Journal of the Botanical Research Institute of Texas

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

History and Dedication

1962—Lloyd H. Shinners

(left), a member of the

Southern Methodist University

(SMU) faculty and a prolific

researcher and writer, published the first issues of Sida,

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

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

botany at SMU and director emeritus of BRIT,

inherited editorship and copyright.

1993—BRIT becomes publisher/copyright holder.

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

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Journal of the Botanical Research Institute of Texas is

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by the Botanical Research Institute of Texas.

VOLUME 9 NUMBER 2 30 NOV 2015

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Table of Contents

SYSTEMATICS

Botrychium michiganense sp. nov. (Ophioglossaceae), a new North American moonwort

Arthur V. Gilman, Donald R. Farrar, and Peter F. Zika 295

Notes on Eritrichium (Boraginaceae) in North America II

David F. Murray 311

Change in taxonomic rank for a Hexastylis (Aristolochiaceae) taxon of the

southeastern United States

Brian R. Keener and L.J. Davenport 317

Calathea galdamesiana (Marantaceae), a new endemic Panamanian species

Helen Kennedy and Rodolfo Flores 319

Casearia draganae, a new species of Samydaceae from western Colombia and Ecuador

Mac H. Alford 325

Irenodendron, a new genus of Samydaceae from South America

Mac H. Alford and Angela D. Dement 331

New species of Senegalia (Fabaceae) from South America

David S. Seigler and John E. Ebinger 335

Nautilocalyx rugosus (Gesneriaceae), a new species from the rlo Cenepa watershed

(Amazonas, Peru)

Rocio del P. Rojas G. and John L. Clark 345

Corrigendum: New combinations in Coryphantha and Escobaria (Cactaceae)

Root Gorelick 351

PALEOBOTANY

Pseudhaplocricus hexandrus gen. et sp. nov. (Commelinaceae) in Mid-Tertiary

Dominican amber

George O. Poinar, Jr. and Kenton L. Chambers 353

Comopellis presbya gen. et sp. nov. (Rhamnaceae) in Mid-Tertiary amber from the

Dominican Republic

Kenton L. Chambers and George O. Poinar, Jr. 361

Klaprothiopsis dyscrita gen. et sp. nov. (Loasaceae) in Mid-Tertiary Dominican amber

George O. Poinar, Jr., Maximilian Weigend, and Tilo Henning 369

Addendum: Prioria dominicana sp. nov. (Fabaceae: Caesalpinioideae), a fossil flower in

Mid-Tertiary Dominican amber

George O. Poinar, Jr. and Kenton L. Chambers 381

DEVELOPMENT AND STRUCTURE

Breeding system and sex ratio variation in mulberries (Morus, Moraceae)

Madhav P. Nepal, Carolyn J. Ferguson, and Mark H. Mayfield 383

BOTANICAL HISTORY

From Cro-Magnon to Krai: A history of botany in Alabama

LJ. Davenport 397

ECONOMIC BOTANY

Birch (Betula, Betulaceae) bark horns and similar instruments in Norway

Torbj0rn Alm 433

FLORISTICS, ECOLOGY, AND CONSERVATION

First collection of Miconia alainii (Melastomataceae: Miconieae) with flowers

Walter S. Judd, Teodoro Clase, and Lucas C. Majure 449

Asteraceae en El Durazno y cercanlas, norte de Tamazula, Durango (Mexico):

Riqueza, distribucion y endemismo

David Ramirez Noya y Yolanda Herrera Arrieta 453

Redescubrimiento de Desmodium angustifolium (Fabaceae) en El Salvador

Pablo GalAn 471

Arachis glabrata (Fabaceae) new to the flora of Louisiana, U.S.A.

Charles M. Allen 475

New co-occurrence of Schoenoplectiella hallii and S. saximontana (Cyperaceae) in Ohio (U.S.A.):

Conservation implications for both species

Paul M. McKenzie, Daniel W. Boone, Marian Smith, and Richard L. Gardner 477

A checklist of vascular plants at the Gulf-shoreline extent of coastal prairie in southeast Texas, U.S.A.

David J. Rosen, Siavash Zamirpour, and Andrew Sipocz 485

In Memoriam

In memoriam: Robert R. “Bob” Kowal (23 April 1939-3 August 2015)

Theodore S. Cochrane 493

Reviewers, Index 503

Book Reviews, Notices, and Announcements 310,316, 324,330,344,350, 360,368, 380, 382,

396,432,452,476

INDEX to new names and new combinations in J. Bot. Res. Inst. Texas 9(2), 2015

Botrychium michiganense W.H. Wagner ex A.V. Gilman, Farrar, & Zika, sp. nov.—300

Calathea galdamesiana H. Kenn. & R. Flores, sp. nov.—319

Casearia draganae M.H. Alford, sp. nov.—325

Comopellis K.L. Chambers & Poinar, gen. nov.—363

Comopellis presbya K.L. Chambers & Poinar, sp. nov.—364

Coryphantha sneedii var. orcuttii (Boed.) Gorelick, comb, et stat. nov.—351

Hexastylis harperi (Gaddy) B.R. Keener & L.J. Davenp., comb, etstat. nov.—317

Irenodendron M.H. Alford & Dement, gen. nov.—332

Irenodendron coriaceum (Spruce ex Benth.) M.H. Alford & Dement, comb. nov.—333

Irenodendron cupulatum (Spruce ex Benth.) M.H. Alford & Dement, comb. nov.—332

Irenodendron ovalifolium (Macbride) M.H. Alford & Dement, comb. nov.—333

Klaprothiopsis Poinar, Weigend, & Henning, gen. nov.—370

Klaprothiopsis dyscrita Poinar, Weigend, & Henning, sp. nov.—370

Nautilocalyx rugosus R. Rojas &J.L. Clark, sp. nov.—345

Pseudhaplocricus Poinar & K.L. Chambers, gen. nov.—354

Pseudhaplocricus hexandrus Poinar & K.L. Chambers, sp. nov.—354

Senegalia duartei Seigler & Ebinger, sp. nov.—336

Senegalia noblickii Seigler & Ebinger, sp. nov.—338

Senegalia phillippei Seigler & Ebinger, sp. nov.—340

BOTRYCHIUM MICHIGANENSE SP. NOV. (OPHIOGLOSSACEAE),

A NEW NORTH AMERICAN MOONWORT

Arthur V. Gilman Donald R. Farrar

Gilman & Briggs Environmental Department of Ecology, Evolution and Organismal Biology

7 Conti Circle, Suite 5 Iowa State University

Barre, Vermont 05641, U.S.A. Ames, Iowa 500 7 7, US.A.

avgilman@together.net

Peter F. Zika

WTU Herbarium

Box 355325, University of Washington

Seattle, Washington 98195, U.S.A.

ABSTRACT

Botrychium michiganense W.H. Wagner ex A.V. Gilman, Farrar & Zika is described as a new moonwort species. It is an allotetraploid, most

likely descended from hybrids between ancestral B. lanceolatum s.l. and B. pallidum. It has a pinnate to pinnate-pinnatifid trophophore (ster¬

ile segment) and is most similar to B. hesperium and B. matricariifolium. These three species can be distinguished by trophophore shape and

dissection and B. michiganense is genetically distinct from the others as evidenced by different allozyme profiles. Botrychium michiganense

ranges from New Brunswick, the St. Tawrence region of Quebec, the Great Takes region of the US and Canada, the Black Hills of South

Dakota, and west through the Rocky Mountains to eastern British Columbia and Washington.

RESUME

Botrychium michiganense W.H. Wagner ex A.V. Gilman, Farrar & Zika est decrit comme une nouvelle espece de botryche. 11 s’agit Tun

allotetraploide issu de croisements entre une lignee ancestrale probable du B. lanceolatum s.l. et le B. pallidum. 11 possede un trophophore

(segment sterile) penne a penne-pennatifide et ressemble beaucoup au B. hesperium et au B. matricariifolium. On peut le reconnaitre par la

forme et la decoupure de son trophophore ainsi que par des caracteres genetiques distincts mis en evidence par des profils alloenzymatiques

differents. 11 se retrouve au Nouveau-Brunswick et dans la vallee du Saint-Taurent au Quebec, dans la region etats-unienne et canadienne des

Grands Tacs, dans les Black Hills au Dakota du Sud, puis a fouest, dans les Montagnes Rocheusesjusque dans Test de la Colombie-Britan-

nique et dans l’Etat de Washington.

INTRODUCTION

The moonworts ( Botrychium Sw. subgenus Botrychium) of North America include about 30 species, most of

which were described in the past 35 years (Wagner & Wagner 1981, 1983b, 1986, 1990a, 1990b, 1993, 1994;

Farrar & Johnson-Groh 1991; Stensvold et al. 2002; Wagner & Grant 2002). Several putative new species re¬

main under study (Stensvold 2008; Farrar & Popovich 2012). Many species are morphologically cryptic, sepa¬

rated by subtle differences of trophophore (sterile segment) shape and lobing, pinna shape, and dissection.

Treatment of these entities as evolutionary species is well-supported by cytological, isozyme, and DNA data

(Wagner 1993; Hauk 1995; Hauk & Haufler 1999; Zika & Farrar 2009).

In this paper we examine plants initially confused with Botchium matricariifolium A. Br. in the Great

Lakes area and with B. hesperium (Maxon & Clausen) Wagner & Lellinger in the Black Hills and Rocky

Mountains. Wagner and Wagner (1990a) reported the apparent discovery of B. hesperium in Ontario and

Michigan, but later they hypothesized these plants represented a new, undescribed species, which they infor¬

mally called “B. michiganense.” This hypothesis received support from the findings of Hauk and Haufler

(1999: Table 3) who reported differences between a Great Lakes population (i.e., “B. michiganense” including

Wagner’s collection 86045) and a western population (i.e., typical B. hesperium) in three allozyme systems ( Pgi-

2, Dia-1, and Mdh-1) of six that they examined. Hauk and Haufler’s (1999) results also demonstrated that

Wagner’s 86045 was, like B. hesperium and B. matricariifolium, an allotetraploid with fixed heterozygosity. We

J. Bot. Res. Inst. Texas 9(2): 295-309.2015

296

Journal of the Botanical Research Institute of Texas 9(2)

used additional allozyme profiles to analyze relationships between B. michiganense, B. hesperium and B. matri-

cariifolium and potential progenitors. We also compared morphologies of this new taxon with the species it had

been confused with, B. matricariifolium in the East and B. hesperium in the West.

MATERIALS AND METHODS

Allozymes

Diagnostic allozyme banding patterns consistent with morphological markers for B. michiganense established

by W.H. Wagner and the authors were identified from populations in Michigan (including the type locality),

Montana and Washington, where B. michiganense co-occurs with B. matricariifolium and with B. hesperium

respectively. Continued study of B. michiganense across the range of its occurrence along with other twice-

dissected (pinnate-pinnatihd) species allowed critical comparison at 20 independently assorting gene loci. For

this comparison, 302 individuals of these three allotetraploids from 37 sites were included (Appendix 1). A

small portion of the base of the common stalk from each specimen was removed for analysis, and the remain¬

der of the plant was pressed as a voucher and deposited at ISC; such samples of B. michiganense are cited in

Additional Specimens under Farrar numbers, where each number represents a single specimen, not a sheet with

multiple specimens.

Ten isozyme systems showing 20 loci were analyzed: aspartate amino transferase ( Aat ), 1-aminocyclo-

propane-l-carboxylate oxygenase (Aco), diaphorase (Did), isocitrate dehydrogenase (Idh), malate dehydroge¬

nase (Mdh\ 6-phosphogluconate dehydrogenase ( 6Pgd ), phosphoglucoisomerase (Pgi), phosphoglucomutase

(Pgm), shikimate dehydrogenase (Skdh) and triosephosphate isomerase ( Tpi ). Procedures and protocols fol¬

lowed those described by Zika & Farrar (2009).

To determine potential progenitors, results were compared to similar data from three diploid species:

Botrychium lanceolatum (S.G. Gmel.) Angstr., B. angustisegmentum Pease & A.H. Moore, and Botrychium palli¬

dum W.H. Wagner.

Morphology

We studied more than 225 individuals of B. michiganense (Appendix 2), including more than 150 for which

allozymes were analyzed, and assessed them for seven morphological variables on the trophophore (Table 1).

Specifically, we assessed samples that were growing together in mixed communities, as recommended by

Wagner and Wagner (1983a). Such communities are naturally occurring common garden experiments, with

all taxa growing under identical conditions. We studied the large populations of B. matricariifolium growing

with B. michiganense at the latter’s type locality, Grand Sable Dunes in Michigan’s Upper Peninsula, and B.

hesperium growing with B. michiganense at Bestrom Meadow, Stevens County, Washington, at Big Prairie (“Big

Meadow”) in Glacier National Park, Glacier County, Montana, and at Great Northern Flats of the North

Branch, Flathead River, Flathead County, Montana. We also compared plants from these populations with oth¬

ers from communities containing only one taxon, located in regions where B. michiganense has not been found:

B. hesperium from Colorado and B. matricariifolium from Vermont. Representative specimens for this assess¬

ment are cited in Appendix 2.

RESULTS

Allozymes

Botrychium michiganense differs from B. hesperium at four of 20 loci tested in the ten enzyme systems, and from

B. matricariifolium at three of 20 loci (Table 2). Allelic patterns indicate that B. michiganense is an allotetraploid

species (2n=180), with fixed heterozygosity displayed at twelve of 20 gene loci. Data indicate that it is related to

a common ancestor of B. angustisegmentum and B. lanceolatum, and to B. pallidum.

Morphology

The genus community tests show that morphological differences observed between multiple taxa growing

at remote locations are retained when they grow in proximity. There are consistent differences between

Gilman et al., Botrychium michiganense sp. nov.

297

Table 1. Morphological variables assessed on more than 300 individual specimens of Botrychium michiganense.

Character

Variation recorded

Trophophore attachment

sessile vs. stalked

Trophophore outline

ovate vs. elongate-ovate

Relative size of pinnae

basal pinnae much larger than distal pinnae vs. proportionately sized

Spacing of medial pinnae

remote vs. overlapping

Incisions of non-basal pinnae

both margins vs. only basiscopic margin

Pinnae shape

lanceolate-rhombic vs. broadly ovate to obovate

Pinnae apex

acute vs. rounded

Table 2. Alleles expressed at enzyme-coding loci in B. michiganense and related species. Allele numbers reflect relative migrating positions within genus Botrychium,

with alleles of the presumed diploid parents color coded. In allotetraploid combinations, the presumed Sect. Lanceolatae contribution is listed first and the presumed

Sect. Lunariae second using the color of the presumed contributing diploid, or black where the contributor is ambivalent. Allele numbers in () are orphan, not

detected in any of the three diploid species, but presumed to have been contributed by the non -Lanceolatae parent. Species are B. angustisegmentum (ang), B.

lanceolatum (Ian), B. matricariifolium (mat), B. michiganense (mich), B. hesperium (hesp) and B. pallidum (pal).

*lt is assumed that tetraploid plants have received two homoeologous copies of each gene, but because these genes are redundant in activity, one of the

homoeologous copies may have become silenced (n). In these data we have not attempted to detect this possibility at loci receiving the same allele from both

parental diploids. Where parental diploids differ in potential contribution but only one allele is expressed, we suggest silencing as the most likely explanation.

**Diploid Botrychium species regularly express banding patterns reflecting four independently migrating loci for diaphorase, but B. pallidum expresses

only 3 bands for this enzyme family. It is possible that Dia-3 has become silenced or that it co-migrates with one of the other loci.

Locus

ang

Ian

mat

mich

hesp

pal

Aat-2

3+(2)

3+(2) or 3+3

3+3

Aat-3

0.2

0.5

0.2+2

0.2+2 or 0.5+2

0.5+2

Aat-4

0.2

0.5

0.2+3

0.2+3 or 0.5+3

0.5+3

Aco-2

3+3

2 or 3

Dia-1

1+2

Dia-2

0.5

1+1

1+1 or 0.5+1

0.5+1

Dia-3

2.5 or 3

3+n

3+n*

n**

Dia-4

5+8

n+8

ldh-1

1+1

Mdh-1

2+1

Mdh-2

3+3

7+3

Mdh-3

2.5 or 3

3+2

3+2 or 2.5+2

3+2

Mdh-4

2+2

6Pgd

4+1

Pgi-2

4+(1) or 4+2 orn+2

4+2

4+2 or4+n

Pgm-1

3+1 or3+n

3+1 orn+1

Pgm-2

1.5

1.5+2

Skdh

2+1 or 2+n

2+1 or 1 +

1+1

Tpi-1

3+(1) or 3+3

3+3

Tpi-2

3+3

Botrychium michiganense, B. matricariifolium, and B. hesperium (Table 3) across the known geographic ranges of

these species.

DISCUSSION

Allozymes

Genetic relationships among twice-dissected allotetraploids of Botrychium are complex. Unquestionably, their

ancestry includes a twice-dissected (pinnate-pinnatihd) diploid member of B. sect. Lanceolatae Clausen, and a

once-dissected (pinnate) diploid member of B. sect. Lunariae Clausen (Clausen 1937; Hauk 1995; Hauk &

298

Journal of the Botanical Research Institute of Texas 9(2)

Table 3. Normal states of morphological characters in B. michiganense and species with which it has been confused. All characters are somewhat variable and may

not discriminate taxa when used alone.

Character

michiganense

Botrychium sp.

matricariifolium

hesperium

Trophophore attachment

sessile

stalked

variable

Trophophore outline

elongate-ovate

ovate to oblong

Ovate

Pinnae relative sizes

disproportionate

proportionate

Medial pinnae

remote

overlapping

2 nd and distal pinnae incisions

basiscopic

both margins

Pinnae shape

lanceolate-rhombic

ovate to oblong

ovate to obovate

Pinnae tip angles

acute

acute to rounded

rounded

Haufler 1999; Hauk et. al. 2012; Dauphin et al. 2014). To decipher the genetics and origin of B. michiganense it

is helpful to first consider the relationship of western B. hesperium to eastern B. matricariifolium.

The allelic composition of western B. hesperium supports an ancestry of B. lanceolatum (=B. lanceolatum

subsp. lanceolatum ) x B. pallidum, whereas the allelic composition of eastern B. matricariifolium supports an

ancestry of B. angustisegmentum (=B. lanceolatum subsp. angustisegmentum (Pease & A.H. Moore) R.T.

Clausen) x B. pallidum, as shown in Table 2. At six of 20 gene loci tested, these two Lanceolatae species differ in

alleles present. At five of these six discriminant loci ( Aat-3, Aat-4, Dia-2, Mdh-2, Skdh), it is the B. lanceolatum

allele that is present in B. hesperium (at the 6 th discriminant locus, Dia-4, no B. sect. Lanceolatae allele is ex¬

pressed in B. hesperium). Botrychium matricariifolium expresses the allele contribution of B. angustisegmentum

at all six discriminant loci. These parentages also correspond to the western North American sympatric distri¬

butions of B. lanceolatum with B. hesperium and eastern North American sympatric distributions of B. angus¬

tisegmentum with B. matricariifolium.

Botrychium michiganense differs from both B. matricariifolium and B. hesperium in combining the dis¬

criminant alleles of both B. angustisegmentum and B. lanceolatum. At Mdh-2, B. michiganense consistently ex¬

presses the allele of B. lanceolatum, whereas at Dia-4 it consistently expresses the allele of B. angustisegmentum.

At the other four discriminant alleles (Aat-3, Aat-4, Dia-2, Skdh) in different populations throughout its range,

B. michiganense expresses either the allele of B. lanceolatum or the allele of B. angustisegmentum. This condition

likely reflects multiple origins of B. michiganense through hybridizations between the same two ancestral dip¬

loid taxa in which an ancestral “B. lanceolatum ” parent possessed both alleles at these four loci, i.e., the

Lanceolatae parent of B. michiganense may have been an ancestor to both current B. lanceolatum and B. angus¬

tisegmentum that existed prior to the current differentiation of those two species. A “multiple origins” scenario

has been demonstrated for other alloploid ferns (Beck et al. 2014; Sigel et al. 2014) and the distributional exten¬

sion of B. michiganense across the ranges of both current taxa is consistent with this hypothesis.

An allozyme survey of the known diploid species of Botrychium sect. Lunariae (Farrar, unpublished) indi¬

cates that B. pallidum is capable of having provided the greatest number of non -Lanceolatae alleles present in B.

michiganense. The necessary non -Lanceolatae allelic contribution was detected in current populations for 19 of

the 20 gene loci analyzed. Only at Aat-2 is an allele expressed in B. michiganense that has not been detected in

either B. pallidum or any member of the B. sect. Lanceolatae complex. This allele, Aat-2=2, is also present in B.

matricariifolium, suggesting that it was likely present in B. pallidum at the time of the formation of B. matricari¬

ifolium, which is also thought to be descended from a cross or crosses between ancestral B. lanceolatum and

ancestral B. pallidum (Farrar 2011). Two additional non -Lanceolatae alleles are also present in B. matricariifoli¬

um at Pgi-2 and Tpi-1 that have not been detected in extant B. pallidum. A simple hypothesis is that ancestral B.

pallidum, like ancestral B. lanceolatum, contained allelic variability not retained in the current taxon. Other

hypotheses explaining its allelic composition seem less parsimonious, as no other species share as many

alleles.

Gilman et al., Botrychium michiganense sp. nov.

299

In summary, the allelic composition of related taxa, based on allelic analysis of gene loci for 20 conserva¬

tive metabolic enzymes, supports recognition of allotetraploid B. michiganense as distinct from both B. hesper-

ium and B. matricariifolium, based on parentage by a distinct set of diploid parent taxa. This analysis suggests

an origin for B. michiganense involving hybridization between B. pallidum and an ancestral member of “B. lan-

ceolatum” genetically different from, and probably ancestral to, the current eastern species B. angustisegmen-

tum and the western species B. lanceolatum.

Morphology

The genus community tests show that morphological differences observed between multiple taxa growing at

remote locations are retained when they grow in proximity. There are consistent differences between

Botrychium michiganense, B. matricariifolium, and B. hesperium (Table 3).

The most consistent characters that distinguish B. michiganense are the relative size of the proximal pin¬

nae, and the incision of the non-basal pinnae. Usually, the basal pinna pair is disproportionately elongate in

relation to the medial and distal pairs, which are abruptly shorter than the basal pair and then proportionately

reduced from one to the next. This character sometimes occurs in B. hesperium but is rare in B. matricariifolium.

The nature of the second degree of dissection is especially apparent in large B. michiganense specimens, where

incisions that are on medial and distal pinnae are deeper on the basiscopic than on the acroscopic margins; in

many average-sized specimens only the basal pinnae and the basiscopic margins of the second pair are incised.

When medial and distal pinnae of B. hesperium and B. matricariifolium are incised, the incisions are equally

deep on both the basiscopic and acroscopic margins. Also consistent within B. michiganense are the narrowly

rhombic-lanceolate, acute and remote pinnae, while B. hesperium pinnae are characteristically obovate, obtuse

and frequently overlapping. Consistent within many B. michiganense are sessile to subsessile trophophores,

compared to the distinctive stalked trophophores of B. matricariifolum.

We present here a key to B. michiganense and sympatric twice-dissected species with which it might be

confused.

1. Plants east of the Great Plains, including northern Alberta.

2. Trophophore sessile or short-stalked to 3 mm, ovate or narrowly ovate; pinnae abruptly reduced in size and cutting

from basal to next pair; pinnae above basal pair showing less cutting on acroscopic margins relative to basiscopic

margins_ B. michiganense

2. Trophophore usually stalked, stalk to 10 mm (rarely sessile), ovate to oblong; pinnae gradually and symmetrically

reduced from base to apex; if pinnae are dissected, incisions equivalent in depth on both basiscopic and acroscopic

margins.

3. Trophophore broadly ovate, pinnae very broadly ovate and rounded, incised_ B. pseudopinnatum

3. Trophophore narrowly ovate to oblong; pinnae narrowly to broadly ovate, in some instances not incised

_B. matricariifolium

1. Plants west of the Great Plains.

4. Trophophore stalk prominent, equal or greater than the distance between the 1 st and 2nd pinna pair; trophophore

broadly triangular_ B. pedunculosum

4. Trophophore sessile to short-stalked, narrowly elongate-ovate to broadly ovate.

5. Sporophore pinnately branched; trophophore lamina lustrous; pinnae always deeply lobed_ B. pinnatum

5. Sporophore ternately branched; trophophore dull; pinnae simple to lobed (incised).

6. Trophophore broadly to narrowly ovate, short stalked; basal pinna usually not disproportionately enlarged, ovate

to rhombic; medial pinnae usually gradually and progressively diminished in size from base, often slightly to

strongly overlapping, broadly ovate to obovate, margins entire to incised and, if incised, then more or less

equally so on basiscopic and acroscopic margins, pinna tips rounded_ B. hesperium

6. Trophophore lanceolate to narrowly ovate, sessile to nearly so; basal pinnae often noticeably longer than me¬

dial pinnae, oblong in pinnae with 3 or more pinnules, ovate with margins crenulate to incised in smaller plants;

medial pinnae separated, lanceolate to narrowly ovate or oblong, entire or, if crenate or dissected, then more

deeply so on basiscopic margins, tips acute or subacute_ B. michiganense

With both allozymes and morphology confirming the distinctiveness of B. michiganense as first recognized by

Herb and Florence Wagner, we consider it to be a new species of allopolyploid origin, and use the name that

Herb suggested.

300

Journal of the Botanical Research Institute of Texas 9(2)

TAXONOMY

Botrychium michiganense W.H. Wagner ex A.V. Gilman, Farrar, & Zika, sp. nov. (Figs. 1-3). Type: MICHIGAN:

Grand Sable Dunes, 27-28Jun 1987, W.H. Wagner 87228A, F.S. Wagner , Farrar &Hauk (holotype: NY; isotypes DAO, GH, ISC, MICH,

MO, US, WTU).

A Botrychio hesperio fronde plus elongo-ovato, pinnatibus basis duobus saepe maiusculis, pinnis mediis remotioribus, atque secundis duo-

bus dissectis basiscopis sed saepe non acroscopis marginibus, differt; a B. matricariifolio laminis saepe sessilibus, pinnis angustioribus, et

illis sigillis supra enumeratis, differt.

Differing from B. hesperium in more elongate-ovate frond, frequently elongated basal pinnae, remote medial

pinnae, and pinna-dissection usually confined to the basal pinnae and the basiscopic margin of the second

pinna-pair. Differing from B. matricariifolium in often sessile to short-stalked trophophore, often elongated

basal pinnae, often proportionately narrower pinnae, 0.4x as wide as long (vs. >0.5x as wide as long) and asym¬

metrical dissection of pinnae above the first pair, with much deeper cutting on the basiscopic side.

Plants fibrous-rooted, non-rhizomatous, herbaceous, deciduous, to 5-21 cm tall, distally with two differ¬

entiated segments: the foliar, non-sporangiate trophophore with pinnately divided lamina and the non-leafy,

ternately much-branched sporophore with numerous sporangia. Proximal common stalk 3-15 cm, diameter

1.6-2.5 mm, trophophore and sporophore elevated well above ground. Sporophore ca. 3-10 cm, often twice

as long as trophophore, with proximal unbranched stalk 0.5-lx as long as trophophore; axis usually (>90%)

medially branched with 3 main branches, each with 3+ pairs of additional branches, and these (in large

specimens) also with 1+ pairs of additional branches (at most 3 orders of branching); all branches ascending.

Sporangia approximate but not clustered, broadly attached, globose, 1.0-1.3 mm in diameter, dehiscing by

distal transverse slit of ca. one-half the circumference. Spores whitish, globose-tetrahedral (38-)41.9(-45)

pm diameter. Trophophore gray-green to medium green and dull in life, sessile or mostly short-stalked, stalk

0.5-3(-5) mm, ovate to usually (>67%) broadly to usually narrowly elongate-ovate (l-)2-2.5(-3) cm at widest

point x (l-)2.5-4(- 5) cm long. Basal pinnae often disproportionately elongated, becoming oblong and paral¬

lel-sided, to 1.5-2x as long as next most distal pinnae, the lamina then with an abrupt transition between the

first two pinnae pairs from long to short pinnae. Basal pinnae usually (always in large specimens) pinnately

divided with 3-5 pairs of pinnules, sinuses as deep on acroscopic as on basiscopic margin, elongate-ovate.

Medial and distal pinnae (2-)3-6(-7) pairs, broadly attached (attachment ca. 0.5x as wide as widest part of

pinna), ascending, remote, gradually decreasing in size upwards (distalmost small and approximate or overlap¬

ping), elongate-ovate to more frequently narrowly lanceolate-rhombic, (0.35-)0.40-0.44(-0.5) times as wide

as long; in most only the second pair incised and then only on basiscopic margin; tips obtuse to usually acute.

Etymology. —From Latinized “Michigan,” where this taxon was first recognized by Herb and Florence

Wagner; referring both to the State of Michigan and to the Great Lakes, originally from Native American

Ojibwe meaning “big lake.”

Habitat. —Habitats supporting B. michiganense are typical for moonworts that favor mesic to xeric mead¬

ows. Sites are sunny, open areas with sporadic or periodical disturbance sufficient to prevent canopy closure

by woody plants. Soils are often calcareous, of neutral reaction, and have minimal organic matter. The overall

communities are usually mid-successional or delayed-successional, often with some bare soil still visible, and

are often diverse in perennial herbs of whatever species are common in the local area. Other moonworts are

often the most faithful associates of B. michiganense, likely due to a shared need for myocorrhizal associations

with certain glomalean fungi (Winther & Friedman 2007). Moonwort species documented to occur with

B. michiganense in “mixed genus communities” include Botrychium ascendens W.H. Wagner, B. campestre

W.H. Wagner & Farrar, B. furculatum Popovich & Farrar ined., B. gallicomontanum Farrar & Johnson-Groh,

B. hesperium, B. lanceolatum, B. lineare W.H. Wagner, B. matricariifolium, B. minganense Viet., B. pallidum, B.

paradoxum W.H. Wagner, B. pedunculosum W.H. Wagner, and B. simplex E. Hitchc. Otherwise, there is no

general association with any particular herb or herbs across its range, although Fragaria L., strawberry, is often

present.

Known sites in the western portion of its range include high elevation prairies and pastures, periodically

Gilman et al., Botrychium michiganense sp. nov.

Fig. 1. Botrychium michiganense, Holotype, unmounted, Wagner87228A, Wagner, Farrar &Hauk (NY). Scale bar 10 cm.

302

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 2. Botrychium michiganense compared to B. hesperium and B. matricariifolium. Three plants at left (1461,1180,1336): B. matricariifolium ; three at

center (17326,2981,18410): B. michiganense; three at right (1854,1808,3857): B. hesperium.

burned sagebrush prairie (the dry extreme of suitable habitats), gravel flats and rarely-flooded river terraces

along high-energy rivers, avalanche meadows, ski areas, and abandoned gravel roads and roadcuts. In the

Black Hills of South Dakota, B. michiganense occurs in open pine and deciduous woodlands and in mountain

meadows as well as on roadsides. Eastern habitats include stabilized dune meadows, abandoned settlements,

dry fields, maintained clearings along utility lines, roadsides, lawns, and mine-tailings. Altitudinal range in

the East is from near sea level to ca. 520 m, in the West from ca. 1500 m to 2600 m or higher, although generally

below treeline.

Distribution.—Botrychium michiganense is known across North America in several general regions (Figs.

4-6). The easternmost specimens seen are from near the Gulf of St. Lawrence in New Brunswick and along the

St. Lawrence River Valley in Quebec. It occurs near the northern Great Lakes (Michigan and Superior) and

extends westward across northern Minnesota. Within the Great Plains it occurs in the Cypress Hills of

Saskatchewan and Alberta, and the Black Hills of South Dakota. West of the Great Plains it occurs on the foot¬

hills and in the Rocky Mountains in Alberta and British Columbia, south to Wyoming, Montana, and

Washington. The westernmost locales are in the Colville National Forest north of Spokane, Washington. In

some areas, including northern Minnesota, the Black Hills National Forest in South Dakota, northwestern

Gilman et al., Botrychium michiganense sp. nov.

303

Fig. 3. East to west morphological variation in Botrychium michiganense over its range. Top row, left to right: eastern Quebec 16332; northern Michigan

1068,562,8264; southern Ontario 17327. Middle row: northern Minnesota 14239,18145; western South Dakota 15027; northern Wyoming 18410,

15018. Bottom row: western Montana 2843; eastern Washington 1864, 1873, 2981,2984. Numbers refer to genetically analyzed specimens (see

Additional Specimens Examined). Scale = 1 cm.

Montana, and northeastern Washington, it appears to be a relatively common species. For example the Black

Hills National Forest has more than 30 currently known populations (Fig. 6), illustrating the results of dedi¬

cated searches in a given area once search images for morphology and habitat are established.

Additional specimens examined.—Note: In addition to the original collector’s numbers, some material sent to Farrar’s lab received addi¬

tional numbers [in square brackets], which represent identification numbers of individual plants analyzed for allozyme study.

CANADA. ALBERTA: Cypress Hills Provincial Park, SE Corner Hwy 41 & Reesor Lake Road, 9 Jul 2002, Williston 3764 & Bartemucci

304

Journal of the Botanical Research Institute of Texas 9(2)

98°W 96°W 94°W 92°W 90°W 88°W 86°W 84°W 82°W 80°W 78°W 76°W 74°W 72°W 70°W 68°W 66°W 64°W

(ALTA). South Dry wood Creek, S facing slope, 21 Jul 1999, Gould, T. Dolman & D. Dolman s.n. (ALTA). La Butte Creek Wildland Provincial

Park, rock outcrop dominated by lichens, 6 Jul 2001, Vujovnic et al, s.n (ALTA). Waterton Lakes National Park, S side of Rte. 6 at the Lewis

Overthrust Scenic Site parking area, 3 Aug 1983, Wagner 83318 & Wagner (MICH); Snowshoe Trail, ca. A mi NW of Red Rock Canyon park¬

ing, large open grassy slope, near pine woods, 5 Aug 1983, Wagner 83328 & Wagner [mixed sheet with B. hesperium] (MICH). Prospect

Creek, Lange Ridge, along trail, open grassy slope, 21 Jun 2008, Fabijan, Dinwoodie & Anderson 02985 (ALTA). Elk Island National Park,

Sand Hills Trail 1,13 Jun 2005, Farrar 12122-12125 (ISC); Elk Island National Park, E corner, along old access road kept open by bison graz¬

ing, 10 Jun 2001, Williston 3423, Cotterill & G. Griffiths (ALTA); Elk Island National Park, Moss Lake Trail, open meadow area, 24 Jun 2001,

Cotterill & G. C. D. Griffiths s.n. (ALTA). Athabasca Co.: Crooked Lake Valley, 28 Jun 2014, D. E. Griffiths s.n. (ALTA). County of Minburn:

Lac La Biche (Birch Lake), 12Jun 2005, G. Griffiths s.n. [=Farrar 12087-12093] (ISC); Lac La Biche, Birch Island, shrubby meadow on summit

of island, 11 Jul 2007, T. M accagno s.n. (ALTA). County of Parkland: Jackfish Lake, 16 Jun 2005, Farrar 12246,12247 (ISC); Jackfish Lake,

29 Jul & 18 Aug 2000, G. C. D. Griffiths s.n. (ALTA). BRITISH COLUMBIA. Arrowhead: Upper Arrow Lake, occasional along gravel road

near church, 10 Jun 1958, Colder 8933 & Savile (DAO). NEW BRUNSWICK. Kent Co.: Kouchibougouac National Park: St. Louis Parish,

near Kelley’s Beach, 26 May 1977, Munro 352 (DAO). ONTARIO. Algoma District: 2.5 mi W of Wawa, just SE of Algoma Central RR bridge

over Rte. 17, 22 Jun 1988, Wagner 88048 & Wagner (MICH); W of Siderite Jet. near Wawa, 23 Jun 1992, Gilman 92070, Wagner & Wagner

(VT); Michipicoten Harbor, near Milmac Mine, sand amongst grass, 12 Jul 1938, Hosie et al. 981 (DAO). Thunder Bay District: Marathon,

21 Jul 2010, Farrar 18274 (ISC); Neys, south side of railway tracks near Neys Provincial Park, E side of park entrance road, open disturbed

meadow, 28 Jun 2009, Oldham & Brinker 36161 [= Larrar 17327] (ISC); Sibley Peninsula, crest at head of Thunder Cape, thin soil in exposed

locations, 14 Jul 1946, Garton 1040 (DAO); Sibley Peninsula, Silver Islet, E of Sibley Creek, dry sandy field, 18 Aug 1950, Garton 1278 &

Campbell (DAO); Sibley Cove, open sandy clearing by old cemetery, 25 Jun 1936, Faylor, Losee & Bannon s.n. (GH). QUEBEC. Rimouski Co.:

Bic Provincial Park, pres du Camp du Cap a l’Orignal, 24 Jun 2001, Cayouette C8840 et al. (DAO); same locale, 23 Jun 2008, Farrar 16332,

16334 (ISC); same locale, 25 Jun 2008, Farrar 16453 (ISC); Bic Provincial Park, Cap a l’Orignal, terrasse et prairie du haute rivages, sable

graveleux et sec, 8 Jul 2008, Cayouette, Farrar & LaBrecque s.n. (DAO); Cap a l’Orignal, beach, 6 Jul 1906, Fernald & Collins 297b (GH); St.-

Denis-sur-Mer, 47°30 , 34.2"N, 69 0 57'24.3"W, pelouse, herbagaie moyenne a basse sur le haut rivage du fleuve St. Laurent, 6 Jul 2008,

Cayouette C9618A (DAO). Cote-Nord: Sept-Iles, Sept-Riviere, aeroport de Sept-Iles, 50°13T5.6"N, 66°15 , 28.7"W, prairie herbacee, 27 Jun

2009, Lynch DL09-041B, DL 09-046 (DAO). SASKATCHEWAN. Cypress Hills: edge of pine woods, 1 Jul 1947, Breitung 4188 (DAO). Cypress

Hills Provincial Park, Centre Block, W side of road N of Lire Lookout Tower, extensive Potentillafruticosa pastures, 28-30 Jul 1983, Wagner

83302 & Wagner (MICH).

UNITED STATES. MICHIGAN. Alger Co.: Grand Sable Dunes, open grassy areas and on slopes with shrubs, 25 Jun 1985, Wagner

85057 & Wagner (MICH); same locale, 28 Jun 1985, Farrar 562 (ISC); same locale, 21 Jun 1995, Farrar 1058 (ISC); shady woods in and around

Lakeshore Trail, 1-2 September 1985, Wagner 85073A & Wagner (MICH); in jack pines, shade form, 21 Jun 1986, Wagner 86042 & Wagner

[mixed sheet with B. matricariifolium] (MICH); 1.4-2.0 mi from beginning of Lakeshore Trail at Grand Sable Lake, 22 Jun 1986, Wagner

86045 & Wagner [mixed sheet with B. matricariifolium ] (MICH); just N of Sable Lake, 22 Jun 1993, Gilman 93090, Wagner & Wagner (VT).

Gilman et al., Botrychium michiganense sp. nov.

305

128°W 126°W 124°W 122°W 120°W 118°W 116°W 114°W 112°W 110°W 108°W 106°W 104°W 102°W 100°W 98°W

Chippewa Co.: Forestry Road 3139 and railroad crossing, 10,17Jun 1988, Wagner 88014, 88016.5 & Wagner (MICH); Bobbygay Lake Road

near Trout Lake, 14Jun 1994, Gilman 94051, Wagner et al. (VT); Bobbygay Road, 19Jun 2003,Jaunzems s.n. [= Farrar 8264] (ISC). Leelanau

Co.: S. Manitou Island, Garden City area, field, 12 Jun 1985, Wagner 85038A & Wagner (MICH). MINNESOTA. Cook Co.: Sawbill Camp, 29

Jun 1998, Farrar 2921, 2922, 2923 (ISC); T61N R4W S8NENWNW, Superior National Forest, Sawbill CCC Camp, 12Jul 2004, Gerdes &

Lawson 618 (MIN); T62N R1E S30 SWNW, Grand Marais Seaplane Base, NE shore of Devil Track Lake, 16 Jun 2007, M cFarlane et al. 9 (MIN);

T64N R01W S10SWNE, Superior National Forest, 0.6 mi NE of Clearwater Lake, 18 Jun 1998, Gerdes 2753 (MIN); T64N R03E S12NENWS,

Superior National Forest, NE of Grand Marais, S Fowl Lake Cliff, 16 Jun 2005, Gerdes et al. 5092 (MIN); T65N R04W S27NWSE, Superior

National Forest, Cross River area, FR 320, 7 Jun 1998, Gerdes 2629 (MIN); T65N R04W S26SENE, Superior National Forest, edge of old

gravel pit and slash burn area, lOJul 1999, Gerdes 3556 (MIN); Pancore Lake, 11 Jul 2003, Greenke s.n. [= Farrar 8667-8670] (ISC). Itasca Co.:

T56 R24 NE A of NE A of Sec 34, taconite tailings basin S of Holman, 30 May 2001, Dahl, Engels & Butler 01006, 01007, 01009 [= Farrar

4926-4928, 4931, 4937-4940, 4942-4944, 4946, 4947] (ISC); taconite tailings basin 2 mi SE of Bovey, 21 Jul 1999, Dahle, Engels & Butler s.n.

[= Farrar 4078] (ISC). Koochiching Co.: Pine Island State Forest, Pine Island Forest Road, N side, SWSW of section, 6 Jun 2011, M acFarlane

& MacFarlane 182 (MIN); Pine Island State Forest, Indian Pines Forest Road, N side, NESE of section. 5 Jun 2011, MacFarlane & MacFarlane

166 (MIN). Lake of the Woods Co.: T160N R34W SWSE34, Beltrami Island State Forest, Nelson Forest Road at State Forest Road 1181,6 Jun

2010, MacFarlane 138 [mixed sheet] (MIN); Beltrami Island State Forest, Bankton Forest Road, 8 Jun 2012, MacFarlane 347 (MIN). Roseau

Co.: Beltrami Island State Forest, Penturen Forest Road, old homestead SESE of Section, 19 Jun 2011, MacFarlane & MacFarlane 267 (MIN).

St. Louis Co.: T57N R20W NE SE 32, mine dump E of Hwy 69,0.5. N of Kitzville, 5 Jun 2007, MacFarlane et al. 25 (MIN); T58N R19W SW

SW 32, 2 mi S of Buhl, N of Hayes Rd., 1 Jul 2006, MacFarlane et al. 19 (MIN); Superior National Forest, T58N R18W NE NW 28, Skibo

Landing, NE side of railroad tracks, 30 Jun 2006, MacFarlane et al. 20 (MIN); T58N R15W NE NW 28, mine dump E of Hwy 138 to Giant’s

Ridge, 6 Jun 2007, MacFarlane et al. 30 (MIN); T58N R15W SW SW 2, on a mine dump ca. 0.5 mi E of Aurora, 8Jun 2008, MacFarlane et al.

306

Journal of the Botanical Research Institute of Texas 9(2)

Black Hills Botrychium michiganense Occurrences

Fig. 6. Occurrence of B. michiganense in the Black Hills National Forest. Map reflects ten years of rare plant search by the Black Hills NF staff, beginning

with a single population discovered in 2002. Map courtesy of Cheryl Mayer, Botany Technician, Forest Service, Black Hills NF.

Gilman et al., Botrychium michiganense sp. nov.

307

70 (MIN); T57N R21W NW NW 15, on top of a mine dump W of Highway 60 & 63 intersection between Kelly Lake and Leetonia, 16 Jun

2007, M acFarlane et al. 42 (MIN); same locale, 7 Jun 2008, M acFarlane et al. 74 (MIN); T58N, in iron ore tailings basin, Off-Highway Vehicle

Recreation Area E of Gilbert, 30 Jun 2004, Dahle et al. 4038 (MIN) [=Farrar 14329] (ISC); Hibbing, 30 Jun 2005, Dahle et al. s.n. [= Farrar

12595-12602] (ISC); taconite tailings ca. 1 mi E of Hibbing, 23 Jun 1999, Dahle et al. 99009 [= Farrar 3610,3611,3612,3613,3614,3617] (ISC);

taconite tailings 2 mi W of Hibbing, 23 Jun 1999, Dahle et al. 99008 [= Farrar 3581-3586] (ISC); Gilbert, 24 Jun 2003, Dahle s.n. [= Farrar

8209] (ISC); Kelley Lake, 9 Jun 2008, Farrar 16078 (ISC); Markmanship Road, 19 Jul 2010, Farrar 18144 (ISC). MONTANA. Flathead Co.:

Glacier National Park, S end of Big Prairie [Big Meadow], elev. 3600 ft, NE of North Fork Flathead River, E of Inside Road, T35N R21W S16,

c. 48°49'N, 114°19'W, 20 Jun 1998, Wagner 98016 & Wagner (MICH); same locale, 14 Jul 2000, Larsen & Larsen 4551-4554,4557-4559 (ISC);

north end of Great Northern Flats, elev. 3300 feet, E side of Forest Service Road 486, W side of North Fork Flathead River, Flathead National

Forest, T33N R20W S35, 20 Jun 1998, Wagner 98023 & Wagner (WTU); same locale, 20 Jun 1998, Gilman 98043, Wagners, Zika, Lesica &

Forest Service botanists (VT); same locale, 19-20 Jun 1998, Farrar 2840-2844 (ISC); Fire History pulloff on Rte 486, S of Polebridge, 20 Jun

2005, Farrar 12393-12397 (ISC). Glacier Co.: a few meters from north shore of west end of Lake Sherburne Reservoir on Swiftcurrent Creek,

Apikuni Flat, elev. 4800 feet, T35N R16W S12, 21 Jun 1998, Zika 13434 & W.H. Wagner (WTU); same locale, 19-20 Jun 1998, Farrar 2884-

2888,2912,2913,2915-2917 (ISC). SOUTH DAKOTA. Custer Co.: Black Hills National Forest, Rocky Road, 30 May 2007, M ergen 06VO35-

36 [= Farrar 14265-14267] (ISC); Custer State Park, Cathedral Spires, 22Jun 2011, Mayer 1403 [= Farrar 18725] (ISC); Bowman, 24 May 2008,

M ergen 08C008B [= Farrar 16048] (ISC); Bowman, below old road of old burn, 29 May 2008, M ergen 08CO23A [= Farrar 16109-16111] (ISC);

Bowman, old burn, 5 Jun 2008, M ergen 08CO28D [= Farrar 16114] (ISC); Star, open slope, 21 Jun 2008, M ergen 08SO17D [= Farrar 16566]

(ISC). Lawrence Co.: Black Hills National Forest, Eagle Cliffs, 19Jun 2009, Farrar 17139 (ISC). Pennington Co.: Black Hills National Forest,

Pine Creek, 16 Jun 2008, Mayer s.n. [= Farrar 16126] (ISC); Calumet, 3-8 Jun 2009, M ergen 09CO10C [= Farrar 17110, 17111, 17141] (ISC);

Windmill, 26 Jun 2008, M ergen 08W002A [= Farrar 16567] (ISC); Harney Peak Trail, 17 Jun 2007, Farrar 18093, 18094, 18110 (ISC); Hat

Mountain, 21 Jun 2011, Farrar 18592 (ISC); Reynold’s Prairie, 16 Jun 2009, Farrar 17119 (ISC); Pactola, 10 Jun 2009, M ergen 09P007A [=

Farrar 17143] (ISC); Sixmile Draw, no date, Mayer 1400 [= Farrar 18724] (ISC); same locale, no date, Mayer & Burkhart s.n. [=Farrar 15027]

(ISC); same locale, no date, M ergen 09P007A [= Farrar 15320] (ISC). WASHINGTON. Stevens Co.: Colville National Forest, Paradise

Meadow, T38N, R41E, S15,26 Jun 1997, Farrar 1863-1877 (ISC); same locale, 14Jun 2002, Ahlenslager s.n. [= Farrar 6329-6333, 6334-6338]

(ISC); same locale, 24 Jul 2005, Legler2868,2869 (WTU); same locale, 30 Jun 1996, Larson 209 (WTU); Colville National Forest, Bestrom

Meadow, T36N R41E S19, 30 Jun 1997, Ahlenslager s.n. [= Farrar 1910-1917] (ISC); same locale, 7 Jul 1998, Farrar 2981-2989 (ISC).

WYOMING. Crook Co.: Black Hills National Forest, Bear Lodge Campground, T54N R62W S20,8 Jun 2004, Farrar 5015-5019 (ISC); same

locale, 16 Jul 2002, Farrar 6381 (ISC); same locale, 6 Jun 2007, Corey BLGC-A [=Farrarl5019] (ISC). Sheridan Co.: Big Horn Mountains: Pole

Creek Road, 17 Aug 2010, Farrar 18410 (ISC).

APPENDIX 1

Number of Botrychium specimens analyzed for allozyme composition. NP = National Park.

Collection sites

B. michiganense

B. hesperium

B. matricariifolium

Canada

Alberta - Birch Lake, Elk Island, Jackfish Lake

Ontario - Lake Superior north shore

Quebec - Bic NP, Kamarouska, St. Denis

United States

Colorado - multiple sites

Minnesota - multiple sites

Montana - Glacier NP, Flathead Lake

New Mexico - Vermejo Ranch

South Dakota - Black Hills, multiple sites

Wyoming - Black Hills, Bear Lodge Mts.

Total

APPENDIX 2

Botrychium specimens inspected for genus community comparisons.

Botrychium hesperium

Colorado: Grand Co.: Winter Park, JK Ski Run, Forror 14490-14501 (ISC)—11 specimens.

Montana: Flathead Co.: Great Northern Flats, Forror2837,2838,2839 (ISC), Zika 13432 (WTU)—13 specimens.

Montana: Glacier Co.: Glacier National Park, Big Prairie, Z/7ca 13439 (WTU); Forror2845-2849;2903-2907 (ISC)—21 specimens.

Washington: Stevens Co.: Bestrom Meadow, Forror2999-3013 (ISC); Forror 6357-6379 (ISC)—38 specimens.

Botrychium matricariifolium

Michigan: Alger Co.: Grand Sable Dunes: Forror 870627A, 876027B, 870627B-1,870627B-2 (all ISC)—26 specimens.

Vermont: Caledonia Co.: Gilmon 04026 (VT)—29 specimens.

308

Journal of the Botanical Research Institute of Texas 9(2)

Botrychium michiganense

Michigan: Alger Co.: Grand Sable Dunes: W.H. Wagner 87228k, F. S. Wagner, Farrar, & Hauk (MICH, GH, NY, DAO UC, US. ISC, VT, WTU, MO,

CAN, MONTU)—47 specimens.

Montana: Flathead Co: Great Northern Flats: Gilman 98043 etal. (VT)—10 specimens.

Montana: Glacier Co.: Glacier National Park: Big Prairie: Wagner 98106 & Wagner (MICH); Larsen & Larsen 4551-4554,4557-4559 (ISC)—16

specimens.

Washington: Stevens Co.: Bestrom Meadow: Farrar 1910-1917,2981-2989 (ISC)—17 specimens.

ACKNOWLEDGMENTS

We thank Kathleen Ahlenslager, Jacques Cayouette, Roger Ferriel, Graham Grifhthst, Karen Larsen, Kirk

Larsen, Michael Oldham, Peter Lesica, Cheryl Mayer, Rich Rabeler, Toby Spribille, Jim Vanderhorst, Herb

Wagnerf, Florence Wagner, and Tara Williams. We are grateful to the curators and staff at the following her¬

baria, who provided access to their collections or loans: ALTA, DAO, GH, ISC, MICH, MINN, VT, and WTU.

Weston Testo provided helpful comments on an early draft of the manuscript. We also thank Warren Hauk

and one anonymous reviewer for careful scrutiny and comments.

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Wagner, F.S. 1993. Chromosomes of North American grapeferns and moonworts (Ophioglossaceae: Botrychium). Contr.

Univ. Michigan Herb. 19:83-92.

Wagner, W.H. Jr. & F.S. Wagner. 1981. New species of moonworts ( Botrychium subg. Botrychium (Ophioglossaceae), from

North America. Amer. Fern J. 71:20-30.

Wagner, W.H. Jr. & F.S. Wagner. 1983a. Genus communities as a systematic tool in the study of New World Botrychium

(Ophioglossaceae).Taxon 32:51-63.

Wagner, W.H. Jr. & F.S. Wagner. 1983b. Two moonworts of the Rocky Mountains: Botrychium hesperium and a new species

formerly confused with it. Amer. Fern J. 73:53-62.

Wagner, W.H. Jr. & F.S. Wagner. 1986. Three new species of moonworts ( Botrychium subg. Botrychium) endemic in west¬

ern North America. Amer. Fern J. 76:33-47.

Wagner, W.H. Jr. & F.S. Wagner. 1990a. Moonworts ( Botrychium subg. Botrychium) of the upper Great Lakes region. Contr.

Univ. Michigan Herb. 17:313-325.

Gilman et al., Botrychium michiganense sp. nov.

309

Wagner, W.H. Jr. & F.S. Wagner. 1990b. Notes on the fan-leaflet group of moonworts in North America with descriptions

of two new members. Amer. Fern J. 80:73-81.

Wagner, W.FI. Jr. & F.S. Wagner. 1993. Botrychium. In: Flora of North America Editorial Committee, eds. Flora of North

America north of Mexico. Oxford University Press, New York U.S.A. and Oxford, U.K. 2:86-101.

Wagner, W.FI. Jr. & F.S. Wagner. 1994. Another widely disjunct, rare, and local North American moonwort

(Ophioglossaceae: Botrychium subg. Botrychium ). Amer. Fern J. 84:5-10.

Wagner, W.FI. Jr. & J. Grant. 2002. Botrychium alaskense, a new moonwort from the interior of Alaska. Amer. Fern J.

92:164-170.

Withner, J.L. & W.F. Friedman. 2007. Arbuscular mycorrhizal symbionts in Botrychium (Ophioglossaceae). Amer. J. Bot.

94:1248-1255.

Zika, P.F. & D.R. Farrar. 2009. Botrychium ascendens W.FI. Wagner (Ophioglossaceae) in Newfoundland and notes on its

origin. Amer. Fern J. 99:249-259.

310

Journal of the Botanical Research Institute of Texas 9(2)

BOOK REVIEW

E. Charles Nelson & DavidJ. Elliott. 2015. The Curious Mister Catesby. (ISBN-13: 978-0-8203-4726-4, hbk).

University of Georgia Press, Main Library, Third Floor, 320 South Jackson Street, Athens, Georgia

30602, U.S.A. (Orders: www.ugapress.org, 1-800-266-5842). $49.95 US, 456 pp., 238 paintings/illus./

photos/maps, 8" x 11".

Meticulous research and high-quality illustrations make The Curious Mister Catesby a comprehensive resource

bound to appeal to a wide audience. The book paints a portrait of the botanical world during Mark Catesby’s

life and in doing so, guides the reader in recognizing his exceptional talent.

Far from a straight-forward biography, the book is the work of nearly two dozen authors of various disci¬

plines, including anthropologists, historians, zoologists, botanists, and more. Every chapter analyzes Catesby’s

work from a different angle, yet each author’s perspective blends seamlessly with the next. The authors build a

detailed story of Mark Catesby’s life and times, which is especially impressive considering the lack of historical

documents about him as an individual. Herbarium labels, governmental records, diaries of Catesby’s contem¬

poraries, and other far-reaching resources track the course of his work.

Rife with high-quality illustrations—including etchings, herbarium specimens, paintings, maps, histori¬

cal documents, and modern photographs— The Curious Mister Catesby is suitable for both leisurely reading

and scholarly studies. The thick, large pages and sturdy binding make the book exceptionally easy to read, and

the plethora of illustrations are captivating. Although I appreciate the variety of illustrations, I would like to see

more of Catesby’s work, especially considering his artistic reputation.

The Curious Mister Catesby also includes chapters about Mark Catesby’s forerunners and contemporaries

and their influence on both him and natural history as a whole. Most notably discussed are Maria Sibylla

Merian, whose artistry Catesby is thought to have admired, and William Dampier, whose dedication and

thoughtful recordings of nature were not unlike Catesby’s.

Mark Catesby owes his great success in North America to a combination of his natural talent and ambi¬

tion and a powerful network of supporters who appreciated his passion for nature. That passion and talent

seeps through the pages of The Curious Mister Catesby, illuminating the man whose tireless excursions and

keen observations led to one of the most important publications of flora and fauna of the New World.— Alyssa

Young, Botanical Research Institute of Texas, Fort Worth, Texas, U.S.A.

J.Bot. Res. Inst. Texas 9(2): 310.2015

NOTES ON ERITRICHIUM (BORAGINACEAE) IN NORTH AMERICA II

David F. Murray

University of Alaska Museum of the North

907 Yukon Drive

Fairbanks, Alaska 99775-6960, US.A.

dfmurray@alaska.edu

ABSTRACT

The common, widespread Eritrichium of the Rocky Mountains is E. argenteum; the importance of the Asian E. sericeum s.l. to the North

American flora is noted; minor issues relating to the type for E. splendens, the treatment of E. chamissonis, and the disposition ofE. howardii

are given.

RESUMEN

El comun Eritrichium de las Montanas Rocosas es E. argenteum; La importancia del asiatico E. sericeum s.l. en la flora norteamericana es

notable; se aportan cuestiones menores relacionadas con el tipo de E. splendens, el tratamiento de E. chamissonis, y la disposicion de E.

howardii.

The purpose of the notes that follow is to provide background information that is more than can be accom¬

modated in comments sections of species accounts in Flora of North America north of Mexico. Additionally,

prior to the appearance of Eritrichium in Volume 15, nomenclatural issues have to be settled (cf., also Gandhi

& Murray 2013 and Murray 2013).

1. A name for the common, widespread Rocky Mountain Eritrichium

Aside from the very distinct Eritrichium howardii (A. Gray) Rydb. the alpine forget-me-not of the Rocky Moun¬

tains was initially named using combinations derived from earlier work in the Arctic: e.g. E. aretioides (Cham.)

DC, Omphalodes nana var. aretioides (Cham.) A. Gray. Rydberg (1900), with E. aretioides var. elongatum Rydb.,

provided the first recognition there might be an entity distinct from E. aretioides, perhaps, he commented, at

the rank of species, but which he published as a variety.

Wight (1902) reviewed the North American taxa, and he noted that the nutlets of Rydberg’s var. elongatum

have smooth margins, whereas some of the material at hand included specimens with nutlets having toothed

margins and different features of leaf indumentum. This he named E. argenteum W. Wight. He then raised var.

elongatum to species rank, and in so doing created a later homonym of E. elongatum (A.DC) A.DC. Wight made

the different types of nutlets the primary dichotomy in his key to species and thus did not work out whether

other characteristics of the two taxa showed a parallel discontinuity. Wight’s taxonomy was used by Rydberg

(1906) and Coulter and Nelson (1909) in what must have been widely used manuals of their day.

Macbride (1916) believed that the nutlet differences proposed by Wight (1902) to be constant and there¬

fore good specific characters. Johnston (1924) used the same basic dichotomy of nutlet morphology as Wight,

but treated those plants with toothed margins as distinct only at the varietal level and published the combina¬

tion E. elongatum var. argenteum (W. Wight) I.M. Johnst., noting that it was a taxon of the more southern por¬

tions of the range of “ elongatum .” This is the taxonomy followed by Harrington (1954).

Years later (1952) Johnston, when commenting on specimens from the Uinta Mountains of Utah received

from Payson, wrote that he saw these as indistinguishable from the “uncommon” var. argenteum, hinted that

they might rise to the rank of species, but ultimately used this material as the basis for E. elongatum var. paysonii

I.M. Johnst.

Cronquist (1959) held the view that, with the exception of E. howardii, all the Rocky Mountain variation

of Eritrichium fell within the limits of E. nanum var. elongatum (Rydb.) Cronq., He also recognized E. nanum var.

aretioides (Cham.) Herder, but not in the Rocky Mountains.

J. Bot. Res. Inst. Texas 9(2): 311 - 315.2015

312

Journal of the Botanical Research Institute of Texas 9(2)

Macbride (1916) had noted the similarity of American plants to E. nanum of Europe and described a dif¬

ference in the flange of the nutlets, which Johnston (1924) was unable to see. Johnston did remark that the

North American material was to him more canescent that E. nanum.

Lechner-Pock (1955-1956), who was familiar with E. nanum through her monographic study of Eritrichi-

um, treated E. nanum as European and distinct from all taxa she knew from North America. In the first couplet

of the key on page 103 of her monograph, Lechner-Pock pointed out that the morphology of the ventral surface

of the nutlets of the two European taxa, E. nanum and E.jankae Simonkai (E. nanum subsp. nanum and E. na¬

num subsp .jankae in the treatment of Chaterl972) differ in the position of the attachment of each nutlet: at the

lower 1/3 versus at the base. This is because the nutlet of E. nanum is enlarged below the attachment, creating

a sac-like form, a feature absent from the North American plants (is this the distinct spreading border or flange

of Macbride?). She illustrated these differences (p. 117, figs. 14-19 for E. nanum, p. 119, contrasting figs. 49-54

for E. “ aretioides ”). Therefore, the name E. nanum is misapplied to North American plants.

Cronquist (1984) wrote that the Rocky Mountain plants are “much like var. nanum except in being usu¬

ally more densely hairy; a small difference in the form of the nutlets has been alleged.” He went on to say,

“Plants of var. elongatum from the northern Rocky Mountains (including the type) have the flange of the nut¬

lets entire, but many of those from the southern Rocky Mountains have the flange evidently lacerate-toothed

with segments up to about 0.5 mm long.” He continued, “These more southern plants with lacerate-toothed

nutlets have often been treated as a distinct species, variety, or form ( argenteum ). Although the nutlets provide

many useful taxonomic characters in the Boraginaceae, in this instance the difference does not appear to be

significant. Plants with the flange entire occur throughout the range of the argenteum phase, without any other

obvious differences.” For more than 50 years the Rocky Mountain specimens have mostly been named E. na¬

num var. elongatum, following Cronquist’s taxonomy. Weber (1967 and in all subsequent editions and revisions

of his floras) named the Colorado plants E. aretioides, but this taxonomy was not widely followed.

The type specimens for Eritrichium argenteum (US!) andE. elongatum (NY!) are sufficiently different from

each other that one can conclude, at least at this starting point, that there are two taxa. Problems arise when

determining a large selection of specimens. The forms of the nutlets while expected, a priori, by Macbride

(1916) and initially by me, to be consistent within taxa were not. Therefore various aspects of leaf indumentum

must be evaluated. Having done that, I conclude that the two type specimens represent extremes of a spectrum

of variation. When many specimens are examined, they do fall into two piles representing an argenteum form

and an elongatum form, but with a third pile of intermediates that can only be assigned arbitrarily to one or the

other pile. If one accepts there is one highly variable species in the Rocky Mountains, then at the rank of spe¬

cies, the correct name is E. argenteum W. Wight.

Synonymy

Eritrichium elongatum var. argenteum (W. Wight) I.M. Johnston

Eritrichium elongatum var. paysonii I.M. Johnston

Eritrichium nanum subsp. villosum var. villosum f. argenteum (W. Wight) Brand

Eritrichium aretioides var. elongatum Rydb.

Eritrichium nanum var. elongatum (Rydb.) Cronq.

Eritrichium elongatum (Rydb.) W. Wight [a later homonym of E. elongatum (A.DC] A.DC.

The next question arising is whether Eritrichium argenteum is distinguishable from E. aretioides. The two spe¬

cies are geographically well separated; however, Wight (1902) and Cronquist (1959) both noted the similarity

of E. argenteum to E. aretioides, citing only dfferences in the length of the teeth on the flange of the nutlets to

distinguish them. Nevertheless, on the strength of these features Cronquist (1959) did maintain as a distinct

entity E. nanum var aretioides (Cham.) Herder. I find that conspicuously pustulate leaves of E. aretioides, nutlets

with longer teeth, and, of ourse, geography separate the Alaska-Yukon material as E. aretioides. Additionally, in

a preliminary molecular study, A. Tribsch (pers. comm.) found that E. argenteum is well separated by genetic

markers from E. aretioides.

Murray, Notes on Eritrichium

313

2. Asiatic origins for species of Eritrichium in North America

There is more variation in boreal and arctic species of Eritrichium in North America than previously reported.

Most of the species found in North America show Asian connections through morphology and distribution.

The Asiatic influence is not surprising, inasmuch as throughout the Tertiary, until the late Miocene flooding of

the Bering Strait, and several times in the Quaternary during glacial maxima and lower sea levels, the exposed

Bering Land Bridge provided a dry land connection between Asia and America. During those times Alaska and

Yukon were physically and biotically an extension of Asia, isolated from the rest of North America by Quater¬

nary ice sheets.

Thus it is not unexpected that Eritrichium villosum (Ledeb.) Bunge, a common tundra species on the Chu¬

kotka Peninsula and westward in Russia, is now known in North America from St. Lawrence Island in the

Bering Strait region of Alaska: vicinity of Savoogna based on Carlson 2012-141 (UAAH !) and Boxer Bay based

on Youngl313 (GH!) [reported in Young (1971) as E. aretioides]. Its closest morphological relatives in North

America are E. aretioides and E. chamissonis from which E. villosum is distinct.

Eritrichium splendens Kearney ex W. Wight is endemic to Alaska and Yukon and morphologically in

sharp contrast with its primary congener, E. aretioides, most visibly by its leaf indumentum consisting of stout,

strigose hairs appressed and aligned toward the apex of the leaves, as is typical of the E. sericeum s.l. complex

as elaborated by Ovchinnikova (2001, 2008) in her study of the Asian members.

Johnston (1924) and then Hulten (1948) pointed out the similarity of E. splendens of Alaska to the Central

Asian E. rupestre (Pallas) Bunge ( =E . altaica Popov) and also to E. pectinatum (Pall.) DC., which is found further

eastward in Siberia, all belonging to the E. sericeum s.l. complex. Gjaerevoll (1967) also compared his collec¬

tion of E. splendens from Alaska with E. rupestre and concluded that the two species were distinct. Scoggan

(1979), on the other hand, named our plants E. rupestre with E. splendens in synonymy.

Ovchinnikova (2001,2008) included E. splendens in her reviews of E. sericeum s.l., thus she recognized its

overall similarity to what she has studied in northeastern Asia, but she maintained E. splendens distinct from

the Asian taxa at the rank of species. Eritrichium splendens should be regarded as but one taxon in a complex of

similar entities in Asia and America for which the stout, strigose hairs are a defining feature.

As collections of Eritrichium with strigose hairs appeared in herbaria they simply had been hied with E.

splendens. Specimens accumulated to the point it become clear that there was more than just E. splendens to be

understood. Examination of these specimens led to the recognition of E. arctisibiicum (V.V. Petrovsky) Khokhr.

(E. sericeum var. arctisibiricum V.V. Petrovsky), new to North America.

Eritrichium arctisibiricum is now known from a locality on the Porcupine River in the boreal interior of

Alaska, at three localities in arctic Alaska, on the Shaviovik River, at Prudhoe Bay, and at Atqasuk (Meade

River) and also in northwestern Yukon. The original description by Petrovsky was based on specimens from

Wrangel Island, but he mapped it as occurring elsewhere, especially in the arctic European Russia, (cf. Petro¬

vsky 1980). The strigose hairs on the leaves are finer than those on E. splendens, but appressed and with strict

orientation toward the leaf apices. The clear differentiation of oblanceolate blade and narrow petiole is a distin¬

guishing feature of E. arctisibiricum as opposed E. splendens, which shows almost no discontinuity between

blade and petiole.

3. Clarification of minor points with respect to the holotype for Eritrichium splendens Kearney ex W.

Wight

There are minor discrepancies between what is on the label of the type specimen for Eritrichum splendens at US

and what was published by Mendenhall (1902) with respect to both the collector and date of collection. The

specimen was not collected by Mendenhall as had been written on the specimen label. As Hulten (1940)

pointed out, the type specimen selected by Wight (1902) is one collected by W.L. Puto, which is clearly stated

in Mendenhall (1902:58)

The locality on the label is given as Old Man Creek, which was the local name given to the Kanuti River

during the period 1898-1913 (Orth 1967). In Mendenhall’s report, p. 64 the account is “Kanuti River. Collected

about 15 miles below Caribou Mountain. July 6,” not July 9 as on the label. The map in Mendenhall (1902)

314

Journal of the Botanical Research Institute of Texas 9(2)

gives the major features, a scale, and the dates and places the expedition stopped. They reached a point roughly

15 miles from Caribou Mountain on July 6, and I take that date of collection to be correct.

4. Eritrichium chamissonis DC.

Eritrichium aretioides var. chamissonis (DC.) V.V. Petrovsky

Eritrichium nanum var. chamissonis (DC.) Herder

Lectotypihcation of the name E. chamissonis and its application have been discussed by Murray (2013). Where¬

as E. chamissonis has been viewed by some as a hybrid between E. villosum and E. aretioides (Popov 1953) or as

merely an ecological modification of E. aretioides (Johnston 1924), this species has features that are sufficiently

constant to distinguish it from other taxa. Furthermore, it has a distinct geography and does not occur sporadi¬

cally within the range of E. aretioides as would be expected if it were simply an expression of local conditions.

The arctic collections of E. chamissonis reported by Wiggins and Thomas (1962) and Cody (1996) are compact

ecological modifications of E. aretioides.

Treatment of E. chamissonis at the rank of species (vs being subsumed under E. aretioides) rests on differ¬

ent chromosome numbers—2n=24 for E. aretioides and 2n=48 for E. chamissonis —and the differences in sculp¬

turing of the nutlet epidermis as revealed by preliminary SEM studies (Garroutte et al. 2010). With the excep¬

tion of St. Lawrence Island, Alaska, the type locality for both species, E aretioides and E. chamissonis are other¬

wise allopatric. The best held characteristic lies in whether the bowers just reach the surface of the leaf mass or

are on stems exerted beyond it. Variation in leaf shape and indumentum reduce key characteristics to a few.

This couplet may be helpful:

Flowers (3)4-8(10), exerted beyond the mass of basal leaves; leaves oblanceolate, conspicuously pustulate, leaf blades

proximally glabrate, distally villous on blades and along margins, marginal hairs to 2 mm, seldom obscuring the termi¬

nal leaves, apices acute or obtuse_ Eritrichium aretioides

Flowers 1 -2(3), rarely exerted beyond the mass of basal leaves; leaves, broadly oblanceolate to obovate, sparingly pustu¬

late, leaf blades villous, densely so at apices especially along margins, marginal hairs to 3 mm, often obscuring the

terminal leaves, apices obtuse to rounded_ Eritrichium chamissonis

5. The disposition of Eritrichium howardii (A. Gray) Rydb.

Eritrichium howardii is a narrow endemic of the Rocky Mountain region (Montana and Wyoming) that is dis¬

tinct from the common, widespread E. argenteum found in the region. By its indumentum, E. howardii is allied

with E. sericeum s.l., therefore its closest relatives in North America are in Alaska and Yukon, but one cannot

rule out that it is even more closely related genetically to Asian species.

Given the diversity of species in Asian Eritrichium and relatively few species in North America, one could

reasonably presume a movement from west to east. Furthermore, when the Boras of Chukotka and Alaska are

compared, movement from Asia to America is well demonstrated. Since E. howardii is so morphologically and

ecologically distinct and geographically isolated, it is not likely to have arrived nor arisen recently. It or its an¬

tecedents have most likely come from E. sericeum s.l. prior to the Quaternary.

Weber (1965, 2003) has sought to explain extraordinary disjunctions of taxa in the mountains of north¬

eastern Asia and northwestern America, postulating a Late Tertiary Bora of which E. howardii or an antecedent

were a part, essentially continuous from Asia to America, but which became dissected and greatly reduced in

area by Quaternary events, e.g., the Ice Ages.

There is no direct evidence for these scenarios; they are speculation but well-reasoned. This species, and

for that matter the entire genus, is an excellent candidate for a phylogeographic study since molecular genetics

might provide answers to questions such as where and when this species arose.

ACKNOWLEDGMENTS

My thanks to Peter Lesica (MTU) for providing me with specimens of Eritrichium from Montana, for his read¬

ing of the manuscript, and for his recommendations for improvement, to Tim Hogan (COLO) for specimens

from Colorado, Matthew Carlson (UAAH) for his collection from St. Lawrence Island, Alaska,. Kanchi N.

Murray, Notes on Eritrichium

315

Gandhi has, as always, been extremely helpful with matters of nomenclature. I thank Svetlana Ovchinnikova

(NSK) and Vladik Petrovsky and Volodya Razzhivin (LE) for responding to my many questions about the ge¬

nus Eritrichium in northeastern Asia.

REFERENCES

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Cody, WJ. 1996. Flora of the Yukon Territory. NRC Research Press, Ottawa, Canada.

Coulter, J.M. & A. Nelson. 1909. New manual of botany of the Central Rocky Mountains. Amer. Book Co., N.Y.

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Gandhi, K.N. & D.F. Murray. 2013. A note on " Eritrichum Schrad. 1819" and Eritrichium Schrad. ex Gaudin 1828

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Weber, W.A. 1967. Rocky Mountain Flora. Univ. Colorado Press, Boulder, Colorado, U.S.A.

Weber, W.A. 2003.The Middle Asian element in the southern Rocky Mountain flora of the western United States: a critical

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Wight, W.F. 1902. The genus Eritrichum in North America. Bull.Torrey Bot. Club 29:407-414.

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316

Journal of the Botanical Research Institute of Texas 9(2)

BOOK REVIEW

Thor Hanson. 2015. The Triumph of Seeds: How Grains, Nuts, Kernels, Pulses, & Pips Conquered the

Plant Kingdom and Shaped Human History. (ISBN-13: 978-0-465-05599-9, hbk). Basic Books, 250

West 57 th St, New York, New York 10107, U.S.A. (Orders: www.basicbooks.com, 1-800-343-4499).

$26.99 US, 304 pp., 5.75" x 8.5".

The book we tend to pick up is mostly the book with an eye-catching cover design and an intriguing title. A

clear validation of such an axiom is Nicole Caputo’s book cover design and Thor Hanson’s The Triumph of Seeds:

How Grains, Nuts, Kernels, Pulses, & Pips Conquered the Plant Kingdom and Shaped Human History.

According to Hanson, the dramatic triumph of seeds poses an obvious question: Why are they so success¬

ful? What traits and habits have allowed seeds, and the plants that bear them, to thoroughly transform our

planet? The answers frame the narrative of Hanson’s work and reveal not only why seeds thrive in nature but

also why they are so vital to us. In the table of contents alone, we learn that seeds nourish, seeds unite, seeds

endure, seeds defend, and seeds travel.

Hanson delivers botanical information with wit and imagination. The text uses only common names of

plants. However, there is a complete list of common and scientific names at the end of the book (Appendix A).

In addition, there are extensive notes to each chapter, a glossary, bibliography, and an index. The Triumph of

Seeds is an excellent addition to the single-issue science genre. It describes how little seeds shaped history in

big ways and why the future of seeds is such a hot button issue.

When are you going to plant The Triumph of Seeds in your library?— Kay M. Stansbery, Ph.D., retired librar¬

ian and BRIT volunteer, Fort Worth, Texas, U.S.A.

Thor Hanson is a conservation biologist, Guggenheim Fellow, Switzer Environmental Fellow, and winner of

the John Burrough Medal for excellence in nature writing and natural history. The author of Feathers and The

Impenetrable Forest, Hanson lives with his wife and son on an island in Washington State.

J.Bot. Res. Inst. Texas 9(2): 316.2015

CHANGE IN TAXONOMIC RANK FOR A HEXASTYLIS (ARIST OLO CHI ACE AE)

TAXON OF THE SOUTHEASTERN UNITED STATES

Brian R. Keener

University of West Alabama (UWAL)

Department of Biological & Environmental Sciences

Livingston, Alabama 35470, U.S.A.

bkeener@uwa.edu

and

Botanical Research Institute of Texas (BRIT)

1700 University Drive, Fort Worth, Texas 76107, U.S.A.

L.J. Davenport

Samford University (SAMF)

Department of Biological & Environmental Sciences

Birmingham, Alabama 35229, U.S.A.

ljdavenp@samford.edu

ABSTRACT

A taxonomic innovation is proposed: Hexastylis harperi (Gaddy) B.R. Keener & T J. Davenp., comb. et. stat. nov. This new species is dis¬

tinct in morphology, habitat, and distribution and shows no intergradation with more widespread and typical, putatively conspecific

populations.

RESUMEN

Se propone una innovacion taxonomica: Hexastylis harperi (Gaddy) B.R. Keener & L.J. Davenp., comb. et. stat. nov. Esta nueva especie es

distinta en morfologia, habitat, y distribucion y no muestra intergradacion con poblaciones mas generalizadas y tipicas, putativamente

conspecificas.

In preparing for the writing of a taxonomic manual for the flora of Alabama, it will be necessary to make

changes in ranks of taxa to address the biodiversity represented. As the Flora of Alabama project continues,

there will undoubtedly be future papers for additional issues as they arise. This paper is the first such

installment.

During June 1927, Roland Harper discovered a plant resembling Hexastylis shuttleworthii (Britten & Baker f.)

Small in a swamp in central Alabama (Autauga County); in 1935, he found a similar-appearing plant in north¬

west Alabama’s Marion County (Harper 1936). Gaddy (1987a) later described Harper’s discovery as a variety of

Hexastylis shuttlew or thii (H. shuttlew or thii var. harperi Gaddy) from the upper Coastal Plain of Alabama and

Georgia. Variety harperi has long rhizomes, which support leaves along their entire lengths, and grows at bog

edges and in acidic hammocks of that Coastal Plain. In contrast, var. shuttlew or thii has short rhizomes with

solely terminal leaves and grows along upland, rocky, forested slopes of the Blue Ridge, Ridge & Valley, and

Appalachian Plateau ecoregions (see Gaddy 1987b and Whittemore & Gaddy 1997 for further details).

The two varieties of Hexastylis shuttlew or thii are distinct in morphology, habitat and distribution. They

are completely allopatric, showing none of the geographical gradations associated with the concept of varieties.

For these reasons, we maintain that species rank is warranted.

Further evidence supporting this change was recently published in broader evolutionary investigations of

Asarum s.l. (Sinn et al. 2015a, 2015b). In both studies, and using different analyses, the authors found that the

two vars. of Hexastylis shuttlew or thii (as Asarum shuttlew or thii Britten & Baker, f) were polyphyletic rather than

“sister” to each other, as would be expected.

It is worth noting that some modern workers (Barringer 1993; Kelly 1997,1998; Sinn et al. 2015a, b) prefer

to subsume Hexastylis into a broadly circumscribed Asarum, but we here follow the generic circumscriptions

used in two recent and influential continental and regional floras, the Flora of North America (Whittemore &

Gaddy 1997) and the Flora of the Southern and Mid-Atlantic States (Weakley 2015).

Hexastylis harperi (Gaddy) B.R. Keener & L.J. Davenp., comb, etstat. nov. Basionym: Hexastylis shuttleworthii (Britten

& Baker E) Small var. harperi Gaddy, Sida 12:54.1987. Type: U.S.A. GEORGIA. Madison Co.: acidic bog under Acer rubrum, Lirioden-

J. Bot. Res. Inst. Texas 9(2): 317 - 318.2015

318

Journal of the Botanical Research Institute of Texas 9(2)

dron tulipifera, and Nyssa sylvatica var. biflora; growing on small hammocks with Osmunda cinnamomea and along bog edges with

Medeola virginiana and Isotria verticillata just N of GA 106, 14.2 mi NE of Athens, 9 May 1986, Gaddy s.n. (holotype: CLEMS; iso¬

types: AUA, GA, GH, MO, NCU, NY, TENN, UNA, VDB).

Hexastylis speciosa R.M. Harper (Harper 1924) and H. harperi have both been called “Harper’s Heartleaf” or

“Harper’s Ginger.” We recommend that “Harper’s Heartleaf” or “Harper’s Ginger” be used for H. harperi, since

those common names are direct translations of the scientific name. For H. speciosa, “Alabama Ginger” and

“Alabama Heartleaf” may be more applicable common names, due to the species’ restriction to central

Alabama.

ACKNOWLEDGMENTS

We thank our reviewers, Alan Weakley and Guy Nesom, for their very timely and helpful suggestions.

REFERENCES

Barringer, K. 1993. New combinations in North American Asarum (Aristolochiaceae). Novon 3:225-227.

Gaddy, L.L. 1987a. Hexastylis shuttleworthii var. harperi (Aristolochiaceae), a new variety of heartleaf from Alabama and

Georgia. Sida 12:51-56.

Gaddy, L.L. 1987b. A review of the taxonomy and biogeography of Hexastylis (Aristolochiaceae). Castanea 52:186-196.

Harper, R.M. 1924. A new heart-leaf and other interesting plants from Autauga County, Alabama. Torreya 24:77-82.

Harper, R.M. 1936. Asarum and Hexastylis in Alabama and neighboring states. Castanea 3:19-24.

Kelly, L.M. 1997. A cladistic analysis of Asarum (Aristolochiaceae) and implications for the evolution of herkogamy. Amer.

J. Bot. 84:1752-1765.

Kelly, L.M. 1998. Phylogenetic relationships in Asarum (Aristolochiaceae) based on morphology and ITS sequences.

Amer. J. Bot. 85:1454-1467.

Sinn, B.T., L.M. Kelly, & J.V. Freudenstein. 2015a. Phylogenetic relationships in Asarum : Effect of data partitioning and a

revised classification. Amer. J. Bot. 102:765-779.

Sinn, B.T., L.M. Kelly, & J.V. Freudenstein. 2015b. Putative floral brood-site mimicry, loss of autonomous selfing, and reduced

vegetative growth are significantly correlated with increased diversification in Asarum (Aristolochiaceae). Molec.

Phylogen. Evol. 89:194-204.

Weakley, A. S. 2015. Flora of the southern and mid-Atlantic states; May 2015 version, http://www.herbarium.unc.edu/

flora.htm.

Whittemore, A.T. & L.L. Gaddy. 1997. Hexastylis. In: Flora of North America Editorial Committee, Flora of North America

north of Mexico. Volume 3, Magnoliophyta: Magnoliidae and Hamamelidae. Oxford Univ. Press, New York, NY. Pp

54-58.

CALATHEA GALDAMESIANA (MARANTACEAE), A NEW ENDEMIC

PANAMANIAN SPECIES

Helen Kennedy

UCR Herbarium, Dept, of Botany & Plant Sci.

University of California Riverside

Riverside, California 92521, US.A.

ganders@mail.ubc.ca

Rodolfo Flores

Smithsonian Tropical Research Institute

Avenida Roosevelt, Edificio Tupper 401

Balboa, Ancon, Rep. de PANAMA

floresr@si.edu, rfloresn84@hotmail.com

ABSTRACT

Calathea galdamesiana H. Kenn. & R. Flores, endemic to Panama, is described as new for inclusion in the Flora Mesoamericana. It occurs

in premontane wet forest and is known only from the type locality in Parque Nacional Santa Fe, Veraguas Province. It is characterized by the

5-8 basal leaves per shoot, in which the petiole is absent and the margins of the leaf sheath are spreading and reflexed; and the 1-4 fusiform

inflorescences borne on a separate, leafless shoot directly from the rhizome. Calathea galdamesiana shares a similar vegetative habit and leaf

morphology with C. panamensis Standi, but differs in the elliptic vs. obovate to obovate-elliptic leaf blade, the 1-4 inflorescences borne on a

separate leafless shoot vs. 1 (rarely 2) borne on the leafy shoot, the red-purple vs. green bracts and chasmogamous vs. cleistogamous flowers.

The flower morphology of C. galdamesiana is most similar to that of C. cleistantha Standi, which also has the inflorescences borne on a sepa¬

rate leafless shoot. Calathea galdamesiana differs from C. cleistantha by the absence of a petiole and the margins of the leaf sheath spreading

to recurved vs. petiole (0-)3.5-15 cm and the margins of leaf sheath clasping, the usually shorter pulvinus (0.5-1 vs. 0.9-2.3 cm), and the

fusiform vs. ovoid inflorescence (length to width ratio 3.2-4.5T vs. <2.5:1).

RESUMEN

Calathea galdamesiana H. Kenn. & R. Flores, endemica de Panama, es descrita como nueva para inclusion en Flora Mesoamericana. Esta

especie se ecuentra en el bosque humedo premontano y esta conocida de solo de la localidad del tipo en Parque Nacional Santa Fe de Provin-

cia Veraguas. Calathea galdamesiana se caracteriza por tener 5-8 hojas basales por brote y la ausencia de peciolo y las margenes de la vaina

son extendidos a recurvados; y inflorescencias 1-4, fusiformes, en un brote aparte, sin hojas, que sale directamente del rizoma. Calathea

galdamesiana tiene habito similar a C. panamensis Standi., pero se distingue por sus laminas foliares elipticas vs obovadas u obovado-elipti-

cas, las inflorescencias 1-4, en un brote aparte, sin hojas vs inflorescencia 1 (raramente 2) en un brote con hojas, las bracteas rojo-purpureas

vs. verdes, y las flores casmogamas vs. cleistogamas. Fa forma de las flores de C. galdamesiana es mas similar a C. cleistantha Standi., la cual

tambien tiene las inflorescencias en un brote aparte, sin hojas. Calathea galdamesiana difiere de C. cleistantha Standi., por la ausencia del

peciolo y los margenes de la vaina extendidos a recurvados vs. peciolos (0-)3.5-15 cm y los margenes de la vaina abrazados, y el pulvinulo

usualmente mas corto (0.5-1 vs. 0.9—2.3 cm), y las inflorescencias fusiformes vs. ovoides (proporcion largo por ancho 3.2-4.5:l vs. <2.5:1).

In preparation for the Flora Mesoamericana treatment, the species of Marantaceae from Panama have been a

special focus as with increased collecting several undescribed species have been discovered recently. With

considerably more collecting since publication of the Woodson & Schery (1945) treatment for Flora of Pana¬

ma, listing 23 species, the total has tripled. By 1972, Dressier (1972:184) reported a total of 35 species for Pan¬

ama. Besides held work, recent herbarium studies at Missouri Botanical Garden and University of Panama

have uncovered additional new species. Just since 2011, 11 more species of Marantaceae have been described

from Panama. Kennedy (2012:49) had reported a total of 63 species whereas, currently, 69 species are recog¬

nized (a 200% increase from the original Flora of Panama treatment). Twenty species are recognized as en¬

demic, including the one described herein plus another as yet undescribed taxon.

TAXONOMIC TREATMENT

Calathea galdamesiana H. Kenn. & R. Flores, sp. nov. (Figs. 1, 2). Type: PANAMA. Veraguas: Parque Nacional Santa Fe,

alrededores de la parcela ubicada por CBMAPII/ANAM, Alto de Piedra, Guabal, 900 m, 8°3T39"N, 81°08 , 58"W (UTM 0483553 E,

0942618 N), 18 May 2015, R. Flores & N. Guerra 3655 (holotype: PMA; isotypes: SCZ, UCH, UCR).

Haec species quoad partes vegetativas Calatheaepanamensi Rowlee ex Standi, similis, sed ab ea lamina foliari elliptica (vs. obovata vel obova-

to-elliptica), inflorescentiis 1 ad 4 (vs. 1 raro 2) surculo aphyllo discreto insidentibus, bracteis rubro-purpureis (vs. viridibus) atque floribus

chasmogamis; quoad partes florales C. cleistanthae Standi, similis, sed ab ea foliis semper sessilibus, vagina florali marginibus patentibus

J. Bot. Res. Inst. Texas 9(2): 319 - 323.2015

320

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 1. Calatheagaldamesiana H. Kenn. & R. Flores. A. Habit. B. Leaf base, pulvinus and apex of alate leaf sheath. C. Inflorescences subtended by cataphyll.

D. Face view of untripped flower. E. Flower on adaxial leaf surface. {Flores & Guerra 3655, PMA, photos by Rodolfo Flores).

Kennedy and Flores, A new endemic Panamanian species of Calathea

321

Fig. 2. Calatheagaldamesiana H. Kenn. & R. Flores. Holotype. Type scan provided by PMA {Flores & Guerra 3655, PMA).

322

Journal of the Botanical Research Institute of Texas 9(2)

usque recurvis (vs. amplexicaulibus), pulvino plerumque breviore (0.5-1 vs. 0.9-2.3 cm longo) atque inflorescentia fusiformi (vs. ovoidea)

longitudinis cum latitudine proportione 3.2-4.5 (vs. ca. 2.5) distinguitur.

Plants rhizomatous, rosulate herbs, 50-80 cm; the inflorescences are borne on a leafless shoot directly from

the rhizome; roots bearing swollen “root tubers” 43-6.5 x 2-3 cm; cataphylls narrowly ovate, apiculate, green.

Leaves 5-8 basal; leaf sheath alate, the wings spreading, recurved, the abaxial surface, both wings and central

back portion, light green, adaxial portion of wings deep green, central portion of leaf sheath lighter, 18-48 cm;

petiole absent; pulvinus light yellow-green to cream-colored, minutely tomentose adaxially, 0.5-1 cm; leaf

blade herbaceous, with pleated appearance (tissue between two adjacent veins planar), elliptic, apex obtuse to

rounded with acumen, base obtuse to rounded, 23.5-46.2 x 19-25.5 cm (length:width ratios [1.34—] 1.81—

2.16:1) lateral veins 11 to 14 per 3 cm (measured at midpoint of each side of blade), vein angles from midrib

35°-42°, measured at midpoint of blade, adaxial surface grass-green, midrib yellow-green, noticeably lighter

than blade, abaxial surface pale grey-green, midrib cream-colored. Inflorescences 1-4 per shoot, the brst ter¬

minal, subsequent ones in the axil of the subtending cataphyll, imbricate, fusiform, 5.8-10.8 x 1.3-2.5 cm;

peduncle red-purple, white if covered by cataphyll, 2.4-6.5 cm, the basal portion, ca. 2 cm, swollen, tissue

growth more on one side, causing the inflorescence to be deflexed onto the ground. Bracts 6-12, spirally ar¬

ranged, ovate, apical ones proportionally longer and narrower, apex acuminate, 4-5.9 cm long, abaxial surface

of bracts red-purple to brownish purple, darker at margins and apex, becoming very dark, almost blackish

purple, in age. Flowers open spontaneously. Sepals red-purple apically, basal half white, ca. 19 mm long. Co¬

rolla tube cream-colored to white, ca. 35 mm; corolla lobes subequal, elliptic, apex obtuse, red-purple abaxi-

ally, white adaxially with the veins seen as bne purple lines, 17-19 x 5-6 mm. Staminodes 3; outer staminode

obovate, apex emarginate, yellow, ca. 14-17 x 8 mm; callose staminode totally callose, apex reflexed, obtuse

with minute acumen to rounded or somewhat irregular, yellow basally, apical 3 dark red-purple, ca. 7 mm

wide; cucullate staminode yellow, ca. 10 mm; stamen pale yellow with lateral petaloid appendage 1-3 mm

wide; ovary pink, ca. 3mm. Fruits and seeds unknown.

Additional specimens: PANAMA. Veraguas: Parque Nacional Santa Fe, alrededores de la parcela ubicada por CBMAPII/ANAM, cerca a la

estacion de ANAM, Alta de Piedra, Guabal, E483553 N942618,3 Sep 2014, R. Flores, R. Vergara, R. Carranza&J. Aguirre 3654 (MO, PMA).

Distribution and habitat.—Calathea galdamesiana is endemic to Panama. It is known only from the type locality

in Parque Nacional Santa Fe, Veraguas Province. It occurs in premontane wet forest habitat. The only collec¬

tions of it were at 900 m elevation.

Discussion.—Calathea galdamesiana belongs to Calathea section Breviscapae Benth. It is characterized by

the 5-8 basal leaves per shoot, the elliptic leaf blades, the absence of a petiole proper, the margins of the leaf

sheath are spreading and reflexed, the 1-4 fusiform inflorescences borne on a separate, leafless shoot directly

from the rhizome, the petals red-purple abaxially, the outer and cucullate staminodes yellow and callose sta¬

minode yellow basally, apical % red-purple. Calathea galdamesiana shares a similar vegetative habit and leaf

morphology with C. panamensis Standi, but differs in the elliptic vs. obovate to obovate-elliptic leaf blade, the

1-4 inflorescences borne on a separate leafless shoot vs. 1 (rarely 2) borne on the leafy shoot, the red-purple vs.

green bracts and chasmogamous vs. cleistogamous flowers. The uncommon leaf morphology, shared with C.

panamensis (alate, spreading, marginally recurved leaf sheath and lack of petiole) together with the inflores¬

cences borne on a separate, leafless, shoot distinguish C. galdamesiana not only from other Panamanian spe¬

cies but from all Central American species and probably South American as well. The Brazilian species, C. albo-

vaginata K. Schum., also has an alate leaf sheath, but there, the inflorescence is borne on the leafy shoot. The

flower morphology of C. galdamesiana is most similar to that of C. cleistantha Standi, which also has the inflo¬

rescences borne on a separate leafless shoot. Calathea galdamesiana differs from C. cleistantha by the absence of

a petiole and the margins of the leaf sheath spreading to recurved vs. petiole (0-)3.5-15 cm and the margins of

leaf sheath clasping, the usually shorter pulvinus (0.5-1 vs. 0.9-2.3 cm), and the fusiform vs. ovoid inflores¬

cence (length to width ratio 3.2-4.5T vs. <2.5:1).

The presence of the swollen “root tubers” in C. galdamesiana together with the thin textured, plicate leaf,

Kennedy and Flores, A new endemic Panamanian species of Calathea

323

are commonly found in deciduous species such as C. panamensis, C. latifolia (Willd. ex Link) Klotzsch and C.

macrosepala K. Schum. Whether this new species is possibly deciduous is not yet known.

Calathea micans (L. Mathieu) Korn., C. microcephala (Poepp. & Endl.) Korn, and C.fucata H. Kenn., in

Calathea section Microcephalum Benth. also exhibit the swollen, and subsequent unequal-sided growth, of the

base of the peduncles observed in C. galdamesiana. The growth of the base of the peduncle acts to deflex the

inflorescence to the ground, usually when in fruit, in the case of C. micans and the related species, which are

dispersed by ants (Horvitz & Beattie 1980). It is possible this is also the case in C. galdamesiana and merits

further observation in the held.

Etymology .—The specific epithet, galdamesiana, is in honor botanist Carmen Galdames, research assis¬

tant at SCZ herbarium, Smithsonian Tropical Research Institute, Ancon, Panama. It is a pleasure to thus recog¬

nize her contribution to our knowledge of the Panamanian Bora through her collections, photographs, co-au-

thorship of the Catalogo de las Plantas Vasculares de Panama, her ethnobotanical studies and her generous

assistance to the authors in the SCZ herbarium, including sharing her botanical knowledge.

ACKNOWLEDGMENTS

We acknowledge the functional and logistic support of the Ministry for the Environment (Miambiente in

Spanish) with regards to the Mesoamerican Biological Corredor of the Panamanian Atlantic watershed (Loan

Contract No. BIRF 7439-PAN and donation agreement for World Environment (FMAM) No. GEF TF 056628)

in light of implementing the Biological Diversity Natural System of information and Monitoring (SNIMDB). We

equally acknowledge the support of the Ministry for the Environment for the necessary permits for collection.

We thank the staff of the PMA Herbarium for the scan of the type. We appreciate the help of the David Mitre of

SNIMDB for managing and streamlining the permits to make the specimens available for study. We acknowl¬

edge the Smithsonian Tropical Research Institute (STRI) through the Nagoya Protocol Project for support to

one of the authors (RF). Last but not least, we acknowledge the help of Alberto S. Taylor, Professor Emeritus of

Botany, University of Panama, for the Spanish translation, Roy Gereau for the Latin diagnosis, and Barney

Lipscomb and the three anonymous reviewers for their helpful corrections and comments.

REFERENCES

Dressler, R.L. 1972. Terrestrial plants of Panama. Bull. Biol. Soc. Wash. 2:179-186.

Horvitz, C.C. & AJ. Beattie. 1980. Ant dispersal of Calathea (Marantaceae) seeds by carnivorous ponerines (Formicidae) in

a tropical rain forest. Amer. J. Bot. 67:321-326.

Kennedy, H. 2012. Calathea rhizanthoides and C. peregrina (Marantaceae), new species endemic to Panama. J. Bot. Res.

Inst. Texas 6:49-54.

Woodson, R.E., Jr. & R.W. Shery. 1945. Marantaceae. In Flora of Panama. Ann. Missouri Bot. Gard. 32:81-105.

324

Journal of the Botanical Research Institute of Texas 9(2)

BOOK REVIEW

Sylvan T. Runkel & Dean M. Roosa. 2014. Wildflowers and Other Plants of Iowa Wetlands. Second Edition.

(ISBN-13: 978-1-60938-285-8, pbk). University of Iowa Press, 119 W. Park Road, 100 Kuhl House, Iowa

City, Iowa 52242-1000, U.S.A. (Orders: www.uiowapress.org, 1-800-621-2736, uipress@uiowa.edu).

$35.00,373 pp., color photos, 6" x 9".

Though the original nomenclature has not changed, this second edition guide includes updated scientific and

common names from Flora of North America North of Mexico and Steyermark’s Flora of Missouri and has all new

color photographs. The most noticeable change, however, is to the overall design of the book, including a

switch from a landscape 8.75" x 5.75" orientation to a portrait 6" x 9" orientation. This has allowed for the ma¬

jority of the plant photos to occupy an entire page, a big boost from the first edition that displayed many verti¬

cal images at a (now) mere 4.25" x 5.75". Plus the new portrait orientation is simply more practical for a held

guide; the increased binding area should translate to greater durability. The page layout is more friendly, with

the standard plant metadata now arranged in a sidebar: common names, etymology, family name (including

updates), and wetland status according to the 2013 National Wetland Plant List (another new feature). The

main text of each species page now contains a cohesive narrative, with a separate “Notes” section at the end for

any updated information since the last edition. As a whole, the updates made to this second edition were

thoughtful and in hindsight perhaps much-needed. I recommend that all owners of the first edition go ahead

and purchase this newest release. You won’t be sorry.— Brooke Best, Ph.D., Botanist and Editor, Botanical

Research Institute of Fexas, Fort Worth, Fexas, U.S.A.

From the Publisher: Originally published in 1999, Wildflowers and Other Plants of Iowa Wetlands was the first

book to focus on the beauty and diversity of the wetland plants that once covered 1.5 million acres of Iowa.

Now this classic of midwestern natural history is back in print with a new format and all-new photographs,

just as Iowa’s wetlands are getting the respect and attention they deserve.

In clear and accessible prose, authors Sylvan Runkel and Dean Roosa provide common, scientific, and

family names; the Latin or Greek meaning of the scientific names; habitat and blooming times; and a complete

description. Plants are presented by habitat (terrestrial or aquatic), then refined by habit (e.g., emergent, float¬

ing, or submerged) or taxonomic group (e.g., ferns and allies or trees, shrubs, and vines). Particularly interest¬

ing is the information on the many ways in which Native Americans and early pioneers used these plants for

everything from pain relief to tonics to soup and the ways that wildlife today use them for food and shelter.

Each of the more than 150 species accounts is accompanied by a brilliant full-page color photograph by bota¬

nist Thomas Rosburg, who has also updated the nomenclature and descriptions for certain species.

After decades of being considered an enemy of the settler, the farmer, and the citizen, Iowa’s wetlands

have come into their own. We are finally caring for these important habitats. Runkel and Roosa’s updated held

companion will be a valuable guide to today’s preservation and restoration initiatives.

J.Bot. Res. Inst. Texas 9(2): 324.2015

CASEARIA DRAGANAE , A NEW SPECIES OF SAMYDACEAE FROM

WESTERN COLOMBIA AND ECUADOR

Mac H. Alford

Department of Biological Sciences

University of Southern Mississippi

Hattiesburg, Mississippi 39402, U.S.A.

mac.alford@usm.edu

ABSTRACT

A new species of Casearia (Samydaceae), C. draganae M.H. Alford, from lowland rainforest of the Pacific basin of Colombia and Ecuador is

described and illustrated. The new species is vegetatively similar to Casearia fasciculata (Ruiz & Pav.) Sleumer but differs in petiole length,

distribution of pellucid-punctations in the lamina, flower size, ovary pubescence, and pericarp thickness.

RESUMEN

Se describe e ilustra una nueva especie de Casearia (Samydaceae), C. draganae M.H. Alford, del bosque inundable de la vertiente del Pacifica

de Colombia y Ecuador. Por sus hojas, la nueva especie es similar a Casearia fasciculata (Ruiz & Pav.) Sleumer pero difiere en la longitud del

peciolo, la distribucion de punteaduras traslucidas en la lamina, el tamano de las flores, la pubescencia del ovario y el grosor del

pericarpio.

Key Words: Casearia, Casearia sect. Casearia species group ‘Singulares,’ Colombia, Ecuador, Flacourtiaceae, Salicaceae, Samydaceae

INTRODUCTION

Casearia Jacq. includes about 180-200 species distributed pantropically and is the largest genus of Samydaceae

(Alford 2005, 2007), or the second or third largest genus of a broadly circumscribed Salicaceae (Chase et al.

2002; APG 2009). It is a common genus of tropical forests worldwide (cf. Phillips & Miller 2002), but speci¬

mens are notoriously difficult to identify to species due to the small size of the flowers, usually <5 mm and oc¬

casionally <2 mm long, and the importance of microscopic floral features such as staminodes and anther con¬

nectives in the keys of major treatments (e.g., Sleumer 1980). Fruits are rather variable in the genus, but few

treatments offer keys that utilize fruit characters (e.g., Olson et al. 1999; Alford 2003; Zmarzty 2007).

Most species of Casearia belong to section Casearia, which was divided into six informal “species groups”

by Sleumer (1980). According to Sleumer (1980), Casearia sect. Casearia species group ‘Singulares’ consists of

two species, C. fasciculata (Ruiz & Pav.) Sleumer and C. combaymensis Tulasne, but with additional collections

of the last 35 years, especially from Ecuador, Peru, and the Guiana Shield, his treatment is now inadequate. In

addition, the type specimen of C. combaymensis is in conflict with his description and instead represents a spe¬

cies in Casearia sect. Casearia species group ‘Decandrae’ (see Mori & Fischer 2002 for details). Several new

species of species group ‘Singulares’ are being prepared for description, and several additional species with

names placed in synonymy by Sleumer (1980) will again be applied in an upcoming revision (M.H. Alford, in

prep., see also Mori & Fischer 2002). One of the new species is described here. It is remarkably similar to C.

fasciculata vegetatively but has shorter petioles, larger flowers, and fruits with thicker pericarps. Casearia fas¬

ciculata, the type of which is from Chinchao (Huanuco), Peru, is distributed on the eastern slopes of the Andes

and in Amazonia; the new species described here is distributed in the lowlands west of the Andes along the

Pacific coast in the Choco biogeographic region.

TAXONOMIC TREATMENT

Casearia draganae M.H. Alford, sp. nov. (Fig. 1). Type: COLOMBIA. Valle del Cauca: Bajo Calima; Concesion Pulpapel/Bue¬

naventura, bosque pluvial tropical, 03°55'N, 77°00'W, 100 m, 23 Nov 1984 (ft, fr.), M. Monsalve B. 541 (holotype: COL).

Casearia draganae differs from Casearia fasciculata (Ruiz & Pav.) Sleumer in its shorter (1-5.5 mm) petioles, larger flowers (sepals 5.0-5.4

mm long), densely hispid ovary, and fruits with thicker (1.2-3 mm) pericarp.

J. Bot. Res. Inst. Texas 9(2): 325 - 329.2015

326

Journal of the Botanical Research Institute of Texas 9(2)

2 mm

5 mm

3 mm

4 cm

Fig. 1. Casearia draganae M.H. Alford. A. Fruiting branch, the upper two leaves showing the abaxial surface and highlighting the raised venation, the

other leaves showing the adaxial surface, one highlighting the tertiary venation perpendicular to the midvein. B. Leaf margin, showing the obscure

leaf teeth, each with a small deciduous cap ("theoid teeth"). C. Pellucid-punctations and tertiary and finer venation of the lamina, as seen with trans¬

mitted light. D. Detail of branch between two nodes, highlighting pubescence and lenticels. E. Flower, in part, longitudinal section, showing 3 of the

5 sepals and 6 of the 10 stamens. F. Cross-section of the fruit, about Vi empty after drying. A-D drawn from Devia A. & Prado 2386 (MO) and F drawn

from Monsalve B. 510 (MO) by Emily J. Lain. E drawn from Monsalve B. 541 (COL) byTharangamala Samarakoon.

Alford, A new species of Casearia (Samydaceae)

327

Shrubs to small trees, 2.5-8 m tall [and probably taller, as many Casearia reach reproductive maturity at

small height], dbh 2.5-8 cm. Young stems angled, often reddish brown to maroon when dry, puberulent, hairs

sometimes more dense on one of the stem faces, with numerous elliptical to narrowly elliptical lenticels; older

stems gray, glabrescent. Leaves alternate, simple, persistent, broadly elliptic to elliptic to obovate, rarely nar¬

rowly elliptic, (7.5-) 10-21 cm long, (3-) 4-8 cm wide, apex abruptly acuminate for 0.5-3 cm, base acute to

attenuate, infrequently obtuse to rounded, subcoriaceous to coriaceous, light green, slightly darker green

above when dry, glabrous, densely pellucid-punctate and -lineate, serrate, sometimes obscurely so, the serra¬

tions minute, each with a small, conical, deciduous cap, ca. 3 teeth/cm in the middle third of the lamina, 4-6

pairs of lateral secondary veins curved ascending, weakly raised above, midvein more distinctly raised within

a trough in the upper 3 A, midvein and secondaries distinctly raised abaxially and contrasting with lamina,

tertiary veins ± perpendicular to the midvein; petiole glabrous or with a few scattered hairs near base, 1-5.5

mm long, 0.7-1.8 mm in diameter; stipules narrowly lanceolate, 5.5 x 0.8 mm, margins entire, 3-veined, gla¬

brous, pellucid-punctate, quickly caducous. Inflorescences axillary glomerules or fascicles, the glomerules

often appearing like 5-6 small balls of bracts, potentially producing >16 flowers in total [only one flowering

individual seen; 3 flowers were present], bracts numerous and closely arranged, chartaceous, broadly ovate, ca.

2 mm long, beige to light reddish brown, glabrous, ultimate peduncles [pedicels below the articulation sensu

Sleumer 1980] short, slightly thicker than pedicel or expanded just below pedicel, articulation pubescent,

pedicels [pedicels above the articulation sensu Sleumer 1980] 4 mm long, 0.3 mm in diameter, densely pubes¬

cent, pubescence golden when dry. Flower buds ellipsoidal, ca. 2 x 1.3 mm. Flowers perigynous, hypanthium

1.5 mm long, sparsely puberulent outside, mostly glabrous inside, with several vein-like ridges on the outer

surface, corresponding to vasculature of stamens; sepals (4—)5, the free lobes lanceolate-oblong, 3.5-3.9 mm

long (total length, base to apex including hypanthium, 5.0-5.4 mm), 1.6-2.0 mm wide at base of lobes, each

lobe with 3 major veins and several minor ones, densely puberulent on both surfaces, white to greenish-yellow,

hairs drying golden; petals lacking; stamens 10, slightly unequal in length, the longer with filaments 2.0 mm

and opposite sepals, the shorter 1.3-1.4 mm and alternating with sepals, filaments sparsely to moderately pi¬

lose, anthers ellipsoidal, 0.4-0.6 x 0.3-0.5 mm, glabrous, dorsihxed, introrsely dehiscent, yellow, lacking or

with a very small apical extension of the connective, staminodes alternating with stamens, oblong to ligulate,

0.6-0.7 x 0.3 mm, densely pilose; pistil 1, ovary superior, ovoid, 1.5 mm tall, 1.3 mm in diameter, hispid and

verrucose on outer surface, possibly pellucid-punctate, placentation parietal, style stout, 0.75 x 0.4 mm, stigma

capitate, smooth, 0.55 mm in diameter; staminodes and stigma presented at about the same height, just above

the rim of the hypanthium. Fruits globular, probably a tardily dehiscent capsule, 2.4-4.8 cm diameter, peri¬

carp coriaceous, 1.2-2.5 (-3) mm thick, outer surface densely verrucose, dark green turning yellow to yellow-

orange when mature, on a moderately puberulent pedicel, 4 mm thick. Seeds 15 or more, flattened-ellipsoidal

with pellucid-punctation streaks, 1.4 x 0.8-0.9 x 0.2 cm, each covered by a clear (colorless), slimy, sweet aril.

Phenology. —With fruits throughout the year. Flowering specimens are known from late November, early

December, and April; flower buds also known from July.

Distribution and habitat. —Lowland rainforest of the Pacific coasts of Colombia (Narino, Valle del Cauca)

and Ecuador (Esmeraldas, Los Rios). Known from primary and secondary forest, where soil is commonly nu¬

trient poor but high in aluminum, 10-350(-1100?) m elevation. Associates include Attalea, Vismia, Isertia, Ce-

cropia, Miconia, Ammandra, Psychotria, Goupia, Pourouma, Vochysia, Mabea, and Inga. A detailed description of

one of the areas where the species has been collected many times may be found in Faber-Langendoen & Gentry

(1991).

Common Name.—“Caimito” (van Rooden et al. 386).

Etymology. —Named in honor of my wife, Dragana Mladenovic Alford, for enduring and supporting my

research endeavors. In the Serbo-Croatian language, Dragana means “sweetheart,” based on the root “drag,”

meaning “dear,” “beloved,” or “precious” (Benson 1994). Given that fruits of this species are noted to be sweet,

the epithet is appropriate.

Conservation Status. —Probably least concern, as the species sometimes inhabits disturbed habitats. How-

328

Journal of the Botanical Research Institute of Texas 9(2)

ever, the Ecuadorian part of its range has very little of its original forest cover (Rodrlguiez-Mahecha et al.

2004).

Additional specimens examined: COLOMBIA. Depto. Narino: Municipio Barbacoas, corregimiento Junin, sitio La Guaraperia a 3 km de

Junin, viaJunm-Tumaco, 1100 m [?], Mar 1995J.L. Ferndndez et al. 12548 (COL). Depto. Valle del Cauca: Rio Calima, La Trojita, 5-50 m,

19 Feb-10 Mar 1944, J. Cuatrecasas 16646 (US); Buenaventura Municipio, region of Bajo Calima, along road between Buenaventura and

Malaga, at km 50.5, 04°09’N, 77°11'W, <100 m, 7 Feb 1990, T.B. Croat &J. Watt 70325 (MO); San Isidro, via a Malaga, km 51, 30 m, 27 Feb

1990, W. DeviaA. 3019 (MO-2); Buenaventura, Corregimiento Bendiciones, via a Aguaclara, calima medio, Region Fitogeografica del Choco,

120 m, 2 Feb 1989, W. DeviaA. & F. Prado 2386 (MO); Bajo Calima Concession, ca. 15 km NW of Buenaventura, 3.7 km fromjuanchaco gate,

1 km past Luchin/Lijal on Luchin, 03°53'N, 77°10'W, 50 m, 17Jun 1987, D. Faber-Langendoen 960 (MO-2); Bajo Calima Concession, ca. 20

km N of Buenaventura, ca. 300 m NW of “Dindo III, Dindo IV” intersection on “Dindo IV,” 03°59'N, 77°04'W, 50 m, 27Jun 1988, D. Faber-

Langendoen & J.A. Hurtado 1495 (MO), 1521 (MO); Bajo Calima Concession, ca. 20 km N of Buenaventura, behind Pulpapel Forestry Station,

03°56'N, 77°03'W, 50 m, 6 Jul 1988, D. Faber-Langendoen & J.A. Hurtado 1620 (MO-2); Bajo Calima Concession, ca. 25 km NW of Buenaven¬

tura, ca. 9 km NW of San Isidro intersection on “Canalete,” near gate, 03°59'N, 77°08'W, 50 m, 8 Jul 1988, D. Faber-Langendoen &J.A.

Hurtado 1667 (MO), 13 Jul 1988, D. Faber-Langendoen & J.A. Hurtado 1761 (MO); Bajo Calima Concession, ca. 20 km NW of Buenaventura

on San Isidro road, 4 km N of San Isidro-Juanchaco Norte intersection, 03°58'N, 77°07'W, 50 m, 4 Aug 1988, D. Faber-Langendoen & J.A.

Hurtado 1976 (MO), 1978 (MO); Buenaventura Municipio: Bajo Calima Concession ca. 16 km NW of Buenaventura and 1 km fromjuanchaco

gate on Tomar Rd., 04°0FN, 77°10'W, 50 m, 25 Jun 1987, D. Faber-Langendoen & F. Renteria 1116 (MO); Bajo Calima Concession, ca. 20 km

N of Buenaventura, behind Pulpapel Forestry Station, 03°53'N, 77°00'W, 50 m, D. Faber-Langendoen & F. Renteria 1311 (MO); Bajo Calima

Concession, ca. 16 km N of Buenaventura, behind Pulpapel Forestry Station, 03°40'N, 77°00'W, 50 m, 24 Apr 1987, D. Faber-Langendoen, E.

Renteria, & M. Monsalve 247 (MO); Rio Naya, upriver from Puerto Merizalde, 03°15'N, 77°25'W, 10 m, 23 Feb 1983, A. Gentry &A.Juncosa

40693 (MO); Across from Pulpapel camp, ca. 15 km N of Buenaventura, 03°59'N, 77°05'W, 50 m, 9 Apr 1987, A. Gentry et al. 56698 (MO); Bajo

Calima, ca. 15 km N of Buenaventura, “new” Dindo area, 03°59'N, 77°02'W, 10 Apr 1987, A. Gentry et al. 56758 (MO); Bajo Calima, Conce-

sion Pulpapel / Buenaventura, 03°55'N, 77°00'W, 100 m, 20 Nov 1984, M. Monsalve B. 510 (CUVC, MO), 13 Dec 1984, M. Monsalve B. 619

(COL), 25 Jul 1989, M. Monsalve B. 3076 (MO); Bajo Calima, Concesion Pulpapel/Buenaventura, carretera gasolina Pt4, 03°55'N, 77°00'W,

100 m, 20 Oct 1987, M. Monsalve B. 2006 (MO); Municipio Buenaventura, forest exploitation in the concession of Carton de Colombia,

03°56'N, 77°10'W, 230 m, 15 Nov-6 Dec 1979,J. van Rooden, B.J.H. ter Welle, & S.M.C. Popper 386 (MO, U). ECUADOR. Prov. Esmeraldas:

San Miguel, Sector Rio Grande, Comunidad Corriente Grande, 00°45'N, 78°47'W, 350 m, 22 Nov 1992, G. Fipaz et al. 2320 (MO). Prov. Los

Rios: Rio Palenque Biological Station, km 56 Quevedo-Santo Domingo, 150-220 m, 11 Feb 1973, C.H. Dodson 5238 (L, US), 2 Oct 1976, C.H.

Dodson&A. Gentry 6455 (L, MO), 8 Oct 1976, A. Gentry & C.H. Dodson 18036 (MO), 17 Jan 1977, C.H. Dodson 6623 (L, MO).

In flower, Casearia draganae is easily recognized as a member of the ‘Singulars’ group due to the well-devel¬

oped hypanthium and the mostly included pistil with a globose or ovoid, rather than ellipsoidal or narrowly

ellipsoidal, ovary Using Sleumer’s key (1980: 304), the user will reach an impasse, as the sepal size falls closer

to the range of C. combaymensis (sensu Sleumer), but the disk lobes are hairy as in C.fasciculata. Other species,

some new and some previously synonymized (e.g., C. bracteifera Sagot, C. singularis Eichler, C. subopaca Triana

& Planch., C. petiolaris Poepp. ex Eichler) can be distinguished from C. draganae in having one of these fea¬

tures: glabrous young branches, leaves with pubescent veins abaxially, entire leaf margins, or long petioles (>1

cm). Species group ‘Singulars’ can also sometimes be recognized by the large pellucid punctations and lines,

which are evenly distributed and are included within a single areola of veins (Sleumer 1980). In the case of C.

draganae and some of the other species, though, this is not always true, as many smaller punctations may exist,

too (see Fig. 1C).

In fruit, Casearia draganae may be confused with other species of group ‘Singulars’ but also with other

large-fruited species with colorless arils in group ‘Decandrae.’ Identification in fruit can be difficult when fruits

are collected immature or maturity is not known. In species group ‘Singulars,’ other species differ from C.

draganae in having one of these features: thinner (<1 mm) or thicker (4-8 mm) fruit walls, longer petioles (8-

24 mm), or entire leaf margins. Among the members of species group ‘Decandrae,’ Casearia pitumba usually

has smaller fruits (2-3 cm diameter) with a hairy remnant of the style base, C. cajambrensis has prominently

ribbed fruits, and C. megacarpa occurs at higher elevations (1250-2200 m). Other large-fruited species either

have smaller mature fruits or longer petioles; most occur in the Amazon basin or in sub-Amazonian Brazil.

Preliminary phylogenetic work based on DNA data indicates that species group ‘Singulares’ may be nest¬

ed within the broader ‘Decandrae’ group (Samarakoon 2015, Samarakoon et al., pers. comm.) and that the

large-fruited species with colorless arils may form a monophyletic group. Additional collections of these spe¬

cies, especially where flowering and fruiting specimens can be obtained from the same individuals, will assist

Alford, A new species of Casearia (Samydaceae)

329

in clarifying the taxonomy of this group. Casearia maynacarpa Liesner & P. Jorgensen (Jorgensen & Liesner

2013), once placed in Mayna Aubl. or Carpotroche Endl. (now Achariaceae), may also belong to this group,

given its large fruit and thick fruit wall.

ACKNOWLEDGMENTS

I thank Rodrigo Bernal (COL) for encouraging me to participate in the project to produce the Catalogo de Plan-

tas y Liquenes de Colombia, which highlighted a need to study the large-fruited species of Casearia, and for

sharing information and many images of Colombian herbarium specimens. I thank the curators of A, BH,

BRIT, COL, DUKE, L, GB, GH, L, MICH, MO, U, US, and USMS for loaning specimens for study or accommo¬

dating my visits, Ron Liesner (MO) and Tharangamala Samarakoon (USMS) for sharing their critical observa¬

tions of the family, and Fabian Michelangeli (NY), John P. Janovec (BRIT, MOL), C. E. Timothy Paine (LSU,

STI), Mercedes S. Poster (US), Pablo Stevenson (ANDES), and Diana C. Acosta Rojas (ANDES) for providing

held images of various large-fruited species that helped to highlight important characteristics. I also thank

Tharangamala Samarakoon for sharing phylogenetic results on Samydaceae, Emily J. Lain and Tharangamala

Samarakoon for preparing the illustrations, Daniel M. McNair (USMS) for assisting in preparation of the

manuscript, and David E. Lemke and Nixon Leonardo Cumbicus Torres for helpful reviews.

REFERENCES

Alford, M.H. 2003. Claves para los generos de Flacourtiaceae de Peru y del Nuevo Mundo. Arnaldoa 10:19-38.

Alford, M.H. 2005. Systematic studies in Flacourtiaceae. Ph.D. dissertation, Cornell University, Ithaca, New York. 290 pp.

Alford, M.H. 2007. Samydaceae. The Tree of Life Web Project. Available at http://tolweb.org/Samydaceae/68361. Ac¬

cessed June 2015.

Alford, M.H. 2015 in press. Salicaceae. In: R. Bernal, S.R. Gradstein & M. Celis, eds. Catalogo de plantas y liquenes de Co¬

lombia. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogota. http://catalogoplantascolombia.

unal.edu.co

Angiosperm Phylogeny Group (APG). 2009. An update of the Angiosperm Phylogeny Group classification for the orders and

families of flowering plants: APG III. Bot. J. Linn. Soc. 161 (2):105-121. doi:10.1111 /j.1095-8339.2009.00996.X

Benson, M., ed. 1994. SerboCroatian dictionary. 3rd ed. Cambridge University Press, Cambridge, U.K. and New York, U.S.A.

Chase, M.W., S. Zmarzty, M.D. LledO, KJ. Wurdack, S.M. Swensen, & M.F. Fay. 2002. When in doubt, put it in Flacourtiaceae: a

molecular phylogenetic analysis based on plastid rbcL DNA sequences. Kew Bull. 57:141-181.

Faber-Langendoen, D. & A.H. Gentry. 1991. The structure and diversity of rain forests at Bajo Calima, Choco region, western

Colombia. Biotropica 23(1 ):2—11.

Jorgensen, P.M. & R.L. Liesner. 2013. Casearia maynacarpa (Salicaceae), a new name for Carpotroche parvifolia. Novon

22(4):422-423.

Mori, S.A. & B. Fischer. 2002. Flacourtiaceae (Flacourtia family). In: S.A. Mori, G. Cremers, C.A. Gracie, J.-J. de Granville, S.V.

Heald, M. Hoff, & J.D. Mitchell, eds. Guide to the vascular plants of central French Guiana. Part 2. Dicotyledons. New

York Botanical Garden Press, New York. Mem. New York Bot. Gard. 76(2):319-328, plates 60-61.

Olson, M., P.E. Berry, &G.A. AymardC. 1999. Flacourtiaceae. In: P.E. Berry, K. Yatskievych, & B.K. Holst, eds. Flora of the Ven¬

ezuelan Guayana. Missouri Botanical Garden Press, St. Louis. 5(Eriocaulaceae-Lentibulariaceae):434-474.

Phillips, O. & J.S. Miller. 2002. Global patterns of plant diversity: Alwyn H. Gentry's forest transect data set. Missouri Bo¬

tanical Garden Press, St. Louis.

Rodriguez-Mahecha, J.V., P. Salaman, P. Jorgensen, T. Consiglio, L. Suarez, F. Arjona, & R. Bensted-Smith. 2004. Tumbes-Choco-

Magdalena. In: R.A. Mittermeier, P. Robles Gil, M. Hoffmann, J. Pilgrim,T. Brooks, C. Goettsch Mittermeier, J. Lamoreux,

& G.A.B. da Fonseca, eds. Hotspots revisited: Earth's biologically richest and most endangered terrestrial ecoregions.

CEMEX, Mexico City, Mexico.

Samarakoon, T. 2015. Phylogenetic relationships of Samydaceae and taxonomic revision of the species of Casearia in

south-central Asia. Ph.D. dissertation, University of Southern Mississippi, Hattiesburg, Mississippi.

Sleumer, H.0.1980. Flacourtiaceae. Flora Neotropica 22:1-499.

Zmarzty, S. 2007. Flora da Reserva Ducke, Amazonas, Brasil: Flacourtiaceae. Rodriguesia 58(3):663-694.

330

Journal of the Botanical Research Institute of Texas 9(2)

BOOK REVIEW

Paul Dowlearn. 2013. The Lazy Man’s Garden: Maximum return; Minimum input. (ISBN-13: 978-149106-

889, pbk). Self-published, 5314 Southwest Parkway, Wichita Falls, Texas 76310, U.S.A. (Orders: www.

wvlandscape.com). $14.95,155 pp., 6" x 9".

Author Paul Dowlearn uses a tongue-in-cheek approach to present sound gardening lore in his newest

book The Lazy Man’s Garden: Maximum return; Minimum input. He shares the wisdom of his own life

experiences from observations and experimentation, using a foundation based on organic methods and

respecting what Mother Nature provides. He thinks outside the box of synthetic fertilizers and toxic

chemicals and follows Nature’s plan which he finds “perfectly suited to the Lazy Man.” Dowlearn stresses

the importance of planning, with an awareness of climate and soil conditions and using plants suitable for

the environment. He applies his best insights for successful gardening to achieving common sense solu¬

tions. Along with a re-awakening of our role as custodians of the planet, Dowlearn’s The Lazy Man’s Gar¬

den touts these as exciting times indeed.— Pat Gerard, Volunteer, Botanical Research Institute of Texas, U.S.A.

J.Bot. Res. Inst. Texas 9(2): 330.2015

IRENODENDRON, A NEW GENUS OF SAMYDACEAE FROM SOUTH AMERICA

Mac H. Alford and Angela D. Dement

Department of Biological Sciences

University of Southern Mississippi

Hattiesburg, Mississippi 39402, U.S.A.

mac.alford@usm.edu

ABSTRACT

Irenodendron (Samydaceae), a new genus for three species of shrubs and trees from South America, is described. Previously recognized as

members of Laetia section Scypholaetia, these species (I. coriaceum, I. cupulatum, and I. ovalifolium) differ from Laetia sensu stricto in

having cupular bracts, an apically divided style, and large rays in the wood. The wood anatomy and style morphology are similar to those of

Ryania and Piparea (=Casearia sect. Piparea), to which we propose the new genus is closely related.

RESUMEN

Se describe Irenodendron (Samydaceae), un nuevo genero para tres especies de arbustos y arboles de Suramerica. Anteriormente reconoci-

das como especies pertenecientes al genero Laetia seccion Scypholaetia, estas especies (I. coriaceum, I. cupulatum, e I. ovalifolium) se

diferencian de Laetia sensu stricto en tener bracteas cupulares, estilos divididos apicalmente, y radios largos en la madera. La anatomia de la

madera y la morfologia del estilo son similares a las de Ryania y Piparea (=Casearia sect. Piparea ), generos a los que se propone el estrecha-

mente relacionado Irenodendron.

Key Words: Flacourtiaceae, Irenodendron, Laetia sect. Scypholaetia, Piparea, Ryania, Samydaceae, South America

INTRODUCTION

Samydaceae (Malpighiales) are a pantropical family of about 256 species of trees and shrubs. They are closely

related to Salicaceae, Lacistemataceae, and Passifloraceae (Alford 2005, 2007; Wurdack & Davis 2009; Xi et al.

2012; Samarakoon 2015) and are sometimes included within a broad Salicaceae (Chase et al. 2002; APG 2009).

Generic circumscriptions in the family are currently being adjusted, as phylogenetic studies utilizing both

morphological and DNA sequence data affirm that the largest genus of the family, Casearia Jacq., is polyphy-

letic (Alford 2005; Samarakoon 2015). One result is that Casearia sect. Piparea (=Piparea Aubl.) (3 spp.) is more

closely related to Ryania Vahl and Trichostephanus Gilg than to other Casearia. This relationship is supported

by staminode position, style morphology, and wood anatomy (Alford 2005). Unlike most other Samydaceae,

the staminodes (or “disk glands” sensu Sleumer 1980) in Ryania, Trichostephanus, and Piparea are located inside

(vs. alternating with) the stamens, the style is apically divided into several branches, each branch with its own

stigma (versus not divided), and the wood consists of dark heartwood with large rays, 3-7 mm high, as wide as

15 cells or 240 pm, and visible to the naked eye (versus light-colored heartwood and small rays, mostly <2 mm

high and no wider than 7 cells) (Miller 1968,1975; Halle & de Wilde 1978).

In preparing a data table for an online interactive key to the species of the family, we noted that three spe¬

cies of Laetia L. have a striking morphological similarity to Ryania and Piparea, including the style and wood

characters noted above. Zmarzty (2007), too, noted a “strong, if superficial, resemblance” between one of these

species of Laetia and one species of Piparea, highlighting the similar leaf shape, leaf texture, capsule shape, and

capsule indumentum. Indeed, specimens of Piparea dentata Aubl. ( =Casearia commersoniana Cambess.) or Pi¬

parea multiflora C.F. Gaertn. (= Casearia javitensis H.B.K.) could easily be confused with these species vegeta-

tively, in flower, or in fruit; the species of Piparea, however, lack the cupular bracts and have a row of stami¬

nodes inside the stamens, the staminodes often ligulate and pubescent, contrasting with the thin and glabrous

filaments.

Although other species of Laetia have been sampled for DNA data, none of these three species has been

included (Alford 2005; Samarakoon 2015). Attempts to obtain fresh samples for DNA extraction or to extract

J. Bot. Res. Inst. Texas 9(2): 331 - 334.2015

332

Journal of the Botanical Research Institute of Texas 9(2)

DNA from herbarium specimens were unsuccessful. These species were placed in Laetia sect. Scypholaetia by

Warburg (1894), and repeated by Gilg (1925), due to the presence of a cup-like bract subtending the flowers,

but Sleumer (1980) in his revision of all Neotropical Flacourtiaceae did not recognize the sectional divisions.

Given the strong morphological evidence, however, we argue that these species should be recognized as a ge¬

nus distinct from Laetia and hypothesize that it is more closely related to Ryania and Piparea.

TAXONOMIC TREATMENT

Irenodendron M.H. Alford & Dement, gen. nov. Type: Irenodendron cupulatum (Spruce ex Benth.) M.H. Alford & De¬

ment (= Laetia cupulatum Spruce ex Benth.).

Laetia sect. Scypholaetia Warb., Die naturlichen Pflanzenfamilien III(6a):49. 1894. Type: Laetia coriacea Spruce ex Benth., J. Proc. Linn.

Soc., Bot 5 (Suppl. 2):84.1861, hie designatus.

Irenodendron is a new genus, similar to Piparea Aubl. and Ryania Vahl. in its divided style and large rays in the wood. Irenodendron differs

from those genera in having bracts fused into a cup-like structure subtending the pedicels and in lacking staminodes or “disk glands.” Ireno¬

dendron differs from Laetia L. sensu stricto in its cup-like bracts, divided style, and large rays in the wood.

Unarmed shrubs or trees, to 20 m tall and 30 cm dbh. Wood anatomy diffuse porous, pores often solitary,

perforation plates simple, intervascular and vessel-ray pitting alternate, both uniseriate homocellular and

multiseriate heterocellular rays with long uniseriate extensions present, multiseriate portion of rays >5 cm

high and up to 240 pm (15 cells) wide, hber-tracheids and libriform fibers present, septate, reddish brown de¬

posits abundant in ray cells and fibrous elements (Miller 1975, based on I. cupulatum, Williams 14194 (F),

MADw 23726). Leaves simple, alternate, distichous, pinnately veined, subcoriaceous to coriaceous, lacking

pellucid-punctations or lines (or practically so?), indumentum, if present, simple, margin entire to obscurely

serrate, crenate, or sinuate-crenate, petiolate, stipulate, stipules early caducous, lamina drying olive-brown

above, brown to reddish-brown below, with black petioles. Inflorescences axillary, rarely supra-axillary, fas¬

cicles of 3-10 flowers, from foliate or defoliate axils, pedicels surrounded at base by connate bracts forming a

cup, the cup sometimes deeply lobed or incised, the cup-like bracts appearing like a calyx and the calyx ap¬

pearing like a corolla in bud. Flowers perigynous, hypanthium <0.5 mm, appearing hypogynous, bisexual,

sepals (4-)5(-6), imbricate, connate at the very base, pale green to white to pink, petals absent, stamens 30-

74(+) in 2-3 close-set whorls, staminodes or “disk glands” absent, ovary superior, placentation parietal, placen¬

tae 3, style branched near apex into 3 branches, each with a minutely capitate stigma. Fruits capsules, dis¬

tinctly 3-sided, valves 3, rusty tomentulose outside, cupular bract persistent. Seeds 1-few, arillate.

Distribution. —Northern Amazonia, in Brazil, Colombia, Guyana, Peru, and Venezuela, and Pacibc coast

of Colombia.

Etymology. —The name is a compound formed from the Greek words 8tpf|vr| (=eirene), meaning peace,

and SevSpov (=dendron), meaning tree, with hopes of an enduring peace within and among the nations of the

Americas.

Comments. —Species delimitation in the genus needs further study. Collections that are available are of

good quality, but the small number of specimens does not permit a good assessment of variation.

KEY TO THE SPECIES OF IRENODENDRON

1. Leaves lanceolate to lanceolate-oblong to ovate, often sinuate-crenate, apex gradually attenuate to acuminate; pedicels

3-4 mm long, barely surpassing the cup of bracts_I. coriaceum

1. Leaves ovate, broadly ovate, elliptic, broadly elliptic, or oblong, often entire or obscurely toothed, apex acute or

abruptly acuminate for 1 (-2) cm; pedicels 7-10 mm long, distinctly surpassing the cup of bracts

2. Young branches and petioles pubescent; cup of bracts not conspicuously lobed or incised (although commonly torn

along one suture in dried specimens)_I. cupulatum

2. Young branches and petioles glabrous or glabrescent; cup of bracts deeply 2-3-lobed_I. ovalifolium

Irenodendron cupulatum (Spruce ex Benth.) M.H. Alford & Dement, comb. nov. Basionym: Laetia cupulata Spruce ex

Benth., J. Proc. Linn. Soc., Bot. 5 (Suppl. 2):84.1861; Guidonia cupulata (Spruce ex Benth.) O. Kuntze, Rev. Gen. Pi. 1:44.1891. Type:

VENEZUELA. Terr. Fed. Amazonas: ad. flum. Casiquiare supra Vasivae ostium, Dec 1853, Spruce 3200 (holotype: K (http://apps.

kew.org/herbcat/getImage.do?imageBarcode=K000471393), isotypes: L [fragment ex W], P, S, W).

Alford and Dement, Irenodendron, a new genus of Samydaceae

333

Distribution .—Amazonian Brazil, Colombia, Guyana, and Venezuela, possibly Peru; mixed evergreen forest,

terra firme, often along streams.

Representative specimens examined: BRAZIL. Amazonas: Alto Rio Solimoes, municipio de Sao Paulo de Olivenga plato ao sul da cidade,

estrada para a localidade do Bom Fim, 25 Nov 1986, C.A. Cid et al. 8546 (INPA, MG).

Irenodendron coriaceum (Spruce ex Benth.) M.H. Alford & Dement, comb. nov. Basionym: Laetia coriacea Spruce ex

Benth., J. Proc. Linn. Soc., Bot. 5 (Suppl. 2):84.1861; Guidonia coriacea (Spruce ex Benth.) O. Kuntze, Rev. Gen. Pi. 1:44.1891. Type:

COLOMBIA. Guainia: Tomo, Aug 1854, Spruce 2730 (lectotype, as designated by Sleumer [1980: 251]: K (http://apps.kew.org/herb-

cat/getImage.do?imageBarcode=K000471398), isolectotypes: B, BR, GOET, P).

Distribution .—Amazonian Brazil, Colombia, and Venezuela; savannah or caatinga on white sand.

Representative specimens examined: BRAZIL. Amazonas: Rio Negro, Communidade Aparecida, opposite mouth of Igarape Tuan, near

abandoned portion of Perimentral Norte, 00°20'N, 67°18'W, 6 Nov 1987, P.J.M. Maas et al. 6932 (F, MG); Savanna, 3 km S of Central Massif

of Serra Araca, 00°49'N, 65°17'W, 100 m, 18 Jul 1985, G.T. Prance et al. 29677 (MICH); 3 km ao Sul da parte central da Serra Araca E 8 km a

Leste do riojauari, 00°49'N, 63°19'W, 60 m, 14 Mar 1984, W.A. Rodrigues et al. 10523 (MICH). Rio Atabapo, Yavita-Pimichin trail near Yavita,

125-140 m, lOJun 1959J.J. Wurdack&L.S. Adderley 42900 (MICH).

Irenodendron ovalifolium (Macbride) M.H. Alford & Dement, comb. nov. Basionym: Laetia ovalifolia Macbride, Can-

dollea 5:389.1934. Type: PERU. Loreto: Mishuyacu near Iquitos, Jan 1930, Klug 757 (holotype: F (http://emuweb.fieldmuseum.org/

web/pages/common/ imagedisplay.php?irn=56562&reftable=efmnh&refirn=257000), isotypes: NY, US).

Distribution .—Amazonian Brazil and Peru, and Pacific of Colombia, collection from Colombia possibly a new

species; lowland forest on white sand.

Representative specimens examined: BRAZIL. Rondonia [Mato Grosso of 1931]: near Tabajara, upper Machado [Ji-Parana] River region,

Nov-Dee 1931, B.A. Krukoff 1393 (MICH). PERU. Loreto: Maynas, Rio Nanay, Caseria Mishana, 30 km SW of Iquitos, Callicebus Biological

Reserve, 4 km S of Mishana, 13 Aug 1980, R.B. Foster 4428 (F); Maynas, Yanamono, Explorama Tourist Camp, halfway between Indiana and

mouth of Rio Napo, 03°28'S, 72°50'W, 130 m, 27Jun 1983, A. Gentry et al. 42229 (MO).

ACKNOWLEDGMENTS

This work was a result of a summer internship by the junior author at the University of Southern Mississippi.

She was supported by NSF grant HRD 0450362, the Alliance for Graduate Education in Mississippi. Funding

for the senior author was through start-up funds and from the American Philosophical Society. Herbaria BH,

COL, F, GH, INPA, L, K, MICH, MG, MO, NY, US, and USMS are thanked for loaning specimens for study or

accommodating our visits, and Ron Liesner (MO) is thanked for sharing his critical observations of the family.

Tharangamala Samarakoon (USMS) is thanked for sharing phylogenetic results on Samydaceae, Cesar Gran-

dez Rios (AMAZ) is thanked for assisting in the held, N. Raul Anzola is thanked for improving the abstract in

Spanish, and Robert Kiger and Wendy Applequist are thanked for helpful reviews.

REFERENCES

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Alford, M.H. 2005. Systematic studies in Flacourtiaceae. Ph.D. dissertation, Cornell University, Ithaca, New York. 290 pp.

Alford, M.H. 2007. Samydaceae. The tree of life web project. Available at http://tolweb.org/Samydaceae/68361.

Accessed June 2015.

Angiosperm Phylogeny Group (APG). 2009. An update of the Angiosperm Phylogeny Group classification for the orders and

families of flowering plants: APG III. Bot. J. Linn. Soc. 161 (2):105-121. doi:10.1111/j.1095-8339.2009.00996.x

Chase, M.W., S. Zmarzty, M.D. LledO, KJ. Wurdack, S.M. Swensen, & M.F. Fay. 2002. When in doubt, put it in Flacourtiaceae: a

molecular phylogenetic analysis based on plastid rbcL DNA sequences. Kew Bull. 57:141-181.

Gilg, E. 1925. Flacourtiaceae. In: A. Engler & K. Prantl, eds. Die naturlichen Pflanzenfamilien, 2nd ed. Duncker & Humblot,

Berlin. 21:377-457.

Halle, N. & JJ.F.E. de Wilde. 1978. Trichostephanus acuminatus Gilg (Flacourtiacees), une approche biosystematique.

Adansonia, ser. 2,18(2):167-182.

Miller, R.B. 1968. Systematic wood anatomy of the American Casearia Jacq. (Flacourtiaceae). M.S. thesis, University of

Wisconsin, Madison, Wisconsin. 56 pp.

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Miller, R.B. 1975. Systematic anatomy of the xylem and comments on the relationships of Flacourtiaceae. J. Arnold

Arbor. 56(1 ):20-102.

Samarakoon, T. 2015. Phylogenetic relationships of Samydaceae and taxonomic revision of the species of Casearia in

south-central Asia. Ph.D. dissertation, University of Southern Mississippi, Hattiesburg, Mississippi. 160 pp.

Sleumer, H.0.1980. Flacourtiaceae. Flora Neotropica 22:1-499.

Warburg, 0.1894. Flacourtiaceae. In: A. Engler & K. Prantl, eds. Die naturlichen Pflanzenfamilien, 1st ed. Wilhelm Engle-

mann, Leipzig. 3(6a):1-56.

Wurdack, K.J. & C.C. Davis. 2009. Malpighiales phylogenetics: Gaining ground on one of the most recalcitrant clades in

the angiosperm tree of life. Amer. J. Bot. 96(8):1551 -1570.

Xi, Z., B.R. Ruhfel, H. Schaefer, A.M. Amorim, M. Sugumaran, KJ. Wurdack, P.K. Endress, M.L. Matthews, P.F. Stevens, S. Mathews,

& C.C. Davis. 2012. Phylogenomics and a posteriori data partitioning resolve the Cretaceous angiosperm radiation

Malpighiales. Proc. Natl. Acad. Sci. U.S.A. 109(43):17519-17524.

Zmarzty, S. 2007. Flora da Reserva Ducke, Amazonas, Brasil: Flacourtiaceae. Rodriguesia 58(3):663-694.

NEW SPECIES OF SENEGALIA (FABACEAE) FROM SOUTH AMERICA

David S. Seigler*

John E. Ebinger

Department of Plant Biology

University of Illinois

Urbana, Illinois 61801, US.A.

seigler@life. illinois. edu

*corresponding author

Emeritus Professor of Botany

Eastern Illinois University

Charleston, Illinois 61920, US.A.

jeebinger@eiu.edu

ABSTRACT

Senegalia duartei Seigler & Ebinger from the state of Rio de Janeiro, Brazil; S. noblickii Seigler & Ebinger, from the states of Bahia, Espirito

Santo and Pernambuco, Brazil; and S. phillippei Seigler & Ebinger from northern South America in Brazil (Amapa and Para), French Gui¬

ana, Guyana, Surinam, and Venezuela (Anzoategui and Bolivar) are described, illustrated, and compared to their probable nearest

relatives.

Keywords: Fabaceae, IUCN Red List, Mimosoideae, Senegalia

RESUMEN

Senegalia duartei Seigler & Ebinger del estado de Rio de Janeiro, Brasil; S. noblickii Seigler & Ebinger, de los estados de Bahia, Espirito

Santo y Pernambuco, Brasill, and S. phillippei Seigler & Ebinger localizada desde el norte de America del Sur desde los estados de Amapa y

Para, Brasil, Guayana Francesa, Guyana, Surinam, y Venezuela desde los estados Anzoategui y Bolivar, son descritas, ilustradas y compara-

das con sus especies afines.

The genus Senegalia currently comprises approximately 102 taxa in the Neotropics (Barros & Morim 2014),

about 68 in Africa, 45 in Asia, and two in Australia (Maslin et al. 2003a, b; Maslin 2015); of which 8 species

occur in two or more areas. Both Old and New World members of the genus are shrubs, trees, or lianas, with

extrafloral petiolar nectaries, and branches and branchlets usually armed with prickles and lacking stipular

spines. The prickles are randomly scattered along the twigs, and also commonly the leaf petiole and rachis

though some species have the prickles in lines along the twigs, and prickles are sometimes fused together into

lines. In a few species prickles are grouped into twos or threes, usually at or near the nodes. Leaves are bipin-

nately compound with one to nearly 50 pinna pairs that contain one to many pairs of leaflets. The petiole and

rachis have variously shaped glands, the position, structure, and shape being consistent within a species and

are of diagnostic importance. Flowers are small, mostly 5-merous, synsepalous, sympetalous, with numerous

stamens (usually 50 to 180), the filaments are not fused and are attached to a more-or-less tubular or circular

nectary disc located at the receptacle of the flowers just below the usually stipitate ovary. Inflorescences are

globose heads or spikes occurring solitary in small clusters in the leaf axils, or grouped into complex axillary

or terminal pseudo-racemes or pseudo-panicles. Pods are usually oblong or broadly linear, with 6-15 uniseri-

ate seed, separating into two valves at maturity or rarely indehiscent or separating into indehiscent one-seeded

loments.

Until recently the genus Senegalia was treated as part of Acacia s.l. (Lewis et al. 2005), but recent morpho¬

logical, chemical and molecular evidence has determined that this large genus represents a polyphyletic unit.

In addition, those studies have helped determine relationships within the genus Acacia s.l., as well as the posi¬

tion of the genera within the Mimosoideae (Maslin et al. 2003a; Miller & Bayer 2003; Luckow et al. 2003; Miller

et al. 2003; Rico-Arce & Bachman 2006; Seiger et al. 2006a; Bouchenak-Khelladi et al. 2010; Gomez-Acevedo

et al. 2010; Murphy et al. 2010; Miller & Seigler 2012; Kyalangalilwa et al. 2013). Based on these data, Acacia s.l.

currently is recognizable as five monophyletic lineages: Acacia s.s., Acaciclla Britton & Rose, Mariosousa Seigler

& Ebinger, Senegalia Raf., and Vachcllia Wight & Arnott (see Miller & Seigler 2012 for an overview of the new

generic classification).

J. Bot. Res. Inst. Texas 9(2): 335 - 343.2015

336

Journal of the Botanical Research Institute of Texas 9(2)

During the course of our work on the genus Senegalia of the New World three undescribed species were

noted from herbarium materials. These taxa are clearly distinct and are here proposed as new species.

Senegalia duartei Seigler & Ebinger, sp. nov. (Fig. 1). Type: BRAZIL. Rio deJaneiro: Arraialdo Cabo, Cabo Frio, Oct 1961, A.P.

Duarte, Ddrdano & Edmundo 5764 isotype: MO).

Senegalia duartei differs from the morphologically most similar species S. bahiensis (Benth.) Seigler & Ebinger in that the leaves are 90-210

mm long (vs. 40-85), the rachis is 70-170 mm long (vs. 20-70), there are 12-26 leaflets/pinna (vs. 6-14), the globose inflorescence is 14-19

mm in diameter (vs. 20-25), the corolla is 2.6-3.6 mm (vs. 3.9-5.2) and the stamens are 6.6-9 mm (vs. 9.5-11.5).

Liana to 4 m tall; bark not seen; twigs light to dark purplish brown, slightly flexuous, terete to ridged, glabrous

to lightly puberulent; short shoots absent; prickles, light to dark purplish brown, sometimes darker above, flat¬

tened, straight to recurved, woody, 1-4 mm long x 1-4 mm wide at the base, glabrous, persistent, widely scat¬

tered along the twig, petiole and rachis. Leaves alternate, 90-210 mm long; stipules brown, ovate and auricu-

late, symmetrical, flattened, straight, herbaceous, 10-25 x 10-20 mm at the widest point, glabrous, early de¬

ciduous; petiole adaxially grooved, 11-35 mm long, glabrous; petiolar gland usually present, located medially

to just below the lowest pinna pair, columnar, 0.7-2.5 mm long, apex 0.4-0.9 mm across, oval, glabrous; rachis

adaxially grooved, 70-170 mm long, glabrous, a sessile globose to columnar gland 0.2-0.5 mm long, some¬

times present between the terminal pinna pair, apex, flattened, glabrous; pinnae 3 to 11 pairs/leaf, 40-115 mm

long, 15-25 mm between pinna pairs; paraphyllidia 0.5-1.1 mm long, early deciduous; petiolule 4.0-6.7 mm

long; leaflets 12 to 26 pairs/pinna, opposite, 2.7-4.0 mm between leaflets pairs, oblong, 8-15 x 2.5-4.0 mm,

glabrous, lateral veins obvious, 3 to 6 veins from the base, base oblique, truncate on one side, margins not cili-

ate, apex acute, midvein subcentral to submarginal. Inflorescence a densely 25- to 60-flowered globose head

14-20 mm across, in axillary to terminal pseudo-racemose or pseudo-paniculate clusters, the main axis to 300

mm long; peduncles 10-27 mm long x 0.4-0.7 mm thick, pubescent; receptacle usually slightly enlarged, el¬

lipsoidal; involucre 1 (rarely 2) small bracts scattered along the peduncle, early deciduous; floral bracts spatu-

late, 0.5-1.4 mm long, puberulent, early deciduous. Flowers pedicellate, stalk 0.5-1.1 mm long, white; calyx

5-lobed, 1.8-2.6 mm long, lightly pubescent; corolla 5-lobed, 2.6-3.6 mm long, glabrous to lightly puberulent,

lobes one-sixth the length of the corolla; stamens 50 to 70, blaments 6-9 mm long, distinct; anther glands ab¬

sent; ovary pubescent, stipe to 1.1 mm long. Legumes straight, flattened, not constricted between the seeds,

oblong, 100-170 x 18-23 mm, chartaceous, transversely striated, glabrous, eglandular, dehiscent along both

sutures; stipe 16-19 mm long; apex acute, beak 0.6-1.1 mm long. Seeds not seen.

Notes.—Senegalia duartei has large ovate stipules (10-25 mm long, 10-20 mm wide) similar to bve other

Senegalia species from Brazil, S. bahiensis (Benth.) Seigler & Ebinger (6-15, 4-10), S. globosa (A. Bocage &

Miotto) de Queiroz (4-7, 0.8-1.3), S. grandistipula (Benth.) Seigler & Ebinger (15-35, 8-25), S. lasiophylla

(Benth.) Seigler & Ebinger (5-10, 3-6) and S. tamarindifolia (L.) Britton & Rose (6-25, 5-20). Although the

sizes and shapes of the stipules overlap to some extent, S. duartei can be distinguished from each of the other

broad-stipule species by other features. S. duartei differs from S. globosa because the latter has globose petiole

and rachis nectaries whereas the glands of S. duartei are columnar. In addition, the leaves of S. duartei are lon¬

ger than those of S. globosa (40-100 mm) and the globose inflorescence is smaller (14-20 vs 20-29 mm). S.

duartei differs from S. grandistipula because the latter has a spicate inflorescence (20-35 x 20-26) mm. In addi¬

tion, the corolla and stamens are longer (6-9 and 9.5-11.5 mm, respectively). S. duartei can be separated from

S. lasiophylla by the smaller stipules (5-10 x 3-6), shorter distance between pinnae (5-11 mm), larger globose

inflorescence diameter, and calyx, corolla, and stamen lengths (2.8-4.1, 3.9-5.2, and 9.5-11.5, respectively).

The anthers of S. lasiophylla possess anther glands which do not occur in S. duartei. Senegalia duartei can be

distinguished from S. tamarindifolia by the number of leaflets/per pinna (9-19 x 12-26), a smaller globose in¬

florescence (20-25 mm), and correspondingly longer calyx, corolla and stamen lengths.

Distribution and ecology —Dry seasonal forests and caatinga, below 500 m in the state of Rio de Janeiro,

Brazil.

Phenology. —Flowering Sep-Dee.

Seigler and Ebinger, New species of Senegalia from South America

337

50 mm

Fig. 1. Senegalia duartei. A. Legume; B. Lower side of leaflet; C. Flower (inverted); D. Stipule; E. Lower leaf surface and stem; F. Petiolar gland; G.

Inflorescences (portion of pseudopanicle). (A, B, C, G L. Riedel 150, NY; D A.P. Duartei & G.M. Barroso 5040, FY; E, F L. Riedel 9, NY).

338

Journal of the Botanical Research Institute of Texas 9(2)

Local names and uses. —None known.

Etymology. —This species is named after A.P. Duarte of the Jardin Botanico de Rio de Janeiro who col¬

lected extensively in the area of Rio de Janeiro.

IUCN Red List category. —DD, data deficient. A rare endemic restricted to the state of Rio de Janeiro, Brazil.

We have seen only 12 specimens including the types and have not found any recent collections of this taxon,

the last being collected in 1961 (IUCN 2001).

Paratypes: BRAZIL: Rio de Janeiro: Rio de Janeiro, 1851, N.Y. Andersson s.n. (S); Guanabara, Barra da Tijuca, 30 Sep 1959, A.P. Duarte 5040

(F); Guanabara, Sacopa, 6 Sep 1960, A.P. Duarte 5277 (F, MO); Morro da Babilonia, Oct 1914, F.C. Hoehne s.n. (HUEFS); Rio de Janeiro, Sep

1829, L. Riedel 9 (NY); Rio de Janeiro, Dec 1831, L. Riedel & B. Luschnatt 150 (A, NY, US). No state listed: Glaziou 8438 (F).

Senegalia noblickii Seigler & Ebinger, sp. nov. (Fig. 2). Type: BRAZIL. Bahia: Iagu, Faz. Smbra, 12°43 , S/40°7 , W, 14 Mar 1985,

L.R. Noblick 3699 (holotype: HUEFS; isotype: NY).

Senegalia noblickii differs from it’s probable nearest relative S. tenuifolia (L.) Britton & Rose which has an orbicular petiolar gland (1.5-4.2

mm long), 10-25 pinna pairs/leaf, leaflets 2.5-5.2 mm long, and 0.5-1.0 mm wide, globose inflorescences 5-10 mm in diameter, and a

transversely striated legume.

Liana to 10 m long; bark not seen; twigs light brown to light purplish brown, not flexuous, terete to slightly

ridged, puberulent; short shoots absent; prickles light purplish brown, the apex usually darker, flattened, re¬

curved, woody, 1-3 x 1-3 mm at the base, glabrous to puberulent, persistent, randomly scattered along the

twig, petiole and rachis. Leaves alternate, 40-150 mm long; stipules green, triangular, symmetrical, flattened,

straight, herbaceous, 1-4 long, 0.7-1.5 mm wide at the base, glabrous, deciduous to persistent; petiole shal¬

lowly adaxially grooved, 15-27 mm long, glabrous to lightly puberulent; petiolar gland solitary, located on the

lower third of the petiole, sessile, mostly oval, 0.4-1.1 mm long, apex flat, sometimes becoming cup-shaped

when mature, glabrous; rachis adaxially grooved, 30-130 mm long, lightly puberulent, an orbicular gland

0.3-0.6 mm across usually between the upper pinna pair, apex depressed and glabrous; pinnae 3 to 13 pairs/

leaf, 24-85 mm long, 6-14 mm between pinna pairs; paraphyllidia 0.4-0.9 mm long, commonly absent; peti-

olule 1.2-2.2 mm long; leaflets 18 to 44 pairs/pinna, opposite, 1.2-2.3 mm between leaflet pairs, oblong, 5.6-

11.0 x 1.3-2.5 mm, glabrous or nearly so on both surfaces, lateral veins obvious, 1 to 4 veins from the base, base

oblique, truncate on one side, margins lightly ciliate, apex broadly acute to obtuse, midvein nearly marginal at

the base, becoming subcentral at the leaflet apex. Inflorescence a densely 15- to 35-flowered globose head

9-14 mm across, in terminal pseudo-paniculate clusters, the main axis to 300 mm long; peduncles 2-8 mm

long x 0.4-0.6 mm thick, puberulent; receptacle not enlarged; involucre absent; floral bracts spatulate, 0.3-0.6

mm long, puberulent, early deciduous. Flowers sessile, white; calyx 5-lobed, 0.8-1.7 mm long, puberulent;

corolla 5-lobed, 1.6-2.5 mm long, glabrous to lightly puberulent, lobes one-third the length of the corolla; sta¬

men filaments 4-6 mm long, distinct; anther glands present; ovary pubescent, stipe to 1.6 mm long. Legumes

straight, flattened, not constricted between the seeds, oblong, 55-120 x 14-23 mm, chartaceous, transversely

striated, puberulent, numerous minute purple glands present, dehiscent along both sutures; stipe 2-5 mm

long; apex acute to obtuse, short beaked. Seeds uniseriate, no pulp, dark brown, orbicular, strongly flattened,

6.5-8.5 x 4.2-5.8 mm across, smooth; pleurogram U-shaped, 1.8-2.4 mm across.

Notes.—Senegalia noblickii shares many features with S. tenuifolia. The relatively small globose inflores¬

cence with small and relatively few flowers suggest a relationship, as does the liana habit, the scattered re¬

curved prickles, the orbicular petiolar gland with a flat apex, and the marginal midvein on the lower part of the

leaflet that becomes submarginal above. Of the species similar morphologically to S. tenuifolia, S. noblickii is

most similar morphologically to S. maschalocephala (Griseb.) Britton & Rose (endemic to Cuba) and S. ebingeri

Seigler (endemic to southeastern Peru) by consistently having < 44-45 leaflets/pinna, but can be distinguished

from the last two species by the larger inflorescence size (S. maschalocephala 12-18 mm, S. ebingeri 13-20 mm)

and the fact that those two species lack anther glands. Although somewhat similar in morphology to S. cathari-

nensis and S. riparia, they differ in that they lack anther glands, the leaflets have central venation, and both

have two glands on the petiole.

340

Journal of the Botanical Research Institute of Texas 9(2)

Distribution and ecology. —Wet tropical forests in alluvial soil, disturbed second growth forest, and caat-

inga from sea level to 500 m in the states of Bahia, Pernambuco and Espirito Santo, Brazil.

Phenology. —Flowering Mar-Jul.

Common name.—None known.

Etymology. —This species is named after L.R. Noblick of the Montgomery Botanical Center, Miami, Flori¬

da, who collected the type specimen. Mr. Noblick is a notable botanist. He has collected extensively in north¬

eastern Brazil and is a recognized authority on palms.

IUCN Red List category. —DD, data deficient. Relatively rare, recently described from about 10 specimens

from eastern Brazil (IUCN 2001).

Paratypes: BRAZIL: Bahia: Povoado Bizamum, 23 k de Tucano, 10°53 , 37"S/38°58 , 2rW, 5 Jun 2004, D. Cardoso 80 (HUEFS); Feira de San¬

tana, Ipuagu, 12 0 13 , 54 ,, S/39 0 4 , 35"W, 200-300 m, 14 Oct 2004, S.F. ConceiQdo, M.F. Borba-Silva, P.R. Almeida, F. Franca & R. Castro 2 (HUEFS,

NY); Monte Santo, 10°26 , 19 ,, S/39°20 , 9 ,, W, 480 m, 27 Feb 2000, AM. Giulietti & R.M. Harley 1839 (HUEFS); Riachao deJacmpe-Rio Toco, 14

km SE da cidade na Br. 324, 12°15 , S/38°58 , W, 10 Jul 1985, L.R. Noblick 4121 (HUEFS, NY); Main', Margem direita do Rio Jacuipe, 29 Mar

1983, E.L.P.G. de Oliveira 656 (HUEFS, MO, NY); 1 km E of Sastro Alves, 12°46 , 21 ,, S/39 0 19 , 43"W, 12 Mar 1993, L.P de Queiroz, M.J.S.L. Castro

& F.S.N. Sena 3079 (NY); Itiuba, Serre de Itiuba, 10°40 , 55 ,, S/39°48 , 46 ,, W, 445 m, 16 Aug 2002, L.P. de Queiroz, C. Correia,J. Costa&J.G. Nas-

cimento 7304 (HUEFS); Maracas, 10 km na estrada para Contendas do Sincora, 26 Feb 2000, M.M. da Silva, R.P. Oliveira, I. Castro, & M.A.S.

Santos 297 (HUEFS). Espirito Santo: Eneruzilhada, margem do Rio Pardo, Matacipo, 26 May 1968, R.P. Belem 3659 (CM). Pernambuco:

Floresta, Inaja, Reserva Biologica de Serra Negra, 8 Mar 1995, G.M. Souza, M. Oliveira, A.PS. Gomes &A. Laurenio 61 (MO, NY, US). State not

known: Brasilia, Blanchet s.n. (BM).

Senegalia phillippei Seigler & Ebinger, sp. nov. (Fig. 3). Type: FRENCH GUIANA: liana in disturbed open wooded area, Saul,

just E of junction La Fontaine and route de Belizon, 2 km S Les Eaux-Claires, 4.70166°N/52.4223° W, 280 m, 26 Sep 1995, L.R. Phil-

lippe,J.L. Crane, S. Mori, C. Grade, B. Wever&R. Yahr 27006A (holotype: ILLS; isotype: NY).

Senegalia phillippei differs from S. hoehnei Seigler, M.P. Morim, M.J.F. Barros, & Ebinger which has the petiole, rachis, and young twigs pu¬

bescent with curled yellow hairs to 0.5 mm long, the globose inflorescence 12-17 mm in diameter, and the ovary glabrous with a stipe up to

1.1 mm long. Senegalia phillippei differs from another similar species S. martiusiana which has a petiole pubescent with reddish-brown hairs,

a densely pubescent calyx, a glabrous ovary, pedicellate flowers (0.9-1.5 mm long) and leaflets 3.5-6.1 in length.

Liana to 35 m tall; bark not seen; twigs dark purplish brown, not flexuous, terete to slightly ridged, usually

densely puberulent; short shoots absent; prickles light brown throughout to dark brown toward the apex, flat¬

tened, mostly recurved, woody, 1-4 mm long, 1-3 mm wide at the base, usually pubescent near base and

sometimes throughout, persistent, scattered along the twig ridges, petiole, and rachis. Leaves alternate, 90-

180 mm long; stipules dark brown to dark reddish brown, narrow triangular, symmetrical, flattened, straight,

herbaceous, 1-4 x 0.5-0.9 mm near the base, puberulent, tardily deciduous; petiole adaxially grooved, 8-16

mm long, puberulent; petiolar glands usually 2, one just below the lowermost pinna pair, columnar, 0.4-1.5

mm long, gland apex commonly expanded, 0.4-1.5 mm across, orbicular to oval, depressed, glabrous; rachis

adaxially grooved, 80-165 mm long, puberulent with a sessile to commonly columnar gland 0.3-1.1 mm long,

between most of the pinna pairs, apex commonly expanded, 0.3-0.8 mm across, orbicular, flat to depressed,

glabrous, pinnae 16 to 33 pairs/leaf, 25-48 mm long, 2-6 mm between pinna pairs; paraphyllidia 0.3-1.1 mm

long, commonly absent; petiolule 0.8-1.4 mm long; leaflets 40 to 70 pairs/pinna, opposite, 0.3-0.7 mm be¬

tween leaflet pairs, linear, 2.5-4.1 x 0.4-0.8 mm, glabrous, lateral veins not obvious, 1 vein from the base, base

oblique, obtuse, margins minutely ciliate, apex obtuse, midvein essentially central. Inflorescence a densely

16- to 38-flowered globose head 9-13 mm across, in terminal pseudo-paniculate clusters, the main axis to 300

mm long; peduncles 4-12 x 0.3-0.5 mm thick, puberulent; receptacle not enlarged, globose; involucre rarely a

single small bract on the peduncle, early deciduous; floral bracts spatulate, 0.3-0.6 mm long, puberulent, early

deciduous. Flowers sessile, cream; calyx 5-lobed, 1.1-2.1 mm long, lightly puberulent; corolla 5-lobed, 2.2-3.2

mm long, usually glabrous, lobes one-quarter the length of the corolla; stamen filaments 5-7 mm long, dis¬

tinct; anther glands absent; ovary pubescent, stipe to 1.6 mm long. Legumes straight, flattened, not constricted

between the seeds, oblong, 110-190 x 25-38 mm, coriaceous, transversely striated, glabrous, eglandular, de¬

hiscent along both sutures; stipe 8-18 mm long; apex acute, beaked. Seeds not seen.

Notes.—Senegalia phillippei is morphologically similar to S. hoehnei , a species of eastern Brazil. With this

Seigler and Ebinger, New species of Senegalia from South America

341

Fig. 3. Senegalia phillippei. A. Legume; B. Inflorescences (portion of pseudopanicle) C. Leaf and stem; D. Petiolar glands; E. Lower side of leaflet; F. Flower

(inverted). (A-F 5. Mori, G. Prance, 1 Boeke, B. Boom 15016, WIS).

342

Journal of the Botanical Research Institute of Texas 9(2)

species it shares 2 columnar petiolar glands, a large number of relatively small leaflets on the pinnae, these

with central to subcentral midveins, and small flowers in small globose heads clustered into large pseudo¬

panicles. Senegaliaphillippei is separated from S. hoehnei by having the petiole, rachis, and young twigs puberu-

lent with curled hairs to 0.5 mm long in S. hoehnei), and rachis glands between the ultimate 10 pinna pairs and

occasionally some lower pinna pairs in S. hoehnei).

Distribution and ecology. —Moist tropical and second growth forests and thickets from near sea level to

1800 m in Brazil (Amapa and Para states), French Guyana, Guyana, Surinam, Venezuela (Anzoategui and

Bolivar states).

Phenology. —Flowering Sep-Nov.

Local names and uses. —None known.

Etymology. —This species is named after L.R. Phillippe of the Illinois Natural History Survey, Champaign,

who was a co-collector of the type specimen.

IUNC Red List category. —DD, data deficient. Probably relatively common but rarely collected (IUCN

2001 ).

Paratypes: BRAZIL: Amapa: 3 km E of Mata Indios, Rio Oiapoque, 2°47 , N/52°27 , W, 24 Sep 1960, H.S. Irwin, J. M. Pires&L.Y.T. Westra 48464

(B, F, NY, US); Rio Araguari, Sierra do Navio, 25 Sep 1961 ,J.M. Pires, W. Rodrigues & G.C. Irvine 51192 (NY, US); Rio Falsino, 10 km upstream

from confluence with Rio Araguari, UO'N/SFM-O'W, 1 Oct 1987, J. Pruski, D. Campbell, D. Williams, R.F. Garcia, J. Freitas, J. Cardoso &. E.

Batista 3335 (MO, NY, US). Para: 10 km N from the state line along BR-010,230 m, 16 Jul 1987, S. Fsugaru & Y. Sano 264 (MO, NY). FRENCH

GUIANA: 40 km from the bridge on the Mahury, S of Cayenne, 4.54802N/52.14872W, 295 m, 21 Sep 2008, C. Feuillet, A. Rossman & S.

Jackman 17015 (NY); Camp No. 3,54°rN/2°36'W, 160 m, 26 Aug 1987J.J. de Granville, L.Allorge, WJ. Hahn, M. Hoff, &N. Weizman 9736 (NY);

Saul, Monts Fa Fumee, 3°37'N/53°12 , W, 200-400 m, 17 Oct 1982, S. Mori, B. Boom, G. Prance&J. Boeke 15106 (CM, MO, NY, WIS); Saul,

3°37 , N/53°12 , W, 200-500 m, 12 Feb 1998, S.A. Mori, C.A. Grade &K.L. Purzycki 24764 (NY); Between airport and Saul, 3°37 , N/53°12 , W, 200

m, 29 Oct 1986, L. Skog, C. Feuillet &A. Rossman 7121 (NY). GUYANA: Km 60 de la route N2 en direction de Regina, 2 Nov 1982, F. Billiet &

B.Jadin 1532 (BR); Potaro-Siparuni region, Iwokrama Rainforest Reserve, 4°45 , N/59°1 , W, 75-400 m, 8 Nov 1995, D. Clarke & B. Hoffman 432

(NY); Camp 4, 220 m, 31 Aug 1987, J.J. Granville, L. Allorge, WJ. Hahn, M. Hoff & N. Weitzjman 9866 (P, US); Marudi Moutains, Mazoa Hill,

2°15 , N/59°10 , W, 400-450 m, 9 Nov 1982, A.L. Stoffers et al. 217 (MO). SURINAM: Marowijne District, Nassau Mountains, 4°49 , N/54°37 , W,

500-550 m, 7 Feb 2003, M J.Jansen-Jacobs, U.P.D.Raghoenandan, A Grant, A. Ramharrakh, H. terSteege & O.Bdnki 6612 (NY); Banks of Tanji-

mama River, 14 Nov. 1944, A.M.W. Mennega 381 (NY). VENEZUELA: Anzoategui: S-facing slopes of Cerro Peonia (Cerro Los Pajaritos),

above Santa Cruz, 1600-1800 m, 20 Mar 1945, J.A. Steyermark 61574 (F, MO, VEN-n.v.); Bolivar: Reserva Forestal Imataca, Carretera Casa

Blanca-San Martin de Turumban, 21 May 1982, B. Stergios, G. Aymard & O. Palacios 3837 (MO, PORT-n.v.).

ACKNOWLEDGMENTS

The authors wish to thank the curators of B, BR, CM, F, HUEFS, MO, NY, P, S, US and WIS for loan of speci¬

mens critical for this study. The authors thank several colleagues for advice concerning questions of nomencla¬

ture and general taxonomic advice. Among these are: J. Lee Crane, K.N. Gandhi, and James Zarucchi. Two re¬

viewers, Bruce Maslin and Gerardo A. Aymard C. provided many helpful comments on the manuscript. We

thank Carlos Leonardo Cespedes for linguistic assistance. This work was partly financed by the National Sci¬

ence Foundation (NSF) through research grants National Science Foundation (NSF BSR 82-15274, NSF-

PCM-82-17114, and NSF DEB 04-15803), grants by the University of Illinois Research Board (1994, 2001), the

United States Department of Agriculture (OICD 58-319R-0-0111), the American Philosophical Society (to DSS,

1992) and a Rupert Barneby Award by the New York Botanical Garden for (DSS) 1997. We wish to thank artist

Alexa Musgrove for preparation of the drawings.

REFERENCES

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39:452-477.

Bouchenak-Khelladi, Y., O, Maurin, J. Hurter, & M. Van Der Bank. 2010. The evolutionary history and biogeography of Mimo¬

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Gomez-Acevedo, S., L. Rico-Arce, A. Delgado Saunas, S. Magallon, & L.E. Eguiarte. 2010. Neotropical mutualism between Acacia

and Pseudomyrmex. Phylogeny and divergence times. Molec. Phylogen. Evol. 56:393-408.

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Gland, Switzerland and Cambridge, England.

Kyalangalilwa, B., J.S. Boatwright, B.H. Daru, 0. Maurin, & M. Van Der Bank. 2013. Phylogenetic position and revised classifica¬

tion of Acacia s.s. (Fabaceae: Mimosoideae) in Africa, including new combinations in Vachellia and Senegalia. Bot. J.

Linnean Soc., accepted and online.

Lewis, G.P., B. Schrire, B. MacKinder, & M. Lock. 2005. Legumes of the world. Royal Botanic Gardens, Kew, U.K.

Luckow, M, J.T. Miller, DJ. Murphy, &T Livschutz. 2003. A phylogenetic analysis of the Mimosoideae (Leguminosae) based

on chloroplast DNA sequence data. In: Klitgaard, B.B. and A. Bruneau, eds. Advances in legume systematics, Part 10:

Higher level systematic. Royal Botanic Gardens, Kew, U.K. Pp. 197-220.

Maslin, B.R. 2015. Synoptic overview of Acacia sensu lato (Leguminosae: Mimosoideae) in east and southeast Asia. Gard.

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Maslin, B.R., J.T. Miller, & D.S. Seigler. 2003a. Overview of the generic status of Acacia (Leguminosae: Mimosoideae).

Australian Syst. Bot. 16:1-18.

Maslin, B.R., A.E. Orchard, & J.G. West. 2003b. Nomenclatural and classification history of Acacia (Leguminosae: Mimosoi¬

deae), and the implications of generic subdivision. Available at: http://www.worldwidewattle.com.

Miller, J.T. & RJ. Bayer. 2003.Molecular phylogenetics of Acacia subgenus Acacia and Aculiferum (Fabaceae: Mimo-

sideae) based on the chloroplast matK coding sequence and flanking trnK intron spacer region. Australian Syst. Bot.

16:27-33.

Miller, J.T. & D.S. Seigler. 2012. Evolutionary and taxonomic relationships of Acacia s.l. (Leguminosae: Mimosoideae).

Australian Syst. Bot. 25:217-224.

Miller, J.T., J.W. Grimes, DJ. Murphy, R.J. Bayer, & P.Y. Ladiges. 2003. A phylogenetic analysis of the Acacieae and Ingeae (Mi-

modoideae: Fabaceae) based on trnK, matk, psbA-trnH, and trnL/trnF sequence data. Syst. Bot. 28:558-566.

Murphy, D.J., G.K. Brown, J.T. Miller, & P.Y. Ladiges. 2010. Molecular phylogeny of Acacia Mill. (Mimosoideae: Leguminosae):

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Rico-Arce, A.M.L. & S. Bachman. 2006. A taxonomic revision of Acacellia. Anales Jard. Bot. Madrid. 63:189-244.

Seigler, D.S., J.E. Ebinger, & J.T. Miller. 2006. New combinations in the genus Senegalia (Fabaceae: Minosoideae) from the

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344

Journal of the Botanical Research Institute of Texas 9(2)

BOOK NOTICE

Tom Kimmerer. 2015. Venerable Trees: History, Conservation, and Biology in the Bluegrass. (ISBN-13: 978-

0813165660, hbk). University Press of Kentucky, 663 South Limestone Street, Lexington, Kentucky

40508-4008, U.S.A. (Orders: www.kentuckypress.com, 1-800-53715487). $39.95, 288 pp., 120 b&w

photos, 118 color photos, 29 maps, 1 figure, 7" x 10".

From the Publisher: When the first settlers arrived in the Bluegrass region of Kentucky, they found an astonish¬

ing landscape of open woodland grazed by vast herds of bison. Farmers quickly replaced the bison with cattle,

sheep, and horses, but left many of the trees to shade their pastures. Today, central Kentucky and central

Tennessee still boast one of the largest populations of presettlement trees in the nation, found in both rural and

urban areas.

In Venerable Trees: History, Biology, and Conservation in the Bluegrass, Tom Kimmerer showcases the

beauty, age, size, and splendor of these ancient trees and the remaining woodland pastures. Documenting the

distinctive settlement history that allowed for their preservation, Kimmerer explains the biology of Bluegrass

trees and explores the reasons why they are now in danger. He also reveals the dedication and creativity of

those fighting to conserve these remarkable three-hundred- to five-hundred-year-old plants—from innova¬

tive, conscientious developers who build around them rather than clearing the land to farmers who use light¬

ning rods to protect them from natural disasters.

Featuring more than one hundred color photographs, this beautifully illustrated book offers guidelines

for conserving ancient trees worldwide while educating readers about their life cycle. Venerable Trees is an in¬

formative call to understand the challenges faced by the companions so deeply rooted in the region’s heritage

and a passionate plea for their preservation.

Tom Kimmerer is chief scientist at Venerable Trees, Inc., a nonprofit dedicated to the conservation of wood¬

land pastures and ancient trees in the Bluegrass. Trained as a tree physiologist and forest scientist, he has been

a researcher and teacher in the United States, Indonesia, and Malaysia.

Contents

Introduction: Two Trees

1. The St. Joe Oak: Finding Venerable Trees

2. The Woodford Groves: The Bluegrass Today

3. The Tlama Tree: The Nashville Basin Today

4. Venerable Tree Species

5. The Ingleside Oak: The Bluegrass and Nashville Basin in 1779

6. The Woodland Pasture

7. The Mother Tree: Reproduction of Venerable Trees

8. The Guardians: Trees in Cemeteries

9. The Toudon Grove: Trees in Public Spaces

10. The Coldstream Tree: Groves, Remnants, and Developments

11. The Elmwood Trees: Growing Old Gracefully

12. The Floracliff Trees: The Long Lives of Venerable Trees

13. The Old Schoolhouse Oak: Extending the Lives of Old Trees

14. The Future of Venerable Trees and Woodland Pastures

Acknowledgments

Appendix: Trees of the Bluegrass and Woodland Pastures

Notes

Index

J.Bot. Res. Inst. Texas 9(2): 344.2015

NAUTILOCALYX RUGOSUS (GESNERIACEAE), A NEW SPECIES FROM THE

RIO CENEPA WATERSHED (AMAZONAS, PERU)

Rocio del P. Rojas G.

Herbario Selva Central Oxapampa (HOXA)

Prolongacion Bolognesi Mz. E, Lote 6

Oxapampa-Pasco, PERU

gesneria@yahoo.com

John L. Clark

Department of Biological Sciences

The University of Alabama, Box 870345

Tuscaloosa, Alabama 35487 U.S.A. and

Science Department, The Lawrenceville School

2500 Main St., Lawrenceville, New Jersey 08648, U.S.A.

jlc@ua.edu

ABSTRACT

A recent botanical inventory of the sandstone outcrops in Tayu mujaji of the rio Cenepa watershed of northeastern Peru (Amazonas prov¬

ince) has resulted in the discovery of a new species, Nautilocalyx rugosus R. Rojas & J.T. Clark of the Gesneriaceae (tribe: Gesnerieae and

subtribe: Columneinae). The new species is distinguished from other species in the genus by a saxicolous habit, basal corolla spur, and mark¬

edly bullate leaves with a single elongate trichome at the apex of each bullation. The habitat of the type locality is a blackwater river on quartz

sandstone in an area known locally as “cerro de los Tayus” and represents one of most botanically unexplored regions of Peru.

Keywords: Nautilocalyx, Gesneriaceae, Sandstone, Cenepa watershed

RESUMEN

El inventario botanico realizado recientemente en el Tayu mujaji (Amazonas, Peru) en la cuenca del rio Cenepa, sobre los afloramientos de

roca de arenisca, ha dado como resultado el descubrimiento de una nueva especie, Nautilocalyx rugosus R. Rojas & J.L. Clark de la familia

Gesneriaceae (tribu: Gesnerieae y subtribu: Columneinae). La nueva especie se distingue de las otras especies del genero por un conjunto de

caracteres que incluyen el habito saxicola, corola con espolon basal, y hojas marcadamente ampolladas, con un solo tricoma alargado en el

apice de cada ampolla. La localidad tipo es conocida localmente como “cerro de los Tayus” y representa una de las areas botanicamente in-

exploradas del Peru.

Palabras Clave: Nautilocalyx, Gesneriaceae, roca de arenisca, cuenca del rio Cenepa

INTRODUCTION

The genus Nautilocalyx Linden ex Hanst., includes 58+ species (Moller & Clark 2013; Feuillet 2008) and is

neotropical in distribution from Mexico to Peru, the Guianas and Amazonian Brazil. The center of diversity for

Nautilocalyx is the Venezuelan Guayana where more than 19 species occur (Feuillet 2008; Feuillet & Steyer-

mark 1999). There are 12 species of Nautilocalyx in Peru (Kvist et al. 2005) and five have been documented in

the rio Cenepa watershed of northeastern Peru (Rojas 2010).

Nautilocalyx is a member of the tribe Gesnerieae and subtribe Columneinae, which is the largest subtribe

with 24 genera and 21% of the total species diversity in the family (Weber et al. 2013). Characters that define

most species of Nautilocalyx include a terrestrial habit, isophyllous leaves, subglobose capsule, and infundibu¬

lar corolla (Wiehler 1983). Recent molecular-based phylogenetic analyses with a broad taxon sampling of

Nautilocalyx strongly support that it lacks monophyly (Mora and Clark, in press). A new generic classification

based on recent phylogenetic results is in the process of being published (Mora & Clark in press; Mora & Clark

2014) and we predict that Nautilocalyx rugosus will nest with other traditionally recognized members of Nauti¬

localyx that are also found in the Amazon Basin or eastern slopes of the Andes.

TAXONOMIC TREATMENT

Nautilocalyx rugOSUS R. Rojas &J.L. Clark, sp. nov. (Figs. 1, 2). Type: PERU. Amazonas: Bagua province, Imaza district,

lower slopes of Cerro Tayu, ca., 1 hour from Chiriaco, vegetation “Kampau” or “Campau,” soil with thick humus layer, 05°15'56"S,

78°22 , 07"W, 900 m, 28 October 2012, H. van der Werff, Rocio Rojas, Luis Valenzuela, Gerry Shareva, Ricardo Apanu, Alcides Roca &

Augusto Reyes Barrantes 24654 (holotype: HOXA; isotypes: HUT, MO, MOL, USM).

J. Bot. Res. Inst. Texas 9(2): 345 - 349.2015

346

Journal of the Botanical Research Institute of Texas 9(2)

UBICACION DEL AREA DE ESTUDIO

Fig. 1. Distribution of Nautilocalyxrugosus R. Rojas &J.L. Clark. Arrows indicate general region of collections and specific collections are indicated by

asterisks (map reproduced from Vasquez et al. 2010 and reprinted with permission from Missouri Botanical Garden Press).

Differs from other species in the genus by a saxicolous habit, basal corolla spur, and bullate leaves with a single elongate trichome at the apex

of each bullation.

Saxicolous herb; appressed to rock wall, stems short, 5-10 cm long, greenish-brown, rhizomes succulent, te¬

rete to slightly sulcate, densely tomentose with uniseriate trichomes; adventitious roots present on the stems.

Leaves opposite, subequal to slightly unequal in a pair, petiolate; petioles terete, 4-20 mm long, tomentose;

blades obovate to narrowly oblong, 4.5-7.5 x 2.2-3.8 cm, base symmetrical to slightly oblique, rounded to

slightly cordate, apex rounded, margin crenate with broad shallow teeth, ciliated along margins, adaxially

dark green to light green when fresh and dark brown when dry, markedly bullate with elongate trichome at

apex of each bullation, bullae 1-3 mm high, blades coriaceous and membranous when dry, abaxially purple,

reticulated, mostly tomentose on veins. Inflorescence a reduced pair-flowered cyme, flowers appearing clus¬

tered in leaf axils, prophyll lanceolate; peduncle 2-3 mm long, in upper leaf axils, 1-5 mature flowers per inflo¬

rescence; pedicels erect, 9-15 mm long, reddish, densely pilose. Calyx lobes fused at base and appearing free,

narrowly ovate, 5—7(—11) mm long, apex narrowly acute, margins with sparse deep serrations, erect, mostly

Rojas and Clark, A new species of Nautilocalyx from Peru

347

Fig. 2. Nautilocalyxrugosus R. Rojas &J.L. Clark. A. Habit. B. Corolla lobes. C. Enlargement of leaf surface showing bullae with elongated trichome on

top of each bullation. D. Corolla tube. E. Corolla spur. (A-E photographic images of the holotype, H. van der Werffetal. 24654).

348

Journal of the Botanical Research Institute of Texas 9(2)

light red with green towards apex, densely pilose on outer surface and glabrous on inner surface. Corolla pos¬

ture slightly oblique relative to the calyx, 2.5-4 cm long, tube funnelform, spurred at base, spur 3-5 mm long,

base-region 2-3 mm in diameter, mid-region 5-7 mm in diameter, throat 10-12 mm wide at apex, uniformly

white, outer surface of corolla tube pilose, throat sparsely pilose, limb glabrous; limb bilaterally symmetrical,

lobes rehexed to patent, nearly equal, lobes suborbicular; upper two lobes 6-8 x 5-10 mm, slightly crenate;

lower three lobes 10-11 x 10-11 mm. Androecium with 4 stamens, didynamous, included; filaments hat,

curved at anthesis, adnate to the base of the corolla, free for 10-18 mm, filaments glabrous; anthers broader

than long, 0.7 x 1.0 mm, dehiscing by longitudinal slits; staminode absent. Gynoecium with dorsal nectary

gland, glabrous, ca. 1 mm long; ovary ovoid, 3x1 mm, densely sericeous, style 2.5 cm long, with glandular

capitate trichomes; stigma bilobed with glandular capitates trichomes. Fruits not observed.

Nautilocalyx rugosus is characterized by a saxicolous habit, succulent to heshy leaf blades, and conspicu¬

ous bullae with a single erect trichome on the apex of each bullation (Fig. 2C). The conspicuous bullations

appear like blisters and reach 3 mm in height. Other Nautilocalyx species with bullate leaves are Nautilocalyx

bullatus (Lem.) Sprague and N. pemphidius L.E. Skog. Nautilocalyx rugosus is differentiated by elongate shoots

(vs. basal rosette in N. pemphidius) and non-hmbriate corolla margins (Fig. 2B) (vs. fimbriate corolla margins

in N. bullatus).

Distribution and habitat.—Nautilocalyx rugosus is endemic to the Tayu mujaji region (Dept. Amazonas,

Prov. Bagua) of northwestern Peru (Fig. 1). Cerro Tayu is a low sandstone mountain largely covered with veg¬

etation that is referred to locally by the Awajun Indigenous people as “Campau” or “Kampau” and corresponds

to an area between 650 and 1200 m. The soils of this region are characterized by a thick layer of humus (litter)

and shallow soils. The area is not suitable for agriculture because of the abundance of exposed sandstone and

shallow soils (Vasquez et al. 2010). Nautilocalyx rugosus is endemic to this region and is mostly found on cliffs

or exposed sandstone. Other plants that grow sympatrically with N. rugosus include Alloneuron ronliesneri B.

Walln. (Melastomataceae), Monolena primuliflora Hook. f. (Melastomataceae), Triolena pluvialis (Wurdack)

Wurdack (Melastomataceae), Peperomia sp. (Piperaceae), Pilea sp. (Urticaceae), Pterozonium reniforme (Mart.)

Fee (Pteridaceae), Guzmania gracilior (Andre) Mez (Bromeliaceae), Cyathea thelypteroides A.R. Sm. (Cyatheace-

ae), and Trichomanes pilosum Raddi (Hymenophyllaceae). This area is considered one of the most botanically

unexplored regions of Peru (Vasquez et al. 2010) and another recently described species from the Gesneriaceae

that is a local endemic is Cremosperma inversum B.R. Keener & J.L. Clark (Keener & Clark 2014).

Etymology. —The specific epithet is in reference to the rugose surface of the leaf blades that are deeply fur¬

rowed like a wrinkle.

Conservation and IUCN Red List category.—Nautilocalyx rugosus is geographically limited to a small area

in the Cenepa watershed of northeastern Peru. According to the IUCN Red List criteria (IUCN 2012) for lim¬

ited geographic range (B2a, less than 10 km 2 and known to exist at only a single location), Nautilocalyx rugosus

should be listed in the category CR (Critically Endangered).

Additional specimens examined: PERU. Amazonas: Bagua province, Imaza district, Quebrada Almendro, along road and in forest on sand¬

stone, 05°14 , 40"S, 78°2r34" W, 400 m, 17 Mar 2001, H. van der Werff Rodolfo Vdsquez & Bruce Gray 16161 (MO); sandstone face, Cerro Tayu,

ca. 1 hour from Chiriaco, 05°15 , 56"S, 078°22 , 07"W, 800 m, H. van der Werff et al 16235 (MO, USM).

ACKNOWLEDGMENTS

The first author is grateful to the Missouri Botanical Garden (MO) and the Herbario Selva Central (HOXA) for

access to their collections; El Servicio Nacional de Areas Naturales Protegidas por el Estado (SERNANP) for

research permits and authorization to work in Amazonas (Peru); Olga Martha Montiel for her support through

Missouri Botanical Garden Peru Program; Rodolfo Vasquez for suggestions on an early version of the manu¬

script; Henk van der Werff for his enormous contributions to botanical research in Peru and comments on the

manuscript. Ricardo Apanu is gratefully acknowledged for guiding us throughout the floristic inventory of the

rlo Cenepa watershed and for bringing us to Tayu. We also thank fellow held botanists Luis Valenzuela, Al-

cides Roca and Gerry Shareva who assisted the 2012 expedition to Tayu Hill. Support for JLC was provided by

Rojas and Clark, A new species of Nautilocalyx from Peru

349

the National Science Foundation (DEB-0841958 and DEB-0949169). We thank the Missouri Botanical Garden

Press for allowing us to reproduce the map of the Cenepa watershed (Figure 1).

REFERENCES

Clark, J.L., M.M. Funke, A.M. Duffy, & J.F. Smith. 2012. Phylogeny of a Neotropical clade in the Gesneriaceae: more tales of

convergent evolution. Inti. J. PI. Sci. 173:894-916.

Clark, J.L., P.S. FIerendeen, L.E. Skog, & E.A. Zimmer. 2006. Phylogenetic relationships and generic boundaries in the Epi-

scieae (Gesneriaceae) inferred from nuclear, chloroplast, and morphological data.Taxon 55:313-336.

Feuillet, C. 2008. Folia Taxonomica 10. New species of Nautilocalyx (Gesneriaceae: Episcieae) from the Venezuela Guay-

ana. J. Bot. Res. Inst.Texas 2:825-836.

Feuillet, C. & J.A. Steyermark. 1999. Gesneriaceae. In: Flora of the Venezuelan Guayana, Volume 5, P.E. Berry, K. Yatskievych,

and B.K. Holst eds., Missouri Botanical Garden Press, St. Louis, Missouri, U.S.A. Pp. 542-573.

Iucn. 2012. IUCN Red List Categories and Criteria: Version 3.1. Second edition. Prepared by the IUCN Species Survival

Commission. Gland, Switzerland and Cambridge, UK. International Union for Conservation of Nature and Natural

Resources (IUCN).

Keener, B.&J.L. Clark. 2014. A new species of Cremosperma (Gesneriaceae) from northeastern Peru. J. Bot. Res. Inst.Texas

8:57-60.

Kvist, L.P. L.E. Skog, M. Amaya-Marquez, & I. Salinas. 2005. Las Gesneriaceas de Peru. Arnaldoa 12:16-40

Moller, M. & J.L. Clark. 2013. The state of molecular studies in the family Gesneriaceae. Selbyana 31:95—125.

Mora, M. & J.L. Clark. 2014. Nautilocalyx erytranthus (Gesneriaceae), a new species from Northwestern Amazonia. Phy-

totaxa 164:183-189

Mora, M. & J.L. Clark, in press. A new generic classification based on recent phylogenetic results. Syst. Bot.

Rojas, R. 2010. Gesneriaceae. In: Vasquez, R., G.R. Rojas, and H. van derWerff, eds., Flora del Rio Cenepa, Amazonas, Peru.

Missouri Botanical Garden Press, St. Louis, Missouri, USA. Pp. 785-808

Vasquez, R., G.R. Rojas, & H. van derWerff. 2010. Flora del Rio Cenepa, Amazonas, Peru. St. Louis, Missouri, USA: Missouri

Botanical Garden Press.

Weber, A., J.L. Clark, & M. Moller. 2013. A new formal classification of Gesneriaceae. Selbyana 31:68-94.

Wiehler, H. 1983. A synopsis of the Neotropical Gesneriaceae. Selbyana 6:1 -219.

350

Journal of the Botanical Research Institute of Texas 9(2)

BOOK NOTICE

Mary Siisip Geniusz. 2015. Plants Have So Much to Give Us, All We Have to Do is Ask: Anishinaabe botani¬

cal teachings. (ISBN-13: 978-0-8166-9676-5, pbk). University of Minnesota Press, 111 Third Avenue

South, Minneapolis, Minnesota 55401, U.S.A. (Orders: www.upress.umn.edu). $22.95,344pp., 37b&w

photos, 7" x 10".

From the Publisher: Mary Siisip Geniusz has spent more than thirty years working with, living with, and using

the Anishinaabe teachings, recipes, and botanical information she shares in Plants Have So Much to Give Us, All

We Have to Do Is Ask. Geniusz gained much of the knowledge she writes about from her years as an oshkaabe-

wis, a traditionally trained apprentice, and as friend to the late Keewaydinoquay, an Anishinaabe medicine

woman from the Leelanau Peninsula in Michigan and a scholar, teacher, and practitioner of native ethnobota-

ny. Keewaydinoquay published little, yet Geniusz has carried on her legacy by making this knowledge acces¬

sible to a broader audience.

Geniusz teaches the ways she was taught—through stories. Sharing the traditional stories she learned at

Keewaydinoquay’s side as well as stories from other American Indian traditions and her own experiences,

Geniusz brings the plants to life with narratives that explain their uses, meaning, and history. Stories such as

“Naanabozho and the Squeaky-Voice Plant” place the plants in cultural context and illustrate the belief in

plants as cognizant beings. Covering a wide range of plants, from conifers to cattails to medicinal uses of yar¬

row, mullein, and dandelion, Geniusz explains how we can work with these botanical beings to create food,

simple medicines, and practical botanical tools.

Plants Have So Much to Give Us, All We Have to Do Is Ask makes this botanical information available to na¬

tive and nonnative healers and educators and emphasizes the Anishinaabe culture that developed the knowl¬

edge and practice.

Mary Siisip Geniusz is of Cree and Metis descent and an oshkaabewis, a traditionally trained apprentice, of

the late Keewaydinoquay. She holds a master’s degree in liberal studies from the University of Wisconsin-

Milwaukee and has taught university courses on ethnobotany, American Indian studies, and American multi¬

cultural studies.

J.Bot. Res. Inst. Texas 9(2): 350.2015

CORRIGENDUM:

NEW COMBINATIONS IN CORYPHANTHA AND ESCOBARIA (CACTACEAE)

Root Gorelick

Department of Biology, School of Mathematics & Statistics, and

Institute of Interdisciplinary Studies

Carleton University, 1125 Raven Road

Ottawa, Ontario K1S 5B6, CANADA

RootGorelick@cadeton.ca

In the previous issue (9, no. 1, 2015) of this journal, I provided two new combinations— Coryphantha sneedii

var. orcuttii and Escobaria sneedii var. orcuttii —based on the same basionym, Escobaria orcuttii Boed. (Gorelick

2015). Shortly thereafter, David Hunt kindly informed me that this contravenes the International Code of No¬

menclature (Melbourne Code) Article 36.2 (McNeill et al. 2012) [previously the International of Botanical

Nomenclature (Vienna Code) (McNeill et al. 2006) Article 34.2], which states:

When, on or after 1 January 1953, two or more different names based on the same type are proposed simultaneously for the

same taxon by the same author (so-called alternative names), none of them is validly published.

Therefore, I hereby correct this error by only proposing one of those two combinations, namely:

Coryphantha sneedii var. orcuttii (Boed.) Gorelick, comb, et stat. nov. Basionym: Escobaria orcuttii Boed., Ein Mammil-

larien Vergleichs-Schliissel 17.1933. Type: U.S.A. New Mexico. Hidalgo Co.: Peloncillo Mountains, near Granite Pass, Mar 1926, J.N.

Rose s.n. (lectotype, designated by Benson 1969: 26: DS).

Coryphantha strobiliformis (Poselg.) Moran var. orcuttii (Rose ex Orcutt) L.D. Benson, Cacti Ariz. ed. 3, 26.1969.

Coryphantha orcuttii (Rose ex Orcutt) Zimmerman, Cact. Succ. J. (US) 44:156.1972.

Coryphantha organensis Zimmerman, Cact. Succ. J. (US) 44:114.1972.

Escobaria organensis (Zimmerman) Castetter, P. Pierce & K.H. Schwer., Cact. Succ. J. (US) 47:60.1975.

Escobaria sneedii Britton & Rose subsp. orcuttii (Boed.) Luthy, Kakt. and. Sukk. 50:278.1999.

Escobaria sneedii Britton & Rose subsp. organensis (Zimmerman) Lu thy, Kakt. and. Sukk. 50:278.1999.

That previous paper in this journal (Gorelick 2015) was predicated on (1) Escobaria orcuttii being a variety,

rather than a subspecies, because of sympatry with other taxa and (2) Escobaria Britton & Rose being either a

subgenus or section of Coryphantha (Engelm.) Lem. Many European botanists have abandoned use of the rank

variety (‘varietas’). The Cactaceae Working Party of the International Organization for Succulent Plant Study

(IOS) has also abandoned use of the rank variety, albeit apparently without complete consensus as reflected by

the opening quote in that paper (Hunt 1999: 23; quoting Meregalli 1993):

Probably one category only, below the species, would do for the great majority of infraspecific taxa, although in (possibly very few)

cases some striking variants of low systematic value (i.e. not related to geographic/ecological isolation) may warrant recognition.

If Escobaria orcuttii is considered a subspecies of Coryphantha sneedii, which I do not advocate because of sym¬

patry with the type variety/subspecies of Coryphantha sneedii at Anthony’s Nose, then someone will have to

propose the new combination Coryphantha sneedii subsp. orcuttii. If Escobaria orcuttii is considered a variety of

Escobaria sneedii, which I do not advocate because of lack of consistent morphological differences between

Coryphantha and Escobaria, then someone will have to propose the new combination Escobaria sneedii var.

orcuttii, which was a nomen nudum in Gorelick (2015). Another plausible combination for this taxon, one for

which I do not agree with the subspecies designation nor the choice of genus, would be Escobaria sneedii subsp.

orcuttii (Boed) Luthy, which has been validly published. A final possibility is that Coryphantha orcuttii is

merely a form (‘forma’) of Coryphantha sneedii. But I am reluctant to invoke this last option without spending

more time in Luna and Hidalgo Counties of New Mexico, especially given the seemingly consistent morpho-

J. Bot. Res. Inst. Texas 9(2): 351 - 352.2015

352

Journal of the Botanical Research Institute of Texas 9(2)

logical differences between Coryphantha sneedii and Coryphantha orcuttii in terms of number of shoots, size of

shoots, and length of central spines (Gorelick 2015 and references therein).

ACKNOWLEDGMENTS

I greatly appreciate David Hunt’s input and help with interpretation of the International Code of

Nomenclature.

REFERENCES

Gorelick, R. 2015. New combinations in Coryphantha and Escobaria (Cactaceae). J. Bot. Res. Inst. Texas 9:25-30.

Hunt, D. R., ed. 1999. What to do about the less-than-species? - resume of workshop 5 at the Cactaceae Concensus Initia¬

tives meeting, Milbourne Port, U.K., 10 April 1994. Cactaceae Consensus Init. 8:23-28.

McNeill, J., F.R. Barrie, W.R. Buck, V. Demoulin, W. Greuter, D.L. Hawksworth, P.S. Herendeen, S. Knapp, K. Marhold, J. Prado, W.F.

Prud'hommeVan Reine, G.F. Smith, J.H. Wiersma, & NJ. Turland, eds. 2012. International code of nomenclature (Melbourne

Code) for algae, fungi, and plants, Regnum Veg. 154.

McNeill, J., F.R. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, K. Marhold, D.H. Nicholson, J. Prado, P.C. Silva, J.E. Skog, J.H.

Wiersma, & N.J. Turland, eds. 2006. International code of botanical nomenclature (Vienna Code), Regnum Veg. 146.

Meregalli, M. 1993. A note on infraspecific terminology in the Cactaceae. I.O.S. Bull. 5:214-216.

PSEUDHAPLOCRICUS HEXANDRUS GEN. ET SP. NOV. (COMMELINACEAE)

IN MID-TERTIARY DOMINICAN AMBER

George O. Poinar, Jr.

Kenton L. Chambers

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

poinarg@science.oregonstate.edu

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

ABSTRACT

Pseudhaplocricus hexandrus gen. et sp. nov. (Commelinaceae) is described from Mid-Tertiary amber from the Dominican Republic. The

trimerous, staminate flower is characterized by a short pedicel, 2 perianth whorls with 3 glabrous, free sepals and 3 glabrous, free, deliques¬

cent petals. The 6 fertile, glabrous stamens are of equal length; their apparent arrangement in a single whorl is due to fusion, during floral

development, of the 2 whorls of 3 which are characteristic of Commelinaceae. The filaments are broadened and united basally. The anthers

are bilocular, dorsifixed and dehisce longitudinally. The fossil establishes a mid-Tertiary lineage of Commelinaceae in Mesoamerica.

RESUMEN

Se describe Pseudhaplocricus hexandrus gen. et sp. nov. (Commelinaceae) del ambar del Terciario Medio de la Republica Dominicana. Las

flores trimeras, estaminadas se caracterizan por un pedicelo corto, 2 verticilos de perianto con tres sepalos libres glabros y tres petalos li-

bres, glabros delicuescentes. Los seis estambres fertiles son de igual longitud; su ordenacion aparente en un verticilo es debida a su fusion

durante el desarrollo floral, de dos verticilos de los tres que son caracteristicos de las Commelinaceae. Los filamentos estan ensanchados y

unidos basalmente. Las anteras son biloculares, dorsifijas y dehiscencia longitudinal. El fosil establece una linea del Terciario Medio de

Commelinaceae en Mesoamerica.

INTRODUCTION

Amber from the Dominican Republic has provided a rare glimpse of the flora that existed in the West Indies

during the mid-Tertiary. The present study describes a fossil member of the Commelinaceae, a pantropical

family of annual or perennial monocotyledonous herbs with some 650 species and 42 genera. While distrib¬

uted worldwide, the family is most abundant in the tropics and subtropics with centers of diversity in the

Americas, Africa and India (Faden 1998; Hardy & Faden 2004). Some species of Commelinaceae, such as

Dayflowers ( Commelina spp.) and Spiderworts ( Tradescantia spp.) are widespread and cultivated as garden,

greenhouse, basket and border plants for their colorful flowers and leaf characters (Bailey 1949).

Only two previously known fossil Commelinaceae have been reported. One is a fossil flower, Commelina-

cites dichorisandroides Caspary, in Eocene Baltic amber (Caspary 1880; Conwentz 1886), and the other in¬

cludes leaves and fruits assigned to the genus Pollia from the Middle Miocene Ngorora Formation in Kenya

(Jacobs & Kabuye 1989). The present specimen is the first New World fossil of the family.

MATERIALS AND METHODS

The fossil originated from amber mines in the northern mountain range (Cordillera Septentrional) of the Do¬

minican Republic between Puerto Plata and Santiago. Amber from mines in this region was produced by the

leguminous tree Hymenaea protera (Poinar 1991). Dating of Dominican amber is still controversial, with the

youngest proposed age of 20-15 Ma based on foraminifera (Iturralde-Vincent & MacPhee 1996) and the oldest

of 45-30 Ma based on coccoliths (Cepek in Schlee 1990). These are considered minimum dates since they are

based on microfossils in the strata containing the amber. Most of the amber is secondarily deposited in turbid-

itic sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994). Dilcher et al. (1992)

stated that “...the amber clasts, from all physical characteristics, were already matured amber at the time of re¬

deposition into marine basins. Therefore, the age of the amber is greater than Miocene and quite likely is as

early as late Eocene.” The issue is further complicated by the discovery of Early Oligocene amber in Puerto Rico

J. Bot. Res. Inst. Texas 9(2): 353 - 359.2015

354

Journal of the Botanical Research Institute of Texas 9(2)

and Maastrichtian-Paleocene amber in Jamaica (Iturralde-Vinent 2001), showing that amber from a range of

deposits occurs in the Greater Antilles. The Dominican amber forest was characterized by Poinar and Poinar

(1999) based on various animal and plant fossils.

Observations and photographs were made with a Nikon SMZ-10 R stereoscopic microscope and Nikon

Optiphot compound microscope with magnifications up to 600 X. Helicon Focus Pro X64 was used to stack

photos for better clarity and depth of held.

DESCRIPTION

Pseudhaplocricus Poinar & K.L. Chambers, gen. nov. (Figs. 1-4). Type Species: Pseudhaplocricus hexandrus Poinar & K.L.

Chambers, sp. nov.

Flower actinomorphic, staminate, pistillode none; perianth with 2 whorls, the outer whorl of 3 glabrous, free

sepals (Fig. 1); inner whorl petaloid with 3 free, glabrous, deliquescent petals lacking a fringed margin (Fig. 2);

fertile stamens 6, of equal length, basally connate in a single whorl at maturity although developing from the

bicyclic androecium characteristic of Commelinaceae (Hardy et al. 2009); anthers bilocular, dorsihxed, de¬

hiscing longitudinally, with narrow connective and spiral-reticulate secondary endothecial thickenings (Fig.

3); filaments glabrous, broadened at base (Figs. 3,4).

Etymology. —From the Greek “pseudo,” false, “haploos,” single, and “krikos,” ring, in reference to the

single whorl of 6 stamens originating from 2 whorls of 3.

Pseudhaplocricus hexandrus Poinar & K.L. Chambers, sp. nov. Type: HISPANIOTA. Dominican Republic: amber mine in

the northern mountain ranges (Cordillera Septentrional), 1986, unknown amber miner s . n . (holotype: catalog number Sd-9-118;

deposited in the Poinar amber collection maintained at Oregon State University, Corvallis, Oregon 97331, U.S.A.).

Flower diameter 3.3 mm across opposing anthers; calculated diameter based on detached petals 8.5 mm; pedi¬

cel length 0.5 mm; sepals ovate-elliptic, length 07(0.5-0.9) mm, width at base 1.0(0.9-1.2) mm; petals de¬

tached in amber, partially liquefied, clawed, only 1 outstretched, length 4.3 mm, length of claw 0.9 mm; sta¬

men filament length 07(0.5-1.0) mm, ring of connected filament bases 1.0 mm in diameter; anther length

1.0(0.9-11) mm, pistil 0.

Etymology. —From the Greek “hexa,” six, and “andro,” man, in reference to the 6 fertile stamens.

DISCUSSION

The original color of the petals and sepals is unknown, since fading of colors occurs over time in amber. Char¬

acters of Pseudhaplocricus hexandrus that are consistent with those of the family Commelinaceae are: unisex¬

ual male flower with 2 trimerous perianth whorls (3 sepals and 3 deliquescent petals), 6 fertile stamens with

glabrous filaments and dorsihxed anthers that dehisce longitudinally, and abortion of the gynoecium (Faden

1998; Hardy & Faden 2004).

Based on the classification of Faden and Hunt (1991), the fossil falls in the tribe Tradescantieae Meisner of

the subfamily Commelinoideae Bruckner. This tribe comprises the New World subtribes Dichorisandrinae

Pichon, Thyrsantheminae Faden & D. Hunt, and Tradescantiinae Rohw. It includes species from Mexico,

Central America, the Caribbean, and South America. Genera in the above subtribes with characters similar to

Pseudhaplocricus hexandrus are Tradcscantia L., whose Bowers are long-pedicelled with some or all Blaments

bearded, Callisa Loefl., whose species mostly have 1-3 stamens, Tripogandra Raf., which has zygomorphic

Bowers, Cochliostema Lem., and Geogenanthus Ule, which have fringed petals. Siderasis and Dichorisandra are

sister genera in the subtribe Dichorisandrinae (Evans et al. 2000) which are noteworthy since both contain

species with actinomorphic flowers, entire petals, and six glabrous stamens with longitudinally dehiscent

anthers as in Pseudhaplocricus. Siderasis consists of one Brazilian species, Siderasis fuscata (Lodd.) H.E. Moore.

The anther connectives of Siderasis, however, are broader than in Pseudhaplocricus, and the degree of filament

fusion, if any, in Siderasis has not been well studied with the microscope but would be slight relative to length

of the filaments in the mature flowers. Dichorisandra consists of over 40 species, second in the New World only

Poinar and Chambers, Pseudhaplocricus hexandrus, a new Commelinaceae from Mid-Tertiary amber

355

Fig. 1. Pseudhaplocricus hexandrus. Apical view of flower, showing 3 sepals and 6 fertile stamens arranged in a single whorl. Scale bar = 0.6 mm.

to Tradescantia , and is distributed from tropical South America to the Caribbean. Although zygomorphy and

poricidal anther dehiscence prevail in the genus, there are some plesiomorphic members with actinomorphic

flowers and longitudinal anther dehiscence (Aona et al. 2012). All members have extremely narrow anther

connectives like Pseudhaplocricus, although the anthers in most Dichorisandra taxa are proportionally longer

relative to the filaments. Additionally, most members of Dichorisandra have very short pedicels. Basal fusion of

the filaments has been documented for at least two members of the genus, although in these instances the fila¬

ments also are fused to the petals.

356

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 2. Pseudhaplocricus hexandrus . Remains of detached, deliquescent petal. Arrows show smooth outer margin of petal. Scale bar = 0.2 mm.

Poinar and Chambers, Pseudhaplocricus hexandrus, a new Commelinaceae from Mid-Tertiary amber

357

Fig. 3. Pseudhaplocricus hexandrus. Stamen showing bilocular anther and glabrous filament whose cells appear empty, due perhaps to enlarged

vacuoles. Scale bar = 0.3 mm.

While many flowers in the Commelinaceae are considered to be insect-pollinated, none are known to

produce nectar, which means that the only reward for potential pollinators is the pollen itself. Pollination in

the Commelinaceae is mainly carried out by social and solitary bees and syrphid flies. The flowers attract in¬

sects by visual aids, e.g., color of foliage and flowers, fringed margins on the petals, filament hairs, or by various

fragrances (Faden 1992). Any features of the flowers that make pollen more accessible would be

advantageous.

The tissue connecting the filaments forms a ring around the center of the flower in Pseudhaplocricus. It is

possible that this dish-like structure provided mechanical support for the weight of a pollinator. From such a

perch, pollinators could easily reach the relatively large anthers on the outstretched filaments. In the case of the

Australian Pollia crispata (R. Br.) Benth., the petals act as landing platforms for bees and flies (Williams &

Walker 2003). The center of the flower of Pseudhaplocricus hexandrus is depressed and contains no trace of an

ovary or pistillode. It is tempting to speculate that it contained some type of liquid reward for pollinating in¬

sects, even though nectar production is unknown in extant flowers of the family (Faden 1992). Insect pollina¬

tors of Pollia crispata in Australia are syrphid flies and halictid and trigonid bees (Williams & Walker 2003).

Representatives of these groups have been recorded from Dominican amber (Poinar & Poinar 1999).

ACKNOWLEDGMENTS

The authors thanks Betsy Gates for determining the family of the fossil and Christopher Hardy for his useful

comments and suggestions.

358

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 4. Pseudhaplocricushexandrus. Apical view of flower showing tissue connections between filament bases of stamen whorl (arrows). Scale bar= 0.6 mm.

REFERENCES

Aona, L.Y.S., R.B. Faden, & M.E. Amaral. 2012. Five new species of Dichorisandra J.C. Mikan (Commelinaceae) from Bahia

State, Brazil. Kew Bull. 66:1 -13.

Bailey, L.H. 1949. Manual of cultivated plants. The MacMillan Company, New York, U.S.A.

Caspary, R. 1880. Commelinacites R. Caspary. Schriften der Koniglichen Physikalisch-Okonomischen Gesellschaft zu

Konigsberg, Sitzungsber. 21:29.

Conwentz, H. 1886. Die flora des Bernstein. Vol. 2. Wilhelm Engelmann Verlag, Leipzig, Germany.

Dilcher, D.L., P.S. Herendeen, & F. Huber. 1992. Fossil Acacia flowers with attached anther glands from Dominican Republic

amber. In: P.S. Herendeen and D.L. Dilcher, eds. Advances in legume systematics: 4. The fossil record. The Royal Bo¬

tanical Gardens, Kew, England. Pp. 33-42.

Draper, G., P. Mann, & J.F. Lewis. 1994. Hispaniola. In: S. Donovan andT.A. Jackson, eds. Caribbean geology: An introduc¬

tion. The University of the West Indies Publishers'Association, Kingston, Jamaica. Pp. 129-150.

Evans, T.M., R.B. Faden, M.G. Simpson, & KJ. Systma. 2000. Phylogenetic relationships in the Commelinaceae. I. A cladistic

analysis of morphological data. Syst. Bot. 25:668-691.

Faden, R.B. 1992. Floral attraction and floral hairs in the Commelinaceae. Ann. Missouri Bot. Gard. 79:46-52.

Poinar and Chambers, Pseudhaplocricus hexandrus, a new Commelinaceae from Mid-Tertiary amber

359

Faden, R.B. 1998. Commelinaceae. In: Kubitzki, K. ed.The families and genera of vascular plants. Vol. 4. Springer-Verlag,

Berlin, Germany. Pp.109-128

Faden, R.B. & D.R. FIunt. 1991. The classification of the Commelinaceae. Taxon 40:19-31.

Gerenday, A. & J.C. French. 1988. Endothecial thickenings in anthers of porate monocotyledons. Amer. J. Bot. 75:22-25.

FIardy, C.R. & R.B. Faden. 2004. Commelinaceae (Spiderwort family) In: N. Smith, S.A.Mori, A.Henderson, D.W. Stevenson,

& S.V. Heald, eds. Flowering plants of the Neotropics. Princeton University Press, Princeton, New Jersey. Pp. 425-429.

Hardy, C.R., L.L. Sloat, & R.B. Faden. 2009. Floral organogenesis and the developmental basis for pollinator deception in

the Asiatic dayflower Commelina communis (Commelinaceae). Amer. J. Bot. 96:1236-1244.

Iturralde-Vinent, M.A. & R.D.E. MacPhee. 1996. Age and paleographic origin of Dominican amber. Science 273:1850-1852.

Iturralde-Vinent, M.A. 2001. Geology of the amber-bearing deposits of the Greater Antilles. Caribbean J. Sci. 37:141-167.

Jacobs, B.F. & C.H.S. Kabuye. 1989. An extinct species of Pollia Thunberg (Commelinaceae) from the Miocene Ngorora

Formation, Kenya. Rev. Palaeobot. Palynol. 59:67-76.

Poinar, G.O., Jr. 1991. Hymenaea protero sp. n. (Leguminosae: Caesalpinioideae) from Dominican amber has African af¬

finities. Experientia 47:1075-1082.

Poinar, G.O., Jr. & R. Poinar. 1999. The amber forest. Princeton University Press, Princeton, New Jersey, U.S.A.

Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C, 28.

Williams, G. & K. Walker. 2003. Pollination of the wet forest herb Pollia crispata (Commelinaceae). Cunninghamia

8:141-146.

360

Journal of the Botanical Research Institute of Texas 9(2)

BOOK NOTICE

Fructuoso Irigoyen-Rascon, with Alfonso Paredes. 2015. Tarahumara Medicine: Ethnobotany and Healing

among the Raramuri of Mexico. (ISBN-13: 978-0-8061-4828-1, hbk). University of Oklahoma Press,

2800 Venture Drive, Norman, Oklahoma 73069, U.S.A. (Orders: www.oupress.com, 1-800-627-7377).

$49.95,336 pp., 22 b&w illus., 3 maps, 4 tables, 6" x 9".

From the Publisher: The Tarahumara, one of North America’s oldest surviving aboriginal groups, call them¬

selves Raramuri, meaning “nimble feet”—and though they live in relative isolation in Chihuahua, Mexico,

their agility in long-distance running is famous worldwide. Tarahumara Medicine is the first in-depth look into

the culture that sustains the “great runners.” Having spent a decade in Tarahumara communities, initially as a

medical student and eventually as a physician and cultural observer, author Fructuoso Irigoyen-Rascon is

uniquely qualified as a guide to the Raramuri’s approach to medicine and healing.

In developing their healing practices, the Tarahumaras interlaced religious lore, magic, and careful ob¬

servations of nature. Irigoyen-Rascon thoroughly situates readers in the Raramuri’s environment, describing

not only their health and nutrition but also the mountains and rivers surrounding them and key aspects of

their culture, from long-distance kick-ball races to corn beer celebrations and religious dances. He describes

the Tarahumaras’ curing ceremonies, including their ritual use of peyote, and provides a comprehensive de¬

scription of Tarahumara traditional herbal remedies, including their botanical characteristics, attributed ef¬

fects, and uses.

To show what these practices—and the underlying concepts of health and disease—might mean to the

Raramuri and to the observer, Irigoyen-Rascon explores his subject from both an outsider and an insider (in¬

digenous) perspective. Through his balanced approach, Irigoyen-Rascon brings to light relationships between

the Raramuri healing system and conventional medicine, and adds significantly to our knowledge of indige¬

nous American therapeutic practices.

As the most complete account of Tarahumara culture ever written, Tarahumara Medicine grants readers

access to a world rarely seen—at once richly different from and inextricably connected with the ideas and

practices of Western medicine.

Fructuoso Irigoyen-Rascon is a psychiatrist in McAllen, Texas. A former researcher at universities in Mexico

and the United States, he has written extensively about Raramuri ethnography and medical conditions.

Alfonso Paredes is Professor of Psychiatry at the University of California-Los Angeles and author of more

than 100 medical papers, including several on the Tarahumara.

J.Bot. Res. Inst. Texas 9(2): 360.2015

COMOPELLIS PRESBYA GEN. ET SP. NOV. (RHAMNACEAE) IN

MID-TERTIARY AMBER FROM THE DOMINICAN REPUBLIC

Kenton L. Chambers

Department of Botany & Plant Pathology

Oregon State University

Corvallis, Oregon U.S.A. 97331

chamberk@science.oregonstate.edu

George O. Poinar, Jr.

Department of Integrative Biology

Oregon State University

Corvallis, Oregon U.S.A. 9733 7

ABSTRACT

Comopellis presbya is described as a new genus and species of Rhamnaceae, based on a fossil flower preserved in amber from Mid-Tertiary

deposits in the Dominican Republic. The fossil consists of a single pentamerous, bisexual flower at anthesis, characterized by narrowly

lanceolate-elliptic petals with involute margins that partially cover the short, appressed stamen, together with a bowl-shaped hypanthium

enclosing the sessile, superior pistil. The thin disc lining the hypanthium has 5 conspicuous appendages, 1 opposite the base of each sepal,

which may have functioned as nectaries. The style is short, not surpassing the hypanthium, and bears a discoid, non-lobed stigma. With

respect to its disc appendages, the genus is most similar to Gouania and Distigouania of the tribe Gouanieae (Medan & Schirarend 2004).

RESUMEN

Se describe Comopellis presbya como Nuevo genero y especie de Rhamnaceae, basados en una flor fosil preservada en ambar de los deposi-

tos del Terciario Medio en la Republica Dominicana. El fosil consiste en una flor simple pentamera y bisexual en antesis, que se caracteriza

por los petalos estrechamente lanceolado-elipticos con margenes involutes que cubren parcialmente el estambre corto y apresado, junto con

un hipanto en forma de taza que encierra el pistilo supero sesil. El disco fino de revestimiento del hipanto tiene 5 apendices conspicuos,

opuestos a la base cada sepalo, que pudieran haber funcionado como nectarios. El estilo es corto, no sobrepasa el hipanto, y lleva un estigma

discoide no lobulado. Con respecto a los apendices del disco, el genero es muy similar a Gouania y Distigouania de la tribu Gouanieae (Medan

& Schirarend 2004).

INTRODUCTION

Knowledge of the flora of tropical forests present in the Caribbean region in the Mid-Tertiary has improved

through the recent description of fossil flowers preserved in amber from the Dominican Republic. Insects and

plants of this forest were well treated by Poinar and Poinar (1999), based on fossils known at the time. Since

then, some 23 new plant species, including a number of previously unknown genera, have been published.

These include 3 species of Fabaceae (Poinar 1991; Dilcher et al. 1992; Poinar & Chambers 2015a), 2 of Poaceae

(Poinar & Judziewicz 2005; Poinar & Columbus 2012), 3 of Arecaceae (Poinar 2002a, 2002b), 1 of

Chrysobalanaceae (Poinar et al. 2008a, revised by Chambers & Poinar 2010), 2 of Lauraceae (Chambers et al.

2011a, 2012), 3 of Meliaceae (Chambers et al. 2011b; Chambers & Poinar 2012), 1 of Burseraceae (Chambers &

Poinar 2013), 1 of Myristicaceae (Poinar & Steeves 2013), 1 of Rhamnaceae (Chambers & Poinar 2014a), 1 of

Ticodendraceae (Chambers & Poinar 2014b), 1 of Commelinaceae (Poinar & Chambers 2015c), 1 possibly of

Moraceae (Poinar et al. 2008b), and 1 of an unknown monocotyledonous family (Poinar & Chambers 2015b).

A previously published fossil of Rhamnaceae from this flora, the genus Distigouania, is notable in being the brst

known member of the family with an irregular corolla and androecium (Chambers & Poinar 2014a).

MATERIALS AND METHODS

Comopellis was collected from an amber mine in the northern mountain ranges (Cordillera Septentrional) of

the Dominican Republic, between Puerto Plato and Santiago. Amber from mines in this region was produced

by the leguminous tree Hymenaea protera (Poinar 1991). The age of the amber is controversial, with dates of

15-20 Ma and 30-45 having been proposed (Iturralde-Vinent & MacPhee 1996; Cepek in Schlee 1990). Both

estimates are derived from microfossils in the marine strata containing the amber, the brst authors deriving

theirs from foraminifera and the latter author utilizing coccoliths. The amber is secondarily deposited in

turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1964). According to

J. Bot. Res. Inst. Texas 9(2): 361 - 367.2015

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Journal of the Botanical Research Institute of Texas 9(2)

Fig. 1 . Comopellispresbya. Apical view of flower. A. petal. B. sepal. C. stamen. D. ovary. E. stigma. F. disc appendages. Scale bar = 1.0 mm.

Dilcher et al. (1992), “... the amber clasts, from all physical characteristics, were already matured amber at the

time of re-deposition into marine basins. Therefore the age of the amber is greater than Miocene and quite

likely is as early as late Eocene.” The discovery of Early Paleocene amber in Puerto Rico and Maastrichtian-

Paleocene amber in Jamaica (Iturralde-Vinent 2001) adds to the known range of such deposits in the Greater

Antilles.

Observations and photographs were made with a Nikon SMA-10 R stereoscopic microscope and Nikon

Optiphot compound microscope with magnifications up to 600X. Helicon Focus Pro X54 was used to stack

photos for better clarity and depth of held.

Chambers and Poinar, Comopellis presbya gen. et sp. nov.

363

Fig. 2. Comopellis presbya. Petal enclosing stamen. Arrow indicates anther. Scale bar = 0.2 mm.

DESCRIPTION

Comopellis K.L. Chambers & Poinar, gen. nov. (Figs. 1-3). Type Species: Comopellis presbya K.T. Chambers & Poinar sp. nov.

Flower bisexual, calyx regular, sepals 5, valvate, spreading, tip not thickened, midline not ridged (Fig. 1), co¬

rolla regular, petals 5, exceeded by sepals, margins involute, partially enclosing stamens, stamens appressed,

ca. 0.75 times as long as petals (Fig. 2), anthers ca. 0.5 times as long as blaments, bilocular, introrsely dehiscent,

pistil superior, basal in hypanthium, ovary globose, carpel number not determined, stigma broadly discoid,

not lobed, included in hypanthium, hypanthium hemispherical, densely hispid abaxially (Fig. 4), disc

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Journal of the Botanical Research Institute of Texas 9(2)

Fig. 3. Comopellispresbya. Apex of disc appendage. Arrow indicates glandular tissue. Scale bar = 0.1 mm.

glabrous, thin, not fleshy, lining hypanthium, the margin forming a rim at base of perianth, with 5 bilobed

appendages opposite sepals (Fig. 3), pedicel puberulent, ± equaling hypanthium (Fig. 4).

Etymology.—From Greek “kome,” hair, and “pellis,” cup or bowl, based on fossil’s hirsute hemispheric

hypanthium.

Comopellis presbya K.L. Chambers & Poinar, sp. nov. (Figs. 1-4). Type: HISPANIOLA. Dominican Republic: amber mine

in the northern mountain ranges (Cordillera Septentrional, between Puerto Plato and Santiago), 2012, unknown amber miner s.n.

(holotype: Catalog number Sd-9-194 , deposited in the Poinar amber collection maintained at Oregon State University, Corvallis,

Oregon 97331, U.S.A.).

Width in apical view 4.6 mm, sepals broadly deltoid-ovate, 1.5-1.7 mm long, 0.9-1.4 mm wide, glabrous, apex

acute, margins entire (Fig. 1), petals narrowly elliptic-lanceolate, 0.9-1.6 mm long, 0.2-0.4 mm wide, glabrous

or lightly floccose apically, stamens 0.8-1.0 mm long, filaments ca. 0.7 mm long, anthers 0.2-0.3 mm long (Fig.

2), ovary 1.6 mm in diameter, style length not determined, stigma 0.7 mm wide (Fig. 1), hypanthium 1.7 mm

wide, 0.8 mm high (Figs. 1,4), appendages of disc ca. 0.8 mm long, glandular apically and possibly nectarifer¬

ous (Fig. 3), pedicel 1.4 mm long.

Etymology .—From Greek ’’presbya,” old, elder.

DISCUSSION

Rhamnaceae are a family of ca. 57 genera and 950 species (Mabberley 2008). The tribal classification was

revised by Richardson et al. (2000a) on the basis of morphological traits, and a phylogenetic analysis was pre¬

sented (Richardson et al. 2000b) derived from plastid DNA sequences. Eleven tribes are recognized by these

Chambers and Poinar, Comopellis presbya gen. et sp. nov.

365

Fig. 4. Comopellis presbya . Lateral view of flower. A. hypanthium. B. pedicel. Scale bar = 0.8 mm.

authors. A key to tribes and genera is given in Medan & Schirarend (2004), together with abbreviated tribal

diagnoses. Vegetative and fruit characteristics, absent in the fossil, are important in the tribal classification;

hence, we have not tried to assign Comopellis to a particular suprageneric category. Distinctive features that

might help in tribal placement of the fossil are the lack of a keeled midrib on the sepals, lanceolate petals later¬

ally enfolding the appressed stamens, the hemispherical hypanthium, the bilobed, gland-tipped appendages of

the hypanthial disc, and the discoid, unlobed stigma. The disc appendages of Distigouania are similarly gland-

tipped, but its flowers are unisexual and irregular, with anthers free from the petals, disc Tiling the hypanthi¬

um, and petals ovate-deltoid and sepaloid in texture (Chambers & Poinar 2014a). The related genus Gouania

has a fleshy disc and inferior ovary. Flowers of Nahinda, an Oligocene fossil from Puebla, Mexico (Calvillo-

Canadell & Cevallos-Ferriz 2007), differ from Comopellis in lacking disc appendages and possessing a thick¬

ened midline rib and apex on the sepals. Based on a comparison of its features with published illustrations and

descriptions (Suessenguth 1953; Liogier 1981; Medan & Schirarend 2004), with a focus on extant genera of

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Journal of the Botanical Research Institute of Texas 9(2)

Rhamnaceae from the Caribbean and the New World in general, we have not been able to bnd a placement for

the fossil, which is here described as new to science.

ACKNOWLEDGMENTS

We thank Melanie DeVore, Melissa Islam, and Steven Manchester for their helpful review comments.

REFERENCES

Calvillo-Canadell, L. & S.R.S. Cevallos-Ferriz. 2007. Reproductive structures of Rhamnaceae from the Cerro del Pueblo

(Late Cretaceous, Coahuila) and Coatzingo (Oligocene, Puebla) Formations, Mexico. Amer. J. Bot. 94:1658-1669.

Chambers, K.L. & 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. 2012. A Mid-Tertiary fossil flower of Swietenia (Meliaceae) in Dominican amber. J. Bot. Res.

Inst. Texas 6:123-127.

Chambers, K.L. & G.O. Poinar, Jr. 2013. A fossil flower of the genus Protium (Burseraceae) in Mid-Tertiary amber from the

Dominican Republic. J. Bot. Res. Inst.Texas 7:367-373.

Chambers, K. L. & G.O. Poinar, Jr. 2014a. Distigouania irregularis (Rhamnaceae) gen. et sp. nov. in Mid-Tertiary amber from

the Dominican Republic. J. Bot. Res. Inst.Texas 8:555-561.

Chambers, K.L. & G.O. Poinar, Jr. 2014b. Ticodendron palaios sp. nov. (Ticodendraceae), a Mid-Tertiary fossil flower in

Dominican amber. J. Bot. Res. Inst.Texas 8:563-568.

Chambers, K.L., G.O. Poinar, Jr., & A.E. Brown. 2011a. 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., & A.E. Brown. 2011 b. Two fossil flowers of Trichilia (Meliaceae) in Dominican amber. J. Bot.

Res. Inst.Texas 5:463-468.

Chambers, K.L., G.O. Poinar, Jr., & A.S. Chanderbali. 2012. Treptostemon (Lauraceae), a new genus of fossil flower from Mid-

Tertiary Dominican amber. J. Bot. Res. Inst.Texas 6:551-556.

Dilcher, D.L., P.S. Herendeen, & F. Huber. 1992. Fossil Acacia flowers with attached anther glands from Dominican amber. In:

P.S. Herendeen & D.L. Dilcher, eds. Advances in legume systematics. Part 4. The fossil record. Royal Botanic Gardens,

Kew, U.K.

Draper, G., P. Mann, &J.F. Lewis. 1994. Hispaniola. In: S. Donovan &T.A. Jackson, eds. Caribbean geology: An introduction.

The University of the West Indies Publisher's Association, Kingston, Jamaica. Pp. 129-150.

Iturralde-Vinent, M.A. 2001. Geology of the amber-bearing deposits of the Greater Antilles. Caribbean J. Sci. 37:141-167.

Iturralde-Vinent, M.A. & R.D.E. MacPhee. 1966. Age and paleographic origin of Dominican amber. Science 273:1850-1852.

Liogier, A.H. 1961. Rhamnaceae. In: Flora of Hispaniola. Part 1. Phytologia Memoirs, Plainfield, New Jersey, U.S.A. Pp.

33-49.

Mabberley, DJ. 2008. Mabberley's plant book. Cambridge University Press, Cambridge, England. P. 733.

Medan, D. &C. Schirarend. 2004. Rhamnaceae. In: K. Kubitzki, ed. The families and genera of vascular plants. VI. Flowering

plants-Dicotyledons. Springer-Verlag, Berlin, Germany. Pp. 320-338.

Poinar, G.O., Jr. 1991. Hymenaea protera sp. n. (Leguminosae: Caesalpinioideae) from Dominican amber has African

affinities. Experientia 47:1075-1082.

Poinar, G.O., Jr. 2002a. Fossil palm flowers in Dominican and Mexican amber. Bot. J. Linn. Soc. 138:57-61.

Poinar, G.O., Jr. 2002b. Fossil palm flowers in Dominican and Baltic amber. Bot. J. Linn. Soc. 139:361 -367.

Poinar, G.O. Jr. & R. Poinar. 1999. The amber forest. Princeton University Press, Princeton, New Jersey, U.S.A.

Poinar, G.O., Jr. & EJ. Judziewicz. 2005. Pharusprimuncinatus (Poaceae: Pharoideae: Phareae) from Dominican amber. Sida

21:2095-2103.

Poinar, G.O. Jr., K.L. Chambers, & A.E. Brown. 2008a. Lasiambixdominicensis gen. et sp. nov., a eudicot flower in Dominican

amber showing affinities with Fabaceae subfamily Caesalpinioideae. J. Bot. Res. Inst. Texas 2:463-471.

Poinar, G.O., Jr., K.L. Chambers, & A.E. Brown. 2008b. Trochanthera lepidota gen. et sp. nov., a fossil angiosperm inflores¬

cence in Dominican amber. J. Bot. Res. Inst.Texas 2:1167-1173.

Poinar, G.O., Jr. & J.T. Columbus. 2012. Alarista succina gen. et sp. nov. (Poaceae: Bambusoideae) in Dominican amber.

Histor. Biol. 1-6.

Poinar, G.O., Jr. & R. Steeves. 2013. Virola dominicana sp. nov. (Myristicaceae) from Dominican amber. Botany 91:530-534.

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367

Poinar, G.O., Jr. & K.L. Chambers. 2015a. Prioria dominicana sp. nov. (Fabaceae: Caesalpinioideae), a fossil flower in Mid-

Tertiary Dominican amber. J. Bot. Res. Inst.Texas 9(1 ):129-134.

Poinar, G.O., Jr. & K.L. Chambers. 2015b. Dasylarynxanomalus gen. et sp. nov., a tubular monocotyledon-like flower in Mid-

Tertiary Dominican amber. J. Bot. Res. Inst.Texas 9(1 ):121—128.

Poinar, G.O., Jr. & K.L. Chambers. 2015c. Pseudhaplocricus hexandrus gen. et sp. nov. (Commelinaceae) in Mid-Tertiary

Dominican amber. J. Bot. Res. Inst.Texas 9(2):353—359.

Richardson, J.E., M.F. Fay, Q.C.B. Cronk, & M.W. Chase. 2000a. A revision of the tribal classification of Rhamnaceae. Kew Bull.

55:311-340.

Richardson, J.E., M.F. Fay, Q.C.B. Cronk, D. Bowman, & M.W. Chase. 2000b. A phylogenetic analysis of Rhamnaceae using rbcL

and trn-F plastid DNA sequences. Amer. J. Bot. 87:1309-1324.

Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C, 28.

Suessenguth, K. 1953. Rhamnaceae. In: H. Flarms, ed. Die Naturlichen Pflanzenfamilien. 20d. Angiospermae: Reihe

Rhamnales. Duncker & Flumblot, Berlin, Germany. Pp. 7-173.

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BOOK NOTICE

T. Scott Bryan & Betty Tucker-Bryan. 2015. The Explorer’s Guide to Death Valley National Park, 3rd ed.

(ISBN-13: 978-1-60732-340-2, pbk). University Press of Colorado, 5589 Arapahoe Avenue, Suite 206C,

Boulder, Colorado 80303, U.S.A. (Orders: www.upcolorado.com, 1-800-621-2736). $23.95,472 pp., 152

illus., 6" x 9".

From the Publisher: Originally published in 1995, soon after Death Valley National Park became the fifty-third

park in the US park system, The Explorer’s Guide to Death Valley National Park was the first complete guidebook

available for this spectacular area.

Now in its third edition, this is still the only book that includes all aspects of the park. Much more than

just a guidebook, it covers the park’s cultural history, botany and zoology, hiking and biking opportunities,

and more. Information is provided for all of Death Valley’s visitors, from first-time travelers just learning about

the area to those who are returning for in-depth explorations.

The book includes updated point-to-point logs for every road within and around the park, as well as more

accurate maps than those in any other publication. With extensive input from National Park Service resource

management, law enforcement, and interpretive personnel, as well as a thorough bibliography for suggested

reading, The Explorer’s Guide to Death Valley National Park, Third Edition is the most up-to-date, accurate, and

comprehensive guide available for this national treasure.

T. Scott Bryan was a seasonal employee at Yellowstone National Park from 1970 through 1986. In addition to

his studies in Yellowstone, he has been to geyser fields throughout the contiguous United States, Mexico,

Japan, Fiji, New Zealand, and the Valley of Geysers on the Kamchatka Peninsula of Russia, leading the first-

ever US study group there in 1991.

Betty Tucker-Bryan is the founder of the Death Valley Hikers Association and has written numerous books

and articles on the outdoors.

J.Bot. Res. Inst. Texas 9(2): 368.2015

KLAPROTHIOPSIS DYSCRITA GEN. ET SP. NOV. (LOASACEAE)

IN MID-TERTIARY DOMINICAN AMBER

George O. Poinar, Jr.

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

poinarg@science.oregonstate.edu

Maximilian Weigend

Nees-lnstitut fur Biodiversitat der Pflanzen

Rheinische-Friedrich-Wilhelms Universitat Bonn

MeckenheimerAllee, 17053115 Bonn, GERMANY

Tilo Henning

Botanischer Garten und Botanisches Museum

Freie Universitat Berlin

Konigin-Luise Str. 6-8, 14195 Berlin, GERMANY

ABSTRACT

Klaprothiopsis dyscrita gen. et sp. nov. is described from two flowers in Dominican Republic amber. The actinomorphic, tetramerous, bi¬

sexual flowers are synsepalous with a short calyx tube and four imbricate persistent lobes rounded at their apices. The four delicate, mem¬

branous petals are inserted at the throat of the calyx. A pair of curved ciliolate staminodes is situated opposite each petal. In addition, the

androecium consists of eight stamens with long, filiform filaments bearing introse anthers with lateral dehiscence. The inferior ovary is

crowned by a minute filiform style with a slightly thickened stigma. The fossils show strong affinities with the tribe Klaprothieae based on

similarity in flower structure but are only tentatively assigned to that group due to some conflicting morphological characters, especially the

antepetalous staminode insertion (vs. antesepalous in extant representatives). These flowers add to the floral diversity of plant communities

that existed in the West Indies in the mid-Tertiary.

Key Words: Klaprothiopsis dyscrita gen. et sp. nov. Toasaceae, Klaprothieae, Dominican amber, paleobotany

RESUMEN

Klaprothiopsis dyscrita gen. et sp. nov. es descrita en base a dos flores preservadas en ambar provenientes de la Republica Dominicana. Las

flores son actinomorfas, tetrameras, bisexuales, sinsepalas con el caliz formando un tubo corto y cuatro lobulos imbricados, persistentes

con apices redondeados. Los cuatro petalos delicados y membranaceos se encuentran insertados en el apice del tubo calicino. Los dos esta-

minodios curvados, ciliolados estan opuestos a los petalos. El androceo consiste en ocho estambres con filamentos largos, filiformes, con

anteras introrsas de dehiscencia lateral. El ovario infero termina en un estilo pequerhsimo con un estigma un poco ensanchado. Los fosiles

son muy parecidos a los de la tribu Klaprothieae en su estructura floral, pero muestran algunas caracteristicas algo aberrantes para el grupo,

especialmente la insercion antepetala de los estaminodios (en contraste con una insercion antesepala en los representantes actuales). Estas

flores representan una importante contribucion a nuestro conocimiento de la diversidad floral de las comunidades vegetales que existian en

las islas de las Indias Occidentales durante la Era del Terciario Medio.

INTRODUCTION

Fossil flowers in amber from the Dominican Republic have provided rare records of Neotropical plant families

and genera that existed in Hispaniola during the mid-Tertiary. Thus far representatives of the families

Fabaceae, Arecaceae, Poaceae, Chrysobalanaceae, Lauraceae, Meliaceae, Burseraceae, Myristicaceae,

Rhamnaceae and Ticodendraceae have been described from Dominican amber (references listed in Poinar &

Chambers 2014). The present study reports the first putative fossil representatives of the family Loasaceae

(Cornales). The Loasaceae is a relatively small family comprising some 350 species in 20 genera of tropical and

subtropical herbs, shrubs, trees and lianas, mainly occurring in the Americas (Weigend 2004a, b) with the

exception of two genera.

MATERIALS AND METHODS

The fossils originated from amber mines in the northern mountain range (Cordillera Septentrional) of the

Dominican Republic between Puerto Plata and Santiago. Amber from mines in this region was produced by

Hymenaeaprotera Poinar (1991) (Fabaceae).

J. Bot. Res. Inst. Texas 9(2): 369 - 379.2015

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Journal of the Botanical Research Institute of Texas 9(2)

The Dominican amber forest was characterized by Poinar and Poinar (1999) based on both animal and

plant fossils.

Dating of Dominican amber is controversial, with the youngest proposed age of 20-15 mya based on

Foraminifera (Iturralde-Vinent & MacPhee 1996) and the oldest of 45-30 mya based on coccoliths (Cepek in

Schlee 1990). These are considered minimum dates as they are based on microfossils in the strata containing

the amber. Most of the amber was secondarily deposited in turbiditic sandstones of the Upper Eocene to Lower

Miocene Mamey Group (Draper et al. 1994). Dilcher et al. (1992) stated that “... the amber clasts, from all

physical characteristics, were already matured amber at the time of re-deposition into marine basins. Therefore,

the age of the amber is greater than Miocene and quite likely is as early as late Eocene”. The issue is further

complicated by the discovery of Early Oligocene amber in Puerto Rico and Maastrichtian-Paleocene amber in

Jamaica (Iturralde-Vinent 2001) showing that amber from a range of deposits occurs in the Greater Antilles.

Observations and photographs were made with a Nikon SMZ-10 R stereoscopic microscope and Nikon

Optiphot compound microscope with magnifications up to 600 X. Helicon Focus Pro X64 was used to stack

photos for better clarity and depth of held.

The two fossil bowers are in separate pieces of Dominican amber. The specimens differ slightly in size and

tint but share the same morphological features and are considered to belong to the same species.

DESCRIPTION

Loasaceaejusseau 1804

Klaprothiopsis Poinar, Weigend, & Henning, gen. nov. (Figs. 1-7). Type Species: Klaprothiopsis dyscrita Poinar, Weigend,

& Henning, sp. nov.

Klaprothiopsis dyscrita Poinar, Weigend, & Henning, sp. nov. (Figs. 1-7). Type: HISPANIOTA. Dominican Republic:

amber mine in the northern mountain ranges (Cordillera Septentrional), 1986, unknown amber miner s.n. (holotype: accession #

Sd-9-108A and paratype (accession # Sd-9-108B) deposited in the Poinar amber collection maintained at Oregon State University,

Corvallis, Oregon 97331, U.S.A.).

Holotype Sd-9-108A (Figs. 1-5): Actinomorphic, tetramerous, bisexual Power, pedicellate; pedicel 2.2 mm long,

set with simple trichomes; ovary inferior; calyx tube wide, calyx lobes persistent, ovate, 1.1 mm long, apex

obtuse, 0.7 mm wide; petals half spreading, narrowly ovate, delicate, membranous, 2.6-2.8 mm long, 1.1 mm

wide, set with simple trichomes at base and margin, base slightly narrowed, apex acuminate, entire; stamino-

dia antepetalous, 2 per petal, curved, ciliolate, 0.6-0.8 mm long, densely set with simple trichomes; stamens 8,

3.0-3.5 mm long, filaments filiform, anthers introse, 0.4-0.6 mm long, 0.2-0.3 mm wide, with lateral dehis¬

cence; style filiform, 0.2 mm long, stigma slightly thickened.

Paratype Sd-9-108B (Figs. 6,7): Actinomorphic, tetramerous, bisexual Power, pedicellate; pedicel 3.0 mm long,

set with simple trichomes; ovary inferior; calyx tube wide, calyx lobes persistent, ovate, 1.2 mm long, apex

obtuse, 0.7 mm wide; petals half spreading, narrowly ovate, delicate, membranous, 2.5 mm long, 1.1 mm wide,

set with simple trichomes at base and margin, base slightly narrowed, apex acuminate, entire; staminodia an¬

tepetalous, 2 per petal, curved, ciliolate, 0.6-0.8 mm long, densely set with simple trichomes; stamens 8, 2.9

mm long, Plaments Pliform, anthers introse, 0.5 mm long, 0.2 mm wide, with lateral dehiscence; style Pliform,

0.2 mm long, stigma slightly thickened.

Etymology. —The generic name rePects the similarity of the fossil to the genus Klaprothia (see Fig. 8). The spe-

cihc epithet is from the Greek “ dyskrita ” = difhcult to determine.

DISCUSSION

Klaprothiopsis dyscrita gen. et sp. nov. is provisionally assigned to the Loasaceae and shows strong similarities

to the tribe Klaprothieae based on its Power morphology. A range of characters, such as Power size, tetramer¬

ous perianth, shape and persistence of the sepals, membranous petal structure and the peculiar free, club-

shaped, papillose staminodes support this placement. This group also has protandrous Powers with a style

that elongates only late in anthesis, indicating that the two fossil Powers may be early anthetic, therefore

Poinar et al., Klaprothiopsis dyscrita gen. et sp. nov. from Mid-Tertiary Dominican amber

371

Fig. 1. Lateral view of side A of Klaprothiopsis dyscrita gen. et sp. nov. Holotype Sd-9-108A in Dominican amber. Scale bar = 0.8 mm.

having only a very short style. Thus, detailed floral morphology and geographic distribution support Klaprothia

spp. as closest relatives of Klaprothiopsis dyscrita gen. et sp. nov. (Table 1).

Table 1 . Comparison of Klaprothiopsis dyscrita gen. et sp. nov. with the two extant species in the genus Klaprothia. All measurements are in millimeters.

Klaprothia mentzelioides

Klaprothia fasciculata

Klaprothiopsis dyscrita gen. et sp. nov.

Length petioles

5-60

10-38

2.2-3

Length sepals

1.0-2.2

0.75-2

1.1-1.2

Length petals

2.0-5.7

1.0-1.75

2.S-2.8

No. of stamens

16-20

4-12

Length of stamens

2.5-4.0

0.75-1.0

2.9-3.5

No. of staminodes

5, three outer and 2 free inner

2*-3, free

2, free

Length of staminodes

1.5-3.0

0.5-0.75

0.6-0.8

Position of staminodes

antesepalous

antepetalous

Style length

2.0-3.0

0.5-1.25

0.2

Shape of ovary

obovoid to oblong, short

cylindrical to obovoid, elongated

minute

* Poston & Nowicke (1990): 2-3 written in the key, 3 in the diagnoses

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 2. Lateral view of side B ofKIaprothiopsisdyscritaqen. et sp. nov. Holotype Sd-9-108A with pedicel and calyx in Dominican amber. Scale bar=1.4 mm.

Poinar et al., Klaprothiopsis dyscrita gen. et sp. nov. from Mid-Tertiary Dominican amber

373

Fig. 3. Top view of center of Klaprothiopsis dyscrita gen. et sp. nov. Holotype Sd-9-108A in Dominican amber. Scale bar = 0.5 mm. Note short pistil sur¬

rounded by stamen bases.

374

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 4. Inside view of Klaprothiopsis dyscrita gen. et sp. nov. Holotype Sd-9-108A showing paired staminodes (S) in front of a petal (P) in Dominican

amber. Scale bar = 0.5 mm.

However, there are some striking morphological differences between the fossil and extant Klaprothia —

and indeed all Loasoideae. Most notable is the antepetalous insertion of the staminodes since both staminodes

and staminodial complexes are universally found in an antesepalous position in extant Loasoideae. Another

character is the receptacle and the ovary. Throughout Loasaceae and indeed all Cornales, the inferior ovary

develops proleptically, i.e., is already largely developed before anthesis begins. Thus, the inferior and the cor¬

responding smaller superior part of the ovary are usually well visible at beginning of the anthesis. The fossil

has a very small ovary and a minuscule style that could be interpreted as a rudimentary female organ in an

otherwise male flower. This would indicate a monoecious or dioecious species, which would argue against

Loasaceae with exclusively hermaphroditic flowers. Finally, although members of the Loasaceae literally ex¬

hibit almost all trichome types known in the plant kingdom, smooth trichomes as found in the fossil flowers

are extremely rare in extant species and not known from Klaprothieae (Poston & Nowicke 1990,1993; Dostert

& Weigend 1999; Weigend 1997a, 2004; Noguera-Savelli et al. 2009). These similarities and differences are

difficult to evaluate, especially in the light of the age of Dominican amber.

In conclusion, the present placement of Klaprothiopsis dyscrita gen. et sp. nov. cannot be confirmed with

absolute certainty due to the above mentioned conflicting characters that question our preliminary assign¬

ment. Overall, the characters observed either support or question placement in Loasaceae, but none of them,

whether observed (smooth trichomes) or assumed (e.g. monoecy vs. proterandry) strongly indicate an alterna-

Poinar et al., Klaprothiopsis dyscrita gen. et sp. nov. from Mid-Tertiary Dominican amber

375

Fig. 5. Lateral view of paired staminodes (arrows) of Klaprothiopsis dyscrita gen. et sp. nov. Holotype Sd-9-1 08A in Dominican amber. Scale bar = 0.3 mm.

tive assignment to any other extant plant family we know. Thus for the present, we treat the fossil as allied to

the Klaprothieae, but refer to it as Klaprothiopsis dyscrita gen. et sp. nov., which reflects our uncertainty of its

placement.

Klaprothieae have repeatedly been retrieved as monophyletic by both, morphological (Weigend 1997a, b,

2004) and molecular data (Hufford et al. 2003, 2005; Weigend et al. 2004) and represent one of the basal clades

of Loasaceae subfam. Loasoideae. In so-called “Higher Loasoideae” the staminodes are united into staminodial

complexes, some of them fused into floral scales. Klaprothieae comprise three small genera ( Klaprothia

Kunth.—2 spp, Plakothira Florence (1997)—3 spp and Xylopodia klaprothioides Weigend—monotypic), with a

heterogenous distribution. The two species of Klaprothia (K. mentzelioides Kunth. and K. fasciculata (C. Presl)

Poston, Poston & Nowicke 1990) are widespread in Central and Western South America (up to Mexico),

Brazil, Caribbean Islands and Galapagos Islands, whereas Plakothira spp. are restricted to the Marquesas

Islands (Polynesia) and Xylopodia is microendemic from the area around Contumaza in northern Peru

(Weigend 1997a, Weigend et al. 2006). In flower size and details of the staminodes, both Xylopodia and

Plakothira differ clearly from the fossil flowers, which are essentially intermediate between the two extant spe-

376

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 6. Dehisced anther of Klaprothiopsis dyscrita gen. et sp. nov. Paratype (Sd-9-108B) in Dominican amber. Scale bar = 0.2 mm.

cies of Klaprothia. Apart from the substantial differences discussed above, Klaprothiopsis dyscrita gen. et sp.

nov. most closely resembles K. fasciculata in terms of the remaining flower morphological characters. This is

also the most plausible placement based on the fossil’s distribution: K. fasciculata is represented in the

Caribbean by two specimens collected in Haiti at low and intermediate elevations (150-1600 m), whereas K.

mentzelioides is montane (1000-3500 m, Poston & Nowicke 1990). Dated phylogenies provide an estimated

age of 44-94 Ma (Xiang et al. 2011) and ca. 60-92 respectively (Schenk & Hufford 2010) for the crown group

of Loasaceae. Schenk & Hufford (2010) also provide an estimated age for subfam. Loasoideae of c. 65 Ma with

the Klaprothieae splitting from the stem lineage between 25-55 Ma ago. Klaprothia fasciculata is tentatively

dated at only ca. 5-10 Ma in their analyses. The age of Klaprothiopsis dyscrita, e.g., ranging between 20-15

(Iturralde-Vincent & MacPhee 1996) and 45-30 (Cepek in Schlee 1990) Ma, is incompatible with the poten¬

tially young age of its possible closest extant relative K. fasciculata (based on this rather broad study on order-

level). Nevertheless, the characters detailed above, including the combination of features found in both extant

species of Klaprothia (Table 1) and the potentially great age of lineages in the Klaprothieae (Schenk & Hufford

2010) justify the assignment of the fossils in the proximity of the Klaprothieae.

An analysis of plant and animal inclusions in Dominican amber showed that the amber forest was a tropi¬

cal moist forest (Poinar & Poinar 1999). Such forests today, which occur throughout the Neotropics, com¬

monly have a dry season of 3 or 4 months, followed by a wet period of 8 to 9 months. The amber forest probably

had several layers of vegetation including canopy, subcanopy, understory, shrub and forest floor strata, which

is typical of moist forests today. Vines and epiphytes would have extended from tree to tree. Such forests no

longer exist in Hispaniola.

Poinar et al., Klaprothiopsis dyscrita gen. et sp. nov. from Mid-Tertiary Dominican amber

377

Fig. 7. Smooth trichomes on surface of petal of Klaprothiopsis dyscrita gen. et sp. nov. Paratype (Sd-9-108B) in Dominican amber. Scale bar = 0.15 mm.

Fig. 8. SEM-photo of top view of the extant Klaprothiafasciculata. Voucher: Peru, Dept. Amazonas, Prov. Bagua. La Peca, trail to El Paraiso, 1000-1300

m, adjacent to banana plantations and along riverbanks. 30.03.1998. Nicolas Dostert 98/76 (M).

378

Journal of the Botanical Research Institute of Texas 9(2)

Klaprothiopsis dyscrita gen. et sp. nov. probably had a herbaceous habit, similar to that of the two extant

species of Klaprothia. Their presence in amber suggests that Klaprothiopsis dyscrita gen. et sp. nov. either grew

closely adjacent to or climbed the trunks of the resin producing trees that produced the amber. These amber

flowers add to the floral diversity that was present in the West Indies in the mid-Tertiary.

ACKNOWLEDGMENTS

Thanks are extended to Hans-Jurgen Ensikat (Bonn) for preparing the SEM-picture of extant K.fasciculata and

to two reviewers whose remarks enhanced the paper.

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Botanical Gardens, Kew, England. Pp. 33-42.

Dostert, N. & M. Weigend. 1999. A synopsis of the Nasa triphylla complex (Loasaceae) including some new species and

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Florence, J. 1997. New species of Plakothira (Loasaceae), Melicope (Rutaceae), and Apetahia (Campanulaceae) from the

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Hufford, L., M.M. Mcmahon, A.M. Sherwood, G. Reeves, & M.W. Chase. 2003. The major clades of Loasaceae: phylogenetic

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Iturralde-Vinent, M.A. & R.D.E. MacPhee. 1996. Age and paleographic origin of Dominican amber. Science 273:1850-1852.

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Noguera-Savelli, E., D. Jauregui, & T. Ruiz-Zapata. 2009. Indument characterization of nine Loasaceae species from

Venezuela. Revista Mex. Biodivers. 80:751-762.

Poinar, G.O., Jr. 1991. Hymenaea protera sp. n. (Leguminosae: Caesalpinioideae) from Dominican amber has African af¬

finities. Experientia 47:1075-1082.

Poinar, G.O., Jr. & R. Poinar. 1999. The amber forest. Princeton University Press, Princeton, New Jersey, U.S.A.

Poinar, G.O., Jr. & K.L. Chambers. 2015. Prioria dominicana sp. nov. (Fabaceae: Caesalpinioideae), a fossil flower in mid-

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Poston, M.E. & J.W. Nowicke. 1990. A reevaluation of Klaprothia and Sclerothrix (Loasaceae: Klaprothieae). Syst. Bot.

15:671-678.

Poston, M.E. & J.W. Nowicke. 1993. Pollen morphology, trichome types, and relationships of the Gronovioideae

(Loasaceae). Amer. J. Bot. 80:689-704.

Schenk, JJ. & L. Hufford. 2010. Effects of substitution models on divergence time estimates: simulations and an empirical

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BOOK NOTICE

Ghillean T. Prance. 2014. That Glorious Forest: Exploring the Plants and Their Indigenous Uses in

Amazonia. (ISBN-13: 978-0-893-27528-0, hbk). New York Botanical Garden Press, Library Bldg, Room

M221, 2900 Southern Boulevard, Bronx, New York 10458-5126, U.S.A. (Orders: www.nybgpress.org,

1-718-817-8721). $69.00, 224 pp., 8.5" x 10".

From the Publisher: After nearly bfty years since his brst journey to the tropics exploring for plants, and a total

of thirty-nine expeditions to the Amazon, Professor Sir Ghillean Prance has gathered in this volume a fascinat¬

ing and diverse collection of accounts from these experiences, along with his thoughts on a lifetime of work

surveying the Amazon flora. A prominent botanist and ecologist, as well as the former Senior Vice President of

Science at The New York Botanical Garden, and subsequently Director of the Royal Botanic Gardens, Kew

(1988-1999), Sir Ghillean has written this book as a tribute to the many people who have helped him to survey

the Amazon region over the last five decades. The main focus in this volume is placed on the process and the

running of the expeditions, as well as collecting specimens. Included at the end of each chapter are plant col¬

lection numbers made on the expeditions described, as well as a list of new species discovered. Professor

Prance is a specialist in a number of plant families, all of which consist of trees and vines widely distributed in

the Amazon region and elsewhere. In recognition of more than a quarter century of his service and ongoing

extraordinary dedication to the mission of The New York Botanical Garden, and to botany and horticulture

internationally, Sir Ghillean was presented in 2008 with the Gold Medal of The New York Botanical Garden,

where he served as a scientist and senior scientific administrator. This medal is the highest honor that the

Botanical Garden confers. In the botanical exploration of Amazonian Brazil, there is probably no contempo¬

rary name more prominent than that of Sir Ghillean Prance.

J.Bot. Res. Inst. Texas 9(2): 380.2015

ADDENDUM: PRIORIA DOMINICANA SP. NOV.

(FABACEAE: CAESALPINIOIDEAE),

A FOSSIL FLOWER IN MID-TERTIARY DOMINICAN AMBER

George O. Poinar, Jr.

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

poinarg@science.oregonstate.edu

In the original publication, the dimensions of the scale

correct values are as follows:

Fig. 1. Scale bar =1.1 mm

Fig. 2. Scale bar = 0.9 mm

Fig. 3. Scale bar = 0.6 mm

Kenton L. Chambers

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

in the 3 bgures were inadvertently omitted. The

J.Bot. Res. Inst. Texas 9(2): 381.2015

382

Journal of the Botanical Research Institute of Texas 9(2)

JOURNAL NOTICE

Neal K. Van Alfen, Jan E. Leach, and Steven Lindlow, eds. 2015 (Aug). Annual Review of Phytopathology,

Volume 53. (ISSN: 0066-4286; ISBN: 978-0-8243-1353-1, hbk). Annual Reviews, Inc., 4139 El Camino

Way, RO. Box 10139, Palo Alto, California 94303, U.S.A. (Orders: www.AnnualReviews.org, science@

annualreviews.org, 1-800-523-8635,1-650-493-4400). $99.00 indiv., 634 pp., 7.5” x 9.25”.

ABOUT THIS JOURNAL—The Annual Review of Phytopathology, in publication since 1963, covers the significant developments in the field

of plant pathology, including plant disease diagnosis, pathogens, host-pathogen Interactions, epidemiology and ecology, breeding for resis¬

tance and plant disease management, and includes a special section on the development of concepts.

Contents of Volume 53:

1. The Wayward Hawaiian Boy Returns Home—D. Gonsalves

2. Playing on a Pathogen’s Weakness: Using Evolution to Guide Sustainable Plant Disease Control Strategies— J. Zhan, PH. Thrall, J.

Papaix, L. Xie, &J.J. Burdon

3. Dissecting the Molecular Network of Virus-Plant Interactions: The Complex Roles of Host Factors—A. Wang

4. Molecular Mechanisms of Nematode-Nematophagous Microbe Interactions: Basis for Biological Control of Plant-Parasitic

Nematodes—J. Li, C. Zou,J. Xu, X.Ji, X. Niu,J. Yang, X. Huang, & K.-Q. Zhang

5. Priming for Enhanced Defense— U. Conrath, G.J.M. Beckers, C.J.G. Langenbach, & M.R.Jaskiewicz

6. Genome-Enabled Analysis of Plant-Pathogen Migration— E.M. Goss

7. Citrus Tristeza Virus: Making an Ally from an Enemy—WO. Dawson, M. Bar-Joseph, S.M. Garnsey, & P. Moreno

8. Practical Benefits of Knowing the Enemy: Modern Molecular Tools for Diagnosing the Etiology of Bacterial Diseases and Understanding

the Taxonomy and Diversity of Plant-Pathogenic Bacteria—C.T. Bull & S.T. Koike

9. Genomics Spurs Rapid Advances in Our Understanding of the Biology of Vascular Wilt Pathogens in the Genus Verticillium —A. Klimes,

K.F. Dobinson, B.PH.J. Thomma, & S.J. Klosterman

10. Soil Health Paradigms and Implications for Disease Management— R.P. Larkin

11. Epidemiology and Population Biology of Pseudoperonospora cubensis: A Model System for Management of Downy Mildews— PS.

Ojiambo, D.H. Gent, L.M. Quesada-Ocampo, M.K. Hausbeck, & G.J. Holmes

12. Identifying and Naming Plant-Pathogenic Fungi: Past, Present, and Future—P.W. Crous, D.L. Hawksworth, & M.J. Wingfield

13. Impact of Diseases on Export and Smallholder Production of Banana— R.C. Ploetz, G.H.J. Kema, &L.-J. M a

14. Evolution of Plant Parasitism in the Phylum Nematoda—C.W Quist, G. Smant, &J. Helder

15. Lipochitooligosaccharides Modulate Plant Host Immunity to Enable Endosymbioses—E. Limpens, A. van Zeijl, & R. Geurts

16. Range-Expanding Pests and Pathogens in a Warming World— D.P. Bebber

17. Sharka Epidemiology and Worldwide Management Strategies: Learning Lessons to Optimize Disease Control in Perennial Plants—L.

Rimbaud, S. Dallot, T. Gottwald, V. Decroocq, E.Jacquot, S. Soubeyrand, & G. Thebaud

18. A Moving View: Subcellular Trafficking Processes in Pattern Recognition Receptor-Triggered Plant Immunity— S. Ben Khaled, J.

Postma, & S. Robatzek

19. Roots Shaping Their Microbiome: Global Hotspots for Microbial Activity— B. Reinhold-Hurek, W. Bunger, C. Sofia Burbano, M. Sabale, &

T. Hurek

20. Identification of Viruses and Viroids by Next-Generation Sequencing and Homology-Dependent and Homology-Independent

Algorithms—Q. Wit, S.-W. Ding, Y. Zhang, &S. Zhu

21. Quantitative Resistance to Biotrophic Filamentous Plant Pathogens: Concepts, Misconceptions, and Mechanisms— R.E. Niks, X. Qi, &

T.C. Marcel

22. Landscape-Scale Disease Risk Quantification and Prediction— J. Yuen &A. Mila

23. Torradoviruses— R.A.A. van der Vlugt, M. Verbeek, A.M. Dullemans, W.M. Wintermantel, W.J. Cuellar, A. Fox, &J.R. Thompson

24. Durable Resistance of Crops to Disease: A Darwinian Perspective— J.K.M. Brown

25. Understanding Plant Immunity as a Surveillance System to Detect Invasion— D.E. Cook, C.H. Mesarich, & B.PH.J. Thomma

26. Leaf Rust of Cultivated Barley: Pathology and Control—R.E. Park, PG. Golegaonkar, L. Derevnina, K.S. Sandhu, H. Karaoglu, H.M.

Elmansour, P.M. Dracatos, & D. Singh

27. Highways in the Sky: Scales of Atmospheric Transport of Plant Pathogens—D.G. Schmale III & S.D. Ross

28. Grapevine Leafroll Disease and Associated Viruses: A Unique Pathosystem—R.A. Naidu, H.J. Maree, &J.T. Burger

Errata

J.Bot. Res. Inst. Texas 9(2): 382.2015

BREEDING SYSTEM AND SEX RATIO VARIATION IN

MULBERRIES ( MORUS, MORACEAE)

Madhav P. Nepal

South Dakota State University

Department of Biology and Microbiology

Brookings, South Dakota 57007, U.S.A.

Madhav.Nepal@sdstate.edu

Carolyn J. Ferguson

Kansas State University

Herbarium and Division of Biology

Manhattan, Kansas 66506, U.S.A.

Mark H. Mayfield

Kansas State University

Herbarium and Division of Biology

Manhattan, Kansas 66506, U.S.A.

ABSTRACT

Flowering plants exhibit a diverse array of sex expression patterns that become of particular interest when considered in the context of co¬

existing native-invasive congeners. This study presents findings on sex expression and sex ratio variation in two congeneric tree species, the

native Morus rubra and the exotic M. alba, in the Flint Hills region of the Great Plains in the United States. Both species exhibited a subdioe-

cious breeding system (with male, female and hermaphrodite individuals co-occurring within populations), and significantly male-biased

sex ratios (i.e., males are more numerous than females). Cumulative sex ratio deviation was higher in the native M. rubra populations than

in the exotic M. alba. Within-species sex ratio did not vary significantly among populations. In one focus study area surveyed over three

years, most individuals were consistent in sex expression, but approximately 10% switched their sex at least once; the vast majority of these

changes were between unisexual and hermaphrodite morphs. There was no size dependence of sex expression based on current sampling.

Our results document subdioecy as the breeding system in these Morus species, and advance the group as an interesting study system for its

reproductive biology.

Key Words: Breeding system, Moraceae, Morus , subdioecy, sex ratio, sex switch

RESUMEN

Fas plantas con f lores exhiben un conjunto de patrones de expresion sexual que tiene particular interes cuando se considera en el contexto

de sus congeneres nativos-invasores con los que coexisten. Este estudio presenta hallazgos sobre la expresion sexual y variacion del ratio

sexual en dos especies congenericas arboreas, la nativa Morus rubra y la exotica M. alba, en la region de Flint Hills de la Gran Tlanura de los

Estados Unidos. Ambas especies exhiben un sistema de cruzamiento subdioico (con individuos masculinos, femeninos y hermafroditas en

las poblaciones), y ratios sexuales significantemente sesgados hacia los masculinos (ej. Eos masculinos son mas numerosos que los femeni¬

nos). Ta desviacion acumulativa de la ratio sexual era mas alta en las poblaciones de la nativa M. rubra que en la de la exotica M. alba. Ta ratio

sexual intraspecifica no varia significativamente entre poblaciones. En un area de estudio estudiada durante tres anos, la mayoria de los

individuos eran consistentes eh la expresion sexual, pero aproximadamente el 10% cambiaban su sexo al menos una vez; la gran mayoria de

estos cambios fueron entre morfos unisexuales y hermafroditas. No hubo dependencia del tamano en la expresion sexual en el muestreo

actual Nuestros resultados documentan subdioecia como Sistema de cruzamiento en estas especies de Morus, y avanzan al grupo como un

sistema de estudio interesante por su biologia reproductiva.

INTRODUCTION

Although an intriguing array of flowering plant sexual systems (= breeding systems) occurs in a hermaphro-

ditism-dioecy continuum, the breeding systems of the majority of plants have not been extensively explored

(Charlesworth 2006). Dioecy, in which male and female plants are separate, occurs in about 6% of angiosperms

(Renner & Ricklefs 1995), and may have evolved from hermaphroditism to prevent inbreeding (Charlesworth

& Charlesworth 1978; Thomson & Barrett 1981). Subdioecy, in which three sexual morphs (male, female and

hermaphrodite) occur, is thought to be a transition in the hermaphroditism-dioecy continuum and has been

documented for several plant species (e.g., Case et al. 2008). A sex ratio of 1:1 (male/female) is expected in an

ideal population if the reproductive cost of being male is equal to that of being female (Fisher 1930). Plants have

evolved several strategies for reproductive assurance that might result in a sex ratio deviation: the extent and

J. Bot. Res. Inst. Texas 9(2): 383 - 395.2015

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Journal of the Botanical Research Institute of Texas 9(2)

pattern of such deviation vary from species to species, among flowering episodes and across populations along

environmental gradients (reviewed in Queenborough et al. 2007). Although several studies have documented

sex ratio variation between flowering episodes (e.g., Nicotra 1998; Morellato 2004), fewer studies have ob¬

served the same individuals for more than two flowering seasons (e.g., Yamashita & Abe 2002; Wheelwright &

Logan 2004), and still fewer studies have included more than one species (e.g., Thomas & LaFrankie 1993;

Queenborough et al. 2007). Sex expression studies over multiple flowering episodes can provide information

on flowering frequency, size dependence, and stability of sex expression (Yamashita & Abe 2002; Nanami et al.

2004).

Spatial distribution of individuals can play an important role in the successful survival and reproduction

of a species (House 1992; Stacy et al. 1996). For example, individuals of a wind-pollinated dioecious species

may experience pollen limitation in areas distant from other individuals (Ashman et al. 2004). Aggregation of

males and females may facilitate pollen transfer (Bawa & Opler 1975) if the habitat is homogeneous in terms of

resources. Many species, however, exhibit spatial segregation of the sexes, where individuals are partitioned

along a resource gradient with females predominantly occurring in the resource rich habitats (Bierzychudek &

Eckhart 1988). There are other factors related to spatial distribution that affect the reproduction of dioecious

species, such as distance between males and females (Mack 1997), flowering frequency (Bawa 1980), effective

population size (Nunney 1995), and pollinator abundance and behavior (Stacy et al. 1996). Some strategies

such as leaky dioecy and parthenocarpy have been suggested for reproductive assurance (Baker & Cox 1984;

Venkatasamy et al. 2007). Production of fleshy fruits (Bawa 1980), woody perennial habit, multi-seeded fruits,

and dispersal by birds are commonly associated with dioecy (Baker & Cox 1984). Size-dependent sex expres¬

sion occurs when male and female individuals exhibit differential reproductive costs. Females generally have

higher reproductive costs and are usually larger in size than males (Lloyd & Bawa 1984). In wind-pollinated

dioecious plants, some studies have reported a higher cost of reproduction in females (Smith 1981; Murakami

& Maki 1992) while other studies have shown a positive correlation between the male investment and plant

size facilitating the dispersal of pollen (Freeman et al. 1980; Solomon 1989). Sex expression patterns in plants

are interesting, as they play important roles in micro-evolutionary processes that involve reproductive success,

inbreeding depression, fixation of deleterious recessive mutations (reviewed in Charlesworth 2006) and plas¬

ticity of sex expression (Delph & Wolf 2005).

Here we present our results on breeding systems and sex ratio variation of two species of the genus Moms:

M. rubra L. and M. alba L. Morus rubra is distributed in eastern North America from the east coast to the east¬

ern margin of the Great Plains, and extends northward to southern Ontario, Canada, where it occurs in iso¬

lated patches of woodlands (Wunderlin 1997). Morus alba, a native to China, was introduced into North

America in the 1600s to establish a silkworm industry. After escaping cultivation, M. alba has successfully

naturalized (Gleason 1952). It often co-occurs with the native M. rubra in forests, and open riparian wood¬

lands. Additionally, M. alba commonly occurs in open areas; and it is considered ecologically invasive (Uva et

al. 1997; Weber 2003). Indeed, the species constitute an interesting study system with many morphological

similarities (and they sometimes hybridize; Burgess et al (2008); although they are not each other’s closest rela¬

tive; Nepal & Ferguson 2012). Relative to the native M. rubra, some studies have found M. alba to produce more

pollen and seeds (Burgess et al. 2008), and better quality of fruits for dispersal by birds (Stapanian 1982). The

invasion success of M. alba remains relatively unexplored, and may in part be attributable to complex repro¬

ductive behavior. Morus species have been treated as dioecious or monoecious in the taxonomic literature (e.g.,

Berg 2001), and their sexual systems have not been extensively explored. Both species produce unisexual flow¬

ers in catkins, and the catkins can be unisexual or bisexual. In the present study, we used census data collected

over a three-year period for the native-invasive pair of Morus in the Flint Hills region of Kansas to address the

following questions. What breeding systems do M. alba and in M. rubra exhibit? Do their sex ratios deviate

from 1:1? And do sexual expression and sex ratios remain constant from a given year to the next?

Nepal et al., Sex expression and sex ratio variation in mulberries (Morus)

385

MATERIALS AND METHODS

We conducted field studies on populations of M. alba and M. rubra (22 populations total) in the Flint Hills re¬

gion of north central Kansas. The Flint Hills region extends throughout an area of 29,600 km 2 in eastern

Kansas and northern Oklahoma, and is a distinct grassland ecoregion in the Great Plains. Although the region

is largely dominated by C 4 grasses, included riparian areas are dominated by woody plant species including

Celtis occidentalism Quercus macrocarpa, and Q. muehlenbergii (Great Plains Flora Association 1986). Morus alba

is common in these habitats and additionally extends onto open prairie, whereas M. rubra persists in relatively

natural riparian areas. Twenty two populations of mulberries from 13 sites near Manhattan, Kansas (nine sites

where populations of M. rubra and M. alba grew together, and four where only M. alba was found), were inves¬

tigated in the spring of 2005: Kings Creek and Shane Creek areas of Konza Prairie Biological Station (KPBS);

Pottawatomie State Fishing Lake No. 1; Pottawatomie State Fishing Lake No. 2; Timber Creek Park and

Farnum Creek Park areas of Milford Lake; Slough Creek Park area of Perry Lake; Frank Anneberg Park and

part of the Linear Park Trail areas in Manhattan, Kansas; southeast and west areas of Tuttle Creek Lake; Three

Mile Creek area of Fort Riley Military Reservation; as well as a privately owned riparian area along Kuenzli

Creek southeast of Alma, Kansas (sect. 16). Representative voucher specimens were labeled and deposited at

the Kansas State University Herbarium (KSC). Although the two species of Morus can hybridize, all of the in¬

dividuals included in this study were readily identified as to species (see Nepal et al 2012).

For each tree, ten to twelve flowering branches growing at varying heights were assessed for sex expres¬

sion. In addition, nearest neighbor distance (NND) and diameter at breast height (DBH) were measured. The

population along Kings Creek was chosen for study of inter-year variation in sex expression; each tree was

tagged with a tree identification number and studied during three flowering episodes (2005, 2006, and 2007).

Sex ratio was expressed as a proportion of males to the sum of males and females. Deviation of sex ratios

from the unity was tested using % 2 statistics. Inter-year variation in sex ratio was analyzed with proportion of

males as the response variable using GENMOD procedure in Statistical Analysis System 9.0 (SAS). To test the

statistical difference between the proportion of males in two species, generalized linear models were used with

binomial distribution and the logit link function (Crawley 1993). Variation of sex ratio within each species was

analyzed across the populations and compared between species. To quantify size dependence of sex expres¬

sion, a GENMOD procedure with gamma distribution was used with DBH as a response variable. Parameter

estimates were analyzed by maximum likelihood and p-values were calculated using % 2 statistics. Similar

analyses were performed in SAS to analyze the relationship between distance and sex expression as hermaph¬

roditism. The relationships between DBH and NND were analyzed using REG procedure in ANOVA with DBH

as the response variable.

RESULTS

Breeding System, Sex Ratios and Lability of Sexual Expression

Out of 408 M. rubra trees studied across nine populations, 42 trees (10.3%) were hermaphrodites, and out of

269 M. alba trees across thirteen sites, 32 trees (12.3%) were hermaphrodites (Table 1)—establishing that both

species exhibit a subdioecious breeding system with males, females and hemaphrodites (see also below, re¬

garding labile sex expression). In addition, the ratio of males to females deviated significantly from a 1:1 ratio

with the proportion of males being greater than that of females for each species (Fig. 1): % 2 1 = 17.36 for M. alba,

% 2 1 = 44.99 for M. rubra, and p = <0.0001 for each species. The cumulative sex ratio between the two species

didn’t differ significantly (% 2 1 = 1.26, p = 0.2623), but was more strongly male-biased in M. rubra. Inter-year sex

ratio variation was not significant within each species (% 2 2 = 1.71, p = 0.4262 for M. alba;y} 2 = 0.06, p = 0.9701

for M. rubra; Table 2); however, the interspecific difference in sex ratio was significant (% 2 1 = 11.26, p = 0.001).

The proportion of males of M. alba increased in 2007 due to mortality of two female trees to fire, and some sex

switching (see below). The sex ratios were consistently male-biased for both species for all the three years.

Sex ratios were consistently male-biased across all populations (Figs. 2, 3). Interpopulation variation in

sex ratio within species was not statistically significant (% 2 12 = 7.70, p = 0.86 for M. alba; % 2 8 = 5.60, p = 0.69 for

386

Journal of the Botanical Research Institute of Texas 9(2)

Table 1. Sampled sites with the information on sample size (N) and number of males (M), females (F) and hermaphrodites (H) for Morns alba (MA) and M. rubra (MR).

Sampled site

Species

Alma: along Kuenzli Creek, Wabaunsee Co.

Anneberg Park: City of Manhattan, Riley Co.

Fort Riley: along Three Mile Creek, Riley Co.

Kings Creek: KPBS, Riley Co.

Linear ParkTrail: along Wildcat Creek, Riley Co.

Farnum Creek: Milford Lake, Geary Co.

Timber Creek: Milford Lake, Clay Co.

Pottawatomie Lake No. 1: Pottawatomie Co.

Pottawatomie Lake No. 2: Pottawatomie Co.

Shane Creek: KPBS, Riley Co.

Slough Creek: Perry Lake, Jefferson Co.

Tuttle Creek SE: Beach Drive below dam, Pottawatomie Co.

Tuttle Creek West: Observation Point Drive west of dam, Riley Co.

Fig. 1. Sex ratio deviation in Morns in the Flint Hills region of Kansas. Sex ratio is expressed as the proportion of males (males / [males + females];

hermaphrodites not included). 95% confidence intervals are indicated. The line at 0.5 on y-axis corresponds a 1:1 ratio of males to females (i.e. equal

numbers of males and females; values above 0.5 indicate more males than females). The letter Vindicates a significant deviation of sex ratio from 1:1.

Nepal et al., Sex expression and sex ratio variation in mulberries (Morus)

387

Table 2. Inter-year sex ratio in Morus at Kings Creek. Sex ratio is expressed as the proportion of males, and the 95% confidence limits (CL) are indicated. Values

denoted by similar letters are not significantly different from each other. Chi Square probability of deviation at 0.5 of alpha, degree of freedom and sample size are

represented by p (y2), df and N, respectively.The hermaphrodites were not included in the analysis. Significant sex ratio deviation from unity is marked with an asterisk.

Species

Year

2005

2006

2007

Morus alba

Sex ratio

0.5937

0.6060

0.7407

Upper CL

0.4192

0.4335

0.5471

Lower CL

0.7474

0.7556

0.8710

P(X2)

0.2917 b

0.2268 b

0.0168 b*

Morus rubra

Sex ratio

0.7666

0.7500

Upper CL

0.6436

0.6258

Lower CL

0.8566

0.8433

P(X2)

<0.0001 a*

0.0002 a*

Fig. 2. Sex ratio variation with 95% confidence intervals across 13 populations of M. alba in the Flint Hills region of Kansas. Sex ratio is expressed as the

proportion of males. The letter "a" indicates a significant deviation of sex ratio from 1:1.

388

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 3. Sex ratio variation with 95% confidence intervals across nine populations of M. rubra in the Flint Hills region of Kansas. Sex ratio is expressed as

the proportion of males. The letter "a" indicates a significant deviation of sex ratio from 1:1.

M. rubra). Two M. alba populations, Kings Creek and Pottawatomie Lake No.l, were significantly male-biased

(% 2 i = 4.80, p <0.05 and % 2 : = 6.03, p <0.05, respectively; Fig. 2); and five M. rubra populations were signifi¬

cantly male-biased: Fort Riley (% 2 : = 4.69, p <0.05), Kings Creek (% 2 : = 14.20, p <0.001), Shane Creek (% 2 : =

8.86, p <0.01), Pottawatomie Lake No. 2 (% 2 : = 7.69, p <0.05), and Slough Creek (% 2 : = 6.54, p <0.05; Fig. 3).

When compared between species, the sex ratio of M. rubra was consistently more strongly male-biased than

that of M. alba.

Out of 75 M. rubra trees observed at Kings Creek across three years (2005,2006 and 2007), changes in sex¬

ual expression (among female, male and hermaphrodite morphs) were rarely observed: 5% (4 trees) switched

sex from 2005-2006, and 8% (6 trees) changed sex from 2006-2007 (Table 3). Of those trees that changed

sexual status, only one of them changed sex both years. Among 38 M. alba trees studied, four trees exhibited

six changes in sex expression (three changes from 2005-2006 and three from 2006-2007; two trees changing

both years). In all but one case (a single observed female to male change in M. alba tree #267), changes in¬

volved switching among unisexual and hermaphrodite conditions (Table 3). Furthermore, while we observed

interesting variation in distribution of male and female flowers on individual hermaphrodite trees (e.g., male

and female flowers on separate catkins vs. mixed catkins with flowers of both sexes), we highlight the varia¬

tion with respect to functional sex expression of the individual plant (male, female, hermaphrodite; Table 3).

Sex Expression Relative to Size and Nearest Neighbor Distance

The size distribution of all sexual morphs in both species is shown in Fig. 4. No significant differences in size

Nepal et al., Sex expression and sex ratio variation in mulberries (Morus)

389

Table 3. Change of sexual status in Morus at Kings Creek between 2005,2006 and 2007. M = male, F = female, MF = hermaphrodite with mixed catkins, and

M+F = hermaphrodite tree with male and female catkins on separate branches. Functional changes (i.e., excluding changes among different expressions of

hermaphroditism: M+F, MF) are shown in bold face font and discussed in the text.

M.alba

Sexual status change

M. rubra

Sexual status change

Tree ID

2005 2006

2006 2007

Tree ID

2005 2006

2006 -*2007

264

M+F+MF M

M^M+F

267

M+F+MF F

F^M

M+F^F

284

F^M+F

M+F

M+F^F

294

F^M+F

M+F M+F+MF

M+F+MF

M+ F MF+M+F

M+F+MF M+F

F^M+F

M+F M+F+MF

M+F+MF ^F

M+F^M

M+F M+F+MF

M+F+MF

M+F M+F+MF

M+F+MF

F^M+F

M+F

100

F^M+F

M+F F

among male, female and hermaphrodite individuals were found within species (% 2 2 = 1.41, p = 0.49 for M. alba ;

X 2 2 = 3.12, p = 0.21 for M. rubra). Males and females of M. alba were statistically larger than those of M. rubra

(Fig. 4). In M. alba, the sex ratio did not differ from 1:1 in the smaller size classes (DBH <20 cm), but differed

390

Journal of the Botanical Research Institute of Texas 9(2)

significantly at larger size classes and was male-biased. In M. rubra, the sex ratios differed from 1:1 in all size

classes. Regression analysis showed no association between the size of the tree (DBH) and the nearest neighbor

distance (NND) for both M. alba (r 2 = 0.0032, p = 0.52) and M. rubra (r 2 = 0.0013, p = 0.46).

The average nearest neighbor distance for hermaphrodites was significantly greater than that for males

and females (% 2 2 = 28.91, p <0.0001 for M. alba ; % 2 2 = 51.41, p <0.0001 for M. rubra; Fig. 5). This pattern was

based on average distances, whereas there were some individual hermaphrodites occurring near other indi¬

viduals, and some males and females occurring at more distant locations. The average nearest neighbor dis¬

tance did not differ between males and females within each species or between the two species.

DISCUSSION

Subdioecy, Sex Ratios and Lability of Sex Expression

Our study documents subdioecy in M. alba and M. rubra. Across 22 populations in the Flint Hills of Kansas (13

of M. alba and nine of M. rubra), the majority of the trees are consistently unisexual, but approximately 10% of

the individuals are hermaphrodites. Moreover, focused study of populations at Kings Creek over three years

found that 10-12% of individuals changed sexual expression (among male, female and hermaphrodite sexual

morphs) at least once across these years; all but one of these observed changes were between unisexual and

hermaphrodite conditions. There was no unidirectional change in sex expression across years (i.e., unisexual

individuals switched from one sex to another or to hermaphroditism, and the hermaphrodites switched to be¬

ing unisexual; Table 3), and all of the changes occurred in plants that expressed hermaphroditism in at least

one year. The subdioecious breeding system in the studied species of Morus might represent a transition in the

hermaphroditism-dioecy continuum, and could be an adaptive strategy, as previously discussed (see Case et

al. 2008), for reproductive assurance by opportunistic selhng, benehtting sexual specialization and avoiding

the effect of inbreeding depression. The evolution of subdioecy is believed to have occurred through a mon-

oecy-paradioecy pathway (where individuals in the population are selected for in such a way that one becomes

increasingly male and the other becomes increasingly female by gradual divergence; Delph & Wolf 2005).

In the present study, both M. alba and M. rubra had some individuals that exhibited sex expression plas¬

ticity. Such a strategy is not considered very common in woody perennials, although it is common in herba¬

ceous dioecious species (Korpelainen 1998). Some examples of tree species with plasticity in sex expression

include Acer rubrum (Sakai 1990), Acer rufinerve (Nanami et al. 2004), Bischofiajavanica (Yamashita & Abe

2002), Clusia nemorosa (Lopes & Machado 1998), Dombeya ciliata (Humeau et al. 2000), Dombeya delislei

(Humeau et al. 1999), Hebe subalpina (Delph 1999), Lindera benzoin (Primack 1985), Myristica insipida

(Armstrong & Irvine 1989), and Thymelaea hirsuta (El-Keblawy & Freeman 1999). Effects of environmental

factors on sex expression were highlighted in all of these studies except in the case of A. rufinerve, where dete¬

riorating plant health resulted in a change from male to female. Males of some plant species are reported to be

inconstant, as in D. ciliata (Humeau et al. 2000), D. delislei (Humeau et al. 1999), and C. nemorosa (Lopes &

Machado 1998). In Morus, while the majority of the trees were male or female, some males and females were

inconstant/labile from a given year to the next. These findings are similar to those of reports on B. javanica

(Yamashita & Abe 2002) and T. hirsuta (El-Keblawy & Freeman 1999), wherein both sexes were inconstant.

This labile sex expression in Morus may have evolved for successful reproduction under unpredictable envi¬

ronmental conditions, ensuring outcrossing and the maintenance of genetic variation within populations.

In some other subdioecious species, studies have shown that sex expression is determined by both genet¬

ics and by genotype - environment interactions. In Atriplex canescens, sex of the majority of unisexual indi¬

viduals was genetically fixed as male or female, while sex varied in other individuals ranging from unisexual

individuals to hermaphrodites with various proportions of male and female flowers (McArthur et al. 1992). In

Rumex nivalis, sex expression was largely determined by genetics, however sex ratios in the progeny depended

on pollination intensity (Stehlik et al. 2008). In Morus, as the majority of the individuals are consistently male

or female and only a small number of individuals are inconstant, as in the case of Atriplex (McArthur et al.

1992), sex expression may be controlled by both genetics and genotype - environment interactions. Further

Nepal et al., Sex expression and sex ratio variation in mulberries (Morus)

391

Fig. 5. Relationships between the nearest-neighbor-distance (NND) and sex types in Morus. The symbols denote mean with 95% confidence intervals.

investigation on the effects of environmental factors on sex determination may provide insights into the main¬

tenance of subdioecy in Morus.

Groundwork for Future Studies Investigating Patterns

In the present study, both species exhibited significant sex ratio deviations. The male-biased sex ratios in mul¬

berries are in line with those reported in other dioecious species (see Bierzychudek & Eckhart 1988; House

1992; Thomas & LaFrankie 1993; Nicotra 1998). These studies have provided proximate causes of male-biased

sex ratios as precocious male flowering, more frequent flowering of males than females (Thomas & LaFrankie

1993; Nicotra 1998) and higher female mortality (Bierzychudek & Eckhart 1988). For most dioecious species

with known sex ratios, sex ratios are consistently male-biased (Delph 1999). Species such as Compsoneara

spucei (Bullock 1982), Myristica insipida (Armstrong & Irvine 1989), and Rhamnus alaternus (Guitihn 1995)

have a stable sex ratio (i.e., 1:1), while species such as Rumex acetosa (Korpelainen 1998) have female-biased sex

ratios. Acer negundo shows male-biased sex ratios in drier habitats and female-biased sex ratios in the moist

areas (Jing & Coley 1990). Unlike in Morus, Yamashita and Abe (Yamashita & Abe 2002) show significant in-

ter-year variation in sex ratios in Bischofiajavanica, with a large number of individuals switching their sex. The

observed inter-year variation in sex expression in both species of Morus studied warrants further

investigation.

Size dependent sex expression has been documented in several dioecious species including Bischofia

392

Journal of the Botanical Research Institute of Texas 9(2)

javanica: the smallest trees were males, medium sized trees inconstant and the largest trees were females

(Yamashita and Abe 2002). We found no size dependence of sex expression in Morus. In M. alba, however, the

frequency of males at the higher size class was higher than those at the smaller size classes (Fig. 4). If male and

female individuals differ in reproductive costs, allocation would be greater in the sex with higher reproductive

costs, resulting in size difference in the sexes (Lloyd & Bawa 1984). Absence of inter-sex difference in size in

Morus suggests that male and female individuals do not differ in reproductive costs (the cost of pollen produc¬

tion in males is comparable to the production in females of fruits). In order to gain insights into reproductive

allocation in Morus, further studies—including identifying the resources that drive expression of one sex

versus the other—are needed.

Stehlik et al. (2008) demonstrated that the proximity of males and females affected the sex ratios in

Rumex nivalis: the closer the males and females, the stronger were the female-biased sex ratios. In Morus, we did

not detect an association between tree size and the nearest neighbor distance. The Morus species are mostly

understory trees with patchy distributions (particularly M. rubra), and they may be experiencing pollen limita¬

tion rather than pollen excess (Ashman et al. 2004). Male-biased sex ratios in these species occurring in a

heterogenous environment may therefore ensure outcrossing and reduce the cost of selhng.

We found no strict spatial segregation of the sexes (SSS) in Morus in the populations studied. In species

that do exhibit SSS such as Acer negundo (Freeman et al. 1997), and Juniperus virginiana (Lawton & Cothran

2000), females often dominate in resource rich habitats (Lloyd & Bawa 1984). Since Morus species are sparsely

distributed understory trees and pollen movement from tree to tree is essential for reproductive success, the

individuals of the same species are less likely to compete for the same limiting resources. Therefore, the species

are less likely to undergo selection for SSS that would further reduce the reproductive success. Our findings are

similar to those for Silene grandiflora, an understory plant that did not exhibit SSS (Bawa & Opler 1975). Some

studies have suggested several factors related to spatial distribution that affect the reproduction of dioecious

species such as distance between males and females (Mack 1997), flowering frequency, effective population

size (Nunney 1995), pollinator abundance and flight behavior (Stacy et al. 1996), etc. Some strategies of plants

such as woody perennial habit (Baker & Cox 1984), production of fleshy fruits (Bawa 1980), production of

multi-seeded fruits and dispersal by birds (Baker & Cox 1984), leaky dioecy, andparthenocarpy (Venkatasamy

et al. 2007) for reproductive assurance have been suggested. The two species of Morus, which are subdioecious,

woody perennials, parthenocarpic (Barbour et al. 1973), and produce multi-seeded fleshy fruits that are dis¬

persed by birds-all potential strategies for reproductive assurance.

Our study found similar reproductive strategies in M. alba and M. rubra in the Flint Hills region of the

Great Plains: each species exhibits a subdioecious breeding system, with a male-biased sex ratio and some la¬

bility in sexual expression. These findings in the two species raise the question of whether similar reproductive

biology may be common in the genus. We did not find evidence for size dependence of sex expression.

Hermaphrodites were, on average, located farther from other plants than were unisexual individuals (although

this general pattern was based on average distances; hermaphrodites sometimes occur in close proximity to

males and females). This work suggests many avenues for future study; in particular, further investigation on

the role of pollen limitation may shed light on the biological significance of patterns and changes in sexual

expression in Morus. Future multi-year studies including additional individuals and populations will be valu¬

able. Documentation of breeding systems of these species advances our basic understanding of the taxa and

furthers Morus as an interesting study system for studies of reproductive and evolutionary biology.

ACKNOWLEDGMENTS

We thank John Barbur, Bill Markley, Marj Markley, Valerie Wright, Jim Larkin, Jim Rivers, Chris Hein, and

Tom Van Slyke for held assistance; and staff and land managers of the various sites for permission to study

plants. David Hartnett and Leigh Murray from Kansas State University provided useful discussion on the

manuscript and assisted in data analyses, respectively. We are grateful to Alan Whittemore and one anony¬

mous reviewer for valuable comments that improved the manuscript. Support to the first author from the

Nepal et al., Sex expression and sex ratio variation in mulberries (Morus)

393

South Dakota Agricultural Experiment Station is gratefully acknowledged; as is support throughout the study

from the Konza Prairie LTER program and the Kansas Agricultural Experiment Station (Contribution No.

15-451-J).

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JOURNAL NOTICE

Sabeeha S. Merchant, Wilhelm Gruissem, and Donald Ort, eds. 2015 (Apr). Annual Review of Plant Biology,

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Contents of Volume 66:

1. The Wayward Hawaiian Boy Returns Home—D. Gonsalves

1. From the Concept of Totipotency to Biofortified Cereals— I. Potrykus

2. The Structure of Photosystem II and the Mechanism of Water Oxidation in Photosynthesis— J.-R. Shen

3. The Plastid Terminal Oxidase: Its Elusive Function Points to Multiple Contributions to Plastid Physiology—WJ. Nawrocki, N.J.

Tourasse, A. Taly, F. Rappaport, & F.-A. Wollman

4. Protein Maturation and Proteolysis in Plant Plastids, Mitochondria, and Peroxisomes— K.J. van Wijk

5. United in Diversity: Mechanosensitive Ion Channels in Plants— E.S. Hamilton, A.M. Schlegel, & E.S. Haswell

6. The Evolution of Plant Secretory Structures and Emergence of Terpenoid Chemical Diversity— B.M. Lange

7. Strigolactones, a Novel Carotenoid-Derived Plant Hormone— S. Al-Babili & H.J. Bouwmeester

8. Moving Toward a Comprehensive Map of Central Plant Metabolism—R. Sulpice & PC. McKeown

9. Engineering Plastid Genomes: Methods, Tools, and Applications in Basic Research and Biotechnology—R. Bock

10. RNA-Directed DNA Methylation: The Evolution of a Complex Epigenetic Pathway in Flowering Plants—M.A. M atzke, T. Kanno, &

A.J.M. Matzke

11. The Polycomb Group Protein Regulatory Network— I. Mozgova & L. Hennig

12. The Molecular Biology of Meiosis in Plants—R. M ercier, C. Mezard, E.Jenczewski, N. Macaisne, & M. Grelon

13. Genome Evolution in Maize: From Genomes Back to Genes— J.C. Schnable

14. Oxygen Sensing and Signaling— JJ. van Dongen & F. Licausi

15. Diverse Stomatal Signaling and the Signal Integration Mechanism—Y. Murata, I.C. Mori, & S. Munemasa

16. The Mechanism and Key Molecules Involved in Pollen Tube Guidance—T. Higashiyama & H. Fakeuchi

17. Signaling to Actin Stochastic Dynamics— J. Li, L. Blanchoin, & C.J. Staiger

18. Photoperiodic Flowering: Time Measurement Mechanisms in Leaves— Y.H. SongJ.S. Shim, H.A. Kinmonth-Schultz, & T. Imaizumi

19. Brachypodium distachyon and Setaria viridis: Model Genetic Systems for the Grasses—T.P. BrutnellJ.L. Bennetzen, & J.P. Vogel

20. Effector-Triggered Immunity: From Pathogen Perception to Robust Defense— H. Cui, K. Fsuda, &J.E. Parker

21. Fungal Effectors and Plant Susceptibility—L. To Presti, D. Lanver, G. Schweizer, S. Fanaka, L. Liang, M. Follot, A. Zuccaro, S. Reissmann, &

R. Kahmann

22. Responses of Temperate Forest Productivity to Insect and Pathogen Disturbances— C.E. Flower & M.A. Gonzalez-Meler

23. Plant Adaptation to Acid Soils: The Molecular Basis for Crop Aluminum Resistance— L.V. Kochian, M.A. Pineros, J. Liu, & J.V. Magalhaes

24. Terrestrial Ecosystems in a Changing Environment: A Dominant Role for Water— C.J. Bernacchi &A. VanLoocke

Errata

J.Bot. Res. Inst. Texas 9(2): 396.2015

FROM CRO-MAGNON TO KRAL: A HISTORY OF BOTANY IN ALABAMA

L.J. Davenport

Biological & Environmental Sciences

Samford University

Birmingham, Alabama 35229, U.S.A.

ljdavenp@samford.edu

ABSTRACT

Due to its great diversity of terrestrial and aquatic habitats, from mountains to the Gulf coast, Alabama supports an inordinate number of

species, including several thousand vascular plants. These species are distributed, from north to south, across the Interior Plateau, Pied¬

mont, Southwestern Appalachian, Ridge & Valley, Southeastern Plains, and Southern Coastal Plain ecoregions.

Alabama’s plant life has been studied and utilized since ancient times. Such studies began with Paleo-Indians, Native Americans, and

European explorers. During the early 1800s, the major botanical “players” were pioneers, settlers, travelers, academics, and medical doc¬

tors. The latter half of the 19 th century was defined by the works of Mobile pharmacist Charles Mohr, culminating in his 1901 magnum opus,

Plant Life of Alabama. Roland Harper, working mainly through the Geological Survey of Alabama, dominated the first half of the twentieth

century. Floristic studies have taken hold since 1950, with “hotbeds” for such studies established at each of the state’s universities.

No current botanist stands as tall as Robert Krai. His voluminous knowledge of Alabama’s flora has been recorded in monographs,

revisions, federal reports, floristic accounts, checklists, and websites. His thousands of Alabama specimens, now housed at BRIT, constitute

a botanical treasure of inimitable value.

RESUMEN

Debido a la gran diversidad de habitats terrestris y acuaticos, de las montanas a la costa del Golfo, Alabama soporta un enorme numero de

especies, que incluyen varios miles de plantas vasculares. Estas especies estan distribuidas, de norte a sur, a traves de la Meseta Interior, de

las ecorregiones Piedemonte, suroeste de los Apalaches, cadena y valle, llanuras del sureste, y llanura costera sur.

La vida vegetal de Alabama ha sido estudiada y utilizada desde los tiempos antiguos. Tales estudios empiezan con los paleo-indios,

americanos nativos, y exploradores europeos. Durante los 1800s tempranos, los mayores actores botanicos fueron los pioneros, colonos,

viajeros, academicos, y doctores en medicina. La ultima mitad del siglo XIX estuvo definida por los trabajos del farmaceutico ambulante

Charles Mohr, que culmina en su opera magna de 1901, Plant Life of Alabama. Roland Harper, trabajando principalmente para el Geological

Survey de Alabama, domino la primera mitad del siglo XX. Los estudios floristicos se han sucedido desde 1950, con “semilleros” para tales

estudios establecidos en cada universidad estatal

Ningun botanico actual sobresale tanto como Robert Krai. Su voluminoso conocimiento de la flor de Alabama se ha recogido en mo-

nografias, revisiones, informes federales, informes floristicos, catalogos, y paginas web. Sus miles de espechnenes de Alabama, actualmente

en BRIT, constituyen un Tesoro botanico de inimitable valor.

Alabama is blessed with a tremendous diversity of species. Although 30th among the United States in total land

area, the state ranks first for its number of known species among the 26 states east of the Mississippi River

(Duncan 2013). A recent checklist by Krai et al. (2011) records 3743 species of Alabama vascular plants alone.

Such high rankings and high numbers are due directly to the diversity of habitats available to be colonized.

GENERAL PHYSIOGRAPHY AND ECOREGIONS

Several attempts have been made to document and categorize Alabama’s plant communities and ecoregions,

including those by Mohr (1901) and Harper (1943a). The current system, devised by Griffith et al. (2001), is

described below (roughly from north to south); a variety of very detailed maps can be downloaded from the

accompanying website (see References).

The Interior Plateau Ecoregion extends from southern Indiana to northern Alabama. Its tablelands and

plains, cut by the west-flowing Tennessee River, developed over Mississippian to Ordovician limestones, sand¬

stones, and shales. Originally covered by deciduous forest, this ecoregion (centered around the city of Hunts¬

ville) was an important agricultural area much favored by early settlers.

The Southwestern Appalachian Ecoregion, stretching to Kentucky, enters Alabama in its northeast cor¬

ner, along with the Tennessee River. The Mississippian to Ordovician limestones, dolomites, and shales of the

J. Bot. Res. Inst. Texas 9(2): 397 - 431.2015

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Journal of the Botanical Research Institute of Texas 9(2)

Sequatchie Valley subunit create an agriculturally rich area. Much of another important subunit—called either

the Shale Hills, Warrior Coal Field, Coal Basin, or Coal Measures—has been strip-mined, with concomitant

alterations of land forms, drainage patterns, soils, and vegetation. The Dissected Plateau subunit contains plant

species, such as eastern hemlock (Tsuga canadensis) and many rare ferns, that migrated south during the Ice

Ages and remain in its cool, protected ravines.

The Ridge & Valley Ecoregion stretches southwest from Pennsylvania, entering Alabama near its north¬

eastern corner and ending near its geographic center. Here parallel ridges and valleys, created by much folding

and faulting, are wedged between the Piedmont to the east and the Interior Plateau to the west. A variety of

geologic materials underlie this ecoregion, including the iron ore of Red Mountain near Birmingham. Drainage

is largely through the Cahaba and Coosa river systems, flowing generally southwest.

The Piedmont is a triangular ecoregion jutting into central Alabama from the eastern (Georgia) border. Its

fine-textured soils have developed over a mixture of Precambrian and Paleozoic metamorphic and igneous

rocks. This ecoregion contains the highest elevations in the state, including Mount Cheaha (2407 ft), and once

housed extensive montane longleaf pine (Finns palustris) forests, which were logged out in the 1890s (Harper

1943a). Drainage is via the Tallapoosa and Coosa rivers, which combine to form the Alabama River east of

Montgomery, the state capital.

The Southeastern Plains Ecoregion developed over the Cretaceous and Tertiary sands, silts, and clays of

former coastlines. Its many subdivisions take up most of the state, running in broad bands from Alabama’s

northwest corner nearly to the Gulf of Mexico in the south. Several distinctive subunits are found here, includ¬

ing the Blackland Prairie or Black Belt “cinching” the midsection of the state. Here the fertile soils developed

from Cretaceous chalks, supporting eastern red-cedar (Juniperus virginiana) and patches of bluestem prairie.

(The prehistoric extent of Alabama’s prairies was apparently not continuous and has led to much debate; see

Rostlund [1957], Jones & Patton [1966], and Barone [2005]). Another unique subunit, the Dougherty Plain or

Wiregrass Region, occupies the southeastern corner of the state. This karst topography contains many sink¬

holes and springs, while the sandy soils once supported park-like growths of longleaf pines maintained by

frequent wildfires.

The Southern Coastal Plain Ecoregion extends from South Carolina through Georgia and Florida to Ala¬

bama, then west through Mississippi and Louisiana. In Alabama, much of this ecoregion consists of the Flood-

plains & Low Terraces subunit of the Mobile or Tensaw Delta, a series of swamps and backwaters based on

river alluvium. A second subunit, the Gulf Barrier Islands & Coastal Marshes, consists largely of saline

marshes, pine forests, and sand dunes. Much of this final subunit—especially the Fort Morgan Peninsula and

Dauphin Island, twin sentinels “guarding” the entrance to Mobile Bay—serve as important refueling areas for

birds and butterflies during their annual trans-Gulf migrations.

NATIVE ALABAMIANS AND THEIR PLANT USES

Alabama has a rich Native American history. The state’s very name—as well as many counties, towns, rivers,

and natural landmarks—bear witness to this history (for examples, see Foscue 1989).

The four main groups of Native Americans occupying prehistoric Alabama were the Chickasaws, Chero-

kees, Creeks (including the Alabamas), and Choctaws. The Chickasaws and Cherokees inhabited the north¬

ernmost portions of the future state, including the Tennessee Valley of the Interior Plateau and Southwestern

Appalachians. The Creeks ruled the diverse ecoregions of the majority of the state except for its western edge,

which was held by the Choctaws.

The hunter-gatherer predecessors of these four tribes left their marks on Alabama, most notably (and

quite literally) in Russell Cave (Fig. 1) in the Southwestern Appalachian Ecoregion near the Tennessee border.

Excavations at this seasonal campsite began in 1951, revealing its use by hunter-gatherers from the Paleo-Indi-

an Period (about 10,000 years ago) through late Woodland times (to 1000 CE). Unfortunately, these excava¬

tions concentrated on point types and potsherds and ignored direct evidence of plant use, like carbonized

seeds and shells. But some indirect evidence exists—pits for storage of nuts and seeds, plus “nut stones” for

Davenport, A history of Botany in Alabama

399

Fig. 1. Mouth of Russell Cave near the Tennessee border, Jackson County, Alabama.

cracking and grinding (Griffin 1974). Nuts likely utilized by Russell Cave inhabitants include hickories

(Carya), oaks (Quercus), and black walnuts (Juglans nigra); available wild fruits were muscadines ( Vitis rotundi-

folia), blackberries (Rubus), pawpaws ( Asimina ), maypops ( Passiflora incarnata), and persimmons ( Diospyros

virginiana). Tubers of the American groundnut ( Apios americana) may have been gathered as well (Heiser

1993) in this part of North America.

In early studies, Miller (1960) recovered a small basket with charred goosefoot ( Chenopodium ) seeds from

Russell Cave. While its exact provenience is now under question, his find lends support for an Eastern Agricul¬

tural Complex that predates the arrival of established agricultural practices from Mesoamerica (see Smith

1985, 1989). Small grains such as goosefoot, marsh elder ( Iva ), and smartweed ( Polygonum ) might have been

grown in plots, easing the transition to later, more intensive farming of the Three Sisters from Mesoamerica—

corn (Zed), beans ( Phaseolus ), and various squashes ( Cucurbita ).

Caddell (1982) provides some evidence for an Eastern Agricultural Complex in Alabama. Using more

modern techniques than those employed at Russell Cave, she analyzed the botanical remains from five sites in

the Tombigbee River Valley of west-central Alabama. Hickory, oak, and walnut shells dominated most levels;

the above-mentioned wild fruits were likewise apparent; and seeds of goosefoot, smartweed, amaranth (Ama-

ranthus), canary grass ( Phalaris ), and many others were represented. Whether these latter seeds were gathered

from the wild or grown in defined plots is, of course, unknown.

By the Mississipian Period (after 500 CE), the agricultural proficiencies of Alabama peoples were manifest

in centers like Moundville along the floodplain of the Black Warrior River, which utilized the full complement

of Mesoamerican foodstuffs. The influence of ancient Mexico upon Moundville is also revealed in its Aztec-like

symbols and the building of temple mounds.

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Journal of the Botanical Research Institute of Texas 9(2)

Although Mississippian culture, as at Moundville, disappeared by 1400 CE, many aspects remained. The

annual Green Corn Ceremony has long been a part of Alabama life, celebrating the successful growth and

harvest of this vital, sustaining grain. Most Southeastern tribes observed such ceremonies, continuing even

after their Trail of Tears deportation to Oklahoma Territory (Swanton 1946; Hudson 1976).

Early European travelers and correspondents were also intrigued by a controversial ritual involving the

Black Drink. The Creek Indians of Alabama and Georgia, especially, used the yaupon holly, Ilex vomitoria —

one of the few Southeastern plants with a high caffeine content—in a dawn ritual that has generated many

humorous anecdotes and some serious academic debate (Hudson 1979; Davenport 2003).

EARLY EXPLORERS AND THEIR DESCRIPTIONS

Starting with Columbus’ first voyage to the New World in 1492, Europeans focused on exploring that world

and exploiting its riches. The first such explorer in the southeastern United States was Hernando de Soto

(15007-1542). De Soto’s exact itinerary through Alabama is only roughly known, and the southwestern Ala¬

bama site (“Mabila”) of his epic battle with the giant Chief Tascaluza (Black Warrior) is still being questioned

(Badger & Clayton 1985; Hall 1987). But apparently De Soto did cross the Coosa River near Childersburg,

whose sign boards (“The Oldest City in America”) proclaim founding by him in 1540. Although four chroni¬

clers recorded De Soto’s travels across Alabama and the Southeast, they provide no real insights or descriptions

of its vegetation.

The Spanish explorer Tristan de Luna (1519-1571) followed in 1560, restricting his incursion to the Ala¬

bama coastline; again, he left no real descriptions of specific plants and vegetation types that he saw. The

French-Canadian colonist Pierre Le Moyne d’Iberville (1661-1706) landed at Dauphin (Massacre) Island,

south of Mobile, in early 1699; there he discovered “all kinds of trees, oaks, elm, ash, pines, and other trees I do

not know, many creepers, sweet-smelling violets, and other yellow flowers ...” (McWilliams 1981). He and his

younger brother, Jean-Baptiste Le Moyne de Bienville (1680-1767), established several outposts, including the

current site of Mobile in 1711 (Atkins 1994).

The first trained naturalist to explore Alabama was William Bartram (1739-1823; Fig. 2). Bartram’s Trav¬

els (1791) show that he entered the east-central part of the state in July 1775 by crossing the Chattahoochee

River near Uchee in Russell County. (For naturalists, the preferred edition of Bartram’s Travels is by Francis

Harper [1958].) Continuing as part of a small caravan, he followed the Old Federal Road on horseback, in

roughly a southwesterly direction. He was fascinated by “the plains” or Black Belt and included this very ac¬

curate description: “... [I]t lies on a deep bed of white, testaceous, limestone rock, which in some places re¬

sembles chalk, and in other places are strata or subterrene banks of various kinds of sea shells,... these dissolv¬

ing near the surface of the earth, and mixing with the superficial mould, render[ing] it extremely productive.”

Bartram continued his overland trek to the edge of the Tensaw Delta northeast of Mobile, then crossed by boat

to that city. He returned to the Delta and explored its northern reaches for several weeks, then finally set sail for

his native Philadelphia.

Bartram’s Travels (1791) provide us with the most vivid and accurate descriptions of primeval Alabama.

Bartram’s knowledge of Southeastern plant life, especially, was unsurpassed. However, his delay in publication

“cost” him several important botanical species, which instead were described by others.

While exploring the Tensaw Delta during his final few weeks in Alabama, Bartram described three new

species; two of these ( Myrica inodora and Magnolia pyramidata) are still credited to him. He greeted a third one,

Oenothera grandiflora, with his characteristic enthusiasm: “... [A] few miles above Taensa, I was struck with

surprise at the appearance of a blooming plant, gilded with the richest golden yellow, ... perhaps the most

pompous and brilliant herbaceous plant yet known to exist.” The seeds collected by Bartram that day became

widely distributed in Europe, prompting Aiton (1789) to describe it from plants growing at Kew. The Dutch

geneticist and co-re-discoverer of Mendelism, Hugo de Vries—due to its common occurrence and obvious

mutagenicity—used it for his seminal studies on mutations (see Cleland 1935). The 1912 visit of De Vries to

Bartram’s “Taensa” (now Dixie Landing), to pay his respects at the type locality of O. grandiflora, is depicted by

Davenport (2011).

Davenport, A history of Botany in Alabama

401

Fig. 2. William Bartram. (Courtesy of the Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, Pennsylvania).

PIONEERS, SETTLERS, AND TRAVELERS

The defeat of the Creek Indians at Horseshoe Bend of the Tallapoosa River in 1814 caused “Alabama fever” to

sweep the young nation. Thousands of settlers converged on the land (then part of Mississippi Territory) sub¬

sequently ceded to the United States—rich bottomland forests and fertile Black Belt prairies, all connected by

navigable waters to ship products worldwide. The land rush was on (for further details, see Atkins 1994).

Many early settlers entered Alabama via the Federal Road from the east, crossing the Chattahoochee and

following Bartram’s path toward Mobile, perhaps veering northwest to the rich Tombigbee Valley. Others ven¬

tured north from Mobile itself. A third group settled around Huntsville in the rich valleys of the Tennessee

River drainage.

402

Journal of the Botanical Research Institute of Texas 9(2)

While few settlers described the countryside, several travelers did. Anne Newport Royall (1769-1854),

traveling through and exploring northern Alabama from 1817-1822, published a volume of “letters” written to

her friend “Matt” back in her native Virginia (Royall 1830). (Such letters, describing the people and places

visited, were a popular literary device of that time.) Unlike other travelers’ notes, Royall’s letters contain a few

descriptions of Alabama native plant life and habitats. Near Melton’s Bluff on the Tennessee River (now inun¬

dated by 20 th century dams), she encountered bottomlands with river cane “as thick as the hairs on your head”;

higher up, “The land is so clear of undergrowth that you may drive a wagon any where through the woods.”

And she considered the area around Courtland (Lawrence County) as “the region of the Carolina pink [ Silene

caroliniana] and Colomba root [ Frasera caroliniensis]. Wagonloads of the latter may be gathered any where in

the woods; it, and the pink, cover the ground” (for other details of Royall’s life and exploits, see Davenport

[2014]).

During the 1830s, several British travelers took stagecoaches from Fort Mitchell (on the Chattahoochee)

to Montgomery, then steamed down the Alabama River to Mobile, essentially following Bar tram’s route except

by water; others took the same journey in reverse (Posey 1938). One such traveler was Scotsman James Stuart

(1775-1849), who noted: “Since passing the Chattahoochee River, the beauty of the country, so far as respect

trees and evergreens, has greatly increased. There are many splendid oaks, tulip-trees, chestnuts, and syca¬

mores skirting the woods. The Magnolia grandiflora is found in great numbers, dog-wood, Cornus florida, and

the red-bud, Cords Canadensis, of great size, covered with a profusion of the most brilliant colours. ...” South

of Montgomery, “the stage passed through the first prairie land that I have seen, consisting of large undulating

pastures, which seem never to have been covered with wood” (Stuart 1833).

Englishman James Silk Buckingham (1786-1855), traveling from Tuskegee to Montgomery, noted the fol¬

lowing changes near Cubahatchee Creek: “The soil now became richer on each side, and the woods were much

more variegated, as, besides the ever-succeeding pine, there was a thick underwood of various flowering

shrubs and trees, including magnolias, yellow jessamines, the dogwood, and the grape-vine, with a very beau¬

tiful tree called the willow-oak” (Buckingham 1842).

A more “royal” traveler than Mrs. Royall and Messrs. Stuart and Buckingham was Sir Charles Lyell

(1797-1875; Fig. 3). Scottish by birth and the premier geologist of that time, Lyell recorded his observations on

the geology, natural history, and people of Alabama during his second trip to the United States. He and his wife

left Columbus, Georgia, by stagecoach in late January 1846. Initially upset by the destruction observed in the

newly cleared country, Lyell still waxed poetic: “The sound of the wind in the boughs of the long-leaved pines

always reminded me of the waves breaking on a distant sea-shore, and it was agreeable to hear it swelling

gradually, and then dying away, as the breeze rose and fell.” At this early point in his visit, he also offered some

less-than-poetic generalizations about the people he met: “As we go southward, we see more cases of intoxica¬

tion, and hear more swearing” (Lyell 1849).

Continuing by railroad to Montgomery, Lyell got his first look at the Black Belt, “a broad zone of calcare¬

ous marl, constituting what is called the prairie, or cane-brake country, bare of natural wood, and where there

is so great a want of water. ...” His party then transferred to a steamboat for a long, winding trip to Mobile, and,

continuing up the Tombigbee and Black Warrior rivers to Tuscaloosa, Lyell admired “the canes on the borders

of the river ..., some of which I found to be thirty feet high.” Returning to Mobile, “on the banks of the Alabama

river the deciduous cypress and cotton[wood] trees were putting out their leaves, and the beautiful scarlet

seed-vessels of the red maple ... enlivened the woods.” And in the city itself, “for the first time, we saw the

beautiful evergreen, the yellow jessamine ( Gelsemium sempervirens), in full bloom ...” (Lyell 1849).

The extensive canebrakes, dominating the alluvial lands of the state, mesmerized pioneers, settlers, and

travelers alike. Their prehistoric extent in Alabama and the Southeast, importance to wildlife (such as bison

and canebrake rattlers), conversion to agriculture, and consequent loss (perhaps 98%) have been summarized

well by Platt and Brantley (1997) and Barone et al. (2008).

Davenport, A history of Botany in Alabama

403

Fig. 3. Charles Lyell. (Courtesy of the Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, Pennsylvania).

404

Journal of the Botanical Research Institute of Texas 9(2)

EARLY TEACHERS AND ACADEMICS

Two itinerant teachers are essential to the story of the early days of Alabama natural history The first, English¬

man Philip Henry Gosse (1810-1888), arrived in Mobile during May 1838 and headed northeast toward Mont¬

gomery by steamboat to seek a teaching position. Along the way, he was hired by a Black Belt planter to teach

in Pleasant Hill, southeast of Selma (Davenport 2010b).

Gosse spent just eight months in Alabama before returning to England. Over twenty years later, he used

his copious notes and line drawings to publish Letters from Alabama (Gosse 1859). His exquisitely detailed

water color paintings, long preserved by Gosse descendants, have recently been brought to light (Mullen &

Littleton 2010).

Gosse’s works are considered to be the first contributions by a trained naturalist residing (albeit briefly) in

Alabama. Although Gosse was interested primarily in insects (especially butterflies), his works do include

descriptions of host plants and key habitats.

A second resident/itinerant naturalist, Samuel Botsford Buckley (1809-1884), served as principal of the

Allenton Academy (in Wilcox County) from 1839-40 (Anonymous 1907). Buckley was mainly interested in

plant life, and, unlike Gosse, he published his Endings soon after his sojourn in Alabama. In his first such pa¬

per, Buckley (1843) described four species that are currently included (Krai et al. 2011) in the Alabama plant

list: Phacelia purshii, Carex styloflexa, Diervilla sessilifolia, and Thalictrum debile. Buckley left Alabama to return

to his native New England, then passed through again in 1859 on his way to Texas, where he later served as

State Geologist. (This was a time, in fact, when many Alabamians left the Old Southwest for the new one, often

inscribing “Gone To Texas” or “GTT” on their cabin walls [Atkins 1994].) During this brief visit, he discovered

Quercus durandii (Buckley 1860) “on the right hand side of the road, about three miles from Allenton” while

walking home from Camden, the county seat (see also Buckley 1881 and Dorr & Nixon 1985).

Three resident academics also contributed to our early knowledge of Alabama’s plant life. In the fall of

1853, Thomas P. Hatch (7-1855) was named Chair of Chemistry & Geology at LaGrange College, which was

located on Little Mountain in northwest Alabama. In early 1855, the school moved across the Tennessee River,

becoming Florence Wesleyan College and, much later, the University of North Alabama (see McDonald 1991).

Hatch died that same year, soon after submitting a “floral calendar” to the American Journal of Science and Arts

recording the blooming periods of local wildflowers (Hatch 1856). Many such calendars were kept during the

1850s, including one by nature philosopher Henry David Thoreau. They contain valuable data to support to¬

day’s concerns about climate change (Nijhuis 2007; Miller-Rushing & Primack 2008).

The Rev. Dr. Reuben D. Nevius (1827-1913) served as rector of Christ Episcopal Church in Tuscaloosa

during the 1850s and 1860s. During the spring of 1857, he and William Stokes Wyman (1830-1915), professor

of Latin at (and later President of) the University of Alabama, discovered an unusual shrub along the Black

Warrior River. Nevius (Fig. 4) sent specimens to Harvard’s Asa Gray, who named it Neviusia alabamensis (Gray

1859) after one of its discoverers. (The naming of this new genus later became controversial, as depicted by

Pollard [1900], Howard [1967], and Davenport [2000].) In that same paper, Gray described the succulent Se-

dum nevii in the reverend-doctor’s honor.

In his introduction to Plant Life of Alabama, Mohr (1901) paid homage to his predecessor Dr. Hezekiah

Gates (7-1849), “a successful apothecary at Mobile, [who] was the first collector of Alabama plants from the

coast region, whence he contributed valuable material to Torrey and Gray for their Flora of North America,

from the year 1836 to the early forties.” Later in that same volume, Mohr named Silphium gatesii in his honor.

Little else is known about Dr. Gates, except that he died in a St. Louis, Missouri, cholera epidemic. The

one-story Creole Plantation-style cottage that he built in 1841, now called the Gates-Daves House, still stands

on Dauphin Street in Mobile.

Davenport, A history of Botany in Alabama

405

Fig. 4. Reuben D. Nevius. (Courtesy of the Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, Pennsylvania).

406

Journal of the Botanical Research Institute of Texas 9(2)

AN ANTEBELLUM HERBAL

On 1 December 1847, twenty-one physicians convened in Mobile to found the Medical Association of the State

of Alabama (Holley 1982). One of the Association’s first acts was to form a committee to study and report back

on the uses of medicinal plants in their respective communities.

Such action was part of a national and regional trend. Similar reports had been published for the nation

(Clapp 1852), South Carolina (Porcher 1849), and Louisiana (Hale 1852). In Alabama’s case, the committee

members’ findings were published individually in the Transactions of the Medical Association of the State of Ala¬

bama from 1851-1855.

These reports differ in quality, quantity, and scientific value. Some authors show solid foundation in the

botanical sciences by following a systematic format, like that of Linnaeus (Smythe 1851), De Jussieu (Denny

1851; Bates 1853), Torrey and Gray (Cabell 1855), or “according to the natural order” (Jenkins 1854). One

(Clanton 1855) is an obvious copy of an earlier one; another (Batchelor 1853) concentrates on just one species,

Gelsemium sempervirens. While most describe the plants and their uses quite fully, Welch (1851) provides few

details. Two years after, Denny (1853) offered an apology for his earlier publication and a plea that “a universal

Congress of Scientific men” will soon “effectually disperse the great clouds of synonyms”—something still

hoped for today. And Smith (1852) included a botanical joke: Aletris aurea “Berry much” resembles the previ¬

ously mentioned A. farinosa.

Combined together, these ten publications by nine authors constitute an antebellum herbal, describing

medical treatments based on 241 species. No other sources record the medicinal use of plants—native, natu¬

ralized, and cultivated—in central Alabama at that time.

1860-1900

One of the most complete and useful of the above treatments of Alabama’s medicinal plants was by Prattville,

Alabama, physician Samuel Parrish Smith (1814-1891), mentioned above. His son, Eugene Allen Smith (1841-

1927), studied geology in Germany after the Civil War, and after completing his Ph.D., he was first appointed

Assistant State Geologist of Mississippi and then State Geologist of Alabama, based in Tuscaloosa. He served in

the latter capacity from 1873 until his death.

As part of his duties in his native state, Smith undertook long summer forays to examine and describe

Alabama’s geology and, especially, its mineral resources. Along the way, he collected plant specimens. By 1878

he had collected enough unusual species to warrant help from an expert, so he turned to Mobile pharmacist

Charles Mohr (1824-1901; Fig. 5).

Born Karl Teodor Mohr in Wurtemburg, Germany, Mohr trained mainly in chemistry (for Mohr’s life

story, see Smith 1901; Davenport 1979a, 1979b). With the revolutionary waves of 1848, he immigrated to the

United States, followed the Gold Rush to California, married a woman from the German community of Cincin¬

nati, and moved to Mexico. Seeking a healthier climate, politically and otherwise, he settled in Mobile, Ala¬

bama, in 1860, where he established a successful pharmaceutical business.

Mohr’s interest in Alabama plants, combined with Smith’s curiosity about the state’s resources, formed a

long and fruitful collaboration. The first “fruit” was a checklist of Alabama’s known plant life (Mohr 1880); the

final one was the 921-page Plant Life of Alabama (Mohr 1901), published jointly by the U.S. National Herbarium

and the Geological Survey of Alabama. The latter remains, to this day, as the only “complete” flora of the state.

Oddly, the first potential collaboration between Smith and Mohr was deferred to Apalachicola, Florida,

physician-botanist Alvan Wentworth Chapman (1809-1899). Smith had collected an unknown shrub with

Elaeagnus-like leaves along the Cahaba River north of Centreville in central Alabama (for more details on this

story, see Davenport 1994 and Wurdack 2006). Mohr sent the specimens on to Chapman, who described Cro¬

ton alabamensis in the second edition of his Flora of the Southern United States (Chapman 1887).

In the twenty years prior to Plant Life , Mohr published extensively on Alabama’s flora. He was especially

interested in new species (Mohr 1897, 1899), adventives on Mobile Bay’s ballast grounds (Mohr 1878), medici¬

nal plants (1890), and forest trees (Mohr 1882,1883,1896a).

Davenport, A history of Botany in Alabama

407

Fig. 5. Charles T. Mohr. (Courtesy of Erik Overbey/Mobile Public Library Collection, University of South Alabama Photographic Archives, Mobile, Alabama).

408

Journal of the Botanical Research Institute of Texas 9(2)

Another sidelight was Mohr’s relationship with Patrick H. Mell (1850-1918), botanist at the Agricultural

& Mechanical College of Alabama (now Auburn University). After corresponding with Mohr about some iden¬

tifications, Mell (1896a) published Part V of his own Flora of Alabama, covering the legume and rose families

(Parts I-IV never appeared). Mohr was shocked, and the American botanical community—which knew of

Mohr’s many years of work on Alabama’s plants—called for Mell’s censure (Anonymous 1896). Despite his

plea of innocence (Mell 1896b), Mell retired from Alabama botanical battles. Mohr (1896b), in a letter to E.A.

Smith, vowed to treat the entire Mell Affair with “silent contempt” and renewed his efforts to finish his opus

(for a light-hearted, football-themed depiction of this episode, see Davenport [2010a]).

Mohr was the first person to consider the overall picture of Alabama’s vegetation. In the introductory

pages of Plant Life, he combined climate factors and average temperatures (basic to Merriam’s Life Zones) to

divide the state into two nearly equal halves—the northern, temperate, Carolinian Area and the southern,

tropical, Louisianan Area (Fig. 6). He then subdivided these areas based on soil, substrate, and other character¬

istics, describing the plant associations within each one.

Mohr’s comrades include Judge Thomas Minott Peters (1808-1888) of Moulton (Lawrence County) in

north-central Alabama. Trained as an attorney, Peters (Fig. 7) served as Chief Justice of Alabama’s Supreme

Court after the Civil War. Before that war, he pursued his avocation of botany, particularly lichens and fungi,

sharing much information with Moses A. Curtis and Henry William Ravenel (Haygood 1987). Gray (1853)

described the rare fern Trichomanes petersii from specimens collected by Peters in Hancock (now Winston)

County. Mohr (1901) was grateful enough to Peters for his contributions on nonvascular plants that he in¬

cluded a short biographical sketch of Peters in Plant Life. Significantly, the Alabama Edition of that book in¬

cludes only two portraits, those of Mohr and Peters (for more on Peters, see Davenport [2014]).

In Plant Life of Alabama, Mohr (1901) also mentioned the vital contributions of Lucien Marcus Under¬

wood (1853-1907) and Franklin Sumner Earle (1856-1929) to his knowledge (and lists) of the state’s fungi,

lichens, and bryophytes. Underwood, a colleague of Mell at the Agricultural & Mechanical College, was also

one of the first professional botanists to visit the Havana Glen (Hale County) hybrid Asplenium site (see Under¬

wood 1896). Underwood left Auburn for Columbia University and was replaced by Earle, who contributed

greatly to Plant Life in the areas of fungi and lichens (Hansen 2003a, 2003b). One year after that publication,

Earle (1902) published a flora of the Alabama Piedmont.

1900-1960

The two decades surrounding the new century featured “incursions” by several botanists from the Biltmore

Herbarium. This herbarium, established in Asheville, North Carolina, sent out teams to comb the Appalachian

Mountains and their environs for new and unusual species (for a history of the Biltmore Herbarium and its

activities and accomplishments, see Anderson [2007]).

The contributions of this group to Alabama botany are numerous, especially in descriptions of new spe¬

cies of forest trees—although most of them are now sunk in synonymy (Davenport 2015). Chauncey D. Beadle

(1866-1950) was particularly prolific for Alabama trees, giving us Quercus boyntonii (Beadle 1901b) and many

“new” hawthorns (Beadle 1901a, 1902a) and cherries (Beadle 1902b). Beadle and Boynton (1901) also de¬

scribed the rare composite Marshallia mohrii, naming it for Charles Mohr, who moved to Asheville to consult

the Biltmore Herbarium and complete work on Plant Life of Alabama. (Sadly, Mohr died two weeks before his

book was published; hence, he is buried in Asheville rather than in his long-time home, Mobile.) Thomas Grant

Harbison (1862-1936) contributed Lrillium stamineum (Harbison 1901) and T. decumbens (Harbison 1902b),

plus general descriptions of the Alabama flora (Harbison 1902a).

Roland McMillan Harper (1878-1966) arrived in Tuscaloosa during late 1905 to begin a seven-decade-

long association with the Geological Survey of Alabama. He commemorated the event, in typical Harper fash¬

ion, by taking a long stroll or “ramble” and noting the plant associations that he saw (Harper 1906).

A Northerner by birth, Harper (Fig. 8) moved south with his school superintendent father and family,

graduating from the University of Georgia before pursuing his doctorate at Columbia University. For his dis¬

sertation, Harper studied the Altamaha Grit region of his adopted state, confirming the connection between

Davenport, A history of Botany in Alabama

409

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Fig. 6. Map of the Floral Areas of Alabama from frontispiece of Mohr's Plant Life of Alabama (Alabama Edition)

410

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 7. Thomas Minott Peters. (Courtesy of the Alabama Supreme Court and State Law Library, Montgomery, Alabama).

Davenport, A history of Botany in Alabama

411

Fig. 8. Roland McMillan Harper. (Courtesy of the Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, Pennsylvania).

412

Journal of the Botanical Research Institute of Texas 9(2)

geology and plants—a theme that he would pursue throughout his life. He was then hired by E.A. Smith “to

continue the work of the late Charles Mohr” on the economic plants of Alabama (Harper 1967). Except for short

stints in Georgia and Florida, Harper remained connected to the Geological Survey of Alabama until his death.

During his long and storied career, Harper published over 600 individual papers. (His life and works have

been summarized by Davenport and Hubbs [1995].) While many of these are impassioned letters-to-the-editor

and statistical “proofs” of societal trends, the rest are legitimately scientific. (An independent spirit, Harper

never published with a co-author.) Harper’s main botanical themes were “plant sociology” or community com¬

position (which he often recorded from train car windows), descriptions of new species (which he often de¬

ferred to others), and the need for fire “un”suppression to maintain Southeastern forests.

Prodigious in memory and tireless in ambulation (since he never learned to drive an automobile), Harper

pursued the most distant and rarest of plants. He produced many works dealing with Alabama’s plant com¬

munities (Harper 1914, 1937, 1939) and newly discovered species, such as Hexastylis speciosa (Harper 1936).

His largest and most Mohr-like works are a series of monographs for the Geological Survey on economic plants,

forest resources, and weeds (Harper 1913,1928,1942,1943a, 1944).

In Forests of Alabama, Harper (1943a) greatly revised Mohr’s 1901 system of plant associations, placing

his emphasis on geological formations rather than on climate. He concluded that “a map of [Alabama’s] forest

regions does not differ much in its broader features from a geological map.” Harper’s forest regions and sub-

regions (Fig. 9), based primarily on geology and soils, are nearly identical to those recently designated by

Griffith et al. (2001) and described above.

There’s a famous story about Harper and his cross-country effort to meet up with Professor de Vries at the

type locality of Oenothera grandiflora, in southern Alabama. De Vries was on a cross-country tour to Houston,

Texas, to deliver an honorary address at Rice University. (This celebratory trip was similar to that of Lyell sev¬

enty years before.) He was feted wherever he went, including Tuscaloosa on 21 September 1912, where he and

H.H. Bartlett of the U.S. Department of Agriculture were met at the railroad station by State Geologist Smith

and by Harper. The next day, the first three gentlemen set out by train and steamboat for Dixie Landing on the

Alabama River in Baldwin County, where Bartram had discovered O. grandiflora in 1775. Harper preferred to

rough it, and he and a colleague set out by train and on foot, camping out in the rain. They arrived in time to

meet the De Vries party retracing Bartram’s voyage up the Mobile Delta. Oddly, Harper—who recorded most of

the details of his adult life in a series of diaries—wrote nothing about meeting the world’s most famous geneti¬

cist (Shores 2008; Davenport 2011).

Harper was far friendlier toward the Benedictine monk Wolfgang Wolf (1872-1950; Fig. 10) of Saint Ber¬

nard Abbey in Cullman; in fact, in her biography of Harper, Shores (2008) devoted an entire chapter to these

“Kindred Spirits.” The pair met in 1927, shortly after the death of E.A. Smith. By that time, Wolf (1918) had al¬

ready published the hybrid oak Quercus bernardiensis (Q. montana x Q. stellata); his final publication (Wolf

1945) was of another oak hybrid, Q. capesii (Q. nigra x Q. phellos). More significantly, he described Talinum

mengesii (Wolf 1920), naming it after his abbot at Saint Bernard (see Plaisance 1958), and later T. appalachianum

(Wolf 1939). He also created the genus Cryptophila (Monotropaceae), which he separated from Monotropsis

(Wolf 1922).

In many ways, Wolf and Harper collaborated on studies of Erythronium in the 1930s, trading information

and specimens (Shores 2008). But rather than a joint paper, the two published separate ones (Harper 1941;

Wolf 1941) in back-to-back issues of Castanea. It was Harper (1951) who announced Brother Wolfgang’s death

to the botanical community.

SWAMP, PRAIRIE, HEMLOCK, AND GLADE STUDIES

The Tennessee Valley Authority (TVA) was established in 1933. During the 1930s and 1940s, several Alabama

ecological studies were conducted by individuals who were either then or formerly employed by TVA. In keep¬

ing with TVA interests, these studies (Penfound & Hall 1939; Hall & Penfound 1943; Penfound et al. 1945)

dealt largely with swamp and mosquito issues. In addition, Isely (1946) published a treatment of aquatic plants

found in TVA reservoirs, later updated by Dennis et al. (1977).

Davenport, A history of Botany in Alabama

413

Fig. 9. Regional Map of Alabama, based largely on geology, from Harper's Forests of Alabama.

414

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 10. Wolfgang Wolf. (Courtesy of Auburn University Special Collections and Archives, Auburn, Alabama).

Davenport, A history of Botany in Alabama

415

As shown above, travelers have long been fascinated by the Blackland Prairie or Black Belt of central Ala¬

bama; and one early resident (McGuire 1834) demonstrated solid understanding of its origin. Since the 1920s,

many studies have built on those initial observations, including ones by Harper (1920), Cocks (1925), Jones

and Patton (1966), Rankin and Davis (1971), Schuster and McDaniel (1973), and Barone and Hill (2007).

A second fascination has been the presence of Canadian hemlock in Alabama’s Southwestern Appala¬

chian Ecoregion. Hemlock studies include those by Harper (1943b), Segars et al. (1951), and Hardin and Lewis

(1980).

Working out of Vanderbilt University and the University of Kentucky, Jerry and Carol (Caudle) Baskin

published extensively on cedar glades and their endemics, including several from Alabama (Baskin & Baskin

1976,1984, 1986a, 1986b; Baskin & Caudle 1967; Baskin et al. 1995; Webb et al. 1992). A later paper (Webb et

al. 1997) listed the species known from the glades and barrens of northwestern Alabama.

Another limestone-based endemic brst noted in northwest Alabama, Jamesianthus alabamensis, was de¬

scribed by S.F. Blake and E.E. Sherff (Sherff 1940) from specimens supplied by Harper. For details of its discov¬

ery and naming, see Harper (1943b), Baldwin (1969), and Dennis (1982).

HALE COUNTY AND OTHER FERN STUDIES

No more famous botanical site exists in west-central Alabama than Havana Glen in Hale County. The site brst

came to light thanks to the efforts of Julia S. Tutwiler (1841-1916; Fig. 11). In 1873, Tutwiler—also noted as a

prison reformer, founding president of the Livingston Normal College (now the University of West Alabama),

and writer of the state anthem—discovered an odd fern in the Glen and recognized it as a hybrid Asplenium (for

more on this story, see Davenport [2007]). The site was later visited and described by Underwood (1896) and

Wherry and Trudell (1930). Because chromosome studies by Wagner (1954) revealed that the hybrid is a via¬

ble, self-reproducing tetraploid, the plant was recently re-named Asplenium tutwilerae (Keener & Davenport

2007).

Summary works on Alabama’s ferns came much later. John W. Short studied the state’s fern bora for his

Master’s thesis at Auburn University (Short 1978) and published several related papers (Short & Freeman

1978a, 1978b; Short 1979). Daniel D. Spaulding joined forces with several other authors to cover the northeast¬

ern portion of the state (Spaulding et al. 2000a, 2000b, 2001a, 2001b, 2001c). Alvin R. Diamond, Jr. and Mi¬

chael Woods did the same for the southeastern portion (Diamond & Woods 2007; Woods & Diamond 2008).

Very recently, Short and Spaulding (2012) produced the debnitive fern guide for Alabama.

BRYOPHYTE AND LICHEN STUDIES

In order to provide a nearly complete boral treatment, Mohr (1901) included non-vascular plants in his Plant

Life of Alabama. Since that time, few Alabama bryophyte papers have been published. Harvill (1950) described

Diphyscium cumberlandianum from King Cove in southwestern Lawrence County; he also supplied a list of

moss species (including new state records) found in that same “remarkable cove” (Harvill 1951). Wilkes (1965)

published a checklist of Alabama mosses; Bowers et al. (1989a, 1989b), Davison and Schotz (1998), and Dia¬

mond et al. (1999) have added many species to that list.

Mohr (1901) also included lichens in his bora, thanks mainly to the efforts of Auburn University’s F.S.

Earle. A century later, Hansen (2003b) updated that catalog and provided a detailed list of lichen collectors and

publications.

1950-2011

The last half of the twentieth century was dominated by boristic studies in Alabama. This emphasis on boris-

tics was part of a state-wide resurgence in natural history led by key individuals at state universities, plus one

very determined amateur.

That amateur was Blanche Evans Dean (1892-1974; Fig. 12). A native of Goodwater (Clay County), Dean

taught science courses at Woodlawn High School in Birmingham (for details of Dean’s life, see Christenson

and Davenport [1997]). But her main love was out-of-doors, specihcally nature study. In 1951 she began a series

416

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 11 . Julia S. Tutwiler. (Courtesy of Julia Tutwiler Library, University of West Alabama, Livingston, Alabama).

Davenport, A history of Botany in Alabama

417

of Outdoor Nature Camps, which morphed into the Audubon Mountain Workshops held at Mentone (DeKalb

County) each spring (for a history of the camps and workshops, see Holliman [1984]). She consolidated her

knowledge of Alabama’s plant life into books on trees, ferns, and wildflowers (Dean 1961, 1969; Dean et al.

1973).

At the University of Alabama, a resurgence of nature study was led by Ralph L. Chermock (1918-1977)

following World War II (see Davenport 2011). An American-born nephew of the Austrian geneticist Erich von

Tchermak (as in Tchermak, Correns and De Vries, the re-discoverers of Mendelism), Chermock mainly

mentored zoology students. (The most famous Chermockian of all is Harvard University’s E.O. Wilson; see

Wilson [1994] for his reminiscences.) As part of this natural history emphasis, Stallard (1950) produced a flo-

ristic study of the Fort Morgan Peninsula under the direction of Chermock’s colleague, A.M. Harvill, Jr.

(1916-2008), who has been mentioned previously for his bryological contributions.

In the 1960s, Joab L. Thomas (1933-2014; Fig. 13) and his students engaged in studies of Alabama endem¬

ics, like Croton alabamensis (Farmer 1962; Farmer & Thomas 1969) and Neviusia alabamensis (Thomas &

Deramus 1964). They also characterized the Black Belt flora (Maginness 1967; Naugle 1967) and that of Tangle-

wood Field Station (Williams 1967), Holt Fock and Dam (Brooks 1969), and Dauphin Island (Deramus 1970).

Thomas left Alabama to become Chancellor of North Carolina State University, later returning to the

University of Alabama as its president. His botanical successor, Robert R. Haynes, conducted a series of studies

on Alabama’s aquatic vascular plants (Haynes 1980; Davenport & Haynes 1981; Wiersema & Haynes 1983;

Haynes & Jacono 2000). In addition, Wiersema (1979) examined the distribution of the Nymphaeaceae in the

state; Hendryx (1996) focused on the aquatic and wetland plants of the Oakmulgee Division of Talladega Na¬

tional Forest; and Keener (1999) studied the flora of Blount County.

With the 1968 arrival of John D. Freeman (1941-1997; Fig. 14), botanical interest at Auburn University

was significantly rekindled (Hansen 2003a). Under the direction of Freeman and others, students completed

many floras and ecological studies, including Reed Brake Research Natural Area (Beckett 1980; Beckett &

Golden 1982), the Red Hills (Diamond 1987), Conecuh County (Diamond & Freeman 1993), Black Belt river

bluffs (Gunn 1985), the Tombigbee River (Crouch 1997; Crouch & Golden 1997), montane longleaf pine com¬

munities (Maceina 1997; Maceina et al. 2000), Horseshoe Bend National Military Park (Petranka et al. 1979),

the Piedmont Plateau (Rutland 1977), the Cahaba River (Sessler 1978), and Alabama’s trees and shrubs (Young-

hance & Freeman 1996).

Botanists at Auburn University and the University of Alabama collaborated on a listing of the state’s en¬

dangered, threatened, and special concern plants (Freeman et al. 1979). Such a list was an expansion of that

proposed by Thomas (1976).

Freeman’s Auburn colleague, Robert S. Boyd, has concentrated on plant ecological studies, especially

those involving rare species: Clematis socialis (Boyd & Hilton 1994), Xyris tennesseensis (Boyd et al. 2011), and

Rudbeckia auriculata (Diamond et al. (2006).

During the 1990s, the rapidly growing plant collections at the University of Alabama and Auburn Univer¬

sity were databased; the data were then combined into a single checklist (Morton et al. 2002). Many of the

“gaps” in that list were Filed by adding specimens from the universities listed below.

Thanks to his dissertation work at the University of North Carolina, Ross C. Clark contributed much to

our understanding of Alabama’s woody plants (Clark 1967, 1969, 1971). Another UNC doctoral student, R.

David Whetstone, concentrated on the Fora of the Cumberland Plateau of our state (Whetstone 1981).

Whetstone then resumed residence at his alma mater, Jacksonville State University. His many students

completed county-wide or local Foras, including the following: Talladega Ranger District (Ballard 1995), Jef¬

ferson County (Barber 1986), Jackson County (Brodeur 1999), Cheaha State Park (Bussey 1983), kittle River

Canyon (Dickson 1992), Etowah County (Hodge-Spaulding 1997), Fimestone County (Hofmann 2000),

Horseblock Mountain (Hruska 1997), Dugger Mountain (Hutchinson 1998), St. Clair County (Jackson 2000),

Randolph County (Nixon 1989), Fake Guntersville State Park (Spaulding 1995, 1999a, 1999b), Madison

County (Threlkeld 1998), and Choccolocco Creek (Weninegar 2002).

At the University of South Alabama, Michel G. belong intensively studied the Fora of the Outer Coastal

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 12. Blanche Evans Dean. (Courtesy of the Alabama Women's Hall of Fame, Judson College, Marion, Alabama).

Davenport, A history of Botany in Alabama

419

mentation, Carnegie Mellon University, Pittsburgh, Pennsylvania).

Fig. 13. Joab L. Thomas. (Courtesy of the Hunt Institute for Botanical Docun

420

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 14. John D. Freeman. (Courtesy of the Freeman family. Auburn, Alabama).

Davenport, A history of Botany in Alabama

421

Plain. He published several works on the general flora and the rare plants of Baldwin and Mobile counties (Le-

long 1977,1988a, 1988b, 1991).

The work of Michael Woods and Alvin Diamond have put Troy University’s botany program securely on

the Alabama map. Woods has directed several legume studies (Woods 2008; Woods & Key 2009), while he,

Diamond, and their students have produced many Boras: Pike County (Diamond 2003), Crenshaw County

(Diamond 2011), Pike County Pocosin Nature Preserve (Diamond et al. 2002), Dale County (Dransfield &

Woods 2004), Coffee County (Martin 2001; Martin et al. 2002), Ech Lake (Rundell & Woods 2001), Pike

County Lake (Woods & Reiss 1998), and Dale County Lake (Woods et al. 2000). In addition to their work on

ferns listed above, Woods and Diamond (2005) collaborated on a treatment of southeastern Alabama

gymnosperms.

Several other theses have been produced by students from out-of-state universities. These schools include

the Catholic University of America (Morgan 1942, Cullman County); the University of North Carolina (Bos¬

tick 1964, 1967; St. Clair County); and Mississippi State University (McDearman 1976, Lawrence County;

Smith 1996, Marion County).

Alabama’s Lorever Wild Land Trust was established by constitutional amendment in 1992; since that

time, over 200,000 acres of land—including some of the state’s most pristine—have been purchased for the

public trust. T. Wayne Barger, botanist with the State Lands Division (which oversees the program), has di¬

rected several studies of these vital Lorever Wild properties, including Indian Mountain (Barger & Holt 2010)

and Coon Creek (Barger & Tenaglia 2008).

The end of the twentieth century also marked a return to herbal studies. John K. Crellin and Jane Philpott

documented the herbal treatments prescribed by Alabama’s leading modern folk practitioner, A. L. “Tommie”

Bass (1908-1996) of Leesburg (Cherokee County). Throughout his long career, Bass combined frontier, Native

American, and African-American traditions. The two-volume set (Crellin & Philpott 1989, 1990) very accu¬

rately records his extensive knowledge, country charm, and wit.

jim allison’s lost world

In the early 1990s, Georgia botanist James R. Allison discovered a “lost world” in Bibb County, Alabama, with

several rare and new taxa inhabiting the severe, exposed landscapes of Ketona dolomite there (Allison & Ste¬

vens 2001). The new taxa included Castilleja kraliana, Coreopsis grandiflora var. inclinata, Dalea cahaba, Erig-

eron strigosus var. dolomiticola, Liatris oligocephala, Onosmodium decipiens, Silphium glutinosum, and Spigelia

gentianoides var. alabamensis (now elevated to species status; see Weakley et al. 2011). In addition, seven state

records were discovered, including Solarium pumilum, which was presumed extinct.

These dolomitic glades are rightly considered “one of the most significant reservoirs of botanical diversity

in the eastern United States” (Allison & Stevens 2001). One of the largest is now preserved as the Kathy Stiles

Preeland Bibb County Glades by the Nature Conservancy.

ROBERT KRAL

But it is the work of Robert Krai that dominates Alabama botany today. Based at Vanderbilt University and

funded by the National Science Poundation, Krai (Pig. 15) undertook intensive study of the flora of the South¬

east, starting in the 1960s. And thanks to several travel grants from the USDA Porest Service, he produced the

massive, two-volume, 1305-page A Report on Some Rare, Threatened, or Endangered Forest-Related Vascular

Plants of the South (Krai 1983a, b). Krai’s intention has always been to produce a flora of Middle Tennessee (the

section directly north of Alabama) and Alabama itself. Toward this end, he published a series of papers on the

rarities that he encountered (Krai 1973,1976a, 1981).

Krai has always taken on the toughest taxonomic assignments, including Xyris (Krai 1966a); Eriocaula-

ceae (1966b); Rhexia (Krai & Bostick 1969); Abildgaardia, Bulbostylis, and Fimbristylis (Krai 1971); and Fuirena

(Krai 1978a). He has produced many treatments of the above groups for the Flora of North America and various

floras of Central and South America. And he has described many new species, including the following from

Alabama: Delphinium alabamicum (Krai 1976b), Xyris tennesseensis (Krai 1978b), Sagittaria secundifolia (Krai

422

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 15. Robert Krai. (Courtesy of the Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, Pennsylvania. Photograph

by Walter Henricks Hodge.).

Davenport, A history of Botany in Alabama

423

1982a), Clematis socialis (Krai 1982b), C. morefieldii (Krai 1987), Fimbristylis brevivaginata (Krai 1992), Ble-

philia subnuda (Simmers & Krai 1992), Solidago arenicola (Keener & Krai 2003), and Xyris spathifolia (Krai &

Moffett 2009).

Krai’s efforts to examine and understand Alabama’s plant life will never be matched. The most prolific

botanist in the history of the state, he has collected over 100,000 specimens—all with his iconic “R. Krai” sig¬

nature. Since 1997, this unsurpassed collection has been housed at BRIT.

Younger botanists—most of whom are mentioned above—met with Krai quarterly from 2002 to 2008 to

create a checklist of Alabama plants (Krai et al. 2011). In January 2012, the on-line Alabama Plant Atlas (http://

floraofalabama.org), based on that checklist, was dedicated, housed at the University of West Alabama under

the directorship of Brian R. Keener. The ultimate goal is to produce the “complete” written flora that Bob Krai

has long envisioned. With points of emphasis now established at Troy, Auburn, Mobile, Montgomery, Livings¬

ton, Jacksonville, Anniston, Birmingham, Tuscaloosa, and Florence, that task is sure to be accomplished.

Thanks, Bob, for carrying us this far.

ACKNOWLEDGMENTS

This paper has been forty years in the making. It started as a card hie of references to early works. Guy Hubbs

“fed” me everything he came across describing early Alabama landscapes, while Tamara Haygood sent her

extensive holdings on Southern botanists.

Injanuary 2012, an outline was presented at the dedication ceremony for the Alabama Plant Atlas. I thank

Brian Keener for forcing me to finally pull my notes together.

I also thank Liz Wells of Samford’s Special Collections for locating obscure genealogy records. Robert

Steen helped me understand the history of LaGrange College. And Wayne Barger directed me to Mike Palmer

and his great list of Boras.

Pulling the portraits together proved to be an arduous task. Dustin Williams and his staff at the Hunt In¬

stitute for Botanical Documentation were truly amazing. I also thank Mike Howell, Bill Mathews, Curtis Han¬

sen, Larry Dorr, Lisa Dodd, Rachel Cohen, Suzanne Oberholster, Alvin Diamond, and Ruth Freeman—wheth¬

er those efforts succeeded or not.

I apologize to any Alabama botanists who feel slighted or left out of this treatment. Please realize that re¬

cent works—those published after the 2011 checklist—have been purposely left out, as have most short papers

on additions to the state Bora. An updated (and, hopefully, complete) list of works is maintained by the Ala¬

bama Plant Atlas. If your work is not cited there, we will make sure it is!

I also apologize to anyone who objects to my use of “Cro-Magnon” in the title. I realize that the term is

both outdated and inaccurate, but I needed its alliterative effect—and to indicate the continuing evolution of

botanical knowledge. It’s a “stretch” that Lloyd Shinners, the founder of this esteemed journal, would

appreciate.

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thesis, University of Alabama, Tuscaloosa, Alabama, U.S.A.

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432

Journal of the Botanical Research Institute of Texas 9(2)

BOOK NOTICE

Kelly M. Hines, Alexander Krings, and Jon M. Stucky. 2015. Guide to the Vascular Flora of Howell Woods,

Johnston County, North Carolina, U.S.A. Sida, Botanical Miscellany 43. (ISSN: 0833-1475; ISBN: 978-

1-889878-47-8, pbk). Botanical Research Institute of Texas Press, Fort Worth, Texas 76107, U.S.A.

(Orders: shop.brit.org, orders@brit.org, 1-817-332-4441). Price to be determined, 276 pp., 7" x 10".

From the Publisher —Howell Woods Environmental Learning

Center (Johnston County, North Carolina; “Howell Woods”) is “the

most significant terrestrial natural area in the county” as designated

by the NC Natural Heritage Program. Comprising 1155 ha, the site

is one of the largest tracts of intact forest remaining in the county.

Howell Woods is home to ten natural plant communities and nu¬

merous rare plant species. Guide to the Vascular Flora of Howell

Woods provides a checklist of the flora compiled from the first au¬

thor’s collections, historic collections, and reports of species from

the site. The illustrated guide is based on the checklist of over 580

species in 123 families and includes 108 new county records. Keys

are provided to all vouchered or reported species and genera.

Habitat, exotic status, flowering and fruiting phenology, abun¬

dance, and synonymy are incorporated. In addition, relevant vouch¬

er information is provided. This is first in a series of Illustrated

Floras of North Carolina Project from the North Carolina State

University Vascular Plant Herbarium (NCSC), designed to be useful

to both botanical specialists and a more general audience.

Kelly Thames (nee Hines) is a wetland scientist at an environmental consulting and habitat restoration firm

continuing her interests working with the public, plants, and the natural world.

Alexander Krings, Ph.D. is Assistant Professor of Plant Biology and Director of the Vascular Plant Herbarium

at North Carolina State University. The central focus of his work is the discovery, taxonomic circumscription,

and monography of plants and the subsequent development of applied resources that facilitate plant identifica¬

tion and related communication. Dr. Krings is also author of the Manual of the Vascular Flora of Nags Head

Woods, Outer Banks, North Carolina (2010) published by the New York Botanical Garden Press.

Jon M. Stucky, Ph.D. is Professor Emeritus of Plant Biology at North Carolina State University. He has taught

undergraduate and graduate level held botany courses for over thirty years, including Systematic Botany,

Grasses, Sedges & Rushes, Wetland Flora, and Local Flora. Dr. Stucky’s research focused on rare plant species

ecology and conservation.

J.Bot. Res. Inst. Texas 9(2): 432.2015

BIRCH ( BETULA , BETULACEAE) BARK HORNS

AND SIMILAR INSTRUMENTS IN NORWAY

Torbjorn Aim

Troms0 Museum

University ofTromso

P . O.Box 6050

Langnes , N-9037 Tromso , NORWAY

torbjorn . alm @ uit.no

ABSTRACT

Wooden horns wrapped in coiled birch bark (Norwegian: lur) have a long history in Norway, dating back at least to the 7th century AD. By

the Vikings, they were used for various signalling purposes, e.g. during battles. More recent uses are generally peaceful—to celebrate the

opening of fairs, announce bishop visitations etc. In the 19th century, playing such instruments was as a popular past-time when herding

cattle, and, if needed, to scare off predators (bears, wolves, etc.) or signal for help. Common throughout the southern part of Norway, such

instruments gained symbolic importance during the 19 th century national revival.

RESUMEN

Los cuernos de madera envueltos en corteza de abedul enrollada (en noruego: lur) tienen una larga historia en Noruega, al menos hasta el

siglo VII AD. Los Vikingos, los usaron con varios propositos de senalizacion, ej. durante las batallas. Los usos mas recientes son general-

mente pacificos—para celebrar el comienzo de ferias, anunciar visitas de obispos etc. En el siglo XIX, tocar tales instrumentos fue un pasa-

tiempo popular cuando se cuidaba el ganado, y, si era necesario, asustar a los predadores (osos, lobos, etc.) o como serial de auxilio. Comunes

por toda la parte sur de Noruega, tales instrumentos ganaron una importancia simbolica durante el renacimiento nacional del siglo XIX.

INTRODUCTION

Plants are versatile raw materials, and wood and bark can be used to produce a vast range of utensils. McCune

and Prendergast (2002) drew attention to the use of birch ( Betula ) for making musical instruments or simple

wooden horns in Europe, commenting on examples from Finland, Norway and Switzerland, based on speci¬

mens in Kew’s Economic Botany Collection. No primary data from Norway were included, and a rather ecclec-

tic selection of other sources was used—but then, very little information is available in English. This paper

aims at a comprehensive review of such wooden, bark-covered horns (Norwegian: lur) in Norway—history,

materials, traditions, and uses.

For a botanist living in the far north of Norway, a striking aspect of the Norwegian birch bark horn tradi¬

tion is it geographical distribution. Such instruments are well known and deeply rooted in folk tradition of the

south, and rare in the north—despite the fact that ideas, traditons and innovations are easilly transmitted

along Norway’s long coast, always a major communication route. The difference is certainly not due to topo¬

graphy, which is equally rugged in the north, suggesting a similar need for signalling instruments. This aspect

has been overlooked in available literature, and trying to explain it is a second goal of this article.

Etymology

The root meaning of the Norwegian word lur (old Norse ludr) is a hollowed-out piece of wood, not necessarily

a musical instrument (Christiansen 1952a, 1952b; Holtsmark 1946). The Norse god Heimdall possessed a fa¬

mous example, Gjallarhorn. According to Norse mythology, and the poems of the younger Edda, “he has a lur

called Gjallarhorn and his lur playing can be heard everywhere.”

A brief definition of such instruments is found in a late 17 th century manuscript by Thomas Bloch, com¬

prising a dictionary of terms used in Fyresdal, Telemark: “Luu, Instrumentum est, ut tuba, factum e ligna ex-

cavato, betulse cortice circumvolutum, qvo pastores sonitum dant ad arcendos a pecoribus ursos et lupos,—is

an instrument, made as a trumpet of hollowed out wood, wrapped in birch bark (Fig. 1), which the shepherds

J. Bot. Res. Inst. Texas 9(2): 433 - 448.2015

434

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 1. A typical Norwegian lur is covered by a coil of birch bark, serving both a decorative purpose and to keep the two halves of the interior wooden

kernel together while blocking leakages.

blow to scarce bears and wolves away from the cattle” (Bloch 1956:23). Knud Leem’s Norwegian dictionary,

compiled in the 1740s, offers a similar explanation: “A luur: an almost one fathom [1.73 m] long instrument

made of wood (Fig. 2), and covered with birch bark, which the peasant girls use to blow, mostly in summer,

when they are herding the cattle, partly for amusement, but also to scare off the bear” (cited from Hannaas

1923:120). An anonymous, late 18 th century dictionary manuscript from Surnadal in Trondelag provides simi¬

lar information: [an instrument] “(...) for blowing, so that the sound can be heard loudly among the surround¬

ing hills, and the bears in particular are scared and driven away from the cattle” (Hagland 1986:47).

The term lur has also been applied to a variety of other objects, e.g., hollow plant stems (Christiansen

1952a, 1952b: Myrvang 2010), which were frequently used as simple pipes, e.g., by children. In toponyms, it

may refer to the occurrence of plant species with this characteristic, e.g. Angelica sylvestris L. (Myrvang

2010:186).

History

As noted by McCune and Prendergast (2002), a wooden horn or lur was found in the Oseberg Viking burial

ship, dendrochronologically dated to 834 AD. The specimen was 107 cm long and had a diameter of 4 cm at the

broad end (Grieg 1928; for an illustration, see Christensen et al. 1994:134 or Vollsnes 2001:49). In a recent ar¬

chaeological review of the Oseberg find (Christensen et al. 1994), the specimen is considered somewhat enig¬

matic, as a “scepter, blowing lur or ???”, but the description (Grieg 1928:270-271) fits a typical lur —a meter-

long, wooden tube, tapering towards one end, consisting of two halves, with a somewhat irregular interior

hollow—and there is hardly any reason for interpreting it otherwise, even though it had not been covered with

Aim, Birch bark horns and similar instruments

435

Fig. 2. This medium-sized instrument (length: 1.53 m) is easily carried, weighing a mere 520 g. Author's collection. Photograph by Mari Karlstad.

bark. Instead, the two halves were kept together by bands, attached in five incised rings. This technique is typi¬

cal of Viking-age or older specimens (Vollsnes 2001:49) but was still used in parts of Norway as long as such

instruments were commonly made, i.e., throughout the 19th century (Gundersen 1994:19).

An even older specimen from Norway was found in the Kvalsund ship burial, unearthed at Heroy in west¬

ern Norway and dated to about 600 AD. It contained a short lur, 72.5 cm long, with a diameter at the widest end

of 4.5 cm (Shetelig & Johannessen 1929:70). The short length may suggest that it had not been used as a musical

instrument but perhaps as a kind of megaphone (Shetelig & Johannessen 1929:39). Similar wooden mega¬

phones were used by fishermen in northern Norway until perhaps a hundred years ago (Christiansen

1952b: 108).

Making and materials

Several authors provide notes on how to make a lur. The most frequently used technique was to make two sepa¬

rate halves from a single piece of wood. It should be noted that Norwegian wooden horns are not exclusively

made from birch wood and birch bark. This choice of material may have been most typical for the coastal dis¬

tricts, where Betula species are the predominant forest trees. Indeed, according to the original publication

(Grieg 1928:270), the Oseberg lur noted by McCune and Prendergast (2002) was made of beech (Fagus sylvatica

L.); more recent authors (Vollsnes 2001:49; Gundersen 1994:19; Sevag 1966:10) state that the material used was

yew (Taxus baccata L.), but no reason for doing so is given, and no revision of the original material has been

carried out (Arne Emil Christensen, pers. comm.).

In many areas, not least inland, conifers were preferred as raw material for the interior wooden tube. In

436

Journal of the Botanical Research Institute of Texas 9(2)

southeastern Norway, spruce (Picea obits (L.) Karst.) was frequently used. Trees that had grown slowly were

preferred. In the Hallingdal area, pine ( Pinus sylvestris L.) was considered the best raw material (Gundersen

1994:22-23), not least because suitable working material could be found as preserved, subfossil stems in the

mires; such material was considered less “fat” than fresh pines. Living pines could also be used; if so, they were

split lengthwise from the bottom end. A single log could produce numerous instruments (Gundersen 1994:23).

The material chosen for the tube had to be of a certain minimum size, as the central part of a stem was

avoided (Gundersen 1994:23). The halves were hollowed out, by knife or other tools. Using a knife would usu¬

ally produce a more or less square interior hollow (Gundersen 1994:23). The two halves were then re-assem¬

bled. Some would glue them together, but others considered this to give instruments of inferior quality (Gun¬

dersen 1994:23). This is why they are wrapped in bark; a long, coiling strip of birch ( Betula ) bark served to keep

the two halves together. Making the lur air-proof was essential, and the bark would also block potential holes

or leakages. In addition, it obviously had a decorative function.

Instruments could also be made from bark only, coiling it up into a small cone, usually some 40 to 60 cm

long (Nupen 1992:16). In addition to birch, bark of rowan ( Sorbus aucuparia L.), alder ( Alnus spp.), and the

larger Salix species could be used (Hooker 1837; Christiansen 1952b; Hoeg 1974; Nupen 1992; Dahl 2005:303).

Even these simple instruments could survive for years but may have served mostly for children’s amusement

(Hooker 1837).

Birch bark for the covering coil of a conventional lur was cut from the trees in spring or early summer

(Gundersen 1994:24; Dahl 2005:303). Straight, tail-grown trees were preferred, as only these would yield bark

of suitable quality—preferably white and with few scars (see discussion). With a knife, a spiral was incised,

starting at suitable height, and the outer layer of the bark peeled off, so that one exposed the green, interior part

(Gundersen 1994:24). The white outer skin was then removed, leaving the detached, yellow-brown part ready

for use or almost so; the margins might need some trimming. The bark was attached to the lur while still fresh.

Winding it onto the lur could start from either end. The start would lock itself. At the far end, the bark strip

could be locked by inserting it under the coil. Small wooden plugs were often used as an additional way of se¬

curing the end. An experienced cutter could make very long coils; Hatledal (1997:6) mentions specimens up to

6 and 7 m long. He also noted that the trees survived, forming what he terms korpebark (“raven bark”) at the

cut.

A detailed description of how to make a lur is provided by Ola Hola, based on traditions in More og Roms-

dal County, western Norway (Hola 2000:66-67). In this case, pines were preferred for the wooden core. The

trees were cut before Christmas, while the moon was waning. They were split lengthwise, placed beneath a

roof, and left drying for one year. From this material, a tube was prepared, some 80 to 90 cm long, with an in¬

terior diameter of about 6 mm at the thinnest end, gradually increasing to 14 mm towards the other end of the

tube, until 14 cm was left. From this point, the diameter increased rapidly towards the outermost part ( sopen ),

which had a diameter of about 10 cm. The wooden frame was 2.5 to 3 mm thick, except at both ends, where it

was thicker. The lur was now covered with birch bark. This could only be done in summer. A fine, even-sur¬

faced, young birch was chosen, and the bark cut spiral-wise downwards with a sharp knife. Each incision

should be about 18 mm apart, so that the bark could be sliced off as long bands. The outermost skin was peeled

off. The initial end of the strip was tapered so that it was only half as wide as the rest. Starting at the mouth¬

piece, the bark was wound onto the tube, with an overlap for each layer of about 3 mm. The bark should be at¬

tached tightly, so that it was stretched a little; this would press the two halves of the tube together. If the bark

needed mending, it was done in the same way as at the start of the tube. The end was attached with three small

wooden pins. Before the lur was used, clean water was poured through it. Instruments could also be made from

a single piece of wood. This would avoid the risk of holes or leakages, but the preparation would be much more

difficult.

Shape and dimensions

The typical shape is that of a long cone, simply because it produced the best sound. A cylindrical lur was a bad

instrument (Gundersen 1994:23). The diameter at the widest end could be some 85 to 100 mm (Gundersen

Aim, Birch bark horns and similar instruments

437

1994:2; Hola 2000:66). The mouth-end had a diameter of 10-12 mm or less (Fet 1991:23). Some horns were

simply cut at the mouth-end; others had a more elaborate mouth-piece. The latter could also be loose and made

of wood, horn, or bone.

The Kew example of a Norwegian lur, depicted by McCune and Prendergast (2002), is 158 cm long. This

is a medium-size instrument according to Norwegian standards. Both much shorter and much longer instru¬

ments were made. The shortest, known as stuttlur “short lur,” notatut etc., could be a mere 30 to 40 cm long

(Nupen 1992:15), while in other areas 60 cm was considered a minimum (Gundersen 1994:24). A typical lange-

lur “long lur” would be about 150 cm, i.e., similar to the Kew specimen. Much longer examples were sometimes

produced. The longest instruments made in the Hallingdal area exceeded 3 m (Gundersen 1994:24), but these

were probably rare exceptions.

The collections at Norsk Folkemuseum (Norwegian Folk Museum) in Oslo contain about twenty speci¬

mens; most are of the medium-sized langelur type, i.e., more or less similar to the Kew specimen; four are of the

stuttlur type. Two deviant, short specimens made of bark only are described as orelur, i.e., made of alder ( Alnus

sp.). Photographs of all these can be seen at museum’s home page (www.norskfolke.museum.no).

At Stange in Hedmark, Embret Msehlum was known for his fine lur playing. In the 1880’s, he made an

unusually large specimen, and it is unlikely that a larger lur was ever made:

“But this large lur, he made after returning home [from America], and he had worked on it for a

whole year. It was completed in 1886. He found a knotless spruce up in Prsestmark’n. He

cleaved it along the middle, and then he started to hollow it out, so that a thin pipe extended all

through it. Then, he wrapped the lur in pieces of birch bark and glued it together. It was very

well done.” (...) “He also made a stand for it, on which he placed the lur when he was playing.

The lur was enormous. It was 5.39 meters long. I believe it must have been the largest lur in the

world but its weight was no more than 3.6 kilos” (Engen 1991:103).

The motivation for making instruments of varying length was two-fold, practical and musical. A very long in¬

strument could not easily be carried around, e.g., when herding livestock. This was a task often carried out by

children, which again would require rather short and small instruments. Musical considerations would rather

motivate long instruments. The longer the instrument, the more different tones it could make (Gundersen

1994:24; Nupen 1992:15). In addition, the loud sound of a large specimen could be heard far away. The sound

produced would still vary a lot between individual instruments, and some were better than others. At least lo¬

cally, holes could be drilled in the side of the lur to provide further musical variation (Morch 1964:182).

Purpose

In the Norse-Icelandic sagas, the lur is more frequently mentioned than any other musical instrument (Volls-

nes 2001:48), though there is nothing to suggest that its use during the Viking age was for entertainment. Most

references point to a military use, as a signal during war and battles, e.g., to board ships, to rally at the king’s

standard, attack, or withdraw. The only old Norse law that explicitly mentions the lur is the hirdskra, a military

code for the royal guard (Sevag 1966:10). At least some Norwegian kings possessed a personal specimen,

konungs ludr, “the king’s lur.” The instrument of king Sverre (c. 1150-1202) even had a name, Andvake, and

seemingly produced a special sound that his soldiers could recognize and know that they were called for

(Vollsnes 2001:51). When Scottish mercenaries—hired by the Swedish king—tried to march through Gud-

brandsdalen in SE Norway towards Sweden in 1612 (see Michell 1886), a lur was used to signal their approach

to the log trap that killed many of them. It was launched “just as they paused to listen to Prillar-Guri, who stood

on top of the slope on the opposite side of [the river] Lagen and blew a lur” (Botten-Hansen 1853:63), or so the

legend says—whether true or not. Local tradition has been busy embellishing the peasants’ victory story with

additional details, e.g. that the Scots took revenge by introduction Cicuta virosa L. to the area—and to Norway;

both claims are at odds with reality (Aim 2015).

If need be, the instrument could also be used to signal the presence of thieves and other criminals (Nys-

tugun 1950:120-121). The lur was also used as a calling signal at more peaceful occasions, e.g., during assem-

438

Journal of the Botanical Research Institute of Texas 9(2)

blies of various kinds. The latter use declined by the 13th century, when the Christianization of Norway had

led to churchbells taking over the old calling function of the lur (Vollsnes 2001:54).

Among the many legends related to the Black Death in Norway, a favorite topic is the desolation met with

afterwards, frequently in terms of a single survivor in each isolated settlement. Their presence was ascertained

by the sight of smoking chimneys or, according to local lore, by signalling with wooden horns, e.g., at Rauland

in Vinje, Telemark (Berge 1940:123), and at Kvikne in Nord-Fron, Oppland (Klonteig 2007:73).

As noted by McCune and Prendergast (2002), wooden horns were frequently used as herding instru¬

ments, in particular in high summer when the herds were moved to separate summer farms (Hornemann

1808:404; Lie 1914:186; Bjerknes 1945:50; Holtvedt 1945:124; Svarteberg 1968:50; Perstolen 1970:90; Svare

1973:241; Sem 1983:11; Engen 1991:103). The horns could signal that the herd had been located or be used to

ward off predators, especially bears and wolves (Hagemann 1889:121; Slingsby 1904:87, 1966, 1998:58, 70;

Hannaas 1923:120; Frolich 1924:161; Sagen 1950:246; Holtvedt 1945:125, 1953:138; Hogasen 1949:139; Per¬

stolen 1969:21-22; Sandaker 1976:61; Lodoen 1989:64; Tomasgard 2004:41; Dahl 2003:185, 2005:303; Nupen

2003:60), or simply to relay signals (Rise 1933:66). A stuttlur “short lur” in the collections of Norsk Folkemu-

seum (depicted in Nupen 1992:17) is inscribed 1862, the year it was made. An accompanying note tells its ori¬

gin: “I also have a lur of birch which my father made in 1862 to scare off wolves and lynx when he was herding

the goats.” According to folk tradition, the bear was scared by the sound of a lur, whereas most claim that he

enjoyed the tunes of the prillarhorn, which was made from a goat, sheep, or cow horn (Skar 1911:96; Morch

1964:181; Perstolen 1970:94-95; Gundersen 1995:73). From the Oslo area, Holtvedt (1953:138) noted that

bears disliked the sound of the lur: “But the best thing to do was to blow the lur close to his ear; he did not like

the sound. He wanted it quiet.” In his childhood in Hornindal (Hordaland, western Norway), before emigrat¬

ing to the U.S.A., I.N. Lodoen used his lur to ward off an attacking bear while herding cattle and sheep:

“The cows had formed a circle and the sheep were tightly grouped. Calves and younger animals

were surrounded for protection. The situation was tense. A big, black bear stood close to me and

was about to catch his prey. What could I do, and what should I do in this critical situation? I

had no other weapon than my lur. It appeared a weak weapon against such a bear. But I started

blowing the lur. Did I blow hard? I must have done so, for I frightened the beast. He ran and ran

without halting, and disappeared up the mountain side at the other side at full speed. The battle

was won and the herd saved. It took some time for the cattle to calm down, and the sheep to

comprehend. After a while, I was able to drive the animals down to the summer farm, and all

was well. The sound of the lur had been heard, and people came running to help me” (Lodoen

1989:61-62).

At least once, the lur was put to more prosaic use in order to chase off a bear, as noted from Vefsn in Nordland,

North Norway:

“The bear took a sheep once grandma was herding. She ran after him and hit him with a long,

large lur she had. The bear was about to carry the sheep across a brook, but now he let the sheep

loose, and attacked grandma instead. He bit her in her thigh, and she had a large wound” (Svare

1973:240-241).

It was obviously better to chase off the bear with the loud sound of the lur. The sound’s ability to carry far is the

point of a humorous folk tale, describing a fabulously large cow:

“(...) and the distance between her horns was so large, that if there was a herder sitting on each

horn blowing a lur —some even said trumpet blowers—they could not hear each other” (Holt¬

vedt 1953:144).

In the 19th century, the sound of the lur was an integral part of the landscapes of southern Norway and en¬

countered almost everywhere (Soegaard 1868:100). It is frequently mentioned (e.g., in tourist travel accounts),

often in more or less romantic settings such as in an account of a visit to mountains of Telemark in 1834: “At

Aim, Birch bark horns and similar instruments

439

first, we passed over a wooden bridge, beneath which the rapids of Maneelven [the river Mana] raged, and so,

while the lurs were sounding up in the mountains, our alpine travel commenced” (Hammerich 1840:36). The

English mountaineer W.C. Slingsby encountered children with wooden horns at several occasions, e.g., at Hel-

lesylt in Stranda, western Norway, in 1875:

“Near Indre Haugen we met at different places children carrying a ‘lure’—a long wooden horn

or trumpet, seven to nine feet in length, formed by two split and hollowed pieces of wood put

together and bound firmly by birch bark, out of which material a bell mouth of five or six inches

diameter is also formed. The boys blew these ‘lurer’ most vigorously in order to frighten away

the bears which had just killed two cows at Haugen” (Slingsby 1904:87).

The instrument is also mentioned in early accounts of the folklore of Norway, e.g., in the legends recorded by

S.O. Wolff in Telemark:

“Countless and differing sounds almost numb the ear, among them the chiming of bells and the

tune of the Luur, which come from the rivulet, where some children are standing with their fish¬

ing rods” (Wolff 1828:74, reprinted in Wolff 2014:137).

“She blew her Langeluur [long lur] so that Veirmaalet [the dwarf’s speech, i.e., the echo] an¬

swered from all the krags, and you could easily tell, where Guro Dalen’s summer farm was situ¬

ated” (Wolff 1833:14, reprinted in Wolff 2014:82-83).

As a herding instrument, the lur was still frequently heard during the last half of the 19th century. It features

prominently among the sounds noted by Ernst Bjerknes in the Krodsherad area of SE Norway during a sum¬

mer visit in 1883:

“For a long time, we had heard the sounds of bells and the mooing sound of the cows from both

sides of the mountains; now the siren calls of the dairy maids were added, and the cows an¬

swered. It turned into a beautiful antiphony, sometimes accompanied by the lur or the roaring

sound of a prillarhorn. It all melted into a glorious symphony of captivating beauty in the still,

warm summer night” (Bjerknes 1945:50).

Tourists visiting the Hjartdal area of Telemark in 1899 heard similar sounds: “Day and night you hear the

sounds of lur , calling [for cows], and chiming bells” (Dahl 2005:264), and the sounds of nearby Fyresdal were

much the same: “There I heard the tunes of a langelur [long lur], and I turned towards the sound, and far away

below Raufjell I saw a fine herd grazing. The shepherd girl stood on a big stone with the lur in her hand, and a

small boy was sitting nearby on a tussock” (Lie 1914:186). Even in the vicinity of Oslo, the capital, the lur was

frequently heard:

“When they were at Oppkuven, they heard all the blowing of lurs on the summer farms at

Krokskaugen—Come home now! Come home now! they signalled to the cow[s] both from

Vakersetra and Heggelisetra and Gagnumsetra and other summer farms as well” (Holtvedt

1945:56).

A lur could be used for any kind of signal and was useful for communicating across the often steep terrains of

Norway (Perstolen 1970:91), not least in foggy weather. In some areas, dairy produce was shipped down the

steep mountain sides attached to wires, and a lur could signal that a new batch was on its way down (Nupen

1992:15). In western Norway, a short wooden horn or notatut was used to signal that shoals of herring had ar¬

rived. The sound of a lur could easily be heard on the other side of a fjord (Fet 1991:23; Nupen 1992). Others

used it to signal that a meal was ready (Holtvedt 1945:125). It was also useful if you got lost in the forest or

mountains (Holtvedt 1945:169-170). Bakke (1990:219) provided some further details from Frsena in More og

Romsdal, western Norway. Here, the instrument was used to signal meals or the end of the working day. Some

would also agree on a way of translating the sounds produced into words, in which case questions, as opposed

to statements, were indicated by repeating the signal twice.

440

Journal of the Botanical Research Institute of Texas 9(2)

As noted by lie (1960), from the 0yer area of interior SE Norway, a lur could serve as a past-time for herd¬

ers but would sometimes be put to more serious use:

“Quite a few herders had a never-lur [birch bark horn], but they were mostly used by boys, rarely

girls. The lur tones could be heard for a long distance. A bokkehaunns-stut [instrument made

from a goat horn] was more common, both with girls and boys. Both the lur and stut were

mostly used for fun and as a past-time. However, if some danger occurred, or an accident struck

(for instance that predators were at large, or a cow had collapsed), so that the herder needed

immediate help, he could sound a signal or alarm. Furthermore, it is likely that the well-known

sounds of lur and stut could help in keeping the cattle collected, and perhaps entice them to re¬

turn home at night” (lie 1960:62).

Opedal (1984) provides two examples of the instrument’s use to keep livestock safe in the Kinsarvik area of

Hardanger (Hordaland, western Norway):

“There was a lot of bears at Tjoflot. At Storahoytrsedet, the dairy maids stayed in a chalet while

herding. The bear came at night, and so they had to go outside with neverlurar and scare him

off” (Opedal 1984:109).

“The bear would ravage the sheep.—At Djonno, they had to keep them in a fence at night. We

could hear the herdsman blow his lur, and how he hit the chalet’s timber walls to scare the bear”

(Opedal 1984:109).

It should be noted that not all references to a lur or neverlur, even in this context, refer to the musical instru¬

ment. Five more records in Opedal’s vast collection of Hardanger folklore may be included to avoid confusion.

In these cases, in Kvam and Ulvik, the kind of neverlur used to chase the bear was not an instrument but simply

a curled-up piece of birch bark, set on fire to scare the beast:

“And now, the bear was walking around the summer farm. Then, another dairy made put a

burning neverlur out through the door opening. This scared him” (Opedal 1984:130).

“At another occasion, the bear came to the summer farm. It was in the evening, and the cattle

got wild. She heard how he padded about sniffing in front of the chalet. Then, she set birch bark

on fire and opened the door. And there the bear stood! And she pointed the lur at him. He

turned around at once (...)” (Opedal 1984:130).

“Afterwards that summer, the dairy maids alighted neverlurar, so that they could keep the bear

away” (Opedal 1984:135).

“At Kleivsseter, my grandmother (born 1834) put neverlurar on fire to ward off the bear. She fol¬

lowed him with the fire and said: “Burn, burn the fur!” Then, the bear got scared and left the

summer farm, running up the mountain slopes” (Opedal 1984:139).

“Every now and then, the bear would haunt the summer farms. The dairy maids alighted never¬

lurar and tied them to long sticks, and pushed it into his fur. Then he ran” (Opedal 1984:121).

At Hornindal in western Norway, a herding boy used his lur to alert his family when both he and his flock of

sheep had been taken by an avalanche during the late 19 th century:

“I had a lur. It was made of birch bark and five foot long. I carried the lur with me everywhere.

The sound it made could be heard for a long distance. I had used this instrument so much that

it was not difficult for me to produce a loud sound. When the avalanche had passed and I saw

what had happened to many of the sheep, I started blowing the lur loudly and for a long time.

And they heard me at home! After a while, help came, and we collected the dead sheep. The

meat and skins could still be used” (Fodoen 1989:62-63).

Aim, Birch bark horns and similar instruments

441

In his account of the “life of a Norwegian peasant family”, Botten-Hansen (1854) noted that children might play

the lur after finishing the day’s herding:

“In particular, they were happy when they returned from the mountains, and the herd, well-fed

by the fat grass and tired of being chased by the horseflies, settled in the summer-farm yard,

ruminating while waiting to be placed in the stall. Then, Ole stood alongside Marit, who was

busy knitting, playing his lur so beautifully that the surfeited cattle, despite their laziness,

would turn their heads against him, (...)” (Botten-Hansen 1854:43).

In southern Norway, the lur was such an integral part of farming practice that even the subterraneans, who

according to folk tradition were also herding cattle, could not do without them. At Krondalen in Jostedalen,

people had “seen a herd belonging to the subterraneans with a dairy maid, dog and pack horse etc. heading for

[the glacier] Krondalsbreen, accompanied by the sound of song and lur tones” (Asbjornsen 1850:69). A legend

from Flatdal in Seljord, Telemark relates how some mythical maidens played lur on a Sunday and enticed a

whole crowd of people attending sermon to leave the church in order to listen. The disturbance lasted until the

vicar also came outside and admonished the disturbing maidens to go into the rock (Schwach 1921:19). Ac¬

cording to the witch trials of Finnmark, northernmost Norway, even the devil played a lur while entertaining

witches at black sabbaths (Bsetzmann 1865:47).

Some were obviously more skilled at blowing a lur than others. Embret Msehlum at Stange in Hedmark,

mentioned above, gained his nickname of Tuter-Embret [“hooter-Embret”] from his skills. He obviously en¬

joyed playing, for

“each year on the 17th of May [Norway’s national day] he climbed up to this vast pine at Stortos-

ti. There, he placed the lur in the cleft of the pine, and stood there playing national songs” (En-

gen 1991:103).

Embret visited numerous other locations with his instruments. Another skilled player, Andreas Ullevalseter,

allegedly could blow his lur in such a way as to accompany traditional dancing (Holtvedt 1945:125). At Horn-

indal in Hordaland, Renda-Hans was another reputed lur blower (Lovlid 2000).

Nowadays wooden horns are mostly used for festive purposes, e.g., to signal the opening of fairs (Sevag

1966:16) or other special occasions (Engen 1991:105) and at the opening ceremony of the 1994 Winter Olym¬

pic Games at Lillehammer. Such use is well rooted in past traditions. There are several accounts of similar use

during the 18th and 19th century, at the opening of banquets and to welcome clerical dignitaries, e.g., at bishop

visitations (Wiel 1802-1805; Mehlum 1891:17; Myhre 1928; Vollsnes 2001:72). During the 19th century na¬

tional revival, the lur served as a kind of national symbol. It is featured on the front cover of a Norwegian ABC

book (Austlid 1880) and in romantic illustrations of peasant life (e.g., 0stgaard 1852, as frontispiece in some

reprint editions). Wooden horns are depicted in some foreigners’ accounts of travels in Norway, e.g., Brace

(1859; plate facing p. 56). W.H. Breton encountered the instrument in the Romsdalen area of western

Norway:

“While engaged in forcing my way on one side of the valley, I heard the notes of a horn, and

discovered that the broken acclivities of the enormous bank, although to the eye ascent seemed

impossible, admitted of pasturage for a few sheep. These were attended by a youthful shepherd,

who performed his rude music upon a horn four feet in length, and made of wood; an instru¬

ment I had seen before” (Breton 1835:272-273).

Ethnicity

All traditions noted above refer to the Norwegian majority population. It should be added, though, that similar

instruments were used by the Finnish ethnic minority of the Finnskogene (“Finn forest”) area of southeast

Norway (Lindtorp 1940; Jenssen 2007:35). A neverlur (Finnish: torvi ) from this area is described as follows:

“For a long time, nceverlurv (torvi) and buck horns ( pukinsarvi ) were the only instruments heard at Finnsko-

442

Journal of the Botanical Research Institute of Texas 9(2)

gene, and some people were masters at using them. Birch bark horns and buck horns were mostly used by

herders in forest and held, and their tunes could be heard for miles among the cliffs” (Lindtorp 1940:97).

DISCUSSION

The birch is hardly mentioned in classical Greek and Roman sources, playing no role in their homelands (de

Cleene & Lejeune 2003:150). A cursory treatment is included in book XVI of Pliny’s Historia naturalis, where it

is first referred to as a Gallic tree (XVI:74), noting its “remarkable white colour” and the use for various utility

purposes (Rackham 1968:437). Pliny also comments on its use for withies (XVI: 176, Rackham 1968:501) and

the quality of the wood (XVI:209, Rackham 1968:525).

The birch is much more important in the north. It features prominently in the traditions of Central and

Northern Europe and northern Asia, both in a religious context and as a utility. In Norse mythology, the tree

was dedicated to Thor, the god of thunder and lightning.

Birch bark has been used for various purposes, e.g., as thatching on roof. Being strong and flexible, it has

also been used to prepare baskets and shoes in Norway. Birch bark baskets (Norwegian: neverkont) were fre¬

quent all over the country and are still sometimes prepared and used. Birch bark shoes were previously much

used by the Finnish ethnic minority of southeastern Norway (Matson 1908; 0stberg 1935). Both these uses are

also well-known in neighboring Sweden (Hasselrot 2005; Rosen 2005). Being highly inflammable, bark was

and still is frequently used to light fires and has also served in torches (de Cleene & Lejeune 2003:157). Like

many other kinds of bark, it could be used for tanning. The thin, inner bark has been used on wounds in both

Norwegian and Sami folk medicine.

Since antiquity, birch bark has also served as writing material (de Cleene & Lejeune 2003:157). The San¬

skrit name bhurja means “the light tree or a tree with bark one can write on.” In Norwegian sources, such use

is first mentioned in the late 16th century by Peder Clausson Friis (Friis 1632, reprinted in Storm 1881). He

noted that bark was used for letters and as practicing material for children who were learning to write. Birch

bark was still used for this purpose in the 18th century (Frimann 1885; Schubeler 1885:482).

A complex and dissected topography may have contributed to the prevalence of birch bark horns in Nor¬

way. They were once common all over southern Norway but seemingly little used further north. The distribu¬

tion map in Fig. 4 is based on the same multitude of sources used for compiling this paper. It is likely to reflect

the general pattern, even if most instruments have of course been made, used, and discarded without making

it into literature or museum collections.

The sources are silent in terms of what kind of birch was used. Given the straight growth and the size of

the stem, one may assume that Betula pendula Roth provided better bark and winding material than the lower,

more crooked and richly branched Betula pubescens Ehrh. This may explain the absence or sparsity of lur-type

instruments in northernmost Norway. The distribution of Betula pendula tapers rapidly out towards the north,

and the species is sparse or absent in the three northernmost countries (Hulten 1971: map 597), mirroring the

distribution of recorded instruments shown (Fig. 4).

The three northern outposts may deserve a comment. They are found, from south to north, in Vefsn and

the Salten (Beiarn-Saltdal) area of Nordland, and in Bardu, Troms. In the Vefsn area, ropes served as a substi¬

tute for Betula bark, tying the two halves of the wooden kernel together (Fig. 3). Further north, the Beiarn-

Saltdal area in Nordland, Saltdal in particular, has a favorable local climate, with hot and dry summers and a

northern exclave of Betula pendula var. lapponica (Lindq.) Hamet-Ahti (see map in Elven 2013). The Bardu area

lacks Betula pendula. It was, however, settled by people migrating north from interior SE Norway from the late

18 th century onwards, thus coming from within the main distribution area of the lur (Fig. 4). They brought

with them the tradition of making wooden horns and perhaps even some instruments. Thus, the distribution

of the birch bark horn tradition within Norway is likely to reflect the availability of suitable winding material

(i.e., bark from Betula pendula) rather than the need for or will to produce such instruments further north.

Similar studies in Sweden or Finland may confirm this pattern.

It should be noted, though, that other predominantly ethnic Norwegian traditions taper out towards the

Aim, Birch bark horns and similar instruments

443

Fig. 3. Norwegian female with a somewhat deviant lur, secured by a coiled-up rope, photographed in 1970 at Mjavatn in Vefsn, Nordland, north Norway,

by Arnt Bakke (photo archive of Tromso museum, TSNF 9731).

north, where local communities are more diverse and may be of either ethnic Norwegian, Sami, or Finnish

origin or a mixture of these. The traditon of planting Rhodiola rosea L. on roofs as a supposed (or apotropaic)

protection againts fire is widely known in southern Norway, extending northwards into Nordland to the Bei-

arn area of Salten (Aim 2004), with a pattern almost identical to the distribution of the lur. The unique Norwe¬

gian tradition of using Linnaea borealis L. to treat shingles (herpes zoster) is also widespread in southern and

central Norway but unknown further north (Aim 2006).

Sweden, with a more gentle terrain but as wide distances, can compete in terms of the number of wooden

horns on record, e.g., in the collections of Nordiska museet (18 of these are depicted in Lid & Solheim 1936).

Kjellstrom (1994) provides a brief, general account of “Scandinavian bark horns.” More or less similar horns

were used in Finland, e.g., to scare off bears (McCune & Prendergast 2002), but also featured in folk music

(Austerlitz 2000).

Kew’s Economic Botany Collections contain examples of birch bark horns from Norway, Finland, and

Switzerland (McCune & Prendergast 2002), thus adding another European country with substantial topo¬

graphic relief. In his lexicon of musical instruments, Sachs (1964) defines the lur simply as “das skandinavis-

che Alphorn” (“the Scandinavian Alp horn”), although the Swiss Biichel horns deviate in shape from the simple

Scandinavian tubes and include a convoluted, more or less trombone-like wind tube (Geiser 1976). Some of the

Swedish birch barks horns depicted by Lid & Solheim (1936) approach this type.

In all these cases, modern means of communication has removed the need for signalling with wooden

horns. Their survival into the present depends solely on their status as traditional instruments arousing some

national pride (Fig. 5), the special sound produced, and the sheer beauty of well-made objects. Modern lurs are

also used by some players of folk music.

444

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 4. Map showing geographical distribution of the Norwegian wooden horns mentioned in text. The approximate northern limit of Betula pendula

as a common feature of lowland forests is indicated by a hatched line, and two disjunct, northern areas of B. pendula var. lapponica by dotted lines.

Aim, Birch bark horns and similar instruments

445

Fig. 5. Early 20th-century postcard showing two Norwegian females in national costumes, one knitting and the other blowing a lur.

446

Journal of the Botanical Research Institute of Texas 9(2)

ACKNOWLEDGMENTS

Professor Arne Emil Christensen, University of Oslo, and responsible curator of the Oseberg material, checked

the records relevant to the Oseberg wooden horn. Sturla Binder, Osteroy museum, provided access to the post¬

card reproduced in Figure 5.1 thank one anonymous reviewer who provided excellent feedback on an earlier

draft.

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FIRST COLLECTION OF MICONIA ALAINII

(MELASTOMATACEAE: MICONIEAE) WITH FLOWERS

Walters. Judd

Department of Biology, 220 Bartram Hall, and

Florida Museum of Natural History

University of Florida

Gainesville, Florida 326 7 7, US.A.

lyonia@ufl.edu

Teodoro Clase

Jardm Botanico National Dr. Rafael M. Moscoso

Apartado21-9

Santo Domingo, REPUBLICA DOMINICANA

ted_clase@hotmail.com

Lucas C. Majure

Department of Research, Conservation, and Collections

Desert Botanical Garden, 1201 North Galvin Parkway

Phoenix, Arizona 85008, U.S.A.

Imajure@dbg.org

ABSTRACT

Miconia alainii, a species previously known only from specimens in vegetative condition or with immature fruits, is now known from flower¬

ing material collected by the second author in the Sierra de Baoruco, Dominican Republic. A revised description of this species, including

previously unknown floral characters, is presented.

Keywords: Dominican Republic, Sierra de Baoruco, Miconia alainii, Miconia sect. Chaenopleura , Melastomataceae

RESUMEN

Miconia alainii es una especie previamente conocida solo de material en condicion vegetativa o con frutos inmaduros, ahora es conocida de

un material con flores que fue colectado por el segundo autor en la Sierra de Baoruco, Republica Dominicana. Se presenta una descripcion

revisada de esta especie en la que se incluyen los caracteres florales antes no conocidos.

Miconia alainii Judd & Skean (Miconieae: Melastomataceae) is a poorly collected species that is endemic to the

moist montane forests of the easternmost peak of the Sierra de Baoruco, a location usually referred to as “Mon-

teada Nueva” in reference to a nearby coffee hnca, but the region actually occupies the summit of Loma Trocha

de Pey (Judd & Skean 1994, Judd 2007). A recent collection of this species by the second author (i.e., Clase

8037, see below) from this locality represents the first collection of this species with flower buds and open flow¬

ers - the species was previously known only from the type collection ( Liogier 25179), which has immature

fruits, and a second gathering (Judd 6569), which is in vegetative condition (and is also a voucher for DNA-

material in silica).

To this point the species has been only provisionally placed within Miconia sect. Chaenopleura because

the putative morphological synapomorphies for that section (as represented in the Greater Antilles) are en¬

tirely reproductive: an actinomorphic androecium of erectly held, obovate anthers that open by two longitudi¬

nal slits, exposing the contents of the four locules, and pale blue berries (see Judd 2007). None of these mor¬

phological characteristics have been available for this species until now. However, the stamens of Clase 8037

are perfectly characteristic of section Chaenopleura: obovate with two longitudinal slits running nearly their

entire length. There can be no doubt, therefore, that this species belongs in section Chaenopleura, which con¬

stitutes a large and morphologically diverse group in the Greater Antilles (Judd 2007). This phylogenetic place¬

ment is also supported by molecular data (F. Michelangeli, unpublished) since DNA nucleotide sequences de¬

rived from Judd 6567 have been included in a phylogenetic analysis of Miconieae, and in this unpublished

analysis M. alainii is placed with other species of this clade.

Morphologically Miconia alainii appears to be quite isolated among the Greater Antillean species of Mico¬

nia section Chaenopleura. It is anatomically nearly unique in having the spongy mesophyll of its leaf blades

J. Bot. Res. Inst. Texas 9(2): 449 - 451.2015

450

Journal of the Botanical Research Institute of Texas 9(2)

strongly lignified (Judd 2007), and it lacks the suites of potential synapomorphies of the nine major infrasec¬

tional clades outlined in the monograph of the group (Judd 2007). Like members of the bullate-leaved clade it

has cordate-based leaves with six secondary veins and the abaxial leaf surface with quaternary veins consis¬

tently raised and prominent. However, it lacks the prominently thickened nodal lenticels, bullate adaxial leaf

surface, and rounded cymes characteristic of this group, which includes species such as M. favosa (Desr.)

Naudin, M. xenotricha Urb. & Ekman, M. howardiana Judd, Salzman & Skean, and M. sintenisii Cogn. Prelimi¬

nary phylogenetic analyses (still unpublished) based upon DNA-sequence data also suggest that it is not

closely related to these species. It may be a fairly early divergent member of Antillean sect. Chaenopleura.

This new material allows us to present here an updated and much more complete description of Miconia

alainii. The description follows the format of Judd (2007) except that the structures here called calyx teeth were

in Judd (2007) called external calyx lobes. The structures here called calyx lobes were referred to as internal

calyx lobes in Judd (2007). In addition, we have specified in the description the condition of both the crown

and the collar at the ovary apex.

DESCRIPTION OF MICONIA ALAINII (SEE ALSO JUDD 2007)

Shrub to 3 m tall. Indumentum of darkly ferrugineous, globular-stellate to irregularly branched or elongate

short-branched hairs, and minute-globular hairs. Young twigs not ridged, ± rectangular in cross section, be¬

coming terete with age, the indumentum dense, of darkly ferrugineous, elongate short-branched to globular-

stellate hairs, these ± persistent; internodes 1.4-10 (-15) cm long. Leaves with petiole 1.5-7.5 cm long, the in¬

dumentum same as that of twigs; blade 4-16 cm long, 3.7-10.5 cm wide, ovate to elliptic, flat, coriaceous, the

apex acuminate or shortly acuminate, the base cordate, the margin plane, entire to sparsely and very shallowly

dentate distally, 0-70% of margin entire, the largest teeth 0.1-1.4 mm long; venation acrodromous, slightly

suprabasal, with prominent midvein and 6 secondary veins, with 4 conspicuous secondary veins, the inner

pair placed 5-23 mm in from margin and the outer pair placed 1.3-5 mm in from margin, and 2 inconspicuous

secondary veins closer to margin, and numerous percurrent tertiary veins, oriented subperpendicular to mid¬

vein, jointed by percurrent-orthogonal quaternary veins; adaxial surface green, the indumentum initially

dense globular-stellate, but quickly glabrescent, the midvein and secondary veins slightly impressed, other

veins ± flat, the surface appearing only very slightly wrinkled after drying, with sparse druse crystals; abaxial

surface light green, sparsely to moderately ferrugineous, globular-stellate to elongate short-branched hairs to

0.1-0.25 mm across, along with minute-globular hairs on midvein and major secondary veins, such hairs very

sparse to sparse on minor secondary, tertiary, and higher order veins, the hairs ± persistent and epidermis

clearly visible, the midvein and major secondary veins prominently raised, minor secondary, tertiary and qua¬

ternary veins raised, higher order veins slightly raised to flat. Inflorescences many-flowered, open-paniculate

cymes of 3-5 major branch pairs, ca. 5-6.5 cm long, 4-5 cm in diameter; proximal segment of lowermost in¬

florescence branches 0.8-1.2 cm long, distal internodes of inflorescence branches increasingly shorter, ulti¬

mate branches 1-9 mm long; moderate ferrugineous, globular-stellate hairs, and minute-globular hairs; pe¬

duncle 1.5-1.9 cm long, with similar indumentum; each inflorescence branch associated with an early cadu¬

cous, ovate to ovate-triangular bract, ca. 1.3-4 mm long, 0.5-1 mm wide, the apices acute; flowers in dichasia,

each subtended by 2 caducous bracteoles, ca. 1-1.5 mm long, 0.25-0.4 mm wide, narrowly triangular to ± lin¬

ear, nearly glabrous, with acute apices. Flowers sessile or nearly so, the pedicel 0-1 mm long. Hypanthium

cylindrical, free portion 0.5-0.8 mm long, the outer surface with sparse to moderate, ferrugineous, globular-

stellate hairs and a few minute-globular hairs, the inner surface glabrous and obscurely 10-ridged, and apices

of the ridges slightly projecting. Calyx teeth 5, 0.3-0.9 mm long, 1-1.7 mm wide, triangular, with acute apex,

indumentum of moderate stellate-globular hairs; calyx lobes 5, 0.3-0.7 mm long, 1-1.7 mm wide, broadly tri¬

angular, with very sparse stellate-globular hairs, with acute to rounded apex, the margin entire; calyx tube

0.3-0.4 mm. Petals 5, 2.4-2.5 mm long, ca. 1.5 mm wide, broadly ovate to elliptic, ± cupped, glabrous, imbri¬

cate and apically interlocking in bud, the apex rounded and with an asymmetrically located notch; margin

entire. Stamens 10, geniculate, positioned in a ring around the flower, thus androecium actinomorphic, with

Judd et al., Flowering specimens of Miconia alainii

451

the filaments flexed back and the anthers held erectly; proximal segment 1.8-2 mm long; distal segment 2-2.3

mm long, with minute dorsal projection on the back of the anther, the anther obovate, 1-1.2 mm long, opening

by 2 longitudinal slits, with fertile portion of anther sacs 0.9-1 mm long, the connective/distal part of filament

extended 0.8-1 mm beyond the base of the anther. Ovary 3-loculate, 2/3-inferior, 2-2.8 mm long, 1.5-2.4 mm

in diameter, globose to ovoid, glabrous and ridged distally, with fluted apical projection (collar) to 0.5 mm en¬

circling the base of style, but crown absent; style ca. 3.3 mm long, straight, glabrous; stigma truncate. Berries

(immature) ca. 3.5 mm in diameter, globose, green with red tinge (but probably blue and slightly larger at ma¬

turity), with sparse to moderate stellate-globular and minute-globular hairs. Seeds 0.7-1 mm long, angular-

obovoid, with a prominent raphe; testa smooth.

Voucher specimens: DOMINICAN REPUBLIC. Prov. Barahona: Sierra de Baoruco, Municipio Polo, lugar denominado Cortico, bosque

nublado, fragmentado, con Brunellia comocladifolia, Prestoea montana, Schefflera tremula, Meliosma impressa, Cyathea sp., Alsophila sp., etc.

UTM265039mEm 2003784mN, elev. 1374 m, 30 Jun 2013 (fl), T. Clase 8037 , withJ.R. Martinez, R. Cdmara and I. Santos (FLAS, JBSD); Sierra

de Baoruco, Loma Trocha de Pey or “Monteada Nueva,” above (E of) Polo, 18 May 1992 (sterile) W.S.Judd 6569 (F, FLAS, JBSD, MO, NY, US);

Barahona: Sierra de Baoruco, Cana Brava, Monteada Nueva, 1300 m, 24 Apr 1976 (young fr), A.H. Liogier 25179 (holotype: JBSD).

It is perhaps surprising that some Antillean species have to this point not been collected in flowering condi¬

tion, as the Greater Antilles are considered to be fairly well collected. However, this is unfortunately not that

uncommon, and for example flowers are still unknown for the recently described Miconia cineana Majure,

Judd, Ionta & Skean (Majure et al. 2014) and Miconia abscondita Majure, Judd & Skean (Judd et al. 2015), two

rare endemics to the southern mountains of Hispaniola (Massif de la Hotte, Haiti) that were described with the

combination of DNA sequence and vegetative morphological data, and flowering material was only collected in

2013 by Eldis Becquer and Fabian Michelangeli (pers. comm.) of a related species (Judd 2007), Miconia turqui-

nensis Urb. & Ekman, of the Sierra Maestra, Cuba. Another very rare species in Miconia sect. Lima has yet to be

described from the Massif de la Hotte, as fertile material will be necessary to distinguish it from close relatives

(Majure et al. unpubl. data). Finally, Miconia alloeotricha (Urb.) Judd, Penneys & Skean, another endemic to the

Massif de la Hotte, was described by I. Urban on the basis of specimens with very young inflorescences in 1929,

and was only collected with mature flowers and fruits in 1989 and 1993 (Judd et al. 2004).

ACKNOWLEDGMENTS

This work was supported, in part, by a grant from the National Science Foundation (BSR-0818399). We thank

Frank Almeda and Eldis R. Becquer for their helpful comments on this paper.

REFERENCES

Judd, W.S. 2007. Revision of Miconia sect. Chaenopleura (Miconieae, Melastomataceae) in the Greater Antilles. Syst. Bot.

Monogr. 81:1-235.

Judd, W.S. & J.D. Skean, Jr. 1994. Miconia alainii (Melastomataceae: Miconieae), a new species from Hispaniola. Novon

4:112-115.

Judd, W.S., D.S. Penneys, & J.D. Skean, Jr. 2004. Rediscovery of Ossaea alloeotricha, an endemic of the high-elevation Massif

de la Hotte, Haiti, and its transfer to Miconia (Melastomataceae: Miconieae). Brittonia 56:159-165.

Judd, W.S., L.C. Majure, J.D. Skean, Jr., & K.M. Neubig. 2015. Miconia abscondita (Melastomataceae: Miconieae), a new spe¬

cies from the Massif de la Hotte, Haiti: rediscovered in herbaria after being hidden for nearly nine decades. Rhodora

117:317-341.

Majure, L.C., W.S. Judd, G.M. Ionta, J.D. Skean, Jr., E.R. Becquer, & K.M. Neubig. 2014. Miconia cineana (Melastomataceae:

Miconieae), a new species from the Massif de la Hotte, Haiti, based on morphological and molecular evidence. Syst.

Bot. 39:906-914.

452

Journal of the Botanical Research Institute of Texas 9(2)

BOOK NOTICE

Stephen C. Meyers, TheaJaster, Katie E. Mitchell, & Linda K. Hardison, eds. 2015. Flora of Oregon, Volume 1:

Pteridophytes, Gymnosperms, and Monocots. (ISBN: 978-1-889878-46-1, hbk). Botanical Research

Institute of Texas Press, Fort Worth, Texas 76107, U.S.A. (Orders: shop.brit.org, orders@brit.org, 1-817-

332-4441). $75.00, 608 pp., 7.5" x 10.5".

From the Publisher —The Oregon Flora Project, Oregon State

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laborated to publish the Flora of Oregon, the first comprehensive flo¬

ra of Oregon in over 50 years—with illustrations!

Flora of Oregon is a three-volume reference that will be the

state’s only flora published in the past half century and the first il¬

lustrated floristic work that exclusively addresses Oregon. Volume 1

presents treatments of the pteridophytes, gymnosperms, and mono¬

cots—1,054 taxa, or 23% of all native and naturalized vascular

plants of Oregon. The taxonomic treatments include dichotomous

keys, family and generic synopses, full taxon descriptions, and il¬

lustrations. A dot map depicting vouchered occurrences and high¬

lighted ecoregions that host the taxon accompanies each descrip¬

tion. There are pen and ink illustrations of 521 taxa, including 86

new works by artist John Myers.

Color photographs accompany chapters describing the state’s

ecology and sites for exploring botanical diversity. Also included are

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Flora of Oregon

J.Bot. Res. Inst. Texas 9(2): 452.2015

ASTERACEAE EN EL DURAZNO Y CERCANIAS, NORTE DE TAMAZULA,

DURANGO (MEXICO): RIQUEZA, DISTRIBUCION Y ENDEMISMO

David Ramirez Noya y Yolanda Herrera Arrieta*

IPNCIIDIR Durango

Sigma 119, Fracc. 20 de Noviembre II

Durango, Dgo., MEXICO 34220

yolah54@gmail.com

RESUMEN

Se estudio la flora de asteraceas y su nivel de endemismo en las cercamas de la comunidad El Durazno, al norte del municipio de Tamazula,

Durango, Mexico, en localidades de dificil acceso y escasamente exploradas de la Sierra Madre Occidental. Se realizaron recorridos y re-

colectas de material botanico en las cuatro estaciones del ano, en el periodo de 2001 a 2014. La riqueza floristica encontrada de la familia

Asteraceae es de 75 generos y 148 especies, de ellas 51% son especies endemicas de Mexico. Se reportan 17 nuevos registros para el estado de

Durango y un endemismo a nivel estatal de 50.33% de generos y 25.75% de especies. Las principales comunidades vegetales de la region es-

tudiada se encuentran en altitudes de entre 670 y 2550 m y estan representadas por bosque de coniferas (2050 a 2550 m) donde se encontro

un 49.34% de las asteraceas registradas en este estudio, bosque de encino-pino (1500-1900 m) con un 39.47% de especies y bosque tropical

caducifolio (600-1000 m) con un 22.37%, por lo que se concluye que la mayor riqueza floristica de esta familia se presenta en el ecosistema

templado subhumedo. De acuerdo con el origen de los taxa, 96% de los generos y 95.27% de las especies son autoctonos y el resto son exoti-

cos. Se registra tambien un 27.7% de riqueza de especies sinantropicas.

Palabras Clave: Asteraceae, Durango, endemismos, nuevos registros

ABSTRACT

We studied the Asteraceae flora and its level of endemism in the vicinity of the community El Durazno, northwest of the municipality

Tamazula, Durango, Mexico. The locations are difficult to access and are poorly explored areas in the Sierra Madre Occidental. Field trips

were conducted to collect botanical material in all four seasons of the year from 2001 to 2014. The floristic richness of the Asteraceae family

consisted of 75 genera and 148 species, of which 51% are endemic to Mexico. We report 17 new records for the State of Durango. Endemism

is high with 50.33% of genera and 25.75% of species. The main plant communities of the region are found at altitudes between 670 and 2550

m and are represented by: 1) coniferous forest (2050 to 2550 m) where 49.34% of the Asteraceae were recorded in this study; 2) Pinus-

Quercus forest (1500-1900 m) with 39.47% of species; and 3) deciduous tropical forest (600-1000 m) with 22.37%. It can be concluded that

the richest area for this family occurs in the subhumid temperate ecosystem. Accordingly, 96% of genera and 95.27% of species are native,

and the rest are exotic. Some 27.7% of the Asteraceae taxa are synantropic species.

Keywords: Asteraceae, Durango, endemism, new records

INTRODUCCION

Las especies endemicas se distribuyen de forma irregular en la superficie terrestre. Un taxon endemico se de¬

fine como “aquel que restringe su distribucion a una region especlfica” (Suarez-Mota y Villasenor 2011), repre-

sentando la fraccion exclusiva de su riqueza biologica. La determinacion de las areas ricas en endemismos es

importante para los estudios biogeograficos historicos, ya que dichas areas se constituyen como probables si-

tios de origen y diversificacion de grupos florlsticos (Rzedowski 1991b).

Villasenor (2003) considera a Mexico como uno de los primeros cinco palses megadiversos por su riqueza

floristica y alto nivel de endemismo en su flora. El endemismo de la flora de Mexico ha sido estudiada a nivel

nacional, estatal y regional; Rzedowski (1991a) estima un 52% de endemismo en Mexico, mientras que Vil¬

lasenor (2003) estima un 65.9%. A nivel estatal los estudios son escasos, resaltan los de Balleza y Villasenor

(2002) para Zacatecas, Castillo-Campos et al. (2005) para Veracruz o Suarez-Mota y Villasenor (2011) para

Oaxaca. El endemismo tambien se discute en trabajos realizados a nivel regional, como los de Hernandez-L.

(1995) en la Sierra de Manantlan, Jalisco-Colima; Mendez et al. (2004) en el Valle de Tehuacan-Cuicatlan,

Puebla-Oaxaca o el de Hinojosa-Espinosa y Cruz-Duran (2010) en el municipio de Atenango del Rio, Guerrero,

entre otros.

J. Bot. Res. Inst. Texas 9(2): 453 - 470.2015

454

Journal of the Botanical Research Institute of Texas 9(2)

La familia de las Compuestas o Asteraceae, es una de las mejor representadas a nivel mundial dentro de

las plantas con flor (Magnoliophyta), dado que se estima que cuenta con 1,535 generos y aproximadamente

23,000 especies (Bremer 1994). De acuerdo con Rzedowski (1991b) es la familia mas diversa en Mexico, con

aproximadamente 314 generos y 2,400 especies (52% de endemismos); posteriormente Villasenor (2003) es-

timo que existen 361 generos y 3,021 especies (65.9% de endemismo) de Asteraceae a nivel nacional. Ambos

autores resaltan que en Mexico se concentra una riqueza elevada de especies de Asteraceae, cuando se compara

con otras regiones del mundo, por lo que proponen que podrla ser su principal centro de diversihcacion. Adi-

cionalmente Villasenor (2003) indica que Asteraceae es la familia de fanerogamas con el mas alto porcentaje de

endemismo en Mexico, en este sentido, llevar a cabo mas estudios a nivel estatal y regional contribuira a con-

firmar dicha teorla y a conocer mas su distribucion florlstica en el pals.

Las Compuestas se desarrollan principalmente en climas templados y secos, la mayorla de ellas como

especies nativas de los ecosistemas que habitan (Rzedowski 1972; Villasenor 1993); sin embargo, un alto por¬

centaje de especies de asteraceas se ve favorecida por la perturbacion (flora sinantropica), por lo que suelen ser

dominantes en medios arvenses y/o ruderales (Villasenor 2012). A la fecha son escasos los estudios de la diver-

sidad de plantas que se distribuyen en la Sierra Madre Occidental, especialmente en sus laderas hacia el Paclf-

ico, por razones complejas como es la combinacion de lo accidentado de la sierra, el diflcil acceso y la lejanla de

centres urbanos, sin mencionar los problemas de inseguridad que aquejan la region. Asi, el objetivo del pre¬

sente estudio fue contribuir al conocimiento de la riqueza y endemismos de la familia Asteraceae en la region

de El Durazno y cercanlas, al norte del municipio de Tamazula, Durango, Mexico. El area explorada se ubica

en la region de las quebradas, ladera oeste de la Sierra Madre Occidental, al noroeste de Durango (Fig. 1), justo

en las proximidades a la convergencia con los estados de Chihuahua y Sinaloa. Se caracteriza por ser particu-

larmente de diflcil acceso, motivo por el cual los muestreos de flora y fauna de estas localidades son escasa-

mente conocidas, lo que reforzo la idea de iniciar el presente estudio, esperando encontrar especies endemicas

y de ocurrencia escasa o rara.

No existen reportes de estudios florlsticos realizados en el area especlhca del presente estudio. Gomez

(2005) establece que los primeros exploradores botanicos en visitar estos rumbos “... viniendo de Alamos,

Son., y cruzando la sierra entre Canelas y Topia hacia Tepehuanes ...” fueron J.M. Mocino y Juan Diego de

Castillo, entre los anos de 1791 y 1792, sin embargo no se conoce la existencia de ejemplares de esa expedicion

que provengan de nuestra area de estudio. Existen registros de ejemplares en los herbarios MBO y US (www.

tropicos.org) de colectas realizadas por J. Gonzalez Ortega entre 1921 y 1922 (compuestas, gramlneas, pottia-

ceas y bignoniaceas) en el municipio de Tamazula, Durango y por Gentry en 1939 (leguminosas), todas ellas

de localidades cercanas a la cabecera municipal de Tamazula, misma que se encuentra a una distancia aproxi-

mada de 55 km al sur del area de estudio en llnea recta. As! tambien, encontramos otros estudios realizados en

localidades ubicadas en un area geograhca radial a distancias menores de 60 km en llnea recta de nuestra area

de estudio, como son los de Greenman (1905); Turner (1986); Nesom (1989a, 1989b, 1990, 1992); Panero y

Villasenor (1993), Vega Avina et al. (2000) y McDonald et al. (2011), quienes mencionan o describen especies

que ser an comparadas con las registradas en este reporte.

El presente estudio fue realizado en el Centro Interdisciplinario de Investigacion para el Desarrollo Inte¬

gral Regional, Unidad Durango (CIIDIR IPN Durango). El primer autor realizo recorridos de exploracion y

muestreo en dos tramos (I y II) de la region, a traves del transito por brechas y veredas, en diferentes meses del

ano, durante el intervalo del perlodo comprendido entre los anos 2001 a 2014; en el marco de estudios de carac-

ter lingulstico, zoologico y botanico que se realizan en el CIIDIR IPN Durango. Concerniente a lo botanico, D.

Ramirez N. se enfoco a la familia Asteraceae por interes personal (y de su especialidad) y con el proposito unico

de contribuir al conocimiento de la biodiversidad de dicha familia en el Estado.

Area de estudio

El Municipio de Tamazula (Fig. 1) esta ubicado al oeste del estado de Durango, colinda con los municipios

Guadalupe y Calvo, Chihuahua y Culiacan, Sinaloa. Registra altitudes desde 190 m, en las cercanlas a la ca¬

becera municipal, hasta 2900 m en localidad contigua a El Durazno. Las cercanlas al Durazno es la region ex-

Ramirez Noya y Herrera Arrieta, Asteraceae en El Durazno,Tamazula, Durango, Mexico

455

• SITIOS DE COLECTA

ASENTAMIENTOS HUMANOS

habitantes

• 0-1

o 2-250

O 251 - 500

ALTITUD (msnm)

^■ 185-200

i_~J 201 - 400

I I 401 - 600

I I 601 - 800

I |801 - 1,000

I 11,001 - 1,200 f)

□ 1,201 - 1,400 ^ 501 ‘ 793

□ 1,401 - 1,600 CAMINOS

[□ 1,601 - 1,800 TIP0

|-1 1,801 - 2,000 """" Terracerfa

| 2,001 - 2,200 . BRECHAS

2,201 - 2,400 □□ LIMITE MUNICIPAL

□I 2,401 -2,600

I I 2,601 - 2,800

|~B 2,801 - 2,945

Kilometres

0 2.5 5

I-1-1-1 I-1—

Mapa en

Coordenadas Geograficas

Sistema de Referencia Horizontal

WGS 84

Fig. 1. Area de los recorridos en El Durazno y cercamas, al norte del municipio Tamazula, Durango, Mexico.

456

Journal of the Botanical Research Institute of Texas 9(2)

plorada y recolectada para el presente estudio (Fig. 1), se encuentra ubicada a 55 km al norte en llnea recta de

la cabecera municipal (Tamazula); es la poblacion mas grande de nuestra area de estudio (por lo cual se toma

como referencia), cuenta con menos de 800 habitantes. Le sigue en importancia el poblado San Juan del Tecu-

an, ubicado a 10 km al norte de El Durazno, con menos de 500 habitantes; otros asentamientos humanos son

comunidades con menos de 250 habitantes, o bien rancherlas de 1 a 10 familias dispersas en el area explorada,

en una porcion de la “region de las quebradas” de la Sierra Madre Occidental.

Las localidades exploradas en el presente estudio pertenecen a la cuenca del Rio Humaya, considerada

por la CONABIO como region terrestre prioritaria para su conservacion (Clasihcacion no. RTP24, Rio Huma¬

ya). El area estudiada cuenta con las siguientes coordenadas extremas: al noroeste 25°33'37"N, 107°04'07" W, a

1512 m sobre el nivel del mar, en la rancherla “El Cordon”; al norte 25°35'21"N, 107°00'42"W, a 2745 msnm, al

sur25°20'20"N, 106°57'05"W, a 669 msnm, proximo al rancho “SantaMatilde”;aleste25°32'24"N, 106°56'58"W

a 2383 msnm; y al oeste 25°33'37"N, 107°04'07"W a 2383 msnm. Las comunidades vegetales que comprende

(Gonzalez et al. 2007) son: bosque de conlferas (de extension restingida) con Abies durangensis, Pinus strobifor-

mis y P. durangensis, encontrado en altitudes de 2425 a 2550 m en canadas humedas; bosque de pino en

lomerlos suaves y mesetas, con Pinus durangensis, P. arizonica, P. leiophyla y P. cooperi, en altitudes de 2026 a

2400 m. Bajando a aproximadamente 1650 m de altitud se encuentra el bosque de pino-encino con Pinus teo-

cote, P. lumholtziiy Quercus spp., con la presencia de Magnolia schiedeanay Juglans sp. en canadas humedas

(justo en la localidad La Presa, de los Aguajes). Cerca de 1525 m de altitud se encuentra el bosque de encino

Quercus spp. y Purser a spp., con Lippia umbellata y Montanoa leucantha bien desarrolladas y muy abundantes en

esta comunidad vegetal. Este bosque parece presentar el area de transicion entre el ecosistema templado y el

tropical, por la presencia de especies de ambos ecosistemas. En las partes mas bajas se encuentra el bosque

tropical caducifolio, en altitudes de 670 a 1400 m con Lysiloma acapulcensis, Acacia pennatula, A. farnesiana,

Ipomoea arborescens, Stemmadenia tomentosa, Bursera spp., Randia spp. y Heliocarpus entre otras, y ya en la

parte mas baja de las quebradas se encuentra Ficus spp., Cochlospermum vitifolium, Tabebuia spp., Acacia spp. y

Lysiloma spp. Las principales localidades (areas de recoleccion intensa) se presentan en la Tabla 1, sin dejar de

recolectar especies diferentes en todo el trayecto.

La descripcion de los recorridos se presenta a continuacion:

a) . Recorrido del tramo I.—ubicado en la parte norte-noroeste de la comunidad El Durazno (Fig. 1), el

punto de partida generalmente fue del poblado Sanjuan del Tecuan, Tamazula, Dgo. (25°34'05"N, 106°57'20"W

y 2400 m de altitud), donde se realizaron colectas al azar caminando por brechas y veredas temporales y de

forma continua en una trayectoria zigzagueante hacia el oeste, que asciende hasta 2550 m de altitud, posterior-

mente descendiendo a la rancherla Aguazul, a 1590 m de altitud; esto con el hn de muestrear la mayor superh-

cie posible. Los recorridos representaron, un total aproximado de 60 km, comprendiendo el bosque de

conlferas ( Abies durangensis, Pinus strobiformis y P. durangensis), bosque de pino ( Pinus durangensis, P. leiophyla

y P. cooperi), hasta el ecotono con el bosque tropical caducifolio. Se reunieron en este tramo un total de 388

registros de ejemplares recolectados, de los que un numero reducido corresponden a ejemplares de fauna

silvestre.

b) . Recorrido del tramo II.—se llevo a cabo al sur de la comunidad El Durazno (Fig. 1). Esta segunda ruta

de exploracion se realizo a partir de las proximidades de La Rancherla San Dario (25°22'36"N, 106 o 55'18"W y

1837 m de altitud), transitando con rumbo oeste-sureste, hasta la parte baja de la Quebrada Camarones,

25 o 21'40"N, 106°5741"W, llegando a bajar hasta 670 m de altitud, en el arroyo Camarones. Las recolectas se

realizaron basicamente siguiendo las brechas y veredas temporales que permiten acceso a pie o con el apoyo de

animales, ya que lo abrupto del terreno limito efectuar caminatas direccionadas. El transecto cubre un total

aproximado de 40 km de recorrido, en comunidades vegetales de bosque de pino ( Pinus durangensis, P. lumholt-

zii), pino-encino ( Pinus teocote, Quercus spp.), a bosque tropical caducifolio de Lysiloma acapulcensis, Acacia

spp., Bursera spp. y Ficus spp. Se reunieron en este tramo un total de 472 registros de ejemplares recolectados,

y al igual que en el Tramo I, un numero reducido de ellas corresponde a ejemplares de fauna silvestre.

Ramirez Noya y Herrera Arrieta, Asteraceae en El Durazno, Tamazula, Durango, Mexico

457

Tabla 1. Localidades muestreadas y taxa encontrados en las cercamas de El Durazno, Tamazula, Durango, Mexico.

Acmello radicans (Jacq.) R.K. Jansen. Predio La Presa, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'27"W, 8 nov 2001,

bosque de Quercus sp v Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2077 (CIIDIR). Arroyo Camarones, Los Aguajes, El Durazno,

Tamazula, Dgo., 25°21 '40"N, 106°57'41 "W, 28 may 2002, bosque tropical caducifolio de Lysiloma spp., Acacia spp., Bursera spp. y Ficus

spp., 710 m, D. Ramirez Noya 2195 (CIIDIR).

Acmella repens (Walter) Rich. Sta Matilde, Los Aguajes, El Durazno,Tamazula, Dgo., 25°21 '21"N, 106°57'53"W, 24 may 2003, bosque tropical

caducifolio de Lysiloma spp., Acacia spp., Bursera spp. y Ficus spp., 670 m, D. Ramirez Noya 2362 (CIIDIR).

Acourtia macrocephala Sch. Bip. Los Bajios, El Tecuan,Tamazula, Dgo., 25°33'15"N, 106°57'10"W, 20 sep 2008. Llanura entre el bosque de

pino, 2425 m, D. Ramirez Noya 2995 (CIIDIR).

Adenophyllum anomalum (Canby & Rose) Sthrother. Proximo al arroyo Camarones, Los Aguajes, El Durazno, Tamazula, Dgo. 25°21 '46"N,

106°57'40"W, 11 die 2014, bosque tropical caducifolio, 809 m, D. Ramirez Noya 4275 (CIIDIR).

Adenophyllum cancellatum (Cass.) Villarreal. Camino a El Caballete, Los Aguajes, El Durazno, Tamazula, Dgo. 25°22'08"N, 106°56'52"W, 8

oct 2014, bosque de Quercus sp. 1481 m. D. Ramirez Noya 4100 (CIIDIR).

Adenophyllum porophyllum (Cav.) Hemsl., orilla de camino, Los Aguajes, El Durazno,Tamazula, Dgo. 25°22'01 "N, 106°56'26"W, 9 nov 2001,

bosque de Pinusteocote, P. lumholtziy Quercus spp., 1520 m, D. Ramirez Noya 2138 (CIIDIR).

Ageratina hederifolia (A. Gray) R.M. King & H. Rob. Cerro Alto, El Tecuan de El Durazno, Tamazula, Dgo., 25°35'20"N, 107°00'50"W, bosque

de Abies durangensis, Pinus durangensis, P. strobifrmis, 28 jun 2005, 2550 m, D. Ramirez Noya 2691 (CIIDIR). La Aguateca, El Tecuan, El

Durazno, Tamazula, Dgo., 25°35'35"N, 106°59'35"W, bosque de Pinus durangensis, P. strobiformis, P. arizonica, 2450 m, 9 abr 2013, D.

Ramirez Noya 3809 (CIIDIR).

Ageratina choricephala (B.L. Rob.) R.M. King & H. Rob. Predio La Canada. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'27"W,

8 nov 2001, bosque de Quercus sp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2075,2426 (CIIDIR).

Ageratina malacolepis (B.L. Rob.) R.M. King & H. Rob.Arroyo Camarones, Los Aguajes, El Durazno,Tamazula, Dgo., 25°21 '32"N, 106°57'45"W, 23

may 2003, bosque tropical caducifolio de Lysiloma spp ., Acacia spp., Bursera spp. y Ficus spp. 715 m, D. Ramirez Noya 2361,2792 (CIIDIR).

Ageratina palmeri (A. Gray) Gage ex B.L. Turner. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'10"N, 106°56'20"W, 15 oct 2003, bosque

de Pinus teocote, P. lumholtzi y Quercus spp., 1650 m, D.Ramlrez Noya 2478 (CIIDIR).

Ageratina pazeuarensis (Kunth) R.M. King & H. Rob. La Aguateca, El Tecuan, El Durazno, Tamazula, Dgo., 25°33'35"N, 106°59'45"W, bosque

de Pinus durangensis, P. arizonica, 3 abr 2008,2250 m, D. Ramirez Noya 2791 (CIIDIR). San Juan, El Tecuan, El Durazno, Tamazula, Dgo.,

25°34'09"N, 106°58'33"W, bosque de Pinus durangensis, P. strobiformis, 10 jun 2008, 2454 m, D. Ramirez Noya 2841 (CIIDIR).

Ageratum corymbosum Zucc. f. corymbosum. Potreritos, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21sep 2008,

bosque de pino abierto2435 m, D. Ramirez Noya 3043 (CIIDIR).

Ageratum corymbosum f. euryphyllum (B.L. Rob.) M.F. Johnson. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 13 oct

2003, bosque de Pinus teocote, P. lumholtziyQ. spp., 1520 m, D. Ramirez Noya 2433 (CIIDIR).

Alloispermum palmeri (A. Gray) C.F. Fernandez & Urbatsch. El tabacal, El Guajolote, El Tecuan, Tamazula, Dgo., 25°33'45"N, 107°04'02"W,

24 jun 2009, bosque de encino, 1600 m, D. Ramirez Noya 3120 (CIIDIR).

Ambrosia ambrosioides (Cav.) W.W. Payne. Sta Matilde, Los Aguajes, El Durazno, Tamazula, Dgo., 25°21 '21 "N, 106°57'53"W, 24 may 2003,

bosque tropical caducifolio de Lysiloma spp., Acacia spp., Bursera spp. y Ficus spp., 670 m, D. Ramirez Noya 2369 (CIIDIR).

Artemisia ludoviciana Nutt. La Aguateca, El Tecuan, El Durazno,Tamazula, Dgo., 25°33'35"N, 106°59'45"W, bosque de Pinus durangensis, P.

arizonica, 3 abr 2008,2250 m, D. Ramirez Noya 2803 (CIIDIR).

Baccharis multiflora Kunth var. herbacea McVaugh. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'08"N, 106°56'52"W, 8 oct 2014, bosque

de Pinus spp. y Quercus spp., 1481 m, D. Ramirez Noya 4098 (CIIDIR).

Baccharispteronioides DC. proximo a El Cordon, El Aguaje, El Durazno, Tamazula, Dgo., 25°21'50" N, 106°55'50"W, 10 oct 2014, bosque de

Pinus teocote, P. lumholtziy Quercus spp., 1790 m, D. Ramirez Noya 4212 (CIIDIR).

Baccharis salicifolia (Ruiz & Pav.) Pers. Cerca a San Dario-Aguajes, 25°23'16"N, 106°55'58"W, 12 oct 2003, bosque de Pinus durangensisyP.

strobiformis, 2071 m, D. RamirezNoya2406 (CIIDIR).

Barkleyanthussalicifolius (Kunth) H. Rob. & Brettell. San Juan, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'42"N, 106°58'32"W, 22 ago 2008,

bosque de Pinus durangensis, P. strobiformis, P. arizonica, 2455 m, D. Ramirez Noya 2935 (CIIDIR).

Bidens aurea (Aiton) Sherff. San Juan, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, Llanura entre el bosque

de pino, 2450 m, D. Ramirez Noya 2973,3953 (CIIDIR).

Bidens bigeloviiA. Gray. Los Aguajes, El Durazno,Tamazula, Dgo., 25°21'40"N, 106°56'40"W, 14 oct 2003, bosque tropical caducifolio, 1400

m, D. Ramirez Noya 2459,2483 (CIIDIR).

Bidens lemmoniiA. Gray. El Tecuan, Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sep 2008, bosque de pino abierto, 2425 m, D. Ramirez

Noya 3009,3015 (CIIDIR).

Bidens ostruthioides (DC.) Sch. Bip. Potreritos, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21 sep 2008, bosque de

pino abierto, 2435 m, D. Ramirez Noya 3042 (CIIDIR).

Bidens pilosa L. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'10"N, 106°56'20"W, 15 oct 2003, bosque de Pinus teocote, P. lumholtziiy

Quercus spp., 1650 m, D. Ramirez Noya 2485 (CIIDIR).

Bidens riparia Kunth. El Rincon, Los Aguajes, El Durazno,Tamazula, Dgo. 25°21'33"N, 106°57'07"W,8oct 2014, bosque tropical caducifolio,

978 m,D. Ramirez Noya 4147 (C\\D\R).

Bidens tenuisecta A. Gray. El Tecuan, Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sep 2008, bosque de pino abierto. 2425 m, D. Ramirez

Noya2996 (CIIDIR).

Brickellia diffusa (Vahl) A. Gray. Proximo a El Caballete, Los Aguajes, El Durazno, Tamazula, Dgo. 25°22'02"N, 106°56'55"W, 8 oct 2014,

bosque de Quercus spp. 1455 m, D.Ramlrez Noya 4115 (CIIDIR).

458

Journal of the Botanical Research Institute of Texas 9(2)

Tabla 1. (continued)

Brickelliofilipes B.L. Rob. Proximo a El Caballete, Los Aguajes, El Durazno,Tamazula, Dgo. 25°22'02"N, 106°56'55"W, 8 oct 2014, bosque de

Quercus sp. 1481 m, D. RamirezNoya 4116 (CIIDIR).

Brickellia oliganthes (Less.) A. Gray. Los Aguajes, El Durazno, Tamazula, Dgo., 25°21'27"N, 106°56'57"W, 14 oct 2003, bosque tropical

caducifolio, 1100 m, D. Ramirez Noya 2472{C\\D\R).

Brickellia oreithales (B.L. Rob) Shinners. San Juan, EITecuan, El Durazno,Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, Llanura en

el bosque de pino, 2450 m, D. RamirezNoya2845 (CIIDIR). El Tecuan, Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sep 2008, bosque

de pino abierto, 2425 m, D. Ramirez Noya 2997 (CIIDIR).

Brickellia secundiflora (Lag.) A. Gray, orilla de camino, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 9 nov 2001,

bosque de Pinus teocote, P. lumholtzii y Quercus spp., 1520 m, D. Ramirez Noya 21 53(CIIDIR).

Brickellia subuligera (S. Schauer) B.L. Turner. La gentilera, Los Aguajes, El Durazno, Tamazula, Dgo., 25°21 '41 "N, 106°57'08.3"W, 8 oct 2014,

bosque tropical caducifolio, 1090 m, D. Ramirez Noya 4129 (CIIDIR).

Carminatia tenuiflora DC. Predio; La Presa. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'27"W, 8 nov 2001, bosque de

Quercus sp., Magnolia sp., Burserasp., 1525 m, D. Ramirez Noya2074 (CIIDIR).

Centaurearothrockii Greenm. Corral de Piedra, EITecuan, El Durazno,Tamazula, Dgo., bosque de pino abierto. 25°34'25"N, 106°58'24"W,

21 ago 2008, 2435 m, D. Ramirez Noya 2913,2959 (CIIDIR).

Chaptaliaruncinata Kunth. Corral de Piedra, EITecuan, El Durazno,Tamazula, Dgo., bosque de pino abierto, 25°34'25"N, 106°58'24"W, 21

ago 2008, 2435 m, D. RamirezNoya2905,3984 (CIIDIR).

Chloracantha spinosa (Benth.) G.L. Nesom var. ialiscensis (McVaugh) S.D. Sundb. Sta Matilde, Los Aguajes, El Durazno, Tamazula, Dgo.,

25°21 '21 "N, 106°57'53"W, 24 may 2003, bosque tropical caducifolio de Lysiloma spp., Acacia spp., Bursera spp. y Ficus spp., 670 m, D.

Ramirez Noya 2365 (CIIDIR).

Chromolaena collina (DC.) R.M. King & H. Rob. Predio La Presa, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'27"W, 8 nov

2001, bosque de Quercus sp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2086 (CIIDIR).

Chromolaena odorata (L.) R.M. King & H. Rob. Los Aguajes, El Durazno, Tamazula, Dgo., 25°21'32"N, 106°56'46"W, 11 die 2014, bosque

tropical caducifolio, 1383 m, D. Ramirez Noya 4281 (CIIDIR).

Chromolaena ovaliflora (Hook. & Arn.) R.M. King & H. Rob. Predio La Presa, Los Aguajes, El Durazno,Tamazula, Dgo.,25°22'01 "N, 106°56'27"W,

8 nov 2001, bosque de Quercus sp., Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2084 (CIIDIR).

Cirsium ehrenbergii Sch. Bip. San Juan, El Tecuan, El Durazno,Tamazula, Dgo., 22 ago 2008, bosque de Pinus durangensis, P. strobiformis, P.

arizonica, 25°34 , 42"N, 106°58 , 32"W, 2455 m, D. Ramirez Noya 2954 (CIIDIR).

Conyza microcephala Hemsl. Cerca a San Dario-Aguajes, 25°23' 16"N, 106°55'58"W, 12 oct 2003, bosque de Pinus durangensisyP. strobiformis,

2071 m, D. Ramirez Noya 2413,3978-b (CIIDIR).

Coreocarpus arizonicus (A. Gray) S.F. Blake var. filiformis (Greenm.) S.F. Blake. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W,

10 die 2014, bosque de Pinus teocote, P. lumholtziy Quercus spp., 1520 m, D. Ramirez Noya 4263 (CIIDIR).

Coreocarpus congregatus (S.F. Blake) E.B. Sm. La Presa, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'27"W, 9 oct 2014,

bosque de Quercus spp. Magnolia sp. Bursera sp., 1525 m, D. Ramires Noya4165-c (CIIDIR).

Cosmos bipinnatus Cav. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, llanura entre el bosque

de pino, 2450m, D. Ramirez Noya s/n. El Tecuan, Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sept 2008, terreno abierto en el bosque

de pino, 2425m,

Cosmospalmeri B.L. Rob. San Juan, EITecuan, El Durazno,Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, Llanura en el bosque

de pino, 2450 m, D. Ramirez Noya 2849 (CIIDIR).

Cosmos parviflorus (Jacq.) Pers. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, Llanura en el

bosque de pino, 2450 m, D. Ramirez Noya 2850,2970 (CIIDIR).

Cosmossulphureus Cav. orilla de camino, Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 9 nov 2001, bosque de Pinus

teocote, P. lumholtziy Quercus spp., 1520 m, D. RamirezNoya2147 (CIIDIR).

Critonia hebebotryaDC. orilla de camino, Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 9 nov 2001, bosque de Pinus

teocote, P. lumholtziy Quercus spp., 1520 m, D. RamirezNoya2125,2423 (CIIDIR).

Critonia quadrangularis (DC.) R.M. King & H. Rob.Predio La Presa, Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'27"W, 8 nov

2001, bosque de Quercus spp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2333 (CIIDIR).

Critoniopsis triflosculosa (Kunth) H. Rob. Los Aguajes, El Durazno, Tamazula, Dgo., 25°21'40"N, 106°56'40"W, 30 jun 2005, bosque tropical

caducifolio, 1400 m, D. RamirezNoya2722 (CIIDIR).

Dahlia coccinea Cav. Proximo al Cordon, El Aguaje, El Durazno,Tamazula, Dgo., 25°21 '50"N, 106°55'50"W, 10 oct 2014, bosque Pinus teocote,

P. lumholtziy Quercus spp., 1790 m, D. Ramirez Noya 4206-b (CIIDIR).

Dahliasherffii P.D. Sorensen. Corral de Piedra, El Tecuan, El Durazno,Tamazula, Dgo., bosque de pino abierto, 25°34'25" 106°58'24", 21 ago

2008, 2435 m, D. Ramirez Noya 2877 (CIIDIR).

Decachaeta ovatifolia (DC.) R.M. King &H. Rob.Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 13 oct 2003, bosque de

Pinus teocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2424,2436 (CIIDIR).

Decachaetascabrella (B.L. Rob.) R.M. King & H. Rob. Predio; La Presa. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'27"W, 8

nov 2001, bosque de Quercus spp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2072 (CIIDIR).

Delilia biflora (L.) Kuntze. Los Aguajes, El Durazno, Tamazula, Dgo., 25°21'40"N, 106°56'40"W, 14 oct 2003, bosque tropical caducifolio,

1400 m, D. Ramirez Noya 2447 (CIIDIR).

Erigeron astranthoides De Jong & G.L. Nesom. Corral de Piedra, El Tecuan, El Durazno, Tamazula, Dgo., bosque de pino, 25°34'25"N,

106°58'24"W, 21 ago 2008, 2435 m, D. Ramirez Noya 2886,2910 (CIIDIR).

Ramirez Noya y Herrera Arrieta, Asteraceae en El Durazno, Tamazula, Durango, Mexico

459

Tabla 1. (continued)

Erigeron coroniglandifer G.L. Nesom. Corral de Piedra, El Tecuan, El Durazno, Tamazula, Dgo., bosque de pino abierto, 25°34'25"N,

106°58'24"W, 21 ago 2008, 2435 m, D. RamirezNoya2882 (CIIDIR). El Tecuan, Tamazula, Dgo., 25°33 , 45"N, 106°57 , 30"W, 20 sep 2008,

bosque de pino abierto, 2425 m, D. Ramirez Noya 2999 (CIIDIR).

Erigeron griseus (Greenm.) G.L. Nesom. Sn Juan, El Tecuan, Tamazula, Dgo., 25°34'30"N, 106°58'20"W, 27 jun 2005, pastoreoen bosque de

pino, 2420 m, D. Ramirez Noya 2648,2653-b (CIIDIR).

Erigeron podophyllus G.L. Nesom. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 22 ago 2008, bosque de pino ( Pinus durangensis, P.

strobiformis, P. arizonica), 25°34 , 42"N, 106°58 , 32"W, 2455 m, D. Ramirez Noya 2934 (CIIDIR).

Erigeron polycephalus (Larsen) Nesom. De amplia distribucion, encontrada en casi todas las localidades.

Erigeron seemannii (Sch. Bip.) Greene. La Aguateca El Tecuan, El Durazno,Tamazula, Dgo., 25°33'45"N, 106°00'18"W, bosque de pino, 3 abr

2008, ( Pinus durangensis, P. arizonica ), 2026 m, D. Ramirez Noya 2805 (CIIDIR).

Erigeron velutipes Hook. & Arn. 2772, 2804 (CIIDIR). Encontrada en casi todas las localidades.

Erigeron wislizeni (A. Gray) Greene. El Tecuan, Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sep 2008, bosque de pino abierto, 2425 m, D.

Ramirez Noya 2999-b (CIIDIR).

Fleischmannia sonorae (A. Gray) R.M. King & H. Rob. Predio La Presa, Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01"N, 106°56'27"W, 8

nov 2001, bosque de Quercus spp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2082,244 7 (CIIDIR).

Galinsoga parviflora Cav.orilla de camino, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 9 nov 2001, bosque de

pino-encino ( Pinus teocote, P. lumholtzi y Quercus spp.), 1520 m, D. Ramirez Noya 2119 (CIIDIR).

Galinsoga quadriradiata Ruiz& Pav. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, Llanura en el

bosque de pino, 2450 m, D. Ramirez Noya 2854 (CIIDIR).

Gamochaeta americana (Mill.) Wedd. Corral de Piedra, El Tecuan. El Durazno,Tamazula, Dgo., 25°34'05"N, 106°58'15"W, 16 oct 2013, pastizal

inducido, 2435 m, D. Ramirez Noya 3983-b (CIIDIR).

Guardiola rosei B.L. Rob. En la orilla de caminos, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 9 nov 2001, bosque

de pino-encino (Pinus teocote, P. lumholtziy Quercus spp.), 1520 m, D. Ramirez Noya 2148,2839,2952 (CIIDIR).

Heterospermapinnatum Cav. De amplia distribucion, encontrada en casi todas las localidades.

Heterotheca chihuahuana B.L. Turner. El Tecuan, El Durazno, Tamazula, Fgo., 25°33'28"N, 106°57'21"W, agostadero en el bosque de Pinus

durangensis, P. strobiformis, 22 may 2003, 2409 m, D. Ramirez Noya 2326{C\\D\R).

Hieracium fendleri Sch. Bip. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'09"N, 106°58'33"W, bosque de Pinus durangensis, P.

strobiformis, 10 jun 2008,2454 m, D. Ramirez Noya 2821 {C\\D\R).

Hieracium schultzii Fries. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'09"N, 106°58'33"W, bosque de Pinus durangensis, P. stro¬

biformis, 27 jun 2005. 2454 m, D. Ramirez Noya 2664 (CIIDIR).

Hofmeisteria schaffneri (A. Gray) R.M. King & H. Rob.Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 27 may 2002,

bosque de Pinus teocote, P. lumholtziy Quercus spp., 1520 m, D. Ramirez Noya 2160 (CIIDIR).

Hymenostephium cordatum (Hook. & Arn.) S.F. Blake. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 7 nov 2001, bosque

de Pinus teocote, P. lumholtziy Quercus spp., 1520 m, D. Ramirez Noya 2067 (CIIDIR).

lostephane heterophylla (Cav.) Benth. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, Llanura en

el bosque de pino, 2450 m, D. Ramirez Noya 2968 (CIIDIR).

Jaegueria hirta (Lag.) Less. Sta Matilde, Los Aguajes, El Durazno, Tamazula, Dgo., 25°21'21"N, 106°57'53"W, 24 may 2003, bosque tropical

caducifolio conLysilomaspp.,Acaciaspp., Bursera spp.y Ficus spp., 670 m, D. Ramirez Noya 2363 (CIIDIR). Potreritos, El Tecuan, El Durazno,

Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21 sep 2008, bosque de pino abierto, 2435 m, D. Ramirez Noya 3020 (CIIDIR).

Jaegueriapurpurascens Rob. EITecuan,Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sep 2008, bosque de pino abierto, 2425 m, D. Ramirez

Noya 3012 (CIIDIR).

Laenecia confusa (Cronquist) G.L. Nesom. El Aguaje, El Durazno,Tamazula, Dgo., 25°21 '36.8"N, 106°56'21.8"W. 9 oct 2014, bosque tropical

caducifolio con Lippia umbellata, Montanoa leucantha e Ipomoea arborescens, 1509 m, D. Ramirez Noya 4199 (CIIDIR).

Laennecia gnaphalioides (Kunth) Cass, San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'05"N, 106°57'30"W, 15 oct 2013, terreno de

cultivo, 2410 m, D. Ramirez Noya 3978 c (CIIDIR).

Laennecia schiedeana (Less.) G.L.Nesom. Cerca a San Dario-Aguajes, 25°23'16"N, 106°55'58"W, 12 oct 2003, bosque de pino {Pinus duran¬

gensis yP. strobiformis ), 2071 m, D. RamirezNoya2414 (CIIDIR).

Laenneciasophiifolia (Kunth) G.L. Nesom. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 15 oct 2003, bosque de Pinus

teocote, P. lumholtzii y Quercus spp., 1650 m, D. Ramirez Noya 2487(01 DIR).

Lagascea decipiens Hemsl. var. decipiens Los Aguajes, El Durazno, Tamazula, Dgo., 25°21'35"N, 106°56'42"W, 14 oct 2003, bosque tropical

caducifolio, 1200 m, D. Ramirez Noya 2461 (CIIDIR).

Lasianthaea ceanothifolia (Willd.) K.M. Becker var. gracilis (W.W. Jones) K.M. Becker.Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01 "N,

106°56'26"W, 7 nov 2001, bosque de Pinus teocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2064 (CIIDIR).

Lasianthaea fruticosa (L.) K.M. Becker var. fasciculata (DC.) K.M. Becker. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W,

13 oct 2003, bosque de Pinus teocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2438 (CIIDIR).

Lasianthaea fruticosa (L.) K.M. Becker var. michoacana (S.F. Blake) K.M. Becker. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N,

106°56'26"W, 13 oct 2003, bosque de Pinus teocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2430 (CIIDIR).

Lasianthaea seemannii (A. Gray) K.M. Becker. Predio La Presa. Los Aguajes, El Durazno, Tamazula, Dgo.,25°22'01"N, 106°56'27"W, 8 nov

2001, bosque de Quercus spp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2096 (CIIDIR).

Leibnitzia lyrata (Sch. Bip.) G.L. Nesom. Corral de Piedra, El Tecuan, El Durazno, Tamazula, Dgo., bosque de pino abierto, 25°34'25"N,

106°58'24"W, 21 ago 2008, 2435 m, D. Ramirez Noya 2866,3011 (CIIDIR).

460

Journal of the Botanical Research Institute of Texas 9(2)

Tabla 1. (continued)

Melompodium bibrocteotum S. Wats. Potreritos, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'17"N, 106°58'30"W,21 sep 2008, bosque de

pino abierto, 2435 m, D. RamirezNoya 3021 (CIIDIR).

Melampodium perfoliatum (Cav.) Kunth. Potreritos, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21 sep 2008, bosque

de pino abierto, 2435 m, D. RamirezNoya 3049 (CIIDIR).

Montanoaleucantha (Lag.) S.F. Blake var. arborescens (DC.) B.L.Turner. Predio; La Presa. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N,

106°56'27"W, 8 nov 2001, bosque de Quercus sp.. Magnolia sp., Bursera sp., 1525 m, D. RamirezNoya2092,2471 (CIIDIR).

Packera bellidifolia (Kunth) W.A. Weber & A. Love San Juan, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'09"N, 106°58'33"W, bosque de

Pinus durangensis, P. strobiformis ), 10 jun 2008,2454 m, D. RamirezNoya2827 (CIIDIR).

Packera candidissima (Greene) W.A. Weber & A. Love. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'09"N, 106°58'33"W, bosque

de Pinus durangensis, P. strobiformis, 10 jun 2008,2454 m, D. Ramirez Noya 2826,2865 (CIIDIR).

Packerascalaris (Greene) C. Jeffrey. Corral de Piedra, El Tecuan, El Durazno,Tamazula, Dgo., bosque de pino abierto, 25°34'25"N, 106°58'24"W,

21 ago 2008, 2435 m, D. Ramirez Noya 2909 (CIIDIR). Potreritos, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21

sep 2008, bosque de pino abierto, 2435 m, D. Ramirez Noya 3038 (CIIDIR).

Pectisprostrata Cav. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 13 oct 2003, bosque de Pinus teocote, P. lumholtzi

y Quercus spp., 1520 m, D. RamirezNoya2435 (CIIDIR).

Perityle microglossa Benth. var. saxosa (Brandegee) A.M. Powell., orilla de camino, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01 "N,

106°56'26"W, 9 nov 2001, bosque de Pinusteocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2115 (CIIDIR). Arroyo Cama rones,

Los Aguajes, El Durazno,Tamazula, Dgo., 25°21 '40"N, 106°57'41 "W, 28 may 2002, bosque tropical caducifolio con Lysiloma spp., Acacia

spp., Bursera spp. y Ficus spp. 710 m, D. Ramirez Noya 2203 (CIIDIR).

Perymenium pringlei B.L. Rob. & Greenm. var .pringlei Los Aguajes, El Durazno, Tamazula, Dgo., 25°21'40"N, 106°56'40"W, 14 oct 2003,

bosque tropical caducifolio, 1400 m, D. Ramirez Noya 2450 (CIIDIR).

Perymenium reticulatum JJ. Fay. Los Aguajes, El Durazno,Tamazula, Dgo., 25°21 '40"N, 106°56'40"W, 14 oct 2003, bosque tropical caducifolio

(ecotono), 1400 m, D. Ramirez Noya 2450 (CIIDIR).

Pinaropappusjunceus A. Gray. El Cordon, El Aguaje, El Durazno,Tamazula, Dgo., 25°21'44.5"N, 106°55'42.1"W.9oct 2014, bosque de Pinus

teocote, P. lumholtzi y Quercus spp., 1850 m, D. Ramirez Noya 4189 (CIIDIR).

Pinaropappus roseus. (Less.) Less. San Juan, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'09"N, 106°58'33"W, bosque de Pinus durangensis,

P. strobiformis, lOjun 2008, 2454 m, D. Ramirez Noya 2819,2831 (CIIDIR).

Pippenalia delphinifolia (Rydb.) McVaugh. El Tecuan,Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sep 2008, bosque de pino abierto, 2425

m, D. Ramirez Noya 3017 (CIIDIR).

Porophyllum linaria (Cav.) DC. Proximo a El Cordon, El Aguaje, El Durazno,Tamazula, Dgo., 25°21'50"N, 106°55'50"W, 10 oct 2014, bosque

de Pinus teocote, P. lumholtzi y Quercus spp., 1790 m, D. Ramirez Noya 4213 (CIIDIR).

Porophyllum macrocephalum DC., orilla de camino, Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 9 nov 2001, bosque

de Pinus teocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2150 (CIIDIR).

Porophyllum pringlei B.L. Rob.La Gentilera, Los Aguajes, El Durazno,Tamazula, Dgo., 25°21 '41 "N, 106°57'08.3"W, 8 oct 2014, bosque tropical

caducifolio, 1090 m, D. Ramirez Noya 4130 (CIIDIR).

Psacalium globosum (B.L. Rob. & Fernald) H. Rob. & Brettell. Corral de Piedra, El Tecuan, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W,

16 oct 2013, pastizal inducido, 2435 m, D. Ramirez Noya 3995 (CIIDIR).

Psacalium sinuatum (Cerv.) H. Rob. & Brettell. Corral de Piedra, El Tecuan, El Durazno,Tamazula, Dgo., bosque de pino abierto, 25°34'25"N,

106°58'24"W, 21 ago 2008, 2435 m, D. Ramirez Noya 2887 (CIIDIR).

Pseudelephantopus spicatus (Juss. ex Aubl.) Rohr. Sta Matilde, Los Aguajes, El Durazno,Tamazula, Dgo., 25°21'21"N, 106°57'53"W, 24 may

2003, bosque tropical caducifolio con Lysiloma spp.. Acacia spp., Bursera spp. y Ficus spp., 670 m, D. Ramirez Noya 2366,2470 (CIIDIR).

Pseudognaphalium oxyphyllum (DC.) Kirb.EI Aguaje, El Durazno,Tamazula, Dgo., 25°21'36.8"N, 106°56'21.8"W, 9 oct 2014, bosque tropical

caducifolio con Lippia umbellata, Montnoa leucantha e Ipomoea arborescens, 1509 m, D. Ramirez Noya 4192 (CIIDIR).

Psedognaphalium semiamplexicaule (DC.) Anderb. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 9 oct 2014, bosque

de Pinus teocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 4161 (CIIDIR).

Ratibida mexicana {S.\Natson)\N.M.Sharp.San Juan, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'45"N, 106°58'20"W,21 ago 2008, Llanura

en el bosque de pino, 2450 m, D. Ramirez Noya 2847 (CIIDIR).

Roldanachapalensis (S. Watson) H. Rob. & Brettell. Predio La Manga, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'10"N 106°57'32", 15 oct

2013, bosque de Pinus cooperi, P. leiophylla y P. teocote, 2420 m, D. Ramirez Noya 3979 (CIIDIR).

Roldana hartwegii (Benth.) H. Rob. & Brettell. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'10"N, 106°56'20"W, 15 oct 2003, bosque de

Pinus teocote, P. lumholtzi y Quercus spp., 1650 m, D. Ramirez Noya 2486 (CIIDIR).

Schkuhriapinnata (Lam.) Kuntze var. wislizenii (A. Gray) B.L.Turner. San Juan, El Tecuan, El Durazno,Tamazula, Dgo., 25°34'45"N, 106°58'20"W,

21 ago 2008, Llanura en el bosque de pino, 2450 m, D. Ramirez Noya 2852 (CIIDIR).

Sclerocarpussessilifolius Greenm. El Aguaje, El Durazno,Tamazula, Dgo., 25°21 '36.8"N, 106°56'21.8"W, 9 oct 2014, bosque tropical caducifolio

con Lippia umbellata, Montnoa leucantha e Ipomoea arborescens, 1509 m, D.RamirezNoya 4193-b (CIIDIR).

Senecio stoechadiformis DC. La Aguateca El Tecuan , El Durazno, Tamazula, Dgo., 25°33'35"N, 106°59'45"W, bosque de Pinus durangensis,

P. arizonica, 3 abr 2008, 2250 m, D. Ramirez Noya 2794,2797 (CIIDIR).

Sigesbeckia jorullensis Kunth. Potreritos, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21 sep 2008, bosque de pino

abierto, 2435 m, D. Ramirez Noya 2048,3060 (CIIDIR).

Simsia amplexicaulis (Cav.) Pers.Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 27 may 2002, bosque de Pinus teocote,

P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2165 (CIIDIR). De amplia distribucion, encontrada en casi todas las localidades.

Ramirez Noya y Herrera Arrieta, Asteraceae en El Durazno, Tamazula, Durango, Mexico

461

Tabla 1. (continued)

Sinclairia palmeri [ A. Gray) B.L.Turner. Potreritos, EITecuan, El Durazno,Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21 sep 2008, bosque de

pino abierto, 2435 m, D. RamirezNoyo 3034b (CIIDIR).

Stenocarpha filiformis (Hemsl.) S.F. Blake. Cerca a San Dario-Aguajes, 25°23'16"N, 106°55'58"W, 12 oct 2003, bosque de Pinus durangensis

y P. strobiformis, 2071 m, D. Ramirez Noya 2417 (CIIDIR).

Stevia aschenborniana Sch. Bip. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 10 die 2014, bosque de pino Pinus

teocote, P. lumholtziy Quercus spp., 1520 m, D. Ramirez Noya 4275 (CIIDIR).

Steviajorullensis Kunth. Potreritos, EITecuan, El Durazno,Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21 sep 2008, bosque de pino abierto,

2435 m, D. Ramirez Noya 3063 (CIIDIR).

Stevia micradenia B.L. Rob. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22' 01 "N, 106°56'26"W, 9 oct 2014, bosque de Pinus teocote, P.

lumholtziy Quercus spp., 1650 m, D. Ramirez Noya 4187 (CIIDIR).

Stevia myricoides McVaugh.Aguazul, El Tecuan, Tamazula, Dgo., 25°33'40"N, 107°01'10"W, 3 abr 2008, bosque de Quercus spp., 1700 m,

D. Ramirez Noya 2802 (CIIDIR).

Stevia origanoides Kunth. Cerca a San Dario-Aguajes, 25°23'16"N, 106°55'58"W, 12 oct 2003, bosque de Pinus durangensisyP. strobiformis,

2071 m, D. Ramirez Noya 2419 (Cl I Dl R).

Stevia ovata Willd. Predio; La Presa. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'27"W, 8 nov 2001, bosque de Quercus

sp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2081,3002-b (CIIDIR).

Steviapurpusii B.L. Rob. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 9 oct 2014, bosque de Pinus teocote, P. lumholtzi

y Quercus spp., 1650 m, D. Ramirez Noya 4162 (CIIDIR).

Steviaserrata Cav. Corral de Piedra, El Tecuan, El Durazno,Tamazula, Dgo., bosque de pino abierto, 25°34'25"N, 106°58'24"W, 21 ago 2008,

2435 m, D. Ramirez Noya 2893,3002 (CIIDIR).

Stevia trifida Lag. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 22 may 2003, bosque de Pinus teocote, P. lumholtziy

Quercus spp., 1520 m, D. Ramirez Noya 2331,3955 (CIIDIR).

Symphyotrichum expansum (Poepp. ex Spreng.) G.L. Nesom, orilla de camino, Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N,

106°56'26"W, 9 nov 2001, bosque de Pinus teocote, P. lumholtziy Quercus spp., 1520 m, D. Ramirez Noya 2122,3955-b (CIIDIR).

Symphyotrichum potosinum (A. Gray) G.L. Nesom. Corral de Piedra, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'05"N, 106°58'15"W, 16

oct 2013, acuatica, creciendo dentro del arroyo permanente, 2435 m, D. Ramirez Noya 4001 (CIIDIR).

Tagetes filifolia Lag. Corral de Piedra, El Tecuan, El Durazno, Tamazula, Dgo., bosque de pino abierto, 25°34'25"N, 106°58'24"W, 21 ago

2008, 2435 m, D. Ramirez Noya 2862,2891 (CIIDIR).

Tagetes foetidissima DC.Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01"N, 106°56'26"W, 27 may 2002, bosque de Pinus teocote, P.

lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2168 (CIIDIR).

Tagetes micrantha Cav. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 9 oct 2014, bosque de Pinus teocote, P. lumholtzi

y Quercus spp., 1520 m, D. Ramirez Noya 4158-b (CIIDIR). De amplia distribucion, encontrada en casi todas las localidades.

Tagetes palmeri A. Gray. Cerca a San Dario-Aguajes, 25°23'16"N, 106°55'58"W, 7 nov 2001, bosque de Pinus durangensis y P. strobiformis,

2071 m, D. Ramirez Noya 2054 (Cl I Dl R).

Tagetes pringlei S. Watson. Corral de Piedra, El Tecuan, El Durazno, Tamazula, Dgo., dentro del arroyo, 25°34'25"N, 106°58'24"W, 21 ago

2008, 2435 m, D. Ramirez Noya 2906 (CIIDIR).

Tagetes subulata Cerv. Cerca a San Dario-Aguajes, 25°23'16"N, 106°55'58"W, 12 oct 2003, bosque de Pinus durangensis y P. strobiformis,

2071 m, D. Ramirez Noya 2418 (Cl I Dl R).

TithoniafruticosaS. Canby &Rose. Los Aguajes, El Durazno,Tamazula, Dgo., 25°21'40"N, 106°56'40"W, 30jun 2005, bosque tropical cadu-

cifolio, 1400 m, D. Ramirez Noya 2721 (CIIDIR).

Tithonia tubiformis (Jacq.) Cass. El Aguaje, El Durazno,Tamazula, Dgo., 25°21 '36.8"N, 106°56'21.8"W, 9 oct 2014, bosque tropical caducifolio

conLippiaumbellata,Montanoaleucanthaelpomoeaarborescens,lS09 m,D. Ramirez Noya 4191 (CIIDIR). Especieescasa en la localidad.

Tridaxmexicana AM. Powell. La Presa. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'27"W, 8 nov 2001, bosque de Quercus

spp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2170 (CIIDIR).

Trigonospermum annum McVaugh & Lask. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'08"N, 106°56'52"W, 8 oct 2014, bosque de

Quercus spp., 1481 m, D. Ramirez Noya 4099 (CIIDIR).

Verbesina longifolia (A. Gray) A. Gray. Potreritos, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'17"N, 106°58'30"W, 21 sep 2008, bosque de

pino abierto, 2435 m, D. Ramirez Noya 3030 (CIIDIR).

Verbesina oncophora B.L.Rob & Seat. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 22 ago 2008, bosque de Pinus durangensis, P. stro¬

biformis, P. arizonica, 25°34'42"N, 106°58'32"W, 2455 m, D. Ramirez Noya 2930 (CIIDIR).

Verbesina parviflora (Kunth) S.F. Blake var. parviflora. San Juan, El Tecuan, El Durazno, Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago

2008, Llanura en el bosque de pino, 2450 m, D. Ramirez Noya 2846,2972 (CIIDIR).

Vernonanthura liatroides (DC.) H. Rob. La Aguateca El Tecuan, El Durazno, Tamazula, Dgo., 25°33'35"N, 106°59'45"W, bosque de Pinus

durangensis, P. arizonica, 3 abr 2008,2250 m, D. Ramirez Noya 2790,2801 (CIIDIR).

Viguiera dentata (Cav.) Spreng. Los Aguajes, El Durazno, Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 13 oct 2003, bosque de Pinus teocote,

P. lumholtziy Quercus spp., 1520 m, D. Ramirez Noya 2425,3952 (CIIDIR).

Xanthismastenolobum (Greene) D.R. Morgan & R.L. Hartm. San Juan, EITecuan, El Durazno,Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21

ago 2008, Llanura en el bosque de pino, 2450 m, D. Ramirez Noya 2966 (CIIDIR).

Xanthocephalum benthamianum Hemsl. San Juan, EITecuan, El Durazno,Tamazula, Dgo., 25°34'45"N, 106°58'20"W, 21 ago 2008, Llanura

en el bosque de pino, 2450 m, D. Ramirez Noya 2967,3951 (CIIDIR).

462

Journal of the Botanical Research Institute of Texas 9(2)

Tabla 1. (continued)

Zinnia americana (Mill.) Olorode & A.M.Torres. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01 "N, 106°56'26"W, 13 oct 2003, bosque de

Pinus teocote, P. lumholtziy Quercus spp., 1520 m, D. RamirezNoya2437 (CIIDIR). Los Aguajes, El Durazno,Tamazula, Dgo., 25°21 '27"N,

106°56'57"W, 14 oct 2003, bosque tropical caducifolio, 1100 m, D. Ramirez Noya 2467 (CIIDIR).

Zinnia angustifolia Kunth var. angustifolia. Predio La Presa. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01"N, 106°56'27"W,8 Nov 2001,

bosque de Quercus spp.. Magnolia sp., Bursera sp., 1525 m, D. Ramirez Noya 2078 (CIIDIR).

Zinnia angustifolia Kunth var. greggii (B.L. Rob & Greenm.) McVaugh. Los Aguajes, El Durazno,Tamazula, Dgo., 25°22'01"N, 106°56'26"W,

7 Nov 2001, bosque de Pinus teocote, P. lumholtzi y Quercus spp., 1520 m, D. Ramirez Noya 2066 (CIIDIR).

Zinnia peruviana (L.) L. Los Aguajes, El Durazno,Tamazula, Dgo., 25°21 '45"N, 106°56'30.11 "W, 9 oct 2014, bosque tropical caducifolio con

Lippia umbellata, Montanoa leucantha e Ipomoea arborescens, 1520 m, D. Ramirez Noya 2147 (CIIDIR).

Zinnia tenuis (S. Watson) Strother. El Tecuan, Tamazula, Dgo., 25°33'45"N, 106°57'30"W, 20 sep 2008, bosque de pino abierto, 2425 m, D.

Ramirez Noya 3010,3016 (CIIDIR).

METODOLOGIA

El presente trabajo se realizo de forma paralela, al colaborar el primer autor (D. Ramirez Noya) con otros

proyectos no botanicos, llevados a cabo en el CIIDIR Durango. El proposito inicial fue realizar un registro de

las especies de Asteraceas que pudieran significar ser elementos novedosos para la entidad, omitiendo la re-

colecta y registro de aquellas especies comunes por su amplia distribucion geografica y recolectando solo 2 a 3

ejemplares de herbario de las especies de distribucion restringida. La idea de integrar un inventario de las as¬

teraceas observadas durante los recorridos, surgio posteriormente, razon por la cual las recolectas de ejem¬

plares no fueron planeadas para ser abundantes y se reconocen las carencias del inventario realizado, sin que

por ello se quiera dejar de reportar los hallazgos registrados.

Los ejemplares recolectados se depositaron en el Herbario CIIDIR, donde fueron identificados crltica-

mente, utilizando floras, monograflas y revisiones (Bremer y Humphries 1993; King & Robinson 1987;

McVaugh 1984; Nesom 1989 a y b, 1992; Sorensen 1969, Strother 1969, 1986; Torres 1964; Turner 1985 a y b,

1996, 1997, entre otros mas). Los ejemplares identificados se cotejaron con las descripciones que proporciono

la bibliografla consultada y posteriormente se compararon con especlmenes del herbario (CIIDIR), as! como

llevando a cabo la consulta de imagenes disponibles en las paginas web:http://www.tropicos.org/; http://www.

ipni.org/ipni/plantnamesearchpage.do; http://www.rad.unam.mx/index.php/search/; http://swbiodiversity.

org/seinet/checklists/; http://www.madrean.org/symbflora/taxa/index. Personal del herbario CIIDIR distri-

buira los duplicados a otros herbarios, una vez terminado el proceso de herborizacion del material. Es impor-

tante mencionar que antes de las recolectas del presente estudio, muchas de las especies que se citan para la

Sierra Madre en Durango y que se incluyen en el presente reporte, no contaban con ejemplares de respaldo en

la coleccion del herbario CIIDIR.

RESULTADOS Y DISCUSION

Se encontraron y se reportan aqul 75 generos y 149 especies de la familia Asteraceae en las localidades

muestreadas de la region de El Durazno y cercanlas, Tamazula, Durango (Tabla 1), 17 de estas especies y una

variedad ( Lagascea decipiens var. decipiens) son nuevos registros para Durango (Tabla 2). En la Tabla 3 se senala

la distribucion de las especies encontradas en los diferentes tipos de vegetacion de la region muestreada, las

especies sinantropicas (Villasenor 2012), la distribucion geografica general de los taxa encontrados, asi como

las especies de distribucion geografica restringida a Mexico (Rzedowski 1991a). En un intento de jerarquizar la

magnitud del endemismo en Mexico se consideran aqul dos niveles: a) especies presentes en mas de 5 estados

de la republica, son endemicas de Mexico, yb) especies presentes en uno a cuatro estados de la Republica Mexi-

cana, son endemismos regionales.

De los nuevos registros es interesante observar que especies conocidas de una sola localidad en: Micho-

acan, Baccharis multiflora var. herbacea, Sinaloa, Coreocarpus congregatus y Estado de Mexico, Perymenium re-

ticulatum son ahora registradas en Durango, lo que amplia su area de distribucion conocida a una region muy

alejada, mientras que especies como Coreocarpus arizonicus var. filiformis y Pinaropappus junceus, que eran

Ramirez Noya y Herrera Arrieta, Asteraceae en El Duraznojamazula, Durango, Mexico

463

Tabla 2. Nuevos registros para el estado de Durango, Mexico

Especies Distribution conocida

Agerotum corymbosum Zucc. f. euryphyllum (B.L. Rob.) M.F. Johnson

Bocchoris multiflora Kunth var. herbacea McVaugh

Brickellia filipes B.L. Rob.

Chromolaena odorata (L.) R.M. King & H. Rob.

Coreocarpus arizonicus (A. Gray) S.F. Blake var. filiformis

(Greenm.) S.F. Blake

Coreocarpus congregatus (S.F. Blake) E.B. Sm.

Gamochaeta americana (Mill.) Wedd.

Hymenostephium cordatum (Hook. & Arn.) S.F. Blake

Lagascea decipiens Hemsl. var. decipiens

Lasianthaea fruticosa (L.) K.M. Becker var. michoacana (DC.) K.M. Becker

Perymenium reticulatum JJ. Fay

Pinaropappus junceus A. Gray

Porophyllum pringlei B.L. Rob.

Pseudognaphalium oxyphyllum (DC.) Kirp.

Pseudognaphalium semiamplexicaule (DC.) Anderb.

Roldana chapalensis (S. Watson) H. Rob. & Brettell.

Stevia aschenborniana Sch. Bip.

Stevia purpusii B.L. Rob.

Ags., Gto., Hgo., Jal., Mor., Qro., S.L.P., Zac.

Mich.

Col., Gro., Jal., Mich., Oax.

America tropical; Africa; Viet Nam; c-s de Mex.

Chi., Sin., Son.

Sin.

Am trop.; Chis., D.F., Gro., Hgo., Jal., Mex., Mich., Oax.,

Pue., Sin., Ver., Zac.

Chis., Hgo., Mex., N.L., S.L.P., Sin., Ver.; CAm

S Arizona, Chih., Gro., Mich., Sin., Son., Oax.

Gro., Jal., Mex., Mich., Mor.

Mex.

Chih., Sin., Son.

Chis., Gro., Jal., Mex., Mich., Mor., Oax., Sin.

Ags., D.F.,Gto., Hgo., Jal., Mor., Pue., Qro., Ver., Zac.

Chis., Mich., Pue., Tams.; CAm

Ags., Col., Gto., Gro., Jal., Mex., Mich., Mor., Oax., Sin., Zac.

Chih., Gro., Jal., Mich., Nay., Sin.

Ags., D.F., Gto., Gro., Hgo., Mex., Mich., Mor., Oax., Pue., Qro.

Regiones: Am= America, CAm= Centroamerica, NAm= Norteamerica, SAm= Sudamerica. Pais : Mex= Mexico. Estados: Chis= Chiapas,

Ch= Chihuahua, Co= Coahuila, D= Durango, G= Guerrero, J= Jalisco, Mi= Michoacan, N= Nayarit, 0= Oaxaca, SLP= San Luis Potosi, Si=

Sinaloa, So= Sonora. Ac= Acuatica.

conocidas de Estados colindantes con Durango (Chihuahua, Sinaloa y Sonora), se registran ahora en el estado.

Las otras 12 especies incluidas en la Tabla 2 se distribuyen en regiones confermadas por mas de 5 Estados y en

muchas de ellas bastante alejados de nuestra region de estudio, lo que representa su registro una importante

ampiacion de su area de distribucion conocida.

Otras especies interesantes de mencionar son aquellas cuya distribucion conocida es muy cercana a la

region de nuestro estudio; por ejemplo Lagasca decipiens var. glandulosa es reportada de Sonora y Sinaloa

(Stuessy 1978), recolectada tambien por D. Ramirez N. de Durango (en San Juan de Camarones, Santiago Pa-

pasquiaro, a 24°55'37"N, 106°24'49"W, 604 m de altitud); mientras que Lagasca decipiens var. decipiens re¬

portada del sur de Arizona, Sonora, Chihuahua, Sinaloa, Michoacan, Guerrero y Oaxaca (McVaugh 1984), es

considerada un caso inusual de distribucion bicentrica, por la carencia de ejemplares provenientes de Nayarit

y Durango. Aqul la reportamos por primera vez de Durango, creciendo en Los Aguajes, El Durazno, Tamazula,

Dgo., a 25 0 21'35" 106°56'42", 1200 m de altitud. La diferencia que presentan los ejemplares no dejan lugar a

dudas de su identihcacion, ya que Lagascea decipiens var. decipiens presenta pedunculos pilosos y no posee las

glandulas estipitadas que si presenta la variedad glandulosa (McVaugh 1984). Esto no concuerda con el posible

bicentrismo propuesto por McVaugh, ampliandose la distribucion geograhca conocida de esta variedad. Otro

dato a observar es la diferencia de altitudes en las que se distribuyen las dos variedades: la variedad glandulosa

a unos 604 m y la variedad tlpica por encima de los 1200 m.

Las preferencias de habitat senalan que el mayor numero de especies de Asteraceas en esta region se dis¬

tribuyen en el Bosque de Conlferas (49.34%), en altitudes de 2050 a 2550 m. Le siguen en importancia las que

habitan en el Bosque de Pino- Encino (39.47%), en altitudes de 1500 a 1900 m y el menor numero se encuentra

en el Bosque Tropical Caducifolio (22.37%), en altitudes de 670 a 1000 m. Resultados similares son reportados

por Suarez y Villasenor (2011) para el Estado de Oaxaca, quienes observaron que la riqueza de especies de as¬

teraceas es mucho mayor en los bosques templados que en los bosques tropicales humedos de Oaxaca. Aun

cuando los ecosistemas de Durango son mas secos (bosques templados sub-humedos y bosque tropical

464

Journal of the Botanical Research Institute of Texas 9(2)

Tabla 3. Listado floristico de las Asteraceas en la region del Durazno y cercamas,Tamazula, Durango, Mexico. Bosque de Comferas= B Comferas, Bosque de Quercus-

Pinus = BQ-P, Bosque Tropical Caducifolio = BTC.

ESPECIES

Coni-

feras

Sinan-

tropica

Distribution

Americana

Endemicas

de:

Distribu¬

tion mas

amplia

Acmella radicans (Jacq.) R.K. Jansen

SAm

Acmella repens (Walter) Rich.

SAm

Acourtia macrocephala Sch. Bip.

NAm

Adenophyllum anomalum (Canby & Rose) Strother

Mex

Adenophyllum cancellatum (Cass.) Villarreal

D,Si

Adenophyllum porophyllum (Cav.) Hemsl.

CAm

Ageratina hederifolia (A. Gray) R.M. King & H. Rob.

D,SLP,Ch

Ageratina choricephala (B.L. Rob.) R.M. King & H. Rob.

Mex

Ageratina malacolepis (B.L. Rob.) R.M. King & H. Rob.

Mex

Ageratinapalmeri (A. Gray) Gage ex B.L. Turner

Mex

Ageratinapazcuarensis (Kunth) R.M. King & H. Rob.

CAm

Ageratum corymbosum Zucc. f. corymbosum

CAm

Ageratum corymbosum f. euryphyllum (B.L. Rob.)

Mex

M.F. Johnson

Alloispermum palmeri (A. Gray) C.F. Fernandez & Urbatsch

D,J

Ambrosia ambrosioides (Cav.) W.W. Payne

Nam

Artemisia ludoviciana Nutt.

NAm

Baccharis multi flora var. herbacea McVaugh

Baccharis pteronioides DC.

NAm

Baccharis salicifolia (Ruiz & Pav.) Pers.

SAm

Barkleyanthussalicifolius (Kunth) H. Rob. & Brettell

NAm

Bidens aurea (Aiton) Sherff

NAm

Bidens bigelovii A. Gray

NAm

Bidens lemmoniiA. Gray

NAm

Bidens ostruthioides (DC.) Sch. Bip.

Bidens pilosa L.

Bidens riparia Kunth (Semiacuatica)

SAm

Bidens tenuisecta A. Gray

NAm

Brickellia diffusa (Vahl) A. Gray

SAm

Brickellia filipes B.L. Rob.

Mex

Brickellia oliganthes (Less.) A. Gray

CAm

Brickellia oreithales (B.L. Rob.) Shinners

Mex

Brickellia secundiflora (Lag.) A. Gray

Mex

Brickellia subuligera (S. Schauer) B.L. Turner

Mex

Carminatia tenuiflora DC.

NAm

Centaurea rothrockii Greenm.

NAm

Chaptalia runcinata Kunth

SAm

Chloracantha spinosa var .jaliscensis (McVaugh)

Mex

S.D. Sundb.

Chromolaena collina (DC.) R.M. King & H. Rob.

NAm

Chromolaena odorata (L.) R.M. King & H. Rob.

Chromolaena ovaliflora (Hook. & Arn.) R.M. King

Mex

& H. Rob.

Cirsium ehrenbergii Sch. Bip.

Mex

Conyza microcephala Hemsl.

CAm

Coreocarpus arizonicus (A. Gray) S.F. Blake var.

NAm

filiformis (Greenm.) S.F. Blake

Coreocarpus congregatus (S.F. Blake) E.B. Sm.

Cosmos bipinnatus Cav.

Cosmos palmeri B.L. Rob.

Ch,D

Cosmos parviflorus (Jacq.) Pers.

NAm

Cosmos sulphureus Cav.

Critonia hebebotrya DC.

CAm

Critonia quadrangularis (DC.) R.M. King & H. Rob.

CAm

Critoniopsis triflosculosa (Kunth) H. Rob.

CAm

Dahlia coccinea Cav.

SAm

Dahlia sherffii P.D. Sorensen

Ch,D,Z

Ramirez Noya y Herrera Arrieta, Asteraceae en El Duraznojamazula, Durango, Mexico

465

Tabla 3. (continued)

ESPECIES

Coni-

feras

Sinan-

tropica

Distribution

Americana

Endemicas

de:

Distribu¬

tion mas

amplia

Decachaeta ovatifolia (DC.) R.M. King & H. Rob.

Mex

Decachaeta scabrella (B.L. Rob.) R.M. King & H. Rob.

Mex

Delilia biflora (L.) Kuntze

SAm

Erigeron astranthoides De Jong & Nesom

Erigeron coroniglandifer G.L. Nesom

Ch,D

Erigeron griseus (Greenm.) Nesom

D,N

Erigeron podophyllus G.L Nesom

Ch,D

Erigeronpolycephalus (Larsen) G.L. Nesom

D,J,Chis

Erigeron seemannii (Sch. Bip.) Greene

D,Ch

Erigeron velutipes Hook. & Arn.

NAm

Erigeron wislizeni (A. Gray) Greene

Ch,D

Fleischmannia sonorae (A. Gray) R.M. King & H. Rob.

CAm

Galinsoga parviflora Cav.

Galinsoga quadriradiata Ruiz & Pav.

Gamochaeta americana (Mill.) Wedd.

Guardiola rosei B.L. Rob.

Ch,D,N

Heterosperma pinnatum Cav.

Heterotheca chihuahuana B.L. Turner

Ch,D

Hieracium fendleri Sch. Bip.

NAm

Hieracium schultzii Fr.

NAm

Hofmeisteria schaffneri (A. Gray) R.M. King & H. Rob.

Mex

Hymenostephium cordatum (Hook. & Arn.) S.F. Blake

CAm

lostephane heterophylla (Cav.) Benth.

Mex

Jaegueria hirta (Lag.) Less. (Acuatica)

SAm

Jaegueria purpurascens B.L. Rob.

Mex

Laennecia confusa (Cronquist) G.L. Nesom

CAm

Laennecia gnaphalioides (Kunth) Cass

SAm

Laennecia schiedeana (Less.) G.L. Nesom

NAm

Laennecia sophiifolia (Kunth) G.L. Nesom

SAm

Lagascea decipiens Hemsl. var. decipiens

NAm

Lasianthaea ceanothifolia (Willd.) K.M. Becker var.

Mex

gracilis (W.W. Jones) K.M. Becker

Lasianthaea fruticosa (L.) K.M. Becker var.

CAm

fasciculata (DC.) K.M. Becker

Lasianthaea fruticosa (L.) K.M. Becker var.

G,Mi

michoacana (S.F. Blake) K.M. Becker

Lasianthaea seemannii (A. Gray) K.M. Becker

So,D,N

Leibnitzia lyrata (Sch. Bip.) G.L. Nesom

CAm

Melampodium bibracteatum S. Watson

CAm

Melampodium perfoliatum (Cav.) Kunth

NAm

Montanoa leucantha var. arborescens (DC.) B.L. Turner

Mex

Packera bellidifolia (Kunth) W.A. Weber & A. Love

Mex

Packera candidissima (Greene) W.A. Weber & A. Love

Ch,D

Packera sea laris (Greene) C. Jeffrey

Ch,D,Co

Pectis prostrata Cav.

Perityle microglossa var. saxosa (Brandegee)

So,D,N

A.M. Powell

Perymenium pringlei B.L. Rob. & Greenm. var. pringlei

D,J

Perymenium reticulatum JJ. Fay

Mex

Pinaropappus junceus A. Gray (Nvo. Reg.)

Ch,Si, So

Pinaropappus roseus. (Less.) Less.

NAm

Pippenalia delphinifolia (Rydb.) McVaugh

Ch,D,Z

Porophyllum linaria (Cav.) DC.

Mex

Porophyllum macrocephalum DC.

NAm

Porophyllum pringlei B.L. Rob.

Mex

Psacalium globosum (B.L. Rob. & Fernald) H. Rob. & Brettell

Ch,D

Psacalium sinuatum (Cerv.) H. Rob. & Brettell

D,J,N,Mi

Pseudelephantopusspicatus (Juss. ex Aubl.) Rohr.

SAm

466

Journal of the Botanical Research Institute of Texas 9(2)

Tabla 3. (continued)

ESPECIES

Coni-

feras

Sinan-

tropica

Distribution

Americana

Endemicas

de:

Distribu¬

tion mas

amplia

Pseudognaphalium oxyphyllum (DC.) Kirp.

Mex

Psedognaphalium semiamplexicaule (DC.) Anderb.

CAm

Ratibida mexicana (S. Watson) W.M. Sharp

Mex

Roldana chapalensis (S. Watson) H. Rob. & Brettell.

Mex

Roldana hartwegii (Benth.) H. Rob. & Brettell

Mex

Schkuhria pinnata var. wislizenii (A. Gray) B.L. Turner

CAm

Sclerocarpus sessilifolius Greenm.

D,J,N,Si

Senecio stoechadiformis DC.

Mex

Sigesbeckia jorullensis Kunth

SAm

Simsia amplexicaulis (Cav.) Pers.

CAm

Sinclairiapalmeri (A. Gray) B.L. Turner

D,N,J,Z

Stenocarpha filiformis (Hemsl.) S.F. Blake

D,Si

Stevia aschenbomiana Sch. Bip.

Mex

Stevia jorullensis Kunth

CAm

Stevia micradenia B.L. Rob.

Mex

Stevia myricoides McVaugh

Mex

Stevia origanoides Kunth

Mex

Stevia ovata Willd.

Stevia purpusii B.L. Rob.

Mex

Stevia serrata Cav.

SAm

Stevia trifida Lag.

Mex

Symphyotrichum expansum (Poepp. ex Spreng.)

G.L. Nesom

Symphyotrichum potosinum (A. Gray) G.L. Nesom

Mex

(Acuatica)

Tagetes fHi folia Lag.

SAm

Tagetes foetidissima DC.

CAm

Tagetes micrantha Cav.

Nam

Tagetes palmeri A. Gray

Mex

Tagetes pringlei S. Watson

Mex

Tagetes subulata Cerv.

SAm

Tithonia fruticosa S. Can by & Rose

Mex

Tithonia tubiformis (Jacq.) Cass.

SAm

Tridax mexicana A.M. Powell

CAm

Trigonospermum annum McVaugh & Lask.

CAm

Verbesina longifolia (A. Gray) A. Gray

NAm

Verbesina oncophora B.L. Rob & Seaton

Mex

Verbesinaparviflora (Kunth) S.F. Blake var. parviflora

Mex

Vernonanthura liatroides (DC.) H. Rob.

D,0

Viguiera dentata (Cav.) Spreng.

NAm

Xanthisma stenolobum (Greene) D.R. Morgan & R.L. Hartm.

D,Ch

Xanthocephalum benthamianum Hemsl.

Mex

Zinnia americana (Mill.) Olorode & A.M. Torres

CAm

Zinnia angustifolia Kunth. var. angustifolia

CAm

Zinnia angustifolia var. greggii (B.L. Rob. & Greenm.)

D,N

McVaugh

Zinnia peruviana (L.) L.

Zinnia tenuis (S. Watson) Strother

D,Ch

Regiones : Am= America, CAm= Centroamerica, NAm= Norteamerica, SAm= Sudamerica. Pais: Mex= Mexico. Estados: Chis= Chiapas,

Ch= Chihuahua, Co= Coahuila, D= Durango, G= Guerrero, J= Jalisco, Mi= Michoacan, N= Nayarit, 0= Oaxaca, SLP= San Luis Potosi, Si=

Sinaloa, So= Sonora. Ac= Acuatica.

Ramirez Noya y Herrera Arrieta, Asteraceae en El Duraznojamazula, Durango, Mexico

467

caducifolio), la riqueza de especies en el ecosistema templado es marcadamente mayor que en el ecosistema

tropical. Solamente se registran 2 especies de habito acuatico: Jaegueria hirta y Symphyotricum potosinum.

Un caso interesante es el comportamiento de Tithonia tubiformis, una maleza ampliamente distribuida en

otras regiones de Mexico. En la zona de estudio solo se registraron unos cuantos ejemplares en el area de estu-

dio (localidad anotada en Tabla 1), lo que sugiere un buen nivel de conservacion de la vegetacion (<>?).

Las observaciones de la flora de Compuestas resultaron ser muy interesantes a lo largo de los recorridos

realizados (Tabla 3), los que aun siendo de pequenas dimensiones, registraron numeros altos de especies de

esta familia. De los 149 generos y 602 especies estimados para Durango (Balleza y Villasenor 2002), 50.3% de

los primeros y 24.7% de las segundas se encontraron en esta reducida region. Setenta y ocho especies de las 149

encontradas (52.3%) restringen su distribucion a Mexico, un aspecto sumamente interesante, ya que una de

cada dos compuestas de nuestra region de estudio, son endemismos a nivel nacional. Dicho porcentaje resulta

ser comparable con el 55.85% de endemismos estimado para el estado de Durango (Villasenor et al. 2004). Los

resultados senalan tambien que 41 especies (27.7%) son arvenses y/o ruderales (especies sinantropicas), las

cuales caracteriza a las Compuestas (Villasenor 2012) como la segunda familia (despues de las Poaceas) con

una proporcion alta de especies arvenses y/o ruderales.

Dos de las 148 especies encontradas presentan distribucion restringida al estado de Durango, dando un

valor de 1.35% de endemismo estatal. Dicha cifra resulta ser del doble que la de Zacatecas, que presenta un

0.8% de endemismo estatal, mientras que representa una decima parte del 11.3% de endemismo estimado para

la flora del estado de Durango (Villasenor 1993). Al analizar el endemismo a nivel regional (Durango, junto con

uno a tres estados colindantes), se observa que 20 especies (Tabla 3) son endemicas en la region compartida de

Durango con un estado colindante, otras ocho especies se conocen de Durango y dos estados colindantes y

cuatro especies de Durango con tres estados colindantes. Lo anterior resulta en un total de 32 especies endemi¬

cas a nivel regional, elevando as! el 1.35% de las especies endemicas a nivel estatal a un 22.97% de endemismo

en la region del noroeste-norte de Mexico. Los resultados ponen de manibesto la importancia del alto ende¬

mismo en la zona de estudio.

Es relevante la riqueza florlstica de Asteraceas encontrada en una region tan pequena como la del presente

estudio, resultado que se esperaba considerando sus caracterlsticas hsiograhcas, lo abrupto del terreno y la

localizacion de estas quebradas en la Sierra Madre Occidental, region considerada por la CONABIO como area

de exclusion. Asl, podemos anotar que nuestros resultados no son comparables con los de estudios florlsticos

en regiones colindantes, teniendo en cuenta la falta de caminos y lo intransitable del area explorada en un in-

tervalo altitudinal de 670-2550 m, que se dice facil, pero que requiere de largas caminatas con el apoyo de ani-

males de carga.

Otros resultados interesantes son las diferencias en el numerode especies con las reportadas en regiones

colindantes. Por ejemplo, Nesom (1989a) en sus estudios del genero Erigeron seccion Polyactis , reconoce 19

especies, citando 6 de ellas para el NW de Durango, de las que cuatro especies se registran en este estudio:

Erigeron coroniglandifer, E. polycephalus, E. seemannii y L. astrahntioides. Existen registros de varias especies de

Asteraceae que han sido descritas de regiones adyacentes a nuestra area de estudio, como son las nuevas espe¬

cies del cerro Mohinora, ubicado en el colindante municipio de Guadalupe y Calvo, Chihuahua: Erigeron cau-

linifolius, E. circulis (Nesom 1989a), L. mohinorensis (Nesom 1989b), L. mcdonaldii (Nesom 1990), L. oreophilus

forma latilobus (Greenman 1905) y Senecio mohinorensis (Greenman 1907). Sin embargo, estas 6 especies en¬

contradas en un area radial aproximada de 60 km en llnea recta de la region de estudio, a pesar de su cercanla

y similitud de condiciones ambientales, no fueron registradas y no se reportan en este trabajo. Una posible

explicacion es el microhabitat especlhco en el que fueron encontradas, causado principalmente por las diferen¬

cias de altitud en un terreno escarpado, provenientes de altitudes cercanas a los 3000 m, versus las recolectadas

para este estudio en altitudes ostensiblemente mas bajas (690-2550 m). En un estudio reciente, McDonald et

al. (2011) reportan para la parte alta del cerro Mohinora un total de 14 especies de Compuestas, de las cuales

solo 2 son compartidas con la region estudiada: Packera scalaris y Verbesina longifolia. Los resultados sugieren

discrepancias en su riqueza florlstica, aun en distancias relativamente cortas (alta diversidad beta). Por otra

468

Journal of the Botanical Research Institute of Texas 9(2)

parte, 3 especies que fueron descritas de los vecinos muncipios de Canelas y Santiago Papasquiaro, Durango, a

una altitud de 2200 m: Galinsoga spellenbergii (Turner 1986), Laennecia spellenbergii (Nesom 1992) y Verbesina

hygrophila (Panero y Villasenor 1993), y una adicional, Verbesina torresii (Turner 1985b) de la localidad El Ca-

torce, municipio Tepehuanes, Durango, a solo 35 km de El Durazno y 1400 m de altitud, no se encontraron

durante los recorridos de campo, aun cuando las altitudes de donde fueron descritas tambien se encuentran

dentro del intervalo altitudinal muestreado, a una distancia lineal menor a 50 km en llnea recta. Nuevamente

estos resultados sugieren que en la Sierra Madre Occidental lo abrupto del terreno confrere una alta riqueza

florlstica en diferencias de coordenadas geografrcas cortas, lo que provee microhabitats diversos que permiten

el desarrollo de endemismos y una alta diversidad beta.

El estudio de Vega Avina et al. (2000), reporta las especies endemicas para la region oeste y noroeste de

Mexico. De ellas, 26 especies en 23 generos son Asteraceas, presentes en el municipio de Culiacan, Sinaloa,

colindante con el municipio de Tamazula y del cual solamente una pequena porcion al noreste son terrenos

colindantes con nuestra area de estudio. De las 26 especies reportadas solamente se encontraron en la zona

estudiada Adenophyllum anomalum, Lasianthaea ceanothifolia var. gracilis, Lasianthaea seemannii y Perityle mi-

croglossa var. saxosa. La explicacion al bajo numero de especies compartidas seguramente es que Vega Avina y

colaboradores exploraron y recolectaron principalmente en la planicie costera del municipio de Culiacan, en

altitudes de 0 a 300 m la mayorla (rara vez por areriba de los 500 m), al parecer excluyendo localidades en las

serranlas del municipio (760 a 860 m), mismas que representan la subida a la Sierra Madre Occidental, en el

area colindante con nuestra area de estudio por encima de 1500 m de altitud.

Los resultados del presente estudiorevelan una alta riqueza de endemismos de la familia Asteraceae en

esta franja de la Sierra Madre Occidental, cadena montanosa donde las pendientes pronunciadas del terreno

unido a los conflictos sociales que le caracterizan, desmotivan la exploracion biologica necesaria para estudios

florlsticos. Se inhere que existen otras porciones de la region de las quebradas de la Sierra Madre Occidental

que se encuentran todavla subexploradas y que seguramente integran un territorio de alta diversidad biologica

y riqueza de endemismos, lo que podra corroborarse con muestreos sistematicos en estudios futuros.

En la Tabla 3 se indica la distribucion geograhca de las especies registradas. Destacan las especies con

ahnidad Neartica (26), distribuidas de Mexico a Norteamerica, mientras que 23 de ellas son de ahnidad Neo¬

tropical, restringidas principalmente a la region mesoamericanas (Davidse et al. 1994). Solamente 17 especies

se distribuyen mas al sur, hasta Sudamerica. Finalmente resulta interesante que 96% de los generos y 95.3% de

las especies se distribuyen solo en America y el porcentaje restante de especies son ahora de distribucion mas

amplia, ya que aparentemente han logrado establecerse como malezas en localidades del Viejo Mundo (Tabla

3).

AGRADECIMIENTOS

Se agradece al CIIDIR, COFAA y SIP del IPN, por su apoyo a los investigadores responsables de este reporte. Al

Instituto de Cultura del Estado de Durango, por el apoyo recibido durante 2003 a traves del Fondo Estatal para

la cultura y las artes. Al PACMYC y la Comision de Apoyo a la Creacion Popular del Estado de Durango

(Proyecto No. 46, ano 2008). Tambien al proyecto Plan de Manejo de la Canada de la Aguateca de El Tecuan,

Tamazula, Durango. As! mismo se agradece la cooperacion y apoyo de los residentes de las comunidades en las

que nos fue permitido instalar nuestro campamento, en Los Aguajes, El Durazno, Tamazula, Durango, a la fa¬

milia Vergara Vergara, en especial al Sr. Narciso Vergara Villarreal, Guadalupe Vergara V., Pedro Vergara V. y Sr.

Anastasio Vergara. Tambien a los senores raramuris nativos del ejido El Tecuan, El Durazno, Tamazula, Du¬

rango: Sra Paulina Gomez Lazcano 1 ) Andrea Gomez Lazcano, Ramona Gomez Lazcano, Monica Vargas Go¬

mez, Ursula Borjas Castillo, Venancio Gomez Lazcano 1 ) Liborio Borjas Castillo. Varios de ellos nos apoyaron

como gulas de campo y nos dieron hospedaje durante los anos de recolecta. Por ultimo, nuestro sincero agra-

decimiento a los revisores del manuscrito, especialmente al Dr. Villasenor, ya que sus acertadas observaciones

nos permitieron enriquecer el reporte final.

Ramirez Noya y Herrera Arrieta, Asteraceae en El Duraznojamazula, Durango, Mexico

469

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Greenman, J.M. 1907. New or noteworthy spermatophytes from Mexico, Central America, and the West Indies. Publ. Field

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Hinojosa-Espinosa, O. & R. Cruz-DurAn. 2010. Compositae in Atenango del Rio, Guerrero, Mexico. Bol. Soc. Bot. Mexico

86:71-75.

King, R.M. & H. Robinson. 1987. The genera of Eupatorieae (Asteraceae). Monogr. Syst. Bot. Missouri Bot. Gard. 22:1 -581.

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center.org/wab/2012/docs/Session%208/1 %20Villasenor.pdf

REDESCUBRIMIENTO DE DESMODIUM ANGUSTIFOLIUM (FABACEAE)

EN EL SALVADOR

Pablo Galan

Herbario LAGU, Jardm Botanico La Laguna

Urbanizacion Industrial Plan de La Laguna, Antiguo Cuscatlan

Apdo. Postal 1197 CG, La Libertad, EL SALVADOR

pgogalan@gmail.com

RESUMEN

Se recolecta nuevamente Desmodium angustifolium para El Salvador, despues de 43 anos de no tener registros de esta especie en el pais. Es un

arbusto debil que se distingue facilmente de otros Desmodium por sus hojas muy angostas y largas. Se encontro en 3 departamentos: 2 al oc-

cidente y uno en la zona oriental del pais.

ABSTRACT

Desmodium angustifolium is collected again in El Salvador, after 43 years of not having records of the species in the country. It is a weak shrub

that is easily distinguished from other Desmodium species by very narrow and long leaves. It was found in three states: two in the west and

the other in the east of the country.

Keywords: Desmodium, Hedysarum, Fabaceae, Morazan, Sonsonate, Santa Ana, El Salvador

INTRODUCCION

En El Salvador, la familia Leguminosae (Fabaceae); conformada por sus tres subfamilias, es una de las mas

numerosas. Para el genero Desmodium Desv. subfamilia Papilionoideae, se reportan 21 especies; ubicadas en

los herbarios LAGU y MHES.

El primer registro de D. angustifolium; es la colectada por Otto Rohweder, el 13 de Agosto de 1950 en la

Hacienda San Jose de la Finca Montecristo en Metapan, Santa Ana; zona noroccidental de El Salvador a una

altitud entre 850 alOOO m.s.n.m. y depositada en el herbario del Missouri Botanical Garden (MO) (Tropicos.

org).

DESCRIPCION DEL GENERO

Desmodium Desv.

Plantas herbaceas a sufrutescentes, postradas, erectas o trepadoras. Follolos 1-3, mucronados, estipelas pre-

sentes; estlpulas generalmente libres o connadas, la mayor la oblicuamente ovadas, largamente atenuadas, es-

triadas, puberulentas en la superficie dorsal, ciliadas. Inflorescencias racemosas o paniculadas, axilares y ter-

minales, bracteas estriadas, ciliadas; caliz bilobado, lobo superior blfido, lobo inferior 3-dentado; petalos pur¬

puras, azules, rojos o blancos (raramente verdosos o amarillentos), estandarte orbicular a obovado, unguicu-

lado, alas falcadas, auriculadas, unguiculadas, carina fusionada; estambres 10, diadelfos, el vexilar fusionado

con el tubo ca 1/3 de su longitud, tubo estaminal persistente en la base del fruto maduro. Lomentos de forma

variada, 1-9 articulados, pianos, enrollados, incurvados, recurvados o incurvado-recurvados, plegados, prin-

cipalmente indehiscentes, sesiles o estipitados (Stevens et al. 2001).

DESCRIPCION DE LA ESPECIE

Desmodium angustifolium Kunth ( Hedysarum angustifolium Kunth) es una hierba escandente o arbusto debil

de hasta 2 m; tallo y corteza lenosa, marron a rojiza; hojas con un follolo lineal, angostos; flores papilionoideas,

rosadas; los frutos son lomentos de hasta 3 mm de largo y 1-3 mm de ancho (Figs. 1, 2).

Distribucion y habitat. —Mexico al norte de Sudamerica, de 700 a 1400 m.s.n.m. (Stevens et al. 2001).

J. Bot. Res. Inst. Texas 9(2): 471 - 474.2015

472

Journal of the Botanical Research Institute of Texas 9(2)

Fig. 1. A. Planta con flores, colectada en A.N.P. San Marcelino. B. Planta con flores y

frutos colectada en A.N.P. La Ermita.

Para El Salvador, los sitios de colecta fueron: Santa ana:

Bosque de robles; en Sonsonate: zona abierta, revestida por

pastizales y con arboles distantes de Pinus; segun MARN 2011,

corresponde al ecosistema denominado: Flujo de lava con escasa

vegetacion. En el segundo sitio, Morazan, predominan los pas¬

tizales y areas semiabiertas con presencia de Pinus y Quercus;

predominando el estrato arboreo. Segun MARN 2011, corre¬

sponde al ecosistema denominado: Bosque tropical semideciduo

mixto submontano bien drenado.

METODOLOGIA

Siguiendo el protocolo de recolecta cientlfica, realizado por el

personal tecnico del herbario deljardln Botanico La Laguna, se

realizaron giras de campo periodicas; con el fin de incrementar

la coleccion de referencia florlstica a nivel nacional.

En agosto de 2012, se visito al Area natural protegida

Complejo San Marcelino; ubicada entre los departamentos de

Sonsonate (Sectores: El Chino-Teshcal y Bosque Las Lajas) y

Santa Ana (Sectores La Presa) en la zona occidental. La primera

Galan, Se recolecta nuevamente Desmodium angustifolium para El Salvador

473

colecta de D. angustifolium, fue en sector El Chino. En octubre de 2013, se visito el Area natural protegida La

Ermita; al nororiente del departamento de Morazan, donde se registro la segunda colecta.

Se revisaron ejemplares de Desmodium en los herbarios LAGU y MHES, encontrando material colectado

unicamente en LAGU. Complementando la consulta en las bases de datos de los sitios Web: Tropicos.org, JS-

TOR Global Plants y Field Museum of Chicago.

Material examinado: EL SALVADOR: Santa Ana: Municipio Candelaria de la Frontera, Cerro El Yupe, vegetacion de robles, 28 Oct 1993,

J.L. Linares & C.A. Martinez 914 (EAP); J.L. Linares & C.A. Martinez 924 (EAP); 06 Oct 1996, J.L. Linares 3652 (EAP, MEXU); Municipio

Metapan, Parque Nacional Montecristo, Mojon del Nance a una altitud de 1186 m.s.n.m. entre las coordenadas 14°21 , 36.65"N 89°

23 , 45.98"W, 03 Sep 2015, P. Galan & S. Garcia 3417 (LAGU). Sonsonate: Municipio Izalco, Canton Las Lajas, A.N.P. Complejo San Mar-

celino, Sector El Chino, Cima de Cerro El Chino, a una altitud de 1206 m.s.n.m. entre las coordenadas 13 0 48'35"N, 89 0 35'22"W, 16 Ago 2012

(fr), P. Galdn, A. Ibdnez, & O. Santamaria 1840 (INB, LAGU, MEXU). Morazan: Municipio Joateca, Canton El Tizate, A.N.P. La Ermita, alre-

dedores de terrenos de Asociacion Administradora de Agua Potable y Saneamiento Ambiental (ASAPSMA), a una altitud de 1061 m.s.n.m.

entre las coordenadas 13°55 , 57"N, 88°4 , 27"W, 09 Oct 2013 (fl, fr), P. Galdn & D. Rodriguez 2542 (B, INB, LAGU, MEXU, MHES, MO).

DISCUSION

Standley y Steyermark (1946) reportan D. angustifolium, desde Mexico hasta Colombia; entre los 100 y 200

m.s.n.m. Considerada una especie poco conspicua y ampliamente distribuida en Guatemala; en El Salvador es

conocida como “lengua de pajaro”.

Schubert (1965) describe el genero Desmodium en Flora of Panama; entre ellos D. angustifolium, con distri-

bucion desde Mexico hasta el norte de Sur America.

Sosa y Gomez-Pompa (1994), en su Lista Florlstica de “Flora de Veracruz”; reportan D. angustifolium y 48

especies mas de Desmodium, para esa region del atlantico mexicano.

Estrada Castillon et al. (2004) realizaron un estudio de las leguminosas en el centro del Estado de Nuevo

Leon, Mexico; siendo Desmodium, uno de los generos con mayor numero de especies; 11 en total, entre ellas D.

angustifolium.

Zamora (2010) reporta para Costa Rica Desmodium angustifolium var. angustifolium, en Bosques humedos

y muy humedos a elevaciones entre 750 y 1450 m.s.n.m.; haciendo mencion que se puede distinguir de otras

especies por sus hojas unifoliadas.

474

Journal of the Botanical Research Institute of Texas 9(2)

CONCLUSIONES

Desmodium angustifolium, probablemente podrla registrarse en otras localidades con habitats similares; para lo

cual se tienen que hacer investigaciones minuciosas. Por otra parte, existe un alto grado de fragmentacion y

deterioro en areas privadas y estatales que poseen habitat adecuado para su desarrollo. Tomando en cuenta la

distribucion restringida y el escaso material colectado, se puede sugerir como una especie con prioridad de

conservation en el pals.

AGRADECIMIENTOS

A Lilian Ferruhno de herbario EAP, Escuela Agricola Panamericana “Zamorano” de Honduras por facilitar

material digital de D. angustifolium, a D. Rodriguez del herbario LAGU por su valiosa revision y correcciones, a

G. CerenyJ. Menjlvar del herbario MHES que amablemente permitieron el ingreso a sus colecciones. Agradec-

emos a Jose Linares y un re visor anonimo por los comentarios.

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Schubert, B.G. 1965. Desmodium. En: J.D. Dwyer & collaborators. Flora of Panama, Part V, Fascicle 4. Family 83. Legumi-

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Sosa, V. & A. Gomez-Pompa. 1994. Leguminosae. Lista Floristica. Flora de Veracruz. Instituto de Ecologia, A.C. Xalapa, Uni¬

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Standley, P.C. & J.A. Steyermark. 1946. Leguminosae. Flora of Guatemala. Fieldiana, Bot. 24(5):1-368.

Stevens, W.D., C. Ulloa, A. Pool, & O.M. Montiel. 2001. Desmodium angustifolium. Flora de Nicaragua. Monogr. Syst. Bot.

Missouri Bot. Gard. 85:993.

Zamora Villalobos, N. 2010. Fabaceae. In: B.E. Hammel, M.H. Grayum, C. Herrera, & N. Zamora Villalobos, eds. Manual de

plantas de Costa Rica. Monogr. Syst. Bot. Missouri Bot. Garden, St. Louis, Missouri, U.S.A. 119(5):395-775.

ARACHIS GLABRATA (FABACEAE) NEW TO THE FLORA OF LOUISIANA, U.S.A.

Charles M. Allen

Colorado State Univ., Fort Polk Station

1645 23rd St

Fort Polk, Louisiana 71459, U.S.A.

ABSTRACT

Rhizome peanut (Arachis glabrata Benth.) (Fabaceae) is reported new to Touisiana and increases the number of states for this introduced

species to four (Alabama, Florida, Georgia, and Touisiana).

RESUMEN

El cacahuet de rizoma (Arachis glabrata Benth.) (Fabaceae) se cita como nuevo para Touisiana e incrementa a cuatro el numero de estados de

esta especie introducida (Alabama, Florida, Georgia, y Touisiana).

Rhizome peanut (Arachis glabrata Benth.) (Fabaceae) is a native of southern Brazil, Paraguay, and extreme

northeastern Argentina (Krapovikas & Gregory 2007) that was apparently first introduced into the United

States in Florida (Isley 1990, 1998). This species has also been reported from Georgia (Carter et al. 2009) and

Alabama (Keener 2012). A recent collection from New Orleans is apparently the first report for Louisiana. The

current distribution map in the USDA Plants Database (http://plants.usda.gov/plants) and BONAP (Kartesz

2015) do not include Louisiana.

Voucher specimens: LOUISIANA. Orleans Parish: yellow flowers, locally abundant in soil area between sidewalk and building at 7446

Garfield St., New Orleans, Louisiana, 20Jun 2015, Charles Allen 22905 (LSU, BRIT, TEX-LL).

REFERENCES

Carter, R., W.W. Baker, &W.M. Morris. 2009. Contributions to the flora of Georgia, U.S.A. Vulpia 8:1-54.

Isley, D. 1990. Vascular flora of the southeastern United States, Vol. 3 Part 2, Leguminosae (Fabaceae). The Univ. North

Carolina Press, Chapel Hill, U.S.A.

Isely, D. 1998. Native and naturalized Leguminosae (Fabaceae) of the United States (exclusive of Alaska and Hawaii).

Monte L. Bean Life Science Museum, Brigham Young Univ., Provo, Utah, U.S.A.

Kartesz, J.T. 2015. The Biota of North America Program (BONAP).Taxonomic Data Center, (http://www.bonap.net/tdc).

Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic synthesis of North America, Version 1.0. Biota of

North America Program (BONAP). (in press)]

Keener, B.R. 2012. Three non-native vascular plant species new to Alabama. Phytoneuron 2012-73:1-3.

Krapovikas, A. & W.C. Gregory. 2007. Taxonomy of the genus Arachis (Leguminosae). Bonplandia 16 (Supl.):1 —205. [Trans¬

lated by D.E. Williams and C.E. Simpson].

USDA, NRCS. 2015. The PLANTS database (http://plants.usda.gov/plants). National Plant Data Center, Baton Rouge,

Louisiana 70874-4490, U.S.A.

J.Bot. Res. Inst. Texas 9(2): 475.2015

476

Journal of the Botanical Research Institute of Texas 9(2)

ANNOUNCEMENTS

2015 DELZIE DEMAREE TRAVEL AWARD RECIPIENT

The 27th Annual Delzie Demaree Travel Award was presented at the 62nd Annual Systematics Symposium

(8-10 Oct 2015) at the Missouri Botanical Garden, St. Louis. One student was presented the Travel Award:

Megan Ruffley, University of Idaho.

The 2015 Travel Award was underwritten by 1) Contributors to the Delzie Demaree Travel Award

Endowment, and 2) Members of the Delzie Demaree Travel Award Committee.

Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the

travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research

Institute of Texas, to Barney Lipscomb, 1700 University Drive, Fort Worth, TX 76107-3400, U.S.A. 1-817-332-

7432; Email: barney@brit.org. Thank you.

THE 2016 APPLICATIONS FOR THE DELZIE DEMAREE TRAVEL AWARD

Applications for the 2016 Delzie Demaree Travel Award should include a letter from the applicant telling how

symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major

professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology

Department, The College of William and Mary, Williamsburg, Virginia 23185-8795, U.S.A. 1-757-221-2799;

Email: dmeware@verizon.net. Applications may also be sent to: Barney Lipscomb, 1700 University Drive, Fort

Worth, Texas 76107-3400, U.S.A. 1-817-332-7432; Email: barney@brit.org. The period for receiving applica¬

tions will end three weeks prior to the date of the symposium if a sufficient number of applications are in hand

at that time. Anyone wishing to apply after that date should inquire whether applications are still being ac¬

cepted before applying. The Systematics Symposium dates for 2016 are 14-15 October 2016 (dates tentative and

subject to change).

The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35

out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discov¬

erer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He also

taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas A&M,

and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of the things he

enjoyed most as a botanist was assisting students with their held botany research.

J.Bot. Res. Inst. Texas 9(2): 476.2015

NEW CO-OCCURRENCE OF SCHOENOPLECTIELLA HALLII AND

S. SAXIMONTANA (CYPERACEAE) IN OHIO (U.S.A.):

CONSERVATION IMPLICATIONS FOR BOTH SPECIES

Paul M. McKenzie

US. Fish and Wildlife Service 1

101 Park De Vi lie Dr.; Suite A

Columbia, Missouri 65203, US.A.

paul_mckenzie@fws.gov

Marian Smith

Daniel W. Boone

3228 Hanna Ave.

Cincinnati, Ohio 45211, US.A.

Richard L. Gardner

Department of Biology Ohio Department of Natural Resources

Southern Illinois University Division of Natural Areas and Preserves

Edwardsville, Illinois 62026, US.A. 2045 Morse Road C-3

Columbus, Ohio 43229, US.A.

ABSTRACT

An examination of achenes from specimens of SchoenoplectieUa saximontana collected at a Pickaway County, Ohio, site in 2011 led to the

discovery of SchoenoplectieUa hallii at the same locality. How long S. hallii has been at the only known site for S. saximontana in Ohio is un¬

known, but is speculated to be of recent occurrence due to the lack of evidence of hybridization between the two species at this location. It is

surmised that the appearance of S. hallii at the Ohio site was due to achene dispersal by migrating waterfowl from established sites in Indi¬

ana, Illinois, Kentucky, or Michigan. The presence of S. hallii and S. saximontana at the Pickaway County site may eventually lead to hybrid¬

ization between the two species, as documented elsewhere within their range, threatening the conservation status of both species in Ohio.

Further monitoring of this site is warranted and additional surveys for both species in Ohio are recommended in areas that may provide

suitable habitat. It is also recommended that surveys be conducted at sites in North America that have either S. saximontana or S. hallii. In

cases where one species greatly outnumbers another, it is recommended that numerous plants be examined from different portions of the

population to detect the rarer species.

RESUMEN

Un examen de los aquenios de especimenes de SchoenoplectieUa saximontana colectados en un lugar del Condado de Pickaway, Ohio, en 2011

dio lugar al descubrimiento de SchoenoplectieUa hallii en la mismo localidad. No se sabe cuanto tiempo ha estado S. hallii en el unico lugar

conocido de S. saximontana en Ohio, pero se especula que sea una ocurrencia reciente debido a la falta de pruebas de hibridacion entre las

dos especies en esta localidad. Se supone que la aparicion de S. hallii en este lugar de Ohio fue debida a la dispersion de aquenios por aves

acuaticas desde puntos de Indiana, Illinois, Kentucky, y Michigan. Ta presencia de S. hallii y S. saximontana en el Condado de Pickaway

puede dar lugar a hibridacion entre las dos especies, como se ha documentado en otras partes, amenazando el status de conservacion de

ambas especies en Ohio. Se necesita mas control en este lugar y se recomiendan estudios adicionales de ambas especies en Ohio en areas que

puedan tener habitats adecuados. Tambien se recomienda que se realicen estudios en otros lugares de Norte America que tengan S. saximon¬

tana o S. hallii. En los casos en que una especie sobrepase mucho en numero a otra, se recomienda que se examinen numerosas plantas de

diferentes partes de la poblacion para detectar la especie mas rara.

INTRODUCTION

SchoenoplectieUa hallii (A. Gray) Lye and S. saximontana (Fernald) Lye are sedge species that were once thought

to be allopatric, with S. hallii occurring primarily in the Midwest and eastern U.S. and S. saximontana largely

confined to the western portions of the country (Gleason & Cronquist 1991; Beatty et al. 2004). Schoenoplecti-

ella hallii has a global ranking of G2/G3 (imperiled/vulnerable): It is listed as “critically imperiled” in seven of

the 10 states in which it is known to be extant, “imperiled” in two, and “imperiled/vulnerable” in one. Her-

dhe findings and conclusions in this article are those of the author(s) and do not necessarily represent the views of the U.S. Fish and Wildlife Service.

Dedication: This paper is dedicated to the memory of Galen Smith whose love and appreciation of the Cyperaceae was a constant source of inspiration to us. He had a

deep conservation concern for members of this family that were threatened with extinction. He was one of true classical taxonomists who understood the importance of

morphology, life history ecology, and natural community/habitat dynamics in separating one species from another. He will be greatly missed.

J. Bot. Res. Inst. Texas 9(2): 477 - 484.2015

478

Journal of the Botanical Research Institute of Texas 9(2)

barium records indicate that it had been reported from GA in 1966, MA in 1931 and IA in 1890 (McKenzie et

al. 2007), but these populations were “possibly or likely extirpated” (NatureServe 2015; McKenzie et al. 2007).

Although S. hallii is reported on NatureServe (2015) as “not ranked/under review” in South Carolina, McKen¬

zie et al. (2007) state that previous reports of S. hallii in SC are based on misidentihcation of S. erecta.

Schoenoplectiella saximontana has a global ranking of G5 (secure) (NatureServe 2015), but it is listed as SI

(critically imperiled) in British Columbia as well as in 7 of the 12 states where it occurs and S3 (vulnerable) in

TX. It has not been ranked or is under review in CA, OK, and SD. It has been reported from two states in Mexico

(Smith 2002a). Throughout its range, S. saximontana is considered to be an uncommon species whose distribu¬

tion is scattered (Smith 2002a; NatureServe 2015).

Schoenoplectiella saximontana was first collected in Pickaway County, Ohio, by Bartley and Pontius in

1936 (OS #3241, #3242, #3243) (https://herbarium.osu.edu/online-data-access; accessed 17Jan 2015) (Schaffner

1937) and subsequently collected at the same site nine times between 1948 and 2008 by several botanists

(http://128.146.250.9/bol/herbarium/Results#; accessed 17 Jan 2015). Most recently, it was collected 7 Aug

2011 by Dan Boone [ Boone 8-7-1229-11 (MO)]. Prior to this report there are no records of S. hallii for Ohio at this

or any other locality in the state (Schaffner 1937).

Pickaway County Site History

The Pickaway County site is located south of Circleville, Ohio, and historically was within a few kilometers or

adjacent to the “Pickaway Plains” native tall grass prairie that was estimated to be 4.8 km long and 11.2 km

wide. This prairie, however, was converted to cropland, and no remnant of this natural community currently

exists. When the site was first botanized by Floyd Bartley in 1936, he named the collection site “George Hitler’s

Pond” after George Hitler who owned the property and whose family originally settled the area in 1799. In the

1940s, the Hitler family sold the property to the List family who subsequently sold the farm in 2013 to the Ap¬

palachia Ohio Alliance, a non-profit conservation organization. The site is now preserved as part of the 95-acre

Floyd Bartley Nature Preserve (Gibbons 2014).

Site Characteristics

The terrain of the general area is gentle, rolling, glacially formed hills with depressions ranging in size from

<0.4 ha to 1 ha. The glacial deposits are Wisconsin age. The soils of Hitler Pond are deep, poorly drained soils

on outwash plains formed over ice contact depositions, with stratified sand and gravel layers. Permeability is

moderately slow in the subsoil but rapid in the substratum (USDA 1980); the slope is 0 to 2 percent. As else¬

where throughout the range of S. hallii, the Pickaway County site is cultivated when dry, but during wet cycles

the ponded water prevents the use of farm equipment necessary for crop production. In years when water is

present at the site, ponding occurs from February to early June (USDA 1980).

Plant associates documented at the site include: Abutilon theophrasti Medik., Amaranthus albus L., Amma-

nia robusta Heer and Regel, Cyperus esculentus L., Echinodorus berteroi (Spreng.) Fassett, Echinochloa crusgalli

(L.) Beauv., Eclipta prostrata (L.) L., Eleocharis engelmannii Steud., E. obtusa (Willd.) Schult., Eragrostis hypnoi-

des (Lam.) BSP, Gratiola neglecta Torr., Eindernia dubia (L.) Pennell, Mollugo verticillata L., Morus alba L. (seed¬

lings), Packera glabella (Poir.) C. Jeffrey, Populus deltoides Bartram ex Marshall (seedlings), Rorripapalustris (L.)

Bresser, R. sessiliflora (Nutt.) Hitchc., Rotala ramosior (L.) Koehne, Sida spinosa L., and Symphyotrichum lanceo-

latum (Willd.) G.L. Nesom.

Circumstances surrounding the discovery of S. hallii at the Pickaway County site

On 30 Sep 2014, Boone delivered specimens of Schoenoplectiella saximontana to the senior author, who broke

the bundle into five separate sheets, shattered the spikelets from a few isolated culms to create achene packets,

and observed that some of the achenes were two-sided rather than 3-sided as would be characteristic of S.

saximontana (Smith & McKenzie 2011). Upon closer examination with a hand lens, McKenzie confirmed that

they were achenes of S. hallii. After removing the specimens from the plant press, he examined all culms to

determine the source of the 2-sided achenes and located two specimens of S. hallii entwined with S. saximon¬

tana culms. The diagnostic difference in achene structure of the two species is depicted in Figure 1. For SEMs

of achene cross sections see Smith and McKenzie (2011).

McKenzie et al., Schoenoplectiella hallii and S. saximontana in Ohio

479

Fig. 1. Comparison of 3-sided achenes of 5. saximontana (A) from Dan Boone8-7-1229and 2-sided achenes of 5. hallii (B) from Dan Boone 8-7-1230. In row

A, note the prominent midrib in 5. saximontana, which creates its distinctive trigonous cross section. See cross sections of achenes (Smith & McKenzie 2011).

The following voucher specimens represent the first documented record of S. hallii for Ohio.

MISSOURI BOTANICAL GARDEN

PLANTS OF OHIO

Schoenoplectiella hallii (A. Gray) Lye

U.S.A. Ohio. Pickaway Co.: Floyd Bartley Nature Preserve, ca. 0.58 mi SW of the inter, of Rts. 41 and 56, or ca. 0.49 mi SSE of the inter, of

Rts. 68 and 56, and ca. 1.2 mi SE of Circleville; 39°34 , 43.66"N, 82°55T1.84"W; NE A Section 32, Circleville Township; elev. 707'. Two plants

mixed in a large population of S. saximontana in wet depression of agricultural field. Discovery made during examination of achenes by Paul

McKenzie on 29 Sep 2014; in association with Ammannia robusta, Echinodorus berteroi, Eleocharis englemannii, E. obtusa, Gratiola neglecta,

Lindernia dubia, Rorippapalustris, R. sessiliflora, and Rotala ramosior. First record for Ohio, 7 Aug 2011, Dan Boone 8-7-1230-1E

MISSOURI BOTANICAL GARDEN

PLANTS OF OHIO

Schoenoplectiella saximontana (Fernald) Lye

U.S.A. Ohio. Pickaway Co.: Floyd Bartley Nature Preserve, ca. 0.58 mi SW of the inter, of Rts. 41 and 56, or ca. 0.49 mi SSE of the inter, of

Rts. 68 and 56, and ca. 1.2 mi SE of Circleville; 39°34 , 43.66"N, 82°55T1.84"W; NE A Section 32, Circleville Township; elev. 707'. Hundreds

of thousands of plants in wet depression of agricultural field; in association with Ammannia robusta, Echinodorus berteroi, Eleocharis engle¬

mannii, E. obtusa, Gratiola neglecta, Lindernia dubia, Rorippapalustris, R. sessiliflora, Rotala ramosior, and Scheonpolectiella hallii ; 7 Aug 2011,

Dan Boone 8-7-1229-11.

480

Journal of the Botanical Research Institute of Texas 9(2)

The sole voucher specimen of S. hallii from the Pickaway County, OH, site is housed at the Missouri Botanical

Garden (accession no. MO-2443050 6604784) (Tropicos database: http://www.tropicos.org/Speci-

men/100755931). One Pickaway County specimen of S. saximontana is at MO (accession no. MO-2443051

6604783) (Tropicos database: http://www.tropicos.org/Specimen/100755932). Duplicates of Dan Boone 8-7-

1229-11 are deposited at CSCN, MICH, OKL, and UWW.

2014 field search of Pickaway County site and examination of herbarium collections

In 2014, Rick Gardner surveyed the site to determine if S. hallii and S. saximontana were present. He found an

abundance of S. saximontana but, no evidence of S. hallii, as had been documented in 2011. The inability to re¬

discover S. hallii at the site may be due to the absence of plants in 2014 or the difficulty in finding the species

among the thousands of culms of S. saximontana. Boone and McKenzie examined all culms of duplicates from

Boone’s 2011 collection of S. saximontana and were unable to find any additional specimens or achenes of S.

hallii. In 2014 and 2015, Gardner examined specimens of S. saximontana collected between 1936 and 2008 at

the Pickaway County site housed at the Ohio State University (OS) herbarium and was unable to confirm any

additional specimens of S. hallii among the specimens of S. saximontana. These observations suggest that ei¬

ther S. hallii is of recent occurrence at the Pickaway County site or has been overlooked due to its rarity at this

locality.

Smith and McKenzie (2011) reported similar fluctuations in species composition for sites of S. hallii and S.

saximontana near Lawton, Oklahoma, on the Wichita Mountains Wildlife Refuge (WMWR) and adjacent ar¬

eas on the Fort Sill Military Reservation (FSMR). During their studies between 2001 and 2013 (see, in part,

Smith & McKenzie 2011, 2013), there were sites where they observed only one of the species; however, either

the second species had been documented previously at each site and/or the achenes were present in the seed

bank. The number of individuals of either species varied significantly from year to year. During some years one

species in low numbers would be difficult to find among the tens of thousands of plants of the second species.

In 2012, Robert Steinauer discovered S. saximontana at a site in Loup County, Nebraska, where previously only

S. hallii had been documented. A survey of the same site by Steinauer, Smith, and McKenzie on 4 Aug 2014

documented S. hallii but not S. saximontana. Since 2004 (Smith et al. 2004), the hybrid, Schoenoplectiella

xmagrathii (Smith & McKenzie 2013), has been reported at the OK sites in densities that fluctuate annually.

Origin of S. saximontana and S. hallii at the Pickaway County site

The origin of S. saximontana at the Pickaway County, OH, site has been an issue of debate. Because S. saximon¬

tana at this site is the only documented locality for this species east of the Mississippi River (Smith 2002a),

some (e.g., Stuckey & Roberts 1982) have postulated that it was adventive from hay transported from the west¬

ern United States where the species is of more regular occurrence. We agree with others, however, who have

suggested migrating waterfowl as the natural dispersal agent responsible for the appearance of S. saximontana

at the Pickaway County site. DeVlaming and Proctor (1968) discovered that ducks and killdeer ingest and

disperse viable achenes of a number of Scirpus species, and Gleason and Cronquist (1964) suggested that the

primary method for long-range dissemination of aquatic species is by attachment of achenes to feathers, bills,

or feet of migrating waterfowl. Vivian-Smith and Stiles (1994) examined 36 species of waterfowl and reported

that 78% of the birds had seeds of wetland plants adhering to their feathers or feet. Additionally, other authors

have demonstrated that achenes with hardened seed coats can remain viable after passing through the diges¬

tive tract of migrating waterfowl (DeVlaming & Proctor 1968; Powers et al.1978; Figuerola & Green 2002).

Others have reported the ability of migratory species of birds to transport seeds long distances externally and

internally (Proctor 1968 and see review in Nathan et al. 2008).

Scirpus species are particularly adapted for both internal and external transport because of their diminu¬

tive size and hard seed coats (Krefting & Roe 1949; DeVlaming & Proctor 1968). In Illinois, Oklahoma, and

Texas, dispersal of Schoenoplectiella is assumed to be facilitated by migrating waterfowl (McClain et al. 1997;

Magrath 2002; McKenzie et al. 2007; Smith & McKenzie 2013) as well as some ungulates, including bison

(Rosas et al. 2008). Some species of waterfowl have been known to fly 1200 km in less than 24 hrs (Preben

Clausen, pers. comm.in Figuerola & Green 2002), and the Pickaway County site is between approximately 371

McKenzie et al., Schoenoplectiella hallii and S. saximontana in Ohio

481

and 611 km from known sites of S. hallii in Illinois, Indiana, Kentucky, or Michigan. Consequently, achenes of

S. hallii could be transported in less than one day by migrating birds between other states with extant sites for

the species and the Ohio locality. As with S. saximontana, we believe that migrating waterfowl were likely re¬

sponsible for the presence of S. hallii at the Pickaway County site, and both species should be considered native

to the state.

Relationship of groundwater to the creation of habitat for Schoenoplectiella

Schoenoplectiella hallii, S. saximontana, and S. xmagrathii are obligate wetland taxa that have similar habitat

requirements: most often sandy, rocky, or gravelly soil, occasionally clay, around the margins of ponds, ditch¬

es, and swales with fluctuating water levels, and a scarcity of other vegetation as competitors (McClain et al.

1997; Smith 2002a; McKenzie et al. 2007; Smith & McKenzie 2013). In Mason County, Illinois, McKenzie et al.

(2010) linked population size and location to a rise in the Illinois River and a concomitant rise in groundwater

that created temporary ponds. Smith (2001, 2002, 2003) conducted a 3-year study in Scott County, Missouri,

and illustrated the dependence of seedling establishment in S. hallii on the presence of groundwater within 1

m of the soil surface and the maintenance of soil moisture at >10%, but less than 100%. Some populations in

Oklahoma contain individuals of all three taxa (Smith & McKenzie 2013), suggesting that they share the same

habitat requirements.

The lack of appearance of S. saximontana at the Pickaway County site between the early 1990s and 2008,

despite repeated trips to the locality to locate the species (Jim McCormac, pers. comm. 2015; Boone pers. obs.),

suggest that the availability of suitable habitat from year to year may be due to multiple factors. Walker et al.

(1965) conducted an analysis of groundwater in areas adjacent to the Scioto River Valley that include the Bart¬

ley Preserve and determined that the permeable sand and gravel deposits in the valley provided one of the most

important sources of large groundwater supplies in Ohio. The authors concluded (Walker et al. 1965:15) that

“natural recharge in the Scioto valley occurs from precipitation, from ground water flowing from the bedrock

at the edges of the valley, and from the Scioto River when it is in flood stage.”

Groundwater and the underlying aquifers are being depleted throughout the United States. In a study of

40 U.S. aquifers, Konikow (2013) determined that aquifer depletion totaled ca. 1000 km 3 from 2000 to 2008

and that the rate increased dramatically after 1950. The aquifer systems with the highest loss include areas

where Schoenoplectiella hallii and S. saximontana are most abundant: the High Plains aquifer (340.9 km 3 ), the

Mississippi embayment aquifer system (182.0 km3), and the Central Valley aquifer system of California (144.8

km 3 ) (Konikow 2013). Zekster et al. (2005) reported that groundwater depletion results in a decline in surface

water and stream flow, and Sophocleous (2002) indicated that such diminutions impact the ecological integrity

of wetlands, resulting in significant loss of habitat and biodiversity. As reported by Smith (2003) for Schoeno¬

plectiella hallii populations in Missouri, Burke et al (1999) suggested that many environmental factors rely on

the accessibility of underlying groundwater remaining within a few meters of the surface, rather than on the

volumetric amount of groundwater theoretically available from deep aquifers. Therefore, wetlands, transient

surface ponds, and stream flows may dry up even when the stored groundwater in a basin remains large.

The hydrological influences on available habitat at the Pickaway County site are not fully understood and

need further study. During years of above-average precipitation, suitable habitat may be present due to above

ground ponding of water or influenced by a rise in an underground aquifer levels. The impact of water with¬

drawals from underground aquifers for agricultural, industrial, or residential uses may be a factor in the avail¬

ability of habitat. During years of below-average precipitation, the added impact of water withdrawn for irriga¬

tion and other uses may prevent the formation of ponded water necessary for germination and growth (Baskin

et al. 2003). Such impacts may not be realized during years when there is an abundant water supply.

Walker et al. (1965) noted that groundwater within the Scioto River Valley flowed from the river to adja¬

cent sand and gravel deposits and suggested that well pumping could cause a reversal in flows. The amount of

groundwater recharge from the Scioto River is dependent upon “the condition of the river bottom, permeabil¬

ity of the unconsolidated material, thickness of the aquifers, and the distance and ground-water gradient be¬

tween the wells and the source of recharge” (Walker et al. 1965:15). Evidence that this aquifer may be impacted

482

Journal of the Botanical Research Institute of Texas 9(2)

due to water withdrawals for industrial use is supported by observations of land owners who live in the vicin¬

ity. The RCA-Thompson plant withdrew an estimated 1 million gallons of water a day when it was in operation

through 2004, and neighbors noted that water levels in the area were noticeably lower than in the years prior

to the plant’s construction and operation (Steve Fleegal, pers. comm., Mar 2015). Reduced water levels due to

water demands by the plant could explain the lack of observations of S. saximontana between the early 1990s

and 2007. Observations of Schoenoplectiella spp. at the Bartley Preserve in 2008, 2011, and 2014 were after the

RCA-Thompson plant closed in 2004 (http://www.epa.ohio.gov/cdo/rca.aspx). With increased needs of

groundwater in the area for commercial, industrial, residential, and recreational uses, further research is

needed to determine the impacts of these demands on habitat availability on the Bartley Preserve. The Appala¬

chia Ohio Alliance recently received an EPA research grant to study the hydrology of the site, and the project is

anticipated to start this year and may provide additional insights into the issue (Fleegal, pers. comm. Feb

2015).

Conservation implications of co-occurrences of Schoenoplectiella hallii and S. saximontana

Magrath (2002) conducted surveys for S. hallii and S. saximontana at multiple sites on the WMWR in Oklaho¬

ma in 2000 and found both species at 20 of the 134 sites that he searched. In 2001, Marian Smith discovered

suspected hybrids between S. hallii and S. saximontana at the same refuge, and this supposition led to an exten¬

sive survey by Smith and McKenzie on the WMWR and adjacent areas of FSMR between 2002 and 2013 (see,

in part, Smith & McKenzie 2011, 2013). These investigations resulted in the documentation of the first re¬

ported hybrid between the two species that included confirmation by genetic analysis (Smith et al. 2004; Smith

& McKenzie 2011, 2013; Esselman et al. 2012). The hybrid was formally described as Schoenoplectiella

xmagrathii Smith and McKenzie (Smith & McKenzie 2013). Smith and McKenzie (2011, 2013) outlined the

potential adverse impacts of hybridization on the conservation of both S. hallii and S. saximontana. As is the

case in Oklahoma, the co-occurrence of S. hallii and S. saximontana in the same population in Ohio may result

in the development of a hybrid swarm, which could threaten both species. Three additional states have sites

where there are mixed populations (NatureServe 2015): Kansas (Craig Freeman pers. comm. 2006), Texas

(Bob O’Kennon, pers. comm. 2007), and Nebraska (Robert Steinhauer, pers. comm. 2012). Although S. saxi¬

montana occurs in eight counties in Oklahoma, it is known to co-occur with S. hallii and the hybrid S.

xmagrathii in only Comanche County (Smith & McKenzie 2013; Oklahoma Vascular Plant Database 2015). It

is not yet known if the hybrid occurs in the Pickaway County site in Ohio, but co-occurrence of S. hallii, S. saxi¬

montana and the hybrid S. xmagrathii at sites in Oklahoma would suggest that it is possible.

Schoenoplectiella saximontana has been known from a number of populations in south Texas for many

years, but the presence of S. hallii in the LBJ Grasslands in North Texas was not reported until 2004 (O’Kennon

& McLemore 2004). Subsequently, both species were discovered in 2007 at Rhodes Lake, Decatur County

(O’Kennon, pers. comm. 2012). As it is less than 193 km from Rhodes Lake to the populations in Comanche

County, Oklahoma, it is plausible that migrating waterfowl could have dispersed seeds from these earlier-es¬

tablished populations to north Texas. Additional S. hallii and S. saximontana sites have been discovered on the

LBJ Grasslands recently, with some individuals having both 2- and 3-sided achenes (O’Kennon, pers. comm.

2012). In addition, S. saximontana populations containing apparent hybrids were found at Enchanted Rock

State Natural Area in south-central Texas in 2012 (O’Kennon, pers. comm.).

Since 1999, numerous new populations of S. hallii have been reported in Nebraska, and a large population

containing both species was discovered there in 2012 (Robert Steinhauer, pers. comm. 2015). Specimens of S.

hallii and S. saximontana taken in 2012 by Steinhauer were examined by McKenzie and Smith in 2014, and no

evidence of hybrids was discovered. A trip to the same site in 2014 by Steinhauer, Smith, and McKenzie yielded

specimens of S. hallii but not S. saximontana. Based on observations by Smith and McKenzie in Oklahoma,

however, the appearance of the hybrid S. xmagrathii may not be immediate. McKenzie and Smith examined

specimens collected by Magrath in 2000 and reported in Magrath (2002) but saw no evidence of hybrids. A

held investigation by Smith in 2001, however, led to the initial observation of possible hybrids (Smith & McK¬

enzie 2011). No hybrids were reported at Rhodes Lake, Texas, in 2007 where both species were recorded, but

McKenzie et al., Schoenoplectiella hallii and S. saximontana in Ohio

483

within five years evidence of hybridization was observed by O’Kennon (pers. comm. 2012). Given these obser¬

vations, it is predictable that S. xmagrathii may occur in Nebraska and Ohio. In a report of the status of hybrids

in Oklahoma, Smith and McKenzie (2013) postulated that S. xmagrathii can displace and out-compete the pa¬

rental species under certain environmental conditions. During their investigations between 2001 and 2012

(Smith & McKenzie 2011, 2013), they observed notable changes in the abundance of the parental species and

the hybrid, and noted that in some years S. xmagrathii greatly outnumbered either S. hallii or S. saximontana.

Recommendations for additional research

Populations of S. hallii and S. saximontana in Pickaway County, Ohio, and other states where both species co¬

occur should continue to be monitored to assess the appearance of hybrids. States where one species is known

to occur, but not the second, should be surveyed to search for the presence of the other. Because one species

may greatly outnumber the second and the rarer species may be limited in its distribution, it is recommended

that multiple individuals from a cross section of the population be examined. Finally, because the species can

be difficult to distinguish morphologically, spikelets on herbarium sheets of both species should be examined

for the presence of 2-sided and 3-sided achenes. The continued occurrence and spread of S. xmagrathii threaten

the conservation and persistence of S. hallii and S. saximontana, but it is not yet known how hybrid population

size varies over time, or if the hybrid can displace one or both of the parental species completely.

ACKNOWLEDGMENTS

We thank Bob O’Kennon for information on populations of S. hallii and S. saximontana in Texas and his assis¬

tance in the held. We are grateful to Bob Steinauer in Nebraska for observations of S. hallii and S. saximontana

in Nebraska and his help in coordinating held visits in 2014 and George Yatskievych and Nancy Parker for as¬

sistance with Fig 1. We also are grateful to Mike Penskar for his thoughtful review of our manuscript.

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A CHECKLIST OF VASCULAR PLANTS AT THE COASTWARD EXTENT OF

COASTAL PRAIRIE IN SOUTHEAST TEXAS, U.S.A.

David J. Rosen and Siavash Zamirpour

Department of Biology

Lee College

Baytown, Texas 77522-0818, U.S.A.

drosen@lee.edu

Andrew Sipocz

Natural Resources, State Parks

Texas Parks and Wildlife Department

14200 Garrett Road

Houston, Texas 77044, U.S.A.

ABSTRACT

The Follets Island Conservation Initiative is a 179 ha parcel on Follets Island in Brazoria County, Texas. Field work from May 2013 through

May 2015 produced a checklist of 130 species of vascular plants representing 41 families and 105 genera from a flora that is 98% native (spe¬

cies) and a protected refuge for fourteen species of conservation interest. The Follets Island site is representative of a strand prairie, a term

used to describe the Gulf-shoreline extent of coastal prairie in southeast Texas.

RESUMEN

Fa Follets Island Conservation Initiative es una parcela de 179 ha en la Isla Follets en el condado de Brazoria, Texas. El trabajo de campo

desde mayo de 2013 hasta mayo de 2015 produjo un catalogo de 130 especies de plantas vasculares representando 41 familias y 105 generos

de una flora que es 98% nativa (especies) y un refugio protegido para catorce especies de interes de conservacion. Fa Isla Follets es represen-

tativa de una pradera de playa, un termino usado para describir la extension la linea de costa del Golfo de la pradera costera en el sureste de

Texas.

INTRODUCTION

Conservation of coastal prairie in Texas (characterized in detail by Smeins et al. 1991) has been a priority for

several decades. An important first step in the conservation of any ecosystem and a necessity for sound man¬

agement decisions is the inventory of botanical resources (Prater et al. 2004). To this end, a few held surveys of

the vascular plants of some of the remaining relatively large (42-120 ha) coastal prairie remnants have been

published (Rosen 2007, 2010, 2014; Singhurst et al. 2014a). These held studies have also provided important

botanical conservation information and progress towards compiling a more complete checklist of the vascular

plants of the ecoregion. Discovery and study of additional prairie remnants and endemic habitat types

throughout the coastal prairie ecoregion continues to be a priority of held botanists and prairie enthusiasts

(e.g., Singhurst et al. 2014b; Rosen unpublished data). Among these sites identihed by the authors is a 179 ha

parcel on Follets Island in Brazoria County, Texas.

Smeins et al. (1991) describe the coastal prairie ecoregion as extending inland from the coast up to 160

km. The Follets Island site was of interest to us as its uplands supported plant species characteristic of remnant

prairies over 55 km from the coast and therefore may represent the Gulf-shoreline extent of coastal prairie and

a contact with other Gulf-shoreline plant communities such as dunes, salt-hats, and salt-marshes. The site also

seemed to shed some light on what has been regionally referred to as “strand prairie” (Hollingsworth 1998).

The origin of this local name is uncertain, but it has been used in reference to upland prairies on Texas’ barrier

islands and Coastal Sand Plain dominated by Schizachyrium littorale (Nash) E.P. Bicknell (absent on Follets

Island) and Paspalum monostachyum Vasey (Diamond et al. 1987; Hollingsworth 1998). In the ecological con¬

text, strand refers to the shoreline of a marine environment, thus the name is htting to describe any Gulf-

shoreline prairie habitat type within the coastal prairie ecoregion. The purpose of the study presented here is

to provide a checklist of vascular plants from the Follets Island site supplemented with conservation notes for

some of our discoveries.

J. Bot. Res. Inst. Texas 9(2): 485 - 492.2015

486

Journal of the Botanical Research Institute of Texas 9(2)

METHODS

Our study site is located in Brazoria County on Follets Island, a peninsula bounded by the Gulf of Mexico and

Christmas Bay and approximately 52 km SW of the City of Galveston (Fig. 1). This 179 ha site was recently

acquired by the Texas Parks and Wildlife Department and will be formally called The Follets Island Conserva¬

tion Initiative and managed as a coastal preserve. The approximate center of this elongated site lies at

29.0196°N, 95.2010°W, and its dimensions are roughly 2.9 km long by 0.3 km wide at its narrowest point and

1.1 km wide at its widest point (Fig. 2). Prairie soils of the site (about 55 ha) are mapped as Mustang fine sand

(Crenwelge et al. 1981). The climate of the site is as described for nearby Galveston Island State Park (Rosen et

al. 2014). The prairies are not inundated by daily tides and occur on the uplands bounded by beach dunes at the

Gulf and salt-flats and salt-marsh on the bay side (Fig. 2). Prairie vegetation also occurs on elevated circular to

oblong mounds surrounded by the saline habitats (Fig 2, Fig. 3).

During ten collecting trips from May 2013 through May 2015 we focused our efforts on prairies through¬

out the site but collected in adjacent habitats as well. A complete set of voucher specimens collected by the first

author (6043-6734 & s.n.) is housed at TEX. Collections by the second author (1-116) are housed at BRIT.

Plant identifications were primarily determined using Flora of North America (1993+), Correll and Johnston

(1970), and Gould (1975). Classification and author names follow Tropicos (2015). The native, introduced, and

conservation status of each species were determined by review of Flora of North America (1993+), Correll and

Johnston (1970), and correspondence with held botanists or specific references mentioned later.

RESULTS & DISCUSSION

Our fieldwork produced a checklist of 130 species of vascular plants representing 41 families and 105 genera

(Appendix 1; Table 1). Speciose families include Poaceae (28 spp. including two non-native), Asteraceae (16

spp.), and Fabaceae (14 spp.). The largest genus, Dichanthelium (Hitchc. & Chase) Gould, contains six native

species. The species Bora is 98% native, while non-native species all rank infrequent-rare in abundance. One

family, Tamaricaceae, is represented by only one non-native species. Fourteen species are of conservation in¬

terest for their endemism, range extreme, and in some cases the discovery of populations on this protected site

(Appendix 2). Eighty-six species are known from inland remnants with vegetation considered representative

of coastal prairie. Five species co-occur at six other inland prairie remnants with nearly complete inventory by

the first author. Thirteen species are shared with other Gulf-shoreline habitats (dunes, salt-marshes, and salt-

hats). Thirty-four species do not occur at six other inland prairie remnants with nearly complete inventory by

the first author (although this could be related to the area of those sites, or that the species simply remain to be

discovered). Insomuch as other Gulf-shoreline habitats are not especially species rich (e.g., Table 2, Rosen et al.

2014), our study suggests the presence of coastal prairie vegetation at the Gulf-shoreline increases local plant

richness (at least at this site). We encourage the continued use of the term strand prairie in conservation efforts

as it emphasizes a unique and interesting phase of the Gulf-shoreline extent of coastal prairie vegetation in

southeast Texas. Discovery and protection of additional strand prairie habitats in southeast Texas should ac¬

company that of inland sites.

APPENDIX 1: CHECKLIST

Families are arranged alphabetically, beginning with monocots and followed by eudicots. Genera, species, and infraspecific names are

arranged alphabetically under families. Annotations indicate native species common to ( § ) or not collected from (♦) any of six other inland

prairie remnants with nearly complete inventory by the first author, non-native species (*) and notes detailing species of conservation

interest O' 14 ). Following each name is an abbreviation representing a frequently used scale of the relative abundance of that species:

r = rare (very difficult to find and limited to one or very few locations or uncommon habitats); i = infrequent (difficult to find with few

individuals or colonies but found in several locations); o = occasional (widely scattered but not difficult to find); f = frequent (easily seen

or found in one or more common habitats but not dominant in any common habitat); and a = abundant (dominant or co-dominant in

one or more common habitats).

Rosen et al., Vascular plants of Follets Island, Brazoria County, Texas

487

Fig. 1. General location of The Follets Island Conservation Initiative, Brazoria County, Texas.

MONOCOTS

AMARYLLIDACEAE

*Zephyranthes chlorosolen (Herb.) D. Dietr., i, 6660,94

ASPARAGACEAE

41 Yucca tenuistylolrel., i, 6512,54

COMMELINACEAE

Commelina erecta L. var. angustifolia (Michx.) Fernald, i, 6576, 71

CYPERACEAE

*Cyperus esculentus L., r, 6732

*Cyperus polystochyos Rottb., o, 6051

Cyperus retrorsus Chapm., o, 6497,40

Eleochoris montevidensis Kunth, o, 6045

Fimbristylis caroliniana (Lam.) Fernald, f, 6615,80

*lsolepis cernuo (Vahl) Roem. & Schult., i, 6289

Rhynchosporo colorata (L.) H. Pfeiff., o, 6046, 55

*Schoenoplectus pungens (Vahl) Palla var. longispicatus (Britton)

S.G.Sm., o, 6415,30

IRIDACEAE

* 2 Sisyrinchium biforme E.P. Bicknell, i, 6286,5

JUNCACEAE

Juncus dichotomus Elliott, i, 6421,34

Juncus marginatus Rostk., o, 6494,37

Juncus roemerionus Scheele, o, 6398, 20

43 Juncus fascinatus (M.C. Johnst.) W. Knapp, o, 6507,50

POACEAE

Andropogon glomeratus (Walter) Britton, Sterns & Poggenb. var.

pumilus (Vasey) Vasey ex L.H. Dewey, o, 6680,108

488

Journal of the Botanical Research Institute of Texas 9(2)

•5

] Kilometers

Digital Georectifed Image: 2014 NAIP (Oct 2014)

Brazoria County, Texas; Christmas Point Quadrangle

Fig. 2. True-color aerial photograph of The Follets Island Conservation Initiative (outlined in green), Brazoria County, Texas.

Aristida longespica Poir. var. geniculata (Raf.) Fernald, r, 6730

Aristido purpurascens Poir. var. purpurascens, i, 6723

*Bothriochloa ischaemum (L.) Keng var. songarica (Rupr. ex Fisch. &

C.A. Mey.) Celarier & J.R. Harlan, r, 6724

Cenchrus spinifex Cav., o, 6666,99

* Dactyloctenium aegyptium (L.) Willd., i, 6661,95

^Dichanthelium aciculare (Desv. ex Poir.) Gould & C.A. Clark subsp.

neuranthum (Griseb.) Freckmann & Lelong, o, 6404, 22

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark subsp. acumi¬

natum, o, 6405, 23

♦Dichanthelium oligosanthes (Schult.) Gould subsp. oligosanthes,

o, 6417,32

Dichanthelium oligosanthes (Schult.) Gould subsp. scribnerianum

(Nash) Freckmann & Lelong, o, 6406, 24

♦Dichantheliumpolyanthes (Schult.) Mohlenbr., o, 6416,31

Dichanthelium sphaerocarpon (Elliott) Gould, o, 6394,17

* 5 Digitaria arenicola (Swallen) Beetle, i, 6727

* 6 Digitaria texana Hitchc., r, 6726

Eragrostis secundiflora J. Presl subsp. oxylepis (Torr.) S.D. Koch, o,

6572,67

Eustachys petraea (Sw.) Desv., o, 6407,25

Muhlenbergia capillaris (Lam.) Trin., f, 6681,109

*Panicum amarum Elliott, i, 6668,101

Paspalum monostachyum Vasey, o, 6656,90

Paspalum setaceum Michx. var. stramineum (Nash) DJ. Banks, o,

6651,85

§ Schizachyrium scoparium (Michx.) Nash var. scoparium, a, 6570,66

Setariaparviflora (Poir.) Kerguelen, i, 6569,65

Spartinapatens (Aiton) Muhl., f, 6561,60

Spartina spartinae (Trin.) Merr. ex Hitchc., f, 6498,41

Rosen et al., Vascular plants of Follets Island, Brazoria County, Texas

489

Fig. 3. Representative photo of strand prairie vegetation on an elevated mound (center) surrounded by other Gulf-shoreline habitat types including

vegetated and un-vegetated salt-flats (foreground, right, and aft) and salt-marsh and open water (background).

Table 1. Taxonomic summary of vascular plants of the 179 ha Follets Island Conservation Initiative. Superscript numbers represent non-native taxa within the

total for each rank.

Monocots

Eudicots

Total

Families:

(D 34

d ) 4 1

Genera:

(5)29

(3) 76

(6) 105

Species:

(4)44

(3) 86

(7) 130

Sphenopholis obtusata (Michx.) Scribn., o, 6400 Sporobolus virginicus (L.) Kunth, f, 6560, 59

Sporobolus pyromidotus (Lam.) Hitchc., o, 6559,58 Vulpio octofloro (Walter) Rydb. var. octofloro, o, 6397

EUDICOTS

AQUIFOLIACEAE

Ilex vomitorio Aiton, r, 6684,112

AMARANTHACEAE

*Atriplexpentandra (Jacq.) Standi., o, 6649,83

APIACEAE

Ammoselinum butleri (Engelm. ex S. Watson) J.M. Coult. & Rose,

i, 6284

7 Polytaenia texono (J.M. Coult. & Rose) Mathias & Constance, f,

6047, 56

*Ptilimnium nuttollii (DC.) Britton, i, 6496, 39

APOCYNACEAE

Cynanchum angustifolium Pers., o, 6513

ARALIACEAE

Hydrocotyle bonoriensis Lam., i, 6665,98

ASTERACEAE

Ambrosia psilostachya DC., f, 6654,88

Aphanostephus skirrhobasis (DC.)Trel. ex Coville & Branner, i, 6056

*Astranthium ciliatum (Raf.) G.L. Nesom, r, 6517

Baccharis halimifolia L., o, 6655,89

Cirsium horridulum Michx., i, 6411

490

Conoclinium coelestinum (L.) DC., o, 6734,

Coreopsis bosolis (A. Dietr.) S.F. Blake var. bosolis, r, 6765

**Erigeron procumbens (Houst. ex Mill.) G.L. Nesom, o, 6390,9

Goillordio pulchello Foug., f, 6287

Gamochaetapurpurea (L.) Cabrera, i, 6396,19

Heterotheca subaxillaris (Lam.) Britton & Rusby, o, 6292

Iva angustifolia Nutt, ex DC., o, 6650,84

Pseudognaphalium obtusifolium (L.) Hilliard & B.L. Burtt, o, 6300

Pyrrhopappus pauciflorus (D. Don) DC., r, 6772

§ Rudbeckia hirta L. var. angustifolia (T.V. Moore) Perdue, o, 6044,28

Solidago sempervirens L. subsp. mexicana (L.) Semple, f, 6682,110

BRASSICACEAE

Lepidium virginicum L. var. virginicum, i, s.n.

CACTACEAE

*Opuntia stricta (Haw.) Haw., o, 6511

CELASTRACEAE

Lepuropetalon spathulatum Muhl. ex Elliott, f, 6281,4

EUPHORBIACEAE

Chamaesyce maculata (L.) Small, o, 6662,96

Croton capitatus Michx. var. lindheimeri (Engelm. & A. Gray) Mull.

Arg., i, 6664,97

Croton glandulosus L. var. var. lindheimeri Mull. Arg., o, 6659,93

4 Croton punctatus Jacq., o, 6504,47

FABACEAE

Acacia farnesiana (L.) Willd., r, 6403,21

4 Astragalus leptocarpusTorr. & A. Gray, o, 6293,8

9 Baptisia bracteata Muhl. ex Elliott var. laevicaulis (A. Gray ex Can by)

Isely, f, 6291,7

Chamaecrista fasciculata (Michx.) Greene, o, 6575, 70

*Dalea emarginata (Torr. & A. Gray) Shinners, i, 6495,38

*Erythrina herbacea L., r, 6516,57

Indigofera miniata Ortega, i, 6055

Mimosa strigillosa Torr. & A. Gray, i, 6663

Neptuniapubescens Benth. var. pubescens, r, 6520

*Rhynchosia americana (Mill.) Metz, i, 6048

*Rhynchosia minima (L.) DC. var. minima, i, 6049

*Strophostyles helvola (L.) Elliott, o, 6667,100

Strophostyles leiosperma (Torr. & A. Gray) Piper, o, 6565,63

Vida ludoviciana Nutt, ex Torr. & A. Gray, o, 6297

GENTIANACEAE

Eustoma exaltatum (L.) Salisb. ex G. Don, o, 6562,61

§ Sabatia campestris Nutt., o, 6393,16

GERANIACEAE

Geranium carolinianum L., o, 6298

LAMIACEAE

Monarda punctata L., o, 6418,33

4 Scutellaria drummondii Benth., i, 6050

*Teucrium cubense Jacq. var. cubense, i, 6054, 29

LINACEAE

4l0 /./nu/7? alatum (Small) H.J.P. Winkl., o, 6295, 27

4l1 /./nu/7? imbricatum (Raf.) Shinners, o, 6052

Linum medium (Planch.) Britton var. texanum (Planch.) Fernald,

o, 6043

LYTHRACEAE

Lythrum alatum Pursh var. lanceolatum Torr. & A. Gray ex A. Gray,

o, 6580, 73

Journal of the Botanical Research Institute of Texas 9(2)

MALVACEAE

Kosteletzkya virginica (L.) C. Presl ex A. Gray var. althaeifolia Chapm.,

r, 6613, 78

MYRICACEAE

Morelia cerifera (L.) Small, r, 6568

ONAGRACEAE

*Calylophus australis Towner & P.H. Raven, i, 6057,18

Oenothera drummondii Hook., f, 6283, 2

4l2 Oenofhera patriciae W.L. Wagner & Hoch, r, 6420

OROBANCHACEAE

Agalinis heterophylla (Nutt.) Small, f, 6648,82

Agalinis maritima (Raf.) Raf., i, 6053

§ Buchnera americana L., i, 6061

Castilleja indivisa Engelm., o, 6290,6

OXALIDACEAE

Oxalis violacea L., r, 6683,111

PHYTOLACCACEAE

Phytolacca americana L., r, 6688,115

PLANTAGINACEAE

413 Plantago hookeriana Fisch. & C.A. Mey., o, 6408,26

Plantago virginica L., o, 6299,10

POLYGALACEAE

4 Polygala verticillata L, i, 6060

POLYPREMRACEAE

Polypremum procumbens L., i, 6059

PRIMULACEAE

Anagallis arvensis L., r, 6769

Anagallis minima (L.) E.H.L. Krause, o, 6280, 3

Samolus ebracteatus Kunth, i, 6392,15

ROSACEAE

Rubus trivial is Michx., f, 6282,1

RUBIACEAE

Diodia teres Walter var. teres, o, 6657,91

Diodia virginiana L. var. virginiana, o, 6658,92

Houstonia pusilla Schopf, i, 6285

Stenaria nigricans (Lam.) Terrell, o, 6503,46

RUTACEAE

*Zanthoxylum fagara (L.) Sarg., r, 6686,113

SAPOTACEAE

Sideroxylon lanuginosum Michx. var. oblongifolium (Nutt.) B.L.

Turner, r, 6687,114

SOLANACEAE

' ]4 Physalis cinerascens (Dunal) Hitchc. var. spathulifolia (Torr.) J.R.

Sullivan, o, 6389,12

*Solanum triquetrum Cav., r, 6689,116

TAMARICACEAE

*Tamarix ramosissima Ledeb., r, 6519

VERBENACEAE

*Lantana camara L., r, 6677,105

*Phyla nodiflora (L.) Greene, i, 6493,36

§ Verbena halei Small, f, 6402

VITACEAE

4 C/ssus incisa Des Moul., r, 6685

Rosen et al., Vascular plants of Follets Island, Brazoria County, Texas

491

APPENDIX 2: CONSERVATION NOTES

1. Yuccatenuistylo: Endemic to coastal Texas from Galveston County to South Texas, inland to Milam County (Keith 2015).

2. Sisyrinchium biforme: In coastal dunes and inland sandy river banks of Texas, Louisiana, and coastal northern Mexico (Cholewa and

Henderson 2002).

3. Juncus fascinatus: Endemic to streams, seeps, and coastal sites in north-central and southeastern Texas (Knapp 2014).

4. Dichanthelium aciculare subsp. neuranthum: Texas represents the western edge of the range of the species where it is "rather rare-a nice

find" (Freckman 2014).

5. Digitariaarenicola: Endemic to deep coastal sands of Texas and is mapped byWipffand Hatch (1994) as seeing its northern-most records from

adjacent Galveston County. Our collection is the first record from Brazoria County, and the second on a protected site in southeast Texas.

6. Digitariatexana: A Sandy-prairie species known only from the Texas'coast and Rio Grande plains (Correll & Johnston 1970).

7. Polytaenia texana: Described by Nesom (2012) as a "near-endemic" restricted to most of Texas and southern Oklahoma and occurring in

most of the prairie habitat types.

8. Erigeronprocumbens: Restricted to coastal sands in Louisiana, Texas, and Tamaulipas (Correll & Johnston 1970).

9. Baptisiabracteata var. loevicoulis: Based on distributions described by Isely (1990) and Correll and Johnston (1970), we interpret this Boptisio

as endemic to the coastal plain of southwest Louisiana and southeast Texas.

10. Linumalatum: Open areas with sandy soils in south-central Texas and northeast Tamaulipas (Correll & Johnston 1970).

11. Linumimbrication: Open areas with sandy soils in east-central Texas (Correll & Johnston 1970).

12. Oenotherapatriciae: A colonial species of open sandy places throughout central and east Texas (as Gouro brochycorpo Small in Correll

& Johnston 1970).

13. Plantago hookeriana: Correll and Johnston (1970) describe this species as "possibly endemic" and occurring from east Texas along the

coast to the Rio Grande Valley, and inland to the Trans Pecos.

14. Physaliscinerascens var. spathulifolia: Sandy beaches and plains along the Texas coast into Lousiana (Correll & Johnston 1970).

ACKNOWLEDGMENTS

We are grateful to David Riskind and Cherie Obrien for their assistance with access and specific information

about the study site. We thank Jennifer Estes for preparing Figures 1 & 2. We are grateful to Eric Keith, Guy

Nesom, and Robert Freckmann (UWSP) for determinations and Bill Carr, Eric Keith, and an anonymous re¬

viewer for helpful suggestions. We thank Shiron Lawrence for assistance in the held.

REFERENCES

Cholewa, A.F. & D.M. Henderson. 2002. Sisyrinchium. In: Flora of North America Editorial Committee, eds. Flora of North

America north of Mexico. Oxford University Press, New York, U.S.A., and Oxford, U.K. 26:351-371.

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

U.S.A.

Crenwelge, G.W., J.D. Crout, E.L. Griff in, M.L. Golden, & J.K. Baker. 1981. Soil survey of Brazoria County, Texas. United States

Dept, of Agriculture, Soil Conservation Service, Washington, D.C.

Diamond, D.D., D.H. Riskind, & S.L. Orzell. 1987. A framework for plant community classification in Texas. Texas J. Sci.

39:203-222.

Flora of North America Editorial Committee, eds. 1993+. Flora of North America north of Mexico. 16+ vols. Oxford University

Press, New York, U.S.A., and Oxford, U.K.

Freckman, R. 2014. Personal communication: e-mail of 28 May.

Gould, F.W. 1975. The grasses of Texas. Texas A&M University Press, College Station, Texas, U.S.A.

Hollingsworth, T. 1998. Galveston Island State Park Resource Management Plan (Draft). Unpublished Document. Texas

Parks and Wildlife Department, State Parks Division, Austin, Texas, U.S.A.

Isely, D. 1990. Vascular flora of the Southeastern United States. Vol. 3, Part 2, Leguminosae (Fabaceae). Univ. North Caro¬

lina Press, Chapel Hill.

Keith, E. 2015. Personal communication: e-mail of 03 June.

Knapp, W. 2014. Juncus fascinatus (Juncaceae), a new combination in Juncus sect. Ozophyllum and notes on morphologi¬

cally similar species. Phytotaxa 174 (5): 243-260.

Nesom, G. L. 2012. Taxonomy of Polytaenia (Apiaceae): P. nuttallii and P. texana. Phytoneuron 6:1-12.

Prather, L.A., O. Alvarez-Fuentes, M.H. Mayfield, & CJ. Ferguson. 2004. The decline of plant collecting in the United States: A

threat to the infrastructure of biodiversity studies. Syst. Bot. 29:15-28.

Rosen, DJ. 2007. The vascular flora of Nash Prairie a Coastal Prairie remnant in Brazoria County, Texas. J. Bot. Res. Inst.

Texas 1(1 ):679-692.

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Rosen, DJ. 2010. The vascular plants of Mowotony Prairie: A small remnant coastal grassland in Brazoria County, Texas.

J. Bot. Res. Inst.Texas 4(1):489-495.

Rosen, D. J., S. K. Lawrence, & A. Sipocz. 2014. The vascular flora of Galveston Island State Park, Galveston County, Texas. J.

Bot. Res. Inst.Texas 8:339-352.

Rosen, DJ. 2014. Addendum to the vascular flora of Nash prairie, Texas, U.S.A. J. Bot. Res. Inst. Texas 8(1 ):381 —382.

Singhurst, J.R., DJ. Rosen, A. Cooper, & W.C. Holmes. 2014a. The vascular flora and plant communities of Candy Abshier

Wildlife Management Area, Chambers County, Texas, U.S.A. J. Bot. Res. Inst.Texas 8(2): 667-677.

Singhurst, J.R., N. Shackelford, W. Newman, J.N. Mink, & W.C. Holmes. 2014b. The ecology and abundance of Hymenoxys

texana (Asteraceae). Phytoneuron 2014-19:1-19.

Smeins, F.E., D.D. Diamond, & C.W. Hanselka. 1991. Ch. 13, Coastal Prairie. In: R.T. Coupland, ed. Ecosystems of the World 8A;

Natural grasslands, introduction and Western Hemisphere. Elsevier, Amsterdam. Pp. 269-290.

Tropicos.org. Missouri Botanical Garden, http://www.tropicos.org. Accessed January 2015.

Wipff, J.K. & S.H. Hatch. 1994. A systematic study of Digitaria - sect. Pennatae (Paniceae) in the New World. Syst. Bot.

19:613-627.

IN MEMORIAM: ROBERT R. “BOB” KOWAL (23 APRIL 1939-3 AUGUST 2015)

Theodore S. Cochrane

Senior Academic Curator Emeritus

Wisconsin State Herbarium

University of Wisconsin-Madison

430 Lincoln Dr .

Madison , Wisconsin 53706 - 1381 , U . S . A .

Robert R. Kowal, October 1986, in his backyard Packera transplant garden.

Dr. Robert Raymond Kowal, affectionately known as “Bob” to his friends, colleagues, and students, died in

Middleton (a satellite of Madison), Wisconsin, on August 3,2015, at the age of 76 after suffering from the effects

of multiple brain tumors. He was a Professor of Botany at the University of Wisconsin-Madison (1971-1997).

Kowal, who was born and raised in Paterson, New Jersey, developed a love of nature as a young boy. He

was captivated not only by the flowers his mother grew in his family’s garden and the many house plants she

kept inside, but also by butterflies, in which he showed a keen interest while in high school. Armed with basic

equipment, he collected butterflies in the vicinity of home, carefully “relaxing” and spreading his specimens

before mounting them. Having led a rather introverted life while a youth, Kowal “really bloomed” (in the words

of his brother Norman) once in the college environment at Cornell University, where he enrolled as a scholar¬

ship student, majoring in botany, minoring in biometry and statistics as well as genetics, and graduating as

class valedictorian in 1960. He continued his academic training at Cornell, pursuing graduate studies under

the guidance of Robert T. Clausen, and in 1968 earned his doctorate in plant taxonomy and ecology. Kowal’s

dissertation was based on three summers of held research conducted on Senecio in Quebec, and among his

publications (listed below) the published version of this work deserves to be mentioned in particular. Kowal

J. Bot. Res. Inst. Texas 9(2): 493 - 501.2015

494

Journal of the Botanical Research Institute of Texas 9(2)

was also greatly interested in cytotaxonomy and numerical taxonomy, and he developed a good working

knowledge of the International Code of Botanical Nomenclature (while it was still known by this longer title). He

held a post-doctoral fellowship in the biomathematics program at North Carolina State University during

1967-69. His first academic appointment was a visiting assistant professorship in biology at Kansas State Uni¬

versity in 1969-71, where he was a colleague of fellow Senecio enthusiast Theodore M. “Ted” Barkley. In the fall

of 1971 he accepted a faculty position in the Department of Botany at the University of Wisconsin-Madison,

retiring in 1997.

His dual interests in botany and statistics served Kowal well, and he was nationally recognized for his

expertise in the use of biomathematics to understand the genetic variability of natural plant populations. Al¬

though traditional in his own working methods, he was always open to new developments and familiarized

himself with modern trends in botanical research.

A dedicated teacher, Kowal regularly taught a high-enrollment general botany course to an entire genera¬

tion of UW students, and he won the appreciation and fondness of undergraduate majors and graduate stu¬

dents who took his advanced courses on the spring flora of Wisconsin, dendrology, advanced plant taxonomy,

community analysis, biological diversity, and plant geography, as well as seminars on plant taxonomy and

workshops on the taxonomy and evolution of the Compositae. His laboratory and held research focused on

plant taxonomy and evolution; he traveled throughout the U.S., southern Canada, and Mexico, researching the

tribe Senecioneae, especially his passion, the aureoid senecios ( Packera ). Although not a prolific collector, he

always insisted on producing complete, good-quality specimens and labels.

Kowal, who long chaired the Department of Botany’s Greenhouses and Garden Committee, eagerly

shared his delight in plants with staff, friends, and neighbors alike. His exuberant home garden was stocked

with a wide range of plants but was most notable both for its ornaments and for the range of daylily, peony, and

hosta cultivars. For his house and office he favored bold plants like Alstroemeria, Amaryllis, Clivia, and Hibis¬

cus, the brightly colored flowers of which demand admiration. Besides being utterly fascinated with plants,

Kowal was an avid swimmer and sometime jogger and flute player, and he loved classical music (especially

opera) and reading. He recognized the need to protect the plants he loved and their habitats and was a long¬

time generous supporter of environmental organizations and other good causes. In accordance with his wish¬

es, Kowal’s body was donated to the University of Wisconsin School of Medicine.

An individualist with his own obsessions, Kowal was unrepentant about his shabby dress, and even

though he disliked bureaucracy and was uncharitable toward shaky ideas, sloppy botany, and slipshod writing,

departmental staff will nonetheless remember him as a remarkably gentle and patient man. He always found

time to assist students and others who came to him with questions and requests. Above all, he was an excellent

botanist.— Ted Cochrane, Senior Academic Curator Emeritus, University of Wisconsin-Madison

PROFESSIONAL SOCIETIES:

The American Association for the Advancement of Science

The American Society of Plant Taxonomists

The Botanical Society of America

The International Association for Plant Taxonomy

The New England Botanical Club

Society for the Study of Evolution

Torrey Botanical Club

HONORS:

Phi Beta Kappa

Honorary Woodrow Wilson Fellow

Sigma Xi

Phi Kappa Phi

Study leave (sabbatical) for the academic year 1989-90, at Kansas State University and in Mexico

Cochrane, Bob Kowal, In Memoriam

495

PUBLICATIONS:

Kowal, R.R. 1968. Senecio aureus and allied species on the Gaspe Peninsula of Quebec. Ph.D. Dissertation, Cornell Univ.,

Ithaca, NY.

Kowal, R.R. 1970. The origin of Senecio gaspensis (Asteraceae) on the Gaspe Peninsula, Quebec. Amer. J. Bot. 57(No. 6,

Pt. 2):749-750. [Abstract.]

Kowal, R.R. 1971. Disadvantages of the generalized variance as a measure of variability. Biometrics 27:213-216.

Kowal, R.R.&T.M. Barkley. 1973. Senecio anonymus Wood, an earlier name for Senecio smallii Britton. Rhodora 75:211-219.

Kowal, R.R. 1975. Systematics of Senecio aureus and allied species on the Gaspe Peninsula, Quebec. Mem.Torr. Bot. Cl.

23:1-113.

Adams, M.S., R. Anderson, & R.R. Kowal. 1975. Temperature dependence of net photosynthesis in Trientalis borealis Raf. in

Wisconsin. Oecologia 18:199-207.

Woelkerling, W.J., R.R. Kowal, & S.B. Gough. 1976. Sedgewick-Rafter cell counts: A procedural analysis. Hydrobiologia

48:95-107.

Kowal, R.R., MJ. Lechowicz, & M.S. Adams. 1976. The use of canonical analysis to compare response curves in physiological

ecology. Flora 165:29-46.

Kowal, R.R., S.A. Mori, & J.A. Kallunki. 1977. Chromosome numbers of Panamanian Lecythidaceae and their use in

subfamilial classification. Brittonia 29:399-410.

Kowal, R.R. & J. Ruffin. 1979. Generic circumscription in the Xanthocephalum complex (Asteraceae, Astereae) using ca¬

nonical analysis. Bot. Soc. Amer., Misc. Ser., Pub. 157:61. [Abstract.]

Kolterman, D.A. & R.R. Kowal. 1983. The origin of hexaploid Senecio pseudaureus var. semicordatus (Asteraceae,

Senecioneae): The bearing of flavonoid chemistry and morphological data. Amer. J. Bot. 70(No. 5, Pt. 2):119.

[Abstract.]

Kowal, R.R. & D.A. Kolterman. 1983. A measure of dissimilarity for multivariate binary (presence/absence) data with

examples of its use on biochemical data from Cnidoscolus (Euphorbiaceae) and Senecio (Asteraceae). Amer. J. Bot.

70(No. 5, Pt. 2):120. [Abstract.]

Phillips, M.N. & R.R. Kowal. 1983. Systematics of Saxifraga pensylvanica and a putative segregate species, S. forbesii

(Saxifragaceae). Amer. J. Bot. 70(No. 5, Pt. 2):125-126. [Abstract.]

Kolterman, D.A. & R.R. Kowal. 1984. The origin of hexaploid Senecio pseudaureus var. semicordatus (Asteraceae,

Senecioneae): The bearing of flavonoid chemistry and morphological data. Bull. Bot. Cl. Wisconsin 16:39. [Abstract.]

Kolterman, D.A, GJ. Breckon, & R.R. Kowal. 1984. Chemotaxonomic studies in Cnidoscolus (Euphorbiaceae). II. Flavonoids

of C. aconitifolius, C. souzae, and C. spinosus. Syst. Bot. 9:22-32.

Kowal, R.R. 1984. Taxonomy and evolution of the Composites. Photo offset. Dept, of Botany, Univ. of Wisconsin, Madi¬

son. [A workbook for workshop and class use, extensively revised and reissued in 1988, 1991,1997, and 2003 with

the title Keys to the Compositae of Wisconsin.]

Newcomb, E.H., J.M.L. Selker, S.R. Tandon, F.-J. Meng, & R.R. Kowal. 1985. Uninfected cells in ureide- and amide-exporting

legume root nodules. In: P.W. Ludden & J.E. Burris, eds. Nitrogen fixation and C0 2 metabolism: A Steenbock Sympo¬

sium in honor of Professor Robert H. Burris. Elsevier Science Pub. Co., Inc., NY. Pp. 31-40.

Newcomb, E.H., S.R. Tandon, & R.R. Kowal. 1985. Ultrastructural specialization for ureide production in uninfected cells of

soybean root nodules. Protoplasma 125(1 —2):1—12.

Kowal, R.R. 1989. Chromosome numbers of Asteranthos and the putatively related Lecythidaceae. Brittonia 41:131-135.

Kowal, R.R., C.C. Freeman, &T.M. Barkley. 1990. Chromosome numbers of the aureoid senecios: A summary. Amer. J. Bot.

77(6, Supplemental42. [Abstract.]

Kowal, R.R. 1991. A new variety of Senecio (Asteraceae: Senecioneae) from the Sierra de Manantlan, Jalisco, Mexico, with

notes on the S. roldana complex. Brittonia 43:102-115.

Knox, E.B. & R.R. Kowal. 1993. Chromosome numbers of the East African giant senecios and giant Lobelias and their

evolutionary significance. Amer. J. Bot. 80:847-853.

Kowal, R.R. 1994. Chromosome numbers of the aureoid senecios ( Packera ) and their evolutionary significance.

International Compositae Conference, Royal Botanic Gardens, Kew, England. [Abstract.]

Ballard, H.E., Jr., KJ. Sytsma, & R.R. Kowal. 1998. Shrinking the violets: Phylogenetic relationships of infrageneric groups in

Viola (Violaceae) based on internal transcribed spacer DNA sequences. Syst. Bot. 23:439-458.

Mahoney, A.M., V.W. Kong, & R.R. Kowal. 2002. Documented chromosome numbers 2002:3. Chromosome counts for

Packera paupercula var. appalachiana (Asteraceae: Senecioneae). Sida 20:605-610.

496

Journal of the Botanical Research Institute of Texas 9(2)

Mahoney, A.M. & R.R. Kowal. 2008. Three new varieties of Packera paupercula (Asteraceae, Senecioneae) in midwestern

and southeastern North America. Novon 18:220-228.

Kowal, R.R., EJ. Judziewicz, & J. Edwards. 2011. Packera insulae-regalis (Asteraceae, Senecioneae), a new species endemic to

Isle Royale, Michigan, USA. Brittonia 63:343-354.

Kowal, R.R., A.M. Mahoney, A. Weakley, & D. Estes. 2015. Validation and lectotypification of Packera crawfordii (Asteraceae).

Phytoneuron 2015-18:1-2. [e-published.]

Kowal, R.R. & A.M. Mahoney. 2015. Comments on the status of Packera crawfordii (Asteraceae, Senecioneae): A neglected

species of southeastern North America. Brittonia (in press).

NEW TAXA AND NEW COMBINATIONS:

ASTERACEAE Senecio galicianus McVaugh var. manantlanensis Kowal—Brittonia 43(2): 109. 1991.

ASTERACEAE Packera paupercula (Michx.) A. Love & D. Love var. savannarum R.R. Kowal— Novon

18(2):222.2008.

ASTERACEAE Packera paupercula (Michx.) A. Love & D. Love var. pseudotomentosa (Mack. & Bush) R.R.

Kowal— Novon 18(2):224.2008.

ASTERACEAE Packera insulae-regalis R.R. Kowal—Brittonia 63(3):345. 2011.

ASTERACEAE Packera crawfordii (Britton) A.M. Mahoney & R.R. Kowal—Phytoneuron 2015-18:1. 2015.

[e-published.]

GRADUATE STUDENTS:

Ballard, Harvey E., Jr. 1996. Phylogenetic relationships and infrageneric groups in Viola (Violaceae) based on morphol¬

ogy, chromosome numbers, natural hybridization and internal transcribed spacer (ITS) sequences. Ph.D. Disserta¬

tion, Dept, of Botany, Univ. of Wisconsin, Madison.

Fields, Douglas M. 1999. The vascular plants of Taylor County, Wisconsin: An annotated checklist. M.S. Thesis, Land

Resources, Univ. of Wisconsin, Madison.

Kallunki, Jacquelyn A. 1979. Reproductive biology of mixed-species populations of Goodyera (Orchidaceae) in northern

Michigan. Ph.D. Dissertation, Dept, of Botany, Univ. of Wisconsin, Madison.

Kolterman, Duane A. 1978. The bearing of flavonoid chemistry on the origin of hexaploid Senecio semicordatus. M.S.

Thesis, Dept, of Botany, Univ. of Wisconsin, Madison.

Kolterman, Duane A. 1982. Flavonoid chemotaxonomic studies in Cnidoscolus section Calyptrosolen (Euphorbiaceae,

Crotonoideae, Manihoteae). Ph.D. Dissertation, Dept, of Botany, Univ. of Wisconsin, Madison.

Mahoney, Alison M. 2000. Contributions to the systematics of the Packera paupercula complex (Asteraceae: Senecione¬

ae). Ph.D. Dissertation, Dept, of Botany, Univ. of Wisconsin, Madison.

Phillips, Marc N. 1977. Systematics of Saxifragapensylvanica and a putative segregate species, S. Forbesii (Saxifragaceae).

M.S. Thesis, Dept, of Botany, Univ. of Wisconsin, Madison.

Price, Roberta. 1979. The Draba crassa complex (Brassicaceae): Systematics and geography. M.S. Thesis, Dept, of Botany,

Univ. of Wisconsin, Madison.

PERSONAL TRIBUTES:

Some personal reminiscences —Bob Kowal and I were colleagues for more than 30 years. He was a next-door

neighbor in Birge Hall for much of that time, during which he graciously and generously shared his expertise

and lab space, often befriending the graduate students who came and went. I first met Bob during my job inter¬

view in the spring of 1978. He was a kindred spirit, our having shared interests in evolutionary diversification,

within-species variation, and using statistics to analyze this variation. He was also welcoming and humble,

qualities to comfort a new assistant professor. I was happy to accept his generous invitation to stay with him in

his apartment overlooking Lake Mendota later while I was looking for a place to live in Madison. I recall an

apartment overrun with a luxuriance of plants, including a blooming hibiscus. It was clear he took a simple

and direct pleasure in all things botanical.

Bob devoted great time and effort to teaching our flagship Botany 130 course, often working the evening

before as well as rising early those mornings. During the first class of the semester, he would describe the parts

Cochrane, Bob Kowal, In Memoriam

497

of a plant, working with a potted specimen. Once through the leaves, stems, and flowers topside, he would ask

the class: “What are we missing?” Then, sometimes with the question, “Should I?” or a pregnant pause, he

would smash the pot down on the counter to expose the soil and roots. Any slumbering student would jolt

awake, and no one could ever forget the lesson. Succeeding lectures had a train of their own props, including

an actual miniature train to hammer home concepts of DNA transcription and translation.

With graduate students and colleagues, Bob was always more than ready to lend his expertise. Those

seeking instruction in cytology knew that Bob was the “go-to” guy for this, even into his retirement years.

Earlier in his career, he was also the department’s authority for biostatistics, capitalizing on his post-doc work

at North Carolina State. When I asked him once why he didn’t pursue National Science Foundation grant fund¬

ing for his own research, he responded that if he did get a grant, he would have to spend time being an admin¬

istrator and would lose time to do the research himself. It was thus gratifying to see him become so happily

engaged with his research once he retired. [Kowal did submit two proposals to the NSF in the 1970s but was

not successful in gaining support.—T.S.C.]

Those of us who socialized with Bob soon learned of his self-deprecating sense of humor and sometimes

also his maladies. On a postcard he sent back to the office staff from a trip to Puerto Rico, he talked about the

tropical wonders of where he was and added: “Alas, I’m still me.” Many of us knew of his difficulties sleeping at

night. He tried various solutions, but I’m not sure any worked. He correspondingly made it a habit to take an

afternoon nap on a cushion he kept rolled up in his office. Being a regular swimmer, Bob was also remarkably

fit. At the pool, I would watch him keep up a steady pace, lap after lap after lap.

When Bob bought a house with a double-size lot on the west side, he was able to take his love of plants

outdoors. The yard was never the same. Bob was always happy to give tours of his garden, providing a graduate

seminar in the art of cultivating daylilies and Senecio relatives along the way.

We will all greatly miss Bob—for his strengths and his foibles. —Don Waller, Professor of Botany, Univer¬

sity of Wisconsin-Madison

Reminiscences —Bob joined the faculty shortly after I did. He was an intellectual in the department when others

are mostly just clever. And he was a good moderating influence. I felt very close to him. What great strength he

had in retiring early, so as to do his work the way he wanted. He was a gentleman scientist, of which there are

so few these days. He and I both eschewed vulgar careerism, and we felt a special resonance. Bob was a great

scientist, showing how the trappings of success do not amount to much. But his work will be cited long after

that of others when their gimmicks are forgotten. Great things are achieved in humility. I loved his Senecio in-

decorus work on the Gaspe Peninsula from his early days, showing that petals cost. I think he knew of my

special affection. I hope so. He had a full life well lived. I am proud to have known him.— Tim Allen, Professor

Emeritus of Botany, University of Wisconsin-Madison

Reminiscences —I was an acquaintance of Bob’s over the years, knowing him first as a reviewer for research

proposals to the National Science Foundation. I could always count on him for conscientious appraisals in his

specialty areas of biomathematics and the Composite Family, especially anything “senecioid.” He was a good

citizen.

During my 1990-91 sabbatical year as a Visiting Professor at the University of Wisconsin, I interacted

only lightly with Bob but learned from many appreciative students of his teaching talent and dedication, and as

well of his strong support for the Department of Botany Greenhouses and Garden. His own large garden was

an idiosyncratic jungle of delights, rich in plants and oddball ornaments. He was a knowledgeable plantsman.

In 2007 Bob allowed me to stay in his spare bedroom while I paid final visits to a friend in hospice care, a

visit that proved all too brief. Bob was gracious and consoling—a generous human being who will be missed

by many.— Jim Rodman, retired, formerly a program director at the National Science Foundation

Some personal reminiscences —I was Bob’s ultimate-in the sense of “last”-graduate student. Bob was very par¬

ticular about using the right word; he once rather firmly reminded me when I’d used a word incorrectly that

498

Journal of the Botanical Research Institute of Texas 9(2)

“words have meaning!” His graduation gift to me was a two-volume Oxford dictionary. It’s in my office at

Mankato State University, where I teach; I consult it all the time.

I earned a bachelor’s degree in studio art with a minor in French at UW-Oshkosh in 1976. But having

loved plants and gardening since childhood, I decided to return to school with the intention of earning a mas¬

ter’s degree in botany with an emphasis on ecology. I met Bob in 1994 as a student in his marvelous Spring

Flora of Wisconsin course, which convinced me that I wanted to be a plant systematist. I asked Harvey Ballard,

who was Bob’s penultimate graduate student and my lab instructor, if he thought Bob might be willing to take

me on as a grad student. Harvey encouraged me to ask him; I will always be grateful that Bob said, “Yes.”

I remember with great pleasure our preparing to take to the held. The two of us took a 10-day driving trip

through the southeastern U.S. in 1995 to collect living Packera plants for our common garden experiment,

which we set up in his backyard. Supplied with locations of Packera populations of interest from herbarium

specimen labels, we’d follow circuitous driving instructions, and eventually Bob would pull his Subaru station

wagon to the roadside. We’d get out and gird up our loins. We’d tuck in our shirts, roll down our sleeves, button

our cuffs, and tuck our pants legs into our socks. Then we smeared a yellow copper sulfate powder that Bob

used to prevent powdery mildew on his plants around our waists and shirt and pants cuffs to prevent chiggers.

I guess it worked. In his breast pocket, Bob carried his hard-cover pocket-sized notebook and a fine-tipped

Rapidograph pen, a fussy instrument that frequently clogged and had to be shaken before it would write. He

strapped on a bag containing his camera, labels on string, and his prized trowel with the wooden handle worn

to the shape of his palm.

Bob had a pile of tough plastic bags into which we put our quarry: Packera plants that were kept alive in

coolers for the duration of our trip. He tied an enormous plastic bag to his belt into which he put our smaller

bags and other plants of interest to be keyed out later. Despite the heat and humidity, Bob’s long legs carried

him tirelessly up and down steep slopes and straight through brush. I scrambled after him. I think we must

have looked a bit like Don Quixote and Sancho Panza.

Bob was exacting, precise, patient, and kind. Just about everything I know about collecting and pressing

plants, keeping records, and writing clearly comes from him. When I teach my Flora of Minnesota class, most

of it comes right out of his course at the UW. I tell my students, when they’re sloppy, “words have meaning,” and

hope that in some small way Bob’s passion for plants and quest for precision in record-keeping and the use of

language pass through me and on to the next generation of botanists. Thank you, Bob! —Alison Mahoney, Pro¬

fessor of Botany, University of Minnesota-Mankato

Reminiscences —I had met Bob during an early visit to the University of Wisconsin campus and reinitiated

contact with him via e-mail prior to my application to the Ph.D. program in the Department of Botany. Bob

expressed interest in serving as my major professor to train me in traditional systematic approaches and sug¬

gested that Ken Sytsma and Hugh litis could fill in the other areas. Immediately upon arriving for my first se¬

mester in Madison, I hunted down Bob’s office and excitedly introduced myself. He seemed eager to have a new

graduate student. Over the next couple of weeks, I checked in with him first thing every day—why, I’m not

sure—and we chatted briefly about what he was doing that day and what I was doing or attempting to do. At

the end of each day, this scenario was repeated. Sometimes these conversations became quite animated when

something one of us mentioned sparked great interest, as we both thoroughly enjoyed discussing taxonomy

and nomenclature, systematic theory, evolution, and specific methods. He never told me he was too busy, and

I never got the sense that I was out of line; but, soon becoming concerned that I might be abusing his accessibil¬

ity, one afternoon I asked him whether I was taking too much of his time or bothering him. He cocked his head

as a thoughtful look passed across his face. “No, no,” he said. “I’ll let you know when I’m too busy. I’m enjoying

our conversations, and it’s been awhile since I’ve had much opportunity to discuss taxonomy or other things

with anybody.” I learned from Bob’s example that students come first unless something desperately important

was looming and requiring my time. I also discovered that opportunities to discuss aspects of our discipline

with Bob and others in the department were just as important for my training and professional development as

coursework, research, and teaching.

Cochrane, Bob Kowal, In Memoriam

499

My teaching assistant positions included Bob’s introductory botany course. There, I witnessed his teach¬

ing style, a mixture of a little overacting, a healthy dose of humor, an occasional skit or demonstration to em¬

phasize a point, and lots of enthusiasm for the subject. I learned from Bob that with new science majors or

non-majors, it is critically important to capture the students’ attention, and he had worked out for himself the

most successful pedagogical approaches. Bob could make every class period exciting and memorable, and even

after teaching the same course for decades, he showed no sign of becoming jaded or bored. This was clear proof

of how much he invested in the course to provide his students with a quality experience. I later incorporated

these lessons into my own teaching.

As my research progressed and I began to write manuscripts, I discovered Bob’s outstanding capacity for

writing and editing. In the first few months of our regular interactions, he bought me Strunk and White’s little

hardbound book The Elements of Style and urged me to read it cover to cover, which I did. I was dismayed at the

extent of his marks and comments on every chapter I gave him for review; sometimes we worked through

several bouts of changes and corrections, but I ultimately benefited from his placing great value on precision

and correctness. It wasn’t that Bob necessarily had to be right, but that he wished everyone to be as precise as

he could. He was especially dismayed by my fervor to use ten words where one would do, as he put it, or to write

florid language instead of plain English. He constantly preached that good writing—scientific or otherwise—

should be terse, precise, and simple to understand. His lessons were strikingly apparent in a paper based on a

dissertation chapter, in which we discussed the evolution of the Hawaiian Viola lineage inferred from DNA

internal transcribed spacer (ITS) sequences. I gave Bob the “semi-final” draft to review. After three rounds of

corrections, he essentially dismantled it and put back together in a quite different format. We submitted the

manuscript to the journal Evolution. One of the reviewers stated that it was one of the cleanest and most beauti¬

fully written he had ever had the pleasure to review. I knew that assessment derived from Bob’s handiwork. I

am more aware of what makes good writing thanks to Bob’s perspective, and when criticizing my own and

others’ writing while wearing my “Bob hat,” I’ve been able to tighten up the prose significantly.

Bob referred to himself as a “biosystematist,” one who applies a wide range of methodologies to delineate

evolutionarily distinct sets of populations, and his having employed statistics, cytogenetics, and observations

from held and test garden studies while studying the difficult group Packera led me to identify him as a possible

mentor. Since high school, I had been engaged in traditional taxonomic studies of plants, especially violets, the

species of which are often difficult to distinguish, and had begun conducting phenetic studies, primitive flavo-

noid identification, and SEM examination of leaves and Bowers of violets when I got to the UW-Madison. Bob

enthusiastically supported my interest in gaining experience with any approach that might prove taxonomi-

cally useful. He was more than willing to provide as much guidance or assistance as one could want, and he

strengthened and extended my statistical understanding and competence. At one point I requested that he give

me some help in obtaining chromosome counts for the very rare Viola frank-smithii. He advised me on possible

Bxation solutions for Bower buds, information I passed along to a collaborator in Utah, who gathered buds and

sent them to us. For the next three months, Bob took time to show me how to prepare tissue for squashes and

how to examine slides for good squashes. Freely acknowledging the limits of his own experience, he was eager

to see me learn molecular systematic methods under Ken Sytsma’s tutelage. I have continued Bob’s legacy by

passing on serious training in traditional methods and nomenclature, as well as molecular approaches, to all

my graduate students and many undergraduates.

While contributing in numerous ways to the professional botanist I am today, Bob also quickly became a

good friend and eventually a very dear friend. During the early weeks of our time in Madison, my partner and

I began meeting Bob and his partner Jim every Friday evening for supper and sometimes a cultural perfor¬

mance or a movie afterwards. Bob and Jim became our “big brothers,” and we relished the socializing and

mutual venting sessions we shared. Bob and I also made many trips to conferences together (during which I

always brought my earplugs with me to muffle his snoring), as well as occasional trips into the field. I shared

Bob’s enthusiasm, nay, childlike delight, in natural diversity, and we had great fun simply studying plants, hike

most visitors to Bob’s home, I was subjected to the inevitable garden tour, but no matter what the season, it was

500

Journal of the Botanical Research Institute of Texas 9(2)

always a treat. Like the man himself, the gardens, which occupied the back, front, and sides of the house, were

slightly disheveled, but their diversity (especially of day lily and peony cultivars) was astonishing for the mod¬

est size of the yard.

While working assiduously at his job, Bob also made time for his partner, his friends, and especially his

beloved garden, and he continued investing regular amounts of time into his research long after retirement.

This strategy, too, has been a continuing lesson I still attempt to apply in my own life. I am grateful to have been

mentored and inspired by Bob Kowal, but I am even more grateful for the close friendship we shared. He was

funny, thoughtful, passionate, compassionate, and unpretentious. He made time for others and took delight in

sharing his knowledge of and enthusiasm for plants. He was an outstanding role model for me, professionally

and personally, and I miss him greatly.— Harvey Ballard, Jr., Associate Professor of Plant Biology, Ohio

University

Remembrances and reflection —Memories I have of Bob include those dating from December 1974 and January

1975, when he accompanied taxonomy graduate students on a plant collecting trip to Central America. Dumb¬

founded customs officials at the international airport of Panama City didn’t know how to react to his bedroll—

a sponge rubber mattress complete with sheets we dubbed “the marshmallow”—taller than he was and bigger

around than he could reach. One of the first plants we learned while there was Scheelia zonensis, a conspicuous

feather palm along the Pacific coast of Panama. Now, Bob was a very intelligent man who was meticulous about

Latin pronunciations, but for some reason he couldn’t learn the name Scheelia. It invariably came out wrong, no

matter how often he spoke it. However, he did take better quality color transparencies than any of the rest of us.

Soon after lunch on the way back to Madison we stopped at the Audubon Society’s famous Corkscrew Swamp

Sanctuary in the western Everglades. While the rest of us were marveling at the old-growth bald cypress trees

and many epiphytes, I could see that Bob was growing more and more anxious. Suddenly, he asked me for the

car keys, turned, and ran back the length of the boardwalk. When he returned a while later, I looked at him

quizzically. “I had to brush my teeth,” he answered. In fact, he brushed his teeth so often that eventually his

dentist told him to cut back; he was damaging the enamel. Among the many other things I remember about

Bob are his digging bomb craters all over his yard in which to plant plants, his diverse collection of baseball-

style caps (many faded or dirty), his creative and hilarious blackberry costume for a Halloween party, his ca¬

pacity for ingesting voluminous amounts of food, the large quantities of olive oil he used when cooking, and

how loudly and continuously he could snore—all night. The first and last night I ever spent in the same room

with him (I made sure of that!), his snoring prevented me from falling asleep for even one second from 11:00

p.m. ‘til 7:00 a.m. Fellow sufferer Andrew Hipp, sacked out on the floor in the adjoining porch, crawled over to

me on his hands and knees at one point in the early morning and exclaimed, “Ted! That’s Bob snoring! I

thought the house was breaking!”

It was a testament to Bob’s commitment to teaching that each semester, year after year, when it was ap¬

proaching 9:00 or later at night and I was on my way out of the building, he would be seated in his office under

that big north-facing window he loved, writing out anew, longhand, the next day’s lecture or the next week’s

exam. He was not one to reuse the same notes from one year to the next. The exams he gave in his advanced

taxonomy class reminded us students of the high standards to which we were being held. After all, he was com¬

mitted to high standards for himself. Bob was a critical thinker. During plant taxonomy seminars he would

quickly see the crux of matters being discussed and express the points he made with great clarity. In depart¬

mental committee meetings he was extremely conscientious, fair-minded, and reasonable.

Bob was one of a kind, a singular individual, his own man, even slightly eccentric. Neither charming nor

cantankerous, he was, well, Bob—a man who was who he was at every moment. The idiom “what you see is

what you get” is an accurate portrayal. No one could be more down to earth, more direct, more honest than Bob

was. What he told you in private was exactly what he would say in public. Bob was absolutely devoid of pre¬

tense, wholly accountable in all that he said and all he supported.

Bob was a friend who showed me more loyalty than I thought I was due. But at the same time he could also

be an unsparing critic, albeit an unreservedly helpful one, when, for example, reading theses and manuscripts.

Cochrane, Bob Kowal, In Memoriam

501

I had enormous respect for Bob’s carefully thought-out positions, and we could talk about anything, as we

did, for example, on long car rides traveling to the Annual Systematics Symposium at the Missouri Botanical

Garden: taxonomic questions; environmental preservation and conservation, including the heedless destruc¬

tion of biological diversity, the mindless consumption of material possessions, and the unsustainable increase

in human population; politics; social injustices; personal finances; music; sex; death. He always spoke the

truth as he knew it, about his views, his feelings, and his life.

A profound joy in living marked Bob’s life, and a remarkable serenity as he lived out his last days. I will

miss his wisdom, pragmatism, and friendship.— Ted Cochrane, Senior Academic Curator Emeritus, University of

Wisconsin-Madison

Remembering Bob Kowal —As a 19-year-old transfer student in 1974,1 picked a major called “Biological Aspects

of Conservation” and was assigned an advisor, whom I was told to go find and talk to about program require¬

ments. The memories of navigating Bascom Hill, finding Birge Hall, and knocking on the door of my advisor,

Bob Kowal, remain unusually vivid. The man sitting across the desk from me was youngish, serious, and, I

suspected, somewhat unenthused about having to speak to another young undergrad seeking advice about

course requirements for a major that was somewhat outside of his usual purview.

Fast forward to 1989, when I was offered the position of Head of the Biology Library in Birge Hall. Bob

came into the library often in those days, and every spring he would bring the staff a vase of beautiful tree peo¬

nies from his garden. I gradually came to know him better and to like him more. I respected him for his schol¬

arly, serious manner and for being one of the more constant faculty users of the library. I liked him for his

quirky, irreverent sense of humor and, as years went on, for being a caring person, a mensch. Our relationship

evolved slowly so that I became happy at the sight of Bob walking through the library door, because we would

converse not only about botanical papers, but also about sleep deprivation, depression, and other more per¬

sonal concerns. Besides the transcendent beauty and wonder of the plant world, we had some kind of mutual

understanding about the vicissitudes of life.

During those years Bob began asking if I would like any daylilies from his garden. “Yes,” I would answer,

and soon he would stop by and leave a plastic bag containing the plants in the shade near the front door. Once,

when we were crossing University Avenue together, I made the mistake of mentioning how fond I’d become of

the “Co-ryd-A-lis” in my garden. Bob startled me when he exclaimed at the top of his lungs, “No, No, No! The

accent is on the antipenultimate syllable! It’s not ‘Co-ryd-A-lis,’ it’s ‘Co-RYD-a-lis.’ Three syllables from the end,

not two! Nothing drives me crazier than people putting the accent on the wrong syllable!” It was a memorable

lesson that I never forgot and still check myself on, even today.

Beginning in 20061 had a series of setbacks to my health. Bob would bring me plants from his garden and

transplant them for me once yard work became hard for me to do. We began going to matinees together on a

somewhat regular basis. I loved going to the movies with Bob, because it meant I had a chance to enjoy his

quirky humor and gentle friendship. I loved it when he laughed spontaneously and loudly at places in the

movie where the rest of the audience was silent. He was so much his own person. Our matinee afternoons were

often followed by a stop at the adjacent Pendleton store (I learned that orange was Bob’s favorite color and that

he was susceptible to spending money on shirts) and dinner out. Bob would pick up the tab, saying that he had

more money than he knew what to do with. One time Bob invited me to go to the opera with him, saying that

I should rest assured he would get the best seats available. They turned out to be in the first row, center.

These are but a few of many memories I have of Bob. He was much loved by all of us, because he was

funny, smart, quirky, kind, and generous. He helped me greatly after I retired and was struggling with physical

pain and isolation. He brightened my life. One could not have had a better friend.— Elsa Althen, former head of

the Biology Library, University of Wisconsin-Madison

Reviewers

Volume 9 (2015)

503

108 reviewers: several individuals reviewed more than one manuscript.

We sincerely apologize if your name was accidently left out.

Thank you for supporting Journal of the Botanical Research Institute of Texas.

Almeda, Frank

Applequist, Wendy

Axelrod, Franklin

Aymard C., Gerardo

Baker, Mark

Barrie, Fred R.

Becquer, Eldis

Brock, Mason T.

Brown, Larry E.

Bruneau, Anne

Cadee, Gerhard C.

Carter, J.R.

Clancy, Keith E.

Clark, Ross C.

CumbicusTorres, Nixon Leonardo

Darwin, Steven P.

Delgado Salinas, Alfonso

Dillon, Michael O.

Dransfield, John

Encina Dominguez, Juan A.

Endress, Peter

Escobar Lechuga, Roberto

Estes, Dwayne

Estrada, Armando

Ezcurra, Cecilia

Foroughbakhch Pournavab, Rahim

Gandhi, Kanchi N.

Gardner, Elliot

Gornall, Richard J.

Grayum, Michael

Hardy, Christopher R.

Ha uk, Warren

Haynes, Robert R.

Herrera, F.

Hoagland, Bruce

Hovenkamp, P.H.

Hufford, Larry

Islam, Melissa

Janovec, John

Judziewicz, Emmet J.

Keener, Brian R.

Keith, Eric L.

Kiger, Robert

Kostel, Grace

Krings, Alexander

Lemke, David E.

Lesica, Peter

Ley, Alexandra C.

Linares, Jose

Longhi-Wagner, Hilda Maria

Lorence, David H.

Lundberg, Anders

MacRoberts, Michael

Manchester, Steven R.

Marsico, Travis

Maslin, Bruce R.

McDonald, Andrew

Meave, Jorge A.

Medel, Alfonso

Morgan, David

Neill, David A.

Nelson, Cyril H.

Nelson, John B.

Nesom, Guy

Norman, Eliane M.

Northrup, Katie

O'Kennon, Bob

Ortiz, Orlando O.

Owens, Chetta

Penskar, Mike

Poindexter, Derick B.

Poole, Jackie

Popescu, loana

Powell, A. Michael

Prance, Sir lain

Riggins, Chance

Rohwer, Jens G.

Rosen, David J.

Sadie, Jimi

Salywon, Andrew

Sanchez-Gonzalez, Arturo

Semple, John C.

Sivadasan, M.

Sousa S., Mario

Spellenberg, Richard

Stern, Stephen

Stevanovic, Vladimir B.

Strother, John L.

Suarez Suarez, Luz Stella

Sundell, Eric

Sundue, Michael

Taylor, Carl

Tucker, Gordon C.

Ulloa Ulloa, Carmen

Urbatsch, Lowell

Valdes Reyna, Jesus

Vanderplank, Sula

Vanderwier, Julie

Villarreal Quintanilla, Jose Angel

Villasenor Rios, Jose L.

Waines, J.G.

Weakley, Alan

Wendt, Tom

Wiersema, John H.

Wilder, George J.

Witsell,Theo

Yatskievych, George

Whittemore, AlanT.

Index to 51 Titles with 102 Authors

Volume 9 (2015)

We know you have a choice.

Thank you for choosing and supporting Journal of the Botanical Research Institute of Texas.

A checklist of vascular plants at the Gulf-shoreline extent of coastal

prairie in southeast Texas, U.S.A. by David J. Rosen, Siavash

Zamirpour, and Andrew Sipocz—9(2):485

A survey of the woody and climbing plants of the Refugio de Aves

Dr. Alexander Skutch, "Los Cusingos," Perez Zeledon Canton,

Costa Rica by Ronald L. Jones, Humberto Jimenez-Saa, and

Allen C. Risk—9(1):149

Addendum to the vascular flora of the Hancock Biological Station,

Murray State University, Calloway County, Kentucky, U.S.A. by

J. Richard Abbott and Ralph L. Thompson—9(1 ):229

Addendum: A gasteroid fungus, Palaeogaster micromorpha

gen. & sp. nov. (Boletales) in Cretaceous Myanmar amber by

George Poinar, Jr., Donis da Silva Alfredo, and luri Goulart

Baseia—9(1 ):135

Addendum: Prioria dominicana sp. nov. (Fabaceae: Caesalpini-

oideae), a fossil flower in Mid-Tertiary Dominican amber by

George O. Poinar, Jr. and Kenton L. Chambers—9(2):381

Arachis glabrata (Fabaceae) new to the flora of Louisiana, U.S.A.

by Charles M. Allen—9(2):475

Asteraceaeen el Duraznoy cercanias, nortedeTamazula, Durango

(Mexico): Riqueza, distribucion y endemismo by David Ramirez

Noya y Yolanda Herrera Arrieta—9(2):453

Birch (Betula, Betulaceae) bark horns and similar instruments in

Norway byTorbjorn Aim—9(2):433

504

Journal of the Botanical Research Institute of Texas 9(2)

Botrychium michiganense sp. nov. (Ophioglossaceae), a new

North American moonwort by Arthur V. Gilman, Donald R.

Farrar, and Peter F. Zika—9(2):295

Breeding system and sex ratio variation in mulberries (Moms,

Moraceae) by Madhav P. Nepal, Carolyn J. Ferguson, and Mark

H. Mayfield—9(2):383

Buddleja davidii (Scrophulariaceae) naturalized populations in

Tennessee (U.S.A.) and its woody associates by Ralph L.Thomp-

son and Katrina Rivers Thompson—9(1 ):213

Calathea galdamesiana (Marantaceae), a new endemic Panama¬

nian species by Helen Kennedy and Rodolfo Flores—9(2):319

Casearia draganae, a new species of Samydaceae from western

Colombia and Ecuador by Mac H. Alford—9(2):325

Changes in taxonomic rank for a Hexastylis (Aristolochiaceae)

taxon of the southeastern United States by Brian R. Keener

and LJ. Davenport—9(2):317

Comopellis presbya (Rhamnaceae) gen. et sp. nov. in Mid-Tertiary

amber from the Dominican Republic by Kenton L. Chambers

and George O. Poinar, Jr.—9(2):361

Corrigendum: New combinations in Coryphantha and Escobaria

(Cactaceae) by Root Gorelick—9(2):351

Cuatro nuevas especies y un nuevo registro de Freziera (Pen-

taphylacaceae) de Ecuadory Peru by Daniel Santamaria-Aguilar

y Laura P. Lagomarsino—9(1 ):89

Cuscatlania vulcanicola (Nyctaginaceae) nuevo registro del genero

y especie para la flora de Honduras by Jose Ledis Linares, Frank

Sullyvan Cardoza Ruizy Patricia Hernandez-Ledesma—9(1 ):167

Cyperus granitophilus (Cyperaceae), a granite outcrop endemic,

new for Texas and Oklahoma (U.S.A.) by Robert J. O'Kennon

and Kimberly Norton Taylor—9(1 ):251

Dasylarynx anomalus gen. et sp. nov., a tubular monocotyledon¬

like flower in Mid-Tertiary Dominican amber by George O.

Poinar, Jr. and Kenton L. Chambers—9(1 ):121

First collection of Miconia alainii (Melastomataceae: Miconieae)

with flowers by Walter S. Judd, Teodoro Clase, and Lucas C.

Majure—9(2):449

Floristics and community ecology of aquatic vegetation occurring

in seven large springs at Ozark National Scenic Riverways,

Missouri (U.S.A.), 2007-2012 by David E. Bowles and Hope R.

Dodd—9(1):235

From Cro-Magnon to Krai: A history of botany in Alabama by LJ.

Davenport—9(2):397

Furrowed blister pods stranded on northern Atlantic Ocean coasts

represent an undescribed Sacoglottis (Humiriaceae) endocarp

most similar to the fossil Sacoglottis costata by Raymond van

der Ham,Tinde van Andel, Linda Butcher, Izumi Hanno,Theo

Lambrechts, Kim Lincicome, Paul Mikkelsen, Jenifer Mina, Bob

Patterson, Ed Perry, and Mike Romance—9(1 ):137

Gnaphaliothamnus nesomii (Asteraceae: Gnaphalieae), a new

species from Guatemala and nomenclatorial changes by

Michael O. Dillon and Federico Luebert—9(1 ):63

In memoriam: Robert R."Bob"Kowal (23 April 1939-3 August 2015)

by Theodore S. Cochrane—9(2):493

Irenodendron, a new genus of Samydaceae from South America

by Mac H. Alford and Angela D. Dement—9(2):331

Klaprothiopsis dyscrita gen. et sp. nov. (Loasaceae) in Mid-Tertiary

Dominican amber by George O. Poinar, Jr., Maximilian Weigend,

and Tilo Henning—9(2):369

Matelea chihuahuensis (Apocynaceae): an addition to the flora

of the United States and a synopsis of the species by Angela

McDonnell, Mark Fishbein, Meg Quinn,Trevor Hare, and Kevin

Keith—9(1 ):187

Micranthes rufopilosa (Saxifragaceae) comb, nov.: an alpine

species from Alaska and Yukon by David F. Murray and Reidar

Elven—9(1 ):7

Nautilocalyx rugosus (Gesneriaceae), a new species from the rio

Cenepa watershed (Amazonas, Peru) by Rocio del P. Rojas G.

and John L. Clark—9(2):345

Nephrolepis cordifolia (Nephrolepidaceae) naturalized in south¬

ern California (U.S.A.): with notes on unintended consequences

of escaped garden plants by Richard E. Riefner, Jr. and Alan R.

Smith—9(1):201

New combinations in Coryphantha and Escobaria (Cactaceae) by

Root Gorelick—9(1 ):25

New combinations in Eumachia (Rubiaceae) for species occur¬

ring on the Guiana Shield by Piero G. Delprete and Joseph H.

Kirkbride, Jr.—9(1 ):75

New combinations in Hexasepalum (Rubiaceae: Spermacoceae)

by Joseph H. Kirkbride, Jr. and Piero G. Delprete—9(1 ):103

New co-occurrence of Schoenoplectiella hallii and S. saximon-

tana (Cyperaceae) in Ohio (U.S.A.): Conservation implications

for both species by Paul M. McKenzie, Daniel W. Boone, Marian

Smith, and Richard L. Gardner—9(2):477

New species of Acanthaceae from Ecuador by Dieter C.

Wasshausen—9(1 ):81

New species of Senegalia (Fabaceae) from South America by David

S. Seigler and John E. Ebinger—9(2):335

Notes on Eritrichium (Boraginaceae) in North America II by David

F. Murray—9(2):311

nrDNA sequence (ITS and ETS) from herbarium specimens reveals

phylogenetic affinities of Erigeron geiseri (Asteraceae)

by Richard D. Noyes, William Caraway, and Dulcinea V.

Groff—9(1 ):11

Nuevas especies de Macherium (Leguminosae: Papilionoi-

deae: Dalbergiae) en Mexico y Centroamerica by Jose L.

Linares—9(1 ):49

Observations about Phaseolus lignosus (Leguminosae: Papilionoi-

deae: Phaseoleae), a bean species from the Bermuda Islands

by D.G. Debouck—9(1 ):107

Phytogeographical relationships and analysis of the flora of South-

Central Texas, U.S.A. byA.A. Saghatelyan—9(1):259

Prioria dominicana sp. nov. (Fabaceae: Caesalpinioideae), a fossil

flower in Mid-Tertiary Dominican amber by George O. Poinar,

Jr. and Kenton L. Chambers—9(1):129

Pseudhaplocricus hexandrus gen. et sp. nov. (Commelinaceae)

in Mid-Tertiary Dominican amber by George O. Poinar, Jr. and

Kenton L. Chambers—9(2):353

Redescubrimiento de Desmodium angustifolium (Fabaceae) en

El Salvador by Pablo Galan—9(2):471

Scleria bellii (Cyperaceae), a distinctive and uncommon nutsedge

from the southern U.S., Cuba, and Mexico by Richard J. LeBlond,

Samantha M.Tessel, and Derick B. Poindexter—9(1 ):31

Structural analysis of shrublands adjacent to the Monterrey

Metropolitan Area, Mexico by Pamela Anabel Canizales-

Velazquez, Oscar Alberto Aguirre-Calderon, Eduardo Alanis-

Rodriguez, Eduardo Javier Trevino-Garza, and Jose Manuel

Mata-Balderas—9(1 ):173

Styrax peltatus (Styracaceae), a new species from Oaxaca, Mexico

by Peter W. Fritsch—9(1):43

The first naturalized occurrence of the genus Forsythia (Oleaceae)

in Arkansas (U.S.A.), with additional noteworthy angiosperm

records for the state by Brett E. Serviss,Tyler L. Childs, Sydney S.

Grant, Tiffany A. Graves, Ethan Holicer, Seth A. McBroom, Logan

Thomas, James H. Peck, and Allen Leible—9(1 ):195

Typification of Sedum rubroglaucum (Crassulaceae) by Peter F.

Zika—9(1 ):1

Index

Index of 102 Authors

Volume 9 (2015)

Thank you for choosing Journal of the Botanical Research Institute of Texas.

505

Abbott, J. Richard—9(1 ):229

Aguirre-Calderon, Oscar Alberto—9(1):173

Alams-Rodriguez, Eduardo—9(1 ):173

Alford, Mac H.—9(2):325, 331

Allen, Charles M.—9(2):475

Aim, Torbjorn—9(2):433

Baseia, luri Goulart—9(1 ):135

Boone, Daniel W.—9(2):477

Bowles, David E.—9(1 ):235

Butcher, Linda—9(1 ):137

Canizales-Velazquez, Pamela

Anabel—9(1 ):173

Caraway, William—9(1 ):11

Cardoza Ruiz, FrankSullyvan—9(1 ):167

Chambers, Kenton L.—9(1 ):121,129;

9(2):353, 361,381

Childs, Tyler L.—9(1 ):195

Clark, John L.—9(2):345

Clase, Teodoro—9(2):449

Cochrane, Theodore S.—9(2):493

Davenport, L.J.—9(2):317, 397

Debouck, D.G.—9(1 ):107

Delprete, Piero G.—9(1 ):75,103

Dement, Angela D.—9(2):331

Dillon, Michael O.—9(1):63

Dodd, Hope R.—9(1 ):235

Ebinger, John E.—9(2):335

Elven, Reidar—9(1 ):7

Farrar, Donald R.—9(2):295

Ferguson,Carolyn J—9(2):383

Fishbein, Mark—9(1 ):187

Flores, Rodolfo—9(2):319

Fritsch, Peter W.—9(1 ):43

Galan, Pablo—9(2):471

Gardner, Richard L.—9(2):477

Gilman, Arthur V.—9(2):295

Gorelick, Root—9(1 ):25; 9(2):351

Grant, Sydney S.—9(1 ):195

Graves, Tiffany A.—9(1 ):195

Groff, Dulcinea V.—9(1 ):11

Hanno, Izumi—9(1 ):137

Hare, Trevor—9(1 ):187

Henning, Tilo—9(2):369

Hernandez-Ledesma, Patricia—9(1 ):167

Herrera Arrieta, Yolanda—9(2):453

Holicer, Ethan—9(1 ):195

Jimenez-Saa, Humberto—9(1 ):149

Jones, Ronald L.—9(1 ):149

Judd, Walter S.—9(2):449

Keener, Brian R.—9(2):317

Keith, Kevin—9(1 ):187

Kennedy, Helen—9(2):319

Kirkbride, Jr., Joseph H.—9(1):75,103

Lagomarsino, Laura P.—9(1):89

Lambrechts, Theo—9(1 ):137

LeBlond, Richard J.—9(1 ):31

Leible, Allen—9(1):195

Linares, Jose Ledis—9(1):49,167

Lincicome, Kim—9(1 ):137

Luebert, Federico—9(1 ):63

Majure, Lucas C—9(2):449

Mata-Balderas, Jose Manuel—9(1 ):173

Mayfield, Mark H.—9(2):383

McBroom, Seth A.—9(1 ):195

McDonnell, Angela—9(1 ):187

McKenzie, Paul M.—9(2):477

Mikkelsen, Paul—9(1 ):137

Mina, Jenifer—9(1 ):137

Murray, David F.—9(1 ):7; 9(2):311

Nepal, Madhav P.—9(2):383

Noyes, Richard D. — 9(1 ):11

O'Kennon, Robert J.— 9(1 ):251

Patterson, Bob— 9(1 ):137

Peck, James H. — 9(1 ):195

Perry, Ed— 9(1 ):137

Poinar, Jr., George O.— 9(1 ):121,129,135;

9(2):353, 361,369, 381

Poindexter, Derick B.—9(1 ):31

Quinn, Meg—9(1):187

Ramirez Noya, David—9(2):453

Riefner, Jr., Richard E.—9(1 ):201

Risk, Allen C.—9(1):149

Rojas G., Rocio del P—9(2):345

Romance, Mike—9(1 ):137

Rosen, David J.—9(2):485

Saghatelyan, A.A.—9(1):259

Santamaria-Aguilar, Daniel—9(1):89

Seigler, David S.—9(2):335

Serviss, Brett E.—9(1 ):195

Silva Alfredo, Donis da—9(1 ):135

Sipocz, Andrew—9(2):485

Smith, Alan R.—9(1 ):201

Smith, Marian—9(2):477

Taylor, Kimberly Norton—9(1 ):251

Tessel, Samantha M.—9(1):31

Thomas, Logan—9(1 ):195

Thompson, Katrina Rivers—9(1 ):213

Thompson, Ralph L.—9(1 ):213, 229

Trevino-Garza, Eduardo Javier—9(1 ):173

van Andel,Tinde—9(1 ):137

van der Ham, Raymond—9(1 ):137

Wasshausen, Dieter C.—9(1 ):81

Weigend, Maximilian—9(2):369

Zamirpour, Siavash—9(2):485

Zika, Peter F.—9(1 ):1; 9(2):295

Botanical Names and Subject Index

Volume 9 (2015)

NEW NAMES AND TYPIFICATIONS (75) IN BOLD FACE

Acacia rigidula—9(1 ):173

Acanthaceae—9(1 ):81

Agave lecheguilla—9(1 ):173

Alaska (U.S.A.)—9(1 ):7

Alexander Skutch Biological Corridor—

9(1 ):149

Alpine—9(1 ):7

Amber—9(1 ):121,129

Apocynaceae—9(1 ):187

Aquatic vegetation—9(1 ):235

Arachis glabrata—9(2):475

Arbuckle Uplift (U.S.A.)—9(1 ):251

Aristolochiaceae—9(2):317

Arkansas (U.S.A.)—9(1 ):195

Asteraceae—9(1 ):11,63; 9(2):453

Atlantic Ocean—9(1 ):137

Bartram, William—9(2):401

Bermuda Islands—9(1 ):107

Betula—9(2):433

Betulaceae—9(2):433

Big Bend Region (Texas, U.S.A.)—9(1 ):259

Birch Bark Horns—9(2):433

Boletales—9(1 ):135

Boraginaceae—9(2):311

Botrychium michiganense—9(2):300

Breeding System (Moraceae)—9(2):383

Buddleja davidii—9(1):213

Cactaceae—9(1 ):25; 9(2):351

Caesalpinioideae—9(1 ):129; 9(2):381

Calathea galdamesiana—9(2):319

California (U.S.A.)—9(1 ):201

Caribbean—9(1 ):129

Carolina Piedmont (U.S.A.)—9(1 ):31

Casearia draganae—9(2):325

Central America—9(1 ):49

Chazaliella—9(1):75

Chionolaena—9(1 ):63

Chytropsis—9(1):75

Climbing plants—9(1 ):149

Coastal Plain (U.S.A.)—9(1 ):31

Coastal prairie (Texas, U.S.A.)—9(2):485

Colombia—9(2):325

Commelinaceae—9(2):353

Community ecology—9(1 ):235

Comopellis —9(2):363

presbya —9(2):364

Cordia boissieri—9(1 ):173

Coryphantha—9(1 ):25

orcuttii—9(1 ):25

sneedii—9(1 ):25

sneedii var. orcuttii —9(1 ):25,

28[invalid]

sneedii var. orcuttii—9(2):351

Costa Rica—9(1 ):149

506

Journal of the Botanical Research Institute of Texas 9(2)

Crassulaceae—9(1 ):1

Cretaceous—9(1 ):135

Cuba—9(1 ):31

Cuscatlania vulcanicola—9(1 ):167

Cyperaceae—9(1 ):31,251; 9(2):477

Cyperus

granitophilus—9(1 ):251

squarrosus—9(1 ):251

Dalbergiae—9(1 ):49

Dasylarynx—9(1 ):122

anomalus—9(1 ):121,122

Dean, Blanche Evans—9(2):418

Diodella—9(1 ):103

Dominican amber—9(1 ):121,129

Dominican Republic—9(1 ):121; 9(2):449

Durango (Mexico)—9(2):453

Dyschoriste ecuadoriana—9(1 ):83

Ecuador—9(1 ):81,89; 9(2):325

Edwards Plateau (U.S.A.)—9(1 ):259

El Durazno (Mexico)—9(2):453

El Salvador—9(2):471

Enchanted Rock State Natural Area

(U.S.A.)—9(1 ):251

Endemic—9(1 ):43,107, 251

Endocarps—9(1 ):137

Erigeron

geiseri—9(1 ):11

strigosus—9(1):11

Eritrichium—9(2):311

aretioides—9(2):314

chamissonis—9(2):314

Escaped garden plants—9(1 ):201

Escobaria—9(1 ):25

orcutti—9(1 ):25

sneedii var. orcuttii—9(1 ):25, 28

Eumachia—9(1):75

abrupta—9(1 ):76

acuifolia—9(1 ):76

albert-smithii—9(1 ):76

astrellantha—9(1 ):76

boliviana—9(1 ):76

cephalantha—9(1 ):77

deinocalyx—9(1 ):77

guianensis—9(1 ):77

kappleri—9(1 ):77

membranacea—9(1 ):76

microdon—9(1 ):77

nana—9(1 ):78

pallidinervia—9(1 ):78

paupertina—9(1 ):78

podocephala—9(1 ):78

wilhelminensis—9(1 ):79

Fabaceae—9(1 ):129,173; 9(2):334, 381,

471,475

Flora of Alabama—9(2):317

Floristics (Missouri)—9(1 ):235

Follets Island (Texas)—9(2):485

Forsythia—9(1 ):195

suspensa—9(1 ):195

viridissima—9(1 ):195

Fossil—9(1 ):121,129,137

Freeman, John D.—9(2):420

Freziera—9(1 ):89

humiriifolia—9(1 ):90

neillii —9(1 ):92

tundaymensis —9(1 ):95

yanachagensis —9(1 ):96

Fungus (gasteroid)—9(1 ):135

Furrowed blister pods—9(1 ):137

Gamochaeta—9(1 ):63

Gesneriaceae—9(2):345

Gnaphalieae—9(1 ):63

Gnaphaliinae—9(1 ):63

Gnaphaliothamnus nesomii —9(1 ):63, 64

Gnaphalium—9(1 ):63

Granitic outcrops—9(1 ):251

Guatemala—9(1 ):63

Guayana Shield—9(1):75

Guiana Shield—9(1):75

Gulf of Mexico—9(1 ):137

Hancock Biological Station (Kentucky,

U.S.A.)—9(1 ):229

Harper, Roland McMillan—9(2):411

Hexasepalum—9(1 ):103

angustatum— 9(1 ):104

apiculatum— 9(1 ):104

gardneri— 9(1 ):104

lippioides— 9(1 ):104

mello-barretoi— 9(1 ):104

radulum— 9(1 ):105

rosmarinifolium— 9(1 ):105

sarmentosum— 9(1 ):105

scandens— 9(1 ):105

serrulatum— 9(1 ):105

Hexastylis harperi— 9(2):317

Hispaniola— 9(1 ):129

History of Botany (Alabama)—9(2):397

Honduras— 9(1 ):1 67

Humiriaceae— 9(1 ):1 37

Hymenaea protera— 9(1 ):129

Invasives— 9(1 ):21 3, 229

Irenodendron— 9(2):332

coriaceum— 9(2):333

cupulatum— 9(2):332

ovalifolium— 9(2):333

Isthmus of Tehuantepec

(Mexico)— 9(1 ):43

IUCN Red List— 9(1 ):43

Kentucky (U.S.A.)— 9(1 ):229

Klaprothiopsis— 9(2):370

dyscrita— 9(2):336

Kowal, Robert R."Bob" (In Memoriam)—

9(2):493

Krai, Robert—9(2):397,422

Leguminosae—9(1 ):49,107

Lima bean—9(1 ):107

Llano Uplift (U.S.A.)—9(1 ):251,259

Loasaceae—9(2):369

Locomapa—9(1 ):167

Los Cusingos (Costa Rica)—9(1 ):149

Louisiana (new state record

U.S.A.)—9(2):475

Lucilia-group

(Gnaphaliothamnus)—9(1 ):63

Lyell, Charles—9(2):403

Macherium—9(1 ):49

excavatum —9(1 ):51

franksullyvanii —9(1 ):58

paucifoliolatum —9(1 ):54

ramosiae —9(1 ):49

rubrinervum —9(1 ):56

Marantaceae—9(2):319

Margaritopsis—9(1 ):75

Matelea chihuahuensis —9(1 ):187

Melastomataceae—9(2):449

Mendoncia

hollenbergiae —9(1 ):81

sericea— 9(1 ):83

Mexico—9(1 ):31,43,49; 9(2):453

Miconia alainii—9(2):449

Miconieae—9(2):449

Micranthes

gaspensis—9(1 ):7

nivalis—9(1 ):7

rufopilosa —9(1 ):7

tenuis—9(1 ):7

Mid-Tertiary Dominican amber—9(2):353,

361,369, 381

Mid-Tertiary—9(1 ):121,129

Milkweed—9(1):187

Missouri (U.S.A.)—9(1 ):235

Mohr, Charles T.—9(2):407

Monocotyledon—9(1 ):121

Monterrey Metropolitan Area, Mexico—

9(1 ):173

Moonwort—9(2):295

Moraceae—9(2):383

Morus—9(2):383

Mulberries—9(2):383

Murray State University—9(1 ):229

Myanmar—9(1 ):135

MycoBank number—9(1 ):135

Naturalized (Buddleja davidii)—9(1 ):213

Nautilocalyx rugosus—9(2):345

Neotropics—9(1 ):5

Neotype (Sedum rubroglaucum)—9(1 ):1

Nephrolepidaceae—9(1 ):201

Nephrolepis cordifolia —9(1 ):201

Nevius, Reuben D.—9(2):405

New Mexico (U.S.A.)—9(1 ):25,187

North American Flora—9(1 ):187

North Sea—9(1):137

Norway—9(2):433

Noteworthy collection—9(1 ):187

nrDNA—9(1 ):11

Nuevo Celilac—9(1 ):167

Nuevo Leon (Mexico)—9(1 ):173

Nutsedge—9(1 ):31

Nyctaginaceae—9(1 ):167

Oaxaca (Mexico)—9(1 ):43

Ohio (U.S.A.)—9(2):477

Oklahoma (U.S.A.)—9(1 ):11,251

Oleaceae—9(1):195

Ophioglossaceae—9(2):295

Orange-eye butterfly bush—9(1 ):213

Ozark National Scenic

Riverways—9(1 ):235

Palaeogaster micromorpha —9(1 ):135

Pamela Lankester Garden—9(1 ):149

Panama—9(2):319

Paniculati (sect.)—9(1 ):107

Papilionoideae—9(1 ):49,107

Index

507

Pentaphylacaceae—9(1 ):89

Peru—9(1 ):89;9(2):345

Peters, Thomas Minott—9(2):410

Phaseoleae—9(1 ):107

Phaseolus lignosus— 9(1 ):107

Phoradendron leucarpum ssp.

leucarpum—9(1 ):229

Phylogenetic affinities—9(1 ):11

Phytogeographical relationships—

9(1 ):259

Piedmont of Alabama—9(1 ):251

Piedmont scrub—9(1 ):173

Pollen—9(1):11

Prioria

copaifera—9(1 ):129

dominicana —9(1 ):129,130

joveri—9(1 ):129

mannii—9(1 ):129

Pseudhaplocricus— 9(2):354

hexandrus— 9(2):354

Pseudognaphalium—9(1 ):3

paramorum— 9(1 ):63, 72

stolonatum— 9(1 ):63, 69

Readea—9(1 ):5

Refugio de Aves (Dr. Alexander Skutch)

—9(1):149

Rhamnaceae—9(2):361

Rhodotypos scandens—9(1 ):195

Rio Cenepa Watershed—9(2):345

Rio General (Costa Rica)—9(1 ):149

Rubiaceae—9(1):75,103

Sacoglottis—9(1 ):137

amazonica—9(1 ):137

costata—9(1 ):137

costata—9(1):137

Samydaceae—9(2):325, 331

San Isidro de El General—9(1 ):149

Saxifragaceae—9(1 ):7

Schistostemon—9(1 ):137

Schoenoplectiella

hallii—9(2):477

saximontana—9(2):477

Scleria bellii —9(1 ):31

Scrophulariaceae—9(1 ):213

Sea-bean—9(1 ):137

Sedum rubroglaucum —9(1 ):1, 4

Selva Zoque (Mexico)—9(1 ):43

Senegalia

duartei—9(2):336

noblickii—9(2):338

phillippei—9(2):340

Sex ratio variation in mulberries—9(2):383

Shrublands (Monterrey, Mexico)—

9(1 ):173

Sierra de los Cuchumatanes—9(1 ):3

South America—9(2):331,334

Spermacoceae—9(1 ):103

Springs (Missouri)—9(1 ):235

Stenostephanus holm-nielsenii —9(1 ):86

Styracaceae—9(1 ):43

Styrax

conterminous—9(1 ):43

peltatus —9(1 ):43

Tamaulipan thornscrub—9(1 ):173

Tennessee (U.S.A.)—9(1 ):213

Texas (U.S.A.)—9(1 ):11,251,259; 9(2):485

Thomas, Joab L.—9(2):419

Trachelospermum jasminoides—9(1 ):195

Tropical Science Center—9(1 ):149

Tutwiler, Julia S.—9(2):416

Type designations—9(1 ):75

Type specimen—9(1 ):107

U.S.A.—9(1 ):31,187,195, 201,213, 229,

235, 251,259

Ulmus alata—9(1 ):229

United States (southeastern)—9(2):317

Uxpanapa-Chimalapa region (Mexico)—

9(1 ):43

Wichita Mountains (U.S.A.)—9(1 ):251

Wild Bermuda bean—9(1 ):107

Wolf, Wolfgang—9(2):414

Woody associates—9(1 ):213

Woody plants—9(1 ):149

Yoro—9(1 ):167

Yosemite National Park (U.S.A.)—9(1 ):1

Yukon (Canada)—9(1 ):7

69 New Names and New Combinations:

Volume 9 (2015)

Botrychium michiganense W.H. Wagner ex A.V. Gilman, Farrar, &

Zika, sp. nov.—9(2):300

Calathea galdamesiana H. Kenn. & R. Flores, sp. nov.—9(2):319

Casearia draganae M.H. Alford, sp. nov.—9(2):325

Comopellis K.L. Chambers & Poinar, gen. nov.—9(2):363

presbya K.L. Chambers & Poinar, sp. nov.—9(2):364

Coryphantha sneedii var. orcuttii (Boed.) Gorelick, comb, et stat.

nov.—9(2):351

Coryphantha sneedii var. orcuttii (Boed.) Gorelick, comb, et stat.

nov.—9(1 ):28 [invalid]

Dasylarynx Poinar & K.L. Chambers, gen. nov.—9(1 ):1 22

anomalus Poinar & K.L. Chambers, sp. nov.—9(1 ):1 22

Dyschoriste ecuadoriana Wassh., sp. nov.— 9(1 ):83

Escobaria sneedii var. orcuttii (Boed.) Gorelick, comb, et stat.

nov.— 9(1):28

Eumachia

abrupta (Hiern) Delprete & J.H. Kirkbr., comb. nov.— 9(1 ):76

acuifolia (C. Wright) Delprete & J.H. Kirkbr., comb. nov.— 9(1 ):76

albert-smithii (Standi.) Delprete & J.H. Kirkbr., comb. nov.

—9(1):76

astrellantha (Wernham) Delprete & J.H. Kirkbr., comb. nov.

—9(1):76

boliviana (Standi.) Delprete & J.H. Kirkbr., comb. nov.—9(1):76

cephalantha (Mull. Arg.) Delprete & J.H. Kirkbr., comb. nov.

—9(1 ):77

deinocalyx (Sandwith) Delprete & J.H. Kirkbr., comb. nov.

—9(1 ):77

guianensis (Bremek.) Delprete & J.H. Kirkbr., comb. nov.

—9(1 ):77

kappleri (Miq.) Delprete & J.H. Kirkbr., comb. nov.—9(1 ):77

membranacea (Gillespie) Delprete & J.H. Kirkbr., comb. nov.

—9(1):76

microdon (DC.) Delprete & J.H. Kirkbr., comb. nov.—9(1 ):77

nana (K. Krause) Delprete & J.H. Kirkbr., comb. nov.—9(1 ):78

pallidinervia (Steyerm.) Delprete & J.H. Kirkbr., comb. nov.

—9(1):78

paupertina (Standi.&Steyerm.) Delprete&J.H. Kirkbr.,comb. nov.

—9(1):78

podocephala (Mull.Arg.) Delprete & J.H. Kirkbr., comb. nov.

—9(1):78

wilhelminensis (Steyerm.) Delprete & J.H. Kirkbr., comb. nov.

—9(1):79

Freziera

humiriifolia D. Santam., sp. nov.—9(1):90

neillii D. Santam., sp. nov.—9(1 ):92

tundaymensis D. Santam., sp. nov.—9(1 ):95

yanachagensis D. Santam., sp. nov.—9(1):96

Gnaphaliothamnus nesomii M.O. Dillon & Luebert, sp. nov.—

9(1):64

Hexasepalum

angustatum (Steyerm.) J.H. Kirkbr. & Delprete, comb. nov.

—9(1):104

apiculatum (Willcl.) Delprete&J.H. Kirkbr., comb. nov.—9(1):104

gardneri (K. Schum.) J.H. Kirkbr. & Delprete, comb. nov.—

9(1 ):104

lippioides (Griseb.) J.H. Kirkbr. & Delprete, comb. nov.—9(1 ):104

mello-barretoi (Standi.) J.H. Kirkbr. & Delprete, comb,

nov.—9(1):104

508

radulum (Willd.) Delprete & J.H. Kirkbr., comb, nov.—9(1 ):105

rosmarinifolium (Pohl ex DC.) Delprete & J.H. Kirkbr., comb,

nov.—9(1):105

sarmentosum (Sw.) Delprete & J.H. Kirkbr., comb. nov.— 9(1 ):1 05

scandens (Sw.) J.H. Kirkbr. & Delprete, comb. nov.— 9(1 ):1 05

serrulatum (P. Beauv.) J.H. Kirkbr. & Delprete, comb. nov.—

9(1 ):1 05

Hexastylis harperi (Gaddy) B.R. Keener & LJ. Davenp., comb, et

stat. nov.— 9(2):317

Irenodendron M.H. Alford & Dement, gen. nov.—9(2):332

coriaceum (Spruce ex Benth.) M.H. Alford & Dement, comb,

nov.— 9(2):333

cupulatum (Spruce ex Benth.) M.H. Alford & Dement, comb,

nov.— 9(2):332

ovalifolium (Macbride) M.H. Alford & Dement, comb. nov.—

9(2):333

Klaprothiopsis Poinar, Weigend, & Henning, gen. nov.—9(2):370

dyscrita Poinar, Weigend, & Henning, sp. nov.—9(2):336

Machaerium

excavatum J. Linares, sp. nov.—9(1 ):51

franksullyvanii J. Linares, sp. nov.—9(1 ):58

paucifoliolatum J. Linares, sp. nov.—9(1 ):54

ramosiae J. Linares, sp. nov.—9(1):49

rubrinervum J. Linares, sp. nov.—9(1 ):56

Journal of the Botanical Research Institute of Texas 9(2)

Mendoncia

hollenbergiae Wassh., sp. nov.—9(1 ):81

sericea Wassh., sp. nov.—9(1 ):83

Micranthes rufopilosa (Hulten) D.F. Murray & Elven, comb. nov.

—9(1):7

Nautilocalyx rugosus R. Rojas & J.L. Clark, sp. nov.—9(2):345

Prioria dominicana Poinar & K.L. Chambers, sp. nov.—9(1 ):130

Pseudhaplocricus Poinar & K.L. Chambers, gen. nov.—9(2):354

hexandrus Poinar & K.L. Chambers, sp. nov.—9(2):354

Pseudognaphalium

paramorum (S.F. Blake) M.O. Dillon, comb. nov.—9(1):72

stolonatum (S.F. Blake) M.O. Dillon, comb. nov.—9(1):69

Scleria bellii LeBlond, sp. nov.—9(1 ):31

Senegalia

duartei Seigler & Ebinger, sp. nov.—9(2):336

noblickii Seigler & Ebinger, sp. nov.—9(2):338

phillippei Seigler & Ebinger, sp. nov.—9(2):340

Stenostephanus holm-nielsenii Wassh., sp. nov.—9(1):86

Styrax peltatus P.W. Fritsch, sp. nov.—9(1 ):43