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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
FAX (936) 294-3940 Subscription Phytologia Information & P.O. Box 2012 Scholarly Exchange Sam Houston State University
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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.
LITERATURE CITED
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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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