© PHYTOLOGIA
An international journal to expedite plant systematic, phytogeographical
and ecological! publication
Vol RAe “February 1998 No. 2
CONTENTS
TURNER, B.L., Podachenium chimalapanum (Asteraceae), a new species from
PASIGERIHOSE MOAR HGd: IVIENIOO: se ne eee 61 —~
~ WALKER, S:B., The status of Obolaria virginica L. (Gentianaceae) in Texas
and surrounding Slates Cece erates pian cosa ah ana Selgt ee es 64
Ee GRANT, V., Classification of the genus Gilia (Polemoniaceae)................. 69
2 - TURNER, B. L., A new species of Neurolaena. (Asteraceae) from Oaxaca,
Ne SAVIO MICO Son rs Sukcan' hain uve pany eae tay Antu guase rewmgtiats center se mveras ot, 87
» WHITE, H.L., JR. BRANCH, & W.C. HOLMES, Comments on_ the
ae distribution of Tillandsia recurvata L. (Bromeliaceae) in Texas.............. 93
| HERSHKOVITZ, M.A., Parakeelya: a new genus segregated from Calandrinia
i ee PONCHO C Rey se), wove er eo neg terse nee Sub rgd CC Et Bi rn antes 98
a NESOM, G.L. & L.E. BROWN, Annotated checklist of the vascular plants of 1
+ Walker, Montgomery, and San Jacinto counties, East Texas................ 107
» MATHIASEN, R.L., C.G. PARKS, B.W. GEILS, & J.S. BEATTY, Notes on
the distribution, host range, plant size, phenology, and sex ratio of two rare
dwarf mistletoes from Central America: Arceuthobium hawksworthii and A.
4 HORAUPEMSE: Opis etegeel eas ay a adg aac has ay war sted yo gtye itp gh 154. 7
- ‘New names in this issue of Phytologia. hicdarde wt nay an AS a eae Lap 165
Beeack issuchiavedable; bert mio oo ere eine iat ce a ie 167
Bas Pnytolopad Memoirs aVallaDle: 5 tare cs Gus fatto li ccs wagonGenioaaeseeoadney 168
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Phytologia (February 1998) 84(2):61-63.
PODACHENIUM CHIMALAPANUM (ASTERACEAE), A NEW SPECIES FROM
EASTERNMOST OAXACA, MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
A new species, Podachenium chimalapanum B.L. Turner, is
described and illustrated from easternmost Oaxaca, México. It is closely
related to P. pachyphyllum and P. chiapanum, but somewhat closer to the
former, occupying a geographical position between these two taxa.
KEY WORDS: Asteraceae, Heliantheae, Podachenium, Oaxaca, México
Routine identification in connection with a forthcoming treatment of the tribe
Heliantheae for the Comps of Mexico (cf. Turner 1997, The Comps of Mexico, [Vol.
10], Phytologia Memoirs) has occasioned the present paper.
PODACHENIUM CHIMALAPANUM B.L. Turner, spec. nov. Figure 1.
TYPE: MEXICO. Oaxaca: Mpio. Sta. Maria Chimalapa, Cerro Piicotzuc, ca. 18
straight-line km SSE of La Laguna Veracruz, top of western peak, granitic ridge
with very steep, in parts, almost vertical slopes (to N and S), growing among
“very dense cloud scrub” 1.0-1.5 m high, very exposed to wind and fog, with
Cavendishia, Gaultheria, etc. (17° 07’ 25” N, 94° 27’ 15” W), 1400 m, 5 Apr
1996, Thomas Wendt et al. 6806 (HOLOTYPE: MEX!; Isotype: TEX!).
Simile P. pachyphyllo (Sch. Bip. ex Klatt) Jansen sed folia parviora et
capitulescentias congestiores cum capitula pauciora habens.
“Shrub” to 1.5 m(?). Stems sparsely pubescent when young, but soon glabrate.
Larger leaves ovate-elliptic, mostly 6-9 cm long, 3-4 cm wide; petioles 5-7 mm long;
blades grading into the petioles, pinnately nervate, glabrous on both sides, the upper
surfaces drying black, the margins serrulate. Heads arranged 3-13 in rather
congested terminal cymose panicles, the ultimate peduncles 1.0-2.5 cm long.
Involucres 3.5-4.5 mm high, ca. 7 mm across; bracts 2-seriate, subequal, the outer
61
62 PHYTOLOGIA _ February 1998 volume 84(2):61-63
UNIVERSITY OF
Exas
HERBARIUM
pba 2 bf Padvchanio oo malepeegen
| Va Toeew
Merbarkam °| Ss Lourvesee State Unverarty, Gotan Rowse
FLORA OF THE CHIMAL APA-UXPANAPA REGION
OAXACA-VERACRUZ, MEXICO
comecstie S ups.
Dovac semrum ehimelaganum St (oer
Si omer
OAXACA: Mow. Sta Mane Crimaiane Cen Pacotzet.
‘ghee ocak mm Centra Serra oe Tos Poos (ca. 12 shape
(ww wr SSE of La Laguna ver) top of western (sigrily
eet, Dt CfANEe (Ste wet) very On oreceeaion soe
wal vey deep 2 pans urnost vertical siopes (to N ang S:
Fost WO" wety denge Cloud torup 1-15 m tat very exposed
fl amo tog wtf Coe Caeranut Gauwenw Lica
Cw blunca Ciaewooraa Svomte '7°9725N
4ST SW cay. 100m
27D CI Very SICeD RCDES. Taya atta. Asc yadow
Fed bea e 7 Celt kee Oana « 7c Oey
Co: T. Wend, Hererto Hernancez G_ P. Tenorio, &
Zea Torres, G. Salazar, ML A Soto, O. Rochs 6806
5 Apri 1356
ersed 0 Ihe Neons Congr sates Receoty. Geant SH, “Ran
Fares) Tree Obversty cs he oceans of Tohoenmpet Bermo”
Fig. 1. Podachemium chimalapanum. isotype TEX.
Turner: New Podachenium from Oaxaca 63
series spatulate and loose, the inner less so. Receptacle moderately convex, ca. 3 mm
across, glabrous; chaff indurate, persistent, their apices acute, somewhat pungent to
the touch. Ray florets ca. 11, neuter; ligules white, 6-8 mm long. Disk florets
numerous; corollas yellow, 2.5-3.0 mm long, the tube ca. 0.5 mm long, the lobes ca.
0.8 mm long. Achenes flattened radially, the body ca. 2.5 mm long, 0.9 mm wide,
the podocarp poorly developed; pappus of 2-5 principal scales 1.0-1.5 mm high,
between these 1 or more smaller scales.
This novelty is closely related to Podachenium pachyphyllum (Sch. Bip. ex Klatt)
Jansen of northcentral Oaxaca, differing from this in having somewhat smaller, ovate-
elliptic leaves, fewer heads (3-13 vs. 15-30) arranged in a much more congested
capitulescence, and having heads with loosely arranged outer phyllaries (vs.
appressed). Turner & Panero (1992) provided a key to the four species of
Podachenium known to them; in this treatment P. chimalapanum will key readily to P.
pachyphyllum, except for its fewer, shorter ligules (ca. 11 vs. 13, 6-8 mm long vs.
15-20 mm long, this latter range inaccurately stated in the table provided by Turner &
Panero). But, as indicated in the above, P. chimalapanum differs notably from P.
pachyphyllum in having a very congested, relatively few-headed capitulescence, the
species standing somewhere between the latter and P. chiapanum B.L. Turmer &
Panero; geographically, it also stands between these two taxa.
The species is named for the Chimalapan region of eastern Oaxaca, to which it
seems confined.
ACKNOWLEDGMENTS
I am grateful to Gayle Turer for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the manuscript.
LITERATURE CITED
Turner, B.L. & J. Panero. 1992. New species and combinations in Podachenium
(Asteraceae, Heliantheae). Phytologia 73:143-148.
Phytologia (February 1998) 84(2):64-68.
THE STATUS OF OBOLARIA VIRGINICA L. (GENTIANACEAE) IN TEXAS
AND SURROUNDING STATES
Suzanne Birmingham Walker
Rt. 2 Box 191, Shelbyville, Texas 75973 U.S.A.
ABSTRACT
A recent find of Obolaria virginica L. from southeastern Texas led to a
review of the status of this species in Texas and surrounding states.
KEY WORDS: Obolaria virginica L., Texas, Gentianaceae
Obolaria virginica L., Pennywort, a northeastern species, reaches its western limit
in southeastern Texas (Correll & Johnston 1970; Johnston 1990; Hatch et al. 1990;
Jones et al. 1997). It is not known from Oklahoma (Taylor & Taylor 1989). In
Arkansas it occurs only in several east central counties (Smith 1988), and in Louisiana
it is reported from fifteen parishes (Thomas & Allen 1998; MacRoberts & MacRoberts
1995).
Obolaria, a monotypic genus, and Bartonia are the only saprophytic genera of the
Gentianaceae found in North America (Maas 1986). Obolaria virginica is found
chiefly in deciduous forests but is also found in rich woods, river banks, mixed
hardwood slopes, and bottom lands (Andreas 1970; Brown 1972; Gillet 1959;
MacRoberts & MacRoberts 1995; Hammond-Soltis 1979). It is said to be common in
rich mesic woods throughout the piedmont and mountainous areas in the southeastern
United States (Kondo 1970). It is easily overlooked in the early stages of growth
(Brown 1972).
On February 4, 1998 while searching for Cardamine concatenata (Michx.) Sw..,
Sanguinaria canadensis L., and Uvularia perfoliata L., 1 found Obolaria virginica
growing in a side slope of a beech-hardwood community in the Angelina National
Forest in Angelina County. It was located within four meters of a dry, shallow,
undeveloped side drain of a larger system. The overstory and midstory species in this
site were Quercus alba L., Q. michauxii Nutt., Fagus grandifolia Ehrh., Ostrya
virginiana (P. Mill.) K. Koch, Magnolia grandiflora L., Ilex opaca Ait., Acer
barbatum Michx., and Pinus taeda L. This appears to be the remnants of perhaps a
once larger hardwood community that was reduced when the surrounding area was cut
and replanted with loblolly pine, in some areas as close as seven meters to the drain.
64
Walker: Status of Obolaria virginica in Texas 65
The duff layer was quite thick and ground cover species found at the time were limited
to Mitchella repens L. and Sanicula sp. Corallorhiza wisteriana Conrad, another
uncommon species in east Texas, was also found near the site.
This type of community has been described in east Texas, where it reaches its
southwestern limit, by McLeod (1972), Diamond et al. (1987), Marks & Harcombe
(1981), Nixon et al. (1980), MacRoberts & MacRoberts (1997). Through restoration,
protection, and improvement, the U.S. Forest Service hopes to achieve a target of
3500 acres of this community in the National Forests and Grasslands in Texas
(NFGT). Currently NFGT claims to have about 2500 acres of beech-white oak forest
in its holdings (USDA 1996a).
I surveyed numerous herbarium collections (TEX, NO, BRIT, ASTC, LTU,
SFRP, LSU, NLU, DUR) for Obolaria from Texas. GH, MICH, and MO have no
specimens of Obolaria from Texas (J. Pringle, pers. comm.). Only two specimens
were found. One is from Jasper County and the other from Sabine County, both in
southeastern Texas. It has been recently reported that a new county record has been
found and a new location was found in an earlier documented county (W. Holmes,
pers. comm.), but the information has not been released. Although MacRoberts &
MacRoberts (1995) located O. virginica in a similar habitat in the Kisatchie National
Forest in Louisiana, floristics of beech-hardwood sites in the Sabine National Forest in
Texas did not include the species (MacRoberts & MacRoberts 1997).
While Obolaria virginica is not on the state rare species list, it is eligible for listing
as it is only known from three sites, fewer than six occurrences being classified as S1,
(i.e., critically imperiled in Texas and especially vulnerable to extirpation from the
state). Beech-Hardwood Forest is rare in east Texas. The Texas Organization for
Endangered Species (TOES) lists it as a threatened community and the Texas Natural
Heritage Program lists it as imperiled. The U.S. Forest Service lists this community
as sensitive, although it has no listing for O. virginica (USDA 1996b).
The distribution of Obolaria virginica in Arkansas, Louisiana, Oklahoma, and
Texas is shown in Figure 1. Documentation is given for only Louisiana and Texas.
DOCUMENTATION
LA: Bienville Parish, Thomas 6666 [NLU]. Caldwell Parish, Marx 474 [LTU];
Thomas 38170 [LTU]; Thomas 51173 [LTU]. Catahoula Parish, Thomas 127
[NLU]; Thomas 128 [NLU]. Claiborne Parish, Lewis 129 [NLU]; Lewis 2650
[NLU]. Grant Parish, Parker 570 [NLU]. Lincoln Parish, Pavlu 50] [LTU].
Morehouse Parish, Pias & Thomas 49776 [LSU]; Thomas 12936 [NLU]; Thomas
58135 [NLU]. Natchitoches Parish, Holmes 4424 [NLU]. Ouachita Parish, Thomas
13906 [NLU]. Rapides Parish, Carr, MacRoberts, & MacRoberts 2168 [VDB];
MacRoberts & MacRoberts 2254 [SFRP]. St. Helena Parish, Allen 15593 [LSU].
Webster Parish, Price s.n [NLU]. W. Feliciana Parish, Cocks s.n [NO]; Penfound
s.n [NO]. Winn Parish, Parker 588 [NLU].
PHYTOLOGIA February 1998 volume 84(2):64-68
ATH EA Sea
Pr Ot SS
tribution of Obolaria virginica
isiana, Oklahoma and Texas.
Walker: Status of Obolaria virginica in Texas 67
TX: Angelina Co., Walker 136 [TEX]. Jasper Co., Lundell 13368 [TEX].
Sabine Co., Correll & Correll 24883 [TEX].
ACKNOWLEDGMENTS
Anne Bradburn, Tom Wendt, James Pringle, Walter Holmes, Mark Mayfield, R.
Dale Thomas, Lindsay Woodruff, Phil Hyatt, Milan Vavrek, and Stanley Rice were
helpful in establishing a distribution of the species. A special thanks to Guy Nesom
for reviewing an earlier version of this paper and for help in the distribution research.
Barbara & Michael MacRoberts were especially helpful in all aspects of this work and
reviewed an earlier version of this paper.
LITERATURE CITED
Andreas, B.A. 1970. The Herbaceous Gentianales of Ohio. Master’s Thesis, Kent
State University, Kent, Ohio.
Brown, C.A. 1972. Wildflowers of Louisiana and Adjoining States. Louisiana State
University Press. Baton Rouge, Louisiana.
Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of Texas.
Texas Research Foundation, Renner, Texas.
Diamond, D.D., D.H. Riskind, S.L. Orzell. 1987. A framework for plant
community classification and conservation in Texas. Texas J. Sci. 39:203-221.
Gillet, J. 1959. A revision of Bartonia and Obolaria (Gentianaceae). Rhodora 61:43-
63.
Hammond-Soltis, G.A. 1979. A Taxonomic Treatment of the Gentianaceae in
Virginia. Master’s Thesis, College of William and Mary, Williamsburg, Virginia.
Hatch, S.L., K.N. Ghandi, & L.E. Brown. 1990. Checklist of the Vascular Plants
of Texas. Texas Agricultural Experiment Station, College Station, Texas.
Johnston, M.C. 1990. The Vascular Plants of Texas. Published by the author.
Austin, Texas.
Jones, S.D., J.K. Wipff, & P.M. Montgomery. 1997. Vascular Plants of Texas: A
Comprehensive Checklist Including Synonymy, Bibliography, and Index.
University of Texas Press. Austin, Texas.
Kondo, K. 1970. The chromosome number of Obolaria virginica L. (Gentianaceae).
Rhodora 72:551-553.
Maas, P.J.M. 1986. Flora neotropica, monograph no. 41-42 0071-5794.
Organization for Flora Neotropica, New York Botanical Garden, New York, New
York.
MacRoberts. B.R. & M.H. MacRoberts. 1995. Noteworthy vascular plant
collections on the Kisatchie National Forest, Louisiana. Phytologia 78:291-313.
MacRoberts, B.R. & M.H. MacRoberts. 1997. Floristics of beech-hardwood forest
in east Texas. Phytologia 82:20-29.
68 PHYeLOLOGIA February 1998 volume 84(2):64-68
MacRoberts, D.T. 1989. A Documented Checklist and Atlas of the Vascular Flora of
Louisiana. Bull. Museum of Life Sciences. No. 9. Louisiana State University,
Shreveport, Louisiana.
Marks, P.L. & P.A. Harcombe. 1981. Forest vegetation of the Big Thicket,
southeast Texas, Ecological Monographs 51:287-305.
McLeod, C.A. 1972. The Big Thicket of eastern Texas: A brief historical botanical
and ecological report. Sam Houston State University, Huntsville, Texas.
Nixon, E.S., K.L. Marietta, R.O. Littlejohn, & H.B. Weyland. 1980. Woody
vegetation of an American Beech (Fagus grandifolia) community in east Texas.
Castanea 45:171-180.
Orzell, S.L. 1990. Inventory of national forests and national grasslands in Texas.
Unpublished report, Texas Natural Heritage Program, Texas Parks and Wildlife
Department. Austin, Texas.
Smith, E.B. 1988. An Atlas and Annotated List of the Vascular Plants of Arkansas.
Fayetteville, Arkansas.
Taylor, R.J. & C.E. Taylor. 1989. An Annotated List of the Ferns, Fern Allies,
Gymnosperms and Flowering Plants of Oklahoma. Southeastern Oklahoma State
University, Durant, Oklahoma.
Texas Organization for Endangered Species. 1993. Endangered, Threatened, and
Watch Lists of Texas Plants. Publication 9. Austin, Texas.
Texas Parks and Wildlife Department. 1997. Special Species List for Angelina,
Jasper, Newton, San Augustine, and Shelby Counties. Austin, Texas.
Thomas, R.D. & C.M. Allen. 1998. Atlas of the Vascular Flora of Louisiana. Vol.
III. Louisiana Department of Wildlife and Fisheries. Baton Rouge, Louisiana.
USDA Forest Service. 1996a. Final Environmental Impact Statement (FEIS) for the
Revised Land and Resource Management Plan, National Forests & Grasslands in
Texas. Tyler, Texas.
USDA Forest Service. 1996b. Final Land and Resource Management Plan. Tyler,
Texas.
Phytologia (February 1998) 84(2):69-86.
CLASSIFICATION OF THE GENUS GI/LIA (POLEMONIACEAE)
Verne Grant
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
The body of knowledge concerning taxonomic characters in Gilia has
increased greatly since the last previous infrageneric classification in 1959.
The older and the newer information is combined to construct a new
infrageneric classification, in which the species are grouped into three
subgenera and seven sections. Five formerly poorly known species which
have traditionally been in Gilia are excluded from the genus. Three of these
are transferred to Tintinabulum, one to Allophyllum, and one to a new genus,
Maculigilia. The very different approaches to classification of molecular
cladistics and evolutionary systematics are currently being applied to Gilia and
related genera; the two approaches are compared and discussed.
KEY WORDS: Gilia, Allophyllum, Maculigilia, Polemoniaceae, polyphyly
It was acommon practice in the nineteenth century to group the temperate species
of Polemoniaceae into four genera: Polemonium, Phlox, Collomia, and Gilia (Meisner
1836-1843; Bentham & Hooker 1876; Gray 1878; Peter 1897). Polemonium and
Phlox have distinctive vegetative and floral characters which define them as genera,
and consequently they have had a relatively stable taxonomic history. To a lesser
extent this is also true of Collomia. Gilia, on the other hand, lacks distinctive
vegetative and floral features, making generic definition difficult. It became a
convenient catchall in the nineteenth century for species that did not fit into the other
genera. Gray (1878), for example, adopted a very broad concept of Gilia which
embraced twelve sections differing greatly from one another; and the treatment of Gilia
by Bentham & hooker (1876) was very similar.
The trend toward all-inclusiveness in Gilia was reversed in the twentieth century as
the plants became better known and more characters were found. The reverse trend
was to segregate subgroups out of the old Gilia as separate genera. This process was
started by Milliken (1904), a student of Jepson, and has continued to the present time.
The net result is that the genus Gilia has been undergoing many and substantial
changes in its constitution throughout its 200-year taxonomic history, unlike
Polemonium, Phlox, and Collomia.
69
70 PHYTOLOGIA February 1998 volume 84(2):69-86
The genus problem in Gilia has two complementary aspects: delimitation of the
genus as a whole, and recognition of natural sections and subgenera. The objectives
are to identify the natural sections or subgenera; to group together those that belong to
the same major monophyletic branch; to exclude subgroups that belong to other
phylogenetic branches; and to use these inclusions and exclusions to determine the
range of variation of the genus.
Three general classification systems of Gilia were proposed in the 1940s and
1950s: those of Mason & A. Grant (1948), Grant & Grant (1956), and Grant (1959).
They all attempted to reflect the natural relationships of the subgroups as understood at
the time. The 1959 system, which is still widely used, recognized five sections:
Giliastrum, Giliandra, Gilia, Saltugilia, and Arachnion. It is in need of revision. The
1948 system contains some elements which were passed over in 1959, but will be
taken up in the present revision.
Much research has been done on Gilia by many workers since 1959. The work
has been carried out on several fronts. Our knowledge of morphological characters
and chromosome numbers has increased. Non-traditional characters such as pollen
morphology, flavonoids, chloroplast DNA, and ribosomal DNA have been introduced
(Stuchlik 1967a, 1967b; Taylor & Levin 1975; Smith et al. 1977; Johnson et al. 1996;
Porter 1977; Day, unpubl.). A sixth section of Gilia, section Kelloggia, has been set
up (Day 1993a, 1993b). Two sections of the 1959 system, Giliandra and Giliastrum
have been segregated as the genera Aliciella and Giliastrum, respectively (Porter
1998a, 1998b).
It is time to restudy and revise the system of classification of Gilia in the light of
the new as well as the old evidence. This task is attempted here.
INTRODUCTION TO THE CLASSIFICATION
A survey of taxonomic characters was carried out in search for relatively
conservative characters that identify groups of related species and distinguish such
groups from one another. The diagnostically useful gross morphological characters
and basic chromosome numbers are emphasized in the following descriptions of
groups. The newer types of characters in pollen, flavonoids, and DNA are presented
following the conventional characters.
A useful pollen character is the distribution of apertures. Two modes occur in
Gilia and its close relatives: pantoporate (apertures scattered over grain) and
zonocolporate (apertures in equatorial zone) (Stuchlik 1967a, 1967b; Taylor & Levin
1975; Chuang et al. 1978; Day, unpubl.).
Three main groups of flavonoids are found in Gilia, and these are designated as
types A, B, and C (Smith et al. 1977). Type A consists of common flavonols
(kaempferol, quercitin, myricetin); type B is 6-methoxyflavonols (patuletin, eupalitin,
eer type C is C-glycosylflavones (apigenin-based and luteolin-based) (Smith
etal. Ty
Grant: Classification of Gilia 7/5
Variation in DNA sequences in a region of a chloroplast gene matK in Gilia and
related genera is plotted in a cladogram (Johnson et al. 1996). Sequence variation in
nuclear ribosomal DNA ITS is plotted on another cladogram (Porter 1997).
Variation in different types of characters is sometimes congruent and sometimes
not. In the latter case, taxonomic judgment and weighting come into play. The
observed distribution of characters, and my interpretation of them in cases of
incongruence, are expressed in the following classification system. This system
groups the species into three subgenera and seven sections. Five species are excluded
from Gilia and transferred to other genera.
Only the essential nomenclature is given here. Additional synonyms for
infrageneric taxa can be found in Grant (1959).
SYSTEM OF CLASSIFICATION
GILIA Ruiz & Pavon, Prodr. Fl. Peruv., 25 andt. 4. 1794. TYPE SPECIES: Gilia
laciniata Ruiz & Pavon.
Plants herbaceous; perennial, biennial or annual, sometimes with a woody or soft-
woody base. Herbage with several types of pubescence, but not with dense coarse
woolly hairs. Leaves alternate, with irregular pinnate teeth, lobes, or divisions, or
entire-margined and linear in reduced forms. Usually spring blooming. Corolla
mostly funnelform, sometimes campanulate, rotate, or salverform. Seeds angular,
mostly sandy-colored, mostly mucilaginous when wet, but not mucilaginous in two
sections. Chromosomes large, basic number x=9 in most sections, x=8 in sect.
Giliandra.
Pollen zonocolporate, except in subg. Kelloggia where pantoporate. Flavonoids
of type A and C widespread in genus; type B flavonoids found only in sections
Giliastrum and Gilmania so far as known. (See preceding section for explanation of
these terms.).
Widespread in western North America, also in temperate South America.
73 species as presently understood. These are grouped here in three subgenera
and seven sections.
For the relationships of Gilia to the Eriastrum-Ipomopsis-Langloisia group and
Leptodactylon-Linanthus group see Grant (1998). The relationships between Gilia,
and Tintinabulum and Macugilia are considered later in the present paper.
{7 PHY TOLGGLA February 1998 volume 84(2):69-86
Key to the Main Subgroups of Gilia
A. Perennials or biennials, with woody base or herbaceous throughout.
B. Corolla campanulate or rotate. Pollen blue. Seeds mucilaginous when wet. ...
x chiara» Adtawepateineets qeapie sb aieaiek ndlle cons trantateate tas mee sect. Giliastrum, in part
BB. Corolla’ funnelform, or occasionally salverform. Pollen yellow or white
(except blue in 1 species of sect. Giliandra). Seeds not mucilaginous when
wet.
C. Plants scapose. Basal leaves pinnately lobed or divided, with strap-shaped
TACHIS, XEG. cc eccne ones ccna s<> os esnstesacnes ea tee enna sect. Giliandra, in part
CC. Plants not scapose. Leaves with broad blade. x=9. Rare.................+
nes daieromdin tations nie Sjasettnele asda ante opine taut eee sect. Gilmania, in part
AA. Annual herbs.
iD: Corolla campanulate.......... oc .c.0..065-nnenane G. incisa group in sect. Giliastrum
DD. Corolla funnelform, or occasionally salverform.
E. Seeds not mucilaginous when wet. Pollen yellow or white.
F. Plants scapose. Basal leaves pinnately lobed or divided, with strap-
shaped rachis. x=9 or 8.......... G. leptomeria group in sect. Giliandra
FF. Plants not scapose. Leaves with broad blade. x=9. Uncommon......
AEURE Da cepad cind diese naencousmehts Sie aeeRete G. latifolia in sect. Gilmania
EE. Seeds mucilaginous when wet. Pollen blue.
G. Leaves with 1 or 2 linear finger-like lobes, or reduced and unlobed.
Pollen pantoporate. Middle and high elevations in mountains. ..........
siosiovineuen ptanwiaie «damienseis apt aa eptaaob selene tata subg. Kelloggia
GG. Leaves dissected but not with lobes as in G. Pollen zonocolporate.
Mainly at lower elevations, uncommon in middle elevations, and rarely
at high elevations... ....--pe9ces swenssiccunaieni Senta aaa ene subg. Gilia
H. Pubescence of interwoven fine cobwebby hairs. Basal leaf
TOSEELE PIeSENE «on a.nhioe -nrscmeesedpesare mien ees sect. Arachnion
HH. Pubescence of straight multicellular hairs and stipitate glands.
Basal leaf rosette present or absent.
I. Upper leaves well developed. No basal leaf rosette.
Inflorescence usually a head or cluster..............+.+. sect. Gilia
II. Upper leaves much reduced. Basal leaf rosette present.
Inflorescence an.Open CYME........<<%- «0 ise panes sect. Saltugilia
I. GILIA subg. GREENEOPHILA Brand, Pflanzenreich 250:144. 1907. TYPE
SPECIES: Gilia rigidula Bentham.
Perennials, sometimes with woody or soft-woody base, biennials, and annuals.
Leaves with once pinnate teeth, lobes, or divisions. Corolla often concolored,
sometimes bicolored. Pollen yellow, white, or blue. Seeds mucilaginous when wet
or not so. Basic chromosome number x=9 and x=8.
Pollen zonocolporate. Flavonoids of types A, B, and C occur in sects. Giliastrum
and Gilmania; no information on flavonoids in sect. Giliandra. Species representing
the sections of this subgenus fall close together in the cladograms for cpDNA and
mDNA. They are remote from species of subg. Gilia on the same cladograms.
Grant: Classification of Gilia 73
Center of distribution of the perennial and biennial members in northern México,
Rocky Mountains, and intermountain region. Annuals widespread in western deserts.
One section occurs also in temperate South America.
Three sections, sects. 1-3 as follows.
1. GILIA section GILIASTRUM Brand, Pflanzenreich 250:147. 1907. TYPE
SPECIES: Gilia rigidula Bentham.
Giliastrum Rydberg, Fl. Rocky Mts., ed. 1, 699 and 1066. 1917.
Perennials with soft-woody base, and annuals. Lower leaves with a broad blade
and once pinnate lobes, the lobes often sharp-tipped, or leaves linear. Corolla
campanulate or rotate. Corolla concolored, violet or whitish, or bicolored, violet with
a yellow center. Pollen blue in some species, no data for other species. Seeds sandy
colored, mucilaginous when wet. x=9, n=6 found in one species.
Type B flavonoid is present.
Center of distribution in northern México and Texas. Secondary centers in Baja
California and temperate South America.
Gilia incisa Benth., G. insignis (Brand) Cory & H.B. Parks, G. foetida Gill. ex
Benth. (S. Amer.), G. gypsophila B.L. Turner, G. ludens Shinners, G. purpusii K.
Brandegee, G. rigidula, and G. stewartii I.M. Johnst. This species list is based on
Turner (1994).
Porter (1998b) is segregating section Giliastrum as a genus separate from Gilia to
eliminate polyphyly. Grouping Giliastrum together with subgenus Gilia results in a
polyphyletic genus according to Porter’s phylogenetic hypothesis based on nrDNA.
According to my phylogenetic hypothesis based on all available characters, section
Giliastrum and subgenus Gilia have diverged widely, but nevertheless from a
monophyletic taxon. See the last two sections of this paper. Separation of Giliastrum
from subgenus Gilia is thus deemed unnecessary from a phylogenetic standpoint, and
in addition, is undesirable taxonomically since it obscures the relationship between the
two phyletic branches.
2. GILIA section GILIANDRA A. Gray, Proc. Amer. Acad. Arts 8:276. 1870.
TYPE SPECIES: Gilia stenothyrsa A. Gray. Aliciella sect. Giliandra (A. Gray)
J.M. Porter, Aliso 17:27. 1998.
Aliciella Brand, Pflanzenreich 250:150. 1907. TYPE SPECIES: Gilia triodon
Eastwood.
Aliciella sect. Aliciella subsect. Subnudae J.M. Porter, Aliso 17:31. 1998. TYPE
SPECIES: Gilia subnuda Torrey ex A. Gray.
Perennial herbs, sometimes woody-based, biennial herbs, and annuals. Plants
with a basal leaf rosette and scapose upper parts. Lower leaves leathery, with a strap-
74 PHYTOLOGIA February 1998 volume 84(2):69-86
shaped rachis and once pinnate lobes or divisions. Corollas mostly funnelform,
sometimes salverform, tubes slender or broad. Corollas showy, concolored, blue,
red, or pink; or small and inconspicuous in most annual species. Pollen yellow or
white, except blue in one species (Gilia tenuis). Seeds sandy colored, not
mucilaginous when wet. x=8 throughout the section, x=9 occurs also in some annual
species.
The species of this section fall into two subgroups. The perennial and biennial
species with showy flowers and x=8 are basal. They occur in Utah, Colorado, and
adjacent areas. The second subgroup consists of annuals, mostly with small flowers,
they have both x=8 and 9 and polyploidy. They are derived and occur in the western
deserts.
The center of distribution of the showy-flowered subgroup is in the Rocky
Mountains and Colorado Plateau. The annuals range widely in the western deserts as
noted above.
Showy-flowered perennial species: Gilia caespitosa A. Gray, G. formosa E.
Greene ex Brand, G. haydeni A. Gray, G. mcevickerae M.E. Jones, G.
pentstemonoides MLE. Jones, G. pinnatifda Moc. & Sessé, G. sedifolia Brandegee,
G. stenothyrsa A. Gray, G. subnuda, and G. tenuis F.J. Sm. & Neese. Annual,
mostly small-flowered species: G. heterostyla S.A. Cochrane & A.G. Day, G.
humillima, G. hutchinsifolia Rydb., G. leptomeria A. Gray, G. lottiae A.G. Day, G.
micromeria A. Gray, G. nyensis Reveal, G. subacaulis Rydb., and G. triodon Eastw.
The list of perennial species is based on Porter (1998a), that of the annual species on
Day (unpubl.).
Porter (1998a) has recently segregated Giliandra and Gilmania as a separate genus,
Aliciella, in order to achieve monophyly. A genus containing Giliandra, Gilmania,
and subgenus Gilia is polyphyletic according to Porter’s phylogenetic hypothesis
based on nDNA. However, this combination is diverse, but monophyletic according
to my phylogenetic hypothesis based on all available characters. See discussion in the
last two sections of this paper. Separation of Aliciella from Gilia is deemed
unnecessary phylogenetically and undesirable taxonomically for the same reasons as
those given above with regard to Giliastrum.
3. GILIA section GILMANIA (Mason & A. Grant) V. & A. Grant, Aliso 3:299.
1956. TYPE SPECIES: Gilia latifolia S. Watson. Gilia subg. Gilmania Mason &
A. Grant, Madrofio 9:205. 1948. Aliciella subg. Gilmania (Mason & A. Grant)
J.M. Porter, Aliso 17:43. 1998.
One species a woody-based perennial, another species an annual. Lower leaves
with a broad blade and lobed margins, the lobes sharp-tipped. Corolla funnelform,
pink. Pollen yellow. Seeds reddish-brown, not mucilaginous when wet. x=9.
Type B flavonoids present.
Deserts from southerm California to Utah.
Gilia latifolia and G. ripleyi Barneby.
Grant: Classification of Gilia 75
Mason & A. Grant (1948) set up a subgenus, Gilmania, for this distinctive small
group; and Grant & Grant (1956) retained it in the rank of section. I later sank
Gilmania into section Giliastrum (Grant 1959); this was a step in the wrong direction.
Porter (1998a) has recently grouped sect. Gilmania with sect. Giliandra in the
segregate genus Aliciella. I regard this as a step in another wrong direction. Gilmania
does not fit well in either section, on conventional taxonomic characters, and is best
treated as a third section coordinate with the other two.
II. GILIA subg. GILIA
Annual herbs. Leaves divided once pinnately to tripinnately, or entire margined
and linear in reduced forms. Corolla mostly funnelform, sometimes salverform.
Corolla concolored, or bicolored or tricolored with spots or UES Pollen blue. Seeds
mucilaginous when wet. x=9.
Pollen zonocolporate. Flavonoids of type B absent so far as known. Species of
the three sections in this subgenus fall close together on the cladograms for cpDNA
and mDNA. They are remote from the species of subg. Greeneophila on the same
cladograms.
Center of distribution in California, occurring in regions with summer-dry climates
and in deserts. Ranging through other parts of western North America, and recurring
in temperate South America.
Three sections, nos. 4-6 as follows.
4. GILIA section GILIA
Annual herbs with well-developed upper leaves. Pubescence of straight
multicellular hairs and stipitate glands. Inflorescence usually a head or cluster.
Corollas funnelform, concolored or tricolored.
Center of distribution in cismontane California. Ranging north to British
Columbia; recurring in temperate South America.
Gilia achilleaefolia Benth., G. angelensis V. Grant, G. capitata Sims, G. clivorum
(Jepson) V. Grant, G. laciniata Ruiz & Pavén (S. Amer.), G. lomensis V. Grant (S.
Amer.), G. millefoliata Fischer & C. Meyer, G. nevinii A. Gray, G. tricolor Benth.,
and G. valdiviensis Griseb. (S. Amer.).
5. GILIA section SALTUGILIA V. & A. Grant, Aliso 3:84. 1954. TYPE
SPECIES: Gilia splendens Douglas.
Annual herbs with well-developed basal leaves and reduced upper leaves.
Pubescence of straight multicellular hairs ana stipitate glands. Inflorescence cymose.
Corollas funnelform, concolored.
76 PHYTOLOGIA _ February 1998 volume 84(2):69-86
Center of distribution in southern California. Ranging to central California
mountains and to southwestern deserts.
Gilia australis (H. Mason & A.D. Grant) V. Grant & A.D. Grant, G. caruifolia
Abrams, G. scopulorum MLE. Jones, G. splendens H. Mason & A.D. Grant, G.
stellata A.A. Heller, and G. yorkii ined. Gilia yorkii is a new species (Shevock &
Day, in press).
6. GILIA section ARACHNION A. & V. Grant, Aliso 3:214. 1956. TYPE
SPECIES: Gilia latiflora A. Gray.
Annual herbs with a basal leaf rosette and scapose upper body. Pubescence of fine
cobwebby hairs especially in leaf axils. Inflorescence cymose. Corollas mostly
funnelform, sometimes salverform. Corollas usually tricolored.
Center of distribution in southern California mountains and Mojave Desert.
Ranging to other areas of western North America; recurring in temperate South
America.
Gilia aliquanta A.D. Grant & V. Grant, G. austro-occidentalis (A.D. Grant & V.
Grant) A.D. Grant & V. Grant, G. brecciarun MLE. Jones, G. cana (M.E. Jones)
A.A. Heller, G. clokeyi H. Mason, G. crassifolia Benth. (S. Amer.), G. diegensis
(Munz) A.D. Grant & V. Grant, G. flavocincta A. Nels., G. inconspicua (Smith)
Sweet, G. interior (H. Mason & A.D. Grant) A.D. Grant, G. jacens A.D. Grant & V.
Grant, G. latiflora (A. Gray) A. Gray, G. leptantha Parish, G. malior A.G. Day & V.
Grant, G. mexicana A.D. Grant & V. Grant, G. minor A.D. Grant & V. Grant, G.
modocensis Eastw., G. ochroleuca M.E. Jones, G. ophthalmoides Brand, G. sinuata
Douglas ex Benth., G. salticola Eastw., G. tenuiflora Benth., G. tetrabreccia A.D.
Grant & V. Grant, G. transmontana (H. Mason & A.D. Grant) A.D. Grant & V.
Grant, and G. tweedyi Rydb.
III. GILIA subg. KELLOGGIA Mason & A. Grant, Madrofio 9:219. 1948. TYPE
SPECIES: Gilia capillaris Kellogg.
Medium-sized to small annual herbs. Herbage glandular-puberulent, or glabrous.
Upper and lower leaves about the same size. Leaves with 1 or 2 linear finger-like
lobes, or unlobed and entire. Corolla funnelform, concolored or bicolored. Pollen
blue, pantoporate, the sexine striated or with spinules. Capsule containing 1-6 seeds
per locule. Seeds mucilaginous when wet. x=9.
No information on flavonoids, and only one puzzling record of DNA sequences.
Center of distribution in mountains of central California, Nevada, and Oregon.
Disjunct populations in Idaho and Colorado.
Grant: Classification of Gilia Wi
7. GILIA section KELLOGGIA Day, Novon 3:332. 1993.
Characters of the subgenus.
Gilia capillaris Kellogg, G. leptalea (A. Gray) E. Greene, and G. sinistra M.E.
Jones.
The species of subgenus Kelloggia are similar in habit, and floral and seed
characters to those of subgenus Gilia, and occur in the same geographical area. For
these reasons the species assigned here to the subgenera Kelloggia and Gilia were
formerly thought to be closely related.
However, the discovery of pantoporate pollen in the species of Kelloggia (Day,
unpubl. data) indicates that it is isolated from the subgenera Gilia and Greeneophila,
which have zonocolporate pollen (Stuchlik 1967a, 1967b; Day, unpubl. data).
What are the true relationships of Kelloggia? Pollen characters and leaf form point
to a relationship with Allophyllum. But more evidence is needed to determine the best
taxonomic disposition of Kelloggia.
DNA evidence might shed light on the problem. The only DNA record of the
section published so far is for cp DNA matK in Gilia sinistra (Johnson et al. 1966). It
places G. sinistra in a group of five Navarrettia species in the cladogram. This result
is out of line with the phenetic evidence concerning both Kelloggia and Navarrettia.
Further molecular studies are needed.
EXCLUSION AND REASSIGNMENT OF FIVE SPECIES
Five species of annuals with reduced vegetative and floral characters, assigned to
Gilia, have been poorly understood throughout this century. The species in question
are G. campanulata A. Gray, G. inyoensis I.M. Johnst., G. filiformis Parry ex A.
Gray, G. maculata Parish, and G. tenerrima A. Gray. The first four are desert annuals
and the last one is montane. The relationships of these species to the main subgroups
of Gilia have not been obvious from the usual taxonomic characters, leaving it up to
successive workers to place them as best they can in the system. Much more has
become known about these plants in the last ten years, however, and it is time for a
reassessment.
The five species do not fit well into any of the seven sections recognized in the
present treatment. If they were to remain in Gilia, additional sections would have to be
set up for them. This is a plan that I considered and tried to implement but in the end
discarded. The species are not only misfits in Gilia, but show signs of relationship to
other genera. Therefore, it seems best to take them out of Gilia and place them in other
genera. I am resurrecting the long dormant genus Tintinabulum of Rydberg (1917) for
G. campanulata, G. inyoensis, and G. filiformis. A new genus Maculigilia is set up
for G. maculata; and G. tenerrima is transferred to Allophyllum.
78 PHYTOLOGIA _ February 1998 volume 84(2):69-86
TINTINABULUM Rydberg, Fl. Rocky Mts., ed. 1, 698 and 1065. 1917. TYPE
SPECIES: Gilia filiformis Parry ex A. Gray. Gilia subg. Tintinabulum Mason &
Grant, Madrofio 9:220. 1948.
Gilia subg. Greeneophila sect. Campanulastrum Brand, Pflanzenreich 250:144.
1907. TYPE SPECIES: Gilia campanulata A. Gray. Gilia subg.
Campanulastrum Mason & A. Grant, Madrofio 9:219. 1948.
Small desert annuals with a spreading habit. Stems very slender. Leaves
alternate, reduced, lanceolate to linear. Flowers solitary on slender pedicels. Corolla
narrowly to broadly campanulate. Corolla yellow or bigolored yellow and white.
Pollen yellow. Seeds mucilaginous when wet. x=9.
Center of distribution northern Mojave Desert, ranging through deserts to Utah and
Arizona. Three species.
Tintinabulum filiforme (Parry ex A. Gray) Rydberg, Fl. Rocky Mts., ed. 1, 698
and 1065. 1917. BASIONYM: Gilia filiformis Parry ex A. Gray, Proc.
Amer. Acad. Arts 10:75. 1874.
Tintinabulum campanulatum (A. Gray) V. Grant, comb. nov. BASIONYM:
Gilia campanulata A. Gray, Proc. Amer. Acad. Arts 9:279. 1870.
Tintinabulum inyoensis (1.M. Johnston) V. Grant, comb. nov. BASIONYM:
Gilia inyoensis 1.M. Johnston, Contrib. Gray Herb. 75:39. 1925.
These three species differ from the rest of Gilia, or from Gilia as delimited here, in
spreading habit with very slender stems, and differ from most sections of Gilia in
campanulate corollas. In my old treatment (Grant 1959) I included them in Gilia sect.
Giliastrum on what now seem to be superficial resemblances. Mason & A. Grant
(1948) had previously assigned them to two adjacent small subgenera of Gilia, subg.
Campanulastrum and subg. Tintinabulum, which was on the right track. They could
be retained in Gilia as a third subgenus.
However, in growth habit and floral characters, the plants resemble Linanthus
sect. Dactylophyllum (the L. aureus [Nutt.] E. Greene group). Furthermore, they
cluster with Leptodactylon and Linanthus in the cladograms for nrDNA (Porter 1997)
and cpDNA (Johnston et al. 1996). *
These indications of a relationship with Linanthus, particularly the DNA evidence,
tilt the scales in favor of segregating the group at the genus level. Rydberg’s (1917)
small genus Tintinabulum is a good place for them. The genus Tintinabulum is
assigned to the tribe Gilieae.
MACULIGILIA V. Grant, genus novum. TYPE SPECIES: Gilia maculata Parish.
Herbae annuae diminutivae, hirsutae. Folia integra, oblongata vel
oblanceolata. flores in capitulis terminalibus. Corolla campanulata, maculata.
Pollen flavus, pantoporatus. Semina rubicunda brunnea, sub aqua immutata.
Chromosomae x=9.
Grant: Classification of Gilia 79
Small annuals with spreading habit. Stems with dense cover of long hairs.
Leaves fleshy, entire, oblong or oblanceolate. Inflorescence a dense cluster. Calyx
lobes free with ciliated membranous margins. Corolla campanulate, corolla lobes
notched at tip. Corolla tricolored with white lobes and throat, yellow tube, and red
spots on lobes. Pollen yellow, pantoporate. Seeds dark reddish brown, not
mucilaginous when wet. x=9.
Colorado Desert, California, rare. One species.
Maculigilia maculata (Parish) V. Grant, comb. nov. BASIONYM: Gilia
maculata Parish, Bull. Torrey Bot. Club 19:93. 1892. Linanthus maculatus
(Parish) Milliken, Univ. California Publ. Bot. 2:55. 1904.
This rare species was described as a Gilia by Parish in 1892, but transferred to
Linanthus by Milliken (1904) and retained there by later students (e.g., Jepson 1943;
Grant 1959). Patterson (1989) rediscovered the species in the wild and made a
thorough study of it. The above diagnosis is based on Patterson’s more detailed
description. Patterson concluded that the species does not belong in Linanthus, but
can be accommodated in Gilia though it is unique there.
Species maculata differs from the present sections of Gilia in its calyx,
pubescence, and leaf blades, and from all sections except Kelloggia in having
pantoporate pollen.
The species does, however, share some distinctive characters in common with
section Gilmania. Namely, seed characters, pollen color, and desert ecology. These
characters suggest a relationship with section Gilmania.
But the molecular evidence points in a different direction. In the cladogram for
nrDNA ITS (Porter 1997), Maculigilia maculata clusters with Leptodactylon and
Linanthus. It is also close to the Tintinabulum campanulatum group which is close to
Leptodactylon and Linanthus in the cladogram.
Nevertheless, Maculigilia maculata is well outside the range of variation of these
genera. Therefore, it is segregated into a monotypic genus of its own which seems to
lie somewhere between Linanthus and Gilia. Maculigilia is assigned tentatively to the
tribe Leptodactyloneae.
Allophyllum tenerrimum (A. Gray) V. Grant, comb. nov. BASIONYM: Gilia
tenerrima A. Gray, Proc. Amer. Acad. Arts 8:277. 1870. Navarrettia tenerrima
Kuntze, Revisio Gen. Pl. 2:433. 1891.
Small annual herb with spreading habit. Leaves, flowers, and capsules much
reduced. Pubescence stipitate-glandular with a slender stalk bearing a black gland.
Leaves alternate, oblanceolate to linear with 1 or 2 lobes, or not lobed and entire.
Flowers solitary, small. Corolla tube and throat white and lobes pale blue. Pollen
grains approaching pantoporate, with fine spinules on the tectum. Capsule globular,
containing usually 1 seed per locule, valves usually falling off at maturity. Seeds
rounded, ovoid, brown; mucilaginous when wet. x=9; 2n=36.
80 PHYTOLOGIA _ February 1998 volume 84(2):69-86
Mid and high elevations in mountains, eastern Oregon and Sierra Nevada of
California to Wyoming and Utah.
This species has been included in Gilia in almost all treatments since Gray (1878).
In some modern treatments it has been placed close to the Gilia leptalea group which is
now Gilia sect. Kelloggia (Mason & A. Grant 1948, 1951; Grant 1959). It is,
however, quite distinct within Gilia.
Stuchlik (1967a, 1967b) and Day (unpubl.) noted that species tenerrimum is
similar to Allophyllum in pollen morphology. Day (unpubl.) noted other similarities to
Allophyllum in pubescence and gross morphology. The trichomes, leaf form,
capsules, and seeds of species tenerrimum are characteristic of Allophyllum but not of
Gilia. This evidence supports the conclusion that the species belongs to Allophyllum,
a conclusion with which Day agrees (pers. comm.). Allophyllum tenerrimum is amply
different from the typical Allophyllum -- A. gilioides (Benth.) A.D. Grant & V. Grant,
A. glutinosum, etc. -- but many of the differences can be attributed to the reduced
nature of A. tenerrimum.
In the DNA cladograms (Johnson et al. 1996; Porter 1997), Allophyllum
tenerrimum occurs close to both Allophyllum and Gilia subg. Gilia.
PHYLOGENY
The characters and character combinations used to delineate the subgenera and
sections of Gilia can be used to infer a probable phylogeny. The two subgenera,
Greeneophila and Gilia, represent two main branches in the inferred phylogenetic tree
(Figure 1). Of these two, the subgenus Greeneophila contains the most primitive
characters.
The Gilia rigidula group (in subg. Greeneophila sect. Giliastrum) exhibits the
primitive life-form or woody-based perennials. It approaches the most primitive genus
Loeselia in life-form and is similar to it in sequences of cpDNA and mDNA (Johnson
et al. 1996; Porter 1997). The G. rigidula group occurs with Loeselia in northern
México and Texas, and is probably derived from a Loeselia-like ancestor in this area
(Grant 1959, 1998).
The perennial members of the other sections of subgenus Greeneophila exhibit
some derived features as compared with section Giliastrum, and appear to be side-
branches derived from a Gilia rigidula-like ancestor. The perennial species of section
Giliandra have colonized the Rocky Mountains and Colorado Plateau. The centers of
distribution of the perennial members of subgenus Greeneophila thus lie in a region
from northern México to the Rocky Mountains and Colorado Plateau.
All three sections of subgenus Greeneophila have given rise to reduced annuals
which have colonized western deserts.
Grant: Classification of Gilia 81
The subgenus Gilia with its three interrelated sections (Gilia, Saltugilia, and
Arachnion) consists entirely of annuals. The showy-flowered species are the basal
members of this subgenus. These have their center of distribution and probable center
of origin in California. The derived small-flowered members are widespread in the
western deserts and mountains where they intermingle with the reduced annuals of
subgenus Greeneophila.
Subgenus Gilia is united with subgenus Greeneophila by some common
characters, but is also amply different from Greeneophila in morphology and center of
distribution. It is also far removed from the subgenus Greeneophila in the cladograms
for cpDNA matK and nrDNA ITS (Johnson et al. 1996; Porter 1997).
It is suggested that the three sections of subgenus Gilia branched off from an
ancestor in or near the G. rigidula group in California in response to climatic changes
toward summer-dry conditions. Such climatic changes occurred in the Middle
Pliocene, Late Pliocene, and Xerothermic phase of the Quaternary (Axelrod 1948,
1950; Raven & Axelrod 1978). The hypothesis presumes that the ancestor
disappeared from the California area as the climate became unfavorable for it, creating
the present systematic gap. Sections Saltugilia and Arachnion then went on to radiate
in the deserts as desert areas expanded in the Late Pliocene and Xerothermic time.
An origin of subgenus Gilia from the perennial G. rigidula group presents no
particular theoretical difficulties. The G. rigidula group has given rise to the annual G.
incisa group in Texas. There is not much difference between G. incisa and reduced
plain-flowered species of subgenus Gilia, such as G. angelensis and G. australis.
The third subgenus Kelloggia does pose a problem. On general morphological
characters, it has been allied to the other far-western annual gilias. However, it has
pantoporate pollen (Day, unpubl. data). This is a primitive condition in the family,
and occurs in Loeselia among other genera, whereas the rest of Gilia including the G.
rigidula group has the derived condition of zonocolporate pollen (Stuchlik 1967a,
1967b; Taylor & Levin 1975; Day, unpubl. data).
This suggests that subgenus Kelloggia is not as closely related to the western
annual gilias as has been thought, is not derived from the Gilia rigidula group, and
may have an independent origin with some unknown ancestor with pantoporate pollen
(Figure 1). If this suggestion is confirmed, Kelloggia does not belong in Gilia.
A number of cases exist in Gilia and its relatives where a given group exhibits
evidence of relationships in two different directions. Some characters of the group in
question, group X, indicate a relationship with group A, other characters of group X
relate it to group B, and A and B are too distantly related to be able to hybridize.
Subgenus Kelloggia is one example of this. Maculigilia is another; one set of
characters suggests a relationship with Gilia section Gilmania, while another set
indicates a relationship with Linanthus. Other such cases are noted in the preceding
sections.
What are the phylogenetic explanations of the apparently bipolar relationships?
One explanation is convergence which is common and widespread. Convergent
evolution has produced desert annuals with reduced vegetative characters and small
flowers in four sections (Giliastrum, Giliandra, Saltugilia, Arachnion), the similar-
82 PHY TOLOGGIA February 1998 volume 84(2):69-86
appearing species were grouped together in the same artificial section Eugilia of older
authors. Another source of bipolar relationships may be the retention of disparate
elements in the genotype from ancient hybridization events; this is a possibility that we
know very little about. We should also consider horizontal gene transfer between
remotely related groups, which may be more common in nature than we realize.
DISCUSSION
Until recently all systematic studies of Gilia and other Polemoniaceae were carried
out within the framework of either traditional taxonomy or evolutionary systematics
(cf. Grant 1998). Studies of DNA sequence variation in Polemoniaceae including
Gilia began in the 1990s, and the DNA evidence has been analyzed and interpreted
according to the procedures of cladistics (Steele & Vilgalys 1994; Johnson et al. 1996;
Porter 1997, 1998a).
The philosophy and methods of traditional taxonomy and_ evolutionary
systematics, on the one hand, and those of molecular cladistics on the other, are very
different, and lead to different conclusions in some cases. I have discussed the
differences as regards the family Polemoniaceae elsewhere (Grant 1998), and will
discuss the differences with respect to Gilia here.
Johnson et al. (1996) and Porter (1997, 1998a) state emphatically that Gilia is
polyphyletic; indeed, “extremely” polyphyletic (Johnson ef al. 1996). Actually, the
phrase “extreme polyphyly” applies better to the historical catchall genus Gilia of the
nineteenth century than it does to the Gilia of recent times. Successive generations of
botanists have labored throughout the twentieth century to make Gilia more natural or
less polyphyletic, and much progress has been made.
What is the standard of reference for determining the polyphyly or monophyly of a
taxonomic group? For Johnson et al. (1996) and Porter (1997, 1998a, 1998b) there is
only one standard. It is the cladogram or gene tree of a particular DNA segment.
Johnson et al. (1996) and Porter (1997) go directly from the clades in their DNA
cladograms to informal taxonomic groups, as I have shown in my analysis of their
work (Grant 1998). Porter 1998a, 1998b) takes it a step further by converting several
of the informal taxa into formal ones. Porter’s (1998a) subdivisions of Aliciella
conform closely to the set of clades and subclades in his nrDNA cladogram (Porter
1997). Gilia species occur in different clades of the DNA cladograms, and it is on this
basis that Gilia is said to be extremely polyphyletic (Johnson et al. 1996; Porter 1997,
1998a).
There is another standard of reference for monophyly/polyphyly, namely, the
system of classification built up by the methods of traditional taxonomy and
evolutionary systematics. These methods include consideration of all possible
characters and weighting of characters in cases of conflict in the evidence. They are
responsible for almost al! of the progress that has been made toward a phylogenetically
natural classification of Gilia and allied genera.
Grant: Classification of Gilia 83
Subg. Subg. Subg.
Greeneophila Gilia Kelloggia
S a &
oo an Sis
Gig es Gugve
Loeselia
Figure 1. Proposed phylogeny of the main
subgroups of the genus Gilia.
84 PHYTOLOGIA _ February 1998 volume 84(2):69-86
In short, all parties can agree that some polyphyly has persisted in Gilia and its
allies into the 1990s. But disagreement exists between the molecular cladists and this
evolutionary systematist as to the extent of the polyphyly, and more fundamentally, as
to whether it can be detected solely from the evidence of one or two genes.
The relationship between DNA evidence and conventional characters in Gilia and
allies falls into three patterns. There are many areas of congruence between the DNA
cladograms of the 1990s and the pre-1990 classification system of Gilia and allied
genera. Here the molecular evidence and traditional evidence are mutually supportive.
There are cases where the DNA evidence calls attention to an anomaly in a sector of the
preexisting system. Reexamination of the anomalous group reveals a previously
unknown congruence between molecular and traditional characters, and leads to a
taxonomic change. Finally, we have cases of incongruence between the DNA
cladograms and preexisting classifications. These force a choice between the
molecular and traditional evidence.
Let us try to place the molecular evidence in some kind of perspective. Natural
relationship is measured by the degree of similarity in the genotypes of two or more
individuals or groups. A single gene or region of a gene represents an infinitesimally
small fraction of the genotype of a higher plant. Furthermore, the DNA of a cell
organelle such as a chloroplast is less central to the genome than a major gene in a
chromosome, and the latter type of genetic element is not being tested in plant
molecular systematics. By contrast, the morphological characters and ecological
preferences of plants represent the expression of scores or hundreds of chromosomal
genes (see Clausen & Heisey 1958; Grant 1975). In any incongruence between the
evidence from one or two genes and that from multifactorial phenotypic characters, the
latter must be given great weight. '
Molecular cladistics is designed for generating cladograms and phylogenies of
genes. It is not designed for constructing classifications. For this we have the time-
tested approaches of traditional taxonomy and evolutionary systematics.
ACKNOWLEDGMENTS
Alva Day furnished valuable unpublished information about several groups of
Gilia and Allophyllum as noted in the text. The manuscript was read by Billie L.
Turner and Karen A. Grant, both of whom made helpful comments. I express my
appreciation to these individuals for their help. The herbarium studies were carried out
at the University of Texas Herbarium.
LITERATURE CITED
Axelrod, D.I. 1948. Climate and evolution in western North America during Middle
Pliocene time. Evolution 2:127-144.
Grant: Classification of Gilia 85
Axelrod, D.I. 1950. Evolution of desert vegetation in western North America.
Carnegie Institute of Washington Publ. no. 590. Washington, D.C.
Bentham, G. & J.D. Hooker. 1876. Polemoniaceae. Genera Plantarum. 2:820-824.
Covent Garden, London, Great Britain.
Chuang, T.I., W.C. Hsieh, & D.H. Wilken. 1978. Contribution of pollen
morphology to systematics in Collomia (Polemoniaceae). Amer. J. Bot. 65:450-
458
Clausen, J. & W.M. Heisey. 1958. Experimental studies on the nature of species.
IV. Genetic structure of ecological races. Carnegie Institute of Washington Publ.
no. 615. Washington, D.C.
Day, A.G. 1993a. Gilia. In: J.C. Hickman (ed.), The Jepson Manual: Higher
Plants of California, pp. 828-839. University of California Press, Berkeley,
California.
Day, A.G. 1993b. New taxa and nomenclatural changes in Allophyllum, Gilia, and
Navarretia (Polemoniaceae). Novon 3:331-340.
Grant, V. 1959. Natural History of the Phlox Family. Martinus Nijhoff, The Hague,
The Netherlands.
Grant, V. 1975. Genetics of Flowering Plants. Columbia University Press, New
York, New York.
Grant, V. 1998. Primary classification and phylogeny of the Polemoniaceae, with
comments on molecular cladistics. Amer. J. Bot. 85:741-752.
Gray, A. 1878. Polemoniaceae. Synoptical Flora of North America, \st ed. 2:128-
151. Ivison, Blakeman, & Taylor, New York, New York.
Jepson, W.L. 1943. Polemoniaceae. A Flora of California. 3:131-222. Associated
Students Store, University of California, Berkeley, California.
Johnson, L.A., J.L. Schultz, D.E. Soltis, & P.S. Soltis. 1996. Monophyly and
generic relationships of Polemoniaceae based on matK sequences. Amer. J. Bot.
83:1207-1224.
Mason, H.L. & A. Grant. 1948. Some problems in the genus Gilia. Madrofio
9:201-220
Mason, H.L. & A. Grant. 1951. Gilia. In: L. Abrams (ed.), Illustrated Flora of the
Pacific States. 3:456-474. Stanford University Press, Stanford, California.
Meisner, C.F. 1836-1843. Polemoniaceae. Plantarum Vascularium Genera
Secundum Ordines Naturales, p. 273. Leipzig, Germany.
Milliken, J. 1904. A review of California Polemoniaceae. Univ. California Publ.
Bot. 2:1-71.
Patterson, R. 1989. Taxonomic relationships of Gilia maculata (Polemoniaceae).
Madrono 36:15-27.
Peter, A. 1897. Polemoniaceae. Jn: A. Engler & K. Prantl (eds.), Die Natiirlichen
Pflanzenfamilien. 4(3a):40-54. Wilhelm Engelmann, Leipzig, Germany.
Porter, JM. 1997. Phylogeny of Polemoniaceae based on ribosomal internal
transcribed DNA sequences. Aliso 15:57-77.
Porter, JM. 1998a. Aliciella, a recircumscribed genus of Polemoniaceae. Aliso
17:23-46.
Porter, JM. 1998b. Nomenclatural changes in Polemoniaceae. Aliso 17:83-85.
Raven, P.H. & D.I. Axelrod. 1978. Origin and relationships of the California flora.
Univ. California Publ. Bot. 72:1-134.
Rydberg, P.A. 1917. Flora of the Rocky Mountains and Adjacent Plains, ed. 1.
Noble Printing Co., New York, New York.
Shevock, J.R. & A.G. Day. In press. A new Gilia (Polemoniaceae) from limestone
outcrops in the southern Sierra Nevada of California. Madrono.
86 PHYTOLOGIA _ February 1998 volume 84(2):69-86
Smith, D.M., C.W. Glennie, J.B. Harbourne, & C.A. Williams. 1977. Flavonoid
diversification in the Polemoniaceae. Biochem. Syst. Ecol. 5:107-115.
Steele, K.P. & R. Vilgalys. 1994. Phylogenetic analyses of Polemoniaceae using
nucleotide sequences of the plastid gene matK. Syst. Bot. 19:126-142.
Stuchlik, L. 1967a. Pollen morphology in the Polemoniaceae. Grana Palynologica
7:146-240.
Stuchlik, L. 1967b. Pollen morphology and taxonomy of the family Polemoniaceae.
Rev. Palaeobot. Palyn. 4:325-333.
Taylor, T.N. & D.A. Levin. 1975. Pollen morphology of Polemoniaceae in relation
to systematics and pollination systems: scanning electron microscopy. Grana
15:91-112.
Turner, B.L. 1994. Taxonomic overview of Gilia sect. Giliastrum (Polemoniaceae)
in Texas and México. Phytologia 76:52-68.
Phytologia (February 1998) 84(2):87-92.
A NEW SPECIES OF NEUROLAENA (ASTERACEAE) FROM OAXACA,
MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
A new species, Neurolaena jannaweissana B.L. Turner, is described
and illustrated from western Oaxaca. Its closest relationship is with N.
balsana, a species of northwestern Guerrero, both taxa known by only a single
collection. A revised key to the Mexican species of Neurolaena is provided,
along with a newly constructed map showing their distributions.
KEY WORDS: Asteraceae, Neurolaena, México, Oaxaca
In my treatment of Neurolaena (Turner 1982), I recognized ten species, most of
these occurring in México. I added two additional Mexican species, N. wendfii in
1985 and N. lamina in 1990; these and the present addition bring to thirteen the
number of taxa now recognized in Neurolaena, ten of these occurring in México. A
key to the Mexican species follows, along with two maps showing their distributions.
KEY TO MEXICAN SPECIES
I pleads radiate Oaxacan ss 3% sce cacdss tate s)-(ss eee Eee R eS Sam 04s acess cas N. oaxacana
|S AP eats GISCOIG: Fo. ..020.cancatbiegess swiedieins TM So 5s RE Gosia ois saree oo en tidveeean eee (2)
2. Pales (chaff) about 2/3 as long or longer than the subtended florets.......... (4)
2. Pales (chaff) of receptacle much shorter than the subtended
florets: leaves: 30-60! cm JONES «. «.\5:-... pes: SPR ois w'sise aeons uera'osie ve alvin es (3)
3. Undersurface of blades prominently resinous-glandular;
mid-ribs densely pubescent with long, dark brown, crisped,
multi-cellular hairs (4-8 septate); pedicels densely
brown-tomentose; achenes glabrous; VeracruZ.................00ce0eeeeee N. venturana
87
88 PHYTOLOGIA February 1998 volume 84(2):87-92
UNIVERSITY OF
EXAS
HERBARIUM
™
YRE OF ow 3 } . .
vi Cure Rene LAR weiANne
PLANTS OF MEXICO
OAXACA
Asenceac
Teevveinen x7
Determined by:
Distrito: Juquila
Municapicx Tataltepec de Valle,
Comuniskek: Taralicpoc de Valdes
Laviong: £16°21A9792' Af: 21000m
pine forest above Tataliepes, Fo C. La Chinche, sos
path towards Llama Wards 400-1200m)
2m Collector: Janea Wessy 988? $ Jon 1994
UNIVERSITY OF TEXAS AT AUSTIN (FEN)
Fig. 1. Neurolaena jannaweissana (holotype).
Tumer: New Neurolaena from Oaxaca 89
Fig. 2. Distribution of Neurolaena lobata in Mexico.
PHY TOLOGIA February 1998 volume 84(2):87-92
90
Pr
., a?
. ”
Ses
wornnnncs 6”
nae? + eg Nn
&
g
3s]
a a ¢
f.8 2_3..
Be BEEEE
>
Sassseaas
Zeger
Fig.:3. Distribution of Neurolaena spp. in Mexico
(other than N. lobata).
Turner: New Neurolaena from Oaxaca 91
3;
Undersurface of blades not, or inconspicuously, glandular
beneath; mid-ribs moderately to sparsely pubescent with
short, white, usually appressed hairs (2-5 septate); pedicels
appressed white-hispid; achenes pubescent; SE Chiapas. ............ N. macrophylla
4(2). Heads 6-12(-14) mm across (pressed); middle series of
inVolucralsbractsslinealsy l-2 mM WiIGe see eeeeene cre eaeeereerncerrensecc scene (5)
4. Heads 14-16 mm across (pressed); middle series of
involucral bracts narrowly ovate to ovate-orbicular,
(2:0 )2-S-4::5 MIM WAKE. Asis siesta coe abo ania. SrinsmemeR ota adeudeatesnnleeiacme (9)
. Undersurface of blades rather evenly soft pubescent (if hairs confined
to the ribs only, the pubescence then fine and appressed). .................20s0000 (7)
Undersurface of blades coarsely hispid along the veins only,
not evenly soft pubescent or densely resinous-glandular. .................20+0s008 (6)
6. Achenes very sparsely atomiferous-glandular; pales 2-3
mm shorter than the subtended florets; Oaxaca. ................ N. jannaweissana
6. Achenes densely atomiferous-glandular throughout; pales
about as long as the subtended florets; Guerrero. ..................005 N. balsana
7(5). Leaves entire to faintly crenate or rarely weakly serrate,
As
never lobed; involucral bracts glabrous or nearly so, the
middle series mostly 1 or 2-nerved; pales exceeding the
subtended florets; branches of capitulescence densely
fulvous(greenish-yellow) tomentose; Chiapas. ..............:.0c0ceeeeseeees N. fulva
Leaves serrate to variously lobed (the uppermost ones sometimes
entire or nearly so); involucral bracts mostly variously
puberulent, usually 3-nerved; pales shorter than the subtended
florets; branches of the capitulescence variously short-hispid,
the hairs brownish; not, preenish-yellow), «40. .agewnasodatncudiiompers sildianile selena tine (8)
8. Leaves thin, very sparsely pubescent beneath, not at all
glandular-atomiferous, the margins unlobed; southern
Meracruz, eastetn Oaxaca) a. 2b. tat benetinen aidteann ee ercaeucemebawialedt N. lamina
8. Leaves thick, densely pubescent beneath and
atomiferous-glandular, the margins frequently deeply
S-lobed: widespread .. omc. sas oss Bos votes aa Srienai emacs andeinncalasigives N. lobata
9(4). Outer involucral bracts appressed, broadly ovate,
a
without terminal hirsute appendages; coastal Veracruz............ N. macrocephala
Outer involucral bracts loose, narrowly lanceolate with
terminal hirsute appendages; easternmost Oaxaca and
Closely: adjacent. VELACINZ: a ..cccsndeticcn ans <lecinssdrine semnenctavell dese eames N. wendtii
NEUROLAENA JANNAWEISSANA B.L. Turner, spec. nov. Figure 1.
TYPE: MEXICO. Oaxaca: Distrito Juquila, Mpio. Tataltepec de Valdez, pine
forest near Tataltepec, E of La Chinche (ca. 16° 21’ N, 97° 33’ W), along path
towards Llano Verde, ca. 1000 m, 5 Jan 1994, Janna Weiss 687 (HOLOTYPE:
TEX):
Similis N. balsanae B.L. Turner sed foliis integris (vice foliorum
dentatorum), achenis glabris aut paene glabris (vice dense glandulosorum), et
paleis flosculis subtentis 2-3 mm _ breviorbus (vice aequalium aut longiorum).
92 PHYTOLOGIA February 1998 volume 84(2):87-92
Suffruticose herb or shrublet to 1 m high or more. Stems about 8-striate,
moderately pubescent with short, somewhat down-curved hairs. Leaves alternate, 12-
21 cm long, 3-4 cm wide; petioles 1-3 cm long, gradually tapering into the blades;
blades pinnately veined, about equally tapered at both ends, the lower veins more
pronounced than the upper, the surfaces moderately hispidulous, mostly along the
major veins, margins entire or nearly so. Inflorescence composed of about 30 heads
arranged in terminal clusters, the latter ca. 8 cm high, 10 cm across, the ultimate
peduncles 8-14 mm long. Heads eradiate; involucres ca. 9 mm high, 14 mm wide
(pressed); involucral bracts about 5-seriate, rather evenly imbricate, the outer series
acute apically, the middle and inner series obtuse or rounded, the middle series linear-
lanceolate, 1-nervate, ca. 1 mm wide. Receptacle ca. 2.5 mm across, ca. 0.5 mm
high, the linear pales scarious, ca. 6 mm long, 2-3 mm shorter than the subtended
florets. Florets numerous (40+) to a head; corollas yellow, glabrous, ca. 6 mm long,
the tubes ca. 3 mm long, throat ca. 2.1 mm long, and the lobes ca. 0.9 mm long.
Achenes ca. 2 mm long, black, the surfaces minutely black-warty, otherwise very
sparsely atomiferous-glandular, and having a few scattered hairs; pappus of 40-50
whitish bristles ca. 6 mm long.
Because of its reduced pales, this taxon will key to, or near, Neurolaena
venturana, in my taxonomic treatment of Neurolaena (Turner 1982). Its closest
affinities, however, appear to be with N. balsana B.L. Turner, a species of
northwestern Guerrero, to date known only by the type. Neurolaena jannaweissana
differs from the latter in having shorter pales (as already noted), entire leaves, and
nearly glabrous achenes (vs. densely atomiferous-glandular throughout).
It is a pleasure to name this species for Janna Weiss, newly crowned doctorate in
botany at the University of Texas, Austin, Texas, her doctoral thesis (Weiss 1998)
being an ethnobotanical study of the native peoples of western Oaxaca.
ACKNOWLEDGMENTS
I am grateful to Tom Wendt and Justin Williams for reviewing the manuscript.
LITERATURE CITED
Turner, B.L. 1982. Taxonomy of Neurolaena (Asteraceae-Heliantheae). Pl. Syst.
Evol. 140:119-139.
1985. A new species of Neurolaena (Asteraceae-Heliantheae) from
southernmost Veracruz, México. Phytologia 58:497-498.
1990. A new species of Neurolaena (Astereae, Heliantheae) from
southern México. Phytologia 69:172-174.
Weiss, J. 1998. Diagnostic concepts and medicinal plant use of the Chatino (Oaxaca,
Mexico) with a comparison of Chinese medicine. Doctoral Thesis. Univ. of
Texas, Austin, Texas, 394 pp.
Phytologia (February 1998) 84(2):93-97.
COMMENTS ON THE DISTRIBUTION OF TILLANDSIA RECURVATA L.
(BROMELIACEAE) IN TEXAS
H.L. White, J.R. Branch, & W.C. Holmes
Department of Biology, Baylor University, Waco, Texas 76798-7388 U.S.A.
ABSTRACT
Recent collections provide new records for the distribution of Tillandsia
recurvata L. (Bromeliaceae) in Texas. An alternate explanation for the
supposed spread of the species, attributed to local or regional environmental
change, is also offered.
KEY WORDS: Bromeliaceae, Tillandsia, Texas, biogeography
McWilliams (1992) chronicled the apparent northeastward expansion (Figure 1) of
the distribution of Tillandsia recurvata L. in Texas, which he attributed to local or
regional environmental change. Citing the observations of professional botanists, he
may have documented the spread and occurrence of the species in Brazos County, but
the remainder of the supposed spread is open to a different interpretation. His
expansion rests primarily upon the eastern limit of the historical distribution given by
Birge (1911) that was determined in the following manner:
“In order to determine the exact distribution of Tillandsia recurvata in the
Southwest[,] specimens were sent to the Superintendents of Schools of the
counties on the supposed border line with inquiry as to whether the plant grew
in that region. The resulting information together with personal observations
was used in making the map showing its distribution in Texas.”
The resulting map (Figure 1) resembles an irregular triangle with the apex reaching
as far north as southern McLennan County, the eastern side intersecting the Rio
Grande at Brownsville (Cameron County), the western side cutting the Rio Grande at
Del Rio (Val Verde County), and the base of the triangle extending along the Mexican
border. As mentioned above, the occurrence of the species in a given area rested upon
the observations of non-botanists near a supposed borderline. Apparently, specimens
were not sent to Birge, only information as to the species’ occurrence. Assuming
accurate recognition of the species by the superintendents, the method would not
account for the occurrence of the species outside of the supposed limits. Birge also
a3
94 PHYTOLOGIA February 1998 volume 84(2):93-97
mentions that the apex of the distribution triangle just touches the 31° north latitude in
Bell County, which is approximately four kilometers north of the present day town of
Salado. Yet she includes within the mapped distribution, the southern tip of
McLennan County, 54 km outside of the stated distribution. Perhaps to account for
error in her method, she extended the limits of the distribution of her map in a northern
direction and.may have done so for the eastern and western borders as well. As a
result, the Birge depiction of the distribution cannot be considered accurate.
Therefore, we believe that McWilliams’ (1992) reliance on Birge’s distribution does
not substantiate the spread of the species, but only documents its occurrence in the
eastern part of its distribution.
This hypothesis is further supported by the following new county records for the
species (see Figure 1).
Specimens cited: TEXAS. Atascosa Co.: rare in Prosopis pasture 31.5 mi S of
Jourdontan on Hwy 16, ca. 50 yd E of roadside park, 6 Apr 1975, Lewis & Joughin
s.n. (TEX-LL). Bastrop Co.: young oak-cedar-yaupon woods in & along ravine of
Wilbarger Creek drainage, ca. 1.3 to 1.4 mi ENE of jct FM 969 & FM 1704, 22 Nov
1986, Carr & Kutae 7964 (TEX-LL). Bell Co.: Moffat Cemetery off Hwy 36, 9 Jun
1998, White 518 (BAYLU). Brown Co.: Camp Bowie Military Reserve, Dept. of
Defense Site, ca. 2.4 mi E of jct US Hwy 377 & FM 45 on FM 45, in live oaks along
spring fed creek, 27 Jul 1998, Loar 00] (BAYLU). Caldwell Co.: roadside along
FM 713, 3.7 mi E of jct FM 86, between McMahan and Delhi, 18 Jul 1986, Lemke
661 (TAMU). Calhoun Co.: open grassy thickets above bay, 4 mi S of Port Lavaca,
on shrubs, Correll 28896 (TEX-LL). Colorado Co.: right side of FM 102, 1 mi S of
Eagle Lake, 12 Apr 1971, Thornton 24 (TAES). Dewitt Co.: 10 mi S of Cuervo on
US Hwy 183, 5 Mar 1993, Holmes 6334 & Yip (BAYLU). Edwards Co.: 14 mi SW
of Rockspring, Reid Ranch, 3 Apr 1983, Johnson 137 (BAYLU). Fayette Co.: Hwy
77, Schulenburg, 27 Mar 1971, Cary 155 (TAES). Gillespie Co.: on oaks in small
park 6 mi S of Fredericksberg on road to Kerrville, 18 May 1962, Correll & Ogden
25311 (TEX-LL). Gonzales Co.: oak trees in Palmetto State Park, camping area near
Park Road 11, 6 Apr 1968, Leonard 2023 (TEX-LL). Guadalupe Co.: 6 mi E of
Seguin, ign. leg. (TAES). Henderson Co.: Geddie farm, 3 mi N of Athens, 17 Nov
1973, Mathis 19 (TAES). Hidalgo Co.: Santa Anna Wildlife Refuge, 6 Apr 1957,
Correll & Schweinfurst 15663 (TEX-LL). Hill Co.: on E bank of Brazos River 0.6
km below Lake Whitney dam, 97.36468° lat. 31.86605° long., 6 Jun 1998, Branch
302 (BAYLV). Irion Co.: 4.3 mi E of Mertzon on Hwy 67 at jct with Spring Creek,
in live oaks and mesquite, 26 Jul 1998, Singhurst 6872 (BAYLU). Jim Wells Co.: in
woods of mesquite, hackberry, anacua, & live oaks near creek, 12.4 miS of Alice, 24
Nov 1954, Johnston 542126 (TEX-LL). Kenedy Co.: Norias Division of King
Ranch, 5.5 mi N of Norias, 3 Nov 1949, Lundell 15033 (TEX-LL). Kerr Co.: in
live oaks in Kerr Wildlife Management Area, 30 Jul 1998, Singhurst 6880 (BAYLU).
Kimble Co.: in live oaks along S Llano River in S Llano River State Park, 27 Jul
1998, Singhurst 6876 (BAYLU). Kleberg Co.: Hwy 771, ca. 1 mi W of water edge
at Riviera Beach, 100 m N of road, 13 Oct 1960, Tranverse 1799 (TEX-LL). Live
Oak Co.: Atascosa River 1.5 mi E of US 281 on FM 99, Whitsett, 19 Apr 1981,
Bensmiller 199 (TAMU). Madison Co.: Navasota River Bank at Hwy 21 Bridge, 27
Aug 1996, Neill 282 (TAMU). Mason Co.: in live oaks at Mason Mt. Wildlife
Management Area, 30 Jul 1998, Singhurst 6879 (BAYLU). McCulloch Co.: FM
1851 & San Saba River. 4.5 mi N of Fredonia, in live oak and mesquite, 24 Jul 1998,
White et al.: Comments on Tillandsia recurvata in Texas 95
Singhurst 6869 (BAYLU). McMullen Co.: Hwy 173, 10 mi N of Tilden, 3 Nov
1962, Gungora, Garza, & McCart 8529 (TEX-LL). Medina Co.: SW side of Hill
Country State Natural Area in live oaks along Bandera Creek, 27 Jul 1998, Singhurst
6878 (BAYLU). Menard Co.: jct of Hwy 864 & San Saba River, 3.9 mi W of Ft.
McKavett, 26 Jul 1998, Singhurst 6973 (BAYLU). Real Co.: Hwy 83, 1 mi S of
Leaky, 27 Mar 1964, Corasco & McCart 9031 (TEX-LL). Refugio Co.: mott near
Melon Creek, Red Well Pasture, Greta Ranch, ca. 8 mi NE of Refugio, 21 Jul 1981,
Hill 10589 (TAES). San Patricio Co.: 500 ft W of Big Lake, Welder Wildlife
Foundation Refuge, 12 Mar 1980, Critchfield 7 (TAMU). San Saba Co.: Colorado
Bend State Park along trail to Gorman Falls, in live oaks adjacent to Colorado River,
24 Jul 1998, Singhurst 6881 (BAYLU). Schleicher Co.: 3.8 mi N of jct FM 2084 &
Hwy 190, jct FM 2084 & Poor Hollow, in live oaks and elms, 25 Jul 1998, Singhurst
6871 (BAYLU). Sutton Co.: ENE of jct FM 3130 & IH-10 towards Roosevelt,
WSW of Cedar Hill Cemetery in live oaks along N Llano River, 26 Jul 1998,
Singhurst 6875 (BAYLU). Tom Green Co.: S Concho River in live oaks, 0.4 mi
NW of Christoval, 25 Jul 1998, Singhurst 6870 (BAYLU). - Williamson Co.: live
oaks W of Round Rock, 17 Aug 1946, York 46292 (TEX-LL).
Depending on location, the new records expand the known distribution of
Tillandsia recurvata presented by Birge (1911), Smith (1944), and McWilliams
(1992), from 100-240 kilometers northward. It also shows that the species is
considerably more widespread in the central, western, and northern parts of the
Edwards Plateau and nearby vegetational regions than previously thought.
The common feature among most of the new records is the proximity to large
bodies of water or permanent creeks, falls, or springs. In Irion County, the species is
fairly localized to Spring Creek in mainly Quercus fusiformis Small (Fagaceae) and
Prosopis (Fabaceae). In Tom Green County, along the South Concho River, T.
recurvata is very abundant in Quercus fusiformis, Prosopis, Celtis (Ulmaceae), and
Ulmus crassifolia Nutt. (Ulmaceae). Similarly, in the southeastern corner of Kimble
County, along the South Llano River, several healthy populations exist in creek
drainages. Menard County’s population, on the San Saba River, is a little more sparse
with only a few plants per tree. In McCulloch County, the species is abundant in Q.
fusiformis, Prosopis, and Juniperus ashei Buchh. (Cupressaceae) along the San Saba
River in the southern two thirds of the county. The number of 7. recurvata in Q.
fusiformis at the Brown County site is described as “plentiful along spring-fed creek.”
At the Hill County location, T. recurvata is abundant, with one to several plants per
tree for a distance of about 400 meters along the edge of a vertical limestone cliff.
Numerous seeps are perched on the cliff, which is 10-15 meters high on the east side
of the Brazos River. Several trees, primarily Juniperus ashei and Quercus fusiformis,
were infested with a hundred or more plants. The sheer number of plants occurring in
Hill County indicates a well-established and presumably old population. No specific
mention of abundance or a creek is made for the Henderson County specimen, but,
based on label information, the location would probably be nearby Caney Creek, three
miles north of Athens.
For regions with less rainfall. Birge (1911) mentions that Tillandsia recurvata “is
in many cases confined entirely to the river valleys.” She also says “. . . its northern
range is determined by the cold of the winters.” It appears moisture and heat retention
from large bodies of water moderate the local environment, forming a microclimate
suitable for this species’ growth.
96
PHYTOLOGIA February 1998 volume 84(2):93-97
Figure |. Shading depicts the distribution of Tillandsia recurvata in Texas reported by
Birge (1911). Closed circles represent records cited by Smith (1944). McWilliams’
(1992) expansion of the distribution shown as triangles. New county records illustrated
as Open circles (see text for explanation).
White et al.: Comments on Tillandsia recurvata in Texas 97
ACKNOWLEDGMENTS
We wish to thank Sidney McDaniel of the Institute for Botanical Exploration,
Mississippi State, Mississippi and Jason Singhurst of the Texas Parks and Wildlife
Department, Austin, Texas for review of this manuscript. This study would not have
been possible without the information and specimens provided by the curators of the
following herbaria: HPC, TAES, TAMU, and TEX-LL. Joe Yelderman of the
Department of Geology, Baylor University provided information about the geology of
the Lake Whitney Dam area of Hill County.
LITERATURE CITED
Birge, W.I. 1911. The Anatomy and Some Biological Aspects of the “Ball Moss,”
Tillandsia recurvata L. Univer. of Texas Bull 194:1-24.
McWilliams, E. 1992. Chronology of the Natural Range Expansion of Tillandsia
recurvata (Bromeliaceae) in Texas. SIDA 15 (2):343.
Smith, L.B. 1944. Bromeliaceae. Jn: Lundell, C.L. et al., Fl. Texas 3 (4):204-205.
Southern Methodist University Press, Dallas, Texas.
Phytologia (February 1998) 84(2):98-106.
PARAKEELYA: A NEW GENUS SEGREGATED FROM CALANDRINIA
(PORTULACACEAE)
Mark A. Hershkovitz!
‘Laboratory of Molecular Systematics, MSC, MRC 534, Smithsonian Institution,
Washington, DC 20560 U.S.A.
ABSTRACT
Extensive analyses of Portulacaceae have demonstrated that traditional
circumscriptions of the genus Calandrinia Kunth were artificial, and that the
included species were intertwined phylogenetically with many other
Portulacaceae lineages. Roger Carolin proposed that the Australian endemic
species of Calandrinia sensu lato, which are not related closely to Calandrinia
sensu stricto, be transferred to Rumicastrum Ulbr., but combinations were
never made. Rumicastrum appears to belong to Chenopodiaceae, however, as
Ulbrich suggested. Here, the genus Parakeelya, based on an aboriginal name,
is established to accommodate the Australian endemic calandrinias. New
combinations are provided for 38 names.
KEY WORDS: Australia, Calandrinia, Parakeelya, Portulacaceae,
phylogenetics, Rumicastrum
In seminal phylogenetic studies of Portulacaceae, Carolin (1987, 1993) determined
that the traditional circumscription of the genus Calandrinia Kunth (100-150 spp.;
e.g., Pax & Hoffmann 1934; cf. McNeill 1974; Nyanyano 1986) was unnatural, and
that the species were intertwined phylogenetically with many lineages of
Portulacaceae. Carolin’s general conclusion was corroborated in subsequent
investigations (Hershkovitz 199la, 1991b, 1993a, 1993b; Hershkovitz & Zimmer
1997, submitted ms.). Hershkovitz (1993a, 1993b) showed that Calandrinia sensu
stricto was a morphologically well-defined lineage comprising fourteen species native
to the Americas, and that these species are not related closely to a group of 35-50
Australian native species classified in Calandrinia sensu lato (though one species of
Calandrinia sensu stricto is adventive in Australia -- see below). These phylogenetic
results have been corroborated by ribosomal DNA sequence data (Hershkovitz &
Zimmer 1997; submitted ms.).
Carolin (1987, 1993) proposed that the Australian endemic calandrinias should be
transferred to the genus Rumicastrum Ulbr., which had been described as a monotypic
98
Hershkovitz: Parakeelya, a new genus in Portulacaceae 99
genus of Chenopodiaceae from Western Australia (Ulbrich 1934). Presumably,
Carolin believed that a specimen identified as Rumicastrum chamaecladum Ulbr.,
George 16288 (PERTH,B; Carolin 1987) actually was an unspecified Australian
calandrinia, notwithstanding that the diagnosis and illustration of Rumicastrum
(Ulbrich 1934) impart no hint of portulacaceous affinity. Carolin’s proposal was
reiterated in papers coauthored by Carolin (Syeda & Carolin 1989, 1990; Kiihn et al.,
1993). Hershkovitz (199la, 1991b, 1993a, 1993b) accepted on faith Carolin’s
proposal.
Nomenclatural realignments reflecting the compelling phylogenetic evidence were
formalized for several members of Calandrinia sensu lato and other Portulacaceae (see
Hershkovitz 1993a; Hershkovitz & Zimmer 1997), but the Australian calandrinias
were not renamed, even in subsequent floristic treatments (see below). Several factors
probably have contributed to the delay in renaming these plants, including inertia and
perhaps the name Rumicastrum itself, which seems perfectly appropriate for a
chenopod but unpalatable for attractive members of Portulacaceae. Also, Hershkovitz’
work concentrated primarily on American plants such that formal names for Australian
plants were dispensable.
Hershkovitz & Zimmer (1997) included an Australian calandrinia (Calandrinia
ptychosperma F. Muell.) in their analysis of ribosomal DNA data for the
portulacaceous alliance. In the original draft of their paper, they had recombined this
species under Rumicastrum. Werner Greuter and Brigitte Zimmer (pers. comm.,
13111997) kindly advised the authors that “Rumicastrum chamaecladum, as evidenced
by the specimen George 16288, which perfectly fits the original description (and
would be a good neotype) has nothing, but absolutely NOTHING to do with
Australian Calandrinia. We are persuaded it is a perfectly good chenopod reasonably
close to Atriplex.” The holotype of Rumicastrum chamaecladum presumably was
destroyed during World War II. Hershkovitz & Zimmer (1997) thus reverted to using
Calandrinia for their plant, as the circumscription of Calandrinia was not the focus of
that paper.
In a manuscript currently in review, Hershkovitz & Zimmer present ribosomal
DNA evidence reinforcing the morphologically-based circumscription of Calandrinia
sensu stricto. The data again show that the sampled Australian taxon isenot related
closely to this genus. In the context of the submitted manuscript, the appellation of
Calandrinia for the Australian plant is awkward and misleading, necessitating a new
generic name.
The present paper proposes the generic name Parakeelya for the Australian
calandrinias. This name derives from the vernacular name “parakeelya” applied
generically to Calandrinia polyandra Benth. by Black (1948), to Calandrinia
(presumably referring to the Australian species only) by West (1986), and, with
specific epithets, to other species of Australian calandrinias by Black (1948) and
numerous floristicians (see below). “Parakeelya” apparently is an alternative spelling
for “periculia,” the aboriginal vernacular for the seed meal of Calandrinia balonensis
Lindl. (Mueller 1876; cf. Poellnitz 1934).
The only purpose of the present paper is to validate names for the Australian
species. A comprehensive generic description is problematic. The Australian
calandrinias comprise a diverse assemblage. For some characters, the species
100 PHYTOLOGIA _ February 1998 volume 84(2):98-106
approach the full range of variation found in Portulacaceae as a whole (e.g., growth
habit, seed geometry and sculpturing, pollen morphology; Carolin 1987, 1993; Syeda
1979; Syeda & Ashton 1989; Syeda & Carolin 1989, 1990; see also floristic
references cited below). There appears to be no known single character that ties the
species together. Some features that occur among these species are unique or unusual
among Portulacaceae, e.g., capsules that split only at the apex, polyforate-operculate
pollen, three-dimensional leaf venation, and anisocytic stomata (Carolin 1987, 1993;
Hershkovitz 1991b). In fact, monophyly of the Australian species has not been
demonstrated. Carolin’s (1987) cladogram of Portulacaceae showed polyforate-
operculate pollen as a synapomorphy of this group, but several species have
pantocolpate pollen (Carolin 1987; Kelley 1973), which plausibly is primitive in this
group (Carolin 1987; Hershkovitz 1993a). Polyphyly of the Australian species is not
supported, either. Specifically, there are no features that link particular species to
otherwise divergent lineages of Portulacaceae. To the contrary, and despite the lack of
a clear synapomorphy, the species appear to be linked to one another by combinations
of features.
Because of the variability in this group, as well as the poor representation of
herbarium material of these taxa in the United States and Europe, the generic diagnosis
will emphasize the traits of the one species for which ribosomal DNA evidence is
available, Calandrinia ptychosperma F. Muell. The diagnosis basically will be that of
Mueller (1876, elaborated from Mueller 1864), amended to include mention of
polyforate-operculate pollen. The diagnosis obviously will omit reference to the range
of variation in the species being transferred, but, at the same time, it will be much
more applicable to the Australian species in general than is that of Calandrinia.
Calandrinia ptychosperma is designated here as the type for the generic name, even
though the vernacular name “parakeelya” refers to either or both of Calandrinia
balonensis and C. polyandra. Calandrinia balonensis and C. polyandra have 12-15-
pantocolpate pollen (Carolin 1987; Kelley 1973), while C. ptychosperma has
polyforate-operculate pollen. To the degree that polyphyly of the Australian plants is
conceivable, these alternative pollen states represent one of the plausible distinctions. I
have observed, however, that C. balonensis is among the Australian species that have
three-dimensional leaf venation (Hershkovitz 1991b), so I suspect that additional
research will show that the Australian plants indeed are monophyletic.
Specific combinations will be provided for those species that otherwise are validly
published and accepted in the most recent Australian floristic and taxonomic works
(Beard 1970; Blackall & Grieve 1988; Chapman 1991; Cunningham et al. 1992; Green
1985; Queensland Herbarium 1993; Syeda 1979, 1981, 1996; Walsh 1996; West
1986, 1987, 1990, 1992). Chapman (1991) listed 60 specific and subspecific
combinations in Calandrinia in Australia, of which 53 pertain to Parakeelya. One
name pertains to Anacampseros (Hershkovitz 1993), and six have been applied in
reference to the American native C. ciliata (Ruiz & Pavén) DC., which has established
adventively in Australia. [All current Australian floras use C. menziesii (Hook.)
Torrey & A. Gray for this species. Discordance with current opinion in American
works (e.g., Kelley 1993) aside, documentation of the identity of the Australian plants
with, specifically, the type of C. menziesii appears to be lacking.] For the group here
circumscribed as Parakeelya, Syeda & Carolin (1990) indicated that there are 34
species, while Carolin (1993) indicated that there are 50. Review of the floristic work
suggests that 33 validly published species are accepted. These plus one problematic
Hershkovitz: Parakeelya, a new genus in Portulacaceae 101
name are transferred here to Parakeelya. Two additional species described by Syeda
(1979) remain unpublished (Syeda & Carolin 1988), and another two apparently are
unidentified/undescribed (West 1990; Queensland Herbarium 1993). Additional
names listed by Chapman (1991) that pertain to Parakeelya are disposed in the
discussion following the taxonomic treatment below.
PARAKEELYA Hershkovitz, gen. nov. TYPE: Parakeelya ptychosperma (F.
Muell.) Hershkovitz (Calandrinia ptychosperma F. Muell.).
Herba humilis, glabra, foliis radicalibus lineari-filiformibus confertis
acutis, caulinis brevioribus, racemis paucifloris, bracteis acutis scariosis,
pedicellis calyce ter aliquotiesve rarius vix longioribus aetate erectiusculis,
sepalis persistentibus rotundo-ovatis acutatis, pollinis grana polyporus-
operculatus, stylo nullo, capsula calycem breviter superante vel aequante apice
quadrivalvi cylindrico-conica, seminibus nitentibus fusco-atris simpliciter
longitudinali-sulcatis fere reniformibus.
PARAKEELYA ARENICOLA (Syeda) Hershkovitz, comb. nov. BASIONYM:
Calandrinia arenicola Syeda, Proc. Linn. Soc. New South Wales 116:153. 1996.
PARAKEELYA BALONENSIS (Lindl.) Hershkovitz, comb. nov. BASIONYM:
Calandrinia balonensis Lindl. in T.L. Mitchell, J. Exped. Trop. Australia. 148.
1848.
PARAKEELYA BREVIPEDATA (F. Muell.) Hershkovitz, comb. nov.
BASIONYM: Calandrinia brevipedata F. Muell, Fragm. 10:69. 1876.
PARAKEELYA CALYPTRATA (Hook. ff.) Hershkovitz, comb. nov.
BASIONYM: Calandrinia calyptrata Hook. f. in Hook., Icon. Pl. 3:296. 1840.
PARAKEELYA COMPOSITA (Nees) Hershkovitz, comb. nov. BASIONYM:
Calandrinia polypetala Fenzl in Endl. et al. var. composita Nees in Lehm., PI.
Preiss. 1:247. 1845.
PARAKEELYA CORRIGIOLOIDES (F. Muell. ex Benth.) Hershkovitz, comb.
nov. BASIONYM: Calandrinia corrigioloides F. Muell. ex Benth., Fl. Austral.
175.1863.
PARAKEELYA CREETHAE (Tratman ex Morrison) Hershkovitz, comb. nov.
BASIONYM: Calandrinia creethae Tratman ex Morrison, J. Bot. 50:165. 1912.
PARAKEELYA DISPERMA_ (J.M._ Black) Hershkovitz. comb. nov.
BASIONYM: Calandrinia disperma J.M. Black, Trans. & Proc. Roy. Soc. South
Australia 45:11, t. If. 1921.
PARAKEELYA EREMAEA (Ewart) Hershkovitz, comb. nov. BASIONYM:
Calandrinia eremaea Ewart, Fl. Victoria 486. 1921. SYNONYM: Calandrinia
pusilla Lindl., nom. illegit., in T.L. Mitchell, J. Exped. Trop. Australia. 360.
1848, non Calandrinia pusilla Barnéoud 1846.
102 PHYTOLOGIA _ February 1998 volume 84(2):98-106
PARAKEELYA GRACILIS (Benth.) Hershkovitz, comb. nov. BASIONYM:
Calandrinia gracilis Benth., Fl. Austral. 1:173. 1863.
PARAKEELYA GRANULIFERA _ (Benth.) Hershkovitz, comb. nov.
BASIONYM: Calandrinia granulifera Benth., Fl. Austral. 1:176. 1863.
All current floristic references (see above) regard Talinum nanum Nees 1845 as
a taxonomic synonym of Calandrinia granulifera Benth. Nee's epithet has priority,
but it was not available for recombination in Calandrinia (see discussion below).
Parakeelya granulifera becomes the valid name for Calandrinia granulifera if it is
considered distinct from Talinum nanum.
PARAKEELYA LEHMANNII (Endl.) Hershkovitz, comb. nov. BASIONYM:
Calandrinia lehmannii Endl. in Lehm., Pl. Preiss. 2:235. 1848.
PARAKEELYA LINIFLORA (Fenzl) Hershkovitz, comb. nov. BASIONYM:
Calandrinia liniflora Fenzl in Endl. et al., Enum. Pl. 52. 1837.
PARAKEELYA NANA (Nees) Hershkovitz, comb. nov. BASIONYM: Talinum
nanum Nees in Lehm., Pl. Preiss. 1:246. 1845. SYNONYMS: Calandrinia
pygmaea F. Muell. nom. illegit., Fragm. 1:175. 1859; Calandrinia neesiana H.
Eichler, Taxon 12:295. 1963.
Calandrinia pygmaea F. Muell. was illegitimate because it was a homotypic
synonym of Talinum nanum Nees. Nonetheless, Calandrinia pygmaea was
adopted in taxonomic and floristic works until the middle of the 20th century (e.g.,
Bentham 1863; Poellnitz 1934; Black 1948), by which time the proper
recombination of Talinum nanum had been preempted by Calandrinia nana Philippi
1894. [Beard (1970) included the name “‘Calandrinia nana (Nees) C.A. Gardn.” in
his checklist, without a reference or basionym.] Calandrinia neesiana H. Eichler
became the new name for Talinum nanum, but this now is widely regarded as a
taxonomic synonym of Calandrinia granulifera Benth. Parakeelya nana has
priority over Calandrinia granulifera when these are regarded as the same species.
PARAKEELYA PAPILLATA (Syeda) Hershkovitz, comb. nov. BASIONYM:
Calandrinia papillata Syeda, Telopea 2:60. 1980.
PARAKEELYA PICKERINGII (A. Gray) Hershkovitz, comb. nov.
BASIONYM: Calandrinia pickeringii A. Gray, U.S. Expl. Exped., Phan. 1:144.
1854.
PARAKEELYA PLEIOPETALA (F. Mutell.) Hershkovitz, comb. nov.
BASIONYM: Calandrinia pleiopetala F. Muell., Fragm. 10:70. 1876.
PARAKEELYA POLYANDRA (Benth.) Hershkovitz, comb. nov. BASIONYM:
Calandrinia polyandra Benth., Fl. Austral. 1:172. 1863. SYNONYM: Talinum
polyandrum Hook. nom. illegit., Bot. Mag. 4833. 1855, non Talinum
polyandrum Ruiz & Pavon, Syst. Veg. Fl. Peruv. Chil. 115. 1798.
PARAKEELYA POLYPETALA (Fenzl) Hershkovitz, comb. nov. BASIONYM:
Calandrinia polypetala Fenzl in Endl. et al., Enum. Pl. 51. 1837.
Hershkovitz: Parakeelya, a new genus in Portulacaceae 103
PARAKEELYA PORIFERA (Syeda) Hershkovitz, comb. nov. BASIONYM:
Calandrinia porifera Syeda, Telopea 2:59. 1980.
PARAKEELYA PRIMULIFLORA (Diels) Hershkovitz, comb. nov.
BASIONYM: Calandrinia primuliflora Diels in Diels & E. Pritz., Bot. Jahrb.
Syst. 35:198, fig. 24 A-F. 1904.
PARAKEELYA PTYCHOSPERMA (F. Muell.) Hershkovitz, comb. nov.
BASIONYM: Calandrinia ptychosperma F. Muell., Fragm. 4:137. 1864.
PARAKEELYA PUMILA (Benth.) Hershkovitz, comb. nov. BASIONYM:
Calandrinia calyptrata Hook. f. in Hook. var. pwmila Benth., Fl. Austral. 1:175.
1863.
PARAKEELYA QUADRIVALVIS (F. Muell.) Hershkovitz, comb. nov.
BASIONYM: Calandrinia quadrivalvis F. Muell., Fragm. 1:176. 1859.
PARAKEELYA REMOTA (J.M. Black) Hershkovitz, comb. nov. BASIONYM:
Calandrinia remota J.M. Black, Trans. & Proc. Roy. Soc. South Australia 47:369.
1923.
PARAKEELYA RETICULATA (Syeda) Hershkovitz, comb. nov. BASIONYM:
Calandrinia reticulata Syeda, Telopea 2:60. 1980.
PARAKEELYA SCHISTORHIZA _ (Morrison) Hershkovitz, comb. nov.
BASIONYM: Calandrinia schistorhiza Morrison, J. Bot. 50:164. 1912.
PARAKEELYA SPERGULARINA (F. Muell.) Hershkovitz, comb. nov.
BASIONYM: Calandrinia spergularina F. Muell., Fragm. 1:175. 1859.
PARAKEELYA SPHAEROPHYLLA (J.M. Black) Hershkovitz, comb. nov.
BASIONYM: Calandrinia sphaerophylla J.M. Black, Trans. & Proc. Roy. Soc.
South Australia 51:378. 1927.
PARAKEELYA STAGNENSIS (J.M. Black) Hershkovitz, comb. nov.
BASIONYM: Calandrinia stagnensis J.M. Black, Trans. & Proc. Roy. Soc.
South Australia 51:379. 1927.
PARAKEELYA STENOGYNA (Domin) Hershkovitz, comb. nov. BASIONYM:
Calandrinia stenogyna Domin, Biblioth. Bot. 22(89):971. 1926
PARAKEELYA STROPHIOLATA (F. Muell.) Hershkovitz, comb. nov.
BASIONYM: — Claytonia_ strophiolata F. Muell., Fragm. 11:82. 1880.
SYNONYM: Calandrinia strophiolata (F. Muell.) Poelln., Feddes Repert. Spec.
Nov. Regni Veg. 35:173. 1934.
PARAKEELYA TUMIDA (Syeda) Hershkovitz, comb. nov. BASIONYM:
Calandrinia tumida Syeda, Proc. Linn. Soc. New South Wales 116:156. 1996.
104 PHYTOLOGIA February 1998 volume 84(2):98-106
PARAKEELYA UNIFLORA (F. Muell.) Hershkovitz, comb. nov. BASIONYM:
Calandrinia uniflora F. Muell., Trans. & Proc. Philos. Inst. Victoria 3:41. 1857.
PARAKEELYA VOLUBILIS (Benth.) Hershkovitz, comb. nov. BASIONYM:
Calandrinia volubilis Benth., Fl. Austral. 1:174. 1863.
Based on the cited taxonomic and floristic references, additional names (excluding
autonyms) in Calandrinia listed by Chapman (1991) are disposed taxonomically as
follows:
Calandrinia calyptrata Hook. f. in Hook. var. pumila Benth, homotypic synonym of
Parakeelya pumila.
Calandrinia caulescens Kunth, synonym of Calandrinia ciliata.
Calandrinia caulescens Kunth var. menziesii (Hook.) A. Gray, synonym of
Calandrinia ciliata.
Calandrinia ciliata (Ruiz & Pavén) DC., accepted.
Calandrinia compressa Schrad. ex DC., accepted, probably does not occur in Australia
(Poellnitz 1934).
Calandrinia cygnorum Diels, synonym of Parakeelya brevipedata.
Calandrinia dipetala J.M. Black, synonym of Parakeelya calyptrata.
Calandrinia liniflora Fenzl in Endl. et al. var. grandiflora Benth., synonym of
Parakeelya liniflora.
Calandrinia maryonii S. Moore, synonym of Parakeelya ptychosperma.
Calandrinia menziesii (Hook.) Torrey & A. Gray, synonym of Calandrinia ciliata.
Calandrinia morrisae Goy, synonym of Parakeelya ptychosperma.
Calandrinia pogonophora F. Muell., synonym of Anacampseros australiana J.M.
Black.
Calandrinia polyandra Benth. var. leptophylla Benth., status uncertain (West 1986; cf.
Black 1948).
Calandrinia polypetala Fenzl in Endl. et al. var. composita Nees, homotypic synonym
of Parakeelya composita.
Calandrinia tepperiana W. Fitzg., synonym of Parakeelya quadrivalvis.
Calandrinia volubilis Benth. var. parvula J.M. Black, status uncertain, not mentioned
in cited floras (including Black 1948).
ACKNOWLEDGMENTS
I thank Urs Eggli and Dan Nicolson for manuscript review and Dan, especially,
for critical discussion.
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South Wales. Vol. 1. New South Wales University Press, Kensington, New
South Wales, Australia.
West, J.G. 1992. Portulacaceae. Pp. 133-138 in J.R. Wheeler (ed.), B.L. R. Rye,
B.L. Koch, & A.J.G. Wilson, Flora of the Kimberly Region. Western
Australia Herbarium, Department of Conservation & Land Management,
Como, Western Australia, Australia.
Phytologia (February 1998) 84(2):107-153.
ANNOTATED CHECKLIST OF THE VASCULAR PLANTS OF WALKER,
MONTGOMERY, AND SAN JACINTO COUNTIES, EAST TEXAS
Guy L. Nesom
Texas Research Institute for Environmental Studies, Sam Houston State University,
Huntsville, Texas 77340-2506 U.S.A.
&
Larry E. Brown
Houston Community College, 1300 Holman, Houston, Texas 77004 U.S.A.
ABSTRACT
An annotated checklist is provided of the vascular plant species of Walker,
Montgomery, and San Jacinto counties of southeast Texas, an area totaling
2450 square miles and completely including the Sam Houston National Forest.
Most of the three-county area is within the “Pineywoods” Texas Natural
Region; smaller portions are within three other Natural Regions: the
“Blackland Prairie,” ““Oak Woods and Prairies,” and “Gulf Coastal Prairie.” A
total of 1373 species in 623 genera and 160 families are recorded. Distribution
records are documented individually by county, either with the herbarium
repository of a voucher specimen or a literature citation. Asteraceae is the
largest family in the flora with 170 species; next largest are Poaceae (155
species), Fabaceae (120 species), and Cyperaceae (105 species). The largest
genera are Carex with (37 species), Cyperus (18 species), Rhynchospora (17
species), Desmodium and Eleocharis (15 species), Dichanthelium and Juncus
(14 species), Eupatorium, Symphyotrichum, and Trifolium (11 species),
Ludwigia (10 species), Hypericum, Paspalum, Solidago, and Viola (9
species), Eragrostis, Helianthus, Panicum, Polygonum, and Ranunculus (8
species), and Chamaesyce, Crataegus, Euphorbia, Lespedeza, Liatris,
Oenothera, Scutellaria, Sisyrinchium, Smilax, and Solanum (7 species).
KEY WORDS: floristics, Texas, checklist
This checklist provides a preliminary summary of the vascular plant species known
to occur in Walker, Montgomery, and San Jacinto counties, Texas. It also provides a
guide to species that may be found within the Sam Houston National Forest, whose
107
108 PHYTOLOGIA _ February 1998 volume 84(2):107-153
boundary lies completely within these three counties. It seems likely that the number
of species in the checklist may be expanded at least 10% as specialized habitats
(especially wetland and deep sand) are studied in more detail. We believe, however,
that the broad floristic outline is indicated here.
The three~county flora includes 1373 species in 160 families and 623 genera. The
average number of species per genus is 2.2. Asteraceae is the largest family in the
flora with 170 species; next largest are Poaceae (155 species), Fabaceae (120 species),
and Cyperaceae (105 species). Other families are significantly smaller (e.g.,
Lamiaceae with 39 species, Euphorbiaceae with 37, Scrophulariaceae with 34,
Apiaceae with 30, Rosaceae with 26, Onagraceae with 24, Fagaceae with 21, and
Ranunculaceae and Rubiaceae with 18). The largest genus, by far, is Carex with 37
species. Other relatively large genera are as follows: Cyperus (18 species),
Rhynchospora (17 species), Desmodium and Eleocharis (15 species), Dichanthelium
and Juncus (14 species), Eupatorium, Symphyotrichum, and Trifolium (11 species),
Ludwigia (10 species), Hypericum, Paspalum, Solidago, and Viola (9 species),
Eragrostis, Helianthus, Panicum, Polygonum, and Ranunculus (8 species), and
Chamaesyce, Crataegus, Euphorbia, Lespedeza, Liatris, Oenothera, Scutellaria,
Sisyrinchium, Smilax, and Solanum (7 species).
Documentation for the occurrence of each taxon is indicated by a symbol for its
county or counties of occurrence. Voucher specimens for most of the records are
deposited in SHST (Sam Houston State University, Huntsville) and SBSC (Spring
Branch Science Center, Houston), the identities of these verified by Nesom and
Brown. Some records (as indicated) are taken from other herbaria and from literature,
and detailed surveys of other Texas herbaria undoubtedly will provide additional
records.
The comprehensive treatment of Correll & Johnston (1970) remains the primary
basis for assessments of the Texas flora. Recent nomenclatural updates (Johnston
1989; Hatch et al. 1990) also have been useful. Nomenclature in the current checklist
mostly follows Jones et al. (1996) and Kartesz (in Kartesz & Meachum 1998). Where
the primary entry in the current checklist differs from the latter two references, other
currently used names are provided as synonyms.
A 1978 fire destroyed most of the plant collection at SHST, which had been
amassed over a number of decades by S.R. Warner (primarily for Walker County) and
served as the basis of knowledge for the local flora. Fortunately, a checklist and
summary of the Walker County plants (including 643 species) was prepared in 1977
by Tom Zimmerman (unpublished), based on Warner’s field notes and herbarium
specimens. Warner’s studies of the Walker County flora included particular attention
to the relationship of plant distribution to soil type, this information included in
Zimmerman’s compilation. Warner made careful identifications, but a small set of taxa
said by Warner to occur in Walker County were probably based on misidentifications
or earlier taxonomic interpretations: these are noted below. Most of the species
recorded by Warner have been recollected for SHST since the 1978 herbarium fire; 61
remain to be recollected. The more recent additions to SHST have been made since
1978 primarily by Claude McLeod, Michael Warnock, Guy Nesom, and various
students. James Kessler added many regional collections of Cyperaceae and other taxa
in the mid 1970’s. Kathy Nelson (1996) added and summarized records for
Asteraceae of Walker County.
Nesom & Brown: Plant checklist for southeast Texas counties 109
Herbarium SBSC is part of the Spring Branch Science Center of the Spring
Branch Independent School District in Houston. It is the largest herbarium in
southeast Texas, with over 31,000 specimens of vascular plants. The herbarium was
begun by R.A. Vines, but more than half of the collections are those of Larry Brown,
as it now serves as the primary repository for vouchers for Brown’s floristic
investigations of the Houston area and southeast Texas. Among these studies, an
annotated checklist of Harris County (including Houston) and the eight counties
contiguous with it (including Montgomery County) is currently in preparation.
A published florula of the Little Thicket Nature Preserve (Peterson & Brown 1983;
San Jacinto County, 650 acres, 485 species, including keys) is very helpful. A
number of records from San Jacinto County on the present list are documented by
Little Thicket vouchers in SBSC.
A checklist of the vascular species of Huntsville State Park (2122 acres, 320
species) is maintained by the Texas Parks & Wildlife Department (Singhurst 1998).
This summary is based originally on a list prepared by Claude McLeod in 1976, with
additions and revisions by TP&WD personnel, including recent collections by Jason
Singhurst. Specimens documenting a number of these HSP records are deposited in
BAYLU (Baylor University) and SHST.
Treatments of the Texas species of legumes (Turner 1959) and ferns and fern allies
(Correll 1956) also have furnished additions to the present checklist not documented
by specimens in SHST, as does a treatment of Asteraceae for Texas (Turner
unpublished). Carr (1994) conducted a floristic study of blackland prairie openings
within the Sam Houston National Forest, these records partly documented by
specimens deposited at TEX. A few records have been taken from various other
published references, as specifically indicated within the list.
Description of the area
Walker County covers about 505,600 acres (790 square miles), Montgomery
County about 697,000 acres (1089 square miles), and San Jacinto County about
365,250 acres (571 square miles), the whole area totaling about 1,568,000 acres
(2450 square miles). Sam Houston National Forest includes about 161 thousand acres
(about 1/10 the total area). Tracts that most nearly approach being “natural areas” (the
least disturbed, with the highest species diversity) are Huntsville State Park (2100
acres in Walker County), Lake Houston State Park (4900 acres partly in Montgomery
County), several relatively small, privately owned nature preserves in San Jacinto
County, and two “Scenic Areas” and one “Wilderness Area” within SHNF that are
currently somewhat exempted from the otherwise heavy economic orientation of
timber management in the National Forest. Otherwise, much of the private land also is
heavily managed for timber production and there is much rural “development” in the
form of housing, ranching, and farming. Conroe and The Woodlands (Montgomery
Co.), and Huntsville (Walker Co.) are the largest urban areas.
The Trinity River flows across the northern tip of Walker County and then forms
the north and eastern boundary of San Jacinto County, where it is impounded as Lake
Livingston. The East Fork and West Fork of the San Jacinto River flow southward
110 Pry TOroGTA February 1998 volume 84(2):107-153
through the area — the latter is the primary input for Lake Conroe in Walker and
Montgomery County. Several variants of bottomland hardwood communities, highly
disturbed for the most part, occur along these and other rivers and creeks.
The climate of the three-county area is hot; daily high temperatures from May
through October commonly are above 90 degrees Fahrenheit. The area receives an
average range of about 43-48 inches of rainfall annually.
Soils in the three-county area were deposited in the Tertiary and Quaternary
periods. The largest areas are Pliocene and Pleistocene sands (Willis, Bentley, and
Beaumont Formations), often with a clay subsoil. Clay prairies in Walker and
Montgomery counties and clay inclusions (all three counties) within the sandy areas of
pine-oak forest are Fleming Formation (Miocene). Sandstone outcrops in Walker
County are Catahoula Formation (Miocene) -- most of the large outcrops have been
removed by quarrying. Alluvial deposits of the larger rivers are extensive in places.
General nature of the vegetation
Parts of Walker County and Montgomery counties are within the “Blackland
Prairie” Natural Region (sensu LBJ School of Public Affairs 1978). Small, discrete
patches of prairie vegetation on clay substrate extend eastward across the region as
inclusions within the pine-hardwood forests. These prairie inclusions are strongly
predictable in floristic composition (Carr 1993) and are most similar in vegetation to
the “Fayette Prairie” (Smeins and Diamond 1983) immediately west of the area. Much
of the western half of Walker County is “Oak Woods & Prairies (Oak Woodlands)”
Natural Region. The southern extremity of Montgomery County lies in the “Gulf
Coastal Prairie” Natural Region.
Most of the three-county area, however, is within the “Pineywoods” Natural
Region. Because of the prevalence of timber management, pines usually are
dominant, either as plantations or mixed with various combinations of hardwoods. A
few white oak and laurel oak dominated stands can be found on mesic slopes in Sam
Houston National Forest. Stands of scrubby post oak - blackjack oak, however,
which typically occur on xeric ridges, have almost all been replaced by roads or
managed pine stands.
Walker and Montgomery counties lie essentially at the southwestern comer of the
Eastern Deciduous Forest, the relatively sharp vegetational change to prairie molded by
rapidly decreasing precipitation (westward) and an abrupt transition from primarily
sandy soils to the prairie clays. A conspicuous set of woody species reaches the
western maximum of its distribution in Walker County (e.g., sweetgum, red maple,
southern magnolia, cherry laurel, nutmeg hickory, the two hornbeam species,
sassafras, fringetree, silverbell, witch hazel, dwarf chinquapin, black oak, white oak,
and swamp chestnut oak). Other Walker County species are near their western
distributional limit (e.g., shortleaf and loblolly pine, dogwood, black gum, white ash,
persimmon, bitternut hickory, water hickory, river birch, southern red oak, willow
oak, and overcup oak). Eastern Montgomery County and a major part of San Jacinto
County lie within the “Big Thicket” region (sensu McLeod 1971). Walker County is
significantly drier than these areas and the geographic distribution of many species
does not cross westward into Walker County (e.g., beech, laurel oak, longleaf pine,
Nesom & Brown: Plant checklist for southeast Texas counties 111
sweetbay, redbay, maple-leaved viburnum, horsesugar, gallberry holly, pepperbush,
titi, and sweetspire (Nesom 1998; Nesom et al. 1997).
ACKNOWLEDGMENTS
Thanks to Stanley Jones (Botanical Research Institute, Bryan), whose records for
Cyperaceae added numerous species to the checklist and numerous new records for
county distribution. Jason Singhurst (Texas Parks & Wildlife Department, Austin)
provided a copy of his floristic summary for Huntsville State Park. And special
thanks to Michael and Barbara MacRoberts (Bog Research, Shreveport) for their
urging and encouragement to publish this checklist, even in what we consider a
preliminary form.
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Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of Texas.
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Correll, D.S. & H.B. Correll. 1975. Aquatic and Wetland Plants of Southwestern
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Do, L.H., W.C. Holmes, & J.R. Singhurst. 1996. New records for Bellardia
trixago (Scrophulariaceae) in Texas. Sida 17:295-296.
Godfrey, R.K. & J.W. Wooten. 1979 & 1981. Aquatic and Wetland Plants of the
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Gould, F.W. 1975. The Grasses of Texas. Texas A&M University, College Station,
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Hartman, D.H. 1974. Forest Structure of the Big Thicket Scenic Area, Texas. M.S.
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Station, Texas.
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Jones, S.D., J.K. Wipff, & P.M. Montgomery. 1997. Vascular Plants of Texas: A
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233.
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by S.R. Warner (1921-1924). Unpublished manuscript.
CHECKLIST
Symbols for documentation of distribution
W = Walker County: voucher SHST; W = voucher SBSC
M = Montgomery County: voucher SHST; M = voucher SBSC
J = San Jacinto County: voucher SHST; J = voucher SBSC
(astc, brch, mich, sbsc, smu, taes, tex) = acronyms for other herbaria of voucher
repository
(mercer) = herbarium of Mercer Arboretum, Humble, Texas
W(reference), M(reference) , or J(reference) = literature report
*J = added from Peterson & Brown (1983)
114 PHY TOL OGTITA February 1998 volume 84(2):107-153
P = added from survey of blackland prairie inclusions (Walker Co.; Carr 1994)
H = added from Huntsville State Park list (Walker Co.; Singhurst 1998)
0 = added from Warner checklist for Walker County (Zimmerman 1977): no SHST
voucher
*[species] = no voucher known for any of the three counties
FERNS
ASPLENIACEAE
Asplenium platyneuron (L.) B.S.P. WMJ
AZOLLACEAE
Azolla caroliniana Willd. WM
BLECHNACEAE
Woodwardia areolata (L.) Moore WM(Correll 1956)J
SYN= Lorinseria areolata (L.) Presl
DENNSTAEDTIACEAE
Pteridium aquilinum (L.) Kuhn WM(Correll 1956)J
DRYOPTERIDACEAE
Athyrium filix-femina (L.) Roth WM(Correll 1956)J
Onoclea sensibilis L. WM(Correll 1956)J
Polystichum acrostichoides (Michx.) Schott WMJ
Woodsia obtusa (Spreng.) Torr. W(Correll 1956)
LYGODIACEAE
Lygodium japonicum (Thunb. ex Murr.) Sw. WMJ
OPHIOGLOSSACEAE
Botrychium dissectum Spreng. WM(Correll 1956)*J
Botrychium lunarioides (Michx.) Sw. W
Botrychium virginianum (L.) Sw. W
Ophioglossum crotalophoroides Walt. WJ
Ophioglossum engelmannii Prantl W
OSMUNDACEAE
Osmunda cinnamomea L. WJ
Osmunda regalis L. WMJ
POLYPODIACEAE
Pleopeltis polypodioides (L.) Andrews & Windham WMJ
SYN= Polypodium polypodioides (L.) Watt
Nesom & Brown: Plant checklist for southeast Texas counties 115
PTERIDACEAE
Cheilanthes alabamensis (Buckl.) Kunze W
Cheilanthes tomentosa Link W
Pellaea atropurpurea (L.) Link W
Pteris multifida Poir. M(Correll 1956)
THELYPTERIDACEAE
Thelypteris kunthii (Desv.) Morton WM(Correll 1956)J
SYN= Thelypteris normalis (C. Chr.) Moxley
Thelypteris x versicolor St.John J(Correll 1956)
SYN= Thelypteris quadrangularis (Fée) Schelpe var. versicolor (St.John) A.
Smith
SYN= Thelypteris hispidula (Decne.) C.F. Reed var. versicolor (St.John)
Lellinger
FERN ALLIES
EQUISETACEAE
Equisetum laevigatum A. Br. WM
ISOETACEAE
Isoetes melanopoda Gay & Durieu ex Durieu W
LYCOPODIACEAE
Lycopodiella appressa (Chapm.) Cranfill J
SYN= Lycopodium appressum (Chapm.) Lloyd & Underwood
SELAGINELLACEAE
Selaginella apoda (L.) Spring WM(Correll 1956)J
Selaginella arenicola Underwood subsp. riddellii (Van Eselt) Tryon W
SYN= Selaginella riddellii Van Eselt
GYMNOSPERMS
PINACEAE
Pinus echinata P. Mill. WMJ
Pinus elliottii Engelm. WM[fide C&J 1970] J
Pinus palustris P. Mill. J
Pinus taeda L. WJ
TAXODIACEAE
Taxodium distichum (L.) Rich. W*J
CUPRESSACEAE
Juniperus virginiana L. WJ
Thuja orientalis L. W
116 PHYTOLOGIA _ February 1998 _ volume 84(2):107-153
MONOCOTS
AGAVACEAE
Manfreda virginica (L.) Salisb. WJ
SYN= Agave virginica L.
SYN2= Polianthes virginica (L.) Shinners
Yucca flaccida Haw. W
Yucca louisianensis Trel. W*J
Yucca treculeana Carr. W
ALISMATACEAE
Sagittaria graminea Michx. WMJ
Sagittaria latifolia Willd. WJ
Sagittaria papillosa Buch. M
AMARYLLIDACEAE
Cooperia drummondii Herb. WJ
Cooperia pedunculata Herb. M
Crinum bulbispermum (Burm.) Milne-R. & Schw. W
Habranthus tubispathus (L'Herit.) Traub WM
SYN= Habranthus texanus (Herb.) Steud.
Hymenocallis liriosme (Rafin.) Shinners WJ
Leucojum aestivum L. W
Manfreda virginica (L.) Salisb.
SYN= Polianthes virginica (L.) Shinners WJ
Narcissus sp. W
ARACEAE
Arisaema dracontium (L.) Schott WJ
Arisaema triphyllum (L.) Schott OJ
Colocasia esculenta (L.) Schott W
Peltandra virginica (L.) Schott W
Pistia stratiotes L. M
Xanthosma sagittifolium (L.) Schott W
ARECACEAE
Sabal minor (Jacq.) Pers. WMJ
BROMELIACEAE
Tillandsia recurvata (L.) L. W
Tillandsia usneoides (L.) L. WMJ
BURMANNIACEAE
Apteria aphylla (Nutt.) Barnh. ex Small J
Burmannia biflora L. J
COMMELINACEAE
Callisia repens (Jacq.) L. W
Commelina communis L. WJ
Commelina erecta L. WM*J
Commelina virginica L. WMJ
Nesom & Brown: Plant checklist for southeast Texas counties 117
Tinantia anomala (Torr.) C.B. Clarke W
Tradescantia hirsutiflora Bush W*J
Tradescantia ohiensis Rafin. W
Tradescantia reverchonii Bush W
CYPERACEAE
Bulbostylis capillaris L.) C.B. Clarke M(Kral 1971)
Bulbostylis ciliatifolia (Ell.) Fern. var. coarctata (Ell.) Kral WM(brch)*J
Carex abscondita Mack. WM(brch)J
Carex albicans Willd. ex Spreng. var. australis (Bailey) Rettig WM)J(taes)
SYN= Carex physorhyncha Liebm. ex Steud.
Carex albolutescens Schwein. WMJ
Carex corrugata Fern. W*J
_ SYN= Carex amphibola Steud. misappl.
Carex crebriflora Wieg. J
Carex basiantha Steud. WJ(smu)
_ SYN= C. willdenowii Schkuhr. ex Willd. misappl.
Carex blanda Dewey W*J
Carex brevior (Dew.) Mack. ex Lunell W(brch)MJ(taes)
Carex bushii Mack. WMJ
Carex caroliniana Schwein. W*J
Carex cephalophora Muhl. ex Willd. H*J
Carex cherokeensis Schwein. WMJ
Carex comosa Boott W
Carex complanata Torr. & Hook. WMJ
Carex crus-corvi Shuttlew. ex Kuntze WJ
Carex debilis Michx. WJ
Carex digitalis Schkuhr ex Willd. var. asymmetrica Fern. WJ
Carex emoryi Dewey W<(brch)
Carex fissa Mack. var. fissa W(mich)
Carex flaccosperma Dewey WMJ
Carex frankii Kunth WMJ
Carex glaucescens Ell. J
Carex hyalina Boott WM(brch)
Carex intumescens Rudge WMJ
Carex joorii Bailey J(brch)
Carex leavenworthii Dewey WM(tex)J(smu)
Carex leptalea Wahl. WMJ
Carex longii Mack. M(taes)
Carex louisianica Bailey WM(mich)J
Carex lupuliformis Sartw. ex Dewey M
Carex lupulina Muhl. ex Willd. WMJ
Carex lurida Wahl. WMJ
Carex meadii Dew. W
Carex microdonta Torr. & Hook. WM(smu)
Carex microrhyncha Mack. W(taes)
Carex muhlenbergii Schkuhr. ex Willd. W*J
Carex oxylepis T. & H. WJ
Carex retroflexa Muhl. ex Willd. WM*J
Carex socialis Mohl. & Schwegm. W(brch)
Carex striatula Michx. W
118 PHYTOLOGIA _ February 1998 volume 84(2):107-153
Carex texensis (Torr.) Bailey W(brch)J(taes)
Carex triangularis Boeck]. WMJ(taes)
Carex tribuloides Wahlenb. var. sangamonensis Clokey WJ
Carex verrucosa Muhl. M(borch)J(brch)
Carex vulpinoidea Michx. M
Cyperus articulatus L. W
Cyperus croceus Vahl M(taes)J(brit)
Cyperus echinatus (L.) Wood WMJ
SYN= Cyperus ovularis (Michx.) Torr.
Cyperus entrerianus Bockler M(taes)
Cyperus erythrorhizos Muhl. WJ
Cyperus esculentus L. WJ
Cyperus filiculmis Vahl HM(Marcks 1972)
Cyperus haspan L. WM
Cyperus hystricinus Fern. WJ
Cyperus iria L. M(brit)
Cyperus ochraceus Vahl H
Cyperus odoratus L. var. squarrosus (Britt.) Jones et al. MJ
Cyperus plukenetii Fern. WJ
Cyperus polystachyos Rottb. WM
Cyperus pseudovegetus Steud. WMJ
Cyperus retroflexus Buckl. var. retroflexus W(taes)M(mich)J
Cyperus retrorsus Chapm. WMJ
Cyperus rotundus L. WMJ
Cyperus squarrosus L. M .
SYN= Cyperus aristatus Rottb.
Cyperus strigosus L. WMJ
Cyperus surinamensis Rottb. M(smu)
Cyperus thyrsiflorus Jungh. W
Cyperus virens Michx. WMJ
Eleocharis acicularis (L.) R. & S. W
Eleocharis albida Torr. W
Eleocharis atropurpurea (Retz.) Pres! M
Eleocharis equisetoides (Ell.) Torr. M
Eleocharis interstincta (Vahl) R. & S. M
Eleocharis palustris (L.) R. & S. WM
SYN= Eleocharis macrostachya Britt.
Eleocharis microcarpa Torr. MJ
Eleocharis montana (Kunth) R.& S. M
Eleocharis montevidensis Kunth WM(smu)J(mich)
Eleocharis obtusa (Willd.) Schultes WMJ
Eleocharis olivacea Torr. W
Eleocharis parvula (R. & S.) Link WM
Eleocharis quadrangulata (Michx.) R. & S. WM(mich)
Eleocharis radicans (Dietr.) Kunth W
Eleocharis tortilis (Link) Schultes WMJ
Eleocharis tenuis (Willd.) Schultes WM .
SYN= Eleocharis verrucosa (Svens.) Harms
Eleocharis vivipara Link W
Fimbristylis autumnalis (L.) R. & S. WMJ
Fimbristylis decipiens Kral M(brch)
Nesom & Brown: Plant checklist for southeast Texas counties 119
Fimbristylis miliacea (L.) Vahl WM(brch)J
Fimbristylis puberula (Michx.) Vahl var. puberula W
Fimbristylis vahlii (Lam.) Link WM(taes)J
Fuirena breviseta (Cov.) Cov. M
Fuirena simplex Vahl WM
Fuirena squarrosa Michx. WM
Isolepis carinata Hook. & Arm. ex Torr. WM
SYN= Scirpus koilolepis (Steud.) Gl.
Isolepis molesta (M.C. Johnston) S.G. Sm. M
SYN= Scirpus molestus M.C. Johnston
Kyllingia brevifolia Rottb. WM(taes)
SYN= Cyperus brevifolius (Rottb.) Hassk.
Kyllingia pumila Michx WMJ
SYN= Cyperus tenuifolius (Steud.) Dandy
Rhynchospora caduca Ell. WMJ
Rhynchospora capitellata (Michx.) Vahl J
Rhynchospora colorata (L.) H. Pfeiffer W
SYN= Dichromena colorata (L.) Hitche.
Rhynchospora corniculata (Lam.) Gray WMJ
Rhynchospora divergens M.A. Curtis M(Correll & Correll 1975)
Rhynchospora filifolia Gray M(tex)
Rhynchospora globularis (Chapm.) Small WMJ
Rhynchospora glomerata (L.) Vahl WMJ
Rhynchospora grayi Kunth W
Rhynchospora harveyi Boott W
Rhynchospora inexpansa (Michx.) Vahl WMJ
Rhynchospora microcarpa Baldw. ex Gray W
Rhynchospora miliacea (Lam.) Gray W
Rhynchospora mixta Britt. M
Rhynchospora nivea Bock. W(us)
Rhynchospora perplexa Britt. MJ
Rhynchospora rariflora (Michx.) Ell. *J
Rhynchospora recognita (Gale) Kral W(taes)
Schoenoplectus pungens (Vahl) Palla var. pungens W(taes)
Scirpus cyperinus (L.) Kunth WM(taes)J
Scirpus pendulus Muhl. WJ SYNE= Scirpus lineatus Michx. misappl.
Scleria ciliata Michx. WM
Scleria oligantha Michx. W*J
Scleria pauciflora (Torr.) A. Nelson W(taes)
Scleria triglomerata Michx. WM
DIOSCOREACEAE
Dioscorea bulbifera L. M
Dioscorea villosa L. WJ
ERIOCAULACEAE
Eriocaulon decangulare L. WM
HYDROCHARITACEAE
Hydrilla verticillata Royale WM
Limnobium spongia (Bosc.) Steud. M
120 PHYTOLOGIA _ February 1998 volume 84(2):107-153
HYPOXIDACEAE
Hypoxis hirsuta (L.) Cov. WMJ
Hypoxis curtisii Rose J(sbsc)
Hypoxis LSS (Baker) Brackett W(tex)J(sbsc)
IRIDACEAE
Alophia drummondii (Grah.) R.C. Foster W
Iris hexagona Walt. MJ
Tris pseudacoris L. M
Tris virginica L. W
Nemastylis geminiflora Nutt. WM
Sisyrinchium angustifolium Mill. *J
Sisyrinchium exile Bickn. WM*J
Sisyrinchium langloisii Greene WMJ
Sisyrinchium minus Engelm. & Gray W
Sisyrinchium rosulatum Bickn. WJ
Sisyrinchium sagittiferum Bickn. WJ
Sisyrinchium texanum Bickn. WM
JUNCACEAE
* Juncus acuminatus Lam. 0
Juncus brachycarpus Engelm. WM
Juncus bufoniusL. M
Juncus capitatus Weigel WM
Juncus coriaceus Mack. WM*J
Juncus dichotomus Ell. WM
Juncus diffusissimus Buckley WMJ
Juncus effusus L. WMJ
Juncus marginatus Rostk. WM*J
Juncus nodatus Cov. J
Juncus repens Michx. WMJ
Juncus scirpoides Lam. WM*J
Juncus tenuis Willd. W*J
Juncus validus Cov. WMJ
Luzula bulbosa (Wood) Rydb. WMI
LEMNACEAE
Lemna perpusilla Torr. W
Lemna valdiviana Phil. W
Spirodela polyrhiza (L.) Schleid. W
Spirodela punctata (Mey.) Thomps. W
SYN= Spirodela oligorhiza (Kurtz) Hegelm.
Wolfia brasiliensis Wedd. W
SYN= Wolfia papulifera Thomps.
LILIACEAE
Allium canadense L. var. canadense WM*J
Allium drummondii Regel W*J
Nesom & Brown: Plant checklist for southeast Texas counties 121
Allium mobilense Regel WM
SYN= Allium canadense L. var. mobilense (Regel) M. Ownbey
Asparagus officinalis L. W
Camassia angusta (Engelm. & Gray) Blankinsh. W
Nothoscordum bivalve (L.) Britt. WMJ
Polygonatum biflorum (Walt.) Ell. *J
Schoenolirion wrightii Sherman W
SYN= Schoenolirion texanum (Scheele) A. Gray
Stenanthium gramineum (Ker-Gawl.) Morong W
Trillium gracile Freeman J
MAYACACEAE
Mayaca fluviatilis Aublet J
SYN= Mayaca aubletii Michx.
NAJADACEAE
Najas guadalupensis (Spreng.) Magnus W
ORCHIDACEAE
Corallorrhiza wisteriana Conrad HOM*J
Habenaria repens Nutt. J
SYN= Platanthera repens (Nutt.) Wood
Listera australis Lindl. W*J
Platanthera ciliaris (L.) Lindl. M
SYN= Habenaria ciliaris (L.) R. Br. ex Ait. f.
Platanthera clavellata (Michx.) Luer J
SYN= Habenaria clavellata (Michx.) Spreng.
Platanthera cristata (Michx.) Luer M(Correll 1944)
Ponthieva racemosa (Walt.) Mohr J
Spiranthes cernua (L.) Richard WJ
Spiranthes gracilis (Bigel.) Beck WJ
SYN= Spiranthes lacera var. gracilis (Bigel.) Luer
Spiranthes tuberosa Rafin. W*J
SYN= Spiranthes grayi Ames
Spiranthes praecox (Walt.) S. Wats. *J
Spiranthes vernalis Engelm. & Gray WM
Tipularia discolor (Pursh) Nutt. HM(sight)J
Triphora trianthophora (Sw.) Rydb. J(Hartman 1974)
POACEAE
“Agrostis elliottiana Schult. *J
Agrostis hyemalis (Walt.) B.S.P WJ
Agrostis perennans (Walt.) Tuckerm. *J
Aira caryophyllea L. *J
Aira elegans Willd. ex Kunth WMJ
SYN= Aira caryophyllea var. capillaris A. Mute
Andropogon gyrans Ashe W
SYN= Andropogon elliottii Chapm.
Andropogon gerardii Vitman W
Andropogon glomeratus (Walt.) B.S.P. WJ
122 PHYTOLOGIA February 1998 volume 84(2): 107-153
Andropogon ternarius Michx. W
Andropogon virginicus L. W
Aristida lanosa Ell. W
Aristida longispica Poir. var. geniculata (Rafin.) Fern. M*J
Aristida oligantha Michx. WMJ
Aristida purpurascens Poir. W
Aristida purpurea Nutt. var. longiseta (Steud.) Vasey W
SYN= Aristida longiseta Steud.
Arundinaria gigantea (Walt.) Muhl. WM(mercer)J
Arundo donax L. M
Avena sativa L. WM
SYN= Avena fatua L. var. sativa (L.) Hausskn.
Axonopus fissifolius (Raddi) Kuhlm. WM
SYN= Axonopus affinis Chase
*Axonopus furcatus (Fluegge ) Hitchc. 0
Bothriochloa ischaemum (L.) Keng W*J
Bothriochloa laguroides (DC.) Herter subsp. torreyana (Steud.) Allred & Gould
WJ
SYN= Bothriochloa saccharoides (Sw.) Rydb.
Bouteloua curtipendula (Michx.) Torr. WJ
Bouteloua hirsuta Lag. WJ
Bouteloua rigidiseta (Steud.) Hitchc. WJ
Briza minor L.
Bromus japonicus Thunb. WM
Bromus pubescens Muhl. ex Willd. W
Bromus secalinus L. WM
Bromus catharticus Vahl WMJ
SYN= Bromus unioloides (Willd.) Kunth
*Cenchrus echinatus L. 0
Cenchrus spiniflex Cav. WMJ
SYN= Cenchrus incertus Curtis
Chasmanthium latifolium (Michx.) Yates WMJ
Chasmanthium laxum (L.) Yates WMJ
Chasmanthium sessiliflorum (Poir.) Yates WMJ
Chloris cucullata Bisch. W
Chloris subdolichostachya Muller W
Coelorachis rugosa (Nutt.) Nash M
Cortaderia selloana (Schultes) Asch. & Graebn. W
SYN= Cortaderia dioica (Spreng.) Spreng.
Cynodon dactylon (L.) Pers. W*J ©
Cynosurus echinatus L. W
Dactylis glomerata L. M .
*Dactyloctenium aegypticum (L.) Willd. 0
Desmazeria rigida (L.) Tutin H
SYN= Catapodium rigidum (L.) Hubbard ex Dony
Dichanthelium aciculare (Desv. ex Poir.) G. & Cl. WJ
SYN= Dichanthelium angustifolium (Ell.) Gould
Dichanthelium acuminatum (Sw.) Gould & Clark WMJ
Dichanthelium boscii (Poir.) G. & Cl. WJ SYN= Panicum boscii Poir.
Dichanthelium commutatum (Schult.) Gould WM
SYN= Panicum commutatum Schult.
Nesom & Brown: Plant checklist for southeast Texas counties 123
Dichanthelium dichotomum (L.) Gould WM*J
SYN= Panicum dichotomum L.
Dichanthelium lanuginosum (Ell.) Gould W*J
SYN= Panicum lanuginosum Ell.
SYNE= Dichanthelium acuminatum var. fasciculatum (Torr.) Freckm.
Dichanthelium laxiflorum (Lam.) Gould WJ
SYN= Panicum laxiflorum Lam.
Dichantheliuwn lindheimeri (Nash) Gould WJ
SYN= Panicum lindheimeri Nash
SYN= Dichanthelium acuminatum var. lindheimeri (Nash) G. & Cl.
Dichanthelium linearifolium (Scribn. ex Nash) Gould WJ
SYN= Panicum linearifolium Scribn. ex Nash
Dichanthelium oligosanthes (Schultes) Gould var. oligosanthes W*J
SYN= Panicum oligosanthes Schultes
Dichanthelium oligosanthes (Schultes) Gould var. scribnerianum (Nash) Gould
WM
SYN= Panicum scribnerianum Nash
Dichanthelium ravenelii (Scribn. & Merr.) Gould. WJ
SYN= Panicum ravenelii Scribn. & Mert.
Dichanthelium scoparium (Lam.) Gould WM
SYN= Panicum scoparium Lam.
Dichanthelium sphaerocarpon (Ell.) Gould. WMJ
SYN= Panicum sphaerocarpon Ell.
SYN= Dichanthelium polyanthes (Schultes) Mohlenbr.
Digitaria ciliaris (Retz.) Koel. W*J
Digitaria cognata (Schult.) Chase W
SYN= Leptoloma cognatum (Schultes) Chase
Digitaria filiformis (L.) Koel. W
Digitaria villosa (Walt.) Pers. WJ
Digitaria violascens Link *J
Echinochloa colona (L.) Link W
Echinochloa crus-gallii (L.) Beauv. WM
Echinochloa walteri (Pursh) Heller WM
Eleusine indica (L.) Gaertn. WM
Elymus canadensis L. W
Elymus virginicus L. WM*J
Eragrostis barrelieri Daveau W
Eragrostis elliottii S. Wats. W
Eragrostis hypnoides (Lam.) B.S.P. M
Eragrostis hirsuta (Michx.) Nees W*J
*Eragrostis pectinacea (Michx.) Nees ex Steud. 0
Eragrostis secundiflora Pres!) WM*J
SYN= Eragrostis oxylepis (Torr.) Torr.
Eragrostis spectabilis (Pursh) Steud. W
*Eragrostis trichodes (Nutt.) Wood 0
Eremochloa ophiuroides (Munro) Hack. W
Erianthus alopecuroides (L.) Ell. W
SYN= Saccharum alopecuroidum (L.) Nutt.
Erianthus giganteus (Walt.) Muhl. WMJ
SYN= Saccharum giganteum (Walt.) Pers.
Erianthus strictus Ell. WM SYN= Saccharum baldwinii Spreng.
124 PAY LOLOG LA February 1998 volume 84(2):107-153
Eustachys petraea (Sw.) Desv. W
SYN= Chloris petraea Sw.
Festuca arundinacea Schreb. MJ
SYN= Lolium arundinaceum (Schreb.) S. Darb.
*Festuca subverticillata (Pers.) Alexeev 0
SYN= Festuca obtusa Bieler
Glyceria arkansana Fern. J
Gymnopogon ambiguus (Michx.) B.S.P. WJ
Hordeum pusillum Nutt. WMJ
Hordeum vulgare L. W
Leersia hexandra Sw. M(mercer)
Leersia oryzoides (L.) Sw. WM
Leersia virginica Willd. WMJ
Leptochloa mucronata (Michx.) Kunth WM
’ SYN= Leptochloa filiformis (Lam.) Beauv. var. attenuata (Nutt.) Steyerm.
Limnodea arkansana (Nutt.) Dewey WJ
Lolium perenne L. WM
SYNE Lolium multiflorum Lam.
Lolium tementulum L. W
Melica mutica Walt. WMJ
Muhlenbergia capillaris (Lam.) Trin. W*J
Oplismenus hirtellus (L.) Beauv. WJ
SYN= Oplismenus setarius (Lam.) R. & S.
Oryza sativa L. W
Panicum anceps Michx. WJ
Panicum brachyanthum Steud. W*J
Panicum capillare L. W
Panicum dichotomiflorum Michx. W
Panicum gymnocarpon Ell. WMJ
SYN= Phanopyrum gymnocarpon (Ell.) Nash
Panicum rigidulum Bosc ex Nees HJ
Panicum verrucosum Muhl. WJ
Panicum virgatum L. W
Paspalum dilatatum Poir. WM*J
Paspalum floridanum Michx. W
Paspalum laeve Michx. W*J
Paspalum langei (Fourn.) Nash W*J
Paspalum notatum Fluegge WMJ
Paspalum plicatulum Michx. WMJ
Paspalum pubiflorum Rupr. ex Fourn. WMJ
Paspalum setaceum Michx. WMJ
SYN= Paspalum pubescens Muhl.
Paspalum urvillei Steud. WMJ
Phalaris caroliniana Walt. W
Phyllostachys aurea Carr.ex Riv. W
Poa annua L. WJ
Poa autumnalis Muhl. ex Ell. WMJ
Rostraria cristata (L.) Tzveley W
SYN= Koehleria gerardii (Vill.) Shinners
Sacciolepis striata (L.) Nash WM
Schedonnardus paniculatus (Nutt.) Trel. W
Nesom & Brown: Plant checklist for southeast Texas counties 125
Schizachyrium scoparium (Michx.) Nash WJ
Secale cereale L. M
Setaria corrugata (Ell.) Schult. W(Gould 1975)
Setaria glauca (L.) Beauv. WM
SYN= Pennisetum glaucum (L.) R. Br.
Setaria parviflora (Poir.) Kerg. WJ
SYN= Setaria geniculata auct. non (Wild.) Beauv.
Setaria viridis (L.) Beauv. W
Sorghastrum elliottii (Mohr) Nash WJ
Sorghastrum nutans (L.) Nash W
Sorghum bicolor (L.) Moench M
Sorghum halepense (L.) Pers. WM
Sphenopholis longiflora (Vasey) Hitchc. M
Sphenopholis obtusata (Michx.) Scribn. WMJ
Sporobolus compositus (Poir.) Merr. var. compositus WMJ
SYN= Sporobolus asper (Beauv.) Kunth
Sporobolus indicus (L.) R. Br. WMJ
*Sporobolus junceus (Beauv.) Kunth H
Sporobolus vaginiflorus (Torr.ex Gray) Wood W
Steinchisma hians (Ell.) Nash WM
SYN= Panicum hians El.
Stenotaphrum secundatum (Walt.) Kuntze W
Stipa avenacea L. WJ
SYN= Piptochaetium avenaceum (L.) Parodi
Stipa leucotricha Trin. & Rupr.
SYN= Nassella leucotricha (Trin. & Rupr.) Pohl WMJ
Tridens flavus (L.) Hitchc. WJ
Tridens strictus (Nutt.) Nash WM
Triplasis purpurea (Walt.) Chapm. *J
Tripsacum dactyloides (L.) L. WM
Trisetum interruptum Buckl. WJ
Urochloa ciliatissima (Buckl.) R. Webster *J
SYNE= Brachiaria ciliatissima (Buckl.) Chase
Urochloa fasciculata (Sw.) R. Webster W
SYN= Panicum fasciculatum Swartz
Urochloa ramosa (L.) R. Webster M
SYN= Panicum ramosum L.
Urochloa texana (Buckl.) R. Webster O*J
SYN= Panicum texanum Buckl.
Vulpia octoflora (Walt.) Rydb. WJ SYN= Festuca octoflora Walt.
Vulpia myuros (L.) Gmel. W
Vulpia sciurea (Nutt.) Henr. J
Zizaniopsis miliacea (Michx.) Doell. & Asch. WM
PONTEDERIACEAE
Eichhornia crassipes (Mart.) Solms W
Heteranthera limosa (Sw.) Willd. M
POTAMOGETONACEAE
Potamogeton diversifolius Rafin. HM(Ogden 1966)
Potamogeton nodosus Poir. WMJ
126 PHYTOLOGIA February 1998 volume 84(2):107-153
SMILACACEAE
Smilax bona-nox L. WJ
Smilax glauca Walt. WMJ
Smilax laurifolia L. WMJ
Smilax pumila Walt. WMI
Smilax rotundifolia L. WMI
Smilax smallii Morong WJ
Smilax tamnoides L. WJ
SYN= Smilax hispida Muhl.
SPARGANIACEAE
Sparganium americanum Nutt. J
TYPHACEAE
Typha angustifolia L. W
*Typha domingensis Pers. 0
Typha latifolia L. W
XYRIDACEAE
Xyris ambigua Bey. ex Kunth WMJ
Xyris iridifolia Chapm. M
Xyris jupicai L.C. Rich. M
Xyris torta J.E. Sm. WJM
Xyris sp. J :
DICOTS
ACANTHACEAE
Dicliptera brachiata (Pursh) Spreng. W*J
Hygrophila lacustris (S. & C.) Nees WM
Justicia lanceolata (Chapm.) Small WM
SYN= Justicia ovata var. lanceolata (Chapm.) R.W. Long
Ruellia brittoniana Leonard M
Ruellia caroliniensis (Gmel.) Steud. WM
Ruellia humilis Nutt. WM*J
Ruellia nudiflora (Engelm. ex Gray) Urb. var. nudiflora WMJ
Ruellia pedunculata Torr. ex Gray J(Turner 1991)
Ruellia strepens L. WM
Yeatesia viridiflora (Nees) Small *J
ACERACEAE
Acer negundo L. WMJ
Acer rubrum L. WMJ
AMARANTHACEAE
Alternanthera philoxeroides (Mart.) Griseb. W
*Amaranthus albus L. O
SYN= Amaranthus graecizans auct. non L.
Nesom & Brown: Plant checklist for southeast Texas counties 127
Amaranthus hybridus L. WM(Reed 1969)
*Amaranthus retroflexus L. 0
*Amaranthus spinosus L. 0
Froelichia floridana (Nutt.) Mog. WM(Reed 1969)J
Gomphrena globosa L. M
Tresine rhizomatosa Standl. WJ
ANACARDIACEAE
Rhus aromatica Aiton WMJ
Rhus copallinum L. WMJ
Rhus glabra L. WM(Barkley 1943)*J
Toxicodendron radicans (L.) Kuntze var. radicans WM*J
SYN= Rhus radicans L.
Toxicodendron pubescens P. Mill. WM(Barkley 1943)J
* SYN= Rhus toxicodendron L.
ANNONACEAE
Asimina parviflora (Michx.) Dun. MJ
APIACEAE
Bifora americana (DC.) Benth. & Hook. W
*Bowlesia incana R. & P. 0
Centella erecta (L.f.) Fern. WMJ
Chaerophyllum tainturieri Hook. WM*J
Cicuta maculataL. WM .
Cyclospermum leptophyllum (Pers.) Sprague WJ
SYN= Apium leptophyllum (Pers.) F.V. Muell. ex Benth.
Cynosciadium digitatum DC. W
Daucus carota L. WM
Daucus pusillus Michx. WMJ
*Eryngium hookeri Walp. 0
Eryngium prostratum Nutt. ex DC. WMJ
Eryngium yuccifolium Michx. W*J
Hydrocotyle ranunculoides L. M
Hydrocotyle umbellata L. WM
Hydrocotyle verticillata Thunb. WMJ
Limnosciadium pinnatum (DC.) Math. & Const. WM
Oxypolis rigidior (L.) Rafin. W
Polytaenia nuttallii DC. W
Polytaenia texana (Coult. & Rose) Math. & Const. *J
Ptilimnium capillaceum (Michx.) Rafin. W
Ptilimnium nuttallii (DC.) Britt. W
Sanicula canadensis L. WMJ
Sanicula odorata (Rafin.) Pryer & Phillippe W SYN= Sanicula gregaria Bickn.
Spermolepis divaricata (Walt.) Rafin. ex Ser. WJ
*Spermolepis echinata (Nutt. ex DC.) Math. & Const. 0
Tauschia texana A. Gray M
Torilis arvensis (Huds.) Link W
*Torilis nodosa (L.) Gaertn. 0
Trepocarpus aethusae Nutt. ex DC. W*J
Zizia aurea (L.) Koch WJ
128 Pry TOL OGIA February 1998 volume 84(2):107-153
APOCYNACEAE
’ Amsonia tabernaemontana Walt. W
Apocynum cannabinum L. W
Nerium oleander L. W
Trachelospermum difforme (Walt.) Gray WMJ
Vinca major L. W
AQUIFOLIACEAE
Ilex ambigua (Michx.) Torrey WMJ
Ilex coriacea (Pursh) Chapm. MJ
Ilex decidua Walt. WM(Lundell 1943)J
Ilex longipes Chapm. ex Trel. WMJ
Ilex opaca Aiton WMJ
Ilex vomitoria Aiton WMJ
ARALIACEAE
Aralia spinosa L. WMJ
Hedera helix L. W
ARISTOLOCHIACEAE
Aristolochia reticulata Jacq. WMSJ
Aristolochia serpentaria L. WJ
Aristolochia tomentosa Sims *J
ASCLEPIADACEAE
Asclepias amplexicaulis J.E. Smith WM
Asclepias tuberosa L. WMJ
Asclepias variegata L. WJ
Asclepias verticillata L. W
Asclepias viridiflora Rafin. WMJ
Asclepias viridis Walt. WMJ
*Cynanchum laeve (Michx.) Pers. H
Matelea cynanchoides (Engelm.) Woods. WM
Matelea decipiens (Alex.) Woods. WM
Matelea gonocarpos (Walt.) Shinners WJ
ASTERACEAE
Achillea millefolium L. subsp. lanulosa (Nutt.) Piper WM
Acmella oppositifolia (Lam.) Jansen var. repens (Walt.) Jansen WMJ
Ageratina altissima (L.) King & Rob. WJ
SYN= Eupatorium rugosum Houtt.
Ambrosia artemisiifolia L. HJ
Ambrosia cumanensis Kunth WJ
SYN= Ambrosa psilostachya DC.
Ambrosia trifida L. W
Amphiachyris dracunculoides (DC.) Nutt. W
Antennaria fallax Greene WM
SYN= Antennaria parlinii var. fallax (Greene) Bayer & Stebbins
Anthemis cotula L. W
Aphanostephus skirrhobasis (DC.) Trel. *J
Nesom & Brown: Plant checklist for southeast Texas counties 129
Arnoglossum ovatum (Walt.) H. Rob. MJ
SYN= Cacalia ovata Walt.
SYN= Cacalia lanceolata Nutt.
Arnoglossum plantagineum Rafin. WMJ
SYN= Cacalia plantaginea (Rafin.) Shinners
Artemisia ludoviciana Nutt. var. ludoviciana M
Baccharis halimifolia L. WJ
Berlandiera pumila (Michx.) Nutt. var. pumila WMJ
Bidens aristosa (Michx.) Britt. WM
Bidens bipinnata L. WJ
Bidens frondosa L. W
Bidens laevis (L.) B.S.P. W.
Bigelowia nuttallii L.C. Anderson W
Boltonia diffusa Ell. MWJ
Brickellia eupatorioides (L.) Shinners var. eupatorioides WMJ
Calyptocarpus vialis Less. WJ
Centaurea americana Nutt. W
Centaurea cyanis L. WM
Chaetopappa asteroides Nutt. ex DC. var. asteroides WM
Chlorocantha spinosa (Benth.) Nesom M
SYN= Aster spinosus Benth.
Chrysopsis pilosa Nutt. WMJ
Chrysopsis texana Nesom W
SYN= Bradburia hirtella Torr. & Gray
Cirsium carolinianum (Walt.) Fern. & Schub. W
Cirsium horridulum Michx. W
Cirsium texanum Buckley W
Conoclinium coelestinum (L.) DC. WJ
SYN= Eupatorium coelestinum L.
Conyza canadensis (L.) Crong. WJ
Conyza bonariensis (L.) Crong. W
Coreopsis basalis (Dietrich) Blake WJ
Coreopsis lanceolata L. WM
Coreopsis tinctoria Nutt. M
Croptilon divaricatum (Nutt.) Rafin. WMJ
Dracopis amplexicaule (Vahl) Cass. W
Dyssodia tenuiloba (DC.) Small var. tenuiloba M
Echinacea atrorubens Nutt. W
Echinacea sanguinea Nutt. WM*J
Eclipta prostrata (L.) L. W
Elephantopus carolinianus Raeusch. WJ
Elephantopus tomentosus L. WMJ
Engelmannia peristenia (Rafin.) Goodman & Lawson WMJ
_ SYN= Engelmannia pinnatifida Gray ex Nutt.
Erechtites hieracifolia (L.) Rafin. ex DC. var. hieracifolia WJ
Erigeron annuus (L.) Pers. W
Erigeron philadelphicus L. W
Erigeron procumbens (Houst. ex P. Mill.) Nesom M
Erigeron strigosus Muhl. ex Willd. WMJ
Erigeron tenuis Torr. & Gray WM
Eupatorium capillifolium (Lam.) Small WJ
130 PHYTOLOGIA _ February 1998 volume 84(2): 107-153
Eupatorium compositifolium Walt. MJ
Eupatorium fistulosum Barr. J
Eupatorium lancifolium (T. & G.) Small WMJ
Eupatorium mohrii Greene M
Eupatorium perfoliatum L. WM
Eupatorium x pinnatifidum Ell. W
Eupatorium rotundifolium L. var. rotundifolium WMJ
Eupatorium semiserratum DC. WMJ
Eupatorium serotinum Michx. W
Eurybia hemispherica (Aiton) Nesom W
SYN= Aster paludosus subsp. hemisphericus (Alexander) Cronq.
SYN= Aster hemisphericus Alexander
Euthamia leptocephala (Torr. & Gray) Greene WMJ
Evax candida (Torr. & Gray) Gray W*J
SYN= Calymmandra candida Torr. & Gray
Evax verna Rafin. W*J
SYN= Evax multicaulis DC.
Facelis retusa (Lam.) Sch.-Bip. WMJ
Fleischmannia incarnata (Walt.) King & Rob. J
SYN= Eupatorium incarnatum Walt.
Gaillardia aestivalis (Walt.) Rock var. aestivalis J
Gaillardia pulchella Foug. WM
Gamochaeta falcata (Lam.) Cabrera W
SYN= Gnaphalium falcatum Lam.
Gamochaeta pensilvanica (Willd.) Cabrera W
SYN= Gnaphalium pensilvanica Willd.
Gamochaeta purpurea (L.) Cabrera WMJ
SYN= Gnaphalium purpureum L.
Grindelia lanceolata Nutt. var. lanceolata WJ
Helenium amarum (Rafin.) Rock WMJ
Helianthus angustifolius L. WJ
Helianthus annuus L. W
Helianthus debilis Nutt. OJ
Helianthus grosse-serratus Martens W
Helianthus hirsutus Rafin. WJ
Helianthus maximiliani Schrad. WM
*Helianthus mollis Lam. H
Helianthus simulans E. Wats. W
Heliopsis helianthoides (L.) Sweet J
Heterotheca subaxillaris (Lam.) Britt. & Rusby WMJ
SYN= Heterotheca latifolia Buckley
Hieracium gronovii L. WM
Hymenopappus artemisiifolius DC. var. artemisiifolius W
Hypochaeris microcephala (Sch.-Bip.) Cabrera var. albiflora (O. Kuntze) Cabrera
WJ
Iva angustifolia DC. W
Iva annua L. WJ
Krigia cespitosa (Rafin.) Chambers WMJ
Krigia dandelion (L.) Nutt. W
Krigia occidentalis Nutt. M
Nesom & Brown: Plant checklist for southeast Texas counties 131
Krigia virginica (L.) Willd. W*J
Krigia wrightii (A. Gray) Chambers ex Kim W*M (Kim & T 1992)
Lactuca canadensis L. WJ
Lactuca floridana (L.) Gaertn. WJ
Lactuca hirsuta Muhl. WJ
Lactuca ludoviciana (Nutt.) Ridd. W
Leucanthemum vulgare Lam. WM
SYN=Chrysanthemum leucanthemum L.
Liatris acidota Engelm. & Gray M*J
Liatris squarrulosa Michx. WMJ
SYN2= Liatris earlei (Walt.) K. Schum.
Liatris cymosa (Ness) K. Schum. W
Liatris elegans (Walt.) Michx. WMJ
Liatris mucronata DC. WJ
Liatris pycnostachya Michx. WMJ
Liatris squarrosa (L.) Michx. var. squarrosa WJ
Lindheimera texana Gray & Engelm. WM
Marshallia cespitosa Nutt. ex DC. var. signata Beadle & Boynt. W
Mikania scandens (L.) Willd. WJ
Ofigoneuron nitidum (Torr. & Gray) Small W(Taylor & Taylor 1984)
Oligoneuron rigidum (L.) Small W
SYN= Solidago rigida L.
Palafoxia hookeriana Torr. & Gray *J
Palafoxia rosea (Bush) Cory var. rosea WM(Turmer & Morris 1976)J
Parthenium hysterophorus L. W
Pityopsis graminifolia (Michx.) Nutt. W
SYN= Chrysopsis graminifolia (Michx.) Ell.
SYN= Heterotheca graminifolia (Michx.) Shinners
Pluchea camphorata (L.) DC. WJ
Pluchea foetida (L.) DC. WMJ
Pluchea odorata (L.) Cass. WMJ
SYN= Pluchea purpurascens (Sw.) DC.
Pseudognaphalium obtusifolium (L.) Hill. & Burtt
SYN= Gnaphalium obtusifolium L. WJ
Pterocaulon virgatum (L.) DC. WM
Pyrrhopappus carolinianus (Walt.) DC. WM
Pyrrhopappus pauciflorus (D. Don) DC. WMJ
Ratibida columnifera (Nutt.) Woot. & Standl. M*J
Rudbeckia fulgida Aiton J
Rudbeckia grandiflora (Sweet) DC. WMJ
Rudbeckia hirta L. [var.] WMJ
Rudbeckia maxima Nutt. WJ
Rudbeckia missouriensis Boynt. & Beadle WJ
SYNE R. fulgida var. missouriensis (Boynt. & Beadle) Cronq.
Rudbeckia nitida Nutt. var. texana Perdue *J
Senecio ampullaceus Hook. WJ
Senecio obovatus Muhl. ex Willd. WMJ
Senecio tampicanus DC. W
SYN= Senecio imparipinnatus Klatt
Silphium gracile A. Gray W*J
Silphium radula Nutt. W SYN?= Silphium asperrimum Hook.
132 PHYTOLOGIA _ February 1998 volume 84(2):107-153
Smallanthus uvedalia (L.) Mackenzie ex Small J
SYN= Polymnia uvedalia (L.) L.
Solidago altissima L. WMJ
SYN= Solidago canadensis var. scabra Torr. & Gray
Solidago delicatula Small WJ
SYN= Solidago ulmifolia var. microphylla A. Gray
Solidago ludoviciana (Gray) Small WMJ
Solidago nemoralis Ait. J
Solidago odora Aiton WJ
Solidago petiolaris Aiton W
Solidago radula Nutt. W(Taylor & Taylor 1984)
Solidago rugosa Aiton WMJ
Solidago speciosa var. rigidiuscula Torr. & Gray W(Taylor & Taylor 1984)
Soliva pterosperma (Juss.) Less. W*J
Sonchus asper (L.) Hill WM*J
Sonchus oleraceus L. W*J
Symphyotrichum divaricatum (Nutt.) Nesom
SYN= Aster subulatus var. ligulatus Shinners
Symphyotrichum drummondii var. texanum (Burg.) Nesom WJ
SYN= Aster drummondii var. texanus (Burg.) A.G. Jones
Symphyotrichum dumosum (L.) Nesom W
SYN= Aster dumosus L.
Symphyotrichum ericoides (L.) Nesom var. ericoides W*J
SYN= Aster ericoides L.
SYN= Virgulus ericoides (L.) Rev. & Keen.
Symphyotrichum lanceolatum (Willd.) Nesom W
SYN= Aster lanceolatus Willd.
Symphyotrichum lateriflorum (L.) Love & Love WMJ
SYN2= Aster lateriflorus (L.) Britt.
Symphyotrichum oolentangiense var. poaceum (Burg.) Nesom WJ
SYN2= Aster oolentangiensis var. poaceus (Burg.) A.G. Jones
Symphyotrichum patens (Aiton) Nesom var. gracile (Hook.) Nesom WJ
SYN= Aster patens var. gracilis Hook.
SYNE= Virgulus patens var. gracilis (Hook.) Rev. & Keen.
Symphyotrichum praealtum (Poir.) Nesom W
SYN= Aster praealtus Poir.
Symphyotrichum pratense (Rafin.) Nesom WJ
SYN= Aster pratensis Rafin.
SYN= Virgulus pratensis (Rafin.) Rev. & Keen.
Symphyotrichum racemosum (Ell.) Nesom WJ
SYN= Aster racemosus Ell.
SYN2= Aster fragilis Willd.
Taraxacum officinale Weber ex Wiggers W
Tetragonotheca ludoviciana (Torr. & Gray) Gray M
Tetraneuris linearifolia (Hook.) Greene var. linearifolia W
SYN= Hymenoxys linearifolia Hook.
Thelesperma filifolium (Hook.) Gray W
Verbesina encelioides (Cav.) A. Gray W
Verbesina virginica L. var. virginica WJ
Vernonia missurica Rafin. WM(mercer)J
Vernonia texana (A. Gray) Small WMJ
Nesom & Brown: Plant checklist for southeast Texas counties 133
Xanthium strumarium L. W
Youngia japonica (L.) DC. W
BERBERIDACEAE
Nandina domestica Thunb. WM
Podophyllum peltatum L. WMJ
BETULACEAE
Alnus serrulata (Ait.) Willd. J
Betula nigra L. WJ
Carpinus caroliniana Walt. WMJ
Ostrya virginiana (Mill.) K. Koch WJ
BIGNONIACEAE
Bignonia capreolata L. WMJ
Campsis radicans (L.) Seem. ex Bureau WMJ
Catalpa bignonioides Walt. *J
Catalpa speciosa (Warder) Warder ex Engelm. WJ
BORAGINACEAE
Heliotropium indicum L. WMJ
Heliotropium procumbens P. Mill. W
Heliotropium tenellum (Nutt.) Torr. WMJ
Lithospermum caroliniense (Gmel.) MacM. WM
Lithospermum incisum Lehm. W*J
Myosotis macrosperma Engelm. WMJ
Myosotis verna Nutt. W
Onosmodium bejariense DC. ex A. DC. WM
SYN= Onosmodium molle Michx. var. bejariense (DC. ex A. DC.) Cronq.
BRASSICACEAE
*Arabis canadensis L. 0
*Camelina microcarpa Andrz. 0
Capsella bursa-pastoris (L.) Medic. W
Cardamine bulbosa (Schreb. ex Muhl.) B.S.P. WMJ
Cardamine hirsuta L. WJ
Cardamine parviflora L. WMJ
Cardamine pensylvanica Muhl. ex Willd. M
Coronopus didymus (L.) J.E. Smith W
Draba brachycarpa Nutt. ex Torr. & Gray *J
*Draba cuneifolia Nutt. ex Torr. & Gray 0
*Lepidium densiflorum Schrad. 0
Lepidium virginicum L. WMJ
Lesquerella gracilis (Hook.) Wats. OM
Raphanus raphanistrum L. W
*Rorippa islandica (Oeder) Borbas 0
Rorippa nasturtium-aquaticum (L.) Hayek. W
Sibara virginica (L.) Roll. W
*Sisymbrium officinale (L.) Scop. 0
134 PHY-ELOLOGIA February 1998 volume 84(2):107-153
CABOMBACEAE
Brasenia schreberi Gmel. MJ(sight)
Cabomba caroliniana A. Gray W
CACTACEAE
* Opuntia engelmannii Engelm. var. lindheimeri (Engelm.) Parf. & Pink. 0
SYN= Opuntia lindheimeri Engelm.
Opuntia ficus-indica (L.) P. Mill. W
SYN= Opuntia compressa (Salisb.) Macbr.
* Opuntia macrorhiza Engelm. P
CALLITRICHACEAE
Calllitriche heterophylla Pursh WM
CAMPANULACEAE
Lobelia appendiculata A. DC. WM(mercer)*J
Lobelia cardinalis L. W*J
Lobelia puberula Michx. WJ
Lobelia reverchonii B.L. Turner M
Lobelia spicata Lam. WMJ
Triodanis biflora (R. & P.) Greene WJ
SYN= Triodanis perfoliata (L.) Nieuw. var. biflora (R. & P.) Bradley
Triodanis lamprosperma McVaugh W
Triodanis perfoliata (L.) Nieuw. WM*J
CANNABINACEAE
Cannabis sativa L. W
CAPPARACEAE
*Polanisia erosa (Nutt.) tis 0
CAPRIFOLIACEAE
Abelia grandiflora (Andre) Rehd. W
Lonicera japonica Thunb. WMJ
Lonicera sempervirens L. WJ
Sambucus canadensis L. WMJ
SYN= Sambucus nigra L. subsp. canadensis (L.) R. Bolli
Symphoricarpos orbiculatus Moench WJ
Viburnum acerifolium L. HMJ
Viburnum cassinoides L. J
SYN= Viburnum nudum L. var. cassinoides (L.) Torr. & Gray
SYN= Viburnum nitidum Ait.
Viburnum dentatum L. WMJ
Viburnum nudum L. WJ
Viburnum rufidulum Rafin. WMJ
CARYOPHYLLACEAE
Agrostemma githago L. W
Arenaria serpyllifolia L. W
Cerastium glomeratum Thuill. WJ
Nesom & Brown: Plant checklist for southeast Texas counties 135
Cerastium fontanum Baumg. var. vulgare (Hartman) Greuter & Burdet WM
SYN= Cerastium vulgatum L.
SYN= Cerastium holosteoides Fries
Loeflingia squarrosa Nutt. WJ
Minuartia drummondii (Shinners) McNeill W
SYN= Arenaria drummondii Shinners
Minuartia muscorum (Fassett) Rabeler W
SYN= Arenaria muriculata Maguire
SYN= Arenaria robusta Steyerm.
SYN= Arenaria patula var. robusta (Stey.) Maguire
Petrorhagia dubia (Rafin.) Lopez & Romo WJ (both from Rabeler 1985)
SYN= Petrorhagia velutina (Guss.) Ball & Heyw.
Polycarpon tetraphyllum (L.) L. W
Sagina decumbens (Ell.) Torr. & Gray W*J
Silene antirrhina L. W*J
Silene gallica L. WM*J
Spergula arvensis L. H
Stellaria media (L.) Vill. WJ
CELASTRACEAE
Euonymus americana L. WMJ
CHENOPODIACEAE
Chenopodium album L. W
Chenopodium ambrosioides L. WMJ
* Chenopodium berlandieri Mog. 0
Chenopodium pallescens Standley W
CISTACEAE
Helianthemum carolinianum (Walt.) Michx. WMJ
Helianthemum georgianum Chapm. W
Helianthemum rosmarinifolium Pursh W
Lechea mucronata Rafin. WJ
SYN= Lechea villosa Ell.
Lechea tenuifolia Michx. WM*J
CLUSIACEAE
Ascyrum hypericoides L. WMJ
SYN= Hypericum hypericoides (L.) Crantz
Ascyrum crux-andreae L. MJ
SYN= Ascyrum stans Michx. ex Willd.
SYN= Hypericum crux-andreae (L.) Crantz
Hypericum drummondii (Grev. & Hook.) Torr. & Gray WJ
Hypericum fasciculatum Lam. WMJ
Hypericum galioides Lam. *J
Hypericum gentianoides (L.) B.S.P. WMJ
Hypericum gymnanthum Eng. & Gray W
Hypericum mutilum L. WJ
Hypericwn prolificum L. WJ
Hypericum pseudomaculatum Bush 0*J
Hypericum punctatum Lam. W
136 PHYTOLOGIA _ February 1998 volume 84(2):107-153
Triadenum walteri (Gmel.) Gleason WJ
SYN= Hypericum walteri Gmel.
SYN= Hypericum petiolatum Walt.
CONVOLVULACEAE
Convolvulus equitans Benth. W
Dichondra carolinensis Michx. WMJ
Evolvulus sericeus Sw. W
Ipomoea indica (Burm. f.) Merr. W
SYN= Ipomoea acuminata (Vahl) R. & S.
Ipomoea cairica (L.) Sw. WM
Ipomoea hederacea Jacq. W
Ipomoea lacunosa L. J
ha pandurata (L.) Mey. WMJ
lpomoea cordatotriloba Dennst. WMJ
SYN= Ipomoea trichocarpa Ell.
Jacquemontia tamnifolia (L.) Griseb. WM
Stylisma humistrata (Walt.) Chapm. W
Stylisma pickeringii (Torr. ex Curtis) Gray W
CORNACEAE
Cornus drummondii C.A. Mey. WJ
Cornus florida L. WMJ
Cornus foemina P. Mill. J
CRASSULACEAE
Penthorum sedoides L. W
CUCURBITACEAE
Melothria pendula L. WJ
CUSCUTACEAE
Cuscuta compacta Juss. ex Choisy J
Cuscuta cuspidata Engelm. W
Cuscuta glabrior (Engelm.) Yunck. W
SYN= Cuscuta pentagona Engelm. var. glabrior (Yunck.) G.,T., & H.
Cuscuta glomerata Choisy *J
DROSERACEAE
Drosera brevifolia Pursh W
SYN= Drosera annua Reed
EBENACEAE
Diospyros virginiana L. WMJ
ERICACEAE
Rhododendron canescens (Michx.) Sweet W
Rhododendron oblongifolium (Small) Millais W*J
Rhododendron prinophyllum (Small) Millais *J
Vaccinium arboreum Marsh. WMJ
Vaccinium corymbosum L. WJ SYN= Vaccinium amoenum Aiton
Nesom & Brown: Plant checklist for southeast Texas counties 137
Vaccinium elliotii Chapm. OJ
Vaccinium stamineum L. WJ
Vaccinium virgatum Aiton WMJ
EUPHORBIACEAE
Acalypha gracilens A. Gray WMJ
*Acalypha ostryifolia Ridd. 0
Acalypha rhomboidea Rafin. WM
SYN= Acalypha virginica L. var. rhomboidea (Rafin.) Cooperr.
*Argythamnia humilis (Engelm. & Gray) Muell. Arg. P
Chamaesyce cordifolia (Ell.) Small W
SYN= Euphorbia cordifolia Ell.
Chamaesyce hypericifolia (L.) Millsp. W
_ SYN= Euphorbia hypericifolia L.
Chamaesyce maculata (L.) Small W
SYN= Euphorbia maculata L.
*Chamaesyce missurica (Rafin.) Shinners 0
SYN= Euphorbia missurica Rafin.
Chamaesyce nutans (Lag.) Small WMJ
SYN= Euphorbia nutans Lag.
Chamaesyce prostrata (Ait.) Small W
SYN= Euphorbia prostrata Ait.
Chamaesyce serpens (Kunth) Small WJ
SYN= Euphorbia serpens Kunth
Cnidosculus texanus (Muell. Arg.) Small WJ
Croton argyranthemus Michx. 0M
Croton capitatus Michx. var. lindheimeri (Engelm. & Gray) Muell. Arg. WMJ
Croton glandulosus L. var. septentrionalis Muell. Arg. WMJ
Croton monanthogynus Michx. WJ
*Croton texensis (K\.) Muell. Arg. 0
Croton willdenowii G. Webster WMJ
SYNE= Crotonopsis elliptica Willd.
Euphorbia cyathophora Murr. W
SYN= Poinsettia cyathophora (Murr.) K1. & Gke.
Euphorbia bicolor Engelm. & Gray WMJ
Euphorbia corollata L. WMJ
Euphorbia dentata Michx. WJ
Euphorbia discoidalis Chapman J
Euphorbia spathulata Lam. W*J
Euphorbia tetrapora Engelm. WJ
Phyllanthus caroliniensis Walt. WJ
Phyllanthus polygonoides Nutt. ex Spreng. WM
Phyllanthus urinaria L. WJ
Sapium sebiferum (L.) Roxb. WJ
Sebastiania fruticosa (Bartr.) Fern. J
Stillingia sylvatica L. WJ
Tragia betonicifolia Nutt. J
*Tragia cordata Michx. 0
Tragia urticifolia Michx. WJ
Vernicia fordii (Hemsl.) Airy-Shaw W
SYN= Aleurites fordii Hemsley
138 PHYTOLOGIA _ February 1998 volume 84(2):107-153
FABACEAE
Acacia angustissima (P. Mill.) Kuntze WM
SYN= Acacia hirta Nutt.
Acacia farnesiana (L.) Willd. WJ
Albizia julibrissin Durazz. WMJ
*Amorpha fruticosa L. H
Amorpha paniculata Torr. & Gray OJ
Amphicarpaea bracteata (L.) Fern. WM(Turner 1959)J
Apios americana Medic J
*Astragalus crassicarpus Nutt. 0
Astragalus distortus Torr. & Gray var. engelmannii (Sheld.) M.E. Jones WJ
Astragalus leptocarpus Torr. & Gray W
Astragalus nuttallianus var. trichocarpus Torr. & Gray OHW(Turner 1959)
Astragalus reflexus Torr. & Gray OW(Turner 1959)
*Astragalus soxmaniorum Lundell 0
Baptisia bracteata Muhl. ex Ell. *var. laevicaulis (Gray ex Canby) Isely 0
SYN= Baptisia laevicaulis (Gray) Small
Baptisia bracteata Muhl. ex Ell. var. leacophaea (Nutt.) Kartesz & Gandhi WM
SYN= Baptisia leucophaea Nutt.
Baptisia nuttalliana Small WMJ
Baptisia sphaerocarpa Nutt. W*J
Centrosema virginianum (L.) Benth. WMJ
Cercis canadensis L. WJ
Chamaecrista fasciculata (Michx.) E. Greene WMJ
SYN= Cassia fasciculata Michx.
Chamaecrista nictitans (L.) Moench WM(Turner 1959)J
SYN= Cassia nictitans L.
Clitoria mariana L. WMJ
Coronilla varia L. W
Crotalaria sagittalis L. WMJ
Dalea compacta Spreng. var. pubescens (Gray) Barneby WMJ
SYN= Dalea helleri Shinners
SYN= Petalostemum pulcherrimus (Heller) Heller
Dalea multiflora (Nutt.) Shinners WMJ
SYN= Petalostemum multiflorus Nutt.
Dalea phleoides (Torr. & Gray) Shinners WM(mercer)J (Turner 1959)
SYN= Petalostemum glandulosus Coult. & Fisher
Desmanthus illinoensis (Michx.) MacM. WMJ
Desmanthus leptolobus Torr. & Gray W(Luckow 1993)
Desmanthus velutinus Scheele W
Desmodium canescens (L.) DC. MJ
Desmodium ciliare (Muhl. ex Willd.) DC. WM(Turmer 1959)J
Desmodium glabellum (Michx.) DC. WMJ
Desmodium laevigatum (Nutt.) DC. WJ
Desmodium lineatum DC. WJ
Desmodium marilandicum (L.) DC. W
Desmodium nudiflorum (L.) DC. MJ
Desmodium nuttallii (Schindl.) Schub. W
Desmodium obtusum (Muhl. ex Willd.) DC. W
Desmodium paniculatum (L.) DC. WMJ
Nesom & Brown: Plant checklist for southeast Texas counties 139
Desmodium pauciflorum (Nutt.) DC. J
Desmodium rotundifolium DC. W(tex) J
Desmodium sessilifolium (Torr.) Torr. & Gray WM
*Desmodium tortuosum (Sw.) DC. 0
Desmodium viridiflorum (L.) DC. W
Erythrina herbacea L. WMJ
*Galactia erecta (Walt.) Vail 0
Galactia macreei M.A. Curtis WJ
Galactia regularis (L.) B.S.P. WJ
Galactia volubilis (L.) Britt. WM
Gleditsia triacanthos L. WMJ
Indigofera miniata Ort. WMJ
Indigofera suffruticosa P. Mill. WJ
Kummerowia striata (Thunb.) Schindl. WJ
' SYN= Lespedeza striata (Thunb.) Hook. & Arn.
Lathyrus hirsutus L. WM
Lathyrus pusillus Ell. WJ
Lespedeza bicolor Turcz. J
Lespedeza cuneata (Dumont) G. Don WJ
Lespedeza hirta (L.) Hornem. WM(Turner 1959)J
Lespedeza procumbens Michx. WJ
Lespedeza repens (L.) Bart. WMJ
Lespedeza stuevei Nutt. WM(Turner 1959)
Lespedeza virginica (L.) Britt. W
Lotus unifoliolatus (Hook.) Benth. W
SYN= Lotus purshianus (Benth.) Clem. & Clem.
Lupinus subcarnosus Hook. W*J
Lupinus texensis Hook. WM*J
Medicago arabica (L.) Hudson W
Medicago lupulina L. WJ
*Medicago minima (L.) L. 0
Medicago polymorpha L. WJ
SYN= Medicago hispida Gaertn.
Medicago sativa L. W
Melilotus albus Medic. W
Melilotus indicus (L.) All. WJ
Melilotus officinalis (L.) Lam. J
Mimosa hystricina (Small ex Britt. & Rose) Turner WMJ
SYN= Schrankia hystricina (Small) Stand.
Mimosa latidens (Small) Tumer M(Tumer 1959)
Mimosa microphylla Dry. WM
SYN= Mimosa uncinata (Willd.) Turner
SYN= Schrankia uncinata Willd.
Mimosa Strigillosa Torr. & Gray WJ
Neptunia lutea (Leavenw.) Benth. WMJ
Neptunia pubescens Benth. var. microcarpa (Rose) Windler W
Pediomelum subulatum (Bush) Rydb. OM(Ockendon 1965)
SYN= Pediomelum hypogaeum (Torr. & Gray) Rydb. var. subulatum (Bush)
Grimes
Prosopis glandulosa Torrey WJ
Psoralidium tenuiflorum (Pursh) Rydb. W(Turner 1959)
140 PHYTOLOGIA _ February 1998 volume 84(2):107-153
Puereria lobata (Willd.) Ohwi W
SYN= Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen & Alm.
Rhynchosia difformis (Ell.) DC. M(Turner 1959)
Rhynchosia latifolia Nutt. ex Torr. & Gray WM(Turner 1959)J
Rhynchosia minima (L.) DC. W*J
Robinia hispida L. W
Robinia pseudo-acacia L. WJ
Senna corymbosa (Lam.) Irwin & Barn. W
Senna marilandica (L.) Link WJ
Senna obtusifolia (L.) Irwin & Barn. M(Turner 1959)J
Senna occidentalis (L.) Link WM*J
Sesbania drummondii (Rydb.) Cory WJ
Sesbania herbacea (P. Mill.) McVaugh WJ
_ SYN= Sesbania macrocarpa Muhl. ex Rafin.
Sesbania vesicaria (Jacq.) Ell. WJ
Strophostyles helvola (L.) Ell. W
Strophostyles leiosperma (Torr. & Gray) Piper WM(Turner 1959)
Strophostyles umbellata (Muhl. ex Willd.) Britt. WM(Tumer 1959)
Stylosanthes biflora (L.) B.S.P. WMJ
Tephrosia onobrychoides Nutt. WM(Turmer 1959)
Tephrosia virginiana (L.) Pers. W*J
Trifolium bejariense Moric. W
Trifolium campestre Schreb. WM
Trifolium carolinianum Michx. W*J
Trifolium dubium Sibth. WMJ
Trifolium incarnatum L. WMJ
Trifolium lappulaceum L. W
*Trifolium polymorphum Poir. 0
SYN= Trifolium amphianthum Torr. & Gray
Trifolium pratense L. J
Trifolium repens L. WMJ
Trifolium resupinatum L. WJ
Trifolium vesiculosum Savi M
Vicia angustifolia L. W
SYN= Vicia sativa L. subsp. nigra (L.) Ehrh.
Vicia dasycarpa Ten. W
SYN= Vicia villosa Roth subsp. varia (Host) Corb.
Vicia leavenworthii Torr. & Gray W
SYN= Vicia ludoviciana Nutt. var. leavenworthii (Torr. & Gray) Lass. &
Gunn.
Vicia ludoviciana Nutt. WM
Vicia minutiflora Dietr. WM(Turner 1959)J
Wisteria frutescens (L.) Poir. J
SYN= Wisteria macrostachya Torr. & Gray
Wisteria sinensis (Sims) DC. WJ
FAGACEAE
Castanea pumila (L.) P. Mill. MJ
Castanea alnifolia Nutt. WJ
Fagus grandifolia Ehrh. MJ
Quercus alba L. WM(mercer)J
Nesom & Brown: Plant checklist for southeast Texas counties
Quercus falcata Michx. WM*J
Quercus incana Bartr. W
Quercus laurifolia Michx. WJ
Quercus lyrata Walt. W
Quercus margarettiae Ashe ex Small WJ
Quercus marilandica Muenchh. WMJ
Quercus michauxii Nutt. WJ
Quercus nigra L. W*J
Quercus pagoda Rafin. WJ
SYN= Quercus falcata Michx. var. pagodifolia EI.
Quercus phellos L. WJ
Quercus shumardii Buckl. WJ
Quercus similis Ashe WM
Quercus sinuata Walt. J
Quercus sinuata Walt. [form 2] W
Quercus stellata Wang. W*J
Quercus velutina Lam. WJ
Quercus virginiana P. Mill. WJ
FUMARIACEAE
Corydalis micrantha (Engelm. ex Gray) Gray WM*J
Fumaria offinialis L. M
GENTIANACEAE
Bartonia texana Correll *J
Centaurium pulchellum (Sw.) Druce W
Eustoma exaltatum (L.) Salisb. ex G. Don WJ
SYN= Eustoma grandiflorum (Rafin.) Shinners
Sabatia campestris Nutt. WM*J
Sabatia gentianoides Ell. M
GERANIACEAE
Geranium carolinianum L. WM(mercer)J
Geranium dissectum L. W
GROSSULARIACEAE
Itea virginica L. MJ
Ribes curvatum Small W
HALORAGACEAE
Myriophyllum aquaticum (Vell.) Verdcourt WM
SYN= Myriophyllum brasiliense Camb.
Myriophyllum heterophyllum Michx. J
Proserpinaca palustris L. M
HAMAMELIDACEAE
Hamamelis virginiana L. WJ
Liquidambar styraciflua L. WMJ
141
142 PHYTOLOGIA _ February 1998 volume 84(2): 107-153
HIPPOCASTANACEAE
Aesculus pavia L. J
HYDROPHYLLACEAE
Hydrolea ovata Nutt. ex Choisy WMJ
Nemophila aphylla (L.) Brumm. WMJ
SYN= Nemophila microcalyx (Nutt.) Fisch. & Mey.
Nemophila phacelioides Nutt. WJ _
Phacelia glabra Nutt. W
Phacelia hirsuta Nutt. W
Phacelia patuliflora (Engelm. & Gray) Gray M
JUGLANDACEAE
Carya alba (L.) Nutt. ex Ell. WMJ
‘ SYN= Carya tomentosa (Lam. ex Poir.) Nutt.
Carya aquatica (Michx. f.) Nutt. WMJ
Carya illinoiensis (Wang.) K. Koch W
Carya myristiciformis (Michx. f.) Nutt. WJ
Carya ovata (P. Mill.) K. Koch W
Carya texana Buckl. WJ
Juglans nigra L. WMJ
KRAMERIACEAE
Krameria lanceolata Torrey W
LAMIACEAE
Ajuga reptans L. W(escaping cult)
Clinopodium brownei (Sw.) Kuntze W
SYN= Micromeria brownei (Sw.) Benth.
Hedeoma drummondii Benth. 0M
Hedeoma hispidum Pursh WM
Hyptis alata (Rafin.) Shinners J
Lamium amplexicaule L. W
Lamium purpureum L. W
*Leonotis nepetifolia (L.) Ait. f. 0
Lycopus rubellus Moench WJ
Lycopus virginicus L. *J
*Mentha piperita L. 0
*Mentha spicata L. 0
Monarda citriodora Cerv. ex Lag. WM*J
Monarda fistulosa L. W*J
Monarda lindheimeri Gray M(mercer)
Monarda punctata L. WMJ
Perilla frutescens (L.) Britt. J
Physostegia angustifolia Fern. M(tex)
Physostegia digitalis Small WMJ
Physostegia intermedia (Nutt.) Engelm. & Gray W
Physostegia pulchella Lundell WJ
Prunella vulgaris L. OHJ
Pycnanthemum albescens Torr. & Gray WMJ
Pycnanthemum tenuifolium Schrad. 0*J
Nesom & Brown: Plant checklist for southeast Texas counties
Rhododon ciliatus (Benth.) Epling M(Turner 1995)
Salvia azurea Michx. ex Lam. WJ
Salvia coccinea Buchoz ex Etlinger 0
Salvia lyrata L. WMJ
Scutellaria cardiophylla Engelm. & Gray WJ
Scutellaria drummondii Benth. var. dr'ummondii MJ
*Scutellaria elliptica Muhl. ex Spreng. 0
Scutellaria integrifolia L. WM*J
Scutellaria racemosa Pers. M
Scutellaria ovata Hill J
Scutellaria parvula Michx. WJ
Stachys crenata Rafin. WJ
Stachys floridana Shuttlew. W
Stachys tenuifolia Willd. WM
Teucrium canadense L. var. canadense W
Trichostema dichotomum L. WM*J
Warnockia scutellarioides (Engelm. & Gray) M. Turner W
SYN= Brazoria scutellarioides Engelm. & Gray
LAURACEAE
Cinnamomum camphora (L.) J. Pres! W
Persea borbonia (L.) Spreng. WMJ
Persea palustris (Rafin.) Sarg. J
Sassafras albidum (Nutt.) Nees WMJ
LENTIBULARIACEAE
Utricularia gibba L. WM
Utricularia radiata Small M
Utricularia subulata L. W
LINACEAE
Linum berlandieri Hook. var. berlandieri WM
SYN= Linum rigidum Pursh var. berlandieri (Hook.) Torr. & Gray
Linum medium (Planch.) Britt. var. texanum (Planch.) Fern. WMJ
Linum striatum Walt. WMJ
LOGANIACEAE
Mitreola petiolata (Gmel.) Torr. & Gray WJ
SYN= Cynoctonum mitreola (L.) Britt.
Gelsemium sempervirens (L.) Jaume St.-Hill WMJ
Polypremum procumbens L. WM*J
Spigelia marilandica (L.) L. *J
LYTHRACEAE
Ammannia coccinea Rottb. WM
Cuphea carthagenensis (Jacq.) Macbr. MJ
Cuphea glutinosa C. & S. J
Laegerstroemia indica L. WM
Lythrum alatwn Pursh var. lanceolatum (Fll.) Torr. & Gray ex Rothr. WJ
Rotala ramosior (L.) Koehne W
143
144 PHYTOLOGIA _ February 1998 volume 84(2):107-153
MAGNOLIACEAE
Liriodendron tulipiferaL. M
Magnolia grandiflora L. WMJ
Magnolia pyramidata Bartr. *J
Magnolia virginiana L. OMJ
MALVACEAE
Abutilon theophrasti Medic. W
Callirhoe involucrata (Torr.) A. Gray W
*Callirhoe leiocarpa R.F. Martin 0
Hibiscus laevis Allioni WM
SYN= Hibiscus militaris Cav.
Hibiscus moscheutos L. subsp. lasiocarpus (Cav.) Blanch. MW
‘ SYN= Hibiscus lasiocarpus Cav.
Malvaviscus arboreus Cav. var. drummondii (T. & G.) Schery WMJ
SYN= Malvaviscus drummondii T. & G.
Modiola caroliniana (L.) G. Don W
Sida ciliaris L. W.
Sida rhombifolia L. WMJ
Sida spinosa L. W
Sphaeralcea angustifolia (Cav.)G. Don M
MELASTOMATACEAE
Rhexia mariana L. var. mariana M
MELIACEAE
Melia azederach L. W*J
MENISPERMACEAE
*Calycocarpum lyonii (Pursh) Gray H
Cocculus carolinus (L.) DC. WJ
MOLLUGINACEAE
Mollugo verticillata L. WMJ
MONOTROPACEAE
Monotropa uniflora L. H*J
MORACEAE
Broussonetia papyrifera (L.) Vent. W
Fatoua villosa (Thunb.) Nakai J
Maclura pomifera (Rafin.) Schneid. WJ
Morus alba L. W
Morus rubra L. WJ
MYRICACEAE
Morella cerifera (L.) Small WMJ
SYN= Myrica cerifera L.
SYN= Myrica pusilla Rafin.
Nesom & Brown: Plant checklist for southeast Texas counties 145
Morella caroliniensis (P. Mill.) Small OJ
SYN= Myrica caroliniensis P. Mill.
SYN= Myrica heterophylla Rafin.
NYMPHAEACEAE
Nuphar lutea (L.) Sm. subsp. advena (Ait.) Kart. & Gandhi WM
SYN= Nuphar lutea (L.) Sm. subsp. macrophylla (Small) Beal
Nymphaea elegans Hook. W
Nymphaea odorata Ait. M(mercer)
NELUMBONACEAE
Nelumbo lutea Willd. M
NYCTAGINACEAE
*Boerhaavia erecta L. 0
Mirabilis jalapa L. W
Mirabilis gigantea (Standl.) Shinners W
Mirabilis albida (Walt.) Heimerl W(Turner 1993)
NYSSACEAE
Nyssa aquatica L. M
Nyssa biflora Walt. J
Nyssa sylvatica Marsh. WJ
OLEACEAE
Chionanthus virginicus L. WMJ
Forestiera ligustrina (Michx.) Poir. WM(Brooks 1977)*J
Fraxinus americana L. WJ
Fraxinus pennsylvanica Marsh. WMJ
Ligustrum japonicum Thunb. W
Ligustrum lucidum Aiton f. WJ
Ligustrum quihoui Carr. W
Ligustrum sinense Lour. WJ
ONAGRACEAE
Calylophus berlandieri Spach subsp. berlandieri MJ
SYN= Calylophus drummondianus Spach var. berlandieri (Spach) Towner &
Raven
Calylophus serrulatus (Nutt.) Raven M(Towner 1977)
Gaura brachycarpa Small W
Gaura lindheimeri Engelm. & Gray J
Gaura longiflora Spach WJ
SYN= Gaura filiformis Small
Gaura sinuata Ser. W*J
Ludwigia alternifolia L. WMI
Ludwigia decurrens Walt. WJ
Ludwigia glandulosa Walt. WJ
Ludwigia hirtella Rafin. WM
Ludwigia leptocarpa (Nutt.) Hara W
Ludwigia octovalvis (Jacq.) Raven WJ
Ludwigia palustris (L.) Ell. WJ
146 PHYTOLOGIA February 1998 volume 84(2): 107-153
Ludwigia peploides (Jacq.) Raven WM
Ludwigia pilosa Walt. J
Ludwigia sphaerocarpa Ell. M
Oenothera heterophylla Spach J
Oenothera laciniata Hill WMJ
Oenothera'linifolia Nutt. W
Oenothera rhombipetala T. & G. J
Oenothera spachiana Torr. & Gray M
Oenothera speciosa Nutt. WMJ
Oenothera triloba Nutt. W
Stenosiphon linifolius (Nutt. ex James) Heynh. W
OROBANCHACEAE
*Orobanche uniflora L. H
OXALIDACEAE
Oxalis corniculata L. var. wrightii (Gray) Turner WMJ
SYN= Oxalis dillenii Jacq. (sensu B.L. Turner 1994)
Oxalis corymbosa DC. WJ
SYN= Oxalis debilis Kunth var. corymbosa (DC.) Lourteig
Oxalis lyonii Pursh WM*J
SYN= Oxalis priceae Small
Oxalis rubra St. Hil. W
Oxalis violacea L. W*(tex) M(tex)
PAPAVERACEAE
Argemone albiflora Hornem. subsp. texana G. Ownbey WM
Sanguinaria canadensis L. *J
PASSIFLORACEAE
Passiflora incamata L. WMJ
Passiflora lutea L. WJ
Passiflora sp. M
PHYTOLACCACEAE
Phytolacca americana L. WMJ
PLANTAGINACEAE
Plantago aristata Michx. WMJ
Plantago elongata Pursh W
*Plantago hookeriana Fisch. & Mey. 0
Plantago major L. W
Plantago rhodosperma Dene. W
Plantago virginica L. WJ
PLATANACEAE
Platanus occidentalis L. WMJ
POLEMONIACEAE
Ipomopsis rubra (L.) Wherry WM(Wherry 1966)J
Phlox cuspidata Scheele WM(Wherry 1966)J
Nesom & Brown: Plant checklist for southeast Texas counties
Phlox drummondii Hook. WMJ
Phlox pilosa L. WMJ
POLYGALACEAE
Polygala mariana P. Mill. WM
Polygala polygama Walt. WM
Polygala verticillata L. WJ
POLYGONACEAE
Brunnichia ovata (Walt.) Shinners W
SYN= Brunnichia cirrhosa Gaertn.
*Eriogonum longifolium Nutt. 0
*Polygonella americana (Fisch. & Mey.) Small 0
*Polygonum aviculare L. 0
Polygonum hydropiperoides Michx. WM
Polygonum lapathifolium L. W
*Polygonum pennsylvanicum L. 0
Polygonum persicaria L. W
Polygonum punctatum Ell. WJ
*Polygonum ramosissimum Michx. 0
*Polygonum tenue Michx. 0
Rumex altissimus Wood M(Rechinger 1937)
Rumex crispus L. W
Rumex hastatulus Baldw. WMJ
Rumex pulcher L. WM
Tovara virginiana (L.) Rafin. W
SYN= Polygonum virginianum L.
PONTEDERIACEAE
*Pontederia cordata L. H
PORTULACACEAE
Claytonia virginica L. WMJ
Portulaca oleracea L. W
Portulaca pilosa L. WJ
*Portulaca umbraticola Kunth 0
Talinum parviflorum Nutt. W
PRIMULACEAE
Anagallis arvensis L. WMJ
Centunculus minimus L. WJ
SYN= Anagallis minimus (L.) Krause
Hottonia inflata Ell. W
Samolus parviflorus Rafin. WMJ
SYN= Samolus valerandi L. subsp. parviflorus (Rafin.) Hulten
RANUNCULACEAE
Anemone berlandieri Pritzel W
Anemone caroliniana Walt. WJ
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148 PHYTOLOGIA _ February 1998 volume 84(2):107-153
Clematis crispa L. J
Clematis pitcheri Torr. & Gray *J
Clematis reticulata Walt. WMJ
Clematis ternifolia DC. W
SYN= Clematis dioscoreifolia Lev\. & Van.
Delphinium ajacis L. W
Delphinium carolinianum Walt. subsp. vimineum (D. Don) M.J. Warnock W
SYN= Delphinium vimineum D. Don
Ranunculus hispidus Michx. var. nitidus (Chapm.) T. Duncan M*J
SYN= Ranunculus carolinianus DC.
Ranunculus fascicularis Muhl. ex Bigelow WMJ
Ranunculus macranthus Scheele W
Ranunculus muricatus L. WM
Ranunculus parviflorus L. W
Ranunculus pusillus Poir. W
Ranunculus sardous Crantz WM
Ranunculus sceleratus L. W
Thalictrum dasycarpum Fisch. & Ave-Lall. *J
Xanthorhiza simplicissima Marsh. *J
RHAMNACEAE
Berchemia scandens (Hill) K. Koch WMJ
* Ceanothus americanus L. 0H
* Ceanothus herbaceus Rafin. 0
Frangula caroliniana (Walt.) Gray WMJ
SYN= Rhamnus caroliniana Walt.
ROSACEAE
Agrimonia microcarpa Wallr. WJ
Agrimonia rostellata Wallr. J
Crataegus brachyacantha Sarg. & Engelm. W
Crataegus crus-galli L. WJ
Crataegus engelmannii Sarg. W*J
Crataegus marshallii Egg]. WMI
Crataegus spathulata Michx. WMJ
Crataegus viridis L. WMJ
* Crataegus warneri Sarg. W(Sargent 1922)
Duchesnea indica (Andrz.) Focke WJ
Geum canadense Jacq. WMJ
Photinia arbutifolia (Lam.) Robertson & Phipps JM
SYN= Pyrus arbutifolia (Lam.) Lf.
Potentilla simplex Michx. H
Prunus angustifolia Marsh. W
Prunus caroliniana (P. Mill.) Ait. WM(mercer)*J
Prunus gracilis Engelm. & Gray WM
Prunus mexicana S. Wats. WMJ
Prunus serotina Ehrh. WMJ
Prunus umbellata Ell. WMJ
Rosa bracteata Wendl. W
Rosa carolina L. WM
Rosa laevigata Michx. *J
Nesom & Brown: Plant checklist for southeast Texas counties
Rosa setigera Michx. OM
Rubus argutus Link WMJ
SYN= Rubus louisianicus Berger
Rubus arvensis Bailey W
SYN= Rubus saepescandens Bailey
Rubus trivialis Michx. WJ
RUBIACEAE
Cephalanthus occidentalis L. WMJ
Diodia teres Walt. WM*J
Diodia virginiana L. WM
Galium aparine L. W*J
Galium pilosum Aiton WMJ
Galium tinctorium (L.) Scop. WMJ
Galium uniflorum Michx. W
Galium virgatum Nutt. W*J
Hedyotis nigricans (Lam.) Fosb. WJ
Houstonia micrantha (Shinners) Terrell W*J
SYN= Hedyotis australis Lewis & Moore
SYN= Hedyotis crassifolia Raf. var. micrantha Shinners
Houstonia pusilla Schoepf WMJ
SYN= Houstonia minima Beck
SYN= Houstonia patens Ell.
SYN= Hedyotis crassifolia Rafin.
Houstonia rosea (Rafin.) Terrell *J
SYN= Hedyotis rosea Rafin.
SYN= Hedyotis tayloriae Fosb.
SYN= Houstonia pygmaea C.H. & M.T. Mueller
Mitchella repens L. WJ
Oldenlandia boscii (DC.) Chapm. W
Oldenlandia uniflora L. WJ
Richardia scabra L. WJ
Sherardia arvensis L. WM
Spermacoce glabra Michx. W
RUTACEAE
Poncirus trifoliata (L.) Rafin. WMJ
SYN= Citrus trifoliata L.
Ptelea trifoliata L. WJ
Zanthoxylum clava-herculis L. WMJ
SALICACEAE
Populus deltoides Bartr. ex Marsh. WJ
Salix nigra Marsh. WM
SAPINDACEAE
Cardiospermum halicacabum L. W
Koelreuteria paniculata Laxm. W
Sapindus drummondit Hook. & Arn. WJ
SYN= Sapindus saponaria L. var. drummondii (Hook. & Arn.) L. Benson
*Ungnadia speciosa Endl. 0
149
150 PHYTOLOGIA _ February 1998 volume 84(2):107-153
SAPOTACEAE
Sideroxylon lanuginosum Michx. WMJ
SYN= Bumelia lanuginosa (Michx.) Pers.
SAURURACEAE
Saururus cernuus L. WMJ
SAXIFRAGACEAE
Lepuropetalon spathulatum Ell. H
Saxifraga texana Buckl. W
SCROPHULARIACEAE
Agalinis fasciculata (Ell.) Rafin. WM
Agalinis gattingeri (Small) Small W
Agalinis heterophylla (Nutt.) Small ex Britt. WJ
Agalinis purpurea (L.) Penn. W*J
Agalinis viridis (Small) Penn. MJ
Aureolaria virginica (L.) Pennell J
SYN= Aureolaria dispersa (Small) Penn.
Aureolaria grandiflora (Benth.) Penn. MJ
Bacopa caroliniana (Walt.) B. Robins. M
Bacopa monnieri (L.) Pennell WMJ
Bellardia trixago (L.) Allm. M(Do et al. 1996)
Castilleja indivisa Engelm. WM*J
Gratiola brevifolia Rafin. HMJ
Gratiola neglecta Torrey W
Gratiola virginiana L. WJ
Lindernia anagallidea (Michx.) Penn. W
Mecardonia acuminata (Walt.) Small *J
Mecardonia procumbens (P. Mill.) Pennell W*J
SYN= Mecardonia vandellioides auct. non (Kunth) Pennell.
Micranthemum umbrosum (Gmel.) Blake WJ
Mimulus alatus Ait. WJ
Nuttallanthus canadensis (L.) D.A. Sutton WJ
SYN2= Linaria canadensis (L.) Don
Nuttallanthus texanus (Scheele) D.A. Sutton WM*J
SYN= Linaria texana Scheele
Pedicularis canadensis L. WJ
Penstemon cobaea Nutt. W
Penstemon digitalis Nutt. ex Sims W
Penstemon laxiflorus Penn. WMJ
SYN= Penstemon australis Small subsp. laxiflorus (Penn.) Bennett
Penstemon murrayanus Hook. OJ
Scoparia dulcis L. WMJ
Verbascum blattaria L. W
Verbascum thapsus L. WJ
Verbascum virgatum Stokes W.
Veronica agrestis L. H
Veronica arvensis L. W*J
Veronica peregrina L. WJ
Veronica persica Poir. M
Nesom & Brown: Plant checklist for southeast Texas counties
SOLANACEAE
Callibrachoa parviflora (Juss.) D'Arcy W
SYN= Petunia parviflora Juss.
Capsicum annuum L. W
*Datura stramonium L. 0
*Physalis angulata L. 0
Physalis heterophylla Nees WJ
*Physalis pubescens L. 0
*Physalis pumila Nutt. 0
* Physalis virginiana Mill. 0
Physalis viscosa L. var. cinerascens (Dun.) Waterf. WJ
Solanum carolinense L. WMJ
Solanum dimidiatum Rafin. WJ
Solanum elaeagnifolium Cav. W
Solanum lycopersicum (L.) Karst. ex Farw. W
SYN= Lycopersicon esculentum P. Mill.
Solanum pseudocapsicum L. J
Solanum ptycanthum Dun. ex DC. WMJ
SYN= Solanum nigrum L. auct.
SYN= Solanum americanum Mill. auct.
Solanum rostratum Dunal OMJ
SPHENOCLEACEAE
Sphenoclea zeylandica Gaertn. W
STERCULIACEAE
Firmiana simplex (L.) W. Wight W
STYRACACEAE
Halesia diptera Ellis WMJ
Styrax americanus Lam. WJ
SYMPLOCACEAE
Symplocos tinctoria (L.) L'Herit. MJ
TILIACEAE
Tilia americana L. WMJ
SYNE Tilia caroliniana P. Mill.
SYNE= Tilia americana L. var. caroliniana (P. Mill.) Castigl.
ULMACEAE
Celtis laevigata Willd. WMJ
Celtis tenuifolia Nutt. W
Planera aquatica Gmel. W
Ulmus alata Michx. WJ
Ulmus americana L. WMJ
Ulmus crassifolia Nutt. WMJ
Ulmus rubra Muhl. W
151
£52 PHYTOLOGIA _ February 1998 volume 84(2):107-153
URTICACEAE
Boehmeria cylindrica (L.) Sw. WJ
Parietaria pensylvanica Muhl. ex Willd. W
*Pilea pumila (L.) Gray 0
*Urtica chamaedryoides Pursh 0
*Urtica urens L. 0
VALERIANACEAE
Valerianella amarella (Lindh. ex Engelm.) Krok. WJ
Valerianella radiata (L.) Dufr. WM
Valerianella stenocarpa (Engelm. ex Gray) Krok W*J
VERBENACEAE
Callicarpa americana L. WMJ
Glandularia bipinnatifida (Nutt.) Nutt. W
SYN= Verbena bipinnatifida Nutt.
Glandularia canadensis (L.) Nutt. WM
SYN= Verbena canadensis (L.) Britt.
Glandularia pulchella (Sweet) Tronc. MJ
SYN= Verbena tenuisecta Briq.
Lantana camara L. WJ
Phyla nodiflora (L.) E. Greene WM*J
Phyla lanceolata (Michx.) Greene HM
Phryma leptostachya L. WJ
Verbena bonariensis L. W .
Verbena brasiliensis Vell. WMJ
Verbena halei Small WMJ
Verbena rigida Spreng. WMJ
Verbena urticifolia L. W
Verbena xutha Lehm. WMJ
Vitex agnus-castus L. W
VIOLACEAE
Viola affinis Le Conte WM*J
SYNE= Viola sororia Willd. var. missouriensis (Greene) McKinney
SYN= Viola missouriensis Greene
SYN= Viola langloisii Greene
Viola bicolor Pursh W
SYN= Viola rafinesquii Greene
Viola lanceolata L. var. lanceolata *J
Viola lanceolata L. var. vittata (Greene) Russell M
SYN= Viola vittata Greene
Viola palmata L. *J
SYNE= Viola triloba Schwein. var. dilatata (Ell.) Brainerd
Viola pedata L. W*J
Viola primulifolia L. J
Viola sagittata Ait. WJ
SYN= Viola triloba Schwein. var. triloba
Viola walteri House WJ
Nesom & Brown: Plant checklist for southeast Texas counties 153
VISCACEAE
Phoradendron tomentosum (DC.) A. Gray W*J
SYN= Phoradendron serotinum (Raf.) M.C. Johnston var. pubescens
(Engelm.) M.C. Johnston :
VITACEAE
Ampelopsis arborea (L.) Koehne WJ
Ampelopsis cordata Michx. W
Cissus trifoliata (L.) L. W
SYN= Cissus incisa auct. non Des Moulins
Parthenocissus quinquefolia (L.) Planch. WMJ
Vitis aestivalis Michx. var. aestivalis W*J
Vitis aestivalis Michx. var. lincecumii (Buck].) Munson W
_ SYNE= Vitis lincecumii Buckley
Vitis cinerea (Engelm.) Millard WJ
Vitis mustangensis Buckley WJ
Vitis palmata Vahl WJ
Vitis rotundifolia Michx. WMJ
Vitis vulpina L. WJ
ZYGOPHYLLACEAE
*Tribulus terrestris L. O
REJECTED or DOUBTFUL
Calylophus serrulata (Nutt.) Raven 0, probably = Calylophus drummondii
Croton floridanus Ferguson 0, =?
Crotonopsis linearis Michx. 0, probably = Croton willdenowii
Forestiera pubescens Nutt. H, probably = Forestiera ligustrina
Gutierrezia texana (DC.) Torr. & Gray 0, probably = Amphiachyris
dracunculoides
Helianthus cucumerifolius T. & G. 0, = Helianthus debilis
Oenothera jamesii T. & G. 0, =?
Palafoxia callosa (Nutt.) T.& G. P, probably = Palafoxia rosea
Parthenium hispidum Rafin. 0, probably = Parthenium hysterophorus
Rhynchospora mixta Britt. ex Small H, =?
Rumex mexicanus Meisn. 0, = ?
Rumex obtusifolius L. 0, probably = Rumex pulcher
Sagittaria lancifolia L. H = perhaps S. platyphylla (Engelm.) J.G. Sm.
Tradescantia subacaulis Bush H, =?
Tridens muticus (Torr.) Nash 0, probably = Tridens strictus
Yucca arkansana Trel. OH, probably = Yucca louisianensis
Phytologia (February 1998) 84(2):154-164.
NOTES ON THE DISTRIBUTION, HOST RANGE, PLANT SIZE, PHENOLOGY,
AND SEX RATIO OF TWO RARE DWARF MISTLETOES FROM CENTRAL
AMERICA: ARCEUTHOBIUM HAWKSWORTHII AND A. HONDURENSE
Robert L. Mathiasen', Catherine G. Parks”, Brian W. Geils’ & Jerome S. Beatty*
'School of Forestry, Northern Arizona University, Flagstaff, Arizona 86011 U.S.A.
Pacific Northwest Research Station, USDA Forest Service, Forest Sciences
Laboratory, 1401 Gekeler Lane, LaGrande, Oregon 97850 U.S.A.
’Rocky Mountain Research Station, USDA Forest Service, 2500 South Pine Knoll
Drive, Flagstaff, Arizona 86001 U.S.A.
‘Forest Insects and Diseases, Pacific Northwest Region, USDA Forest Service, 16400
Champion Way, Sandy, Oregon 97055 U.S.A.
ABSTRACT
New information on the distribution, host range, plant size, flowering
periods, and sex ratio of Arceuthobium hawksworthii and A. hondurense are
provided based on plant collections, field-observations and roadside-surveys
conducted in 1998 in Belize, Guatemala, and Honduras, Central America.
Arceuthobium hawksworthii parasitizes Pinus caribaea var. hondurensis and
P. oocarpa vat. ochoterenai in Belize, but the latter pine is less susceptible and
classified here as a secondary host. Our surveys in the P. caribaea var.
hondurensis forests near Poptun, Guatemala indicate that A. hawksworthii
does not occur there, and therefore, this mistletoe probably does not occur in
Guatemala. Our surveys of the Mountain Pine Ridge Region of western Belize
indicate that A. hawksworthii occurs there at elevations between 520-900 m
and is distributed in an area of approximately 250 sq. km. Therefore, this
dwarf mistletoe probably has the most restricted geographic range of any
known dwarf mistletoe. No new populations of A. hondurense were
discovered in Honduras, but our examination of previously reported
populations of this dwarf mistletoe indicates that it only parasitizes P. oocarpa
vars. oocarpa and ochoterenai and not P. maximinoi as previously reported.
Additional plant measurements of A. hawksworthii and A. hondurense are
reported, including new maximum records of plant height for both species.
The sex ratio of both dwarf mistletoes is 1:1, but the sample size for A.
154
Mathiasen et al.: | Summary of two Arceuthobium in Central America 155
hondurense was small (64 plants) and additional sampling will be needed to
confirm this finding.
KEY WORDS: Hawksworth’s dwarf mistletoe, Honduran dwarf mistletoe,
Caribbean pine
BACKGROUND
Dwarf mistletoes (Arceuthobium spp., Viscaceae) are parasitic flowering plants
that parasitize members of the Pinaceae in the New World and Pinaceae and
Cupressaceae in the Old World. Although the dwarf mistletoes are widespread and
common parasites of the Pinaceae in the United States and México, there are only a
few taxa known from Central America. Little is known regarding the distribution,
host range, ecology, and phenology of the Central American taxa (Hawksworth &
Wiens 1996).
Two of the rarest dwarf mistletoes known occur in Belize and Honduras:
Arceuthobium hawksworthii Wiens & C.G. Shaw, III and A. hondurense Hawksw.
& Wiens. Arceuthobium hawksworthii is known only from the Mountain Pine Ridge
Region (MPR) of western Belize (Wiens & Shaw 1994) and A. hondurense is only
known from two areas of Honduras (Hawksworth & Wiens 1972, 1996).
Hawksworth & Wiens (1996) speculated that A. hondurense may be so rare in
Honduras that it might be in danger of extinction if the extensive harvesting of pine
forests in Honduras continues.
Specimens of Arceuthobium hawksworthii have been collected by various
investigators from the MPR since 1959, but this taxon was not described until recently
(Wiens & Shaw 1994). It was previously considered to represent a disjunct
population of Arceuthobium globosum Hawksw. & Wiens (Hawksworth & Wiens
1972) and later was classified as A. aureum Hawksw. & Wiens subsp. aureum
(Hawksworth & Wiens 1977). However, a detailed analysis of the Belizean
populations by Wiens & Shaw (1994) indicated these populations are distinct from the
A. aureum subsp. aureum populations found in the western and central highlands of
Guatemala. Wiens & Shaw (1994) speculated that A. hawksworthii might occur in
eastern Guatemala in a disjunct population of its principal host, Pinus caribaea Morelet
var. hondurensis (Senecl.) Barr. & Golf., that occurs in the vicinity of Poptun (Perry
1991). They also indicated that they did not observe A. hawksworthii below an
elevation of 700 m in the MPR although previous reports place it as low as 520 m in
this Region. Specimens of A. hawksworthii collected by A. Carty at high elevations
(above 900 m) near Cooma Cairn in 1981 were tentatively identified as hybrids of
Pinus caribaea var. hondurensis and Pinus oocarpa Schiede var. ochoterenai Mart.
Wiens & Shaw (1994) reported they were unable to confirm if A. hawksworthii was
parasitizing Pinus oocarpa var. ochoterenai or hybrids of these species at this location.
Arceuthobium hondurense was described in 1970 from populations southeast of
Tegucigalpa, Honduras, but only one additional population of this rare dwarf mistletoe
has been discovered since then (Hawksworth & Wiens 1970, 1972, 1984, 1996). It
is presently known from only two general localities in Honduras; in the Piedra Herrada
156 PHYTOLOGIA February 1998 volume 84(2):154-164
Mountains, approximately 22 km southeast of Tegucigalpa and from approximately 20
km northwest of San Pedro Sula in Cusuco National Park (Hawksworth & Wiens
1996). Thus far, this dwarf mistletoe has only been reported parasitizing Pinus
oocarpa var. oocarpa and P. maximinoi H.E. Moore (Hawksworth & Wiens 1996).
So little is known about its host range and distribution in Central America that it is
considered to be the rarest dwarf mistletoe described from the New World. It may
also occur in El Salvador in the vicinity of Montecristo (Santa Ana Province) near the
borders of Honduras and Guatemala, but this tentative report (Hawksworth & Wiens
1996) needs to be confirmed.
SAMPLING AND FIELD OBSERVATIONS
Distribution
From March 3--4, May 29--June 1, and August 18--20, 1998 we conducted
roadside-surveys in the pine forests of the MPR of western Belize. Arceuthobium
hawksworthii is common in the MPR east of Augustine, particularly along the Baldy
Beacon, Hidden Valley Falls, Brunton Trail, Orchid Hill, and Granite Cairn Roads.
We observed populations of A. hawksworthii at an elevation of 520 m at several
locations, but the mistletoe is most common above 600 m. The reasons for the lower
elevational limitation are unclear, because its pine hosts are continuously distributed
throughout the MPR at elevations below 500 m. In addition, as was reported by
Wiens & Shaw (1994) we did not observe A. hawksworthii in the lower elevation
Pinus caribaea var. hondurensis forests closer to the coast of Belize. Our surveys now
provide adequate information to estimate the distribution of this rare dwarf mistletoe in
the MPR. Because we did not find populations of A. hawksworthii below 520 m in
the MPR, we used this lower elevational limit and our roadside-survey observations to
estimate the approximate distribution of A. hawksworthii (Figure 1); it only occurs in
an area of approximately 250 sq. km. Because the highlands of the MPR are
geologically distinct and have evidently been isolated from other highland regions of
Central America for several thousand years (Bateson 1972; Bateson & Hall 1977;
Means 1997), A. hawksworthii has been separated from other dwarf mistletoe
populations long enough to have evolved into a distinct taxon, endemic to the MPR.
On March 6, 1998 we surveyed the pine populations near Poptun, Guatemala.
These pine populations are typical Pinus caribaea var. hondurensis as reported by
Perry (1991). We estimate the overall forest area dominated by pine near Poptun to be
restricted to about 300 sq. km; located adjacent to the Belizean border, extending 38
km west; and south of the Machaquila River, extending 8-12 km south of the river
(Perry 1991 and personal communication with Placido Castellanos of Machiquila,
Guatemala). Because of this limited area, we were able to survey the majority of the
pine forest (elevation 500-700 m) in the Poptun area. We did not observe
Arceuthobium hawksworthii in these forests and we believe it does not occur there.
Therefore, we are doubtful that A. hawksworthii occurs in Guatemala.
In Honduras we relocated the population of Arceuthobium hondurense northwest
of Cofradia (approximately 20 km west of San Pedro Sula) in Cusuco National Park
(Figure 2) at an elevation of 1400 m (ca. 3 km northwest of the village of Buenos
Mathiasen et al.: © Summary of two Arceuthobium in Central America 157
Aires). This is the same population of A. hondurense sampled by J.S. Beatty in 1991
that is reported in Hawksworth & Wiens (1996). Our limited roadside surveys in this
area did not reveal any other populations of this rare dwarf mistletoe in northwestern
Honduras, but we suspect it occurs elsewhere within and in the vicinity of Cusuco
National Park.
Host Range
We examined host material (needles and mature female cones) collected from
several pines at different elevations in the MPR in an attempt to clarify the identity of
the hosts parasitized by Arceuthobium hawksworthii. Specimens of foliage and
female cones of dwarf mistletoe-infected pines were collected and forwarded to J.P.
Perry, Jr. for confirmation of our tentative field identifications, including a sample
taken from a putative hybrid (Pinus caribaea var. hondurensis X Pinus oocarpa var.
ochoterenai). In all cases our identification of the host agreed with those of Mr. Perry,
except that he felt the putative hybrid was best classified as Pinus oocarpa vat.
ochoterenai. Our field observations from the MPR confirm that Pinus caribaea var.
hondurensis is the principal host of A. hawksworthii in this area. This pine is severely
infected in many areas and tree mortality is common in severely infested pine stands.
We were unable to find locations with sufficient numbers of Pinus oocarpa var.
ochoterenai growing near severely infected P. caribaea var. hondurensis to gather
quantitative data on the comparative susceptibility of these hosts to Arceuthobium
hawksworthii. However, based on our observations at several locations where both
P. caribaea var. hondurensis and P. oocarpa var. ochoterenai were both parasitized by
A. hawksworthii, we believe the latter pine is less susceptible to this dwarf mistletoe.
In some areas where many P. caribaea var. hondurensis were severely infected and
where many dead P. caribaea var. hondurensis had evidence of past mistletoe infection
(witches--brooms), infection of occasional, large (> 10 m in height) P. oocarpa var.
ochoterenai was much less severe. In addition, we observed infection of small P.
caribaea var. hondurensis (less than 2 m in height) under larger, infected P. caribaea
var. hondurensis, but small P. oocarpa var. ochoterenai growing in the same areas
were not infected. We tentatively classify P. oocarpa var. ochoterenai as a secondary
host of A. hawksworthii using the host susceptibility classification system of
Hawksworth & Wiens (1972).
Wiens & Shaw (1994) reported that a species of Psittacanthus is common on Pinus
caribaea var. hondurensis in the lowlands of Belize and in the MPR. We frequently
observed this mistletoe on pines in these areas as well and identified the mistletoe as
Psittacanthus pinicola Kuijt. This species of Psittacanthus is distinguished by its red
to orange flowers with yellow-green tips which are about 4 cm long with a
conspicuous ligule at their base (Kuijt 1987). Another distinctive feature is that it has
leaves whose margins taper into the petiole. We also observed both Psittacanthus
pinicola and Arceuthobium hawksworthii parasitizing the same tree, and in some cases
the same branch, in the MPR as was reported by Wiens & Shaw (1994).
The taxonomic status of the Pinus oocarpa populations in the MPR is still in
question and some investigators consider these populations to be more representative
of P. tecunumanii (Schw.) Eguiluz & Perry (Farjon & Styles 1997). Other
158 PHYTOLOGIA _ February 1998 volume 84(2):154-164
investigators consider these populations to represent P. oocarpa var. ochoterenai (Hunt
1962; Perry 1991). We have chosen to follow the classification of Perry (1991).
Hawksworth & Wiens (1996) reported the host of Arceuthobium hondurense in
northwestern Honduras is Pinus maximinoi, but this host classification was tentative
and based on information provided to one of us (J.S. Beatty) by Honduran Forestry
personnel. Our examination of the pines being parasitized in northwestern Honduras
indicates that the host is P. oocarpa var. ochoterenai and not P. maximinoi as reported
by Hawksworth & Wiens. Therefore, we conclude that A. hondurense has only been
found parasitizing P. oocarpa vars. oocarpa and ochoterenai in Honduras, thus far.
Because so little is currently known about the distribution and host range of A.
hondurense, it is likely that once its distribution is better documented in Central
America, it will be found parasitizing other species of Pinus.
Plant Measurements
Additional measurements of male and female plants were made on fresh specimens
of Arceuthobium hawksworthii in order to add to the small amount of data available
for this species. Plant height (nearest 0.1 cm), the width of the base of shoots (nearest
0.1 mm), and the length (nearest 1.0 mm), and width of the third internode (nearest
0.1 mm) were measured on the dominant shoots from 150 infections of each sex.
Means, standard deviations, maximums, and minimums for these four morphological
characters are presented below:
Loionieers 1A 9.7
Po Mean | S_ Dev. | Max. | Min._|
ie re latte
[en ite ere
el aa al
ie lan tl
Third Internode:
Ree ele A
Length (mm)
Alea?
Width (mm)
Height (cm)
The largest plants we found were males (approximately 33 cm) and slightly
exceeded the maximum plant height (30 cm) reported by Wiens & Shaw (1994);
otherwise, these sizes and ranges are consistent with those reported by Wiens &
Shaw. It is interesting to note that based on our measurements of 300 plants, which is
the largest set of plant measurements completed for a dwarf mistletoe of which we are
aware (see Hawksworth & Wiens 1996), the set of morphological characters we
Mathiasen et al.: | Summary of two Arceuthobium in Central America 159
measured for female and male plants of Arceuthobium hawksworthii are the nearly the
same.
Additional measurements of male and female plants were made on fresh specimens
of Arceuthobium hondurense collected on March 10 and August 21, 1998.
Measurements were the same as for A. hawksworthii (above), but only 25 infections
of each sex were measured for A. hondurense. Means, standard deviations,
maximums, and minimums for these four morphological characters are presented
below:
Male
ae
Height (cm) 13.6
ma. 2 4
=e
Third Internode:
See
The sizes and ranges of these characters for female plants are nearly identical to
those reported for this taxon (both sexes combined) by Hawksworth & Wiens (1972,
1996), but the sizes and ranges for male plants are larger. For instance, the largest
plant(s) reported by Hawksworth & Wiens (either sex) was only 21 cm compared to
the 32 cm male plant we measured from southeast of Tegucigalpa.
Phenology
Male plants of Arceuthobium hawksworthii were still flowering as of March 5 at
elevations over 800 m in the MPR. As was reported by Wiens & Shaw (1994) male
flowers had perianths that were predominantly 3-merous and rarely 4-merous, but we
did not observe male flowers with the vivid red color inside the perianth lobes
described by Wiens & Shaw. The male flowers we observed were the same color on
the inside as on the outside--a slightly darker shade of yellowish-brown than the
staminate spike shoot color. Female flowers were in an early stage of fruit
development on many female plants and no mature fruits were observed on female
plants in early March. Wiens & Shaw (1994) speculated that A. hawksworthii
dispersed seed in June. However, our observations of fruit in early June indicated that
they were not near maturity. Our observations of fruit on August 18--20, 1998 found
that only a few fruits were nearing maturity, but no fruit were dispersing seed then and
attempts to manually press seeds out of fruit were unsuccessful. Anthesis was not
160 PHYTOLOGIA _ February 1998 volume 84(2):154-164
occurring in either June or August. Although Wiens & Shaw (1994) speculated that
A. hawksworthii has multiple periods of anthesis and seed dispersal, this taxon
appears to only have one period of anthesis annually (mid January through early
March) and one annual seed dispersal period that we estimate is from mid September
through October based on our observations in August, 1998. Therefore, fruit
maturation requires 6--7 months.
Male plants of Arceuthobium hondurense had begun anthesis and some fruits on
female plants were beginning to disperse seed on August 21. Therefore, flowering
and seed dispersal of A. hondurense starts slightly earlier than September which was
reported by Hawksworth & Wiens (1972, 1996) as the period of anthesis and seed
dispersal. The inside of male flower perianths of A. hondurense are bright red as
previously reported (Hawksworth & Wiens 1972, 1996), but perianth lobes become
the same color as male shoots when dried. Male and female plants of A. hondurense
demonstrate a high degree of sexual dimorphism. Male plants are more open and
spreading, and female plants are compact and more densely branched.
Sex Ratio
Dwarf mistletoes are dioecious and most species have a 1:1 sex ratio (Hawksworth
& Wiens 1996), but some species are reported to have a female-biased sex ratio
(Hawksworth & Wiens 1996; Wiens et al. 1996). On May 31 and August 19 we
systematically examined separate infections of Arceuthobium hawksworthii and
determined their sex based on flower morphology of mature plants. Twenty infected
Pinus caribaea var. hondurensis were felled and infections on each branch, starting at
the bottom of the tree, were examined for infections with mature plants. We sexed
1066 infections; 522 were male and 544 were female. Therefore, the sex ratio of A.
hawksworthii is 1:1, like most species of the genus.
On August 21 we systematically examined infections on five Pinus oocarpa and
determined the sex of 64 mature plants of A. hondurense. Mature plants that could be
sexed from the ground on small trees (less than 6 m in height) using binoculars were
included in this sample and only one larger tree (10 m in height) with several infections
with mature plants was destructively sampled. The sex ratio of this small sample of
mature plants was 1:1 (31 females and 33 males), but a larger sample is needed to
confirm this preliminary finding.
Plant Collections
Specimens of male and female plants of Arceuthobium hawksworthii from the
MPR were deposited at the Deaver Herbarium, Northern Arizona University,
Flagstaff, AZ, the US National Herbarium, Washington, D.C. and the Forestry
Herbarium, Ministry of Agriculture, Belmopan, Belize. Specimens of male and
female plants of A. hondurense were collected from Cusuco National Park and from
southeast of Tegucigalpa and deposited at the Deaver Herbarium, Northern Arizona
University, Flagstaff, AZ, the US National Herbarium, Washington, D.C., the
Herbario Paul C. Standley, Escuela Agricola Panamericana, Tegucigalpa, Honduras,
and at Southern Illinois University, Carbondale, IL.
Mathiasen et al.: | Summary of two Arceuthobium in Central America 161
Cayo District
@©an Ignacio
Mountain Pine Ridge
%4 ‘>
oS SI
SLR)
Guatemala
Guatemala
Cayo
District
as SS —_,
Figure |. Location of the Mountain Pine Ridge Region (MPR) in Belize and
the approximate distribution of Arceuthobium hawksworthii within
the MPR. Thick line in blow-up represents the MPR and the cross-
hatched area represents the distribution of A. hawksworthii
(approximately 250 sq. km). Approximate area of pine forests
surveyed in Guatemala is indicated by dark line around Poptun.
162 PHY TOLOGTIA February 1998 volume 84(2):154-164
Guatemala
El Salvador Nicaragua
75 150 Km
Figure 2. Location of the only known populations of Arceuthobium
hondurense in Honduras: Cusuco National Park (circle)
and in the Piedra Herrada Mountains southeast of
Tegucigalpa (star).
Mathiasen et al.: © Summary of two Arceuthobium in Central America 163
Because the nuclear ribosomal DNA cistron has not been sequenced for
Arceuthobium hondurense as has been done for several other species of Arceuthobium
(Nickrent et al. 1994; Nickrent 1996), specimens of plant material of A. hondurense
from both of its known locations in Honduras were sent to Dr. D.L. Nickrent,
Southern Illinois University, Carbondale, IL. Dr. Nickrent will complete a molecular
analysis of the internal transcribed spacer regions for A. hondurense and determine if
this information provides insights into the phylogenetic relationships of this dwarf
mistletoe with other North American species, particularly A. bicarinatum Urban which
occurs on the island of Hispaniola and is morphologically similar to A. hondurense.
LITERATURE CITED
Bateson, J.H. 1972. New interpretations of the geology of Maya Mountains, British
Honduras. Amer. Assoc. Petroleum Geologists Bull. 56:956-963.
Bateson, J.H., & I.H.S. Hall. 1977. The Geology of the Maya Mountains, Belize.
Institute of Geological Sciences, London, Great Britain.
Farjon, A., & B.T. Styles. 1997. Pinus (Pinaceace). Flora Neotropica Monogr. 75,
New York Botanical Garden, New York, New York.
Hawksworth, F.G. & D. Wiens. 1970. New taxa and nomenclatural changes in
Arceuthobium (Viscaceae). Brittonia 22(3):265-269.
Hawksworth, F.G. & D. Wiens. 1972. Biology and Classification of Dwarf
Mistletoes (Arceuthobium). USDA For. Serv. Ag. Handb. 401.
Hawksworth, F.G. & D. Wiens. 1977. Arceuthobium (Viscaceae) in Mexico and
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Symposium. USDA For. Serv. Gen. Tech. Rep. RM-111.
Hawksworth, F.G. & D. Wiens. 1996. Dwarf Mistletoes: Biology, Pathology and
Systematics. USDA For. Serv. Ag. Handb. 709.
Hunt, R.S. 1962. Some notes on the pines of British Honduras. Empire Forestry
Review 41:134-145.
Kuijt, J. 1987. Novelties in Mesoamerican mistletoes (Loranthaceae and Viscaceae).
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Means, D.B. 1997. Natural History of Mountain Pine Ridge, Belize. Tallahassee,
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Nickrent, D.L., K.P. Schuette, & E.M. Starr. 1994. A molecular phylogeny of
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Hawksworth, F.G. & D. Wiens, Dwarf Mistletoes: Biology, Pathology and
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Wiens, D. & C.G. Shaw, III. 1994. Arceuthobium hawksworthii (Viscaceae), a new
species of dwarf mistletoe from Belize. J. Idaho Acad. Sci. 30(1):25-32.
Wiens, D., D.L. Nickrent, C.G. Shaw, F.G. Hawksworth, P.E. Hennon, & E.J.
King. 1996. Embryonic and host-associated skewed adult sex ratios in dwarf
mistletoe. Heredity 77:55-63.
Phytologia (February 1998) 84(2):165-166.
NEW NAMES IN THIS ISSUE OF PHYTOLOGIA
As a result of the International Botanical Congress in Tokyo in 1993, the
International Association of Plant Taxonomy has been tasked with exploring the
feasibility of registration of plant and fungi names. In accordance with terms of the
pilot implementation of the registration concept, new names and combinations
produced in this issue of PHYTOLOGIA are listed below.
New name or combination
Podachenium chimalapanum B.L. Turner
Tintinabulum campanulatum (A. Gray) V. Grant
Tintinabulum inyoensis (I.M. Johnston) V. Grant
Maculigilia V. Grant
Maculigilia maculata (Parish) V. Grant
Allophyllum tenerrimum (A. Gray) V. Grant
Neurolaena jannaweissana B.L. Turner
Parakeelya Hershkovitz
Parakeelya arenicola (Syeda) Hershkovitz
Parakeelya balonensis (Lindl.) Hershkovitz
Parakeelya brevipedata (F. Muell.) Hershkovitz
Parakeelya calyptrata (Hook. f.) Hershkovitz
Parakeelya composita (Nees) Hershkovitz
Parakeelya corrigioloides (F. Muell. ex Benth.) Hershkovitz
Parakeelya creethae (Tratman ex Morrison) Hershkovitz
Parakeelya disperma (J.M. Black) Hershkovitz
Parakeelya eremaea (Ewart) Hershkovitz
Parakeelya gracilis (Benth.) Hershkovitz
Parakeelya granulifera (Benth.) Hershkovitz
Parakeelya lehmannii (Endl.) Hershkovitz
Parakeelya liniflora (Fenzl) Hershkovitz
Parakeelya nana (Nees) Hershkovitz
Parakeelya papillata (Syeda) Hershkovitz
Parakeelya pickeringii (A. Gray) Hershkovitz
Parakeelya pleiopetala (F. Muell.) Hershkovitz
Parakeelya polyandra (Benth.) Hershkovitz
Parakeelya polypetala (Fenzl) Hershkovitz
Parakeelya porifera (Syeda) Hershkovitz
Parakeelya primuliflora (Diels) Hershkovitz
Parakeelya ptychosperma (F. Muell.) Hershkovitz
Parakeelya pumila (Benth.) Hershkovitz
Parakeelya quadrivalvis (F. Muell.) Hershkovitz
Parakeelya remota (J.M. Black) Hershkovitz
165
Page Number
166 PHYTOLOGIA _ February 1998
Parakeelya reticulata (Syeda) Hershkovitz
Parakeelya schistorhiza (Morrison) Hershkovitz
Parakeelya spergularina (F. Muell.) Hershkovitz
Parakeelya sphaerophylla (J.M. Black) Hershkovitz
Parakeelya stagnensis (J.M. Black) Hershkovitz
Parakeelya strophiolata (F. Muell.) Hershkovitz
Parakeelya tumida (Syeda) Hershkovitz
Parakeelya uniflora (F. Muell.) Hershkovitz
Parakeelya volubilis (Benth.) Hershkovitz
volume 84(2):165-166
103
103
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