SIDA susie

VOLUME 7 NUMBER 1 MAY 1977
CONTENTS

Studies in the Ranunculaceae of the southwestern United

States. VI. Miscellaneous genera. Carl S. Keener. 1
‘The pollen morphology of Dalea section Cylipogon

(Psoraleae: Leguminosae). Blanche W. Meeson. 13
A new species of Chaetopappa (Asteraceae-Astereae) from

north central Mexico. B. L. Turner. 22
Rare and endangered aquatic vascular plants of Ohio:

an annotated list of the imperiled species.

Ronald L. Stuckey and Marvin L. Roberts. 24
A new species of Rhynchospora (Cyperaceae) from

southwestern Georgia. Robert Kral. 42
Vascular flora of the Ragland Hills area, Forrest and Perry

counties, Mississippi. Ken E. Rogers. 5]
Chromosome numbers of gypsophilic plant species of the

Chihuahuan Desert. A. Michael Powell and Shirley A. Powell. 80

NOTES. New and verified additions to the Nebraska vascular flora. I]. 91—
Sarcostemna cynachoides (Asclepiadaceae) in Arkansas. 92—Psilotaceae

symposium reprints available. 93

US ISSN 0036-1488
SIDA, CONTRIBUTIONS TO BOTANY
Founded by Lloyd H. Shinners, 1962

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SMU Herbarium

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©
Sida, Contributions to Botany, Volume 7, Number 1, pages 1-93
Copyright 1977
by Wm. F. Mahler

STUDIES IN THE RANUNCULACEAE OF THE
SOUTHEASTERN UNITED STATES.
VI. MISCELLANEOUS GENERA”

CARL S. KEENER
Department of Biolo
The Pennsylvania State University
University Park, Pa. 16802

In providing a treatment of the Ranunculaceae for the forthcoming Vascu-
lar Flora of the Southeastern United States, a number of nomenclatural and
taxonomic decisions were made which call for additional clarification (see
Keener, 1975, for a general rationale governing these studies). This treat-
ment of various smaller genera is deliberately conservative and I have at-
tempted to note those species deserving careful biosystematic study.

Within the southeastern United States, Ranunculaceae is represented by
18 genera and 93 species which compares to 50-70 genera and 2000-4000 spe-
cies for the world (Buchheim, 1964; Zimmermann, 1965). The following key
to the genera native or naturalized within the southeastern United States
also includes three genera (Adonis, Helleborus, Nigella) which occasionally

nunculus, and Thalictrum; these genera, although included in the key given
below, are not discussed further.

KEY TO GENERA

1. Flowers actinomorphic, the sepals mostly equal; fruit vario
2. Leaves opposite; styles of mature fruits usually elongate a eal
atis.
2. Leaves whorled, alternate or basal; styles of mature fruits not aici
and plumose.
3. Fruit an achene or utricle; carpels 1-ovuled.
4. Sepals short-spurred; leaves basal, linear; fruiting receptacles more
than 1 cm long yosurus,
4. Sepals not spurred; "leaves various, rarely basal and linear: fale
receptacles usually less than - m long
d. Petals present, usually show

sed on a manuscript and notes compiled for the forthcoming Vascular Flora of the
Southeastern rae — The area covered in this treatment is bounded by and i e
Louisia Arkan ntucky, West Virginia, Maryland, and Delaware. In aeer the
format ie Radfor ‘4 et al. (1967). Any suggestions for improving this treatment should
be sent to me so that necessary corrections can be made before the Vascular Flora is in press.
? Contribution No. 134 from the Department of Biology, The Pennsylvania State Uni-
versity.

5"
aQ
c
Q
a

SIDA 7(1): 1-12. 1977,

3.

6. Petals basally nectariferous; leaves not finely dissected (except
in several aquatic species); terrestrial and aquatic
‘ ies
6. Petals lacking basal nectaries; “leaves: finely dissected:
trial (introduced species sporadically escaped from per eeer ne
toe ee ee ee) 6Adonis.
5. Petals absent, the sepals or filaments often well-developed.
Leaves ternately compound or decompound, the ate usually
distinctly petiolulate; flowers often unisexual Thalictrum.
Leaves simple, often deeply parted, or if compound, the leaflets
searcely petiolulate; pears usually bisexual.
8. Leaves cauline, alternate; involucral leaves (bracts) absent;
fruit a dehiscent utricle . . . . .). Trautvetteria.
8. Leaves basal, except for the opposite or whorled involucral
leaves subtending the flowers; fruit an achene.
9. Involucral leaves entire, nee closely subtending (less
than 3 mm) the petaloid ca : Hepatica.
9. Involucral leaves compou he or de seply divided (the Meads
or segments variously toothed, lobed or dissected), rem
om the flowers : ieee
Fruit a follicle or berry; carpels at least 9. ovule od.
Plants small shrubs; inflorescences paniculate, drooping
yd ee a ee . Nanthorhiza.
10. Plants herbaceous; inflorescences not paniculate, “the flowers race-
mose or solitary.
11. Largest leaves simple, variously toothed to deeply palmately
lobed.
12. Sepals 3; ovules 2; fruit a berry Hydrastis.
12. Sepals more than 3: ovules more than 2: fruit a . follicle.
13. Petals absent; sepals petaloid, usually bright yellow; Paha
simple, merely shallowly toothed toe eee.) 6Caltha.
13. Petals (staminodia) 2-lipped, hollow: sepals See green-
ish-yellow; leaves palmate or re (introduced es oc-

casionally escaped from cultivation) Helleborus,
11. Largest leaves ternately compound iG decompound. or finely dis-
sected.

14. Leaves variously ane ee — generally broad, toothed or

lobed leaflets; carpels dis
15. Petals distinctly nase | eee variously colored
Aquilegia.

15. oe absent, or if present, not spurred; flowers usually

whit
16. coe small, not petaloid, caducous; flowers many, race-

mose.
17. Carpels 1-8, becoming saa flowering racemes clong-
ate, more than 15 em lo wee ww.) 6Cimicifuga.

17. mee 1, becoming a ae flowering racemes. short,
s than 5 cm long claed.
16. Sepals conspicuous, pet taloid, ao cadena: dowers solitary

i a als ne Dah small; follicles oo
tate; cauline leaves abse Coptis

18. an anne follicles ae ine Tenge present
: Enemion.

14. Leaves finely dissected into narrow segments; carpels a

ally connate (introduced species sporadically esca a fro

ultivation) seit,
1. Flowers hh anata the ‘upper (posterior) sepal distinctly ae the

others; fruit a follic

19. Upper sepal ho are or helmet-shaped, not spurred; petals 2(—5), en-
tirely concealed by the hooded sepals Aconitum.
19. ea aes spurred, not hooded; petals 2 « or 4, with at least the hmbs

20. Cunpel 1; petals 2, united: introduced annuals . Consolida.
20. Carpels 2 or more; petals 4, distinct; native perennials
; . . Delphinium.
ACTAEA L., Baneberry
The baneberry genus Actaea includes about 10 species widely distributed
in the northern hemisphere (Tamura, 1966). Two species are in eastern North
America.

1. Berries white (rarely red); mature eed stout, more than 1 mm thick;
PA

stigmas during anthesis 1-2 mm broad, sessile 1. A. CHYPODA.

1. Berries red (rarely white e); mature pedicels slender, a than 1 mm
thick; stigmas during anthesis 0.5-1 mm broad, subse -.

. F A. RUBRA.

1. A. PACHYPODA Ell., White Baneberry, Doll’s-eyes

Rich open woods and thickets; all prov. SE except LA. [ALL except Tex.
and Okla.]. Incl. forma rubrocarpa (Killip) Fern.—Fernald (1950), a form
with red fruits; A. alba sensu Small (1933), Gleason and Cronquist (1963),
non Miller (1768).

The taxonomy of the eastern North American species of Actaea is fairly
well understood. There are two species which differ chiefly in pedicel thick-
ness, and although both species have red and white fruited forms, within a
given species one color predominates (cf. key above). Of the two species,
however, the nomenclature for the widespread thick-pedicelled species has
not been satisfactorily resolved. At issue is whether the name A. alba Miller
or A. pachypoda Ell. should be applied to this species.

When Linnaeus (Sp. Pl. I: 504. 1753) described A. spicata var. alba he
used no specimens, but merely cited two works in which the plant was pre-
viously figured: t. 77 of Cornut’s Canadensium Plantarum Historia (1635)
and ii. fol. 1, t. 2, fig. 7 of Morison’s Plantarum Historia Universalis (1680)
both reproduced by Fernald (1940). Later, when Miller (1768) published the
name A. alba he gave a brief diagnosis and merely cited Morison (but not
Linnaeus) and as Mackenzie (1928) pointed out, ‘“ it is not correct
therefore to write Actaea alba (L) Miller.” According ie. Mackenzie (1928),
although Miller grew the plant, no specimen apparently has been preserved.

ackenzie (1928) maintained that the illustrations by Cornut and Morison
(who obviously copied his illustrations from Cornut—see Fernald, 1940) rep-

resented an Actaea with ovoid racemes and slender pedicels. He concluded
that Cornut, Morison, Linnaeus, and Miller all described the slender-pedi-
celled white-fruited form which is currently called A. rubra (Aiton) Willd.
forma neglecta (Gillman) Robinson. But Mackenzie never followed through
on the nomenclature. Instead of applying the name A. alba Miller to the
white-fruited form of the slender-pedicelled species which would then have
become the name for the species, he regarded the red-fruited form to be
more natural (‘‘undoubtdly the specific type’’) than the rarer white form.
Consequently, Mackenzie used the name A. rubra (Aiton) Willd. for the
species. Finally, there being no earlier available name, Mackenzie applied
Elhott’s A. pachypoda to the thick-pedicelled plants.

Fernald (1940) thoroughly reviewed the nomenclatural history of Actaea
spicata var, alba and concluded that Cornut’s illustration was not that of a
Canadian plant. Moreover, Fernald pointed out that although Cornut’s sketch
apparently was based on the European A. spicata (a species with black
ovcid fruits), the phrase “‘baccis niveis’’ seemingly referred to North Amer-
ican material. Fernald’s conclusion was that inasmuch as ‘“‘the basis of
A. alba Mill. was confused,’’ the unequivocal name A. pachypoda Ell. should
be used instead.

Gleason (1944) agreed with the facts set forth by Mackenzie and Fernald,
but he differed sharply with their conclusions. In the first place, Gleason
argued that greater weight should be given to the description of the fruit.
Gleason noted that in the description of his species (‘‘Aconitum baccis
niveis & rubris’’), Cornut included a statement to the effect that ‘‘the white
berry became ‘orbicularis’ at maturity and had a purple spot at the sum-
mit.”’ This seemingly implied that the plant was from eastern North Amer-
ica, not from Europe. Furthermore, Linnaeus, having no specimen of the
North American thick-pedicelled, white-fruited baneberry, based his variety
alba entirely on ey description and illustrations. Gleason believed that
Linnaeus’ diagnosis CB: Aconitum baccis niveis. Corn. canad. 76. t. 77°’)
clearly singled out the North American white-fruited baneberry by eliminat-
ing the European black-fruited A. spicata and the American red-fruited
A. rubra. Consequently, Gleason felt that ‘‘A. alba (L.) Miller’? should be
the name applied to the thick-pedicelled, white-fruited baneberry of eastern
North America.

After reviewing the evidence, I have adopted A. pachypoda Ell. for the
thick-pedicelled eastern North American baneberry for the following rea-
sons:

1. Fernald (1940) clearly demonstrated that the sketches of both Cornut
and Morison illustrate the European A. spicata far better than they illustrate
either of the two eastern North American species.

2. Linnaeus’ diagnosis (Sp. Pl. I: 504. 1753) for variety alba (“B. Aconi-
tum baccis niveis’’) is too brief. It could be applied equally well to the com-
mon white-fruited thick-pedicelled species (A. pachypoda) or the white-
fruited form of the thin-pedicelled species (A. rubra).

3. Because no type material for var. alba is known to have been preserved,
and in view of Linnaeus’ inadequate diagnosis, I am relying on the illustra-
tions by Cornut and Morison to clarify matters. Because the illustrations and
diagnosis apparently refer to two different taxa, it is best to regard Lin-
naeus’ variety alba as a nomen ambiguum

4. Miller’s diagnosis (Miller, 1768) is also es brief. The only definite points
in his diagnosis are the phrase ‘‘racemo ovato” and a single reference to
Morison’s illustration. Because no type material has been preserved so far
as is known (Mackenzie, 1928), one is again forced to rely on an illustration
which more accurately figures the European A. spicata than it does the
eastern North American plants.

2. A. RUBRA (Aiton) Willd., Red Baneberry

Rich woods and thickets; chiefly mts. Reported for W.Va.; to be sought
elsewhere in the SE. [ALL except Tex. and Okla.] Incl. forma neglecta
(Gillmar) Robinson—Fernald (1950), a form with white fruits; A. pachypoda
Ell.—sensu Radford (1968), pro parte.

Actaea rubra ranges over the cooler parts of North America and probably
is a species which recently (in post-Pleistocene time) migrated into eastern
North America (Kane, 1966). It has been reported for West Virginia (Fer-
nald, 1950), and Tennessee and North Carolina (Small, 1933), but I have
seen no specimens. It should be sought in the cool mountainous regions of
Maryland, Virginia and West Virginia (cf. Strausbaugh and Core, 1971).

AQUILEGIA L., Columbine
This genus of about 70 species (Munz, 1946) is widely distributed in the
northern hemisphere. There are two species in the southeastern United

ates, one native and one an introduced escape.

1. Spurs straight to curved, not ee oa perianth ae yellow or
ted

both; stamens conspicuously exse 1. A. CANADENSIS
1. Spurs strongly she As oe en pie: pure, a or white;
stamens scarcely exserted fue ee oe A. VULGARIS.

1. A. CANADENSIS L., Wild Columbine

Rich rocky woods, ledges, pastures and roadside banks; all prov. SE
except La. and Miss. [ALL]. Incl. A. coccinea Small—Small (1933), A. c.
var. coccinea (Small) Munz—Fernald (1950), A. australis Small—Small
(1933), A. c. var. latiuscula (Greene) Munz—Fernald (1950).

Munz (1946), who revised the genus on a worldwide basis, recognized five
varieties of A. canadensis although he admitted that ‘‘there is abundant
intergradation and many specimens are difficult to place.’’ Until thorough
biosystematic studies involving comparative population analyses are under-
taken to assess the overa!l geographic variability, I prefer to recognize

A. canadensis as a wide-ranging polymorphic species.

2. A. VULGARIS L.
A garden escape occasionally established along roadsides, waste places
and fields; mts. and pied. NC., W.Va. [Pa., N.J.].

Although Munz (1946) provided a key and description to 24 varieties of
this species, most seem to be mere trivial variants based on flower color
and size, leaflet shape, size and dissection, pubescence, etc. Consequently,
no attempt has been made to account for them in our flora.

CALTHA L., Marsh-Marigold
One of the most primitive genera within the Ranunculaceae, Caltha is a
worldwide, generally extra-tropical genus of ca. 20 species (Tamura, 1966;
Buchheim, 1964), one of which occurs throughout arctic and eastern North
America and Eurasia.

1. C. PALUSTRIS L.

Wet woods, wet meadows, marshes and bogs; chiefly mts. Del. cp., Md.,
N.C., Tenn., Va., W.Va. [Ill., Ind., Ohio, Pa., N.J.].

This species varies considerably with respect to ‘‘habit, size, shape of leaf
and follicle and in chromosome number” (Tutin, 1964). Kootin-Sanwu and
Weodell (1971) analyzed the cytogenetic relationships of various chromosome
races and concluded that C. palustris 1) is sexually reproducing with the
plants self-incompatible, 2) consists of a number of chromosome races with
a base number of 8 [the 2n—32 race is common in North America; 2n=—56
race the only one in Britain; other races in Europe include 2n=16, 28, 48,
60, 72 plus many aneuploids], 3) by means of polyploidy, hybridization and
obligate outbreeding coupled with vegetative reproduction has become a
highly successful species now widely established in wet habitats throughout
the northern hemisphere, and 4) ‘‘is interesting in that there exist well-
defined, partially or completely isolated chromosome races which if mor-
phologically distinct could be regarded as separate species, but which must,
on present evidence, be regarded as one.”’ In view that the eastern North
American plants are an apparent uniform 2n=32 race, it would be of interest
to analyze those populations consisting of slender plants with small flowers
and broadly reniform leaves formerly recognized as C. flabellifolia Pursh
(Pursh, 1814) [= C. palustris var. radicans (Forst.) Beck—see Smit, 1973].
For a comprehensive treatment of Caltha, see Smit (1973).

CIMICIFUGA L., Bugbane
Ramsey (1965), who studied the genus from a comprehensive perspective,
recognized 12 species widely distributed throughout the northern hemisphere,
of which three are in eastern North America.

1. Carpels 1(-2), aes Lees firm-walled.

. Follicles ca. 7 m , the seeds smooth; petals (staminodia) present;
leaflets 20 or aoe ee ‘lea 1. C. RACEMOSA.,
. Follicles ca. 16 ae long, the seeds scaly: petals (staminodia) absent;
leaflets 3-9(17) per leaf se a 2 (CG; RUBIFOLIA.
1. Carpels 3-8, ane tae thin, papery an 3. C. AMERICANA.

1. C. RACEMOSA (L.) Nuttall, Black Cohosh

Rich moist woods, wooded slopes, ravines and thickets; all prov. SE ex-
cept La. and Miss. [ALL except Tex. and Okla.]. Incl. C. r. var. cordifolia
(Pursh) Gray—sensu Fernald (1950), pro parte, and forma dissecta (Gray)
Fern.—Ferna 1950).

Plants with extremely dissected leaves have been named forma dissecta
(Gray) Fern., but such variants appear to be unworthy of taxonomic recog-
nition. With respect to variety cordifolia, see the discussion under no. 2
below

2. C. RUBIFOLIA Kearney

Wooded limestone river bluffs, ravines and coves: chiefly mts. Tenn. mts.
and IP, Va. [Ill.]. Incl. C. racemosa (L.) Nuttall var. cordifolia (Pursh)
Gray—sensu Fernald (1950), pro parte.

Cimicifuga rubifolia, a relatively rare and poorly understood species, was
described by Kearney (1897) to include the unicarpellate, sessile-ovaried
plants with large deeply cordate leaflets which he collected near Knoxville,
Tennessee.

Much earlier, Pursh (1814) described C. cordifolia which over the years
was applied (e.g. by Gray, 1895, and Fernald, 1950) to robust forms of
C. racemosa with subcordate leaflets as well as elements referable to C.
rubifolia Kearney (Ramsey, 1985). However, even though Pursh’s C. cordi-
folia was apparently based on a plant having deeply cordate leaflets, the
type description (Pursh, 1814) clearly points to a species earlier described
by Michaux as C. americana (Ramsey, 1965). Hence, Pursh’s C. cordifolia
is a synonym of C. americana and Gray (1895) and Fernald (1950) simply
misapplied the name.

3. C. AMERICANA Michaux

Rich moist wooded slopes and coves; mts., rarely pied. Ga., Ky., Md.,

N.C., 8.C., Tenn., Va., W.Va. [Pa.].

COPTIS Salisbury, Goldthread
Apparently a relict genus, Coptis consists of about 14 species widely dis-
tributed throughout the cooler parts of the north temperate zone (Tamura,
1968).
1. C. TRIFOLIA (L.) Salisbury
Rich damp mossy woods and bogs; mts. Md., W.Va. [Ind., Ohio, Pa.,
N.J.]. Our material can be segregated as var. groenlandica (Oeder) Fassett

[C. groenlandica (Oeder) Fern.—Fernald (1950) |.

On the basis of available herbarium material, Fernald (1929) treated the
plants of Greenland and Atlantic North America as C. groenlandica and dis-
tinct from C. trifolia which he considered restricted to eastern Asia and
Alaska. Fernald did remark, however, that possible transitional populations
might lead to a reclassification. Hultén (1937), in comparing specimens from
the Aleutians and southeastern Alaska, concluded that plants from the Aleu-
tians coincided with material from Kamtchatka, whereas those from south-
eastern Alaska apparently were more closely related to plants from Green-
land. Because of these more or less transitional populations, Hultén treated
the material from southeastern Alaska to British Columbia, eastern North
America, and Greenland as C. trifolia subsp. groenlandica (Oeder) Hultén
which was later reduced by Fassett (1946) to varietal status. Although I
prefer to regard C. trifolia as a single wide-ranging species, those who
recognize regional variants can segregate our material as C. trifolia var.
groenlandica (Oeder) Fassett.

ENEMION Raf., False Rue Anemone

The delimitation of taxa within the Tribe Isopyreae Schrédinger, demarked
by small chromosomes (x=7) and follicular fruits (Tamura, 1968), has long
been unsettled. Drummond and Hutchinson (1920), who thoroughly reviewed
the taxonomic history of Isopyrum and its close allies, recognized seven
genera, including Enemion which was originally described by Rafinesque
(1820) to accommodate newly discovered plants somewhat resembling the
common Rue Anemone (Thalictrum thalictroides (L.) Eames & Boivin).
Although Torrey and Gray (1838, p. 29) initially recognized Rafinesque’s
genus, in the Supplement to their Flora (vol. 1, p. 660, 1838) they transferred
Enemion biternatum to Isopyrum as well as firmly delimiting it from the
European I. thalictroides L. of which it had at one time been considered
a mere apetalous form (Drummond and Hutchinson, 1920). In the latest
revision of this group, Tamura and Lauener (1968) likewise recognized
Enemion, along with four other closely related genera (Dichocarpum, Iso-
pyrum, Leptopyrum and Paraquilegia). Of these, only Enemion lacks petals,
which, along with its several free carpels having transverse veinlets, led
Tamura (1968) to consider it as the most primitive genus within the Iso-
pyreae.

However, not all recent workers accept the generic segregation of ne-
mion. For example, Calder and Taylor (1963) believe Drummond and Hutch-
inson (1920) misinterpreted the petaloid structures of Isopyrum as petals
instead of interpreting them as staminodia and moreover placed “‘. . . too
much stress on their value as characters for segregation of the genera.”’
Nevertheless, irrespective of whether one wishes to call these structures
staminodia or petals (cf. Tamura, 1965; Cronquist, 1968: 81), the taxonomic
question is whether the absence of these structures is sufficient for a generic
segregation of Enemion from Isopyrum sen. lat.

As recognized in this paper, Enemion has six species, distributed in North
America and eastern Asia (Calder and Taylor, 1963); one of these, E. biter-
natum Raf., is in eastern North America.

1. E. BITERNATUM Raf.

Rich moist woods, rare; chiefly cp. and pied. Ala., Ark. mts., Fla., Ky. IP,
N.C., $.C., Tenn. pied., W.Va. [Okla., Mo., Ill., Ind., Ohio]. Isopyrum biter-
es (Raf.) T. & G.—Small (1933), Fernald (1950), Gleason and Cronquist
(1963), Radford (1968).

HEPATICA Miller, Liverleaf, Hepatica

Although some authors unite Hepatica with Anemone (e.g. Hiroe, 1957:
Hara, 1962; Ohwi, 1965), I regard Hepatica as a distinct genus, chiefly on
the basis of the position of its involucral bracts which closely subtend the
flowers.

There has been some difference ef opinion regarding the taxonomic treat-
ment of the eastern North American hepaticas. Current regional manuals
(e.g. Fernald, 1950; Gleason and Cronquist, 1963; Radford, 1968) generally
recognize two species, H. acutiloba and H. americana. However, critical
studies by Hara and Kurosawa (1958) and by Julian and Cora Steyermark
(1950) established that H. nobilis Miller sensu lato is widely distributed in
three disjunct regions in the north temperate zone (Europe, eastern Asia,
eastern North America). The European plants are only slightly variable
whereas those in eastern Asia are the most polymorphic of all and include
the only known tetraploid race (Hara, 1962).

With respect to the eastern North American populations there are two
relatively distinct forms, which apparently hybridize where they are sym-
patric (Steyermark and Steyermark, 1960). On the basis of their herbarium
studies, the Steyermarks (1960) concluded that “‘it is, therefore, a matter
of difficulty to distinguish some of the European forms of H. nobilis from
one or the other of the two American taxa.’ Intensive biosystematic re-
search, includ'ng comparative breeding studies should be undertaken to re-
solve the evolutionary relationships within the H. nobilis complex. However,
pending such research, I am following the preliminary work of the Steyer-
marks in recognizing the eastern North American hepaticas as regional
varieties of the European H. nobilis.

1. H. NOBILIS Miller
Two varieties are recognized following Steyermark and Steyermark (1960).

la. Var. OBTUSA (Pursh) Steyermark

Leaves cleft to about the middle of the blade; apex of leaf lobes and bracts
rounded to obtuse. Sepals typically bluish (sometimes white or pink). Rich
woods and dry upland slopes, often in acid soils; all prov. SE except La.
and Miss. [ALL except Tex. and Okla.]. H. hepatica (L.) Karsten—Small

10

(1933); II. americana (DC.) Ker.-Gawl.—Fernald (1950), Gleason and Cron-
quist (1963), Radford (1968).
lb. Var. ACUTA (Pursh) Steyermark

Leaves parted to below the middle of the blade; apex of leaf lobes and
bracts acute. Sepals typically whitish. Upland woods and rocky slopes, often
in calcareous soil; chiefly mts. and pied. Ala., Ark., Ga., Ky., Md., N.C.,
S.C., Tenn., Va., W.Va. [Mo., Ill., Ind., Ohio, Pa.]. H. acuta (Pursh) Brit-
ton—Small (1933); H. acutiloba DC.—Fernald (1950), Gleason and Cronquist
(1963), Radford (1968).

HYDRASTIS Ellis ex L., Golden Seal, Orangeroot

This monotypic genus (Janchen, 1949; Tamura, 1968), restricted to eastern
North America, was once widely used in homeopathic medicine. For exam-
ple, Lloyd and Lloyd (1884), who devoted over one hundred pages in dis-
cussing its botanical characteristics and medicinal properties, estimated
that the total annual production (ca. 1880) reached nearly 75 tons of rhizomes
and reots. Due to such extermination plus its rather exacting ecological re-
quirements, Hydrastis is now quite rare and definitely needs protection.

1. H. CANADENSIS L.
Rich woods, rare; chiefly mts. SE except Ala., Fla., La. [Mo., lL, Ind.,
Ohio, Pa.].

MYOSURUS L., Mouse-tail

The specialized and phyletically advanced genus Myosurus consists of
about 16 species widely distributed in all continents, although it is absent
in eastern Asia (Tamura, 1967). Campbell (1952) recognized four species
native to North America, one of which (M. minimus) also occurs in South
America, Hurope, Australia, and New Zealand. For a study of reproductive
biclogy, meiotic and mitotic behavior, see Stone (1959, 1960).
1. M. MINIMUS L.

Low fields in alluvial or calcareous soil; chiefly ep. and pied. Ala., Ark.,

IP, La., N.C., S.C., Tenn., Va. ep. |Tex., Okla., Mo., Ill., Ind. ].

According to Campbell (1952), M. minimus consists of eight subspecific

taxa, one of which (subsp. minimus) alone occurs in eastern North America,
TRAUTVETTERIA Fischer & Meyer, False-Bugbane

This genus is disjunct in eastern Asia, western and eastern North America

(Tamura, 1967), and consists of one to three or more species, depending on

the taxonomic treatment.

1. T. CAROLINENSIS (Walter) Vail
Wooded seepage slopes, stream banks and bogs; chiefly mts. Ala., Ga.,
Ky., Md., N.C., S.C., Tenn., Va., W.Va. [Mo., Pa.].

XANTHORHIZA Marshall, Yellow-root

This monotypic genus, confined to the southeastern United States, is dis-
tinguished by its woody habit, drooping paniculate inflorescences and yellow
weed (whence the name),

Although a number of authors (e.g. Buchheim, 1964) use the spelling
AXanthorrhiza (following recommended transliteration of classical Greek in
the formation of compound words), I am retaining the spelling originally
adcpted by Marshall (1785), a practice reeommended by Stern (1966, p. 269).

1. X. SIMPLICISSIMA Marshall, Yellow-root

Shaded stream banks, moist woods, thickets and rocky ledges; all prov.
Ala., Ga., Ky. mts., Md. cp., Miss., N.C., $.C., Tenn., Va., W.Va.

ACKNOWLEDGMENTS
Appreciation is expressed to R. A. Pursell, Monte Manuel, and Paul Roth-
rock for their critical comments on an initial draft of this paper. Special
thanks are due to curators of the following herbaria who provided access
to specimens for these studies: FSU, GA, NCU, NCSC, PAC, PENN, PH,
SMU, US, VPI, and WVA.

REFERENCES

BUCHHEIM, G. 1964. Ranunculaceae, p. 133-137. In H. Melchior (ed.), = Engler’s Syl-
el as Pflanzenfamilien, Band Il. Auflage 12. pe Borntraeger, Berlin-Nikolassee.
CAL J. A. and R. L. TAYLOR. 1963. A new species of tinea ee to the

oe igs of British Columbia and its relation to other species in the genus.
flo 69

CAMPBELL, G. oe The genus Myoswrus L. (Ranunculaceae) in North America.
Aliso 2: 398-403.

CRONQUIST, A. 1968. The evolution and classification of flowering plants. Houghton
Mifflin Co., ee on.

DRUMMOND, J. and J. HUTCHINSON. 1920. A revision of Isopyram (Ranuncula-
ceae) and its nearer allies. Kew Bull. 1920: 145-169

FASSETT, N. C. 1946. Preliminary reports on the floes of Wisconsin, No. 33. Ranuncula-
ceae. Tens. Wisc. Acad. Sci. 38: 189-210.

FERNALD, M. L. 1929. Copfis “trifolia and its eastern American representative. Rhodora
LY 13

ie i Some spermatophytes in eastern North America. Rhodora 42: 260-265,

—— . Gray’s manual of botany, 8th ed. American Book Co., New York.

GLEASON, ie - 1944, Actaca alba versus Actaea pachypoda. Rhodors 46: 146-148.

‘ _ and A. CRONQUIST. 1963. Manual of vascular soa of northeastern United

States and adjacent Canada. D. Van Nostrand Co., Princeton
GRAY, A. 1895. See bp. 1-57. In B. oe oe ed. Synoptical flora of
North America, Vol. I, pt. American Book C

HARA, H. 1962. Racial M fferences In Ww oa. species with special reference to those
common to Japan and North America. Amer. J. Bot. : -652.

—__ and S. KUROSAWA. 1958. Differentiation within Anemone hepatica L. of
Japan. J. Jap. Bot. 33: 265-274.

HIROE, M. 1957. A cytotaxonomical study on Anemone hepatica L. (Ranunculaceae) of
apan. Bot. Mag. Tokyo 70: 4-7.

HULTEN, E. 1937. The flora of the Aleutian Islands and westernmost Alaska peninsula.
Bokforlags Aktiebolaget Thule, Stockholm.

12

SS
aA ke

JANCHEN, E. 1949, Die systematische Gliederung der ae as und ieee
Denkschr. Wiss. Wien. Wien. Math.-naturwiss. Kl. 108, . 4, Abh

KANE, J. M. 1966. Biosystematies of the genus Acfaca in North ee alee:
Amer. J. Bot, - 634 (abst.

KEARNEY, T. H. 1897. New or otherwise interesting plants of eastern Tennessee. Bull.
Yorrcy Bot. Club 24: 560-575.

KEENER, C. S. 1975. Studies in the Ranunculaceae of the southeastern United States. I.
Anemone LL. eae 40: 36-44

-82.

KOOTIN- SANW U, and 8. R. J. WOODE — 1971. The cytology of Caltha palustris:
cytogenetic rel ane Heredity 26: 121-13
LLOYD, J. U. and C. G. LLOYD 1884 ee Drugs and medicines of North America,

Vol. 1. Ranunculaceae. Cincinnati.
MACKENZIE, K. K. 1928. White-fruited bane-berries. Torreya 28: 51-53.
MARSHALL, H. 1785. Arbustum americanum. Philadelphia.
MILLER, P. 1768. The gardeners dictionary, ed. 8. ea
MUNZ, P. A. 1946. Aguile, the cultivated and wild columbines. Gentes Herb. 7: 1-150.

OHWI1, J. 1965. Flora of Japan. Smithsonian Institution, Washington, D.C
PURSH, F. 1814. Flora americae septentrion i ondon.
aa A. E. 1968. Ranunculaceae, p. -468. In A. E. Radford, H. E. Ahles ane

Bell, pees al of the vascular flora of ue Carolinas. Univ. of North Carolina Pre
Cc on a
‘ “bru, J. W. HARDIN, and R. WILBUR. 1967. Contributor’s guide
for the ae flora of the southeastern United States. Dept. of Botany, University of
orth Carolina, Chapel Hill, N.C.
RAFINESQUE, C. S. 1820. Description d’un nouveau genre de plantes, Envemion, et re-
marques botaniques. J. Phys. 91: 70-72

RAMSEY, G. 1965. A biosystematic eudy of the genus Cimicifuga (Ranunculaceac).
Ph.D. Thesis, University of ‘Tennessee.
SMALL, J. K. 1933, Manual of the pole flora. Publ. by the author, New York.
SMIT, P. G. 1973. A revision of Ca i (Ranunculaceae). a sa 21: 119-150.
STERN, W. T. 1966. a ve Hafner Publ. Co., Nev
STEYERMARK, J. A. and C. S. STEYERMARK. 1960. ee hae in’ North America,
3-232

STONE, D. FE. 1959. A unique balanced breeding system in the vernal pool mousetails.
Evolution - 151-174
9

960. Nuclear cytology of the California mousetails (Myoswrus). Madrofio

ST ae eee H, D. and E. L. CORE, 1971. Flora of West Virginia, pt. TH, ed. 2. West
Virginia Univ. B rat as ane n, W.Va
TAMURA, M. 1965. Morphology, eenlony and
Rep. Osaka Univ. 14: 53-71.
1966. ies lees. ecology and phylogeny of the Ranunculaceae VI. Sci. Rep.
Osaka Univ. 15: 13-35
. 1967. Morphology, ecology and phylogeny of the Ranunculaceae VII. Sci. Rep.
Osaka Univ. 16: 21-43
968. omihalaes, ecology and phylogeny of the Ranunculaceae VIIL. Sci. Rep.
Osaka Gite 17: 41-56.
and L. A. LA \UF NER. 1968. A revision of Isopyrum, Dichocarpum and their
: 267-273

shylogeny of the Ranunculaceae TV. Sei.

alhi Not : .
TORREY, J. and A. GRAY. 1838. A fee: of North America, Vol. 1. Wiley & Putman,
New York. [Facsimile edition, 1969. Hlafner Publ. Co., New York.
— T. G. 1964. Caltha L., p. 201. Ia T. G. ‘Tutin, et al. (eds.) Flora Europaea, Vol.
ambridge Univ. Press, London.
Bey Stan ce W. 1965. Familie oe p. 33-77. In G. gi, Illustrierte
Flora yon Mittel-Europa, ed. 2. Bd. IIT (Teil Lief. 1). Carl Hanser, ie

THE POLLEN MORPHOLOGY OF
DALEA SECTION CYLIPOGON:
(PSORALEAE: LEGUMINOSAE)

BLANCHE W. MEESON?
Herbarium, Department of Biology
Southern Methodist University
Dallas, Texas 75275

ABSTRACT—Pollen grains of Dalea section Cylipogon are described by
scanning electron microscopy. This section consists of ten species; nine
are discussed here. In this section there are three definite pollen types. Two
types are clearly distinct while the third is intermediate. The first type is
spherical, highly reticulate with anomotreme apertures. The second is pro-
late, finely reticulate with tricolpate apertures. The intermediate type has
three different forms, two prolate and a subprolate form.

The genus Dalea is a heterogenecus group which is presently being revised
by R. C. Barneby. An investigation of the pollen morphology of Cylipogon
was initiated for two reasons; first, to support the segregation of these spe-
cies into a section and second, to clarify the species boundaries of these
taxa, if possible.

The proposed section Cylipogon is composed of ten species with similar
morphological characteristics. The pollen of the following Dalea species
have been examined:

D. aurea Nutt. D. hallii Gray

D. jamesii T. & G. D. laniceps Barneby
D. luisana S. Wats. D. nana Torr.

D. wrightit Gray D. parrasana Brandg.

il
VY

. prostrata Ort.
The tenth undescribed species is _known only by the fruiting form. Voucher
specimens are listed in Table

MATERIALS AND METHODS

Pollen grains were prepared by a modification of the standard acetolysis
method devised by Erdtman (1952, 1960). The pollen material was not cen-
trifuged but was allowed to settle out of the medium. This technique greatly
increased the quantity and quality of the pollen yield. Pollen prepared for
scanning electron microscopy was washed in 759% ETOH until all visible
traces of the acetolysis solution were gone.

A description of Dalea section Cylipogon will be ates in a revision of genus Dalea
by R. C. Barneby, currently in manuscript (Mem. N.Y Gard. vol. 27

a nee of Biological Sciences, University of ee ue i aks California
9310

~

SIDA 7(1): 13-21. 1977.

14

The washed pollen was deposited on the nonadhesive side of metallic
tape, attached to aluminum stubs and allowed to air dry. The desiccated
pollen was then coated with gold-palladium and examined with a MSM-2
Mini-SEM and AMR-1000.

RESULTS

The pollen morphology of Cylipogon is of three types. D. aurea and D.
nana Characterize the first pollen type. These species possess very similar
but not identical pollen grains, The pollen of both species is highly reticulate
with spherical structures underlying and connecting the reticulations. The
grains of both species are spherical with anomotreme apertures. The spe-
cific aperture patterns of the pollen grains can be used to distinguish these
species. D, aurea is characterized by at least two forked apertures where-
as D. nana has only one forked aperture and a donut-shaped aperture.
(Figs. 1, 2 and 5)

The second pollen type is represented by D. halli, D. jamesti, D. laniceps,
D. prostrata, D. parrasana and D. wrightii. The pollen grains of these spe-
cies are almost identical. They cannot be separated morphologically by
shape, by reticulation or by aperture type, position and number. Polar
lengths may be helpful but these measurements are not available. The pol-
Ien of this group is prolate, tricolpate with a finely reticulate sexine. (Figs.
3 & 5)

D. luisana characterizes the third pollen type. The pollen of this species is
predominately prolate but some grains are subprolate. All mature grains
have a reticulated sexine similar to the second pollen type. Three distinct
aperture arrangements have been found. One of the patterns is a tricolpate
grain similar to pollen group two. (Figs. 4 & 5) The second aperture ar-
rangement is formed by three colpate apertures merging at a single equa-
torial point producing a forked effect. The third form seems to be a distor-
tion of the previous arrangement. Three undulating apertures merge at the
approximate pole of the subprolate grain rather than at the equator. The
apertures appear similar to those of D. aurea and D. nana because of the
undulations and the forked junction.

DISCUSSION

The pollen morphology of the section Cylipogon shows a continuum of
pollen types from a prolate, tricolpate grain (D. hallii) to a spherical ano-
motreme grain (D. aurea). D. luisana appears to be the intermediate pollen
type because of its variability and aperture distortion, Certain pollen grains
of D. luisana are prolate tricolpate while others are distorted to subprolate
anomotreme grains. D. luisana pollen grains represent the intermediate
forms connecting the two extremes. Thus, section Cylipogon seems to be
a natural taxon.

Three of the nine species examined can be conclusively identified on the
bases of their pollen morphology. They are separated in the following man-

or:

Fig. 1. Dalea aurea. A. 1000x, B. Note that the grain has cae ve slightly, 1250x, Cc.
425

Ox, D. 1125x, FE. Note two forked apertures,
are the same grain from the rear and front.

00x, Note that A. & C.

16

Fig. 2, Dalea nana. A. 1375x, B. Note donut-shaped aperture, 1875x, C, 1125x, D. Note
forked aperture, 1250x, E. Note forked aperture, 1375x, F. 1500x.

%

33

ate

4

cae

2

me
et

phe
ook

*

Some
‘
*

ig &
Fe PM ae
ae at

a ee

laniceps, 1425x, D. D. prostrata,

c
fe)
Ss
ae)
gS
°
hae
para
—~
u
2
(
aU
Oo -w
oS
SOR
Pa
leas l
ac)
ae
oom
—
oo
an
o ™™
St one
i ie
Q
Vv.
sl an
5m
Ay
a
Or
aT
nan
[avi
2S
5 8
u 5
wy
aH
es
of
fat
cz
om
=
So:
o <
“

SX.

>

wrightii, 16

dD.

E.

1. Spherical; sexine highly reticulate; spherical protrusions between the
reticulations.
2. Two forked apertures present; donut-shaped aperture absent

in GA ee eS D. aurea.
2. One forked aperture; one donut-shaped aperture present . D. nana.
Prolate to subprolate; sexine finely reticulate; spherical sexine pro-

—

trusion absent.
3. Grains tricolpate . . . . . . Dz. hallit, D. jamesii, D. laniceps,
D. prostrata, D. parrasana, D. wrightit.
3. Grains with three merging colpate apertures and grains with tricol-
ined D. luisana.

pate apertures . . ...

1

Fig. 4. Dalea Inisana. A. Note tricolpate apertures, 2000x, B. Note colpi merging at the
E aperture, 1688x, D. Note

Xx

equator, 1125x, C. Note the number of furrows and the
undulating colpi merging and the shape of the grain, 1875

Fig. 5. Reticulation in section Colibogon. A. Dalea hallii, 6125x, B. D. jamesii, 6125x, C.
aniceps, 6500x, D. D. prostrata, 11875x, FE. D. bee 6875x, F. D. Iuisana,
11800x, G. D. aurea, Nore spherical protrusions between ridges, 12500x, H. D.

nana, Note spherical protrusions between ridges, 10625x

Table I.

Voucher Specimens for Dalea section Cylipogon

Taxa Collector Location Herbarium
D. aurea Nutt. C.L. York, 54389 Bell Co., Texas SMU
M.J. Trlica & D.V. Sellers, 115 Carson Co., Texas NY
S. Hewitt, 23 Comanche Co., Texas SMU
D.S. & H.B. Correll, 12995 Cooke Co., Texas SMU
D. halliti: Gray B.L. Turner, 1845 Dallas Co., Texas SMU
D. jamesti T. & G. J. & C. Taylor, 3715 Cimarron Co., Oklahoma SMU
D. laniceps Barneby Ripley & Barneby, 14962 Coahuila, Mexico NY
D. luisana S. Wats. Ripley & Barneby, 14768 San Luis Potosi, Mexico NY
D. nana Torr. F.R. Waller, 1412 Deaf Smith Co., Texas SMU
J.W. Thieret, 30891la Brewster Co., Texas SMU
L.H. Shinners, 32284 Menard Co., Texas SMU
D. parrasana Brandg. Ripley & Barneby, 13282 Coahuila, Mexico NY
D. prostrata Ort. Ripley & Barneby, 13940 Chihuahua, Mexico NY
D. wrightti Gray SMU-DMNH, 102 Brewster Co., Texas SMU

21

Absolute measurements of the major axis of the prolate, tricolpate grains
may provide information which can be used to separate these taxa.

ACKNOWLEDGEMENTS
I would like to thank Wm. F. Mahler, R. C. Barneby,
Laury for their assistance.

V. Allison and L.

REFERENCES
ERDTMAN, G. 1952. Pollen anon and plant taxonomy-Angiosperms Vol. 1. Alm-
qvist and oe Stockho

. The ae method. A revised description. Svensk Bot. Tidskr. 54(4):

er 564,

A NEW SPECIES OF CHAETOPAPPA
(ASTERACEAE-ASTEREAE) FROM
NORTH CENTRAL MEXICO

B. L. TURNER
Department of Botany
University of Texas, Austin 78712

Collections in the Chihuahuan Desert region of north central Mexico by
Prof. M. C. Johnston and collaborators have revealed a number of novelties,
one of the rarer being the new species described here.

CHAETOPAPPA plomeensis Turner, sp. nov. (Fig. 1).

Herbae perennes 9-13 cm altae. Laminae foliorum spathulatae supra gla-
brace. Ramuli inflorescentiae primarii secundariique tenues sinuati foliis
paucis squamiformibus. Setae pappi ca. 25, 2-3 mm longae.

Small perennial, 9-13 cm tall. Leaves alternate, mostly basal, on tufted
offshoots from a slender, ramifying rcot system; blades spatulate, sparsely
pubescent beneath, glabrous above, the margins appearing ciliate. Flower-
ing stems delicate, bearing 2-13 heads, in habit very similar to C. parryi
Gray except that the primary (and secondary) branches are spindly or
sinuate, these bearing fewer and much-reduced, scale-like, leaves. Technical
characters of the head are also surprisingly like that species except that
the cup-like crown of short scales is wholly replaced by a pappus of ca. 25
setae which are 2—3 mm long.

HOLOTYPE: MEXICO. Coahuila; La Cuesta del Plomo on the Musquiz-
Boquillas highway (28°44’ N., 102°31’ W.), 1750—777 m, steep limestone
slopes and canyons. I’. Chiang, T. Wendt & M. C. Johnston 9218b. 14 Sept.
1972 (TEX; known only from the holotype which comprises 5 plants; each
of these has the characteristic leaves and pappus).

At first glance the collections concerned would appear to be depauperate
forms of the fairly widespread Chaetopappa parryi. However, the pappus
is strikingly different and, combined with habital and leaf differences, the
taxon seems worthy of recognition. It is apparently a local, bluff-inhabiting,
endemic, much like C. hersheyi Blake, which is known only from the Guada-
lupe Mountains of western Texas. In Shinners’ (1946) revision of the genus,
C. plomoensis would key to C. parryi, but the latter has strongly erect, quite
leafy, flowering branches, leaves which are pubescent on both surfaces and,
of course, a small, squamellate, crown-like pappus, the latter being consist-
ent in the numerous specimens examined, both at the University of Texas
(TEX) and elsewhere (MO, US)

Iam grateful to Prof. M. C. Johnston for the Latin diagnosis and partial
support from his NSF grant.

REFERENCES
SHINNERS, L. H. 1946. Revision of the genus Chaecfopappa DC. Wrightia 1: 63-81.

SIDA 7(1): 22-23. 1977.

RARE AND ENDANGERED AQUATIC
VASCULAR PLANTS OF OHIO: AN ANNOTATED
LIST OF THE IMPERILED SPECIES

RONALD L, STUCKEY
Department of Botany and Center for Lake Erie Area Research
The Ohio State University, Columbus 43210

MARVIN L. ROBERTS
Department of Botany
University of Wyoming, Laramie 82071

In Ohio and elsewhere, considerable concern is growing for the identity
and protection of rare and endangered, or imperiled, species of plants.
Man’s impact on the environment has caused drastic changes that are now
affecting organisms to the extent that these imperiled species are extinct
or becoming extirpated, or their survival is endangered or threatened. Lists
of rare and endangered species of plants for various states are now begin-
ning to appear, for example, Arkansas (Tucker, 1974), Missouri (Holt,
Keefe, Lewis, Pflieger, and Sullivan, 1974), New Jersey (Fairbrothers and
Hough, 1973), and Texas (Anonymous, 1974), and a national report on the
endangered and threatened plant species of the United States recently has
been presented to the Congress of the United States (Ayensu and others,
1974). Although a list of Ohio’s rare vascular plants was published 30 years
ago (Jones, 1943), we here present a documented report on the imperiled
species of aquatic vascular plants for Ohio, which to our knowledge is the
first report for this group of plants for any given state. We are aware that
reports of this type have certain limitations and will require revision as
more information becomes available.

The indigenous or native aquatic vascular plants of Ohio are those plants
that live and complete all or most of their life history in water or in a habi-
tat that is wet throughout part of the year, such as a marsh, bog, or wet
prairie, These plants generally grow submersed, floating, submersed with
floating leaves, or emersed in shallow water. The inclusion of plants as
aquatic and the information on their habitat comes from (1) available label
data on herbarium specimens, (2) a knowledge of the biology of the species
from our field studies in the state over the past eight years, and when
necessary, (3) floristic literature for Ohio and adjacent states or from mono-
graphs of specific genera. Herbarium specimen records were obtained prin-
cipally from the collections at The Ohio State University, Kent State Uni-
versity, The University of Cincinnati, and occasionally from Bowling Green
State University and Miami University. Records for several groups studied
by specialists (Ball for Salia, Mackenzie or Hermann for Carex, for exam-

SIDA 7(1): 24-41. 1977.

20

ple) were extracted from the card files of E. Lucy Braun at The Ohio State
University Herbarium. We sometimes had great difficulty characterizing the
habitat of those species which have not been located in the state 1 fifty
years or more.

Ohio’s imperiled species of aquatic vascular plants can be grouped into
four categories based principally on (1) their distribution and abundance
geographically, (2) the changes in population numbers through time, (3)
the abundance and quality of existing suitable habitats in the state, and
(4) the peculiarities of the biology of the species. The following categories
are established and defined for use in the annotated list presented in Table 1.

EXTINCT. An extinct species is one that has been completely eliminated
not only from Ohio, but from all other areas of its range within historic
time. None of Ohio’s aquatic vascular plants belong in this category.

EXTIRPATED (EX). An extirpated species is one based on records known
from one or more counties 70-80 years ago, but which has not been reported
in Ohio since. These species may be expected to survive in areas nearby to
Ohio. In some cases our knowledge of site conditions and the species toler-
ances leads us to believe they may not exist in Ohio ered > re-
cent records. Forty species are considered to be extirpated i io

ENDANGERED (EN). An endangered species is one nr on records
known from one or more counties 70-80 years ago, but which has been re-
corded in Ohio from only one or two counties since 1950. These species are
in immediate jeopardy in Ohio and could readily become extirpated through
destruction or environmental changes of a few critical sites. A few species
without recent records have been considered as endangered because we be-
lieve they are probably present in unstudied existing suitable sites. Fifty-
four species are considered to be endangered in Ohio.

HREATENED (TH). A threatened species is one known from o!der rec-
erds and has recentiy been recorded in Ohio from three to no more than
seven counties since 1950. These species are believed to have continued to
survive at population levels comparable to their previous abundance, but
they could become endangered or extirpated if drastic environmental and
habitat changes occur. Fifty-two species are placed in this category.

hese 146 imperiled species of Ohio aquatic vascular plants, if not extir-
pated already, exist in such small population numbers that special attention
should be given for their survival as a part of the state flora. We believe
that the change in the number of populations through time illustrates the
effects the environment has had on the survival of a species. The data for
the species considered in Table 1 are presented to substantiate and explain
their inclusion. The most appropriate time periods selected to show these
changes in the number of populations for each species are (1) the number
of counties with at least one population record before about 1900, (2) the
number of counties with at least one population record since about 1950,
(3) the year of the earliest known record, and (4) the year of the latest
known record. A list of the counties where each species has been recorded

26

is also included (Table 1), and the number of imperiled species known for
cach county is stated in Table 2. The trends cbserved and recorded should
be predictive if the daia base is reasonably accurate and substantial, and if
habitat destruction occurs at about the present pace. An alternative list of
species tolerant of changes and increasing in population numbers could also
be presented. Of the 146 imperiled species, eight have never been locat-
ed in Ohio since 1900 and 320 additional ones have not been seen or reported
in the past 25 years.

The habitats of Ohio’s imperiled species of aquatic vascular plants are
quite specialized and complex, such as bogs, fens, marshes, swamps and/or
ccmbinations of these with ponds, lakes, streams, and/or mudflats. In order
to simplify and condense information on the habitat, we have listed one
or two of the most characteristic kinds of habitats in which each one of
these imperiled species may be expected to occur. The aquatic or wetland
habitat types selected along with a note on their chief distinguishing feature
are: pond or lake (quiet, open water), stream (flowing water), fen (calcare-
ous substrate conditions), bog (acidic substrate conditions), marsh (domi-
nated with herbaceous vegetation), swamp (dominated with woody vegceta-
tion), and mudflat (exposed soil conditions that were formerly covered with
water earlier in the growing season).

Considerable changes in the aquatic vascular plant flora of particular
sites (Stuckey, 1971; Judd & Taub, 1973; Lowden, 1969; Moore, 1973) and
for the genus Najas (Wentz & Stuckey, 1971) have been documented in Ohio.
Our data indicate that the aquatic habitats in northeastern Ohio and the
marshes along the shoreline of Lake Erie in northwestern Ohio have the
greatest diversity of aquatic vascular plants in the state. These areas also
contain the largest number of imperiled species of aquatic vascular plants
(Table 3). The glacial topography of northern Ohio contains most of the
bog, fen, and lake flora of the state. The marsh, swamp, stream, and mud-
flat flora is as well or even more represented here as elsewhere. The north-
ern portion of Ohio has been subjected to considerable agricultural develop-
ment, including extensive drainage and channelization. The northeastern
section is currently undergoing rapid urbanization with a severe impact on
the bog, pond, and lake flora from residential and industrial development.
This impact has been particularly significant in the Cleveland-Akron-Canton
metropolitan areas in the counties of Portage, Summit, Stark, Lake, and
Geauga. The counties bordering Lake Erie in northwestern Ohio (Irie,
Lucas, and Ottawa) have been severely affected by shoreline development
which has disturbed the flora of the marshes, swamps, and mudflats. It is
fortunate that these areas of the state that are now currently most severely
affected by habitat degradation were centers of field botanical research
70-80 years ago before such changes became apparent. The information
available from the beginning of the century has, therefore, been invaluable
or documenting the depletion of the aquatic vascular plant flora. These
trends are clearly evident, and, therefore, we believe in a strong program

=

=-

27

for further acquisition and protection of Ohio’s aquatic vascular plants and
their habitats in wetland sites.

Much of our field work on the aquatic vascular plant flora of Ohio has been
supported by the Ohio Biological Survey during the seasons of 1967-1973.
Thanks are also expressed to Mr. Thomas Duncan, Dr. Robert R. Haynes,
Mr. Ronnie Johnson, and Mr. W. Alan Wentz, who have aided in our studies
of the state’s aquatic plant flora. Miss Edna Kirby contributed in assisting
with obtaining data from the specimens in The Ohio State University Her-
barium

Table 1. Annotated Checklist of Imperiled Aquatic Vascular
Plants Indigenous in Ohio.

ood
Sa E
o © S Eas
nG& & 2 §
er Fa 2
oe. s
SS = ca =
ae . = 3
CS § @ 8g
. saa =
= _ Pos o ~
E g5552 2 2
n ei a S&S OG
co) 5 Cage, 4
J Zw On BG a oH
—_ Vg i) ro) °
: gef28 2 2
=) Name of Plant Soe s List of Counties
i and Habitat eS ZOZR Dm Ss With Recordst
JSOETACEAE
EX Isoetes braunii Dur. 1 0 0 1913 1935 67
Lake. Quillwort
EN Isoetes engelmannii A. Br. 4 1 1 1838 1973 11*, 12%, 50, 67
Pond & Lake. Quillwort
SPARGANIACEAE
TH ae ata aa.
(Engelm.) Moro 6 1 4 1894 1962 3, 8t, 22*, 5Ot, 677, 77+
Marsh’ Se: eed
EN Sparganium Gicwes
pum Rydb. 3 1 2 1892 1969 I1t, 25*, 867
Bog. Bur-reed
POTAMOGETONACEAE
EN tac i filiformis
0 1 1939 1970 G2+
Lake. Filiform Pondweed
t The se refer to the counties as listed in Table 2. An asterisk after the faci
indicates a record previous to 1900; a dagger after the ene ae one a record sinc

1950.

oe
rH

Potamogeton friesii
a 3 2 0 1895 1901 22*, 62*, 77
Lake. Pondweed
Potamogeton gramineus
4s 6 4 2 1888 1970 22*, 267, 487, 62%, 75%,
Pond & Stream. Pondweed 85"
Potamogeton hillii Morong 1877 1913 4%, 67
Pond & Stream. Pondweed
Potamogeton perfoliatus

i)
a
[—

is 1 0 1898 1898 62%
Lake. Pondweed
Potamogeton praelongus

Wul

fen. 5 1 1 1898 1969 4, 22, 62", 76, 867

Lake. White-stem

Pondweed
Potamogeton pulcher

Tuckerm, 6 3 1 18388 1970 4, 11%, 12*, 25, 407, 85*
Pond. Spotted Pondweed

Potamogeton richardsonii

Rydb. 8 5 2

. . 1835 1970 4, 107, 16*, 22*, 43,
Lake. Red-head pondweed G62*r, 67%, TT
oe robbinsil
2 2 1894 1966 22%, 67+, 77**

ae Ppaaoad
Potamogeton spirillus

Tuckerm. 6 2 4 189- 1971 4*, 28*, 477, 507, 67%,
Pond. Pondweed 784
Potamogeton strictifolius

r. benn. 1 0 1895 1898 22%
Lake, Pondweed
Potamogeton tennesseen-
sis Fern. 0 2 19382 1967 157, 40, 82+

Pond & Stream. Pondweed
Potamogeton vaseyi
ins 3 0 0 1900 1935 4, 67, 78
Lake. Pondweed
Bie sear et zosteri-

formis Fern. 10 6 4 1835 1970 11¥, 22%, 28, 45%,
Lake. "Flat. stem Pond- 62*7, 64*, 677, i ya
NAJADACEAE
Najas gracillima (A.Br.)

Magnus 3 2 0 1898 1918 62*, 67, 85*

Lake. Naiad

JUNCAGINACEAE
Triglochin eae L.
Fen. Arrow-g

Triglochin panes L. 6 1
Fen. Arrow-grass

for!
=
w

1835 1972 11*t, 12", 43, 46%, 76*%,
77*

Ww

1840 1965 11%, 22*, 49, 71, 72%,
76%

E

Z

ae
an

ab

an

E

Z

E

~

T

=

T

=n

EN

E

*

TH

T

xq

T

x

Scheuchzeria palustris L.
Bog.

ALISMATACEAE
Echinodorus rostratus
(Nutt.) Engelm.
Mudflat. Bur-head
Sagittaria australis
(J.G. Sm.) Small
Mudflat. Arrowhead
Sagittaria cuneata

heldon

Mudflat. Arrowhead
Sagittaria graminea

Pond. Grass-leaf Arrow-
head

pecitlana eens Willd.
ns (Muhl.)

Marsh. Common Arrow-
head

GRAMINEAE
Calamagrostis inexpansa
A.
Fen. Northern Reed
Glyceria acutiflora Torr.
Swamp. Manna Grass
Glyceria borealis (Nash)
Batchelder
Pond. Northern Manna
Grass
Glyceria grandis S. Wats.
Bog. Tall oe Grass
Givesie melica
(Michx.) F.T. "Hubbard
Swamp. Manna Grass
Glyceria pallida (Torr.)

Pond. Manna Grass
Zizania aquatica

7

oO

aL. 15
Marsh. Annual Wild Rice

CYPERACEAE

Carex alata Torr.
Bog & Marsh. Sedge
Carex aquatilis Wahl.
Marsh. Sedge

bo

for)

bo

—

w

oo

bo

ra)

~1

we

or)

w

ome]

fo]

1840

1905

1875

1894

1838

1961

1946

1842

1925

1842

1889

1840

1833

1887

1879

1973

1973

1972

1972

1951

1967

1967

1971

1925

1969

1961

1971

1972

1899

1939

29

3, 4, 23, 28%, 45*7, 47%,
67

31, 657, 667, 71

1t,

12*,. 13°,
20%, oe 40+, 53; 667

6, 227, 40, 45, 62* 7, 85*

11*, 18*, 48, 88

157, 347, 417

227, 487, 717
387, 45, 597
48

4+, 157, 287, 677, 74,

76"+, 857

4", 18",
43*, 67%,

23, 287, 40,
TT LON

18, a8t,
677, 76%,

381, 404, fi?

77+, 78t,

‘ 12, 22%, 37*, 43, .
48*, 57", 62", 64%,

a. 76, 77, 79%

18*, 43*, 76", 7%, 85*

6%, 22%,
48%, 62"

43*, 46%, 47%,

30

EEN

TN

eX

Fry
ip
SA

KX

Bog & Fen

Carex aurea Nutt.

Fen & Bog. Sedge
Carex baileyi Britt.
Marsh. Sedge

Carex bebbii (Bailey)

Carex atherodes Spreng.
lge

Bog. Sedge

Carex brunnescens
(Pers.) Poir.

Bog & Swamp. Sedge

Carex cephalantha
(Bailey) Bickn.

Bog. Sedge

Carex oo
Shutt

Fen. ae

Carex cryptolepis Mack.

Bog & Fen. Sedge

Carex docomposita Muhl.

Bog.

Carex oa Schrank
Bog. Sedge

Carex flava L.

Fen ge

Carex haydenii Dew
Fen & Marsh. Sedge
Carex howel Mack,
Bog. Sedge

Carex interior Bailey
Fen & Bog. Sedge
Carex sastoCaEDe Ehrh.
Bog. Sedge

Carex limosa L.

Bog. Sedge

Carex oligosperma Michx.
Bog. Sedge

Carex projecta Mack.

Bog. Sedge

Carex pseudo-cyperus L.

Bog. Sedge

Carex retrorsa Schwein.

Marsh. Sedge

Carex sartwellii Dew

Fen. Sedge

Carex oe Willd.

Fen. Sedg

Carex shi aminea Willd.
Marsh. Sedge

He

o

eo

oO

co

lor)

bo

bo

bo

—

bo

_

—_)

S

=

—)

oS

=

i]

—_

—)

—

—)

—

=)

oS

—

—

bo

1897

1898

1934

1893

1871

1883

1840

1871

1838

1840

1840

1894

1892

1886

1889

1842

1898

1894

1890

189-

1893

1840

1935

1972

1970)

1934

1935

1915

19138

1935

1971

1910

1935

1966

1922

1958

1950

1938

1945

1968

1912

1922

189-

1934

1965

1963

197, 22°, 627, 85*

4*, 22", 48, 777

43", 45, 78

28, 43°, 47", 67*, 76
6", 11*, 20*, 71, 87
lv, 4, 48*, 487, 67%
Or
25%, 31%, 45%
22*, 25, 43%, 45%, 47%,
67, 76, 77*
11*i
22*, 47*, 67, 78
287, 40, 4 5 AT
64, 67, 76*, 77 a”
157, 18+, 22*, 31, 40,
43*, 47", 7
45", 47%, 70, 76%, 77%,
85
3, 4%, 23, 45, 85*
20%, 677
43%, 47*
22* 45
48*
22*, 487, 71

11*+, 22*, 49, 76

374, 40%

EX

EN

T

a

T

en

f6

an]

T

an

fh

se)

T

x

T

an)

ft

an

TH

TH

E

Zz

T

mm

Carex suberecta (Olney)
Britt

ritt.

Marsh. Sedge

Carex trisperma Dew.

Bog. Sedge

Carex viridula Michx.

Fen. Sedge

Cladium mariscoides
(Muhl t

Fen. Twig- Rush

Cyperus diandrus Torr.

Mudflat. Umbrella Sedge

Cyperus engelmannii

eud.
Mudflat. Umbrella Sedge
Eleocharis compressa
Sulliv.
Fen. Flat-stemmed Spike-
rush

Eleocharis elliptica Kunth
Fe ik

Eleocharis intermedia
Schultes
sls a Matted Spike-

ee olivacea Torr.
Bog & Mudflat.
Olivaceous Spike-rush
Eleocharis pauciflora
(Lightf.) Lin
Fen. Spike-rush
Eleocharis quadrangulata
(Michx.) R. & S.
Marsh. ay -angled
Spike-rush
Eleocharis rostellata
r

orr.
Fen. Spike-rush
Eriophorum virginicum L.
Bog. Tawny Cotton-grass
Eriophorum viridi-cari-
natum (Engelm.) Fern.
Bog. Cotton-grass, Bog-
cotton
Hemicarpha micrantha
(Vahl) Pax
Mudflat
Rhynchospora alba (L.)
Vahl

Fen. White Beak-rush

~]

a
S&S

So

>

for)

~J

oS

yy

oo

=

He

co

w

or

bo

y

1898

1871

1893

1893

1898

1897

1840

1892

1840

1894

1929

1891

1892

1887

1884

1928

1840

1963

1938

1972

1969

1969

1971

1969

1969

1974

1970

1967

1973

1973

1973

1969

1967

1971

G*, 11*, Wt, 23, 25%,
, 36, 71

29+, 31

3, 4, 28*, 43°, 47, 67*,
77*, 78, 86%

11", 22*+, 624, 774

11*7, 22*1, 28, 481, 67
88

157, 20*, 22+, 47, 487,

62T

4, 6*t, 227, ae 46*¥,

487, 621, 76, 85%

He 227, 20%, , 487,
, olf, 62° +f, — 71,

rt

7, 22*+, 47%, 48*+, 627,

877
15f, 207, 23, 25*, 297,
457, 487, 6277 ; , 8bt
11}, 18*, 627, 77*, 86T
11, 71, 867

li, 3, 8, 40+, 44%, 67
77%

11*¥, 224, 387,
46*¥, 55, 677, 764, 77+
3, 4* 20° , 28*, 39, 45*7,
47%, 671 5 10; 76t, 85
4, ae . Ty ey ees
43%, 4 , 674, 76, 77*+

4, 40, 487, 65, 867

4, 11*t, 22", 28*, 38%,
39, 40, 45*, 47*, 67%,
70, 76, 77*+, 85%

EX

E

Z

T

x

T

en)

E

Zz

E

rs

E

Z

T

<=

T

<=

E

Z

T

=

E

va

E

a

E

~

E

a

T

rm

Rhynchospora globularis
hapm.) toe

Fen. Beak-ru

Scirpus Hetil ae Fern.
Marsh.

Scirpus cick Fern.

Bog. Bulrush

Scirpus purshianus Fern.

Mudflat. sh

Scirpus smithii Gray

Mudflat. Bulrush

Scirpus torreyi Olney

Marsh "US

Scleria pauciflora Willd.

Fen, Nut-rus

Scleria verticillata Willd.

Fen sh

ARACEAE

Calla palustris L.

Bog. Wild Calla

LEMNACEAE

Lemna valdiviana Phillipi

Pond. Pale Duckweed

Wolffia ee C.H.
Thomps

Pond. Pointed Wolffia

Wolffiella floridana (J.D.
Sm.) C.H. Thompson

Pond. Star Wolffiella

XYRIDACEAE

Xyris difformis Chapman

Bog. Yellow-eyed-Grass

Xyris tor Sm

Mudilat. Yellow. -eyed-

ERIOCAULACEAE
Eriocaulon septangulare
With.

Mudflat. White-buttons,
Duckgrass

PONTEDERIACEAE
Heteranthera reniformis
R. & P.

Mudflat. Mud-plantain

JUNCACEAE

Juncus alpinus Vill.

ush

bh

2

bo

So

bo

ww

ww

_—

i]

—_

on

a

_

ww

i—)

So

bo

—)

o

bo

1897

1898

1887

1885

1895

1889

1897

1840

1837

1901

1932

1906

1890

1892

1913

1837

1840

1946

1973

1954

1973

1970

1896

1960

1969

1971

1958

1973

1930

1891

1961

1915

1849

1972

22*, 48, 71
387, 427, 62*, 70*, 85

6*, 18, 43%,
651, 77%, 78,
147, 40+, 43*,
66, 77*, 82+
62*}

47, 517,
85

62*, 65,
22*, 43*, 62*
ly, 22*, 40, 48, 66, 71

11*t, 12, 22*f, 24, 25*,
297, 65, 67+, 717

aPy,. 181, 287, 39°, OF 1,
POP ty bee ee

131, 43

40, 477, 677, 837

23, 45, 64, 67, 77

28*, 67*

277, 40, 487, 73, 87*

67, 77

31", 57*

18*, 22t, 43*, 48, 62*1

Zz

a0)

=

x

Zz

~*~

a

ee)

Zz

a0

q

Juncus greenei Oakes &

Tuckerm.
Mudflat. Rush

Juncus interior Wieg.
. Rush

Marsh

Juncus subcaudatus (En-
gelm.) Coville & Blake
h. Rush

Marsh.

LILIACEAE

Tofieldia glutinosa
(Michx. rs.

Fen. False Asphodel

IRIDACEAE

Iris brevicaulis Raf.

Marsh. Leafy Blue Flag

ORCHIDACEAE
Arethusa bulbosa
Bog. Are

L.
thusa; Dragon’s
Cypripedium candidum
Willd

illd.
Fen. White Lady’s-slipper
Habenaria eee

(Willd.) Hoo
Bog. White pepe
Orchid

Habenaria ciliaris (L.)
R.

Bog. Orange or Yellow
hid

Fringed Orc

Habenaria leucophaea
: . Gra

Nutt.) A
Bog. Prairie White
Fringed Orchid

Spiranthes lucida (H.H.
Eat.) Am

; es
Fen & Marsh. Shining
Ladies’ Tresses

Spiranthes romanzoffiana
Ch

am.
Fen. Hooded Ladies’
Tresses
SALICACEAE
ae bebbiana Sarg.

g & Fen. Long-beaked

Willow

Salix candida Fluegge

Bog & Fen. Sage-leaf
Willow

oo

bo

bo

S

i)

Oo

oo

bo

487
49, 727
377, 657, 717

11*+, 12*, 467, 48, 57*,
67+, 76*t, 77

6*, 197, 25, 31*, 35, 36,
51, 62, 65, 71, 80, 81%,
837

45*+, 47*, 67*+

11*, 12*, 22*1, 48, 57*,
677, 78, 88

4*, 28, 48, 67*, 77*
1, 26*, 48, 48*, 71, 73
GF, dae; 22) 20 Oe

Tt, 1, 13%, 18*, 29%,
43, 48, 52%, 66, 67%,
71t, 77+, 78

4*}, 22", 28, 43%, 52*,
764

4*, 29%, 26%, 357, 39,
43*, 47*, 48*, 62*, 77*,
87*, 8g*

15+, 22*, 26", 67+, 76*,
i 88"

867,

T

=

T

as

T

an

E

Z

T

on

T

a0)

T

an,

Salix pedicellaris Pursh 12
Bog & Fen. Bog Willow

Salix petiolaris J.E.
Smith 8
Bog & Fen. Willow
Salix serissima (Bailey)
Fern.

Bog & Fen. Autumn
Willow

x Salix subsericea (An-
derss.) Schneid.

MY RICACEAE
Myrica pensylvanica
,oisel, 3
Boe Bayberry
BETULACEAE
Betula pumila L. i
Fen. Low Birch
URTICACEAE
Pilea fontana (Lunell)
Rydb.
Marsh. Clearweed
POLY GONACEAE
Polygonum pensylvani-
var. eglandu- 2
losum Myers
Mudflat. Smartweed
CHERATOPHYLLACEAE
Ceratophyllum echinatum
Gray
Pond. Hornwort, Coontail
NYMPHAEACEAE
Nuphar variegatum
Im.
Pond. Yellow Water-lily,
Spatter-dock
RANUNCULACEAE
Ranunculus pusillus Poir. 4
ars Mudflat.
Water Crowfoot
CRUCIFERAE
Armoracia aquatica
ieg

(Eat.) 6

Stream. Lake Cress

or

w

c=

_

w

sy

bo

ol

o

1891

1879

1899

1899

189-

1835

1900

1940

1913

1894

1889

1969

1969

1969

1969

1961

1972

1974

1975

1974

1970

1950

1936

3, 4, 11, 28+, 39, 45%,
64*, 67*, 76, 77+, 85°
86"

6*, 1st, 22*, 26*, 48*,
677, 777, 86, 87*

15+, 67+, 764, 777, 86*

4* 11*, 47", 774
4, 67%, 77

ty, 22 67, 76%, 77°,
867,

15%,
7

f, 424, 67+, 72%, 76%,
+ ‘ . y

22%, 62%

4+, 107, 25, 397, 40,
471, 627, 67,. 70

22", 487, 627

4, 13, 277, 71

16*, 45%, 48, 49%, 64%,
5

TH

a

an]

E

Zz

T

a8)

T

as

T

se)

T

x

T

a0

E

Zz

Zz

E

E

a

T

xq

EN

Rorippa_ sessiliflora

(Nutt.) Hitche. 7 #1
Mudflat. Yellow-cress
SARRACINEACEAE

Sarracenia purpurea L. 14 9
Bog. Pitcher-plant

DROSERACEAE
Drosera intermedia
Hayne 4 |]
Bog. Sundew
ROSACEAE
Potentilla palustris (L.)
14

Bog. Marsh Cinquefoil

CALLITRICHACEAE
Callitriche terrestris

Mudflat, Water Starwort
Callitriche verna L.
ond & Stream. Starwort

RHAMNACEAE
Rhamnus eae L’Her. 10 3
Fen. Buckthor

HYPERICACEAE
Hypericum kalmianum L. 6 4
Fen. St. John’s-wort

HALORAGACEAE
Myriophyllum exalbes-

cens Fern. 12 10
Pond & Lake. Water-

milfoil
Myriophyllum hetero-

phyllum
Pond & Stream. Water-

milfoil
Myriophyllum verticil-

atum 2
Lake. Water-milfoil

UMBELLIFERAE
Hydrocotyle americana L. 12 6
Swamp. Water-pennywort

Hydrocotyle umbellata L. 4 2
Pond. Water-pennywort

ichx. 1

aay

ay

—

ws

raw)

bo

~]

w

=

fon)

bo

1878

1835

1899

1838

1890

1888

1840

1891

1891

1835

1838

1885

1891

1970

1971

1967

1960

1969

1971

1972

1960

1972

1973

1840

1971

1971

1}, 31*, 547, 627, 657,
66, 71

’

a —

3, 4%, 20*, 28*7, 39,
45+, 46%, 47*, 67%, 70,
(Oh; 77%, g5* 86%

287, 43, 48, 85%

3, 4*, 11*, 187, 287, 38,
39, 45%, a7*, 67%, ise
eo; 78, 85*

137, 21, 20; 31, 31, 40;
60, 677, 71, 3

4, 107, 18*, 25, 28%,
A ON

47, 11*7, 157, 18*, 28,

43", 481, 671, 764, 777
22*+, 26+, 48*, 62*, 76%,
i

6*, 18%, 22*, 23%, 38%,
45*, 46", 624 , 64%,
67%, 76%,

4, 11, 36, 45*, 467, 78,
807

11*, 67*
15+, 18*, 287, 381, 43%,
Uy, 16"), 7,

457, 677,

4, 287, 67*1, 76

EN

EX

ERICACEAE
Andromeda _ glaucophylla
Link

Bog. Bog-rosemary
Chamaedaphne calycu-

lata (L.) Moench
Bog. Leather-leaf
Vaccinium oxycoccos L.
Bog. Small cranberry
PRIMULACEAE
Hottonia inflata Ell.
Pond. Featherfoil
GENTIANACEAE
Menyanthes trifoliata L,
Bog. Buckbean

SCROPHULARIACEAE
Gratiola virginiana L.

Mudflat. Hedge-hyssop
Gl seas viscidula Pen-

Mudflat.

LENTIBULARIACEAE
Utricularia

Michx
Fen & Bog.
Utricularia

Hayne
Fen & Bog. Bladderwort
Utricularia mir

en & Bog. Bladderwort

Hedge-hyssop

cornuta

Bladderwort
intermedia

PLANTAGINACEAE

Plantago Sages Lam.
Stream. Plant

CAPRIFOLIACEAE

oars Sey Gronov.

. Twin

VALERIANACEAE

Valeriana ciliata Torr.
Gray

Fen. Valerian

COMPOSITAE
Megalodonta beckii
(Torr.) Greene

Lake. Bur. marigold

bo

fom]

es

bo

fod

a

bo

ew)

_

ww

bo

a

—)

—

_

1878

1871

1889

1891

1835

1897

1932

1839

1893

1893

1835

1900

1837

1860

1929

1970

1961

1974

1970

1973

1974

1964

1972

1972

1973

1924

1964

1966

4, 28*, 76, 77, 85"

4, 18%, 20", 22, 28*%,
45,604, 1081. (7%) 80%,
86"

3, 28, 45%, 47*, 674,
76, 77*, 85"

4*, 734

3, 4, 11%, 25, 28, 39,
43%, 45°F, 67, 76, 77%,
85*

13, 22", 27%, 401

274, 40%, 73%

10), 97% 85

11, 43", 677, 761, 85"

11*t, 157, 45, 67

BG oo
49

46, 47%,

48,

37

Table 2. ee List of Ohio Counties With Numbers of Imperiled Species
County.

n Each
Number of Number of
Imperiled Imperiled
Number County Species Number County Species
1 Adams 8 45 Licking 27
2 Allen 0 46 Log 10
3 Ashland 9 47 Lorain 27
4 Ashtabula 37 48 Lucas 33
5 Athens 0 49 Madison 5D
6 Auglaize 12 50 Mahoning 3
ff Belmont 2 dl arion 3
8 rown 2 52 Medina 3
9 Butler 0 D3 Meigs 1
10 Carroll 2 54 Mercer 0
11 Champaign 35 By) Miami 1
12 ‘lark 10 D6 Monroe 0
13 Clermont D 57 Montgomery 5
14 inton 1 08 Morgan 1
15 Columbiana 14 59 Morro 1
16 Coshocton 1 60 Muskingum 0
17 Crawford 0 61 oble 0
18 Cuyahoga 17 62 Ottawa 31
19 Darke 2 63 Paulding 0
20 Defiance 6 64 rry 6
21 Delaware 1 65 Pickaway 8
22 Erie 47 66 ike 6
23 Fairfield 7 67 Portage 60
24 Fayette 1 68 Preble 0
25 Franklin 14 69 Putnam 0
26 Fulton 7 70 Richland 5
27 Gallia 4 71 Ross 17
28 Geauga 28 72 Sandusky 3
29 Greene 4 73 Scioto 4
30 Guernsey 0 74 Seneca 1
31 Hamilton 6 75 Shelby 0
32 Hancock 0 76 Stark 43
33 Hardin 0 77 Summit 49
34 Harrison 1 78 Trumbul 14
30 enry 2 79 Tuscarawas 4
36 Highland 4 80 ion 2
37 Hocking 4 81 Van Wert 1
38 Holmes 8 82 Vinton 2
39 ron 8 83 Warren 3
40 Jackson 20 84 Washington 0
41 Jefferson 2 85 ayne 29
42 Knox 2 86 Williams 15
43 Lake 30 87 Wood 0
44 Lawrence 0 88 Wyandot 6

Table 3. Counties with the Largest Numbers of Imperiled Species.

Rank County Number of Imperiled Species Portion of the State
1 Portage 60 Northeastern
2 Summit 49 Northeastern
3 Erie 47 Northwestern
4 rk 43 Northeastern
5 Ashtabula 37 Northeastern
6 Champaign 30 West-central
i Lucas 33 Northwestern
8 Ottawa 31 Northwestern
9 , 30 Northeastern

10 Geauga 28 Northeastern

11 wicking 27 entr

12 Lorain 27 Northeastern
APPENDIX

Those species of very limited occurrence in the state which we consider
as non-indigenous (non-native) to the flora of Ohio are enumerated in part 1
of the appendix. In certain situations, our concepts of which species are
non-indigenous differ somewhat with previously published statements, hence
the need for this separate list. For those individuals who consider some of
these species in this list as indigenous (native), they would have to be
placed in table 1. In part 2 of the appendix, we list all of those species
known to us which have been reported for Ohio as having a very limited
occurrence in the state, but which have been eliminated from consideration
because of the reasons that are stated for each species.

Appendix Part 1. Annotated List of Rare Aquatic Plants
Non-Indigenous in Ohio.

Number of Counties with Records
Number of Counties with Records
Year of Earliest Known Record
Year of Latest Known Record

Total Number of Counties

S
308
: _
w 7
Ss) w
® oO
=~ cB)
Name of Plant fe List of Counties
and Habitat OAH With Recordst
MARSILEACEAE
Marsilea quadrifolia L. 2 0 1 1941 1970 287, 25

Pond. Water Clover

SALVINIACEAE

Azolla caroliniana Willd.
¢) Water-velve

NAJADACEAE

Najas marina L.

d. Naiad
GRAMINBAE
Calamagrostis cinnoides

Muhl.) Bart.

Marsh. Reed Bentgrass

CYPERACEAE

Eleocharis caribaea (Rottb.)
.F. Blake

Mudflat. Spike-rush
Eleocharis wolfii Gray
Fen. Spike-rush
Ericphorum gracile W.D.J.

Bog. Cotton-grass

Scirpus saximontanus Fern.

Mudflat. Bulrush

JUNCACEAE

Juncus diffusissimus Buckl.

Marsh. Rush

NYMPHAEACEAE

Cabomba oo Gray

Stream. Fanwort

ELATINACEAE

Elatine brachysperma Gray

Mudflat. Waterwort

LYTHRACEAE

Lythrum hyssopifolia L.

Mudflat. Loosestrife

ONAGRACEAE

Ludwigia decurrens Walt.
(Jussiaea decurrens
(Walt.) DC.)

Mudflat. Primrose-willow

rho leptocarpa (Nutt.)
Ha

(a ea leptocarpa Nutt.)

Mudflat. Primrose-willow

HALORAGACEAE

Myriophyllum aquaticum
(Vellozo) Verdc.

—

—

1901

1949

1931

1936

1954

1933

1935

1935

1951

1951

1949

1974

1971

1972

1971

1935

1955

1974

1973

1959

227, 25, 31, 43%,
627, 67, 76%, 7
227, 627

45*

627

71

25

65+

407, 447, 66%,
43, 777, 78%
657

657

li, 81, 277, 44

Tf. Ot

737

48%

9+

5)

(M. brasiliense PaMiness:)
Pond & oe am. Parrot’s
Feather
UMBELLIFERAE

Hydrocotyle ranunculoides

Marsh. Water-pennywort

PRIMULACEAE

Centunculus minimus L.

Mudflat. Chaffweed

MENYANTHACEAE

Ny aa pevtatum
1.)

(Gm
oe

COMPOSITAE

Senecio glabellus Poir.

Fiat eheaet

2 1959 1971
0 1935 1935
0 1930 1930
1 1926 1974

107, 387
40, 65
4

627, 68

Mudflat. Butterwort
Appendix Part 2. Species Excluded
Species Locality Reference Reason for Exclusion
Hippuris vulgaris Ohio Muenscher (1944) No specimen
L. located
Lemna minima Paulding County — (1937), No specimen
hillipi Braun (1967) located
Cuyahoga County Walters (1950) No specimen
located
Ohio Muenscher (1944) No specimen
located
Lemna perpusilla Mercer County Hicks (1937) Mercer County
Torr. eae not
Knox and Mercer’ Braun (1967) locat Knox
Counties County oe
is L.

Mayaca aubleti
Michx,

Myriophyllum al-
terniflorum
Myricphyllum
humile (Raf.)
Morong

Peplis diandra
Nutt. (Didiplis

Auglaize County

Ohio
Ohio

Ohio

Lake County

Gleason (1905)

Muenscher (1944)

Muenscher (1944)

Muenscher (1944)

Schaffner (1932),
Jones (1943),

ceca re-
orted according
to Gleason (1952,
Vol. 1, p. 377)

No specimen
located

No specimen
located

No specimen
located

Specimen is
Callitriche

diandra (Nutt.) Blackwell (1970) heterophylla

Wood)

Podostemum Pickaway County Althaus (1967) Specimen is

ceratophylum Riccia fluitans
ichx,

REFERENCES

ALTHAUS, R. A. 1967. aia members of the Papaveraceae, Fumariaceae, Ca
ridaceae, Resedaceae, Sarreceniaceae, Droseraceae, and Podostemaceae in Ohio. M.S. Thesis,

ANONYMOUS. 1974. Rare i Endan gered Plants Native to Texas. 3rd edition. Mimeo-
graphed. Rare Plant Study Center, The University of Texas, Austin. 12 pp.

AYENSU, E. S. and. others. a Report on aera and Threatened Plant Species
of the United States. Presented to the Congress of the United States of America by t
Secretary, oo oT Smithsonian Institution, Washington, al 200 pp.
Sle 75 for the of the Committee on Merchant Sar and Fisheries, Serial

94-A, cae oa re Session, House Document No. -51. United States Govern-

ment Printing oa Washington, D.

BLACKWELL, W. , JR. 1970. The Cychbaceke of Ohio. Ohio J. Sci. 70: 346-352

sere: E. L. fee Te oe [of Ohio]. Cat-tails to Orchids. With Sie
Gra by Clara G. Weishaupt. The Ohio State University Press, Columbus. 464

EAIRBROTHERS, D. E. and M. Y. HOUGH. 1973. Rare or Be one Read see

New Jersey. New Jersey State Museum Scienc ce Notes No “ren 53

GLEASON, H. A. 1905. Notes from the Ohio State Herbarium, wv en "Nat. 6: 397- 398,
«1952. The New Britton and Brown Illustrated Flor the Northeastern amiey
- States ui Adjacent Canada. Volume 1. [The New "York See Garden, New York].

ae
or)

ae L. E. 1937. The Sees of Indiana. Amer. Midl. Nat. 18:
. SWIS, F

. F. KEEFE, .#H.

VAN, editorial committee. oe Rare & Endangered Species of Missouri. Missouri De-
partment of Conservation ee United States Department of Agriculture Soil Conservation
Service. Pages not numbere

JONES, C. H. 1943. Studies in Ohio. floristics. Il. Rare aan of Ohio. Castanea 8: 81-108.
UDD, J. H. TAUB. The effects - ecological ieee on Buckeye Lake,
Ohio, with emphasis on aaa bass and aquatic ee plants. Ohio Biol. Surv.,
ee Notes pp.

OWDEN, R. M. i960 co in the marsh flora since 1901, pp. 263-267. Im A vascular
flora Z Winou Point, Ottawa and Sandusky Counties, Ohio. Ohio J. Sci. 69: 257-284.

MOORE, D. L. 1973. Changes in the aquatic vascular plant flora of East Harbor State
Park, homily Coun Ohio, since 1895. M.S. Thesis, The Ohio State University, Colum-
bus.

MUENSCHE R, W. C. 1944, Aquatic Plants of the United States. Comstock Publishing Co.,

York. 374 pp. eo 1948, 1964, 1967)
SCHAFENER, J. H. 1932. Revised catalog of Ohio vascular plants. Ohio Biol. Surv. 5:

87-21 ae . ‘ -

STUCKE 971. Changes of vascular aquatic flowering plants during 70 years in
Put-in- ae aren Lake Erie, Ohio. Ohio J. Sci. 72: 321-342.

TUCK E. 1974. Threatened native plan ts of Arkansas, pp. 39-65. In Charles T.

R, G. 4.
Sais Arkanses Natural st a State of Arkansas. Arkansas Department of Planning,
eat M. B. 195 re Sepidiien and Lemna minima in Ohio. Ohio J. Sci. 50:

WE STUCKEY. eh The changing distribution of the genus
Najas eee in Ohi: Ohio J. Sci. 71: 292-302

A NEW SPECIES OF RHYNCHOSPORA
(CYPERACEAE) FROM SOUTHWESTERN
GEORGIA

ROBERT KRAL
Department of eiagy, Vanderbilt University
ashville, Tenn. 37235

In 1947, during his extensive floristic survey of southwestern Georgia, Dr.
Robert Thorne (now at RSA) discovered in Baker County what appeared to
him to be a new species of Rhynchospora (see Castanea 16: 35. 1951). Speci-
mens from this collection, together with some other Cyperaceae, have been
sent to me by Dr. Thorne for definitive study. The Rhynchospora is indeed a
novelty, and is described as follows:

Rhynchospora thernei Kral, sp. nov.

Planta perennis glabra. Rhizoma squamulosa breve ramosissima implexa.,
Culmi plures caespitosi filiformes 10-15 cm alt., 0.2-0.3 mm lati, infime tere-
tiusculi vel obtusanguli sursum trigoni, scabridiusculi. Folia culmo breviora
filiformia 10-13 cm longa, laminis basibus planis superne trigonis vel canali-
culatis scabridiusculis. Cymae 2-3 valde distantes perlaxi, anguste-turbina-
tae, simplices vel semicompositae, pauciramosae, paucispiculosae. Pedun-
culi filiformi scabridiusculi vel laevi. Spiculae lanceolato-ovoideae vel fusi-
formes acuminatae, 2.5-3.0 mm longae, 1-2-nucigerae. Squamae ovatae sub-
scariosae ca. 1.5 mm longae, anguste-acutae, integrae vel minute erosae,
breve mucronatae, lampro-aureo-ferrugineae. Setae 6 minute-antrorsae-
scabrae subtiles usque ad 14-24 nuceis aequantes. Stamina 1-2, anthera ca.
0.2. mm longa. Achenium (cum tuberculo) ca. 1 mm longum _ subobovato-
ellipticum, biconvexum, reticulatum, Mmarginatum, cancellatum, cancellis
verticalis rectangularis. Tuberculum breviconicum, acutum, ca. 0.15 mm
longum,

Type. U.S.A. Georgia. Baker Co.: damp sand of Emory University Field
Station Experimental Area near Mossy Pond, 23 June 1947, Robert F. Thorne
4881. Holotype RSA. Isotype V

This species, still known only from the type locality, is named after its
discoverer, whose extensive field work during the 1940’s, particularly in
southwestern Georgia, has contributed much to our present knowledge of
the southeastern flora

the description andicates. R. thornei is a low, very slender-leaved
baa -culmed sedge superficially very much resembling either R. pusilla (R.
intermixta) or R. divergens. Like them it has quite small spikelets, of simi-
lar coloration. However, it has a definite perianth of bristles and a smaller
number of florets per spikelet and therefore belongs in the Section aie

SIDA 7(1): 42-50. 1977.

43

chospora”’ Griseb. rather than in the Section Psilocarya (Torr.) Clarke to
which R. pusilla and R. divergens belong (see G. Ktkenthal, Vorarbeiten zu
einer Monographie der Rhynchosporideae, Botanische Jahrbucher 74: Heft
3, 1949 and subsequent numbers for treatment of these!). If the excellent
key to series of Eurhynochospora done by S. Gale (1944) is applied to the
preblem, the new beak-rush is found to belong to series ‘‘Rariflorae’’ Gale.
This series contains, in Gale’s treatment, but 2 species, namely R. rariflora
(Michx.) Ell. and R. stenophylla Chapm. The former is widespread in the
southeastern United States, the Caribbean and Central America; the latter
is rare and local from N.C. south into §.C. with disjunctions in northwest
Florida and southern Alabama. While R. thornei is a lower, more slender
plant than either of these, the vegetative characters agree rather well, par-
ticularly those of scape and leaf and those of the formation of dense mats
through the close interweavings of short, scaly-rhizomatous offshoots.

However the case, there is wide divergence of opinion between Gale,
Kiikenthal and many other cypcrologists when it comes to placement of
species in sections or series of the subgenus ‘‘Diplostylae’’ Bentham, par-
ticularly when it is realized that almost all authors concentrate mainly on
akene and perianth bristle characters. In light of this difficulty and because
of my own inexperience with the genus on a broad scale, I shall present
below a small treatment of those few North American Rhynchospora that
most closely resemble the new species.

1. Perianth bristles present; plants forming mats of tufts by ee ia
ing, scaly rhizomes; akene, weer tubercle, 1 mm long o
. Akene and tubercle ca. 1 mm long; akene body distinctly ed
plants not more than 1.5 dm a cea not longer than 3 mm
ae a G thornei Kral
2. Akene and tubercle mostly 2-3 mm long aken ne “body not distinctly
margined; plants rarely as low as 2 dm; ae 3.5-5.0 mm long.

3. Perianth bristles rarely exceeding middle of akene body, never reach-
ing to tubercle base; spikelets 3.5-4.5 mm long; eee short-triangu-
lar rariflora Ell.

3 Perianth bristles mostly reaching at least the tubercle le spikelets
ca. - mm long; tubercle narrowly triangular-acuminate, 1.0-1.5 mm
lon 3. R. stenophylla Chapm.

1. Perianth bristles absent: ‘plants lacking short, branching scaly rhizomes;
akene including tuber cle not longer than 0.6 mm
4. Akene body nearly smooth, aca reticulate

=

4. R. divergens M. A. Curtis
4. Akene body transversely rugose and finely Sa lined
5. R. pusilla M. A. Curtis

1. RHYNCHOSPORA THORNEI Kral, Fig. 1.

The plant densely tufted, perennial, glabrous, forming mats by means of
short, ascending, scaly-rhizomatous offshoots. Lowest leaves shortest, most-
ly sheath; longest leaves 10-13 cm long, 0.2-0.3 mm broad, with blades
spreading-ascending, proximally flat, then grading upward from involute to
semiterete or channelled, then trigonous except for the somewhat flattened

ee;
BD es

"a
4 os

ts:

, aa, GE
<5 LP

‘igure 1. Rhynchospora thornei. a. Habit sketch. b. Inner surface of cau-
line leaf at sheath orifice, mi ade, and apex. c. Inflorescence. d. Akene,
showing perianth bristles, cross-section of body, and body with surface
markings. From Thorne 4881.

45

narrow apex, there scabrous-edged. Culms mostly 10-15 cm long, somewhat
flexuous, filiform, mostly 0.2-0.8 mm broad, proximally obtusangulate, multi-
costate, becoming mostly sharply trigonous distally. Cymes mostly 1-2, if
2, distant, each subtended by at least 1 foliaceous, setaceous bract, narrowly
turbinate, open and with few spikelets, the peduncles few and of varying
lengths, these simple or with 2 or 3 spikelets each. Spikelets lance-ovoid or
fusiform, 2.5-3.0 mm long, acuminate, usually 2-flowered, each subtended by
9-3 lanceolate, barren bractlets shorter than the fertile scales. Fertile scales
ovate, subscarious, ca. 1.5 mm long, acute, entire or slightly erose, 1-nerve

and slightly keeled, the mid-nerve excurrent as a short mucro, the surfaces
smcoth, lustrous golden-brown. Perianth bristles 4-6, antrorsely minutely
barbellate, the shortest vestigial, the longest shorter than the akene body.
Stamens 1, (—2), the anthers ca. 0.2 mm long, short-linear. Akene (including
tubercle) ca. 1 mm long, the body ellipsoidal, biconvex, a lustrous pale
brown, produced along the edge into an evident though narrow, wirelike
roargin, the surface minutely but evidently cancellate, the cancellae nar-
rowly to broadly rectangular, vertically oriented in wavy horizontal rows
(more isodiametric and irregularly arranged proximally and distally!) and
also transversely rugulose; tubercle short-conic, barely 0.15 mm long.

2. RHYNCHOSPORA i cena (Michx.) Ell., Sk. Bot. S. Car. &

Ga. i., ig.

Schoenus rariflorus oe , Fl. Bor.-Am. I. 1803.

Phaeocephalum rariflorum House, Am. ae Nat. 6. 202. 1902.

Perennial, glabrous, densely tufted from closely forking scaly compact

rhizomes. Leaves linear-filiform, the lowest mostly sheathing, the longest
with blades mostly 10-15 cm long, between 0.7 and 1.0 mm broad, proximally
flattish, the backs with several raised nerves, often becoming strongly revo-
lute upward and triangular, acute-angled and sulcate toward the apex, there
with the margins finely scabrid. Culms (1.5-) 2.0-5.0 (-6.0) dm tall, wiry-
filiform, rather weak, terete and multicostate proximally, distally with a
strong sulcus below the nodes. Inflorescences mostly 2, distant, each cyme
open, simple or compound the (usually) few spikelets divaricate-ascending
on peduncles of various lengths, each prophyllate; lowermost cymes sub-
tended by a foliaceous bract that is longer, the uppermost by a similar
bract but this equalling or shorter, and each cyme if compound with smaller
bracts subtending each cymule. Spikelets 3.5-4.5 mm long, narrowly ovoid
or fusiform, acuminate, mostly with (—3) 2 fertile scales these subtended by
2-3 (-4) smaller, barren bracts. Fertile scales smooth, ovate to oblong, 2-3
mm Icng, obtuse to broadly acute, subentire, scarious, a rich golden-reddish-
brown, the backs rounded, with 1 prominent medial nerve, this excurrent as
a short mucro. Perianth bristles mostly 6, dark reddish-brown, upwardly
scabrous, unequal, the longest rarely longer than mid-akene. Stamens usual-
ly 2, anthers ca. 2 mm long, narrowly linear. Akene with tubercle ca. 2 mm
long, ellipsoidal to obovoid, biconvex to somewhat plano-convex (the side

46

facing inward often flattish), the body yellowish or reddish-brown, trans-
versely wavy-rugose, with several irregular rows of vertically oriented,
nearly linear, fine cancellae. Tubercle grayish-crustose, blunt-triangular, ca.
0.8 mm high, somewhat dorsiventrally flattened,

Q.

\

Figure 2. Rhynchospora rariflora. a. Habit sketch. b. Inner surface of leaf
at sheath orifice; back of leaf at same level; leaf blade apex. ec. Two views
of inflorescence, that on left at fruiting stage, that on right showing a portion
of a younger inflorescence (slightly enlarged above scale!). d. Akene and
stylar apparatus, cross-section through body. From Kral 24154.

47

Moist to wet sands or sandy peats of low savanna, bogs, seeps, Coastal
Plain from New Jersey and southeast Virginia southward into peninsular
Florida and westward into Texas with isolated stations inland as far as
northern Georgia and Middle Tennessee, scattered in Caribbean and Central
America.

R. rariflora, together with the much more local R. stenophylla both tend
to form small ‘‘sods’’ of densely caespitose tufts and thus in habit are very
similar to R. thornei.

3. RHYNCHOSPORA STENOPHYLLA Chapm., FI. S.U.S. p. 525. 1860.
Similar in almost all ways to R. rariflora but taller, the culms often

from 6 to 9 dm tall. Spikelets narrower, more narrowly lanceolate in out-
line, longer, ca. 5 mm long, usually with but 1 fertile floret. Akene with
tubercle longer; akene body ca. 1.5 mm long, narrowly biconvex, sculptured
as in R. rariflora; tubercle ca. 1.5 mm long, narrowly triangular-acuminate,
somewhat dorsiventrally flattened. Perianth bristles usually 6, unequal, an-
trorsely barbellate, arising from a lobed short-hairy basal joint and some
nearly reaching or exceeding the tubercle tip.

In habitats similar to those of R. rariflora but with known range restricted
to the eastern parts of the Carolinas, northwestern Florida and southern
Alabama. Fig. 3.

4, RHYNCHOSPORA eee Chapm. ex., M.A. Curtis, Amer. Journ.
ae r, 2, 7, p. 409. 1849. Fig
divergens Chapm ee, 1850.
Ses seis divergens House, i Mi dl. Nat. 6, p. 201. 1920.

Perennial, glabrous, densely tufted through close-set, thick-based offshoots.
Leaves linear-filiform, the lowest sheathing, the sheaths tubular, multi-
costate, scarious on side opposite the blade, the blades 5-15 cm long, 0.3-0.5
mm wide ascending or spreading, flat just above the sheath, upwardly be-
coming strongly involute, then channelled, then sharply trigonous toward
the narrowly acute apex, there scabrous on the angles and on the slightly
flattened tip. Culms linear-filiform, (1.5-) 2.0-5.0 (-6.0) dm tall, erect or
spreading and arching from the clump, proximally terete, multicostate, up-
wardly becoming channelled below the sheaths on the side opposite the
leaf blades, Inflorescences (1-) 2-3 (—4) narrowly to broadly turbinate, each
subtended by a long-linear-filiform-bladed bract this exceeding the cyme.
Cymes compound, paniculiform, the spikelets numerous, the peduncles sca-
brid, of varying length, a few primary peduncles simple, most zig-zag, with
each node producing a lanceolate bract, a chaffy-tubular prophyll, as a
shert-stalked spikelet and a prolongation of the primary peduncle. Spikelets
asymmetrically lance-ovoid or fusiform, ca. 3 mm long, acuminate, usually
subtended by 2-3 lanceolate sterile scales shorter than the 4 or more fertile
scales, the lowermost sterile scales abscising, then progressively the fertile
abscising (akenes often persisting after) as the spikelet elongates, thus the
spikelets producing several fruits but keeping their original length and

Imm.

\ i) pt ELLE
2) pail” TACO
7 set “tl di IY,
i we qc
yi le

— ae

See
= =

6) i

Figure 3. Rhynchospora stenophylla. a. Habit sketch. b. Inflorescence.
c. Akene and cross section through body. From Godfrey 56716.

Dy
CSP

5

wk

PUL Mom nt
POI i
sal pO

Mller
le,
”

Figure 4. Rhynchospora divergens. a. Habit sketch. b. Inner surface of
leaf at sheath orifice; blade apex. c. Inflorescence. d. Inflorescence branch,
slightly enlarged. e. Akene and cross section through body. From Kral 47024.
f. R. pusilla—akene and cross section through akene body. From Kral 17578.

30

shape. Fertile bracts ovate, ca. 1.5-2.0 mm long, acute, entire, all scales a
lustrous reddish-brown, with the single mid-nerve excurrent as a short
mucro. Perianth bristles absent. Stamens 1, ca. 1 mm long, the anther nar-
rowly linear, ca. 0.2 mm long. Akene including tubercle 0.7-0.8 mm long,
obovoid, short-stipitate, strongly biconvex, with a low but evident, rounded
margin, a lustrous pale brown, finely but evidently reticulate, the arcoles
mostly rectangular, vertically oriented in irregular rows. Tubercle de-
pressed-conic, apiculate, ca. 0.1 mm _ long.

Sands, sandy-peats and silts of swales in savannas, ditchbanks, seeps,
bogs and pondshores in the Coastal plain of the U.S. from S.C. southward
through peninsular Florida and west with decreasing frequence into Texas;
in the Bahamas, Cuba, Santo Domingo. Flowering and fruiting from June
to frost, all year sou

.. RHYNCHOSPORA PUSILLA i a ex. M. A. Curtis, Amer. Journ.

Se. ser. 2. No. 7, p. 409. 1849. Fig.
R. pusilla Chapm., FI. S.U.S. p. a 1860.

intermixta C. Wright in Sauv., Fl. Cub. p. ae 1873.
Phacocephalum pusillum House, Am. Midl. Nat. 6, p. 201. 1920.

Similar to R. divergens Curtis in all ways except the fruit. Akene ca. 0.5
mm long, oblong-obovate, short-stipitate, biconvex, narrowly margined, pale
brown to nearly white, lustrous, strongly transversely wavy-rugose, the
concavities between the ridges minutely longitudinally striate.

Moist sands, sandy-peats and silts of low meadows, savannas, bogs, seeps
and damp shores, mostly in the Coastal Plain from N.C. southward into
peninsular Florida, westward into eastern Texas, with localities scattered
inland to northern Alabama: Carribean. Flowering from June to frost, all
year in frost-free climates.

Curtis (l.c.) appears to have been sent a manuscript description by Chap-
man, from which he extracted a mere diagnosis, but the identification is un-
mistakable. Chapman, (l.c.), subsequently published this and R. divergens
without reference to the Curtis publication. Most authors have treated R.
pusilla as R. intermixta.

REFERENCES
CHAPMAN, A. W. 1860. Flora of the southern United States. New

ork.
CURTIS, M. A. 1849, a and rare plants, chiefly of the en Amer. J. Sci. Arts
XX

ser. 2, Lee

GALE, SHIRLEY. on beac section a enruene in Canada, the United
States and the West Indies. Contr. Gray Herb. CL

HOUSE, H. D. 1920. A consideration of certain genera proposed by Ehrhart. Am. Midl.

KUEKENTHAL, G. 1949-1952. Vorarbeiten zu einer Monographie der Rhynchosporideae.,
Bot. Jahrb. Syst. Band 74, 75,

THORNE, ROBERT F. 1951. Vascular plants previously unreported from Georgia. Cas-
tanea 16 8.

VASCULAR FLORA OF THE RAGLAND HILLS
AREA, FORREST AND PERRY COUNTIES,
MISSISSIPPI

KEN E. ROGERS
Department of Horticulture, University of Maryland
College Park, Maryland 20742

ABSTRACT. Thorough field studies of the Ragland Hills area, Forrest and
Perry counties, Mississippi began in 1969 and were continued until autumn

consideration. About 250-300 acres of these ravines, although disturbed by
man, probably represent the largest and best remnant of the beech-magnolia
forest in Mississippi. One-thousand nineteen species of vascular plants, vari-
ously distributed among 452 genera in 130 families, are recorded for the

The Ragland Hills, located in Forrest and Perry Counties in south Mis-
sissippi, are approximately 12 miles south-southeast of Hattiesburg (Fig. 1).
It is a maturely dissected area and includes some of the most rugged terrain
in south Mississippi. The area is divided by U.S. Highway 98 and is bounded
by Caraway Creek on the east, the Leaf River on the north, Weldy Creek
on the west, and the Camp Shelby Military Reservation on the south. Ap-
proximately 3600 acres, encompassing parts or all of fifteen sections (Sec-
tions 9, 10, 13, 14, 15, 16, 21, 22, 23, 24, 25, 26, R12W, T3N, Forrest County;
Sections 18, 19, 30, R11W, T3N, Perry County) are included. All land is pri-
vately owned with about 90% of the total acreage being in Forrest County.

Most of the state of Mississippi lies within the Physiographic Province
known as the Gulf Coastal Plain (Fenneman, 1938). Most of the southern
one-third of the state, including Forrest and Perry counties, is in the sub-
division referred to as the Longleaf Pine Region or Pine Hills (Lowe, 1921).
This subdivision is named for the extensive forest of pine which is dominated
by the Longleaf Pine. This region is a maturely dissected highland sloping
to the southeast, with elevations ranging from approximately 500 feet to 100
feet (U.S. Geological Survey, 1956)

The Ragland Hills are a part of the north-facing slope of the divide be-
tween the Leaf River and Black Creek. They are erosional in origin having
been formed by the Leaf River and its tributaries which drain the northern
half of Forrest and Perry counties. Elevations range from about 300 feet
along the crest of the divide to a little below 100 feet along the Leaf River
(U.S. Geological Survey, 1949).

Geologically most of the area is underlain by clays of Miocene Age be-
longing to the Hattiesburg Clay Formation. Portions of the crest of the
divide are underlain by sands and gravels of the Plio-Pleistocene Citronelle

SIDA 7(1): 51-79. 1977.

D2

Formation. These formations are in part overlain by Pleistocene to Recent
terrace and alluvial deposits (Mississippi Geological Society, 1969; Foster,
1941).

The soils are primarily sands and sandy loams which are rather sterile
(Personal Communication, Soil Conservation Service; Vanderford, 1962).
The pH ranges from 4.5 to 5.5. Deep colluvial silty clay loams and organic
muck soils are associated with the swamps and poorly drained depressions.

The mean annual precipitation is between 60 and 62 inches. It is rather
evenly distributed throughout the year with the autumn being the driest sea-
son. The average annual temperature is approximately 67° F with the average
July temperature between 81° F and 82° F and a mean January temperature

<
=z
~]
2
i YArIOO MADISON

BCOTT HEWTON LAUDERDALE

_———}

_—_—_—_—
JASFER

CLARKE
A
WAYNE
COVINGTON
FORREST PERRY GREENE
PEARL RIVER GEORGE
STORE
JACKSON

HARRISON

HARCOCK
<4 oe oS

Fig. 1. ihe in which study was conducted is indicated by a circle in For-
»st and Perry counties, Mississippi.

03

of 51° F to 52° F (McWhorter, 1962). Temperatures range from approxi-
mately 7° F to 106° F.

The effects of human activities are manifest throughout the area. A few
dwellings occur but these are restricted mainly to areas adjacent to the
highway. There are no dwellings in the ravines. Cutting of the forest began
in the 1800's and has continued in varying degrees to the present. Currently
both timber and pulpwood are being harvested. A number of access roads
crisscross the area, and several sand and gravel pits occur. Recent ex-
ploration for oil has resulted in further damage to the forests.

The study area includes not only the ravines but portions of the adjacent
divide crest as well as the floodplain of the Leaf River. The plant communi-
ties consequently are quite diverse and include river swamps, hardwood
bottoms, savannahs, flatwoods, sandhills, bays, and upland forests of pine,
mixed pine-hardwood, and hardwood. Also included are the various habitats
associated with human activities such as those around dwellings, artificial
lakes, in pastures and fields, and along roads and trails. Several species of
aquatic plants are found in oxbow lakes, swamps, and artificial lakes and
ponds.

The most floristically rich and outstanding plant community is the beech-
magnolia forest associated with the mesic ravines along the north slope of
the divide. Approximately 250-300 acres of the steepest ravines (N4, Sec.
24, R12W, T3N, Forrest County; NW1!4 of the NW, Sec. 19, R11W, T3N,
Perry County) have escaped to some degree the ee that has befallen
the native vegetation surrounding these ravines. This is probably the largest
and richest remnant of the beech-magnolia forest in Mississippi.

I have attempted to provide as nearly as possible a complete record of
the vascular plants which occur in the ravines as well as those which are
associated with the adjacent uplands and bottom lands. The collection of
specimens and observations of the plants and the area began in 1969 and
were continued until autumn 1973. These have been made during all seasons
of the year. The annotated list which follows is arranged alphabetically by
families and within families. In most cases the scientific names are in ac
cord with those used in the Manual of the Vascular Plants of Texas (Correll
and Johnston, 1970).

Herbarium specimens are deposited in the Herbarium of the University
of North Carolina. Duplicate specimens, when available, were distributed to
various other herbaria including GH, MISSA, MO, NY, SMU, US, TENN, and
VDB.

The pteridophytes and seed plants of the area are distributed among 452
genera in 130 families. Of the 1019 species which were found in the area,
approximately 131 (12.9%) are intreduced. A number of the introduced spe-
cies are of the type that persist around old homesites and probably do not
reproduce themselves in this region.

The five largest families which together comprise almost half of the total
number of species are: Gramineae (153), Compositae (136), Cyperaceae

54
(79), Leguminosae (73), and Labiatae (27).

ANNOTATED LIST OF THE VASCULAR FLORA OF RAGLAND HILLS

ACANTHACEAE
Dicliptera brachiata (Pursh) Sprengel. Locally abundant along the Leaf River. Native.
Justicia ovata (Walter) Lindau. Infrequent in swampy places. Native.
Ruellia caroliniensis (Walt.) Steudel. Occasional in pinelands, dryish woods, and old fields.

ative.
ACERACEAE
Acer rubrum L. (Incl. A, Sang var. ie (H. & A.) Sarg.). Frequent in a variety
of woodland habitats. Native.

Acer saccharinum L. Occasional along floodplain of river. Native.

AIZOACEAE

Mollugo verticillata L. Common in disturbed places. Introduced.

ALISMATACEAE
Echinodorus cordifolius (L.) Griseb. Infrequent in shallow water
swamps. Native.
Sagittaria graminea Michx. Infrequent in shallow water of ditches, ponds, lakes, streams,
and swamps. Native.
Sagittaria ie ‘folie Willd. Locally abundant in shallow water and wet places. Nativ
Sagittaria longirostrata
ative.

tt

»f ponds, lakes, and

p
Micheli) J. G. Smith. Infrequent in shallow water and wet hes

—

AMARANTHACEAE
see ies philoxeroides (Martius) Griseb. Common in ponds, lakes, swamps, and some-
times in wet waste places. Introduced.
Pleat hbybridus L. Common in cultivated and fallow fields.
ge alder viridis L. Common in cultivated and fallow fields, ane along roadsides.

ed.

AMARYLLIDACEAE
Hymenocallis occidentalis (LeConte) Kunth. Frequent in low moist woods.
Polianthes virginica (L.) Shinners. Infrequent in open pine and mixed ae 'N, ative,

ANACARDIACEAE

Rhus copallina L. Common in mixed woods, pinewoods, ing old fields. Native.

Rhus glabra L. Occasional along em ondland borders. Natiy
Rhus radicans L. Common in woodlands, disturbed Be ne along aece ae
Rhus toxic — ndron L. Common in upland mixed w eons and oe elds.
Rhus vers » Occasional or frequent in low wet plac ive
ANNONACEAE

Asimina parviflora’ (Michx.) Dunal. Occasional in usually dryish mixed woods. Native.

APOCYNACEAE
Trachelospermum difforme (Walter) Gray. Infrequent in low woods. Native.

AQUIFOLIACEAE
Hex ambigua (Michx.) Torr. Infrequent in upland mixed woods. Native
Hex amelanchier M. A. Curtis. Infrequent or rare in low rich woods. Native.
Hex coriacea (Pursh) Chapman. Common in moist or wet low places. Native.
Hex decidua Walter. Occasional in swamps, ravines, and alot ong streams. Native.
Ilex glabra (L.) Gray. Common in open neuioods and moist places. Native
Hex longipes Chapman. Infrequent or rare in upland areas. Nati

Ilex opaca Aiton. Frequent in mixed woods and ravines. Native.

Ilex verticillata (L.) Gray. Occasional in low woods. Native.

Ilex vomitoria Aiton. Common in low woods, pinelands, and along wood borders. Native.
ARACEAE

Arisaema dracontinm (L.) Schott. Infrequent in ravines and low woods. Native

Arisaema tripbyllum (L.) Schott. Os in low woods and ravines. Native

Orontium aquaticum L. Infrequent in sandy streams and open ponds. Native

ARALIACEAE
Aralia spinosa L. Frequent in mixed woods and disturbed ravines. Native.
ARISTOLLOCHIACEAE
ge uate I oe A. bastata Nutt.). Occasional in mesic upland woods and
er slopes of ravines.

P
Ae hia tomentosa Sims. “Occoxdl along floodplain of river. Nativ
Hex ee arifolia (Michx.) Small. Frequent in moist or dryish mixed soak: and ravines.
at

ASCLEPIADACEAE
ponies amplexicaulis Smith. Infrequent in dry mixed woods. Native.
A s humistrata Walter. Infrequent in sandy loam soils of open, dry upland woods.

re,
oe a Elliott. Infrequent in Pinevoods and along roadsides. Native.

Asc a s per s Walter. Infrequent in shallow water of swamps. Nati
Ascle pias naeee L. eee in open anee woods and borders of low woods. Native.
ie lepias verticillata L. Infrequent along roadsides and in open upland mixed woods. Native.

Cynanchum laeve (Michx.) Persoon. Infrequent in open mixed woods. ‘Nati v

BALSAMINACEAE

Impatiens capensis L. Locally abundant in wet places bordering cypress-tupelo swamps.
Native

BETULACEAE
Alnus eta eee Willd. rae Sone streams. Native.
Betula nigra ent along river floodplain. Nativ
pee eran Walter, aa in mesic woodla ae Native.
Istrya virginiana (Miller) K. Koch. Frequent in rich woodlands. Native.

BIGNONIACEAE
Eee se shaase L. Frequent in mesic woodlands. Native.
Campsis radicans (L.) Seemann. Common along roads, trails, fences, and open woods. Native.
oe pane Walter. Infrequent along streams, roadsides, ‘od near dwellings. Native,
eae indicum L. Infrequent in open moist places. Introduced.
Myosotis macrosperma Engelm. ee at the edge of old fields. Native.
Onosmodium pee (L.) DC. Infrequent in open pinelands. Native.

BRASSICACEAE

Cardamine bulbosa (Schreber) BSP. Local in moist deciduous woods. Native.
Cardamine hirsuta L. Frequent in disturbed places. oo

Lepidinm virginicum L. Frequent in disturbed places. Nat

Raphanus rapbanistrum L. Infrequent in disturbed aes iainee.

BROMELIACEAE

Tillandsia usneoides L. Common in deciduous (rarely evergreen) trees. Native.

BURMANNIACEAE

Apteria aphylla (Nutt.) Barnhart. Local in mesic woods. Native.

CACTACEAE

Opuntia drummondii Graham. Rare in sandy well-drained soil. Native.
CALLITRICHACEAE

Callitriche terrestris Raf. Infrequent on moist sandy loam in open areas. Native.

CAMPANULACEAE

Lobelia cardinalis vcal in wet soil bordering swamps and ponds. Native
Lobelia glandulosa oe Infrequent in moist soil bor

ordering open swamps. ‘Nat tive.
Lobelia puberula Mi Frequent or infrequent in moist to dryish open places. Native.
Lobelia spicata Lam. Infrequent in open woodlands. Native.
Triodanis biflora (R. & P. ireene. Frequent in disturbed places. Native.
Triodanis per Nie Frequent in disturbed places. Native.

Hiata (L.) evw. ur
Wahlenbergia marginata (Thunberg) DC. Common in disturbed places. Introduced.

CAPPARIDACEAE
Cleome houtteana Raf. Occasional escape from cultivation to disturbed places. Introduced.
CAPRIFOLIACEAE

Lonicera japonica TV es Occurs in a variety of habitats, sometimes overrunning native
vegetation. renee ed.

~
a

Ont sempervirens os Occasional in oak-pine woodlands. Native.

Sambucus seo Me) L. Locally abundant in open moist places. Native
Dur »» Frequent in alluvial woods and low woods. Native.

1 s and w ds. Native.

wet eee Native.

ry upland woods. Native.

Locally numerous in swamps
Viburnum af dulce Raf. Occasional in

CARYOPHYLLACEAE

Cerastinm ie nea oo Frequent in disturbed places. Introduced.
G.

Sagina decumbens (Ell) ° Frequent in pomee places. Native
Silene pe ts Ls ee along roadsides. Native
Silene ovata Pursh. Local on upper slopes of ravines. a

ative.
SteHaria media (L.) Cyrill. Occasional in disturbed places. Introduced.
CELASTRACEAE
Enonymus americanus L. Occasional in low woods. Native
CERATOPHYLIACEAE
Ceratophyllum demersum L. Locally abundant in swamps. Native.
CHENOPODIACEAE
Chenopodium album L. a in cultivated
Chenopodium ambrosioides
Chenopodium pumilio R.

and other disturbed places. Introduced.
Infrequent in disturbed places. Introduced.
a. Rare in disturbed places. Introduced

CISTACEAE
ee oe (Walter) Michx, Infrequent In| open a Native.
chea minor L. Occasional along pond ies and in clearings. Native

pie tefl Mich Occasional in dry

sandy loam soils of upland areas. Native.
Lechea villosa . Frequent in clear

rings a upland open woods. Native
COMMELINACEAE

ae caroliniana Walter. Infrequent in disturbed places. Native

Commelina communis L. Common in disturbed eee usually near dwellings. Sep eaiaia
rae i — een f. Infrequent in disturbed places near dwellings. Nat
Commelina 1 L. Infrequent in disturbed bles Native.

Cc Gronalea oe reinica L,

ocally numerous in rich woods. Native.
Tradescantia obiensis ae

Sees along ae Nat

COMPOSITAE

Acanthospermum australe (Loefl.) O. Keze. Locally numerous in sterile soil in open places.
Introduced.
Ambrosia oe L. eae or sometimes abundant in cultivated fields, fallow
fields, and urbed places. Native
Aesbao . LL. - aie abun dane slong floodplain of river.
Antennaria fallax Greene. Infrequent or locally abundant on upper stones of ravines. Native.
Nat

Aster concolor L. Occasional in pine or mixed woods. Native.
Aster gee L. Frequent in mixed and pinewoods, along roadsides and various disturbed

places. ve.
Avte interflora (L.) Britton, Frequent or infrequent along roads, trails, and in mixed
ods.
ree anes L. Occasional in ps pine or mixed woods.
Aster paludosus Ait. Infrequent in usually dry soil of open an one Native
Aster patens Ait. Frequent or eee in upland mixed woods and pinewoods. Native.

pe pilosus Willd. Infrequent along roads, wood borders, ang edges of fields. Native.
- pracalfus Poir. Infrequent in low open places. Native
pe simplex Willd. Localized in low moist woods and Aaa river floodplain. Native.
Aster undulatus L. Infrequent in open upland mixed woods. Native.
Baccharis halimifolia L. Frequent ae roads and in low moist places. Native.
Bidens bipinnata L. Occasional in disturbed places. Native.
Bidens discoidea (T. & G.) Britt. Occasional in moist low ae Native.
Bidens frondosa L. Frequent in low woods 4 = moist hers Nativ
Bidens mitis (Michx.) Sherff. Common t places. Native.
Bigelowia nudata (Michx.) DC. Occasional in moist eer Native.
Bolo nia diffusa Ell. Frequent or infrequent along roads and in low usually open places.
Native.
Brintonio discoidea (Ell.) Greene. Frequent in mixed woods. Native.
Cacalia clliottii (Harper) Shinners. Infrequent in moist woods and moist open places. Native.
Cacalia eis Nutt. Infrequent in low pinelands. Native.
aie ia fomentosa Vent. Infrequent in open low wet places. Native.
Chrysogonnn virginianum L. Frequent in dry mixed woods and open places. Native.
Cirsi carolinianum (Walter) Fern. & Schubert. Infrequent in open disturbed upland
woods. Native.
Cirsium borridulum Michx. Occasional along roads and in old fields. Native.
Conyza canadensis (L.) Cronquist. Common in disturbed places. Native.
Conyza floribunda H.B.K. oe in disturbed places. Introduced.
ae caren (bC.) T. & G. ee along roads and in other disturbed

*

—

eae ae L. Infrequent in disturbed places. Native.

Coreopsis major Walter. Common in dry open woods. Native.

Coreopsis stenophylla F. E. Boynton. Rare in open disturbed places. Native.

Coreopsis finctoria Nutt. Infrequent in various disturbed places. Native

Coreopsis tripteris L. Infrequent in open low ground. Native.

pica divaricatum (Nutt.) Raf. Abundant in sandy loam or sand of open usually dis-
bed places. Native

Peli — Lae Hasskarh. Occasional at the edges of ponds, lakes, swamps, and in wet
tches,

Pee ae Willd. Frequent in mesic or dry mixed woods. Native.

Elephantopus elatus Bertoloni. Frequent or infrequent in open pinew woods. Native.

Elephantopus nudatus Gray. Occasional in sandy loam soils of pine and mixed woods. Native.

aimee. tomentosus L. Occasional in open pine and mixed woods. Native

Erechtites f haar (L.) Raf. Occasional in disturbed places. Native.

Erigeron annuus (L.) Persoon. Frequent to infrequent along roads and trails. Native.

Erigeron ph Tatelbbrc us L. Infrequent in open disturbed places. Native.

Erigeron strigosus Muhl. ex Willd. Frequent along roads, trails, and in open disturbed places.

Native

palo album L. Infrequent in dry pine or mixed woods. Native.

08

Eupatorium altissimum L. Infrequent in — ungs in dry mixed ia aag Native.
Eupatorium aromaticum L. Frequent in pine and mixed woods. Native

n
Eupatorium re ade (Lam.) Small. ee yon in disturbed plac ative.
Eupatorium coelestinum L. Infrequent in wet ditches and openings in low woods. Native.
Eupatorium en Gee Walter. Infrequent along roads and in other disturbed places.
ative.
Eupatorium fistulosum a Localized in low wet areas, Native.
Eupatorium hyssopifolium L. Frequent. or infrequent in open pinewoods, dry to moist
clearings, and fields. Native.
Eupatorium leucolepis (DC.) T. & G. Open usually low slag not common. Native.
oo bea aera L. Occasional or sometimes common in low woods, borders of
swamps, ds, and lakes, and in wet ditches. Nativ
ite Mie ‘hlov Walter. Infrequent or rare in open pinewoods. Native.
Eupatorium recurvans Serial. Infrequent or locally numerous in moist pinewoods. Native.
Eupatorinm ean L. Common in open pine and mixe -d woods. Native.
Eupatorium semiserratum DC. Frequent or infrequent in moist ae or mixed woods. Native.
Eupatorium serotinum Michx Frequent in open distu hed eed Native.

Euthamia Sige CL: ) ae Infrequent or sometimes locally abundant in moist open
ces. Nativ

Euthaniia epochal Ris G.) Greene. Infrequent in open sandy soil. Native.

Euthamia minor (Michx.) eae Infrequent or sometimes locally numerous in moist open
places. Native.

Facelts retusa (Lam.) Sch.-Bip. Infrequent in pastures and along roads and trails. Introduced.

Gaillardia aestivalis (Walter) H. Rock. Infrequent in dry sandy loam soils of upland mixed
woods. Native.

Gnaphalium ie atum Lam. Infrequent or locally numerous along roads and in fields and
pastures. tive.

Guaphalinm bale ri Britton. Infrequent in dry pinewoods. Native.

Guaphalium obtusifolium L. Fr ei or infrequent in fields open mixed or pinewoods,
and along roads and trails. Nati

Guaphalinm pensylvanicum Willd. lesen! or infrequent in disturbed places. Native.

Guaphalium purpurenm WL. Frequent in disturbed places. Native.

Helenium amarum (Raf.) H. Rock. Infrequent or locally numerous in pastures, fields,
. al iV

g :
Helenium flexuosum Rat. Infrequent in moist places. Nat
Helianthus angustifolius L. Frequent in wet or moist soil in open eek. along roadside
ditches, and edges of ponds and marshy areas. Native.
Helianthus annuus L. Infrequent in disturbed places. i
Helianthus atrorubens L. Infrequent in open mixed woods and at wood borders. Native.
Helianthus i cee: Nutt. Infrequent in open pinewoods. Native.
eee hirs: Raf. — numerous or occasional in open upland mixed woods and
ads trails.
Helantbns mcroceal 7 Ps G. Infrequent and local in mixed woods and at wood

bo

ae. rabuiles (Pursh) T. & G. Occasional in open dry pinewoods.

Helianthus resinosus Small. Infrequent or locally numerous in open eee nee and
at wood borders. Native.

Helianthus strumosus L. Frequent in upland woods and along roads and trails.

Heliopsis gracilis Nutt. Infrequent in openings and at borders of upland eon aie

ative.

Heterotheca oe (Michx.) Shinners. Frequent in sandy soils along railroads, roads,
and in dr oods. Native.

Heterotheca mariana (L.) Shinners. Infrequent in dry woods. Native.

Heterotheca nervosa (Willd.) Shinners. Infrequent or frequent in sandy loam soils of pine-
woods and mixed woods. Native.

Heterotheca hie (Nutt.) Shinners. Infrequent or rare in disturbed places. Native.

Heterotheca subaxillaris (Lam.) Britt. & Rusby. Infrequent along railroads, in sandy fields
and waste places. Native

Heterotheca trichophylla (eines Shinners. Infrequent in dry upland wooded sandhills.
Native.

Hieracium gronovii L. Infrequent in mixed woods and pinelands. Native.

Iva ciliata Willd. Locally = fees railroad. Native.

Krigia dandelion (L.) Nutt. infrequent along roads. Native.

Krigia oppositifolia Raf. ee in ae ae places. Native.

Krigia virginica (L.) Willd. eee along roads, in fields and in lawns. Native.
Kubnia SS L. Infrequent in dr mixed woods. Native.

Lactuca canadensis L. Fre eek or infrequent along roads, rai ge ame a Native.
Lactuca ieee ana (L.) Gaertner. Infrequent in disturbed plac ativ

Lactuca graminifolia Michx, Infrequent along roads in upland . wo mn Nativ

datris spicata (L.) Willd. Frequent or infrequent in low moist open areas, low pinclads,
and ditches. Native.

Liatris squarrosa (L.) Michx. Frequent in sandy loam ee of upland woods. Native.

Liatris squarrulosa Michx. Infrequent or rare in dry woods. Native

u

Marshallia trinervia (Walter) on ex Branner & Coville. Teal i low moist places at
edges of mixed woods. Nativ

si sc anton ce Willd. i cacae at borders of lakes, ponds, and swamps and in low

t pla Nat
Pluchea ae are (L) DC. Frequent or infrequent in marshy places and ditches. Native.
Pluchea foctida (L.) DC. Frequent in low wet places, edges of swamps, and in wet ditches.
ative
ee uvedalia L. Infrequent in rich wo ae eo
Prenanthes altissima L. Infrequent in mesic ravines. ive.
Prenanthes serpentaria Pursh. Infrequent in open m nixed woodlands. Native.

Pyrrbopappus ai (Walter C. Common in disturbed places. Native.

Rudbeckia birt . Frequent along roads, trails, and in sandy woodlands. Native.

a shbte Poiret. Abundant in low moist woodlands. Native

Sericocar pus Pee (Walter) Porter. Frequent in open binned and upland mixed
woods.

Sericocar pus ies (L.) BSP. Frequent or infrequent in dry oe Native.

Silphiune gatesii Mohr. Occasional in dry, open upland mixed woods. Nat

Soli dago ae ssima L. Common in old fields, along roads and trails, and in ae open places.
Native.

Solidago boottii Hooker. Fre equent in moist sandy loam soils in mixed woods. Native.
Solidago caesia L. Infrequent in beech-magnolia woods in ravines. Native.

Solidago gigantea Ait. ae es or locally abundant in low open moist places. Native.
Solidago odora Ait. Fre en dryish woods. Native.

Solidago patula Muhl. ae in low, wet open areas. iv

g streams, and in open wet seis and ditches. Native.

Solidago ulmifolia Muhl. ex Page Infrequent along river floodplain. Native.

Soliva pferosperma (Juss.) Infrequent in lawns. Introduced.

Sonchus asper ( Hill. Infrequent in disturbed places usually near dwellings. Introduced.

Sonchus bili L. Infrequent in disturbed places usually in the vicinity of dwellings.
ntroduced.

ei americana (Mutis) Hieron. var. repens (Walter) A. H. Moore. ce in often
den rowths in ee ditcl ae along _ ban s, ae margins of swamps. Nativ

Taraxacum officinale ggers. Frequent in a variety of disturbed places. (esate ed.

Teh sgonotbee | ie o a eee in ee mixed woods and disturbed woods.

Priva ‘adoration (Walter ex J. F. Gmelin) Cassini. Frequent in pine and mixed woods.
Verbesina pn en Infrequent or rare in open mixed woodlands. Native.
Verbesina virgini . Frequent at the borders of pastures, low woods, and old fields. Native.
Verbesina walteri ee fee uent along floodplain of river. Native.

l

Vernonia altissima Nutt. Infrequent in alluvial woods and low ground along roadsides.
aah

Xanthium strumarium L. Infrequent to frequent along river banks. ie
Umi japonica (L.) DC. Infrequent in disturbed places. Introduce

CONVOLVULACEAE

Bonamia patens (Desr.) Shinners. Occasional in deep sandy soils of upland areas. Native.

60

Cuscuta compacta Juss. Local in moist places on both deciduous woody and_ herbaceous
plants. Native.
Cuscuta gronovii Willd. ex R. & S. Local in moist places on woody and herbaceous plants.

ative.
Dichondra carolinensis Michx. Local in lawns and pastures. Native.
Ipomoea coccinea L. Infrequent in disturbed places. Introduced.

p

Ipomoea hederacea Jacq. Infrequent at edges of streams and low woods and borders of fields.
Ir ase ae

Ipom nosa L. ee along streams, open thickets, and roadside ditches. Native.

I pomoca sales (L.) G. W. Meyer. Infrequent in disturbed places. Native

Ipomoea ae Ls eae in disturbed places. Introduced.

Ipomoea trichocarpa Ell. Infrequent along roads and at wood borders. Native.
i Sea hg (L.) Griseb. Brequent in cultivated ground and various other
irbed places. tive.

CORNACEAE

Cornus Pe Ly ts Infrequent isa streams. Nat
fornus florida L. Common in variou odland h a ative
Cornus eels Lam. Infrequent in swamp i Native.

CRASSULACEAE

Penthorum sedoides L. Local in wet ditches, edges of swamps, and along streams, Native.

CUCURBITACEAE
Cayaponia grandifolia (T. & G.) Small. Infrequent or rare in rich woods. Native.
Melothria pendula L. Infrequent in alluvial woods. Nativ

CUPRESSACEAE
Juniperus virginiana L. Infrequent in mixed woodlands. Native

CYPERACEAE

Bulbostylis barbata (Rottboell) Clarke. Locally numerous in cultivated ground. Introduc
Bulbostylis capillaris (L.) Clarke. Frequent in dry sandy loam aie especially in disturbe d

places. Native.
Hsin te ia AS II.) Fern. Frequent in dry sandy a Native.

abscondita Ma nzie. Common in low rich woods.
aes albolutescens aoe Comm in various moist pone ive.
Carex atlantica Bailey (Incl. C. Fema rae C. incomperta Bickn.). Infrequent at the
edges open swamps. Nati
Carex complanata Torr. Hooks, Fr eae in moist sandy woods and ditches. Native.
Carex crebriflora Wieg. Infrequent in low rich woods. Native.

Carex debilis Michx. Infrequent or eee common along woodland streams and low moist

Carex digitalis Willd. en in rich woods. Native.

Darex flaccosperma Dewey. ie saien in rich wooded ravines. Native.

Carex folliculata L. snes or infrequent in wet places. Native

Carex He nkii Kunth. Locally abund dant in moist places along roads, on river floodplains, and

oa

a

re.

at s of ponds and lakes I
Cares nese Ell. fener in alluvial woods, wet ditches, and at edges of streams.

oe ‘bendals Mackenzie. Infrequent in low woods. Native.

Caren ignota Dewey. Infrequent or rare in ma rich woods. Native.
Carex intumescens Rudge. Frequent i low moist places. Native.
Carex laxiflora Lam. Infrequent in reese eadenel a ravines. Native.
Carex leavenworthii Dewey. Infrequent along roads and in sas Nativ
Carex leptalea Wahlenberg. Frequent in organic muck in low wet woods. Navivs:
Jaren ies Bailey. Infrequent in low woods. Nati

rex lur Wahlenberg. Frequent in various moist ole Native.

Car
Carex ia rgii Schkuhr. Infrequent in sandy woods. Native.
Carex nigro-marginata Schweinitz. Infrequent in beech-magnolia forest in ravines. Native.

Carex oblita Steudel. Infrequent in wet woods and along streams. Native.
Carex Be Schkuhr, Infrequent in rich woods. Native.

Carex pic eudel. Occurs in colonies on the upper slopes of ravines. Native.
Carex reforms ame ca oe or rare in i moist eens Native.
Carex striatula Michx,. nal in rich woodlands. Native.
Carex pee Buckley. sa ar nae in rich woodlands.
Carex tenax Chapman. Locally frequent in wooded sandhills. Native
Carex triangularis Boeckler. Infrequent or rare in low woodlands. Native.
a enusta Dewey. Infrequent in wet woods a along streams. Nativ
arex cs al ii oe Infrequent in lo ich woo ative.
a yperus compress nt or infrequent along roads and in fields.
Cyperus ele eee Infrequent in wet ditches and on ae oY aed Native.

Cyperus esculentus L. Locally abundant in sandy fields Native.
rt iculmi a E ative.

Cyperus globulosus Aublet. Frequent in disturbed places, Native.

Cyperus haspan L. Frequent in wet ditches at edges of ponds and swa es ive.

Cyperus iria L. Occasional in wet cultivated ground and depresvions ie du

Cyperus ovularis (Michx.) Torr. Infrequent along roads, trails, and in dry aro and fields.
Native.

Cyperus plukenetii Fern. Infrequent in dry woods a sandy eae Sy nia Native.

Cyperus psendovegetus Steudel. Locally frequent in moist place

Cyperus refrorsus Chapman. Infrequent or locally numerous mn ae i grounds. Native.

Cyperus rotundus L. Locally common in lawns, sandy fields a ee as Introduced.

lig eee (Torr.) Mattfield & Kikenthal. Locally abundant in moist an. loam
soils about akes, and in ditches. Native.

gies ee — ee quent or infrequent in various moist disturbed places, and at borders
of ponds and swamps. Native.

Cyperus eee ae pe bapag in nae soil. Native
Cyperus virens Michx. Loca fre in various st or wet plac ativ

Eleocharis microcarpa Torr. Occur 2 ithe ee of ee and lakes, ee in water forming
dense mats of attenuated stems. Nat

Eleocharis obtusa (Willd. ) ere. Locally abundant in moist sandy places. Native

Eleocharis fortilis (Link) Schultes. Locally abundant in wet soil in open deciduous weeds
Na

Eleocharis tuberculosa (Michx.) R. & S. Locally frequent in moist to wet sandy loam soils.
Native

Fimbristyis aufumnalis (L.) R. & S. Locally numerous in open disturbed sandy loam soils.
Nat

Fimbrist ylis miliacea se Vahl. Infrequent in cultivated ground and at borders of ponds
and lakes. Introduc

oe puberula “ich Vahl. Frequent or infrequent in open pinewoods, low
grounds, and along ro tive

Fuirena squarrosa Michx. cae . in wet pinelands, moist sandy soil and swamps. Native.

Rhynchospora caduca Ell, Frequent in low sandy woods, swamps, and savannahs. Native.

Rhynchospora cephalantha Gray var. Pee a Fern. & Gale. Locally numerous in low
wet pinewoods. Native.

Rbynchosbora compressa Carey. Local in low pinewoods. Nat
Rhynchospora _cornic ulata (Lam.) Gray. Locally abundant in Ae swamps, and low wet
places. Nat
ee vie A. Dietr. Often abundant in low ground, ditches, and around ponds.
ative

Rbynebospora A ee (Chapman) Small. Common in open moist sandy places low pine-
woods, ditches. Native.

Rincon globularis (Chapm.) Small var. recognita Gale. Frequent in open moist sandy
places. Native.

Rhynchospora glomerata (L.) Vahl. Local in wet ditches and edges of ponds. Native.

Rhynchospora gracilenta Gray. Infrequent to frequent in low pinelands. Native.

ve.
Rhynchospora inex pansa “ hx.) Vahl. Abundant in sandy loam soils of low eer
and along roads. Nativ

62

Rhynchospora miliacea (Lam.) Gray. Infrequent or rare in swamp e ive.
Rhynck ag nixta Britt. ex Small. Locally numerous in low plicwinels and swamp. for-

ests.
Riynchospor hein Britt. ex a faethe in low pinelands and swamps. Native.
Rhynck ene rariflora (Michx.) Ell. Locally numerous in low pinewoods and boggy places.
Ts

Scirpus eeaas var. rubricosus (Fern.) Gilly. Locally abundant in shallow water of
swamps, ponds, and in ditches. Native.
Scirpus koilole pis (oteudel) Gleason, Local in moist sandy open places, frequently in culti-

Scleria ciliata Michx. Locally numerous in pinelands and dry mixed woods. Native.

leria flaccida Sten Locally abundant in open wet woods. Native
ales hae ichx. Frequent in moist sandy loam of woods aise clearings. Native.
uciflora vei ex Willd. Infrequent or locally numerous in moist sandy loam of

plese mixed woods, and in clearings. Native.
Scleria friglomerata Michx. Locally abundant in moist to dry sandy loams of open woods
and clearings. Native

CYRILLACEAE
Cyrilla racemiflora L. Frequent at the edges of open swamps, in low pinelands, ditches,

and alluvial ail Native

DIOSCOREACEAE

1Oist oj woods of ravines. Native.

=

Dioscorea quaternata (Walt.) J. F. Gmel. Scattered it

Dioscorea villosa L. Scattered in) moist rich woods of ravines. Nativ
EBENACEAE

Diospyros virginiana L. Frequent in woodlands and_ fields. Native
ERICACEAE

Epigaea repens L. Rare in dry sandy loam soil of pine-oak-hickory woods. Native.

Gaylussacia dumosa (Andrz.) T. & G. Qccasional in moist to dry pinewoods and clear-
ings. Native.

Ralniiz latifolia L. reas in beech-magnolia for in ravines. Native.

Leucothoe axillaris, (Lam.) D. Don. Local in Fae forest of ravines and along
streams and swamp forests. ative.

Leucothoe racemosa (L.) Gray. Infrequent in low woods. Native

Lyonia ligustrina (L.) DC. Infrequent in low moist woods. Native.

Lyonia lucida au K. Koch. Infrequent or locally numerous in low wet woods or in
shallow swam Native.

Oxydendrum Be pone o ) DC. Occasional or frequent in drier portions of ravines and
upland mixed woods. Native.

Rhododendron canescens ~Ailieh es Sweet. eneaes in low mixed woods. Native.

Vaccinium arboreum Marshall. Locally abundant in dry open woods. Native.

Vaccinium Hees Camp. Frequent in ene and 2 open woods. Native.

Vaccinium elliottii Chapman. Frequent or infrequent in alluvial woods, swampy places, and
ravines. Nativ

esac vane um L. Frequent in mixed woodlands, less frequent in pinelands and ra-
vine

Go eee Ait. Infrequent in open mixed woods. Native.

ERIOCAULACEAE
Lachnocaulon anceps (Walter) Morong. Localized in ditches and low pinelands. Native.

EUPHORBIACEAE
Acalypha gracilens Gray. Frequent or abundant locally in disturbed places. Native
a rhomboidea Raf. Infrequent along roads, trails, borders of fields and oer and
ound dwelli eo a
Poe virginica L, Infrequent in disturbed places. Native.
Aleurites fordit Hemuley. Infrequent escape from cultivation to usually disturbed habitats.
Introduced.

[on

63

pean silence Bs sia Engelm. & Gr Occasional in dry upland woods. leis
ton capitatu . Common in sandy fields a various other disturbed places. |
c se eee - var. septe ‘ntrionalis Muell. Arg. Frequent along roads, in Esllow fields,
and other disturbed habitats. Nat
Crotonopsis elliptica Willd. locally + numerous in open sandy places. Native.

Euphorbia corollata L. Frequent in dry yods. Native.
penis maculata L. (E. supina Raf.) Pecan in open disturbed places, especially culti-
ated . Native.

a He oo. - preslii Guss.) Common in yarious disturbed habitats, especially
cultivated fields. Na

tive.
Phyllanthus caroliniensis “Walter, Infrequent or loca

ly numerous, usually in’ disturbed
places such as fields. Native.
aa wrinaria L. Locally abundant in disturbed areas. Introduced

ee fruticosa (Bartr.) Fern. Frequent in alluvial woods, at borders of swamps, and
occasionally in ravines. Native.

Tragia acest bare in dry deep sand of sparsely wooded upland areas, Native.

Tragia cal in deep sand of Lara Mees sandhills. Native

Tragia ow nae ee or rare at the edges of upland mixed ever Native.

FAGACEAE

Castanea alnifolia Nutt. var. floridana ae Infrequent in upland mixed woods. Native.
Castanea pumila (L.) Mill. var. ashei Sudw. Infrequent in upland mixed woods. Native.
Fagus grandifolia Ehrhare. Connon in pee and low moist woods. Native
Quercus alba L. Frequent in moist or somewhat dry woodlands. Native
Ouercus austrina Small. Rare on banks of river. Native
Quercus coccinea Muench. Occasional in upland aaa woods. Native.
Quercus falcata Michx. Common in dry enone: Native.
Quercus bemisphaerica Bartram. Frequent in us sually gon es Native

nC j n. Frequent oodec ae s.

s daevis Wi :
Quercus laurifolia Michx. Infrequent in low moist woods or wet depressions. Nati
Quercus lyrata Walter. Infrequent in moist woods along streams, or in wet tee
N

a

Quercus piapearelle Ashe. Frequent in wooded sandhills and open ae woods. Native.
Quercus marilandica’ Muenchh. Frequent in dry upland woods. Nativ

Ouercus michauxii Nuttall. Occasional in ravines and low. rich aoe News:

Quercus nigra L. Pieeuent in alluvial woods, ravines and along streams. Native.

Quercus pagoda Raf. Occasional in ravines and low woods. Native

Ouercus phellos L. Occasional in low moist ground, Native.

Ouercus shumardii Buckley. Infrequent in ravines and low woods. Native.

Quercus stellata Wang. Frequent in upland mixed woods. Native.

Ouercus velutina Lam. Infrequent in upland mixed woods. Native.

rENTIANACEAE
Gentiana saponaria L. Infrequent or rare in recently cleared and burned low woods. Native.
ee ee (L.) Pursh. Frequent in low places along roads and in low open woods.
Sibat co diata Ell. Infrequent in moist ground of open pinelands and along roadways.
ve.
ee campanulala (L.) Torrey. Infrequent in open wet places. Native.

GERANIACEAE

Geranium carolinianum L. Frequent in disturbed places, especially roadsides. Native.

GRAMINEAE
Agrostis elliottiana Schultes. Occasional in upland disturbed open places. Native.
Agrostis byemalis (Walter) BSP. Common along roads and trails and in fields. Native.
Agrostis perennans (Walter) Tuckerman. Frequent in old fields, openings in mixed woods,
along roads and trails. Native.
Alopecurus carolinianus Walter. Localized in moist or wet open places. Native.

j=

2

64

Andropogon elliottit, Chapman. eae in rather ar open woods. Native.

Andropogon gerardii Vitman. Infrequ in open dry places. Native.

Andropogon glomeratu (Walter) BSP. iy ace in low epee moist areas. Native.
Andropogon ternarius as Frequent in dry open woods. Native

a Anita . Common in open woods, old fields and paceuees and along roads

an re

rails. Native.
Per aie alee (Michx.) Beauy. Infrequent in open dry pinewoods and = savannahs.

lative.
Aristida eee (Schultes) Kunth. Infrequent in lov — pha Native.
Aristida dichotoma Michx. Common in dry open place Nativ
Aristida lanosa Muhl. ex Ell. Fr requent or infrequent in i ere dein soils. Native.

Aristida longespica Poiret. Frequent in open, sandy places. Native.

Aristida oligantha Michx. Frequent in open, dry ace in sandy loam soils. Nat

Aristida purpurascens Poiret. Frequent in usually dry soil in open pine and vied woods,
open ground, and at wood borders. Native.

Aristida simpliciflora Chapman. Rare in open, dry upland woods peeuye:

Aristida virgata Vrinius. Localized in open, moist sandy places. Native

pa ale oe (Walter) Muhl, Frequent along streams, a of swamps, and in low
Woods. ativ

koa hd legis (Walter) Muhl. meas in boggy, open gna Native.

Avena sativa L. Occasional escz Upe from cultivation to roadsides oduced.

Axonopus affinis C ve Common in open Een along roads and ee ; and | in various other
disturbed plac

Brachiaria Heyl "(Geiseb..) Nash. Frequent in disturbed sandy soils, especially culti-
vated gro und. tive.

é

a)

Briza minor L. oe along roads and various other disturbed areas. Introduced.
Bromus commutatus Schrad. Occasion val or localized along roads. Introduced.
Bromus japonicus Vhunberg. Infrequent along roads. Introduced.
Bromus unioloides (Willd.) H. B. K. Occasional or locally abundant along roads. Intro-
duced.
ee ee (Hackel) Fern. Locally numerous or scattered i
place atiyv
eee aie ‘latifolinm (Michx.) Yates. Occasional in low woods and along. streams.
ative,

1°)

open, dry sandy

Chasmanthinm laxum (L.) Yates. Occasional in open moist woods. Native.

Chasmanthinm sessiliflorum (Poir.) Yates. Frequent or infrequent in deciduous woods.
Native.

Crentum aromaticun (Walter) Wood. Occasi onal in open pinewoods. Native.

Cynodon dactylon (L.) Persoon. Common in various disturbed places. Introduced.

Dactyloctenium acgyptium (L.) Beauv. eee fake in cultivated oe Introduced.

Danthonia sericea Nutt. Frequent or infrequent in dry, open woods. Native.

Digitaria jiliformis (L.) Koeler. Occasional or locally abundant in sandy soil along roads,
railways, and other open places. Native

Digitaria En ee (Schreber) Schreber. Tesegt in open disturbed areas. Introduced.

Digitaria sanguinalis (L.) Sco sal Common along trails, roads, and in fields. roan

Digitaria villosa (Walter) Pers. Occasional in sandy soils in open disturbed places. Nati

Digitaria violascens Link. Occasional in sandy soils in open disturbed places. Native.

Echinochloa crusgalli (L.) Beauv. oe or locally abundant
Introduced.

1 wet places and fields.

Eleusine indica (L.) Gaertner. Common along roads and in fields. Introduced.

Elymus virginicus L. Occasional or locally abundant in moist woods and ditches. Native.

Eragrostis amabilis (L.) Wight & Arnott ex Nees. Infrequent in pastures. Introduced.

Eragrostis cilianensis (All) Lutati. Infrequent in open disturbed places. Introduced.

Eragrostis ciliaris (L.) R. Br. Infrequent in pastures. Introduced.

Eragrostis glomerata (Walt.) L. H. Dewey. Occasional in wet sand or mud along rivers.
Native.

Eragrostis hirsuta (Michx.) Nees. Occasional along roads, trails, and in open woods. Native.

Eragrostis oxylepis (Torr.) Torr. Common in sandy soil in open areas. Native.

Eragrostis abs (L.) Beauy. Frequent or infrequent along roads and in open ground. In-
troduc

65

Eragrostis refracta’ (Muhl.) Scribner. Frequent in open woods and various sandy soils.

ative.
ke spectabilis (Pursh) Steudel. Frequent along trails, roads, and other open areas.
Native.
ara a aie (L.) EI. Locally abundant in open moist Dies Native
Erianthus brevibarbis Michx. Localized in open low places. Nativ

Erianthus on ee ras ex El. Locally abundant in low eae ditches, and other
moist areas. Native
Erianthus gig ae uS (Walter) ae Locally abundant in open moist places. Native

Erianthus strictus Baldwin. Abunda in open wet grounds. Native.
Festuca clatior L. (Inel. F. nee Scribn. ) Infrequent along roads. Introduced.
Gy ie ee ambiguus (ich, BSP. Occasional in open dry woods. Native.
Gymnopogon brevifolins Trinius. Infrequent in open pine and mixed woods. Native.
i sal pusilla a Nurs “Commer in open disturbed habitats. Native.
ydroc hloa caroliniensis Beauv. Localized in mud and shallow water of ponds and_ lakes.

Na re

Imperata brasiliensis Trin. Locally abundant in open disturbed places, principally along
roads. Introduced.

Leersia lenticularis Michx. Frequent or infrequent at the borders of swamps, lakes, and in
ditches. Native.

Leersia oryzoides (L.) Swartz. Frequent or abundant in mud and shallow water of swamps,

oynds, and ditches. Native.

Leersia virginica Willd. Frequent in moist woods and open moist places. Native

Lolium perenne L. (nel. L. multifiorum Lam.) Frequent along roads. Introduced.

Manisuris rugosa (Nutt.) Kuntze. Infrequent in open moist pinewoods. Native

Melica mutica Walter. Infrequent in upland mixed woods. Native.

Muhlenbergia expansa (DC.) Trinius. Common in pine-woodlands. Nat

Muhlenbergia schreberi J. F. Gmelin. Locally abut — in open eee areas. Native.

ce

Oplismenus setarius (Lam.) R. & S. ae in moist woods. Native.

Panicum aciculare Desv. Cor in open sandy nee and open ahi Native.

Panicum anceps Michx. oe along roads, trails, at the . f fields, and in ditches.
Native.

Panicum anceps var. rhizomatum (Hitch. & Chase) Fern. Infrequent in moist pinewoods.

ative.
Panicum SE Ell. Frequent in open dry woodlands. Native
Panicum boscii Poiret. Occasional in dry woods. Native

Panicum commatatom Schultes (Incl. P. joorii Vasey: Frequent in low woods, mixed
woods, and ood borders. Native.
Panicum eee var. ashei (Pearson) Fern. Occasional in dry woods. Native
anicum curtifolinm Nash. Infrequent in open moist places.
Panicum depauperatum Muhl. Infrequent in dry woods. Native
Panicum enone Michx. Frequent in various open ou places and low woods.
Jative

hen un “lichotomun L. (Incl. P. barbulatuin Michx., P. nitidum Lam., P. mattamuskee-
Ashe, P. roanokense Ashe, P. microcarpon taht, P. yadkinense Ashe, P. lucidum
rie . Frequent in mesic and mixed woods and various open areas. Native

Panicum a an Baldw. ex Ell. Occasional in open moist places. Navive:
Panicum gymmnocar pon a Pa abundant in mud and shallow water in cypress-tupelo
swamps and lakes. Nat
Panicum hians Ell. Frequent or infrequent in moist open places. Native.
Panicun 1 lanuginosum Ell. Frequent in open dryish woods, fields, and along roads and trails.
re,

a
Panicum laxiflorum Lam. Frequent in open mesic woods. Native,
Panicum lindbeimeri Nash. Frequent in usually moist open woods and along roads and trails.
Native.
Panicum longifolinm Torr. Frequent in moist open areas and savannahs. Native.
Panicum mutabile Scribn. & Smith. Occasional in dry mixed woods. Nativ
Panicune ne Schultes. Occasional in open dry areas and in chat mixed woods.
atiy

Panicum elie Schultes. Infrequent in moist openings. Native

66

Panicum ramosum L. Infrequent or rare in disturbed places. Introduced.

Panicum ravenelii Scribn. & Merr. Infrequent in open dry woods. Native.

Panicum las Nees. (2. agrostoides Sprengel). Common in moist or wet usually open
places. Nat

Panicum aan a Ell. Common in marshy places, along streams, at edges of lakes and
ponds, and in ditches. Native.

Panicum scabriusculum var. cryptanthum (Ashe) Gleason. Common in low wet open places.
Native.

Panicum scoparium Lam. Occasional or abundant in moist open habitats. Native.

Panicum ee EN. Frequent along roads, trails, and various other open dry or moist
place ative.

Panicum ae peeps Nash. Infrequent at the edges of lakes, ponds, swamps, and in ditches.
Native.

Panicum strigosum Muhl. Infrequent in open moist ground. Native

Panicum fenue Muhl. Cncl. P. albomarginatum Nash, P. aan Nash, P. flavovirens

fash). Common in low, moist ground in open areas. Native

Panicum peau Buckl, Rare along roads. Introduced

Panicum verrucosum Muhl. Occasional or locally abundant in open moist places. Native.

Panicum villosissimum Nash (Incl. P. psendopubescens Nash). Frequent in open dry woods
and along wood borders. Native.

Panicum virgatum L. Frequent or locally abundant in open moist ground. ane:

Paspalum bosciannm Flugge. Infrequent in open usually wet areas. Native

Paspalion ciliatifolium Muhl. Occasional in disturbed places. Nativ

Paspalum di atatin oir. — along roads, trails, and in fields. Introduced.

Badin ah (El : aon Infrequent in mud or floating i water. Native.

Paspalum laeve Michx. (Incl. P. longipilum Nash, P. cireulare Nash). Frequent in open
places such as fields, along roads and trails. Native

ere ee notatum Flugge. Common along roads, eile and in fields ene pastes Introduced.

Paspalum plicatulum Michx. Infrequent in open moist ground. Nativ

— um praecox Walt. nel. Pe as Lam.). Infrequent in moist open pinewooc

1s.
Paspalnn sefaceum Michx. hoe pubescens Muhl., P. debile Michx., P. stramineum
Nash, P. supinim Bose., 1 ong lata LeConte). Oscasional or frequent in ope
woods, ae s, along trails and roads, and in other disturbed places. Nati

Paspalum urvillei Steudel. Common He a. trails, and at ares of fields. Introduced.
Paspalum vaginatum Swartz. Infrequent in faaehes Native.
Phyllos — aurea Riv, Local along streams and in ravines. Introduced.
Poa ant L. Frequent in various disturbed places. Introduced.
Poa pre Muhl. ex Ell. Occasional in ravines.
Poa chapmaniana Scribner. Occasional or frequent in
and trails. Nativ
Schizachyrium scoparia (Michx.) Nees. Common in upland woods, well-drained open
places and along roads and trails. Native.
Schizachyrium fenerum Nees. Common in ae savannahs. Native
ale

awns, vacant lots, and along roads

Secale cereale L. Infrequent escape from cultivation to roadsides. Introduced.

Sefaria geniculata (Lam.) Beauv. Infrequent in moist soil in various open areas, Native.

Setaria glauca (L.) Beauv. Infrequent in disturbed places. Introduced.

Sefaria viridis (L.) Beauy. Infrequent along roads and in vacant lots and railroad yards.
Introduced.

Sorghastrum avenaceum (Michx.) Nash. Infrequent in open dry mixed woods. Native
reas elliottii, (Mohr) Nash. Infrequent in open upland woods and along roads.
ative,
sorghum ges (L.) Moench. Occasional escape from cultivation to roadsides, railroad
yards, and vacant lots. Introduced.
Soe bale (L.) Persoon. Occasional or sometimes common in various disturbed
places. Introduced.
oar Aifotant Sealy Scribner. Occasional in open dry woods. Native.
Sa obtusata’ (Michx.) Seribn. var. major (Torr.) Erdman. ee, Ve moist
zround along roads, an 7 other open areas. Native

~

67

Sphenopholis obtusata (Michx.) Scribn, var. obtusata. Frequent along roads and trails.

Native.

Sporobolus saben bee er) Hitche. Infrequent in open
Q Kunth. Common in pine savannahs. Nativ

sandy loam soil in open pine and

=

dry uaa Native
Shovobolue junc Michx.) Ni
Sporobolus macer eae Hitche. Infrequent or rare in
pine-hardwoc ive.
Sporobolus indicus (L.) R. Br. Frequent in disturbed places. Introdu
Sti Infrequent in well-drained i loam soil in cee oa mined woods. Native.
ative.

S H :
Tridens ambiguus (EIL.) ee Infrequent in moist oper

Tridens carolinianus (Steud.) Henrard. Infrequent in open
Tridens chapmanii (Small) Chase. Infrequent in open upland mixed woods. Native
Tridens flavus Hitche. Common along roads, trails and in open Ris Native
Tridens strictus (Nutt.) Nash. Infrequent in open eually moist ground. Native
Triplasis americana Beauv. Infrequent in dry sandy places. Native

Chapman. Frequent or infrequent in is sandy places. Native

kes purpurea (Walter)
rificum aestivum L. aaa oe escape from cultivation to disturbed ha
Vali myuros (L.) C. C. Gmelin. Infrequent in vacant lots, railroad yards, along roads.
Introduced.

Vulpia octoflora (Walter) Rydberg. Common along roads and
places. Native.

abitats. Introduced.

n various other disturbed

HALORAGACEAE

Myriophyllum brasiliense Camb. Local in swamps, lakes and ponds. Introduced.

Proserpinaca palustris L. Local in marshy places and shallow water. Native.

Proserpinaca pectinata Lam. Local in shallow water of ponds, in ditches, and marshy areas.
ative

HAMAMELIDACEAE

in mixed woods and ravines. Native

es, low woods. Native

Hamamelis virginiana L. Frequent
Liquidambar styraciflua L. Frequent in moist ravin
HIPPOCAST ANACEAE
Aesculus pavia L. Frequent in mesic woods. Native.
HYDROPHYLLACEAE
Hydrolea quadrivalvis Walter. Locally abundant in shallow aoe ang pools. Native.
Hydrolea uniflora Raf. Locally abundance in shallow ponds. Nati

HY PERICACEAE

Ascyrum hbypericoides L. Frequent in open pine and hardwood forests. a

Aseyrum stans Michx. Occasional in open pine and mixed woods. Nat

Hypericum ies nahin W alte Infrequent in low pine-flatwoods. Nodee.

Hypericum drummondii (Grev. & Hooker) T. & G. Frequent along roads and in dryish
open woods ne fields. Naave

Hypeviewm ge ae (L.) BSP. Frequent along roadways, trails and at edges of fields.

ative
aia um gates Lam. Frequent along streams, wet depressions, at the edges of swamps,
an pinewoods. Native.
O jpetes un sion L. Frequent in open wet places around swamps, bogs, ditches, and
ponds ative.
Hoy perienin nudiflorum
Hypericum punctatum Lam. Infrequent in woods an 5 iy
Hypericum sefosum L. Occasional in low pine- flatwoods. Native.
tubulosum Walter. Frequent at the edges of swamps and in ieee woods.
Hypericum virginicum L. Frequent around swamps, marshy areas, and ponds. Native.
Hypericum walteri Gmelin. Frequent in openings in low moist woods. cee :

Michx. ex Willd. Infrequent or rare along = streams. Native.
d fields. Nat
Native.

ILLICIACEAE

Frequent understory shrub in ravines and on floodplains. Native.

Ulicinm floridanum Fllis

IRIDACEAE
Sisyrinchiim angustifolium Miller. Infrequent in moist open woodlands. Native
Sisyrinchium rosulatum Michx. Occasional in lawns and along roads. Introduced.

JUGLANDACEAE
Carya aquatica (Michx. f.) Nutt. Occasional in swamp forests. Native.
Carya glabra (Miller) Sweet. Occasional in well-drained woods. Native.
Carya ra msis (Wang.) K. Koch. Frequent in fields, pastures, and at old homesites where
plan
Carya paiee (Ashe) Engler & Graebner. Frequent in well-drained eu woods. Native.
Carya tomentosa Nutt. Frequent in well-drained mixed woods. Native

JUNCACEAE
Juncus acuminatus Michx. Infrequent in open wet ditches ane wet depressions. Native
Juncus biflorus Ell. Frequent in open moist places. Nativ
Juncus brac Nenere Engelm, Common locally in open moist ground. Native.
Juncus coriaceus Mackenzie. Common in ditches, wet depressions, and about ponds. Native
Juncus debilis pane Infrequent and local in open wet ground. Native.

ci r

Juncus dichotor hoes Common in open dryish or moist places. Native.

Juncus (issn | uckley. Frequent in usually open wet places about streams, ponds,
and in ditche ee 2.

Juncus effusus L. Common in wet depressions, ditches, and around ponds. Native.

Juncus elliottii —. Frequent in open wet areas. Native.

Juncus gymmnocarpus Coville. Infrequent and local on the borders of lakes. Native.

Juncus marginatus Rostk. Frequent in open moist places. Native.

Juncus polycephalus Michx. Frequent in wet soil in ditches, depressions, and in
water in ponds. Native

2)

allow

Juncus repens Michx. Cons at edges of ponds and in wet low places, frequently sub-
merged in ponds. Native.

Juncus scirpoides Lam. Frequent in moist open places. Native.

Juncus tenuis Willd. Frequent in moist or dryish soils in openings, Native.

Juncus trigonocarpus Steudel. Frequent in mud or shallow water, ponds and ditches. Native.

Juncus validus Coville. Infrequent or locally frequent in open wet a Native

Lusula bulbosa (Wood) Rydberg. Infrequent in open upland woods. Native

LABIATAE
Collinsonia canadensis L. Locally frequent in ravines. Native.
Collinsonia serotina Walt. Rare in deciduous woods along streams. Native.
Glechoma hederacea L. Infrequent in open bottom land and woods. ne
Hedeoma hispidum Pursh. Frequent ee roads and in pastures.

Nat
— — (Raf.) Shinners. Common in moist ground along roads ne in marshy places.

es eae? (A. Rich.) Brig. Abundant locally in open woods along Leaf River. In-
troduced.

Lamium am plexicaule e i. uent along roads and in pastures and lawns. Introduced

Lamium oe L. Infrequent in lawns and pastures. Introduced.

Lycopus rubellus Mc tee Frequent in swamps, marshy ground, ditches, and at edges of
ead Native.

copus virginicus L. Infrequent in low moist woods and around swamps. Se

Menards sig fata L. Local in dry sandy soil at borders of mixed woods t

Perilla frutescens (L.) Britton, Locally numerous in open disturbed Seo oa and pastures.
Introduced.

Physostegia pe a (L.) Bentham. age in low areas along roads. —
learis L.

Prunella vulga Infrequent along roads, in clearings, fields, and pastures. Nativ
anes eae ees pies ns T. & G. ees or infrequent in pinelands and ne pine-
hardwoods. Native.

Les mun reed Schrader. Occasional in es son places. Native.
Salvia azurea Lam. asional in open upland woods. tiy
Salvia ee L; cate in lawns and along roads. ean

aie elliptica a oe in ixed woods. Native.
sancti integrifoli Frequent or ae about lakes and ponds, in ditches, and
pen low ground. ets
Seutlln ia lateriflora L. Infrequent or rare in low moist woods. Native.
‘utellaria ovata Hill var. bracteata (Benth.) Blake. Rare in open bottom land woods. Na-

tive.
Stachys floridana Shuttlew. Abundant locally in eet and along open streams. Introduced.
Stacy fenuifolia Willd. Infrequent or rare in tom land ie tiv

Teucrium canadense L. Infrequent in bottom Ba woods. peel

Tric ae ma dichotomum L. Frequent in ce dry places. Nativ

Trichostema sefaceum Houttuyn. Frequent in dry open places. Nee

LAURACEAE
Persea borbonia (L.) Spreng. Frequent in ravines, low woods, and borders of swamps. Native.
aan albidum (Nutt.) Nees. Frequent in open upland woods, slopes of ravines, old
fields, and wood borders. Native.

LEGUMINOSAE

Albizia julibrissin Pena Frequent escape from cultivation to roadsides and other dis-
turbed habitats. Introduce
ge ee fruticosa L. ek along river banks and in open low bottom land woods.
tive.
Amphicarba Olae teata a Fern. Infrequent in open upland mixed woods. Native.
Apios americana Medicus. Infrequent in low moist woods or thi ckets. Native.
is f

Arachis ae wea L. Tidceauent escape from cultivated to w aste areas. Introduced.
Astragalus distortus T. & G. Rare in dry open pinewoods. Na
T. 7

Baptisia lencant ci & Occasional at wood borders and i. open woods. Native.
lassia fasciculata Michx, Bee ent along roads, trails and around field ative.
Cassia nictifans equent along roads, trails, and about fields. Natiy

Cassia obtusifolia L, A common weed of cultivated fields. Native.

Cassia occidentalis L. Frequent in cultivated fields. Introduced.

Centrosema virginianum (L.) Bentham. Frequent in ce mixed woods. Native
Cercis canadensis L. Frequent in mixed woods and ravines. Native.

Clitoria mariana L, Frequent in open well-drained Fees ae woods. Native.

Crotalaria angulata Miller. Frequent in dry open woods and « hs sandy places. Native.

Crotala igittalis oe in open dry woods. Nati
Crotalaria spectabilis Roth, Infrequent along roads and in fields. Introduced.
Desmoc lina fee ae ex Willd.) DC. Frequent along wood borders, roads, and edges

of fields
ne ae a nm ey DC. Frequent in open areas such as fields, borders of woods,
and along trails. Nativ
Desmodium glutinosum (Muhl. ex Willd.) Wood. Infrequent in ravines. ei
Desmodium laevigatum (Nute.) DC. Scattered in open upland aac ne
Desmodium lineatum DC. Infrequent in dry open mixed woods, Nativ
; jun (L. Infrequent in ravines. Native.
Desmodium obtusum (Muhl. ex Willd.) DC. Frequent in open mixed woods and along
woods and _ fields. i

»
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nt

Desmodium paniculatum DC. Frequent in fields and open dryish woods. Nat

pues perplexum Schubert. Infrequent in open mixed woods and at borders be “fields.

Desn ae rotundifolinm DC. Frequent or occasional in open mixed woods. Native.
Desmodium strictum (Pursh) DC. Infrequent in wooded sandhills. Native
Desmodium tortuosum ieee DC. Abundant locally in fallow fields. ieromee

Desmodium viridiflorum (L.) DC. Occasional or frequent in well-drained open mixed woods.
Native.
Erythrina berbacea L. Frequent in upland mixed woods. Nativ
Galactia macreei M. A. Curtis. Infrequent in mixed woods, new clearings, and thickets.
ative,
Galactia regularis (L.) BSP. Occasional in wooded sandhills. Nati
G

alactia volubilis (L.) Britton. Frequent along wood borders, in open woods, and thickets.
Ive

70

Glycine max (L.) Merrill. An occasional escape is cultivation to roadsides. Introduced.

Lathyrus hirsutus 1. Infrequent along roads. Introduced

Lespedeza capitata: Michx. Frequent in open sar occasional in dryish mixed woods
and at edges of fields. Native.

Lespedeza cuneata (Dumont) G. Don. Infrequent or frequent pow oo Introduced.

Lespedeza a (L.) Hornemann. Frequent in dry open woods.

Lespedeza x nuttallii Darl. Infrequent or rare on roadsides nea

alan as ie vm Michx. Infrequent in open woods. Native.
espedeza repen 2% arton, Frequent or infrequent in dry upland woods and along roads
and trails. SCs

Lespedeza striata (Vhunb.) H. & A. Locally abundant in disturbed places. Introduced.

Lespedeza stuever Nutt. Infrequent in open upland woods. Native.

Lespedeza violacea (L.) Persoon. Infrequent in openings of upland woods. Nativ

Lespedeza virginica (L.) Britton. Frequent at the borders of fields, along roads and trails.
Natiy

si INUS "louis Willd. Local in dry open woods. Native.
efalostemum carolinianum (Lam.) Sprague. Frequent in well-drained upland woods, prin-
cipally in sandhills. Native.

Phascolus polystachios (L.) BSP. Infrequent in i Native

Pueraria lobata (Willd.) Ohwi. Abundant locally alae eae eens

Rhynchosia pitcheria Burkhart. Infrequent. in deep, well- drained sand. Native

N:

Rhynchosia reniformis DC. Frequent in dry open _ ati
Rhynchosia tomentosa (L.) H. & A, Infrequent in open oe aauile Native.
Se leis microphylla (Solander ex Smith) MacBride. Frequent in dry open mixed woods.
ative
Sesbania ee ed wv.) Benth. Rare on river banks. Introduced.
Strophostyles helve L.) Ell. Occasional in clearings and dry open woods. Native.
fs

Strophostyles sa (Muhl. ex Willd.) Britt. Infrequent in upland mixed woods and
clearings. Native.

Sfylosanthes biflora (L.) BSP. Frequent in disturbed woodland. Native.

Pepa florida (Dietrich) C. E. Wood. Frequent in open dryish pine and mixed woods.

ative
Pa ee Spicata’ (Walter) ‘T. & G. Infrequent in open woods. Nati
Tephrosia virginiana (L.) Pers. Frequent in open pine and mixed ecdlenil Native.
Trifolium arvense 1. Abundant locally in open disturbed areas. Introduced

olium campestre Schrel Frequent along roads and in lawns and pastures. Introduced
Trifolium dubium Sibthorp. Frequent in disturbed places. Introduced
Prifolium incarnatum L. Infrequent on roadsides. Introduced
Trifolium repens oc in ae an astures. Introduced.
Vrifolium vesiculosu . Infrequent ire along roads. Intre :
Vicia angustifolia Reicha. ea. elle along roads and ; ie: of nelds, i TmmeBouucete

Vicia caroliniana Wa Infrequent at borders of upland ae woods. Nativ
Vicia dasycarpa Tenore. Occasional along roads. Introduced

Vicia tetrasperma (L.) Moench. Local along roads. Introduced.

Wisteria frutescens (L.) Poi edges of swamps. Native.
Wisteria sinensis (Sims) Sweet. Rare along roads. Introduced.

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ps

LEMNACEARF
Lemna stad Phillipi. Frequent or infrequent in swamps and ponds. Native
Spirodela o eis a (Kurtz

~~

Hegelm. Frequent or iaeegnes in swamps and ponds. Native.

Spirodela ah (L.) Schleid. Frequent or infrequent in lakes and ie Native.
Wolftella eh (J. D. Smith) ‘Thompson. Locally numerous in swamps. Native.

LEN TIBULARIACEAE
Utricularia biflora Lam. Locally abundant or oceasional in shallow water or marshy places.
Native.
LILIACEAE
Aletris aurea Walter. Occasional in moist pinclands. Native.
Aletris farinosa L. Occasional in pinelands. Native

Allium canadense L. Frequent along roads. Native

Allium canadense var. mobilense (Regel) Ow abe Common along roads. Native
Chamaclirium luteum (L. ray. Occasional in ravines. Native.

Hemerocallis fulva L. Infrequent at old homesites. Introduced.

Lilinm michauxii Poir, Occasional on upper slopes of ravines. Native

Medcola virginiana L. Frequent in ravines, Native.

Melanthinm virginicum L, age ae or fe equent in wet grass-sedge communities. Native.
ca bivalve (L.) Britt. Common on roadsides, in lawns, and at old homesites.

Native.
ee bona-nox L. Occasional in thickets, upland woods, and on floodplains. Nat
Smilax ecirrhata (Engelm.) Watso son var. Duger? (Small) Ahles. Occasional in ravines eee
Smilax glauca Walter. Frequent variety of habita Nacive

Smilax hispida Muhl. Infrequent in bottom land woods. Native.
Smilax laurifolia L. Frequent in alluvial woods, around swamps and lakes, and various other
habitats. Native.

Smilax pulverulenta Michx. Infrequent in ravines. Native

Smilax pumila Walter. Frequent in dry open mixed and cao woodland. Native
ee a. . Frequent in moist thickets, deciduous woods, along fence rows, and
wood borders. Native.
Smilax smallii ee Frequent or infrequent in mixed moist woods and along streams.
Jative.

Smilax walteri Pursh. Occasional in bays and marshy places.
Trillium cuneatum Raf. Infrequent to locally abundant in ravines and low rich woods.

~

Native.

Uvularia perfoliata L. Frequent in ravines. 2G

Uvularia sessilifolia L. Frequent in ray . Nati

Yucca smalliana Fernald. Infrequent in a a ae soil in dryish open places. Native

LINACEAE

Linum medium (Planchon) Britt. Frequent in open places about fields and along roads and
trails. Native.

Linum. striatum Walter. Infrequent in low open places, ditches, and borders of boggy areas.
Native

LOGANIACEAE

Gelseminm sempervirens (L.) Aiton f. Frequent in open areas in woods. Native.
oe rankinii Small. Scattered and not frequent in open woods and along trails.
Native
spi mienaa die a L. Infrequent or locally numerous on slopes of ravines and at edge of
mixed woods. Native
LYCOPODIACEAE
Lycopodium alopecuroides L. Occasional in wet ditches. Native.
LYTHRACEAE

ane. Infrequent around lakes and in wet depressions, Native.

Rotala ramosior (L.) Koe

MAGNOLIACEAE

Liriodendron tulipifera L. Frequent in ravines and alluvial woods. Native.
Magnolia acuminata L, Frequent or infrequent in ravines. Native.
gnolia grandiflora L. Common in ravines, occasional in low rich woods. Native.
Magnolia macrophylla Michx. Common in ravines. Native.
Magnolia pyramidata Bartram ex Pursh. Frequent in ravines. Native.
Magnolia virginiana L. Frequent in wet low areas of ravines, bays, occasional along streams.
Native.
MALVACEAE
Callirhoe bepaver (Cav.) A. Gray. Frequent but ie along aie Native.
Hibiscus aculeatus Walter. Frequent along roads and at edges of low pinewoods. Native.
Modiola a (L.) G. Don. Infrequent or rare in open ee near dwellings and

along roads. Native.
Sida rhombifolia I .. Rather common along roads and at edges of fields. Introduced.
Sida spinosa L. Frequent along roads and around fields. Native
MAYACACEAE

Mayaca aubletii Michx. Abundant locally in shallow water and mud. Native.

MELASTOMATACEAE
Rhexia alifanus Walter. Frequent. in open pine-flatwoods and openings in upland mixed
woods. Nativ
lies mariana “L. Locally numerous ditches, open pine-flatwoods, and borders of ponds.
lative,
he via nashii Small. Infrequent in wet ditches and edges of marshy areas. Native.
Rhexia virginica L. Local in ditches, moist pinewoods, and borders of ponds. Native.

MELIACEAE
Melia azedarach L, Naturalized in openings, wood borders and_ roadsides. Introduced.
MENISPERMACEAE

Calycocarpum lyoni (1 — Gray. In ifrequent or rare bottom land woods. Native
Coceulus carolinus (L.) DC. Frequent in disturbed woods and along roads and trails. Cities:

MORACEAE

Morus rubra L. Frequent in ravines, borders of swamps, and low woods. Native.

MYRICACEAE
Myrica cerifer Common in flatwoods, savannahs, and edges of a and streams. Native.
Myrica a Raf. Local in pine-flatwoods and bays. Natiy

NYMPHAEACEAE
aelin. Comn in) some sae and lakes. Native.
Cabomba carolina Gry. Abundant cally in) swamps. Native.
. oui ei (L.) Sibthorp & Sm oe uae mac ee (Small) E. O. Beal. Common
loca In open swamps and lakes. Nati

Brasenia schreb

NYSSACEAEF
Nyssa aquatica L. Abundant in swamps. Nativ
Nyssa biflora Walter. Common in bays, various swamps and borders of streams. Native
Nyssa sylvatica Marshall. Common in moist or dry woods. Native

OLEACEAE
Chionanthus panne us L. Occasional on the upper slopes of ravines and in upland mixed
woods. Native

n low wet woods, borders of streams, and

Forestiera ac shined ies Poir. Occasional

edges of swamps. Nativ
Fraxinus americana L, Oecadonal In ravines and other mesic woods. Native.
Fraxinus caroliniana Miller. Infrequent or rare in’ sy I ative
Fravinus pennsylvanica Mars Jecasional in low mesic woods. Native
Ligustrum japonicum Vhunberg frequent in open ravines. Introd |
Ligustrum sinense Lour. Frequent in mee rows and along wood borders. MES OE USE:
Osmanthus americana (L. xray. Frequent in moist or dry mixed woods. Native

ONAGRACEAE
Gaura biennis L. Occasional along roads and trails. Native.
Gaura filipes Spach, Occasional in open saith, woods. Native.
era aration L. Frequent in wet ditches, low moist woods, and borders o

f ponds.
tive.
ae ge sii Walter. Frequent in open moist woods, borders of ponds, and i n marshy
places. Native.

Ludwigia Pana Walter. Occasional in low woods and marshy places. Native.

73

ts lebincan ne (Nutt.) Hara. Occasional around ponds, marshy areas, and in wet
ditches. Nativ
udwigia “Winans Ww alter. Frequent in low moist open areas and around ponds. Native.
i ;
Native.
Ocnothera bia unis L. Infrequent or locally numerous along roads and in old fields. Native
Ocnothera fruticosa L. Common in cultivated and fallow fields and along roads. Na

OPHIOGLOSSACEAE
oe biternatum (Sav.) Underwood. Oceasional in ravines and
lative

ia. ght m virginianum (L.) Sw
Ophiaglosinm crotalophoroides Walter

bottom land forests.

Occasional in low woods and ravines. Natiy
Infrequent or rare in sandy loam soil along ous
ORCHIDACEAE
Calopogon. pallidus Chapman. Infrequent in wet ditches and moist grass-sedge communities.
ative.
Se sca odontorhiza (Willd.) Nutt. Infrequent or rare in ravines. Native.
Corallorhiza wisteri jana Conrad. Infrequent or rare in ravines. Nate:
j (L.) R. Br. Infrequent in open pine-flatwoods. Native.
ponds and open swamps. Native
Listera australis Lindley. Infrequent in ravines. Native.
Malavis unifolia Michx. Rare in ravines. Native.
Ponthieva racemosa (Walter) Mohr. Infrequent or rare in ravin Native.
j oy L.) Richard. Occasional at borders of swamps and lakes, Native.
in moist woods and along “athe Native.
< Gray. Occasional in moist open places. Nativ

Tipularia discolor (Pursh) Nutt. Rare in ravines. Native
OROBANCHACEAE

Epifagus virginiana (L.) Barton, Infrequent or frequent in ravines. Native.

OSMUNDACEAE

Osmunda cinnamomed - Frequent in bays, about swamps, and wet areas of ravines. Native.
Osmunda regalis Le v ae (Willd.) Gray. Locally numerous in open wet arcas of
ive.

ravines, a. noist wood around swamps. Nat

a
=

OXALIDACEAE
Ovalis dillenii Jacquin. Frequent in old fields, gardens, and

Onalis priceae 7 Infrequent or rare at edge of mixed woods. Native.
An escape from cultivation and established locally in disturbed places.

along roads and trails. Native.

Ovalis stricta L. Frequent along roads and in lawns and fields. Na
Oxalis violacea L. Infrequent in disturbed mesic woods and on road oe Native.

PALMAE
oe hystrix (Fraser) H. W ae Rare along streams in ra Native.
Sabal minor (Jacq.) Pers. Occasional in ravines and bottom land heeds Native.

PAPAVERACEAE

Sanguinaria canadensis L, Occasional in beech-magnolia forest in ravines. Native.

PASSIFLORACEAE
Passiflora incarnata L. Frequent along roads, trails and fence rows and in old fields. Native.
od ive

Passiflora lutea L. Infrequent in moist woods. Nativ

PHRYMACEAE

Phryma leptostachya L. Frequent in beech-magnolia forest in ravines. Native.

PHYTOLACCACEAE
Phytolacca americana L., Common on river floodplain, occasional or frequent in a variety of
disturbed places. Native
PINACEAE
Pinus ee Miller. Frequent or common in well-drained sandy loam soils in upland areas.

Pinus Sie Walter. Frequent in moist woods on river terraces and in ravines. Native.
Pinus palsies Miller. Common in savannahs, flatwoods; occasional in well-drained woods.
la

PLANTAGINACEAE
Plantago aristata Michx. Peden along roads and trails and other disturbed places. Native.
Plantago bybrida Bart. Local in moist open ground. Native
Plantago lanceolata L. Frequent in various disturbed places eueki as fields, open woods, and
roads ils Ive
Plantago rugelii Dene. Infrequent in open low woods. Native.
Plantago virginica L. Common along roads and trails, Native.
POLEMONIACEAE
Phlox pilosa L. Frequent in ings, Open mixed woods, and along reads. Native.

Phlox pilosa subsp. oe ee Wherry. Occasional in clearings. Native.

POLY GALACEAE

Polygala cruci L. Infrequent or rare in wet open flatwoods. Natiy
Se piso ee Walter. Occasional in upland mixed woods. nave.
Po incarnata L. Occasional in open pinewoods. Native.

Sea mariana Miller. Infrequent in open wet places. Native.
Polygala nana (Michx.) DC. Frequent in) moist sandy woods, Native.

POLYGONACEAE

— ovata (Walter) Shinners. Locally abundant along rivers, creeks, and about
swam ative.

tae. _basdropiperoites (Michx.) Small. Frequent about ponds, lakes, swamps, and in
ditches ative.

Persicaria sen aldiiea (L.) Small. Infrequent in moist open ground. Native.

Persicaria punctata (FIL) Small. Abundant locally or scattered in open wet places or shal-
ow water. Native.

Poly ge convolvulus Ly Infrequent in open dryish woods. Introduced.

Rumex crispus L. Common in disturbed areas. Introduced.

nee Dastatulas 3 Baldwin ex Ell. Frequent along roads, trails, and along fields and wood

ati

Rumen sie r L. Infrequent in open moist woods and along roads. a
Tovara virginiana (L.) Raf. Frequent in low woods and moist ravines. Nati

POLYPODIACEAE

Asplenium oe uron (L.) Oakes. Frequent on upper ope of ravines, in dryish woods,
and along roads, fence rows, and wood borders. Native

Athyrium aie nioides’ (Michx.) A. A. Eaton, an in ravines and low wet places.
Native

Pnrnsaria arcolata (L.) Presl. Frequent in various moist or wet places. Nat

One le a sensibilis L. Frequent in moist woods, borders of ponds and swamps, ane in ditches.

oipscde poly podioides (L.) Watt. Localized on trunks and branches of deciduous trees.

N
Native.
Polysticbuni acrostichoides Satake Schott. Frequent in ravines. Nati
Preridium aquilinum (L.) Kuhn. Common. in’ well-drained eel eae ead: trails, and
wood borders ative.

Thely pteris deniate (Forsk.) E. St. John. Infrequent in ravines. Native.
Thelypteris hexagonoptera ees Weatherby. Common in ravines. Native.

1B)

Thely pteris normalis (C. Chr.) Moxley. Common along streams in ravines and in ditches.
ative
eve ie Schott. Common in wet ditches and other wet or seasonally inundated
tive.
Thely pee ee (Gaudich. ) Alston. ai a or rare along — streams. Native.
Woodwardia virginica (L.) Smith. Frequent in low moist places. Native.
PORTULACACEAE
Portulaca oleracea L. Infrequent in open sandy soil. Introduced.
ee

Potamogeton berchtoldii Fieber. Infrequent in la ative.
Potamogcton diversifolius Raf. Infrequent in lakes swamps. Native

kes,

PRIMULACEAE

Centunculus minimus L. Very local in upland wet depressions. Native

RANUNCULACEAE

Clematis virginiana L. Infrequent in open low woods and along streams. Native.
ene Ser ee Walter, Infrequent in open dryish places. Native.
— = abortivus L. Frequent in lawns, along roads and trails, and various other places.
ativ

a ee fascicularis Muhl. ley in open sandy loam soils. Native.
Ranunculus parviflorus L. Infrequent in disturbed places. Introduced.

Ranunculus pusillus Poir. Frequent in seasonally inundated pools. Native
Thalictrun polygamum Muhl. Infrequent or rare in low rich onde. along Leaf River.
ative.

Xanthorhiza simplicissima Marshall. Rare in sandy soil along Weldy Creek. Native.

RHAMNACEAE

Berchemia scandens (Hill) K. Koch. Frequent in mesic veods; Native
Ceanothus americanus L. Frequent in upland woods. Native
Rhamnus caroliniana Walter. Occasional in ravines and ee hardwoods, Native.

ROSACEAE

Agrimonia microcarpa Wallroth. Infrequent in upland mixed woods. Native.
pubescens Wallroth. ange in upland mixed ies Native.
ichx. f.) Fern. Frequent in Es. vines. Nati
Crataegus alma Beadle. Frequent in upland woods. Native
Cratacgus marsballii Eggl. Frequent in deciduous and ee woods. Native.
— opaca Hook. & Arn. Infrequent in wet open depressions. Native.
D snea indica (Andrz.) Focke. Infrequent or common in areas near dwellings and open
places. Introduced
Fragaria virginiana Duchn. Infrequent or rare in open low woods. Native
Prunus angustifolia Marshall. Common in open dry woods. Native
Prunus Nici tele Ait. is boc in low woods and ravines. Native.

wna §. Wats. Rare in dry open woods. Native.

a
<
eS

nee be ersica (L.) Barsch, Rare escape to roadsides. Introduced.

serotina Ehrhart. Frequent in various eb ay habitats. Native

Prunus umbecllata Ell. he aoe or infrequent in -drained open mixed woods. Native.
Pyrus angustifolia Ait. Infrequ - mixed set ne at wood borders. Native

yrs aul (L.) L. f. focally numerous in bays and marshy olace Native.

Rosa punehaate Wendland. Locally abundant along Sra Introduced.

Rosa carolina L. Infrequent in open upland woods, Nat

eee wrgntn Link. Frequent at borders of low woods, oe of fields, and in open woods.

o.
a

eee benaifoliie Small. Frequent in moist woods. Nati
Rubus flagellaris Willd. Infrequent or rare in upland eee oods. Native.

76

Rubus trivialis Michx. Common along railroads, trails, and in a variety of disturbed habi-
tats. Native

RUBIACEAE
Cephalanthus occidentalis L. Frequent about lakes and swamps, and in shallow water. Native.
Diodia feres Walter. Frequent in dry, open areas. Native
Diodia virginiana L. Frequent around ponds and other wet — Native.
Galinm aparine L. Common in open disturbed places. Nat
Galinm circaezans Michx. ee in ravines. Native.
Galinm pilosum Ait. Occasional in ixed Seeds A ravines. Native.

<=
~
SS
=
>
ae
SH
Y
A
ta
coy
a
i
= 5
S
2
Neos
5
°
=
<=
=
=
5
°
a
eg
2
5
fe)
»
a

ae ees ae ee Lam. Infrequent In open moist ground. Introduced.
Hedyotis uniflora (L.) Lam. Infrequent in moist sandy places. Native.
Houstonia nos umbens (J.P. Gmelin) Standley. Occasional in sandy loam in pine and mixed

woods. Native.
Houstonta purpurea L. Frequent in mesic woods. Native.
Houstonia ied schesot Frequent in a sr loam soils. Native.
Mitchella repens L. Common in ravin

non it ati

Richardia ra L. Common in cultiv ee i ‘fall ow fields. Introduced.
RUTACEAE

Citrus trifoliata L. Rare along the Leaf River. Introduced.

Ptelea trifoliata L. Infrequent along the Leaf River. Native.

SALICACEAE
Populus deltoides Marshall, Frequent along Leaf River, rarely in wet soil at sand and gravel
vits. Native
Salix nigra Maxshall; Common along Leaf River and about lakes and swamps. Native.
SAPOTACEAE
Bumelia lycioides (L.) Pers. Rare long Leaf River. Native.

SAURURACEAE

Qu.

Saururus cernuus L. Common around lakes and swamps. Native.
SAXIFRAGACEAE

Decumaria barbara L. Frequent in ravines. Native

Hydrangea ai a Bartr. Frequent in ravines. Native.
Itea virginica L. Frequent in shallow water or wet soil about swamps, ponds, and_ bays.
Native

SCHISANDRACEAE
Schisandra glabra’ (Brickell) Rehder. Rare in ravines. Native.

SCHIZAEACEAE

ad japonicum (Thunb.) Swartz. Occasional in disturbed woods in low areas. In-
uced.

SCROPHULARIACEAE

Agalinis setacea (J.F. Gmelin) Raf. cae in open dry woods. Native

C:

Agalinis tennifolia (Vahl) Raf. Occasional in old fields and clearings. Native.

Aureolaria dispersa (Small) Pennell. Gecasonal along roads and in open upland woods.
Native.

Aureolaria dae (Nutt.) Pennell. Infrequent in dry open weads. Native.

Aurcolaria virginica (L.) Pennell. Infreque nt in mixed woods. Native

Buchnera americana L. Frequent. in — flatwoods. Native.

Buchnera floridana Gandoger. Occasional in pine- Hagen MEN

Gratiola pilosa Michx. Infrequent pine-flatwoods. Nativ

Gratiola ss Nl L. Infrequent in wet open places. Native

Linaria canade L.) Dumont. Common in a variety of open me cae Soe Native.
Lindernia nasal a@ (Michx.) Pennell. Local in open wet areas. Nat
Lindernia dubia (L.) Pennell. Infrequent along streams. Native

Mecardonia acuminata (Walter) Small. Occasional in moist open places. Nat

Micranthe ions umbrosum (Walter) Blake. Common in wet soil or shallow water. hadee

Mimulus alatus Ait. Infrequent around swamps and lakes. Native.

Pedicularis oe L. Local in moist open sandy loam soil. Native.

Penstemon australis Small. Frequent in various open mixed and pinewoods and _ clearings.
ative.

Scoparia dulcis L. Infrequent in open sandy loam soils. Nat

Seymeria cassioides (Walter) Blake. Occasional in open vpland re aede Native.

Native.

Seymeria pectinata Pursh. Occasional in upla nixed w
erbascum thap Rare at garbage es Introduced
eronica arvensis L. Occasional in and lawns. Native

Veronica peregrina L. Infrequent in moist open areas. Native.

SELAGINELLACEAE

Selaginella apoda (L.) Spring. Infrequent in sand along streams. Native.

SOLANACEAE

Datura stramonium L. Rare at garbage dumps. Introduced.
di

Se aed esculentum Miller. Occasional in disturbed areas. Introduced.

Petuni« atkinsiana D. Don ex London gael escape from cultivation. Introduced.

Phoysalis pee L. Frequent in cultivated ground. Native

bea pubescens L. Infrequent in cultivated i Native.

Physalis virginiana Miller. Occasional in disturbed places. Native.

Solanum americanum Miller. Occasional in open deciduous ne and various waste places.
Native.

Solanum a - Occasional in a variety of disturbed nae: Native.

Solanum nigrun nfrequent in fields and at wood borders. Native
Solanum pase nie Lam. Rare in old fields. Introduced

SPARGANIACEAE

Sparganium americanum Nutt. Infrequent or frequent in shallow water of ponds and lakes.
Native

STERCULIACEAE

Melochia corchorifolia L. Common in cultivated fields. Introduced.
STYRACACEAE
Halesia diptera Ellis. Frequent in ravines. Native.
Styrax americana Lam. Occasional in wet woods ane around lakes and swamps. Native
Styrax grandifolia Ait. Frequent in ravines. Nativ
SYMPLOCACEAR
Symplocos tinctoria (L.) L’Her. Frequent in mixed woods. Native.
TAXODIACEAE
Taxodium distichum (L.) Richard. Common in cypress-tupelo swamps. Native.
THEACEAEF
Stewartia malacodendron L. Infrequent in ravines. Native.
TILIACEAE
Tilia caroliniana Miller. Frequent in ravines. Native.
ULMACEAE

Celtis laevigata Willd. Occasional in alluvial woods. Native.
Celtis tenuifolia Nutt. var. georgiana (Small) Fern. & Schubert. Occasional in open upland

nixed woods. Native.
anerad aquatica eee J. F. Gmelin. Occasional in shallow water or about lakes and
npy woo Nativ
Ulnins alata Michx. en in mixed woods. Native.
Ulmus americana L. Occasional in low woods. Native.
Ulmus rubra Muhl. Infrequent or rare in ravines. Native.

—=

ey,

UMBELLIFERAE

ree leptophyllum ae F. Mueller. Occasional in moist open areas. Native.

a astatica (L.) ban. Occasional in moist sandy loam or shallow water. Native.
aii tanta Hooker. Frequent along wee and about dwellings. Nacive,
Daucus carota L. Infrequent along roads. Introduced

Eryngium sana Nutt. Infrequent in open wet places. Native.

a Sis yuccifolium Michx. (Incl. E. synchaetum (Gray) Rose). Frequent in open pine-
oods, Native.

eee age wmbellata L. Occasional in shallow water of ponds. Native.

Hydrocotyle verticillata Thunb. Infrequent in shallow water of lakes. Native.

Oxypolis fliformis (Walter) Britt. Infrequent in open wet areas. Native

Oxypolis rigidior (L.) Raf. Infrequent in bays and pe places. Native.

Ptilimnium capillaceum (Michx.) Raf. Occasional in ditches and other open wet soils.
Native.

Sanicula canadensis L. Infrequent in mixed woods. Native.

Sanicula smallii Bicknell. eum or infrequent in ravines and low woods. Native
Thaspium barbinode (Michx.) Nutt. Infrequent in ravines and low woods. Native.
Thaspium trifoliatum (L.) Gray var. flavum Blake. Infrequent in mixed woods and ravines.
ative.
Trepocarpus acthusae Nutt. Local along Leaf River. Native.
URTICACEAE

oe iis (L.) Swartz. Frequent in swamps, ditches, and around lakes and
onds. Nat

tahoe a pei ee nsis (L.) Weddell. Occasional and local in ravines and me woods. Native.
Pilea pumila (L.) . ray. Occasional in ravines and low woods. Native

VALERIANACEAE

Valerianella radiata (L.) Dufr. Common along roads. Native
VERBENACEAE
Lantana camara L. Infrequent escape from cultivation or persisting around old homesites.
tro
Callic ar pa americana L. Common the upper slopes of ravines and in upland woods. Native

eri bonariensis L. Infrequent or rare along roads. Introduced.
Verbena brasiliensis Vellozo. Common in a variety of sean es Introduced.

Verbena carnea Medicus. Infrequent in upland mixed a ‘Native.
Verbena halei Small. Frequent along and trails. Native.

Verbena rigida Sprengel. Local in dry open places. Introduced.
Verbena tenuisecta Briquet. Common son roads and in other open sandy sites. Introduced.
p y
Verbena urticifolia L. take requent or e in disturbed low woods. Native
VIOLACEAE
Viola esculenta Ell. Infrequent in moist pinewoods. Native.
Viola floridana Brainerd. Rather common in ravines and low woods. Native.
Viola palmata Occasional in ravines. Native.

ies Wav eaders Pursh. Infrequent in open disturbed areas. Native.
Yiola pedata V.. Occasional in upland mixed woods. Native.

ea brimulifolia L. Frequent in moist sandy soils or muck about ponds, swamps, and in
wet ditches. Native.

Viola rafinesquii Greene. Frequent along roads and in fields. Native.

Viola friloba Schweinitz. Occasional in ravines and low woods. Native.

Viola walteri House. Common in rayines and other mesic woods. Native.
VISCACEAE
Phoradendron serotinum (Raf.) M. C. Johnston. Frequent on a yariety of deciduous trees.
ative
VITACEAE
Anmtpelopsis arborea (L.) Koehne. Frequent about swamps, lakes, and various other wet
areas. Native.
Ampelopsis cordata Michx. Infrequent and local along railroad. Native.
Parthenocissus quinquefolia (L.) Planchon. Common in various mesic woodland habitats.
Native.
Vitis acstivalis Michx. var. argentifolia (Munson) Fern. (Incl. V. bicolor LeConte). Fre-
quent along stream banks and in low woods. Native.
Vitis cinerea Engelm. ex Millardet. Occasional along streams and in low woods. Native.
_ aged Michx. Common in a variety of woodland habitats ranging from moist
Vitis sie i "igeeaieat in low woods and along streams. Native.
XYRIDACEAE
Xyris caroliniana Walt. Infrequent in moist pine-flatwoods. Nat
Xyris difformis Chapm. Occasional at the borders of ponds and ie Native.
Xyris fimbriata Ell, Frequent in sand or mud about lakes and in wet ditches. Native.
Xyris iridifolia Chapman. Frequent at the borders of ponds a in wet ditches. Native.
Xyris jupicai Richard. Locally abundant in moist or wet sand in open discanied places.
V ative.

ACKNOWLEDGMENTS

I wish to thank several persons who assisted me during the study. In-
formation on soils was obtained largely in the field with Rex Davis of the
Soil Conservation Service. I am grateful to F. J. Hermann, Robert Kral,
Michael G. Lelong, and Sidney McDaniel for their help in resolving certain
problems of plant identification. I would also like to express my appreciation
to The Society of Sigma Xi for a Grant-in-Aid of Research which contributed
to the support of this study. Special gratitude is due M. M. Roberts of Hat-
tiesburg, Mississippi, for his understanding and generosity in support of this
endeavo

REFERENCES
CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the vascular plants of Texas.
Texas Reareh Foundation, Renner, Texas.
FENNEMAN, N. M. 1938. Physiography of eastern United States. McGraw-Hil
Company, New York, N.
FOSTER, V. M. 1941. Pores County mineral resources. Geology. Bull. Miss. St. Geol.

—

Book

—

1-87.
LOWE, E. N. 1921. Plants of Mississippi. A list of flowering plants and ferns. Bull. Miss.
St. Geol. Sur. 17: 1-29
MCWHORTER, J. C. 1962. Climatic patterns of Mississippi. Miss. St. Univ. Agr. Exp. Sta.
Bull. 650: 1-24.
MISSISSIPPI GEOLOGICAL SOCIETY. 1969. Geologic map of Mississippi. Jackson, Mis-
sissippi.
S. GEOLOGICAL SURVEY. 1949. New Augusta Quadrangle, Mississippi, 15 minute
series aes map). Washington, D. C.
956. State of Mississippi elevation map. Nae Sage D.C.
VANDE One: H. B. 1962. Soils of Mississippi, Miss. St. Univ., Univ. Agr. Exp. Sta.,
Mississippi State, Miss.

CHROMOSOME NUMBERS OF
GYPSOPHILIC PLANT SPECIES OF
THE CHIHUAHUAN DESERT

A. MICHAEL POWELL and SHIRLEY A. POWELL
Department of Biology, Sul Ross State University
and Chihuahuan Desert Research Institute
Alpine, Texas 79830

This reports original chromosome counts for some dicotyledon species
which occur on or are associated with gypsum soils in the Chihuahuan
Desert region of North America. Many taxa are absolutely restricted to
gypsum while others may also grow on surrounding non-gypsum habitats
(Johnston, 1941; Waterfall, 1946). These obligate gypsophiles and facultative
gypsophiles include species that constitute unique gypsum floras of the Chi-
huahuan Desert and elsewhere in western United States and northern Mex-
ica (Powell and Turner, 1975).

The chromosome numbers presented herein were accumulated since 1969
when a chromosomal survey of gypsophilic species was initiated. Major
cbjectives of the chromosomal studies were to compare the percentages of
polyploid species on gypsum and non-gypsum substrates and to evaluate the
polyploid percentage in the entire Chihuahuan Desert flora. Elsewhere we
report (Powell and Sloan, 1977) that the gypsum floras comprise fewer poly-
pleids (17.8%) than non-gypsum vegetation (32.7%), and we calculated an
overall polyploid percentage of 30.2 for the Chihuahuan Desert flora. Table 1
presents a large portion of the counts that have contributed to the percent-
ages. A limited number of other counts for gypsophilic taxa have been re-
ported by Turner (1972a, 1972b, 1972c, 1973a, 1973b), Turner, Powell and Wat-
son (1973), Bacon (1974a, 1974b), and a few other workers. We hope even-
tually to increase the number of counts for gypsophilic and non-gypsophilic
species of the Chihuahuan Desert and therefore to refine the accuracy of
polyploid percentages in the edaphic-floras. Several hundred species of

about 150 genera and 50 families have been estimated to occur on gypsum

outcrops of the Chihuahuan Desert. Chromosome numbers are known for
about 125 species, about 55 genera and about 25 families, of which 78 species

representing 54 genera and 17 families are presented in Table 1,

Information regarding previous reports for the species listed in Table 1
can be found in the chromosome number indexes (Darlington and Wylie,

1956; Cave, 1956-64; Ornduff, 1965-1967; Moore, 1967-1972; Federov, 1969)

and other recent literature. Therefore not all of the taxa are commented

upon in discussion.
Buds for meiotic analyses were fixed in Modified Carnoy’s Solution

SIDA 7(1): 80-90, 1977.

81

(4: 3: 1), and chromosomes were stained with acetocarmine during appli-
cation of standard squash techniques (Turner and Johnston, 1961). Vouchers
are deposited at SRSC, TEX, or US. In Table 1 collection numbers preceded
by P are by Powell, SI are by Sloan, Si are by Sikes, O are by Olsen, Tu are
by Turner, and To are by Tomb.

Table 1 reflects the predisposition of certain families, genera, and species
to gypsum habitats. Our observations of gypsum vegetation encourage us
to draw conclusions about the most prominent families on the unusual sub-
strate, even though this is not necessarily revealed through the taxa for
which we present chromosomal data. In terms of number of genera and
species, the families most prominently represented on gypsum habitats in-
clude Asteraceae, by far the largest, and Cruciferae, Hydrophyllaceae,
Boraginaceae, Gramineae, Loasaceae, and Nyctaginaceae. The Nyctage
family is not represented in Table 1 because we have not been able to ob-
tain unequivocal chromosome counts for any of its gypsophilic members.
Families perhaps less prominent but characteristic on gypsum exposures
are Euphorbiaceae, Chenopodiaceae, Leguminosae, Caryophyllaceae, Zygo-
phyllaceae, Onagraceae, Malvaceae, Frankeniaceae, and Amaranthaccac.
As suggested in Table 1 certain gypsophilic families and genera contain
more polyploid or aneuploid taxa than others. For example, in Asteraceae
we have calculated ca. 10% polyploidy for the gypsophilic species, while
ca. 27% polyploidy was found for the non-gypsophilic species (data in
Table 1, and published elsewhere). Our analysis of the chromosome number
reports for Euphorbiaceae by Urbatsch et al. (1975) revealed ca. 60% poly-
ploidy for both the gypsophilic and the non-gypsophilic species of this family
in the Chihuahuan Desert region. Eventually we hope to draw conclusions
regarding the chromosomal evolution of gypsophilic taxa, but as yet suf-
ficient data are not available to warrant such a discussion.

Sesuvium verrucosum (n = 8) occurs in gypseous and non-gypseous,
saline habitats and thus the taxon should be regarded as a facultative gyp-
sophile, or perhaps more accurately as a halophytic gypsophile, according
to the terminology proposed by Johnston (1941) to denote those gypsophilic
taxa that can tolerate concentrations of salts and alkali.

Chromosome numbers are listed for 50 species of Asteraceae, easily the
phylad with the most gypsophilic taxa. Most of these species (Table 1) are
cither dwarf shrubs (19 spp.) or herbaceous perennials (18 spp.), life forms
which predominate on gypsum exposures. Large shrubs or trees and an-
nuals are less common on gypsum, although annual facultative gypsophiles
are more numerous.

Dicranocarpus parviflorus (n = 10) is one of the few annuals that seem-
ingly has an absolute requirement for gypsum. The species ranges through-
out most of the Chihuahuan Desert, exhibits considerable exomorphic vari-
ability, particularly in fruit characters, but is not known to vary in chromo-
some number (Turner and Johnston, 1961; Table 1). Dyssodia acerosa
(n — 8) is one of the many facultative gypsophiles that does not exhibit

82

chromcesomal (except n = 13 in some non-gypsum habitats) or superficial
morphological variability whether growing on gypseous or non-gypseous sub-
strates. High (1974) has demonstrated that cryptic ecotypic differentiation
does eccur in populations of facultative gypsophiles, and that subtle genetic
differences have evolved in some isolated populations of obligate gypso-
philes, but studies of this sort are generally lacking for gypsophilic vege-
tation (Powell and Turner, 1975; Parsons, 1975).

Of the five species of Flaveria (« = 18) reported in Table 1, only F.
anomala is considered to be an obligate gypsophile. The other species are
commonly found in mixed gypsum-saline habitats.

Turner (1972c) reported tetraploid counts (n = 34) for Gaillardia multi-
ceps var. microcephala, one of the relatively few polyploid gypsophiles. In
Table 1 we also list a hexaploid count (n = 51) for one population of var.
microcephala. Gaillardia sp. is an undescribed entity seemingly restricted
to gypsum in the area of Chihuahua indicated by the localities in Table 1.

The collections of Machaeranthera cf. pinnatifida (n = 4,8), all from
“pure” gypsum or gypseous substrates (Table 1) are superficially similar
to members of this taxon (rn — 4,8) found on non-gypseous soils. As stated
earlier, the occurrence of facultative gypsophiles such as Machacranthera
is possibly correlated with edaphic adaptation, but it has been difficult to
detect.

Pseudoclappia arenaria was reported earlier (Powell and Turner, 1963)
as n — 18+1, and our present count (Table 1) is also not unequivocal. A
positive number is difficult to ascertain because the chromosomes are small,
heteromorphic, univalents or fragments may be present, and one ¢

O

r more
bivalents tend to separate early. Although Pseudoclappia watsonii is listed as
n — 19, the chromosome number of this taxon also needs verification.

The species of Sartwellia, all nm = 18 (Table 1), are exclusively gypsophil-
ous, with the possible exception of S. puberula which occasionally grows in
soils that are weakly or questionably gypseous.

Any distributional or taxonomic significance for the diploid (n = 11) and
tetraploid (n = 22) populations of Thelesperma megapotamicum is not im-
mediately apparent. Mostly tetraploid and a few diploid populations of this
species also occur Gn non-gypseous soils (Powell and Sloan, in preparation).
The gypsophilic and nen-gypsophilic populations are not consistently dis-
tinguishable except that gypsophiles in Texas occasionally more closely re-
semble the Mexican T. ramosius (2 — 11) than other T. megapotamicum.

The chromosome number obtained for Petalonyx crenatus (n = 22) differs
from the n = 23 that was determined for the other four species of the genus
(Davis and Thompson, 1967). Petalonyx crenatus, a Coahuila endemic on
gypsum, is isolated from the other species which are widespread in the west-
ern North America deserts, and so its chromosomal difference is not sur-
prising.

The count listed for Tetraclea coulteri (n = 21) is a first report for the

83

genus which has been placed variously in the Verbenaceac and in the Labi-
ateae. In both families there are species with « = 7, and both families ex-
hibit taxa with an array of base numbers which could give rise to n = 2
The chromosome number of Tetraclea may be of value in placing the genus
if related genera were studied cytologically.

A chromosome number for the monotypic Mexican endemic Sericodes
greggit (n = ca. 15) is reported in Table 1. The approximate count was ob-
tained from but three cells which revealed small, light-staining chromo-
somes, but perhaps it will contribute toward understanding the origin of the
related genus Larrea (x — 13), a North American-South American disjunct
(Porter, 1974).

_—

ACKNOWLEDGEMENTS
We are grateful to Prof. B. lL. Turner for field companionship and assist-
ance, and for stimulating discussions concerning plant gypsophily. We also
thank A. Spencer Tomb, Sam Sikes, and Steve High for field assistance, bud
collections, and for contributing counts, John Averett for information con-
cerning Chamaesaracha, and Henry J. Thompson for identifications of Ment-
zelia species. The study was supported in part by NSF Grant GB-37674.

1. ORIGINAL CHROMOSOME COUNTS FOR OBLIGATE AND FACULTATIVE
PLANT GYPSOPHILES OF THE CHIHUAHUAN DESERT

Table

Taxon ‘Game sate c No. Locality
AIZOACEAE
*Sesuvinm verrucosim Raf. § TEX: Reeves Co. Toyah Lake 4 mi SE Pecos,
P 1911; 1 mi N of Pecos, S/ 79.
8 MEX: Coah. 12 mi E of Cuatro Cienegas, P &
Tu 2279

ASTERACEAE

Aster snbulatus Michx. § TEX: Culberson Co. 20 mi W of Orla, S/ 88.
*Baccharis cf. wrightii Gray 9* TEX: Brewster Co. near Terlingua Tw s.n

bu csothioi Benth
. dealbata Gra

2 TEX: Culberson Co. 26 mi W of — SI 100;
Reeves Co., 6 mi W of Orla, S/
mingly obligate Sempnr

halbphyee sypsop phi

* — previously einen chromosome number

Aame a onl oe or 2 = 8 HW + 1 UL

— T+ i. chromosomes.

stay ba ai. — — lus 2 fragments

5 re

ith occasional multivalents
| carly pes heteromorphic bivalents, 1-2 fragments, one multivalent, could be
n= 17 II LJ,2 = 18 H, or vw = 19 IL.
® possibly » = 18 IT + 117.
N consis tently # == 18 Il + 2 rings of four

aaa ulate and univalents
T, Sf 24

ae ‘apomictic, bivalents rarely observed.

84

{Bartlettia scaposa Gray

*Brickellia sf
Erigerou pinkavii ‘Turner

tDicranocarpus parviflorus Gray

Dyssodia acerosa DC.

tEricamera triantha
(Blake) Shinners

Eupatorium cf. gregeii
Gray

Flaveria anomala
3

L. Robins

chloraefolia

Flaveria
Gray

Flaveria op positifolia
(DC.) Rydb.

Flaveria palmeri
J. R. Johnse.

Flaveria trinervia

(Spreng.) C. Mohr

leaasli Witenes

Greene
microcephala Turner

11

\o

Xo

io =]

“so

m
oo

—
wo

a
o

ro
oo

TEX: Eludspeth Co. just E of Tommy’s Town,
P 2418.

MEX: Chih. 5 mi W of Presa Granero, P & Tu
2027.

MEX: Coah. 18 mi E of Cuatro Cienegas, P &
Tu 2270

MEX: . L. 7.5 mi S of San Roberto junct.,
P& To 2560; 8S. L. P., 28 mi § of Matehuala,

P & To 2581; Zac., 8.5 mi NE of Condigeiin
del Oro, P & To 2598,

TEX: Culberson Co. 26 mi W of Orla, S/ fee

MEX: Chih. 8.1 mi W of Camargo, p To
2650

TE Brewster Co. near Terlingua, P 2390;

EX
Ward Co., 5 mi S of Pyote, S/ 69.
MEX: 4 mi
2281,

NW of Cuatro Cienegas, P & Tx
5 mit W of Cuatro Cienegas, P & Tn 2293,
mi

MEX: Coah. 8 S of Cuatro Cienegas, Si, O,
&

P 849

oo Coah. ta mi N_ of Concepcion del Oro,
To 259 N. L. 7.5 mi S$ of San Roberto

junect., P& 7 2559; 34.4 mi S of San Roberto

junct., P & To 2566; 1 i N of te-

huala, P & To 2568: Si, O & P 818; 28 mi §

of Matehuala, P & To 2579; a 3.5 mi NE of

Concepcion del Oro, P & To 2

MEX: Coah. 12 mi E of Cuatro Cienegas, P &

Tu 2278,

TEX: 20 mi W of Orla, S/ 73;

met

MEX: Coah. 39 mi N of. San

Culberson Co.
ca.

Reeves Co. Y% mi E of Toyahvale, P 2799.

border, S/, O

fee 4
3]; f Saltillo, P &

9 mt N of Dgo- Zac.
N. L.

mi S of San

oO
; Zac. SY mi NE of junct. 49-54, P &

S: eaters P & To
2611; juste W at Cuatro Cienegas, P & To 2621;

67 mi SW of Cuatro Cienegas, P & To 2631;
near Matamoros, S/, O, & P 838: 15 mi E of
Cuatro Cienegas, $4, O, & P 851

MEX: a 3 mi W. of C amargo, P & To
2658; 2-3 mi N of Meoqui, P & To 2660;

Ss. L. PL. ca. “ mi N_ of vom aa Si, O, & P

NE -

P

TEX: Reev
Williamson
Si 812

Co., 1

s mi N_ of cos, P 2204
Cox “ea. 1

mi N - reere hall,

aie Chaves Co. 1-4 mi N of Dexter,

ahs i ard Co,

—

6 mi S of Pyote, P 1876.

Gaillardia pinnatifida

Torr 17
+*Guillardia furneri 17

Averett & Powell

Gutierrezia microcephala

(DC.) Gray 4
Gutierrezia cf. fexana
Dc.) T. & G. 4
rrezia sp. 16
Hari greggii Gray
ar. greg 18

Huaploesthes greggii var. fexana

Coult.) I. M. Johnst.

foo)

Huaploesthes greggi cf. var.
(Coult.) M. Johnst.

fenana
l

XHaploesthes robusta
I. M. Johnst

18

TH ynrenoxys odorata DC. 15
aes es

ry) Gre 6

Isocoma wrightii
(Gray) Rydb. 6

Isocoma wrightii
iray) Rydb

ca.12
Leucelene ericoides
(Torr.) Greene 8
Mac deta es ra cf. pinnatifida
Hook.) Shinners 4
ie - ranthera cf. pinnatifida
ok.) Shinners 8

TEX: Culberson Co. 8-15 mi W of Orla, SI 26.
MEX: Chih. 14 mi W of Presa Granero, &
Tu 2025; 1.7 mi § of Placer de Guadalupe,
P& nu 2055

20 mi W of Orla, S/ 90;

TEX: Culberson Co.
Ward Co. 7 mi § of Pyote, S/

between Monclova and Monterrey,

MEX: N. L.

P & Tu 2295.
TEX: Reeves Co. 7.4 mi E of Pecos, S! 40.
MEX: Coah. ca. mi W of Oballos,

2252: 1 mi N of “leadc P & Tu 2725
Portrero Sk P®& 4 2331; near epi aeea
P& Tu

P& Tu
IN.

NEW eee orro Co. 48.5 mi FE of San An-
tonio, P

TEX — . ear junct. 1053-329, P 2777;
Crosby Co. near "White River Lake Dam, P
2787; Ward Co., just S$ Ais Pyote, P 1899; 6 mi
E of Grandfalls, P 235

EX: Brewster of Te slings
High & High 99; aa x ee just N of Lang

ry, P 2678.

MEX: Coah. 3-4 mi SW of Cuatro Cienegas,

P & To 2619.

se KX: N. L. 45-46 mi SE of Saltillo, P & To
i4

MEX: ca. 15 mi E of Cuatro Cienegas, P & Tu

O27 ls

TEX: Culberson Co. 16 mi of Orla, S/ 357;

29 mi N of Van Horn, P 2788; Reeves Co. 1

mi N of Pecos, S! 58

TEX: Co. ca. 1 mi E of Tommy’s

Hurspeth
Town, P 2791.

EX: oe rson Co. 34 mi N of Van Horn,
: 239
MEX: Coah. 35.5 mi N of Concepcidn del Oro,
P & To 2601; 67 mi SW of Cuatro Cienegas,
P & To 2632; 45 mi SW of Cuatro Cienegas,
Tu 6003, Chih. 11.5 mi W of Camargo, P &
To 2641; 8.1 mi W of Camargo, P & To 2651.
N. L. Garcia Caves, Tu pee
TEX: Culberson Co. 36 mi of Van Horn,
P 1943; Reeves Co. 4 mi W [ Orla, S! 45;
mi SE of Pecos, S/ rie

Ojinaga, P, Tu, &

MEX: Chih. 20 mi W of
Si 2458

86

Machae a ‘ra tanacetifolia
H.B.K.) Nees 4 TEX: Reeves Co. 7.4 mi SE of Pecos, SI 35,
. m ee m lencanthum

10 TEX: Brewster Co. near Terlingua, P 2388; Jeff
Davis Co. Near Brack’s Tunnel, - & ‘ube ock
312; Reeves Co. 6 mi W of Orla,

Perityle parryi Gray 17) MEX: Chih. 6 mi W of a oe & Tu
2030; 2049,

tPerityle vaseyi Coult. 17) MEX: Chih. 6.5 mi § of Ojinaga, P & Tu 2002
TEX: Brewster Co. 1.5 mi Wo Teena

High & Fligh 97.

ee arenaria
Rydk 18° TEX: Reeves Co. 1 mi N of Pecos P 1907; 2142.
* ye se red ha watsonii
rell & ca, 19" “TEX: Hudspeth Co. just E of Tommy’s Town,
P 2415

Psilostrophe cf. tagetina

(Nutt.) Greene 16 ‘TEX: Culberson Co. 25.6 mi Su of Orla, SI 77;
34 mi of Van Horn, P 2394; Reeves Co. 7.4
mi SE of Pecos, S/ 36; Ga Co. 6 mi E of
Grandfalls, P 2368.
tSarfwellia flaveriae
Gray 18 NEW MEX: Chaves Co. 1-4 mi N of Dexter,
P 2796, TEX: Crane Co. 13 mi E of Grandfalls,
Meyer 30 = Culberean Co. 45 mi N of Van Horn,
P 2192s Reeves C5. 8 ani W of Orla, S/ 31; 4
mi W of Orla, S! 46; 5 mi W of Onl. P 1917.
tSartwellia mexicana
ray 18 MEX: Coah. 0.5 mi S of Est. Hermanas, P &
To 2548; §. L. P. 1 mi N of Matehuala, P &
To 2570; 28 mi S$ of Matehuala, P & To 2580;
Zac. 8.5 mi N of oe - Oro, P = To
2593; $5 mi NE unct. 49- P& To 2592,
tSarfwellia puberula
Rydb 18 MEX: Coah. 61 mi W of Saltillo, P & To 2609;
40 mi 7 of Sal Pedro, P & To 2613; 38 mi SW
of Cuatro Cienegas, P a To 2626; 68 mi SW of
Cuatro care P& » 2634; 67 mi N of San
Pedro, Si, O, & P ih Chik, 12 mi W of Ca-
margo, P - To 26353; ca. 27 mi NE of El Mor-
rion, P & To 2666
+ pees ee
Pow & Te 18 MEX: Chih. ca. 5 mi W of Presa Granero, P
2536
+* Senecio warnockii
Shinners 20 ‘TEX: Culberson Co. 20 mi W of Orla, S/ 93,
Stephanomeria panciflora
Torr.) A. Nels 8 MEX: Chih. 6.5 mi § of Ojinaga, P & Tu 2003,
TEX: Presidio Co. NW of Candelaria, High &
Gallagher 79,
Strotheria gy psophila
‘Turner 8 MEX: N. L. 15.7 mi S of San Roberto junct.,
P & To 2565.
T ie sperma longipes
Gri 10 TEX: Brewster Co. near Terlingua, P 2385.
L he le as rma megapotamicum
Ope g.) Kuntz 11) TEX: Culberson Co. 22 mi W of Orla, S/ 96,
Thele ns rmaome ene wan
AS aa Kuntz. ca. TL) TEX: Ward Co. 7.6 mi S$ of Pyote, SI 85

Thelesperma me gapofamicum

(Spreng.) Kuntz.

Thelesperma megapotamicum

(Spreng.) Kuntz.

+Thelesperma raniosius
Blake

+Thelesperma scabridulum
Blake

Varilla mexicana Gray
Verbesina encelioides
Cav.) Gray
+X ylorbiza wrightii
(Gray) Greene
BORAGINACEAE
+*Coldenia bispidissima
Torr yray

CARYOPHYLLACEAE
+*Drymaria cf. lyropetala

+*Drymaria sp.

CHENOPODIACEAE

X= Snaeda suffrutescens
Wats

CRUCIFERAE

Lepidium montanum
Nutt

Nerisyrenia cantporum
(Gray) Greene

Nerisyrenia cam porum
we

(Gray) Green

+Nerisyrenia gypsophila
Bacon

tNerisyrenia a ae
(Wats.) Gree

he Ner risyrenia linearifolia

(Wats.) Greene

on

Xo

mM

bo

—

bo

\o

—

\o

Co

\o

\o

_
oe

87

MEX: Coah. 1 mi N of eae P & Tu 2726,
ete ME X: Chaves Co. 1- N of Dexter,
> 97

MEX: Chih. 15 mi SE of Fl Morrion, P 2432.

TEX: Culberson Co. 34 mi N of Van Horn,

P 2404; Reeves Co. 7.4 mi SE of Pecos, S/ 37;
16

MEX: Coah. 1 mi S of Est. Hermanas, P & Tu
2264.

MEX: Coah. 35.5 mi N of Concepcidn del Oro,
P& To 2600,
MEX: Coah. Las Delicias, P & Tu 2698.

>

TEX: Reeves Co. 7.4 mi > of Pecos SI! 43;
Ward Co., 5 mi S of Pyote, S/ 67

MEX: Chih. 10.4 mi S of Ojinaga, P & Tu 2017

TEX: Culberson Co. 36 mi N of Van Horn,
P 1939; Reeves Co. 5 mi W of ae io 1914;
Ward Co. 6 mi CE of Grandfalls, P 23

MEX: N. L. between Monclova and Monterrey

I ;
MEX: Chih. ca. 7 mi NW = of Presa Granero,
33

TEX: Reeves Co. 4 mi W of Orla, S/ 47.

NEW MEX: cee Co. 48.5 mi E of San An-
tonto, P 253

TEX: ne Co., 20 mi W of Orla, S/ 91;
34 mi N of Van Horn, P 2400; Hudspeth Co.,
\ I Reeves 8

oO Sn Mts., 2409; e o., §
mi W S ; 7.4 mi SE of Pecos, S/
41; W

; Ward Co., 's mi S of Pyote, S/ 62; 6 mi E
of Grandfalls, P 2357

MEX: Chih. 11.5 mi W of Camargo, P & To
2640

NEW ME ofia side San An-
dres Mts., ee : oe 264

MEX: Chih. near Presa Granero P & Tn 2026.

TEX: Culberson Co., 20 mi a of Orla, S/ 74;
34 mi N of Van Hein. P 239

o Reeves Co., 7.4 mi SE of Pecos, §
Ward Co. 5 mi § os Pyote, S/ 63

88

*Streptanthus carinatus
Wright

EUPHORBIACEAE
Cay.
Cav.

Crofton dioicus

Croton diovicus

GENTIANACEAE
ye OF:

ntarium sp.

HY DROPHYLLACEAE

Nama hispidum Gray

*Phacelia @y psogenia

/L. ohnst.

t* Phacelia cf.

Vorr

integrifolia

LINACEAE
“Linum puberulum

(Engelm.) Heller

LOASACEAE

Encnide lobata (Hook.) Gray

+Mentzelia humilis
(Gray) Darl

Mentselia multiflora (Nutt.)
multiflora

Mentre tia
H

saxicola

Thoms. & Zavor.

Menfzelia sp.
+° Petalonyx crenatus
: cS vats

ray

MALVACEAE
t*Sphacralcea subbastata
Coult

ONAGRACEAE
Calylophus hartwegii Benth.
aven )
subsp. filifolius (Eastw.)
Towner & Raven

us

N

21

bo
ie

MEX: Chih. 20 mi W of Ojinaga, P, Tu, & Si
2457

TEX: Ward Co. 6 mi E of Grandfalls P 2372.
TEX: Culberson Co. 26 mi W of Orla, S/ 99.
MEX: Chih. 27 mi NE of El Morrién, P, Tu,
& Si 2487.

MEX: Chih. 20 mi W of QOjinaga, P, Tu, &
Si 2460,

MEX: Chih. ca. 6 mi SE of EL Morrién, P &
Tu 2689,

: Brewster Co., near ‘Terlingua, 2384;
Presidio Co. ca. 10 mi W ‘of il eae High &
Gallagher 90; Reeves Co. mi SE of Pecos,
SI 115
TEX: Ward Co. 7 mi § of Pyote, S/ 120.

MEX: N. L. Puerto Chico, near Monterrey, P
& Tu 2332.

TEX: Culberson Co. 34 mi N of Van Horn,
P 2399; Reeves Co. 6 mi W of Orla, S! +49;
5 mi W of Orla, P 1912.

TEX: Reeves Co. 7.4 mi SE Pecos, S/ 34.
eos Chih. 27 mi Pal of El Merrion, PL Tu,
& Sr 2498; 8.1 mi W of Camargo, P & To 2647.
TE ve Hudspeth o 22 mi § of Tommy’s Town,
P

MEX: _Coah. 2 mi off hwy. toward Las Delicias,
P 269

TEX: ieee Co. 34 N of Van sans
P 2396; Ward Co. 5 mi S. - Pyote, SI 6

TEX: Culberson Co. 34 N of Van Horn,
P 2405; Reeves Co. 6 mi W of Orla, SI 52

SCROPHULARIACEAE

2 MEX: N. L. between Monclova and Monterrey,
P & Tu 2306; Chih. 15 mi SE of El Morrison,

—
~~

“Castilleja sp.

P 2435
SOLANACEAE
Chamaesaracha villosa Rydb. 12 MEX: Chih. 20 mi W of Ojinaga, P, Tu, &
Si 2465.

MEX: N. L. between Monclova and Monterrey,
P & Tu 2307

m
ho

Physalis bederaefolia
Gray

VERBENACEAE
*Tetraclea coulteri: Gray 21 MEX: Coah. ca. 30 mi NE of San Pedro, P &
Tu 2690,

ZYGOPHYLLACEAE
+*Sericodes greggit Gray ca. 15 MEX: Coah. ca. 54 mi N of San Pedro, Si, O, &
P S842,

Larrea tridentata
°.) Cov 13) TEX: Hudspeth Co. 1 mi S of Tommy’s Town,
P S6(

REFERENCES

BACON, J. D. 1974a. Chromosome numbers and taxonomic notes in the genus Nama

ia ie Brittonia 26: LOL-105.
974b. The genus Nerisyrenia (Cruciferae): A sg as and cyto-

taxonomic eee Ph. D. Thesis, The University of Texas, Aust

CAVE, M. S. [ed.]. 1956-1964. inde to plant chromosome cuehae. University of North
Carolina Press, Chapel Hill.

DARLINGTON, D. C. and A. P. WYLIE. 1956. Chromosome Atlas of Flowering Plants.
George Allen and Unvin Ltd., London.

DAVIS, W. S. and H. J. THOMPSON, 1967. A revision of Pefalonyx (Loasaceae) with
a consideration A affinities in subfamily Gronovioideae. Madrofio 19: I-18.

FEDEROV, A. [ed.]. 1969, Chromosome Numbers of Flowering Plants. Acad. Sci.
U.S.S.R., ee Botanical Institute, Leningrac

HIGH, S. D. 1974. A pain sdy of ea gypsum restricted taxa in Trans-Pecos
Texas. M. S. Thesis, Sul Ross State University, Alpine, Texas.

JOHNSTON, I. M. 1941. Gypsophily among Madi can desert plants. J. Arn. Arb. 22:

45-170.
MOORE, R. J. [ed.]. 1967-1971, 1972. Index to plant chromosome numbers. penn en
Bureau Plant Taxonomy and eae ee Utrecht, Netherlands. Regnum Ve
9

90,
Gee R. [ed.]. 1965-1967. Index to plant chromosome numbers. International
Bureau - Plant Taxonomy and Nomenclature, Utrecht, Netherlands. Regnum Veg.
-
PARSONS a ; . Gypsophily in plants—a review. Amer. Mid. Nat. 96: 1-20.
RTE _ - . cen Disjunct diecaciusions in the New World Zygophyllaceae. Taxon
33

rte . M. and S. A. SLOAN. 1975. Polyploid ear in gypsum and non-
gypsum floras of the Chihuahuan Desert. Sci. Biol. 37-38.
., and B. L. TURNER. 1963. penis oe ers in the Compositae. VII.
Additional species from the southwestern United States and Mexico. Madrofio 17: 128-140.
. 1977. Aspects of the plant Melee of the gypsum outcrops of the Chihuahuan
Desert. In Transactions—Symposium on the Biological Resources of the Chihuahuan
Desert Region, U. S. and Mexico. National Park Service, Washington, D. C.
TURNER, B. L. 1972a. Two new species of Isoconra (Compositae—Astereae) from north
central Mexico. Sida 5: 23-25

197 2b. — ai a new monotypic genus from north

central Wastied: Amer, J. Bot. 59: 180-182.
: 97 20: Two new exmsophiln species of Gaillardia (Compositae) from north
central Mexico. Southwe a 81-190.
1973a. Two new 5 gypaophilow species of Machaeranthera (Asteraceae—Astereac)
from north central Mexico. Phytologia 26: 116-120.
’ i Machacranthera retin (Asteraceae) a bizarre new gypsophile from
north mene Mexico. Amer. J. Bot. 60: 836-838.
. , and M. C, J¢ INSTON, ae Chromosome numbers in the Compositae. HI.
Certain Mexican species. Britt 13: -69,
, A. M. POWER LL, a oa J. W ATSON. 1973. Chromosome numbers in Mexican

r. is 2-596.

» EJ. D. BACON, R. L. HARTMAN, M. C. JOHNSTON, 1 J. WAT-

SON, and G. L. WEBSTER. 1975. Checreceas numbers for North ee Euphor-
biaceae. Amer. J. Bot. 62: 494-500.

WATERFALL, U. T. 1946. Observations on the desert gypsum flora of southwestern Texas
and adjacent New Mexico. Amer. Mid. Nat. 36

NOTES

NEW AND VERIFIED ADDITIONS TO THE NEBRASKA VASCULAR
FLORA. II.—In a previous paper Churchill (Sida 6: 116-117. 1975) reported
7 new and 1 verified vascular taxa for Nebraska. Further field and her-
barium work during 1975 has produced additional new records for Nebraska.
All specimens cited are deposited at the Nebraska State Herbarium (NEB)
with a duplicate set of the Churchill and Shildneck collections deposited at
the University of Kansas (KANU). New taxa for the state are marked with
an asterisk (*).

* Juncus effusus L, Richardson Co.: NW. of Verdon (Sec. 34, T3N, R15E),
in shallow water of roadside ditch, 26 June 1975, Shildneck C-7624. This ex-
tends the range of this rush north from Kansas and Missouri.

* Aristida purpurascens Poir. Jefferson Co.: 5 miles S. of Fairbury on

0 & j eroded sandstone ravine, 1 October 1975, Churchill 6811.
This extends the range north from Kansa

* Alliaria officinalis Andrz. Webster ee 1 mile S. of Red Cloud, flood-
plain woods along Republican River, 28 May 1975, Churchill 5427. This is
the first report for the state of this introduced and naturalized taxon from
Europe.

Rorippa austriaca (Crantz) Besser. Cuming Co.: 6 miles SE. of Beemer
on Hwy US 275, wet roadside ditch, 31 May 1975, Churchill 5511. Stuckey
(Sida 4: 279-430. 1972) reported a single collection of this adventive taxon
from Europe for Nebraska. Thus, this collection is the second report but
the first specimen for the Nebraska State Herbarium.

* Potentilla plattensis Rydb. Cheyenne Co.: E. of Potter, wet grassland,
20 June 1953, Kiener 29423. This extends the range east from Colorado and
Wyoming.

Rosa multiflora Thunb. Jefferson Co.: 5 miles S. of Fairbury on Hwy 15

just E., on steep banks of shaded sandstone ravine, 1 October 1975,
Churchill 6776. This is the first verified collection of this Asia rose collected
in the wild. Several collections in NEB by J. M. Bates of this taxon are
marked with the abbreviation ‘‘cult.’’, but other duplicates of the same
number give no indication whether or not they were collected under culti-
vation.

Desmodium sessilifolium (Torr.) T. & G. Jefferson Co.: 5 miles S. of Fair-
bury on Hwy 15 & just E., grassy W-facing sandstone bluff, 1 October 1975,
Churchill 6791. This verifies an earlier collection from Jefferson Co.: Fair-
bury, 20 July 1891, Price s. n. This extends the range north from eastern
Kansas.

Hypericum mutilum L. Jefferson Co.: 5 miles S. of Fairbury on Hwy 15
& just E., shaded Quercus tributary creek, 1 October 1975, Churchill 6745.
This verifies two earlier collections from Jefferson Co.: Endicott, 22 August

SIDA 7(1): 91. 1977.

92

1893, Woods & Saunders 2092; Fairbury, 7 July 1890, Price s. n. These col-
lections extend the range north from eastern Kansas.

* Lobelia inflata L. Burt Co.: 5 miles S, of Decatur & 1.5 miles W., slope
on grazed upland woods, 5 October 1975, Churchill 6850. Richardson Co.:
Rulo-White Cloud road (NE, Sec.26, TIN, R18E), edge of wooded slope
next to road, 13 September 1974, Shildneck C-7151. Two earlier collections
were reported by Winter (An Analysis of the Flowering Plants of Nebraska.
Univ. of Nebr. 1936), one specimen was incorrectly identified and the other
cultivated. This extends the range east from Iowa and north from Kansas.

I wish to thank Dr. Ronald McGregor for verification of Rorippa and
Potentilla.—Steven P. Churchill, School of Life Sciences, University of
Nebraska, and Division of Botany, University of Nebraska State Museum,
Lincoln, Nebraska 68588.

SARCOSTEMMA CYNANCHOIDES (ASCLEPIADACEAE) IN ARKANSAS
—Sarcostemma cynanchoides Dene. is widely distributed from central Mex-
ico to southern Utah and from extreme southern California to about 97 de-
grees west latitude in Texas and Oklahoma. As interpreted by Holm (Ann.
Mo. Bot. Gard. 37: 528. 1950.), this species may be divided into eastern and
western subspecies. The eastern subspecies, S. cynanchoides subsp. cynan-
choides, was recently found to occur in Arkansas where it is now known
from two localities. One station from the north central part of the state rep-
resents an extension of about 275 miles from the previous easternmost col-
lection from near Stillwater, Oklahoma. This collection of Thomas and Reid
was distributed under the label of Northeast Louisiana State College, Mon-
roe, as Cynanchum laeve (Michx.) Pers., a species superficially resembling
S. cynanchoides in its climbing habit and cordate leaf shape. The flowers
of the two are strikingly different, those of S. cynanchoides being much
larger, and having a corona of 5 inflated segments. The flowers of C. laeve
are small, never more than 4 mm in diameter and have a corona of delicate
membranous scales. Subsequent to the first draft of this note, Dr. Mahler
informed me of a second Arkansas record for S$. cynanchoides. This collec-
tion, made by Demaree, is from western Arkansas and is from a locality
nearly halfway between the collection cited above and the acregsaune sta-

Reid 20640 (MO, SMU). Franklin County. High bottoms of Arkansas River.
Common. P.O. Ozark. Elev. 380 ft. 3 Sept. 1966. Delzie Demaree 54246 (SMU).
—D.L. Spellman, Missouri Botanical Garden, St. Louis, MO. 63110

SIDA 7(1): 92, 1977.

PSILOTACEAE SYMPOSIUM REPRINTS AVAILABLE

The recent symposium entitled ‘‘Taxonomic and Morphological Relation-
ships of the Psilotaceae,’’ which was presented during the 1976 AIBS meet-
ings at Tulane University, will be published in the January-March issue of
BRITTONIA. This symposium, under the chairmanship of Richard A. White
(Duke University) and sponsored by the American Fern Society, the Ameri-
can Society of Plant Taxonomists, and the Pteridological, Paleobotanical,
Structural and Systematic Sections of the Botanical Society of America,
summarizes the major points of the current controversy surrounding this
enigmatic plant group.

Contributors are: David W. Bierhorst (University of Massachusetts), The
systematic position of Psilotum and Tmesipteris; Patricia G, Gensel (Uni-
versity of North Carolina), Morphologic and taxonomic relationships of the
Psilotaceae relative to evolutionary lines in early land vascular plants;
Donald R. Kaplan (University of California—Berkeley), Morphological status
of the shoot systems of Psilotaceae; and Warren H. Wagner, Jr. (University
of Michigan), Systematic implications of the Psilotaceae. The set of papers,
together with introduction and discussion, will be printed as a unit and
become available in April as a separate through the New York Botanical
Garden. Copies can be obtained by sending your order, along with $3.00 per
copy ($2.50 each in quantities of 30 or more), to: BRITTONIA, New York
Botanical Garden, Bronx, NY 10458. No orders will be filled unless accom-
panied by payment.

SIDA 7(1): 93, 1977.

SIDA ‘usin

VOLUME 7 NUMBER 2 DECEMBER 1977

CONTENTS

Pollen exine morphology and its adaptive significance.
Walter H. Lewis. oo

A new species of Hedeoma (Labiatae) from north central
Mexico. Robert S. Irving. 103

Case studies: a new approach to teaching plant systematics.
Tod F. Stuessy. 107

Annotated list of vascular plants in Mobile, Alabama.
Michel G. Lelong. 118

Synopsis of Suaeda (Chenopodiaceae) in North America.
Christine O. Hopkins & Will H. Blackwell, Jr. 147

Taxonomy and distribution of Gentiana (Gentianaceae)
in Mexico and Central America. |. Sect. Pheumonanthe.
James S. Pringle. 174

NOTES: Juvenile leaves in Oklahoma Marsilea (Marsileaceae). 218—Ammoselinum butleri
(Umbelliferae), new to North Carolina. 220—Additions to the Louisiana flora. 220—New
combinations in Mexican Pavonia (Malvaceae). 223—Seed germination and seedlings of

Krameria lanceolata (Krameriaceae). 224.

NOTICE. 226.

US ISSN 0036-1488

SIDA, CONTRIBUTIONS TO BOTANY
Founded by Lloyd H. Shinners, 1962

Editor & Publisher
Wm. F. Mahler
SMU Herbarium

Dallas, Texas, 75275

Associate Editor Assistant Editor

John W. Thieret Barney L. Lipscomb
Northern Kentucky University SMU Herbarium
Highland Heights, Kentucky, 41076 Dallas, Texas, 75275

Contributors to Sida should refer to the latest issues for style, follow the
CBE Style Manual (38rd ed.), and use the ‘‘adopted”’ journal abbreviations

given in Botanico-Periodicum-Huntianum.

Subscriptions: Libraries—$6.00 (U.S.) per year; individuals—$8.00 (U.S.)

per volume; numbers issued twice a year.

©
Sida, Contributions to Botany, Volume 7, Number 2, pages 95-226,
Copyright 1977
by Wm. F. Mahler

POLLEN EXINE MORPHOLOGY AND ITS
ADAPTIVE SIGNIFICANCE

WALTER H. LEWIS
Missouri Botanical Garden and Washington University
St. Louis, Missouri 63130

The contribution of the research papers? presented in this symposium,
“Systematics and Evolution of Pollen Characters in Angiosperms,”’ could be
summarized as the potential gift that new advances in electron microscopy
can make to systematics.

I am reminded of an earlier stepping stone in palynological research
by Professor G. Erdtman, who developed the acetylosis procedure, a tech-
nique that allowed the first concise and intimate view of pollen wall morph-
ology. However much progress in this field we attribute to Professor Erdt-
man, he in turn told me some time ago that his inspiration in palynology
had really been the outcome, the natural evolution, if you will, of the
pioneering research of Dr. R. P. Wodehouse, whose drawings of stained
pollen grains remain classic examples of what may be accomplished with
a minimum of instrumentation and much dedication. This modesty was
nevertheless characteristic of a great scientist whose impact on palynology
is everlasting, and appropriately this symposium is dedicated to Professor
Erdtman’s memory.

Let me capsulate the research papers presented to you this morning
and add a few remarks of my own on potential adaptive significance of the
diversity we have observed

Recent research by Richard C. Keating (1973) on the Flacourtiaceae
using only light microscopy has now been supplemented by both trans-
mission and scanning electron microscopy, greatly enhancing observations
of wall structure and grain surface. Keating considers the triaperturate

"Summary address, Pollen Symposium, A.I.B.S. meetings, Arizona State University, Tempe,
July, 1974,

2 In sequence of presentation are:

Keating, R. C. 1975 [1976]. ‘Vrends of specialization in Sone of Sora with
gee ope ations of Cochlospermaceae and Bixaceae. Gran 9-49,

owicke, J. W. 1975 [19761]. Pollen morphology in the order ee rmae. Grana 15:
oS

Taylor, T. N. and D. A. Levin. 1975 [1976]. Pollen morphology of Polemoniaceae in
relation to systematics and pollination systems: scanning electron microscopy. Grana 15:
1-112

Tomb, A. S. 1975 [1976]. Pollen morphology in tribe Lactuceae (Compositae). Grana
9-89.

Skvarla, J. J.. B. L. Turner, V. C. Patel and A. S. Tomb. 1977. Pollen ee in

the Compositae and in miGepholaneslly related families. In V. H. eed ie (ed.) Biology

and Chemistry of the Compositae. Academic Press, New York (in 1

SIDA 7 (2): 95—102. 1977.

96

grain, which is typical of many genera in the Flacourtiaceae, primitive.
This opinion correlates well with vegetative and floral features, and thus
places the family among the more primitive Angiosperms. Therefore, he
assigns a basal position to the family in the Violales, for none of the
other members of the order shows so primitive a pollen morphology or so
little diversity. Of what variation does exist, however, Keating made the
important observation that it is not unidirectional in all groups. For ex-
ample, there is a tendency for near solid tecta to have been derived from
finely reticulated tecta, as well as a tendency to form well developed infra-
tectal spaces among both primitive and advanced taxa. These suggest non-
homologous derivations and signal caution to the phylogenist who would
base hypotheses on exine characters alone. Here, too, are examples of
parallel evolution that again flash caution to those who would interpret
like-morphologies invariably as having an immediate, common ancestory.

Keating outlined an interesting microperforate tectal sequence in the
Flacourtiaceae from those pollen having numerous micropores and_ flat
tecta to those with micropores clumped into orifices between high ridged
tecta. This condition masks a reticulated exine, for if it were true reticula-
tion, the bacula would be organized below the ridges only, which is not
the case. This sequence is valuable not only because it suggests an origin
of reticulated exines from essentially tectate one, but it emphasizes the
wide occurrence of micropores, a subject I shall return to shortly.

An undoubted feature of this symposium is the enormous scope of the
papers, perhaps epitomized by the report of Joan W. Nowicke who char-
acterized palynologically 16 families in the Centrospermae. This mass
approach to pollen morphology is not one that I am criticizing. Quite the
opposite, for I hope we can encourage such studies involving extensive
research and synthesis of as many parts of as many organisms as we
possibly can in our quest for an understanding of broad evolutionary
pathways.

The report by Nowicke is, therefore, possibly a unique feat for shear
number of taxa involved, but it is also much more. For example, the find-
ing of spinulose, perforated exines universally in the Centrospermae is
perhaps as important a unifying character as the curved embryo or the
unique sieve tube plastids now used to distinguish members of this order.
Moreover, these new data do not preclude and perhaps even support the
inclusion of the Caryophyllaceae and Molluginaceae, which otherwise do
not possess betalains, with betalain-possessing families, and, thus, placing
in the Centrospermae taxa having either anthocyanins or betalains.

Another significant contribution of her work is finding pollen grains in
the Gyrostemonaceae and Achatocarpaceae distinct from those of the
Phytolaccaceae. This confirms their separation from the latter, and ques-
tions their retention in the Centrospermae. In Nowicke’s opinion, the
pollen characters of Theligonum are sufficiently different from all other

97

Centrospermae known to suggest, in correlation with sporophytic features,
its removal from the order, and perhaps, as suggested by Wunderlich
(1971) placed in the Rubiaceae.

These more primitive families served as a springboard for the report
by Thomas N. Taylor and Donald A. Levin who surveyed the more ad-
vanced Polemoniaceae. They found that grains with colporate and pan-
toporate apertures predominate, although tropical genera are typically
pantoporate. Among former genera they observed a trend from large to
small size pollen. Among those with pantoporate pollen, they noted an
increase in aperture number with a decrease in pollen size from a tropical
genus (Bonplandia) to one with strong temperate affinities (Polemonium).
These and other data were compatible with a morphological and chromo-
somal phylogeny proposed by Verne Grant (1959). Some differences exist,
however, and these need exploration.

Looking beyond comparative morpholog »y, Taylor and Levin investigated
the intriguing question of correlations between, for example, pollen size,
which is so variable in the Polemoniaceac, and pollinator. They found a
significant relationship between the large pollen transported by bats as
well as between much smaller grains pollinated by various insects. They
do not believe that these correlates are due to coadaptation directly,
but rather to style length in which selection favors pollen. size having
enough metabolites for pollen tube growth to reach the embryo sac. How-
ever, we cannot dismiss the observation that a direct correlation may
exist between the large, hexaploid pollen and large flowers of Cobaea,
and bat pollination due to mechanisms as yet far from understood.

With regard to exine architecture, however, Taylor and Levin saw no
apparent association with pollination mechanism and environmental pa-
rameters, a point I shall return to subsequently. They concluded that
parallel evolution, at least in the Polemoniaceae, remains unidentified in
pollen morphology, an interesting possibility in so variable a family
palynologically.

The remainder of the symposium was devoted to the highly evolved
and palynologically diverse Asteraceae. In his discussion of chicory exines
(tribe Lactuceae . spencer Tomb typified the wall structure as
echinolophate, that is to say, with elaborate systems of ridges and spines

isposed around and between apertures and pseudoapertures. This feature
is the outstanding condition of the Lactuccac, for all subtribes recognized
possess such grains. Some, but not all, subtribes also have echinate pollen,
a much less elaborate spinulous pattern in which the spines are usually
seattered over the surface and obvious gaps (lacunae) are absent. Tomb
suggested subtribal modifications in classification based on these and other
characters of the pollen correlated with morphological and chromosomal
data. He also presented the probable evolutionary pathways for these pollen
from echinate to echinolophate, from moderate-sized cavea to those exines

without a gap between the sexine and nexine, from tectate, perforate
lacunal surfaces to nontectate lacunae, and from large sized grains to
those having smaller sizes. These may be unidirectional trends, although
some caution is necessary. Via polyploidy, for example, the more evolved,
smaller grains could have independently given rise to much larger grains
in various subtribes, and these, therefore, would be of more recent origin
than the smaller, less ploid grains. Tomb also made this observation while
discussing evolutionary direction involving the origin of echinolophate
exines, namely, that less elaborate echinate exines may have evolved by
reduction from more elaborate ones and that this atavism must not be
confused with a single origin. In other words, the echinate condition can be
both primitive and advanced.

The second paper on asteraceous pollen was presented by John J. Skvarla
as a general survey of research on the tribe Inuleae, a tribe whose pollen
exine is much less dramatically claborate than that of the chicories, but
none-the-less intriguing. He recognized two basic pollen types in the
Inuleae, the first having simple or ramified bacula usually with abundant
infrabaculate micropores (a type clearly related to the Senecionecae and
Heliantheac), and the second in which the bacula are very complex having,
for example, several levels of internal tecta, but lacking infrabaculate
micropores and cavea. The second morphology is characteristic of the
subtribe Tarchonantheae, as well as several genera scattered throughout
the tribe (Adenocaulon, Gymnarrhena, Osmites-Osmitopsis), a morphology
which the author notes is common among the tribe Anthemideae. As a
result of this study, Skvarla suggests that much of the infratribal classifica-
tion of the Inuleae is artificial.

We have observed among these surveys, involving largely comparative
morphology of the exine, the substantial value of palynology in systematic
studies. Clearly, palynological evidence may be useful in placing taxa of
uncertain affinity, of suggesting taxonomic rearrangement, as well as cor-
roborating other lines of evidence in order to establish evolutionary
hypotheses.

Exine diversity is extraordinary in the sample we have just observed,
illustrating surely the tremendous competence of flowering plants in de-
signing and building these walls. But why, we might ask, have these plants
bothered? Are we to assume that what we have observed are but mean-
ingless manifestations; if not, can we recognize that there are adaptive
elements at play and that diversity has resulted from selective forces
(Heslop-Harrison 1971)? Clues do exist of the adaptive value of pollen wall
structure. Basically the wall serves to protect contents from desiccation,
it facilitates or permits efficient germination, and it can also be expected
to have adaptive roles concerned with dispersal. In relation to these func-
tions, let us examine the most conspicuous feature of the wall so univer-
sally stressed during this symposium, the aperture.

99

Apertures form the preferred paths of exit for the pollen tube, but their
function extends beyond providing an escape route. For example, they
act as regulators controlling the gain and loss of water. Very briefly, the
plates, plugs and granules which cover many apertures consist of typically
impermeable exine which can be observed to close on dehydration caused
by an infolding of the intine at the aperture site. Imbibing water, however,
brings about a gaping of the aperture with the exine being carried apart
on the surface of the extended intine or left at the aperture margins. A
further aid to this regulation is the so-called pollenkit or lipid of tapetal
origin which on desiccation forms a coating over the apertures effectively
obstructing water loss. As regulatory devices, this apertural system seems
simple enough, but they do illustrate regulatory functions that have adap-
tive values and are therefore likely to have evolved under the pressure of
selection.

A conspicuous feature of apertures is their numerical difference. Why,
for example, has the polyaperturate condition apparently evolved from
the more common. triaperturate dicotyledonous pollen? Could aperture
number itself be adaptive? If so, there must be functions other than pro-
viding for the exit of a single pollen tube. One function is certainly related
to the release of enzymes and other proteins from moistened grains to
stigmatic surfaces. Among these proteins emitted from the apertural intine
are lytic enzymes, suggesting a function associated with the digestion of
substrate on the stigma and thus with the initial nutrition of the tube.
Other substances are concerned with compatibility reactions which de-
termine what pollinations will proceed to fertilization and the next genera-
tion and which ones will not. They are recognition substances which may
or may not be recognized by sites on the protein pellicle of stigmatic
papillae (Mattsson et al. 1974). Therefore, aperture adaptation is not only
related to germination, but to the control of the breeding system and the
physiology of the emerging tube as well. When each aperture is seen as a
site for storage and release of physiologically significant materials, and
not just an area for potential pollen tube exit, this places the evolution of
aperture number in a new perspective.

What of those parts of the exine beyond the apertures? Early observa-
tions by Wodehouse (1935) are still appropriate: pollen grains from those
plants having reverted to anemophily differ from those of their related
entomophilous species by possessing grains with thinner and smoother
exines. For example, grains of anemophilous Populus are smooth com-
pared to the reticulated exines of Salix, a genus largely of entomophilous
species. Or, on discussing the wide differences in pollen encountered in
the tribe Anthemideae (Asteraceae), Wodehouse noted that the insect
pollinated species of Anthemis (Leucanthemum) and Chrysanthemum have
grains with well developed spines and_ thick exines, whereas those of
Artemisia, Crossostephium and Picrothamnus, which are wind pollinated,

100

have grains in which the spines are vestigial or absent and the exines are
much less thick. He concluded that members of this tribe may be divided
rather sharply into two groups consisting of those which are echinate
grained and insect pollinated and those with smooth or nearly smooth
grained and wind pollinated.

There are also additional characteristics of pollen correlated with mode
of pollination, Clearly the lipoidal component of the exine differs, for it is
absent or present in only small amounts in anemophilous species, while
being a most conspicious adjunct of the pollen of entomophilous plants.
This pollenkit provides not only much of the visual and olfactory stimulus
needed to guide insects, but it also provides the means for affixing the
grains to the pollinator’s body (Heslop-Harrison 1971).

But what of the elaborate architectural detail of the sexine which I
should mention is largely of sporophytic, not gametophytic, origin?
Proteinaceous material can now be recognized in the intra- and interbacular
spaces of the wall as well as in the cavea between the sexine and the
nexine among asteraceous pollen, This material is entirely of sporophytic
origin, having been injected into the cavities of the sexine following
derivation from membrane-bounded cisternae released from the tapetum
of the anthers during its dissolution (Heslop-Harrison et al. 19738). On
moistening, these sexinous-held sporophytic proteins are discharged by
the pollen through the micropores in the tectum, micropores which we
have observed as a very common feature among the diverse families
observed today. This discharge has been particularly well demonstrated
by its release within 5 to 30 seconds following pollen moistening of numerous
antigens and allergens from the micropores at the base of spines of
Ambrosia and Cosmos (Knox and Heslop-Harrison 1971, Howlett et al. 1978).
Therefore, there is a functional significance to the complex internal archi-
tecture of the tectal and baculate layers of the exine: it is basically a
storage site for materials derived from the tapetum which, through micro-
pores, provides passage for injection of stored materials and for their
release when pollen is moistened. In nontectate exines, the spaces between
the muri are also concerned with the storage of tapetum derived materials.
In the Asteraceae, the exine storage capacity is further enhanced by the
development of cavea, which provide enormous reservoirs for products
transferred from the sporophyte.

The materials undoubtedly function in several ways. They possess a role
in inter- and infraspecific incompatibility systems where the genetic con-
trol resides with the parental sporophyte (Heslop-Harrison 1967). This has
been shown for Populus (Knox et al. 1972) where proteins from the pollen
of one species have been used to make pollen of an incompatible foreign
species acceptable, and in another example for Cosmos bipinnatus (Howlett
et al. 1973), extracts of compatible pollen make self-incompatible pollen
fully self-pollen acceptable by the stigmas. In addition, exine-held fractions

101

have recently been shown associated with incompatibility responses in the
Brassicaceae (Heslop-Harrison et al. 1974). It seems clear, that for a wide
range of flowering plants the exine is the site of recognition materials in
incompatibility systems. I should note that the allergenic response in
man to these outer wall-based proteins, as well as those held by the intine,
is no more than an unfortunate accident of evolution, for their role is to
act as recognition substances during pollen-stigma interactions, not to
harass those among us who produce immunoglobulin E on contact followed
by alarming allergic symptoms.

Briefly, we have glimpsed at several, newly discovered roles involving
the pollen wall, largely thanks to the research of Professor J. Heslop-
Harrison and his team at the Royal Botanic Gardens, Kew, and to Pro-
fessor John Rowley and his associates at the Palynological Laboratory,
Stockholm. These functional roles of pollen can be related to a number
of morphological characters only recently exposed clearly by electron
microscopes. For example, the widespread asteraceous cavea could be
considered adaptively important, as indeed could the frequency, size and
position of the micropores, and the diverse baculate spaces of the sexine.
All relate to the storage and conveyance of significant proteinaceous com-
pounds from one generation to the next, yet before the advent of electron
microscopy, these features were virtually or entirely unobservable. We
need more observations and experiments before the full significance of
exine sculpturing is realized. It is now apparent that such diversity is
not all incidental, rather, that it is more likely to be accidentally interwoven
with adaptive elements which has given us a highly intricate and in many
cases selected covering to the male gametophyte of higher plants. More
physio-morphological research similar to that described today and taking
place at Kew and in Stockholm is imperitive before the relevance of this
minute plant, the pollen grain, is fully understood and the selective ad-
vantages of its innumerable diversities are fully appreciated.

REFERENCES

BELIN, L. and J. ROWLEY. 1971. Demonstration of birch pollen allergen from isolated
pollen grains using immunofluorescence and a_ single radial imal used technique.
Int. Arch. Allergy Appl. Immunol. 754-769,

GRANT, V. 1959. Natural History of a Phlox family. Volume 1 Systematic Botany.
Ree Nijhoff, The oS 280
ESLOP-HARRISON, J. 1967. Ribosome sites and S gene ees Nature 218: 9

- 1971. Sporopollenin in the biological context, 1-20. . Brooks, P, oor nt,
M. Muir, P. van Guzel & G. Shaw (eds.) Sporopollenin., reac Press, London and

New York

ac]

Boe ey ESLOP- hea R. B. KNOX and B. HOWLETT. 1973. ae
wall peaueias: ene ic’ _and ‘sporophytic’ fractions in the pollen walls of the Mal-
vaceac. Ann. Botany 37: -412

KNOX and 'Y. HES SLOP-HARRISON, 1974. Pollen-wall proteins: exine-
a eld gato ae sociated with the incompatibility response in Cruciferae. Theor. Appl.
Genet. 44: 133-137.

102

HOWLETT, B. J., . KNOX and J. HESLOP-HARRISON, 1973.
rele ase of the eee eae E from intine
and Cosmos. J. Cell Sci. 13: 603-619,

Pollen-wall proteins:
and exine sites in pollen grains of ragweed
N 1973. Pollen penne and relationships of the Flacourtiaceae. Ann.
Missourt Bot. Gard. 60: -305.

KNOX, R. B. and J. eee HARRISON. 1971. Pollen-wall proteins: localization of
antigenic and ve es proteins in the pollen-grain walls of sa vous spp. (ragweeds).

Cycobios Had) -54,
: WILLING ae A. E. ASHFORD. 1972.

Role of pollen-wall proteins as
Kacbe Alcon pee in int Nat 2

yecific incompatibility in poplar ure 237: 381-383
— N, O., R. B. KNOX, a HESLOP-HARRISON and Y. HESLOP- HARRISON.
974. Protein pellicle of stigmatic papillae as a probable recognition site in incompatibility
reactions. Nature 247: 298-300.
WODEHOUSE, R. P. 1935. Pollen Grains. McGraw-Hill, New York.
WUNDERLICH, R. 1971. Die systematische Stellung von a Ossterr. Bot. Z.
119: 329-394

=

A NEW SPECIES OF HEDEOMA (LABIATAE)
FROM NORTH CENTRAL MEXICO

ROBERT S. IRVING
Department of Biology, University of Arkansas, Pine Bluff 71601

Hedeoma is a widespread genus distributed principally in southwestern
United States and northern Mexico. Many species of Hedeoma are abundant
and somewhat weedy herbaceous annuals and perennials and make up what
might be termed the central core of the genus. These species are seemingly
evolutionarily active and their phenetic coherence is easily perceived.
Other subgroupings within Hedeoma are quite different. Their species are
robust perennials or semi-shrubs, confined largely to isolated mountains
in the Chihuahuan and Lower Sonoran deserts. As a group they have been
poorly collected and our understanding of their patterns of variation and
evolutionary trends is likewise meager. In recent years, with more inten-
sive field work in our desert systems, a number of new taxa and many new
collection records have been added to these groups (Irving, 1967; Irving,
1970; Moran, 1969). This work has revealed that intraspecific variation is
much wider than was once delineated (Irving, 1968) and that many ‘‘clear-
cut” taxa must now be viewed more cautiously. Moreover, the genus
Poliomintha, allied to the shrubby hedeomas of the Chihuahuan Desert, is
questionably distinct. Although no changes are proposed here, the recent
discoveries of new taxa and the greater understanding of morphological
variation, especially in such species as H. palmeri and H. montanum, have
left the generic boundary between Hedeoma and Poliomintha tenuous.

Field efforts toward the forthcoming flora of the Chihuahuan Desert
(Johnston, Hendrickson and collaborators) have uncovered still another
new and heretofore uncollected desert species.

HEDEOMA johnstonii Irving sp. nov. Fig. 1.

Herbae perennae ad 35 cm altae caulibus multis praeter modos comosos
elabris. Folia glabra ovato-oblanceolata 10-20 mm longs 4-7 mm lata dentata
base attenuata apice acuta petioli breves. Cymulae 1-3-florae. Calycis tubus
8-12 mm longus tubulosi-infundibuliformis limbus bilabiatus, dentes superi
labium reflexum formantes dentes inferi adscendents. Corolla roseoviolacea
tenuis ca 45 mm longa. Mericarpia badia ca 2 mm longa 1 mm lata.
Chromosomatum numerus ignotus.

Wiry perennial herbs up to 30.0 cm tall; shoots ascending or decumbent,
numerous, branching primarily from the base and rooting at the lower
nodes, glabrous except for a minute tuft of retrorsely curling hairs at each
node. Leaves spreading or ascending, stiff membranous in texture, glabrous,
conspicuously glandular punctate on the lower surface, ovate-oblanccolate,

SIDA 7(2): 103—106. 1977.

104

A

Fig. 1. Hedeoma johnstonii. A. Habit, X %; B. Leaf undersurface, X 5;
C. Corolla, X 5; D. Calyx, X 5.

105

10.0-20.0 mm long, 4.0-7.0 mm wide, dentate with 4-5 pairs of teeth termi-
nating the costae, apex acute, base attenuate to a short petiole (ca, 1.5
mm long), midrib and secondaries conspicuously elevated. Cymules 1-3
flowered, well-spaced along the upper one-third of the stem; primary
peduncles ca. 1.0 mm long; primary pedicels ca. 3.0 mm long; bracteoles
lanceolate, equal to or slightly longer than the pedicels. Calyx 10.0-15.0 mm
long (base to tip of upper teeth), tubular-funnelform, not gibbous and dilated
upwardly, sparsely hirsute; upper teeth partially united forming a small
but conspicuously reflexed lip, the lobes triangular ca. 1.0 mm long, sparsely
ciliate; lower teeth deltoid below ca. 1.0 mm wide, abruptly tapering to
an aristate apex, ca. 1.0-2.0 mm long, hirsute-ciliate; annulus located just
below the teeth, included. Corolla showy, long and slender, up to mm
long from its base to the tip of the upper lip, pink-violet, broadly annulate
within where seated in the calyx; upper lip short, ca. 4.0 mm long, 3.0 mm
wide, emarginate; the lower lip spreading, ca. 5.0 mm long, ca. 3.0 mm
wide, Nutlets 2.0 mm long, 1.0 mm wide. Chromosome number unknown.

Type: MEXICO. Coahuila: Canyon Hundido on north side of Pico de
Centinela, Sierra del Jardin, 8 km e. of Rancho El Jardin. “Steep canyon
through igneous sierra, gravelly and sandy loam derived from extrusive
igneous rocks, 1500-2250 m.’’ 27 July 1973: M. C. Johnston, F. Chiang, T. L.
Wendt and D. Riskind 11803. (Holotype, TEX; Isotypes to be distributed)

Additional Collections. MEXICO. Coahuila. Sierra del Jardin e. of Rancho
El Caballo. 16 Sept. 1972. F. Chiang, T. L. Wendt and M. C. Johnston 9341.
Sierra Maderas del Carmen, Canyon El Dos. 3 Aug. 1974. T. L. Wendt and
A, Adamcewicz 469.

Hedeoma johnstonii is a distinctive species apparently endemic to the
steep igneous slopes of the Sierra del Jardin, Coahuila, Mexico. As with
many of the narrowly endemic species of Hedeoma its affinities are puzzl-
ing. Its habit is very reminiscent of Hedeoma bellum, an endemic of the
mcuntains of the western coast of Mexico, but, in the details of its calyx
and corolla, it is more closely allied to Hedeoma costatum and its allies,
It is readily separated from all species of Hedeoma by its glabrous stems,
its glabrous, dentate leaves, and its exceptionally long and showy corollas.

The specific epithet honors Marshall C. Johnston who not only first col-
lected the species and judged it to be the “most beautiful of all Hedeomas”’
but who also has been a continuing source of encouragement and support
for many years.

I should like to also thank Dr. Johnston for his Latin diagnosis and ex-
press my gratitude to my Research Associate, Susan Brenholts, for her help
in many ways. Acknowledgement is also made to Brenda Mahler for her
illustrations.

106

REFERENCES

IRVING, R. S. 1968. The systematics of Hedeoma. Ph.D, Thesis. University of Texas,

Austin,
. 1967. Hesperozygis pusilla (Labiatae): a new species from Mexico. Brittonia 19:

245-247,

- —, 1970. Novelties in Hedeoma (Labiatac). Brittonia 22: 330-345

MORAN, ete new dicots from Baja California, Mexico. Trans. Bin Diego Soc.
Nat. Hist. 15 (17): 95.

CASE STUDIES: A NEW APPROACH TO
TEACHING PLANT SYSTEMATICS

TOD F. STUESSY
Department of Botany, The Ohio State University, Columbus, Ohio 43210

The teaching of plant systematics is difficult at best. Because of the
broad scope of the subject matter, one of the major problems is deciding
exactly what types of information to include and what to ignore. The avail-
able options range from a very descriptive approach emphasizing familial,
generic, and/or specific recognition to a perspective heavily laced with
evolutionary theory and slanted toward expcrimental systematics or bio-
systematics. Somewhere between these two extremes lies the majority of
existing courses in plant systematics, with the exact emphasis depending
upon the interests and background of the professor involved.

Whatever the approach used in teaching plant systematics, in most cases
little opportunity is available for the students to learn to make taxonomic
decisions. Conspicuously lacking is some format or set of exercises for the
student to apply his new understanding of how to assess variation with
classical and/or experimental methods. He comes to know well the ways
to go about gathering data from plants, but he lacks the experience and
confidence to make sound taxonomic judgments.

Such a deficiency in students completing my own course in “Experimental
Taxonomy” has been embarassingly apparent (to me as well as to the
students) on the final examinations, in which I usually include a_ hypo-
thetical set of morphological, cytological, and chemical data and ask the
students to evaluate these for their taxonomic significance in the particular
case study. The students performed poorly, and they raised the question
of why this same type of exercise wasn’t given to them throughout the
course?

As a result of this student criticism, I have developed over the past few
years a set of exercises that are designed to help the student gain con-
fidence and proficiency in evaluating comparative data and in making
taxonomic decisions. These exercises, or ‘“‘case studies,’ are taken from
recent systematic literature and are chosen to reflect different taxonomic
problems as well as different types of comparative data. It is from the
belief that the case study method has proven valuable in my own classes
that I offer the following commentary for general appraisal. The purpose
of the present paper, therefore, is: (1) to describe in detail exactly what
a case study is; (2) to suggest possibilities for effective use of the case
studies in the classroom; and (3) to comment on the beneficial results of
using case studies, as judged from my own experience.

SIDA 7(2): 107—117. 1977.

108

DESCRIPTION OF A CASE STUDY

The way in which case studies are structured relates directly to the two
main learning objectives for the students: (1) they should be able to recog-
nize discontinuities in sets of comparative data, whether morphological,
cytological, chemical, etc: and (2) they should be able to evaluate these
observed discontinuities with respect to the various categories or ranks in
the taxonomic hierarchy. Other secondary objectives prevail, such as the
demonstration of confidence in making decisions and defending viewpoints,
but these aspects will be discussed later.

The particular case studies are selected with two criteria in mind: (1)
the papers should contain different types of data which should be displayed
in different ways, such as in bar graphs, scatter diagrams, and histograms,
and (2) the data presented must be displayed in such a way that taxonomic
decisions can be made from these figures and tables alone. The first cri-
terion is emphasized to give the student as much practice as possible with
handling and interpreting different types of data. The second criterion is
necessary so that the students may work on the comparative data them-
selves without relying on the author’s biased discussion. Papers are not
desirable, therefore, that treat stimulating problems but have only limited
amounts of displayed data. The studies selected need not all be excellent
from the standpoint of data-gathering or interpretation either; those with
deficiencies also can be used pedagogically to good effect.

The case study is divided into four sections: (1) statement of the prob-
lem; (2) materials and methods; (3) data; and (4) abbreviated reference.
The first section, the statement of the problem, indicates approximately at
what taxonomic level the study focuses. For example, it would be neces-
sary for the student to know that the problem at hand deals with infra-
specific rather than intergeneric delimitation. If some guidelines are not
given on the approximate rank involved, especially early in the course, the
students sometimes approach the problem from very different perspectives.
This often results in generating more confusion than understanding. The
second section on materials and methods is believed necessary so that the
students may see how the comparative data have been gathered (the
laboratory of the course may be oriented this way if so desired) and so
that they may evaluate critically the appropriateness of the methods for
the situation at hand. In most papers the section on methods is set off by
itself, which allows for easy removal from the paper proper; in other
papers this information must be extracted from the introductory remarks
or from the discussion. The third section of the case study contains the
data, which are the most important part of the exercise because they serve
as the basis for making the taxonomic decisions. As the author usually
arranges the data in tables and/or figures to suggest the relationships as
he views them, it is useful to scramble the data as presented and give un-

109

suggestive labels to the initial groups, such as Taxon A, Taxon B, Taxon C,
etc. In this way the student does not become prejudiced toward the author’s
viewpoint. The fourth and final section of the case study includes the
abbreviated reference of the paper from which the data were taken. The
title is omitted deliberately, because often it can give clues to the rank of
the taxa involved or other suggestions that might bias the student.

To give a better idea of what constitutes a case study, an example is
included below. It is important to remember that this is only one example
from out of hundreds that could have been selected; it is also a relatively
simple exercise. Some case studies, especially those involving cytogenetic
data, can be very complex, but they are long and space limitations prohibit
their inclusion here,

1. Problem: Provide a classification for the following taxa (A-D) using
any or all of the following categories: genus, species, subspecies, variety,
and form.

2. Materials and Methods: The taxa under study have been collected rather
sparingly. Specimens borrowed from 12 herbaria yielded a total of only
72 collections. This material was supplemented by field observations and
mass collections from 13 populations (Fig.

Buds collected in the field for chromosomal studies were fixed in a modi-
fied Carnoy’s solution (4 parts chloroform: 3 parts absolute alcohol: 1 part
glacial acetic acid) and refrigerated immediately. They were transferred
after 24 hours to 70% ethyl alcohol ‘colored’ with iron acetate and left
under refrigeration until squashes were made some months later. An aceto-
orcein stain (0.5% orcein in a solution of 1 part glacial acetic acid: 1 part
lactic acid: 1 part water) was employed. Mitotic counts were made from
germinating seeds following colchicine treatment.

A single count of n = 12 for Taxon B has been reported by Raven &
Kyhos (1961, Amer. J. Bot, 48:842). This was corroborated here by both
meiotic and mitotic counts for plants in population 698. Meiotic counts of n
= 12 were obtained for populations 700, 703, and 705, all of these being
referable to Taxon A (Fig. 5). Meiosis was regular in all pollen cells
examined except for occasional bridges at first anaphase in population 703.
Bivalents ordinarily formed two terminalized chiasmata at late diakinesis.

After thorough morphologic investigation of all 13 populations surveyed
in this study, certain characteristics of stems and leaves were found to
be of greater diagnostic value than others in rendering taxonomic judg-
ments. Figure 15 presents quantitative measurements of these characters.
For these data a minimum of 20 individuals from each population was
measured. One typical lower leaf per stem was used for blade width,
length, and length/width measurements. The total upper surface of these
leaves was examined with care in order to determine the minimum and
maximum trichome lengths. The single longest leaf per individual stem

—
~—

110

was utilized for maximum leaf length, and stem heights were accounted
for in attempting to evaluate the variation in the size of the plants. Table 1
supplements the data presented in Fig. 15. In it is summarized the remain-
ing most significant morphologic variation that was found.
3. Data: Figs. 1-15; Table 1.
4. Reference: Ellison, W. L. 1971, Brittonia 23: 269-279.

EMPLOYMENT OF CASE STUDIES IN THE CLASSROOM

Before the case studies can be used effectively in the classroom, an
introduction to concepts of taxonomy and classification (including grouping
and ranking) must be presented. If the students have not already learned
operational definitions for recognizing taxa and their subsequent ranking,
they will not be able to work the case studies. In addition, some introduc-
tion to different types of data-gathering and display techniques is recom-
mended. Only in this manner can the student begin to make progress in
the critical evaluation of the data included in the case studies. Some con-
siderable attention, therefore, must be given to the introduction of these
several aspects of taxonomy before assigning the case studies to the class.

I have found it helpful to begin working first with the students on case
study examples. These are structured exactly as the exercises described
earlier in this paper, but a fifth section is added containing a discussion
of the problem and a tentative classification, either drafted by the profes-
sor or extracted from the paper itself. In this way the students can check
their own classifications against the one provided. When the students have
mastered the examples, then they are ready to begin the exercises.

A number of mechanisms might be used for actually presenting the case
study exercises to the students, but I have found the following sequence
workable. An exercise is assigned to the class on one day with the an-
nounced intent of discussing the classifications on another day. At that
time, the students are asked to describe their systems, usually accompanied
by outlines on the blackboard, for the benefit of and discussion by the rest
of the class. If the class is large, time may preclude having every person
present his or her classification, but ideally everyone should have the
opportunity. After discussion of the individual classifications, with par-
ticular emphasis on the prevailing similarities and differences, the profes-
sor can present his own and the author’s views. Finally, the original paper
can be put on library reserve or other accessible location for additional
study by the students at their convenience.

BENEFICIAL RESULTS OF USING CASE STUDIES

In my own experience of teaching plant systematics with case studies,
I have noticed several beneficial results that accrue to the students. It is
realized that the observations to follow have not been quantified or experi-

111

mentally verified in any way; they are simply syntheses of feedback from
students on formal and informal course evaluations both before and after
using case studies. It is believed, however, that the benefits mentioned are
real.

1 DA HO

Fig. 1. Map showing ae of the taxa under study. Taxon A, tri-
angles; Taxon B, dots; C, circles; intermediates between taxa B
and C, half- closed circles; me D, squares. Numbers indicate locations
of mass collection

Figs. 2-8. Taxa A (Figs. 2-5) and D (Figs. 6-8). Habit, Figs. 2 & 6, X 0.5;
he ad, Figs. 3 & 7 (part only), i - 5; dise floret, Figs. 4 & 8, X 5; chromo-
somes in diakinesis, Fig. 5, X 1

113

The first beneficial result is that the students seem to finish the course
with a higher level of understanding and appreciation for plant systematics
than without employment of the case studies. The objectives of being able to
recognize discontinuities in sets of comparative data and being able to
evaluate these discontinuities with respect to the ranks of the taxonomic
hierarchy are achieved in large measure for almost everyone in the class.
The students gain first-hand experience in struggling to apply concepts of
taxa and categories, and this seems to cement these concepts more firmly.
The students also develop a more positive attitude toward plant systematics
in general, perhaps due to a better understanding of the difficulties in-
volved with making classifications.

Figs. 9-14. Taxa B (Figs. 9-11) and C (Figs. 12-14). Same scale and views
as in Figs, 2-8.

114

LONGEST LEAF (BLADE+PETIOLE) (mm)

692

693 Ee 'D

695 —_— — =

696 C

697 _ <a -— :

698 —o :

699 eorcper B

703 ll A

704 — el -

705 — |
+ erry) a a 1b hohe TH at J,

rear TRICHOMES fim)
"

692 ——
693 > —
694 —— oo
695 ms ef —
696 — a
i i ee
698 a -
699 _ —_
700 oe Lr , ‘1
703 — — ee ee
704 — a
705 = — -
ha Gk Sk a a,

BLADE WIDTH OF LOWER LEAF (mm)

693 a : p
694 hs cera

695 mbm -

696 - =o C
697 od ——

ze me ] B
6990

703 — - A
704 ——

Ce) | Un ieee .

692 ———

693 —— ~ D
694 a

69 mm

69E _ C
697 ———o-

698 a aed ]
699 = B
r0C ed ]

6 oukm

= A
705 eee

BLADE LENGTH/WIDTH

692 ——— 692 a eed

693 — D 693 = ‘D

694 ae 694 a - |

695 —— — 695 mbes = _

696 ee C 696 ote =n C

697 = — ; 697 a |

698 a B 698 ~ =p 7 B

699 ——=_ : 699 ——_

700 =a 700 == —

703 ——_ 703 a

704 — A 704 = A

705 705 — |
See ae A oe ee L

Fig. 15. Quantitative measurements o

taxa (A-D) in this study.

the standard
rectangle.

error

f
he range of variation observed
the length of the horizontal line. The resulting unweighted mean and

some characteristics of the four
is indicated by

wice

are indicated by the vertical line and accompanying

Table 1

Significant Comparative Morphological Charactera of the Taxa under Study.

Taxon A

Taxon 0

Taxon D

stem Jeafiness
lower stem surface
es
upper stem surface
processes
peduncles & bract surface
es

petiole length (mm)
peduncle length (mm)

bract apex

mostly basal or occasionally
extending midway up stem

glabrous to mlocletately hie
cent w/fe ew-nt
ands

moderately pubescent ie
umerous stipitate glan

numerous short stipitate
ands

(3) 12-45
(2.5) 8-55(65)

obtuse, acute

Taxon B
both basal & cauline,
extending up stem 15-60% o
stem height

lightly to heavily eae
w/few globules of sur

exudate

lightly to heavily pubescent
w/no-few stipitate glands

to moderate pubescence
ae a sessile or short stipi-
tate glands; occasionally
glabrate
(7) 13-40
11-140

caudate-attenuate or cuspidate
(rarely acuminate)

both basal & cauline, leaves
eg 20-60% of stem

lightly to moderately pubes
cent w/stipitate glandulos ave
increasing markedly w/height

glabrous to moderately pubes-
cent w/numerous stipitate
glands

numerous stipitate glands

16-60
6-140

obtuse, acute

mostly ne to apex or rarely
only m

glabrous to heavily Sia
short stipitat

oe to moderately pubes-
w/no-few sessile or short
ae glands

lightly to heavily pubescent
w/fcw-numerous stipitate
glands

8-35

8-55(75)

obtuse, acute, acuminate

number of disc florets 41-83 40-60 40-75 25-58

disc fleret throat 2.3-3.8 1.4-3.5 1.5-4.0 2.0-2.8
length (mm)

achene length (mm) 10-5.1 2.0-6.2 3.5-6.0(8.0) 3.0-4.6
pappus scale length (mm) (0.6) 1.0-3.2 0.7-3.2 0.7-3.7 1.0-2.4(2.8)

cIl

116

The second beneficial result of using case studies is the positive carry-
over to other science courses, and especially to those in other areas of
biology. The students gain a better appreciation of the utility of any man-
made classification for both animate and inanimate objects. They also are
much better prepared to make taxonomic decisions of their own or criti-
cally evaluate those of professional taxonomists in other courses that re-
quire such efforts (c.g., Introductory Microbiology, Survey of the Animal
Kingdom). The students also develop increased proficiency in evaluating
methods of data collection, synthesis, and display, which has strong carry-
over to all other areas of science.

The third beneficial result of using case studies is the improved intel-
lectual self-confidence that is generated within most students. Occasionally,
of course, a student will remain confused and baffled throughout the entire
course and will never really understand how to work the case studies; but
this occurrence is rare. In general, the students gain much self-confidence
in making taxonomic decisions as evidenced by their increasingly percep-
tive comments throughout the course. Perhaps even more important is that
the students gain self-confidence in presenting and defending their own
viewpoints before their peers. Sometimes, especially early in the term,
the resultant student classifications are widely divergent, and this leads
to lively and constructive discussion.

CONCLUSION

From my experience in using case studies in plant systematics courses,
I firmly believe that they are an extremely valuable part of the curriculum.
The case studies can be used at the undergraduate or graduate levels, with
the complexity of the problems adjusted according to the backgrounds and
abilities of the students involved. It may have been noticed that the previous
section of the paper stressed only beneficial results of using case studies;
this was done deliberately, because I believe no detrimental effects accrue,
except for the loss of time that alternatively could have been spent on
other topics.

Because it involves considerable efforts to extract and rearrange the
data from original papers for class use as case studies, it might be worth
mentioning that I have assembled a collection of case study examples and
exercises for publication in the near future. Along with these case studies
have been written several chapters that hopefully will provide at least an
introduction to the concepts that are necessary for working effectively the
exercises themselves.

ACKNOWLEDGMENTS
Sincerest thanks go to the numerous students who have offered criticisms

in the development of the case study concept presented in this paper.
Thanks are also due Dr. E. D. Rudolph for reading the final manuscript.

117

Costs associated with manuscript preparation and publication have been
borne in part by NSF Grant GB-37678. Permission to reproduce figures
and tables and to quote portions of the text in the case study used here
from Brittonia has been granted generously by The New York Botanical
Garden.

ANNOTATED LIST OF VASCULAR PLANTS
IN MOBILE, ALABAMA

MICHEL G. LELONG
Department of Biological Sciences, University of South Alabama
Mobile, Alabama, 36688

The need for a reasonably complete and recent list of plants in the
Mobile area has been felt in the teaching of some of our biology courses
and in other activities. Therefore a floristic survey of the University of
South Alabama property and of the adjacent Municipal Park area located
in the northwestern part of Mobile, was conducted from 1968 to the end
of 1970

The study area covers approximately three sections of land, and con-
sists primarily of rolling, sandy, Longleaf pinewoods typical of the region.
It is traversed by Three Mile Creek and adjacent swamp forests and wet
savannahs. The elevation of this area ranges from about 80 feet to 220
feet. The upland pinewoods is dominated by longleaf pine, and a diversity
of oaks such as laurel oak (Quercus hemispherica), southern red oak
(Q. falcata), turkey oak (Q. laevis), and sand post oak (Q. margaretta).
The understory includes ericaceous shrubs such as the blueberries (Vac-
cinium arboreum, V. elliottii, and V. darrowii), yaupon (Ilex vomitoria),
winged sumac (Rhus copallina), flowering dogwood (Cornus florida), and
numerous vines such as the green brier (Smilax auriculata) and yellow
jessamine (Gelsemium sempervirens). The ground layer in this pinewoods
consists of a great variety of herbs belonging primarily to the grass, the
legume, and the aster families. Representatives of these families are
Andropogon tener, A. scoparius (little bluestem), the dropseed grass
(Sporobolus junceus), the love grasses (Eragrostis spectabilis and E. refrac-
ta), sand lupine (Lupinus diffusus), the rattleboxes (Crotalaria angulata and
C. purshii), the beggars ticks (Desmodium laevigatum and D. viridiflorum),
Rhynchosia galactioides, the sunflower (Helianthus radula), deer tongue
(Trilissa odoratissima), the goldenrod (Solidago odora), the ironweed (Ver-
nonia angustifolia) and Coreopsis major. Herbs belonging to other families
occurring commonly in the lonfleaf pinewoods include scarlet basil (Cala-
mintha coccinea), the sage (Salvia azurea), Queen’s delight (Stillingia syl-
vatica), Hibiscus aculeatus, Verbena carnea, flowering spurge (Huphorbia
corollata) and spurge nettle (Cnidoscolus stimulosus). The swamp forest bor-
dering Three Mile Creek is dominated by sweet bay (Magnolia virginiana),
tupelo gum (Nyssa sylvatica var. biflora), red maple (Acer rubrum) and tu-
lip tree (Liriodendron tulipifera). The understory consists of a few shrubs
such as Virginia willow (Itea virginica), star-anise (Illicium floridanum),
Leucothoe axillaris, and strawberry bush (Euonymus americanus), The

SIDA P(2oe 118146... 1977,

119

ground layer in the Bay Forest is scant consisting largely of shade-tolerant
plants such as netted chain-fern (Woodwardia areolata), Gratiola virginiana,
and Lindernia dubia.

The transition zone between the Longleaf pine forest and the Bay Forest
is occupied by a rather narrow strip of wet savannah including a great
diversity of plants. Common shrubs in this area are gallberry (Ilex glabra),
wax myrtle (Myrica cerifera), black titi (Cliftonia monophylla) and sweet
pepperbush (Clethra alnifolia). The herb layer is very dense and varied
including numerous sedges such as beak rushes (Rhynchospora spp.), sun-
dews (Drosera spp.), meadow beauties (Rhevia alifanus, R. lutea, R. mari-
ana), milkworts (Polygala lutea, P. ramosa), violets (Viola primulifolia,
V. lanceolata), the orchids (Calopogon pulchellus and Spiranthes praecox),
and numerous panic grasses such as Panicum ciliatum, P. ensifolium, P.
scabriusculum and P. strigosum.

Other plant associations represented to a lesser extent in the study area
include the pond margins and the floating and submerged plants at Mobile
Municipal Park, as well as the fast-growing weedy habitats of roadsides,
clearings, lawns and pathways.

The climate of Mobile is subtropical and greatly influenced by the Gulf
of Mexico. Based on a forty year record (U.S. Dept. Agric, 1941), the aver-
age July temperature is about 82°F: and the average January temperature
is about 53°F. The average growing season lasts 298 days without killing
frost from February 17 to December 12. The average annual precipitation
is about 60 in. with 33 in. falling during the warm season,

Before the turn of our century, few botanists had investigated the flora
of southern Alabama. In 1857, Charles Mohr established himself in the
drug business in Mobile where he remained until 1900, shortly before his
death. During this time, he collected plants extensively throughout. the
State and particularly in the Mobile Bay region. His efforts culminated in
the publication of his book on the plant life of Alabama in 1901. Earlier,
Hezekiah Gates, a pharmacist in Mobile, collected plants of the coast region
from 1836 to 1842.

During our century, much interest has been shown in the flora of the
islands separating the Mississippi Sound and the Gulf of Mexico. F. E.
Lloyd and S. M. Tracy (1901) surveyed the insular flora of adjacent
Mississippi and Louisiana. W. T. Pennfound and M. E. O’Neill (1934)
studied the vegetation of Cat Island. L. J. Pessin and T. D. Burleigh (1941)
recorded observations on the forest biology of Horn Island, and later E.
Avery Richmond (1962, 1968) compiled a list of the plants on this island.
G. J. Miller and S. B. Jones Jr. (1967) reported on the vascular flora of
Ship Island, Mississippi. Rebecca Deramus (1970) conducted a floristic sur-
vey of vascular plants on Dauphin Island, in the extreme south of Mobile
County.

We should also briefly note some of the numerous contributions of Roland

120

M. Harper to a better general knowledge of the plants of our region,
especially his catalogue of the trees, shrubs and vines of Alabama (1928).
Recently Ross C. Clark (1971) greatly expanded our understanding of the
distribution of woody plants in the State.

During our floristic survey, approximately 3500 specimens were collected
and identified. One set of specimens is kept in the Natural History Collec-
tion of the University of South Alabama. Another set is being sent to the
University of North Carolina Herbarium. This sampling represents 876
species and varieties, 414 genera included in 125 families of vascular plants.

The grass family (Poaceae) is best represented in the study area with
135 species and varieties, followed by the composite family (Asteraceae)
with 110 species and varieties, the sedge family (Cyperaceae) with
species, and the family Fabaceae with 65 species.

In the following checklist, the nomenclature and taxonomy follow pri-
marily that of Radford, Ahles and Bell (1968). Small (1933) and other major
plant manuals were also consulted as well as recent treatments of plant
groups well represented in our area such as those by Kral dealing with
Xyris (1966) and with Bulbostylis and Fimbristylis (1971).

The synonymy is that of Mohr (1901), the latest comprehensive book
treating the flora of our region, To render the list more useful, a brief
statement of habitat and of the frequency of occurence is included.

One hundred and twenty-five species and varieties included in the follow-
ing list were not reported in Mohr’s catalogue. The names of these plants
are preceded by a single asterisk. Twenty-eight additional species and
varieties on this list were reported by Mohr to occur in the State outside
of Mobile County. The names of these plants are preceded by a double
asterisk. Plants which were not reported by Mohr fall largely into two
major categories: 1) species described after 1901 and closely related to
plants reported by Mohr; their recognition may merely reflect a “narrower”
species interpretation than that of Mohr; 2) introductions or ‘‘aggressive”
weeds which have recently spread to our region such as Stachys floridana
and Phyllanthus urinaria. Naturalized plant species are indicated by ‘‘N”
at the end of the entry pertaining to them in the following list.

I would like to express my appreciation to Dr. Sidney McDaniel for
annotating a number of ‘‘troublesome’’ specimens, especially those belong-
ing to the genus Vaccinium. I am grateful to Dr. Albert E. Radford for
reading the preliminary list and suggesting improvement in its presenta-
tion. I am also grateful to Dr. Robert Kral for making numerous helpful
suggestions for improvement of the original manuscript. I am thankful to
Dr. Grady Webster for verifying the identity of Phyllanthus urinaria and to
Dr. Murray Evans for verifying the determination of Thelypteris torresiana
and T. dentata.

I am indebted to Dr. Joab Thomas of the University of Alabama, to Dr.
Joseph Ewan of Tulane University, and to Dr. J. Ray Watson of Mississippi

121

State University for allowing me to consult the herbaria of which they
have charge. Dr. Ray Watson and Dr. Ken Rogers also kindly allowed me
to benefit generally from their knowledge of our Southeastern flora.

Finally I am thankful to the research committee of the University of
South Alabama for supporting this project.

ANNOTATED LIST

LYCOPODIACEAE
LYCOPODIUM ALOPECUROIDES L. Foxtail Clubmoss. Common in wet sandy pine-
lands.
LYCOPODIUM ADPRESSUM (Chapm.) Lloyd & Unders
requent in wet sandy pinelands.
LYCOPODIUM CAROLINIANUM L. Slender Clubmoss. Rare in wet sa andy pinelands.
LYCOPODIUM CERNUUM L. Nodding Clubmoss. Rare in moist, open sandy area n

swamp,

. Southern Bog Clubmoss. In-

ear

OPHIOGLOSSACEA
OPHILOGLOSSUM CROTALOPHOROIDES Walt.

and inconspicuous in lawns and meadov

Tuberous Adders-tongue. Infrequent

OSMUNDACEAE
ie eae CINNAMOMEA L. Cinnamon Fern
of swamp forest.
OSMUNDA REGALIS L. Royal Fern.

. Common in moist pinelands and along
Infrequent in moist open areas.
SCHIZAEACEAE

*LYGODIUM ae leases (Thunb.) Sw. Japanese Climbing Fern

woods, ditch

. Infrequent in moist

PTERIDACEAE
PTERIDIUM AQUILINUM (L.) Kuhn. Bracken Fern.
woods.

Common in dry, sandy open

ASPIDIACEAE
ON ASPLENIOIDES (Michx.) A.A

. Eat. Southern Lady Fern. Infrequent in

amp fore
ONOCLEA SENSIBI LIS L. Sensitive Infrequent along margins of swamp.
*“THELYPTERIS DENTATA (Forsk.) Chr. Rare in moist pinelands.

THELYPTERIS NORMALIS Cae Moxley.
Common along margin . swamp forest
T = LYPTERIS PALUSTRIS Schott. M aah Fern.
requent in wet savannahs,

a | ee LYPTERIS TORRESIANA Alston. Infrequent it

oe patens (SW.) Kuntze]

[Dryopteris thelypteris (L.) Gray]

1 swamp. forest.
BLECHNACEAE

W Ca Cee AREOLATA (L.) Moore. Netted Chain-fern. Common in swamp

WOODWARDIA VIRGINICA (L.) J. E. Smit

1. Common in wet savannahs.
ASPLENIACEAE
ASPLENIUM PLATYNEURON (L.) Oakes. Ebony Spleenwort.
sandy woods.

Infrequent in mesic,

POLYPODIACEAE
oe POLYPODIOIDES (L.) WATT.
ee

Resurrection Fern. Infrequent on
large

122

PINACEAE
PINUS ELLIOTTH Engelm, Slash Pine. Infrequent in wet pinelands.
PINUS PAL pane Mill. Longleaf Pine. Common in sandy upland woods and along

margin of swamp forest.

PINUS TAE DA L. Loblolly Pine. Infrequent in open, sandy areas.

CUPRESSACEAE

CHAMAECYPARIS THYOIDES (L.) BSP. White Cedar. Infrequent in swamp forest.
JUNIPERUS VIRGININIANA L. Red Cedar. Infrequent in dry, sandy soil.
TYPHACEAE
TYPHA LATIFOLIA L. Common Cat-tail. Infrequent in wet areas and ditches.
SPARGANIACEAE
SPARGANIUM AMERICANUM Nutt. [S. androcladum Gee ha Morong] Bur-
reed. Infrequent in opening within swamp forest and along stre
POTAMOGETONACEAE
POTAMOGETON DIVERSIFOLIUS Raf. Pondweed. Infrequent, floating in’ stream.
ALISMATACEAE
“© SAGITTARIA ENGELMANNIANA J.G. Sm. Arrowhead. Rare on edges of
opens FALCATA Pursh. [S. era falcata (Pursh) J.G. Smith].

on margins of nd.
SAGITTA ARIA LATIFOLIA Willd. Wapato. Common in wet savannahs and along pond

pond.
ia ccadat

margins.
HY DROCHARITACEAE
*EGERIA DENSA Planch. Elodea. Infrequent on sandy stream bank,
POACEAE
AGROSTIS ELLIOTTIANA Schultes. Silvery hair grass. Infrequent in meadow
AG ae HYEMALIS (Walt.) BSP. Bent Grass. Common on roadsides, ae a and
AGROSTIS- PERENNANS (Walt.) Tuckerm. [A. intermedia Scribner] Autumn Bent
srass. Common in moist sand along stream.
*AIRA ELEGANS Willd. ex Gaudin. Hairgrass. Rare
ALOPECURUS C or aa Walt. [A. geniculatus L.]

»

ong roadside.
Foxtail Grass. Rare in

moist lawns and_ fields.
ANDROPOGON GERARDI Vitman.
pee ON GLOMERATUS (Walt.) BSP. Bushy beardgrass.

Big Bluestem. Infrequent in dry, open woods.

Infrequent in moist
open a

ANDROPOGON SCOPARIUS Michx. Little Bluestem. Common in

*ANDROPOGON SUBTENUIS Nas . Beardgrass. Common on dry, sandy soi

ANDROPOGON TENER (Nees) Kunth. Common in dry, open woods.

hr aaa TERNARIUS ee [A. argyraeus Schult.] Infrequent in dry, sandy

dry, open woods,
il.

oil.
ANDROPOG ON VIRGINICUS L. [A. capillipes Nash] Broom Sedge. Infrequent in dry,
open ods.

ANTHAE "NANTIA VILLOSA (Michx.) Beauv. Common in dry woods.

ee INTERMEDIA. Scribn. and Ball. Three Awn Grass. Infrequent in dry,

dy woods.
A ARISTIDA. LANOSA Muhl. ex EIL [A
ARISTIDA LONGESPICA Poir. [A. gracilis EI. frequent in dry,
ARISTIDA MOHRIL Nash. Infrequent in dry, sandy woods. |
ARISTIDA PURPURASCENS Poir. Common in dry, open
*ARISTIDA VIRGATA Trin. [close to A. palustris (Chap.) Vasey in Mohr] Infrequent
in moist Open areas.

ARUNDINARIA TECTA (Walt.) Muhl. Cane. Infrequent a in low woods.
*ARUNDO DONAX L. Giant Reed. Rare on roadside, escaped. N.

. lanata Poir.] Common in dry, sandy woods.
ih ] In open sites.

123

AVENA SATIVA L. Oats. Rare in waste places, N.
ee ee ao agi ea compressum (Sw.) Nees] Carpet Grass. Infre-

uent in low, open areas, cultiv
*BRACHIARIA PL A'TYPHLYLLA (Gri) Nash. Infrequent in open, a areas.
BRIZA MINOR L. Quaking Grass. Rare oist, open disturbed are

BROMUS be enero — - ee (Willd.) H.B.K.] one Cae Infre-
quent b

*BROMUS COMMUTATUS ied [B. racemosus L.] Rare in dry waste places. N.

“BROMUS TECTORUM L. Rare on dry roadside.

CENCHRUS ECHINATUS L. Sandspur. Common on dry roadsides and waste places.

“CENCHRUS GRACILLIMUS Nash. Infrequent on dry roadsides

CENCHRUS PAUCIFLORUS Benth. [C. incertus M.A. Curtis] Ceimed on dry sandy

soil.

eee LAXA (L.) Yates. [Uniola laxa (L.) B.S.P.] Infrequent in mesic
woo

CHASMANTHIUM oo (Poir) Yates. [Uniola longifolia Scribner] Infre-
a in woodland border

— PETRAEA Sw. c. swartziana Doell] Finger Grass. Rare on dry, sandy dis-

= : tes

CY NODON DACTYLON Gs ) Pers. [C eee dactylon (L.) Kuntze] Bermuda Grass.
Common on roadsides te places, lawns

DACTYLIS GLOME RATA. L. Orchard oan cer re in cultivated Soe N.

DACTYLOCTENIUM AEGYPTIUM (L.) Beauv. Crowfoot Grass. Common on road-

N.

DANTHONIA SERICEA Nutt. Oat Grass. Common in dry, open woods.
ee ee ISCHAE MUM (Schreber) Schreber ex a thl. oe yerisma lineare (Krock.)

ash] Smooth rass. ea in’ disturbe
DIG ITARIA PILIFORMIS (L.) Koel. y — I Pi (Wale, ) Fern. [Syntherisma villo-
n Walt.] Crab Grass. Common in i andy areas,
DIGITARIA SANGUINALIS (L. . es [Syncherivn sanguinale (L.) Nash] Common
ass. Common on_roadsi waste places, cultivated ground.

rab
pIGIT ARIA SEROTINA a. che: fe eee file Walt.] Rare lawn
ved.
ECHINOCHLOA COLONUM (L.) Link. [Panicum colonum L.] Jungle Grass. Rare
N.

side
ELEUSINE INDICA (L.) Gaertn. Goose Grass. Common on roadsides, waste places
el pears CURVULA (Schrad.) Nees. Love Grass. Infrequent in dry, onde open

FRAGROST 5 ris IT S. Wats. [E. nitida (Ell) Chap.] Infrequent on dry road-
sides and w plac

FRAGROSTIS. HIRSUTA (Michx.) Nees. Commnen in dry open wees roadsides.

ERAGROSTIS OXYLEPIS aes Torr. [E. secundiflora Presl.] Rare on dry roadsides.

BLAChOstE REFRACTA (Muhl.) Scribn. Gan lon in moist pine can

ERAGROSTIS Pee (Pursh) Steud. [probably corresponds to E. pectinacea

(Michx.) Steud. Mohr] Common in dry open woods, roadsides.
al & REMOCE ae OPHIUROIDIS (Munro) Hack. Centipede Grass. Infrequent in dry,
places. N.
FRIANTHUS ALOPECUROIDES (L.) El. Plume Grass. Common in woodland borders,
nahs

PRIANTHUS, GIGANTEUS (Walt.) Muhl. [E. saccharoides Michx.] Infrequent in
moist nahs.

“FESTUCA ARUNDINACEA Schreb. Reed Fescue. Rare on roadside

i‘ UCA MYUROS L. Fescue. Infrequent on cultivated and waste erciads

SR OCTOFLORA Walt. Infre equent on cultivated and waste grounds.

GYMNOPOGON AMBIGUUS (Michx.) BSP. Windmill Grass. ommnon in dry woods.

GYMNOPOGON BREVIFOLIUS Trin. Common in dry woo

HACKELOCHLOA GRANULARIS (L.) Kuntze. Rare in open, disturbed area. N.

HORDEUM PUSILLUM Nutt. Little Barley. Infrequent on waste grounds

HYDROCHLOA CAROLINIENSIS Beauv. [H. fluitans (Michx.) ashi Infrequent
aquatic.

124

*IMPE RATA BRASILIENSIS Trin. Infrequent on open marshy area

N.
LEERSIA HEXANDRA Swartz. [Homalocenchrus hexandrus (Sw.) eee Cut Grass.
Infrequent in — shaded areas.
LEERSIA ORYZO § (L.) Swartz. [Homalocenchrus oryzoides (L.) Pollich] Common
on margins of me
LEERSIA VIRGI NICA Willd. [Homalocenchrus virginicus (Willd.) Britton] Infrequent

in moist sha

ed :
*LEPTOCHLOA UNINERVIA (Presl.) Hitchc. Sprangletop. Rare along dry, sandy

roadside

*LEPTOLOMA COGNATUM — Chase. [Panicum cognatum Schult.] Witch-
s

Grass. Rare on dry, open sandy

*LOLIUM MULTIFLORUM Lam. Rye G Grass. Common on waste places, lawns

LUZIOLA BAHIENSIS (Steud.) Hitche. [L. alabamensis Chapm.] Rare on errr
f swamp and along stream. N
MUHLENBE RGIA EXPANSA (DC) Trin. [M. trichopodes (Ell.) Chap.] Muhl.

Common in dry, open pineland

**MUHL ENB ERS SCHREBERI LE. Gmel. [M. diffusa Schreb.] Nimblewill. Rare in

shady waste places
a LISMENUS SE TARIUS (Lam.) Roem. & Schult. Rare in mesic woods.
ANICUM ACICULARE Desv. ex Poir. [P. arenicola Ashe] Panic Grass. Common on

o
i]
Q
an

open, sandy woods, rc des.
PANICUM AGROSTOIDES Spreng. [Includes P. elongatum ramosior Mohr] Common
in low woods, along margins * Baer and streams.
Cc

PANICUM aE § Michx. var. ANCEPS. [Includes P. rostratum Muhl.] Common

sic
*PANICUM "ANC SEPS var. RHIZOMATUM. (Hitche. & Chase) Fern. Infrequent on

roadsides,, moist pinelatr
PANICUM ANGUST IFO UM Ell. Common in dry, open woods.
PANICUM BARBUL TUM Mic hx. Infreque nt on sa da stream banks.
fot ; -

pecans DICHOTOMIFLORUM ich [P. sualbecuin Lam] Common in low
ods, waste place
PANICUM DICHOTOMUM L. [Includes P. ludicum Ashe] Infrequent in low pinelands,
stream banks.
“PANIC UM ENSIFOLIUM Baldw. ex EIl. Infrequent in moist, open a
PANICUM HIANS Ell. [P. melicarium Michx.] Infrequent along margins of stream

PANICUM LANCEA \RIUM Trin. [P. nashianum Scribne r | Rare in low sandy woods.
PANICUM LANUGINOSUM Ell. (sensu lato) Common in ees open woods, roadsides.
— ia LAXIFLORUM Lam. [Includes P. pyriforme Nash] Infrequent in mesic

«PANIC UM pees IMERI Nash. [Possibly corresponds to P. paucipilum Nash.] Infre-
sen woods, roadsides.
PANICUM LONG IFOLIUM ‘Torrey. gon on moist open streambank.
PANICUM OLIGOSANTHES Schule. Infre equent in dry, open woods.
*PANICUM Spooner Nash. Rare in div. open areas.
*PANICUM RAMOSUM L. Rare on roadside. N.
*PANICUM RAVENELII 2. & Merr. Common in dry pinelands.
PANICUM REPENS L. Torpedo Grass. Common along margins of ponds.
cents SCABRIUSCULUM Ell. Common i
onds.
PANICUM SCOPARIUM Lam. [Includes also P. pubescens Lam. and P. viscidum FIL]
Infrequent in moist open areas.
PANICUM SPHAEROCARPON Ell. Common in dry, open woods, roadsides.
PANICUM STRIGOSUM Muhl. [P. longipedunculatum Scribner] Rare in moist open
areas.

1 Wet sav annahs, ditches, margins of

125

a anes oe [Includes P. trifolium Nash and P. albomarginatum Nash]
Cor dry urbed eee and woods

PANICUM eee Muhl. Common in borders of swamps = wet ditches.

PANICUM VILLOSISSIMUM Nash. [Includes also P. haemacarpon Ashe, P. xantho-
spermum Scribner and Mohr, and P. pseudopubescens Nash] on in dry, open

Ww ands,
PANICUM VIRGATUM L. Switchgrass. Infrequent in low open woods.
era a BIFIDUM (Bertol.) 1 Nash. Infrequent in sandy pine and oak woods.
ASPALUM CONJUGATUM Mae Rare lawn weed.
PASPALUM DEBILE ye [Included by Mohr under P. longipedunculatum Le Conte]
nfre en, disturbed area.
ayy! DIFFOR ME Te Conte. Rare in moist, sandy soil.
PASPALUM DILATATUM Poir. Dallis Grass. Common on roadsides, waste places. N.
PASPALUM FLORIDANUM Michx. Common on roadsides, waste places.
PASPALUM LAEVE Michx. Infrequent in ope ae waste grounds.
*PASPALUM NOTATUM Flugge. Bahia Grass. mon on omni waste places.
woods.

alt. in. low ee
PASPALUM SE TACE UM Michx. [P. viliatifolium Michx.] Common in dry, open woods,
roadsides, waste ground.
PASPALUM SUPINUM Bose. [P. ciliatifolium dasyphyllum (EIL) Chap.] Infrequent in

dry, open grou coe

*PASPALUM URVILLEI Steud. Vasey Grass. Common on roadsides, waste ground. N.
PHALARIS CAROL iM ANA Walt. Canary Grass. Infrequent in waste places.
POA ANNUA L. Blue ss. Common. on - wns waste places. N.

petit ieee MONSPE ae NSIS (L.) Desf. Rabbit-foot Grass. Rare in marshy area
along stream. N.
SACCIOLEPIS STRIATA (L.) Nash. [Panicum gibbum EIl.] Common along border of

st ponds.

SECALE ec aes = Rye. Rare along roadside. N

SETARIA GENICULATA (Lam.) Beauy. Fenaeesahien imberbis eis ae and
Chaetochloa oe (H.B.K.) Scribn. and Merr.] Bristlegrass. in places.

SORGHASTRUM ELLIOTTIL (Mohr) Nash. [Chrysopogon elliottii Mohs] Indian
Grass. Common in dry, open woods.

ee NUTANS (L.) Nash. [Chrysopogon nutans linneanus Doell] In-

dry, open woods.
ion ae HALE PENSE (L.) Pers. Johnson Grass. Common on roadsides, waste places.
N.
SPHE aoe IS FILIFORMIS (Chapm.) Seribn. [Eatonia filiformis (Chap.) Vasey]
We Gr Common in dry, open woods
SPHE NOPHOLIS OBTUSATA (Michx.) Serb . [| Eatonia obtusata (Michx.) Gray var.

Repens (Walt.) A.H. Moore} Common on moist roadsides, waste places.
he nae CLANDESTINUS (Bieler) Hitche. ee ed. Rare in dry, open woods.
SPOROBOLUS JUNCEUS (Michx.) Kunth. Common in dry, open woods.
SPOROBOL ie POIRETTL (Roem. & Schult.) Hitche. [probably aes in S. indicus
ds. N.

r.] Smut Grass, Se ae on roadsides, cultiv ee and waste gr ee
ee cor INIANUS (Steud.) Henr. Commo ry, open
pet FLAVUS (L.) Hitche. var. CHAPMANII i shinee ne velingia chap-
manii Small] Purple Top. Common in woodland margins, roadsides.
meer ee RICANA Beauv. rlesliaers americana (Beauy.) Kuntze] Rare in

dry, sandy area.
“TRITICUM. AESTIVUM L. Wheat. Rare on waste ground. N.

CYPERACEAE
BULBOSTYLIS Hennes Age Clarke. Infrequent in moist, sanc
BULBOSTYLIS CILIATIF (ELL) Fern. [Stenophyllus eae ‘(EIL) Mohr]
Common in dry, sa ae areas.
CAREX ALBOLUTESCENS Schwein. Sedge. Common in low, moist. areas.
**CAREX AMPHIBOLA Steud. Infrequent in low, swampy woods

126

CAREX ATLANTICA neces Infrequent in swamp for
CAREX COMPLANATA T & Hook. [C. hirsuta Willd. ] Infrequent in low woods.
CAREX DEBILIS Michx. eee in low eroade.
CAREX ELLIOTTII ar ies and Torr. Infrequent in low woods.
CAREX FESTUCACEA vkuhr. Infrequent in low woods.
CAREX corre 1. var. AUSTRALIS Bailey. nae aces in low woods.
CAREX GLAUCESCENS Ell. Infrequent in wet sayannahs.
CAREX INTUMESCENS Rudge. Infrequent in open, wet areas.
CAREX LEAVENWORTHIL Dewey. Infrequent along shady roadside.
Sere LEPTALEA Wahl. var. HARPERI (Fern.) Stone. Infrequent in low woods.
AREX LURIDA Wahl. Common in open wet areas.
Cc,

es X VENUSTA Dev [C. oblita Steud.] Infrequent in low woods.

— PERUS COMPRE SSUS ne Common in moist, disturbed places.
YPERUS CUSPI ree cae Rare weed on campus lawn.

. PERUS ESC Nut ae Common on roadsides, a

ULENTUS
CYPERUS FILICULMIS a Infrequent in lawns, cultivated S.
CYPERUS GLOBULOSUS Aublet. [C. echinatus (EII.) Wood. | nee in sandy
wasteplaces.

CYPERUS oo L. Infrequent in moist places.

3
ry

CYPERUS IRIA re in wet, open areas,
CYPERUS open US [C. speciosus Vahl. } Common in wet a. areas, ditche
ee ey POLYS ae Rottb. var. TEXENSIS (Torr.) Fe [C. ee cats

Torrey] Infrequent in low, disturbed places.
=“ ‘CYPE RUS PSEUDOVEGETUS Steud. Infrequent in wet areas.
CYPERUS RETROFRACTUS Engelm. ex Boeck. Infrequent in dry woods.
PERUS RE t RORSUS Chapm. [C. cylindricus (EIl.) Britton] Common in cultivated

punds, ands.
CY PERUS ROTUNDUS L. Nut Grass. Common in moist, cultivated grounds. N.
CYPERUS SESQUIFLORUS (Torr.) Mattf. & Kukenth. [Kyllinga sesquiflora ‘Torr.]
Common in ditches, moist lawns.
CYPERUS RIGOSUS L. Common in low waste plac
CYPERUS TENUIFOLIUS (Stued.) Dandy. [Killinga unis Michx.] Infrequent
low fields
DULICHIUM ARUNDINACEUM (L.) Britt. Infrequent in swamp.
*ELEOCHARIS ne FOIDES (Ell) Torr. [Included by Mohr under EF. interstincta]

n

Spike-rush. Rare on border of stream.

ELEOCHARIS EL. AVE SCE ie aad Urb: [E. ochreata (Nees) Steud and E. olivacea
Torr nfrequent in wet relands, te of ponds and stream.

ELEOCHARIS MICROCARPA Torr. Infrequent along border of ponds and_ stre

ELEOCHARIS OBTUSA (Willd.) Schult. [E. ovata (Roth.) Roem. & Schult. ] ee
along border of pon and strean

oo ROBBINSII Oakes hate partly submerged along stream. bank.
ELEO ARIS TUBERCULOSA Uiichie, ) R. & S. Common in wet open areas and along
er ny ponds and stream.

Rana ae ANNUA ae, R. and S. Infrequent in moist grassy areas.

FIMBRISTYLIS AUTUMN ) R. an Common in moist, open areas
FIMBRISTYLIS pee. (L 2 age Infrequent in moist op rea
*FIMBRISTYLIS MILIACEA (L.) Common in moist, open areas. N

*FIMBRISTYLIS TOMENTOSA ae pea in moist, Open areas.

FUIRENA SCIRPOIDEA Michx. Rare along stream bank.

FUIRENA SQUARROSA Michx. Umbrella-grass. Common in wet open areas.
ane ARYA NITENS (Vahl) Wood. Bald Rush. Infrequent in moist open sandy

RHY NCHOSPORA CADUCA Ell. Beak-rush. Rare in open marshy area.
CHOSPORA CEPHALANTHA Gray. [R. axillaris (Lam.) Britton]
PRE : NCI TOSPORA CHALAROCEPHALA Fern. & Gale. Infrequent in open marshy

RHYNCHOSPORA CHAPMANIDE M.A. Curtis. Rare in open marshy area.
RHYNCHOSPORA CILIARIS (Michx.) Mohr. Infrequent in moist open areas.

127

RHYNCHOSPORA CORNICULATA (Lam.) Gray. Common in wet Ls n area
RHYNCHOSPORA FASCICULARIS (Michx.) Vahl. Infrequent in open vn hy areas
RHYNCHOSPORA GLOBULARIS (Chapm.) Small. [Rhynchospora cymosa (Willd. )

Ell. and R. cymosa globularis Chap.] Infrequent on margin of ponds.
RHYNCHOSPORA GLOMERATA (L.) Vahl. Common in open marshy area.
RHYNCHOSPORA GRACILENTA Gray. Infrequent in wet open areas.
RHYNCHOSPORA GRAYI Kunth. Uncommon on dry een oe
*RHYNCHOSPORA HARVEYI W. Boott. Infrequent in open marshy a
RHYNCHOSPORA INTERMIXTA E. - right. peeerncaes pusilla Chapm.] Rare in

open marshy areas.

RHYNCHOSPORA MACROSTACHYA Torrey. Infrequent along open margins of pond.

RHYNCHOSPORA MEGALOCARPA Gray. Infrequent in dry, sandy a elands.

RHYNCHOSPORA RARIFLORA (Michx.) Ell Uncommon a small stream.

SCIRPUS CYPERINUS (L.) Kunth. Bulrush. Common in moist. areas.

SCIRPUS ETUBERCULATUS (Steud.) ae [S. cylindricus “aS Britton] Rare
on margin of pond,

SCIRPUS KOILOLEPIS (Steud.) Gleason.
in wet, open areas.

SCLERIA CILIATA Michx. Nut- ‘ne Common in dry, open woods.

eee Cece aie ae Michx. var. pubescens Britton. [S. torreyana Walp.] Infre-

in moist pineland
SCLERIA RCrOM ERE TA Michx. Common in sandy pinelands, pond margins.

7

S. carinatus (Hook. and Arn.) Gray] Rare

—

ARECACEAE
SERENOA REPENS (Bartr.) Small. [S. serrulata (Michx.) Benth. and Hook.] Saw
Palmetto. Common in low pinelands.
ARACEAE
*COLOCASIA ESCULENTA Schott. ‘Varo. Rare in ditches near park. N.
ORONTIUM AQUATICUM L. Golden Club. Common along margins of stream and

n open moist area along

ponds.
PELTANDRA SAGITTAEFOLIA (Michx.) Morong. Rare
stream.
PELTANDRA VIRGINICA (L.) Kunth. Infrequent in swamps.
LEMNACEAE

*LEMNA PERPUSILLA Torrey. [possibly included by Mohr under L. Minor L.
along margins of pond.

Rare

MAY ACACEAE
MAYACA AUBLETTI Michx. Bog Moss. ng ees in wet, sandy open areas.
MAYACA FLUVIATILIS Aubl. [included by Mohr, perhaps judiciously under M.

aubletii] Infrequent in wet, sandy open areas.

XYRIDACEAE

XYRIS AMBIGUA Beyr. Yellow-eyed Grass. Common in moist’ pinelands.

XYRIS CAROLINIANA Walt. [X. flexuosa Muhl.] Common in moist pinelands.

*XYRIS DIFFORMIS Chapm. var. DIFFORMIS. Jipeobably included by Mohr in X.
communis Kunth.] Infrec ae on pond margit

XYRIS FIMBRIATA Ell. Rare in wet pine ee

XYRIS IRIDIFOLIA Chapm, Ta dies nt in openings in swamp.

XYRIS JUPICAL Rich. EX. communis Kunth.}] Common in moist pinelands pond mar-
gins.

ERIOCAULACEAE
ERIOCAULON DECANGULA L. Pipewort. Infrequent along pond mare
LACHNOCAULON ANCEPS a oraae Bog-buttons. Infrequent in moist, open
sandy areas.

128

BROMELIACEAE
TILLANDSIA USNEOIDES L. Spanish Moss. Infrequent epiphyte.

COMMELINACEAE
*ANEILEMA NUDIFLORUM (L.) Kunth. Infrequent in moist cultivated sites. N.
COMMELINA DIFFUSA eee [C. nudiflora L. ac ower. Infrequent in waste places.
COMMELINA ERECTA L. Common in waste pla
TRADESCANTIA HIRSUTICAULIS Small. Rare: in or woods.
TRADESCANTIA OHIENSIS Raf. [T. reflexa Raf. ] ae Infrequent in moist

roadsides, waste plac

JUNCACEAE
JUNCUS ACUMINATUS Michx. Rush. Common in moist open are
JUNCUS ‘epee EML [J. marginatus clue (Michx.) Coville} Infrequent in
moist, Oper
*JUNCUS CORIACEUS Mackenzie. Rare along stream and pond margins
JUNCUS oe Gray. [J. acuminatus debilis (Gray) Engelm.] eed in moist,
open area
JUNCUS DICHOTOMUS Ell. common in moist, see areas.
JUNCUS DIFFUSISSIMUS Buckl. Common in wet are alone stream and ponds.
JUNCUS EFFUSUS L. Infrequent in edges of swamp ne ponds.
JUNCUS ELLIOT Chapm. Rare in wet, open areas
JUNCUS GYMNOCARPUS Coville. Rare in wet, open areas.
JUNCUS MARGINATUS Rostk. Common in moist, sandy soil.
UNCUS POLYCEPHALUS Michx. Infrequent along stream and = ponds.
UNCUS REPENS Michx. Infrequent ane pond —

st a
JUNCUS TENUIS Willd. Common in waste and cultiv i. grounds
JUNCUS TRIGONOCARPUS Steud. Infrequent In moist, open areas.
*JUNCUS VALIDUS Coville. Infrequent in moist, open :

LILIACEAE
ALETRIS AUREA Rae Infrequent in moist. pinelan
ALETRIS FARINO . Unicorn-root. Common in . pinelands.
ALLIUM Seas re Kuntze. Pee poeta bivalve (L.) Britton] False Garlic.
Common in roadsides, cultivated and waste grounds.

ALLIUM CANADENSE L. Wild Onion. Pome on roadsides, waste grounds.
*ALLIUM INODORUM Aiton. Uncommon in dis ae areas at park. N.
AMIANTHIUM MUSCAETOXICUM (Walt.) Gray. [Chrosperma muscaetoxicum
(Walt.) Kuntze] Fly Poison. Common on wooded slopes.
CHAMAELIRIUM LUTEUM (L.) Gray. Blazing Star. Rare in mesic woods.
SMILAX AURICULATA Walt. Greenbrier, Catbrier. Common in dry, open woods.
SMILAX BONA-NOX L. Common in disturbed areas, roadsides.
SMILAX GLAUCA Walt. Common along border of swamp forest.
SMILAX LAURIFOLIA L. Common in swamp forest.
SMILAX PUMILA Walt. Common in dry, open woods.
Rare ist’ area i

*SMILAX ROTUNDIFOLIA in’ mo rea along creek.
SMILAX SMALLIL Morong. Is lanceolata L.] oe in mesic woods.
SMILAX WALTERI Pursh. Rare in moist areas along creek.

TOFIELDIA RACEMOSA pee t.) BSP. _ ‘Asphodel Infrequent in moist pinelands.
YUCCA ALOIFOLIA L. Spanish—Bayonet. Rare in woodland border.
YUCCA FILAMENTOSA L. Bear-grass. ne in dry, open woods.
DIOSCOREAECEAE
DIOSCOREA VILLOSA L. var. VILLOSA. Wild Yam. Rare along border of swamp
forest

AMARYLLIDACEAE
CRINUM BULBISPERUM Milne-Redhead & Schweickerdt. [C. americanum L.] Rare

along open stream ban {.

129

*“HYPOXIS MICRANTHA aa Yellow Star-grass. Common in dry pinewoods.

“HYPOXIS SESSILIS L. Infrequent in dry woods.

ZEPHYRANTHES ATAMASCO “13 Herb. [Atamosco atamasco (L.) Greene] Rain
Lily. Rare in roadside ditch.

HAEMODOR ACEAE
ee ee CAROLINIANA (Lam.) Dandy. [Gyrotheca capitata (Walt.) Mo-
ong] Redroot. Infrequent in moist pinelands.
IRIDACEAE
*IRIS FULVA Ker. Red Iris. Rare along creek.
ae HEXAGONA Walter. Blue Flag. ia age ae pond.
IRIS VERNA L. Dwarf Iris. Common in dry, woods,
Sr SRUCE ENG ARENICOLA Bichn. [S. ea Bicknell and S. carolinianum Bick-
nell] Blue-eyed Grass. Common in dry, sandy a woods
*SISYRINCHIUM EXILE Bicknell. Frequent lawn
a ee MUCRONATUM Michx. var. ATLANTICUM (Bicknell) Ahles. Com-
on in dry, sandy woo

mn
SISYRINCHIUM ROSUL AT UM Michx. Common in dry, sandy waste grounds, lawns.

BURMANNIACEAE
APTERIA APHYLLA (Nutt.) Barnhart. [A. setacea Nutt. ] Infrequent in low woods.
BURMANNIA CAPITATA (Gmelin) “Martius. Infrequent and inconspicuous in wet,
sandy areas along stream.

ORCHIDACEAE
CALOPOGON PULCHELLUS (Salisb.) R.Br. [Limodorum tuberosum L.] Grass Pink.
Common in moist pinelands.
CLEISTES DIVARICATA (L.) Ames. [Pogonia divaricata (L.) R. Br.] Rosebud Or-
chid. Rare in moist woodlands.
HABENARIA CILIARIS (L.) R. Br. Yellow Fringed Orchid. Infrequent in open,

marsh

HABENARIA REPENS Nutt. Water ae soca Rare on Rains of lake.
sare OPHIOGLOSSOIDES (L.) Ker-G ose Pogonia. Infrequent in wet
savannah.
SPIRANTHES GRACILIS (Bigel.) Beck var. FLORIDANA (Wherry) Correll. [Gyro-
achys gracilis (Bigel.) Bee ee in low pinelands.
SPIRANTHES PRAECOX (Wal Wa [Gyrostachys praecox (Walt.) Kuntze]

adies Tresses. ieee ae in moist pinelands.

s-leaved I
igen VERNALIS Engelm. and Gray. Spring Ladies Tresses. Rare in open,

SAURURACEAE
SAURURUS CERNUUS L. Lizard’s Tail. Rare along stream banks.
SALICACEAE
POPULUS DELTOIDES Marsh. Cottonwood. Infrequent along stream banks.
SALIX NIGRA Marsh. Black Willow. Common on stream banks and other moist areas.
MYRICACEAE
MYRICA CERIFERA L. Wax Myrtle. Common in moist pinelands.
MYRICA HETEROPHYLLA Raf. [M. carolinensis Mill.} Bayberry. Infrequent in moist
pinelands and along sie of bay est.
MYRICA INODORA Bartram. Rare along margin of bay forest.
JUGLANDACEAE
CARYA ILLINOENSIS (Wang.) K. Knock. [Hicoria pecan (Marsh.) Britton] Pecan.
Rare on roadsides, escaped.
CARYA TOMENTOSA Soba Nutt. [Hicoria alba (L.) Britton] Mockernut Hickory.
Common in dry, open woo

130

FAGACEAE
*CASTANEA ALNIFOLIA Nutt. var. FLORIDANA Sarg. Florida Chinquapin. Rare in
dry woods.
CASTANEA ae (L.) Mill. Chinquapin. Common in dry, sandy upland woods.
QUERCUS ALBA L. ys Oak. Rare in mesic woodland.

QUERCUS ese Michx. [Q. digitata (Marsh.) Sudworth] Southern Red Oak,
Spanish Oak. Conon in dry, open woods.
*QUERCUS a MINA’ Small. Dwarf Live Oak. Infrequent in dry, upland woods.

UERCUS HE enero ae Bartr. [Q. laurifolia Michx.] Laurel Oak. Common in
dry, open woods to swamp margins.
QUERCUS INCANA Bartr. [Q. brevifoli
in dry, sandy soil.
sk RCUS LAEVIS Walt. [Q. catesbaci Michx.] Turkey Oak. Common in dry, sandy

ia (Lam.) Sargent] Blue-Jack Oak. Common

*QUE RC US MARGARETTA Ashe. Scrubby Post Oak. Common in eit open woods.
QUERCUS NIGRA L. Water Oak. Common in bord ers of low

QUERCUS STELLATA Wang. Post Oak. Rare in mesic woods.

QUERCUS VIRGINIANA Mill. Live Oak. Infrequent in open woods.

MORACEAE
i won ae PAPYRIFERA (L.) Vent. Paper Mulberry. Rare, escaped near rail-
Res trac

TOUA V ILLOSA (Thunb.) Nakai. Rare weed in cultivated grounds. N.
on ALBA L. White Mulberry. Rare in cultivated and waste grounds. N
MORUS RUBRA L. Red Mulberry. Infrequent in low woods.

CANNABACEAE

BOEHMERIA CYLINDRICA (L.) Sw. False Nettle. Common in low woods along stream.

LORANTHACEAE
PHORADENDRON SEROTINUM (Raf.) M.C. Johnston. Mistletoe. Infrequent parasite

on deciduous trees.

POLYGONACEAE
POLYGONUM HYDROPIPEROIDE ’ Michx. Smart-weed. Common along stream banks.
POLYGONUM LAPATHIFOLIUM L. Infrequent along stream banks.
FORTEONEM. PE NSYLVANICUM : Infrequent in moist disturbed areas and along

stream banks.
POLYGONUM PUNCTATUM EIL Common in swamp fo
**POLYGONUM SETACEUM Baldw. ex Ell. Infrequent in swamp for
RUMEX CRISPUS L. Sour Dock. Infrequent on moist roadsides, open stream banks. N.
RUMEX HASTATULUS Balw. ex Ell. Wild Sorrel. Common on roadsides, waste places.

CHENOPODIACEAE
CHENOPODIUM ALBUM L. Pigweed. Rare in waste places.
CHENOPODIUM AMBROSIOIDES L.
N.

Mexican-tea. Infrequent in waste places, roadsides.

AMARANTHACEAE
ALTERNANTHERA PHILOXEROIDES (Mart.) Griseb. [Telanthera philoxeroides acu-
tifolia Moq.] Alligator-weed. Frequent along pond margins. N.
AMARANTHUS HYBRIDUS L. Pigweed. Rare on moist roadside.
AMARANTHUS SPINOSUS L. Thorny amaranth, ain pea along roadside. N.
*AMARANTHUS VIRIDIS L. Rare weed, in flower

NYCTAGINACEAE
oe JALAPA L. Four-o’clock. Rare in) wast ces. N.
PHYTOLACCA AMERICANA L. [P. decandra Ly coe Common 1

pes

a waste

131

AIZOACEAE
MOLLUGO VERTICILLATA L. Carpet-weed. Infrequent in cultivated and waste
grounds, N.
POR TULACACEAE
PORTULACA OLERACEA L. Rare in cultivated grounds. N.
CARYOPHYLLACEAE
** ARENARIA Beta pene L. Rare along roadside. N.
CE oe GLOMERATUM $ Thuillier. [C. viscosum L.] Mouse-Ear Chickweed.
lawns, waste ee sie along roadsides.
oa DECUMBENS (ElL.) T G. Pearlwort. Common on lawn, waste places.
SILENE ANTIRRHINA L. Sleepy ca Rare along dry, sandy roadsi
STELLARIA MEDIA (L.) Cyrill,
N.

grounds, waste places.

e.
[ Alsine media L.] Chickweed. ates in cultivated

=

NYMPHAEACEAE
NYMPHAEA ODORATA ar oe odorata (Dryand.) Woodv. & Wood] Water-
lily. Common in pond at
CABOMBACEAE
BRASENIA SCHREBERI oe [B. purpurea (Michx.) Casp.] Water Shield. Infrequent,

floating in pond at

RANUNCULACEAE

CLEMATIS CRISPA L. ather-Flower. a margins of swan

a SOR eaTe Levl. and = Vanio re in open, presen area by
eek. N.

R ANUNCUI US MURICATUS L. Buttercup. oe in ei wet meadows.

RANUNCULUS PUSILLUS Poir. Infrequent in moi cas
XANTHORHIZA SIMPLICISSIMA Marshall. enh apiifolia peers, Rare along
shaded stream banks.

MENISPERMACEAE
**COCCULUS CAROLINUS (L.) DC. [Cebatha carolina (L.) Britton] Coralbeads. Rare
1

in cultivated ground.

MAGNOLIACEAE
ILLICIUM FLORIDANUM Ellis. Star Anise. Common in bay forest as understory shrubs.
LIRIODENDRON TULIPIFERA L. Tulip Tree. Common in low woods.
Le eneray GRANDIFLORA L. [M. foctida (L.) Sargent] Southern Magnolia. In-
requent along border of swamp _ forest,

MAGNOLIA VIRGINIANA L. Sweet Bay. Common in swamp forest.

ANNONACEAE
ASIMINA PARVIFLORA (Michx.) Dunal. Dwarf Pawpaw. Common in sandy woods.

ALYCANTHACEAE
CALYCANTHUS ie L. [Butneria florida (L.) Dearney] Sweet Shrub. Rare
d

in mesic Woo lanc

LAURACEAE
*CINNAMONUM CAMPHORA (L.) Nees. and Eberm. Camphor-Tree. Infrequent along
bor of woods. N.
PE RSEA oe? USTRIS Sa Sarg. [P. pubescens (Pursh) Sargent] Red Bay. Common

and its margins.
SASSAFRAS ALBIDUM. ne Nees. [S. sassafras (L.) Karst.] Sassafras. Common

in rolling pinelands.

132

FUMARIACEAE
CORYDALIS MICRANTHA la: Gray ssp. AUSTRALIS (Chapman) Ownbey.
Infrequent along moist roadsi
BRASSICACEAE
BRASSICA NAPUS L. [B. campestris L.] Turnip. Rare in disturbed open areas. N.
CAPSELLA BURSA-PASTORIS (L.) Medic. [Bursa bursa- ae ag (L.) Britton] Shep-
herd’s purse. Common weed around dwellings and law
d kb

*CARDAMINE HIRSUTA L. [probably included by Mohr under C. pennsylvanica

cena eats paren (L.) Sm. a ent on ist roadsides, N.
EPIDIUM ICUM L. a, Giana? in eile vated a waste grounds.
. wera nh eaten UM Occasional along roadsides

RORIPPA WALTERI (EI) ie Yellow Cress. Infrec equent in wet meadows
SIBARA VIRGINICA (L.) Rollins [Arabis virginica (L.) Trelease] Infrequent in dis-
turbed areas.
DROSERACEAE
DROSERA CAPILLARIS Poir. Sundew. Common in moist pinelands.
DROSERA INTERMEDIA Hayne. Infrequent in moist pinelands, border of ponds.
DROSERA LEUCANTHA Shinners. [D. brevifolia Pursh] Common in moist pinelands,

order of ponds.

SAXIFRAGACEAE
DECUMARIA aaa Li oh oe Common in swamp forest.
ITEA VIRGINIC . Virginia Willow. Common in low woods and along stream.
LE pon on er scaavena even EN. Infrequent and inconspicuous in
damp, grassy areas.

HAMAMELIDACEAE
HAMAMELIS VIRGINIANA L. Witch-hazel. Comm in moist woods.
LIQUIDAMBAR STYRACIFLUA L. Sweet-gum. fede couene in mesic woods.

=
>

PLATANACEAE
PLATANUS OCCIDENTALIS L. Sycamore. Infrequent in moist sand along creek.

AGRIMONIA PUBESCENS Wallr. var. MICROCARPA Wallr. [A. pumila Muhl.] Agri-
mony. Rare in upland mixed woods.
AMELANCHIER ARBOREA (Michx. F.) Fern. June-berry. Infrequent near stream

banks.
CHRYSOBALANUS OBLONGIFOLIUS Michx. Gopher Apple. Common in dry, open
woods.
sche sii INDICA (Andr.) Focke. Indian “Strawberry. Infrequent on roadsides,
plac
M AL 'US "ANGUSTIFOLIA (Ait.) Michx. [Pyrus angustifolia Ait.] Crab Apple. Rare
along path in mesic woods.

PRUNUS ANGUST IFOLIA Marsh. Chickasaw Plum. Common on _ roadsides, woodland

PRUNUS CAROLINIANA Ait. Laurel Cherry. Rare on waste grounds, escaped.

PRUNUS PERSICA (L.) Batsch. Peach. Rare on roadsides, escaped. N.

oe SEROTINA Ehrh. Black Cherry. Common in woodlands, waste grounds.

*PYRACANTHA CRENATO-SERRATA Rehd. Rare, escaped in open waste grounds. N

Seen ers ATA Wendl. Macartney Rose. Rare in roadside thicket, escaped, N.
VIGATA Michx. Cherokee Rose. Infrequent along woodland margins, road-

4

ROSA Ore Thunb. Rose. Rare in waste places; former home sites. N.
*ROSA WICHURAIANA Crepin. Memorial Rose. Rare along railroad ee N.
*RUBUS SE TULIFOLIUS Small. [probably corresponds at least partly to R. argutus

Link in Mohr] Blackberry. Common on roadsides, and in moist. pinelands.

133

eee CUNEIFOLIUS Pursh. Infrequent on margins of ponds.
**RUBUS FLAGELLARIS Willd. Dewberry. Rare in open waste places.
RUBUS TRIVIALIS Michx. Common on dry roadsides, waste places.
SORBUS ARBUTIFOLIA (L.) Heynh. [Aronia arbutifolia (L.) Ell.] Chokeberry.

Common in wet savannahs,

FABACEAE
ALBIZIA JULIBRISSIN Durazz. Mimosa. Common in open woods, roadsides. N.
4 pe oo Medic, [Apios apios (L.) MacMillan] Gredednie: Infrequent

Pod CnnGe VILLOSUS Michx. Rare along dry sandy roadside.
CASSIA FASCICULATA Michx. [Chamaecrista fascicularis (Michx.) Greene] Partridge
Pea. Common on roadsides, woody places.
CASSIA NICTITANS L. [Chamaecrista nictitans (L.) Moench and Chamaecrista multi-
pinnata (Pollard) Greene] Wild Sensitive Plant. Common on roadsides, weedy places.
ides. N.

CASSIA OCCIDENTALIS L. Coffee Senna. Infrequent on roadsides.

CENTROSEMA VIRGINIANUM (L.) a [Bradburya virginiana (L.) Kuntze] But-
terfly Pea. Common in dry, open wo

CLITORIA MARIANA L. Common in ae open wood

CROTALARIA ale ATA Mill. [C. rotundifolia Wale.) Poir.] Rattlebox. Common
in dry, open woo

*CROTALARIA INTERMEDIA Kotschy. Rare on roadside. N.

CROTALARIA PURSHII DC. Infrequent in open woods.

**“CROTALARIA SAGITTALIS L. Rare in open woods,

*CROTALARIA SPECTABILIS Roth. Common on_ roadsides

i id ae PUNICEA (Car.) DC. Infrequent on bores of bay forest and along

DE sMANTHUS ae NSIS (Michx.) Mac. M. [Acuan illinoense (Michx.) Kuntze]

Rare in waste
DE SMODIUM CILIARE (Muhl. ex Willd.) DC. [Meibomia obtusa (Muhl.) Vail] Beg-
ce. Common in pinewoods, lawns.

r Lic
DE "SMODIUM GLABELLUM (Micl 1x.) DC. [Meibomia dillenii (Darl) Kuntze] Common
on dry roadsides, open woods.
** DESMODIUM ie heme (Nutt.) DC. [Meibomia laevigata (Nutt.) Kuntze]
Common in dry, open woo
DESMODIUM LINEATUM DC. [| Meibomia arenicola Vail] Common in dry, oe woods.
DE oe ae aia (L.) DC. [Meibomia paniculata (L.) Kuntze] Com-

ss

ods.
DE ‘SMODIUM TENUIFOL — T. & G. [Mcibomia tenuifolia Torr. and (Gray) Kuntze]
Infrequent in moist ar
*DESMODIUM TORTUO: uM (Sw.) DC. oe on dry roadsides, waste plac
DESMODIUM VIRIDIFLORUN, (L.) DC. [Meibomia vividiflora (L.) Ree: ae
quent on roadsides, open
GALACTIA ERECTA (Walt.) oe Milk Pea. eae in ie open woods.
*GALACTIA ee Michx. Common in dry, exe)
GALACTIA VOLUBILIS (L.) Britt. Common i ry, open woods.
GLOTTIDIUM VESICARIUM (Jacq.) Paes, ae pod. eee in moist,
areas near stream. N.
LESPEDEZA CAPITATA Michx. Common in dry, ey roadsides and woodland borders.
*LESPEDEZA CUNEATA (Dumont) G. Don. Sericea. Rare on dr ry roadside. N.
LESPEDEZA HIRTA (L.) Hornem. Common in cs open woods.
LESPEDEZA REPENS (L.) Bart. Common in dry, upland woods, roadsides
LESPEDEZA le (Thunb.) HH. & A. Japanese Clover. Govaman in Sale eed and

open

waste groun N.
*LESPEDEZA aoe Nutt. pases in dry, open. sites.
LESPEDEZA VIRGINICA (L.) Britt. Common in dry, open margins of wood, roadsides.

ate DIFFUSUS Nutt. heices Common in dry, open woods.
LILOTUS ALBA Desr. White Sweet Clover. Rare on roadsides, waste places. N.
sides. N.

MELILORUG INDICA (L.). All. Sour Clover. Rare on roads

134

sie faacags LUPULINA L. Black Medick. Common on roadsides, lawns, and waste

PE TALOSTEMUM GRACILIS Nutt. [Kuhnistera gracilis (Nutt.) Kuntze] Rare in

relands.
PE TALOSTE MUM haa INIANUM (L am.) Sprague. Vaal pinnata (Walt.)
Kun mer Farewell. Common in dry, open woo
*PUE RARIA Hae “CWill d.) sue Kudzu. Common on ae des, waste grounds.
ROBINIA PSEUDO-ACACIA L. Back Locust. Rare in woodland borders, probably es-
caped.
RHYNCHOSIA GALACTIOIDES Endl. Common in dry, open woo
RHY NCH RENIFORMIS DC. [R. simplicifolia (Walt.) sis Common in dry,

op
RHY NCHOSIA TOMENTOSA (L.) H. and A. [R. erecta (Walt.) DC.] Rare along
dry, sand dside.
SCHRANKIA MICROPHYLLA (Soland ex Sm.) Macbr. [Morongia angustata (Torr.
and Gray) Britton] Sensitive Brier. Common in dry, open woods.
SESBANIA EXALTATA ner A. W. Hill. [Sesban macrocarpum Muhl.] Infrequent
in moist, disturbed area
STYLOSANTHES BIFLORA (L.) BSP. Pencil Flower. Common in dry, open woods,
adsides
ST ROPHOSTY LES HELVOLA (L.) EIL [Phaseolus helvolus L.] Common in dry, open
ods, roadsides.
STROPHIOSTYLES _— (Muhl. ex Willd.) Britton. [Phaseolus umbellatus
(Muhl) Britton] Rare dry, open woods.
TEPHROSIA CHRYSOPHYLLA. Pursh. [Cracca chrysophylla (Pursh) Kuntze] Com-
non in dry, of woods.
“TEPHROSIA FLORIDA (E.G. Dietr.) C.E. Wood. [possibly same as Cracca smallii

(Small) Vail in Mohr] Common in dry, open woods.
alt.) T. G

TEPHROSIA SPICATA (Walt.) & G. [Cracca spicata (Walt.) Kuntze] Rare in
p pinela ANC ds.

*TRIFOLIUM Ae STRE Schreb. [T. procumbens L.] Low Hop Clover. Common on
lawns,

TRIFOLIUM CAROLINI ANUM Michx. Clover. Common on roadsides, cultivated and
waste grounds.
**TRIFOLIUM DUBIUM ana Hop Clover. Infrequent in cultivated and waste places. N.
sei ea INCARNATUM L. Crimson Clover. ane on pone
TRIFOLIUM REPENS _ White Clover. Common in cultivated and waste crannies: N.
ee aa RESUPINATUM L. Persian Clover. Rare ions roads de.
ca! ANGUS EOS Reich. Vetch. Common on_ roadsides, muleacal and waste
place
oe “DASY CARPA Ten. ae in waste places. N.
VIC TETRASPERMA (L.) Moench. Common in cultivated and waste grounds. N.
IW ee RIA SINENSIS — Wisteria. Infrec quent on roadsic des. old homesites. N.
ZORNIA BRACTEATA Walt. ex Gmel. Infrequent in dry, sandy soil.

a

LINACEAE
LINUM STRIATUM Walt. Common in moist. plac
LINUM VIRGINIANUM L. var. MEDIUM Planch. © jones in dry, open areas.
OXALIDACEAE
*OXALIS DILLENII Ue Wood Sorrel. Common in cultiv | and waste grounds.
*“OXALIS RUBRA St. Hil. Infrequent in disturbed, shady sites. N.
OXALIS STRICTA L. sy a in cultivated and waste eround
OXALIS VIOLACEA L. Violet. Infrequent in mesic wooc
GERANIACEAE
GERANIUM CAROLINIANUM L. Cranesbill. Common in cultivated and waste grounds.
MELIACEAE
MELIA AZEDARACH L. China-berry. Commonly cultivated, occasionally escaped on
woodland borders. N.

POLYGALA ea N
POLYGALA CRUCIATA
POLYGALA eS CDRE LOR:
POLYGALA ete
POLYGALA LUTE
POLYGALA RGA Mill.
POLYGALA NANA (Michx.)
POLYGALA RAMOSA Ell. In

POLYGALA VERTICILLATA

ACALYPHA GRACILENS Gray.

vated and waste grounds.
*ACALYPHA RHOMBOIDEA
*ACALYPHA SETOSA A. Ric

AS URITES FORDII Hemsley

eniborcouts STIMULOSUS (Michx.) Engelm. & Gray.
ods.

Spurge Nettle. Common in
CROTON
CROTON GLANDULO

1 waste places

sides an p
**CROTONOPSIS LIN

OSUS L.

Es Infrequent in
Infrequent in open,

1. Rare in waste places.

CAPITATUS Michx.

135

POLY GALACEAE

utt. Milkwort. a in wet pinelands.
Infrequent in olst, Open areas.
Wale. Csoson in dry,

open woods.
dry, open ‘penee woods.
marshy area,

Infrequent in soit ey place
DC. Common in open, ae wi
frequent in moist, open shee
Infrequent in dry, open woods.
EUPHORBIACEAE
Three-seeded Mercury. Common on roadsides, culti-
Raf. Rare in open area along stream.

N
Rarely escaped along stream and on roac

dside.

. Tung Tree.
[Jatropha stimulosa Michx.]
dry,
on roadsides waste

nd
var. SE PTE NTRIONAL Is Muell. Arg.

places.
Common on road-

EARIS ee Infrequent on roadside, woodland borders
EUPHORBIA COROLLATA Flowering Spurge. Common in dry, open w en
*EUPHORBIA HETEROPHY in L. Painted Leaf. Ra yn railroad right of way
EUPHORBIA HIRTA L. [probably same as E. oilulifera discolor Engelm in Mohr]
Common in cultivated and waste ground.
EUPHORBIA MACULATA L. Eyebane ae in waste grounds.
See SUPINA Raf. [E. humistrata Engelm.] Common on roadsides, waste places.

PHYL L KG THUS URINA
SAPIUM SEBIFERUM
Infrequent escaped in was
STILLINGIA SYLVATICA Ga
*TRAGIA SMALLII Shinners.

**CALLITRICHE DEFLEXA A.

NTHUS CAROL Pe NSIS W:

Ci: Rox. Chinese Tallov

Common in dry

ult. ees, in cultivated and waste grounds:
Common in oe rated and waste grounds.

Tree (Locally known as Doped £66).
grounds. N,

Common in

te

rden. dry, open sandy woods.

, open woods.

CALLITRICHACEAE
Infrequent

and inconspicuous in damp field.

RHUS COPALLINA L.

RHUS RADICANS L. Poiso:
TOX ICODENDRON
VERNIX L. Poison §

RHUS
RHUS

CLIFTONIA ae -

and border bay
CYRILLA R Ron: fie

ILE x ee (Michx.)
oods.

ASSINE

ILE
se x cOnIACEA’ aan Cha

LEX GLABRA (L.) Gray.

Dwarf Wingec
RHUS GLABRA L. Smocth or Cuma, Sumac.

Braun. [C. austinii Engelm.] Water Starwort.

mI DIACEAE

or { Sumac. Common in dry, open woods.
Kare on m_ bank.

Infrequent in
umac. Common ret pinelands— near str

ACEAE
Black Titi.

CYRILI
(Lam.)

Britton. Common in low pinelands

Swamp Cyrilla, Titi. Common in border of bay forest.

AQUIFOLIACEAE
(I. Walt

Torr. caroliniana .] Holly. Infrequent in upland

moist woodland borders

pm. Large Gallberry. Cima understory shrub in bay

Bitter Gallberry. Common in wet pinelands.

136

ILEX OPACA Ait. Holly. er in low swampy wo
ILEX VOMITORIA Ait. Yaupon. Common along ae. a bay forest.
CELASTRACEAE
EUONYMUS AMERICANUS L. Strawberry Bush. Infrequent in swamp forest.
ACERACEAE
ACER RUBRUM L. Red Maple. Common in low woods.
ACER RUBRUM var. DRUMMONDII (Hook. Arn.) Sarg. [A. drummondii Hook
& Arn.] Red Maple. Infrequent in low ods.
AMNACEAE
cae arenee SCANDENS (Hill) K. Kock. Rattan Vine. Infrequent in swamp forest

ear st
CE ANOTHUS AMERICANUS L. New Jersey Tea. Common in dry, open woods.
VITACEAE

AMPELOPSIS ARBOREA (L.) Koehne. Pepper-vine. Common in low woods, roadsides.

PART ENOCH QUINQUEFOLIA (L.) Planch. Virginia Creeper. Infrequent in
mesic woods

VITIS AE STIVAL AS Michx. Summer Grape. Common in shady blanc

VITIS ROTUNDIFOLIA Michx. Muscadine. Common in woodlan

VITIS VULPINA L. Frost Grape. Rare in low woo ce

MALVACEAE
HIBISCUS ACULEATUS Walt. Common in dry woods.
HIBISCUS MUTABILIS L. Rarely escaped along roadside.
HIBISCUS SYRIACUS L. Rose of Sharon. Rarely esc ae or persisting on old home
sites
MODIOLA CAROLINIANA (L.) G. Don. Common in cultivated and waste grounds.
SIDA RHOMBIFOLIA L. Common ee roadsides and in waste ground. N.

STERCULIACEAE
fect Seta A PL ATANIFOLIA (L.f.) Marsili. Japanese Varnish Tree. Rare, escaped from

cultivat
ME LOCHIA, CORCHORIFOL IA L. Chocolate-weed. Infrequent in waste places. N.

THEACEAE
GORDONIA LASIANTHUS (L.) Ellis. Loblolly Bay. Rare in bay forest.

HY PERICACEAE
*HYPERICUM BRACHYPHYLLUM (Spach) Steud. [probably eae to H. as-
palathoides Willd. in Mohr] St. John’s Wort. Rare in wet a eland
**HYPERICUM CANADENSE L. Infrequent on border of la
HYPERICUM CISTIFOLIUM Lam. Common along border es lake and low pinelands.
HYPERICUM GALIOIDES Lam. Rare in low pinelands.
or GENTIANOIDES (L.) BSP. Pineweed. Common on_ roadsides, waste

place
HYPE RICUM HYPERICOIDES (L.) Crantz. [Ascyrum hypericoides L.] St. Andrew’s
Cross mon in dry woods.

HYPE Na MUTILUM L. Common in wet, shady are

HYPERICUM PSEUDOMACULATUM Bush. [H. ae Walt.] Rare in open,
sandy woods.

HYPERICUM SETOSUM L. in savanna

HYPERICUM STANS (Mic On Adams a Rohn. tha stans Michx.] St. Peter’s

v Common in A to mesic woods.
*HYPERICUM STRAGALUM P. hdeiss and Robson. Infrequent along margins of swamp.
HYPERICUM ence L. [Triadenum virginicum (L.) Raf.] Common in wet,

shady areas.

137

CISTACEAE
pecan a CAROLINIANUM (Walt.) Michx. Frost-weed. Common in dry,
open
HELIANTHEMUM si aaeertar a Chapm. Infrequent in very dry, aon areas.
LECHEA MINOR L. ia Common in dry, open woods, disturbed place
LECHEA PATULA ons Infrequent in very dry, open areas.
LECHEA VILLOSA EI. ees in waste places, lawns.

VIOLACEAE
ie oes Se Brainerd. [probably corresponds to V. papilionacea Pursh and V.
palm L. in Mohr] Violet. Infrequent in pace
VIOLA LANCEOL ATA L. Infrequent in wet, sandy a
bee cece a L. [V. primu laefolia. ee " Pollard] Common in moist,

* VIOLA ‘SE PTE 7MLOBA House. [probably alpen V. brittoniana Pollard and V.
vicinalis Greene in Mohr] Common in dry, open oe pinelands
PASSIFLORACEAE

PASSIFLORA INCARNATA L. Maypop. Infrequent sp roadsides and in waste ee
PASSIFLORA LUTEA L. Yellow Passionflower. Rare along woodland border

CACTACEAE
ca mous COMPRESSA (Salisb.) Macbr. [O. humifusa Raf.] Prickly Pear. Rare in
dry, open woods.
ELAEAGNACEAE
*ELAEAGNUS UMBELLATA Thunb, Silverberry. Rare in waste places. N.

LYTHRACEAE
UPHEA CARTHAGENSIS (Jacq.) Macbr. Waxweed. Infrequent in wet pinelands.
LAGERSTROEMIA INDICA L, Crape-myrtle. Infrequent, escaped on old homesites. N.

>

MELASTOMATACEAE
RHEXIA ALIFANUS Walt. [R. glabella Michx.] Meadow-beauty. Common in low

RHEXIA TEA Walt. Infrequent in low pinelands, margins of ponds.
RHEXIA Ne L. var. MARIANA. Common in wet meadows, borders of pond,

es.
*“RHEXIA MARIANA L. var. PURPUREA Michx. Occasional in moist a n areas,
RHEXIA PETIOLATA Walt. [R. ciliosa Michx.] Rare in wet, open are
RHEXIA VIRGINICA L. var. VIRGINICA. Occasional in moist, open areas
RHEXIA VIRGINICA var. PURSHII TSnvenoel) C.W, James. [R. stricta Boreh Rare

in wet, Open areas,

ONAGRACEAE

AURA ANGUSTIFOLIA Michx. Rare on roadsides
: ae FILIPES Spach. [G. na auxil eee Coauied in dry, open woods.
LUDWIGIA ALTERNIFOLIA L. Common along margin of stream and ponds.
*LUDWIGIA BONARIE ais (Micheli) Hara. Rare along margins of pond.
LUDWIGIA ea nee Walt. [Jussiaea decurrens (Wale.) DC oaks along

margin of stream and ponds.
Bee GL ANDULOSA Walt. Common in wet pinelands, borders of stream and
pon

LUDWIGIA ea (Nutt.) H. Hara. [Jussiaea pilosa H.B.K.] Common along
tream and pond ma

LUDWIGIA LINE ARIS. Walter, Infrequent in moist, open areas.

LUDWIGIA PALUSTRIS (L.) Ell. [Isnardia palustris L.] Infrequent along edge of
tream and ponds

LUDWIGIA PILOSA Walt. Common along stream and pond margins.

138

LUDWIGIA REPENS Forster. Rare along stream.

OENOTHERA BIENNIS L. [Onagra biennis (L.) Scop.] Evening Primrose. Common
on roadsides, waste places.

OENOTHERA FRUTICOSA L. [Kneifha fruticosa (L.) Raimann] Infrequent in dry,
open woods.

OENOTHERA LACINIATA Hill. Common in cultivated and waste plac

OENOTHERA SPECIOSA Nutt. [Xylopleurum speciosum (Nutt.) Raimann] Infrequent
on roadsides. N

HALORAGACEAE

Bore cre BRASILIENSE Camb. Parrot-feather. Rare, partly submerged along
dge of s

PROSE RPINACA PE CTINATA Lam. Infrequent in moist, open areas and ditches.

ARALIACEAE
ARALIA SPINOSA L. Hercules Club. Common in wet open places.
APIACEAE
ANGELICA VENENOSA (Greenway) Fern. Rare in mesic woods.
APIUM LEPTOPHYLLUM (Pers.) F. Muell. [A. anmi (L.) Urban] Marsh Parsley.
Infrequent in disturbed places. N.
CENTELLA ASIATICA (L.) Urban. Common in wet, open areas.
Shee ROPHYLLUM TAINTURIERI Hook. Wild Chervil. Infrequent in waste places.
AUCUS PUSILLUS Michx. Wild Carrot. Rare in waste places.
Bec INTEGRIFOLIUM Walt. pao in wet pinelands.
ERY NGIUM Sp aun Michx. var. SYNCHAETUM Michx. Button Snakeroot.
Infrequent in dry ods.
ee BONARIE NSIS Lam. [Possibly included by Mohr under H. verticil-
ata] Marsh Pennywort. Common in moist, open areas.
ea. RIGIDIOR (L.) Raf. Cowbane. Rare in moist, open area
PTILIMNIUM CAPILLACEUM (Michx.) Raf. Common in moist, Sioa grounds
and waste places
SANICULA CANA \DENSI S L. Sanicle. Rare in mixed deciduous woo
_— See i DIVARICATA (Walt.) Raf. [S. divaricatus pu ie Britton] Rare on

adsides
SPERMOL EPIS ECHINATA (Nutt. ex DC.) Heller. [S. echinatus (Nutt.) Britton] In-
waste places.

TREPOCARPUS AETHUSAE Nutt. Infrequent along moist, sandy stream banks.
NYSSACEAE

NYSSA BIFLORA Walt. Black Gum. Common in open areas along bay forest.

NYSSA SYLVATICA Marsh. Black Gum. Infrequent in dry, shaded woods.
CORNACEAE

CORNUS FLORIDA L. Flowering Dogwood. Common i

1 woodlands.

CLETHRACEAE
CLETHRA ALNIFOLIA L. var. TOMENTOSA (Lam.) Michx. Sweet Pepperbrush.
Common on margins of creek and bay forest.
ERICACEAE
EPIGAEA REPENS L. Trailing Arbutus. Rare in wooded slope along stream.
GAYLUSSACIA DUMOSA (Andr.) T. & G. Dwarf Huckleberry. Common in dry,
open woods.
GAYLUSSACIA MOSIERI Small. [G. hirtella (Ait. f.) Klotzsch] Infrequent in sandy
border of stream.
GAYLUSSACIA NANA (A. Gray) es [Included in G. frondosa tomentosa Gray
in Mohr] tea Gay, in dry, open woods.
KALMIA LATIFOLIA L. Mountain Lau rel. Infrequent in shaded stream banks.
LEUCOT HOE AXILLARIS (Lam.) D. Don. Downy Fetter-bush. Common in bay forest

and along strea

139

LYONIA LUCIDA (Lam.) K. hae nitida (Bar a Benth. and Hook.] Shining

Fetter-bush. Common — is nelands and along. str
MONOTROPA UNIFLOR tedien Pipe. Infrequent in ae pinewoods.
se nme nay (L.) DC. Sourwood. Common in wooded slopes along

*RHODODENDRON CANESCENS (Michx.) Sweet. [probably included under Azalea
|

nudiflora L. in Mohr] Wild Azalea. Infrequent in low pinelands.

ODODENDRON VISCOSUM (L.) Torr. var. SERRULATUM (Small) ae
[Az alea viscosa L.] Sw ial Honeysuckle, Swamp Azalea. Common on stream ma
gins and in low pinelar

a.

VACCINIUM A AROR UAE Marsh, se eberry. Common in dry, open wooc
*VACCINIUM Faas II ens [prob ine corresponds to V. myrsinites Lam. in > Mohr]
Common in open woods and pineland

be ei ELLIOTTH ies eens Common in shaded woods along bay

VACCINTOM pa ala L. var. SERICEUM Mohr. Squaw Huckleberry. Common
in

VACCINIUM TE NELLUM Ait. Blueberry. Rare in dry, open woods.

VACCINIUM VIRGATUM Ait. Highbush Blueberry. futeaeel along border of bay
forest

PRIMULACEAE
CENTUNCULUS MINIMUS L. Chaffweed. Infrequent in moist, open areas.

FBENACEAE
DIOSPYROS VIRGINIANA L. Persimmon. Common in dry woods.
SYMPLOCACEAE
SYMPLOCOS oleae (L.) L’Her. Horse-Sugar, Sweetleaf. Common in shaded
woods along bay fores
STYRACACEAE
STYRAX AMERICANA Lam. (S. pulverulenta Michx.] Storax. Rare on stream bank.
OLEACEAE
pat aes RUM SINENSE Lour. Privet. ae in low woods, waste places. N.
oe AMERICANA (L.) Gray. Wild Olive. ee in open woods along
es fores
LOGANIACEAE
CYNOCTONUM MITREO (L.) Britton, oe wort. Rare in wet, open pineland.
TONUM SE Soro J.P. Gmel. Infrequent in open pinelands.
Sy MIUM SEMPERVIRENS (L.) Ait. Yellow oe ye in dry, open

oods.

POLY PREMUM PROCUMBENS L. Common in cultivated and waste grounds.
GENTIANACEAE

GENTIANA VILLOSA L. Gentian. Rare in moist area along swamp.

SABATIA ANGULARIS (L.) Pursh. Rose Pink. Infrequent in wet pinelands.

SABATIA BRACHIATA Ell. [S. angustifolia (Michx.) Britton] Common in dry, open

woods.

SABATIA CAMPANULATA (L.) Torrey. Rare in low pinelands.
ASCLEPIADACEAE

ASCLEPIAS AMPLEXICAULIS Smith. [A. obtusifolia Michx.] Milkweed. Infrequent in

dry woods, roadsides.
ASCLEPIAS HUMISTRATA Walt. Infrequent in dry open woods.
ASCLEPIAS MICHAUXII Dene. Common in dry, open woods.
ASCLEPIAS OBOVATA EIL. Infrequent in dry, open woods.
ASCLEPIAS TUBEROSA L. Butterfly-weed. Infrequent in dry, open woods.

140

**ASCLEPIAS VERTICILLATA L. Rare in dry, open woods.
CONVOLVULACEAE
*BONAMIA PATENS (Desr.) Shinners. Common in dry, sandy woods.
CUSCUTA CAMPESTRIS anaes Dodder. 7 eet parasite in moist areas

CUSCUTA COMPACTA Juss. Common parasite on woody plants in open, moist areas.
DICHONDRA CAROLINENSIS Michx. [D. evolvulacea (L.f.) Britton] Common in

»

oist lawns and waste

lac
JACQUEMONTIA TAMNIFOLIA (L.) Griseb. Infrequent in open waste places, road-
JN.

en A COCCINEA L. Red Morning-glory. Rare in waste places. N.
IPOMOEA HEDERACEA (L.) Jacq. Morning-glory. Common in dry, open woods and

IPOMOEA LACUNOSA L. Rare in dry, open woods and waste places.

IPOMOEA PANDURATA (L.) G.F.W. Mey. Man-root. Common in dry, open woods
and waste places.

sue PURPUREA (L.) Roth. Common Morning-glory. Infrequent in waste

plac
IPOMOE A QUAMOCLIT L. Cypress Vine. Infrequent in cultivated and waste places. N.
IPOMOEA TRICHOCARPA EIL. [I. carolina (L.) Pursh] Infrequent in dry, sandy areas.

POLEMONIACEAE
PHLOX PILOSA L. [Including P. pilosa detonsa Gray in Mohr] Common in dry, open
woods and_ pinelands.
BORAGINACEAE
ONOSMODIUM VIRGINIANUM (L.) A. DC. False-Gromwell. Common in dry, sandy
open woods

VERBENACEAE
CALLICARPA AMERICANA L. French eae Common in open woods, thickets.
LANTANA CA cee L. in waste places. N.
LIPPIA NODIFLORA (L.) Michx. Common in moist, open sandy places, lawns.
*VERBENA Sen Ei Common in waste places, roadsides. N.
VERBENA CARNEA Medicus. [V. carolina L.] Common in dry, open woods.
VERBENA OFFICINALIS L. Rare on roadsides,
VERBENA RIGIDA Spreng. Infrequent on roadsides, Hg places. N.
*VERBENA TENUISECTA Briq. Rare in dry, open woods. N.

LAMIACEAE
CAL pe haa ae A ey Benth. in DC. [Clinopodium coccineum (Nutt.)
Kuntze] Co dry, open woods.
DRACOCE PHALUM VIRGINIANUM L. [Physostegia virginiana (L.) Benth.] Obedient

Pla re dit roadsides.
“HE DE ae HISPIDUM. Pursh, Infrequent along dry roadside
HYPTIS ALATA (Raf.) Shinners. [Mesosphaerum rugosum (L. ) Pollard] Common in

woods.
*HYPTIS MUTABILIS (A. aya Briq. Rare along moist shady roadside. N.
LAMIUM AMPL . eee L. Henbit. eee on roadsides, sna waste cee N.
LYCOPUS RUBELLUS vee Common along margins of mp for

**TYCOPUS VIRGINICU US L. Rare along margin of swamp ie.
i >n woods,

: f
PERILLA FRUTESCENS (L.) Britt. Infrequent along shaded, disturbed stream banks.
PYCNANTHEMUM INCANUM (L.) Michx. [Koellia albescens (Torr. & Gray)
r

SALVIA AZUREA Lam. Sage. © in ee open woods.

SALVIA LYRATA L. Common in cultivated and waste grounds.

SCUTELLARIA ELLIPTICA Muhl. [S. pilosa Michy. 1 Skullcap. Common in dry, open
woods.

SCUTELLARIA INTEGRIFOLIA L. Infrequent along border of pond.

141

*STACHYS yak ass Shuttlew. Locally known as Rattlesnake Weed. Common in
cultivated and waste grounds and along stream banks.

TRICHOSTEMA DICHOTC OMUM L. Blue Curls. Common in dry, open woods.

TRICHOSTEMA SETACEUM Houtt. [T. lineare Nutt.] Infrequent in dry, open woods,
roadsides.

SOLANACEAE
DATURA STRAMONIUM L. Jimson Weed. Rare in disturbed habitats. N.
PHYSALIS ANGULATA L. Ground Cherry. Rare on waste grounds. N.
PHYSALIS PUBESCENS L. [P. barbadensis Jacq.] Uncommon weed in campus lawn.
PHYSALIS VISCOSA L. Infrequent on dry, sandy roadsides.
SOLANUM AMERICANUM Mill. Nightshade. Infrequent in waste places.
SOLANUM CAROLINENSE L. Common on roadsides, waste places

SOLANUM SISYMBRIFOLIUM Lam. Rare in waste places. N

°

SCROPHULARIACEAE

AGALINIS ee (EI.) Rat. [Gerardia fasciculata Ell.] Infrequent in dry,
open

*AGAL INIS. ‘OBTUSIFOLIA Raf. Rare in dry, open woods.

AGALINIS PURPUREA (L.) Pennell. [Gerardia purpurea L.] Common in dry, open
woods.

AGALINIS SETACEA (J.F. Gmel) Raf. Common in dry, open wo

** AURE oe FLAVA (L.) Farwell. [Dasystoma flava (L.) W Mok aoe in dry,
open we

eae ARIA oo (Nutt.) Pennell. [Dasystoma pectinata (Nutt.) Benth.]

nmon in dry ods.

secann RA FLORIDANA Gandoger. ee close to B. americana and B. elongata Sw.
in’ Mohr] Infrequent in moist savannahs.

ere PILOSA Michx. Hedge Hy: ssop. Infrequent in moist open areas along stream

nd ponds.
GRAT [OLA VIRGINIANA L, [G. sphaerocarpa Ell] Infrequent in moist, open areas.

LINARIA CANADENSIS (L.) Dumont. Blue Toad-flax. Common on roadsides, weedy
places, lawns

LINDE RNIA DUBIA (L.) Pennell. [Ilysanthes gratioloides (L.) Benth.] Rare in wet,
muddy margins of stream.

*MAZUS ae, (Thunberg) Kuntze, Rare in moist lawns. N.

MECARDONIA ACUMINATA (Walt.) Small. ican: acuminata (Walt.) Kuntze]
pane non in wet, open areas near stream.

as sides UMBROSUM (J.F. Gmel.) Blake. [M. orbiculatum Michx.] In-
frequer et, sandy open areas.

PE DICULARIS CANADE NSIS L. Lousewort, Wood Betony. Rare in shaded pinelands
near swamp for

*PENSTE MON AUSTRALIS Small. [ Probably corresponds at least partly to P. hirsutus

Willd. in ee Beard-tongue. Common in dry, open woods.

*SCOPARIA DULCIS L. Sweet-broom, Goat Weed. Rare in cultivated grounds. N.

VERBASCUM THAPSUS L. Wooly Mullein. Rare in waste places. N

VERONICA ARVENSIS L. Corn Speedwell. Common in cultivated mee waste grounds. N

VERONICA PEREGRINA L. Infrequent in cultivated and waste grounds.

BIGNONIACEAE
ie oe ATA (L.) Dupre [Bignonia crucigera L.] Cross Vine.
alon

fc
Saves RADICANS 0 Seem. oe radicans L.] Trumpet Vine. Infrequent in

BIGNONIOIDES Walt. [C. catalpa (L.) Karst.] Infrequent in low, dis-
eae woodlands.
OROBANCHACEAE
*“OROBANCHE UNIFLORA L. Cancer-root. Rare in moist, sandy, open area near stream.

142

LENTIBULARIACEAE
PINGUICULA PLANIFOLIA — Butterwort. Rare in low’ pinelands.

in low. pinelands.
UTRICULARIA BIFLORA Lam. Blad fdey wort. Infrequent in shallow margins of ponds.
UTRICULARIA JUNCEA Vahl. Infrequent in shallow a of ponds.
UTRICULARIA SUBULATA L. Common in wet pineland

ACANTHACEAE
RUELLIA CILIOSA Pursh. Common in dry, open woods.

PLANTAGINACEAE
de eo ARISTATA Michx. Plantain. Infrequent in dry, sandy roadsides, railroad
ballast.
PL ANTAGO HETEROPHYLLA Nutt. Common on_ roadsides, cultivated and waste

*PLANTAGO a RIANA Fisch. & Mey. var. NUDA (Gray) Poe. Infrequent on
railroad ballast.

PLANTAGO LANCEOLATA L. Infrequent in waste grounds. N.

PLANTAGO MAJOR L. Rare lawn weed.

PLANTAGO VIRGINICA L. Common in cultivated and waste grounds.

RUBIACEAE
CEPHALANTHUS OCCIDENTALIS L. Button Bush. Infrequent in low areas, margins
of creek and ponds.
DIODIA TERES Walt. Common in dry roadsides, waste places, open woods.
DIODIA VIRGINIANA L. Common in moist, cultivated ground and swamp forest mar-
gins.
GALIUM APARINE L. Bedstraw. Infrequent in moist roadsides and waste pl
GALIUM HISPIDULUM Michx. Common in “ ah open woods.
GALIUM eens Ait. Common in dry, 1 woods, roadsides.
GALIUM ‘TINCTORIUM L. Infrequent ane presen of pond.
HOUSTONIA Seine NS (J.-F. Gmel) Standley. [H. rotundifoli ia Michx.] Common
in’ sandy pinelands.
|! 1OUSTO ae cig Schoepf. [H. minor (Michx.) Britton] Infrequent in cultivated
and waste unds.
MITCHELL Ns RE PENS L. Partridge Berry. Infrequent in mesic woods.
OLDENLANDIA BOSCIL (DC) Chapman. Infrequent a ie of ponds and streams.
*OLDENLANDIA ee MBOSA L. Rare in waste pl

aces.

OLDENLANDIA FASCICULATA a rtol.) Small. 0. cele Mohr] Infrequent in
moist, open areas,
OLDENLANDIA UNIFLORA L. Infrequent in moist, open areas

areas
*RICHARDIA BRASILIENSIS (Moqg.) Gomez. Rare on roadsi des. N.
RICHARDIA SCABRA L. Common in cultivated and waste grounds N.
*SHERARDIA ARVENSIS L. Infrequent in cultivated grounds.

CAPRIFOLIACEAE
LONICERA JAPONICA Thunb. Japanese Honeysuckle. Common in cultivated and
ste 3 ‘ N.
LONICERA SEMPERVIRENS L. Coral) Honeysuckle. Rare in’ pinelanc
pie enna CANADENSIS L. Elderberry. Infrequent in open aaa
eam
VIBURNUM NUDUM

stream banks.

areas along

—

Possum-Haw Viburnum. Common in bay forest and along

VALERIANACEAE
VALERIANELLA RADIATA (L.) Dufr. Corn Salad.

places.

Infrequent in moist, open waste

CUCURBITACEAE
MELOTHRIA PENDULA L. Creeping Cucumber. Infrequent in moist cultivated grounds.

143

CAMPANULACEAE
*LOBELIA GEORGIANA roe Lobelia.
LOBELIA A tee oe Wal
Infrequent pin
LOBELIA PUBERULA. Mics font in dry to moist open woods.
SPECULARIA BIFLORA (R. & P.) Fisch. & Mey. [Legouzia biflora (Ruiz & Pav.)

I quent in low pinelanc
ae eeiene a to L. Paludosa a in Mohr]

i ] Venus’ Loo rN glass. Common _ in ae d and waste grounds.
SPE ere ages (. ? A.DC, oes perfoliata (L.) Britton] Common
in cultivated and waste groun
*WAHLENBE RGIA MARGINATA (Thunb.) DC. Infrequent in cultivated and waste
grounds. N.,

ASTERACEAE
ACHILLLEA MILLEFOLIUM L. Yarrow. Rare
AG ERATINA AROMATICA (L.) RK. & R.

in waste places. N.

[Eupatorium aromaticum L.] Common

aoe ARTE MISHIFOLIA L. Ragweed. Common on roadsides, waste places.
MBROSIA TRIFIL L. Giant Ragweed. oe on roa pie

oe R Na rin ae Aster. Commo

7 R nae TATUS (Walt.) Ahles.

1 woods.

ASTE R CONCOLOR L. Common in dry,

ASTER DUMOSUS L. Common in dry, o

ASTER EXILIS Ell. Infrequent in moist, open. sites.

ASTER LINARIIFOLIUS L. [Tonactis linariifolius (L.) Greene] Common in dry, open

WC ca
ASTER PATENS Ait. var. GRACILIS Hook. Common
ASTE . PILOSUS Willd. [A.

waste places.
in dry, open we
[Sericoc arpus bifoliacus, (Wa alt.) Porter] Common

open woods.

ICN sites.

in woodland border

ericoides pilosus (Willd.) Porter] Frost Pace Rare in
woodland bor

der
ma ER SOL IDAGINE US Michx.
Aster. Infrequent in dry —
ASTE R UMBELLATUS Mill. TARE V ISQUAMUS Fern.
Britton] Rare along margin o
BA eet HALIMIFOLIA L.

iad linifolius (L.) B.S.P.]) White Topped
[ Doellingeria humilis ( Willd.)
amp. fores
ome eee: Sea Myrtle. Infrequent in wet, open
ve DUINA UNIFLORA Nutt.
mon in moist savannahs.
BIDENS BIPINNATA L. Spanish Needles.
BIDENS FRONDOSA L. Begger-ticks.
BIDENS MITIS nee) Sherff. [Ve

[Actinospermum uniflorum (Nutt.) Barnhart] Com-

Infrequent in moist open areas.
Common in wet open. areas.

ery close to B. Coronata (L.) Fisch. in Mohr]

Common in ypen area,
mae EL ee NUDA FA (Michx.) DC. Leet a nudata (Michx.) Britton]
ayless Goldenr Infrequent in open moist

oe DISCOIDEA (ELL)
*CALYPTOCARP /TALIS Less. Rare in waste erie
CARDUUS SPINOSISSIMUS Walt. Thistle.
CHAPTALIA TOMENT OSA Vent.

Sun-bonnets.. Common

Infrequent on roadsides, waste plac
a semiflosculare (Walt.) eer

rshy area.
CONN ie ess ae Lest INUM (L. ) DC. [Eupatorium coelestinum L.] Mist Flower.
Common pen ditches.
CORFOPSIS “BASALIS (Die) Blake. [C. drummondii (D.Don) Torr. and Gray]
Ra in vacant lot. N

CORE ‘OPSIS MATOR Ws te Common in dry,

COREOPSIS TINCTORIA Nutt. Infrequent along roadsides. N.

*CREPIS JAPONICA (L.) Benth. Hawk’s beard. Infrequent in disturbed habitats, flower
L +

seds. N.
CROPTILON DIVARICATUM (Nutt.) Raf.

open woods,

[Isoppapus divaricatus (Nutt.) Torr.
and Gray] Common on roadsides, waste places

ECLIPTA ALBA (L.) Hassk. Rare on stream Gane

*ELEPHANTOPUS ELATUS Bertoloni.

Common in dry, open woods.

144

**ELEPHANTOPUS NUDATUS Gray. anni in dry, — woods.
ELEPHANTOPUS TOMENTOSUS L. Infrequent in dry woods.
E - CHTITES HIERACIFOLIA (L.) Raf. Fireweed. hee eat in moist, disturbed

ERIGE RON BONARIENSIS L. Common on roadsides, waste plac
ERIGERON CANADENSIS L. Horseweed. oe on roads oe waste places.
ERIGERON PHILADELPHICUS L. Daisy Flea . Common on ye ground,
ERIGERON STRIGOSUS va ex Willd. = ramosus (Walt.) B.S.P.] Common on
roadsides, waste places.

ERIGE RON VERNUS (L.) T. & G. Infrequent in moist, open areas.
EUPATORIUM ALBUM L. Pei ada Common in dry, of ods.
EUPATORIUM CAPILLIFOLIUM oe Small var. CAPIL LIF OL TUM.

Dog Fennel.

Common in roadsides, disturbed areas
pore On IUM CAPIL LIF OLIUM var. ~ LEPTOPHYLLUM (DC.) Ahles. Infrequent on
roadsides
E sein r ORIUM COMPOSITIFOLIUM Walt. Rough Dog Fennel. Common on roadsides,
plac

FUPATORIOM CUNEIFOLIUM 9 Willd. [E. linearifolium Walt.
Shapm.] Common in dry open woods, waste places.

E UPATORIUM FISTULOSUM $ Barratt. [E. maculatum L.] Joe-Pye Weed.
along margin of swamp forest.

EUPATORIUM PERFOLIATUM L. Boneset. Infrequent along margin of as forest.

*EFUPATORIUM RECURVANS Small. Thorou ughwort. nfrequent in dry woodlands.

EKUPATORIUM ROTUNDIFOLIUM L. Common j ;

EUPATORIUM ar RRATUM DC. Infrequent in moist woodland

*PACELIS RETUSA (Lam.) Sch.-Bip. Common in eet sandy roadsides, lawns. N.

GAILLARDIA AEST Soe IS (Walt. ) Rock. var. LANCEOLATA (Michx.) Ahles. [G.
lanceolata: Michx.] Common in dry, open woo as

GNAPHALIUM OBTUSIFOLIUM L. R abbit Tobacco.

unds.

GNAPHAL IUM PURPUREUM L. Cudweed. Common in cultivated and waste grounds.

GNAPHALIUM PURPUREUM var. SPATHULATUM (Lam.) Ahles. [G. spatulathum
Lam.] Common in cultivated ee Ww re groun

HELENIUM AMARUM ia, H. . [ H.
on roadsides, waste place

HELENIUM FLE XUOSUM. Raf. [H. nudiflorum Nutt. ]

and E. tortifolium

Infrequent

_

n

Common in cultivated and waste

ids.
tenuifolium Nutt.] Bitter-weed. Common

Sneeze-weed. Rare in moist,
“HE L [ANT HUS ANNUUS L. pias, Rare in waste place

HELIANTHUS ANGUSTIFOLIUS L. Common in wet. savannahs.
HELIANTHUS HETEROPHYLL . pues Infrequent along moist,
HELIANTHUS RADULA (Pursh) T.

open stream banks.

. Common in nek open ne
HETE ee THECA 7 Shee OLIA ve hx : Shi [Chrysopsis | graminifolia
(Michx.) Nutt. ] aiden Aster. Common in y; ods.
— nee RO THE os Teer (L.) Shinners. a Bao HEe (L.) Nutt.] Com-

dry ods.
HETEROTHECA. TRICHOPHYLLA (Nutt.) Shinners
Infrequent in dry woods.
HIERACIUM GRONOVII L. Hawkweed. Infre
*HYPOCHAERIS ELATA ee 2 Asta
a RIS RADICATA lav

. [Chrysopsis trichophylla Nutt.]

equent in dry woods.

ioe lawn weed, roadside.

wn weed.

IVA NUA L. [I. ciliata wa oe along roadside.

ae. OPPOSITIFOLIA Raf. pposifli (Raf.) Kuntze] Dwarf Dandelion.
Common on_ roadsides, cultivated gro

KRIGIA VIRGINICA (L.) Willd. Palonaten carolinianum (Walt.) Britton] Common
on roadsides, cultivated grounds.

KUHNIA MOSIERI Small. [K. kuhnia (Gaert.) Mohr] False Boneset. Rare in dry pine-
woods.

LACTUCA > vacbabinne Ly [es
waste plac

LACTUCA FLORIDANA (L) Gaertn.

sagittifolia Ell.] Wild Lettuce. Common on roadsides,

Infrequent along disturbed stream banks.

145

LIATRIS ene ages Michx. [Lacinaria elegans (Walt.) Kuntze] Blazing Star.
Common in oods.
ee GRACILIS. Pakh: [Lacinaria gracilis Pursh] Infrequent in dry, open wood-

an
LIATRIS genes Willd. [Lacinaria graminifolia pilosa (Ait.) Britton] Com-
mon in
LIATRIS SPICATA. (Ly “Will . [Lacinaria spicata (L.) Kuntze] Rare in dry, open site.

me

ae oe na ane hx. eens ae (L.) Hill] Infrequent in
dr

MIKANTIA " SCANDENS (L.) Willd. [Willughbaeya scandens (L.) Kuntze] Climbing
oneset. Common in moist open areas.

PARTHENIUM HYSTEROPHORUS L. Rare in cultivated and waste grounds. N.

ee A ad gene (L.) Dc. Iptae hfleabarie, eee along margin of creek

nd ponds in wet savannah,

PLUCHE A FOE TIDA (L.) DC. Common along margin of creek

PRENANTHES SERPENTARIA Pursh. [Nabalus serpentaria eee Lae Gall-of-
the-earth. Infrequent in dry woodlands.

PYRRHOPAPPUS CAROLINIANUS (Walt.) DC. [Sitilias caroliniana (Walt.) Raf]
False Dandelion. Common on roadsides, lawns, waste places.

RUDBECKIA HIRTA L. Black-cyed Susan. Common in dry, open woods, waste places.

seer GLABELLUS Poir. [S. lobatus Pers.] Butterweed. Rare in moist, cultivated

inds.

**SILPHIUM DENTATUM Ell. var. GATESIL (Mohr) Ahles. [S. gatesii Mohr] Infrequent

along margin of swamp for

SOLIDA GO ALTISSIMA L, Is. pore ce L.] Goldenrod.
place

SOLIDAGO FISTULOSA Mill. i. are In moist pine elands.

SOLIDAGO GRAMINIFOLIA (L.) Salisb. Rare in moist, open area

*SOLIDAGO MICROCEDHALA (Greene) Bush. [P easibly ee to Euthammia
caroliniana (L.) Greene in Mohr] Common in wet, Open areas.

SOLIDAGO ODORA Ait. Common in dry, open woods.

SOLIDAGO PATULA Muhl. ex Willd. var. STRICTULA T. & G. [S. salicina EIll.]
Rare in moist, open area.

SOLIDAGO PUBERULA Nutt. var. PULVERULENTA (Nutt.) Chapm. Infrequent in
dry, open woods.

SOLIDAGO RUGOSA Mill. Infrequent in dry woods, roadsides.

*SOLIDAGO RUGOSA var. CELTIDIFOLIA (Small) Fern. Infrequent in moist areas

Common on roadsides, waste

mans

near stream.
SOLIDAGO STRICTA Ait. Infrequent in moist pee eads
*“SOLIDAGO TORTIFOLIA FIl. Infrequent in y, open woods.
SOLIVA PTEROSPERMA (Juss.) Less. pec in moist lawns.
SONCHUS AS ey (L.) Sow-thistle. Common on roadsides, w aste places. N.
ACEUS L. Common on roadsides, Hs plac
SPILANTHES eiCane (Mutis) Hieron. var. REPENS Ww ules A.H. Moore [S.
repens (Walt.) Michx.] Rare on stream bank

TARAXACUM OFFICINALE Wiggers [T. taraxacum (L.) Karst] Dandelion, Infrequent

ie}

on lawns. N.
TETRAGONOTHECA HELIANTHOIDES L. euler in dry, open woods.
TRILISA ODORATI pols (J.G. Gmel.) Cass. Deer’s Tongue. Common in dry, open
ids.

a
[any

wn

woods
VERBESINA OCCIDENTALIS (L.) Walt. Crownbeard, Wingstem. Common on road-

sides ers,
VERBESINA VIRGINICA L. Rare in dry, open wo
VERNONIA ALTISSIMA Nutt. [V. gigantea oleen Speeoal Ironweed. Infrequent
on border of swamp forest.

VERNONIA ANGUSTIFOLIA Michx. [V. graminifolia Walt.] Common in dry, open

woods.
XANTHIUM STRUMARIUM L. var. GLABRATUM (DC.) Cronquist. Cocklebur. In-
ks.

frequent along open stream ban

REFERENCES
CLARK, R. C. 1971. The woody plants of Alab

. Ann. Missouri Bot. Gard.
DERAMUS, R. 1970. Studies on the flora

5
of ihe ‘ssc plants of Dauphin fdand, Mobile
iology, University of Alabama, University.
Catalogue of the trees,
Monogr. 9. 357 pp.

shrubs and vines of Alabama, Geol. Surv. Ala.
RA . 1966. Xyris (Xyridaceae) of the continental United States and Canada. Sida

2(3): 177-260.
- 1971. A treatment of Abildgaardia, Bulbostylis and Fimbristylis (Cyperaceae)
for Roreh America. Sida 4(2): 57-227
LLOYD, F. E. and S. M. TRACY. (1901,

Bull. Torrey Bot. Club 28: 61-10
a ,G. J. and S. B. JONES, a 1967.
Castanea 32: 84-99,
MOHIR, CHARLES 1901. Plane life of Alabama. Cont
PENFOUND, W. T. and M. E. O'NEILL. 1934,
Ecology 15: ae
PESSIN, L. J. and T. D. alae IGH. 1941.
Mississippi. Ecology 22 (1 0-78
RADFORD, A. E., H.E. AHL 7 cad Cc. BELL. 1968. Manual of the vascular flora of
the Carolinas. University of North a Press, Chapel Hi
RICHMOND, E. A. 1962. The fauna and flora of Horn Island, Wisdasont Gulf Research
Reports aye 59-106.

8.

The insular flora of Mississippi and Louisiana.
The vascular flora of Ship Island, Mississippi.

6S. Natl. Herb. 6: 1-921
The vegetation of Cat Island, Mississippi.

Notes on the forest biology of Horn Island,

A eee to the fauna and flora of Horn Island, Mississippi. Gulf
Research Repor 23 13-254.

SMA J. K. 1933. eee of the Southeastern flora. University of North Carolina Press,
preety lll.

SYNOPSIS OF SUAEDA (CHENOPODIACEAE)
IN NORTH AMERICA

CHRISTINE O, HOPKINS! AND WILL H. BLACKWELL, JR.
Herbarium, Department of Botany, Miami University
Oxford, Ohio 45056

ABSTRACT

Tne study of North American elie es of se halophytic genus Suaeda
has led to recognition of 14 species and five varieties plus two species of
doubtful status. Taxonomic pro slong associated with the succulent habit of
Suaeda and the historical development of the genus are discussed. Dis-
tribution, typification and systematics of the species are presented with
citation of specimens from each county or district recorded,

Suaeda, a chenopodiaceous genus of sea coasts and inland saline soils,
has long been a source of confusion to taxonomists and generally ignored
by collectors. It is a relatively unattractive plant and species’ character-
istics are not easily noted without magnification. Dried specimens do not
always show the same leaf and calyx features as fresh, succulent plants,
and leaf shape in cross section varies with the degree of salinity of the
habitat.

The plants seem to exhibit phenotypic plasticity which has not previously
been taken into account by taxonomists in treatments of Suaeda. Succulence
and roundness of the leaf in cross section increases with increasing salinity
of habitat. In his study on halophytes Ungar (1974) found that even the
habit of Suaeda, particularly S. depressa and S. maritima, varies according
to the salinity of the soil. He reported that in highly saline soils S. depressa
will be dwarfed with a prostrate growth form, or appear as a weak, single-
stemmed upright plant; both forms under 20 cm. long. In conditions of
lower salt concentration the plants exhibit a more robust growth form. A
single main shoot or several strong shoots 30 to 100 cm. long may be pro-
duced. We observed similar habit differences for S. depressa around alkaline
lake beds in northern California and south central Oregon.

There is also some question about the effects of heat and moisture on
the degree of pubescence. Observations made by Henrickson (personal
communication) of the very pubescent shrub S. suffrutescens in northern
Mexico seem to indicate that the plants are pubescent in very dry years
and glabrous or nearly so in wetter years. The phenotypic expression in
relation to habitat conditions needs further investigation, perhaps before a
world-wide monograph on Suaeda can be attempted

’ Present address: 409 So. G St., Lakeview, Oregon 97630.

SIDA GQ) 14717. Torz,

148

Characters that are constant, and presumably reliable at the species
level, appear to be: herbaceous versus suffrutescent habit, inflorescence
types (including density of glomerules), leaf shape and internode length,
calyx features and seed size. There are five basic calyx types noted in
North American specimens (sepals thin and rounded; sepals thickened and
hooded at the apex; sepals sharply horned [corniculate] and hooded at the
apex; sepals hooded at apex and inflated vertically or keeled [carinate];
sepals hooded at apex, or corniculate, and inflated vertically as well as
basally [forming both a keel and transverse wing]).

It must be kept in mind that these plants are very succulent, so that
the calyx and leaves will flatten or shrivel upon drying. Our examination
of fresh and dry specimens has shown that the basic calyx remains constant.
The following glossary to sepal characters and Figure 1 may be helpful
in interpreting terminology related to calyx types employed in the key and
descriptions.

Carinate: keeled, with a vertical ridge or dorsal inflation from base to apex

Corniculate: horned, pointed, with one or more projecting dorsal append-
ages; may be furrowed or folded upon drying

Crispate: undulating or very wavy dried keel

Cucullate: hooded, rounded at the apex over the seed, usually thickened
and not horned nor keeled

Inflated lobe: dorsal thickening of sepal or swelling to form a vertical
ridge, basal swelling or tubercle-like projection

Rounded: not inflated dorsally nor greatly thickened at apex, usually a thin
sepal closely fitting over seed

Transverse wing: flattened horizontal projection, usually circumbasal and
irregular; a dry, basally inflated lobe.

Suaeda might be confused with other genera of Chenopodiaceae and the
following comparisons may help differentiating among them:

Suaeda: flat, spiral embryo; endosperm scant or lacking; pubescence short
or absent; leaves simple, linear; usually with three scarious floral bracts;
fruiting calyx rounded, cucullate, corniculate, or may be transversely
winged at base

Halogeton: spiral embryo; endosperm scant or lacking; pubescence pilose,
especially in inflorescence; leaves linear, tipped with a short conical

_ tubercle prolonged to a bristle; flowers bracteolate; fruiting calyx hori-
zontally winged near the apex

Kochia: annular embryo; endosperm present; pubescence simple, villous to
tomentose in inflorescence; leaves simple, linear; no scarious_ bracts
present; fruiting calyx rounded, enclosing fruit, becoming carinate to
winged horizontally toward the base, cleft no more than halfway to base

149

Bassia: annular embryo, endosperm present; pubescence pilose or tomen-
tose, especially in inflorescence; leaves linear to lanceolate; bracts one
to two, greenish; fruiting calyx cleft more than halfway to base, armed
with a short and blunt to elongated and hooked spine from the back of

each lobe.

Figure 1. Calyx types of Suaeda in North America.

_—
—

<

. Sepals thin, rounded, not inflated, as S. maritima (A).

Sepals thickened at apex, hooded (cucullate), as S. torreyana (B—fresh,
C—dried, mature).

. Sepals inflated vertically, keeled (carinate) as S. linearis (D—fresh,
E—dri e).

ied, mature

. Sepals inflated pee as well as obliquely to the base, forming a keel

and a transverse wing upon drying, as S. jacoensis and S. es
(F, G), and S. ae which is corniculate and winged (H

. ee appendaged dorsally with horn-like projections Hae as
S. depressa (J, K), and S. americana (lL, M).

150

Historically, species that were eventually named Suaeda were segregated
from the Linnaean (1753) genera Chenopodium and Salsola by Meyer (1829),
who included them in his new genus Schoberia. Moquin-Tandon (1840), in
his Chenopodearum Monographica Enumeratio, divided Chenopodiaceae into
two ‘‘suborders” and created six more synonyms for Suaeda based upon
such relatively inconstant characters as seed vertical or seed horizontal.

In 1896 Druce published the names Dondia fruticosa (Forsk.) and D. mari-
tima (Dum.), transferring them from Suaeda to revitalize Dondia Adanson
1763. The use of Dondia was retained by Heller in 1898 and 1910 with his
transfer of ten more species from Suaeda, and the name was also used by
Standley in his publication on Chenopodiales in North American Flora (1916).
In 1918 Macbride transferred Dondia back to Suaeda, and was followed by
Standley in 1929 and 1930.

In 1954 action by the International Botanical Congress conserved Suaeda
against Dondia and also against Lerchea Rueling 1774; the latter an
illegitimate orthographical variant of Lerchia A. Haller ex Zinn 1757,
which was also unavailable because of Lerchea Linnaeus 1771 nom. cons.
Dondia was a superfluous name for Lerchia, so was also illegitimate. This
congress conserved Suaeda, based on Forskal, but did not provide a generic
description. To avoid conserving a nomen nudum, Brenan (1954) proposed
the solution by amending the name and citing the first valid publication for
Suaeda, which is Scopoli (1777). As finally accepted and published by Ricketts
and Stafleu ae the ee for Suaeda is: Suaeda (Forskal, Fl. Aegypt.
Arab. 69 t. 775) ex Scopoli, Intr. 333. 1777, excl. verba falsa ‘“‘Capsula
ah ae ee oe The type species is S. vera Forskal ex J. F.
Gmelin (1791).

The first revision of North American Chenopodiaceae was published by
Sereno Watson in 1874. He delimited seven species of Suaeda, amending
and adding to his 1871 publication, which was based upon specimens col-
lected on the geological exploration of the 40th parallel under Clarence
King. Watson brought together the American-occurring species, which had
previously been placed under genera as Salsola, Chenopodium, Schoberia,
and Chenopodina. John Torrey (1828, 1843, 1853, 1857) was one of the first
American botanists to examine plants collected from western America. He
apparently found plants of Swaeda which did not exactly fit the existing
species’ descriptions. Watson resolved the problem somewhat by describing
new species of Suaeda, including S. depressa, S. torreyana, S. diffusa, S.
suffrutescens, S. californica, and S. occidentalis. He also named S. linearis
var. ramosa of northeastern America.

In 1907 M. L. Fernald published a work on northeastern American Suaeda,
including S. maritima, S. linearis var. ramosa, and two new species, S. richii,
and S. americana.

Major changes in the taxonomy of the genus occurred in 1916 with Paul
C. Standley’s work on Chenopodiales. He recognized 20 species of Dondia,

151

including D. minutiflora, named by Watson in 1883 (as a species of Suaeda),
D. insularis Britton (1906), and eight new species that he named: D. nigra,
D. fernaldii, D. mexicana, D. ramosissima, D. palmeri, D. taxifolia, D.
tampicensis, and D. brevifolia.

Three new species and two varieties from Texas and Mexico were added
in 1943 by I. M. Johnston: S. jacoensis, S. nigrescens, S. nigrescens var.
glabra, S. suffrutescens var. detonsa, and S. duripes. Recent floristic works
(e.g. Correll and Johnston [1970], Fernald [1952], Gleason [1952], Hitchcock
et al. [1969], Lundell [1969], and Munz [1974]) have included the above
species, with some lowered to the varietal level or included in synonymy.

Trends in morphology and distribution of the more than 100 species of
Suaeda were examined on a world-wide basis by Iljin (1936) who believed
the genus to have originated not later than by the beginning of the Creta-
ceous period. Iljin published a map of the range of Swaeda showing the
cosmopolitan nature of its distribution, plus a discussion of his seven
sections based upon stigma types. Three of those sections (Platystigma,
Heterosperma, and Limbogermen) can be distinguished among North Ameri-
can species of Suaeda.

The present treatment is based upon our examination of North American
specimens of Suaeda from the following herbaria: A, CSULA, DAV, F, GH,
MO, MU, NY, OSU, TEX, UC, UC-JEP, US, and UT. Several Old World
specimens were also examined from F, GH, MO, NY, and US. Curators of
these herbaria are gratefully acknowledged for loans of specimens. We are
also deeply appreciative of the loan of specimens, slides, and information
on personal observations receive from Dr. James Henrickson of California
State University at Los Angeles. Personal collections from northern Cali-
fornia and southern Oregon are deposited at MU.

Following the species description is a citation of one specimen per county
or district recorded which displays mature floral and vegetative character-
istics. Measurements and other data given in the descriptions and key are
based primarily upon dried specimens. Unless otherwise noted, all type
specimens seen are indicated.

GENERIC DESCRIPTION

Annual or perennial herbs, or suffrutescent perennials, more or less
fleshy, glabrous to short pubescent; leaves alternate, narrow and often
terete, never spine-tipped; flowers bisexual or unisexual (the plants some-
times monoecious or polygamous), sessile, in clusters of (1-) 3 (-9) in the
upper axils; bracts usually 3, scarious; calyx 5-cleft more than halfway
to the base, the lobes narrow, keeled (carinate), transversely winged, or
rounded on the back, sometimes thickly hooded (cucullate) at the apex, or
horned (corniculate); stamens 5, the short filaments usually exserted at
maturity; ovary subglobose or depressed, one-celled, the ovule solitary,
basal, the styles (2-) 3; utricle enclosed by the infolding sepals, pericarp
usually free from seed; seed horizontal or vertical; endosperm scanty or
none; embryo coiled in a flat spiral.

152

SPECIES OF SUAEDA IN NORTH AMERICA

A. Glabrous herbaceous annuals or perennials; mature sepals thin, rounded

and som

metimes inflated vertically and basally; if inflated, sepals may dry

with sharply Niobe apical ae (corniculate), rounded hoods, vertical
keels and/or transverse wings at the base.
B. One or all sepals saa Sted or corniculate, sometimes with trans-
verse wing developm
C. One or two sepals iy pointed; along north Atlantic Coast.
1.

S. americana

C. All sepals sharply pointed and/or transversely winged; interior wes-
5.

tern

D. Sepals transversely winged and corniculate; leaves narrow at

base; flowers and leaves not crowded; stems thin and flexu

D. Sepals usually not transversely winged, but corniculate;

broa ae ie base; flowers usually in dense terminal inflorescences;

branches
E. Seeds LO. 1. 3 mm. broad; naa! of interior wester n American
_ Plains d. 5. depressa
. Seeds generally larger “(12 2.0 mm, broad); plants of inland and
south central California . . depressa var. erecta

B. Sepals rounded, may be Saunas eid: Re develop transverse

g when greatly inflated to the base of the sepal.

F. Sena: rounded, not usually carinate; branches decumbent; flowers
in few-flowered ae along stem.

G. Seeds 1.8-2.0 mm. broad; leaves glaucous, acute, 0.5-5.0 cm. long:

sepals rounded, may be simone carinate; Atlantic Coast and

Q

coastal Euro ' 2. S. maritima

. Seeds 1.25-1. 5D mm. _ broad: leaves dark- -green, blunt- tipped, 0.3-1.5

cm. eeu pees rounded, not maaan, along north Atlantic Coast

of Am a 3. S. richii

F. Sepals sounded and. vine. may ice be ioieyorsely inflated or

winged at the base; branches erect; flowers in panicles or subspicate
inflorescences.

H. Sepals narrowly and evenly carinate, drying with the keel not

unevenly crispate nor transversely winged; leaves linear, acute

not acer aat nor apiculate; along Atlantic Coast, Gulf of Mexico,
West Indie 4. S. linearis

H. Sepals broadly inflated: prelonecd off obliquely at hase: drying with

transverse wing and/or crispate keel nae acuminate or apicu

a
late; primarily “of north central Mex xic

i

lomnl

Plants branching aa ie base, 3-12 a tall; ee es not opposite,
1-4 cm. long, acu or apiculate; sepals not irregularly cris-
pate on keel, nor developing corky-thickened appendages on keel
or ee seed ca. 0.8 mm. broad; plants of northern and central
Mexi 16. S. mexicana

. ee not Smanelne aenesieuously: ae the base xe 3 dm. tall; lower

leaves opposite, 1-2 cm. long, apex obtuse, ap iculate: sepals ir-
regularly crispate on keel, often also transversely wing ee and
developing corky-thickened appendages; seed 1.1-1.3 mm. broad;
endemic on Salt flats at south end of Lake Jaco, es Mexico

17. S. jacoensis

153

gee -based (suffrutescent) perennials, glabrous or pubescent; sepals
ucullate (usually pea st at apex and broadly rounded over seed, not
corniculate nor cari ).
J. Plants pubescent ork maturity, although calyx or leaves may not
be.

K. Calyx nearly or pea glabrous

L. Flowers 1-3 per axil; leaves elaucous, rounded at apex, 0.3-0.7 cm.
long; plants ee become completely glabrous with age; seed c

1.5 mm. broad. (See also under lead Q) . S. ee

L. Flowers 3-9 per axil; leaves green, acute, 0. 3- 1- 3 cm. long: leaves

t remain pubescent: seed less t

M. Flowers 1.5-2. b globose-obovoid; stem pubescent;

a. 1.0 mm. broad; along coast of southern Texas and

eastern Mexico . . 18. S. tampicensis

M. Flowers 1.0-1.5 mm. broad, globose: leaves and stem pubescen
ee ae 0.7 mm. broad; southwestern U.S. and northwestern
Mex S. suffrutescens var. detonsa
K. Entire een including calyx ‘and leaves, pubescen
Pubescence short; branches of inflorescence slender, spreading and
flexuous; leaves linear, contracted at base, not crowded; flowers

1-3 p . torreyana var. ramosissima

N. Pubescence tomentulose or villous; — stout, erect; leaves
not pnaee ted at a wded, f not, flowers 3-9 per axil.

. Flowers 2.5-3.0 m ad 1-2 we ae 1.0-2.5 em. long,

Sei leaving Renee oneen leaf ba
1. S. alioaniea var. Caius
O. Flowers 1-2 mm. br ‘oad, more ‘than 2 per axil; leaves 0.3-1.3
long paay leave ‘knobby leaf bases,
P. Leave s 0.3-0.8 ecm. long, obtuse or rounded apex, crowded, leav-
ee racnt knobby leaf bases; meas: 1-3 per axil; coastal
California and Lower California, Mexic
12.

al

S. californica var. pubescens

P. Leaves 0.5-1.3 cm. long, ‘acute, not crowded, reduced in the
oe not leaving prominent moses leaf bases; flowers
3-9 per axil, many staminate; southwestern U. : and northern
io... . ee 14. S. suffrutescens
J. Plants glabrous eee mature, or glaucous,
eaves cm. or longer, acute, gree
. Plants densely oe branches sto a leaves lanceolate, 1.5-3.0 cm

long; leaving prominent knobby leaf bases; flowers 2-3 mm. broad,
1-2 per aa seed 1.5-2.0 mm. broad.
woe ew ee ee. «10. SS. californica
R. Plants not densely leafy, branches more slender and _ flexuous;
leaves linear, acute, often reduced in inflorescence, contracted at
the base, not leaving prominent knobby leaf bases; flowers 1.0-1.5
mm. broad, usually 3 per axil; sced 0.8-1.5 mm :
8. S. torreyana
Q. Leaves 0.2-0.8 cm. long, rounded at apex, glaucous,
Plants Site glabrous, glaucous; see 1.0 mm. broad; along
Atlantic Coast of Texas and Mexico, West “Indies.
S. conferta
S. Plants pubescent with glabrous calyx when may becom
completely glabrous; seed ca. 1.5 mm. broad; an ees Mexico.
13. S. palmeri

154

1. SUAEDA AMERICANA (Pers.) Fernald, Se 9: 146. 1907.
Salsola salsa B americana Pers., Syn. Pl. 1: 296. 1805.
Chenopodium salsum 3 americanum (Pers.) Roca and Schult.,

270. 1820.

Veg. 6

Syst.

Decumbent annual, glabrous; branches 2-3 dm. long, only the abundant
flowering ones ascending; leaves green, becoming red as the plant matures
in autumn; lower lIcaves linear, acute, to 2.0 em. long, shorter and broader
in the inflorescence; flowers in dense subspicate inflorescences, ¢
having an irregular calyx with one or
hocded and corniculate than the others, occasionally with development of
transver se wing at the base; seed horizontal, 1.0-1.5 mm. broad, black at

Type: From the lower St. Lawrence River, Michaux s.n. (Paris, Photo-

Matted sea-blite, American seepweed.

Occurs in salt marshes along the Atlantic Coast, Nova Scotia to New
Jersey. Plants mature in late August to November.

TYPIFICATION: The type specimen, described by Michaux in 1803 as
Salsola salsa ? L., was collected by Michaux at the mouth of the St. Lawr-
ence River and examined by Fernald (1907) in the summer of 1903, at which
time he noted:

Loosely branching Rear: very immature, but from the crowding of
the flowers seems e the same as the Norwood Cove and Wells Beach
(Maine) species of ae September and October . . his plant of Nor-
wood Cove and Wells Beach is well characterized not aie by its late
fruiting and sub- prostrate habit but by its densely crowded flowers on
spiciform ranches, very irregular calyx, and its rich claret-color
in autumn; and it is apparently the plant intended by Persoon as
Salsola salsa, var. ? americana
In Michaux’s 1803 publication he questionably identified the specimen from
the St. Lawrence River as Salsola salsa L. and made the observation that
it differed from the typical plant of Linnaeus in its low, subdecumbent
habit with dense clusters of flowers. This was also the notation made by
Persoon when he renamed the specimen var. ? americana in 1805.

Watson (1874) included Salsola salsa sensu Michaux, from the mouth of
the St. Lawrence River, as a synonym under his Suaeda linearis var.
ramosa,. This had been treated as a synonym by Moquin-Tandon, and
others, who doubtfully placed it under the more southern S._ linearis.
Fernald’s exploration of the coast of Maine, and the lower St. Lawrence
valley, and our examination of ae specimens, have shown that the
tall, small-seeded S. linearis does not occur in central and northern Maine,
nor the maritime provinces of Canada. We have found that typical S. linearis
rarely occurs further north than New York, and in that state S. linearis
and S. americana were often difficult to separate.

—~

155

Britton apparently misapplied Persoon’s name to the southern erect
plant (S. linearis) under his combination Dondia americana (Pers.) Britton
in 1896. Fernald was apparently correct in redefining the subdecumbent
spiciform plant of the American North Atlantic coast as a new species of
Suaeda through elevation of Persoon’s variety and in employing the type
and description referred to by Persoon.

SPECIMENS EXAMINED: CANADA, PRINCE EDWARD ISLAND:
Prince Co.: Cape Aylesbury, Fernald, Long and St. John 7416, Aug. 29,
1912 (F, GH, MO, NY, UC). NEW BRUNSWICK: Gloucester Co.: Shippigan

hi

Sept. 22, 1913 (GH). Northumberland Co.: Fox I., Blake 5694, Sept. 18, 1913
(GH, US). Westmoreland Co.: Moncton, Svenson ‘and Fassett 1092, Aug. 21,
1923 (GH). NOVA SCOTIA: Colchester Co.: mouth of Salmon R., Truro,
Fernald and Wiegand 3323, Sept. 11, 1910 (GH). QUEBEC: Bonaventure,
Marie--Victorin, Rolland-Germain and Jacques 33348, Aug. 13, 1930 (GH,
NY). River du Loup Co.: Isle Verte, Lepage 13975, Sept. 2, 1957 (GH, US).

UNITED STATES, MAINE: Norwood Cove, Fernald s.n., Sept. 18, 1892
(GH). York Co.: Wells Beach, Furbish s.n., Sept. 1898 (GH). Hancock Co.:

ee Co.: Stratford, Eames S.N., Aug. 22, ie (NY, US). New Haven
Co. Haven, Co sles s.2., Aug. 12, 1892 (MU). MASSACHUSETTS:
Nantucket Co.: Nantuc ket L, Bicknell s.n., Sept. 11, 1907 (NY). Dulkes Co.:
Martha’s Vineyard, Bicknell 4023, Sept. 29. 1929 (NY), Middlesex Co.:
Cambridge, Churchill ., Aug. 27, 1912 (MO). Essex Co.: Lynn, Morong
s.n., Sept. 7, 1875 (MO, NY). Suffolk Co.: S. Boston, Dean s.n., Oct. 4, 1909
(GH). Barnstable Co.: Barnstable Harbor, Yarmouth, Fernald 452, Sept. 7,
1928 (F, GH, MO, NY, TEX, UC, US). Bristol Co.: Swansea, Touisset,
Deane and Rand s.n., Sept. 20, 1909 (GH). Plymouth Co.: Hingham ea ae
Forbes s.n., Oct. 6, 1907 (TEX, UC). RHODE ISLAND: Newpo 70.:
iia vee Greenman 1729, Sept. 27, ee (GH, MO). NEW YORK: Richmond
Co.: Staten I., Kearney s.n., Oct. 1893 (NY). Suffolk Co.: Aquebogue,
see T. “You ae S.N., sept 1873 ca NY, US). Brooklyn itt

Hdwards $.n., Sept. 3. 1939 (NY). Cape Ma ay Co.: Cape hee Witte s.n.,
Sept. 20, 1930 (NY). Ocean Co.: Pt. Pleasant, Mackenzie 4819, Oct. 2, 1910
(NY). Monmouth Co.: Keansburg, Mackenzie 4480, Sept. 26, 1909 (MO, NY).

2. SUAEDA MARITIMA (L.) Dumort., Fl. Belg. 22. 1827.
PI. 221. 1753.

Chencpodina maritima Gs Mog. in DC., Prodr. 13(2): 161. 1849.

ee ee Nutt. ex. Moq. in DC., Prodr. 13(2): 161. 1849, as
onym

Gia a ina Biome a vulgaris Moq. in DC., Prodr. 13(2): 161. 1849.

Lerchea itima (L.) Kuntze, Rev. pe 549.

Dondia ne (L.) Druce, Ann. Scot. Nat. Hist. “49. 1896.

Erect or decumbent annual (more robust than S. richii), 0.5-5.0 dm. tall;

156

leaves glaucous, linear, acute, semiterete, 0.5-3.0 (-5.0) em. long, those of
the inflorescence shorter yet much exceeding the 1-3 (-4) axiliary flowers;
sepals pale-green, rounded or obscurely carinate; seed 1.8-2.0 mm. broad,
red-brown to black at maturity.

Type locality: Seashores of Europe. Type material at Linnaean Herbarium
and at British Museum of Natural History, not seen (phototype GH!).

White sea-blite.

Coastal, mostly growing in wet salt marshes, high tidal beaches or mud
flats, Quebec to Florida, and coastal Europe.

SPECIMENS EXAMINED: CANADA, PRINCE EDWARD ISLAND:
Souris R., Marie-Victorin 9907, Aug. 20, 1919 (F, GH, NY, US). Queens Co.:
Rocky Point, Fernald, Long and St. John 7414, Aug. 31, 1912 (GH). Prince
Co.: Summerside, Fernald, Long and St. John 7412, Aug. 7, 1912 (GH, US).
NEW BRUNSWICK: Charlotte Co.: Grand Manan, Weather by and Weather-
by 7315, Aug. 14, 1944 (GH, NY, US). Westmoreland Co.: Moncton, Blake
5739, Sept. 30, 1913 (GH, NY, US). NOVA SCOTIA: Lunenburg Co.: Crescent
Beach, Zinck 1124, Aug. 29, 1941 (F). Shelburne Co.: Villagedale, Fernald,
Long and Linder 21165, Aug. 7, 1920 (GH). Halifax Co.: Purcell’s Cov
Halifax Harbor, Howe and Lang 1572, Sept. 2, 1901 (GH, NY). Yarmouth
Co.: Fernald, Long and Linder 21170, Sept. 7, 1920 (GH). Digby Co.: Digby,
Macoun 83976, Aug. 26, 1910 (GH). Annapolis Co.: ae Royal, Fernald
2620, Sept. 4, 1932 (GH). Cape Breton Co.: Cape Breton I., Nichols 1585,
July 15, 1915 (GH). Kings Co.: Port Williams, Hoag s.n., Aug. 23, 1902
(GH). QUEBEC: Carlton Co.: Bonaventure, Marie-Victorin, Rolland-
Germain and Jacques 33318, Aug. 12, 1930 (GH, MU, NY).

UNITED STATES, MAINE: Washington Co.: Kelly Point, Pembroke,
Fernald 1744, July 21, 1909 (US). York Co.: Old Orchard Beach, Ocean Park,
Wells s.n., Oct. 12, 1968 (MU). Knox Co.: Glencove, Friesner 6188, Aug. 10,
1933 ( , TEX, >, UT). Lincoln Co.: Penobscot Bay, Steyermark 2178,
July 25, 1930 (F). PENNSYL VANIA: Greenwich Pt., Philadelphia, Parker
s.n., Sept. 9, 1874 (MO, NY). MASSACHUSETTS: Norfolk Co.: Hingham,
Churchill s.n., Oct. 6, 1888 (MO). Essex, Co.: Essex, Knowlton s.n., Sept.
20, 1908 (UC). Suffolk Co.: Oak Island, Revere, Brooks 1051, Sept. 11, 1912
(UC). Barnstable Co.: Woods Hole, Peters s.n., Aug. 1884 (NY). NEW
HAMPSHIRE: Strafford Co.: cobble beach, Colony Cove, Durham Pt.,
Durham, Hogdon, Smith and Smith 1137, Sept. 13, 1941 (GH, MO, NY, OSU,
TEX, UC, US). Rockingham Co.: Hampton, Robinson 752, Sept. 29, 1901
(F, GH). RHODE ISLAND: Providence Co.: Pautucket Neck, Collins and
Chamberlain s.n., Oct. 21, 1899 (GH, US). CONNECTICUT: Fairfield Co.:
Housatonic R., Eames 72, Sept. 11, 1898 (GH). New Haven Co.: Milford,
Harger s.n., Aug 20, 1990 (GH). NEW YORK Richmond Co.: Prince’s Bay,
Staten I, Kearney s.n., Oct. 6, 1894 (F, NY, US). Nassau Co.: Long L.,
Muenscher and Curtis 6123, Sept. 1, 19388 (US). Suffolk Co.: Huntington,
Muenscher and Curtis 6118, Aug. 30, 1988 (GH). NEW JERSEY: Ocean .
Ocean City, Benner s.n., Sept. 18, 1912 (GH). VIRGINIA: Ft. Monroe,
Vasey s.n., 1879 (MO). FL ORIDA: Wakulla Co.: St. Marks Wildlife Reface
ao 60361, Sept. 30, 1960 (GH , UC). Hillsborough Co.: 12 mi. n.w.

Tampa, Double Branch Bay, Ray i Lakela 11122, July 15, 1962 (GH).

3. SUAEDA RICHI Fernald, Rhodora 9: 145.
Dondia richii (Fernald) A. Heller, ee 6: 83. 1910.

157

Decumbent annual, in mats 1-5 dm. in diameter; leaves dark-green, not
glaucous, linear to linear-oblong, blunt, subterete, dorsally compressed,
the lower leaves to 1.5 cm. long, those subtending fascicles of flowers
broader and shorter (0.3-0.5 cm. long); sepals rounded on back, not carinate;
seeds black at maturity, 1.25-1.50 mm. broad.

Type collection: Wells Beach, York Co., Maine, J. C. Parlin and M. L.
Fernald s.n., July 23, 1898 (US #1086836 !).

Rich’s sea-blite.

Occurs in salt marshes along North Atlantic Coast, Newfoundland to
Massachusetts.

SPECIMENS EXAMINED: CANADA, PRINCE EDWARD ISLAND:
Tracadie Beach, Churchill s.n., July 31, 1901 (MO). NEWFOUNDLAND:
Avalon Peninsula, Fernald, Long and Dunbar 26645, Aug. 26, 1924 (GH).
NOVA SCOTIA: Lehave R., Macoun 83977, Aug. 6, 1910 (F, GH). Picton Co.:
Loch Broom, Robinson 153, Aug. OL. 1905 (NY).

UNITED STATES, MAINE: York Co.: Wells, Deane 207, Sept. 28, 1907
(F, GH, MO, NY, TEX, UC, US).

4. SUAEDA LINEARIS (Ell.) Moq., Chenop. ae 130. 1840.
Chenopodium ae Pursh, FI. 198. 1814, not L.
Salsola linearis (El.) Sk. 1: 332. 1817.
Suaeda maritima Torr., Fl. N. Y. 2: 141. 1843, not Dumort
Chenopodina linearis (Ell.) Mogq. in DC., Prodr. 13(2): 164. 1849,
Chenopodina maritima Gray, Man. ed. 2, 366. 1856, not q.
Suaeda linearis (Ell.) Wats., Proc. Amer. Acad. yore Sci. 9: 87. 1874,
in part.
Suaeda linearis var. ramosa Wats., Proc. Amer. Acad, Arts Sci. 9: 87.
1874, excl. syn. Salsola salsa Michx. and S. salsa var. americana Pers.
Dondia americana (Pers.) Britton in Britt. and Brown, Ill. Fl. 1: 584.
t. 1393. 1896, excl. syn. Salsola salsa var. americana Pers
Dondia linearis (Ell.) Heller, Cat. N. Amer. Pl. 3. 1898.
Dondia carinata Millsp., Field Columb. Mus. Publ. Bot. 2: 297. 1909.
(Type: New Providence, Bahamas, Northrop 150, Jan. 1890, NY, not
seen. Isotype, Northrop 194, NY!).

Erect or ascending annual, someiimes persisting in warm regions, 2-9
dm. high, profusely branched, the slender branches ascending or spreading,
not decumbent; leaves narrowly linear, acute, not glaucous, dark-green, to
4 cm. long, shorter in slender elongated flowering branches; inflorescence
usually a short, loose panicle; sepals equally carinate; seed black, 1.0-1.5
mm. broad.

Type: S. Carolina, as “Chenopodium maritimum Walt. Jul-Sept. in
scirpetis maritimis’”, Walter s.n. (CHARL, not seen; phototype US!)

Southern sea-blite.

Occurs along the Atlantic Coast, New York to Yucatan peninsula of
Mexico, and West Indies.

SPECIMENS EXAMINED: NEW YORK: Kings Co.: Bergen Beach,
Svenson 6365, Sept. 28, 1934 (GH, MO, UC). Suffolk Co.: ae Long

158

I., Clute 309, Sept. 3, 1898 (NY). NEW JERSEY: Cape May Co.: Cape May,
Witte s.n., Sept. 28, 1930 (NY). Atlantic Co.: Brigantine Beach, Fogg 9535,
Aug. 26, 1935 (F). PENNSYLVANIA: Greenwich Point, Philadelphia,
Parker s.n., Sept. 9, 1874 (GH, MO, NY). NORTH CAROLINA: nels,
Co.: Swansboro, Wood 6383, July 28, 1946 (GH) Dare Co.: Hatteras, Radford

peg e and Miller 7616, Aug. 27, 1958 (GH). SOUTH CAROLINA: Beaufort
Co.: ing I., Bell 4776, Sept. 6, 1956 ae Georgetown Co.: Georgetown,

U

Harris "C19699, July 28, 1919 (¢ S). GEORGIA: McIntosh Co.: Sapelo L.,
Duncan 20565, Sept. 19, 1956 (F, GH, US). “Chatham Co.: Tybee I., Harper
47, Sept. 29, 1900 (NY, US). FLORIDA: Lee Co.: Sanibel IL. Tracy 7624,

May 18, 1905 (F, GH, MO, NY, US). Dade Co.: Matheson Hammock,
Hawkes s.n., Sept. 18, 1947 (UC). Brevard Co.: Titusville, Nash 2310, July
30, 1895 (F, GH, MO, NY, US). Pinellas Co.: Long Key, Thorne 9412, Dec.
21, 1949 (UC). Indian River Co.: Vero Beach, Lemaire 192, Apr. 6, 1957
(GH). Collier Co.: Cape Roman, Lakela and Barilotti 31047, Aug. 16. 1967
(GH). Franklin Co.: a Harbor, Kral 2784, July 18, 1956 (GH, NY).
Sarasota Co.: Sarasota Bay, Perkins s.n., Dec. 14, 1942 (GH). Wakulla Co.:
St. Mark’s Wildlife Refuge, a and Kral 54106, Oct. 6, 1955 (GH, NY).
Monroe Co.: Saddlebunch mee Moldenke — 20, 1930 (MO, MU, NY).
ALABAMA: Mobile Co.: Mon Luis I., Mohr , July 1867 (F, US). LOUISI-
ANA: Plaguemines Parish: econ L., ae ee Lloyd 492, Aug. 18, 1900
F, MO, NY, US). Cameron Parish: Cam eron, Cocks 1724, pay 1906 (GH).
Terrebonne Parish: Timbalier L., Wurzlow s.n., July 25, 1914 (NY). TEXAS:
Cameron Co.: 14 mi. from Pt. Isabe 1, Brownsville Road, F ae and Duncan
3085, Aug. 1, 1921 (MO, NY). Kenedy Co.: El Toro L, Tharp 49290, June ee
1949 (MO, NY, TEX, UC, US). Willacy Co.: Red Fish Bay, Cory s.n.,
16, 1940 (GH), Galveston Co.: Galveston Bay, Lindheimer s.n., July- res
1892 (GH, MO). San Patricio Co.: Ingleside, Cory 45323, July 25, 1944 (TEX).
Jefferson Co.: 8 mi. west of Sabine Pass, Cory 11056, Oct. 5, 1934 (GH).
Brooks Co.: Falfurgias, Innes 317, Nov. 23, 1941 (GH). Calhoun Co.: Port
Lavaca, Gentry 37, Aug. 3, 1946 (F, GH, TEX). Nueces Co.: Mustang L.,
Innes 378, Nov. 29, 1940 (GH). Brazoria Co.: Brazoria Nat’] Wildlife Refuge,
Canal, Fleetwood 9078, July 20, 1967 (TEX). Hidalgo Co.: East Donna,
Whitehouse 44238, Aug. 20, 1944 (GH). Pecos Co.: E. Escondido Spr., Tharp
and Baker 3, Sept. 29, 1943 (as S. Tharpii) (F, MO, TEX, UC, US).
MAS: Red Bay, Andros, Northrop and Northrop 45 5, Apr. 15, 1890
(F, GH, NY). North Bimini, Howard and Howard 9990, May 1948 (GH, NY).
Eleuthera, Rock Sound, Britton and Millspaugh 5563, Feb. 2 1907 (F, NY).
: Haiti, Ekman a rae 9 GH).
CUBA: Havana, Baker Aug. 26, 1904 (GH, NY). Salina de la Prin-
cipal, Cayo Romano, eee Shafer 2629, Oct. 20, 1909 (F, GH, NY, US).
eae PENINSULA: Yaxactun, Gaumer 23253, March 1916 (F, GH
US). Chichankanab, Gaumer 2219, 1916 (F, GH, MO, UC, US).
seen Schott 398, May 12, 1865 (I). Progresso, Steere 3090, Aug. 11,
1932 (IF).

UAEDA DEPRESSA (Pursh) hie Bot. King’s Expl. 294. 1871.

Salsola depressa Pursh, FI. . Sept. 197. 1814.

Chenopodium americanum nee y Spreng., Veg. 1: 922. 1825, excl.
rs

fi mM.

Chenopodium calceoliforme oe Fl. Bor, Amer. 2: 126. 18388 (Type:
“About Carlton House Fort’, T. Drummond Son, !

Suaeda calceoliformis Pane "Mogq., Chen, Enum, 128. 1840.

che wiih depressa (Pursh) Moq,. in DC., Prodr. aay. 164. 1849.

159

Schoberia se (Pers.) C. A. Meyer in Ledeb., Fl. Alt. 1: 402.

1829, as synonym.
Suaeda Wicieas Nutt. ex Moq. in DC., Prodr. 13(2): 164. 1849. (Type:
“Voleanic plains of the Missouri, Nuttall s.n., not seen, description

Schoberia calceoliformis (Hook .) Moq. in DC., Prodr. 13(2): 166. 1849.
af (Hook

nd
Dondia calceoliformis (Hook.) Rydb., Bull. Toes Bot. Club 39: 3138.
1912.

Glabrous, simple to (more commonly) freely branched, procumbent to
erect annual up to 8 dm. tall; leaves linear, mostly 1-3 (-4) cm. long,
semiterete, reduced greatly in the inflorescence (where the leaves are
mostly 2-3 mm. long), ovate to broadly lanceolate, usually somewhat
broader than the lower leaves and sometimes hiding the flowers; spikes
slender; flowers congested, 3-7 per axil in an often narrow panicle or in
narrow stout spikes arising along the main branches most of their distance;
sepals distinctly unequal, about 1.5 mm. long, all strongly hooded, often
corniculate or corrugate-corniculate transversely upon drying; seed hori-
zontal, about 1.0-1.3 mm. broad.

Type locality: ‘On the volcanic plains of the Missouri River,” Nuttall s.n.
(PH or BM ?, not seen).

Pahute weed, Pursh’s sea-blite, Pursh’s seepweed.

Occurs on usually highly saline or alkaline soil, Alaska and Yukon, south
into California, east of Cascades in Washington to Saskatchewan, Minnesota
to Texas. Matures in July to September.

TYPIFICATION: The Nuttall specimen was not located, but most of
those cited by Watson (1871) were seen, which included many plants col-
lected on the American plains and very closely resembled descriptions by
Pursh and Hooker.

All plants previously called S.
the name S. depressa, with the exception of the California specimens note
under the treatment of S. depressa var. erecta. The varietal designation is
retained for those plants because of their strictly erect habit, simpie stems,
and strict narrow inflorescences enclosed by the long, lanceolate leaves
which allow a collector to distinguish them from typical S.

depressa var. erecta were placed under

depressa.

SPECIMENS EXAMINED: CANADA, N.W. TERRITORY: Ft. Smith,
Mackenzie District, Loan 306, Aug. 21, 1950 (GH); Raup Lake, Raup 2261,
. 19, 1928 (GH). YUKON TERRIT TORY: Tagish, Calder 28347, Aug. 17,
1960 (UC). SASKATCHEWAN: Round Lake Valley, Macoun and Herriot
76792, Aug. 6, 1906 (GH, MO, NY). ALBERTA: Rosedale, Moodie 1212,
Aug. 18, 1915 (F, GH, MO, NY). ae District, Brinkman 735, July
17, 1922 (GH). BRITISH COLUMB IA: Pea R. District, Bear L.,
893, Sept. 11, 1939 (UC); Richter Pass, nes McCabe 4625,
1937 (UC); Queen Charlotte I., Calder and Taylor 35938,
MO, UC). MANITOBA: Winnipeg, Macoun 12866, Aug. 12, 1896 (NY);
Carroll, Senn and Gordon 3131, Aug. 21, 1946 (MO),

160

UNITED STATES, ALASKA: Kenai Peninsula, Calder 6681, Aug. 138, 1951
(UC); Anchorage, Murie s.n., Aug. 2, 1922 (US). MINNESOTA: Clay Ce.:
Barnesville, Moore 23639, Sept. 25, 1957 (US). Kittson Co.: Clow Township,
Moore and Moore 11576, Aug. 9, 1939 (GH). Lac Que Parle Co.: Odessa,
Gleason 9398, mae 13, 1940 (NY). MISSOURI: Sheffield (introduced), Bush
3885, May 28, 6 (GH, NY). NEBRASKA: Lancaster Co.: near Horse Cr.,
Scotts Bluff, a. 327, Aug. 1, 1891 (NY, US). Kearney Co.: Mindo on,
Hopeman 900, Aug. 1895 (GH, US). Da awson Co.: Lexington, ee 4762,
Oct. 11, 1908 (GH). KANSAS: Republic Co.: salt marsh, Sect. 29,

Morley 1002, Sept. 5, 1960 (NY, UC). Barber Co.: Hazelton, eee 15088,
Aug. 31, 1959 (US). Reno Co.: Hutchinson, Benke 2195, Oct. 7, 1918 (F).
SOUTH DAKOTA: Spink Co.: Redfield, Ricksecker 105, Aug. 10, 1908 (

Stephens 45405, Sept. 17, 1970 (NY, UC). Harding Co.: Antelope Valley,
Visher 512, Aug. 6, 1912 (F). NORTH DAKOTA: Nelson Co.: Petersburg,
Stephens 45188, Sept. 18, 1970 (NY). Lamour Co.: Edgeley, Moore and
Moore 10074, Aug. 11, 1987 (GH, NY, UC). Sargent Co.: Gwinner, Stephens
45048, Sept. 11, 1970 (NY). Ward Co.: Minot, Rider 359, July 30, 1896 (F).
Benson Co.: Leeds, Lunell s.n., Sept. 1, 1899 (MU). Grand Forks Co.:
Kellys Slough, Grand Forks, Mabbott 504, Sept. on 1917 (US). Cass Co.:
Fargo, Stevens 3044, Sept. 4, 1967 (UC, US). Pierce Co.: Lake n.e. of Petri-
fied L., Mabbott 440, Aug. 31, 1917 (US). Sioux Co.: Solen, Stevens 1385, Aug.
29, 1952 (UC, US). Burleigh Co.: McKensie Slough, Bismark, Metcalf 351,
ms 24, 1917 (MO). Mercer Co.: Ft. Mandan, Stevens 917, Aug. es 1941

US). NEW MEXICO: Chaves Co.: Roswell, Earle and Earle 311, Aug.
“a 1900 (MO, NY, US). San Juan Co.: Farmington, Standley ae July
7, 1911 (GH, NY, US). Bernalillo Co.: Albuquerque, Herrick s.n., Sept.

24, 1945 (UC). “OKL AHOMA: Cleveland Co.: Norman, Shead s.n., Sept. 17,
1954 (GH, UC). Jackson Co.: Altus, Hopkins 1012, Oct. 24, 19836 (GH, US).
Alfalfa Co.: Salt Plains Wildlife Refuge, Nighswonger im Oct. 6, 1968
(TEX). COLORADO: Alamosa Co.: Alamosa, Talbot 353, Aug. 19, 1926 (UC).
Montrose Co.: Montrose, Shear 4930, July 22, 1897 (NY, US), Fremont Co.:

Canyon City, Shear 3448, Aug. 7, 1896 (NY). Chaffee Co.: Buena Vista
Sheldon 255, July 20, 1892 (US). Larimer Co.: Ft. Collins, Cowen 2228, Sept.
3, 1893 (MO, NY). Mesa Co.: Grand Junction, Underwood and Selby 508,
Sept. 15, 1901 (NY). Park Co.: Hartsel Jet., Weber 12930, Aug. 11, 1965
(UT). Gunnison Co.: Stevens Cr., Gunnison, Hall 251, July 4, 1961 (UT).
WYOMING: Albany Co.: alkali flats, Laramie, Nelson 8677, Sept. 1902 (F,
GH, MO, NY, UC, Rts Carbon Co.: alkali flats, Hanna, Nelson 8161, Aug.
31, 1990 (GH, MO, US). Sweetwater Co.: Lost Creek, Ownbey and Lang
1087, Aug. 9, 1936 aoe Uinta Co.: Ft. Bridger, Hopeman s.n., Sept. 2, 1919
(MO). Johnson Co.: Buffalo, Tweedy 3286, Sept. 19090 (NY). MONTANA:
Big Horn Co.: Crow Agency, Hayden and Porter 67, Aug. 8, 1871 (GH, NY).
Dawson Co.: Colgate, near Glendive, Sandberg 1034, Sept. 6, 1892 (GH, NY,
US). Lewis and Clark Co.: Helena, Kelsey s.n., Sept. 21, 1891 (MU, NY).
Lake Co.: Polson, Umbach 260, Aug. 19, 1901 (F, NY, US). Philips Co.:
Malta, Alkali L., Blankinship 126, Aug. 24, 1890 (GH, US). Blaine Co.:

Havre, small 11471, Sept. 5, 1987 (MO, NY) IDAHO: Bear Lake Co.

Bear Lake, avis 1822, Sept. 1, 1987 (F). Custer " Co. : May, Hitchcock and
eens oe ae 20, 1944 (NY, UC). Owyhee Co.: Grandview, Davis 4232,
Aug. 19, 1941 (F, UC). Lemhi Co.: Salmon, Christ 5854, July 20, 1934 (NY),

161

Bannock Co.: Pocotello, Christ 8369, July 12, 1937 (NY). Franklin Co.:

Preston, Davis 4062, Aug. 21, 1941 (F). Bingham Co.: Springfield, Davis
1493, July 29, 1939 (F, UC). Gem Co.: Emmett, Christ 8550, Aug. 6, 1937
(NY). Payette Co.: Falk Station, Christ and Christ 16862, June 14, a i (NY).
NEVADA: Washoe Co.: Glendale, Kennedy 1952, Sept. 26, 1912 (F , MO,
NY, US). Elko Co.: Salt plains, Hitchcock 1123, Aug. 25, 1897 hee MO,
NY). Churchill Co.: Carson Sink, en ee 1700, Oct. 1907 (F, GH, MO, NY,
UC, US). aes Co.: northern Nevada, Griffiths and Morris s.n., July
1901 (NY). AH: Rich Co.: Woodruff Flowers F3-33, Aug. 19, 1933 (F,
UT). Cache ae McHugh s.n., July 29, 1955 (NY). Davis Co.: Farmington
Bay Refuge, Piranian and Hobson 14836, Sept. AB, 1936 (UC). Box Elder Co.:
Bear R. Refuge, Hobson 14805, Sept. 10, 1936 (UC). Salt Lake Co.: State
Prison, Arnow 3856, Sept. 30, 1973 (UT). Millard Co.: Scipio, Goodding 1506,
Aug. 13, 1935 (UT). Utah Co.: Provo, Galway and Harrison 10836, Sept. 13,
1944 (F, UC). Sanpete Co.: Ephraim, Eggleston 10261, July 25, 1941 (US).
Summit Co.: Park City, James s.n., Aug. 15, 1887 (GH). Weber Co.: sand
flat, n. of Little Mt., Arnow 803, Oct. ‘95, 1967 (UT). ase: Co.: Duchesne,
Harrison and Garrett 8889, Sept 2, 19388 Cr, UC). Carbon Co.: Saline flats,
Price, Flowers 836, Sept. 18, 1927 (UT). Grand Co.: Arches Nat’l Mon.,

Welsh and Moore 2722, Sept. 28, 1968 (NY). WASHINGTON: Whatcom Co.:

Birch Bay, Muenscher 7753, Aug. 23 1937 (F, GH, MO, NY, US). Grant Co.:

Soap ite Eyerdam 6365, Aug. 12, 1943 (UC). Lincoln Co. : Irby, Elmer 1241,
sept. 1898 (US). Okanogan Co.: alkali soil between Omak and Okanogan,
Fiker 1425, Sept. 15, 1933 (NY, US). Klickitat Co.: Bingen, Suksdorf s.n.,
Aug. 1906 (NY). Kitsat Co.: Keyport, Eyerdam 1181, Aug. 7, 1937 (MO),
Island Co.: Coupeville, Gardner s.n., 1899 (UC, UC-JEP). Mason Co.: Hoods-
port, Jones 8625, Oct. 12, 1935 (GH). OREGON: Lake Co.: Summer L.,

Coville 1257, Aug. 7, 1902 (US). Harney Co. Harney Valley, Cusick 2044,
July 18, 1898 (F, GH, MO, UC, US). CALIFORNIA: Modoc Co.: McGinty
Point, Goose Lake, Wheeler 3942, sept. 4, ag (GH, MO, NY). Siskiyou ee

Lower Klamath Lake, Hopkins s.n., Sept. 1974 (MU). Mono Co.: Bridge-

port, aay and Smith 1691, Aug. 81, 1949 UC). Santa Cruz Co.: San
Juan Bautista, Belshaw 2199, May 22, 1936 (UC). Santa Barbara Co.:
carnal Pollard s.n., Sept. 9, 1964 (TEX). Ventura Co.: Oxnard, Davy
7788, 1901 (UC). Santa Clara Co.: Milpitas, McMinn 117, Sept. 20, 1924 (UC).
Inyo Co.: Death Valley, Ash Meadows Road, Roos and Roos 6236, Sept. 14,
1954 (UT). Lassen Co.: Honey Lake Valley, Davy 3350, June 17, 1897 (UC).

6. SUAEDA DEPRESSA var. ERECTA Wats., Proc. Amer. Acad. Arts Sci.

9: 90. 1874.

Suaeda minutiflora Wats., Proc. Amer. Acad. Arts Sci. 18: 194. 1883.
(Type coll.: Santa Barbara, Calif., Cooper s.n., 1879, GH!).

Dondia minutiflora (Wats.) Heller, Cat. N. Amer. Pl. 3. 1898.

Dondia depressa var. erecta (Wats.) Heller, Cat. N. Amer. Pl. 3. 1898.

Dondia erecta (Wats.) A. Nels., Bot. Gaz. 34: 364. 1902.

Suaeda erecta (Wats.) A. Nels. in Coult. and Nels., Man. Bot. Rocky
Mts. 169. 1909.

Glabrous, 3-7 dm. high, the branches mostly simple below, erect; leaves
linear or lance-linear, broadest at base, 1.5-4.0 cm. long: inflorescence
leaves shorter and broader at base; inflorencence branches stout, short
and erect, mostly crowded; flowers clustered in axils; sepals irregularly

162

corniculate, strongly hooded; seed black, 1.2-2.0 mm. broad.
Type: Santa Barbara, Calif., E. Cooper s.n., 1879 (GH!).
Occurs in inland areas of central and southern California.

SPECIMENS EXAMINED: CALIFORNIA: Santa Barbara Co.: Sa
Barbara, Pollard s.n., Oct. 8, 1957 (NY, UC). Los Angeles Co.: Los enn
Nevin 727, 1882 (NY, UC). Orange Co.: e. of Long Beach, Bryant Ranch,
Wolf 3786, June 28, 1932 (UC, US). Inyo Co.: Amargosa Desert, Roos and
Roos 6236, Sept. 14, 1954 (GH, NY, UC, ee an san eee Co.: San
Bernardino, Parish 4200, Sept. 25, 1896 (F, MO, NY, UC, US). et
Co.: near Santa Ana R., Riverside, Hall ee 26, 1905 (I, GH, MO,

OSU, UC).

7. SUAEDA OCCIDENTALIS Wats., Proc. Amer. Acad. Arts Sci. 9: 90. 1874.
Schoberia occidentalis Wats., Bot. King’s Expl. 295. ce
Dondia occidentalis (Wats.) Heller, Cat. N. Amer. 3. 1898.

Simple and erect to freely branched and _ spreading-erect, glabrous,
yellowish-green annual, 1-3 dm. tall; branches flexuous, ascending or
spreading; leaves numerous, linear, subterete, 0.5-3.0 cm. long, 1.0-1.5 mm.
bread, acute to acuminate, reduced gradually in the inflorescence, the
leaves there little if at all broader than the lower leaves, 2-4 mm. long;
spikes ultimately open, the glomerules of 2-3 flowers not congested along
the slender branches; sepals usually distinctly unequal, one or more usually
more prominently cross-corrugate than the others, but all somewhat hooded,
and often irregularly transverse winged in age, about 1 mm. long, obtusely
lobed; seed horizontal, about 1 mm. broad.

Type: From Ruby Valley, Nevada, Watson 999, Sept. 1868 (US #49417!).

Slender sea-blite, western seepweed.

Occurs in saline or alkaline flats and marshes in the sagebrush areas of
the Great Basin, southeastern Oregon, central Washington, eastern Utah
through eastern Idaho, to southern Wyoming, Colorado. Matures in late
June to August.

SPECIMENS EXAMINED: WASHINGTON: Okanogan Co.: Tonasket,
Thompson 8688, July 2, 1937 (GH). IDAHO: Canyon Co.: Caldwell, Jensen

( REGON: Harney Co.: eastern Oregon near Nevada line,
Cusick 2031, July 12, 1898 (F, GH, MO, UC, US). Lake Co.: Silver L., Cusick
2726, Aug. 5, 1901 (F, GH, MO, NY, UC), CALIFORNIA: Lassen Co.:
Litchfield, Hoover 4641, Sept. 29, 1940 (UC, US). NEVADA: Elko Co.:
Humboldt Valley, Watson 997, Sept. 1868 (GH, NY, US). Washoe Co.:
Spanish Spring, Hendrix 876, Aug. 16, 1938 (UC). Douglas Co.: alkali flat,
Wellington Quadrangle, Johannsen me Sept. 20, 1937 (UC). Nye Co.: Potts
Ranger Station, Smith s.n., Aug. 10, 1989 (NY). Lander Co.: Austin, Goodner
and Henning 873, July 27, 1937 (TEX). UTAH: Salt Lake Co.: salt flats, Salt
Lake City, Jones 1329, Aug. 25, 1879 (F, NY). Davis Co.: west of se

992, Aug. 8, 1929 (UT). WYOMING: Gros Ventre River, Forwood $.N., July
31, 1881 (US). Lincoln Co.: at Hams Fork and La Barge, Curtis s.n., July
13, 1900 (NY).

163

8. SUAEDA TORREYANA Wats., Proc. Amer. Acad. Arts Sci. 9: 88. 1874.
Chenopodium maritimum Torr., Ann. Lyc. N. . 1828, not S.
maritima (L.) Dumort. (Type: upper part of Canadi lan R., New
i t

es 2, NY!
Chenopodium nigrum Raf., “Atl. Jour. 146. 1832. (nomen nudum
Chenopodina linearis Torr. in Stansbury, Expl. Utah 394. 1853, not C,
linearis Moq. 1849. Type: from mountains on west shore of Great Salt
Lake, Utah, Stansbury [Salt me Expedition] s.n., May 30, 1850,
NY!).

Suaeda fruticosa var. ?multiflora Torr., Pac. R. R. Rep. 4: 130. 1857.
(Type: Llano Estacado, Ft. Smith to. Rio Grande (Tucumcari), New
Mexico, Bigelow s.n., Sept. 15, 1853, US #63412!).

Chenopodina moquini Torr.., Pac. R. R. Rep. 7(3): 18. ae homon
Suaeda fruticosa Wats., Bot. King’s Expl, 294, 1871, t S. ieee
Forsk. 1775. (Type: central Wyoming, Sweetwater R. Fremont 622,

NY!

Suaeda maritima (Torr. Hs Wats., Bot. King’s Expl. 294. 1871, not S. mari-

tima (L.) Dumort.
Pia diffusa Wats., es Amer. Acad. Arts Sci. 9: 88. 1874 (Type:
m Truckee Valley, nw, ede Bailey (Watson) 996, Aug. 1867,

Pe intermedia Wats., Proc. Amer. Acad. Arts ol. 14: 296. 1879.

Suaeda moquini (Torr.) Greene, Pittonia 1:
Dondia multifiora (Torr.) Heller, Cat. N. Amer, Pi. 3. 1898.
Dondia diffusa (Wats.) Heller, Cat, N. Amer. Pl. 3. 1898.

Dondia wilsonii Millsp., Field Columb. Mus. Publ. Bot. 2: 297. 1909.
Type: an m S. Caicos I., Bahamas, Percy Wilson 7616, Dec. 14-16,
1907,

Dondia fpusicoit (Wats.) Standley, N. Amer. Fl. 21: 90. 1916.

Dondia nigra (Raf.) Standley, N. Amer. Fl. 21: 89. 1916.

m 90. 1916.
Suaeda nigra (Raf.) Macbride, Contrib. Gray Herb, n.s. 56: 50. 1918.

Perennial, glabrous, 2-8 (-10) dm. high, freely branched, ascending slen-
der branches from a short woody base; leaves linear, with a contracted
base and acute tip, 0.5-3.0 em. long, not crowded (usually at least 2 mm.
between nodes), usually reduced in the somewhat paniculate inflorescence,
although some plants have glomerules of flowers at the base of the larger
leaves below; flowers 1-5 per axil (usually 3, with the center flower larger
and functionally staminate); calyx glabrous, cucullate, the lobes cleft more
than halfway; seeds 0.8-1.2 (-1.5) mm. broad, black, shiny, vertical or
horizontal.

Lectotype (selected by Lundell, 1969): From mountains on west shore of
Great Salt Lake, Utah, Stansbury Expedition s.n., May 30, 1850 (NY!).

Torrey sea-blite, alkali seepweed.

Occurs in alkaline soil of interior plains, Oregon to Alberta, south to
Baja California, Mexico, east to Texas and West Indies. Plants mature
from July to September.

164

TYPIFICATION: Standley (1916) separated this complex of the American
plains into three species: Suaeda nigra, S. fruticosa, and S. torreyana.
Examination of types and alher cited specimens revealed no essential differ-
ences in the calyx types, floral arrangement, leaf characters, and habit.
Whether or not the species are annual or perennial is a difficult character,
since most of the type specimens are fragments of floral branches. Exami-
nation of sets of entire specimens revealed the plants were herbaceous from
a woody base. The thickness of the leaf varies with alkalinity of the habitat,
but all specimens have acute leaf tips and narrowed bases, whether or
not the leaf is very flat, semiterete, or round. Floral characters seem con-
sistent over the range, with a deeply cleft, cucullate calyx. Within a
glomcrule, the central flower is usually functionally staminate, larger, and
has acute sepals, while the two lateral pistillate flowers have more obtuse
tips. Seed characters appear consistent.

In his original publication Watson (1874) named several collections to
represent his species S. diffusa (an annual) and S. torreyana (a perennial).
Of these, Lundell (1969) selected the plant collected on the Stansbury expedi-
tion as lectotype, although the specimen does not have mature flowers. This
specimen was also noted by Torrey (1853), but misnamed Chenopodina
linearis. Suaeda diffusa was not selected as the species name since it has
been used as a synonym for S. nigra, a superfluous name, by Rafinesque as
Chenopodium nigrum, The specimen taken as type for C. nigrum was col-
lected along the upper Canadian River in New Mexico and named Cheno-
podium maritima by Torrey (1828); it is a sterile fragment. The desig-
nation S. torreyana is preferred in this treatment over S. fruticosa, an
extensively ranging Old World species, until their true relationship is
established.

SPECIMENS ee CANADA, ALBERTA: Milk River, Macoun
12934, July 10, 1895 (F, NY).

UNITED STATES, Nor DAKOTA: Billings Co.: Sullys Spring, aay
1290, Aug. 10, 1951 (US). McKenzie Co.: n. T. Roosevelt Park, Stevens 722,
is 16, 19438 (MO, UC), SOUTH DAKOTA: womans Co.: Cave Hills, Visher

Aug. 2, 1910 (F). Pennington Co.: Badlands, Canby s.n., Aug. 12, 1882
ce NY). MONTANA: Wheatland Co.: 10 mi. e. of Harlowton, "Hitehooak
2426, Aug. 1934 (MO). Rosebud Co.: Birney, Bennett s.n., July 28, 1957 (F,
NY, UC, Owyhee Co.: Bruneau, Christ 11130, June 1, 1940
(NY). Caen Co.: Caldwell, Christ 14190, July 2, 1943 (NY). Elmore Co.:
Glenns Ferry, Christ and Ward 10312, June 22, 19389 (NY). Ada Co.: Grand-

view, Davis 2062, June 18, 1940 (F). ORE GON: Harney Co.: Malheur R.,
Cusick 1939, June 18, 1898 (F, GH, MO, UC, US). NEBRASKA: Ft. Union,
Hayden s.n., 1854 (MO). WYOMING: Albany Co.: Big Hollow, Goodman
4448, July 11, 1947 (TEX). Sheridan Co.: Ucross, Nelson 9758, Aug. 1916
(MO). Fremont Co.: Green River, Shear 3346, June 1895 (NY). Sweetwater
Co.: n. Rock Springs, Ownbey and Lang 1146, Aug. 19, 1936 (MO)

RADO: Moffat Co.: Yampa R., 10 mi. w. of Maybell, Porter and Porter
3674, July 11, 1945 (GH, MO, TEX, UC). Montrose Co.: Naturita, Payson
and Payson 3880, June 28, 1924 (GH, MO, UC). Montezuma Co.: Mancos,

165

Baker, Earle and Tracy 421, July 8, 1898 (GH, MO, NY, US). Mesa Co.:
Grand Junction, Baker 925, Aug . 29, 1901 (GH, MO, NY, UC). El Paso Co.:
Manitou Springs, Engelmann es Sept. 1, 1881 (MO). Fremont Co.: n.
Canon City, Bacigalupi 1053, Aug. 29, 1924 (GH). UTAH: Box Elder Co.:
Brigham City, Zundell 247, June 19, 1910 (NY). Tooele Co.: 14% mi. s. of
Timpie Station, Arnow 511, Aug. 6, 1967 (MO, UT). Juab Co.: Troutcreek,
Maguire and Richards 2576. Aug. 27, 1933 (UC). Millard Co.: Hatton, Davis
D2572, June 4, 1930 (UT). Kane Co.: Warm Cr. Rd. near Lake Powell,
Collotzi 1046, Aug. 30, 1967 (NY). Garfield Co.: Escalente, Welsh and Welsh
9412, Sept. 5, 1969 (NY). Wayne Co.: Fruita, Maguire 19296, July 1, 1940
(NY). Emery Co.: San Raphael Bridge, Harrison 9809, Aug. 21, 1940 (US).

Salt Lake, Garrett 5009, Oct. 7, 1928 (IF, UC, Co flats,
Flowers 208, July 1924 ). Duchesne Co.: Duchesne R., yton,
elsh, urdock and Shaw 9435, Sept. 20, 1969 (NY, UT). Carbon Co

Helper, Coville and Kearney 2602, Aug. 1899 (US). Uintah Co.: 20 mi. s.
Vernal, Graham 6112, June 19, 1931 (F, MO). Sevier Co.: 7 mi. e. Salina,
Anderson 951, Aug. 25, 1958 (NY, UC). San Juan Co.: Navajo Spring, Cutler
3013, Aug. 31, 19389 (GH, MO, NY). NEVADA: Elko Co.: Salt Spring, Pilot
Mt., Holmgren 1601, July 31, 1941 (GH, NY, UC, US). Humboldt Co.:
Humboldt L., Watson 998, May 1868 (GH, NY). Washoe Co.: Pyramid L.,
Train 2605, Oct. 28, 1938 (NY). Churchill Co.: Carson L., Tidestrom 10784,
July 30, 1919 (NY, US). Nye Co.: Amargosa, Beatley 9350, July 16, 1969
NY). Esmeralda Co.: Fish Lake Valley, Archer 7948, Sept. 25, 1938 (F,
MO, NY). Lincoln Co.: Crystal Spring, Train 2412, Aug. 30, 1938 (NY).
Storey Co.: Truckee Valley, Bailey (Watson) 996, Aug. 1867 (GH, NY, UC,
US). CALIFORNIA: Lassen Co.: Honey Lake Valley, Davey 3325, June 17,
1897 (UC). Contra Costa Co.: Byron Hot Springs, Wiggins 4585, April 17, 1930
(UC). Tulare Co.: Goshen, Congdon s.n., Sept. 1881 (NY). San Joaquin Co.:
Lathrop, Congdon s.n., Aug. 26, 1887 (U C). Alameda Co.: Livermore, Howell
11441, July 20, 1933 (F, GH, MO, US). Madera Co.: Chowchilla, Hoover 1611,
Oct. 1, 1936 (UC). Fresno Co.: near Kerman Jct., Bacigalupi 2668, July 29,
1941 (GH, UC). Merced Co.: Los Banos Wildlife Refuge, Nobs and Smith 22,
July 8, 1948 (UC). Inyo Co.: Deep Spring Lake, Duran 3308, June 17, 1932
(UC). Kern Co.: Delano, Wieslander 484, May 1, 1934 (UC). Los Angeles Co.:
Long Beach, McClatchie s.n., May 30, 1891 (NY). San Luis Obispo Co.:
Cuyama Ranch, Peterson 353, July 9, 1936 (UC). San Bernardino Co.: alka-
line flats, Loma Linda, Munz and Johnston 8909, July 29, 1924 (GH, NY).
Riverside Co.: Perris, Howell 4786 ay 10, 1980 (MO). Orange Co.: New-
port Bay, Booth 1247, July 4, 1932 (UC, UC-JEP). San Diego Co.: Chula

sta, Abrams 4191, Sept. 10, 1904 (F, GH, MO, NY). Ventura Co.: Pt. Magu,
Howell 3789, May 19, 1928 (NY). Santa Barbara Co.: Cuyama Beach, French
720, Feb. 26, 1936 (UC). ARIZONA: Maricopa Co.: Guadalupe to Montezuma
Head, Killip 40848, Jan. 19, 1951 (US). Pinal Co.: Sacaton, Harrison and
Kearney 9056, Aug. 31, 1932 (F). Pima Co.: Tucson, Griffiths 2104, Nov. 15,
1900 (NY). Coconino Co.: Tuba City, Kearney and Peebles 12878, Sept. 27,
1935 (US). Navajo Co.: Winslow, Griffiths 5003, July 8, 1903 (MO). Yuma
Co.: Mohawk, Peebles 12923, Oct. 12, 1935 (US). Gila Co.: saline bottoms,
Whites Mills, Gila R., Palmer 216, Sept. 8, 1867 (GH, MO). NEW MEXICO:
San Juan Co.: Shiprock, Standley 7206, July 25, 1911 (US). Otero Co.:
Tularosa, Birum s.n., Oct. 12, 19381 (UC). Chaves Co.: Hagerman, Benke
5023, April 27, 1929 (F). TEXAS: Oldham Co.: Tascosa, Reverchon 2946,
June 24, 1902 (MO). Gonzales Co.: Tharp 78, May 21, 1936 (MO, UC).

166

Pecos Co.: Llano Estacado, Bigelow s.n., Sept. 15, 1853 (US). _ Co.:
Boca Chica, Muenscher and Muenscher 14403, Feb. 26, 1939 (N

MEXICO, SINALOA: Escuinapa, Ortega 5180, 1923 (US). aire one mi.
Punta Prieta, Wiggins 7739, Feb. 23, 1935 (F, US). SONORA: Altar, Wiggins
5992, Oct. 29. 1932 (F, US). CHIHUAHUA: Bolson de Mapini and Cienega
Grande, Saladillo, Gregg 458, Sept. 20, 1848 (GH, re

CUBA: Salinas de Caimanera, Hioram 3941, Aug. 24, 0 (NY).

BAHAMAS: S. Caicos L., Wilson 7616, Dec. ‘14. 16 1907 i GH, NY).

9. SUAEDA TORREYANA var. RAMOSISSIMA (Standley) Munz, Man. S.
Calif. Bot. 144. 1935.
Suaeda suffrutescens Wats., Bot. Calif. 2: 59, 1880, in part.
Dondia ramosissima Standle ey, N. Amer. FI. 21: 91. 1916.
ey da ramosissima (Standley) Johnston, Proc. Calif. Acad. Sci. Ser. 4.
12(30): 1017. 1924.

Densely short-pubescent throughout, otherwise similar to typical S. tor-

Type: Lees Ferry, Arizona, E. W. Nelson 62, Aug. 24, 1909 (US #564515! ).

Bush seepweed.

Occurs in alkaline areas in the interior hot dry deserts, Oregon to Nevada,
Utah, California, and northern Mexico.

SPECIMENS EXAMINED: ee ee STATES, OREGON: Maiheur Valley,
Griffiths and Morris 709, Aug. 15, 1901 (MO). NEVADA: Clark Co.: Las
Vegas, Gullion 473, May 4, 1953 (UC). Nye Co.: Amargosa Drainage, near
Spring Meadows Farm, Beatley 9794, Oct. 11, 1969 (NY). Washoe Co.: Pyra-

mid L., Tidestrom 10680, July 24, 1919 (NY, US). Douglas Co.: Topaz, Hen-
drix 1091, June 16, 1939 (UC). Humboldt Co. northern Nevada, Train 15910,
June 17, 1934 (MU). UTAH: Tooele Co.: Garrett 2773, June 17, 1914 (F, UT).
Salt Lake Co.: Salt Lake, Harrison 317H, July 12, 1939 (UC). Millard Co.:
Painter Spring, Fautin 8930, June 29, 1939 (UC). Washington Co.: St. George,
Cottam 38379, June 21, 1928 (UT). NEW MEXICO: Lincoln Co.: ¢
Tularosa, Wooton and Standley s.n., Aug. 19, 1907 (US). ARIZONA: Yuma
Co.: Yuma, oe a s.n., Sept. 20, 1906 (US). Navajo Co.: Holbrook, Rusby
382, Aug. 20, 3 (NY). Coconino Co.: Lees Ferry, Nelson 62, Aug. 24,
1909 (US). eo Los Angeles Co.: Lancaster, Elmer 3672, June
1902 (F, GH, NY). Inyo Co.: Lone Pine, Jepson 5122, July 24, 1912 (UC-JEP).
San Bernardino Co.: 1 mi. w. Old Woman Spring, Munz and Johnston 11199,
Sept. 5, 1928 (F). Fresno Co.: 23 mi. n.-n.w. of Fresno, San Joaquin R.,
Quibell 2292, June 9, 1953 (UT). Kern Co.: Bakersfield, Mastwood s.n., Oct.
4, 1894 (GH). Riverside Co.: Salton Basin, Indio, Parish 8268, Oct. 8, 1912
(GH, UC, UC-JEP). Imperial Co.: Holtzville, Keller s.n., Nov. 11, 1937
(GH, UC). Tulare Co.: Earlmont, Ferris and Rossbach 9670, May 21, 1938
(GH). Merced Co.: Los Banos, Jussel s.n. Pee 1921 (UC-JEP). Stanislaus
Co.: Modesto, Hoover 118, Oct. 20, 1934 (UC, US).

MEXICO, SONORA: Guayamas, Jones oo Nov. 6, 1930 (NY, UC).
SINALOA: Labradas, Ferris and Mexia 5229, Sept. 21, 1925 (GH). BAJA:
Loreto, Jones 27558, Oct. 16, 19830 (GH, MO, NY, UC

10. SUAEDA CALIFORNICA Wats., Proc. Amer, Acad. Arts Sci, 9: 89. 1874.
Dondia californica (Wats.) Heller, Cat. N. Amer. Pl. 3. 1898.

Stout, densely leafy, shrubby perennial, 2-8 dm. high; leaves deciduous

167

below, leaving prominent knobby leaf bases; branches ascending, stout,
but herbaceous; leaves subtercte, 1.5-3.5 cm. long, acute, lanceolate, little
reduced in the inflorescence; flowers 1 or 2 per axil, 2.0-3.0 mm. wide; calyx
deeply cleft, rounded on back, cucullate; seed 1.5-2.0 mm. broad, shining
black.

Type: San Pablo Landing, California, Bolander 412, May 1865 (GH!).

California sea-blite

Occurs in salt marshes along the Pacific Coast, San Francisco Bay, Cali-
fornia to Lower California, Mexico. Plants mature July to October.

eee EXAMINED: UNITED STATES, CALIFORNIA: Alameda
Co.: Bay Farm I., Davy s.n., Sept. 24, 1898 (UC). Sonora Co.: San Pablo
Landing, Baie 412, May 1865 (GH, NY, US). San Luis Obispo: sea cliffs,
Ferris and Ferris 7670, April 16, 1929 (UC). Santa Cruz Co.: Anacapa L.,
Clokey 4921, June 4, 1930 (NY, UC). Santa Barbara Co.: Santa Cruz L.,
ae 8 Sept. 21, 1930 (F). Ventura Co.: Ventura R., Pollard s.n.,
. 10, 2 (TEX). Los Angeles Co.: Topango Canyon

San Diego Co.: . San Diego Bay, Gander 5519, May 7, 1938 (UC).
MEXICO, BAJA: Hassler Cove, San Martin I., Moran 3062, May 5, 1948
(UG);

11. SUAEDA CALIFORNICA var. TAXIFOLIA (Standley) Munz, Calif. FI.
384, 1959.

Suaeda californica var. pubescens Jepson, FI. Calif. 447. 1914, in part.
Dondia taxifolia Standley, N. Amer. Fl. 21:
Suaeda eas Fea Standley, Publ. Field Mus. Nat. Hist. Bot.

Much-branched suffrutescent perennial, densely tomentulose or short-
villous throughout; branches very stout, 2-10 dm. long, densely leafy;
leaves narrowly linear, 1.0-3.0 cm. long, acute to acuminate, terete, those
of the inflorescence crowded and little reduced (exceeding flower clusters),
leaving prominent knobby leaf bases; flowers 1-4 per axil, 2.5-3.0 mm. broad;
calyx cucullate, densely pubescent: seed 1.5-2.0 mm. broad, black.

Type: From salt marshes, Playa del Rey, Los Angeles Co., Calif., LeRoy
Abrams 2490, June 10, 1902 (US #614215! Isotype NY!).

Woolly sea-blite.

Occurs in salt marshes along coast of California, Santa Barbara Co. to
Baja California, Mexico.

SPECIMENS EXAMINED: UNITED STATES, CALIFORNIA: Santa
Barbara Co.: Santa Cruz I., Hoffmann 546, Sept. 21, 1930 (F). Kern Co.:
Rosamond, Davy 2264b, May 9, 1896 (UC). Ventura Co.: beach, 8 mi. n

, MU . ( o
Ballast Point, Santa Catalina L., Nuttall 805, ‘Aug. 21, 1920 (F, MO). San
Diego Co.: beach at Del Mar, Peebles 315, Aug. 29, 1930 (A). Orange Co.:
Santa Anas River bridge, Newport Beach, Keller s.n., Nov. 26, 1937 (GH,
UC

F
MEXICO, BAJA: Laguna Mormona, Johnson 130, March 25, 1973 (DAV).
San Martin L, Howell 10716, Aug. 19, 1932 (F, MO, NY, UC, US).

168
12, SUAEDA CALIFORNICA var. PUBESCENS Jepson, Fl. Calif. pt. IV:
447. 1914

Dondia brevifolia Standley, N. Amer. Fl. 21: 92. 1916, not Suaeda
brevifolia aan 1895. (Type: Neer Calif., Davidson 1779, Sept.
1907 US #692799!).

Suaeda patos (Standley) Eastwood, Proc. Calif. Acad. Sci. Ser. IV.
18(14): 4

Sued brachyphit Standley, Publ. Field Mus. Nat. Hist. Bot. Ser. 8:
10. , (nom nov. for D. brevifolia Standley) not Phil. 1895.

eee ie ssp. brevifolia Abrams, Fl. Pac. States 2: 96. 1944.

Leaves short (0.3-0.8 cm.), obtuse to rounded at apex, with persistent
knobby leaf bases; flowers smaller than those of the typical species (1.0-1.5
mm. broad); entire plant covered with very short, dense pubescence.

Type: Newport, Calif., Davidson 1779, Sept. 1907 (US #692799! ).

Occurs in salt marshes along the coast of California, Sonoran Zone, Santa
Barbara Co. and south into Baja and Lower California, Mexico, and Sinaloa,
Mexico.

SPECIMENS EXAMINED: UNITED STATES, CALIFORNIA: ie
Co.: Newport Beach, Davidson 1779, Sept. 1907 (US). Los Angeles Co.
Santa Monica, Hasse s.n., Aug. 19, :

EXICO, BAJA: Magdalena Bay, Mason 1910, May 29, 1925 (GH).
SONORA: Puerto Libertad, Wiggins 6073, Oct. 25, 1932 (US). SINALOA:
Cerros de Navachiste, Gentry 14358, Sept. 26, 1954 (US).

13. SUAEDA PALMERI F cueuaa ) Standley, Publ. Field Mus. Nat. Hist.
t. Ser. 8: 193

Dondia palmeri ies N. Amer. Fl. 21: 91. 1916.

Glaucous, much branched perennial, suffrutescent; branches stout, ascend-
ing or somewhat spreading, mostly simple, densely white-tomentulose when
young, often glabrate in age; leaves terete, 0.3-0.7 cm. long, 1.5-2.0 mm
thick, rounded at apex, narrowed at base, white-tomentulose when young,
spreading, those of the inflorescence little reduced; flowers globose-obovoid,
1.5-2.0 mm. broad, 1 or 2 per axil; bracts rounded or obtuse at apex; calyx
glaucous, glabrous or nearly so, cleft to the middle or lower, the lobes
obtuse, rounded on back, cucullate, fleshy; seed usually horizontal, 1.5 mm.
broad, black; styles 3, ‘feathery’.

Type: Near Parras, Coahuila, Mexico, E. Palmer 1168, June 1880 (US
#48315! ).

Occurs in north central Mexico.

SPECIMENS EXAMINED: MEXICO, COAHUILA: 4 mi. w. of Cuatro
Cienegas, Johnston 7138, Aug. 24, 1988 (GH, TEX). ZACATECAS: Concepcion
del Oro, Henrickson 14370, Sept. 26, 1974 (CSULA).

14. SUAEDA SUFFRUTESCENS Wats., Proc. Amer. Acad. Arts Sci. 9: 88.
1874

Dondia suffrutescens (Wats.) Heller, Cat. N. Amer. Pl. 3. 1898.

Much branched suffrutescent perennial 3-10 dm. high, copiously tomen-

169

tulose throughout, the pubescence loose and spreading; branches ascending,
paniculately branched; leaves terete, mostly 0.5-1.3 cm. long, much reduced
in the inflorescence, all acute, not crowded; flowers 1.0-1.5 mm. broad, 1-6
(-9) clustered in axils, crowded in dense spikes; calyx cleft to below middle,
the lobes obtuse, relatively thin, cucullate, densely short pubescent as the
leaves and stems; seed 0.7-1.0 mm. broad, black.

Lectotype (selected by Lundell, 1969): From the Rio Grande, Maj. Wm.
H. Emory s.n., Oct. 6, 1846 (NY!)

Occurs in alkaline gypseous plains and valleys, western Texas, New
Mexico, Oklahoma, Arizona, south into northern Mexico.

SPECIMENS EXAMINED: UNITED STATES, OKLAHOMA: Major Co.:
Orienta, Waterfall 7544, July 11, 1947 (MO, NY, TEX). Tillman Co.: Hollis-
ter, Waterfall 7296, June 27, 1947 (MO, TEX). TEXAS: Crane Co.: Buena
Vi sta , Warnock 15608, July 13, 1957 (TEX). — Co.: Ft. Stockton, Mears
1546, ‘May 28, 1967 (TEX). Reeves Co.: Pecos to Barstow, Tharp and Gim-
bride 51-994, July 7, 1951 (TEX). ane Co.: gypsum soil, east of
oo Waterfall 5189, July 20, 1948 (GH, MO, NY). Hudspeth Co.:

t. Quitman, Cory 31045, Oct. 24, 19388 (GH). El Paso Co.: El Paso, corner
- Country Club Rd. and Montoya, Blackwell 2329, July 14, 1976 (MU).
Jeff Davis Co.: Chispa, Waterfall 5319, July 26, 1943 (F, GH, :
eg Co.: along Rio Grande R., Porvenir, York 48249, July 7, 1948

EX). Brewster Co.: Lajitas, Warnock 953, July 17, 1937 (GH, TEX, US).
aa Co.: east of Pecos, Waterfall 5504, Aug, 6, 1943 (MO, NY). NEW
MEXICO: Chaves Co.: Roswell, Waterfall 4314, Aug. 23, 1942 (GH). Eddy

o.: Carlsbad, Standley 40337, Aug. 12, 1924 (US). Lincoln Co.: White Mts.,
Wooton 191, July 22, 1897 (GH, MO, NY, UC, US). Dona Ana Co.: sandy
bank at end of Camino Real just beyond El Paso city limits, Blackwell 2334,
July 14, 1976 (MU). Otero Co.: White Sands, Alamogordo, Fisher 54, Ang.
6, 1931 (F). Catron Co.: Bat Cave Wells, Smith 63, July 2, 1948 (GH).
ARIZONA: Navajo Co.: Holbrook, Zuck s.n., July 10, 1897 (NY, US).

MEXICO, COAHUILA: Laguna del C oyote, Henrickson 14269a, Sept. 22,
1974 (CSULA). Laguna de Leche, cee 8629, Aug. 30, 1941 (GH, MO).
CHIHUAHUA: Rio Grande, Paso del Norte, Pringle 1997, Sept. 9, 1888 (F,
MO, NY, UC, US).

15. SUAEDA SUFFRUTESCENS var. DETONSA Johnston, Jour. Arnold

Arb. 24: 230. 1943.
Suaeda nigrescens Johnston, Jour. Arnold Arb. 24: 228-229. 1943. (Type:
from La Ventura, Coahuila, Mexico, Johnston 7650, Sept. 12, 1938.

Suaeda nigrescens var. glabra Johnston, Jour. Arnold Arb. 24: 228-229.
1943. (Type: from Ft. Quitman, Hudsp eth Co., Texas, Waterfall
3993a, Aug. 18, 1942 GH!).

Leaves glabrous, green to glaucous, 0.3-1.3 cm. long, obtuse to acute
tips; stems more or less pubescent, the leaves and calyx usually glabrous;
other characters like typical S. suffrutescens.

Type: From 3 mi. west of Cuatro Cienegas, saline flats, loosely and
widely branched, 1-5 ft. tall, frequently supported by other bushes, Johnston
7128, Aug. 24-26, 1938 (GH!).

170

Occurs in saline gypseous soils in trans-Pecos Texas, along the Rio Grande
and Pecos valleys in adjacent New Mexico and south into southern Chi-
huahua and Coahuila, Mexico.

DISCUSSION: It is difficult to distinguish between this variety and
specimens identified as S. nigrescens and its variety glabra, since leaf
characters, inflorescence, branching patterns and distribution are very
similar. There is some difference in degree of stem pubescence, and it may
be shown later that S. nigrescens and S. nigrescens var. glabra are distinct
from S. suffrutescens var. detonsa genetically. Until these differences, if
they exist, can be shown taxonomically, doubtful specimens will be placed
under S. suffrutescens var. detonsa. The flowers are also more densely
clustered than on typical S. torreyana, with which members of this variety
might also be confused.

SPECIMENS EXAMINED: UNITED STATES, TEXAS: Gonzales Co.: 50
mi. e. of San Antonio, salt flats near Pilgrim, Cory 19213, Aug. 23, 1936
(GH). Brewster Co.: Tornillo Creek, Hot Springs, Sperry 1637, Mar. 26, 1939
(GH). Andrews as ae L., Tharp s.n., July 10, 1941 (TEX). Hudspeth
Co.: Rio Grande, Ft. Quitman, Waterfall 3993a, Aug. 18, 1942
NEW MEXICO: Oe Co.: White Sands, Archer 7334, Oct. 19, 1938 (GH).
Chaves Co.: Hagerman, Benke 5023, Apr. 27, 1923 (GH).

“XICO, COAHUILA: Monclova to Cuatro Cienegas, Johnston 7128, Aug
24-26, 1938 (GH). SAN LUIS POTOSI: alkaline flat, San Miguel, aes
7619, Sept. 11-12, 1938 (GH). CHIHUAHUA: Meoqui, LeSeuer 197, Aug. 24,
1935 (GH, TEX). ZACATECAS: west of Guadalupe Carceron, Johnston,
Wendt and Chiang 11597, July 3, 1973 (CSULA).

16. SUAEDA eae (Standley) Standley, Publ. Field Mus. Nat. Hist.
Bot. Ser. 4: 208.
Dondia mexicana aay N. Amer. Fl. 21: 89. 1916.

Annual herbs, glabrous, 3-12 dm. tall, mostly branched at the base;
branches numerous, ascending or nearly erect, elongate; leaves numerous
but not crowded, linear, semiterete, the lower leaves 1.2-4.0 cm. long, 1.2
mm, broad, acuminate or attenuate, those of the inflorescence shorter;
inflorescence paniculately branched, the branches erect; flowers crowded
in the axils and forming spikes 4-5 mm. thick; calyx deeply cleft, the lobes
rounded, becoming more enlarged at the base, strongly cucullate-carinate
in age, the fruiting calyx 3 mm. broad; stamens exserted at maturity;
seed ca. 0.8 mm. broad, horizontal, shining dark brownish-red; styles 2,
‘feathery’.

Type: From San Luis Potosi, Hacienda Angostura, alkaline plain near
San Bartolo Station, Mexico, C. G. Pringle 3788, July 15, 1891 (US #48314!).

Mexican seepweed.

Occurs in western Texas, south into San Luis Potosi, Mexico.

SPECIMENS EXAMINED: MEXICO, COAHUILA: 3 mi. west of Cuatro

Cienegas, Henrickson 14292, Sept. 23, 1974 (CSULA). SAN LUIS POTOSI:
Las Tables, Pennell 18047, April 9-10, 1934 (GH, US).

171

17. SUAEDA JACOENSIS Johnston, Jour. Arnold Arb. 24: 228. 1948.

Annual or perennial succulent, 1-3 dm. tall, glabrous, pale green, branch-
ing from small caudex crowning a coarse persisting root; stems slender,
somewhat purplish verrucose (warty), numerous, erect, rigid, simple or
sparsely branched; leaves linear, 1-2 cm. long, ca. 1.2 mm. wide, com-
pressed succulent, the apex obtuse, apiculate; the lower leaves conspicu-
ously opposite, the others alternate; flowers axillary, subsessile, clusters
distant (usually 2-3 mm. apart), subspicate; bracts hyaline, triangular or
oblong, dentate or inconspicuously lacerate-dentate; calyx in fruit irregular
crispate and carinate, some almost appendaged with corky tubercles, or
with a vertical as well as a transverse basal wing that may be corky-
thickened and prolonged off obliquely at the base; seed horizontal, 1.1-1.3
mm. broad, shiny black-brown; styles 2, ‘feathery’.

Type: From salt flats at southeastern end of Laguna de Jaco, common,
western Coahuila, Mexico, Stewart and Johnston 1975, Oct. 4-5, 1941 (GH!).

Apparently an endemic species on flats at southeastern end of Laguna
de Jaco, Coahuila, Mexico, where the soil is strongly saline and gypsiferous.

SPECIMENS EXAMINED: MEXICO, COAHUILA: Laguna de Jaco, Hen-
rickson 14215, Sept. 20-21, 1974 (CSULA).

18. SUAEDA TAMPICENSIS ao Standley, Publ. Field Mus. Nat.
Hist. Bot. Ser. 8: 10. 1930
Dondia tampicensis Standley, N. Amer. Fl. 21: 91. 1916.

Green, much branched perennial, the branches stout, ascending or
decumbent, suffrutescent, copiously short-villous, paniculately branched;
leaves terete, 7-15 mm. long, acute or acuminate, spreading, those of the
inflorescence little reduced; flowers 1.5-2.0 mm. broad, globose-obovoid,
1-5 per axil; bractlets acuminate; calyx glabrous or nearly so, cleft to
below the middle, the lobes rounded at the apex, fleshy, cucullate; seed
usually horizontal, 1 mm. broad, black, shiny.

ype: Along the coast near La Barra, Tamaulipas, 8 km. east of Tampico,
Mexico, Edward Palmer 262, Feb. 1910 (US #463200! ).

Occurs in coastal sandy areas, southern Texas and eastern coast of
Mexico, also in the West Indies.

SPECIMENS EXAMINED: UNITED STATES, TEXAS: Cameron Co.:
mi. east of Brownsville, Correll and Johnston 17957, July 13, 1957 re
Starr Co.: Rio Grande City, Tharp 5783, June 16, 1928 (TEX, US). Kenedy
Co.: Padre I., Tharp 5567, June 1922 (TEX).

MEXICO, TAMAULIPAS: Matamoras, Wooton s.n., June 29, 1919 (US).

DOMINICAN a i. Isla de Cabritos, Laguna de Enriquillo, Ekman
9860, April 16, 1928 (GH, US).

HAITI: Bocazelle, een Plains, Sweet s.n., 1925 (NY, US).

19. SUAEDA CONFERTA (Small) Johnston, Jour, Arnold ee - 230. 1943.
Dondia conferta Small, Bull. N. Y. Bot. Gard. 1: 280

172

Lerchea conferta (Small) K. Schumann in Just., Bot. Jahrb. 27(1): 482

Dondia insularis Britton, Bull. N. Y. Bot. Gard. 4: 138. 1906. (Type:
from Grand Turk I., Bahamas, Nash and Taylor 3873, Aug. 27-Sept.
1, 1905, US #847951! ),

Perennial suffrutescent shrub, stems erect or ascending, 4-10 dm. long,
the branches prostrate or spreading, forming dense tufts, very brittle,
glabrous; leaves numerous, glabrous, the leaf-blades fleshy and blue-gray
(glaucous), mostly less than 1 cm. long, 1-2 mm. broad, oblong; flowers
solitary or clustered in the axils of the rather approximate leaves, especially
numerous on the branchlets; calyx lobes obtuse, cucullate, seed about 1
mm. broad, black.

7 From coastal flats, Corpus Christi, A. A. Heller 1827, June 2-6,
1894 (US #213983! ).
Along the Atlantic Coast of Texas and Mexico, also in the West Indies.

SPECIMENS EXAMINED: UNITED STATES, TEXAS: Nueces Co.:
Corpus Christi, Young s.n., Mar. 14, 1914 (MO, TEX). San Patricio Co.:
Ingleside, Cory oo July 25, 1944 (GH). Cameron Co.: Yucca I., Johnston
ae Dec. 23, 1954 (TEX). Hidalgo Co.: Clover 1189, May 24, 1933. Kenedy
Co.: Padre L, Cutak 29, July 26, 1938 (MO). Aransas Co.: Rockport, Rever-
chon 1788, July 1893 (MO). Kleberg Co.: Laguna Madre, King Ranch,
Johnston s.n., July 3, 1953 (TEX).

AITI: Plains de la Artibonite, Ekman H3342, Feb. 27, 1925 (GH

DOMINICAN REPUBLIC: Santo Domingo, Isla de Bauritas, Ekman
ee Apr. 16, 1928 (A).

AHAMAS: Grand Turk L, Nash and Taylor 3873, Aug. 27, 1905 (F, US).

DOUBTFUL AND EXCLUDED SPECIES

1. Suaeda duripes Johnston, Jour. Arnold Arb, 24: 231. 1943. (Type: from
exas, Pecos River, salt soil, Geo. Thurber 114, Nov. 1850 GH!).
= ? S. suffrutescens var. detonsa. Known only from the type collection,
cava by Johnston (1943) to be endemic to Reeves and Pecos Cos., Texas
2. Suaeda fernaldii (Standley) Standley, Publ. Field Mus. Nat. Hist. Bot.
Ser. 4: 203. 1929. (Type: from red sandstone alluvium near mouth of
Salmon River, Truro, Colchester Co., Nova Scotia, Fernald and eile
3324, Sept. 11, 1910 GH!, fragment and photo US!, photo NY!), a
Dondia fernaldiit Standley, N. Amer. FI. 21: 88. 1916. Known only ae
the type collection. It may be a local variant of S. americana.

REFERENCES
BRENAN, J. P. M. 1954. Proposal #157, correction to #2261. Suaeda. 3: 65.
BRITTON, N. L. rae Contribution to the flora of the cae | TIL ‘Bull, N. Y.
Bot. ae 4: 136-14
and A. BROWN, 1896. Illustrated flora of the United States, Canada and the
British possessions. New York.
CORRELL, D. S. and M. . eee Smee 1970. Manual of the vascular plants of Texas.

Texas Recoureh: Foundation, Renner, Tex

TORREY, J. 1828.

7 of New York

UNGAR, I
. J. and — 4 Queen, eds. Academic Press, New
wo S. 1871. vy
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= 8
Arts. See 9:
18

173

DRUCE, G. C. 1896. Notes on the ninth edition of the London catalogue of British plants
Ann. Scot. Nat. Hist. 5: 42.
FERNALD, M. L. 1907. The genus Swaeda in northeastern oe oc 9:
1952. wee manual of botany. Ninth ed. American Boo w York.
GLE ASON, H. A., ed. 1952. The new Beiteon: am hae mn lensed fe i: norehesscrn
United States and adjacent Cana . Ve - aster Lancaster
GMELIN, J. F. 1791. Caroli 4 Lin ema natura . - eerie. 2: a
R, _ A. 1898, Coe of North y eneeae meas moet of Mex
10. New eee III. deep acieccs 6: 83.
ircrecce C..Ls NQUIST, M. EY, and J. W. THOMPSON. 1969.
eee plants of fang: ee Northwest. Univ. Wash. Press. Vol. 17 of Univ. Wash.
Sea

Publ.
TEIN; aM. M. 136 Contribution a i a ae du genre Suaeda et de la tribu Suacdeae.
Sove nik. Leningrad. -49

43

140-146.

>

ttle.

arene species from Mexico and adjacent United States.
ae Arn ae res 24: 227-23

1.
ae es ae Ae 1753. Species plentaeu m Ud:
LU? , ed. 1969, A flora ofa s Research ee Renner, Texas.
cae 7 0 8. Pees American sperm: ie ce new or rred. Contr. Gray
Herb. n. s. :
MICHAUX, i‘ cane mee Boreali-Americana. Vol. 1. Paris.
MOQUIN-TANDON, C. B. A. 18

1840. Chenopodearum monographica enumeratio. Pari
NZ, P. A. 1974. A as of southern California, and supplement. Univ. of Calif. fee.
if,
PERSOON, C. H. 1805.

Synopsis plantarum.
RICKETTS, H. W. and F.

5-296.
S, A. STAFLEU. 1960. Nomina generica conservanda #2261.
Suaeda. Taxon 8: 267.
SCOPOLI, J. A. 1777. Introductio ad historiam naturaiem erle, Pragae.
ST ee i Ae 1916. Order Scenic teen in’ North Nero Flora. 21(1): 86-92.
. Studies of American plants. I. Publ. Field Mus. Nat. Hist.
Ser. 4: 3 Sia

Chicago. Bot.

—. 1930. Studies of American plants. HI. Publ. Field Mus. Nat. Hist. Chicago.
0.

Some account of a collection of plants made during a journey to and

from = Rocky Mts. in the summer = a, by Edwin P. James, M.D. Asst. Surg.,

U ao aa Lyc. Nat. Hist. Y 39.

. A flora of the state er ca

York. Vol. 2. (Part 2 of Natural History

otany (Appendix D) in Howard Stansbury. 1852. An expedition to the

valley of he Great $ Sale Lake of Utah. Sampor- ne Son and Co., London.

2 Reports of explorations and surve ain the most practicable

and a route for a ea from i Miss. R. to the Pacific Ocean. Route near
h p eae 5 = red by Lt. A. Whipple, 1853-1854. 4: 130.

. A. 1874. Tnland halophistes of ce U

to ascert

in "Ecology of halophytes. Reimold
York.

Sy 294-295.

A revision of the North American Chenopodiaceae. Proc. Amer. Acad.
82-126.

. Contributions of American Botany. Proc. Amer. Acad. Arts Sci. 18: 194.

TAXONOMY AND DISTRIBUTION OF
GENTIANA (GENTIANACEAE) IN MEXICO AND
CENTRAL AMERICA. I. SECT. PVEUMONANTHE'

JAMES S. PRINGLE
Royal Botanical Gardens, Hamilton, Ontario, Canada

This paper is the first of two that together will constitute a taxonomic
monograph on Gentiana L. in Mexico and Central America. No such mono-
graph has previously been published, nor does any modern floristic work
provide for identification of all these species.

Following Gilg’s (1895) treatment of the Gentianaceae, as modified by
Wagenitz (1964), Gentiana is included in the tribe Gentianeae, subtribe Gen-
tianinae, Tribal and subtribal traits have been given by these authors.

The circumscription of Gentiana accepted here follows that in publications
on the Gentianaceae by Smith (1936; 1967), Fabris (1960), Gillett (1963),
Toyokuni (1963), Mason & Iltis (1966), Nilsson (1967), Tutin (1972), and
Weaver & Rudenberg (1975). As thus delimited, Gentiana is distinguished
from other genera in the Gentianinae by the following combination of char-
acters: calcium cxalate crystals in the leaves (except in sect. Calathianae
Froel.); a continuous intracalycular membrane (except in at least part of
sect. pele Cc as ea three primary veins per petal; appendages,
or plicae (discussed below), between the primary divisions of the corolla
(except in G. ot ae basifixed anthers; naked nectar-secreting glands
borne on the gynophore; septicidal, capsular fruits; no fringes, scales, or
nectaries on the interior surface of the corolla; and no marginal fringes on
the corolla lobes. This circumscription of Gentiana is narrower than that in
most North American floras in that it excludes those North American species
included by Gillett (1957) in Gentianella Moench. It is, however, more in-
clusive than that of Love & Léve (1975), who restricted Gentiana to the five

a na

are treated as distinct genera, the species discussed in the present paper
would be placed in the genus Tretorhiza Adans.

Although a complete generic description of Gentiana is not included here,
it does appear to be necessary to discuss certain features of the corolla
because of their unusual nature and because of the diversity of terms which
have been used in describing them. The corolla of Gentiana consists of five
(occasionally four or six, rarely more) united petals, each of which termi-
nates in a corolla lobe. Within the corolla tube, each of the primary divisions

‘Contribution No. 22 from the Royal Botanical Gardens, Hamilton, Ontario, Canada.

SIDA 7(2): 174—217. 1977.

175

is separated from the next by a triangular segment of corolla tissue, which
is invaginated in bud and to some extent at anthesis, and which is vascular-
ized only distally by branches from the lateral veins of the two adjacent
petals. These structures are called ‘‘appendages’”’ in this paper, following
such works as the Flora Europaea (Tutin, 1972); in some other publications
on Gentiana, they have been termed “‘plicae”’ or ‘“‘plaits

The corollas of Gentiana, dextrorsely contorted in bud, emerge from the
calyces long before anthesis. Until the corolla opens, part of each lobe is
overlapped by part of the next lobe clockwise, except in a few species with
greatly fen lobes. The margin of the corolla lobe exposed in bud is
termed the in’? in this ae and the other margin is termed
the ‘‘inner margin.’’ This use of ‘‘outer’’ and
from references to the exterior and interior (i.e., abaxial and adaxial) sur-
faces of the corolla.

Kusnezow’s (1894) monograph of Gentiana ‘‘subgenus Eugentiana’’ and
his (1895) treatment of Gentiana in Engler & Prantl’s Die natiirlichen Pflan-
zenfamilien provide the generally accepted basis for the division of Gentiana
into sections, although some later authors have proposed refinements. Kusne-
zow’s “‘subgenus Engentiana’’ is $ equivalent to Gentiana as the genus is Cir-

This paper deals with the taxonomy and distribution of sect. Pneumonanthe.
Key to the Sections of Gentiana in Mexico and Central America
la. Small plants, usually less than 8 cm tall, with slender, fibrous roots;
leaves less than 1 cm long; flowers less than 2 cm in length or diameter;
free portions of corolla appendages nearly symmetrical, 2/3 to fully
as long as lobes; gynophore slender, elongating in fruit so that the ovary
is largely or completely exserted from the marcescent corolla.
Ya ae sect. Chondrophyllae.
. Plants usually well over 8 cm tall; os — large and persistent, or re-
placed by several stout, fleshy secondary roots; larger leaves over 2 cm
long; flowers over 2 cm long; free portions of corolla appendages more
or less oblique, much exceeded by lobes (in Mexican and Central Amer-
ican species); gynophore variable, but not elongating sufficiently to ex-
sert the entire ovary from the marcescent corolla. sect. Pneumonanthe.
GENTIANA [sect.] PNEUMONANTHE Gaudin, Fl. Helv. 2:269. 1828. Type

species (Art. 22, ICBN): Gentiana pneumonanthe L.

Pneumonanthe Gled. (pro gen.), Syst. Pl. p. 238. 1764. Type species (only
species definitely included): Gentiana pneumonanthe L., as Pneumo-
nanthe [sp.].

Gentiana Pneumonanthe Link (sine ord.), Enumeratio Pl. Hort. Reg.
1:258, :

Gentiana subgenus Pneumonanthe Raf., Med. FI. 1:208. 1828. Type species
(Art. 22, ICBN): Gentiana pneumonanthe L

—
io”

176

Xolemia Raf. (pro gen.), Fl. Tellur, 3:22, ‘‘1836’’ [1837]. Lectotype species
(hoc loc.): Gentiana saponaria L.

Xolemia subgenus Cutlera Raf., Fl. Tellur. 3:23, ‘1836’’ [1837]. Lectotype
species (hoc loc.): Gentiana catesbaei Walt., as Xolemia catesbei [sic]
(Walt.) Raf,

Gentiana sect. Cyane Griseb. in Hook., Fl. Bor.-Amer. 2:54. ‘1840’ [1837].
Type species (Art. 7, ICBN; see Pringie, in press): Gentiana pneumo-
nanthe L.

Perennials. Roots either several-many, thick, fleshy, and individually usu-
ally of short duration, or, in other species, consisting of a stout, persistent
tap root and several smaller secondary roots. Rhizomes short and erect
(geocorms). Stems I-several, terete, generally over 5 cm long and in some
species often over 1 m, ascending to erect, or in some species commonly
decumbent at base, usually unbranched or with short, peduncle-like branch-
es only. Rosettes of leaves absent except in seedlings. Leaf and calyx-lobe
margins entire, minutely denticulate, or ciliolate, in most species neither
cartilaginous nor scarious, Flowers erect (or somewhat declined in G. cali-
culata Lex.), 2.5-7.5 cm long. Calyx tubes either 15-nerved or 10-nerved
through fusion of lateral sepal veins, cylindrical to vase-shaped, or in some
species either occasionally or regularly cleft and spathelike. Intracalycular
membrane generally well developed. Corollas ranging from completely
closed to wide open at anthesis, open corollas closing at night and in cloudy
weather but not thigmonastic. Corolla tubes cylindric to funnelform, vase-
shaped, or urceolate. Corolla lobes shorter than tubes, deltoid to ovate-
elliptic, or, in G. andrewsii Griseb., reduced to mucros. Free portions of
corolla appendages usually well developed, basically bifid, usually oblique,
with the sinus next to outer margin of each corolla lobe being deeper than
the one next to inner margin, and the appendage segment next to the shal-
lower sinus being narrower, but sometimes longer, than the other segment;
in some species with the segment next to the inner corolla-lobe margin
greatly reduced and deflexed, appearing as a projection from the lobe, or
obsolete. Free portions of appendages in a few species symmetrical but well
developed, in a few other species reduced to 2 small teeth or nearly obso-
lete. Summits of appendages (when well developed) ranging from subentire,
erose, or minutely serrate (in species with one segment reduced) to shal-
lowly to deeply laciniate, or shallowly fimbriate to deeply setose-fimbriate.
Corollas of most species predominantly dark to light blue or violet-blue,
with occasional forms having lilac, rose-violet, or white corollas; corollas
of some species regularly violet, white, yellowish-white, or greenish-white;
those of G. caliculata Lex. and G. mirandae Paray red. Darker stripes
usually present on the interior surface of the tube, between primary veins
of petals. Stamens equal, with straight filaments. Anthers connate (in all
species with closed or nearly closed corollas, and in some species with open
corollas) or free. Pollen grains oblate-spheroid to prolate, with sexine dis-
tinctly and often coarsely striate. Ovary compressed lance-ovoid, gradually

177

tapering into a short style, the style not sharply demarcated at anthesis
and not spirally twisted (except tardily in G. caliculata). Stigmatic lobes
linguliform or in a few species more slender, recurving. Gynophore rela-
tively short and stout, usually 0.5-1.5 cm long and 1.5-2 mm thick at anthesis
(longer and more slender in G. mirandae), elongating somewhat in fruit
but not so as to elevate the entire ovary above the marcescent corolla. Car-
pels each with 1 dorsal and 6 ventral vascular bundles. Ovules distributed
all over interior surface of ovary except in the immediate vicinity of su-
tures. Seeds with an obscurely to prominently reticulate testa, the testa
in most species being expanded into a membranous wing partially or com-
pletely surrounding body of seed. 2n = 26 in all taxa for which chromosome
counts have been published. Sectional descripticn compiled from Kusnezow
(1895), Lindsey (1940), Toyokuni (1963), Nilsson (1967), standard references
to published chromosome counts, and my own observations.

The foregoing description does not provide for G. asclepiadea L. because
various lines of evidence suggest that this species may be misplaced in
sect. Pneumonanthe. The generic name Dasystephana Adans., typified by
G. asclepiadea, has therefore been omitted from the synonymy, along with
the illegitimate modification Dasistepha Raf. Gentiana newberryi A. Gray

and G. tiogana Heller have likewise been excluded from consideration, since
their inclusion in this section also appears to be questionable.

The generic names Ciminalis Adans., Diploma Raf., Ericoila Borkh, (nom.
illeg., altered to Ericala by some authors), Hippion F.W. Schmidt (nom.
illeg.), and Tretorhiza Adans. cmend, Live & Love are typified by species
in other sections of Gentiana. Therefore, they have been excluded from the
synonymy of sect. Pneumonanthe, although some AUTHORS have transferred
species in sect. Pneumonanthe to these ‘genera.’

Considerab‘e variation exists in the orthography of the name of this sec-
tion and in the authorship to which it has been attributed. Reasons for citing
Gaudin (1828) as the author of the name Gentiana sect. Pnewmonanthe have

een given previously (Pringle, in press)

The formal subdivision of sect. Pnewmonanthe, which would require con-
sideration of variation patterns throughout its range, is not within the scope
of this paper. The Mexican and Central American species of this section
can, however, be informally divided into two groups. The first group, com-
prising G. bicuspidata, G. ovatiloba, and G. hooperi, is characterized by
irregular, unpaired branching, if any, and by ebractcolate flowers. The
other group, which includes G. spathacea, G. laevigata, and G. caliculata,
differs in having isocladous branching, if any, with the flowers being borne
in symmetrical, usually condensed cymes, and in the presence of paired
bracts subtending each flower. Gentiana mirandae, because of the paired
bracts subtending the flowers, can also be included in the latter group,
although its branching is not always strictly ae dou

In its entirety, sect. Pneumonanthe comprises about e species, native to
North America from Guatemala north to Alaska and Labrador; to most of

178

Europe, except for the warmer, lowland regions; and to much of Asia,
including the Caucasus, the Himalaya, central and southern Soviet Asia,
central, southern, and eastern China, and Japan. Throughout the range of
this section, its representatives are largely confined to humid or subhumid,
warm-temperate to boreal regions. Those native to tropical latitudes, in-
cluding the Mexican and Guatemalan species, are restricted to high eleva-
tions.

In the present paper, references to the physiographic provinces in which
Gentiana species occur follow Raisz (1964). References to vegetational zones
in Mexico follow Leopold (1950) and Flores Mata et al. (1971). Most Mexican
species of sect. Pneumonanthe occur in the coniferous forest (bosque de
coniferas) of Flores Mata et al. Its lowest subdivision, the pine-oak forest
(bosque de pino-encino) of both Leopold and Flores Mata et al., is extensive
in Mexico, occupying much of the higher terrain in the Sierra Madre Occi-
dental, Sierra Madre Oriental, Neovoleanic Plateau, and Sierra Madre del
Sur. It was further subdivided by Leopold, with the pine-oak woodland prob-
ably being the most important subdivision as a habitat of species in sect.
Pneumonanthe, although they also occur in openings in the pine forest.
Gentiana species are also present in the vegetational zones at higher alti-
tudes on the Neovoleanic Plateau, including the transitional pine-alder-fir
forest of Leopold (ca. 2200 m to 2300-2600 m), the fir forest (bosque de
oyamel) (up to 2900-3200 m), and the relatively open Pinus hartwegii Lindl.
forest (bosque de pino), which extends to treeline about 3600 m. Treeless
alpine meadows (zacatonales of Flores Mata et al.) on the highest moun-
tains are the habitat of one species in sect. Pnewmonanthe.

In the Sierra Madre del Sur and the Chiapas-Guatemala Highlands and
along the eastern rim of the Neovoleanic Plateau, the cloud forest of Leo-
pold—termed deciduous forest (bosque de caducifolio) by Flores Mata et al.,
although it includes some pines—prevails above 1500 m and constitutes the
principal habitat of the representatives of sect. Pneumonanthe in these
ranges. In the highest mountains, Pinus rudis Endl. reaches higher eleva-
tions than other tree species, forming open woodlands in which Gentiana
is also present.

The vegetation and major ag ea features of Guatemala, the only
Central American country in which sect. Pneumonanthe is represented,
have been mapped by ene te (1950). Two physiographic regions in
Guatemala are mentioned in the present paper, namely, the Sierra de los
Cuchumatanes, which is a part of Los Altos of Raisz (1964), and the Sierra
Madre of Guatemala, which is continuous with the Sierra de Chiapas of
southern Mexico. Gentiana species occur in the temperate-cold upland for-
est of Steyermark (equivalent, within the context of the present study, to
Leopold’s cloud forest), and in the alpine zones.

Populations of G. bicuspidata, G. hooperi, G. mirandae, G. ovatiloba, and

_ spathacea have been studied in the field. Also, specimens have been
examined from the following herbaria (abbreviations from Holmgren & Keu-

179
ken, 1974): A, ASU, BH, CAN, CAS, COLO, CU, DS, DUKE, ENCB, F, GH,
A,

TAES, TENN, TEX, UC, US, VT, WTU, ac WVA. Only a representative
selection of specimens has been cited for each species, except for the newly
described G. hooperi and the rarely collected G. mirandae, for which all
known collections are listed. A complete list of specimens examin

been placed in the Depository of Unpublished Data, National Science "
brary, Ottawa, Canada, from which copies can be obtained, as well as in
the library of the Royal Botanical Gardens, Hamilton, Ontario.

The chromosome counts reported in this paper are the first for any Mex-
ican species of Gentiana. For G. bicuspidata and G. spathacea, the voucher
specimens cited are those from which seed was obtained in the field. Chrom-
osome counts were obtained from the seedlings. Root tips were pretreated
in a saturated solution of q@-bromonaphthalene for 2.5 hr at 18-20°C, fixed in
Farmer’s fluid, and stained with acetocarmine. Root tips of G. ovatiloba
were obtained from plants collected in the field and were immediately fixed
in Farmer’s fluid. The illustrations of chromosomes are tracings from pho-
tomicrographs.

Species concepts in the present work take into account the remarkable
extent to which interfertility has persisted among morphologically diverse
species in sect. Pneumonanthe. The frequency of natural interspecific hybrid-
ization in this section is correlated more closely with the similarity of the
geographic and ecological ranges of the parental species than with eau
logical similarities. Consequently, the mere existence of intermediates ca
not be accepted as a practical basis for the amalgamation or reduction in
rank of two taxa that would otherwise be treated as two species. Species
have been accepted at this rank if the great majority of specimens seen
could definitely be identified as belonging to one species or the other, with
any intermediates evidently representing exceptional local populations
rather than true intergradation. Experience with sect. Pneuwmonanthe in
Canada and the United States has indicated that species meeting this cri-
terion, even if the differences between them are not great, are neverthe-
less likely to be more sharply defined than any of the more inclusive taxo-
nomic units one might attempt to assemble.

Key to the Mexican and Central American Species of Sect. Pnewmonanthe
la. Corollas red; stamens exserted.

2a. Calyx lobes arcuate-recurved; corolla lobes reflexed; corolla append-

ages bifid, ca. 1 mm long; gynophore much shorter than ovary at an-

thesis. . 6. G. caliculata
. Calyx and epistles jobes erent or eaves SO; aon appendages oblique-
ly triangular, ca. 2 mm long; gynophore about as long as ovary at

hesis. Coe eee ke kee ew 67. G. mirandae.
1b. Corollas blue, violet, or nearly white; stamens included.

3a. Branching, if any, irregular, the branches usually only one at a node;

flowers solitary at ends of stems or branches, or in raceme-like in-

bo
ao

180

florescences, not individually subtended by small bracts; corolla tubes
vase-shaped or broadly campanulate.
4a. Corolla lobes violet-blue, contrasting with the white appendages;
calyx lobes linear-subulate, 2-5 mm long, noticeably shorter than
tube; corolla tube slender below, flaring only in upper half.
be ds ca OS . G, hooper.
4b. Corolla lobes and appendages concolorous, “hie. wolee or nearly
white; calyx lobes linear to ovate-elliptic, about as long as tube or
longer; corolla tube flaring from near base.
da. Leaves linear; upper internodes longer than middle and lower
ones, often over 2 cm long; calyx lobes linear; corolla lobes more
or less orbicular, apiculate; flowers solitary, each terminating an
unbranched stem or a distinct branch usually over 1 cm long
oe iw aoe 4 1. G. eee
. Leaves elliptic, closely spaced throughout; eine lobes elliptic to
ovate; corolla lobes ovate, rounded at apex when fully expanded;
flowers solitary or in rather dense racemes of 2-8, the peduncle-
like branches usually less than 1 cm long. . . 2. G. ovatiloba.
3b. Branching, if any, isocladous; flowers clustered in compact cymes or
heads, individually subtended by small bracts; corolla tubes cylindric
to narrowly urceolate-campanulate.
6a. Leaves broadly lanceolate to ovate, with 3-7 prominent veins, neither
falcate nor strongly conduplicate; calyx usually largely hyaline,
deeply cleft, with minute lobes; corolla lobes elliptic, apiculate.

nn
io”

. oe we ae G. spathacea.
6b. Leaves linear to narrowly oblong, usually iter ani the midvein
prominent, upper leaves usually ear genes falcate and = con-
duplicate; calyx usually green and uncleft, with lobes 2-7 mm long;
corolla lobes deltoid-ovate, usually oes or rounded at the apex
a ae L & . G. inevieate.
1. GENTIANA BICUSPIDATA (G. Don) Briq., eondoiles 4: 324. 1931. Fig. 1.
Pneumonanthe bicuspidata G. Don, Gen. Hist. 4:194. ‘51838’ [1837].
ased on Gentiana assurgens Sessé & Mocifio, ined. Lectotype (hoc
loc.): MEXICO: no further locality data, Mocifio & Sessé s.n., no date,
OXF, photos in HAM!; probable duplicates, Sessé et al. 1368 (number
assigned by curator at MA), no date, F!, MA, and (unnumbered) GH!
Gentiana assurgens Sessé & Mocifo ex G. Don, Gen. Hist. 4:194. ‘1838’’
[1837], pro syn
Gentiana ye gens Cerv. ex Griseb. in Hook., Fl. Bor.-Amer. 2:57.
“1845” 837] (in ablative, ‘“‘adsurgente’’), nomen nudum; Cerv. e
Griseb., ai Sp. Gentianearum p. 286. ‘1839’ [1838], with ot
Type collection: Sesse et al. 1368, cited under Pnewmonanthe bicuspidata
G. Don, above.
Gentiana adsurgens [var.] 8 uniflora Kusn., Trudy S.-Peterburgsk. Obshch.
Estestvoisp., Otd. Bot. 24(2):61. 1894. Syntypes: MEXICO: [Distrito

181

—) &
i cust |
\ >)
a, Me

Uy
A HOND SNe
f aes

Fig. 1. Gentiana bicuspidata. a, flowering stem; b, interior surface of
corolla, and c, exterior surface of calyx, slit longitudinally and pressed; d,
pistil; e, s tic chromosomes; f, ranges of G. autumnalis (aut), G. bicus-
pidata (bic), G. ovatiloba (ova), and G. pennelliana (pen).

182

Federal]: Desierto Viejo, Valle de México, Bourgeau 1123, 17 Oct 1866,
B (probably not extant), LE (uncited duplicates in GH!, MICH!, US!,
and probably elsewhere; and MEXICO: no further locality data, Ehren-
berg s.n., B (probably not extant), LE (uncited duplicate in MEXU!).

Gentiana angusta M.E. Jones, Contr. W. Bot. 12:52. 1908. Probable type:

MEXICO: Chihuahua: Mound Valley, Sierra Madre Mts., alt. 7000 ft,
Jones s.n., 18 Sep 1903, DS!

Tap root usually well developed and persistent, 0.8-1.5 em in diameter at
the summit and 0.5-1.5 dm long, often forked, or occasionally accompanied
or replaced by 2-several subequal secondary roots. Stems usually 1-6, oc-
casionally up to 12 or more, decumbent to suberect, 1-4 dm long, papillose-
puberulent in lines below the leaf bases, otherwise glabrous, usually simple,
occasionally with 1-4 branches. Leaves bright green, linear or lowest ones
narrowly elliptic, longest near middle of stem, only moderately reduced
above and below, except near soil level, upper internodes often appreciably
longer than lower and middle ones. Upper leaves acute at apex, lower
leaves obtuse, all tapering at base. Leaves mostly 1.5-5.5 em long, upper
ones usually 20-30 times as long as broad, occasionally broader, median
ones 8-15 times as long as broad, lowest ones highly variab!e in proportions.
Leaf margins minutely and shallowly denticulate, usually more or less
revolute. Flowers solitary at ends of stems or branches, without involucres,
the uppermost leaves usually well separated from base of flower. Calyx
tube uncleft, 7-17 mm long, glabrous. Calyx lobes erect, linear to narrowly
oblong, 5-10(-20) mm long, acute. Corolla 3-5.5 cm long, the tube funnelform,
increasing gradually in diameter from the base upward. Corolla lobes
spreading, elliptic or sometimes obovate, (5.5-)7-12 mm long, apices abruptly
acuminate to rounded-apiculate, margins sparingly erose-serrate to sub-
entire, sinus adjacent to outer edge of each lobe slightly deeper than the
other. Free portions of corolla appendages 2-5 mm long, subequally bifid,
primary divisions often further cleft, ultimate segments acuminate at-
tenuate. Lower 14-34 of corolla tube pale, exterior surface of the petals
proper suffused with bronze-purple, interior surface with purplish-blue
stripes between the central and lateral veins of petals and with small pur-
plish-blue spots elsewhere, Upper 4-% of corolla tube, lobes, and free por-
tions of appendages pale to deep blue throughout, with the purplish suffu-
sions on exterior surface extending to outer margins of lobes and a green
suffusion extending to all but inner margin, and prominent yellowish-green
spots on interior surface of the lower half of lobes and adjacent portions
of tube. Stamen filaments becoming free a little below middle of corolla
tube, free portions 7.5-10.5 mm long. Anthers ca. 5 mm long, not cohering.
Fruits slightly to more than half exserted at anthesis. Seeds ca. 2 mm long,
0.9 mm wide, completely winged. 2n = 26 (voucher specimen: MEXICO:
Hidalgo: 4.8 km N of Agua Blanca Iturbide, J.S. Pringle 1303a, 28 Jan 1972,
HAM

Most populations of G. bicuspidata are in the Neovolcanic Plateau and in

183

®@ G bicuspidata

a G. hooperi

—

Fig. 2. Documented distribution of Gentiana bicuspidata and G. hooperi.

184

the adjacent southernmost portion of the Sierra Madre Oriental (Sierra de
Pachuca), from Hidalgo west to northeastern Michoacan, with disjunct pop-
ulations occurring in the Sierra Madre Occidental in Durango and southern
Chihuahua (Fig. 2). Its usual habitat is a consistently moist meadow, often
near a stream or pond, most frequently in the pine-alder-fir zone, sometimes
in the fir forest zone, or, in its easternmost populations, in the northern out-
liers of the cloud forest. Specimens have been collected in flower from Sep-
tember through March.

Representative specimens examined:

MEXICO: Chihuahua: near Guacochic, Goldman 179 (GH, US). Distrito
Federal: Cerro del ane cerca del Puerto de Las Cruces, Rzedowski
21727 (DS, ENCB, F, MEXU, MICH, TEX, WIS). Durango: 8 mi NW of Es-
taci6n Coyotes, Breedlove 18778 (MICH). Hidalgo: Sierra de Pachuca, C.G.
Pringle 15007 (ASU, CAS, CM, COLO, F, GH, MICH, MSC, TEX, UC, US,
VT). México: Valley of Toluca, C.G. Pringle 4309 (CAS, F, GH, MEXU,
MICH, MO, MSC, MU, NY, RSA, UC, US, VT). Michoacan: 6-7 mi N of
San Pedro Aguaro, McVaugh & Wilbur 9991 (MICH). Morelos: head of Lake
Zempoala, Correll 14282 (DUKE). Veracruz: Palo Bendito, Huayacocotla,
Herndndez M. & Yolanda de Hdez. 974 (GH). Zacatecas: Coulter 951 (K;
photos in HAM seen).

The names Pneumonanthe bicuspidata G. Don and Gentiana adsurgens
Cerv. ex Griseb. appear to have been based on the same collection, although
on different specimens. This collection was made during the period 1788-1804
at some unspecified Mexican locality by the botanical expedition headed by
Martin Sessé L., which included J.M. Mocino and Vincente Cervantes among
its members (Standley, 1920). Sessé & Mocino designated this collection
Gentiana assurgens in herb., but this name did not appear in their pub-
lished works.

Don (1837) based the name Pneumonanthe bicuspidata on Gentiana assur-
gens Sessé & Mocifio. His acquaintance with unpublished names proposed by
Sessé & Mocino was based on duplicates of their collections in the herbari-
um of A.B. Lambert (as indicated in Don, 1837, passim). Sessé & Mocino
collections from Lambert’s herbarium are now in BM, G, and OXF (Miller,

970). Of these three herbaria, only OXF contains a specimen of their
Gentiana assurgens. Since this appears to be the only specimen of this col-
lection likely to have been seen by Don, it has been selected as the lecto-
type of the name Pneumonanthe bicuspidata G. Don.

According to Miller (1970), Lambert received his Sessé & Mocino speci-
mens from J.A. asi who had selected duplicate material from that left
at MA by Mocino. The annotation ‘‘Guayaquilensis’? on the specimen of
“Gentiana assurgens’’ at OXF may have been added by someone who con-
fused Gentiana assurgens Sessé & Mocino, ined., with Selatium assurgens
D. Don ex G. Don, which was based on a specimen in the Lambert her-
barium collected by Ruiz & Pavon or Tafella at Guayaquil, and which was
equated with Gentiana guayaquilensis Griseb. in Index Kewensis. Although

185

obviously erroneous, this annotation does help to link the OXF specimen to
Lambert’s herbarium and thus to Don

Additional specimens that may be duplicates of the same collection by the
Sessé expedition are at MA and F; those at F were examined during the
present study. These bear Sessé & Mocifio’s label ‘‘Gentiana assurgens
N[obis]”? and the number 1368, which was assigned by a curator at MA.
Another collection, or possibly additional duplicates of the same collection,
bear the label “‘Gentiana adsurgens’’ and the number 682. In addition to the
replicates of Sessé et al. 1368 and 682 at MA and F, there is an unnumbered
specimen in the type collection at GH labeled ‘‘Gentiana assurgens sp. n.
Mexico” and, in another hand, ‘“‘Cervantes’’ following the binomial, and
“Fielding his writing?’’ This specimen is probably a duplicate of Sessé et
al. 1368 or 682. All of the specimens in F, GH, and OXF cited above clearly
represent the species treated here as G. bicuspidata.

Grisebach (1838), in publishing the name G. adsurgens, attributed the
origin of this name to Cervantes, Cervantes never published this name, but
presumably used it in herb. or in ms. The relationship between names pro-
posed or published by Cervantes and those proposed or published by Sessé
& Mocino has been discussed by McVaugh (1945), who concluded that ‘‘we
are probably justified in supposing that the Sessé and Mocifio plants [in
MA and F] are to be regarded as the types of Cervantes’ species.’’ The
name G. adsurgens Cerv. ex Griseb., therefore, seems to be best typified
by the Sessé expedition collection numbered 1368. The specimen seen by
Grisebach may have been in B, where Grisebach worked on his Genera et
Species Gentianearum, in which case it would not be extant

Whether the specific epithet bicuspidata has priority over adsurgens has
been the subject of some discussion (Briquet, 1931; Williams, 1968). Accord-
to Stafleu (1967), the first part of Vol. 4 of Don’s General History of the Dich-
lamydeous Plants (which probably included Pneumonanthe bicuspidata) was
published in 1837, and the entire volume had been published by 8 Apr 1838
or earlier. Grisebach’s Genera et Species Gentianearum, in which this spe-
cies was first described under the name Gentiana adsurgens, was published
in early Oct 1838. Therefore, I accept bicuspidata as being the correct spe-
cific epithet for this taxon.

Gentiana bicuspidata is morphologically similar to G. autumnalis L., a
species native to the Atlantic Coastal eas from South Carolina to new

proportionately lower and much divided; the margins of the corolla lobes of
G. bicuspidata are entire or only slightly erose, while those of G. autumnal-
is are prominently erose-serrate; the calyx lobes of G. bicuspidata are
abruptly acute, while those of G. autumnalis are acuminate; the tap root of
G. bicuspidata is usually persistent and well developed, while that of G.
autumnalis is soon replaced by several lesser roots; and the seed wing of

186

G. bicuspidata is much more prominent than that of G. autumnalis. G. bi-
cuspidata averages lower in stature and probably higher in number of stems
per plant than G. autumnalis, but there is considerable overlapping in the
ranges of variation in both of these traits. The somatic chromosome number
of 26, reported here for G. bicuspidata, has also been reported for G. au-
tumnalis (as G. porphyrio J.F. Gmel.) by Rork (1949).

Gentiana bicuspidata and G. autumnalis, along with G. ovatiloba of Mex-
ico and Guatemala and G. pennelliana Fern. of Florida, appear to be closely
retated, all having the characteristics of the ‘‘first group’? of species recog-
nized above, as well as having widely open corollas with large lobes and
deeply cleft appendages, free anthers, more or less decumbent stems, and,
with the exception of G. ovatiloba, linear leaves. The distribution of this
group of closely related species (Fig. 1f) corresponds to a pattern recog-
nized by many biogeographers, in which identical or closely related taxa
cecur on the Atlantic Coastal Plain of the United States and disjunctly in
the highlands of central and southern Mexico. Species or groups exhibiting
the distribution pattern may have different phytogeographic histories. In
the case of the G. autumnalis-G. bicuspidata complex, however, it appears
probable that its colonization of Mexico and Guatemala occurred subsequent
to its establishment in northern North America, and that the Mexican and
ruatemalan species were derived from an ancestral complex centered in
more northern latitudes, rather than vice versa. This interpretation is indi-
cated by the fact that sect. Pneumonanthe is distributed predominantly in
North Temperate, rather than tropical, latitudes, with its southernmost
populations being restricted to relatively cool, montane habitats, and both
of the groups distinguished above are more widely distributed and more di-
verse in northern North America than in Mexico and Guatemala. The ex-
tent of the distribution of this complex both north and south of the disjunc-
tion in its range and its differentiation into two species in each segment of
its range indicate that this complex has had a relatively long history in both
regions, rather than having recently colonized one of these regions through
long-distance dispersal.

The scattered populations of G. bicuspidata in the Sierra Madre Occi-
dental may be remnants of a more extensive and more nearly continuous
distribution in the past, when climatic conditions permitted greater develop-
ment of mesic vegetation. The assumption that the northwestern outliers of
G. bicuspidata are relictual could account for patterns of variation in this
species. Such traits as relatively short corolla lobes in some specimens
from Chihuahua may be due to past introgression of genetic material from
the G. affinis complex when both species-groups were more widely distri-
buted in western North America; reciprocal introgression may account for
the occurrence of long-branched individuals of G. affinis (= G. interrupta
Greene) near the Mexican-United States border.

Plants of this species bearing only one flower per stem were segregated
by Kusnzow (1894) as G. adsurgens [var.] B uniflora. Herbarium and field

187

studies of G. bicuspidata indicate that such variation commonly occurs
among plants in a single population and even among the stems on a single

Gentiana angusta was described by Jones (1908) from plants found 18
Sep 3 “in meadows along with other gentians, Mound Valley, Sierra
Madre Mts., Chihuahua . . . at 7000 feet alt.’ In the paper in which he
described G. angusta, Jones stated that the type specimens of all new names
published therein were in his personal herbarium. No such specimen is at
POM, which later acquired Jones’s herbarium, but a collection reasonably
compatible with Jones’s description of G. angusta, and bearing identical
data, is at DS, one of the many herbaria to which Jones sent specimens
during his lifetime. This collection comprises three plants mounted on one
sheet, all of which are entirely characteristic of G. bicuspidata, except that
the corolla lobes are relatively short for this species. This collection is
labeled “‘Gentiana linifolia,’’ with no author’s name. It seems probable that
this is the type collection for the name G. angusta. Gentiana linifolia may
have been Jones’s first choice for a name for these plants and may later
have been replaced in his paper but not corrected on the label when he
found that this name was preoccupied. These specimens do indicate clearly
that a collection of G. bicuspidata was made by Jones at Mound Valley on
18 Sep 1903 and was considered by him to represent a taxon to which no
name then in use should be applied (the names G. linifolia Salisb. and G.
linifolia Willd. ex Schult., based on European plants, having long since

passed into obscurity). For these reasons, and because no other Jones spe-

cimens of Gentiana from Mound Valley appear to exist, G. angusta M.E.

Jones is treated here as being a taxonomic synonym of G. bicuspidata (G.

Don) Briq.

2. GENTIANA OVATILOBA Kusn., Trudy Imp. S.-Peterburgsk. Bot. Sada
(Acta Horti Petrop.) 13:60. 1893. Type: MEXICO: [Distrito Federal]:
Desierto Viejo, Valle de México, Bourgeau 1123-bis, 17 Oct 1866, holotype
LE, isotype (not cited by Kusnezow) GH!

Gentiana caespitosa Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles
11(1):370. 1844, non G. cespitosa Willd. ex Schult. in Roem. & Schult.,
Syst. Veg. 4:185. 1819, nec G. caespitosa R. Graham, Edinburgh New
Philos. J. 10:367. 1830. Type: MEXICO: Veracruz: Pic d’Orizaba, 10000
[Paris ft], Galeotti 1483, ‘184-"’ (date incomplete, probably actually
1838), P, photos in F!, !, NY!

Gentiana guatemalensis Standley & Steyerm., Publ. Field Mus. Nat. Hist.,
Bot. Ser. 23:75. 1944. Type: GUATEMALA: Huehuetenango: alpine mea-
dow, region of Chémal, Sierra de los Cuchumatanes, alt. about 3300 m,
Standley 81113, 28 Dec 1940, holotype F!, isotype GH!

Gentiana lewisiae Standley & Steyerm., Publ. Field Mus. Nat. Hist., Bot.
Ser. 23:76. 1944. Type: GUATEMALA: Totonicapan: damp meadow, Pa-
caja, region of Desconsuelo, mountains above Totonicapan, alt. 3100-
3200 m, Standley 84556, 23 Jan 1940,

188

Fig. 3. Gentiana ovatiloba. a, root of plant from deep soil; b, flowering
stem; c, portion of plant from dry, shallow soil; d, interior surface of corol-
la, and e, exterior surface of calyx, slit longitudinally and pressed; f, pistil;
g, somatic chromosomes.

ps \
4GO —~* /
A |
}
le MEX : \
A ee
o~ TAK. ~ y
@oF 8 - 7 a
e . GH
ee Pe enna a
MOR PUE. ‘ : " $ sie
; < ew ; oe CAMP .
= 7 ae - |
\ : pene ag OE ee
\ ‘ 5 a ee aa r
5
s ‘
° \ |
° | iyanee
. f
GRO.
e
+ OAX
. : Bo ge
Pe = CHIS, pom
_s E /
— B eS e
‘
ae
SS
N
) GUATEMALA
[ WW

Fig. 4. Documented distribution of Gentiana ovatiloba.

Tap root usually persistent, 1-1.8 cm in diameter at summit and 0.7-1.5
dm long, often forked distally and producing branch roots; older plants often
bearing many stout secondary roots from branches and divisions of the geo-
corm, these divisions sometimes becoming essentially independent. Stems
cespitose, usually 2-12, occasionally more, decumbent to erect, 1-6 dm long,
glabrous except for a few minute projections in lines below leaf bases, sim-
ple or with 1-7 short branches near summit. Leaves somewhat succulent,
bright green, elliptic, largest near middle of stem, only the lowest much re-
duced, evenly and closely spaced, obtuse to rounded at apex, rounded at
base. Leaf margins smooth, flat or somewhat revolute. Largest leaves 1-3
cm long, 2.5-3.5 times as long as broad. Flowers solitary or in racemose or
corymbose clusters of 1-8 through the production of short branches near
summit of main stem. Flowers usually closely subtended by uppermost pair
of leaves, but without differentiated bracts. Calyx tube uncleft, 6.5-10 mm
long, glabrous. Calyx lobes erect, oblong to narrowly elliptic-ovate, 5-12 mm
long, obtuse, sometimes apiculate. Corolla 2.5-4.5 cm long, the tube funnel-
form. Corolla lobes spreading, ovate-triangular or broadly ovate to elliptic-
rhombic, 3.5-8.5 mm long, sparingly erose-serrate to nearly entire, the apices
cbtuse to acute, the sinuses subequal. Free portions of the appendages 2-4
mm long, basically bifid with the segments variously toothed or cleft, the

190

ultimate divisions acute to attenuate. Lower 34 of corolla tube pale, increas-
ingly suffused with blue upward, with bronze-purple suffusions on exterior
surfaces of petals proper, and purplish stripes (varying in distinctness) on
interior surfaces between midvein and lateral veins. Upper part of corolla
tube, lobes, and free portions of appendages blue, exterior surface of lobes
suffused with green except near inner margin, and with bronze-purple near
outer margin, Blue pigment sometimes very pale, or replaced by violet.
Intericr corolla surfaces unspotted or (in northern populations) with a few
yellowish-green spots near base of lobes. Stamen filaments becoming free
about 4 the height of coro!la tube, free portions 4.5-6 mm long. Anthers ca.
3.9 mm long, not cohering. Fruit slightly exserted from marcescent corolla
at maturity. Seeds similar to those of G. bicuspidata, 2n = 26 (voucher
specimen: MEXICO: México: Canada de Alcalican, SW side of Ixtaccihuatl,
J.S. Pringle & T. Duncan 1509, 22 Jun 1974, HAM).

Gentiana ovatiloba grows in the highest mountains in the Neovolcanic
belt from western Veracruz west to the Distrito Federal, and in the Sierra
Madre del Sur in Oaxaca. Disjunct populations are present in the mountains
of western Guatemala (Fig. 4). Reports of ““G. adsurgens’’ in Chiapas, e.g.
by Standley & Williams (1969), which one might expect to have been based
on G. ovatiloba, actually appear to have been based on specimens of G.
laevigata. Gentiana ovatiloba is usually found in the alpine grasslands just
above treeline, with occasional populations extending to open, exposed sites
in the Pinus hartwegii or P. rudis zones. Most plants of G. ovatiloba are
found along streams and in seasonally moist depressions, associated more
with rushes and mesophytic forbs than with the dominant bunchgrasses.
Occasional plants, however, occur in shallow, dry soils near rock outcrops.
The flowering season extends from August to March, and sporadically
through the summer in some populations.

Representative specimens examined:

GUATEMALA: Huechuetenango: 7 mi N of Santa Eulalia, Breedlove 8597
(DS, F, US). Solo!a: 10-12 km NW of Los Encuentros, Williams et al. 27307
(Ff, NY). Totonicapan: about 12 km SW of Totonicapan, Williams et al.
25462 (I, NY, US). MEXICO: México: Canada de Alcalican, Ixtaccihuatl,
Rzedowski 21569 (DS, ENCB, MICH, WIS). Oaxaca: Zempoaltepetl, Camp
2666 (A, MICH, NY, RSA, UC). Puebla: lddera NE de Ixtaccihuatl, Ern
300 (ENCB). Veracruz: Cofre de Perote, NW side of mountain, Beaman
2189 (GH, MSC).

Collections of G. ovatiloba from Guatemala have consistently been recog-
nized as being specifically distinct from G. bicuspidata. Standley & Steyer-
mark (1944) and Standley & Williams (1969) considered G. guatemalensis
and G. lewisiae, as they designated these plants, to be endemic to Guate-
mala, evidently having accepted the Mexican representatives of this com-
plex as being ‘‘G. adsurgens.’’ In recent years, specimens from Oaxaca
have, with increasing frequency, been equated taxonomically with the Gua-
temalan material, usually as G. guatemalensis. To date, however, specimens

191

virtually indistinguishable from the Guatemalan plants have generally been
included in G. bicuspidata (or “G. adsurgens’’) if they were collected north
of Oaxaca

The most conspicuous differences between G. ovatiloba and G. bicuspidata
are the following. The leaves and calyx lobes of G. ovatiloba are propor-
tionately much wider than those of G. bicuspidata. The leaves of G. ova-
tiloba are more closely spaced than those of G. bicuspidata, especially
toward the summit of the stem, where the leaves of G. bicuspidata are
often widely separated. Gentiana bicuspidata sometimes produces flowering

ranches several cm long, with two or more pairs of leaves, whereas the
branches of G. ovatiloba are almost always short and peduncle-like. The
corolla lobes of G. ovatiloba are ovate to ovate-triangular and are blunt;
those of G. bicuspidata are elliptic to suborbicular, widest near or above
the middle, and are generally apiculate. The intericr surface of the corolla
of G. bicuspidata is conspicuously spotted with yellowish-green, especially
near the base of the lobes; such spots are usually absent from the corollas
of G. ovatiloba, although not invariably so in the northernmost part of its
range. The free portions of the coro'la appendages of G. bicuspidata tend
to be proportionately longer than those of G. ovatiloba. Both the stamen
filaments and the anthers of G. bicuspidata are longer than those of G.
ovatiloba, with the anthers of the former thus being nearer the throat of
the corolla.

Despite this rather extensive list of differences, G. ovatiloba and G._ bicus-
pidata appear to be very closely related. Most plants of this complex can be
readily and unequivocally assigned to one or the other of these species, but
occasional specimens deviate from the typical form of one species in having
one or more traits, usually vegetative, like those of the other species. For
example, Lyonnet 458 (GH, NY, US), from the Desierto Viejo, Distrito Fed-
eral, where both G. bicuspidata and G. ovatiloba have been collected, is
typical of the former species in general habit and in corolla morphology but
has exceptionally broad, blunt leaves and calyx lobes. A duplicate of this
collection in F, however, falls within the usual range of variation in G. bi-
cuspidata. A joint, but independently numbered, collection by Pringle &

Pringle 4985, GH, MICH, US, VT; Smith 752, MO, NY), and Nelson 1400

H, US), from the same general area, consist of plants that are typical of
G. ovatiloba in habit, leaf spacing, and floral morphology but have narrower
leaves (mostly 2-4 mm wide) than those ordinarily found in this species.

Most of the alpine zones constituting the habitat of G. ovatiloba are at
elevations that resulted from Pliocene and Pleistocene uplift and volcanic
activity, with the exception of those in the somewhat older Sierra de los

uchumatanes in the southern part of its range. This association with rela-
tively recently formed habitats suggests that G. ovatiloba is probably the
latest product of evolution in the G. bicuspidata complex, representing an
adaptation to alpine conditions.

192

Standley & Steyermark (1944), as noted above, treated the Guatemalan
representatives of G. ovatiloba as two species, G. guatemalensis and G.
lewisiae, which were originally contrasted as follows: G. guatemalensis—
flowers 1-3 per stem; calyx tube ca. 8 mm long; calyx lobes ca. 10 mm
long; corolla ca. 2.5 cm long; segments of the free portions of the corolla
appendages acute to acuminate, sparsely serrate. G. lewisiae—flowers soli-
tary; calyx tube 5-6 mm long; calyx lobes 4-7 mm long; corolla ca. 2 cm
long; segments of the free portions of the appendages rounded, entire. In
1969, however, both G. guatemalensis and G. lewisiae were described by
Standley & Williams as having 1-few flowers per stem. These authors then
noted that G. guatemalensis and G. lewisiae ‘‘are perhaps too much alike
and the distinctions tend to disappear with additional collections.”

As mapped by Steyermark (1950), the alpine zones, which are the habitat
of G. ovatiloba, are restricted to two areas in Guatemala: the Sierra de los
Cuchumatanes, the source of the more northern records for Guatemala
mapped in Fig. 4; and the Sierra Madre, the source of the records for south-
ern Guatemala. Thus isolated from each other as well as from the Mexican
populations, these Guatemalan populations do appear to have differentiated
o the extent that plants from the Sierra Madre tend to be smaller, in stature
and in the size of their leaves and flowers, than plants from the Sierra de
los Cuchumatanes or from Mexican populations. However, the study of ad-
ditional Guatemalan material which has become available since the descrip-
tion of G. guatemalensis and G. lewisiae, the inclusion of specimens from
Mexico as well as from Guatemala in the present study, and field observa-
tions of sizeable populations occupying diverse microhabitats indicate that
G. guatemalensis and G. lewisiae cannot justifiably be maintained as sepa-
rate species. Quantitative variation appears to be essentially continuous
within this species, with no evident discontinuity separating the Sierra
Madre plants from those of other regions. No qualitative differences were
discerned between the type specimens of G. lewisiae and G. guatemalensis,

r between specimens from the Sierra Madre and those from the Sierra
s los Cuchumatanes or from Mexico. The divisions of the corolla append-
ages appear to be as acute in the type collection of G. lewisiae, and in
other specimens from the Sierra Madre which have been identified as G.
lewisiae, as in the type collection of G. guatemalensis or in other specimens
from Guatemala or Mexico.

Limited local variation also appears to have developed among the Mex-
ican populations of G. ovatiloba. As 1 observed on Ixtaccihuatl, for exam-
ple, stems of large and small plants alike bore oy one flower per stem,
whereas in the Sierra Juarez, Oaxaca, flowers were almost invariably
borne in racemes of two or more. Within the Ixtaccihuatl popu ation, vari-
ation not evidently correlated with environmental factors was note co-
rolla color and lobe length. Microenvironmental factors also appear i con-
tribute significantly to variability of G. ovatiloba; plants in relatively dry,
exposed sites differ conspicuously in stem height and in spacing and condu-

198

plicateness of the leaves from those growing among dense vegetation in
wet depressions (Figs. 3b and 3c).
3. NTIANA heooperi Pringle, species nova. Fig. 5.

erba perennis, glabra. Caules 1-6, ascendentes, 0.3-2 dm longi. Folia
griseo-viridia, aliquanto succulenta, obtusa, maxima 2-4.5 cm longa in parte
caulis inferiore. Flores vulgo solitarii, interdum racemosi 2-4, exinvolu-
crati. Tubus calycis infissus, 8-16 mm longus. Lobi calycis linear-subulati,
acuminati, 2-5 mm longi. Corolla infundibuliformis, 3.5-5.5 cm longa. Lobi
corollae elliptico-obovati, patentes, 6-10 mm longi, 4.5-9 mm lati, integres,
obtusi, violaceo-lazulini, albomarginati, e viridi-purpureo extra. Partes li-
brae plicarum 2-4 mm longae, 3-5 mm latae, bifidae segementis laciniatis,
medium tubi corollae; partes librae 5-7 mm longae.

Type: MEXICO: Aguascalientes: Municipio San José de la Gracia, high
plateau in the Sierra Fria ca. 8 km SW of La Congoja, ca. 22° 10’ N, 102°
33’ W, 7 Feb 1972, J.S. Pringle 1320 (ho'otype HAM, isotypes CAN, HAM, K,
MEXU, MICH, NY, S, TENN). Topo-paratype, from the same locality, ex-
pressed as “ ‘Sierra Fria’ 15 mi W of Presa Calles’: Hooper 10 Mar 1953
(MICH). Additional paratypes: MEXICO: Chihuahua: vicinity of Minaca,
Rose 11645 (US). Durango: Sierra de San Francisco, Foshag Feb 194i
(MEXU); Sierra de Carcaria, near Rio Verde, Foshag 28 Feb 1941 (ENCB

US).

Several subequal roots 1.5-8 mm in diameter usually present. Stems 1-6,
decumbent to ascending, 0.3-2 dm long, glabrous, simple or with 1-4 short
branches. Leaves gray-green, often suffused with purple, rather succulent,
scmewhat conduplicate, especially upper ones, linear or lower ones nar-
rowly elliptic to oblanceolate, obtuse at extreme apex, tapering at base,
upper leaves strongly ascending, median and lower leaves moderately as-
cending or spreading. Leaf margins very narrowly cartilaginous, minutely
denticulate proximally, entire distally, the cartilaginous margin usually
abruptly widened in distal 1-3 mm. Leaves largest on lower half of stem.
upper leaves gradually reduced to 1/3-2/3 the length of the largest leaves,
only those near and below the soil line scalelike. Largest leaves 2-4.5 em
long, (4-)8-30 times as long as wide. Flowers solitary at end of main stem
and branches, occasionally appearing racemose if several reduced branches
are present, exinvolucraie or subtended only by uppermost pair of leaves.
Calyx tube uncleft, 8-16 mm long, g’abrous. Calyx lobes erect, linear-sub-
ulate, 2-5 mm long, acuminate. Corolla 3.5-5.5 cm long, tube funnelform,
with lower half slender, widening very gradually upward, upper half more
abruptly flaring. Coro!la lobes spreading, elliptic-obovate, 6-10 mm_ long,
4.5-7(-9) mm wide, entire, obtuse, sinuses about equal on both sides. Free
portions of appendages 2-4 mm long, 3-5 mm wide, bifid, divisions further
irregularly toothed or cleft, ultimate segments acute to attenuate. Corolla
tube greenish-white below, increasingly suffused upward with violet-blue
and externally with green, in the petals proper. Corolla lobes violet-blue
(often similar in color to corollas of typical Vinca minor L., sometimes

194

_ 5. Gentiana hooperi. a, flowering plant; b, interior surface of corolla,
a . exterior surface of calyx, slit longitudinally and pressed; d, pistil.

paler or bluer, occasionally nearly white), narrowly white-margined, the
exterior surface suffused with green except near inner margin, and with
purple near outer margin. Dark blue-violet, usually interrupted, streaks
present on inner surface of the upper 4-% of corolla tube between midrib
and lateral veins of each petal, extending into the lobes as rows of dots;
additional spots or short streaks occasionally present at juncture of petals
proper and appendages. Midribs narrowly outlined with green on both sur-
faces. Appendages white throughout, or fused portions with a faint to prom-
inent blue patch on each side near summit. Stamen filaments becoming
free at or slightly below middle of corolla tube, free portions 5-7 mm long.
Anthers free at anthesis. Fruit and seeds not seen.

Gentiana hooperi is named for Emmet T. Hooper, Curator of Mammals,
University of Michigan Museum of Zoology, whose collection of this species
was the first I saw, and whose directions enabled me to visit the type
locality.

To date, G. hooperi has been collected only at four widely separated lo-
calities, all in the lower, eastern ranges of the Sierra Madre Occidental
(Fig. 2). At the type locality in the Sierra Fria in northwestern Aguascali-
entes, some plants of G. hooperi were found in swampy sites (dry at flower-

195

ing time) among relatively dense stands of Juncus species and other grami-
noids. Other plants grew on higher ground where only a sparse herbaceous
layer was present, among widely scattered small trees of Juniperus deppe-
ana Steud., some of the gentians being in seasonally moist depressions,
others in very dry, exposed sites. Gentiana hooperi was just coming into
bloom when the type collection was made in early February; the other col-
lections, for which dates are given above, also comprise plants in flower.

The slender corolla tube of G. hooperi, which flares widely only near the
summit, and the linear, succulent, gray-green leaves give this species an
aspect superficially similar to that of some species in sect. Frigidae Kusn.
The leaf margins also resemble those of species in sect. Frigidae. The habit
of growth of G. hooperi, however, clearly indicates that its affinity is with
sect. Pneumonanthe, since species in sect. Frigidae have slender, creeping
rhizomes from which either overwintering or permanent rosettes arise. The
well-developed, bifid free portions of the corolla appendages of G. hooperi
are also more characteristic of sect. Pnewmonanthe than of sect. Frigidae.

The closest relatives of G. hooperi are probably G. bicuspidata and G.
ovatiloba. It is, however, well differentiated from these species and may
have had a relatively long history of isolation. Although additional popula-
tions of G. hoopevi may remain to be discovered, its distribution is obvious-
ly sporadic and disjunct and is probably best regarded as comprising the
remnants of a more nearly continuous distribution in the past.

Gentiana hooperi is generally of lower stature than G. bicuspidata, even
when plants growing in apparently similar habitats are compared. The
leaves of these species differ markedly in color and texture. The acuteness
of the upper leaves of G. bicuspidata also contributes to the difference in as-
pect between it and G. hooperi. The calyx lobes of G. hooperi are shorter
and, in contrast to the leaves, more sharply pointed than those of G. bicus-
pidata.

The corolla of G. hooperi is significantly different is shape from that of
G. bicuspidata, the corolla tube of G. hooperi being prolonged and slender,
as noted above, in contrast to the proportionately broader corolla tube of
G. bicuspidata. The corolla lobes of G. bicuspidata are usually apiculate,
whereas those of G. hooperi are obtuse (although the inrolling of the mar-
gins, before the corolla has fully expanded, may temporarily create the
appearance of apiculate lobes on newly opened corollas). The illustrations
of these species show the marked difference in the levels at which the sta-
mens become free from the corolla tubes.

Two aspects of corolla pigmentation present especially obvious differences
between G. hooperi and G. bicuspidata. The basic color of the corollas of
G. hooperi is a slightly grayed periwinkle-blue, with an appreciable violet
content, while that of the corollas of G. bicuspidata is a true blue, varying
in intensity but with very little violet content. The predominantly white free
portions of the appendages of G. hooperi are conspicuous and contrast with
the blue lobes. In G. bicuspidata, the lobes and appendages are similar in

196

color. Other differences between G. hooperi and G. bicuspidata can be noted

from the descriptions and illustrations of these species.

4. GENTIANA SPATHACEA H.B.K., Nov. Gen. Sp. 3:173 (quarto text).
1

Type: MEXICO: [Veracruz]: in declivitate orientali montium Mexi-
canorum, prope urbem Xalapae, alt. 700 hex., Humboldt s.n., no date,
P, photos in F!, MO!, US! Fig. 6.

Gentiana plicata Willd. ex Schult. in Roem. & Schult., Syst. Veg. 6:185.
1820. Type: duplicate of type collection of Gentiana spathacea, cited
above, B-herb. Willd., photos in HAM!

Coilantha sessaei D. Don ex G. Don, Gen. Hist. 4:185. 1837. Type: MEX-
ICO: no further locality data published, Sessé et al. collection formerly
in herb. A.B. Lambert, not located, probably not extant. (x char.)

Coilantha mocinni D. Don Ex G. Don, Gen. Hist. 4:185. 1837. Type: MIX-
ICO: no further locality data published, Sessé et al. collection formerly
in herb. A.B. Lambert, not located, probably not extant. (Ex char.)

Ericala spathacea (H.B.K.) G. Don, Gen, Hist. 4:191. 1837.

Gentiana ovalis Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 11(1):369.
1844. Type: MEXICO: [Veracruz]: Xalapa, 6000 |Paris ft], Galeotti
1486, no date, G, photos in F!, GH!, MICH!, NY!, US!

Gentiana sessaei (G. Don) Griseb. in DC., Prodr. 9:112. 1845.

Gentiana spathacea {var.] B Benthami Griseb. in DC., Prodr. 9:118. 1845.
ype: MEXICO: [Hidalgo]: Velasco, prope Real del Monte, Hartweg
349, 1839, holotype K, photos in F!, MICH!, isotype NY!

Gentiana coerulea Sessé & Mocino, Naturaleza (Mexico City), ser. 2,
1(App.):44. 1888, etiam G. caerulea [sic] Sessé & Mocifo ex G. Don,
ten. Hist. 4: 185. ‘11838 [1837], pro syn., et G. coerulea Sessé & Mocino
ex, Griseb. in DC., Prodr. 9:112. 1845, pro syn., non G. coerulea Ruiz
& Pavoén ex Griseb., Gen. Sp. Gentianearum p. 235, ‘1839 [1838], pro
syn. Type collection: MEXICO: no further locality data, Sessé et al.
897 (no. assigned by a later curator at MA), no date, MA, F!

Gentiana spathacea [var.] y integra Kusn., Trudy §.-Peterburgsk. Obshch.
Estestvoisp., Otd. Bot. 24(2):53. 1894. No type indicated.

Pneumonanthe spathacea (H.B.K.) Greene, Leafl. Bot. Observ. Crit. 1:71.
190

nN

Dasystephana spathacea (H.B.K.) Arthur, Torreya 19:49. 1919.

Tap root persistent, usually 1-2 cm in diameter at summit and 1.5-2 dm
long, generally forked and bearing branch roots distally, often accompanied
by several-many thick secondary roots arising from the elongating, some-
times much branched and divided geocorm. Stems 1-20(-50 or more), as-
cending to erect, 1-10 dm tall, minutely papillose-puberulent or glabrous.
Leaves bright green, lanceolate to broadly ovate, prominently 3-nerved
(the larger ones also having 1 or 2 pairs of less prominent nerves), round-
ed at base, acute at apex, with margins entire and minutely revolute. Larger
leaves 2.5-8 ecm long, 2-5(-6) times as long as wide. Leaves largest a short

Gentiana spathacea. a, underground portions; b, flowering stem;
Cc, iteelor surface of corolla, and d, exterior surface of cee slit longitud-
inally and pressed; e, pistil; f, somatic eee fie . documented dis-
tribution of G. spathacea (open circles), bic a (closed circles),
and mixe populations ae hybrids Fae acca. ae in the Distrito
Federal and vicinity.

197

198

distance below summit of stem, slightly reduced above, gradually reduced
to small scales near base of stem. Flowers solitary or in small sessile or
short-pedunculate groups in upper 1-8 (rarely more) axils and in terminal
clusters of 3-30, those in clusters subtended by pairs of reduced leaves or
linear bracts (see discussion below). Calyx tubes typically cleft to base on
one side, hence spathelike, 5-10 mm long, herbaceous below, hyaline above,
glabrous. Calyx lobes usually subulate, less than 1 mm long, closely spaced
at summit of the tube opposite the cleft. Occasional plants with calyces
larger, herbaceous throughout, less deeply or not at all cleft, with oblong-
oblanceolate lobes up to 6 mm long and more evenly spaced. Corollas
2.5-4(-5) cm long, tubes urceolate-campanulate. Corolla lobes spreading,
broadly ovate-elliptic to orbicular, 4-8(-10) mm long, 2.5-6.5 mm wide, en-
tire, apiculate, sinuses on each side almost equally deep. Free portions of
appendages erect, 3-6 mm long, 2.5-4 mm wide, tapering into 2 subequal,
closely spaced, attenuate teeth, sometimes with lesser teeth or irregularities
along the margin. Lower 2/5 of corolla tube whitish, remainder of corolla
blue, becoming deeper in color upward, especially in the petals proper,
the deep blue of lobes extending downward on interior surface of the tube
in stripes between the midrib and lateral veins. Petals below the lobes,
and the outer margin of each lobe, very dark and suffused with purple on
exterior surface. Appendages lighter, but blue throughout. Stamen_ fila-
ments becoming free at ca, 3/8 the height of corolla tube, free portions ca.
6 mm long. Anthers loosely coherent or free at anthesis, ca. 3 mm long.
Fruits only slightly exserted from corolla at maturity. Seeds ca. 2 mm long,

mm wide, winged, wing very narrow on one side, 2n = 26 (voucher
specimen: Hidalgo: 2.6 km N of Mexico Hwy. 105 on road to Mineral del
Chico, J.S. Pringle 1305a, 28 Jan 1972, HAM).

The range of G. spathacea largely coincides with the boundaries of the
Neovoleanic Plateau, extending from Veracruz west to Jalisco, although it
reaches the southernmost portions of the Sierra Madre Oriental in Hidalgo
and southern San Luis Potosi and has been collected at one locality in the
northwestern part of the Sierra Madre del Sur in Guerrero (Fig. 7). Most
populations of G. spathacea are in the upper reaches of the pine-oak forest
and in the pine-alder-fir zone; a few extend to the fir forest zone. Within
these zones, the usual habitat of G. spathacea consists of partially shaded,
generally well-drained slopes, often in a small opening or near the edge of
the woods. Specimens have been collected in flower from August through
April. Plants in the drier microhabitats usually bloom in autumn or early
winter and may be found with ripe fruits and desiccated stems in late win-
ter, while plants in moister sites are then in bloom.

Representative specimens examined:

MEXICO: Distrito Federal: Sierra de Ajusco, C.G. aa 6221 (CAS,
CM, F, GH, MEXU, MICH, MO, MSC, NY, TEX, UC, US, VT). Guanajuato:
2miS of Santa Rosa, Harker & Mellowes 145 (ENCB, WIS). Hidalgo: Sierra
de Pachuca, C.G. Pringle 15006 (ASU, CAS, COLO, GH, MICH, MSC, TEX,

Fig. 7. Documented distribution of Gentiana spathacea.

UC, US, VT). Jalisco: Nevado de Colima, McVaugh & oo 11615 (MEXU,
MICH, US). México: Meson Viejo, Hinton et al. 8973 (ENCB, F, GH, MICH,
MO, NY, US). Michoacan: Zitacuaro - Copandiro, Hinton a 13551 (MICH,
NY, US). Morelos: Toro, iad 21923 (MEXU). Puebla: near La Venta,
Share 441853 (GH, HAM, XU, MO, NCU, NY, TENN). San Luis Potosi:
5 km al N de Soledad de ee Rzedowski 7095 (ENCB, MEXU, TEX).
Tlaxcala: am Fuss der Ruines Cacaxtla, Seler & Seler 3556 (GH).
Gentiana spathacea appears to be most closely related to the G. affinis
complex of western North America north of Mexico, although not exhibit-
ing so close a relationship as that of G. bicuspidata to G. autumnalis. This
group of species is also more widely distributed, and includes more taxa,
in northern North America than in Mexico. At times during the Tertiary,
northern, mesophytic species-groups were evidently more widely distributed
in the Southern Rocky Mountains and the Sierra Madre Occidental; it seems
likely, therefore, that the range of the G. affinis-G. spathacea group was
more nearly continuous in the past, and variation within this group may
formerly have been clinal. The present degree of differentiation of G. spa-
thacea from related species, however, along with its extensive distribution
and variability on the Neovoleanic Plateau and in adjacent regions, indi-

200

cates that this species has probably had a relatively long history in Mexico.

Gentiana rusbyi Greene ex A. Gray, from the Mogoll6n Mountains of New
Mexico, was described (Gray, 1886) as being intermediate between G. affinis
(as G. bigelowii A. Gray, a taxonomically insignificant variant) and G. spa-
thacea. Examination of the type collection, Rusby 263 (MICH, MO, NY),
however, indicates that plants so designated are entirely typical of G. affinis
in habit, foliage, and corolla morphology, differing only in having reduced
calyx lobes. According to Hitchcock (1959), such plants occur sporadically
throughout much of the range of G. affinis and do not warrant taxonomic
recognition. It appears, therefore, that G. rusbyi should be included within
G. affinis and need not be interpreted as connecting G. affinis with G. spa-
thacea.

Variants of G. spathacea have been recognized pee by ie
authors. It was first treated as more than one species by Don (1837),
divided it into three species in two genera. Coilantha sessaei was ae
on a Sessé & Mocinfo collection in the herbarium of A.B. Lambert, which
had been designated G. coerulea by its collectors. The specimen seen by
Don in Lambert’s herbarium could not be located in BM, G, or OXF, the
three herbaria known to have received Sessé expedition specimens from the
Lambert herbarium. Examination of a duplicate of this collection in F, how-
ever, has confirmed that G. coerulea Sessé & Mocifho, and hence C. sessaei
G. ai are synonymous with G. spathacea H.B.K.

Don’s (1837) description of Ericala spathacea is simply a paraphrase of
the description of G. spathacea by Humboldt et al. (1819), and contains no
indication that Don had examined any specimens thus identified. It is not
clear from Don’s descriptions what he considered to be the most significant
differences between C. sessaei and I. spathacea or why he placed them in
different genera. The only point in which they were actually contrasted was
that C. sessaei was said to have axillary flowers, whereas thos E. spa-
thacea were described as being in terminal clusters. When aes (1845)
transferred C. sessaei to Gentiana, he distinguished it from G. spathacea
chiefly by its lacking paired bracts beneath the flowers, according to Don’s
description of this taxon.

Don (1837) also described Coilantha mocinni, which was said to differ
from C. sessaei in that it did have paired bracts beneath each flower. His
description of C. mocinni as having 5-nerved, ovate-lanceolate leaves and
a spathaceous calyx, and his statement that it, too, had been identified as
G. coerulea Sessé & Mocino in Lambert’s herbarium, indicate that it was
also based on a specimen of G. spathacea, perhaps another specimen from
the collection on which the name C. sessaei had been based. No specimen
appropriate for the typification of the name C. mocinni could be found in
BM, G, or OXF. Grisebach’s (1845) treatment of this name as a synonym
of G. caliculata Lex. was based only on Don’s description of C. mocinni
and was evidently due to Grisebach’s admitted unfamiliarity with G. cali-
culata, of which he had then seen no specimens. Certainly the elements of

201

the description of C. mocinni noted above are inapplicable to G. caliculata.

To discuss the status of these segregates, it is necessary to consider the
nature of the inflorescence of G. spathacea, which is basically a dichasial
cyme. Within G. spathacea, there is a trend toward condensation of the in-
florescence. Plants at one end of this trend have the entire inflorescence
relatively open, so that at least the lower flowers may appear to be pedi-
cellate and to be subtended only by “‘leaves,’’ with no small bracts being
present. In other plants, the pedicels, cyme branches, and upper internodes
are very short or obsolete, and the leaves and bracts in the inflorescence
are much reduced. In the most extreme expression of this trend, the inflores-
cence is more or less capitate, with the inner (theoretically upper) bracts,
or some of them, minute or obsolete. Within a single population of G. spa-
thacea, a wide range of variation in condensation of the inflorescence may
be encountered. Small plants in the driest, most exposed microhabitats gen-
erally have small, terminal, capitate inflorescences, whereas plants in moist-
er, more shaded sites are likely to have more longated inflorescences, with
flower clusters or short flowering branches arising from several of the upper
axils of the main stems. Otherwise, there is little obvious correlation be-
tween microhabitat and inflorescence form. Obviously, minor variation in
such features as the length of cyme branches and pedicels, or the develop-
ment of bracts, does not constitute a sound basis for taxonomic division of
G. spathacea.

Martens & Galeotti (1844) segregated a solitary-flowered individual or
individuals of G. spathacea with relatively broad leaves as G. ovalis. The
solitary flower can almost certainly be attributed to the age of the plant or
to adverse growing conditions. Leaf shape is variable within populations of
G. spathacea and is evidently related to environmental factors. Plants in
dry, sunny situations have relatively narrow leaves, which have a tendency

tionately wider, wide-spreading, an at. The leaves of the type specimen
of G. ovalis do not significantly exceed the range of variation in leaf shape
commonly found in populations of G. spathacea.

An unnamed variant of G. spathacea was described by Bentham (1840)
as having larger calyx teeth and frequently entire corolla appendages. This
variant was given formal taxonomic status as G. spathacea var. benthamii
("5 Benthami’’) by Grisebach (1845). Later, Kusnezow (1894) described
var. integra as having an uncleft calyx with unequal, subulate teeth. (The
calyx tube of var. benthamii was not described by Bentham or Grisebach
but was evidently assumed by Kusnezow to have been cleft.)

Variability in calyx form exists throughout the range of G. spathacea.

with minute lobes. Specimens approaching both extremes of variation, with

202

calyces ranging from hyaline and deeply cleft, with minute lobes, to her-
baceous and uncleft, with lobes up to 6.5 mm long, may sometimes be found
within a single collection, e.g., Hinton 524 (US), from Temascaltepec, Méxi-
co, Hinton et al. 13540 (F, ae MICH, MO, NY, US) from the vicinity of
Zitacuaro, Michoacan, or Hinton et al. 15749 (DS, F, MEXU, NY, POM, UC,
US), from Barroloso, Michoacan. There appears to be a trend toward less
frequent and less extreme cleavage of the calyx tube and reduction of the
calyx lobes in the western part of the range of G. spathacea, but plants
from as far west as the vicinity of Autldn, Jalisco, exhibit considerable
variation in these traits, even among the plants from a single population
(McVaugh et al. 10285 and 21320, MICH). Except that large calyx lobes are
generally associated with shallowly cleft or uncleft calyx tubes, the traits by
which var. benthamii and var. integra were distinguished do not appear to
be correlated with other morphological features. Neither the corolla lobes
nor the free portions of the appendages of the holotype and isotype speci-
mens of var. benthamii appear to differ appreciably from those of typical
G. spathacea. Because of the sporadic occurrence of plants with relatively
large calyx lobes and/or uncleft calyx tubes, formal taxonomic recognition
of these variants seems to be unwarranted.

Plants collected in the vicinity of Patzcuaro, Michoacan (C.G. Pringle
3982, MEEXU, MSC, MU, NY, POM, UC, US, VT) are exceptional in that the
stems had continued to clongate after the devclopment of the lower axillary
flower buds, so that, at the time of collection, clusters of flowers or buds,
opening in acropetal sequence, were present in as many as 14 successive
axils. Moran 10075 (DS), from El Carmen, Hidalgo, has unusually large
corollas for this species, ca. 5 cm long, with acuminate lobes ca. 10 mm
long.

Gentiana spathacea is also variable in other, less conspicuous aspects.
Plants in some populations have minutely papillose-puberulent stems, while
plants in other populations have glabrous stems. (Most Gentiana species
studied by me are consistent in having either puberulent or glabrous stems,
but some, e.g., G. saponaria L., are similarly variable in this respect.)
Within a single population of G. spathacea, one may encounter variation in
whether the anthers are coherent or free; this variation may be correlated
to some degree with the age of the flowers. Inrolling of the margins of the
obes of wilted or marcescent corollas may affect the superficial aspect of
specimens prepared from such material.

The name G. plicata Willd. ex Schult. was not applied to a proposed seg-
regate of this species but instead to the same collection that typifies the
name G. spathacea, Schultes and Kunth having been working simultaneously
on different sets of Humboldt’s collections. In the present study, examination
of photographs of the type specimen of G. plicata has confirmed that Hum-
boldt et al. (1823) were correct in treating this name as a synonym of their
own G. spathacea.

2038

9. GENTIANA LAEVIGATA Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles
11(1):370. 1844. Type: MEXICO: Oaxaca: Cordillera, Sierra, 5-7000
[Paris ft], Galeotti 1481, Nov-Apr 1840, G, photos in F!, MICH!, NY! (du-
plicates may be elsewhere). Fig. 8

Tap root, if present, 2.5-5 mm thick, usually accompanied or replaced by
several well-developed secondary roots. Stems 1-6, at least the basal por-
tions decumbent, the distal portions varying from flexuous to erect, 1.5-5(-9)

dm long, minutely but copiously papillose-puberulent below, sparsely papil-

lose-puberulent to smooth above. Stems and leaves usual y much suffused

with reddish-purple. Lower leaves, or nearly all the leaves on short stems

(exclusive of reduced leaves near soil line), lanceolate to oblong, nearly flat

and widely spreading to moderately conduplicate and arcuate, with 1 or 3

prominent veins, 1.5-5 cm long, 3.5-6.5 times as long as wide, obtuse to sub-

acute at apex, abruptly tapering at base. Lower internodes 0.8-3 cm long.

Upper leaves similar in shape but increasingly conduplicate and arcuate,

1.5-3 cm long. Upper internodes of well-developed stems 4-9 em long. Involu-

cral leaves subtending terminal flower cluster in 1 or 2 pairs, lanceolate to

ovate, 2-3.5 cm long, (3-)8-14 mm wide, acute at apex, abruptly tapering to
strongly rounded at base, strongly conduplicate, arcuate, enveloping the
lower part of flower cluster. Leaf margins entire, revolute. Flowers solitary
or in clusters of 2-6, confined to the terminal cluster or also borne on short

(up to 2.5 cm), peduncle-like branches from uppermost 1-4 nodes. Individual

flowers subtended by linear bracts usually 10-15 mm long, occasionally larg-

er and foliaceous. Calyx tubes typically uncleft, 7-12 mm long, glabrous, the
portions enveloped by the involucral leaves often somewhat hyaline. Calyx

lobes erect or nearly so, typically linear-subulate to narrowly oblong, 2.5-8

mm long, 0.2-2 mm wide, acute. Calyces of occasional plants deeply cleft,

with greatly reduced lobes. Corollas (2.5-)3-4 em long, tube urceolate-cylin-

dric, gradually expanding upward. Corolla lobes spreading, ovate-triangular,

3.0-7 mm long, as wide or slightly wider than long, minutely erose-serrate

to nearly entire, obtuse or occasionally apiculate, sinuses subequal. Free

portions of appendages asymmetrical, consisting of 2 nearly equal, attenu-
ate teeth each 1.5-2 mm long and 0.5-1.2 mm wide, and an erose shoulder

1.8-3 mm wide extending to outer edge of the next lobe clockwise. Lower

half of corolla tube whitish, tube increasingly suffused with blue upward,

uppermost part of the tube, the lobes, and the free portions of the append-
ages deep blue throughout. Exterior surface of corolla with purplish suffu-
sions as in G. spathacea. Blue pigment extending downward on interior sur-
face of tube as prominent stripes between central and lateral petal veins.

Stamen filaments becoming free at ca. 2/5 the height of corolla tube, free

portions 10-12 mm long. Anthers loosely connate or free. Summit of ovary

about even with tips of marcescent corolla in mature fruit. Seeds ca. 1.8

mm long and 0.8 mm long and 0.8 mm wide, winged, wing very narrow along

one side.

Gentiana laevigata is evidently endemic to the Sierra Madre del Sur in

Oaxaca, and to Los Altos, a subdivision of the Chiapas-Guatemala Highlands

204

Fig. 8. Gentiana laevigata. a, flowering stem; b, interior surface of corol-
la, and c, exterior of calyx, slit longitudinally and pressed; d, pistil; e, docu-
mented distribution.

205

in Chiapas (Fig. 8e). Its usual habitat consists of grassy slopes in the pine-
oak zone and in the lower portions of the cloud forest. Flowering occurs
from August through February.

Representative specimens examined:

ME : Chiapas: between Las Casas and Ejido “El Triunfo,’ Sharp
45994 (MEXU, NY, TENN); barrio of Tuk, paraje of Matsab, Municipio of
Tenejapa, Breedlove 12443 (DS, F, MICH): 9 mi SE of San Cristébal Las
Casas, Breedlove & Raven 13421 (DS, F); near Zinacantan center, Laughlin
2705 (DS, US). Oaxaca: Santa Inés del Monte, Conzatti 1348 (GH, MEXU):
near Tlaxiaco, Camp 2264 (NY); halfway from Guelatao to Llano de las
Flores, Vilas 332A, 332B (WIS)

Gentiana laevigata is represented by relatively few collections. It occurs
in small, widely scattered populations that exhibit some local differentiation
in morphological traits. The majority of specimens, however, including all
those cited above, are very similar to one another and to the type specimen.
These specimens, therefore, were regarded as being most representative of
G. laevigata in the preparation of the description of this species.

Gentiana laevigata appears to be a derivative of G. spathacea or its im-
mediate ancestor, which became established in southern Mexico after oro-
genic processes had created suitable habitats. Its range in the mountains of
Oaxaca and Chiapas is isolated from that of G. spathacea by the intervening

Gentiana laevigata and G. spathacea differ most conspicuously in their
general habit and foliage. The stems of G. spathacea are relatively stout
and are ascending to erect, or decumbent only near the base. Mature plants
growing in favorable sites usually produce many stems. The stems of G.
laevigata, in contrast, are slender and often flexuous, Laughlin 2705 (DS,
US) having been described as a ‘‘vine,’”’ and seldom number more than
three or four per plant. Gentiana spathacea is usually densely leafy, where-
as the upper leaves of G. laevigata are widely spaced. Also, the leaves of
G. laevigata are proportionately wider than those of G. spathacea. The
upper leaves of G. laevigata are generally somewhat arcuate and strongly
conduplicate, in contrast to those of G. spathacea, which are not arcuate
and seldom strongly conduplicate except in sun form

The corolla lobes of G. laevigata are broadly ee. ovate, and the apices
are usually obtuse or rounded. Thus they contrast with the elliptic, strongly
apiculate corolla lobes of G. spathacea. Also, the free portions of the corolla
appendages of G. laevigata are definitely asymmetrical, consisting of two
teeth and an erose “‘shoulder,’’ whereas those of G. spathacea are nearly
symmetrically divided into two teeth only. Gentiana laevigata differs further
from G. spathacea in its proportionately longer stamens.

Both G. laevigata and G. spathacea are variable in calyx form. Typical
plants of G. laevigata differ from those of G. spathacea in having uncleft
calyces with well-developed, subequal lobes. Occasional plants of G. lae-
vigata, e.g., C.G. Pringle 5647 (GH, MICH, VT), collected near Las Sedas,

206

Oaxaca, differ from the typical form of this species in having cleft calyx
tubes. Ghiesbreght 718 (MO), from the vicinity of San Crist6bal Las Casas,
Chiapas, and MacDougall 2616 (NY), collected between La Cumbre and
Ixtlan de Juarez, Oaxaca, include flowers with cleft and uncleft calyces on
the same sheet.

Specimens collected between Tamazulapa and Ayutla, Oaxaca (Camp
2725, NY) exhibit unusual branching and have proportionately wider leaves
than are commonly encountered in G. laevigata, although the flowers are
entirely characteristic of this species. On one of the plants in this collection,
there are three slender branches 1-1.5 dm long, plus several leafy tufts,
borne near the base of the main stem, in addition to branches up to 3 cm
long from the upper five axils. The branches terminate in clusters of rela-
tively small flowers. Another plant on the same sheet has branches ca. 1.5
cm long from the upper three axils only.

6. GENTIANA CALICULATA Lex. in La Llave & Lex., Nov. Veg. Descr.
p. 18. 1824. Type: MEXICO: |Michoacan]: prope Santa Maria, oppidum
indianum juxta oe no further collection data, not located,
probably not ex . (ex ar.) Neotype (hoc loc.): MEXICO: [Méx-
ico]: in sylvis prope ae Ehrenberg s.n., no date (probably
Apr 1831), MEXU! Fig. 9.

Gentiana salpinx Griseb., Linnaea 22:44. 1849. Type: MEXICO: [México]:
in sylvis prope Temascaltepec, Ehrenberg 450, Apr 1831, holotype form-
erly in B, probably not extant; neotype of Gentiana caliculata, cited
above, is probably an unnumbered duplicate of the same collection;
other duplicates may be elsewhere.

Root system comprising several subequal, fleshy roots (according to Lex-
arza, in La Llave & Lexarza, 1824). Stems 1-few, ascending to erect, 0.4-1.6
m tall, minutely papillose-puberulent. Lower leaves (except for scalelike
leaves near soil line) ovate to elliptic, flat, 3-5-nerved, 3-9 cm long, 1-4 cm
wide (length/width ratio variable), acute at apex, rounded at base, often
marcescent at flowering time. Lower internodes 2-3 cm long. Median leaves
smaller and more distantly spaced, elliptic-oblong, flat to moderately con-
duplicate, 1-3-nerved, acute at apex, truncate at base. Upper leaves linear,
strongly conduplicate and arcuate, 1l-nerved, 1.5-5 em long, 1-6 cm wide,
acute to acuminate at apex, tapering at base. Upper internodes usually
5-15 cm long. Leaf margins entire, revolute. Cymes terminal and on short
(or occasionally elongate, up to 1.5 dm long) branches from the upper 2-8
nodes, the branches often arching toward tip, the two branches arising from
opposite sides of the same node usually being curved in the same direction,
the inflorescence, or at least its lower part, thus being more or less secund.
Flowers solitary or in small, dense clusters (often in pairs), individually
short-pedicellate or sessile. Cymes, their divisions, and individual flowers
subtended by linear strongly arcuate and conduplicate, acuminate, minutely
ciliclate bracts, those subtending cymes usually 2-3 cm long, 2-4 mm wide,
those subtending individual flowers or pairs of flowers 1-2 cm long, 1-2 mm

207

\\ - Y ea ; if
wi s V4 vv {
\ Va Y
: ie
d Wi a
Gentiana caliculata. a, flowering stem; b, a pair . flowers with
ee bracts; c-d, interior surfaces of corollas, with stamens present

(c) and stamens removed On and e, exterior surface of es slit longi-
tudinally and pressed; f, pisti

208

wide. Calyx tubes uncleft, 6-16 mm long, minutely but copiously puberulent.
Calyx lobes linear, strongly arcuate-recurved, 6-20 mm long, puberulent and
ciliolate, Corolla tubes 2.5-4 cm long, flaring just above summit of calyx
tube, otherwise nearly cylindrical. Corolla lobes reflexed, 6.5-15 mm _ long,
oblong-ovate to nearly orbicular, apiculate, margins often erose-serrate,
especially near base, sinuses subequal. Free portions of appendages con-
sisting of 2 approximately equal, acuminate to attenuate teeth each 1.5-2
mm long and ca. 1 mm wide, usually largely concealed by lobes. Lower 1/3
of corolla tube, lobes, and free portions of appendages crimson throughout,
interior surface with small yellowish-green spots on lower portion of lobes
and adjacent portions of tube, the red pigment extending downward along
veins and as spots or short streaks on interior surface between central and
lateral petal veins. Stamen filaments becoming free a little below middle of
corolla tube. Anthers free, the tips reaching nearly to the level of tips of
the corolla lobes of unopened or pressed flowers, the anthers thus being ex-
serted 6-9 mm from mouth of intact corollas with reflexed lobes. Pistil with
ovary tapering upward into a distinct, slender style, which bifurcates below
the stigmatic surfaces, the stigmas being exserted 2-10 mm beyond mouth
of corolla tube, and with the non-placental zones adjacent to the sutures
relatively wide, otherwise typical of the section. Both anthers and_ style
branches spiraling in age. Fruit slightly exserted at maturity. Seeds ca. 2
mm long, 0.4 mm wide, prominently winged at one end, wing otherwise nar-
row but complete.

Gentiana caliculata is endemic to a region comprising the western portion
of the Neovoleanic Plateau, from Morelos west to Jalisco, and extending
south to the Sierra Madre del Sur in Guerrero (Fig. 10). Here it grows on
grassy slopes in the pine-oak zone, flowering from March through June.

Representative specimens examined:

MEXICO: Guerrero: Puerto Rico, Hinton et al. 14169 (DS, F, GH, MEXU,
MICH, NY, POM, UC, US, WTU). Jalisco: Nevado de Colima, C.G. Pringle
5513 (GH). México: Rincon, Hinton 11201 (CU, F, GH, ILL, MO, NY, TEX,
US, WTU). Michcean: Sierra Torrecillas, Hinton et al. 13690 G&NCB, GH,
NY, US). Morelos: near Cuernavaca, C.G. Pringle 7767 (MEXU, VT).

No type specimen for the name G. caliculata Lex. has been located, and
it is extremely unlikely that any such specimen exists. The only herbarium
known to contain specimens collected by La Llave & Lexarza prior to the
publication of their Novorum Vegetabilium Descriptiones is that of the Con-
servatoire et Jardin Botanique, Genéve (G) (Chaudri et al., 1972; R. Mc-
Vaugh, personal communication). According to information graciously sup-
plied by Dr. M. Dittrich, G. caliculata is not represented among the La
Llave & Lexarza collections there.

The only other herbarium known to house La Llave collections is that of
the Botanische Staatssammlung, Miinchen (M), at which there are four spe-
cimens of Mexican gentians collected by La Llave in 1830, after the Novorum
Vegetabilium Descriptiones had been published. These include one specimen

209

) VER
Nw : (
GTO. Qro. ‘
/ \ i
sek ‘\ HGO. Kiar
JAL. oe a Me es : )
eae, )
MICH.
MEX TLAX.
D.F.
e
® °° .
° MOR
PUE.
GRO
OAX .
a
@ G. caliculata ee. Pi
4 G. mirandae NE

Fig. 10. Documented distribution of Gentiana caliculata and G. mirandace.

of Gentiana bicuspidata and two of G. spathacea, all from Anganguco,
Michoacan, and one of Gentianella amarella (L.) Bérner ssp. hartwegii
(Benth.) J.M. Gillett, from Las Cruces, Distrito Federal. Since none of these
specimens was identified to species by La Llave, they do not indicate to
what species La Llave & Lexarza applied any of the new specific names
published by them.

In the absence of a type specimen for the name G. caliculata Lex.,
whether this species is the same as G. salpinxy Griseb. has long remained
in doubt. Specimens have generally been filed under the name G. salpinx in
herbaria, but some collectors and curators, including C.G. Pringle and, more
recently, E. Matuda and L.B. Smith, have called this species G. caliculata.

The type specimen of G. salpinx had been labeled ‘‘Gentiana calyculata”’
[sic] by Ehrenberg, according to Grisebach (1849) and Bullock (1935), as
had the probable replicate in MEXU. Ehrenberg did not indicate whether
he had identified these specimens as G. caliculata on any basis other than
that of Lexarza’s (in La Llave & Lexarza, 1824) published description of this
species. However, since Ehrenberg was a contemporary of Lexarza’s col-
league, La Llave, in Mexico, was acquainted with the leading authorities on
the Mexican flora at the time, and traveled quite widely in Mexico (Urban,
1897), the possibility that he had some further basis for the identification of

210

this collection seems not unlikely. Accordingly, I have selected an extant
replicate of this Ehrenberg collection as the neotype of Gentiana caliculata
Lex.

Although Grisebach (1849), Kusnezow (1894), and Bullock (1935) all as-
sumed that G. caliculata Lex. and G. salpinx Griseb. were different species,
each of these authors admitted to being acquainted only with the latter.
This assumption appears to have been based almost entirely on Lexarza’s
(in La Llave & Lexarza, 1824) description of the corolla of G. caliculata as
being “‘calyce paulo longior.” In all other respects, Lexarza’s description of
G. caliculata appears to constitute an accurate and relatively detailed de-
scription of the species subsequently called G. salpinx. No other gentiana-
ceous species having red corollas with reflexed lobes, exserted anthers, and
the other traits attributed to G. caliculata by Lexarza is known from the
vicinity of Morelia (formerly Valladolid, latinized Vallisoletum) in Michoa-
can, Consequently it appears virtually certain that the name G. caliculata
was applied to the same species as the later name G. salpinx, despite the
presence of the one discordant element pertaining to the relative lengths
of the calyx and corolla in the original description of G. caliculata,

A number of authors on pollination ecology, notably Grant & Grant (1968),
have observed that certain floral features are commonly associated with
hummingbird pollination, especially in North America. Some of these traits,
such as the prolonged corolla tube, the abundance of nectar, and the eleva-
tion of the ovary above the nectar by a gynophore, prevail throughout sect.
Pneumonanthe and in much of the genus Gentiana. It seems likely, there-
fore, that other Mexican species of Gentiana may occasionally be visited by
hummingbirds, although the effectiveness of these birds as pollinators is
doubtful. In G. caliculata, however, there are a number of floral traits that
appear to be specific adaptations to pollination by hummingbirds. These in-
clude the red corolla; the reflexed corolla lobes, which eliminate a landing
platform that might be used by insects; the exserted anthers and stigmatic
lobes; and the deviation of the flowers from the strictly vertical position
prevailing in the genus.

The relationships of G. caliculata are obscured by the modifications of flor-
al structure involved in the ‘“shummingbird-pollination syndrome.”’ The red
corolla, the relatively long style, and the frequent presence of well-developed
branches within the cyme are distinct departures from the prevalent mor-
phology of sect. Pnewmonanthe, The vegetative portions of G. caliculata,
especially with smaller plants, do, however, resemble those of G. laevigata,
and the free portions of the corolla appendages are similar to those of cer-
tain species in sect. Pneumonanthe native to the Pacific Coastal regions of
northern North America and northern Asia, such as G. platypetala Griseb.
and G. sikokiana Maxim. These features, along with the bracteation of G.
caliculata and its relatively short and stout gynophore, indicate that this
species is correctly placed in sect. Pneumonanthe.

211

7. GENTIANA MIRANDAE Paray, Bol. Soc, Bot. México 21:15. 1957. Type:
MEXICO: Guerrero: faldas del cerro Tlacotepec, Mpio. Tlacotepec,
Paray 2013, 30 Mar 1956, holotype MEXU! isotype ENCB!, photos HAM!
Fig. 11

Tap root persistent, long and slender (ca. 4.3 mm in diameter at summit
and 4 dm long, with 1 fork, in Pringle 1753). Stems 1-3, arching or extending
nearly horizontally from steep slopes (as seen in Hidalgo) to erect (as de-
scribed by Paray, 1957), 2-9 dm long, with short, peduncle-like branches
from upper 1 or 2 nodes, the longer stems also with short leafy branches
from lower nodes, minutely papillose-puberulent in lines near nodes, other-
wise glabrous, often suffused with purple. Leaves medium to dark green,
sometimes suffused with purple, lanceolate to ovate, lower ones obtuse to
acute, upper ones acute to acuminate at apex, rounded at base, those be-
tween the middle of the stem and base of inflorescence largest, 3-8 cm long
and 1.8-3.5 times as long as wide, with 3 or 5 prominent veins. Lower leaves
gradually smaller, the lowest mere scales. Leaf margins minutely denticu-
late, very narrowly revolute except sometimes near base. Upper 2-5 inter-
nodes 2-7 cm long, lower ones gradually shorter. Flowers occasionally soli-
tary, usually in terminal, sometimes imperfectly developed, cymes of 3-5,
and sometimes also solitary or paired at ends of short branches from the
first 1-6 nodes below the terminal cymes, the 2 flowers borne on opposite
cyme branches or terminating the same branch reaching anthesis asyn-
chronously. Each flower subtended by a pair of linear to lanceolate bracts
1-3.5 cm long. Calyx tube uncleft, 7.5-20 mm long, 5-ridged, glabrous. Calyx
lobes erect, linear to oblong, 6-14 mm long, 1-2 mm wide, narrowly carinate,
abruptly acuminate, with margins minutely denticulate proximally, entire
distally. Corolla tube cylindric, 4.5-7.5 cm long, 8-12 mm in diameter. Corolla
lobes erect to slightly divergent, ovate-triangular, 4-6 mm long and about
as wide as long, entire, acute to acuminate, the sinus adjacent to outer edge
of each lobe slightly lower than the other. Free portions of corolla append-
ages obliquely triangular, ca. 2 mm long, acute, entire or sparingly erose.
Corolla tube above summit of calyx tube (except sometimes for lowest 1-10
mm), along with lobes and free portions of appendages yellow in bud,
orange-red at anthesis, becoming scarlet, then purplish brick-red when the
stigma is receptive. Corolla tube not striped on interior surface. Stamen
filaments becoming free at 0.33-0.38 times length of corolla tube; free por-
tions very slender, elevating anthers so that their tips approach or slightly
exceed the level of the tips of corolla lobes. Anthers ca. 3.5 mm long, not
cohering. Ovary 1.8-3 cm long when stigma is receptive, elevated by a gyno-
phore about the same length. Style deeply 2-cleft, the divisions slender, con-
tinuing to elongate and coiling after anthesis (the flowers being strongly
protandrous), the stigmatic surfaces exserted beyond tips of corolla lobes.
Fruits and seeds not seen.

Gentiana mirandae is known only from the type collection (from Guer-
rero) and from the three collections from one Hidalgo locality cited below,

212

Fig. 11. Gentiana mirandae. a, flowering stem; b-c, interior surfaces of
corollas, with stamens present (b) and stamens removed (c), and d, calyx,
slit longitudinally and pressed; e, pistil.

213

which were identified during the present study (Fig. 10). The type collection
was made in openings on wooded slopes with Pinus and Alnus dominant,
at 2200-2500 m altitude, and those from Hidalgo on a steep, rocky roadside
bank in Pinus-Alnus woods about 2100 m. All four collections were made in
March, at which time the plants were in full bloom.

Additional specimens examined:

MEXICO: Hidalgo: El Estribo, carretera Tulancingo-Tenango, km 42,
Gimate L. 509 (EF NCB); carretera Tulancingo-Tenango, km 44, Gimate L.
542 (ENCB); 18.4 km by road NE of Metepec on road to Tenango de Doria,

Like G. caliculata, G. mirandae appears to Ne adapted to pollination by
hummingbirds, as it, too, has long-tubed, red corollas and exserted anthers
and stigmatic lobes. Its corolla lobes, although erect rather than reflexed,
likewise fail to provide a landing platform. The branching pattern of G.
mirandae, its corolla shape and color, and its elongate gynophore, which is
unique among species in sect. Pneumonanthe, give this species an aspect
similar to that of certain species in the Asiatic sect. Stenogyne Franch. The
carpels of G. mirandae, however, have seven vascular bundles each, with
the ovules being distributed over much of the interior surface, and thus con-
form to the pattern of ovarian anatomy reported for sect. Pneumonanthe by
Lindsey (1940) and differ from that of sect. Stenogyne.

The disjunction in the known range of G. mirandae is not readily explic-
able, since pine-alder forests, which constitute the habitat of G. mirandae as
far as is known, occur extensively between the stations of this species in
Guerrero and Hidalgo. Although additional populations may be found in the
future, it seems certain that G. mirandae is genuinely rare and widely dis-
junct in its occurrence. Some differentiation may exist between the Guerrero
and Hidalgo populations, as indicated by Paray’s (1957) description of the
Guerrero plants as being erect and as having larger flowers than those seen
in Hidalgo. Specimens from both regions, however, are so similar in details
of leaf, calyx, and corolla morphology that their taxonomic separation would
be unwarranted.

Interspecific hybrids

Although, in view of the ei ed interfertility among species of sect.
Pneumonanthe, it seems unlikely that the Mexican and Central American
species are separated by ene of incompatibility, hybridization among
these species seems to be extremely rare. Some species are ecologically
isolated from others; G. ovatiloba, for example, being virtually restricted to
alpine habitats, rarely occurs in proximity to species growing at lower ele-
vations. Genetic exchange among the species of the pine-oak of the Mexican
mountains appears to be minimized by the tendency of these species to occur
in small, scattered populations and by the fragmented distribution of the
habitat itself.

214

Specimens cited above as examples of variation within one species in the
direction of its nearest Mexican relative should probably not be considered
to represent the results of interspecific hybridization. Some of these speci-
mens may simply represent intraspecific variability that by chance happens
to result in similarity to a related species in one or two traits. Others may
represent patristically intermediate populations persisting in isolated, rela-
tively stable habitats as remnants of ancestral complexes that existed be-
fore the differentiation of present-day species was complete.

True hybridization between well-differentiated species of Gentiana in Mex-
ico appears to be represented only by a series of collections by L. Paray,
from the vicinity of Contreras, Distrito Federal, mapped in Fig. 6g. Paray
1247 (ENCB) and two of the stems in Paray 1248 (ENCB) are typical of
G. spathacea. Paray 1249 (IKNCB), from the same area, represents G. bi-
cuspidata, with some of the specimens on this sheet showing evidence of
hybridization. Paray 3451 (DS, ENCB, MEXU, MICH, MSC) and the remain-
ing components of Paray 1248 are collections of evident hybrids between
G. bicuspidata and G. spathacea, as Paray suspected in the case of 3451
(in sched.). These specimens have decumbent to ascending stems 2-4 dm
long, some bearing small, leafy branches or tufts in some of the leaf axils.
The leaves are ovate, the larger ones 2.2-4.5 em long, 2.5-3 times as long as
wide, with the apices of the lower leaves obtuse, those of the upper leaves
varying from obtuse to acute. The flowers are borne in small terminal clus-
ters and singly at the ends of peduncle-like branches up to 30 mm long,
the branches usually in pairs, but one member of the pair sometimes termi-
nating in an abortive flower only. The flowers are individually subtended by
ovate bracts 12-20 mm long. The calyces are uncleft, with broadly linear to
ovate-oblong lobes 4-7 mm long. Corolla length ranges from 2.7-3 cm on
one stem to ca. 4 cm on another, The corolla lobes are orbicular or nearly
so, 4-6 mm long, obtuse or minutely apiculate. The free portions of the ap-
pendages are bicuspidate. Paray 367 (ENCB), from the Sierra de Las Cru-
ces, about 10 km to the west of Contreras, consists of highly similar plants
that are probably of the same origin.

Unconfirmed Report

The range of Gentiana affinis Griseb. in Hook., Fl. Bor.-Am. 2:56. ‘‘1840”’
[1837] was said to extend from British Columbia and Alberta south to Cali-
fornia, Arizona, and northern Mexico by Hitchcock (1959). No specimens
were located during the present study that would substantiate this report of
G. affinis from Mexico. There are specimens of G. affinis (as G. interrupta,
a taxonomically insignificant variant of this species) and G. parryi A. Gray
in F, collected by Brother G. Arséne et al. in 1926, that bear labels with the
printed heading “‘Plantes du Mexique.’’ The handwritten locality data, ‘‘Las
Vegas,’ however, doubtless refer to Las Vegas, San Miguel County, New
Mexico, where Arséne is known to have collected plants in 1926 (Ewan,
1950), rather than to a Mexican locality.

215

Kearney et al. (1942) have reported G. affinis (as G. bigelowti, another
minor variant) occurring as far south as Cochise and Pima counties in
southeastern Arizona. Gentiana affinis, therefore, might be expected in the
Sierra de San Bernardino and the Sierra de San Luis in adjacent northeast-
ern Sonora

Detailed descriptions of G. affinis, accompanied by illustrations, have
been published by Hitchcock (1959) and by Gillett (1963).

ACKNOWLEDGMENTS

I express my sincere thanks to: the curators and staff of the herbaria
listed in the introduction of this paper, for the loan of specimens or for their
hospitality and cooperation during my visits to their institutions; to those
at B, K, M, and OXF, for supplying photographs of nomenclaturally im-
portant specimens; to the Consejo Nacional de Ciencia y Tecnologia, for
permission to collect specimens of Gentiana in Mexico; to Drs. John M
Gillett and Rogers McVaugh, for their reviews of a preliminary version of
the manuscript, and for valuable information and suggestions; to Dr.
Thomas Duncan, for his cooperation and assistance with field work; and to
the Board of the Royal Botanical Gardens, for its contributions to the sup-
vort of the research and to the publication of this paper.

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GAUDIN, “I.” .F.G.P.]. 1828-1833. Flora Helve “ee Zurich: Orelli, Fuesslin et

Soci. 7 vols. (liane in vol. Es ne
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Dept. Ade. Publ. 99
GRANT, K. A. and = Cue "1968. Hummingbirds and their Flowers. New York: Co-
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a

NOTES

JUVENILE LEAVES IN OKLAHOMA MARSILEA (MARSILEACEAE)—
During June 1973, while teaching at the University of Oklahoma Biological
Station, I collected specimens of Marsilea that were unusual, at least to
mc, because of the diversity of form exhibited by their leaves. Long a
subscriber to—and a teacher of—the maxim that Marsilea leaves are quadri-
foliolate, I was surprised to see simple, bilobed, trilobed, and other un-
expectedly-shaped leaves as well as ‘‘normal’’ quadrifoliolate ones. The
present note and illustration call attention to the unusual leaves, which
apparently are seldom seen.

The plants, collected 21 June 1973, were rooted in the muddy bottom of
a clearwater pond about 1 mile north of Headquarters, Tishomingo National
Wildlife Refuge, Johnston County, Oklahoma. Growing with them were
Chara, Typha seedlings, Potamogeton diversifolius, P. nodosus, Najas guada-
lupensis, Echinodorus rostratus, Elatine triandra, and Bacopa rotundifolia.
Water at the collection site was about 12 inches deep. The blades of the
quadrifoliolate leaves—one per specimen—were floating; the petiole of
each arose from a rhizome that was very shallowly covered with mud of
the pond bottom. No rhizome I collected exceeded about 8 cm in length.
The ‘‘unusual” leaves, arising each at a different node, were small, not
over 1 cm long (including the petiole). Unfortunately, because I preserved
only individual leaves after removing them from rhizomes, I do not know
if leaf shape corre!ated with position on a rhizome.

These leaves were certainly ‘juvenile’? leaves such as those described
(pp. 74, 75) and illustrated (p. 37) by K. M. Gupta in his book Marsilea
(Bot. Monogr. 2. Council of Scientific & Industrial Research, New Delhi,
1962). According to Gupta, new ‘‘sporelings’’ [i.e., sporophytes] are pro-
duced by germination of perennating organs ‘‘when the water in the ponds
settles down to a suitable depth of two to three feet. As usual the young
plants pass through the well-known [!] juvenile stages producing unifoliate,
bifoliate, trifoliate, quadrifoliate submerged and finally the floating leaves
characteristic of the genus.”

Eames, in Morphology of Vascular Plants. Lower Groups (1936) wrote:
“The earliest leaves of Marsilea are subulate, like those of Pilularia; then
follow, in turn, spatulate; two-lobed, as in Regnellidium; four-lobed; and
finally the four-leaflet type.”’

Later in the 1973 season, as water level in the pond fell, I collected
typical M. mucronata A. Br. on the drying pond margin.—John W. Thierct,
Department of Biological Sciences, Northern Kentucky University, Highland
Heights, KY 41076.

SIDA 7(2): 218. 1977.

219

Figure 1. Marsilea mucronata. Upper row and lower left and center, ju-
venile leaves, 4X; lower right, in square, adult leaf, 4X.

SIDA 7(2): 219, 1977.

220

AMMOSELINUM BUTLERI (UMBELLIFERAE), NEW TO NORTH
CAROLINA—In early April of 1975 I collected an unfamiliar plant of the
Umbelliferae. With the help of Dr. J. R. Massey, Curator of the University
of North Carolina Herbarium, it was identified as Ammoselinum butleri
(Iengelm.) Coult. & Rose and represents a new addition to the flora of
North Carolina. The previous range of this species is given by Mathias
and Constance (1944-45) as Arkansas and Oklahoma to Texas. Correll and
Johnston (1970) extended the range southward to Nuevo Leon and west to
New Mexico.

In North Carolina the plant was found growing on a weed covered slope
with several other plants which it superficially resembles: Alchemilla
microcarpa BOSS: . Reut., Soliva pterosperma (Juss.) Less. and Coronopus
didymus (L.) Sm

To be certain that Ammoselinum was truly established the locality was
revisited in the spring of 1976 by Emily Wood and it was found that the
plants were not only as abundant as in the previous year but had in fact
spread beyond their original boundaries.

Collection data for the Ammoselinum are as follows: Orange County,
North Carolina. University of North Carolina campus, Chapel Hill. On
weed-covered slope south of Kenan Stadium. Flowers white. Plants prostrate
or reclining on other vegetation. D. EF. Boufford 15932, 2 April 1975; Lawn
behind Kenan Stadium, adjacent to Ram’s Head Parking Lot. . W. Wood
2901, 28 March 1976. Vouchers have been deposited at North Carolina Uni-
versity and duplicates will be distributed through the North Carolina Uni-
versity exchange program.— David E. Boufford, Biology Department,
Washington University, St. Louis, Missouri, 63130.

REFERENCES
CORRELL, D. S$. and M. C. JOHNSTON. 1970. Manual of the vascular plants of ‘Texas.
Texas Research Foun tution. Renner, Texas.

MATHIAS, M. E. and L. CC te TANCE. 1944-45. North American flora 28 B: 43-295,

coe i., H Apes aid C. R. BELL. 1968. Manual of the vascular flora of the

Carolinas. Unenay of North Carolina Press, Chapel Hill.

ADDITIONS TO THE LOUISIANA FLORA—During the late 1960’s and
early 1970’s Thieret published several lists of additions to the Louisiana
flora culminating in 1972 with a check-list of vascular plants through the
monocots. Reported additions have been scanty since then except for Allen’s
addition of 19 grasses in 1974. The following note adds 16 new monocots, one
definite and two tentative dicots. Lack of a complete check-list is a serious
handicap to Louisiana collectors particularly those in the poorly covered
northwestern section of the state.

Vouchers on all plants are in the herbarium of Louisiana State University

SIDA 7(2): 220, 1977.

221

at Shreveport (LSUS). All were collected in Caddo Parish.

Sparaganium androcladum (Engelm.) Morong. Very uncommon; in ditch
at Ellerbe and Fluornoy-Lucas roads, 7 miles south of Shreveport. July 6,
1976, MacR. 1838.

Agrostis gigantea Roth. (Agrostis alba of authors, not L.) Uncommon on
shores of Cross Lake at State Fish Hatchery. July 5, 1976, MacR. 1836.
Anthoxanthum artistatum Bios. Locally scattered in sparse woodland on
shores of small lake off Ellerbe road, 9 miles south of Shreveport. May 27,
1976, MacR. 1742

Bromus macrostachys Desf. This oddly distributed species, previously re-
ported from Yonkers, N.Y. and College Station, Texas (Hitchock, Gould)
is not uncommon along a drainage ditch in Betty Virginia Park, Shreveport.
June 12, 1976, MacR. 1770. Identified by Dr. Frank W. Gould.

Cynosurus echinatus L. Uncommon; local on Wallace Lake road about 1%
miles from Wallace Lake. An erratically distributed grass reported for
Arkansas and Oklahoma by Hitchcock but not for Texas by Gould. June 13,
1976, MacR. 1773. Identified by Dr. Goul

Elymus canadensis lL. Common to aeuneant in roadside ditches, open fields
and thin woods. Ellerbe and Norris Ferry roads, 9 miles south of Shreve-
port. June 6, 1976, MacR. 1761.

Vulpia bromoides (L.) S. F. Gray (Vulpia dertonensis (All.) Gola.) Common
along shores of small lake off Ellerbe road, south edge of Shreveport. May
1, 1976, MacR. 1684.

Lolium temulentum L. Common in damp roadside ditches; Norris Ferry
road at DeBroeck road, 13 miles south of Shreveport. April 25, 1976, MacR.
1673.

Sclerochloa dura (L.) Beauv. Eurasian grass widely introduced and estab-
lished through the western states and in New York. Lawn of Willis-Knighton
Hospital, Shreveport. Nov. 1, 1975, MacR. 1596.

Sorghum almum Parodi. Considered to be a hybrid, with S. halepense as
one parent, it is now given specific rank by Gould. Single location but
apparently persistent. Railroad siding at Forbing. Aug. 23, 1975, MacR. 1466.
Eleocharis acicularis (L.) R.&S. Common in shallow running water below
dam on small lake off Ellerbe road, 9 miles south of Shreveport. Aug. 11,
1975. Mac.R. 1438.

Zebrina pendula Schnizl. var. quadrifolia Bailey. A common ornamental,
escaped and growing with Commelina sp. in roadside ditch, Norris Ferry
road, 9 miles south of Shreveport. July 2, 1976, MacR. 1824.

Trillium texanum Buckley has been reported from a few scattered locations
in xas (Nixon, Lewis & Freeman). It is considered a threatened, endemic
species. We can now report an extension of its range into Louisiana. Very
rare in low wet bogs, 2 miles southwest of Rodessa. March 23, 1977, MacR.
2160.

SIDA 7(2): 221, 1977,

222

Hymenocallis caroliniana (L.) Herb. Dry sandy upland woods off Wallace
Lake road. Probably a relic of cultivation but long persistent. July 8, 1976,
Mack. 1848.
Dioscorea quarternata (Walt.) Gmel. Thieret shows this in synonomy with
D. villosa. Following Correll & Johnston it is given species status. Common
in dense woods along Bayou Pierre, south Shreveport. May 6, 1975, MacR.

89.
Habenaria lacera (Michx.) Lodd. Rare in dense woods off Norris Ferry
road, 9% miles south of Shreveport, June 6, 1976, MacR. 1759.
Trifolium hirtum All. This weedy clover is widely but erratically distributed
throughout the United States. It was not included in ‘‘Legumes of Louisiana”
by Alex Lasseigne who identified the plant for me. Very locally abundant
on Wallace Lake road about 1% miles from Wallace Lake. April 26, 1974,
Mack, 1018.

The following two species may not be new to Louisiana but are sufficiently
outside their normal range to be noted:
Hieracium scabrum Michx. Found in a single location at south edge of
TIXast Ridge Country Club, south side of Shreveport. Aug. 13, 1976, MacR.
1923.
Monarda pectinata Nutt. A western species not expected east of western
Texas. Uncommon on median strip of Clyde Fant Parkway, Shreveport.
June 8, 1976, MacR. 1753.—D. T. MacRoberts, Louisiana State University,
Shreveport, 71115.

REFERENCES
ALLEN, C. M. 1974. Nineteen species of grasses (Poaceae) new to Louisiana. La. Acad. of
Science, XXXVII: ae

CORRELL, D. S. and } JOHNSTON, 1970. Manual of the vascular plants of Texas.
Texas Research Fondation Renner, Texas.
oma FLW. 19 The grasses of Texas. Texas A & M University Press, College Station,

ac HC OCK, A. S. rev. A. CHASE. 1950. Manual of the grasses of the United States.
Dover Pobhhin ak New York.
LASSEIGNE, A 973. Louisiana legumes. University of Southwestern Louisiana, Lafayette,

NIXON, B.S.) N. LEWIS, and J. D. FREEMAN. Trillhums in Trouble. Texas Parks &
\" ee XXXV. #1, Jan. 1977
mae ST, J. W. 1963. Additions to the Flora of Louisiana ee 28: 169-170.
1964. More additions to the Louisiana flora, Sida 1: 294-2
1966. Additions to the Louisiana flora. Sida 2: 264-265.
. 1967. Thirty additions to the Louisiana flora. Sida 3: 123-12
. 1968. Additions to the vascular flora of Louisiana. La. Acad. eeiciue XXXII:
91-97,
969, Twenty-five species of vascular plants new to Louisiana. La. Acad. of
Science, XXXIL 78-82.
972. Checklist of the vascular flora of Louisiana. Part 1: Ferns and on allies,
gymnosperms, and monocotyledons. Lafayette Natural ai cory Pees Lafayette, La

223

NEW COMBINATIONS IN MEXICAN PAVONIA (MALVACEAE)—
PAVONIA anisaster (Standley) Fryxell, comb. nov.
Based on: Hibiscus anisaster Standley, Publ. Field Mus. Nat. Hist., Bot.
Ser. 4: 230. 1929. Type: Nayarit, between Tepic and Santiago, alt. 1000
m, 15 Sep 1926, Mexia 634 (GH! UC! US!).

Standley placed this plant in the wrong genus because the material he
studied and described did not have mature fruits, nor did he evidently count
the stigmas (five in Hibiscus, ten in Pavonia). The opportunity to examine
specimens with mature fruits, collected more recently from the same area
(Feddema 820, McVaugh 13375, McVaugh 18836), which are clearly conspe-
cific with the Mexia collection that is the basis for Standley’s name, makes
it clear that the species is correctly placed in Pavonia.

Pavonia anisaster is allied to P. firmiflora Schery, P. racemifera Hooker
& Arnott, P. palmeri (HE. G. Baker) Schery, P. ortegiana (Standley) Stand-
ley, P. spicata Cavanilles, and P. rhizophorae Killip. It differs from these
species most noticeably in having a dark red corolla.

PAVONIA oxyphylla (Mocifio & Sessé ex DeCandolle) Fryxell, comb. nov.

Based on: Hibiscus oxyphyllus Mocino & Sessé ex DeCandolle, Prodr.

1: 455, 1824. Type: in Mexici montibus Xochipici (fl. mex. ic. ined.—

IDC microfiche, 86).

Synonyms:

Pavonia glandulosa Presl, Rel. Haenk. 2: 129. 1835. Type: Mexico,
Haenke s.n. (PR—4 sheets! ).

Pavonia melanommata var. pringleana R. E. Fries, K. Svensk. Vet.
Akad. Hand], 42: 43. 1907. Type: Morelos, near Yautepec, Pringle 8718
(MEXU! MO! UC! US!).

Pavonia glandulosa Presl has been regarded previously as a doubtful
name (Contr. U. S. Natl. Herb. 23: 773, 1923; Leafl. W. Bot. 7: 122. 1954) and
Hibiscus oxyphyllus has generally been retained in Hibiscus, usually as a
synonym 0 . brasiliensis L. (Annuaire Cons. Jard. Bot., Genéve 4:
1900; Contr. U. S. Natl. Herb. 23: 781. 1923; Leafl. W. Bot. 7: 283. 1955).
examination of type material of Presl’s name and of the description ne
illustration upon which DeCandolle’s name is based requires the presentation
of the above new combination in specific rank and the following new com-
bination in varietal rank.

PAVONIA oxyphylla (Mog. & Sess. ex DC.) Fryx. var. melanommata (Rob-
inson & Seaton) Fryxell, stat. et comb. nov.
Based on: Pavonia melanommata Robinson & Seaton, Proc. Amer. Acad.
Arts 28: 104. 1893. Type: Michoacan: Monte Leén, Pringle 4343 (ARIZ!
GH! MO! NY! UC! US—2 sheets!).

SIDA 7(2): 223, 1977.

224

Discussion of the distributional and morphological differences between
these two varieties and further specimen citations may be found in Brittonia
95: 83-85. 1973.—Paul A. Fryxell, Agronomy Field Laboratory, Texas A&M
University, College Station, Texas 77843.

SEED GERMINATION AND SEEDLINGS OF KRAMERIA LANCEOLA-
TA (KRAMERIACEAE)—Krameria lanceolata Torr. (=k. spathulata
Small), sandspur, a suffrutescent low-growing perennial, has thin, wiry rhi-
zomes and large, fleshy storage roots. The small, lanceolate, canescent
leaves are deciduous.

Parasitism was first reported in the genus Krameria in 1910 (Cannon
1910), but the parasitic behavior of Kk. lanceolata was not recorded until
recently (Musselman 1976). Like many species of Scrophulariaceae, Olaca-
ceae, and Santalaceae, Krameria lanceolata is a hemiparasite, forming haus-
toria on roots of neighboring plants. It likewise has a broad host range, at-
taching to trees, shrubs, grasses, and other plants.

Krameria lanceolata occurs from Kansas south to Arizona, Texas, Chi-
huahua, and Coahuila, Mexico with disjunct populations in Florida and Geor-
gia. In Georgia it occurs on sandy ridges along coastal rivers, and in Florida
it is found mostly on deep sands in several northern and central countries.
The largest populations we found were near Tampa, Florida.

Very little is known about the life history of these plants. In a review of
previous work on this genus Kuijt (1969) states: ‘“‘We know virtually nothing
abcut the process of germination’? and, further, ‘“‘the entire process of
germination . . . is unexplained.’’ Our note records, apparently for the first
time, information on germination and seedling development. Observations
were made during a cooperative study between Old Dominion University
and the Southern Forest Experiment Station of the U. S. Forest Service to
survey all root parasites in the Southeast relative to their potential as pa-
thogens of commercial tree species.

METHODS AND MATERIALS

Fruits were collected in June 1976 at the ecology study area of the Uni-
versity of South Florida in Tampa and immediately sent to the Forest Serv-
ice laboratory, Pineville, Louisiana, where they were planted with approxi-
mately 20 commercial tree species. Voucher specimens are deposited in
the herbarium of Old Dominion University (ODU) and the herbarium of the
University of South Florida (USF). Twenty surplus fruits were soaked in
water for 48 hours and the seeds were excised. Eleven seeds enlarged due
to imbibition of water; nine remained the same size. The 11 enlarged seeds
were placed on moist fiber in a germination room maintained at 24° C with
16 hours of light and 8 hours of dark. Only five seeds germinated; they were
planted in pots with Pinus taeda seedings.

SIDA 7(2): 224. 1977.

225

OBSERVATIONS AND DISCUSSION

The ovary of K. lanceolata contains two ovules, one of which always
aborts. We found nothing resembling the situation in kK. grayi reported by
Kuijt (1969) where apparently mature fruits did not contain well-developed
seeds. We estimate that as many as 70% of the fruits had fully developed
seeds in them. The morphology of the seed is similar to that illustrated by
Kuijt as a fruit [sic] of K. parviflora var. glandulosa (Kuijt 1969: 156). The
fleshy cotyledons contain an unusually large amount of starch. A thin testa
covers the seed. None of the seeds planted in pots with potential hosts germi-
nated. We attribute this to the low temperature (24° C) at which pots were
maintained and to the soil used in the pots. Therefore, the information we
present on germination is based on seeds removed from the fruits. How-
ever, this form of germination appears to be identical to that observed in
the few fruits that dehisced and germinated in petri dishes. Germination in
petri dishes was epigeal, in contrast to the suggestion of Kuijt (1969).

After 3 days on moist fiber, the radicle began to elongate and exhibited
positive geotropism. At this stage, the seedling had a well-developed root
cap but no root hairs. Secondary growth began about a week after germi-
nation with a resultant sloughing of cortical cells on the oldest part of the
primary root. Secondary roots developed within 2 weeks of germination.
While only a few seedlings were available for study, no haustorial attach-
ments were noted in those seedlings that had been with their host plant for
10 days. Thus, it appears that the early stage of seedling development re-
sembles many other root parasites in being autotrophic.

Germination and seedling development of Kk. lanceolata is similar to
those of other root parasites such as those Scrophulariaceae, Santalaceae,
and Olacaceae examined in our studies (Musselman and Mann, unpublished).
Except for Striga asiatica (Scrophulariaceae), a germination stimulant is

The one striking feature of the seedling of K. lanceolata is the apparent
absence of root hairs.—Lytton J. Musselman, Department of Biological Sci-
ences, Old Dominion University, Norfolk VA 23508 and William F. Mann,
Jr., U.S.D.A. Forest Service, Southern Forest Experiment Station, Pineville,
LA 71360.

REFERENCES
CANNON, W. A. 1910. The root habits and parasitism of Krameria canescens Gray.
Carn. Inst. Wash. Publ. 129: 5-
KUIJT, J. 1969. a biology of parasitic flowering plants. Univ. Calif. Press, Berkeley.
MUSSELMAN, L. J. 1975. ne ee and haustorial structure in Krameria ae olata (Kra-
meriaceae): A preliminary study. Phytomorphology. 25: 416-422 (pub. 1976).

SIDA 7(2): 225. 1977.

226
NOTICE

The Herbarium of the University of North Carolina-Chapel Hill has a num-
ber of back volumes and issues of the Journal of the Elisha Mitchel Scien-
tific Society. Write Herbarium, Coker Hall 010-A, University of North Caro-
lina, Chapel Hill, NC 27514 for list.

SIDA 7(2): 226, 1977.

7

SIDA susie"

VOLUME 7 NUMBER 3 JUNE 1978
CONTENTS
Sidus Sidarum—ll. Paul A. Fryxell. 227

Sectional and subgeneric names in Gentiana (Gentianaceae).
James S. Pringle. 232

Synopsis of Kochia (Chenopodiaceae) in North America.
Will H. Blackwell, Jr., Mark D. Baechle and Gene Williamson. 248

Seed characters in some native American vetches.
J. Stuart Lassetter. 255

Biosystematics of the Texanae Vernonias (Vernonieae:
Compositae). G. Caywood Chapman and Samuel B. Jones. 264

Mississippi flora: a guide to the ferns and fern allies.
A. Murray Evans. 282

The Chihuahuan Desert species of Ericameria (Compositae:
Astereae). Lowell E. Urbatsch. 298

NOTES: Lobelia nuttallii (C | ) in southeastern Louisiana. 304—Notes on

Coreopsis. 304—Cyperus surinamensis (Cyperaceae): new to Arkansas, Kansas and Okla-

homa. 30

NOTICES 308

US ISSN 0036-1488

SIDA, CONTRIBUTIONS TO BOTANY
Founded by Lloyd H. Shinners, 1962

Editor & Publisher
Wm. F. Mahler
SMU Herbarium

Dallas, Texas, 75275

Associate Editor Assistant Editor
John W. Thieret Barney L. Lipscomb
Northern Kentucky University SMU Herbarium
Highland Heights, Kentucky, 41076 Dallas, Texas, 75275

Contributors to Sida should refer to the latest issues for style, follow the
CBE Style Manual (3rd ed.), and use the ‘adopted’? journal abbreviations

given in Botanico-Periodicum-Huntianum.

Subscriptions: Libraries—$6.00 (U.S.) per year; individuals—$8.00 (U.S.)
per volume; numbers issued twice a year.

©
Sida, Contributions to Botany, Volume 7, Number 3, pages 227-308.
Copyright 1978
by Wm. F. Mahler

SIDUS SIDARUM—II’

PAUL A. FRYXELL
Agricultural Research Service, USDA, Texas A&M University
College Station, TX 77843

NOTE 1. A NEW SPECIES OF SIDA FRGM MEXICO.

Both Standley (1923) and Kearney (1954) state that Sida xanti A. Gray
eccurs both in Baja California and in Sonora in the vicinity of Guaymas.
The plant is localized and seldom collected in the latter area. Careful exami-
nation of specimens collected near Guaymas shows that they differ in
several characters from the plants that grow in the Baja California penin-
sula and deserve recognition as a distinct species, which is described below.

1. SIDA hyalina Fryxell, spec. nov.
Suffrutex erectus. Caules teres, dense atque minute stellato-puberuli. Folia
ovata vel lanceolata, e basi grosse dentata, ad basim truncata, apice acuta,

solitarii, 1.0-2.5 cm longi, supra medium articulati, molli- spabetili, floribus

saepe apicem versus aggregatis; pedicelli 2-5 mm longi. Calyces 8-10 mm

longi, obscure 10-costati, molli-puberuli, semi-divisi in 5 lobis triangularibus,

ad basim late rotundati ubi fructiferi. Petala alba (luteola ubi marcida),
gues m

dum ciliata. Columna staminalis pallida, oe stellato-pubescens, 3-5
mm longa, apice staminifera; filamenta ca. m longa, pallida, in 5 tur-
mis disposita ut videtur; antherae numerosae Te 100), pallidae, ca. 0.7
mm longae. Styli ca. 9, per 3-4 mm colu m staminalem excedentes,
pallidi; stigmata capitata, areas stylorum vix excedentia, fuscata.
Fructus schizocarpium ca. 9 mericarpiorum; mericarpia 4 mm alta, partibus
inferiore manifeste reticulatis in parietibus lateralibus et dorsalibus,

elon re
aleeibes laevibus et apicem versus hyalinis; semen solitarium in basi
mericarpiorum.
TYPE: Sinaloa: Cerros de Navachiste about Bahia Topolobampo; rocky
volcanic slopes with coastal Thorn Forest, 26—30 Sep 1954, H. S. Gentry
14301 (holotype: LL; isotypes: DES, US): road from Los Mochis to Topo-
lobampo, talus slopes and rocky cliffs above salt flats, just north of Topo-
lobampo; diffuse subshrub to 0.8 m tall; at 10:15 a.m. flowers not yet open,
but corolla from previous day apparently yellow, 25 Oct 1973, D. M. Bates
& J. L. Vivaldi 3350 (paratypes: BH, pf); near Topolobampo, 15—25 Sept.
1897, Palmer 199 (paratype: US)

‘The first of this series was published in Sida 6: 1-6. 1975.

SIDA 7(3): 227-231. 1978.

Sida hyalina is named to emphasize the distinctiveness of the hyaline
nature of the mericarp wall, especially in the elongated apical portion,

Distinguishing features of the new species and S. xanti principally concern
characters of the mericarps, which are glabrous to sparsely pubescent in
S. vanti and retrorsely barbed in S. hyalina; the mericarps differ morpho-
logically as depicted in Fig. 1. In addition, S. xanti tends to have longer
peduncles, coarser pubescence, often has viscid pedicels, and usually has
yellow-orange petals fading rose, whereas those of S. hyalina are white
fading yellow. Mericarps of these two species and of S. salviifolia Presl are
illustrated in Fig. 1. The latter species, which occurs from Sinaloa in
western Mexico south to Colombia, is included in the discussion because it
is evidently part of the same species-alliance and is in close geographical
proximity to S. hyalina. A comparison of their mericarps shows a strong
similarity in the retrorsely barbed spines and in the darkened marginal
ribs of the apical part of the mericarp, which ribs by extension become
the spines. Various other species are also a part of this alliance (including,
but not confined to, the North American species S. elliottii Torrey & Gray,
S. tragiifolia A. Gray, and S. lindheimeri Engelmann & Gray and the South
American species S. angustissima St.-Hilaire, S. cerradoensis Krapovickas,
S. regnellii R. E. Fries, S. variegata (Grisebach) Krapovickas, and S.
chalchaquensis Rodrigo). S. chalchaquensis of northwestern Argentina de-
serves direct comparison with S. hyalina because its mericarps also have

Figure 1, Individual mericarps, from left to right, of Sida xanti, S. hyalina,
and §. salviifolia (vouchers: Moran 7025, Gentry 14301, and Fryxell 1155a,
respectively). The arrow notes the hyaline area of the upper mericarp

= 1 mm.

229

a distinctive hyaline area at the apical end, although the two species are
otherwise clearly distinct.

Characters common to members of this species alliance include:
(a) leaf form—a truncate or rounded base, remote dentation extending to
the very base of the blade, and a more or less elliptical shape, varying from
narrowly linear (S. angustissima) to broadly ovate-elliptic in several species;
(b) calyx—similarity of the disposition of the nerves and of pigmentation;
(c) filaments—organized more or less into five groups or phalanges as they
emerge from the apex of the staminal column;
(d) corolla—a tendency to rose pigmentation:
(e) mericarps—usually 8—12 in number. prominently reticulate basally, the
apical (dehiscent) portion relatively wells -developed, broad, and with or
without spines, the spines when present developed as extensions of marginal
costae and retrorsely barbe
The mericarps of this group of species are highly diverse, as Fig 1 shows,
but they seem to be united by certain common morphological features, as
I have indicated. Chromosome numbers are n—=7 for S. angustissima, S.
regnellii, and S. variegata and n=14 for S. lindheimeri (Krapovickas, 1957,
1969). The chromesome number is not known for S. hyalina.

NOTE 2—THE TYPE OF SIDA PROCUMBENS SWARTZ

Kearney (1954, 1958) accepts as S. procumbens Swartz a species commonly
found from the southern United States to northern south America, includ-
ing Mexico, Central America, and the West Indies. He lists the following
syncnyms for S. procumbens: S. diffusa H.B.K., S. filicaulis Torrey & Gray,
S. filiformis Moricand (non Jacquin), S. ae Cavanilles, S. pilosa Cava-
nilles, S. supina L’Heritier, and perhaps S. caespitosa Helwig. Kearney
Sanice (Kearney & Peebles, 1942) expressed doubt that S. diffusa and
S. procumbens are distinct, but later, as stated above, concluded that they
represent the same species, for which he adopted the older name.

Although most recent floras follow Kearney’s usage, a few do not and
adopt alternative names for this species—S. diffusa H.B.K. (Camargo, 1970)
or S. filicaulis Torrey & Gray (Gould, 1989: Correll & Johnston, 1970; Jones,
1975). Shinners (1953) adopts the name S. filiformis Moricand, giving S
filicaulis as a synonym, ignoring S. procumbens Swartz. Swartz’s name is
the oldest of those given and therefore has priority over the others. To
adopt an alternative name, as for example Correll & Johnston do, citing
‘S. procumbens of authors’ as a synonym, implies either that Swartz’s
name is illegitimate or misinterpreted or that it represents a different
species. I can find no serious contention in the literature nor does my
field experience with the plant suggest that more than one species is in-
volved. The validity of Swartz’s name is not in doubt. However, it was
necessary to establish the identity of the plant described by Swartz. There-
fore, the type of Sida procumbens Swartz was examined through the co-
cperaticn and courtesy of the Director of the Stockholm herbarium, where

230

Figure 2. Type of Sida procumbens Swartz.

231

Swartz’s types are preserved.

The type material of Sida procumbens Swartz is a mixed gathering. There
are five separate branches affixed to the sheet (Fig. 2), the collection com-
ing from Hispaniola. Three of these branches represent the species in
question and, in the interest of nomenclatural stability, are chosen as
lectotype material of S. procumbens. The other two branches represent
Sida spinosa Linnaeus and are excluded from the typification of S. pro-
cumbens. Moreover, Swartz’s description conforms to the chosen specimens
and not to the discordant specimens of S. spinosa in two particulars: ‘‘foliis

villosis’’ and ‘‘caulibus procumbentibus hirsutis.’? On the basis of this
isolie tee I conclude that the correct name for this species is Sida
procumbens Swartz.

REFERENCES
CAMARGO G., LUIS A. 1970. Catalogo illustrado de las plantas de Cundinamarca. vol.
IV.
CORRELL, D. S. 1M. C. JOHNSTON. re Manual of the vascular plants of Texas.

Texas Research ed en, Renner. xv-+ 1881 pp.
GOULD, F. W. 1969. Texas plants—a Sane and eco

ogical summary. Texas Agric.
ee F, Ee ee Bids of the Texas Coastal Bend. Welder Wildlife Foundation: Sinton.

KE "ARNEY, oa a Cece A tentative key to the North American species of Sida L. Leafl.
W. Bot. os 138-170.
E ——.. 1958. A tentative key to the South American species of Sida L. Leafl. W. Bot.

" ——— and R. H. Peebles. 1942. Flowering plants and ferns of Arizona. U.S. Dept.
Agric. Misc. Publ. 423. U.S. Govt. Printing Office: W elds 1065
KRAPOVICKAS, A. 1957. Numeros efomosoricos ae ta ceas americanas de la tribu
veae. Rev. Agron. Noroeste Arge entina 5-
1969, aie citotaxonomicas i ee Aceas. “Bonplan 3: 9-24,
SHINNERS, L. H. 1953. [Not te on Sida.] Field & Lab. 21:
STANDLEY, P. C. i 923. Sida, in: Trees and shrubs of ee Contr. U.S. Natl. Herb.
B53) « 761-767

SECTIONAL AND SUBGENERIC NAMES
IN GENTIANA (GENTIANACEAE)
JAMES S. PRINGLE

Royal Botanical Gardens
Hamilton, Ontario, Canada

The genus Gentiana L., as traditionally circumscribed, constituted a large
and heterogeneous group of species. The removal of Crawfurdia Wall.,
Gentianella Moench, Megacodon (Hemsl.) H. Sm., and Tripterospermum
Blume—with Gentianella further divided by some authors into Gentianella
s. str., Comastoma (Wettst.) Toyokuni, Gentianopsis Ma, and Pterygocalyx
Maxim.—is now widely accepted, this restriction of Gentiana having been
adopted in such standard references as Flora Europaea (Tutin 1972) and
Hortus Third (Staff... 1976), and in treatments of Gentianaceae in several
national and regional floras. The species remaining in Gentiana are general-
ly recognized as constituting a natural group, sharing the morphological
features listed in Pringle (1977), with these species being more closely re-
lated to each other than to the segregated groups.

A much narrower circumscription of Gentiana has been advocated by
Holub (1973) and Love & Léve (1975), who restricted the genus to the five
species usually treated as its nominate section. A few authors, e.g., Toyo-
kuni (1965), have taken intermediate positions. These further restrictions
of Gentiana, however, have much less often been accepted. It remains ap-
propriate, therefore, to discuss nomenclature of Gentiana as this genus is
circumscribed in Flora Europaea.

Most 20th century authors on Gentiana have applied the rank of section
to the major subdivisions of the genus. There has been close agreement on
the number of sections recognized and on the assignment of species to sec-
tions. Consequently, it is possible to regard some sectional names as “‘op-
erational,” in that they are the correct names for the sections considered
taxonomically acceptable in recent monographs. These sectional names ap-
pear in capitals in this paper. Other names published at the rank of section
appear in italics, following the discussions of the accepted names with
which, respectively, they are considered synonymous. The sequence of ac-
cepted names follows Flora Europaea. Names of sections not represented
in this flora follow the names of the European sections to which they appear
to be most closely related.

The scope of this paper is strictly nomenclatural. Problems of classifica-
tion, such as transfers of species from one section to another, or the estab-
lishment of additional sections for anomalous species, are not dealt with

‘Contribution No. 19 from the Royal Botanical Gardens, Hamilton, Ontario, Canada.

SIDA 7(3): 232-247. 1978.

233

here. Also excluded is any formal discussion of subdivisions of sections.
Several authors have recognized such taxa, but none of their treatments has
been widely followed, some being too locally oriented for general applica-
tion, or having been established without adequate data on basic chromo-
some numbers. An extensive search of literature on these taxa has dis-
closed none having a name that would affect the legitimacy of the sectional
names accepted here,

IMPORTANT SOURCES OF SECTIONAL NAMES

Reneaulme’s (1611) Specimen historiae plantarum was the source of sev-
eral epithets used by later authors as sectional names. Unitary names in
this work were cited in synonymy at the species level in Linnaeus’ (1753)
Species plantarum. Sectional names derived from these unitary names may,
in most cases, be typified by the species to a according to Linnaeus,
the unitary names were applied by Reneaulm

The status of unitary names published in oe s (1790) Elementa botani-
ca is discussed in Article 20 of the International code of botanical nomencla-
ture (ICBN), 1972 ed. Although incorrectly attributed to Necker, sectional
names derived from names in this work have sometimes been validly pub-
lished by later authors.

The four names published for subdivisions of Gentiana in Froelich’s
(1796) De Gentiana libellus sistens have sometimes been rejected as having
been published sine ord., but Brizicky (1969) has pointed out that they were
validly published as sectional names, noting that the subdivisions were re-
ferred to as sections in the introduction.

The sections into which Gentiana was divided by Froelich (1796) were ac-

to Froelich, although they were altered from plural adjectives to nouns.

Link has often been cited as the author of sectional names in Gentiana.
In Link’s (1821-1822) Enumeratio, names of generic subdivisions were pub-
lished sine ord., and must be rejected (Brizicky 1969). Sectional names
were validly published in Link’s (1827-1833) Hortus Regius, but Gentiana
was not subdivided in this work. Link’s (1829-1833) Handbuch does contain
validly published sectional names in Gentiana, but none is the earliest
available for any section included in the genus as circumscribed here.

In Dumortier’s (1827) Florula Belgica, Gentiana was divided into five
sections, three of them within the present generic circumscription. Brizicky
(1969) and Stafleu & Cowan (1976) have accepted the generic subdivisions
preceded by the symbol § in this work as validly published, formal sections.
Suggestions that the Florula Belgica might have been published later than
1827 have been rejected as unsubstantiated by Stafleu & Cowan (1976); 1827
must therefore be accepted as its date of publication (Article 30, ICBN).

In Gaudin’s (1828-1833) Flora Helvetica, Gentiana was divided into ten

234

subdivisions, six of them still referable to the genus. In this seven-volume
work, the treatment of each genus begins with a key in which species are
grouped according to morphological features. Beginning with Volume 2,
groups of species in large genera, which were described in the keys, were
given names printed in small capitals. In Volume 5, such a subdivision of
Hieracium is referred to as a section (Latin sectio). A recent amendment
to Article 35 of ICBN (Stafleu & Voss 1975, Voss 1976) clearly indicates that
all names for subdivisions of genera designated by the same format in this
flora should be accepted as having been published at the rank of section.

Bunge’s (1829) monograph on Gentiana, Grisebach’s (1837) treatment of
the Gentianaceae in Hooker’s Flora Boreali-Americanae, and Grisebach’s
(1838, 1845) two major treatises on the Gentianaceae of the world brought
the subdivision of Gentiana close to modern practice. In all these works,
the sections are described and their rank is clearly indicated. Some epithets
used as sectional names by Grisebach had previously been used as generic
names by Adanson (1763). These names are interpreted here as new names
rather than as new combinations, since Adanson’s use of the epithets was
not cited by Grisebach; the epithets could have been taken directly from
pre-Linnaean sources; and Grisebach’s (1845) use of the epithet Dasyste-
phana differed greatly from Adanson’s.

Groups of species given Latin names in Huxley’s (1888) phylogenetic
study of the “natural order Gentianeae” (i.e., the family Gentianaceae)
have sometimes been interpreted as subdivisions of Gentiana because they
cut across the traditional circumscription of the genus, some species being
assigned to each of his two ‘‘series.’’ Actually, however, these groups are
suprageneric. Huxley’s Perimelitae included not only a portion of the tradi-
tional Gentiana but also Halenia and Swertia. His even more inclusive
Mesomelitae comprised not only the remainder of Gentiana but also many
other genera, e.g., Sabatia, Voyria, and even Menyanthes. No suggestion
was made that all of these genera should be included in Gentiana; instead
Huxley recommended that Gentiana s. lat. be divided. The two “‘series”’
were further divided, but the subdivisions were assigned no rank and in
some cases comprised more than one accepted genus.

The last of the major contributors to division of Gentiana into sections
was Kusnezow. His monograph of ‘‘subgenus Eugentiana’’ (1894) and his
treatment of Gentiana in Die natiirlichen Pflanzenfamilien (1895) have served
as the basis for most subsequent treatments of Gentiana at the sectional
level.

NAMES OF SECTIONS

Sect. GENTIANA

The first restriction of Gentiana pertinent to its typification was made by
Adanson (1763), who included only G. lutea L. in this genus and transferred
other Linnaean species of Gentiana to new genera. Gentiana lutea has been

235

accepted as the type species of the generic name in all recent publications
on the genus. Any section including G. lutea must therefore be called sect.
Gentiana; any other name is illegitimate.

[Sect.] Asterias Gaudin, Fl. Helv. 2:269. 1828.

Sect. Asterias, as established by Gaudin, included only G. lutea and G.
hybrida Schleich. ex DC., the latter interpreted as a probable hybrid be-
tween G. lutea and G. purpurea L. Previously, the epithet Asterias had been

sed by Reneaulme (1611) as a unitary name for G. lutea and by Bork-
hausen (1796) as the name of a genus consisting only of G. lutea.

[Sect.] Coelanthae Froel., De Gentiana libellus p. 15. 1796. Type species

(Article 7, I : Gentiana lutea

The sectional epithet Coelanthae is derived from Coilantha, a unitary
name used by Reneaulme (1611) for G. purpurea L. However, since Froe-
lich (1796) did not directly equate his sect. Coelanthae with Coilantha Re-
neaulme, his choice of an epithet hardly constitutes a definite indication of
a type. Because sect. Coelanthae, as circumscribed by Froelich, included
G. lutea, its name is superfluous, and G. lutea must be accepted as its type.

[Sect.] Coelanthae Bunge, Nouv. Mém. Soc. Imp, Naturalistes Moscou 1:206.

1829, non Froel., 1796.

Gentiana purpurea and three closely related species are usually placed in
the same section as G. lutea, which they resemble in general habit and with
which they are interfertile, e.g., by Kusnezow (1894, 1895) and Tutin (1972).
However, because of the unique deeply lobed corolla, without appendages,
of G. lutea, some authors, including Gaudin (1828), Bunge (1829), and
Grisebach (1838, 1845), have placed this species in a monotypic section.

Bunge (1829) retained G. purpurea and two related species in sect. Coe-
lanthae (attributed to Froelich) but transferred G. lutea to sect. Asterias.
Since sect. Coelanthae Froel. must be typified by G. lutea, its application
to a section from which G. lutea is excluded is illegitimate. Gaudin (1828)
treated this group as sect. Gentiana, and later authors, e.g., Grisebach
(1838, 1845), generally used the sectional name Coelanthe [sic]. It appears,
therefore, that no legitimate name has been published for a section com-
prising G. purpurea and closely related species but not including G. lutea.
[Sect.] Gentianotypus Dumort., Florula Belg. p. 51. 1827.

This is a superfluous name for a section in which only G. lutea was listed.
[Sect.] PNEUMONANTHE Gardin, Fl. Helv. 2:269. 1828. Type species (Arti-

cle 22, BN): Gentiana pneumonanthe L

Greene (1904) has reviewed the early history of the epithet Pneuwmonanthe.
Tutin (1972), Pringle (1977), and other recent authors have recognized Gau-
din (1828) as the first to have validly published the combination Gentiana
sect. Pneumonanthe, its earlier use by Link (1821) having been sine ord.

236

Sect. Cyane Griseb. in Hook., Fl. Bor.-Am. 2:54. ‘1845”’ [1837]. Type species

(Article 7, ICBN): Gentiana pneumonanthe L.

The epithet Cyane is derived from the unitary name Cyana, applied by
Reneaulme (1611) to Gentiana pneumonanthe. Although Grisebach (1837)
first published the combination Gentiana sect. Cyane in a work in which
only North American species were described, his frequent mention of G.
pneumonanthe and his attribution of the epithet Cyane to Reneaulme clearly
indicate that this species was included in his original concept of this section.
Therefore, G. pneumonanthe must be accepted as the type species of this
sectional name.

Sect. KUDOA (Masamune) Satake & Toyokuni ex Toyokuni, J. Jap. Bot.
35:202. 1960. Based on Kudoa Masamune, J. Trop. Agric. 2:29. 1930 (pro
gen.). Type species: Gentiana yakushimensis Makino, as Kudoa yakushi-
mensis (Makino) Masamune.

Section Kudoa, which was accepted by Toyokuni (1963) but not by Smith
(1967), consists of Asiatic species that in most earlier works were placed
in sect. Frigidae. At least the type species, however, was recently placed
in sect. Pneumonanthe by Smith (1967). The reduction of Kudoa from gen-
eric to sectional status met current requirements of the ICBN.

Sect. STENOGYNE Franch., Bull. Soc. Bot. France 31:375. 1884. Type spe-
cies: Gentiana primuliflora Franch.

This sectional name was published in accord with current requirements.
The only species placed in sect. Stenogyne by Franchet (1884) was G. primu-
liflora. The sectional circumscription was considerably expanded by Kusne-
zow (1894), but G. primuliflora has consistently been included in this section.

Sect. FRIGIDAE Kusn., Trudy Imp. S.-Peterburgsk. Bot. Sada 13:61. 1898,
“Frigida.”’ Type species (Article 22, ICBN): Gentiana frigida Haenke.
Article 21 of ICBN, on the names of subdivisions of genera, requires

that the epithets be ‘‘either of the same form as a generic name, or a plural

adjective.’ Since Frigida is a singular adjective, its orthography must be
changed to Frigidae (Article 32, Note 1, ICBN). Otherwise, the publication
of this sectional name met current requirements.

Sect. Isomeria Kusn., Trudy S.-Peterburgsk. Obshch. Estestvoisp., Otd. Bot.
24(2):198. 1894. Lectotype species (hoc loc.): Gentiana depressa D. Don.
Section Isomeria, when established by Kusnezow (1894), comprised five

species: G. cachemirica Decne., G. delavayi Franch., G. depressa D. Don,

G. emodi Marq. (as G. amoena C.B. Clarke, non Wedd.), and G. loderi

Hook. fil. (now generally included in G. cachemirica). Smith (1961) con-

sidered this grouping to be unnatural, and assigned G. cachemirica (includ-

ing G. loderi) to sect. Pneumonanthe and the remaining species to sect.

Frigidae.

237

Although the sectional name Isomeria is not required in currently accept-
ed treatments of Gentiana, the name is not illegitimate. It seems appropri-
ate, therefore, to designate a lectotype in case any of the species named

(1894, 1895) generalized descriptions of sect. Isomeria most closely. It is,
moreover, a well-known species, which also typifies the name of the series
(sect. F'rigidae subsect. Sympodiae series Depressae H. Sm.) to which it
and G. emodi were assigned by Smith (1961)

[Sect.] CRUCIATA Gaudin, Fl. Helv. 2:269. 1828. Type species (Article 22,

ICBN): Gentiana cruciata L

Although the epithet Cruciata may appear to be a singular adjective, it is
actually “‘of the same form as a generic name.’’ Cruciata was a pre-Lin-
naean generic or unitary name variously applied to species with decussate
leaves or with leaves in whorls of four. Since 1753, it has in fact been used
as a generic name in both the Gentianaceae and the Rubiaceae. Its associa-
tion with Gentiana cruciata can be traced back to Bauhin & Cherler’s (1651)
designation of this species as ‘‘Gentiana minor, seu vulgi Cruciata,’’ and
to other pre-Linnaean applications of ‘“Cruciata,”’ as a noun, to this species
(listed by Bauhin & Cherler 1651).

[Sect.] Erythaliae Bunge, Nouv. Mém. Soc. Imp. Naturalistes Moscou
1:207. 1829. Type species (Article 7, ICBN): Gentiana cruciata L
As circumscribed by Bunge (1829), sect. Erythaliae included G. cruciata.
This sectional name, therefore, is superfluous.

Sect. Tretorrhiza Griseb. in Hook., Fl. Bor.-Am. 2:60. ‘1845’ [1837]. Lecto-

type species (hoc loc.): Gentiana cruciata L.

The epithet Tretorrhiza is derived from a unitary name applied by Re-
neaulme (1611) to G. cruciata. Because sect. Tretorrhiza was first described
in a work on North American plants, G. cruciata was not included in this
publication. Gentiana douglasiana Bong. was the only North American spe-
cies assigned to this section. Grisebach’s (1837) attribution of the sectional
name to Reneaulme, however, constitutes a direct link between G. cruciata
and the original description of this section. Fifteen months later, when
Grisebach (1838) published on Gentianaceae of the world, sect. Tretorrhiza
appeared as a natural group of species, of which G. cruciata was represen-
tative, except that G. douglasiana then constituted a discordant element.
Gentiana douglasiana does not appear to have been of great importance in
the preparation of the original description of sect. Tretorrhiza, since the
section was said to consist of perennial herbs, although G. douglasiana is
annual. Therefore, G. cruciata is interpreted as having been included in all
circumscriptions of sect. Tretorrhiza, and, in accord with Article 7 of the
ICBN, this species is regarded here as the type of the sectional name. This

238

sectional name is consequently considered to be illegitimate.

Sect. Kurroo C.B. Clarke, J. Linn. Soc., Bot. 14:433. 1875. Type species

(Article 22, ICBN): Gentiana kurroo Royle.

Section Kurroo, as established by Clarke (1875), comprised G. kurroo and
three other rosette-forming Himalayan species that Clarke correctly realized
had been misplaced in sect. Pneumonanthe (by Grisebach 1845). Kusnezow
(1894), however, noting the similarity of these species to G. cruciata, placed
them in his sect. Aptera (= sect. Cruciata). As long as G. kurroo and G.
cruciata are included in the same section, the sectional name Kurroo is
not required.

Sect. Aptera Kusn., Trudy Imp. S.-Peterburgsk. Bot. Sada 13:62. 1893.

Since the paper in which sect. Aptera was first described (Kusnezow
1893) dealt only with new taxa, only G. kaufmanniana Regel & Schmalh.
var. affghanica Kusn. and G. biflora Regel ex Kusn. were described under
this sectional heading in this work. The following year, Kusnezow (1894)
indicated the full scope of the section, which included G. cruciata. In this
work, the sectional name Aptera was clearly superfluous. Even if G. kauf-
manniana or G. biflora should be designated the lectotype, it is highly un-
likely that the name Gentiana sect. Aptera would find any legitimate usage,
since these species have consistently been regarded as being consectional
with G. cruciata.

[Sect.] CHONDROPHYLLAE Bunge, Nouv. Mém. Soc. Imp. Naturalistes
Moscou 1:207. 1829. Lectotype species (hoc loc.): Gentiana aquatica L.
Gentiana aquatica is here designated the lectotype of the name Gentiana

sect. Chondrophyllae because it is a well-known, Linnaean species, which

was illustrated and described in detail by Bunge (1829), and from which

Bunge distinguished other species assigned to this section. Bunge’s descrip-

tion of G. aquatica corresponds especially closely to his description of this

section, and the cartilaginous leaf margins, from which the sectional epithet
is derived, are shown in Bunge’s illustration of this species only.

Sect. Eurythalia Griseb., G6tt. Gel. Anz. 1840-815. 1840, non [sect.] Erytha-
liae Bunge, 1829. Lectotype species (hoc loc.): Gentiana coronata Royle.
Eyrythalia was used by Reneaulme (1611) as a unitary name for Genti-

anella campestris (L.) Borner. Borkhausen (1796) applied this epithet to a

genus initially comprising this and three other species, two of them refer-

able to Gentianella s. str. Don (1837) ascribed the generic name Eurythalia

(an orthographic variant, both y and u being used as transliterations for the

Greek letter y) to Reneaulme and Borkhausen. The two species assigned to

Eurythalia by Don, Gentiana carinata (D. Don) Griseb. and G. coronata
oyle, along with G. hugelii Griseb., have corolla traits similar to those of

some Gentianella species as well as those characteristic of Gentiana. It

may be assumed, therefore, that Don was placing newly described species

239

in Eyrythalia Borkh., rather than establishing a new genus at variance with
Borkhausen’s concept of Eyrythalia.

Grisebach (1840) considered his Gentiana sect. Eurythalia to be based on
Eurythalia sensu D. Don but broke the nomenclatural continuity by restrict-
ing the section to the three species named above. The species placed in
Eyrythalia by Borkhausen, except the incongruous G. septemfida Pallas,
were placed in sect. Amarella by Grisebach (1838, 1845). Therefore, Genti-
ana sect. Eurythalia Griseb. must be regarded as a new name rather than
a new combination.

The three species placed in sect. Eurythalia by Grisebach were trans-
ferred to sect. Chondrophyllae by Clarke (1883) and have generally been in-
cluded in sect. Chondrophyllae by later authors. Further studies, however,
may indicate that this group should be restored to sectional status, in which
case a new epithet will be required unless Eurythalia and Erythaliae are
deemed to be sufficiently distinct.

[Sect.] CIMINALIS (Adans.) Dumort., Florula Belg. p. 51. 1827. Based on
Ciminalis Adans., Fam. pl. p. 504. 1763 (pro gen.). Lectotype species
(Holub 1973): Gentiana acaulis L.. as Ciminalis [sp.].

Three species—Gentiana acaulis L., G. pneumonanthe L., and G. verna
L.—were originally included in Ciminalis by Adanson (1763). Each of these
is now placed in a different section of Gentiana. Holub (1973), in designating
G. acaulis the lectotype, noted that the genus Ciminalis had in effect been
restricted by later authors, citing Gleditsch’s (1764) publication of the gen-
eric name Pneumonanthe, typified by G. pneumonanthe, and Moench’s
(1794) listing only G. acaulis, as Ciminalis longifolia Moench, under Cimi-
nalis. Moench did not state that the other species placed in this genus by
Adanson were excluded, but his description of the stigma of Ciminalis was
not applicable to G. pneumonanthe.

Ciminalis was more definitely restricted by Rafinesque (1837), who equated
Ciminalis with Thylacitis Reneaulme (1611), i.e., G. acaulis, and assigned to
it “C. grandiflora, angustif[olia], alpina &c which were Gentliana] aphyla
[evidently a lapsus calami for ‘acaulis’] and 2 var. of Authors.’ Both G.
pneumonanthe and G. verna were transferred to other genera. Since G.
acaulis was retained in Ciminalis, and since Rafinesque said that his Cimi-
nalis was that of Moench, who had attributed its authorship to Adanson,
Rafinesque was restricting Ciminalis Adans. rather than reapplying the epi-
thet. Placement of ‘“‘Ciminalis Raf.” in synonymy with Voyria Aubl. (Index
Kewensis; Merrill 1949) is incorrect, “Gent. aphyla’’ having been inter-
preted as Gentiana aphylla L. (= Voyria aphylla (L.) Pers.) rather than
as an error. Gentiana grandiflora Lam. (now included in G. acaulis), G.
angustifolia Vill., and G. alpina Vill. all apply to closely related taxa in sect.
Ciminalis; their specific epithets have never been associated with Voyria.
Recently, Love & Léve (1975) restricted Ciminalis to species having a base

240

chromosome number nine, including G. acaulis (n = 18), but excluding G.
pneumonanthe (n = 13) and G. verna (n = 14).

Sect. Megalanthe Gaudin, Fl. Helv. 2:270. 1828.

This name has been adopted in some recent publications, but, since its
original circumscription by Gaudin (1828) included G. acaulis, the type
species of the earlier sectional name Ciminalis (Adans.) Dumort., the name
is superfluous.

Sect. Thylacites Griseb. in DC., Prodr. 9:115. 1845.

The epithet Thylacites is derived from Thylacitis Reneaulme (1611), a
unitary name for G. acaulis. Since Grisebach’s (1845) original circumscrip-
tion of sect. Thylacites included the type species of the earlier sectional
name Ciminalis (Adans.) Dumort., this name is superfluous.

[Sect.] CALATHIANAE Froel., De Gentiana libellus p. 57. 1796. Lectotype

species (hoc loc.): Gentiana nivalis L.

As noted by Bauhin & Cherler (1651), the word “Calathiana’’ appeared
in pre-Linnacan names for several low-growing gentians. Particularly prom-
inent in this work were the Calathiana verna of Dalechamps, evidently
equivalent to Gentiana verna L., and Bauhin & Cherler’s own Gentianellis
affinis, Calathiana verna Dalechampu, cum albis staminibus, which was
probably the closely related G. nivalis. These pre-Linnaean names were
presumably important in relation to Froelich’s (1796) use of Calathianae
as a sectional epithet. As described by Froelich (1796), sect. Calathianae
comprised mostly low-growing, alpine species, characterized by infundi-
buliform corollas, whereas species in sect. Coelanthae were said to have
campanulate corollas. Sect. Calathianae sensu Froelich was equivalent to
sect. Calathianae in its present circumscription plus sect. Chondrophyllae
and sect. Ciminalis.

Section Calathianae Froel., with its epithet altered to Calathia, was ac-
cepted by Lamarck & de Candolle (1805). Notably, however, they trans-
ferred G. acaulis, the only species in their work that would now be placed
in sect. Ciminalis, from sect. Calathianae to sect. Coelanthae. The residual
sect. Calathianae sensu Lamarck & de Candolle was thus equivalent only
to sect. Calathianae s. str. plus sect. Chondrophyllae.

Bunge (1829) further narrowed the circumscription of sect. Calathianae.
Gentiana nivalis and closely related species were retained in this section;
G. aquatica and its relatives were transferred to the new sect. Chondro-
phyllae.

Because G. nivalis was included by Froelich (1796) in his sect. Calathi-
anae and fits his description of this section, and because it also figured
prominently in Bunge’s (1829) redefinition of sect. Calathianae, this species
is here designated the lectotype of the name Gentiana sect. Calathianae.
This lectotypification, moreover, is compatible with Holub’s (1793) and

241

Love & Léve’s (1975) treatment of G. nivalis as the type of generic name
Calathiana Delarbre, this being the only species placed in this genus by
Delarbre (1800)

[Sect.] Ericoila Dumort., Florula Belg. p. 51. 1827. Type species (Article

7, ICBN): Gentiana nivalis L

Gentiana sect. Ericoila Dumort. was based on Ericoila Borkh. (pro gen.),
which is an illegitimate name because Borkhausen’s (1796) circumscription
of Fricoila included the type species of the earlier generic name Tretorhiza
Adans. (Holub 1973). The sectional name must therefore be treated as a
new name originated by Dumortier (Article 72, ICBN) rather than as a new
combination, and Gentiana cruciata need not be regarded as its type, even
though it must be accepted as the type of Ericoila Borkh. Dumortier’s
(1827) citation of ‘‘Ericoila Bork.’ does suffice as a reference to a published

description, and his inclusion of two relatively dissimilar Belgian species,
G. cruciata and G. bavarica L., in this section did not indicate any reduction
in its scope. Thus Gentiana sect. Ericoila by implication included G. nivalis,
which is highly similar to G. bavarica in floral morphology and which had
also been transferred to Ericoila by Borkhausen (1796). This name, there-
fore, is regarded as illegitimate.

[Sect.] Gentianella Gaudin, Fl. Helv, 2:270. 1828, non Dumort., 1827.

Section Gentianella Gaudin (1828) comprised G. nivalis and G. utriculosa
L. This sectional name is superfluous because the section included the type
species of sect. Calathianae Froel., and also because it is a later homonym
of sect. Gentianella Dumort.

[Sect.] Hippion (F.W. Schmidt) Gaudin, Fl. Helv. 2:270. 1828. Based on
Hippion F.W. Schmidt, Fl. Boém. Inchoata 2:18. 1793 (pro gen.). Lectotype
species (hoc loc.): Gentiana verna L., as Hippion aestivum F.W. Schmidt.
Bauhin & Cherler (1651) reviewed the early use of “Hippion’’ in the names

of Gentiana species. Their Gentianella quae Hippion was equated by Lin-

naeus (1753) with his Gentiana verna. The genus Hippion F.W. Schmidt

(1793) included G. verna in its original circumscription. Gaudin’s (1828)

Gentiana sect. Hippion was obviously based on Hippion F.W. Schmidt, as

indicated by his citation of this generic name and combinations made there-

in by Schmidt. Gaudin, however, restricted Hippion to three Swiss species,
of which G. verna seems to have been most important in the preparation
of the sectional description.

Subsequent authors have not followed Gaudin in placing G. verna and
G. nivalis in different sections. The basic chromosome number seven pre-
vails in both groups, although, as noted by Gaudin, the groups do differ in
longevity and to some extent in habit.

242

Sect. Cyclostigma Griseb., Gen. Sp. Gentianearum p. 259. ‘1839’ [1838].

This sectional name is illegitimate because the section thus named in-
cluded G. nivalis, the type species of Gentiana sect. Calathianae Froel.,
as well as the type species of other, earlier sectional names.

Sect. OTHOPHORA Kusn., Trudy S.-Peterburgsk. Obshch. Estestvoisp., Otd.

Bot. 24(2):102. 1894. Type species: Gentiana otophora Franch.

Originally, only one species, G. otophora Franch., was placed in sect.
Othophora by Kusnezow (1894). Although the sectional epithet differs in
spelling from the specific epithet from which it was derived, Kusnezow’s
(1894, 1895) consistent use of this spelling indicates that it was intentional,
rather than a typographical error. It may be suspected that Kusnezow
employed ‘‘Othophora”’ as a singular adjective, but it coincidentally has the
form of a generic name, comparable to Otophora Blume (Sapindaceae) and
several other generic names ending in ‘‘-phora.”

—

NAMES OF SUBGENERA

The rank of subgenus in Gentiana has generally been used to designate
groups that are excluded from the genus in its present circumscription.
Since this rank has not been used in any recent works in which this restric-
tion of the genus has been adopted, only an informal discussion of sub-
generic names is presented here.

The first subgenera comprising species still referable to Gentiana were
established by Rafinesque. In his Medical flora (1828), Gentiana was divided
into three subgenera: Rotularia, typified by G. lutea, its name therefore
being superfluous; Pnewmonanthe, its epithet indicating G. pneumonanthe
as the type species; and Eublephis, comprising species now excluded from
Gentiana.

Reichenbach also published the name Gentiana subgenus Pneumonanthe
in 1828. Since Reichenbach’s Conspectus was published in December of that
year (Stafleu 1967), it seems probable that it was antedated by Volume 1
of Rafinesque’s Medical flora, although the month in which this volume was
published has not been determined. Reichenbach (1828) also ie the
name Gentiana sect. Hippion, based on Hippion F.W. Schmidt (pro gen.)
and therefore typified by G. verna

In Rafinesque’s (1837) Flora Telluriana, Gentiana s. lat. was divided into
several genera, some of which were further divided into subgenera. Four
of these subgenera fall within the present concept of Gentiana. Rafinesque
established three subgenera within the residual genus Gentiana. Subgenus
Picriza was typified by G. lutea, its name therefore being superfluous (the
earlier name Rotularia was not mentioned). For subgenus Pleuroglossa
G. punctata and G. purpurea were cited as cotypes. Merrill (1949) listed
G. punctata as the type, but his arbitrary listing of the first of two or more
ie as the type does not constitute binding lectotypification under Arti-

e 8 of the ICBN. Rafinesque’s description of subgenus Pleuroglossa was

243

clearly based almost entirely on G. purpurea, containing elements inapplic-
able to G. punctata, and therefore G. purpurea is designated the lectotype
here. Subgenus Tulbela was typified by G. septemfida Pallas (as G. rossica
Raf.), which was incongruously retained in Gentiana even though closely re-
lated species were removed to segregate genera.

The fourth pertinent subgeneric epithet in the Flora Telluriana is Cutlera.
This epithet had previously been used by Rafinesque as a generic name,
although never validly published as such, and in some works appeared as
“Cuttera,’’ evidently a typographical error. The first species transferred to
this proposed genus were Gentiana saponaria L. and G. villosa L., with
G. catesbaei Walt. being added later (Merrill 1949). In the Flora Telluriana,
however, Cutlera appeared as a subgenus in the new genus Xolemia Raf.
It is not clear which of the species assigned to Xolemia were considered
to be the ‘“‘true Xolemias”’ and which constituted subgenus Cutlera. Gentiana
catesbaei, however, seems to correspond most closely to Rafinesque’s diag-
nosis of subgenus Cutlera, in having relatively open corollas

Rafinesque (1837) also published the name Amarella ene Hippion
in Flora Telluriana. The generic name Hippion F.W. Schmidt was cited in
synonymy, but Gentiana viscosa Ait. (= Ixanthus viscosus (Ait.) Griseb.)
was expressly stated to be the type of this subgeneric name. This species
had never been included in Hippion by Schmidt. Moreover, G. verna, ac-
cepted here as the type species of Hippion F.W. Schmidt, was placed in
Ricoila by Rafinesque. Amarella subgenus Hippion must therefore be re-
garded as a new name rather than a new combination. It could not now be
transferred to Gentiana, as it would form an illegitimate homonym.

House (1924) recognized three subgenera in his treatment of Gentiana in
New York, two of which would no longer be included in Gentiana. The third,
subgenus Dasystephana, was evidently equivalent to much of the present
circumscription of Gentiana. Dasystephana Adans. (pro gen.), indicated to

e the basionym, had comprised only G. asclepiadea L. when established
by Adanson (1763); this species, therefore, must be regarded as the type
species of Gentiana subgenus Dasystephana (Adans.) House. This name is
illegitimate, having been antedated by Gentiana sect. Dasystephana Griseb.,
applied (Grisebach, 1845) to a section comprising only the species now
known as Gentianella thyrsoidea (Hook.) Fabris. House’s inclusion of Pneu-
monanthe Gilib. among the synonyms of Gentiana subgenus Dasystephana
indicates that this name is also superfluous, as a later synonym of Gentiana
subgenus Pneumonanthe Raf.

Holub (1970), who restricted Gentiana to five species, divided the genus
into two subgenera, subgenus Gentiana, consisting of G. lutea only, and
subgenus Coilantha (Borkh.) Holub. Subgenus Coilantha was based on Coil-
antha Borkh. (pro gen.), to which Borkhausen (1796) had assigned only G.
purpurea. Since Holub’s Gentiana subgenus Coilantha included both cotypes
of Rafinesque’s subgenus Pleuroglossa, the latter name should have been
adopted.

244

EXCLUDED NAMES

The names listed below were originally published as the names of sections
or subgenera of Gentiana. However, since none of the species included in
the original circumscriptions of these taxa would now be placed in Gentiana,
and since the epithets (except for Gentianella) have not otherwise been as-
sociated with species included in the present circumscription of this genus,
these names have not been discussed in this paper. The genus to which
each of these taxa would now be referred is given following the name. The
names of genera segregated from Gentianella s. lat. have been employed for
the sake of clarity, although I do not accept the generic status of all these
segregates without reservation.

[Sect.] Amarella Gaudin, Fl. Helv. 2:270. 1828. = Gentianella.

Subgenus Amarelloides Torr. & Gray ex A. Gray, Man. ed. 1 p. 358. 1848.
= Gentianella and Gentianopsis.

Sect. Andicola Griseb., Gen. Sp. Gentianearum p. 213. 1838 [18397]. =
Gentianella.

Sect. Antarctophila Griseb., Gen. Sp. Gentianearum p. 235. 1838 [*‘1839"’].
= Gentianella.

Subgenus Anthepogon (Raf.) Toyokuni, Hokuriku J. Bot. 6:33. 1957, pro syn.
= Gentianopsis.

Sect. Arctophila Griseb., Gen. Sp. Gentianearum p. 250. 1838 [‘1839"]. =
Gentianella.

[Sect.] Chionanthe Gaudin, Fl. Helv. 2:270. 1828. = Comastoma.
Sect. Comastoma Wettst., Oesterr. Bot. Z. 45:174. 1896. = Comastoma.

[Sect.] Crossopetalae Froel., De Gentiana libellus p. 109. 1796. = Gentianop-
sis.

Sect. Dipterospermum (C.B. Clarke) Marquand, Bull. Mise. Inform, 1931: 69.
1931. = Crawfurdia and Pterygocalyz.

Sect. Dasystephana Griseb, in DC., Prodr. 9:117. 1845. = Gentianella.

[Sect.] Endotrichae Froel., De Gentiana libellus p. 86. 1796. = Comastoma,
Gentianella, and Lomatogonium.

Subgenus Eublephis Raf., Med. fl. 1:208. 1828. = Gentianopsis.
[Sect.] Eyrythalia (Borkh.) Dumort., Florula Belg. p. 51. 1827, ‘“‘Eurytalia,”

sensu Dumortier, non [sect.] Erythaliae Bunge, nec sect. Eurythalia (D.
Don) Griseb. = Gentianella.

245

[Sect.] Gentianella (Borkh.) Dumort., Florula Belg. p. 51. 1827, sensu Du-
mortier. = Gentianopsis.

sine ee Gentianella Reichenb., Herbarienbuch p. 133. 1841; Kusn. ee oes
Imp. $.-Peterburgsk. Bot. Sada. 13:58. 1898, superfluous comb. = Coma-
Gane. Gentianella, Gentianopsis, and (sensu Kusnezow) Megacodon and
Pterygocalyx.

Subgenus net (Ma) Toyokuni, Hokuriku J. Bot. 6:33. 1957. = Gen-
tianopsis

Sect. Imaicola Griseb. in DC., Prodr. 9:94. 1895. — Gentianella.
Sect. Megaccdon Hemsley, J. Linn. Soc., Bot. 26:137. 1890. — Megacodon.

Sect. Stylophora C.B. Clarke in Hook. fil., Fl. Brit. India 4:118. 1893. —
Megacodon.

Subgenus Stylophora (C.B. Clarke) Toyokuni, Symb. Asahikawenses 1:143.
1965. = Megacodon

Sect. Tripterospermum (Blume) Marquand, Bull. Misc. Inform. 1931:70.
. = Tripterospermum.

[Sect.] Urananthe Gaudin, Fl. Helv. 2:270. 1828. — Gentianopsis.

ACKNOWLEDGMENTS

Much of the research for this paper was done in the library of the Royal
Botanical Gardens. I am grateful to all whose contributions to the develop-
ment of this library have made such studies possible. Additional research
was carried out at the Lloyd Library and at the libraries of the Gray Her-
barium and Arnold Arboretum, Harvard University; McMaster University;
the University of Michigan; the Missouri Botanical Garden: and the Uni-
versity of Toronto. The hospitality and cooperation of the staff at these li-
braries is greatly appreciated.

Special thanks are extended to Dr, William G. D’Arcy, who called my at-
tention to important literature and provided valuable suggestions. I am also
grateful to Dr. Carroll E. Wood, Jr., for information and discussions that
encouraged development of this paper.

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tion on nomenclature proposals. Taxon 25;

SYNOPSIS OF KOCHIA (CHENOPODIACEAE)
IN NORTH AMERICA

WILL H. BLACKWELL, JR., MARK D. BAECHLE, AND
E WILLIAMSON
Herbarium, Department of Botany, Miami University
Oxford, Ohio 45056

ABSTRACT

Kochia in North America as revised includes three species, two native

and one introduced. A historical summary of work on the genus in North

merica is presented, and typification, distribution, and systematics of the
species are discussed. Specimens from each county recorded are cited for
the native species.

As with most genera of Chenopodiaceae in North America, Kochia has
long been neglected taxonomically and has become a persistent source of
error, Not only are misidentifications frequent in herbaria, but doubt has
continued to exist as to the number of taxa that should be recognized, the
rank they should be assigned, and the distributional pattern they exhibit;
nomenclatural questions also exist. It is toward solution of these problems
that the present study is directed.

Kochia was described by Roth in 1800 based on a single European species,
K. arenaria (= K. laniflora, cf. Flora Europaea 1: 99). Since 1800, additions
to the genus have been substantial; Airy Shaw (1973) provides the recent
estimate of 90 species, these occurring primarily in Europe, Asia, and Aus-
tralia. The complement of species found in North America is small, and
consists of both native and introduced elements.

Kochia first received serious taxonomic attention in North America in
1874 when Watson described two taxa: K. americana [var, americana] and
K. americana var. vestita. Later (1882), Watson described a second native
species, K. californica. In , Rydberg added yet another by elevating
Watson’s var. vestita to the level of species without explanation. The next
milestone for the genus in North America was Standley’s revision (1916),
incorporated in his treatment of the Chenopodiaceae in North American
Flora, Standley recognized four species: K. americana, Kk. vestita, K. cali-
fornica, and the introduced K. scoparia; he recognized no varieties.

Most floristic treatments of Kochia in the United States have accepted
the conclusions of Standley’s revision. A few exceptions are apparent, how-
ever, Weber (1972), in his Rocky Mountain Flora, suggested that Kk. iranica
Bornmueller is a conspicuous component of the Colorado flora and that it
has been confused with K. scoparia. However, we have studied one of
Weber’s collections, Weber 12991 (GH, UC), of ‘‘iranica’”’ and find it to be

SIDA 7(3): 248-254. 1978,

249

merely another specimen of K. scoparia. In fact, no specimens identifiable
as K. iranica turned up in any of the North American collections we ex-
amined. It is thus our conclusion that K. iranica is not a part of the flora
of the United States. Kochia hirsuta Nolte, reported by Davis (1952) as in-
troduced in Idaho, is probably not a Kochia. The short-conic tubercles or
spines of the calyx indicate that it is more likely Bassia. We have examined
the type of Kochia alata Bates; it is just another specimen of K. scoparia.
The varieties of K. scoparia recognized by Steyermark (1963) seem unwar-
ranted as intergradation between them is continuous and geographic pat-
tern lacking.

Although several genera of Chenopodiaceae in North America have a
similar appearance, we encountered the greatest difficulty, particularly at
the outset, distinguishing Kochia from Suaeda. However, we now know that
several characteristics are reliable. The embryo of Kochia is annular and
usually associated with endosperm; Suaeda’s embryo is spirally coiled, the
endosperm lacking, In the absence of good embryos, distinction can still be
made. Suaeda has three scarious bracts subtending each axillary flower or
flower group, these in turn subtended by a leaf or larger herbaceous bract:
in Kochia, scarious bracts are lacking. Kochia stems (native species) are
generally floriferous or potentially so throughout their length; in Suaeda,
the flowers are usually restricted to distal inflorenscences of various sorts.
The calyx of Kochia is cleft no more than half way to the base, while that
of Suaeda is cleft nearly to the base. Finally, in Kochia distinct transverse
wings develop on the back of the calyx lobes in fruit, a feature usually
(though not always) lacking in Suaeda. Number of flowers per leaf axil and
alternate vs. opposite leaves (Davis, 1952), will not provide separation be-
tween these genera. The number of flowers per leaf axil in Kochia is 1-3 (5)
and in Suaeda is 1-3(-9). In both genera the leaves are typically alternate.
We acknowledge Mrs. Christine O. Hopkins for her assistance in distin-
guishing these two genera.

To determine the number of taxa of Kochia occurring in North America
(including introduced, naturalized taxa), the rank they should be assigned,
their distribution, and their typification, North American specimens were
studied from the following herbaria: A, F, GH, MO, MU, TEX, UC, US,
and UT. For comparison, Old World specimens were examined from MU,
TEX, UC, and UT. Australian collections were borrowed from MO and US.
We acknowledge the curators of these herbaria for their cooperation in
loaning Kochia specimens. In the species treatments, based on our herbari-
um and literature study, only those synonyms causing confusion in North
America are listed, in addition to basionyms.

GENERIC DESCRIPTION
Annuals, perennial herbs (often woody at the base), or subshrubs. Leaves
mostly alternate, flat or terete, sessile or petiolate. Flowers small, sessile,
1-5 per axil, bisexual or unisexual (native U.S. species often polygamous),

250

in terminal leafy spikes or the entire branches floriferous; scarious bracts
lacking; calyx 5-lobed, cleft not more than half way to the base, developing
dorsal, horizontal, membranous wings in fruit; corolla lacking; stamens D,
usually exserted; stigmas 2 or 3. Utricle depressed-globose, closely sur-
rounding but not attached to the seed, eventually disintegrating. Seed hori-
zontal and occupying most of the space within the utricle, remaining more
or less enclosed in the calyx; embryo annular around the scanty to copious
endosperm.

SPECIES OF KOCHIA IN NORTH AMERICA
A. Annuals: leaves often petiolate, thin and flattened; flowers restricted to
terminal leafy spikes or spaced one the branches; fruiting calyx wings

0.4-1 mm long, to 1.5 mm broad Kk. scoparia
Pac leaves sessile, terete or essentially flattened: branches po-
tentially floriferous throughout; fruiting calyx wings 1.3-2.5 mm long,
2.5-4 mm broad.

B. ee Loner branched throughout, the pubescence brownish (at
least “-b ees specimens); leaves more or less flattened (only
sehay fleshy), 1.25-3.25 mm broad 2. K. californica

. Br oes ee to the base, the ‘bra nches ‘ascending, ads
simp ic escence white or grayish | or ee — leaves terete
or cee 0.5-1.25 mm broa 3. kK. americana

1. Baer SCOPARIA (L.) Schrader, Neues Journal fiir die Botanik 3(3):
809.

jy cores scoparia L., Sp. Pl. 1: 221. 1753
Kochia alata Bates, Amer, Bot. 24: 52. 1918. (Type: Bates 6767 US !).

Glabrous to puberulent or villous, branching annual, often turning red in
Fall. Leaves petiolate to subsessile, lanceolate or oblanceolate to linear,
3-70 mm long, 0.5-7.5 mm broad, thin and flattened, not at all fleshy. Flow-
ers sessile, in short, leafy spikes or spaced along the branches; anthers
0.5-1 mm long; calyx 1-2 mm long, the lobes 0.5-1.5 mm broad, in fruit de-
veloping membranous wings 0.4-1 mm long and up to 1.5 mm broad. Seed
1-1.8 mm broad.

Kochia scoparia, introduced from Europe as an ornamental, has escaped
from cultivation in the United States, becoming naturalized in a number of
localities. It could be expected to occur anywhere in the United States as
a weed along roadsides or in waste places except in the Southeast. It has
doubtless been cultivated for its red foliage, hence the name “‘fire-bush’”’

r “burning-bush.”’

Variation is extensive in K. scoparia and has led to the proliferation of
both species and varietal names. In attempting to account for the variation
of K. scoparia in Missouri, Steyermark (1963) recognized three varieties:
var. culta Farw., with thread-like leaves; var. pubescens Fenzl, with broad-
er leaves and downy stems; and var. scoparia, with broader leaves and
sparsely hairy to glabrous stems. We have encountered these and other
“forms” of K. scoparia in our examination of specimens, but we find the
variation to be essentially continuous and distinct geographical patterns

251

lacking. We do not see the necessity cr even the advisability of formalizing
the variation encountered in North America with infraspecific Latin epithets.

2. KOCHIA CALIFORNICA S. Watson, Proc. Amer. Acad. Arts 17: 378. 1882.
K. americana var. californica M. E. Jones, Contr. W. Bot. 11: 19. 1903.
Brownish, sericeous perennial from a woody base, paniculately branched
throughout, the branches spreading or ascending, often striated. Leaves
sessile, flat or somewhat fleshy, 2.5-17 mm long, 1.25-3.25 mm broad, dense-
ly hispid or sericeous, the apex more or less acute. Flowers axillary, soli-
tary or in clusters of 2-5, often tomentose; anthers 0.5-1 mm long; calyx
1-2 mm long, the lobes 0.5-1.3 mm broad, becoming winged in fruit, the
wings 2-2.5 mm long and 2.5-3 mm broad. Utricle pubescent, the seed ca. 2
mm in diameter.

TYPIFICATION: In connection with his original description, Watson
(1882) cited two collections without preference: Parry [275], ‘‘near Colton;”’
and Parish [1348], “at Rabbit Springs, San Bernardino County.” By citing
the type locality as ‘‘Near Colton,” Standley (1916) in effect preferenced
the Parry collection as lectotypic. Whether or not Standley meant to make
this choice is nomenclaturally irrelevant. Regardless, we have found no
reason to overturn his lectotypification. Since duplicates of Parry 275 exist
in at least two herbaria (GH, F 134932), and since neither Standley nor
anyone else has selected one as the lectotype, it would seem appropriate
to do so herein. The Field Musum specimen is a mixed collection, Parish
1348 (F 134882) occupying the left side of the sheet and a fragment of Parry
275 the right. Although bearing the same collection number as the Gray
Herbarium specimen, the label of the Parry collection reads ‘‘Mojave,
California,’’ rather than Colton. On the other hand, the Gray pois
specimen of Parry 275 is more ample (two good branches on the sheet),
is not a mixed collection, and the label clearly states ‘‘Colton.’’? Thus, oo
Gray Herbarium specimen seems preferable as the lectotype, and we so
designate it. It is moot to nomenclatural considerations that Colton may in
fact be an erroneous locality. Mojave is more likely the actual collection
cite since Colton is cismontane. It is interesting, and puzzling, that Parry
275 is the only collection, among all the K. californica specimens we ex-
amined, found to be in the fruiting condition, although a number of flowering
specimens have been collected.

DISTRIBUTION: Kochia californica, more restricted in its distribution
than the other native species, K. americana, appears to be limited to south-
eastern California and adjacent southwestern Nevada. The localized distri-
bution, coupled with the dearth of fruiting specimens, led us initially to sus-
pect that it was an introduced species, perhaps persisting locally, but not
setting seed. Australia was considered as a possible source of introduction.
However, our study of Australian and Old World Kochia specimens revealed
no other species closely similar to K. californica. Dr. Paul G. Wilson of the
Western Australian Herbarium, who has recently completed a manuscript

202

on Kochia in Australia, was kind enough to confirm our judgment in this
matter based on color transparencies of herbarium specimens of K. cali-
fornica, which we supplied him. Our appreciation is extended to Dr. Wilson
for his assistance in the solution of this problem. We can only continue to
conclude that K. californica is a native to North America, and that it does
not occur elsewhere.

SPECIMENS Pear (At least one specimen from each county recorded in our
sey is cited): CALIFORNIA: Fresno Co.: 9 mt of Kerman, Hoover 2325 (US); al-

kaline plains, 2.5 1 of Tranquility Junction, along California hwy. 180, Bac igalupi,
Wiggins and Ferris 2670 3 3 nyo Co.: Shoshone Grinnell sv UC). Kerr
near Bakersfield, Rogers s.n ; Rio Br Buena Vis ake, Short 288 (UC) Ls

Angeles Co.: Lancaster, Pee es Abrams and McGregor 509 (GH, US); Lancaster,
Elmer 3717 (F, MO, US); uncultivated alkaline area 5 mi W of Lancaster on Antelope
Valley Road, Ferris and Rossbach 9475 (F, UC). Madera Co.: mi SW of Chowchilla,
Hoover 2567 (UC, US); 16 mi from Madera on Binshaweh Road, a 2619 (UC, US);
San ela Valley, Kennedy s.n. (UC). San Bernardino Co.: 3.5 mi SE of | i
Axelrod 337 (UC); dry lake bed, Rabbit Springs, Mojave Desert, Jepson 5938 (UC);
Victor, rae sn. (MU); Barstow, Jones 387 (GH); Victor, Palmer 224 (US); sandy arid
hillside of Victorville, Mojave — Wheeler 2077 (UC); Desert Wells, Mojave Desert,
Purpus 5731 (UC, US). NEVADA: Nye Co.: 0.5 mi E of house on Ash Meadow Ranch,
Coville and Funston 369 (US)
3. KOCHIA AMERICANA S. Watson, Proc. Amer. Acad. Arts 9: 93. 1874.

x. americana var. vestita Watson, loc. ci

k. vestita (Watson) Rydberg, Flora of Colorado 119. 1906.

Glabrate to gray-white hispid or sericeous perennial with more or less
erect and often simple branches originating at or near the woody base, the
branches frequently stramineous. Leaves fleshy, terete or subterete, 4-25
mm long, 0.5-1.25 mm broad, sessile. Flowers axillary, solitary or in groups
of 2 or 3, often pubescent; anthers 1-1.5 mm long; calyx 2-2.5 mm long, the
ovate lobes 1.3-2 mm broad, crenately winged in fruit, the mature mem-

ranous wings 1.3-2.5 mm long and 2.5-4 mm broad. Utricle glabrate to
villous, the seed 1-1.8 mm broad.

TYPIFICATION: Watson (1874), in his protologue, cites the following col-
lections: Torrey 465, Watson 992, Green s.n., and Wheeler s.n. These should
be regarded as syntypic collections. As in the case of Kk, californica, Stand-
ley (1916) effected lectotypification of K. americana by citing ‘““Western Ne-
vada’ as the type locality. Only the Torrey collection (from Humboldt
Mountains, Nevada) fits this locality, and since the only specimen of Torrey
465 we have been able to locate is at the Gray Herbarium, we accept it as
lectotype of the name K. americana var. americana. Unfortunately, it occurs
on the same herbarium sheet with Watson 992, Wheeler s.n., and an addi-
tional specimen collected by Parry (Parry 260). However, this should not
preclude its service as lectotype.

Following his description of K. americana var. vestita, Watson listed two
specimens: Watson 991 and Burgess s.n. When Rydberg (1906) elevated va-
riety vestita to a species, he did not specify a type. Once again we must
look to Standley who, by indicating the type locality as ‘‘Shores of Great
Salt Lake, Utah,’’ was wittingly or unwittingly choosing a lectotype collec-

253

tion, Watson 991. The Burgess collection is probably from Nevada. We have
seen the U.S. National Herbarium specimen of Watson 991 (US 63408) and
designate it as lectotype of the name Kochia americana var. vestita (iso-
lectotype GH, but mixed with the Burgess collection)

The type of var. vestita is merely one of many variably pubescent indi-
viduals of K. americana we have seen, the variation being essentially con-
tinuous. We do not believe that this random variation in pubescence should
be accorded varietal status, much less that of a species.

DISTRIBUTION AND SYSTEMATIC CONSIDERATIONS: Kochia amer-
icana, known as green-molly or gray-molly, has a much broader distribu-
tion than K. californica. According to our herbarium studies, K. americana
ranges from southwestern Wyoming, southwestern Idaho, and southeastern
Oregon, through Utah and Nevada, to southeastern California, northeastern
Arizona, western Colorado, and southern New Mexico. Unlike K. californica,
fruiting specimens of K. americana are frequent throughout its range.

Prior to Watson’s description of K. americana, specimens in American
herbaria were usually identified as K. prostrata Schrader, a broadly dis-
tributed Eurasian species. This, for example, was the case with Watson
991 (the type of var. vestita). It remains to be finally demonstrated just how
distinct K. americana and K. prostrata are. Watson (1874) suggests the fol-
lowing rather technical characters for separation: the pubescence of the
utricle, the amount of albumin, and the prolongation of the radicle sheath.
Dr. Paul G. Wilson of Australia has agreed that K. americana and K. pro-
strata are similar, but has declined to pronounce on their synonymy in the
absence of more information about K. prostrata. Though K. americana and
K. prostrata are unquestionably similar, we believe them to be distinct spe-
cies based on habit, geographical distribution, and the characters discussed
by Watson.

SPECIMENS EXAMINED or least one specimen from each county re-
corded in our study is cited): ARIZONA: Apache Co.: 9 mi NE of Rock
Point, Peebles and Smith 13539 aS. Navajo Co.: near Aeathiee Eastwood
Howell 6634 (US). CALIFORNIA: Inyo Co.: Death Valley, Bee Mesa,

ro from Kieler to Darwin, Coville and Funston 905 GH, , US);
ee a flats of hwy. 190, 5 mi E of road to Darwin, Muntz re (UT):
Death ae region, W side of Funeral Range, Boundary Canyon, Wolf 6661
(GH). Lassen Co.: Honey Lake Mae Davy 3329 (UC). San Bernardino
Co.: eedoe Brandegee (UC). COLO RADO: Mesa Co.: Gunnison water-
shed, Grand Junction, Baker 928 vices MO, UC); Colorado National Monu-
ee 3 mi S of Fruita, a of we approach to monument, Weber 3770

guire 5871 (GH). Lander Co.: mouth of Birch Creek Canyon, 14 m i from ae
tin, Henning 78 (F, MO, UC); Eastgate, Train s.n. (MU). Lincoln on Cathe-

254

dral Gorge State Park, 3 mi NW of Panaca, Train 2480 (UT). Lyon Co.:
6 mi SE of Rockland, Wellington, Hendrix 644 (UC). ineral Co.: near
reservoir at head of Pickhandle Gulch, Ferris 13106 (GH, UC); 2 mi S-SW
of Sonoma, Bridgeport, Graham 281 (UC). Nye Co.: 2 mi NW of Penelas
Mill, 15 mi NW of Ione, Beach 818 (UC, US); 2 mi below age of the north
rk of Twin a Henning 100 (UC). Washoe Co.: 20 E of Reno,
Archer 6231 (F, MO, UC); northern base of Granite ee 15 mi NE of
Gerlach, ee 5917 (UC). White Pine Co.: 26 mi SW of Ely, shee
D2619 (UT). NEW MEXICO: Dona Ana Co.: Mesquite, roadside, Low
Sonoran Zone, Fosberg $3452 ie OREGON: Harney Co.: Island Ranch,

near Burns, Griffiths and Morris 720 (MO); Narrows, Peck 1657 (G
Malheur Co.: forks of te - eur, Cusick 1111 (US). UTAH: Box Elder ae
h

Garrett 5389 (F, UT). ery Co.: hillsides of East Ferron, Cottam 5200

Millard Co.: Painter Spring, Fautin 8931 (); Milford, Jones 1491 (A F,
one Piute Co.: along Sevier River, Rydberg and Carlton 6929 (US). Salt
Co.: near Garfield, Flowers 952 (UT). San Juan Co.: 0.5 mi E of
nee and Jaques Trading Post, 3 or 4 mi W of seats National Monu-
ment, Cutler 2757 (GH, MO). Sevier Co.: Glenwood, Ward 154 (MO, US).
Tooele Co.: Stansbury Island, Great Salt Lake, Garrett 5357 (F); W of
Grantsville, Harris C2056 (MO). Utah Co.: near railroad track between
Bringhurst and Lehi, Garrett 5206 (F, UT). Uin tah Co.: desert SE of Jenson,
Graham 8297 (F). Washington Co.: Valley of Virgin, near St. George, Parry
H, MO, US). Wayne Co.: Blue Valley, Stanton 515 (UT). WYOMING
Carbon Co.: Fort Steele, Tweedy 4496 (US). Sublette Co.: plains between
Eden and Big Piney, E. and L. Payson 2590 (MO, UC). Sweetwater :
Rock Springs, Degener ner Reiler 16101 (MO); Poin t of Rocks, Merrill and
Wilcox 649 (GH, US). Uinta Co.: near Hampton, Patterson and Beaty s.n.
(F).

REFERENCES
AIRY SHAW, K. 1973. J. C. Willis: A Dictionary of the Flowering Plants and Ferns,
8th edition. Cambri University Press, London.
BALL, P. W. in T. G. TUTIN ef al. 1964. Kochia in Hiss Europaea 1: 98-99.
DAVIS, R. J. 1952. Flora of Idaho. Brigham Young Univers ity Press, Provo, Utah.
ROTH, W. A. 1800. Kochia. Journal fiir die Botanik 1: 306-308.
RYDBERG, P. A. 1906. Flora of Colorado. Bulletin of the Colorado Agricultural Experi-

=

1 00.
STANDLEY, P. C. 1916. ea in North American Flora 21(1): 3-93
STEYERMARK, J. A. 1963 . Flora of Missouri. Iowa State University Press, Ame
WATSON, S. 1874. A revision oF ie North American Chenopodiaceae. Proc. hee Acad.

82. | Contributions of American botany. Lists of plants se ge gee
Texas and northern Mexico, collected chiefly by Dr. E. Palmer in -80. loc
17: 316-361.

WEBE R, W. 1972. Rocky Mountain Flora. University of Colorado Press, Boulder.

SEED CHARACTERS IN SOME NATIVE
AMERICAN VETCHES’
J. STUART LASSETTER
Department of Biological Sciences, Eastern Kentucky University
Richmond, Kentucky 40475

INTRODUCTION

Hermann (1960) and Gunn (1965, 1970, 1971) have used seed characters
for identifying species of Vicia (vetch). Hermann used the relative hilum
ength, which is a percentage figure calculated as hilum length/seed cir-
cumference x 100. Gunn (1970) lists four seed characters as being reliable
in identification: seed circumference, relative hilum length, hilum shape,
and relative pesition of the lens to the hilum. The lens or strophiole is a
protuberance or swelling on the seed margin between the hilum and chalaza
and is derived from integumentary tissue (Kopooshian and Isely, 1966).

The relationship of Vicia ludoviciana Nutt., V. leavenworthii T. & G., V.
exigua Nutt. and the varieties of each have been interpreted differently by
recent workers (Shinners, 1948: Turner. 1956; Isely, unpublished manu-
script; Lassetter, 1972). During a taxonomic study of this native complex
cecurring from Alabama to California, and northern Mexico into the Rocky
Mountains, Lassetter (1972) examined seed characters to evaluate their
taxonomic usefulness. The features studied were relative hilum length,
seed diameter (substituted for circumference because the seeds are essen-
tially spherical), hilum shape (determined by measuring the hilum width
at each end), color of testa, and color of mottled portion of testa (if present).
The distance between the lens and hilum was not measured because it is
essentially identical in these vetches.

MATERIALS AND METHODS
A total of 60 seed samples was analyzed, Except for four samples, each
one contained field-collected seeds from many plants of a single population.
The samples, then, were collected from populations and were not just seed
from a single plant. The remaining four samples were obtained from green-
house specimens grown from seeds taken from herbarium sheets.

Each sample was poured into a petri plate with 25 evenly spaced marks,

* Journal Paper No. J-8496 of the Iowa Agriculture and Home a hee Experiment
Station, Ames, Iowa. Project 1814. Field work was supported by N.S.F. Grant #GB-27935,
5 ; S .

‘ the
barium, supported by the Sciences and Elomantees Research Institute, were used in the
preparation of this paper.

SIDA 7 (3): 255-263. 1978.

Table 1. Relative hilum lengths in the V. ludoviciana complex.

Taxon

Hermann Gunn

(1960 (1965,

1971)

V. ludoviciana var. ludoviciana 25—33 30
(= defunct var. typica)

V. ludoviciana var. laxiflora none

given

V. ludoviciana var. texana 23

V. leavenworthii var. leavenworthii ca. 25 22
(= defunct var. typica)

V. leavenworthii var. occidentalis 19—21

Louisiana type
V. exigua var. exigua 20 14—17

Lassetter

25.9—37.9

16.2—21.8

17.7—25.8

19.0—22.6

10.0—14.7

45.6

19.5—21.6

9&6

Number
of
samples

16

257

and data were taken from the 25 seeds on or nearest the marks. Observa-
tions were made at 10x or 20x through a stereoscopic microscope equipped
with an ocular micrometer. Measurements were recorded to the nearest
0.1 mm except for hilum width, which was recorded to the nearest 0.01 mm

The colors were subjectively observed and numerically recorded (see
Lassetter, 1972)

A correlation coefficient matrix was computed to show related variation
between the following: seed diameter, relative hilum length, hilum length,
hilum width at wide end, hilum width at narrow (ens) end, base color of
testa, and color of mottling. Means of the data were used as input. The
longitude and latitude of the mass collection site were also entered as data
to determine if any correlation between site and seed characters existed.

RESULTS AND DISCUSSION

Table 1 contains the relative hilum length as reported by Hermann (1960),
Gunn (1965, 1971), and from this study. Values from Hermann (1960) are
for species only. In Gunn’s data, values for V. ludoviciana var. ludoviciana
and V. leavenworthi var. leavenworthii are from Gunn (1971); values for
cther taxa are from Gunn (1965). The number of samples refers to Las-
setter’s data, which are the minimum and maximum sample means for
each taxon. The Louisiana type is a vetch previously recognized as V. ludo-
viciana var. ludoviciana, but which has characteristics closely resembling
V. leavenworthii var. leavenworthii, and it was studied as a separate entity.

The names used by Hermann, Gunn, and Lassetter in Table 1 represent
the same taxa concept except for one case. Vicia exiguda var. exigua sensu
Hermann occurred from California to Texas. Vicia exigua var. exigua sensu
Gunn occurred from California to Arizona, New Mexico, and Utah, and was
not present in Texas, the southern Texas plants being assigned to V. leaven-
worthii var. occidentalis. Vicia exigua var. _exigua sensu Lassetter occurred

the southern Texas plants, being assigned to V. leavenworthii var. occi-
dentalis. Some of the differences in Table 1 between my data and that of
previous workers may be due to differences in taxa concept and in applica-
tion of taxa names.

The overlap of relative hilum length is clearly shown (Fig. 1) when the
mean values for each population sample are arranged in graphical form.
Three main breaks in the values are present. The Louisiana type is distinct,
with 46 percent. All the other samples with values above 30 percent are
V. ludoviciana var. ludoviciana, and only one sample of this taxon was less
than 30 percent. The three samples with values of 10-15 percent are V.
pint tiece var. occidentalis. The largest group of samples falls into the
16-26 percent range and includes all types except V. leavenworthii var.
occidentalis and the Louisiana type.

If a plus-one standard deviation and a minus-one standard deviation of
each sample are considered, the resulting overlap of values is great enough

VICIA LUDOVICIANA VAR. LAXIFLORA

VICIA LUDOVICIANA VAR. TEXANA

VICIA LEAVENWORTHII VAR. LEAVENWORTHII

VICIA LEAVENWORTHII VAR. OCCIDENTALIS
VICIA EXIGUA VAR. EXIGUA

2) LOUISIANA TYPE — [EEE

0) | EH VICIA LUDOVICIANA VAR. LUDOVICIANA

Y/|\|I

NUMBER OF POPULATIONS
on)

lO l2 14 I6 18 20 22 24 26 23 30 32 34 36 38 40 42 44 46

RELATIVE HILUM LENGTH

Figure 1. Means of relative hilum lengths.

8&6

259

to produce a continuum of values from 9 percent (V. leavenworthii var.
occidentalis) to 40 percent (V. ludoviciana var. ludoviciana) (Lassetter,

97 If the total range of variation in each sample is considered, the
breaks in Fig. 1 at 13 percent and 40-45 percent disappear (Lassetter, 1972).

From Table 2, the diameter of most seeds is between two and three mm,
with only a few types having larger or smaller seeds. The overlap that exists
precludes delimiting these vetches by seed diameter.

Gunn (1970, 1971) gives the hilum shape of V. ludoviciana and V. exigua
as oblong (with parallel sides) and, of V. leavenworthii, as wedge (with
tapering sides). My data confirm the oblong shape for the first two species,
but variation in hilum shape within individual samples of V. leavenworthii
was observed. Some population samples contained seeds with oblong hila,
as well as seeds with wedge-shaped hila. Because of intrapopulational
variation, this character is not wholly useful inasmuch as all three species
can have oblong hila.

The base color and mottling also varied within individual samples. Some
samples were uniformly colored, and others consisted entirely of mottled
sceds. Other samples, however, contained a mixture of mottled and uni-
formly colored seeds. I believe that color variation is not due to the stage
of seed maturity, because seeds were collected from plants with com-
pletely ripened legumes.

Table 3 contains correlation coefficients of longitude, latitude, and seven
seed characteristics. The highest correlation is between hilum length and
relative hilum length. This correlation indicates that these two characters
are related, but are not dependent on the diameter of the seed, with which
they correlate at 0.36 and 0.06, respectively. The wide-end and narrow-end
width of the hilum correlate at 0.92. The usefulness of hilum shape was
discussed earlier. The wide-end width of the hilum correlates at 0.50 with
seed diameter, indicating that hilum width is somewhat related to seed size.

The distribution of the V. ludoviciana complex is primarily east-west, and
longitude correlated with hilum length and relative hilum length at 0.50 and
0.66 respectively. The samples with the shortest hila and shortest relative
hilum length are from the arid southwestern United States. This correlation
may indicate an adaptive character of survival value in that environment.
Many papilionoid legumes are termed hard-seeded because their seed coats
are impervious, and do not imbibe water (Hyde, 1954). The hilum of hard-
seeded legumes functions in producing the proper level of seed dessication.
Once the moisture content has dropped to about 14 percent, the seed coat
beccmes impermeable and further dehydration is accomplished by moisture
loss through the hilar fissure (Hyde, 1954). A small hilum could be ad-
vantageous in reducing excessive moisture loss from seed in extremely
arid environments.

According to Gunn (1971), other native North American vetches occurring
in the same general xeric areas also have rather small relative hilum
lengths: V. americana, 20-33%; V. leucophaeca, 23%; and V. pulchella, 26%.

Table 2. Seed diameter in mm.

Taxon Min. Minus One Mean Plus one

S‘d. Dev. Std. Dev. Max.

V. ludoviciana var. ludoviciana 1.80 2.08 2.32 2.56 2.68
(= defunct var. typica)

V. ludoviciana var. laxiflora 1.92 1.99 2.17 2.35 2.30

V. ludoviciana var. texana 1.80 1.97 210 2.33 2.35

V. leavenworthii var. leavenworthii 1.92 2.07 2.30 2.53 2.78
(= defunct var. typica)

V. leavenworthii var. occidentalis 2.12 2.09 2.43 2.77 2.90

Louisiana type 2.60 2.62 2.77 2.92 3.30

V. exigua var. exigua 2.75 2.67 2.94 3.21 3.23

09%

Table 3. Correlation coefficients of longitude, latitude, and seven sced characteristics.

Longitude Latitude Seed Base Mottling Hilum Hilum Hilum Relative
diameter color color length width at width at hilum
wide end narrow end length

Longitude 1.00

Latitude 37 1.00

Seed diameter 34 43 1.00

Base color 24 O01 .O1 1.00

Moitling color Al .08 05 10 1.00

Hilum length .O2 10 36 .06 27 1.00

Hilum width at wide end .10 387 36 27 Ag 33 1.00

Hilum width at narrow end .07 38 50 .26 .28 39 .92 1.00

Relative hilum length .66 .02 .06 .03 we .94 22 a | 1.00

196

262

Additional native species with a relative hilum length of 75% (V. acutzfolia,
V. caroliniana, V. floridana, V. gigantea, V. minutiflora, V. ocalensis) are
mostly of the eastern and southeastern United States, where habitats are
more mesic. Vicia cracca (33%) is circumpolar, and V. humilis (30%) is of
wet pine woodlands and meadows of Mexico.

CONCLUSIONS

All the characters evaluated are variable. The overlap between the three
species and between varieties within species is evident. In addition, seed
coloring and hilum shape varied within individual samples, and because of
intrapopulational variation cannot be used effectively to differentiate the
taxa.

The relative hilum length and other seed characters may be useful for
identification of other groups of Vicia species and are valuable in combina-
tion with other morphological characteristics for identification purposes.
Because of the overlap due to wide variation demonstrated in these charac-
ters, I believe it unwise to base identification of taxa in this complex on seed
characters alone.

The distribution of most taxa of the complex is centered in a different
area (Lassetter, 1972). Portions of ranges often overlap, but the centers of
distribution usually do not, Occurrence of the taxa of this complex is in
part related to edaphic factors (Shinners, 1948: Turner, 1956; Lassetter,
1972). The intraspecific differences in relative hilum length of varieties may
indicate an ecotypic adaptation of those taxa to other nonedaphic environ-
mental factors.

ACKNOWLEDGEMENTS

I express sincere gratitude to Duane Isely for his suggestion of the initial
dissertation problem and his guidance throughout this study, which repre-
sents a portion of a dissertation submitted as partial fulfillment of the the
requirements for the Degree of Doctor of Philosophy in Botany, Iowa State
University, Ames, 1972. I also appreciate his review of this manuscript.

Mr. Jim Cheek, of the Eastern Kentucky University computer center pre-
pared the correlation matrix.

REFERENCES
GUNN, C. R. 1965. The Vicia americana complex (Leguminosae). Ph.D. Dissertation.
lowa gee Univ., es aes. (No. 66-2961. Uniy. Microfilms, Ann Arbor, Mich
970. A key and cece for the ae ae one hundred ccs of Viel ia (Le-
peTTTen Proc. Int. Seed Test. Assoc. 34:733—79
1971. Seeds of native and ante ee of North America. U. S. Dep.
Agric. oes Handb. 392
oe RM. ANN, F. J. 1960. Weeehes of the United States
S. Dep. Agric. Agric. Handb. 168.
ae E. O. C. 1954. The function of the hilum in some Papilionaceae in relation to
ripening of the seed and the permeability of the testa. Ann. Bot. 18:241—256
KOPOOSHIAN, H., and D. ISELY. 1966. Seed character relationships in the Leguminosae.
Proc. lowa Acad. Sci. 7 :

native, naturalized and cultivated.

263

ee re S. 1972. A biosystematic study of the Vicia ludoviciana complex (Legu-
minosae). Ph.D. Dissertation. Iowa State Univ, Ames. (No. 73-9458. Kerox Univ. Mi-
crofilms, pe Arbor ich.

SHINNERS, - = 19 48. The weeks and pea vines (Vicia and Lathyrus) of Texas. Field
Lab. 16:18—

TURNER, B. a 1956. The legumes of Texas. Univ. of Texas Press, Austin.

BIOSYSTEMATICS OF THE TEXANAE
VERNONIAS (VERNONIEAE: COMPOSITAE):

G. CAYWOOD CHAPMAN? AND SAMUEL B., JONES
Department of Botany, The University of Georgia, Athens, Ga. 30602

RACT. In the most recent monograph | of Vernonia in North America,

comparative morphology, biochemical systematics, transplant studies, hy-
bridization experiments, and geographical distribution is presented and
used to establish a new pene ace Two species are now recognized in
the group: V. texana, from the south central United States, and V. greggii
with four subspecies from the Sierra Madre Oriental in Mexico.

INTRODUCTION

The genus Vernonia Schreb. is largely tropical with about 800 to 1,000
species distributed both in the Old and in the New World. Gleason (1906)
divided the North American members of section Lepidoploa into two sub-
sections and seven series. The species from the eastern United States and
northeastern Mexico were included in subsection Paniculatae series Verae.
Gleason recognized a number of species-groups within series Verae, each
characterized by its own morphological features and geographical distribu-
tion. Four species were included in the Texanae species-group: V. ervend-
bergii Gray, V. greggii Gray, V. schaffneri Gray, all from the Sierra Madre
Oriental of northeastern Mexico, and V. texana (Gray) Small, from the
south-central United States (Gleason, 1906, 1922). The loose inflorescences,
the hemispheric to broadly campanulate involucres, and the southwestern
distribution characterize this group (Gleason, 1906).

Gleason (1923) suggested that the North American Vernonias could be
traced back to an origin in South America. His theory was based on the
increase toward the south in the number of species and on what he con-
sidered to be the more primitive morphology of the tropical species. He
theorized that Vernonia followed two migration routes into North America.
The eastern route was through the Windward Islands into the Greater An-
tilles. The western route was through Central America and Mexico into the
southwestern United States through Texas and, hence, north into the east-
ern one-half of the United States. Gleason (1923) believed that the Texanae

1wWe thank the curators of the herbaria who kindly loaned specimens (UARK, GA, GH,
MO, MEX, NY, SMU, LAF, ore US). This study is part of the Ph.D. ee of
the senior wiathGe presented to the Graduate School of the University of Georgia. It was
supported by a NSF Doctoral eee Improvement Grant and by NSF grants to SBJ.
2 Present address, Gordon Junior College, Barnesville, Georgia 30204,

SIDA 7(3): 264-281. 1978.

265

species-group is the most primitive in series Verae.

e taxonomic history of the group is not complicated. Torrey and Gray
(1841) cited a X form of Vernonia angustifolia Michx. as having leaves
which were longer than the q@ and 8 form. Also, they recognized the form
as occurring in Louisiana and Texas, while the q and § ranged from Florida
to North Carolina. Gray (1844) followed the earlier treatment of Torrey and
Gray (1841) for V. angustifolia var. texana. He separated this variety pri-
marily on the shape of the involucral bract tips. Trelease (1888) changed
the name of V. angustifolia to V. granienifolia (sic) var. texana in Branner
and Coville’s (1891) ‘‘Plants of Arkansas’’; however, V. gramienifolia is a
later homonym. Kuntze (1891) changed the generic name of Vernonia to
Cacalia and applied the name C. angustifolia var. texana to a specimen from
Cairo, Mississippi, located in the extreme northeastern corner of the state.
Twelve years later Small (1903) elevated var. texana to the rank of species.

Vernonia ervendbergit and V. schaffneri were described as species by
Gray (1882) in his ‘‘Novitiae Arizonicae’’. In this same publication Gray
recognized two extremes of V. greggii as var. greggii and var. palmeri.
HKighteen years later, Gleason (1906) eliminated var. palmeri, and in 1937,
Standley described V. taylorae; a name that has more or less been ignored.
Throughout the experimental sections of this study, Vernonia ervendbergii,
V. greggii, V. schaffneri, and V. texana, are referred to sensu Gleason
(1922).

There has been a great deal of confusion and uncertainty with regard to
the identifications of the three species from Mexico. This lack of under-
standing is due primarily to their similar morphology and to lack of collec-
tions. There also has been some difficulty in identification due to hybridiza-
tion between Vernonia texana and several sympatric species in the United
States. The chromosome numbers for Vernonia greggii and V. texana have
been previously reported as n = 17 by Hunter (1964) for the former and by
Jones (1970) for the latter. The chromosome numbers of V. schaffneri and
V. ervendbergii have not been previously reported. Experimental hybrids
had not been made previously between Vernonias from the United States
and Mexico.

The purposes of this study were: (1) to investigate their cytotaxonomy;
(2) to determine whether or not the Mexican Texanae could be intercrossed
with each other and with other members of series Verae native to the United
States; (3) to better delimit the geographical ranges of the Mexican taxa;

from anatomical, biochemical, cytological, ecological, hybridization, and
morphological studies.
RESULTS
Anatomy and morphology. The upper and lower leaf blade epidermis,
leaf trichomes, leaf venation, leaf anatomy, pollen grains, achenes an
pappus were compared among all four taxa (cf. Appendix), The observed

266

differences in these features were slight among the taxa. Leaves of Verno-
nia texana have two palisade layers whereas the three Mexican taxa have
a single palisade layer. The achenes of V. texana tend to be densely cov-
ered with trichomes whereas the others are not. The anatomy and morpholo-
gy of the three Mexican taxa are quite similar and in general resembie
that found in the other members of subsection Paniculatae series Verae
(Jones, 1972; Urbatsch, 1972; Faust, 1972; King and Jones, 1975).
Chromosome numbers. Acetocarmine squashes of pollen mother cells
from 26 collections of the four taxa all yielded n = 17 (cf. Appendix). These
additional counts confirm earlier reports (Hunter, 1964 and Jones, 1970) and
conform to that found in series Verae in eastern North America.
Chemotaxonomy. Flavonoids were examined from 31 populations (cf. Ap-
pendix) of these four taxa. There was some variation in compounds among
the local populations of Vernonia texana but all four taxa have the same
flavonoid compounds. The flavonoid variation in V. texana seems to be in
part geographical and also due to hybridization with other sympatric taxa.
The taxa of this study have the greatest number of flavonoid compounds in
series Verae (Mabry et al., 1975). If reduction in compounds is an advanced
feature then the flavonoid evidence supports Gleason’s (1923) hypothesis
that the Texanae species-group is primitive in series Verae. Mabry et al.
(1975) report that all taxa have the sesquiterpene lactone glaucolide-A.
Experimental hybridizations. Crossing experiments of the Texanae species
group with other species in series Verae gave 80 to 90% mean pollen stain-
ability in F,, F,, and F, generations. Details of the 154 crosses attempted
in 105 combinations of taxa are reported in Chapman (1973) [Xerox copies
are available from G.C.C.]. Generally highly fertile (as indicated by pollen
stainability) first, second, and third generation and backcross hybrids were
obtained. Some of the progeny from the second and third generation hybrids
were weak and had abnormal development similar to that reported by
Faust (1972), but no significant hybrid sterility barriers were observed. One
of the more significant aspects of this study is that the taxa of series Verae
from the Sierra Madre Oriental of northeastern Mexico can be crossed with
other Vernonias from the eastern United States despite many years of geo-
cies isolation. Hybrids between the Mexican Texanae and members of
the xican series Umbelliformes produced hybrids with ca. 20-25% stained
soieae (Jones
Nioephiotorical analysis of local population samples. Local population sam-
ples of ten to twenty-six plants were collected from several locations
throughout the ranges of each taxon (see Fig. 1). Twelve morphological
characters (see Table 1) were measured on each plant of the population
samples and the values obtained were punched onto computer cards. The
mean, standard deviation, and range of each character for every population
was calculated, using a BMDO2R—Step-wise Regression Program from
the Health Sciences Computing Facility, UCLA (April 13, 1965 version).
A multivariant, univariant discriminant analysis of irregular data program

e
°
a JH
=|
fe}
°
MONTERREY 9&
a f}
*.
o “44
fo)
°
° t)
°
& VICTORIA
a
SAN LUIS
*% POTOS!
a,
\
a So
Oo

W MEXICO city

F I. Distribution 7 Rg ee texana (closed circle);
greggii (open circle),

p. steele (triangle), ssp.
square), and ssp. jeter (closed square).

V. greggii ssp.
schaffneri (open

268

(MUDAID—6400) written by Dr. R. E. Barghman of the University of Geor-
gia Statistics Department was used to compare these four taxa. Individual
populations of a taxon were combined into a single group for the computer
analysis. For example, the Vernonia ervendbergii group contained 75 indi-
vidual plants from 3 population samples; V. greggii, 100 (4 populations);
V. schaffneri, 97 (4 populations); and V. texana, 389 (18 populations). In
addition, the features of each taxon were compared, using dice-grams for
each of the twelve characters. Polygonal graphs showing eight characters
were constructed for each population and overlaid on or near their collec-
tion site on a base map. A hybrid population of V. texana was analyzed
with scatter diagrams.

It was assumed that much of the variation among the species-group and
certain correlations of characters might be better understood if only the
three Mexican taxa were analyzed together. Hence, another analysis was
made that excluded Vernonia texana using the same MUDAID—6400 pro-
gram as before. The MUDAID program included a canonical test of correla-
tion for each of the twelve morphological characters to each other and a
comparison of each character to the standard error.

The results shown in Table 2 indicate that the number of flowers/head
is the most useful character to distinguish the populational means of Ver-
nonia texana, V. ervendbergii, V. greggii, and V. schaffneri. The next best
characteristic is involucre length, followed in decreasing order by inner
bract length, involucre width, leaf width, leaf length/width, outer bract
length, inner bract width, outer bract width, leaf length, outer bract length/
width, and inner bract length/width. When V. texana is removed from the
statistical comparison, there is a shift in the order of significant characters.
For example, in the comparison without V. texana, the inner bract is the
most significant character and the number of flowers per head dropped to
the fourth best character.

The variation pattern in eight characters of 24 populations of Vernonia
texana from throughout most of its geographical range is shown by poly-

TABLE 1. Morphological eae Measured or Scored on the Local
Popul

ation Samp]
Characteristic Characteristic
(LELT) Leaf length, mm (IBL) Inner bract length, mm
(LEWI) Leaf width, mm (IBW) Inner bract width, mm
(LL/W) Leaf length/width (IL/W) Inner bract length/width
(INLT) Involucre length, mm (OBL) Outer bract length
(INWI) Involucre width, mm (OBW) Outer bract width

(FLS) Flowers per head (OL/W) Outer bract length/width

269

gonal graphs in Figure 2. Of the eight characters used in the polygons, leaf
width and leaf length/width appear to vary the most from population to
population. Leaves tend to be narrower in the southeastern part of the range
and wider toward the north and northwest. Much intra-populational variation
is revealed in the polygons, and no definite geographical correlation is appar-
ent.

The polygonal graphs constructed from sixteen population samples of the
three Mexican taxa, Vernonia ervendbergii, V. greggii, and V. schaffneri,
are compared in Figure 2. There appear to be no major differences in their
basic shape, but there is a considerable difference in the size of the graphs.
The eastern side of the Sierra Madre has the smallest polygons, the western
side has the largest, and to the south are found the intermediate polygons
showing a geographical correlation.

The dice-grams (Figure 3) demonstrate that the ranges of all twelve char-
acters overlap when composite populations of Vernonia ervendbergii, V.
greggii, V. schaffneri, and V. texana are compared.

The MUDAID computer program produced a table showing correlations
based on matrix E (correlation based on corrected > from ANOVA) among
the twelve characteristics used in the morphological studies. Table pre-
sents each character and the five characters most correlated with it for all
four taxa, Vernonia texana, V. ervendbergii, V. greggii, and V. schaffneri.
Those characters showing a value of .500 or higher are considered signifi-
cant and are underlined. The inner bract length shows a correlation between

TABLE 2. F vs. Standard Error for Characteristics Measured on Local
Populations of the Texanae Species—Group of Vernonia.

Mexican Taxa Mexican Taxa + V. Texana

Character* F Value Character* F Value
IBL 197.6 FLS 750.3
INWI 193.4 INLT 613.8
INLT 181.3 IBL 368.0
FLS 169.7 INWI 365.1
OBL 144.0 LEWI 305.4
LL/W 89.4 LL/W 192.6
LELT 67.6 OBL 155.9
OBW 60.9 IBW 103.6
IBW 27.6 OBW 70.6
LEWI 17.9 LELT 54.2
IL/W 17.5 OL/W 16.0
OL/W 13.0 IL/W 10.0

* Refer to Table 2 for explanation of abbreviation.

270

the involucre width, inner bract width, inner bract length/width, and outer
bract length; the inner bract width to the inner bract length/width and the
outer bract length to the outer bract width.

Natural hybridization between Vernonia texana and V. baldwinii is docu-
mented by scatter diagrams in Figure 4. The hybrid population sample
(15843) includes some of both parental types and many intermediate forms.
Putative backcrosses were also recognizable within the population. Artifi-
cially produced F, hybrids (crosses 10C, 11C) are morphologically inter-
mediate between ie two parental types, V. texana (15839) and V. baldwinii
(15841).

The morphological analysis of the local population samples indicates that
Vernonia ervendbergii, V. greggii, and V. schaffneri are related to V. tex-
ana: however, V. texana can be easily distinguished from the other three
taxa. The smaller number of flowers per head and the narrow-linear to
narrow-lanceolate leaves of V. texana are the most useful key characters
for separating V. tevana from the three Mexican taxa. It is not possible to
clearly separate the three Mexican taxa from each other with morphological
features.

The morphological characters evaluated for the local population samples
of Vernonia texana vary in their mean values as shown by the polygonal
graphs. This variation can be expected since the taxon ranges over a rather

TABLE 3. Correlation of the Five Most Correlated Characteristics Based
atrix E for rnonia ervendbergii, V. greggti, V. schaff-
neri, and V. texan

Character* Correlation

Higher Lower

LELT LEWI .388 INWLI .133 IBW .108 OBW .102 FLS .092
LEWI LL/W .492 LELT .388 OBW .320 OBL .276 INWI .232

LL/W LEWI .492 OBL .265 OBW .261 INWI .196 IBL .160
INLT IBL .371 OBL .299 FLS .278 OBW .201. LEWI .128
INWI IBL .509* FLS .384 OBL .350 LEWI .232 IL/W .212
FLS INWLI .384 INLT .278 OBL .264 IBL .241 OBW .220
IBL OBL .624 IL/W 539 INWI .509 NLT .371 OBW .313
IBW IL/W .779 OBW .312. OL/W .248 LEWI .186 FLS .172
IL/W IBW .779 IBL .539 FLS .396 OL/W .345 OBL .285
OBL IBL .624 OBW .557) OL/W .397 INWI .349 INLT .299
OBW OBL .557 OL/W 397 LEWI .320 IBL .313 IBW .312
OL/W OBL .397 OBW 397 IL/W_ .345 IBL .277 IBW .248

* Refer to Table 2 for explanation of abbreviation.
** Underlined characters show high correlation.

271

Fig. 2. Polygonal graphs of eight pene characteristics from local
Sele of Vernonia texana (in U.S.) a V. ervendbergii, V. greggii,
V. schaffneri (in Mexico). The Se inside polygon is a collection
ae (cf. Appendix). Axes represent A, number of flowers per head:
involucre length (mm); C, inner bract len ngth (mm); D, involucre width
mm); E, leaf width (cm); F, leaf length/width; G, outer bract length
(mm); H, inner bract width (mm).

272

A
8
Cc
Dd

—fe——

=. —h. pee eee

° 2 4
OUTER BRACT LENGTH mm

o
o

08 14
OUTER BRACT WIDTH mm

0
OUTER BRACT L/W

—_

ay

a

oO 5
INNER BRACT LENGTH mm

4
05 10
INNER BRACT WIDTH mm

10

rn . “

15 20 26

re a

INNER BRACT L/W

6 12

rf OOW>

INVOLUCRE LENGTH mm

t rr =)
0 90 wo
INVOLUCRE WIOTH mm
i
—E
—_lm

4 L A A 4

40 60 80

0 20
FLOWERS / HEAD

Ee

[_ eee - *
Y 9 18 2
LEAF LENGTH cm

————_—
—=
-—_
=——

Q
LEAF WIOTH om

>

=—

—
oO 4 i
LEAF LW ae $0

Fig. 3. Mean, range, and standard deviation for selected characters of
Vernonia ervendbergii (A), V. greggii ee mes i a (C), and V. texana
(D) from combined populations of each t

273

> LEAF WIDTH CM
= 7 , : te
bb b
bb e

OV3H /SH3MOT14
c

o@
bye foe
ee

| e 2 2
°
re)

beo 7 0 2

al uzro °

(oe) Uae

E56
Zo
> 2
a] =
oO
ee) LEAF WIDTH CM
m re = - >
es Oo $ & a ep anf)
mn
oO
=
5
NO
os oe) fe)
re)
x o “oS ?
> ro)
re} fe)
ay oOo fe} (eo)
a fe)
ra Oo
oO
a
4
2 re) o (~O ro)
(o)

ig. Scatier diagrams of Vernonia texana and V. baldwinii. A, typical
V. terana (19839), artificial F, V. texana X V. baldwinii hybrids (X10C,
X11C), and typical V. baldwinii (15841); B, local population sample (15843)
is a ate population which includes putative hybrids and both parental
types

274

large geographical area but the variation is due to natural hybridization
with other sympatric Vernonias.

The eight morphological characters of Vernonia ervendbergii, V. greggii,
and V. schaffneri compared in polygonal graphs (see Fig. 2) show some
geographical correlation. Although the taxa appear sympatric, the moun-
tainous region where these taxa occur provides diverse habitats. Two popu-
lations could be only a few airline miles apart, yet be isolated by a moun-
tain range, with one site being dry and the other wet. The isolation available
in the area would allow local populations to diverge, since they are prob-
ably not sharing the same gene pool with many adjacent populations.

The hybridization experiments demonstrated that hybridization can occur
in the greenhouse and some evidence of natural hybridization is suggested
by the polygonal graphs of populations 85 and 86. These two populations
are intermediate in size between those of V. ervendbergii (87) and V. greg-
git (83). The two intermediate populations (85, 86) were collected along a
road cut through the Santa Rosa Canyon connecting the ranges of V. ervend-
bergii and V. greggii. Scatter diagrams of V. greggii, V. ervendbergii, and
putative hybrid populations confirm that natural hybridization does occur
between the Mexican taxa (Fig. 5). Most of the individuals in the hybrid
population appeared to be F, hybrids rather than backcrosses to either
parent.

Vernonia schafjneri populations generally are intermediate between V.
ervendbergii and V. greggii and resemble closely the population (86) dis-
cussed above. This possibly could be the result of past hybridizations, with
the hybrids now occupying slightly different habitats than either V. ervend-
bergit or V. greggit.

Transplant studies and ecology. In the autumn of 1970 and the summer of
1971, rootstecks of Vernonia ervendbergii, V. greggii, and V. schaffneri were
collected from the field, potted, and grown in the greenhouse and transplant-
ed into the Boiany Plant Growth Facility Garden in Athens in the spring of
1971 and 1972. Rootstocks of V. texana were collected in the summer of 1971
and transplanted to the garden in the spring of 1972.

Transplants of Vernonia ervendbergii, V. greggii, and V. schajjneri from
Mexico, and V. texana from the south central United States became estab-
lished and were successfully over-wintered for four years at Athens until
termination of the experiment. Vegetative shoots began emerging from the
underground rhizomes during late April and early May. Floral development
in the three Mexican taxa began in late May with some anthesis by the first
week in June and by mid-June they were at the peak of flowering. The
flowering pericd normally lasted for 3-4 weeks. Vernonia texana began
erowing vegetatively about the same time as the Mexican taxa but did
not reach peak flowering until the middle of August. The garden populations
of V. terana showed a variation of 7-10 days in the time of anthesis, with
the populations from Arkansas flowering before those from southeast Texas.

275

10.
= 8.
=
= e e
2
e® © e
a 6s .
= °.
2 © $d, °
- e
eq 44
Ww ) e e 86
Z
Zz
20 30 40 50 60 70
FLOWERS / HEAD
109
go §
= 8: © ome) ° oO
= re) re) 0° oOo
x fe) re) O50
o . 2
a e
ar 6+ e e e e
- e
5 AR a
< a e
a AAA
oO A ERVENDBERGI|
>, A AA
4 @ HYBRID
x rN A
4 o GREGGII
Zz
20 30 40 50 60 70
FLOWERS /HEAD

Fig. 5. Scatter diagrams of hybridization betwcen Vernonia greggii and
is

V. ervendbergii. Local population sample

86 (top) is a eal els ceorate

which includes primarily hybrid type plants. lower diag compares

typical V. greggii (83), artificial F, V. greg
7).

(X35C), and typical V. ervendbergii (8

7X VY. ce hybrids

276

The Mexican taxa are found primarily in the open oak-pine vegetative
zone at ca. 4500 ft to 7500 ft in the Sierra Madre Oriental on limestone par-
ent material. They also invade semi-disturbed habitats such as roadsides
or pastures and sometimes form large local populations. The soils from
habitats of Vernonia ervendbergii and V. greggii are of similar pH and type.
There are many exposed limestone boulders in these areas, and the soil is
very dark colored with a pH of 7-8. Vernonia ervendbergi occupies the
moist, slightly alkaline soils in the open oak-pine zone of the eastern slopes
of the Sierra Madre Oriental. Vernonia greggii is found on the western side
of the Sierras in slightly alkaline soil of the open pine zone where, due to
the rain shadow, there is less moisture. Vernonia schaffneri, found in the
extreme southern part of the species-group’s range in Mexico, inhabitats
severely eroded, over-grazed pastures. Much more precipitation falls in this
area, as evidenced by the lush vegetation and severely eroded pastures.
Some parental limestone rocks are exposed, but the soil is not as well-
drained as that of V. ervendbergii or V. greggii. This soil is red with a pH
of 4-5.

Vernonia texana is associated with the pine-oak communities of east Texas
to Mississippi, north to the mixed hardwocd forest of central Arkansas.
These habitats are typically dry, on sandy soil with little organic matter and
a pH of approximately 5.5. Plants of V. texana are widely scattered in open-
ly grazed areas, woodlands, and along road and railroad right-of-ways.

SYSTEMATICS AND TAXONOMIC TREATMENT

Vernonia texana is uniform throughout its range in chromosome number
and leaf anatomy. It is variable in a few morphological characters (leaf
width and leaf length/width); however, no sharp discontinuities were de-
tected in the morphological variation. Interspecific hybridization also causes
some local morphological and flavonoid variation. The transplant studies
indicated a slight variation in the phenology of local populations on a north-
south gradient. Although variation was noted within V. texana, no distinct
subpopulations that merit naming were detected.

The three Mexican taxa, Vernonia ah eal V. greggti, and V. schajf-
neri, are more closely related to each other o V. texana. Cytologically
and phytochemically, the three Mexican taxa are identical as all have the
sesquiterpene lactone glaucolide-A and the same flavonoid compounds
(Mabry, et al., 1975). Also, the leaf venation, trichome complement, and
internal leaf anatomy are similar; nevertheless, slight variation in the leaf
epidermal pattern exists among the taxa. The vegetation e wh a
three taxa grow in the Sierra Madre Oriental is similar (Leopold, 1950),
although their micro-habitats differ. With the exception of one character,
the number of flowers per head, the three taxa are not easily distinguished
from one another. The values obtained for almost all morphological char-

6400 (discriminant analysis) program did not provide any combination of

277

characters that clearly separated one taxon from another. There was, how-

ever, a rather distinct geographical correlation with the morphological vari-

ation,

The four taxa in the Texanae species-group are closely related. The phyto-
chemical data, both flavonoid and sesquiterpene, was identical. There were
no differences in the chromosome number or basic leaf anatomy since the
same kinds of trichomes, leaf venation, and leaf epidermal patterns were
found in all members of the group. A discriminant analysis of morphological
characters from local population samples throughout the range of each
taxon suggested that Vernonia texana is separable from the three Mexican
taxa. In contrast, the three Mexican taxa were not distinctly separable from
each other. Artificial hybridization experiments indicated that no gross
chromosomal differences existed among the four taxa. Fertility was high
among the F,s, and no hybrid breakdown was found within the group. Al-
though the habitats and geographical ranges are diverse, all taxa were able
to survive for four years in a uniform garden in the Piedmont of the South-
eastern United States. However, as Miranda and Sharp (1950) first suggest-
ed, the climates of the uplands of the southeastern United States and of the
Sierra Madre Oriental of northeastern Mexico are surprisingly similar.

The four taxa were probably derived from a common ancestral line. Ver-
nontia texana should be maintained as a species and the three Mexican taxa
treated as subspecies. Since V. ervendbergii, V. greggi, and V. schaffneri
were described by Asa Gray (1882) in the same publication, the choice of
which name to apply is entirely a matter of preference for the author (Sta-
fleu, 1972) and V. greggii was selected.

During the course of this study a herbarium specimen from Mexico was
discovered which could not be assigned to the Mexican subspecies, The
specimen’s collection location (Muzquiz, Coahuila) is most interesting, since
this site is directly along the likely migration route that Vernonia used to
reach the United States (Jones, 1976). We feel that this most likely repre-
sents a different population and should, therefore, be treated as a subspecies
of V. greggii. Ideally, a systematist should have more material on which to
describe a subspecies; however, at this time this is impossible. A trip to
Muzquiz was made in August 1973 to search for additional plant material of
this taxon but the trip was unsuccessful due to the remoteness of the area
and mechanical troubles with the vehicle.

1. Heads 11-35 flowered (usually ca. 20); middle cauline leaves 0.2-3.4 cm
wide, stiff; involucres 4-9 mm high; pinelands, E Texas, SE Okla., Ark.,
La., S iss. foe eee ee we ee Ud OV. étexana.

. Heads 15-75 flowered (usually over 27); middle cauline leaves 0.7-5 em
nee flexible; involucres 5-15 mm high; oak-pine zone, Sierra Madre

—

Oriental 2. V. greggit.
1, VERNONIA TEXANA (Gray) Small, FL. S. E. U. S. 1160. 1903.
Vernonia angustifolia Michx. var. texana Gray, Syn. Fl. N. Am. 1: 91.

1884. TYPE: Louisiana, Dr. Hale s.n. (HOLOTYPE: NY!)
ee angustifolia T. & G. Fl. N. Am. 2: 59. 1841. TYPE: Louisiana,
. Hale s.n. (HOLOTYPE: NY!)

278

Vernonia granienifolia [sic] Walt. var. texana So Trel. ex Branner
Coville, Rep. Geol. Surv. Ark. for 1888. 189.
Cacalia angustifol ia (Michx.) Kuntze var. a Gray, Rev. Gen. Pl.
891.

Perennial herbs; stems erect, puberulent to glabrate, 4-10 dm high; leaves
scattered, cauline; blades of middle stem leaves 0.2-1.4 ecm wide, 6.5-13.5
cm long, linear to linear-lanceolate or sometimes lanceolate, scabrous above,
scabrous to puberulent beneath, apically acute, basally attentuate, margins
revolute, remotely serrate, or sometimes prominently toothed; petioles 1-4
mm long, puberulent; inflorescence loose and open; heads 15-25 flowered;
involucre broadly campanulate, 4-6 mm high, 5-7 mm wide; phyllaries im-
bricate, slightly appressed, greenish-purple, ciliate; inner phyllaries oblong
to oblong-lanceolate, 2.9-6 mm long, 1-1.7 mm wide with acute to short-
acuminate tips ca. 0.1-0.2 mm long; pappus brownish to straw colored, often
tinged with purple, inner bristles 5.5-7 mm long; outer scales ca. 0.6 mm
long; corollas 9-11 mm long; achenes 1.5-3 mm long; flowering from late
July to August; chromosome number n = 17.

This species is distributed from Arkansas and SE Oklahoma south to
E Texas and SW Mississippi (Fig. 1).

Vernonia texana grows in well-drained soil of pinewoods and along road-
sides, This species is known to hybridize in nature with V. baldwinii, V.
gigantea and V. missurica.

2. VERNONIA stu eres Gray, Proc. Am. Acad. 17: 204. 1882. Typification
listed under subspec

Perennial herbs; pi oe puberulent to glabrate, 2-10 dm high; leaves
scattered, cauline; blades of midd!e stem leaf 0.7-5 cm wide, 3.3-12.5 cm
long, lanceolate to linear-lanceolate, elliptic or ovate, scabrous above,
brous to puberulent beneath, apically acute to acuminate, or obtuse, ie
narrowed to nearly sessile, margins serrulate to remotely serrulate; petioles
1-3 mm, nearly sessile, puberulent; inflorescence loose and open, sometimes
flattened: heads 14-75 flowered; involucre campanulate to hemispheric,
5-15 mm high, 418 mm wide; phyliaries imbricate and appressed, with
acute to obtuse or long acuminate tips, glabrous to thinly ciliate; achenes
3.5-10 mm long; flowering from early June to late July; chromosome num-
ber n = 17.

This species is distributed throughout the Sierra Madre Oriental of Mex-
ico (Fig. 1). The subspecies are characterized by the following key:

1. Heads ca. 75 flowered; outer phyllaries long acuminate.

2d. ag faustiana.

1. Heads ar than 70 flowered; outer phyllaries not long acuminate.

2. Mid stem leaves lanceolate to linear-lanceolate, heads Aare over

40 eee: western slopes of Sierra Madre Oriental.
2a p. greggy
2. Middle stem raves elliptic - ovate, heads usually ‘less fer 40
ered; eastern slope of the Sierra Madre Oriental.
3. Heads 27 (14 36) Howexed, phylaries ae with purple tips, pap-
pus whitish to straw colored. 2b. ssp. ervendbergii.

279

a 36 (21-50) flowered, phyllaries ae pappus brownish-tawny
colo 2c. ssp. schaffneri.
2a. VERNONIA GREGGII Gray ssp. GREG GIL
T > Mexico: Nuevo Leon, valley near Saltillo, 1848, Gregg 102
(HOLOTYPE: MO!)
Vernonia greggii Gray var. palmeri Gray, Proc. Am. Acad. 17: 204.
1882. TYPE: Mexico: Nuevo Leon, 1880, Palmer 753 (HOLOTYPE:
Y!
ne taylorae Standl., Field Museum Pub, Botan. 17: 224.
19 7. ge Mexico: Hacienda Pabillo, Galeana, Nuevo Leon, 1963,
Taylor 9 (HOLOTYPE: F!, ISOTYPE:

Vernonia greggii ssp. greggii is distributed ak the drier western side
of the Sierra Madre Oriental from about Dr. Arroyo, Nuevo Leon, to the
mountains immediately west of Monclova, Coahuila. This subspecies grows
in rocky limestone soil, along roadsides in full sun, but is often found in
semi-shade under sparse oak-pine-juniper forest. Sometimes robust plants
are found along wet ditches or dry stream sides. This taxon is somewhat
variable throughout its geographical range. The range of this taxon over-
laps with the range of V. greggii ssp. ervendbergti in the Santa Rosa Can-
yon east of Galeana, Nuevo Leon, and hybridization has been documented
from populations in this area.
2b. VERNONIA GREGGIL Gray ssp. ervendvergii (Gray) Chapman and

Jones stat. no
Vernonia ee a, Proc. Am. Acad. 17: 203. 1882,
exi W Monclova, Coah., 1880, Palmer 750 (SYN-
GH! NY!)

Vernonia greggii ssp. ervendbergii grows in the oak-pine vegetation along
the eastern escarpments of the Sierra Madre Oriental. Its range extends
from about Monterrey, Nuevo Leon, southward to immediately north of
Xilitla, S.L.P. It is also found to the east on the outlying Sierra de Tamau-
plipas in the Coastal Plain of Mexico. This subspecies invades semi-disturbed
areas such as roadsides and cut-over forests. The soil in which it grows is
black, moist, and slightly alkaline.
2c. VERNONIA GREGGII Gray ssp. sehaffneri (Gray) Chapman and Jones

stat. nov.

Vernoma schaffneri Gray, Proc. Am. Acad. 17: 204, 1882.
TY exico: near man Luis Potosi, $.L.P., 1877, Schaffner 347 (SYN-
TYPES: NY!, GH!,

Vernonia greggii ssp. oe has the most restricted range of the three
subspecies. This taxon is found only in the vicinity of Jacala, Hidalgo, SW
of Xilitla, S.L.P., Hidalgo in the state of Queretaro, and the vicinity of Las
Rusias east of San Luis Potosi, $.L.P. This subspecies seems to be associ-
ated with grazed-over pastures that are eroded into gullies. The soil is
acidic, red, and gummy.

Although natural hybridization between V. greggii ssp. schaffneri and ssp.
ervendbergii was not documented in this study, it is speculated that there
may be gene flow via the mountains W of Xilitla, S.L.P. to San Luis Potosi,
S.L.P. Unfortunately no roads traverse this area.

280
2d. VERNONIA GREGGII Gray ssp. faustiana Chapman and Jones ssp.

TYPE: Mexico: Canyon in Sierra Hermosa de Santa Rosa, 25 mi NW
Melchor Muzquiz, Coahuila, 1963, Latorre 13 (HOLOTYPE: TEX!)
Herbe perennes erectae, 0.3-0.6 metralis, caulibus tomentosis; folia 9-13
cm longa, 1.5-2.5 cm lata, linearis-lanceolatus; scaberalus supra et infra,
apicibus acutis, basibus cuneatis, marginibus dentatis, petioli 2-4 mm longi;
inflorescentiae dense compactis; capitula ca. 75 florata; involucra cam-
paniformis-hemisphericae, 9-10 mm longa, 11-15 mm lata; phyllaria im-
bricata, appresus, stramineo-purpurea; phyllaris interioribus linearis, 8—9
mm longis, 1.0-1.3 mm latis; phyllaris externis acumnatis longis, apicem-
purpurea, 1-2 mm longis; pappi stramineus; setae interiores 7-8 mm longis;
setae externae ca. 1.0 mm longis: corolla purpurea, 10-11 mm _ longis;
achaenia fere glabris, 4-8 mm longis.
This subspecies is named in honor of Dr. W. Z. Faust, who was properly
introduced to Mexico during the unsuccessful search for additional material
of this taxon.

APPENDIX
COLLECTION LOCATIONS FOR SAMPLES OF VERNONIA

Voucher specimens (deposited in GA) of materials used in the study of
flavonoids (ann pated f), cytology (c), hybridization (h), and morphology
(m). Collection numbers are those of the senior author unless otherwise
noted.

Vernonia greggii ssp. greggit
MEXICO: NUEVO LEON: 2.1 miS oe 51 _ f, he nN mi N Dr. Arroyo,
80 (c,f,h,m); 30 mi N Dr. Arroyo, 81 (c,m); : i N Dr. Arroyo at
LaJoya, 82 (c,f,m); 18 mi S Galeana, A Ne: ne a mi E Galeana, 85 (m);
14 mi E Galeana, 86 (m).

Vernonia greggii ssp. ervendbergit
MEXICO: NUEVO LEON: Horsetail Falls, 45 (c,f,h); Horsetail Falls,
87 ! m).—QUERETARO: 15 mi S Jalpan, 71 (c,m). Km 226, hwy. 120,
15 Landade Matamoros, 72 (m).—SAN LUIS POTOSI: 20 mi W Valles,
75 (c,m); Km 71, 42 mi W Valles, 76 (c,m).—TAMAULIPAS: Km 153, hwy.
101, Victoria, 46 (c,f,h): km 153, hwy. 101, Victoria, 60 (m).

Vernonia greggii ssp. schaffneri
MEXICO: HIDALGO: 5 mi S Jacala, 49 SE Pe 111, hwy. 85, Puerto
La Zorra, 66 (c,f,m); 10 mi S. age la, 68 (c,m); is Jacala, 69 (m).—
SAN LUIS POTOSI: Km 233, hw - 120, 20 mi S She 73 (c,m); ca. 31 mi
E San Luis Potosi, at Las eee 79 (c).

Vernonia texana

UNITED STATES: ARKANSAS: ASHLEY CO.: 10 mi W Parkdale, 91
(c,f,m); 20 mi W Parkdale, 92 (f,m). CLEVELAND CO.: 8 mi N New Edin-
burg, 94 (m). DALLAS CO.: 1 mi S Princeton, 96 (f,m). HOT SPRING CO.:
near DeRoche, 97 (f,m); 3 mi E Bismarck, 98 (f,m); ca. 9 mi NW Malvern,
Jones 15839 (c,f,hk,m); ca. 9 mi MW Malvern, Jones 15840 (f); ca. 9 mi NW
Malvern, Jones 15842 (f,m); ca. 9 mi NW Malvern, Jones 15843 (f,m). POLK
CO.: 5 mi N Grannis, 99 (f,m); 2 mi N Grannis 100 (f,m).—LOUISIANA:
VERNON Pa.: 3 mi N Slagle, 56 (c); 5 mi N Leesville, Jones 17658 (f,h,m).

281

—MISSISSIPPI: AMITE CO.: 1.3 mi E Lincoln—Amite Co. line, — 2293
(m). FRANKLIN CO.: Clear Springs Lake, Ba 2292 (c,h iy ae KE CoO.:
mi Magnolia, Jones 19645 (c,h,m,f)—TEXAS: CHERO Sane 2 mi
N Jacksonville, 104 (f,m); 8 mi N ae 105 (c,m). Ngee CO.:
7 mi W Columbus, 90 (c,f). GOLIAD CO.: 10 mi SW Victoria, 59 oo
JASPER CO.: 6 mi NE Burkeville, Jones: 17659 a LEON CO.: 1m
N Buffalo, 106 (f, os MILAN CO.: 13 mi N Rockdale, 107 (c,f,m). RUSK
CO.: 4 mi NW Tatum, 102 (f,m); 12 o SW Tatum, 103 (f,m). TYLER CoO.:
5 mi W of Woodville, Vones 17661 (f,m

REFERENCES
eee ~ C. & F. V. COVILLE. 1891. A list of the plants of Arkansas. Ark. Geol.
Sur

Gren. a C. 1973, Biosystematics of the Texanae Species-Group of Vernonia (Com-
positae). Ph.D. Dissertation, niv. of Georgia, Athens
FAUST, W. Z. 1972. A biosystematic study of the Taceriows species group of the genus
Vernonia Saueween Brittonia 24: 363—378.
A.

GLEASON, H. 906. A revision of the North American Vernoniecac. Bull. N. Y. Bot.
Gard. 4: 203—-2
——. 1922. Vernonieae. North American Flor

23. amen and. Boost shical are rene of oe genus Vernonia in North
Agnetics. Amer. J. Bot I
GRAY, A. 1844, Synoptic floss ae North America. 1: 91. New York,
_ Novitae Arizonicae. Proc. Am. Acad. 17: 203—204.
HUNTE R, G. EB nee Che omosome numbers in ee section Lepidoploa: subsection
Panic sileste Verac. S. Naturalist 9: 239—24
JONES, S. B. 1970, Ea numbers in Danes Bull. Torrey Bot. Club 97:
nua’
72. A systematic study of the fasciculatae group of Vernonia (Compositac).
Bretonts oe 28—45.
i . 1976. Cytogenetics and affinities of ees See a) from the Mexican
Eiehlande eastern North America. Evolution. 30: 462
KING, B. L. S. B. JONES. 1975. The Vernonia ey ie complex (Compositae).
Brittonia 27: ae
KUNTZE, O. 1891. Pes — Plantarum. 2:
LEOPOLD, A. S. 1950. Vegetation zones of Mexico. Boolory 31:
MABRY, T. J., Z. ABDEL- BASET, W. G. PADOLINA and S. B. ey on Systematic

sae of flavonoids and sesquiterpene lactones in species of Vernonia. Biochem.
Syst. Eco. 2: 18

MIRANDA, F. & A. J. ARP. cons Characteristics A the vegetation in certain tem-
perate regions 7 eastern avers co. Ecology 31: 313—33

SHARP, A. J. 1951. The relation - re Eocene Wilcox rea to some modern floras. Evo-
lution 5: 1—5,

ee 7 1903. Flora of the southeastern United States. New York: the author.

i+ io pp. 1160—1161).

STAFLEU, _ a a 972. International code of botanical ecarmen a Veg. 82.

STANDLEY, P. C. Pees Studies of American en Field N n Pub. 17: 224.

TO ee J. & A. GRAY. 1841. Flora of North America. pe New me Wiley and

Putnan
TRE PASE, W. as In: J. C. Branner & F. V. Coville. A list of the plants of Arkansas.
Ark. a Sur. 4: 189.
eyo L. E. on stematic study of the ra and Giganteae species-group
of the genus Vernonia aie itae). Brittonia 24: 229—238

MISSISSIPPI FLORA: A GUIDE TO THE
ERNS AND FERN ALLIES

A. MURRAY EVANS!
Department of Botany, The University of Tennessee, Knoxville, TN. 37916

ABSTRACT
Keys, distribution maps, habitats, nomenclature, and notes are given for
the ferns and fern allies known to occur naturally or naturalized in Missis-
sippi.

INTRODUCTION

This paper represents my contribution to the Flora of Mississippi Project,
an effort being undertaken primarily by Drs. Sam Jones, (University of
Georgia), Tom Pullen, (University of Mississippi), and Ray Watson (Mis-
sissippi State University). Hopefully, this information will stimulate efforts
to improve knowledge of the ferns and fern allies of the state, so that any
additions or corrections can be incorporated before the proposed compre-
hensive guide to the flora of Mississippi is published. Any overlooked spe-
cies, additional collections, needed modifications, or suggestions are wel-
comed so that appropriate revisions can b

The primary sources of information for this treatment are the herbarium
collections of the University of Mississippi, Mississippi State University,
and the University of Southern Mississippi, and only those taxa supported
by voucher materials or unquestionably substantiated in the literature are
included. References made to some of the standard manuals of the south-
eastern flora are abbreviated as follows: Small, 19388 (S); Fernald, 1950
(F); Gleason and Cronquist, 1963 (G); Evans, in Radford et al., 1968 (R).
Other taxonomic works used in preparation of this treatment are cited in
the text.

The physiographic regions of Mississippi, based on those of Lowe (1921)
are outlined in Fig. 1. The abbreviations used in the text are: Tennessee
River Hills (TRH); Northeastern Pine Belt (NPB); Pontotoc Ridge (PR);

Loess Bluff Hills (LBH); Yazoo-Mississippi Delta (YMD); Longleaf Pine
Region (LPR); and Coastal Pine Meadows (CPM).

PTERIDOPHYTA
1. Plant free-floating on water or resting on mud; spores of 2 sizes, mega-
spores and microspores . . 13. Azollaceae.
1. Plants rooted to soil or mud: s pores of « on tw
2. Plant nee jointed; ne leaves feed Chee ‘at least basally

‘Contr. Bot. Lab., Univ. Tenn., N.S. 497.

SIDA-7 ii) = Geeo9 7 res.

283

into sheaths; sporangia borne below peltate plates in terminal cones.
1E

quisetaceae.

2. Plant not jointed; the leaves all separate and not forming sheaths;
sporangia borne variously but not on peltate plates.

3. Sporangia in axils of scale- ue or buried in basal pockets of soft

pene rush-like leaves; leaves with only a single vein or midrib,

4.

4,

"Plant ‘tufted, rush-like, the stem forming a lobed corm; leaves 5—50
m long; spores of two sizes soetaceae,
Plant coarse moss; the stems elongate and ‘branching: leaves

0.1-1.5 ¢
5. Plant foe “usually 8-40 cm tall; the shoots usually 5-20 mm
broad; spores of one size . 2. Lycopodiaceae.
d. Plant slender, usually low, 1 rarely more than 5 em tall; shoots
ally 2-6 mm broad; spores of 2 sizes . . 3. Sela ginellaceae.

usu

3. Sporangia on long, simple or branching fertile spikes”? arising from
the leaf, or borne upon surfaces of normal or contracted leaf blades;
leaves simple or divided, with lateral veins arising from the midribs.

or)

. Sporangia 0.5-1 mm broad, exposed on panicle-like cluster or sun-

ken in spike-like Lenn from the blade base or petiole base
but not on separate modified non-green leaf. 5. Ophioglossaceae,
b

. Sporangia 0.1-0.4 m road, borne directly upon leaf blade or on

non-green skeletonized leaf or aes apex (Osmundaceae), or inrolled
bead-like leaflets ca. 2— 7 mm in diam. of specialized non-green leaf
in Onocl (Aspleniaceac).
7. Leaves specialized ints “‘vine-like’’ organs which climb upon other
vegetation; sporangial annulus apica 7. Schizaeaceae.
7. Leaves typically fern-like, not ‘‘vine- like” sporangial annulus
lateral, or as a more or less vertical bow.
8. Bases of leaves with stipules or thin lateral wings; sporangia
with only a lateral patch of thickened annular ce ot
ee undaceae.
8. Bases of leaves lacking stipules: sporangia with a eat defined
bow-shaped annulus.

9. Leaf blade delicate, filmy, only 1 cell thick ewe the veins;
sporangia in outward-facing cup-like or ee involucres
at the leaf margin. menophyllaceae.

. Leaf blade thick, at least 2 or more cells hick between veins;
sporangia dorsal or marginal, but not in an outward- facing
marginal cup
10. Sori marginal, covered by revolute marginal leaf tissue.

Rhizome and leaf bases clothed with hairs only; sorus

continuous along leaf margin, petioles green or stramine-
ous, coarse and tough. ennstaedtiaceae.

. Rhizome and leaf bases clothed with re rus con-
tinuous along leaf margin, or, if round, the petiole ebony
colored and brittle. 10. Adiantaceae.

10. Sori on the underside of the leaf blade : ‘if marginal the in-

dusia kidney-shaped and not continuous with the leaf mar-

co

a
_

—
_

gin
12. Sorus round, or linear along the veins; indusium present
r, if lacking, the leaf pinnate or more dissected.
11. Aspleniaceae.
12. Sorus round; indusium Jacking; ‘leaf ae ate
. Polypodiaceae.

284

1. EQUISETACEAE
EQUISETUM L. Scouring-Rush
1. E. hyemale var. affine (Engelm.) A. A. Eat. May-Aug. Railroads, road-
sides, stream banks; LBH, NCP, YMD. E. praealtuwm Raf.—S. Fig. 1

2. LYCOPODIACEAE
LYCOPODIUM L. Clubmoss

1. Upright shoot diffusely branched; strobili nodding. 1. L. cernuum.

1. Upright shoot unbranched or few-branched, terminating a etew erect
strobili.

2. Leaves of the ae oe shoot sharply differentiated between pe-
duncle and strobilus; leaves of the peduncle not ese eines horizontal
stems strongly Ao ee L. carolinianum,

2. Leaves of the fertile upright shoots with similar sporopiyl and vege-
tative leaves; leaves of the peduncle densely imbricate; horizontal
stems only slightly dorsiventral. (Hybrids with mieeneainte morphology
are typically ee between the following three taxa when in
mixed populat
3. Upright ee (including leaves) 4 mm or less in diam.; leaves of

the peduncle and strobilus appressed and entire, or rarely with a few
cilia at the leaf base; horizontal stems somewhat dorsiventral and

rooting to the ground throughout. 3. L. appressum.
3. Upright shoots ee leaves) 6 mm or mor e in diam.; leaves of
oS uncle ascending a incurved and those of re strobilus spread-

ng, leaves ciliate A cachoue horizontal stems not, or only slightly,
dorsiventral either flat and rooting continuously or arching and root-
ing aay ittentl
or

ntly.
4. ontal stems rooting continuously along the ground, the rhizome
ie cere or less i ., the leaves usually spreading
almost at right angles to the rhizome; gro the current year

t .
usually with only one (occasionally 2) erect fertile oe and usu-
ally without additional stiffly erect but sterile shoo

_ prostratum.
4. Horizontal stems strongly arching ‘and rooting ‘only intermittently,
the rhizome 3 mm or more in diam., the leaves usually ascending

and/or incurved; growth of the current year usually with 2 or more

(occasionally only one) erect fertile shoots, and usually with 1-

several additional stiffly erect sterile shoots. . 5. L. alopecuroides.

.L. cernuum L., — ing C. All year. Wet loamy roadbanks; George
aca Jackson Cos. Fig

.L. carolinianum . Slender C. Jul-Nov. Wet bogs, sandy meadows,
pinelands, and faagaiee. CPM, LPR, and NCP. Fig. 1.

3. L. appressum (Chapm.) Lloyd & Underw., Southern Bog C. Jul-Nov.
Bogs, wet meadows, pinelands, roadsides, and lake shores; CPM, LPR,
NCP, and TRH. L. inundatum var. bigelovii Tuckerm.—F, G. Hybridizes fre-
quently with nos. 4 and 5.

4. L. prostratum Harper, Prostrate Foxtail C. Jul-Nov. Bogs, wet road-
sides, lake shores, savannahs and pinelands; CPM, LPR, and NCP. L. alope-
curoides L.—F, G, R, in part. Hybridizes frequently with nos. 3 and 5. Fig.
1.

285

a :
a do

\

PHYSIOGRAPHIC REGIONS

i a ee

cena
5s | we ra
apn oe ele ad
¢ ¥ cn a Le oe
ec ee ae

a 7 | Lo
yo. | @8e@
& : @

LYCOPODIUM ALOPECUROIDES

carolinianum,
donee S. @

fy

ti Se

wise ie

< < , 7 nd

f _@@
(@@
eo ~

LYCOPODIUM APPRESSUM

At |

ees ro AY

SELAGINELLA APODA

SELAGINELLA LUDOVICIANA

var.

Fig

in Mississi i of ae um hyemale
appressum, L. prostratum

da,

ffi
L. alopecuroides

el. Physiographic regions of Mississippi (see text) and distribution
affine, Lycopodium cernuum,
Selaginella

5. L. alopecuroides L., Foxtail C. Jul-Nov. Bogs, wet roadsides, lake
shores, savannahs, and pinelands; CPM, JP, LPR, and NCP. Hybridizes
frequently with nos. 3 and 4. Fig. 1

Species nos. 3, 4, and 5 represent a confusing complex of taxa which are
presently under investigation. Intermediates which we interpret to be hy-
brids are encountered when two or more grow in proximity; pure stands
appear uniformly to conform to the species standards outlined here (with
J. C. Odenwelder, pers. com.).

3. SELAGINELLACEAE
SELAGINELLA Beauv. Spikemoss

1. Leaf blades with wide hyaline margins with 2-4 rows of oo hyaline

cells; strobilus 10-20 mm long; plant mostly ascending. ludoviciana,.
1. Leaf blades with narrow hyaline margins with 0-1 row of a. fle

cells; strobilus 6-15 mm long; plant ae creeping. 2. S. apoda

1. S. ludoviciana A. Br., Louisiana S, Jun-Oct. Moist deciduous and pine
wocds; CPM and LPR. Diplostachyum aes (A. Br.) Small—s.

2. S. apoda (L.) Spring., Meadow. S. Jun-Oct. Wet roadsides, deciduous

wocds, bogs, and stream banks; CPM, LPR, NCP, and TRH. Diplostachyum
sa (L.) Beauv.—S; S. apoda (L.) Fern.—F. Fig

This treatment follows that of Somers and Buck (1975),

4. ISOETACEAE
ISOETES L. Quillwort
1. I. engelmannii A. Br., Engelmann’s Q. Apr-Nov. Small pond margin;
Tishomingo Co. First collected for the state in 1975. Fig. 6.

5. OPHIOGLOSSACEAE

. Sterile segment of leaf dissected; fertile spike branched. 1. Botrychium.
1. Sterile segment of leaf simple; fertile spike simple. . 2. Ophioglossum.

_

1. BOTRYCHIUM Sw. Grapefern

1. Leaves herbaceous, or coriaceo leaves and spores maturing in late
winter or spring; fertile nen Beene from the petiole 0.5 cm or
less below sterile pi e.
2. Leaves periec cae. 8-80 cm high, sterile blade raised well above the
ground. virginianum.
2. oe coriaceous, 6-14 « ecm high, sterile blade lying close to the ground.
2. = lunarioides.
1. Leaves coriaceous, evergreen or appearing “only. in abecs r or fall;
spores maturing in late summer ae fertile eae joining petiole
more than 0.5 em below sterile pi
3. Ultimate segments with meee veins ae ite or absen

leaves often two or more per season. . B. ala Roe
3. Ultimate Ogee with conspicuous central veins; ‘leaves normally
one per

4. Blades poe tripinnate, the outline of the small lateral lobes of

287

pinnae, or pinnules of more dissected pinnae, rhomboidal and angu-
lar, or whole sterile portion highly dissected. . B. dissectum.
the outline of the small lateral lobes of
innae, or pinnules of the more dissected pinnae, oblong and some-

what rounded. 4. B. biternatum.

1. B. virginianum ‘5. Sw., ’ Rattlesnake- fern. "Mar. June, Rich deciduous
woods, swamps, ravines, and upland woods; throughout except CPM. Fig. 2.

2. B. lunarioides Sw., Prostrate G. Mar-May. Disturbed grassy areas,
cemeteries; Clark, Cacece. Greene, Jones, Lauderdale, Wayne Cos. B. biter-
natum sensu Small, not Underwood—S. Fig.

. B. alabamense Maxon, Alabama G. Sep- Oct. Old fields, rare; George
Co. Fig. 2.

4. B. dissectum Spreng., Common G. Sep.-Oct. Deciduous woods: uncom-
mon and usually confused with the following species; Lafayette, Marshall
Cos. includes B. obliquum Muhl.—S in part; B. dissectum var. obliquum
(Muhl.) Clute—G. Fig

do. B. biternatum Gav) Underw., Southern G. Aug-Oct. Moist pine and de-
ciduous woods, and ravines; scatiened throughout except CPM and YMD.
B. dissectum var. tenuifolium (Underw.) Farw.—F, G; B. obliquum Muhl.—
S, in part; not B. biternatum sensu Small—S. Fig. 2

—
mes!
_—
wo.
Qu
@
n
So
n
i
7
=e
so
—e
=)
SD
job)
co
@

2. OPHIOGLOSSUM L. ADDER’S-TONGUE

1. Leaf blades Hoe i 3.0 cm long, 2 or more per plant; rhizome short
eelindaie to glo
2. Rhizome BiGRece: epulbose: 4-15 mm in diam.; leaf blades 1-3.5 cm long.
O. crotalophoroides.
2. Rhizome short-cylindric, 2-3 mm in diam.: leaf blades 0.5-1.5 cm long.
udicaule.
1. Leaf blades more than 3.5 cm long, mostly 1 per plant: ‘rhizome cylindric.
3. Main veins forming large primary areoles with ets smaller veins
forming secondary areoles; leaf apex acute to apic
engelmanni.
3. Main veins eee areoles not enclosing finer secondary areoles; leaf
apex rounded to a
4. Plants clonal with horizontal stolons producing numerous new plan
usually with 4-8 parallel veins passing through base of sterile Rede.

producing a slender basal midvein. . petiolatum.
4. Plants not clonal, without proliferous ‘stolons: usually with 8-20 veins

passing through blade base, producing a heavy basal — ein
D. vulgatum.

1. O. crotalophoroides Walt., Bulbous A. Mar- Aug. Sandy lawns and ceme-
teries; LPR, NCP. Frequent but little collected. Fig. 2

2. O. nudicaule L. f., Least A. Mar-Aug. Lawns, pastures and cemeteries;
CPM, LPR, NCP. O. tenerum Mett.—S. Frequent but little collected. Fig. 2.

3. O. engelmanni Prantl, Limestone A. Mar-Aug. Pastures and open woods,
on pains Chickasaw, Lafayette, Lee and Madison, Oktibbeha, Warren
Cos. Fig

4. O. Latin Hook., Stalked A. Feb-Aug. Lawns and cemeteries; LPR.

ig. 2

5. O. vulgatum L. pycnostichum Fern., Southern A, Mar-Aug. Rich
Lafayette, Leflore, Marion, Monroe, Tate, Tisho-

woods and grassy areas;
mingo, and Wilkinson Cos. Fig. 8.
Many distribution records in the Ophioglossaceae were contributed by Dr

288
var.

|

Go “
R. Dale Thomas (Pers. Comm
gorgre arr ry
vi a @ & xy ad
J 4 @ege fi | |
‘@.% ee by | 5
ed YY \ | uk |_| | an |
89 . @ yi @ s |
Lf | - : eo 7 |
® é ee ¢
(eo o- ey tN @ =
Cy @ |e
Vb \ one \ ~~)
BOTRYCHIUM VIRGINTANUM BOTRYCHIUM LUNARIOIDES BOTRYCHIUM ALABAMENSE
Ss oe 4, mae
ra ® te oo | - ee
if | 4 |
a e Hota é. ée |
6 Cla co 1 @®
: | bys 0@ a le
43 G eo a
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s A, fe A e ae :
f | co. e ees ©e bad
{ | ‘e. o | < ) Pees ©
| fc eo :
Vesna Yes aa 9.9.2)
BOTRYCHIUM DISSECTUM BOTRYCHIUM BITERNATUM OPHIOGLOSSUM CROTALOPHOROIDES
Page re eae anne ae a a Fa
Kal ae 1, 1 ;
y | - @ @) Z
! fre SO | J
eee g mal seria. a
wr ec :
of VI ® ey ty ae,
P 4 e@ e 9 Cf rr | 5 Ang 4
{ aS eo Fs . jr: ae { Riines -
i ee 806 ed als ee ...
0 © a0 STUN | Le | |
383 eau 333
_ <eacase Veet ‘oe! .@)
OPHTOGLOSSUM NUDICAULE OPHIOGLOSSUM ENGELMANNI OPHIOGLOSSUM PETIOLATUM
ure 2. Distribution in Mississippi of pds virginianum, B. lu-
amense, dissectum, B. biternatum, pene cro-

Fig
narioides, B. alab ;
talophoroides, O. nudicaule, O. engelmanni, O. ieee

289
6. OSMUNDACEAE
OSMUNDA L

1. Leaves 2-pinnatifid, dimorphic, the fertile leaves without lamina.

J. cinnamomea.
1. Leaves 2-pinnate, the fertile pinnae “confined to the terminal portion of
the blade. 2. O. regalis.

1. O. cinnamomea L., Cinnamon Fern. Mar- “May. Bogs, swamps, pine, and
deciduous forests; throughout, except LBH and YMD. Fig. 3.

2. O. regalis L. var. spectabilis (Willd.) Gray, Royal Fern. Mar-June.
Woods, bogs, wet savannahs: throughout, except LBH and YMD. Fig. 3.

7. SCHIZAEACEAE
LYGODIUM Sw. Climbing Fern
1. L. japonicum (Thunb.) Swartz, Japanese C. F. May-Nov. Woods and
readsides; chiefly CPM and LPR. Naturalized from Asia. Fig. 3.

8. HY MENOPHYLLACEAE
TRICHOMANES L. Filmy-Fern
. T. petersii Gray, Dwarf Filmy Fern. All year. Wet shaded sandstone,
rare; Amite, Simpson, and Smith Cos. Fig. 3

9. DENNSTAEDTIACEAE
PTERIDIUM Gleditsch Bracken
1. P. aquilinum (L.) Kuhn. Jun-Sept. Old fields, open woods and roadsides;
throughout, except YMD. Pteris latiuscula Desv.—S; P. caudata L.—S (in
part). Fig. 3

10. ADIANTACEAE
1. Leaf blades pedate or ovate in outline; sori and indusioid leaflet margin
discontinuous. 1. dle
1. Leaf blades ovate or eee in outline: leaflet margin reflexed
ducing continuous indusioid
2. Leaves villose, ultimate se see usually 1-4 mm long. 2. Cheilanthes.
2. Leaves glabrous, ultimate segments usually 8 mm or more lon ng.

3. Pteris.

1. ADIANTUM L.
1. Leaf dichotomous in a fan-like pattern. . . . . . . - A. pedatum.
1. Leaf pinnate with a prominent central axis. . 2. A. capillus-veneris.

1. A. pedatum L., Maidenhair Fern. Jun-Aug. Rich mesic woods; LB
NCP, PR, NPB, and TRH. Fig. 3.

2. A. capillus-veneris L., Venus’-hair Fern. Jun-Aug. On limestone; Clarke
and Wayne Cos. Fig. 3.

2. CHEILANTHES Sw. Lip-Fern
1. C. lanosa (Michx.) D.C. Eat., Hairy L. F. Jun-Aug. On sandstone;
Lafayette and Tishomingo Cos. Fig. 3

290

3. PTERIS L. Brake
Spider B. May-Oct. Rocky woods;
Pycnodoria multifida (Poir.) Small—sS

Hinds, Jeffer-

1. Pteris multifida Poir.,
son, Warren and Wilkinson Cos.

Fig. 4.

|

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3°38,
ee0e

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‘>

f

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A =~

ae wel
( (© ofees ” © @ense
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OSMUNDA REGALIS

ery

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fas . K eo |
4 | e 2 |
er eh
, . | ‘ol ; .
é at fi

J ! r
Vio \ La
Cone ee

CHETLANTHES LANOSA

ADIANTUM PEDATUM ADIANTUM CAPILLUS-VENERIS

Figure 3. Distribution in Mississippi . id ea ig vulgatum var. pyc-
ectabilis, Lygodium

nostichum, Osmun is cinnamomea
japonicum, nes petersit, Peeridinm pes Adiantum pedatum,
capillus-veneris, ‘Cheilanthes lanos

291

11. ASPLENIACEAE*
1. Leaves dimorphic, the fertile leaves with sporangia borne inside of rolled

up, much contracted leaflet segments; sterile leaves coarsely lobed with
netted veins throughout. 5. Onoclea
1. Leaves not dimorphic, the sporangia borne exposed ¢ on mainly unmodified

segments, the fertile segments never rolled up; veins free, or only par
tially anastomosing (or veins anastomosing and sori elongate and irregu-
larly shaped following along the veins in Asplenium rhizophyllum)
2. Sori round, without indusia; leaves herbaceous, ey triangular,
Thelypteris.
2. Sori round or elongate, wae indusia; leaves of various textures, nar-
]

oe

3. Sori elongate, linear or ee eeonai ned: indusium attached to leaf

along one

4. Sori in parallel lines along the midrib of the Sapa or the pin-
nules, ns anastomosing along the midrib and free near the lobe
mar 8. Woodwardia.

4. Sori diverging at an acute angle from the midrib of the pinnae or
innules; veins free.

5. Scales of the rhizome with blackish, thick secondary lateral cell
walls; at least the base of the stipe polished black or brown;
sori straight. 6. Asplenium.

d. Scales of the rhizome with similar ‘thin lateral, dorsal and ventral
cell wall faces; stipe base dull green, reddis sh or stramineus:
sori straight, arched or hooked. 4, Athyrium.

3. Sori round or kidney-shaped, indusium attached beneath sorus.
3. Indusia composed of more or less separated flaps or filaments,
these beneath and around the eens cluster like a cup
2. 0dsia.
6. Indusia composed of single units not divided into flaps or Fre
the shapes variou

7. Indusium hood- like, attached by a broad base beneath the sorus

and arching over the sorus from one side, not ea or reniform
: . Cys stopteris.

7. Indusium peltate or renifor
8. pcteaeee peltate, ntiached: to leaf by a median stalk through

sorus; sori confined to terminal constricted lea — ts.

7. Polystichum.
8. Indusium reniform, attached to leaf f along the sinus; sori on
unmodified leaflets. . 1. Thelypteris.

1. THELYPTERIS Schmidel
1. Blade triangular

2. Leaves 1- pinnatifid, the pinnae lobed and with adnate wings along the
rachis joining the pinnae. T. hexagonoptera.
2. sor es 3-, 4-pinnatifid, the pinnae without ‘wide ‘attachment to the
achis 2. T. torresiana.

1. Blade lanceolate to oblong- lanceola te,
3. Blade base with 3-7 pairs of oe! reduced to eat pinnae.
3. noveboracensis.

The i name “‘Aspidiaceae,” or Polypodiaceae sensu Jato, has often been ap-
olied to this grou

292

3. Blade base lacking rudimentary pinnae, the lowest pinnae only slightly
reduced or not at all.
re Basal veinlets of adjacent pinna lobes running toward the sinus with-
out uniting, or uniting at the base of the sinus.

5. Rhizome short-creeping to suberect; leaves spreading, with soft
pubescence, tending to become light green or yellowish at maturity,
usually less than 0.5 m tall; veins simple.

4. quadrangularis var. versicolor.

5. Rhizome short- or - long- creeping to suberect; leaves stiffer, dark
to light hanes at maturity, usually more than 0.5 m tall; veins
simple or for
6. Leaves light green; margins of fertile fronds usually revolute;

lateral veinlets of oe mostly forked; segmen ee cut nearly

to the pinna axis. 5. T. palustris.

. Leaves dark green; mar -gins usually flat: lateral veinlets of seg-

ments simple; segments cut only 0.5 to 0.75 of the way to the

T. thit.

4. Basal veinlets: of adjacent pinna lobes uniting below the sinus with
a co n vein extending to the base of the

7. Petiole each leaves tending to - slightly aoe the earliest

maller, wider, spreading and sterile, the later ones more erect,

wer and fertile; the united vein to the sinus ae Ae pinnae

for)

being shallowly lobe dentata.
7. Petiole yellowish; leaves “not dimorphic: “united ° vein ees the pin-
nae deeply lobed. 4. T. quadrangularis var. versicolor.

1. T. hexagonoptera (Michx.) Weath., Broad Beech Fern. Apr-Sept. Rich
hardwood forests, ravines and loess hills; scattered throughout except YMD.
Phegopteris hexagonoptera (Michx.) Fé ris hexagonoptera
(Michx.) C. Chr.—F. Fig. 4

2. T. torresiana (Gaud.) Alston, Mariana Fern, May-Oct. Wet woods and
stream banks; scattered in LPR, and Marshal and Warren Cos. Dryopteris
setigera (Bl.) Ktze.—S(misapplied). Fig. 4.

3. T. novaboracensis i) Nieuwl,, New York Fern. May-Oct. Rich woods;
TRH. Dryopteris no acensis (L.) Gray—F. Fig. 4.

4. T. Sinden “(Fée) Schelpe var. mene (R. St. John) A. R.
Smith, Variable Maiden Fern, May- ae Rich deciduous woods and ravines;
scattered in LBH, LPR, and Scott Co. T. versicolor R. St. John—S. Fig. 4.

5. T. palustris Schott, Marsh Fern. at Wet roadsides, stream banks,
marshy lake shores, bogs and wet woods; CPM, LPR, and Attala Co. Dryop-
teris thelypteris (L.) Gray-F; T. thelypteris (L.) Nieuwl.—S. Fig. 4.

;. T. kunthii (Desv.) Morton, Widespread Maiden Fern. Apr-Nov. Wet
aonds: roadsides, stream banks, and sandy flatwoods; CPM; LBH, ‘LPR,
and NCP. T. normalis (C.Chr.) Moxley—S. Fig. 4.

7. T. dentata (Forssk.) C. Chr., Tapering Tri-vein Fern. Jun-Aug. Edge of
mixed woods and sandy loam; Forrest, Lauderdale and Pike Cos. Fig. 4.

This treatment follows that of Smith (1971).

2. WOODSIA R. Br.
1. Woodsia obtusa (Spreng.) Torr., Blunt-lobed Woodsia. May-Aug. De-

293

ciduous woods, sandy soil and sandstone outcrops; scattered in LBH, LPR,

NCP, NPB, and TRH.

ree. o_@
Pee oe? i |
a a x J fr = |
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wets

THELYPTERIS HEXAGONOPTERA

eA oo Tit Ty)
~ aaa Pa i
C Lap ye a
eo Cot
ee gif te
ASV | \,®
oom Ee fj. ee. ©
(ee * eo. (9 Ce
/ (es @ 8

THELYPTERIS QUADRANGULARIS THELYPTERIS PALUSTRIS

THELYPTERIS NOVABORACENSIS
Daa eal Oe aia aa ee
| @ @
Ful es J ari
oy J
ri °
ee! |} | fi ee
2 eo
Pp |
ko o ie:
pro] | ‘ @
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gee on

WOODSIA OBTUSA

THELYPTERIS KUNTHII THELYPTERIS DENTATA

igure 4. Distribution in Mississippi of Pteris multifida, oo hexa-
gonoptera, T. torresiana, T. novaboracensis, T. quadrangi . versi-
color, T. palustris, T. kunthii, T. dentata, Woodsia obtusa

294

3. CYSTOPTERIS

Bernh. Bladder Fern

Spreading B.F. Apr-Jul.

Rich woods,

1. C. protrusa (Weath.) Blasdell,
ravines and loess bluffs; LBH and JP. C. fragilis (L.) Bernh. var. protrusa
Weath.—F, G; fragilis (L.) Bernh.—S. Fig.

@ ] ; a |
JD { | f ry i
e hs 4 | Fe + |
s {—_ __}—* ‘2 ; { |
C Lda aa
of eh ot et
4 Ra ttt .@ |
®e50@ ‘oe
> A {
CYSTOPTERIS PROTRUSA ATHYRIUM ASPLENIOIDES ATHYRIUM PYCNOCARPON
¢®@ -@ eile e e @
f ae Pat @ J |
i @ @ = ee | | ¢
s e di F + ‘ | 5S
4 @ : | % { . ; ia &
- @e@ ©@ gia eT br |
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ONOCLEA SENSIBILIS ASPLENIUM RHIZOPHYLLU
ray ene
+ @ ‘si |
- oe Se aepR sae
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so °os ly
Be eee . |
ee oe
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glo
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*,
— AJ L ~
ASPLENTUM PLATYNEURON ASPLENTUM TRICHOMANES POLYSTICHUM ACROSTICHOIDES

Figure 5. Seas in — of Cystopteris protrusa, Athyrium
es ocarpon, oclea sensibilis, Asplenium rhizophyllum,
Polystichum acrostichoides

asplenioid
A. pinnatifidum, A. ee on, A. trichomanes,

295

4. ATHYRIUM Roth
1. Leaves 2-3 pinnate; sori crescent-shaped, ee 1 mm or less long.
. asplenioides.
1. Leaves 1-pinnate; sori nearly straight, usually 2 mm or more long
pycnocarpon.
1. A. asplenioides (Michx.) Desv., Southern Lady Fern. May- Oct. Swamp
forests, stream banks, marshes and lake shores, wet woods and ravines;
throughout except YMD. A. filix-femina (L.) Roth var. asplenioides (Michx.)
Farw.—F, G. Fig. 5.
2. A. pycnocarpon (Spreng.) Tidestr., Glade Fern. Jul-Aug. Rich woods
and ravines; Claiborne, Lafayette, Tishomingo, Warren, and Yazoo Cos.
Homalosorus pycnocarpus (Spreng.) Small—S. Fig. 5

3. ONOCLEA L. Sensitive Fern
1. O. sensibilis L. Jul-Nov. Swamps, bogs, stream banks, wet roadsides,
wet woods and ravines; throughout except YMD, CPM, FW, and PR. Fig. 5.

6. ASPLENIUM L. Spleenwort
1. Leaves undivided or lobed —— = veins ee a
: 1. A. rhizophyllum
1. Leaves deeply lobed or pinnate: veins free

2. Blades deeply lobed at most, or pinnate at the base only; apex narrowly
caudate. . . 2. A. pinnatifidum.

2. Blades pinnate throughout; apex not ‘cauda te,
3. Pinnae alternate, sessile, the bases usually overlapping the rachis;
leaves somewhat tae the fertile ones upright and taller than

the spreading sterile leav 3. A. platyneuron.
3. Pinnae opposite, short steed. “the bases not overlapping Ba rachis;
s monomor 4. A. trichomanes.

1. A. rhizophyllum L., Walking Fern. May- Oct. Sandstone eee Tisho-
mingo and Union Cos. Guinea soni rhizophyllus (L.) Link—F, G, S. Fig. 5.

2. A. pinnatifidum Nutt., Lobed S. May-Oct. Dry sandstone ledges; Tisho-
mingo Co. Fig. 5

3. A. platyneuron (L.) Oakes, Ebony S. May-Oct. Woods, ae. road-
sides, pastures, sandy soil and sandstone outcrops; throughout.

4. A. trichomanes L., Maidenhair S. May-Oct. Sandstone su ee Tisho-
mingo and Noxubee Cos. Fig. 5.

7. POLYSTICHUM Roth
1. P. acrostichoides (Michx.) Schott, Christmas Fern. Apr-Aug. Rich de-
ciduous pine and mixed woods, ravines, swamp woods, stream banks, loess
and sandy soil; throughout except YMD. Fig. 5

8. WOODWARDIA Sm. Chain-Fern

1. Leaves dimorphic, the fertile much contracted, the sterile pinnatified to
vag leaflet bases ee adnate except at blade base; veins form-
etwork throughou 1. W. areolata.

: Sterile ana fertile leaves nee pinnate- -pinnatifid, ‘the leaflet ee fully

a

296
contracted throughout; veins free except along midribs of segments.
2. W. De eer.
1. W. areolata (L.) Moore, Netted C. F. Aug- Nov. Wet places, lake shores,
low woods and ravines; throughout except
. Fig. 6.

stream banks, swamps, bogs
YMD and LBH. Lorinseria areolata (L.) Presl—sS. Fig

ate ee.
POLYPODIUM POLYPODIOIDES

WOODHARDIA VIRGINICA
mi

“S82 S|

WOODWARDIA AREOLATA

filet ias
@
\PtSse Agen
AZOLLA CAROLINIANA ISOETES ENGELMAN!

ss ie > aaa Coe aes if + RES 5 T
¢ f ) { .]
peo) cae oe tee
Pa ey Pa aul 2 | |
é aren oj. eee ¢ ihe A
f |} . ‘e Ss . Sa ee a 4 } p ‘|

5 Le reel tet PL ff $4) inom tes

wh eee ee a ee ee

oA gd f by Pye lee

AN Sart ra b~r Tw] |
| Bete fo] tty \

e 6. Distribution in Mississippi of Woodwardia areolata, W. virginica,

Spee an polypodioides, Azolla caroliniana, Isoetes engelmanni

297

2. W. virginica (L.) Sm., Virginia C. F. May-Aug. Wet roadsides, pine bar-
rens, bogs, swamp forests and wet woods; CPM, LPR, and NCP. Anchistea
virginica (L.) Presl.—S. Fig. 6

12. POLYPODIACEAE sensu strictu
POL UM L. Polypody
1. P. polypodioides (L.) Watt, Resurrection Fern. All year. On trees and
rocks, hardwood, pine and mixed forests, cypress swamps, pastures, flood-
plains, and ravines; throughout except YMD. Marginaria arn (L.)
Tidestr.—S., P. polypodioides var. michauxianum Weath.—F.

13. AZOLLACEAE
AZOLLA Lam. Mosquito Fern

1. A. caroliniana Willd. Jun-Sept. Floating or resting on mud, lakes,
swamps, and ponds; Bolivar, Noxubee and Pearl River Cos. Fig. 6

REFERENCES
ee J. K. 1938. Ferns of the southeastern states, 1964 facsimle edition. Hafner, New

Son M. L. 1950. Gray’s manual of botany, 8th Ed. pmenean Book Co., New York.
GLEASON, H. A. si A. CRONQUIST. 1963. Manual of vascular plants of Pel sears
United States and adjacent race Van Nostrand-Reinhold Co, New Yor
. , A. E., oe E. Ahles, and C. R. Bell. Manual of the vas-
cular flora of the ea. University of North Carolina Press, Chapel F
y W. R. BUCK. 1975. mee Seis S. apoda a their hybrids
in the southeastern United States. Amer. Fern 76-82.
SMITH, A. R. 1971 ere o the Ran species of Thelypteris Sec. Cyclosorus.
Univ. Calif. Publ. Bot. 59: 1-13

THE CHIHUAHUAN DESERT SPECIES OF
ERICAMERIA (COMPOSITAE: ASTEREAE)

LOWELL E. URBATSCH
Department of Botany
Louisiana State University, Baton Rouge, LA. 70803

The purpose of this paper is to treat the Chihuahuan Desert species of
Ericameria in advance of the flora being prepared for that region by M.
Johnston et al. In this report one new combination is made and the taxon-
omy and distribution of the other four indigenous species are given. I con-
cur with taxonomists who give Ericameria generic status (e.g. Johnston,
1970) rather than with those who treat it as a section in the genus Haplopap-
pus (e.g. Hall, 1928). Ericameria comprises ca. 20 species of shrubs (Ur-
batsch, ined.), and California is its center of greatest species diversity.

The Chihuahuan Desert species of Ericameria are novelties. They stand
apart from one another and from their California relatives in that each has
a unique flavonoid complement and one or more extraordinary morpho-
logical features (Urbatsch, ined.). However, two characters, their zygomor-
phic disk corollas and their relatively long, thick style branches, give the
Chihuahuan Desert species unity. Perhaps the pattern of diversity expressed
by these species suggests that they are relicts that evolved from a common
extinct ancestral group which also gave rise to the California species. Alter-
natively, the Chihuahuan Desert species may be recent evolutionary prod-
ucts that rapidly evolved in response to drastically changing environmental
conditions. Certainly, the apparent morphological and chemical remoteness
among the Chihuahuan Desert species and between them and the other Eric-
amerias offers a challenge to cladistic formulation.

Ericameria is closely allied with the genus Ch t} s, and it is often
confused with the genera Isocoma and Sia lun: Morphological fea-
tures that distinguish the four genera are given in Table

Key to Species of Chihuahuan Desert oammeria
1. Leaves acicular, non-resinous; ray flowers absent. . 5. E. purpusii.
1. Leaves linear, not rigid and ee like, usually resinous: ray flowers ab-
sent or present.

2. Ray flowers present; heads with 7 or more flowers, phyllaries imbricate.
3. Middle and upper he elliptic to obovate with a green to tan
subapical patch.

Leaves widely spaced, internodes 5— “la mm long, stems appearing

‘Jeafless’’; heads with 7-14 flower 1. E. pseudobaccharis.

4. Leaves crowded, internodes 1-4 mm long, stems leafy; heads with

ers. E. parrasana,

3. Middle ee subulate with medial, prominent, dark- Soke costa.

3. E. laricifolia.

2. el flowers absent; heads with 3-7 ‘flowers: phyllaries in subver rtical
S. 8 4

antha.,

SIDA 7(3): 298-303. 1978.

299

Taxonomic Treatment

1 ER haris (Blake) Urbatsch, comb. nov.—Hap-
lopappus- -pseudobaccharis Blake, J. Wash. Acad. Sci. 40:47. 1950. TYPE:
MEXICO: Coahuila: arid limestone hillside, Valle Seco, Sierra de la Paila,
General Cepeda, 4 July 1944, G. B. Hinton et al. 16546 (HOLOTYPE, US!;
ISOTYPES, GH!, NY!)

Ericameria pseudobaccharis is known only from south central Coahuila,
1200-1500 m; it apparently flowers when conditions are favorable.

Ericameria pseudobaccharis was placed in synonymy with E. triantha by
Johnston (1970) who had only limited material available. This species re-
mained relatively unknown until James Hendrickson and co-workers redis-
covered it at several different localities in Coahuila. The morphological
range of variation for E. pseudobaccharis and E. triantha is now better un-
derstood and they are readily distinguishable. In E. pseudobaccharis the
heads are solitary or disposed in racemes, radiate, 7-14 flowered, and have
imbricate phyllaries, whereas in FE. triantha the heads are in cymose Clus-
ters, eradiate, 3-7 flowered, and have phyllaries in subvertical files. The
two taxa have only one flavonoid in common while they differ in twelve

TABLE 1. Comparison of Key Morphological Features that Distinguish
Ericameria from Similar Genera.

Genus’ Ericameria Chrysothamnus Isocoma Xanthocephalum
Character
entire entire serrate, entire
margins lobed or
entire
phyllary imbricate to vertical files imbricate imbricate
arrange- sub-vertical
ment files
is gradually gradually abruptly gradually
corolla ampliate ampliate inflated ampliate
tubes
pappus bristles bristles bristles scales or
ent
heads discoid or discoid discoid radiate

radiate

300

compounds (Urbatsch, ined.). The species are adapted to different habitats,
I. pseudobaccharis occurs on limestone slopes and sometimes in gypseous
areas, while E. triantha is commonly found in gypseous soils on desert flats.

Additional specimens exa mined: eee eee ca. 18 (air) mi NE
of Tlahualilo, in Sica de Tlahu ee ca. 9 mi NW of Los Charcos de Risa,
on small limestone peak, ca. 1. i E of main ae 30 Sep 173, Hendrickson
13720 (LL); ca. 32 (air) mi NE ae Tlahualilo, in the NW portion of the Sierra
de las Delicias, in the first canyon south of the Puerto de las Sardines, on
limestone, infrequent, 9 Aug 1973, Hendrickson 12216 (LL); ca. 67 (air) mi
SW of Cuatro Ciénegas, 1.5 mi SW of Las Delicias in FE side of Sierra de las
Delicias, > Aug. 1973, re-neere-reems 12467 (LL); ca. 62 (air) mi WsW of

of Guestade Gallo, 8 Aug 1973, esenloentonnn 12110 (LL); ca. 9.2 (rd) mi §
of El Hundido, in Izotal, frequent, 29 Sep. 1973, Hendrickson 1376a (LL).

2. ERICAMERIA PARRASANA Blake, Contr. Gray Herb. 52:26. 1917.
TYPE: MEXICO: Coahuila, rocky slopes, Sierra de Parras, March 1905,
Purpus 1005 (HOLOTYPE, UC!; ISOTYPE, GR!). zn NORGE parrasanus
(Blake) Blake Contr. U.S. Natl. Herb. 23:1490. 1926

Ericameria parrasana is known only from the ‘alice obscure type locality
and the one cited below. It apparently flowers when conditions are favorable.

Additional specimens examined: MEXICO: ZACATECAS: 2 km §S of San
Miquel on road to Cedros, 24° 55’ 50”N, 102° 07’W, 1570m; 1 Jul 1978, John-
ston, Wendt & Olnane 11542 ee same location as preceding; 22 Aug 1974,
Urbatsch et al. 1942

Ericameria parrasana was known only from the type collection until re-
cently rediscovered by M. C. Johnston. Hall (1928) maintained that E.
parrasana and Haplopappus linearifolius comprise Haplopappus sect. Steno-
topsis, which Hall regarded as remote from sect. Ericameria. In my judg-
ment both species in Stenotopsis belong to Ericameria. Although E. parra-
sana and H. linearifolius are similar in pappus color, head size, and leaf
glands, they differ markedly in several other features such as style-branch
morphology, capitulescence type, and flavonoid complements. The latter spe-
cies appears to be closely allied to E. cooperi (Urbatsch, ined.). The affini-
ties of FE. parrasana are not readily apparent. Its flavonoids are more like
those of EF. palmeri. Clearly more study is needed before the relationships
of BE. — are known with certainty.

3.E MRIA LARICIFOLIA (A. Gray) Shinners, Field and Lab 18:27.
sere lariciiolitis Gray, Pl. Wright. 2 (Smithsonian Contribution
Vol 5, art. 6): 80. 1853. TYPE: U.S.A.: NEW MEXICO, Hidalgo Co.: Guada-
lupe Pass, 5 Oct 1851, Wright, accession number 1188 (HOLOTYPE, GH!;
ISOTYPES, MO! US!). Aster laricifolius (A. Gray) Kuntze, Rev. Gen. Pl.
318. 1891. Chrysoma laricifolia (A. Gray) Greene, Erythea 3:11. 1895.
Bigelowia nelsonii Fernald, Proc. Amer. Acad. Arts 36:505. 1901. TYPE:
MEXICO: CHIHUAHUA: Sierra Madre, 29 Sep. 1899, Nelson 6494 (HOLO-
TYPE, GH!; photos of the type NY! UC!). Ericameria nelsonii (Ferna!d)
Blake, Contr. Gray Herb. 52:26. 1917.

Ericameria laricifolia ranges from eastern San Bernardino Co., California

301

to Mohave Co., Arizona southeastward to northeastern Sonora, Mexico and
Presidio Co., Texas. This species is often locally common, usually growing
in stony soils of foothills and from crevices of canyon walls, 600-2500 m.
Flowering is from mid-September into November; occasionally a few some-
what larger vernal heads are produced in April and May. Ericameria larici-
folia is not closely related to the other Chihuahuan Desert species, but is
allied to the California species, E. pinifolia and E. brachylepis.
Representative Chihuahuan Desert region specimens examined: MEXICO:
CHIHUAHUA: mtns., near Ciudad Juarez, 1911, Stearns s.n. (NY); SW
slope of Sierra de la Rancheria (on Rancho Candelaria) from the top down
to a marble quarry at base, 31° OVN, 106° 21’W, elev. 1500-2180 m, among

near Silver mak elev. 6000 ft., 19 Sep. 1903, Metealfe 74 (ARIZ, GH, MO,
NY, UC, US); Dofia Ana Co:: ‘Ore an Mts., 18 Oct 1903, Wooton s.n. (ARIZ,
MICH, NY): Hidalgo Co.: Ae an N. fe of Big Hatchett Mts., 18 Aug
1954, Castetter 9372 (SMU); a Filonda eke ‘Ss Slope, alt. 5800 ft,
8 Sep 1908, Goldman 1485 aan XAS: El Pas : on mtns. above Mc-
Kelligon Canyon, Franklin Mts., a about 3 rt ei 16 Oct 1952, Correil

Vv
northside Chinati Mts., old Woods ranch, 18 mi NW of Shafter, 10 Nov 1946,
aan & Warnock 46961 (GH, MO, TEX, UC).

a \NTHA (Blake) Shinners, Field and Lab. 19:133.
= Heplonaner: trianthus Blake, J. Wash. Acad. Sci. 28:485. TYPE: U.S.A.:
Brewster Co., frequent along road from Study Butte to Terlingua, Chisos
Mtns. area. 31 Aug 1937, Warnock 1126 (HOLOTYPE, US!; ISOTYPE, LL!).
Ericameria triantha occurs from Brewster Co., Texas southward through-
out most of Coahuila, eastern Chihuahua, northeastern Durango, and north-
western Nuevo Leén from 700-1500 m. It is locally frequent in gypseous, cal-
carous, or saline habitats with gravelly to fine alluvial soils and flowers

August to October and to some extent throughout the entire year.

The nearly vertical alignment of phyllaries of E. triantha suggests that
it may well belong in the genus Chrysothamnus. However, anatomical studies
by Anderson (1970) show that E. triantha, unlike Chrysothamnus, has the
well developed secretory canals in its achenes and corollas aharaciencnc of
species in Ericameria. The flavonoid complement of triantha is most similar
to that of E. diffusa (a species centered in Baja California) which Blake
(1938) had suggested as its closest ally (Urbatsch, ined.).

Representative specimens examined: MEXICO: COAHUILA: west side of
Luguna del Rey; 18 Oct 1971, Bacon & Leverich 1197 (TEX); Laguna Salada,
salt flats about lake; 11 Aug 1967, Cole, Minckley & Pinkava 3645 (ASU);

mi W of La Vibora on small gypsum knoll near railroad, in Bols6n de
Mapimi region of Chihuahuan Desert; 21 Sep 1972, Henrickson 7868 (LL);
10 mi S of Castill6n, 12 Aug 1970, Johnston & Muller 195 (CAS, GH, LL,
MICH): CHIHUAHUA: road from Castillon to Mula, via S. Salvador and
Piramide, 4 mi N of Mula; 21-22 Sep me ees & Muller 1439 (GH, LL);
> mi E. of Carrillo, municipio de Jim 15 Sep 1939, Muller 3319 (GH,
MICH, TEX, UC); DURANGO: road en aa illo to Zaragoza and north
to Mohovano, 31 mi N of Zaragoza; 20 Sep 1938, Johnston 7799 (GH, UC);

302

NUEVO LEON: 3 km from EI Milagro and 10 km from Icamole on winding
rd. between, 25° 55’ 30”N, 100° 47’°W, 720 m, 5 Jul 1973, Jonston, Wendt, Chi-
ang 11612 (LL): U.S.A.: TEXAS: Brewster Co.: near Hot Springs, Big Bend
Nat’l Park, limestone flag hills; 13 Sep 1961, Correll & Jonston 94557 (LL,

5. ERICAMERIA PURPUSII Brandegee, Univ. Calif. Publ. Bot. 4: 191.
1910. TYPE: MEXICO: COAHUILA: Cerro de Macho, June 1910, Purpus
4479 (HOLOTYPE, UC!) Haplopappus purpusii (Brandegee) Blake, Contr.
U.S. Natl. Herb. 23: 1491. 1926.

Ericameria purpusii is distributed from western Coahuila and southwestern
Chihuahua south to central Durango at 1100-1200 m. It is sometimes locally
frequent but usually uncommon on limestone hills, and sometimes in gypse-
ous soils; it flowers in May and Aug-Sep.

The singularity of this species is manifest by its acicular, non-resinous
leaves, and unique flavonoid complement (Urbatsch, ined.). E. purpusii is
not readily accommodated in any existing genus, but its habit, involucral
and pappus features, and chromosome number, n=9, place it closest to
Ericameria. No useful purpose would be served by erecting a monotypic
genus next to Ericameria for this taxon.

Hall (1928) doubtfully assigned E. purspusii to Haplopappus sect. Asiris
(a taxon centered in the Rocky, Sierra, and Cascade mountains) based
mainly on a single technical feature, lack of impressed resin pits. However,
corolla and style branch features and consideration of its geographic occur-
rence place it closer to the other Chihuahuan Desert species of Ericameria.

Additional specimens examined: MEXICO: CHIHUAHUA: southern part
of Las Pampas Ranch, Preson El Lindero, 27° 13’ 30’”N, 104° 42’W, 1650 m,
26 Aug 1972, _ Chiang, Wendt, Johnston 8936H (LL); 6 km ESE of Rancho

104° 37°W, 1575 m, 29 Aug 1972, Johnston, Wendt, Chiang 8984 (LL); COA-
HUILA: 5.5 mi E of Los Americanos, Fas 103° 15’°W, 1080 m, 2 Sep
1972, Chiang, Wendt, Johnston 9117 (LL); . 29 rd. mi ESE of Esmaralda,
27° 10’N, 103” 17’°W, 20 Sep 1972, Hendrickson 7846 (LL); road from San An-
ie de los Alamos, 30 km E through low limestone hills to Buenavista,
27° 30’'N lat., W of Buenavista, 21 Aug 1941, Johnston 8308 (LL); S of
oe de Lec he, 30 Aug 1941, Johnston 8614 (LL); S of Matrimonio Viejo,
22 Sep 1941, Johnston 9365 (LL); E of Americanos, 23 Sep 1941, Johnston
9380 (LL); 9.5 m WNW of Hacienda Zacatosa, 27° 02’ 30”N, 102° 46’W, 1225
m, 7 May ae es Wendt, Chiang 10906 (LL); 7.5 km N of Rancho
Presa de Lumbre on rd. to Zacatosa, 27° 59’N, 102° 53’ 30”W, 1225 m, 7 Aug
1973, Johnston, Chiang, Wendt, Hendrickson 12105 (LL); DURANGO: 52.5
km W of Ceballos on rd. to Villa Hidalgo, a few km due S of Buen Dia, 24°
19’N, 104° 30’W, 1400 m, 31 Oct 1972, Chiang, Wendt, Johnston, 9984 (LL).

ACKNOWLEDGMENTS
I am grateful to B. L. Turner, M. J. Johnston, James Hendrickson, and
Ronald Hartman for their help with various aspects of this study. Also, I
thank the curators of the various herbaria for their generous loans of speci-
mens. Financial support received from NSF Grant GB-40362, a grant-in-aid

303

of research from the Society of the Sigma Xi, the LSU Department of Bot-
any, and the Department of Botany, University of Texas, Austin, is acknowl-
edged.

REFERENCES
ANDERSON, - C. 1970. Floral anatomy of Chysothamnus (Astereae, Compositae) Sida
3: 466—50
BLAKE, S. F. ioe: Eleven new American Aste Jour. Wash. Acad. Sci. 28: 478—492,
gee H. M. 1928. The genus Haplopeppae—& paveres study in the Compositae.
negie ve ods of Washing 2
Sor OM M. C. 1970. In Correll, D. S. ae ML C. Johnston. Manual of the vascular
plants of Texas. Texas Research Foundation, Renner, TX.

NOTES

LOBELIA NUTTALLIL (CAMPANULACEAE) IN SOUTHEASTERN
LOUISIANA.—On 4 Jun 1976, Lobelia nuttallii R.&S. was discovered 12 miles
northeast of Chalmette, Orleans Parish, Louisiana, during the botany field
trip of the 1976 AIBS New Orleans meetings. The plants are characterized
by tiny blue flowers, narrow stem leaves, and pedicels bracteolate at the
base. This station extends the species distribution approximately 100 miles
west from southern Alabama according to MeVaugh (‘‘North American
species of Lobelia.’ Rhodora 38: 352-353, 1936) and Prof. Joseph Ewan,
Tulane University, New Orleans (pers. comm.). Small (Manual of the south-
eastern flora, 1933) reported the species for Mississippi, but specimen docu-
mentation is apparently lacking.

The plants—fewer than 50 of them—were scattered along a sandy road-
side bank above a saline marsh bordering the waters separating lakes
Pontchartrain and Borgne. The annual habit of the species and the type
of habitat in which the plants were found suggest that the species may have
been unintentionally introduced from some more easterly region. The speci-
men (Brooks 12099 and Hauser) is on deposit at the University of Kansas
Herbarium (KANU).—Ralph E. Brooks and Larry A. Hauser, State Biologi-
cal Survey of Kansas and University of Kansas Herbarium, Lawrence, KS
66044.

NOTES ON COREOPSIS—Since the publication of my summary paper on
eastern United States and Canadian Coreopsis (Sida 6(3): 123-215. 1976),
a quantity of type material from PH has been called to my attention and a
number of important distributional records have been collected. [ am pub-
iishing this note to mere critically document my summary paper and to add
the distributional records to the literature. My thanks to the curators of
APSC, FSU, GA, NCU, NO, PH, and WIS, who kindly asked me to annotate
their material. Special thanks to Dr. J. A. Mears, who called my attention
to the type material at PH, and to Dr. A. E. Radford, who pointed out the
rosea collections from S. Carolina.

>

I. Important specimens at PH

1. C. cardaminefolia (DC.) Nutt. var. angustiloba T. G.
PH Acq. No. 918074 HOLOTYPE! (the n name ee to the plant on the
right). This = = C. tinctoria Nutt. var. tinctor

Horn Island, Miss. S. Mi. Tracy 8542. 25 May 1903. U. Penn. Acq.
90382. ISOTYPE! This = C. pubescens Ell. var. debilis (Sherff) E. B.

ith
3. C. X delphinifolia Raf. var. ae see ma concolor Sherfft
Tatnall Co., Ga. Moist swale in sandhills 3 1 of Reidsville.
A. Cronquist 5598. 3 Aug 1948. PH Acq. No. "899747. ISOTYPE! This =
C. X delphinifolia Raf.

w

SIDA 7(3): 304. 1978.

—

_

—

~

~

305

4. C. oe (D. Don) T. & G. var. wrightii Gra
Wri vight Texas. (N. Mex.) 1851. PH Acq. No. 918087. ISOTYPE!
is = Phelesperma a
0. C. floridana E. B. i
Low pine barrens, Te tae bieele. Florida. Biltmore Herbarium 2066°.
22 Sep 1879. I mentioned a duplicate of this at NY as “clear rly ce a
’ in my summary paper. This is also C. floridana E. B. Sm
grandiflora Hogg ex Sweet var. harveyana (Gray) Sherff ae de-
mareei Sherff
Bradley Co., Arkansas. Prairies. P. O. Warren. Elev. 150 ft. D. demaree
1926A. 7 Jun 1939. PH Acq. No. 816060, ISOTYPE! This = as determined.
7. C. grandiflora Hoge ex Sweet var. pilosa Sherff
PH Acq. No. 556249. ISOTYPE! This — C. lanceolata L.
. C. harveyana Gra
Cliffs near Ft. Smith, Arkansas. F. L. Harvey s.n. June 1885. PH Acq.
No. 918071. ISOTYPE! This = C. a. Nutt. var. tinctoria
9. C. heierogyna Fern
Rich alluvial woods and thickets back of sand- beach of James River,
below Sunken Meadow Beach. M. L. Fernald & B. Long 8506. 14 Jun 1938.
PH Acq. No. 918086. ISOTYPE! This = C. lanceolata L.
10. C. heterophylla Nutt.

H Acq. No. 918072. HOLOTYPE! This = C. lanceolata L. (the plant
on the left side of the sheet). Note: The name C. heterophylla Nutt.
should be listed as a synonym of C. lanceolata L. (not C. grandiflora
Hogg ex Sweet var. grandiflora tte oe as in my summary
paper). Red River, Arkansas. Nuttall s.n. PH Acq. No. 918092. ISOTYPE!

is = C. grandiflora Hogg ex ee var, harveyana (A. Gray) Sherff
forma harveyana

1. C. leavenworthii T. & G. var. curtissii as
Dry ground along Suwanee River, Florida. H. Curtiss 6734. Oct. 1900.
PH Acq. No. 918085. ISOTYPE! This — C. seen T. & G.

. C. linifolia Nutt.
rates s.n. Alabama. PH Acq. No. 918070 and 917971 (2 sheets). ISO-
TYPES! This = as determined,

3. C. nudata Nutt.
St. Mary’s, W. Florida. Baldwin s.n. PH Acq. No. 918069. HOLOTYPE!
and 918084 ISOTYPE! These = as determined.

oe

co

bo

rn.
Nansemond Co., Virginia. Springy roadside bank north of Factory Hill.
M. L. Fernald & B. Long 9655. 13 Oct. 1938. PH Acq. No. 918083. TOPO-
TYPE! This = C. linifolia Se

9. C. oniscicarpa her var. sim s Fer

Nansemond Co., Virginia. aie ‘ditch ¢ 4 border of low woods northeast
of Baines Hill School. M. L. Fernald & B. Long 13827. PH Acq. No.
918079, 918080, and 918081 (3 sheets). ISOTYPES!
Same location. M. L. Fernald & B. Long 14061, 12 Sep 1941. TOPOTYPE!
All of See = C. linifolia Nutt

6. C. palmat utt.
(St. Lou fiey eee PH Acq. No. ran HOLOTYPE! (plant on the
right side of the sheet). This = as determined. Note: This takes prece-

dcnce over the neotype that I named in ae a paper.

SIDA 7(3): 305. 1978.

306

17.

—
io 2]

bo
co]

bo
—

QO

S)

'o)

a

QO

me He Nut
tt.) N. So PH ae, No. 918067. HOLOTYPE! This = as are
ee Leavenworth s Georgia. PH Acq. No. 917913. This =
termined, This een collection indicates that the range i C.
rosea did (but probably does not now) exte tend to Georgia, as stated by
Nuttall

._ C. saxicola Alex

DeKalb Co., Georgia. Dry, light soil, Stone Mountain. I’. W. Pennell 4029.
2 Aug 1912. U. Penn. Acq. No. 61145. Ren: This = C. grandiflora
Hogg ex Sweet var. saxicola (Alex.) E. B. Sm

_ C. stellata Nutt.

PH Acq. No. 918065. HOLOTYPE! es plant on the left). This = C.
major Walt. forma oemleri (Ell.) She

wes pd Nutt

(Nutt.) A PH ‘Ac eq. No. 918064. HOLOTYPE! Red River, Ark. PH Acq.
No. 918073. ee These — C. tinctoria Nutt. var. tinctoria

. C. wrayit

i Nutt.
(Nutt.) N. Carol. PH Acq. No. 918063. HOLOTYPE! This = C. X del-
phinifolia Raf. Note: C. wrayit Nutt. should be listed as a synonym of
C. X delphinifolia Raf., not as a a m of C. major Walt. forma
oemleri (Ell.) Sherff as in my su ary paper. This specimen indicates
that N. Carolina is (or was) in the — of C. X delphinifolia Raf.

II. Important distributional records
grandiflora Hogg ex Sweet var. grandiflora Sages grandiflora
FLORIDA: Liberty Co.: Godfrey 75183 (FSU, UARK), first record for
Florida. MISSISSIPPI: Alcorn Co.: Temple a (GA); Lawrence Co.:
Temple 12283 (GA), first records for Mis eee

_ grandiflora Hogg ex Sweet var. saxicola (Ale

ith
MISSISSI a Alcorn Co. Temple 6597 (GA), first record Mississippi.

. nudata Nut

Poin St. Tammany Parish: Lemaire 1013 (NO), first record for
Louisiana.

rosea Nutt.

S. CAROLINA: Aiken Co.: Massey & Massey 2983 (NCU, 2 sheets; GA),
Massey & Massey 3020 (NCU). These rather surprising collections ex-
tend the present range of C. rosea to S. Carolina.

_ tinctoria Nutt. var. tincto

FLORIDA: Dixie Co.: Gots 75270 (FSU, UARK), Kral 5407 (NO);
ylor GA).

Sumter Co.: Kral 6846 (GA); Ta > Co.: Lazor 3758
Note: All four of these collections from Florida have entirely yellow
ligules, suggesting contamination with C, leavenworthii T. & G. A speci-

men of C. leavenworthii T. & G. I observed recently from Hernando Co.

Florida (Kral 6795, GA) had the upper leaves pinnate and the achene
aristae rather short, suggesting contamination with C. tin ctoria var.
tinctoria. This northwestern area of the Florida penninsula includes the
type locality of ‘‘var. curtissii Sherff,” and is evidently an area of in-
tergradation between C. leavenworthii and C. tinctoria var. tinctoria,.

SIDA 7(3): 306. 1978.

307

Correction
On p. 148 of my summary paper, in line 8a, ‘“‘Bot. S. C. & Ga. 2:44.”’ should
read: Bot. S. C. & Ga. 2:441.—Edwin B. Smith, Department of Botany &
Bacteriology, University of Arkansas, Fayetteville, Arkansas, 72701.

CYPERUS SURINAMENSIS (CYPERACEAE): NEW TO ARKANSAS,
KANSAS, AND OKLAHOMA—Collections of Cyperus surinamensis Rottb.
in the United States are thus far known only from the Coastal Plain, ranging
from South Carolina to Florida, west to Texas. Additional collections can
now be reported from inland states of Arkansas, Kansas, and Oklahoma.
ARKANSAS. Jefferson Co.: growing in mud near water, on NW corner of
Boyd’s Island, Arkansas River, Pine Bluff, Locke, 31 Aug 1975 (UARK).
KANSAS. Harvey Co.: wet roadside ditch, sandy soil, 3 mi E and 2 mi N
of Burton, Stephens 19167, 11 Oct 1967 (SMU, UARK). Reno Co.: SW of
Hutchinson, Shanks, 27 May 1945 (SMU). OKLAHOMA, Choctaw Co.: above
water line of Roebuck Lake in sandy soil, Murphy, 8 Jul 1952 (SMU). Mar-
shall Co.: infrequent, margins of Lake Texoma, Univ. of Okla. Biol. Sta.,
Goodman 5761, 4 Aug 1953 (SMU). Texas collections are common from the
coast and inland to Robertson Co. but rare from north central Texas (Tar-
rant & Dallas Cos., ca. 400 miles inland). The Oklahoma collections are ca.
80 miles north of this; those in Kansas, ca. 300 miles. This puts Kansas col-
lections ca. 750 miles inland, somewhat out of the expected range of the
species. Specimens reported here were misidentified as C. acuminatus, a
species superficially resembling C. surinamensis—Barney Lipscomb, Her-
barium Southern Methodist University, Dallas, TX 75275.

SIDA 7(3): 307. 1978.

308

NOTICES

The Department of Botany, Smithsonian Institution, was recently asked by
the Soil Conservation Service of the United States Department of Agricul-
ture to produce an updated, revised, and corrected edition of the National
List of Scientific Plant Names, which was originally issued by the Service
in 1971. The Department of Botany has agreed to undertake the revision of
this checklist and looks forward to bringing the list to the attention of a
much wider circle of botanists and conservationists.

A special support staff has been assembled to work on this project for a
period of two years, after which the Soil Conservation Service and Smith-
sonian Institution will publish the results as an authoritative catalogue of
correct scientific names of all vascular plant taxa (including subspecies and
varieties) in North America, Hawaii, Puerto Rico and the Virgin Islands.

In the process of revising the list, intensive use will be made of the peri-
odieal literature and floras which have appeared in the intervening years
since 1971. A very significant aspect of this work will be our invitation to
the t axonomie community to participate in this endeavor. We will be solicit-

the highly valued cooperation of numerous botanical colleagues early
Aan

The need for a standard, efficiently produced, and more widely available
checklist of North American vascular plants has long been felt by all. The
opportunity now exists for filling this vacuum with a publication in a rea-
sonable amount of time. In the published revision, full acknowledgment will
be given to the people making individual contributions of data.

Please address your correspondence regarding the checklist to: The
Chairman, Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D. C. 20560

The magnifications given in the legend of the illustrations of Marsilea
mucronata (SIDA 7:219. 1977) should be changed to 10X and 1.25X.

SIDA 7 (3): 308, 1978

SIDA ‘csotany

VOLUME 7 NUMBER 4 DECEMBER 1978

CONTENTS

A synopsis of Fuirena (Cyperaceae) for the Americas north
of South America. Robert Kral. 309

A revised description of the salt marsh rush, Juncus
roemerianus, Lionel N. Eleuterius. Shays)

Additions to the vascular flora of Oklahoma.
R. John Taylor and Constance E. Taylor. 361

Taxonomy and geography of Nicolletia (Compositae:Tageteae).
John L. Strother. 369

Chromosome numbers in Erigeron and Conyza (Compositae).
Guy L. Nesom. 375

A synopsis of North American Corispermum (Chenopodiaceae).
Nita J. Maihle and Will H. Blackwell, Jr. 382

NOTES. Additions to the Texas flora. ie alee vascular plant records
Willd. (Compositae), 396—Additions to Alabama Lythraceae. 396—Nomen-
clatural correction in Clematis. 397

NOTICE 397

INDEX TO VOLUME 7. Barney L. Lipscomb. 398

US ISSN 0036-1488

SIDA, CONTRIBUTIONS TO BOTANY
Founded by Lloyd H. Shinners, 1962

Editor & Publisher
Wm, F. Mahler
SMU Herbarium

Dallas, Texas, 75275

Associate Editor Assistant Editor
John W. Thieret Barney L. Lipscomb
Northern Kentucky University SMU Herbarium
Highland Heights, Kentucky, 41076 Dallas, Texas, 75275

Contributors to Sida should refer to the latest issues for style, follow the
CBE Style Manual (8rd ed.), and use the ‘‘adopted”’ journal abbreviations

given in Botanico-Periodicum-Huntianum,

Subscriptions: Libraries—$6.00 (U.S.) per year; individuals—$8.00 (U.S.)

per volume; numbers issued twice a year.

Sida, Contributions to Botany, Volume 7, Number 4, pages 309-421.
Copyright 1978
by Wm. F. Mahler

A SYNOPSIS OF FUIRENA (CYPERACEAE) FOR
THE AMERICAS NORTH OF SOUTH AMERICA

ROBERT KRAL
Department of Biology, Vanderbilt University
Nashville, Tennessee 37235

Fuirena (Cyperaceae) comprises an estimated 30 species, mostly warm-
temperate or tropical, in both hemispheres. All are plants of high hydro-
period soils, most are heliophytes. Of the 13 taxa treated here one is a rare
ruderal, 6 are found mostly in grass-sedge formations such as found in
interdunal swales toward the seacoasts or in ‘‘sweet’’ marshes or seeps in-
Jand and the remainder are on highly organic sands or peat mucks of
swamps and bogs. Several are wetlands weeds. Only two are annual. No
economic value is known for the species.

Some cyperologists consider Fuirena to be a part of Scirpus. Whether
this is true or not is not belabored here; the plants are distinguishable from
other sedges through a combination of all or most of the following charac-
teristics: (1.) Culms generally leafy, the leaves with erect, scaly, ciliate
ligules and in 3 ranks; spikelets either in terminal clusters or in a compound
of sessile or pedunculate, both terminal and axillary, glomerules. (2.) Scales
of spikelets numerous, spirally arranged, awned, with the awn of many spe-
cies spreading or recurved, imparting a ‘‘bristly’ look to the spikelet. (3.)
Perianth usually of 6 bristles, 3 of which are broad-bladed, or all 6 lacking
blades, or of 3 bladed members with 3 reduced to nubs, or vestigial: most
of the species in this treatment have some bladed perianth members, but it
is interesting to see in the same small group of species a complete range of
reduction in perianth. (4.) Style deciduous, but tip of akene produced up-
ward into a definite beak whose apex is frequently tuberculate-papillose or
scabrid. (5.) Akene trigonous, usually stipitate.

The most current treatment of North American Fuirena is that done by
Svenson (1957). This is a very conservative approach based primarily on
perianth characteristics. My own work is preliminary to a biosystematic
study, and is intended merely to clarify some nomenclature, to present some
additional observations on the gross morphology and the ecology, and to
provide some additional distributional information not available to Svenson.

Acknowledgments
This preliminary study has been based primarily on field work stretched
over a period of more than 10 years and concentrated in the southern United
States and Mexico, During this period several hundred duplicate specimens
have been collected and these will be distributed at the close of the entire

SIDA 7 (4): 309—354.

310

study. Also during this period, and thanks to curators and staffs of the fol-
lowing named herbaria, more hundreds of specimens have been provided
through loans, namely from: C, CHARL, FSU, GA, GH, KANU, LAF, LSU,
_ NY, P, PH, POM, RSA, SMU, TENN, TEX, UC, US. Several spe-
cial collections made by Dr. Sidney McDaniel and Dr. D. Demaree have also
been examine
Grateful acineuledaient is here given to the following people: Dr. L.H.
Shinners and Dr. W. Mahler who obligingly made available several valuable
references and comments; to Dr. A. Skovsted and to Dr. A. Lourteig of C
and P respectively, who either provided basic type materials or took time
and trouble to make comparison with these: to all curators, herbarium
staffs of institutions mentioned above for their kindness not only in provision
of loans but for their patience in extending time periods of loans.
This investigation was supported in part by research grants GB-159,
GB-3255 and GB-6688X to the author from the National Science Foundation.

Morphology

Habit. Fuirena is made up of both annual and perennial species, but only
two of the taxa considered here are truly annual (F. pumila, F. simplex var.
aristulata). Of these the former invariably dies at the end of one growing
season, while the latter indicates at the southern portion of its range a
tendency to sprout from very contracted overwintering rosettes. Root sys-
tems of all species are fibrous and diffuse. Seed of all species tends to germ-
inate freely within two weeks of sowing in the greenhouse. Viability decreas-
es sharply after one year from harvesting.

Rootstock. The two essentially annual taxa are weakly to strongly caespi-
tose, the size of the clump and indeed the individual shoots are strongly
dependent on a combination of soil moisture and (probably) on organic con-
tent of substratum. These, as would be suspected, have the slenderest roots
and the shallowest root systems. The perennial species are all rhizomatous,
but the growth patterns of these are of two sorts. In F. scirpoidea, F. longa
F. repens and F. simplex var. simplex the rhizomes are of relatively con-
sistent width, are sheathed by short, scaly leaves, these at well separated
or approximate nodes, in the case of the latter therefore imbricated. In the
remaining perennials the rhizomes produce at approximate or distant inter-
vals contracted ovoid, or nearly round cormous offshoots. These are pro-
gressively formed as the rhizome elongates and are particularly noticeable
and full toward fall and in winter. Actually the whole structure is contract-
ed culm plus an apex with very contracted preformed shoot leaves and all
stages of formation and expansion are to be found on a single vigorous plant
at any stage of the growing season. In the ‘“‘bud”’ or ‘‘cormose’”’ stage only
the outer leaf is visible with the culm base, distended culm, the sheath and
its minute blade enfolding the shoot apex. Through this pushes the rapidly
elongating culm as the shoot elongates.

=~

311

Culm. The culm of Fuirena is slenderest toward its base with the inter-
nodes contracted and sheathed. Its habit may be erect or quite lax and
leaning. It is solid at the nodes, internodes terete or obtusely trigonous,
progressively elongating upward, green, strongly and numerously longi-
tudinally ribbed, with the vascular tissue peripheral and the large central
area spongy-lacunar. Stomatal areas are confined to the narrow intervals
between the ribs.

Leaves. The leaves are in 3 ranks and are comprised of a loose to rather
tight cylindrical or 3-angled sheath, this multiribbed as in the culm, and a
spreading to ascending linear or lance-linear, acute or narrowly tapering,
blade. The sheaths are closed except toward the orifice, where there is pro-
duced an erect, usually pale brown, scarious, ciliate scale, the ligule. The
upper and lower surfaces of the leaf blades are usually markedly different.
The upper surface is as a rule smoother (though in some species it may still
be quite trichomiferous and also minutely scabrid or papillose) with numerous
longitudinal nerves moderately to slightly impressed, marked usually by a
narrow and overlying band. Between the nerves are several rows of short to
long-rectangular, longitudinally oriented, epidermal cells, these with side and
end walls variously papillose, sometimes with end walls produced upward
into dome-shaped processes; cell contents are comparatively translucent or
made opaque with mucaceous or farinose material; stomata are not in evi-
dence. If trichomes are present these may be short, unicellular or elongate-
tapering, multicellular, glassy like hollow icicles: they may appear over the
lumen or toward the endwalls. The lower leaf blade surface appears as an
extension of the sheath surface, hence is strongly multicostate, the costal
area comprised of shorter, glassy-walled cells, these often drawn out into
short or elongated trichomes; cells of the intercostal areas are longer and
thinner-walled, are longitudinally oriented in rows, these interrupted at more
or less regular intervals by guard cell pairs and subsidiary cells, these like-
wise vertically oriented and appearing singly or in short transverse rows,
slightly sunken or raised depending on the species. The lower surface of the
leaf, in trichomiferous species, is usually hairier than the upper with the
longer hairs again glassy and mostly found arising from costal cells.

Inflorescence. Spikelets in all species are comprised of very many ascending
or erect, rather loosely imbricate, cymbiform scales, all but the lowest 2—3
bearing florets in their axils. The lowermost sterile bracts show transition to
uppermost inflorescence bractlets, thus are more leaf-like and longer-tipped.
The fertile bracts are rather uniform through the genus, tending to conceal
all the floret except the tips of filaments and the anthers at anthesis together
with the 3 hispidulous long-linear stigmas; most fertile bracts are of a broadly
obovate, oblong to ovate type, broadly scarious margined, the backs rounded,
with strongly raised nerves medially which converge to form a stiffish, promi-
nent, erect to spreading or recurved mucro. Bract apices are rounded or
slightly retuse, scarious, continuous across the mucro base adaxially (see

312

plates) and thus in position homologous to ligule. Bract margins and backs
are in many species stiff-hairy, the hairs of various lengths but all straight
and sharp. Common spikelet shapes are ovoid, lance-ovoid or cylindrical.
Sequence of flowering within the spikelet is acropetal. When the fruit is ma-
ture the lowermost scales begin to absciss, exposing raised, ‘“‘V’’ shaped
scars, Old spikelets toward the end of a season therefore show a considerable
length of rachis, in that flowering is indeterminate, but (rather interestingly)
the spikelet tends to retain the same overall shape and dimensions, While ex-
tremes of F. scirpoidea may produce but a single spikelet at the culm tip,
most produce them either in a terminal cluster or this plus pedunculate clus-
ters from 1 or more of the subtending nodes. Arrangement of spikelets is
basically cymose, with central spikelets of a group or unit maturing first. The
smallest unit or cluster within the inflorescence is here termed a ‘“‘glomerule.”
A single glomerule is subtended by 1 or more variously awned bracts the
bodies of which are not much different than those of the spikelet; it is made
up of few to many sessile or subsessile spikelets. The glomerule may be soli-
tary or may form a compound with others in a diffuse system, usually with
the primary axis or axis branch terminating in a sessile glomerule, this ex-
ceeded by pedunculate ones on peduncles of varying lengths. Each peduncle
is sheathed at its base by a thin-tubular, oblique-orificed ‘‘prophyll’’ which
by position is actually a ligule in that it is located at the inside of the junction
between bract sheath and bract blade

Flowers. The flower of Fuirena is perfect, sessile in the axil of a single
bract (see plate for F. breviseta, fig. 4). A perianth is present and is basically
biseriate. The commonest situation in our species is for the outer set of
3 to be simple bristles, these smooth or retrorsely barbellate, while the inner
set of 3 bear broadish blades. In F. incompleta there are no bladed members,
only 6 simple bristles or (in var. obliterata) the bristles very reduced or ab-
sent. In F. camptrotricha, F. umbellata, F. repens only 3 members develop
prominently and these bear blades; the other set of 3 is reduced to short,
tubercle-like processes or is absent. The most unusual perianth situation is
displayed by the introduced F. wallichiana, whose perianth has a combination
of short unbranched bristles and longer, flatter sparingly pinnately-branched
segments.

The stamens range from 1 in the annual species to 3 in the perennials, the
hypogynous, long-linear, flattened filaments at anthesis projecting the linear
or oblong, basifixed, 2-locular anthers beyond the bract tips. The pollen is in
cryptotetrads. Anthers are smallest in the annuals.

The ovary is tricarpellate, sessile or stipitate, the beaked apex prominent,
jointed to an elongate smoothish style, this terminating in 3, exserted, long-
linear, hispidulous stigma lobes.

The mode of flowering is similar to that observed for Fimbristylis, Bulbo-
stylis, Abildgaardia. In a given flower the stamens are later to shed pollen
than are the stigma lobes to receive. In a given spikelet, usually in a given

313

inflorescence, a series of spirals of florets will either be presenting or receiv-
ing pollen but not doing both simultaneously. The opportunity for selfing ap-
pears to be remote.

Fruit. The fruit is rather uniform through all our species, being a more or
less stipitate akene, whose apex is a prominent linear or narrowly triangular
beak (this sometimes papillose), whose body is broadly elliptic and trigonous
with prominent rounded angles. The surfaces of the akene body between the
wirelike angles are usually glassy, sometimes very delicately striated and
cross-striated, only in F’. wallichiana being cancellate.

Cytology

Some preliminary studies have been made of microsporogenesis in a few
species, sufficient to convince this writer that, at least in North America the
mode of spore production is uniformly cryptosporic. Microsporogenesis pro-
ceeds as has been observed in Fimbristylis and related genera (Tanaka, 1939,
1941; Kral, 1971). The nucleus of the microspore mother cell meiotically di-
vides and a tetrad is formed. Though low sutures appear to develop in the
mother cell wall, cytokinesis does not take place. Instead, 3 of the 4 haploid
nuclei move to one end of the sporocyte, lose both size and distinctness,
while the fourth nucleus divides mitotically. Of these two products, one loses
size and joins the other 3 reduced genomes. The remaining, or fifth, nucleus is
thus the only functional product. (See illustrations of the various stages.)

My own counts are still rather limited, and more will be published later
on the subject as additional chromosomal studies are made. They now reveal
a haploid number of 23 for F. breviseta, F. longa, F. pumila, F. scirpoidea, F.
squarrosa, and F. camptotricha (mistakenly reported as F. wmbellata in a
prior, 1971, paper! ). This count is so unusual and so unexpected that one won-
ders at the derivation of this condition; certainly it needs to be examined
further. My limited inspection of chromosomes of F. simplex var. simplex
has revealed two conditions for what appear to be phenotypically very similar
sorts. In Texan material of the variety I have a haploid count of 15; in ma-
terial of the same variety from the Gulf Coast of Mexico, the ‘“‘conventional”’
count of 23 was made. As stated earlier, this information is so peculiar and
conflicting as to warrant a more ample study.

Pollination. The flowers of Fuirena appear to be, with their exserted his-
pidulous stigma lobes and anthers, anemophilous. However, I was able to
make one observation of what appeared to be bee pollination in a large pop-
ulation of Fuirena scirpoidea near Port St. Joe, Gulf County, Florida in July
1958. The bee was the common honeybee, and there was no question of abund-
ance and persistence of these bees on and around the spikelets. It should
probably be assumed that the visits were for pollen.

Hybrids. Some coastal plain habitats have up to 4 species of Fuirena grow-
ing together or in proximity. Yet the differences between the taxa, however,
fine, appear to be consistent over the range. Such variation as occurs is also

314

a”
a
w® é
ler
a
Om a an
8 : a
4
e
,er
°e
e
e q
Pe ° + se , 6 e
“ee ¢ e 6 tee,
a 2 «& a
© “ e.
Pr _* e ‘. . *e
*, g or
aa, ces ,

MICROSPOROGENESIS IN FUIRENA. 1. F. pumila (Kral 22886, third

division). 2. F. pumila (Kral 22411, prophase 1). 3. F. breviseta (Kral 22905,
third division), 4. F. breviseta (Kral 22887, prophase 1). 5. F. scirpoidea
(Kral 22961, prophase 1). 6. F. scirpoidea (Kral 23005, prophase 1). 7. F,
longa (Kral 22962, third division. 8. F. longa (Kral 23004, third division).
9. F. squarrosa (Kral 20668, prophase 2). 10. F. mal ae (Kral 24997, pro-

. simplex (Kral 27810, third division). F. longa (Kral
22962, prophase 1). 138. F. scirpoidea (Kral 22961, eres se l).

2°
[S*
fy
oO
=
ny

315

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@ ee . &
as ae | 7 . 7
Ps "5 Bre: 6 Py ‘
8
i z ¢
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consistent over the range. Further study is desirable in the case of F. longa,
which has a morphology strikingly intermediate between that of F. breviseta
and F’. scirpoidea and may be of comparatively recent hybrid origin.

Taxonomic Treatment

In the treatment which follows, keys and descriptions are based on material
collected from within the area of study, namely America north of South Amer-
ica. Generic limits are drawn only in the sense of the species treated. Keys
and descriptions, while based in part on vegetative characteristics, are still
centered on the characteristics of fruiting spikelets and healthy plants. Di-
mensions of spikelets are drawn only from the scale-bearing portion of rachis
from which the scales have fallen, and this is not included in the measure-
ment. Species are numbered in a single continuous sequence,

316

The large number of specimens examined in the course of the project makes
it necessary to cite these collections separately. A list of citations is being
prepared and, hopefully, will be available upon request to the writer at the
time this paper appears

The illustrations are numbered to correspond with the text treatment. Each
plate shows a whole plant reduced together with enlarged views of florets,
fertile bracts, nodes showing inner surfaces of lower parts of leaf blades. All
illustrations are by the author.

FUIRENA, Rottb., Descr. & Ic. 70. 1773.
Vaginaria Pers., Syn. Pl. 1:70. 1805.

Perennial or annual, usually pubescent, caespitose or rhizomatous herbs,
if rhizomatous with rhizomes short and thickened or extensively creeping and
forking or producing in addition axillary cormous offshoots from which culms
arise. Culms erect or leaning to 1 m tall, simple, stiff or wand-like. Sheaths
of leaves loosely or closely investing the culm, strongly costate; blades either
reduced to scale-like appendages or of a linear type, stiffly spreading or erect.
Spikelets usually 1 cm or more long, ovoid, short-cylindrical, or ellipsoidal,
arranged singly at culm apices or in terminal and axillary clusters, these
made up of 1-several, sessile or variously pedunculate glomerules, or (less
usual) the spikelets more pinnately arranged. Scales of spikelets numerous,
spirally arranged, lowest 1—3 barren, the fertile mostly ovate, oblong or obo-
vate, thinnish, the backs rounded, usually hairy, medially strongly to faintly
costate, costae merging subapically to form a conspicuous, erect, or spread-
ing stiffish, usually scabrid, awn. Perianth of most species 6-parted, with 3
simple bristles forming an outer whorl and 3 stipitate-bladed members form-
ing an inner whorl; in some species the perianth is reduced to 3 bladed mem-
bers, or comprised only of bladeless members or (rarely) lacking. Stamens
(1-) 2—3 (-6), the filaments elongate, flattish, the anthers basifixed, oblong or
linear, 2-locular, the pollen in ‘‘cryptotetrads’’. Style 3-cleft, disarticulating

ate gynoecial beak, stigma lobes linear, papillate-scabrid.
oe eas to prominently stipitate, usually strongly 3-angled, body in
outline rhombic or broadly ellipsoidal, the angles prominent, wirelike, the
faces flat or slightly convex or concave, smooth or faintly cross-lined or can-
cellate, the apex a prominent narrowly triangular or linear beak, this some-
times expanded distally and there scabrid or papillose.

Key to Fuirena of the New World north of South America

1. Perianth in part or solely of eas bearing broad blades; akene surface
lustrous, smooth or very finely stri
2. Perianth comprised both of bladed and bladeless members, daa
bristles reaching at least to the middle of the stipe of the a
3. The Pee nue — producing corm-like sie. coer an-
thers at least
4. Culms oe wan Se smooth, on elongate, creeping, e-cormous
rhizomes; mucro of fertile scales erect, mostly less than 14% of the

317

length of the scale body; sheaths of leaves either essentially blade-
less or with longest blades rarely re aching 5 cm; backs of fertile

». Blades of upper culm leaves short, mostly mucro- like: apices of
perianth scales mostl acute; spikelets mostly ovoid and 1—5,
sessile in a terminal glomerule, with subtending bract shorter
than the spikelets 1. F. scirpoidea Michx.,

. Blades of upper culm leaves longer, ae plane, to 5 cm long:
apices of perianth scales acuminate; spikelets mostly lance-ovoid
or lance-cylindric, narrowly acute; spikelets sessile in a terminal
glomerule and frequently with pedunculate clusters as well; sub-
tending bract usually longer than the spikelets.

F. longa Chapm., p. 319.

. Culms usually stouter, usually with at least some ee oe long
hairs on ves eee and/or sheaths, on elongate to short, cormous
rhizomes; cro of fertile bracts erect or spreading-recurved; leaf
blades ole ee sca: the longer ones (at least on healthy plants)
rarely as short as 5 cm; backs of fertile scales usually with only 3
prominent medial nerves.

6. Mucro of fertile bracts at least 24 as long as the scale ee

pean spreading or recurved-tipped; rhizomes cormous
corm

Or

7. Se producing corm-like shoot eee _perianth scale blades

with apex obtusely angled, acute, acu te or narrowly in-
curved-conic, not bearing a subapical apis or bristle.
8. Perianth bristles (sepals) short, incurved, th, their tips

00
not reaching the bases of the perianth blade. (petals); lower-
most sheaths of leaves hispid, those of mid-culm or above be-

coming smooth; apex of is blades ABicesneled or
acute, not ee or acumin
. breviseta “(Coville) Coville in Harper, p. 324.
2 Perianth eee longer, more erect or even slightly spread-
ing, smooth or retrorsely barbellate, their tips reaching at
to the bases of perianth blades: all sheaths of leaves usually
hispid or hispidulous or hirsute; “ape x of perianth blades either
conic and very tumid, or acuminate.
9. Petal blades with an eer thinnish or thickened,
curved apex; perianth bristles retrorsely barbellate; an-

Co

Coastal Plain and inland to adjacent provinces east of the
Mississippi River, with the cormous rhizome buds usually
close-set. 3. F’. squarrosa Michx 7.
. Petal blades cumid at ete urity, narrowing distally to be
conic and erect, sometimes apiculate as well; perianth bris-
tles smooth; anthe ers ca. 2 mm long; plants of the Gulf

va)

with cormous parts of the rhizome often separated by dis
tinct Giewais of narrow see longer than the corm
width. . PF. bushii Kral, p. 331.
7. Rhizomes simple, iach: corm- aie ee buds; perianth scale
blades with apex flattish, or tumid, conic, blunt or emarginate,
but usually producing a short or elongate, subapical dorsal
bristle. - 1 . . . . 7. FL simplex Vahl, p. 332.

6. Mucro of fertile scales erect, seldom as long as % the length
of the scale body; rhizomes cormous; robust plants from southern
Mexico southward; also Cuba.

10. Broader blades of leaves usually at least 1.5 cm wide, these
and the sheaths mostly smooth except for stout-based spread-

acuminate, usually incurved, lacking a short, incurved, sub-
apical dorsal bristle. 3. F. robusta Kunth, p. 322,
Broader blades of leaves ‘usually | less than 1.5 cm wide,
a the sheaths usually hairy; apex of perianth blades usually
mid, generally blunter, and bearing a short, incurved, sub-
ee dorsal bristle
=. 4 _ extremes of I’. camptotricha C. Wr., p. 342.
3. The plants ene usually annual; anthers mostly 0.5—0.6
(-0.7) mm long.
11. Apex of Sar orith scale obtuse to retuse, bearing a dorsal- subapical
seals e, this retrorsely barbellate; plants mostly of the prairie
provi

—
com]

7a. F. simplex Vahl var. aristulata (Torr.) Kral, p. 336.

11. Apex of perianth scale acuminate, usually very narrowly so, usu-

ally incurved, not bearing a subapical bristle; plants mostly of the

Atlantic and Gulf Coastal Plain with outliers in the Lake States.

8. F. pumila (Torr.) Spreng., p. 338.

2. The perianth comprised only of bladed members (very reduced, tubercle

or papilla-like bristles may sometimes be located in cycles inward or
outward from the bladed members! ).

12. Claw of petal blades crimped; robust plants with stout cormous rhi-
zomes and culms at least 0.5 ecm thick; inflorescence often diffuse,
comprised of many glomerules of spikelets.

13. Mature petal blades flattish or thickened only marginally, over the
nerves, and apically, the apex generally retuse-notched, from this
notch arising a slender, coiled bristle.

F. umbellata Rottb., p. 340.
; a petal blades with distal half tumid, inflated, the apex broad-
ly conic or obtuse, papillose-scabrid, the bristle, if present, shorter,

Saal subapical, erect or incurved.

—
ow

10. F. camptotricha C.Wr., p. 342.
12. Claw of petal blades. straight; slender lowish plants from ‘slender,
diffuse-c ener e-cormous, elongate rhizomes, the culms never as
thick a orescence usually a terminal cluster of sessile or
ic srikelet 11. F. repens Boeckler, p. 346.
1. Perianth comprising pene bristles: or with some producing narrow, lineal
or lineal-lobed blades, or perianth essentially absent.
14. Akene surface smoothish; perianth bristles simple or lacking.
15. Perianth bristles usually 6, subequal, most of them well exceeding
e middle of the akene body
l2a. F. incompleta Nees var. incompleta, p. 348.
15. Perianth bristles absent or nase to short, unequal lengths, rarely
exceeding the middle of the aken y.
2b. F. ep pet a var. obliterata Kral, p. 350.
14. Akene ‘euace. strongly cancellate; some all of the ao bristles
bearing narrow, a pinnate or linear -lobed
13. F. batlichians Kunth, p. 390.

319

1. FUIRENA SCIRPOIDEA Michx., Fl. Bor. Am. 1:38. 1803.
Vaginaria richardi Pers., Syn. Pl. 1:70. 1805.
Scirpus scirpoideus (Michx.) T.Koyama, Journ. Fac. Sci. Univ. Tokyo
IIT (7):287. 1958.

Essentially glabrous perennial, 2—6 dm tall, from an elongate, often fork-
ing system of rhizomes just beneath the substrate surface, these pale, stout-
ish, with darker brown oblong scales overlapping the lower part of each in-
ternode. Culms erect, wand-like, arising like parts of a picket fence, usually
very many and rather approximate along the creeping rhizome; internodes
rather short at the culm base, elongating toward the inflorescence, subterete,
multicostate, nodes swollen. Leaves mostly sheath, particularly those prox-
imal on the culm, the sheaths loosely investing the internode, multicostate,
tubular, firm, almost bladeless, producing at the oblique orifice a shor
(rarely longer than 3—4 mm) incrassate-margined, cusplike blade and an
erect, scarious entire or short-ciliate, brownish oblique ligular orifice.
Sheaths becoming shorter, internodes progressively longer upward, the ulti-
mate one terminating in a single spike or a tight cluster of 2—5, subtended
by a short-bladed bract shorter than the spikelets. Spikelets ovoid to lance
ovoid, mostly 7—10 mm long (—1.5 cm), blunt or (rarely) acute. Fertile
scales mostly ovate, oblong or obovate, the body 2.5—3.5 mm long, ciliate,
thin, mostly tan, greenish-brown or reddish-brown, the backs rounded, pu-
berulent or hirsutulous, with at least 5 strong median nerves, 3 convergent
to form a short, stiff, erect mucro mostly less than 1% as long as the scale
body and scabrid. Florets mostly 1.5—2.0 mm long, perianth 6-parted, calyx
bristles retrorsely barbellate, reaching to the tips of the petal claws or
slightly beyond; petal blades mostly ovate, about as long as the claws,
thickened distally but somewhat compressed between the 3, raised basal
nerves, apex usually compressed-conic, apiculate d also often slightly
scabrid. Anthers ca. 2 mm long. Akene slender-stipitate, body trigonous
with the angles pale, wirelike and the faces flat to slightly convex, a deep,
lustrous reddish-brown or castaneous; stylar apiculus distally papillate or
minutely scabrid. Fig. 1

Sandy or sandy-peaty marshes, swales and seeps, sometimes in slightly
brackish situations, along the seacoast from N.C. south to southern penin-
sular Florida and west in the Gulf Coastal Plain to southern Texas: Cuba.
Map 1

Type: ‘“Florida.”’ Michaux. Holotype at P compared with material sent
from VDB. Dr. Alicia Lourteig, who made the comparison, found a good
match.

2. FUIRENA LONGA Chapm., FI. S. US. ed. 3:541. 1897.

Similar to F. scirpoidea but with rhizomes shorter, more branched and
with culms frequently appearing tufted. Sheaths of mid and upper culms
producing flat, short-linear blades, these longest at mid-culm or above,

320

Fig. 1. F. scirpoidea (Kral 32626).

321

¢ SYKA HLOG
© LLYTLLSIAV YA
XFTAWIS AYA

© YINOT FS

© YICIOAYIIS F

Map 1.

ulent. Culms

Spikelet clust
ly as long as to longer than

mostly cylindric-lanceolate or lance

there 1.5—5.0 cm long, smooth or with the upper surface puber

at level of inflorescence frequently hispidulous.
rules) often 2, subtended by bracts from near

the spikelets. Spikelets narrow

ers (glome-

?

ovoid
mostly

nerved medially as in F. scirpoidea
sometimes slightly more than \% the length

’

’

acute, more greenish. Fertile scales with body similar in length

obovate, thin, hirtellous or puberulent

but the excurrent mucro longer

’

3

322

of the scale body, Perianth essentially as in F. scirpoidea but with petal
blades often with a longer, narrower point. Anthers 1.3—1.5 mm long. Fruit
as in F’. scirpoidea. Fig. 2.

In habitats similar to those of F. scirpoidea and frequently mixed with it,
in Florida and west along the coast to southern Mississippi; southern Texas.
Map 1.

Type: Florida. Franklin Co.: ‘‘Apalachicola, A. W. Chapman’, GH! MO!

This, so often placed in synonymy of F. scirpoidea, has a very different
look in the field. I have seen much of it in peninsular Florida and in the
Gulf Coastal Plain west to southern Alabama. When seen mixed with F.
scirpoidea it stands out at once because of its more appropriate shoots on
shorter, more branching rhizomes, its longer leaf and bract blades, its
narrower, more bristly, greener spikelets which appear in larger, more
numerous clusters on the culms. I have yet to see it where there was not
also some F’. breviseta nearby and suspect without proof as yet that this
entity has probably arisen as a hybrid between F’. scirpoidea and F’.. bre-
viseta or a similar species such as F. squarrosa. Certainly it is a vegeta-
tive intermediate, Tending to support this notion is the fact that but one
locality for F. longa is presently known for Texas (Aransas County), dis-
junct from the furthest west Mississippi county by several hundred miles.
From this same locality have also been collected F. scirpoidea and F.
breviseta, these also representing considerable extensions of known range
southwest.

3. FUIRENA ROBUSTA Kunth, Enum. PI. 2:185. 1837.
F. bahiensis Lindl. & Nees; Nees in Mart. Fl. Bras, 2(1):108. 1842

Brasi
* latifolia Steud. Syn. Cyp. 126. 1855 oo Brasil).
: schizophylla C. Wright in Sauv. Anal. Acad. Ci. Habana 8:32. 1871.

uelta de Abajo, Cuba)

Perennial, with thick, spongy, erect culms to 16 dm tall, caespitose or

leaves shortest, sheathing with oblique, ciliate orifices or with short-tri-
angular blades; blades of mid-culm longest, oblong-linear or lance-linear,
to 30 cm long and 3 cm wide, usually tapering from a cordate-clasping base
to a narrowly acute apex, margins at least proximally with upward ap-
pressed, stiff, long trichomes, otherwise tuberculate-scabrid or entire, the
surfaces smooth; sheaths smooth or rarely glabrescent. Inflorescence of
3—6 axillary systems of pedunculate, cymose clusters of glomerules from
the upper nodes, the system interruptedly cylindrical, primary peduncles
hirsutulous, triangular. Spikelets ovoid or lance-ovoid, ca. 5—6 mm long,
reddish-brown or greenish-reddish-brown. Fertile scales ovate or obovate,
rounded-ciliate, ca. 3 mm long, including a rigid, erect cusp shorter than
the scale body, the backs with 3, rarely 5, strong medial nerves, these
converging into an erect cusp usually less than 4% as long as the scale

323

Fig. 2. F. longa (Kral 23139).

324

body which is overall glabrous to appressed-puberulent or strigillose.
Perianth of 6 members, the outer set of 3 simple bristles, these about as
long as the akene body or slightly longer and smoothish, the inner 3
(petals) bladed, clawed, ca. 2 mm long, the blades thickened, oblong or
lance-oblong, obscurely 3-nerved, distally tumid, the apices incurved-
apiculate or mucronate, Anthers normally 3, ca. 1.5 mm long. Akene ca.
1.5 mm long, stipitate-trigonous, body with angles prominent, wirelike,
faces flat or slightly convex, smooth, lustrous, pale brown. Fig. 3.

Swamps and bogs in lowlands in Cuba, Panama; South America. Map 4.

Type: Type locality, southern Brasil. Material not examined by this
writer. The description rendered by Nees in Martius (1842) of F. bahiensis,
which is this species, is very good and matches in all regards. The type
description by Kunth, based on material collected by Sellow, is brief but
adequate to separate this from F. umbellata with which it might be
confused,

F. robusta differs from F. umbellata mainly as follows: (1.) It is smoother
than the latter, the sheath and leaf surfaces (very often even the in
florescence) smooth; in F. umbellata the same surfaces are often anenuiene
more often hispidulous or hirsute. (2.) The margins of leaves, at least to-
ward the base, have ascending or more often appressed, long, stiff hairs;
in F. umbellata hairs of this area, if long, are less thick and are spreading.
(3.) The spikelets are larger, wider, smoother. (4.) There is but 1 set of

a slender, coiled mucro; in F. robusta both sets of perianth are present,
with stipes of bladed members mostly straight and with the usually swollen
blade tips terminating acuminately in an incurved, not coiled, awn.

4. FUIRENA BREVISETA (Coville) Coville in Harper, Bull. Torrey Bot.
ub 28:466. 1
F. squarrosa Michx. var. breviseta Coville, Bull. Torrey Bot. Club 17:6.
1890.

Perennial from scaly rhizomes, the shoots arising from axillary, cormous
offshoots. Culms to 1 m tall, usually 5 dm or lower, tufted, or approximate
along the creeping, superficial rhizome, erect to ascending or leaning, inter-
nodes terete or something slightly angulate, multicostate, the lowest shortest,
the uppermost longest, usually smooth except toward and in the inflores-
cence, where hispidulous. Leaf sheaths loosely cylindrical, eo ribbed,

culm. Ligular orifice short-cylindric, submembranaceous, oblique, short-
ciliate. Largest leaves at mid-culm or above, the blades spreading to re-
flected or erect, broadly lineal or linear-lanceolate, 5—15 cm long, 3—10 mm
broad, flat, usually gradually narrowing from near the cordate-clasping
base to the attenuated apex, margin slightly thickened-cartilaginous, usually

325

Fig. 3. F. robusta (Bailey 256).

326

liate at least proximally, the upper and lower surfaces

ispid-ci

spreading-h

smooth to sparsely strigose or strigillose or puberulent (usually nearly

x

smooth), more strongly nerved beneath. Inflorescence a terminal dense to

© YUWid 7

© KLISITAG FS

Map 2.

327

open system of sessile and pedunculate glomerules of spikelets, primary
and secondary peduncles angulate, hispidulous, often subtended by 1—3
additional ascending-pedunculate clusters of glomerules, all glomerules
usually from 3—5 or more, digitately spreading spikelets. Spikelets ovoid,
lance-ovoid, or cylindrical, mostly 1.0—1.5 (—2.0) em long. Fertile scale
body mostly oblong to obovate, 3.0—3.5 mm long, rounded to slightly emargi-
nate, ciliate, thin, greenish brown or reddish-brown, backs rounded, hir-
sutulous or minutely scabrid, with usually 3, strongly raised, tan or greenish
nerves medially, these converging to a stiffish, slender and spreading-
recurved mucro ca. as long as the scale body and scabrid or hispidulous.
Perianth of 6 members, the sepal bristles usually smooth, stoutish, incurved,
short, rarely as long as the claw of the petal, rarely longer than the stipe
of the akene. Petals ca. 2 mm. long, blade long-clawed, ovate or short -
oblong, thickened except for the base, where showing 3 raised nerves,
apically often incurved, broadly acute to obtuse, sometimes apiculate,
usually minutely tuberculate-scabrid. Anthers 1.0—1.3 long. Fruit trigonous-
Stipitate, body with the three angles pale, wirelike, the faces flattish or
slightly convex, a deep, lustrous brown, the stylar apiculus slightly broad-
ened and hispidulous distally. Fig. 4.

ogs, wet sandy places, seeps, savanna ditches, mostly Coastal Plain
from A a Virginia south through Florida and west, in the Gulf
Coastal Plain to southern Texas; Cuba. Map 2.

Lectotype: Florida. Duval Co.: low grounds, A. H. Curtiss 3068. DUKE!
F! GH! MIN! NY! PH! TENN! A large set soon to be distributed and which
compares well with the type is as follows: Alabama: Baldwin Co.: peaty
pockets in sandy slash pine flatwoods just E of Orange Beach by Ala. 180,
22 Oct 1969, R. Kral 38239.

This species is placed by many authors in synonymy of F. squarrosa,
species admittedly very similar in habit. However, as Coville (1890) long
ago recognized, it may be sorted easily from amongst a mass of mixed
herbarium material of both, on the basis of its distinctive perianth char-
acters, namely the long-stipitate, bluntish petal blades (in contrast to those

en m
Eieuished in the field by its stiffer, smoother foliage. While in F. squarrosa
the leaf sheaths are pilose-hispid from base to apex of the culm, those of
F. breviseta have only the lowermost sheaths hispid, the upper are almost
invariably smooth. (In perhaps 1 instance in 100 the upper sheaths may be
hispid!) The exceptions may represent chance hybrids, but even these
show the perianth character of F. breviseta, and aré so annotated.

5. FUIRENA SQUARROSA Michx., Fl. Bor. Am. 1:37. 1803.
F. squarrosa var. hispida Chapm., Fl. S. US. 514. 1860.

Perennial from scaly rhizomes, shoots arising from axillary, cormous

328

ta (Kral 38239).

Fig. 4. F. brevise

Fig. 5. F. squarrosa (Kral 23225).

329

330

offshoots. Culms to 1 m tall, usually lower, erect to ascending or leaning
on other plants, culms strongly multicostate, smooth except in the inflore-
cence. Lowest leaves scaly, mostly sheath, blades progressively longer as
mid-culm is approached, then gradually diminishing in size into the in-
florescence; sheaths cylindrical or gradually inflated distally, strongly
multicostate, hispid-hirsute, sometimes also hirsutulous (in a few extremes
with uppermost sheaths or a culm nearly smooth); leaf blades spreading
to erect, linear-attenuate, the larger mostly 8—20 cm long, 4—10 mm broad,
mostly tapering from near the clasping base to the apex, strongly margined,
upper surface scabro-puberulent and finely nerved, lower surface puberulent
and hispid-hirsute, strongly raised-nerved, the margin spreading hispid-
hirsute; ligule short-cylindric, ciliate, oblique, scarious. Systems of glo-
merules in a rather dense terminal cluster, often also from the next lower
1—2 nodes on stiffly erect or ascending, angulate, hirsute or hirsutulous
penduncles. Glomerules subcapitate, spikelets crowded, digitately spread-
ing, the clusters subtended by leaf-like bracts, 1 usually longer than
cluster subtended. Spikes ovoid to cylindric-lance-ovoid, to 2 cm _ long.
Fertile scales obovate, 2.5—3.5 mm long, rounded or retuse, ciliate, with
reddish or greenish-brown sides and a pale green or tan midzone, longi-
tudinal ribs 3 (—5) medially, the central strongest, all merging into a
scabro-hispidulous, spreading-recurved mucro more than 44 as long as the
scale body. Florets 2.0—3.5 mm long, perianth 6-parted, calyx bristles
reaching at least to the base, often to the middle of the petal blades,
antrorsely barbellate. Corolla scales long-stipitate, claws slightly shorter
than to nearly as long as the blades, blades narrowly to broadly ovate,
usually incurved-acuminate, frequently minutely prickly apically on the
inner face, bases cordate, truncate, broadly cuneate or abruptly attenuate,
dorsal surface with a strongly raised longitudinal rib continuous with the
scale apex and with at least 1 pair of spreading lateral nerves; scale body
either of a consistent thickness or distally tumid. Anthers 3, ca. 1.0 mm
long. Stipe of akene shorter than the petal claws; akene with 3 edges pale,
wirelike, faces lustrous, deep brown or castaneous, faintly cross-lined.
Fig. 5.

Open bogs, shores, ditches, seeps and wet peaty clearings, usually on
substrates derived from sands, sandy alluvium, siliceous rock or shales,
U.S.A. from Long Island southward in the Coastal Plain to peninsular
Florida and west into eastern Texas; inland throughout the Piedmont and
contiguous portions of the southern Appalachians and into the Mississippi
embayment north to southern Arkansas and western Tennessee. Map 2.

Type: ‘“Georgia’: Presumably collected by A. Michaux. Holotype: P,
material from Florida, Mississippi and Louisiana compared with holotype
by Dr. Lourteig matches well.

To the north of its range in the Atlantic Coastal Plain are forms which
are quite tumid in the distal 1/2—1/3 of the petal blade. This characteristic
shows up less frequently southward in the Coastal Plain and inland, is quite

331

rare even in southern Alabama populations, and is not found west, where
all F. squarrosa populations have flatter, frequently more slender petal
blades. The elongate, retrorsely barbellate calyx bristles appear to be quite
constant in character throughout the range of the species. This same taxon
is sometimes parametered so as to include F. pumila, which is annual, has
anthers but 1/2 as long as those of F. squarrosa, and which has tumid,
slenderly pointed petal blades as well as much longer calyx bristles. Neither
is this to be confused with F. bushii (F. ciliata B. F. Bush), a southwestern
Coastal Plain species which is often taller, has anthers about twice as long
as those of F. squarrosa, and which has a characteristically longer-ciliate
leaf margin together with conspicuously longer awns of the fertile scales.
The petal blades of this last are at maturity much more swollen even than
the N.J. extremes of F. squarrosa.

6. FUIRENA bushii Kral, nom. nov.
IF’, ciliata Bush, Mo. Bot. Gard. pee 16:91. 1905.
not ea ex Steud., Syn. Pl. 2:126. 1855.

Rhizome elongate, slender, with rather distant cormous offshoots. Culms
1-several, sometimes appearing tufted, erect to leaning on other vegetation,
to 10 dm tall, subterete to angled, multicostate, proximally smoothish,
distally hirsute. Lowest leaves scale-like, mostly sheath; medial culm-
leaves largest, sheaths mostly overlapping on the culms, strongly hirsute,
multicostate, cylindrical, rather loosely investing the culm, ligule scarious,
reddish-brown, erect, oblique, ciliate with long hairs; leaf blades linear,
spreading to ascending, the largest 6—15 (—20) cm long, 5—10 mm broad,
tapering from the rounded-clasping base to the attenuate apex, upper sur-
face sparsely to copiously strigose-pilose, lower surface heavily so, margins
incrassate, stiffly spreading-long-ciliate. Upper culm leaves gradually re-
duced toward the inflorescence, more distant, sheaths not overlapping and
culm exposed for longer intervals. Spikelets in tight clusters (in depauperate
specimens peduncles may terminate in a single spikelet) on slender, angu-
late, stiffish, hirsute to hirsutulous primary and secondary peduncles, each
cluster from one to several glomerules, some stalked, and arranged either
terminally or also with additional clusters from the next lower 1—2 nodes,
each cluster of glomerules substended by 1—3 involucral bracts similar to
leaf blades but smaller, the longer ones often longer than the spikelet
cluster. Fertile scales obovate, 3.0—3.5 mm long, greenish or reddish-brown,
marginally subscarious, rounded or retuse, ciliate, backs hirsute and/or
hirsutulous, medially strongly 3-nerved, these converging distally to form
a spreading-recurved, stiffish, scabrid mucro nearly as long as the scale
body. Fruiting florets ca. 2.0—2.5 mm long. Sepals smooth, usually extend-
ing to the base of the petal blades. Petal claws often fully 1/2 the total
length, blade above the base of the akene body; blades ovate, basally
rounded, cordate or short-attenuate, flattish and triple-nerved, distally

332

becoming very tumid, often subterete and terminating in a short-conic apex,
eae smooth or with a slight scabrosity adaxially distally. Anthers 3,

. 2mm long. Akene with stipe shorter than the petal claws, surfaces of
sharply trigonous body a lustrous pale to deep brown, angles wirelike,
glassy, paler; stylar end sometimes nearly 1 mm long with a papillose-
hispidulous apex. Fig. 6.

Acid, usually boggy or seep situations, generally in full sun in arenaceous
soil districts, Coastal Plain and Interior Highlands, Louisiana westward
into central Texas, northward into the Ozarks of Arkansas and Oklahoma.

ap 1

Type: Texas: Smith Co., swamps, Swan, J. Reverchon 2911. 10 Jun 1902,
Holotype: MO; Isotypes: GH!, NY!

This species vegetatively and in character of petals is closest to F’. squar-
rosa, In Louisiana and in east Texas the two are sometimes found in the
same bog. As is true of some other sedge complexes where closely related
species may occupy what seems to be the same habitat and not intergrade,
these two appear not to hybridize.

7. FUIRENA SIMPLEX Vahl, Eclog. 2:8. 1798.
F, oe Vahl, Eclog. 2:8. 1798.
F. schiedeana Ku nth, Enum. Pl. 2:183. 1837.
F. penne var. macrostachya Britt., Bull. Torrey Bot. Club 11:87.
1884,

F. cylindrica Bush, Mo. Bot. Gard. Rep. 16:91. 1905.
F. zacapana Bartlett in Rob. & Bartl. Proc. Am. Acad. 43:50. 1907.
F. primiera M.E. Jones, Contr. W. Bot. 18:25. 1933.

Perennial, to 1 m tall, usually 2—4 dm, from stout and short to long and
slender rhizomes, without axillary cormous offshoots. Shoots erect to ascend-
ing or leaning on other plants, tufted or lined out along the rhizome, inter-
nodes terete to somewhat angled, multicostate, usually smooth except
toward the upper culm. Leaf sheaths somewhat inflated, multicostate,
those of mid-culm usually smooth, those of upper and lower culm hispid or
hirsute; ligule short-cylindric, oblique, membranaceous, pale brown, ciliate;
leaf blades linear or lance-linear, larger ones from 5—20 cm long, 3—7 mm
broad, spreading or ascending to erect, usually tapering gradually from
the clasping base to the attenuated apex, the margin usually hispid ciliate,
the upper surface smooth to pilose, the lower surface hispid, pilose, or
smooth. Spikelet clusters grading from solitary and terminal to 2—3 (—5),
the lower or hirsutulous, slender, slightly spreading or erect, axillary
peduncles and all subtended by 1—3, narrowly linear bracts similar to leaf
blades, the longest longer than the subtended spikelet cluster. Spikelets
ovoid, lance-ovoid or cylindrical, 0.8—1.5 (—2.0) cm long, usually acute.
Fertile scales with body mostly obovate or oblong, 2.5—3.5 mm long, thin-
nish, usually tan with tints or maroon and green, or olivaceous, rounded or
slightly emarginate, ciliate, the back scaberulous, puberulent, hispid or

Fig. 6. F. bushit (Kral 24477).

333

3 mm,

334

nearly smooth, usually the lowermost scales of a spikelet with the longest
and most abundant hairs; scale back medially with 5—7 strong, raise
nerves, 3 strongest convergent to form a stiffish but spreading, usually
scabrid, pale green mucro, usually 2/3 or more the length of the scale.
Florets mostly 2—3 mm long (exclusive of bristle tips), calyx bristles
reaching at least to the bases of the petal scales, retrorsely barbellate,
sometimes with narrow blades (i.e. Marsh 1740, Monclova, Coahuila,
Mexico). Petal blades longer or shorter than their slender claws, mostly
vate with ices acute to rounded or retuse, margins sometimes bristly
distally, bases attenuate or truncate, cordate, rounded; bases of pera!

but distally usually tumid, with the mid-nerve dorsally and subapically
excurrent as a short to elongate, erect or incurved mucro. Anthers usually
3, mainly from 0.9—1.2 mm long. Akene prominently stipitate, angles wire-
like, glassy, usually pale, faces flat to slightly concave or convex, a deep,
glossy brown, reddish-brown or even yellowish; stylar apex narrow, usually
papillose-tipped. Fig. 7.

Sunny swamps, swales, bogs, seeps, ditches and wet places, America from
southern Kansas and Illinois southward through the Interior Highlands and
plains of the U.S. into Texas, New Mexico, thence southward through
Mexico (incl. Baja California) into Nicaragua; in the Caribbean in Cuba,
Jamaica, Puerto Rico. Venezuela? Map 1, 3.

This is perhaps the most variable in perianth of all American Fuirena.

Type: Jamaica: ‘‘Bocachica, V. Rohr 27’. “American meridionali’’. Holo-
type: C! The type specimen shows only the leafy culm and the spikelets,
but these are ample, the eee unmistakable.

The common form of the species has 3 calyx bristles and 3 bladed petals
but there are individuals throughout the range that produce a third set of
very short bristles just inward on the receptacle from the calyx bristles.
The calyx bristles are always retrorsely hispidulous or barbellate, their
length varying but usually longer than the petal claws. The blade of the
petal varies over most the range of the species. In the United States, at
least through the Ozark system southwest through Texas, these are usually
deltoid or ovoid, triple-nerved and cordate-truncate basally, becoming
thickened toward the tumid apex, there somewhat emarginate and with a
prominent, subapical-dorsal extension of the mid-nerve, this strongly pro-
jecting and retrorsely barbellate. Such plants are consistently perennial,
strongly rhizomatous, usually tallish, and agree the most with material
from most of Mexico and Central America. To the west in the United States
and rarely in northern Mexico is a low, mostly e-rhizomatous, mostly annual

Fig. 7. F. ete (plant habit from Demaree 61188A) fertile scale and
ret just below from Demaree 61188A; perianth scales at mid-
right from York 46180; perianth scales at lower right from Kral

25288)

339

336

plant with anthers 1/2 as long which appears to be a distinct geographical
variant, here treated as var. aristulata (Torr.). Other variants showing
slight perianth anomalies but otherwise blending well into the matrix of
forms of this quite variable species, have been described as species, to wit:
(1.) F. zacapana Bartlett. Guatemala: Swamp, Gualan, C. C. Deam 423,
13 Jan 1905 (GH! F!). (2.) F. primiera Jones. Mexico: Baja California.
Primiera Agua, near Loreto, 19 Oct 1930, M. E. Jones 27604 (POM! NY! UC!)

Some of the examples of Latin American F. simplex that give the most
trouble are the ones which have the tumid petal scale apex projecting
conically and at the same time have a very reduced subapical bristle.
These must often be examined through all stages of their development in
order properly to be sure of identification.

Sea-coastal races and those from brackish marshes inland show a ten-
dency to produce the most extensively creeping rhizomes, often develop
clones of gigantic size. Such plants are also smoother, shorter and harder-
leaved, stiffer in habit.

7a. FUIRENA SIMPLEX var. aristulata (Torr.) Kral, comb. nov.
F. squarrosa var. aristulata Torr., Ann. Lyc. Nat. Hist. N.Y. 3:291.
1836.

Mostly annual, mostly without rhizomes, the slender culms single or in
small tufts, erect or spreading, rarely taller than 3 dm. Lowest leaves
mostly sheathing and bladeless or with mucro-like blades, smooth to sparsely
hispid or hispidulous. Larger leaf blades ascending or spreading, lance-
linear, 10 cm or less, attenuate, the margins hispid-ciliate at least proxi-
ally, the surfaces smooth to hispid or hispidulous, particularly along the
costae; sheaths multicostate, terete or angled, smooth to sparsely or copi-
ously hispidulous. Bot and blades becoming progressively more
pubescent upward on the culm. Upper portion of culm leafless, multicostate,
usually hirsutulous. Spikelets in 1—3 clusters per culm, usually the terminal
largest, the lower on slender, erect or ascending hispidulous peduncles, at
least the uppermost cluster subtended by 1—3 linear-subulate bracts, these
spreading or reflexed, 1 usually longer than the cluster. Spikelets mostly
lance-ovoid, ca. 1 cm long or less, acute. Fertile scales with body obovate,
rounded, 4.0—4.5 mm long, puberulent, brownish-green or olivaceous, mid-
nerves 3, strongly raised, converging to a spreading or squarrose but stiffish,
seaberulous or hispidulous mucro fully 2 mm long. Perianth 6-parted, the
calyx bristles 1—2 mm long, retrorsely scabrid, the bladed members ca.
1.5 mm long with blades longer than the claws, oblong or ovate, truncate
or retuse, flattish or distally tumid, the awn slightly subapical, dorsal,
erect, very short to nearly 1 mm long and itself retrorsely barbellate;
nerves of scale 3, the laterals arcuate. Stamens 1—3, the anthers mostly
0.5—0.6 mm long. Akene stipitate-trigonous, including the stylar end 1.5—1.7
mm long, the body broadly elliptic in outline, the angles prominent, wire-

‘g dew

CARIBBEAN

AMERICA No. 113

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338

like, the faces flattish and glassy, usually pale brown or near white, the
narrow stylar end smoothish or apically papillate, ca. 0.5 mm long. Fig. 7a.

andy or sandy-peaty low places in prairies, river and stream bottoms and
seeps, Nebraska (mostly in the Platte and Loup systems) southward through
Kansas and northwestern Missouri through Oklahoma and Texas. Rare and
local in northern Mexico; New Mexico. Map 1,3.

Type: ‘Arkansas’? Long’s Expedition to the Rocky Mountains, coll. by
Dr. James. Holotype: NY, but Coville (1890) cites this, together with ma-
terial I have on loan, to wit: New Mexico, A. Fendler 877; Texas: Comal
Co., New Braunfels, F. Lindheimer 185, 186; Comanche Springs, F. Lind-
heimer 206, 1244.

One might question the advisability of retaining this as a distinct variety,
particularly in context of a species as variable as F. simplex. My reasons
are that this taxon appears to occupy a definite geographic area and is
rather consistently a lower, more slender, and almost always a tufted e-
rhizomatus plant. Its anthers at maturity almost always range between
0.5—0.6 mm in length while those of the rest of F. simplex are mostly
around 1 mm long. There are some anomalies, i.e. material from Oklahoma:
Greer Co., G.W. Stevens 1018.1, 18 Jun 1913, which has rhizomes. Such plants
are occasional in southern Kansas, Oklahoma, and in Texas. But this might
be in keeping with a variety of comparatively recent derivation from F.
simplex. I hope to be able to study the chromosomes of this, essentially
prairie plant, in order to see if there are real genomal differences.

8. FUIRENA PUMILA (Torr.) Spreng., Syst. Veg. I: 237. 1825.
F. squarrosa var. pumila Torr., Fl. N. & Midl. States I: 68. 1824.
F. torreyana Beck, Bot. U.S. 429. 1833.

Caespitose annual 0.8—6.0 dm tall, mostly 3 dm or lower. Culms erect or
spreading, stiffish, but slender, the internodes multicostate, lower ones
smoothish, the upper toward and in the inflorescence hispid. Lower leaves
mostly sheath, those toward mid-culm with sheaths loosely cylindrical,
smoothish to hispid, somewhat dilated distally, there with a pale to reddish-
brown, thin, oblique, ciliate, ligule; blades spreading or ascending, linear-
lanceolate or linear, mostly 5—12 cm long, 3—5 mm broad, mainly tapering
from near the clasping base to the narrowly acute apex, margin somewhat
cartilaginous-thickened, usually spreading-hispid or scabrid, surfaces stri-
gose-hispid or smooth, multicostate beneath. Leaf blades thence somewhat
reduced into the inflorescence, topping longer internodes, and usually hispid.
Inflorescence a terminal cluster of spikelets, leafy-bracted, and often with
pedunculate similar clusters from the next-lowest 1—2 nodes (in depauperate
specimens the culm may produce but 1 spikelet!). Spikelets ovoid or ovoid-
cylindric, mostly 0.7—1.5 cm long, the lowest bracts hairiest, often quite
hispid or hispidulous. Fertile scales with bodies mostly obovate, ca. 3 mm
long, rounded or slightly emarginate, ciliate, then, mostly greenish-brown,

339
backs rounded, hirsute, hispidulous or hispid, with usually 3 strongly raised
6 members, calyx bristles retrorsely barbellate, extending at least to the
base of the petal scales and often to near their tips. Petal blades long-

clawed, mostly ovate, slenderly incurved-acuminate into apical bristles, the
base rounded cordate or truncate, body thin and 3—5-nerved basally but

LOM.

Fig. 7a. F. simplex var. aristulata (McGrath 6016).

340

medially and distally thickened. Anthers 0.5—0.7 mm long. Fruit stipitate,
body trigonous, edges wirelike and pallid, faces a deep, lustrous brown or
reddish-brown; apiculus narrow, somewhat papillose and thickened distally.
Fig. 8.

Moist to wet, usually sandy or sandy peaty places, sometimes weedy,
toward the coast from Massachusetts southward through the Coastal Plain
to southern peninsular Florida, west in the Gulf Coastal Plain, but increas-
ingly infrequent, to southern Texas with a disjunction inland in the Great
Lakes lowlands of Michigan and Indiana. Map. 2.

Type: New York: Long Island, J.R. Torrey. Holotype: NY!; Isotype:
GH!

Many have considered this as being but an annual variety of F. squarrosa,
explaining that that species becomes annual north in its range. However,
F. pumila has the same annual habit however far south it grows (and is
annual in the greenhouse as well!), has a consistently different petal blade
character, has consistently smaller, shorter anthers, and never produces
the cormous rhizome buds as in F.. squarrosa.

The nomenclature of the species has been a bit clouded because of the
reluctance of some authors to cite the basionym. Thus the species has often
been credited to Sprengel, though he himself made it plain that the name
originated with Torrey.

9. FUIRENA UMBELLATA Rottb., Descr. & Ic. 70. 1773.

F. paniculata L.f., Suppl. 105. 1781.

F. tereticulmis Presl, Rel. Haenk. 1:180. 1828.

Scirpus umbellatus (Rottb.) O. Ktze., Rev. Gen. 3(2):337. 1898.

Perennial, the thick, scaly, fibrous-internoded rhizomes producing cor-

mous offshoots. Culms stiffly erect or leaning, 5—11 dm tall, internodes mul-
ticostate and terete or frequently sharply 3—5-angled, toward the stout base
(1.0—1.5 cm thick), smoothish or becoming hirsute or hispid at or toward
the inflorescence. Leaf sheaths longest and overlapping at or near culm
base, loosely investing the internode, cylindric, multicostate, smooth to hir-
tellous or hispid, particularly toward or at the inflorescence level; ligular
orifice thin, brownish, erect, oblique, short-ciliate; blades short, scale-like
at culm base, becoming longest at mid-culm or above, there linear-lanceo-
late, linear-oblong, 10—30 cm long, 0.8—2.5 cm broad, narrowly acute to
attenuate, the margins smooth or scaberulous, the base clasping, upper
surface smooth to sparsely or finely hispid, strigose, or scaberulous, finely
and evenly nerved, lower surface smooth or variously hairy, strongly multi-
costate, the nerves often hispid. Leaf blades gradually reduced in size into
the inflorescence, this a compound of distant to approximate clusters of
sessile and pedunculate glomerules of spikelets, the lowest clusters on
slender, elongate, usually hispidulous primary peduncles, the whole struc-
ture interruptedly cylindric and each cluster of glomerules subtended by

Fig. 8. F. pumila (Kral 44735).

341

342

1—3 leaflike hairy bracts, the longest longer than the cluster. Spikelets
lance-ovoid or cylindrical, 5—8 (—10) mm long. Fertile scales oblong to
obovate, 2.5—3.0 mm long, rounded or retuse, distally ciliate, thin, the backs
greenish-brown, smooth to puberulent or strigillose, medially 3-nerved,
these converging to a subapical, stiffish, flattish or subterete, ascending to
spreading, scabrid mucro ca. 1/3 as long as the scale body. Florets 2.0—2.5
mm long. Perianth of but 3 scales (petals) the blades on crimped claws,
short-oblong or obovate, 1.5—2.0 mm long, obtuse, rounded or emarginate,
incurved-aristate, the arista often coiled apically, the blade body 3—5-
nerved, thinnish. Stamens 2 or 3, anthers 0.7—1.0 mm long. Akenes 1.0—1.5
mm long, body angles wirelike, paler than the brownish, shining faces;
apiculus apically papillose or smooth. Fig. 9.

Swamps, bogs, seeps, marshes and low, wet meadows, throughout the
warmer, more humid parts of mainland Latin America and in the Carib-
bean. Old World tropics. Map 4.

Type: Surinam, Rolander. Not examined. The plate (from Rottb., Tab.
XVII. Fig. 3) shows a very good rendition of the slender, long arista on
the petal scales, this unmistakably identifying the species.

10. FUIRENA CAMPTOTRICHA C. Wright in Sauv., Anal. Acad. Si. Habana
8:32
F. bulbipes Blake, Contribs. U.S. Nat. Herb, 24:2. 1922.
F. umbellata var. unguiculata Kukenth., Report. Sp. Nov. 23:200. 1926.

In rhizome, culm, leaf, and generally in habit much like F. wmbellata.
Leaf blades mostly 5—20 cm long, 0.5—1.5 ecm broad, usually spreading
hispid-ciliate at least toward the bases, the ligular sheath-orifice short-
ciliate, sometimes also strigillose. Inflorescence somewhat sparser, often
more open. Spikes ovate to lance-ovoid or cylindrical, mostly 0.5—1.0 cm
long, reddish or greenish-brown. Fertile scales as in F. umbellata with
mucro seldom more than 1/3 the scale body length, erect or slightly spread-
ing. Florets as in F. umbellata, the perianth scales on crinkled stipes, but
the distal 4% of the blade tumid, hispidulous, and with a shorter, subapical
mucro or awn (this in a few cases very minute or absent). Fig. 10

Bogs, swamps, wet sunny places, near the southern coasts of and in the
Isthmus of Mexico, southward into Panama and South America; Cuba.

ap

Type: Cuba. Havana: en sabanas humedas cerca de Dayaniguas, Juris-
diccion de los Palacios, C. Wright 3778; GH!, NY! This material shows the
short, slender, subapical awn on the petal scale, together with the tumid
scale apex typical of the species.

As Svenson (1957) has commented, this is so close to F. umbellata in
many respects that it well could have arisen from that more widespread
species. It may well be that a world revision of the genus would reduce
such as F. camptotricha to varietal rank. However, in perianth character

yy

Si

Fig. 9. F. umbellata (Curtiss 299).

343

344

Fig. 10. F. camptotricha (Kral 25066).

‘5 dew

F UMBELLATA

at

FE REPENS ©

EINCOMPLETA ®
VAR. OBLITERATA 4

GPE

346

it does show, with its subapical petal, bristle and tumid petal blade, some
overlap with F. simplex, another species with which it is sympatric. Type
material under the C. Wright number 3778 has been interpreted variously
in that it also contains some F’. robusta. But the Wright description on the
one hand declares ‘‘setis nullis,’’ thus eliminating F’. robusta, and on the
other includes the swollen perianth scale character, which eliminates F.
umbellata, the species to which Svenson (1957) assigns the Wright speci-
mens. A particularly bad specimen to interpret is Nee and Mori 3636 from
Colon, Panama, ‘‘0.7 mi NE of bridge over Rio Piedras on road from Porto-
belo to Pilon.’’ This has petal blades crinkle-stiped, swollen distally and
with a short subapical bristle as in F. camptotricha; on the other hand
there are small bristles also, the base of the leaf blades are prominently
and stiffly ciliated and the spikelet has larger dimensions, all characters of
F. robusta. Such specimens (however rare) are an indication that the next
phase of the study has to be an investigation of the cytology and breeding
system in these Fuirena.

11. FUIRENA REPENS Boeckler, Bot. Jahrb. 7:277. 1886.

Slender low perennial, the rhizomes creeping, slender but swollen at ir-
regular intervals into narrowly fusiform thickened zones, pale reddish-
brown, internodes frequently 2 cm long, often forking or 3-branched or
producing compact, narrowly fusiform, several-noded rhizomatous off-
shoots, these developing apically into leafy shoots. Shoots erect, low, rarely
reaching 2.5 dm, the lowermost part scaly, smoothest, the leaves of mid-
culm longest; sheaths tubular-cylindric, rather loosely investing the culms
and often overlapping, multicostate, hirsute, the ligular orifice scarious,
oblique, long-ciliate, pale brown; larger leaf blades narrowly linear, as-
cending or spreading, to 5 cm long, rarely broader than 5 mm, attenuate
from near the base to the slender apex, smooth to sparsely or copiously
hirsute, sparingly spreading ciliate proximally. Culm smooth proximally
multicostate, with internodes becoming spreading hirsute upward, terminat-
ing in a subcapitate cluster of (1-) 3—7 spikelets subtended by 1—3 leaflike
bracts, these shorter than to twice the length of the spikelets, spreading,
hispid-hirsute proximally, smooth to scabrid distally. Spikelets lance-ovoid,
greenish or olivaceous, the longest ca. 1.0 cm long. Fertile scales obovate,
thin, ca. 2.5 mm long, the backs hispidulous and sparingly strigose, medial-
ly 3-nerved, these converging to form a spreading-ascending mucro to 1.5

m long. Perianth with calyx bristles reduced to minute sharp-tipped tu-
bercles; petals ca, 2.0—2.5 mm long, the claw ca. 0.4 mm long, the blade
ovate, proximally flattish and triple-nerved, the distal Y% inflated and taper-
ing into a slender, erect to incurved awn. Anthers ca. 2 mm long. Akene
ca. 1.5 mm long altogether, the stipe shorter than to nearly as long as the
sharply trigonous body, the stylar end slender elongate, slightly broadening
and scaberulous-papillose distally; akene surface pale, minutely cancellate-
punctate, the 3 edges smooth, wirelike, glassy. Fig. 11; map 4

347

1a

Fig. 11. F. repens (Schaffner 196).

348

Type: Mexico: San Luis Potosi: “San Louis Potosi, leg. J.G. Schaffner
196, 1879. Herbarium V. A. Vigener.’’ Holotype presumably at Berlin and
destroyed. Neotype: NY!; isoneotype: C

This species, so far as I know, has only been collected from its original
pag and only two numbers, both collections of J.G. Schaffner are ex-
tant. I have designated ee specimen at NY as the neotype. The label of
a Schaffner 567 (GH! PH!) adds somewhat to the sparse information
about he species, a te ‘in paludosis, San Rafael, ex convalli San Luis
Potosi

A tieci: there is little information on which to base retention of this
as a species. However, it differs from the others in so many respects as to
deserve reconsideration. Hopefully the plant will be refound.

12, FUIRENA INCOMPLETA Nees in Mar., Fl. Braz. 2(1):107. 1842.
F. hexachaeta Schlecht. Linnaea 19:69.
Scirpus incompletus (Nees in Mart.) T. Koyama, Journ. Fac. Sci. Univ.
Tokyo, II, 7:287. 1958.

Perennial 4—14 dm tall, from creeping rhizomes, the culms erect, ascend-
ing or leaning on other vegetation, nearly caespitose or set closely together
in lines. Internodes usually smooth except in the inflorescence, usually
sharply 3-angled. Lowest leaves mostly sheathing and bladeless or with
short-triangular blades, usually smooth or puberulent, mid-culm leaves
with sheaths loosely enfolding internodes, mostly sharply 3-angled, smooth
to puberulent or hispid, the ligular orifice very short-cylindrical, oblique,
scarious, reddish-brown, smooth to hispidulous; leaf blades at mid-culm
longest, linear, mostly tapering gradually from base to apex, 4—17 cm long,
4—7 mm broad, the edges smooth and cartilaginous thickened, sometimes
remotely ciliate proximally, the surfaces smooth or strigillose along the
nerves beneath, particularly along the strongly raised median. Inflorescence
with central axis usually hispidulous, either a single terminal cymose com-
pound of glomerules of spikelets, or a series of 2—4, well-separate com-
pound of glomerules, the lowest usually on short to elongate, slender, erect
to slightly divergent primary peduncles, these densely hispidulous, sharply
3-angled (sometimes the spikelets of clusters tending to be compactly pin-
nately disposed!). All clusters of glomerules subtended by leaflike bracts,
the lowermost longest, usually equal to or longer than the subtended spike-
let cluster, then gradually shorter, grading to smaller ones subtending in-
dividual glomerules, thence to the lowermost fertile scales. Spikelets nar-
rowly oblong-cylindric to lance-linear, 0.7—1.4 cm long, usually acute. Fer-
tile scale body broadly elliptic, oblong, or obovate, 3.0—3.5 mm long, round-
ed, short-ciliate or nearly entire, very thin, the backs mostly greenish-
brown or olivaceous, rounded, appressed-puberulent or hispidulous, usually
with 3 greenish, slightly to strongly raised median nerves convergent to
an erect or slightly spreading mucro % as long to nearly as long as the
scale body and strigillose. Perianth of 4—6 subequal, retrorsely barbellate

349

73).

F. incompleta (Killip 45

Fig. 12a.

300

bristles, these extending to or exceeding the style tip of the akene. An-
thers mostly 3, 0.8—1.5 mm long. Akene short-stipitate, 1.5—2.0 mm long,
the body trigonous, broadly or narrowly rhombic in outline, angles promi-
nent and wirelike, faces concave or flattish, lustrous brown or greenish-
brown, stylar apiculus short, distally papillose. Fig. 12.

Marshes, bogs, seeps, usually in full sun and at elevations of 5000 ft. or
more in the mountains, Mexico southward into southern South America.
Map 4.

Type: Brazil: Sta. Maria. Goyaz, Pohl. The type specimen has not been
examined by this writer, but the description leaves no doubt. Two varieties
are distinguished by Nees, mainly based on pubescence character. The
variety ‘‘alpha,’’ according to Nees is pubescent, the sheath angles hirsute:
ciliate, the leaves puberulent above, hirsute beneath. The variety ‘‘beta’
is, for the most part, smooth. Material I have seen from north of South
America all appears to fit the former variety.

Toward the north of the range of the species, in Mexico from the state
of Mexico north and west to Sonora in the Sierra Madre Occidental, a geo-
graphical variant appears which merits description, namely:

12a. FUIRENA INCOMPLETA var. obliterata Kral, var. nov.

F. incompleta Nees var. alpha affinis, a qua imprimis differt perianthii
absenti vel vestigiali, setis redactis inaequalibus achenii stipite breviore

Similar to the var. ‘alpha’ of F. incompleta but from which it differs
particularly by the perianth absent or vestigial, with the reduced bristles
unequal, shorter than to slightly longer than the stipe of the akene. Fig.

While this variety overlaps F’. incompleta proper in the state of Mexico.
it is the sole representative of the species northward. It appears from the
type description of F. hexachaeta Schlecht., that it does not vary signifi-
cantly in any way from that rendered for the species F’. incompleta by Nees
ab Esenbeck.

Known localities for the new variety are as follows: Type: Mexico: State
of Mexico: north end of Tenancingo; large seep near Tenancingo Park by
Mex. hwy. 55, R. Kral 25234, 30 Jul 1965. Holotype: US!; Isotypes to be
dictabuted: Other records: Mexico: Jalisco State: Sierra del Tigre 3 mi
S Mazamitla, R. McVaugh & W. N. Koelz 410 (MICH); sandy clay of seep-
age by Mex. hwy. 15, 17.3 mi E Guadalajara, el. 5000’, R. Kral 25622. Mex-
ico State: Temascaltepec, Penon, 1700 meters, Hinton 4406. Nayarit State:
rocky sloping oak savanna, el, ca. 4000’, 9.5 mi WNW Chapallilla by Mex.
hwy. 15, R. Kral & J. Murrell 27555; seepage area in red sandy clay soil,
oak type with scattered pine, between Chapallilla and Ixtlan beside Mex.
hwy. 15, R. Kral 25648. Sonora State: Tepopa, Rio Mayo, rooted in mud of
springside, H.S. Gentry 1412 (F, GH, MICH, UC). Map 4.

1. FUIRENA WALLICHIANA Kunth, Enum, Pl. II. as 1837.

Below is a description based solely on a collection made by Dr. Clyde

F. Reed, seemingly from a chance (temporary?) introduction appearing

2mm

Fig. 12b. F. incompleta var. obliterata (Gentry 1312).

/2 b

301

352

in “wet areas of chrome ore piles’? in Newport News, Warwick County,
Virginia. The collection, deposited at PH, is described as follows.

Perennial to 5 cm tall, smoothish, culms tufted, close-set along branched,
thickish, scaly rhizomes. Internodes smooth, strongly few-costate, slender,
longest toward the inflorescence. Sheaths tubular, slightly dilated around
the shoot, the lowest lacking blades, oblique, scarious-margined, converging
to small cusps. Sheaths at mid and upper culm shorter, their orifices bear-
ing an oblique, scarious, low-ciliate, short-cylindric ligule. Largest blades
toward mid-culm or above, these 5—10 mm long, narrowly linear, no wider
than 3 mm, gradually attenuated from near the blade base, smooth except
for the scabrid, slightly involuted, margin, the lower surface rather strong-
ly costate and with a prominently raised mid-rib. Inflorescence either a
single terminal spikelet or an inequilaterally arranged cymose system of
spikelets, usually few, digitately spreading or in some cases spreading pin-
nately along secondary peduncles, the spikelet clusters usually subtended
by a linear-filiform-bladed bract, this longer than the cluster; all peduncles
angulate, scabrid. Spikelets lance-ovoid or lance-cylindric, acute, ca. 0.7—1.0
cm long, greenish brown. Fertile scales ovate or oblong, ca. 3 mm long,
thin, appressed-puberulent, bearing medially 3 rather indistinct nerves
converging to a slender, smoothish or papillate-scabrid, pale green, erect
to slightly spreading mucro % or more as long as the body. Perianth bris-
tles 2—6, of various lengths, often geniculate, sometimes distantly and ir-
regularly pinnately few-branched, even somewhat webbe etween the
branches on largest bristles, the branches and axis tips marginally long-
papillate. Anthers 3, ca. 2 mm long. Fruit trigonous, ca. 1.5 mm long, ellip-
soidal in outline, the stipe short, the angles sharp, pale papillate-serrulate,
the faces convex, strongly cancellate, the stylar apiculus short-triangular,
pale, papillate. Fig. 13

The webbing between the sparse branches of the petal bristles approach-
es the description of F’. coerulescens Steud., but the habit of the plant, the
character of its fertile bracts, the tendency for the bristles to be shorter
and the strongly cancellate akene faces suggest F. wallichiana Kunth, a
species of the East Indies. This particular material is interesting in that
it shows a transition between a species such as F. incompleta whose peri-
anth is entirely of bristles and those which show the conventional Fuirena
perianth comprised at least in part of bladed members. Certainly one sam-
ple alone is not basis enough for identification, particularly in that the speci-
men comes from a very extreme habitat

REFERENCES

BECK, L.C. 1833. Botany of Northern and Midland States. Albany.
BOECKLER, ©. 1886. Fuirena in Bot. Yahrb. 7:277.

aa wee neuer cyperaceen. Allg. . Zeits. 2:77-79.
BRIT TON, N.L. and A. BROWN. 1913. ca ae — the Northern United States,
Canada, and fhe Br ‘ich Possessions, Vol, New York.

BOSE B.F. 1905. The north American species of Fuirena. Mo. Bot. Gard. Rep. 16:87-99.

393

SS,

Sim,

=e
Sex
i @

/

6,
Reo’ Ne 544
Wo

13

Fig. 13. F. wallichiana (Reed 44066).

304

CHAPMAN, A.W. 1897. Flora of the Southern United States, ed. Nev
CLARKE, C.B. 1909. iceacon of Cyperaceae. William aad oo. the a
COVILLE, F.V. 1890. eee of the United States species of the genus ae Bull.

(1) :1
ELLIOTT, S. 1813. Nee of the Botany of South Carolina and Georgia. Charleston.
GONZALES-MAS. re 1964. Cyperaceae of Puerto Rico. Dissertation.
HARPER, R.M. 1901. Collection of plants in Georgia. eed - . Club 28:466.
JONES, M.E. 1933. Fausrena in Contribs. to Western Botar
KRAL, R. 1971. A treatment of Abid gaardia, Bulbostylis a eile (Cy peraceae)
for Ua eae Sida 4(2):57-227.
KUKENTHAL, G. 1926. Repert. Spec. Nov. edde
KUNTH, se 1822. Synopsis Plantarum I. e ris.
7 . 1837. ava aa Plantarum Hl. Stutgardiae et Tubingae
LANJOUW, J. and F. A. STAFLEU. 1964. Index Herbariorum i The Herbaria of the
67.

World, I-xvi: 1-1

23:183-222.

LINNAEUS, C. 1781. Se ae alan ann eae Systematis Vegetabilium 13. Brunsvigae.
MICHAUX, A. 1803. Flora eali Ame I. Paris.

NEES VON ESENBECK, oe ees ae in Martius, aa Brasiliensis, Vol. 2.
ALLA, E. 1908. Densissens K. K. Akad. Wiss Bosch: Wien.

Sp.
PRESL, J.S. 1828. Fasrena in Rel. Haenk. I.
ROTTBOLL, C.F. 1773. Descriptionem et Iconem Rariores. Hauniae.
SCHLECHTENDAL, D.F. 1847. Fuirena in Linnaea 19:68-70.
SMALL, J.K. 1933. Manual of the Southesatern Flora. New York.
SPRENGEL, K. 1825. Systema Vegetabilum I: 303-304.
STEUDEL, E.G. 1855. Synopsis Plantarum Glumacearum, part IJ. Stutgart.
SVENSON, H.K. 1957. North American Flora 18 (9) :505-507.
TANAKA, N. 1939. Chromosome Studies in Cyperaceae VII cae Mag. of Japan: 480-488.
1941, Chromosome Studies in Cyperaceae XII. Bot. Mag. of ie 649-660.
TORREY, ER. 1836. nee — North American Com nen Ann. Lyc. N.Y. 3:239-356.
VAHL, M. fae Enumeratio Plat um I. Hauniae.
= -

: : . Eclogae Americanae 2. Hauniae.
WRIGHT, - je Sauv. Anal. Acad. Sci. Habana 8.

A REVISED DESCRIPTION OF THE
SALT MARSH RUSH, JUNCUS ROEMERIANUS.

LIONEL N. ELEUTERIUS
Gulf Coast Research Laboratory
Ocean Springs, Miss. 39564

ABSTRACT
A reevaluation of nine taxonomic characteristics presently used to identify
Juncus roemerianus, in view of recent morphological, populational and dis-
tributional information, indicates that existing descriptions are inadequate
or incorrect. A new description of the species is given

INTRODUCTION

Juncus roemerianus Scheele is found only on the east coast of North
America (Weimarck 1946, Eleuterius 1975), where it produces a copious
vegetational cover over tidal marsh on the south Atlantic and Gulf Coasts
of the United States (Eleuterius 1976a). During a recent autecological study
of the species two distinct plant types were recognized based on flower
morphology. One plant type was hermaphroditic, bearing perfect flowers
and the other was female, bearing imperfect, pistillate flowers (Eleuterius
and McDaniel 1974, Eleuterius 1974). The plants spread rapidly by means
of vigorous rhizome growth and a single plant type may dominate several
acres or more of tidal marsh (Eleuterius In review a). In view of these
and other recent findings the biological characteristics presently used to
identify and describe J. roemerianus need reevaluation and clarification.
Manuals used to describe this plant have erroneous descriptions. The salient
taxonomic features of the plant species have not been previously recognized,
thus inadequate descriptions prevail. Although I have studied herbarium
specimens of other members of the Junci Thalassi of which Juncus roemer-
ianus is a member, I feel that this ‘‘classical’’ approach may perpetuate
existing errors in the literature. This paper covers nine taxonomic features
and the reasons for the needed change. A revised description of the tidal
marsh rush is given.

Historical sketch. Scheele (1849) first described and named Juncus roe-
merianus from plants bearing flowers collected on Mustang Island at Gal-
veston, Texas. Prior collections of the species from North America were
considered Juncus maritimus, the European form named by Lamarck
(1789). However, Chapman (1860) discovered an isolated colony of true J.
maritimus on Long Island, New York. Later authorities such as Coville
(1894) cite Chapman as the author of J. maritimus. Although Scheele dis-
covered what he considered a new species, he did not describe the distinc-
tive features of J. roemerianus. Engelmann (1861-1868) described the plant

SIDA 7(4): 355—360.

356

and pointed out some of the distinct characteristics which separate it from
the closely allied J. maritimus. This separation was based primarily on seed
morphology. He also stated that he found ‘‘a rare form of J. roemerianus

where both circles of stamens were suppressed or rather dnaeraevelopee
and in a rudimentary state so that those plants became unisexual.’’ He also
noted that J. roemerianus was the only species of Juncus which produce
unisexual plants. Corresponding male plants were not seen, but he sug-
gested that they may exist. The specimens that he observed were from
Georgia and Florida. Other early workers such as Coville (1894), Chapman
(1897), Small (1903), and Blankinship (1903) listed J. roemerianus in their
papers or manuals.

Breeding system. Chapman (1897) and Fernald (1950) stated that Juncus
roemerianus Was dioecious and Small (1903, 1933) suggested that the ‘‘flow
ers were usually dioecious.’’ Radford et al. (1968), Correll and Johnston
(1970), Correll and Correll (1975), Gleason (1968) and Jones (1975) consid-
ered the species to be hermaphroditic, having perfect flowers. However,
zleason (1968) stated that the stamens were ‘‘usually none in fertile flowers.”
Long and Lakela (1972) described the flowers as unisexual, but do not
clarify their eal Thus, the species may be assumed to be monoe-
cious or dioec

In recent oe only perfect and pistillate flowers have been found, al-
though thousands of inflorescences of Juneus roemerianus have been ex-
amined (Eleuterius and McDaniel 1974). Collections were taken from Mis-
sissippi, Texas, Louisiana, Alabama, Florida, Georgia, South Carolina and
North Carolina. Extensive study of the rhizomes during anthesis show:
that they bear culms with inflorescences composed ae ee of pistillate
or perfect flowers. A given rhizome produced only one flow ype, never
both. Transplanted clonal material from unisexual and tea plants have
for five years consistently produced only the respective flower type (Eleu-
terius and McDaniel 1974, Eleuterius 1974, In review b). Plants grown from
seed have also consistently produced a single flower type (Eleuterius, In
review ¢c).

I have also searched in vain for staminate flowers in plants comprising
mature stands and in the progeny of known parental types from widely
separated areas. Only plants with pistillate or perfect flowers are appar-
ently produced. These data offer sufficient proof that the species is gyno-
dioecious.

Involucral bract. The terminal bract is erect and terete and appears to
e a continuation of the stem. Correll and Johnston (1970) and Correll and
Correll (1975) state that the terminal bract is 3 times as long as the in-
florescence. Observations of plants in various habitats indicate that. this
feature is unreliable as a taxonomic characteristic since the bract may
range from 2—90 cm in length and may be from %4—10 times the length of
the inflorescence.

357

Inflorescence morphology. Small (1903, 1933) described the culm as a
scape, while Correll and Johnston (1970), Correll and Correll (1975) state
that the inflorescence is a panicle, with compound branching, 7—12 cm in
length. Buchenau (1906) described the inflorescence as a panicle, but did
not indicate size. Gleason (1968) stated that inflorescence was 6—15 cm in
leng

Recent work corroborates paniculate descriptions of the inflorescence;
however, the form and size are extremely variable between populations
(Eleuterius 1974). The inflorescence may vary from a few- to a many-
branched panicle. The length of the inflorescence branches may vary, pro-
ducing in some instances tufted inflorescences. The reason for this phe-
nomenon is unknown. In most instances the inflorescence is ae branched,
but ranges from 2—45 cm in length. The variations within populations is
relatively small; however, that between certain populations is considerable
differing by 100 orders of magnitude or more (Eleuterius, In revi
cause of this variation is presently unclear, but under ea

Flower cluster. Correll and Johnston (1970) and Correll and Correll (1975)
indicate that the flowers occur in clusters of 2—5. Gleason (1968) states that
the sessile clusters occur in groups of 2—4, with two to six flowers in each.
I find that the sessile clusters occur in groups of 2—6, with from 2—8 flow-
ers per cluster. The solitary flowers referred to by Jones (1975) are obvi-
ously immature, since flower development occurs sequentially in each clus-
ter iicuiers. 1974,

Bracteole. Eleuterius (1974), Gleason (1968) and Correll and Johnston
(1970), Correll and Correll (1975) point out that each flower is subtended
by a short, ovate bract. I have not observed any occasional extra bract
mentioned in the latter two references cited above which may represent
a feature peculiar to a localized population on the Texas coast.

Capsule. Descriptions of the capsule in the major taxonomic reference
manuals are highly variable. This is understandable since the mature cap-

flcrescences with occasional capsules containing 6—8 carpels, suggesting
genetic aberration.

Seeds. Only Gleason (1968), Long and Lakela (1972), and Radford et al.
(1968) present descriptions of the seeds, although they are obviously im-
portant in separating Juncus roemerianus from closely related species.
Many seeds, especially from hermaphroditic plants, apparently do not reach
maturity; these are undersized, transparent and have low viability. Since
seed development is rapid, reaching maturity in 3—5 weeks (Eleuterius
1975), and each capsule encloses seeds in different stages of development,
the difference, although slight, is sufficient to merit special attention in
identification. The seeds may be elongate, wedge-shaped or round, generally
not caudate, but often with a slight point at one end. Englemann (1861-

398

1868) stated that the only apparent difference between J. roemerianus and
the European Juncus maritimus Lam. is the presence in the latter of a
minute hooked appendage at one end of the seed. This is questionable and
merits further study.

Plant form and size. Correll and Johnston (1970) and Correll and Correll
(1975) state that Juncus roemerianus is a tufted perennial. Long and Lakela
(1972) state that it occurs in large tufts with the culms spaced in rows on
horizontal scaly rhizomes. While these statements are essentially true,
J. roemerianus generally does not grow in tufts, like Juncus effusus L., be-
cause the erect shoots are widely separated by long rhizomes. The inter-
shoot distance along the rhizomes of J. effusus is very short, producing a
closely arranged, dense mass of erect shoots or tuft. The elongated, spread-
ing rhizome of J. roemerianus is apparently unique among rushes (Eleu-
terius 1976b), especially in context of species found in dry ground or fresh-
water marshes. Although I have examined Juncus gerardii L. from New Eng-
land, I have not examined adequate material of the European J. maritimus
or the South American J. acutus var. leopoldii (Parlatore) Buchenau an
other taxa inhabiting saline tidal marshes.

Small (1908, 1933), Correll and Johnston (1970), Gleason (1968) state that
the height of Juncus roemerianus varies from 5—12, 5—15, 4-10 dm, re-
spectively. Jones (1975), Long and Lakela (1972) and Radford et al. (1968)
state that the species is 1, 1.5 and 0.5—1.5 m tall respectively. Mature
plants (bearing flowers) have been collected ranging in height from 0.3—2.5
m. Plant height is generally uniform within a given stand, but very different
between stands. The ‘‘average’’ height of J. roemerianus observed through-
out the range of the species probably lies somewhere between 10.5—13.5 dm.

Period of anthesis. Correll and Johnston (1970) state that flowering occurs
in the spring, while Long and Lakela (1972) state that it occurs in the fall.
Radford et al. (1968) indicated that anthesis covers a period from May to
October while Jones (1975) indicates the period extends from March to June.
Direct observation on flowering over a wide range of habitats and through a
number of years, study of herbarium collections and personal communication
with a number of coastal researchers have provided adequate information to
clarify the above inconsistencies. In North Carolina the species produces
flowers from January to June, in extreme south Florida the species flowers
during March and April, while in Mississippi it occurs from January to April.
This longitudinal gradient suggests that the length of time that the plants
are subjected to low temperatures (followed by increasing day length) is
a critical factor influencing the duration of anthesis. The longer cold periods,
apparently, correspond to longer periods of anthesis. The above reference
indicating that flowering occurs in the fall is erroneous, although the dead
culm with intact capsules may exist for a year or more (Eleuterius 1976b).
When wet the empty capsules close. Plant workers less familiar with the spe-
cies probably collected specimens with this persistent culm during the fall,
thus, the irregularities.

399

The following revised taxonomic description summarizes the preceding
subject discussions and presents other morphological details of value in iden-
tifying Juncus roemerianus.

JUNCUS ROEMERIANUS Scheele

Stout, rigid, densely spreading, gynodioecious perennial, with erect shoots
1.5—23.0 dm high; leaves all basal, terete, longer than the culm, the sheaths
inflated with free margins, dark green with grey or brownish cast, auricles
well developed to several mm high, cartilaginous; culms terete, pungent, less
rigid than leaves; involcucral bract, stout, spinescent, varying in length from
a few cm to 3 dm or more, conspicuous, generally longer than the inflor-
escence, resembles leaf in cross-section; inflorescence paniculate, scape
branches very unequal, 2—30 mm long; flowers perfect and pistillate on sep-
arate plants, 2—8 in small clusters, each flower subtended by bracteole;
perianth segments of perfect flowers about 4 mm long, pale brown, glossy,
indurate, the outer broadly lanceolate, obtuse to acutish, with a broad scari-
ous margin, often subtended by a minute auricle, the inner tepals shorter,
acute to obtuse at the scarious-margined apex; stamens, 6 nearly equaling
the perianth, the anthers 1.5—2.5 mm long, about five times longer than stout
filaments; stigmas yellow or light brown; capsule subglobose, obtuse, mucro-
nate, 5 mm or less long, the valves rigid, equal or shorter than the perianth,
seeds from perfect flowers obvoid, acute to slightly tailed at one end, finely
reticulate to striate, pale yellow to light brown to brown, 0.3—0.6 mm; peri-
anth segments of pistillate flowers same as above, except shorter, 3 mm long,
reddish brown with a short staminoid at the base of each tepal, stigmas much
exceeding the perianth, red; capsule about 5.5 mm long, much exceeding the
perianth; seeds from pistillate flowers obvoid, acute to slightly tailed at one
end, finely reticulate, brown to dark brown, 0.4—0.6 mm

Tidal marshes from Delaware to south Florida and westward to Texas. An-
thesis occurs from January to June in North Carolina, the period becoming
shorter southward and restricted to March and April in extreme south Flori-
da. Westward from northern Florida into Texas, anthesis occurs from Janu-
ary to April. Seeds are shed about four weeks after the flowers reach ma-
turity.

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ee ane y J. W. 1903. The plant formations of eastern Massachusetts. Rhodora

BUCHENAU. F. 1906. Juncaceae. In Engler. Das Pflanzenreich. 25 (1V.36):1—284

CHAPMAN, A. W. 1860. Flora of the couchiern United States... Ist ed. Cambridge, Mass.
493p.

897. pee of the southern United States. Cambridge, Mass. 3

CORR PUL, D.S. & B. COR R ELL, 1975. Aquatic and wetland pants ae Seite scons
United States. Sanford University Press, Stanford, California. 177

__ _ & MC. JO HINSTON, 1972, Manual of the Texas flora. Texas Research Founda-
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COVILLE, F. V. 1894. eee Mem. Torr. Bot. Club 5:105—108.

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DAVIS, P. H. & V. H. HEYWOOD. 1967. Principles of angiosperm taxonomy. D. Van
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ELEUTERIUS, L. N. 197 4. = naia| study of Juncus roemerianus Ph.D, dissertation,
Mississippi State University.
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merlanus, eeeen 39:103—10
= ; 74. Flower morphology and plant types within Juncus roemertanus. Bull.
Mar. Sci, 2 Hes :493—497.
rae 1975. The life history of the salt marsh rush, Juncus roemerianus. Bull. Torry
Bot. Chib oe 35—140.
6a. The distribution of Juncus roemerianus in the salt marshes of North
pees “Cheng Sei. 17 (4) :289—292.
6b, eee jae ae and anatomy of the salt marsh rush, Juncus
roemerianus, Cale Res. Reports 5(2):1—10.
- . In review a. Pertlation structure of the gynodioecious salt marsh rush, Juncus
roemerianus.
In review b. Transplantation of the salt marsh rush, Juncus roemerianus to
apenas sail,
. In review c. Sex distribution in the progeny of the gynodioecious rush, Juncus
roeme ae 5.
ENGELMANN, G. i 1868. A. revision of — North American species of the genus

Jus eae s. Acad. . St. Louis 2:424—
FE RNALD, M. L. 1950. oon s manual of a 8th Ed. American Book Company, New
ork. 767p.

GLEASON, H. A. 1952. Illustrated flora of the northeastern United States and adjacent
Canada. The New York Botanical Garden. Hafner Publishing Co., Inc. New York.
1

GRANT, V. 1975. Genetics of flowering plants. Columbia University Press, New York.
5
JONES, F. B. 1975. Flora of the Texas coastal bend. Misson Press, Corpus Christi, Texas.

2p.
LAMARCK, J. B. 1789. veer salsa la Botanique.
LONG, R

& O. LAKELA. 1972. a of the flora of he Florida. University of
Miami Press. ie — er ae
RADFORD, A. HLES, & C. “7 BELL. 1968. Manual of the area flora of
the Carolinas. Univ ren oa. Press. Chapel Hill, North origi 11 :

SCHEELE, A. 1849, Junceae. 1. Juncus roemerianus. Linnaea 22:3

SMALL, J. K. ed oes of the southeastern United States. cae 256p
1933. Flora of the southeastern United States. North Carolina ee Chapel
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os IMARK, 1946. Studies in wide ae os reference to the species in Ethiopia
and the ae Svensk. Bot. Tidskr. 131—

ADDITIONS TO THE VASCULAR FLORA
OF OKLAHOMA

R. JOHN TAYLOR AND CONSTANCE E. TAYLOR
Biology Department, Southeastern Oklahoma State University
Durant, Oklahoma 74701

ABSTRACT
Forty-seven new taxa are reported to the Oklahoma flora and specimens of
four additional taxa are cited which verifies their present occurrence in the
state. The fifty-one taxa are as follows: Lycopodium adpressum, Potamoge-
ton natans, Cenchrus echinatus, Panicum verrucosum, Eleocharis smallit,
Eleocharis tortilis, Scirpus supinus, Rhynchospora gracilenta, Rhynchospora

capitatus, Melanthium ‘virginicum, Burmannia capitata, Habenaria repens,
Spiranthes odorata, Glinus radiatus, Paronychia Drummondii, Tillaea aqua-
tica, Parnassia grandifolia, Aeschynomene indica, Amorpha paniculata, Co-
lutea Fo caus Geranium dissectum, Polygala cruciata, Bowlesia incana,
Eryngium in tegrifoli ium, Cynoctonum sessilifolinm, noe quadriflora,
Phlox eee Scutellaria integrifolia, Linaria canadensis var. canadensis,

Bacopa monnieri, Epifagus virginiana, ee ee Utric ~ularia subt-
lata, Aster ptarmicoides, Cirsium muticum, Euthamia graminifolia, Euthamia
leptocephala, Evax candida, Onopordum acanthium, Solidago arguta var.
Boottii, Solidago es: Solidago gigantea var. gigantea, Solidago patula
var. strictula.

The following taxa are thought to be newly reported to the Oklahoma flora.
Four of these taxa are listed as occurring in the state by other sources, but
no specimens were cited or no specimens can be located in Oklahoma her-
baria. Further, they are not treated as occurring in our flora or they are
treated as doubtful in the most recent manual (Waterfall, 1969). All specimens
cited were collected by John and Connie Taylor unless otherwise indicated,
and are deposited at the Southeastern Ok'!ahoma State University Herbarium
(DUR). Duplicates of most have been sent to SMU, NELU, VDB, and KANU.

Most of these species were collected from eight hillside bogs in Atoka,
Chectaw, McCurtain, and Pushmataha counties in southeastern Oklahoma.
These bogs have developed mainly on sandy formations of the old Gulf Coast-
al Plain. They form along the flanks of very small drainage systems near
their upper ends, and some of them contain several feet of peat. The bogs
range in size from about an acre to over 40 acres. They may be forested,
but grasses, sedges, and other herbaceous species compose the vegetation of
most of them. There are 20 of these bog habitats known to the authors in
southeastern Olahom

The remainder of a selec were collected from various habitats through-
out the state. Most of the taxa occur in the Gulf Coastal Plain area of Texas
and represent extensions of their known range northward and westward into

SpA 7 S61 ser.

362

Oklahoma. We have relied on three manuals for nomenclature: Correll &
Johnston, 1970; Correll & Correll, 1972; and Steyermark, 1963.

LYCOPODIUM ADPRESSUM (Chapm.) Lloyd & Underw. ATOKA CO.:
edge of bog, 0.5 mi NW of Boehler, 26 Mar 1975, J. & C. Taylor 18217; and
bog 0.5 mi SW of Boehler, 10 Jun 1976, J. & C. Taylor 21709. This species is
very similar to L. alopecuroides and it has been treated as a variety of that
species.

POTAMOGETON NATANS L. MCCURTAIN CO.: Yashaw Creek, 2.5 mi
S of Broken Bow, 20 Jul 1976, J. & C. Taylor 22750; PUSHMATAHA CO.:
small creek 2 mi SE of clayton and about 130 m W of US hwy. 271, 9 Aug
1976, J. & C. Taylor 23047. This species is listed for Choctaw Co., Oklahoma
by Correll & Correll (1972) but we found no Oklahoma specimens in our state
herbaria.

CENCHRUS ECHINATUS L. MCCURTAIN CO.: edge of blind lake along
the Red River, 3.5 mi E and 8.5 mi S of Idabel, 2 Aug 1976, J. & C. Taylor
22837. This is a weedy species of the coastal plain.

PANICUM VERRUCOSUM Muhl,. CHOCTAW CO.: wooded bog, 4 mi NE
of Swink, 4 Sep 1975, J. & C. Taylor 20445. This species is similar to P.
brachyanthum which grows in similar habitats, but can be differentiated
from that species by its smaller spikelets and lack of pubescence. Both pa
cums are species of the southeastern United States.

ELEOCHARIS SMALLII Britt. CHOCTAW CoO.: collected from a sandy
well drained roadside 2 mi E of Hugo, 10 May 1969, J. Askew 116. The species
was determined by Barney Lipscomb. It is a wide ranging species of the
northeastern United States, west to Illinois and southern Missouri.

ELEOCHARIS TORTILIS (Link) Schult. ATOKA CO.: bog about 0.5 mi
SW of Boehler, 10 Jun 1976, J. & C. Taylor 21711. This species was determined
by Barney Lipscomb. It is another plant of the coastal plain area of the
United States.

SCIRPUS SUPINUS L. var. HALLII (Gray) Gray. ATOKA CO.: edge of
a shallow water filled sandy depression in an open forest, 0.25 mi S of Boeh-
ler, 23 May 1975, J. & C. Taylor 18541. The very small form of Echinodorus
tenellus grew with this species along the edge of these depressions. Species
was determined by Barney Lipscomb. Scirpus saximontanus, which also oc-
curs in Oklahoma, has been treated as a second variety of this species of
the coastal plain area.

SHOSPORA GRACILENTA Gray. CHOCTAW CO.: bog 11 mi NW of
Hugo, 22 Aug 1976, J. & C. Taylor and J. R. Wright 23235. PUSHMATAHA
CO.: bog 3 mi S and 7.5 mi W of Antlers, 23 Jul 1976, J. & C. Taylor 22774.
This is a species of bogs and similar habitats of the coastal plain area of the
United States.

RHYNCHOSPORA RARIFLORA (Michx.) Ell. ATOKA CO.: from a bog,
0.5 mi SW of Bochler, 10 Jun 1976, J. & C. Taylor 21710; PUSHMATAHA CO.:
bog on the north side of the county road, 3 mi S and 7.5 mi W of Antlers,

363

23 Jul 1976, J. & C. Taylor 22773. The range and habitat is similar to the
preceding species.

SCLERIA MUHLENBERGII Steud. ATOKA CO.: bog 0.5 mi NW of Boehler,
13 Aug 1976, J. & C. Taylor 23202; PUSHMATAHA CO.: bog 5.5 mi W of
Antlers, 1 Sep 1976, J. Taylor 23387; CHOCTAW CO.: bog 11 mi N of Hugo,
22 Aug 1976, J. & C. Taylor and J. R. Wright 23238, and from a bog 11.5 mi
NW of Hugo, 2 Oct 1976, J. & C. Taylor 23803. This distinctive member of
the genus Scleria is another species of coastal plain bogs and similar habi-
tats. In bogs of Atoka and Pushmataha counties, this species, the two Rhyn-
chosporas above, and Dulichiwm arundinaceum grow in mixed stands.

SCLERIA VERTICILLATA Muhl. CHOCTAW CO.: bog 11 mi W of Hugo,
22 Aug 1976, J. & C. Taylor and J. R. Wright 23236. This coastal plain species
is equally rare in Texas (Correll & Correll, 1972) and in Missouri (Steyer-
mark, 1963), but seems to be most abundant in the coastal plain habitats
throughout the eastern half of the United States.

CAREX DECOMPOSITA Muhl. ATOKA CO.: edge of a bog 0.5 mi NW of
Boehler, 6 May 1975, J. Taylor 18447. At this location it grew in a few inches
of standing water with Hottonia inflata and Peplis diandra. This relatively
large Carex of the eastern United States is reported to be rare (Correll &
Correll, 1972).

CAREX LEPTALEA Wahlen. ATOKA CO.: bog 0.5 mi SW of Boehler,
15 May 1976, J. & C. Taylor 21305; CHOCTAW CO.: bog 4 mi NE of Swink,
2 Aug 1976, J. & C. Taylor 22879. At the Atoka Co. location C. leptalea grew
at the edge of clumps of Osmunda cinnamomea, O. regalis, Lorinseria areo-
lata and Onoclea sensibilis. These four fern species occur in most of the bogs
we have studied, the last two found as far west as Marshall County.

CAREX SWANII (Fern.) Mack. MCCURTAIN CO.: floodplain forest along
the west fork of the Glover River, 0.5 mi W of Battiest, 16 May 1972, J. & C.
Taylor 10375. This species was determined by Dr. David Castaner. Its habitat
is moist forests of the northeastern United States.

WOLFFIA PUNCTATA Griseb. ATOKA CO.: open water of a bog 0.5 mi
NW of Boehler, 1 Nov 1976, J. & C. Taylor 23965. It was found growing with
Azolla caroliniana, Limnobium spongia, and species of Lemna and Spirodelia.
Correll & Correll (1972) list McCurtain Co., Oklahoma as part of its range,
but we found no Oklahoma specimens in state herbaria.

ERIOCAULON DECANGULARE L. PUSHMATAHA CO.: bog on the south
side of a county road, 3 mi S and 7.5 mi W of Antlers, 7 Jul 1976, J. & C. Tay-
lor 22410; and a bog 5.5 mi W of Antlers 22 Aug 1976, J. & C. Taylor 23252.
At both of these locations it was found growing with the small, rare, and
endangered FE. kornickianum, and with Peltandra virginica and Iris virginica.

JUNCUS CAPITATUS Weigel. MCCURTAIN CO.: along the moist, sandy
roadside, 0.5 mi S of Moon, 29 Apr 1975, J. Taylor 18405. Species determined
by Barney Lipscomb. This small Juncus is an introduction from Europe. It
is reported to be rare in Texas where it is infrequently collected.

364

MELANTHIUM VIRGINICUM L. CHOCTAW CO.: from a bog 11 mi NW
of Hugo, 22 Aug 1976, J. & C. Taylor and J. R. Wright 23213. These plants
were about 4 feet tall and grew with Spartina pectinata, Verbena hastata,
and Veronicastrum virginicum.

BURMANNIA CAPITATA (Walt.) Mart. PUSHMATAHA CO.:: in bog 5.5 mi
W of Antlers, 1 Sep 1976, J. Taylor 23390. This species was growing among
grasses and sedges with such species as Selaginella apoda, Eriocaulon kor-
nickianum, and Bartonia paniculata.

HABENARIA REPENS Nutt. ATOKA CO.: bog at edge of beaver pond,
0.5 mi NW of Boehler, 26 Oct 1974; J. & C. Taylor 17262. This is a large
bog orchid of the coastal plain area in the United States.

SPIRANTHES ODORATA Nutt. CHOCTAW CO.: bog 11.5 mi NW of Hugo,
2 Oct 1976, J. & C. Taylor 23799. Species determined by Dr. Larry Mcgrath.
While treated as a variety of S. cernua by Correll (1950), Leur (1975) feels
that this taxon should be considered a separate species

GLINUS RADIATUS (R. & P.) Rohrb. MCCURTAIN CO.: Big Grassy
Lake (presently being drained) 4.5 mi E, 3.4 mi S of Tom, 2 Aug 1976,
J. & C. Taylor 22871. Species determined by Wm. F. Mahler.

PARONYCHIA DRUMMONDII T. & G. ATOKA CO.: in an open sandy
upland forest 130 m from the Boehler intersection, 6 May 1975, J. & C.
Taylor 18437. The soil at this location is derived from the Antlers Sand
Formation, with a very light, sandy, topsoil. The forest is composed of
widely spaced trees with Quercus incana, Sassafras albidum, and stands of
Vaccinium arboreum being abundant. The vegetation of the open areas is
sparse and composed mainly of low herbaceous species. This type of habi-
tat is very common on the Antlers Sand Formation of Atoka, Choctaw
Pushmataha Counties. Some of the species that grow with P. pate s
at the Atoka Co. location are as follows: Selaginella riddellii, Triplasis pur-
purea, Aristolochia reticulata, A. serpentaria, Eriogonum multiflorum, Poly-
gonella americana, Leoflingia squarrosa, Polanisia erosa, Crotonopsis line-
aris, and Penstemon murrayanus.

TILLAEA AQUATICA L. ATOKA CO.: edge of shallow, water filled sandy
depression in an open forest, 0.25 mi S of the Boehler intersection, 23 May
1975, J. & C. Taylor 18542A.

PARNASSIA GRANDIFOLIA DC. CHOCTAW CO.: abundant in bog 11
mi NW of Hugo, 22 Aug 1976, J. & C. Taylor and J. R. Wright 23223; 14 Sep
1976, J. & C. Taylor 23596; and 2 Oct 1976, J. & C. Taylor 23797.

AESCHYNOMENE INDICA L. MCCURTAIN CO.: edge of a rice field
near the dam of Barney Ward Lake, 1.5 mi SW of Tom, 19 Oct 1974, J. & C.
Taylor 17208. Growing with this species was Cuscuta attenuata and Ipomoea
wrightii.

AMORPHA PANICULATA T. & G. MCCURTAIN CO.: grazed field east
of Barney Ward Lake, about 1.3 mi SW of Tom, 4 Jun 1976, J. & C. Taylor
21590.

365

COLUTEA ARBORESCENS L. CIMARRON CO.: rocky draw about 0.5
mi E of Kenton, population covering about one acre, 19 Jun 1976, J. Taylor

2040.

GERANIUM DISSECTUM L. MCCURTAIN CO.: edge of a rice field near
the dam of Barney Ward Lake, 1.5 mi SW of Tom. 29 Apr 1975, J. & C. Tay-
lor 18413.

POLYGALA CRUCIATA L. PUSHMATAHA CO.: bog 5.5 mi W of Antlers,
22 Aug 1976, J. & C. Taylor and J. R. Wright 23243.

BOWLESIA INCANA Ruiz. & Pav. COTTON CO.: base of Tamarix gallica
growing in floodplain of the Red River just W of the US hwy. 277 and 281
bridge, SW of Randlett, 2 May 1976, J. & C. Taylor 21166.

ERYNGIUM INTEGRIFOLIUM Walter. Reported by Goodman (1974)
based on E. J. Palmer 8998 (MO). We have collected it from PUSHMATAHA
CO.: bog on west side of road, 10.5 mi N and 1 mi W of Soper, 4 Sep 1975,

Taylor 20461; and bog 5.5 mi W of Antlers, 1 Sep 1976, J. Taylor
23379, CHOCTAW CO.: bog 11 mi NW of Hugo, 14 Sep 1976, J. & C. Taylor
603

3603.

CYNOCTONUM SESSILIFOLIUM Walter ex J. F. Gmelin. ATOKA CO.:
bog 0.5 mi NW of Boehler and 20 mi SE of Atoka, 13 Aug 1976, J. & C. Taylor
23200.

LYSIMACHIA QUADRIFLORA Sims. CHOCTAW CO.: bog 11 mi NW of
Hugo, 22 Aug 1976, J. & C. Taylor and J. R. Wright 23237; and J. & C. Tay-
lor 23614, 14 Sep 1976. Ray (1956) lists this species for Arkansas.

PHLOX PANICULATA L. MCCURTAIN CO.: southwest bank of the Mt.
Fork River about 8 mi NE of Smithville, 21 Oct 1976, J. Taylor 23958. Listed
by Waterfall (1969) as seen only from cultivation.

SCUTELLARIA INTEGRIFOLIA L. MCCURTAIN CO.: from 2.7 mi N of
Tom, 4 Jun 1976, J. & C. Taylor 21605. The plants were growing with Eu-
stylis purpurea.

LINARIA CANADENSIS (L.) Dum. var. CANADENSIS. ATOKA CO.::
sandy soil near the edge of a bog 0.5 mi NW of Boehler, 6 May 1975, J. & C.
Taylor 18450, and from an open sandy upland forest 130 m S of the Boehler
intersection, 23 May 1975, J. & C. Taylor 18521. This taxon may be easily
distinguished from L. texana (lL. c. var. texana) by its smaller purple flow-
ers and in our area always in very sandy soil.

BACOPA MONNIERI (L.) Penn. MCCURTAIN CO.: along the 1941 cut-
off blind lake of the Red River, 3.5 mi E and 8.5 mi S of Idabel, 2 Aug 1976,
J. & C. Taylor 22826. It was found growing with Stenotaphrum secundatum.

EPIFAGUS VIRGINIANA (L.) Bart. MCCURTAIN CO.: southwest bank
of the Mt. Fork River about 8 mi NE of Smithville, 21 Oct 1976, J. Taylor
23960. Two clumps were found growing under a large beech tree. It has
been known previously close by in Arkansas.

UTRICULARIA INFLATA Walt. ATOKA CO.: very abundant in a shallow
water filled sandy depression in an open forest 0.25 mi S of the Boehler

366

intersection, 6 May 1975, J. & C. Taylor 18433. An additional sheet of this
species was found in our herbarium from MCCURTAIN CO.: swamp, 2.5
mi S of Eagletown, 17 Jun 1967, Johnson 90.

UTRICULARIA SUBULATA L. ATOKA CO.: bog 0.5 mi NW of Bochler,
6 May 1975, J. Taylor 18449; PUSHMATAHA CO.: bog 5.5 mi W of Antlers,
1 Sep 1976, J. & C. Taylor 23392. This small terrestrial species was growing
among Lycopodium adpressum at the Atoka location and with Burmannia
capitata and Bartonia paniculata in Pushmataha County.

ASTER PTARMICOIDES (Nees) T. & G. JOHNSTON CO.: established
on the Baum Limestone, 2.9 mi NW of Mansville, 7 Sep. 1973, J. & C. Taylor
14856. At OKL, two additional sheets were found in the Aster innomen folder
from the same location: Perino & Massey 846, 13 May 1971; and J. Williams
525 (marked A. ptarmicoides), 4 Sep 1971. This species was not present in
the area in 1960 when the senior author was researching the Grassland
Communities on the Baum Limestone (Taylor & Penfound, 1961). Anderson
and Creech (1975) believe this species is more closely related to Solidago
than Aster.

CIRSIUM MUTICUM (Michx.) Pers. CHOCTAW CO.: bog 11 mi NW of
Hugo, 22 Aug 1976, J. & C. Taylor and J. R. Wright 23229; and 14 Sep 1976,
J. & C. Taylor 23609.

EUTHAMIA GRAMINIFOLIA (L.) Nutt. CHOCTAW CO.: bog 11 mi NW
of Hugo, 22 Aug 1976, J. & C. Taylor and J. R. Wright 23216; PUSHMATAHA
CO.: bog 5.5 mi W of Antlers, 1 Sep 1976, J. & C. Taylor 23370. Very isolated
and disjunct from its distribution in the northern and eastern United States,
the closest population occurring in northern Missouri

EUTHAMIA LEPTOCEPHALA (T. & G.) Greene. ‘CHOCTAW CO.: at
Unger between US hwy. 70 and the railroad track, 5 Nov 1971, J. & C. Tay-
lor 9416; MCCURTAIN CO.: Barney Ward Lake, 2 mi W of Tom, 22 Jul
1972, J. & C. Taylor 11121. We include the citation of these specimens to
clarify the status of this species in Oklahoma. Waterfall (1969) includes
this taxon as part of the E. gymnospermoides complex. Sieren (1970) and
Taylor (1975) offer conclusive evidence of its distinctness. At the Unger
location, both FE. leptocephala and E. gymnospermoides were growing in-
termixed together. Occupation of the same habitat has been noted both in
Texas and in Louisiana, but no evidence of intergradation has been ob-
served.

EVAX CANDIDA (T. & G.) Gray. ATOKA CO.: sandy open forest about
130 mS of the Boehler intersection, 23 May 1975, J. & C. Taylor 18523.

ONOPORDUM ACANTHIUM L. ROGER MILLS CO.: roadside about 2.5
mi W of Strong City, 18 Jun 1976, J. Taylor 21868. This weedy thistle is
known from Kansas and Texas, so its occurrence in Oklahoma was to be
expected. It is a native of Europe.

SOLIDAGO ARGUTA Ait. var. BOOTTIL (Hook.) Pal. & Stey. Not current-
ly considered a part of the Oklahoma flora, but reported as early as 1900 as

367

S. Boottii by Bogue. Solidago arguta was used in Van Vleet’s list in 1902.
Our specimen J. & C. Taylor 14928 from LEFLORE CO. north flank of Blue
Bouncer Mt. along a branch of Eagle Fork Creek, 11 mi NE of Octavia,
18 Sep 1973, documents this taxon as valid member of the Oklahoma

ora. It has since been collected from TCCuEAIN CO. along Mt. Fork
River, Beavers ae State Park, 18 Sep 1976, J. & C. Taylor 23628; CHERO-

E CO.: stream below McSpadden Falls, 6 Sep 1976, J. & C. Taylor 23401.
It has been seen and identified elsewhere in the Ouachita Mts. and Ozark
Plateau of Oklahoma. It should not be considered as rare.

Bogue had annotated his S. Bootii “Frequent in dry shaded situations,”
the name evidently being misapplied to S. ulmifolia (S. delicatula) which is
very common throughout east central Oklahoma in the dryer blackjack-
post oak forests. The var. Boottii is found in the moister habitats of stream
bottoms in oak-hickory forests and blooms later in the fall, after the late
summer flowering of S. ulmifolia is nearly terminated.

SOLIDAGO FLEXICAULIS L. MCCURTAIN CO.: along the west bank
of the Mt. Fork River, 8 mi NE of Smithville, 21 Oct 1976, J. & C. Taylor
2

3961.

SOLIDAGO GIGANTEA Ait. var. GIGANTEA. MCCURTAIN CO.: road-
side ditch, 2.8 mi N of Tom, 14 Sep 1974, J. & C. Taylor 16996; CHOCTAW
CO.: bog 11 mi NW of Hugo, 22 Aug 1976, J. & C. Taylor 23226. The pu-
bescence on the stems and leaves separates this variety from the very
common glabrous stem and leaved S. g. var. leiophylla.

SOLIDAGO PATULA Muhl. var. STRICTULA T. & G. MCCURTAIN CO.:
disturbed roadside with windrows of bulldozed trees, along hwy. 87, 2 mi S
and 0.4 mi W of Tom, 18 Sep 1976, J. & C. Taylor 23641.

ACKNOWLEDGEMENTS

We are grateful to Wm. F. Mahler and Barney Lipscomb, Southern Meth-
odist University Herbarium, for their assistance. Mr. Lipscomb accom-
panied the senior author on field trips. Dr. Larry Magrath accompanied us
on field trips and assisted in some species determinations. Dr. J. R. Wright
flew the senior author over southeastern Oklahoma, which resulted in loca-
tion of three of the larger bogs. Some of the field work was supported by
NIH Grant No. 5506RR08005.

REFERENCES

ANDERSON, L. C. and J. B. CREECH. 1975. Comparative leaf anatomy of Solidago and
related eee: Amer. J. Bot. 62:486-493

BOGUE, E. E. 1900. An annotated catolog . thie ferns and flowering plants of Oklahoma.
Oklahoma Agric. <s Sta. Bull. 45. 48

CORRELL, D. S. 1950. Native Orchids of Toni America north of Mexico. Chromica Bo-
tanica Co., Waltham, Mass

CORRELL, D. S. and H. B. CORRELL. 1972. ae and wetland ao of southwestern
United States. Environmental Protection Agency, Washington

CORRELL, D. S. and M. C. JOHNSTON. 1970. ee of the ae plants of Texas.

Texas Research Foundation, Renner, Texas

368

GOODMAN, G. J. 1 . Notes on Oklahoma plants. Proc. Oklahoma Acad. Sci. 54:98-99.
LUER, C. A. 1975, whe native Orchids of the United States and Canada pete Florida,

New York Botanical Garden, New York.
a 24:1-160.

RAY, J. D. 1956. The genus Teytiiae hia in the New World. a Biol.
STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Universit ress, Ames.
taxonomic revision of the genus Exthamia. Ph. Dissertation,

Uni-

SIEREN, D. J. 197
versity of Illinois, Urban

TAYLO E. 1975, Fatt &ymnospermoides (Compositae). Ph.D. Dissertation. Uni-
ae oF Ohlaome. Norm

TAYLOR, R. J. and W. T _ PENFOUND, 1961. The grassland communities on the Baum
nee in Johr fe ee Oklahoma. Southw. Naturalist 6:98-

VAN VLEET, A. H. 2 hn of Oklahoma, Terr. of Okla. Dept. Geol and Nat. Hist.

2 Keys ¢ to the flora of Oklahoma. Published by the author, Okla-

homa State Cae, Sdillware

TAXONOMY AND GEOGRAPHY OF
NICOLLETIA (COMPOSITAE: TAGETEAE)

JOHN L. STROTHER
Department of Botany, University of California, Berkeley 94720

Nicolletias are easily distinguished from other Tageteae by pinkish to
magenta stripes or bands on the laminae of the white to pink ray corollas, a
peculiar pappus of fascicles of coarse bristles alternating with and subtend-
ing lanceolate, awn-tipped squamellae, and, apparently, a novel chromo-
some base number, x = 10. Geographic ranges of the three species are al-
lopatric and each is restricted to a single floristic province (Fig. 1): Nicol-
letia edwardsii in Chihuahuan Desert, N. occidentalis in Mojave Desert,
and N. trifida in Sonoran Desert of Lower California.

Position of Nicolletia within Tageteae is puzzling. Morphologically, its
nearest relative is Leucactinia Rydb., a little known monotype with white
to pale yellow ray corollas, a pappus of coarse bristles (the innermost some-
times narrowly hyaline-margined at base), and chromosome number un-
known. Leucactinia is known from only five localities in Nuevo Leon, Coahui-
la, San Luis Potosi, and Aguascalientes. Isolation of this pair of genera
within Tageteae probably reflects an early derivation from ancestral Tage-
teae (Strother, 1977).

CHROMOSOMES

All species of Nicolletia have been reported to have 2n — 10 II in four
to six populations of each. The base chromosome number for the genus is
taken to be x = 10. Base numbers for other genera of Tageteae are: 7, 8, 9,
11, 12, 18, and 15; base numbers of seven of the 16 genera remain unknown.

One population of N. trifida (30 miles south of San Filipe, Powell and
Turner 1739, DS, NY, US) has been reported to have n = 20. Additional
counts of 2n = 10 II from N. occidentalis are reported here: CA, Los An-
geles Co., ca. 11 and 12 miles east of Palmdale, Strother 1221 and 1222, UC.

TAXONOMY
NICOLLETIA A. Gray n Frémont, Se exped. Rocky Mts. p. 315. 1845.
T

Annuals or perennial herbs; stems ae to erect, 5—30 (—50) cm
high at anthesis; glabrous, often glaucous. Leaves mostly alternate (lower-
most opposite); pinnately divided into 3—9 (0—11) stout to linear-filiform
lobes, these usually bristle-tipped and bearing a subterminal, orangish, pel-
lucid gland containing strongly scented oil. Heads terminating branches,
subsessile or pedunculate. Involucres turbinate to fusiform or campanulate;
12—18 mm high. Calyculus of 0—6 short, lanceolate to triangular bractlets.

SIDA:7 (4): 369—374.

370

Phyllaries 8—12; free to base; imbricate; membranous, narrowly scarious-
margined; most bearing 1—5, oval to linear, pellucid glands; persistent and
becoming reflexed. Receptacle convex to hemispheric or conical; favose to
alveolate; sparsely and minutely hispid to glabrous. Ray florets 8 (7—12);
pistillate, fertile; corollas white to pink, laminae usually with bands or
stripes of magenta or pink, very showy, apically retuse or shallowly 3-lobed.
Disc florets 15—100-+; perfect, fertile; corollas pale to bright yellow, some-
times tipped with purple, narrowly cylindrical, 7—9 mm long, tube much
shorter than the throat, lobes 5, deltoid to lanceolate; anthers minutely
sagittate; style branches long, slender, tipped with filiform, hispidulous,
astigmatic appendages. Achenes slender, somewhat clavate, blackish, close-
ly pubescent with short, white to rufous hairs. Pappus of 5 (—6) fascicles
of 7—15 fine to coarse, hispidulous bristles subtending and alternating with
5 (—6) lanceolate, awn-tipped squamellae.

Key to Nicolletia
a. Leaves mostly 3—5-lobed, rachis scarcely wider than lobes; laminae of
ray corollas 7—16 mm long; disc florets mostly 15—50 per hea
b. Leaf lobes mostly patent; peduncles 5—30 mm long, heads held scarce-
ly above foliage; laminae of ray corollas a 9—16 = long; Chi
huahuan Desert. 1. N. edwardsii
bb. Leaf lobes mostly antrorse: “peduncles 10—70 mm Fea heads held
well above foliage; laminae of ray corollas 7—11 mm long; Lower

California. 2. N. trifida
aa. Leaves mostly 7—9- lobed, ‘rachis mostly 2-3 times as wide as —
laminae of ray corollas 4—8 mm long; disc florets —— 60—100-+ p

head; Mojave Desert. : : 3. N. secitoniae
1. NICOLLETIA EDWARDSII A. Gray, ag eee Contr. Knowl. 3(5):119
(Pl. wright. pt. 1) 1852. Type: Mex Chihuahua, ‘‘near Guajuquilla”

(present Cd. Jimenez, near 27°N, 105° W), Aug, Dr. Edwards s.n. Holo-
: NY!

Annuals (or short-lived perennials); spreading, mostly 5—25 cm_ high,
weakly succulent, sometimes weakly glaucous. Leaves 1—6 cm long, pin-
nately parted into 3—5 linear to filiform, usually patent lobes (or simple
and filiform), sometimes with additional short, lateral lobes. Peduncles

rowly turbinate to weakly fusiform. Calyculus of 4—6 triangular bractlets
1—2 mm long. Phyllaries mostly 6—8, lanceolate, acute 10 attenuate, 11—

8 (7—9); tube of corolla slender, 4.2—6.2 mm long; laminae linear to nar-
rowly elliptic, 7—16 mm long, 3—6 mm wide; style branches exserted
0.1—1.1 (—2.2) mm. Disc florets 15—25 (—50); corollas creamy yellow, 7—8
mm long, tube 1.5—1.7 mm long, throat narrowly cylindrical, ca. 5 mm
long, lobes 0.4—0.7 mm long, deltoid to lance-triangular, papillose and
sparsely glandular-puberulent; anthers ca. 3.5 mm long, including basal
collar (0.3—0.4 mm long) and apical appendage (0.4—0.5 mm long); style

371

branches 2.5—2.7 mm long, including appendage (1.7—2.2 mm long). Achenes
5—7 mm long. Pappus bristles 3—5 mm long; squamellae 4.8—7.2 mm long,
including awn (1.8—3.3 mm long).

Distribution (Fig. 1): Chihuahuan Desert from trans-Pecos Texas south
and east to northern Zacatecas. Locally common to abundant on sandy to
silty, oe Ean or gypseous, soils on flats or playas in Larrea Scrub;
8 m. Flowering mostly Aug—Oct, following summer rains; occasion-
al through the year (Fig. 1).

2 NICOLLETIA TRIFIDA Rydb., N. Amer. FI. 34:180. 1915. Type: Mexico,
Baja California, Bahia de los Angeles (ca. 29°N, 113°35’W), Apr—Dec
1887, E. Palmer 569. Holotype: NY!; isotypes: GH!, NY!, UC!, US!

Annuals or short-lived perennials: ascending to stiffly erect, 1—3 (—5)
dm high, often succulent and glaucous, Leaves 3—5 (1—7) cm long, pin-
nately parted into 3 (0—5) linear-terete, antrorse (30—45°) lobes. Peduncles
2 (1—7) cm long, heads usually held well above foliage. Involucres narrowly
turbinate to somewhat fusiform or campanulate, Calyculus of 0—2 lance-
triangular bractlets 1—2 mm _ long. Phyllaries 8 (7—10), lance-linear to
cblanceolate, attenuate or abruptly acute, 12—16 mm long, 2.5—3.8 mm
wide, very thin, hyaline-margined, each usually bearing a conspicuous sub-
apical gland and, rarely, with 2—4 lateral, linear glands. Ray florets 8
(—11); tube of corollas 4.8—$.5 mm long; laminae ovate-elliptic, 7—11 mm
long, 2.5—5.2 mm wide; style branches exserted 0.8—2.2 mm. Dise florets
30—50 (—80); corollas yellow, 7.2—8.9 mm long, tube 1.8—2.7 mm long,
throat 4.5—5.1 mm long, gradually dilated, lobes 0.9—1.1 mm long, narrowly
triangular, papillate, nct glandular-puberulent; anthers 3.5—3.9 mm _ long
including collar (0.4—0.6 mm long) and apical appendage (0.6—0.8 mm
long); style branches 3.1—3.8 mm long, including appendage (1.1—1.5 mm
long). Achenes 5—8 mm long. Pappus bristles 3—6 mm _ long, very fine;
squamellae 4.2—7.7 mm long including awn (0.7—2.2 mm long).

Distribution (Fig. 1): Sonoran Desert portion of Lower California between
24° and 30°30’ N; 0—1300 m. Flowering mostly Feb—May and Oct—Nov,
following rains; occasional through the year (Fig. 1).

Achenes of N. trifida seem eminently suited for disperal and much of
western Sonora and the islands of the Gulf of California seem to provide
ample hospitable habitats, yet N. trifida has been reported from only one
of the islands (Isla Inez, very near the coast of Baja California Sur at ea.
27° N, I.M. Johnston 3654, CAS, GH) and not at all from Sonora.

3. eee OCCIDENTALIS A. Gray in de Rep. exped. Rocky
Mts 316. 1845. Type: California, ‘“‘banks of Mohahve river, growing
in rata sands,’ Apr 1844, Frémont ‘‘358’’. ease. NY!; ‘isotypes:

H!, NY!

Perennials; strictly erect from deep taproots; 12—29 cm high; glaucous.
Leaves at very base (soil level and below) reduced to scales, normal leaves
2—7 cm long, pinnately parted into 7—9 (5—11) short, patent or antrorse

372

a N. occidentalis
N. trifida
N. edwards:

Fig. 1. Distributions of Nicolletia spp. Histograms indicate flowering times as determined
from cumulative herbarium records.

373

lobes, fleshy and often crowded below the heads. Peduncles mostly 2—10
mm long, heads surpassed by foliage. Involucres broadly turbinate to cylin-
drical. Calyculus of 2—4 lanceolate to triangular bractlets 4—8 mm long.
Phyllaries 8 (—12), linear to ovate, abruptly acute to acuminate, 14—18 mm
long, 3.5—5.5 mm wide, somewhat fleshy, narrowly scarious-margined, often
tinged with purple, each bearing 0—1 subapical and 0—4 submarginal
glands. Ray florets 8—12; tube of corollas 5.5—6.5 mm long; laminae nar-
rowly ovate-elliptic, 4.5—8.5 mm long, 2.5—3.8 mm wide: style branches
exserted 2.8—3.8 mm. Disc florets (30—) 60—100+; corollas yellow, often
tipped with purple, 8.2—9.5 mm long, tube 2.2—3.0 mm long, throat 4.8—5.5
mm long, lobes 0.8—1.1 mm long, deltoid, papillate, not glandular-puberu-
lent; anthers 3.8—4.5 mm long including collar (0.4—0.5 mm long) and
apical appendage (0.7—0.8 mm long); style branches 3.2—3.5 mm long, in-
cluding appendage (0.9—1.2 mm long). Achenes 7—9 mm long. Pappus bris-
tles 3—7 mm long; squamellae 6—8 mm long, including awn (1—2 mm long).

Distribution (Fig. 1): California, Mojave Desert from northeastern Kern
Co. south into northeastern Los Angeles Co. and east and south across
western San Bernardino Co. into north central Riverside Co. (Joshua Tree
National Monument). Locally common in deep sands, often in floors of
washes; 600—1400 m. Flowering mostly Apr—Jun, following winter rains
(Fig. 1).

The often cited occurrence of N. occidentalis in San Diego Co., California,
is apparently based on mislabelled specimens (S.B. and W.F. Parish 193).
Some of these (GH, MO, US) read ‘‘sandy banks of Mojave River, May
1882.’’ On others (DS, F, NY, PH, US), printed labels read ‘‘San Diego Co.,
San Felipe’’ and either May or June, 1882. Two of the latter have the print-
ed locality struck through and ‘‘Mojave R”’ (NY) or ‘‘Mojave Desert’? (US)
written in. I have seen no other collections attributed to San Diego Co.

GEOGRAPHY

Distances separating geographic ranges of species pairs in Nicolletia are
considerable: edwardsii/occidentalis—1000 km, edwardsii/trifida—650 km,
and occidentalis/trifida—400 km (Fig. 1). Historical development of this dis-
tribution pattern invites speculation. The ancestral population of Nicolletia
probably resided in central Mexico, center of diversity for Tageteae (Stroth-
er, 1977). But, how did N. trifida get to Lower California? How did N. occi-
dentalis get to Mojave Desert? Long distance dispersal and colonization fol-
lowed by differentiation in response to selection? Migrations of ancestral
stock and subsequent differentiations and extinctions?

Comparative morphology of Nicolletias offers little help in reaching a
conclusion, in making a choice between alternative hypotheses. No single
species seems to be markedly more ‘‘primitive’’ or ‘‘archetypal’’ than the
others. In Dyssodia subg. Clomenocoma, however, there is a group of spe-
cies that form a definite, linear, morphological sequence from primitive to

374

advanced (Strother, 1969). Geographical distribution of this series of species
has implications for interpreting the biography of Nicolletia.

In subg. Clomenocoma, Dyssodia aurantia and D. appendiculata are mor-
phologically primitive and closely approximate a hypothetical archetype for
Dyssodia in particular and Tageteae in general. Dyssodia aurantia is re-
stricted to Veracruz; D. appendiculata ranges across southern Mexico from
Chiapas to Colima. Dyssodia squamosa is closely related to D. appendiculata
and ranges northwest of the latter from Jalisco through Nayarit into Sina-
loa, Next in this series is D. speciosa, which is restricted to southern Baja
California Sur. It is followed by D. porophylloides, which ranges from mid-
peninsular Lower California into California, Arizona, and Sonora. The series
ends with D. cooperi, which ranges north from D. porophylloides in Califor-
nia and Arizona into southern Nevada (Strother, 1969). This morphological/
geographical sequence in Dyssodia correlates closely with a continuum of
increasing aridity of habitats (Walter and Lieth, 1967). Dyssodia appendicu-
lata occupies rather mesic habitats and the habitat of D. cooperi is the most
xeric of the series.

Nicolletias cannot be placed so readily into a linear evolutionary sequence,
nor do they cover such a broad spectrum of habitats. Still, a history of mi-
gration from the Chihuahuan Desert region into the Sonoran and Mojave
deserts followed by differentiations and extinctions seems to be a plausible
explanation for present distributions of these species.

ACKNOWLEDGMENTS
For loans of specimens or assistance during visits, I thank curators and
staff of the following herbaria: A, ASU, BM, CAS, DS, E, F, GH, JEPS, K,
LL, MICH, MO, NO, NY, PENN, PH, POM, RM, RSA, SD, SMU, TEX,
UC, and US

REFERENCES
STROTHER, J.L., 1969. a of Dyssodia Cavanilles (Compositae: Tageteae). Univ.
Calif. Publ. Bot, 48:1—8
. 1977. Ta oe ee natic review : He Paes ee V.H., et al., eds. The biology
and Sa of the Compositae. London ee mic Pre
WALTER, H. and H. LIETH. 1967. Klimadiagramm——Weltatlas. Jena: G. Fischer.

CHROMOSOME NUMBERS IN ERIGERON
AND CONYZA (COMPOSITAE)

GUY L. NESOM
Department of Botany, The University of North Carolina
Chapel Hill, N.C. 27514

Though reports of chromosome numbers in the Compositae are extensive,
the genus Erigeron has received relatively little attention. Of the approx-
imately 147 currently recognized North American species north of Mexico,
only about 66 have been studied cytologically, and infraspecific variation in
chromosome number is known in about 26 of these. The counts reported
here are from collections made mostly during the last three years and are
peripheral to a more inclusive study of Erigeron.

Chromosome numbers are presented for 74 collections representing 1
species of Conyza and 23 of Erigeron; 9 are for previously unreported taxa
and 1 differs from an earlier report for the same taxon. Counts were made
either from buds fixed in the field with Farmer’s solution (acetic alcohol
3:1; Sass, 1958) or from radicles grown from achenes in the laboratory,
pretreated for 4 hours in 8-hydroxyquinoline, and fixed in Farmer’s solution.
Vouchers are deposited at NCU or at Jacksonville State University, Ala-
bama; duplicates are at TEX and DUKE. Thanks go to Miklos Treiber for
collecting buds and vouchers of several species and to David Whetstone,
from whose Alabama collections achenes of E. strigosus, E. annuus, and C.
canadensis were taken.

Table I. Species of Erigeron and Conyza examined for chromosome number.

Taxon Chromosome Locality and Collection
number 2n
CONYZA:
C. canadensis (L.) 18 (1)+ ALABAMA: Cullman Co., just SE
Cronq. of Oak Level. Whetstone 4845.
18 (8) ALABAMA: Marshall Co., Gunters-
ville. Whetstone 6335.
9. LOUISIANA: LaSalle Parish, Olla.
Nesom R313.
9. NORTH CAROLINA: Jackson Co.,
Cashiers. Nesom R309.
ERIGERON:
E. annuus (L.) Pers. 27 (1) ALABAMA: DeKalb Co., 5 mi S of

t. Payne. Whetstone 3520.

SIDA 7(4): 375—381.

376

*K, basilobatus Blake
E.. bellidiastrum Nutt
ar. bellidiastrum

FE. eatoni A. Gray subsp.
eatoni

E. eximius Greene

*K, formosissimus Greene
var. formosissimus

*E, Pea leg Greene
s (Rydb.)

Feet Etowah ie Walnut
rove. Whetstone 409
pela Franklin a 1 mi S of
Russellville. Whetstone 4474.
ALABAMA: Walker Co., 4 mi W
of Jasper. Whetstone 4728.
NORTH CAROLINA: Orange Co.,
Chapel Hill. Nesom R319.
VIRGINIA: Pittsylvania Co.,
Gretna. Treiber 1476.
MEXICO; NUEVO LEON: Cerro
otosi, 8 mi WNW of Galeana.
Nesom R566
MEXICO; NUEVO LEON: Pena
Nevada, ca. 35 E of Doctor
Arroyo. Nesom R574.
TAH: eld C 11 mi E of
Escalante. Nesom R38
EW MEXICO: Torrance Co., 19
mi SE of Willard. Nesom R656.
COLORADO: Mesa Co., 7 mi § of
Mesa. Nesom R11B.
COLORADO: Grand ee 5 mi § of
Grand Lake. Nesom R.
COLORADO: Canson 2s Sapi-
nero. Nesom
NEW MEXICO: Taos Co., 1.5
E of Taos a Valley. Nesom RSI.
NEW MEXICO: Bernalillo Co., E
side of Sandia Peak summit Ne-

, som R669.

NEW MEXICO: Taos i 9 mi SE
of Rodarte. Nesom R68

NEW MEXICO: Sandoval Co., 22
mi W of Los Alamos. Nesom R241.
NEW MEXICO: Rio Arriba 2
mi W of Tres Piedras. Nesom R672.

NEW MEXICO: Sandoval Co., 18
mi W of Los Alamos. Nesom R240.

NEW MEXICO: Colfax Co., 11 mi
SSW of Eagle Nest. Nesom R213A.
NEW MEXICO: Bernalillo Co., 6
mi NW of Sandia Peak. Nesom
R665

NEW MEXICO: Rio Arriba Co., 10
mi NW of Tres Piedras. Neco
R676

*F’. leiomerus A. Gray

E. lonchophyllus Hook.

"Ei, palmeri A. Gray
E. peregrinus (Pursh)

Greene subsp. calli-

E. philadelphicus L.

E. pinnatisectus (A.
ray) A. Nels.

E. platyphyllus Greene

*F. potosinus Standley

E. pulchellus Michaux

377

NEW MEXICO: Valencia Co.
Spellenberg 4912.

COLORADO: Chaffee Co., 7 mi E
of Buena Vista. Nesom R314.

MEXICO, NUEVO LEON: NE of
Pena Nevada, 39 mi E of Doctor
Arroyo. Nesom Ro78.

NEW MEXICO: Taos Co., ca. 6 mi
of Red River. Nesom R680.

LOUISIANA: tect Parish,

Natchitoches. Nesom R32

NORTH CAROLINA: Bu im Co., 6

mi S$ of Jonas Ridge. Treiber 1411.

res CAROLINA: kel ae
mi FE of Boone. Treiber

Sune tee ee

Co., Carlisle. Treiber 1478

COLORADO: Clear Creek Co.,
Guanella Pass. Nesom R258.

NEW MEXICO: Lineoln Co., Sierra
Blanca Ski Area, Nesom R654.
NEW MEXICO: Otero Co., 9 mi
NE of Cloudcroft. Nesom R650.

MEXICO, NUEVO LEON: Cerro
otosi, 8 mi WNW of Galeana.

Nesom R567.

MEXICO, NUEVO LEON: Cerro
Potosi, 8 mi WNW of Galeana.

Nesom R568.

NORTH CAROLINA: Alleghany
Co., Blue Ridge Parkway, 1 mi S
of jet with Hwy 18. Treiber 1445.
NORTH CAROLINA: Buncombe
Co., Blue Ridge Parkway, 2 mi
SEE of Asheville. Treiber 1432.
NORTH CAROLINA: Jones Co.,
Island Creek, 6 mi S of New Bern.
Nesom R315.
NORTH CAROLINA: eae i
Co., Blue Ridge Parkw 1.5 mi
SW of ject with Hwy 80. Treiber 1429,
NORTH CAROLINA: Montgomery
1 mi WNW of Uwharrie. Ne-
som R316.

378

E. pumilis Nutt. subsp.

concinnoides Cronq.
I. rusbyi A. Gray

E.. strigosus Muhl. ex
Willd.

*E. subtrinervis Rydb.
subsp. subtrinervis

NORTH CAROLINA: oe Co.,
Chapel Hill. Schram
VERMONT: Windso oan. 2 mi W
of Woodstock. Treiber 1466.
COLORADO: Gunnison Co., 10 mi
NE of Almont. Spongberg s.n
NEW MEXICO: Otero Co., 1 mi §$
of Cloudcroft. Nesom R644.
ALABAMA: Marshall Co., 7.3 mi
E of Douglas. Whetstone 3218.
ALABAMA: Morgan Co., 3 mi N
of Hulaco. Whetstone 2916.
ABAMA: Morgan Co., Falkville.
Whetstone 2995.
ALABAMA: Tuscaloosa Co., Holt
2 mi NE of Tuscaloosa. Whetstone
4666.
ALABAMA: Winston Co., 6 mi ESE
of Haleyville. Whetstone 4279.
ALABAMA: Winston Co., Haley-
ville. Whetstone 4499.
FLORIDA: Liberty Co., 5 mi E of
Torreya State Park. Nesom R325.
KENTUCKY: Hopkins Co., e mi N
of Madisonville. Nesom R30
MISSISSIPPI: Harrison a
Gulfport. Nesom R324,
TEXAS: Kaufman Co., 24 mi E of
Dallas. Nesom R317.

COLORADO: Alamosa Co., Great
Sand Dunes Natl. Mon. Nesom
248.

COLORADO: Park Co., 1 mi E of
Weston Pass. Nesom R288.
COLORADO: eee Co., 4 mi E of
Grant. Nesom
COLORADO: — Co., 8 mi SW
of Lake George. Nesom R272.
NEW MEXICO: Bernalillo Co., E
side of Sandia Peak summit. Ne-
som R667.
NEW MEXICO: Colfax Co., 0.5 mi
ESE of Red River Pass. Nesom
BY

W MEXICO: Rio Arriba Co.,
San Antonio Peak, 16 mi N of Tres
ie m R247.

NEW MEXICO: Taos Co., 11 mi E

379

of Taos. Nesom R207.

94 NEW MEXICO: Taos Co., 3 mi SW
of Taos Ski Valley. Nesom R223.
“BE. tenuis T. & G, a: LOUISIANA: Natchitoches Pari
1 mi W of Natchitoches. Nesom
18,
36 (1) MISSISSIPPI: es Co., Wave-
land. Nesom R32
tHE. utahensis A. Gray 18 (1) UTAH: Garfield Co., 2 mi E
Tropic. Nesom s.n. (no voucher).
*F. vetensis Rydb. Se COLORADO: Grand Co., 5 mi S
of Grand Lake. Nesom R354.
E. vernus (L.) T. & G. 18 (4) GEORGIA: Worth oF 7 mi N of
Sylvester. Nesom R32
18 (1) MISSISSIPPI: Han ‘ee Co., 3 mi
W of Waveland. ecoa R322.
a MISSISSIPPI: Pearl River Co.,

Picayune, Nesom R323.

* First report for the taxon.

+ First cee of this chromosome number for the taxon.

+ Numbers in parentheses after mitotic counts refer to the number of radi ae for which
the see ee number was ae All radicles for each collection were grown from
achenes of a single head.

DISCUSSION

The first counts for Erigeron formosissimus (n=9, n=18) are given here.
There are no apparent characters in the tetraploid plants from Bernalillo
Co., N. M., which would distinguish them from known diploids of this taxon.
However, immediately adjacent to the diploid clone of E. formosissimus var.
viscidus from Colfax Co., N. M. (Nesom R213A), a clump of the same spe-
cies (Nesom R213B) was Peciieatad which had produced pollen with extreme-
ly low stainability and much irregularity in size, indicating that it is prob-
ably polyploid. This putative polyploid clone has typical glandularity on the
phyllaries and upper peduncle but differs from the adjacent, more typical,
var. viscidus in having a noticeably more strigose involucre.

The tetraploid counts for Erigeron eximius (=E. superbus Greene, see
Weber, 1973) in the Sangre de Cristo Range are interesting in that the pre-
vious tetraploid report for this species is from the Front Range in Boulder
Co., Colo. (Love and Kapoor, 1967). A diploid count has been reported from
Cochise Co., Ariz., by Watson (1973). The presence of tetravalents in the
Bernalillo Co., N. M., plants suggests that they may be of autoploid origin.

Erigeron potosinus is Known from Cerro Potosi in southern Nuevo Leon
and from two additional localities in Tamaulipas, Mexico. It appears to be
extremely closely related to FE. eximius, and the two may eventually prove
to be conspecific. Plants of both taxa produce herbaceous rhizomes, but

380

those of E. potosinus do so more prolifically and produce more dense clonal
colonies. Both collections of EF. potosinus reported here were made on the
ENE side of Cerro Potosi in a subalpine meadow with scattered pines at
about 3200-3300 meters in elevation, below the summit of about 3650 to 3820
meters (Beaman and Andresen, 1966). The second collection (R568) was
made about 100 meters below the first (R567). From the first collection tetra-
ploid counts were obtained from two plants; in both of them multivalent as-
sociations were observed and pairing ranged from 8,,’s + 2,,'s to 18,,’s
(though some loose pairing may have been present in the latter cell). Pol-
len from eight other plants from this locality was found to be over 96%
abortive. Tetrads were formed most commonly, though some pentads were
noted, but micrograins were commonly formed with the tetrads to incor-
porate individual laggard chromosomes not reaching either pole. The more
normal sized pollen grains were malformed and mostly devoid of cytoplasm.
Gross meiotic abnormalities do not appear to account for the high pollen
sterility in these tetraploids. From the second collection of E. potosinus
(R568) clear diploid counts were obtained from three plants with meiosis
and tetrad formation normal in each. Of seven other plants from this lo-
cality from which pollen has been examined, two have over 90% viable
pollen and are probably diploid. The other five have from 95% to 35% abor-
tive pollen; micrograins commonly occur in three of these plants and all
five are probably polyploid.

Lack of morphological distinction between the diploids and_ tetraploids
of Erigeron potosinus, other than perhaps a slight size difference, and the
presence of multivalents in the tetraploids are strong circumstantial evi-
dence that autoploids are being formed on Cerro Potosi. The only other spe-
cles of Erigeron seen in this area were E. basilobatus Blake and EF. cf.
nudiflorus Buckley; both were growing at a much lower elevation than BP.
potosinus and are morphologically distinct. The two collections of FE. poto-
sinus known from Tamaulipas are Stanford, Taylor, and Lauber 2501A
(TEX and SMU) and Stanford, Taylor, and Lauber 2673 (SMU). Gross ir-
regularities in the pollen examined of two plants from each of these locali-
ties indicate that they are probably polyploid.

Both collections of Erigeron basilobatus (n=27,,, n=36,,) have noticeably
aborted pollen, although pairing was apparently normal in all the cells ob-
served. Pollen from an isotype of this species (Muller 2934, SMU) showed
a lowered viability, indicating that these plants are probably also polyploid.
Muller’s collection was made in Nuevo Leon as were the two reported here.

The first chromosome counts for Erigeron tenuis (n=9,,, n=18,,) are pre-
sented here. Besides the tetraploid count near the southeastern extremity
of the range of the species in Hancock Co., Miss., four other plants exam-
ined from Oktibbeha, Pearl River, and Rankin Cos., Miss., are also pro-
bably polyploid—an estimate based on their production of extremely abor-
tive pollen. Of these four only the collection from Rankin Co. (Jones 18591.2)
is atypical of the species in any apparent way; on the herbarium label

381

(NCU) it is noted by Dr. Jones as being unusual and has exceptionally large
basal and lower cauline leaves.

The discovery of diploid Erigeron strigosus from Mississippi, Alabama,
and Florida confirms the report by Turner and Flyr (1966) of diploid LE.
strigosus from Florida and establishes this southern area as a center of
sexual populations for the species. Checks of pollen size and stainability
from other locations in Alabama show that diploids also occur in Franklin,
Blount, and Cullman Counties. Numerous other chromosome counts over
the range of this widespread colonizer have been polyploid. The production
of relatively high percentages of achenes with euploid chromosome numbers
different from that of the megasporangiate parent is a phenomenon being
documented in greater detail in Erigern flagellaris A. Gray, Erigeron diver-
gens T. & G., and their relatives (Nesom, in progress). Erigeron strigosus
apparently hybridizes freely with E. annuus, but plants of the latter species
have not yet been found at a level other than triploid.

REFERENCES
BEAMAN, J. H. and J. W. ANDRESEN. 1966, The yee ee and phytogeog-
Sys )e

raphy of the summit of Cerro Potosi, Mexico. Am. Midl.
CRONQUIST, A. ue Sas of the North Remar species 7 Erigeron, north of

eae Brittonia 6: -302.

LOVE, A. and B. M. KAPOOR. 1967. In IOPB chromosome number reports XIV. Taxon
16: 552- 7
ASS, E 958. Botanical Microtechnique. 3rd ed. Iowa State U. Press, Ames

STADE 5 L. and D,. Flyr. 1966. Chromosome numbers in the one X. North
American species. Amer. J. Bot. -33

d W. A. 1973. age to “the Caloride flora, V, with nomenclature revisions.
Southw. Nat. 18: 317-3

WATSON, T. J. 1973. Chromosome numbers in Compositae from the southwestern United
States. Southw. Nat. 18: 117-124.

A SYNOPSIS OF NORTH AMERICAN
CORISPERMUM (CHENOPODIACEAE)
NITA J. MAIHLE? AND WILL H. BLACKWELL, JR.
Herbarium, Department of Botany, Miami University
Oxford, Ohio 45056

ABSTRACT
Corispermum (Chenopodiaceae) is an Eurasian genus of annual herbs
containing members which have become established in the North American

flora. The taxonomy of the eo estar: of this genus occurring in North
America has not received careful attention for some time, The most recent
taxonomic revision of North American Corispermum was actually that of
P. C. Standley (1916) in which the following species were recognized: C.
ae alee L., C. nitidum Kitaibel ex Schultes
nd C. villosium. in ydbere. Standley considered the first two species to be
introduced and the latter two native. In the present study, herbarium speci-
mens were examined from 14 different herbaria. On the basis of literature
and Gila pa of these herbarium specimens, only three species of Coris-
permum, all introduced, are recognized in North America: C. hyssopifolium
C. nitidum Kitaibel ex Schultes, and C. orientale Lamarck. A study of
distr ibutional data acquired from herbarium specimens and state floras sug-
gests that the geographic range for all three species has increased with time.

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INTRODUCTION AND METHODS

Corispermum Soe a genus of branched, annual herbs, is
not native to N America. However, it has become well established in
the North pie flora. It is commonly found on dunes and sandy hills,
beaches or sand flats, and along roadsides and railroad ballasts. The flowers
are small, inconspicuous, and arranged in spikes located in axils of the
bracts. The ‘‘bug-like’’ fruit, exposed in the bract axils, is often a promi-
nent feature.

The genus is sometimes confused with several other chenopodiaceous taxa
Kochia scoparia (L.) Schrader resembles Corispermum in its habit, but the
calyx of Kochia is lobed not more than half-way to the base and forms
dorsal, membranous wings in fruit. The calyx of Corispermum is scarious,
lobed to the base, and does not develop wings. Salsola kali L. is also super-
ficially similar to Corispermum, but has a spirally coiled embryo with no
endosperm, and the tips of the leaves and bracts are commonly spinulose.
The embryo of Corispermum is annular, surrounding copious endosperm,
and the leaves and bracts, though narrowed, are seldom spinulose.

"This work was supported in part by a grant (W. H. Blackwell, Principal Investigator)
from the American Philosophical Society (No. ae abit Fund) and from the Willard

Sherman Turrell Herbarium Fund of Miami a aae SILY
esent address: Department of Botany, Ohio State cece srsity, Columbus, OH 43210.

SIDA 7(4): 382—391.,

383

The genus Corispermum contains 50-60 species, 34 of which (along with
several varieties) are included in Flora of the U.S.S.R. (ljin, 1936). Al-
though North American taxa of Corispermum are few in number, they have
not received careful taxonomic attention for some time. A brief summary
of the literature relevant to taxa found in North America will serve to sup-
port this poi

Schultes a. first published Corispermum nitidum, which had earlier
been described, but not validly published, by Kitaibel. Several superfluous
names were subsequently given to this species, including C. tenue Link
(1820) and C. microspermum Host (1827); however, no North American vari-
ants have been proposed.

Corispermum hyssopifolium was described by Linnaeus (1753). In 1818
Nuttall published a North American variety of this species, C. hyssopifolium
var. americanum. Later, he elevated this variety to the status of species,
viz. C. americanum Nuttall (1834). In 1838 W. J. Hooker added several other
North American varieties, including C. hyssopifolium var. gracile, C. hysop-
ifolium var. robustius, and C. hyssopifolium var. rubricaule. Watson (1874)
urther increased the list of North American taxa with C. hyssopifolium
var. microcarpum.

Around the turn of the century Rydberg described three supposedly new
North American species: Corispermum villosum in 1897, C. marginale in
1903, and C. emarginatum in 1904. Macbride (1918) suggested that C. villo-
sum Rydberg and C. emarginatum Rydberg actually represented the same
taxon which he considered to be merely a variety of C. orientale (an Old
World species), i.e. C. orientale var. emarginatum (Rydberg) Macbride. A
few ae after eee descriptions, two additional North American spe-

ere sed: C. imbricatum Nelson (1909) and C. simplicissimum
Sit ae ee (1909) di ssed the introduction of Corispermum in
North America and eicaled ae a. distribution at that time was ‘‘through-
out the middle west.’

The most recent revision of the North American taxa of this genus was
that of Standley (1916), who recognized four species. These species and
those names that Standley considered their synonyms are as follows: 1) Cor-
ispermum hyssopifolium: C. hyssopifolium var. americanum, C. american-
um, C. marginale, C. imbricatum, C. simplicissimum,; 2) C. nitidum: C tenue,
C. microspermum, C. hyssopifolium var. microcarpum; 3) C. emarginatum;
and 4) C. villosum. The first two species he thought to be introduced, while
the latter two he considered native. Standley also mentioned C. pilosum,
first described by Rafinesque (1836), but treated it as a doubtful species.

Standley’s work, while admirable for its time, leaves many questions un-
answered. Herbarium specimens are frequently misidentified, and the num-
ber of taxa which should be recognized, as well as the rank which should
be accorded them, is uncertain. Furthermore, the native versus introduced
status of various taxa is subject to question, and the geographic distribution

384

given for each species in 1916 has become outdated. The present study is
thus an updated taxonomic revision of the North American species of Coris-
permum, including a reconsideration of each species’ geographic range.

The conclusions of this study as to the number of taxa of Corispermum
in North America and their taxonomic rank are based on literature and on
personal observations of herbarium specimens. Measurements and other
data included in the descriptions are from dried specimens. Over 950 her-
barium specimens were examined, including both New and Old World ma-
terial. The type specimens of most of the proposed North American taxa
were examined. In all, specimens were studied from 14 different herbaria.
Abbreviations are after Holmgren and Keuken, 1974: F, GH, MIN, MO, MU,
NY, OSC, OSU, PH, RM, TEX, UC, US, and UT. The curators of these her-
baria are gratefully acknowledged for the loan of specimens of Corispermum.

After a study of previous descriptions, the species descriptions herein
were written to emphasize those specimens examined which represented the
broadest range of morphological variation. Type citations (for all types
seen) are included with the synonyms (pertinent to North America) follow-
ing the correct name of each species.

By surveying state and regional floras and examining labels of herbarium
specimens, a more current geographic range was established for each taxon.
Specimens were selected for citation on the basis of this distributional sur-
vey. They are thus representative of the range of Corispermum in North
America as it is now known. A card file will be maintained for all specimens
examined. Each specimen’s card typically includes information on the fol-
lowing: name of specimen, herbarium from which it was received, accession
number, collector, collection number, date of collection, and habitat.

GENERIC DESCRIPTION
CORISPERMUM L., Species Plantarum 1:4. 1753.

Branched annual herbaceous plants, frequently slightly woody, glabrous
to pubescent, the pubescence of stellate hairs; height of mature plant highly
variable in response to climatic and edaphic factors. Leaves green (more
rarely tinged with purple), alternate, simple, linear, sessile, entire, often
deciduous early, those of the inflorescence reduced to bracts (which are
slightly scarious-margined). Flowers perfect, axillary, arranged in spikes;
perianth segments sepalloid, hypogynous, whitish, scarious, 1—3 (when 3
the posterior segment largest, the 2 anterior ones smaller); stamens 1—3,
hypogynous, becoming exserted; ovary superior, the stigmas 2. Fruit achene-
like, strongly exserted from perianth, flattened, oval to rounded-oval, often
convex on the dorsal surface and concave on the ventral surface; pericarp
somewhat hardened and closely surrounding the seed, the margin winged
or not. Seed vertical; embryo annular, surrounding copious endosperm.
Flowers and fruits developing late summer through autumn.

Type species: Corispermum hyssopifolium L., Species Plantarum 1:4,
753

385

The name Corispermum is derived from the Greek words coris bedbug,

and sperma, seed. Vernacular names include: bugseed, bugweed, and tick-
seed.

SPECIES OF CORISPERMUM IN NORTH AMERICA

There are three species of Corispermum in North America; all three have
been introduced. They are regarded as introduced species on the basis of
their similarity to Old World material, the time of their recorded appearance
in the North American flora, and their ruderal pattern of distribution. State-
ments in the literature also support their introduced status. Distributional
data accumulated in this study indicate, in comparison with previously avail-
able information, an increasing range for all three species of Corispermum
in North America with time.
A. Fruit une cuously wing-margined.

B. Spikes dense; bracts closely overlapping and all at least as broad a

the

C.h BS onienim
B. eons loose: bracts not distinctly overlapping, especially along lower-
ost portion of spike, and at least the lower bracts much narrowe

rane the fruit. nitidum

A. Fruit lacking conspicuous wing- margins. oo 5 See =) C. orientale

1. pe ae [pean L., Sp. Pl. 1:4, 1753,
Lin n Herbarium, IDC No. 12!

americanum Nuttall, Trans. a Philos. Soc. 5:165. 1834.

Hee te ar. americanum Nattall, The Genera of North Amer-
n Plants 1:3. 1818. (Type: Nuttall s.n., PH!, labeled C. hyssopi-

rm i var. pubescens)

hyssopifolium var. gracile Hooker, Flora Boreali-Americana 2:125.

1838

(Phototype:

aa

hyssopifolium var. robustius Hooker, Flora Boreali-Americana 2:125.
1838.

hyssoptfolium var. rubricaule Hooker, Flora Boreali-Americana 2:125.
1838

Q OQ 2 2

imbricatum A. Nelson, Coulter’s New Manual of Botany of the Central
Rocky Mountains, revised ed., 163. 1909. (Type: Elias Nelson 733,
M!

a

marginale peat Bull. Torrey Bot. Club 30:247. 1903.
Herrick s.n.,
C. simplicissimum “Lune Amer. Midl. Naturalist 1:207. 1910.
Lunell 395,

(Holotype:
(Type:

Stems slender to stout, glabrous to pubescent, the pubescence sparse to
very dense. Leaves green, broadly linear, 0.5—9.0 cm long, 1.0—5.0

mm
wide. Bracts linear to ovate, closely

overlapping, at least as wide as fruit,
as long as or longer than fruit, narrowed at the apex. Spikes densely flow-
ered, up to 9 cm long, the spike axis not exposed. Fruits 1.0—4.0 mm wide,
strongly wing-margined, the wing pale, up to 0.5 mm wide.

The specific name is derived from the Greek hyssopos, an aromatic herb.

Type locality: along the Volga River, U.S.S.R

The North American specimens of Corispermum hyssopifolium do not

386

differ distinctly from Old World specimens of this species. Also, previous
descriptions of C. hyssopifoliwum from both North America and the Old World
are quite similar. Thus, the various specific and varietal names which have
been recognized as applying to distinct taxa are reduced to synonyms.

Representative specimens: CANADA: ALBERTA: Rees tare railway gravel, 23 Sep
1939, Groh 1099 (TEX, UC); Cardston, gravel soil, Lee’s Creek, 8 Aug 1914, Mooi 206
(NY); Craigmyle District, in grain crop, 30 Aug i Ue 5326 (OSC, RM); Fort
Saskatchewan, 4 Sep 1939, Turner os (GH). MANITO BA: Glenboro, 35 | SE of
Brandon, prairie sandhills, 24 Aug 1951, Scoggan 10545 (GH); Sidney, three mi E of here,
5 Sep 1954,Rossbach 78 (UC). peice sl eat Bay District, railroad ballast, north
shore of Lake Superior, 13 Aug 1937, Hosie, Losec, and Bannan 1461 (GH); Rainy River
District, on railroad ballast, prostrate, 15 Aug 1961, Garton 9438 (MIN); Welland Co.:
Point Abino, 23 Aug 1886, Coville sn. (MO). QUEBEC: Abitibi Co.: Villemontel, railroad
ballast, 31 Aug 1952, Edldwe and Breitung 4320 (GH); Labelle Co.: 8 Oct 1949, Giroux
sam. (MU). NORTHWEST TERRITORIES: Mackenzie District, disturbed sand along road-
side on ee one mile south of Fort Simpson Island, 3 Aug. 1955, Cody and Matte
9130 (F, GH, MIN); Mackenzie District, five miles upstream from Norman Wells, rare
on gravel beach . nage River, 27 Jul 1953, Cody and Gutteridge 7603 (F); Mac-
kenzie District of Fort —— windblown sand along river channel, 22 Aug
1939, Ae ond Soper 9868 ne RM

EX of ana Quarry ails 1912, Stearns sin. (MO).

ane aay : ALASKA: Porcupine ie 15 mi from Cun, 22 Aug 1926,
Murie 49 (US) ; Yukon River between Rampart and Tanana, 9 Aug 193 = eae 13 (US).
ARIZONA: Navajo Co.: Holbrook, 4 Oct ees Zuck sn. (US). cares IFORNIA: Inyo

Co.: scattered on lower slopes of sand dunes at mouth of Marble Canyon, 13 May ;
ue 6361 (UC). COLORADO: Jackson Co.: San Luis Valley, 1875, Brandegee s.n. (UC);

rimer Co.: North Park, 5 Sep 1899, Osterbout 3201 (RM); Moffat Co.: three mi NW
e Sunbeam, sand Bs out, 2 Sep 1970, Weber 14246 (US, UT); Weld Co.: 1909, Johbns-
ton 618 (RM). AHO: Kootenai Co.: near Horse Plains Mountain, 25 Aug 1895,
Leiberg 1602 ‘Mo, NY, UC); Nez Perce Co.: sandy island in Clearwater River, op-
posite Lewiston, 12 Oct 1924, St. Jobu 6792 (NY, UC). ILLINOIS: Cook Co.: a.

—_~

Sep 1888, sat sm. (UC); Lake Co.: Winthrop Harbor, 30 Aug 1909, Gates 3226
(F). INDIANA: Howard Co.: Kokomo, 1942, Ek sn. (GH, MO, MU, NY, TEX,

La Porte Co.: Micl City, 30 Sep , Benke 2749 (F); Porter Co >den Diines,
4 Oct 1930, Standley 57479 (F). IOWA oe Co.: moist sandy alluvium alo is-
sourt River, 9 Aug 1952, Fay 4703 (UC). CHIGAN: Berrien Co.: in sandy soil close to

Lake M Michigan in Bridgman, 20 Sep 1952 eee 1891 om eelauna Co.: on open and ex-
posed places, 13 Sep 1917, McCoy 1037 (ys Manistee Co. Manistee, in sand on lake shores,
21 Aug 1882, Morong s.n. (NY); Mason Co.: sand-flats at Ludington, shore of Lake Michi-
gan, 2 Oct. 1949, McVaugh 11198 (MO). MINNESOTA: Polk Co.: dry sand about one mi
SW of Fertile, 22 Aug 1947, Terrell 1827 (M S); Recs Co.: ballast of railroad
one and three tenths mi N of Longworth, 2 Aug 1939, Veove and Moore 11370 (MIN,
NY, UC); St. Louis Co.: Duluth, Minnesota Point, 6 Sep 1936, Lakela 1841 (FP, ;
PH); Todd Co.: Staples, sandy soil, 29 Aug a. "Chon cuss! sa. (MO). MISSOURI:
Clay Co.: Quo River, rare, 21 Sep 1895, Mackenzie 1053 (NY); Perry Co.: Wittenburg,
sand dunes along Mississippi River flood plain, 31 Jul 1934, Steyermark 14056 (NY);
St. Louis Co.: Chain of Rocks, Missouri River sand dunes, 5 Oct 1933, Kellogg s.n. (MO).

T Cascade Co.: Great Falls, Aug 1895, Anderson s.n. (MU, NY); Dawson
Co.: sandy soil near Glendive, 6 Sep 1892, Sandberg, MacDougal and Heller 1013 (F,
GH, NY, US). NEBRASKA: Kearney Co.: Minden, sandy soil, 20 Sep 1907, Hapeman
saa, (MIN); Lancaster Co.: Lincoln, Aug 1898, Eat gi ate san. (PH). NEW JERSEY
Nassau Co.: Bayville, 7 Jul 1935, Murray s.n. NEW YORK: Erie Co.: Buffalo,
Clinton sn. (F, NY). NORTH DAKOTA: Varna Co.: sand blow-out, four to five
mi W of Denbigh, 13 Sep 1932, Hotchkiss 4505 (US); Pierce Co.: Rugby, railroad,
30 Jul 1912, Bergman 2588 (MIN, MO); Richland Co.: Leonard, sand dunes, 19 Sep

F, k,

1937, Stevens 79 NY); Stark Co.: a on river ban 18 Jul 1911, Bergman
1257 (F). OHIO: Huron Con freight yards, Jul Shuman 965.1 (OSU). OREGON:
Baker Co.: 28 Nov 1952, Wright sa. (OSC); cae Co.: Grants Pass, Sep 1887,

387

Howell sn. (OSC); Lake Co.: near Alkali Lake, 18 Jul 1933, Thompson 12,172 (F, GH,

O, NY, UC, US); Wasco Co.: along Columbia River opposite The Dallas, ie Evans
saa. (OSC). SOUTH DAKOTA: Fall River Co.: sandbars Cheyenne River, 12 Sep 1924,
Over 15943 (UC); Walworth Co.: sandbars Missouri River, 28 Aug 1921, Over sin.
(US). TEXAS: Webb Co.: Laredo, Rio Grande River, 1-20 Aug 1879, Palmer 1165
(MO). UTAH: Juab Co.: Lynndyl sand dunes, 8 Sep 1965, ioe and Moore 5127 (MIN,

NY); Millard Co.: juniper association, sand dune, 18 Sep 1926, Guarreff 1010 (RM).
WASHINGTON: Douglas C je F Crab and Wilson Le 25 Jul 1893, Sandberg
and Leiberg 309 (PF, GH, NY, S); Grant Co.: sandy coos plains near
Quincy, 15 June 1931, T bom pson 6764 (G GH, MO); Klickitat Co of th
Columbia River, 11 Sep, 2 ct 1893 Siege he e GH MIN M é

US); Whitman Co.: sandy bank of Snake River at Wawa 15 Oct 1939, Ownbey and

Ownbey sn. (MIN, NY, RM, TEX, UC). SG Keno Co.: moist prairie
along Lake Michigan, 16 Oct 1961, I/tis 19,461 (TEX); Milwaukee Co.: Milwaukee,
sandy lake beach, 1866, Lapham s.n. (MO); Racine Co.: Racine, 10 Sep 1878, Davis 41
(MIN, MON, OSU); Sherboyean Co.: sandy beach of ee Michigan 10 Sep 1956, Iltis and
Koeppen 8247 (UC). WYOMING: Sweetwater Co.: on sand dune one mi W of Steamboat
Mountain, 2 Sep 1936, Lang 99 (RM).

2. ene oy Kitaibel ex Schultes, Osterreichs Flora, re-
sed ed., 1:7. 1814.
C. hyssopifoliom var. microcarpum Watson, Proc. Amer. Acad. Arts
9:122. 1874. (Lectotype: Fendler 711, ie
C. microspermum Host, Flora Austriaca 1: 1827
C. tenue Link, Jahrbticher He aa 1:27. 1820.

Stems slender, glabrous or more rarely sparsely pubescent, the pubes-
cence denser in younger plants, Leaves green, narrowly linear, 0.5—6.0 cm
long, 0.5—3.0 mm wide. Bracts commonly not imbricate, exposing the spike
axis, only rarely slightly overlapping at apices or slightly telescoped in
immature plants, the lower bracts much narrower than the fruit, frequently
longer than fruit. Spikes not congested, up to 15 cm long. Fruits green to
black, 1.0—3.0 mm wide, strongly wing-margined, the wing pale, up to 0.3
mm wide

The specific name is from the Latin nitidus, bright.

Type locality: Hungary.

This plant is recognized in both the Old and New World as an excellent
source of forage for cattle. Many members of the Chenopodiaceae, including
Corispermum, are at least somewhat halophytic or salt-tolerant. As a re-
sult, this plant can establish itself in many areas where other species can-
net. It is especially common along roadsides and in waste plac

The selection of the lectotype of Corispermum hyssopifolium var. micro-
carpum Watson is made herein from among the syntypic collections cited
by Watson (1874).

Representative specimens: CANADA: henge eae Grande Clariete, W of Hartney,
dry sandy clearings in dune area, 29 Jul 1951, Scoggan 10142 (MIN); St. Lazare, 70 mi
N of Br eee in valley of Assini ieee River opposite mouth of Qu’Appelle River,
prairie sandhills, 7 Aug 1951, Scoggan 10246 (GH, MIN). ONTARIO: Rainy River Dis-
trict, on panne of sand dunes, ug 1961, Garton 9148 (UC); TI istri

ae pSEouD nd, staff parking lot Lakehead niversiey: 10 Aug 1972, Garton 15175 (UC).
XICO: State of eee sand dunes, 10-19 Oct 1935, LeSueur 285 (F, MO,
UC).

TE -

388

UNITED a basis rep teh ee Apache Co.: Adamana, 6-15 Aug 1903, Griffiths 5074

ty, Se { 1 ,N
Osterbout 1156 (RM). ILLINOIS: Cook Co.: Chicago, ve Sep 1954, Tiere 1033 (F);
M d Co.: Athens, 1861, Hall sn. (MO). INDIANA: Lake Co.: in sandy beach along
ee Deas 1746 (NY, RM, US); La Porte
lamin Jr. 1886 (F). ad Barber Co.: Medicine

1890 ge 300 (GH, NY); ee Co.: Tribune,
2 Sep 1893, Reed s.n. (MU, : US): Lo ogan Co.: sa ee aol Jul 1895, He :

oks Co.: Rockport, 1889-1894, habolime SM. yy

ICHIG
Lansing 2884 (GH); Calhoun Co.: Dp 1
Ludington, Hamlin Lake, moving dune, 14 Sep 1910, C
INNESOTA: Clearwater Co.: Squaw Lake, 25 Aug 1936, Grant 6830 (MIN); Goodhue
Co.: sandy soil Cannon Falls, Jul 1893, Scofield s.1. (GH); Hennepin Co.: Minneapolis,
Jul 1890, roe sm. (RM); Ottertail Co.: Battle Lake, Aug 1892, Sheldon s.n.
MO, OSU, UC). “NE BRASKA: Hooker Co.: 1
1893, Rydberg 1647 (NY, US); Kearney Co.: §
( di : Middle Loup River near Maric: on Paced bank, 11 Sep
Rydberg 1647 (NY, US). ae Legs Dona Ana Co. esilla Walley; 20 Sep 1907,
MO, OSC, RM); Gren ie dunes of quartz sands,
1 A

c Zt

S
Wooton and 1 ry 3194 (F,

14 m larosa, 13 Oct an ’ Brum s.a. (UC); San Juan Co.: Farmi
ot. Wooton. 2764 (US); Socorro Co.: 27 Aug 1965, Hess, Dunn and Spellman 419
along railroad track, 15 Sep 1921,

(GH). NEW YORK: Monroe Co.: tae Rochester
T

. NORTH DAKOTA: Ranson Co.: Anselm, 30 Aug 1918, Sfevens

Baster 5499 (GH,

Som N). OHIO: ae Co.: Chillicothe, B & O Railroad, 12 Sep 1965, Bartley 1078

( , OSU). OKLAHOMA: Canadian floodplain of tl outh Canadian Ri I

of Bridgeport, 6 Oct 1940, Waterfall 2 MO, NY); ales Co.: sand near river
); Woods Co.: on sandy

10 mi NW Norman, 8 Sep 1946, ee 4080 (MO, RM :
ar Alva, 24 Sep 1913, Peet 2849 (GH, MIN, MO, ,

bank of Sale Fork River 1

US). SOUTH DAKOTA: Clay Co.: sandy flood a of Re lion Riv 25 Sep 1914,

Over 5110 (US); Harding Co.: Cave , steppe, ep 1912, Visher 668 (F). TE

El Paso Co.: on dunes about 10 mi NW El af », W of Franklin Mountains, 30 Oct

1962, Correll 26570 (GH); Hemphill Co avel dikes in sand hills two mi S of Glazier,
deep sands of sand hills one mi S of

8 Oct 1965, Rowell 10919 (GH); Lipscomb Co.:
‘ , Rowell 10908 (GH); Wheeler Co.: flood plain, north fork of
River Sep 1950, Tharp and Miller 51-376 (MIN, RM, TE UTAH: Emergy Co.:
vicinity of Molly’s Castle, San Rafael Desert, 15 Aug 1957, Mitchell and Grover s.m.
Salt Lake Co.: Bingham Canyon, 27 Aug 1879, Jones 1341 MIN, NY, RM,
); i ie 15 Sep 1964, Welsh, Moore, and Olsen 3763 (MIN).
WASHINGTON: Grant Co.: dry sand SW of N June 1921, St. John, Court-
ney, and Parker 4948 (GH); Klickitat Co.: high sandy river bank,
3675 (GH); es Co.: woody bank of Snake River, 4 a 1924
WISCONSIN: Racine Co.: 10 Sep 1878, Davis s.n. (F heboygan Co.: occasional on
dunes and sandy bee hes, Lake Michigan area, 3 Sep 1939, * Pohl 1401 (PH).

3. Saar pen er ORIENTALE Lamarck, Encyclopédie méthodique bo-
que 2:110. 1786.

C. Aer od Rydberg, Bull.
Aven Nelson 4282, GH, MO,

—
io)
a
oO
a
—
*)
a

Torrey Bot. Club 31:404. 1904. (Type:
, RM!

Cc. — var. emarginatum (Rydberg) Macbride, Contr. Gray Herb.
new series, 53:12. 1
C; rene Rydberg, Bull. Torrey Bot. Club 24:191. 1897. (Lectotypic
S!)

collection: Rydberg 2623, NY, U

Stems glabrous or more rarely sparsely pubescent. Leaves linear, green

389

or occasionally tinged with purple, 0.5—6.0 cm long, 0.5—3.5 mm wide.
Bracts overlapping, exposing the axis along the lower portion of the in-
florescence, at least as broad as and commonly slightly longer than the
fruit, narrowed at apex. Spikes densely flowered, up to 13 cm long. Fruits
1.0—3.5 mm wide, lacking distinct wing margins though the margin of the
fruit may be sharpened or acute.

The specific name is derived from the type locality: ‘‘in Oriente.’ The
type specimen is maintained in Paris (P).

The selection of the lectotypic collection of C. villosum was actually made
by Standley (1916) when he chose Manhattan, Montana, as the type locality.

Immature specimens of C. orientale and C. hyssopifolium are somewhat
similar in appearance; however, C. hyssopifolium can be distinguished from
C. orientale in immature stages by the degree of pubescence on the in-
florescence. The inflorescence of C. hyssopifolium (especially in younger
plants) tends to be very densely pubescent, whereas that of C. orientale is
seldom pubescent, even during early stages of development.

The New World and Old World specimens of C. orientale are virtually
identical. Although Macbride (1918) was correct in proposing that C. villo-
sum and C. emarginatum are not native species, that they do in fact repre-
sent the same taxon, and that they are actually C. orientale, he suggested
that North American specimens differed enough from Old World material
to be recognized as a distinct variety, viz. C. orientale var. emarginatum
(Rydberg) Macbride. Careful attention was given to Macbride’s description,
but no distinguishing features were observed among the specimens which
would support the proposed variety. Consequently, only one nonwing-mar-
gined species (without varieties) should be recognized among the North
American taxa of Corispermum.

Representative specimens: CANADA: ALBERTA: Cardston, vicinity of Calgary, 8 Au
1914, Moodie 6 (F); Fort Macleod, : Aug 1895, Macoun 12,93) (F). ONTARIO: L ai
.: Port Franks, 15 Oct 1906, ee sam. (MU, TEX); Rainy River ae oe
railroad ballast, 15 Aug 1961, Garton 9383 (MIN); Thunder Bay Distric Po rthur,

ae ae 6 Sep 1952, Garton 2209 (MO). SASKATCHEWAN: Medic ne 78
Aug 18 Macon 12936 (NY); Webb, sand dunes, 29 Aug 1947, Breitung 5 5842 (MO).
MEX ios. State of Chihuahua: much ean forming sand tuft, 1852, Thurber s.n.

(F).

UNITED STATES: ARIZONA: cerns Co.: Canyon de Chelly National Monument,
dry sandy areas in canyon, 17 May 1 Demaree 38425 (UT); Coconino Co.: Flagstaff,
5 Sep 1943, Schallert sn. (MO, ae Coro ie. Chaffee Co.: Salida, 31 Aug 1892,
ee SM. cae E] Paso Co.: Colorado a 11 Aug 1873, Porter s.n. (RM);
Gunnis : Gunnison, along irrigation ditches and adjacent dry ground, 12 Sep 1901,
cee ae Selby 401 (NY); Moffat Co.: infrequent on on sandy areas one-quarter
f

mi W
MICHIGAN: Marquette Co.: Marquette, 1 Sep 1923, Sher ff SM, (F)._ MINNESOTA:

Woods Co.: Morris Point Osea a a growing on sandy shore, 29 Aug 1939, Miers and
Moore 12226 (GH, RM); St. Louis Co.: Duluth, in gravel common on the beaches of the

lake and ba ug 1936, Lake 1770 Clee US). MISSOURI: Clay Co.: common, 21
Sep oe Macken: ie 422 (M Y); Jackson . Kansas OI; sands, 24 Sep 1909, Bush
5923 (GH, } , MO, NY, a Ray Co.: ne ars and alluvial depressions a ie ee
River, 5.5 mi aoe of Henrietta, 9 Oct 1956, Sfeyermark $3150 (MO). MONTA Daw-

390

son Co.: Colgate, 6 Sep 1892, Sandberg, MacDougal, and Heller 1013 (US); Hill Co.:
Havre, 4 Sep 1903 ” Blenkinsbip sn. (RM). NEW MEXICO: Bernalillo Co.: Albuquerque,
Aug 1884, ie 4936 (NY); se aie Co.: San Ysidro, 18 Aug 1926, Arsene and Bene-
dict 16476 (F, US); Santa Fe Co.: Santa Fe, 10 Sep 1881, Engelmann s.n. (MO); Valencia
Co.: Grants, I Oct 1884, Jones oe (US). NORTH DAKOTA: Billings Co.: Medora,
in sand near top of butte, 17 Sep 1911, Bergman 1284 (RM); Cass Co.: Fargo, in sand on
railroad grade, 12 Sep 1942, Stevens 669 (MO, NY, UC); Morton Co.: Mandan, 10 Sep
1891, Wright ne (NY, RM). OREGON: Walla-Walla Co.: 17 Sep 1894, Leiberg 912
(GH, MO, NY, US). TEXAS: Wilbarger Co.: Sep 1880, Bol! 1232 (F). WISCONSIN:
Ashland Co.: railroad tracks, 10 Sep 1927, Penokee 16282 (MO). WYOMING: Albany
Co.: Laramie, i cree bad 22 Aug 1913, Machride 2699 (MO); Sweetwater Co.:
sand dunes near Steamboat Moauniesin, 1 Sep 1936, Ownbey and Lang 1122 (MO); Uinta
Co.: Diamondville, on the loose railroad grades, Nelson 8109 (GH, MIN, MO, NY, RM
S).

DOUBTFUL SPECIES
Corispermum pilosum Rafinesque, New Flora and Botany of North America
4:46. 1836.

Originally collected by Kin (or Kinn) in Florida (Rafinesque, 1836), no
specimens of this species have been collected in recent years. Standley
(1916) suggested that this plant may be C. hyssopifolium L., if it was in fact
correctly placed in this genus.

DISTRIBUTIONAL HISTORY OF CORISPERMUM IN NORTH AMERICA
The increased distribution of all three species of Corispermum with time
appears to be associated with two major factors of seed dispersal, water-
ways (rivers, streams, lakes) and transport along thoroughfares. A third,
but relatively minor (operating within a smaller range), factor is wind.
Wind, particularly for those populations which have become established
on sand dunes, could account for seed dispersal across a distance of several
miles.
In Canada the range of Corispermum appears to have increased north-
ward and westward with time. This statement is based on herbarium speci-
mens examined, Many of these Canadian specimens were collected on or
near military camps, and the increase in range of Corispermum may have
accompanied the establishment of these camps.

The range of Corispermum in Mexico seems fairly limited; however, all
three species are present and apparently have been for some time (i.e.
since at least 1912 for C. hyssopifolium, 1935 for C. nitidum and 1852 for
C. orientale). Thus, it is probable that their distribution in Mexico is greater

r findings indicate. This could be a result not only of the herbaria
selected for this study, but also of the general incompleteness of representa-
tive herbarium material available from this region.

n the basis of specimens examined it is not possible to indicate a certain
point of origin for each species in the United States nor exact pathways of
establishment. However, during the course of this study several locations
of early establishment became apparent. Again, these are not necessarily
indications of points of origin, but are only suggested population centers

391

for each species early in the history of Corispermum in the United States.

It is postulated that the earliest representatives of the genus (i.e. C.
hyssopifolium) in the United States became established just prior to 1818
(Nuttall), somewhere in the midwest. The establishment of all three species
since that time has been successful throughout a major portion of the coun-
try. Populations of both C. nitidwm and C. hyssopifolium in the Great Lakes
region and in several states along the Missouri River were some of the

centers, the species’ rapid spread would have been facilitated by major
waterways. Based on numbers of herbarium specimens examined, both
C. nitidum and C. hyssopifolium appear better established than C. orientale,
and C. hyssopifolium, of the first two, is probably the more widespread.
Earliest specimens of C. orientale were from New Mexico and Texas in the
late 1800’s (circa 1880). However, C. orientale is now rather widely distri-
buted throughout the United States. The establishment of C. orientale in
many areas seems to be associated with transport routes, particularly rail-
ways, and is thus more recent than that of the other two species. Conse-
quently, it is postulated that, with time, C. orientale will become as suc-
cessful in North America as C. nitidum and C. hyssopifolium.

REFERENCES
HOOKER, W. J. 1838. ae Boreali-Americana 2:125-126, Reprint, 1960. Hafner Pub-
lishing renee New ork,
HOST, N 827. Flora ees: 1:31 Vienn
ILJIN, M. ee In: V. L. Komarov id 7. ie Shishkin, Flora U.S.S.R. 6:105-122. Aka-

LAMARCK, J. 1786. Encyclopédie se asieae ee :110-111. Paris.
LINK, H. F. 1820. Jahrbiicher der Gewichskunde 1:27-31. Berlin.

} arum 1:4.
NELL, J. 1910. New plants from North Dakota. Amer. Mid]. Naturalist 1:207
MACBRIDE, J. F. 1918. New or otherwise interesting plants, mostly North Sa
Liliaceae and Chenopodiaceae. oath Gray Herb., new series, 53:12-13.
NELSON, A. 1909. Coulter’s new manual of botany of the Central Rocky Mountains, re-
vised ed., 163-164. American Book Company, Chicago.
NUTTALL, ze 1818. The genera of North American Plants 1:
aaa ae Collections toward a flora of the een of Arkansas. Amer. Phil.

RATINESOUE. @ S. 1836. New flora and botany of North America 4:46-47. ean
RYDBERG, me a 1897. Rarities from vente Bull. Torrey Bot. Club 24:19

. Studies on the Rocky Mountain Flora. Bull. heey Bot. Club ous

. Studies on the oo Mountain Flora. Bull. Torrey Bot. Club 31:404.
SCHUL ae “a i 1814. Osterreichs Flora, revised e ed... “Pe. Seicausbure Company, Vienna.
STANDLEY, P. C. 1916. Iv: North American lors 21(1):79-80. New York Botanical

Garden, New York.
WATSON, S. 1874. A revision of the North American Chenopodiaceae. Proc. Amer. Acad.
Arts 9:82-126,

NOTES

ADDITIONS TO THE TEXAS FLORA—Recent collections sent to the SMU
Herbarium for identification from the Texas Parks and Wildlife Depart-
ment have yielded a number of additions to the Texas flora. I acknowledge
help from David Riskind of that Department and from the New York Bo-
tanical Garden for ae of a sass men

Holcus lanatus L. Co.: Ca ddo Lake State Park, 27 Jul 1976,
Fleetwood 11746 a aie at TAES. This was reported by Silveus
(1933) from Beaumont, Texas, and was listed for Texas by Gould (1975),
who cited no voucher specimen. A duplicate collection was sent to Frank
Gould of Texas A&M University for verification.

Eleocharis baldwintt (Torr.) Chapm. Harrison Co.: moist site along boat
ramp in Caddo Lake State Park, 15 Jul 1975, Fleetwood 11833 (SMU). This
species, included in Correll & Johnston (1970) although no collections were
known for the state, was to be expected in eastern Texas. Svenson (1937)
cited a collection from Caddo Parish, La.: sandy silt on margin of Caddo
Lake near Oil City, 23 Sep 1934, Uhler & Kubichek (B). The western part
of this lake extends into Harrison County, Texas.

Cardamine pensylvanica Muhl. This crucifer was included in the state’s
flora by Cory & Parks (1937) and by Gould (1969). Correll & Johnston (1970)
noted that the species had been reported for the state, but they excluded
it because they had not seen specimens from Texas. The following Texas
collections are now available. Dallas Co.: weed under lath, mostly along
walks in clay, Northhaven Gardens, north Dallas, 6 Nov 1960, Shinners
29514 (SMU). Harris Co.: sandy soil of backyard, 400 block Southchester
Road, Houston, 16 Mar ne Smith 15 (SMU). Liberty Co.: along roadside
of hwy. 787 near Melvid, 25 Mar 1971, Amerson 335 (SMU). This is a species
similar to C. parviflora var. arenicola but with the terminal leaflet larger
than the lateral ones and the stems ped hispid near the base.

Plantago rugelii Dene. This is a widespread perennial species primarily
in the eastern half of the United Stat a southern Canada. Although
early reported for Texas by Coulter een -1894) and later by Cory & Parks

(1937), it has been consistently omitted from more recent treatments of the
state’s flora, e.g., Shinners (1950, 1958), Gould (1969), Correll & Johnston
(1970), and Correll & Correll (1972). Small (1933) and Steyermark (1963)
included Texas in the range of P. rugelii.

The species is morphologically similar and closely related to the more
widespread but less common P. major. They may be separated as follows
(Steyermark, 1963).

Mature fruit peerage transversely between the base and the middle,

seeds 4—9; sepals an racts subacute to acute; bracts Ae
lanceolate; base of petioles usually dark red-purple. . ugelii

SIDA 7 (4): 392. 1978.

393

Mature fruit dehiscing transversely near the middle, seeds 6—15; sepals
and gees uae or obtuse; bracts oes ovate; base of petioles
usually g . . P. major

Originally I was going to report this species for Texas based on a single
collection from Titus County. While going through Bassett’s Plantains of
Canada (1973), I noted on a distribution map two dots in Texas. Correspond-
ence with Mr. Bassett, made in an attempt to locate these collections, elicit-
ed the response that he had not kept records and thus was uncertain of the
herbarium or herbaria in which Texas specimens of P. rugelii were de-
posited.

The following voucher collections will document the long-time presence
and the continued existence of this species in Texas. Dallas Co.: damp
woods, Dallas, Jul 1877, Reverchon s.n. (NY). Grayson Co.: Denison, 31
Aug 1936, Tharp s.n. (SMU). Titus Co.: near Swanano Creek N of hwy. 11
between Cason and Pittsburg, 16 Jun 1973, Amerson & Riskind 1812 (SMU).

Helianthus strumosus L. This widespread, summer-flowering species of
central and southeastern United States was previously reported from Texas
by Heiser (1969). Correll & Johnston (1970), however, did not include H.
strumosus in the Texas flora. The following collections are voucher specimens
from northeastern Texas. Harrison Co. Caddo Lake State Park, 27 Jun 1975,
Fleetwood 11699 (SMU); 30 Jun 1975, 11761 (SMU); 15 Jul 1975, 11783 (SMU);
30 Jul 1975, 11791, 11796 (SMU).

Helianthus strumosus is a perennial species closely related to H. tuberosus
but differing primarily in having generally smoother stems, in the absence
of glands on the lower leaf surface, and in the rhizomes not being tuberous.

Senecio vulgaris L. This was reported for Texas by Cory & Parks (1937)
and by Gould (1969). It is, however, omitted from Correll & Johnston (1970).
A recent collection of S. vulgaris in north central Texas has led to the dis-
covery of earlier collections of this species in the SMU herbarium. The fol-
lowing are voucher specimens. Dallas Co.: weed in flower bed in black
clay, University Park, Dallas, 13 May 1958, Shinners 27305 (SMU); weed
under lath, sandy loam, Northaven Gardens, north Dallas, 6 Nov 1960, Shin-
ners 29156 (SMU); in bed of evergreens, 1950 N Industrial Blvd., 28 Apr
1970, Flyr 1382 (SMU); open field behind Faircrest St. near Lake Highlands
High School, 7 Mar 1977, McGehee 90 (SMU). Wichita Co.: lawn weed next
to driveway, W side of City Lake in Iowa Park, 10 Apr 1971, Mahler 6420
(SMU).—Barney Lipscomb, Herbarium, Southern Methodist University,
Dallas, TX 75275

REFERENCES
ee I. J. se The plantains of Canada. Monograph 7, Research Branch, Canada
c

ry D. S. a M. Cc. aaa ae 1970. Manual of the vascular plants of Texas.
Texas Research Foundation, Renner, Tex

SIDA 7(4): 393. 1978.

394

CORRELL, D. = and H. B. CORRELL. 1972. Aquatic and wetland plants of southwestern
United States. Environmental Protection Agency. Washington, D.C.

CORY, V. L. and H. B. PARKS. 1937. Crealenie of the flora of the state of Texas. Tex.
Agric. Exp. Sta. Bull. No. 550.

COULTER, J. M. 1891-1894, Botany of western Texas. Contrib. U.S. Natl. Herb.

GOULD, F. W. 1969. Texas plants—A checklist and ecological summary. ‘Tex, eee
Exp. Sta. so ae aga

. The ee . Texas. Texas A ae Univ. Press, College Station.

HEISER, ne i . SMITH, S. B. EVENGER and W. C. MARTIN,
1969. ae Sees ies ae eo vs Torrey Bot. Club 22(3): 1-21
SHINNERS, L. H. 1950. The North Texas species of Pimiazo (Plantaginaceae). Field :

Lab. 18(3): 113-119.
1958. Spring flora of the Dallas-Fort ae area Texas. Publ. by author, Dallas
SILVEUS, W, A. 1933. Texas grasses. Publ. by author, San Antonio.
SMALL, J. K. 1933. Manual of the southeastern ae Publ. by aes New ork
STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univ. Ames
SVENSON, H. K. 1937. Monographic a of the genus Sas IV. Rhode 39:

NOTEWORTHY VASCULAR PLANT RECORDS FROM TEXAS—Recent
collections from units of the state park system as well as from natural re-
source investigations involving the Texas Parks and Wildlife Department
have yielded several distribution records of note.

Sporobolus silveanus Swallen. This species was previously known, in
Texas, from the southeastern counties of Brazos, Hardin, and Orange.
(Gould, The grasses of Texas, 1975). Sporobolus silveanus is reported here
as occurring in far northeastern Texas in Lamar County, the following col-
lection representing a disjunction of at least 350 km north of the nearest
known Texas locale: Lamar Co.: Tridens prairie at intersection of hwy. 82
and farm road 32 ca. 7 miles W of Paris, Nov 1971, Collins s.n. (LL). Spo-
robolus silveanus constitutes ca. 57% basal cover in the Tridens prairie
community. Associated species included Andropogon gerardi, Desmanthus
illinoensis, Bevcninus hirsutus, H. maximilianti, Manisuris cylindrica, Pas-
palum floridanum, Sorghastrum avenaceum, Tridens strictus, and Tripsacum
dactyloides.

Menispermum canadense L. Recent collections from Bandera and Real
counties document the rediscovery of Menispermum in Texas—where it had
not been collected in over 70 years. The first known Texas collection is:
Dallas Co.: rare in woods, 30 Aug 1901, Reverchon 2733 (GH, SMU). Shinners
included the species in his Spring flora of the Dallas-Fort Worth area, Texas
(2nd ed., 1972). However, Correll & Johnston (Manual of the vascular plants
of Texas, 1970) and Stanford (Keys to the vascular plants of the Texas Ed-
wards Plateau and adjacent areas, 1976), do not report this taxon in their
manuals. The following collections document M. canadense in Texas: Ban-
dera Co.: Lost Maples State Natural Area, clambering over herbs and rocks
and climbing into shrubs and trees in rocky stream bottom. Abundant within
ca. 0.5 hectare. Associated with Aesculus pavia var. pavia, Diospyros, tex-

SIDA 7 (4): 394. 1978.

399

ana, Juglans major, Juniperus ashei, Prunus serotina, Quercus texana,
Tilia floridana, and Verbesina virginica, 23 Apr 1974, Snyder 232 (LL);

Jun 1976, Riskind & Riskind 2002 (SMU). Real Co.: climbing over shrubs,
herbs, and rocks in bed of Can Creek in Lost Maples State Natural Area,
18 Jun 1975, Smith 693 (LL). The nearest known populations of M. canadense
are in Caddo and Canadian counties, Oklahoma (Rice, The microclimate

homa, Ecology 41:445-453.1960; Little, The vegetation of the Caddo County
canyons, Oklahoma, Ecology 20:1-10.1939). The species is also known from
the mountains above Monterrey, Nuevo Leén, Mexico. No collections of
Menispermum from intervening points are known to me. Populations of
Menispermum in the Lost Maples State Natural Area have been observed
for 2 years, and no fruiting individuals seen; the population appears to be
unisexual. All collections made were from staminate plants.

Acer grandidentatum Nutt. Canyon big-tooth maple is a common tem-
perate deciduous small tree in mesic canyon woodlands of moderate eleva-
tions in mountains of northern Mexico and southwestern and western Unite
States, north to Utah and Wyoming. The species is localized but well repre-
sented in certain deeply entrenched, mesic limestone canyons on the south-
ern portion of the Balcones Escarpment in Texas as far east as Kendall
County. Most A. grandidentatum of the Edwards Plateau and neighboring
areas has been referred to the var. sinuosum (Rehd.) Little. Recently dis-
covered stands of A. grandidentatum in Bell County, ca. 220 km north north-
east of the Balcones Escarpment populations, appear closest to the western
var. brachypterum, based upon leaf morphology. Bell Co.: exposed slopes
and creekbeds of rocky limestone soils in intermittent tributary of Bear
Creek, Fort Hood Military Reservation, 6 Jun 1976, Riskind & Snyder 1993,
1997, 1999 (NY, SMU); 6 Jun 1976, 2000 (NY).

_igncnia capreolata L. Bignonia capreolata is a common high-climbing

the Balcones Escarpment is unusual and noteworthy: Bandera Co.: Sabinal
Canyon ca. 7 miles N of Vanderpool, Warren Murphey Ranch, on steep
limestone bluff ca. 100 ft above river, 8 Apr 1972, Welborn s.n. (LL). This
area is now within Lost Maples State Natural Area. Welborn’s collection
represents a disjunction of ca. 410 km from the nearest known collection in
Walker County and at least 270 km from a disjunct population along the
Guadalupe River in Victoria County, 29 May 1932, Tharp s.n. ye

Gailum correllii Dempst. Heretofore known only from the type locality
on limestone walls of Langtry Canyon (Mile Canyon) in Val Verde County
(Correll & Rollins 32612) and limestone cliffs above Rio Grande in north-
western Coahuila, Mexico (Johnston, Wendt, & Chiang 10607). This species
has recently been collected from vertical walls of a tributary canyon of
the Rio Grande. Brewster Co.: local in limestone crevices at Waterworks

=

SIDA 7 (4): 395. 1978.

396

Canyon at Asa Jones, 1 Apr 1973, Riskind 1016 (LL); 19 Oct. 1975, 1861a
(LL). Associated plants were Cirsium turneri, Penstemon baccharifolius,
and Perityle cf. parryi. These collections represent range extensions up-
stream along the Rio Grande of ca. 100 km.

I acknowledge assistance from: Debbie Balser, Harold Beaty, Marshall
Johnston, Barney Lipscomb, Larry Lodwick, Wm. F. Mahler, Jackie Smith,
Bruce Snyder, and Carroll E. Wood, Jr.—David H. Riskind, Resource Man-
agement Section, Texas Parks and Wildlife Department, Austin, TX 7874

CHROMOSOME NUMBERS OF ASTER ERICOIDES L. AND ASTER
PILOSUS WILLD. (COMPOSITAE)—For the widespread, common Aster
ericoides, there is but a single report of chromosome number: 2n = 32
(Huziwara. Karotype analysis in some genera of Compositae. VIII. Further
studies on the chromosomes of Aster. Amer. J. Bot. 49:116-119. 1962). In light
of this single report, I undertcok to confirm the determination. In two local
populations (Harriman 1278, Harriman 14,178, in herb. OSH), I found re-
peatedly 2n = 10 in root tips. Dr. Huziwara (of Kobe University, Japan)
has kindly loaned to me the voucher specimen for his determination; the
plant with which he worked proves to be A. pilosus, not A. ericoides. My
report, then, establishes a correct number for A. ericoides but leaves open
the question of whether A. pilosus should have as one of its chromosome
numbers Huziwara’s count of 2n = 32, an anomalous number for a species
in which 2n = 48 and n = 12 have been reported (Van Faasen & Sterk,
Chromosome numbers in asters. Rhodora 75:26-33. 1973). I examined one
local population of A, pilosus (Harriman 8822, in herb. OSH) and likewise
found 2n = 48. I suggest that Huziwara’s count is best treated as an error
in Aster, perhaps stemming from some inadvertent mixing of seeds and
specimens sent him by workers at the Montreal Botanical Garden.—Neil
A. Harriman, Biology Department, University of Wisconsin-Oshkosh, Osh-
kosh, WI 54901

ADDITIONS TO ALABAMA LYTHRACEAE.—Shirley A. Graham (Sida
6: 80-103. 1975) recently revised Lythraceae for the forthcoming Vascular
Flora of the Southeastern United States (A. E. Radford et al., editors). Re-
cent collections, listed below, are additions to known ranges and supplement
existing physiographic province records. These specimen e deposited in
the Herbarium of the University of North Carolina at aad a (NCU).

Ammannia coccinea Rottb, ALABAMA. INTERIOR LOW PLATEAU:
Franklin Co., Whetstone & Massey 4426; Lawrence Co., Whetstone & Rad-
ford 7126. CUMBERLAND PLATEAU: Cullman Co., Whetstone & Atkinson
3610; Etowah Co., Whetstone & cals 6868; qacicson Co., Whetstone &
Radford 7489. COASTAL PLAIN: Barbour Co., Kral 33202 (NCU 451006).
Heretofore, the documented distribution of this species included all provinces

SIDA 7(4): 396. 1978,

397

of the southeast with the notable exception of Alabam

Lythrum salicaria L. ALABAMA. CUMBERLAND A Tate Madison
Co., Whetstone & Marx 8761. The nearest known station for this species was
Watauga County, North Carolina (Taylor & Evans 56, NCU 382589).—R.
David Whetstone, George R. Cooley Research Laboratory, Department of
Botany, University of North Carolina, Chapel Hill, NC 27514

NOMENCLATURAL CORRECTION IN CLEMATIS.—During a revision of
Clematis (see Sida 6: 36. 1975) for the forthcoming Vascular Flora of the
Southeastern United States, I accepted the name Clematis maximowicziana
Franchet & Savatier for the commonly cultivated clematis native to Japan
which is now casually escaped throughout eastern North America. Mean-
while, Dr. L. A. Lauener (in litteris) kindly indicated that Hiroshi Hara
(J. Japanese Bot. 50: 155-158. 1975) established the identity of Clematis ter-
niflora DeCandolle (Syst. 1: 137. 1817), and which has priority over C. mazi-
mowicziana. After studying Hara’s paper, I am prepared to accept his con-
clusion that C. terniflora is the correct name for the clematis passing various-
y as C. paniculata Thunberg (Trans. Linn. Soc. 2: 337. 1794, non J. F. Gme-
lin, Syst. 873. 1791), C. maximowicziana Franchet & Savatier (Enum. Pl.
Japon. 2: 261. 1878), or C. dioscoreifolia Léveillé & Vaniot (Fedde, Repert.
Nov. Sp. 7: 339. 1909). For a complete list of synonyms, consult Hara’s
paper.—Carl S. Keener, 202 Buckhout Laboratory, The Pennsylvania State
University, University Park, PA 16802

ERRATA

In my Notes on Coreopsis (SIDA 7:304-307. 1978), I accidentally gave cre-
dit to Rafinesque for the combination C. X delphinifolia. In the entries 3 and
21, this should be corrected to C. X delphinifolia Lam.—Edwin B. Smith,
Department of Botany & Bacteriology, University of Arkansas, Fayetteville,
Arkansas, 72701.

SIDA 7 (4): 397. 1978.

SID

CONTRIBUTIONS
TO BOTANY

Index to Volume 7

Names of contributing authors are in capital letters. New scientific names
are in boldface. Synonyms are in italics. Page numbers in italics are those
pages on which a name is used as a synonym.

Abildgaardia 146, 312

Acalypha gracilens 62, 135; rhom-
boidea 62, 135; setosa 135; virgi-
nica 62

Acanthaceae 29, 54, 142

Acanthospermum australe 57
cer drummondii 136; grandidenta-
tum 395, var. brachypterum 395,
var. sinuosum 395; rubrum 54,
118, 136, var. drummondii 54, 136;
saccharinum 54

Aceraceae 54, 1:

Achillea millefolium 143

Aconitum 1, 3, 4

Actaea 2, 3, 4, 12; alba 3, 11; pa-
chypoda 38, 4, 11, f. rubrocarpa 3;
rubra 4, 5, f. neglecta 4; spicata
4, 5, var. alba 3, 4, 5

Actinospermum uniflorum 143

Acuan illinoense 1:

diantaceae 283, 289

Adiantum capillum-veneris 289,
290; pedatum 289, 290

Adonis 1, 2

Adopogon carolinianum 144

Aeschynomene indica 361, 364

Aesculus pavia 67, 394

Agalinis setacea 76; fasciculata
141; obtusifolia 141; purpurea
141; setacea 141; tenuifolia 76

Ageratina aromatica 143

Agrimonia microcarpa 75; pubes-
cens 75, var. microcarpa 132;
pumila 132

Agrostis alba 221; elliottiana 63,
122; gigantea 221: hyemalis 638,

122; intermedia 122; perennans
63,
Aira elegans 122
Aizoaceae 54, 83, 131
Albizia julibrissin 69, 133
Alchemilla microcarpa 220
Aletris aurea 70, 128: farinosa 70,

Aleurites fordii 62, 135

Alismataceae 54, 122

Alliaria officinalis 91

Allium bivalve 128; canadense 71,
128, var. mobilense 71; inodorum

Alnus 213; serrulata 55

Alcpecurus carolinianus 63, 122;
geniculatus 122

Alsine media 131

Alternanthera philoxeroides 54, 130

Amaranthaceae 54, 130

Amaranthus hybridus 54, 130; spin-
osus 130; viridis 54, 130

Amaryllidaceae 54, 128

02; artemisiifolia

Amianthium muscaetoxicum 128

Ammannia coccinea 396

Ammoselinum butleri 220

Amorpha fruticosa 69; paniculata
361, 364

Ampelopsis arborea 79, 136; cor-

Amphicarpa bracteata 69
Anacardiaceae 54, 135
Anchistea virginica 297

Andromeda glaucophylla 36

parius 118, 122; subtenuis 122;
tener 118; ternarius 64, 122; vir-
ginicus 64,
Aneilema nudiflorum 128
Anemone 1, 2, 12; hepatica 11
Angelica venenosa 138
Anisostichus capreolata 141
Annonaceae 54, 131
Antennaria fallas 57
Anthaenantia villosa 64, 122
Anthemis 99
Fee ee ee artistatum 221
Apiaceae 138
Apios americana 69, 133; apios 133
Apium anmi 138; leptophyllum 78,
138

Apocynaceae 54

Apteria aphylla 55, 129; setacea 129

Aquifoliaceae 54,

Aquilegia 2, 5, 9, 12; australis 5;
canadensis 5, 6; coccinea 5, var.
coccinea 5, var. latiuscula 5; vul-
garis 5,

Arabis virginica 132

Araceae 32, 55, 127

Arachis hypogaea 69

Aralia spinosa 55, 138

Araliaceae 55, 138

Arecaceae 127

Arenaria serpyllifolia 131

Arethusa bulbosa

Arisaema dracontium 55; triphyl-
lum 55

Aristida affinis 64; dichotoma 64;
gracilis 122; intermedia 122; la-
nata 122; lanosa 64, 122; longespi-
ca 64, 122; mohrii 122; oligantha
64; palustris 122; purpurascens
64, 91, 122; simpliciflora 64; vir-
gata 64, 122

Aristolochia hastata 55; reticulata

399

364; serpentaria 55, 364; tomen-
tosa 59

Aristolochiaceae 55

Armoracia aquatica 34

Aronia arbutifolia 133

Artemisia 9

Arundinaria gigantea 64; tecta 64,

Arundo donax 122

Asclepiadaceae 55, 92, 139

Asclepias amplexicaulis 55, 139;
humistrata 55, 139; michauxii
139; obovata 55, 139; obtusifolia
139; perennis 55; tuberosa 139;

Ascyrum hypericoides 67, 136, stans

Asimina parviflora 54, 131

Aspidiaceae 121

rene 121, 283, 291

Asplen 291, 295; pinnatifidum
294, cone platyneuron 74, 121, 294,
995: rhizophyllum 291, 294, 295;
trichomanes 294, 2

Aster adnatus 57, 143; bifoliatus
143: concolor 57, 143; dumosus
57, 143; ericoides 396, var. pilosus
143; exilis 148; laricifolius 300;

linariifolius 57,

366; simplex 57; solidagineus 143;
subulatus 83; umbellatus var.
brevisquamus 143; undulatus 57
Asteraceae 83, 143
Astragalus distortus 69; villosus 133
Atamosco atamosco 129
Athyrium 291, 295; asplenioides 74,
121, 294, 295; felix-femina var. as-
plenioides 295; pycnocarpon 294,

Atriplex maritima 155
Aureolaria dispersa 76; flava 141;
pectinata 76, 141; virginica 76

400

Avena sativa 64, 123

Axonopus affinis 64, 123

Azalea nudiflora 139; viscosa 139

Azolla caroliniana 39, 296, 297, 363

Azollaceae 282, 297

Baccharis halimifolia 57, 143;
wrightii 83

Bacopa monnieri 361, 365; rotundi-

olia
BAECHLE, MARK D., 248
Bahia absinthifolia var. dealbata

Balduina uniflora 143

Balsaminaceae 55

Baptisia leucantha 69

Bartlettia scaposa 84

Bartonia paniculata 364, 366

Bassia 149

Berchemia scandens 75, 136

Betula nigra 55; pumila 34

Betulaceae 34, 55

Bidens bipinnata 57, 143; coronata
143; discoidea 57; frondosa_ 57,
143; mitis 57, 1438

Bigelowia nelsonii 300; nudata 57,
143

Bignonia capreolata 55, 395; cruci-
gera 141

Bignoniaceae 55, 141

Bixaceae 95

BLACKWELL, WILL H, JR. 147,
248,

Blechnaceae 121

Boehmeria cylindrica 78, 130

Boltonia diffusa 57

Bonamia patens 59, 140

Bonplandia 97

Boraginaceae 87, 140

Botrychium alabamense 286, 287,

8

oides 286, 287, 288; obliquum 287;

virginianum 73, 286, 287, 288
BOUFFORD, DAVID E. 220

Bowlesia incana 361, 365
Brachiaria platyphylla 64, 123
Bradburya virginiana 133

Brasenia purpurea 131; schreberi

Brassica campestris 132; napus 132
Brassicaceae 55, 132

Brichellia 84

Brintonio discoidea 57, 143

Briza minor 64, 123

Bromeliaceae 55, 128

crostachys 221; racemosus 123;
tectorum 123; unioloides 64, 123
BROOKS, RALPH E. 30
Broussonetia papyrifera 130
Brunnichia ovata 74
gee americana 76, 141; elong-
a 141; floridana 76, 141
cna 120, 146, 312; barbata
60, 125; capillaris 60; ciliatifolia
60, 125
Bumelia lycioides 76
Burmannia capitata 129, 361, 364,

Burmanniaceae 55, 129

Bursa bursa-pastoris 132

Butneria florida 131

Cabomba caroliniana 39, 72

Cabombaceae 131

Cacalia angustifolia 278;
57; lanceolata 57

Cactaceae 56, 137

Calamagrostis cinnoides 39; inex-
pansa 29

Calamintha coccinea 118, 140

Calathia 240

Calathiana verna 240

Calla palustris 32

Callicarpa americana 78, 140

Callirhoe papaver

Callitrichaceae 35, 56, 135

Callitriche austinii 15; deflexa 135;
heterophylla 40; terrestris 35, 56;

elliottii

verna 35

Calopogon pallidus 73; pulchellus
119, 129

Caltha 2, 6, 12; flabellifolia 6; pa-
lustris 6, 12, var. radicans 6

Calycanthaceae 131

Calycanthus floridus 131

Calycocarpum lyoni 72

Calylophus hartwegii ssp. filifolius
88

Calyptocarpus vialis 143

Campanulaceae 56, 143, 304

Campsis radicans 55, 141

Camptosorus rhizophyllus 295

Cannabaceae 130

Capparidaceae 56

Caprifoliaceae 36, 56, 142

Capriola dactylon 123

Capsella bursa-pastoris 132

Cardamine bulbosa 55; hirsuta 55,
132; parviflora var. arenicola 392;
pensylvanica 132, 392

Carduus spinosissimus 143

Carex 24; abscondita 60; alata 29;
albolutescens 60, 125; amphibola
125; aquatilis 29; atherodes 30;
atlantica 60, 126; aurea 30; baile-
yi 30; bebbii 30; brunnescens 30;
cephalantha 30; complanata 60,
126; crebriflora 60; crus-corvi 30;
cryptolepis 30; debilis 60, 126; de-

i ; diandra

tralis 126; frankii 60; glaucescens
60, 126; haydenii 30; hirsuta 126;
hirsutella 60; howei 30, 60; ignota
60; incomperta 60; interior 30; in-
tumescens 60, 126; lasiocarpa 30;
laxiflora 60; Jleavenworthii 60,

61, 36 ‘
30; louisiani-

gli 60; nigro-marginata 60; oblita

401

61, 126; oligocarpa 61; oligosper-
ma 30; picta 61; projecta 30;
pseudo-cyperus 30; reniformis 61;
retrorsa 30; sartwellii 30; sterilis
30; straminea 30; striatula 61;
styloflexa 61; suberecta 31; swa-
nii 361, 363; tenax 61; triangularis
61; trisperma 31; venusta 61, 126;
viridula 31; willdenowii 61

Carpinus caroliniana 55

Carya aquatica 68; glabra 68; illi-
noensis 68, 129; pallida 68; to-
mentosa 68, 129

Caryophyllaceae 56, 87, 131

Cassia fasciculata 69, 133; nictitans
69, 133; obtusifolia 69, 133; occi-
dentalis 69, 133; tora 133

Castalia odorata 131

Castanea alnifolia var. floridana 63,
130; pumila 130, var. ashei 63

Castillija 89

Catalpa bignonioides 55, 141; catal-
pa 141

Cayaponia grandifolia 60

Ceanothus americanus 75, 136

Cebatha carolina 131

Celastraceae 56, 136

Celtis laevigata 77; tenuifolia var.
georgiana 77

Cenchrus echinatus 123, 361, 362;
egracillimus 123; incertus 123;
longispinus 64; pauciflorus 123

Centarium 88

Centella asiatica 78, 138

Centrosema virginianum 69, 133

Centunculus minimus 40, 75, 139

Cephalanthus occidentalis 76, 142

Cerastium glomeratum 56, 131; vis-
cosum 131

Ceratophyllaceae 34, 56

Ceratophyllum dermersum 56; echi-
natum 34

Cercis canadensis 69

Chaerophyllum tainturieri 78, 138

Chaetochloa gracilis 125; imberbis

402

125
Chaetopappa 22; hersheyi 22; par-
ryi 22; plomoensis 22, 23
Chamaecrista fascicularis 133; nic-
titans 133
Chamaecyparis thyoides 122
Chamaedaphne calyculata 36
Chamaelirium luteum 71, 12
Chamaesaracha 83; villosa 89
CHAPMAN, CAYWOOD G. 264
Chaptalia tomentosa 57, 148
Chara 218
Chasmanthium latifolium 64; laxum
64, 123; sessiliflorum 64, 123
Cheilanthes lanosa 289, 29
Chenopodiaceae 56, 87, 180, 147, 248,

Chenopodina 150; depressa_ 158;
linearis 157, 168, 164; maritima
99, 157, q@ vulgaris 155; moquini

Chenopodium 150; album 56, 1380;
americanum 158; ambrosioides
56, 130; calceoliforme 158; mari-
timum 155, 157, 163; nigrum 163,
164; pumilio 56; salsum B ameri-
canum 154; scoparia 250

Chionanthus virginicus 72

Choris petraea 123; swartziana 123

Chondrophora nudata 148

Chrosperma muscaetoxicum 128

Chrysanthemum 99

Chrysobalanus oblongifolius 132

0
Chrysopogon elliottii 125; nutans

Chrysopsis graminifolia 144; mari-
ana 144; trichophylla 144

Chrysothamnus 298

CHURCHILL, STEVEN P. 92

Cimicifuga 2, 6, 7, 12; americana 7;
cordifolia 7; racemosa 7, var.
cordifolia 7, f. dissecta 7; rubi-
folia 7

Ciminalis 177, 239; alpina 239; an-
gustifolia 239; grandiflora 239;
longifolia 239

Cinnamonum camphora 131

Cirsium carolinianum 57; horridu-
lum 57; muticum 361, 366; tur-

Cistaceae 56, 13

Citrus trifoliata 76
Cladium mariscoides 31
Cleistes divaricata 129

flora 397; virginiana 75
Cleome houtteana 56
Clethra alnifolia 119, var. tomento-
sa 138
Clethraceae 138
Cliftonia monophylla 119, 135
Clinopodium coccineum 140
Clitoria mariana
Cnidoscolus et ieais 63, 118, 135
Cobaea 97
Cocculus carolinus 72, 131
Cochlospermaceae 95
Coilantha mocinni 196, 200, 201;
sessaei 196, 200
Coldenia hispidissima 87
Collinsonia canadensis 68; serotina

Colocasia esculenta 127

Colutea arborescens 361, 365
Comastoma 232, 244

Commelina caroliniana 56; commu-

Conceiniin ani 143

Consolida

Convolvulaceae 59, 140

Conyza 875; canadensis 57, 375;
floribunda 57

Coptis 2; groenlandica 8; trifolia 1,
1, var. groenlandica 7,
Gorallorhisa odontorhiza 73; wis-

teriana
Coreopsis 304, 397; basalis 143;
cardaminifolia 57, var. angustilo-
a 304; corninsularis 304; X del-
phinifolia 304, 397, var. delphini-
folia f. concolor 304; drummon-
dii 143, var. wrightii 305: florida-
na 305; grandiflora var. grandi-
flora f. grandiflora 306, var. har-
veyana f. demareei 305, f. har-
veyana 305, var. pilosa 305, var.
saxicola 306; harveyana 305; he-
terogyna 305; heterophylla 305;
lanceolata 57, ; leavenworthii
, var. curtissti 305, 306;
linifolia 305; lucida 216; major 57,
; 3, f. oemleri 306; nudata
305, 306; oniscicarpa 305, var.
simulans 305; palmata 305: pu-
bescens var. debilis 304; rosea
306; saxicola 306; stellata 306;
stenophylla 57; tinctoria o7, 148,
304, 306; tripteris 57

Corispermum a americanum

bustius ae 385, var. ribricauié
5 mbricatum 383, 385;
marginale , 385; microsper-

383,
ane 383, 387; villosum 383, 388.

Cornaceae 60, 138
Cornus alternifolia 60; florida 60,

403

118, 138; stricta 60

Coronopus didymus 132, 220

Corydalis micrantha ssp. australis
132

Cosmos bipinnatus 100

Cracca chrysophylla 134;
134; spicata 134

Crassulaceae 60

Crataegus alma 75; marshallii 75;
opaca 75

Crawfurdia 232, 244

Crepis japonica 143

Crinum americanum 128: bulbi-
sperum 1

Croptilon divaricatum 57, 148

Crossostephium 99

Crotalaria angulata 69, 118, 133; in-
termedia 133; purshii 118, 133:
rotundifolia 133; sagittalis 69,

smallti

Croton capitatus 63, 135; dioicus

88; glandulosus var. septentri-
onalis 638, 18
Crotonopsis elliptica 63: linearis

135, 364
Cruciferae 34, 87
Ctenium aromaticum 64
Cucurbitaceae 60, 142

Cuscuta attenuata 364; campestris
140; compacta 60, 140; gronovii

Cynanchum laeve 55, 92

Cynoctonum mitreola 139; sessili-
folium 139, 361, 365

Cynodon dactylon 64, 123

Cynosurus echinatus 221

Cyperaceae 29, 39, 53, 42, 60, 125,
307, 309

<ypenis aCuMnaLOS 307; compres-

natus 126; eieelinannn 3l,
esculentus 61, 126; filiculmis 61.

404

126; globulosus 61, 126; haspan
61, 126; iria 61, 126; microdontus
126; odoratus 126; ovularis 61;
plukenetii 61; polysta
texensis 126; pseudovegetus 61,
126; retrofractus 126; retrorsus
61, 126; rotundus 61, 126; sesqui-
florus 61, 126; speciosus 126;
strigosus 61, 126; surinamensis
307; tenuifolius 61, 126; virens 61
Cypripedium candidum 33
Cyrilla racemiflora 62, 185
Cyrillaceae 62, 185
Cystopteris 291, 294; fragilis 294,
var. protrusa 294; protrusa 294
Dactylis glomerata 123
Dactyloctenium aegyptium 64, 123
Dalea 13;; aurea 13, 14, 15, 18, 19,
20: hallii 18, 14, 15, 18, 19, 20;
jamesii 13, 14, 15, 18, 19, 20; lani-
ceps 13, 14, 15, 18, 19, 20; luisana

20; wrightii 13, 14, 15, 18, 19, 20
Danthonia sericea 64, 123
Dasystephana 177; spathacea 196
Dasystoma flava 141; pectinata 141
Datura stramonium 77,
Daubentonia punicea 133
Daucus carota 78; pusillus 138
Decumaria barbara 76, 182
Delphinium 1, 3; carolinianum 75
Dennstaedtiaceae 283, 289
Desmanthus illinoensis 133, 394
Desmodium ciliare 69, 133; glabel-

lum 69, 133; glutinosum 69; laevi-

gatum 69, 118, 133; lineatum 69,

69; rotundifolium 69; sessilifoli-
um 91; strictum 69; tenuifolium
133; tortuosum 69; viridiflorum
69, 118, 133

Dichocarpum 8, 12

Dichondra carolinensis 60, 140;
evolvulacea 140

Dicliptera brachiata 54

Dicranocarpus parviflorus 81, 84

Didiplis diandra 41

Digitaria filiformis 64, var. villosa
123; ischaemum 64, 123; sangui-
nalis 64, 123; serotina 123; villosa
64; violascens 64

Diodia teres 76, 142; virginiana 76,

Dioscorea quaternata 62, 222; vil-
losa 62, 128,

Dioscoreaceae 62, 128

Diospyros texana 394; virginiana
62, 139

Diploma 177

Diplostachyum apodum 286; ludo-
vicianum 286

Doellingeria humilis 148
ondia americana 155, 157; brevi-
folia 151, 168; calceoliformis 159;
californica 166; carinata 157; con-

laris 132; intermedia 35, 132; leu-
cantha 132

Droseraceae 35, 1382

Drymaria lyropetala 87

Dryopteris hexagonoptera 292; no-
vaboracensis 292; patens 121; se-
tigera 292; thelypteris 121, 292

Duchesnea indica 75, 132

Dulichium arundinaceum 126, 363

Dyssodia 373; acerosa 81, 84; ap-
pendiculata 374; aurantia 374;
ccoperi 374; porophylloides 374;
squamosa 374

Eatonia filiformis 125; obtusata var.
repens 125

Ebenaceae 62, 139

Echinochloa colonum 123; crusgalli

Echinodorus cordifolius 54; rostra-
tus 29, 218; tenellus 362

Eclipta alba 57, 148

Egeria densa 122

Elaeagnaceae 137

Elaeagnus umbellata 137

Elatinaceae 39

Elatine brachysperma 39; triandra

Eleocharis acicularis 221; baldwinii

361, 362; tortilis 61, 361, 362; tu-
berculosa 61, 126; wolfii 39

Elephantopus carolinianus 57; elat-
us 57, 143; nudatus 57, 144; to-
mentosus 57, 144

Eleusine indica 64, 123

Elymus canadensis 221; virginicus

Enemion 2, 8, 9, 12; biternatum 8,
9

Epifagus virginiana 73, 361, 365;
repens 62, 138

Equisetaceae 283, 284

Equisetum hyemale var. affine 284,
285; praealtum 284

Eragrostis amabilis 64; cilianensis

405

64; ciliaris 64; curvula 123: elli-
ottii 123; glomerata 64; hirsuta
64, 123; nitida 123; oxylepis 64,
123; pectinacea 123; pilosa 64; re-
fracta 65, 118, 123; secundiflora
123; spectabilis 65, 118, 123
Erechitites hieracifolia 57, 144

ganteus 65, 123. Oe oides

2, 138
Ericala 177; spathacea 196, 200
Ixricameria 298; brachylepis 301;
ceoperi 300; diffusa 301; laricifo-

triantha 84, 298, 299, 300, 301
Ericoila 177
Erigeron 375; annuus 57, 375, 381:
basilobatus 376, 380;

1;
376; eximius 376, 379; flagellaris
381; formosissimus var. formo-
sissimus 376, 377, var. viscidus
376, 379; leiomerus 377; loncho-
phyllus 377; palmeri 377; pere-
grinum ssp. callianthemus var.
callianthemus 377; philadelphicus
07, 144, 377; pinkavii 84; pinnati-
sectus 377; platyphyllus 377; po-
tosinus 377, 379, 380; pulchellus
377; pumilis ssp.
378; ramosus 144;
strigosus 57, 144, 375, 378, 381;

superbus 379; denuke 379, 380;
vernus 144, 379;

Eriocaulaceae 32, 62, 127

406

Eriocaulon decangulare 127, 361,
363; kornickianum 363, 364; sep-
tangulare 32

Eriogonum multiflorum 364

Eriophorum gracile 39; virginicum
31; viridi-carinatum 31

Eryngium integrifolium 138, 361,
365; prostratum 78; synchaetum
78; yuccifolium 78, var. synchae-

tum
Erythrina herbacea 69
Eucnide lobata 8
Eunonymus americanus 56, 118, 136
Eupatorium album 57, 144; altissi-
mum 58; aromaticum 058, 143;
capillifolium var. capillifolium 58,
144, var. leptophyllum 144; coel-
estinum 58, 143; compositifohum
58, 144; cuneifolium 144; fistulo-
sum 58, 144; greggii 84; hyssopi-
folium 58; leucolepis 58; lineari-
folium 144; maculatum 144; per-
foliatum 58, 144; pilosum 58; re-
curvans 58, 144; rotundifolium
, 144; semiserratum 58, 144;
serotinum 58
uphorbia corollata 63, 118, 135;
irt

nutans 63; ies 135; piel
63; supina 63, 135
Euphorbiaceae 62, 88, 135
Eurythalia 238
Eustylis purpurea 365
Euthamia caroliniana 145; grami-
nifolia 58, 361, 366; gymnosperm-
oides 366, 368; leptocephala 58,

Vv
Fabaceae 133

Fatoua villosa 130

Festuca arundinacea 65, 123; elati-

ma 126; miliacea 61, 126; puber-
ula 61; tomentosa 126

Firmiana platanifolia 136

Flacourtiaceae 95

Flaveria anomala 82, 84; chlorae-
folia 84; oppositifolia 84; palmeri
84; trinervia 84

Forestiera acuminata 72

Fragaria virginiana 75

Fraxinus americana 72; caroliniana
72; pensylvanica 72

FRYXELL, PAUL A, 224, 227

scirpoidea 126, 310, 313, 314, 315,

hispida 327, var. macrostachya
332, var. pumila 338; tereticulmis
340; torreyana 338; umbellata

312, 318, 318, 324, 340, 342, 345,
346, var. unguiculata 342; walli-
chiana 312, 313, 350, 352, 353; za-

e
Gaillardia 90; aestivalis 58, var.
lanceolata 144; lanceolata 144; mul-
ticeps var. microcephala 82, 84;
pinnatifida 85; turneri 85
Galactia erecta 133; macreei 69;
mollis 133; regularis 69; volubilis

, 133

Galium aparine 76, 142; circaezans
76; correllii 395; hispidulum 142;
pilosum 76, 142; tinctorium 76,
142; uniflorum 76

Gaura angustifolia 137; biennis 72;
filipes 72, 137; michauxii 137

eee dumosa 62, 138; fron-
dosa var. tomentosa 138; hirtella
138; mosieri 138; nana 138

Gelsemium rankinii 71; sempervir-
ens 71, 118, 139

Gentiana 174, 232; acaulis 239, 240;
adsurgens 180, 184, 185, 190, 191,
B uniflora 180, 186; affinis 186,

piadea 177, 243; ingens 180,

guat
BrayAuileneie 184; hooweri 177,

407

178, 179, 180, 183, 193, 194, 195,

215; saponaria 63, , 202, 243
septemfida 239, 243; sessaei 196
sikokiana 210; cea 177

thami 196, 201, 202, yg integra 196,
201, 202; tiogana 177; verna 239,
240, 241, 242; villosa 139, 243: ya-
kushimensis 236

Gentianaceae 36, 63, 88, 139, 174,

Gentianella 174, 232, 244: amarella
209; campestris 238: utriculosa

Gentianellis affinis 240

Geranium carolinianum 63, 134;
dissectum 361, 365

Gerardia fasciculata 141; purpurea

Glinus radiatus 361, 364

Glyceria acutiflora 29; borealis 29;

408
grandis 29; melicaria 29; pallida

29

Glycine max 70

Gnaphalium falcatum 58; obtusifo-
lium 58, 144; pensylvanicum 58;
purpureum 58, 144, var. spathula-
tum 144; spathulatum 144

Gordonia lasianthus 136

Gramineae 29, 39, 53, 63

Gratiola pilosa 76, 141; sphaerocar-
pa 141; virginiana 36, 77, 119,
141; viscidula 36

Gutierrezia microcephala 85; tex-
ana 85

Gymunarrhena 98

Gymnopogon ambiguus 65, 123;
brevifolius 65, 123

Gyrostachys gracilis 129; praecox

1

Gyrotheca capitata 129

Habenaria ciliaris 73, 129; lacera

22; leucophaea 33; repens 73,
129, 361, 364

Hackelochloa granularis 123

Haemodoraceae 129

Halenia 234

Halesia diptera 77

Halogeton 148

Haloragaceae 35, 39, 67, 138

Hamamelidaceae 67, 132
amamelis virginiana 67, 132

sr hp greggii var. greggii 85,

t 85; robusta

ace 298; laricifolius 300;
linearifolius 300; parransanus
300; purpusii 302; trianthus 301

HARRIMAN, NEIL A. 3

HAUSER, LARRY A. 304

Hedeoma 103; bellum 105; costa-
tum 105; hispidum 68, 140; john-
stonii 103, 104, 105; montanum
103; palmeri 103

Hedyotis boscii 76; corymbosa 76;
uniflora 76

Helenium amarum 58, 144; flexuo-

bo

sum 58, 144; nudiflorum 144; tenu-
ifolium 144

Helianthemum = carolinianum _ 56,
137; georgianum 137

Helianthus angustifolius 58, 144;
annuus 58, 144; atrorubens 58;
heterophyllus 58, 144;; hirsutus
58, 394; maximilianii 394; micro-
cephalus 58; radula 58, 118, 144;
resinosus 58; strumosus 393; tu-
berosus 393

Heliopsis gracilis 58

a eigeaes indicum 55

Hellebor

aie fulva 71

Hemicarpha micrantha 31

Hepatica 2, 9, 12; acuta 10; acutilo-
ba 9, 10; americana 9, 10; hepati-
ca 9; nobilis 9, var. acuta 10,
var. obtusa 9

Hesperozygis pusilla 106

Heteranthera reniformis 32

Heterotheca graminifolia 58, 144;

lana ; nervosa 58; pi-

losa_ 58; eee 98; tricho-
phylla 58, 144

Hexastylis arifolia 55

Hibiscus aculeatus 71, 118, 136;
brasiliensis 223; mutabilis 136;
oxyphyllus 223; syriacus 136

Hicoria alba 129; pecan 129

Hieracium 234; gronovii 59, 144;
scabrum

Hippion 177; aestivum 241
ippocastanaceae 67

Homalocenchrus hexandrus 124;
oryzoides 124; virginicus 124

Homalosorus pycnocarpus 295

HOPKINS, CHRISTINE O. 147

Houstonia minor 142; procumbens
76, 142; purpurea 76; pusilla 76,

142; rotundifolia 142

Hydrangea quercifolia 76

Hydrastis 2, 10; canadensis 10

Hydrocharitaceae 122

Hydrochloa caroliniensis 65, 123;
fluitans 123

Hydrocotyle americana 35; bonari-
ensis 138; ranunculoides 40; um-
bellata 35, 78; verticillata 78, 138

Hydrolea quadrivalis 67; uniflora
67

Hydrophyllaceae 67, 88

Hymenocallis caroliniana 222; occi-
dentalis 54

Hymenophyllaceae 283, 289

Hypericum Peele liuitles 136; bra-
chyphyllum 136; canadense 136;
cistifolium 136; denticulatum 67;
drummondii 67; gentianoides 67,
136; galioides 67, 136; hpericoides

136; mutilum 67, 91, 136; nudi-
florum 67; pseudomaculatum 136;
punctatum 67; setosum : 6;
stans 136; stragalum 136; tubu-
losum 67; virginicum 67, 136;
walteri 67

Hypochaeris elata 144; radicata 144

Hypoxis micrantha 129; sessilis 129

Hyptis alata 68, 140; mutabilis 68,

Ilex ambigua 54, 135; amelanchier
54: carolinana 135; cassine 135;
coriacea 54, 135; decidua 54; gla-
bra 54, 119, 135; longipes 54;
opaca 55, 136; verticillata 55; vo-
mitoria 55, 118,

Illiciaceae 67

Ilicium floridanum 67, 118, 131

Ilysanthes gratioloides 141

Impatiens capensis 55

Imperata brasiliensis 65, 124

Ionactis linariifolius 148

409

Ipomoea carolina 140; coccinea 60,
140: hederacea 60, 140; lacunosa
60, 140; pandurata 60, 140; pur-
purea 140; quamoclit 60, 140; tri-
chocarpa 60, 140; wrightii 364

Tridaceae 33, 68, 129

Iris brevicaulis 33; fulva 129; hexa-
gona 129; verna 129; virginica
363

IRVING, ROBERT 5S. 103

Isnardia palustris 137

Isocoma 89, 298; coronopifolia 85;
wrightii 85

Isoetaceae 27, 283

Isoetes braunii 27; engelmannii 27,
286, 296

Isopappus divaricatus 143

Jacquemontia tamnifolia 60, 140
Jatropha stimulosa 135

JONES, SAMUEL B. 264

de nee 68, 129

Juglans major 395

Juncaceae 382, a 68, 128
Juncaginaceae 28

Junci thalassi 355

Juncus 195; acuminatus 68, 128,

poldii 358; alpinus 32; biflorus 68,
128: brachycarpus 68; capitatus
a0 363; coriaceus 68, 128; debilis

128: dichotomus 68, 128;
diffusissimus 39, 68, 128; effusus
68, 91, 128, 358; elliottii 68, 128;
gerardii 358; greenei 331; gymno-
carpus 68, 128; interior 33; mar-
ginatus 68, 128, var. aristulatus
128; maritimus 355, 356, 358; poly-
cephalus 68, 128; repens 68, 128;
roemerianus 355, 356, 357, 358,
359, 361; scirpoides 68, 128; sub-
caudatus 33; tenuis 68, 128; trigo-
nocarpus 68, 128; validus 68, 128

410

Juniperus ashei 395; deppeana 195;
virginiana 60, 1
Jussiaea decurrens 137; leptocarpa

Kneiffia fruticosa 138

Kochia 148, 248; alata 249, 250:
americana 248, 250, 252, 253, var.
californica 251, var. vestita 248,
252, 253; arenaria 248: californica
248, 250, 251, 252, 253; hirsuta 249:
iranica 249; laniflora 248; prostra-

ta 253; scoparia 248, 249, 250, 382,

u

var a , Var, pubescens
200, var. scoparia 250: vestita
248, 252

Koellia albescens 140

KRAL, ROBERT 42, 309

Krameriaceae 224

Krameria canescens 225; grayi 224;
lanceolata 224, 225; spathulata

Krigia dandelion 59; oppositifolia
59, 144; virginica 59, 144
Kudoa yakushimensis 236

Kuhnia eupatorioides 59; kuhnia

144; mosieri 144

Kuhnistera gracilis 134; pinnata
sesquiflora

Kyllinga pumila 126;
126

Labiatae 27, 68
ac ern caroliniana 129

Lachnocaulon anceps 62, 127

Lacinaria elegans 145; gracilis. 145;
graminifolia var. pilosa 145; spi-
cata 145; squarrosa 145

Lactuca canadensis 59, 144; flori-

59, 144; graminifolia 59;

sagittifolia 144

Lagerstroemia indica 137

Lamiaceae 140

Lamium amplexicaule 68, 140; pur-

pureum 68
Lantana camara 78, 140
Laportea canadensis 78
Larrea 83, 371; tridentata 89
LASSETER, J. STUART 255
Lathyrus 263; hirsutus 70

Lechea minor 56, 137; patula 137;
tenuifolia 56; villosa 56,

Leersia hexandra 124; lenticularis
65; oryzoides 65, 124; virginica
65, 124

Legouzia biflora 148; ;

Leguminosae 138,

LELONG, MICHEL G. 118

Lemna 363; minima 40, 127; pur-
pusilla 40, 127; valdiviana 32, 70

Lemnaceae 32, 70, 127

Lentibulariaceae 36, 70, 142

Leoflingia squarrosa 364

Lepidium montanum 87; virginicum

perfoliata 143

Leptochloa uninervia 124

Leptoloma cognatum 124

Leptopyrum 8

Lepuropetalon spathulatum 132

Lerchea 150; calceoliformis 159;
conferta 172: maritima 155

Lespedeza capitata 70, 133; cuneata
70, 133; hirta 70, 133; X nuttallii
70; mbens 70; repens 70,
133; peer 70, 133; stuevei 70,
33; violacea 70; virginica 70, 133

Leucactinia 369

Leucanthemum 99

Leucelene ericoides 85

Leucothoe axillaris 62, 118, 138

LEWIS, WALTER H. 95

Liatris elegans 145; gracilis 145;
graminifolia 145; spicata 59, 145;
squarrosa 59, 145; squarrulosa 59

Ligustrum japonicum 72; sinense

9

72, 18
Liliaceae 33, 70, 128
Lilium michauxii 71

Limnobium spongia 363
Limodorum tuberosum 129

1

Linaria canadensis 77, 141, var.
canadensis 361, 365, var. texana
365; texana 365

Lindernia anagallidea 77; dubia 77,
119, 141

Linnaea borealis 36

Linum medium 71; puberulum 88;
Striatum 71, 134; virginianum
var. medium 134

Lippia nodiflora 140

LIPSCOMB, BARNEY 307, 393

Liquidambar styraciflua 67, 132

Liriodendron tulipifera 71, 118, 131

Listera australis 73

Loasaceae 88

Lobelia cardinalis 56; georgiana
143; glandulosa 56, 143; inflata
92; nuttallii 304; paludosa 1438;
puberula 56, 143; spicata 56

Loganiaceae 71, 139

Lolium multiflorum 67, 124; perenne
65; temulentum 221

Lomatogonium 244

Lonicera yaponies 36, 142; semper-

1
Lorinseria areolata 4, 296, 363

2,
137; glandulosa 72, 137; leptocar-
pa 39, 73, 137; linearis 73, 137;
palustris 73, 137; pilosa 73, 137;
repens 138
Lupinus diffusus 118, 133; villosus
70

Luziola alabamensis 124; bahiensis
Luzula bulbosa 68

Lycopersicon esculentum 77

362, 366; alopecuroides 71, 121,

411

284, 285, 286; appressum 284, 285;
carolinianum 121, 284, 285; cer-
nuum 121, 284, 285; inundatum
Vv bigelovi

Lycopus rubellus 68, 140; virginicus
68,

Lygodium japonicum 76, 121, 289,
290

Lyonia ligustrina 62; lucida 139

Lythrum hyssopifolia 39; salicaria
397

MACROBERTS, D.T. 222
Machaeranthera 90; pinnatifida 82,
85: restiformis 90; tanacetifolia

Magnolia acuminata 71; foetida
131; grandiflora 71, 131; macro-
phylla 71; pyramidata 71; virgi-
niana 71, 118, 131

Maenoliaceae 71, 131

MAIHLE, NITA J. 382

Malaxis unifolia 73

Malus angustifolia 132

Malvaceae 71, 88, 136, 223

Manisuris cylindrica 394; rugosa

MANN, WILLIAM F. 225

Marginaria polypodioides 297

Marshallia trinervia 59

Marsilea 218, 219; mucronata 308;
quadrifolia 38

Marsileaceae 38, 218

Mayaca aubletii 40, 72, 127; fluvi-
atilis

Mayacaceae 72, 127

Mazus japonicus 141

Mecardonia acuminata 77, 141

Melampodium leucanthum 86

Medeola virginiana 71

, BLANCHE W. 13
Megacodon 232, 245
Megalodonta beckii 36

412

Meibomia arenicola 133; dillenii
133; laevigata 133; obtusa 133;
paniculata 133; tenuifolia 133;
viridiflora 133

Melanthium virginicum 71, 361, 364

Melastomataceae 72, 137

Melia azedarach 72, 134

Meliaceae 72, 134

ca mutica 65
ia re 133; indica 133
elochia corchorifolia 77, 136
Melothria pendula 60, 142
Menispermaceae 72, 131
Menispermum canadense 394, 395
Mentzelia 83; humilis 88; multiflora
ssp. multiflora 88; saxicola 88
Menyanthaceae 40
Menyanthes 234; trifoliata 36
Mesosphaerum rugosum 140
Michanthemum orbiculatum 141;
umbrosum 77, 141
Mikania scandens 59, 145

Monarda pectinata 222; punctata
68, 140

Monniera acuminata 141

Monotropa uniflora 139

Moraceae 72, 130

Morongia angustata 134

Morus alba 130; rubra 72, 130

Muhlenbergia diffusa 124; expansa
65, 124: schreberi 65, 124; tricho-
podes 124

MUSSELMAN, LYTTON J. 225

Myosotis macrosperma 55

Myosurus 1, 10, 11, 12; minimus 10,
ssp. minimus 10

Myrica carolinensis 129; cerifera
72, 119, 129; heterophylla 72, 129;
inodora 129; pensylvanica 34

Myricaceae 34, 72, 129

Myriophyllum = alterniflorum 40;
aquaticum 39; brasiliense 40, 67,
138; exalbescens 35; heterophyl-
lum 35; humile 40; verticillatum
35

Nabalus serpentaria 145

Najadaceae 28,

Najas 26; gracillima 28; guadalu-
pensis 218; marina 39

Nama 89; hispidum 88

Nerisyrenia 89; camporum 87;
gypsophila 87; linearifolia 87

NESOM, GUY L. 375

Nicolletia 369; edwardsii 369, 370,
72, 373: occidentalis 369, 370,
371, 372, 373; trifida 369, 370, 371,
372, 37

Nigella 1,

Nothoscordum bivalve 71, 128

Nuphar luteum ssp. macrophyllum
72; variegatum 34

Nyctaginaceae 130

Nymphaea odorata 131

Nymphaeaceae 34, 39, 72, 131

Nymphoides peltatum 40

Nyssa aquatica 72; biflora 72, 138;
sylvatica 72, 138, var. biflora 118

Nyssaceae 72,

Oenothera biennis 73, 138; fruticosa
73, 138; laciniata 138; speciosa
138

Oldenlandia boscii 142; corymbosa
142; fasciculata 142; littoralis

f US
Onagraceae 39, 72, 88, 137
Onoclea 291, 295; sensibilis 74, 121,
294, 295, 363
Onopordum acanthium 361, 366
Onosmodium virginianum 55, 140
Ophioglossaceae 73, 121, 283, 286

ni 287, 288; nudicaule 287, 288;

petiolatum 287, 288; tenerum 287,
288; vulgatum var. pycnostichum
287, 288,

Oplismenus setarius 65, 124

Orontium aquaticum 55, 127

Osmanthus americana 72, 139

Osmites-Osmitopsis 98

Osmunda cinnamomea 73, 121, 289,

0, 3638; regalis 121, 363, var.

spectabilis 73, 289, 29

Osmundaceae 73, 121, 283, 289

Ostrya virginiana 55

Oxalidaceae 73, 134

Oxalis dillenii 73, 134; priceae 73;
rubra 73, 134; stricta 73, 134;
violacea 73, 134

Oxydendrum arboreum 62, 139

Oxypolis filiformis 78; rigidior 78,

Palmae 73

Panicum aciculare 65, 124; agros
toides 66, 124; albomarginatum
66, 125; anceps 65, , var. rhi-
zomatum 65, 124; angustifolium
65, 124; arenicola 124; barbula-
tum 65, 124; boscii 65; brachyan-
thum 362; ciliatum 119, 124; cog-
natum 124; colonum 123: com-
monsianum 124; commutatum 65,
124, var. ashei 65; curtifolium 65;
depauperatum 65; dichotomiflor-
um 65, 124; dichotomum 65, 124;
elongatum var. ramosior 124; en-
sifolium 65, 119, 124; flavovirens
66; gibbum 125; gymnocarpon 65;
haemacarpon 125; hians 65, 124;
joorit 65; lancearium 124; lanugi-

lindheimeri 65, 124; longifolium
65, 124; longipedunculatum 124;

413

lucidum 65, 124; mattamuskee-

65, 124; patentifolium 124; pauci-
pilum 124; polyanthes 65; proli-
ferum 124; pseudopubescens 66,
125; pubescens 124 pyriforme 124;
ramosum 66, 124; ravenelii 66,

roanokense 65; rostratum 124;

thospermum 125; yadkinense 65
Papaveraceae 73
Paraquilegia 8
Parnassia grandifolia 361, 364
Paronychia drummondii 361, 364
Parthenium hysterophorus 145
Parthenocissus quinquefolia 79, 136
Paspalum bifidum 125; boscianum
66; ciliatifolium 66, 125, var. da-
syphyllum 125; circulare 66; com-

laeve a 235: lentiferum 66:

Passiflora incarnata 73, 137; lutea
73, 137

Passifloraceae 73, 137

Pavonia 223; anisaster 223; firmi-
flora 223; glandulosa 223; mela-

nommata 223, var. pringleana

223; ortegiana 223; oxyphylla 223,
var. melanommata 223; palmeri
223; racemifera 223; rhizophorae
223; spicata 223
Pedicularis canadensis 77, 141
Peltandra sagittaefolia 127; virgini-

Penstemon australis 77, 141; bac-
charifolius 396; hirsutus 141;
murrayanus 364

Penthorum sedoides 60

Peplis diandra 40, 363

Perilla frutescens 68, 140

Perityle parryi 86; vaseyi 86

Persea borbonia 69; palustris 131;
pubescens 131

Persicaria hydropiperoides 74; pen-

tata

carolinianum — 70,
134; gracilis 134

Petunia X atkinsiana 77

Phacelia gypsogenia 88; integrifolia
88

Phaeocephalum divergens 47; pusil-
lum 50; rariflorum 45

Phalaris caroliniana 125

Phaseolus helvolus 134; polystachi-
os 70; uwmbellatus 134

Phegopteris hexagonoptera 292

Phlox paniculata 361, 365; pilosa
74, 140, var. detonsa 140, ssp. de-
tonsa 74

Phoradendron serotinum 79, 130

Phryma leptostachya 73

Phrymaceae 73

Phyllanthus caroliniensis 63, 135;

ea 66
77, 141; barba-
1; hederaefolia 89; pu-
bescens 77, 141; virginiana 77;
viscosa 141
Physostegia virginiana 68, 140
Phytolacca americana 74, 130; de-

andra 130
Phyt olaccaceae 74

Pinguicula lutea 142; planifolia 142

Pinus 213; echinata 74; elliottii
122; glabra 74; hartwegii 178,
190; palustris 74, 122; rudis 178,
199; taeda 122

Planera aquatica 78

Plantaginaceae 36, 74, 142

Plantago aristata 74, 142; cordata

rugelii 74, 392, 393; virginica 74,
142

Platanaceae 132

Platanus occidentalis 132

Pluchea camphorata 59, 145; foeti-
da 59,

Pneumonanthe bicuspidata 180, 184;

Podostemum ceratophyllum 41
Pogonia divaricata 129; ophioglos-

soides 129
Polanisia erosa 364
Polemoniaceae 74, 140
Polemonium 97

ae virginica 54

as a

olygala brevifolia 135; cruciata 74,
65; grandiflora 74,

—
i)
io) ¢

Polygonella americana 364

Polygonum convolvulus 74; hydro-

ae 130; lapthifolium 130;
ensylvanicum 34, 130; puncta-

aa 130; ee 130

Polypodiaceae 74, 121, 283, 297

Polypodium polypodioides 74, 121,
296, 297, var. michauxianum 297

Polypogon monspeliensis 125

Polypremum procumbens 139

Polystichum 291, 295; acrosti-
choides 74, 294, 295

Pontederiaceae 32

Ponthieva racemosa

Populus 99, 100; ae 76, 129

Portulaca oleracea 75,

Portulacaceae 75, 1

Potamogeton een 75; diver-
sifolius 75, 122, 218; filiformis 27:
friesii 28; gramineus 28; hillii 28;
natans 361, 367; nodosus 218; per-
foliatus 28; praelongus 28; pul-
cher 28; richardsonii 28; robbinsii
28; spirillus 28; strictifolius 28:
tennesseensis 28; vaseyl 28; zos-
teriformis 28

Potamogetonaceae 27, 75, 122

Potentilla 92; palustris 35; plat-
tensis 91

POWELL, A. MICHAEL 80

POWELL, A. SHIRLEY 80

Prenanthes altissima 59; serpen-
taria 59, 145

Primulaceae 36, 40, 75, 139

PRINGLE, JAMES S. 174, 232

Proserpinaca palustris 67; pectina-
ta :

Prunella vulgaris 68

Prunus angustifolia 75, 132; caro-
liniana 75, 132; mexicana 75; per-
sica 75, 132; serotina 75, 132, 395;
umbellata 75

Pseudoclappia arenaria 82, 86;
watsonii 82, 86

Psilocarya nitens 126

Psilostrophe tagetina 86

415

Psilotum 93

Ptelea trifoliata 76

Pteridaceae

Pteridium aquilinum 74, 121, 289,

Pteris 289; caudata 289; latiuscula
289; multifida 290, 293
Pterygocalyx 232,
Ptilimnium capillaceum 78, 138
Pueraria lobata 70, 134
Pycnanthemum albescens 68; inca-
num 140; tenuifolium 68
Pycnodoria multifida 289
Pyracantha crenato-serrata 132
Pyrrhopappus carolinianus 59, 145
Pyrus angustifolia 75, 132; arbuti-
folia 75
Quercus alba 63, 130; austrina 63;
brevifolia 130; catesbaei 130; coc-
cinea 63; ser 130; falcata 63,
18, 130; eminata 130; hemi-
sphaerica Ps 118, 130; incana 63,
130, 364; laevis 63, 118, 130; lauri-

Ranunculus 1, 2. ‘ghontivus 79;
fascicularis 75; :
parviflorus 75; pusillus 34, 75, 131

Raphanus raphanistrum 55, 132

Regnellidium 218

Rhamnaceae 35, 75, 136

Rhamnus alnifolia 35; caroliniana

75

Rhapidophyllum hystrix 73

Rhexia alifanus 72, 119, 187; ciliosa
137; glabella 137; lutea 119, 137;
mariana 72, 119, 137, var. pur-
purea 137; nashii 72; petiolata
137; stricta 137; virginica 72, 187,
ar. purshii 137

416

Rhododendron canescens 62, 139;
viscosum var. serrulatum 139
Rhus copallina 54, 118, 135; glabra
54, 135; radicans 54, 135; toxico-
dendron 54, 135; vernix 54, 135
Rhynchosia CneEG, 134; galactioides
118, 134; pitcheria 70; reniformis

, 134; pees 134; tomen-

esa , 134
asmichosnor 42, 119; alba 31: az-
illaris 126; caduca 61, 126; cepha-
lantha 126, var. pleiocephala 61;
chalarocephala 126; chapmanii

127, 361, 362; slenoohvila 43, 47,
48; thornei 42, 43,

Riccia fluitans 41

Richardia brasiliensis 142; scabra

76, 142
RISKIND, DAVID H. 396
ROBERTS, MARVIN L. 24
Robinia pseudo-acacia 134
G 1

Rorippa 92; austriaca 91; sessili-
flora 35; walteri 132

Rosa bracteata 75, 132; carolina 75;
laevigata 132; multiflora 91, 132;

Rotala ramosior 71

Rubiaceae 76, 142

Rubus argutus 75, 132: betulifolius
75, 132; cuneifolius 133; flagellaris

79, 133; trivialis 75, 133
Rudbeckia hirta 59, 145
Ruellia caroliniensis 54; ciliosa 142
astatulus

Sabatia 234; angularis 63, 139; an-
gustifolia 139; brachiata 63, 139;
campanulata 63, 139

Sacciolepis striata 125

Sagina decumbens 56, 131

Sagittaria australis 29; cuneata 29;
engelmanniana 122; falcata 122;
graminea 29, 54; lancifolia 122;
latifolia 29, 54, 122; longirostrata

Salicaceae 33, 76, 129

Salix 24, 99; bebbiana 33; candida
33; nigra 76, 129; pedicellaris 34;
petiolaris 34; serissima 34; su
sericea

Salsola 150; depressa 158; kali 382;
linearis 157; maritima 155; salsa
157, B americana 154, var. amer-
icana 157

Salvia azurea 68, 118, 140; lyrata
68, 140

Salvinaceae 39

Sambucus canadensis 56, 142

Sanguinaria canadensis 73

Sanicula candensis 78, 138: smallii

Sapium sebiferum 135

Sapotaceae 76

Sarcostemma cynanchoides 92

Sarracenia purpurea 35

Sarraceniaceae 35

Sartwellia flaveriae 86; gypsophila
6; mexicana 86; puberula 82, 86

Sassafras albidum 69, 131, 364; sas-

Saxifragaceae 76, 132

Scheuchzeria palustris 29

Schizachyrium scoparium 66; ten-

erum 66
Schizaeaceae 76, 121, 283, 289

Schoenus rariflorus 45
Schrankia microphylla 70, 134

362; scirpoideus 319; smithii 32;
supinus 361, 362; torreyi 32; um-
bellatus 340

Seleria ciliata 62, 127; flaccida 62;

127; torreyana 127; triglomerata
62, 127; verticillata 32, 361, 363

Sclerochloa dura 221

Scoparia dulcis 77, 141

Scrophulariaceae 36, 76, 89, 141

Scutellaria elliptica 69, 140; inte-
grifolia 69, 140, 361, 365; lateri-
flora 69; ovata var. bracteata 69:
pilosa 14

Sebastiania fruticosa 63

Secale cereal 5

Selaginella anode 77, 285, 286, 297,
364; ludoviciana 285, 286, 297
riddellii 364

Selaginellaceae 77, 283, 286

Selatium assurgens 184

Senecio glabellus 40, 59, 145; lo-
batus 145; vulgaris 393; war-
nockii 86

Serenoa repens 127; serrulata 127

Sericocarpus' bifoliatus 59, 143;
linifolius 148

Sericodes greggii 83, 89

Serinea oppositifolia 144

Sesban macrocarpum 134

Sesbania punicea 70

Sesuvium verrucosum 81, 8

Setaria geniculata 66, 125; glauca
66; viridis 66

Seymeria cassioides 77; pectinata

Sherardia arvensis 142

Sibara virginica 132

Sida 227; angustissima 228, 229.
caespitosa 229; cerradoensis 228:
chalchaquensis 228; diffusa 229;
elliottii 228; filicaulis 229; fili-
formis 229; hyalina 227, 228; lind-
heimeri 228, 229; ovata 229; pi-

upina 229; tragiifolia
i vaniesnta 228, 229; xanti 227,

Sidus sidarum 227

Sieglingia americana 125; chap-
manit 125

Silene antirrhina 56, 131; ovata 56

Silphium dentatum var. gatesii 145:
gatesii 59, 145

Sisyrinchium angustifolium 68; are-
nicola ; carolinianum 129;
exile 129; fuscatuwm 129; mucro-
natum var. atlanticum 129; rosu-
latum 68, 129

Sitilias caroliniana 145

Smilax auriculata 118, 128; bona-

Solanaceae 77, 89, 141
Solanum americanum 77, 141; caro-

418

linense 77, 141; nigrum 77; sisym-
briifolium 77, 141
Solidago altissima 59, 145; arguta
367, var. boottii 361, 366; boottii
i canadensis

flexicaulis 361, 367; gigantea 59,
var. gigantea 67, var. leio-
phylla 367; graminifolia 145; mi-
crocephala 145; odora 59, 118,
145; patula 59, var. strictula 145,
361, 367; puberula var, pulveru-
lenta 145; rugosa 59, 145, var.
celtiifolia 145: salicina 145; stric-
a 145; tortifolia 145; ulmifolia 59,

Soliva pterosperma 59, 145, 220
Sonchus asper 59, 145; oleraceus
5

59, 14

Sorbus arbutifolia 133
orghastrum avenaceum 66, 394;
elliottii 66, 125; nutans 125

Sorghum almum 221; bicolor 66;
halepense 66, 125, 221

Sparganiaceae 27,

Sparganium americanum
androcladum 27, 122, 221; chloro:
carpum 27

Spartina pectinata 364

Specularia biflora 1438; perfoliata

SPELLMAN, D.L. 92

Spermolepis divaricata 138; echi-
nata

Sphaeralcea subhastata 88

Sphenopholis filiformis 66, 125; ob-
tusata var. major 66, var. ob-
tusata 67, 125

Spigelia marilandica 71

Spilanthes americana var. repens
59, 145; repens 145; cernua 73,
364; gracilis 129; laciniata 73;
lucida 33; odorata 361, 364; prae-
cox 119, 129; romanzoffiana 33;
vernalis 73, 129

Spirodelia 363; oligorhiza 70; poly-
rhiza 70

Sporobolus clandestinus 67, 125;
indicus 67, 125; junceus 67, 118,
125; macer 67; poiretti 125; sil-
veanus 394

Stachys floridana 69, 120, 141; tenu-
ifolia 69

Stellaria media 56, 131

Stenophyllus ciliatifolius 125

Stenotaphrum secundatum 365

Stephanomeria pauciflora 86

Sterculiaceae 77, 136

Stewartia malacodendron 77

Stillingia sylvatica 118, 135

Stipa avenacea 67

Streptanthus carinatus 88

Striga asiatica 225

Strophostyles ae 70; umbellata
70

STROTHER, JOHN L. 369
Strotheria 90; gypsophila 86

Stylosanthes biflora 70, 134
Styraceae 77, 139
Styrax americana 77, 139; grandi-

olia 77
Suaeda 147, 249; americana 149
150

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erecta 161; fernaldii 172; fruti-
cosa 163, 64, var. multiflora 163;

1
157; maritima 147, 149, 152, 155,
157, 163; mexicana 149, 152, 170;
minutiflora 161; moquini 168; ni-

gra 163, 164; nigrescens 151, 169,

Symplocaceae 77, 139

Symploces tinctoria 77, 139

Syntherisma lineare 123; sangui-
nale 123; serotinum 123; villosum

Tamarix gallica 365
araxacum officinale 59, 145; tara-
xacum 145

Taxodiaceae 77

Taxodium distichum 77

TAYLOR, CONSTANCE E. 361

TAYLOR, JOHN R. 361

Tecoma radicans 141

Telanthera philoxeroides var. acu-
tifolia 130

Tephrosia chrysophylla_ 134; florida

Thaspium barbinode 78; trifoliatum
var. flavum 78

Theaceae 77, 136

Thelesperma 305; longipes 86; me-
gapotamicum 82, 86, 87; ramosius
82, 87; scabridulum 87

Theligonum 96,

Thelypteris 291; dentata 74, 120,

; ; ; hexagonoptera 74,

291, 292, 2938; kunthii 292, 293;

419

normalis 75, 121, 292: novebora-

Thyrsanthema semiflosculare 148

Tilia caroliniana

Tiliaceae 77

Tillandsia usneoides 55, 128

Tila floridana 395

Tillaea aquatica 351, 364

Tipularia discolor 73

Tmesipteris 93

Tofieldia glutinosa 33; racemosa 128

Tovara virginiana 74

Trachelospermum difforme 54

Tradescantia hristicaulis 128: ohi-
ensis 56, 128; reflexa 128

Tragia smallii 63, 135; urens 63;
urticifolia 63

Trautvetteria 2, 10; carolinensis 10

Trepocarpus aethusae 78,

Tretorhiza 174, 17

Triadenum Ut OE 136

lineare 141; setaceum 69, 141
Tridens ambiguvs 67; carolinianus
67, 125; chapmanii 67; flavus 67,
var. chapmanii 125; strictus 67,
394
Trifolium arvense 70; So pe otc

Triglochin maritimum 28; palustre
28

Trilisa odoratissima 59, 118, 145
Trillium cuneatum 71; texanum 221
Triodanis biflora 56; perfoliata 56

420

Triplasis americana 67, 125; pur-
purea 67, 364

Tripsacum dactyloides 394

Tripterospermum 232, 245

Triticum aestivum 67, 125

TURNER, B.L. 22

Typha 218; latifolia 122

Typhaceae 122

Ulmaceae 77

Ulmus alata 78; americana 78; ru-

ra 78

Umbelliferae 35, 40, 220

Uniola laxa 123; longifolia 123

URBATSCH, LOWELL E. 298

Urticaceae 34,

Utricularia biflora 70, 142; cornuta
36; inflata 361, 365; intermedia
36: juncea 142; minor 36; subu-
lata 142, 361, 366

Uvularia perfoliata 71; sessilifolia
71
Vaccinium 120; arboreum 62, 118,

64, darrowii 62, 118, 139; el-
liottii 62, 118, 189; myrsinites 139;
oxycoccos 36; stramineum :
var. sericeum 139; tenellum 139;
virgatum 62,

Vaginaria 316; richardi 319

Valeriana ciliata 36

Valerianaceae 36, 78, 142

Valerianella radiata 78, 142

Varilla mexicana 87

Verbascum thapsus 77, 141

Verbena bonariensis 78; brasilien-
sis 78, 140; canadensis 78; carnea
78, 118, 140; carolina 140; el
78; hastata 364; officinalis 140;
rigida 78, 140; tenuisecta 78, 140;
urticifolia 78

Verbenaceae 78, 89, 140

Verbesina encelioides 87; helian-
thoides 59; occidentalis 145; vir-
ginica 59, 145, 395; walteri 59

Veronia 264; altissima 59, 145; an-
gustifolia 118, 145, 265, 277, var.

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—

texana 265, 277; Daldwann 270,

276, 277, 279; gigantea 145, 278;
graminifolia 145; .granienifolia
265, 277; greggii 264, 268, 269, 270,
271, 272, 274, 275, 276, 277, 278,
ssp. ervendbergii 267, 278, 279,
280, ssp. faustiana 267, 278, 280,

schaffnert 267, 279, 280, var. greg-
palmeri 265, 279;
lindheimeri 281; missurica 278;

65,
267, 270, ae 272, 273, 274, 276,
277, 278, 280
Veronica arvensis 77, 141; peregri-
na 77,
Veronicastrum virginicum 364
Viburnum dentatum 56; nitidum 56;
nudum 56, 142; rufidulum 56
Vicia acutifolia 262; americana 259,

gantea 262; humilis 262;
worthii 255, 257, 258, 259, 260, var.
occidentalis 257, 258, 259, 260;

laxiflora 258, 260, var. texana 258,
260; minutiflora 262;
262; pulchella 259; tetrasperma

Viola brittoniana 137; esculenta 78;

paniesquil 78: sepieminna 137;
triloba 78; vicinalis 137; walteri

Violaceae 78, 137

Viscaceae 79

Vitace

itis aestivalis 136, var. argentifo-

tundifolia 79, 136; vulpina 79, 136
Voyria 234; aphylla 239
Vulpia bromoides 221; dertonensis
221; myuros 67; octoflora 67
Wahlenbergia marginata 56, 143
WHETSTONE, R. DAVID 397
WILLIAMSON, GENE 248
Willughaeya scandens 145
Wisteria frutescens 70; sinensis 70,

Wolffia papulifera 32; punctata 361,

363

Wolffiella floridana 32, 70

Woodsia 291, 292; obtusa 292, 293

Woodwardia 291, 295; areolata 119,
121, 295, 296; virginica 75, 121,
296, 297

Xanthium strumarium 59, var. gla-

bratum 145

Barney L. Lipscomb
Herbarium

Southern Methodist University
Dallas, Texas 75275

421

Xanthocephalum 298
Xanthorhiza 2, 11; simplicissima 11,

Xolemia catesbei 176
Xylopleurum speciosum 138

ambigua 127; caro-
liniana 79, 127; communis 127;
difformis var. difformis 32, 79,
127; fimbriata 79, 127; flexuosa
127; iridifolia 79, 127; jupicai 127;
torta 32

Youngia japonica 59

Yucca aloifolia 128;
128; smalliana 71

Zanthorhiza apitfolia 131

Zebrina pendula var. quadrifolia

filamentosa

Zephyranthes atamasco 129
Zozania aquatica 29

Zornia bracteata 134
Zygophyllaceae 89