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PHYTOLOGIA

An international journal to expedite plant systematic, phytogeographical and
ecological publication

Vol. 86 February 2004 No. 1

B. L. TURNER AND J. K. WILLIAMS A letter from the new owner and the new
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E. L. KEITH, Dimorphotheca sinuata and Zinnia violacea (Asteraceae), two
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PHYTOLOGIA

(ISSN 00319430)

Phytologia, a journal of plant systematics, phytogeography and
vegetation ecology, is published three times a year by the Warner
Herbarium, a research unit of the Department of Biological Sciences,
Sam Houston State University.

Editor Justin K. Williams
e-mail: bio_jkw @shsu.edu
PHONE: (936) 294-1552

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Copyright 2004 by the Warner Herbarium, Sam Houston
State University. Printed by the Sam Houston Press.

Phytologia (Feb 2004) 86(1)

A LETTER FROM THE NEW OWNER AND THE NEW
EDITOR OF PHYTOLOGIA

PHYTOLOGIA was established in 1933 by the late Henry A.
Gleason (1882-1975) and Harold N. Moldenke (1909-1996). In their
forward to the first issue the two entrepreneurs noted that their journal
was “designed to expedite botanical publication.” They further stated
“In these days of depression, the announcement of a new botanical
magazine is made with trepidation and will doubtless be received with
suspicion.”

To this end the journal fared well maintaining a steady stream of
rapid publications. With Gleason’s retirement as one of the editors in
1951, H. A. Moldenke and his wife Alma continued the enterprise, this
with much effort and attrition of their time: they truly felt that they
were serving the greater botanical community. For many years no
financial gain accrued to the managers and publication costs per page
were remarkably low, an original mission of the founding managers.

The Moldenkes continued as managers through the year 1988,
whereupon they sold PHYTOLOGIA to Billie L. Turner, with the
understanding that he would see to it that the journal continued to serve
the botanical community as intended by its founders.

Almost immediately after acquiring the Journal, one of B. L.
Turner’s academic children, Michael J. Warnock, at the time a member
of the faculty at Sam Houston State University, Huntsville, Texas,
offered to take over stewardship of PHYTOLOGIA with the
understanding that he would carry out the responsibilities entrusted to
Turner by the Moldenkes. This he agreed to do, and the journal was
sold to Warnock.

So it was that Warnock took over stewardship of PHYTOLOGIA in
November of 1988, continuing this through the year 2003 (editing and
publishing volumes 66-85). Unfortunately, newly assumed
administrative duties as Director of Research at the University of
Missouri, Columbia, precluded any sustained stewardship of the

MISSOURI BOTANICAL

MAR 2 3 2004
GARDEN LIBRARY

Phytologia (Feb 2004) 86(1)

Journal, and after a lapse of several years he agreed to resell
PHYTOLOGIA back to the B. L. Turner.

In an effort to reestablish and maintain a tradition instituted by the
two visionary and founding botanists, H. A. Gleason and H. N.
Moldenke, we B. L. Turner and J. K. Williams present volume 86 of
PHYTOLOGIA*.

i a
pL l i I
B. L. Turner J. K. Williams
Prof. Emeritus Assistant Professor
University of Texas, Austin Sam Houston State University
Owner of PHYTOLOGIA PHYTOLOGIA, Editor

photo by T. L. Wendt

*The cover date for the previously published volume and issue of
PHYTOLOGIA, Vol. 85(6), was December 1998, although the official
publication date is August 2003 (PHYTOLOGIA 85: 511). In an effort
to eliminate any future potential nomenclatural complications resulting
from a question or confusion in priority, the cover date for Volume
86(1), is February 2004, consistent with the publication date. This will
result in a 5 year publication gap in the serial publication of
PHYTOLOGIA.

Phytologia (Feb 2004) 86(1) ]

FLORISTICS OF BAYGALLS IN CENTRAL LOUISIANA
Barbara R. MacRoberts and Michael H. MacRoberts

Bog Research, 740 Columbia, Shreveport, LA 71104 and Herbarium
Museum of Life Sciences, Louisiana State University in Shreveport
Shreveport, LA 71115

Lynn Stacey Jackson
U.S.D.A. Forest Service, Kisatchie National Forest, Winn Ranger
District, Winnfield, LA 71483

ABSTRACT

Headwater plant communities --- baygalls and pitcher plant bogs
--- were Studied in the northern part of the Winn Ranger District,
Kisatchie National Forest, Louisiana. Floristically baygalls, discussed
in this paper, form a fairly uniform community type across most of the
West Gulf Coastal Plain.

KEY WORDS: baygall, West Gulf Coastal Plain, floristics, Kisatchie
National Forest, Louisiana.

INTRODUCTION

Baygalls are a characteristic wetland community of the Atlantic
and Gulf Coastal Plains extending through western Louisiana into
eastern Texas (Allen et al. 1988; Bridges and Orzell 1989; Brooks et al.
1993; Christenson 1988; Harcombe et al. 1993; Nesom et al. 1997;
Weakley et al. 1998). They are variable over their range but are
generally characterized by such woody species as Magnolia virginiana
(sweet bay) and Ilex coriacea (gallberry holly), for which the
community is named. Other common associates are Acer rubrum, Ilex
opaca, Nyssa biflora, Persea palustris, Rhododendron spp., and

2 Phytologia (Feb 2004) 86(/)

Vaccinium spp. In western Louisiana and eastern Texas, baygalls are
generally headwater communities occurring at streamheads or on short
slopes that receive enough seepage to be permanently saturated. They
are recognized as a natural community in both Louisiana and Texas
(Diamond et al. 1987; Louisiana Natural Heritage Program 1994; Texas
Natural Heritage Program 1993; Turner et al. 1999; Weakley et al.
1998; Van Kley 1999a,b) and have been described for various localities
in the West Gulf Coastal Plain (Ajilvsgi 1979; Brooks et al. 1993;
Marks and Harcombe 1981; Martin and Smith 1991; Nesom et al.
1997; Nixon et al. 1980, 1983; Nixon and Ward 1988; Orzell 1990;
Van Kley 1999a,b; Watson 1979).

As with most plant communities, descriptions usually center on
woody species because they can be identified year round and are
relatively easily quantified. The herbaceous layer is often briefly
mentioned but seldom described in detail. In this study we describe
both.

SITE DESCRIPTION

This study was conducted on the northern part of the Winn Ranger
District, Kisatchie National Forest, in northern Natchitoches and Winn
parishes, Louisiana. All study sites are located within what is generally
classified as riparian forest habitat that is adjacent to, and often
surrounded by, upland longleaf pine forest (Martin and Smith 1991).
The area was originally dominated by longleaf pine (Eldredge 1934;
Martin and Smith 1991), notably in forest/savanna form on uplands, but
today in some areas the longleaf pine has been replaced with off-site
species such as loblolly pine. The longleaf pine forest community type
has received detailed ecological and distributional attention in the West
Gulf Coastal Plain (Bridges and Orzell 1989; Evans 1997; Frost 1993;
Harcombe et al. 1993; Van Kley 1999a,b).

The area has a subtropical humid climate with hot summers and
mild winters. The average annual precipitation is about 140 cm spread
evenly throughout the year. The average annual temperature is about

Phytologia (Feb 2004) 86(1) 3

20C (extreme range is about -7C to 38C). Thunderstorms average about
60 days per year and are concentrated in the growing season (Olson and
Platt 1995). Fire was an important element in the evolution of the plants
and animals of the area, many communities being dependent on
growing season burns ignited by lightning strikes (Bridges and Orzell
1989; Drewa et al. 2002; Frost 1998; Harcombe et al. 1993; Olson and
Platt 1995; Platt 1999).

Baygalls are situated on loamy fine sand, fine sandy loam, or fine
loamy sand (Malbis, Betis, Ruston, Sacul). All are below uplands
(ridgetops) of similar soils but with gentler gradients (1-5 percent as
opposed to 5-12 percent). All are Paleudults and Hapludults (suborder
= Udults, Order = Ultisols) (Martin et al. 1990). There is little
topographic relief except for slightly rolling hills. The study area is on
the older geologic surfaces, notably the Sparta formation of Tertiary
age (Groat and Roland 1984).

The study area in general consists of two north-south ridges
divided by Saline Bayou. The watershed of the western area flows into
Black Lake Bayou to the west and the Saline Bayou to the east. The
watershed of the eastern ridge flows into Saline Bayou to the west and
Dugdemona River to the east. All ultimately discharge into the Red
River.

METHODS

1. We made monthly visits to each of six baygalls (numbers 5
through 10) between March and November 1999 to obtain complete
floristic lists for each site. The study sites are listed in Appendix | by
name and number. All sites were between 0.1 and 0.2 ha in size.
Voucher collections for many of the species are deposited at BRIT and
LSU. Plant nomenclature follows Kartesz and Meacham (1999);
scientific authorities for all names used in this paper can be read from
that reference.

4 Phytologia (Feb 2004) 86(1)

Species Number of Mean Range (cm)
Individuals Diameter (cm)
Magnolia virgin. 32 10.5 2-30
Nyssa biflora 30 8.0 2-40
Persea palustris 29 2 Be 2-10
Viburnum nudum 9 2.0 2
Acer rubrum vi 6.1 2-10
Ilex opaca 3) 2.4 2-3
Chionanthus virg. 4 4.0 2-10
Vaccinium sp. 4 2 2-3
Liquidambar styra. 4 4.0 2-6
Toxicodendron v. 3 2.7 2-4
Taxodium distich. 3 pSr7 12-17
Pinus taeda l 12.0 12
Alnus serrulata ! 2.0 2
Quercus alba | 18.0 18

Table 1. Dominant woody species.

2. In May and June 1999 we established ten, 5 X 5 meter quadrats
(25 m sq.), one in each of ten baygalls; we added four sites to the
original six to increase our sample size. Frequency and diameter of all
woody species with a dbh of 2 cm or greater were recorded.
Herbaceous species presence was described by identifying all species in
the plots; herbaceous cover was measured and percent cover estimated
for major species. Canopy cover was estimated for the quadrat.

3. A soil sample was collected from the upper 15 cm of each of the
ten 5 X 5 meter plots and analyzed by A & L Laboratories, Memphis,
Tennessee.

RESULTS

Appendix 2 summarizes the total species composition of six

Phytologia (Feb 2004) 86(1) 5

Species

Athyrium filix-fem.
Bignonia capreolata
Carex atlantica

Carex debilis

Carex leptalea 21
Chasmanthum laxum
Dichanthelium dich.
Dichanthelium sp.
Eleocharis tortilis
Gelsemium semper.
Liverworts 2
Lycopus rubellus
Mitchella repens 2
Osmunda cinnam.
Osmunda regalis 1
Platanthera clav.
Rudbeckia scabrifolia
Scleria sp.

Scleria triglomerata
Solidago patula
Sphagnum sp.

Viola primulifolia Z
Woodwardia areolata 22
Woodwardia virginica
Xyris laxifolia

Percent by plot

19

oes" 10

m= N
i)

—

—
Ww
i=)
-
No

10
1
30 20 88 50 50 30 60
By LO
1

Total 30 "5. 52 .69° 98 82 70° 80.8! 54
Table 2. Percent herbaceous ground cover by plot.

baygalls. Appendix 3 shows occurrence by plot for each species in the
5 X 5 mplots. Table | summarizes the larger woody vegetation in ten 5
X 5 m plots. Table 2 gives the percent of herbaceous ground cover by
dominant species in each plot. Only species accounting for 1% or more

6 Phytologia (Feb 2004) 86(1)

in any plot are included. Table 3 gives percentage frequency of all
species from the baygalls as rated in Reed’s (1988 and update)
hydrophytic classification. It also gives the prevalence index for the
entire sample (see Cowardin et al. 1979; Federal Manual 1989;
MacRoberts and MacRoberts 2001; Reed 1988; U.S. Corps of
Engineers 1987 for an explanation of terms and measures). Table 4
gives information on soil samples from the ten sample sites.

DISCUSSION

There were 40 families, 61 genera, and 89 species in the six
baygalls. The average baygall had 52 species (range 39 to 68). Dicots
accounted for 54 percent of the total species. All sites had liverworts
and Sphagnum, which are not included in the totals. Sorenson’s Index
of Similarity among the six baygalls ranges from 62 to 87, indicating
that they are all floristically the same community.

Magnolia virginiana, Nyssa biflora, and Persea palustris were
clearly the dominant woody species, both in number of individuals and
in basal area within the 5 X 5 m plots. Several species, for example,
Smilax laurifolia, were present in many plots, but since the stems were
always less than 2 cm diameter, they are not listed among those in
Table 1. Canopy cover of the ten plots ranged from 80% to 100% and
averaged 94%.

Overall herbaceous cover ranged from 5% to 98% and averaged
65%. Ferns and monocots dominated. The following species are listed
in order of dominance in the herbaceous layer: Woodwardia areolata
(35.1%), Carex leptalea (5.6%), Mitchella repens (4.1%), Carex debilis
(3.7%), Osmunda cinnamomea (2.9%), Woodwardia virginica (2.3%),
Carex atlantica (1.9%), Eleocharis tortilis (1.9%), liverworts (1.1%),
Osmunda regalis (1.0%), Sphagnum sp. (1.0%). All the other species in
these plots accounted for less than 1% each of the total herbaceous
layer.

Phytologia (Feb 2004) 86(1) F

OBL FACW FAC FACU- UPL Prev. Sample

Index Size
All species oy 6 39 24 #10 0 27 89
Woody species 9 38 35 18 0 2.62') 34
Herb. species 38 =6.40 16 6 0 1.89 55

Table 3. Species percents by wetland categories, prevalence index,
and species sample size (see text for category explanation).

The flora of these baygall sites is clearly dominated by
hydrophytes. About 80% are OBL, FACW, and FAC (50% or higher
qualifies a site as wetland). Counting all species for these sites, the
prevalence index is 2.17, meaning that baygalls are wetlands (1.0
means that all species are obligate wetland species, 5.0 means that all
species are obligate upland [dry] species). Woody and non-woody
species differed in prevalence: the prevalence index for woody species
was 2.62 and that for herbaceous species was 1.89. This suggests either
that woody vegetation has a greater moisture gradient than non-woody
species or that the species are somewhat misclassified.

The soils are acidic and low in nutrients and organic matter, falling
within the normal range of soil chemistry of baygalls and pitcher plant
bogs in east Texas and west Louisiana except that the pH for the
present group of baygalls is slightly lower (MacRobderts and
MacRoberts 2001; MacRoberts et al. 2002; Nesom et al. 1997; Nixon
and Ward 1986).

The West Gulf Coastal Plain is ecologically part of the eastern
flora. Using Kartesz and Meacham (1999), we determined the
distribution of the species that occur in our study sites. We found 95%
to be eastern, 3% cosmopolitan (found throughout North America), and
2% endemic. The endemics are Rudbeckia_ scabrifolia and
Rhododendron oblongifolium. The cosmopolitan species are Carex
leptalea, Juncus effusus, and Athyrium filix-femina.

8 Phytologia (Feb 2004) 86(1)

Site Exchangeable ions (ppm)
pH P K Ca Mg OM%

i BSeP SLT S2 “322°. 46 1.9
2 43 10“ Si "458" 69 i)
3 44 10 28 196 52 aa
+ Be 5 BB 62. 3B oe 4h 8 48
5 AY S45. 304147) “36. ° 9 19
6 A like OC 225 Gee 4522
fi ALG! oA StH 99 ISG SG eics
8 Ae AG 1031S) 23
9 4.3), 28) 1 259 "86,028" © 256

10 AOR 1133) 2135-569),".44
Table 4. Soil chemistry.

Baygalls in the West Gulf Coastal Plain occur in association with
pyrogenic communities (Bridges and Orzell 1989; Harcombe et al.
1993). But unlike these communities, baygalls are not pyrogenic. We
have witnessed in many instances that prescribed fire, both in growing
and non-growing season, does not penetrate far, if at all, into baygall
communities but at most fingers into the edges. These fires, which
historically probably occurred once every few years and which
probably occurred in the growing season and were caused by lightning,
cleared the surrounding lands but usually left these baygall “islands”
intact.

Comparing baygalls with other plant communities in the West Gulf
Coastal Plain indicates that, while distinct, they are most closely related
to pitcher plant bogs as indicated by ordination (Van Kley 1999a) and
direct species comparisons (MacRoberts and MacRoberts 2001; Nixon
and Ward 1986). For example, comparing the total floristic list for six
baygalls to four small pitcher plant bogs on the Kisatchie District of the
Kisatchie National Forest with comparable species numbers gives an
Index of Similarity (Sorenson’s) in the mid-30s (MacRoberts and

Phytologia (Feb 2004) 86(1) 9

MacRoberts 2001). Thus, while sharing many species, baygalls and
bogs do not share enough to be considered the same plant community.

Martin and Smith (1991) have distinguished between "wooded
seeps" and "bayhead swamps" but little in their description separates
these entities; these authors admit that the two communities are very
similar and intergrade. Brooks et al. (1993) found that wooded seeps
graded into wet creek bottoms or bayhead swamps. On the basis of
quantitative data, Van Kley (1999a) and Turner et al. (1999) found that
these "different" communities were clearly one. Our work supports the
conclusion that they be considered the same.

This does not mean that baygalls are uniform. Most of our baygalls
are the same community type described by Nixon et al. (1983) as
"wet," which occurs at branch heads, creek heads, wet creek bottoms,
and seepages. However, some show similarities to bogs (Nixon and
Ward 1988). Notable is site 10, which showed bog similarities and may
have been a bog at one time.

While we found Nyssa biflora and Viburnum nudum important,
Nixon et al. (1980) and Nixon and Ward (1988) in their study of wet
creek bottoms in Nacogdoches County and in the Trout Creek drains on
the Angelina National Forest in Texas did not. As they point out, at
least in their Trout Creek location, the herbaceous species are a mixture
of baygall and seepage bog species, which may partly explain the
difference (Nixon and Ward 1986). In the Trout Creek area, baygalls
and bogs are often contiguous and the species from one grade into the
other in ecotonal zones.

Brooks et al. (1993) studied baygalls on a north-south gradient in
east Texas from southern Rusk to central Hardin counties. They found
that, while Nyssa biflora and Magnolia virginiana were important in
both northern and southern examples, Acer rubrum, Liquidambar
styraciflua, and Morella carolinensis were important in the north and
Cyrilla racemiflora, Ilex coriacea, and Quercus laurifolia were
important in the south. They also found that Persea palustris (which

10 Phytologia (Feb 2004) 86(1)

was important in the Winn District baygalls) was not paticularly
important in either group.

Nixon et al. (1983) in their study of different segments of a single
streamside in San Augustine County found that different areas of the
stream had different species composition, depending on moisture.

Nesom et al. (1997) found that in Walker and Anderson counties
on the western edge of the baygall community range, such trees as
Magnolia and Persea have entirely dropped out of the flora and such
species as Viburnum nudum dominate.

ACKNOWLEDGMENTS

This work was supported by Chief's Grant 08-99-06-CCS-003,
National Forest Foundation 08-99-06-CCS-007, and National Fish and
Wildlife Foundation, Headwater Community Plant Restoration 99-100-
011. Thanks to Guy Nesom and Jim Neal for reviewing an earlier
version of the paper.

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Forestry, Stephen F. Austin State Univ., Nacogdoches, Texas.

Nixon, E.S. and J.R. Ward. 1988. Vegetation of a wet creek bottom site
in eastern Texas. Texas J. Sci. 40:358-361.

Olson, M.S. and W.J. Platt. 1995. Effects of habitat and growing season

fires on resprouting of shrubs in longleaf pine savannas. Vegetation
119:101-118.

Orzell, S.L. 1990. Texas Natural Heritage Program inventory of
National Forests and Grasslands in Texas. Unpublished report. Texas

Natural Heritage Program, Austin, Texas.

Platt, W.J. 1999. Southeastern pine savannas. Pp. 23-51. In R.C.

14 Phytologia (Feb 2004) 86(1)

Anderson, J.S. Frailsh, and J.M. Baskin (Eds.), Savannas, barrens,
and rock outcrop communities of North America. Cambridge
University Press, New York.

Reed, P.B. 1988 (and 1996 update). National list of plant species that
occur in wetlands: national summary. U.S. Fish and Wildlife
Service, Biological Report 88(24). Washington, D.C.

Texas Natural Heritage Program. 1993. Plant communities of Texas
(series level). Unpublished report. Texas Natural Heritage
Program, Austin, Texas.

Tumer, R.L., J.E. Van Kley, L.S. Smith, and R.E. Evans. 1999.
Ecological classification system for the national forests and
adjacent areas of the West Gulf Coastal Plain. The Nature
Conservancy, Nacogdoches, Texas.

U.S. Army Corps of Engineers. 1987. Corps of Engineers wetlands
delineation manual. Technical Report Y-87-1. U.S. Army Eng.
Waterway Exp. Stn., Vicksburg, Mississippi.

Van Kley, J.E. 1999a. The vegetation of the Kisatchie sandstone hills,
Louisiana. Castanea 64:64-80.

Van Kley, J.E. 1999b. The vegetation of the high terrace rolling
uplands, Louisiana. Castanea 64:3 18-336.

Watson, G. 1979. Big Thicket plant ecology. Big Thicket Museum,
Saratoga, Texas.

Weakley, A.S., K.D. Patterson, S. Landall, and M. Pyne. 1998.
International classification of ecological communities: terrestrial
vegetation of the southeastern United States. Working Draft. The
Nature Conservancy, Chapel Hill, North Carolina.

Phytologia (Feb 2004) 86(1) 3,

Appendix 1. Site numbers, locations, and common names. All sites are
on the Winn Ranger District, Kisatchie National Forest.

1) Strange Road Baygall. Natchitoches Parish, Compartment 18.
TI2ZNR6WS1/12.

2) Lewis Klein Baygall. Natchitoches Parish, Compartment 18.
TI3NR6WS35/36.

3) Thalictrum Baygall. Natchitoches Parish, Compartment 18.
TI3NR6WS25/26/35/36.

4) Road 506 Baygall. Winn Parish, Compartment 5. T1I3NRSWS9.

5) Compartment 18 Baygall. Natchitoches Parish, Compartment 18.
TI3NR6WS26.

6) Compartment 19 Baygall. Natchitoches Parish, Compartment 19.
TI3NRO6WS2S.

7) Rudbeckia Baygall. Natchitoches Parish, Compartment 1.
TI3NR7WS2.

8) Snake Baygall. Natchitoches Parish, Compartment 1. T13NR6WS7.
9) Cypress Branch Baygall. Winn Parish, Compartment 5.
TI3NRS5WS8/9.

10) Road WSA Baygall. Winn Parish, Compartment 5. T1L3NRSWS4.

16 Phytologia (Feb 2004) 86(/)

Appendix 2. Vascular plants occurring in six baygalls (x =
present, -=absent)

FAMILY/ SITE
SPECIES

ACERACEAE
Acer rubrum SEO 8G Cr KG
ANACARDIACEAE
Toxicodendron vernix Bo aC a Senex
APIACEAE
Oxypolis rigidior oe eK
Ptilimnium capillaceum - = = = =
AQUIFOLIACEAE
Tlex opaca De tier? Sel a Sie 29'S a gn
I. vomitoria 2 lig
ARACEAE
Arisaema triphyllum SS aa ex
ASTERACEAE
Doellingeria
sericocarpoides a KK
Elephantopus nudatus = SS
Eu patorium perfoliatum Se XM Xe ne
E. rotundifolium = (RP PSS) es x
Helianthus angustifolius = et a xX
Liatris pycnostachya eS SP ee EX
Rudbeckia scabrifolia = =x = = =
Solidago patula
var. strictula > >, CP, Ge, ame, <
S. rugosa var. aspera - - = = = x
Symphyotrichum
lateriflorum ac ace =? ae oc ac
BETULACEAE
Alnus serrulata 6 Se = oe oe
BIGNONIACEAE

Bignonia capreolata ay > a nec

Phytologia (Feb 2004) 86(1)

FAMILY/
SPECIES

BLECHNACEAE

Woodwardia areolata

W. virginiana
CAPRIFOLIACEAE
Viburnum nudum
CLUSIACEAE

Hypericum crux-andreae
CORNACEAE

Cornus florida

Nyssa biflora
CUPRESSACEAE

Taxodium distichum
CYPERACEAE

Carex atlantica

C. crebriflora
C. crinata

C. debilis

C. glaucescens
Cc. intumescens

Cc. leptalea

Gy elurida

Eleocharis tortilis

E. tuberculosa
Rhynchospora caduca

R. glomerata

R. gracilenta

Scleria oligantha

S. triglomerata
DRYOPTERIDACEAE
Athyrium filix-femina
ERICACEAE

Rhododendron canescens

R. oblongifolium

x mm mh mM OOM

x e Mm OM

SITE

*

*

*

10

oe

17

18 Phytologia (Feb 2004) 86(1)

FAMILY/
SPECIES

Vaccinium fuscatum

V. corymbosum
ERIOCAULACEAE
Eriocaulon decangulare
FAGACEAE

Fagus grandifolia
Quercus alba

Q. nigra

GENTIANACEAE

Bartonia paniculata
HAMAMELIDACEAE
Liquidambar styraciflua
IRIDACEAE

Sisyrinchium mucronatum
JUNCACEAE

Juncus effusus
LAMIACEAE

Lycopus rubellus
LAURACEAE

Persea palustris
LILIACEAE

Melanthium virginicum
LOGANIACEAE

Gelsemium sempervirens
MAGNOLIACEAE

Magnolia virginiana
MELASTOMATACEAE

Rhexia mariana

R. petiolate
MYRICACEAE

Morella cerifera

M. carolinensis
OLEACEAE

SITE

10

Phytologia (Feb 2004) 86(1)

FAMILY/
SPECIES

Chionanthus virginicus
ORCHIDACEAE
Calopogon tuberosus
Platanthera ciliaris
P. clavellata
Pogonia ophioglossoides
OSMUNDACEAE

Osmunda cinnamomea
O. regalis

PINACEAE

Pinus palustris

P. taeda

POACEAE

Chasmanthium laxum
Dichanthelium commutatum
D. dichotomum

var. dichotomum
Leersia virginica
Panicum verrucosum
Saccharum giganteum
ROSACEAE

Photinia pyrifolia
RUBIACEAE
Mitchella repens
SMILACACEAE

Smilax bona-nox

S. glauca

S. laurifolia

S. rotundifolia
VERBENACEAE
Callicarpa americana
VIOLACEAE

Viola primulifolia

*

*

*

SITE

Pane tees Pe

19

2() Phytologia (Feb 2004) 86(1)

FAMILY/
SPECIES

5
VITACEAE
Vitis rotundifolium x
XYRIDACEAE

Xyris ambigua -
Ke laxatolia =
Sphagnum

Liverworts

SITE

10

Phytologia (Feb 2004) 86(1)

Appendix 3. Species/plot occurrence.

Species

Vaccinium sp.
Woodwardia areolata
Magnolia virginiana
Osmunda cinnamomea
Persea palustis
Michella repens
Nyssa biflora

Carex leptalea
Liquidambar styr.
Viburnum nudum
Carex debilis

Ilex opaca

Liverworts

Morella carolinensis
Osmunda regalis
Acer rubrum
Dichanthelium sp.
Eleocharis tortilis
Lycopus rubellus
Smilax laurifolia
Rhododendron sp.
Solidago patula
Callicarpa americana
Athyrium filix-fem.
Chasmanthium laxum
Chionanthus virgin.
Ilex vomitoria
Morella cerifera
Pinus taeda
Toxicodendron vernix

Scleria sp.

Occurrence by Plot

l 3
x * x
x PS x
x x
1 § x
x
x x x
x x
x x
» «
x x x
x
x x x
x
x
x x
x
x x

~~ ~

~

= 00

~

a i!

10

22 Phytologia (Feb 2004) 86(1)

Appendix 3. Species/plot occurrence (cont.).

Species Occurrence by Plot

Woodwardia virginica

Viola primulifolia x G

Alnus serrulata 55

Photinia pyrifolia

Bignonia capreolata

Carex atlantica x
Eupatorium rotund.

Gelsemium semper. x.
Platanthera clave.

Quercus alba 5 5
Rhynchospora caduca KG

Rudbeckia scabrif. x
Sphagnum sp. G

Taxodium distichum x
Vitis rotundifolia 5

Xyris laxifolia x

Phytologia (Feb 2004) 86(1) 253

THE STATUS OF CAREX TENAX CHAPMAN (CYPERACEAE)
IN THE WEST GULF COASTAL PLAIN

1 2
Michael H. MacRoberts , Suzanne Birmingham Walker , &
1
Barbara R. MacRoberts

1
Bog Research, 740 Columbia, Shreveport, Louisiana 71104, U.S.A. &
Herbarium, Museum of Life Sciences, Louisiana State University in
Shreveport, Shreveport, Louisiana 71115, U.S.A.

2
14671 State Highway 87 South, Shelbyville, Texas 75973, U.S.A.
ABSTRACT

Carex tenax Chapman is uncommon over much of its range.
Recent discoveries of new locations of this species in east Texas
prompts this review of its status in the West Gulf Coastal Plain, where
it is now known from six counties in east Texas and three parishes in
adjacent Louisiana. It occurs in xeric sandylands and arenic dry
uplands, and its absence in many apparently suitable sites suggests that
the species is truly rare in the West Gulf Coastal Plain.

KEY WORDS: Carex tenax Chapman, Cyperaceae, West Gulf
Coastal Plain.

Carex tenax is endemic to the Atlantic and Gulf Coastal plains,
extending from North Carolina to east Texas (Ball 2002). It is reported
as being rare in Florida, Louisiana, and North Carolina (Kartesz &
Meacham 1999). In the West Gulf Coastal Plain, it is known from
Texas (Correll & Johnston 1970; Turner et al. 2003) and Louisiana
(Thomas & Allen 1993) but has not been found in Arkansas (Hyatt
1998) or Oklahoma (Taylor & Taylor 1989). In Louisiana, it is listed as
an §2 species (Louisiana Natural Heritage Program 1999) and is
reported from Natchitoches and Vernon parishes (MacRoberts &

24 Phytologia (Feb 2004) 86(/)

MacRoberts 1995a; Thomas & Allen 1993). In Texas, it has been
reported from Hardin County only (Correll & Johnston 1970; Matos &
Rudolph 1985; Turner et al. 2003). It is not currently listed as rare in
Texas (Texas Natural Heritage Program 1995; Texas Organization of
Endangered Species 1993), although it once was (Matos & Rudolph
1985).

We recently located C. tenax in Shelby and San Augustine
counties, Texas, and at a new location in Hardin Co., Texas. In Shelby
Co., we found it at three sites within a one mile radius centering on 31°
39’ 50”N and 94° 14’ 40”W, which is about 9.5 miles SSW of Center.
The single San Augustine Co. site is about 13 miles south of Center at
31° 35’ 54” N and 94° 11’ 37” W. The new Hardin Co. site is in the
Turkey Creek Unit of the Big Thicket National Preserve on the
Sandhill Loop of the Kirby Nature Trail. These new sites ranged in
elevation from 75 to 426 feet. All are found in xeric sandylands
(grossarenic dry uplands, arenic dry uplands) (MacRoberts &
MacRoberts 1995a; Turner et al. 1999). Carex tenax was found almost
invariably near or under an oak or pine (several areas had pine
plantations) in partial shade. Our observations on the species in
Louisiana and Texas are similar: the species, while inhabiting open
xeric sandylands, apparently prefers to be in partial shade and is an
“edge” species. Plant associates include Berlandiera pumila,
Bulbostylis ciliatifolia, Carya texana, Cnidoscolus texanus, Croptilon
divaricatum, Croton glandulosus, C. michauxii, Cyperus grayioides, C.
retrofractus, Dalea villosa, Diodia teres, Eriogonum longifolium, E.
multiflorum, Froelichia floridana, Helianthus debilis, Liatris elegans,
Loeflingia squarrosa, Matelea cynanchoides, Mimosa _hystricina,
Mirabilis albida, Monarda punctata, Opuntia humifusa, Paspalum
setaceum, Penstemon murrayanus, Paronychia drummondii, Polanisia
erosa var. erosa, Quercus incana, Q. margaretta, Selaginella
arenicola ssp. riddellii, Stylisma pickeringii, Tradescantia reverchonii,
Yucca louisianensis (authorities can be read in Kartesz & Meacham
1999). The habitat in which C. tenax occurs --- xeric sandylands and
arenic dry uplands--- is found virtually throughout the West Gulf
Coastal Plains (McBryde 1933; Bridges & Orzell 1989; MacRoberts &
MacRoberts 1994, 1995b, 1995b, 1996; MacRoberts et al. 2002a,

Phytologia (Feb 2004) 86(1) 25

aes

*
t
& re,

|

Figure |. Distribution of Carex tenax in the West Gulf Coastal Plain.

2002b; Turner et al. 1999). Searches for C. tenax in northwestern
Louisiana have failed to locate a population (MacRoberts &
MacRoberts 1995b) as have searches for the species in Texas in the
Post Oak Savanna region, e.g., xeric sites in Anderson County
(MacRoberts et al. 2002a; Singhurst et al. in prep.). The absence of C.
tenax in apparently suitable sites in east Texas and west Louisiana
suggests that the species is truly rare in the West Gulf Coastal Plain.

We searched, or had curators search, ASTC, BRCH, BRIT, VDB,
TAMU, SBSC, LSU, LSUS, SFRP, and TEX. We also contacted
knowledgeable individuals who might have additional information. As
a result we located specimens of C. tenax from Bienville Parish,
Louisiana (Phil Hyatt, pers. comm.) and Newton and Tyler counties,
Texas (Edwin Bridges and Steve Orzell, pers. comm.). These
specimens also come from xeric sandyland and arenic dry upland

26 Phytologia (Feb 2004) 86(1)

habitat. Figure 1 shows the known county/parish locations for C. tenax
in the West Gulf Coastal Plain.

DOCUMENTATION: LOUISIANA: Bienville Parish: (Hyatt
8368 [SFRP] 8370 [MICH]), Natchitoches Parish: (MacRoberts &
MacRoberts 2282 [VDB], 2294 [NLU], 2334 [LSU], 2356 [LSUS]),
Vernon Parish: (Thieret 30144 [LAF], Hyatt 8224 [MICH]). TEXAS:
Hardin Co.: (Orzell & Bridges 8900 [MICH], Brown 4502 [ASTC].
Matos & Rudolph 556 [ASTC]), MacRoberts & MacRoberts 5631
[TEX], Singhurst 2494 [BAYLU]), Jasper Co.: (Singhurst & Bridges
12,419 [BAYLU]), Newton Co.: (Orzell & Bridges 6322 [to be
deposited]), San Augustine Co.: (MacRoberts, MacRoberts & Walker
4907 [TEX]), Shelby Co.: (MacRoberts, MacRoberts & Walker 4695
[VDB]), Tyler Co.: (Orzell & Bridges 9105 [MICH]).

ACKNOWLEDGMENTS
Billie Turner, Larry Brown, Stanley Jones, Phil Hyatt, Steve

Orzell, and Edwin Bridges aided with this study as did the curators and
staff of all herbaria cited.

LITERATURE CITED

Ball, P.W. 2002. Carex Linnaeus sect. Hallerianae. Pp. 487-489. In:
Flora of North America Editorial Committee (ed.). Flora of North
America. Vol. 23. Oxford University Press, New York.

Bridges, E.L. & S.L. Orzell 1989. Longleaf pine communities of the
west gulf coastal plain. Natural Areas Journal 9:246-263.

Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of
Texas. Texas Research Foundation, Renner, Texas.

Hyatt, P.E. 1998. Arkansas Carex (Cyperaceae): a briefly annotated
list. Sida 18:535-554.

Phytologia (Feb 2004) 86(1) ay

Kartesz, J.T. & C.A. Meacham. 1999. Synthesis of North American
Flora. Version 1.0. North Carolina Botanical Garden, Chapel Hill,
North Carolina.

Louisiana Natural Heritage Program. 1999. Rare plant species of
Louisiana. Unpublished report. Louisiana Department of Wildlife
and Fisheries. Baton Rouge, Louisiana.

MacRoberts, M.H. & B.R. MacRoberts. 1994. Floristics of a xeric
sandyland in western Louisiana. Phytologia 77:414-424.

MacRoberts, M.H. & B.R. MacRoberts. 1995a. Noteworthy vascular
plant collections on the Kisatchie National Forest, Louisiana.
Phytologia 78:291-313.

MacRoberts, B.R. & M.H. MacRoberts. 1995b. Floristics of xeric
sandhills in northwestern Louisiana. Phytologia 79:123-131.

MacRoberts, B.R. & M.H. MacRoberts. 1996. Floristics of xeric
sandhills in east Texas. Phytologia 80: 1-7.

MacRoberts, B.R., M.H. MacRoberts, & J.C. Cathey. 2002a. Floristics
of xeric sandylands in the Post Oak Savanna region of east Texas.
Sida 20:373-386.

MacRoberts, M.H., B.R. MacRoberts, B. Sorrie, & R. Evans. 2002b.
Endemism in the West Gulf Coastal Plain: importance of xeric
habitat. Sida 20:767-779.

McBryde, J.B. 1933. The vegetation and habitat factors of the Carrizo
Sands. Ecol. Monogr. 3:247-297.

Matos, J.A. & D.C. Rudolph. 1985. The vegetation of the Roy E.
Larsen Sandylands Sanctuary in the Big Thicket of Texas.
Castanea 50:228-249.

28 Phytologia (Feb 2004) 86(1)

Singhurst, J.R., J. Cathey, D. Prochaska, H. Haucke, & W.C. Holmes.
in prep. Vascular flora of Gus Engeling Wildlife Management
Area, Anderson County, Texas. Unpublished Report, Texas Parks
and Wildlife, Austin, Texas.

Taylor, R.J. & C.E.S. Taylor 1989. An annotated list of the ferns, fern
allies, gymnosperms and flowering plants of Oklahoma.
Southeastern Oklahoma State Univ., Durant, Oklahoma.

Texas Natural Heritage Program.1995. Special plant list. Unpublished
report. Texas Parks and Wildlife Department, Austin, Texas.

Texas Organization for Endangered Species. 1993. Endangered,
threatened and watch lists of Texas plants. Publ. 9, Austin, Texas.

Thomas, R.D. & C.M. Allen. 1993. Atlas of the vascular flora of
Louisiana. Louisiana Dept. Wildlife and Fisheries, Baton Rouge,
Louisiana.

Turner, B.L., H. Nichols, G. Denny & O. Doron. 2003. Atlas of the
vascular plants of Texas. Sida, Bot. Misc. 24:1-888.

Turmer, RL... J.E Van _Kley, -L:S.Smith, & RE.) Evanss71999.
Ecological classification system of the national forests and
adjacent areas of the West Gulf Coastal Plain. The Nature
Conservancy, Nacogdoches, Texas.

Phytologia (Feb 2004) 86(1) 29

DIMORPHOTHECA SINUATA AND ZINNIA VIOLACEA
(ASTERACEAE), TWO ESCAPED CULTIVARS, NEW TO
FLORA OF TEXAS

Eric L. Keith
Raven Environmental Services
P.O. Box 6482
Huntsville, Texas 77342, U.S.A.
eric.keith @excite.com

ABSTRACT

Two cultivated annual composites are reported as new to the
Texas flora: Dimorphotheca sinuata DC., Zinnia violacea Cav.

KEYWORDS: Dimorphotheca, Zinnia, Asteraceae, Flora, Texas.

Dimorphotheca sinuata DC., glandular cape marigold, has been
reported as escaped from cultivation in California and Oregon
(Hickman 1993, USDA 2002). Until now, the species has not been
reported for Texas (Correll & Johnston 1970, Hatch et. al. 1990, Jones
et. al. 1997, Nesom & Brown 1998, Sloan-Nelson 1996, Turner et. al.
2002). Approximately a dozen plants were found blooming in shades of
orange and yellow along a recently constructed roadside with black
clayey soil. These plants may be the result of direct seeding or
contaminants in grass seeds, however, several plants were present at the
same location in spring 2002.

Voucher specimen: Walker Co.: Roadside of Veteran’s Memorial
Parkway app. | mi S of TX Hwy 30, 19 Feb 02, Keith 94 (BRIT).

Zinnia violacea Cav., Elegant Zinnia, has been reported for nine
states in the eastern United States (Gandhi & Thomas 1989, USDA
2002, Wunderlin & Hansen 2003). Approximately, 10 individuals
were found in full bloom along a black, clayey roadside and adjacent
pasture. This species has not been previously reported as escaped from

30 Phytologia (Feb 2004) 86(/)

cultivation in Texas (Correll & Johnston 1970, Hatch et. al. 1990,
Nesom & Brown 1998, Nelson 1996, Turner et. al. 2002), however,
Jones et. al. (1997) list this species as cultivated. The plants appear to
have successfully reproduced because the area where they occurred is
on the edge of a cattle pasture where intentional cultivation was
unlikely. In addition, two plants were found at the same locality in July
2003.

Voucher specimens: Walker Co.: Blackland pasture at intersection of
Veteran’s Memorial Parkway and Smither Ave., 18 Jul 02, Keith 200
(BRIT); Blackland pasture at intersection of Veteran’s Memorial
Parkway and Smither Ave, 4 Aug 02, Keith 206 (SHST).

ACKNOWLEDGMENTS

Thanks to Guy Nesom at the Botanical Research Institute of Texas
for identifying Dimorphotheca sinuata and verifying the identification
of Zinnia violacea.

LITERATURE CITED

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

Gandhi, K. N. and R. D. Thomas. 1989. Asteraceae of Louisiana. Sida
Botanical Miscellany No. 4. SMU Herbarium, Dallas. p.179.

Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1990. Checklist of the
vascular plants of Texas, MP-1655. Texas Agric. Exp. Sta.,
College Station.

Hickman, James C. (Ed.). 1993. The Jepson manual: higher plants of
California. Berkeley, CA: University of California Press. p. 246.

Jones, S.D., J.K. Wipff, and P.M. Montgomery. 1997. Vascular Plants
of Texas. A comprehensive checklist including synonomy,
bibliography, and index. University of Texas Press. Austin.

Phytologia (Feb 2004) 86(1) 34

Nesom, G. L. and L. E. Brown. 1998. Annotated checklist of the
vascular plants of Walker, Montgomery, and San Jacinto Counties,
East Texas. Phytologia 84 (2): 107-153.

Sloan-Nelson, K. B. 1996. Floristic Study of Walker County, Texas:
Asteraceae. Masters of Arts, Sam Houston State University.

Turmer, B.L., H. Nichols, G. Denny & O. Doron. 2003. Atlas of the
vascular plants of Texas. Sida, Bot. Misc. 24:1-888.

USDA, NRCS. 2002. The PLANTS Database, Version 3.5
(http://plants.usda.gov). National Plant Data Center, Baton Rouge,
LA 70874-4490 USA.

Wunderlin, R. P., and B. F. Hansen. 2003. Atlas of Florida Vascular
Plants (http://www.plantatlas.usf.edu/).[S. M. Landry and K. N.
Campbell (application development), Florida Center for
Community Design and Research.] Institute for Systematic
Botany, University of South Florida, Tampa

32 Phytologia (Feb 2004) 86(1)

KEYS TO THE FLORA OF FLORIDA -- 9, OXALIS
(OXALIDACEAE)

Daniel B. Ward

Department of Botany, University of Florida, Gainesville, Florida
32611, U.S.A.

ABSTRACT

Oxalis (Oxalidaceae) is represented in Florida by 8 species, with
O. corniculata treated as consisting of 2 varieties and O. dillenii of 2
subspecies. Extended commentary is provided regarding the correct
typification of O. stricta, with brief discussion addressing the
nomenclature of other species. An amplified key is given to the
Florida taxa. Two species are excluded.

KEY WORDS: Oxalis, Oxalidaceae, Florida flora.

Oxalis (Oxalidaceae) in Florida is a moderate sized, readily
recognized genus with some species sharply defined and others subtly
distinguished and intergrading, some names consistently applied and
others wholly conflicting depending on which student of the genus is
considered authoritative. Though K. R. Robertson (J. Amold Arbor.
52:649-665. 1971) excellently summarized the then-current
nomenclature of the southeastern species as seen by G. Eiten (Taxon
4:99-105. 1955; Amer. Midl. Nat. 69:257-309. 1963; see also: Ph.D.
diss., Columbia Univ. 1959), a later worker, A. Lourteig (Phytologia
42:57-198. 1979), has significantly challenged the application of
certain important species. It is believed the present study may be the
first subsequent effort to resolve the differences in interpretation
between these two authors.

Phytologia (Feb 2004) 86(1) 33

Eiten (1955) has typified Oxalis stricta L. (1753) as the largely
northern, erect plant with septate hairs and slender fleshy rhizomes. A
very different conclusion was reached by Lourteig (1979).

Two species are involved: an erect septate-haired plant common in
both America and Europe (and thus likely familiar to Linnaeus), here
called Oxalis stricta (by Lourteig called O. fontana); and a lax
stemmed nonseptate-haired plant native to America and introduced
rarely to Europe, here called O. dillenii (by Lourteig called O. stricta).
[The plants are unmistakably different on a significant suite of
characters. The hair character, not the most obvious, is used here only
as a convenient label.] Linnaeus (1753:435) cited with his new O.
stricta the pre-Linnaean publications of Gronovius (1743), Tournefort
(1700), and Morison (1680) and, by implication, their associated
specimens and plates; at that time he possessed no specimens of his
own. The underlying specimens and illustrations differ; it is agreed
(Eiten, Lourteig) that the Gronovius reference is to a nonseptate
specimen within the circumscription of O. dillenii, while Morison's
plate corresponds to the septate-haired taxon. [The authors either did
not search the herbaria of Morison (OXF) and Tournefort (P), or found
nothing. Their (unstated) presumption appears to have been that
Morison left no specimen and Tournefort had neither an illustration nor
surviving specimen.] Later, after publishing O. stricta, Linnaeus
obtained a specimen of the septate-haired plant (LINN 600.33) which
he marked as "stricta."

Conflict centers on application of I-C.B.N. (Art. 9.2), the criteria
for selecting a lectotype when more than one specimen or illustration
was available to the original author. Eiten (1955) noted that B. L.
Robinson (J. Bot. 44:386. 1906) had selected as lectotype a Virginia
specimen of Gronovius (BM: Clayton 474), the nonseptate-haired
plant, but argued that this selection was contrary to Linnaeus's intent.
He then selected as lectotype the Morison plate (t. 17, f. 3) of the erect
septate-haired plant. This action led him to use O. stricta and O.
dillenii Jacq. (1794) for the septate-haired and nonseptate-haired plants
respectively. Lourteig (1979:60) was not persuaded. She -- without
reference to the actions by Robinson or Eiten -- argued that Linnaeus

34 Phytologia (Feb 2004) 86(1)

had worked with Gronovius (and, by implication, was familiar with his
specimen), while it was "unlikely he had in mind the specimens of
Tournefort and of Morison;" she selected the Gronovius specimen as
lectotype. She then used O. fontana Bunge (1833) and O. stricta for
the septate- and nonseptate-haired plants, respectively.

Although a lectotype may be superceded if original type materials
are rediscovered or if one can show that the lectotype is in serious
conflict with the protologue (I-C.B.N., Art. 9.17), it is not common for
authors to propose reversal of their predecessor's lectotypifications.
The present situation is further unusual in that argument is made for a
double reversal -- Eiten (1955) of Robinson (1906), and Lourteig
(1979) of Eiten (1955). It must tax even the nomenclaturally adept
reader to follow the arguments and to decide whether to use O. stricta
and O. dillenii for the septate and nonseptate plants, as did Eiten, or use
O. fontana and O. stricta, as did Lourteig. Practice, as in so many
other examples, has been for writers to take the easier pathway and
docilely to follow the conclusions of the latest author. Thus, after Eiten
(1955, 1963), American writers commonly used O. stricta for the
septate-haired plant, and after Lourteig (1979) -- without a whimper of
protest or, for that matter, indication of understanding -- have
employed that name for the nonseptate-haired plant. Eiten's perhaps
excessively detailed style, and Lourteig's failure to mention Eiten's
contrary action or to refute the specifics of his argument, is not helpful.

But even with the complexity of the arguments as presented, still
other factors may be mentioned. Though a type may be a specimen or
an illustration (I.C.B.N., Art. 8.1), the superiority of actual plant
material is well appreciated; had no lectotype yet been chosen,
selection of the nonseptate Gronovius specimen over the Morison plate
would be favored. That preference, though not in itself determining
typification, may have influenced Robinson (1906) in his selection as
type the Virginia specimen rather than the European illustration. Of
greater weight, however, and pointing in the opposite direction, is the
probability, not well brought out by either author, that Linnaeus would
likely have known the plant often found as a weed in European
gardens, added later to his herbarium, and illustrated by Morison, while

Phytologia (Feb 2004) 86(1)

Ww
N

he would have encountered the nonseptate-haired American
introduction only by brief contact with the Gronovius specimen, if at
all. Further, his epithet is apt only for the European plant; that species
is quite erect, or strictus (at least when young), while the plant
represented by the Gronovius specimen is invariably lax and sprawling.
Additional importance can be attributed to the dominant (but not
universal) European practice of recognizing the plant common there as
O. stricta.

One thus has the choice of accepting the earlier Robinson
lectotypification and typifying O. stricta as a plant Linnaeus scarcely
(or never) knew, or rejecting that lectotypification and re-lectotypifying
O. stricta as the plant familiar in Europe and probably to Linnaeus.
The advantages of the latter choice are obvious. Stability without
doing violence to nomenclatural precision is best attained by rejection
of the early Robinson (1906) lectotypification of the nonseptate-haired
plant (per Eiten, 1955), and selection of the erect, septate-haired plant
common both in Europe and eastern North America (and introduced
into Florida) as the basis for the name Oxalis stricta L.

The judgment of Eiten (1963) is cautiously accepted here that
variation within Oxalis dillenii is best apportioned by recognition of
ssp. dillenii and ssp. filipes. In contrast, Lourteig (1979) held these
taxa at specific rank, with different names; she recognized O. stricta
(as discussed above) and O. florida Salisb. The differences (in
pubescence, and size and robustness of the plants, as noted in the key)
are appreciable and intermediates seem few. In respects ssp. filipes is
closer to O. corniculata than to ssp. dillenii (Eiten, 1963:268).
Lourteig's unwillingness to address Eiten's arguments leaves one
reluctant to endorse her conclusions and nomenclature. Yet it would
not appear unreasonable for one to recognize the two taxa as species --
as have essentially all previous authors -- with the names O. dillenii
and O. florida. (Eiten (1963:301) was incorrect in claiming O. florida
is superfluous (I.C.B.N., Art. 52.1). The name cited in synonymy by
Salisbury (1796) was pre-Linnaean which, since not available for his
use, does not disturb the legitimacy of O. florida.}

36 Phytologia (Feb 2004) 86(1)

The assignment of Oxalis corniculata L. (1753) is now established
by Eiten's (1955) lectotypification of the name. In Florida two distinct
forms occur: leaves green, variable in size, fruits glabrous; and leaves
bronze-purple, small, fruits long-pubescent. Eiten (pers. comm., Oct
1967) has been unwilling to assign varietal names. Lourteig (1979)
treated these taxa as O. corniculata var. corniculata, and var.
atropurpurea Planch. (incl. Xanthoxalis Langloisii); she also
recognized var. villosa (Bieb.) Hohen.

Decision has been deferred as to proper treatment of Oxalis
priceae Small. This name represents a complex of perennial
(rhizomatous), relatively large flowered, little-known taxa usually
restricted to undisturbed woodlands and too easily dismissed as waifs
of O. dillenii or other more common species. Eiten (1963) divided O.
priceae into three subspecies distinguished on features of stem and
pedicel pubesence. Two, ssp. colorea (Small) Eiten, and ssp. texana
(Small) Eiten, were reported to reach Florida. Lourteig (1979) -- again
without reference to the arguments of Eiten -- recognized three species,
one with two subspecies. Comparison is difficult because this second
author interpreted both the taxa and their types quite differently and
permitted no taxa to bear the names assigned by the first author.
Further study may justify some degree of separation. [D. M. Mulcahy
(Amer. J. Bot. 51:1045-1050. 1964), incidental to his primary interest
in flower form, reported variations in growth patterns and heterostyly
among the three taxa.] But, at least within Florida, the difference are
so subtle and intergrading that they are not believed worthy of
recognition.

The renaming of Oxalis priceae as O. lyonii Pursh (1813), by
Lourteig (1979), is in error. The type of O. lyonii was from
Cumberland Island, an early-settlement plantation in southeastern
coastal Georgia. Lourteig, finding Pursh's type to be lost and lacking
other collections from near the type locality that she could assign to O.
lyonii, neotypified her interpretation of the name with a Mississippi
specimen of Eiten's ssp. texana. However that entity, a rather
uncommon perennial of dry, usually undisturbed woodlands,
apparently does not extend eastward to Florida, much less the Atlantic

Phytologia (Feb 2004) 86(1) S|

coastal plain. [Reports of its presence in Florida are based on
misidentified specimens (Cooley - USF) or specimens of ssp. colorea.]|
Pursh's description of O. lyonii is mostly language appropriate only to
the genus, but contains phrases ("...caule ramoso decumbente...siliquis
tomentosis...") exactly fitting O. dillenii, a species common on
agricultural soils of eastern Georgia. [O. priceae is erect above its
rhizomatous base, with finely pubescent fruits.| Though claimed
otherwise by Lourteig, no part of Pursh's description uniquely implies
he had in hand any member of the O. priceae complex. True O. lyonii
is probably a synonym of O. dillenii, as has been supposed by previous
authors (Eiten, 1963.; etc.). Lourteig's neotype is clearly in conflict
with the protologue of O. lyonii, and should be set aside. If the
complex is left undivided, O. priceae retains its priority.

Oxalis violacea is so clearly a native in the northern states that it is
easily assumed to be so in Florida. Yet its habitat here is essentially
confined to roadsides and other disturbed, widely separated areas.
Northward, its coherent range largely stops in mid-Georgia (S. B. Jones
& N.C. Coile, Distribution of the Vascular Flora of Georgia. 1988). It
is best treated as introduced.

The introduced South American species of Sect. Jonoxalis are
perhaps best addressed by M. E. Denton (Publ. Mus. Michigan State
Univ. Biol. 4:455-615. 1973). Introduced species are denoted by an *.

The "amplified key" format employed here is designed to present
in compact form the basic morphological framework of a conventional
dichotomous key, as well as data on habitat, range, and frequency.
This paper is a continuation of a series begun in the 1970s (vide
Phytologia 35:404-413. 1977). I wish to thank David W. Hail and
Kent D. Perkins for constructively reviewing the manuscript, and to
acknowledge extended correspondence with Dr. George Eiten and his
assistance with certain identifications, though all nomenclatural
interpretations are my own.

38 Phytologia (Feb 2004) 86(1)

Oxalis L. Wood Sorrels

1. Petals yellow to orange-yellow; sepals without orange dots
(tubercles) at apex; plants annual or short-lived perennial; base
fibrons’orof thin Thizomes2 2202 A Sect. Corniculatae

2. Hairs of petioles and stem septate (multicellular); stems erect,
arising from a slender shallow rhizome; annual or short-lived
perennial herb. Floodplains. Mid-panhandle (Liberty County:

Apalachicola River); rare. Summer-fall......... *Oxalis stricta L.
[O. europaea Jord.; O. fontana Bunge; Xanthoxalis cymosa
Small]

2. Hairs of petiole and stem non-septate (unicellular); stems
creeping or decumbent from fibrous roots, or erect from a
perennial base (if erect, corolla >12 mm. long).

3. Corolla <11 mm. long; longer internal flower structures
(either stamens or pistil) <S mm. long; stems creeping or
decumbent, from fibrous roots.

4. Mature capsules gradually tapering to apex; seeds and their
transverse ridges uniformly brown; stems creeping, rooting
at nodes; perennial herb. Moist soil of gardens, lawns. All
WCAG: AEF 2.: Hath Kt dooaee, eee *Oxalis corniculata L.

4a. Leaflets ca. 1.5 cm. broad, green; stems both creeping
and ascending-erect. Throughout Florida: common

SEE BULA I BR I, OA: var. corniculata
[Xanthoxalis corniculata (L.) Small]

4a. Leaflets ca. 1 cm. broad, dark maroon; stems closely
creeping. Throughout Florida; common...................

seihs OS METAS: AS var. atropurpurea Planch.
[Xanthoxalis langloisii Small]

Phytologia (Feb 2004) 86(1) 39

4. Mature capsules with parallel sides, abruptly tapering to
apex; seeds brown with gray or white crests or distinct spots
on transverse ridges; stems often decumbent at base but
only rarely rooting at nodes; annual or perennial herb.
Gardens, pastures, moist disturbed areas. Spring-summer.
DE PON EEE CE PEPE OEE PORE PETER CECER Oxalis dillenii Jacq.

4b. Capsules densely pilose; stem hairs strictly appressed
upwards; plants relatively robust (stems 1.5-2 mm.
thick). Nearly throughout (south to Poik, Brevard
counties; excl. south peninsula); common...ssp. dillenii

[O. lyonii Pursh; Xanthoxalis stricta, misapplied]

4b. Capsules partially or completely glabrous; some or all
stem hairs retrorse; plants relatively slender (stems 1-
1.5 mm. thick). Throughout Florida; common........
peut A sa beth teeth. sect. ch aadnemvews, ssp. filipes (Small) Eiten
[O. florida Salisb.; Xanthoxalis brittoniae Small;
Xanthoxalis filipes Small]

3. Corolla 12-14 mm. long; longer internal flower structures
(either stamens or pistil) 6-8 mm. long; stems erect from base,
with short rhizomes; perennial herb. Mesic woodlands.
Panhandle (Escambia County, east to Gadsden, Liberty
counties); infrequent. Spring. Restricted to undisturbed
habitats and often overlooked............... Oxalis priceae Small
[O. lyonii, misapplied; Xanthoxalis colorea Small;
Xanthoxalis macrantha, misapplied; Xanthoxalis priceae
Small; Xanthoxalis recurva, misapplied]

1. Petals pink to lavender; sepals usually with orange dots (tubercles)
at apex; plants perennial; base stocky, tuberous or bulbous.........
blast? tists. Gk Speeds sexe Ta eke s huis. aces parents pl aesten reels Sect. Ionoxalis

40 Phytologia (Feb 2004) 86(1)

5. Leaflets angular obcordate or obdeltoid; petals pinkish purple;
sepals 4 mm. long, with distinct orange dot at tips; perennial
herb. Moist waste areas, fencerows. Peninsula (Duval, Alachua
counties, south to Dade County); infrequent. Spring-summet......
Bas Le aa sear al tes ea Eee oes * Oxalis intermedia A.
Rich.

[O. latifolia, misapplied; Jonoxalis intermedia (A. Rich.) Small]

5. Leaflets rounded obcordate.

6. Plant arising from a segmented, woody taproot; leaflets <2.5
cm. broad; sepals densely appressed-pubescent; petals red or
pinkish red (rarely white); inflorescences delicate, usually
compound, often many (to 100) per plant; perennial herb.
Moist disturbed areas. West and mid-panhandle (Escambia to
Leon, Franklin counties); rare. Spring...* Oxalis rubra St.-Hil.

6. Plant arising from a fleshy bulb or cluster of bulblets; sepals
glabrous or sparsely pubescent toward base; inflorescences
usually few (1-10) per plant.

7. Leaflets 2.5-5.0 cm. broad; inflorescence compound; petals
purplish pink; sepals 5 mm. long, with no or indistinct
orange dot (tubercle) at tips; perennial herb. Fencerows,
dooryards, moist disturbed areas. Throughout; mostly near
habitations, where often common. Winter-spring............
eA tas cee es, Ae * Oxalis corymbosa DC.
[O. debilis HBK. var. corymbosa (DC.) Lourteig; O.
martiana Zucc.; lonoxalis Martiana (Zucc.) Small]

7. Leaflets 1.5-2.0 cm. broad; inflorescence simple; petals
violet; sepals 3 mm. long, with distinct orange dot (tubercle)
at tips; perennial herb. Dry waste areas, rocky outcrops.
Mid-panhandle (Calhoun, Jackson counties, scattered east to
Putnam County); rare. Spring, fall. Not persisting.
Jones asaiseicle ds VIOLET WOOD SORREL..* Oxalis violacea L.

Phytologia (Feb 2004) 86(1) 4]

[lonoxalis violacea (L.) Small]
Excluded names:

Oxalis pes-caprae L. Buttercup Oxalis
Oxalis cernua Thunb.

Bolboxalis cernua (Thunb.) Small
Reported for "waste-places and cult. grounds, N. Fla." (Small, 1933).
Perhaps once cultivated. No Florida specimens are known.

Oxalis triangularis St. Hil.

Reported [as ssp. papilionacea (Hoffmanns. ex Zucc.) Lourteig] for
Leon Co. (Wunderlin, Hansen & Anderson, 2002), based upon a 2001
specimen (Anderson 19691 - FSU). Stated as "several plants," but
without clear evidence of naturalization.

42. Phytologia (Feb 2004) 86(1)

BIOLOGICAL STATUS OF THE NAMES ABRONIA
CARLETONII AND A. NEALLEYI (NYCTAGINACEAE)

B. L. Turner

Plant Resources Center University of Texas Austin, TX, 78713,
U.S.A.

ABSTRACT

The taxonomic history of Abronia carletonii Coulter & Fisher and
A. nealleyi Standl. is reviewed. Galloway (1975), in his revisionary
treatment of the genus Abronia for the North America desert region,
concluded that the two names were synonymous, A. carletonii having
priority. After examination of the holotype of the latter, this collected
in eastern Colorado, I conclude that it is a late-flowering atypical
element of the widespread, highly variable, A. fragrans Nutt. ex
Hook.f Abronia nealleyi is typified by material from Reeves Co.,
Texas. It is a localized species, occurring on bare gypsum outcrops in
northern Culberson and Reeves counties of Trans-Pecos, Texas and
closely adjacent New Mexico (Eddy Co.).

KEY WORDS: Abronia, Nyctaginaceae, Texas

Abronia carletonii was proposed by Coulter & Fisher in 1892,
basing this upon a single specimen collected in "eastern Colorado" in
1891 by M. A. Carleton (holotype: Carleton459, FM!). Its authors
thought their new species to be:

Most closely related to A. turbinata Torr., having the
coriaceous double wing of the section, but differing from
that species in having slender white glabrous (but minutely
glandular) stems, more numerous flowers, broader rose-
colored attenuate or cuspidate bracts, and the perianth and
its lobes not so deeply cut.

Phytologia (Feb 2004) 86(1) 43

M. E. Jones subsequently reduced A. carletonii to varietal status
under A. turbinata. Standley (1909), however, retained A. carletonii
(at the time this still known only by the type), noting that "It is not the
same as A. angustifolia [sic] as Mr. Jones claims, but seems to me
much nearer A. fragrans."

Galloway (1975) also retained A. carletonii but included in this A.
nealleyi, the latter based upon a single collection made by Nealley in
1893 from "Screw Bean," Reeves Co., Texas. Standley (1909) has
provided an excellent sketch of A. nealleyi.

In addition to the types mentioned in the above, Galloway cited
and/or mapped eight additional collections of A. carletonii, as follows:

New Mexico. Eddy Co.: two collections mapped, but not cited.
San Miguel Co.: San Jeronimo, Benedict 2334 (US).

Texas. Brewster Co. [sic]: " About 25 mi. E of W end of
Paso-Tex pipe line road, Correll & Rollins 23192
(LL). The locality concerned is actually in northern
Culberson or Hudspeth counties.

Culberson Co.: two collections mapped, but not cited.
El Paso Co.: two collections mapped, but not cited.

Hudspeth Co.: "Ables", Parks & Cory 1504 (TAES).

I consider all of the above cited specimens to be A. nealleyi
(except for the collection from San Miguel Co., New Mexico, which is
probably an aberrant specimen of A. fragrans; at least it does not
belong to A. nealleyi as conceived here).

My interest in the several names of Abronia mentioned in the
above was stimulated by my attempt to place a name upon an erect
perennial herb having lanceolate leaf blades that seemed confined to

44 Phytologia (Feb 2004) 86(1)

bare gypsum outcrops of north-cenrtral Trans-Pecos, Texas and
closely adjacent New Mexico (Eddy Co). Field work revealed
numerous populations of the taxon concerned along state highway 652
from its entry into New Mexico eastwards to Orla, Texas, this also
attested to by numerous vouchers on file at SRSC and LL-TEX.
Careful study of this material, along with a study of the literature,
strongly suggested that these populations are best referred to A.
nealleyi.

Application of the name A. carletonii is moot. Its distribution as
portrayed by Galloway (as noted above) made little
morphogeographical sense, nor did he map the taxon as occurring in
Colorado, where the type was reportedly collected. To resolve the
problem I borrowed the type of A. carletonii. A brief description of the
type and comments upon its likely biological status follows.

Comments upon the type of Abronia carletonii

The holotype lacks a root and appears to be a late-flowering sprig
taken from a prostrate stem devoid of primary leaves. Its flowering
secondary branches possess relatively small flowers. Except for its
prostrate stems, the plant superficially resembles A. nealleyi. Closer
inspection, however, shows that the stems and leaves are lacking the
villous vestiture of A. nealleyi and, along with its depauperate state
and sporatic distribution as displayed by Galloway, I have no
hesitation in referring the type material of A. carletonii to A. fragrans.
The latter species is common throughout the western portions of the
central grasslands, namely Wyoming, Colorado, and New Mexico, as
is obvious from the dot maps of the species provided by Galloway
(1975) and Barkley (1976). Indeed, an immature anthocarp from the
type of A. carletonii (now deposited in a packet on the type sheet
itself) shows this to possess five compressed wings, the latter not
dilated at their apices. Using Galloway's key to the species of Abronia,
if annual, the type itself would key to A .fragrans. Regardless, for the
Atlas of Texas Plants (Turner et al. 2003) I should have taken up the
name A. nealleyi for material occurring on the gypsum out crops of
Culberson and Reeves counties formally called A. carletonii by most

Phytologia (Feb 2004) 86(1) 45

workers (e.g.. Reed 1969; Correll and Johnston 1970; Johnston 1988;
and yet others). The present contribution calls attention to the
misnomer concerned.

ACKNOWLEDGEMENTS

I am grateful to the staff at Field Museum (FM) for the loan of
type material of A. carletonii, and to Mike Powell and Gayle Turner
for reviewing the manuscript.

LITERATURE CITED

Barkley, T. [ed.] 1976. Atlas of the Flora of the Great Plains. Iowa
State Univ. Press, Ames.

Correll, D.S. and M.C. Johnston 1970. Manual of the Vascular Plants
of Texas. Texas Research Foundation, Renner, Texas.

Galloway, L.A. 1975. Systematics of the North American desert
species of Abronia and Tripterocalyx (Nyctaginaceae). Brittonia27:
328-347.

Johnston, M.C. 1988. The Vascular Plants of Texas [an updating],
privately published by the author, Austin, Texas.

Reed, C.F. 1969. Nyctaginaceae, in Flora of Texas 2: 151-220.

Standley, P.C. 1909. The Allionaceae of the United States , with notes
on the Mexican species. Contr. U. S. Nati Herb. 12: 303-389.

Turner, B.L., H. Nichols, G. Denny & O. Doron. 2003. Atlas of the
vascular plants of Texas. Sida, Bot. Misc. 24:1-888.

46 Phytologia (Feb 2004) 86(1)

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