mentzelia longiloba J. Dari. var. yavapaiensis J .J. Schenk & L. Hufford [M. multiflora (Nutt.) A. Gray subsp. longi-

l oba 0- Dari.) Felger. Blazing star; fegapega

Cool-season ann«ak and perhaps sometimes short-lived perennials. Flowers bright yellow.

Lower canyons and washes in the Tinajas Altas Mountains; apparently not common in the flora area.

Frontera Canyon, 18 Mar 1998, Felger (observation).

Mentzelia puberula J. Dari. Silver blazing star

Facultative annuals to short-lived perennials, the plants often bushy; leaves often silvery gray-green. Flowers pale yel-

Canyons and rocky slopes; widely scattered but seldom common. A fossil from the Butler Mountains dates from about

3800 years ago.

Butler Mts, Van Devender 27 Mar 1983. Tinajas Altas, cliff face, Reichenbacher 480. tButler Mts, twigs, seeds, 3820 ybp.

Petalonyx linearis Greene. Narrowleaf sandpaper plant

Small, densely branched, rounded shrubs, usually less than 0.5 m tall. Flowers white; flowering at least in March and April

Localized on widely scattered arid, gravelly-sandy washes and rocky slopes. It is documented for the flora area for 9

millennia.

Tinajas Altas, 1 Dec 1938, Goodding 5335. Tinajas Altas Mts, 1.7 km WNW of Tinajas Altas Peak, wash of granitic sand

and rocks, infrequent, 340 m. Baker 13307 (ASU). tTinajas Altas, leaves, 1230 & 9230 ybp.

MALVACEAE— Mallow Fam

f Abutilon sp./spp.

This genus is documented for the flora area from 5900 to about 11,000 years ago. Three species of small shrubs

in this genus occur in Cabeza Prieta Refuge: A. incanum (Link) Sweet, A. malacum S. Watson, and A. palmeri S.

Watson; and a fourth one, A. abutiloides (Jacq.) Garke ex Britton & P. Wilson, occurs in Organ Pipe Monument,

but none presently occur in the flora area.

tTinajas Altas, mericarp fragments, 5860-11,040 ybp (3 samples).

Hibiscus denudatus Benth. var. denudatus. Rock hibiscus

Perennial subshrubs. Corollas white to pinkish with maroon spots in the center, flowering at various seasons, espe-

cially during warmer months.

Widespread and common in the Tinajas Altas Mountains; pediments, canyons, and slopes to higher elevations, "fl*

hibiscus has been in the Bora area for at least 15,700 years.

1 mi N of Tinajas Altas, Kurtz 1157. Tinajas Altas, ridge above tanks, 1420 ft, Felger 04-91. tButler Mts, stems, leaf N -

ments, 740 & 8570 ybp. tTinajas Altas, leaf fragments, seeds, 5860-15,680 ybp (5 samples).

HORSFORDIA— Velvet mallow

Tall, slender and sparsely branched shrubs. Flowering at various seasons except during coldest and driest weather-

Horsfordia alata (S. Watson) A. Gray. Pink velvet-mallow

Spindly shrubs (1.7) 2.5-4 m tall. Flowers pale lavender-white to pink, drying bluish to pale lavender.

Small to large washes in upper bajadas, canyons, and rocky slopes to higher elevations.

Large arroyo in front of the Tinajas, Felger 04-73 . 1 mi N of Tinajas Altas, Kurtz 1158. Observations: Cam

E of Raven Butte, 10 Jan 2002, Felger, Granitic hills at SW side of Tinajas Altas range, 10 Jan 2002, Fd

ainodelDi^

felgeretal., Flora ofTinajas Altas, Arizona

231

Horsfordia newberryi (S. Watson) A. Gray. Orange velvet-mallow

Few-branched shrubs to 2.5+ m tall, often considerably shorter. Flowers bright yellow-ora

Canyons and rocky slopes in the mountains.

Borrego Canyon, 16 Jun 1992, Felger (observation).

SPHAERALCEA— Globemallow; mal de o;o

it section short and stubby, less than half i

v Raven Butte Tank, Felger ]

iable in size, from less than 1

/alley floors, bajadas, and lc

Robust, large plants, mostly on sandy

soils, have been called S. orcuttii; other than the size of the plant, there are no apparent differences (Felger 2000).

Coyote Water, banks of wash, 1.5-1.8 (2) m tall, many, few-branched erect stems, Felger 05-141. High Tanks Gate, Felger

98-111. Camino del Diablo at Coyote Wash, 10 Jan 2001, Felger (observation).

Sphaeralcea emoryi Toxt. ex A. Gray. Emory globemallow; mal de ojo

Annuals or short-lived perennials. Flowers orange to red-orange; spring and summer-fall rainy seasons.

Locally in fine-textured soils at the south end of Coyote Wash in the Lechuguilla Valley.

Camino del Diablo at Coyote Wash, localized in the mesquite “forest,” Felger 10-167.

Sphaeralcea sp./spp. (may be, or includes, S. ambigud)

tButler Mts, twigs, leaf fragments 8 mericarps, 740-11,250 ybp (4 samples). TTinajas Altas, leaf fragments, mericarps,

seeds, 4010-10,950 ybp (10 samples).

MARTYNIACEAE— Devils Claw Family

Ptoboscidea altheifolia (Benth.) Decne. Desert devils claw; cuernitos, urn de gato ; ban 'ihugga

Strongly scented, herbaceous perennials from a deeply buried, large, tuberous root. Flowers showy, often 4 cm long,

yellow with yellow-orange and bronze markings; growing and flowering with hot weather.

^ndy to silty soils of washes, floodplains, and valley plains.

Qun ®o del Diablo, SE of Raven Butte, Felger 04-18.

MOLLUGINACEAE— Carpetweed Family

***JJ“»» eerviana (L.) Ser. Thread-stem carpet-weed, Indian chickweed

snail-shaped with dark ridges (striae) on the dorsal side.

“ soils along Coyote Wash in the Lechuguilla Valley. Widespread in the Sonoran Desert and elsewh<

Water, Felg^-50 ^

NYCTAGINACEAE —Four O’clock Family

or perennial herbs or subshrubs. Sepals petal-like, the petals none. Fruits 1-seeded and achene- or nut-

’ encl °sed by the calyx tube and technically an anthocarp, the collective structure here called the “fruit.”

Journal of the Botanical Research Institute of Texas 6(1)

Abronia viUosa S. Watson var. villosa. Sand-verbena; verbena de la arena

Cool-season annuals; highly variable in size. Flowers diurnal, showy, very fragrant, pinkish purple to pale magenta.

Fruits winged and beaked in a globose cluster.

Common and widespread on sandy plains and dunes at the western margin of the flora area near the base of the Butler

Mountains. Although not documented elsewhere in the flora area, it is common and widespread on sandy soils in

many nearby areas.

Butler Mts, Van Devender 27 Mar 1983.

ING FOUR-O’CLOCK

Short-lived perennials with slender, trailing stems, and also flowering in the first season. Flowers clustered in trios re

sembling a single flower, violet-rose or magenta; flowering non-seasonally in wanner months with sufficient soil

Widespread across the flora area in many habitats including gravelly floodplains, washes, canyons, flats, and rocky

slopes.

Coyote Water, Felger 04-23. Camino del Diablo E of Raven Butte, Felger 01-583. Tinajas Alias Pass, Reeves R5415 (ASU)-

BOERHAV1A — Spiderling; jauniupin

Summer annuals and one perennial; the plants at first usually upright, later often spreading ai

decumbent or even prostrate. Flowers open in the early morning and collapse with daytime heat.

d flowering branches may be

n long, chunky (broadly obovate), all or mostly 4-ar

iribaeajacq.] Scarlet si

-than the other

.

;hwest<rn V' rlh

with sufficient soil moisture. Flowers small and bright red-purple, remaining open lc

boerhavias in the region. Fruits glandular-pubescent.

Localized and well established at the base of the tinajas; not seen elsewhere in the flora ar

sites regionally. This weedy species, now cosmopolitan in distribution, is probably

America but does not seem to be native in the Tinajas Altas and nearby regions (Felger 2000).

Tinajas Altas, near the lower tinaja: 15 Jun 1992, Felger 92-609; 29 Nov 2001 , Felger 01-582 ; 28 Mar 2010, Felger 10-

5; juantiupIn; makkumI ha-jeved

rate hairs, the flowering bra

Vash.

Coyote Water, Felger 04-28.

Felgeretal., Flora ofTinajas Altas, Arizona

233

Boerhavia spicaia Choisy [B. coulteri (Hook, f.) S. Waison. B. coulteri var. palmeri (5. Watson) Spel

Choisy var. palmeri S. Watson] Palmer’s spiderung; juantiupIn

Summer annuals; flowering branches conspicuously glandular-sticky, sticking to shoes, socks, and pai

very small, white to pale pink, on racemose branches. Fruits 5-angled.

Various habitats: sandy to rocky soils; washes, bajadas, flats, and slopes; widespread and often very com

Coyote Water, Felger 04-29.

Boerhavia wrightii A. Gray. Large-bract spiderung

Summer annuals, densely glandular sticky including the flowering branches. Flowers pinkish white tt

mose branches. The plants are often rather robust and the floral bracts, flowers, and fruits are largf

the other bocrhavias in the region. Fruits 4-angled.

Sandy to rocky soils in many habitats; seasonally common and nearly ubiquitous across the flora area. 1

men about 8200 years old from the Butler Mountains.

Gamino del Diablo, SE of Raven Butte, Felger 04-08. Coyote Water, Felger 04-30. Tinajas Altas, 26 Oct 2l

servation). tButler Mts, fruit, 8160 ybp.

MIRABILIS— Four O’clock

Herbaceous perennials; flowei

r pale pink, collapsing with daytime heat, t

Mirabilis laevis (Benth.) Curran var. villosa (Kellogg) Spellenb. [M. bigelovii A. Grayl Desert four o’clock

Herbaceous perennials, with glandular-pubescent herbage. Flowers white or pale pink, flowering at various seasoT

Widely scattered on rocky slopes to higher elevations, canyons, washes, valley plains, and bajadas. There is a sped

about 11,100 years old from the Butler Mountains.

Borrego Canyon, canyon slopes, Felger 92-614. Surveyors Canyon, Felger 10-199. Tinajas Altas, Goodding 1517. Tinaj

). Coyote Water, Felger 98-119. Camino del

Alas Pass, Reeves 5:

:r (observation). tButler 1

fruit, 11,060 ybp-

j,SE of Raven Butte, 25 Oct :

Canyons in Tinajas Altas Mountains. The only other known locality for this speries in Arizona is in the Kofa Mountains.

The plants differ from M. laevis in being more robust, and generally having larger, thicker, and more yellowish

Borrego Canyon, common in canyon bottom, Felger 92-613. Surveyors Canyon, Felger 10-200. Frontera Canyon, Felger

98-108. Tinajas Altas Mts, Goodding 07 Mar 1940 (ASU). Yuma Co: Palm Canyon, ca. % way to the top, 8 Oct 1977,

ONAGRACEAE— Evening-Primrose Family

AU members of this family in the flora area grow and flower during the cool seasoi

pertine (opening near sunset) and collapse after sunrise depending on temperature

Tinajas Altas, Harrison 3608. Canyon above Tinajas Altas, Felger 04-83.

Chylismia claviformis (Torr. & Frem.) A. Heller subsp. peeblesii (Munz) W.L. Wagner & Hoch [Camissom

claviformis (Torr. & Fr&n.) P.H. Raven subsp. peeblesii (Munz) P.H. Raven] Browneyes

Annuals. Flowers white, sometimes with pink tinges, and with a red-brown center, the flowers becoming pink with age

and often purplish when dry; the stamens, style, and stigma white.

Widespread, especially on sandy to gravelly soils; washes, valley plains and bajadas, canyons, and rocky slopes.

Tinajas Altas, Van Devender 5 Mar 1983. Canyon below Raven Butte Tank, Felger 10-222. Tinajas Altas Pass, McLaughlin

lerioides (A. Gray) W.L. Wagner & Hoch [(

enerioides (A. Gray) P.H.F

sunset, usually collapsing

the floral structure and m

Is. Flowers very small, at first white, often becoming pink with age; opening m

ir sunrise. This is the smallest-flowered evening primrose in the Sonoran Des<

ions are characteristic of self-pollinated flowers,

pread, especially on rocky slopes from low to high elevations.

5 Altas, rocky slope, 1900 ft. Van Devender 5 Mar 1 983. Tinajas Altas, Goodding 6 Mar 1940 (ASU). Surveyors Can-

i, canyon bottom, Felger 10-211. Above the tinajas, 19 Mar 1998, Felger (observation).

alifomic

becoming orar

Widespread acros;

Tinajas Altas, Var

Surveyors Can

erect and often sparsely branched, the leaves usually <

inge with age and often drying pink.

tn Devender 05 Mar 1983. Tinajas Altas Canyon, above

nyon, canyon bottom, Felger 10-206. Vicinity of Coyote

l by the time of flowering. Flowers yellow,

., 19 Mar 1998, Felger (obs

Oenothera primiveris A. Gray. Yellow desert evening-primrose

Annuals. Flowers relatively large and lemon-yellow. Capsules woody, persisting on the dried stems long after the plants

perishes.

Sandy gravel or silty-ckyish soils of washes, sand flats, canyons, and sometimes in soil pockets on rocky slopes. Speci-

mens in the region are generally identifiable as subsp. bufonis (M.E. Jones) Munz, distinguished by larger and gener-

ally cross-pollinated flowers.

Coyote Water, Felger 04-54. Canyon below lowermost Tinajas Altas, 19 Mar 1998, Felger (observation).

OROBANCHACEAE — Broomrape Family

tCASTILLEJA sp./spp.

There are records for the genus dated at 11 millennia in the flora area and 20.5 millennia in the Ajo Mountains

in Organ Pipe Monument. The nearest present-day members are C. exserta (A. Heller) T.l. Chuang & Heckard

subsp. exserta (owl’s clover) at Jose Juan Charco in Cabeza Prieta Refuge and C. lanata A. Gray (Indian paint-

brush) in the Ajo Mountains.

tTinajas Altas, seeds, 11,040 ybp. Ajo Mountains, 20,490 ybp.

Orobanche cooped (A. Gray) A. Heller. Desert broomrape; mo’otadk

Apparently annuals; parasitic on roots of Ambrosia shrubs including A. dumosa and sometimes A. ilicifolia and A. salsda

and also reported on Larrea (Wiggins 1964). Stems very thick and succulent, appearing in spring, flowering and

withering by the end of April or sooner. Flowers purple and white, and the throat marked with yellow.

Often locally common on sandy flats and bajadas, sandy-gravelly washes, and canyon bottoms. Widespread across the

flora region. The young plants are edible after thorough roasting.

Near jet of Camino del Diablo and Cipriano Pass Road, Tinajas Altas Mts, Von Devender 6 Mar 1983. Frontera Canyon,

Felger 98-104. Camino del Diablo (West Rte), Davis Plain W of Tinajas Altas Mts, Reeves R5427 (ASU).

PAPAVERACEAE — Poppy Family

Eschscholzia minutiflora S. Watson. Little gold-poppy

i annuals. Flowers small and yellow.

Cool-!

Felgeretal., Flora of Tinajas Altas, Ariz

235

Seasonally common during years of favorable rains in the Tinajas Altas Mo

t jiff so many other species of ephemerals in the more arid regions of th

ing dry years. In spring 1983 Tom reported it as being rare, and Richai

spring 2010 field trip, when it was widespread and common.

Vicinity of Tinajas Altas, rare, under Larrea , Van Devender 5 Mar 1983. Tin

Felger 10-188. Surveyors Canyon, canyon bottom, Felger 10-206.

PLANTAGINACEAE (includes Scrophulariaceae in part)— Plantain Famii

Generally herbaceous perennials, probably short-lived, also flowering in first :

t, February to Apri

Known from the flora area by a single plant and therefore not collected; it was not a perennial.

Tinajas Altas, wash just below the lower tank, rare, 10 Jan 2002, Felger (observation).

4.E. Jones var. pseudospectabilis. Mojave beard-tongue

is perennials, also flowering in the first year. Flowers bright rose-purple; observed in fl

Canyons and at higher elevations in the mountains, generally not common.

Tinajas Altas, frequent in wash above tanks, 487 m, Hodgson 2723 (DES). Canyon above Tinaja*

and lower N-facing slopes among rocks, Felger 04-77. Frontera Canyon, 18 Mar 1998, Felger (

tPenstemon sp.

tTinajas Altas, fruits, 10,950 & 11,040 ybp.

Plantago ovata Forssk. var. fastigiata (E. Morris) S.C. Meyers & Liston [P. insularis Eastw. NotP. insularis Nyman

' ex Briq. P. fastigiata E. Morris. P insularis var. fastigiata (E. Morris) Jeps.] Desert woolly plantain, Indian wheat;

pastora; mumsa

Cool-season annuals; highly variable in size, flowering stems often 5-20 cm tall. Flowers small, straw-colored and

Papery. Seeds mucilaginous when wet and clinging tenaciously when dry.

Abundant and widespread, especially on sandy soils; washes, valley plains and bajadas, canyons, and soil pockets on

rocky slopes. The seeds were significant food and medicinal resources. It has been in the flora area for more than

10,000-year-old fossils from the Homaday Mountains in the Pinacate region.

Nought-stressed plants can be as small 6.4-12 mm tall, with persistent cotyledons to 12 mm long, only 2 leaves, each 7

long, and a peduncle bearing a single fruit.

present-day population is the inland North American variety (Meyers & Liston 2008). The infraspecific status of

the fossils is not known.

Coyote Water, Felger 05-123. Cipriano Pass, Reeves R5448 (ASU). Tinajas Altas, Felger 10-193. Observations: Canyon

above Tinajas Altas, 19 Mar 1998, Felger, Frontera Canyon, 18 Nov 1998, Felger. tButler Mts, seeds, 740-11,250 ybp

(6 samples). tOrgan Pipe Monument: Ajo Mts, Montezuma’s Head, seeds, 20,490 ybp.

Pseud °rontium cyathiferum (Benth.) Rothm. [Antirrhinum cyathiferum Benth.] Desert snapdragon

N ° n- a easonal annuals, viscid glandular-hairy (sticky) and foul smelling. Flowers small and dark purple.

Known in the flora area from a single locality but probably more widespread.

Anion g rocks in the bottom of Frontera Canyon, 18 Mar 1998, Felger (observation).

236

e northern margin of the flora area.

3 fall with daytime heat. The small flowers and small anthers point to a selfing

NE end of Tinajas Alias Pass, Van Devender 6 Mar 1983. Coyote Water, 18 Mar 1998, Felger (observation).

Gilia stellata A. Heller. Star giua

Basal rosette leaves well developed, stem leaves greatly reduced. Flowers white tinged with violet.

Common and widespread across the flora area; washes, bajadas, hills, and mountains.

Tinajas Alias Pass, Van Devender 5 Mar 1983. Tinajas Alias, Felger 10-189. Coyote Water, Felger 05-146.

Langloisia setosissima (Torr. & A. Gray) Greene subsp. setosissima. Bristly langloisia

Plants bristly and compact, 2-5 cm tall. Corollas bright lavender-pink; anthers and pollen white or blue.

Sandy to rocky soils; washes, valley plains and bajadas, hills, and mountains.

E side Tinajas Alias Pass, Felger 98-102.

Linanthus jonesii (A. Gray) Greene, jones’ desert-turmpet

Stems wire-like, the plants usually taller than broad. This species c

shortly after dusk and powerfully sweet-fragrant, sometimes nai

Washes, valley plains and bajadas, canyons, hills, and mountains at

Tinajas Alias, Van Devender 26 Mar 1983 (det. J. Mark Porter 2012). 1

presumably the same species).

trized by simple leaves, very coarse glandular

nail fruit. Flowers white, nocturnal, opening

losing before sunrise.

:ra Canyon, 18 Mar 1998, Felger (observation,

►eseliastrum schottii (Torr.) Timbrook [Langloisia schottii (Torr.) Greene] Schott’s cauco

Low dunes near the west side of Tinajas Altas Mountains and perhaps elsewhere in the western part of the flora area.

Butler Mts, Van Devender 27 Mar 1 983.

POLYGONACEAE— Buckwheat Family

CHORIZANTHE— Spine-flower

Small spring annuals; first leaves in a basal rosette; flowers minute, white or yellow and barely protruding <*

Felger etal., Flora of Tinajas Altas, Arizona

hidden in firm, spinescent bracts. The larger leaves in a bas

early March. Flowering mostly February to March, the plani

full flower with onset of fruiting, the weather is wai

t breaking apart at

Chorizanthe brevicornu Torr. subsp. brevicomu. Brittle spineflower, short-torn spineflower

Cool-season annuals. Flowers white and minute. As the plants mature and dry in April and early May, they completely

break apart, the inflorescence branches breaking into small seed-bearing segments that have tiny grappling-hook

Sandy, gravelly, and rocky soils; washes, canyons, upper bajadas, pediments, and lower mountain slopes. It was in the

flora area 10,400 years ago.

Tinajas Altas, Van Devender 5 Mar 1983. Steep canyon slope N of Tinajas Altas, 19 Mar 1998, Felger (observation). tButler

Mts, fruit-bearing segments (with hooks), 10,360 ybp.

Widespread; bajadas and valley plains, washes, pediments, and lowe

8200 years.

Camino del Diablo SE of Raven Butte, Felger 05-33. NNE of parking ai

SW side of Tinajas Altas Mts, 10 Jan 2002, Felger, observation. tBi

s been in the flora for at It

ERIOGONUM— Wild buckwheat

Eriogonum fasciculatum Benth. var. polifolium (Benth.) Torr. & A. Gray. Flat-top buckwheat

Small shrubs; flowers whitish or pink; February to May.

U Pper bajadas and washes, canyons, and rocky slopes in the mountains, especially at higher elevations in canyoi

east- and north-facing rocky slopes. The history of this variety in the flora area extends to 11,100 years, an

species has been in the area at least 18,700 years.

Tinajas Altas, Shreve5942. Tinajas Altas Mts, NE of upper tanks, Felger 04-74. tButler Mts, leaves

samples). tVariety unknown: Tinajas Altas, leaves, 5080-18,700 ybp (7 samples).

Erio S on «*m inflatum Torr. & Frem. Desert trumpet, bladder stem

Perennial herbs with a hard, knotty base; leaves basal, the flowering stems leafless, the first internt

the u PPer part inflated (swollen and hollow) or not without apparent pattern. Flowers small J

spring and again with the summer rains.

Uanyons, rocky hills and mountain slopes. It grew in the Butler Mountains 8200 years ago.

s, 8160-11,060 ybp (3

238

Canyon below Raven Butte Tank, Felger 10-226. Cipriano Pass, N end of Tinajas Alias Mts, Reeves 5441 (ASU). Observe

lions: Canyon above Tinajas Alias, 19 Mar 1998, Felger, Granitic hills at SW side of Tinajas Alias range, 10 Jan 2002,

Felger. tButler Mts, fruit with calyx, 8160 ybp.

Eriogonum thomasii Torr. Thomas’ wild buckwheat

Small, delicate, cool-season annuals. Flowers pink and yellow; flowering branches and involucres glabrous, the outer

Washes, valley plains and bajadas, and canyons.

Coyote Water, Felger 05-153. Vicinity of Tinajas Altas, 1900 ft, Van Devender 5 Mai

Tank, Felger 10-227.

Eriogonum thurberi Torr. Thurber’s \

Small, delicate, cool-season annuals.

outer tepals as wide or wider than long, broadest towards the tip, and i

E edge of Davis Plain, west branch of Camino del Diablo, Felger 05-93.

Eriogonum trichopes Torr. Little desert trumpet

Cool-season annuals; highly variable in size. Flowers yellow.

Sandy, gravelly soils; valley plains, bajadas, and washes.

Camino del Diablo, SE of Raven Butte, Felger 05-28. Coyote Wash at Camino del Diablo, 10 Jan 2002, Felger (observa-

Eriogonum wrightii Torr. ex Benth. var. nodosum (Small) Reveal [E. wrightii var. pringlei (J.M. Coult. & Fisher)

Reveal] Bastard sage

Small, scruffy subshrubs with whitish stems and small, quickly drought-deciduous leaves. Flowers white or pink; flow-

ering response apparently non-seasonal, or perhaps not during summer.

Mountains, especially north-facing slopes and canyons. This species has been in the flora area for more than 43,000 years.

Tinajas Altas, 5 Dec 1935, Goodding 1449. Frontera Canyon, 18 Mar 1998, Felger (observation). tVariety unknown.

Tinajas Altas, leaves, 5860-15,680 (14 samples), & >43,000 ybp.

x>l-season annuals with very slender, delicate

green or white and pink woolly bracts.

.mes and sand flats west of the Tinajas Altas V

itler Mts, Van Devender 27 Mar 1983.

PORTULACACEAE— Purslane Family

Portulaca halimoides L. [P. parvula A. Gray] Silk-cotton purslane

Summer annuals, the plants small and succulent. Sepals reddish and relatively persistent; petals, anthers, and stign»

golden yellow but these are only evident while the flowers are open— generally from a few hours after sunrise until

Coyote Water, Felger 04-61.

RANUNCULACEAE— Ranunculus Family

Inemone tuberosa Rydb. Desert windflower

Small, herbaceous root perennials; growing and flowering in spring.

It was at Tinajas Altas nearly 10,000 years ago. The nearest present-day population occurs in Arizona Upland in the/

Mountains, especially at higher elevations. It is generally not known from the Lower Colorado Desert region, w

the exception of a few records from the Whipple Mountains (De Groot 2007).

TTinajas Altas, seeds, 9900 ybp.

RESEDACEAE — Mignonette Family

O ligomeris tinifolia (Vahl) J.F. Macbr. Desert cambess

Annuals, recorded in the region from October through May. Flowers minute and inconspicuous, white and green.

239

Journal of the Botanical Research Institute of Texas 6(1)

Coyote Water, on Prosopis glandulosa, Felger 04-59. Observations: Camino del Diablo near Coyote Wash, on Asclqrn

albicans, 29 Dec 2001, Felger; Tinajas Alias, on Olneya, 19 Mar 1998, Felger. tButler Mts, fruits, 740-8160 ybp (3

SCROPHULARIACEAE, see OROBANCHACEAE and PLANTAGINACEAE

SOLANACEAE — Nightshade Or Potato Family

Annuals or perennials includin

bases (resembling a woolly pad

4. Fruiting calyx loosely growing around the berry like a paper bag; corolla:

Chamaesaracha coronopus (Dunal) A. Gray. False nightshade

Herbaceous perennials from deeply buried roots, the stems often parti

Locally common along Coyote Wash.

Coyote Water, Felger 04-35. Coyote Wash at Camino del Diablo, Felger (

a discolor Bemh. Desert datura; towache

i-seasonal annuals but frost sensitive and responding poorly to cooler weather; hig

occasionally 1 m tall. Herbage stinky. Flowers usually 8-17 cm long, white with a ]

nal and fragrant. Fruits globose, spiny capsules, turning down at maturity,

lely scattered across the flora area; washes, bajadas and valley plains, and canyon 1

est of any plant in the region, but are comparatively small within the genus,

nino del Diablo, SE of Raven Butte, Felger 04-20. Coyote Wash at Camino del Di

Tinajas Alias, bajada, 19 Mar 1998, Felger ; Frontera Canyon, 18 Mar 1998, Felger.

>, Felger 04-64. Observations:

LYCIUM — Wolfberry; sauoeso

The four species in the region are hardwood shrubs often 1-2 m tall and armed with spines or thorn-tipped

twigs. Leaves semi-succulent, the margins entire.

Lycium fremontii is morphologically gynodioecious: different plants have (1) smaller flowers that are

male-sterile, with well-developed female parts and reduced or sterile stamens and anthers — these plants pro-

duce fruit, or (2) larger flowers that appear bisexual with well-developed male and female parts, however, these

plants may be largely or functionally male and have substantially reduced or no fruit production (Miller &

Venable 2002). The other three species in the flora area produce bisexual flowers. The fruits are edible (except

L. macrodon ), although only L. fremontii seems to have been a significant food plant (Felger 2007; Felger &

Moser 1985; Hodgson 2001).

Lycium andersonii A. Gray var. andersonii [Landersonii var. deserticola (C.L. Hitchc.) Jeps.] Anderson’s wotfBi**'

Spinescent shrubs to 2+ m tall. Leaves slender and glabrous. Flowers small, slender, and lavender; non-seasonaL Fruits

orange, fleshy, and edible.

Canyons and washes along the east side of the Tinajas Altas Mountains (Fig. 15).

Arroyo 2 mi SE of Tinajas Altas, Felger 08-196. Borrego Canyon, 3 Feb 1990, Felger (observation).

Lycium fremontii A. Gray var. fremontii. FrSmont wolfberry; saucieso; kuavutJ

Readily recognized by the glandular hairs on the herbage, pedicels, and calyx; long pedicels; dimorphic flowers (male

and female flowers on separate plants); and relatively large, soft, red-orange, fleshy, and edible fruits. Flowers laven-

der. The wet foliage smells like a wet dog.

Widely scattered and often locally common along washes.

Coyote Water, Felger 04-48. Arroyo 2 mi SE of Tinajas Altas, Felger 08-197. Below Tinajas Altas, wash, McLaughlin 1967.

Camino del Diablo, E of Raven Butte, 29 Nov 2001, Felger (observation).

Lycium macrodon A. Gray. Desert wolfberry; s-cuk kuavulI

Long-shoot nodes often bearing stout spines; leaves often moderately glaucous and sometimes unusually large on long

shoots. Flowers white and green, February to April. Fruits hard, glaucous, and constricted (appearing pinched)

below the middle.

Washes, sand flats, alluvial flats, and sometimes on north-facing slopes at lower elevations. It grew in the Butler Moun-

tains 8200 years ago but is not found there today. The fossil fruits are hard and bony.

Baseof Tinajas Altas Mts, Harrison & Kearney 6571. Tinajas Altas, Gila Mountains, 6 Mar 1937, Harbison 16816. Camino

del Diablo at Coyote Wash, Felger 10-164. tButler Mts, twigs, leaves, fruits, 8160 ybp.

Lycium parishii A. Gray. Parish wolfberry; saucieso

Readily recognized by the markedly glandular-pubescent herbage, pedicels, and sepals, pale gray-green foliage, and pale

lavender flowers with highly variable calyx lobes often much longer than the calyx tube. Corollas lavender; flower-

ing response non-seasonal after rains. Fruits orange, fleshy, and marginally edible.

This is the most common and widespread lycium in the flora area. Washes, flats, canyons, and rocky slopes.

Coyote Water, abundant, Felger 04-49. Tinajas Altas, Van Devender 9 Mar 1980. Tinajas Altas Pass, Reeves R 5369 (ASU).

Observations: Granitic hills at SW side of Tinajas Altas range, 10 Jan 2002, Felger, Frontera Canyon, 18 Mar 1998,

tLyciumsp./spp.

These fossil samples may represent more than one species. These seeds ai

of various lycium species, but not L. macrodon.

trmajas Altas, seeds, 4010-10,950 ybp (4 samples).

te flattish, oval, and orange-brown typical

Glandular-pubescent annual or perennial herbs; fruits of capsules with n

} ; mostly sandy to gravelly soils of washes, bajadas, and desert

:s 10913. Coyote Water, Felger 05-127. Tinajas Altas Pass, Reeves R5407 (ASU).

. Martens & Galeotti [Njrigonophylla Dunal] Coyote tobacco, desert tobacco; i

Perennial herbs or subshrubs, and

Easlsid e of Tinajas Altas Mountaii

sifalia Benth. var. versicolor (Rydb.) Waterf.l Desert ground-cherry; tomaitud d

flowering in the first season. Flowers dull yellow, at vanous seasons.

242

Tinajas Altas, Goodding 4902. Vicinity of Tinajas Altas, Van Devender 5 March 1983. Wash, 2 mi SE of Tinajas Alias, F ti-

ger 08-206. Frontera Canyon, 18 Mar 1998, Felger (observation).

tPhysalis sp./spp.

Physalis has been a member of the local flora for more than 43,000 years. The seeds are diagnostic only to genus.

tButler Mts, seeds, 740-11,250 ybp (5 samples). tTinajas Altas, seeds, 1230 to > 43,000 ybp (20 samples).

tSolanum hindsianum Benth. [or possibly S. elaeagnifolium Cav.] Hu

Sparsely branched shrubs, variously spiny or not. Flowers showy and

The nearest present-day populations are in mountains in nearby nortl

al. 2007b).

'JDS nightshade; mala muter

lavender.

iwestem Sonora and Organ Pipe Monument. Ik

nined by freezing weather (Felger 2000; Felger et

nightshade was in Tinajas Altas 11,000 years ago and in the Aji

tTinajas Altas, seeds, 10,950 ybp. tOrgan Pipe Monument: Ajo Me

day flora area but it is a com-

ither flat, lens-shaped seeds

ompare well with the fossil seeds. The fossil

ins 20,500 years ago.

Montezuma’s Head, seeds, 20,490 ybp.

URTICACEAE— Nettle Family

Parietaria hespera B.D. Hinton var. hespera. Desert pelutory

Delicate cool-season annuals. Flowers minute and green.

Seasonally common along Coyote Wash and in the mountains, especially in protected niches among rocks and beneath

Coyote Water, Felger 04-55. Tinajas Altas, Felger 10-192. Canyon below Raven Butte Tank, Felger 10-236.

VERBENACEAE— Verbena Family

tGLANDULARIA sp ,/spp.

Non-seasonal annuals or herbaceous perennials, present at Tinajas Altas 8700 years ago. The nearest present-

day occurrences of the genus are G. bipinnatifida (Nutt.) Nutt, and G. gooddingii (Briq.) Solbrig in Organ Pipe

Monument, and the latter is also in the eastern margin of Cabeza Prieta Refuge.

tTinajas Altas, nutlets, 8660 ybp.

VISCACEAE, see SANTALACEAE

ZYGOPHYLLACEAE— Caltrop Family

FAGON1A

Low-growing, spinescent perennials and also flowering in the first season. Leaves 3-foliolate. Flowers laven-

der-pink; flowering with warm weather at various seasons except during extended drought. No other genus is

so wide ranging yet so closely restricted to the hot, arid deserts of the world. The seeds, which become muci-

laginous when wet and adhere tenaciously upon drying, are probably a major factor in the unique and widely

disjunct distributions (Bray 1898).

Fagonia laevis Standi. [F. califomica subsp. laevis (Standi.) Wiggins] Fig. 20.

Herbage glabrous or essentially so; spines short and moderately curved.

Rocky, arid slopes, canyons, and upper bajadas; not seen on open desert flats.

Fdgeretal., Flora ofTinajas Altas, Arizona

243

FUO. Fagonialaevis. Tinajas Altas Canyon just above the tinajas. 28 April 2010. Photo by JM.

Borrego Canyon, Felger 93-195. Surveyors Canyon, Felger 10-202. Canyon below Raven Butte Tank, Felger 10-230.

Fagonia pachyacantha Rydb. [F. calif omica var. glutinosa Vail]

Arid, rocky slopes and canyons of hills and mountains, and upper bajadas.

Tinajas Altas Mts, 1200 ft, Lindquist 25 Mar 1983.

KaUstroemia californica (S. Watson) Vail. Baiburin, maldeojo

Summer annuals. Stems generally trailing; leaves pinnate. Flowers small, yellow to yellow-orange.

Documented from washes and sand flats along Coyote Wash and expected elsewhere.

Coyote Water, Felger 04-47.

divaricata Cav. subsp. tridentata (Sesse & Moc, ex DC.) Felger & C.H. Lowe [L tridentata (Sesse & Moc.

“DC.) Coville] Creosotebush; hediondiua, gobernadora; segai, segoi

Cong-lived shrubs to 2 + m tall with very hard wood and gummy, resinous herbage. Flowers yeUow, produced at various

it 1570 ft. Creosotebush has been a

dicinal plant in the Sonoran Desert.

*** the Sonoran Desert, Lama was restricted to elevations below 1000 ft in the last glacial pern

Probably many earlier glacial periods in the Pleistocene. The 18,700 ybp sample tram Tinajas Altas is the oldest

■“ocarbon dale tor Larrea in Norrh America. This is a landem acceler.ro, radiocarbon dam taker , direcrly on lo

■«es and leaves (Van Devender 1990). Kim Hunter relared smmaul cell sire to ploidy level, enabling her ,o de

244

hat early date (Hunter et al. 2001). The Mojave Desert hexaploids were only found in Holocene sam-

1 as they migrated out of the Lower Colorado River Valley. Considering that 18,700 ybp was

the Wisconsin full glacial, it is likely that creosotebush desertscrub was in the Lower Colorado Valley for much of

the Pleistocene — after creosotebush immigrated from South America to the Chihuahuan Desert at some unknown

tmino del Diablo, SE of Raven Butte, Felger 04-17. Mesa E of Tinajas Altas, Felger 08-187. Canyon above Tinajas Altas,

rocky bench just above arroyo bottom, 1570 ft (not seen at higher elevations), 26 Oct 2004, Felger (observation).

tButler Mts, twigs, leaves, fruits, 740-11,250 ybp (7 samples). tTinajas Altas, twigs, leaves, fruits, 4010-18,700 ybp.

MONOCOTYLEDONS

Agave deserti Engelm. subsp. simplex Gentry. Desert agave; lechugwia, mezcal; ’a’ud

Medium-sized agaves, solitary or forming small colonies from offsets. Leaves thick, dull colored, and with margins

spines. Flowers yellow, mostly April to June; fruits ripening June and July.

Widely scattered in the mountains, and occasional on upper bajadas on the eastern sides of the mountains. These agave

served as major food resources and the leaves yielded fiber. ^

Holocene. It is well represented in the fossil record from Tinajas Altas for than more than 18,700 years. Most of th

fossil agave specimens were identified by Tony Burgess. His identifications were based in part on examination a»

n Camino del Diablo and Borrego Canyon, Felger 90-19. Tinajas Altas, Goldman 2310 (US, image seen); Van Do-

er 86-142. tTinajas Altas, leaf fragments (epidermis), prickles, seeds, 1230-18,700 ybp (17 samples).

A. Gray. Ajo uly, desert uly; ajo silvestre; a:sos

easons from a single and rather large corm (“bulb”). Bowers white, 6-8 cm wide

n or early evening, and partially closing with daytime heat of the next morning

izes, in drier years the flower stalks may scarcely be visible but in wet years tney

may be more than 1.5 m tall, and in the driest years the plants remain as dormant bulbs.

Mostly on deep, sandy soils of the Lechuguilla Valley, plains and bajadas, and sometimes on rocky pediments. The rela-

tively large corns were baked or boiled and also eaten fresh, probably in spring (Castetter & Bell 1951; Felger 2007;

Hodgson 2001), but are rather slimy when fresh (also see Rea 1997).

Coyote Water, Felger 05-119.

Growing and flowering during cool i

Flowering stalks highly variable

245

Hesperoyucca whipplei (Torr.) Baker ex. Trel. [Yucca whipplei Torr. Y. newberryi McKelvey] Spanish bayonet

A gave-like rosettes more than 1 m wide, the plants dying after flowering.

This species was widespread across the Sonoran Desert in early Ice Age times (Van Devender 1990) and is docum<

for Tinajas Altas from 1 1 ,000 to more than 43,000 years ago. It is missing from most Sonoran Desert middle am

Holocene midden samples, indicating that the main range reduction was due to drying climatic conditions, bi

food resource: The hearts and the young emerging flower stalks were pit-baked like those of agaves, and the flc

and seeds were also eaten (Felger 2007). The harvesting methods preclude reproduction of the plants. A small

tual population occurs in the Sierra del Viejo (immediately south of Mexico Highway 2 in adjacent Sonora son

the Tinajas Altas Mountains, a place where there is no freshwater source (Felger 2000). It is otherwise absent a<

the final demise of this plant across most of the Sonoran Desert?

tTinajas Altas, leaf fragments, 11,040, 18,700, & >43,000 ybp.

Nolina bigelovii (Torr.) S. Watson. Desert tree-beargrass, Bigelow beargrass. Fig. 21A, B.

Yucca-like plants developing a woody trunk and large, tall flowering stalks. Flowers small, white and green, in June, and

The young flower stalks were pit-baked by the Cahuillas (Bean & Saubel 1972). The fibrous leaves of other Nolina spe-

cies were used for basketry and other practical uses, and the seeds prepared for food, but evidence is lacking for such

use for N. bigelovii (Felger 2007). Nolina bigelovii has been in the Tinajas Altas Mountains for more than 43,000 years.

Spectacularly tall nolinas occur along the bottom of Frontera Canyon (Fig. 21 A). Some of these giants have trunks 4-5

m tall, and one fallen, dead nolina had a trunk slightly more than 6 m long (measured 18 Mar 1998). Most of these

giants appeared to be declining and had leaves substantially shorter than smaller, “normal-sized” plants. Scattered,

few nolinas, mostly in the canyon and higher slopes above the Tinajas Altas tanks, have trunks 3-4+ m tall and may

Tinajas Altas, Shreve 6233. Tinajas Mts, major tinajas, 390 m, Hodgson 6975 (DES).

tTinajas Altas, leaf fragments, 4010-18,700 (16 samples), & >43,000 ybp.

Triteleiopsis palmeri (S. Watson) Hoover [Brodiaea palmeri S. Watson] Blue sand-lily

Growing and flowering during cooler seasons from a cluster of small cormlets (bulblets). Flowers attractive, deep blue.

Sand flats and dunes west of the Tinajas Altas Mountains and widespread westward to Yuma and in the nearby western

Part of Cabeza Prieta Refuge and the Gran Desierto in adjacent Sonora.

The cormlets are tasty, eaten fresh or cooked (Felger 2007; Felger & Moser 1985).

Butler Mts, Van Devender 27 Mar 1983. Near Border Monument 198, Morrison 73 (ASU).

Wucca brevifolia Engelm. var. brevifolia and/or var.jaegeriana McKelvey [Y.jaegeriana (McKelvey) L.W. Lenz]

Joshua tree

■Hus unique tree, emblematic of the Mohave Desert, was common in Organ Pipe Monument and the flora area from at

least 43,000 to about 11,000 years ago. The nearest present-day occurrences of this species are in west-central and

TTinajas Altas, leaf fragments, 11,040, 18,700, & >43,000 ybp (abundant in these samples).

CYPERACEAE— Sedge Family

C yperns sqnarrosus L. [C. aristatus Rottb. Mariscus squarrosus (L.) C.B. Clarke] Dwarf sedge

Found along Coyote Wash following rains that resulted in exceptionalflow t

200 °)- Bird-dispersal is implied since the nearest populations are not cor

Coyote Water, Felger 04-41 Cabeza Prieta Refuge: Las Playas, Monson 9.

(CAB).

through the wash. The nearest known popu-

mected to Coyote Wash by surface flow.

Cabeza Prieta Tanks, Monson 25 Sep 1955

246

Journal of the Botanical Research Institute of Texas 6(1)

the Tinajas Altas Mountains above the tinajas, 25 October 2009. Photos by RSF.

NOLINACEAE, see ASPARAGACEAE

POACEAE — Grass Family

Grasses are the second largest family in the flora. The 18 species in the present-day flora represent 8% of the

total flora, as compared to the 14% of the flora of adjacent northwestern Sonora (Felger 2000). Three additional

species are known from the flora area only by fossils. Eight present-day grass species in the flora are annua

(ephemerals) and 7 are tufted perennials, although some of the perennials may become reproductive in the first

season. Four species are native to the Old World.

Thirteen grass species have been found in the fossil packrat middens. Three of them, Bouteloua repens,

Festuca microstachys, and Setaria macrostachya, are no longer present in the flora area and today are found in

less harsh areas to east of the flora area. Identifiable fragments of the fossils include fragments of inflorescences

and reproductive structures that may include spikelets or even spikes or spikelet clusters, florets, caryopses

(“grain”), or combinations or pieces of these parts.

-sided spikes (spicate branches), the spikes 2 or

Felgeretal., Flora ofTinajas Altas, Arizona

247

s; spikes symmetrical or laterally compressed, but r

). Spikelets usually 3-awned; without cl

ARISTIDA— Thrffawn; zacate tres b areas

Annuals and perennials. Spikelets 1-flowered, readily breaking off above the glumes (often lodging in socks —

people might be supplanting animals as dispersal agents). Lemma hard at maturity, slender and terete, narrow-

he awns reduced or absent.

iual Aristida in the So-

noran Desert. It has been in the region for more than 10,650 years.

Tinajas Altas, Felger 04-69. Tinajas Altas Pass, Reeves R 5416 9 (ASU). Camino del Diablo at Coyote Wash, F

tButler Mts, 8160-10,615 ybp. tTinajas Altas Mts, 5080-10,070 ybp (3 samples; Van Devender et al. 19<

Aristida purpurea Nutt. var. nealleyi (Vasey) Allred [A. glauca (Nees) Walp.1 Purple threeawn; tres a

Med perennials; flowering response non-seasonal.

Tinajas Altas and Butler Mountains. The history of this variety extends to 9700 years in the Tinajas Altas Mo

this or a similar species was in the Buder Mountains 11,300 years ago.

Butler Mts, Von Devender 27 Mar 1983. Tinajas Altas, Vorhies 16 Apr 1924. tBuder Mts, 6490 ybp (Van De

1990b: 342). tTinajas Altas Mts, 4010 & 9700 ybp (Van Devender et al. 1990b: 341). tA. cf. purpurea

6490 to 11,250 ybp (3 samples, Van Devender et al. 1990b: 342).

Cochlea barbinodis (Lag.) Herter [Andropogon barbinodis Lag.) Cane bluestem; zacate popamix)

Robust tufted perennials; growing and reproductive during warm weather.

°ue small population was seen in Frontera Canyon and other small populations are known from a few seal

holes in Cabeza Prieta Refuge and the Gila N

■ w,.

^TUOUA-Grak,,

ltle Ptesent-day grama

i, 19 Mar 1998, Felger (observation).

a (most arid portion) of the Sonoran Desert are sun

248

often short-lived and with weakly developed root systems, and the basal leaves sparse and the internodes be-

tween them readily visible.

Bouteloua aristidoides (Kunth) Griseb. [B. aristidoidesvar. arizonica M.E. Jones] Six-weeks needle grama; aceotia,

Summer annuals, the roots often weakly developed.

the region for at least 11,300 years.

Coyote Water, Felger 04-31. Tinajas Alias, above the tinajas, 19 Mar 1998, Felger (observation). 150 m N of Raven Peak

(700 m SSW of Raven Butte), steep draw, Walter & Morrison 584 (ASU). tButler Mts, 610-11,250 ybp (6 samples; Van

Devender et al. 1990b: 342). tTinajas Alias Mts, 4010 & 8700 ybp (Van Devender et al. 1990b: 341).

Bouteloua barbata Lag. var. barbata. Six-weeks grama; navajtta, zacate uebrero; s-cuk mudadt-kam

Widespread from the Lechuguilla Valley and to at least 1700 ft in mountain canyons.

Coyote Water, Felger 04-32. Tinajas Alias, Felger 04-70.

tBouteloua barbata var. barbata &/or var. rothrockii (Vasey) Gould [B. rothrockii Vasey]

One or perhaps both varieties of this grama grass are represented in the fossil record from about 5000 and 8700 yeais

ago in the Tinajas Altas Mountains (in Organ Pipe Monument for at least 21,900 years). The two taxa are distin-

guished by growth form (annual for var. barbata, perennial for var. rothrockii), not by reproductive structures, and

cannot be distinguished by the fossils, which are reproductive fragments.

tTinajas Altas Mts, 5080 & 8700 ybp (Van Devender et al. 1990b: 341).

tBouteloua repens (Kunth) Scribn. & Mem [B.filiformis (E. Fount.) Griffiths] Slender grama; navajita delgam

Tufted perennials, generally growing with warm weather.

Documented for 8200 years ago in the Butler Mountains. The nearest present-day populations are in the Puerto Blanco

and Ajo Mountains and locally elsewhere in Arizona Upland portions of Organ Pipe Monument.

tButler Mts, 8160 ybp (Van Devender et al. 1990b: 342).

-. subsp. rubens (L.) Husn. [B. rubens L.] Red b:

12-26 mm long, stiff, stout, straight or slightly curved

This invasive grass, native to the Old World, has become abundant in southern Arizona and a

Sonora, and is thoroughly established in Organ Pipe Monument, Cabeza Prieta Refuge, a

produce dormant seeds and does not maintain a seed bank in the soil (Salo 2004).

Camino del Diablo at Coyote Wash, Felger 02-8. Tinajas Altas, Felger 05-483.

inal lobes, and an awn

ass the northern part of

lals, red brome does not

*Cenchrus ciliaris L. [Pennisetum ciliare (L.) Link] Buffelgrass; zacate buffel (

Buffelgrass is well established along Mexico Highway 2 across northwestern Sonora and has spread northward into

larger washes and canyons at the southern tip of the Tinajas Altas Mountains. As of 2010 this usually invasive spe-

cies had been seen in the Tinajas Altas region only along the international border.

Frontera Canyon, international border, not common, 18 Mar 1998, Felger (observation).

Dasyochloa pulchella (Kunth) Willd. ex Rydb. lErioneuronpulchellut

Hitchc.] Fluff grass; zacate borrecuero

Low, tufted perennials with slender, arching stolons. Flowering with w

cruitment apparently occurs with summer rains. Seedling? may grow

Tateoka. Tridens pulchdtus (Kund^

last of oki.di? or dead plants. “*‘ dl

Felgeretal., Flora ofTinajas Altas, Arizona

Widespread in dry, open sites on mountains, hills, and upper bajadas. This is one of the most arid-inhabiting p<

grasses in the Sonoran Desert. It has been in the region for more than 10,300 years.

amino del Diablo, SE of Raven Butte, Felger 04-11. Tinajas Altas, above upper tinajas, Felger 04-92. tButler Mi

11,250 ybp (9 samples; Van Devender et al. 1990b: 342). tTinajas Altas Mts, 4010-10,300 ybp (5 samples; V

Digitaria califomica (Benth.)

ifomica iTrichachne califomica (Benth.) Chase] Arizona cottontop;

Tufted perennials; panicles narrow and densely flowered, the spikelets silky-cottony.

Only two plants of this grass were seen in the flora area, but perhaps it occurs at higher elevations in the me

Tinajas Altas, one plant seen among rocks at base of tinaja slopes, 29 Dec 2005, Felger (observation); 28 Mar

10-185 (one panicle collected).

Coyote Water, Felger 04-21.

2010. Fdga

tFestuca microstachys Nutt. I Vulpia microstachys (Nutt.) Munro] Small fescue

Cool-season annuals. Documented in the flora area from 11,000 years ago. The nearest present-day populations are in

the Ajo Mountains in Organ Pipe Monument. Four varieties are sometimes recognized, but they are not geographi-

cally well marked and do not seem worthy of recognition (Felger 2007b; var. ciliata A. Gray ex Beal is recorded for

the Ajo Mountains).

tTinajas Altas Mts, 10,950 ybp (Van Devender et al. 1990b: 340).

Festuca octoflora Walter [F. octoflora var.

flowered fescue

hirtella Piper. Vulpia octoflora (Walter) RydbJ Six-weeks fescue, eight-

in size, 2.5-15+ cm tall.

tains to at least mid-elevations. It has been in the flora area for at least 11,300

Three varieties are sometimes recognized but are of doubtful significance (Felger 2000; Felger et al. 2007b). Present-day

specimens are var. hirtella, characterized in part by pubescent spikelets (lemmas).

Tinajas Altas, 1450 ft, Van Devender 10 Mar 1980. Tinajas Altas, near lowermost tinaja, Felger 10-186. Coyote Water,

Felger 05-136. tButler Mts, 11,250 ybp (Van Devender et al. 1990b: 342). F. cf. octoflora: TTmajas Altas Mts, 8970-

10,950 ybp (4 samples; Van Devender et al. 1990b: 340).

x Roem. & Schult. [Andropogon contort

ist-colored and persistent. Spikelets beai

leteropogon contortus (L.) P. Be

Robust tufted perennials; dry lea

*5-7+ cm long. Reproductive during warmer months.

Canyon bottoms in the Tinajas Altas Mountains. Although some authors have claimed it is

it has been here at Tinajas Altas for at least 7900 years (Van Devender et al. 1990b).

Tinajas Altas Mts, 1200 ft, Van Devender 25 Mar 1983. tTinajas Altas Mts, 5860-7860ybp.

“^ngida (Thurb.) Benth. ex Scribn. [Pleuraphis rigida Thurb.] Big

..] Tanglehead; zacate Colorado

ftedpen

, during the wanner months. Large, inflated

ss valley plains and bajadas, and sometimes

rocky slopes. It h

Coyote Water, Fd^r M^^Su^eyors^a^on, Felger 10-201. Canyon above Tineas Altas, 19 Mar 1998, Felger (observa-

tion). tButler Mts, 3820-10,400 ybp (3 samples; Van Devender et al 1990b:

i Little-seed muhly; liendrilla chzca

stooped during cooler seasons. Spikelets 1-flowen

lizing and nor opening); terminal panicles branched a

rrmac (.rmnsprH with awns 14—28 mm long.

Fdgeretal., F

THEMIDACEAE, see ASPARAGACEAE

TYPHACEAE— Cattail Family

T ypha domingensis Pers. Southern cattail; tvle; ’uduvhag

Robust perennial herbs; winter dormant. Leaves upright, to 1.8 m long. The tiny, lightw

thrir associated hairs are produced in prodigious quantities and may be airborne o\

Cattails were seen at tinajas #4 (see Bryan 1925: 133) during the 1970s a

(Fig. 22). This is the only wetland plant species known from the Tii

terholes in nearby regions, including the Cabeza Prieta Refuge and i

Tinaja or pool #4 is a “shelf” Tinaja (about 25 ft perpendicular to flow), narrow (5 ft parallel to flow), nearly inacces

pool midway up the plunge-pool series and above, not beside, the large wedged boulder. It is filled with sedii

and Broyles has only once seen standing water in it, up to 60 cm deep, but the sediment undoubtedly holds moi:

for a long time. Typha has not been seen at any other pool, for they may be too frequently scoured to support wet

is, and a specimen was obtained in 2008

as Mountains. It occurs at scattered wa-

stern Sonora, and is abundant along the

plants

. A label for Cheilanthes parryi

Litas, “cattails at tanks,” 12 Feb

i in damp soil at the edge of the

DOUBTFUL, ENIGMATIC, AND POTENTIAL RECORDS

AMARANTHACEAE— Amaranth Family

Amaranthus palmeri S. Watson. Careless ml

Summer annuals; leaf blades mostly ovate. M

firm and often sharp; sepals not fringed; st

Strangely, there are no records for it in the flor

nearby localities.

C0NVOLVULACEAE— Morning Glory Fan

ages during the day.

: exception, the Ajo Mounta

Theej

gan Pipe Monument is the ir

:ollected by Andrew Alexant

Nic” Nichol on 25 April 1938 at the “South

iry to include the Tinajas Altas Mountains

lichol collected Muhlenbergta dumosa in the “Mohawk

The muhly grass is likewise an enigmatic and tantaliz-

a plausible place to support such plants, or perhaps

in the concept of the Gila Mountains. On tl

Mountains” and Aristida purpurea in the “Gi

ing “extralimital” collection. In addition to a

canyon in Spook Canyon in the Gila Moun

Nichors locality data is in error.

— Grass Family

California threeawn; ires barbas oe California

. nnials also flowering in the first season. Tliis grass, characteristic of sandy soils, should be sought in

Portions of the Lechuguilla Mountains and sand soils on the west side of the Tinajas Altas Mountain

on sand flats and dunes at the west side of Cabeza Prieta Refuge and ir

flora area (Felger 2000; Felger et al. 2007b).

o of Sonora just south of the

^ ^que grass has relatively large burs bearing sharp spines. Not known for tl

n the nearby Gran Desierto al

Fig. 22. Cattail ( Typha domingensis, Wilder 08-383) at Tinajas Altas, tinaja #4. 20 November 2008. Photo by Benjamin T. Wilder.

ACKNOWLEDGMENTS

Over the years many different friends and colleagues have provided information and assistance, or accompa-

nied us in the field. In this regard we thank Kelly W. Allred, A. Elizabeth (Betsy) Arnold, Daniel F. Austin,

Thomas Bowen, Brad Boyle, Steven M. Buckley, Tony L. Burgess, J. Travis Columbus, Mark A. Dimmitt, Luke

Taylor Evans, Mark Fishbein, Exequiel Ezcurra, Judy Ann Gibson, Edward Erik Gilbert, W. Eugene Hall ^

phen F. Hale, Gayle Harrison Hartmann, James L. Heard, James Henrickson, Wendy Caye Hodgson, PhfljP ’

Jenkins, Sarah Lynn Johnson, the late Cynthia A. Lindquist, Elizabeth Makings, Curtis McCasland, Michel

McMahon, J. Mark Porter, Jeff Pralle, Adrianne G. Rankin, Jon P. Rebman, the late Charlotte Reeder and Jo n

Reeder, Karen Reichhardt, Rusty Russell, Susan Rutman, Andrew Salywon, Andrew C. Sanders, Silke Schnd

der, Richard Spellenberg, Victor Steinmann, Larry Toolin, Sula Vanderplank, Benjamin T. Wilder, and George

Yatskievych. Cathy Moser Marlett drafted maps and Heidi Orcutt-Gachiri and Matt Schultz helped with an-

other map. Adrian Quijada-Mascarenas translated the abstract. Sarah J. De Groot, Philip D. Jenkins, and Fran-

cisco Molina-Freaner provided a careful review of an earlier draft of the manuscript.

The authors thank the United States Marine Corps for encouragement of research on the Gol wa

Range and generous support of this publication. Richard writes: 1 especially appreciate Bill Broyles introducing

253

me to this great place and for good times in the field. Portions of the earlier work as well as more recent efforts

were supported by the Wallace Research Foundation. Tom writes: I thank Paul S. Martin for stimulating my

interest in packrat middens and secrets of the Ice Ages. Bill applauds the inclusive community of professional

and amateur researchers who enthusiastically share their deep love and vast knowledge of the desert. Jim

Malusa thanks the generous support of the United States Marine Corps, made possible by the efforts of Ronald

L. Pearce, Abigail Rosenberg, Robert Palmer, Traci Allen, and Robert Lovich. Thanks, too, to Karen Reichhardt

of the Yuma Field Office, Bureau of Land Management, who first introduced Malusa to the wonders of the

western Goldwater Range.

Adams, D.K./

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258

BOOK REVIEW

Eric Mader, Matthew Shepherd, Mace Vaughan, Scott Black, and Gretchen LeBuhn. 2011. Attracting Native

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— Amanda Stone Norton, Ph D.,

Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, Texas 76107-3400, U.SA

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i. Bot Res. Inst Texas 6(1): 258. 2012

A FLORISTIC INVENTORY OF DISMAL KEY AND FAKAHATCHEE ISLAND-

TWO SHELL MOUNDS SITUATED WITHIN THE TEN THOUSAND ISLANDS

REGION IN THE GULF OF MEXICO (COLLIER COUNTY, FLORIDA)

GJ. Wilder

MJ. Barry

Naples Botanical Garden

4820 Bayshore Drive

Naples, Florida 341 12-7336, USA

gwilder@naplesgarden.org

Institute for Regional Conservatio

22601 Southwest 152nd Avenue#

Miami, Florida 33170, U.SA

barry@regionalconservation.org

RESUMEN

taxa of vascular

INTRODUCTION

This is the third of a series of papers focused on the flora of southwestern Florida (Wilder «

Wilder & Roche 2009). Herein, we present the results of a study of the native and nonnati

Pknts growing wild on Dismal Key and Fakahatchee Island.

Dismal Key and Fakahatchee Island are, simultaneously, islands and massive shell mounds situated 3.7

miles apart within the Ten Thousand Islands region in the Gulf of Mexico, in Collier Co., Honda (Figs. 1, 2).

The two islands are centered atN 25 0 53’27" and W 81°33'28", and at N25°52'15" and W81°29'25", respectively.

Hk ™ of each island is, approximately, 73 acres (Beriault et al. 2003). Beriault el al. (2003) ranked both is-

together with Chokoloskee Island and Russell Key, as “ . . . the four largest a

mound complexes in the Ten Thousand Islands.” On each of the two islands the max

20 feet (South Florida Water Management District 2010). ,

Prehistoric Indians constructed both islands from shells over a prolonged time period. The islands served

as platforms for habitation and, possibly, as waste dumps. Schwadron (2010) concluded, based on data

foot radio-carbon dating, that for large mounds ("shell work sites’) within the Ten Thousand Islands region,

construction seems to have occurred from about 1900 to 900 years before the present, with a possible

l** in construction at ca. 1600 to 1300 years before the present; however, parts of Dismal Key and Fa a-

takhee Island might be considerably older than that, according to studies of Horr’s Island (an island wit

Ten Thousand Islands region situated 5.6 miles northwest of Dismal Key). Based o

dating, Russo (1991) concluded that Indians occupied the v

““ween 3000 and 4000 years before the present, and to a lesser exte

atls were likely the last of a

n data derived from radio-

n of Horr’s Island primarily

r and later. The Calusa Indi-

housand Islands.

Non Indians

wicted the Indians and settled Dismal Key and Fakahatchee Island. Beria.

Teased): 2

260

Journal of the Botanical Research Institute of Texas 6(1)

262

et al. (2003) reported historic artifacts and refuse from Dismal Key dating from the 1870s onward, and they

indicated evidence for at least three distinct historic home sites on the Island. They also quoted from observa-

tions made by Hrdlicka, in 1918, that much “high ground” on Dismal Key was then under cultivation. Also,

until approximately 1980, a succession of hermits occupied a house on Dismal Key that was constructed, per-

haps, in the 1920s or 1930s (Seely 2010). Fakahatchee Island was reportedly settled as early as 1870 (Tebeau

1966) and was subjected to . . long and intensive use by historic white settlers” (Beriault et al. 2003). The in-

habitants farmed, raised livestock on, and fished near the Island. They constructed houses (including a school-

house), cisterns, and a small cemetery, and they left behind various artifacts. They departed from Fakahatchee

Island, probably during the early-to-mid-1950s (Beriault et al. 2003; Seely 2010).

Today the islands are uninhabited and are included within a group of overlapping preserves. Dismal Key

belongs both to the Ten Thousand Islands Aquatic Preserve and to the Ten Thousand Islands National Wildlife

Refuge. Fakahatchee Island is included within the Ten Thousand Islands Aquatic Preserve. The Ten Thousand

Islands Aquatic Preserve and Rookery Bay Aquatic Preserve, jointly, compose the Rookery Bay National Estua-

rine Research Reserve (Burch 1998).

We undertook the present study for three main reasons. First, we wished to compile a list of the plant spe-

cies present on both islands. There existed no thorough, previously published flora or list of voucher specimens

for either Island; however, Beriault et al. (2003) had produced a brief, undocumented list of species for each

island; Burch (1998) had published an inventory of the vascular plants of the Rookery Bay National Estuarine

Research Reserve; and, Barry (2009) had compiled unpublished lists of endangered, threatened, and exotic

species of the Ten Thousand Islands National Wildlife Refuge. Second, we wished to document the island

floras by vouchering inventoried species. Third, by implementing study immediately, rather than by delaying

until the future, we desired to preempt the continuing rise in sea level that is likely to alter, and to eventually

inundate, both islands.

Herein, besides addressing the three objectives, aforementioned, we briefly characterize the topography

of the islands and certain relicts of human habitation, we describe the main kinds of habitats present on each

island and we indicate the species that occupy each kind, and we compare Dismal Key and Fakahatchee Island,

floristically, to other areas.

METHODS

George Wilder visited Dismal Key 23 times, from May 8, 2010 through Jan. 7, 2012, and Fakahatchee Island 24

times, from Aug. 6, 2009 through Dec. 10, 2011. Beginning in the early 1990s, Michael Barry independently

visited the Islands, focusing greatest attention on them during 2005 through 2009. For each island, during our

entire study each month of the year was represented by one or more visits; however, in any year we did not al-

ways visit an island during any two successive months. George Wilder vouchered species with dried herbari-

um specimens that were deposited at the Herbarium of Southwestern Florida (SWF), housed at the Naples

Botanical Garden (Appendix 1). We characterize species as native, exotic, and endemic to Florida, according to

Wunderlin and Hansen (2011), with the exception of Carica papaya, where we follow Ward (2011) and con-

sider this species to be native. Mostly, our nomenclature follows Wunderlin and Hansen (2011); however, Ap-

pendix 1 (footnote 1) specifies nomenclatural differences between that work and the present paper.

RESULTS AND DISCUSSION

Topography and relicts of former habitation

Beriault et al. (2003) and Schwadron (2010) have provided detailed descriptions of these aspects. Presented

here is a brief account that reflects personal observations. Dismal Key and Fakahatchee Island exhibit low-ly-

ing, relatively flat substrate, but additional topographic features interrupt the flat areas. For example, each is-

land exhibits a prominent curved band of elevated substrate connecting two sides of the island; each band

consists of conjoined, short ridges oriented perpendicular to the band’s long axis. The islands are characterize

by additional ridges. Some ridges occur inland, but on Dismal Key there also exists a massive ridge (an app^'

263

ent seawall) situated just landward of the island’s southern margin (i.e., in a position facing open ocean; Fig. 1,

S); this seawall manifests a long extension that continues northeastward from the main body of the island.

Fakahatchee Island exhibits a less obvious seawall (Fig. 2, S). An isolated tall mound known as Youman’s

Mound occurs within the westernmost hammock on Fakahatchee Island. Also, regularly spaced trenches—

interpreted as the remnants of Indian canals — are oriented perpendicular to the shoreline, both along the

western portion of Dismal Key and along the eastern and southern parts of Fakahatchee Island (Figs. 1, 2, Ca).

A long trench extends northeastward across Fakahatchee Island. Mangrove vegetation dominates all trenches.