Full text of "Rhodora"
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JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by
ALBION REED HODGDON, Editor-in-Chief
ALBERT FREDERICK HILL
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON Associate Editors
RADCLIFFE BARNES PIKE
ELIZABETH ANNE SHAW
STEPHEN ALAN SPONGBERG
Vol. 74 March, 1972 No. 797
CONTENTS:
Revision of the Genus Melampodium (Compositae:
Heliantheae). Tod F. Stues8y nouses 1
In Memoriam Charles Schweinfurth, 1890-1970.
MEM rc ueniens siet tec qu eia s HERE 71
Nomenclatural Changes in Lesquerella.
Reed C. Rollins and Elizabeth A. Shaw .............. ne 76
The Parker Cleaveland Herbarium of Bowdoin College.
Edward Hehre, A. R. Hodgdon and R. B. Pike ............. 80
Studies on the Audouinella microscopica (Naeg.) Woelk.
Complex (Rhodophyta). William J. Woelkerling ............ 85
Nomenclatural Clarification of Two Species of Verbesina
(Compositae) Endemic to Florida. James R. Coleman ... 97
(Continued on Inside Back Cover)
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JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUD
Vol. 74 March, 1972 No. 797
REVISION OF THE GENUS MELAMPODIUM
(COMPOSITAE: HELIANTHEAE)'
Top F. STUESSY
Probably every plant collector in Mexico at some time
during his travels has come upon large fields of weedy
yellow-headed Compositae. More often than not, these herbs
belong to the genus Melampodium. Such abundance of num-
bers, however, is not matched at all by our knowledge of
the group, and in fact, the only comprehensive treatment
of the genus is B. L. Robinson's synopsis published in 1901.
Although this study brings together partial synonymy and
provides a key for the group, all taxa are not adequately
described nor does the key allow for easy and accurate iden-
tification. Many of the recent specimens being collected in
Mexico and Central America have been extremely difficult
if not impossible to identify satisfactorily. This incomplete
understanding of relationships within Melampodium sug-
gested that a thorough revision should be undertaken.
The present investigation is based on five years of study,
including four months of field work and an examination of
over 8000 herbarium specimens. The treatment presented
here is essentially a straight-forward revision of Melampo-
dium. Evolutionary considerations regarding the whole
genus have been kept to a minimum as these evaluations
have been the subject of a separate paper (Stuessy, 1971a).
*Publication No. 795 from the Department of Botany, The Ohio
State University, Columbus.
2 Rhodora [Vol. 74
Comments on probable phyletic relationships among se-
lected species, however, are interspersed throughout the
text, and data from experimental studies either in progress
or already published are brought to bear on the relation-
ships when appropriate. It is realized that many system-
atic problems still persist within Melampodium, but it
is hoped that the present study dispels the confusion that
has prevailed.
TAXONOMIC HISTORY
The first description of Melampodium appeared in Lin-
naeus’ Hortus Cliffortianus (1738) which was the refer-
ence cited in the Species Plantarum of 1753. Linnaeus in
his Critica Botanica of 1737 (p. 76) clearly indicated that
the name he gave to the genus was derived from “Melam-
pus medicus graecus." Apparently overlooking this ex-
planation, many later workers (e.g., Gray, 1884; Cockerell,
1905) erroneously have believed the name to come from
the Greek words meaning ''black-foot."
From the time of Linnaeus' original description of M.
americanum in 1738, the number of the species described
in Melampodium has increased to over 88. Difficulties of
specific cireumscription and synonymy, resulting from the
many described taxa, have been surpassed by the more
perplexing problems dealing with generic and sectional de-
limitation. Because discussions have already been pub-
lished on the history of generic problems associated with
Melampodium (Stuessy, 1969b, 1970a), the following com-
ments will emphasize the development of attempts to sub-
divide the genus.
Humboldt, Bonpland, and Kunth (1820) established the
first subdivisions of the genus. Based primarily on the
genera that were included in synonymy, they designated
three groups (here recognized as subgenera) : Melampodia,
Dysodia (Rich. in Pers.) H.B.K., and Alciniae (Cav.)
H.B.K. No descriptions or comments were added to explain
the rationale for the groupings beyond the inclusion of
species.
1972] Melampodium — Stuessy 3
Morphological criteria for the sections later recognized
by Candolle (1836) were first stated by Cassini (1829).
Cassini recognized the separate genera, Melampodium, Al-
cina, and a new genus, Zarabellia, all of which had inner
phyllaries tightly enclosing and fused with a single ray
achene, but each of which possessed distinctive characters
as follows:
Melampodium L. (1) Inner phyllaries each crowned with
a large hooded appendage (2) Sterile disc ovaries very
large.
Zarabellia Cass. (1) Inner phyllaries with no hood but
with two small valve-like processes (2) sterile disc ovaries
nearly or completely absent.
Alcina Cav. (1) Inner phyllaries with no hood or valve-
like processes and smooth at the sides (2) Sterile disc
ovaries nearly or completely absent.
Candolle (1836) put Zarabellia and Alcina into Melam-
podium but maintained the taxa as three sections based on
Cassini’s bract differences: Humelampodium DC., Zara-
bellia (Cass.) DC., and Alcina (Cav.) DC. Later, Bentham
and Hooker (1873) and Hoffmann (1890) did not recognize
sections in Melampodium, perhaps to stress the morpholog-
ical unity of the genus (Turner and King, 1962). Baker,
treating Brazilian species (1884), suggested two subgen-
era: Dysodium (Rich. in Pers.) H.B.K., and Unzia (L.f.)
Baker, which were characterized by pedunculate and sub-
sessile heads, respectively. Baillon (1882) departed signif-
icantly from previous authors by lumping Acanthospermum
Schrank and Lecocarpus Dene. into Melampodium, and
these included genera formed the basis of his two recog-
nized sections: Acanthospermum (Schrank) Baillon and
Lecocarpus (Dene.) Baillon.
Robinson in his synopsis of Melampodium (1901) sub-
merged section Alcina (containing the single species, M.
perfoliatum) into section Zarabellia. The two other sec-
tions of Candolle were retained on the basis of the hood
versus non-hood bract character. Robinson acknowledged
the variability of this feature but wrote that (p. 455)
4 Rhodora [Vol. 74
* ., the presence or absence of a hood can usually be
determined readily, and the two sections Eumelampodium
and Zarabellia may conveniently be retained.”
Using cytological data, Turner and King (1962) re-
opened considerations of sectional subdivision. Based on
chromosome counts from 89 populations of 21 species’, the
taxa of section Melampodium (Humelampodium DC.) were
shown to be on a base of « — 10, and the species within
section Zarabellia to be multibasic with x = 9, 11, 12, 16°
and 23+. In addition to bract and chromosome base num-
ber criteria, “small plants" and “densely pubescent or to-
mentose (rarely merely hispid) foliage" were suggested as
characterizing section Melampodium, with “heterogeneous”
morphology and “hispid pubescence” more indicative of
section Zarabellia.
No one has ever proposed series within the two recog-
nized sections of Melampodium. In fact, Robinson experi-
enced frustration in attempting to create such order (1901,
p. 455): “Much difficulty has been experienced in giving
the species a natural sequence, and after many efforts the
hope of securing such an arrangement has been abandoned.”
Turner and King (1962) suggested that within the more
morphologically heterogeneous section Zarabellia some for-
mal subdivisions might be desirable after further study.
They mentioned several informal groups such as: (1) “the
small-headed, annual species having a chromosome base
of x = 9"; (2) “those rhizomatous species centering about
M. montanum with chromosome numbers on a base of x —
11"; and (3) "those annual species with large involucral
bracts and smooth achenes having chromosome numbers of
n = 11 and 12."
*See Stuessy (1971b) for a reassignment of these chromosome
counts to the presently recognized taxa.
‘Due to the generic transfer of M. camphoratum (n = 16) back
into Umzia (Stuessy, 1969b), this number no longer belongs in
Melampodium.
‘Because of recent counts of n — 12 for M. dicoelocarpum (Stuessy,
1971b), the n — 23 count for this species seems best interpreted as
an aneuploid at the tetraploid level on a base of x — 12.
1972] Melampodium — Stuessy D
Figs. 1-4. Disc florets of representative species of Melampodium
showing the two different types of abortive ovaries. Figs. 1 & 2,
M. leucanthum, Stuessy 752 (OS); Figs. 3 & 4, M. divaricatum,
Stuessy 547 (OS). Drawn from fresh material. The corollas and
stamens in Figs. 2 and 4 have been removed, but they are the same
florets as in Figs. 1 and 3, respectively. The arrow in Fig. 1 points
to a marked annular constriction at the point of ovary and corolla
tube juncture, which is characteristic of species in sect. Melampodium.
That same marked constriction is absent from the florets of species
in the other five sections of the genus (Fig. 3). Both types of ovaries
are capped by ovarian discs (Figs. 2 & 4).
In the present treatment I depart significantly from pre-
vious workers by recognizing six sections in the genus.
Based on coherence of morphology and chromosome num-
bers, section Melampodium seems a phyletic unit and there-
fore is retained, although here for the first time it is di-
vided into five series. Section Zarabellia s.l. of previous
authors, however, has been circumscribed in the past by
6 Rhodora [Vol. 74
the absence of hoods of the inner phyllaries (to be dis-
cussed in detail later) and by the possession of chromo-
some numbers other than on a base of x = 10 (i.e, xz = 9,
11, and 12). The only positive feature shared by all taxa
of the section is the rudimentary ovary of the disc florets
which contrasts markedly with the more conspicuous ovary
of those taxa in section Melampodium (Figs. 1-4; these
ovary differences mentioned earlier by Cassini, 1829, as
having generic significance). The diversity of other mor-
phological features and chromosome numbers in taxa of
section Zarabellia s.l., however, suggests that the one uni-
fying ovary feature may have been derived by parallel
evolution (Stuessy, 1971a). From this phyletic viewpoint
section Zarabellia s.l. seems an artificial unit and is there-
fore divided into five separate sections based on chromo-
somal and morphological differences. Two of the sections,
Zarabellia, s.str. and Rhizomaria, correspond respectively
to the first two informal groups mentioned by Turner and
King (1962). A list of the recognized sections, series, and
included taxa follows:
Synopsis of Classification of Melampodium L.
I. Section Melampodium
1. Series Melampodium
1. M. americanum L. 2. M. diffusum Cass. 3. M.
pilosum Stuessy 4. M. longipes (A. Gray)
Robins. 5. M. linearilobum DC.
2. Series Leucantha Stuessy
6. M. leucanthum Torr. & A. Gray 7. M. cine-
reum DC. Ta. M. cinereum DC. var. cinereum
Tb. M. cinereum DC. var. hirtellum Stuessy
Tc. M. cinereum DC. var. ramosissimum (DC.)
A. Gray 8. M. argophyllum (A. Gray ex
Robins.) Blake
3. Series Sericea Stuessy
9. M. sericeum Lag. 10. M. pringlei Robins. 11.
M. strigosum Stuessy 12. M. longicorne A.
Gray 13. M. nayaritense Stuessy
1972] Melampodium — Stuessy 7
4. Series Cupulata Stuessy
14. M. cupulatum A. Gray 15. M. appendiculatum
Robins. 16. M. sinuatum Brandg. 17. M. rosei
Robins. 18. M. tenellum Hook. & Arn. 19. M.
glabribracteatum Stuessy
5. Series Longipila Stuessy
20. M. longipilum Robins.
II. Section Zarabellia (Cass.) DC.
21. M. longifolium Cerv. ex Cav. 22. M. mimuli-
folium Robins. 23. M. gracile Less. 24. M.
microcephalum Less. 25. M. paniculatum
Gardn.
III. Section Serratura Stuessy
26. M. divaricatum (Rich. in Pers.) DC. 27. M.
costaricense Stuessy 28. M. dicoelocarpum
Robins. 29. M. tepicense Robins. 30. M. sina-
loense Stuessy
IV. Section Bibractiaria Stuessy
31. M. bibracteatum S. Wats. 32. M. repens Sessé
& Moc.
V. Section Rhizomaria Stuessy
33. M. montanum Benth. 33a. M. montanum
Benth. var. montanum 33b. M. montanum
Benth. var. viridulum Stuessy 34. M. aureum
Brandg.
VI. Section Alcina (Cav.) DC.
35. M. perfoliatum (Cav.) H.B.K. 36. M. glabrum
S. Wats. 37. M. nutans Stuessy
DISTRIBUTIONAL SUMMARY
Melampodium is a tropical and subtropical genus mostly
restricted to Mexico and Central America (Fig. 5) with
four species located in the southwestern United States, and
three species scattered in Colombia and Brazil. Although
the Brazilian distributions could indicate South American
origins for the two species, M. paniculatum and M. divari-
catum, the fact that these species are found abundantly
also in Central America and Mexico suggests that their
presence in Brazil is due to recent introductions by man.
8 Rhodora [Vol. 74
Fig. 5. Map of North and South America showing generalized
distribution of Melampodium. Collections from Burma and the Philip-
pine Islands not shown.
The exceedingly weedy nature of M. divaricatum and
M. perfoliatum has allowed successful introductions of the
genus to Burma, Cuba, Puerto Rico and the Virgin Islands.
Another Mexican species, M. diffuswm, has become estab-
lished near Manila in the Philippine Islands.
1972] Melampodium — Stuessy 9
Melampodium occupies primarily moist habitats from
mountain pine-oak to tropical deciduous forests. Only two
species, M. leucanthum and M. cinereum, are clearly xero-
morphic, found in the drier regions of northern Mexico and
adjacent United States.
SPECIFIC AND VARIETAL CATEGORIES
It would be a formidable task to treat 37 species of a
genus exhaustively with many experimental approaches to
discover all the effective isolating mechanisms. As a re-
sult, in the present study a morphological species concept
has been stressed. With the exception of chromosome num-
bers for many taxa, only in a few cases have data from ex-
perimental work been used to define more accurately the
reproductive limits of each taxon. It is assumed, however,
that the qualitative and quantitative morphological discon-
tinuities formally recognized in this study do represent
genetic differences that in some fashion are responsible for
maintaining the integrity of each specific and varietal unit.
An attempt has been made to treat specific and varietal
categories consistently with regard to the data available
in each case. Varieties are regarded as morpho-geograph-
ical subdivisions of a species (Kapadia, 1964) that pre-
sumably reflect genetic differences.
MORPHOLOGY AND TAXONOMIC CRITERIA
As an aid to understanding the specific and varietal cate-
gories used in the present study, the taxonomic value of
various morphological features is discussed below:
Habit. — Both herbaceous annuals and suffruticose or
rhizomatous perennials are found in the genus. The peren-
nial habit is the less common condition found only in the
following seven species of sections Melampodium and Rhi-
zomaria: M. americanum, M. leucanthum, M. cinereum, M.
argophyllum, M. sinuatum, M. montanum, and M. aureum.
Roots. — All taxa of the genus are tap-rooted except for
the two rhizomatous perennial species of section Rhizo-
maria: M. montanum and M. aureum.
10 Rhodora [Vol. 74
Leaves. — The shape, size, vesture, and type of margin
of leaves are often useful as discriminating taxonomic
characters at various levels in the hierarchy from section
to variety. However, in some taxa leaf shape and size can
be quite variable, especially in the widespread species, and
this plasticity has been the cause of recognition of forms
as species or varieties by previous workers (e.g., in M.
divaricatum).
Vesture. — The surface of various organs ranges from
glabrous to copiously sericeous, and the vesture often can
be used as a reliable taxonomic character.
Heads. — Many features of the outer and inner involu-
cral bracts and the ray florets are of diagnostic value.
Within limits, the number of disc and ray florets is some-
times useful as a specific indicator, and as mentioned pre-
viously, the shape of the abortive ovaries of the disc florets
is useful as a criterion for delimiting section Melampodium.
“Fruit” in this treatment refers to the mature ray achene
and enveloping inner involucral bract (Robinson, 1901).
The fruits are often capped by extensions of the inner in-
velucral bracts of varying shapes and sizes upward from
the achene apexes (Figs. 6-9). The presence of one partic-
ular type of extension called a “hood” (the term apparently
first used by Cassini, 1829, and followed by most subsequent
authors) correlates well with other features of section
Melampodium. Within most species, however, the hood
size, shape, and appendage vary considerably. In fact, the
amount of variation prevalent among fruits in different
plants of a single population (Fig. 10) and sometimes even
among bracts in a single head (cf. Fig. 9E in Stuessy,
1970b) is astonishing. Such permutations, especially in the
sculpturing of the lateral outer surfaces of the inner phyl-
laries, have been judged by some students of the group
(e.g., Robinson, 1901) as indicative of varieties or even as
distinct species. However, with few exceptions (such as in
M. divaricatum and M. dicoelocarpum) the lateral bract
surface features usually are not reliable as taxonomic char-
acters.
1972] Melampodium — Stuessy Bi
Figs. 6-9. Representative fruits of Melampodium illustrating
variation in achenal apexes. Fig. 6, M. glabrum, apex moderately
sculptured, Stuessy 709 (OS); Fig. 7, M. longifolium, apex with
abaxial protuberance, Sprengel s.n. (P); Fig. 8, M. longipilum, apex
with adaxial awn, Stuessy 634 (OS); Fig. 9, M. leucanthum, apex
with hood, Stuessy 752 (OS).
Fig. 10. Representative fruits of Melampodium linearilobum from
a single population in Michoacán, Mexico (Stuessy 698, OS) illus-
trating variations in structure of the inner involucral bracts.
12 Rhodora [Vol. 74
TAXONOMIC TREATMENT
MELAMPODIUM L.
Melampodium L. [Hort. Cliff. 425. 1738.] Sp. Pl. 921.
1753. Gen. Pl. ed. 5. 392. 1754. Type species: Melampo-
dium americanum L.
Cargilla Adans. Fam. 2:130. 1763. nom. superfl. in-
cluding Chrysogonum L. and Melampodium L.
Alcina Cav. Ic. 1:10. t. 75. 1791. Type species: Alcina
perfoliata Cav. = Melampodium perfoliatum (Cav.) H.B.K.
Dysodium Rich. in Pers. Syn. 2:489. 1807. non Dys-
sodia Cav. 1802. Type species: Dysodium divaricatum Rich.
in Pers. = Melampodium divaricatum (Rich. in Pers.) DC.
Melampodium L. subg. Alcina (Cav.) H.B.K. [as subg.
Alciniae]. Nov. Gen. Sp. 4:274. 1820.
Melampodium L. subg. Dysodium (Rich. in Pers.) H.B.K.
[as subg. Dysodia]. Nov. Gen. Sp. 4:273. 1820.
Melampodium L. subg. Melampodium [as subg. Melam-
podia]. H.B.K. Nov. Gen. Sp. 4:272. 1820.
Camutia Bonat. ex Steud. Nom. Bot. 146. 1821. pro syn.,
nom. nud. Based on Camutia perfoliata Bonat. ex Steud.
Carnutia [attributed to Bonat. ex Steud. by] Baker in
Martius, Fl. Bras. 6(3) :159. 1884. Orthogr. var.
Zarabellia Cass. Dict. Se. Nat. 59:240. 1829. [non
Neck. 1790. invalid fide Art. 20, Intern. Code Bot. Nomen.]
Type species: Zarabellia rhomboidea Cass. = Melampo-
dium longifolium Cerv. ex Cav.
Annual herbs to perennial subshrubs. Tap roots in all
but 2 species (M. aureum and M. montanum with fibrous
roots and rhizomes). Stems decumbent to erect, terete to
finely striate, dichotomously branched with peduncles aris-
ing from the middle and upper dichotomies. Peduncles very
short and stout to long and filiform. Leaves opposite, de-
cussate, linear to ovate-rhombic, at apex acuminate to ob-
tuse, at base attenuate to auriculate-connate, with upper
surfaces glabrous to pilose, with lower surfaces glabrous
to sericeous, at margin entire to toothed. Heads solitary.
Receptacle paleaceous, convex, sometimes elevated on a
short cylindrical stalk 2-3 mm above the outer involucre.
1972] Melampodium — Stuessy 13
Involucre biseriate. Outer involucre spreading to cupulate;
bracts 2-5, more or less equal, with margins entire, separate
to connate more than 2/3 their length, with adaxial surface
glabrous. Innermost phyllaries each enclosing a single ray
achene, completely covering or sometimes partially open at
apex, often extending upward into a hood or other apical
appendage (Figs. 6-9). Ray florets 3-13; ligules varying
shades of yellow or cream-white (2 species), at the apex bi-
or tri-dentate, attached on adaxial side of achene apex; tube
nonexistent or very short; style branches filiform, flattened,
obtuse at the apex, with stigmatic surfaces marginal and
enlarged and running the length of the branches, recurved
when fruit maturing; achenes asymmetrically obovoid and
laterally compressed. Disc florets 3-110; corollas yellow-
green to yellow-orange, regular, 5-lobed (rarely 4-lobed),
with lobes partially reflexed or extended at right angles,
throat salverform-funnelform; anthers brown, exserted
from corolla 1/3-1/2 their length; style linear-capillaceous,
unbranched, exserted from within anthers; ovary sterile,
capped by a disc; pappus absent. Paleae scarious, con-
duplicate around disc corollas. Chromosome numbers, n =
9, 10, 11, 12, 18, 20, 23, 25 + 1, 27, 30, and 33.
ARTIFICIAL KEY TO TAXA
a. Ligules white. (b)
b. Outer phyllaries connate more than half their
length. (c)
c. Stems and leaves strigillose; leaves with margins
usually entire-sinuate (less often pinnately
lobed) ; heads 20-37 mm diam‘; ligules 7-13 mm
long, 2.5-8 mm wide; plants 15-50 em tall
MEE cs qx onda se sees TATEM 6. M. leucanthum
c. Stems and leaves canescent-tomentose; leaves
with margins usually pinnately lobed-parted
(less often entire) ; heads 10-23 mm diam; lig-
ules 2.5-9 mm long, 1.5-4.5 mm wide; plants 12-
22 em tall ................ 8. M. argophyllum
'In the keys and descriptions head diameter excludes the outer
involueral bracts.
14 Rhodora [Vol. 74
b. Outer phyllaries connate a third or less their
length. (d)
d. Leaves with basal margins hispid with hairs 0.6-
d.
1.5 mm long; stems strigose-hispid with hairs
0.2-1.5 mm long . 7b. M. cinereum var. hirtellum
Leaves with basal margins strigose with hairs
0.3-0.6 mm long; stems strigose with hairs 0.1-
0.8 mm long. (e)
e. Leaves linear, 7-32 mm long, 2-5 mm broad;
heads 3-5 mm tall; outer involucre 4.5-9 mm
diam; outer phyllaries 2.7-4 mm long, 1.5-
2.8 mm wide; hood 0.8-1.5 mm tall ........
ZEN 7c. M. cinereum var. ramosissimum
e. Leaves linear-oblong, 12-55 mm long, 1-14
mm broad; heads 5-8 mm tall; outer involucre
7-14.38 mm diam; outer phyllaries 3.5-7.5 mm
long, 2.3-5 mm wide; hood 1.3-2.9 mm tall ..
es Ta. M. cinereum var. cinereum
a. Ligules yellow. (f)
f. Outer phyllaries 2. (g)
g.
g.
Leaves oblong to obovate-oblanceolate; ray flor-
ets 5-6 (rarely 3); stems erect to decumbent;
plants subaquatic ........ 31. M. bibracteatum
Leaves obovate; ray florets 2-3; stems prostrate ;
plants terrestrial .............. 32. M. repens
f. Outer phyllaries 3-5. (h)
h.
Outer phyllaries 3. (i)
i. Heads 3-5.5 mm diam; ligules usually less
than 2 mm long; peduncles copiously stipi-
tate-glandular ........ 25. M. paniculatum
i. Heads 5-15 mm diam; ligules usually more
than 2 mm long; peduncles strigillose to his-
pid to weakly stipitate-glandular. (j)
j. Leaves subauriculate at base ..........
ee ee ee 23. M. gracile
j. Leaves attenuate to obtuse at base. (k)
k. Leaves with tapering petioles 1-3 cm
long; inner phyllaries open in a small
1972] Melampodium — Stuessy 15
hood at achene apex, often extended
Into-awn ... 0 . 9T. M. nutans
k. Leaves sessile or with petioles 0.1-0.4
cm long; inner phyllaries completely
closed over achene apex, with no ex-
tended appenddgsg 8541 eas
Po ote alg s E 24. M. microcephalum
h. Outer phyllaries 5. (1)
l. Leaves sericeous beneath. (m)
m. Outer phyllaries with the margin scari-
OUS ................. 5. M. linearilobum
m. Outer phyllaries with the margin herba-
ceous. (n)
n. Leaves narrowly ovate to lanceo-
late. (0)
o. Plants perennial; heads 11-17 mm
diam; ligules 4-6 mm long ......
i las ate RE l. M. americanum
o. Plants annual; heads 6-11 mm
diam; ligules 2.5-4.5 mm long. (p)
p. Plants 8-28 cm tall; stems pil-
ose; outer phyllaries ovate to
narrowly ovate; leaves copious-
ly sericeous beneath .........
PNIS 3. M. pilosum
p. Plants 25-40 cm tall; stems stri-
gillose-hispidulous; outer phyl-
laries ovate; leaves moderately
sericeous beneath ..... .....
DENN ee 2. M. diffusum
n. Leaves linear, oblanceolate, or deeply
pinnately divided. (q)
q. Ligules 2 mm long or less. (r)
r. Leaves linear and entire to
deeply pinnately divided ; ligules
ca 2 mm long; heads 8-11 mm
diam; disc florets 15-25 ......
E esuna 13. M. nayaritense
16
Rhodora [Vol. 74
r. Leaves linear-oblong to oblance-
olate, often lobed or parted;
ligules ca 1 mm long; heads 4-
8 mm diam; disc florets 2-
12. (s)
s. Peduncles 5-22 mm long;
dise florets 5-12; ligules
on undersurface yellow at
apex; outer involucre 6-11
mm diam .. 9. M. sericeum
s. Peduncles 1-2 mm long; disc
florets 2-3; ligules on under-
surface purple at apex;
outer involucre 3-4 mm diam
ENDE 10. M. pringlet
q. Ligules more than 2 mm long. (t)
t. Plants annual; stems yellow to
green ......... 4. M. longipes
t. Plants perennial; stems purple
to green .... 1. M. americanum
l. Leaves glabrous or strigose beneath. (u)
u. Outer phyllaries with the margin narrow-
ly scarious. (v)
V.
Peduncles 1.3-1.7 cm long; outer phyl-
laries abaxially glabrous .....
DEM 19. M. glabribracteatum
Peduncles usually more than 2 cm
long; outer phyllaries abaxially sub-
glabrous, strigose or pilose. (w)
w. Plants perennial with fibrous roots
and rhizomes. (x)
x. Heads 19-38 mm diam; ligules
yellow-orange to yellow, 5-12
mm long; disc corollas yellow;
fruit 2-2.3 mm long*; chromo-
*Fruit length in the key and descriptions excludes hoods and ap-
pendages.
1972] Melampodium — Stuessy 17
yellow, 4-7.5 mm long; disc
corollas yellow-green; fruit 1.6-
1.8 mm long; chromosome num-
ber, n = 11. (y)
y. Ligules on undersurface
light green at apex and on
veins .. 33b. M. montanum
var. viridulum
y. Ligules on undersurface
dark purple at apex and on
veins .. 33a. M. montanum
var. montanum
w. Plants annual or perennial with
tap roots. (z)
Z.
Plants perennial; leaves on un-
dersurface copiously tomentose,
at margin markedly sinuate ..
DER op oe Pune 16. M. sinuatum
Plants annual; leaves on under-
surface strigose, at margin en-
tire, lobed or divided. (aa)
aa.Leaves at base attenuate.
(bb)
bb. Outer phyllaries ovate to
narrowly ovate, 4-6 mm
long; heads 7-8 mm tall ..
........ 14. M. cupulatum
bb. Outer phyllaries ovate to
orbieulate, 2.5-3.5 mm
long; heads 3.5-5. mm
tall .... 18. M. tenellum
aa. Leaves at base obtuse to sub-
auriculate. (cc)
cc. Outer phyllaries connate
2/3 their length ........
18
Rhodora [Vol. 74
.. 15. M. appendiculatum
cc. Outer phyllaries connate
less than 1/4 their length
ZEE 17. M. rose
u. Outer phyllaries with the margin herba-
ceous. (dd)
dd. Outer involucre 15-32 mm diam ; outer
phyllaries 6-20 mm long ; fruits 4-7 mm
long; lower leaves conspicuously per-
foliate at base .... 35. M. perfoliatum
dd. Outer involucre 3-10 (—-14) mm diam;
outer phyllaries 2-6(—6.8) mm long;
fruits 1.6-3 (—4) mm long; lower leaves
attenuate to subauriculate at base. (ee)
ee. Leaves markedly petiolate. (ff)
ff. Ligules 3.5-7 mm long. (gg)
gg. Fruit at apex with a flattened
adaxial appendage extending
upward into a cirrhous awn
(Fig. 8) .... 20. M. longipilum
gg. Fruit at apex variously ridged
and sculptured but not as above
EE 26. M. divaricatum
ff. Ligules less than 3 mm long. (hh)
hh. Fruit on lateral surfaces with
2 deep oval cavities ..........
. 28. M. dicoelocarpum
hh. Fruit on lateral surfaces vari-
ously ribbed, tubercled, and
sculptured but not as above.
(ii)
ii. Heads 6-10 mm diam; stems
usually 2-4 mm diam ......
ZEN 27. M. costaricense
ii. Heads 3-5 mm diam; stems
usually less than 2 mm diam.
(jj)
jj. Peduncles 0.8-8 mm long;
1972] Melampodium — Stuessy 19
disc florets 3-7(-12) .....
hee 29. M. tepicense
jj. Peduncles 10-29 mm long;
disc florets ca 14 ........
NS 30. M. sinaloense
ee. Leaves sessile or subsessile. (kk)
kk. Plants subaquatic; stems thick
and yellowish .... 36. M. glabrum
kk. Plants terrestrial; stems greenish
to greenish-purple. (ll)
ll. Ligules less than 2 mm long;
outer phyllaries lanceolate.
(mm)
mm. Fruits with an abaxial pro-
tuberance (Fig. 7) ......
`... 21. M. longifolium
mm. Fruits with hoods (Fig. 9)
or minutely sculptured at
apex. (nn)
nn. Plants usually 15-20 cm
tall; stems ascending;
peduncles usually 0-3 mm
long; leaves at margin
entire-pinnately lobed ;
heads 3-4 mm tall, 4-8
mm diam; outer phyl-
laries lanceolate-elliptic .
e i 11. M. strigosum
nn. Plants usually 20-40 cm
tall; stems erect; pedun-
cles usually 10-20 mm
long; leaves at margin en-
tire to dentate; heads 4-7
mm tall, 7-13 mm diam;
outer phyllaries elliptic to
obovate-rhombic .......
T" 12. M. longicorne
ll Ligules more than 2 mm long;
20 Rhodora [Vol. 74
outer phyllaries ovate or nar-
rowly ovate. (00)
oo. Leaves attenuate at base ....
DENEN 24. M. microcephalum
oo. Leaves subauriculate to ob-
tuse at base. (pp)
pp. Stems copiously pilose near
apex; outer phyllaries
ovate-lanceolate, 5-6 mm
long, at apex acute ......
22. M. mimulifoliwm
pp. Stems moderately hispid-
pilose and weakly stipitate-
glandular near apex; outer
phyllaries ovate, 3.5-5 mm
long, at apex acuminate
INE 23. M. gracile
I. MELAMPODIUM section Melampodium
Melampodium L. sect. Eumelampodium DC. Prodr. 5:518.
1836. Type species: Melampodium americanum L.
Tap-rooted annuals or perennials; leaves linear to ovate,
sessile to markedly petiolate, with margins entire or lobed
or obscurely serrate; outer involucre cupulate, with bracts
5, at margins herbaceous or scarious; ovaries of the disc
florets linear, 1.2-2.3 mm long; tube of disc florets with a
marked basal annular constriction at point of ovary junc-
ture (Fig. 1, arrow) ; fruits usually capped (Fig. 9) with
a hood (absent in M. pringlei, and M. sinwatum; sometimes
absent in M. americanum, M. hispidum, M. longicorne, and
M. sericeum; rudimentary in M. cupulatum) or with a flat-
tended adaxial awn (M. longipilum, Fig. 8) ; chromosome
base number, x — 10. Species 1-20.
1. Series MELAMPODIUM
Annual herbs or suffruticose perennials (in M. ameri-
canum) ; leaves narrowly ovate to linear, often pinnately
divided, sericeous beneath, ligules orange-yellow, more than
1972] Melampodium — Stuessy 21
or rarely about 2 mm long; margins of outer phyllaries her-
baceous or scarious (in M. linearilobwm) ; fruits usually
hooded. Species 1-5. Type species: Melampodium ameri-
canum L.
1. Melampodium americanum L. [Hort. Cliff. 425. 1738.]
Sp. Pl. 921. 1753. TYPE: MEXICO: Veracruz, near Vera-
cruz, Mar 1731, W. Houstoun s.n. (Holotype, BM; photo-
graph of holotype, Ny! US!; photograph of BM isotype, US!).
Calendula decumbens Mill. Gard. Dict. ed. 8. n. 9. 1768.
nom. superfl., based on type of Melampodium americanum.
Melampolium heterophyllum Lag. Gen. et Sp. Nov. 33.
1816. TYPE: “NOVA HISPANIA": grown in Roy. Bot. Gard.
Madrid, source and collector of seeds uncertain (Holotype,
MA?; tracing of DC isotype [?], GH!).
Melampodium sericeum H.B.K. Nov. Gen. Sp. 4: 272,
t. 398. 1820. non Lag. 1816. nom. illegit. TYPE: MEXICO:
Guerrero, between Tehuilotepec and Taxco, 5520 ft, Apr
1803, A. J. Bonpland 3968 (Holotype, P; isotype, P!; frag-
ment of holotype, P! US!; photograph of P isotype, OS! TEX!;
photograph of probable B isotype, TEX!).
Melampodium angustifolium DC. Prodr. 5:519. 1836.
TYPE: PERU [?]: “in Peruviae montanis, ad Oronocum,"
1790-91, T. Haenke s.n. (Holotype, PR; isotypes, P [2]!;
probable isotype, F!; photograph of G-DC isotype, F! NY!
us! Ipc! 800. 927: IIL1!; photograph of P isotype, os!
TEX!). If the collection actually is from Peru, then the
range of M. americanum would be extended much further
south than its present southern limit in Guatemala. How-
ever, apparent mixing of label data of Haenke's specimens
has been reported in at least three instances by other
"The method of citing specimens on Interdocumentation (IDC)
microfiches described by Hepper (1968) is used here throughout the
text. Taking the above citation as an example, the first number
(800) refers to the specific herbarium in the IDC collection (here
G-DC) followed by the numbers of: the individual microfiche (927),
the particular line on the card (III), and the specific frame (—speci-
men) on the line (1).
22 Rhodora [Vol. 74
workers (Hitchcock, 1909; Merrill, 1925; Tryon, 1955).
Because Haenke also collected in Mexico in the region from
Acapulco to Mexico City (Safford, 1905; Alston, 1934)
where M. americanum is common, it seems likely that mix-
ing of label data also has occurred here.
Melampodium kunthianum DC. Prodr. 5:519. 1836. nom.
nov. Based on type of M. sericeum H.B.K.
Meiampodium nelsonii Greenm. Proc. Amer. Acad. Arts
& Sci. 41:260. 1905. TYPE: MEXICO: Michoacán, Volcano
of Jorullo, 28 Mar 1903, E. W. Nelson 6939 (Lectotype
chosen, GH!; isotypes, NY! US!; photograph of US isotype,
TEX !).
Perennial subshrubs, 15-60 cm tall. Stems ascending, 1-2
mm diam, strigose to hispid-pilose with hairs 0.2-1 mm long.
Peduncles 2.5-7.5 em long. Leaves sessile, linear to ovate,
9-6 em long, 0.2-2.7 cm wide, at apex acute-obtuse, at base
attenuate to obtuse, with upper surface strigose with hairs
0.2-0.5 mm long, with lower surface sericeous; margin
entire or lobed to pinnately divided into linear segments,
at base usually pilose-hirsute with hairs up to 1 mm long.
Heads 7-8 mm tall, 11-17 mm diam. Outer involucre cupu-
late, 7-10 mm diam; bracts 5, slightly connate at base, im-
bricate 2/3 their length (rarely separate), ovate to rhombic,
5-7 mm long, 3-4 mm wide, at apex acuminate, with abaxial
surface strigose with hairs 0.5 mm long near apex, near
base pilose to sericeous, at margin herbaceous. Fruits 2-3
mm long, with lateral surfaces smooth with ribs and few
tubercles to verrucate-aculeate; hood apex mucronate (rare-
ly muticous) to cirrhous, with tapering appendage up to
2 mm long. Ray florets 8-13; ligules yellow-orange, oblong-
elliptic, 4-6 mm long, 1.5-3 mm wide. Disc florets 50-100;
corollas yellow-orange, 2.1 mm diam, with throat and tube
each 1 mm long. Paleae oblanceolate, 4 mm long, 1 mm
wide; apex yellow, with margin entire to undulate becom-
ing dentate-erose laterally; midrib prominent, pubescent
with hairs 0.4 mm long. Chromosome number, n = 10.
1972] Melampodium — Stuessy 23
Fig. 11. Map of Mexico and adjacent Guatemala showing distri-
bution of Melampodium americanum (dots), M. glabribracteatum
(square), and M. pringlei (triangle). Three collections of M.
americanum near Tepic, Nayarit, not shown.
Habitats ranging from savannas to pine-oak forests in
Mexico on the eastern slope of the Sierra Madre Oriental
and on the western slopes of the Sierra Madres Occidental
and Sur, with extensions into Guatemala (Fig. 11), 210-
2330 m. Flowering dates, Jan-Aug.
This species is very widespread, ranging from north in
Tamaulipas, Mexico, to south into Guatemala and is found
in diverse habitats from mountain pine forests to lowland
savannas. As might be expected from such geographical
24 Rhodora [Vol. 74
and ecological diversity, much morphological variability
also is present. Although no formal varieties are proposed
at this time, morpho-geographic trends do exist that need
further study: (1) plants with short leaves with few and
broad lobes and large heads found in the coastal lowlands
near Alvarado, Veracruz; (2) plants with long leaves, pin-
nately divided into narrow lobes, found in the mountains
of Chiapas and Guatemala; (3) short plants with small
heads and a tendency toward the annual habit found in the
mountains of the western Sierra Madre in Jalisco, Micho-
acan and Guerrero.
Three collections (Feddema 832; King 3699; Rose, Stand-
ley and Russell 14307) found in the northwestern range of
this species (north of Tepic, Nayarit) also may deserve
formal taxonomic status. The outer involucral bracts of
these collections are more rhombic with narrower atten-
uate apexes as compared with the more typical ovate
bracts of the rest of the species, and the ligules are longer
and narrower than usual. A collecticn from Guerrero, how-
ever, Hinton 9134, approaches these Nayarit collections and
seems to vitiate clear varietal or specific recognition. Ob-
viously additional work, especially in the Tepic area, is
needed.
9
REPRESENTATIVE SPECIMENS: Guatemala. BAJA VERAPAZ: ca 3 mi
S of Salamá, 10 Jul 1960, King 3260 (DS, NY, TEX, UC, Us); ca
14 mi W of Salama, 14 Jul 1960, King 3358 (DS, NY, TEX, UC, US);
ca 15 mi S of Rabinal, 14 Jul 1960, King 3364 (DS, NY, TEX, UC, US).
EL PROGRESO: 35 mi NE of Guatemala, 30 Jul 1966, Stuessy 602
(TEX). HUEHUETENANGO: Uaxackanal, 1 Aug 1896, Volkem 2973
(F [photograph of US specimen], GH, Ny, US). Mexico. CHIAPAS:
ca 10 mi E of the Oaxaca-Chiapas border on rte 190, 23 Jun 1960,
King 2980 (DS, NY, TEX, UC, US); ca 12 mi E of Cintalapa, 23 Jun
1960, King 2987 (DS, NY, TEX, UC, US); ca 33 mi S of Tuxtla Gutiérrez,
27 Jun 1960, King 3105 (Ds, NY, TEX, UC, US); 30 mi SE of Comitán,
26 Jul 1966, Stuessy 571 (TEX); 1 mi N of Entronque Santa Isabel,
10 Aug 1966, Stuessy 632 (TEX). COLIMA: Colima, 24 Oct 1910,
Orcutt 4516 (ps, F, GH); Colima, 9 Jan-6 Feb 1891, Palmer 1172 (G,
GH, NY, UC, US); 5 mi NW of rte 110 on rd to Alzada, 25 Aug 1966,
Stuessy 724 (TEX); Alzada, 25 Aug 1966, Stuessy 727 (TEX); Man-
zanillo, 1863-64, Xantus s.n. (F, GH [2], Ny [2], vs [2]). GUERRERO:
1972] Melampodium — Stuessy 25
Placeres, Puerta, 22 Jul 1936, Hinton 9134 (ARIZ, TEX, US); ca 9 mi
NW of Taxco, 14 Mar 1961, King 4168 (F, NY, TEX, UC, US); ca 25 mi
NE of Acapulco, 14 Mar 1961, King 4178 (F, NY, TEX, UC, US). JALISCO:
Río Cuvianes, 13 Jun 1892, Jones 274 (pom [2], US); Río Cuvianes,
13 Jun 1892, Jones 369 (POM, US); % km S of Puente San Pedro on
rd from Colima to Ciudad Guzmán, 31 Jul 1960, Koeppen & Iltis 618
(TEX, UC); 10 mi S of Autlán toward La Resolana, 28 Jun 1949,
R. & C. Wilbur 1895 (us). MÉXICO: 8 km SW of Luvianos, 2 Sep
1965, Rzedowski 20748 (08). MICHOACÁN: Coalcomán, 5 Jan 1939,
Hinton et al. 12871 (Ny, US); Uruapán, Tancitaro, 26 Oct 1940,
Hinton et al. 15587 (ARIZ, GH, TEX, US); ca 32 km N of Playa Azul
(near Los Encinos), 25-31 Oct 1961, King & Soderstrom 4992 (NY,
SMU, TEX, UC, US); Apatzingán, canyon below Acahuato, 15 Aug
1941, Leavenworth & Hoogstraal 1589 (F, GH); 7 mi S of Ario de
Rosales, 20 Aug 1966, Stuessy 688 (TEX). NAYARIT: Mirador del
Águila, ca 14 mi N of Tepic, 21 Aug 1959, Feddema 832 (MICH [2]);
ca 25 mi N of Tepic, 12 Aug 1960, King 3699 (DS, NY, TEX, UC, US);
vicinity of Acaponeta, Tepic, 10 Apr 1910, Rose, Standley & Russell
14307 (vs); Ixtlan del Río — San Marcos, 5 Aug 1921, Thompson
s.n. (NY). OAXACA: Estación San Marcos, Istmo de Tehuantepec,
21 Jan 1907, Conzatti 1106 (F, GH, US) ; vicinity of Yalalag [Hidalgo
Yalalag], Jul 1894, Nelson 947 (US); rd between Llano Grande &
Pinotepa, 19 Feb 1895, Nelson 2339 (US); between Guichocovi &
Lagunas, 27 Jun 1895, Nelson 2740 (GH, US). SAN LUIS POTOSÍ: San
Dieguito, 13-16 Jun 1904, Palmer 459 (US). TAMAULIPAS: 10 km
NW of El Progreso, 22 Aug 1941, Stanford, Retherford & Northeraft
1077 (ARIZ, DS, GH, NY, UC). VERACRUZ: La Purga, 27 Jan 1906,
Greenman 279 (F, GH, NY); ca 26 mi E of Cuitláhuac, 6 Jun 1960,
King 2682 (DS, NY, TEX, UC, US); ca 14 mi SE of Alvarado, 7 Jun
1960, King 2718 (Ds, NY, TEX, UC, US); vicinity of Pueblo Viejo,
2 km S of Tampico, 1-2 Jun 1910, Palmer 535 (GH, NY, US); 63 mi
S of jet rtes 110 & 105, 6 Jul 1966, Stuessy 471 (TEX).
2. Melampodium diffusum Cass. Dict. Sci. Nat. 59 :238.
1829. TYPE: PHILIPPINE ISLANDS: Manila, summer 1825,
F. L. Busseuil s.n. (Holotype, P?).
Melampodium manillense Less. Linnaea 6:155. t. 2, f.G
(fruit). 1831. TYPE: PHILIPPINE ISLANDS: “in Luconia"
[Luzon], 1831, A. Chamisso s.n. (Holotype, B?; photograph
of G-DC isotype (?), Ipc 800. 927:I. 6!).
Melampodium lanceolatum DC. Prodr. 5:519. 1836.
TYPE: PHILIPPINE ISLANDS or MEXICO: locality and date
unknown, L. Née s.n. (Holotype, G-DC; isotype, MA!; photo-
26 Rhodora [Vol. 74
graph of holotype, Us!, pc 800. 927:1I. 6!; photograph of
MA isotype, OS! TEX!).
Melampodium diffusum Cass. var. lancelatum (DC.)
Robins. Proc. Amer. Acad. Arts & Sci. 36:460. 1901.
Annual herbs, 25-40 cm tall. Stems erect, 1-5 mm diam,
subglabrous at base to strigillose-hispidulous above with
hairs up to 0.8 mm long. Peduncles 0.8-6.5 em long. Leaves
usually sessile (sometimes with petioles up to 3 mm long),
lanceolate to narrowly ovate, 2.5-5.5 cm long, 0.5-1.5 (—3)
cm wide, at apex acute, at base attenuate to somewhat
obtuse, with upper surface strigose with hairs 0.3 mm long,
with lower surface sericeous; margin entire to 1-3 cleft,
hispid with hairs up to 1.5 mm long. Heads 5-7 mm tall,
6-11 mm diam. Outer involucre cupulate, 5-9 mm diam;
bracts 5, slightly connate at base, imbricate 1/2 their length,
ovate, 2.5-6 mm long, 1.5-4 mm wide, at apex acuminate,
with abaxial surface strigillose with hairs 0.3 mm long, at
margin herbaceous. Fruits 1.8-2.3 mm long, with lateral
surfaces smooth and longitudinally ribbed to tuberculate-
aculeate; hood apex mucronate to cirrhous (rarely muti-
cous)^, with appendage up to 2 mm long. Ray florets 8-13;
ligules yellow-orange, oblong-elliptic, 2.5-4 mm long, 1-3
mm wide. Disc florets 40-60; corollas yellow-orange, 1.3 mm
diam, with throat 0.6 and tube 0.4 mm long. Paleae ob-
lanceolate, 3 mm long, 0.9 mm wide; apex yellow, with
margin entire to moderately erose; midrib prominent, most-
ly glabrous. Chromosome number, n — 10.
Tropical deciduous forests in Colima and near Acapulco,
Guerrero, Mexico, and Manila in the Philippine Islands
(Fig. 12), 30-300 m. Flowering dates, Jun-Jan.
One collection, Winbery & Rowell 2492, approaches the
perennial habit in this characteristically annual species. Be-
cause this collection also possesses heads and leaves larger
than usual, the suggestion of hybridization with nearby M.
americanum, especially at higher altitudes in Guerrero, can-
not be excluded.
*One collection, Palmer 3 (F), has fruits with hoods and no hoods
in one head of the single plant.
1972] Melampodium — Stuessy 27
Fig. 12. Map of Mexico, adjacent Guatemala, and the Philippine
Islands (inset), showing distribution of Melampodium diffusum
(triangles), M. longifoliwm (closed squares), M. longipilum (dots),
M. mimulifolium (open square), and M. pilosum (circles).
Robinson (1901, p. 456) previously noticed the unusual
bicentric distribution of this species and offered the follow-
ing plausible explanation:
*As the genus is otherwise American, the occurrence of
this species in a region so remote has always been prob-
lematie, and it has been a matter of no small interest to
find the Philippine plant closely matched by specimens
recently collected by Dr. Edward Palmer, about Acapulco,
Mexico. There can therefore be scarcely a doubt that the
genus is in reality of New World origin, and that a single
Mexican species was accidentally introduced into the
Philippines, where it attracted scientific attention before
it was recognized in America. This seems the more likely
from the circumstance that Mexico and the Philippines
were under the same national control, and early con-
28 Rhodora [Vol. 74
nected by a certain amount of oceanic traffic [demon-
strated by Merrill (1954)]. This being the case, the
transference of seed from Acapulco, the most important
Pacific port of Mexico, to the neighborhood of Manila,
presents no inherent improbability."
REPRESENTATIVE SPECIMENS. Mexico. COLIMA: 5-10 mi N of Teco-
man, 25 Dec 1958, Thompson & Fields 332( TEX). GUERRERO: above
Playa Hornos ca 1 mi E of Acapulco, 9 Jan 1944, Barkley 14142
(F, GH, POM, TEX, UC, US); near Pie de la Cuesta NW of Acapulco,
5 Jan 1944, Barkley 14163 (TEx, us); 20 mi NE of Acapulco, 20 Aug
1947, Barkley, Webster & Paxson 17M724 (TEX); between Acapulco
& Pueblo Nuevo, 13 Nov 1882, Hancock 25 (F, K); 8 mi E of Acapulco,
22 Jun 1952, Mockford & Rowell 2769 (SMU); near Acapulco, 25-31
Oct 1895, Palmer 3 (F, GH, Ny, UC, US); near Acapulco, Dec 1895,
Palmer 281 (GH, US); 26 mi W of Acapulco, 19 Aug 1961, Powell &
Edmondson 758 (F, TEX); 5 mi SW of Tierra Colorada, 17 Jun 1952,
Winbery & Rowell 2492 (sMU); 1 mi N of Acapulco, 27 Aug 1965,
Stuessy 366 (TEX); near S shore of Acapulco bay, 28 Aug 1965,
Stuessy 369 (TEX). Philippine Islands. LUZON: Cavite, Aug 1905,
Foxworthy 167 (NY, US); Cavite, Binacayan, 21 Jan 1917, Merrill
10641 (NY, US); Cavite, Bacoor, Jul 1910, Robinson 11835 (F);
Laguna, 17 Apr 1953, Canicosa 1103 (US); Nueva Vizcaya, Bam-
bang, Jan 1924, Clemens s.n. (UC); Nueva Vizcaya, Benquet, Dec
1908, Curran & Merritt 15837 (US); Nueva Vizcaya, Jan 1913,
MeGregor 20181 (vs).
9. Melampodium pilosum Stuessy, Brittonia 22:115. f. 6.
1970. TYPE: MEXICO: Guerrero, Montes de Oca, Vallecitos,
28 Sep 1937, G. B. Hinton 11428 (Holotype, NY!; isotypes,
G! GH! uc! us!).
Annual herbs, 8-28 cm tall. Stems erect to decumbent,
1-2 mm diam, pilose with hairs up to 1.5 mm long. Pedun-
cles 3-3.7 cm long. Leaves with petioles 2 mm long, lance-
olate to narrowly ovate, 3-6 cm long, 1-1.3 cm wide, at apex
acute, at base obtuse, with upper surface near apex strigose
with hairs 0.3 mm long, near base pilose with hairs 1.5 mm
long, with undersurface markedly sericeous; margin en-
tire to rarely 1-2 lobed, irregularly ciliate with hairs 0.3
mm long, toward base pilose with hairs up to 2 mm long.
Heads 6-7 mm tall, 7.5-11 mm diam. Outer involucre cupu-
late, 6.5-9 mm diam; bracts 5, slightly connate at base, im-
1972] Melampodium — Stuessy 29
bricate 1/2 their length, ovate to narrowly ovate, 4-5 mm
long, 2-2.5 ram wide, at apex acuminate, with abaxial sur-
face pilose with hairs 1 mm long, at margin herbaceous.
Fruits 1.6-2.5 mm long, with lateral surfaces 3-ribbed and
smooth to very weakly tuberculate; hood apex mucronate
to cirrhous?, with tapering appendage up to 2 mm long.
Ray florets 7-8; ligules yellow-orange, ovate-oblong, 3-4.5
mm long, 2-3 mm wide. Disc florets 55-80; corollas yellow,
1.2 mm diam, with throat and tube each 0.7 mm long. Paleae
oblanceolate, 4 mm long, 1 mm wide; apex yellow, with
margin erose becoming laciniate laterally; midrib promi-
nent, hirsutulous with hairs 0.3 mm long. Chromosome
number, » — 10.
Tropical deciduous forests in Michoacán and Guerrero,
Mexico (Fig. 12), 910-1220 m. Flowering dates, Aug-Sep.
REPRESENTATIVE SPECIMENS. Mexico. GUERRERO: Coyuca, Chaca-
merito, 14 Aug 1934, Hinton 6436 (F, G, GH, NY [2], US). MICHOACÁN:
ca 1 mi N of Huacana, 11 Aug 1965, Melchert & Sorensen 6085
(ENCB); 25 mi S of Ario de Rosales, 20 Aug 1966, Stuessy 695
(TEX).
4. Melampodium longipes (A. Gray) Robins. Proc. Amer.
Acad. Arts & Sci. 36:459. 1901.
Melampodium sericeum Lag. var. longipes A. Gray.
Proc. Amer. Acad. Arts & Sci. 22:423. 1887. TYPE:
MEXICO: Jalisco, Tequila, “base of mountains," Aug 1886,
E. Palmer 391 (Holotype, GH ! ; isotypes, G! K! Mo! Ny 12]!
US!; photograph of Us isotype, TEX!).
Annual herbs, 10-46 cm tall. Stems erect, 1-2 mm diam,
hispid-pilose with hairs up to 1 mm long. Peduncles 5-8
cm long. Leaves sessile, linear and entire to markedly pin-
nately parted, 3-7 cm long, 0.3-3.5 cm wide, at apex acute-
obtuse, at base mostly attenuate (less often obtuse), with
upper surface strigose with hairs 0.3-1 mm long, with lower
surface sericeous; margin entire, at base hispid-pilose with
hairs up to 2 mm long. Heads 6.5-8.5 mm tall, 10-12 mm
*The holotype has fruits possessing hoods with both mucronate and
cirrhous apexes.
30 Rhodora [Vol. 74
Fig. 18. Map of central Mexico showing distribution of Melam-
podium dicoelocarpum (triangles), M. glabrum (dots), M. longipes
(squares), and M. nutans (circles). One collection of M. nutans from
Oaxaca not shown.
diam. Outer involucre cupulate, 7-8 mm diam; bracts 5,
slightly connate at base, imbricate 2/3 their length, obovate,
4-5 mm long, 2.5-3 mm wide, at apex acuminate, with
abaxial surface pilose with hairs 1 mm long (more dense
near base) ; margin herbaceous. Fruits 2-3 mm long, with
lateral surfaces tuberculate with 3-4 longitudinal ribs; hood
apex mucronate to cirrhous, with tapering appendage up
to 3 mm long. Ray florets 8-10; ligules yellow-orange, ovate-
oblong, 3-5 mm long, 1-3 mm wide. Disc florets 50-80;
corollas yellow-orange, 2 mm diam, with throat 0.8 mm and
tube 0.7 mm long. Paleae oblanceolate, 3.5 mm long, 0.8
mm wide; apex yellow, with margin entire to undulate;
midrib distinct, mostly glabrous. Chromosome number,
n = 10.
1972] Melampodium — Stuessy 31
Dry hillsides in Jalisco, Nayarit and Sinaloa, Mexico
(Fig. 13), ca 1220 m. Flowering dates, Aug-Sep.
This species is very similar to both M. americanum and
M. linearilobum. M. longipes usually can be distinguished
from either of these two taxa by its annual habit (peren-
nial in M. americanum) and herbaceous outer involucral
bracts (conspicuously scarious in M. linearilobum). The
leaves divided markedly into pinnatifid segments and the
yellowish hue of the stems also serve as additional features
to help recognize M. longipes from among its close relatives.
One collection, Rose 3183, from Sinaloa tends more to-
ward a perennial habit, and the leaves are more distinctly
pinnatifid. Nevertheless, these specimens appear to fall
within the limits of M. longipes.
REPRESENTATIVE SPECIMENS. Mexico. JALISCO: ca 2 mi NW of
Tequila, 8 Aug 1960, King 3662 (DS, MICH, NY, TEX, UC, US); Tequila,
22 Sep 1893, Pringle 4598 (F, G [2], GH, MICH, MSC, NY, P, UC, US);
2 mi NW of Tequila, 3 Sep 1965, Stuessy 397 (TEX), 26 Aug 1966,
Stuessy 738 (TEX). NAYARIT: El Maguey, Sierra Madre, Tepic,
26 Aug 1905, Goldsmith 132 (Ds, F, GH, NY, UC, US). SINALOA: be-
tween Rosario & Colomas, 13 Jul 1897, Rose 3183 (GH, US).
5. Melampodium linearilobum DC. Prodr. 5:518. 1836.
TYPE: MEXICO: “ad Guilotepec et Las Bacas," Sep 1831,
L. Alamán s.n. (Holotype, G-DC; isotype, G!; photograph of
holotype, inc 800. 927: II. 2!; photograph of G isotype,
F! os! TEX! us!).
Melampolium canescens Brandg. Zoe 5:222. 1905. TYPE:
MEXICO: Sinaloa, Cofradia [ca 30 mi E of Culiacán near
Durango border], 22 Oct 1904, T. S. Brandegee s.n. (Lecto-
type chosen, UC!; isotypes, GH! POM! US!).
Annual herbs, 6-50 cm tall. Stems erect, 1-3 mm diam,
strigillose-strigose with hairs 0.8 mm long. Peduncles 3.5-
5.5 em long. Leaves sessile, linear and entire to markedly
pinnately parted, 3-5.5 cm long, 0.3-3 em wide, at apex
acute-obtuse, at base attenuate to obtuse, with upper sur-
face strigose with hairs 0.2-1 mm long, with lower surface
sericeous; margin entire, at base hispid-pilose with hairs
up to 2 mm long. Heads 5-7 mm tall, 7-11 mm diam. Outer
32 Rhodora [Vol. 74
involucre cupulate, 6-10 mm diam; bracts 5, slightly con-
nate at base, imbricate 1/2 their length, narrowly ovate to
somewhat obovate, 3.5-5.5 mm long, 2.5-3 mm wide, at apex
acute, with abaxial surface pilose-strigose with hairs 1-1.8
mm long; margin conspicuously scarious. Fruits 2-3 mm
long, with lateral surfaces smooth and striate to tubercu-
late; hood apex muticous to cirrhous, with appendage up
to 2.5 mm long. Ray florets 7-8; ligules yellow-orange,
ovate-orbicular (often with imbricate margins), 2-4 mm
long, 2-5 mm wide. Disc florets 45-75; corollas yellow-
orange, 2.4 mm diam, with throat 0.9 mm and tube 0.5 mm
long. Paleae oblanceolate, 4 mm long, 1 mm wide; apex
yellow, with margin entire (rarely erose) ; midrib promi-
nent, mostly glabrous. Chromosome number, » — 10.
Tropical deciduous and pine forests of the western Sierra
Madres of Mexico and into Central America to Costa Rica
(Fig. 14), 20-1620 m. Flowering dates, Jul-Oct.
The conspicuous scarious margins (colored orange-yel-
low when fresh) of the outer involucral bracts of M. linear-
ilobum. The type of M. canescens from Sinaloa has linear,
members of series Melampodium. Were it not for this char-
acteristic bract feature, M. linearilobum would be very dif-
fieult to distinguish from M. longipes.
In spite of considerable vegetative variability, no intra-
specific categories have been recognized within M. linear-
ilobum. The type of M. canescens from Sinaloa has linear,
entire leaves, as do other collections from nearby areas, but
plants with similar leaves also appear in Central America
and in scattered localities in Mexico. As a further indica-
tion of the plasticity of leaf shape, in many plants the upper
leaves are linear and entire, while those lower on the stem
are once or twice pinnatifid. Due to this vegetative plastic-
ity, M. canescens is considered herein to be synonymous
with M. linearilobwm.
REPRESENTATIVE SPECIMENS. Costa Rica. GUANACASTE: Bolson
River, 16 Jul 1918, W. & H. Rowlee 179 (Ny, US). El Salvador.
AHUACHAPÁN: 1921, Padilla 1 (US), 1921, Padilla 242 (us), 1923,
Q2
e»
1972] Melampodium — Stuessy
Fig. 14. Map of Mexico and Central America showing distribu-
tion of Melampodium linearilobum.
Padilla 586 (US). MORAZAN: ca 8 km SW of Montecristo, 3 Dec
1941, Tucker 455 (F, GH, NY, SMU, UC, US). SANTA ANA: near
Chalchuapa, 1922, Calderén 979 (GH, NY, US). USULUTAN: 24 mi
E of turnoff to San Vicente on rte 1, 3 Aug 1966, Stwessy 613 (TEX).
Guatemala. CHIQUIMULA: near Ipala, 23 Oct 1939, Steyermark 30309
(F). HUEHUETENANGO: between San Ildefonso Ixtahuacán & Cuilco,
16 Aug 1942, Steyermark 50724 (F, NY). JUTIAPA: between Suchitán
& Sta Catarina, Jul 1870, Bernoulli 709 (NY); near Jutiapa, 20 Dec
1938, Standley 60493 (F); between Jutiapa & Plan de Urrutia, 28
Oct 1940, Standley 75620 (F, US); 8 mi NE of Jutiapa, 2 Aug 1966,
Stuessy 607 (TEX). QUICHE: 1942, Ignacio 1495 (F). ZACAPA: near
divide on rd between Zacapa & Chiquimula, 9 Oct 1940, Standley
73774 (F); between Río Hondo & Santa Cruz, 11 Oct 1940, Standley
74071 (F); between Río Hondo & Santa Cruz, 11 Oct 1940, Standley
74125 (F, US); near La Fragua, 14 Oct 1940, Standley 74813 (F);
near Sta Rosalia, 4 Oct 1939, Steyermark 29041 (F). Honduras.
CHOLUTECA: vicinity of Pespire, 18-27 Oct 1950, Standley 27175
(F). FRANCISCO MORAZÁN: El Chile, Tegucigalpa, 18 Sep 1950,
34 Rhodora [Vol. 74
Standley 26708 (F). OLANCHO: near Rio Telica, 20 Nov 1963,
Molina 13346 (F). VALLE: 21 Jan 1956, Molina 5899 (F). Mexico.
CHIAPAS: Chicomucelo, 14 Jul 1941, Matuda 4444 (4504) (F);
valley of Jiquipilas, 16-18 Aug 1895, Nelson 2949 (GH, US). DURANGO?:
Lodiego [25°N, 106*45'W; McVaugh, 1956], 9-15 Oct 1891, Palmer
1609 in part (vs [2]). GUERRERO: Coyuca de Catalán, Pungarabato,
10 Jul 1934, Hinton et al. 6270 (F, GH, NY, US); Mina, Placeres,
18 Aug 1936, Hinton et al. 9118 (ARIZ, GH, TEX, US); near Iguala,
26 Apr 1900, Pringle 9162 (F, GH, Ny, US); Los Amates Station,
28 Sep 1905, Pringle 10065 (ARIZ, F, G, NY, SMU, UC, US); ca 50 mi
S of Cuernavaca, 26 Aug 1965, Stuessy 361 (TEX). MÉXICO: Tema-
scaltepec, Limones, 7 Nov 1932, Hinton 2513 (GH, NY [2]); Tema-
scaltepec, Palmar, 9 Aug 1934, Hinton et al. 6420 (Ny, US); San
Antonio, Tlatlaya, 20 Jul 1954, Matuda et al. 31141 (vs); Palmar
Chico, Amatepec, 24-25 Aug 1954, Matuda et al. 31314 (Us).
MICHOACAN: Zitacuaro, Zitacuaro-Tiamaro, 9 Sep 1938, Hinton et al.
13214 (ARIZ, GH, TEX, US); Tancitaro, above Apatzingan, 15 Aug
1941, Leavenworth & Hoogstraal 1480 (F, GH, NY); Apatzingán,
below Acahuato, 15 Aug 1941, Leavenworth & Hoogstraal 1588
(F); 1 mi N of Apatzingán, 21 Aug 1966, Stuessy 697 (TEX); 4 mi
N of Nueva Italia, 21 Aug 1966, Stuessy 698 (OS, TEX). OAXACA:
Salina Cruz, 23 Aug 1935, Fisher 35321 (ARIZ, F, NY, SMU, US);
ca 37 mi W of Tehuantepec, 23 Jul 1960, King 3454 (DS, Ny, TEX,
UC, US) ; ca 3 mi NE of Huajuapán de León, 28 Jul 1960, King 5539
(TEX); Playa de Puerto Ángel, Oct 1917, Reko 3812 (vs); 24 mi
S of Sola de Vega, 18 Aug 1966, Stuessy 650 (TEX). SINALOA: Cerro
Colorado, near Cofradia, 2 Nov 1904, Brandegee s.n. (GH, UC);
Cofradia, 25 Nov 1939, Gentry 5039 (ARIZ, GH, NY); Las Mesas,
Sierra Surotato, 25 Aug 1941, Gentry 6153 (ARIZ, GH, NY); La Noria,
10 Oct 1925, Mexia 240 (uc), 14 Oct 1925, Mexia 349 (POM, UC, US).
Nicaragua. GRANADA: 1 mi S of Granada, 6 Aug 1966, Stuessy 619
(TEX).
2. Series Leucantha Stuessy, ser. nov.
Plantae perennes suffruticosae; folia anguste ovata vel
lineata, saepe lobata, subtus strigosa; ligulae eburneae, plus
quam vel raro circa 2 mm longae; involucrum extimum
marginibus herbaceis; fructi cucullati. Species 6-8. Typus:
Melampodium leucanthum Torr. & A. Gray.
6. Melampodium leucanthum Torr. & A. Gray, Fl. N. Amer.
2(2) :271. 1842. TYPE: TEXAS: without locality or date,
J. L. Riddell s.n. (Holotype, Nv?). Type not located, but the
following collection cited by Gray (1852) clearly is repre-
1972] Melampodium — Stuessy 35
sentative: “Hills near El Paso," Apr 1849, C. Wright 311
(GH! UC! UST):
Perennial subshrubs, 15-60 cm tall. Stems ascending, 1.1-
2.5 mm diam, strigillose (rarely hispid) with hairs 0.1-0.2
mm long. Peduncles 3-7 cm long. Leaves sessile, linear-
oblong, 2.1-4.5 em long, 0.13-1 cm wide, at apex and base
obtuse, with both surfaces strigillose with hairs 1-2 mm
long; margin entire to pinnately 6-lobed, at base strigillose
with hairs 0.1-0.2 mm long. Heads 6-8 mm tall, 20-37 mm
diam. Outer involucre cupulate, 10-13 mm diam; bracts 5,
connate 1/2 to 3/4 their length, ovate, 5-7 mm long, 4-5.1
mm wide, at apex acute, with abaxial surface strigillose-
strigose with hairs 0.1-0.7 mm long; margin herbaceous.
Fruits 1.5-2.6 mm long, with lateral surfaces aculeate-ver-
rucate; hood apex muticous (very rarely mucronate). Ray
florets 8-13; ligules cream-white, oblong-elliptic, 7-13 mm
long, 2.5-8 mm wide. Disc florets 25-50; corollas yellow,
1 mm diam, with throat 1.2 mm and tube 0.8 mm long.
Paleae oblong-elliptic, 2.9 mm long, 1.1 mm wide; apex
yellow, with margin laciniate; midrib prominent, often
strigillose with hairs 0.1 mm long. Chromosome numbers,
» — 10 and 20.
Caleareous soils throughout western and central Texas
(stopping at the Edward's Plateau), Arizona, New Mexico,
and portions of Oklahoma, Kansas, Colorado, and northern
Mexico (Fig. 15), 490-2590 m. Flowering dates, Apr-Sep.
This species is the familiar ‘‘blackfoot daisy” of the
southwestern United States that has been gathered exten-
sively by many different collectors. Despite its abundance,
however, the relationships to M. cinereum and M. argophyl-
lum have been unclear for many years. Recent studies have
shown (Stuessy, 1971c; present treatment) that M. leucan-
thum has a separate distribution that comes only within
ten miles of M. cinereum at the edge of the Edward's
Plateau.
Both 2n and 4n cytological races have been reported for
this species (Turner and King, 1962; Stuessy, 1971b & c),
36 Rhodora [Vol. 74
Fig. 15. Map of the southwestern United States and adjacent
Mexico showing distribution of Melampodium lewcanthum.
and they are morphologically indistinguishable. Evidence
for treating these cytotypes as informal races having been
derived by spontaneous autopolyploidy has been reviewed
by Stuessy (1971c).
REPRESENTATIVE SPECIMENS. Mexico. CHIHUAHUA: 7 mi N of
Mestenas, 25 Sep 1938, Johnston 7959 (GH, US); Organos Mts, 8 Sep
1937, LeSueur 1456 (F, TEX); Colonia Diaz, 20-21 Sep 1899, Nelson
6447 (GH, US); hills near Chihuahua, 5 Oct 1886, Pringle 1019 (F,
NY, US); 20 mi S of Ciudad Jüarez, 21 Jul 1937, Shreve 7918
(ARIZ, GH, US); 59 mi N of Villa Ahumada, 23 Aug 1967, Stuessy
1122 (TEX). COAHUILA: Del Carmen Mts, 6 Sep 1936, Marsh 855
(F, GH, TEX); Cañon del Indio Felipe, close to the Chihuahuan
boundary, 27-29 Sep 1940, Stewart 84 (GH). SONORA: Colonia
Morelos, 15 Sep-4 Oct 1941, White 4587 (ARIZ, GH); 5 mi E of
Esqueda on rd to Río de la Tierra, 10 May 1948, Wiggins 11765
1972] Melampodium — Stuessy 37
(Ds, US). UNITED STATES. Arizona: COCHISE CO., near Douglas,
16 Aug 1907, Goodding 2399a (DS, GH, US). GILA CO., ca 3 mi N of
Dripping Springs turnoff, 11 Apr 1965, Niles 572 (ARIZ, TEX);
GREENLEE CO., 7 mi N of Metcalf, 6 Jun 1935, Maguire, Richards &
Moeller 11849 (ARIZ, GH, NY); MARICOPA CO., 3 mi N of Sunflower,
9 Apr 1960, Russell 11511 (smu [2], UC); MOHAVE co., Kingman,
18 Apr 1935, Kearney & Peebles 11133 (ARIZ, GH, US); PIMA CO.,
3 mi N of Greaterville, 17 Aug 1967, Turner 5735 (TEX); YAVAPAI CO.,
10 mi SW of Congress, 21 Apr 1962, Turner 4790 (DS, FSU, SMU,
TEX); YUMA CO. Squaw Canyon, Harquahala Mts, 9 Sep 1952,
Wright 42-54 (ARIZ). Colorado: BACA co. 25 mi S of Pritchett,
11 Jul 1947, Porter 4262 (GH, RSA, SMU, TEX); EL PASO CO., below
Colorado Springs, 29 May 1878, Jones 117 (F, POM [2]); FREMONT
co., Canon City, 2 Jul 1920, Clokey 3946 (DS, F, GH, NY, POM, UC, US) ;
PROWERS CO. 31 mi S of Lamar, 6 Jun 1967, Irving 825 (TEX);
PUEBLO CO., 12 mi S of Pueblo, 7 Jun 1922, Wiegand & Upton 4439
(NY). Kansas: FORD CO., 3 mi S of Dodge City, 9 Jul 1950, Horr
3444 (GH, US); HAMILTON CO., vicinity of Syracuse, 15 Sep 1912,
Rose & Fitch 17043 (NY, US); MEADE CO., 16 mi SE of Meade, 16
Aug 1939, Horr & Franklin E278 (F, GH, SMU, TEX, UC, US) ; SEWARD
CO., Jul 1891, Carleton 331 (ARIZ, F, US). New Mexico: BERNALILLO
CO. ca 2 mi E of Albuquerque, 1915, Kammerer 48 (DS, TEX);
CHAVES CO., 20 mi S of Roswell, Aug 1900, F. & E. Earle 295 (NY,
POM, US); DE BACA CO., 6 mi E of Yeso, 8 Aug 1967, Turner 5673
(TEX); DONA ANA CO., Organ Mts, 9 Jul 1897, Wooton 117 (ps, GH,
NY, POM, UC, US) ; GRANT CO., Mangas Springs, 18 mi NE of Silver
City, 15 May 1903, Metcalfe 66 (ARIZ, DS, NY, POM, UC [2], vs);
LINCOLN CO., 15 mi E of Capitan, 25 Jul 1938, Hitchcock, Rethke & van
Raadshooven 4244 (DS, GH, UC); ROOSEVELT CO., 5 mi NE of Portales,
14 Jul 1930, Goodman & Hitchcock 1130 (ps, GH, NY, UC) ; SAN MIGUEL
co., 10 mi SE of Las Vegas, 27 Jul 1924, Bacigalupi 607 (Ds, GH,
UC) ; SANTA FE CO., 20 mi S of Golden, 24 Jul 1938, Hitchcock, Rethke
& van Raadshooven 4214 (DS, GH, UC); SIERRA CO., Kingston, 29 Aug
1904, Metcalfe 1271 (F, GH, NY, POM, UC, US). Oklahoma: BEAVER
co., near Knowles, 5 May 1913, Stevens 325 (ps, NY, SMU, US);
CIMARRON CO., Kenton, low sides of Black Mesa, 28 Jul 1936, Demaree
13344 (GH, NY, POM, SMU); JACKSON CO. El Dorado, 18 Apr 1936,
Demaree 12210 (SMU, US). Texas: ANDREWS CO, 5 mi WSW of
Andrews, 15 Sep 1966, Shinners 31559 (SMU); BELL CO. 2% mi S
of Nolanville, 7 Mar 1954, C. & G. York 54037 (SMU, TEX); BEXAR
co., 5 mi NW of San Antonio, 4 May 1948, Burr 135 (ps, Ny [2]);
BLANCO CO. 10 mi N of Johnson City, 18 Apr 1958, Thompson &
Graham 18 (SMU, TEX); BREWSTER CO. Marathon, 15 Jul 1965,
Stuessy 230 (TEX); BROWN CO., 7 mi S of Brownwood, 24 Apr 1966,
Guthrie 43 (SMU, TEX); CALLAHAN CO., ca 2 mi W of Baird, 27 Mar
38 Rhodora Vol. 74
1968, Henderson 63-88 (FSU, SMU, TEX); COLEMAN CO., 1 mi SE of
Santa Ana, 9 Jul 1957, Shinners 26390 (SMU); COMAL CO., Comanche
Spring, Mar 1849, Lindheimer 949 (ARIZ, F, GH, NY, TEX, UC, US);
COMANCHE CO. 7 mi W of Comanche, 6 Apr 1966, Teeters 18 (LL,
SMU, TEX); CORYELL CO., 4 mi S of Gatesville, 2 Oct 1965, Baize 5
(LL, SMU, TEX); CROCKETT CO., 31 mi W of Ozona, 8 Jul 1965, Stuessy
147 (TEX); CULBERSON CO. 1 mi S of Texas-New Mexico line E of
Guadalupe Mts, 14 Aug 1942, Waterfall 3781 (ARIZ, GH, NY) ; DAWSON
co., plains between Lamesa & Tahoka, 29 Apr 1925, Small & Wherry
12134 (NY, TEX); ECTOR CO., 5.8 mi S of Odessa, 8 May 1966, Shinners
31236 (SMU); EL PASO CO., 28 mi SE of El Paso, 24 Aug 1967,
Stuessy 1127 (TEX); ERATH CO., Stephenville Pk, 9 mi N of Stephen-
ville, 2 Apr 1950, Shinners 12190 (SMU); FLOYD CO., intersec-
tion of hwys 97 & 2009, 22 Apr 1962, Melchert 180 (TEX); GIL-
LESPIE C0., 4.8 mi NE of Lawrence Jung Ranch, 31 Mar 1967,
Mears 1383 (TEX); HAMILTON CO. 17 mi S of Hico, 16 Apr 1945,
Shinners 7183 (GH, LL, SMU, UC); HARDEMAN CO., 11 mi S of Quanah,
4 Apr 1966, Turner 7 (LL, TEX); HAYS CO., 1.5 mi E of Dripping
Springs, 30 May 1966, Stuessy 414 (TEX); HEMPHILL Co., 18 mi SW
of Canadian, 15 Sep 1950, Tharp & Miller 51-312 (RSA, TEX);
HUDSPETH CO., ca 20 mi N of Allamore, 29 Jul 1943, Waterfall 4818
(GH, NY, SMU); JEFF DAVIS CO. gravel flat between Big & Little
Aguja Canyons, Davis Mts, 16 Jun 1931, Moore & Steyermark 3115
. (DS, GH, NY, UC, US); KENDALL CO. 5 mi W of Sisterdale, 24 May
1965, Flyr 489 (ns, SMU); KERR CO., Kerrville, 19-25 Apr 1894, Heller
1632 (ARIZ, F, GH, NY [2], POM, SMU, UC [2], US); KNOX CO., 5 mi
E of Benjamin, 22 Apr 1962, Melchert 169-C (TEX); LLANO CO.,
Marble, 31 Mar 1967, Mears 1413 (TEX); LOVING CO, 1 mi W of
Mentone, 10 Jul 1965, Stuessy 180 (TEX); LUBBOCK CO., Lubbock,
4 May 1930, Demaree 7594 (DS, GH, US); NOLAN CO., Sweetwater,
5 May 1927, Stanfield s.n. (NY, TEX); OCHILTREE CO., 7.8 mi SE of
Perryton, 13 Jul 1957, Wallis 4807 (ARIZ, SMU); OLDHAM CO., 16 mi
N of Vega, 17 May 1967, Turner 5632 (TEX); PECOS CO., 10 mi E
of Ft. Stockton, 19 Apr 1946, Warnock 46217 (FSU, TEX); POTTER
co., 1 mi N of Canadian River Bridge, 19 May 1945, Jespersen
2678 (DS, F, NY, SMU, UC, US); PRESIDIO CO., 7.5 mi N of Candelaria,
17 Apr 1947, MceVaugh 7986 (ps, F, LL [2], SMU, TEX [2]) ; RANDALL
co., Palo Duro State Pk, 20 Oct 1945, Cory 50422 (GH, NY, SMU);
SHACKELFORD CO., 2 mi NE of Jones Co. line, 27 Mar 1963, Henderson
63-98 (SMU, TEX); STEPHENS CO. 3 mi N of Ranger, 29 Apr 1939,
Culwell & Timmons 3091 (SMU, TEX); TAYLOR CO, 5 mi SW of
Abilene, 30 Jun 1962, Turner & Melchert 4840 (TEX); TRAVIS CO.,
7 mi SW of Zilker Pk, Austin, 13 Oct 1966, Stuessy 752 (0s, TEX) ;
UVALDE CO., Garner State Pk, 21 Jun 1958, Sullivan & Turner 34
(FSU, TEX); VAL VERDE CO, 13 mi W of Langtry, 23 Mar 1941,
1972] Melampodium — Stuessy 39
Innes & Warnock 589 (DS, GH, NY, TEX); WARD CO., Barstow, 15 Apr
1902, Tracy & Earle 23 (F, GH, NY [2], TEX, US), WILBARGER CO.,
16.9 mi S of Electra-Waggoner pastures, 12 May 1945, Whitehouse
9841 (SMU, TEX); WINKLER CO., 1 mi W of Kermit, 10 Jul 1965,
Stuessy 167 (TEX).
7. Melampodium cinereum DC. Prodr. 5:518. 1836.
Perennial subshrubs, 14-20 cm tall. Stems suffruticose,
ascending. Leaves sessile, at apex and base obtuse, at mar-
gin revolute when dry. Outer involucre cupulate; bracts
5, connate 1/6 to 1/3 their length, ovate, at apex acute, with
abaxial surface strigose with hairs 0.2-1 mm long, at mar-
gin herbaceous. Fruits with lateral surfaces aculeate-ver-
rucate. Ligules cream-white, oblong-elliptic. Disc corollas
yellow. Paleae oblong-oblanceolate; apex yellow, with mar-
gin laciniate; midrib prominent, glabrous or puberulous
with hairs 0.1 mm long.
Ta. Melampodium cinereum DC. var. cinereum
Melampodium cinereum DC. Prodr. 5:518. 1836. TYPE:
MEXICO[?]: San Fernando de las Presas, Oct 1830, J. L.
Berlandier 2243 [—823] (Holotype, G-DC; isotypes, G! GH!
K!; photograph of holotype, Ipc 800. 927: II. 4!; photo-
graph of K isotype, B! US!; photograph of G isotype, F! Us!).
San Fernando is a town about 83 miles SW of Matamoros,
Tamaulipas, Mexico, an area visited by Berlandier (Geiser,
1948). However, this variety is known to occur only near
Laredo and the surrounding territory in southcentral Texas,
indicating that the locality may have been interchanged
with M. ramosissimum as suggested by Gray (1884) and
Robinson (1901).
Stems 0.8-1.3 mm diam, strigose with hairs 0.1-0.8 mm
long. Peduncles 1.5-7 cm long. Leaves linear-oblong, 1.2-
5.0 em long, 0.12-1.4 cm wide, with both leaf surfaces
strigose with hairs 0.2-0.6 mm long; margin entire to pin-
nately 10-lobed or parted, near base strigose with hairs
up to 0.6 mm long. Heads 5-8 mm tall, 9-23 mm diam.
Outer involucre 7-13 mm diam; bracts 3.5-7.3 mm long,
2.3-5 mm wide. Fruits 1.4-2.2 mm long; hood apex muti-
40 Rhodora [Vol. 74
Fig. 16. Map of southern Texas and adjacent Mexico showing
distribution of Melampodium cinereum var. cinereum, small form
(circles), large form (dots); M. cinereum var. hirtellum (open
squares); M. cinereum var. ramosissimum (closed squares); and
M. argophyllum (triangles). Shaded area indicates generalized south-
eastern distribution of M. leucanthum.
cous to cirrhous, with appendage up to 2.2 mm long. Ray
florets 7-13; ligules 3-8.2 mm long, 1-3.6 mm wide. Disc
florets 30-50; corollas 1.1 mm diam, with throat 0.8 mm
and tube 0.7 mm long. Paleae 2.5 mm long, 1.1 mm wide.
Chromosome numbers, » — 10 and 20.
1972] Melampodium — Stuessy 41
Mesquite-grasslands of Rio Grande Plains of Texas south
to Cameron Co., east to Jim Wells Co., north to Bexar Co.
and west to Maverick Co., extending into Tamaulipas and
Nuevo Leon, Mexico, 30-240 m (Fig. 16). Flowering dates,
Mar-Nov.
All the varieties of M. cinereum can be distinguished
most easily from M. leucanthum and M. argophyllum by
the outer phyllaries that are connate only 1/4 their length
at the base. The outer involucral bracts of the latter two
species always are fused at least 1/2 or even 2/3 their
length. The usefulness of this feature has not been pre-
viously noted, but it is the best diagnostic morphological
character that I have discovered for separating M. cinereum
from the other taxa of the white-rayed complex.
Scattered within the range of var. cinereum are two
morphological forms (small and large) that are weakly
differentiated by quantitative features of habit and head
size. These informal units presumably correspond to the
2n and 4n autopolyploid chromosomal races reported within
this variety by Turner and King (1962) and Stuessy (1971b
& c). Additional information concerning the presumptive
autopolyploid origin of these races can be found in Stuessy
(1971c).
REPRESENTATIVE SPECIMENS (SMALL FORM). Mexico: NUEVO LEÓN :
S of Nuevo Laredo, 8 Jun 1935, Clark 6635 (NY); 15 mi N of
Sabinas Hidalgo, 12 Jun 1967, Stuessy 854 (TEX). TAMAULIPAS:
5 km S of Nuevo Laredo, 11 Nov 1961, Domínguez 63 (TEX); 24 km
S of Nuevo Laredo, 24 Mar 1962, Dominguez & McCart 8217 (SMU,
TEX); 59 mi N of Sabinas Hidalgo, 12 Jun 1967, Stuessy 857 (TEX).
UNITED STATES. Texas: CAMERON CO., Santa Ana National Wild-
life Refuge, 26 Sep 1961, Fleetwood 3840 (TEX); DIMMIT CO., 3 mi
W of Bigwells, 21 Apr 1945, Shinners 7395 (GH, LL, SMU, UC) ; DUVAL
co., 4 mi NE of Freer, 8 Oct 1954, Tharp & Johnston 541 [802A]
(TEX); FRIO CO., 5.5 mi S of Pearsall, 1 May 1954, Johnston, Tharp
& Turner 3472 (SMU, TEX); HIDALGO CO., 6 mi E of Sullivan City,
8 Mar 1959, Turner 4490 (TEX); JIM HOGG CO., 8 mi E of Hebbron-
ville, 12 Jun 1952, Jones 719 (SMU); JIM WELLS CO., near Alice,
21 Jun 1935, Drushel 10441 (NY); LA SALLE CO., 1 mi W of Encinal,
21 Apr 1963, Sánchez 105 (SMU, TEX); MAVERICK CO. 1 mi E of
Eagle Pass, 22 May 1898, Bray s.n. (TEX); STARR CO., 5.8 mi E of
42 Rhodora [Vol. 74
Rio Grande City, 9 Oct 1954, Tharp & Johnston 541922 (TEX);
WEBB CO., 22 mi NW of jct rtes 83 & 35, 5 Jul 1967, Stuessy 868
(TEX); ZAPATA CO., 13 mi N of San Ygnacio, 31 Jan 1954, Shinners
17659 (SMU); ZAVALA CO., 6 mi S of Batesville, 6 May 1964, Turner
5006 (SMU, TEX).
REPRESENTATIVE SPECIMENS (LARGE FORM). Mexico: TAMAU-
LIPAS: 3 mi S of Nuevo Laredo, 11 Nov 1961, Escalante 21 (TEX);
50 mi SE of Nuevo Laredo, 28 Mar 1964, A. & R. Garcia 47 (ARIZ,
SMU, TEX) ; 12 mi SE of Nuevo Laredo, 17 Mar 1962, C. & L. de la
Garza 46 (SMU, TEX). UNITED STATES. Texas: BEXAR CO., High-
land Hills, San Antonio, 1 Apr 1963, Martinez & García 13 (SMU);
BROOKS CO., 15 mi E of Hebbronville, 17 Mar 1968, Ramirez 44 (UL,
TEX); CAMERON CO., 12 mi N of Brownsville, 18 Apr 1965, Ríos &
Cavazos 264 (LL, SMU); DUVAL CO., 25 mi N of Hebbronville, 20 Jun
1966, Stuessy 429 (TEX); FRIO CO. 9 mi S of Moore, 20 Nov 1967,
Stuessy & Renold 1284 (TEX); HIDALGO CO., 1 mi E of Sullivan City,
1 Apr 1941, C. & A. Lundell 9866 (LL [3], RSA); JIM HOGG CO., 5 mi
E of Hebbronville, 20 Jun 1966, Stuessy 425 (TEX); JIM. WELLS CO.,
2 mi E of San Diego, 25 Nov 1962, Dohnke 3 (SMU, TEX); KINNEY
co., [?], Bracket [Brackettville?], 21 Mar 1900, Canby 133 (vs);
LA SALLE CO., 1 mi N of Encinal, 16 Mar 1963, Solis 124 (SMU, TEX) ;
LIVE OAK CO., ca 32 mi S of Whitsett, 3 Jun 1967, Stuessy 771 (TEX);
MCMULLEN CO., 22% mi SW of George West, 17 Apr 1965, Ríos &
Cavazos 231 (LL, SMU); MEDINA CO., ca 3 mi S of Devine, 28 Oct
1952, Correll 15204 (LL, US); NUECES CO., ca 4 mi W of Mathis,
8 May 1957, Jones 1359 (SMU); STARR CO., 3 mi N of Roma, 31 Jan
1954, Shinners 17708 (SMU); UVALDE CO., Leona River, 7 mi SE of
Uvalde, 23 Jun 1935, Munz 13303 (POM); WEBB CO., 14 mi NE of
Laredo, 9 Mar 1963, Cisneros 33 (LL); ZAPATA CO., 10 mi N of San
Ygnacio, 7 Apr 1963, Arzola 178 (LL, SMU).
7b. Melampodium cinereum DC. var. hirtellum Stuessy,
Sida 3:348. 1969. TYPE: MEXICO: Nuevo León, 5 km S of
Sabinas Hidalgo on rd to Monterrey, 21 Apr 1939, T. C. &
E. M. Frye 2415 (Holotype, US!; isotypes, DS! GH! NY!
RSA! UC!).
Stems 0.8-1.3 diam, strigose-hispid with hairs 0.2-1.5
mm long. Peduncles 0.7-8 cm long. Leaves linear-oblong,
1.1-3.4 em long, 0.2-1 cm wide, with both surfaces strigose
with hairs 0.3-0.8 mm long, with midrib on undersurface
hispid with hairs up to 1.5 mm long; margin pinnately 8-
lobed or parted, near base hispid with hairs 0.4-1.5 mm
long. Heads 4.5-5 mm tall, 9-18 mm diam. Outer involucre
7-10 mm diam; bracts 3.3-6 mm long, 2.2-3.6 mm wide.
1972] Melampodium — Stuessy 43
Fruits 1.3-2 mm long; hood apex muticous to mucronate,
with appendage up to 0.3 mm long. Ray florets 8-13; ligules
3-6 mm long, 1.3-2.8 mm wide. Disc florets 25-50; corollas
1.1 mm diam, with throat 0.7 mm and tube 0.5 mm long.
Paleae 3.2 mm long, 0.9 mm wide. Chromosome number,
n — 10.
Mesquite-grasslands in eastern Coahuila, northwestern
Nuevo León and Tamaulipas, Mexico, and into Val Verde,
Kinney, Uvalde and Maverick Cos. of Texas (Fig. 16),
210-520 m. Flowering dates, Feb-Oct.
REPRESENTATIVE SPECIMENS. Mexico: COAHUILA: Saltillo, Aug
1913, Adole 29 (F, GH, NY, US); Sabinas, 16 Sep 19957, Kenoyer 5
(F); 30 mi S of Monclova, 14 Aug 1948, Kenoyer & Crum 2594 (A);
Muzquiz, Apr 1938, Marsh 1176 (F, GH, TEX); Soledad, 25 mi SW
from Monclova, 9-19 Sep 1880, Palmer 556 (GH, NY, US); Rio
Grande Valley near Diaz, 17 Apr 1900, Pringle 9008 (F, GH, US);
4 mi W of Nueva Rosita, 14 Aug 1967, Stuessy 902 (TEX); 21 mi S
of Monclova on rte 57, 14 Aug 1967, Stuessy 912 (TEX); 23 mi N
of Sabinas, 23 Aug 1959, Waterfall 15805 (F, SMU); Monclova,
25 mi SW of Sabinas, 19 Jun 1936, Wynd & Muller 214 (ARIZ, FSU,
GH, NY, US). NUEVO LEÓN: Monterrey, Obispado, Jul 1911, Abloon
187 (us); 36 mi NE of Sabinas Hidalgo, 24 Mar 1944, Barkley
14575C (GH, NY); between Laredo & Monterrey, 8 Feb 1945, Bonner
55 (F); 12 mi N of Sabinas Hidalgo, 26 Mar 1944, Heard & Barkley
14542B (TEX); Cerro del Obispado, 20 Oct 1946, Lacás 58 (F); 8 mi
S of Sabinas Hidalgo, 8 Feb 1964, May 13 (sMU); 12 mi W of
Monterrey, 27 Feb 1944, Painter, Lucas & Barkley 14297 (TEX); 47
mi S of Nuevo Laredo, 28 Apr 1962, Reséndez 52 (SMU, TEX) ; 17 mi
NE of Sabinas [Hidalgo], 20 Mar 1963, Rodriguez 72 (SMU);
26 mi N of Sabinas Hidalgo, 12 Jun 1967, Stuessy 855 (TEX);
38 mi N of Sabinas Hidalgo, 12 Jun 1967, Stuessy 856 (TEX).
UNITED STATES. Texas: KINNEY C0., 9 mi W of Brackettville,
28 Mar 1947, McVaugh 7694 (DS, F, LL [2], SMU, TEX); MAVERICK
co., 30 mi SE of Eagle Pass, 14 Mar 1964, Bruni 15 (LL, SMU, TEX) ;
UVALDE CO. 2 mi N of Cline, 8 Jul 1936, Hedrick 156 (UC); VAL
VERDE C0., Del Rio, 20 Apr 1930, Jones 25916 (DS, POM).
7c. Melampodium cinereum DC. var. ramosissimum (DC.)
A. Gray, Smithson. Contrib. Knowledge 3, Art. 5: 103.
1850.
Melampodium ramosissimum DC. Prodr. 5:518. 1836.
TYPE: TEXAS[?]: between the Nueces River & Laredo,
44 Rhodora [Vol. 74
Jul 1829, J. L. Berlandier 2017 [—607] (Holotype, G-DC;
isotypes, F! G! GH[2]! HAL! K! Mo[2]! w[2]! wis!; photo-
graph of holotype, Ipc 800. 927: II. 1!; photograph of K
isotype,US!; photograph of HAL isotype, OS! TEX!).
Melampodium cinerascens S. F. Blake, Contrib. U. S.
Nat. Herb. 22:605. 1924. TYPE: MEXICO: Tamaulipas,
Hacienda Buena Vista, ca 20 mi E of Abasolo, 18 Jun 1919,
E. O. Wooton s.n. (Holotype, US!; photograph of holotype,
TEX!).
Stems 0.7-0.8 mm diam, strigose with hairs 0.1-0.8 mm
long. Peduncles 0.7-3.8 cm long. Leaves linear, 0.7-3.2 cm
long, 0.2-0.5 em wide, with both leaf surfaces strigose with
hairs 0.2-1 mm long; margin entire to pinnately 10-lobed
or cleft, near base strigose with hairs up to 1 mm long.
Heads 3-5 mm tall, 5-14 mm diam. Outer involucre 4.5-9
mm diam; bracts 2.7-4 mm long, 1.3-2.8 mm wide. Fruits
1.5-1.7 mm long; hood apex mucronate to cirrhous (rarely
muticous), with appendage up to 1.5 mm long. Ray florets
7-8; ligules 2-3.8 mm long, 1-2.3 mm wide. Disc florets
25-35; corollas 1.8 mm diam, with throat and tube each
0.4 mm long. Paleae 2.5 mm long, 0.9 mm wide. Chromo-
some number, n —10.
Mesquite-grasslands of northern Tamaulipas, Mexico, and
adjacent Hidalgo, Cameron and Webb Cos. of Texas (Fig.
16), 15-90 m. Flowering dates, Jun-Dec.
There is some character intergradation where the three
varieties of M. cinereum overlap near the Rio Grande Val-
ley of southern Texas (Fig. 16). Although no detailed
studies have been carried out to clarify the nature of this
intergradation, hybridization is likely to be occurring.
REPRESENTATIVE SPECIMENS. Mexico. TAMAULIPAS: 20 mi E of
San Fernando-Santander Jiménez hwy on rd to Loreto (11 mi W
of Loreto), 15 Sep 1960, Crutchfield & Johnston 5527 (TEX); 13 mi
E of Abasolo turnoff on the Santander-Jiménez-Pesca rd, 15 Dec
1960, Crutchfield & Johnston 6141b (TEX); 48 mi S of Reynosa,
19 Oct 1959, Graham & Johnston 4878 (TEX); 1 mi N of San
Fernando, 4 Jul 1966, Stuessy 450 (TEX); ca 2 mi SE of Reynosa,
4 Jun 1967, Stuessy 778 (TEX); 27 mi S of Reynosa, 4 Jun 1967,
1972] Melampodium — Stuessy 45
Stuessy 787 (TEX); 3 mi S of San Fernando, 18 Nov 1967, Stuessy
& Renold 1261 (TEX). UNITED STATES. Texas: HIDALGO CO.,
Mission, Lomita Alta, 13 Jul 1937, Cameron 248 in part (TEX) ;
McAllen, 10 Aug 1937, Cameron 248 in part (TEX); La Joya gravel
pit, 27 Aug 1942, Walker 93 (LL, TEX); WEBB CO., 11 mi W of
Encinal, 5 Jul 1960, H. & F. Iltis & Koeppen 35 (wIs).
8. Melampodium argophyllum (A. Gray ex Robins.) S. F.
Blake, Contrib. U. S. Nat. Herb. 22:606. 1924.
Melampodium cinereum DC. var. argophyllum A. Gray
ex Robins. Proc. Amer. Acad. Arts & Sci. 36:458. 1901.
(A. Gray in S. Wats. Proc. Amer. Acad. Arts & Sci. 18:
104. 1883. nom nud., but based on the type collection)
TYPE: MEXICO: Coahuila Sierra Madre S of Saltillo [ca
60 km SE of Saltillo; McVaugh, 1956], Feb-Oct 1880, FE.
Palmer 2068 (Holotype, GH!).
Melampodium leucanthum Torr. & A. Gray var. argo-
phyllum (A. Gray ex Robins.) Stuessy, Sida 3:348. 1969.
Perennial subshrubs, 12-22 cm tall. Stems ascending, 0.8-
1.2 mm diam, canescent-tomentose with hairs 0.2-1 mm
long. Peduncles 3.5-7.7 cm long. Leaves sessile, linear-ob-
long, 1-2.8 cm long, 0.2-1.5 cm wide, at apex and base obtuse,
with both surfaces tomentose with hairs 0.2-1 mm long;
margin pinnately 8-lobed or parted (rarely entire), near
base strigose-hispid with hairs 0.5-0.7 mm long. Heads
4-7 mm tall, 10-23 mm diam. Outer involucre cupulate,
6.2-12 mm diam; bracts 5, connate 1/2 to 3/5 their length,
ovate, 3.5-6 mm long, 2.5-5 mm wide, at apex acute, with
abaxial surface tomentose with hairs 0.1-1 mm long; mar-
gin herbaceous. Fruits 0.8-1.9 mm long, with lateral sur-
faces aculeate-verrucate; hood apex muticous to cirrhous
(usually mucronate). Ray florets 8-9; ligules cream-white,
oblong-elliptic, 2.5-9 mm long, 1.5-4.5 mm wide. Disc florets
25-50; corollas yellow, 1 mm diam, throat and tube each
0.8 mm long. Paleae oblong-elliptic, 2.6 mm long, 1.2 mm
wide; apex yellow, with margin laciniate; midrib promi-
nent, often strigillose with hairs 0.1 mm long. Chromosome
number unknown.
46 Rhodora [Vol. 74
Arid mountains in western Nuevo León and adjacent
Coahuila, Mexico (Fig. 16), 1830-2440 m. Flowering dates,
Feb-Oct.
Although M. argophyllum was first described as a variety
of M. cinereum, in subsequent years the status of the
former taxon has been changed twice. Morphologically M.
argophyllum shows similarity to both M. leucanthum and
M. cinereum but seems to have a greater total resemblance
to the fcrmer species. Due to recent chromatographic stud-
ies on M. argophyllum and other taxa of the white-rayed
complex (Stuessy, 1971c), however, I have reversed my
earlier judgment (1969a) based solely on morphological
criteria which treated M. argophyllum as a variety of M.
leucanthwm. I now concur with Blake's (1924) recognition
of M. argophyllum as being a distinct species. Refer to my
recent paper (1971c) for additional discussions on the sta-
tus of M. argophyllum and on hypothetical evolutionary
relationships within the entire white-rayed complex.
REPRESENTATIVE SPECIMENS. Mexico. COAHUILA: G. Cepeda, Sierra
de la Paila (V. Seco), 6 Jul 1944, Hinton et al. 16563 (GH, NY, US);
45 km SW of Monterrey, 28 Feb 1946, Johnson & Barkley 16250M
(TEX); Castanos, “Puerto de San Lázaro," Sierra de San Lazar,
30 Aug 1939, Muller 3040 (F, GH, UC); Sierra de la Pata Solana,
Mar 1905, Purpus 1003 (F, GH, NY, UC), Feb 1905, Purpus 1003a
(uc); Sierra de la Paila, Oct 1910, Purpus 4730 (uc); Sierra de
Parras, Mar 1905, Purpus s.n. (UC); Cañon Espantosa, W slope of
Sierra de Saa Vicente, ca 20 km ESE of Cuatro Cienegas, 25 Mar
1941, Schroeder 120 (GH); 9 km S of Parras on Sierras Negras,
3 Jul 1941, Stanford, Retherford & Northcraft 186 (ARIZ, DS, F,
GH, NY [2], UC); Castanos, San Lázaro, “rocky slopes of El Puerto
de San Lázaro," 16 Jun 1936, Wynd & Muller 132 (ARIZ, US). NUEVO
LEÓN: Las Salinas, 1924, Orcutt 1395 (us); 24 km W of Icamole,
3 Feb 1907, Safford 1264 (us).
3. Series Sericea Stuessy, ser. nov.
Herbae annuae; folia lanceolata vel pinnatim divisa, sub-
tus sericea vel strigosa; ligulae luteae, minus quam vel raro
circa 2 mm longae; involucrum extimum marginibus her-
baceis; fructi plerumque cucullati. Species 9-13. Typus:
Melampodium sericeum Lag.
1972] Melampodium — Stuessy 47
9. Melampodium sericeum Lag. Gen. et Sp. Nov. 32. 1816.
non H.B.K. 1820. TYPE: “NOVA HISPANIA": 1787-1804,
seeds collected by M. Sessé & J. M. Mocino s.n. (Holotype,
MA?; isotype, G!; photograph of G isotype, F! os! TEX! US!;
photograph of G-DC isotype, IDC 800. 937: I. 8!).
Melampodium hispidum H.B.K. Nov. Gen. Sp. 4:273.
t. 399. 1820. TYPE: MEXICO: Michoacán, between Pazcuaro
& Valladolid [Morelia], 6000 ft, Sep 1803, F. H. A. von
Humboldt & A. J. Bonpland. s.n. (Holotype, P; fragments
of holotype, P!).
Melampodium sericeum Lag. var. brevipes A. Gray, Proc.
Amer. Acad. Arts & Sci. 22: 423. 1887. TYPE: MEXICO:
Guanajuato, León, W of Guanajuato, 1829, “Menzies”
[probably J. Méndez] s.n. (Holotype, GH!; photograph of
G-DC isotype, TEX!; tracing of probable isotype [G?], GH!).
Melampodium sericeum Lag. var. exappendiculatum
Robins. Proc. Amer. Acad. Arts & Sci. 36:459. 1901. TYPE:
MEXICO: San Luis Potosí, “in mountains near Morales,"
Aug 1876, J. G. Schaffner 271 (Lectotype chosen, GH!).
Annual herbs, 3-40 cm tall. Stems ascending to erect,
0.4-2.5 mm diam, hispid-pilose with hairs up to 1 mm long.
Peduncles 5-22 mm long. Leaves sessile, linear-oblong to
oblanceolate, 2-6.5 em long, 0.2-3 em wide, at apex acute-
obtuse, at base attenuate to obtuse, with upper surface
strigose with hairs 0.5 mm long, with lower surface serice-
ous; margin entire to pinnately parted with 1-4 lobes, ir-
regularly ciliate with hairs 0.2 mm long, at base hispid with
hairs 1-1.5 mm long. Heads 4-5 mm tall, 4-8 mm diam.
Outer involucre cupulate, 6-11 mm diam; bracts 5, slightly
connate at base, separate to imbricate, narrowly rhombic
to obovate, 3-7 mm long, 2-4 mm wide, at apex acute, with
abaxial surface pilose to hispid with hairs 1-1.5 mm long;
margin herbaceous. Fruits 2.5-3.2 mm long, with lateral
surfaces smooth and striate to strongly verrucate; hood
apex muticous to cirrhous, with tapering appendage up to
2 mm long. Ray florets 5-7; ligules yellow, narrowly ovate,
0.8-1.2(2.2) mm long, 0.6-0.8(—1.2) mm wide. Disc florets
Cruz, Sep 1851, G. Thurber 937 (Lectotype, GH! K! Mo!
48 Rhodora [Vol. 74
Fig. 17. Map of Mexico and adjacent Central America showing
distribution of Melampodium sericeum.
(—5) 8-12; corollas yellow, 1.1 mm diam, with throat 0.6 mm
and tube 0.5 mm long. Paleae oblanceolate, 3.5 mm long,
0.3 mm wide; apex yellow, with margin dentate-erose; mid-
rib prominent, glabrous. Chromosome number, n = 30.
Pine-oak forests of Mexico, Guatemala and El Salvador
(Fig. 17), 1100-2590 m. Flowering dates, Jul-Sep.
This is a widespread, variable species. Although no
varieties are recognized here in M. sericeum, there are sev-
eral morphologically distinct collections that should be
noted. One specimen from Guatemala, Steyermark 50530,
is unusually robust and has brown-purple disc florets. Also
from Guatemala are several collections, Morales 1300, Rojas
86, Standley 59769 and 82849, which possess fruits with
1972] Melampodium — Stuessy 49
broad, wide-spreading hoods in contrast to the much nar-
rower hoods typical of the species.
REPRESENTATIVE SPECIMENS. El Salvador. SANTA ANA: near Chal-
chuapa, 1922, Calderón 1019 (GH, US). Guatemala. BAJA VERAPAZ:
Santa Rosa, Oct 1912, Tuerckheim 3955 (US). ESCUINTLA: 1942,
Ignacio 1567 (F). GUATEMALA: Laguna, Amatitlan, 1929, Morales
1300 (F); Chilloni, 21 Jun 1921, Rojas 86 (GH, NY, US); Finca
Bretafia, rd between Guatemala & Riscal, 12 Dec 1938, Standley
59769 (F, NY, SMU). HUEHUETENANGO: ca 3 mi S of Huehuetenango,
18 Jul 1960, King 3416 (NY, TEX, UC); Huehuetenango, 9 Nov 1934,
Skutch 1592 (GH); between San Sebastian H. & Rio San Juan,
9 Jan 1941, Standley 82849 (F, NY, US); between Huehuetenango &
San Sebastian H., 12 Aug 1942, Steyermark 50392 (F, GH, US);
between San Sebastián H. & San Rafael Pétzal, 14 Aug 1942,
Steyermark 50530 (F). QUICHE: 1942, Ignacio 1404 (F). Mexico.
CHIHUAHUA: La Mesa Colorada, 14 Oct 1939, Gentry 550 [550M]
(ARIZ, DS, F); Loreto, Río Mayo, 1 Sep 1936, Gentry 2553 (ARIZ, GH,
F, UC, US); Temosachie, Cañón Huahuatán, 10 mi SE of Madera,
23 Sep 1939, Muller 3484 (GH, UC). COLIMA: Alzada, 25 Aug 1966,
Stuessy 730 (TEX). DISTRITO FEDERAL: Olivar, 15 Aug 1910, Orcutt
3605 (F); above Santa Fe, 4 Sep 1901, Pringle 8609 in part (F, GH,
NY, POM, UC, US [2]). DURANGO: vicinity of Durango, Iron Mt,
Oct 1896, Palmer 926 (GH, US). GUANAJUATO: 20 mi NE of Irapuato,
28 Aug 1947, Barkley, Rowell & Paxson 735 (TEX); near Guanajuato,
Sep [year?], Dugés 44 (GH [2]); Guanajuato, 1909, Furness s.n.
(F); near Acambaro, 6 Oct 1892, Pringle 5309 (GH); 14 mi SE of
León, 16 Aug 1957, Waterfall & Wallis 13886 (F, SMU). GUERRERO:
Petaquillas, 27 Aug 1965, Stuessy 364 (TEX). JALISCO: 56 mi SE
of Guadalajara, 21 Aug 1953, W. & M. Manning 531212 (GH, TEX);
ca 11 mi SE of Lagos de Moreno, 7 Sep 1952, McVaugh 12822 (SMU,
US); Guadalajara, Jul 1886, Palmer 260 (GH, US); near Guadalajara,
24 Aug 1901, Rose & Hay 6275 (GH, US); 2 mi NW of Tequila,
26 Aug 1966, Stuessy 740 (TEX). MEXICO: Temascaltepec, Plaza de
Gallos, 22 Jul 1934, Hinton et al. 6324 (GH, NY, US). MICHOACÁN:
ca 10 mi W of Morelia, 5 Aug 1960, King 3626 (Ds, NY, TEX, UC, US) ;
Cerro Potrerillos, ca 5 mi N of Cotija, 5-9 Oct 1961, King & Soder-
strom 4584 (NY, SMU, TEX, UC, US) ; 0.8 mi NW of Zitacuaro, 1 Sep
1965, Stuessy 377 (TEX); 7 mi S of Ario de Rosales, 20 Aug 1966,
Stuessy 690 (TEX); 11 mi NW of Zamora, 23 Aug 1966, Stuessy 703
(TEX). MORELOS: Cuernavaca, 31 Aug 1910, Orcutt 3887 (F).
NAYARIT: 5 mi SE of Ixtlán del Río, 16 Aug 1961, Waterfall 16342
(SMU, UC). OAXACA: 53 mi S of Tehuacán, 13 Aug 1961, Powell &
Edmondson 660 (F, TEX); Valley of Oaxaca, 5 Jul 1897, Pringle
6728 (F, GH, NY, UC, US [2]) ; 38 mi S of Zimatlán, 12 Aug 1966,
50 Rhodora [Vol. 74
Stuessy 642 (TEX); 4 mi S of Sola de Vega, 13 Aug 1966, Stuessy
647 (TEX); % mi NW of Las Sedas, 15 Aug 1966, Stuessy 660 (TEX).
PUEBLA: Los Molinos between Puebla & Atlixco, 18 Sep 1944, Sharp
44961 (NY). QUERÉTARO: Jul 1904, Kuntze 27465 (Ny [2]); 6 mi
W of Querétaro, 6 Aug 1961, Powell & Edmondson 579 (F, TEX);
4.7 mi N of Querétaro, 8 Aug 1959, Rock M-442 (TEX). SAN LUIS
PoTOsÍ: 15-16 mi E of Ciudad del Maiz, 20 Sep 1960, Crutchfield &
Johnston 5664 (TEX); 3 mi E of Ciudad Maiz, 2 Sep 1948, Kenoyer
& Crum 4093 (GH); near San Luis Potosí, 1877, Schaffner 270 in
part (NY). SONORA: ca 2 mi W of Tres Ríos (ca 88 mi W of Vieja
Casa Grandes, Chihuahua), 29 Aug 1952, Tucker 2565 (ARIZ, US);
Cañón de Huépari, N of Aribabi, 2-3 Sep 1939, White 2642 in part
(Gi). ZACATECAS: between Colotlán & Plateado, 31 Aug 1897,
Rose 3617 (F, US).
10. Melampodium pringlei Robins. Proc. Amer. Acad. Arts
& Sci. 36:461. 1901. TYPE: MEXICO: Oaxaca, Las Sedas,
1850 m, 15 Sep 1894, C. G. Pringle 5722 (Holotype, GH!).
Annual herbs, 30 em tall. Stems ascending, 1-2 mm diam,
hirsute-hispid with hairs up to 1 mm long. Peduncles 1-2
mm long. Leaves sessile, linear-oblong to oblanceolate, 2-
2.8 em long, 0.7-1.6 em wide, at apex acute-obtuse, at base
attenuate to obtuse, with upper surface strigose with hairs
1 mm long, with lower surface copiously sericeous; margin
entire to 2-lobed, irregularly ciliate with hairs 0.3 mm long,
near base hirsute with hairs 1-2 mm long. Heads 3-4 mm
tall, 3.5-4 mm diam. Outer involucre cupulate, 3-4 mm
diam; bracts 5, slightly connate at base, separate, lanceo-
late, 3-3.5 mm long, 1 mm wide, at apex acute-acuminate,
with abaxial surface hirsute with hairs 1 mm long; margin
herbaceous. Fruits 3 mm long, with lateral surfaces 3-
ribbed with large tubercles. Ray florets 3-4; ligules yellow,
with undersurface near apex light purple, ovate, 1 mm
long, 0.5 mm wide. Disc florets 2-3; corollas yellow, 1.1 mm
diam, with throat 0.6 mm and tube 0.5 mm long. Paleae
oblanceolate, 2.5 mm long, 0.8 mm wide; apex purple, with
margin erose-dentate; midrib prominent, weakly pilose with
hairs 0.3 mm long. Chromosome number unknown.
Known only from the type collection from pine-oak for-
ests near Las Sedas, Oaxaca, Mexico (Fig. 11), 1860 m.
Flowering date, Sep.
1972] Melampodium — Stuessy 51
Although this taxon is very similar to M. sericeum, it
seems sufficiently distinct to be maintained as a separate
species at the present time. Additional collections, however,
may indicate that varietal status is preferable. I have
searched the countryside around Las Sedas, the type lo-
cality, but have been unsuccessful in rediscovering the
taxon. Characters (in addition to those listed in the key)
which distinguish M. pringlei from its closeset relative are:
M. sericeum M. pringlei
1. Heads 4-5 mm tall, 4-8 1. Heads 3-4 mm tall,
mm diam. 3.5-4 mm diam.
2. Outer involucral bracts 2. Outer involucral bracts
narrowly rhombic to lanceolate, 3-3.5 mm
obovate, 3-7 mm long, long, 1 mm wide.
2-4 mm wide.
3. Ray florets 5-7. 3. Ray florets 3-4.
4. Paleae yellow-tipped. 4. Paleae purple-tipped.
11. Melampodium strigosum Stuessy, sp. nov.
Melampodium coronopifolium Sch. Bip. ex Hemsl. Biol.
Centr. Am. Bot. 2:145. 1881. nom. nud. Based on several
specimens including C. C. Parry & E. Palmer 444 1/2 (GH!
K! MO! NY! US!).
Herbae annuae, 3-35 cm altae. Caules ascendentes, 0.6-2
mm diametro, moderate hispido-strigillosi, pilis 0.3-1 mm
longis. Pedunculi 0-3(-11) mm longi. Folia sessilia, ob-
longo-linearia vel oblanceolata, 0.6-5 cm longa, 0.2-1.5 cm
lata, apice acuto-obtusa, basi obtuso-subauriculata, utrinque
strigosa pilis 0.2-0.6 mm longis, marginibus integris vel
pinnatim 2-4 lobatis, basalibus strigoso-hispidis, pilis 0.3-
1.1 mm longis. Capitula 3-4 mm alta, 4-8 mm diametro.
Involucrum extimum cupulatum, 3-8 mm diametro; bracteae
9, basi leviter connatae, separatae, lanceolato-ellipticae, 2.7-
6 mm longae, 1-2.9 mm latae, apice acutae, extus strigoso-
hispidae in nervis principalibus, pilis 0.3-0.8 mm longis,
marginibus herbaceis. Fructus 2.2-3 mm longi, lateribus
striatis vel verrucatis, cucullis apice muticis vel cirrhosis,
52 Rhodora [Vol. 74
Fig. 18. Map of Mexico and adjacent United States showing dis-
tribution of Melampodium strigosum (dots), M. sinaloense (square),
and M. tepicense (triangles). One collection of M. strigosum from
Colorado not shown.
1972] Melampodium — Stuessy 53
mucrones 2-2.5 mm longis. Flosculi radii 5-8; ligulae flavae,
oblongo-ovatae, 0.6-1.1 mm longae, 0.5-0.8 mm latae. Flos-
culi disci 4-6; corollae flavae, 1-2 mm diametro, faucibus 0.6
et tubis 0.5 mm longis. Paleae oblongo-ellipticae, 2.3 mm
longae, 0.8 mm latae, apice flavae, marginibus dentatis, cos-
tis conspicuis, glabris vel puberulis pilis 0.1 mm longis.
Chromosomatum numerus, n = 20.
TYPUS: MEXICO: Chihuahua, ca 5 mi E of Ciudad Guer-
rero on rte 16, 21 Aug 1967, T. F. Stuessy 1054 (Holotype,
US!; isotypes, F! GH! MICH! NY! os! SMU! TEX! UC!).
Pine-oak forests of central and northwestern Mexico,
reaching Cochise, Pima and Santa Cruz Cos. of southeast-
ern Arizona, Jeff Davis Co. of Texas, and Chaffee Co. of
Colorado (Fig. 18), 1310-2590 m. Flowering dates, Aug-
Oct.
This new species is a common taxon that has masquer-
aded under the name of M. hispidum H.B.K. for many
years. Dr. Rogers McVaugh first mentioned to me that the
holotype of M. hispidum in the herbarium at Paris seemed
conspecific with the even more common species, M. sericeum
Lag. Having obtained additional information from Dr.
Alicia Lourteig at Paris regarding the holotype of M. his-
pidum, I believe it is definitely conspecific with M. sericeum.
The placement of M. hispidum in synonymy with M.
sericeum leaves M. coronopifolium Sch. Bip. ex Hemsl. as
the only other name applicable to this taxon, but this latter
epithet is a nomen nudum and therefore not validly pub-
lished. By reference to three specimens cited in the proto-
logue as being representative, however, M. coronopifolium
is clearly referable to this species. I could now validate
this available epithet, but I prefer to name this undescribed
Species with an epithet that emphasizes the distinctive
strigose pubescence of the leaves in contrast to the mark-
edly sericeous leaves of the morphologically similar M.
sericeum.
REPRESENTATIVE SPECIMENS. Mexico. CHIHUAHUA: Parral, 19 Sep
1898, Goldman 110 (NY, Us); San Diego Canyon, Sierra Madre Mts,
16 Sep 1903, Jones s.n. (POM); Sta Clara Mts, 10-19 Oct 1935,
54 Rhodora [Vol]. 74
LeSueur 341 (ARIZ, F, GH, TEX); near Chihuahua, Sep-Oct 188»,
Pringle 297 (F, GH, NY, US [2]), 1 Jun 1885, Pringle 10 (F, GH, K,
NY [2], US); mts near Chihuahua, Sep 1886, Pringle 754 (F, NY
[2], uc [2], us [2]); near Chihuahua, 12 Sep 1886, Pringle 1045
(ps, NY, UC); ca 2-3 mi W of Parral, 18 Aug 1967, Stuessy 1016
(TEX); 2 mi S of Cuauhtémoc, 20 Aug 1967, Stuessy 1038 (TEX) ;
near Colonia García in Sierra Madre Mts, 26 Sep 1899, Townsend
& Barber 351 (GH, NY, US). DISTRITO FEDERAL: Pyramid of Cuicuilco,
Tlalpan, 1 Sep 1936, MacDaniels 747 (F); Valley of Mexico, 9 Sep
1896, Pringle 6491 (F, GH, NY, UC, US); above Santa Fe, Valley of
Mexico, 4 Sep 1901, Pringle 8609 in part (GH); near Club Golf de
Chapultepec, 4 Sep 1946, Zamora, Paxon & Barkley 16M905 (TEX).
DURANGO: vicinity of Durango, Apr-Nov 1896, Palmer 486 (F, GH,
NY, UC, US). GUANAJUATO: Guanajuato, Aug 1899, Dugès s.n. (US) ;
Guanajuato, 1909, Furness s.n. (F). HIDALGO: near El Salto, 17 Sep
1901, Pringle 9331 (GH, NY, US); near Landa Station, 3 Sep 1903,
Pringle 11548 (ARIZ, F, GH, SMU, US). QUERÉTARO: 22 mi NE of
San Juan del Río, 10 Sep 1962, Turner & Powell 1116 (TEX) ; 14 mi
SE of San Juan del Río, 17 Aug 1957, Waterfall & Wallis 13960
(SMU, US). SONORA: Cañón de Huépari N of Aribabi, 2-3 Sep 1939,
White 2642 in part (ARIZ), 7 Sep 1939, White 2788 (ps). UNITED
STATES. Arizona: COCHISE CO. Chiricahua Mts, 18 Sep 1907,
Blumer 1665 (ARIZ, F, GH, NY, US); Portal, Chiricahua Natl Forest,
17-19 Sep 1914, Eggleston 10729 (us); Lanner Canyon, Huachuca
Mts, 24 Aug 1910, Goodding 814 (ARIZ, GH, US); Huachuca Mts,
20 Aug 1893, Holzner 1962 (vs); Chiricahua Mts, 22 Sep 1931,
Jones s.n. (POM); Huachuca Mts, near Ft Huachuca, Sep 1882,
Lemmon 2777 (F, GH, K, NY, UC); PIMA CO., Rincon Mts, Dec 1907,
Goodding 18 (ARIZ); SANTA CRUZ CO., Nogales, 25 Oct 1926, Jones
22681 (pom); Patagonia Mts, 18 Aug 1928, Kearney 5587A (us);
Nogales, 7 Aug 1927, Peebles, Harrison & Kearney 4614 (us).
Colorado: CHAFFEE CO., Buena Vista, Jones s.n. (POM). Texas: JEFF
DAVIS co., Davis Mts, Madera Canyon, Aug 1936, Hinckley 620
(NY), 8 Oct 1936, Hinckley 881 (ARIZ, F, GH, LL, NY [2], SMU);
Davis Mts, Madera Creek, old Fisher Ranch House, 16 Sep 1944,
Hinckley 3293 (LL, US); Madera Canyon, Mt Livermore, Aug 1936,
Hinckley sm. (TEX); Madera Canyon, Davis Mts, 25 Aug 1967,
Stuessy 1128 (TEX); 11 Sep 1959, Turner 4620 (SMU, TEX) ; N of
Merrill ranch on rd to Mt Locke, 13 Aug 1950, Warnock 9229 (SMU) ;
upper Madera Canyon, Davis Mts, 5 Oct 1955, Warnock 13507 (LL) ;
Davis Mts, 18 Sep 1918, Young s.n. (TEX).
12. Melampodium longicorne A. Gray, Mem. Amer. Acad.
ser. 2. 5:321. 1855. TYPE: MEXICO: Sonora, near Santa
1972] Melampodium — Stuessy 55
Cruz, Sep 1851, G. Thurber 937 (Lectotype, GH!; isotypes,
GH! K! MO! NY!; photograph of K isotype, USE
Annual herbs, 12-60 cm tall. Stems erect, 0.9-2.8 mm
diam, subglabrous to hispid-strigose with hairs 0.5-1 mm
long. Peduncles 4-30 mm long. Leaves sessile, linear-ob-
long to oblanceolate, 3-5 em long, 0.3-1.2 cm wide, at apex
obtuse, at base truncate-subauriculate, with both surfaces
strigose with hairs 0.3-0.8 mm long; margin entire to ob-
scurely dentate, at base hispid-hispidulous with hairs 0.5-1
mm long. Heads 4-7 mm tall, 7-13 mm diam. Outer in-
volucre cupulate, 6-8(—10) mm diam, bracts 5, slightly con-
nate at base, usually imbricate 2/3 their length (less often
separate), elliptic to obovate-rhombic, 4.3-6.8 mm long, 2-4
mm wide, at apex acute, with abaxial surface moderately
strigose with hairs 0.5-1 mm long; margin herbaceous.
Fruits 5-3.6 mm long, with lateral surfaces granulate to
aculeate-verrucate; hood apex cirrhous, with appendage
over 4 mm long. Ray florets 7-12; ligules yellow, oblong-
elliptic, 1.2-1.5 mm long, 0.7-0.9 mm wide. Disc florets 8-10;
corollas yellow, 1 mm diam, with throat 0.5 mm and tube
0.4 mm long. Paleae oblong-elliptic, 3.4 mm long, 1.1 mm
wide; apex yellow, with margin bidentate; midrib promi-
nent, strigillose with hairs 0.1 mm long. Chromosome num-
ber, n = 30.
Mountains of southeastern Arizona in Cochise, Pima and
Santa Cruz Cos., and in adjacent Sonora, Mexico (Fig. 19),
1130-1830 m. Flowering date, Sep.
Found exclusively in southeastern Arizona and neighbor-
ing Sonora, this species is often conspicuous with its long
coiled awns on the hoods of the ray achenes. Just as often,
however, the awns are absent, making separation from the
morphologically similar M. strigosum difficult. In the area
of sympatry in Arizona, there is a trend in M. strigosum
toward longer peduncles, more entire leaves, and more erect
stems, suggesting that hybridization may be occurring be-
tween the two taxa.
Asa Gray seems to have experienced some difficulty in
delimiting M. longicorne, even though he himself described
56 Rhodora [Vol. 74
Fig. 19. Map of northwestern Mexico and adjacent United States
showing distribution of Melampodium appendiculatum (squares),
M. cupulatum (triangles), M. longicorne (circles), and M. sinuatum
(dots).
it as new. He first (1855) confused a Wright 1205 collec-
tion (distinct M. longicorne) with M. strigosum and later
(1859) emended his original description to include a Schott
72 collection, the latter being treated subsequently as be-
longing to M. appendiculatum (Robinson, 1901; present
treatment).
1972] Melampodium — Stuessy 57
Confusion has existed regarding the correct spelling of
the specific epithet, and some workers (Robinson, 1901;
Tidestrom and Kittell, 1941; Gentry, 1942) have elected to
use the form longicornu rather than longicorne as originally
published. Even Gray himself ceased to use the original
adjectival spelling and reverted to the substantive form
longicornu in his subsequent publications of 1859 and 1884.
However, as longicorne is the original correct spelling, I
have used this form in the present treatment, as have other
workers in recent times (Kearney and Peebles, 1942, 1964;
Shreve and Wiggins, 1964).
REPRESENTATIVE SPECIMENS. Mexico. SONORA: 2 mi E of Agua
Prieta-Nacozari rd to Oputo & Angostura, 7 Sep 1961, Turner, Dodge
& Mason 2068 (ARIZ, DS); near Santa Cruz, 1851, Wright 1205 (G, GH
[3], K, NY, US). UNITED STATES. Arizona: COCHISE CO., 10 mi
N of Chiricahua Station, 8 Sep 1942, Barneby 5135 (NY); Chiri-
cahua Mts, N of Wilgus Ranch, 2 Sep 1907, Blumer 2128 (F, GH,
NY); Ft Huachuca Military Reservation, 29 Sep 1962, Goodding
267-62 (ARIZ); Huachuca Game Preserve, Garden Canyon, 21 Sep
1949, Goodding 580-49 (ARIZ [2]) ; Montezuma Canyon, Huachuca Mts,
17 Oct 1958, Goodding 699-58 (ARIZ, UC); Ramsey Canyon, Huachuca
Mts, 29 Sep 1929, Jones 25045 (Ny [2], POM, TEx [2], vc [2]);
Huachuca Mts, 3 Sep 1903, Jones s.n. (NY, POM); near Apache Pass,
Chiricahua Mts, Sep 1881, Lemmon 331 (F, Uc [2]); Huachuca Mts,
Aug 1882, Lemmon 2795 (F, G, NY, US); 4% mi S of Sonoita &
Parker Canyon rds, 10 Oct 1965, Reese 94 (ARIZ); Bear Canyon,
Huachuca Mts, 28 Aug 1936, Shreve 7709 (ARIZ); near Ft Huachuca,
Aug 1894, Wilcox 335 (us); Ft. Huachuca, Aug 1893, Wilcox s.n.
(NY); PIMA CO, Oak Tree Canyon, Thurber Ranch, Santa Rita
Mts, 14 Oct 1940, Benson 10598 (ARIZ); Madera Canyon, Santa
Rita Mts, 6 Sep 1944, Clark 12377 (ARIZ); 17 mi S of Tucson-
Benson hwy on rte 83, 21 Oct 1944, Gould 2883 (ARIZ); Thurber
Ranch, Santa Rita Mts, 14 Sep 1945, Gould & Haskell 3268 (ARIZ,
DS, GH, NY, UC, US); Santa Rita Mts, 24 Aug 1903, Jomes s.n.
(DS, POM); Madera Canyon, Santa Rita Mts, 24 Aug 1926, Loomis
& King 2890 (ARIZ); Santa Rita Mts, 16 Sep 1884, Pringle 55 (GH,
SMU); Greaterville, 16 Sep 1916, Shreve 4965 (ARIZ) ; Empire Ranch,
26 Aug 1936, Shreve 7685 (ARIZ, SMU); Santa Rita Mts, Rozemont,
20 Sep 1915, Thornber 7404 (ARIZ), 10 Sep 1915, Thornber 8121
(ARIZ), 20 Sep 1915, Thornber 9050 (ARIZ); SANTA CRUZ CO., Syca-
more Creek, ca 28 mi NW of Nogales, 20 Sep 1964, Cronquist 10003
(NY); Sycamore Canyon, near Ruby, 30 Sep 1944, Darrow & Haskell
2012 (ARIZ, NY, UC, US); near Canoa Tank, Summit Motorway, on
58 Rhodora [Vol. 74
Nogales-Ruby rd, 14 Oct 1944, Darrow & Haskell 2256 (ARIZ);
Patagonia, 14 Sep 1959, Goodding 248-59 (ARIZ); Sonoita Creek,
30-31 Aug [year?], Harrison 8181 (F); Nogales, 30 Aug 1931,
Harrison & Fulton 8153 (ARIZ, Us); Sonoita, 2 Sep 1928, Harrison
& Kearney 5702 (us); Patagonia Mts, 15 Sep 1934, Kearney &
Peebles 10052 (uc); Sycamore Canyon, SW of Atascosa Mts & SE
of Ruby, 3 Oct 1951, Parker 7707 (ARIZ, F, RSA, UC, US); Patagonia
Mts (Nogales), 28 Aug 1927, Peebles & Harrison 4680 (ARIZ, F);
Harshaw, 22 Aug 1932, Shreve 6003 (ARIZ, F).
13. Melampodium nayaritense Stuessy, Brittonia 22:113.
f. 5. 1970. TYPE: MEXICO: Nayarit, Canon de Jesús María,
1000 ft, 29 Aug 1905, P. Goldsmith 133 (Holotype, NY!;
isotypes, F! GH! MO! Uc! US!).
Annual herbs, 10-30 cm tall. Stems ascending to erect,
1-2 mm diam, hirtellous with hairs 0.5 mm long. Peduncles
1.5-3.6 em long. Leaves sessile, linear and entire to deeply
pinnately divided, 3-4.5 cm long, 0.2-3.7 em wide, at apex
acute, at base attenuate to obtuse, with upper surface finely
strigose with hairs 1 mm long, with lower surface sericeous ;
margin at base pilose-hispid with hairs up to 1.5 mm long.
Heads 5-6 mm tall, 8-11 mm diam. Outer involucre cupu-
late, 7-10 mm diam; bracts 5, slightly connate at base, sep-
arate, obovate-rhombie, 1.5-5.5 mm long, 3-3.5 mm wide, at
apex acute-acuminate, with abaxial surface pilose with
hairs 1 mm long; margin herbaceous. Fruits 3 mm long,
with lateral surfaces ribbed and smooth to markedly tuber-
culate; hood apex cirrhous, with tapering appendage up
to 2 mm long. Ray florets 5-6; ligules yellow, ovate-orbicu-
late, 2 mm long, 2 mm wide. Disc florets 15-25; corollas
yellow, 1.2 mm diam, with throat 0.8 mm and tube 0.6 mm
long. Paleae broadly oblanceolate, 3.5 mm long, 1.6 mm
wide; apex yellow, with margin entire and undulate; mid-
rib prominent, weakly pubescent with hairs 0.3 mm long.
Chromosome number unknown.
Known from woodland hills in northeastern Nayarit, and
adjacent Durango, Mexico (Fig. 20), 180-300 m. Flowering
dates, Aug-Sep.
*In one head of an isotype collection (UC) are found both smooth
and roughly tuberculate fruits.
1972] Melampodium — Stuessy 59
Fig. 20. Map of Mexico showing distribution of Melampodium
bibracteatum (dots), M. nayaritense (squares), M. repens (circles),
M. rosei (closed triangles), and M. tenellum (open triangles). One
collection of M. bibracteatum from Guatemala not shown.
REPRESENTATIVE SPECIMENS. Mexico. DURANGO: Huasemota, 14 Aug
1897, Rose 3476 (US). NAYARIT: Valley of Río Jesús María, near
village of Jesus María, 17 Sep 1960, Feddema 1209 (MICH).
4. Series Cupulata Stuessy, ser. nov.
Herbae annuae vel perennae (M. sinuatwm); folia an-
guste ovata (saepe lobata), subtus strigosa; ligulae luteae
vel flavo-aurantiacae, plus quam vel raro circa 2 mm longae;
involucrum extimum marginibus scariosis; fructi plerum-
que eucullati. Species 14-19. Typus: Melampodium cupula-
tum A. Gray.
60 Rhodora [Vol. 74
14. Melampodium cupulatum A. Gray, Proc. Amer. Acad.
Arts & Sci. 8:291. 1870. TYPE: MEXICO: Sonora, Rio Yaqui
[27937' N, 110°38’ W; McVaugh, 1956], Nov-Dec 1869, FE.
Palmer 20 (Holotype, GH!; isotypes, NY! US[2]!; photo-
graph of US isotype, TEX!).
Eclipta pusilla M. E. Jones, Contrib. West. Bot. 18:70.
1933. TYPE: MEXICO: Baja California, Loreto, Arroyo
Undo Ranch, 26 Oct 1930, M. E. Jones 27739 (Holotype,
POM!).
Annual herbs, 10-35 cm tall. Stems erect, 1-4 mm diam,
moderately puberulent to hispidulous with hairs 0.1-0.3 mm
long. Peduncles 3.5-8 em long. Leaves sessile or with short
petioles 2-3 mm long, ovate-lanceolate, 3.3-7.5 cm long, 0.8-
2.1 em wide, at apex acute, at base attenuate, with both
surfaces strigillose with hairs 0.2-0.8 mm long; margin en-
tire to broadly crenate or sinuate on upper 3/4 of blade.
Heads 7-8 mm tall, 9-17 mm diam. Outer involucre cupu-
late, 6.5-10 mm diam; bracts 5, connate 1/2 their length,
ovate to narrowly ovate, 4-6 mm long, 2.7-3.3 mm wide, at
apex acute-acuminate, with abaxial surface strigillose with
hairs 0.2-0.8 mm long; margin scarious. Fruits 3-3.4 mm
long, with lateral surfaces with small tubercles and reticula-
tions." Ray florets 8-12; ligules yellow-orange, oblong-el-
liptie, 3-6 mm long, 1-3 mm wide. Disc florets 40-60;
corollas yellow-orange, 2 mm diam, with throat 0.7 mm and
tube 0.3 mm long. Paleae oblanceolate, 3.5 mm long, 1 mm
wide; apex yellow, with margin erose; midrib prominent,
puberulous with hairs 0.1 mm long. Chromosome number,
n — 10.
Arid regions of Baja California, Sinaloa, Sonora, and
Nayarit, Mexico (Fig. 19), 30-790 m. Flowering dates, Sep-
Mar.
REPRESENTATIVE SPECIMENS. Mexico. BAJA CALIFORNIA: vicinity of
La Cumbre de Alta Gracia, 1 Nov 1964, Carter 4880 (Ds, UC); Mesa
del Potrero de San Javier, 19 Sep 1965, Carter 4964 (Ds, UC); Cerro
Gabilán, S of Portezuelo de Gabilán, 30 Sep 1965, Carter 5055 (UC);
"Although hoods are usually rudimentary in this species, a few
coiled awns on small hoods are found in Feddema 1219 from Nayarit.
1972] Melampodium — Stuessy 61
NE end of Valle de Los Encinos (S side of Cerro Giganta), 28 Sep
1967, Carter & Moran 5314 (uc); 2 mi E of Comondü on rd to
Loreto, 28 Aug 1955, Chambers 965 (pns, UC); N of Comondú, 3 Oct
1941, Hammerly 182 (ns, POM); 8 mi S of Querétaro, 29 Nov 1946,
Wiggins 11507 (Ds, GH, UC, US). NAYARIT: near Jesús Maria, 18 Sep
1960, Feddema 1219 (MICH). SINALOA: 5 mi W of Culiacán on rd
to Altata, 28 Jan 1964, Flyr 112 (TEX); Cerro Tecomate, W of
Pericos, 28 Feb 1940, Gentry 5750 (ARIZ, DS, GH, NY, UC); Maratón,
12 mi W of Culiacán, 9 Mar 1944, Gentry 7005 (DS, F, GH, NY, UC,
US); near Culiacán, Oct [year?], Schaffner s.n. (GH). SONORA: near
rr station, Masiaca, 13 Sep 1930, Abrams 12811 (Ds); Ciudad Obregón,
29 Sep 1933, Gentry 290 (Ds); Alamos, 26 Mar-8 Apr 1890, Palmer
726 (GH); 4 mi S of La Puerca, SW of Hermosillo toward Tastiota,
4 Sep 1941, Wiggins & Rollins 281 (DS, GH, NY, UC, US).
15. Melampodium appendiculatum Robins. Proc. Amer.
Acad. Arts & Sci. 36:457. 1901. TYPE: MEXICO: Chihua-
hua, near Frailes [ca 12 km NE of Guachochic; McVaugh,
1956], ca 7400 ft, Oct 1885, E. Palmer 245 (Holotype, 3H !;
isotype, US!; photograph of US isotype, TEX!).
Melampodium appendiculatum Robins. var. leiocarpum
Robins. Proc. Amer. Acad. Arts & Sci. 36:457. 1901. TYPE:
MEXICO: Sonora, Alamos, 16-30 Sep 1890, E. Palmer '126
(not 726, 26 Mar-8 Apr 1890) (Holotype, GH!; isotypes,
NY! US!).
Melampodium appendiculatum Robins. var. sonorense
Robins. Proc. Amer. Acad. Arts & Sci. 36:457. 1901. TYPE:
MEXICO: Sonora, Cochuto, 5100 ft, 2 Oct 1890, C. V. Hart-
man 71 (Holotype, GH!).
Annual herbs, 10-40 em tall. Stems erect, 1-3 mm diam,
subglabrous at base to hispid or pilose above with hairs
0.5-1.5 mm long. Peduncles 1-8 cm long. Leaves sessile,
linear to lanceolate, 2.5-8.3 cm long, 0.4-1.8 em wide, at
apex acute, at base subauriculate, with both surfaces stri-
gose-hispid with hairs 1-1.5 mm long; margin entire to
obsoletely crenate, at base hispid with hairs up to 2 mm
long. Heads 5-8 mm tall, 7-17 mm diam. Outer involucre
strongly cupulate, 4.5-8 mm diam; bracts 5, connate 2/3
their length, ovate, 3.5-4.5 mm long, 2-3 mm wide, at apex
acute, with abaxial surface pubescent-pilose with hairs 0.5
62 Rhodora [Vol. 74
mm long (longer near base) ; margin scarious. Fruits 2.3-
2.8 mm long, with lateral surfaces smooth and glabrous to
very tuberculate and striate; hood apex cirrhous (rarely
muticous or mucronate) ?, with appendage up to 4 mm long.
Ray florets 8-13; ligules yellow-orange, oblong-elliptic, 2-8
mm long, 1-2.9 mm wide. Disc florets 25-40; corollas yellow,
1.3 mm diam, with throat 0.7 mm and tube 0.4 mm long.
Paleae oblancolate, 3.2 mm long, 1 mm wide; apex yellow,
with margin dentate-laciniate; midrib prominent, puberu-
lent with hairs 0.1 mm long. Chromosome number un-
known.
Pine-oak forests in mountains of Sonora, Chihuahua and
northwestern Durango, Mexico (Fig. 19), 790-1390 m.
Flowering dates, Jul-Nov.
Robinson (1901) described two new varieties within this
species based on inner involucral bract differences. From
an examination of specimens that indicate intra- and inter-
populational variability, however, it seems clear that these
described varieties represent simply forms within the mor-
phological range of one species.
REPRESENTATIVE SPECIMENS. Mexico. CHIHUAHUA: Guasaremos,
Río Mayo, 5 Aug 1935, Gentry 1570 (ARIZ, F, GH, NY, UC, US), 3 Aug
1936, Gentry 2353 (ARIZ, F, GH); Rio Aros, 23 Jul 1937, LeSueur
1466 (TEX), 28 Jul 1937, LeSueur 1472 (F); Sierra Madre Mts near
Seven Star Mine, 27 Aug 1899, Townsend & Barber 376 (F, GH, NY,
POM, UC, US [2]). DURANGO?: Lodiego [ca 25° N, 106°45' W; Mc-
Vaugh, 1956], 9-15 Oct 1891, Palmer 1609 in part (F, GH, NY, US).
SONORA: 20 mi NE of Ures, 16 Nov 1939, Drouet, Richards & Lock-
hart 3609 (F); Baviácora, W of Rio de Sonora, 17 Nov 1989, Drouet,
Richards & Lockhart 3624 (F); Alamos, Quirocoba, 13 Nov 1933,
Gentry 790M [790] (ARIZ, DS); San Bernardo, Río Mayo, 14 Jul
1935, Gentry 1492 (F, Gu) ; 6 mi S of Nogales, 7 Aug 1965, R. & J.
Matthews 480 (TEX); San Bernardino Ranch, 22 Aug 1893, Mearns
1997 (US); ca 12 mi from Cananea at km 20 on rd to Bacanuchi,
25 Aug 1963, Moreno MS-198 (ARIZ); Nácori Chico, Pie de la Cuesta,
6 Oct 1939, Muller 3659 (F, UC) ; Aduana, 10 Aug 1930, Russell &
Souviron 2 (Us); Sta Magdalena, Nov 1855, Schott 72 (F, GH, NY),
"In the collection Shreve 6758 (US), most flowering heads have
fruits with long cirrhous appendages, but in one head all the achenes
are muticous or only very slightly mucronate,
1972] Melampodium — Stuessy 63
Schott s.n. (F); 21 mi SE of Moctezuma, 25 Sep 1934, Shreve 6758
(ARIZ, US); Arispe, 18 Aug 1958, Turner 192 & Lowe 2074 (ARIZ);
91 mi S of Nogales along Rio de los Alisos, 8 Sep 1934, Wiggins
7021 (Ds, TEX [2], US); ca 5 mi S of Babiácora along Sonora River,
21 Sep 1934, Wiggins 7388 (ARIZ, ps, TEX [2], US); 21 mi S of
Moctezuma, 25 Sep 1934, Wiggins 7444 (ARIZ, DS, TEX, US); 8 mi
S of Nogales & 2 mi W of rr, 3 Oct 1934, Wiggins 7533 (Ds, Us);
8 mi from Matape toward Batuc, 9 Sep 1941, Wiggins & Rollins 433
(ARIZ, DS, GH, NY); Puerto de Huépari, NW of Aribabi, 7 Sep 1939,
White 2784 (ARIZ, GH, US); Cañón de la Gallina, 25 Aug 1940, White
3516 (ARIZ, GH); El Rancho de la Nacha, 25 mi W of La Angostura,
14-20 Aug 1941, White 3908 (GH); Cañón de la Palomita, N of El
Tigre, 29 Aug 1941, White 4153 (GH).
16. Melampodium sinuatum Brandg. Proc. Calif. Acad. Sci.
ser. 2. 3:144. 1891. TYPE: MEXICO: Baja California, San
José del Cabo, 16 Sep 1890, T. S. Brandegee 302 (not 302,
Sep 1891; not 302, 10 Mar 1892) (Lectotype chosen, UC!).
Suffruticose perennials, 8-35 cm tall. Stems erect, 1-2
mm diam, puberulent-canescent with hairs 0.2-0.5 mm leng.
Peduncles 3-9 em long. Leaves sessile, linear to elliptic,
2-4.5 cm long, 0.5-1.6 cm wide, at apex obtuse, at base
subauriculate, with both surfaces densely strigillose with
hairs 0.2-0.3 mm long; margin sinuate-undulate (rarely
repand) to irregularly lobed. Heads 6-7 mm tall, 10-20 mm
diam. Outer involucre cupulate, 6-9.5 mm diam; bracts 5,
connate 3/4 their length, orbiculate, 3-4.5 mm long, 2.5-4.5
mm wide, at apex obtuse, with abaxial surface strigillose-
tomentose with hairs 0.4-0.7 mm long; margin slightly
scarious. Fruits 2.9-3.2 mm long, with lateral surfaces
tuberculate-striate or reticulate. Ray florets 9-13; ligules
yellow-orange, elliptic-linear, 4-9 mm long, 1.5-3 mm wide.
Disc florets 50-80; corollas yellow-orange, 1.3 mm diam,
with throat 1 mm and tube 0.5 mm long. Paleae broadly
oblanceolate, 3.5 mm long, 0.8 mm wide; apex yellow, with
margin erose-laciniate; midrib prominent, puberulent with
hairs 0.1 mm long. Chromosome number unknown.
Low granitic hills near San José del Cabo, Baja Califor-
nia, Mexico (Fig. 19), ca 60 m. Flowering dates, Sep-Mar.
64 Rhodora [Vol. 74
M. sinuatum is one of the three species in the genus,
along with M. cupulatum and M. divaricatum that are
found in Baja California. The distributional and morpho-
logical similarity of the former two taxa, and the restricted
distribution of M. sinuatum at the tip of Baja California
suggest that this species may have been derived from a
mainland ancestral population of M. cupulatum.
REPRESENTATIVE SPECIMENS. Mexico. BAJA CALIFORNIA: San José
del Cabo, Sep 1891, Brandegee 302 (GH, US); Sierra de la Trinidad,
Nov 1902, Brandegee s.n. (vc [2], US); 12.2 km SW José del Cabo,
16 Dec 1947, Carter, Alexander & Kellogg 2216 (DS, UC, us); San
José del Cabo, Mar 1901, Purpus 416 (uc, US) ; b mi W of San José
del Cabo, 17 Dec 1958, Wiggins 14377 (DS, GH, TEX, UC).
17. Melampodium rosei Robins. Proc. Amer. Acad. Arts &
Sci. 36:461. 1901. TYPE: MEXICO: Sinaloa, between Rosa-
rio & Concepcion, 28 Jul 1897, J. N. Rose 3271 (Holotype,
us!; photograph of holotype, TEX!).
Melampodium arenicola Robins. Proc. Amer. Acad. Arts
& Sci. 36:457. 1901. TYPE: MEXICO: Sinaloa, Mazatlan, Isla
Piedra, 31 Dec 1894, F. H. Lamb 361a (Holotype, GH!; iso-
types, G[2] ! Mo! NY! U$!; photograph of US isotype, TEX !).
Melampodium rosei Robins. var. subintegrum Robins.
Proc. Amer. Acad. Arts & Sci. 36:462. 1901. TYPE:
MEXICO: Sinaloa, Rosario, 7 Jul 1897, J. N. Rose 1568
(Holotype, US!; isotypes, F! GH!; photograph of holotype,
TEX!; photograph of K isotype, US!).
Annual herbs, 10-50 cm tall. Stems erect to decumbent,
1-3 mm diam, glabrous near base to hispidulous above with
hairs up to 0.5 mm long. Peduncles 2-5 em long. Leaves
sessile, lanceolate to oblong-ovate, 3-5.5 em long, 0.5-2 cm
wide, at apex obtuse to acute, at base subauriculate (rarely
obtuse), with both surfaces strigose with hairs 0.3-1 mm
long; margin subentire to strongly lobed. Heads 4-6 mm
tall, 7-12 mm diam. Outer involucre cupulate, 5-8 mm
diam; bracts 5, connate 1/4 or less their length at base,
separate to imbricate 3/4 their length, orbiculate to ovate,
2.5.3.8 mm long, 1.5-3.5 mm wide, at apex broadly rounded
to acute, with abaxial surface subglabrous to strigillose
1972] Melampodium — Stuessy 65
with hairs 0.3 mm long (longer near base) ; margin scari-
ous. Fruits 1.1-2.3 mm long, with lateral surfaces smooth
to striate (sometimes very tuberculate); hood apex mu-
ticous or less often mucronate to cirrhous, with appendage
up to 3 mm long. Ray florets 8-9; ligules yellow-orange, ob-
long-elliptic, 3-6 mm long, 1-2.3 mm wide. Disc florets 40-
60; corollas yellow-orange, 1.6 mm diam, with throat 0.7
mm and tube 0.5 mm long. Paleae broadly oblanceolate,
2.7 mm long, 0.8 mm wide; apex yellow, with margin lacini-
ate; midrib prominent, weakly puberulent with hairs 0.2
mm long. Chromosome number, n = 10.
Tropical deciduous and thorn forests in Sinaloa and ad-
jacent Durango, Mexico (Fig. 20), 0-400 m. Flowering
dates, Aug-Jan.
Some of the specimens on Isla Piedra off the coast of
Mazatlán (type locality of M. arenicola) possess more lobed
leaves, a tendency toward decumbent stems, larger heads,
and narrower and longer ligules than is typical for the
species. Because these forms exhibit considerable char-
acter intergradations and are found also intermixed in at
least one population on the mainland (personal observa-
tions), they are not here formally recognized. With M.
arenicola and M. rosei regarded as being conspecific and
because both names were published at the same time, one
of the epithets must be selected for use. M. rosei has been
chosen because this name has been applied almost exclu-
sively in identifications of previous collections of this taxon.
Collections from Ymala (Imala), Sinaloa, at the north-
west portion of the range of this species morphologically
approach M. cupulatum, a species found nearby (Fig. 19).
Hybridization between the two taxa cannot be exluded.
REPRESENTATIVE SPECIMENS. Mexico. DURANGO: La Bajada, Tama-
zula, Nov 1921, Ortega 4446 (US). SINALOA: Mazatlán, 3 Nov 1893,
Brandegee s.n. (US) ; 2 mi SE of Mazatlán, 4 Aug 1938, Eyerdam &
Beetle 8687 (ARIZ, UC); Mazatlán, 15 Sep 1927, Ferris 5020 (ps);
vicinity of Labradas, 18 Sep 1925, Ferris & Mexia 5078 (ps, GH);
near Mazatlán, 29 Jan 1964, Flyr 138 (TEX); ca 2 mi N of Escuinapa,
13 Aug 1960, King 3707 (Ds, NY, TEX, UC, US); ca 12 mi N of Escui-
napa, 13 Aug 1960, King 3709 (DS, NY, RSA, TEX, UC); ca 13 mi N
66 Rhodora [Vol. 74
of Rosario, 13 Aug 1960, King 8710 (Ds, NY, TEX, UC, US); ca 21 mi
N of Rosario, 13 Aug 1960, King 8712 (DS, NY, TEX, UC, US); ca
3 mi S of Mazatlan, 13 Aug 1960, King 3715 (DS, NY, TEX, UC, US) ;
ca 10 mi NE of jet rtes 15 & 40, 14 Aug 1960, King 3716 (DS, NY,
TEX, UC); Mazatlán, on Isla Piedra, 31 Dec 1894, Lamb s.n. (DS, UC,
US); Mazatlán, 2 Jan 1895, Lamb s.n. (DS, GH, NY, UC, US); San
Agustín, 1921, Ortega 4008 (Us) ; Escuinapa, 1926, Ortega 6125 (us),
1933, Ortega 7004 (F); Ymala [Imala], [25°42’ N, 107^15' W; Mc-
Vaugh, 1956], 16-25 Aug 1891, Palmer 1457 (F, GH, US), 25 Sep-8 Oct
1891, Palmer 1757a (ARIZ, GH, US); 10 mi E of jet rtes 15 & 40,
26 Aug 1961, Powell & Edmondson 908 (TEX); near Colomas, 19 Jul
1897, Rose s.n. (US); between Mazatlán & Villa Unión, 14 Dec 1936,
Shreve 7821 (ARIZ, F); 4.5 mi N of Escuinapa, 4 Sep 1965, Stuessy
404 (TEX); Isla Piedra, Mazatlán, 28 Aug 1966, Stuessy 748 (TEX);
4 mi S of Mazatlán, 28 Aug 1966, Stuessy 750 (TEX); 12-15 km SE
of Mazatlán, 4 Aug 1938, Worth & Morrison 8807 (vc, US) ; Mazatlán
& vicinity, Dec 1888, Wright 1213 (ps, F, UC, US).
18. Melampodium tenellum Hook. & Arn. Bot. Beech. Voy.
299. 1838. TYPE: MEXICO: Nayarit, Tepic, 1825-28, G. T.
Lay & A. Collie et al. s.n. (Holotype, K!; photograph of
holotype, F! MICH! OS! TEX! US!; photograph and fragment
of holotype, US!).
Annual herbs, 6-30 cm tall. Stems erect (rarely decum-
bent), 1-2 mm diam, subglabrous to hispidulous with hairs
0.3 mm long. Peduncles 3-7 cm long. Leaves sessile or with
petioles 2-10 mm long, lanceolate to narrowly ovate, 2-5.5
cm long, 0.5-1.4 em wide, at apex obtuse (less often acute),
at base attenuate (rarely obtuse), with both surfaces scat-
teredly strigose with hairs 0.3 mm long; margin subentire
to shallowly lobed. Heads 3.5-5.5 mm tall, 6-14 mm diam.
Outer involucre cupulate, 3.5-7 mm diam; bracts 5, connate
1/4-1/2 their length, separate to imbricate, orbiculate to
ovate, 2.5-3.5 mm long, 2.5-3 mm wide, at apex acute to
rounded, with abaxial surface strigillose-hispidulous with
hairs 0.8 mm long; margin scarious. Fruits 1.8-2.6 mm
long, with lateral surfaces usually with 4 longitudinal rows
of tubercles (rarely smooth); hood apex muticous. Ray
florets 8-9; ligules yellow-orange, oblong-elliptic, 2-5 mm
long, 1-2.8 mm wide. Disc florets 40-60; corollas yellow-
orange, 1.3 mm diam, with throat 0.8 mm and tube 0.5 mm
1972] Melampodium — Stuessy 67
long. Paleae oblanceolate, 2.5 mm long, 0.7 mm wide; apex
yellow, with margin laciniate; midrib prominent, glabrous.
Chromosome number, n — 10.
Tropical deciduous forests and savannas in Nayarit, Jalis-
co, and Michoacán, Mexico (Fig. 20), 30-920 m. Flowering
dates, Jul-Nov.
M. tenellum is morphologically similar to M. rosei, but
each species does possess unique morphological and geo-
graphic features. As in M. rosei, however, M. tenellum
also has considerable variation in vegetative and outer phyl-
lary merphology. Much additional work will be needed
before this intraspecific variation can be understood satis-
factorily.
REPRESENTATIVE SPECIMENS. Mexico. JALISCO: Puerto Vallarta, 20
Jul 1932, Howell 10303 (US). MiCHOACÁN: 24 km S of Arteaga,
29 Nov 1968, Rzedowski 26618 (os). NAYARIT: E of Tepic along
Río Tepic, 22-24 Aug 1935, Pennell s.n. (GH); Santiago Ixc., Mezcal-
titan, Jan 1926, González 6125 (ps, US); ca 38 mi S of Nayarit-
Sinaloa border on rte 15, 12 Aug 1960, King 3703 (Ds, NY, TEX, UC,
US); ca 28 mi S of Nayarit-Sinaloa border on rte 15, 12 Aug 1960,
King 3704 (DS, NY, RSA, TEX, UC); ca 21 mi S of Nayarit-Sinaloa
border on rte 15, 13 Aug 1960, King 3705 (DS, NY, TEX, UC, US);
ca 1 mi S of Nayarit-Sinaloa border on rte 15, 13 Aug 1960, King
9706 (DS, NY, RSA, TEX, UC); 27.9 mi SE of Nayarit-Sinaloa border
on rte 15, 4 Sep 1965, Stuessy 401 (TEX); 10 mi NW of jct rte 15
& rd to Tuxpán, 27 Aug 1966, Stuessy 745 (TEX).
19. Melampodium glabribracteatum Stuessy, Brittonia 22:
112. f. 3. 1970. TYPE: MEXICO: Oaxaca, Cerro de San
Antonio, Ocotlán, 1600 m, 29 Aug 1936, C. Conzatti 5169
(Holotype, K!; photograph of holotype, os! TEX!).
Annual herbs, 15-22 cm tall. Stems erect to decumbent,
1-1.5 mm diam, subglabrous at base to pilose above with
hairs up to 1 mm long. Peduncles 1.3-1.7 cm long. Leaves
sessile, blades lanceolate, 2-3.3 em long, 0.4-0.7 cm wide,
at apex obtuse, at base obtuse-subauriculate, with both sur-
faces sparingly strigose with hairs up to 1 mm long; margin
entire or rarely 2-lobed, near base pilose with hairs up to
1 mm long. Heads 5-6 mm tall, 6-9 mm diam. Outer in-
volucre cupulate, 7-8 mm diam; bracts 5, slightly connate
58 Rhodora [Vol. 74
at base, imbricate 3/4 their length, elliptic-orbiculate, 3-4
mm long, 2.4-3.2 mm wide, at apex rounded, with abaxial
surface glabrous; margin narrowly scarious, ciliate with
hairs up to 0.6 mm long. Fruits 2.1-2.5 mm long, with
laterai surfaces ribbed and tuberculate-aculeate; hood apex
cirrhous, with tapering appendage up to 2 mm long. Ray
florets 6-8; ligules yellow, ovate-oblong, 3.5 mm long, 2-2.5
mm wide. Disc florets ca 30; corollas yellow, 1 mm diam,
with throat 1 mm and tube 0.7 mm long. Paleae oblance-
olate, 3.3 mm long, 0.8 mm wide; apex yellow, with margin
smooth-erose; midrib prominent, glabrous. Chromosome
number unknown.
Known only from the type collection from pine-oak for-
ests near Ocotlán, Oaxaca (Fig. 11), 1600 m. Flowering
date, Aug.
5. Series Longipila Stuessy, ser. nov.
Herbae annuae; folia late ovata, subtus sericea; ligulae
flavae-aurantiacae, plus quam 2 mm longae; involucrum
extimum marginibus herbaceis; fructi apice arista adaxiali
complanato cirrhoso. Species 20. Typus: Melampodium
longipilum Robins.
20. Melampodium longipilum Robins. Proc. Amer. Acad.
Arts & Sci. 27:173. 1892. TYPE: MEXICO: San Luis Potosi,
San José Pass, 11 Jul 1890, C. G. Pringle 3639 (Holotype,
GH!).
Melampodium villicaule Greenm. Field Col. Mus. Pub.
Bot. 2:345. 1912. TYPE: MEXICO: Guerrero, Rio Baisas,
26 Aug 1910, C. R. Orcutt 4386 (Holotype, F!).
Annual herbs, 7-30 cm tall. Stems erect, 0.7-4 mm diam,
villous with hairs 2 mm long. Peduncles 2-11.5 cm long.
Leaves with short and broad petioles (rarely sessile) 3-6
mm long, with blades ovate to elliptic, 2-7.5 cm long, 0.5-3
em wide, at apex acute, at base attenuate to partially sub-
auriculate, with both surfaces pilose with hairs 1 mm long;
margin entire. Heads 5-7.5 mm tall, 7-12 mm diam. Outer
involucre markedly cupulate, 6.5-9.5 mm diam; bracts 5,
1972] Melampodium — Stuessy 69
slightly connate at base, imbricate 1/2 their length, ovate,
3.5-5 mm long, 2-3.5 mm wide, at apex acute, with abaxial
surface pilose with hairs 1 mm long; margin herbaceous.
Fruits 2.2-2.8 mm long, with lateral surfaces tuberculate or
striate with tapering cirrhous appendages up to 8 mm long.
Ray florets 7-8; ligules yellow-orange, ovate-elliptic, 3.5-5.5
mm long, 1.8-4 mm wide. Disc florets 40-70; corollas yel-
low-orange, 1.2 mm diam, with throat 0.8 mm and tube
0.5 mm long. Paleae oblanceolate, 3.5 mm long, 0.8 mm
wide; apex yellow, with margin laciniate-dentate; midrib
prominent, pubescent with hairs 0.5 mm long. Chromosome
number, n = 10."
Scattered in tropical deciduous and pine-oak forests of
western Guatemala and the Mexican states of San Luis
Potosi, Hidalgo, Puebla, Guerrero, and Oaxaca (Fig. 12),
1870-3050 m. Flowering dates, Jul-Nov.
In my opinion this species is morphologically closest to
M. tenellum, although Turner and King (1962) have sug-
gested that its relationship is with M. divaricatum (in sec-
tion Serratura). 'The unusual adaxially flattened appendage
(Fig. 8) makes M. longipilum unique within the genus, and
this feature along with other characters such as large ovate
leaves and markedly cupulate outer involucres suggests
separation of the species as a monotypic series. Considered
somewhat rare before the present treatment, M. longipilum
is quite common especially throughout Oaxaca.
REPRESENTATIVE SPECIMENS. Guatemala. HUEHUETENANGO: Cumbre
Papal between Cuilco & Ixmoquí, 19 Aug 1942, Steyermark 50916
(F, NY). Mexico. GUERRERO: 19 mi N of Chilpancingo, 29 Aug 1965,
Stuessy 374 (TEX). HIDALGO: Metztitlán, SE of Metztitlán, 24 Nov
1942, Moore 2283 (GH). OAXACA: ca 64 mi SE of Oaxaca, 23 Jul
1960, King 3461 (DS, NY, TEX, UC, US); 13 mi NW of Tehuantepec,
23 Aug 1965, Stuessy 328 (TEX); 3.8 mi NW of Huajuapán de León,
25 Aug 1965, Stuessy 344 (TEX) ; 10 mi NW of Tehuantepec, 11 Aug
1966, Stuessy 634 (os, TEX); 51 mi NW of Tehuantepec, 11 Aug
1966, Stuessy 636 (TEX); 87 mi NW of Tehuantepec, 11 Aug 1966,
Stuessy 637 (TEX); 28 mi S of Zimatlán, 12 Aug 1966, Stuessy 639
(TEX). PUEBLA: Tehuacán, Sep 1911, Purpus 5619 (GH, NY, UC); near
"See Stuessy (1971b) for a discussion of the n — 11 count for
this species.
70 Rhodora [Vol. 74
Tehuacán, 36 Aug-8 Sep 1905, J. N. & J. S. Rose & Painter 10157
(US); Asunción de Chila, 25 Aug 1965, Stuessy 346 (TEX); 32 mi
NW of Huajuapán de León, 25 Aug 1965, Stuessy 347 (TEX); 2 mi
NW of Tehuitzingo, 16 Aug 1966, Stwessy 667 (TEX). SAN LUIS
POTOSÍ: Villar, 14 Sep 1893, Pringle 4537 (F, G, NY, UC, US [2]).
(To be continued)
IN MEMORIAM
CHARLES SCHWEINFURTH, 1890-1970
November 16, 1970 marks the end of an outstanding era
in the annals of American orchidology.
Charles Schweinfurth was born on April 13, 1890 in
Brookline, Massachusetts, in one of the oldest and most
71
72 Rhodora [Vol. 74
historic regions of the United States. During his formative
years he attended public schools in Brookline where he
quickly developed an intense devotion to various aspects of
natural history. While his interest in bird watching and
plant collecting started as an avocation, they became the
driving force for his whole life soon after graduation from
college.
He entered Harvard University in 1909 and majored in
chemistry, graduating in 1913 with a degree of A.B., cum
laude. 'This was a remarkable achievement, for in his sopho-
more year he succumbed to polio which paralyzed his right
arm. To regain the use of his arm, with a self discipline
that eharacterized his whole life, he soon started daily ex-
ercises, aided by his devoted parents. He maintained this
discipline almost to his last days. When he was informed
by his doctors that he must abandon chemistry, he found
a life-long career in his avocation.
Charles’ commitment to botany became officially regis-
tered on April 5, 1912 when he was elected a member of
the New England Botanical Club. He faithfully supported,
as well as promoted, its interest, often beyond the call of
duty. It was, indeed, a rare occurrence when Charles would
succumb to some more compelling business than the attend-
ance of the club’s monthly meetings. It was in this same
spirit that he accepted the burdensome responsibility of
corresponding secretary in 1949, a post he held in the club
until 1965, when he had to resign from it due to his doctor’s
order. His memberships included, among others, also the
American Association for the Advancement of Science, the
International Association for Plant Taxonomy, the National
Audubon Society, the Appalachian Mountain Club, and the
Boston Museum of Science.
Charles had many avocations in addition to his ardent
bird watching. During the summer months he enjoyed
traveling, since he was greatly interested in the classical
aspects of our cultural past. Consequently photography
became a second nature to him in recording precious im-
pressions, especially of paintings, sculptures and architec-
1972] Schweinfurth — Garay 73
tural designs for his leisure viewing at home or for sharing
them with others at some later time. During winter months
he spent much of his extra-curricular time on writing for
local newspapers, as well as in pursuit of winter sports,
notably ice skating.
The professional life for Charles Schweinfurth started
in 1914 when he accepted a position offered to him by Pro-
fessor Oakes Ames to tend the living orchid collection at
North Easton, Massachusetts. Within less than a year Pro-
fessor Ames, recognizing Charles’ great ability for remem-
bering scientific names and fine details, took him on as a
personal assistant to work in his private “Ames Botanical
Laboratory", also in North Easton.
It was then that his orchidaceous career started. From
January, 1915, Charles Schweinfurth was an orchidologist.
He worked intensively on the orchids of the Philippines,
those from Mt. Kinabalu, British North Borneo and various
Pacifie islands, followed by studies of the orchids of Hon-
duras, Costa Rica and Panama. In 1936, he co-authored
with Ames and Hubbard an extensive monographic study,
“The Genus Epidendrum in the United States and Middle
America".
These works alone would suffice to record him among out-
standing orchidologists. Charles, however, has built an even
more imposing monument for posterity. True to his good
nature and encouraged by Professor Ames, in 1922 he ac-
cepted an offer made by J. Francis Macbride to write up
the orchid family for his proposed “Flora of Peru". He
labored year after year and on April 9, 1958, the first of
the four parts of his monumental work appeared in pub-
lication. This work was the first descriptive treatise on
orchids of any Andean country.
Charles, like many of the outstanding botanists, while
working on a given project, concurrently undertook others.
He accepted the very onerous task of identifying the nu-
merous colored plates prepared under the supervision of
Celestino Mutis during La Real Expedición Botániea del
Nuevo Reino de Granada, between 1760 and 1817. The first
74 Rhodora Vol. 74
volume of this labor of love was published in 1963, while
the second volume appeared in 1969.
In 1958, Charles attended the 3rd South American Bo-
tanieal Congress in Lima, Peru, where he was awarded
Catedratico Honorario (Honorary Professor of Botany)
by Universidad Mayor de San Marcos, the oldest university
of the Americas. It was during that year, when he also
visitel Colombia, that the picture reproduced here was
taken by his colleague, Dr. Hernando Garcia-Barriga of
Universidad Nacional, Bogota, during a visit to the Paramo
de Chipaque.
Charles received another award of Catedratico Honorario
in 1962, from Universidad de Cuzco. In 1964, he was hon-
ored by the American Orchid Society with a Gold Medal
inscribed “For outstanding contributions to orchidology”,
and on July 12, 1966, he was elected Miembro Honorario
by Sociedad Colombiana de Orquideologia.
On November 29, 1965, the staff of the Botanical Museum
of Harvard University honored him on the occasion of his
Golden Jubilee in Orchidology with a scroll, inscribed as
follows: “Fifty years have passed since our colleague
Charles Schweinfurth initiated his productive career in
orchidology. Dean of the world’s orchidologists, like his
predecessors Lindley, Rolfe, Reichenbach, Schlechter,
Kraenzlin, Smith and Ames, he has made invaluable con-
tributions to our knowledge of the systematics of the orchids
of both the Old and the New World. His great number of
publications, his elucidation of the intricate structure of
orchid flowers, his augmentation to our knowledge of the
phytogeography of orchids, his acute powers of observation
for minute details, his loyalty and his dedication to his
chosen field stand as an inspiration to both present and
future students of orchidology. In grateful recognition of
these fifty years of service to orchidology, we of the Botan-
ical Museum of Harvard University present this scroll.”
The most meaningful event in Charles’ life, however, oc-
curred in December 1963, when, after a visit to the Holy
Land, he married the lifelong friend of the Schweinfurth
1972] Schweinfurth — Garay 75
family, Miss Maria Elizabeth Westergren of Stockholm,
Sweden. The blessings of this mature companionship he
openly cherished throughout the rest of his life.
The orchid world is vastly richer today for there was a
Charles Schweinfurth intimately associated with it. And
for those few who had the privilege of sharing his meaning-
ful friendship, not only the orchid world, but the whole
world was just a little better to live in.
LESLIE A. GARAY
ORCHID HERBARIUM OF OAKES AMES
BOTANICAL MUSEUM
HARVARD UNIVERSITY
NOMENCLATURAL CHANGES IN LESQUERELLA
REED C. ROLLINS AND ELIZABETH A. SHAW
During the last twenty years there has been a gradual
introduction into the International Code of Botanical No-
menclature of provisions for the automatic recognition of
tautonymous names and epithets when a taxon in a cate-
gory of infrafamilial rank (other than genus or species)
is segregated from one of the next higher rank. In the
Code adopted at Stockholm (Reg. Veg. 3: Arts. 34, 35.
1952) it was clearly stated that valid publication of the
name of a taxon of infraspecific rank which does not include
the type of the name of the taxon of the next higher rank
automaticaliy establishes in parallel the name of a second
taxon of the same subordinate rank, this having the same
type as does the name of the taxon of higher rank and the
same epithet. At Paris similar provisions were made for
categories of infrageneric rank (Reg. Veg. 8: Art. 22.
1956) and at Montreal for categories of infrafamilial rank
(Reg. Veg. 23: Art. 19. 1961). However it soon became
apparent that the very broad application of this useful
principle to the name of any taxon which includes the
nomenclatural type of the name of the taxon next higher
in rank can give rise to complications.
Although the International Code of Botanical Nomencla-
ture (Reg. Veg. 46: Arts. 2, 3. 1966) makes it clear that
every Recent plant belongs to a species, a genus, and a fam-
ily (as well as to an order, a class, and a division) it
nowhere provides that a plant must belong to a subspecies
or to a section, a series, or to a tribe. The use of such cate-
gories of intermediate, and optional, rank reflects only a
recognition of the possibility of using more hierarchical
steps within the framework of the required classification.
The sweeping extension of the principle of automatic rec-
ognition of tautonyms to names of taxa in such optional
ranks creates the possibility that a taxon with a particular
circumscription, position and rank might have, according
76
r
1972] Lesquerrella — Rollins and Shaw Ti
to the classification used, two correct names. Classification
and nomenclature are thus entangled in a way contrary to
Principle IV of the Code. Such distressing situations were
discussed by Wood & Webster (1968) and a series of care-
fully worded proposals intended to clarify matters was sub-
mitted to, and accepted by, the 11th International Botanical
Congress in 1969. In essence, these modifications restrict
the automatic recognition of tautonyms in the optional
ranks to names of only those taxa which do include the
type of the name of a species, a genus, or of a family. In
the other optional ranks, strict priority of publication ap-
plies.
Difficulties arising from the entanglement of classification
and nomenclature have come to light more often in dealing
with categories ranked between family and genus, or be-
tween genus and species, than with those below the rank
of species, and Wood & Webster were not entirely satisfied
with the example used with their proposal for modification
of Article 26, which deals with names of taxa of infra-
specific rank, in the then current edition of the Code. In
the course of our revision of the North American species
of Lesquerella (in press), we made certain decisions result-
ing in our recognition of two subspecies of Lesquerella
lasiocarpa (Hook. ex Gray) Watson, and, in that subspecies
not including the type of the name of the species, of two
varieties. We realized that one taxonomic decision in par-
ticular had produced precisely the situation Wood & Web-
ster had in mind when proposing the change in Article 26.
Since this is being used as an example in the forthcoming
edition of the Code, we wish to make the necessary nomen-
clatural changes here and to provide some explanation of
them. Our classification of L. lasiocarpa is summarized as
follows:
(1) Lesquerella lasiocarpa subsp. lasiocarpa
Vesicaria lasiocarpa Hook. ex Gray, Smithson. Con-
trib. Knowl. 5: 13. 1853. Type: Texas. between Bexar
& Trinity River, Berlandier in 1828 (K)
78 Rhodora [Vol. 74
(2) Lesquerella lasiocarpa subsp. berlandieri (Gray) Rol-
lins & Shaw, comb. nov.
Synthlipsis berlandieri Gray, Bot. Mex. Bound. Sur-
vey 34. 1859. Lectotype (here designated): Mexico.
Tamaulipas, Matamoros, Berlandier 3017 (GH), iso-
types (MO, US)
(22) L. lasiocarpa [subsp. berlandieri] var. hispida
(Wats.) Rollins & Shaw, comb. nov.
Synthlipsis berlandieri var. hispida Wats.,
Proc. Amer. Acad. 17: 321. 1882. Holotype:
locality uncertain, Palmer 26 (GH)
S. berlandieri Gray (see above)
L. lasiocarpa var. berlandieri (Gray) Payson,
Ann. Missouri Bot. Gard. 8: 189. 1922
L. lasiocarpa var. ampla, Rollins, Rhodora 57:
245. 1955. Holotype: Mexico. Tamaulipas,
vicinity of Victoria, Palmer 41 (GH), isotype
(NY)
(2b) L. lasiocarpa [subsp. berlandieri] var. hetero-
chroma (Wats.) Rollins, Rhodora 57: 245. 1955
Synthlipsis heterochroma Wats., Proc. Amer.
Acad. 17: 321. 1882
According to the provisions of the 1966 edition of the
Code, it would have been necessary to use the name L. lasio-
carpa, [subsp. berlandieri] var. berlandieri [without an au-
thor— not *(Gray) Payson"] for that variety (2a) of
subsp. berlandieri which includes the type of the name of
the subspecies, no matter how it might be circumscribed,
when recognizing taxonomically, as we do, this optional
category. Still following the 1966 edition of the Code, if
one chose nct to adopt the category “subspecies” in classify-
ing the infraspecific taxa of Lesquerella lasiocarpa, but
used only “varietas,” the correct name for this taxon, still
with the same circumscription, position, and rank, would
be L. lasiocarpa var. hispida (Wats.) Rollins & Shaw. This
is the earliest available epithet in the rank of variety for
a taxon which does not include the type of the name of the
taxon of next higher rank. Thus, depending upon the classi-
1972] Lesquerrella — Rollins and Shaw 79
fication used (whether subspecies or not in the present in-
stance), this one variety could have two correct names, a
situation contrary to Principle IV.
The crux of the matter is our decision to circumscribe the
variety which includes the type of the name L. lasiocarpa
subsp. berlandieri so as to include Synthlipsis berlandieri
var. hispida Watson. Article 26 as modified at Seattle now
provides that use of a tautonymous epithet without citation
of an author’s name applies only to the names of taxa of
infraspecific rank which do include the type of the name of
the species to which they are assigned. Since variety 2a
does not include the type of the name L. lasiocarpa, the
earliest available epithet in varietal rank, if one such exists,
must then be used for it, and in this case it is “hispida”. In
Payson’s classification, based upon different criteria, S. ber-
landieri var. hispida was not placed in the synonymy of S.
berlandieri, and he was thus quite correct in his use of the
name L. lasiocarpa var. berlandieri (Gray) Payson. How-
ever, according to the Code as modified at Seattle, the taxon,
as circumscribed by us, is to be called Lesquerella lasio-
carpa var. hispida (Wats.) Rollins & Shaw.
LITERATURE CITED
Woop, C. E. & G. L. WEBSTER. 1968. Tautonyms and Confusion in
tbe International Code. Taxon 17: 645-651.
GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS 02138
THE PARKER CLEAVELAND HERBARIUM OF
BOWDOIN COLLEGE
EDWARD HEHRE, A. R. HODGDON, AND R. B. PIKE!
The Herbarium of Bowdoin College, Brunswick, Maine,
which was assembled in the 19th century under the direc-
tion of Parker Cleaveland (1780-1858) was stored away in
the attic of Searle’s Science Building at Bowdoin College
for over half a century. Professor Cleaveland was a dis-
tinguished naturalist in the nineteenth century tradition.
He is best known for the collection of minerals housed for
many years in the *Cleaveland Cabinet" in Massachusetts
Hall at Bowdoin College. Cleaveland had studied under
Goethe who recommended him highly when he began his
career in Brunswick in 1805. That he had much knowledge
of botany and of botanists is suggested by the important
collections that came his way. However, the Cleaveland
Collection, like many herbaria in small colleges, ceased to
grow and became isolated and inaccessible after its aus-
picious beginning, and because of neglect, failed to fulfill
the purpose for which it had been assembled.
Through the kindness of the President and overseers of
Bowdoin College, the Cleaveland Herbarium is now on a
ten-year loan to the University of New Hampshire.
By means of support provided by Mr. Sumner Pike of
Lubec, Maine, the collection of approximately 9,400 speci-
mens was given a thorough and painstaking rehabilitation
by the senior author. Most specimens were in good condi-
tion with a minimum of damage by mold and insects but
nearly all required remounting and, for a great many, some
nomenclatural updating. The entire collection now is incor-
porated in the University of New Hampshire Herbarium
(NHA), each sheet being stamped with the Bowdoin Seal
for ready recognition.
"Published with the approval of the Director of the New Hampshire
Agricultural Experiment Station as scientific contribution number
610.
80
1972] Herbarium — Hehre, Hodgdon & Pike 81
For a relatively small collection, the Cleaveland Her-
barium is notable for its representation of important
collectors. One can speculate that Professor Cleaveland
intended at the beginning to create an important botanical
center at Bowdoin.
The first two successful collectors who worked for Gray
in the 1840s are represented with 124 specimens by Ferdi-
nand Lindheimer (1801-1879) and 207 from New Mexico
by Augustus Fendler (1813-1883) of which 39 are isotypes
of species described by Gray and others. There are even
duplicate isotypes for 13 of the taxa. Dupree (1959) briefly
and McKelvey (1955) at much greater length discuss the
activities of these two botanical collectors of the American
Southwest.
There follows a partial and by no means complete list
of other botanists whose collections are to be found in the
Cleaveland Herbarium:
Michael S. Bebb (1833-1895) — 157 specimens, preponder-
antly from Illinois.
Joseph Blake (1814-1888) — nearly 1500, chiefly from
Cumberland County in Maine and some in Cóos, Carroll,
Grafton and Belknap Counties in New Hampshire.
William Boott (1805-1887) — 16 from the White Mountains
of New Hampshire.
William N. Canby (1831-1904) — 256 from Delaware and
adjoining areas.
Parker Cleaveland (1780-1858) — about 500, mostly from
Brunswick, Maine.
Chester Dewey (1784-1867) — 4 of Carex.
Asa Gray (1810-1888) — 16 (ex herbario W. S. Poor)
John S. Henslow (1796-1861) — father-in-law of Joseph
Dalton Hooker and whose recommendation led to Dar-
win's accepting the post of naturalist on the “Beagle” —
271 from Great Britain.
Marcus E. Jones (1852-1934) — 66 collected about 1880 in
Rocky Mountain area.
Peter D. Knieskern, M.D. (1798-1871) —119 from New
Jersey.
82 Rhodora [Vol. 74
William Oakes (1799-1848) —52 from Massachusetts and
New Hampshire, without date.
Thomas C. Porter (1822-1901) — 129 from Pennsylvania.
Henry P. Sartwell (1792-1867) — 114 from western New
York State, 87 of these being Carex.
Frank Lamson Seribner (1851-1938) — the first U.S.D.A.
Plant Pathologist — 140 from Manchester, Maine.
Edward Tuckerman, Jr. (1817-1886) — 250 mostly from
New England, a few from central Europe.
George Vasey (1822-1893) — 160 from Illinois.
Oliver R. Willis (1815-1902) — 335 from New Jersey.
Nearly all the specimens of any particular collector are
of different species and since many of the collectors were
specialists on certain groups and worked in different parts
of the world, the collection contains a very great number
of species.
The Herbarium of the University of New Hampsnire
(NHA) is now being computerized and it will soon be pos-
sible to provide interested persons with considerable infor-
mation as to the collectors represented as well as informa-
tion on the labels of the plants they collected. For the
Parker Cleaveland collection, in the case of each important
collector, we have made lists of all species represented.
Thus we are in a position now to supply this information.
Wooton & Standley (1915) give type localities for all taxa
and often specify the collectors. For Fendler, usually the
collection number is cited making it comparatively easy to
locate his isotypes. A large percentage of Fendler’s isotypes
of New Mexican plants is represented in the Cleaveland
collection.
List of Fendler isotypes in the Parker Cleaveland Her-
barium:
1. Selaginella Underwoodii Hieron., (in duplicate), Fend-
ler 1024.
Cheilanthes Fendleri Hook., Fendler 1015.
Poa Fendleriana (Steud.) Vasey, (in duplicate), Fend-
ler 932.
oo po
Herbarium — Hehre, Hodgdon & Pike 83
P. Bigelovii Vasey & Scribn., Fendler 931.
Aristida Fendleriana Steud., Fendler 973.
A. longiseta Steud., (in duplicate), Fendler 978.
Cyperus Fendlerianus Boeckel, Fendler 865.
Salix Fendleriana Anderss., Fendler 816.
S. irrorata Anderss., Fendler 812.
Quercus novemexicana (A.DC) Rydb., Fendler 809.
. Arceuthobium cryptopodium Engelm., Fendler 283.
Phoradendron juniperinum Engelm., Fendler 281.
Abronia Fendleri Standley, Fendler 739.
Lesquerella intermedia (S. Wats.) Heller, (in dupli-
cate), Fendler 38.
. Draba neomexicana Greene, (in duplicate), Fendler
43,
. Arabis Fendleri (S. Wats). Greene, Fendler 27.
Potentilla propinqua Rydb., Fendler 198.
Astragalus ceramicus Sheld., (in duplicate), Fendler
161.
A. lonchocarpus 'Torr., Fendler 160.
Euphorbia Fendleri Torr. & Gray, (in duplicate),
Fendler 800.
Sphaeralcea Fendleri A. Gray, Fendler 78.
Galpinsia Fendleri (A. Gray) Heller, Fendler 230.
Cymopterus Fendleri A. Gray, Fendler 274.
Daphnidostylis Fendleriana Klotzsch, Fendler.
Dodecatheon radicatum Greene, Fendler 549.
Mertensia Fendleri A. Gray, Fendler 625.
Oreocarya fulvocanescens (S. Wats.) Greene, (in du-
plicate), Fendler 632.
Oreocarya multicaulis (Torr.) Greene, Fendler 636.
Hydrophyllum Fendleri (A. Gray) Heller, Fendler 642.
Lycium pallidum Miers., Fendler 670.
. Physalis Fendleri A. Gray, (in triplicate), Fendler.
. P. similas A. Nels., Fendler 575.
Eupatorium Fendleri A. Gray, Fendler 347.
Erigeron cinereus A. Gray, (in duplicate), Fendler
914.
84 Rhodora [Vol. 74
35. Actinella argentea A. Gray, (in duplicate), Fendler
457.
36. Actinella Richardsonii Nutt. var. floribunda A. Gray
(in duplicate), Fendler 460.
37. Diplopappus ericoides Torr. & Gray var. hirtella A.
Gray, (in duplicate), Fendler 348.
38. Bidens tenuisecta A. Gray, Fendler 449.
39. Agoseris purpurea (A. Gray) Greene, Fendler 487.
For many years there has been a diminishing support
for small herbaria, some of which, like the Cleaveland Col-
lection at Bowdoin College, are of historic importance and
have scientific value. The space they occupy has been en-
croached upon by classrooms and by modern research
setups. This does not mean that the collections are less
important than they ever were. It does suggest that the
custodians of such herbaria should take steps, as Bowdoin
College has done, to make their specimens available to the
scientific community.
LITERATURE CITED
DUPREE, A. HUNTER. 1959. Asa Gray, Harvard University Press,
505 p.
MCKELVEY, SUSAN DELANO. 1955. Botanical Exploration of the
Trans-Mississippi West 1790-1850. The Arnold Arboretum of
Harvard University, 1144 p.
Wooton, E. O. and PAUL STANDLEY. 1915. Flora of New Mexico,
Contrib. U.S. Nat. Herb., Vol. 19.
DEPARTMENT OF BIOLOGY
SOUTHAMPTON COLLEGE OF LONG ISLAND UNIVERSITY
SOUTHAMPTON, NEW YORK 11968
DEPARTMENT OF BOTANY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824
DEPARTMENT OF PLANT SCIENCE
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824
STUDIES ON
THE AUDOUINELLA MICROSCOPICA (NAEG.)
WOELK. COMPLEX (RHODOPHYTA)
WILLIAM J. WOELKERLING
Several recent studies (Abbott 1962, West 1968, Woelker-
ling 1970, 1971) have indicated that the taxonomic status
of numerous species in the Audouinella complex (Acrochae-
tium-Rhodochorton complex) of the red algae is in need of
review. Many of these taxa have been described from
meagre material and (or) without regard for possible in-
traspecific variation, and recent work on several species
(Abbott 1968, West 1969, Woelkerling 1970) has resulted
in a reduction of some taxa to synonomy.
Preliminary work of the author on the audouinelloid
algae of the New England coast of North America has led
to a detailed consideration of the relationships of Audowi-
nella microscopica (Naegeli) Woelkerling to six closely re-
lated taxa: Acrochaetium crassipes Boergesen (1909, p. 1,
Fig. 1; 1915, p. 20, Figs. 11-13), A. catenulatum Howe
(1914, p. 84, pl. 31, Figs. 12-18), A. microfilum Jao (1936,
p. 240, pl. 10, Figs. 1-5), Kylinia collopoda (Rosenvinge)
Kylin (see Rosenvinge 1898, p. 41, Figs. 10-11; 1909, p. 81),
K. compacta (Jao) Papenfuss (see Jao 1936, p. 241, pl. 10,
Figs. 6-14), and K. moniliformis (Rosenvinge) Kylin (see
Rosenvinge 1909, p. 98, Figs. 28-29). (It should be noted
here that although the taxonomic proposals of Woelkerling
(1971) have been adopted in this study, older generic
names have been employed in cases of probable synonomy
in order to avoid making new and unnecessary nomencla-
tural combinations). A. crassipes and A. catenulatum were
described respectively from material collected in the Vir-
gin Islands (Boergeson 1909) and Peru (Howe 1914) while
the other four taxa have been described from or are re-
ported to occur along the New England and adjacent coasts.
These seven species have been distinguished from one
another on slight differences in habit, branching, cell size,
85
86 Rhodora [Vol. 74
chromoplast shape, and spore size. These apparent differ-
ences, however, may not be as taxonomically reliable as
formerly thought, and Woelkerling (1971) has suggested
that several or all of the taxa under discussion may be con-
specific. The aims of the present investigation have been:
1) to critically examine and compare morphologically the
type collections and other populations of these seven taxa,
and 2) to clarify taxonomic limits within this species com-
plex, particularly with reference to the New England flora.
MATERIALS AND METHODS
The morphological techniques employed in these studies
have been detailed elsewhere (Woelkerling 1970). Line
drawings have been made with the aid of a Leitz drawing
head microscope attachment; herbarium abbreviations fol-
low Lanjouw and Stafleu (1964).
Wherever possible, results have been based on the study
of populations (Table 1) rather than isolated individuals.
This approach has been facilitated by the nature of the
material; i.e., audouinelloid algae sometimes occur in large
numbers on various substrates and by virtue of their small
size, dozens or hundreds of individuals may be present in
a single collection. The data presented here in most cases
represents the results of study on numerous individuals
within each population; an exception is the type collection
of Acrochaetium crassipes Boergesen which is represented
in C only by several drawings. Data on this taxon has been
taken from the accounts of Boergesen (1909, 1915).
MORPHOLOGY
Audouinella microscopica, the earliest described member
of this complex, was first characterized by Kuetzing (1849),
and later Naegeli (1861) discussed and illustrated it in
somewhat greater detail. Although Kuetzing (1849, p. 640)
cited the Bay of Naples as the type locality, Hamel (1927;
1928) has indicated that the type collection came from
Torquay, England and is represented by specimen 454 in
Hauck and Richter's ^Phykotheca Universalis" (as Chan-
1972] Audouinella — Woelkerling 87
| M
Figs. 1-14. Audouinella microscopica (Naegeli) Woelkerling. Figs.
1-6. Monosporangial plants from type collection. Note variation in
habit and in development of basal cell wall. Figs. 7-8. Spermatangial
plants from type collection. Figs. 9-11. Cystocarpic plants from type
collection. Note remains of trichogyne and an apparent transversely
divided carpogonium (Fig. 9). Figs. 12-14. Sexual plants from Den-
mark removed from the host, Chordaria. Note variation in develop-
ment of basal cell wall and carpogonium with attached spermatium
(Fig. 13).
88 Rhodora [Vol. 74
transia secundata (Lyngbye) Thuret). The type specimen
in the Kuetzing collections at L contains only plants col-
lected by Naegeli in England (identical to those distributed
in Hauck and Richter) and none from the Bay of Naples.
The location given by Kuetzing (1849) is, therefore, ap-
parently in error.
Plants from the type collection of A. microscopica in L
as well as isotypes in FH and NY have been examined during
this study. The plants form a dense population on Enter-
omorpha (Chlorophyta), are 40-100 (-200) p tall, and are
attached to the host by unicellular bases with or without
enlarged lower cell walls (Figs. 1-11). One or occasionally
several erect filaments arise from the basal cell (Figs. 3,
4), and these may remain unbranched (Fig. 1-2) or bear
several secundly or irregularly arranged laterals (Figs. 3-
11). Cells are doliiform to cylindrical in shape, 6-8 (-12) &
wide and 6-12, (1-2 diameters) long; terminal hairs up to
40u long occur occasionally.
Both monosporangial and sexual plants are present in
the material examined. The monosporangia occur singly
or in pairs, are 7-10, long and 5-7, wide, are sessile or
stalked, and are scattered over the erect filaments (Figs. 1-
6). Spermatangia are ovoid, up to 4u long, occur singly or
in pairs, are sessile or stalked and are scattered over the
erect filaments (Figs. 7-8). Unfertilized carpogonia have
not been observed definitely in the type collection material,
but the remains of at least one apparently transversely
divided fertilized carpogonium has been seen (Fig. 9).
Gonimoblasts are several celled and give rise to terminal
or lateral carposporangia 7-10, long and 6-8, wide (Figs.
10-11). Only the clustered arrangement of the carpospor-
angia distinguishes the gonimoblast from a monosporangial
bearing branch. Tetrasporangial individuals have not been
observed but are reported by Schiffner (19531).
Hamel (1927, 1928) previously reported sexual individ-
uals, and Lund (1942) noted possible antheridia in Danish
plants. One Danish collection on loan from Dr. Lund has
been examined and found to contain numerous sexual in-
1972] Audouinella — Woelkerling 89
dividuals including specimens with carpogonia bearing at-
tached spermatia (Figs. 12-14). Lund (1942) referred the
Danish material to Chantransia (= Kylinia) collopoda, a
taxon here considered conspecific with Audouinella micro-
scopica.
Monosporangial plants more or less agreeing with the
above description of A. microscopica occur along the New
England and adjacent coasts, but up to the present time,
they have been referred (Edelstein & McLachlan 1966;
Edelstein et. al. 1967; Jao 1936; Taylor 1937, 1957) to
four other species (see above) including two (A. microfilum
and K. compacta) with type localities in the Cape Cod
region. An analysis (Table 1) of a number of New England
populations including those cited by the above authors,
strongly indicates that they agree in all essential features
with A. microscopica and are therefore justifiably referred
to that species.
The analysis further indicates that while the range in
basal attachment, height, branching, number of erect axes,
chromoplast shape, cell size, and spore size may vary some-
what from one population to the next (probably attribu-
table to variation in environmental factors and age), con-
siderable overlap in these characters exists between various
collections and in no cases can distinct specific limits be
drawn. Consequently, it appears that all these plants are
best regarded as members of a single, variable species —
A. microscopica. Woelkerling (1971) has found similar vari-
ation in southern Australian populations of this species.
Ecological data on A. microscopica in New England re-
mains scant. Specimens have been collected from July
through February, but the species probably occurs through-
out the year and has thus far escaped detection by virtue
of its small size. Sexual plants have not been reported to
date in New England, and indeed have been recorded only
twice from European waters. A. microscopica has been
found growing on a number of algae in New England waters
(Chaetomorpha, Chondria, Chondrus, Chordaria, Clado-
phora, Entromorpha, Polysiphonia, Porphyra) all of which
90 Rhodora [Vol. 74
occur in the sublittoral or in the drift. The species no doubt
enjoys a much wider host distribution and is to be sought
particularly on old and heavily epiphytized algae.
SYSTEMATIC IMPLICATIONS
The results of this study again (see Woelkerling 1971)
raise the question as to whether a number of taxa closely
related to Audowinella microscopica are really distinct
species. The type collections of six of these taxa
have been available for study, and a detailed analysis
(Table 1) strongly indicates that taxonomic distinctions
cannot be made among them on the bases of height, number
of erect axes, cell size, or spore size. As is the case for the
various New England collections, considerable overlap in
the above characters is evident in the type collection pop-
ulations, and specific limits cannot be clearly drawn. More-
over, a comparison of the type collection illustrations (see
Boergesen 1909; Howe 1914; Jao 1936; and Rosenvinge
1909) also indicates the great similarity of these taxa.
In addition to the above characteristics, apparent differ-
ences in habit, development of the basal cell wall, degree
of branching, origin of laterals, chromoplast shape, pres-
ence or absence of hairs, and position of sporangia have been
used in making specific distinctions. As Woelkerling (1971)
has shown in a lengthy review, the degree of branching,
presence or absence of hairs, and sporangial position are
not taxonomically reliable in general for making specific
distinctions in the Awdouinella complex, and the present
study supports this view. Thus, for example, the type col-
lection of A. microscopica contains plants which vary in the
degree of branching, may or may not have unicellular hairs,
and possess both terminal and lateral sporangia (Figs. 1-
11).
Hamel (1927, 1928) attempted to distinguish A. micro-
scopica from other members of this complex on the basis of
lateral branch origin. Thus, according to Hamel, the first
lateral branch in A. microscopica always arises from the
first cell above the base. The type collection of this species,
1972] Audouinella — Woelkerling 91
however, also contains unbranched plants (Figs. 1-2) and
plants in which the lateral arises from the basal cell (Fig
4) or from two or more cells above the base (Figs. 9-11).
Woelkerling (1971) found similar variation in Australian
populations of A. microscopica, and this variation also oc-
curs in New England populations. Thus it appears that
specific distinction based on origin of lateral branches is
not taxonomically reliable.
The presence of a much enlarged basal cell wall has been
used (Rosenvinge 1898, 1909; Lund 1942) to distinguish
Kylinia collopoda from Audouinella microscopica. However,
considerable variation in basal cell wall development occurs
in the type collection of A. microscopica (Figs. 1-11) as
well as collections made by Lund (Figs. 12-14), and Woel-
kerling (1971) reported a similar situation in southern
Australian populations of this species. This variation sug-
gests that this character also is not taxonomically reliable
for delimiting species in this complex.
Chromoplast shape (parietal vs. stellate) has also been
used (Taylor 1957) to distinguish these species. However,
recent work (West 1968, p. 92, 95; Woelkerling 1971) has
indicated that plastid shape shows considerable intraspe-
cific variation and therefore is not a generally trustworthy
taxonomic criterion. Some variation has already been re-
corded in plastids of A. microscopica (Woelkerling 1971),
and studies (e.g., Abbott 1962, p. 100; Boergesen 1937,
p. 39, 41; Drew 1928, p. 156, 176, 177, 182; Feldmann 1962,
p. 220; Levring 1937, p. 94) have noted similar variation
in other species. It appears, therefore, that the taxa under
discussion cannot be distinguished from one another solely
on apparent differences in chromoplast shape.
This study has not revealed any other criteria by which
these taxa may be reliably separated into distinct species;
consequently all are regarded here as conspecific with Au-
douinella microscopica.
The relationships of A. microscopica to four other taxa
remain uncertain, primarily because the type collections
have not been available for examination. The taxa include
92 Rhodora [Vol. 74
Chantransia meditierranea Levring (1942, p. 30, Figs. 1a-
g), C. minutissima Reinsch (1874-5, p. 33, tab. V, Fig. 2a,
tab. XI, Fig. 3a; not of other authors), C. trifila Buffham
(1892, p. 24, pl. 3, Figs. 1-4), and Kylinia scapae Lyle
(1929, p. 245, Figs. 6-7). A comparative examination of the
type collections of these taxa will almost certainly show
them to be conspecific with A. microscopica. The andro-
phores described and illustrated by Lyle (1929) probably
represent young, unelongated unicellular hairs.
COLLECTIONS EXAMINED
Types and Isotypes: DENMARK: Kattegat Channel, 17.
vii. 1890, Rosenvinge (C, Rosenvinge 863, Algae marinae
Danicae, type of Chantransia moniliformis Rosenvinge).
ENGLAND: Torquay, 1845, Naegeli (L 940285 .. . 306, type
of Audouinella microscopica (Naegeli) Woelkerling). Tor-
quay, 1845, Naegeli (FH, No. 454, *Phykotheka Univer-
salis", isotype of A. miscroscopica). Torquay 1845, Naegeli
(NY, No. 454, *Phykotheka Universalis", isotype of A. mi-
croscopica). GREENLAND: Holstenborg, 9. vii. 1895, Hanson
(c, type of Chantransia collopoda (Rosenvinge) Rosen-
vinge). PERU: La Punta, region of Callao, 25. i. 1907, Coker
(NY, type of Acrochaetium catenulatum Howe). UNITED
STATES: Black Rock, Sconticut Neck, New Bedford, Massa-
chusetts, 25. vii. 1934, Jao (MICH, Woods Hole, No. 275,
type of Acrochaetium compactum Jao). Norton Point,
Martha’s Vineyard, Massachusetts, 3. viii. 1934, Jao (MICH,
Woods Hole, No. 280 [not 274 as reported by Jao 1936,
p. 240], type of Acrochaetium microfilum Jao). VIRGIN
ISLANDS: St. Thomas (The Harbour), i. 1906, Boergesen
(c, type of Acrochaetium crassipes (Boergesen) Boerge-
sen).
Other collections: DENMARK: Vorupor, NW coast of
Jutland, 30. vii. 1929, Lund (C). MASSACHUSETTS: Cape
Codder Point (Falmouth), 19. xi. 1969, Woelkerling (2292,
personal collection). West Falmouth Harbor, 17. x. 1970,
Woelkering (2826, personal collection). Woods Hole
(Nobska Point), 4. ii. 1970, Woelkerling (2320, personal
1972] Audouinella — Woelkerling 93
collection). NOVA SCOTIA: Cranberry Cove, 12. ix. 1965,
Edelstein (Nat. Res. Council. Herb., Halifax, No. 1867, as
Acrochaetium microfilum Jao). Herring Cove, 18. i. 1966,
Edelstein (Nat. Res. Council. Herb., Halifax, No. 2217, as
Kylinia compacta (Jao) Papenfuss). Ketch Harbour, 7. ii.
1966, McLachlan & Edelstein (Nat. Res. Council. Herb.,
Halifax, No. 2244, as Kylinia collopoda (Rosenvinge) Pap-
enfuss). Peggy’s Cove, 26. viii. 1965, Edelstein (Nat. Res.
Council. Herb., Halifax, No. 2105, as K. collopoda).
SUMMARY
The relationship of Awudouinella microscopica to six close-
ly related taxa has been investigated with particular ref-
erence to the New England and adjacent coasts. The mor-
phology of monosporangial and sexual plants in the type
collection of A. microscopica is discussed and illustrated.
Collections from the New England region previously re-
ferred to Acrochaetium microfilum Jao, Kylinia collopoda
(Rosenvinge) Kylin, K. compacta (Jao) Papenfuss, and K.
moniliformis (Rosenvinge) Kylin have been found to rep-
resent specimens of Audouinella microscopica. A compara-
tive study of the type collections of these five taxa as well
as Acrochaetium crassipes Boergesen and A. catenulatum
Howe indicates that all taxa are conspecific with Audouinella
microscopica. Chantransia meditierranea Levring, C. min-
utissima Reinsch non. al., C. trifila Buffham, and Kylinia
scapae Lyle are regarded as probable synonyms.
ACKNOWLEDGMENTS
Sincere thanks are due Dr. T. Edelstein (Atlantic Re-
gional Laboratory, National Research Council, Halifax,
Nova Scotia), Dr. I. Mackenzie Lamb (FH), Dr. Soren Lund
(c), Dr. W. F. Prud’homme van Reine (L), Dr. Clark
Rogerson (NY), and Dr. W. R. Taylor (MICH) for the loan
of herbarium material. Miss Susan Heller kindly aided in
the preparation of the drawings. This work was supported
in part by National Science Foundation Grant GB-13250
to the Systematics-Ecology Program.
94 Rhodora [Vol. 74
Contribution No. 242 from the Systematics-Ecology Pro-
gram, Marine Biological Laboratory.
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ABBOTT, I. A. 1968. An examination of the type specimens of some
species of Acrochaetium (Rhodophyceae) Taxon 17: 518-520.
BOERGESEN, F. 1909. Some new or little known West Indian Flor-
ideae. Bot. Tidsskr. 30: 1-19.
BOERGESEN, F. 1915. The marine algae of the Danish West Indies.
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BOERGESEN, F. 1937. Contributions to a south Indian marine algal
flora. J. Indian bot. Soc. 16: 1-56, pl. 1.
BUFFHAM, T. H. 1892. Chantransia trifila, a new marine alga. J.
Quekett microsc. Club (Ser. 2) 5: 24-26.
Drew, K. M. 1928. A revision of the genera Chantransia, Rhodo-
chorton, and Acrochaetium, etc. Univ. Calif. Publs. Bot. 14(5) :
139-224, pl. 37-48.
EDELSTEIN, T. and MCLACHLAN, J. 1966. Species of Acrochaetiwm
and Kylinia new to North America. Br. phycol. Bull. 3: 37-41.
EDELSTEIN, T., J. MCLACHLAN, and J. S. CRAIGIE. 1967. Investiga-
tions of the marine algae of Nova Scotia. II. Species of Rho-
dophyceae new or rare to Nova Scotia. Can. J. Bot. 45: 202,
pl. 1-8.
FELDMANN, J. 1962. The Rhodophyta order Acrochaetiales and its
classification. Proc. IX Pacif. Sci. Congr. 4: 219-221.
HAMEL, G. 1927. “Recherches sur les genres Acrochaetium Naeg.
et Rhodochorton Naeg.” Dissertation. (Paris).
HAMEL, G. 1928. Sur les generes Acrochaetium Naeg. et Rhodo-
chorton Naeg. Revue algol. 3: 159-210.
Hauck, F., and P. RICHTER. 1894. *Phykotheka Universalis”. X.
Leipzig.
HowE, M. A. 1914. The marine algae of Peru. Mem. Torrey bot.
Club 15: 1-185, pl. 1-66.
Jao, C. C. 1936. New Rhodophyceae from Woods Hole. Bull. Torrey
Bot. Club 63: 237-257.
KUETZING, F. T. 1849. “Species Algarum" (Leipzing).
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The herbaria of the world. Ed. 5. Regnum Vegetabile 31: 1-251.
LEVRING, T. 1937. Zur kenntnis der algenflora der Norwegischen
Westkueste. Acta Univ. Lund (N.F. Avd. 2) 33(8): 1-147.
LEVRING, T. 1942. Meeresalgen aus dem Adriatischen meer, Sizilien,
und dem golf von Nepal. K. fysiogr. Sallsk. Lund Forh. 12(3):
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1972] Audouinella — Woelkerling 95
LUND, S. 1942. Chantransia collopoda K. Rosenv., a new rhodo-
phyceous alga of the Danish waters. Bot. Tidsskr. 46: 53-57.
LvLE, L. 1929. Marine algae of some German warships in Scapa
Flow and of the neighboring shores. J. Linn. Soc. (Bot). 48:
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giam". 1. Nuernbert.
RosENVINGE, L. K. 1898. Deuxieme memorie sur les algues du
Groenland. Meddr. Groenland 20: 1-125.
RosENVINGE, L. K. 1909. The marine algae of Denmark. I. Rho-
dophyceae 1. K. danske Vidensk. Selsk. Skr. Afd 7 Raekke 7(1):
1-151.
ScHIFFNER, V. 1931. Neue und bemerkenswerte meeresalgen. Hed-
wigia 71: 139-205.
TAYLOR, W. R. 1937. “Marine Algae of the Northeastern Coast of
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North America", Ed. 2. (Ann Arbor).
WEST, J. A. 1968. Morphology and reproduction of the red alga
Achrochaetium pectinatum in culture. J. Phycol. 4: 89-99.
WEST, J. A. 1969. The life histories of Rhodochorton purpureum
and R. tenue in culture. J. Phycol. 5: 12-21.
WOELKERLING, W. J. 1970. Acrochaetium botryocarpum (Harv.) J.
Ag. (Rhodophyta) in Southern Australia. Br. phycol. J. 5(2):
159-171.
WOELKERLING, W. J. 1971. Morphology and Taxonomy of the Au-
douinella complex (Rhodophyta) in Southern Australia. Aust. J.
Bot., Suppl. Ser., Suppl. 1(1971): 1-91.
DEPARTMENT OF BOTANY
UNIVERSITY OF WISCONSIN
MADISON, WISCONSIN 53706
[Vol. 74
Rhodora
96
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NOMENCLATURAL CLARIFICATION OF
TWO SPECIES OF VERBESINA (COMPOSITAE)
ENDEMIC TO FLORIDA:
JAMES R. COLEMAN
Two species of Verbesina are evidently endemic to Flor-
ida. One of these is restricted to a few coastal counties of
the Apalachicola region of western Florida whereas the
second has been collected from several counties in north-
eastern Florida. The species are disjunct by approximately
150 miles and are readily distinguished from each other by
several morphological features. Most notably, the eastern
species has decurrent leaves which form winged stems and
has rayed heads whereas the western species has wingless
stems and rayless heads. I have examined approximately
50 collections of these species and have observed no excep-
tions to these differences.
In 1822 Nuttall was the first to describe the eastern spe-
cies, naming it Actinomeris pauciflora and in 1841 he elab-
orated on his original description. Actinomeris pauciflora
was described as having decurrent leaves and 3-4 rays.
The description was based on a collection made by Ware.
I have been unable to locate this specimen and do not know
whether it still exists. However, there is no doubt as to the
species being described.
In 1872 Chapman applied the name A. pauciflora Nutt.
to a coastal species from western Florida. The species was
described as having wingless stems and the florets as hav-
ing “the marginal ones abortive.” Chapman’s lucid descrip-
tion leaves no doubt that he was erroneously applying the
name A. pauciflora Nutt. to the species from the Apalach-
icola region. In 1878 Chapman, evidently still unaware of
his error, redescribed the eastern species as new, giving
it the name Actinomeris heterophylla. He described A.
heterophylla as having winged stems, 5-10 rays and as oc-
"This investigation was supported by National Science Foundation
Grant GB-13923.
97
98 Rhodora [Vol. 74
curring in eastern Florida. I have examined the type and
it clearly belongs to the eastern species.
The next event in the nomenclatural history of these spe-
cies occurred in 1883 when Gray transferred the eastern
species to Verbesina forming the combination V. hetero-
phylla (Chapm.) A. Gray. Although no description was
provided, Gray did list Actinomeris heterophylla Chapm. as
asynonym. The description Gray provided in his Synoptical
Flora (1884) clearly reveals that he was referring to the
eastern species. In his 1883 publication Gray also trans-
ferred A. pauciflora Nutt. to Verbesina. However, since the
specific epithet was preoccupied by a Mexican Verbesina
described by Hemsley in 1881, he formed the new name
Verbesina warei A. Gray and gave Actinomeris pauciflora
Nutt. as a synonym. Gray’s description of Verbesina waret
in his Synoptical Flora is confusing in that he described
the stems as wingless but described the heads, in quotes, as
being 3-4 rayed. He stated that the species occurs near the
coast but did not specify the eastern or western coast. Col-
lections by Ware and by Chapman were cited; however,
Chapman collected both species and, as far as I am aware,
Ware collected only the eastern. There is, therefore, serious
doubt as to whether Gray was applying the name Verbesina
warei to the eastern or western species. These species had
been very infrequently collected prior to 1884 and it is pos-
sible that Gray never saw specimens of one or the other
or either species. It appears probable that Gray compiled
his description of V. warei largely or entirely from Nuttall’s
and Chapman’s descriptions of Actinomeris pauciflora Nutt.
It should be recalled that Chapman’s description was ac-
tually of the eastern species. Were this assumption true, it
could account for the mixing of characteristics of both spe-
cies in Gray's description of Verbesina warei, for his use
of quotes in referring to the rayed condition and to his fail-
ure to indicate whether the species occurred on the east or
west coast as the two descriptions conflict on this point.
In citing the collections of Ware and Chapman, Gray was
probably again simply following Nuttall and Chapman.
1972] Verbesina — Coleman 99
In their revision of the genus Verbesina, Robinson and
Greenman (1899) applied the name Verbesina heterophylla
A. Gray to the eastern species and listed Actinomeris heter-
ophylla Chapm. in synonymy. The western species was
called Verbesina warei A. Gray and Actinomeris pauciflora
Nutt. was given as a synonym.
Small (1903), in his Flora of the Southeastern United
States, also called the eastern species Verbesina hetero-
phylla (Chapm.) A. Gray, but formed a new combination,
Verbesina pauciflora (Nutt.) Small, for the western species
and listed both Actinomeris pauciflora Nutt. and Verbesina
warei A. Gray in synonymy. Small also was evidently un-
aware that Nuttall’s name was originally applied to the
eastern species and was either unaware of, or ignored, the
fact that the combination Verbesina pauciflora was pre-
occupied.
Alexander, in treating these species in Small’s Manual
of the Southeastern United States (1933), transferred both
to the genus Pterophyton, forming the new combination
P. pauciflorum (Nutt.) Alex. for the western species and
P. heterophyllum (Chapm.) Alex. for the eastern species.
The genus Pterophyton Cass. had been reduced to a section
of Verbesina by Torrey and Gray in 1842. This reduction
appears justified when the genus Verbesina is considered
in its entirety rather than on a regional basis.
The question arises as to the correct names for the two
species under consideration. Although the earliest binom-
ial applied to the eastern species is Actinomeris pauciflora
Nutt. (1822), the epithet pauciflora cannot be used in com-
bination with Verbesina since the combination V. pauci-
flora is preoccupied by Hemsley’s Mexican species. The
second oldest binomial given to this species is Actinomeris
heterophylla Chapm. (1878), the specific epithet of which
was transferred by Gray to form the correct name for the
eastern species, Verbesina heterophylla (Chapm.) A. Gray.
Although Verbesina warei A. Gray has been used fairly
commonly for the western species, Gray did give Actinom-
eris pauciflora Nutt. as a synonym in making the transfer.
100 Rhodora [Vol. 74
The name Verbesina warei A. Gray is, therefore, associated
with the type of Actinomeris pauciflora which belongs to
the eastern species. Verbesina warei must then be con-
sidered a synonym of V. heterophylla (Nutt.) A. Gray.
Verbesina heterophylla (Chapm.) A. Gray, Proc. Amer.
Acad. 19: 12, 1883.
Actinomeris pauciflora Nutt., Sill. Journ. 5: 301, 1822;
Trans. Amer. Phil. Soc. 7: 364, 1841. Type: East Florida,
Ware s.n. (location unknown).
Actinomeris heterophylla Chapm. in Coult., Bot. Gaz. 3:
6, 1878. Type: East Florida, Chapman s.n., 1871. (NY!)
Verbesina warei A. Gray, Proc. Amer. Acad. 19: 12,
1883. nom. nov. for A. pauciflora Nutt., not Verbesina
pauciflora Hemsl. 1881.
Pterophyton heterophyllum (Chapm.) Alex. in Small,
Man. S. E. Fl. 1444, 1933.
The earliest description of the western species was given
by Chapman in 1872; however, he failed to recognize it
as a new species and misidentified it as Actinomeris pauci-
flora Nutt. Probably the most commonly used name for
this species is Verbesina warei A. Gray. However, this
name belongs to the eastern species. Since the western
species has never been given a valid name, it is described
as a new species.
Verbesina chapmanii J. R. Coleman, sp. nov. Perennis
ca 0.5-1.1 m alta; caulis simplex non alatus; folia opposita
vel alterna, sessilia vel subsessilia, lanceolata, elliptica vel
lata, scabra; capitula 1-5 terminalia pedunculata, plerum-
que 1-2 cm lata, discoidea; involucra 2-seriata, ca 8-12 mm
elliptico-oblonga, plerumque 5-9 cm longa, usque ad 3.2 cm
alta; flores lutei; achaenia alata, oblongo-obovata, plerum-
que 6-8 mm longa, 4-5 mm lata, glabra vel subglabra, epap-
posa vel pappi setis usque ad ca 1 mm longis praedita.
Perennial herbs mostly 0.5-1.1 m tall with heavy rhi-
zomes; stems numerous, unbranched below the inflores-
cence, wingless, hispidulous near the heads, otherwise
glabrous or subglabrous; leaves alternate or opposite, be-
1972] Verbesina — Coleman 101
coming bractiform above, rigid, sessile or subsessile, lance-
olate, elliptical or elliptic-oblong, serrulate to subentire,
obtuse or less frequently acute, to ca 12 cm long, commonly
5-9 cm long, to ca 3.2 cm wide, scabrous; inflorescence
solitary or to 5-headed; heads mostly 1-2 cm wide, discoid;
phyllaries 2-seriate, ca 8-12 cm tall; florets yellow, mostly
12-14 mm tall; paleae ca 8-10 mm tall, mostly red-purple
apically; achenes mostly red-purple, oblong-obovate, mostly
6-8 mm long, 4-5 mm wide, glabrous or sparingly hirtel-
lous apically, lateral wings to ca 1 mm wide, continuous
about the crown to form a shallow cup, pappus of 2 short
horns or lacking.
Type: open prairie-like pine-cypress savanna, 3 mi N
of Orange, Liberty Co, Florida, McDaniel 4468 (GH!)
DEPARTMENT OF BOTANY
THE UNIVERSITY OF GEORGIA
ATHENS, GEORGIA 30601
LITERATURE CITED
CHAPMAN, A. W. 1872. Flora of the southern United States. Ivi-
son, Blakeman, Taylor and Co., N. Y. 621 pp.
GRAY, A. 1884. Synoptical Flora of North America. Vol. I. Part
II. Smithsonian Institution, Washington, D. C.
HEMSLEY, W. B. 1881. Biologia centrali-americana. Bot. Vol. II.
London.
SMALL, J. K. 1903. Flora of the southeastern United States. Pub-
lished by the author, N. Y. 1370 pp.
. 1933. Manual of the southeastern flora. Univ. of
North Carolina Press, Chapel Hill. 1554 pp.
ROBINSON, B. L. and J. M. GREENMAN. 1899. A synopsis of the
genus Verbesina, with an analytical key to the species. Proc.
Amer. Acad. 34: 534-5064.
TORREY, J. and A. GRAY. 1842. A Flora of North America Vol. 2
pt. 2. Wiley and Putnam, N. Y.
SOMATIC CHROMOSOME NUMBERS
FOR SOME ASTERACEAE
MILOSLAV KOVANDA
Few plant groups can claim the attention of botanists
that the American Asteraceae have been enjoying in recent
years. In addition to thorough monographic studies an un-
precedented series of chromosome number reports has ap-
peared, and it may seem trivial to publish additional counts
for a handful of miscellaneous taxa. Yet the need for re-
peated chromosome sampling cannot be stressed enough.
Previous studies have revealed that intraspecific polyploidy
is widespread, and chromosome counts from as many popu-
lations as possible throughout the range of each species are
required to gain an idea of the nature and significance of
this diversity. Many of the taxa discussed below are wide-
ranging, polymorphous entities that have been counted but
once or twice, making the "cytofloristic" approach impera-
tive. The counts for Artemisia saxicola and Haplopappus
lyalli are believed to be the first to be published for these
species. A new ploidy level is established for Chrysopsis
fulcrata.
MATERIALS AND METHODS
All chromosome counts were obtained from squashes of
root-tips of young seedlings grown in the laboratory.
Achenes were collected in the field in the late summer and
fall of 1970. All germinated without special treatment in
2 to 9 days when placed on moist filter paper in Petri dishes
and maintained at room temperature. Excised root-tips were
pretreated in a saturated solution of 8-hydroxyquinoline
for 3 hours, fixed in 1 : 3 acetic alcohol, and stained in lacto-
propionic orceine (for details, see Kovanda 1970). The
counts were verified in five or more plantlings. Voucher
specimens were collected for all plants sampled and will
be deposited at the Herbarium of the National Museum in
Prague (PR). Duplicates, when available, will be deposited
102
1972] Kovanda — Asteraceae 103
in the United States National Herbarium (US). A portion
of the material examined has been planted in the experi-
mental plot of the Botanical Institute, Czechoslovak Acad-
emy of Sciences, Prague, for future observations. All col-
lections were made by Kovanda or by Kovanda and S. G.
Shetler.
OBSERVATIONS
TRIBE VERNONIEAE
1. Elephantopus carolinianus Willd.
Maryland, Montgomery County: thicket along Chesapeake
& Ohio Canal near Glen Echo, vicinity of Washington, D.C.,
Kovanda 3558. 2n = 22.
This species recently has been examined cytologically
from two widely separated areas: Virginia (Baldwin &
Speese, 1955) and Texas (Lewis et al., 1962). In both cases,
the authors report the same chromosome numbers, 2n — 22
and n = 11, respectively. The entire genus seems to have
the base number x — 11, and all species counted heretofore
have proved to be diploid. The count 2n = 44 for E. tomen-
tosus L., ascribed by Fedorov et al. (1969) to S. B. Jones
(1966), is in error, Jones having counted n — 11 and listed
the earlier count of 2n = 22 made by Baldwin & Speese.
TRIBE EUPATORIEAE
2. Eupatorium coelestinum L.
Maryland, Montgomery County: dry woodland near Cabin
John Bridge, vicinity of Washington, D.C., Kovanda 3529.
2n — 20.
This number was reported previously by Grant (1953)
for material from Pennsylvania, Kentucky, and Tennessee.
By its chromosome morphology this species differs from all
other species of Hupatorium having z = 10 and would be
best referred perhaps to the genus Conocliniwm DC. (see
Grant 1953).
9. Liatris punctata Hook.
Colorado, Larimer County: Rist Canyon, grassy roadside
about 8 mi. W. of Bellevue, 6800 ft., Kovanda & Shetler
3169. 2n = 40.
104 Rhodora [Vol. 74
In her study of the genus Liatris, Gaiser (1950) has dem-
onstrated that two chromosome races exist in this species,
which she was able to identify with previously described
infraspecific taxa. Diploids (2n — 20) have been referred
to var. nebraskana Gaiser, and the tetraploid level (2n =
40) is represented in var. punctata (var. typica Gaiser)
and var. mexicana Gaiser. The two cytotypes are not al-
ways easily recognizable on a morphological basis but show,
in addition to marked differences in the beginning of
anthesis, also a clear geographical separation. Tetraploids
seem to occur throughout most of the extensive range of
the species (with var. mexicana, being limited to its south-
ern part) while the diploid variety is obviously localized.
The Rist Canyon population certainly falls within the typ-
ical variety but approaches somewhat var. nebraskana by
its leaves which are almost entirely devoid of cilia.
TRIBE ASTEREAE
4. Aster occidentalis (Nutt.) T. & G.
Colorado, Larimer County: brushy hillside north of Stove
Prairie School, S. of Poudre River, 6500 ft, Kovanda &
Shetler 3183. 2n — 10.
The count reported here is consistent with those pub-
lished by Huziwara (1958, 1959) and Solbrig et al. (1969).
The Western Aster is one of the few aster species not in
the x = 9 series. The number x = 8, likewise x = 5 in
A. exilis Ell. and A. foliaceus Lindl., and x — 13 in A. chil-
ensis Nees, is clearly secondary (Solbrig et al., 1969). Other
species with « — 8 include, for instance, A. adscendens
Lindl., A. ericoides L., A. hirtifolius Blake, A. lateriflorus
(L.) Britt., and A. puniceus L.; most of them are tetra-
ploid, with 2n = 32. It is interesting to note that base num-
bers other than x — 9 are not known to occur in species
native to the Old World, which supports the contention that
the center of origin of this huge genus is America.
5. Chrysopsis fulcrata Greene
Colorado, Clear Creek County: abandoned campground in
Clear Creek valley, about 12 mi. south of Georgetown, 11,440
1972] Kovanda — Asteraceae 105
ft., Kovanda & Shetler 3331. 2n — 18.
Colorado, Larimer County: Rist Canyon, grassy roadside
about 8 mi. west of Bellevue, 6800 ft. Kovanda & Shetler
3172. 2n = 36.
Counts by Raven et al. (1960), Turner et al. (1962), Sol-
brig et al. (1964), Chuksanova et al. (1968), and Solbrig
et al. (1969) indicate that both diploids and tetraploids are
present in the Chrysopsis villosa species aggregate. In C.
fulcrata, a segregate defined mainly by leaf-like bracts sub-
tending the heads, glandular phyllaries and sessile leaves,
only the diploid level (» = 9) has been sampled (Solbrig
et al. 1964, 1969). An examination of material from Rist
Canyon clearly showed 36 somatic chromosomes. The col-
lection was compared with Greene’s type material of C. ful-
crata in the U. S. National Herbarium, and no specific dif-
ferences could be seen except for the leaves. These are
sometimes slightly tapering at the base in my plants but
are distinctly semi-amplexicaul in the type specimen, col-
lected in the Organ Mountains, New Mexico. In the Clear
Creek diploid this tapering is even more conspicuous, mark-
ing an intergradation with C. hispida (Hook.) DC. and C.
viscida (A. Gray) Greene. The taxonomy of the entire com-
plex is not very well understood, and there appears to be
little correlation between the cytological findings and mor-
phological segregation of the several species.
6. Chrysothamnus nauseosus (Pallas) Britt. subsp. nau-
seosus
Wyoming, Albany County: stony roadside just east of Cen-
tennial, 8000 ft., Kovanda & Shetler 3205. 2n = 18.
The Rabbitbrush was the subject of a detailed cytotax-
onomic study by Anderson (1966) who demonstrated that
polyploidy is rare in this genus. He found only the diploid
number 2n — 18 in many plants of C. nauseosus divided
among 12 subspecies. The same number has been counted
by Raven et al. (1960) and Solbrig et al. (1964). The count
reported here is only a further proof (if one is needed) that
the astonishing morphological variation shown by this spe-
cies cannot be credited to polyploidy.
106 Rhodora [Vol. 74
7. Erigeron speciosus (Lindl.) DC.
Wyoming, Teton County: Teton Pass, 9000 ft., Kovanda &
Shetler 3252. 2n = 18.
Previous counts seem to indicate that this species com-
prises diploid and tetraploid cytotypes. Ferri (1961), Zhu-
kova (1964, 1967) and Huziwara (1965) give, alternatively,
^ — 9 or 2n — 18, contrasting with 2» — 36 counted by
Vilmorin and Chopinet (1954). Unfortunately, all these
counts were made on cultivated material of uncertain origin,
and it remains to be seen whether both the races occur in
natural habitats. The Teton Pass population proved to be
diploid and appears referable to var. macranthus (Nutt.)
Cronquist, which has been counted as E. macranthus Nutt.
by Bergman (1942) and Zhukova (1964). Both counts were
2n — 18, too, but the sources of the material were not given.
8. Grindelia aphanactis Rydb.
New Mexico, Rio Arriba County: Rio Grande Canyon,
about 5 mi. NE. of San Juan Pueblo, 5800 ft., Kovanda 3404.
2n — 24.
The only previous count for this species was n — 12 by
Raven et al. (1960) on material from Arizona. Grindelia
Willd. is a natural genus, and, chromosomally, a very homo-
geneous one, with the base number only x — 6. The majority
of species seem to be diploid, with 2n = 12 (see also Whit-
aker and Steyermark 1935, Dunford 1964, 1970).
9. Haplopappus lyallii A. Gray
Colorado, Clear Creek County: Loveland Pass, 11,900 ft.,
Kovanda & Shetler 3382. 2n — 18.
No previous cytological work seems to have been done
on this tiny goldenweed. Two members of the section Tones-
tus, H. eximius Hall and H. peirsonii (Keck) Howell, have
been shown to have 2n — 18 and ca. 90, respectively (Steb-
bins in Howell 1950), which would indicate that the base
number for that group is 9. But considering the variety
of chromosome numbers encountered in other species (see
e.g., Raven et al. 1960, Solbrig et al. 1964, 1969), any such
statement would be premature. In the genus Haplopappus,
1972] Kovanda — Asteraceae 107
as broadly conceived by Hall (1925), « = 9 as a basic
chromosome number is not uncommon; however, it is puz-
zling to discover that it is largely associated with the
shrubby groups of the southwestern deserts, such as sec-
tions Ericameria and Stenotopsis. The only other herba-
ceous species hitherto known to have the base number « =
9 is H. clementis (Rydb.) S. F. Blake of the section Pyr-
rocoma. The genus Tonestus was proposed by Nelson (1904)
to accommodate low perennial herbs from the Rocky Moun-
tains area and “to bring together under one name these
species [H. laceratus Henderson, H. lyallii A. Gray, and
H. pygmaeus T. & G.] which are allied by habit and mor-
phological characters to each other and are aberrant in any
recognized genus or genera in which they can be placed."
When more cytological information is available, it may be-
come necessary to reconsider the generic and sectional
limits.
10. Solidago flexicaulis L.
Maryland, Harford County: woodland along Susquehanna
River below Deer Creek, between Schweers Landing and
Lapidum, Kovanda & Shetler 3638. 2n — 18.
The same diploid number has been found in plants from
several Canadian localities (see Beaudry and Chabot 1959,
Kapoor and Beaudry 1966, Kapoor 1970). A tetraploid
cytotype was reported from Michigan (Solbrig et al. 1964).
11. Solidago multiradiata Ait.
Colorado, Clear Creek County: Clear Creek valley, about
12 mi. west of Georgetown, 11,400 ft., Kovanda & Shetler
3872. 2n — 18.
This arctic-alpine species was once thought to be diploid
(Beaudry and Chabot 1959, Beaudry 1963) but has recently
been shown to have tetraploid races, 2n — 36, in western
Canada and Alaska (Taylor 1967, Johnson & Packer 1968,
Packer 1968). The count recorded here is the first made on
material from the Southern Rockies where the species
reaches its southernmost limit. The collection is var. scopu-
lorum A. Gray.
108 Rhodora [Vol. 74
12. Solidago sempervirens L.
Delaware, Sussex County: salt marsh about 2 mi. S. of
Rehoboth Beach, Kovanda 3589. 2n — 18.
Several counts have been made on this halophyte, reveal-
ing the presence of diploid and tetraploid levels. However,
these races do not correspond with the two major segments
recognized taxonomically within S. sempervirens. The
southern entity, var. mexicana (L.) Fern., was first re-
ported to be diploid but was later found to have also 2n =
36, and the same holds for the more northern var. semper-
virens (Goodwin 1937, Beaudry and Chabot 1959, Beaudry
1963, Kapoor 1970). Likewise, the geographical distribu-
tion of the cytotypes does not seem to form a definite
pattern, but there are too few counts to allow any generali-
zation. Their morphological differences and ecological pref-
erences, if any, also await future research.
TRIBE HELIANTHEAE
13. Rudbeckia hirta L.
Maryland, Frederick County: Catoctin Mountains, old field
near cabin of David Scott, west of Thurmont, Kovanda 2896.
2n = 38.
Colorado, El Paso County: Pikes Peak, 9000-foot level,
streamside, Kovanda & Shetler 3398. 2n — 38.
Rudbeckia hirta is an extremely variable species, and
many local variants have been given varietal or specific
rank. Perdue (1957) greatly reduced their number by
recognizing five geographical varieties: var. hirta, var.
pulcherrima Farwell, var. corymbifera Fernald, var. angus-
tifolia (T. V. Moore) Perdue, and var. floridana (T. V.
Moore) Perdue. Most controversial perhaps is the treat-
ment of var. pulcherrima, which was first described as R.
serotina Nutt. and to which both the populations sampled
by the present author belong. Fernald (1948) considered
it a western entity which had spread eastwards, becoming
thoroughly naturalized throughout most of the United
States, and advocated its specific status. His ideas, however,
received little support from later workers (Perdue 1957,
Core 1962). In contrast to its intricate morphological struc-
1972] Kovanda — Asteraceae 109
ture, the species complex appears remarkably uniform in
cytology, 2n = 38 being the only chromosome number
so far obtained (Battaglia 1946, 1947, Perdue 1959). The
same chromosome number has also been reported for R.
serotina (Battaglia 1947, Mulligan 1959).
14. Viguiera multiflora (Nutt.) S. F. Blake
Utah, Daggett County: Uinta Mountains, between Cub
Creek and Y Creek, ca. 6 mi. W. of Red Canyon Road, 8500
ft., Kovanda & Shetler 3275. 2n = 16.
Two earlier counts for this species give the gametic
chromosome number as n = 8 (Heiser and Smith 1955,
Heiser 1963).
TRIBE HELENIEAE
15. Hymenoxys richardsonii (Hook.) Cockerell
New Mexico, Santa Fe County: 2 mi. NE. of Santa Fe,
7200 ft., Kovanda 3428. 2n — 30.
The chromosome number of this species has been deter-
mined repeatedly as either n — 15 or 2n — 30 (Speese and
Baldwin 1952, Strother 1966). The count 2n — 28, made by
Taylor and Brockman (1966) on a collection from Saskatch-
ewan, suggest that there are two chromosome races in this
species whose morphology and geographical distribution re-
quire further study. Material examined by the present au-
thor belongs to var. floribunda (A. Gray) Parker and was
collected in or near the type locality (“rocky hills, as well
as plains and creek bottoms, around Sante Fe", A. Gray in
Mem. Amer. Acad., n.s. 4: 101, 1849).
16. Pericome caudata A. Gray
New Mexico, Santa Fe County: Sangre de Cristo Moun-
tains, along the Chamisa Trail, 7600 ft, Kovanda 3426.
2n = 36.
Pericome caudata has recently been counted by Turner
and Flyr (1966) and Powell (1968). Both obtained a game-
tic chromosome number of » — 18. Raven and Kyhos
(1961) have found the same number in the closely related
P. glandulosa Goodm. It would appear that the small genus
Pericome has the base number x = 9.
110 Rhodora [Vol. 74
TRIBE ANTHEMIDEAE
17. Artemisia borealis Pallas
Colorado, Clear Creek County: Loveland Pass, 11,990 ft.,
Kovanda & Shetler 3376. 2n = 18.
Artemisia borealis is taxonomically complex in the Old
and New World, and attempts to subdivide it into more
natural entities have not been very successful. Diploids and
tetraploids, based on x — 9, are known to exist in this group.
Both these races have been shown to occur in North Amer-
ica, but their disposition is not at all clear. Diploids (2n =
18) have been discovered in Greenland (Jörgensen et al.
1958), northern Quebec (Hedberg 1967), and Alberta
(Packer in Johnson and Packer 1968), but tetraploids
(2n = 36) have so far been found only in Alaska (Johnson
and Packer 1968). The Loveland Pass record is important
because the count was made on material from the most
southern portion of the species range. The plants examined
have basal leaves three times pinnatifid, densely hairy on
both surfaces; stems almost totally glabrous; heads erect,
arranged in a narrow, spike-like inflorescence, somewhat
interrupted in the lower half; phyllaries villous; achenes
silky-hairy.
18. Artemisia frigida Willd.
Colorado, Park County: 2 mi. N. of Fairplay on Route 9,
9900 ft., Kovanda & Shetler 3379. 2n = 18.
This number was obtained previously by Lóve and Lóve
(1964) on plants from Manitoba and by Knaben (1968)
on plants from Alaska. Kawatani and Ohno (1964) give
2n — 18 for garden material of unspecified origin.
19. Artemisia saxicola Rydb.
Colorado, Grand County: alpine tundra above Berthoud
Pass, 12,400 ft., Kovanda & Shetler 3324. 2n — 18.
Colorado, Summit County: near Pass Lake, just S. of
Loveland Pass, 11,000 ft., Kovanda & Shetler 3367. 2» =
18.
These are probably the first counts for the Rocky Moun-
tains segregate of A. arctica Less. This species, ranging in
northeastern Asia and northwestern North America and
1972] Kovanda — Asteraceae 111
itself a segregate of the European A. norvegica Fries, has
been counted many times on Asiatic material and is known
to have two chromosome numbers, 2n — 18 and 36. (Kita-
mura 1957, Sokolovskaya 1963, Kawatani and Ohno 1964,
Zhukova 1964, 1965, 1967). Only two records are available
from North America (Johnson and Packer 1968, Taylor
and Mulligan 1968). Both counts are tetraploid and were
made on plants from Alaska and the Queen Charlotte
Islands, respectively. It is extremely interesting to discover
that the Rocky Mountain populations are diploid and, there-
fore, more primitive than the northern A. arctica. This,
combined with the geographical isolation and a degree of
morphological differentiation, would justify the specific rank
assigned to this race by Rydberg but disregarded by most
later students.
TRIBE SENECIONEAE
20. Senecio longilobus Benth.
New Mexico, Santa Fe County: near Bishop's Lodge, N.
of Santa Fe, 6900 ft., Kovanda 3425. 2n — 40.
This number conforms to the findings of Jackson (1959)
and Ornduff et al. (1967) who determined n = 20 on plants
from New Mexico and Texas. Powell and Turner (1963)
have reported the same chromosome number for Mexican
material under the name of S. filifolius Nutt. The section
Suffruticosi, confined to the western United States and
northern Mexico, appears chromosomally uniform. All spe-
cies hitherto examined have proved to have » — 20 (Stout-
amire and Beaman 1960; Turner, Ellison and King 1961;
Ornduff et al. 1963, 1967), and Ornduff et al. (1963) have
proposed that the base number is either x = 10 or x = 20.
ACKNOWLEDGMENTS
This work was carried out during my appointment as a
Visiting Research Associate in the Department of Botany
at the Smithsonian Institution, Washington, D.C., 1970-71.
It is my pleasant duty to express my appreciation to S. G.
Shetler, Associate Curator, Department of Botany, for ar-
ranging the Rocky Mountain trip, making my stay in the
112 Rhodora [Vol. 74
United States an eventful and interesting one, and for many
stimulating ideas. My sincere thanks are due to W. A.
Weber, University of Colorado, Boulder, who kindly checked
or made identifications of my collections from the Colorado
Rockies; however, I accept full responsibility for the tax-
onomic opinions expressed here. The fieldwork was sup-
ported in part by the Smithsonian Research Foundation
(Sg-0653061/C2).
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L. RUDENBERG. 1964. Chromosome numbers in Compositae. V.
Astereae II. Amer. J. Bot. 51(5): 518-519.
, , , and ?
1969. Chromosome numbers in Compositae VII. Astereae III.
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SPEESE, B. M. and J. T. BALDWIN. 1952. Chromosomes of Hyme-
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1965. Kariologischeskaya kharakteristika neko-
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INSTITUTE OF BOTANY
CZECHOSLOVAK ACADEMY OF SCIENCES
PRAGUE
INTERACTIONS BETWEEN APIS MELLIFERA
(HYMENOPTERA: APIDAE)
AND TRIBULUS CISTOIDES (ZYGOPHYLLACEAE)
DANIEL F. AUSTIN
In southern Florida a common weed of disturbed areas
is Tribulus cistoides L. (Burr Nut or Puncture Vine). A
frequent visitor to this plant is the Honey Bee, Apis melli-
fera L. Little is known about the reproductive biology of
this weedy plant (Porter, 1971; pers. comm.), but observa-
tions in Florida suggest some interesting interactions be-
tween Apis and Tribulus.
According to Grant (1950) there is a division of labor
among field bees of Apis mellifera. Some of the workers
carry water; others gather pollen, nectar or propolis, while
others search for new food sources. The constancy of an
individual to her job and to a particular plant species has
been amply verified by numerous authors and summarized
by Grant (op. cit.).
While I have observed Hemiptera, Lepidoptera, Diptera,
and other Hymenoptera on Tribulus in Florida, the most
frequent visitors observed on the plants near Boca Raton,
Palm Beach County, are Apis mellifera. There is a marked
behavioral division among the Honey Bee individuals visit-
ing the plants.
Most of the Honey Bees visiting the flowers approach in
the “normal” bee fashion described by Meeuse (1961) and
Fagri & Pijl (1966). Instead of landing on the stamens
and stigmas as they should, however, they circle around
the flower and land on the outside of the calyx and corolla
(Figs. 1-4). After landing they separate two petals with
their front legs, insert their tongues, and sip nectar (Fig.
3). They continue this procedure around the flower until
they have collected nectar between all the petals. Once a
flower has been utilized, the bees fly to another and repeat
the process. Several flowers are usually visited before the
bee disappears.
117
118 Rhodora [Vol. 74
Figure 1-4. Apis mellifera visiting the flowers of Tribulus cis-
toides from the back. Note the tongue of the bee in Fig. 3.
Other Honey Bees approach and land within the corolla
in the *normal" manner. In so doing they position their
bodies over the stigma and stamens (Figs. 5-7). Normally
they insert their tongue between the petals and the nectary
(Fig. 7), sip nectar, and turn around the flower to drink
nectar from other nectaries. They dust much of their body
with pollen as they turn (Figs. 5-6). After the nectar has
been collected, they often brush the stamens with their
front legs to gather pollen.
After nectar has been taken and pollen gathered with the
front legs, the bees may fly to another flower to repeat
the process. It is common, however, for an individual to
brush pollen, fly off and hover in front of the flower, re-
turn to brush more pollen, and hover again. This may be
done three or four times before the process is repeated at
another flower.
1972] Tribulus cistoides — Austin 119
|
"
t
LÀ Tn ;
Figure 5-8. Bees visiting the flowers of T'ribulus cistoides from
the front. Fig. 5-7. Apis mellifera. Fig. 8. Chloralictus.
Those individuals which exhibit this hovering behavior
usually have their pollen baskets full. The bees with pollen
in the pollen baskets have little or no pollen adhering to
the hairs of their legs and body. Bees which have no pollen
in their baskets rarely hover, and their bodies and legs
usually are dusted with pollen. The individuals illustrated
here (Figs. 5-7) were photographed between 3:40 and
4:40 p.m. in December. Although their bodies are dusted
with pollen, the pollen baskets are empty.
There has been a seasonal shift in the labor of the in-
dividuals visiting the plants. From September until Decem-
ber about 18 out of 20 Honey Bees gathered nectar. None
of the nectar gathering bees were ever seen collecting pollen.
In February the visitation had dropped from about 20 bees
to about 4 bees per hour. All of those bees seen in Febru-
ary were gathering pollen.
120 Rhodora [Vol. 74
Associated with the reduction of visits by Honey Bees
was an increase of visits by other insects. During Febru-
ary the flowers were visited by five bee species, three Dip-
tera, one Hemiptera, and one Lepidoptera in addition to
the Honey Bees. Equivalent observations in December re-
vealed visits by Honey Bees with one other bee species
(Chloralictus — Fig. 8), two Diptera, and one Lepidoptera.
Honey Bees were most common in December; other bees
(Agapostemon, Chloralictus (2 species), Halictus, and Lasi-
oglossum) in February.
The reason for the apparent preference for Tribulus nec-
tar over pollen during the period from September to De-
cember is not clear. Nor is it clear that these observations
are the result of a rigid division of labor. There must be
some strong force (attractant?) which guides the Honey
Bees around the visual guides usually followed in flowers.
Since much of the underside of some Honey Bees is
dusted with pollen, self- or cross-pollination could easily be
accomplished. The plants have not been tested for autog-
amy or self-compatibility, but the ample fruits present
suggest that Honey Bees or other visitors may successfully
complete pollination. Porter (pers. comm.) indicates that
T. cistoides is protandrous and thus probably outcrossing.
Self-compatibility, however, should not be ruled out until
demonstrated.
Tribulus cistoides is native to tropical and subtropical
Africa (north to Cape Verde on the west and Mozambique
on the east). Since it is now widely dispersed throughout
the drier tropics, it often becomes a bothersome weed.
Apis mellifera, probably native to the Mediterranean area,
is also naturalized in the New World. The interactions
between Apis and Tribulus in Florida probably did not
evolve as the result of foraging activities of the Honey Bee
in the Old World since 1) Apis is polytropic by nature, and
2) their distributions do not permit a long history of co-
evolution. It must have been only recently that Apis learned
to utilize Tribulus as a nectar and pollen source, especially
since Apis is capable of rapid learning (Meeuse, 1961;
1972] Tribulus cistoides — Austin 121
Adams, pers. comm., 1971). Regardless of where and when
Apis learned, pollen is frequently transported from one
Tribulus flower to the other by the visits of Apis.
An important factor in the reproduction of the plants is
the approach used by the Apis individuals. Those bees ap-
proaching from the back of the flowers merely “steal” nec-
tar without transferring pollen, while those workers enter-
ing the flowers from the front effect pollination.
The majority of species other than Apis visit the flowers
of Tribulus in the “normal” front approach. Deviations
from this are the one Lepidoptera (Melachroia) which took
nectar from the back of the flower, and occasional individ-
uals of the Dance Fly (Diptera: Empididae) species. Most
of the Dance Flies utilized the flowers in the normally ex-
pected front approach. The flies are too small, however, to
accomplish much pollination.
The largest visitor observed on the flowers was a Diptera,
probably a Syrphid fly, but no collection was made. Sev-
eral flowers were visited by the fly before attempts at
photography frightened it away. Another fly, a Tachinid
Fly, has been seen occasionally throughout the period of
observation. This species is commonly prey to yellow preda-
tory spiders in the flowers.
Hymenoptera other than Apis have been seen on the
flowers. One, a small Chloralictus bee, crawled under the
stamens and stigma (Fig. 8). Although the individual in
the photograph was not captured, others with the same
behavior have been collected. These small insects moved
among the stamens as they went from nectary to nectary.
Due to the visits among the stamen their bodies often be-
came heavily dusted with pollen (Fig. 8). In spite of this
none of them has been seen in contact with the stigma;
pollination by these small bees must be rare. An Ash-Gray
Leaf Bug (Hemiptera: Piesmatidae) exhibited much the
same behavior as the small Chloralictus bee.
Four bee species other than Apis are probably effective
in the pollination of Tribulus. Agapostemon, Chloralictus
nymphaerum, Halictus, and Lasioglossum all collect pollen
122 Rhodora [Vol. 74
and nectar in the “normal” manner, and the undersides of
their bodies are heavily dusted with pollen. A fifth species
was observed on the plants, but not captured. This large
wasp-like Hymenoptera visited the plants near Boynton
Beach and gathered only nectar on the single visit observed.
PLANT COLLECTIONS: FLORIDA: PALM BEACH COUNTY: hammock
end north of public beach in Boynton Beach, Austin 4895 (FAU);
campus of Florida Atlantic University, Austin 4422 (FAU, MO).
INSECT COLLECTIONS: HEMIPTERA: PIESMATIDAE: Austin 4422-
15, 4422-16 (FAU). LEPIDOPTERA: GEOMETRIDAE: Melanchroia
cephise (Cramer), Austin 4422-24 (FAU). DIPTERA: EMPIDIDAE:
Austin 4422-18, 4422-14 (FAU); TACHINIDAE: Austin 4422-27 (FAU).
HYMENOPTERA: APIDAE: Apis mellifera L., Austin 4422-1, 4422-2,
4422-3, 4422-17, 4422-18, 4422-19, 4422-20 (FAU); HALICTIDAE: Agapos-
temon splendens Lef.. Austin 4422-21 (FAU) ; Chloralictus nymphaerum
Robertson, Austin 4422-5, 4422-23 (FAU); Chloralictus aff. marinis
Crawf., Austin 4422-26 (FAU); Halictus sp., Austin 4422-4, 4422-28
(FAU); Lasioglossum sp., Austin 4422-6, 4422-23 (FAU).
ACKNOWLEDGEMENTS
Thanks are due Ralph M. Adams (Florida Atlantic Uni-
versity, Boca Raton, Florida) and Duncan M. Porter (Mis-
souri Botanical Garden, St. Louis, Missouri) who read the
manuscript and offered suggestions. The Lepidoptera were
determined by Thomas E. Pliske (University of Miami,
Miami, Florida); the Hymenoptera were determined with
a reference collection donated by Calaway H. Dodson (Uni-
versity of Miami, Miami, Florida). The assistance given
by these men is gratefully acknowledged. The remaining
insects were determined with Borror and DeLong (1954)
and Borror and White (1970). This study was supported
by a grant from the Florida Atlantic University Division
of Sponsored Research.
LITERATURE CITED
Borror, D. J. and D. M. DELoNG. 1964. An Introduction to the
Study of Insects, Holt, Rinehart & Winston, New York.
and R. E. WHITE. 1970. A Field Guide to the Insects
of America North of Mexico. Peterson Field Guide Series,
Houghton Mifflin Co., Boston.
FAGRI, K. and L. VAN DER PIJL. 1966. The Principles of Pollination
Ecology. Pergamon Press, New York.
1972] Tribulus cistoides — Austin 123
GRANT, V. 1950. The Flower Constancy of Bees, Bot. Rev. 16: 379-
398.
MEEUSE, B. J. D. 1961. The Story of Pollination. Ronald Press Co.,
New York.
PORTER, D. M. 1971. Notes on the Floral Glands in Tribulus (Zygo-
phyllaceae). Ann. Missouri Bot. Gard. (in press).
HERBARIUM
DEPARTMENT OF BIOLOGICAL SCIENCES
FLORIDA ATLANTIC UNIVERSITY
BOCA RATON, FLORIDA 33432
REDEFINITION OF CEDRELA OAXACENSIS
C. DC. & ROSE
C. EARLE SMITH, JR.
During the course of preparation of “A Revision of Ced-
rela (Meliaceae)” (Fieldiana: Bot. 29: 295-341. 1960) dif-
ficulties in defining inadequate collections of Spanish cedar
from Mexico and Central America led me to place many
species in synonymy. Among these, I recognized a grouping
of specimens with (for the genus) exceptionally large cap-
sules under the name Cedrela oaxacensis C. DC. & Rose.
Subsequently, Dr. Faustino Miranda kindly showed me his
well-collected specimens with very good notes which indi-
cated that the ecological disparity among the large-capsuled
cedrelas was, indeed, solidly based on the confusion of at
least two species. A large-fruited species in higher eleva-
tions from Chiapas to Panama bears several capsules in
a hanging inflorescence. Another species, bearing single
erect capsules, grows in less well watered habitats primarily
in the semi-deciduous Pacific slope forests from Mexico
southward. In Flora of Panama, VI. Family 92, Meliaceae
(Ann. Mo. Bot. Gard. 52: 60 seq. 1965), I partially untangled
this skein of my own making by recognizing C. tonduzii
C. DC. as the upland, moist-forest species with large cap-
sules in hanging inflorescences. I included in this C. sal-
vadorensis Standl. and I have no evidence at the present
time which leads me to change this opinion.
At the time that I visited the type locality of C. oaxacensis
(Monte Alban, Oaxaca) in 1957 in search of material identi-
fiable with Pringle’s collection 4802, the arborescent veg-
etation was only beginning to recover from a severe clearing
which had accompanied archaeological work at this impor-
tant pre-Conquest site. Unable to find Cedrela in the area
to corroborate the original collection, I mistakenly concluded
that the capsules accompanying the Pringle collection were
anomalous. I have since made three collections of Cedrela
oaxacensis in the Oaxaca Valley (one at Monte Alban in
124
1972] Cedrela oaxacensis 125
sprout clumps from original stumps on the site) which prove
that the foliage and capsules of Pringle 4802 are indeed
related. I am now redefining C. oaxacensis to exclude all
of the large-capsuled collections which I had previously
placed there.
Cedrela oaxacensis C. DC. & Rose. Contrib. U. S. Nat.
Herb. 5: 190. 1899.
Lectotype collection: C. G. Pringle 4802. Monte Alban,
Oaxaca. Aug., 1894. (US, B, F, MO, NY, PH).
C. discolor Blake. Proc. Biol. Soc. Wash. 33: 108. 1920.
Type collection: Palmer 184. San Ramon, Durango. Tree
to 10 m tall. Branchlets thick, lenticels small, scattered.
Leaves with, usually, 7 pairs of leaflets to 19 cm long,
7 cm wide, ovate, base rounded to obtuse to abruptly trun-
cate, rarely slightly acute, apex shortly acuminate, acute,
blade markedly pubescent beneath; rachis and petiolules
frequently pubescent, petiolules 4 mm long or less. In-
florescence moderately open, to ca. 25 cm long in flower,
to ca. 30 cm long in fruit, shorter than the leaves, puberu-
lent; bracts early deciduous; flowers pinkish, calyx to 2.5
mm deep, 5-lobed, lobes to 1.75 mm wide at base, puberu-
lent, petals 5.0 to 6.0 mm long, lanate without, puberulent
within, pinkish, but fading toward the thinner margins;
filaments fleshy, broad at point of attachment and tapering
gradually upward to the anther, connective broad, apiculum
short, obtuse; anthers ca. 1.0 mm long; ovary ovoid, gla-
brous, style to 2.0 mm long, thick, tapering very gradually
to the thick, capitate stigma. Fruit 3.5 to 4.0 em long, valves
thin, less than 1.0 mm thick, outer surface lenticellate,
outer layer peeling away from the inner as the capsule
opens; central column with wings extending only to the
margin of a distinct rostrum 6.0 to 8.0 mm in diameter, the
scars from the seed attachments 5.0 to 8.0 mm long imme-
diately beneath the rostrum between the wings.
Trees of hillslopes in the Oaxaca Valley, north and west
as far as Durango and probably to the southeast along the
Pacific slopes.
126 Rhodora [Vol. 74
Isotype collections of Pringle 4802 and Palmer 184 were
recently reexamined in the U. S. National Herbarium, con-
firming their conspecificity. Unfortunately, other material
was not then available and I was unable to place other col-
lections previously cited as C. oaxacensis. However, I have
made the following collections in Oaxaca: Rd. to Guelatao
ca. 5 km above intersection with Pan American Highway.
Stream margin. Smith & Schoenwetter 4346. Aug. 6, 1966.;
Monte Alban, on side of mound. Smith & Kitchen 4823.
July 2, 1968.; Barranca above molino. San Gabriel Etla.
Smith & S. Kitchen 4825. July 4, 1968.
I leave unresolved the disposition of C. saxatilis Rose and
C. poblensis Miranda. I have already indicated my previous
placement of C. salvadorensis Standl. with C. tonduzii. I
believe that Miranda felt that C. poblensis is a synonym
of C. salvadorensis (annotations on specimens in U. S. Na-
tional Herbarium). Neither the specimens nor the localities
clearly indicate the proper place of this species. Collections
made on the Pacific slope are more apt to be the short tree
with the single capsules. However, with increase in eleva-
tion, many areas support humid forests in which the larger,
multi-capsuled species might be found. Additional field col-
lecting with careful documentation is needed to delimit the
ranges of the large-capsuled species. I was unable to see a
specimen of Pringle 11806 (type collection of C. saxatilis)
and I make no disposition, although I suspect that it belongs
with C. oaxacensis.
I thank the staff of the U. S. National Herbarium for
their generous cooperation and my colleague, Joab Thomas,
for his patience and helpful comments.
PROFESSOR OF ANTHROPOLOGY AND BIOLOGY
UNIVERSITY OF ALABAMA
TUSCALOOSA, ALABAMA 35486
AN ABNORMAL SPECIMEN OF
LAMINARIA DIGITATA (L.) LAMOUROUX
ARTHUR C. MATHIESON, EMERY F. SWAN
AND RICHARD A. FRALICK!
The present report describes the occurrence and mor-
phology of an abnormal specimen of Laminaria digitata (L.)
Lamouroux. The plant was found (by Emery Swan) de-
tached in a deep tide pool near mean low water at Odiornes
Point, Rye, New Hampshire, U.S.A. (70°42’42” W longi-
tude, 43°03’33” N latitude) on May 16, 1971. Figures 1
and 2 illustrate the general morphology of the plant; a
single stipe and two attached blades are evident. The overall
length of the plant was 208 cm. The maximum dimensions
of the fronds were 120X 98 and 76 X 74 cm respectively,
exclusive of the stipe. The larger blade was darker in
color, healthier in appearance, and it seemed to be more
actively growing, as evidenced by its segmentation (Fig. 2),
than the smaller frond.
Laminaria digitata is a common plant in the vicinity of
Odiornes Point. It grows abundantly on solid rocks from
0 to 14 m below mean low water, with maximum biomass
occurring from -2 to 10 m (Mathieson, Hehre and Rey-
nolds, in press). It is speculated that the plant broke off
its holdfast and then regenerated a second frond.
Recent studies by Markham (1968) demonstrated exten-
sive regenerative capacities in Laminaria sinclairii (Har-
vey) Farlow, Anderson e£ Eaton. Detached portions of its
haptera, as short as 2.5 mm in length, were capable of de-
veloping into entire plants. Laminaria digitata also exhibits
extensive regenerative capacities (unpublished data, A.
Mathieson), for new blades can be produced from intact
stipes if they are injured or destroyed. Similar regenera-
tive capacities are probably evident in healthy, pelagic
specimens. Figure 3 shows a conspicuous swelling of the
'Published with the approval of the Director of the New Hampshire
Agricultural Experiment Station as Scientific Contribution Number
127
128 Rhodora [Vol. 74
t $ RINI
LLUTETELECLEETERLETULELTELTLELTIEE]
1972] Laminaria — Mathieson, Swan and Fralick 129
stipe in the abnormal specimen; it probably represents the
site of regeneration of the second frond. The swelling is
several cm below the transition zone —the area of “ac-
tive” cell division. Thus, regeneration occurred in an area
presumably restricted to elongation and differentiation of
cells. The older appearance of the smaller frond suggests
that it is the original one. However, in order to preserve
the intact specimen a critical examination (section) of
the stipe was not made.
Burrows (1956, 1958) reported extensive populations of
loose-lying algae (particularly Chorda filum, Desmarestia
aculeata, Laminaria saccharina and Saccorhiza polyschides)
from the subtidal zones at Port Erin Bay, Isle of Man —
particularly after periods of calm weather and high insola-
tion. Most of the plants were initially attached. Extensive
SCUBA observations near Odiornes Point have never shown
quantities of loose-lying L. digitata or any other algae —
perhaps because of the exposed nature of the site. The
accidental deposition of the specimen in a deep tide pool
may have provided a relatively stable habitat and allowed
regeneration to occur. According to Burrows (1958) mem-
bers of the loose-lying populations may serve as a means of
restocking habitats devoid of reproductive plants. No re-
productive sori were found on the specimen of L. digitata.
The vegetative stae of the plant may have been due to the
time of its collection, for L. digitata exhibits limited re-
production during the spring in New England (Mathieson,
Hehre and Reynolds, in press). It should also be noted that
several pelagic algae (e.g. Sargassum, Focus and Gracil-
aria) are reported (Fritsch, 1959) to have limited repro-
ductive capacities.
Figure 1. Intact specimen of Laminaria digitata, with a single
stipe and two attached blades.
Figure 2. Close up of stipe and fronds, showing darker and
healthier appearance of larger (left) frond. Active segmentation of
the larger frond is also visible.
Figure 3. Conspicuous swelling of the stipe, that may represent
the site of regeneration of the second frond.
130 Rhodora [Vol. 74
According to van Overbeek (1940 a, b) auxins may be
found in Macrocystis pyrifera and other known brown
algae. If this were true for L. digitata, then the loss of
polarity, due to a pelagic state, may explain the regenera-
tion of a second frond rather than a holdfast. Extensive
regeneration and the loss of polarity must also occur in
other unattached algae that produce spherical morphologies,
such as Cladophora holsatica Kiitzing described by Smith
(1950) and the “beach form” of Ascophyllum nodosum (L.)
Le Jolis ecad Mackaii (Turner) Cotton reported by Gibb
(1957).
In conclusion the occurrence of an abnormal specimen of
L. digitata poses a variety of questions concerning the
growth and ecology of the species. Additional studies should
be carried out to determine the growth, reproduction and
ecological significance of pelagic specimens of L. digitata.
LITERATURE CITED
Burrows, E. M. 1956. A preliminary survey of the sublittoral algae
of Port Erin Bay, Isle of Man. Phycol. Bull. 4: 14-15.
, 1958. Sublittoral algae populattions in Port Erin
Bay, Isle of Man. J. Mar. Biol. Assoc. U.K. 37: 687-703.
FRITSCH, F. E. 1959. The structure and reproduction of the algae.
Vol. II. Cambridge University Press, 939 pp.
GIBB, D. C. 1957. The free-living forms of Ascophyllum nodosum
(L.) Le Jol. J. Ecol. 45: 49-83.
MARKHAM, J. W. 1968. Studies on the haptera of Laminaria sin-
clairii (Harvey) Farlow, Anderson et Eaton. Syesis 1: 125-131.
MATHIESON, A. C., HEHRE, E. J. and ReyNoups, N. B. Investigations
of New England marine algae I: A floristic and descriptive
ecological study of the marine algae at Jaffrey Point, New
Hampshire, Nova Hedwigia (in press).
OVERBEEK VAN, F. 1940a. Auxin in marine algae. Plant Physiol.
15: 291-299.
SMITH, G. M. 1950. The fresh-water algae of the United States,
McGraw-Hill Book Co., N. Y., 719 pp.
JACKSON ESTUARINE LABORATORY
OF THE UNIVERSITY OF NEW HAMPSHIRE
DURHAM, N.H., U. S. A. 03824
RANGE EXTENSIONS OF VASCULAR AQUATIC
PLANTS IN NEW ENGLAND
C. BARRE HELLQUIST
During 1970 and 1971 extensive collecting of vascular
aquatic plants was conducted in New England. New
records for some states as well as interesting extensions
of range were noted. In New England the aquatic flora
has been neglected during the past twenty years, so these
ranges may have expanded or may have been overlooked.
Such botanists as M. L. Fernald and A. S. Pease did exten-
sive collecting in the time span of 1920 to 1955. Since then
no single person has devoted much time to the aquatics in
New England except William Countryman who has done
extensive collecting in Vermont.
The plants collected and observed are discussed below.
Representative specimens have been deposited in the
author’s personal herbarium as well as those of Boston
State College, the University of New Hampshire and the
New England Botanical Club.
Potamogeton filiformis Pers. var. borealis (Raf.) St. John
Both Potamogeton filiformis and its variety borealis are
found in New England only in the extreme north. A plant
of calcareous waters, (St. John, 1916), this plant has been
collected from several areas in Aroostook County, Maine,
and in northern Vermont (Seymour, 1969). A specimen
in the University of New Hampshire herbarium (S B.
Krochmal, 142a) from Piermont, New Hampshire, Grafton
County was examined by this author and keyed out as
P. filiformis Pers. var. Macounii Morong, which has never
been reported from New England. This identification is
questionable and more specimens of this plant would be
desirable for absolute identification. In October 1970,
several specimens of the variety borealis were collected for
the first time in New Hampshire in Lombard Pond, Cole-
brook, Cóos County (Hellquist 93 and 94). This pond has
191
132 Rhodora [Vol. 74
the highest alkalinity of any in the state with a Methyl
Orange ppm. reading of 90 (Newell, 1960). Of all the
ponds in New Hampshire this is one of the few where this
Potamogeton would find the requirements of an alkaline
body of water in the northern portion of the state. Lime
Pond in Colebrook and Ladd Pond in West Stewartstown
might also harbor this species.
Potamogeton crispus L.
This has increased its range extensively in the past fifty
years. Records in the Gray Herbarium and the New Eng-
land Botanical Club Herbarium include very few reports
of the species. Seymour (1969) noted this plant from
Middlesex County in Massachusetts, Chittenden County in
Vermont, Providence County in Rhode Island and a number
of areas in Connecticut. Collections of this plant were made
from a number of ponds and lakes in Massachusetts and
a couple from northwestern Connecticut. Ogden (1943)
noted it from Spy Pond in Arlington, Massachusetts, Fresh
Pond in Cambridge, Massachusetts and the Sudbury River
in Concord, Massachusetts. All of these are in Middlesex
County. In Connecticut, the Housatonic River in Litchfield
County was the only area noted.
From the recent collections by the present author it is
evident that Potamogeton crispus L. has increased its
range extensively, especially in the hard water regions of
Berkshire County, Massachusetts. This plant was ex-
tremely common in the Stockbridge Bowl, Stockbridge,
Massachusetts where it was the most abundant pondweed
and along with Myriophyllum spicatum L. var. exalbescens
(Fernald) Jepson made up about 80% of the vascular vege-
tation. This plant was also extremely common at Lake Buel
in Monterey and New Marlboro, Massachusetts, Mill Pond,
South Egremont, Massachusetts and the Mill Pond in Shef-
field, Massachusetts. A new record in Middlesex County
for this species is Fisk Pond, Framingham, Massachusetts.
From Connecticut it was found in Mudge Pond and Indian
Lake, Sharon, Litchfield County. In all the above mentioned
96
1972] Aquatic Plants — Hellquist 133
areas except Stockbridge Bowl it was not the predominant
species but was nevertheless common. This species has
been known to be more tolerant of polluted waters (Fassett,
1966). After observing the water where collections were
made this statement would seem to hold true. The ponds
from western Massachusetts and Connecticut were in alka-
line regions where this plant does well (Moyle, 1945). This
plant should be looked for in almost all hard water regions
of New England especially where pollution may occur.
Potamogeton strictifolius Ar. Benn. Var. rutiloides Fern.
This is one of the narrow-leaved pondweeds of the section
Pusilli and is characteristic of basic or alkaline waters of
the northern United States and Canada (Fernald, 1932).
Fernald at that time noted it from Lake Champlain, Ver-
mont. Seymour (1963) recorded the variety rutiloides
from three other areas of western Vermont not far from
Lake Champlain as well as from Berkshire County, Massa-
chusetts.
A new eastern record for Potamogeton strictifolius Ar.
Benn. Var. rutiloides Fern. in New England is the Mill
Pond in Windsor, Vermont, Windsor County. It was
abundant in some of the shallow areas on the eastern side
of the pond. This plant, upon first observation, was mis-
taken by the writer for Potamogeton Friesii Rupr. with
which it may be confused easily. Potamogeton strictifolius
and its variety generally have three veins, but this speci-
men had three to five. Fernald (1932) noted that P. stricti-
folius may rarely have five veins. The Mill Pond has the
typical vegetation of the more alkaline lakes including
Najas flexilis (Willd.) Rostk. & Schmidt., Vallisneria
americana Michx. and Potamogeton pectinatus L. (Moyle,
1945).
Potamogeton longiligulatus Fern.
One of the rarer pondweeds in the United States as well
as New England is P. longiligulatus. Only two specimens
have been reported and both were from northwestern Con-
necticut. Fernald (1932) noted it from Twin Lakes, Salis-
134 Rhodora [Vol. 74
bury, Connecticut, where it was collected by C. H. Bissell
in 1906. A second report was from Indian Lake in Sharon,
Connecticut, (Seymour, 1969). This specimen was de-
posited in the New England Botanical Club Herbarium but
after careful study by the writer it was found to be a very
narrow-leaved form of Potamogeton zosteriformis Fernald.
The second actual record for this specimen comes from
Evarts’ Pond (Lake Runnymede) in Windsor, Vermont,
Windsor County. Evarts’ Pond is a privately owned pond
which is of interest in that it was one of the ponds where
Potamogeton Hillii Morong. had been reported in 1875
(Fernald, 1932). More recent collections of P. Hillii from
Evarts’ Pond in the late thirties are in the New England
Botanical Club Herbarium. In an attempt to find Pota-
mogeton Hillii, permission was received from the owners
to collect in the pond.
A narrow-leaved pondweed believed to be Potamogeton
Hillii was collected by the author at the northern portion
of the northeast bay and on the southwestern end of the
pond. In both areas it was found in water approximately
three feet deep. Upon examining these collections care-
fully, it was determined that the two rare Potamogetons,
Hillii and longiligulatus were both found. Fernald (1932)
commented on much confusion in identifying these two
species. Both have sharply acute, cuspidate leaves. Pota-
mogeton Hillii has three veins while P. longiligulatus has
five to nine. Both specimens were in flower at this time
so they had the same initial appearance.
In talking with Mr. Evarts, one of the owners, it was
learned that the state of the pond had deteriorated over the
past two summers. The water previously had been very
clear, but now becomes cloudy as the water warms up in
the summer. He stated that corn has been planted along
the north side of the pond, and is fertilized extensively with
chemicals. This most likely has led to the turbid water
with its poor light penetration so it was rocommended that
the corn crop be discontinued in 1972. Hopefully this would
clear up the problem. Since this pond has no inlets, but
1972] Aquatic Plants — Hellquist 135
is spring-fed, it shouldn’t take long for the water to become
clear again. It would be a shame for the ecology of this
pond to be so disturbed that these two valuable pondweeds
would be destroyed. Nymphaea tuberosa Paine was also
discovered here and will be discussed later in this paper.
Potamogeton gemmiparus Robbins.
This uncommon narrow-leaved pondweed has been re-
ported from only four New England states: Maine, Massa-
chusetts, Connecticut and Rhode Island (Fernald, 1932).
Since it is from eastern New England it is found in more
acid waters. This plant had never been reported in New
Hampshire, where it should occur. Seymour (1969) records
it as near as Fryeburg, Maine, where it is found in the
Saco River.
Two stations of this plant were found by the author in
Carroll County, New Hampshire, both in the Saco River
drainage. One station where it was somewhat rare was
the southern end of Conway Lake in Eaton, New Hamp-
shire. Here it was found with other acid water plants such
as Potamogeton Berchtoldii Fieber, P. Berchtoldii Fieber,
var. tenuissimus (Mert. & Koch) Fern., P. epihydrus Raf.
var. ramosus (Peck) House, P. gramineus L. var. grami-
neus and P. capillaceus. Bearcamp Pond in Sandwich, New
Hampshire was the other site of collection. Here the speci-
men was collected at the inlet of the Bearcamp River.
Again, it was not plentiful. Potamogeton gemmiparus was
growing along with P. natans L., P. gramineus L. var.
myriophyllus Robbins. P. gramineus L. var. gramineus,
P. confervoides, Reichenb., P. capillaceus Poir. and P.
Spirillus Tuckerm.
Egeria densa Planchon
Egeria has been reported from three different towns in
New England — two in Massachusetts and one in Vermont
(Seymour, 1969). The reports from Massachusetts were
from Abington in Plymouth County and Quincy in Norfolk
County. In Vermont it was collected from Townshend,
Windham County. It was observed by the writer to be very
136 Rhodora [Vol. 74
common in Hemenway Pond, Milton, Massachusetts, Nor-
folk County, near the Boston city limits where it was
growing with Elodea Nuttallii (Planch.) St. John, Pota-
mogeton epihydrus Raf. var. ramosus (Peck) House,
Myriophyllum humile (raf.) Morong and Utricularia
geminiscapa Benj. This plant was introduced and natu-
ralized from South America and is one to be looked for
around cities because it is being sold in pet shops.
Wolffia columbiana Karst.
This plant is continuing to spread its range to the north
in New Hampshire. Previously reported from Hillsborough
County, (Countryman, 1968) and Rockingham County
(Colt, et. al., 1971) it was discovered by the author in
Durham, New Hampshire, Strafford County. In Durham,
it was reported from two new locations — the Mill Pond
and Beard’s Creek Pond. This plant was not observed in
either place in 1970, but was very abundant at Beard’s
Creek Pond and common at the Mill Pond in 1971. Wolfia
punctata Griseb., which usually occurs with the former,
was not found but should be looked for in the future.
In talking with Dr. Albion Hodgdon at the University
of New Hampshire in Durham, it was agreed that this
plant probably is rapidly extending its range. One explana-
tion for the rapid spread may be the passage of ducks in
the region moving between ponds. Wolffia would stick to
their bodies and be carried around easily. Also anyone
transferring a boat from ponds could easily introduce the
plants into new bodies of water.
Nymphaea tuberosa Paine
Seymour (1969) recorded this plant from six stations
in western Vermont, Winchester, Massachusetts, and three
townships in Connecticut. This species has been found by
the writer to be quite abundant. In Vermont, a new eastern
record for the state is herein reported from Evarts’ Pond,
Windsor County. Here it was the only member of the
Nymphaeaceae and was found extensively around the pond.
The first two records for this plant in New Hampshire are
1972] Aquatic Plants — Hellquist 137
Lake Kanasatka, Moultonboro, Carroll County and Beard’s
Creek Pond, Durham, Strafford County. This pond lily was
found at the southern end of Lake Kanasatka near the gift
shop “The Horse of Another Color" along Route 25. It
appears that this plant was introduced here some time ago,
due to its localized range at the southern end of the lake.
It was found with other members of the same family in-
cluding Nymphaea odorata Ait., Nuphar variegatum
Engelm., and Brasenia Schreberi Gmel. This is a lake
with a higher alkalinity than most other lakes in Carroll
County as shown by the presence of Potamogeton zosteri-
formis Fernald and P. praelongus Wulfen. At Beard’s
Creek in Durham Nymphaea tuberosa Paine was found
near Route 4 where it has spread throughout the southern
end of the pond. The water is probably in much the same
condition as Taylor River in Hampton, New Hampshire
(Colt, et al., 1971) since this also is a tidal estuary which
has been blocked off by a dam. Much salt in the bottom
might tend to raise the alkalinity of the water. Other plants
of interest growing along with N. tuberosa here are Myrio-
phyllum verticillatum L. var. pectinatum Wallr. and Elodea
Nuttallii (Planch.).
In Massachusetts this plant was discovered by the writer
in three locations, all within Middlesex County. These were
the Sudbury and Concord Rivers in Concord, Massachusetts
and a small pond by Upper Mystic Lake in Winchester,
Massachusetts which is near the previously reported site.
Mr. Richard Eaton of Lincoln, Massachusetts told the
author in 1971 that Nymphaea odorata used to be very com-
mon in these rivers and was almost completely destroyed
due to the pollution. Nymphaea tuberosa is found in the
highly productive lakes of the midwest and seems to be
more tolerant of polluted waters. In the Sudbury River
Nymphaea odorata was found in limited numbers while
N. tuberosa was more common. The area observed in the
Concord River had Nymphaea tuberosa as the only water
lily and it was found at only one location.
The location at Winchester, Massachusetts was a small
138 Rhodora [Vol. 74
pond at the recreation area on the east side of Upper Mystic
Lake. It appears that this lily was planted here as it does
not extend into Upper Mystic Lake which connects with
this pond.
This plant may be more common than previously ob-
served. Many of the specimens reported as Nymphaea
odorata Ait. var. gigantea Tricker may actually be N. tu-
berosa. The leaves of N. tuberosa are generally larger
and green underneath, or with only a slight purplish tinge.
Probably the best identifying characteristic of this plant
is the large green petiole with brown stripes as opposed
to the purple petiole of Nymphaea odorata. Fernald (1950)
noted that the flowers are odorless or barely fragrant which
may be misleading. The flowers have a noticeable odor,
but when compared with N. odorata they are less fragrant.
Another interesting feature found on a number of speci-
mens was the presence of pubescence on the petiole near
the blade. In some this was so marked that the striping in
the petiole was obscured. Other characteristics which aid
in identification are the more rounded petals and the ten-
dency of the leaves to become elevated above the water by
as much as one foot.
Nymphaea alba L. forma rosea Hartm.
A European species, this pond lily has not been reported
previously from New England. It was found by the writer
to be relatively abundant in Winkley Pond, Barrington,
New Hampshire, Strafford County. It occurs around the
perimeter of the Pond where it grows with Nymphaea
odorata and a possible hybrid between N. odorata and N.
alba f. rosea which has the pointed more numerous petals
and is a light pink. Winkley Pond is an acid body of water
in a semi-bog condition with many ericaceous plants. Of
particular interest as an associated plant is Potamogeton
confervoides Reichenb. which is extremely common there.
Nymphaea alba f. rosea is characterized by 12-24 rounded
deep pink petals (Fernald, 1950). The leaves are similar
to N. odorata with deep purple undersides. The following
1972] Aquatic Plants — Hellquist 139
observations were made on N. alba f. rosea: many of the
petioles were pubescent as seen in N. tuberosa and the
leaves were elevated above the surface of the water along
the edge of the pond. Because of the attractive blossom,
this water lily may have been planted more extensively
than observed. Many specimens labeled Nymphaea odorata
Ait. forma rubra Guillon may actually be N. alba f. rosea.
Myriophyllum alterniflorum D. C.
Except for the present report M. alterniflorum has been
found in New England only from numerous areas in
Vermont and Maine and from Cóos County in northern
New Hampshire (Seymour, 1969). Last year the first
record of the species south of Cóos County in New Hamp-
shire was noted in Upper Danforth Pond, town of Freedom,
Carroll County, (Hellquist, 1971). Two other locations for
this plant have also been found in Carroll County by the
writer. One is from Cooks Pond in Madison, New Hamp-
shire where it was scarce. Isolated plants were found in
this acid-water pond which also included Potamogeton
confervoides Reichenb. (Hodgdon, et. al, 1946). Lake
Kanasatka, Moultonboro, New Hampshire was the other
station. The Myriophyllum was found at the south end of
the lake near the previously reported Nymphaea tuberosa
Paine. As more extensive studies are carried out in central
New Hampshire, this plant should prove to be more
abundant.
Myriophyllum heterophyllum Michx.
This plant has been quite uncommon in New Engiand
until recently if the few herbarium specimens are a fair
indication. It had been recorded from Bridgeport, Con-
necticut, Uxbridge and Sutton, Massachusetts (Seymour,
1969). A specimen from Sandy Pond, Ayer, Massachu-
setts, collected by Edward Richardson in 1940 is on deposit
in the University of Massachusetts herbarium.
In southern Worcester and Middlesex Counties, Massa-
chusetts, the writer found M. heterophyllum to be extremely
common in a number of ponds and lakes, where: it forms
140 Rhodora [Vol. 74
large beds around the shore. It has been found in White-
hall Reservoir and North Pond in Hopkinton, Middlesex
County; Pratt Pond, Upton; Silver Lake, Grafton; and
Hopedale Pond, Hopedale, Worcester County. Houghton’s
Pond (Hoosiwhisick Pond), Canton, Norfolk County, Mill
Pond, Wareham, and Agawam River, Wareham, Plymouth
County are other areas where this plant was found to be
abundant. Fassett (1966) noted this plant to range from
Virginia to Florida, west to Ontario and Minnesota, so it
appears that this specimen is spreading rapidly in southern
New England. One cause of this recent and rapid spread
may be the fact that the plant is sold extensively in pet
shops for aquarium use. It may be discarded into ponds
and streams as has Egeria densa Planchon.
LITERATURE CITED
Corr, L. C., JR., C. BARRE HELLQUIST and W. J. L. ZUBRIN. 1971. An
Interesting Association of Rare Aquatic Plants from New Hamp-
shire. Rhodora 73: 296-297.
CoUNTRYMAN, W. D. 1968. Wolfia in New Hampshire. Rhodora
70: 491.
Fassett, NoRMAN C. 1966. A Manual of Aquatic Plants. ( Appen-
dix by E. C. Ogden). University of Wisconsin Press. Madison,
Wisconsin. 405 p.
FERNALD, M. L. 1932. The Linear-leaved North American Species
of Potamogeton, section Axillares. Memoirs of the American
Academy of Arts and Sciences Vol. 17, Part I. 183 p.
1950. Gray's Manual of Botany, eighth edition.
American Book Company, New York. 1632 p.
HELLQUIST, C. BARRE, 1971. Vascular Flora of Ossipee Lake, New
Hampshire and its Shoreline. Rhodora 73: 249-261.
Hopepon, A. R., P. GIGUERE, S. B. KROCKMAL and A. RIEL. 1954.
New Potamogeton Records in New Hampshire. Rhodora 54:
237-246.
MovLE, Jonn B. 1946. Some Chemical factors Influencing the Dis-
tribution of Aquatic Plants in Minnesota. Am. Midl. Nat. 34:
402-420.
NEWELL, ARTHUR E. 1960. Biological Survey of the Lakes and
Ponds in Coos, Grafton, and Carroll Counties. Survey Report
no. 8a.New Hampshire Fish and Game Department. 297 p.
OGDEN, E. C. 1943. The Broad Leaved Species of Potamogeton of
North America North of Mexico. Rhodora 45: 57-105, 119-163,
171-214.
1972] Triphora trianthophora — Eastman 141
SEYMOUR, FRANK C. 1969. The Flora of New England. The Charles
Tuttle Co., Rutland, Vermont. 596 p.
St. JoHN, HaROLD. A Revision of the North American Species of
Potamogeton of the Section Coleophylli. Rhodora 18: 121-138.
BIOLOGY DEPARTMENT
BOSTON STATE COLLEGE
625 HUNTINGTON AVENUE
BOSTON, MASSACHUSETTS 02115
A SECOND OCCURRENCE FOR TRIPHORA TRIAN-
THOPHORA (SW.) RYDB. IN MAINE. Another station of
the orchid, Triphora trianthophora, the second to be found
in the last few years, was discovered on August 31, 1971
at Evans Notch, Batchelders Grant, Oxford County by C.
Paul Wight and me. This location is about three miles in
a direct line from the Stow station (Rhodora 71: 509,
1969). This new station consists of many individuals grow-
ing on a wooded hillside in depressions which are filled with
thickly matted beech leaves. It is difficult to estimate how
many of the orchids occur at this station, as they tend to
grow singly, spreading out over a large area, unlike the
Stow station, where the orchids were growing in large
groupings. We were fortunate in finding many of the plants
still in flower for the majority had already lost their blos-
soms. Other interesting plants were found among the
Triphoras, namely, Corallorhiza maculata, Conopholis amer-
icana, Epifagus virginiana, and members of the Botrychium
group.
Specimens have been deposited at the Herbaria of the
University of Maine and the New England Botanical Club.
LESLEY M. EASTMAN
OLD ORCHARD BEACH, MAINE 04064
ILLINOIS FIELD AND HERBARIUM STUDIES
ROBERT H. MOHLENBROCK AND DAN K. EVANS
Continued field and herbarium studies of Illinois plants
have resulted in the discovery of several unreported or pre-
viously undocumented taxa for the state. These new addi-
tions to the flora are reported in this paper, in addition to
locality records for other rare taxa in southern Illinois.
All specimens cited are in the herbarium of Southern Ili-
nois University (SIU), unless otherwise indicated.
TAXA NEW TO ILLINOIS
Cyperus iria L., an introduced southern sedge, was known
to range in North America from Virginia east to southeast-
ern Missouri, southwest to Texas, and east to Florida. Now
Illinois may be added to the range. In Illinois, this species
may be found sparingly in the wet meadows at Horseshoe
Lake, Alexander County.
Of the twenty-one taxa of Cyperus found in Illinois, this
species may be most likely confused with those in the fol-
lowing key.
a. Scales 1.0-1.5 mm long; achenes 0.8-1.0 mm long. b.
b. Achenes white; spikelets about 1 mm broad; scales
closely arranged ................ C. erythrorhizos
b. Achenes brown or black; spikelets about 1.5 mm
broad; scales somewhat remote from each other .
a C. iria
a. Scales 1.5-4.5 mm long; achenes 1.0-2.8 mm long. c.
c. Scales remote, the tip of one just reaching the base
of the one above; base of the achene enclosed in a
corky sheath ............ sees C. engelmannit
c. Scales approximate and overlapping ; no corky sheath
present at base of achene. d.
d. Some or all of the mature spikelets reflexed;
spikelets subterete .......... C. lancastriensis
d. None of the spikelets (except sometimes the low-
est pair) reflexed; spikelets flattened. e.
142
1972] Illinois Studies —- Mohlenbrock & Evans 143
e. Rhizomes scaly and usually ending in a tuber;
scaletips slightly spreading .... C. esculentus
e. Rhizomes absent or merely hard and corm-
like; scaletips appressed. f.
f. Plants: annual without rhizomes; scales
ferrugineous or golden-brown, 1.7-3.0 mm
long; achenes obovoid-oblongoid . ......
M ar ee: C. ferruginescens
f. Plants perennial with hard, corm-like
bases; scales stramineous, 3.5-5.0 mm
long; achenes linear ..... .. C. strigosus
COLLECTION DATA: Alexander County: Horseshoe Lake,
wet meadows, July 30, 1968, E. Estes s.n.
Ulmus procera Salisb. Several specimens of the English
elm oecur in the vicinity of an abandoned farm along the
“Giant City blacktop” about 1 1/4 miles north of Giant City
State Park. The colony is apparently spreading by suckers.
This elm is distinguished by its scabrous upper leaf surface
and the presence of corky wings on the branches. COLLEC-
TION DATA: Jackson County: along blacktop road between
Carbondale and Giant City State Park, June 26, 1970, R.
H. Mohlenbrock 19256.
Amaranthus caudatus L. This often cultivated Tassel
Flower was collected from a spontaneous colony in waste
ground. Although it has undoubtedly escaped elsewhere
in Illinois, this is the first report of such an occurrence. COL-
LECTION DATA: Jackson County: waste ground, Oakland
Street, Carbondale, July 29, 1970, D. Evans 1112.
Ranunculus repens L. var. pleniflorus Fern. The garden
form of the creeping buttercup, with most of its stamens
transformed into petals, was collected in a grassy area at
the edge of the campus of Southern Illinois University,
Carbondale. COLLECTION DATA: Jackson County: Carbon-
dale, August 14, 1967, R. H. Hohlenbrock s.n.
Nigella damascena L. The Love-in-a-mist, a member of
the Ranunculaceae, is occasionally grown as a flower garden
ornamental and is rarely escaped from cultivation. How-
ever, a few specimens were observed at the edge of an
144 Rhodora [Vol. 74
abandoned lot at the rear of 411 South 20 Street, Murphys-
boro, during the summer of 1968. The deeply divided in-
volucre which subtends the blue flowers accounts for the
common name. COLLECTION DATA: Jackson County: lot,
rear of 411 S. 20 Street, Murphysboro, July 27, 1968, R.
H. Mohlenbrock 16121.
Rorippa islandica Oeder var. islandica. This variety of
Yellow Cress, new to the Illinois flora, was collected from
mud and sand flats along the Mississippi River. In that
habitat it occurs with the more abundant variety fernald-
iana and a similar second species, R. sessiliflora.
Although normally found in the northeastern United
States, Quebec, and Greenland, Steyermark (1960) reports
a station in Harrison County, Missouri. COLLECTION DATA:
Jackson County: Mississippi River mud and sand flats,
Grand Tower, November 1, 1969, D. Evans 656.
Rubus procerus P. J. Muell., a native of Europe, is re-
ported by Fernald (1950) as naturalized in the United
States from Delaware to Virginia. Steyermark (19623) re-
ports a Palmer collection from Newton County, Missouri.
The authors collected this Illinois record from a roadside
habitat in Randolph County, where it grows in abundance.
The large drupelets are most edible. Among the many tax-
onomically difficult species of Rubus one encounters in Illi-
nois, only three have leaflets with the underside whitened.
These may be identified in the field by the following key:
a. Stems glaucous . .. ............... R. occidentalis
a. Stems not glaucous. b.
b. Mature fruit red; prickles small, bristle-like or ab-
sent; leaves ovate-lanceolate ... ................
NENNEN R. idaeus var. strigosus
b. Mature fruit black; prickles abundant, flat with a
broad base; leaves broadly ovate ...... R. procerus
COLLECTION DATA: Randolph County: roadside, 10 miles
south of Chester, Illinois, July 21, 1970, R. Mohlenbrock &
D. Evans 1113.
Croton texensis (Klotzsch) Muell. Jones and Fuller
(1955) report that although Ferguson (1901) cites a Hall
1972] Illinois Studies — Mohlenbrock & Evans 145
collection of this species from Athens, Illinois, it is excluded
from the Illinois flora since no specimen could be found to
verify this report. However, a specimen of this species,
collected by Hall from Athens, is on deposit in the her-
barium of the Missouri Botanical Garden. This poisonous
species normally ranges from South Dakota, south to Mex-
ico, with introductions known from western Missouri, Ala-
bama, and New England. COLLECTION DATA: Menard
County Athens, Hall 514.
Hibiscus syriacus L. The Rose-of-Sharon is a small tree
often planted, particularly in the past, as an ornamental.
Occasionally specimens may be found around abandoned
homesteads, although they generally do not appear to be
spreading. A small grouping of specimens occurs along an
abandoned road on the Little Grassy camp facilities of
Southern Illinois University. At this site, there is indication
that the plants have multiplied since their original planting
some forty years ago. COLLECTION DATA: Jackson County:
along dirt road, Little Grassy camp, Southern Illinois Uni-
versity, August 14, 1969, R. H. Mohlenbrock s.n.
Ascyrum hypericoides L. var. hypericoides. The range
of the typical variety of A. hypericoides generally lies to
the south and east of Illinois. On the other hand, var. multi-
caule (Michx.) Fern., is a relatively common taxa in dry
woods, on slopes, and along ridges across the southern tip
of Illinois. Variety hypericoides has never been reported
from Illinois although there is a specimen of it, housed in
the herbarium of the Missouri Botanical Garden, annotated
by Dr. Preston Adams, the most recent monographer of
the group. It is interesting to note, however, that Adams
fails to attribute this variety to Illinois in his monograph
(1957). Variety hypericoides differs from var. multicaule
by its more ascending habit and by its narrower leaves
which attain widths of only 4(-5) mm. COLLECTION DATA:
Hancock County: Augusta, July, 1842, S. B. Mead s.n.
Hypericum punctatum Lam. var. pseudomaculatum
(Bush) Fern. When a typical specimen of the Large Spotted
St. Johns-wort is encountered, the first reaction is to con-
146 Rhodora [Vol. 74
sider it a distinct species since the large flowers are so
strikingly different from the similar but smaller-flowered
var. punctatum. However, no other characters seem fo ex-
ist which can reliably distinguish var. pseudomaculatum
from var. punctatum. A frequently used character to sep-
arate these two entities is the uppermost leaves obtuse in
var. punctatum and acute in var. pseudomaculatum. Al-
though all specimens which we have observed of var. pseud-
omaculatum do have acute upper leaves, a few specimens
of var. punctatum also have acute upper leaves. Jones
(1963) and Jones, et al. (1955) attribute Hypericum pseud-
omaculatum to Illinois, but do not indicate localities. Since
we have never before observed a specimen of this plant
from Illinois, we are documenting its occurrence in Illinois
by the following collection. COLLECTION DATA: Jackson
County: dry woods, July 26, 1964, J. Ozment & DeFilipps
1726.
Hypericum densiflorum Pursh. This shrubby St. Johns-
wort previously was known from southern Missouri, but
the collection cited below is apparently the first from Illi-
nois. This species closely resembles H. lobocarpum from
which it differs only by the presence of three styles and a
3-celled capsule. Since these seem to be fundamental dif-
ferences to us, we are treating the two as separate species.
Other botanists prefer to treat H. lobocarpum as a variety
of H. densiflorum. Although Fernald (1950) indicates that
the tip of the leaf may be used to separate H. densiflorum
from H. lobocarpum, we do not find this character to be
reliable. The other shrubby St. Johns-wort in Illinois with
three styles is H. spathulatum, a species usually larger in
all respects. COLLECTION DATA: Alexander County: swampy
woods near Miller City, August 28, 1964, J. Ozment & R.
DeFilipps 2712.
Hypericum sphaerocarpum Michx. var. turgidum (Small)
Svenson. This southern variant of H. sphaerocarpum is
recognized by its narrow, revolute leaves which have no
apparent lateral nerves. In some cases, it looks more like
H. denticulatum Walt. or H. dolabriforme Vent. than H.
1972] Illinois Studies — Mohlenbrock & Evans 147
sphaerocarpum var. sphaerocarpum. It is distinguished
from H. denticulatum by its united styles and from H.
dolabriforme (which does not occur in Illinois) by its
shorter sepals and somewhat smaller flowers. COLLECTION
DATA: Jackson County: railroad prairie, 6 miles north of
Murphysboro, June 13, 1953, R. H. Mohlenbrock s.n.; Ma-
coupin County: railroad prairie southwest of Carlinville,
August 15, 1968, J. White 335; Monroe County: limestone
bluffs, one mile south of Fults, August 7, 1962, J. Ozment
s.n.
Hypericum sphacrocarpum Michx. var. turgidum (Small)
(1945) indicated that the Northern St. Johns-wort was
known from Illinois, we are unaware of any Illinois collec-
tions of this species prior to the 1964 collection cited here.
Since the overall range of H. boreale reaches from New-
foundland across to eastern Iowa, its occurrence in north-
ern Illinois is not unexpected. 'This species is similar in
appearance to the other small-flowered species of Hyperi-
cum in Illinois, but differs in the bracteal leaves which are
similar to the cauline leaves in shape and size. COLLECTION
DATA: Iroquois County: Iroquois County Conservation
Area, August 17, 1964, J. Ozment & R. DeFilipps 2298.
Aesculus glabra Willd. var. leucodermis Sarg. This
variety differs from typical A. glabra in its distinctly
whitened lower leaf surfaces and its whitish bark. At its
only Illinois location, it grows in a mesophytic woods. COL-
LECTION DATA: Jackson County: along Kinkaid Creek, three
miles south of Ava, July 14, 1970, R. H. Mohlenbrock s.n.
Acer rubrum L. f. tomentosum (Desf.) Dansereau. This
form is enigmatic in that the densely tomentose lower leaf
surface of this upland taxon is identical with that of the
swamp-inhabiting A. rubrum L. var. drummondii (H. & A.)
Sarg. Only the fruits can be used reliably to separate these
two taxa. Forma tomentosum has samaras up to 2.5 cm
long, while var. drummondii has samaras over 3.0 cm long.
COLLECTION DATA: Union County: upland woods, Pine
Hills, July 3, 1970, R. H. Mohlenbrock s.n.
Euonymus fortunei (Turcz. Hand.-Maz. This climbing
148 Rhodora [Vol. 74
evergreen ornamental has become rampant in a low woods
in Giant City State Park. COLLECTION DATA: Jackson
County: low woods, Giant City State Park, February 6,
1971, R. H. Mohlenbrock 21626.
Elaeagnus umbellatus Thunb. is previously unreported
from Illinois. Collections from this small tree were made
from an island in Lake-of-Egypt, Williamson County. At
this site a large number of plants are well established and
are spreading spontaneously. No evidence of a homesite
exists on the island. Elaeagnus umbellatus differs from E.
angustifolia, also a rare adventive in Illinois, by displaying
both brown and silver scales on the leaves and twigs, while
the latter has only silver scales present. COLLECTION DATA:
Williamson County: island in Lake-of-Egypt, August, 1970,
J. Swayne s.n.
Cynosciadium digitatum DC. represents a genus new to
the Illinois flora. Collections of this species were made from
the Pin Oak flats in the Greentree Reservoir, Jackson
County. This area, long set aside as a waterfowl preserve,
has seldom been botanized during the growing season since
it is mostly innundated and infested with mosquitos. Col-
lections made in 1969 and again in 1970 from widely sep-
arated areas indicate this species is well established at this
southern Illinois site. Associated species include Glyceria
striata, Carex squarrosa, Quercus palustris, and Ptilim-
nium costatum. The overall range of this species previously
reached from Mississippi and Texas north to Oklahoma,
and east to southeastern Missouri. COLLECTION DATA: Jack-
son County: Greentree Reservoir, Pin Oak flats, June 6,
1969, R. Anderson & D. Evans 1001.
Scutellaria ovata Hill var. rugosa (Wood) Fern. This tax-
on is very different in appearance from the other varieties
of S. ovata (var. ovata and var. versicolor) in Illinois. Its
small, rather sprawling habit is contrasted sharply with
the large, erect stature of the other varieties. Perhaps the
most striking difference exhibited by var. rugosa is found
in the leaves, which are strongly wrinkled and seldom
reach a length greater than 4 cm. In general, var. rugosa
1972] Illinois Studies — Mohlenbrock & Evans 149
occupies dry, rocky slopes, whereas var. ovata and var.
versicolor occur more frequently in more mesic woodlands.
Fernald (1950) indicates that var. rugosa is further dis-
tinguished by its smaller corolla (about 1 cm long). Ilinois
as well as Missouri material assignable to var. rugosa fre-
quently has the corolla up to 1.5 cm long, while some
corollas in var. ovata and var. versicolor may be less than
1.5 em long. COLLECTION DATA: Monroe County: south of
Fults, Illinois, August 14, 1952, W. Bailey & J. Swayne
2818.
Hypochoeris glabra L. This introduced species is not re-
corded in Gray’s Manual of Botany, 8th edition (Fernald,
1950). The station closest to Illinois seems to be eastern
South Carolina as reported by Radford, et al. (1964). In
Illinois, this species was collected from waste ground where
it grew with other pioneer-type weeds. The beakless outer
achenes of H. glabra distinguish it from H. radicata, also
a rare species in Illinois, in which all the nutlets are beaked.
COLLECTION DATA: Jackson County: rear of 1401 Tripoli,
Carbondale, July, 1969, P. Thomson s.n.
ADDITIONAL LOCALITIES FOR RARE
SOUTHERN ILLINOIS TAXA
Carex hystricina Muhl. This species is relatively common
in the northern two-thirds of the state, but is known else-
where in Illinois only from Pulaski and now Union County.
Union County: LaRue Swamp, June 28, 1969, D. Tindall &
S. Wunderle s.n.
Lychnis alba Mill. This Union County collection is from
the farthest south locality for this species in Illinois. Union
County: along Route 51, near Cobden, May 15, 1969, R. H.
Mohlenbrock & D. Evans s.n.
Ranunculus carolinianus DC. The achenes of this species
are larger (over 3.5 mm long) than those of R. septen-
trionalis and have a conspicuous broad, high keel near the
margin of the achene. Previously this species was known
from collections in Champaign and Hancock counties in the
central part of the state. The collection reported here is
150 Rhodora [Vol. 74
the first for southern Illinois. Union County: McCann
Springs, May, 1968, C. Ott s.n.
Lotus corniculatus L. This adventive is becoming increas-
ingly more common in Illinois. There are now three south-
ern Illinois counties for this species, as well as several
northern counties. Johnson County: roadside, near Gore-
ville, July, 1969, D. Evans & R. H. Mohlenbrock 1046.
Trifolium pratense L. f. leucochraceum Aschers. & Prantl.
Only a specimen from McLean County was previously
known from Illinois. Randolph County: roadside, north of
Prairie de Rocher, July 21, 1970, D. Evans 1108.
Hibiscus trionum L. The collection reported here is the
farthest south for this species in Illinois. Jackson County:
mud and sand flats of the Mississippi River, Grand Tower,
October 9, 1968, D. Evans 441.
Myriopsyllum exalbescens Fern. This species was pre-
viously known only from the northern half of Illinois.
Johnson County: Lake-of-Egypt, Beaver Neck, September
6, 1968, J. Swayne s.n.
Lysimachia terrestris (L.) BSP. This species was not
known from the southern half of the state until it was
collected at the edge of a swamp. Union County: LaRue
Swamp, June 25, 1968, M. Sadler s.n.
Verbena hastata X urticifolia Pepoon. This hybrid is
scattered in Illinois, but its only southern Illinois localities
previously had been from Hardin and Wabash Counties.
Jackson County: roadside, Southern Illinois University
campus, August 22, 1969, D. Evans & R. H. Mohlenbrock
1010.
Physalis pendula Rydb. The only previously known col-
lection of this species is from Union County. Jackson
County: mud flats of the Mississippi River, near Fountain
Bluff, October 26, 1968, D. Evans 543.
Centaurea solstitialis L. Jackson County can now be
added to Massac County for a second southern Illinois
locality for this adventive. Jackson County: along Illinois
Central Railroad, 1 mile S of Elkville, August 11, 1970,
D. Evans 1091.
1972] Illinois Studies — Mohlenbrock & Evans 151
LITERATURE CITED
ADAMS, P. 1957. A revision of the genus Ascyrum (Hypericaceae).
Rhodora 59: 73-95.
FERGUSON, A. M. 1901. Crotons of the United States. Annual Re-
port of the Missouri Botanical Garden 12: 33-73.
FERNALD, M. L. 1945. An incomplete flora of Illinois. Rhodora
47: 204-219.
1950. Gray's Manual of Botany, ed. 8. The Amer-
ican Book Co., New York. 1632 pp.
JONES, G. N. 1963. Flora of Illinois, ed. 3. The University of Notre
Dame Press, Notre Dame, Indiana. 401 pp.
, G. D. FULLER, G. S. WINTERRINGER, H. E. AHLES, & A.
FLYNN. 1955. Vascular Plants of Illinois. The University of
Illinois Press, Urbana, and the Illinois State Museum, Springfield.
593 pp.
RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1964. Guide to the
Vascular Flora of the Carolinas. The Book Exchange, Univer-
sity of North Carolina, Chapel Hill. 383 pp.
STEYERMARK, J. A. 1963. Flora of Missouri. The Iowa State Uni-
versity Press. Ames. 1725 pp.
DEPARTMENT OF BOTANY
SOUTHERN ILLINOIS UNIVERSITY
CARBONDALE, ILLINOIS 62901
CHRYSOPSIS MARIANA IN NEW ENGLAND: Until
very recently, the genus Chrysopsis was believed to be rep-
resented in New England by the single species C. falcata
(Pursh) Ell. It is a plant of the coastal plain ranging from
southeastern Massachusetts (notably Cape Cod) to New
Jersey, according to Gray’s Manual, 8th edition.
Another species, the wider ranging C. mariana (L.)
Ell. is reported from Block Island, Washington Co., Rhode
Island, by Mr. Robert Marks of Attleboro, Massachusetts,
and authenticated by specimens collected by him in 1969
and 1971. In a recent letter to me (November, 1971), he
states that three hundred or more plants occur in an area
of approximately one quarter mile on the Lewis farm in
the south-west corner of the island. There appears to be
a gradual decrease in abundance from near the ocean north-
ward and eastward. Vouchers are being processed for in-
clusion in the herbarium of the New England Botanical
Club.
It is not surprising that this species of Golden Aster
should turn up on Block Island. It has been known from
near-by Long Island, New York, for more than one hun-
dred years, e.g., Hempstead, D.C. Eaton, 1860, (NY) Judg-
ing from specimens in the herbarium of Brooklyn Garden
dated between 1886 and 1912, it seems to have been a fairly
common plant in all sections of the island including the
eastern end. I learn from correspondence that there is a
recently collected specimen at the New York Botanical Gar-
den from Montauk at the eastern tip (Andrews 3-84, 1962).
What is surprising is that such a conspicuous plant should
have escaped the attention of previous visitors to Block
Island, as early as ninety years ago, and especially M. L.
Fernald and companions who collected extensively there in
'W. W. Bailey in Notes on the Flora of Block Island (Bull. Torr.
Bot. Club 20: 231-239, 1893) published a list of 294 vascular plants
found by himself and his *youthful companion J. F. Collins" between
July 19 and August 31, 1892. If seen by them, C. mariana should
have been recognizable at the end of August.
152
1972] Chrysopsis — Eaton 153
August and again in September 1913.' It may have been
present at that time, mostly in a vegetative state, having
been suppressed by over-grazing by cattle, and only re-
cently is recovering or re-capturing its former habitat
among thickets of bayberry and other shrubs and stunted
trees now replacing the closely-cropped turf of former pas-
ture-land. Alternatively, Block Island may constitute a re-
cent extension of range. I think this is unlikely. Block
Island is downwind from nearby Montauk and presumably
has been exposed to invasion by wind-blown seeds of Chry-
sopsis for several millenia.
However, the colony should be watched. Mr. Marks knows
the island intimately and is confident that the Chrysopsis
is localized in its southwestern corner. If it gradually
spreads throughout, then recent colonization may be an
attractive supposition.
RICHARD J. EATON
22 DIVINITY AVENUE
CAMBRIDGE, MASSACHUSETTS 02138
ERIOGONUM HARPERI GOODMAN IN TENNES-
SEE: Eriogonum harperi was described by George J. Good-
man in 1947 (Bull. Torr. Bot. Club 74(4) : 329-331). The
description was based on material collected on a “marly
glade” in Colbert County, Alabama (Roland M. Harper
3944 in 1943). Two other locations for the species, based
on Harper's notes, were reported by Goodman. One was
“a calcareous prairie area", also in Colbert County, and
the other was “a small cedar glade” in the vicinity of Rus-
sellviile in adjacent Franklin County. These three locations
are within fifteen miles of each other.
Our specimens (Rogers and Watson 4073, August, 1970,
MISSA, TENN and University of Southern Mississippi) were
collected on a limestone bluff overlooking the Caney Fork
River where it intersects 1-40, about 7 to 8 miles east of the
Carthage exit, Wilson County, Tennessee. Perhaps a half
dozen plants were growing along the bluff near the high-
way.
This Tennessee locality is apparently the first new site
to be discovered since the description of the species and the
first site outside of Alabama. It extends the known range
of the species approximately 150 miles to the Northeast.
J. RAY WATSON, JR.
DEPARTMENT OF BOTANY
MISSISSIPPI STATE UNIVERSITY
STATE COLLEGE, MISSISSIPPI
KEN E. ROGERS
DEPARTMENT OF BIOLOGY
UNIVERSITY OF SOUTHERN MISSISSIPPI
HATTIESBURG, MISSISSIPPI 39401
LATHYRUS APHACA L. NEW TO TENNESSEE AND
THE SOUTHEAST: Lathyrus aphaca L. has been collected
in Madison County, Tennessee, as a wild plant by the senior
author. This species is a new record for Tennessee and,
apparently, for the Southeast as well. Its identification was
sought in all the manuals of the Eastern United States in
addition to Mahler’s recent monograph (1970) to no avail.
The bright yellow flowers always lead to Crotalaria in these
keys which it surely is not. Dr. Robert L. Wilbur of Duke
University kindly agreed to examine a specimen, and he
reports the identification to be L. aphaca. According to
Dr. Wilbur this is the first record of the species in the
Southeastern United States. Specimen cited: "Tennessee.
Madison County. 2.2 mi. S of Jackson on side of highway
45S. North bank. 15 June 1970. Richard L. Beardsley,
591. A specimen is deposited in the Memphis State Univer-
sity Herbarium, and others are to be distributed elsewhere.
Observations in 1971 indicate that this species is persistent
and may be spreading some from the original collection
site. The origin of the species in Tennessee can be only
speculative.
LITERATURE CITED
MAHLER, W. F. 1970. Manual of the legumes of Tennessee. J.
Tenn. Acad. Sci. 45(3): 65-96.
RICHARD L. BEARDSLEY
EDWARD T. BROWNE, JR.
DEPARTMENT OF BIOLOGY
MEMPHIS STATE UNIVERSITY
MEMPHIS, TENNESSEE 38111
RECORDS ON THE FLORA OF WISCONSIN: Leonurus
Marrubiastrum L. (Labiatae), a weedy adventive from Eu-
rope, is reported to range from Pennsylvania and Delaware
to Florida and Illinois. This is to announce an extension
of its range into southern Wisconsin: Dane County, waste
places in Kegonsa State Park, section 18, R11E, T6N,
Pleasant Springs Township, Hansen 164, 27 September
1970. One sheet is deposited in the herbarium, Wisconsin
State University-Oshkosh ; no duplicates were collected. Dr.
Hugh Iltis has kindly confirmed the identification; there
are no Wisconsin records for the plant at WIS, MIL, or UWM.
R. C. Koeppen (1957. Preliminary Reports on the Flora
of Wisconsin. No. 41. Labiatae — Mint Family. Trans. Wis.
Acad. Sci. Arts and Letters, 46: 115-140) does not report
the plant for Wisconsin.
Aethusa Cynapium L. (Umbelliferae), another weedy ad-
ventive from Europe, is reported by Fernald (Gray's Man-
ual, 8th Edition, 1950) to range from Nova Scotia to south-
ern Ontario and Minnesota, south to Delaware, Pennsyl-
vania, and Ohio. Gleason & Cronquist (Manual of Vascu-
lar Plants, 1963) assign it a more restricted range, from
Nova Scotia and Maine to Pennsylvania and Ohio. Dr.
Gerald Ownbey at MIN informs me that there are indeed
two Minnescta records for the plant in their collections, the
latest from 1902. However, the curators at WIS, MIL, and
UWM have no Wisconsin records for the plant, which can
now be tentatively added to the state's flora: Winnebago
County, along fence in back yard at 212 Oxford Avenue,
in the city of Oshkosh; growing as a weed in a fenceline
separating two vegetable gardens. The residents on either
side testify that it was certainly not planted there; Mis-
terek 037, 23 September 1970. Dr. Iltis has confirmed my
identification; the single specimen, deposited in the her-
barium here, has abundant fruits and perhaps may become
more than a mere casual element in our flora.
Linaria dalmatica (L.) Mill. (Scrophulariaceae) is re-
ported by Fernald (loc. cit.) to range from Cape Breton
156
1972] Wisconsin — Harriman 157
Island to Pennsylvania and Ohio. This is to report its
rather extensive occurrence in Wisconsin: Bayfield County,
abundant and well established on a sandy roadside on county
trunk J, ca. 1.5 miles N of its junction with county trunk
I, section 2, R4W, T50N, Bayfield Township; Harriman
7124, 17 July 1971. One sheet is deposited in the herbarium
here, with duplicates at UW and MIL. Iltis at UW informs
me (personal communication) that there are three rec-
ords for the plant from Bayfield County deposited there,
plus one each from Vilas, Sawyer, Sauk, Dane, and Green
counties. It is certainly to be included now in the Wiscon-
sin flora; it was not reported by P. J. Salamun (1951. Pre-
liminary Reports on the Flora of Wisconsin. No. 36. Scro-
phulariaceae. Trans. Wis. Acad. Sci, Arts and Letters. 40:
111-138).
NEIL A. HARRIMAN
BIOLOGY DEPARTMENT
UNIVERSITY OF WISCONSIN
OSHKOSH 54901
LOMENTARIA CLAVELLOSA (TURNER) GAILLON:
AN ADDITION TO THE MARINE ALGAL FLORA OF
NEW HAMPSHIRE. On 13 August 1971, several plants of
Lomentaria clavellosa were collected at Dover Point, Dover,
New Hampshire. Attached plants were found on rocks
about 3 feet below mean low water near the concrete sup-
port of the northbound section of the General Sullivan
Bridge. It was reported in North America for the first
time in 1963, from three stations in Boston Harbor, Massa-
chusetts (Wilce and Lee, 1964). Since 1963, it has also
been reported from Marthas Vineyard and Woods Hole in
Massachusetts. The occurrence of L. clavellosa at Dover
Point is not only a northward extension of its previously
known North American range but it is the first record of
this species from an estuarine environment in North Amer-
ica. However, although Dover Point is located several miles
from the open coast (it is the confluence of the Piscataqua
River and Little Bay), its algal flora is unique, with definite
coastal as well as estuarine species present (Mathieson,
Reynolds & Hehre, in press). Voucher specimens are de-
posited in the Algal Herbaria at Southampton College, the
University of New Hampshire and the University of Puerto
Rico at Mayaguez.
I would like to thank the following people for their
assistance in obtaining specimens of Lomentaria clavellosa:
Messrs. R. Shepp, R. Szita, D. Fallon, Miss Dorothy Meyers
and Miss Debbie Near.
LITERATURE CITED
MATHIESON, A. C., REYNOLDS, N. B., and HEHRE, E. J. Investiga-
tions of New England Marine Algae II. Species Composition,
Distribution and Zonation of Seaweeds in the Great Bay Estuary
System and adjacent Coastal Waters. Nova Hedwigia (in
press).
WILCE, R. T. and LEE, R. W. 1963. Lomentaria clavellosa in North
America. Botanica Marina 6(3/4).
EDWARD J. HEHRE
SOUTHAMPTON COLLEGE
SOUTHAMPTON, NEW YORK 11968
158
INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA
Manuscripts must be double-spaced or preferably triple-
spaced (not on corrasable bond), and a list of legends for
figures and maps provided on a separate page. Footnotes
should be used sparingly, as they are usually not necessary.
Do not indicate the style of type through the use of capitals
or underscoring, particularly in the citations of specimens,
except that the names of species and genera may be under-
lined to indicate italics in discussions. Specimen citations
should be selected critically especially for common species
of broad distribution. Systematic revisions and similar
papers should be prepared in the format of “The System-
atics and Ecology of Poison-Ivy and the Poison-Oaks,”
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu-
larly with reference to the indentation of keys and syno-
nyms. Papers of a floristic nature should follow, as far as
possible, the format of "Contribution to the Fungus Flora
of Northeastern North America. V.," H. E. Bigelow & M. E.
Barr, Rhodora 71: 177-208. 1969. For bibliographic cita-
tions, a recommended list of standard journal abbreviations
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot.
Club 85: 277-300. 1958.
Volume 74. No. 797. including pages 1-159, was issued April 7. 1972.
CONTENTS: — continued
Somatic Chromosome Numbers for Some Asteraceae.
Miloslav Kovanda eieaeoeaii a a oee a 102
Interactions between Apis mellifera (Hymenoptera:
Apidae) and Tribulus cistoides (Zygophyllaceae)
PO o eL mmerterénni blctlaodoneanadeteussiiuuliiaaseatene TI
Redefinition of Cedrela oaxacensis C. DC. & Rose.
C. Earle Smith, Jr. AEN A EE E E 124
An Abnormal Specimen of Laminaria digitata (L).
Lamouroux. Arthur C. Mathieson, Emery F. Swan and
Richard A. Fraliel ennea eene eaae o inten etta tatto sette aeta P eods 127
Range Extensions of Vascular Aquatic Plants in New Eng-
land. C. Barre Hellquist........... esee 131
A Second Occurrence for Triphora trianthophora (SW.)
Rydb. in Maine. Lesley M. Eastman ........... ee 141
Illinois Field and Herbarium Studies.
Robert H. Mohlenbrock and Dam K. Evang ............ ee 142
Chrysopsis mariana in New England. Richard J. Eaton .... 152
Eriogonum harperi Goodman in Tennessee:
J. Ray Watson, Jr. and Ken E. Rogers ...... eee 154
Lathyrus aphaca L. New to Tennessee and the Southeast.
Richard L. Beardsley and Edward T. Browne, Jr. ........ 155
Records on the Flora of Wisconsin. Neal A. Harriman .... 156
Lomentaria clavellosa (Turner) Gaillon: An Addition to
the Marine Algal Flora of New Hampshire.
Edward J. Hehe ...cccorccssocsscccsssssscsssssccessssssssesoncgasovanconcstennseasere 158
Hovdova
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by
ALBION REED HODGDON, Editor-in-Chief
ALBERT FREDERICK HILL
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON »Associate Editors
RADCLIFFE BARNES PIKE
ELIZABETH ANNE SHAW
STEPHEN ALAN SPONGBERG
Vol. 74 June, 1972 No. 798
CONTENTS:
Revision of the Genus Melampodium (Compositae: Heli-
antheae). Tod F. Stwessy essent
Spores, Chromosomes and Relations of the Fern Pellaea
atropurpurea. Alice F. Twryom viccccccccsssssssssssssscsssescsssesscscescece
The Genus Coriaria (Coriariaceae) in the Western Hemi-
Sphere. Lawrence E. Skog essees.
List of the Cladoniae of Southeastern Massachusetts, with
Special Reference to the Collection of Charles Albert
Robbins. Lawrence B. Mish ciccccccccccssscsssssssssssssssssssssscessaene
On the Marine Algae of Kent Island, Bay of Fundy.
Kenneth L. Koetzner and R. D. Wood aese
( Continued on Inside Back Cover)
242
Che New England Botanical Club, Bue.
Botanical Museum, Oxford St., Cambridge, Mass. 02138
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Rbodora
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Vol. 74 June, 1972 No. 798
REVISION OF THE GENUS MELAMPODIUM
(COMPOSITAE: HELIANTHEAE)'
Top F. STUESSY
(Continued from page 70)
II. Melampodium section Zarabellia (Cass.) DC.
Melampodium L. sect. Zarabellia (Cass.) DC. Prodr. 5:
519. 1836. Zarabellia Cass. Dict. Sc. Nat. 59:240. 1829.
Type species: Melampodium longifolium Cerv. ex Cav.
Tap-rooted annuals; leaves rhombic to deltoid (rarely
narrowly ovate), sessile or shortly petiolate, with margins
entire to obscurely serrate; outer involucre usually cupu-
late or less often spreading, with bracts 3-5, at margins
herbaceous; ovaries of the disc florets ovoid, less than 0.7
mm long, rudimentary (Figs. 3 & 4) ; fruits at apex nearly
smooth or moderately sculptured, or sometimes with an
abaxial protuberance (in M. longifolium, Fig. 7) ; chromo-
some base number, x — 9. Species 21-25.
21. Melampodium longifolium Cerv. ex Cav. Anal. Cien.
Nat. 6:333. 1803. non Brouss. ex Willd. 1809. TYPE:
"NUEVA HISPANIA": grown from seed in the Royal Bot.
Gard. Madrid, flowering Jun-Sep, V. Cervantes s.n. ( Holo-
type, MA?; isotype, G!; photograph of G-DC isotype, IDC
800. 927: III. 6!; photograph of G isotype, F! us!).
Melampodium longifolium Brouss. ex Willd. Enum. Plant.
Hort. Berol. 934. 1809. nom. illegit. non Cerv. ex Cav. 1803.
161
162 Rhodora [Vol. 74
TYPE: MEXICO: locality, date, and collector unknown, culti-
vated in Bot. Gard. Berlin and harvested in 1808 (Holotype,
B; photograph of holotype, TEX!).
Zarabellia rhomboidea Cass. Dict. Sc. Nat. 59:241. 1829.
TYPE: cultivated in Bot. Gard. Paris, source of seeds un-
known (Holotype, P?; photograph of possible G-DC isotype,
Ipc 800. 927: III. 7!).
Dysodium divaricatum Hort. ex DC. Prodr. 5:520. 1836.
non Rich. in Pers. 1807. nom. illegit., pro syn.
Melampodium rhomboideum (Cass.) DC. Prodr. 5:520.
1836.
Melampodium parvulum Brandg. Zoe 5:262. 1908. TYPE:
MEXICO: Puebla, Cerro Santa Lucia, Jul 1907, C. A. Purpus
2813 (not 2813’) (Holotype, UC!; isotypes, F! GH! NY! US!;
photograph of US isotype, TEX!).
Annual herbs, 8-45 cm long. Stems erect, 1.5-4 mm diam,
moderately hispid near base to copiously hispid above with
hairs 0.8 mm long. Peduncles 0-21 mm long. Leaves sessile,
rhombic (rarely elliptic, younger leaves often lanceolate-
ovate), 2.5-6 cm long, 0.5-2.7 cm wide, at apex acute, at base
subauriculate, with both surfaces moderately pubescent
with hairs 0.3 mm long; margin obscurely serrate. Heads
3-4 mm tall, 5-7 mm diam. Outer involucre spreading, 7-10
mm diam; bracts 5 (rarely 4), separate, lanceolate, 3-5 mm
long, 1.2-2 mm wide, at apex acute-acuminate, with abaxial
surface glabrous to moderately pubescent (very pubescent
at base) with hairs 0.7 mm long; margin herbaceous.
Fruits 2.2-3 mm long, with lateral surfaces with 2 large
irregular cavities. Ray florets 5-8; ligules yellow, ovate-
elliptic, 1.3-1.5 mm long, 0.8-0.9 mm wide. Disc florets 10-
15; corollas yellow, 0.8 mm diam, with throat 0.7 mm and
tube 0.6 mm long. Paleae oblong, 1.4 mm long, 0.7 mm
wide; apex colorless, with margin erose; midrib distinct,
glabrous. Chromosome number, » — 9.
Scattered throughout pine-oak forests in the Mexican
states of Nuevo Léon, San Luis Postosi, México, Distrito
1972] Melampodium — Stuessy 163
Federal, Puebla, and Oaxaca (Fig. 12), 1830-2740 m. Flow-
ering dates, Aug-Sep.
REPRESENTATIVE SPECIMENS. Mexico. DISTRITO FEDERAL: Tlalpan,
Pedregal, Valley of Mexico, 20 Aug 1896, Harshberger 176 (GH);
Tlalpan, Pyramid of Cuicuilco, 15 Aug 1935, MacDaniels 53 (F);
Valley of Mexico near Tlalpan, 20 Aug 1896, Pringle 6455 (F, GH,
NY, UC, US); Pena Pobre near Tlalpan, 8 Aug 1944, Sharp 44208
(NY). MEXICO: near El Oro, 14 Sep 1901, Pringle 9325 (F, GH, US).
NUEVO LEON: Hacienda Pablillo, Galeana, 8 Aug 1936, Taylor 115
(DS, F, NY, RSA, TEX). OAXACA: N limits of Las Sedas, 15 Aug 1966,
Stuessy 659 (TEX). PUEBLA: vicinity of Puebla, 9 Sep 1906, Arséne
76 (vs), 27 Oct 1907, Arsène 1199 (US), 10 Sep 1910, Nicolas 5447
(GH, US), s.n. (NY); Teocalli de Cholula, near Puebla, 25 Sep 1909,
Nicolas s.n. (NY); Rancho Posadas, near Puebla, 25 Sep 1909, Nico-
las sn. (NY); vicinity of San Luis Tultitlanapa, Aug 1908, Purpus
2813’ (F, GH, NY, UC, US). SAN LUIS POTOSÍ: vicinity of San Luis
Potosi, 1878, Parry & Palmer 444 (F, GH, NY, US); 22 mi E of San
Luis Potosí, 5 Aug 1961, Powell & Edmondson 551 (F, TEX).
22. Melampodium mimulifolium Robins. Proc. Amer. Acad.
Arts & Sci. 36:462. 1901. TYPE: MEXICO: Oaxaca, Totonto-
pec, 1700-2150 m, 15 Jul 1894, E. W. Nelson 740 (Holotype,
US!; photograph of holotype, TEX !).
Annual herbs, over 30 cm tall. Stems erect, 2-3 mm diam,
subglabrous below to pilose-villous above with hairs 1 mm
long. Peduncles 3-4.2 cm long. Leaves sessile, lanceolate-
elliptic, 4.5-6.8 cm long, 1.5-2.2 cm wide, at apex acuminate,
at base subauriculate to auriculate, with both surfaces stri-
gose with hairs 0.4-1 mm long; margin entire to serrulate.
Heads 6-7 mm tall, 8-10 mm diam. Outer involucre spread-
ing to shallowly cupulate, 8-9 mm diam; bracts 5, slightly
connate at base, ovate-lanceolate, 5-6 mm long, 2-2.5 mm
wide, at apex acute, with abaxial surface moderately pilose
with hairs 0.5-1 mm long; margin herbaceous. Fruits
2-2.1 mm long, with lateral surfaces smooth-striate with
enlarged tuberculate margins. Ray florets ca 8; ligules
yellow-orange, ovate, 3-4 mm long, 1.5-3 mm wide. Disc
florets ca 50; corollas yellow, 1 mm diam, with throat 0.8
mm and tube 1.4 mm long. Paleae oblanceolate-obovate, 2.2
mm long, 0.8 mm wide; apex yellow, with margin erose-
164 Rhodora [Vol. 74
laciniate; midrib weak, glabrous. Chromosome number un-
known.
Known only from the type collection from pine-oak for-
ests at Totontopec, Oaxaca, Mexico (Fig. 12), 1700-2150 m.
Flowering date, Jul.
M. mimulifolium is very similar to M. gracile, and in fact,
additional collecting may disclose transitional specimens
that might suggest merger of the two species. At present,
however, M. mimulifolium seems treated best as a distinct
species that can be distinguished from M. gracile by the
former's shorter peduncles, narrower leaves, longer and
narrower outer phyllaries, and absence of stipitate-glandu-
lar hairs anywhere on the plant.
23. Melampodium gracile Less. Linnaea 6:407. 1831. TYPE:
MEXICO: Veracruz, Papantla, Jan 1829, C. J. W. Schiede &
F. Deppe 1254 (Lectotype, HAL!; isotypes, HAL! P! w! Mo!
photograph of lectotype, 0S! TEX!; photograph of HAL iso-
type, OS! TEX!; photograph of P isotype, oS! TEX!).
Melampodium oblongifolium DC. Prodr. 5:519. 1836.
TYPE: MEXICO: Veracruz, “rancho de los huevos” [from
label] near Tantoyuca, Jan 1832, J. L. Berlandier 2153 [=
733] (Lectotype chosen, G-DC; isotypes, F! G [2]! GH! K [2]!
MO! P!; photograph of lectotype, IDC 800. 927: III. 2!; trac-
ing of lectotype, GH!; photograph of G-DC isotype, IDC 800.
927: IIL 3!; photograph of K isotypes, vus [2]!).
Melampodium gracile Less. var. oblongifolium (DC.) A.
Gray, Proc. Amer. Acad. Arts & Sci. 5:182. 1861.
Melampodium microcarpum S. F. Blake, Contrib. U. S.
Nat. Herb. 22:606. 1924. TYPE: MEXICO: Tamaulipas, vi-
cinity of Gómez Farias, ca 350 m, 13-21 Apr 1907, E.
Palmer 319 (Holotype, US!; isotype, GH! MO!; photograph
of holotype, TEX! UC!).
Annual herbs, 17-50 cm tall. Stems erect, 1.2-4 mm diam,
hispid-pilose and weakly stipitate-glandular with hairs
0.1-2 mm long. Peduncles 5.5-11.5 cm long. Leaves sessile,
1972] Melampodium — Stuessy 165
eri
WA ]
n »
k bs.
/ Va
Fig. 21. Map of Mexico and adjacent Guatemala and British
Honduras showing distribution of Melampodium gracile (triangles),
and M. microcephalum (dots).
usually deltoid (younger leaves often lanceolate), 5-10 em
long, 2-4 cm wide, at apex acute to acuminate, at base nar-
rowed and dilated near stem to subauriculate (rarely ob-
tuse), with both surfaces strigillose with hairs 0.2 mm long;
margin obscurely crenate to irregularly cleft. Heads 6-7
mm tall, 8-15 mm diam. Outer involucre spreading to shal-
lowly cupulate, 7-10 mm diam; bracts 3 (sometimes 4 or
5), slightly connate at base, imbricate 1/3 their length,
ovate, 3.5-5 mm long, 2.5-3.5 mm wide, at apex acuminate,
with abaxial surface strigose with hairs 0.3-1 mm long near
apex, near base stipitate-glandular with hairs 0.1-0.2 mm
long; margin herbaceous. Fruits 2.3-2.5 mm long, with lat-
eral surfaces reticulate and striate to tuberculate. Ray
florets 5-8; ligules yellow-orange, ovate, 2-5 mm long, 3-4
mm wide. Disc florets 30-45; corollas yellow-orange, 1.5
166 Rhodora [Vol. 74
mm diam, with throat and tube each 0.9 mm long. Paleae
oblanceolate-obovate, 2.2 mm long, 0.8 mm wide; apex
yellow-orange, with margin entire to dentate-laciniate lat-
erally; midrib absent. Chromosome number, n — 9.
Pine-oak, tropical deciduous, tropical evergreen, and rain
forests on the Gulf side of the Sierra Madre Oriental and
the Yucatan Peninsula, and on the Pacific sides of the
Sierra Madres Occidental and Sur, with extensions into
British Honduras and northwestern Guatemala (Fig. 21),
30-1830 m. Flowering dates, Jul-Oct.
Melampodium gracile is very similar morphologically
and geographically to M. microcephalum, but the two differ
in several features such as habit, and the shapes of leaf
bases, outer involucral bracts, and ligules. Single popula-
tions of both species have been crossed reciprocally in the
greenhouse, and although robust F, hybrids are obtained,
they are almost completely sterile. Detailed analyses of
hybridization among all three species in the genus that
have stipitate-glandular peduncles (M. gracile, M. micro-
cephalum and M. paniculatum) currently are in progress
and the results will be reported in a separate paper.
REPRESENTATIVE SPECIMENS. British Honduras. EL CAYO: San Agus-
tin, Jul-Aug 1936, Lundell 6759 (ARIZ, GH, NY, TEX [2], Us). Guate-
mala. HUEHUETENANGO: between San Sebastián H. & San Rafael
Pétzal, 14 Aug 1942, Steyermark 50539 (F); between Nentón &
Miramar, 29 Aug 1942, Steyermark 51452 (F, UC). PETÉN: vicinity
of La Libertad, Aug-Nov 1933, Aguilar 25 (F [photograph of US
specimen], NY, US). Mexico. CAMPECHE: Champotón, 7-15 Jul 1932,
Steere 1805 (us); 1 mi W of Champotón, 18 Jul 1966, Stuessy 531
(TEX); 16 mi N of Champotón, 18 Jul 1966, Stuessy 533 (TEX); 12
mi S of Campeche, 25 Jul 1966, Stuessy 546 (TEX). CHIAPAS: ca
28 mi SE of Comitán, 25 Jun 1960, King 3042 (DS, NY, TEX, UC, US);
ca 17 mi S of Tuxtla Gutiérrez, 27 Jun 1960, King 3096 (DS, NY, TEX,
uc, Us); 10 mi S of Bochil, 25 Jul 1966, Stuessy 562 (TEX); 32 mi
SE of Comitán, 26 Jul 1966, Stuessy 574 (TEX). COLIMA: 7 mi S of
Colima, 25 Aug 1966, Stuessy 722 (TEX). GUERRERO: Acuitlapan,
24 Sep 1937, Abbott 439 (GH); Rio Balsas, Aug 1910, Orcutt 4365
(F); near Iguala, 14 Sep 1900, Pringle 9177 (F, GH, SMU, UC, US),
10 & 12 Aug 1905, J. N. & J. S. Rose, & Painter 9308 (GH, US).
HIDALGO: Jacala, 20 Oct 1937, Kenoyer 427 (F). MÉXICO: Temas-
1972] Melampodium — Stuessy 167
caltepee, Anonas, 24 Jul 1934, Hinton 6338 (US). MICHOACÁN: ca
4 mi E of Jiquilpán, 6 Aug 1960, King 3636 (DS, NY, TEX, UC, US) ;
ca 5 mi E of Zamora, 6 Aug 1960, King 3646 (DS, NY, TEX, UC, US);
30 mi NW of Zamora, 2 Sep 1965, Stuessy 393 (TEX); 25 mi S of
Ario de Rosales, 20 Aug 1966, Stuessy 694 (TEX) ; 2 mi S of jct rte 15
& rd to Cotija, 24 Aug 1966, Stuessy 713 (TEX). MORELOS: ca 11 mi
8 of Cuernavaca, 14 Mar 1961, King 4165 (TEX) ; 6 mi W of Yautepec,
14 Aug 1950, Pipes 81 (SMU); valley near Jojutla, 17 Jun 1901,
Pringle 8466 (F, GH, NY, POM, UC, US); near Yautepec, 27 Oct 1902,
Pringle 9937 (F, GH, Ny, US); 7 mi NW of Cuautla, 25 Aug 1965,
Slessy 356 (TEX). SAN LUIS POTOSÍ: El Pujal, Río Tampaón valley,
21 Jul 1939, Chase 7533 (ARIZ, F, GH, NY); El Salto, 15 Jun 1951,
Holman & Dittman 791 (SMU) ; El Salto, 14 Jun 1951, Kay & Higgins
202 (SMU); El Salto, 20 Feb 1961, King 3887 (F, NY, TEX, US); 48 mi
N of Tamazunchale, 23 Aug 1957, Waterfall & Wallis 14288 (smu).
TAMAULIPAS: 7 mi N of Padilla, 9 Jun 1951, Paynet & Hulan 1213
(SMU, TEX); 10 km NW of El Progreso, 22 Aug 1941, Stanford,
Retherford & Northeraft 1078 (ARIZ, DS, GH, NY) ; 15 mi SW of Ciudad
Mante, 9 Aug 1965, Stuessy 274 (TEx [2]); ca 3-4 mi E of Nuevo
Morelos, 5 Jul 1966, Stuessy 463 (TEX); 3 mi E of Gómez Farias,
8 Jun 1967, Stuessy 826 (TEX). VERACRUZ: Huasteca, Wartenberg,
near Tantcyuca, 1858, Ervendberg 92 (GH); 13 mi W of Orizaba,
18 Nov 1959, Graham & Johnston 4777 (TEX); 7 mi SW [SE] of
Morelos, 12 Aug 1961, Powell & Edmondson 646 (F, TEX): Jalapa,
1894, Smith 1605 (au, UC); 18 mi S of ject rtes 110 & 105, 6 Jul
1966, Stuessy 466 (TEX). YUCATAN: Progreso, Gawmer 2349 (F);
Izamal, 1895, Gaumer & sons 789 (Ds, F, GH [2], Ny [2], pom, vc [3],
US); at km 13 on Merida-Progreso rd, May-Aug 1938; C. & A.
Lundell 7972 (ps, F, TEX, UC, US); 13 km N of Merida, 20 Jul 1966,
Stuessy 537 (TEX); 86 mi W of Valladolid, 21 Jul 1966, Stuessy 544
(TEX [2]).
24. Melampodium microcephalum Less. Linnaea 9:268.
1854. TYPE: MEXICO: in cultivation near Hacienda de la
Laguna, Jul 1829, C. J. W. Schiede 217 (Holotype, HAL!;
isotype, P!; photograph of holotype, os! TEX!; photographs
of P isotype, F! os! TEX! US!).
Melampodium lanceolatum Sessé & Moc. Fl. Mex. ed. 2.
196. 1894. nom. illegit. non DC. 1836. TYPE: “calidoribus
Novae Hispaniae regionibus,” Jul 1795-1804, M. Sessé et al.
“3978” (Lectotype chosen, MA! [drawing of stem with two
pairs of leaves on reverse side of label] ; isotypes, MA [3]!).
168 Rhodora [Vol. 74
Annual herbs, 15-75 cm tall. Stems erect to decumbent
and rooting at nodes, 1-2 mm diam, villous and weakly
stipitate-glandular with hairs 0.1-1 mm long. Peduncles
2.5.6.5 cm long. Leaves sessile or with petioles 1-4 mm
long, ovate-lanceolate, 3-6.5 cm long, 0.9-3.5 cm wide, at
apex acute to obtuse, at base attenuate to obtuse, with both
surfaces strigose with hairs 0.2-1 mm long; margin entire
to obscurely crenate. Heads 3-4 mm tall, 5-10 mm diam.
Outer involucre spreading to shallowly cupulate, 5-8 (-10)
mm diam; bracts 3 (sometimes 4 or 5), slightly connate
at base, separate, narrowly ovate, 3-4.5 mm long, 1.5-3 mm
wide, at apex acuminate, with abaxial surface pilose-villous
and stipitate-glandular with hairs 0.1-1 mm long; margin
herbaceous. Fruits 1.7-2 mm long, with lateral surfaces
rugose-reticulate with enlarged tuberculate margins. Ray
florets 5-8; ligules yellow-orange, elliptic, 2.5-4 mm long,
2.9 mm wide. Disc florets 35-50; corollas yellow-orange, 1.6
mm diam, with throat 1 mm and tube 0.7 mm long. Paleae
oblanceolate-obovate, 2 mm long, 0.8 mm wide; apex yellow,
with margin serrulate-laciniate; midrib absent. Chromo-
some number, n = 9.
Mostly pine-oak, tropical deciduous and tropical ever-
green forests in many states of Mexico in the Mesa Central
and the Sierra Madres Occidental and Sur, and in adjacent
Guatemala (Fig. 21), 50-1740 m. Flowering dates, Jun-
Nov.
M. microcephalum is often confused with M. gracile and
M. paniculatum. Although all these taxa possess three outer
involueral bracts, stipitate-glandular stems and peduncles,
and chromosome numbers on a base of x — 9, the size of
the heads and the shapes of the leaf bases can be used to
distinguish easily one from another (see key).
In most taxa of Melampodium the number of outer in-
volucral bracts is very constant, but in both M. microcepha-
lum and M. gracile, as seen from field and greenhouse
studies, plants occasionally are found with four or five
bracts instead of the usual three.
1972] Melampodium — Stuessy 169
REPRESENTATIVE SPECIMENS. (Guatemala. HUEHUETENANGO: ca 6
mi S of Huehuetenango, 18 Jul 1960, King 3425 (DS, NY, TEX, UC, US).
Mexico. CHIAPAS: 10 mi SE of Tonalá, 10 Aug 1966, Stuessy 627
(TEX). GUANAJUATO: near Acambaro, 18 Oct 1904, Pringle 13069
(F, GH, SMU, UC). GUERRERO: Taxco, 28 July 1937, Abbott 304 (GH);
20 mi NE of Acapulco, 20 Aug 1947, Barkley, Webster & Paxson
17M732 (TEX); Mina, Manchón, 7 Aug 1936, Hinton 9205 (ARIZ);
Mina, Río Frio, 25 Sep 1937, Hinton et al. 10719 (ARIZ, GH, POM,
TEX, US). JALISCO: Cerro Viejo, above Zapotitan de Hidalgo, ca
45 mi S of Guadalajara, 27 Jun 1956, Gregory & Eiten 205 (SMU).
MÉXICO: Temascaltepec, Volcán, 10 Sep 1932, Hinton 1661 (F, GH,
NY [2], US); Temascaltepec, Bejucos, 6 Oct 1932, Hinton 2008 (Ds,
F, NY [2], US) ; Temascaltepec, Ixtapán, 22 Jun 1934, Hinton et al.
6207 (F, GH, US). MICHOACÁN: 7 mi W of La Piedad, 6 Oct 1962,
Cronquist 9613 (NY, TEX, US) ; ca 3 mi S of Ciudad Hidalgo, 4 Aug
1960, King 3607 (Ds, NY, TEX, UC, US); ca 22 mi S of Uruapan,
Cerro de Carboneras, 16-22 Oct 1961, King & Soderstrom 4804 (NY,
SMU, TEX, UC, US); Lake Cuitzeo, 26 Oct 1892, Pringle 4322 (F, GH
[2, one a tracing], Ny, vc [2], US); 6 mi NW of Tuxpán, 1 Sep 1965,
Stuessy 383 (TEX). MORELOS: El Teycal, 9 km SW of Tepoztlan,
30 Aug 1955, Clausen s.n. (NY); Cuantla, Aug 1904, Kuntze 23527
(NY [3]) ; Yautepec, 30 May 1939, Nagel 8030 (GH); Barranca, near
Cuernavaca, 2 Nov 1896, Pringle 7321 (GH, US); Moyotepec, 8 Nov
1964, Ripley & Barneby 13708 (NY). OAXACA: Cerro de San Felipe,
26 Sep 1897, Conzatti & González 550 (GH, US); 10-12 km E of
Niltepec, 18 Jul 1959, King 1833 (NY, SMU, TEX, UC, US) ; vicinity of
Choapám, 28-29 Jul 1894. Nelson 870 (US); 1 mi E of Monte Alban,
12 Aug 1966, Stuessy 638 (TEX); 29 mi S of Zimatlán, 12 Aug 1966,
Stuessy 640 (TEX). PUEBLA: Mt Orizaba, Botteri 809 (F, GH); near
Puebla, Jul 1888, Hoffmann 832 (GH); Orizaba, 27 Mar 1887, Mills-
paugh s.n. (F); Mt Orizaba, 24 Aug 1891, Seaton 461 (F, GH, NY,
US). VERACRUZ: Tantoyuca, Dec 1830, Berlandier s.n. (GH); valley
of Córdoba, 20 Jan 1866, Bourgeau 1628 (GH, US) ; valley of Córdoba,
23 Apr 1865-66, Bourgeau 1629 (US); Pacho, near Jalapa, Jan 1894,
Smith 1655 (F, NY, RSA).
25. Melampodium paniculatum Gardn. Hook. Lond. Jour.
Bot. 7:287. 1848. TYPE: BRAZIL: Goias, near Arraias, Apr
[Feb?] 1840, G. Gardner 3844 (Holotype, K!; isotypes, F
[2]! c [2]! Ny [2]! P! w!; photograph of holotype, us!;
photograph of G isotype, F! US!).
Melampodium laxum Sch. Bip. Ber. Bot. Zeit. 3:155. 1845.
nom. nud. Based on G. Gardner 3844.
170 Rhodora [Vol. 74
Melampodium brachyglossum J. D. Smith, Bot. Gaz. 13:
74. 1888. TYPE: GUATEMALA: Alta Verapaz, Coban, 4300 ft,
May 1886, H. von Tuerckheim 114 (Lectotype chosen, US!;
isotype, US!; photograph of holotype, TEX!). K, NY & P
specimens that bear exactly the same printed label as the
lectotype are clearly Jaegeria hirta (Lag.) Less. (see
Torres, 1968, for agreement). It is probably a labeling
error.
Annual herbs, 13-70 em tall. Stems erect, 0.7-5 mm diam,
hispid and stipitate-glandular with hairs 0.1-1 mm long.
Peduncles 1.1-5 cm long. Leaves sessile or with petioles up
to 5 mm long, ovate-rhombic (younger leaves approaching
lanceolate), 3-9 cm long, 1-5 cm wide, at apex acute to
acuminate, at base attenuate, often dilated almost to sub-
auriculate, with both surfaces strigose with hairs 0.3-1 mm
long; margin entire to irregularly serrate-crenate. Heads
2.2-4.5 mm tall, 3-5.5 mm diam. Outer involucre cupulate,
5-7 mm diam; bracts 3, separate, ovate, 1.8-4 mm long,
1.5-2.8 mm wide, at apex acute-acuminate, with abaxial
surface strigose and stipitate-glandular with hairs 0.1-0.8
mm long; margin herbaceous. Fruits 2-2.8 mm long, with
lateral surfaces with longitudinal ribs and striations. Ray
florets 3-5; ligules yellow, ovate, 1-2 mm long, 1-2 mm wide.
Disc florets 10-15; corollas yellow, 1.1 mm diam, with throat
and tube each 0.8 mm long. Paleae elliptic, 1.3 mm long,
0.7 mm wide; apex yellow-orange, with margin laciniate;
midrib weak, glabrous. Chromosome numbers, n = 8 and
27.
Low tropical mountains of Chiapas, Mexico, Central
America (excluding British Honduras, Nicaragua, and Pan-
ama), and isolated points of Colombia and Brazil in South
America (Figs. 22 & 23), 370-2010 m. Flowering dates,
Jan-Dec.
Adding to the chromosomal diversity already existing
in Melampodium are the new counts in M. paniculatum of
n = 27 reported from two different populations in Brazil
1972] Melampodium — Stuessy 171
Fig. 22. Map of Central America showing distribution of Me-
lampodium paniculatum. See Fig. 23 for South American localities.
172 Rhodora [Vol. 74
Fig. 23. Map of South America showing distribution of Melampo-
dium divaricatum (dot), and M. paniculatum (circles).
by Coleman (1970)'*. I have examined the cited voucher
specimens, and due to an absence of significant morpholog-
ical differences between the » — 18 and n — 27 cytotypes,
it seems best at this time to consider these chromosomally
distinct populations as simply polyploid races of a single
species. It is interesting to note, however, that the tetra-
ploid counts of n = 18 prevail in Central America while
the chromosomally derived hexaploid populations of m
— 27 are found only in Brazil where M. paniculatum prob-
ably has been recently introduced by man. M. paniculatum
thus joins M. cinereum, M. dicoelocarpum and M. leucan-
thum as the fourth species in the genus known to have
intraspecific euploidy.
"These new counts actually were listed as » — 26-27 and n — 26.
In recent correspondence regarding these counts, however, Dr. Cole-
man states that “On restudying the slide, I have decided that the
correct number of the material I counted is n — 27."
1972] Melampodium — Stuessy 173
REPRESENTATIVE SPECIMENS. Brazil. FEDERAL DISTRICT: Córrego
Landim, ca 20 km N of Brasilia, 15 Dee 1965, Irwin, Souza & Santos
11303 (TEX); Corallino, 1817-21, Pohl 1368 [1276] (NY). GOIAS: near
Arraias, Apr 1840, Gardner 287 (NY). MATO GROSSO: Sta Terezinha,
12 Feb 1948, Macedo 1408 (US). MINAS GERAIS: 1838, Clausen s.n.
(F, NY [2], Us); Belo Horizonte, Pampulha, Feb 1945, Williams
5643 (vc, US). RÍO DE JANEIRO: Río de Janeiro, Feb 1882, Glaziou
12794 (K). SAO PAULO: São Joao de Boa Vista, 6 Sep 1893, Lofgren
& Edwall 16695 (Us). Colombia. CUNDINAMARCA: Bogotá, Triana
1331 (NY, US). Costa Rica. CARTAGO: 24 Aug 1924, Torres 50 (Us).
GUANACASTE: coffee plantations of San Francisco de Guadalupe, Jul
1892, Pittier 6963 (GH); Hatillo, 4 Jan 1935, Solís 52 (F). HEREDIA:
Río Virilla on hwy to Heredia, 1 Dec 1937-1 Jan 1938, Allen 585 (F);
Río Virrilla near La Uruca, Aug 1888, Pittier 429 (F, GH); Río
Virilla near San Juan, Jun 1896, Tonduz 10144 (vs [2]). SAN JOSE:
San José, 4 Sep 1938, Orozco 260 (F); vicinity of San José, 6 Nov
1938, Orozco 418 (F); vicinity of Las Pavas, 29 Feb 1924, Standley
36058 (US); vicinity of San José, Feb 1924, Standley 38971 (vs);
Desamparados, 27 Jun 1923, Stevens 136 (vs). EI Salvador. AHUA-
CHAPÁN: vicinity of Ahuachapán, 9-27 Jan 1922, Standley 20260
(GH), 16-25 Jan 1947, Standley & Padilla 2755 (F). LA LIBERTAD:
Finca Paraíso, 4 mi S of Sta Tecla, 14 Jan 1946, Carlson 106
(F, UC, US) ; Sta Tecla, 20 Jul 1941, García 51 (UC). SAN SALVADOR:
Volcán de San Salvador, Apr 1922, Calderón 459 (GH, NY, US) ; San
Salvador, 1922, Calderón 779 (GH, NY, US); vicinity of San Salvador,
30 Mar-24 Apr 1922, Standley 22687 (GH, US); Volcan de San Salva-
dor, 7 Apr 1922, Standley 22864 (GH, NY, US); San Salvador, Nov
1906, Velasco 8946 (US). SANTA ANA: Cerro Miramundo, above
Hacienda Los Planos, NE of Metapán, 25 Feb 1946, Carlson 873 (F);
Finca of Los Naranjos, E of Cerro de los Naranjos, 27 Apr 1942,
Tucker 1313 (F, NY, UC, US). SAN VICENTE: vicinity of San Vicente,
7-14 Feb 1947, Standley & Padilla 3630 (F). Guatemala: ALTA VERA-
PAZ: ca 4 mi NE cf San Pedro Carcha, 12 Jul 1960, King 3329 (ps,
NY, TEX, UC, US); 28 mi E of San Miguel Uspantán, 28 Jul 1966,
Stuessy 589 (TEX); 1 mi NE of San Pedro Carchá, 29 Jul 1966,
Stuessy 594 (TEX): Coban, Jul 1885, Tuerckheim 761 (us), Mar 19083,
8417 (F. GH, NY, US). CHIMALTENANGO: near Río Pixcavó between
Chimaltenango & San Martin Jilotepeque, 3 Feb 1939, Standley 64303
(F, NY); Finca La Alameda near Chimaltenango, 11-22 Dec 1940,
Standley 79869 (F). EL QUICHE: Nebaj, 19 Nov 1934, Skutch 1709
(F, GH); 4 mi E of Cunén, 28 Jul 1966, Stuessy 585 (TEX); 11 mi E
of San Miguel Uspantan, 28 Jul 1966, Stuessy 587 (TEX). ESCUINTLA:
Finca Monterrey, Volcán de Fuego, 5 Feb 1939, Standley 64595 (F,
NY); Los Diamantes, 9 Mar 1896, Volkem 2564 (GH, NY). GUATE-
MALA: near Finca La Aurora, 1938-39, Aguilar 322 (F). HUEHUE-
174 Rhodora [Vol. 74
TENANGO: ca 3 mi S of Huehuetenango, 18 Jul 1960, King 3417
(DS, NY, TEX, UC, US); vicinity of Cuilco near Rio Cuileo, 17 Aug 1942,
Steyermark 50773 (F, GH); 2 mi S of Huehuetenango, 26 Jul 1966,
Stuessy 578 (TEX); 1 mi W of Aguacatán, 27 Jul 1966, Stuessy 582
(TEX); Chaculá, 20 Aug 1896, Volkem 2774 (GH, NY, US). JALAPA:
vicinity of Jalapa, 7-18 Nov 1940, Standley 76429 (F); between
Jalapa & San Pedro Pinula, 12 Nov 1940, Standley 77073 (F); mts
near Chahuite, NW of Jalapa, 16 Nov 1940, Standley 77440 (F);
between Jalapa & Montana Miramundo, 7 Dec 1939, Steyermark 32847
(F). QUEZALTENANGO: vicinity of San Francisco de Miramar, Costa
Cuca, Apr 1905, Pittier 57 (F, US), Pittier 60 (F, US); between
Colomba & Las Mercedes, 20 Feb 1941, Standley 87933 (F); San
Francisco Miramar, 15 Mar 1921, Tonduz & Rojas 203 (Us).
RETALHULEU: Finca Helvetia, 6 May 1937, Muenscher 12467 (F).
SACATEPÉQUEZ: Antigua, 12 Sep 1959, O. & I. Degener 26504 (US);
Antigua, 13 Feb 1905, Kellerman 4501 (F, US); Antigua, Apr 1890,
Smith 2334 (us); near Antigua, Nov 1938-Feb 1939, Standley 60326
(F, NY); Barranca above Duefias, 21 Jan 1939, Standley 63265 (F,
NY). SAN MARCOS: Río Cabüs near Malacatán, 15 Mar 1939, Standley
68866 (F). SANTA ROSA: near Cerro Redondo, 20 Dec 1938, Standley
60413 (F); La Joya de Limón, E of Cuilapa, 25 Nov 1940, Standley
78268 (F). soLotA: Panajachel, near Lake Atitlán, 11 Sep 1959,
O. & I. Degener 26559 (US) ; behind Lago de Atitlán near Panajachel,
5 Jul 1960, King 3242 (ps, NY, TEX, UC, US). SUCHITEPÉQUEZ: Finca
Moca, 27 Oct 1934, Skutch 1550 (GH). Honduras. CHOLUTECA: vi-
cinity of San Marcos de Colón, 12-22 Jan 1949, Standley 15756 (F).
COMAYAGUA: vicinity of Siguatepeque, 14-27 Feb 1928, Standley
56290 (vs), 56522 (F, US), 25 Mar-5 Apr 1947, Standley & Chacón
6625 (F), 6660 (r); Rittenhouse's hacienda, 30 Jun 1936, Yuncker,
Dawson & Youse 5519 (F, GH). EL PARAÍSO: Güinope, Dec 1943,
Valerio 1662 (F). INTIBUCA: vicinity of La Esperanza & Intibucá,
31 Jan-12 Feb 1950, Standley 25149 (F). MORAZÁN: Río Yeguare,
22 Jul 1948, S. F. Glassman 2003 (r, Nv), 26 Aug 1951, Williams
18257 (F); near Río Yeguare, below El Zamorano, 10 Aug 1947,
Standley 12105 (F); Cerro de Uyuca between Las Flores & Tatumbla,
17 Aug 1949, Standley 22718 (F); La Leona, Tegucigalpa, 6 Sep
1951, Standley 28678 (F); Zamorano, 7 Jan 1946, Valerio 3735 (F,
GH). OLANCHO: vicinity of Juticalpa, 5-16 Mar 1949, Standley 17710
(F). Mexico. CHIAPAS: Tenejapa, Pokolum, 10 Jul 1964, Breedlove
6085 (ps, F); Chamula, Toh Tik on rd to Chenalho, 12 Dec 1964,
Breedlove 7861 (ps, F); Los Lagos, 3 mi NW of Rancho San José,
18 Jan 1952, Carlson 2254 (F).
1972] Melampodium — Stuessy 175
III. Melampodium section Serratura Stuessy, sect. nov.
Herbae annuae radicibus palaribus; folia rhombica vel
deltoidea (raro anguste ovata), petiolata, marginibus ser-
ratis (raro integris); involucrum extimum cupulatum vel
interdum patens, bracteis 5, marginibus herbaceis; ovaria
flosculi disci ovoidea, minus quam 0.7 mm longa, rudimen-
tales (Figs. 3 & 4) ; fructus apice fere laeves vel moderate
sculpti et sine cucullis, vel complanata abaxiali arista (M.
tepicense) ; chromosomatum numerus basicus, x = 12.
Species 26-30. Typus: Melampodium divaricatum (Rich.
in Pers.) DC.
26. Melampodium divaricatum (Rich. in Pers.) DC. Prodr.
5:520. 1836.
Dysodium divaricatum Rich. in Pers. Syn. 2:489. 1807.
non Hort. ex DC. 1836. TYPE: COLOMBIA: “Gairam, prope
St. Martham," 1785-89, L. C. M. Richard s.n. (Lectotype
chosen, P!; isotype, P!; photograph of holotype, os! TEX!;
photograph of P isotype, os!).
Wedelia ovatifolia Willd. Enum. Suppl. 61. 1814. TYPE:
source of seed unknown, apparently grown in Bot. Gard.
Berlin, “2421” (Holotype, B?; isotype, w!; photograph of w
isotype, OS! TEX!; photograph of probable G-DC isotype, IDC
800. 928: I. 4!).
Alcina ovatifolia (Willd.) Jacq. f. Eclog. P1. 1:115. t. 78.
1815.
Wedelia minor Hort. ex Hornem. Enum. Hort. Hafn.
2:855. 1815. TYPE: introduced into the Bot. Gard. Copen-
hagen in 1812 from Berlin Bot. Gard., source of seed
unknown (Lectotype, c!; isotypes, C [2]!; photograph of
lectotype, oS! TEX!).
Alcina ovalifolia Lag. Gen. et Sp. Nov. 32. 1816. TYPE:
"NOVA HISPANIA" [MEXICO] : seeds cultivated and grown in
Bot. Gard. Madrid, 1805, M. Sessé s.n. (Holotype, MA!;
isotype, G!; photograph of holotype, os! TEX!; photograph
of G isotype, OS! TEX!).
176 Rhodora [Vol. 74
Melampodium paludosum H.B.K. Nov. Gen. Sp. 4:287.
1820. TYPE: PANAMA: “in humidis prope ostia fluminis sinu,
juxta litora Dariensis," Mar 1801, F. H. A. von Humboldt
& A. J. Bonpland 1421 (Holotype, P; isotype, P!; photo-
graph of P isotype, OS! TEX!).
Melampodium ovatifolium Rchb. Icon. Bot. Exot. 30. t. 42.
1824. nom. superfl. Based on type of Dysodium divaricatum
Rich. in Pers.
Melampodium berterianum Spreng. Syst. Veg. 3:619.
1826. TYPE: “IND. occ.” [COLOMBIA]: near Sta Martha, C.
G. Bertero s.n. (Holotype, P!; probable isotype, GH!; photo-
graph of holotype, os! TEX [2]! US!; photograph of G-DC
isotype, IDC 800. 928: I. 3!).
Alcina minor Cass. Dict. Sc. Nat. 59:243. 1829. TYPE:
MEXICO: cultivated in Bot. Gard. Paris (Holotype, P?).
Dysodium radiatum Hort. ex Desf. Cat. Hort. Paris. ed. 3.
182. 1829. pro syn.
Melampodium pumilum Benth. Pl. Hartw. 64. 1840. TYPE:
MEXICO: exact locality unknown, 1830, G. J. Graham 5
(Holotype, K!; isotype, GH!; photograph of holotype, os!
TEX! US!).
Melampodium tenellum Hook. & Arn. var. flaccidum
Benth. Bot. Voy. Sulphur 115. 1845. TYPE: MEXICO: Nay-
arit, Tepic, 1844, A. Sinclair s.n. (Lectotype chosen, K!;
isotype, K !; photograph of lectotype, US [3] !).
Melampodium divaricatum (Rich. in Pers.) DC. var.
macranthum Schlecht. Linnaea 24:198. 1851. TYPE: GUATE-
MALA: locality and date unknown, seeds collected and
grown in Bot. Gard. Turin, J. von Warscewicz s.n. (Holo-
type, HAL; isotype, W !; photograph of w isotype, 0S! TEX!).
Melampodium flaccidum (Benth.) Benth. in Oerst. Kjoeb.
Vidensk. Meddel. 5-7:86. 1852.
Melampodium copiosum Klatt, Engl. Bot. Jahrb. 8:41.
1887. TYPE: GUATEMALA: Alta Verapaz, near Cobán, 1300
1972] Melampodium — Stuessy 177
m, Feb [May?] 1882, F. C. Lehmann 1434 (Holotype, GH!;
isotypes, F! G [2]! K! US!; photograph of K isotype, US!;
photograph and fragment of K isotype, US!).
Melampodium panamense Klatt, Engl. Bot. Jahrb. 8:42.
1887. TYPE: PANAMA: “ad margines silvarum," 20 Aug
1880, F. C. Lehmann 69 (Holotype, GH!; isotypes, G! K!
US!; photograph of K isotype, US!).
Spilanthes guatemalensis Vatk. ex J. D. Smith, Enum. PI.
Guat. 1:23. 1889. nom. nud. Based on H. von Tuerckheim
124 (F! GH! NY! US [2]!).
Melampodium rhombifolium Sessé & Moc. Fl. Mexic. ed. 2.
195 [first on page]. 1894. nom. illegit. non Sessé & Moc.
1890. nec Sessé & Moc. [second on page]. 1894. TYPE:
MEXICO: Michoacan, Temascaltepec, Jul 1790-1791, M. Sessé
et al. “3983” (Holotype, MA!; isotype, F!; photograph of
holotype, OS! TEX!).
Eleutheranthera divaricata (Rich. in Pers.) Millsp. Field
Col. Mus. Pub. Bot. 1:53. 1895.
Annual herbs, 15-100 cm tall. Stems erect, with lateral
stems sometimes decumbent and rooting at the nodes, 1.3-6
mm diam, glabrous to pubescent with hairs up to 0.7 mm
long. Peduncles 1.5-13.5 cm long. Leaves with petioles 2-20
mm long, ovate to rhombic (younger leaves approaching
lanceolate), 1.5-15 cm long, 0.5-9.5 cm wide, at apex acumi-
nate to obtuse, at base attenuate (rarely obtuse to cordate),
with both surfaces infrequently strigillose with hairs 0.2
mm long; margin entire to coarsely dentate-crenate. Heads
5-10 mm tall, 10-21 mm diam. Outer involucre cupulate, 6-9
mm diam; bracts 5, connate 1/3-1/4 their length, imbricate,
ovate-orbiculate, 3.5-6 mm long, 3-5 mm wide, at apex
obtuse, with abaxial surface glabrous to tomentose near
peduncle; margin herbaceous, near base tomentose. Fruits
2.8-4 mm long, with lateral surfaces with diagonal stria-
tions and enlarged margins. Ray florets 8-13; ligules yel-
low-orange, oblong-elliptic, 3.5-7 mm long, 1.6-3 mm wide.
Disc florets 40-70; yellow-orange, 2 mm diam, with throat
Fig. 24. Map of Mexico, Central America, northern South Amer-
ica, and the Caribbean region showing distribution of Melampodium
divaricatum. See Fig. 23 for the one Brazilian locality. Collections
from Burma and the Virgin Islands not shown.
1 mm and tube 0.5 mm long. Paleae obovate, 2.5 mm long,
0.9 mm wide; apex yellow-orange, with margin dentate-
erose; midrib weak, glabrous. Chromosome number, n
= 12.
Very widely spread in many subtropical habitats through-
out Mexico, Central America, and northwestern Colombia,
also in eastern Brazil, and introduced into Cuba, Burma,
Puerto Rico and the Virgin Islands (Figs. 23 & 24), 15-
2990 m. Flowering dates, Jan-Dec.
This is the most widespread and weedy species of the
genus. Although field and garden studies have shown leaf
shape to be extremely variable, which partially accounts
for the many synonyms accorded this taxon, the morphology
of the outer involucral bracts is very constant and allows
the species to be recognized without difficulty. One plant
from Nicaragua, Stwessy 615, has fruits with an anomalous
hooded condition; other plants from the same population
are hoodless.
REPRESENTATIVE SPECIMENS. Brazil. BAHÍA: 1842, Glocker 19 (US);
Marau, 22 Jan 1965, Pereira 9616 & Pabst 8505 [34972] (F). British
1972] Melampodium — Stuessy 179
Honduras. EL CAYO: El Cayo, near Camp 6, 11 Mar 1938, Gentle 2325
(ARIZ, F, GH, NY). Burma. MANDALA: Maymyo, Nov 1950, White
100 (US). Colombia. ANTIOQUÍA: near Dabeiba, 20 Dec 1947, Barkley
& Gutiérrez 1777 (F). BOLIVAR: vicinity of Turbaco, 6-22 Nov 1926,
Killip & Smith 14169 (F, Ny, US). CORDOBA: W of Monteria, near
Río Sinu, 6 Jan 1949, Cervantes, Molina & Barkley 19Bo061 (vus).
MAGDALENA: Sta Marta, Bonda, 9 Nov 1898-1899, Smith 514 in part
(NY [3], Uc); Sta Marta, Jul 1898-1901, Smith 514 in part (F, RSA,
SMU, TEX, UC, US). Costa Rica. ALAJUELA: between San Josecito &
La Garita, 18 Aug 1963, Jiménez 1073 (F, US). GUANACASTE: Nicoya,
Dec 1899, Tonduz 13615 (NY, US). PUNTARENAS: 21 km SE of
Puntarenas, near bridge over Río Barranca, 20 Aug 1938, Stork &
Morrison 8920 (GH, UC). Cuba. HAVANA: Vedado, shore of Arigu-
anabo River, San Antonio de los Baños, 3 Sep 1939, Léon, Victorin
& Alain 17344 (NY). El Salvador. AHUACHAPÁN: vicinity of Ahua-
chapán, 9-27 Jan 1922, Standley 19722 (GH, US). LA LIBERTAD: Finca
Paraíso, 4 mi S of Sta Tecla, 14 Jan 1946, Carlson 79 (F, UC, US).
MORAZÁN: Río San Miguel opposite Barrios Mine, ca 5 km W of
Montecristo, 19 Dec 1941, T'ucker 585 (F, NY, UC, US). SAN MIGUEL:
39 mi W of La Unión, 7 Aug 1966, Stwessy 623 (TEX). SAN SALVADOR:
18 mi E of San Salvador, 2 Aug 1966, Stuessy 611 (TEX). SANTA ANA:
4 mi SE of Sta Ana, 2 Aug 1966, Stuessy 609 (TEX). SAN VICENTE:
vicinity of San Vicente, 2-11 Mar 1922, Standley 21648 (GH, NY, US).
SONSONATE: vicinity of Sonsonate, 18-27 Mar 1922, Standley 21789
(GH, NY, US). Guatemala. ALTA VERAPAZ: ca 2 mi E of Coban, 12
Jul 1960, King 3322 (ps, NY, TEX, UC, US); 28 mi E of San Miguel
Uspantán, 28 Jul 1966, Stuessy 588 (TEX); 1 mi NE cf San Pedro
Carchá, 29 Jul 1966, Stuessy 592 (TEX). BAJA VERAPAZ: ca 1 mi S
of Rabinal, 14 Jul 1960, King 3359 (DS, NY, TEX, UC, US). CHIMAL-
TENANGO: San Martín J., 1928, Morales 1221 (F). CHIQUIMULA:
Montana Nonojá, 3-5 mi E of Camotán, 11 Nov 1939. Steyermark
31670 (F). EL PROGRESO: between Finca Piamonte & Finca San
Miguel, 9 Feb 1942, Steyermark 43745 (F, UC). EL QUICHE: 11 mi
E of San Miguel Usnantaén, 28 Jul 1966, Stuessu 586 (TEX). ESCU-
INTLA: 9 mi NE of Escuintla, 8 Aug 1966, Stuessy 624 (TEX).
GUATEMALA: ca 4 mi E of Guatemala, 9 Jul 1960, King 3249 (ps,
NY, TEX, UC, US). HUEHUETENANGO: Chacula, 29 Jul 1896, Volkem
3013 (GH, NY, US). IZABAL: Cristina, 23 May 1919, Blake 7632
(US). JALAPA: vicinity of Jalapa, 7-18 Nov 1940, Standley 77149
(F). JUTIAPA: 25 mi E of Cuilapa, 2 Aug 1966, Stuessy 605 (TEX);
8 mi NE of Jutiapa, 2 Aug 1966, Stuessy 608 (TEX). QUEZALTENANGO:
Colomba, 31 Dec 1934, Skutch 2041 (F, GH). RETALHULEU: 1 mi W
of jet rtes 2 & 9S, N of Retalhuleu, 10 Aug 1966, Stuessy 625 (TEX).
SANTA ROSA: Volcán Jumaytepeque, Dec 1892, Heyde et Lux 4213
(F, GH, NY, US [2]). SUCHITEPEQUEZ: Mazatenango, 28 Feb 1905,
180 Rhodora [Vol. 74
Kellerman 4958 (SMU, US). ZACAPA: Gualán, 18 Jan 1905, Deam
247 (F, GH, NY, US). Honduras. EL PARAÍSO: Llano de Lizapa, 24
Aug 1948, Molina 1082 (F). FRANCISCO MORAZÁN: near San Francisco
along Yeguare River, 16 Nov 1947, Williams & Molina 13398 (F, GH).
VALLE: Amapala, 11 Sep 1945, Valerio 3341 (F [2]). Mexico. BAJA
CALIFORNIA: San José del Cabo, 12 Sep 1890, Brandegee 301 (UC).
CAMPECHE: 1 mi W of Champotón, 18 Jul 1966, Stuessy 532 (TEX) ;
14 mi N of Champotón, 18 Jul 1966, Stuessy 534 (TEX); 14 mi E
of Campeche, 18 Jul 1966, Stuessy 535 (TEX); 12 mi S of Campeche,
22 Jul 1966, Stuessy 545 (TEX). CHIAPAS: ca 15 mi S of Suchiate,
20 Jul 1960, King 3434 (DS, NY, RSA, TEX, UC); 19 mi S of Pichucalco,
23 Jul 1966, Stuessy 550 (TEX). COLIMA: 4 mi E of Colima, 25 Aug
1966, Stuessy 723 (TEX); 7 mi NW of rte 110 on rd to Alzada, 25 Aug
1966, Stuessy 725 (TEX); Alzada, 25 Aug 1966, Stuessy 728 (TEX).
DISTRITO FEDERAL: Sta Rosa, 4 Oct 1952, Matuda 26621 (US). GUANA-
JUATO: valley near Irapuato, 20 Sep 1889, Pringle 2819 (F, GH).
GUERRERO: vicinity of Acapulco, Oct 1894-Mar 1895, Palmer 186
(F, GH, NY, US) ; 4 mi S of Taxco, 17 Aug 1961, Powell & Edmondson
756 (F, TEX); S limits of Petaquillas, 27 Aug 1965, Stuessy 365
(TEX). HIDALGO: Jacala, 25 Jun 1939, Chase 7122 (F). JALISCO: Río
Cuvianes, 13 Jun 1892, Jones 273 (PoM [2], US). MEXICO: Tema-
scaltepec, 2 Sep 1932, Hinton 1520 (NY). MICHOACÁN: ca 45 mi W
of Morelia, 5 Aug 1960, King 3634 (DS, NY, RSA, TEX, UC); 16 mi
NW of Zitacuaro, 19 Aug 1966, Stuessy 677 (TEX); S city limits
of Uruapán, 21 Aug 1966, Stuessy 702 (TEX). MORELOS: ca 11 mi
S of Cuernavaca, 14 Mar 1961, King 4164 (F, NY, TEX, UC, US) ; 35.5
mi NW of Izácar de Matamoros, 25 Aug 1965, Stuessy 348 (TEX) ;
6 mi NW of Cuautla, 25 Aug 1965, Stuessy 351 (TEX); 10 mi S of
Cuernavaca, 26 Aug 1965, Stuessy 359 (TEX). NAYARIT: ca 16 mi
NW of Ixtlán del Río, 10 Aug 1960, King 3678 (Ds, NY, TEX, UC, US) ;
1 mi N of Tepie, 27 Aug 1966, Stuessy 743 (TEX); 10 mi NW of
jet rte 16 & rd to Tuxpán, 27 Aug 1966, Stuessy 746 (TEX). NUEVO
LEÓN: Villa de Santiago, Las Adjuntas, 21 Jun 1940, Leavenworth
157a (F). OAXACA: ca 1 mi W of the Oaxaca-Chiapas border on
rte 190, 9 Jun 1960, King 2743 (DS, NY, TEX, UC, US); 0.7 mi N of
Huajuapán de León, 24 Aug 1965, Stuessy 342 (TEX); 40 mi S of
Zimatlán, 13 Avg 1966, Stuessy 645 (TEX). PUEBLA: just SE of the
Morelos-Puebla border on rte 115, 19 Jun 1960, King 2917 (DS, NY,
TEX, UC, US); Asunción de Chila, 25 Aug 1965, Stuessy 345 (TEX).
QUERÉTARO: 2 mi W of Landa, 13 Dec 1960, Crutchfield & Johnston
6120b (TEX). QUINTANA ROO: Coba, Jun-Jul 1938, C. & A. Lundell
7879 (US). SAN LUIS POTOSÍ: ca 10 mi S of Tamazunchale, 23 Mar
1961, King 4239 (F, NY, TEX, UC, US); Cardenas, 3 Nov 1891, Pringle
3923 (DS, F, GH, NY, vc [2], Us); 2 mi off rte 80 on rd to Quadal-
cazar, 10 Aug 1965, Stuessy 282 (TEX). SINALOA: ca 15 mi N of
1972] Melampodium — Stuessy 181
Rosario, 13 Aug 1960, King 3711 (DS, NY, RSA, TEX, uc); 4.5 mi N
of Escuinapa, 4 Sep 1965, Stuessy 403 (TEX). TABASCO: 57 mi SE
of Coatzacoalcos, 17 Jul 1966, Stuessy 527 (TEX); 84 mi E of Coat-
zacoalcos, 17 Jul 1966, Stuessy 528 (TEX); 5 mi W of Villa Hermosa,
23 Jul 1966, Stuessy 547 (0S, TEX); 37 mi S of Villa Hermosa,
23 Jul 1966, Stuessy 548 (TEX). TAMAULIPAS: ca 4-5 mi W of Ciudad
Mante, 18 Feb 1961, King 3839 (F, NY, TEX, US); 4 mi S of Juamave,
6 Jul 1949, Stanford, Taylor & Lauber 2318 (DS, GH, NY, RSA, SMU,
TEX, UC, US [2]) ; 1 mi E of Gómez Farias, 8 Jul 1967, Stuessy 829
(TEX). VERACRUZ: ca 25 mi SE of Poza Rica, 10 Mar 1961, King 4138
(F, NY, TEX, US); 23.2 mi SE of Alvarado, 21 Aug 1965, Stuessy 318
(TEX); 12 mi S of Tantoyuca, 6 Jul 1966, Stuessy 473 (TEX);
49 mi SE of Catemaco, 16 Jul 1966, Stuessy 526 (TEX). YUCATÁN:
Izamal, 1895, Gawmer 563 (F, GH [2], NY [2], us); Progreso, 20 Jul
1966, Stuessy 543 (TEX). Nicaragua, CHINANDEGA: vicinity of
Chichigalpa, 12-18 Jul 1947, Standley 11226 (F). GRANADA: Volcan
Mombacho, 20 Feb 1903, Baker 2482 (GH, POM, UC, us); NE limits
of Granada, 6 Aug 1966, Stuessy 620 (TEX). MANAGUA: 1 mi SE
of Managua, 5 Aug 1966, Stwessy 616 (TEX); 5 mi E of Managua,
6 Aug 1966, Stuessy 621 (TEX). MASAYA: near Masaya, SW slopes
of Santiago Voleano, 5 Jul 1923, Maxon 7694 (US). MATAGALPA:
Sebaco, 5 Aug 1966, Stuessy 614 & 615 (TEX). PUNTARENAS: 21 km
SE cf Puntarenas, 20 Aug 1938, Stork & Morrison 8920 (GH, UC).
Panama. CANAL ZONE: vicinity of Frijoles, 9 Jul 1962, King 5226
(UC, US). DARIÉN: trail between Pinogana & Yavisa, 17 Mar 1937,
Allen 241 (cH [3], NY, US). PANAMA: near Arraiján, 21 Jul 1938,
Woodson, Allen & Seibert 1870 (GH, NY, US). PEARL ARCHIPELAGO:
San José Is, Naval Station, 22 Jul 1945, Erlanson 500 (DS, GH,
TEX, US). Puerto Rico: Mayaguez, 14-15 Feb 1914, Britton & Cowell
1561 (NY). Virgin Islands. ST. CROIX: near Mt Pleasant, 19 Aug
1896, Ricksecker "S" (F, NY).
27. Melampodium costaricense Stuessy, Brittonia 22:118.
f.7. 1970. TYPE: COSTA RICA: Alajuela, Naranjo, Cerro del
Espíritu Santo, 1200 m, 11 Jul 1941, A. Smith 2922 (Holo-
type, F!).
Annual herbs, 15-35 em tall. Stems erect, with lateral
stems often decumbent, 0.8-4 mm diam, glabrous to tomen-
tose with hairs 0.8 mm long. Peduncles 0.5-7 cm long.
Leaves with petioles 4-25 mm long, ovate-rhombic (younger
leaves approaching lanceolate), 2.5-7 cm long, 1.5-5.5 cm
wide, at apex acuminate to obtuse, at base obtuse-attenuate,
with upper surface infrequently strigose with hairs 0.6 mm
182 Rhodora [Vol. 74
long, with lower surface glabrous; margin obscurely to
coarsely serrate. Heads 4-5 mm tall, 6-7 (10) mm diam.
Outer involucre cupulate, 5-10 mm diam; bracts 5, connate
1/3-1/4 their length, imbricate, ovate-orbiculate, 2.2-5 mm
long, 2-4 mm wide, at apex obtuse, with abaxial surface
glabrous; margin herbaceous, ciliate with hairs 0.3 mm
long. Fruits 2.9-3.1 mm long, with lateral surfaces with
diagonal ridges and enlarged margins. Ray florets 5-8;
ligules yellow-orange, elliptic, 1-1.5 mm long, 0.5-1 mm
wide. Disc florets 15-25; corollas yellow, 1.1 mm diam, with
throat 0.8 mm and tube 0.5 mm long. Paleae oblong-elliptic,
2 mm long, 0.7 mm wide; apex yellow, with margin erose;
midrib weak, glabrous. Chromosome number, n = 25 + 1.
Tropical habitats in British Honduras, Nicaragua, Costa
Rica and Panama of Central America, and Colombia, South
America (Fig. 25), 0-1980 m. Flowering dates, Jan-Dec.
REPRESENTATIVE SPECIMENS, British Honduras. EL CAYO: El Cayo,
Jun-Aug 1936, Lundell 6107 (F, GH, NY, US). Colombia. CUNDINA-
MARCA: Caqueza to Rio Sananie, 24 Aug 1917, Pennell 1335 (Ny).
Costa Rica. ALAJUELA: banks of the Río Rosales y Carret, 8 Dec
1933, Brenes 17471 (NY); San Pedro de San Ramón, 29 Jul 1935,
Brenes 20572 (F). CARTAGO: above Tres Ríos, 19 Mar 1965, Godfrey
67156 (FsU); Turrialba, Instituto Interamericano de Agrícolas,
20 Aug 1962, King 5348 (TEX, UC, US); Finca Las Concavas, 7-8
Dec 1925, Standley 41531 (US); rr below Turrialba, 9 Jun 1928,
Stork 2486 (F, US). GUANACASTE: Hatillo, 4 Jan 1935, Solís 56 (F).
LÍMON: marshes of Suerre & Dos Bocas, 3 Oct 1951, Shank & Molina
4236 (F, GH); Hamburg Finca, on the Río Reventazon below Cairo,
19 Feb 1926, Standley & Valerio 48814 (US). SAN JosÉ: Alajuelita,
8 Sep 1946, Echeverría 628 (F); San Isidro, 26 Feb 1965, Godfrey
6664 (FSU); San Pedro de Montes de Oca, 2 Aug 1938, Orozco 294
(F); El General [San Isidro del], Jan 1939, Skutch 3968 (GH, K,
NY, US); between San Pedro de Montes de Oca & Curridabat, 2 Feb
1924, Standley 32757 (us); San José, Feb 1924, Standley 39021
(US). Nicaragua. GRANADA: near Granada, 1845-48, Oersted 9010
(photograph of K specimen, US). Panama, BOCAS DEL TORO:
Changuinola Valley, 15 Mar 1924, Dunlap 557 (F, US). CHIRIQUT:
25 mi N of Concepción, 29 Jul 1962, King 5290 (TEX, UC, US).
28. Melampodium dicoelocarpum Robins. Proc. Amer. Acad.
Arts & Sci. 44:619. 1909. TYPE: MEXICO: Guerrero, El Cala-
1972] Melampodium — Stuessy 183
Fig. 25. Map of Central America and neighboring countries show-
ing distribution of Melampodium costaricense.
bazal, 300 m, 20 Oct 1898, E. Langlassé 482 (Holotype, GH!;
isotypes, G! K! P! US!; photograph of K isotype, US!; photo-
graph of US isotype, TEX!).
Annual herbs, 10-50 cm tall. Stems erect, 0.6-1.6 mm
diam, glabrous to moderately pilose with hairs 0.3-1.3 mm
long. Peduncles deflected, 0.4-7.5 cm long. Leaves with
petioles 2-10 mm long, ovate to somewhat deltoid (younger
leaves often ovate-lanceolate), 2-6 cm long, 0.7-3 cm wide,
at apex acuminate-attenuate, at base obtuse-attenuate, with
both surfaces moderately pilose with hairs 1 mm long;
margin obscurely to coarsely serrate. Heads 2-4 mm tall,
2.5-8 mm diam. Outer involucre spreading, 2-7 mm diam;
bracts 5 (rarely 3 or 4), slightly connate at base, separate,
ovate-lanceolate, 1.5-4.5 mm long, 1-2.5 mm wide, at apex
acuminate, with abaxial surface moderately strigose with
184 Rhodora [Vol. 74
hairs 0.6-1 mm long; margin herbaceous. Fruits 1.7-3 mm
long, with lateral surfaces with 2 deep oval depressions.
Ray florets 3-5; ligules yellow-orange, oblong-elliptic, 2.2-
2.5 mm long, 1.2-1.8 mm wide. Disc florets 25-35; corollas
yellow-orange, 0.8 mm diam, with throat 0.7 mm and tube
0.5 mm long. Paleae oblong-elliptic, 2 mm long, 0.7 mm
wide; apex yellow, with margin erose to laciniate; midrib
weak, glabrous. Chromosome numbers, n — 12 and 23.
Tropical deciduous forests in the Mexican states of Jalis-
co, Michoacán, Guerrero and México (Fig. 13), 200-1890 m.
Flowering dates, Jun-Oct.
The cytological vouchers for the two chromosome levels
are morphologically distinguishable, the aneuploid at the
tetraploid level being larger in most characters. However,
without intensive field and garden studies, it seems prema-
ture to propose formal recognition of these forms.
Among all the taxa of Melampodium, M. dicoelocarpum
can be distinguished easily by the two deep oval cavities on
the lateral sides of the fruit.
REPRESENTATIVE SPECIMENS. Mexico, GUERRERO: Mina, Zihuagio,
17 Oct 1936, Hinton 9714 (G, GH, MICH, NY, P, UC, US); Galeana,
Atoyac-San Juan, 12 Oct 1939, Hinton et al. 14614 (ARIZ, F, GH, NY,
US). JALISCO: Tecalitlán, Barranca de San Juan de Dios, ca 15 km
E of Pihuamo, 24 Oct 1963, Feddema 2204 (MICH). MÉXICO: Temas-
caltepee, Anonas, 24 Jun 1934, Hinton et al. 6334 (GH, MICH, NY).
MICHOACÁN: ca 5 mi N of Cotija, 5-9 Oct 1961, King & Soderstrom
4646 (MICH, NY, SMU, TEX, UC, US); 25 mi S of Ario de Rosales,
20 Aug 1966, Stuessy 693 (TEX); 15 mi S of jet rte 15 on rd to
Cotija, Stwessy 715 (TEX).
29. Melampodium tepicense Robins. Proc. Amer. Acad.
Arts & Sci. 44:620. 1909. TYPE: MEXICO: Nayarit, Tepic,
"
5 Jan-6 Feb 1892, E. Palmer 1814 (Holotype, GH!; iso-
types, F! NY! Us!).
Melampodium cornutum S. F. Blake, Jour. Wash. Acad.
Sei. 16:421. 1926. TYPE: MEXICO: Colima, Alzada,4 Nov
1910, C. R. Orcutt 6601 (Holotype, Us!; photograph of holo-
type, TEX!).
1972] Melampodium — Stuessy 185
Annual herbs, 7-35 cm tall. Stems decumbent to erect,
0.5-1.3 mm diam, mostly glabrous to puberulent with hairs
0.1 mm long. Peduncles 0.8-8 mm long. Leaves with nar-
rowly marginate petioles 1-18 mm long, ovate to rhombic,
0.8-5.7 cm long, 0.7-3.5 cm wide, at apex acute, at base at-
tenuate to obtuse, with both surfaces sparsely strigose with
hairs 0.3-1 mm long; margin crenate to serrate, near base
entire. Heads 2-3 mm tall, 3-5 mm diam. Outer involucre
cupulate, 3.5-7 mm diam; bracts 5, slightly connate at base,
separate, elliptic, 2-4 mm long, 1.2-2.2 mm wide, at apex
obtuse, with abaxial surface glabrous; margin herbaceous.
Fruits 1.6-2 mm long, with lateral surfaces tuberculate to
aculeate-reticulate; apex with a flattened abaxial append-
age’, with tapering cirrhous awn up to 4 mm long. Ray
florets 3-7; ligules pale-yellow, ovate-oblong, 0.8-1.5 mm
long, 0.4-0.8 mm wide. Disc florets 3-7(12) ; corollas yel-
low-green, 0.8 mm diam, with throat 0.4 mm and tube 0.7
mm long. Paleae oblong-oblanceolate, 1 mm long, 0.5 mm
wide; apex pale-yellow, with margin irregularly dentate;
midrib weak, glabrous. Chromosome number unknown.
Pine-oak and tropical deciduous forests in Nayarit, Jal-
isco, Colima, and Michoacan, Mexico (Fig. 18), 100-1740
m. Flowering dates, Aug-Feb.
Recent collections frem Nayarit, Mexico, show much var-
iation and intergradation in the leaf and inner involucral
bract characters used by Blake (1926) to separate M. tepi-
cense from M. cornutum. In my opinion they are conspe-
cific.
REPRESENTATIVE SPECIMENS. Mexico. COLIMA: SW foothills of
Nevado de Colima, 1-1.5 mi S of Hacienda San Antonio, 11 Aug 1957,
McVaugh 16112 (MiCH). JALISCO: ea 10 km NE of Puerto Vallarta
near Milagro on rd to Mascota, 15 Nov 1963, Feddema 2577 (MICH);
10 mi S of Autlán, 19 Aug 1949, R. & C. Wilbur 2416 (MICH).
MICHOACÁN: Coalcoman, Parotas, 28 Sep 1938, Hinton et al. 12276
(MICH, NY, UC, US). NAYARIT: 10 mi SE of Ahuacatlán on rd to Bar-
"Similar to the appendage in M. longipilum, except here in M.
tepicense it is on the abaxial rather than the adaxial side of the
achene apex.
186 Rhodora [Vol. 74
ranca del Oro, 11-12 Aug 1959, Feddema 279 (MICH); 1 km N of El
Cuatante, ca 40 km NNE of Puerto Vallarta, 18 Nov 1963, Feddema
2651 (MICH); 8 mi W of Tepic, 10 Sep 1960, McVaugh 18906 (MICH) ;
9 mi N of Compostela, 12 Nov 1959, McVaugh & Koelz 542 (MICH).
30. Melampodium sinaloense Stuessy, Brittonia 22:122.
f. 10. 1970. TYPE: MEXICO: Sinaloa, Quebrado de Mansana,
Sierra Surotato, 4000-4500 ft, 10-14 Sep 1941, H. S. Gentry
6507-A (Holotype, GH ! ; isotypes, ARIZ! DS! MICH! MO! NY!).
Annual herbs, 25-35 em tall. Stems decumbent, 1-2 mm
diam, sparsely tomentose to moderately hispidulous with
hairs 0.1-0.3 mm long. Peduncles 1-2.9 cm long. Leaves
with petioles 10-15 mm long, ovate, 3.5-5 cm long, 2-2.5 em
wide, at apex acute, at base attenuate, with both surfaces
sparingly strigose with hairs 0.3-1.2 mm long; margin ser-
rate. Heads 3-4 mm tall, 3-4 mm diam. Outer involucre
cupulate, 5-10 mm diam; bracts 5, slightly connate at base,
separate, narrowly elliptic to elliptic, 2.5-6 mm long, 1.2-
1.5 mm wide, at apex obtuse, with abaxial surface with
2-5 hairs 0.3 mm long on midrib and major lateral veins;
margin herbaceous. Fruits 1.8-2 mm long, with lateral
surfaces ridged. Ray florets 6; ligules pale-yellow, elliptic,
1.5-Z mm long, 1-1.2 mm wide. Disc florets ca 14; corollas
pale-yellow, 0.8 mm diam, with throat 0.7 mm and tube
0.6 mm long. Paleae elliptic, 1.3 mm long, 0.6 mm wide;
apex pale-yellow, with margin irregularly and weakly den-
tate; midrib weak, glabrous. Chromosome number un-
known.
Known only from the type collection from oak forests in
Sinaloa, Mexico (Fig. 18), 1220-1370 m. Flowering date,
Sep.
IV. Melampodium section Bibractiaria Stuessy, sect. nov.
Herbae annuae radicibus palaribus; folia ovata vel ob-
ovata, sessilia vel subsessilia, marginibus integris; invol-
ucrum extimum cupulatum, bracteis 2, marginibus herba-
ceis vel leniter scariosis; ovaria flosculi disci ovoidea, minus
quam 0.7 mm longa, rudimentales (Figs. 3 & 4) ; fructus
apice fere laeves vel moderate sculpti et sine cucullis ;
1972] Melampodium — Stuessy 187
chromosomatum numerus basicus ignotus. Species 31 &
32. Typus: Melampodium bibracteatum S. Wats.
31. Melampodium bibracteatum S. Wats. Proc. Amer.
Acad. Arts & Sci. 26:140. 1891. TYPE: MEXICO: México, Del
Rio [ca 13 mi N of Toluca], 30 Aug 1890, C. G. Pringle
3230 (Holotype, US!; isotypes, DS! F! G[2]! GH! MA! MO!
MSC! NY [2]! P! uc! US! W!; photograph of holotype, TEX!;
photograph of K isotype, US!).
Annual herbs, 2-32 cm tall. Stems ascending to decum-
bent, 1.8-3.5 mm diam, glabrous below to pubescent above
with hairs 0.2 mm long. Peduncles 0-3.5 mm long (heads
often sessile). Leaves sessile, with blades oblong to ob-
ovate or oblanceolate, 0.9-4 cm long, 0.3-1.8 cm wide, at
apex obtuse-acute, at base obtuse-subauriculate to some-
what connate, with both surfaces with a conspicuous midrib
(very large basally) extending 1/3-1/2 of blade, with upper
surface glabrous, with lower surface subglabrous; margin
entire to 6-lobed, moderately ciliate with hairs 0.1 mm long.
Heads 3.5-4.5 mm tall, 3.5-4.8 mm diam. Outer involucre
cupulate, 4-7 mm diam; bracts 2, separate, ovate, 3-4.2 mm
long, 1.5-2.5 mm wide, at apex acute, with abaxial surface
glabrous; margin herbaceous to weakly scarious, rarely
with several hairs 0.2-0.5 mm long. Fruits 2-2.7 mm long,
with lateral surfaces smooth with several conspicuous
nerves. Ray florets (3-) 5-6; ligules yellow, elliptic-ovate, 1
mm long, 0.4 mm wide. Disc florets 4; corollas yellow, 0.8
mm diam, with throat 0.7 mm and tube 0.5 mm long.
Paleae oblanceolate, 1.5 mm long, 0.6 mm wide; apex color-
less, with margin irregularly laciniate; midrib very weak,
glabrous. Chromosome number unknown.
Mainly in low, moist areas in pine-oak forests of central
Mexico (Hidalgo, México and Puebla) but populations oc-
eur also in southwestern Durango and in Guatemala (Fig.
20), 2140-3360 m. Flowering dates, Aug-Oct.
REPRESENTATIVE SPECIMENS. Guatemala. HUEHUETENANGO: Sierra
de los Cuchumatanes, 1.5 mi W of Llano de San Miguel on trail to
Todos Santos, 2 Aug 1960, Beaman 3977 (GH, TEX, UC). Mexico.
188 Rhodora [Vol. 74
DURANGO: 10 mi W of El Salto, 2 Oct 1962, Cronquist 9585 (NY); 3 mi
SW of El Salto, 12 Aug 1961, Waterfall 16198 (SMU); 57 mi SW
of Durango, 13 Aug 1957, Waterfall & Wallis 13702 (smu, Us).
HIDALGO: near Buena Vista Station, 10 Aug 1904, Pringle 13068
(ARIZ, F, GH, SMU, US). MEXICO: Tultenango Station, 14 Sep 1901,
Pringle 9301 (F, GH, K, NY, US); !? mi W of Station Del Rio, 18 Aug
1966, Stuessy 675 (TEX); 7 mi W of Toluca, 19 Aug 1966, Stuessy
676 (TEX). PUEBLA: near Cerro El Pinar, 28 Aug 1945, Alexander &
Hernández 2229 [XA 218] (Nv); Laguna de San Baltasar, vicinity
of Puebla, 20 Sep 1966, Arsène 294 (US).
32. Melampodium repens Sessé & Moc. Fl. Mex. ed. 2. 195.
1894. TYPE: MEXICO: Veracruz, *Habitat in medietate Vul-
cani altissimi de Orizava," Jun 1795-1804, M. Sessé et al.
“3981” (Holotype, MA!; photograph of holotype, 0S! TEX!).
Melampodium arvense Robins. Proc. Amer. Acad. Arts
& Sci. 36:464. 1901. TYPE: MEXICO: Distrito Federal, Val-
ley of Mexico, 7500 ft, 19 Oct 1896, C. G. Pringle 7327
(Holotype, GH!; isotypes, F! POM! US!; photograph of US
isotype, TEX!).
Annual herbs, 2-30 cm tall. Stems prostrate (often form-
ing a mat), 0.8-1.8 mm diam, puberulent with hairs 0.2
mm long. Peduncles 0-3 mm long (heads often sessile).
Leaves sessile or with petioles 2 mm long, obovate, 0.5-2.5
cm long, 0.3-1.6 cm wide, at apex acute-cbtuse, at base at-
tenuate-obtuse to somewhat connate, with both surfaces
with a conspicuous midrib (very large basally) extending
less than 1/3 of blade, with upper surface glabrous, with
lower surface subglabrous; margin obscurely crenate,
sometimes moderately ciliate with hairs 0.1 mm long. Heads
2-3 mm tall, 3.5-4 mm diam. Outer involucre cupulate, 4-5
mm diam; bracts 2, separate, cvate, 3.5 mm long, 1.8-2.2
mm wide, at apex acute, abaxial surface glabrous; margin
herbaceous. Fruits 2-2.5 mm long, with lateral surfaces
smooth with several conspicuous nerves. Ray florets 2-3;
ligules yellow, elliptic-oblong, 0.8-1.2 mm long, 0.3 mm wide.
Disc florets 6; corollas yellow, 0.5 mm diam, with throat
0.4 mm and tube 1.0 mm long. Paleae oblanceolate, 1.2 mm
long, 0.4 wide; apex colorless, with margin laciniate; mid-
rib very weak, glabrous. Chromosome number unknown.
1972] Melampodium — Stuessy 189
Pine-oak forests in the Mexican states of Distrito Fed-
eral, México, Michoacan, and Veracruz (Fig. 20), 1380-
3150 m. Flowering dates, Aug-Sep.
The two species of section Bibractiaria are perhaps the
most unusual taxa of the whole genus, particularly in their
over-all diminutive appearance. Chromosome counts have
not been obtained yet from these two species, although
collections have been made and counts attempted. In view
of the wide diversity of chromosome numbers already found
in Melampodium, it will be interesting to learn if new num-
bers may be obtained from these two taxa.
REPRESENTATIVE SPECIMENS. Mexico, DISTRITO FEDERAL: Cima, 24
Aug 1910, Orcutt 3780 (F); Eslava, Valley of Mexico, 7 Sep 1901,
Pringle 8610 (ARIZ, F, G, GH, K, MSC, NY, POM, SMU, TEX, UC, US).
DURANGO [?]: “in mountains near Santa Angela,” Sep 1855, Schaff-
ner s.n. (GH). MÉXICO: Parque Nacional de Laguna Zempoala,
26 Jul 1947, Barkley, Webster & Rowell 7438 (TEX); Temascaltepec,
Cucha, 4 Sep 1933, Hinton 4628 (GH, NY, us); Temascaltepec, Teju-
pilco, 27 Sep 1933, Hinton 4826 (GH, K, NY, US); Toluca, 26 Sep
1892, Pringle 5257 (GH); Salto de Agua, Oct 1905, Purpus 1809
(F, UC); Popocatépetl, 22 Aug 1901, Rose & Hay 6244 (NY, US);
Llano Grande, near Rio Frio, 28 Jul 1944, Sharp 44116 (NY); 2 mi
E cf Cuajimalpa, 18 Aug 1966, Stuessy 670 (TEX). MICHOACAN:
SW side of Cerro San Andres, ca 12 km N of Ciudad Hidalgo, 6 Sep
1960, Beaman 4352 (GH). VERACRUZ: Loma Grande, 27 Aug 1938,
Balls & Gourlay B5372 (x).
V. Melampodium section Rhizomaria Stuessy, sect. nov.
Plantae perennae rhizomatosae; folia anguste ovata vel
oblonga, sessilia vel interdum sessilia, marginibus integris
vel subintegris; involucrum extimum patens vel aliquan-
tum cupulatum, bracteis 5, marginibus scariosis; ovaria
flosculi disci ovoidea, minus quam 0.7 mm longa, rudimen-
tales (Figs. 3 & 4); fructus apice fere laeves vel moderate
sculpti et sine cucullis; chromosomatum numerus basicus,
x = 11. Species 33 & 34. Typus: Melampodium montanum
Benth.
33. Melampodium montanum Benth. Pl. Hartw. 64. 1840.
Suffrutescent perennials. Stems erect to ascending.
Leaves usually sessile, ovate to elliptic, at apex obtuse
190 Rhodora [Vol. 74
(rarely acute), at base obtuse to subauriculate. Outer in-
volucre spreading or somewhat cupulate; bracts 5, connate
1/5 their length, imbricate, ovate, 4-5 mm long, 4-5 mm
wide, at apex acute, with margin scarious. Fruits 1.6-1.8
mm long, with lateral surfaces smooth to nervose or mar-
ginally tuberculate. Ray florets 9-13; ligules yellow, nar-
rowly elliptic-oblong. Disc florets 80-110; corollas yellow-
green, with tube 0.8 mm long. Paleae oblanceolate, 2 mm
long, 0.6 mm wide; apex yellow, with margin entire; mid-
rib prominent, glabrous.
33a. Melampodium montanum Benth. var. montanum
Melampodium montanum Benth. Pl. Hartw. 64. 1840.
TYPE: MEXICO: Oaxaca, “In summo jugo (Cumbre) inter
Oaxaca et la Sierra,” 1839, C. T. Hartweg 475 (Holotype,
K!; photograph of holotype, os! TEX! US!).
Melampodium liebmanii Sch. Bip. ex Klatt, Leopoldina
23:89. 1887. TYPE: MEXICO: Oaxaca, “Cumbre de Estepa
et Yavesía," Jun 1842, F. M. Liebmann 232 (Lectotype, C;
isotype, P!; photograph of lectotype, US!; photograph of C
isotype, F! NY! us [2] !; photograph of P isotype, 0S! TEX!;
drawing and presumed fragment of lectotype, GH !).
Plants 6-28 em tall. Stems 0.8-1.2 mm diam, copiously to
sparsely villous with hairs 0.8-1.5 mm long. Peduncles 3-
7.5 cm long. Leaves sessile or with petioles 1 mm long, 1.6-4
cm long, 0.8-1.7 em wide, with both surfaces strigose with
hairs 0.8 mm long; margin entire to obscurely denticulate.
Heads 5-6 mm tall, 11-21 mm wide. Outer involucre 8-13
mm diam; bracts at apex purple-tinged, with abaxial sur-
face villous with hairs 1 mm long. Ligules at apex and on
veins on undersurface dark purple, 4-7.5 mm long, 1.5-2.5
mm wide. Disc florets 1 mm diam, with throat 1.4 mm
long. Chromosome number, n = ca 11.
Pine-oak forests in north-central Oaxaca, Mexico (Fig.
26), 1740-3360 m. Flowering dates, Jul-Aug.
1972] Melampodium — Stuessy 191
Fig. 26. Map of Mexico and adjoining Guatemala showing dis-
tribution of Melampodium aureum (squares), M. montanum var.
montanum (dots), and M. montanum var. viridulum (triangles).
REPRESENTATIVE SPECIMENS. Mexico, OAXACA: Cerro San Felipe,
18 Aug 1921, Conzatti 4169 (US) ; Jul-Aug 1900, Conzatti & González
998 (GH); vicinity of Cerro Zempoaltepet| [Zempoaltepec], 10 Aug
1950, Hallberg 900 (Us); ca 10 mi N of jet rtes 190 & 175, 26 Jul
1960, King 3492 (ps, NY, TEX, UC, US); NW side of summit of Mt
Zempoaltepec, 9 Jul 1894, Nelson 655 (US); Sierra de San Felipe,
28 May 1894, Pringle 4666 (GH, Us).
33b. Melampodium montanum Benth. var. viridulum Stues-
Sy, var. nov.
Plantae 10-35 cm altae. Caules 0.8-1.4 mm diametro, vil-
losi pilis 1-1.5 mm longis (raro glabri). Pedunculi 1-6.5
cm longi. Folia subsessilia (vel petiolis 2-3 mm longis),
laminae 1.6-3.3 longae, 0.7-1.4 em latae, utrinque strigosae
vel moderate vilosae pilis 1-1.5 mm longis (subtus raro
glabrae); margines integri. Capitula 4.5-6 mm alta, 12-17
mm diametro. Involucrum extimum 8-14 mm diametro;
bracteae apice interdum purpurascentes, extus villosae pilis
192 Rhodora [Vol. 74
1 mm longis (interdum subglabrae). Ligulae subtus apice
venaque viridulae, 4-5 mm longae, 1.2-1.9 mm latae. Flos-
culi disci 0.8 mm diametro, faucibus 1.2 mm longis. Chro-
mosomatum numerus, n = 11.
TYPUS: MEXICO: Chiapas, ca 5 mi E of San Cristóbal
de Las Casas on rte 190, 11 Jun 1960, R. M. King 2801
(Holotype, US!; isotypes, Ds! NY! TEX! UC!).
Scattered in pine-oak forests in several Mexican states
in the Sierra Madres Oriental, Occidental and Sur, and in-
cluding Chiapas and northwestern Guatemala (Fig. 26),
610-2740 m. Flowering dates, Jun-Oct.
REPRESENTATIVE SPECIMENS. Guatemala. HUEHUETENANGO: Sierra
de los Cuchumatanes, between kms 324 & 325 on rte 9N (between
Chemal & San Juan Ixcoy), 4 Aug 1959, Beaman 3043 (GH, NY, UC);
San Pedro Coloma, 20 Aug 1934, Skutch 1036 (F, GH, US); SW
slopes of Sierra de los Cuchumatanes, between Chiantla & Patio de
Bolas, 6 Jul 1942, Steyermark 48234 (F); along Río Selegua, oppo-
site San Sebastián H., !3 Aug 1942, Steyermark 50490 (F, GH).
Mexico. CHIAPAS: Jitotol, 3 mi S of Jitotol, 12 Feb 1965, Breedlove
8906 (ns); Tenejapa, Tuk, Matsab, 30 Sep 1965, Breedlove 12522
(ps); Comitán, 30 Apr 1904, Goldman 914 (US) ; ca 17 mi W of San
Cristóbal de Las Casas, 9 Jun 1960, King 2796 (DS, NY, TEX, UC, US) ;
ca 1 mi E of Teopisca, 12 Jun 1960, King 2843 (TEX); San Cristóbal
de Las Casas, Cerro San Cristóbal, 11 Jun 1966, Laughlin 1050 (DS) ;
Amatenango del Valle, 12 Jun 1945, Matuda 6002 (F); 34 mi S of
Ishuatán on gravel rd to Tuxtla Gutiérrez, 24 Jul 1966, Stuessy 559
(TEX); 20 mi S of San Cristóbal, 25 Jul 1966, Stuessy 566 (TEX);
8 mi S of San Cristóbal, 25 Jul 1966, Stuessy 567 (TEX); 2 mi SE
of Teopisca, 25 Jul 1966, Stuessy 568 (TEX); 9 mi SE of Tecpisca,
25 Jul 1966, Stuessy 570 (TEX). GUERRERO: Taxco, 23 Jul 1936,
Abbott 162 (GH). JALISCO: Sierra del Tigre, 3 mi S of Mazamitla,
18 Sep 1952, MeVaugh 13033 (SMU, US). MÉXICO: Temascaltepec,
Bejucos, 7 Oct 1932, Hinton 2017 (ARIZ, F, MA, NY [2], vs).
MICHOACÁN: Morelia, Cerro Azul, 1910, Arséne 6575 (US). OAXACA:
15 km S of Sola de Vega, on rd to Puerto Escondido, 30 Sep 1965,
Rzedowski 21322 (TEX). PUEBLA: vicinity of Puebla, woody hill
farther than Cerro Tepoxuchitl, 16 Sep 1907, Arséne 1896 (GH, NY,
us); hills near Amozoc, 10 Sep 1901, Pringle 9361 (F, GH, US).
QUERÉTARO: 8 km W of El Lobo, municipio de Landa, 31 Aug 1957,
Rzedowsk: 9320 (ENCB). SAN LUIS POTOSÍ: “barranca of Las Canoas,”
18 Aug 1891, Pringle 3818 (ps, F, GH [2], Ny [2], vc [2], US).
TAMAULIPAS: Gómez Farías Region, between Rancho del Cielo &
Charco de los Perros, spring 1965, Webster 140 (TEX).
1972] Melampodium — Stuessy 193
34. Melampodium aureum Brandg. Univ. Calif. Pub. Bot.
4:94. 1910. TYPE: MEXICO: Puebla, Cerro de Gavilan, 8000-
9000 ft, Aug 1909, C. A. Purpus 3822 (Holotype, UC!; iso-
types, F! G! GH! MO! NY! US!; photograph of US isotype,
TEX!).
Suffrutescent perennials, 10-35 cm tall. Stems erect to
ascending (occasionally decumbent), 0.8-2 mm diam, mod-
erately villous with hairs 0.5-1 mm long. Peduncles 4.5-11.5
cm long. Leaves usually sessile or with petioles 2-3 mm
long, ovate to weakly spatulate, 2.7-6 em long, 1-2.7 em
wids, at apex obtuse to acute, at base obtuse to subauricu-
late, with both surfaces strigose with hairs 0.5-0.8 mm long;
margin entire to serrulate, ciliate with hairs 0.3 mm long.
Heads 6-7 mm tall, 19-38 mm diam. Outer involucre spread-
ing or somewhat cupulate, 10-14 mm diam; bracts 5, con-
nate 1/5 their length, imbricate, ovate, 4-6 mm long, 3.5-
6 mm wide, at apex acute and sometimes purple-tinged,
with abaxial surface villous with hairs 0.5 mm long (some-
times mostly glabrous) ; margin scarious. Fruits 2-2.3 mm
leng, with lateral surfaces smooth to rugose-aculeate. Ray
florets 11-14; ligules yellow-orange to yellow, narrowly el-
liptic-oblong to oblanceolate, 5-13 mm long, 2-3 mm wide.
Disc florets 80-110; corollas yellow, 1 mm diam, with throat
1.8 mm and tube 0.7 mm long. Paleae oblanceolate, 3.2
mm long, 0.6 mm wide; apex yellow, with margin entire;
midrib prominent, glabrous. Chromosome number, n =
Scattered in pine-oak forests in Nuevo León, Michoacán,
Oaxaea, and Pueb!a, Mexico (Fig. 26), 1800-2500 m.
Flowering dates, Jul-Sep.
The morphology of the hexaploid M. aureum suggests
that it may have been derived from the diploid M. mon-
tanum by autopolyploidy, perhaps through the union of un-
reduced gametes from diploid and tetraploid plants. The
two taxa are so similar, in fact, that experimental studies
may indicate that M. aureum should be treated as simply
a polyploid race of M. montanum. The extant chromosomal
voucher specimens, however, do show a considerable amount
194 Rhodora [Vol. 74
of morphological divergence albeit of a quantitative na-
ture. In view of these morphological differences and in
the absence of intensive field and laboratory studies, I
prefer to maintain the two taxa as distinct species at the
present time.
REPRESENTATIVE SPECIMENS. Mexico. MICHOACAN: vicinity of
Morelia, Cerro San Miguel, 15 Sep 1910, Arsène 5219 (GH, NY, US);
ca 20 mi W of Ciudad Hidalgo, 4 Aug 1960, King 3617 (DS, NY, TEX,
uc, Us); 23 mi NW of Ciudad Hidalgo, 19 Aug 1966, Stuessy 682
(TEX); 21 mi NW of Ciudad Hidalgo, 19 Aug 1966, Stuessy 684
(TEX). NUEVO LEÓN: Dulces Nombres, 20 Jul 1948, Meyer & Rogers
2828 (vs), 10 Aug 1948, 2911 (US). OAXACA: Nochistlán, Cuesta
Blanca, rd to Montelobos, 22 Jun 1907, Conzatti 1865 (F); ca 8 mi
SE of Nochistlan, 27 Jul 1960, King 3529 (DS, NY, TEX, UC, US);
7 mi SE of Nochistlán, 15 Aug 1966, Stuessy 663 (TEX). PUEBLA:
Cerro del Oro, Aug 1909, Purpus 3822a (F, GH, NY, UC, US).
VI. Melampodium section Alcina (Cav.) DC.
Melampodium L. sect. Alcina (Cav.) DC. Prodr. 5:520.
1836. Alcina Cav. Ic. 1:10. t. 15. 1791. Type species:
Melampodium perfoliatum (Cav.) H.B.K.
Tap-rooted annuals; leaves ovate or oblong, sessile or
petiolate, with margins entire or weakly serrate; outer in-
volucre spreading, with bracts 3-5, at margins herbaceous,
ovaries of the disc florets ovoid, less than 0.7 mm long, rudi-
mentary (Figs. 3 & 4); fruits at apex nearly smooth or
moderately sculptured and without hoods; chromosome
base number, x — 11. Species 35-37.
35. Melampodium perfoliatum (Cav.) H.B.K. Nov. Gen.
Sp. 4:274. 1820.
Alcina perfoliata Cav. Cav. le. 1:11. f. 15. 1791. TYPE:
“IMPERIO MEXICANO": cultivated in Bot. Gard. Madrid,
flowering 29 Nov (Lectotype chosen, MA!; isotypes, MA
[2] !; photograph of lectotype, OS! TEX!).
Wedelia perfoliata (Cav.) Willd. Sp. Plant. 3(3) :2335.
1803.
Polymnia perfoliata (Cav.) Poiret, Encycl. Méthod.
Botan. 5:506. 1804.
1972] Melampodium — Stuessy 195
Camutia perfoliata Bonato ex Steud. Nom. Bot. 146. 1821.
pro syn.
Melampodium connatum Sessé & Moc. ex DC. Prodr.
5:521. 1836. pro syn.
Polymnia revoluta Autor. ex Steud. Nom. Bot. ed. 2. 2:
377. 1841. pro syn.
Annual herbs, 20-150 cm tall. Stems erect, 1-11 mm diam,
glabrous below to very minutely puberulent above. Pe-
duncles 0.6-11 em long. Leaves sessile, rhombic-deltoid
(rarely ovate-obovate) and tapering to an auriculate-con-
nate base (sometimes obtuse on upper leaves), 2-21 em
long, 0.9-15 em wide, at apex acute, with both surfaces
strigose with hairs 0.2-0.3 mm long; margin obscurely to
strongly irregularly serrate. Heads 5-7 mm tall, 8-11 mm
diam. Outer involucre spreading, 15-32 mm diam; bracts
5, slightly imbricate and connate at base, oblong-elliptic,
6-20 mm long, 4-11 mm wide, at apex obtuse, with abaxial
surface occasionally strigose with hairs 0.2-0.5 mm long ;
margin herbaceous. Fruits 4-7 mm long, with lateral sur-
faces smooth to moderately striate or nervose. Ray florets
8-13; ligules yellow-orange, oblong-elliptic, 2.5-4 mm long,
0.9-2.3 mm wide. Disc florets 30-45; corollas yellow, 1.5
mm diam, with throat 1 mm and tube 0.8 mm long. Paleae
oblong-elliptic, 2.1 mm long, 0.7 mm wide; apex light yel-
low, with margin dentate-laciniate; midrib distinct, gla-
brous. Chromosome numbers, n = 11 & 12.
Tropical deciduous and pine-oak forests in Mexico, Guate-
mala, and Costa Rica, introduced to Cuba and Los Angeles,
California (Fig. 21), 610-2440 m. Flowering dates, Jan-
Dec.
M. perfoliatum is the second most widely spread species
in the genus (after M. divaricatum), but apparently it has
been introduced only at one point in the United States
(near Los Angeles). Because the city of Los Angeles and
surrounding suburbs have grown tremendously in the last
196 Rhodora [Vol. 74
Fig. 27. Map of Mexico, Central America and neighboring coun-
tries showing distribution of Melampodium perfoliatum.
60 years, it is likely that M. pei foliatum, however weedy
and aggressive, has not managed to survive. Specimens ex-
amined from Los Angeles Co., California, support this con-
tention, as there are no records later than 1902.
At maturity M. perfoliatum is very easy to recognize
because the large fruits remain temporarily in a conspicu-
ous ring within the spreading foliaceous outer phyllaries.
In the field the bases of the lower leaves are markedly per-
foliate and also help distinguish the species. On herbarium
specimens, however, this latter characteristic is less help-
ful because the bases of the upper leaves are sometimes
obtuse, and such leaves, due to their more convenient size,
most frequently are selected and pressed by collectors.
1972] Melampodium — Stuessy 197
An interesting nomenclatural problem is associated with
the earliest date of publication for the combination Melam-
podium perfoliatum. It has generally been attributed to
H.B.K. in 1820. An anonymous author (perhaps M. A.
Valdes, fide Langman, 1964) in the Gazeta de México of
1792-93 (vol. 5, p. 243) reported the events taking place at
a public botanical examination in La Real y Pontifica Uni-
versidad in Mexico City on December 7, 1792. At this
gathering a student, Sebastian Gomez Moron, read a paper
in which he “. . . determinó y describió con la mayor prop-
iedad el Melampodio reunido, o Melampodium perfoliatum.
... A footnote elaborates in some detail the circumstances
surrounding this new combination which precedes that of
H.B.K. by at least 27 years. According to the anonymous
author, it was the “opinion” of the “Expedición” (i.e.,
Sessé, Castillo, and Mocino) traveling through Mexico, that
Alcina perfoliata of Cavanilles should not be generically
separated from the well-known Melampodium. The stu-
dent, Gómez Moron, apparently looked into the matter more
closely and agreed with this contention. The author adds
another sentence in the footnote, however, that changes the
nomenclatural complexion of the situation: ‘‘Los Botanicos
que tuvieren ocasion de cotejar la Planta, 6 la exacta de-
scripcion de este autor [Cavanilles] con qualquiera de los
Melampodios conocidos, pesarán los fundamentos que me-
dian entre una y otra parte, y fixarán el nombre con que
debe conocerse en lo sucesivo esta nueva especie." The au-
thor states that botanists who might compare the species
with the existing taxa of Melampodium will have to weigh
the facts and decide what the correct name should be. I
interpret this hesitation to mean that the author did not
accept the new combination at this time, which therefore
precludes its validation (Article 34, Provisions 1 and 2,
Intern. Code Bot. Nomen.; Lanjouw et al. 1966).
REPRESENTATIVE SPECIMENS. Costa Rica. ALAJUELA: between San
Josecito & La Garita, 18 Aug 1963, Jiménez 1074 (F). CARTAGO:
vicinity of Turrialba, 5 Feb 1965, Godfrey 66184 (FSU); Turrialba,
198 Rhodora [Vol. 74
Instituto Interamericano de Agricolas, 20 Aug 1962, King 5350 (TEX,
uc, US); 4 km E of Cartago, 28 Aug 1962, King 5407 (TEX, UC, US).
SAN JOSE: Rio Torres, San José, 31 Jan 1965, Godfrey 66064 (FSU).
Cuba. LA HABANA: Santiago de las Vegas, 30 Jun 1904, Baker &
Wilson 516 (F, NY, UC, US); Laguna de Ariguanabo at Cayo La Rosa,
17 Jul 1923, Ekman 16932 (NY); Santiago de las Vegas, 10 Oct 1904,
Hermann 148 (F, NY, US); Laguna de Castellano, 27 Dec 1910,
Wilson 9553 (NY [2]). PINAR DEL RIO: near Potrerito, Sumidero,
19 May 1951, Alain 1819 (NY). Guatemala. GUATEMALA: city limits
of Guatemala, 9 Jul 1960, King 3248 (DS, NY, TEX, UC, US); Guate-
mala City, 1 Aug 1966, Stuessy 603 (TEX). HUEHUETENANGO: ca 1
mi S of Huehuetenango, 18 Jul 1960, King 3410 (DS, NY, TEX, UC, US) ;
2 mi S of Huehuetenango, 26 Jul 1966, Stuessy 580 (TEX). QUICHÉ:
1942, Aguilar 1942 (F). Mexico, AGUASCALIENTES: below Presa
Calles, 31 Aug 1939, Shreve 9278 (ARIZ, GH). CHIHUAHUA: Arroyo
Hondo, Sierra Charuco, 11 Sep 1985, Gentry 1752 (ARIZ, F, GH, NY,
uc, US); Municipio de Guadalupe & Calvo, Nabogame, 11 Oct 1959,
Gentry, Correll & Arguelles 17970 (US). COLIMA: Colima, 9 Jan-
6 Feb 1891, Palmer 1163 (GH, NY, US); Alzada, 25 Aug 1966, Stuessy
729 (TEX). DISTRITO FEDERAL: Mixcoac, Olivar, 7 Sep 1913, Arsène
s.n. (US); San Andrés, Aug 1930, Lyonnet 569 (US); Pyramid of
Cuicuileo, Tlalpan, 15 Aug 1935, MacDaniels 41 (F); near Tlalpan,
1 Sep 1936, MacDaniels 723 (F); San Ángel, 12 Aug 1910, Orcutt
3548 (F). DURANGO: vicinity of Durango, Jun 1896, Palmer 286
(F, GH, NY, UC, US) ; ca 30 mi N of Durango, 12 Nov 1959, Gentry &
Arguelles 18152 (US). GUANAJUATO: Guanajuato, Dugés 461 (GH),
462 (GH); Obregón, 13 Oct 1895, Volkens 1134 (GH, NY). GUERRERO:
Taxco, 28 Sep 1937, Abbott 445 (GH), 5 Sep 1959, Degener 26220
(NY); Mina, Zihuaqueo-Filo Mayor, 20 Aug 1936, Hinton et al. 9293
(GH, US). JALISCO: La Barranca, 24 Nov 1930, Jones 27773 (ns [2],
F, NY, POM, UC); NE slopes of Nevado de Colima, below Canoa de
Leoncito, 17 Oct 1952, McVaugh 13576 (SMU, Us); near Santa Monia,
12-13 Nov 1952, MeVaugh 14107 (smu, Us); near Guadalajara,
29 Oct 1888, Pringle 1767 (F, GH, NY [2], UC, US); 2 mi NW of La
Barca, 23 Aug 1966, Stuessy 705 (TEX). MEXICO: Valley of México,
Tizapán, 12 Feb 1865-66, Bourgeau 841 (GH); Temascaltepec, Temas-
caltepec, 2 Sep 1932, Hinton 1521 (NY); Temascaltepec, Chorrera,
25 Sep 1932, Hinton 1838 (GH, NY, US); Temascaltepec, Ixtapán,
2 Nov 1932, Hinton 2480 (Ny, UC, US); % mi N of Tepetlixpa, 16
Aug 1966, Stuessy 668 (TEX). MICHOACÁN: 13 mi S of Uruapán,
28 Oct 1962, Cronquist 9745 (NY, SMU, TEX, US); ca 11 mi W of the
Michoacán-Mexico state border on rte 15, 3 Aug 1960, King 3600
(DS, NY, SMU, TEX, UC, US); E of San Juan Nuevo (ca 8 km S of
Uruapán), 11-15 Oct 1961, King & Soderstrom 4707 (NY, SMU, TEX,
UC, US); 2 mi W of Ciudad Hidalgo, 21 Aug 1961, Powell & Edmondson.
1972] Melampodium — Stuessy 199
816 (DS, F, TEX); 0.8 mi NW of Zitacuaro, 1 Sep 1965, Stuessy 379
(TEX). MORELOS: vicinity of Cuernavaca, 28 Jan 1932, Asplund 517
(NY), 31 Aug 1910, Orcutt 3892 (F, US). NAYARIT: vicinity of Jalisco,
10 Nov 1925, Ferris 5890 (DS, GH, US); Tepic, 14 Feb 1927, Jones
23398 (pom), 10 Feb 1927, Jones 23402 (POM). PUEBLA: vicinity of
Puebla, Manzanilla, Nov 1908, Arsène 3523 (US); ca 5 mi W of
Puebla, 29 Jul 1960, King 3560 (NY, TEX, UC, US) ; vicinity of Puebla,
Hacienda Batan, 20 Dec 1911, Nicolas 6150 (GH, US); Teteles, 9 Jul
1966, Stuessy 497 (TEX); 3 mi W of Puebla, 14 Jul 1966, Stuessy
503 (TEX). OAXACA: Cerro San Felipe, 20 Sep 1908, Conzatti 2291
(F); vicinity of Totontepec, 15-20 Jul 1894, Nelson 715 (US); Sierra
de San Felipe, Oct 1894, Smith 351 (F); N city limits of Zimatlán,
14 Aug 1966, Stuessy 654 (TEX). SAN LUIS POTOSÍ: 32 mi E of San
Luis Potosí, 19 Sep 1960, Crutchfield & Johnston 5643a (TEX); near
San Luis Potosí, Aug 1876, Schaffner 270 in part (GH, NY). SINALOA:
Ocurahui, Sierra Surotato, 1-10 Sep 1941, Gentry 6350 (ARIZ, DS,
GH, NY). VERACRUZ: Zacualpán, Dec 1907, Purpus 2849 (F, GH, NY,
UC, US) ; Mt Orizaba, 22 Aug 1891, Seaton 54 (F, GH, NY, us); 8 mi
NW of Jalapa, 9 Jul 1966, Stuessy 494 (TEX) ; 34 mi NW of Tehuacán,
14 Jul 1966, Stuessy 505 (TEX); 4% mi W of San Juan, near Pico de
Orizaba, 15 Jul 1966, Stuessy 511 (TEX). UNITED STATES. Cali-
fornia: LOS ANGELES CO.: Cienega, Aug 1902, Braunton 625 (DS, UC
[2], us); Los Angeles, Aug 1902 [037], Grant 763 (ARIZ, F, UC),
Sep 1879, James s.n. (NY, US); between Los Angeles & Compton,
Nov 1895, McClatchie s.n. (DS, NY, POM); Los Angeles, Oct 1882,
S. & W. Parish 1171 (ps, F, GH, NY, US [2]).
36. Melampodium glabrum S. Wats. Proc. Amer. Acad.
Arts & Sci. 26:139. 1891. TYPE: MEXICO: Guanajuato, val-
ley near Irapuato, 20 Sep 1889, C. G. Pringle 2821 (Holo-
type, GH!).
Annual herbs, 25-60 cm tall. Stems erect, 1-10 mm diam,
somewhat soft and watery when fresh, glabrous near base
and covered with bladdery hairs toward apex. Peduncles
2.5-5.5 em long. Leaves sessile, oblong-oblanceolate, 2.5-6
cm long, 0.7-2.1 em wide, at apex obtuse, at base subauricu-
late, with both surfaces waxy and glabrous; margin sinu-
ate-dentate. Heads 4.3-5 mm tall, 7-11 mm wide. Outer
involucre spreading, 8-10(14) mm diam; bracts 5, fleshy,
slightly connate at base, imbricate, ovate-orbiculate, 4.2-6
mm long, 3.8-4.5 mm wide, at apex obtuse with abaxial sur-
face glabrous; margin herbaceous. Fruits 2.5-2.9 mm long,
200 Rhodora [Vol. 74
with lateral surfaces smooth to moderately striate or ner-
vose. Ray florets 6-8; ligules yellow, oblong-ovate, 1.3-3.4
mm long, 0.7-1.4 mm wide. Disc florets 30-50; corollas
yellow, 1 mm diam, with throat 0.9 mm and tube 0.8 mm
long. Paleae oblanceolate-elliptic, 1.6 mm long, 0.5 mm
wide; apex yellow, with margin entire; midrib prominent,
glabrous. Chromosome number, n = 11.
Very moist areas in pine-oak forests or high altitude
grasslands in the Mexican states of Guanajuato, Jalisco,
and Michoacan (Fig. 13), ca 1890 m. Flowering dates, Aug-
Sep.
This species is an attractive, fleshy, little herb found
often in moist depressions on roadsides. The large folia-
ceous outer involucral bracts and conspicuous blackish
fruits as well as the n — 11 chromosome number indicate
that M. glabrum is allied closely to M. perfoliatum.
REPRESENTATIVE SPECIMENS. Mexico. GUANAJUATO: near San
Miguel Allende, Aug 1947, Kenoyer 2421 (GH). JALISCO: valley of
the Lerma near La Barca, 16 Sep 1891, Pringle 3863 (DS, G, GH
[2], MA, Msc, NY [2], P, vc [2], US, w); 3 mi N of La Barca, 24 Aug
1966, Stuessy 708, 709 (TEX). MICHOACÁN: vicinity of Morelia, 13
Sep 1910, Arsène s.n. (US); 9 mi S of jet rte 15 on rd to Cotija,
24 Aug 1966, Stuessy 714 (TEX).
37. Melampodium nutans Stuessy, Brittonia 22:122. f. 9.
1970. TYPE: MEXICO: Colima, 12 mi S of Colima on rte 110,
25 Aug 1966, T. F. Stuessy 721 (Holotype, US!; isotypes,
DS!F!GH! MICH! NY! TEX! UC!).
Annual herbs, 16-40 em tall. Stems erect, 1.2-2 mm diam,
hispidulous-strigillose with hairs 0.2-0.5 mm long. Pedun-
cles 5.5-6.5 cm long. Leaves ovate to rhombic, with taper-
ing petioles 1-3 cm long, blades 4-5 cm long, 2-4 cm wide,
at apex acuminate, at base attenuate, with both surfaces
strigose with hairs 0.5 mm long; margin subentire to ser-
rate. Heads 4-6 mm tall, 7-8(10) mm diam. Outer in-
voluere spreading, 8-14 mm diam; bracts 3 (rarely 4),
slightly connate at base, separate, ovate to narrowly ovate,
4-7 mm long, 2.5-4 mm wide, at apex acute, with abaxial
1972] Melampodium — Stuessy 201
surface strigose with hairs 0.3-0.5 mm long; margin her-
baceous. Fruits 1.8-2.1 mm long, with lateral surfaces
ribbed and reticulated; hood at apex muticous to cirrhous'*,
with tapering appendage up to 6 mm long. Ray florets 5-7;
ligules yellow-orange, orbiculate, margins imbricate, 2-3 mm
long, 2-4 mm wide. Disc florets 40-80; corollas yellow-
orange, 1.7 mm diam, with throat 0.7 mm and tube 1 mm
long. Paleae narrowly obovate, 2 mm long, 1 mm wide;
apex yellow, with margin laciniate; midrib distinct, pube-
rulent with hairs 0.1 mm long. Chromosome number,
n — 11.
Tropical deciduous forests of Colima, Jalisco, Michoacán
and Oaxaca, Mexico (Fig. 13), 50-500 m. Flowering dates,
Aug-Nov.
The hoods on fruits of M. nutans are shorter and
thicker than are the hoods of species of section Melampo-
dium. Such a difference is not unexpected, however, be-
cause M. nutans differs in many other respects from the
taxa in that section. Due to the spreading outer involucral
bracts, rudimentary disc ovaries, herbaceous habit, and
chromosome number of » — 11 of this species, it clearly
belongs in section Alcina near M. perfoliatum and M. gla-
brum.
REPRESENTATIVE SPECIMENS. Mexico. COLIMA: ca 11 mi SSW of
Cd Colima, 21 Sep 1958, McVaugh 18070 (MICH), 25 Nov 1959,
McVaugh & Koelz 1054 (MICH); 12 mi S of Cd Colima, 25 Aug 1966,
Stuessy 720 (TEX). JALISCO: between Sierra de los Corales & Tepal-
catepec, Michoacán, 26 Oct 1963, Feddema 2238 (MICH); Tecalitlan,
26 Oct 1963, Rzedowski 17514 (MICH, OS). MICHOACÁN: ca 5 km
NW of Aquila, 23 Nov 1963, Feddema 2700 (MICH); 15 mi SSW of
Apatzingán, 2 Aug 1940, Leavenworth 431 (F, GH, NY), 432 (F);
8 km NW of Aquila, 23 Nov 1963, Rzedowski 17921 (MICH, 0s).
OAXACA: 3 km SW of Puerto Ángel, 24 Sep 1965, Rzedowski 21155
(os).
“Isotype material in my preserved personal collection has fruits
with both muticous hoods and hoods with long cirrhous appendages
in the same head (cf. Fig. 9E in Stuessy, 1970b).
202 Rhodora [Vol. 74
DOUBTFUL SPECIES
Melampodium angulatum Sessé & Moc. Pl. Nov. Hispan.
149. 1890. TYPE: MEXICO: Guerrero, Ayahualtempa, Aug
1789, M. Sessé et al. s.n. (Holotype, MA?). From the de-
scription and locality data this taxon seems most similar to
M. perfoliatum, although the leaf undersurface in the latter
species is never “tomentose.”
Melampodium dichothomum Sessé & Moc. Pl. Nov. His-
pan. 149. 1890. TYPE: MEXICO: Guerrero, Mazatlán [ca 15
km S of Chilpancingo; Sprague (1926), and Rickett
(1947) ], Jun 1789, M. Sessé et al. “3977.” MA has a mixed
sheet with both M. montanum and M. divaricatum and an-
other sheet with M. divaricatum. A sheet at F has a frag-
ment of only M. divaricatum. The description more closely
resembles M. montanum, but both M. divaricatum and M.
montanum are found in Guerrero, and neither seems ex-
plicitly excluded by the description.
Melampodium diversifolium Sessé & Moc. Fl. Mex. ed. 2.
197. 1894. TYPE: locality unknown, 1787-1804, M. Sessé et
al. s.n. (Holotype, MA?). The description seems to refer to
M. linearilobum or M. longipes, but not mentioned are sev-
eral diagnostic characters such as the leaf undersurface
and margins of the outer involucral bracts.
Melampodium pinnatum Sessé & Moc. Pl. Nov. Hispan.
149. 1890. TYPE: MEXICO: Michoacan, Apatzingán, Oct
1790-91, M. Sessé, J. M. Mocino & J. Castillo s.n. (Holo-
type, MA?). The description is too general to allow any
specific placement.
Melampodium rhombifolium Sessé & Moc. Pl. Nov. His-
pan. 149. 1890. TYPE: MEXICO: México, San Augustín,
Sep 1787-1804, M. Sessé et al. s.n. (Holotype, MA?). The
character “Receptaculum nudum,” if correct, would exclude
this taxon from Melampodium.
Melampodium vhombifolium Sessé & Moc. Fl. Mexic. ed.
2. 195 [second on page]. 1894. nom. illegit. non Sessé &
Moc. 1890. nec Sessé & Moc. [first on page]. 1894. TYPE:
1972] Melampodium — Stuessy 203
MEXICO: “Hortis Sancti Augustini, prope Mexicum," Aug-
Sep 1787-1804, M. Sessé et al. s.n. (Holotype, MA?). Spe-
cific affinity cannot be determined from the description.
EXCLUDED NAMES
Melampodium L. sect. Acanthospermum (Schrank)
Baillon, Hist. Plant. 8:231. 1882. =Acanthospermum
Schrank, Pl. Rar. Hort. Acad. Monac. 53. 1820.
Melampodium achillaeoides (Less.) Hemsley, Biol. Centr.
Am. Bot. 2:145. 1881. —Villanova achillaeoides (Less.)
Less. Syn. Gen. Comp. 256. 1832. TYPE: MEXICO: “los Llanos
de Perote," Sep 1828, C. J. W. Schiede 354 (Holotype,
HAL!; photograph of holotype, os! TEX !).
Melampodium anomalum M. E. Jones, Contr. West. Bot.
18:72. 1933. TYPE: MEXICO: Jalisco, near Guadalajara, La
Barranca, 17 Nov 1930, M. E. Jones 27727 (Lectotype,
POM; isotype, MO! NY!; photograph and fragment of lecto-
type, US!). =Tragoceras schiedeanum Less. Linnaea 9:269.
1834. fide Blake (1945) and Torres (1963).
Melampodium australe Loefl. Iter Hisp. 268. 1758. TYPE:
VENEZUELA: vicinity of Barcelona, 15 Feb 1755, P. Loefling
"151" (Holotype, LINN). —Acanthospermum australe
(Loefl.) Kuntze, Rev. Gen. Pl. 1:303. t. 22, h-m. 1891.
Melampodium baranguillae Spreng. Syst. 3:619. 1826.
TYPE: COLOMBIA: “Ad fl. Magdalenae," date unknown, C.
G. Bertero s.n. (Holotype, P-co). Blake (1930) saw the
type of M. baranguillae in the Schultz-Bipontinus Her-
barium and found “it to be identical with Sclerocarpus co-
lumbianus Rusby & Blake."
Melampodium bonairense Boldingh, Fl. Neder. West-Ind.
Bil 393. 1913. Fl. Dutch West Ind. Is. 2:107. t. 9. 1914.
TYPE: DUTCH WEST INDIAN ISLANDS: Island Curacao, Mal-
pais, 28-30 Oct 1909, I. Boldingh M.14 (Holotype, U; para-
type [Boldingh 7401], P!; fragment of P paratype, Us!;
photograph of K paratype [Boldingh 7401], us!). —Balti-
mora sp.
204 Rhodora [Vol. 74
Melampodium camphoratum (L.f.) Baker in Martius, Fl.
Bras. 6(3) :161. 1884. —Unzia camphorata L.f. Suppl. 368.
1781. TYPE: SURINAM: “locis arenosis," date unknown, C.
G. Dalberg s.n. (HOLOTYPE, LINN; photograph of holotype,
os!).
Melampodium digynum (Steetz in Seem.) Jackson, Index
Kew. 2:188. 1895. Unxia digyna Steetz in Seem. Bot. Voy.
Herald. t. 30. 1853. text 154. 1854. TYPE: PANAMA: “about
Panama [City]," Nov 1846, B. C. Seemann 46 (HOLOTYPE,
BM; photograph and fragment of K isotype, us!; photo-
graph of K isotype, F! us [2]!). =Unxia camphorata L.f.
Suppl. 368. 1781.
Melampodium? dombeyanum DC. Prodr. 5:520. 1836.
TYPE: PERU: locality and date unknown, J. Dombey s.n.
(Holotype, G-DC; isotype, P!; photograph of holotype, IDC
800. 923: III. 6!; photograph of P isotype, OS! TEX!). S.
F. Blake (1930) saw the type in the Prodromus Herbarium
and notes that it “has shown its identity with Sigesbeckia
flosculosa” L’Her. Stirp. Nov. 37. t. 19. 1785-86.
Melampodium durandi (A. Gray) M. E. Jones, Contrib.
West. Bot. 15:156. 1929. Hemizonia durandi A. Gray, Proc.
Amer. Acad. Arts & Sci. 6:549. 1865. TYPE: CALIFORNIA:
Nevada Co., “.. . in the vicinity of Nevada [City]" [Du-
rand, 1855], ca. 3000 ft., 1851, H. Pratten s.n. (Holotype,
PH?). —Madia minima (A. Gray) Keck, Madroño 10:22.
1949.
Melampodium geminatum Brandg. Zoe 5:223. 1905. TYPE:
MEXICO: Sinaloa, Cofradia [ca 30 mi E of Culiacán near
Durango border], 24 Oct 1904, T. S. Brandegee s.n. (Lecto-
type chosen, UC!). = Baltimora sp.
Melampodium? hidalgoa DC. Prodr. 5:521. 1836. nom.
superfl. of Melampodium ternatum (Llave & Lex.) DC.
—Hidalgoa ternata Llave & Lex. Nov. Veg. Descr. 1:15.
1824.
1972] Melampodium — Stuessy 205
Melampodium hirsutum (Rich.) Jackson, Index Kew. 2:
188. 1895. Unwia hirsuta Rich. Act. Soc. Hist. Nat. Par.
1:105. 1792. TYPE: FRENCH GUIANA: locality unknown,
1792, J. B. Leblond 346 (Holotype, P; isotype, G!; photo-
graph of G isotype, OS! TEX!). =Unxia camphorata L.f.
Suppl. 368. 1781.
Melampodium humile Sw. Prodr. Veg. Ind. Occ. 114.
1788. TYPE: JAMAICA: “Domingo,” 1783-1787, O. Swartz
s.n. (Holotype, S; photograph of B isotype, TEX!). —Acan-
thospermum humile (Sw.) DC. Prodr. 5:222. 1836.
Melampodium L. sect. Lecocarpus (Dcne.) Baillon, Hist.
Plant. 8:231. 1882. —Lecocarpus Dene. Bot. Voy. Venus,
t. 14. 1846.
Melampodium minimum (A. Gray) M. E. Jones, Con-
trib. West. Bot. 15:156. 1929. Hemizonia minima A. Gray,
Proc. Amer. Acad. Arts & Sci. 6:549. 1865. TYPE: IDAHO:
"Dry soil, near Soda Springs," 8600 ft., 1860-1862, W. H.
Brewer s.n. (Holotype, GH?). —Madia minima (A. Gray)
Keck, Madrofio 10:22. 1949.
Melampodium minutiflorum M. E. Jones, Contrib. West.
Bot. 18:72. 1933. TYPE: MEXICO: Guadalajara, La Bar-
ranca, 17 Nov 1930, M. E. Jones 27738 (Lectotype, POM;
isotype, MO! US!; photograph and fragment of lectotype,
US!). —Galeana pratensis (H.B.K.) Rydberg, Fl. N. Amer.
94(1) :42. 1914.
Melampodium paludicola Taub. Engl. Bot. Jahrb. 21:455.
1896. TYPE: BRAZIL: Goias, “in paludibus ad flumen Parana-
hyba [Paranaiba]," Feb 1893, E. Ule 2978 [—397] (Holo-
type, P!; isotype, F!; fragment of holotype, US!; photo-
graph of holotype, F! os! TEX! Us!). Definitely not a Mel-
ampodium; “the description of which suggests a Sclerocar-
pus" (Robinson, 1901).
Melampodium? ruderale Sw. Fl. Ind. Occ. 3:1372. 1806.
TYPE: JAMAICA: “in ruderatis Jamaicae australioris," date
206 Rhodora [Vol. 74
unknown, O. Swartz s.n. (Holotype, 8?). — Eleutheran-
thera ruderalis (Sw.) Sch. Bip. Bot. Zeit. 24:259. 1866.
Melampodium suffruticosum Baker in Martius, Fl. Bras.
6(3):162. 1884. TYPE: VENEZUELA: “prope Esmeralda in
ditione fluminis Orinoco," Dec 1853, R. Spruce 3225 (Holo-
type, K!; isotypes, G! NY! P!; photograph of holotype, os!
TEX! US!). =Unxia suffruticosa (Baker) Stuessy, Brit-
tonia 21:319. 1970.
Melampodium? ternatum (Llave & Lex.) DC. Prodr. 5:
521. 1836. —Hidalgoa ternata Llave & Lex. Nov. Veg.
Descr. 1:15. 1824. TYPE: MEXICO: Veracruz, “. .. in mar-
gine densissimarum silvarum, prope ultimam cataractam
fluminis Blanco, non longe à S. Jose del Corral [near Ori-
zaba]," Apr [year?], P. La Llave s.n. (Holotype, MA?).
Melampodium L. subg. Unxia (L.f.) Baker in Martius,
Fl. Bras. 6(3) :162. 1884. —Unzia L.f. Suppl. 56. 1781.
ACKNOWLEDGEMENTS
This paper represents a modification of a dissertation
submitted to the Graduate School of the University of
Texas at Austin in partial fulfillment of the requirements
for the degree of Doctor of Philosophy. Sincerest thanks
go to Dr. B. L. Turner for the initial suggestion of the
problem and for his guidance throughout the development
of the study. Appreciation is also owed to Drs. Marshall
C. Johnston, Rogers MeVaugh, Lloyd H. Shinners, and Mr.
Imre J. Eifert for their help with literature and nomen-
clature; Mr. Geza Knipfer for preparation of most of the
base maps used for the species distributions; Messrs. Ar-
nold B. Birdsong, Imre J. Eifert, John B. Schoenhofen and
Drs. Robert S. Irving and Walter Renold for their help and
companionship on field trips; Dr. C. E. B. Bonner for cam-
era lucida drawings of diagnostic features of the type of
Melampodium angustifolium; Dr. Alicia Lourteig for in-
formation concerning type-material at Paris; Dr. J. F. M.
Cannon for information regarding Calendula decumbens;
1972] Melampodium — Stuessy 207
Dr. Harold Robinson for help in interpreting ovarian disc
structures; Dr. Rogers MeVaugh for special help with typi-
fication of the Sessé & Mocifio names; Miss Suzanne Moor-
head for drawing Figs. 6-9 and 10; and Drs. Rogers Mc-
Vaugh, Ronald L. Stuckey, Rolla Tryon, and B. L. Turner
for critically reading the entire manuscript. Finally, many
thanks are due my wife, Carol, who courageously typed
cards for representative specimens and spent countless
hours typing the drafts of the manuscript. Field work was
partially supported by NsF Traineeship 4128 (to W. F.
Blair), NSF grant GB-1458 (to H. J. Arnott), and a Sigma
Xi Grant-in-Aid.
Collective appreciation is expressed to the various cura-
tors and directors of herbaria from which loans of speci-
mens were made as indicated in the following list (abbre-
viations after Lanjouw and Stafleu, 1964) : A, ARIZ, B, C,
DS, ENCB, F, FSU, G, GH, HAL, K, LL, MA, MICH, MO, MSC, NY,
P, POM, RSA, S, SMU, TEX, UC, UPS, US, W, WIS.
LITERATURE CITED
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collected by Thaddeus Haenke and described by Karel Borewog
Presl. J. Bot. (Lond.) 72: 223-226.
BAILLON, H. E. 1882. Histoire des Plantes, vol. 8, p. 231. Paris.
BAKER, J. G. 1884. Melampodiwm, p. 159-162. In C. F. P. von
Martius, Flora Brasiliensis, vol. 6(3). Munich.
BENTHAM, G., and J. D. HOOKER. 1873. Genera Plantarum, vol. 2,
p. 348-349. Reeve & Co., London.
BLAKE, S. F. 1924. New American Asteraceae. Contrib. U. S. Nat.
Herb. 22: 587-661.
1926. Five new American Melampodinae. J. Wash.
Acad. Sci. 16: 418-422.
1930. Notes on certain type specimens of American
Asteraceae in European herbaria. Contrib. U. S. Nat. Herb.
26: 227-268.
1945. Asteraceae described from Mexico and the
southwestern United States by M. E. Jones, 1908-1935. Contrib.
U. S. Nat. Herb. 29: 117-137.
CANDOLLE, A. P. DE. 1836. Prodromus, vol. 5, p. 517-521. Paris.
Cassini, A. H. G. 1829. Melampodium, Zarabellia, and Alcina, p.
237-244. In G. Cuvier (ed.), Dict. Sci. Natur., ed. 2, vol. 59.
Paris.
208 Rhodora [Vol. 74
COCKERELL, T. D. A. 1905. The name Melampodium. Torreya 5: 70.
COLEMAN, J. R. 1970. Additional chromosome numbers in Brazilian
Compositae. Rhodora 72: 94-99.
DURAND, E. 1855. Plantae Prattenianae. J. Acad. Nat. Sci. Philad.
n.s. 3: 79-104.
GEISER, S. W. 1948. Naturalists of the Frontier, ed. 2, p. 30-54.
Southern Methodist Univ, Press, Dallas.
GENTRY, H. S. 1942. Rio Mayo Plants. Carnegie Inst. Wash. Publ.
527: 262.
GRAY, A. 1852. Plantae Wrightianae Texano-neo-Mexicanae. Smith-
son. Contrib. Knowl. 3(5): 103.
1855. Plantae Novae Thurberianae. Mem. Amer. Acad.
ser. 2, 5: 321.
1859. Melampodium, p. 85. In J. Torrey, Botany of the
Boundary, United States and Mexican Boundary Survey, vol.
2(1). Washington, D. C.
1884. Synoptical Flora of North America, vol. 1(2),
p. 238-239. Ivison, Blakeman, Taylor, and Co., N. Y.
HEPPER, F. N. 1968. A standard citation of microfiches. Taxon
17: 604.
HITCHCOCK, A. S. 1909. Catalogue of the grasses of Cuba. Contrib.
U. S. Nat. Herb. 12: 183-256. (cf. p. 210)
HOFFMANN, O. 1890. Melampodium, p. 218. In A. Engler and K.
Prantl, Die natürlichen Pflanzenfamilien, vol. 4(5). Leipzig.
HUMBOLDT, F. H. A., A. J. BONPLAND, and C. S. KUNTH. 1820. Nova
Genera et Species Plantarum, vol. 4, p. 272-274. Paris.
KAPADIA, Z. J. 1964. Varietas and subspecies: a suggestion to-
wards greater uniformity. Taxon 12: 257-259.
KEARNEY, T. H., and R. H. PEEBLES. 1942. Flowering Plants and
Ferns of Arizona. U. S. D. A. Misc. Publ. No. 423: 941.
, and . 1964. Arizona Flora, ed. 2,
p. 810. Univ. Calif. Press, Berkeley.
LANGMAN, I. K. 1964. A Selected Guide to the Literature on the
Flowering Plants of Mexico, p. 756. Univ. Penn. Press, Phila-
delphia.
LANJOUW, J., et al. (eds.). 1966. International Code of Botanical
Nomenclature. Regnum Veg. 46: 37-38.
, and F. A. STAFLEU. 1964. The herbaria of the world.
Index Herbariorum, part 1, ed. 5. Regnum Veg. 31: 205-228.
LINNAEUS, C. 1737. Critica Botanica. Leiden.
1738. Hortus Cliffortianus, p. 425. Amsterdam.
1753. Species Plantarum, vol. 2, p. 921. Stockholm.
McVauaGH, R. 1956. Edward Palmer: Plant Explorer of the
American West. Univ. of Oklahoma Press, Norman.
MERRILL, E. D. 1923. An Enumeration of Philippine Flowering
1972] Melampodium — Stuessy 209
Plants, pp. 1-128. Philip. Dept. Agr. and Nat. Sci., Publ. No. 18,
vol. 1(1). Manila. (cf. p. 49)
1954. The botany of Cook's voyages. Chron. Bot.
14: 161-384.
RICKETT, H. W. 1947. The royal botanical expedition to New Spain.
Chron. Bot. 11: 1-86.
ROBINSON, B. L. 1901. Synopsis of the genus Melampodium. Proc.
Amer. Acad. Arts & Sci. 36: 455-466.
SAFFORD, W. E. 1905. The useful plants of the island of Guam.
Contrib. U. S. Nat. Herb. 9: 1-416. (cf. p. 25)
SHREVE, F., and I. L. WiagGINS. 1964. Vegetation and Flora of the
Sonoran Desert, p. 1522. Stanford Univ. Press, Calif.
SPRAGUE, T. A. 1926. Sessé & Mocifio’s Plantae Novae Hispaniae
and Flora Mexicana. Kew Bull. Misc. Inform. 9: 417-425.
STUESSY, T. F. 1969a. A new variety and new combination in
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1970a. The genus Acanthospermum (Compositae-
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1970b. Six new species of Melampodium (Com-
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1971a. Chromosome numbers and phylogeny in
Melampodium (Compositae). Amer. J. Bot. 58: 732-736.
1971b. Chromosome studies in Melampodium
(Compositae-Heliantheae). Madroño 20: 365-372.
1971c. Systematic relationships in the white-rayed
species of Melampodium (Compositae). Brittonia 23: 177-190.
TIDESTROM, I., and T. KrrTELL. 1941. A Flora of Arizona and New
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TORRES, A. M. 1963. Revision of Tragoceras (Compositae). Brit-
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263-269.
DEPARTMENT OF BOTANY
THE OHIO STATE UNIVERSITY
COLUMBUS, OHIO 43210
210
TAP ON E
SERERERESEEEEREEESEEEEE
Rhodora
LIST OF TAXA OF MELAMPODIUM
americanum L.
diffusum Cass.
pilosum Stuessy
longipes (A. Gray) Robins.
linearilobum DC.
leucanthum Torr. & A. Gray
cinereum DC.
M. cinereum DC. var. cinereum
M. cinereum DC. var. hirtellum Stuessy
M. cinereum DC. var. ramosissimum (DC.)
A. Gray
. argophyllum (A. Gray ex Robins.) Blake
. sericeum Lag.
. pringlei Robins.
. strigosum Stuessy
. longicorne A. Gray
nayaritense Stuessy
. cupulatum A. Gray
appendiculatum Robins.
. sinuatum Brandg.
rosei Robins.
. tenellum Hook. & Arn.
glabribracteatum Stuessy
. longipilum Robins.
longifolium Cerv. ex Cav.
mimulifolium Robins.
. gracile Less.
. microcephalum Less.
. paniculatum Gardn.
. divaricatum (Rich. in Pers.) DC.
. costaricense Stuessy
. dicoelocarpum Robins.
[Vol. 74
1972]
29. M. tepicense Robins.
30. M. sinaloense Stuessy
31. M. bibracteatum S. Wats.
32. M. repens Sessé & Moc.
33. M. montanum Benth.
Melampodium — Stuessy
211
33a. M. montanum Benth. var. montanum
33b. M. montanum Benth. var. viridulum Stuessy
34. M. aureum Brandg.
35. M. perfoliatum (Cav.) H.B.K.
36. M. glabrum S. Wats.
37. M. nutans Stuessy
D. Doubtful species
E. Excluded taxon
INDEX TO EXSICCATAE
Numbers in parentheses represent the taxa recognized in this study
(see preceding list).
Abbott, R. Q. 162 (33b); 304
(24); 439 (23); 445 (35).
Abloon, Bro. 137 (7b).
Abrams, L. R. 12811 (14).
Adole, Fr. L. 29 (7b).
Aguilar H., M. 25 (23); 322
(25); 1404 (9); 1495 (5);
1567 (9); 1942 (35).
Alain, Bro. 1819 (35).
Alamán, L. s.n. (5).
Alexander, E. L. & E. Hernández
X. 2229 [XA218] (31).
Allen, P. H. 241 (26); 585 (25).
Arséne, Bro. G. 76 (21); 294
(31); 1199 (21); 1896 (33b);
3523 (35); 5219 (34); 6575
(33b); s.n. (35), (36).
Arzola, O. 178 (7a).
Asplund, C. 517 (35).
Bacigalupi, R. C. 607 (6).
Baize, R. M. 5 (6).
Baker, C. F. 2482 (26).
Baker & Wilson 516 (35).
Balls, E. K. & W. B. Gourlay
B5372 (32).
Barkley, F. A. 14142, 14163 (2);
14575C (7b).
Barkley, F. A. & G. Gutierrez V.
1777 (26).
Barkley, F. A., C. M. Rowell, Jr.
& J. B. Paxson 735 (9).
Barkley, F. A., G. L. Webster &
J. B. Paxson 17M724 (2);
17M732 (24).
Barkley, F. A., G. L. Webster &
C. M. Rowell, Jr. 7488 (32).
Barneby, R. C. 5135 (12).
Beaman, J. H. 3043 (33b); 3977
(31); 4852 (32):
Benson, L. 10598 (12).
Berlandier, J. L. 2017 [607]
(Tc); 2153 [733] (23); 2243
[8238]. (Ta) "sa. :(24):
Bernoulli, C. G. 709 (5).
Bertero, C. G. s.n. (26) ; s.n. (E).
212
Blake, S. F. 7632 (26).
Blumer, J. C. 1665 (11); 2128
(12).
Boldingh, I. M.14 (E).
Bonner, Mrs. J. 55 (Tb).
Bonpland, A. J. 3968 (1).
Botteri, M. 809 (24).
Bourgeau, E. 841 (35);
1629 (24).
Brandegee, T. S. 301 (26); 302
(16); sn. (5), (16), (17).
Braunton, E. 625 (35).
Bray s.n. (Ta).
Breedlove, D. E. 6085, 7861 (25);
8906, 12522 (33b).
Brenes, A. M. 17471, 20572 (27).
Brewer, W. H. s.n. (E).
Britton, N. L. & J. F. Cowell
1561 (26).
Bruni, T. 15 (7b).
Burr, R. D. 135 (6).
Busseuil, F. L. s.n. (2).
1628,
Calderón, S. 459, 779 (25); 979
(5); 1019 (9).
Cameron, E. R. 248 in
(7c).
Canby, W. M. 133 (7a).
Canicosa, E. 1103 (2).
Carleton, M. A. 331 (6).
Carlson, M. C. 79 (26);
873, 2254 (25).
Carter, A. 4880, 4964, 5055 (14).
Carter, A., A. M. Alexander &
L. Kellogg 2216 (16).
Carter, A. & R. Moran
(14).
Cervantes, J., J. A. Molina &
F. A. Barkley 19Bo061 (26).
Cervantes, V. s.n. (21).
Chambers, K. L. 965 (14).
Chamisso, A. s.n. (2).
Chase, V. H. 7122 (26); 7533
(23).
Cisneros, F. 33 (7a).
part
106,
5314
Rhodora
[Vol. 74
Clark, O. M. 6635 (Ta); 12377
(12).
Clausen, P. s.n. (25).
Clausen, R. T. s.n. (24).
Clemens, M. S. s.n. (2).
Clokey, I. W. 3946 (6).
Conzatti, C. 1706 (1); 1865
(34); 2291 (35); 4169 (33a);
5169 (19).
Conzatti, C. & V. González 550
(24); 998 (33a).
Correll, D. S. 15204 (Ta).
Cory, V. L. 50422 (6).
Cronquist, A. 9585 (31); 9613
(24); 9745 (35); 10003 (12).
Crutchfield, J. & M. C. Johnston
5527 (Te); 5643a (35); 5664
(9); 6120b (26); 6141b (Te).
Culwell & Timmons 3091 (6).
Curran, H. M. & M. L. Merritt
15837 (2).
Dalberg, C. G. s.n. (E).
Daniel, Bro. 2290 (26).
Darrow, R. A. & H. S. Haskell
2012, 2256 (12).
Deam, C. C. 247 (26).
Degener, O. 26220 (35).
Degener, O. & I. 26504, 26559
(25).
Demaree, D. 7594, 12210, 13344
(6).
Dohnke, W. C., Jr. 3 (7a).
Dombey, J. s.n. (E).
Domínguez M., M. 63 (7a).
Domínguez M., M. & W. McCart
8217 (Ta).
Drouet, F., D. Richards & W. A.
Lockhart 3609, 3624 (15).
Drushel, J. A. 10441 (7a).
Duges, A. s.n. (11); 44 (9); 461
(35).
Dunlap, V. C. 557 (27).
Earle, F. S, & E. S. 295 (6).
Echeverría, J. A. 628 (27).
1972]
Eggleston, W. W. 10729 (11).
Ekman, E. L. 16932 (35).
Erlanson, C. O. 500 (26).
Ervendberg, L. C. 92 (23).
Escalante, L. 21 (7a).
Eyerdam, W. J. & A. A. Beetle
8687 (17).
d);
2204
2651
Feddema, C. 279 (29); 832
1209 (18); 1219 (14);
(28); 2238 (37); 2577,
(29); 2700 (37).
Ferris, R. S. 5020 (17);
(35).
Ferris, R. S. & Y. Mexia 5078
(17).
Fisher, G. L. 35321 (5).
Fleetwood, R. J. 3840 (7a).
Flyr, D. 112 (14); 138 (17);
489 (6).
Foxworthy, F. W. 167 (2).
Frye, T. C. & E. M. 2415 (7b).
DE xn): En.
5890
Furness,
(11).
García, A., Jr. & R. 47 (7a).
García, C. 51 (25).
Gardner, G. 287, 3844 (25).
Garza, C. & L. de la 46 (7a).
Gaumer, G. F. 563 (26); 2349
(23).
Gaumer, G. F. & sons 789 (23).
Gentle, P. H. 2325 (26).
Gentry, H. S. 290 (14); 550
[550M] (9); 790 [790M], 1492,
1570 (15); 1752 (35); 2353
(15); 2553 (9); 5039 (5);
5750 (14); 6153 (5); 6350
(35); 6507-A (30); 7005 (14).
Gentry, H. S. & J. Arguelles
18152 (35).
Gentry, H. S., D. S. Correll &
J. Arguelles 17970 (35).
Glassman, S. F. 2003 (25).
Melampodium — Stuessy
213
Glaziou, A. 12794 (25).
Glocker, E. F. von 19 (26).
Godfrey, R. K. 6664 (27) ; 66064,
66184 (35); 67156 (27).
Goldman, E. A. 110 (11); 914
(33b).
Goldsmith, P. 132 (4); 133 (13).
González O., J. 6125 (18).
Goodding, L. N. 13 (11); 248-59,
267-62, 580-49, 699-58 (12);
814 (11); 2399a (6).
Goodman, G. J. & C. L. Hitchcock
1130 (6).
` Gould, F. W. 2883 (12).
Gould, F. W. & H. S. Haskell
3268 (12).
Graham, G. J. 5 (26).
Graham, J. & M. C. Johnston
4378 (7c); 4777 (23).
Grant, G. B. 763 (35).
Greenman, J. M. 279 (1).
Gregory, D. P. & G. Eiten 205
(24).
Guthrie, N. 43 (6).
Haenke, T. s.n. (1).
Hallberg, B. 900 (33a).
Hammerly, B. J. 182 (14).
Hancock, W. 25 (2).
Harrison, G. J. 8181 (12).
Harrison, G. J. & H. J. Fulton
8153 (12).
Harrison, G. J. & T. H. Kearney
5702 (12).
Harshberger, J. W. 176 (21).
Hartman, C. V. 71 (15).
Hartweg, C. T. 475 (33a).
Heard, N. T. & F. A. Barkley
14542B (7b).
Hedrick, D. C. 156 (7b).
Heller, A. A. 1632 (6).
Henderson, N. C. 63-88, 63-98
(6).
Hermann, H. A. van 148 (35).
Heyde, E. T. & E. Lux 4213 (26).
214
Hinckley, L. C. 620, 881, 3293, s.n.
(11).
Hinton, G. B. 1520 (26); 1521
(35); 1661 (24); 1838 (35);
2008 (24); 2017 (33b); 2480
(35); 2513 (5); 4628, 4826
(32); 6207 (24); 6270 (5);
6324 (9); 6334 (28); 60338
(23); 6420 (5); 6436 (3);
9113 (5); 9134 (1); 9205
(24); 9293 (35); 9714 (28);
10719 (24); 11428 (3); 12276
(29); 12871 (1); 13214 (5);
14614 (28); 15587 (1); 16563
(8).
Hitchcock, C. L., R. V. Rethke &
R. van Raadshooven 4214, 4244
(6).
Hoffmann, O. 832 (24).
Holman, R. S. & A. F. Dittman
791 (23).
Holzner, F. X. 1962 (11).
Horr, W. H. 3444 (6).
Horr, W. H. & L. H. Franklin
E278 (6).
Houstoun, W. s.n. (1).
Howell, J. T. 10303 (18).
Humboldt, F. H. A. von & A. J.
Bonpland 142 (26); s.n. (9).
Iltis, H. & F., & R. Koeppen 35
(7c).
Innes, R. R. & B. H. Warnock
589 (6).
Irving, R. 825 (6).
Irwin, H. S., R. Souza & R. Reis
dos Santos 11303 (25).
James, J. F. s.n. (35).
Jespersen, B. & H. 2678 (6).
Jiménez M., A. 1073 (26); 1074
(35).
Johnson, J. C. & F. A. Barkley
16250M (8).
Johnston, I. M. 7959 (6).
Rhodora
[Vol. 74
Johnston, M. C., B. C. Tharp &
B. L. Turner 3472 (7a).
Jones, F. B. 719, 1359 (7a).
Jones, M. E. 117 (6); 273 (26);
274, 369 (1); 22681 (11);
23398, 23402 (35); 25045
(12); 25916 (Tb); 27727,
27738 (E); 27739 (14); 27773
(35); s.n. (11); s.n. (12).
Kammerer, A. L. 48 (6).
Kay, V. W. & J. E. Higgins 202
(23).
Kearney, T. H. 5587A (11).
Kearney, T. H. & R. H. Peebles
10052 (12); 11133 (6).
Kellerman, W. A. 4501
4958 (26).
Kenoyer, L. A. 5 (Tb); 427 (23) ;
2421 (36).
Kenoyer, L. A. & Crum 2594
(1b); 4093 (9).
Killip, E. P. & A. C. Smith 14169
(26).
King, R. M. 1833 (24);
2718 (1); 2743 (26);
(25) ;
2682,
2796,
2801, 2843 (33b);
2980, 2987 (1);
(23); 3105 (1);
3948 (35); 3249
(1); 3322 (26);
3358 (1); 3359
(1); 3410 (35);
3417 (25); 3425
(26); 3454 (5);
3492 (33a); 3529
(5); 8560, 3600
(24); 3617 (34);
3634 (26); 3636,
3662 (4); 8678
(1); 3703, 3704,
(18); 3707, 3709,
3711 (26); 3712,
(17); 3833 (26);
2917 (26);
3042, 3096
3242 (25);
(26); 3260
3329 (25);
(26); 3364
3416 (9);
(24); 3434
3461 (20);
(34); 3539
(35); 3607
3626 (9);
3646 (23);
(26); 3699
3705, 3706
3710 (17);
3715, 3716
3887 (23);
1972]
4138, 4164 (26); 4165 (23);
4168, 4178 (1); 4239 (26);
5090 (27); 5226 (26); 5348
(21); 5350, 5407 (35).
King, R. M. & T. R. Soderstrom
4584 (9); 4646 (28); 4707
(35); 4804 (24); 4992 (1).
Koeppen, R. & F. Iltis 618 (1).
Kuntze, O. 23527 (24); 27465
(9).
Lacás, M. M. 58 (7b).
Lamb, F H. 361a, s.n. (17).
Langlassé, E. 482 (28).
Laughlin, R. M. 1050 (33b).
Lay, G. T. & A. Collie s.n. (18).
Leavenworth, W. C. 157a (26);
431 (37).
Leavenworth, W. C. & H. Hoog-
straal 1480, 1588 (5); 1589
t
Leblond, J. B. 346 (E).
Lehmann, F. C. 69, 1434 (26).
Lemmon, J. G. 331 (12); 2777
(11); 2795 (12).
Léon, Bro. M. Victorin & Bro.
Alain 17344 (26).
LeSueur, H. 341 (11); 1456 (6) ;
1466, 1472 (15).
Liebmann, F. M. 232 (33a).
Lindheimer, F. J. 949 (6).
Llave, P. La s.n. (E).
Loefling, P. “151” (E).
Löfgren, A. & G. Edwall 16695
(25).
Loomis, H. F. & C. J. King 2890
(12).
Lundell, C. L. 6107 (27); 6759
(23).
Lundell, C. L. & A. A. 7879 (26) ;
7972 (23); 9866 (7a).
Lyonnet, E. 569 (35).
MacDaniels, L. H. 41 (35); 53
(21); 723 (35); 747 (11).
Melampodium — Stuessy
215
Macedo, A. 1408 (25).
Maguire, B., B. L. Richards & T.
Moeller 11849 (6).
Manning, W. E. & M. S. 531212
(9).
Marsh, E. G., Jr. 855 (6); 1176
(7b).
Martinez, P. & A. Garcia 13
(7a).
Matthews, R. W. & J. R. 480
(15).
Matuda, E. 4444 [4504] (5);
6002 (33b); 26621 (26);
31141, 31314 (5).
Maxon, W. R. 7694, 7696 (26).
May, J. 13 (7b).
McClatchie, A. J. s.n. (35).
McGregor, C. 20181 (2).
McVaugh, R. 7694 (7b); 7986
(6); 12822 (9); 13033 (33b);
13576, 14107 (35): 16112
(29); 18070 (37); 18906 (29).
McVaugh, R. & W. N. Koelz 542
(29); 1054 (37).
Mearns, E. A. 1997 (15).
Mears, J. A. 1383, 1413 (6).
Melchert, T. E. 169-C, 180 (6).
Melchert, T. E. & P. Sorensen
6085 (3).
Mendez, J. [^Menzies"] s.n. (9).
Merrill, E. D. 10641 (2).
Metcalfe, O. B. 66, 1271 (6).
Mexia, Y. 240, 349 (5).
Meyer, F. G. & D. J. Rogers 2828,
2911 (34).
Millspaugh, C. F. s.n. (24).
Mockford, J. P. & C. M. Rowell,
Jr. 2769 (2).
Molina, J. A. & F. A. Barkley
19Bo150 (26).
Molina R., A. 1082 (26); 5899,
13346 (5).
Moore, H. E., Jr. 2283 (20).
Moore, J. A. & J. A. Steyermark
3115 (6).
216
Morales, R., H. 1221 (26); 1300
(9).
Moreno, E. M. MS-198 (15).
Muenscher, W. C. 12467 (25).
Muller, C. H. 3040 (8); 3434
(9); 3659 (15).
Munz, P. A. 13303 (7a).
Nagel, O. 8030 (24).
Née, L. s.n. (2).
Nelson, E. W. 655 (33a); 715
(35); 740 (22); 870 (24);
947, 2339, 2740 (1); 2949 (5);
6447 (6); 6939 (1).
Nicolas, Bro. 5447 (21); €150
(35); s.n. (21).
Niles, W. E. 572 (6).
Oersted, A. S. 9010 (27).
Orcutt, C. R. 1395 (8); 3548
(35); 3605 (9); 3780 (32);
3887 (9); 3892 (35); 4365
(23); 4386 (20); 4516 (1);
6601 (29).
Orozco, J. M. 260 (25);
(27); 418 (25).
Ortega, J. G. 4008, 4446, 6125,
7004 (17).
294
Padilla, S. A. 1, 242, 586 (5).
Painter, J. T., W. D. Lucas &
F. A. Barkley 14297 (7b).
Palmer, E. 3 (2); 20 (14); 186
(26); 245 (15); 260 (9); 281
(2); 286 (35); 319 (23); 391
(4); 459 (1); 486 (11); 535
(1); 556 (7b); 726 in part
(14), (15); 926 (9); 1163
(35); 1172 (1); 1457 (17);
1609 in part (5), (15) ; 1757a
(17); 1814 (29): 2068 (8).
Parish, S. B. & W. F. 1171 (35).
Parker, K. F. 7707 (12).
Parry, C. C. & E. Palmer 444
(21); 444% (11).
Rhodora
[Vol. 74
Paynet, A. M. & H. D. Hulan
1213 (23).
Peebles, R. H. & G. J. Harrison
4680 (12).
Peebles, R. H., G. J. Harrison &
T. H. Kearney 4614 (11).
Pennell, F. W. 1335 (27); sn.
(18).
Pereira, E. & G. Pabst 8505
[34972] (26).
Pipes, D. 81 (23).
Pittier, H. 57, 60, 429, 6963 (25).
Pohl, J. E. 1368 [1276] (25).
Porter, C. L. 4262 (6).
Powell, A. M. & J. Edmondson
551 (21); 579 (9); 646 (23);
660 (9); 756 (26); 758 (2);
816 (35); 908 (17).
Pratten, H. s.n. (E).
Pringle, C. G. 10 (11); 55 (12);
297, 754 (11); 1019 (6); 1045
(11); 1767 (35); 2819 (26);
2821 (36); 3230 (31); 3639
(20); 3818 (33b); 3863 (36);
3923 (26); 4322 (24); 4537
(20); 4598 (4); 4666 (33a);
5257 (32); 5309 (9); 5722
(10); 6455 (21); 6491 (11);
6728 (9); 7321 (24); 7327
(32); 8466 (23); 8609 in part
(9), (11); 8610 (32); 9008
(7b); 9162 (5); 9177 (23);
9301 (31); 9325 (21); 9331
(11); 9361 (33b); 9937 (23);
10065 (5); 11548 (11); 13068
(31); 13069 (24).
Purpus, C. A. 416 (16); 1003,
1003a (8); 1809 (32); 2813,
2813’ (21); 2849 (35); 3822,
3822a (34); 4730 (8); 5619
(20); s.n. (8).
Ramírez, J. 44 (Ta).
Reese, J. A. 94 (12).
Reko, B. P. 3812 (5).
1972]
Reséndez, O. I. 52 (7b).
Richard, L. C. M. s.n. (26).
Ricksecker, A. E. *S" (26).
Riddell, J. L. s.n. (6).
Ríos, A. & H. Cavazos 231, 264
(Ta).
Ripley, H. D. & R. C. Barneby
13708 (24).
Robinson, C. B., Jr. 11835 (2).
Rock, H. F. L. M-442 (9).
Rodríguez, J. J. 72 (7b).
Rodriguez, J. V. 1662 (25); 3341
(26); 3735 (25).
Rojas 86 (9).
Rose, J. N. 1568 (17); 3183 (4);
3271 (17); 3476 (18); 3617
(9); s.n. (17).
Rose, J. N. & W. R. Fitch 17045
(6).
Rose, J. N. & R. Hay 6244 (32);
6275 (9).
Rose, J. N., J. H. Painter & J. S.
Rose 9308 (23); 10137 (20).
Rose, J. N., P. C. Standley &
P. G. Russell 14307 (1).
Rowlee, W. W. & H. E. 179 (5).
Russell, N. H. 11511 (6).
Russell, P. G. & M. J. Souviron
2 (15).
Rzedowski, J. 9320 (33a) ; 17514,
17921, 21155 (37); 20748 (1);
21322 (33b); 26618 (18).
Safford, W. E. 1264 (8).
Sánchez, A. 105 (7a).
Schaffner, J. G. 270 in part (9),
(35); 271 (9) ; s.n. (14), (32).
Schiede, C. J. W. 217 (24); 354
(E).
Schiede, C. J. W. & F. Deppe
1254 (23).
Schott, A. 72, s.n. (15).
Schroeder, A. H. 120 (8).
Seaton, H. E. 54 (35); 461 (24).
Seemann, B. C. 46 (E).
Melampodium — Stuessy
217
Sessé, M. “3977” (D); “3978”
(24)3.5 489817 1982) 5: 39831,
s8m(26)5 s.n: Gb):
Sessé, M. & J. M. Mocino s.n.
(9).
Shank, P. J. & A. Molina R.
4236 (27).
Sharp, A. J. 44116 (32); 44208
(21); 44961 (9).
Shinners, L. H. 7183 (6); 7395
(7a); 12190 (6); 17659, 17708
(7a); 26390, 31286, 31559 (6).
Shreve, F. 4965, 6003 (12) ; 6758
(15); 7685, 7709 (12); 7821
(17); 7918 (6); 9278 (35).
Sinclair, A. s.n. (26).
Skutch, A. F. 1036 (33b); 1550
(25); 1592 (9); 1709 (25);
2041 (26); 3968 (27).
Small, J. K. & E. T. Wherry
12134 (6).
Smith, A. 2922 (27).
Smith, C. L. 351 (35);
(23); 1655 (24).
Smith, H. H. 514 in part (26).
Smith, J. D. 2334 (25); 2382
(26).
Solís, F. 52 (25); 56 (27).
Solís, M. J. 124 (7a).
Spruce, R. 3225 (E).
1605
Standley, P. C. 11226 (26);
12105, 15756, 17710 (25);
19722 (26); 20260 (25);
21648, 21789 (26); 22687,
22718, 22864, 25149 (25);
26708, 27175 (5); 28678 (25);
32757 (27); 36058, 38971
(25); 89021, 41581 (27);
56290, 56522 (25); 59769 (9);
60326, 60413 (25); 60493 (5);
63265, 643808, 64595, 68866
(25); 73774, 74071, 74125,
74813, 75620 (5); 76429, 77073
(25); 77149 (26); 77449,
218
78268, 79869 (25); 82849 (9);
87933 (25).
Standley, P. C. & J. Chacón P.
6625, 6660 (25).
Standley, P. C. & E. Padilla V.
2755, 3630 (25).
Standley, P. C. & J. Valerio
48814 (27).
Stanfield, Mrs. V. E. s.n. (6).
Stanford, L. R., K. L. Rether-
ford & R. D. Northcraft 186
(8); 1077 (1); 1078 (23).
Stanford, L. R., L. A. Taylor &
S. M. Lauber 2313 (26).
Steere, W. C. 1805 (23).
Stevens, F. L. 136 (25).
Stevens, G. W. 325 (6).
Stewart, R. M. 84 (6).
Steyermark, J. A. 29041, 30509
(5); 31670 (26); 32847 (25);
43745 (26); 48234 (33b);
50392 (9); 50490 (33b);
50530 (9); 50539 (23); 50724
(5); 50773 (25); 50916 (20);
51452 (23).
Stork, H. E. 2486 (27).
Stork, H. E. & J. L. Morrison
8920 (26).
Stuessy, T. F. 147, 167, 180, 230
(6); 274 (23); 282, 318 (26);
328 (20); 342 (26); 344 (20);
345 (26); 346, 347 (20); 348,
351 (26); 356 (23); 359 (26);
361 (5); 364 (9); 365 (26);
366, 369 (2); 374 (20); 377
(9); 379 (35); 383 (24); 393
(23); 397 (4); 401 (18); 4053
(26); 404 (17); 414 (6); 425,
429 (Ta); 450 (Tc); 463, 466
(23); 471 (1); 473 (26); 494,
497, 503, 505, 511 (35); 526,
527, 528 (26); 531 (23); 582
(26) ; 533 (23) ; 534, 535 (26);
537 (23); 543 (26); 544 (23);
545 (26); 546 (23); 547, 548,
Rhodora
[Vol. 74
550 (26); 559 (33b); 562
(23) ; 566, 567, 568, 570 (33b) ;
571 (1); 574 (23); 578 (25);
580 (35); 582, 585 (25); 586
(26); 587 (25); 588 (26) ; 589
(25); 592 (26); 594 (25); 602
(1); 603 (35); 605 (26); 607
(5); 608, 609, 611 (26); 613
(5); 614, 615, 616 (26); 619
(5); 620, 621, 623, 624, 625
(26); 627 (24); 632 (1); 634,
636, 637 (20); 638 (24); 639
(20); 640 (24); 642 (9); 645
(26); 647 (9); 650 (5); 654
(35); 659 (21); 660 (9); 663
(34); 667 (20); 668 (35); 670
(32) ; 675, 676 (31) ; 677 (26);
682, 684 (34); 688 (1); 690
(9); 693 (28); 694 (23);
695 (3); 697, 698 (5); 702
(26); 703 (9); 705 (35);
708, 709 (36); 713 (23); 714
(36); 715 (28) ; 720, 721 (37);
722 (23); 723 (26); 724 (1);
725 (26); 727 (1); 728 (26);
729 (35); 730 (9); 738 (4);
740 (9); 743 (26); 745 (18);
746 (26); 748, 750 (17); 751,
752 (6); 771 (7a); 778, 787
(Te); 826 (23); 829 (26); 854
(Ta); 855, 856 (Tb); 857, 868
(Ta); 902, 912 (7b): 1016,
1038, 1054 (11); 1122, 1127
(6); 1128 (11).
Stuessy, T. F. & W. Renold 1261
(7c); 1284 (7a).
Sullivan, W. & B. L. Turner 34
(6).
Swartz, O. s.n. (E).
Taylor, M. 115 (21).
Tetters, S. 18 (6).
Tharp, B. C. & M. C. Johnston
541 [802A], 541922 (7a).
Tharp, B. C. & Miller 51-312
(6).
1972]
Thompson, B. & B. Fields 332
(2).
Thompson, B. & A. Graham 18
(6).
Thompson, E. s.n. (1).
Thornber, J. J. 7404, 8121, 9050
(12).
Thurber, G. 937 (12).
Tonduz, A. 10144 (25); 13615
(26).
Tonduz, A. & Rojas 203 (25).
Torres R., R. 50 (25).
Townsend, C. H. T. & C. M.
Barber 351 (11); 376 (15).
Tracy, S. M. & F. S. Earle 23
(6).
Triana, J. J. 1331 (25).
Tucker. J. M. 455 (5); 585 (26);
1313 (25); 2565 (9).
Tuerckheim, H. von 114 (25);
124 (26); 761 (25); 3955 (9);
8417 (25).
Turner, B. L. 4490 (7a); 4620
(11); 4790 (6); 5006 (Ta);
5632, 5673, 5735 (6).
Turner, B. L. & T. E. Melchert
4840 (6).
Turner, B. L. & A. M. Powell
1116 (11).
Turner, C. 7 (6).
Turner, R. M., R. A. Dodge &
C. T. Mason, Jr. 2068 (12).
Turner, R. M. & C. H. Lowe, Jr.
2074 (15).
Ule, E. 2978 [397] (E).
Velasco, L. V. 8946 (25).
Volkem 2564, 2774 (25); 2973
(1); 3013 (26).
Volkens, G. L. A. 1134 (35).
Walker, M. L. 93 (Tc).
Wallis, C. 4807 (6).
Warnock, B. H. 9229,
(11); 46217 (6).
Warscewicz, J. von s.n. (26).
Waterfall, U. T. 3781, 4818 (6);
13507
Melampodium — Stuessy
219
15805 (7b); 16198 (31); 16342
(9).
Waterfall, U. T. & C. S. Wallis
13702 (31); 13886 (9); 13960
(11); 14288 (23).
Webster, M. B. 140 (33b).
White, O. E. 100 (26).
White, S. S. 2642 in part (9),
(11); 2784 (15); 2788 (11);
3516, 3908, 4153 (15); 4587
(6).
Whitehouse, E. 9841 (6).
Wiegand, M. C. & G. B. Upton
4439 (6).
Wiggins, I. L. 7021, 7388, 7444,
7533 (15); 11507 (14); 11765
(6); 14377 (16).
Wiggins, I. L. & R. C. Rollins 281
(14); 433 (15).
Wilbur, R. L. & C. R. 1395 (1);
2416 (29).
Wilcox, T. E. 335, s.n. (12).
Williams, L. O. 5643, 18257 (25).
Williams, L. O. & A. Molina R.
13398 (26).
Wilson, P. 9553 (35).
Winbery, J. E. & C. M. Rowell,
Jr. 2492 (2).
Woodson, R. E., Jr., P. H. Allen
& R. J. Seibert 1370 (26).
Wooton, E. O. 117 (6) ; s.n. (7c).
Worth, C. R. & J. L. Morrison
8807 (17).
Wright, C. 311 (6); 1205 (12).
Wright, J. T. 42-54 (6).
Wright, W. G. 1213 (17).
Wynd, F. L. & C. H. Muller 132
(8); 214 (7b).
Xantus, L. J. s.n. (1).
York, C. L. & G. 54037 (6).
Young, M. S. s.n. (11).
Yuncker, T. G., R. F. Dawson &
H. R. Youse 5519 (25).
Zamora, F., J. B. Paxon & F. A.
Barkley 16M905 (11).
SPORES, CHROMOSOMES AND RELATIONS
OF THE FERN PELLAEA ATROPURPUREA
ALICE F. TRYON
The Purple Cliff-Brake, Pellaea atropurpurea, is the only
American species of the large complex of cheilanthoid ferns
that has a range extending from the tropics into northern
regions, including New England. Only triploid plants are
known, which evidently are hybrids, and they are apog-
amous with 87 chromosomes in the sporophyte and spore
mother cells. The species is a tool of some biological interest
as there appears to be no evidence of sexual reproduction,
although the specimens are highly variable throughout the
geographic range. Northern Mexico and the adjacent
southwestern United States is regarded as a center of dis-
persal of Pellaea section Pellaea in America, for eleven of
the fourteen species have distributions that converge there
or have close relatives that do. Studies of the cytology and
spores of P. atropurpurea and two other species, P. notabilis
and P. ternifolia, that occur in this central area, provide
new data for establishing relationships among these species,
and insights on the possible origin of P. atropurpurea.
MATERIAL AND METHODS
The general geographic distributions of the taxa are
outlined on the maps; localities more than 600 miles from
the main range are indicated by separate dots. Ranges are
based on data from my revision of Pellaea section Pellaea
(A. Tryon, 1957) and from recent reports of range exten-
sions. The wide cytological sampling of populations is
shown on the maps. The new cytological] records are com-
bined with earlier ones in Table 1. Meiosis was studied in
sporangia fixed in 3:1 absolute ethyl alcohol: glacial acetic
acid. Mitosis was examined in root tips fixed in the same
solution following pretreatment for three hours with
saturated aqueous solution of paradichlorobenzine.
Scanning electron microscope photographs were obtained
from spores undercoated with carbon prior to coating with
an alloy of palladium-gold while rotating and tilting the
220
221
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1972]
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222 Rhodora [Vol. 74
sample. The original magnifications of the photographs
of whole spores were taken at 1000 X and portions in
greater detail at 5000 X. Spore measurements are of the
greatest diameter of spores in lactic acid, including the
outer perine layers. Range and mean measurements are
based on samples of 100 spores — 25 from each of four
collections — except for Pellaea notabilis, in which 50
spores were measured from each of two collections.
Measurements of the pinnae are of the longest fertile
pinnae in the central portion of the lamina. Data are drawn
from the revisionary study cited and additional specimens
in collections of the Gray Herbarium, the Field Museum of
Natural History, and the Smithsonian Institution.
ACKNOWLEDGMENTS
I am most appreciative of the help of Drs. Donald M.
Britton and Rolla Tryon for reading and helpful comments
on the manuscript. I am also grateful to Dr. Robert H.
Mohlenbrock for the use of the Cambridge Stereoscan
scanning electron microscope, at the University of Southern
Illinois, Carbondale and the assistance there of Judith
Murphy with Figs. 31-33. Other spores were examined with
an AMR instrument model 900, Harvard University; the
help of G. R. Pierce in the McKay Laboratory is much
appreciated. Drawings of the habit and leaf detail were
done mostly by Sarah Babb Landry but also by Mary
Robbins.
GEOGRAPHY AND ECOLOGY
The leaves of Pellaea atropurpurea (L.) Link are di-
morphic, and several specimens from Mexico have leaves
with mostly simple pinnae as in Fig. 1. The species ranges
from Guatemala northward along the Sierra Oriental of
Mexico, northeast across the United States to Vermont,
southern Quebec and Ontario, and there are disjunct
records from areas in southern British Columbia and
Alberta and also north of Lake Athabaska, Saskatchewan
(Fig. 3). It occurs on exposed or shaded ledges and in
creviees of limestone cliffs, on talus slopes, masonry walls,
1972] Pellaea atropurpurea — Tryon 223
Figs. 1, 2. Habit of Pellaea: Fig. 1. P. atropurpurea with leaves
of most simple form, Puebla, Arséne 3548 (GH); Fig. 2. P. notabilis
with dimorphic leaves, Nuevo Leon, Knobloch 1963, (GH), a. Young
plant, b, Fertile leaves on mature plant; both from spores. All X !4.
224 Rhodora [Vol. 74
or in rocky loam in woods, at 300-5200 meters. At its
southernmost station in Guatemala it grows in crevices of
ancient Indian pyramids. At Campbell’s Bay, in Quebec,
near Ottawa, it occurs on precambrian rocks in which lime-
stone is incorporated. In Ontario it is reported by Rigby
(1968) to occur in sheltered locations on talus or limestone
paving, rather than exposed cliffs.
Leaves of mature plants of Pellaea notabilis Maxon are
usually dimorphic and the pinnae simple. The first leaves
are entire and cordate, and the older ones rarely have a
compound lamina as in Fig. 2. Collections are known from
the vicinity of Victoria, Tamaulipas, at 320 m. and from
Nuevo Leon (Fig. 3). In the latter state it grows on cal-
careous rocks at 1100 m. in San Francisco canyon, south
of Monterrey, and P. atropurpurea has also been collected
there. Plants of P. notabilis, grown from spores of two
collections from this canyon, produced fertile leaves in six
months. The rapid and vigorous growth of these plants in
culture suggests that scme external factors are responsible
for the limited distribution of the species.
Peliaea ternifolia (Cav.) Link car. ternifolia is the widest
ranging of all the American pellaeas occurring from south-
ern Texas to Argentina and Hawaii, at 200-4000 m. Both
diploids and tetraploids (Figs. 5, 6) are known in the
variety, and beth occur at the same localities in the states
of Durango 2nd Michoacán as well as the stations reported
here. Ecological data from these collections as well as field
observations show that the tetraploids grow in mcre shaded,
wetter sites than the diploids. The localities for the three
cytologically documented plants are shown on the map of
the general range of the variety in continental North
America (Fig. 7). In addition to these tetraploids other
specimens which are morphologically similar, including
those reported by Pray (1967) from Mexico and Texas, are
included on the map.
The leaves of Pellaea ternifolia vav. Wrightiana (Hook.)
A. Tryon (Fig. 4) most closely resemble the form of the
diploids in var. ternifolia, and variation in these is noted
Fig. 3. Pellaea atropurpurea and P. notalilis: P. atropurpurea
outlined dots-disjunct stations, circles-cytological records; P. notabilis
stars-the upper a cytological record.
226 Rhodora [Vol. 74
Figs. 4-6.
Habit of Pellaea ternifolia: Fig. 4. var. Wrightiana,
Burro mts, New Mexico, Rusby in 1880 (GH); Figs. 5, 6. var. terni-
folia: Fig. 5. Diploid, Puebla, Riba 408 (GH). Fig. 6. Tetraploid
from culture, San Luis Potosí, Rollins & Tryon 58218 (GH). All
x lé.
1972] Pellaea atropurpurea — Tryon 227
7
Figs. 7, 8. Range of Pellaea ternifolia: Fig. 7. var. ternifolia
outlined, dot-disjunct station, stars and circles-tetraploids, circles-
cytological records, half circles-localities of both diploids and tetra-
ploids; Fig. 8. var. Wrightiana, circles-tetraploids, star-triploid,
228 Rhodcra [Vol. 74
in the section under morphology. The geographic range
of var. Wrightiana extends from southern Oklahoma south-
west to Arizona and across the Mexican border in northern
Chihuahua, Coahuila and Baja California, at 1400-2300 m.
(Fig. 8). A disjunct station is also reported about 1000
miles eastward in Alexander Co. in western North Caro-
lina. This latter record may indicate a formerly more
continuous distribution of these plants across the southern
United States similar to the present range of P. atropur-
purea, or possibly long distance dispersal from populations
farther west. Specimens are mostly from exposed situa-
tions, among sandstone or granitic rocks or rarely on lime-
stone. A few collections of somewhat larger specimens are
from shade, in moist forested sites.
CYTOLOGY
Mitosis and meiosis were examined in plants of Pellaea
atropurpurea from Chipinque Mesa, south of Monterrey,
Mexico. In the mitotic cells many chromosomes have sub-
terminal centromeres as shown in Fig. 11. Differences in
the position of centromeres, in chromosome size, and the
occurrence of satellites, are diagnostic aspects useful in
karyotype analyses. In these apogamous plants most spor-
angia have eight spore-mother-cells, and some sixteen-celled
sporangia are also formed. In meiotic nuclei, the nucleoli
are conspieuous and persist through diakensis. Their struc-
ture can be distinguished from the bivalents in Fig. 9 by
the circular form and lighter stain. In the eight-celled
sporangia the chromosomes are usually associated in bi-
valents (Figs. 9, 16), and this is also reported by Manton
(1950) and Rigby (1968). In apogamous plants the
chromosome number in sporangia with eight spore-mother-
cells, according to Manton, undergoes premeiotic doubling
to accommodate reduction. In sixteen-celled sporangia, at
meiosis, Manton (1950) and Britton (in Tryon & Britton,
1958) report approximately 29 pairs and 29 single chromo-
somes. The pairing associations in several] cells from both
types of sporangia were studied by Rigby. In these, two
cells from sixteen-celled sporangia (a specimen from Que-
a
Figs. 9-14. Meiotic and mitotic nuclei: Figs. 9-11. Pellaea
atropurpurea: Fig. 9. Nucleus from 8 celled sporangium with 87
bivalents, the nucleolus at upper right, X 660; Fig. 10. Early stage
in sporogenesis, micronucleus and 4 nuclei at center, X 600; Fig. 11.
Mitotic chromosomes from young sporangium. All triploid, n=87,
Rollins & Tryon 5860 (GH). Fig. 12. Meiosis, P. notabilis, diploid,
n—29 Marroquin 1855 (GH) 660. Figs. 13, 14. Meiosis, P. ternifolia
var. ternifolia: Fig. 13. Diploid P. ternifolia var. ternifolia; Fig.
13. Diploid, »—29, Riba 408 (GH) X 1000; Fig. 14. Tetraploid, »—58,
Riba 412 (MEXU) X 1000.
230 Rhodora [Vol. 74
bec) had equal numbers of pairs and univalents, but one
had nine and the other six trivalents. A second collection
(from British Columbia) had 31 bivalents, 22 univalents
and one trivalent. Affinities between the parental genomes
are expressed in these cells by the associations of more than
29 bivalents and also the trivalent formations. The varia-
tions in chromosome associations in these cells and also
the lack of multivalents in the eight-celled sporangia, which
have doubled the chromosome number, raise problems in
the interpretation of chromosome pairing in these plants.
There are frequent irregularities as shown in the early
stage of sporogenesis in Fig. 10, in the sixteen celled
sporangia, with an extra micronucleus.
Two collections of Pellaea notabilis from Nuevo Leon
uniformly had 29 bivalents at meiosis as in Figs. 12, and
15. This establishes the species as a diploid with the basic
chromosome number for Pel/laea. The number is consistent
with the species being proposed as one of the elements
involved in the origin of the triploid P. atropurpurea.
Both diploid and tetraploid plants of Pellaea ternifolia
var. ternifolia were studied from the same localities near
San Luis Potosi and also farther south near Puebla. There
are earlier records of these two polyploid levels, reported
by Britton, from localities near Mexico City but not at the
same site. The diploid with 29 bivalents and the tetraploid
with 58, in Figs. 17, 18 are collections from Puebla. Con-
trast in the number of bivalents at late metaphase in these
two polyploid levels is readily apparent in the figures. The
terminal portions of the chromosomes in the centromeric
regions in Figs. 12, 14 are stretched thinner than the re-
mainder of the chromosomes forming the bivalents. These
attenuated portions show tension on the late metaphase
chromosomes as they prepare to move to the poles. The
tetraploid plants generally resemble the diploids, but the
pinnae tend to be entire and the rachises more pubescent
(Figs. 5, 6) as compared to the diploids, which have ternate
pinnae and nearly glabrous leaves. The drawing of the
tetraploid in Fig. 6 represents an especially robust plant
1972] Pellaea atropurpurea — Tryon 231
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showing bivalents in focus in black, others in outline, nucleolus stip-
pled, from the same cells except P. atropurpurea: Fig. 15. P. nota-
bilis, Fig. 16. P. atropurpurea, n=87, Rollins & Tryon 5860 (GH);
Figs. 17, 18. P. ternifolia var. ternifolia with thin portions omitted:
Fig. 17. Diploid, Fig. 18. Tetraploid.
232 Rhodora [Vol. 74
that was grown in culture for several months. At Hacienda
Batán, near Puebla, the tetraploids are rare, occurring in
moist, shaded places while the diploids grow in exposed,
rocky crevices. Other specimens, which appear to be tetra-
ploids on the basis of the characters noted above, are quite
widely distributed as is shown by the stars on the map in
Fig. 7.
The cytological records of Pellaea ternifolia var. Wright-
iana are based on three earlier reports from localities in
the United States. Tetraploids with » — 58 were noted
from the western part of the range in New Mexico by Lloyd
(1968) and from the disjunct station in North Carolina
by Wagner (1965). These plants were considered to be
amphiploid hybrids of the diploids P. ternifolia var. terni-
folia and P. truncata Goodd. (P. longimucronata Hook. ).
A triploid plant of var. Wrightiana with 29 bivalents and
29 univalents from Arizona was reported by Knobloch &
Britton (1963). This plant was considered to have origi-
nated as a hybrid between the diploid P. truncata and a
tetraploid plant of var. ternifolia. Knobloch & Britton
report the same chromosome number in both mitotic and
meiotic cells and also 64 irrgular spores in the sporangia.
A specimen (Pringle, May 6) from the Santa Catalina
mountains, which also appears to be a variant as noted in
the survey of the spores, indicates the need for further
cytological sampling of var. Wrightiana.
SPORES
In Pellaea atropurpurea only spores of the 32-spored
sporangia mature, and this reduced number reflects the
apogamous type of reproduction. The spores are globose
or somewhat ellipsoidal (Fig. 19) and unusually large,
ranging from 54-92 „ in diameter. The triradiate scar
scarcely projects, and is apparent only in cleared material,
with arms extending 34, or more of the radius. The perine
ridges are similar to those in P. notabilis but are sharper
and more prominent. At higher magnification the perine
surface consists of compacted sporopollenin particles (Fig.
20).
1972] Pellaea atropurpurea — Tryon 233
Figs. 19-22. Spores cf Pellaea: Figs. 19, 20. P. atropurpurea:
Fig. 19. Nuevo Leon, Rollins & Tryon 5860 (GH), X 660; Fig. 20.
Perine surface, Hac. Batán, Arsène 3548 (GH), X 3300. Figs. 21:22.
P. notabilis: Fig. 21. Knobloch 1963 (GH), X 660; Fig. 22. Perine
surface, X 3300.
234 Rhodora [Vol. 74
In Pellaea notabilis sporangia have 64 well formed spores
that are globose or somewhat ellipsoidal (Fig. 21) and
range from 40-62 , in diameter. The slightly larger spore
size in this diploid, relative to diploid spores in P. terni-
folia var. ternifolia, may be partly due to the inclusion of
the prominent ridges in the measurements. These dense
ridges obscure the triradiate scar, but in cleared material
it projects slightly and the arms extend ?4 of the spore
radius. At higher magnification the perine deposition ap-
pears uniformly verrucate on both floor and ridge surface
(Fig. 22).
In both diploid and tetraploid forms of Pellaea ternifolia
the sporangia uniformly have 64 spores. Those of the
tetraploid are consistently larger, ranging from 46-62 , as
compared to the diploids which range from 38-58 ,. The
size differences are especially apparent in Figs. 23 and 25
and have been reported also in other collections by Pray
(1968). Spore shape is generally spherical or tetrahedral-
globose especially in the tetraploid which has broader sur-
faces between the arms of the triradiate scar. These arms
extend 34, or more of the spore radius. The perine is
generally similar in spores of both polyploid levels, but
variation occurs in density and prominence of the ridges.
These may be especially prominent in the diploid spores
(Fig. 24), and on the smoother surface of the tetraploid
spores (Fig. 26) there are often numerous sporopollenin
particles.
Spores of Pellaea ternifolia var. Wrightiana were ex-
amined from populations throughout its range. Sporangia
usually have 64 spores of relatively uniform size. A speci-
men (Pringle, May 6, 1883, GH) collected in the Santa
Catalina mountains in Arizona had many irregular spores
and some small ones as shown in Fig. 34. This specimen
may represent another triploid, similar to that reported
from the same area by Knobloch and Britton (1963). The
spores of var. Wrightiana are usually globose with a promi-
nent triradiate scar extending 9/4 or more of the spore
radius. The form of the perine is relatively uniform as
Figs. 23-28. Spores of Pellaea ternifolia: Figs. 23-26. var. terni-
folia: Fig. 23. Diploid, X 660; Fig. 24. Perine surface, X 3300, both
San Luis Potosí, Rollins & Tryon 58219 (Gu); Fig. 25. Tetraploid,
X 660; Fig. 26. Perine surface with portion of triradiate scar, X
3300, both Rollins & Tryon 58218 (GH); Figs. 27, 28. var. Wright-
iana: Fig. 27. Gillispie Co., Texas, Correll & Correll 12762 (Gu),
X 660; Fig. 28. Santa Catalina mts. Arizona, Phillips & Reynolds
2943 (GH), X 660.
236 Rhodora [Vol. 74
Figs. 29-34. Spores of Pellaea ternifolia var. Wrightiana: Fig. 29.
Lower spore with portions of perine eroded, smooth exine surface at
right, X 660; Fig. 30. Perine surface with portion of triradiate
scar, X 3300, both, Coahuila, Johnston & Miller 1509 (GH); Figs. 31-
33. Alexander Co., North Carolina, Bozeman et al. 45152 (stu):
9
Fig. 31. X 660, Fig. 32. Perine surface, X 3300, Fig. 33. Spore with
partly eroded perine and subtending scabrous layer, X 660, Fig. 34.
Santa Catalina mts. Arizona Pringle, May 6, 1883 (GH), X 660.
1972] Pellaea atropurpurea — Tryon 237
shown in the series of specimens from Mexico, Arizona,
Texas, and North Carolina (Figs. 27-34), although there
is some variation in size and prominence of the perine. The
specimens from North Carolina have an especially rugose
perine with stratification as shown in Fig. 33, showing a
coarse outer rugose layer subtended by a strongly scabrous
one. Similar stratification is also evident in the specimen
from Mexico in Fig. 29, which has some scabrous deposition
on the smooth surface of the spore exine where the perine
has been abraded.
The differences in prominence of the triradiate scars in
these species may imply distinctions in spore development.
The prominent triradiate scars with scant perine deposition
characteristic of the spores of P. ternifolia indicate that
these may have stronger tetrad associations. In contrast
to these, the obscure triradiate scars in P. notabilis and
P. atropurpurea suggest the tetrads may be disassociated
before completion of the perine deposition.
MORPHOLOGY
Comparisons of the morphology of the species and
varieties are made in Table 2. Similarities between Pellaea
notabilis and P. atropurpurea (Figs. 1, 2) are especially
evident in the dimorphic leaves, terete shape of the petiole,
longer pinnae, and concolorous rhizome scales. These two
species are readily distinguished from P. ternifolia by these
characters. There are also marked similarities among the
three kinds of P. ternifolia (Figs. 4-6) in the petiole shape,
monomorphic leaves, and bicolorous rhizome scales. The
form of the perine, in addition to the characters shown in
the chart, also establishes a close relationship among them.
These similarities express relationships that, by compari-
son to other species of Pellaea, can appropriately be recog-
nized by including them in a single species.
Morphological variation in collections of Pellaea ternifolia
var. Wrightiana was surveyed over the whole geographic
range. Most of the specimens have leaves most closely
resembling the form of the diploid var. ternifolia (Figs.
4,5). Some variation is shown in the division of the leaves,
{ Vol. 74
Rhodcra
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especially on the same plant, and this appears to be corre-
lated with leaf size. In these specimens the leaves with
shorter pinnae rachises and nearly ternate pinnae, re-
sembling those of var. ternifolia, are usually smaller and
seem to be produced before the more highly divided leaves
resembling those of P. truncata. A similar correlation
between leaf size and division is also evident in the plant
from North Carolina figured by Wagner (1965). Thus,
variation in leaf form in plants of var. Wrightiana shows
types intermediate between var. ternifolia and P. truncata
and also differences in developmental sequence.
DISCUSSION
The report of 29 bivalents in Pellaea notabilis establishes
this as one of the basic species among the American
pellaeas. Similarities of the spores and other morphological
characters indicate that it is probably one of the elements
involved in the origin of the widely distributed triploid,
P. atropurpurea. On the basis of this new information
several possibilities may be considered for the origin of
P. atropurpurea.
The strong morphological resemblances between the two
species suggest that Pellaea atropurpurea may be an auto-
ploid of P. notabilis. The triploid could have originated
from two gametes of P. notabilis, one of them unreduced.
The studies of P. atropurpurea by Rigby show that affinities
do exist in the two parental genomes.
However, several unique characters of Pellaea atropur-
purea must also be considered. The broad leaves up to
three times pinnate with densely pubescent rachises sug-
gest that a second distinctive parent is involved in addition
to P. notabilis. The cytological and morphological survey
of P. ternifolia, the most likely parent in northern Mexico,
shows that none of its variants completely fills the morpho-
logical requirements of the second parent. The spores of
P. ternifolia are especially diagnostic, and the perine form
is quite different from that of P. atropurpurea. The unique
morphological features of Pellaea atropurpurea are in-
consistent with an autoploid origin from P. notabilis, and
240 Rhodora [Vol. 74
the characters of its closest relatives near the present range
of that species are inconsistent with an amphiploid origin
involving one of them. The most reasonable conclusion at
this time is that either the second parental element of P.
atropurpurea belongs to another species group possibly not
within the geographic center, or it may be extinct. The
rare occurrence of P. notabilis, its closest relative in north-
ern Mexico, suggests that the other parent may no longer
be extant.
The extensive distribution of P. atropurpurea is un-
doubtedly partly based on the absence of syngamy. Elimi-
nation of the conditions required for fertilization results
in more broadly adapted apogamous gametophytes. The
large spores may also provide additional reserves, increased
survival capacity, and a greater energy source for more
rapid development of apogamous gametophytes and spore-
lings. In Pellaea ovata (Desv.) Weath., which has both a
sexual (2x) and an apogamous (32) race, the range of
the latter is over 4000 miles greater than that of the sexual
race. The effectiveness of the apogamous reproductive
system is shown by the wider geographic distribution of
the apogamous race in that species as well as in P. atro-
purpurea.
GRAY HERBARIUM,
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS 02138.
1972] Pellaea atropurpurea — Tryon 241
LITERATURE CITED
KNOBLOCH, I. W. & D. M. BRITTON. 1963. The chromosome number
and possible ancestry of Pellaea Wrightiana. Amer. Journ. Bot.
50: 52-55.
LLovp, R. M. 1966. I.O. P. B. Chromosome number reports. Taxon
15: 283.
MANTON, I. 1950. Problems of Cytology and Evolution in the
Pteridophyta. Cambridge University Press.
Pray, T. R. 1967. Notes on the distribution of some American
cheilanthoid ferns. Amer. Fern Jour. 57: 52-58.
RIGBY, S. J. 1968. An investigation of Pellaea glabella and Pellaea
atropurpurea and their relationships. University of Guelph,
Guelph, Ontario. M. S. thesis, 120 pp.
Tryon, A. F. 1957. A revision of the fern genus Pellaea section
Pellaea. Ann. Missouri Bot. Gard. 44: 125-193.
TRYON, A. F. & D. B. BRITTON. 1968. Cytotaxonomic studies on the
fern genus Pellaea. Evolution 12: 137-145.
WAGNER, W. H. 1965. Pellaea Wrightiana in North Carolina and
the question of its origin. Jour. Elisha Mitchell Sci. Soc. 81:
95-103.
THE GENUS CORIARIA (CORIARIACEAE)
IN THE WESTERN HEMISPHERE’
LAURENCE E. SKOG
With the recent awakening of research in hallucinogenic
plants, some attention has been given to the genus Coriaria.
Plants of this genus are known to be toxic to animals when
taken internally, as well as being hallucinatory in small
quantities. Studies of its toxicity and the application of
the toxic elements in treatments of barbituate poisoning
have made necessary a basic taxonomic study of Coriaria.
This paper deals with taxonomic studies of the plants
which occur in the Western Hemisphere. Coriaria inhabits
warm temperate regions of southern South America, and,
in the tropical areas, high elevations of the northern Andes,
ranging north through Central America to the Sierra
Madre Occidental in Mexico. Coriaria is known also from
southern Europe, North Africa, the Himalayas, islands of
the Pacific coast of Asia and the western Pacific Ocean, in-
cluding New Zealand. The genus makes an interesting
study of geographical distribution and dispersal which has
resulted in highly disjunct populations. The plants of the
Western Hemisphere have close relatives on the other side
of the Pacific Ocean, but these are excluded from considera-
tion here.
PHYLOGENY OF THE GENUS
As the only genus in the family Coriariaceae, Coriaria
has no close relatives and no close morphologic affiliation
with any other family. In 1816 De Candolle placed Coriaria
in the Rhamnaceae. Since that time the position of Coriaria
in a phylogenetic system has changed often with various
authors, ranging from alliances with the Urticaceae (Max-
imowicz, 1881) to a much more advanced position near the
Araliaceae (Russow, 1884). Most often the genus has been
‘Based on a thesis submitted to the Graduate School of the Uni-
versity of Connecticut in partial fulfillment of the requirements for
the degree of Master of Science.
242
1972] Coriaria — Skog 243
placed near the Anacardiaceae as was done by Bentham
and Hooker in 1862. Hutchinson in 1964 placed Coriaria
in its own order, Coriariales, between the Dilleniales and
the Rosales, and derived from the former. Cronquist
(1968), with a note of doubt, has placed the Coriariaceae
in the Ranunculales.
The phylogenetic relationships of the Coriariaceae can-
not be determined accurately without, perhaps, the dis-
covery of some “pre-Coriaria” which would link the family
to another group. It is apparently not closely related to any
single other family; it has long been distinguished and
kept separate from other families and probably should not
be closely associated with others in the same order. There
is, however, no single character of the family that sets
it apart, but a combination of traits. If the dicots are
viewed as polyphyletic, Coriaria could be one of the distinct
lines which branched from the main line of evolution some-
time in the Cretaceous, retaining many primitive characters
while evolving other more advanced or specialized traits.
If one lists the characteristics of dicotyledonous families
and then attempts to place a family among them, correla-
tions become evident which can sometimes be interpreted
as showing evolutionary relationships; with Coriaria this
has not been successful. Various characters of dicots are
considered as being either primitive or advanced. These
often determine the position of the family in a phylogenetic
order. Maximowicz (1881) attempted to draw conclusions
about relationships by using morphological characters of
various families, then weighting them as to their import-
ance in Coriaria. After totaling the values, Maximowicz
concluded the genus was most closely allied to the Sima-
roubaceae, and also had affinities with the Phytolaccaceae.
If one had to select another family in which Coriaria could
be placed, one might choose the Rosaceae. The characters
of Coriaria fall within the broad limits of this family, as
defined or delimited by Lawrence (1951) and Hutchinson
(1964). Within the Rosaceae, though, there is no single
genus to which Coriaria is closely related. The genus fits
244 Rhodora [Vol. 74
less well in the Simaroubaceae or Erythroxylaceae, and
only superficially in the Brunelliaceae. Paralleling this
viewpoint, Hutchinson has placed Coriaria in a distinct
order between the Brunelliaceae and Rosaceae, and Hallier
(1905) placed the Coriariaceae in the Rosales, but allied
to the Simaroubaceae.
TAXONOMY
Coriaria L., Genera Plantarum, ed. 5. 459. 1754. Type:
Coriaria myrtifolia L.
Heterocladus Turez., Bull. Soc. Imp. Naturalistes Moscou
20: 152. 1847. Type: H. caracasanus Turcz. Non Hetero-
cladia Dnd. as Hererocladia Dnd. ex Turcz., sphalm. ex
Jacks.
Heterophylleia Turez., Bull. Soc. Imp. Naturalistes
Moscou 21: 591. 1848. Type: H. caracasana (Turcz.)
Turcz.
Subshrubs to small trees. Roots fibrous, with prominent
nitrogen-fixing nodules. Stems rhizomatous, the branches
sharply quadrangular, becoming terete, eventually arching
to pendulous, the lenticels corky. Clusters of axillary buds
produced in the axils of the leaves of the main axes. Leaves
simple, whorled to opposite, appearing distichous through
alternate twisting of the branches, often approximate
above, short-petiolate to subsessile, the minute stipules
withering and persistent or caducous, the blades palmately
veined, lanceolate to ovate, entire, cordate, truncate to
rounded at the base, the apex rostrate to mucronate.
Racemes erect to pendent, terminating main or lateral
axes. Flowers small, basically pentamerous, actinomorphic,
or the calyx slightly zygomorphic, perfect, polygamo-
monoecious, andromonoecious or appearing so through
marked proterogyny, hypogynous, green, red or dark pur-
ple, the pedicels ebracteolate to several-bracteolate, the
sepals at fertilization, but accrescent and prominently fleshy
subulate, persistent, the petals distinct, shorter than the
sepals at fertization, but accrescent and prominently fleshy
in fruit, ultimately black, red or yellow, stamens 10 in two
1972] Coriaria — Skog 245
whorls, with filiform filaments 2 mm. long, each anther
sagittate, smooth to verruculose, 4-celled, dehiscing by
longitudinal slits, the pollen spheric, 3-colporate, pistils 5
to 10 (-12), distinct, simple, the styles fasciculate and
strict or divergent, papillose and stigmatic over their
entire surfaces. Ovule solitary, pendant. Fruits dorsally
keeled achenes, each bilaterally compressed, all enclosed
by the fleshy accrescent petals. Seeds with a single mem-
branous testa and scanty endosperm.
Key to the species and subspecies of Coriaria. (All known
species included here for completeness. )
a. Bases of lateral branches nearly always without bud
scales; inflorescences produced terminally, either on the
main stem or on branches, but on the current flush of
growth; racemes sparsely to very densely pubescent;
bracts subtending the flowers lanceolate to elliptic, en-
tire; flowers strictly perfect; inner whorl of stamens
free; achenes 1 to 2 mm. long. (b)
b. Stems herbaceous, dying to the ground following
maturation of fruits; leaves of the flowering axes
ovate to suborbiculate, with 7 to 9 main basal veins;
fruit pedicels more than 1.5 cm. long. Himalayas of
Tibet to China. .... 4478 Coriaria terminalis
b. Stems woody, perennial; leaves of the flowering axes
lanceolate to elliptic, with 3 to 5 main basal veins;
fruit pedicels less than 1 cm. long. .. V.
Coriaria ruscifolia
b e oo o o D e $4.» o o we. 68) Q ^. 9 a ee
bearing numerous short branches bearing inflor-
escences terminally; leaves 0.5 to 3.1 cm. long.
Western slopes of the Andes from Mexico to
Peru, and cool mountains of New Guinea and
New Zealand. ose nun PO aa ea "e
Coriaria ruscifolia subsp. microphylla
c. Large shrubs to small trees; main axes bearing
inflorescences in the axils of the leaves or rarely
on short branches; leaves 1.0 to 7.5 cm. long.
Temperate coasts of Chile, New Zealand and
246 Rhodora [Vol. 74
other Pacific islands. ....................00.
ZEE Coriaria ruscifolia subsp. ruscifolia
a. Bases of lateral branches surrounded by persistent bud
scales; inflorescences produced from the axils of the
leaves of the previous flush of growth; bracts subtend-
ing the flowers ovate to orbiculate; flowers usually
andromonoecious; inner whorl of stamens epipetalous.
Shrubs of the Northern Hemisphere. (d)
d. Racemes finely puberulent to densely pubescent;
achenes 1.5 to 2.5 mm. long; bracts subtending the
flowers subentire to erose. Northern India, Nepal
and Western China. ......... Coriaria nepalensis
d. Racemes glabrous; achenes more than 3 mm. long:
bracts subtending the flowers entire. (e)
e. Accrescent petals far exceeding the mature
achenes and enclosing them; leaves ovate-lanceo-
late; styles linear to fusiform. Japan, Formosa
and the Philippines. ....... Coriaria japonica
e. Accrescent petals not exceeding the exposed
mature achenes; leaves subelliptic; styles fili-
form. West-Mediterranean region. ............
2*2 25 2 2-222929» Coriaria myrtifolia
Coriaria ruscifolia L., Species Plantarum, ed. 1. 1037.
1753.
Plants fruticose; new branches sprouting and developing
rapidly, fleshy, at first terete, green to purple to gray,
secondary growth producing a quadrangular stem which is
brown or gray at maturity, cork developing early, lenticels
present only on the previous flush of growth, scaly terminal
buds not formed.
Leaves subsessile, glabrous and dark green above, puberu-
lent on the emersed veins and paler beneath ; leaves of the
main stem orbiculate, with 5 to 9 main basal veins, those
of the lateral branches elliptic to lanceolate, with 3 to 5
main basal veins, acute to cordate at the base, the apex
acute to mucronate.
Racemes few to numerous, pendent, finely to densely
pubescent, the pedicels ebracteolate, 3 to 6 mm. long, at
1972] Coriaria — Skog 247
maturity less than 1 cm. long, with entire, lanceolate to
subulate bracts. Flowers 2 to 3 mm, in diameter, usually
pentamerous, regular, perfect, proterogynous, sepals ovate,
green to red, 1.5 to 2 mm. long, 1 to 1.5 mm. wide, petals
smaller, ovate, anthers yellow to reddish, styles linear,
fasciculate, green to purple, matured achenes enclosed by
the accrescent fleshy, black petals, 1 to 2.5 mm. long.
The numerous populations of Coriaria found in the West-
ern Hemisphere are here regarded as two subspecies of a
single species, polymorphic in leaf size and shape and habit
in response to geographic location and altitude. The char-
acters which unite all these plants into one species are
listed below:
1. The inflorescences are produced on wood of the cur-
rent growing season. These racemes are either borne
terminally on the lateral leafy branches or from the axils
of the large orbiculate leaves on the main axes.
2. The leaves on the lateral branches have 3 to 5 veins.
3. The flowers have the same morphology throughout
the distributional range and are always perfect, a trait
shared only with Coriaria terminalis of the Himalayas.
The two subspecies have heretofore been maintained as
separate species since they were first described. Reducing
these taxa from species to subspecies reflects their simi-
larity and at the same time illustrates the differences in
morphology which are ecological adaptations to altitudinal
changes. At higher elevations, 600 m. to 4000 m. in the
Andes, small-leaved forms occur while larger-leaved forms
grow at elevations from sea level to 300 m. in southern
South America. Another character which varies with alti-
tude and which is noticeable in dried specimens as well as
in the living plants is the increased numbers of lateral
branches occurring with increased elevation. The correla-
tion with altitude of these variations, independent of genetic
constitution, has been demonstrated by comparisons of
seedlings of South American forms grown under identical
greenhouse conditions at the University of Connecticut.
Seeds were collected in Colombia at 3200 m. (Skog 1000)
248 Rhodora [Vol. 74
and Chile at 5 m. (Skog 1075), germinated and grown to-
gether in the greenhouse. The resulting plants were similar
in leaf size and shape, and bore few lateral branches.
(Herbarium vouchers are on file at CONN of the repre-
sentative plants, Skog 1375, 1376.) Studies of herbarium
specimens and field observations indicate that when plants
otherwise assignable to the small-leaved subspecies, C.
ruscifolia subsp. microphylla, are found at lower eleva-
tion (less than 1000 m.) the leaves are larger and the
individuals have fewer lateral brances.
Coriaria ruscifolia L. subsp. ruscifolia
Coriaria ruscifolia L., Species Plantarum, ed. 1. 1037.
1755. Type: Feuillée, Journal des observations physiques,
mathématiques et botaniques 3 [Histoire des plantes méde-
cinales . . . du Perou & du Chily]: 17, t. 12. 1725?
Large shrubs to small trees. Main axes to 7 m. long.
bearing whorled or opposite leaves 1.0 to 7.5 cm. long,
0.8 to 3.2 em. wide. Racemes arising from the axils of the
leaves of the main axes, or at the apices of the few lateral
branches, to 25 cm. long.
Vernacular names: CHILE: deu, dehue-lahuen, deó,
hiuque, mataratones, veu.
Distribution and habitat in the Western Hemisphere:
River banks or lake shores in western Argentina in the
province of Neuquén, and in central and southern Chile in
the provinces of Arauco, Biobio, Cautin, Chiloé, Concepción,
Llanquihue, Malleco, Maule, Nuble, Osorno, Santiago and
Valdivia.
Representative specimens examined: ARGENTINA. PROVINCE OF
NEUQUÉN: Lago Nahuel Huapi, Isla Victoria, Cordini 8 (F, US).
CHILE. PROVINCE OF ARAUCO: Contulmo, Gunckel 21784 (CSG) ;
Santa Cruz s. n., 1.835 (BH). PROVINCE OF BIOBIO: Fundo ‘Los Prados,’
Riegel s. n., Jan. 1, 1955 (CSG). PROVINCE OF CAUTIN: Rio Quepa,
Temuco, D. Bullock s. n., 31 Jan. 1906 (BM); Quebrada del Río Pal-
quin, Gunckel 15413 (csc); Puerto Saavedra, Hollenmayer 8 (CSG, M).
PROVINCE OF CHILOÉ: Chiloé, J. Anderson 61 (BM); Piruquina,
Junge s. n., Mar. 25, 1932 (csG) ; Junge 386 (M, MO); near Puerto
Varas, 50 m., Morrison 17537 (BH, GH, MO) ; Castro, 1-10 m., Pennell
1972] Coriaria — Skog 249
12600 (F, GH, SGO, US); cerros cerca de Chonchi, Ricardi 5308 (CONC,
CONN). PROVINCE OF CONCEPCION: Laraquete, Castillo s. n., Feb.
1946 (csG); Conception, Cuming 1461 (BM); Tomé, Germain s. n.,
Nov. 1855 (GH, sco); Talcahuano, Póppig 131 (BM, MO); near
Tumbes, 5 m., Skog 1040 (CONN). PROVINCE OF LLANQUIHUE: Lago
Llanquihue, Calvert s. n. in 1912 (BM); Petrohue, Gunckel 9227
(csG); Piedra Azul, Ricardi 5286 (CONC, CONN); Casa Panque, Shan-
non & Shannon 24 (US); near Los Riscos, 200 m., Skog 1073, 1074,
1075 (CONN); near Ensenada, 100 m., West 4678 (GH, MO). PROVINCE
OF MALLECO: Puren, Claude-Joseph 3029 (US). PROVINCE OF MAULE:
Constitución: Burnier s. n. Sept. 1958 (csG), Gunckel 21785 (csa),
Volekmann s. n. (SGO). PROVINCE OF NUBLE: Valle Rio Renegado:
Alvares Ramirez s. n., Mar. 1954 (csG), Cuming 146 (E, GH); Valle
de Alico, Jarpa s. n., Feb. 10, 1935 (csG); Reed s. n., 1871 (BM, GH);
Sargent s. n., Jun. 29, 1906 (A). PROVINCE OF OSORNO: Fundo Río
Blanco, southwest of Purranque, Eyerdam 10758 (F, US); Lago
Llanquihue, F. Phillipi & Barchers s. n., 15/1/85 (BM); north shore
of Lago Llanquihue, Senn 4635 (MO, US); southwest of Puerto
Klocker, near Lago Llanquihue at base of Volcán Osorno, Skog 1062,
1064 (CONN). PROVINCE OF SANTIAGO: Claude-Joseph 957 (GH, US).
PROVINCE OF VALDIVIA: Valdivia, Bridges 612 (BM); Uferbüsch des
Calle-calle, Buchtien s. n., Sept. 17, 1896 (E, GH, M, US) ; ad margines
rivulorum, Gay 1019 (sao); Isla del Ray, Carbonares, Gunckel 1097,
1149 (cse); Valdivia, Buchtien s. n., 12.X1.1904 (LD); Valdivia,
Lechler s. n. (M).
Coriaria ruscifolia L. subsp. microphylla (Poir.) L. Skog
stat. nov.
Coriaria microphylla Poir., Encyclopédie méthodique.
Botanique 4: 87. 1804. Type: J. Jussieu s. n., P-JU.
Coriaria thymifolia Humb. & Bonpl. ex Willd., Species
Plantarum, ed. 4. 4: 819. 1805. Type: Humboldt & Bon-
pland. 3018, B, P.
Coriaria phylicifolia Humb. & Bonpl. ex Willd., Species
Plantarum, ed. 4. 4: 819. 1805, ex char.
Coriaria atropurpurea DC., Prodromus systematis na-
turalis regni vegetabilis 1: 740. 1824. Type: De Candol!e,
Calques des dessins de la flore du Mexique de Mocino et
Sessé, t. 1167.
Heterocladus caracasanus Turcz. Bull. Soc. Imp. Natu-
ralistes Moscou 20: 152. 1847. Type: Galeotti 293, LE.
Heterophylleia caracasana (Turez.) Turez., Bull. Soc.
Imp. Naturalistes Moscou 21: 591.
250 Rhodora [Vol. 74
Coriaria cuneifolia Sessé & Moc., Plantae Novae His-
paniae, ed. 1. 173. 1890, ex char.
Suffruticose herbs to large shrubs. Main axes to 4 m.
long bearing numerous short lateral branches. Leaves op-
posite to approximate, 0.5 to 3.1 cm. long, 0.2 to 1.6 cm.
wide. Racemes terminating the lateral branches, to 15 cm.
long.
Vernacular names: COLOMBIA: rebentadera. ECUADOR:
pinan, shanshi, shanchi or zhanzhi, tinta. GUATEMALA:
moco tinto, moco de chompipe. MEXICO: helecho de tierra,
tlalocopetate, tlalocopatlatl. PERU: mio-mio, saca-saca,
mio-venenosa, raqui-raqui. VENEZUELA: tisis, helecho de
playa, helecho-uvite.
Distribution and habitat in the Western Hemisphere:
Moist montane forest openings and hillsides in Mexico,
Guatemala and Panama and on the western slopes of the
Andes in Colombia, Venezuela, Ecuador and Peru.
The plants referred to this subspecies have long been
named C. thymifolia Humb. & Bonpl. ex Willd. Coriaria
microphylla Poir. predates C. thymifolia by almost two
years, however, and should be used if this group is main-
tained as a separate species.
Representative specimens examined: COLOMBIA. ANTIOQUIA: La
Sierra, 18 km. north of Medellín, 2000 m., Archer 1307 (BM, COL, US).
BOGOTÁ: Holton 27, 808 (GH); Triana 5596 (BM, COL); Cordillera de
Bogotá, 2650 m., Triana s. n. (BM, MEXU). BOYACA: Grubb, Curry &
Fernández-P. 519 (COL, US). CALDAS: Killip & Hazen 8956 (GH, US) ;
Laguneta, Salento, 2500 m., von Sneidern 3149 (COL, MO). CAUCA:
Cordillera Central, vertiente occidental, 2470 m., Cuatrecasas 19369
(A, 2 sheets, F); Puracé, Fernández-P. 6218A (COL); entre Popayán
y Puracé, 2500 m., Yepes Agredo 390 (COL, F, US). CUNDINAMARCA:
Salto de Tequendama, 2100 m., Alston 7409 (BM); García-Barriga
130, 17488 (COL); al sur de Usme, entre La Regadera y El Hato,
3000-3100 m., Idrobo, Jaramillo, Mesa-Bernal & Smith 399 (COL, MO,
US). HUILA: Cordillera oriental, east of Neiva, 1300-1800 m., Rusby
& Pennell 1008 (GH, MO, US). MAGDALENA: Van der Hammen 1165
(COL). NARIÑO: Ferndndez-P., et al. 1083 (coL); Pasto, Cebadal,
2700-2800 m., Schultes & Villarreal 7939 (COL, F, US). NORTE DE
SANTANDER: Cuatrecasas & Garcia-Barriga 10064 (COL, US).
PUTOMAYO: Garcia-Barriga 7766 (COL, US); Valley of Sibundoy,
1972] Coriaria — Skog 251
Sibundoy, 2225-2300 m., Schultes & Villarreal 7714 (COL, F).
SANTANDER: Killip & Smith 16784, 19227 (A, GH, US). TOLIMA:
Cuatrecasas 9361 (COL, US). VALLE: Cordillera Central, vertiente
occidental, 2270-2320 m., Cuatrecasas 18140 (F). COSTA RICA.
CARTAGO: Volcán Irazi: 10000-11300 ft., Allen 676 (F, MO); Four-
nier 1202 (CRU); Potrero Cerrado road, 6500 ft., Lankester s. n., 1925
(F, US); 2400 m., Skog 1339 (CONN); Smith 4761 (GH, US, 2 sheets) ;
1400 m., Torres-R. 288, s. n., 1924 (su). ECUADOR. AzuAy: Rio
Tarqui, 8300 ft., Camp E-3905 (F, GH, US). BOLIVAR: entre Guaranda
y Vinchoa, 2800 m., Acosta Solis 5927 (F). CANAR: near the village
of Marcos, Camp E-2488 (MO). CHIMBORAZO: 2450 m., Acosta Solis
5534 (F); Huigra, Rose & Rose 22182 (BM, US). IMBABURA: Shanchi-
pamba, 2750 m., Acosta Solis 14514 (F, 2 sheets). LOJA: 2200 m.,
Dodson & Thein 567 (MO); near Loja, between 2300-2700 m., Skog
1162, 1167, 1168, 1169, 1172, 1192 (CONN). PINCHINCHA: Cord. occ.,
los Alpes, 2800-3000 m., Acosta Solis 7079 (F); vicinity of Quito,
3000 m., Asplund 6157 (LD, US); above Quito, 10300 ft., Balls B5846
(BM, E, MO, US); Quito, Couthouy s. n., 1855 (GH); Quito, Jameson
206 (BM); Alrededores de Quito, 2850 m., Paredes 23 (Q). TUNGU-
RAHUA: entre Leito y la Coma, Cord. oriental, 2700-3000 m., Acosta
Solis 8977 (F); Volcano Tungurahua, 2200-2400 m., Dodson & Thein
1886 (M0); Rio Ambato Valley, W. of Ambato, Fagerlind & Wiborn
965 (LD). LOCALITY NOT DEFINITE: Heinricks 634 (M); interandine
highland, 2000 m., Rimbach 97, (A, F). GUATEMALA. CHIMALTEN-
ANGO: Barranco de La Sierra, southeast of Patzüm, about 2100 m.,
Standley 61508 (A, F). EL PROGRESSO: slopes adjacent to Finca
Piamonte, 2500-3000 m., Steyermark 43711 (A, F). ESQUINTLA:
Texcuaco, 150(?) m., Morales 1061 (F). GUATEMALA: near Finca
La Aurora, 1500 m., Agwilar 248 (F); near Guatemala, Kellerman
5252 (MEXU, US); Pinula, 4200 ft., J. Donnell Smith 1928 (M). HUE-
HUETENANGO: San Juan Atitan, 8400 ft., Skutch 1166 (A, F). JALAPA:
between Miramundo and summit of Montaña Miramundo 2000-2500
m., Steyermark 32753 (F). QUEZALTENANGO: Sunil, Hartweg 524
(BM, GH, LD); near Santa Maria de Jesus, 6400 ft., Skutch 901
(A, F, US). SACATEPEQUEZ: Near Antigua, 6000 ft., Kellerman 1510
(F, 2 sheets, US); near San Lucas, 2300 m., Williams & Molina
11833 (F, GH). SAN MARCOS: southwest of San Marcos, 8000 ft.,
Morley 725 (F, GH, US). SOLOLÁ: Volcán Atitlán, Beaman 4091 (GH);
Volcán Santa Clara, 2100-3000 m., Steyermark 46991 (F). JUNCTION
OF HUEHUETENANGO, TOTONICAPÁN AND QUEZALTENANGO: Sierra
Madre mountains, 2400-2600 m., Williams, Molina & Williams 22608
(F). LOCALITY NOT DEFINITE: Fosberg 19055 (coL, Us). MEXICO.
CHIAPAS: Puerto Viento, Rayón, Breedlove 9003 (BM); Mt. Pasitar,
near San Cristobal, Matuda 461 (F, MEXU, MO, US); Mt. Tacana,
1000-2000 m., Matuda 2417 (A4, F, MEXU) ; near San Cristobal de las
252 Rhodora [Vol. 74
Casas, above 7000 ft., Pfeifer, Kremer & Abendroth 2501 (CONN,
2 sheets); Rio Prospero, Seler 2269 (GH, US). GUERRERO: Hinton
14201, 15398 (GH, US); Distrito Mina, Petlacala, Ynes Mexia 8972
(F, GH, MO). JALISCO: Rich canyons, mountains near Lake Chapala,
Pringle 2437 (BM, F, GH, M, MEXU, MO, US). MEXICO: ravines near
Ozumba, 8000 ft., Pringle 9713 (CU, F, GH, MEXU, MO, US); Pringle
11971 (CU, F, GH, US); along brooks, Amecameca, Purpus 1768 (F, GH,
MO, US); Ozumba, Purpus 3057 (BM, E, F, GH, MO, US). MICHOACAN:
San Juan Viejo, Beaman 4393 (GH); near Paracutin, along road
into Angahuan, 7500 ft., Bratz M-806 (MEXU); Tancitaro, Uruapan,
Hinton 15448 (GH, US); 2 km. N. & NE. of Puentacillar, Iltis,
Koeppen & Iltis 387 (BM); above Tancitaro, 7000 ft., Leavenworth
270 (F, GH, MO). MORELOS: Jojutla a de Juárez, D. de Juárez,
Salazar s. n., August 8 1912 (MEXU, 2 sheets). SINALOA: Along
Mex. #40, between Durango and Mazatlan, steep mountain sides,
Pfeifer & Skog 3073 (CONN). PANAMA. Volcán de Chiriqui Potrero
Muleto, Boquete district, 10400 ft., Davidson 1013 (A, F, US); be-
tween El Hato and Cerro Punta, Llanos, Ebinger 800 (Mo); Petrero
Muleto to summit de Chiriqui, Woodson & Schery 416 (GH, US).
PERU. APURIMAC: Prov. Abancay, arriba Abancay, 3300-3500 m.,
Ferreyra 9801 (USM); Abancay, Vargas 1954 (CUZ) ; between Huan-
carama and Cochacaya, 3400 m., West 3761 (GH, MO). AYACUCHO:
Aina, between Huanta and Rio Apurimac, 750-1000 m., Killip &
Smith 23119 (F, US). cuzco: Machu-Piechu, 7600 ft., Gourlay 87
(E); Machu-Picchu, 2200 m., Skog 1144, 1146, 1147 (CONN); Machu-
Picchu, 6000 ft., Stafford 1053 (BM, F); Urubamba, 2000 m., Vargas
1871 (cuz, GH); Pillahuata, cerro de cusilluyoc, 3000-3300 m., Pen-
nell 14131 (F, GH, US); Prov. Quispicanchis, Dist. Marcapata, 3000
m., Vargas 9701 (^, F, MO); Ollantaytambo, about 3000 m., Cook &
Gilbert 607 (^, F, GH, US, 2 sheets). HUANCAVELICA: east of Surcu-
bamba, Prov. Tayacaja, 2500 m., Stork & Horton 10343 (A, F, MO).
HUANUCO: Prov. Ambo, cerca a San Rafael, 2600 m., Ferreyra 1961
(USM, US); Pampayacu to Huanuco, Kanehira 230 (A, F); Chaglia,
Woytkowski 5293 (F, MO). JUNIN: east of Huasahuasi, 2400-2500
m., Hutchinson 1115 (BH, F, GH, M, MO, US, USM). LA LIBERTAD:
Prov. Otuzco, cerca a Usquil, 3000-3100 m., Ferreyra 7668 (US,
USM). PUNO: Prov. Carabaya, Ollachea, 2700 m., Vargas 6918 (cuz).
LOCALITY NOT DEFINITE: Jameson 577 (E); “Perou,” J. Jussieu s. n.
(P-JU); Ruiz & Pavon s. n. (BM, MO). VENEZUELA. MÉRIDA:
Mérida, Alston 6673 (BM, US); Plaza de Mucuqui, 2200 m., Bernardi
189 (MER, VEN); northeast cf Mérida along road to Valencia, 1850
m., Breteler 3300 (CONN, IFLA, 3 sheets, US); San Rafael, 3185 m.,
Gehriger 9 (^, GH, MO, Us); Distrito Campo Elías, San José de
Acequias, 2400 m., López-Palacios 774 (CONN, MER); El Cucharito
near Tabay Village, 1800 m., Skog 1247 (CONN); between Tabay and
1972] Coriaria — Skog 253
Mérida, along Rio Chama, 1820 m., Steyermark 55865 (F). TACHIRA:
Distrito Uribante, San José, antes de la poblacion de Pregonero,
1440 m., Marcano-Berti 1680 (CONN, MER). TRUJILLO: Jajó, Burkart
16686 (VEN).
ACKNOWLEDGEMENTS
The author gratefully acknowledges the aid and advice
of Dr. H. W. Pfeifer during the work, Dr. W. J. Dress for
reading the manuscript and support from the National Sci-
ence Foundation Grant to the University of Connecticut for
graduate student travel, which enabled the author to visit
herbaria and collect in Central and South America. Also,
the author is grateful to the curators of the herbaria cited
for allowing him to examine material of Coriaria. The
abbreviations for herbaria are from Index Herbariorum,
ed. 5 by Lanjouw and Stafleu, except for the following,
which are not included: CRU — The herbarium of the Uni-
versity of Costa Rica, San José, Costa Rica; csa — The
personal herbarium of Professor Hugo Gunckel L. of San-
tiago, Chile; IFLA — The herbarium of the Instituto Forestal
Latino-Americano, Mérida, Venezuela.
LITERATURE CITED
BENTHAM, G. & J. D. HOOKER. 1862. Genera Plantarum .
1: 429.
CANDOLLE, A. P. DE. 1816. Essae sur les propriétes medicales des
plantes, ed. 2. p. 350-351.
CRONQUIST, A. 1968. The Evolution and Classification of Flowering
Plants. 396 p.
HALLIER, H. 1905. Provisional scheme of the natural (phylo-
genetic) system of Flowering Plants. New Phytologist 4: 151-
162.
HUTCHINSON, J. 1964. The Genera of Flowering Plants. 1: 172-
173.
LAWRENCE, G. H. M. 1951. Taxonomy of Vascular Plants. 823 p.
MaxriMOWICZ, C. J. 1881. De Coriaria, Ilice et Monochasmate,
hujusque generibus proxime affinibus Bungea et Cymbaria.
Mém. Acad. Imp. Sci. Saint Pétersbourg, sér. 7. 29(3): 1-13.
Russow, E. 1884. Uber den anatomischen Bau der Laubsprosse der
Coriarieen. Sitzungsber. Naturf.-Ges. Univ. Dorpat 6: 88.
L. H. BAILEY HORTORIUM
CORNELL UNIVERSITY
ITHACA, NEW YORK
LIST OF THE CLADONIAE OF
SOUTHEASTERN MASSACHUSETTS,
WITH SPECIAL REFERENCE TO THE COLLECTION
OF CHARLES ALBERT ROBBINS
LAWRENCE B. MISH
After Charles Albert Robbins died in 1930, S. F. Blake
(1933) wrote of the long and methodical collecting and
identifying done by this quiet man from Wareham, Massa-
chusetts, compiling the largest private collection of the
genus Cladonia up to that time. Robbins’ collection for the
small town on Cape Cod was more extensive than the col-
lection of Evans (1930) for the whole state of Connecticut.
For almost forty years, Robbins’ Cladonias have been housed
in the Farlow Herbarium and his approximately 46 species
are unknown to the world except for those discussed in his
five papers, published between 1924 and 1929.
While surveying the plants of Southeastern Massachusetts
during a sabbatical leave in 1968-69, it was found that some
of the plants in the Robbins collection were not included in
the work of Thomson (1967). It would seem wise, there-
fore, to publish a list at this time. The numbers appearing
in front of each taxon correspond to those of Thomson
(1967). Where no connection could be made with the taxa
in Thomson’s work, Zahlbruckner (1922-1940) was referred
to. The letters following the taxa indicate the collection
locality, viz.:
B — Bournedale NF — North Falmouth
pr — Bridgewater o | — Orleans
pre — Brewster P — Plymouth
c — Carver R — Rockland
p — Duxbury S —Stoughton
E — Eastham sh — Sharon
H — Hingham w — Wareham
M — Marshfield wf — Wellfleet
Na — Nantucket wH — Woods Hole
254
1972] Cladoniae — Mish
255
Those citations marked with an asterisk (*) are for
plants not collected by Mr. Robbins.
SCIENTIFIC NAME
1. C. papillaria (Ehrh.) Hoffm.
f. molariformis
(Hoffm.) Schaer.
f. papillaria Thoms.
f. stipata Flk.
5. C. coccifera (L.) Willd.
v. frondescens (Nyl.) Vain.
v. stemmatina Ach.
9. C. deformis (L.) Hoffm.
f. crenulata Ach.
11. C. pleurota (Flk.) Schaer.
12. C. incrassata Flk.
f. clavata Robb.
f. squamulosa (Robb.) Evans
14. C. floerkeana (Fr.) Flk.
v. intermedia Hepp.
f. fastigiata Laur.
v. intermedia
f. peritheta Robb.
f. intermedia (Hepp.) Thoms.
f. carcata (Ach.) Thoms.
f. floerkeana Thoms.
f. squamosissima T. Fr.
15. C. bacillaris (Ach.) Nyl.
f. attenuata Robb.
f. peritheta (Wallr.) Zahlbr.
f. fruticaulescens Vain.
f. obtusa Wallr.
f. bacillaris Thoms.
16. C. cristatella Tuck.
f. aurantiaca Robb.
PLACE
B
*
Un
gam zzz
nm
p
a
um ue dU m Mo EUIS (S
Uu
TEE
ive]
W,S
Widespread
WwW
256
21.
22.
23.
25.
30.
əl.
32.
36.
3T.
39.
f.
rh Ph bh Th kh rh Ph
Rhodora
lepidifera (Vain.) Robb.
pleurocarpa Robb.
squamulosa Robb.
squamosissima Robb.
cristatella Thoms.
ochrocarpa Tuck.
ramosa Tuck.
vestita Tuck.
C. macilenta Hoffm.
f.
macilenta Thoms.
C. didyma (Fée) Vain.
var. muscigena (Eschw.) Vain.
C. leporina Fr.
C. piedmontensis Merr.
f.
lepidifera (Vain.) Robb.
phyllocoma Robb.
obconica Robb.
piedmontensis Thoms.
intermedia Robb.
squamossisima Robb.
C. strepsilis (Ach.) Vain.
f.
f.
f.
f.
abortiva Robb.
strepsilis Thoms.
coralloidea (Ach.) Vain.
glabrata Vain.
C. robbinsii Evans
f. squamulosa (Evans) Evans
C. capitata (Michx.) Spreng.
f.
capitata, Thoms.
C. clavulifera Vain.
C. apodocarpa Robb.
C. polycarpoides Nyl.
f.
epiphylla (Robb.) Thoms.
æ
TEE
~
TELE
Z
fo
~
zzz" Q2QAQ9009 Ss
EI
æ
TERREN
[Vol. 74
Ua
1972]
40.
46.
4T.
49.
56.
5T.
58.
Cladoniae — Mish
f. squamulosa..(Robb.) Thoms.
f. polycarpoides Thoms.
C. cariosa (Ach.) Spreng.
f. squamulosa
(Müll. Arg.) Vain.
C. mateocyatha Robb.
f. squamulata Robb.
C. gracilis (L.) Willd.
var. dilitata (Hoffm.) Vain.
f. dilacerata (Flk.) Vain.
var. elongata (Jacq.)
Fr. f. elongata Thoms.
C. verticillata (Hoffm.) Schaer.
f. pallida Robb.
var. cervicornus Flk.
f. phyllophora Flk.
phyllocephala Flot.
aggregata (Del.) Oliv.
apoticta (Ach.) Vain.
phyllocephala (Flot.) Oliv.
f. verticillata Thoms.
C. simulata Robb.
C. pityrea Flk.) Fr.
var. pityrea Thoms.
pityrea Thoms,
squamulifera Vain.
subacuta Vain.
cladomorpha (Flk.) Vain.
scyphifera (Del.) Vain.
C. pyxidata (L.) Hoffm.
var. pyxidata Thoms.
f. centralis Oliv.
f. marginalis Hoffm.
f. prolifera Wallr.
257
33
* =
a: MEE.
a a
33333
EEEE:
og
5
m
e
len
258 Rhodora [Vol. 74
f. lophyra (Ach.) Korb.
f. homodactyla Wallr.
59. C. chlorophaea (Flk.) Spreng.
f. costata (Flk.) Arn.
f. simplex (Hoffm.) Arn.
f. prolifera (Wallr.) Arn.
60. C. grayi Merr.
peritheta Evans
cyathiformis Sandst.
centralis Evans
prolifera Sandst.
squamulosa Sandst.
pallida Robb.
epiphylaa Robb.
foliosa Robb.
. clavata Robb.
64. C. fimbrriata (L). Fr.
f. stenoscypha Evans
f. conista (Ach.) Robb.
f. pycnotheliza (Nyl.) Vain.
f. simplex (Weiss) Vain.
66. C. nemoxyna (Ach.) Nyl.
f. ambigua Wallr.
f. phyllocephala Arn.
f. scyphifera Robb.
f. nemoxyna Thoms.
s444 xz
a a
Ph rh rh rh rh r5 rh rh rh
2zzzzzzz22
a
wees
w
e
Ui
68. C. coniocraea (Flk.) Spreng.
f. ceratodes (Flk.) Thoms.
f. phyllostrota (Flk.) Vain.
f. truncata (Flk.)
DT. & Sarnth.
f. actinota FIk.
f. pycnotheliza (Nyl) Vain.
f. peritheta Robb.
72. C. turgida (Ehrh.) Hoffm.
f. conspicua (Schaer.) Nyl.
TEREE
vn
2 4444
*
ise]
e$
1972]
Cladoniae — Mish
76. C. furcata (Huds.) Schrad.
var. furcata Fk.
TT.
79.
81.
82.
83.
86.
88.
C. scabriuscula (Del. in Duby) Nyl.
a
DU DEP led
fissa (Flk.) Aigr.
furcata Thoms.
subclausa (Sandst.) Evans
rigidula Mass.
subulata (Ach.) Vain.
. corymbosa (Ach.) Vain.
var. palamaea (Ach.) Vain.
var. paradoxa Vain.
var. pinnata (Flk.) Vain.
f.
f:
P^ Ph rh Ph Ph
f. recurva (Hoffm.) Sandst.
foliolosa (Del.) Vain.
turgida Scriba ex Sandst.
elegans Robb.
squamulosa Robb.
. adspersa Flk.
farinacea (Vain.) Sandst.
scabriuscula Thoms.
. parasitica (Hoffm.) Hoffm.
f.
f.
f.
congesta Robb.
stertlia Robb.
parasitica Thoms.
floridana Vain.
beawmontii (Tuck.) Vain.
caespiticia (Pers.) Flk.
f.
epiphylla (Arn.) Sandst.
multiformis Merr.
carassensis Vain.
E
subregularis Vain.
f. digressa Vain.
f.
f. spectabilis Robb.
regularis Vain.
*3 4344
2443 43445
-
243223 3 Z3 4 8
JEEEEELTILLLE
-
æ
is
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259
260 Rhodora [Vol. 74
f. obliqua Robb.
m. spectabilis Robb.
f. subregularis Vain.
m. spectabilis Robb.
f. subirregularis Robb.
z
89. C. squamosa (Scop.) Hoffm.
f. pityrea Arn.
f. fascicularis (Del.) Nyl.
f. denticollis (Hoffm.) Vain.
f. squamosa Thoms.
var. laevicorticata Sandst.
f. degenerascens Anders.
f. laevicorticata
(Sandst.) Evans
f. carneopallida Sandst.
var. muricella (Del.) Vain.
f. sarmentosa (Tayl.)
Müll Arg.
f. ventricosa (Del.) Oliv.
var. phyllocoma Robb.
f. muricella Sandst.
pseudocrispata Sandst.
rigida (Del.) Nyl.
subesquamosa Nyl.
subtrachynella Vain.
f. turfacea (Arn.) Vain.
90. C. subsquamosa (Nyl.) Vain.
f. subrigida Robb.
f. luxurians (Nyl.) Vain.
f. degenerata Robb.
f. sublevis Robb.
f. subrigida Robb.
f. scabrida Robb.
91. C. crispata (Ach.) Flot.
f. subascypha Savicz
var. dilacerata
(Schaer.) Malbr. W
zzzzz xz
a
z
zz
zzz
Ph rh rh rh
4 tt ot
2a 355
3434334344
=
1972] Cladoniae — Mish 261
f. elegans (Del.) Vain.
f. infundibulifera
(Schaer.) Thoms.
var. divulsa (Del.) Arn.
f. crispata Thoms.
var. virgata Ach.
f. kairamoi Vain.
92. C. cenotea (Ach.) Schaer.
f. exaltata Nyl.
f. cenotea Thoms.
93. C. glauca Flk.
97. C. uncialis (L.) Wigg. f. humilior Fr.
44 449 3 444 4
J
f. turgescans (Del. E. Fries w
f. obtusata (Ach.) Nyl. W, E
f. uncialis Thoms. Ww
f. setigera Anders S W
f. soraligera (Robb.) Sandst. W
f. subobtusata (Coem.) Arn. Ww
101. C. boryi Tuck. W, P, D, WH,
E, Bre, Q
f. prolifera Robb. Ww
f. lacunosa (Bory) Evans W, S, D, Sh
102. C. caroliniana Schwein, ex Tuck. W
107. C. alpestris (L.) Rabh. W, Br, E H
108. C. impexa Harm. f. prolifera Robb. w
f. impexa Thoms. Ww
f. subpellucida Harm. W
109. C. terrae-novae Ahti W
110. C. tenuis (Flk.) Harm. W: Ss, R
f. prostrata Robb. Ww
f. prolifera Robb. Ww
111. C. subtenuis ( Abb.) Evans W, S, E
262 .. Rhodora [Vol. 74
113. C. mitis Sandst.
f. pallida Robb.
tenuis Sandst.
divaricata Sandst.
prolifera Sandst.
soralifera Sandst.
f. prostrata Sandst.
115. C. sylvatica (L.) Hofim.
f. sphagnoides (Flk.) Oliv.
f. congesta Robb.
m. fissa Flk.
f. gracilior Vain.
f. arbuscula Wallr.
f. decumbens Flk.
f. scabrida Robb.
m. sorediata Sandst.
f. pygmaea Sandst.
116. C. rangiferina (L.) Wigg. Widespread
f. incrassata (Schaer.) Anders W
f. tenuior (Del.) Vain. wW
R, Nonquit
-
rh rh rh Ph
Z
"i
4444446
[^ Uu.
Z
"rj
aL
zZ
£5
44444444248
Ua
Not included in Thomson's book:
C. poculifera Robb.
f. simplex Robb.
C. pseudopyxidata Robb.
f. clavata Robb.
f. Simplex Robb.
f. squamulosa Robb.
f. lateralis Robb.
f. prolifera Robb.
C. pyxioides. (Wallr.) Britz. f. simplex Britz.
== 222222 22 z
C. spumosa Flk.
99
1972] Cladoniae — Mish 26
LITERATURE CITED
BLAKE, S. F. 1933. Charles Albert Robbins, 1874-1930. Rhodora
35: 140-145.
Evans, A. W. 1930. The Cladoniae of Connecticut. Trans. Conn.
Acad. Arts & Sci. 30: 357-510.
THOMSON, J. W. 1967. The Lichen Genus Cladonia in North Amer-
ica. U. Toronto Press.
ZAHLBRUCKNER, A. 1922-1940. Catalogus Lichenum universalis,
vols. 1-10. Leipzig.
DEPARTMENT OF BIOLOGY
BRIDGEWATER STATE COLLEGE
BRIDGEWATER, MASSACHUSETTS 02324
ON THE MARINE ALGAE OF KENT ISLAND,
BAY OF FUNDY
KENNETH L. KOETZNER and R. D. Woop
The extreme tidal amplitudes found in the Bay of Fundy
are world famous, and biologists are intrigued by the rela-
tionship between organisms and altitudes of such extreme
tides. Whereas much of the benthic region of the upper
reaches of the bay consists of clean-swept mud, there are
scattered rock outcrops along the bay where attached algae
can be studied. One such spot is Kent Island, New Bruns-
wick, a fairly small island in the mouth of the Bay of
Fundy. It lies just southeast of Grand Manan Island and
is one of the Three Island group, located at approximately
44°35’ N lat and 66°46’W long.
The junior author (rdw) visited the Bowdoin College
Biological Station on Kent Island from June 28 to July 3,
1957, to study the marine algae. The results were compiled,
but publication was deferred at the request of a colleague.
Now, after nearly a decade during which no apparent con-
flict of interest was apparent, further delay seemed un-
justified; and, at the request of other colleagues, the report
is now issued.
During the five days on the island, collecting was done
around the entire coastline; but, as work was sharply
limited by rains, effort was concentrated in four areas: —
on the north shore (N), a boulder-strewn beach ; — on the
east shore (E), a cobble beach with rocks and tide pools at
the low water level; — on the south shore (S), a rocky cliff
region; — on the west shore in the Basin (B), a cove ex-
posed as mud flats at extreme low tides. Specimens, both
dried and fluid (5% of neutral formalin in seawater), were
prepared and are on deposit at the University of Rhode
Island. Examination of the specimens and compilation of
the data was the primary responsibility of the senior author
(klk) ; while the field work, observations, and final prepa-
ration of manuscript were the responsibility of the junior
264
1972] Marine Algae — Koetzner and Wood 265
author (rdw). Classification and nomenclature follow Tay-
lor (1957).
Perhaps the first report of the algae of the island was
by Bowers (1942) who identified 35 species, 15 of which
were not noted during the present study. Other early re-
ports of algae from the region but not from the island itself
are those by Hay (1882) of 33 species and Hay and McKay
(1886) for the Bay of Fundy. In addition, there are records
by Eaton (1873) for Maine, Klugh (1917) for New Bruns-
wick, Roscoe (1931) for Nova Scotia, Bell and MacFarlane
(1933a, b) of 66 species for the Bay of Fundy, Humm
(1950) for Newfoundland, and Taylor (1937; 1957) who
treated in detail the algae of the entire northeastern coast
of North America, including the Bay of Fundy.
The junior writer (rdw) is indebted to Dr. James Moul-
ton of Bowdoin College for the idea and arrangements for
the visit; Dr. Charles Huntington, Director of the Bowdoin
College Biological Station, and his wife, for their kindness
and hospitality; and to that fabulous down east captain
who “punched” his way through the impenetrable fog and
found Kent Island “by ear.”
(E = east shore; B = Basin; N — north shore;
S — south shore; see text for descriptions)
Chlorophyceae
Ulotrichaceae
l. Ulothrix flacca (Dillw.) Thuret. E, S. Abundantly
matted on rocks at + 11 ft.
Ulvaceae
2. Enteromorpha clathrata (Roth) J. Ag. — N, E, S, B,
N of Basin.
Variable in habit. Not common.
(2a. E. compressa (L.) Grev. — (Bowers, 1942) ).
3. E. erecta (Lyngb.) J. Ag. — E. Rare.
4. E. intestinalis (L.) Link — N, E, S, B, N of Basin.
On rocks and stones. Common.
9. E. linza (L.) J. Ag. — E. Rare.
(5a. E. marginata J. Ag. — (Bowers, 1942) ),
266 Rhodora [Vol. 74
6. E. minima Nag. — E. Attached to stones.
7. E. prolifera (Müll.) J. Ag. — E, B. Masses attached
to rocks in lower littoral. (also Bowers, 1942).
8. Monostroma fuscum (Post. & Rupr.) Witt. (f. blytii
(Aresch.) Collins — N. In wash, rare.
9. Ulva lactuca L.—N, E, S. Lower littoral. (also
Bowers, 1942).
Prasiolaceae
10. Prasiola stipitata Suhr — N, E. Found high in spray
zone.
Cladophoraceae
11. Chaetomorpha melagonium (Web. & Mohr) Kütz. —
Found in midlittoral zone and lower tide pools.
(11a. C. area (Dilw.) Kütz. — Bowers, 1942).
19. Cladophora gracilis (Griff. ex Harv.) Kütz. — N, E,
S, B. Abundant throughout mid and lower littoral
zone.
18. C.ruprestris (L.) Kütz. — N, B. Uncommon, in lower
littoral.
14. C. refracta (Roth) Kütz. — N, B. Uncommon. (also
Bowers, 1942).
15. Rhizoclonium tortuosum Kütz. — N. With Clado-
phora sp. in midlittoral tidepools.
16. Spongomorpha arcta (Dillw.) Kütz.— E, S. Form-
ing thick, stiff tufts up to 6 in. in diameter, abun-
dant in lower littoral zone.
17. S. spinescens Kiitz. — S. Uncommon, in association
with S. arcta. At + 3 ft.
Phaeophyceae
Ectocarpaceae
18. Ectocarpus confervoides (Roth) Le Jol.—N, B.
Epiphytic on coarser brown algae.
(18a. Giffordia granulosa (J. E. Smith Hamel — (Bower,
1942, as E. granulosus) ).
19. Pylaiella littoralis (L.) Kjellm. — B. On rocks, Fucus
and Ascophyllum.
1972] Marine Algae — Koetzner and Wood 267
Ralfsiaceae
20. Ralfsia fungiformis (Gunn.) Setch. et Gard. —N, E,
S. Forming crusts several inches in diameter on
rock, upper littoral zone.
Elachisteaceae
21. Elachistea fucicola (Vell.) Aresch. — N, E. S. At-
tached in dense tufts to Ascophyllum.
Chordariaceae
22. Chordaria flagelliformis (Müll) C. Ag.—E, B.
Found awash, often mixed with Dumontia.
23. Sphaerotrichia divaricata (C. Ag.) Kylin. — N of
Basin. Occasional, + 9 to + 13 feet.
Desmarestiaceae
24. Desmarestia aculeata (L.) Lamour. — N, S, B. Occa-
sional, found in lower littoral.
25. D. viridis (Müll.) Lamour. — E. On shells of lower
littoral, + 4 ft.
Punctariaceae
(25a. Asperococcus echinatus (Mert.) Grev.— (Bowers,
1942) ).
26. Scytosiphon lomentaria (Lyngb.) C. Ag. —S. At-
tached to rocks, + 2 ft.
27. Punctaria latifolia Grev. — E, S, B. Common at or
below low water.
Dictyosiphonaceae
28. Dictyosiphon foeniculaceus (Huds.) Grev. — E, B.
Common near low water.
Laminariaceae
29. Agarum cribrosum (Mert.) Bory — N. Occasional,
in sublittoral region. (also Bowers, 1942).
30. Alaria esculenta (L.) Grev. — E, N, S. Fairly com-
mon near or below low water. (also Bowers, 1942).
31. Laminaria agardhii Kjellm. — N, E, S. Common,
found with Alaria. (also Bowers, 1942).
268 Rhodora [Vol. 74
32. L. digitata (L.) Lamour. — E, S. Common below
low water. (also Bowers, 1942).
(32a. L. intermedia Fosl. — (Bowers, 1942) ).
33. L. longicruris De La Pyl.— B. Found attached to
weir. (also Bowers, 1942).
34. L. platymeris De La Pyl. — E, B. In tide pools on
beach at + 2 ft.
(34a. L. stenophylla (Kütz.) J. Ag. — (Bowers, 1942) ; —
L. digitata fide Taylor (1957: 185). ]
35. L. sp. —B. A form similar to L. agardhii but with
flat stipe, 0.5-2 em. thick; blade thinner and wider.
Fucaceae
36. Ascophyllum nodosum (L.) Le Jol. — N, E, S, B.
Common in the entire littoral zone in most areas.
(also Bowers, 1942).
(36a. A. mackaii (Turn.) Holms et Batt. — (Bowers,
1942) ).
37. Fucus edentatus De La Pyl. — N, E. Form with ex-
tremely flattened and broad receptacles. (also
Bowers, 1942).
38. F. evanescens C. Ag., prox. — S of Basin. Occasional
on rocks in sand. (also Bowers, 1942).
39. F. filiformis Gmelin — N, E. Common in high tide
pools. (also Bowers, 1942).
40. F. spiralis L. — N, E, S, B. Common in all areas of
the upper littoral zone, often mingling with strand
vegetation.
41. F. vesiculosus L. — N, E, S. B. Common throughout
the littoral zone (also Bowers, 1942).
Rhodophyceae
Bangiaceae
49. Porphyra umbilicalis (L.) J. Ag. — E, S. Occasional
in lower littoral. (also Bowers, 1942).
Acrochaetiaceae
43. ?Aerochaetium daviesii (Dillw.) Nàg. — N.
(43a. Kylinia secundata | (Lyngb.) Papenf. — (Bowers,
1942 as A. secundatum) ).
1972]
44,
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
Marine Algae — Koetzner and Wood 269
Dumontiaceae
Dumontia incrassata (Müll.) Lamour. — E, S. At-
tached to rock in tide pocls, commonly with Chor-
daria.
Squamariaceae
Hildenbrandia prototypus Nardo — E. Common in
tide pools in the upper littoral.
Corallinaceae
Corallina officinalis L.— E, S. Abundant at or
slightly above low water level.
Kallymeniaceae
Euthora cristata (L.) J. Ag. —S, W. Found awash.
Rhodophyllidaceae
Cystoclonium purpureum (Huds.) Batt. — N, S. On
rocks at low water, with Porphyra. (also Bowers,
1942).
Phyllophoraceae
Ahnfeltia plicata (Huds.) Fries — E, S. Common in
the mid to lower littoral zone.
Phyllophora brodiaei (Turn.) J. Ag. — E. S. Occa-
sional.
Gigartinaceae
Gigartina stellata (Stackh.) Batt. — E, S. Common to
abundant in the mid to lower littoral.
Rhodymeniaceae
Halosaccion ramentaceum (L.) J. Ag. —N, S. Occa-
sional.
Rhodymenia palmata (L.) Grev. — E. Common at
low water, often attaining length of 1.5 m. (also
Bowers, 1942).
Ceramiaceae
Ceramium rubrum (Huds.) C. Ag. — S. Uncommon.
Ptilota serrata Kütz. — S, B. In wash.
270 Rhodora [Vol. 74
. Delesseriaceae
56. Phycodrys rubens (Huds.) Batt. — N, S, W. Attached
in the lower littoral.
Rhodomelaceae
57. Polysiphonia harveyi Bailey — N. Uncommon.
58. P. lanosa (L.) Tandy — E, S. Epiphytic on Asco-
phyllum, forming dense masses.
59. P. nigrescens (Huds.) Grev. — E, S. Forming dense
growths on rocks in lower littoral.
60. P. urceolata (Lightf.) Grev. — N. Uncommon.
61. Rhodomela confervoides (Huds.) Silva — N, S. At-
tached to rocks at lower water.
LITERATURE CITED
BELL, H. P. and MACFARLANE, C. I. 1933a. The marine algae of
the Maritime Provinces of Canada, I. List of species with their
distribution and prevalence. Can. Jour. Res. 9: 265-279.
BELL, H. P. and MACFARLANE, C. I. 1933b. The marine algae of the
Maritime Provinces of Canada, II. A study of their ecology.
Ibid. 9: 280-293.
Bowers, C. H. 1942. Algae of Kent Island. Bowdoin Sci. St. Bull.
8: 36-37.
EATON, D. C. 1873. List of marine algae collected near Eastport,
Maine, in August and September, 1873, in connection with the
work of the U. S. Fish Commission under Prof. S. F. Baird.
Trans. Conn. Acad. Arts & Sci. 2: 343-350.
Hav, G. U. 1882. Marine algae of the Bay of Fundy. Bull. New
Brunswick Nat. Hist. Soc. 1: 62-68.
Hay, G. U. and MacKay, A. H. 1886. Marine algae of the Bay of
Fundy. Ibid. 1: 32-33.
HuMM, H. J. 1950. Notes on the marine algae of Newfoundland.
Jour. Tenn. Acad Sci. 25: 229.
KLUGH, M. V. 1917. Marine algae of the Passamaquoddy region,
New Brunswick. Contrib. Can. Biol, Suppl. 6th Ann. Rep.
Dept. Naval Serv., Fisheries Bur., pp 79-85.
1972] Marine Algae — Koetzner and Wood 271
Roscoe, M. V. 1931. The algae of St. Paul Island. Rhodora 33:
127-131.
TAYLOR, W. R. 1937. Marine algae of the northeastern coast of
North America. Univ. Michigan Press, Ann Arbor. 427 p.
TAYLOR, W. R. 1957. Ibid., 2nd ed. 509 p.
NEW YORK STATE DEPARTMENT OF
ENVIRONMENTAL CONSERVATION,
4175 VETERANS MEMORIAL HIGHWAY,
RONKONKOMA, N.Y. 11779
and
DEPARTMENT OF BOTANY, UNIVERSITY OF RHODE ISLAND,
KINGSTON, R.I. 02881
A WHITE-FLOWERED FORM OF IRIS LACUSTRIS
FROM ONTARIO. In an open dune hollow within fifty
yards of the shoreline north of McNab Point in Bruce
County, Ontario, a white-flowered form of the Dwarf Lake
Iris, Iris lacustris Nutt., was discovered amongst a few
hundred blooming plants of the typical form.
It is proposed that this new form be named Iris lacustris
Nutt. forma albiflora Cruise & Catling, forma nova; differt
forma typica quad flores albos non caeruleos habet.
Presently known only from the type collection.
Type (Specimen and Color Photograph): Associating
with many of the typical form and with Oryzopsis asperi-
folia and O. pungens, in moist sand in first dune hollow
behind bay north of McNab Point (44° 28’ N., 81° 24’ W.),
U. T. M. G. zone 17, 692243, ca. 2 miles south of South-
ampton, Saugeen Township, Bruce County, Ontario. 22
May, 1971, P. M. Catling & S. M. McKay, (TRT 168074).
J. E. CRUISE & P. M. CATLING
DEPARTMENT OF BOTANY
UNIVERSITY OF TORONTO
TORONTO 181, ONTARIO, CANADA
NEOMIRANDEA ALLENII, A NEW EPIPHYTIC
COMPOSITE OF THE AMERICAN RAIN FOREST
R. M. KING and H. ROBINSON
Members of the Eupatorian genus Neomirandea, ranging
from southern Mexico to Eucador, might well be called the
Orchids of the Asteraceae. These usually epiphytic plants
with rather succulent stems and showy inflorescences have
only recently been recognized as a distinct genus (King &
Robinson, 1970). Collectors have often commented, some-
times extensively, on the unusual habit of the plants, and
Professor B. L. Robinson (1918) in his description of a
Colombian species, N. sciaphila (B. L. Robinson) R. M. King
& H. Robinson, mentioned the horticultural potential of the
plant. It is rare that labels do not mention the vining habit,
epiphytic nature, or showy reddish-purple flowers. We
take occasion here to call attention to some of the interest-
ing problems of this unique group and to describe a four-
teenth species from Panama and Colombia.
The ecology of the genus Neomirandea presents a par-
ticularly important area for future investigation. Most of
the species, even some of the largest ones, are referred to
as epiphytic. The habit prompted the name Eupatorium
parasiticum Klatt, and a collection of the type species of
the genus, N. araliaefolia (Lessing) R. M. King & H.
Robinson was described by Skutch, “shrub or small tree
with branches 25 ft. long and 6 inches in diameter.
Epiphytic on an oak tree, 8 ft. above ground, the roots
clasping the trunk in the manner of those of a strangling
fig, concrescent where touching each other. A single large
root descends along the trunk to the ground. Flowers
white, rare. More about this interesting plant in my
journal under March 4, 1933." The new species described
here is cited by Allen as “fleshy branching epiphytic shrub
114 meters. In tops of tallest trees." The question remains,
what of the few remaining members of this well marked
genus not known to be epiphytic? But for their relation-
ship, these species might never be examined properly to see
272
1972] Neomirandea — King and Robinson 273
if their substrate is in any way specialized, possibly old
stumps or rotten logs. Experiments with the seeds of these
plants are also essential to determine what adaptation they
have to their substrate.
Among the Eupatorieae, Neomirandea is one of the few
genera of particular interest cytologically. The chromosome
picture in the Eupatorieae is for the most part rather
simple. Most groups show a haploid number of 10 chromo-
somes or multiples of that number. Fleischmannia normally
has n — 10, and a few species with » — 4 (Baker, 1967).
A whole related complex of genera which we call Agera-
tinoid (Ageratina, Oxylobus, etc.) shows base numbers of
x = 17, and x = 16. Thus far, two species of Neomirandea
have been counted, (Turner & King, 1964), N. angularis
(B. L. Robinson) R. M. King & H. Robinson, n = 25 and
N. costaricensis R. M. King & H. Robinson, » — 17 as
Eupatorium aff. eximium. 'These two counts suggest some
cytological complexity in this genus which is in some ways
between the Ageratinoid complex and other members of
the Eupatorieae.
The new species is represented by two collections which
show a range from Central Panama to the foothills of the
northern Andes in Colombia. The characters of the species
are as follows:
Neomirandea allenii R. M. King & H. Robinson, sp. nov.
Frutex epiphyticus carnosus. Folia opposita majuscula
subglabra integra, petiolo perbreve; laminae late ellipticae
7-9 cm. longae, 4-6 cm. latae, apice anguste obtusatae, nervis
lateralibus prominulis pinnatis. Involucri squamae ca. 12
subaequilongae late lanceolatae extus parce pubescentes.
Flores ca. 10 in capitulo ca. 4.5 mm. longi; corollae anguste
infundibulares extus distincte glanduliferae, faucibus intus
glabris; styli non nodulosi; achaenae setiferae et glandi-
ferae inferne dense glanduliferae et setiferae; setae pappi
scabrae, cellulis apicalibus setarum nonnullarum valde
obtusis.
Fleshy branching epiphytic shrubs. Stems terete, min-
utely pubescent. Leaves opposite, petioled; petioles glabrous
274 Rhodora [Vol. 74
ca. 5 mm. long, blades fleshy, ovate, entire, glabrous on
both surfaces, up to 9 cm. long and up to 6 cm. wide, lateral
veins prominent. Inflorescence a corymbose panicle. Phyl-
laries ca. 12, subequal, broadly lanceolate, pubescent, in
2-3 series. Receptacle flat or slightly convex, glabrous.
Heads ca. 10 flowered, flowers purple or pink, ca. 4.5 mm.
long (excluding style branches) corollas narrowly funnel-
form, lobes about as long as wide, outer surface of corolla
with numerous short stalked glands, cells broadly oblong
with straight walls, inner surface of corolla glabrous;
anther appendages large, anther collars slender, composed
mainly of rectangular to quadrate cells, walls not orna-
mented. Style base not enlarged, glabrous, stylar append-
ages not enlarged, mamillose. Achenes prismatic, 4-5 ribbed,
with a few short stalked glands and uniseriate multiseptate
hairs, glands and hairs numerous at base and apex, car-
popodia distinct, cells quadrate, thin walled, pappus of ca.
30 scabrous setae ca. 5 mm. long, some setae with very
blunt apical cells.
PANAMA: PROVINCIA DE COCLE: Region north of El
Valle de Anton, alt. 1000 meters, August 21, 1946. Fleshy
branching epiphytic shrub, 115 m. In tops of tallest trees.
Leaves leathery. Flowers purple, showy. Paul H. Allen
3643 (Holotype NY). Additional collection: COLOMBIA:
ANTIOQUIA: Above Llano Grande, elevation 2000 m. Earl
L. Core 497 (us).
The new species by its lack of hairs inside the corolla,
and by the lack of an enlarged style base proves to be a
member of the subgenus Critoniopsis related to N. eximia
(B. L. Robinson) R. M. King & H. Robinson and N. scia-
phila (B. L. Robinson) R. M. King & H. Robinson. Among
the prominent distinctions of N. allenii are the larger leaves
with prominent secondary veins, the distinct glands on the
surface of the corolla, the hairs and glands near the base
of the achenes, and the frequent interspersed pappus setae
with very blunt apical cells.
1972] Neomirandea — King and Robinson 275
ACKNOWLEDGEMENT
This study was supported in part by the National Science
Foundation Grant GB-20502 to the senior author.
LITERATURE CITED
BAKER, H. G. 1967. The evolution of weedy taxa in the Eupatorium
microstemon species aggregate. Taxon 16: 293-300.
KING, R. M. & H. R. RoBINSON. 1970. Studies in the Eupatorieae
(Compositae). XXI. A new genus, Neomirandea. Phytologia
19(5): 305-310. ON
ROBINSON, B. L. 1918. A descriptive revision of the Colombian
Eupatoriums. Contr. Gray Herb. 55: 264-330.
TURNER, B. L. & R. M. KING. 1964. Chromosome numbers in the
Compositae. VIII. Mexican and Central American species.
Southw. Nat. 9: 27-39.
DEPARTMENT OF BOTANY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560
STUDIES IN THE GUTTIFERAE.
III. AN EVALUATION OF SOME PUTATIVE
SPONTANEOUS GARDEN HYBRIDS IN
HYPERICUM SECT. MYRIANDRA'
PRESTON ADAMS
For many years several species of Hypericum have been
in cultivation at the Arnold Arboretum, Jamaica Plain,
Massachusetts. Some of these are woody shrubs of Sect.
Myriandra native to eastern North America. Around 1909
Alfred Rehder, a long-time member of the staff, discovered
in the Arboretum garden plants of Hypericum which ap-
peared to him to be intermediate in many characteristics
between some of these native American species. He thought
that these “atypical” individuals were spontaneous hybrids
and, being interested in new and different plants for horti-
cultural purposes, gave them binomials. The specimens
which he collected are now in the herbarium of the Arnold
Arboretum.
During monographic studies of the woody specimens of
Hypericum in eastern North America (Adams, 1962), I
examined Rehder’s specimens. Having observed little evi-
dence of hybridization among these plants in nature, I
welcomed the opportunity to study putative examples of
spontaneous crossing under garden conditions. The present
contribution is an evaluation of the possible hybrid nature
of Rehder’s specimens.
Hypericum X Arnoldianum Rehder, Mitteil. Deutsch.
Dendrol Ges. 19: 253. 1910 [19112]. Rehder applied this
name to plants which he believed had originated from a
cross between H. lobocarpum Gattinger and H. galioides
Lam., with the latter being the maternal parent. During
1910 Rehder made several collections of this putative
‘Financial support for this study was provided by the Graduate
Council Research Fund of DePauw University and the Fernald Fund
for Field Study in Systematic Botany at Harvard University, estab-
lished by the late Mr. F. W. Hunnewell of Wellesley, Mass. I want
to thank Dr. Tod F. Stuessy of Ohio State University for helpful
comments on the manuscript.
276
1972] Hybrids in Hypericum — Adams 277
hybrid. These plants persisted in the Arboretum for many
years, there being collections made in 1916, 1921, and 1927
in the herbarium of the Arboretum.
Study of Rehder’s specimens of H. X Arnoldianum lend
support to his hypothesis of a hybrid origin for these plants.
The most convincing evidence is the presence of a very poor
seed set in the mature fruits on a collection made by Rehder
on October 9, 1910. In each capsule examined nearly all of
the ovules had aborted early in their development and only
a few well-formed seeds were present in each fruit.
The exact parental origin of H. X Arnoldianum is prob-
lematical. Some substantiation for Rehder's belief that
H. lobocarpum may have served as one of the parents of
this species is provided by analysis of the style numbers.
The 5-styled condition is characteristic of the majority of
fruits of H. lobocarpum: some 79% of 2014 capsules from
23 collections of wild plants had 5 styles (Adams, 1962).
While the 5-styled condition was found in only 3% of the
63 capsules of H. X Arnoldianum examined, about 16%
had 4 styles. The 4-styled condition was present in about
19% of the fruits of H. lobocarpum examined. In addition,
the deeply lobed condition so characteristic of the capsules
of H. lobocarpum is approached by the partially lobed
fruits of H. Arnoldianum. It seems reasonable, therefore,
to consider H. lobocarpum to have participated in the
ancestry of H. X Arnoldianum.
The identity of the maternal parent of H. X Arnoldianum
is less clear. Study of Rehder's specimens of this species
does afford support for his hypothesis that H. galioides
was involved. Evidence against this hypothesis is provided
by analysis of style numbers. Of a total of 63 mature fruits
of H. X Arnoldianum examined, 16% possessed 4 styles,
3% had 5 styles, and the remaining 81% were 3-styled.
Capsules of the putative maternal parent, H. galioides, are
almost invariably 3-styled, with 4-styled fruits being seen
very rarely.
What might have been the maternal parent of H. X
Arnoldianum? A clue may be provided by the striking
278 Rhodora [Vol. 74
resemblance between specimens of this species and plants
of H. densiflorum Pursh, especially those obtained from
populations of the latter species growing in eastern Ten-
nessee. In both plants the leaves are 10-15 times longer
than wide. In addition, the seed size and coat ornamenta-
tion of H. X Arnoldianum compare very well with that
present in H. densiflorum. Plants of H. densiflorum were
in cultivation at the time that Rehder described H. X
Arnoldianum and these could well have served as one of
the parents of this species. The close resemblance of H. X
Arnoldianum to plants from the eastern Tennessee popu-
lations of H. densiflorum could be explained by postulating
that some of the Arboretum’s living collections of the latter
species might have been originally obtained from the Ten-
nessee portion of the geographic range of H. densiflorum.
The available evidence, therefore, suggests that the spon-
taneous cross that is hypothesized to have produced H. X
Arnoldianum most likely occurred between plants of H.
densiflorum and H. lobocarpum. Rehder’s suggestion that
H. galioides may have been involved may have been due to
misidentification of specimens. Such an error is under-
standable, since several collectors of Hypericum in eastern
Tennessee, especially during the late 19th and early 20th
centuries, identified specimens of H. densiflorum as H.
galioides. However, both Svenson (1940) and Adams
(1962) have shown that the latter species is a native of the
lower Atlantic and Gulf Coastal plains.
Further support for the hypothesis that H. lobocarpum
and H. densiflorwm may have been the parents of H. X
Arnoldianum is provided by chromosome analysis. In
plants from the Arnold Arboretum identified as H. X
Arnoldianum (perhaps the same individuals studied by
Rehder), Hoar and Haertl (1932) found a gametic number
of 9. They reported “no irregularity in chromosome be-
havior nor morphological sterility of pollen" and concluded
that this hybrid “apparently came from compatible par-
ents.” Since they did not preserve voucher specimens, their
identification cannot now be verified. Jf they did indeed
1972] Hybrids in Hypericum — Adams 279
study the chromosomes of true H. X Arnoldianum, then
the observed chromosome regularity and viable pollen
would constitute strong support for the hypothesis that
the putative parents were indeed H. lobocarpum and
H. densiflorum. 'These two species are very closely related.
In fact, H. lobocarpum, a plant of Arkansas and adjacent
portions of surrounding states, has been considered as a
variety of H. densiflorum (Svenson, 1940), but the geo-
graphic ranges of each of these species are not known to
overlap, there being at least 80 miles distance between the
nearest known populations (Adams, 1962).
Hypericum X Dawsonianum Rehder l. c. The plants of
this putative hybrid were hypothesized by Rehder to have
orginated from a spontaneous cross between H. lobocarpum
Gattinger and H. prolificum L., with the latter as the
maternal parent. Two collections, both in mature fruit,
were made by Rehder in the Arboretum gardens during
the fall of 1910.
That Rehder's plants of H. X Dawsonianum are most
likely indeed of hybrid origin is strongly suggested by the
very poor seed set in the fruits, with most of the ovules
having failed to develop to maturity. Additional support
for this hypothesis is provided by an array of features
and eonditions more or less intermediate between the two
putative parents, H. lobocarpum and H. prolificum. The
seed size and coat ornamentation of the few seeds that are
present in the mature fruits are distinctly intermediate
between the two presumed parental species. The style
number of Rehder's collection of October 9 of H. X Daw-
sonianum is also intermediate between these two species:
about 50% are 3-styled and 50% are 4-styled. Almost the
same ratio of 3- and 4-styled capsules is present in Rehder's
collection of October 10; a very few fruits, however, had
5 styles. The 5-styled condition eccurred in about 79%
of 2014 capsules from 23 collections (Adams, 1962) of H.
lobocarpum. Fruits with 3 styles are the rule in H. pro-
lificum, with only an occasional capsule having 4 or, very
rarely, 5 styles. In addition, the fruit size of H. X Daw-
280 Rhodora [Vol. 74
sonianum is well within the range of H. prolificum. The
sulcate to deeply lobed capsules of H. X Dawsonianum,
however, are strikingly similar to those of H. lobocarpum.
Further substantiation of the hypothesis of hybridization
between H. lobocarpum and H. prolificum is the observa-
tion that interbreeding may occur occasionally between
plants of these two species growing in the field. I have
seen 3 collections from Ripley Co., Missouri (Steyermark
66885, F; Bush 282, GH, NY ; Makensie 395, NY) and one
from Howell County (Steyermark 40010, F, MO) which
I cannot assign definitely to either species. Several factors
favor the occurrence of hybridization between these two
species. The distributional range of the more southern
H. lobocarpum overlaps that of the more northern H. pro-
lificum in central Arkansas, forming a zone some 3 or 4
counties wide running from the southwestern corner of
the state diagonally across to southeastern Missouri
(Adams, 1962). Plants of both species flower at the same
time, and there is little obvious difference in their habitat
requirements. As living specimens of H. lobocarpum and
H. prolificum had been cultivated in the Arnold Arboretum
for many years, it seems reasonable to conclude that
Rehder’s H. X Dawsonianum specimens did indeed origi-
nate as the result of spontaneous crossing between these
two species under garden conditions.
The meiotic chromosome behavior of an individual plant
of H. X Dawsonianum was reported by Hoar and Haertl
(1932) as displaying no irregularity. Neither did they
observe any evidence of pollen sterility. Based on their
findings, they concluded that the putative parents of the
hybrid were genetically “compatible.” As with the case
of H. X Arnoldianum, however, no voucher specimens were
preserved and, therefore, it is not now possible to verify
the original identification.
Hypericum X nothum Rehder, l. c., 254. On October 10,
1910 Rehder collected specimens in the Arboretum which
he interpreted to be the product of a cross between H.
kalmianum L. and H. densiflorum Pursh, with the latter
as the maternal parent. There is little evidence, however,
1972] Hybrids in Hypericum — Adams 281
to support his hypothesis of hybrid origin of these plants.
The few 4- and 5-styled fruits present on the specimens of
the putative hybrid are suggestive of H. kalmianum, which
is typically 5-styled although 4 styles are not infrequently
encountered. However, most of the fruits on Rehder's
specimens have only 3 styles, a characteristic of H. densi-
florum. Other close resemblances to H. densiflorum include
seed shape, size and coat ornamentation, inflorescence form,
and fruit shape and size. These features strongly suggest
that Rehder's specimens of H. X nothum are most likely
not of hybrid origin. They can be readily assigned to
H. densiflorum.
Hypericum X VanFleetii Hort. ex Rehder, Man. Cult.
Trees and Shrubs, ed. 2, 640. 1940. According to Rehder,
this binomial refers to a horticultural form believed to be
a hybrid between H. prolificum and H. frondosum Michx.
There is no evidence that the several specimens of culti-
vated plants in the herbarium of the Arnold Arboretum
bearing this name are of hybrid origin. Instead, I believe
that they are readily referable to H. prolificum.
It is doubtful if any of these putative hybrids are still
being maintained by horticulturists. There were none
present in the gardens of the Arnold Arboretum in 1959.
This is not surprising since there are several introduced
species of Hypericum which make better garden plants,
with larger and more showy flowers and other more de-
sirable features. In the living collections at the Kew
Gardens in England there are several woody shrubs of
Hypericum Sect. Myriandra originally obtained from east-
ern North America. According to Dr. N. K. B. Robson of
the British Museum of Natural History (personal commu-
nication), these plants *do nct seem to fit exactly into any
of the described species." It is entirely possible, therefore,
that when two or more of the woody species are brought
into the garden, some hybridization may take place. In
nature, however, there appears to be very little interspecific
hybridization among the woody members of Hypericum
Sect. Myriandra.
282 Rhodora [Vol. 74
LITERATURE CITED
ADAMS, PRESTON. 1962. Studies in the Guttiferae. I. A synopsis
of Hypericum Section Myriandra. Contrib. Gray Herb. 189, 1-51.
Hoar, C. S. and E. J. HAERTL. 1932. Meiosis in the genus Hyperi-
cum. Bot. Gaz. 93: 197-204.
SvENSON, H. K. 1940. Plants of the southern United States. II.
Woody species of Hypericum. Rhodora 42: 8-19.
DEPARTMENT OF BOTANY
DEPAUW UNIVERSITY
GREENCASTLE, INDIANA 46135
A NEW SPECIES OF THE ORCHID
GENUS HOFFMANNSEGGELLA
H. G. JONES
The genus Hoffmannseggella was established by the
author (Jones, 1968) to accommodate the species which
were formerly assigned to the section Cyrtolaelia (Schlech-
ter, 1917) of the genus Laelia. In his review of Laelia
published in 1952, the late Dr. F. C. Hoehne, who prob-
ably knew these plants better than anyone else, wrote:
*Indubitavelmente, elas se distinguem tanto das que con-
stituem as outras seccoes, que poderiam ser agrupadas
como género distincto"; and later in the same paper;
“Acreditamos, entretanto, que haverá vantagem . . . e
razao para separar a seccao Cyrtolaelia de Laelia como
género distincto."
Apart from Laelia, one other generic name has been
associated with Hoffmannseggella cinnabarina (Batem.)
H. G. Jones, which is the generic type of Hoffmannseggella:
in 1843 Hoffmannsegg had transferred this species to his
new genus, Amalias. Unfortunately, in the original pub-
lication of Amalias (1842), Hoffmannsegg had made only
one specific combination, and this was based upon Laelia
anceps Lindl., which is a true species of Laelia, not belong-
ing to the section Cyrtolaelia. Consequently, since Amalias
anceps (Lindl) Hoff. automatically becomes the type-
species of Amalias, this name must be treated as a generic
synonym of Laelia, and was not available for use for the
species of section Cyrtolaelia, when the latter were raised
to generic rank.
The genus Hoffmannseggella falls naturally into two
fairly distinct sections: in the typical section (sect. Hoff-
mannseggella), the pseudobulbs are short and thick —
conical, subfusiform or ovoid in shape — and the flower-
scapes quite tall, usually much longer than the leaves. In
the species of sect. Pleurothalloides, however, the pseudo-
bulbs are reduced to slender cylindrical stems; and the
flower-scapes are much shorter, about equal to or very
283
284 Rhodora [Vol. 74
slightly longer than the leaves. This gives the plants a
rather distinct resemblance to the genus Plewrothallis —
hence the sectional name — a resemblance which has been
commented upon by Reichenbach (1874) and by Hoehne
(1930). In the course of delimiting the species of the
latter section for my projected monograph of the genus,
I have found it necessary to recognize one new species,
which is described and named below.
Hoffmannseggella brevicaulis H. G. Jones, sp. nov.
Species affinis H. harpophyllae a qua caulibus multo
brevioribus, floribus majoribus labello apice anguste atte-
nuato facile distinguitur. Epiphytica, erecta, circa 24 cm
alta; rhizomate valde abbreviato; radicibus albescentibus,
filiformibus, flexuosis, glabris; caulibus cylindraceis, uni-
foliatis, circa 7 cm longis; folio erecto oblongo, coriaceo,
apice obtuso vel subacuto, circa 17 cm longo, medio circa
2 em lato. Racemus erectus vel suberectus, 2-3-florus, quam
folio multo brevior; floribus luteis, circa 6.5 cm diametri-
bus: sepalis oblongis, apice acutis, circa 3.5 cm longis,
medio circa 9 mm latis; petalis quam sepalia aequimagnis
vel paulo brevioribus; labello circa 3 cm longo, medio circa
1.5 em lato, parte superiore profunde trilobo, carinis 2
paralellis elevatis, acutis; lobis lateralis subovatis, acutis;
lobo mediano elongato, anguste attenuato, apice multo
acuminato. Columna crassa subcylindrica, circa 1 cm longa,
medio circa 4 mm lata ; ovario pedicellato glabro, circa 4 cm
longo.
Type: Brazil habitat not recorded — flowered unde:
cultivation in Barbados, Feb. 1967. Herb Jones. H (L) /19a.
This taxon has been previously referred to by the author
in another paper (Jones 1970). It first came to my atten-
tion during the course of examining the specimens of H.
harpophylla (Rchb.f.) H. G. Jones, preserved in Reichen-
bach's herbarium (Naturhistorischen Museums, Vienna),
when I noticed that two of the sheets included under this
name were obviously atypical. These were subsequently
found to agree with cultivated specimens from Brazil which
1972] Hoffmannseggella — Jones 285
undoubtedly represented an undescribed species, somewhat
related to H. harpophylla, but easily distinguished by the
characters described in the above diagnosis, which may be
summarized as follows: —
H. HARPOPHYLLA
1. Plants tall and rather loose; pseudobulbs longer than
the leaves.
2. Leaves narrow, tapering to a slender, acute point.
3. Inflorescence numerous, 6-8-flowered.
4. Flowers rich orange-red.
5. Sepals and petals narrow, usually less than 6 mm
wide.
6. Front lobe of the labellum broadening outwards from
the base, apex obtuse.
H. BREVICAULIS
1. Plants shorter and more compact; pseudobulbs shorter
than the leaves.
2. Leaves broader, apex obtuse or subacute.
3. Inflorescence 2-3-flowered.
4. Flowers yellow.
5. Sepals and petals broader, 7-9 mm wide.
6. Front lobe of the labellum very narrow, tapering to
a long, slender point.
H. brevicaulis is also related to another quite recently-
discovered species, H. Kautskyi (Pabst) H. G. Jones
(Hoffmannseggella Kautskyi (Pabst) H. G. Jones, comb.
nov. Basionym: Laelia Kautskyi Pabst in Orch. Rev. 78:
321. 1970). The latter species resembles H. brevicaulis
in the color of the flowers, but the habit, the form of the
labellum, and the flowering period are all different. All the
species of Hoffmannseggella are confined geographically
to Brazil.
REFERENCES
HoEHNE, F. C. 1930. Album de Orchidáceas Brasileiras. 264 pp.,
Secretario da Agricultura, São Paulo.
HOEHNE, F. C. 1952. Algo concernente ao género Laelia Lindl.
das Orchidáceas e uma nova espécie para éle do Estado de Minas
Gerais. Arquivos de Botánica do Estado de Sáo Paulo 2: 157-167.
286 Rhodora [Vol. 74
HOFFMANNSEGG, G. 1842. Verzeichniss der Orchideen für 1842.
22 pp., C. Heinrich, Dresden.
HOFFMANNSEGG, G. 1843. Verzeichniss der Orchideen für 1843. 45
pp. H. M. Goltschalck, Dresden.
Jones, H. G. 1968. Studies in Neotropical Orchidology. Acta
Botanica Academiae Scientiarum Hungaricae 14: 63-70.
Jones, H. G. 1970. Orchidaceae Americanae. Caldasia 10: 491-495.
REICHENBACH, H. G. 1874. New Garden Plants. Gardener’s Chroni-
cle. 5: 542-543.
SCHLECHTER, R. 1917. Die Einteilung der Gattung Laelia und die
Geographische Verbreitung ihrer Gruppen. Orchis 11: 87-96.
POST OFFICE BOX 111
BRIDGETOWN, BARBADOS
WEST INDIES
CHROMOSOME NUMBER IN LIQUIDAMBAR
FRANK S. SANTAMOUR, JR.!
The genus Liquidambar belongs to the Hamamelidaceae,
although most authorities consider this genus so distinct
that they place Liquidambar, along with some other genera
such as Bucklandia and Disanthus, in the sub-family
Liquidambaroideae. Cytologically, all genera of Hamameli-
daceae (sub-family Hamamelioideae) reported thus far
have a basic chromosome number of x—12 (Darlington and
Wylie, 1955). Liquidambar is the only genus of the sub-
family Liquidambaroideae that has been studied cytologic-
ally.
Anderson and Sax (1935) studied meiosis in L. styra-
ciflua L. growing at the Arnold Arboretum in Jamaica
Plain, Massachusetts. They concluded that L. styraciflua
was a diploid (2»—30) with a basic number of z=15
chromosomes. They also noted marked meiotic irregu-
larities and high pollen sterility in their material, and
ascribed these irregularities to cultivation in an area north
of the natural species range.
Pizzolongo (1958) reported that the diploid number of
L. styraciflua (from shoot apices) and L. orientalis L.
(from root tips) was 2n—32, and concluded that the
haploid (and basic) number was »—16. However, the
only countable meiotic figure he found in L. styraciflua
was an anaphase I, showing groups of 15 and 16 chromo-
somes. Meiotic irregularities and consequent pollen sterility
apparently depended “upon some univalent chromosomes
which do not respect the metaphasic congression and cause
an unequal chromosome distribution among the pollen
grains."
This study was conducted in Italy, where the trees, as
in the Anderson and Sax work, were growing outside their
native range. Even with the irregular meiotic divisions,
"Research Geneticist, U. S. National Arboretum, Plant Science Re-
search Division, Agricultural Research Service, U.S. Department of
Agriculture, Washington, D.C. 20002.
287
288 Rhodora [Vol. 74
FA
‘fo
V
e.,
*a
LI +
Figure 1. Meiotic stages in Liquidambar styraciflua L.;
(a) Diakinesis with »—16 (540 X)
(b) Anaphase I with two groups of 16 chromosomes (1200 X).
however, Pizzolongo did not believe that the species had
two biotypes, with 2»—30 and 2n=32 chromosomes.
Work with Liquidambar in the project on “Cytogenetics,
Breeding, and Evaluation of Shade Trees" at the U.S.
National Arboretum has stressed interspecific hybridiza-
tion. We have obtained hybrids between L. stryaciflua and
both Asiatic species, L. orientalis and L. formosana Hance.
Thus, it was important to determine the number and
meiotic behavior of the chromosomes of our parent trees,
especially L. styraciflua, to decide if detailed cytological
analysis of the progenies would be necessary.
1972] Liquidambar — Santamour 289
In the spring of 1971, inflorescences of L. styraciflua
were collected from 4 native trees on the grounds of the
Arboretum in Washington, D. C. Two of these trees were
used as parents in the breeding work. The flowers were
fixed in 1:3 acetic-alcohol for 24 hours and stored in 80%
ethanol. Meiosis in the pollen-mother-cell was studied using
standard aceto-carmine squash techniques. Pollen was col-
lected from the same trees and pollen abortion was deter-
mined from a random sample of 200 grains stained with
aceto-carmine. Pollen size was based on measurements of
50 sound pollen grains from each tree. Mitosis in root
tips of seedlings of L. styraciflua X L. orientalis and L.
styraciflua X L. formosana parentage was also examined
using aceto-carmine squash techniques.
Accurate chromosome counts at first metaphase of
meiosis were, as ncted by previous authors, impossible to
achieve. The chromosomes at this stage are clumped to-
gether and no more than 10 distinct bivalents could be
observed. Usually one, and frequently two, bivalents were
found outside the metaphase grouping and exhibited pre-
cocicus separation. Pizzolongo attributed the high pollen
abortion he observed to this irregular chromosome behavior.
In my material it was not difficult to determine n=16
chromosomes at diakinesis or late anaphase I. Many diads
were counted with this number in both nuclei. At the
4-nucleate stage (anaphase II), nuclei with n=16 chromo-
somes could also be determined, but in no case could all
four nuclei be counted.
The major problem in documentation was obtaining
figures in which all the chromcsomes were in the same
plane for photographic purposes. The picture of diakinesis
(Fig. 1a) shows 16 chromosomes, with at least three and
perhaps four chromosomes attached to the nucleolus. This
configuration explains the multinucleolate microspores
noted by Pizzolongo. At anaphase I (Fig. 1b), two groups
of 16 chromosomes are shown.
290 Rhodora [Vol. 74
Pollen abortion in the four trees studied ranged from
6% to 20% and averaged 11%. Pollen grain diameter
averaged about 39 microns, and did not differ significantly
among trees. Although abnormal meiotic behavior may
explain the rather high pollen abortion sometimes found
in this species, the degree of meiotic irregularities probably
depends on individual tree characteristics.
Chromosome counts on root tips of hybrids of L. sty:a-
ciflua with L. orientalis and L. formosana gave 2n-—82.
These facts imply that all three species have the same
chromosome number.
LITERATURE CITED
ANDERSON, EDGAR & KARL Sax. 1935. Chromosome numbers in the
Hamamelidaceae and their phylogenetic significance. Jour.
Arnold Arb. 16: 210-215.
DARLINGTON, C. D. & A. P. WYLIE. 1955. Chromosome Atlas of
Flowering Plants (ed. 2) George Allen and Unwin Ltd., 519 pp.
PIZZOLONGO, PAOLO. 1958. Ricerche cario-tassonomiche su alcune
Hamamelidales. Ann. Bot. (Roma) 26: 1-18.
RECENT DISCOVERIES IN THE
NEW HAMPSHIRE FLORA
A. R. HODGDON AND FREDERIC L. STEELE?
Our field work in New Hampshire during the 1971 season
consisted only of a few day-excursions, no real effort having
been made to search for unusual plants. However, several
discoveries were made of which the following are most
significant.
Scirpus robustus Pursh. Saltmarsh at Adams Point,
Durham, Strafford County, A. R. Hodgdon, G. Brooke,
J. Wicks, et al., September 29, 1971 #19145. It is often
true that several rare and local species may occur in close
proximity. It happens that the halophytic composite Iva
frutescens var oraria, previously reported from Durham
(Hodgdon & Krochmal 1950), reaches its northeastern
known limit of range, aside from its disjunct occurrence
in Nova Scotia, within a stone’s throw of the Bulrush. The
ample size of the Scirpus colony shows that it must have
been growing here for a long time. Thus it must have been
overlooked by the senior author many times during the
past twenty or thirty years, perhaps because attention was
directed too much to the Iva. The nearest reported stations
for Scirpus robustus are in Newbury and Cambridge,
Massachusetts, making the Durham locality the initial
record for New Hampshire and the first north of Massa-
chusetts. It is a conspicuous tall sedge, leafy nearly to the
inflorescence, with big ovoid spikelets of rich brown color,
the plant in size somewhat resembling the also robust
Scirpus maritimus var Fernaldi which, however, typically
has elongated spikelets. In the herbarium of the New Eng-
land Botanical Club there are specimens from coasta] areas
of southwestern Maine which have regularly been called
S. maritimus var Fernaldi, but which have ovoid spikelets
or otherwise approach S. robustus in some characteristics.
'Published with the approval of the Director of the New Hampshire
Agricultural Experiment Station as Scientifie Contribution No. 616.
291
292 Rhodora [Vol. 74
Presumably the separation of these two taxa is not always
an easy matter.
Pycnanthemum Torrei Benth. Southern slope of Jeremy
Hill, Pelham, Rockingham County, A. R. Hodgdon and
Frederic Stecle, August 4, 1971 #19148. Jeremy Hill lies
within a mile or two of the Massachusetts line. The south-
ern slope where we fcund the Pycnanthemum has been
known for a long time as a rich collecting area; many
species of Desmodium and Lespedeza, Phryma leptostachya
and other plants of interest grow along the road which
extends upward on the slope into the woods. Arthur Stanley
Pease collected there and his success in finding unusual
species so far north led the senior author also to visit the
place a number of years ago. It has been known as the
northeastern known limit of range for Desmodium mari-
landicum (L.) DC., Lespedeza virginica (L.) Britton and
Pycnanthemum incanum (L.) Michx. while at least ten
other species are found on this slope that have not been
reported in Maine. On our August visit we explored the
rocky slope just above the road a little way into the woods
and came upon a clump or two of a big mint with which
we were unfamiliar but which we assumed to be a Pycnan-
themum and which proved to be P. Torrei. The calyx-teeth
of P. Torrei are acuminate and are about 1.5 mm long
serving readily to separate it from P. verticillatum or P.
virginianum. In addition the leaves are thin and mem-
branaceous and the heads are few at the tips of somewhat
elongated slender branches. The leaves of P. Torrei are
considerably wider than those of P. tenuifolium. This is
a somewhat remarkable extension of range all the way
from New Haven County, Connecticut where there are
two stations for it known in New England. It was col-
lected in New Haven by C. H. Bissell and R. W. Woodward
in August 1910 and in Cheshire also in New Haven County
by E. B. Harger in August 1929. Specimens from these
two localities are present in both the Gray and the New Eng-
land Botanieal Club Herbaria. It seems unlikely that this
mint, being scarce even in southern Connecticut, has moved
1972] New Hampshire Flora — Hodgdon and Steele 293
to Jeremy Hill in recent times. On the other hand, it is
surprising that it was overlooked by the botanists who
have visited the area unless indeed their attentions were
too much diverted by all the other interesting plants.
Lechea tenuifolia Michx. Along cleared trail to fire tower,
Jeremy Hill, Pelham, A. R. Hodgdon and Frederic L. Steele
#19146. Our success that August day, in finding a species
of Pycnanthemum seemingly new to New Hampshire, en-
couraged the first author to assume that other significant
species might also be discovered on Jeremy Hill. It was
decided therefore to follow a trail upward to the fire tower.
Plants of Viola pedata were scattered along the trail and
higher up, where the trail widened into a cleared roadway
there were plants of Hypoxis hirsuta in good flower. A few
depauperate plants of the pinweed, Lechea tenuifolia were
growing nearby. In 1879 Kate Furbish found this species
in Wells, York County, Maine. In the Herbarium of the
New England Botanical Club one can find this Wells speci-
men but none from New Hampshire. In the Gray Her-
barium however, there are two specimens from New
Hampshire both being F. W. Batchelder collections from
Pelham, one from the Gumpas District, the other with no
precise record of locality. It is likely then that ours is the
first record from Jeremy Hill and since Batchelder’s more
recent collection was made in 1902, a contemporary com-
munication about it may be justified to show that it still
is to be found north of the Massachusetts line.
Another pinweed, Lechea villosa', is found on Jeremy
Hill and indeed quite generally in townships close to the
Massachusetts line in New Hampshire, Durham being a
somewhat isolated outpost for it to the northeast.
‘It seems preferable to retain the binomial Lechea villosa of Elliot
which is clearly typified and about which there is no question instead
of the vague, poorly defined and dubious L. mucronata of Rafinesque
who published several other names that might be interpreted as
L. villosa and who demonstrated in his published study of the genus
that he failed to understand the group.
294 Rhodora [Vol. 74
Cyperus Houghtonii Torr. Sandy roadside by pond near
Silver Lake, Hollis, Hillsboro County, A. R. Hodgdon and
Frederic L. Steele, August 25, 1971 #19144. This rare
species is not new to New Hampshire, having previously
been collected in Bath and Hinsdale in the Connecticut
Valley. Its distribution in New England is somewhat un-
usual, however, and therefore deserves comment. We found
it growing along with Cyperus strigosus, Fimbristylis
autumnalis, Bulbostylis capillaris and Panicum philadel-
phicum at the edge of a road near a small pond not far
from Silver Lake in Hollis. It was difficult to sort out the
depauperate plants of Cyperus from the other plants with
which it grew, indicating that it may be more common in
suitable situations, obscured at times by associated plants,
than it now seems. Its New England range includes Win-
slow (Kennebec Valley), Maine where it was discovered
by Ralph Bean on July 22, 1915, Bath and Hinsdale, New
Hampshire, Chittenden and Rutland Counties in western
Vermont and several scattered localities in Massachusetts,
there being no records reported by Seymour (1969) from
Connecticut and Rhode Island.
Specimens of the four principal species discussed in this
article have been deposited in the herbaria of the Univer-
sity of New Hampshire and the New England Botanical
Club.
LITERATURE CITED
Hopepon, A. R. and STANLEY KROCHMAL. 1950. Range — extensions
and clarifications in New Hampshire. Rhodora 52: 162-163.
SEYMOUR, FRANK C. 1969. The Flora of New England. Charles E.
Tuttle Co., Rutland, Vermont. 596 p.
DEPARTMENT OF BOTANY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM 03824
and
ST. MARY’S-IN-THE-MOUNTAINS
LITTLETON, NEW HAMPSHIRE
SOME ALGAL INVADERS OF
THE NORTHWESTERN FRINGES
OF THE SARGASSO SEA
WILLIAM J. WOELKERLING
The Sargasso Sea encompasses a wide but not sharply de-
limited area (biologically speaking) of the North Atlantic
Ocean situated roughly between 20 and 40 degrees north
latitude and 35 and 75 degrees west longitude (Marmer
1928) and owes its name to the presence of two species of
brown algae: Sargassum fluitans Boergesen and S. natans
(L.) J. Meyen. On a recent cruise (during May 1970), the
author observed numerous plants of Ascophyllum nodosum
(L.) LeJolis and Fucus vesiculosus L. adrift with Sargas-
sum along the northwestern fringes of the Sargasso Sea.
The area visited includes the region from 69°24’ west long-
itude to 69°48’ west longitude and 38°53’ north latitude to
39°11’ north latitude (ca. 375 square miles) and lies rough-
ly 125 miles south of Nantucket Island, 150 miles south of
Cape Cod, Massachusetts, 170 miles east-southeast of Mon-
tauk Point, Long Island, and 75 miles north of the Gulf
Stream.
The occurrence of macroscopic algae within the region
appeared to be sporadic; densities as high as one plant per
square meter were encountered in some places while at
others virtually no weed was observed. Entangled masses
of Aescophyllum and Fucus similar to those reported for
Sargassum (Winge 1923) were not observed.
Samples taken at several stations with the aid of a long-
handled dip net revealed both epiphytized and unepiphytized
plants of Ascophyllum and Fucus. In all, 14 species of al-
gae were collected (Table 1) including 2 Chlorophyta, 10
Phaeophyta, and 2 Rhodophyta. None of the epiphytic spe-
cies has been recorded from the Sargasso Sea previously
although isolated plants of Ascophyllum and Fucus have
been reported (Collins 1917, Collins and Hervey 1917,
Winge 1923). Many individuals of Ascophyllum and Fucus
295
296 Rhodora [Voi. 74
examined appeared fresh and may still have been growing
at the time of collection, but others bore very distended re-
ceptacles and showed signs of vegetative decay. Most of
the epiphytes appeared somewhat moribund and lighter in
color than their mainland counterparts and probably were
not actively growing. These observations suggest that at
least the epiphytized plants encountered represent detached
specimens which have drifted out from the east coast of the
United States rather than true pelagic forms.
Both Ascophyllum and Fucus, however, are reported to
propagate vegetatively in the drifting state in European
waters (Levring 1940, Oltmanns 1889, Reinke 1892, Sauva-
geau in Collins and Hervey 1971, p. 78) and further study
appears warranted to determine whether reproduction of
any sort similarly occurs in the northern Sargasso Sea. The
presence of apparently healthy, unepiphytized plants of As-
cophllum and Fucus in the region visited certainly suggests
the possibility of vegetative propagation. In fact, Collins
and Hervey (1917, p. 79) have concluded (without experi-
mental data) that Ascophyllum grows actively and occasion-
ally reproduces sexually in the Sargasso Sea. Until more
direct evidence comes to hand, however, it seems best to re-
gard Ascophyllum, Fucus and particularly their associated
epiphytes as invaders rather than permanent components of
the Sargasso Sea flora.
Sincere thanks are due Gordon Volkmann of the Woods
Hole Oceanographic Institution for arranging passage on
the WHOI research vessel ATLANTIS II. One of the prepared
sets of herbarium specimens has been retained by the au-
thor while duplicates have been deposited in the Farlow
Herbarium of Harvard University and the University of
California Herbarium at Berkeley. The work described
here was supported by Grant GB-13250 from the National
Science Foundation to the Systematics-Ecology Program.
SEP Contribution No. 231.
1972] Algal Invaders — Woelkerling 297
REFERENCES
COLLINS, F. S. 1917. The Sargasso Sea. Rhodora 19: 77-84.
COLLINS, F. S. and A. B. HERVEY. 1917. The algae of Bermuda.
Proc, Am. Acad. Arts. Sci. 53(1): 1-195.
LEvRING, T. 1940. “Studien über die Algenvegetation von Blekinge,
Südschweden". Dissertation. Lund.
MARMER, H. A. 1928. The Sargasso Sea. U. S. Naval Inst. Proc.
54(1): 33-35.
OLTMANNS, F. 1889. Beiträge zur kenntnis der Fucaceen. Bibl.
Bot. 14: 1-94, pl. 1-15.
REINKE, J. 1892. Uber giste der Ostseeflora. Ber. dt. bot. Ges.
10: 4-12.
WINGE, O. 1923. The Sargasso Sea, its boundaries and vegetation.
Rep. Dan. oceanogr. Exped. 1908-1910 3(2): 1-34.
TABLE 1. Algae Collected Along Northwestern Fringes
of Sargasso Sea, May 1970.
SPECIES REMARKS
Chlorophyta :
Monostroma pulchrum On Fucus
Farlow
Spovgomorpha arcta
(Dillwyn) Kuetzing On Fucus
Phaeophyta :
Ascophyllum nodosum Specimens up to 1.0 m long
(L.) LeJolis collected
Chordaria flagelliformis Immature; on Fucus
(Müller) C. Ag.
Elachistea lubrica On Ascophyllum and Fucus
Rupr.
298 Rhodora [Vol. 74
Fucus vesiculosus L. Specimens up to .5 m long
collected
Isthomoplea sphaerophora On Polysiphonia lanosa in
(Harvey in Hooker) turn on Ascophyllum
Kjellman
Punctaria latifolia Grev. On Fucus
Punctaria plantaginea On Fucus
(Roth) Grev.
Pylaiella littoralis L. On Fucus
Kjellman
Sargassum natans L. Pelagic
J. Meyen
Scytosiphon lomentaria On Fucus
(Lyngbye) C. Ag.
Rhodophyta :
Erythrotrichia carnea On Fucus
(Dillwyn.) J. Ag.
Polysiphonia lanosa L. On Ascophyllum
Tandy
DEPARTMENT OF BOTANY
THE UNIVERSITY OF WISCONSIN
MADISON, WISCONSIN 53706
INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA
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nyms. Papers of a floristic nature should follow, as far as
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Volume 74, No. 798. including pages 161-299. was issued June 30, 1972
299
CONTENTS: — continued
A White-Flowered Form of Iris lacustris from Ontario.
J. E. Cruise and P. M. Catling crsscccsccccssssseeessecsesseereesennenens 271
Neomirandea allenii, A New Epiphytic Composite of the
American Rain Forest.
R. M. King and H. Robinson .sssesssrrrereserererereereererererereees 272
Studies in the Guttiferae. III. An Evaluation of Some
Putative Spontaneous Garden Hybrids in Hypericum
Sect. Myriandra. Preston Adams .........emnm 276
A New Species of the Orchid Genus Hoffmannseggella.
H. G. JONCS ceccccssscecccsssssccecscessecseessssssscessseeeeeeeeenaeesssseseeegeceeesees 283
Chromosome Number in Liquidambar.
Frank S. Santamowr, J.. eene nnne nnn nennen 287
Recent Discoveries in the New Hampshire Flora.
A. R. Hodgdon and Frederic L. Steele .........../:2 291
Some Algal Invaders of the Northwestern Fringes of the
Sargasso Sea. William J. Woelkerling |... eene 295
Instructions to Contributors. .............. eene 299
Hodova
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by
ALBION REED HODGDON, Editor-in-Chief
ALBERT FREDERICK HILL
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON > Associate Editors
RADCLIFFE BARNES PIKE
ELIZABETH ANNE SHAW
STEPHEN ALAN SPONGBERG
Vol. 74 September, 1972 No. 799
CONTENTS:
Intraspecific Variation in Chromosomes of Sedum in the
Southwestern United States.
D IL de EST RR
The Rachilla in Cape Cod Species of Carex with Notes on
the History of the Perigynium and Rachilla.
Henry K. Svensom verccsssssssssssssssrsrssesevsrssssstssssesssesssavscarasasscessesee
Distribution of Native and Introduced Knapweeds (Cen-
taurea) in Canada and the United States.
EE NN E TNE E AEA o E E D
A New Variety of Leucophyllum laevigatum (Scrophu-
lariaceae) from Mexico.
Robert W. Kiger wcscscssssssssscsssssssssssssssssscsssesesessssassessecesssssssessees
(Continued on Inside Back Cover)
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Rbodora
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Vol. 74 September, 1972 No. 799
INTRASPECIFIC VARIATION IN CHROMOSOMES OF
SEDUM IN THE SOUTHWESTERN UNITED STATES
CHARLES H. UHL
Many species of Sedum include plants with different
chromosome numbers (Uhl, 1961, 1970b). This intraspe-
cific variation in number occurs in two patterns, here desig-
nated as established variation and casual variation. Estab-
lished variation is represented by clear-cut geographical or
ecological chromosome races, where any population clearly
has a standard chromosome number, but populations in
other areas or habitats may differ in the number of chromo-
somes in their sets (dysploidy) or in their number of
chromosome sets (polyploidy). Casual variation is more
random and accidental and is represented by individual
plants that depart from the locally established chromosome
race in ways that usually reflect some cytological instabil-
ity or irregularity. The commonest examples are trisomy
(2n + 1) and the presence of a variable number of small,
extra, accessory, or “B” chromosomes.
Each established variant in chromosome number has a
balanced, more or less stable and true-breeding outfit of
chromosomes that has been tested and proved and that oc-
cupies a geographical or ecological portion of the range
of the species. Most casual variants, on the other hand,
have an unstable outfit of chromosomes that does not breed
true to its own chromosomal type. Casual variants repre-
sent a sort of cytological “noise” or "static", and where
501
302 Rhodora [Vol. 74
they are common they may sometimes blur or obscure the
pattern of established variation. They may originate as
segregants from hybrids with related species or between
different established variants (chromosome races), or their
occurrence may simply indicate that the population has not
yet evolved to chromosomal stability. Doubtless each estab-
lished variant first arose as a casual variant that was suc-
cessful and eventually stabilized, and this process must
still be going on. A relatively high background of casual
variation might be expected in species with a significant
amount of established variation. On the one hand, this
reflects the continuation of the processes (instabilities)
that originally led to the chromosome races (established
variants), and on the other hand it may result from the oc-
currence of hybrids where the isolation was not yet com-
plete.
The species of Sedum in the southwestern United States
offer good examples of both types of chromosomal varia-
tion. This paper describes the chromosomes and the dis-
tribution of the chromosome races in five of these species:
S. cockerellii Britton (including S. griffiihsii Rose), S.
niveum Davidson, S. stelliforme S. Wats., S. wrightii A.
Gray, and S. havardii Rose. Probably all of these species,
as well as other related species, occur in adjacent areas of
northern Mexico, but material from there has been limited,
except for S. wrightii. Living plants of these species are
easy enough to identify, but many specimens of them are
incorrectly identified in herbaria. In particular S. cocker-
ellii, S. stelliforme and S. wrightii have been confused, and
ranges given for these species in some Floras cannot be
depended upon.
Most plants were collected in the field but grown in cul-
tivation for a time before study. Counts were made from
conventional] aceto-carmine squash preparations of pollen
mother cells. The photographs are from permanent prep-
arations, all X 2000. Voucher specimens are in the Wie-
gand Herbarium or Bailey Hortorium of Cornell Univer-
sity.
1972] Sedum — Uhl 303
Clausen and Uhl (1943) classified S. cockerellii and S.
griffithsii, as then known, into separate subsections, largely
on the basis of a difference in chromosome number »—16
and n=—14 or 29, respectively). However, the two are very
closely related, and in my opinion the additional chromo-
somal and morphological variation known today make it
impractical to separate them. Therefore S. cockerellii is
conceived broadly here, to include also S. griffithsii and
the various other binomials earlier considered synonymous
with one or the other of them (Clausen and Uhl, 1943).
Thus conceived, S. cockerellii has at least four chromo-
some races (established variants), with 14, 16, 29 or 30
gametic chromosomes. Probably the lone Texas population,
with n=15, represents a fifth chromosome race. Each of
these races occupies a definite geographic area, and in no
case was more than one chromosome race found in the same
range of mountains (Fig. 29). One or more extra chromo-
somes were found in plants from six widely separated lo-
calities. These casual variants occurred in populations in
which the standard numbers (as confirmed by counts from
other plants in four of the six populations) were «—14
(2 populations), n=15 (1 population), and n=16 (3 pop-
ulations). In all but one case the extra chromosomes were
smaller than the regular members of the set; they some-
times differed among themselves in size (e.g., Fig. 4) and
occasionally paired with each other, although usually not
with members of the standard set. Most of them seem to
represent typical examples of accessory or “B” chromo-
somes. Their origin and function, if any, are completely
unknown.
In general, plants of S. cockerellii from south central
Arizona and northern Sonora have 14 pairs of chromo-
somes (Fig. 1). These include typical S. griffithsii (C5176).
One plant from the Huachuca Mountains (U887A) had two
small extra chromosomes that formed a bivalent in 7 of
the 23 metaphase I plates examined (Fig. 2) but remained
unpaired in the remaining 16 in the same anther (Fig. 3).
Another plant from the same population (U887B) had the
304 Rhodora [Vol. 74
standard 14 pairs. Still another plant, from Sycamore
Canyon, southeast of Ruby (UC65.135), had three small,
extra chromosomes (Fig. 4), but six other plants collected
at various times from the same locality (U884, U1824) all
had the standard 14 pairs.
The lone Texas population, which I believe is a new spe-
cies record for the state, has n=15 (Fig. 5), although one
of the four plants studied had a small, extra chromosome,
possibly an accessory (Fig. 6). One pair of chromosomes
is large, perhaps the result of combining most of two
chromosomes from an ancestor with n=—16.
Plants of S. cockerellii ranging from Oak Creek Canyon
in north central Arizona to the White Mountains in the
east central part of the state, mostly along or near the
Mogollon Rim, as well as most collections from New Mex-
ico, have n—16 (Fig. 7). These include plants from the
vicinity of the type locality (U1315, U1316, C5254). Con-
strictions, apparently kinetochores, are detectable in some
chromosomes of mitotic metaphases (2n—32, Fig. 8). Thus
there is no reason to believe that diffuse or multiple kine-
tochores are related to the dysploidy in this species. A
plant from the White Mountains of east centra] Arizona
had a small accessory chromosome (U1155, Fig. 9), but
plants from six other localities in the same mountains all
had the standard 16 pairs. One plant from the Jemez
Mountains of north central New Mexico had a small ac-
cessory chromosome (U1147G) and another from the same
locality (C5281) had several (six in Fig. 10), but seven
other plants from the same population had the standard
16 pairs (U1147A, B, C, D, E, F, H). Still another plant,
from the Sangre de Cristo Range in northern New Mex-
ico, had probably four extra chromosomes (U1133, Fig.
11). Since these were large enough to represent standard
chromosomes, and since they appeared often to enter into
multivalent formation with regular chromosomes (Fig. 12),
the plant is possibly a quadruple trisomic (2n+4), per-
haps descended from a triploid.
In the Mule Mountains (near Bisbee), in the Chiricahuas,
1972] Sedum — Uhl 305
on Mount Graham, and on Kitt Peak in the Quinlan Moun-
tains, all in southern Arizona, S. cockerellii has n=29
(Fig. 13). Plants from the Sierra Blanca Range in south-
ern New Mexico have n—30 (Fig. 14). In these two tetra-
ploid chromosome races univalents and multivalents are
formed occasionally and make it difficult to detect any cas-
ual variation in chromosome number. None was noted in
the 18 tetraploid collections studied.
S. niveum is closely related to S. cockerellii (Clausen
and Uhl, 1943). High in the San Bernardino and Santa
Rosa Mountains of southern California this endemic spe-
cies has n—16 (Fig. 15), as does one race of S. cockerelli.
S. niveum has recently been discovered about 200 miles to
the south, high in the Sierra San Pedro Martir of north-
ern Baja California (Moran, 1969). Here it is octoploid,
with n—64 (Fig. 16), but occasionally forming a few mul-
tivalents and univalents at meiosis.
At one of its two type localities, Carr Peak in the Hua-
chuca Mountains of southeastern Arizona, S. stelliforme
has n=20, plus a variable number of accessory chromo-
somes, two to seven in different plants, (Figs. 17, 18). An-
other plant, probably this species, from more than 400
miles to the southeast in southern Chihuahua, Mexico, also
had »—20 (Fig. 19), plus one or two extra, accessory
chromosomes in some cells. Very similar plants from the
White Mountains of east central Arizona and from the
Black Range in southwestern New Mexico are considered
for the present to be also the same species, but they all
have n=12 (Fig. 20). The cause and significance of the
two different, apparently unrelated, chromosome numbers
(established variants) in the species is not at all clear;
perhaps closer morphological study may support their tax-
onomic separation. One plant from the Black Range
(U1625A) had 8 small accessory chromosomes (six are
shown in Fig. 21), but two other plants from the same
population and plants from seven other populations in the
same mountains all had »—12.
S. wrightii includes plants with four levels of ploidy
306 Rhodora [Vol. 74
based on z—12, diploid through octoploid (n=12, 24, 36,
and 48; Figs. 22, 24-26). No plants with accessory chromo-
somes or other casual variants were definitely noted, but
one collection (UC52.1680) clearly had 13 pairs of chromo-
somes (Fig. 23), none of which was distinctively smaller
than the others. Four plants collected later from what was
presumed to be the same locality (U879) all had the stand-
ard diploid n=12. It is not certain whether the plant with
n=13 is better considered another established variant or a
casual variant. Univalents, multivalents, and close second-
ary associations are common enough to make exact counts
difficult in some hexaploids (n=36) and octoploids (n=
48). Probably these are autopolyploids. Autopolyploidy has
been reported in S. ternatum (Baldwin, 1936; Uhl, 1970b),
and, on the basis of evidence from chromosome pairing in
hybrids, it appears to be common in the Mexican Crassu-
laceae (Uhl, 1970a, and unpub.).
The four principal chromosome types (established vari-
ants) of S. wrightii are not as discrete geographically
(Fig. 30) as those of S. cockerellii and S. stelliforme (Fig.
29). Only diploids were found in southern New Mexico,
and they occur also in the Davis Mountains of western
Texas and in the vicinity of Saltillo, Coahuila, Mexico.
Tetraploids (n=24) were found at the presumed type lo-
cality (U882, near El Paso), near Marathon, Texas, in the
Figs. 1-20. Sedum chromosomes in pollen mother cells at meta-
phase I (except Figs. 8, 16), 32000. Arrows identify univalents
and/or accessory (“B”) chromosomes. Figs. 1-14. S. cockerelli.
Fig. 1. U883, »—14. Fig. 2. U887A, n=15. Fig. 3. U887A, n—14--
2, Fig. 4. UC65.135, w—14--3B. Fig. 5. U880C, w—15. Fig. 6.
U880B, »—15-4-1B. Fig. 7. U900, »—16. Fig. 8. U900, mitosis in
anther wall, 25—32. Note constrictions (kinetochores) in some
chromosomes. Fig. 9. U1155, w—16--1B. Fig. 10. C5281, n—16+
6B. Fig. 11. U1133, »—16--4B. Fig. 12. U1133, 24, (identified by
lines) --14&--2, Fig. 18. U888, »—29. Fig. 14. U1307, n—30.
Figs. 15-16. S. niveum. Fig. 15. M345, n—16. Fig. 16. M15333, n—
64 (Metaphase II). Figs. 17-20. S. stelliforme. Fig. 17. U1680A,
n—20--2B. Fig. 18. U1680C, n—20+7B. Fig. 19. U2051, n=20.
Fig. 20. U1312, »—12.
1972]
Sedum — Uhl
507
308 Rhodora [Vol. 74
Figs. 21-28. Sedum chromosomes in pollen mother cells at meta-
phase I, X 2000. Arrows identify univalents and/or accessory (“B”)
chromosomes. Fig. 21. S. stelliforme. U1625A, n—12--6B. Fig. 22-26.
S. wrightii. Fig. 22. U879, n—12. Fig. 23. UC52.1680, »—13. Fig.
24. UC53.407, n—24. Fig. 25. U877, n—36. Fig. 26. M6317, »—48.
Figs. 27-28. S. havardii. Fig. 27. UC52.1679, n—54, Fig. 28. U876,
m—81.
Sierra del Carmen (across the Rio Grande from Big Bend)
and in the Sierra Madre Oriental in southern Nuevo Leon,
Mexico. Hexaploids (»—36) were found at Big Bend and
135 miles to the east, near Del Rio, Texas, and octoploids
(n—48) only in southern Nuevo Leon.
S. havardii is apparently of Mexican affinities. It is
known in the United States only in the Big Bend region of
western Texas, where plants with 54 and 81 gametic
chromosomes have been found (Figs. 27, 28). Presumably
the basic karyotype of this species consists of 27 chromo-
somes, but no collection with this number has yet been
found. The only collection with n—81 (U876) was made
only 11/2 miles from another (U875) with n=54. Thus
it is not yet clear whether the plants with the different
numbers really represent established chromosomal vari-
1972] Sedum — Uhl 309
1
S. cockerellit
© n=14 ascen
— — A nsi5
D n=!6 asa
e n-29 D uU
mn-3O mu di m
S. stelliforme Dt
. n=12 m by
x n=20 n
[ ]
He
E
A x :
Fig. 29. Distribution of chromosome races (established variants)
of Sedum cockerellii (s. lat.) and S. stelliforme in New Mexico,
Arizona and adjacent regions.
310 Rhodora [Vol. 74
nzl2
n=13
n=24
n=36
n=48
$
o NEW MEXICO
-eonpo
a TEXAS
(2
Fig. 30. Distribution of chromosome races of Sedum wrightii in
New Mexico, Texas and northeastern Mexico,
1972] Sedum — Uhl 311
ants (races) or only two samples from a large population
containing casual variants in ploidy. Two of the most sim-
ilar species are S. diffusum (x—19) of northeastern Mex-
ico and S. alamosanum (x—18) of northwestern Mexico,
both of which also have polyploid races.
Plants of S. cockerellii with n=16 (U1690) have been
found within inches of S. stelliforme with n—12 (U1691)
in the White Mountains, and plants of S. griffithsii (here
included in S. cockerellii) with n—14 on Carr Peak in the
Huachuca Mountains (U1679) are no more than a mile
from plants of S. stelliforme that have n—20 + accessories
(U1680). At two localities on opposite sides of Red River
Pass in the Sangre de Cristo Range of northern New Mex-
ico, S. cockerellii with n=16 (U1134, U1138) was found
within inches of the yellow-flowered, tetraploid S. lanceo-
latum, also n=16 (U1135, U1139), with both species in
flower together. This locality is near the northern limit of
the range of S. cockerellii, near the southern limit for S.
lanceolatum.
S. cockerellii was never found close to S. wrightii, al-
though perhaps the two might occur together in the Davis
or Organ Mountains. S. cockerellii generally occurs in the
Transition and Canadian Zones, nearly always near streams
that flow all year, whereas S. wrightii is generally a plant
of the warmer and drier Sonoran Zone (Clausen and Uhl,
1943). However, in the Sacramento Mountains about
Clouderoft, New Mexico, diploid S. wrightii (n—12) oc-
curs at higher elevations (four collections: 8000-8400’)
than does S. cockerellii 20-30 miles north in the Sierra
Blanca Range about Ruidoso (five collections: 7100-7600’).
Each species was found in only the one range in this area
and not found in the other. S. wrightii (hexaploid, U873)
occurs with S. havardii (U875) in the Chisos Mountains
of Big Bend National Park and with other species in Mex-
ico.
Each of the five species has two or more chromosome
races (established variants). Polyploidy definitely occurs
in four of the five, and it may also account in some degree
312 Rhodora [Vol. 74
for the difference between the races of S. stelliforme with
n—12 and n=20 + B's. Extra chromosomes, usually ac-
cessories, were found in some diploid plants in three spe-
cies. Probably this sort of casual variation should be ex-
pected where so much established variation also occurs.
In the same region Graptopetalum rusbyi of Arizona also
includes polyploids and variations from strict diploidy
(Uhl, 1970a).
Apparently quite a few structural changes have occurred
in the chromosomes of these species. Structural rearrange-
ments must have been involved in the evolution of the dif-
ferent numbers (14, 15, 16) in diploid S. cockerellit.
Chromosome bridges at anaphase, which usually indicate
heterozygosity for an inversion, were not especially looked
for, but they were common in two plants of this species
(U887A and C5176).
Differences in chromosome number interfere with free
interchange of genetic material and thus they represent a
means of reproductive isolation that may eventually lead
to new taxonomic species. However, anointing each estab-
lished variant (chromosome race) as a different taxonomic
species, as some (e.g., Love, 1951) have favored for other
groups, seems undesirable here. It would increase the num-
ber of species reported on from five to at least 15, and
many of the splinter species probably could not be distin-
guished without cytological study. Thus it seems better to
take a broad concept of these species. The chromosomal
variants are considered to indicate that evolutionary
changes are actively occurring in the five species, but the
changes are considered not yet to have crossed the thresh-
old of the practical, taxonomic species. More careful mor-
phological study, such as that now being carried out by
my colleague, Professor R. T. Clausen, may some day re-
quire modification of this conclusion for one or more of the
five species.
The large proportion of chromosomal variants in these
species finds a parallel in certain other species of Sedum
in the United States, e.g., S. glaucophyllum and S. terna-
1972] Sedum — Uhl 313
tum (Uhl, 1970b), and also S. lanceolatum (Uhl, unpub.).
However, it contrasts sharply with the situation in still
others. For example, in 78 collections of S. spathulifolium,
ranging from Vancouver Island to southern California and
exhibiting considerable variation in such characters as leaf
thickness, width, crowding and glaucosity, no established
variants were found, and all plants but two had seemingly
identical karyotypes with n—15 (Uhl, unpub.). The only
two exceptions were both casual variants: a plant with
n=14 in a population that also included a standard plant,
and a triploid. Seventy-nine other collections, representing
five other species related to S. spathulifolium, also all had
n=15, with no variants, either established or casual. How-
ever, one related species had n=30 and another n—45. It
is not clear whether the relative scarcity of non-standard
chromosome complements in these species means that they
are produced less frequently or that they survive less well,
or both. At any rate, the karyotype here has remained
stable in number and probably also in form during the
evolution of enough morphological diversity that six spe-
cies are recognized and while spreading 1100 miles apart.
This stability must reflect differences in the nature of the
genetic and chromosomal changes that have contributed to
evolution in S. spathulifolium and its relatives (subgenus
Gormania), contrasted with those prevailing in most of the
species described earlier.
Summary
Variation in chromosome number is found within each
of five species of Sedum of the southwestern United States.
Established variation is represented by polyploid and/or
dysploid geographic chromosome races, two to five for each
species. Casual variation, represented by the occurrence of
occasional plants that are trisomic or that bear accessory
chromosomes, was found in two, possibly three, species.
Successful casual variants can become established variants
(races). Although differences in chromosome number are
accompanied by some degree of reproductive isolation, rec-
314 Rhodora [Vol. 74
ognition of each chromosomal variant as a different species
does not seem appropriate in these actively evolving cases.
Acknowledgement
I thank those who have provided some of the plants re-
ported on here, especially Dr. Reid Moran, of the Natural
History Museum, San Diego (M nos.), and Mr. Paul Hutch-
ison, formerly of the University of California Botanical
Garden, Berkeley (UC nos.). Publication was assisted by
the L. W. Sharp Fund of Cornell University.
COLLECTIONS STUDIED
S. cockerellii Britton
Arizona:
U894 n—14 Pima Co. 4 mi. N. of road summit, Mt. Lemmon,
Santa Catalina Mts., 6240'.
U893 n—14 Pima Co. 0.8 mi. below Mt. Lemmon Lodge, 7520’.
UC52.1674 n—14 Same locality, (E. R. Blakely).
U892 n—14 Pima Co., Bear Canyon, 15 mi. by road S. of Mt.
Lemmon Lodge, 5480’.
U883 m—14 Santa Cruz Co. Madera Canyon, Santa Rita Mts.,
6400’.
C5176 m—14 Santa Cruz Co. near summit, Baldy Peak, Santa
Rita Mts., 8900’. ? Topotype of S. griffithsii Rose,
(R. T. Clausen).
U884 n=l4 (5 plants) Santa Cruz Co., Sycamore Canyon, 5 mi.
S.E. of Ruby, Atasco Mts., 3960’.
U1824 n=l4 Same locality, (A. M. Phillips III).
UC65.135 n—14+3B Same locality, (D. M. Hutt).
U885 n—14 (2 plants) Santa Cruz Co., Sonoita Cr., 3 mi. S.W.
of Patagonia, 3920'.
U886 «—14 Santa Cruz Co., Mile 16 N.E. of Nogales, 2.8 mi
S.W. of Patagonia, 3900’.
U887A m—14--2, Cochise Co. Ramsay Canyon, Huachuca Mts.,
6000'.
U887B n»—14 Same locality.
U1681 14 Cochise Co., Reef Mine, Upper Carr Canyon, Hua-
chuca Mts., 7100'.
U1679 mn—14 Cochise Co., N. slope of Carr Peak, Huachuca Mts.,
7700’.
C5189 m—14 Cochise Co. Same locality, (R. T. Clausen).
1972] Sedum — Uhl 315
UC52.1678 n=l4 Cochise Co., Guadalupe Canyon, Guadalupe Mts.,
4300’, (E. R. Blakely).
Mexico:
M14849 w—14 Sonora. Cerro Colorado, S.E. of Cananea, (C. Glass
and D. Gold).
Arizona:
U900 n—16 Coconino Co, Oak Creek Canyon, 12 mi. N. of
Sedona, 5600'.
U1617 m—16 Coconino Co., Macks Crossing, E. Clear Cr., 15 mi.
N.E. of Clints Well, 6260'.
UC52.1673 n—16 Gila Co., N. peak of Four peaks Mt., Mazatzal
Mts., 7645', (E. R. Blakely).
U1688 m—16 Gila Co., Workman Creek Falls, 44 mi. N. of Miami,
Sierra Ancha, 6600’.
U1618 »—16 Apache Co. near dam, Hawley Lake, White Mts.,
8200'.
U1694 n—16 Apache Co., Snake Cr., 19 mi. E. of McNary, White
Mts.
U1158 m—16 Apache Co., 3.8 mi. N. of Greer, White Mts., 8400’.
U1690 n»—16 Apache Co., 14 mi. W. of Alpine, White Mts., 8500'.
U1157 m—16 Greenlee Co., 6.6 mi. S. of Alpine, 7950’.
U1156 a—16 Greenlee Co., 10.3 mi. S. of Alpine, 7700’.
U1155 m^—16--1B Greenlee Co., 2.7 mi. N. of Hannagan Meadows,
White Mts., 8800'.
New Mexico:
U1152 5»—16 Catron Co., 10 1/2 mi. W. of Reserve, San Francisco
Mts., 7200'.
U1151 5—16 Catron Co. Tularcsa R., 1 mi. E. of Aragon, 6750’.
U1150 n»—16 Catron Co. Tularosa R., 3 mi. E. of Aragon, 6900’.
U1621 m»—16 Catron Co., Wall L., 9 mi. S. of Beavernead, Black
Range, 6400’,
U1619 m—16 Sierra Co. Poverty Can., 19 mi. N.W. of Winston,
Black Range, 7500'.
U1153 m—16 Grant Co., 4 mi. W. of Mule Cr., San Francisco Mts.,
5800’.
U1622 n—16 Grant Co. Black Canyon Forest Camp, 32 mi. N.
of Mimbres, Black Range, 6770’.
U1309 n=—16 Grant Co. 2.5 mi. N. of Pinos Altos, Pinos Altos
Range, 6600’.
U1310 n=16 Grant Co. Iron Cr. 6.6 mi. W. of Emory Pass,
Black Range, 6800’.
U1311 m—16 Grant Co., Iron Cr., 5.1 mi. W. of Emory Pass, 7200’.
U1314 5—16 Torrance Co., Red Canyon Forest Camp, 7 mi. W.
of Manzano, Manzano Mts., 7800'.
316 Rhodora [Vol. 74
U1148 »—16 Sandoval Co., Jemez R. at Soda Spring, 1.6 mi. N.
of Jemez Springs, 6350’.
C5281 n—16+3-6B Sandoval Co. 4 mi. N. of Battleship Rock,
near LaCueva, Jemez Mts., 7700’, (R. T. Clausen).
Ull47 n=16 (7 plants) »—16--1B (1 plant) Sandoval Co., La-
Cueva, (Probably same locality as preceding).
U1146 n—16 Sandoval Co. Jemez Falls, 7850'.
U1145 n—16 Sandoval Co., Las Conchas Forest Camp, Jemez Mts.,
8400'.
U1149 a—16 Sandoval Co., Las Huertas Cr., 6 mi. S. of Placitas,
Sandia Mts., 7830'.
C4633b n—16 (2 plants) Sandoval Co., Ellis Brook, N. side Sandia
Mts., (R. T. Clausen).
C4633a n=16 Sandoval Co., Sandia Crest, Sandia Mts., 10,400’,
(R. T. Clausen).
U1315 n—16 Santa Fe Co., 6 mi. N.E. of Santa Fe, 7500’.
U1316 n—16 Santa Fe Co., 9 mi. N.E. of Santa Fe, Black Canyon
Forest Camp, 8300'.
Ul1143 m—16 San Miguel Co., E. side Pecos R. at bridge 6.4 mi. N.
of Pecos, 7200'.
C5254 n—16 San Miguel Co., Pecos R., 5 mi. S. of Terrero, 7500’,
(R. T. Clausen).
Uli42 a—16 San Miguel Co. Gallinas Cr., 1/2 mi. W. of Mon-
tezuma, 6750’.
U1317 n=16 Rio Arriba Co. Rio Vallecitos, 5.5 mi. N. of La-
Madera, Ortega Mts., 6900’.
U1138 n=16 Taos Co., S. side Red River, 2.7 mi. E. of Questa,
7550’,
U1137 n=16 Taos Co. S. side of Red River, 4.5 mi. W. of Red
River village, 8150’.
U1136 n=i6 Taos Co., Mouth of Mallette Canyon, village of Red
River, 8650’.
U1140 n—16 Taos Co., S. side Rio Pueblo at Comales Forest
Camp, 7850’.
U1141 n—16 Taos Co. N. side Rio Pueblo, 2.2 mi. N.W. of Tres
Ritos, 8250'.
U1134 a—16 Colfax Co., 2.6 mi. E. of Red River Pass, Sangre de
Cristo Range, 8900'.
U1133 n»—16--4 Colfax Co., Cimarron Cr., 5 mi. E. of Eagle Nest,
7850’.
U1132 n=16 Colfax Co., Cimarron Canyon, 5 mi. W. of Ute Park,
7750’.
U1131 n=16 Colfax Co, Cimarron Cr. 1 mi. W. of Ute Park,
7550’.
1972] Sedum — Uhl 317
Texas:
U880 "—15 (2 plants) n=15+1 (1 plant) Jeff Davis Co., Ma-
dera Canyon, 25 mi. W.NW. of Fort Davis, Davis
Mts., 5800’. 1961.
U1633 m—15 Same locality, 1967.
Arizona:
U1685 752—329 1.6 mi. SE of Columbine Ranger Sta., Mt. Graham,
9215’.
U1684 m—29 Graham Co., Grant Creek, Swift Trail, Mt. Graham,
8750’.
U1683 n=29 Graham Co., Near Snow Flat, Swift Trail, Mt. Gra-
ham, 9000’.
U1686 n=29 Graham Co., 1.7 mi. N.W. of Ladybug Saddle, Swift
Trail Mt. Graham, 8800’.
U1682 mn—29 Graham Co. 1 mi. below Turkey Flat, Swift Trail,
Mt. Graham, 7300’.
U1926 ^—29 Graham Co. Marijilda Canyon, Mt. Graham, 8200',
(A. M. Phillips III).
U1927 m—29 Graham Co., Marijilda Canyon, Mt. Graham, 4800',
(A. M. Phillips III).
C5224 n=29 Cochise Co. S. of Rustler Park, Chiricahua Mts.,
8500', (R. T. Clausen).
UC65.144 n=29 Cochise Co., 1/2 mi. below Onion Saddle, Chiri-
cahua Mts., 7300', (D. Hutt 1260).
U1678 n=29 Cochise Co., S. Fork, Cave Cr, 3 mi. S.W. of
Portal, Chiricahua Mts., 5150’.
U1253 n=29 Cochise Co., Chiricahua Mts., (C. English).
U888 5»—29 (2 plants) Cochise Co., 1/2 mi. N. of Mule Pass,
Mule Mts., N. of Bisbee, 5840’.
New Mexico:
U1629 n=30 Lincoln Co., Bonito Cr., 7.2 mi. W. of Angus, Sierra
Blanea, 7500’.
U1628 n=30 Lincoln Co., 3.1 mi. W. of Angus, 7100’,
U1307 m—30 Lincoln Co. Eagle Cr., 2.5 mi. W. of Alto, Sierra
Blanca, 7600’,
U1304 n=30 Same, 1.5 mi. W. of Alto, 7450’.
U1630 m—30 Otero Co., Carrizo Cr., Mescalero Apache Indian Res.,
2.2 mi. W. of Ruidoso, Sierra Blanca, 7200'.
S. niveum Davidson
California:
C4759 n—16 San Bernardino Co., Sugar Loaf Peak, S. Bernardino
Mts., 7000', (R. T. Clausen).
C4759b »—16 Same, Dollar Lake, 9300', (R. T. Clausen).
318
U919
M345
M15346
M15333
M15682
Rhodora [Vol. 74
n—16 San Bernardino Co., above Jenks Lake, 8000’, (D.
Verity).
1n—16 Riverside Co., Santa Rosa Peak, Santa Rosa Mts.,
(P. A. Munz).
n—16 Riverside Co., 1 mi. N.W. of Toro Peak, Santa Rosa
Mts., 7450', (R. Moran).
Mexico:
n—64 Baja California, Observatory Peak, Sierra San
Pedro Martir, 9300', (R. Moran).
n—64 prob. Same, N. slope, 8400'. (R. Moran).
S. stelliforme S. Wats.
U1680
C5192
U1693
U1689
U1691
U1692
U2051
U1623
U1624
U1625
Arizona:
n—20+2-7B (5 plants) Cochise Co., N.E. slope Carr Peak,
Huachucha Mts., 8300. TOPOTYPE
n—20+2B Same Locality, (R. T. Clausen).
n—12 Apache Co., N. side Crescent L., White Mts., 9100’.
n—12 Apache Co., Cienega Redondo, 16 mi. W. of Alpine,
White Mts., 8950’.
n—12 Apache Co. 11/2 mi, W. of Three Forks, Black R.,
14 mi. W. of Alpine, 8550’.
n—12 Apache Co. Williams Valley, 7.2 mi. W. of Alpine,
8650’.
Mexico:
n—20--1 or 2B Chihuahua, 55 mi. W. of San Francisco del
Oro, (M. Kimnach and F. K. Brandt 1058).
New Mexico:
n—12 Grant Co., 1.4 mi. S. of Black Canyon Forest Camp,
Black Range, 7000’.
»—12 Grant Co., 2.6 mi. S. of same, 7300’.
n—12 (2 plants) Grant Co., 3.8 mi. N. of Rocky Canyon
Forest Camp, Black Range, 7800'.
U1625A n—12-4-8B Same locality.
U1626
U1627
U1312
U1313
U1620
n—12 Grant Co., 1.2 mi. N. of same, 7600’.
1—12 Grant Co., 2.9 mi. S. of same, 7400’.
n—12 Grant Co. 2.8 mi. W. of Emory Pass, Black Range,
7600’.
n—12 Sierra Co., 1 mi. N. of Emory Pass, Black Range,
9000’.
n—12 Sierra Co., N. side of Boiler Peak, 3.2 mi. W. of
Continental Divide, Black Range, 7900’.
S. wrightii A. Gray
U1631
New Mexico:
n—12 Otero Co., Silver Springs Canyon, 7.8 mi. N.E. of
Clouderoft, Sacramento Mts., 8000’.
1972] Sedum — Uhl 319
U1303 n=12 Otero Co., Silver Springs Canyon, 5 mi. N.E. of
Cloudcroft, 8400’.
U1302 n=12 Otero Co., Cox Canyon, 5.2 mi. S.E. of Cloudcroft,
8150’.
U1632 n=12 Otero Co. Upper Penasco Canyon, 13 1/2 mi. S. of
Cloudcroft, 8400’.
C5241 n=12 Dona Ana Co, Canyon at La Cueva, Organ Mts.,
(R. T. Clausen).
Texas:
U882 n—24 El Paso Co, McKelligon Canyon, N. of El Paso, S.
side of Franklin Mts., 5400’, TOPOTYPE.
C364 . n—12 Jeff Davis Co., Fern Canyon, Davis Mts.
UC52.1680 n—13 Jeff Davis Co., 20 mi. N. of Alpine, Davis Mts.,
4900', (E. R. Blakely 1378).
U879 n—12 (4 plants) Jeff Davis Co. 5 mi. S. of Fort Davis,
Davis Mts., 4800', (Same locality as preceding?).
U878 «—12 Brewster Co., Sunny Glen, 6 mi. W.NW. of Alpine,
4800'.
U1676 m—24 Brewster Co., 3.4 mi. S. of US 90 at Marathon, 4100'.
U877, U1675 n=36 Brewster Co. Green Gulch, N. of Panther
Gap, Chisos Mts., Big Bend National Park, 5600’.
U873 n—36 Upper Boot Canyon, Chisos Mts., 6900’.
UC52.1672 m—36 E. side of Emory Peak, Chisos Mts., 7000’, (E.
R. Blakely 1402).
C5298 »n—36 Chinese Wall, Chisos Mts., (R. T. Clausen).
U868 n—36 Val Verde Co., E. side of Devils River, 8 mi. N.W.
of Del Rio, 1100’.
UC54.1334 n=36 Same locality, (J. B. Perry).
Mexico:
UC53.407 n—24 Coahuila, 8 mi. S.W. of Piedra Blanca, Sierra del
Carmen, 7000’, (Ward Russell).
U1497 m—12 Coahuila, Chorro Canyon, 27 km. S.E. of Saltillo, ca.
6500'.
U1501 n=12 Coahuila, Chorro Canyon, 33 km. S.E. of Saltillo, ca.
1000'.
M7825 m—12 Coahuila, Chorro Canyon, (R. Moran).
M6317 n=48 Nuevo Leon, San Jose de Raices, (R. Moran).
U1494 n=45-48 Nuevo Leon, 6 km. S. of San Jose de Raices,
6100’.
U1916 n—48 Nuevo Leon, 3 km. W. of Iturbide, 5400’.
U1914 n—24 Nuevo Leon, 16 mi. N. of Ascension, 7900’.
U1912 n=?24 Nuevo Leon, 30 mi. N. of Doctor Arroyo, 4 mi. N.
of La Escondida, 5900'.
320 Rhodora [Vol. 74
Sedum havardii Rose
Texas:
U875 n—54 Brewster Co. Chisos Mts., Big Bend National Park:
N. of Boot Spring, 6900’.
U876 n—81 11/4 mi. S. of Boot Spring, 7000’.
UC52.1679 n—54 E. side of Emory Peak, 7000’, (E. R. Blakely).
U1674 m—54 Above E. side of Panther Pass, N. side of Casa
Grande, 6200'.
C363 n=54 Green Gulch, 5500’.
LITERATURE CITED
BALDWIN, J. T., Jr. 1936. Polyploidy in Sedum ternatum. J.
Hered. 27: 241-248.
CLAUSEN, R. T., and C. H. UHL. 1943. Revision of Sedum cocker-
ellii and related species. Brittonia 5: 33-46.
Love, A. 1951. Taxonomical evaluation of polyploids. Caryologia
3: 263-284.
Moran, R. 1969. Sedum in Baja California. J. Cactus & Succu-
lent Soc. Am. 41: 20-25.
UHL, C. H. 1961. Some cytotaxonomic problems in the Crassula-
ceae. Evolution 15: 375-877.
1970a. Chromosomes of Graptopetalum and Thomp-
sonella (Crassulaceae). Am. J. Bot. 57: 1115-1121.
1970b. Heteroploidy in Sedum glaucophyllum. Rho-
dora 72: 460-479.
DIVISION OF BIOLOGY
CORNELL UNIVERSITY
ITHACA, NEW YORK 14850
THE RACHILLA IN CAPE COD SPECIES OF CAREX
WITH NOTES ON THE HISTORY OF
THE PERIGYNIUM AND RACHILLA
HENRY K. SVENSON
This account takes up the local occurrence of the pecu-
liar bristle-like or spikelet-bearing structure within the
perigynium of Carex, known as the “rachilla”. There is
a review of various opinions, especially those of Duval-
Jouve, Holttum, and Raymond. There is also a discussion
of the relationship of Cymophyllus to Carex and to the
West Indian Bisboeckelera. A plate with many figures il-
lustrates the various interpretations of the rachilla.
It was a surprise to find on Cape Cod, in southeastern
Massachusetts, Carex with development of the rachilla.
First detected was Carex Emmonsii, which appeared pecu-
liar to the naked eye. It grew in rich soil with skunk
cabbage along the so-called Marston Mills River. The micro-
scope showed that spikelets, developing within the perigy-
nium, had extended out of it, a characteristic of Schoenozi-
phium. This is a genus of sedges mainly South African.
Further observation showed that this condition is not rare
in Carex Emmonsii in dry sandy fields. C. Emmonsii is
the commonest Carex on Cape Cod, and easily recognized
by the tufts of long recurving leaves, which remain over
winter. It grows especially well where the soil has been
disturbed. The same rachilla structure is found, though
less abundantly, in robust specimens of the closely related
Caiex pensylvanica. In a sandy field in Osterville, both
species, together with rachilla-bearing C. scoparia, grew in
a field being overrun by Robinia viscosa, Prunus serotina,
and Ewonymus europaea. Further good examples of C.
pensylvanica were found in sandy fill near Johns Pond in
Mashpee.
This axis within the perigynium assumes various forms,
most commonly as a small bristle, but also as the schoenoxi-
phoid condition mentioned above (see fig. 7). On Cape Cod
I have seen the rachilla in C. scoparia, C. Swanii, C. debilis
321
322 Rhodora [Vol. 74
1972] Carex — Svenson 323
var. Rudgei, C. bullata, C. canescens, C. Emmonsii and C.
pensylvanica. There are undoubtedly more. The figure of
C. complanata, collected by me in Tennessee, was drawn by
Miss Purdy about 1940. It has a rachilla reminiscent of a
hookless Uncinia.
The rachilla in Carex was first noted by Kunth (1788-
1850), who correctly interpreted it. The problem is illus-
trated in figs. 11-15, where rachillas are shown in the series
Schoenoxiphium to Carex, based on figures from Kukenthal
(1909), Das Pflanzenreich: Cyperaceae-Caricoideae; and
from R. S. Snell (1936). Anatomy of the spikelets and
flowers of Carex, Kobresia, and Uncinia, in Bull. Torrey
Club 63: 277-295. In Carex, as in fig. 15, the rachilla is
commonly reduced; but see fig. 4, C. complanata, and fig.
5, C. scoparia. There are many examples in Boott’s (1858),
“Illustrations of the Genus Carex". In many of his illus-
trations the rachilla is flat with a serrated margin. That
of C. Fraseri is strap-shaped without serrations. That of
S. scoparia (my fig. 5) is perhaps closest to C. Camposii
(t. 574) from the Sierra Nevada of Spain, which Kueken-
Figs. 1-17. The rachilla (except in 4) is black. Axis of spikelet,
a; scale (glume, b; perigynium, c; achene, d. Fig. 1. Uncinia unci-
noides, X 10, Hawaii, Svenson, Sept. 5, 1960; 2. style-base enlarged.
3. Rachlla of Carex tonsa, Ostervile, Mass., Svenson, June 10,
1967. 4. Carex complanata, X 10, Altamont, Tennessee, Svenson,
Oct. 3, 19835 (drawn by Maud H. Purdy). 5. Carex scoparia, X 10,
Osterville, Svenson 2933, June 23, 1970. 6. Carex pallescens, X 5,
from Duval-Jouve, pl. vii, f. 8. 7. Carex Emmonsii, X 5, Svenson,
June 16, 1967 (lowest scales removed). 8. Same collection as 7 (all
scales removed). 9. Carex pallescens, X 10, from Duval-Jouve, pl.
vii, f. 9 (perigynium bisected longitudinally). 10. Carex debilis var.
Rudgei, Osterville, Svenson, Aug. 18, 1967. 11-15. Diagrammatic
sections to show rachilla of 11, Schoenoxiphium; 12. Cobresia; 13.
Uncinia; 14. Carex microglochin; 15. Carex (with reduced rachilla) ;
all from Snell (1936) and Kuekenthal (1909). 16. Schoenoxiphium
rufum, X 6, from C. B. Clarke, t. 141, f.2 17. The same, with peri-
gynium removed, f. 3.
Dissections and samples of material from Cape Cod are filed on
a sheet in the cover of Carex Emmonsii from Barnstable County in
the herbarium of the New England Botanica] Club.
324 Rhodora [Vol. 74
thal places in the section Elatae, not represented in Amer-
ica. But none of Boott's plates illustrate a rachilla in the
Ovales. By Barros (1935), Cyperaceae of Argentina, in
Anales Mus. Ciencias Nat. Hist., the rachilla is shown in
4 species.
The best account of these contradictory structures is by
Duval-Jouve (1864): Sur la signification morphologique
d'une anomalie qué présentent les utricules de quelques
Carex, in Bull. Soc. Bot. France. 1864. 19 pp. 1 pl. Since
the structures are similar to those that I have found, and
since many species treated by him are also in New England,
I have given his account the most extended treatment. In
the spring of 1864, Duval-Jouve first found Carex palles-
cens with peculiar compound spikes, and spikelets issuing
from the perigynia. Altogether he records a list of 24
species with a rachilla, obtained from the vicinity of Arles
and Strasbourg. The anomalous spikelets were found most
frequently toward the base of the spike, sometimes in the
middle (C. acuta, C. pseudocyperus, but only rarely toward
the apex. In C. flacca?, C. Oederi, C. flava, and C. hirta,
compound spikelets were so abundant that in some local-
ities it was almost impossible to find a plant without them.
In C. flacca the rachilla appeared most often as the Schoe-
noxiphium-type. In C. flava the perigynia were often split,
'N. J. Duval-Jouve (1810-1883). For biography see Flahault, Bull.
Soc. Bot. France. His work is typical of the best of French botanists.
He is perhaps best known for his work, illustrated in color, of Equi-
setum hiemale. Allis as though inlaid with gold. His work on Carex
is equally as good. As a forerunner of botanical work he wrote a
"Traite de Logique, ou Essai sur la theorie de la Science", Paris
(1848), and a second edition, which I have seen, in 1885. It treats
of various kinds of observations, perceptions, evidence. etc., things
which often today tend to be forgotten. Surely no other botanist was
so well prepared for his scientifie task. I did not, however, find a
history of the transition of nomenclature from logie to biology, in
which I was particularly interested.
"Now a very abundant and variable species throughout the gypsi-
ferous region (about Windsor), often growing in grassy pastures
near the gypsum cliffs. Roland, Fl. Nova Scotia (1947), p. 230.
1972] Carex — Svenson 325
resembling Elyna spicata. All species were in places highly
modified by pastures, ditches, animals, and in many other
ways. They were best developed on terrain newly dis-
turbed, or where traces of modification were apparent. On
Cape Cod, Carex pallescens? is common in grassy places,
but I have yet to find a specimen with a rachilla.
The achene in Carex represents the lowest element of a
reduced axillary inflorescence, and the achene occupies a
basal lateral position at the lower part of the perigynium.
This axis (the rachilla) when present is commonly bris-
tle-ike, or it may have several flowers as in Schoenozi-
phiwm. Presence and development of the rachilla and the
origin of the perigynium have been the subject of intensive
study — one of the unfilled crossword spaces in the Cypera-
ceous organization. It is generally conceded that the perigy-
nium consists of a single scale, and not the union of two
scales. Torrey, in North American Cyperaceae (1838), p.
404, noted in a specimen of C. Willdenowii that the perigy-
nium of the lowest flower enclosed the peduncle of a sep-
arate staminate spike. The two remaining pistillate flowers
had within the perigynium, in place of a staminate spike,
a green squamaceous rudiment. “This curious plant, there-
fore, seems in a manner to connect Schoenoxiphium N. ab
E. with Uncinia, and to explain the nature of the setaceous
body in the latter genus".
Theodore Holm (1854-1932) provides a bibliography of
55 items on the perigynium, together with an excellent
account of terminology, in Am. Journ. Sci. ser. II. 214-
220 (1896) entitled "The cladoprophyllum and anthopro-
"The var. neogaea Fernald (Rhodora 44: 306. 1942) is supposed
to be without a short beak at the apex of the perigynium; but Boott
(1858, pp. 39, 40) says the beak may be present or absent, and he
shows a beak in t. 450, illustrating a specimen from Rhode Island.
Kuekenthal (1909, p. 433) describes the perigynium as erostrate ex-
cept in var. glaberrima which has “utriculae interdum brevissime
rostratae”. The American var. neogaea therefore does not appear to
amount to much. On Cape Cod, and otherwise in Massachusetts as
I have seen it, Carex pallescens does not have the appearance of a
native species.
326 Rhodora [Vol. 74
phyllum in Carex”, and the accompanying plate shows
schoenoxiphoid spikelets in C. crinita and C. Sprengelii.
Holm was one of the ablest of American botanists, who
combined precise taxonomy with morphology; he was also
an excellent artist. One of the latest accounts is by Holt-
tum: “The spikelet in Cyperaceae” in Bot. Review, vol
14 (no. 8) 525-541. 1948. He, likewise, reviews the history
of the perigynium. Much of the interest in the subject
comes from the fact that Carex is one of the world’s largest
genera, and because it is abundant in the north temperate
region. 267 species are listed in Gray Man. ed. 8; 793 spe-
cies in Kuekenthal’s world-wide treatment (1909).
The development of the perigynium is almost like a sec-
ond evolution of the seed plants. The developing embryo
has been protected within the perigynium. Dispersal of
mature perigynia is by floating, flying, attachment to men
or animals, or in many other ways. This protection has
emancipated Carex from the tropics, which (according to
Holttum and to Raymond) is its primitive home, and al-
lowed its penetration into areas freed from the Pleistocene
ice. Raymond (1959): Carices indochinenses necnon
siamenses. Mém. Jard. Bot. Montreal, no. 53, mentions the
Careyanae and Pandophyllae as perhaps the most primitive
elements of Carex. The Careyanae were segregated by him
from the Laziflorae, which are characteristic sedges of the
Appalachian forest of southeastern United States.
As Fernald says (Rhodora 5: 249. 1903) the genera of
the Cariceae are not very good: “Besides the genus Carex
the members of the Cariceae have been grouped by differ-
ent modern authors into various ill-defined genera varying
with the personal equation". But that is also true of other
genera in the Cyperaceae. The presence of the rachilla in
widely diverse species of Carex led Duval-Jouve to suspect
that the rachilla might not be of phylogenetic significance,
but merely atavistic remnants appearing sporadically.
Thus the occurrence of the rachilla in many species of
Primocarex led to the early idea that Primocarex was prim-
9
1972] Carex — Svenson 327
itive. It is interesting in this respect to compare the rachilla
of C. complanata (fig. 4) with Uncinia (fig. 1).
Of interest is the lack of an articulation or joint in the
style of the Vesiculares (in which the perigynium is highly
adapted for floating), and in the Ovales* (in which it is
wind-distributed). The Ovales have their great distribu-
tion on the Atlantic coastal plain and in Pacific North
America, cf. chart by Kuekenthal (1909) p. 14, and the
recent fine treatment by Hermann (1970): “Carices of
the Rocky Mountains and Colorado Basin". Nelmes: “Facts
and Speculations on phylogeny in the tribe Cariceae" in
Kew Bull: (1961) 427-436, explained that some unispicate
species were of uncinoid origin, and he distributed most of
Kuekenthal's Primocarex variously in the genus. And to
emphasize the ill-defined genera, Kuekenthal (1940) de-
scribed Schoenoxiphium kobresioides from material col-
lected by Van Steenis in Sumatra, and termed it the “Ver-
bindungslied zwischen Schoenoxiphium und Kobresia".
Holttum thought Mapania to be the most primitive genus.
See also my comments on Mapania macrophylla in Bull.
Torr. Club 75: 95. 1948, with fig. 3. The breakdown of the
tubular translucent corolla provides us with a possible ori-
gin of the perigonial bristles in the Cyperaceae. I was incor-
rect in ascribing M. macrophylla to Santo Domingo as well
as British Guiana. Meyer’s plants all came from the Es-
sequibo River in British Guiana.
Mackenzie set up Cymophyllus, based on Carex Fraseri.
By Pursh it was considered a Mapania, but I believe it to
be a true Carex, illustrated by Boott (t 484) with a strap-
shaped rachilla. There is a reduction series in the West
Indies from Mapania and Hypolytrum to Bisboeckelera
(Hoppia) and Calyptrocarya, which simulates Carex; but
I think it is an unrelated parallel series, as shown by the
various illustrations in Clarke’s “Illustrations of Cypera-
*The style is said to be jointed by Mackenzie, and is so illustrated
in North American Cariceae (1940), but I have not found it so. See
fig. 5).
328 Rhodora [Vol. 74
ceae" (1909). In my illustration of Mapania sylvatica, the
type of the genus, in “Cyperaceae of Panama", the corolla
segments are inrolled around the stamens. This I am sure
is the correct interpretation, and the flower is a normal
flower of the Cyperaceae, and not an assemblage of “‘par-
tial inflorescences”. (Ann. Missouri Bot. Gard., 1943)
Another interesting structure in Carex is the style-base,
which is sometimes swollen and articulated. Duval-Jouve
described C. oedipostyla, i.e. with a “swollen foot” in Mém.
Acad. Sci et Lettres de Montpellier (1870), including a
plate with many figures. The articulating style-base is wide-
spread in Eleocharis and some other genera, but has little
real significance in their delineation. It is one of the his-
torical straws grasped by the drowning botanist. Some
decades ago I thought that the swollen style-base in the
Laxiflorae might be a transition to broadleaved Cyperus
(Mariscus) of the tropical rain forest, and especially to
Cyperus diffusus. The structure of the achene and prophyl-
lum in Carex (including perhaps Schoenoxiphium), is
closer to Cyperus than to any other genus. In visiting Nic-
aragua last spring, I found Cyperus diffusus to be a road-
side and plantation weed, and not a rain forest type. It
is probably different in the Old World.
Schoenoxiphium is generally considered to be the most
primitive of the Caricoideae, and the accompanying figures
from Kuekenthal, Snell, and Clarke show the ideas pre-
sented. It is a small genus centering on South Africa, and
extending to the mountains of Madagascar, southern Si-
beria just north of Afghanistan (Krechetovich in Flora
Tadzhikskoi USSR 2: 10-12. 1963), and to Sumatra. D.
M. Napper, in Journ. East Afr. Hist. Soc. and Corydon
Museum, Nairobi, 24 (no. 2); 1963, p. 12, says *predomi-
nantly South African, about 15 species. The normal habi-
tats are damp forests and wooded or open grasslands. It
is unusual for Schoenoxiphium to occur at the higher alti-
tudes (over 7,000 or 8,000 ft.) where Carex usually oc-
curs”.
1972] Carex — Svenson 329
The Caricoideae include Schoenoxiphium Nees (1832),
Kobresia® Willd. (1809), Elyna Schrader (1806), Uncinia
Persoon (1807), and Carex? Linnaeus (1753).
Kobresia has according to Kukenthal (1909) its origin
and central point in the high mountains of central Asia
from Turkestan to Central China. 14 of the species are
peculiar to the Himalayas, especially the western part, and
5 to the high mountains of Central China. The few North
American species are well treated (with photographs) by
Dunan in Bull. Torrey Club 83: 192-195. 1956; and in K.
simpliuscula rachillas are shown. According to Mackenzie,
Kobresia extends from Newfoundland and Greenland to
Colorado and Alaska. K. elachycarpa Fernald was de-
scribed in 1897 from banks of the Aroostook River at Fort
Fairfield in Maine, but later treated by him as Carex elachy-
carpa, which goes to show how tenuous are the limits of
these genera.
Uncinia has been well treated by Kuekenthal (1909) and
consists of 24 species mainly of the southern hemisphere
from the Antarctic Islands through the South American
Andes to Mexico and Jamaica. The remainder is in Aus-
tralia, Tasmania, New Zealand, New Guinea, and the
Hawaiian Islands. The illustrated specimen (fig. 2) was
collected by me in the Hawaiian Islands.
Carex microglochin Wahlenberg is illustrated in fig. 4.
It extends from Greenland to Alaska, southward to Quebec
and Colorado. It is also known in the Patagonian region
and in Eurasia. In describing the occurrence in Newfound-
land, Fernald in Rhodora 28: 61. 1926, says “It is one of
the rarest and taxonomically most interesting of species,
having the primitive rachilla, which is obsolete in most
"According to Kuekenthal, named for the botanophile Paul de
Cobres of Augsburg. It is sometimes spelled with a “C”.
*Duval-Jouve (1864, footnote p. 16): The word Carex was used
by Virgil, Columella, Calpurnia, etc., to designate hard, sharp, or
cutting herbs . . . perhaps tall Juncus, Typha, or Sparganiwm, or
perhaps dry thickets at the borders of pastures, i.e. “Frondibus hir-
sutis et carice pastus acuta" thickets which the shepherds of Proven-
cal still call carigues.
330 Rhodora [Vol. 74
northern Carices, projecting from the top of the perigy-
nium; in other words, C. microglochin is an excessively
local relic-species now known only in a few remote corners
of the globe, and about as well placed in the Antarctic
genus Uncinia as in the genus Carex, which abounds in
the northern hemisphere . . . its prickly little fruit had
obviously been spread by pedestrians with their high skin
boots."
Now, with all these contradictory views, and the ques-
tionable relationship of the various elements, it is hoped
that the reader will go out to collect some rachilla mate-
rial for himself, and will then turn to Fernald's amusing
story of Carex microglochin in the twenty-eighth volume
of Rhodora.
OSTERVILLE, MASSACHUSETTS 02655
DISTRIBUTION OF NATIVE AND
INTRODUCED KNAPWEEDS (CENTAUREA)
IN CANADA AND THE UNITED STATES!
R. J. MOORE
The genus Centaurea (family Compositae, tribe Car-
dueae) is a large and taxonomically complex group of at
least 500 species, centered in the Mediterranean region.
The great majority of species are Eurasiatic but two spe-
cies are native to the southwestern United States and a few
others are native in northern South America. A number
of Old World species are naturalized, to varying degrees,
in Canada and the United States.
During the course of studies of the species found in
Canada, data concerning the genus in the United States
have been brought together from the literature and from
herbarium collections examined. Floras of various areas
of the United States treat the species found in their range
but there is no complete inventory of United States spe-
cies. Moreover, there is obvious disagreement between au-
thors regarding the nomenclature and taxonomy of certain
species and uncertainty as to the occurrence of a few spe-
cies in the country. Boivin (1966-67) has listed the spe-
cies found in Canada and we have discussed the morphol-
ogy and distribution of these species in a study of the tribe
Cardueae (Moore and Frankton, 1972).
In the following key is given the diagnostic characters
of the taxa which have been reported or have been found
to occur in Canada and the United States. Brief notes re-
garding taxonomy, common synonymy and a summary of
the distribution are added.
The identification and distribution of Canadian plants
have been verified from herbarium collections, notably
those of the following institutions: Department of Agri-
culture, Ottawa (DAO); National Museum of Canada
(CAN); Montreal Botanic Garden (MTJB) ; Botanical In-
stitute , University of Montreal (MT) ; University of Brit-
‘Contribution no. 892 of the Plant Research Institute.
331
332 Rhodora [Vol. 74
ish Columbia (UBC); British Columbia Provincial Mu-
seum, Victoria (V). Information regarding United States
species has been compiled from the commonly known re-
cent floras, some of which are cited here, and from her-
barium specimens examined in the course of studies of the
Cardueae of Canada. Specimens have been borrowed from
the Gray Herbarium of Harvard University (GH), New
York Botanical Garden (NY), and the University of Ver-
mont (VT) to resolve certain specific problems. Verifica-
tion of United States records which appear to be credible
has not been attempted. The various North American in-
troductions have been compared with European specimens
of the taxa in our herbarium (DAO). Specimens of all
the taxa discussed here have been seen, but in the cases
of a few rarer species (C. eriophora, C. diluta, C. nervosa)
only Old World specimens were available for study.
Since most of the species are adequately described in
available floras, descriptive comments are here appended
only in cases where there is disagreement with the con-
cepts expressed by most authors or to emphasize points of
distinction between species.
The genus Centaurea is here treated sensu lato, as is the
most common practice in North America. As indicated by
synonymy, some of the species are often placed in segre-
gate genera.
DESCRIPTIVE CHARACTERS
Some discussion of the characters used in the following
key may be helpful. The phyllaries (involucral bracts)
take many forms in the genus and are very useful as diag-
nostic characters since they are present and easily exam-
ined in all flowering specimens. The innermost row of
phyllaries is frequently unlike the outer rows and descrip-
tions here refer to the more conspicuous outer 2 or 3 rows,
unless otherwise stated. An apical spine is considered to
be a rigid pointed apical structure that is distinctly longer
than any lateral points that may be present on the phyl-
lary. Phyllary margins (as distinct from the phyllary
1972] Centaurea — R. J. Moore 333
apex) may also bear spines, which are pointed rigid struc-
tures, unlike the wiry, more or less flexible marginal proc-
esses seen in the condition described as pectinate, or the
serrations of membranaceous or chartaceous margins. In
many species the apical portion of the phyllary is expanded
into a structure (appendage) that is distinctly broader
than the phyllary base (blade). In our species the appen-
dage is either pectinate or chartaceous and with entire or
lacerate margins. The flower heads may consist wholly of
perfect tubular florets, all similar in size, or of shorter,
perfect central florets surrounded by a peripheral row of
longer, sterile tubular florets, which simulate ray florets.
In the key, these conditions are described as eradiate (all
florets similar) or radiate (short central and longer periph-
eral florets — actually a falsely-radiate condition).
KEY TO SPECIES
a. Outer and mid involucral phyllaries terminated by a
distinct spine (very short in C. salmantica and C. di-
luta) ; with or without marginal spines as well. (b)
b. Marginal spines or processes absent. (c)
c. Apical spines more than 5 mm long (usually 10-
25 mm). (d)
d. Stems winged (decurrent leaf bases) ; flowers
yellow (Sect. Mesocentron DC.) (e)
e. Apical spines pinnately spinose (bearing
several lateral spinules on the shaft);
phyllaries arachnoid. (f)
f. Apical spines 5-10 mm long; involucre
about 15 mm high 1. C. melitensis
f. Apical spines more than 10 mm long;
involucre approx. 20 mm high ......
p ) 2. C. eriophora
e. Apical spine palmately spinose (without
spinules on the shaft but with 2-5 pairs
of palmately arranged basal spinules) ;
phyllaries glabrous. (g)
g. Heads 35-40 mm high, involucre 20-
334
Rhodora [Vol. 74
25 mm high, 15 mm broad; tips of in-
ner phyllaries not dilated; achenes
approx. 4.5 X 3 mm; pappus dark
brown ............. 3. C. sulphurea
£ Heads 20 mm high, involucre 15 mm
high, 10 mm broad; tips of inner phyl-
laries chartaceous and dilated; achenes
approx. 3 X 1.5 mm; pappus white .
MEM" 4. C. solstitialis
d. Stems not winged; flowers pink to purple
(Sect. Calcitrapa Cass.). (h)
h. Involucres longer than broad (length 12-
17 mm; L/W approx. 2.5) ; heads sessile
or subsessile; pappus absent ..........
Lecce eee eee eee eae 5. C. calcitrapa
h. Involucre globular, about as long as
broad (length 11-14 mm) ; heads pedicel-
late (pedicels 10-50 mm long); pappus
about 2 mm long ......... 6. C. iberica
c. Apical spines 1-5 mm long (i)
i. Apical spines 3-5 mm; phyllaries sericeous,
margin dark pigmented .... 7. C. muricata
i. Apical spines 1-2 mm; phyllaries glabrate,
margins not pigmented. (j)
j. Phyllary tips dark, terminated by a 1 mm
Spine or sometimes merely mucronate;
margin entire ......... 8. C. salmantica
j. Phyllary tips not pigmented, with a 1-2
mm spine and a small chartaceous, pal-
mate-lacerate appendage; margin narrow-
ly scarious ... 9. C. diluta
b. Marginal spines or processes (at least 4 pairs) pres-
ent, in addition to the apical spine (Sect. Acrolo-
phus Cass.) (k)
k. Heads radiate (but peripheral florets only slight-
ly larger than the inner florets) ; marginal proc-
esses straight or curved and somewhat flexible,
1972]
Centaurea — R. J. Moore 335
phyllary tip and margin pigmented, forming a
dark V-shaped mark. (1)
l. Involucres 6-8 mm broad, globose and rounded
at the base; heads borne in corymbs or
corymbose panicles, the branches not strik-
ingly straight and stiff in appearance .
uri AG c GA, zu 10. C. maculosa
l. Involucre 4-6 mm broad, distinctly longer than
broad and the base usually tapering; heads
borne in panicles on stiffly diverging branches
Sp ass i oH cs DE 11. C. paniculata
Heads eradiate; marginal spines straight and
stiff, phyllary margins not pigmented in a V-
shaped marking but occasionally with a dark ir-
regular apical mark. (m)
m. Heads 14-16 mm high; achenes 2.3-2.5 mm
long, pappus absent or short (1 mm) ; flowers
usually white but sometimes yellow, pink or
mauve no ui M ME 12. C. diffusa
m. Heads 12-13 mm high; achenes 2.5-3 mm
long; pappus present (to 2.5 mm); flowers
pink or maüve — 10 5 b SIL Les VAS
M a 4 13. C. virgata var. squarrosa
a. Outer phyllaries not terminated by a conspicuous, stiff
spine which is longer and more prominent than any
marginal processes (margins pectinate, lacerate or en-
tire)
(n)
n. Phyllaries bearing an apical appendage (an abrupt
expansion of the phyllary blade, broader than the
base of the blade) (o)
o. Heads small (to 4 cm broad) (p)
p. Involucres approximately as long as broad;
phyllary appendages broad and completely
covering the bases of adjacent phyllaries (q)
q. Appendages brown and papery, with lac-
erate margins ............ 14. C. jacea
q. At least some of the appendages pectinate
336
Rhodora [Vol. 74
(wiry marginal processes), appendages
black or dark brown (r)
r. Apex of appendages extended into a
very long, recurved process and with
numerous long lateral processes, feath-
er-like in appearance (s)
S.
Plants rarely branched, bases of
upper leaves truncate and almost
clasping; involucres 20-24 mm
high; achenes 4-5 mm, pappus 2.5-
3 mm long .. . 15. C. nervosa
Plants usually branched; leaf bases
tapering; involucres about 15 mm
long; achenes 3-3.5 mm long, pap-
pus 1 mm . 16. C. austriaca
r. Apex of appendages appressed and not
extended into a long recurved tip (t)
t.
t.
Heads eradiate; pappus present
(short, 0.5 mm but fully devel-
oped); appendages black, wiry
processes long, 2-3 X width of the
blade, completely covering the ad-
jacent phyllaries ... 17. C. nigra
Heads eradiate or radiate; pappus
absent or vestigial (few short
bristles) ; appendages brown (not
dark brown) and variable from
pectinate to chartaceous-lacerate,
appendages broad, covering adja-
cent phyllaries . ..........
Lee eee 18. C. X pratensis
p. Involucre longer than broad (L/W = 1.5);
phyllary appendages smaller and not com-
pletely obscuring adjacent phyllaries (u)
u. Appendages black or dark brown, small
and usually triangular, not concealing the
contrasting green phyllary blades; rameal
1972]
Centaurea — R. J. Moore 337
leaves narrow and smaller than the cauline
leaves Durum cnt 19. C. nigrescens
u. Appendages dark brown, to twice the
width of the phyllary base and almost
covering adjacent phyllary bases; rameal
leaves not clearly differing in size from
the cauline leaves ........ 20. C. dubia
o. Heads large (over 4 cm broad) (v)
v. Phyllary appendages chartaceous and pal-
mately lacerate; heads eradiate, flowers yel-
low .. i . 21. C. macrocephala
v. Appendages pinnately pectinate; heads radi-
ate, flowers purplish (w)
w. Phyllaries brown, appendages bearing 8-
12 pairs of cilia .. ... 22. C. rothrockii
w. Phyllaries straw-colored and sometimes
purple-tinged, appendages bearing 3-8
pairs of cia. 0 23. C. americana
n. Phyllaries lacking an apical appendage (x)
x. Phyllary margins papery, either entire or lacerate
but not pectinate (y)
y.
Phyllary margins essentially entire, not lacerate
(z)
z. Heads to 2 cm high, eradiate .. 24. C. repens
z. Heads to 4 cm high, radiate . 25. C. moschata
Phyllary margins regularly dentate-lacerate (aa)
(Sect. Cyanus DC.)
aa. Leaves elliptie, oblanceolate, 10 mm or more
wide; heads over 3 cm broad ............
NIRE sesto ae Oe. DE MOntAanNa
aa. DT linear, lanceolate, to 3 mm wide;
heads to 3 cm broad ... 21. C. cyanus
x. Phyllaries pectinate (with wiry marginal processes)
(Sect. Lopholoma Cass.) (bb)
bb. Heads eradiate; foliage hoary .... .....
. 28. C. cineraria
bb. Hos radiate; tod glabrous s
29. C. scabiosa
io; oe Kee sie CAR RAN tel. ie: 6 vw. dps ea WW eal ee MP.
338
1.
Rhodora [Vol. 74
NOTES ON SPECIES AND THEIR DISTRIBUTION
C. melitensis L. Maltese Centaury, Tocalote, Croix de
Malte; common: U. S.— west coast, Ariz., N. M.,
Texas, established in the western states, occasional
and perhaps not persisting in the central and eastern
states (Steyermark, 1963; Fernald, 1950) ; Canada —
Vancouver Is., B. C.
C. eriophora L. Rare: U. S.— Cal. Only one 1909
collection from Los Angeles is reported (Howell,
1959).
C. sulphurea Willd. (C. sicula of Cal. authors, accord-
ing to Howell (1959) ; rare: U. S. — Cal.
C. solstitialis L. Barnaby's Thistle, Yellow Star-this-
tle, Centaurée du Solstice; common: U. S. — probably
in all states, weedy in the southwest; Canada — sw
Ont., Man., Sask.
C. calcitrapa L. Purple Star-thistle, Caltrope; com-
mon: U. S.— southeastern states to N. Y. (Fernald,
1950), west coast (Abrams & Ferris, 1960), Ariz.
(Howell, 1959), Ill. (Jones & Fuller, 1955) ; Canada
— sw Ont., Vancouver Is., B. C.
Howell (1959) records that a similar plant, possibly
to be called C. calcitrapoides L., had been collected in
Riverside Co., Cal. We have a collection from Hum-
boldt Co., Cal. (Tracy 16769 — DAO) which may be C.
caleitrapoides, since a sparse pappus is present. This
species differs from C. calcitrapa in the taller habit
and the presence of a short pappus.
C. iberica Trev. in Spreng. Iberian Star-thistle;
rare?: U. S. — Cal. Howell (1959) reports this spe-
cies as rare in 3 counties of Cal., first collected in 1924.
Several west coast floras list it for Cal. only.
This species is similar to C. calcitrapa and speci-
mens identified as C. calcitrapa may be C. iberica. Two
such cases have been revised: Soil Conservation Col-
1972] Centaurea — R. J. Moore 339
lection, Sept. 10, 1955, Converse Co., Wyo. — DAO;
Jones s.n., July 13, 1946, Labette Co., Kans. — DAO).
To our knowledge, C. iberica has not been reported in
Kansas or Wyoming. Iberian Star-thistle may occur
in states other than California, in ranges reported for
C. calcitrapa.
7. C. muricata L. (Amberboa muricata (L.) DC., Vol-
untarella muricata (L.) Benth. & Hook.) ; rare: U. S.
— Santa Barbara, Cal.
8. C. salmantica L. (Microlonchus salmanticus (L.) DC.,
Mantisalea salmantica (L.) Briq. & Cavill.); rare:
U. S. — Cal. (Howell, 1959), Ariz., (Kearney & Peeb-
les, 1960)
9. C. diluta Ait. Rare: U. S. — Cal.
10. C. maculosa Lam. Spotted Knapweed, Centaurée;
common: U. S. — all states except possibly the south-
eastern; Canada — N. S., N. B., Que., Ont., B. C. The
plant is a serious weed in some areas, forming infes-
tations.
This taxon is treated in many different ways by
European authors. Some take up the prior name C.
stoebe L., which may or may not apply. Others treat
maculosa as a variety of C. paniculata or divide macu-
losa into three subspecies, which by other authors, are
raised to species.
Among North American collections the ssp. micran-
thos (Gmel.) Gugler is distinguished by smaller heads
(involucre 10-11 mm high) and fewer (4-6 pairs) and
shorter phyllary processes and by the black or brown
pigmentation of the phyllary margin and processes,
contrasting with the green blade. The ssp. rhenana
(Bor. Gugler and ssp. maculosa have larger heads
(involucre 11-14 mm) and 5-10 pairs of longer mar-
ginal processes which are black — dark brown in
rhenana but brown and partially white in the typical
subspecies.
340
11.
12.
13.
14.
15.
Rhodora [Vol. 74
C. paniculata L. Rare: Canada Vancouver Is., B. C.
This species is similar to C. maculosa and there has
been dispute whether these B. C. plants are paniculata
or maculosa. Howell (1959) has restored C. paniculata
to the B. C. Flora and his decision seems to be justi-
fied.
C. diffusa Lam. Tumble Knapweed; common: U. S.
— apparently widespread in the north, above 40°,
probably common in western states and occasional
elsewhere; Canada — Alta., B. C.
C. virgata Lam. var. squarrosa (Willd.) Boiss. (C.
squarrosa Willd.) ; rare: U. S. — Cal, Utah (Howell,
1959).
C. jacea L. Brown Knapweed; common: U. S. —
widely reported for northeastern and west coast states,
probably midwest also; Canada — Que., Ont., B. C.
Specimens of C. pratensis (nigra X jacea) are fre-
quently identified as C. jacea. Probably both of these
occur within the range commonly stated for C. jacea.
Phyllary appendages are brown, chartaceous and en-
tire or subentire but not deeply lacerate as in the hy-
brids. Heads are radiate.
C. nervosa Willd.
Not known to occur in Canada or U. S. This species
has been reported only by Fernald (1950) from one
collection (Burnham, July 30, 1916, margin of grain-
field, east of lower falls, N. Beaver Creek, Vaughns,
north of Hudson Falls, Washington Co., N. Y.—
GH!). We have revised this specimen to C. austriaca.
Plants of C. nervosa are 10-40 cm tall, usually un-
branched, with a single head, the stem leafy to the
top; lower leaves oblanceolate, base tapering, middle
and upper leaves oblong, base truncate or shallowly
clasping; involucre 20-24 mm high; phyllaries lanceo-
late to 2 cm long, recurved, plumose, black or dark
brown.
1972] Centaurea — R. J. Moore 341
16. C. austriaca Willd. (C. phiygia ssp. austriaca (Willd.)
Gugler); rare: U. S. — N. Y., Vt., O.
This species has been reported only by Seymour
(1969) on the basis of a collection from Dorset, Ben-
nington Co., Vt. (Gilbert s.n., Sept. 22, 1966 — VT!).
We agree with this identification, This collection and
that of Burnham (see C. nervosa) are from adjacent
counties of New York and Vermont, from locations ap-
proximately 25 miles apart. Gilbert notes that the
plants had persisted near farm buildings for several
years but did not seem to mature seed. However, it
seems that the introduction has survived for half a
century in this area. An additional collection, probably
this species, from Ohio (Webb 1525, Sept. 18, 1921,
Portage Co., O. — GH!) has been seen.
These plants are 15-80 cm tall, branched and bear-
ing many heads per plant; cauline leaves narrow, up-
per leaves not clasping the stem; heads smaller than
in C. nervosa, ovate, approximately 14 X 16 mm; in-
ner phyllaries rounded at tip, middle and outer phyl-
laries long and plumose, black or dark brown, recurved.
17. C. nigra L. Black Knapweed, Centaurée noire; (C.
jacea var. nigra Briq. C. consimilis sensu Piper &
Beattie, Flora Southeastern Washington, 1914) ; com-
mon: U. S. — widespread in the northern states, prob-
ably south to about 37° N; Canada — Nfld., P.E.I.,
N: SS N. Ba One. Ont, B. C.
The type specimen of C. nigra has eradiate heads
(Marsden-Jones & Turrill, 1954) and it seems that
plants that are typical in other characters as well are
always eradiate. Descriptions and illustrations in some
floras indicate that C. nigra may have radiate or eradi-
ate heads. The radiate plants are, in our opinion, C. X
pratensis, a hybrid between jacea and nigra which is
highly variable and sometimes closely approaches the
parental species.
In typical C. nigra the phyllary appendages are
342
18.
19.
Rhodora [Vol. 74
black (not dark brown), the wiry cilia very long, al-
most matted, covering adjacent phyllaries. A white-
flowered form (C. nigra f. pallens Spenn) has been
collected in N. S. and Que., Canada.
C. X pratensis Thuill. Protean Knapweed; common:
U. S. — probably throughout the ranges reported for
C. nigra and C. jacea (northern states); Canada —
N. S., Que., Ont., B. C.
This is a hybrid between C. nigra and C. jacea and
is apparently included in the description of C. nigra
in some floras. Abrams & Ferris (1960) and Howell
(1959) report the occurrence of the species on the west
coast of U.S.A. Specimens from Mass., Me., N. Y.,
Pa., R. I., Vt., (GH, VT) have been examined in this
study.
This hybrid shows great variation in the form of
the phyllary appendage: wiry and pectinate, dark
brown but not black, cilia long and covering adjacent
phyllaries; some appendages pectinate and others part-
ly pectinate and partly chartaceous-lacerate and show-
ing many intermediate conditions between pectinate
(nigra character) and chartaceous (jacea character) ;
sometimes all chartaceous and subentire. Thus the ap-
pendage form may closely approach those of the typ-
ical parent species but usually shows some evidence of
an intermediate condition. Heads are usually radiate
but may be eradiate in specimens which closely ap-
proach C. nigra.
C. nigrescens Willd. Short-fringed Knapweed; (C.
vochinensis Bernh., C. transalpina Schleicher p.p.) ;
common: U. S. — probably widespread in the north-
ern states; Canada — N. S., Que., Ont., B. C.
Kerner (1881, pp. 81-85) selected the type of C.
nigrescens in Willdenow’s herbarium and found it to
be the same as the later-described C. vochinensis.
Many U. S. floras recognize C. vochinensis, some take
up C. nigrescens and a few list both species. Appar-
1972] Centaurea — R. J. Moore 343
ently C. dubia is included. There is confusion in the
application of these names. Collections of C. nigres-
cens from Conn., D.C., Ind., Mo., N. J., N. Y., Neb.,
Ore., Pa., Va., Wisc., (DAO, GH, NY) and from the
above-listed provinces of Canada have been seen.
The heads separate C. nigrescens from the general-
ly similar C. nigra. The involucre of C. nigrescens is
longer than broad and the small, triangular, black,
short-fringed appendages contrast strongly with the
green phyllary blades which are not covered by the
appendages. The upper (branch) leaves tend to be
noticeably smaller than the lower leaves. Heads of
this species and of C. dubia are usually eradiate.
20. C. dubia Suter (C. nigrescens Willd. p.p., C. transal-
pina Schleicher s. str., C. jacea var. transalpina Briq.,
C. dubia ssp. eu-dubia (Suter) Gugler & Thellung, C.
nigrescens var. dubia Hermann); common: U. S.—
Mass., N. J., N. Y., O., Ore., Va., Vt.
According to Cronquist (1955; Gleason & Cronquist,
1963) who is the only author to recognize C. dubia in
North America, the species is widely established in
northeastern United States and southeastern Canada
and occasional in northwestern U.S.A. We have seen
specimens (NY, GH) from the above states ; none
from Canada are known. The Species is closely re-
lated to C. nigrescens and it is difficult to separate
some specimens.
These plants are similar to C. nigrescens but the
phyllary appendages have longer cilia which almost,
and sometimes completely, cover the adjacent phyllary
bases. There is no distinct difference in size between
upper and lower leaves and the latter may tend to be
broader.
21. C. macrocephala Puschk. in Willd. Rare: Canada —
Que. a casual adventive from cultivation, probably
not persisting (Moore & Frankton, 1972).
344
22.
23.
24.
26.
27.
28.
29.
Rhodora [Vol. 74
C. rothrockii Greenm. Native species: U. S.:— sw
N. M., se Ariz.
C. americana Nutt. Native species: U. S. — Ark. &
Ia. to Ariz., occasionally an escape from cultivation in
the northeast.
C. repens L. Russian Knapweed; (C. picris Pall.,
Acroptilon repens (L.) DC., A. picris (Pall.) Boiss.) ;
common: U. S.— reported in all states except the
southeastern; Canada — Ont., Man., Sask., Alta., B. C.
This species is frequently a serious weed in our
area. There is cytological and morphological justifica-
tion for segregating this species as Acroptilon repens.
C. moschata L. Sweet Sultan; (Amberboa moschata
(L.) DC.) ; rare: U. S. — northeast, Ill., Cal.; Canada
— Vancouver Is., B. C.
This garden escape seems to persist in the north-
ern states but is never common.
C. montana L. Mountain Bluet; rare: Canada —
Nfld., N. B., Que., Ont., B. C.
Gleason & Cronquist (1963) state that this species
is a rare escape in the northern part of the range of
their flora (northeastern states and adjacent Canada).
C. cyanus L. Bluebottle, Cornflower, Barbeau, Casse
lunette; (Leucacantha cyanus Nwd. & Lunnel, Cyanus
segetum Bauh.) ; common: U. S. — probably all states :
Canada — all provinces except Sask.
This garden escape is reported in all regions. The
variety denudata Suksdorf, reported for Wash., is a
less pubescent variant.
C. cineraria L. Dusty Miller; rare: U. S. — Cal.
C. scabiosa L. Greater Centaurea, Hardheads, Cen-
taurée scabieuse; rare: U. S.— O., Ia., Mont., Can-
ada — Que., s. Ont.
This escape is apparently not widely established; it
1972] Centaurea — R. J. Moore 345
is mentioned for only three states by several floras.
The reported occurrence in N. B., Canada (Boivin,
1966-67) is an error due to a mislabelled specimen
(Boivin, personal comm.).
SUMMARY
Twenty-six introduced and two native species of Cen-
taurea are known to occur in the area Canada-United
States. Sixteen of the adventive species are found in Can-
ada and three of these in Canada alone. An additional ten
introduced and two native species occur in the United
States, making a total of 23 introduced and two native
species in that country.
Probably eleven of the introduced species can be classed
as widespread: C. solstitialis, C. calcitrapa, C. maculosa,
C. diffusa, C. jacea, C. nigra, C. X pratensis, C. repens,
C. cyanus and perhaps C. nigrescens and C. dubia. With
the exception of the last, these species are known from
both countries and generally the ranges are continuous,
although more extensive in the United States. A number
of the rarer species are known only in the southwestern
states, notably California.
Five species (C. diffusa, C. jacea, C. maculosa, C. nigra,
C. repens) are regarded as weeds in Canada (Frankton &
Mulligan, 1970), C. repens being the most serious. Rus-
sian Knapweed has been reported to cause sheep and horse
poisoning in the Old World and C. repens and C. solstitialis
have recently been shown (Young, Brown & Klinger, 1970)
to cause Chewing Disease in horses in the western United
States.
ACKNOWLEGEMENTS
Appreciation is expressed to the curators of herbaria
who have made their collections available for study.
LITERATURE CITED
ABRAMS, L., & R. S. Ferris. 1960. Illustrated flora of the Pacific
states. Vol. IV. Stanford University Press.
346 Rhodora [Vol. 74
Boivin, B. 1966-67. Enumeration des plantes du Canada. Natu-
raliste canad. 93: 253-274; 371-437; 583-646; 989-1063. 94: 131-
157; 471-528; 625-655.
CRONQUIST, A. 1955. Vascular plants of the Pacific Northwest.
Part 5. University of Washington Press.
FERNALD, M. L. 1950. Gray's Manual of Botany. Ed. 8. American
Book Co.
FRANKTON, C., & G. A. MULLIGAN. 1970. Weeds of Canada. Can-
ada Dept. Agriculture Publ. 948. Queen's Printer, Ottawa.
GLEASON, H. A., & A. CRONQUIST. 1963. Manual of vascular plants
of northeastern United States and adjacent Canada. Van Nos-
trand.
HowELL, J. T. 1959. Distributional data on weedy thistles in west-
ern North America. Leaflets Western Botany 9: 17-29.
Jones, G. N., & G. D. FULLER. 1955. Vascular plants of Illinois,
University of Illinois Press.
KEARNEY, K. H., & R. H. PEEBLES. 1960. Arizona Flora. Univer-
sity California Press.
KERNER, A. 1881. Schedae ad floram exsiccatam Austro-Hungari-
cam. I. Vienna.
Moore, R. J., & C. FRANKTON. 1972. Tribe Cardueae (Thistles) in
Canada. Canada Dept. Agriculture (in press).
MARSDEN-JONES, E. M., & W. B. TunmRILL. 1954. British Knap-
weeds. Ray Society. London.
SEYMOUR, F. G. 1969. The flora of New England. C. E. Tuttle
Co., Rutland, Vt.
STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univer-
sity Press.
YouwxG, S., W. W. Brown, & B. KLINGER. 1970. Nigropallidal en-
cephalomacia in horses caused by ingestion of weeds of the
genus Centaurea. J. Amer. Vet. Med. Assoc, 157: 1602-1605.
PLANT RESEARCH INSTITUTE
CANADA DEPARTMENT OF AGRICULTURE
OTTAWA, ONTARIO K1A0C6
CANADA
A NEW VARIETY OF
LEUCOPHYLLUM LAEVIGATUM
(SCROPHULARIACEAE) FROM MEXICO
ROBERT W. KIGER
In the course of field work in northern Mexico during
the summer of 1971, we (J. L. Reveal, W. J. Hess and the
author) collected Leucophyllum laevigatum Standl. from
the Carneros Pass area of Coahuila. Our collection differs
in several respects from the typical phase of the species as
it is found in San Luis Potosi, Durango and Chihuahua.
Aside from ours, the only other collection from Coahuila,
based on a study of specimens at the National Museum of
Natural History at the Smithsonian Institution and the
herbarium of the Field Museum, is one made by Pringle in
the same area in 1890. It also differs from the typical
phase and resembles our collection closely. In view of the
morphological differences found in the Coahuila specimens,
their geographical separation from the rest of the known
range of the species, and the evidence that the same geo-
graphical and morphological correlation has existed over
at least the past 80 years, a new variety of L. laevigatum
is here proposed. This is the first intraspecific taxon to be
described for the species, itself first described in 1924.
Leucophyllum laevigatum Standl.
var. coahuilense Kiger, var. nov.
A var. laevigato foliis plerumque late oblanceolato-
spathulatis, ad 16 mm longis ac 8 mm latis, frequente ac
profunde emarginatis, nunquam acutis, saltem subtus per-
sistenter stellato-tomentulosis, sepalis lineari-lanceolatis,
corolla caerulea, 2-2.5 cm longa, tubo latius campanulato
diagnoscenda.
Low spreading shrub, the young branches densely white
stellate-tomentulose with the pubescence thinning with age;
leaves mostly broadly spatulate-oblanceolate, to 16 mm
long and 8 mm wide, the apex rounded or more frequently
markedly emarginate, never acute, often mucronulate, the
347
348 Rhodora [Vol. 74
young petioles densely stellate-tomentulose, the leaf-blades
sparsely stellate-puberulent above and moderately so be-
low when young, the pubescence persistent at least below ;
pedicels 3-5 mm long, stellate-tomentulose ; calyx cleft near-
ly to base, the sepals linear-lanceolate, to 4 mm long, acute,
stellate-tomentulose outside, often glandular within and
without; corolla blue, yellowish mottled low in tube, 2-2.5
cm long, campanulate, glabrous and semi-lustrous outside,
the tube to 13 mm long, moderately to sparsely arachnose-
pubescent inside, the limb unequally 5-lobed, the lobes obo-
vate, the 3 larger broadly so, rounded at the apex, mod-
erately villous-sericeous within, the margin cilate, the
stamens didynamous, the filaments adnate to the corolla
tube for 1/2 their length, to 7 mm and 8 mm long respec-
tively, the stigma included in the corolla throat at anthesis
about 1 mm below the mouth; mature capsules to 5 mm
long.
TYPE. MEXICO: Coahuila: Carneros Pass area, along Mex-
ico Highway 54, 0.7 mile west of highway, about 0.5 mile
south of Estación Carneros, 23 miles south of Saltillo, in
low canyon associated with Yucca, Agave, Opuntia and
Juniperus, 5 August 1971, Reveal, Hess & Kiger 2617. Holo-
type US! Isotypes to be distributed from US.
ADDITIONAL SPECIMENS EXAMINED. MEXICO: Coahuila:
Carneros Pass area, Pringle 3460 (F, US).
The typical variety of Leucophyllum laevigatum has
leaves proportionately more narrowly spatulate-oblance-
olate, to 20 mm long and 6 mm wide, more often rounded
and less frequently and deeply emarginate, sometimes
acute, and becoming glabrous or at least glabrate with age.
The sepals are linear and usually 2-3 mm long, rarely
reaching 4 mm in length. The most striking differences
between the two varieties are associated with the corolla.
In the typical variety, as in other species of Leucophyllum
in Mexico, the corolla is purple to violet, whereas in var.
coahuilense it is a definite blue. In the latter the corolla
is consistently from 2 to 2.5 cm long at anthesis, most often
1972] Leucophyllum — Kiger 349
closer to 2.5 em, while in var. laevigatum it is from 1.3 to
2 cm long, usually about 1.5 cm. The tube is proportionate-
ly longer in var. coahwilense and is more broadly cam-
panulate than in the typical variety where it is narrowly
so. Though capsules as long as those usually found in var.
laevigatum (to 6 mm) have not been observed in the new
variety, the difference is probably not significant since the
specimens of the latter have only a few mature capsules.
Too, while the capsules of the typical] variety reach 6 mm
in length, they are often smaller and thus overlap the range
observed for var. coahuilense.
The type of Leucophyllum laevigatum var. laevigatum
was collected by Nelson (4689) in Durango, Mexico, be-
tween Ramos and Indé (holotype, Us!). Other specimens
of this variety examined include — MEXICO: San Luís
Potosí: Charcas, Lundell 5512. Durango: Huarichic, north-
east of Pedricena, Pennell 18604; Indé, Reko 5207; 5 miles
northeast of Yerbanis, Moran 6248; 10 miles north-north-
west of Rodeo, Morley 630; 20 miles south of Zarca, Gen-
try 8614. Chihuahua: near La Boquilla, Shreve 8072. All
specimens examined are at US.
Field studies in Mexico were supported by an NSF grant,
GB-22645, to James L. Reveal for studies on the genus
Eriogonum and the Intermountain Flora.
DEPARTMENT OF BOTANY
UNIVERSITY OF MARYLAND
COLLEGE PARK, MARYLAND 20742
VARIATION WITHIN CALOCHORTUS VENUSTULUS
(LILIACEAE)
JAMES L. REVEAL AND WILLIAM J. HESS
While collecting specimens of a new Hriogonum in the
high mountains west of Hidalgo del Parral of northern
Mexico, we made small collections of routine nature in
order to better understand the local vegetation. In addi-
tion to the new Eriogonum, we found an unknown Castil-
leja which Dr. Noel Holmgren of the New York Botanical
Garden will describe and a variant of Calochortus venus-
tulus Greene which likewise is undescribed.
Calochortus venustulus Greene var. imbricus Reveal &
Hess, var. nov. A var. venustulo petalis 20-28 mm longis,
pedunculis 10-15 cm longis, plantis 3-4.5(6) dm altis dif-
fert. Stems 3-4.5(6) dm high; peduncles erect, slender,
10-15 em long; petals obovate, 2-2.8 cm long.
TYPE. MEXICO: Durango: Sierra Madre Occidental, along
the dirt road from Hidalgo del Parral toward El Vergel
out of San Francisco del Oro, about 60 road miles west of
Parral and 18.5 road miles west of Ojito, on forest floor
and slopes near limestone outcrops, associated with Quercus
reticulata, Pinus ponderosa and P. leiophylla, Arctostaphy-
los, Arbutus and numerous other low shrubs and herbs, at
about 8000 feet elevation, 11 August 1971, Reveal, Hess &
Kiger 2741. Holotype, us! Isotypes to be distributed from
US.
The var. imbricus (from the Latin imbricus, rainy, al-
luding to the rainy day that we discovered this plant) is
a lovely addition to a genus already well noted for its
beautiful members. Particularly, in the darkness of the
rain and heavy clouds on that day, its large, golden flowers
stood out in marked contrast from the dark greens and
browns of the pine-oak forest floor.
During our investigations of this taxon, we found that
Calochortus venustulus, as defined by Ownbey (1940),
should be further subdivided.
The typical variety, var. venustulus, is common in the
Sierra Madre Occidental west of Durango where it occurs
350
1972] Calochortus — Reveal and Hess 351
in the forests and dry meadows. However, to the north,
west of Chihuahua and hence northward nearly to the
United States, a second element is common in open plains
and scattered oak forests. This phase was named C. mad-
rensis by Watson just a few months after Greene proposed
C. venustulus and has been placed in synonymy under the
latter name. However, the northern population is consis-
tently taller, with longer peduncles and shorter leaves in
relationship to the inflorescence. This pattern of distribu-
tion is somewhat similar to that found in Eriogonum atro-
rubens Engelm. in Wisliz. (Reveal, 1967), and likely is
expressed by other species as well. Thus, we feel the north-
ern element should be recognized as C. venustulus var.
madrensis (S. Wats.) Reveal & Hess, based on C. madren-
sis S. Wats., Proc. Amer. Acad. Arts 23: 283. 1888.
These variants may be distinguished as follows:
A. Petals (10) 12-16 (18) mm long. (B)
B. Plants 1-2 dm high; leaves mostly longer than the
inflorescence; peduncles 3-6 cm long; Durango
var. venustulus
B. Plants (1.5) 2.5-4.5 dm high; leaves mostly as long
to slightly shorter than the inflorescence; peduncles
5-8 cm long; Chihuahua var. madrensis
A. Petals 20-28 mm long; plants 3-4.5 (6) dm high; leaves
mostly shorter than the inflorescence; peduncles 10-15
em long; extreme northern Durango .. var. imbricus
Field studies in Mexico were supported by an NSF
grant, GB-22645 to Reveal for studies on the genus Eriogo-
num and the Intermountain Flora.
LITERATURE CITED
OwNBEY, M. A. 1940. A monograph of the genus Calochortus.
Ann. Missouri Bot. Gard, 27: 371-560.
REVEAL, J. L. 1967. Notes on Eriogonum — II. Variations in Erio-
gonum atrorubens. Sida 3: 82-86.
UNIVERSITY OF MARYLAND
COLLEGE PARK, MARYLAND 20742
and
NEWARK STATE COLLEGE
UNION, NEW JERSEY 07083
NEW FORM OF THUJA OCCIDENTALIS
RESEMBLING KNOWN CULTIVARS
Mary I. MOORE
In 1963 the author visited the canyon of the Barron
River in Algonquin Provincial Park, Ontario at latitude
45954' N and longitude 77°38’-39’ W. Normal Thuja occi-
dentalis L. grows on slopes at the foot of cliffs in this mile
long fault in granitie gneiss. On several talus formations
there are dwarf forms, barely a meter high, with dead
tops, compact, prostrate branching and normal leaf devel-
opment. One dwarf shrub of very unusual form was found
(Moore, 1964) on a north facing rock slide covered with a
carpet of chasmophytic ferns, Cladonia and bryophytes
such as Hedwigia and Polytrichum. A specimen was sent
to the Plant Research Institute, Dept. of Agriculture, Ot-
tawa. Drs. W. G. Dore and C. Frankton were unfamiliar
with the shrub but established the anatomy of the stem as
that of a conifer and the odor as that of Eastern white
cedar (Frankton in litt., 1963). In 1968 Dr. R. E. Beschel,
Queen's University, Kingston, Ont. visited the shrub and
noted its resemblance to certain bizarre cultivars. Both he
and Mr. E. Perem of the Wood Anatomy Group, Eastern
Forest Products Laboratory, Ottawa considered that the
wood anatomy corresponded with the description given for
T. occidentalis (Arbor vitae).
DESCRIPTION
The shrub has a central stem 0.8 m high, with eight
branches up to 2 m in length on the lower half. Of these,
two come from beneath the rock and are in a prostrate
position, The branches may have three orders of branch-
ing and bear live or dead leaves. They are basically of
three types:
(1) About 75% are whip-like, sparingly ramified, with
four rows of equal, appressed-decussate, ovate, keeled and
pointed leaves 3-5 mm long (Fig. 1; 2A, 2B. Fig. 2A).
The overall tetragonal habit of the branches resembles
strongly some appressed species of Cassiope D. Don.
352
1972] Thuja occidentalis — M. Moore 353
Figure 1. Branch and leaf form of typical Thuja occidentalis and
the dwarf conifer in the Barron canyon. A — side view of branchlet,
B — end view of branchlet: 1 — typical T. occidentalis, 2 — tetrag-
onal, decussate form on dwarf conifer, 3 — lycopodiaceous form on
dwarf conifer.
(2) Several branches bear fastigiate clusters of twigs
with rows of divaricate, needle-shaped leaves 6-8 mm long.
These appear juvenile but some are stiffer and more sharp-
ly pointed. They may develop in a gradual sequence out
of the first type and revert again to it (Fig. 2B).
(3) A few small sprays bear leaves similar to but usually
smaller than (1). These branchlets are flattened, without
the constricted segments of normal T. occidentalis, and are
intensely branched in the manner of Lycopodium trista-
chyum Pursh (Fig. 1; 3A, 3B. Fig. 2C).
DISCUSSION
The shrub bears no external resemblance to a normal
Thuja occidentalis. A search of botanical and horticul-
Q9
or
nS
Rhodora [Vol. 74
Figure 2. Branches from dwarf conifer in Barron canyon. A —
tetragonal, whip-like branch, sparingly ramified, B — fastigiate clus-
ter with juvenile-type needles, C — lycopodiaceous form of branch-
ing.
tural literature reveals no such specific entity. The forma
prostrata of Vict. & Rousseau has normal adult leaf struc-
ture and differs mainly in habit. Var. ericoides Beiss. &
Hochst., which is also a cultivar, bears only small resem-
blance to parts of the shrub with juvenile-type needles. The
cultivar, 'Douglasi? Rehder, given as a synonym for ‘Fili-
formis’ Beiss., has some similarity with it and is described
as a compact shrub with slender, pendulous branches often
flattened at the tip. Bailey (1923) says these branches are
partly four-angled with sharply pointed leaves, but his il-
lustration shows very typical T. occidentalis foliage. Reh-
der (1901), the authority for the variety, states that the
stronger shoots are round, covered with widely-spaced, long,
pointed and somewhat projecting leaves, but that there is
1972] Thuja occidentalis — M. Moore 355
no juvenile foliage. The illustration of Welch (1966) con-
firms this. 'Ohlendorfi? has tetragonal branches with de-
cussate leaves but the illustration in Welch shows a very
compact shrub with upright branches.
The leaves of the tetragonal and lycopodiaceous branches
do show an interesting resemblance to those of T. orien-
talis L., species of Cupressus and Chamaecyparis (Dalli-
more & Jackson, 1948) as well as to forms of Juniperus.
Horticultural forms of dwarf and other abnormal coni-
fers are developed by mutation, self-pollination and hy-
bridization. According to Welch most, if not all, cultivars
of Arbor-vitae were originally the result of seed mutation.
Such seedlings can be cultivated in a nursery, but in nature
their chances of survival are not great. ‘Spaeth? (Anony-
mous, 1893), a synonym of ‘Ohlendorfi’, was developed in
this way from a seedling of T. occidentalis.
CONCLUSION
The question has been asked, is the shrub a monstrosity
induced by the particular habitat in which it occurs, or is
it a genetic form that arose as a result of self-fertilization,
hybridization or mutation.
The shrub is close to the river where in some years it
could be subject to damage from ice and high water; a visit
in 1970 showed that this had recently occurred with dam-
age to many of the long, tetragonal branches. The climate
of the canyon is, in part, abnormally cool; several rare
arctic-alpine species occur there (Moore, 1964) and on the
first visit in late May two large ice falls were still pres-
ent. The lower branches of cedars along the river are
browsed. However, no other known cedars have abnor-
ma] leaves.
Abnormal leaf forms may appear on young conifers,
usually under thirty years of age, after which they usually
outgrow all or most of the characteristics (A. R. Buckley,
Horticulture Division, Plant Research Institute, pers.
comm., 1969). The shrub in question must be at least fifty
356 Rhodora [Vol. 74
years old since Mr. Perem gave the age of a small branch-
let 2.3 mm in diameter as more than twenty years.
The plant presumably could not have been a hybrid be-
cause no other species of Thuja grows naturally in On-
tario. The canyon could not have been cultivated and what
farms may have been in this area of the precambrian
shield have returned to forest. It could have been due to
self-pollination. However, it seems most probable that the
shrub is the result of a spontaneous mutation. Unfortu-
nately there is no evidence of flowering or cone develop-
ment.
Mr. John Santon, Petawawa Forest Experiment Sta-
tion, has rooted tiny cuttings of both juvenile and tetrag-
onal foliage. Growth the first winter was generally of the
tetragonal, decussate form common to the shrub but pres-
ent summer growth is mostly of the juvenile type. Mr.
Santon considers that this alteration of form may be
seasonal but commented that a return to juvenile growth
is frequent in cuttings.
The dwarf conifer in the Barron canyon appears to be
a unique form. Its wood anatomy defines the shrub as
Thuja occidentalis and the odor is characteristic of that
species. It can be assumed that this Eastern white cedar
was not induced by humans although the shrub does re-
semble in part several bizarre cultivars of the species pre-
viously described.
ACKNOWLEDGMENTS
I wish to thank all those who gave help with the manu-
script and especially those who helped to identify the
shrub.
SELECTED BIBLIOGRAPHY
ANONYMOUS. 1893. Thuya occidentalis Spaethi. Gartenflora, Ber-
lin, 42: 539.
BAILEY, L. H. 1923. The cultivated evergreens. MacMillan Co.,
New York. 224-25,
1944. The standard cyclopedia of horticulture.
MacMillan Co., New York. 3335-36.
1972] Thuja occidentalis — M. Moore 357
CARRIERE, E. A. 1880. Revue du genre Retinospora. Revue Hor-
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DALLIMORE W., and A. B. JACKSON. 1948. A handbook of coniferae.
Edward Arnold Ltd., London. 686 pp.
FERNALD, M.L. 1950. Gray’s manual of botany. American Book
Co., New York. 1632 pp.
HORNIBROOK, Murray. 1923. Dwarf and slow-growing conifers.
Scribner, New York. 165-75.
KRÜSSMANN, G. 1960. Die Nadelgehólze. Paul Parey, Berlin and
Hamburg. 295-303.
Moore, Mary I. 1964. Some interesting plants in the Barron can-
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REHDER, ALFRED. 1901. Thuya occidentalis var. Douglasi. Móeller's
Deutsche Gártner-Zeitung. Erfurt, Germany. Vol. 16: 30.
1940. Manual of cultivated trees and shrubs
hardy in North America. MacMillan Co., New York. 996 pp.
SILVA Tarouca, E. and C. SCHNEIDER. 1923. Unsere Freiland-
Nadelgehólze. Leipzig. 288-90
SYNGE, P. M. (editor). 2nd edition. Dictionary of gardening. Royal
Hort. Soc., Oxford. 4: 2104.
WELCH, H. J. 1966. Dwarf conifers. Faber and Faber, London.
334 pp.
PETAWAWA FOREST EXPERIMENT STATION
CHALK RIVER, ONTARIO
SNOW COVER AND THE DIAPENSIA LAPPONICA
HABITAT IN THE WHITE MOUNTAINS,
NEW HAMPSHIRE!
WESLEY N. TIFFNEY, JR.
The alpine zones of the White Mountains, New Hamp-
shire, provide a convenient study area for investigators
interested in the ecology of alpine plants. The White Moun-
tains are a series of peaks, heavily glaciated during the
Pleistocene Epoch, 46 of which exceed 4,000 ft. The highest
of these is Mt. Washington (6,288 ft.) in the Presidential
Range. During the Wisconsin substage of Pleistocene
glaciation the climate allowed arctic plants to become estab-
lished in the area. The present harsh climate typical of the
alpine zones of the White Mountains has maintained these
plants as an arctic outpost in a temperate area. Treeline,
here defined as the upper limit of continuous tree cover, is
located at about 4,800 ft. on the west side of the Presi-
dential Range and at about 5,200 ft. on the east (Antevs,
1932; Bliss, 1963). Prevailing winds are from the west.
The best developed alpine zone in the White Mountains is
found on the Presidential Range; it extends from Mt.
Madison (5,363 ft.) in the north to Mt. Clinton (4,312 ft.)
in the south. The area is about eight miles long and two
miles wide at its greatest extent and encompasses about
7.5 square miles (Bliss, 1963). A smaller alpine zone exists
in the Franconia Range some 20 miles west of the Presi-
dential Range; it extends from Mt. North Lafayette (5,000
ft.) to Mt. Haystack (4,600 ft.), having as its highest point
Mt. Lafayette (5,249 ft.). This zone encompasses about 1.5
square miles.
The climate of the alpine zone has been summarized by
several investigators (Antevs, 1932; Bliss, 1963; Pease,
1964). Weather information has been gathered by the Mt.
‘This paper constitutes a part of a doctoral dissertation submitted
to the University of New Hampshire.
358
1972] Diapensia — Tiffney 359
Washington Observatory located on the summit of that
peak. In general, the climate is characterized by high
winds, low temperatures, high precipitation with heavy
winter snow and extensive fog and cloud cover.
This paper represents a portion of a larger investigation
involving the distribution of White Mountain plants rela-
tive to winter snow cover. The purpose of the present study
is to define the ecological relationship between snow cover
and a habitat typified by Diapensia lapponica L., an Angio-
sperm cushion-forming plant, used by Bliss (1963) to
characterize the association he termed the “Diapensia
community".
In both the arctic and in the alpine zones of the White
Mountains Diapensia is reported in exposed and wind-
swept areas (Antevs, 1932; Hadley and Bliss, 1964; Lóve
and Lóve, 1966; Courtin, 1968, 1968a). Several investiga-
tors have further suggested that the plant is limited to
areas blown clear of winter snow (Hadley and Bliss, 1964;
Courtin, 1968, 1968a). Bliss (1963) reports that Diapensia
lapponica is a prime colonizer of bare ground and is found
in ". . . areas that are subjected to frost action because
of the lack of winter snow cover". Later, (1966) he reports,
"Diapensia communities are found where winter snow
cover is thin, or non-existent, and melts early in spring".
Bliss (1969) recorded snow depth for two winters in three
locations; it is not stated if any of these locations were in
Diapensia areas. However, he states, “During much of the
winter, large areas of Alpine Garden are blown free of
snow, areas dominated by Diapensia communities".
My objective in this study was to analyze the species
composition of the Diapensia habitat and to determine the
degree of interdependence among its component species.
First, it was necessary to test the snow cover hypothesis
described above by gathering extensive records of winter
snow depths in Diapensia habitats. I observed a number
of habitats over several seasons to find if the plants did
inhabit snow-free areas and if these areas were sub-
stantially free of snow throughout the winter season. A
360 Rhodora [Vol. 74
system of summer sampling was designed to accumulate
data for statistical analysis of interdependence among com-
ponent species of the habitat.
Snow Depth and Winter Dessication
For the winter snow depth study 14 plots were estab-
lished and marked with wire stakes at the margins of the
greatest Diapensia concentrations. Two plots were located
in each of the following areas: near the fourth peak of
Mt. Adams, at the Lake-of-the-Clouds Hut, on the summit
cone of Mt. Washington, on the section of the Chandler
Ridge termed Cape Horn, at the lip of Huntington Ravine
in Alpine Garden, at Lion Head in Alpine Garden and at
Cow Pasture on the Automobile Road. These plots were
visited throughout the winters of 1966-1967, 1967-1968 and
1968-1969 on as nearly a regular basis as the harsh winter
weather of the Presidential Range permitted. Each plot
was visited at least once each winter and some aS Many
as eight times. During the three winters 14 trips were
made to check snow depths in plots.
After a snowfall in the White Mountains the wind fre-
quently rises, usually from the west. Most of the snow
which has accumulated during the storm in the open alpine
zone is blown away and collects below treeline on the east
side of the mountains. This is particularly true on the
Presidential Range and leads to the massive accumula-
tions of snow common in the large glacial valleys on the
east side of the range, such as Tuckerman and Huntington
Ravines. Snow remaining in the alpine zone is packed into
the sparce vegetation or in the lee of large wind obstruc-
tions. In ascending the mountain increasing snow depth
is encountered until treeline is reached; above treeline snow
cover conforms to the microrelief of the alpine zone. In
the Diapensia habitat there are few obstructions and virtu-
ally all the snow is blown clear throughout the winter
season (Fig. 1).
A trip to check snow depth made on February 19-26,
1967, will serve as an example of the snow cover /wind
1972] Diapensia — Tiffney 361
Fig. 1. Snow-free ridge crest, about one-half mile long, near Lion
Head in Alpine Garden, Presidential Range, New Hampshire. The
area is dominated by Diapensia lapponica. March 3, 1969.
relationship on the Presidential Range. On the night of
February 20 a storm deposited a recorded 2.7 inches of
snow at the summit of Mt. Washington. Drifts of two to
three feet accumulated in the vicinity of the Weather
Observatory. On the morning of February 21 the wind
rose from the west and reached a sustained velocity of
60 to 70 MPH with a peak gust of 105 MPH. Virtually all
the snow, including the large drifts, was removed from
the summit area. On February 23-24 another storm
developed and deposited a recorded 19 inches of snow at
the summit. Winds rose during the evening of February 24
and reached a sustained velocity of 70 to 80 MPH. The
wind persisted through February 26. Again, almost all ac-
cumulated snow was removed from the summit area. Visits
to six Diapensia habitats on February 26, varying from
6,000 to 4,000 feet elevation, showed that all were devoid
362 Rhodora [Vol. 74
SR ag Ge 777 0
v de iiim ae, - es Wr
LI
Fig. 2. Winter aspect of a Diapensia habitat at Cape Horn,
Chandler Ridge, Mt. Washington. Surrounding areas are covered
with snow. The ice axe is about 40 inches long. Feb, 26, 1968
of snow after a total fall of 21.7 inches over the previous
six days (Fig. 2).
Late wet snowfalls, although more dense than dry mid-
winter snows, are also blown away by strong winds and
do not accumulate in the Diapensia habitat nor do they
delay flowering. On May 26, 1967, a 12.8 inch snowfall was
followed by winds of 70 MPH. Visits to several habitats
on May 30 and 31 disclosed no snow in the areas. Less
than three weeks later on June 18, 1967, Diapensia plants
were in full bloom throughout the Presidential Range
(Fig. 3).
Winds are not as strong where plants are actually grow-
ing on the range as they are where recorded on a 39 foot
instrument tower on the summit of Mt. Washington. A
wind velocity of 100 MPH on the summit is equivalent, on
the average, to a speed of 30 MPH at plant level (Courtin,
1972] Diapensia — Tiffney 363
Fig. 3. Plants of Diapensia lapponica in bloom, Cape Horn, Mt.
Washington, New Hampshire. June 17, 1968. The lateral area
covered by the photograph is about 18 inches long.
1968). Such wind velocities are stil] sufficient to remove
snow from exposed areas and to promote desiccation in
winter-exposed plants.
In an average winter on Mt. Washington the Weather
Observatory records 221.4 inches of snow. The winter of
1966-1967 totaled 260.6 inches, the winter of 1967-1968,
211.1 inches and the record winter of 1968-1969 yielded
566.4 inches of snow. These variations in annual snowfall
have little effect on the total amount of snow adhering to
the ground in the alpine zone. My observations indicate
that the snow holding capacity of the zone is reached early
in the winter. Additional snow is blown away by strong
winds. This study spanned two relatively normal snowfall
years and one record year. Amounts of snow in Diapensia
habitats and other areas checked remained constant. This
minimal snow cover melts rapidly once warm -weather
arrives and melts at about the same time each year. Even
364 Rhodora [Vol. 74
Fig. 4. Winter condition of Diapensia lapponica plants, Cape
Horn, Chandler Ridge, Mt. Washington, New Hampshire. Meltwater
has formed ice about the bases of the plants but the crowns are
fully exposed. Jan. 26, 1967. The ice axe is about 40 inches long.
late-lasting drifts supporting specialized snowbank com-
munities (Bliss, 1963) disappear at about the same time
each season. The flowering time of Diapensia lapponica
and other spring-blooming plants on the Presidential Range
is not greatly influenced by heavy or light snow years or
by heavy snowfalls late in May. In the three years included
in this study alpine plants were at the peak of their bloom
between June 15 and June 25 (Fig. 3).
The examples above, drawn from the winter of 1966-
1967, are typical of results of other visits in 1967-1968 and
1968-1969. In some cases meltwater had formed ice about
the bases of Diapensia plants but in all cases observed the
major portion of the plant was exposed (Fig. 4). Areas
dominated by Diapensia were those areas blown clear of
snow throughout the winter. Accumulations of as little
as a few inches of snow marked the margin of the Diapensia
1972] Diapensia — Tiffney 365
habitat and the beginning of a transition to another form
of plant cover.
Desiccation is a major factor in the distribution of alpine
plants as Tranquillini (1964) notes when he states, “Ex-
tremely short snow cover can only be tolerated by plants
that are resistant not only to cold but also to long periods
of drought . . .", and Sakai (1970) notes that “. . . desic-
cation damage in winter constitutes the greatest limiting
factor for growing plants in cold climates and in high
mountains . . .". Plants in the Diapensia habitat appear
well suited to resist the effects of desiccation. Several in-
vestigators have noted that the low mat or cushion form
exhibited by plants such as Diapensia is of survival value
in these conditions (Holttum, 1922; Hadley and Bliss,
1964; Courtin, 1968a). The low profile of these plants
offers less resistance to strong winds and their desiccating
effect is minimized.
Information on the freezing resistance of several plants
of the Diapensia habitat has been accumulated by Sakai
and Otsuka (1970). They note that several species acquire
a resistance in the fall at —15? C. that allows them to with-
stand winter temperatures of —70° C. After acquiring such
resistance Vaccinium uliginosum could withstand tempera-
tures of —50° C., V. vitis-idaea and Loiseleuria procumbens
could survive at —70° C. The most cold resistant plant
tested was Diapensia lapponica which, after full harden-
ing, could survive immersion in liquid nitrogen at —-196? C.
(Sakai and Otsuka, 1970). The lowest recorded tempera-
ture for Mt. Washington is —44? C. (ATS EJ.
Species Composition
Twenty-two Diapensia habitats were chozen for sampling
to determine species composition and major sample areas
were located in them. Plots were placed in the center of
the habitat. The sampling procedure used is similar to
that of Bliss (1963). An area 4 by 8 meters (320,000 sq.
cm.) was marked with surveying tape and stakes. The
4-meter axis was located parallel to the slope of the ground.
366 Rhodora [Vol. 74
1 2 3 4 5 6 7 8
TRANSECT LOCI - METERS
Fig. 5. Arrangement of quadrats within a major sample area.
The transects on which the quadrats are located are randomly
chosen. The design is similar to that of Bliss (1963).
Five rectangular quadrats 25 by 40 cm. (1,000 sq. cm.)
were established along a sequence of transects based at
four randomly chosen locations on the 8-meter axis. Alter-
nate quadrat transects were begun on the 8-meter axis,
and others 0.5 meters from it. Twenty rectangular quadrats
were established in each major sample area for a total
sampled area of 20,000 sq. cm. or 6.3% of the total area
sampled (Fig. 5). Quadrat size and number corresponded
with recommendations for plot size and sample design in
alpine vegetation made by Eddleman et al. (1964).
Measurements were taken of the ground area covered by
all vascular plants in each 25 by 40 cm. quadrat. Measure-
ments were taken also for cover of moss and lichen and
for uncolonized ground and rock. These were expressed
as percent cover for each sample area, and grand totals
for all 22 areas sampled were compiled. Areas sampled
for species composition were, in general, the same as those
used for winter observations. Results are given in Table 1.
My results expressed as percent cover are roughly com-
parable to those of Bliss (1963) expressed as importance
1972] Diapensia — Tiffney 367
Table 1
Standard
Type Mean Range Deviation
Rock 31.1 1.4-66.6 19.5
Diapensia lapponica 27.6 1.8-54.0 15.0
Lichen (all types) 13.0 0—46.4 14.1
Vaccinium uliginosum 5.8 0-18.5 5.2
Juncus trifidus 5.3 0-23.0 6.4
Uncolonized Ground 4.3 0-18.2 4.6
Moss (all types) 3.2 0—31.1 6.7
Carex bigelowii EM 0—11.8 3.6
Salix uva-ursi 1.5 0-7.9 2.5
Vaccinium vitis-idaea 1.1 0-7.2 2.0
Potentilla tridentata 1:0 0—4.8 1.3
Loiseleuria procumbens 0.8 0-7.8 1.9
Rhododendron lapponicum 0.6 0—4.3 1d
Arenaria groenlandica 0.5 0-2.5 0.7
Results of sampling major plant components of the
Diapensia habitat, White Mountains, New Hampshire.
Means, ranges and standard deviations for all types
sampled are expressed as percent ground cover of the total
area sampled.
Other plants were present in quantities of less than
0.5% ; these minor components were not subjected to sta-
tistical analysis. These were, together with their means:
Solidago cutleri 0.4% ; Poa alpigena 0.3%; Scirpus cespi-
tosus var. callosus 0.2%; Betula ssp. 0.1%; Vaccinium
angustifolium 0.05% ; Abies balsamea 0.05% ; Empetrum
nigrum 0.03% ; Prenanthes boottii 0.01% ; and Lycopodium
selago 0.002%. These minor forms totaled 1.1% of the
total area sampled. The nomenclature used is that of
Fernald (1950).
368 Rhodora [Vol. 74
values. He recorded higher values for rock and uncolonized
ground as well as for Juncus trifidus; slightly higher values
for Potentilla tridentata, Loiseleuria procumbens, Rhodo-
dendron lapponicum, and Arenaria groenlandica; similar
values for Diapensia lapponica and Vaccinium uliginosum,
and a lower value for lichen. Results of Bliss’s analysis
are based on 2 to 9 sample areas.
Results of my sampling indicate that 47.3% of the area
in the Diapensia habitats was occupied by vascular plants,
35.4% was occupied by rock and uncolonized ground and
16.2% by moss and lichen. The typical plant of these
habitats was Diapensia lapponica which was about five
times as concentrated as the next ranking vascular plant.
Wide ranges and large standard deviations were found
for all component species tested (Table 1). This suggests
that it is not possible to predict with accuracy the concen-
trations of the components to be found in a given Diapensia
habitat. The minimum concentration of Diapensia lap-
ponica, 1.8% of the area sampled, was encountered in a
plot located on the west flank of Mt. Madison. Other sample
plots placed in nearby Diapensia habitats yielded results
of 30.1%, 35.0% and 47.2% cover of D. lapponica. The
maximum concentration encountered, 54.0% was in a plot
located on the west side of Mt. Lafayette in the Franconia
Range. Nearby sample plots in this area showed concentra-
tions for D. lapponica of 15.6%, 22.0% and 33.4%. Values
for other component species of the habitat were equally
variable.
Analysis of Component Species
To assess the degree of interdependence existing among
the component species of the Diapensia habitat statistical
testing was necessary. Tests based on frequency data such
as the “C.” correlation of Hurlbert (1969) were rejected
due to the difficulty of collecting accurate frequency data
in alpine vegetation. As Woodin (1959) has indicated,
frequency information is difficult to obtain in alpine areas
of mat-form vegetation as it is hard to tell where one plant
1972] Diapensia — Tiffney 369
ends and another begins. Goodall (1970) notes that, “Many
plants, however, have methods of vegetative reproduction
which make distinctions between individuals highly arbi-
trary. Biomass per unit area is a measure which is more
consistently objective’. For these reasons estimates of
biomass in terms of percent ground cover were used here.
The purpose of this part of the study is to assess the degree
of interdependence present among plants of the Diapensia
habitat. Sokal and Rohlf (1969) have said, “. .. when we
wish to establish the degree of association between pairs
of variables in a population sample, correlation analysis
is the proper approach.” Hence, the product-moment cor-
relation coefficient has been used here.
In computing the correlation coefficient, data are ranked
in two sets. The first set (Y,) is arranged in increasing
rank order, the second set (Y.) is paired with the first. The
correlation coefficient (7) indicates the degree of associa-
tion between the two sets (Y, and Y.) and may be expressed
in percent. The value of r may vary from -1 (minus 100%)
to +1 (plus 100%). A value approaching -1 indicates
negative association between Y, and Y,, a value approach-
ing +1 indicates positive association between the two sets
of variables. A value of r approaching 0 indicates little
interdependence between Y, and Y.. Reciprocal tests yield
the same results. For each correlation coefficient (value
of r) significance tests are performed against the hypoth-
esis that r — 0.
Each of the 14 major species components of the Diapen-
sia habitat (those having means greater than 0.5% of the
total area sampled) was tested against the other 13 major
components. This required a total of 91 correlation co-
efficient tests, which were performed with the aid of a
computer and using Leasco Systems and Research program
no. CL-00001.023-00. Results are given in Table 2.
Ninety-one tests were performed; only eight yielded
positive or negative correlation coefficients of 50% or
greater. The strongest relationship observed was between
moss and Arenaria groenlandica with a value of 68.0%.
[Vol. 74
Rhodora
370
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1972] Diapensia — Tiffney 371
None of the relationships tested could be considered strong
(in excess of 90%).
A negative correlation coefficient resulted from the test
between rock and lichen (-66% ). Hence, rock and lichen
covary negatively 66% of the time. Lichens sampled in
this study were of the foliose and fruticose types. Crustose
or rock-inhabiting forms were rare and were not included.
A rock habitat is not suitable for the growth of lichens in
the areas tested. With the exception of the negative rela-
tionship with lichens mentioned here, the large quantities
of rock present in the Diapensia habitat (31.1% of the
total area sampled) do not greatly influence the plants
growing in the habitat.
Negative correlations resulted from tests of Diapensia
lapponica with Vaccinium uliginosum (56%) and with
Juncas trifidus (-54%). This indicates that D. lapponica
is negatively associated with V. uliginosum and J. trifidus
about 55% of the time. The latter two plants are com-
ponent species of an aggregation termed the “dwarf-shrub-
rush-heath community” by Bliss (1963). This aggrega-
tion is commonly found at the margins of the Diapensia
habitat. As these margins are reached snow cover increases
slightly and V. uliginosum and J. trifidus begin to dominate.
These two species also characterize areas of slightly greater
snow cover, as in the lee of small obstructions dispersed
throughout the Diapensia habitat itself. A gradient was
established by Bliss (1963) with Diapensia at the snow-
free extreme and the snowbank community at the other
extreme. “Dwarf-shrub-rush-heath” is placed next in order
with increasing snow cover to the Diapensia aggregation
on this model. Rather than indicating that competitive
exclusion is operating among D. lapponica and V. uligino-
sum and J. trifidus, I conclude that the negative correla-
tions indicate that the margin of the Diapensia habitat
has been reached in these sample situations and that a
change to dominance by other forms is occurring. In this
case the optimum snow cover situation for Diapensia has
372 Rhodora [Vol. 74
been exceeded, and the optimum for the other two species
is being approached.
This situation poses some possibilities for future experi-
mental work. Snow depth in selected Diapensia habitats
could be increased artificially by erecting several wind-
breaks in the habitat. Vegetation could be mapped prior
to establishing windbreaks, then re-mapped in successive
growing seasons. At question would be changes in the
composition of plant aggregates growing in the altered
habitat. A project of this nature is planned.
The test between Vaccinium uliginosum and lichen yielded
a positive correlation of 61%. This indicates positive inter-
dependence between V. uliginosum and lichen 61% of the
time. Lichens, lacking roots, are not as firmly anchored
as other plants and are frequently blown away by strong
winds, an occurrence noted also by Antevs (1932). Most
plants in the Diapensia habitat are mat-forming or cushion
plants. An exception is V. wliginosum which, particularly
in areas of slightly greater snow cover, adopts a more up-
right stature. Under these conditions V. uliginosum consists
of a number of interwoven branches which provide anchor-
age for lichens. Possibly, the vascular plant provides sup-
port for lichens of several types and prevents them from
being blown away by strong winds.
Positive results were obtained from tests between Are-
naria groenlandica and uncolonized ground (54%), and
moss (68%) and Carex bigelowti (61%).
The *sedge-meadow community” of Bliss (1963) was
composed of two vascular plants, Carex bigelowii and Are-
naria groenlandica. In addition to keeping snow cover
records for the Diapensia habitat I accumulated informa-
tion on winter snow depth in areas of sedge meadow on
the summit cone of Mt. Washington. About one to two
inches of snow accumulate in these areas, the rest being
blown away. This amount is sufficient to cover the ground
level meristems of C. bigelowii, although dead culms from
the previous growing season frequently protrude from the
thin snow cover. Sedge meadow occupies a similar posi-
9
1972] Diapensia — Tiffney 373
tion on the snow cover gradient to the *dwarf-shrub-rush-
heath community” of Bliss (1963). It is a marginal form
to the Diapensia habitat, being supported by slightly in-
creased snow depth. Again, experimental work should be
performed to see if this form increases in concentration
as a result of induced increases in snow cover in selected
Diapensia habitats. Carex bigelowii and A. groenlandica
are associated about 60° of the time in my samples.
There is doubt as to whether Arenaria groenlandica is
ahnual or perennial (Gleason and Cronquist, 1963). The
plant is described as being associated with mossy areas by
Antevs (1932) and is regarded as being an early colonizer
of bare ground by Antevs (1932) and by Bliss (1963).
The plant is noted as growing on disturbed ground and
particularly in hiking trails by Harris (1964). An annual
or biennial life cycle would offer some explanation for the
affinities of this plant for mossy areas and areas of open
ground. In my tests it was positively associated with un-
colonized ground 54% of the time and with moss 68% of
the time. Open ground provides space to grow in the alpine
zone otherwise occupied by perennial plants or rock. Mossy
areas may promote seed germination and seedling survival
due to greater water availability. Observations made in
this study indicate that A. groenlandica is little controlled
by snow cover, being distributed widely in a number of
habitats which show wide variation in their amount of
winter snow.
Loiseleuria procumbens is a cushion-forming plant very
similar in form to Diapensia lapponica. It is rarely found
in other than Diapensia habitats in the White Mountains.
But, the test of L. procumbens with D. lapponica yielded
a correlation coefficient of only —22% indicating that the
plants were negatively associated —22% of the time. This
value of r (—22%) is not significantly different from 0 at
the .05 (5.0%) conference level.
Of ten vascular plants tested (Table 2) five showed
association with other vascular plants. These were Dia-
pensia lapponica, Vaccinium uliginosum, Juncas trifidus,
374 Rhodora [Vol. 74
Carex bigelowii and Arenaria groenlandica. One of these
five, V. uliginosum covaried with lichen and one, A. groen-
landica, showed interdependence with moss and with un-
colonized ground. One group of non-vascular plants,
lichens, covaried with rock. Five vascular plants showed
no interdependence with other components of the habitat.
These were Salix uva-ursi, Vaccinium vitis-idaea, Potentilla
tridentata, Loiseleuria procumbens and Rhododendron lap-
ponicum. Of ninety-one potential positive or negative inter-
actions tested only eight occurred; none of these eight could
be considered intense.
Discussion
The community concept involves the ideas that there are
interactions among the component plants of the community
and that the species structure of the community follows
some predictable pattern. As Odum (1971) has said, “Com-
munities not only have a definite functional unity with
characteristic trophic structures and patterns of energy
flow but they also have compositional unity in that there
is a certain probability that certain species will occur
together". Both Savile (1960) and Bliss (1962) have
noted that competition in severe habitats is often secondary
to the effects of the physical environment. The result
", .. may be the random occurrence of plants with few
distinct associations" (Bliss, 1962). Savile (1960) sug-
gests that arctic vegetation may better be described by
habitats than by associations. I have used the term habi-
tat, here defined as the place where a plant grows, in this
analysis of an area dominated by Diapensia lapponica.
Sakai (1970) has noted that winter desiccation is a major
limiting factor to alpine plant growth. Sakai and Otsuka
(1970) have shown that Diapensia lapponica, Vaccinium
uliginosum, V. vitis-idaea and Loiseleuria procumbens can
survive very low temperatures. I suggest that Diapensia
and other plants associated with it can survive in this
snow-free and exposed habitat because they can tolerate
very low temperatures and resist desiccation from strong
1972] Diapensia — Tiffney 375
winter winds. Other plants are excluded because they are
not adapted to survive in this harsh physical environment.
Further investigation may provide more information on
such relationships in the alpine zone.
Conclusions
The occurrence of the Diapensia habitat is predictable
in relation to winter snow cover. Diapensia lapponica is
found in areas blown clear of snow throughout the winter
season. Increasing snow cover results in increasing domi-
nance by other forms of plants. The Diapensia habitat is
located at one extreme of the snow cover gradient, and this
study suggests that aggregations characterized by Vac-
cinium uliginosum, Juncus trifidus and Carex bigelowii
may be next in rank with increasing snow cover.
Wide variation was found among concentrations of all
species components over the 22 habitats tested. These
habitats do not consist of predictable percentages of species
components.
Interdependence among the species components of the
Diapensia habitat are few. Of ninety-one potential inter-
actions tested eight occurred at significant levels and none
of these could be considered intense. Control in the habitat
is primarily exercised by the harsh microclimate. Prime
factors are low temperatures and high winds combining
to promote severe winter desiccation in this snow-free area.
Plants which are able to survive in this harsh microenviron-
ment do so and grow here. Others are not fitted to survive
in these conditions and may be excluded by the micro-
environment.
The unpredictable concentrations of plants in the areas
tested, together with their lack of interactions, renders the
use of the term habitat more appropriate than the term
community to describe the aggregate found in this location.
Acknowledgements
I should like to give particular thanks to Dr. Albion R.
Hodgdon for his continued support of this work. Thanks
376 Rhodora [Vol. 74
are also due to the Mt. Washington Observatory for pro-
viding living facilities; and to S. Tiffney, M. Carriker, J.
Roughgarden and J. Davis for their discussion and assist-
ance.
LITERATURE CITED
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Buss, L. C. 1962. Adaptations of arctic and alpine plants to
environmental conditions, Arctic 15: 117-144.
1963. Alpine plant communities of the Presidential
Range, New Hampshire. Ecology 44: 678-697.
1966. Plant productivity in alpine microenvironments
on Mt. Washington, New Hampshire. Ecol. Monog. 36: 125-155.
1969. Alpine community patterns in relation to en-
vironmental parameters in Essays in plant geography and
ecology. K. N. H. Greenridge, ed. Nova Scotia Museum, Halifax.
pp. 167-184.
CourTIN, G. 1968. Microclimate of Mt. Washington. Mt. Wash-
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1968a. Evapotranspiration and energy budget of two
alpine environments, Mt. Washington, New Hampshire. Ph.D.
thesis, Univ. of Ill, Urbana. 172p.
EDDLEMAN, L. E., E. E. REMMENGA and R. T. WARD. 1964 An evalu-
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FERNALD, M. L. 1950. Gray’s manual of botany, eighth ed. Ameri-
can Book Co., N.Y. 1632p.
GLEASON, H. A. and A. CRoNQUIST. 1963. Manual of the vascular
plants of Northeastern United States and adjacent Canada.
D. Van Nostrand Co. Inc., N.Y. 810p.
GoopaLL, D. W. 1970. Statistical plant ecology im Ann, Rev. Ecol.
and Syst, R. Johnson, P. Frank and C. Michener eds. Ann.
Revs. Inc., Palo Alto, Calif. pp. 99-124.
HADLEY, E. B. and L. C. Briss. 1964. Energy relationships of
alpine plants on Mt. Washington, New Hampshire. Ecol. Monog.
34: 331-357.
Harris, S. K. 1964. Mountain flowers of New England. Appala-
chian Mountain Club, Boston. 147p.
HoLTTUM, R. E. 1922. 'The vegetation of West Greenland. Jour.
Ecol. 10: 87-108.
HURLBERT, S. H. 1969. A coefficient of interspecific association.
Ecology 50: 1-9.
1972] Diapensia — Tiffney 377
LóvE, A. and D. LóvE. 1966. Cytotaxonomy of the alpine vascular
plants of Mt. Washington. Univ. Colo. Series in Biol, No. 24.
Univ. Colo. Press. 74p.
PEASE, A. S. 1964. A flora of Northern New Hampshire. New
England Botanical Club Inc., Cambridge, Mass. 278p.
OpuM, E. P. 1971. Fundamentals of ecology, third ed. W. B.
Saunders Co., Philadelphia. 574p.
SAKAI, A. 1970. Mechanism of desiccation damage of conifers
wintering in soil-frozen areas. Ecology 51: 657-664.
. and K. OrsuKA. 1970. Freezing resistance of alpine
plants. Ecology 51: 665-671.
SAVILLE, D. B. A. 1960. Limitations of the competitive exclusion
principle. Science 132: 1761.
SoKAL, R. R. and J. RoHrr. 1969. Biometry. W. H. Freeman &
Co., San Francisco, Calif. 776p.
TRANQUILLINI, W. 1964. The physiology of plants at high altitudes.
Ann. Rev. Plant Physiol. 15: 345-362.
WooDIN, H. E. 1959. Establishment of a permanent vegetation
transact above timberline on Mt. Marcy, New York. Ecology
40: 320-322.
BIOLOGY DEPARTMENT
UNIVERSITY OF MASSACHUSETTS — BOSTON
100 ARLINGTON STREET, BOSTON, MASS. 02116
NEW AND INTERESTING PLANTS
FROM THE GREAT PLAINS
RONALD L. MCGREGOR, LAWRENCE K. MAGRATH
AND RONALD R. WEEDON
As a result of field and herbarium studies, a number of
new or interesting records of plants have been found for
Kansas, Nebraska, South Dakota and Wyoming. A new
form of Sida spinosa L. with white flowers that has been
known in Kansas for several years is described. All speci-
mens cited, unless indicated otherwise, are located at KANU.
Paspalum distichum L. KANSAS: NEOSHO CO.: 5 1/4 min
Erie on US 59, edge of farm pond in water, 18 October
1970, Holland 2854. This represents an extension of the
range of the species northward from Oklahoma.
Panicum philadelphicum Bernh. ex Trin. KANSAS: CHAU-
TAUQUA CO.: 2 mi se Sedan, sandy open scrub oak area,
scattered to common, 2 October 1969, McGregor 20708.
GREENWOOD CO.: 1 mi ne Fall River, sandy open oak
wooded area, abundant in local area, 2 October 1969, Mc-
Gregor 20691. This represents an extension of the range
of the species northward from Oklahoma and westward
from Missouri.
Andropogon intermedius R. Br. NEBRASKA: WEBSTER
co.: 2 mi s, 3 w Red Cloud, reseeded prairie, common, 11
August 1965, MeGregor 19466. In Kansas this species is
known from CLOUD CO.: 1 September 1952, Fraser 956,
and REPUBLIC CO.: 10 September 1970, McGregor 23514.
This species is widespread in tropical and subtropical
areas of the Old World and is introduced in the United
States.
Eleocharis quadrangulata (Michx.) R. & S. var. crassior
Fern. KANSAS: BOURBON CO.: Bourbon Co. St. Lake
No.3, 1 mie & 2 n of Hiattville, common along se side of
lake, soggy area of shoreline over limestone, 7 August
1970, Magrath & Weedon 5750. COFFEY CO.: 6.5 min &
378
1972] Great Plains — McGregor, Magrath & Weedon 379
1.5 w of Westphalia, side of a large pond, muddy loam,
three small clumps, 19 September 1970, Magrath 6250,
NEOSHO CO.: 2 mi s & 2 w of St. Paul, muddy margin of
pond in native meadow, 13 July 1965, Holland 1065.
Scirpus olneyi A. Gray. KANSAS: CHEROKEE CO.: 1.5 mi
w, 0.5 s Baxter Springs, shallow water and muddy mar-
gin of pond, large colony, 17 July 1970, Bare, Johnson &
McGregor 2530; 6 August 1970, Magrath & Weedon 5719
(duplicates determined by T. M. Koyama). This extends
the range of the species northward from Texas and
westward from Missouri where it is known only from
Jefferson County.
Carex melanostachya M. B. ex Willd. KANSAS: DOUGLAS
CO.: 0.5 mi n of Lone Star, roadside ditch, a few large
colonies, 14 May 1964, Kolstad 1193; 25 May 1964, Kol-
stad & Harms 2150 (duplicates determined by F. J. Her-
mann); 24 May 1971, Weedon 6938. This is the first
report of this species for North America. Komarov
(1964) describes the habitat of this species as steppes,
meadows and grassy places on saline soils, and gives its
general distribution as Central Europe, the Mediterra-
nean, Balkans, Asia Minor, Iran, Kulja, and the Caspian
coast of the Caucasus.
Rumex stenophyllus Ledeb. NEBRASKA. BUFFALO CO.: 1
mi s of Elm Creek, sandy flood plain of the Platte River,
14 July 1968, Bare, Johnson & McGregor 1670. CUSTER
co.: 6.5 mi se Anselmo, prairie roadside bar ditch, in
water, sandy soil, common, 20 July 1968, Stephens &
Brooks 25182. DAWES CO.: 1.5 mi ne Crawford, wooded
creek bank, wet sandy soil, weedy area, common, 14 July
1968, Stephens & Brooks 24637. DAWSON CO.: 2 mi s of
Gothenburg, sandy flood plain of the Platte River, par-
tially wooded, 14 July 1968, Bare, McGregor & Johnson
1594. GARDEN CO.: 1 mis & 0.5 mi e of Lewellen, low wet
prairie in the North Platte River valley, 13 July 1968,
Bare, McGregor & Johnson 1443. GREELEY CO.: 10 mi w of
Spalding, sandy banks of Cedar River, common, 28 Au-
380 Rhodora [Vol. 74
gust 1967, Bare & McGregor 419. HARLAN CO.: 4 mis of
Alma, sandy area near lake, locally common, 30 August
1965, McGregor 19568. HOLT CO.: 8 mi s of O'Neill, wet
roadside area, sandy soil, 13 July 1966, Barker 2968.
HOOKER CO.: 16 mi w Mullen, low spot in sandhill prairie,
moist sandy soil, several plants, all small, 27 August 1968,
Stephens 28267. KEITH CO.: 0.5 mi s of Brule, bank of
South Platte River, locally abundant, 28 August 1966,
McGregor 20175. KIMBALL CO.: 6 mi w of Kimball, mar-
gin of lake, common in local area, 30 August 1966, Mc-
Gregor 20269. LINCOLN co.: 4 mi w of Sutherland,
sandy banks of the Platte River, 14 July 1968, Bare,
Johnson & McGregor 1542. SCOTTS BLUFF CO.: 0.5 mi n
of Melbeta, bank of North Platte River, sandy soil with
some white alkali present, 27 September 1968, Weedon
4776. SHERMAN CO.: 3 mi se Litchfield, bank of Beaver
Creek, small colony, 2 September 1965, McGregor 19784.
WEBSTER CO.: 4 mi w of Red Cloud, bank of the Repub-
lican River, large colony, 25 June 1953, McGregor 7545;
3 mi w of Red Cloud, 1 August 1964, McGregor 18676.
Polygonum sawatchense Small. NEBRASKA: SIOUX CO.:
6 mi nw Fort Robinson, open pine forested hillside, 24
June 1970, McGregor 22358. This is an extension of the
range of the species southward from the Black Hills of
South Dakota.
Suckleya suckleyana (Torr. Rydb. KANSAS: CHEYENNE
co.: 0.5 mi w of St. Francis, sandy bank of river, locally
common, 17 August 1971, McGregor 24027. HAMILTON
CO.: 1 mi s of Syracuse, muddy bank of water hole, local
colonies, 18 August 1971, McGregor 24091. KEARNY CO.:
3 mi ne Lakin, muddy shore of Lake McKinney, locally
abundant, 18 August 1971, McGregor 24101. STAFFORD
CO. : Quivira Natl. Wildlife Refuge, open areas in Distich-
lis flats, locally common, 19 August 1971, McGregor
24129.
Paronychia depressa Nutt. KANSAS: CHEYENNE CO.: 12
mi n St. Francis, rocky ledge, 9 June 1967, Stephens
1972] Great Plains — McGregor, Magrath & Weedon 381
11462. This is an extension of the range of the species
southward from Nebraska and eastward from Colorado.
Stellaria aquatica (L.) Scopoli. KANSAS: DONIPHAN CO.:
1 mi n White Cloud, willow thicket, bank of Missouri
River, several large colonies, 29 August 1970, MeGregor
23467. This species is native to Europe. In North Amer-
ica it is known from Quebec and Ontario to North Caro-
lina and Louisiana, also in British Columbia. It is not
known from any of the states adjacent to Kansas.
Cerastium glomeratum Thuillier. KANSAS: CHEROKEE CO. :
6 mi e Baxter Springs, open rocky hillside, 13 Apri] 1946,
McGregor 2735. NEOSHO CO.: 9 mi n, 0.5 e Erie, pasture,
3 May 1965, Holland 876. In the central plains states this
species has been reported from South Dakota, Illinois and
Texas.
Draba lanceolata Royale. SOUTH DAKOTA: LAWRENCE
CO.: 2 mi s, 8 w Walnut, Crooks Tower, top of peak, ca.
7180 ft, open woods, 12 August 1969, Stephens 35914.
This is an extension eastward from the Rocky Mountains
and Montana.
Alyssum desertorum Stapf. SOUTH DAKOTA: FALL RIVER
CO.: 2.5 mi e, 1 s Oelrichs, flat prairie at Limestone
Butte, dry rocky gravel soil, few plants, 20 May 1970,
Stephens 38265. WYOMING: CROOK CO.: 17 mi s Sun-
dance, open prairie hillside, dry rocky clay soil, few
plants, 13 June 1970. WESTON CO.: 7 mi se Osage, flat
prairie pasture, dry gravel clay soil, 12 June 1970,
Stephens & Brooks 39597. This plant is known from
collections in Oregon, Washington, Idaho and Montana.
This is a weed introduced from the Old World, which is
now spreading in North America.
Lesquerella gordonii (Gray) Wats. KANSAS: BARBER CO.:
4 mis Sun City, gypsiferous soil, grassland, 6 May 1956,
Lathrop 2249; 11 mi w Medicine Lodge, red soil, pasture,
2 June 1968, Birkholz 2241. CLARK CO.: 6 mi n Ashland,
sandy soil, 28 April 1939, Horr E250; e of Clark Co. St.
382 Rhodora [Vol. 74
Lake, grassland, dry sandy loam, common, 18 June 1951,
Horr 3969. COMANCHE CO.: 4 mi se Coldwater, rocky
level grassland, 28 April 1939, Horr, s.n.; 2 mi s of Cold-
water, stationary sand dune bank, 28 May 1957, Lathrop
& McGregor 3518. KIOWA CO.: 2 mi e, 3 n Belvidere,
sandy grassland, common, 13 June 1951, Horr & Mc-
Gregor 3808. MEADE CO.: 2 mi s, 11⁄4, e of Meade, over-
grazed rocky pasture, common, 5 May 1957, Horr 4950;
Meade Co. St. Park, sandy grassland, 16 June 1965,
Bare 4. SEWARD CO.: 9 mi ne Liberal, sandy prairie,
locally abundant, 27 April 1965, McGregor 19057. This
is an extension of the range of the species northward
from western Oklahoma.
Saxifraga occidentalis Wats. var. occidentalis. SOUTH
DAKOTA: LAWRENCE CO.: 2.5 mi e, 5 n Savoy at Bridal
Veil Falls, Spearfish Canyon, moist steep rocky bank,
22 June 1970, Stephens 40481. According to Hitchcock,
et al. (1961), the nearest other localities are northwestern
Wyoming and western Montana.
Croton capitatus Michx. NEBRASKA: SARPY CO.: 0.8 mi
n&3 w (on St. Spur N 150) & 2 s on N 50 of Louisville,
loess over sandstone, dry roadside ditch, rare 27 Septem-
ber 1970, Magrath 6362. This extends the range of the
species northward from Kansas.
Croton glandulosus L. var. septentrionalis Muell. NE-
BRASKA: ANTELOPE CO.: 1.4 mi nw of Clearwater on
US 275, prairie roadside & railroad right-of-way, loam,
extremely dry, one plant seen, 27 September 1970,
Magrath 6390A. CASS CO.: 0.5 mi n Louisville, Louisville
Park, flat area around small lake, dry sandy soil, abun-
dant, 5 September 1970, Stephens 44634. These collec-
tions extend the range of the species northward from
Kansas.
Croton monanthogynus Michx. NEBRASKA: ANTELOPE
CO.: 1.5 mi w (on gravel rd) & 1.5 s & w (on paved rd)
of Oakdale, dry roadside near oak woods, sandy loess,
1972] Great Plains — McGregor, Magrath & Weedon 383
scattered, 27 September 1970, Magrath 6381. FRANKLIN
co.: 1 mi w on N 4 & 1.5 s of Campbell, roadside ditch,
dry loam, rare, 29 September 1970, Magrath & Hays
6472. PAWNEE CO.: 1 mis, 5 e Burchard, weedy prairie
pasture, dry rocky soil, 5 September 1967, Stephens
17523. RICHARDSON CO.: 1.4 mi e of Humboldt on N 4,
roadcut & roadside ditch, rocky loam, common, 26 Sep-
tember 1970, Magrath 6335. SARPY CO.: 0.8 min & 3 w
on St. Spur N 150 & 2 s of Louisville on N 50, loess over
sandstone, roadside ditch, uncommon, 27 September 1970,
Magrath 6363. THAYER CO.: 2 mi w of Chester on N 8,
roadside ditch, loam, rare, 29 September 1970, Magrath
6481. WEBSTER CO.: 1.4 mi e of Webster-Franklin Co.
line on N 4, disturbed area in an open prairie hay mea-
dow, loam, rare, 29 September 1970, Magrath & Hays
6475. These collections extend the range of this species
northward from Kansas.
Callitriche terrestris Raf. emend. Torr. KANSAS: MONT-
GOMERY CO.: 12 mi sw Independence, at Camp Wilder-
ness, along damp draw in oak woods, 20 June 1970, Hol-
land 2812c, in part. This is an extension of the range of
the species westward from Missouri and northward from
Oklahoma.
Anoda cristata (L.) Schlect. KANSAS: NEOSHO CO.: in
South Mound, Lincoln Township, farmlot, 30 September
1970, Holland 2848. This species is native to South
America and the southwestern United States. In the
midwest, it is known to occur in Iowa and in McDonald
Co., Missouri.
Sida spinosa L. f. albiflora Magrath, form. nov.
A forma spinosa differt petalis albis.
TYPE: KANSAS: DOUGLAS CO.: West campus of Uni-
versity of Kansas in Lawrence, growing along e edge of
experimental greenhouse in loam soil, colony of 50-70
plants, 7 September 1970, Magrath 6147 (Holotype:
KANU!). The white flowered plants were scattered in a
large colony of plants that had typical yellow flowers.
384 Rhodora [Vol. 74
This white flowered form of S. spinosa was first observed
in this same area by Vernon Harms in 1962. Harms
(1965) performed crossing studies on this taxon and
concluded that the white flowered form represented a
homozygous recessive condition for a single gene.
Hibiscus lasiocarpos Cav. KANSAS: BOURBON CO.: 1.5 mi
e & 3 n of Hiattville, Bourbon Co. St. Lake, growing in
marshy area along w side of lake, common, 25 August
1969, Magrath 4588; 7 August 1970, Magrath & Weedon
5746. CHEROKEE CO.: 2 mi s of Galena, in mixed woods
of the flood plain woods of Shoal Creek, 5 August 1956,
Lathrop 2863; 1 mi s of Riverton, low swampy area,
abundant in local area, 12 August 1960, McGregor 15924;
swampy area along Spring River, 26 August 1969, Bare
2169. The Bourbon Co. collections represent the first
area in Kansas other than se Cherokee Co. where this
species is known to occur.
Thymelaea passerina (L.) Cosson & Germ. KANSAS:
CLOUD CO.: 1 mi n Concordia, flood-plain of Republican
River, sandy soil, willow thicket, scattered, 11 September
1970, McGregor 23572. This species has been reported
for North America from west-central Iowa, northeastern
Nebraska and extreme western Nebraska. European
manuals indicate that the species is a common weed in
dry soils and appears commonly in grain fields of south
and central Europe and western Asia.
Centunculus minimus L. KANSAS: MONTGOMERY CO.: 12
mi sw Independence, at Camp Wilderness, along damp
draw in oak woods, 20 June 1970, Holland 2812c, in part.
This represents an extension of the species westward
from Missouri and northward from eastern Oklahoma.
Glaux maritima L. NEBRASKA: SIOUX CO.: n edge of
Agate, flat prairie swale near Niobrara River, sandy
soil, 16 August 1970, Stephens 43966. This species occurs
in North Dakota, but is not known from South Dakota.
1972] Great Plains — McGregor, Magrath & Weedon 385
Sabatia campestris Nutt. f. albiflora D. M. Moore. KAN-
SAS: CHEROKEE CO.: 0.4 mi w & 1 s of Baxter Springs,
in prairie in soil over limestone, abandoned rock quarry
area, 17 July 1969, Johnson 2259, in part. MONTGOMERY
co.: 2 mi s & 0.2 w of Sycamore, prairie hay meadow,
sandy clay soil, rare, 12 July 1969, Weedon & Magrath
5516. WILSON CO.: 4.5 mi e Neodesha, sandy blue stem
prairie, scattered, 4 July 1970, McGregor 22386.
Eustoma grandiflorum (Raf.) Shinners f. album (Hol-
zinger) Waterfall KANSAS: RICE CO.: 0.5 mi s of
Sterling, heavily grazed pasture, sandy soil, one plant
seen, 9 August 1970, Magrath & Weedon 5754. STAFFORD
co.: T218, R11W, Sec 21, found in dense Spartina area,
very rare, 13 August 1958, Ungar 666a.
Apocynum medium Greene. KANSAS: DOUGLAS CO.: 14.
mi s of Lone Star, thin soil over limestone, abundant on
old slope of roadcut, 11 June 1969, Johnson 2081; 14
June 1970, Johnson 2496. This extends the range of this
species westward from Missouri.
Asclepias incarnata L. var. incarnata f. albiflora Heller.
KANSAS: COFFEY CO.: 6.5 mi n & 1.5 w of Westphalia,
growing near edge of large pond, moist loam, one plant,
5 September 1970, Magrath 6136. DOUGLAS CO.: in old
roadbed at southern end of Baldwin Lake, 15 August
1961, Richards 3686; 1 mi s & 1.5 e of Baldwin City,
growing at edge of lake, six plants, 19 August 1969,
Bare 2114. NEBRASKA: DODGE CO.: Fremont Lakes
Rec. Area, 3 mi w of Fremont on US 30, growing along
se edge of lake, one white flowered plant in large colony
of pink flowered plants, sandy loam, 21 August 1970,
Magrath 5978.
Hydrophyllum appendiculatum Michx. NEBRASKA: DOUG-
LAS CO.: Omaha, wooded ravine s of Hummel Park,
6 June 1968, Sutherland 1689. This extends the range
of this species northward from Kansas and northwest-
ward from Missouri.
386 Rhodora [Vol. 74
Scutellaria brittonii Porter. NEBRASKA: KIMBALL CO.:
2 mi w Kimball, upland prairie, dry sandy-gravel soil,
14 June 1967, Stephens 11664; 8 mi sw, 1.5 w Bushnell,
open pine-prairie hilltop, dry rocky clay soil, few plants,
5 June 1969, Stephens & Brooks 31479; 29 June 1970,
Stephens & Brooks 40859. This species also occurs in
Wyoming, Colorado and New Mexico.
Leonurus marrubiastrum L. NEBRASKA: DOUGLAS CO.:
4 mi e of Venice on N 92, Elkhorn River, slip-off bank
s of bridge, muddy silt with some sandy spots, rare, 21
August 1970, Magrath 5968. NEMAHA CO.: 8.1 mi s of
Auburn on US 73 & 75, riverbed e of bridge, slip-off bank,
silty-sandy clay, scattered, Magrath 5931. SOUTH DA-
KOTA: HUTCHINSON CO.: 0.5 mi e Olivet, bank of James
River, moist sandy soil, abundant, 8 September 1970,
Stephens 44866. This extends the range of the species
northward from Kansas and northwestward from Mis-
souri.
Lamium amplexicaule L. f. albiflorum D. M. Moore. KAN-
SAS: DOUGLAS CO.: City of Lawrence, 2 blocks n of
23rd St. & 1 block e of Pennsylvania St., one plant, grow-
ing in pasture, loam, 30 April 1971, W. & T. Vollmer, s.n.
COFFEY CO.: 1 mi n, 7 w on US 50 of Waverly, roadside
park, growing in ditch, loam, uncommon, 22 May 1972,
Magrath 7407.
Solanum sarrachoides Sendl. KANSAS: NEOSHO CO.: 4 mi
s&be Erie, in farmyard near fence rows, 29 September
1970, Holland 2840. This is a South American species
that has been introduced into North America. It is
known otherwise only from Jasper and McDonald Coun-
ties in Missouri and northeastern Oklahoma.
Mimulus alatus Ait. f. albiflorus House. KANSAS: FRANK-
LIN CO.: Ferndale, 1.5 mi s, 1 e & 1!4 s of Homewood,
muddy creek floodplain near base of sandstone bluffs, one
plant, 29 August 1970, Magrath 6125. LINN CO.: stream
channel just n of Mound City, just e of bridge on K 7,
1972] Great Plains — McGregor, Magrath & Weedon 387
surfacing limestone, rare, 7 August 1970, Weedon &
Magrath 6546.
Agalinis heterophylla (Nutt.) Small. KANSAS: CHAU-
TAUQUA CO.: 1.6 mi ne of Sedan, oak woods over sand-
stone, rare, 14 September 1970, Magrath 6224. CHEROKEE
CO.: 2.1 mi w of Jct of US 166 & 69 on 166, 1.6 s & 0.2
e, abandoned rock quarry, rocky limestone soil, common,
14 September 1970, Magrath 6243; 1 mi w Baxter
Springs, scattered in prairie, 3 October 1970, MeGregor
22641. GREENWOOD CO.: 0.6 mi e of Sallyards on US 54,
rocky prairie hay meadow, one plant found, 13 Septem-
ber 1970, Magrath 6201A. This extends the range of the
species westward from Lawrence and Dunklin Counties
in Missouri and northward frcm Oklahoma.
Orobanche multiflora Nutt. KANSAS: SCOTT CO.: 12 min
& 3 w of Scott City, gravelly prairie hillsides with caliche
outcrops, only one plant seen, 3 October 1970, Bare 2605.
This extends the range of the species northward from
Oklahoma and eastward from Colorado.
Triodanis biflora (R. & P.) Greene. NEBRASKA: JOHN-
SON CO.: 3.2 mis of Cook on N 50, grazed pasture, loam,
rare, 28 June 1970, Magrath & Hays 5639A. PAWNEE
CO.: 0.6 mi s of Pawnee-Johnson Co. line on N 50, prairie
hay meadow, in disturbed area, loam, rare, 28 June 1970,
Magrath & Hays 5642. This is an extension northward
of this species from Kansas.
Lobelia siphilitica L. var. siphilitica f. albiflora Britt.
KANSAS: DOUGLAS CO.: 1.5 mi e of Lecompton, Kansas
River bluff, moist ditch, 19 September 1968, Bare 1944.
Iva axillaris Pursh. KANSAS: FINNEY CO.: river bank,
18 July 1912, Wilson & Miller, s.n. HAMILTON CO.: 1
mi s, 1 e of Coolidge, prairie roadside right-of-way, moist
sandy soil, locally abundant, area 25 ft. across, 3 August
1971, Stephens 50368. These are extensions of the range
of this species southward from Nebraska and eastward
from Colorado.
388 Rhodora [Vol. 74
Helianthus mollis Lam. NEBRASKA: RICHARDSON CO.:
1.4 mi e of Humboldt on N 4, prairie pasture, loam, un-
common, 26 September 1970, Magrath 6336A. WEBSTER
co.: 2 mi s, 3 w Red Cloud, sandy prairie hillside, rare
in the area, 11 August 1965, McGregor 19479. This is
an extension of the range of the species northward from
Kansas.
Bidens comosa (Gray) Wieg. WYOMING: GOSHEN CO.:
side channel of North Platte River w of city park
in Torrington, sandy to clay loam soil, scattered, 28
September 1968, Weedon 4787; North Platte River on
s edge of Torrington, both banks of river, sandy soil,
scattered, 28 September 1968, Weedon 4789. WASHAKIE
co.: Worland, in a wet saline area, 16 July 1966, Tresler
322 (RM). According to Sherff (1937), the distribution
of this species is from Maine, Quebec and North Dakota
southward and southwestward to North Carolina, Ten-
nessee, New Mexico and Utah. It has also been reported
from eastern Colorado.
Artemisia annua L. KANSAS: DONIPHAN CO.: 1 mi n
White Cloud, bank of Missouri River, willow thicket,
several large colonies, 29 August 1970, McGregor 23467.
This is the second known record for the state.
LITERATURE CITED
BARKLEY, T. M. 1968. A Manual of the Flowering Plants of
Kansas. The Kansas St. Univ. Endowment Assoc. Manhattan,
Kansas.
Harms, V. L. 1965. Observations on the Breeding System of Sida
spinosa (Malvaceae). Trans. Kan. Acad. Sci. 68(1): 123-24.
HircHCOCK, C. L., A. CRONQUIST, M. OwNBEY, J. W. THOMPSON.
1961. Vascular Plants of the Pacific Northwest. Part 3: Saxi-
fragaceae to Ericaceae. Univ. of Washington Press, Seattle.
Komarov, V. L. ed. 1964. Flora of the U.S.S.R. Vol. III. pp. 328-
29. Pub. for Smithsonian Inst. and NSF by the Israel Program
for Scientifie Translations, Jerusalem.
SHERFF, E. E. 1937. The Genus Bidens. Field Mus. Nat. Hist.
Bot. Ser. 16: 251.
DEPARTMENT OF BOTANY
UNIVERSITY OF KANSAS,
LAWRENCE 66044
NEW RECORDS OF NORTH DAKOTA PLANTS
O. A. STEVENS
In 1961 I gave a ten-year supplement to my Handbook
(Stevens 1950), which was incorporated as an (unpaged)
appendix to the third printing of the book (1963). The
intention was to provide a second supplement in 1970, but
at that date active collecting was proceeding, especially in
the western part of the state and this supplement was post-
poned. In the meantime several partial lists have appeared
and it seems desirable not to postpone this one longer.
In addition to species which are reported for the first
time (marked with *), a few significant additional loca-
tions for other species are included. Mere changes in names
are usually not mentioned.
Botrychium lunaria L. was found in Burke County in
1971 by Glen Hegstad.
*Equisetum palustre L. was reported by Barker (1971)
for Ransom County.
*Equisetum scirpoides Michx. was reported by Stewart
(1970) from Barnes County.
*Pinus banksiana Lamb. Several cone-bearing and seed-
ling trees were found near McLeod, in Richland County in
1970 (Gerald Seiler 1775). This is presumably a forest
planting but the record has not been found.
*Bromus squarrosus L. Abundant in an abandoned,
sandy field, Kidder County in 1971, Stevens 3292, 3294.
*Chloris virgata Sw. This probably was an escape from
S C S nursery at Mandan, Morton County, September 13,
1962, Geo. Karey.
*Cinna arundinacea L. was collected at two locations in
Ransom County, Stevens 2742, 2761.
*Eragrostis spectabilis (Pursh) Steud. was reported by
Barker (1971) from Ransom County.
*Polypogon monspeliense L. is recorded from LaMoure
County, H. Kantrud, September 30, 1970; Slope County in
1970, Zaczkowski 4000; Williams County in 1971, G. D.
Hegstad.
389
390 Rhodora [Vol. 74
*Triplasis purpurea (Walt.) Chapm. was collected in two
places in Richland County in 1964 (W. Wanek 124).
*Sporobolus airoides Torr. is confirmed by N. Zaczkow-
ski 3012, 3017, and 4257.
*Carex alopecoidea Tuckerman was reported by Stew-
art (1970). I had tentatively placed here a specimen from
Richland County, Stevens 88. O. A. Kolstad at the Univer-
sity of Kansas, who examined all of our North Dakota
Carex in 1965, put this as C. stipata but part of a sheet
of stipata in C. alopecoidea. Presumably all five stalks were
from one clump (which proves nothing). Kolstad referred
to alopecoidea one specimen from Barnes County, Stevens
in 1950.
*Carex capillaris L. Bottineau County in 1961, Stevens
2499.
Carex davisii Schw. & Torr. Our specimens reported as
C. gracillima were placed here by Kolstad. Stewart (1970)
reported *C. buxbaumii, *C. lacustris, *C. lasiocarpa, *C.
limosa and *C. vesicaria.
*Iris missouriensis Nutt. A specimen was received, but
not saved, from Mrs. Paul DeKrey, Tappen, Kidder County
in 1968. G. D. Hoag visited the place and reported a num-
ber of clumps in a depression of the prairie. Richard Wil-
liams found it in northeastern Emmons County in 1971.
*Juglans nigra L. can be regarded as naturalized because
it is often planted and squirrels carry the nuts to other
places. At least three seedlings appeared in our yard, pre-
sumably from nuts from a tree across the street (one, re-
planted about 15 years ago is still growing) and I have
seen others at several places.
*Juncus vaseyi Engelm. A specimen from Bottineau
County was determined by L. J. Harms.
Quercus ellipsoidalis E. J. Hill. Little (1971, map 163
E) shows this extending into Richland County but we have
no knowledge of such (may be planted).
Atriplex glabriuscula Edmonst. The specimens reported
as this (Stevens, 1963) are now referred to A. hetero-
sperma Bunge by Aellen and by Frankton (1968) who
1972] North Dakota Plants — Stevens 391
uses A. subspicata (Nutt.) Rydb. for what we had called
A. hastata, but retained one specimen from Richland
County, Bell 377, as A. hastata, also one from Clay County,
Minnesota.
*Lychnis chaleedonica L. We have two Specimens, pre-
sumably escapes. Benson County, J. Lunell in 1917 “bor-
dering a slough”, and Dickey County, K. M. Larson in
1965.
*Salix pedicillata Pursh was collected in McHenry
County, Stevens 3225, in the fen cited by Stewart (1970).
Eriogonum gordoni Benth. was reported as E. trichopes
(Stevens 1960) but apparently is E. gordoni or close to it.
Further specimens are from Burke and Mountrail Counties
in 1970 by Hegstad.
Anemone multifida Poir. from Golden Valley County was
a surprise, N. Zaczkowski 364, 3481.
* Ranunculus cardiophyllus Hook. A specimen collected
in McKenzie County in 1967 by C. A. Barr seems to be this
species.
Alyssum desertorum Stapf. apparently is the proper
name for that reported (Stevens 1963) as A. alyssoides.
Better specimens are from the same county by Zaczkowski.
*Arabis glabra Gray. Two specimens from Dunn County,
Stevens & Moir 2255 and Sept. 14, 1956, and one from
Burke County, Stevens, July 20, 1966, seems to belong here.
Chorispora tenella (Pall.) D.C. A second record is from
Billings County, Zaczkowski 3333.
“Erysimum repandum L. A single specimen collected on
the campus in 1967 seems to be this species.
*Berberis repens Lindl. was reported by Barker (1971)
from Bowman County.
*Ribes cereum Dougl. A sterile specimen was collected
by Jack Nelson at Pretty Butte, western edge of Slope
County, in 1960. Nick Zaczkowski collected flowering
specimens there in 1970.
*Potentilla palustris L. was reported by Stewart (1970)
from McHenry County.
*Astragalus purshii Dougl. was reported by Zaczkowski
392 Rhodora [Vol. 74
(1970). It was identified by Fernald from a specimen
which I collected in McKenzie County July 15, 1943. An-
other was collected in Emmons County in 1945. Walsh
(1960) referred these to A. lotiflorus.
Caragana arborescens Lam. Glen Hegstad, who has been
studying our northwestern counties extensively, reports
apparent establishment. This area had many farmsteads
abandoned after 1934.
Lotus corniculatus L. has become quite common along
streets and lawns in the Fargo area.
Polygala senega L. Three collections by Hegstad in
Burke County, 4770, 5413, 8038, extend the known range
of this species.
* Euphorbia stictospora Engelm. A specimen, Bell 1072,
said to have been collected in the dooryard of a ranch in
1907 in McKenzie County was determined as this by Rich-
ardson (1968). It must have been a chance introduction.
*Rhamnus davurica Pall. In 1965 I saw in Ransom
County a windbreak of this, containing great numbers of
seedlings. A second record is from O. J. Lung in 1964
from Emmons County (“came up in garden”). The fruits
seem not usually eaten by birds.
Acalypha rhomboidea Raf. A second specimen is from
Stutsman County, Sept. 3, 1964, Mrs. Amy Rolling, “came
up in flower bed".
*Acer saccharinum L. is often planted as a street tree.
Two to three year old seedlings were collected by a hedge
near a large maple tree in 1970, Stevens 3285.
*Lavatera cretica L. was collected in Divide County on
Aug. 20, 1969 by L. M. Mitich and by Hegstad (6239) in
Burke County in 1970; escaped from, or persisting from,
gardens.
Hypericum perforatum L. A second specimen, Stevens
3152, in Ransom County on the Sheyenne River bank by an
old farm house, may have been planted. There were sev-
eral plants but no new ones.
*Gaura parviflora Dougl. reported by Barker (1971)
1972] North Dakota Plants — Stevens 393
from Golden Valley County is probably a railroad intro-
duction.
Conium maculatum L. was collected in Golden Valley and
Billings Counties in 1969 and 1970 by Zaczkowski.
Osmorhiza claytoni (Michx.) C. B. Clarke. A second
record was by V. Facey, Grand Forks County in 1958.
*Halenia deflexa (Sm.) Griseb. was reported by Barker
(1971) from Pembina County (KANU).
Phlox alyssifolia Greene was collected on Bullion Butte,
Stark County, Zaczkowski 2329 in 1970.
Physostegia parviflora Nutt. I am inclined to think that
our plant should be considered a variety of P. virginiana
L. It is uniformly leafy, the leaves 1-2 (—5) em wide, sharp-
ly toothed, flowers 15 (—18) mm long. Specimens of P.
formosior Lunell, so labeled by Lunell, do not differ mate-
rially, but one from the same area labeled P. parviflora has
the lower leaves narrowly oblong with blunt, low teeth.
One from McLean County (V. T. Heidenreich 210) has
leaves about 7 mm wide and some from Burke County
(Hegstad 8458) have narrow leaves and blunt teeth but
his 5943 from within ten miles of the same place agrees
with our eastern specimens.
*Lythrum salicaria L. is much planted and a consider-
able colony develeped in a low, roadside area at Fargo
(Stevens 2332). I found one plant at a similar place about
one mile distant.
Stachys palustris L. 'The prevalent form in open, usually
moist ground, has rather densely, evenly pilose stems (S.
scopulorum Greene?). Along wooded streams in the east-
ern edge of the state the mid and lower stems often are
nearly glabrous, with a few, retrorse, coarser, often some-
what pustulate, hairs on the angles and a few or more soft
hairs on the sides. The calyx varies from sparsely to more
densely pilose, rarely with short, glandular hairs.
*Collinsia parviflora Lindl. was found in Slope County,
Zaczkowski 3072.
*Pentstemon procera Dougl. was reported by Barker
(1971) from Burke County.
394 Rhodora [Vol. 74
*Chaenorrhinum minus (L.) Lange was collected along a
formerly used railway track at Valley City, Barnes County
in 1963, Stevens 2670. The area was pastured and plants
not seen in 1971.
Veronicastrum virginicum (L.) Farwell. Roys Willen-
bring, collecting in Pembina County 1969-71, found a few
plants near Cavalier, Pembina County, 799.
*Galium verum L. One specimen is from Dickinson,
Stark County in 1912, C. H. Waldron 130. I recall that
perhaps about 1935, Dr. L. R. Waldron showed me a frag-
ment of it. In 1967 I discovered a colony of it in a yard
near his house, and distributed several sheets, Stevens
3028. In 1970 I looked again but did not find it.
*Sambucus pubens Michx. In 1964 I visited a farm grove
near Colfax, Cass County, where this plant had apparently
been introduced and spread through the grove and along
fence lines. Some years ago it had spread through the uni-
versity’s woody plantings but was eradicated.
* Ambrosia (Franseria) tomentosa (A. Gray) Payne. A
colony appeared in garden plots of the University at Fargo
in 1963 (Stevens 2693). Another specimen is from Grand
Rapids, LaMoure County, Bill Shak, Aug. 21, 1921.
Aster ciliolatus Lindl. This name apparently should be
used for what has been called A. sagittifolius L. I have
been unable to make satisfactory conclusions.
Bidens acuta (Wieg.) Britton. I still feel that this is a
better “species” than many others. In our area it is quite
striking, usually with sessile leaves, heads the size of those
of B. vulgata with 2-3 cm long, leafy, outer bracts. The
long-petioled leaves of B. comosa are lacking.
Centaurea maculosa Lam. A large colony of an acre or
more was found along State Highway 1 in Nelson County
at the outlet of Stump Lake in 1962, Stevens 2962.
*Chaenactis douglasii (Hook.) H. & A. This is another
extension from the west in Bowman County, Zaczkowski.
Crepis tectorum L. I had used the name C. capillaris,
first on a specimen by Lunell from Bottineau County, but
I believe it is C. tectorum which is quite common in the
1972] North Dakota Plants — Stevens 395
Cavalier-Pembina area and spreading more widely. It is
also abundant in western Becker County, Minnesota.
*Erigeron canbyi. Specimens from the Killdeer Mts.,
Dunn County, by C. A. Barr, May 8, 1963, were considered
this form by Mr. Barr.
Hymenopappus tenuifolius Pursh. Collections from Ben-
son County, W. T. Barker 5567, make a notable eastern
range extension.
*Senecio vulgaris L. was received from Dickinson, Stark
County in 1954; Stevens 1558 in 1955. Another garden lo-
cation is at Fargo, Cass County in 1962, Stevens 2648,
2870. It had been found at Hawley, Clay County, Minne-
sota.
*Thelesperma marginatum Rydb. was collected in Wil-
liams and Divide Counties, Hegstad 3095, 3552, 5'798, 8030,
in 1969-71.
REFERENCES
BARKER, W. T. 1971. Vascular plants new to North Dakota.
Rhodora 73: 58, 59.
FRANKTON, C. and I. J. BassETT. 1968. The genue Atriplex in
Canada. I. Three introduced species — Canad. Jour. Bot. 46:
1309-1313.
LITTLE, E. J. 1971. Atlas of United States Trees. VoL 1. UTS.
Dept. Agr. Misc. Publ. 1146.
RICHARDSON, J. W. 1968. The genus Euphorbia in the high plains
of Kansas and prairie plains of Kansas, Nebraska, South and
North Dakota. Univ. Kan. Bull. 48: 45-112.
STEVENS, O. A. 1950. Handbook of North Dakota Plants. Institute
of Regional Studies, North Dakota State University, Fargo.
- 1961. New records of North Dakota plants. Rho-
dora 63: 39-46.
1963. Handbook of North Dakota Plants. Insti-
tute of Regional Studies, North Dakota State University, Fargo.
(Third printing)
STEWART, R. E. 1970. New records of vascular plants in North
Dakota. The Prairie Naturalist 3: 35-48,
Warp, R, and W. T. BARKER. 1971. Cytotaxonomic studies of the
Scirpus lacustris complex in North Dakota. Proc. N. Dakota.
Acad. Sci. 25: 32.
396 Rhodora [Vol. 74
WELsH, S. L. 1960. Legumes of the north-central states: Galegea.
Iowa St. Coll. Jour. Sci. 35: 111-249.
ZACZKOWSKI, N. K. and W. T. BARKER. 1971 (abstr.). Floristic
survey of southwestern North Dakota. Proc. N. Dakota Acad.
Sci. 25: 32.
NORTH DAKOTA STATE UNIVERSITY
FARGO, N. D. 58102
PRIORITY OF
THE BINOMIAL PYROLA CHLORANTHA
ERICH HABER
Some confusion exists in the literature concerning the
usage and priority of the binomials P. chlorantha Swartz
and P. virens Schreber in Schweigger & Koerte. The most
commonly used binomial for the green-flowered shinleaf in
Europe up to and including the present time, and in North
America until 1941, was P. chlorantha (Swartz, 1810).
Fernald's short note (1941) in which he cited P. virens as
antedating P. chlorantha by six years resulted in the com-
mon acceptance of the former name in most North Amer-
ican publications. However, the Flora Erlangensis of 1804
does not mention Pyrola virens.
It becomes clear, however, that P. chlorantha has priority
over P. virens when the publication containing the orig-
inal description of P. virens is closely examined ( Schweig-
ger & Koerte, 1811). The 1811 publication, Flora Erlan-
gensis, is composed of two parts plus an Addenda et Cor-
rigenda to Part I. The pagination of the addendum is
sequential to that in Part I which is taken directly from a
doctoral dissertation by Schweigger (1804).
A. F. Schweigger and F. Koerte published Flora Erlan-
gensis in 1811 at which time the addendum as well as Part
II were bound with a facsimile of the 1804 publication.
The title page for the addendum bears neither of the au-
thors’ names, whereas the author of Part I is clearly indi-
cated as being Schweigger and the authors of Part II as
being Schweigger and Koerte. In the preface to the com-
1972] Pyrola — Haber 397
plete volume written by Koerte, it is stated that all the
additions which are preceded by an asterisk are the work
of J. C. D. von Schreber, who had reviewed the manuscript.
P. virens on p. 154 of the addendum is preceded by an
asterisk. There appears to be no question that the valid
date of publication of P. virens is 1811, one year after the
publication of the binomial P. chlorantha. The correct ci-
tation for the former is as follows: P. virens Schreber in
Schweiger & Koerte, Fl. Erlang. I, add. p. 154. 1811.
The author would like to express his appreciation to Dr.
John W. Grear Jr. for his assistance in obtaining repro-
ductions of pertinent sections of the origina] publications
by Schweigger and Koerte.
LITERATURE CITED
FERNALD, M. L. 1941. Transfers in Pyrola, Rhodora 43: 167.
ScHWEIGGER, A. F. 1804. Specimen Florae Erlangensis. Erlangen,
Hilpert.
ScHWEIGGER, A. F. & F. KoERTE. 1811. Flora Erlangensis. Erlan-
gen, Palm.
Swartz, O. P. 1810. Sv. Vet. Acad. Stockh. Handl. 31: 190.
NATIONAL MUSEUM OF NATURAL SCIENCES
BOTANY DIVISION
OTTAWA, ONTARIO K1A0MS8
CHROMOSOME NUMBERS OF
SCIRPUS PURSHIANUS AND S. SMITHII
ALFRED E. SCHUYLER
Two eastern North American aquatic bulrushes, Scirpus
purshianus Fern. and S. smithii Gray, superficially resem-
ble one another and have been confused in taxonomic lit-
erature. Beetle (1942) stated that, “Apparently S. smithii
is no more than an aquatic extreme of S. debilis [an earlier
name applied to plants of S. purshianus]," while Fernald
(1942) maintained, “. .. all living botanists who accurately
know living plants in the field find them abundantly dis-
tinct." Some botanists (Gleason and Cronquist, 1963)
have continued to treat these species in accordance with
Beetle's statement and others (Seymour, 1969) have con-
formed with Fernald's statement. The characteristics used
by Fernald to delineate S. purshianus, S. smithii, and their
forms are accepted here and the following discussion is in
accordance with his treatment of these taxa in Gray’s
Manual (Fernald, 1950).
Part of the taxonomic confusion with S. purshianus and
S. smithii is probably caused by the similar pattern of in-
fraspecific variation of each species. Each has ecologically
differentiated forms characterized morphologically by the
presence or absence of well-developed barbed bristles at-
tached to the achenes. S. smithii, which often appears to
be associated with somewhat alkaline substrates, has three
described forms: f. levisetus (Fass.) Fern. (having nearly
smooth bristles), f. setosus (Fern.) Fern. (having well-
developed barbed bristles), and f. smithii (lacking bris-
tles). S. purshianus, which often appears to be associated
with more acid substrates, has two forms: f. purshianus
(having well-developed barbed bristles) and f. williamsii
(Fern.) Fern. (lacking bristles).
In both S. purshianus and S. smithii, the presence or ab-
sence of barbed bristles appears to be correlated with the
stability of the habitat. For example in eastern New York,
New Jersey, eastern Pennsylvania, and Delaware, S. smithii
398
1972] Chromosome Numbers — Schuyler 399
f. smithii is almost completely restricted to the specialized
conditions of fresh inter-tidal zones of the Hudson, Raritan,
and Delaware Rivers while in the same area f. setosus
occurs in and around lakes, ponds, and quagmires (see
Long, 1910, for a discussion of these forms in the vicinity
of Philadelphia). S. smithii f. levisetus is common in the
estuaries of the Kennebec River in Maine and the St. Law-
rence River in Quebec. Although bristles are present in
f. levisetus, they lack the prominent barb-like teeth char-
acteristic of f. setosus. Thus it appears that plants of S.
smithii which lack barbed bristles are better adapted to
the periodic fluctuation in water level characteristic of the
estuarine environment than are those which have barbed
bristles. A similar pattern emerges for S. purshianus f.
purshianus and f. williamsii, although tidal fluctuation in
water level is not involved since neither form grows in
tidal estuaries. However, f. williamsii is frequently found
on the sandy shores of lakes in Cass, Kalamazoo, and Van
Buren Counties in southwestern Michigan while the more
widespread f. purshianus is infrequently found in this area.
There is (or was) considerable water level fluctuation in
these southwestern Michigan lakes from spring to fall, a
factor which probably is a barrier to plants adapted to
lake shores with more uniform water levels, This may be
what excludes f. purshianus from many southwestern
Michigan lakes since, throughout most of its range, it often
grows along lake shores with relatively uniform water
levels. Thus the forms of both species having barbed bris-
tles appear better adapted to relatively stable conditions
associated with little or moderate water level fluctuation
while the forms lacking barbed bristles appear better
adapted to relatively unstable conditions associated with
substantial water level fluctuation.
Despite the ecological and morphological resemblance be-
tween forms of these two species, I have always found that
the morphological criteria used by Fernald to distinguish
them are reliable. In addition, the chromosome numbers
determined for the forms of each species provide evidence
[Vol. 74
Rhodora
‘(9224 Lajfinyog) ijs ‘S Jo soMOsOWOLYyD 9n019]p— "6 SIT
'(e9et «4ojfim[og) snupu[S4nd “gS Jo səwosowoaryəð 2120I9]A
T SA
1972] Chromosome Numbers — Schuyler 401
supporting Fernald’s treatment. 19 meiotic units (Fig. 1)
were observed in pollen mother cells of S. purshianus f.
purshianus and f. williamsii (Table 1). 20 meiotic units,
with one unit consistently larger than the rest (Fig. 2),
were observed in pollen mother cells of S. smithii, f. levi-
setus, f. setosus, and f. smithii (Table 1). Ordinarily,
chromosome numbers are consistent for a given species of
Scirpus and provide evidence corroborating distinctions
among them. The chromosome numbers reported here sup-
port the specific recognition of S. purshianus and S. smithii
and indicate that the infraspecific taxa are associated with
the proper species in Fernald's treatment.
TABLE 1. Chromosome Numbers Determined for Forms
or Scirpus purshianus and S. smithii
Number of
Taxon Meiotic Units Voucher Specimens?
S. purshianus
f. purshianus 19 Massachusetts: Barnstable Co.,
4265; Pennsylvania: Northamp-
ton Co. 3819; Pennsylvania:
Berks Co., 4088
S. purshianus
f. williamsii 19 Michigan: Kalamazoo Co., 3903
S. smithii
f. levisetus 20 Maine: Sagadahoc Co., 4276
S. smithii
f. setosus 20 New Jersey: Sussex Co., 4201
S. smithii
f. smithii 20 New Jersey: Burlington Co.,
4203
^A]] are collections by the author preserved at the Academy
of Natural Sciences.
ACKNOWLEDGMENTS
I am grateful to George H. Lauff for use of facilities at
the W. K. Kellogg Biological Station during the summer of
402 Rhodora [Vol. 74
1966. Vincent Abraitys, Florence M. Givens, Patricia R.
Schuyler, and Henry K. Svenson helped in connection with
field work, Karen Keil did chromosome squashes, and
Helena Greenwood typed the manuscript.
LITERATURE CITED
BEETLE, A. A. 1942. Studies of the Genus Scirpus L., V. Notes on
the Section Actaeogeton. Am. Journ. Bot. 29: 653-656.
FERNALD, M. L. 1942. Transfers in Scirpus $ Actaeogeton. Rho-
dora 44: 479-484.
1950. Gray's Manual of Botany. 8th ed. Ameri-
can Book Co. New York. lxiv + 1632 pp.
GLEASON, H. A. & A, CRONQUIST. 1963. Manual of Vascular Plants
of Northeastern United States and Adjacent Canada. D. Van
Nostrand Co., Ine., Princeton. li + 810 pp.
LONG, B. 1910. Range Extension of Scirpus smithii var. setosus.
Rhodora 12: 155-156.
SEYMOUR, F. C. 1969. The Flora of New England. Charles E.
Tuttle Co., Rutland. xvi + 596 pp.
DEPARTMENT OF BOTANY
ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA
PHILADELPHIA, PENNSYLVANIA 19103
TUSSILAGO FARFARA L.
(SENECIONEAE-ASTERACEAE)
IN VIRGINIA
MILES F. JOHNSON
Tussilago farfara, the Coltsfoot, a perennial rhizomatous
herbaceous native of the Old World with a single yellow
head terminating a scaly bracted scape in early spring, and
developing large cordate leaves later, was seen in a newly
developed City Park in Richmond, Virginia, in April, 1971.
A survey of the manuals for this area (Fernald, 1950;
Gleason & Cronquist, 1963; Harvill, 1970) showed T. far-
fara absent from Virginia. However, a survey of other
literature and herbaria proved that the Richmond speci-
men was not a new state record.
This paper is based upon collections at herbaria of the
following institutions: Virginia Polytechnic Institute and
State University, Blacksburg, Lynchburg College, Lynch-
burg, College of William and Mary, Williamsburg, Virginia
Commonwealth University, Richmond, and the United
States National Museum, Smithsonian Institution, Washing-
ton, D.C. I wish to acknowledge aid of the directors and
(or) curators at these herbaria.
The first known collections of Tussilago in Virginia in-
clude Freer 2627 (Lynchburg, US), from route 39 near
McClung, Bath County, Freer 2628 (Lynchburg, US) col-
lected from the west side of Lantz Mountain, Highland
County, Freer 2686A (VPISU) from Rockbridge County and
Gwynn s.n. (VPISU, US) from Warren County; all specimens
were collected in 1962. See Fig. 1. The first known pub-
lished report of Tussilago in Virginia is that of Freer
(1968) from Rockbridge County.
A general migration of Tussilago farfara in the United
States from north to south is noted in Fernald (1.c.) and
Gleason & Cronquist (1.c.). Fernald (1.c.) lists the species
ranging from Newfoundland to its southern limits in New
Jersey, Pennsylvania and Ohio though Reed (1951) re-
cords Tussilago in Baltimore County, Maryland, as early
405
404 Rhodora [Vol. 74
as 1903 and Strausbaugh and Core (1964) report this spe-
cies in West Virginia in 1933. Gleason and Cronquist
(1.c.), publishing 13 years later than Fernald, indicate a
southern range extension to Maryland and West Virginia.
It is apparent that the southern migration of Tussilago
is continuing at present. This species entered West Vir-
ginia in 1933 in Monongahela County which borders on
southwestern Pennsylvania and by 1964 was abundant in
Kanawha County in southern West Virginia and probably
in every county of the state (Strausbaugh and Core, 1.c.).
Migration continued and Tussilago seems to have entered
Virginia from the west in the early 1960’s (Fig. 1).
The migration pattern within Virginia perpetuates the
trend southward and eastward (Fig. 1). It is noted that
the earlier collections were made in the mountains and
that the majority of the recent ones have come from the
Piedmont and Coastal Plain. The plant in Richmond was
seen in sandy, disturbed soil immediately adjacent to the
James River. The river, the headwaters of which rise in
western Virginia, may well have transported the diseminule
from which the specimen grew. Thus, the range of Tus-
silago farfara in the United States should be extended to
include Virginia and this interesting species should become
part of the treatment of the Flora of Virginia now in prep-
aration.
Figure 1. Distribution of Tussilago farfara in Virginia. Dates
indicate the earliest known collection from each county.
1972] Tussilago — Johnson 405
LITERATURE CITED
FERNALD, MERRITT LYNDON. 1950. Gray’s Manual of Botany eighth
edition. American Book Company. New York. 1632 p.
FREER, R. S. 1968. Plants of the Central Blue Ridge: Supplement
II. Castanea 33: 163-193.
GLEASON, H. A., AND ARTHUR CRONQUIST. 1963. Manual of Vascular
Plants of Northeastern United States and Adjacent Canada.
D. Van Nostrand Company, Inc. Princeton. 810 p.
HARVILL, A. M. 1970. Spring Flora of Virginia. McClain Printing
Company. Parsons, West Virginia. 240 p.
REED, C. F. 1951. An Early Record of Tussilago farfara in Mary-
land. Castanea 16: 11.
DEPARTMENT OF BIOLOGY
VIRGINIA COMMONWEALTH UNIVERSITY
RICHMOND, VIRGINIA 23220
INVESTIGATIONS OF NEW ENGLAND
MARINE ALGAE V.
THE ALGAL VEGETATION OF THE
HAMPTON-SEABROOK ESTUARY AND THE
OPEN COAST NEAR HAMPTON, NEW HAMPSHIRE!
ARTHUR C. MATHIESON AND RICHARD A. FRALICK
Most botanical studies of New England salt marshes have
been restricted to Massachusetts (e.g. Chapman, 1940; Drum
and Webber, 1965; Webber, 1967, 1968; Webber and Wilce,
1971) and Connecticut (e.g. Miller and Egler, 1950) and
little is known of these valuable wetlands in New Hampshire
salt marches (i.e. Davis, 1956; Vagenas, 1969). Davis sum-
marized the general ecology of the Crommet Creek Estuary,
which is a small tributary of Great Bay, New Hampshire.
Vagenas studied the seasonal occurrence, composition and
zonation of algae and flowering plants at five locations in
the Hampton-Seabrook salt marsh. For the past seven years
we have conducted a variety of floristic and ecological stud-
ies of the marine algae on the open coast of New Hampshire
and the Great Bay Estuary System (Mathieson, Hehre and
Reynolds, in press; Mathieson, Reynolds and Hehre, in
press; Hehre and Mathieson, 1970). The present investi-
gation was initiated, at the request of Normandeau Associ-
ates Incorporated, in order to provide a broad “baseline” of
information regarding the species composition, distribution
and ecology of the benthonic marine algae of the Hampton-
Seabrook Estuary and the adjacent open coast of New
Hampshire (Fig. 1).
Collections and observations of marine algae were made
at fifty-four stations in the Hampton-Seabrook Estuary and
the adjacent open coast of New Hampshire (Fig. 1 and
Table I) during the summer and fall of 1969. Prior to 1969
seasonal collections had been made on the open coast near
‘Published with the approval of the Director of the New Hampshire
Agricultural Experiment Station as scientific contribution number
534.
406
1972] Marine Algae — Mathieson and Fralick 407
Bound Rock, and limited collections were made in the es-
tuary (during the fall and winter of 1966). The majority
of the stations were established by Normandeau Associates
in conjunction with their ecological survey of the estuary.
The estuarine sites were identified by a pair of numbered
stakes on opposite sides of the channel, and most of the col-
lections were restricted to the side of the channel numbered
on the map (Fig. 1) unless conspicuous differences were
noted on the two sides. The locations were primarily shore
sites, but a few offshore outcrops of rock were also studied
(Fig. 1). An aluminum boat and outboard motor were
used at most stations.
Representative samples of all conspicuous algal species
were obtained at each station. Collections were made in the
littoral (on foot) and sublittoral zones (by SCUBA). Ex-
tensive collections were made in the sublittoral zones on the
open coast, but because of poor visibility, tidal currents and
limited substrate for seaweed attachment, fewer sublittoral
samples were available in the Estuary. A complete set of
nearly 700 herbarium voucher specimens has been deposit-
ed in the Algal Herbarium of the University of New Hamp-
shire (NHA). The nomenclature of the Second Revised
British Checklist (Parke and Dixon, 1968) has been applied
in most cases. The vertical distribution of the major algal
components described in this article is given according to
the biological classification of Lewis (1964).
A record of surface water salinity and temperature was
made at most stations with a thermometer and a set of hy-
drometers. All of the salinity values were corrected to 15 C.
Diurnal variations of temperature, salinity and current
speed were recorded at one location (station C-9) during a
twelve hour period. The current speed was recorded with
a Little Captain boat speedometer (Swift Instrument Com-
pany of Boston, Massachusetts), which was modified with
a six-foot well tube.
Table II summarizes the surface water temperature and
salinity values recorded (1966-1969). The hydrographic
conditions on the exposed open coast (Bound Rock) are
408
Rhodora
[Vol. 74
HAMPTON— SEABROOK AREA, NEW HAMPSHIRE
42°56 |
.
YS a
A17 A9 T TN
AB D
Gad k
S E
m .
- .
eS .
A3 b
Al
»
f^ Pa PR, WI
fy we
Ep uA HAMPTON BEACH
e x 2
Bil
B2 We B7 B6 Z BIS a
e oe * e
FS) BIO B9 A
AS BS 0
°B4 cS H4 e s -
B8 *n | H3 !
ASS 82 HBIa*
[LÀ 4 x
*N e// se HAMPTON HARBOR 71
ES HB2*,*
ES Hi e 2" "HB3
^ FAJ . HB
- B3 SS .
BOUND x
ROCK a"
D2 Di = Rocks
H2
Y M 7 p^
4 S.
& cio
a
0.9 qe .
f= a SEABROOK BEACH
"3
(GJ e CI N
e 0
is s :
al E c
5
2 9» A
c3 .
le
C4 C5
y sN .
oe) ce ao
d ! Mil
N EN ile
——
c8
c9
. 42 52
70 51 oot
f 704
n 1 8
1972] Marine Algae — Mathieson and Fralick 409
more uniform than in the Estuary — particularly at the
headwaters of the tidal rivers. Gross fluctuations (both
daily and seasonally) of temperature and salinity are evi-
dent at any estuarine location. Figures 2 and 3 summarize
the results of a twelve hour study made at station C-9 on
the Blackwater River. The location has a pronounced tidal
current; thus, vertical mixing of the water column is evi-
dent and there is no stratification. Diurnal variations of
temperature and salinity varied from 13.2 to 16.0 C and
11.5 to 14.5 0/00, respectively (Fig. 2). The maximum sa-
linity and the minimum temperature were recorded at
slack water. Diurnal variations of tidal currents are shown
in Figure 3. The current varied from 0-4.8 knots.
A variety of substrates is evident at the stations. In gen-
eral there is a gradient from sandy beaches with rock out-
crops (i.e. Bound Rock which is on the exposed open coast)
to sandy-muddy locations (i.e. at the mouth of the Estuary)
to muddy and peat-like substrates (i.e. at the head of the
tidal rivers). Only limited quantities of solid substrates
(rock outcrops, boulders and breakwaters) are present in
the Estuary.
There are pronounced seasonal fluctuations of sand levels
on the open coast near Bound Rock, and the abrasive action
of sand limits both the species diversity and biomass of
algae. Even so there are many more species at Bound Rock
than within the Estuary. The conspicuous vegetation at
Bound Rock is composed of Cladophora sericea, Enteromor-
pha intestinalis, Pseudendoclonium marinum, Ulva lactuca,
Fucus distichus ssp distichus, F. vesiculosus, Petalonia fas-
cia, Ralfsia verrucosa, Scytosiphon lomentaria, Ahnfeltia
Figure 1. The Hampton-Seabrook Estuary lying between latitudes
42° 51’ 30" to 42° 55’ 55" north and longitudes 70° 49’ 30" to 70° 51’
. 80" west within the townships of Hampton, Hampton Falls and Sea-
brook, New Hampshire. Note that all of the tidal waters enter and
leave via the Hampton Harbor Inlet and that five rivers (Taylor,
Hampton Falls, Hampton, Brown and Blackwater) as well as many
smaller creeks and brooks are present within the Estuary. Dots with
numbers show the stations.
410 Rhodora [Vol. 74
DIURNAL VARIATION OF TEMPERATURE ANO SALINITY
AT STATION C79
l6 4
4 c
H 28
K
l4 4
x m ~
=
»
E K - æ E
: i
= ri
= 12 4 a ^
m 24
10 7
OCT.9, 1969 [
T T T T T T T T T T T T
7 9 n 13 15 7
TIME
Figure 2. Diurnal variation of temperature and salinity at station
C-9.
DIURNAL VARIATION OF CURRENTS
AT STATION C-9
5 4
4 4
a 931
c
m
m
m
z 2 4
A
2 14d
o
=
wu
O 4
OCT.9,1969
T T T T T T T T T T T T
7 9 1 13 15 17
TIME
Figure 3. Diurnal variation of currents at station C-9.
1972] Marine Algae — Mathieson and Fralick 411
plicata, Chondrus crispus, Corallina officinalis, Hilden-
brandia prototypus and Porphyra umbilicalis. Some species
(e.g. Polyides rotundus and Ahnfeltia plicata) are restrict-
ed to such sandy areas. Other plants which are not “sand
loving” forms are often reduced in size, and they are only
found in cracks and crevices. Several species are restricted
to tide pools, which are somewhat protected from the
fullest force of the sand and surf. Fucus vesiculosus domi-
nates the eulittoral zone. Ascophyllum nodosum is con-
spicuously absent. Several common sublittoral species such
as Alaria esculenta, Laminaria saccharina, L. digitata and
Desmarestia viridis are rare at Bound Rock. However, they
are present in greater abundance in the offshore rocks near
Hampton Beach (station HB1-HB4 in Fig. 1).
A comparison of the common plants at Bound Rock with
those of a “typical” estuarine site shows several differences.
A variety of fucoid (Ascophyllum nodosum, A. nodosum f.
scorpioides and Fucus vesiculosus var. spiralis) green
(Cladophora sericea, Enteromorpha erecta, E. intestinalis,
Monostroma oxyspermum, Percursaria percursa, Rhizo-
clonium riparium and Ulva lactuca), blue green (Lyngbya
spp, Oscillatoria spp and Gomphosphaeria spp.) and yellow
green algae (Vaucheria sp.) dominate the littoral zone. In
addition Spartina alterniflora, S. patens and Distichlis
spicata are abundant in the upper portion of the shore.
Results
A total of 118 taxa of benthonic marine algae was col-
lected from the fifty-four stations, including 30 Chlorophy-
ceae, 35 Phaeophyceae and 53 Rhodophyceae. A detailed
examination of the Cyanophyceae, Bacillariophyceae and
Xanthophyceae was beyond the scope of the present investi-
gation. Table III summarizes the number of seaweed taxa
found at each of the stations. The largest number of taxa
was found at Bound Rock with fewer towards the head of
the Estuary. Gross fluctuations of species numbers were
recorded at different locations within the Estuary.
The following catalogue of Species includes some synony-
412 Rhodora [Vol. 74
my, notes on collection sites and habitats and noteworthy
taxonomic features. The occurence of each species is de-
signated numerically as follows according to its habitat
(Hehre and Mathieson, 1970; Mathieson, Hehre and Rey-
nolds, in press; Mathieson, Reynolds and Hehre, in press) :
#1 Coastal — restricted to the open coast
#2 Estuarine — restricted to the estuarine environment
#3 Cosmopolitan — present in both estuarine and open
coastal environments.
Several of the cosmopolitan species may not occur in both
habitats in the Hampton-Seabrook area, but they have been
recorded from either the Great Bay Estuary or the open
coast adjacent to Bound Rock.
List of Species
Chlorophyceae :
Blidingia marginata (J. Agardh) P. Dangeard:
equals Enteromorpha marginata J. Agardh in Taylor,
1957)
Found once on rocks in the lower eulittoral zone at Bound
Rock. #3
Blidingia minima (Nàgeli ex Kützing" Kylin:
(equals Enteromorpha minima Nageli in Taylor, 1957)
Found once in the estuary (B9) on a muddy bank in the
upper littorcl zone. Abundant on the exposed open coast
(Bound Rock) where it forms fringe zones. #3
Bryopsis plumosa (Hudson) C. Agardh
Locally abundant in the estuary (A3, A5, A9, A15, A17, B1,
B2, B4 to B6, B8, and H3) on muddy surfaces in the sub-
littoral and lower eulittoral zones. Not found on the open
coast. #2
Chaetomorpha aerea (Dillwyn) Kützing:
Collected three times from the lower eulittoral (in tide
pools) at Bound Rock. #1
Chaetomorpha linum (O. F. Muller) Kützing:
(including Chaetomorpha atrovirens Taylor, in Taylor,
1957)
1972] Marine Algae — Mathieson and Fralick 413
According to our interpretation C. atrovirens and C. linum
are not distinct for there is a continuous gradient of size
and color between the two. Chaetomorpha linum is the old-
er name and it should be retained. The plant is common as
an entangled mass amongst various algae in the lower eulit-
toral and sublittoral zones of the estuary (A2, All, A18,
B1, B2, B4 to B6, B9, B10, B12, C1, C2, C6, C7, and C9) and
the exposed open coast (Bound Rock, HB1 and HB4). £3
Chaetomorpha melagonium (Weber et Mohr) Kützing:
Occasional on rocks in the lower eulittoral and sublittoral
zones of the exposed open coast (Bound Rock, HB1 and
HB4). £3
Cladophora sericea (Hudson) Kützing sensu van den Hoek,
1963:
Locally abundant in high tide pools in the estuary (A6, AT,
A9, A10, B3, B7, B10, B12, C3 to C5, and H3) and on the
exposed open coast (Bound Rock). £3
Codiolum pusillum (Lyngbye) Kjellman in Foslie:
Locally abundant on rocks in the upper littoral zone of the
exposed open coast (Bound Rock). Often mixed with
Bangia f uscopurpurea and various blue green algae. It may
be the sporophyte stage of one or more local species of
Urospora (Scagel, 1966). £1
Enteromoprpha compressa, (L.) Greville:
Found once on rocks in the lower eulittoral zone at Bound
Rock. £3
Enteromorpha erecta (Lyngbye) J. Agardh:
Abundant on muddy surfaces of the eulittoral zone through-
out the estuary (Al to A9, A13, A14, A16, A17, B1 to B7,
B9, B13, C1 to C6, H1 and H3) ; occasionally present as an
epiphyte on fucoid algae and Spartina alterniflora. Uncom-
mon on the exposed open coast (Bound Rock). #3
Enteromorpha intestinalis (L.) Link:
Common on rocks and muddy surfaces of the eulittoral zone
throughout the estuary (Al, A2, A4, A6 to A10, A13, A17,
B1 to B5, B8, C1, C3, C5 to C7, C9, and H3) and on the
open coast (Bound Rock). #3
414 Rhodora [Vol. 74
Enteromorpha linza (L.) J. Agardh:
Found sporadically in the estuary (C9 and H1) and the ex-
posed open coast (Bound Rock and HB1) ; present on mud,
rocks and as an epiphyte in the lower eulittoral and upper
sublittoral zones. #3
Enteromorpha plumosa Kitzing:
Found twice in the estuary (A18 and B12) on muddy sur-
faces in the lower eulittoral zone. #3
Enteromorpha prolifera (O. F. Müller) J. Agardh:
Infrequent in the estuary (B2, C2, C3, C5, C6 and C11);
present on muddy surfaces of the eulittoral zone. Mixed
with Enteromorpha erecta but never occurring as abun-
dantly. #3
Monostroma fuscum (Postels et Ruprecht) Wittrock:
Found sporadically throughout the estuary (A15, B2, B4,
B6, B8, C1 and C3) on mud and rocks in the lower eulittoral
and sublittoral zones. Abundant in localized areas on the
exposed open coast (Bound Rock and HB4), and with the
same vertical distribution. #3
Monostroma grevillei (Thuret) Wittrock :
Infrequent on rocks in the mid and lower eulittoral zones of
the estuary (A5, A15, B6 and B10). Abundant on the ex-
posed open coast (Bound Rock) and with the same vertical
distribution. #3
Monostroma leptodermum Kjellman:
Found once adrift at A3. #3
*Monostroma oxyspermum (Kiitzing) Doty:
Locally abundant throughout the estuary (A2 to A4, A9,
B1, B4, B6, B8, Cl, C2, and C4 to C7) — particularly in
areas of low salinity. It forms a distinct band on high ver-
tical (muddy) banks and occasionally occurs as an epiphyte
on Spartina alterniflora and other vascular plants. #2
Monostroma pulchrum Farlow:
Common (during the summer) on rocks and on various al-
gae in the lower eulittoral and sublittoral zones of the ex-
posed open coast (Bound Rock). #3
*Within range but not previously recorded for New Hampshire.
1972] Marine Algae — Mathieson and Fralick 415
Percursaria percursa (C. Agardh) Rosenvinge:
Locally abundant in the estuary (Al, A2, A16, B1, B3, B6,
B7, and C3 to C5) as free-floating masses in tide pools and
attached to muddy surfaces in the upper eulittoral zone.
Often mixed with Rhizoclonium riparium and Cladophora
sericea. #3
Pseudendoclonium marinum (Reinke) Aleem et Schulz:
(equals Protoderma marinum Reinke in Taylor, 1957)
Abundant on rocks from the mid eulittoral to the sublittoral
zones on the exposed open coast (Bound Rock) ; found once
in the estuary (All). £3
Rhizoclonium riparium (Roth) Harvey:
Abundant throughout the estuary (A1 to A5, A7 to A9,
A15, B1 to B4, B6, B8 to B10, B11 to B13, C1 to C6, C8,
C10, C11, and H3) as free floating masses in tide pools and
attached to muddy surfaces in the upper littoral zone. Often
mixed with Cladophora sericea and Percursaria percursa.
#3
Rhizoclonium tortuosum Kützing :
Abundant during the summer on the exposed open coast
(Bound Rock) ; found once in the estuary (C9). Present as
entangled masses amongst various algae (particularly
Chondrus erispus and Gigartina stellata) in the lower eu-
littoral and sublittoral zones. £3
Spongomorpha arcta (Dillwyn) Kützing:
Abundant on rocks (rarely as an epiphyte) in the mid-
lower eulittoral zones of the exposed open coast (Bound
Rock) ; most conspicuous in the late winter and spring.
#3
Spongomorpha spinescens Kiitzing:
Abundant on rocks in the mid to lower eulittoral zones of
the exposed open coast (Bound Rock) ; most conspicuous in
the spring and summer. #3
Ulothrix flacca (Dillwyn) Thuret in Le Jolis:
Locally abundant (during winter and spring) on rocks in
the upper littoral zone of the exposed open coast (Bound
Rock). #3
416 Rhodora [Vol. 74
Ulva lactuca L:
Ubiquitous throughout the estuary (being found at all sta-
tions except A6, A11, A15, A16, C8, D2, D3, H2, and H4)
on mud and any solid substrates in the lower eulittoral and
sublittoral zones. Present on the exposed open coast (Bound
Rock and HB2 and HB4) but not as abundant as in the
estuary. £3
Urospora collabens (C. Agardh) Holmes et Batters:
Locally abundant (particularly during the winter and
spring) on rocks in the upper littoral zone at Bound Rock.
#3
Urospora penicilliformis (Roth) Areschoug:
The abundance and distribution (both seasonal and verti-
cal) of this species is essentially similar to that of U.
collabens — except that it tends to appear later than U.
collabens. #3
Urospora speciosa (Carmichael ex Harvey in Hooker)
Leblond et Hamel:
Occasional (particularly during the winter and spring) on
rocks in the upper littoral zone at Bound Rock. £1
Phaeophyceae:
Agarum cribrosum (Mertens) Bory:
Common on rocks in the sublittoral zone of the exposed
open coast (Bound Rock, HB1, HB3, and HB4) ; not found
in the estuary. #1
Alaria esculenta (L.) Greville:
Locally abundant on rocks in the sublittoral zone of the
exposed open coast (Bound Rock, HB2 and HB3); not
found in the estuary. #1
Ascophyllum nodosum (L.) Le Jolis:
Common throughout the estuary (A2, A4, A11, A13, A14,
A17, A18, B1, B2, B9, B10, C1 to C4, C10 and C11 and
H3) on any solid substrate from the upper sublittoral to
the mid eulittoral zones. It is a rare plant on the exposed
open coast at Bound Rock. #3
Ascophyllum nodosum (L.) Le Jolis f. scorpioides (Horne-
mann) Reinke:
1972] Marine Algae — Mathieson and Fralick 417
Common on high muddy banks throughout the estuary
(Al to A3, A5 to AT, A12 to A14, A17, B1 to B3, B5, B6,
B8, B9, B13, C1 to C3, C5, C6, C11, D2 and D3) ; entangled
amongst Spartina alterniflora and other vascular plants.
#2
Chorda filum (L) Stackhouse:
Found once on pier pilings in the estuary (H3); another
time on rocks and shells on the exposed open coast (Bound
Rock). In both cases the plants were present in the sub-
littoral zone. #3
Chorda tomentosa Lyngbye:
Locally abundant (during the summer) on scattered rocks
in the sublittoral zone of the exposed open coast (Bound
Rock and HB3). #3
Chordaria flagelliformis (O. F. Müller) C. Agardh:
Found twice in the estuary (H1 and H3) ; in both cases it
was growing on pier pilings in the lower eulittoral — sub-
littoral zones. The plant is common (during the summer)
on the exposed open coast (Bound Rock and HB2) and has
the same vertical distribution as in the estuary. £3
Desmarestia aculeata (L.) Lamouroux:
Common on rocks in the sublittoral zone of the exposed
open coast (Bound Rock, HB2 to HB4). #1
Desmarestia viridis (O. F. Müller) Lamouroux:
Occasional on rocks in the sublittoral zone of the exposed
open coast (Bound Rock, HB1, HB3 and HB4). #1
Dictyosiphon foeniculaceus (Hudson) Greville:
Found once on mud covered rocks in the lower eulittoral
zone of the estuary (C5). #3
Ectocarpus confervoides (Roth) Le Jolis:
(equals E. siliculosus (Dillwyn) Lyngbye in Parke and
Dixon, 1968)
Occasional on rocks and larger algae in the eulittoral and
sublittoral zones of the estuary (A14, A17, B9, C9 to C11
and H1) and the exposed open coast (Bound Rock, HB1 to
HB4). The estuarine plants were more robust in stature
than the plants from the open coast. #3
418 Rhodora [Vol. 74
Ectocarpus siliculosus (Dillwyn) Lyngbye:
Found once as an epiphyte on Spartina alterniflora in the
upper littoral zone of the estuary (C5). #3
Elachista fucicola (Velley) Areschoug :
An occasional epiphyte on Ascophyllum nodosum and Fucus
vesiculosus var spiralis in the estuary (Al, B12, C1, C3,
C5 and H3); very common on Fucus vesiculosus on the
exposed open coast (Bound Rock and HB2). #3
Fucus distichus (L.) emend Powell subsp. distichus Powell:
Locally abundant in high tide pools of the exposed open
coast (Bound Rock). £1
Fucus distichus (L.) emend Powell subsp. edentatus (De
la Pylaie) Powell:
Common on rocks in the lower eulittoral — sublittoral zones
of the exposed open coast (Bound Rock and HB2). Found
once in a similar habitat near the mouth of the estuary
(H1). £3
Fucus distichus (L.) emend. Powell subsp. evanescens (C.
Agardh) Powell:
Locally abundant on rocks in the lower eulittoral — sub-
littoral zones of the exposed open coast (Bound Rock).
#3
Fucus vesiculosus L:
Abundant on semi-exposed rocks from the mid to the lower
eulittoral at Bound Rock. #3
Fucus vesiculosus L. var spiralis Farlow:
Ubiquitous throughout the estuary (at all stations except
A12, A15, A16, C8, H1, H2 and H4) on mud, rocks, shells
and any other solid substrates in the mid to the upper eulit-
toral zone. It is one of the most conspicuous species on the
upper banks of the salt marshes, where it is associated with
Ascophyllum nodosum f. scorpioides, Spartina alterniflora
and various other vascular plants. #2
Giffordia granulosa (J. E. Smith) Hamel:
Found once in the estuary (A7) on mud-covered rocks in
the lower eulittoral zone. #2
1972] Marine Algae — Mathieson and Fralick 419
Laminaria digitata (Hudson) Lamouroux:
Present on rocky substrate in the sublittoral zone of the
exposed open coast (Bound Rock, HB1, HB3 and HB4)
and at the mouth of the estuary (H1). Each specimen had
a consistent +-— anatomy (i.e. mucilage ducts are present
in the blade and absent from the stipe, Wilce, 1965). #3
Laminaria saccharina (L.) Lamouroux sensu Wilce, 1965:
Its distribution was essentially similar to that of L. digitata
except that it was found at one other coastal (HB2) and
estuarine location (A18). All of the specimens were the
—— ecotype of Wilce, 1965 (ie. L. agardhii Kjellman in
Taylor, 1957). £3
Leathesia difformis (L.) Areschoug:
A common epiphyte (during the summer) on Chondrus
crispus and other algae in the lower eulittoral-upper sub-
littoral zones of the exposed open coast (Bound Rock and
HB2). £1
Petalonia fascia (O. F. Müller) Kuntze:
Common throughout the estuary (A4, A5, A7, A9, A14,
A15, A17, B2, B4, B6, B10, B13, C2, C6, C11, D2 and D3)
and on the exposed open coast (Bound Rock). It is present
in the eulittoral zone on rocks (often in tide pools), mud,
and occasionally as an epiphyte on large algae. #3
Pilaiella littoralis (L.) Kjellman:
Common on rocks and as an epiphyte on Fucus vesiculosus
in the mid-lower eulittoral zones of the exposed open coast
(Bound Rock). Occasionally present in the estuary (Cl,
C5 and C6) and with the same vertical distribution. #3
*Pseudolithoderma extensum (Crouan frat.) S. Lund:
(equals Lithoderma extensum (Crouan) Hamel in Tay-
lor, 1957)
Occasional on sublittoral stones and boulders of the ex-
posed open coast (HB2 and HB4). #1
Ralfsia borneti Kuckuck:
Occasional on stones and boulders in the lower eulittoral
— sublittoral zones of the estuary (A8, A10, B10, B11, and
C1). According to Edelstein, Chen and McLachlan (1970)
420 Rhodora [Vol. 74
R. borneti is a stage in the life history of Petalonia fascia,
and it is not a valid taxa. #3
Ralfsia clavata (Harvey in Hooker) Crouan frat.:
Found once in the estuary (A-8) on mud covered rocks in
the lower eulittoral zone. It is also described (Edelstein,
Chen and McLachlan, 1970) as a stage in the life history
of Petalonia fascia. #3
Ralfsia fungiformis (Gunner) Setchell et Gardner:
Found once on rocks in the sublittoral zone of the exposed
open coast (HB4). #1
Ralfsia verrucosa (Areschoug) J. Agardh:
Found twice on rocks and shells in the upper sublittoral
and eulittoral zones of the estuary (B3 and C1). Abundant
in the same zones on the exposed open coast (Bound Rock).
#3
Saccorhiza dermatodea (De la Pylaie) J. Agardh:
Occasional on rocks in the sublittoral zone of the exposed
open coast (HB1 and HB3). #1
Scytosiphon lomentaria (Lyngbye) Link:
Present on rocks (often in tide pools), mussels shells, mud
and occasionally epiphytic on various plants in the eulittoral
zone of the estuary (A5, A14, A15, A17, B1, B2, B4, B10,
C9, C11 and D3) and the exposed open coast (Bound Rock).
#3
**Sorapion kjellmanii (Wille) Rosenvinge:
Found once on rocks in the sublittoral zone of the exposed
open coast (HB2). £1
**Sphacelaria plumosa Lyngbye:
(equals Chaetopteris plumosa (Lyngbye) Kiitzing in
Taylor, 1957)
Occasional on sand covered rocks in the mid-lower sublit-
toral zone of the exposed open coast (HB1). #1
Sphacelaria radicans (Dillwyn) C. Agardh:
Occasional on muddy or sandy surfaces in the mid-lower
eulittoral zone of the estuary (A9, B1 and B10); also
**Range extension and a new record for New Hampshire.
1972] Marine Algae — Mathieson and Fralick 421
present on sand covered rocks in the sublittoral zone of
the exposed open coast (HB1, HB3 and HB4). £3
Spongonema tomentosum (Hudson) Kützing:
(equals Ectocarpus tomentosus (Hudson) Lyngbye in
Taylor, 1957)
Found once as an epiphyte on Laminaria saccharina in the
sublittoral zone of the exposed open coast (HB3). #1
Rhodophyceae :
Ahnfeltia plicata (Hudson) Fries:
Locally abundant on sand-covered rocks and boulders in
the lower eulittoral — sublittoral zones of the exposed open
coast (Bound Rock, HB1, HB2 and HB4). #3
Antithamnion floccosum (O. F. Müller) Kleen:
Found once on sand covered rocks in the sublittoral zone
of the exposed open coast (HB3). #3
Audouinella membranaceae (Magnus) Papenfuss:
Epiphytic on species of Sertularia which in turn may be
epiphytic (commonly on fucoid algae) or saxicolous in the
eulittoral zone of the exposed open coast (Bound Rock). #3
* Asterocytis ramosa (Thwaites in Harvey)
Gobi ex Schmitz:
Found once as an epiphyte on Cladophora sericea in a high
marshy tide pool (C4). Growing in association with Per-
cursaria percursa and various blue green algae. #2
Bangia ciliaris Carmichael:
Found once as an epiphyte on Cladophora sericea in a
high marshy tide pool (C5) in the estuary. #3
Bangia fuscopurpurea (Dillwyn) Lyngbye:
Abundant (particularly during the winter and spring) on
rocks in the upper littoral zone of the exposed open coast
(Bound Rock). #3
Callithamnion baileyi Harvey:
Found once in the estuary (H3) on a styrofoam float. #3
Callithamnion corymbosum (J. E. Smith) C. Agardh:
Found once in the estuary (A17) on rocks in the upper
sublittoral zone. #2
422 Rhodora [Vol. 74
Ceramium rubrum (Hudson) C. Agardh:
Present on rocks and epiphytic on large macroscopic algae
in the lower eulittoral and sublittoral zones. Occasional
within the estuary (B1 to B3, B7, C2, C5, and H3), but
more abundant on the exposed open coast (Bound Rock,
HB1 to HB4). #3
Ceramium strictum Harvey:
Abundant throughout the estuary (A1 to A4, AT, A8, A10,
A12, A17, B1 to B6, B9, C1, C3 to C7, C11 and H1) on
muddy surfaces in the lower eulittoral — sublittoral zones.
Uncommon on the exposed open coast (Bound Rock). #3
Chondrus crispus Stackhouse:
Common throughout the estuary (Al to A3, A5, A9 to
A15, A17, A18, B2, B4 to B6, B8 to B11, Cl, C9, C11, D3
and H1) and the exposed open coast (Bound Rock, HB2
to HB4); present on any solid substrate in the lower
eulittoral and sublittoral zones. The stature of the estuarine
plants is much larger than the plants from the open coast.
#3
Clathromorphum circumscriptum (Strömfelt) Foslie:
(equals Phymatolithon compactum (Kjellman) Foslie
in Taylor, 1957)
Abundant on rocks and shells in the lower eulittoral and
sublittoral zones of the exposed open coast (Bound Rock,
HB1 to HB4). Found once in the estuary (A10) on a sub-
tidal population of mussels. £3
Corallina officinalis L.:
Locally abundant on rocks and boulders (often in tide
pools) in the eulittoral and sublittoral zones of the exposed
open coast (Bound Rock, HB1 to HB4). £1
Cystoclonium purpureum (Hudson) Batters var. cirrhosum
Harvey:
Present on rocks and as an epiphyte on larger algae in the
sublittoral zone of the exposed open coast (Bound Rock,
HB1 to HB4). #3
Dermatolithon pustulatum (Lamouroux) Foslie:
1972] Marine Algae — Mathieson and Fralick 423
(equals Lithophyllum pustulatum (Lamouroux) Foslie in
Taylor, 1957)
An occasional epiphyte on various algae (particularly
Chondrus crispus and Gigartina stellata) in the lower
eulittoral-sublittoral zones of the exposed open coast
(Bound Rock, HB2 and HB4). #3
Dumontia incrassata (O. F. Müller) Lamouroux:
Abundant on rocks in the mid and lower eulittoral zones
of the exposed coast (Bound Rock). Occasional throughout
the estuary (A5, A14, A15, A17, A18, B6, B10, and D3)
and with the same vertical distribution as on the open
coast. #3
Euthora cristata (C. Agardh) J. Agardh:
Present on rocks and occasionally epiphytic on various
plants (e.g. Phyllophora spp.) in the sublittoral zone of the
exposed open coast (Bound Rock, HB1, HB3 and HB4).
#1
Gigartina stellata (Stackhouse) Batters:
On rocks in the lower eulittoral and sublittoral fringe zones
of the exposed open coast (Bound Rock). #3
Gloiosiphonia capillaris (Hudson) Carmichael ex Berkeley :
Found twice on rocks in the sublittoral zone of the ex-
posed open coast (HB1 and HB4). £3
Hildenbrandia prototypus Nardo:
Common on rocks in the eulittoral and sublittoral zones of
the exposed open coast (Bound Rock and HB2). Less com-
mon in the estuary (A8, A10, A11 and B10) but with the
same vertical distribution. #3
Kylinia secundata (Lyngbye) Papenfuss:
A common epiphyte on various algae in the eulittoral zone
of the exposed open coast (Bound Rock). #3
Lithophyllum corallinae (€rouan frat.) Heydrich:
Found once in the sublittoral zone of the exposed open
coast (HB4) ; a specific epiphyte on Corallina officinalis.
#1
Lithothamnium glaciale Kjellman:
Common on shells and rocks (often in tide pools) in the
424 Rhodora [Vol. 74
lower eulittoral and sublittoral zones of the exposed open
coast (Bound Rock, HB1 to HB4). #1
Melobesia lejolisti Rosanoff :
(equals Fosliella lejolisii (Rosanoff) Howe in Taylor,
1957)
An occasional epiphyte on Phyllophora spp. in the sub-
littoral zone of the exposed open coast (Bound Rock and
HB3). #3
Membranoptera alata (Hudson) Stackhouse:
A common epiphyte on various algae (occasionally on rock)
in the sublittoral zone of the exposed open coast (Bound
Rock, HB1, HB3 and HB4). #1
Nemalion helminthoides (Velley in Withering) Batters:
Uncommon on rocks in the lower eulittoral zone of the
exposed open coast (Bound Rock). #1
Petrocelis middendorfii (Ruprecht) Kjellman:
Present on rocks and shells in the sublittoral zone of the
exposed open coast (HB2 and HB4). #3
Peyssonelia rosenvingii (Schmity in Rosenvinge) :
Occasional on rocks in the sublittoral zone of the exposed
open coast (HB1 and HB4). #1
Phycodrys rubens (L.) Batters:
Common on rocks and as an epiphyte on various algae in
the lowest eulittoral and sublittoral zones of the exposed
open coast (Bound Rock, HB1, HB3 and HB4) and at the
mouth of the estuary (H1). #3
Phyllophora brodiaei (Turner) Endlich:
Common on rocks in the sublittoral zone of the exposed
open coast (Bound Rock, HB1 to HB4). #1
Phyllophora membranifolia (Goodenough et Woodward)
J. Agardh:
The vertical and horizontal distribution of P. membrani-
folia is essentially similar to that of P. brodiaei. It was
found at stations HB2, HB3 and Bound Rock. #3
*Phymatolithon laevigatum (Foslie) Foslie:
Present on rocks and shells in the sublittoral zone of the
exposed open coast (HB2 to HB4). #1
1972] Marine Algae — Mathieson and Fralick 425
Phymatolithon lenormandi (Areschoug) Adey:
Present on rocks in the sublittoral zone of the exposed open
coast (HB4). #3
Plumaria elegans (Bonnemaison) Schmitz:
Common on vertical rock faces under overhanging fucoids
in the lower eulittoral zone of the exposed open coast
(Bound Rock). It was found once in the estuary (A14)
with the same vertical distribution. #3
Polyides rotundus (Hudson) Greville:
Occasional on sand covered rocks in the sublittoral zone of
the exposed open coast (Bound Rock, HB2 to HB4). #3
Polysiphonia denudata (Dillwyn) Greville ex Harvey in
Hooker:
Found twice in the estuary (B6 and B10) on muddy sur-
faces in the sublittoral zone. #3
Polysiphonia elongata (Hudson) Sprengel:
Found twice in the sublittoral zone of the estuary (A7 and
H3). #2
Polysiphonia fibrillosa (Dillwyn) Sprengel :
Common throughout the estuary (Al, A2, A4, A10, B2 to
B6, C2, C4 to C6, C9 and H3) on muddy surfaces in the
lower eulittoral and sublittoral zones. #2
Polysiphonia lanosa (L.) Tandy:
Hemiparasitic on Ascophyllum nodosum on the open coast
(Bound Rock) and at the mouth of the estuary (A2 and
A4). £3
Polysiphonia nigra (Hudson) Batters:
Present on rocks in the lower eulitoral-sublittoral zones of
the estuary (A15, B2, B3, B6, B7 and H3) and the exposed
open coast (Bound Rock). #3
Polysiphonia nigrescens (Hudson) Greville:
On rocks and shells in the lower eulittoral and sublittoral
zones of the estuary (A4, AT, All, A12, A15, B2, BA, B5,
B10, C1, C2, C5, H1 and H3) and the exposed open coast
(HB1 and HB4). £3
426 Rhodora [Vol. 74
Polysiphonia novae-angliae Taylor:
Present on rocks in the sublittoral zone of the exposed
open coast (Bound Rock, HB1 and HB2). #3
Polysiphonia subtilissima Montagne:
Found once in the estuary (A12) ; growing on mud in the
lower eulittoral zone. #1
Polysiphonia urceolata (Lightfoot ex Dillwyn) Greville:
Common on rocks in the lower eulittoral and sublittoral
zones of the exposed open coast (Bound Rock, HB1 to HB4).
Found once in the estuary at (B6) with the same vertical
distribution. #3
Porphyra miniata (C. Agardh) C. Agardh:
Common (particularly during the summer) on rocks and
epiphytic on various plants in the upper sublittoral zone
of the exposed open coast (HB1 to HB4). #3
Porphyra umbilicalis (L.) J. Agardh:
Common on rocks, mud and on various algae in the eulit-
toral zone of the estuary (Al, A3 to A5, A8, A12, A14,
A17, C3, C5, C6, C9, H1 and H3) and the exposed open
coast (Bound Rock). #3 The forma epiphytica Collins
was only found at Bound Rock. #3
Ptilota serrata Kiitzing:
Present on rocks and epiphytic on various algae in the
sublittoral zone of the exposed open coast (Bound Rock,
HB1 and HB3). #1
Rhodochorton purpureum (Lightfoot) Rosenvinge:
Present on vertical rock faces under overhanging fucoids
in the midlower eulittoral zone of the exposed open coast
(Bound Rock). #3
*Rhodophysema elegans (Crouan frat. ex J. Agardh)
Dixon: (equals Rhododermis elegans Crouan in Taylor,
1957)
Occasional on rocks and shells in the sublittoral zone of
the exposed open coast (HB2 and HB4). #1
Rhodomela confervoides (Hudson) Silva:
Locally abundant on sand-covered rocks in the lower eulit-
1972] Marine Algae — Mathieson and Fralick 427
toral and sublittoral zones of the exposed open coast (Bound
Rock, HB1 and HB3). £1
Rhodophyllis dichotoma (Lepeschkin) Gobi:
Found once adrift at Bound Rock. #1
Rhodymenia palmata (L.) Greville:
Relatively common on rocks and various algae (particularly
Laminaria spp.) in the sublittoral zone of the exposed open
coast (Bound Rock, HB1 to HB4) and the mouth of the
estuary (H1). #3
DISCUSSION
Eight of the taxa of marine algae recorded in the present
study represent new records for the state of New Hamp-
shire (Monostroma oxyspermum, Asterocytis ramosa,
Phymatolithon laevigatum, Polysiphonia fibrillosa, Rhodo-
physema elegans, Pseudolithoderma extensum, Sorapion
kjellmanii and Sphacelaria plumosa). Sorapion kjellmanii
and Sphacelaria plumosa also represent extensions of
known distributional range on the northeast coast of
North America, for they were both previously recorded
from Ellesmere Island to Nova Scotia (as summarized
in Taylor, 1957 and Cardinal, 1968). It should be noted
that both species were collected in the mid-lower sublittoral
zones of the open coast.
The primary factors restricting the growth and distri-
bution of seaweeds in the Hampton-Seabrook Estuary are
a lack of stable substrate and adverse hydrographic condi-
tions. As a consequence gross fluctuations of species num-
bers occurred at different locations, and a gradual reduction
in species diversity and biomass was evident from the open
coast to the head of the estuary. Green algae showed the
widest distribution throughout the estuary, for they grew
on a variety of substrates and in a wide range of tempera-
tures and salinities.
A greater diversity and biomass of seaweeds were re-
corded in the Great Bay Estuary complex (Mathieson,
Reynolds and Hehre, in press) than in the Hampton-
428 Rhodora [Vol. 74
Seabrook Estuary. Thus, 98 taxa of algae were found in
the former area, while only 57 were found in the latter.
The variety of species found in the estuarine environments
of Great Bay is primarily due to its large size (over 15,000
acres of tidewater) and its greater variety of habitats
(particularly tidal rapids) and substrates. Peak popula-
tions of seaweeds were found at Dover Point and Adams
Point in the Great Bay System (Mathieson, Reynolds and
Hehre, in press). Both of the sites are exposed to strong
tidal currents and their floras are “open coastal" in nature.
More than half of the 76 cosmopolitan species found in
the Great Bay area were either restricted to the two tidal
rapid sites or they did not extend inland beyond Dover
Point or Adams Point. Only one major tidal rapid was
observed in the Hampton-Seabrook area (station C-9),
and it was so shaded that few algae grew there. The dis-
crepancy in species numbers between the two estuaries is
primarily accounted for by the cosmopolitan component of
Great Bay.
All of the species found in the Hampton-Seabrook
Estuary, except Monostroma oxyspermum, Chorda filum,
Dictyosiphon foeniculaceus, Elachista fucicola, Ralfsia
borneti, R. clavata, Sphacelaria radicans, Asterocytis
ramosa, Plumaria elegans and Polysiphonia fibrillosa,
were also found in the Great Bay area. The occurrence
of P. elegans and S. radicans in estuarine waters was
particularly striking, for both of the species are most
abundant on the open coast.
ACKNOWLEDGEMENTS
The author wishes to thank the following individuals for
their help: Mr. John Shipman, Dr. Norman Reynolds and
Dr. Edward Hehre for their help during the collection and
processing of specimens; Dr. Donald Normandeau, of Nor-
mandeau Associates, who provided a boat, outboard motor
and other support to complete the investigation.
1972]
A-12
A-13
A-14
A-15
Marine Algae — Mathieson and Fralick 429
Table I
Stations
At the mouth of the first major tributary SE of Tide Mill
Creek on the Hampton River, Hampton Falls, Hampton,
N. H.* (1.3 miles inland)
The northeast portion of an “island” formed at the mouth of
the Taylor River, Blind Creek and the upper part of the
Hampton River, Hampton Falls, Hampton, N. H. (1.91 miles
inland)
Approximately 1500 feet NE of the mouth of Tide Mill Creek
and the Hampton River, Hampton, N. H. (1.3 miles inland)
Approximately 1800 feet NW of Station A-2 on the Hampton
River, Hampton-Hampton Falls, N. H. (2.3 miles inland)
Opposite a small brook, which empties into the first tributary
above the mouth of the Hampton Falls River, Hampton, N. H.
(3.3 miles inland)
At the junction of the first oxbow NE of station A-5 on the
Taylor River, Hampton, N. H. (3.2 miles inland)
Hampton Landing on the Taylor River, Hampton, N. H. (3.1
miles inland)
At the mouth of Nudds Canal and Blind Creek, Hampton,
N. H. (2.31 miles inland)
Tide Mill Creek, by the route 101 bridge, Hampton, N. H.
(2.56 miles inland)
Hampton Falls River south of Depot Avenue and near the
Boston and Maine Railroad bridge, Hampton Falls, N. H.
(5.2 miles inland)
Approximately 2000 feet SW of the Boston and Maine “sub-
station” which is between Lafayette and Landing Roads.
Station A-11 is adjacent to the B & M railroad tracks, and
on the Taylor River, Hampton, N. H. (4.4 miles inland)
End of Depot Avenue on the Hampton Falls River, Hampton
Falls, N. H. (5.0 miles inland)
Middle of the southernmost oxbow near the mouth of the
Taylor River and the Hampton Town Line, Hampton, N. H.
(3.0 miles inland)
A bend in the first tributary above (north) the Hampton Falls
River where the river crosses the railroad tracks, Hampton,
N. H. (3.7 miles inland)
Same as A-14, except the land side of the railroad tracks,
rather than the harbor side as A-13, Hampton, N. H. (3.7
miles inland)
* Distance inland from exposed open coast
Rhodora [Vol. 74
A salt marsh on a point of land made by the Hampton River
and the first tributary above the Hampton Falls River,
Hampton, N. H. (3.5 miles inland)
Hampton Landing, Taylor River, Hampton, N. H. (3.01 miles
inland)
The Willows; at the mouth of Tide Mill Creek and the Hamp-
ton River, Hampton, N. H. (1.6 miles inland)
On a small “island” opposite Eastman’s Slough and about
2500 feet west of Locke Point State Park area, Hampton
Falls, Seabrook, N. H. (1.0 miles inland)
The southernmost portion of Eastman Slough, near Halftide
Rock and at the mouth of the Brown River, Hampton Falls-
Seabrook, N. H. (1.7 miles inland)
Just inside the mouth of Hunts Island Creek and at the
junction of the Brown River, Seabrook, N. H. (1.8 miles
inland)
Approximately 500 feet NW of the first tributary past Hunts
Island Creek, Hampton Falls-Seabrook, N. H. (2.0 miles in-
land)
Approximately 700 feet NE of the mouth of Swain's Creek,
Hampton Falls, N. H. (2.2 miles inland)
Robbins Point, Hampton Flats, Hampton Falls-Seabrook,
N. H. (2.3 miles inland)
Browns River, first tributary upstream from Swain's Creek,
Hampton Falls, N. H. (2.5 miles inland)
Approximately 800 feet upstream from Robbins Point, Hamp-
ton Falls, N. H. (2.4 miles inland)
Approximately 700 feet upstream from station B-8, Hampton
Falls-Seabrook, N. H. (2.53 miles inland)
End of Rocks Road, on the Browns River, Hampton Falls-
Seabrook, N. H. (2.8 miles inland)
Near the mouth of the first major tributary east of the head
waters of the Browns River, Hampton Falls-Seabrook, N. H.
(3.0 miles inland)
Approximately 1500 feet upstream (west) from station B-11,
just before a major oxbow, Hampton Falls-Seabrook, N. H.
(3.3 miles inland)
Swain’s Creek, neck of the first oxbow, Hampton Flats,
Hampton Falls, N. H. (2.6 miles inland)
At the mouth of the Blackwater River; near the first tributary
SW of Mills Point, Seabrook, N. H. (1.5 miles inland)
Approximately 1200 feet SW of the first tributary past River-
side, Seabrook, N. H. (2.2 miles inland)
Approximately 1200 feet south of C-2, Seabrook, N. H. (2.3
miles inland)
1972] Marine Algae — Mathieson and Fralick 431
C- 4 Approximately 500 feet SE of C-3, Seabrook, N. H. (2.4 miles
inland)
C- 5 Approximately 800 feet SE of C-4, Seabrook, N. H. (2.5 miles
inland)
C- 6 Approximately 800 feet SE of C-5 and near the first tributary
southeast of C-2, Seabrook, N. H. (2.7 miles inland)
C- 7 Approximately 1200 feet SE of C-6 and near a large white
rock, Seabrook, N. H. (3.0 miles inland)
C-8 Approximately 1500 feet SE of C-7, Seabrook, N. H. (3.3
miles inland)
C-9 By the route 286 bridge which crosses the Blackwater River,
Seabrook, N. H. (3.6 miles inland)
C-10 Mills Point — mouth of the Blackwater River, Seabrook, N. H.
(1.3 miles inland)
C-11 Riverside, Seabrook, N. H. (1.7 miles inland)
D-1 At the mouth of the creek between Knowles Island and main-
land, Seabrook, N. H. (1.34 miles inland)
D- 2 Before the first bend near the mouth of Mill Creek, Seabrook,
N. H. (1.6 miles inland) l
D- 3 Walton Landing, end of Walton Road, Seabrook, N. H. (2.1
miles inland)
H- 1 At the junction of the middle piling of the toll bridge in
Hampton Harbor and the tower at Hampton, N. H. (0.4 miles
inland)
H- 2 In the channel of Hampton Harbor near the mouth of the
Blackwater River and at the junction of the imaginary line
between the Seabrook Marina and Knowles Island, Seabrook,
N. H. (1.0 miles inland)
H- 3 Smith and Gilmore Marina, Hampton, N. H. (0.8 miles inland)
H- 4 Hampton Marina — mouth of Hampton River, Hampton, N. H.
(0.97 miles inland)
HB-Bound
Rock The area in the immediate vicinity of Beckman's Point, which
is near the mouth of Hampton Harbor, Hampton, N. H. Ex-
posed open coast.
HB-1 A series of offshore rocks in the immediate proximity of the
red marker flag, and in 35-40 feet of water, Hampton, N. H.
Exposed open coast
HB-2 The southernmost rock outcrop (a rookery) before the Hamp-
ton Breakwater; in 15-20 feet of water and the sheltered side,
Hampton, N. H. Exposed open coast.
HB-3 The exposed (seaward) side of the same rock outcrop as
HB-2; 30-40 feet of water, Hampton, N. H. Exposed open
coast.
HB-4 The southernmost outcrop which is opposite the Hampton
State Beach Park Breakwater; in 20-25 feet of water, Hamp-
ton, N. H. Exposed open coast.
432
Station
Al
A2
A3
A4
A5
A6
AT
A8
A9
A10
A11
A12
A13
A14
A15
A16
A17
A18
B1
B2
B3
B4
B5
B6
Temperature
(C)
16 - 20
19 - 19.5
16 - 20.5
19 - 20.8
18 - 21.5
20.5
20.5 - 29.5
15.0 - 19.8
(-1.0) - 22.5
24.0
21.5
9.0 - 10.0
8.0
7.0
8.0
9.0
0.0
0.0
16.0
17.0 - 18.0
17.0 - 19.0
16.5
16.0 - 16.5
2.0 - 17.0
18.0
18.0
16.0 - 16.5
16.5 - 20.0
16.0 - 19.0
16.0 - 19.0
Rhodora
Table II
Surface Water Temperatures and Salinities
Salinities
(0/00)
31 - 33.0
31 - 31.5
31 - 32.0
31 - 32.0
31 - 31.3
32.0
30.5 - 32.0
20.2 - 30.0
33.5
31.3
28.6 - 30.4
27.4 - 27.6
24.0
23.9
30.6
24.7 - 24.9
30.6 - 30.8
26.3
22.8 - 30.9
30.5 - 30.9
30.5
24.4 - 30.3
20.7 - 29.9
28.9
28.9
17.2 - 18.2
16.2 - 31.6
14.2 - 30.9
13.2 - 30.9
28.9
29.4
28.5
25.6
26.0
25.2
25.2
25.6
[Vol. 74
Dates*
12-21-69
10-04-66
11-18-66
11-18-66
11-18-66
11-18-66
11-13-66
11-13-66
12-06-69
12-06-66
1972] Marine Algae — Mathieson and Fralick 433
C8 15.5 24.5
C9 13.2 - 16.2 11.5 - 14.5 10-09-69
C10 10.0 27.2 10-13-66
C11 11.0 26.8 10-22-66
D2 13.0 23.6 11-11-66
D3 13.0 20.2 11-11-66
H1 16.5 30.5
H2 16.8 90.5
H4 1.0 30.5 12-21-66
Bound Rock 0.0 - 19.0 29.5 - 32.0 7-18-66
* All dates from August and September, 1969 except those specified.
Table III. Number of Seaweed Taxa of Various Stations
Stations Total Chlorophyceae Phaeophyceae Rhodophyceae
Al 19 5 3 4
A2 14 1 3 4
A3 11 6 2 3
A4 13 5 3 5
A5 12 5 4 3
A6 5 3 2 -—
AT 12 5 4 3
A8 10 4 3 3
A9 11 T 3 1
A10 11 3 3 5
A11 7 2 2 3
A12 7 1 1 5
A13 8 3 3 2
A14 12 2 6 4
A15 8 4 = 4
A16 2 2 Es =
A17 15 4 6 5
A18 8 3 3 2
B1 15 8 5 2
B2 19 8 5 6
B3 13 6 3 4
B4 15 8 3 4
B5 E 5 2 4
B6 19 9 3 1
B7 10 6 2 2
B8 7 4 1 2
B9 11 5 4 2
B10 16 5 6 5
434 Rhodora [Vol. 74
Bll 5 2 2 1
B12 T 5 2 —
B13 6 3 3 0
C1 17 7 7 3
C2 13 6 4 3
C3 14 8 4 2
C4 11 6 2 3
C5 19 8 5 6
C6 15 7 5 3
C7 6 4 1 1
C8 1 1 — —
C9 10 5 2 3
C10 5 2 3 —
C11 11 3 6 2
D1 — — — —
D2 4 — 4 —
D3 6 — 4 2
H1 14 3 5 6
H2 1 — — 1
H3 18 6 5 7
H4 0 - — —
HB-Bound
Rock 79 23 21 35
HB1 30 3 8 19
HB2 30 1 10 19
HB3 30 0 10 20
HB4 38 4 9 25
REFERENCES
CARDINAL, A. 1968. Répertoire des Algues Marines Benthiques de
L’est du Canada. Cahiers d’Information #48, Sta., Biol. Mar.,
Grand Riviere, mimeographed, 213 pp.
CHAPMAN, V. J. 1940. Studies in salt marsh ecology. VI and VII.
J. Ecol. 28: 118-152.
Davis, R. B. 1956. An ecological study of a tidal salt marsh and
estuary. M. S. thesis, University of New Hampshire, 102 pp.
DRUM, R. and E. E. WEBBER. 1965. Diatoms from a Massachusetts
salt marsh. Bot. Mar. 9: 70-77.
EDELSTEIN, T. L., C. M. CHEN and J. MCLACHLAN, 1970. The life
cycle of Ralfsia clavata and Ralfsia borneti. Can. J. Bot. 48:
527-531.
1972] Marine Algae — Mathieson and Fralick 435
HEHRE, E. and A. C. MATHIESON. 1970. Investigations of New Eng-
land marine algae III: Composition, seasonal occurrence, distri-
bution and reproductive periodicity of the marine Rhodophyceae
in New Hampshire. Rhodora 72: 194-239.
HOoEK, C. VAN DEN. 1963. Revision of the European species of
Cladophora. xi 4- 248 pp, Leiden.
LEWIS, J. R. 1964. The ecology of rock shores. xii + 323 pp. Eng-
lish University Press Ltd., London.
MATHIESON, A. C. E. HEHRE and N. REYNOLDS. Investigations of
New England marine algae I. A floristic and descriptive eco-
logical study of the marine algae at Jaffrey Point, New Hamp-
shire. Nova Hedwigia (in press).
MATHIESON, A. C., N. REYNOLDS and E. HEHRE. ibid II. The species
composition, distribution and zonation of seaweeds in the Great
Bay Estuary System and the adjacent open coast of New Hamp-
shire. Nova Hedwigia (in press).
MILLER, W. R. and F. E. EGLER. 1950. Vegetation of the Wequete-
quock-Pawcatuck tidal marshes, Connecticut. Ecol. Mon. 20:
143-172.
PARKE, M. and P. S. Drxon. 1968. Check-list of British marine
algae — second revision. J. mar. biol. Ass. U. K. 48: 783-832.
ScAGEL, R. F. 1966. Marine algae of British Columbia and northern
Washington, Part I: Chlorophyceae (green algae). Nat. Mus.
Canada, Biol. Ser. No. 74, Bull. No. 207; viii + 257.
TAYLOR, W. R. 1957. Marine algae of the northeastern coast of
North America. viii + 509 pp. Univ. Michigan Press, Ann Arbor.
VAGENAS, G. 1969. An ecological study of the Hampton-Seabrook,
New Hampshire salt marshes. M. S. Thesis, University of New
Hampshire, 74 pp.
WEBBER, E. E. 1967. Blue-green algae from a Massachusetts salt
marsh. Bull. Torrey Bot. Club. 94: 99-106.
1968. Seasonal occurrence and ecology of salt
marsh phanerogams at Ipswich, Massachusetts, Rhodora 70:
442-450.
WEBBER, E. E. and R. T. WILCE. 1971. Benthic salt marsh algae
at Ipswich, Massachusetts. Rhodora 73: 262-291.
WILCE, R. T. 1965. Studies in the genus Laminaria III. A revision
of the North Atlantic species of the Simplices Sections of Lami-
naria. Botanica Gothoburgensia III, pp. 247-256.
DEPARTMENT OF BOTANY
AND
JACKSON ESTUARINE LABORATORY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824
EDGEWORTHIA (THYMELAEACEAE)
NEW TO THE WESTERN HEMISPHERE
WILBUR H. DUNCAN AND MARIE MELLINGER
Edgeworthia chrysantha Lindl, a shrub previously
known to be native only from eastern Asia, has been found
growing wild in Rabun County in northeastern Georgia in
the Blue Ridge Mountains. Plants were first observed 15
September 1971 by the junior author. They are now known
to occur at intervals along 2.2 miles of Wolf Creek in de-
ciduous woods at elevations of about 1200-1600 feet. The
woods have had selective cutting but have never been
cleared.
Many colonies of the Edgeworthia occur along Wolf
Creek. One consists of at least 500 plants and extends along
the banks of the stream for about 1/4 mile. The largest
plant is 3.5 m tall and the stem 4 cm in diameter 15 cm
above the ground. Plants vary in size, those on the stream
bank being the most robust with smaller plants extending
6 m from the stream bank. Plants 75 cm or more tall
usually bear inflorescences.
Open flowers were found on some plants on 26 February
1972. These flowers and vegetative material collected be-
fore frost enabled us to identify the plants positively to
species. Specimens with full sized but immature fruits
were collected 21 May 1972. All characteristics of the
plants compare well with those of 23 specimens which we
have seen from Japan. Edgeworthia chrysantha was intro-
duced into Japan for cultivation as a source of paper and
is now widely escaped there (Ohwi, 1965).
Is Edgeworthia native to the Southeastern States? We
have found no reports to substantiate or repudiate such a
theory. It is especially significant that neither Small (1933)
nor Nevling (1962) reports the genus for this area. Ap-
parently it has been reported as native only from China to
Himalaya (Hooker, 1885; Nakai, 1924; Hamaya, 1955;
Makino, 1957; Hutchinson, 1967). Fairchild (1903) re-
ports the introduction of the species into the United States
from Japan and later (1908) says that planting trials at
Washington, D.C., and elsewhere indicate it will prove
436
1972] Edgeworthia — Duncan and Mellinger 437
hardy as far north as the Carolinas and will seed freely.
Bailey and Bailey (1941) state that E. chrysantha “— is
introduced in N. Amer.” The Rabun County plants may
be from one of these introductions, however, we have no
data to indicate the species was introduced there or that it
has escaped cultivation from elsewhere. Yet, it is incred-
ible to think that the plants are an eastern Asiatic dis-
junct.
A few vascular plant species occur in both the Southeast-
ern United States and China and/or Japan, e.g. Mitchella
repens L. All are more widespread in both hemispheres
than is the Edgeworthia in Rabun County. In addition,
there are many genera in the Southeastern United States
that have different species occurring in Asia, but no pair
in any genus has the strong similarities as do our Edge-
worthia plants and the 23 specimens we have seen from
Japan. We are continuing our search for information which
might indicate the Rabun County plants are introduced or
escaped. If such data are lacking, then, because of the cir-
cumstances under which the plants are growing, we think
the colony could be native. We solicit any information that
might be of help.
Obviously descriptions of E. chrysantha are not readily
available to most persons in the United States. Therefore,
a short description is presented below along with a partial
synonomy.
Edgeworthia chrysantha Lindl.
(E. papyrifera (Sieb.) Sieb. et Zucc., E. tomentosa
(Thunb.) Nakai, E. gardneri of authors, not Meissner)
Deciduous shrub to 3.5 m tall, usually ternately branched.
Twigs yellow-brown, appressed-pubescent when young. Leaf
scars raised about 45° at the lower end, with one crescent-
shaped bundle scar. Leaves petioled, the blades lanceolate
to sometimes oblong; acute to shortly acuminate or rarely
obtuse at the apex; cuneate to acuminate at the base; 8-25
em long; thinly silky, especially below; the hairs .25-.75
mm long, appressed, and pointing toward the tip. Inflores-
cences are in axillary peduncled heads which develop be-
fore the leaves fall and are conspicuous during the winter.
438 Rhodora [Vol. 74
Flowers fragrant, 20-50 in each head. Heads subtended by
several densely silky ovate to oblong bracts which usually
fall before the flowers open. The heads droop early, at
least by the time that all flowers have opened. Flowers
silky hairy on outside, apetalous, 10-18 mm long, the calyx
lobes 2-3 mm long and yellow inside. Stamens 8 in two
series of 4 on inside of floral tube, filaments very short.
Ovulary nearly sessile, hairy, 1-locular. Styles 3-4 mm
long. Stigma clavate. The entire pistil 7-9 mm long. The
flowers open as early as late February. The fruits mature
in summer. They are hairy at the apex, 6-7 mm long, el-
lipsoid to ovate, dry, green, and often surrounded by the
somewhat persistent floral tube. Voucher specimens are in
the University of Georgia Herbarium: Mellinger, 15 Sep-
tember 1971; Duncan 23751, 21 May 1972 (fruiting); in
deciduous woods along Wolf Creek, Rabun Co., Georgia,
Blue Ridge Province.
Edgeworthia as presently interpreted apparently consists
of four species. They have been separated mostly on the
basis of vestures or the lack of any, flower colors, and pe-
duncle lengths. E. longipes Lace, which occurs in upper
Burma, has peduncles 9-11 cm long. In other species they
are only 1-2.5 em long. E. albiflora Nakai, which occurs
in Yunan and Szechwan Province, China, may be recognized
by the whitish interior of the calyx and glabrous under-
sides of the leaves. In the remaining two species, E. gard-
neri Meisner and E. chrysantha, the leaves are hairy
beneath (sometimes faintly so) and the flowers are yellow
inside. The flowers appear with the leaves in the former
and before the leaves in the latter. A much more reliable
character apparently has not been used previously; the in-
volucral bracts are ovate to oblong in E. chrysantha and
linear (up to 35 mm long) in E. gardneri. There seems to
be a short-bracted and a long-bracted form of E. gardneri.
These forms may deserve taxonomic recognition and should
be studied. The long-bracted form occurs from Nepal into
northern Burma, and the short-bracted form from eastern
Nepal into southwestern China. Edgeworthia chrysantha
occurs in the southern half of China and as an adventive in
1972] Edgeworthia — Duncan and Mellinger 439
Japan. It is considered by Nakai (1929) as two species
which are separated by stoutness of twigs and size of leaf
scars. We, however, agree with Hamaya (1955) that there
is a gradual transition in sizes and there is no basis for
division into two species.
We acknowledge with appreciation the loan of specimens
from several herbaria, and information provided by Drs.
Lorin I. Nevling, Richard A. Howard, and Frederick G.
Meyer.
LITERATURE CITED
BAILEY, L. H. and E. Z. BAILEY. 1941. Hortus Second. Macmillan
Co. N.Y.
FAIRCHILD, Davip. 1903. Mitsumata, a Japanese Paper Plant.
U.S.D.A., Bureau of Plant Industry, Bull. No. 42: 9-11.
1908. Cultural Directions for the Mitsumata,
a Japanese Paper Plant, U.S.D.A. Bureau of Plant Industry.
Misc. Leaflet. 2 p.
HAMAYA, ToSHIO. 1955. A Dendrological Monograph of the
Thymelaeaceae Plants of Japan. Bull. of the Tokyo University
Forests. No. 50: 246-248.
HOOKER, Sir J. D. 1885. Flora of the British India, v. 5. Cheno-
podiaceae to Orchidaceae. L. Reeve & Co., Ltd., Ashford, Kent,
England.
HUTCHINSON, JOHN. 1967. The Genera of Flowering Plants. Ox-
ford at the Clarendon Press. 659 p.
MAKINO, ToMiLARO. 1957. An Illustrated Flora of Japan with
Cultivated and Naturalized Plants (Enlarged Ed.) The Hoku-
ryukan C., Ltd., Tokyo. 1304 p. & indices.
NAKAI, T. 1924. Some New and Noteworthy Ligneous Plants of
Eastern Asia. Journal of the Arnold Arboretum 5: 81-83.
NEVLING, LoRIN L, JR. 1962. The Thymelaeaceae of the Southeast-
ern United States. Journal of the Arnold Arboretum 43: 428-
434.
OHWI, JISABURO. 1965. Flora of Japan. Smithsonian Institution,
Washington, D.C. 1067 p.
SMALL, J. K. 1933. Manual of the Southeastern Flora. Published
by the author. N.Y. 1554 p.
DEPARTMENT OF BOTANY
UNIVERSITY OF GEORGIA
ATHENS, GEORGIA 30601
ROUTE 1, TIGER, GEORGIA 30576
REQUEST FOR UNNEEDED COPIES OF RHODORA
FOR DECEMBER 1971, NO. 796
The error on the cover of the fourth number of Rhodora
for 1971, indicating it as September 1971, no. 795, has
resulted in many persons discarding it as a duplicate. Had
they looked inside the cover they would have detected the
error immediately. However so many requests have come
in for replacements that Dr. Herman Sweet requests that
any subscribers who have this December issue, and do not
regularly keep a file of Rhodora, send this particular copy
to him at the Botanical Museum, Oxford Street, Cambridge,
Mass. 02138.
INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA
Manuscripts must be double-spaced or preferably triple-
spaced (not on corrasable bond), and a list of legends for
figures and maps provided on a separate page. Footnotes
should be used sparingly, as they are usually not necessary.
Do not indicate the style of type through the use of capitals
or underscoring, particularly in the citations of specimens,
except that the names of species and genera may be under-
lined to indicate italics in discussions. Specimen citations
should be selected critically especially for common species
of broad distribution. Systematic revisions and similar
papers should be prepared in the format of “The System-
atics and Ecology of Poison-Ivy and the Poison-Oaks,"
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu-
larly with reference to the indentation of keys and syno-
nyms. Papers of a floristic nature should follow, as far as
possible, the format of “Contribution to the Fungus Flora
of Northeastern North America. V.," H. E. Bigelow & M. E.
Barr, Rhodora 71: 177-203. 1969. For bibliographic cita-
tions, a recommended list of standard journal abbreviations
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Pot.
Club 85: 277-300. 1958.
Volume 74, No. 799, including pages 301-440, was issued September 29, 1972.
440
CONTENTS: continued
Variation within Calochortus venustulus (Liliaceae).
James L. Reveal and William J. Hess ..................... eee 350
New Form of Thuja occidentalis Resembling Known Cul-
tivars.
Mary- NIE. MERE T 352
Snow Cover and the Diapensia lapponica Habitat in the
White Mountains, New Hampshire.
Weasley N. Tifney; JY. ————— 358
New and Interesting Plants from the Great Plains.
Ronald L. McGregor, Lawrence K. Magrath and
Ronald R. Weedon ........ ,2 eee nennen ntn eet nennt 378
New Records of North Dakota Plants.
O. A. Stevens eese eese sanete tesa asta stet e sess aae ee eee eese eee bes ee 389
Priority of the Binomial Pyrola chlorantha.
Erich Haber PP S 396
Chromosome Numbers of Scirpus purshianus and S. smithii.
VU ME ONES] 11771 1 RR 398
Tussilago farfara L. (Senecioneae-Asteraceae) in Virginia.
Miles F. Johmsom crsccccccccssssesssscessscceccccevesenssecsssseeceesceeeeeeeseneeees 408
Investigations of New England Marine Algae V. The Algal
Vegetation of the Hampton-Seabrook Estuary and the
Open Coast near Hampton, New Hampshire.
Arthur C. Mathieson and Richard A. Fralick .................. 406
Edgeworthia (Thymelaeaceae) New to the Western Hem-
isphere.
Wilbur H. Duncan and Marie Mellinger ............................ 436
Request from Editor ................... eere eene 440
Instructions for Contributors to Rhodora .................................. 440
Hovdova
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by
ALBION REED HODGDON, Editor-in-Chief
ALBERT FREDERICK HILL
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON > Associate Editors
RADCLIFFE BARNES PIKE
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
Vol. 74 December, 1972 No. 800
CONTENTS:
Taxonomic Fern Notes, VI— New Species of American
Cyatheaceae.
Rolle, Tryon .srececssscseccssccscrsserssscececevsssesscscecececscsocsonscnesenossnssesesese 4. 441
New Species in the Phacelia crenulata Group
(Hydrophyllaceae)
N. Duane Atwood E AAA A E 451
Rediscovery of Tilia neglecta Spach.
Henry K. Svenson aaa ana tte Des 469
The Ecology of Benthic Salt Marsh Algae at Ipswich, Massa-
chusetts I. Zonation and Distribution of Algal Species.
E. E. Webber and R. T. Wile ette 475
(Continued on Inside Back Cover)
The Nef England Botanical Club, Ine.
Botanical Museum, Oxford St., Cambridge, Mass. 02138
STATEMENT OF OWNERSHIP, MANAGEMENT AND
CIRCULATION
Title of Publication: RHODORA
Date of Filing: Sept. 25, 1972
Frequency of Issue: Quarterly
Location of Known Office of Publication:
Botanical Museum, Harvard University, Oxford Street,
Cambridge, Mass. 02138
Location of the Headquarters of the Publishers:
Botanical Museum, Harvard University, Oxford Street,
Cambridge, Mass. 02138
Names and Addresses of Publisher, Editor, and Managing Editor:
Publisher:
New England Botanical Club, Inc., Botanical Museum,
Oxford Street, Cambridge, Mass. 02138
Editor:
Albion R. Hodgdon, Dept. of Botany,
University of New Hampshire, Durham, N. H. 03824
Managing Editor:
Herman R. Sweet, Botanical Museum, Harvard University,
Oxford Steet, Cambridge, Mass. 02138
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complete. Herman R. Sweet, Business Manager
MANUFACTURED BY
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Rhodora
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Vol. 74 December, 1972 No. 800
TAXONOMIC FERN NOTES, VI — NEW SPECIES OF
AMERICAN CYATHEACEAE.
ROLLA TRYON
The many ample collections of tree ferns made during
explorations in South America by Julian Steyermark, Bas-
sett Maguire, John Wurdack, Ramón Ferreyra, Benkt
Sparre, and Paul Hutchison and Kenneth Wright are an
essential source of information for studies of the Ameri-
can Cyatheaceae now in progress. Several of the new
species among them are described here in addition to the
species of tree fern that is characteristic of the mossy
forest of Puerto Rico.
Some of the endemism in the Cyatheaceae in Guayana is
reflected in the four new species from that region. Al-
though endemism in the ferns in this area is not as strong
as in many groups of flowering plants, it is significant
especially with respect to the remarkable morphological
innovations.
The species are described under the genera recognized
in my recent paper on the classification of the family
(Tryon, 1970) and special terms employed are defined
there. The illustrations have been prepared by Sally
Landry, Mary Robbins, and Lydia Wunsch.
Sphaeropteris. sipapoensis Tryon, spec. nov. Figs. 1-4.
Petiolus sine spinis squamis multis persistentibus struc-
tura conformi circa 1-2 cm longis brunneolo-albis manifeste
ciliatis; lamina bipinnata pinna apicali subconformi; rhachis
441
442 Rhodora [Vol. 74
squamata; pinnae ad 40 cm longae petiolulatae; pinnulae
integra vel aliquantum lobatae subcoriaceae articulatae
breviter petiolulatae base late cordata apice subacuto vel
acuminato pagina superiore glabra pagina inferiore squamis
valde dissectis ; venae liberae; sori submarginales plerumque
a margine circa 0.5 mm exindusiati paraphysibus longiori-
bus quam sporangiis albidis circa 1 mm longis. Holotypus:
Mixed forest at Intermediate Camp, Cerro Sipapo (Pará-
que), Terr. Amazonas, Venezuela, 2 Feb. 1949, B. Maguire
and L. Politi 28165, NY. Isotypus: GH.
Sphaeropteris sipapoensis is closely related to S. margi-
nalis (Kl) Tryon of British Guiana and Surinam. The
two species share the following characters: large, brown-
ish-white, long-ciliate petiole scales, stalked pinnules, a
nearly conform terminal pinna at the lamina apex and the
absence of an indusium. Sphaeropteris marginalis has the
sori farther (mostly 1.2-2.0 mm) back from the margin
than S. sipapoensis and the paraphyses are shorter (ca.
0.5 mm long) and tan to brown in color. Sphaeropteris
marginalis also has smaller, less coriaceous, and more deeply
lobed pinnules than S. sipapoensis. Further collections
are required in order to assess the importance of these
pinnule characters.
The related S. marginalis has been reported from Mexico
(for example, by Maxon and Morton, p. 70, 1948) but this
is based on an erroneous association of a specimen of the
species at Kew ! with the label of Jurgensen 915, Sierra
San Nolasco, Mexico. Fournier (p. 135, 1872) cites
Jurgensen 915 as Hemistegia decurrens [= Cnemidaria
decurrens (Liebm.) Tryon], a species of Mexico and un-
doubtedly the one properly associated with the label.
Sphaeropteris Atahuallpa Tryon, spec. nov. Figs. 5-8.
Petiolus parce tuberculatus squamis structura conformi
ad 6 cm longis 1 cm latis brunneolo-albis basem versus
densissimis persistentibus squamis minoribus longe ciliatis
apice filiformi concolori squamellis multis valde dissectis
etiam instructus; lamina bipinnato-pinnatifida vel juxta
1972] Cyatheaceae — Tryon 443
rhachim bipinnato-pinnatisecta apice sensim acuminato;
pinnae ad 70 cm longae subsessiles; pinnulae subsessiles
segmentis minime crenato-serratis costulis inferne squamis
albidis; venae liberae; sori ad furcam venarum; indusium
subfuscum sphaeropteroideum. Holotypus: Cerros Calla
Calla, above Balsas on road to Leimebamba, 3000-3100 m,
Prov. Chachapoyas, Dept. Amazonas, Peru, 14 Oct. 1964,
P. C. Hutchison and J. W. Wright 6922, GH. Paratypus:
Summit of Puma-urcu, southeast of Chachapoyas, 3100-3200
m, Prov. Chachapoyas, Dept. of Amazonas, Peru, 3 July
1962, J. J. Wurdack 1153, GH.
Sphaeropteris Atahuallpa is a distinctive species, related
by its sphaeropteroid (globose) indusium only to the species
of the Sphaeropteris horrida group (Tryon, 1971). From
these, however, it differs in the petiole scales which lack
a dark apical seta and dark rigid marginal teeth. The
largest petiole scales are five to six cells thick near their
base. Atahuallpa, a son of Huayna Capac, was Lord-Inca
of the Kingdom of Quito. He was captured at Cajamarca
by Francisco Pizarro, then ransomed with silver and gold
to the present value of perhaps fifty million dollars, and
finally betrayed and murdered by Pizarro.
Alsophila bryophila Tryon, spec. nov. Figs. 9-10.
Petiolus brevissimus modice tuberculatus squamis struc-
tura marginata brunneis apice seta minima brunnea fragili;
lamina pinnato-pinnatisecta vel juxta rhachim bipinnata
basem versus modice vel sensim deminuta apice sensim
acuminato; rhachis pubescens plus minusve squamata;
pinnae ad 15-25 cm longae sessiles costa pagina superiore
pubescenti pagina inferiore pubescenti saepe squamis sub-
planis sine setis fuscatis; segmenta integra coriacea utrin-
que longe pubescentia costulis basem versus squamis paucis
albidis bullatis sine setis; venae liberae; sori ad furcam
venarum; indusium hemitelioideum vel meniscoideum.
Holotypus: Near West Peak, Luquillo National Forest,
Puerto Rico, 12 Nov. 1966, G. J. Gastony 12, GH. Para-
typi (all from Puerto Rico): Howard, et al. 15645, 15722,
444 Rhodora [Vol. 74
EN
US
NN:
NW
Y
a
L / // ]] |]
(IJ
Wy
XM NOI
Figs. 1-4. Sphaeropteris sipapoensis, (Maguire and Politi 28765).
Fig. 1, basal pinna, X 1/3, GH. Fig. 2, central pinnules of central
pinna, X 1/3, Ny. Fig. 3, portion of fertile pinnule (paraphyses
removed from receptacles to left), X 1, NY. Fig. 4, articulate pin-
nule base, enlarged, NY.
Figs. 5-8. Sphaeropteris Atahuallpa (Hutchinson and Wright 6922,
GH). Fig. 5, apical portion of crozier, X 1/3. Fig. 6, portion of
petiole, X 1/3. Fig. 7, central pinnules of central pinna, X 1/3.
Fig. 8, fertile segments: sori and venation (left), with scaly indu-
ment (right), X 1 1/3.
Figs. 9-10. Alsophila bryophila (Gastony 12, GH). Fig. 9, cen-
tral pinna, X 1/3. Fig. 10, fertile segments: indusia (upper right),
with pubescence (upper to left), venation and receptacles (lower),
x 1 1/3.
1972] Cyatheaceae — Tryon 445
15725, A; Howard and Nevling 15740, A; Shafer 3305,
GH, MO; Shafer 3632, F, NY; Sintenis 1480, GH.
Alsophila bryophila grows in wet forests in Puerto Rico
at elevations of about 900-1000 m. The species is char-
acteristic of the mossy (elfin) forest of the Luquillo
Mountains and the specific epithet is derived from this
association.
Alsophila bryophila is close to A. minor (D. C. Eaton)
Tryon, A. hotteana (C. Chr. and Ekman) Tryon and A.
Brooksii (Maxon) Tryon, from which it can be distin-
guished by the hemitelioid to meniscoid indusium and the
absence of dark setae on the scales of the under surface of
the pinnae. The three related species all have a cup-shaped
indusium and some, frequently many, scales or squamules
on the under surface of the pinnae with dark setae on the
margins and a dark seta at the apex. They also have the
upper surface of the segments glabrous to rarely sparingly
pubescent, which in A. bryophila the segments are usually
strongly pubescent above. Alsophila Brooksii, also in Puerto
Rico, is fully bipinnate through most of the lamina. Some
materials of A. hotteana have a meniscoid indusium but in
these specimens it is not certain that the structure is com-
plete. They do have the distinguishing dark-armed indu-
ment on the axes beneath.
The other species of American Alsophila: A. Abbott
(Maxon) Tryon, A. capensis (L. f.) J. Sm., A. confinis
(C. Chr.) Tryon, A. dryopteroides (Maxon) Tryon, A.
Engelii Tryon, A. Nockii (Jenm.) Tryon, A. paucifolia
Baker, A. Salvinii Hook. and A. Urbanii (Brause) Tryon,
differ from A. bryophila in having a cup-shaped indusium
and (or) in having a fully bipinnate or more complex
lamina, and in different indument on the under surface of
the pinnae.
Alsophila bryophila has long been known under the
name of Cyathea pubescens Kuhn. That species, however,
is Nephelea pubescens (Kuhn) Tryon and is endemic to
the Blue Mountains of Jamaica (Gastony, 1972). Cyathea
446 Rhodora [Vol. 74
pubescens has also been ascribed to Cuba and Hispaniola
(Maxon, p. 386, 1926), but the specimens from those
islands belong to Alsophila minor.
Trichipteris Steyermarkii Tryon, spec. nov. Figs. 11-12.
Petiolus basem versus acute tuberculatus squamis struc-
tura marginata; lamina bipinnata pinna apicali conformi;
pinnae circa 25-35 cm longae articulatae breviter petiolu-
latae; pinnulae integrae vel subintegrae coriaceae articu-
latae breviter petiolulatae base cordata apice subacuto vel
acuto pagina superiore glabra pagina inferiore squamis pau-
cis brunneolis subplanis; venae liberae; sori ad furcam vena-
rum exindusiati paraphysibus brevioribus quam sporangiis.
Holotypus: Bosque nublado virgen en la cumbre, Cerro de
Humo, Peninsula de Paria, Estado Sucre, Venezuela, 2 Mar.
1966, J. A. Steyermark 94923, VEN. Isotypi: GH, U.
Paratypus: Selva nublada en la cumbre plana, Cerro
Patao, Peninsula de Paria, Estado Sucre, Venezuela, 19 Jul.
1962, J. A. Steyermark and G. Agostini 91048, VEN.
Trichipteris Steyermarkii is related in its lamina archi-
tecture and articulate pinnae and pinnules to T. sagittifolia
of Trinidad and to T. cordata of the states of Aragua and
Carabobo, Venezuela. Both species differ from T. Steyer-
markii in having bullate scales on the lower surface of the
pinnules (T. sagittifolia also has some flattish ones).
Futher differences are the many long slender spines at the
base of the petiole of T. sagittifolia and the pinnately lobed
pinnules of T. cordata.
Hooker (p. 37, 1865) questioned the source of the Cruger
collection upon which Alsophila sagittifolia was based:
"Trinidad ? (possibly the opposite coast of Venezuela)".
The type collection (Kew !) has bullate scales on the pin-
nules and corresponds with the Trinidad species (for ex-
ample, Fendler 142 and 143).
Trichipteris Cyclodium Tryon, spec. nov. Figs. 13-14.
Petiolus sine spinis squamis structura marginata brun-
neis albido-limbatis dense ciliatis; lamina 1-pinnata pinna
1972] Cyatheaceae — Tryon 447
apicali conformi articulata; pinnae circa 15-18 cm longae
valde ascendentes integrae late crenatae vel minime pin-
natilobatae valde coriaceae articulatae petiolulatae utrinque
glabrae vel subglabrae base late cuneata apice obtuso; venae
liberae utrinque elevatae; sori plerumque in venis simpli-
cibus 3-4 seriati exindusiati paraphysibus longioribus quam
sporangiis. Holotypus: Camp Savanna, Cerro Sipapo
(Paráque), Terr. Amazonas, Venezuela, 6 Dec. 1948, B.
Maguire and L. Politi 27541, NY. Isotypus: GH.
Trichipteris Cyclodium is a very distinctive species. The
lamina has a close resemblance to that of Cyclodium menis-
cioides (Willd.) Presl (a species that can also be placed
in Stigmatopteris or Dryopteris, depending upon how
generic lines are drawn). Trichipteris Cyclodium is related
to T. Williamsii (Maxon) Tryon of Panama and to the
next species T. Maguirei by its simple, articulate and stalked
pinnae and by its lamina with a conform terminal pinna.
It differs from T. Williamsii in its very coriaceous, strongly
ascending, obtuse pinnae and whitish-bordered petiole
scales. Trichipteris Williamsii has the pinnae only moder-
ately thickened, patent and long-acuminate and the petiole
seales are entirely whitish or have a brown stripe toward
the apex. A comparison with T. Magwuirei is made under
that species.
Trichipteris Maguirei Tryon, spec. nov. Figs. 15-16.
Petiolus sine spinis squamis structura marginata brunneis
albido-limbatis dense ciliatis ; lamina 1-pinnata pinna apicali
conformi articulata vel apice binato; pinnae circa 8-12 cm
longae integrae coriaceae articulatae utrinque glabrae
pinnis basilaribus circa 7 mm petiolulatis base cuneata
apice abrupte acuminato serrato; venae liberae vel aliquot
junctae utrinque elevatae; sori plerumque in venis simpli-
cibus 1-2 seriati exindusiati paraphysibus longioribus quam
sporangiis. Holotypus: Frequent, terrestrial, talus forest
near first escarpment, Cerro Sipapo (Paráque), Terr.
Amazonas, Venezuela, 11 Dec. 1948, B. Magwire and. L.
Politi 27597, NY. Isotypus: GH. Paratypi: (all from
448 Rhodora [Vol. 74
Figs. 11-12. Trichipteris Steyermarkü (Steyermark 94923, VEN).
Fig. ll, central pinna, X 1/3. Fig. 12, fertile pinnule, with recep-
tacles and scales, X 1.
Figs. 13-14. Trichipteris Cyclodium (Maguire and Politi 27541,
NY). Fig. 18, central pinnae, X 1/3. Fig. 14, portion of fertile pinna,
with sori, X 2/3.
Figs. 15-16. Trichipteris Maguirei (Maguire and Politi 27597,
GH). Fig. 15, central pinnae, X 1/3. Fig. 16, portion of fertile pinna,
with sori, X 1.
Figs. 17-19. Cyathea Steyermarkii (Steyermark 105194, GH),
Fig. 17, lamina apex, X 1/3. Fig. 18, central pinnules of central
pinna, X 1/3. Fig. 19, portion of fertile pinnule, with receptacles,
X 1.
1972] Cyatheaceae — Tryon 449
Terr. Amazonas, Venezuela) Terrestrial, uncommon, wet
montane moss forest, Phelps Camp to North Savanna,
1400 m, Cerro Sipapo (Paráque), 17 Dec. 1948, B. Maguire
and L. Politi 27752, NY; terrestrial, wet mixed montane
forest, slope above Cano Grande, 1500 m, Cerro Sipapo
(Paráque), 21 Jan. 1949, B. Maguire and L. Politi 28493,
NY; dense woodland along right fork of Cano de Dios,
1900 m, Cerro Huachamacari, Río Cunucunuma, 13 Dec.
1950, B. Maguire, R. S. Cowan and J. J. Wurdack 30188,
NY.
Trichipteris Maguirei differs from the preceding species,
T. Cyclodium, in its smaller pinnae that are cuneate at the
base and abruptly acute to acuminate at the apex, and in
the lamina that often has a pair of terminal pinnae. The
larger pinnae of T. Cyclodium are broadly cuneate at the
base and obtuse at the apex, and the lamina has a single
terminal pinna. Tvichipteris Maguirei is perhaps more
closely related to T. Williamsii, but differs from it in the
smaller pinnae, ca. 8-12 cm. long, the shorter pinna stalks,
those of the basal pinnae being ca. 7 mm long, and the
petiole scales with whitish borders. In T. Williamsii, the
pinnae are ca. 15-20 cm long, the stalks of the basal ones
are ca. 13 mm long, and the petiole scales are entirely
whitish or have a brown stripe toward the apex.
Cyathea Steyermarkii Tryon. spec. nov. Figs. 17-19.
Petiolus basem versus modice muricatus squamis struc-
tura marginata brunneis brunneolo-albis-limbatis; lamina
bipinnata pinna apicali conformi articulata; pinnae circa
25-30 cm longae articulatae petiolulatae apice sensim
acuminato; pinnulae non articulatae crenatae vel aliquan-
tum pinnatilobatae chartaceae base cuneata apice obtuso vel
subacuto; venae liberae; sori plerumque in venis simplicibus
inter costam et marginem medii; indusium parvum hemi-
telioideum. Holotypus: Cumbre del Cerro Autana, 4° 52”
N. lat. 67° 27” W. long., 1230-1270 m, Terr. Amazonas,
Venezuela, 21-22 Sept. 1971, J. A. Steyermark 105194, GH.
450 Rhodora [Vol. 74
Cyathea Steyermarkii is related to three other species
with a hemitelioid indusium: Hemitelia conformis Tryon,
H. petiolata Hook. and H. Woronovii Maxon and Morton.
I placed the first two in Cnemidaria (Tryon, 1970) ; how-
ever, Robert Stolze, who is monographing the genus, in-
forms me that these species and H. Woronovii are to be
excluded and properly belong in Cyathea. Hemitelia petio-
lata is the only one of these hemitelioid species that has
been placed in Cyathea (as C. panamensis Domin). The
three species have the sori close to the margin, while in
C. Steyermarkii the sori are about half way between the
costa and the margin. Hemitelia petiolata and H. Woro-
novii have the apex of the lamina gradually reduced to the
tip, while in C. Steyermarkii the apex is a conform termi-
nal pinna. Hemitelia conformis is further distinguished
by the pinnae that have a conform, articulate terminal
pinnule. In C. Steyermarkii the pinnae are gradually re-
duced at the apex.
LITERATURE CITED
FOURNIER, E. 1872. Mission Sci. Mex. Amér. Cent. Recherches Bo-
taniques, I. Cryptogamie (Mex. Plantas, I. Cryptogamie). Paris.
GASTONY, G. J. 1972. A revision of the fern genus Nephelea. Con-
trib Gray Herb. 203. (in press)
Hooker, W. J. 1865. Synopsis Filicum. London.
Maxon, W. R. 1926. Pteridophyta of Porto Rico and the Virgin
Islands. Sci. Surv. Porto Rico and V. I. 6 (3): 378-521.
MAxon, W. R. and C. V. MORTON. 1948. Pteridophyta, i» B. Ma-
guire, et al., Plant explorations in Guiana in 1944. Bull. Torrey
Bot. Club 75: 66-80.
Tryon, R. 1970. The classification of the Cyatheaceae. Contrib.
Gray Herb. 200: 1-53.
1971. The American tree ferns allied to Sphaeropteris
horrida. Rhodora 73: 1-19.
GRAY HERBARIUM,
HARVARD UNIVERSITY,
CAMBRIDGE, MASS. 02138
NEW SPECIES IN THE PHACELIA CRENULATA
GROUP (HYDROPHYLLACEAE)'
N. DUANE ATWOOD
While preparing a taxonomic revision of the Phacelia
crenulata group it was necessary to investigate the rela-
tionships of several taxa in southern Utah, northern
Arizona and Mexico. These investigations demonstrated
the presence of several previously undescribed taxa. These
areas are well known for their varied geology, topography
and climate. They have also been some of the least bota-
nized and most poorly understood localities within the
range of the P. crenulata group. Therefore, it is not sur-
prising that new species are present, particularly in a
genus which contains many taxa with restricted distribu-
tion. The purpose of this paper is to present the diagnoses,
descriptions, distributions, and illustrations of the pro-
posed new taxa.
The herbaria where specimens are deposited are abbre-
viated according to Lanjouw and Stafleu (1964). Those
herbaria not listed in Lanjouw and Stafleu are as follows:
Dixie Junior College, St. George, Utah (DIX), Weber State
College, Ogden, Utah (WSC), and West Texas State Uni-
versity, Canyon, Texas (WTS).
The author is grateful to the curators of herbaria that
were visited or who sent loan material for this study, and
to Mrs. Kaye Thorne for preparation of the illustrations.
Phacelia constancei Atwood sp. nov.
A P. palmeri differt caulibus brevioribus, foliis angusti-
oribus et revolutioribus, et caulibus rubellis pilioribus et
ramis lateralibus frondosioribus.
Herba biennis erecta, 1.5-4.8 dm alta, frondosa omnino;
folia petiolata (pro parte maxima), 1.0-10 cm longa, 0.3-
"This study was supported in part by the Department of Botany
and Range Science, Brigham Young University, Provo, Utah, and
by Cost of Education Funds from a National Defense Education Act
Title IV Fellowship.
451
452 Rhodora [Vol. 74
1.5 em lata, involuta, e undulata ad pinnatifida, lineares
lanceolata, hirsutula supra, pilis glandularis dispersis;
caules crasses, simplices vel ramosi omnino, rubelli, e
hirsutuli ad hirsuti, et minute glanduliferi; inflorescentia,
cymarum scorpiodorum compositorum; pediceli usque ad
1 mm longi; sepala e elliptica ad oblanceolata, 3-4 mm longa,
hirsutuli ad hirsuti et stipitata glandulifera; corolla tubu-
laris, 5-6 mm longa; stamina exserta 3-4 mm; stylus fur-
catus pubescens infimus 1/3, exsertus 3-4 mm stamina
longior; capsula subglobosa, hirsutula et glandulifera
omnino, sepala brevior; semina 4, nigricantia, 2.5-2.8 mm
longa, 1.0-1.2 mm lata, elliptica, margines corrugati, pagina
ventralis foveolata, excavata et divisa porca prominens,
porca lata una corrugata, pagina dorsalis foveolata.
TYPE: Arizona: COCONINO COUNTY: 1 mi north of Fre-
donia, on clay hills, growing with Sarcobatus, Lycium,
Astragalus, and Phacelia corrugata, 27 May 1968, D. At-
wood 1385A (HOLOTYPE: BRY; ISOTYPES: ARIZ, ASC, CAS,
DIX, GH, NY, POM, RM, UC, US, UT, UTC, WSC).
Additional materials examined: Arizona: MOHAVE COUNTY: ca 19
mi south of Kanab Indian Reservation (southern boundary) on road
to Tuweep junction, 27 May 1969, D. Atwood 1765, (BRY); 5 mi
northeast of Tuweep, 17 August 1946, K. F. Parker, et al. 6254,
(GH, NY, POM, US); ca 4 mi north of Tuweep junction, 27 May 1969,
D. Atwood 1761, (BRY, CAS); 1 mi north of Fredonia, 5 June 1968,
D. Atwood 1529, (BRY, GH, UC) ; 1 mi north of Fredonia, 27 May 1969,
D. Atwood 1785, (ARIZ, BRY, CAS, NY, US, UTC, WSC); Utah: KANE
COUNTY: ca 30 mi east of Kanab along Hwy 89, mi post 30, 6 June
1968, D. Atwood 1532, (BRY, CAS, GH, NY, US); 15 mi east of Kanab,
5 June 1969, D. Atwood 1739A, (ARIZ, BRY, US, UT); 15 mi east of
Kanab, 5 June 1942, Ripley & Barneby 4836, (CAS, RSA); 20 mi east
of Kanab, 5 June 1969, D. Atwood 1792B, (BRY, CAS, GH, NY, POM,
RM, US); ca 32 mi east of Kanab along Hwy 89, 6 June 1968, D.
Atwood 1532A, (BRY, POM, RSA); ca 3.7 mi east of Skutumpah-Alton
junction at head of Johnston Canyon (R.4. 5W., T.408.), 7 June 1969,
D. Atwood 1801A, (BRY).
Taxonomically, Phacelia constancei (Figure 1, Map 1)
appears to be most closely related to P. palmeri Torr. ex
S. Wats., but is distinguished from that species by the
smaller growth form, narrower and more revolute leaves,
the stems are reddish with shorter and fewer hairs and
the lateral inflorescence branches are more leafy.
1972] Phacelia — Atwood 453,
Fig. 1. Phacelia constancei. Habit and seed of plant.
454 Rhodora [Vol. 74
^i
l
Map 1. Southern Utah and northern Arizona. Range of P. con-
stancei.
The latter occurs from Mohave County, Arizona, west to
Washington and Iron County, Utah, and in Clark County,
Nevada. The new taxon is known only from restricted
areas in Mohave County, Arizona north to Kane County,
Utah. Both species grow on alkaline clay bluffs and flats
of the Moenkopi Formation, late May to mid August.
Chromosome voucher specimen (n=11): Tuweep Valley,
5 mi ne. of Tuweep, 17 August 1946, K. F. Parker et al.
6254, (UC).
This species is named in honor of Dr. L. C. Constance,
University of California at Berkeley, who has made nu-
merous important contributions in the genus Phacelia.
Phacelia rafaelensis Atwood sp. nov.
A P. utahensis differt corolla tubularo diluta et canali-
culata lobis aliquatum expansis, staminibus et stylis exsertis
tantum 3-5 mm, filis et antheris tristis, semine porca inter-
dum corrugata.
1972] Phacelia — Atwood 455
Herba biennis, erecta, 0.8-5.4 dm alta; caules crassi
simplices vel interdum ramosi basi, e olivacei ad brunnei
glandiferi et hirsuti; folia basalia petiolata dentata ad
pinnatifida, 2-7 cm longa, 0.5-1.5 em lata, strigosa ad hir-
suta, folia caulina sessilia, undulati-crenata ad dentata,
oblonga ad lanceolata, 1.0-10 cm longa, 0.5-3.5 cm lata,
strigulosa ad hirsuta et sparsim stipitati-glandulifera ;
inflorescentia terminalis proparte maxima interdum axil-
laris, paniculata; flores fere sessiles; sepala oblonga ad
spathulata, 3-4 mm longa in fructis, 1.0-1.7 mm lata, gland-
ulifera et hirsuta; corolla tubularis pallida, 5-6 mm longa;
stamen et stylus exsertus 3-5 mm; stylus 34 longitudis,
pubescens infernus 14; capsula globosa, 4-5 mm longa,
stipitati-glandulifera et hirsuta; semina 4, elliptica ad
oblonga, 3.4-4.0 mm longa, 1.5-2.0 mm lata, pagina ven-
tralis alveolata pallidior pagina dorsalis, excavata et divisa
porca prominenti, porca lata una corrugata, margines
integrae, pagina dorsalis brunnea et alveolata radose.
TYPE: Utah: WAYNE COUNTY: Capital Reef National
Monument, red clay hills (Moenkopi Formation) southeast
of visitor center, 12 June 1969, D. Atwood & L. C. Higgins
1834, (HOLOTYPE: BRY; ISOTYPES, ARIZ, ASC, B, BRY, CAS,
COLO, DIX, GH, JEPS, MEXU, NY, POM, RM, RSA, UC, US, UT,
UTC).
Additional materials examined: Arizona: COCONINO COUNTY: Lee's
Ferry, 16 June 1890, M. E. Jones s.n, (POM); 2 mi w. of Navajo
Bridge, 12 May 1955, Ripley & Barneby 12664, (CAS); MOHAVE
COUNTY: 2 mi e. of Fredonia, 6 June 1942, Ripley & Barneby 4362,
(RSA); ca 4 mi w. of Fredonia, mi post 22, 27 May 1968, D. Atwood
1390, (ARIZ, ASC, BRY, CAS, POM, US); 1 mi n. of Fredonia, 5 June
1968, D. Atwood 1530, (BRY, B, POM); ca 4 mi s. of Fredonia-Pipe
Springs road, 13 May 1952, E. McKlintock 52-600, (CAS); Utah:
EMERY COUNTY: Chinle Formation, at jct of Buckhorn Wash & San
Rafael River Canyon north of river, 30 May 1970, Welsh, Atwood &
Moore 9819, (ARIZ, ASC, B, BRY, CAS, GH, NY, RM, RSA, UC, US, UT, UTC,
WSC); 2 mi s. of San Rafael River Bridge, 16 June 1969, D. Atwood
1843, (ARIZ, ASC, B, BRY, GH, NY, US, WSC) ; 11 mi n. of Goblin Valley
turn off on Temple Mountain road from US 24, thence 43 mi s.w. on
mining road, 17 June 1969, D. Atwood 1855, (BRY, RM); R.9E.,
T.26S., San Rafael Swell, 17 June 1969, D. Atwood 1847, (ARIZ,
456 Rhodora [Vol. 74
ASC, B, BRY, CAS, GH, NY, UC, US, UT, UTC); ca 13 mi e. of Buckhorn
Wash, north of San Rafael River, 30 May 1970, Welsh, Atwood &
Moore 9846, (BRY); ca 5 mi e. of Buckhorn Wash along road north
of San Rafael River, 30 May 1970, Welsh, Atwood & Moore 9844,
(ARIZ, B, BRY, CAS, GH, NY, US); ca 2 mi w. of San Rafael River
Bridge at jet. of Buckhorn Wash and San Rafael River, 1 June
1970, Welsh, Atwood & Moore 9903, (BRY, NY, US, WSC); WASHING-
TON COUNTY: 11 mi w. of Hurricane, 2 May 1942, F. Gould 1681,
(CAS, UC); near Virgin, 10 May 1941, Eastwood & Howell 9200,
(CAS); e. of Bloomington, 10 April 1970, Welsh, Atwood & Matthews
9535, (BRY, NY, UC, WSC); 2 mi e. of Hurricane, 27 May 1968,
D. Atwood 1409, (BRY, CAS); Rockville, 12 May 1941, Eastwood &
Howell 9327, (CAS, GH); 4 mi s.w. of Saint George, 27 May 1968,
D. Atwood 1417, (BRY); Moenkopi Formation at Bloomington, 3 May
1970, Welsh, Atwood & Matthews 9694, (BRY); clay hills n. of
Bloomington, 25 May 1969, D. Atwood 1698, (ARIZ, ASC, B, BRY, CAS,
GH, JEPS, NY, RM, UC, UTC, WSC); ca 5 mi s.w. of St. George, 7 April
1968, Higgins 1244, (BRY); WAYNE COUNTY: Capital Reef Wash
2 mi s.e. of Fruita, 6 May 1940, B. Maguire 18146, (CAS, NY, UC, UTC) ;
3 mi s. of Highway 24 on Notom Road, 17 June 1969, D. Atwood 1860,
(ASC, BRY, CAS, GH, RSA, UC, UT, US, WSC); Moenkopi Formation at
Fruita, 10 May 1969, Welsh Atwood & Higgins 8968, (BRY, CAS, RM,
RSA, UC, US, UT); 3 mi w. of Fruita, 4 June 1953, R. McVaugh, 14438,
(CAS, NY, UC).
Phacelia rafaelensis appears to be related to P. utahensis
but differs from that species as indicated in the diagnosis.
P. utahensis is endemic to the Arapien Shale Formation
in Sevier and San Pete County, Utah, whereas P. rafae-
lensis (Figure 2, Map 2) occurs from Emery County south
to Kane County, Utah and Mohave County, Arizona and
east to Washington County, Utah. It grows on clay hills.
of the Moenkopi Formation. mid April to mid June.
This species is named for the San Rafael Swell, a floristic-
ally unique area, in Emery County, Utah.
Phacelia howelliana Atwood sp. nov.
A. P. corrugata differt corollis pusillioribus et tubo albo,
seminibus diluti-marginatis dorsalibus laevibus, plantis
brevioribus et plerumque ramosis et frondosis basi.
Plantae annuae, 0.9-2.3 dm altae; caules pro parte max-
ima et frondosi ad basin versus, glandulifera et aliquot pili
longiores simplices praesenta ; folia e late oblonga ad ovales,
Phacelia — Atwood 457
1972]
Phacelia rafaelensis. Habit and seed of plant.
Fig. 2.
[Vol. 74
Rhodora
458
2”
=. = = o m mo ae m m aa a e e m ml
tS
Lj
r
1
I
l
/
——— ——————————————-—h co
m--——-—-—---—------2—----2--2-2-2--4----
eee eee ee
Southern Utah and northern Arizona. Range of P. rafae-
Map 2.
lensis.
1972] Phacelia — Atwood 459
2-6 cm longa, 1.0-2.5 cm lata, irregulariter crenata ad lobata,
strigosa et leviter glandulifera, petiolus usque ad 5.0 cm
longa; inflorescentia cymarum scorpiodorum compositorum,
pedicelli usque ad 2.0 mm longi; sepala linearia ad anguste
oblanceolata, 3.5-4.0 mm longa, 1.0-1.5 mm lata, glanduli-
fera et hirsuta; corolla 5-6 mm longa, 6-7 mm lata, rotata
ad infundibuliformis, lobi pallidi violacei, tubus albus,
stamina et stylus exsertus 3-4 mm, stylus breviores stamina
furcatus 34, longitudo pubescens infirmus !4; capsula ob-
longa ad subglobosa glandularis et hirsutulosa apprime
versus apex; semina 4, brunnea, 3.2-4.0 mm longa, 1.4-1.8
mm lata, elliptica, margines corrugati usque involuti ad
complanati, pagina ventrali foveolata, excavata et divisa
porca prominens, interdum curva lata una et vix corrugata,
pagina dorsalis prophyreus laevis et margo pallida circum-
tineta.
TYPE: Utah: SAN JUAN COUNTY: ca 0.4 mi n. of Bluff,
Utah on highway 163, in mouth of canyon on rocky-sandy-
clay soil, 13 May 1970, D. Atwood 2454, (HOLOTYPE: BRY;
ISOTYPES, ARIZ, ASC, B, BRY, CAS, GH, NY, RM, UC, US, UTC).
Additional materials examined: Utah: GRAND COUNTY: 13 mi n.
of Moab, 15 June 1944, A. H. Holmgren & S. Hansen 3319, (NY);
Moenkopi Formation, T.25S., R.23E., in Castle Valley, 5 June 1970,
Welsh, Atwood & Moore 9957, (ARIZ, BRY, CAS, WTS) ; Arches National
Monument, vicinity of Courthouse Wash, 2 May 1963, G. L. Pyrah,
et al. 71, (BRY); Potash Plant, 20 April 1962, S. L. Welsh, 1670A,
(BRY); SAN JUAN COUNTY: Bluff, near the Navajo Twins, 29 April
1961, D. Bright 55, (BRY); Bluff, 24 May 1919, M. E. Jones s.n.,
(POM); along road just w. of Monument Valley Hospital, 15 May
1970, D. Atwood 2511, (ARIZ, ASC, B, BRY, CAS, DIX, GH, JEPS, NY,
POM, RM, RSA, UC, US, UTC, WSC); Goulding Trading Post); 26
June 1948, J. T. Howell 24687, (CAS); canyon of the San Juan
River between Bluff and the Colorado River, P. Thompson s.n., May
1941, (CAS); dark canyon-Cataract Canyon, 1 July 1938, E. Clover
& M. Jotter 2143A, (GH); ca 2 mi e. of Bluff, 12 May 1944, A. H.
Holmgren 3187, (UTC).
This entity is related to P. corrugata A. Nels. which
ranges throughout most of Utah and extends into Colorado,
northern New Mexico and northern Arizona. It is distin-
guished from P. corrugata by its low much branched growth
460 Rhodora [Vol. 74
form, smaller corolla and white tube. The leaves are mostly
basal and the seeds are different as indicated in the diag-
nosis and general description. P. howelliana (Figure 3,
Map 3) is known only from San Juan and Grand Counties,
Utah where it grows in red sandy, gravelly or sandy-clay
soils at ca 4500 to 5000 feet. It probably occurs in Colo-
rado near Moab and undoubtedly in Monument Valley in
-
Fig. 3. Phacelia howelliana. Habit and seed of plant.
1972] Phacelia — Atwood 461
a ee ee ee —— — —
ee
o
Bluff
y —?e
Map 3. Southern Utah, Grand and San Juan Counties, Range of
P. howelliana.
462 Rhodora [Vol. 74
Arizona, although no specimens have been seen from either
area.
This species is named in honor of John Thomas Howell
at the California Academy of Sciences, who has published
many important works within the genus Phacelia.
Phacelia vossii Atwood sp. nov.
Plantae perennis, 2.1-7.3 dm altae; caules erecta e caudice
ligneo, hirsutis ad hirsutulis et stipitati-glanduliferis pilis;
folia lineara ad lanceolata, 2.0-11 cm lata, revoluta, pagina
ventralis glandulifera dense, pagina dorsalis hirsuta et
stipitate-glanduliferis dispersis, margines dentatae ad ir-
regulariter dentatae, petiolata, petiolus usque ad 1.5 cm
longa; inflorescentia terminalis vel secus caules dispersa
usque ad 14 longitudo vel, minus, simplex ad cymarum
scorpiodorum compositorum, racemi individua usque ad
7.5 cm longa per fructus, pedicelli usque ad 2.0 mm longa
per fructus, glandulifera et hirsuta; corolla tubularis
(campanulata ?), diluta purpurascens ad albida (?), 5.5
mm longa ; sepala oblanceolata ad spathulata, 4-6 mm longa,
1.5-2.0 mm lata, glandulifera et hirsuta; stamina exserta ;
stylus exserta circa 2.0 mm stamina longior, circa 9.0 mm
longa, furcatus 94 longitudo, inferna 14, pubescens; capsula
ovata, 3.7 mm longa, 2.5-2.7 mm lata, glandulifera et stri-
gosa; semina 4, elliptica ad oblonga, 2.5-3.1 mm longa, 1.0-
1.4 mm lata, margines integra ad corrugata parte, pagina
dorsalis foveolata, pagina ventralis foveolata et porca
prominentia centrale corrugata, porca excavata utrinque.
TYPE: Mexico: NUEVO LEON: Sierra Madre Oriental,
caleite and limestone hills beyond Pablillo toward Santa
Clara, about 15 mi s.w. of Galeana, sparse in arroyos,
flowers light purple, 18 July 1934, C. H. & M. T. Mueller
1075, (HOLOTYPE: GH; ISOTYPE, MEXU).
Additional materials examined: Mexico: NUEVO LEON: Hacienda
Pablillo, Galeana, 13 August 1936, Mary Taylor 156, (UC, NY).
Phacelia vossii (Figure 4, Map 4) is known only from
the type collections by Galeana in the State of Nuevo Leon,
Mexico. It apparently grows in calcite and limestone soils.
463
Phacelia — Atwood
1972]
A
Fig. 4. Phacelia vossii. Habit and seed of plant.
464 Rhodora [Vol. 74
COAHUILA ;^ \
p
EN /
^ €
} E
1 \
I t
{ )
J A
r ^
. `
\, oN "d
IN t "a " e
MU -^77N i
S
N
`
[
{
\
\ -
\ /
t l
p
-2
v
Map 4. Mexico, States of Coahuila and Nuevo Leon. Range of
P. vossit.
The flower color is not apparent in the specimens at hand,
but the label of the type material indicates that it is a light
purple. Other members of the crenulata group which pos-
sess a pale corolla are often also whitish at times.
This taxon appears to be related to P. pinnatifida Griseb.
According to Brand (1913) the latter occurs in the Andes
of South America in Peru, Boliva, and Argentina, from
3000 to 3800 meters. The specimens cited were identified
as P. integrifolia Torr., however P. vossti is easily distin-
1972] Phacelia — Atwood 465
guished from the former by its evident perennial habit, pale
more or less tubular corollas, smaller corrugated seeds and
more densely pubescent herbage.
This species is named in honor of John Voss who revised
the crenulata group in 1987.
Phacelia welshii Atwood sp. nov.
Plantae annuae, 1.0-4.5 dm altae; caules plus minusve
flavo-virentes, simplices vel ramosi frondosi, hirsuti et
dense tecti multi-cellularis stipitatis, glandibus; folia
oblonga ad lanceolata, 1.5-8.0 em longa, 0.5-2.7 cm lata,
hirsuta et dense glandulifera, margines undulatae crenatae
vel dentatae saepe revolutae, folia basales fasciculata,
petiolata, petiolus 2.0 cm longus or minus, folia caulina
sessiles vel fere sic, saepe cordata ad basim; inflorescentia
cymarum scorpiodorum compositorum, terminalis ad ex-
trema principalis caulis et rami lateralis, dense glandulifera
et hirsuta, racemi individui congesta, sed laxe per fructus,
usque ad 1.0 dm longa, pedicelli usque ad 1.5 mm longa;
sepala spat lata glandulifere; corolla campanulata, pur-
purascens ad violacea, 5.0-6.0 mm longa et lata, pubescentia ;
anthera lutea; stylus exserta circa 8.0 mm, furcatus 94
longitudo, inferna 1⁄4, hirsutulosa et glandulifera; semina 4,
oblonga, ventraliter brunnea ad prophyreus dorsale, foveo-
lata, 2.8-3.4 mm longa, 1.3-1.5 mm lata, pagina ventrali
pagina dorsalis pallidior quam et porcis prominentis, porca
corrugata lata una, margines corrugatae et plus minusve
revolutae.
TYPE: Arizona: COCONINO COUNTY: along highway 89
north of Gray Mountain, on red shale (also common south
of town on red shale), 19 May 1970, D. Atwood 2608, (HOLO-
TYPE: BRY; ISOTYPES, ARIZ, ASC, B, BRY, CAS, COLO, DIX, GH,
NY, POM, RM, RSA, UC, US, UT, UTC, WSC, WTS).
Additional] materials examined: Arizona: COCONINO COUNTY: 2.5
mi w. of highway 89-64 jct, red shale, 19 May 1970, D. Atwood 2611,
(ARIZ, ASC, B, BRY, CAS, COLO, DIX, GH, NY, UC, US, WSC, WTS); 10 mi
n. of Wuptaki National Monument jet with highway 89, 19 May
1970, D. Atwood 2605, (ARIZ, ASC, B, BRY, CAS, GH, NY, POM, RM RSA,
UC, US, UT, UTC, WTS) ; ca 1.5 mi n.w. of Headquarters of Wuptaki Na-
466
Fig. 5.
Phacelia welshii. Habit and seed of plant.
[Vol.
74
1972] Phacelia — Atwood 467
FLAGSTAFF
Map 5. Coconino County, Arizona. Range of P. welshii.
468 Rhodora [Vol. 74
tional Monument along highway on red shale, growing with Ephedra,
Hilaria, Oryzopsis, blue grama, Atriplex and apache plume, 18 May
1970, D. Atwood 2601, (ASC, BRY, CAS, US, WSC); rocky limestone
areas above the Little Colorado River, 6 May 1961, D. Demaree 44391,
(CAS, UC); red shale hills along road ca 0.5 mi n. of headquarters
of Wupatki National Monument, 18 May 1970, D. Atwood 2598,
(ARIZ, B, BRY, CAS, COLO, DIX, GH, NY); Willow Springs n. of Cameron,
7 June 1942, Ripley & Barneby 4875, (CAS) ; red shale 11 mi s. of
Cameron, 20 June 1948, Howell 24397, (CAS).
P. welshii (Figure 5, Map 5) is probably most closely
related to P. corrugata and P. utahensis. It can be distin-
guished from the former by its broader, dark brown seeds,
long (up to 1.3 mm long), flattened, multicellular, stipitate
glands and different leaves. Typically P. corrugata is a
more northern plant but is not uncommon in Coconino
County. The new taxon can be differentiated from P. uta-
hensis by its more open inflorescence, corrugated seeds,
and broader, shorter, densely glandular leaves. P. utahensis
is endemic to the Arapien Shale Formation in Utah.
This species is named in honor of Dr. Stanley L. Welsh,
Brigham Young University, for his rich contributions
toward the flora of the Intermountain area, and for his
encouragement and invaluable help throughout the course
of this study.
DEPT. OF BOTANY AND RANGE SCIENCE
BRIGHAM YOUNG UNIVERSITY
PROVO, UTAH 84601
REDISCOVERY OF TILIA NEGLECTA SPACH
HENRY K. SVENSON
Tilia neglecta has always been a questionable species of
Linden. It was described in 1834 from “ambulacris Hort.
Bot. Paris” and was stated to be of North American origin.
The range is from Montreal to Washington and southwest-
ward. Due to variability, and absence of a “type” with
which the name could be correlated, it has remained one
of the least tangible species in a taxonomically difficult
a ; delis y ee
Fig. 1. Four views of Tilia neglecta, Sandy Neck, Cape Cod,
Massachusetts.
469
470 Rhodora [Vol. 74
genus. In Rhodora 72: 339-344. 1970, I described a “forest”
of Tilia neglecta in a dune hollow on Cape Cod, Massachu-
setts (fig. 1), and reviewed its subspecific synonyms, but
at that time I had not seen the fine treatment of the Ameri-
can species by G. N. Jones in Illinois Biological Monographs
no. 39, 1968.
In the spring of 1970, on a visit to the herbarium of the
Jardin des Plantes I found in the European folders a speci-
men of T. neglecta labeled in Spach’s hand, dated 1834, and
from the “Menagerie”. Then, after making detailed
sketches of this specimen, I went over to the western en-
trance of the zoological garden. Just a short distance within
the gate and opposite the old viper pit, stood a gigantic
weatherbeaten linden, which in its leaves corresponded
exactly with the herbarium specimen, evidently the long-lost
Tilia neglecta. There were no records, but Dr. H. Heine
said that it was not a European linden, and suggested that
the tree originated from seeds sent by Michaux, which
would have produced flowers and fruit by 1834. From my
visit in 1937 I remember vividly the rocky viper pit. These
vipers are now in elegant quarters in the new reptile house,
and the rocks are given over to some harmless snakes. The
exhibit, I think, has suffered from wear, and there is a
sign in French (a universal language) “Please do not
throw stones at the snakes." At the opposite entrance of
the *Menagerie" there is a large unlabeled tree of T.
heterophylla of about the same vintage as T. neglecta.
In the summer of 1970 Dr. Heine made for me a fine
series of herbarium specimens of the Menagerie tree (see
fig. 2), showing foliage, flowers, and sprout growth. These
I have given to the Gray Herbarium. They confirm in
microscopic detail the Tilia neglecta specimen mentioned
above, and Dr. Heine has labeled this tree as the “holotype”.
Accompanying the specimen to be retained by the Gray
Herbarium, I have placed the detailed sketches I made of
the 1834 specimen.
André Michaux (1746-1802) ranged far in the eastern
United States. He had two stations for assembling seeds
1972] Tilia — Svenson 471
ee E e a EE EL
Fig. 2. Leaf (upper surface) and flower cluster from the type
tree of Tilia neglecta (Jardin des Plantes, Paris), X ca. 2/3 (actual
size of leaf 11.5 X 9.5 em).
and roots; one at Charleston, South Carolina, the other on
the east bank of the Hackensack River at New Durham in
New Jersey. An account of the New Jersey garden, which
covered 8 or 10 acres now occupied by the Hoboken Ceme-
tery, is given by Rusby in Bull. Torrey Club 11: 88-90.
1884. Zenkert in Flora of the Niagara Frontier Region,
1934 says that the younger Michaux (1770-1855), having
on his first expedition (1801-1803) traveled in the terri-
tory west of the Alleghenies, during the three years of his
second journey explored the Atlantic Coastal States from
Maine to Georgia and made five trips to the interior, one
472 Rhodora [Vol. 74
of which, undertaken in 1806 or 1807, took him from New
York to Lakes Ontario and Erie. The Michaux Herbarium
at Paris consists of selected plants to illustrate the “Flora
boreali-americana" published in 1803.
The Paris herbarium is also well represented by other
early collectors. In addition to the Michaux Herbarium,
there are four folders of American lindens, in which there
are many Michaux specimens. I saw in the third folder
the specimen collected by W. Riehl in Missouri, labeled as
T. neglecta by Spach. It was sent to Jones and correctly
annotated as T. americana. Among the early collectors are
Tuckerman*, Pearson, Cooper, Torrey, Oakes, and others.
In the European folders are three additional sheets of T.
neglecta, so named by Spach “fleurs 12 juillet, 1850; fr.
Sept. 1851; cultivée dans les promenades de Carlsruhe
(grand duché de Baden)". There are also the varieties of
Tilia published by A. Braun. Both the American and Euro-
pean collections would be well worth going over. For help
at the Jardin des Plantes I am also greatly indebted to
the Director, Dr. Jean Jacques Leroy, and to Dr. Alicia
Lourteig who has charge of the Michaux Herbarium.
To return to a survey of American Literature on the
subject, we find that Hough, Handbook of Trees... (1907,
and reprint 1947), says that T. Michauxii is similar to
T. heterophylla, and also confounded with T. pubescens
*Edward Tuckerman (1817-1886) a graduate of Union College in
Schenectady, was the first, and I think the only curator of their her-
barium. In his fine classification of the genus Carex, published in
Schenectady and dedicated to John Torrey, he mentions himself as
«Mus. et Herb. Concord Cur.". While I was teaching at Union,
I came upon a trunk or two filled with Tuckerman specimens, and
wrote an account of Tuckerman in the Union Alumni Bulletin
(about 1924). The plants consisted primarily of collections by Jona-
than Pearson and Lewis Beck from the Schenectady region; plants
collected by Tuckerman in Europe, chiefly in the Epping Forest of
England; and a large series of European plants given to him by
Grisebach. Of Tuckerman's Carices and other interesting plants of
eastern America, I found none, and suppose that he took these col-
lections with him on leaving Union. I understand that the Union
College Herbarium is now dismantled.
1972] Tilia — Svenson 473
Aiton, and not well understood. T. neglecta is not treated
by Deam in Flora of Indiana, but L. Braun in The Woody
Plants of Ohio (1961) publishes a map, and considers it
“a wide-ranging species or species-complex, perhaps inter-
mediate between 7. americana and T. heterophylla, but not
synonymous with either.” House in Annotated list of the
ferns and flowering plants of New York State, 1924, men-
tions T. neglecta as frequent in central New York; the
lower Hudson River Valley, Long Island, and Staten Island.
Wiegand and Eames in Flora of the Lake Cayuga Basin,
New York, 1926, always careful in their judgments, note
that “the pubescence on the lower surface of the leaf is
highly variable, and on it several species have been founded.
The pubescence, though always close, fluctuates very gradu-
ally in density, and no other structural characters occur.
It seems better, therefore, to recognize this closely pubescent
form as simply a variety, T. americana var. heterophylla
Loudon. Sargent’s T. neglecta is transitional to typical
T. americana’. Jones’ detailed treatment recognizes only
three species in eastern United States: T. americana, T.
heterophylla, and T. caroliniana. Sargent, in Bot. Gaz. 66:
494-496. 1918 states “I now consider T. neglecta to have
been the T. Michauxii of Nuttall, which is the T. argentea
of Michaux . . . The pubescence on the lower surface of
the leaves is so constant and so persistent throughout the
season, it seems best to consider it a species rather than a
pubescent form of T. glabra. The base of the style in T.
neglecta is furnished with long hairs, and that of T. glabra
appears to be quite glabrous".
In the Paris tree the petals are 7.5 X 3 mm with nar-
rowed base; the staminodia about 4.0-4.5 X 1.0 mm. In a
microscopic comparison of the Cape Cod and Paris ma-
terial, we see that the calyx is of the same shape, but in
the Paris specimen the pubescence is not dense. Pedicels.
are thin and elongate, about 10 X 0.5 mm (see fig. 2) in
the Paris material; but in the Cape Cod specimens densely
stellate and about 4-5 X 0.7 mm. For pubescence of the
under surface of the leaf, I have gone over the Cape Cod
474 Rhodora [Vol. 74
material more carefully. Under a magnification of 40X,
boiled fragments 2 mm square were cut with a razor blade
directly on the 0.5 mm grid eyepiece. Simple hairs could
then be readily counted for 1 sq. mm. The count of hairs
or trichomes from an area, not dense, for 5 mm. square of
lower leaf surface: 4-pronged 15-38 (10-25%) ; 3-pronged
1-6 (1-5%); 2-pronged (Malpighian) 6-10 (1%); 6-8-
pronged (none); simple 100-150 (90%). This analysis
(cf. the footnote on p. 343 of Rhodora, vol. 72) shows that
there is a considerable amount of variation.
For the Paris tree (in several counts) there was an
average of about 375 hairs on 5X5 mm of lower leaf sur-
face: 4-pronged (5%); 3-pronged (1%); 2-pronged
(396) ; 5-pronged (less than 1%); simple (90%). In this
Paris material the 2-pronged hairs are merely bifurcate;
the simple hairs are mainly set at an angle on the veins,
giving the veins a fuzzy appearance, whereas in the Cape
Cod material the larger veins are practically glabrous.
As an exeuse for this statistical treatment, I may cite
the statement of Jones (p. 110) that much of the practical
identification and classification of Tilia must be made on
the amount and kind of pubescence, particularly on the
underside of the leaves. Probably this statistical method
is not the best one, but cutting and counting is not as diffi-
cult as it might seem. Perhaps the pattern of veining of
the upper surface of the leaf would be more adaptable.
At any rate, I have begun at the beginning, as Fernald
would say, and the different opinions of capable botanists
and the variations observed by them in these leaf struc-
tures, shows how difficult the subject is. I have accumu-
lated notes and sketches of several specimens in the Paris
Herbarium, and I have gone over casually the specimens
of T. neglecta in the Gray Herbarium. I do not have the
inclination to pursue the subject further. — Osterville,
Mass. 02655.
THE ECOLOGY OF BENTHIC SALT MARSH ALGAE
AT
IPSWICH, MASSACHUSETTS
I. ZONATION AND DISTRIBUTON
OF ALGAL SPECIES
E. E. WEBBER AND R. T. WILCE
In a previous publication (Webber & Wilce, 1971) we
treated the salt marsh algae at Ipswich in light of their oc-
currence, morphology, cytology, seasonal and reproductive
periodicities, based on both field and laboratory investiga-
tions. In this paper the descriptive ecology of these plants
is discussed.
The ecological nomenclature used to describe vertical and
horizontal distributions of algae has varied considerably.
Rather than adopt new terms describing the zoned character
of the salt marsh algal vegetation, we have selected estab-
lished terminology common to terrestrial ecological studies
which applies also to flowering plants of the marsh environ-
ment (Weaver & Clements, 1938; Feldmann, 1951; Oosting,
1956; den Hartog, 1959). Accordingly, we have attempted
to describe both the physiognamy and the vegetationa] units
at each station, the latter in relation to their successional
character.
Superficially, the zoned arrangement of an algal salt
marsh vegetation often is not easily detected. The flat ex-
panse of varying shades of green which cover the salt marsh
surface contrasts sharply with the distinct and frequently
colorful patterns of algal zonation of the rocky coast (see
Frontispiece, Lewis, 1964). In order to discuss algal zona-
tion and distribution at the Ipswich salt marsh, it was
necessary to identify levels on the marsh surface with which
to relate the presence and vertical range of algal species.
Marsh phanerogams, owing to their abundance, distinctive-
ness, and regular and sharp zonation, served as reference
points to this end. From the seaward edge of the marsh
extending landward, the following seed plant associations
475
476 Rhodora [Vol. 74
were recognized: Spartina alterniflora var. glabra, Spar-
tina patens, Spartina patens-Scirpus americanus (or Spar-
tina patens-Distichlis spicata) mixture, and Juncus gerardi.
Algal zonation in the salt marsh becomes more apparent
and more readily understood when studied in relation to
the distribution of these phanerogams.
As stressed by Lewis (1964) and suggested earlier by
the Stephensons (1949), a discussion of zonation along any
shore should be based upon biological criteria, rather than
relying strictly upon tidal limits. These limits may not be
coincident with natural biological zones, especially when
one considers the upper limits of vegetation in relation to
tide levels. On shores exposed to wave action and spray,
the distribution of those organisms which typically form
the uppermost biological zones will be extended even higher
than where a splash and spray zone are not present (c.f.,
Lewis, 1964, fig. 52). In the salt marsh environment at
Ipswich, one encounters an approximation of a spray zone
only near Station 1 (Webber & Wilce, 1971, p. 265) ; here,
the presence of wood pilings provides a steep vertical face
which contrasts with the gently sloping character of the
surrounding marsh surfaces. This change in topography
results in a stronger local wave splash, as compared with
wave action on the marsh surfaces; thus, the upper dis-
tributional limit of Calothrix crustacea, for example, oc-
curs at a higher level on these pilings than on the surround-
ing marsh. Therefore, using essentially biological criteria
in relation to tide levels we recognize in the marsh environ-
ment a sublittoral, a littoral, and a less well defined, but
nonetheless recognizable, supralittoral zone.
The sublittoral zone of the Ipswich salt marsh, when
compared with the same zone characteristic of the open
coast, is essentially non-existent. The marsh drains twice
daily through a series of canals which, at low tide level,
are mostly devoid of water. Where the marsh is closest to
the sea and in the major trunk canals, water with salinity
characteristics of open coastal water remains at low tide.
It is in these salt marsh areas, the openings to the sea and
1972] Benthic Algae — Webber and Wilce ATT
the major canals, where a true sublittoral zone can be rec-
ognized, and, correspondingly, a sublittora] flora.
We stress the fully marine character of the sublittoral
benthic vegetation. Also, in our list of species from the
Ipswich salt marsh (Webber & Wilce, 1971), we cite a num-
ber of taxa commonly represented in the marsh sublittoral
as drifting plants, e.g., Chaetomorpha melagonium and
Phycodrys rubens, to name but two; these plants, of vari-
able longevity in the marsh environment, are transients
from the open coast sublittoral. Their frequent appearance
in the marsh sublittoral in conjunction with the attached
marsh sublittoral vegetation is a manifestation both of the
marine character of the marsh subtidal zone and the prox-
imity of the marsh entrance to the open coast.
The littoral zone at the Ipswich marsh contains those
crganisms which are regularly submerged and exposed by
the rise and fall of the tides. Biologically, this zone ex-
tends from the upper limit of the sublittoral, as indicated
by Fucus distichus ssp. evanescens, to that uppermost por-
tion on the shore populated by Juncus gerardi. Algae of
the littoral zone show a zoned pattern of distribution as con-
trolled by local factors of, e.g., habitat competition, avail-
ability of substrates, and topographic differences resulting
in degree of exposure to insolation, desiccation, tidal action,
and fresh water influence. These algal assemblages of pre-
dictable major constituents we interpret as associations,
folowing Borgesen, 1905, Davis, 1913, Feldmann, 1951,
Wi!ce, 1959, den Hartog, 1959 and Jorde Klavestad, 1963.
However, as Chapman (1956) points out, one characteristic
of salt marsh vegetation is its relative instability; thus, the
term “associes” is a more apt designation of the vegeta-
tional status in these specific environments.
The supralittoral zone is characterized by those species
which are predominantly terrestrial and which are not
merely extensions of upper littoral marine species. The or-
ganisms which are regularly submerged and exposed by the
tide are in fact, considered “maritime” (Lewis, 1964) rather
than strictly marine. While a comparatively clear-cut dis-
478 Rhodora [Vol. 74
tinction between those plants and animals of the upper lit-
toral and those of the supralittoral may be possible on a
rocky coast, this distinction is not always apparent at the
Ipswich salt marsh. For example, Calothrix crustacea clear-
ly occurs on the marsh surface (Station 1) in the upper lit-
toral zone. In addition, this species along with the upper
littoral green alga Pseudendoclonium submarinum, colonize
wood pilings near Station 1. Owing to moderate wave
splash on these pilings, both algal species extend vertically
higher than the level of the surrounding marsh surface.
Yet, at Ipswich, the position of C. crustacea and P. sub-
marinum on the pilings near Station 1 is comparable to the
position on the marsh surface populated by such terrestrial
or “maritime” plants as Juncus balticus var. littoralis, Pani-
cum virgatum, and Solidago sempervirens. Indeed, on the
pilings with Calothrix and Pseudendoclonium one always
finds the lichen Lecanora chlarotera which, according to
Hale (personal communication), is not marine in its dis-
tribution, but rather is a common terrestrial species. Thus,
the vegetation at this level is dominantly maritime-terres-
trial, and does not represent solely an upward extension of
the marine flora. We recognize, therefore, a supra-littoral
zone biologically distinct in its composition of vascular
plants and associated cryptogams which lies invariably im-
mediately above the uppermost limit of salt marsh phanero-
gams, i.e., above Juncus gerardi.
ZONATION AND DISTRIBUTION OF ALGAL SPECIES
Before describing the species composition at each sta-
tion, it is appropriate to identify and describe briefly the
stations from which the algal collections and related data
were taken (see also Webber & Wilce, 1971).
Station 1: typically marine, little influenced by fresh
water runoff; shoreline a vertical mud bank with substra-
tum consisting of mud, small stones, and the Spartina
grasses; seasonal salinity range of 18-30°/o0.
Station 2: Similar to Sta. 1, but with a gently sloping
1972] Benthic Algae — Webber and Wilce 479
shoreline; substrates predominantly cobble, shells of Midio-
lus demissa and Spartina grasses.
Station 3: Tidal ditch outflow, 0.3-1 m deep, seasonally
influenced by fresh water runoff; substrates of small stones
and wood pilings; seasonal salinity range 3-33? /oo.
Station 4: Section of a major tidal zreek beneath a
highway bridge panning Fox Creek, moderately influenced
by fresh water runoff; substrate mostly of stone and shell;
seasonal salinity range 13-329/00.
Station 5: The innermost extremity of the Ipswich salt
marsh, consisting of two substations due to the habitat
character in the tidal creek in this region of the marsh;
seasonal salinity range for both substations 0-27°/00.
Substation 5a: A transect through the tidal creek near
Northgate Rd. where at high water the bottom is cov-
ered to a depth of 0.6 m, and mostly exposed at low
water; markedly influenced by fresh water runoff ; sub-
strates of small stones, mud, and plants of Ruppia
maritima; this area is characterized also by a twice
daily rapid flow of water.
Substation 5b: Differs from the latter by having a
mud bottom, continual water cover at a 0.6 m depth,
and a generally slow water displacement downstream.
Sublittoral zone:
Stations 1 and 2. Species common to the sublittoral of
these stations were not unlike those one might encounter in
the sublittoral on a rocky coast. These algae included:
Enteromorpha intestinalis Enteromorpha flexuosa
Enteromorpha linza ssp. pilifera
Enteromorpha linza Kornmannia leptoderma
var. oblanceolata Ulva gigantea
Enteromorpha ahleriana Chaetomorpha linum
Enteromorpha clathrata Ralfsia verrucosa
Enteromorpha flexuosa Ralfsia clavata
ssp. flexuosa Scytosiphon lomentaria
480 Rhodora [Vol. 74
Petalonia fascia
Chorda filum
Laminaria saccharina
Dumontia incrassata
Hildenbrandia prototypus
Agardhiella tenera
Gracilaria foliifera
Gracilaria verrucosa
Chondrus crispus
Ce:amium diaphanum
Ceramium rubrum
Ceramium rubriforme
Polysiphonia denudata
Polysiphonia lanosa
Polysiphonia nigra
Polysiphonia nigrescens
The majority of these species were of late spring and
summer occurrence, while the following attained maxi-
mum vegetative and reproductive development during the
winter and early spring months:
Enteromorpha linza
var. oblanceolata
Kornmannia leptoderma
Ralfsia clavata
Scytosiphon lomentaria
Petalonia fascia
Laminaria saccharina
Dumontia incrassata
Ralfsia verrucosa
Conspicuous perennials at the sublittoral-littoral inter-
face were Fucus distichus ssp. evanescens, Fucus vesiculo-
sus and Chondrus crispus.
Station 3: There was a decrease in numbers of sublit-
toral species at Station 3 as compared with Stations 1 and
2. This reduction in taxa is due likely to the comparative
lack of rocky substrate, the pronounced fresh water in-
fluence in the winter and spring months, and the shallow
water at this station. Those algae common to the sublit-
toral of Station 3 throughout the summer were:
Enteromorpha intestinalis
Enteromorpha flexuosa
ssp. pilifera
Ulva rigida
Cladophora liniformis
Bryopsis plumosa
Ceramium diaphanum
Ceramium fastigiatum
Polysiphonia denudata
The winter and early spring algal vegetation was dom-
inated by four species:
Kornmannia leptoderma
Scytosiphon lomentaria
Petalonia fascia
Melosira juergensii
1972] Benthic Algae — Webber and Wilce 481
The only sublittoral perennials in this station were Ralf-
sia clavata, Ralfsia verrucosa, and Chondrus crispus.
Station 4. The submerged algal vegetation here was
scant. From November through March Kornmannia lepto-
derma, Pentalonia fascia, and Dumontia incrassata com-
prised the macroscopic algae at this station in the sublit-
toral zone. These taxa were continually vegetative and
greatly reduced in size, as compared with their larger and
normally reproductive counterparts at Stations 1-3.
The sublittoral perennials were the same as those at
Station 3, i.e., Ralfsia clavata, Ralfsia verrucosa, and Chon-
drus crispus.
Station 5,a and b. The vegetation in the sublittoral here
had the smallest number of algal species of any station.
We have considered this to be a reflection of the reduced
salinities occurring here through most of the year. During
the spring and autumn, the following species dominated:
Capsosiphon fulvescens Ectocarpus confervoides
Enteromorpha intestinalis var. dasycarpa
Polysiphonia urceolata
In early spring only the stalked diatom, Gomphonema
olivaceum, constituted the macroscopic algal vegetation,
covering the numerous small stones in this tidal creek.
Owing to a thick ice cover, samples of winter algae were
not obtained from this station.
Littora! Zone
Stations 1 and 2. Fucus vesiculosus and Ascophyllum
nodosum occurred in the littoral zone wherever stone, rock,
or other solid substrate was present. The landward extent
of these species differed, however, between Stations 1 and
2. For example, at Station 1 the marsh surface is essen-
tially 1 meter higher above high water neap tide levels
than is that of Station 2, owing to the presence of a ver-
tical mud bank at the former station. Fucus vesiculosus
became established on this bank, the plants extending
about 3/4 of the height of this vertical surface. At this
482 Rhodora [Vol. 74
uppermost level, Fucus thalli tended toward a spiral growth
habit, and they were smaller than those plants in the mid-
littoral. By contrast, the lower level and more sloping top-
ography of Station 2 resulted in both Fucus and Ascophyl-
lum extending onto the marsh surface, where they devel-
oped as typical marsh fucoids, i.e, F. vesiculosus var.
spiralis and A. nodosum f. scorpioides, both at the bases
of Spartina alterniflora var. glabra.
Immediately beneath the seaward marsh edge, a dark
green algal zone, the Enteromorpha associes (Fig. 1), was
conspicuous on the mud at both stations. The summer and
autumn algae comprising this associes were:
Enteromorpha ahlneriana Cladophora sericea
Enteromorpha flexuosa Rhizoclonium riparium
ssp. paradoxa f. riparium
Enteromorpha prolifera f. validum
Percursaria percursa
Throughout the winter and spring months this Entero-
morpha associes assumed a pronounced brown hue owing
to increased numbers of Pylaiella littoralis, with lesser
amounts of Ectocarpus confervoides var. arcta. Associated
with these plants at this time were irregularly distributed
clusters of:
Enteromorpha flexuosa Percursaria percursa
ssp. paradoxa Vaucheria intermedia
Monostroma oxyspermum
Covering the mud in the Spartinetum alternifloretum at
Station 1 were conspicuous green tufts of Rhizoclonium
riparium (f. riparium and f. polyrhizum), Rhizoclonium
implexum, and Vaucheria intermedia.
While these algae were similarly located at Station 2,
their distribution at the latter station was limited to the
more seaward edge of the Spartinetum. We attribute this
local restriction in vertical distribution to competition for
habitat; marsh fucoids and the ribbed mussel, Modiolus
demissa, thoroughly blanketed much of the available mud
substrate beneath Spartina alterniflora var. glabra, thus
Benthic Algae — Webber and Wilce 483
1972]
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484 Rhodora [Vol. 74
prohibiting any further upshore spread of Rhizoclonium
riparium, R. implexum, and Vaucheria intermediwm. A
variety of filamentous bluegreens was collected throughout
the year from the mud of this Spartinetum at both Sta-
tions 1 and 2 (Webber, 1967).
Moving landward, the next algal associes cecurred be-
neath Spartina patens and consisted almost entirely of
Vaucheria arcassonensis (Webber, 1968). Growth of this
alga, with lesser amounts of V. intermedia, was particularly
evident during the summer and autumn months, forming
dense, felt-like mats on the soil. Numerous filamentous
bluegreens also were common at this time on the sandy
peat soil of the Spartinetum patentis.
Old culms of Spartina patens were epiphytized through-
out the year mainly by Calothrix confervicola, as tufts to
3 mm tall Often entangled with C. confervicola were
plants of Microcoleus tenerrimus ; the latter species never
occurred alone as a dominant. Blum (1968), working on
several Cape Cod (Massachusetts) marshes, concluded that
a species of Calothrix (likely C. confervicola) is the most
important colonizer in the moist micro-habitats on dead
S. patens culms. He also demonstrated a marked interde-
pendence between specific salt marsh grasses and their un-
derlying algal species. Blum further elucidated the rela-
tionship between the mature form of the graminoid zone
and such environmental parameters as light penetration
and intensity in the zone, filtration and detritus retention,
and drainage, as all these events are influenced by the
morphology of a specific stand of grass.
The brownish-green tubular thalli of Capsosiphon fulves-
cens were present also in the Ipswich Spartinetum paten-
tis. While mocroscopic individuals of this species were
epilithic in the sublittoral at Substation 5a (Webber &
Wilce, 1971, p. 269), the macroscopic thalli of Capsosiphon
(the type commonly encountered elsewhere) grew only in
a single, small, water-retaining marsh depression at Sía-
tion 1. The discovery of two populations of Capsosiphon,
distinctly different in their morphologies, seasonality, and
1972] Benthic Algae — Webber and Wilce 485.
habitat requirements, suggests further questions as to the
character of the species and the role of the environment
in determining the expression of that character. Labora-
tory and field experiments designed to elucidate this mat-
ter are forthcoming.
The highest band of salt marsh phanerogams at Stations
] and 2 was the Juncus gerardi association; the bases of
Juncus and its associated algal species were wetted by sea
water only during periods of spring tides. The algal veg-
etation on the sandy soil here was predominantly Tolypoth-
rix tenuis and Rhizoclonium riparium f. validum. On patches
of soil not colonized by Tolypothrix and Rhizoclonium,
Calothrix crustacea formed brownish-blue masses through-
out the year, being especially abundant during the autumn,
winter, and spring months.
Station 3. The littoral zone of Station 3 consisted of
portions of the marsh surface and several wood pilings lo-
cated in the outflow of the tidal ditch at this station.
Bright green patches of Pseudendoclonium submarinum
were always apparent, occurring essentially in the top 20
cm of these pilings throughout the year. Monostroma oxy-
spermum occurred just beneath P. submarinum on the pil-
ings’ surfaces; this species first appeared in September as
small foliose clusters, and developed abundantly through-
out the winter until late April, after which it was no longer
apparent at this station. The lowermost extent of M. oxy-
spermum coincided with low water neap tide levels. Ulo-
thrix subflaccida, mixed with Monostroma, was first evident
in December, and most common during the spring months.
Similar to M. oxyspermum, U. subflaccida was absent dur-
ing the summer (July through September).
The algal vegetation of the marsh littoral at Station 3
consisted of few species; these were Vaucheria intermedia,
Rhizoclonium implexum, Percursaria percursa, and a mix-
ture of filamentous bluegreens. It was quite similar in its
species composition to the associes of comparable zonation
at Stations 1 and 2, being essentially a telescoped version
of the same zone at these stations.
486 Rhodora [Vol. 74
Station 4. Wooden bridge supports over Fox Creek and
the small stones and larger rocks in the creek bed imme-
diately beneath the bridge were substrates in the littoral
zone of this station. A scant algal vegetation characterized
the littoral zone here, with only Pseudendoclonium sub-
marinum and a few filaments of Calothrix crustacea appar-
ent on the bridge supports through the year. Ralfsia clav-
ata and R. verrucosa were yearly inhabitants also, as epi-
lithic crusts in the most damp and most shaded areas of
the creek bed along with large quantities of the barnacle,
Balanus balanoides. The creek bed and that portion of the
littoral zone with Pseudendoclonium and Calothrix could
also be characterized as the Balanus zone. Balanus was
present on virtually all solid substratum at this station,
and was the dominant competitor for substrate coloniza-
tion. Perhaps this is the reason for so few species of at-
tached algae here. In other upper shore locations one
usually finds numerous algal species in the Balanus zone.
While the obvious factors contributing to this associes of
so few species appear to be competition for substrate, ex-
posure to insolation and to desiccation, the specific ecologi-
eal factors responsible for this causal relationship are yet
to be discovered.
Station 5, a and b. The littoral zone of Station 5 con-
sisted of muddy tidal creek banks which contained an algal
flora throughout the spring, summer, and autumn months
of Vaucheria compacta var. koksoakensis mixed with fila-
mentous bluegreens.
Owing to an uneveness in the creek bed coupled with an
irregular accumulation of variously sized stones, a segment
of Substation 5a drained completely at low tide. Thus, a
small and well-defined littoral region was present in the
bed itself. Small stones here were covered with attached
plants of Monostroma oxyspermum from September
through December. This species also was encountered at
Stations 1-3 from late October through the winter, per-
sisting until June. Conover (1958) reported a similar au-
tumn-winter (October to January) reoccurrence of M.
1972] Benthic Algae — Webber and Wilce 487
oxyspermum in addition to its spring (March to June)
growth in Great Pond Estuary, Massachusetts. His data
shows comparable spring and autumn light intensities, and
he suggested this factor as responsible for his observed
pattern of seasonal distribution. While Conover’s hypothe-
sis might well be involved in an explanation of the pres-
ence of M. oxyspermum in Substation 5a at Ipswich, our
field data indicate that salinity levels may also affect the
seasonal appearance of these plants. Salinities at Substa-
tion 5a during the September to December period ranged
from 25-27?/oo, a variation similar to that, 24-309/oo,
measured at Stations 1-3 during the period of maximum
spring growth of Monostroma oxyspermum. Thus, the
seasonal patterns of occurrence of this species at Ipswich
coincide with low light intensities and high salinity levels.
Winter collections were not possible at Station 5 be-
cause of a thick ice and snow cover.
Supralittoral zone.
This zone was recognized only in conjunction with the
wood pilings near Station 1. The algal vegetation which
occurred in and on these pilings was uniform in species
composition throughout the year, and consisted of:
Pseudendoclonium Urococcus foslieanus
submarinum Branched filamentous Chry-
Calothrix crustacea sophyte undescribed
Schizothrix calcicola
SUMMARY COMMENTS
In this paper we have attempted to summarize our de-
scriptive ecological data in an overview interpretation of
the ecology of the salt marsh algae at Ipswich. The algal
taxa are characterized relative to their horizontal and ver-
tical zonation on the marsh, and the seasonality of the
dominant species is described.
LITERATURE CITED
BLUM, J. L. 1968. Salt marsh Spartinas and associated algae.
Ecol. Monogr. 38: 199-221.
BQRGESEN, F. 1905. The algal-vegetation of the Faeróes. coasts,
488 Rhodora [Vol. 74
with remarks on the phytogeography. In: Warming, Botany of
the Faróes. 2: 679-845, pls. 18-24.
CHAPMAN, V. J. 1956. The marine algae of New Zealand. J. Linn.
Soc. Lond. (Bot.) 55: 333-501.
Conover, J. T. 1958. Seasonal growth of benthic marine plants
as related to environmental factors in an estuary. Inst, Mar.
Sci, Univ. of Texas. Port Aransas. 5: 97-197.
Davis, B. M. 1913. General characteristics of the algal vegetation
of Buzzards Bay and Vineyard Sound in the vicinity of Woods
Hole. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fish. 31: 443-
544. (1911).
DEN HarTOG, C. 1959. The epilithic algal communities occurring
along the coast of the Netherlands. Wentia. I: 1-241.
FELDMANN, J. 1951. Ecology of marine algae. 313-333. In: G. M.
Smith (ed.), Manual of Phycology. Chronica Botanica, Wal-
tham, Mass.
HALE, M. (Personal communication. Smithsonian Institute. Wash-
ington, D.C.).
JogpE, L, AND N. KravEsTAD. 1963. The natural history of the
Hardangerfjord. 4. The benthonic algal vegetation. Sarsia 9:
1-99.
Lewis, J. R. 1964. The Ecology of Rocky Shores. The English
Universities Press, Ltd. London. 323 pp.
OosTING, H. J. 1956. The Study of Plant Communities. 2nd ed.
W. H. Freeman Co. San Francisco. 440 pp.
STEPHENSON, T. A., and A. STEPHENSON. 1949. The universal fea-
tures of zonation between tide marks on rocky coasts. J. Ecol.
37: 289-305.
Weaver, J. E., and F. E. CLEMENTS. 1938. Plant Ecology. 2nd ed.
McGraw-Hill Book Co., Inc. N. Y. 601 pp.
WEBBER, E. E. 1967. Bluegreen algae from a Massachusetts salt
marsh. Bull. Tor. Bot. Club, 94: 99-106.
1968. New England salt marsh Vaucherieae. Rho-
dora. 70: 274-277.
, AND R. T. Witce. 1971. Benthic salt marsh algae
at Ipswich, Massachusetts. Ibid. 73: 262-291.
WincE, R. T. 1959. The marine algae of the Labrador Peninsula.
and northwest Newfoundland (ecology and distribution). Bull..
Nat. Mus. Can. 158, iv + 103.
DEPT. OF BIOLOGY
KEUKA COLLEGE
KEUKA PARK, NEW YORK 14478
and
DEPT. OF BOTANY
UNIVERSITY OF MASSACHUSETTS
AMHERST, MASSACHUSETTS 01002
LUPINUS MEXICANUS CERV. EX. LAG.
Davip B. DUNN
Since Lupinus mexicanus Cerv. ex. Lag. (1816) was the
first taxon in the genus Lupinus named for Mexico it is
imperative to determine which lupine the name was applied
to. As the earliest name it would take priority over all
the later names. In all of Charles Piper Smith’s works on
the lupines of Mexico he literally ignored the name. In
my own studies it has been a perplexing problem for quite
a number of years. The curator of the herbarium at
Madrid, Spain, searched, but was unable to locate a speci-
men of Lupinus mexicanus of any vintage. Since seeds
were sent to England by Lagasca, and the plants grown
were utilized to prepare a much more detailed and accurate
description for Edwards Botanical Register (no. 457, in
1820) than that originally provided by Lagasca, Dr. Walters
at Cambridge, England, also searched the herbarium there
but was unable to find a specimen of the material used in
either of the original descriptions provided by Lagasca and
Edwards. In 1832, Maund was attempting to identify
Lupinus mexicanus and stated that the species had been
lost to science, but that he thought that it had been re-
gained. He supplied an illustration as had Edwards.
Edwards suggested that L. mexicanus was probably a
biennial and that it had flowered in the “stove” (glass-
house) in February, which suggests that they had planted
the seeds the previous season, but flowering had not oc-
curred until February. Maund’s plant material is indicated
as a perennial, which was probably another taxon, and
nothing is given which helps identify L. mexicanus.
The curators of the herbaria of Kew, Paris, Berlin,
Zurich and others searched but were unable to find any
specimens of the period involved. Since there is apparently
no type specimen and since the description of Lagasca could
apply to several taxa it is necessary to select some element
to fix the application of the name until such time as an
489
490 Rhodora [Vol. 74
authentic specimen may be discovered. The illustration in
Edwards Botanical Register (no. 457, 1820) serves this
purpose and I select it to serve in lieu of a type specimen
for Lupinus mexicanus Cerv. ex Lag. It should also be
pointed out that there is no accurately labeled material of
Lupinus mexicanus in any of the herbaria and the material
distributed by Pringle as L. mexicanus has been a major
source of error, since he applied the name to, at best, a
variant of L. aschenbornii, which is an alpine perennial.
During the preparation of the treatment of Lupinus for
the ‘Flora de Valle de Mexico” (for Dr. Rzedowski) the
lupines of the area have been studied in considerable detail.
Several field trips to the area have been conducted and
numerous collections made. Among the collections are
plants collected along the toll road between Mexico City
and Queretaro, the latter, the site of the palace of the
Emperor Maximilian. This route is probably not far from
the original route the Spaniards traveled, since it is on the
most direct route between the two early cities. In the area
near Tula, a number of collections of lupines were made,
which closely match the detailed description in Edwards
Botanical Register. 'The collection made in July by Giles
Waines contained both flowering and fruiting specimens.
Plants collected along the toll road a few miles northwest
of the Tula turnoff by both Harmon and Dunn on three
different years in late December were in full fruit, as
well as having a few branches in flower. Seeds from both
the Waines collection and the Dunn collection were grown
in the research greenhouse at the University of Missouri.
What had been assumed to be the same taxon turned out
to be two different taxa of the same complex — one an
annual flowering in the fall and early winter and the second
a biennial germinating at the same time as the annual but
requiring cold treatment during the winter before flowering
the following season. By planting the seeds of the annual
in January and the biennial in the early fall of the previous
year the two were brought into flower at the same time.
Attempts to self-pollinate the biennial all failed, so the
1972] Lupinus — Dunn 491
taxon is obligate outcrossing. Most of the attempts to cross
the annual and the biennial failed but a few produced some
seeds. Thus there are two distinct breeding populations
isolated primarily by their phenology, which remain dis-
tinct in nature, but which have not fully achieved inter-
sterility, which is not uncommon in plants. After becoming
aware of this difference, the colonies along the toll road
were examined and found to contain many plants with only
caespitose clusters of basal leaves with no flowering shoots,
in late December, while the annuals were in fruit with the
lateral branches still in flower. The biennial is Lupinus
mexicanus and the annual is Lupinus bilineatus Benth.
The following translations of the Latin descriptions of
both Lagasca and Edwards are given for determining the
identity: “288. Lupinus mexicanus Cerv. ex. Lag. Gen. et
Sp. Nov. 22, 1816.
Calyces alternately bracteolate (appendaged) upper-lip
semibifid, lower lip obscurely tridentate.
Lupinus mexicanus Cerv., near L. termis L.
Leaves ternate at the base, remains of 5-7 leaflets; leaf-
lets lanceolate, mucronate, upper surface glabrous, lower
surface pilose. Stipules setaceous, pilose as the stems.
Peduncles opposite the leaf, spicate-racemose above. Bracts
setaceous, deciduous. Flowers alternate (scattered), short
stalked, blue. Calyx bracteoles setaceous, short. Legumes
pilose.
Habitat in New Spain (Mexico). Seeds sent with D.
Vincent Cervantes."
From Edwards Bot. Reg. 457. 1820, the following trans-
lation: (Note, familial and generic portions are not given
and interpretations are included in parentheses). “Plants,
except for the corolla, entirely shaggy-pilose. Leaflets 3?-
5?, 7-8, elongate-cuneate, narrow, short pointed, tapered
for a considerable way down, glabrous above, rendered
white beneath by shaggy-pilose pubescence, longest about
2 inches, shorter than the long-piled petioles; stipules
linear-subulate, erect, long-piled. Racemes elongate spi-
cate, laxly many-flowered, flowers scattered, sparse, la-
492 Rhodora [Vol. 74
vender to purplish-blue, with darker streaks (veins),
bracteose before anthesis; peduncles opposite the leaves
(true of all lupines that have the top lateral bud elongate
as a branch) ; pedicels hirsute, shaggy, ascending, shorter
than the calyx, bracts linear-sublate, very narrow, filiform,
longer than the calyx, caducous. Calyces shaggy-hirsute,
green, with divaricate lips, the upper spilt at the end into
a notch. Banners folded, reflexed, astride the upper margins
of the wings by a deep sulcus; wings hatchet shaped,
pointed, cohering by the front edges; keels pale ascending
upward, narrow subulate-falcate, as long as the wings;
with a long deep-purple tapered point, — etc.”
The distinctive traits from which L. mexicanus can be
determined are the shaggy-pilose hairs, the upper surface
of the leaflets glabrous, while the lower is shaggy-pilose,
and the corolla is glabrous as intimated by Edwards, the
long setaceous bracts and short setaceous bracteoles at the
lateral sinuses of the calyx, the shaggy-hirsute pedicels,
the deep sulcus of the banner, the narrow subulate-falcate
keel with the long tapered point, and Edward's term of
pointed hatchet-shaped wings. His discussion of flowering
time and suggestion that the taxon is a biennial apply.
The field location is that of the most direct route between
the two early major cities of that time. All of the traits
apply to the biennial plant material collected by Giles
Waines. The greenhouse plants grown from seed more
closely resemble the illustration provided in Edwards, as
would be expected, since greenhouse plants were used in
preparing the description. The field material, in nature,
has denser foliage and a denser raceme, which would
represent the effects of high light intensity at high eleva-
tions.
The specimen collected by Giles Waines Lupin #2 (UMO-
88263) (Fig. 1) is considered as typical material of Lupinus
mexicanus Cerv. ex. Lag. A field duplicate has been sent
to Madrid (MA) and a duplicate grown in the greenhouse
has been sent to Cambridge (CGE) ; others will be distrib-
uted as available. The close relatives are Lupinus bilineatus
1972] Lupinus — Dunn 493
id
eo
2*
Lapin dI
ce ld act e
fact. ged
Asper Quest As
i per- os Q4 p
i Tum pole [Mesie - Ga wu
1 ds "CA. ea owt e Ale
i ke Tase pila en Dek,
i /
i rdi trad .
: Vlas, Dt bee.
i fy í
i WM L i
H
due got ead tal Z id Bel he
pu ur bola E
guy uet zy
a
riw
Plants
Figure 1. A field specimen of Lupinus mexicanus Cerv. ex. Lag.,
a biennial collected in July by Giles Waines, Lupine #2 (deposited
at UMO; duplicate sent to MA; a greenhouse grown duplicate sent to
CGE).
494 Rhodora [Vol. 74
Benth (1839), L. hartwegii Lindl. (1839), and L. persis-
tens Rose (1905).
The taxon which has been misinterpreted as Lupinus
mexicanus by several authors is Lupinus aschenbornü.
While the latter taxon has glabrous flowers and a glabrous
upper leaflet surface and pilose hairs, the hairs do not match
Edward's term of shaggy nor are the leaflets shaped right
nor is the keel slender, long-faleate. The keel is almost
straight on the upper edge and the banner is narrow and
without the deep sulcus. In addition L. aschenborni is
alpine and it is a perennial, which goes dormant every
winter. It is a most unlikely candidate for a greenhouse
plant or a plant that could be grown outside in Madrid,
Spain, at the Botanical Garden. In addition the flower size
and shape are entirely wrong for the material illustrated
by Edwards from seeds sent by Lagasca.
DIVISION OF BIOLOGICAL SCIENCES
UNIVERSTY OF MISSOURI-COLUMBIA
COLUMBIA, MISSOURI 65201
NOMENCLATURAL AND TAXONOMIC NOTES
ON MEXICAN COMPOSITAE
ROGERS McVAUGH
The following notes have accumulated during the prepa-
ration of a treatment of the family Compositae for a pro-
posed Flora Novo-Galiciana. Included are one new com-
bination, discussions of points of nomenclature, mention
of notable range-extensions, and comments on taxonomic
decisions, all of which are somewhat out of place in a
formal floristic treatment. The area covered by the pro-
posed flora includes the Mexican States of Jalisco, Colima,
and Aguascalientes, and some adjoining territory; for a
fuller description of it see Brittonia 13: 145-147. 1961,
Contr. Univ. Mich. Herb. 9: 1-7. 1966, or Contr. Univ.
Mich. Herb. 9: 207-357. 1972. Another paper preliminary
to the Flora Novo-Galiciana, including descriptions of a
number of taxa of Compositae new to science, appeared
in Contr. Univ. Mich. Herb. 9: 359-484. 1972. For general
support of the field-work and other activities on which
this paper is based, I am grateful to the National Science
Foundation (Grant no. GB-5218X).
In the course of revision of the various groups of Com-
positae for the Flora, I have seen most of the relevant types
(or in some instances isotypes). When I have seen and
studied a specimen, this is indicated in the text below in
the conventional way [!]. To the many persons and insti-
tutions that have permitted and assisted my studies of the
valuable collections in their charge, my sincere thanks.
Many of the remarks on nomenclature refer to the names
published by Kunth in the fourth volume of Nova Genera
et Species Plantarum (1818), or those published by De-
Candolle in the fifth and sixth volumes of his Prodromus
(1836, 1838). To the authorities at Paris, where I have
studied most of the types of Humboldt & Bonpland, and at
Geneva, where similarly I have seen and studied the types
of DeCandolle, I am most grateful.
495
496 Rhodora [Vol. 74
It is necessary to comment on the dates of publication of
the fourth volume of the Nova Genera et Species, which
dealt wholly with the Compositae. Folio and quarto editions
of this were published simultaneously (as they were for
the other 6 volumes of the set). Volume 4 appeared in
1820, in 5 installments, on 17 Apr, 22 Mai, 31 Jul, 18 Sep
and 26 Dec, respectively. Apparently the quarto edition
was not available to the public before these dates, nor was
the folio edition generally distributed. The printing of the
folio text was completed, however, in September 1818, and
at least 4 copies were distributed and became more or less
accessible to the botanical public by 1 December 1818. The
authors presented a copy to the Institut in Paris on 26
October 1818. Under the code of nomenclature now in
force, this constitutes effective publication as of that date,
of all the new names proposed in the volume. The fact that
the authors considered the work unpublished until it was
generally distributed in 1820, is irrelevant. In citations of
names published in the Nova Genera. et Species, botanical
custom has been to cite the page-numbers of the quarto
edition, as this has been more generally available. Some
authors have cited the pages of both quarto and folio edi-
tions, on the assumption that the two were published simul-
taneously. Names of Compositae, however, are correctly
cited as from the folio edition only, because publication of
the quarto pages was delayed for a year and a half to two
years. For summary of the dates of publieation of both
editions and the cireumstances surrounding the distribu-
tion of volume 4, see Stafleu, F., Taxonomic Literature
(Regnum Veg. 52: 225-226. 1967). Many of the circum-
stances were first made publie by Henri Cassini, who was
given a copy of the folio text by Kunth, and who criticized
it in his own published articles before the dates of general
distribution in 1820. Cassini's remarks, originally pub-
lished in the Journal de Physique, de Chimie, d'Histoire
Naturalle et des Arts, early in 1819, were reprinted in his
Opuscules Phytologiques, vol. 1, pp. 324, 339, etc. 1826.
1972] Mexican Compositae — McVaugh 497
Ageratella microphylla (Sch. Bip.) A. Gray in S. Wats.
Proc. Am. Acad. 22: 419. 1887.
Ageratum microphyllum Sch. Bip. in Seem. Bot. Voy. Her-
ald 298. 1856.
Ageratella microphylla var. seemannii and var. palmeri A.
Gray in S. Wats. Proc. Am. Acad. 22: 419. 1887.
Ageratella palmeri (A. Gray) Rob. Proc. Am. Acad. 41:
272. 1905.
The type of A. microphylla (from the “Sierra Madre",
i.e. perhaps from Cerro del Pinal, Sinaloa, Seemann 2043!,
at K) was taken from a plant with unusually broad leaves,
as well shown in the illustration published by Hemsley
(Biol. Centr. Am. Bot. 5: pl. 42. 1881). The specimen
evidently represents a portion of a plant in which several
long leafy branches arose from a larger stem. As is often
the case in the Compositae, the leaves are opposite at the
lower nodes of such branches, but sometimes alternate
above. Similarly abnormally branched specimens, of other-
wise typical “palmerv’, have the same leaf-arrangement
but with fewer opposite leaves (e.g. Gaiser 62, at MICH,
from near Guadalajara). Nothing exactly like the type of
microphylla has been found by any subsequent collector,
and we can but surmise what it may represent, until it
can be found again in the field.
Gray (1887) distinguished what he called two varieties
on the basis of the broad, incised-dentate leaves and
subspicate inflorescence of var. seemannii (i.e. typical
microphylla), and the narrow, subentire leaves and looser
inflorescence of var. palmeri. Robinson (Proc. Am. Acad.
41: 271-272. 1905), and Blake, in the Trees and Shrubs
of Mexico, noted these features, and stated also that the
leaves were alternate in palmeri but opposite in microphylla.
Both these authors treated the two taxa as distinct species.
Blake's key is based entirely on leaf-characters. In fact it
seems that neither position nor shape of leaves is completely
diagnostic. In the original specimens of var. palmeri
(Palmer 587!, at GH, from near Guadalajara), and in most
498 Rhodora [Vol. 74
flowering specimens, the narrow leaves of sterile axillary
shoots are much in evidence, giving the impression of a
narrow-leaved plant. There is much individual variation
in this respect, but the main cauline leaves subtending
axillary shoots are seldom very narrow, and often much
toothed or lobed.
The differences between “spikelike” and “loose” inflores-
cences also appear to be related to individual differences
in branching. When vigorous lateral leafy branches (e.g.
those 10-30 cm long) produce a number of short-peduncled
axillary clusters of flower-heads, the effect is of a spike;
when (as in most specimens), the lateral branches are
reduced in length, and in number of nodes, and bear few
or no heads except small clusters at the tips, the effect is
of a small loose panicle. Various intermediates can be
demonstrated.
Archibaccharis hieraciifolia Heering, Jahrb. Hamburg.
Wiss. Anst. 21, Beih. 3: 40. 1904.
Baccharis hieraciifolia Hemsl. Biol. Centr. Am. Bot. 2:
129. 1881, not B. hieracifolia Lam., 1783.
Hemibaccharis hieraciodes Blake, Contr. U.S. Nat. Herb.
20: 547. 1924.
Archibaccharis hieracioides (Blake) Blake, Jour. Wash-
ington Acad. Sci. 17: 60. 1927.
Blake (1927) discussed the nomenclature of this species
and that of Archibaccharis hirtella. Both were assigned
names by Heering in 1904, but both were called ''n.spec."
in spite of the fact that both names were derived from
previously published names. As Blake said, “it seems ad-
visable to treat these two names of Heering as representing
new combinations". Under the International Code of
Botanical Nomenclature (Art. 72) a new combination
derived from a previously published but illegitimate name
(e.g. Archibaccharis hieraciifolia from Baccharis hieracit-
folia Hemsl.) may be treated as a new name. It is there-
fore proper to use the name originally proposed by Heering,
not Archibaccharis hieracioides, the substitute proposed by
Blake.
1972] Mexican Compositae — McVaugh 499
Baccharis sulcata DC. in DC. Prodr. 5: 419. 1836.
Baccharis potosina A. Gray, Proc. Am. Acad. 15: 33. 1879.
Blake (in the Trees and Shrubs of Mexico, 1926) treated
Baccharis sulcata as a synonym of B. thesioides H.B.K.,
but the type of B. sulcata (Villalpando, Méndez! in G-DC)
certainly represents another species. The narrow leaves
(up to 3-3.5 mm wide) vary from quite entire to toothed,
with up to 4 or rarely 6 short teeth per cm of margin. In
other respects, including the characters of heads and in-
florescence, and the habit, the plants are typical of what
has been called B. potosina (San Luis Potosí, Parry &
Palmer 410! at GH, the type).
Bidens acrifolia Sherff, Bot. Gaz. 94: 591. 1933.
Bidens polyglossa Sherff, Brittonia 16: 61. 1964.
Southern Sinaloa (Concordia, Dehesa 1532!, K, the type),
and western Jalisco (northwest of Cuautla, McVaugh
13633!, MICH, type of B. polyglossa).
The type of Bidens acrifolia is an immature and incom-
plete specimen, but in all observable details it agrees pre-
cisely with the abundant available material of B. poly-
glossa. The apparent differences mentioned by Sherff
(Brittonia 16: 62. 1964) are insignificant or (those per-
taining to the leaf-pubescence and to the phyllaries) non-
existent.
Chaptalia runcinata H.B.K. Nov. Gen. & Sp. 4 [ed. fol.]:
5. pl. 303. 1818.
Sonora (Pennell 19647), western Durango (Maysilles
7452, 7809, 7848; Cronquist 9560), northern Nayarit (Rose
2022, at US). Costa Rica; Venezuela and Colombia (Hum-
boldt & Bonpland!, at P, the type) ; Bolivia to southeastern
Brazil and northern Argentina. Not previously reported
from north of Costa Rica, but abundant on the summer-
wet, high pine plains of western Durango (cited specimens
from Sonora and Durango all at MICH). The seasonal
forms with reduced or filiform marginal flowers, reported
by Burkart for South American representatives of this
species, have not been found in Mexico, but otherwise the
500 Rhodora [Vol. 74
Mexican material agrees perfectly with that from further
south.
Conyza viscosa Mill. Gard. Dict. ed. 8. Conyza no. 8. 1768.
Conyza lyrata H.B.K. Nov. Gen. & Sp. 4 [ed. fol.] : 55. 1818.
The type, from Veracruz (Houstoun!, BM), is a pale-
pilose plant resembling what has been called Conyza lyrata
var. pilosa Fernald, Proc. Am. Acad. 36: 506. 1901 (Chia-
pas, Seler 1879!, GH, the type), whereas the type of C.
lyrata H.B.K. (Guayaquil, Hwmboldt & Bonpland!, P) is
much less hairy, with more conspicuous glands.
Erigeron longipes DC. in DC. Prodr. 5: 285. 1836.
Erigeron scaposum DC. in DC. Prodr. 5: 287. 1836.
Erigeron scaposum p latifolium DC. in DC. Prodr. 5: 287.
1836.
Erigeron affine DC. in DC. Prodr. 5: 289. 1836.
From Coahuila and perhaps Durango southward nearly
throughout southern Mexico to Central America, variable
in habit, originally described as subscapose. The leaves,
especially in some parts of eastern Mexico, may be grouped
toward the base of the stem, whereas in Nueva Galicia the
leafy part of the stem is usually elongated and the leaves
well spaced along it. I have not seen the type (Karvinskt,
s.n., at M), which is from some unknown locality in Mexico,
but judging from fragments in DeCandolle's herbarium
(G-DC!), it represents the same species as the other names
cited above. Subscapose forms seem to be especially
abundant in Oaxaca, and it may be that Karvinski's speci-
mens came for that state. Hrigeron scaposum and E.
longipes were described by DeCandolle as perennials,
whereas E. affine, the type of which was a plant with
elongated leafy stems, was erroneously described as an
annual. The type of E. affine, from near the City of Mexico
(Berlandier 522!, G-DC), almost certainly represents a
perennial species. The basal parts of the plant are much
like those of E. scaposum, and in fact the two supposed
species are essentially similar except that in E. affine the
hairs are longer on both leaf-surfaces. The type of E. sca-
1972] Mexican Compositae — McVaugh 501
posum came from near Toluca (Andrieux 277!, G-DC), and
that of B latifolium from near Mexico City (Berlandier
375!, G-DC).
Erigeron velutipes Hook. & Arn. Bot. Beech. Voy. 434. 1841.
Erigeron alamosanum Rose, Contr. U.S. Nat. Herb. 1: 102.
1891.
Western Mexico, from southern Sonora (Palmer 348!, vs,
the type of E. alamosanum) to Chihuahua, Sinaloa, southern
Zacatecas, Nayarit, Jalisco, and Michoacán. The type (Sin-
clair s.n.!, at K) was collected between San Blas and Tepic.
Plants of E. velutipes have often been misidentified with
Erigeron tenellum DC. in DC. Prodr. 5: 288. 1836, the
type of which came from Matamoros, Tamaulipas. (Ber-
landier 2129! in G-pc, lectotype). E. tenellum, like E. velu-
tipes, is a slender branched annual, but the herbage is
eglandular and the base of the plant is only moderately stiff-
hairy. The name E. alamosanum was based on specimens
a little larger than average for the species. Similarly
vigorous plants are not infrequent along the Pacific Slope
from Sonora to Nayarit.
Eupatorium albicaule Sch. Bip. ex Klatt, Leopoldina 20: 89.
1884.
Eupatorium albicaule var. laxius Rob. Proc. Am. Acad.
35: 330. 1900.
Eupatorium leucoderme Rob. Proc. Am. Acad. 41: 274.
1905.
Eupatorium ymalense Rob. Contr. Gray Herb. II. 75: 14.
1925.
Lowlands, sea-level to 300 m. in elevation, Sinaloa
(Ymala, Palmer 1474!, GH, type of var. laxius and of E.
ymalense), Nayarit, Jalisco, Colima, Michoacán (Chuta,
Langlassé 183!, GH, type of E. leucoderme) ; Tabasco and
Chiapas; Yucatán Peninsula; north to Veracruz (Papantla,
Liebmann 88!, P, an isotype), San Luis Potosí, and Tamau-
lipas. This geographical range is like that of many other
species of tropical Mexico; see the maps in Arboles Tropi-
cales de México, by Pennington & Sarukhan (published by
502 Rhodora [Vol. 74
the Instituto Nacional de Investigaciones Forestales, and
the FAO, pp. vii, 413. México, 1968).
Plants growing in western Mexico were distinguished
from the original Eupatorium albicaule, first as the var.
laxius and later as an independent species, E. ymalense.
The only significant difference between the two populations
seems to lie in the shape of the phyllaries, which are acute
or attenuate in the plant of the Pacific lowlands, and usu-
ally, but not always, blunt or subacute in the plant of
eastern Mexico and the Yucatán Peninsula. Robinson re-
ported the heads in E. albicaule as *about 7 or 8-flowered",
but they are usually 10-13-flowered, as in the supposed E.
ymalense.
Eupatorium collinum DC. in DC. Prodr. 5: 164. 1836, var.
collinum.
Eupatorium stillingiaefolium DC. in DC. Prodr. 5: 160.
1836.
This variety appears to be restricted to eastern Mexico.
The type (Berlandier 2162! in G-DC), from near Tantoyuca
in the Atlantic lowlands of Veracruz, apparently does not
differ significantly from the type of E. stillingiaefolium
(Berlandier 2142! in G-DC), which was collected in Tamau-
lipas. In these plants, and in modern specimens from the
same region, the phyllaries are narrow and attenuate or
acute (the middle ones 1 mm wide), often nearly all equal,
sparingly ciliate, evidently resinous-dotted and more or
less densely short-pubescent, but scarcely if at all arachnoid-
tomentose. In a second variety ranging widely through
central and western Mexico into Central America, the
plants are more pubescent and less conspicuously glandular,
the phyllaries are more strongly graduated, and some or
all of them obtuse and evidently closely fimbriate-ciliate.
I can detect no great variation within the limits of this
second population, except that rather densely pilose indi-
viduals seem to be more frequent in Chiapas and Central
America than from Guerrero westward. The second variety
is the following:
1972] Mexican Compositae — McVaugh 503
Eupatorium collinum DC., var. mendezii (DC) McVaugh,
comb. nov.
Eupatorium mendezii DC. in DC. Prodr. 5: 160. 1836.
Eupatorium neaeanum DC. in DC. Prodr. 5: 160. 1836.
?Eupatorium nigrescens Hook. & Arn. Bot. Beech. Voy.
297. 1838.
Sonora to Jalisco, Guanajuato (León, Mendez!, in G-DC,
the type), Morelos, Guerrero (Acapulco, Née !, in GDC;
type of E. neaeanum), Oaxaca, Chiapas and Central
America.
DeCandolle grouped E. neaeanum, E. mendezii and E.
stillingiaefolium with other species having the heads 25- 30-
flowered, but E. collinum was described as having the heads
90-flowered, and accordingly was not closely associated
with the others in the Prodromus. I suspect that some
error was involved here, although Robinson (in the Trees
and Shrubs of Mexico, p. 1448) states that the heads in
collinum may have as many as 46 flowers. The number
of flowers usually varies from 22 to 28; it is rarely as low
as 18 or 20, and even more rarely 30 or more; I have seen
only one specimen in which the heads were about 36-
flowered, and none with a larger head.
Robinson separated E. collinum from E. mendezii and
E. neaeanum partly on the basis of pubescence and leaf-
shape. The latter seems quite useless as an indicator of
Specific lines in this group of species. Pubescence in the
group is highly variable, as noted above, but the types of
E. mendezii and E. neaeanum are not strongly pubescent
in comparison with other specimens from central and
western Mexico, and the differences between them and
typical collinum are no more than might be expected of
regional populations in one species.
Eupatorium hebebotryum (DC.) Hemsl. Biol. Centr. Am.
Bot. 2: 95. 1881.
Critonia hebebotrya DC. in DC. Prodr. 5: 141. 1826.
Hebeclinium tepicanum Hook. & Arn. Bot. Beech. Voy.
434. 1841.
504 Rhodora [Vol. 74
Eupatorium tepicanum (Hook. & Arn.) Hemsl. Biol. Centr.
Am. Bot. 2: 101. 1881.
Nayarit (Tepic, Sinclair!, K, type of H. tepicanum),
Jalisco, Michoacan, Morelos, Guerrero (Haenke!, in G-DC,
the type probably from this state) ; Central America. The
distinctions used to separate the two supposed species,
namely that between sessile and pedicellate heads, and
that in the color of the dried leaves, appear to be inconse-
quential.
Eupatorium ovaliflorum Hook. & Arn. Bot. Beech. Voy. 297.
1838.
Eupatorium. bertholdii Sch. Bip. in Seem. Bot. Voy. Herald
299. 1856.
Southern Sonora, ? western Durango (*Sierra Madre",
Seemann 2011! at K, isotype of E. bertholdii), Sinaloa,
Nayarit (Tepic, Beechey!, at K, the type), Jalisco, Michoa-
cán.
According to Robinson (in Trees and Shrubs of Mexico,
p. 1433), Eupatorium bertholdii is distinguished from E.
ovaliflorum by having the heads 10- to 13-flowered and the
involucre 2-2.6 mm thick, as against heads 20- to 40-flowered
and the involucre 4-5 mm thick. In the material at hand
these differences seem not to hold; I have not seen any
head with more than 22 flowers, and none as much as 4mm
thick. The flowers are usually about 17-20, and the in-
volucre 2.5-3 mm thick.
Eupatorium polybotryum DC. Prodr. 5: 174. 1836.
Nothites ovatifolia DC. in DC. Prodr. 5: 187. 1836, not
Eupatorium ovatifolium Hieron., 1908.
Ophryosporus ovatifolius (DC.) Hemsl. Biol. Centr. Am.
Bot. 2: 79. 1881.
Eupatorium petraeum Rob. Proc. Am. Acad. 41: 2775. 1905.
Ophryosporus petraeus (Rob.) Rob. Contr. Gray Herb. II..
75: 4. 1925.
Decachaeta ovatifolia (DC.) King & H. Rob. Brittonia 21:
282. 1969.
Southeastern Jalisco, western Michoacán, Edo. de México,.
1972] Mexican Compositae — McVaugh 505
Guerrero (Langlassé 565!, GH, type of E. petraeum). A
related species ranging from western Michoacán north-
westward to Sinaloa and Chihuahua is Eupatorium sca-
brellum Rob. Proc. Am. Acad. 35: 339. 1900 [E. micro-
cephalum A. Gray, Proc. Am. Acad. 21: 384. 1886, not of
Regel; Ophryosporus ovatifolius sensu Rob. in Standl.
Contr. U.S. Nat. Herb. 23: 1469. 1926, not Nothites ovati-
folia DC.]. E. scabrellum and E. polybotryum are separable
as follows:
1. Phyllaries 12-15, 2- 4-seriate, the outer gradually
shorter, the innermost (2-5 in each head) longer
and narrower than the others and completely or
partially modified into pales, deciduous with the
achenes or before; flowers (7-) 10-12 (-15). ....
D TOR AEHROE COUPES OTT E. polybotryum.
1. Phyllaries 6-8 in 1-2 nearly equal series (in addition
to 2-3 smaller outer bracts), the innermost neither
prolonged and resembling pales, nor deciduous;
lowers 4.9... oaa m as E. scabrellum.
The nomenclature of these species has become confused.
In the Trees and Shrubs of Mexico, Robinson (1926, p.
1469) took up the name Ophryosporus ovatifolius but mis-
applied it to the species correctly known as Eupatorium
[Ophryosporus] scabrellum, at the same time recognizing
as independent species O. scabrellus (Rob.) Rob. and O.
petraeus (Rob.) Rob. Robinson said of his Ophrysporus
ovatifolius, “Typical material of this species, early collected
by Haenke in Mexico but without indication of locality, has.
never been precisely matched except by specimens collected
by Seemann, also without recorded locality". Presumably
Robinson referred to two collections studied by him at the
Gray Herbarium; the Seemann collection is the type of
Eupatorium microcephalum A. Gray (= E. scabrellum
Rob.). The Haenke specimen consists of a few heads and
the tip of a branch of an inflorescence, annotated by Gray
as “Nothites ovatifolia DC.', and by Robinson as Eupa-
torium polybotryum DC. [ie. Ophryosporus ovatifolius
506 Rhodora [Vol. 74
of Robinson’s treatment in 1926]. Unfortunately Robin-
son seems to have misinterpreted this fragment, which
certainly represents Eupatorium petraeum, not E. micro-
cephalum. The heads are only T- 8-flowered, but the in-
volucre is several-seriate, and the narrow paleaceous inner
phyllaries are unmistakably those of "E. petraeum". The
original Haenke specimens at G-DC, the types of Eupa-
torium polybotryum and Nothites ovatifolia, respectively,
I have seen through the courtesy of Prof. J. Miége. These
represent one and the same species, as Robinson long ago
suggested, but that species is “Eupatorium petraeum”, not
Ophryosporus ovatifolius in the sense of Robinson. Both
the Haenke collections represent a species with 12-15
graduated phyllaries of which the inner are deciduous at
maturity; with 1-3 narrow pales on the receptacle; and the
flowers 8-11 in each head.
King and H. Robinson (1969) first formally combined
Eupatorium polybotrium [sic] and Nothites ovatifolia,
under the name Decachaeta ovatifolia. The correct epithet
for the combined taxa is therefore ovatifolia except in
Eupatorium where the name is preoccupied.
Gnaphalium canescens DC. in DC. Prodr. 6: 228. 1838.
Gnaphalium wrightii A. Gray, Proc. Am. Acad. 17: 214.
1882.
There seems to be no significant difference between the
plants of nothern Mexico and southwestern United States
that have been called G. wrightii (western Texas, Wright
394!, the type), and the plants of central Mexico, from
Durango and Aguascalientes to Jalisco, Guanajuato,
Hidalgo and the Valley of Mexico, that have been called
G. canescens (León, Méndez, in G-DC, the type). The leaves
in this species tend to be broadest near the middle or above.
A very similar species is Gnaphalium roseum H.B.K. Nov.
Gen. & Sp. 4 [ed. fol]: 63. 1818, in which the leaves tend
to be oblong or broadest at base. The type (from Guana-
juato, Humboldt & Bonpland!, P), is a coarse woolly plant
with immature heads.
1972] Mexican Compositae — McVaugh 507
Gnaphalium inornatum DC. in DC. Prodr. 6: 225. 1838.
This species was based upon four collections made by
Berlandier, viz. no. 1195 (*ad montem de Las Cruces"),
no. 740 (from the Valley of Mexico), no. 309 (between
Tampico and Real del Monte) and no. 1146 (from Guchi-
laque, between Mexico and Cuernavaca). DeCandolle de-
scribed it as a woolly plant with narrow erect shortly
decurrent leaves, pale reddish (‘‘refescentibus”) phyllaries,
about 10 hermaphrodite flowers and 30-40 pistillate flowers.
Judging from the specimens in the Prodromus herbarium
(cf, also IDC microfiche 1066), and from duplicates (at P)
of all the cited numbers, it seems that three different species
are represented by the syntypes of G. inornatum. The
description in the protologue does not exactly fit any of
the species. Nos. 1146 and 1195 apparently represent the
same species, in which the number of flowers in a head is
100 or more and the perfect flowers are 7-11 or more.
DeCandolle's report of 10 perfect flowers presumably was
based on one of these specimens, and one of them may
appropriately be designated as lectotype. No. 740 may be
eliminated from consideration as the heads are about 50-
flowered, and the perfect flowers are 4 in number. De-
Candolle's specimen is so immature that it seems unlikely
that he dissected a head (cf. Field Mus. neg. 28713). The
remaining syntype, no. 309, seems to represent a third
species, also one in which the heads are about 50-flowered
and the perfect flowers about 5.
Berlandier 1146 is designated as lectotype since in the
Prodromus herbarium it appears to be a somewhat better
specimen (than no. 1195) ; since from the position of speci-
mens in the herbarium it seems to have been the one on
which DeCandolle's report of 10 perfect flowers was based;
and since from Berlandier's notes it appears this collection
was more widely distributed than his no. 1195.
Gnaphalium sphacilatum H.B.K. Nov. Gen. & Sp. 4[ed. fol.] :
67. 1818.
Gnaphalium pedunculosum I. M. Johnst. Contr. Gray Herb.
II. 68: 99. 1923.
508 Rhodora [Vol. 74
Durango (Palmer 411! in GH, the type of G. pedunculo-
sum), northern Jalisco, San Luis Potosi, D.F., Edo. de
México (Teotihuacán, Hahn; beween the City of Mexico
and Huehuetoca, Humboldt & Bonpland!, the type, at P).
Similar, narrow-leaved plants with uniformly gray thin
tomentum occur in various other areas both in North and
South America, and have been called by various other
names, either treated as distinct species or as varieties of
Gnaphalium purpureum. The type of G. sphacilatum is
an immature but otherwise typical plant with the charac-
teristic bracts, pappus, glabrous acute phyllaries, and linear
thinly silky leaves.
Gnaphalium stramineum H.B.K. Nov. Gen. & Sp. A[ed. fol]:
66. 1818.
Gnaphalium chilense Spreng. Syst. Veg. 3: 480. 1826.
Gnaphalium sprengelii Hook. & Arn. Bot. Beech. Voy. 150.
1833.
Gnaphalium berlandieri DC. in DC. Prodr. 6: 223. 1838.
Material from central and eastern Mexico can usually
be distinguished from California specimens of Gnaphalium
“chilense” because of the somewhat narrower leaves and
the tendency for the uppermost leaves to become bractei-
form. The types of G. stramineum (between Moran and
Omitlán, Humboldt & Bonpland!, at P), and G. berlandieri
(D.F., Berlandier 471!, in G-DC) are representatives of the
Mexican population. The Humboldt & Bonpland specimens
were apparently badly wilted before drying, and perhaps
taken originally from slender or starved plants, but the
heads have the characters of this species (flowers 175 or
more, hermaphrodite flowers 15-16, phyllaries obtuse, the
inner ones about 23). An isotype at Paris (Bonpland 4108)
seems clearly to represent the same taxon. The character-
istic yellow color of the phyllaries can no longer be seen in
the original material of G. stramineum.
Gnaphalium viscosum H.B.K. Nov. Gen. & Sp. 4[ed. fol.]:
64. 1818.
Gnaphalium hirtum H.B.K. Nov. Gen. & Sp. 4[ed. fol.] : 64.
1818.
1972] Mexican Compositae — McVaugh 509
27Gnaphalium gracile H.B.K. Nov. Gen. & Sp. A[ed. fol.]:
65. 1818.
Gnaphalium tenue H.B.K. Nov. Gen & Sp. 4[ed. fol.]: 65.
1818.
Gnaphalium leptophyllum DC. in DC. Prodr. 6: 226. 1838.
A plant primarily of the Central Plateau of Mexico,
widely distributed from western Texas to northeastern
Sonara, south at moderate elevations except in the extreme
deserts to Oaxaca and Central America. Cronquist (in
Vasc. Pl. Pacific N.W. 5: 204. 1955; and Man. Vasc. Pl.
N.E. U.S. & Can. 737. 1963) has applied the name Gnapha-
lium viscosum very broadly, so as to include the plant of
temperate North America that has been better known as
G. decurrens Ives, or G. macounii Greene. This seems to
be unrealistic; G. viscosum has a characteristic Texano-
Mexican range distinct from that of G. macowunii and, com-
pared with other currently accepted species of Gnaphalium,
it is morphologically very distinct. It differs from G. ma-
counii vegetatively in its far more numerous and narrower
leaves. The inner phyllaries in G. macounit are usually
about 21, the flowers fewer (125-150 in a head) but often
larger than those of G. viscosum. At least until the com-
plex of glandular, decurrent-leaved fragrant species of
Gnaphalium can be revised as a whole, G. viscosum may
well be circumscribed narrowly. Nothing exactly referable
to G. macounii has been found in Nueva Galicia, but nu-
merous collections from the Sierra Madre Oriental suggest
that G. macounii ranges well south into eastern Mexico.
In Chihuahua G. viscosum may be confused with G. leuco-
cephalum A. Gray, in which the phyllaries are pearly
white, relatively dull and opaque and symmetrically gradu-
ated in length.
This is the only Mexican species having the herbage
strongly glandular-pubescent throughout, the leaves very
numerous (100 or more in well developed plants) and
linear or narrowly sagittate, the flowers 200 or more in a
head, the mature receptacle 3-4 mm wide, and the plants
strictly annual, not forming a basal rosette. The types of
G. viscosum, G. hirtum, G. tenue and G. leptophyllum seem
510 Rhodora [Vol. 74
without question to represent the same taxon. The type of
G. gracile (Guanajuato, Humboldt & Bonpland! at P) is
apparently from a weak, depauperate plant with relatively
few leaves and numerous heads.
Heterotheca inuloides Cass., var. rosei Wagenknecht, Rho-
dora 62: 69. 1960.
Heterotheca leptoglossa DC. in DC. Prodr. 5: 317. 1836.
In Wagenknecht’s revision, Heterotheca leptoglossa is
maintained as a distinct species, distinguished from H.
inuoides by “its annual habit, narrow leaves, smaller capi-
tular [sic], and linear phyllaries". The type of H. lepto-
glossa. (León, Méndez!, in G-DC and cf. Intern. Doc. Center
microfiche, Prodromus herbarium, no. 857), is the upper
part of what appears to be a vigorous plant of H. inuloides,
with leaves normal for that species. As many plants of
H. inuloides flower the first year, the differences between
annual and perennial habit in this group are of question-
able significance. In the type of H. leptoglossa neither
phyllaries nor ligules are atypical of H. inuloides. The
ligules are not linear as described by DeCandolle but ob-
lanceolate, 2.5 mm wide by 10-11 mm long. The heads are
of about average size for H. inuloides, and the phyllaries
are not unusually narrow.
Montanoa [*Montagnaea"] karvinskii DC. in DC. Prodr. 5:
565. 1836.
Montanoa olivae Sch. Bip. ex. K. Koch, Wochenschr. Gaertn.
7: 406. 1864.
Montanoa gracilis Sch. Bip. ex K. Koch, Wochenschr.
Gaertn. 7: 407. 1864.
Montanoa subtruncata A. Gray, Proc. Am. Acad. 22: 424.
1887.
?Montanoa affinis Blake, Contr. U.S. Nat. Herb. 22: 612.
1924.
A common and widely distributed plant, from Sinaloa
to Jalisco, Guerrero and Oaxaca, easily recognized by the
wingless petioles, the glabrous lobes and throat of the
disk-flowers, and the pilose anthers. It was long known
1972] Mexican Compositae — McVaugh 511
as M. subtruncata, until Blake (Contr. U.S. Nat. Herb.
26: 246. 1930) confirmed the identity of that plant with
M. olivae and M. gracilis. After examination of the type
of M. karvinskii (without locality, Karvinski s.n. in G-DC),
I can confirm Blake's tentative suggestion that this is a
still older name applying to the same species. Apparently
the plant called M. affinis is merely a nearly glabrous form
of the same species.
Oxypappus scaber Benth. Bot. Voy. Sulph. 118. 1845.
Chrysopsis scabra, Hook & Arn. Bot. Beech. Voy. 434. 1841.
not C. scabra, Ell., ?1823.
Pectis seemannii Sch. Bip. in Seem. Bot. Voy. Herald 309.
1856.
Oxypappus seemannii (Sch. Bip.) Blake, Contr. U.S. Nat.
Herb. 26: 261. 1930.
The name Oxypappus scaber was a new combination
based on the illegitimate name Chrysopsis scabra, Hook. &
Arn. According to the International Code of Botanical
Nomenclature (Art. 72) such a combination is not to be
rejected, but is to be treated as a new name. See also above
under Archibaccharis hieracifolia.
Tragoceros americanus (Mill.) Blake, Contr. U.S. Nat. Herb.
26: 240. 1930.
Calendula americana Mill. Gard. Dict. ed. 8. Calendula
no. 10. 1768.
Tragoceras schiedeanum Less. Linnaea 9: 269. 1834.
The original material of Calendula americana was sent
to Miller from Veracruz by William Houstoun, between
1729 and 1733. Blake (1930) reported, after examination
of the type, that it represented what had been called
Tragoceros microglossus DC. (DC. Prodr. 5: 533. 1836),
a rather local species of Jalisco and Guanajuato. According
to Torres, however (Brittonia 15: 290-302. 1963), the only
species of Tragoceros known from the Atlantic lowlands
of Mexico is the one called T. schiedeanus. The most east-
erly localities known for T. microglossus (T. “americanus” )
are more than 500 km from Veracruz, in quite a different
512 Rhodora [Vol. 74
vegetational zone. It seemed highly unlikely that a dis-
tinctive inland species like T. microglossus should have
been found near Veracruz about the year 1730, but never
have been recollected there. In April, 1970, therefore, I
reexamined the type of Calendula americana (Houstoun,
at BM). It is true, as Blake remarked, that the heads are
sessile or practically so, as in T. microglossus, but the
ligules of the ray-flowers are the narrow, tapering and
conspicuously bifid ones of T. schiedeanus, not the obtuse
or barely emarginate ones of T. microglossus. As such
bifid rays occur in this genus only in T. schiedeanus and
the related T. zinnioides (which has larger heads), there
can be little doubt that the Houstoun specimen represents
what has been called T. schiedeanus. It is therefore un-
fortunately necessary to use the name T. americanus for
this species, and relegate the well-known T. schiedeanus
to synonymy.
The original spelling of the generic name was Tragoceros,
and according to the International Code of Botanical No-
menclature this spelling must be kept, although various
authors have changed it to Tragoceras in an attempt to
conform to the spelling of the classical Greek word for
horn, from which the -ceros part of the name was derived.
The Code recommends (Rec. 75A) that names ending in
-ceras be treated as neuter in the future, but this does not
affect names already published, and in no way authorizes
anyone to alter the original spelling of any such name.
There is ample biological precedent for names ending in
-ceros (not -ceras), as pointed out by Bentham a century
ago (Gen. Pl. 2, pt. 1: 356. 1873). Such names as Antho-
ceros and Rhinoceros have been consistently treated as
masculine since the time of Linnaeus. Tragaceros has been
used in its original form by some authors (e.g. Hemsley
and Asa Gray), who have treated it as masculine. Others
(notably Lessing, DeCandolle and most recently Torres)
who have used the -ceras spelling have treated the name
as neuter. There would seem to be no justification for this
latter course under the Code, and the neuter forms of
1972] Mexican Compositae — McVaugh 513
specific epithets published under Tragoceras are to be
treated as orthograhic errors.
Trigonospermum adenostemmoides Less. Syn. Gen. Comp.
214. 1832.
Since the appearance of the note on this species by Mc-
Vaugh and Laskowski (Contr. Univ. Mich. Herb. 9: 498-
500. 1972) I have seen the type of the name, through the
courtesy of the Director of the Institut für Systematische
Botanik und Pflanzengeographie of the Martin Luther
University, Halle (HAL). The specimen bears Schiede's
original label: “Composita. Herba annua caule ultraorgyali.
Jun. 29", and an annotation in the hand of Lessing:
“Trigonospermum n[.] g[.] adenostemoides n[.] sp [.]"
The ample fruiting and flowering specimen clearly repre-
sents the taxon treated as T. adenostemmoides in our paper
cited above.
Vernonia triflosculosa H.B.K. Nov. Gen. & Sp. 4[ed. fol.]:
31. 1818.
Vernonia barbinervis Sch. Bip. in Seem. Bot. Voy. Herald
297. 1856.
Vernonia (?) palmeri Rose, Contr. U.S. Nat. Herb. 1: 101.
1891.
Vernonia chacalana Blake, Contr. Gray Herb. II. 52: 19.
1917.
Southern Sonora (Palmer 387!, at US, type of V.(?)
palmeri), Sinaloa, Durango (Chacala, Goldman 333!, at
GH, type of V. chacalana), Nayarit, Jalisco, Colima, Guer-
rero (Acahuizotla, Humboldt & Bonpland!, the type) ;
Central America. The type of V. barbinervis (Seemann
1998!, at P), came from the “Sierra Madre", i.e. probably
from Durango, Sinaloa, or northern Nayarit.
According to both Gleason and Blake, Vernonia palmeri
is a distinct species ranging from Sonora to Tepic, char-
acterized by the (usually) relatively abundant pubescence
of the lower leaf-surface. Vernonia barbinervis, known
only from the type-region, has the foliage almost glabrous
except that the leaves are densely tomentose along the
514 Rhodora [Vol. 74
midvein beneath. In typical Vernonia triflosculosa, sup-
posed to range from Colima to Costa Rica, the leaves are
said to be glabrous to “thinly tomentulose" beneath. Both
glabrous- and pubescent-leaved plants occur in Nueva
Galicia, but I do not find any other features correlated
with differences in pubescence, and I believe only one
species can be recognized.
Viguiera puruana Paray, Bol. Soc. Bot. Méx. 22: 4. 1958.
Viguiera blakei McVaugh, Contr. Univ. Mich. Herb. 9:
454. 1972.
My attention was called to the similarity between
Viguiera puruana and V. blakei by a specimen collected
in Michoacán (6 km south of Tuxpan on the road to Zitá-
cuaro, Rzedowski 25153, MICH) and correctly identified
as V. puruana. Since then, through the kindness of Dr.
Ramón Riba y Nava Esparza, I have seen the type of V.
puruana (Michoacán, San José de Purta, Paray 1780,
MEXU). There can be no doubt it represents the taxon
more recently described under the name of V. blakei.
Xanthocephalum sericocarpum A. Gray, Proc. Am. Acad.
15: 31. 1879.
Two very similar species have long been confused under
this name. One is a perennial of arid grasslands and their
borders, ranging from Querétaro and San Luis Potosi to
northern Jalisco and the plains near the City of Durango.
The other is an annual, chiefly of pine forests, from western
Chihuahua to Aguascalientes and northern Jalisco. The
ranges of the two overlap, as far as known, only in Aguas-
calientes and Jalisco:
1. Phllaries 16-18 (-25); disk-flowers commonly about
80-85; plants perennial, from a woody taproot. ..
MNT X. sericocarpum.
1. Phyllaries 35-50; disk-flowers 100-200; plants annual.
peep e ween peessdessaveesaseneess X. conoideum.
The type of X. sericocarpum (San Luis Potosi, Parry
& Palmer 369!, GH) evidently represents the perennial
species. The heads are small, the phyllaries 20-25, broad
1972] Mexican Compositae — McVaugh 515
and relatively firm (not lanceolate or narrowly rhombic
with broad hyaline margins as in X. conoideum). A col-
lection by Schaffner, mounted on the type-sheet by Gray,
apparently represents the same species.
The annual species is X. conoideum Hemsl. Biol. Centr.
Am. Bot. 2: 110. 1881. The type (Coulter 299!, at K) was
collected somewhere in central Mexico, “between Real del
Monte and Zacatecas".
Zinnia bicolor (DC.) Hemsl. Biol. Centr. Am. Bot. 2: 153.
1881.
Mendezia bicolor DC. in DC. Prodr. 5: 533. 1836.
Zinnia tenella Rob. Proc. Am. Acad. 63: 39. 1907.
Western Chihuahua, Durango (Tejamen, Palmer 500!,
at GH, type of Z. tenella) , southern Zacatecas, northern and
eastern Jalisco, Guanajuato (León, Méndez! in G-DC, the
type), San Luis Potosí.
The ligules of the ray-flowers are white or yellow (the
two colorforms sometimes mixed in the same population),
or yellow with a red spot at base. A plant with yellow
ligules red-spotted at base formed the basis for Z. tenella.
According to Torres Z. tenella differs from Z. bicolor also
in having “lanceolate to elliptic’ rather than “linear to
lance-oblong" leaves, and “obovate to cuneate” rather than
"linear-elliptic" ray-achenes. In Nueva Galicia the differ-
ences in leaf-shape, in achene-shape, and in ray-color,
appear to represent individual variations, and it seems
futile to try to distinguish more than a single species in
the group.
Zinnia violacea Cav. Ic. 1: 57. pl. 81. Dec 1791.
Zinnia elegans Jacq. Ic. P1. Rar. 3: 15. pl. 589. 1793; Coll.
Bot 5: 152 791;
Essentially all authors have taken up the name Zinnia
elegans in preference to Z. violacea. The place of publica-
tion of Z. elegans was correctly cited by a few contemporary
authors, as by Willdenow (Sp. Pl. 3, pt. 3: 2140. 1803)
and by Sims (in Bot. Mag. pl. 527. 1801). In DeCandolle’s
revision of the Compositae, however, through a typo-
516 Rhodora . [Vol. 74
graphical error the reference was given as “Jacq. Coll. 3.
p. 152” (DC. Prodr. 5: 536. 1836). This error has been
perpetuated by subsequent authors who have merely copied
from DeCandolle without looking up the original reference.
Thus in Hemsley’s treatment of Zinnia in the Biologia
Centrali-Americana, and in the Index Kewensis, the cita-
tion is of volume 3 of the Collectanea, not volume 5. As
volume 3 was published late in 1791 (cf. Stafleu, F.,
Taxonomic Literature, p. 232. 1967) it may well have en-
joyed priority over the first volume of Cavanilles' Icones,
which appeared in December of the same year. The fifth
volume of the Collectanea, however, did not appear in
print until 1797. It seems clear that the name Zinnia
violacea Cav. has priority of more than a year over Z.
elegans Jacq., which was published first in 1793 and again
in 1797.
UNIVERSITY HERBARIUM
UNIVERSITY OF MICHIGAN
ANN ARBOR, MICHIGAN 48104
THE COMBINATION PELTANDRA VIRGINICA (L.)
SCHOTT & ENDLICHER: The indigenous eastern North
American genus Peltandra was described by Rafinesque
(J. Phys. Chim. Hist. Nat. Arts 89: 103, 1819) when he
based it on a simultaneously described new species, P. un-
dulata Raf. in a somewhat confusing manner, Rafinesque
states that “Calladium sagittaefolium" (= Arum sagittae-
folium L., a member of the genus Xanthosoma according to
Fernald, Rhodora 50: 59, 1948) and “C. virginicum” (pre-
sumably — Arum virginicum L.) are related to Peltandra,
but he apparently did not intend their inclusion in the
genus. Rafinesque (New Flora and Botany of North Amer-
ica 1: 87, 1836) later conceded that P. undulata was
probably identical to Arum virginicum. However, he re-
1972] Peltandra virginica — Blackwell 517
tained the name P. undulata, failing to employ the correct
combination P. virginica, Index Kewensis wrongly at-
tributes the combination P. virginica to Rafinsque.
Most authors who have dealt with Peltandra virginica
credit the name to Kunth (Enumeratio Plantarum 3: 43,
1841). Included in this list are: Morong (Mem. Torrey
Bot. Club 5: 102, 1894), Tidestrom (Rhodora 12: 48, 1910),
Blake (Rhodora 14: 104, 1912), Small (Manual of the
Southeastern Flora 246, 1933), Barkley (Madrofio 7: 133,
1944), Gleason (New Britton and Brown Illustrated Flora
1: 368, 1952), Huttleston (Taxon 2: 33, 1953), Fassett
(A Manual of Aquatic Plants, second ed., 164, 1957), and
Radford et al. (Manual of the Vascular Flora of the Caro-
linas 257, 1968). On the other hand, Fernald (Rhodora
50: 56, 1948; Gray's Manual of Botany, eighth ed., 383,
1950) and Merrill (Index Rafinesquianus 81, 1949) recog-
nize Scott and Endlicher (Meletemata Botanica 19, 1832)
as having made the combination. Clearly, authorship of
this combination is a point of nomenclatural confusion.
A study of the original literature of concern revealed
that the combination Peltandra virginica was first used
in print by Steudel (Nomenclator Botanicus 1: 603, 1821)
when he mistakenly attributed the combination to Rafin-
esque. Steudel should not receive credit for a new com-
bination in this instance as he considered P. virginica a
member of the genus Arwm, indicating it as a synonym of
Arum virginicum. The only species accepted by Steudel as
belonging to Peltandra is P. undulata. Steudel (Nomen-
clator Botanicus, second ed., 1: 249, 1840) later discon-
tinued recognition of the genus Peltandra and included
it in the synonymy of Caladium. Merrill (1949) notes
Steudel’s (1821) use of the name P. virginica and rightly
does not credit him with the combination. Schott and
Endlicher (1832) accord Peltandra generic status and
accept P. undulata and P. virginica as distinct species,
attributing both names to Rafinesque. Kunth (1841) cites
Schott and Endlicher’s treatment and similarly recognizes
both species, also crediting Rafinesque with the names. At
518 Rhodora [Vol. 74
a considerably later date, Schott (Synopsis Aroidearum 50,
1856) maintains the same opinion concerning the taxa of
Peltandra and their authorship.
Although there can be little or no support for regarding
P. virginica and P. undulata as distinct, Schott and End-
licher (1832) validly make the combination Peltandra
virginica for the first time, even though they erroneously
attribute the combination to Rafinesque. Fernald (1948,
1950) and Merrill (1949) are thus vindicated in their
designation of authorship as Peltandra virginica (L.)
Schott & Endlicher. By the new wording of Recommenda-
tion 46C of the International Code of Botanical Nomen-
clature, as proposed by Yeo (Regnum Vegetabile 60: 62,
1969, Proposal 281) and amended and accepted by the
Eleventh International Botanical Congress (see Taxon 19:
48, 1970), the citation of authorship as Peltandra virginica
(L.) Rafinesque ex Schott & Endlicher would be equally
correct, though in my opinion less desirable since the
ascription to Rafinesque is apparently a mistaken one.
I wish to gratefully acknowledge the American Philo-
sophical Society for grant support of my current systematic
investigations of the genus Peltandra.
WILL H. BLACKWELL, JR.
DEPARTMENT OF BOTANY
MIAMI UNIVERSITY
OXFORD, OHIO 45056
SPONTANEOUS HYBRIDS BETWEEN CERASTIUM
TOMENTOSUM LINN. AND C. ARVENSE LINN.
J. K. MORTON
The “White Rock" or “Snow in Summer" of our gardens
— Cerastium tomentosum Linn. is a native of southern
Europe, now commonly cultivated as a rock garden plant
throughout the temperate world. It is a very vigorous peren-
nial which rapidly spreads and takes over the whole of the
rock garden, hence its rhizomes are frequently dug up and
thrown onto roadsides, river banks and rubbish dumps
where it sometimes becomes established and competes suc-
cessfully with the native flora. C. arvense Linn. is a native
species in North America where the most widely occurring
form is diploid. However the common European form is
tetraploid (2n = 72 as in C. tomentosum) and has been
introduced into northeastern North America where it fre-
quently becomes a weed along roadsides, riverbanks, and
in old pasture and quarries etc. Whether its introduction
to this continent was deliberate or accidental is uncertain,
though the former is not improbable for it makes an at-
tractive rock garden plant, apart from its propensity to
spread too vigorously.
In the mid and upper reaches of the Grand River valley
in southern Ontario, both these species occur; C. tomen-
tosum as a common garden plant persisting in derelict
gardens and occasionally established as a garden throwout;
and C. arvense as a well established weed on road, rail and
river banks etc. Hybrids between the two species have
been discovered at Fergus, Waterloo and Galt. At Fergus
the hybrid plants are growing amongst concrete slabs on a
grassy roadside bank at the Elora end of the town. The
area probably originated as a rockery but has long been
neglected, and both parent species are growing alongside
the hybrids. Further plants of the hybrid occur below the
road, on old ballast used in constructing the road and
dumped on the river bank. At Waterloo the plant was
519
520 Rhodora [Vol. 74
found in an old rockery of a long since derelict farm house,
and has since been transplanted into several rock gardens
in the town. Neither parent now grows in the immediate
vicinity of the old farm house and the nearest colony of
C. arvense is several miles away. At Galt the hybrid plants
are growing amongst grass on cindery ground on an old
rubbish tip, and in the lawn of an adjoining house on the
Brantford side of the town between route 24 and the Grand
River. Neither parent is growing in the immediate vicinty
but C. arvense occurs along the Grand River about a mile
distant.
The Fergus and Waterloo plants appear to be the F;
hybrid and are intermediate between the two parents, but
somewhat closer to C. tomentosum in general appearance,
looking like a rather poorly developed plant of that species
in which the white indumentum is less dense. The Galt
colony apparently consists of F» segregates probably result-
ing from selfing of the F, hybrid. They are smaller, very
variable plants and several of them are nearer to the C.
arvense parent in general appearance.
These hybrids flower copiously and produce a number of
short capsules, the teeth of which barely exceed the calyx.
Most of the ovules in these abort, but frequently one or
two, occasionally more, very large seeds are formed. These
are fertile and the progeny show extensive segregation,
ranging from close to one or other of the parents, to various
combinations of characters intermediate between the
parents. It is probable that these seeds result from selfings
rather than pollination from either parent.
Hybrids between C. tomentosum and C. arvense are
rarely encountered in Europe where the species are native
and sometimes grow in close proximity. The only reference
in the literature to such hybrids, that I have been able to
locate, is Ascherson and Graebner 1919 where they are
referred to C. maureri and C. rigoi without authority or
supporting information. Neither of these names appears
to have been taken up by subsequent workers, perhaps be-
cause of the very rare occurrence of this hybrid.
1972] Cerastium Hybrids — Morton 521
REFERENCE
ASCHERSON, P. and GRAEBNER, P. 1919. Synopsis der Mitteleuro-
paischen flora. 5, 1:636.
DEPARTMENT OF BIOLOGY
UNIVERSITY OF WATERLOO
WATERLOO, ONTARIO, CANADA
AMPHIPRORA ORNATA BAILEY —
A SECOND STATION IN NEW ENGLAND
L. GC. COLT, Jg.
During continuing studies of the phytoplankton of the
Connecticut River I collected a single specimen of Amphi-
prora ornata Bailey. As the accompanying figure shows
the species is distinct enough in structure so as to make
it readily recognizable.
The specimen was recovered from material taken by a
vertical haul with a Wisconsin-type sampler in 46 feet of
water. The collecting station is located 0.1 miles north
of the dam at Vernon, Vermont. All of the stations being
utilized during this study are within five miles of Vernon.
The collection was made just before 12 Noon on June 5,
1970. At this time the air temperature was 68 degrees F.,
and the water temperature was 66 degrees F.
Although the collecting station is physically closer to
Vermont than New Hampshire, ownership of the Con-
nectieut River below high water mark is vested in the
State of New Hampshire. Thus, Amphiprora ornata Bailey
is reported for the Town of Hinsdale, Cheshire County,
New Hampshire.
This appears to be only the second collection for New
England. The previous station being somewhere in the
vicinity of New Haven, Conn., reported by Terry (1907).
Other reported stations include the type from Florida by
522 Rhodora [Vol. 74
I-/2 u~
Amphiprora ornata Bailey. Camera lucida drawing by the author.
1972] Amphiprora — Colt 523
Bailey (1850), Boyer (1827) in Pennsylvania, and Whit-
ford (1969) in North Carolina.
LITERATURE CITED
BAILEY, J. W. 1850. Microscopial observations made in South
Carolina, Georgia, and Florida. Smithsonian Contributions to
Knowledge, Volume II, Art. 8.
Boyer, C. S. 1927. Synopsis of North American Diatomaceae.
Proc. Acad. Nat. Sci. Phila., Vol. 79(2): 229-583.
TERRY, W. A. A partial list of Connecticut diatoms with some
account of their distribution in certain parts of the state. Rho-
dora 8(104): 125-140.
WHITFORD, L. A., & G. J. SCHUMACHER. 1969. A Manual of the
Fresh-water Algae in North Carolina. North Carolina Agric.
Exp. Station Tech. Bull. 188.
BOSTON STATE COLLEGE
BOSTON, MASS. 02115
THE YELLOW MANDARIN,
A NOTABLE ADDITION TO
THE FLORA OF ARKANSAS
PAUL L. REDFEARN, JR.
Although the yellow mandarin, Disporum lanuginosum
(Michx.) Nickols, is widespread in the eastern United
States, it was, according to Robert G. Johnson (personal
communication), not known west of Davidson County,
Tennessee, except for one record of its occurrence in Ar-
kansas which was most likely in cultivation. Dr. E. B.
Smith (personal communication) also informs me that a
single specimen from Texarkana, Miller County, Arkansas,
is present in the herbarium of University of Arkansas and
is marked “probably cultivated." The discovery of a new
locality for the yellow mandarin in Arkansas is therefore
notable for two reasons. First, because of the area where
it was discovered, there can be no doubt now that it is a
native, though rare, member of the flora of Arkansas. Sec-
ond, it adds to the growing list of southern and Appalach-
524 Rhodora [Vol. 74
ian species present in the relic mesophytic forests com-
mon throughout the Boston Mountains (Braun, 1950).
This list includes such vascular plants as Fagus grandi-
flora, Liquidambar styraciflua, Magnolia tripetala and Ly-
copodium lucidulum, and bryophytes such as Hookeria
acutifolia, Brotherella tenuirostris, Schwetschkeopsis fab-
ronia, Sciaromium lescurii, Sphagnum capillaceum var.
tenerum and Bryoxiphium norvegicum. Except for this
last species, which occurs in several nearby ravines, all of
these taxa are located in the immediate vicinity of the hab-
itat of Disporum lanuginosum. Another notable species in
this area is Dodecatheon frenchii. Like D. lanuginosum,
it also occurs along the base of a north-facing sandstone
bluff; otherwise, this species is known from one other lo-
cality in Arkansas (Olah & DeFillipps, 1968) and in the
Ozarks of southern Illinois (Voigt & Mohlenbrock, 1964).
The specific collection data for Disporum lanuginosum
in the Boston Mountains of Arkansas is: Newton County.
On rocky soil at base of north-facing sandstone bluff, upper
reaches of Terrapin Branch, sect. 26, T. 14 N., R. 23 W.,
alt. ca. 700 ft., Redfearn 27381 (ARK, SMS, NCU, UMO).
LITERATURE CITED
BRAUN, E. Lucy. 1950. Deciduous forests of eastern North America.
The Blakiston Co., Philadelphia. p. 170.
MOHLENBROCK, RoBERT H. and Joun W. VoicT. 1959. A Flora
of Southern Illinois. Southern Illinois University Press, Car-
bondale. p. 266.
OLAH, L. V. and R. A. DEFiLLIPPS. 1968. A cytotaxonomic study
of French's shooting star. Bull. Torrey Bot. Club. 95: 168-198.
DEPARTMENT OF LIFE SCIENCES
SOUTHWEST MISSOURI STATE UNIVERSITY
SPRINGFIELD, MISSOURI 65802
RHAPIDOPHYLLUM HYSTRIX
IN MISSISSIPPI
J. RAY WATSON
Rhapidophyllum hystrix (Pursh) Wendland & Drude,
the Needle Palm, is a rare shrub of the southeastern
Coastal Plain. It occurs sporadically from central Florida
and southern Georgia westward to Mississippi (Small,
1923; Harper, 1928; Small, 1933). Recent field studies have
extended its known distribution in Mississippi; herbarium
specimens will be deposited in MISSA.
The northernmost site known in Mississippi is in Lauder-
dale County, approximately 10 miles south of Meridian
(Watson 7314), where it was discovered by Leslie Hu-
bricht. Southward, it has been found in Clarke (Watson
7439) and George (Denier 1273) counties and reaches its
southern limit in Jackson County (Watson 7870). It ex-
tends westward to Simpson County where it was noted by
R. B. Channel in the early 1960’s (personal communica-
tion). A recent search of this site to obtain herbarium
specimens was, however, unsuccessful. Recently it has been
found in Forrest County (Watson 8115, October 25, 1969),
approximately 60 miles east of the Simpson County local-
ity.
Within the state, the Needle Palm is known to occur
either along the flood-plains of small streams or along the
drainages of rich, wooded ravines.
LITERATURE CITED
Harper, R. M. 1928. Economic Botany of Alabama, Part 2. Cata-
logue of the trees, shrubs and vines of Alabama, with their
economic properties and local distribution. Geol. Surv. Ala.
Monogr. 9.
SMALL, J. K. 1923. The needle palm — Rhapidophyllum hystrix.
Jour. N. Y. Bot. Gard. 24: 105-114.
1933. Manual of the Southeastern Flora. The author,
New York.
DEPARTMENT OF BOTANY
MISSISSIPPI STATE UNIVERSITY
STATE COLLEGE, MISSISSIPPI 39762
525
EQUISETUM FROM PENINSULA FLORIDA
DON R. REYNOLDS AND KEN CALHOUN
The first report of Equisetum in Florida was made in
1932 when E. praealtum was discovered in Northern
Florida (Small, 1938). E. robustum is recorded from 1929
growing along the Apalachicola River at Alum Bluff. Cor-
rell (1938) lists E. praealtum as occurring along wet sandy
stream banks in Liberty County, Florida. Schaffner (1939)
- maps the locality of Equisetum in Florida as occurring in
the pan-handle of the State. Wherry (1961) lists E.
hyemale var. elatum as occurring in Gadston county along
the Georgia border and westward to Texas. He lists the
habitat as, “sandy shores and seemingly barren areas
where moisture and nutrients can be found at depth. In-
vading disturbed soil, fills . . ." Heretofore Equisetum
is known only from the pan-handle area of Northwest
Florida. We wish to report Equisetum from peninsula
Florida.
Two collections have been identified as Equisetum
hyemale var. affine, following Hauke (1963). This determi-
nation has been confirmed by a comparison with the speci-
men of G. Engelmann in the Missouri Botanical Garden, St.
Louis, which is the type of this variety.
SPECIMENS EXAMINED: Florida Technological Uni-
versity Herbarium:
8 February 1971. Ken Calhoun and Don R. Reynolds,
Brooksville Limestone Quarry, Liberty County, Florida.
The specimens were growing in moist clay in several places
in the quarry. The quarry is approximately 14 mile long
and 300 yards wide and 50 feet deep. It was abandoned in
1967. There was little competition from other plants for
the large Equisetum clones which flourished along a large
water filled pit that ran the length of the quarry. The
area immediately surrounding the quarry and other likely
areas within a 25 mile radius were searched on several
526
1972] Equisetum — Reynolds and Calhoun 527
occasions. Although several apparently suitable habitats
were located, Equisetum seems to be restricted to the con-
fines of the Brooksville Quarry.
9 May 1971. Don R. Reynolds and Ken Calhoun, Sand
Quarry, approximately 10 miles east of Clermont on US
Highway 50, Lake County, Florida. The specimens were
growing in sandy soil which was intermittently wetted
from water being pumped into the quarry as part of the
dredging operation. The quarry covered about one square
mile; a large part was covered with water. No vegetation
was growing in the immediate proximity of the Equisetum
clone.
Hauke (1963) has demonstrated the clinal variation be-
tween the southern states of E. hyemale var. affine, based
on teeth retention, height of plants and ridge angularity.
The clines of Mississippi, Alabama and Northern Florida
are similar to the clines of Georgia except that the per-
centage of teeth retention in the Georgia clines is much
lower. According to the percentage of teeth retention in
the two specimens cited (80-100%), the Equisetum from
Liberty and Lake Counties represent a southerly extension
of the Mississippi, Alabama, North Florida clines rather
than the Georgia one.
A third collection of Horsetail was recently given to Dr.
Robert Long (Biology Department, University of South
Florida) which was found growing in a ditch in Dunedin,
Pinellis County, Florida. Dr. Long (personal communica-
tion) identified the plants as Equisetum hyemale. This
collection represents the southernmost record on the Florida
peninsula.
LITERATURE CITED
CoRRELL, D. S. 1938. A county check-list of Florida ferns and fern
allies. American Fern Journal 28: 11-15.
HAUKE, RICHARD. 1963. A taxonomic monograph of the genus
Equisetum subgenus Hippochaete. J. Cramer. Stuttgart. 123 p.
SCHAFFNER, J. H. 1939. Distribution of exclusively North American
species of Equisetum. American Fern Journal 29: 45-47.
528 Rhodora [Vol. 74
SMALL, J. K. 1938. Ferns of the southeastern states. Hafner Co.
N. Y. Reprint Edition. 517 p.
Wuerry, E. T. 1961. The southern fern guide. Southern, eastern
and midland United States and adjacent Canada. Doubleday and
Co., Garden City, N. Y. 318 p.
DEPARTMENT OF BIOLOGICAL SCIENCES
FLORIDA TECHNOLOGICAL UNIVERSITY
ORLANDO, FLORIDA 32816
WIND AND THE WINTER-EXPOSED PLANT. In his
paper “Snow Cover and the Diapensia lapponica habitat in
the White Mountains, New Hampshire” (Rhodora 74: 358-
377), W. N. Tiffney, Jr. makes repeated reference to the
prevalence of high winds in the alpine zone which, in his
belief, *. . . promote desiccation in winter-exposed plants".
The idea that winter desiccation is significantly influenced
by high winds is shared by at least one other ecologist (Lind-
say, 1971), but this conclusion seems to be based on the
application of summertime wind effects to the wintertime
situation. The energy budget of the winter-exposed plant
is, however, quite different from that of the more freely
transpiring plant in a summer microclimate, and the preva-
lence of high winter winds may instead forestall damaging
water loss.
Transpiration from a leaf surface, at any time, is directly
proportional to the water vapor concentration gradient be-
tween the leaf and air, and is inversely proportional to the
diffusive resistances offered by the leaf and boundary layer
of air adjacent to the leaf (Gates, 1965). Hence T DL 7 =
1 a
where T is the transpiration rate, c; — c, is the water vapor
concentration difference between the intercellular spaces of
the leaf and the bulk air outside the leaf boundary layer,
and r, and r, are respectively the leaf and boundary re-
sistance to gaseous transfer. It should be noted that the
driving force for transpiration, c; — Ca, is strongly influenced
by the temperature difference between the leaf and air,
since saturation vapor pressure is a function of tempera-
ture. The greater the elevation of leaf temperature above
1972] Winter-exposed Plants — Marchand 529
air temperature, the greater the vapor concentration gradi-
ent between the leaf and air.
Leaf resistances to the diffusion of water vapor are pro-
vided by the stomates and cuticle, the relative magnitude
of these being somewhat species dependent. The minimum
stomatal resistance of the alpine species Ledum groenlandi-
cum is for, example, less than 2 sec cm-t, as determined for
plants growing in both a northern bog (Small, 1972) and
above timberline on Mt. Washington, New Hampshire
(Marchand, unpublished data). The stomatal resistance
of conifers may be somewhat higher, perhaps near 20 sec
cm- (Waggoner and Turner, 1971). In contrast, cuticular
resistance is usually several times greater in magnitude,
having been reported (Holmgren et al., 1965) as high as
460 sec cm"! for the European species Quercus robur. In
addition, cuticular resistance has been found to increase
sharply with decreasing temperature (Holmgren et al.,
1965).
Boundary layer resistance is provided by a transfer zone
of air in contact with (and influenced by) the leaf. The
thinner this surrounding layer, the more rapid will be the
heat convection or vapor transfer through this zone, since
heat and vapor concentration gradients between the leaf
and bulk air will be steeper. Single leaf boundary layer
resistances have been found to be generally less than 0.9
sec cm-! for severa] deciduous woodland species (Holmgren
et al., 1965), although this will vary directly with leaf size
and shape and inversely with windspeed (Gates, 1965).
It is through reduction of boundary layer resistance that
the influence of wind currents is important. The effect of
wind is two-fold; (1) it facilitates removal of moist air
from the leaf surface, thereby increasing the rate of trans-
piration, and (2) it increases the rate of heat transfer from
the leaf, through forced convection, thus tending to main-
tain temperature equilibrium between the leaf and air. In
the latter case, c; — c, is reduced and consequently transpira-
tion is decreased. The relative importance of these two
processes will depend on other microenvironmenta] factors
as well as on the physiological behavior of the plant.
530 Rhodora [Vol. 74
During the summer growth period, the most significant
leaf resistance to loss of water (under non-stress condi-
tions) is that of the open stomates. Cuticular transpiration
is negligible as long as the stomates remain open. At this
time of year then, the boundary layer resistance is closer in
magnitude to the leaf resistance and any reduction of r,
by turbulent exchange becomes significant in terms of in-
creasing transpiration. Under these circumstances, the ef-
fect of wind on leaf temperature seems to be of minor
importance.
In the wintertime, the relative importance of the diffusive
resistances is changed significantly. Stomatal opening has
not been reported to occur during the winter (Tranquillini,
1964; Schulze et al., 1967) as apparently it is prevented by
low temperatures (Staefelt, 1962). As a result, vapor dif-
fusion is largely via cuticular pathways and the leaf re-
sistance thus becomes very high.* It is on this point that
my argument is based. As seen from the above transpiration
model, when r, is high, r, becomes negligible and any re-
duction of the boundary layer resistance by high winds is
unimportant in terms of increasing vapor transfer. So the
dominant effect of wind in the wintertime is, in my opinion,
related to the consequences of forced convection, and these
are (1) maintenance of leaf temperatures below the freez-
ing point of cell water when air temperatures are very low,
and (2) reduction of temperature differences between the
leaf and air with consequent reduction of e, — ca. In the first
case, water loss will be limited to the process of sublimation
from frozen tissues, requiring greater energy input than
for the evaporation of free water. In both cases, the net
effect of wind would be to reduce, rather than increase
water loss.
Where reference has been made to the work of Tranquil-
lini (1964) and Sakai (1970), citing conditions of winter
drought and desiccation damage in alpine areas, it should
*The discussion here is pertinent only to evergreen species, and
since heavy cutinization is characteristic of evergreens, the value of
cuticular resistance is assumed to be very high relative to r..
1972] Winter-exposed Plants — Marchand 581
be noted that such conditions are the combined result of
frozen soils and the strong heating of plant parts above
freezing, due to high direct and reflected radiation load on
exposed plants. Thus it is most likely that winter-exposed
plants will experience damaging water loss on days which
are clear and windless. Whether or not winter desiccation
in the alpine areas of the White Mts. of New Hampshire
can be substantiated, the conclusion that “. . . low tempera-
tures and high winds combine to promote severe desiccation
in this snow-free area" seems unwarranted on the basis of
simple physical considerations.
LITERATURE CITED
Gates, D. M. 1965. Energy, plants, and ecology. Ecology 46: 1-13.
HOLMGREN, P., P. G. JARVIS, and M. S. JARVIS. 1965. Resistances
to carbon dioxide and water vapor transfer in leaves of different
plant species. Physiol. Plant. 18: 557-573.
LINDSAY, J. H. 1971. Annual cycle of leaf water potential in Picea
engelmannii and Abies lasiocarpa at timberline in Wyoming.
Arctic and Alpine Res. 3: 131-138.
SAKAI, A. 1970. Mechanism of desiccation damage of conifers win-
tering in soil-frozen areas. Ecology 51: 657-664.
ScHULZE, E. D., H. A. Mooney, and E. L. DUNN. 1967. Wintertime
photosynthesis of bristlecone pine (Pinus aristata) in the White
Mountains of California. Ecology 48: 1044-1047.
SMALL, E. 1972. Water relations of plants in raised sphagnum peat
bogs. Ecology 53: 726-728.
STALFELT, M. G. 1962. The effect of temperature on opening of the
stomatal cells. Physiol. Plant. 15: 772-779.
TIFFNEY, W. N., JR. 1972. Snow cover and the Diapensia lapponica
habitat in the White Mountains, New Hampshire. Rhodora 74:
358-377.
TRANQUILLINI, W. 1964. The physiology of plants at high altitudes.
Ann. Rev. Plant Physiol. 15: 345-362.
WAGGONER, P. E. and N. C. TURNER. 1971. Transpiration and its
control by stomata in a pine forest. Conn. Agr. Exp. Sta., Bull.
726, 87 p.
PETER J. MARCHAND
DEPARTMENT OF BOTANY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM 03824
XII INTERNATIONAL BOTANICAL CONGRESS
JUNE 23-30, 1975
The Closing Plenary Session of the XI International Bo-
tanical Congress held at Seattle, U.S.A., in 1969, accepted
an invitation issued by the Academy of Sciences of the
U.S.S.R. to convene the XII International Botanical Con-
gress in the City of Leningrad in 1975. In 1971 the Organiz-
ing Committee was appointed: consisting of a chairman
(A. L. Takhtajan) ; four vice-chairmen (A. A. Prokofiev,
A. A. Theodorov, N. V. Tsitsin, A. A. Yatsenko-Khmelev-
Sky) ; a secretary-general (O. V. Zalensky) ; a scientific sec-
retary (N. S. Snigirevskaya) ; and a number of members
at large. The XII International Botanical Congress is in-
tended to facilitate interdisciplinary communication among
botanists as well as an informal exchange of ideas. A num-
ber of sections are planned, including special ones to accom-
modate mycologists (also lichenologists), phycologists, and
bryologists.
The Congress will be divided between organized half-day
symposia and half-day contributed paper sessions. In ad-
dition to the opening and closing plenary sessions, two even-
ing lectures are being scheduled. All special interest groups
wishing to apply for space and time during the Congress
should do so by writing as soon as possible to the secretary-
general, Dr. Oleg Zalensky, Komarov Botanical Institute of
the Academy of Sciences of the U.S.S.R., 2, Popov Street,
Leningrad 197022, U.S.S.R.
The sessions of the Nomenclature Section will take place,
as usual, immediately before the opening of the Congress —
in this instance June 20-23. Four days are set aside in
order to enable the Section to convene for six to eight ses-
sions of two to four hours each.
A meeting of the International Association of Botanic
Gardens (President Academician N. V. Tsitsin) will be held
in Moscow at the Main Botanical Garden of the Academy of
Sciences of the U.S.S.R. on June 20.
A tentative schedule of scientific field trips has been
532
1972] Botanical Congress — Takhtajan 533
planned for the immediate pre-Congress and post-Congress
periods. The principal purpose of these trips is to acquaint
visiting botanists with as many interesting and unique fea-
tures of the flora and vegetation of various regions of the
U.S.S.R. as possible. Some specialized trips for phycologists,
lichenologists, bryologists, and palaeobotanists are also
planned.
The double postcards announcing the XII International
Botanical Congress will be mailed during the last months
of 1972. Those who wish to receive further information on
the Congress should return their interest cards by March 1,
1973, so that they will be placed on the mailing list for the
First Information Circular expected to be published June-
July 1973.
CHAIRMAN, ORGANIZING COMMITTEE
A. TAKHTAJAN
REQUEST FOR UNNEEDED COPIES OF RHODORA
FOR DECEMBER 1971, NO. 796
The error on the cover of the fourth number of Rhodora
for 1971, indicating it as September 1971, no. 795, has
resulted in many persons discarding it as a duplicate. Had
they looked inside the cover they would have detected the
error immediately. However so many requests have come
in for replacements that Dr. Herman Sweet requests that
any subscribers who have this December issue, and do not
regularly keep a file of Rhodora, send this particular copy
to him at the Botanical Museum, Oxford Street, Cambridge,
Mass. 02138.
Dovora
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by
ALBION REED HODGDON, Editor-in-Chief
ALBERT FREDERICK HILL
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON >Associate Editors
RADCLIFFE BARNES PIKE
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
VOLUME 74
1972
Che Nef England Botanical Club, Ine.
Botanical Museum, Oxford St., Cambridge, Mass. 02138
INDEX TO VOLUME 74
New scientific names and combinations are printed in bold face type
Abies balsamea 367
Abronia Fendleri 83
Acalypha rhomboidea 392
Acanthospermum australe 203;
humile 205
Acer rubrum, var. drummondii
147; f. tomentosum 147; sac-
charinum 392
Acrochaetium catenulatum 92, 93;
compactum 92; crassipes 92,
93; daviesii 268; microfilum 93
Acroptilon picris 344; repens 344
Actinella argentea 84; Richard-
sonii, var. floribunda 84
Actinomeris heterophylla 97-100;
pauciflora 97-100
Adams, Preston, Studies in the
Guttiferae III. An Evaluation
of some Putative Spontaneous
Garden Hybrids in Hypericum
sect. Myriandra 276
Aesculus glabra 147; var. leuco-
dermis 147
Aethusa cynapium 156
Agalinis heterophylla 387
Agardhiella tenera 480
Agarum cribrosum 267, 416
Ageratella microphylla 497; var.
palmeri 497; var. seemannii
497; palmeri 497
Ageratum microphyllum 497
Agoseris purpurea 84
Ahnfeltia plicata 269, 409, 421
Alaria esculenta 267, 411, 416
Alcina minor 176; ovalifolia 175;
ovatifolia 175; perfoliata 12,
194, 195
Algae of Kent Island, Bay of
Fundy. On the Marine 264
Algal Invaders of the Northeast-
ern Fringes of the Sargasso
Sea, Some 295
Algal Vegetation of the Hamp-
ton-Seabrook Estuary and the
Open Coast Near Hampton,
New Hampshire, The 406
Alsophila Abbottii 445; Brooksii
445; bryophila 443-445; capen-
sis 445; confinis 445; dryopte-
roides 445; Engelii 445; hot-
teana 445; minor 445, 446;
Nockii 445; pauciflora 445;
sagittifolia 446; Salvinii 445;
Urbanii 445
Alyssum alynoides 391; deserto-
rum 381, 391
Amalias anceps 283
Amaranthus caudatus 143
Amberboa moschata 344; muri-
cata 339
Ambrosia tomentosa 394
Amphiprora ornata 521, 522;
Bailey, A Second Station in
New England 521
Andropogon intermedius 378
Anemone multifida 391
Anoda cristata 383
Antithamnion floccosum 421
Apocynum medium 385
Aquatic Plants in New England,
Range Extensions of Vascular
131
Arabis Fendleri 83; glabra 391
Arceuthobium cryptopodium 83
Archibaccharis hieraciifolia 498,
511; hieracioides 498
Arenaria groenlandica 367-370,
372-374
Aristida Fendleriana 83; longi-
seta 83
537
538
Artemisia annua 388; arctica 110,
111; borealis 110; frigida 110;
norvegica 111; saxicola 102, 110
Arum sagittaefolium 516; virgin-
icum 516, 517
Asclepias incarnata, var. incar-
nata, f. albiflora 385
Ascophyllum mackaii 286; nodo-
sum 268 295, 297, 411, 416,
418, 425, 481, 483; f. scorpi-
oides 411, 416, 418, 482; ecad.
Mackaii 130
Ascyrum hypericoides, var. hy-
pericoides 145; var. multicaule
145
Asperococcus echinatus 267
Aster adscendens 104; chilensis
104; ciliolatus 394; ericoides
104; exilis 104; foliaceus 104;
hirtifolius 104; lateriflorus 104;
occidentalis 104; puniceus 104;
sagittifolius 394
Asteraceae, Somatic Chromo-
some Numbers for Some 102
Asterocytis ramosa 421, 427, 428
Astragalus ceramicus 83; loncho-
carpus 83; lotiflorus 392;
purshii 391
Atriplex glabriuscula 390; has-
tata 391; heterosperma 390;
subspicata 391
Atwood, N. Duane, New Species
in the Phacelia crenulata
Group (Hydrophyllaceae) 451
Audouinella catenulatum $85, 92,
96; crassipes 85, 86, 96; mem-
branacea 421; microfilum 85,
89, 92, 96; microscopica 85-93,
96; (Naeg.) Woelk. Complex
(Rhodophyta), Studies on 85
Austin, Daniel F. Interactions
between Apis mellifera (Hy-
menoptera: Apidae) and Tribu-
lus cistoides (Zygophyllaceae)
117
Baccharis hieracifolia 498; hie-
Rhodora
[Vol. 74
raciifolia 498; potosina 499;
sulcata 499; thesioides 499
Bangia ciliaris 421; fuscopur-
purea 413, 421
Beardsley, Richard L. and Edw.
T. Browne, Jr., Lathyrus
aphaca L. New to Tennessee
and the Southwest 155
Berberis repens 391
Bidens acrifolia 499; acuta 394;
comosa 388, 394; polyglossa
499; tenuisecta 84; vulgata 394
Blackwell, Will H., The Combina-
tion Peltandra virginica (L.)
Schott & Enlicher 516
Blidingia marginata 412; minima
412
Botanical Congress, XII Interna-
tional 532
Botrychium lunaria 389
Bowdoin College, The Parker
Cleaveland Herbarium of 80
Brasenia schreberi 137
Bromus squarrosus 389
Brotherella tenuirostris 524
Browne Edw. T., see Beardsley,
Richard L.
Bryopsis plumosa 412, 480
Bryoxiphium norvegicum 524
Bulbostylis capillaris 294
Camutia perfoliata 195
Caragana arborescens 392
Calendula americana 511, 512; de-
cumbens 21, 206
Calhoun, Ken, see Reynolds, Don
R.
Calladium sagittaefolium 510;
virginicum 516
Callithamnion baileyi 421; corym-
bosum 421
Callitriehe terrestris 383
Calochortus madrensis 351; venu-
stulus 350, 351; (Liliaceae),
Variation Within 350 var. im-
bricus 350, 351; var. madrensis
351; var. venustulus 350, 351
1972]
Calothrix confervicola 484; crus-
tacea 476, 478 485-487
Cape Cod Species of Carex, The
Rachilla in, With Notes on the
History of the Perigynium and
Rachilla 321
Capsosiphon fulvescens 481, 484
Carex acuta 324; alopecoidea 390;
bigelowii 367, 370, 372-375; bul-
lata 323; buxbaumii 390; Cam-
posii 323; capillaris 390;
canescens 323; complanata 323,
327; crinita 326; davisii 390;
debilis, var. Rudgei 321, 323;
elachycarpa 329; Emmonsii
321, 323; flacca 324; flava 324;
Fraseri 323; gracillima 390;
hirta 324; hystricina 149; lacus-
tris 390; lasiocarpa 390; limosa
390; melanostachya 379; micro-
glochin 322, 329, 330; Oederi
324; oedipostyla 328; pallescens
323-325; var. glaberrima 325;
var. neogaea 325; pensylvanica
321, 323; pseudocyperus 324;
scoparia 321, 323; Sprengelii
326; squarrosa 148; stipata
390; Swanii 321; tonsa 323;
vesicaria 390; Willdenowii 325
Carex, the Rachilla in Cape Cod
Species of, with Notes on the
History of the Perigynium and
Rachilla 321
Catling, P. M., see Cruise, J. E.
Cedrela discolor 125; oaxacensis
124-126; C. DC. & Rose, Re-
definition of 124; poblensis
126; salvadorensis 124, 126;
saxatilis 126; tonduzii 124 126
Centaurea americana 337, 344;
austriaca 336, 340, 341; calci-
trapa 334, 338, 339, 345; calci-
trapoides 338; cineraria 337,
344; consimilis 340; cyanus 337,
344, 345; var. denudata 344;
diffusa 335, 340, 345; diluta
Index to Volume 74
539
332-334, 339; dubia 337, 343,
345; ssp. eu-dubia 343; erio-
phora 332; iberica 334, 338;
jacea 335, 340-342, 345; var.
nigra | 340; var. transal-
pina 343; macrocephala 387,
343; maculosa 335, 339, 340,
345, 394; ssp. maculosa 339;
ssp. micranthos 339; ssp. rhe-
nana 339; melitensis 333, 358;
montana 337, 344; moschata
337, 344; muricata 334, 339;
nervosa 332, 336, 340, 341;
nigra 336, 340-342, 345; f. pal-
lens 342; nigrescens 337, 342,
343, 345; var. dubia 343; panic-
ulata 335, 339, 340; picris 344;
phrygia, ssp. austriaca 341; X
pratensis 336, 340-342, 345;
repens 337, 344, 345; rothrockii
337, 344; salmantica 333, 334,
339; scabiosa 337, 344; sicula
338; solstitialis 150, 334, 338,
345; squarrosa 340; stoebe 339;
sulphurea 334, 338; transalpina
342, 343; virgata, var. squar-
rosa 335, 340; vochinensis 342
Centunculus minimus 384
Ceramium diaphanum 480; fasti-
giatum 480; rubriforme 480;
rubrum 269, 422, 480; strictum
422
Cerastium arvense 519, 520; glo-
meratum 381; maureri 520;
rigoi 520; tomentosum 519,
520; Linn. and C. Arvense
Linn., Spontaneous Hybrids be-
tween 519
Chaenactis douglasii 394
Chaenorrhinum minus 394
Chaetomorpha area 266, 412;
atrovirens 412, 413; linum 412,
413, 479; melagonium 266, 413,
A477
Chaetopteris plumosa 420
540 Rhodora
Chantransia collopoda 89, 92;
mediterranea 92, 93; minutis-
sima 92, 93; moniliformis 92;
secundata 88; trifila 92, 93
Chaptalia ruminata 499
Cheilanthes Fendleri 82
Chloris virgata 389
Chondrus crispus 411, 415, 419,
422, 423, 480, 481, 483
Chorda filum 129, 417, 428, 480;
tomentosum 417
Chordaria flagelliformis 267, 297,
417
Chorispora tenella 391
Chromosome Numbers from some
Asteraceae, Somatic 102
Chrysopsis fulcrata 102, 104,
105; hispida 105; mariana 152;
In New England 152; scabra
511; villosa 105
Chrysothamnus nauseosus, ssp.
nauseosus 105
Cinna arundinacea 389
Cladonia alpestris 261; apodo-
carpa 256; bacillaris 255; f.
attenuata 255; f. bacillaris 255;
f. fruticaulescens 255; f. ob-
tusa 255; f. peritheta 255;
beaumontii 259; bonyi 261; f.
lacrimosa 261; f. prolifera 261;
caespiticia 259; f. epiphylla
259; carassensis 259; f. digres-
sa 259; f. obliqua, m. specta-
bilis 260; f. regularis 259; f.
spectabilis 259; f. subirregu-
laris 260; f. subregularis 259;
f. subregularis m. spectabilis
260; capitata 256; f. capitata
256; cariosa 257; f. squa-
mulosa 257; caroliniana 261;
cenotea 261; f. cenotea 261;
f. exaltata 261; chlorophaea
258; f. costata 258; f. pro-
lifera 258; f. simplex 258;
clavulifera 256; coccifera 255;
var. frondescens 255; var.
[Vol. 74
stemmatina 255; coniocraea
258; f. actinota 258; f. cera-
todes 258; f. peritheta 258; f.
phyllostrota 258; f. pycnothe-
liza 258; f. truncata 258;
crispata, f. crispata 261; f.
elegans 261; f. infundibulifera
261; f. subascypha 260; var.
dilacerata 260; var. divulsa
261; var. virgata, f. kairamoi
261; cristatella 255; f. auran-
tiaca 255; f. cristatella 256; f.
lepidifera 256; f. ochrocarpa
256; f. pleurocarpa 256; f.
ramosa 256; f. squamosissima
256; f. squamulosa 256; f. ves-
tita 256; deformis, f. crenulata
255; didyma, var. muscigera
256; fimbriata, f. conista 258;
f. pycnotheliza 258; f. simplex
258; f. stenoscypha 258; floe-
rheana, f. carcata 255; f. floe-
rheana 255; f. intermedia 255;
f. squamosissima 255; var. in-
termedia, f. fastigiata 255; f.
peritheta 255; floridana 259;
furcata 259; f. corymbosa 259;
f. fissa 259; f. foliolosa 259; f.
furcata 259; f. rigidula 259; f.
subclausa 259; f. subulata 259;
f. turgida 259; var. furcata
259; var. palamaea 259; var.
paradoxa 259; var. pinnata,
f. recurva 259; glauca 261; gra-
cilis, var. dilitata 257; f. dila-
cerata 257; var. elongata, f.
elongata 257; grayi 258; f. cen-
tralis 258; f. clavata 258; f.
cyathiformis 258; f. epiphylea
258; f. foliosa 258; f. pallida
258; f. peritheta 258; f. proli-
fera 258; f. squamulosa 258;
impexa, f. impexa 261; f. proli-
fera 261; f. subpellucida 261;
incrassata 255; f. clavata 255;
f. squamulosa 255; leporina
1972]
256; macilenta 256; f. maci-
lenta 256; mateocyatha 257; f.
squamulata 257; mitis 262; f.
divaricata 262; f. pallida 262;
f. prolifera 262; f. prostrata
262; soralifera 262; f. tenuis
262; multiformis 259; nem-
oxyna 258; f. ambigua 258;
nemoxyna 258; f. phyllocephala
258; f. scyphifera 258; papil-
laria 255; f. molariformis 255;
f. papillaria 255; f. stipata
255; parasitica 259; f. congesta
259; f. parasitica 259; f. sterilia
259; piedmontensis 256; f. in-
termedia 256; f. lepidifera 256;
f. obconica 256; f. phyllocoma
256; f. piedmontensis 256; f.
squamossisima 256; pityrea,
var. pityrea 257; f. cladomor-
pha 257; f. pityrea 257; f.
scyphifera 257; f. squamulifera
257; f. subacuta 257; pleurota
255; poculifera 262; f. simplex
262; polycarpoides 256; f. epi-
phylla 256; f. polycarpoides
257; f. squamulosa 257; pseudo-
phyxidata 262; f. clavata 262;
f. lateralis 262; f. prolifera
262; f. simplex 262; f. squamu-
losa 262; pyxidata 257; f.
centralis 257; f. homodactyla
258; f. lophyra 258; f. margin-
alis 257; f. prolifera 257; var.
pyxidata 257; pyxioides, f.
simplex 262; rangiferina 262;
f. incrassata 262; f. tenuior
262; robbinsii, f. squamulosa
250; scabriuscula 259; f. ad-
spersa 259; f. elegans 259; f.
farinacea 259; f. scabriuscula
259; f. squamulosa 259; simu-
lata 257; spumosa 262; squa-
mosa 260; carneopallida 260; f.
denticollis 260; f. fascicularis
260; f. laevicorticata 260; f.
Index to Volume 74
541
muricella 260; f. pityrea 260;
f. pseudocrispata 260; f. squa-
mosa 260; f. subtrachynella
260; f. turfacea 260; f. ven-
tricosa 260; var. laevicorticata,
f. degenerascens 260; var.
muricella, f. sarmentosa 260;
var. phyllocoma 260; strepsilis
256; f. abortiva 256; f. coral-
loidea 256; f. glabrata 256; f.
strepsilis 256; subsquamosa, f.
degenerata 260; f. luxurians
260; f. scabrida 260; f. sublevis
260; f. subrigida 260; subtenuis
261; sylvatica 262; f. arbuscula
262; f. congesta 262; m. fissa
262; f. decumbens 262; f. graci-
lior 262; f. pygmaea 262; f.
scabrida 262; f. sphagnoides
262; m. sorediata 262; tenuis
261; f. prolifera 261; f. pro-
strata 261; terrae-novi 261;
turgida 258; f. conspicua 258;
uncialis, f. turgescans 261; f.
humilior 261; f. obtusata 261;
f. setigera 261; f. soraligera
261; f. subobtusata 261; f. un-
cialis 261; verticillata 257; f.
aggregata 257; f. apoticta
257; f. pallida 257; f. phyllo-
cephala 257; f. verticillata
257; var. cervicornis, f. phyl-
lophora 257
Cladoniae of Southeastern Mas-
sachusetts with Special Refer-
ence to the Collection of
Charles Albert Robbins, List
of the 254
Cladophora gracilis 266; holsat-
ica 130; liniformis 280; re-
fracta 266; rupestris 266; seri-
cea 409, 411, 413, 415, 421
Clathromorphum circumscriptum
422
Cleaveland Herbarium of Bow-
doin College, The Parker 80
542
Cnemidaria decurrens 442
Codiolum pusillum 413
Coleman, James R., Nomencla-
tural Clarification of Two Spe-
cies of Verbesina (Compo-
positae) Endemic to Florida
97
Collinsia parviflora 393
Colt, L. C., Jr., Amphiprora or-
nata Bailey — A Second Sta-
tion in New England 521
Compositae, Nomenclatural and
Taxonomic Notes on Mexican
495
Conium maculatum 397
Conopholis americana 141
Conyza lyrata 500; var.
500; viscosa 500
Corallina officinalis 269, 411, 422,
423
Corallorhiza maculata 141
Coriaria atropurpurea 249; (Cori-
ariaceae) in the Western Hem-
isphere, The Genus 242; cune-
ifolia 250; japonica 246; mi-
crophylla 249, 250; myrtifolia
244, 246; nepalensis 246; phyli-
cifolia 249; ruscifolia 245, 246,
248; ssp. microphylla 245, 249;
ssp. ruscifolia 246, 248; ter-
minalis 245, 247; thymifolia
249, 250
Crepis capillaris 394; tectorum
394
Critonia hebebotrya 503
Croton capitatus 382; glandulo-
sus, var. septentrionalis 382;
monanthogynus 382; texensis
144
Cruise, J. E. and P. M. Catling,
A White-flowered Form of Iris
lacustris from Ontario 271
Cyanus segetum 344
Cyathea panamensis 450; pubes-
cens 445; Steyermarkii 448-
450
pilosa
Rhodora
[Vol. 74
Cyatheaceae, New Species of
American 441
Cyclodium meniscioides 447
Cymopterus Fendleri 83
Cynosciadium digitatum 148
Cyperus diffusus 328; esculentus
143; engelmannii 142; erythro-
rhizos 142; Fendlerianus 83;
ferruginescens 143; Houghtonii
294; iria 142; lancastriensis
142; strigosus 143, 294
Cystoclonium purpureum 269;
var. cirrhosum 422
Daphnidostylus Fendleriana 83
Decachaeta ovatifolia 504, 506
Dermatolithon pustulatum 422
Desmarestia aculeata 129, 267,
417; viridis 267, 411, 417
Desmodium marilandicum 292
Diapensia lapponica 359, 361,
363-365, 367, 368, 370, 371,
373-375, 528; Habitat in the
White Mountains of New
Hampshire, Snow Cover and
358
Dictyosiphon foeniculaceus 267,
417, 428
Diplopappus ericoides, var. hir-
tella 84
Disporum lanuginosum 523, 524
Distichlis spicata 411, 476
Dodecatheon frenchii 524; radi-
catum 83
Draba lanceolata 381; neomex-
icana 83
Dumontia incrassata 269, 423,
480, 481
Duncan, Wilbur H. and Marie
Mellinger, Edgeworthia (Thy-
melaceae) New to the Western
Hemisphere 436
Dunn, David B., Lupinus mexi-
canus Cerv. ex Lag. 489
Dysodium divaricatum 12,
175, 176; radiatum 176
162,
1972]
Eastman, Lesley M., A Second
Occurrence for Triphora trian-
thophora (SW.) Rydb. in
Maine 141
Eaton, Richard, Chrysopsis ma-
riana in Nw England 152
Eclipta pusilla 60
Ectocarpus confervoides 266, 417;
var. dasycarpa 481; siliculosis
417, 418; tomentosus 421
Edgeworthia albiflora 438; chry-
santha 436-438; gardneri 438;
longipes 438; papyrifera 437;
(Thymelaeaceae) New to the
Western Hemisphere 436; to-
mentosa 437
Egeria densa 135, 140
Elachistea fucicola 267, 418, 428;
lubrica 297
Elaeagnus angustifolius 148; um-
bellatus 148
Eleocharis quadrangulata,
crassior 378
Elephantopus carolinianus 103
Eleutheranthera ruderalis 206;
divaricata 177
Elodea Nuttallii 136, 137
Elyna spicata 325
Empetrum nigrum 367
Enteromorpha ahleriana 479,
482; clathrata 265, 479; com-
pressa 265, 413; erecta 265,
411, 413, 414; flexuosa, ssp.
flexuosa 429; ssp. paradoxa
482; ssp. pilifera 479, 480; in-
testinalis 265, 409, 411, 413,
479-481; linza 265, 414, 479;
var, oblanceolata 479, 480;
marginata 265, 412; minima
266, 412; plumosa 414; pro-
lifera 266, 414
Epifagus virginiana 141
Equisetum from Peninsula Flor-
ida 525
Equisetum hyemale 324, 527;
var. affine 526, 527; var. ela-
var.
Index to Volume 74
543
tum 526; palustre 389; praeal-
tum 526; robustum 526; scir-
poides 389
Eragrostis spectabilis 389
Erigeron affine 500; alamosanum
501; canbyi 395; cinereus 83;
longipes 500; macranthus 106;
scaposum 500; Jf latifolium
500, 501; speciosus 106; var.
macranthus 106; tenellum 501;
velutipes 501
Eriogonum atrorubens 351; gor-
donii 391; harperi 154; Good-
man in Tennessee 154; tri-
chopes 391
Erysimum repandum 391
Erythrotrichia carnea 298
Euonymous europaea 321; for-
tunei 147
Eupatorium albicaule 501, 502;
var. laxius 501, 502; bertholdii
504; coelestinum 103; collinum
503; var. collinum 502; var.
mendezii 503; aff. eximium
273; Fendleri 83; hebebotryum
503; leucoderme 501; mendezii
503; microcephalum 505, 506;
neaeanum 503; nigrescens 503;
ovaliflorum 504; ovatifolium
504; parasiticum 272; petraeum
504, 506; polybotryum 504-506;
scabrellum 505; stillingiaefo-
lium 502, 503; tepicanum 503;
ymalense 501, 502
Euphorbia Fendleri 83; stricto-
spora 392
Eustoma grandiflorum, f. album
385
Euthora cristata 269, 423
Evans, K., see Mohlenbrock,
Robert H.
Fagus grandifolia 524
Fimbristylis autumnalis 294
Fosliella lejolisii 424
544
Fralick, Richard A., see Mathie-
son, A. C.
Franseria tomentosa 394
Fucus distichus, ssp. distichus
409, 418; ssp. edentatus 418;
ssp. evanescens 418, 477; 480,
483; edentatus 268; evanescens
268; filiformis 268; spiralis
268; vesiculosis 268, 295, 298,
409, 411, 418, 419, 480, 481,
483; var. spiralis 411, 418, 482
Fundy, On the Marine Algae of
Kent Island, Bay of 264
Galeana pratensis 205
Galium verum 394
Galpinsia Fendleri 83
Garay, Leslie A., In Memoriam
Charles Schweinfurth, 1890-
1970 71
Gaura parviflora 392
Giffordia granulosa 266, 418
Gigartina stellata 269, 415, 423
Glaux maritima 384
Gloiosiphonia capillaris 423
Glyceria striata 148
Gnaphalium berlandieri — 508;
canescens 506; chilense 508; de-
currens 509; gracile 509, 510;
hirtum 508, 509; inornatum
507; leptophyllum 509; leuco-
cephalum 509; macounii 509;
pedunculosum 507, 508; pur-
pureum 508; roseum 506; spha-
cilatum 507; sprengelii 508;
stramineum 508; tenue 509;
viscosum 508, 509; Wrightii
506
Gomphonema olivaceum 481
Gracilaria foliifera 480; verru-
cosa 480
Graptopetalum rusbyi 312
Great Plains, New and Interest-
ing Plants from the 378
Grindelia aphanactis 106
Haber, Erich, Priority of the Bi-
Rhodora
[Vol. 74
nomial Pyrola chlorantha 396
Halenia deflexa 393
Halosaccion ramentaceum 269
Haplopappus clementis 107; exi-
mius 106; laceratus 107; lyalli
102, 106, 107; peirsonii 106;
pygmaeus 107
Harriman, Neil A., Records on
the Flora of Wisconsin 156
Hebeclinium tepicanum 503, 504
Hehre, Edw. J., Lomentaria
clavellosa (Turner) Gaillon. An
Addition to the Marine Algal
Flora of New Hampshire 158;
A. R. Hodgdon and R. B. Pike,
The Parker Cleaveland Her-
barium of Bowdoin College 80
Helianthus mollis 388
Hellquist, C. Barre, Range Ex-
tensions of Vascular Aquatic
Plants in New England 131
Hemibaccharis hieraciodes 498
Hemistegia decurrens 442
Hemitelia conformis 450; petio-
lata 450; Woronovii 450
Hemizonia minima 205; durandi
204
Hess, William J., see Reveal,
James
Heterocladus caracasanus 244,
249
Heterophylleia caracasana 249
Heterotheca inuloides, var. rosei
510; leptoglossa 510
Hibiscus lasiocarpa 384; syria-
cus 145; trionum 150
Hidalgoa ternata 204, 206
Hildenbrandia prototypus 269,
411, 423, 480
Hodgdon, A. R. and Frederic L.
Steele, Recent Discoveries in
the New Hampshire Flora 291;
see also Hehre, Edw. J.
Hoffmannseggella, A New Spe-
cies of the Orchid Genus 283
Hoffmannseggella brevicaulis 284,
1972]
285; cinnabarina 283; harpo-
phylla 284, 285; Kautskyi 285
Hookeria acutifolia 524
Hydrophyllum appendiculatum
385; Fendleri 83
Hymenopappus tenuifolius 395
Hymenoxys richardsonii 109; var.
floribunda 109
Hypericum Arnoldianum 277; X
Arnoldianum 276-280; X Daw-
sonianum 279, 280; densiflorum
146, 278-281; denticulatum 146,
147; dolabriforme 146, 147;
frondosum 281; galioides 276-
278; Kalmianum 280, 281; lobo-
carpum 146, 276-280; X no-
thum 280, 281; perforatum 392;
prolificum 279-281; punctatum,
var. pseudomaculatum 145,
146; var. punctatum 146; Sect.
Myriandra, An Evaluation of
Some Putative Spontaneous
Garden Hybrids In 276; spathu-
latum 146; sphaerocarpum, var.
sphaerocarpum 147; var. turgi-
dum 146, 147; X Van Fleetii
281
Hypochoeris glabra 149; radicata
149
Hypoxis hirsuta 293
Illinois Field
Studies 142
Ipswich, Massachusetts, The Ecol-
ogy of Benthic Salt Marsh
Algae at, I. Zonation and dis-
tribution of Algal Species 475
Iris lacustris 271; From Ontario,
A White-flowered Form of 271;
f. albiflora 271; missouriensis
390
Isthmoplea sphaerophora 298
Iva axillaris 387; frutescens, var.
oraria 291
and Herbarium
Jaegeria hirta 170
Index to Volume 74
545
Johnson, Miles F., Tussilago far-
fara L. (Senecioneae-Astera-
ceae) in Virginia 403
Jones, H. G., A New Species of
the Orchid Genus Hoffmann-
seggella 283
Juglans nigra 390
Juncus balticus, var. littoralis
478; gerardi 476-478, 483, 485;
trifidus 367, 368, 370, 371, 373,
3875; vaseyi 390
Kansas, The Yellow Mandarin, A
Notable Addition to the Flora
of 523
Kiger, Robert W., A New Vari-
ety of Leucophyllum laeviga-
tum (Scrophulariaceae) from
Mexico 347
King, R. M. and H. Robinson,
Neomirandia allenii A New
Epiphytic Composite of the
American Rain Forest 272
Knapweeds (Centaurea) in Can-
ada and the United States, Dis-
tribution of Native and Intro-
duced 331
Kobresia elachycarpa 329; sim-
pliuscula 329
Koetzner, Kenneth and R. D.
Wood, On the Marine Algae
of Kent Island, Bay of Fundy
264
Kornmannia leptoderma 479-481
Kovanda, Miloslav, Somatic
Chromosome Numbers for
some Asteraceae 102
Kylinia collopoda 85, 91, 93, 96;
compacta 85, 89, 93, 96; monili-
formis 85, 93, 96; scapae 92,
98; secundata 268, 473
Laelia anceps 283
Laminaria agardhii 267; digitata
127, 129, 130, 268, 411, 419;
intermedia 268; longicruris
546
268; platymeris 268; Sinclairii
127; saccharina 129, 411, 419,
421, 480, 483; stenophylla 268
Lamium amplexicaule, f. albiflo-
rum 386
Lavatera cretica 392
Lathyrus aphaca 155; L. New
to Tennessee and the South-
west 155
Leathesia difformis 419
Lecanora chlarotera 478
Lechea tenuifolia 293; villosa 293
Ledum groenlandicum 529
Leonurus Marrubiastrum
386
Lespedeza virginica 292
Lesquerella gordonii 381; lasio-
carpa 77, 79; var. ampla 78;
ssp. berlandieri 78, 79; var.
heterochroma 78; var. hispida
78; var. berlandieri 78, 79; in-
termedia 83; var. hispida 79;
ssp. lasiocarpa 77; Nomenclatu-
ral Changes in 76
Leucantha cyanus 344
Leucophylum laevigatum 347,
348; (Scrophulariaceae) From
Mexico, A New Variety of 347;
var. coahuilense 347-349; var.
laevigatum 349
Liatris punctata 103; var. ne-
braskana 104; var. mexicana
104; var. punctata 104; var.
typica 104
Linaria dalmatica 156
Liquidambar, Chromosome Num-
bers in 287; formosana 288,
290; orientalis 287, 290; sty-
raciflua 287, 288, 290, 524; X
L. formosana 289; X L. ori-
entalis 289
Lithoderma extensum 419
Lithophyllum corallinae
pustulatum 423
Lithothamnion glaciale 423
156,
423;
Rhodora
[Vol. 74
Lobelia siphilitica, var. siphilitica,
f. albiflora 387
Loiseleuria procumbens 365, 367,
368, 370, 373, 374
Lomentaria clavellosa 158;
(Turner) Gaillon: An Addi-
tion to the Marine Algal Flora
of New Hampshire 158
Lotus corniculatus 150, 392
Lupinus ashenbornii 490, 494;
bilineatus 491, 492; hartwegii
494; mexicanus 489-494; Cerv.
ex Lag. 489; persistens 494;
termis 491
Lychnis alba 149;
391
Lysimachia terrestris 150
Lycium pallidum 83
Lycopodium lucidulum 524; se-
lago 367; tristachyum 353
Lythrum salicaria 393
chalcedonica
McGregor, Ronald L., Lawrence
K. Magrath and Ronald Wee-
don, New and Interesting
Plants from the Great Plains
378
Macrocystis pyrifera 130
McVaugh, Rogers, Nomenclatu-
ral and Taxonomic Notes on
Mexican Compositae 495
Madia minima 204, 205
Magnolia tripetala 524
Magrath, Lawrence K., see Mc-
Gregor, Ronald
Mandarin, The Yellow, A Notable
Addition to the Flora of Kan-
sas 523
Mantisalea salmantica 339
Mapania macrophylla 327; syl-
vatica 328
Marchand, Peter, Wind and the
Winter-exposed Plant 528
Mathieson, Arthur C., Emery
Swan and Richard A. Fralick,
An Abnormal Specimen of
1972]
Laminaria digitata (L.) La-
mouroux 127; and Richard A.
Fralick, Investigations of New
England Algae V. The Algal
Vegetation of the Hampton-
Seabrook Estuary and the
Open Coast Near Hampton,
New Hampshire 406
Melampodium achillaeoides 203;
americanum 2, 6, 9, 12, 15, 16,
20-23, 31, 210; angulatum 202;
angustifolium 21, 206; anoma-
lum 203; appendiculatum 7, 18,
56, 61, 210; var. leiocarpum
61; var. sonorense 61; areni-
cola 64, 65; argophyllum 6, 9,
13, 35, 40, 41, 45, 46, 210; ar-
vense 188; aureum 7, 9, 12, 17,
191, 193, 211; australe 203;
baranquillae 203; berterianum
176; bibracteatum 7, 14, 59,
187, 210; bonairense 203;
brachyglossum 170; camphora-
tum 4, 204; cinerascens 44;
cinereum 6, 9, 35, 39, 41, 44, 46,
172, 210; var. cinereum 6, 14,
39, 40, 210; var. hirtellum 6, 14,
40, 42, 210; var. ramosissimum
6, 14, 40, 43, 210; (Compositae;
Heliantheae), Revision of the
Genus 1, 161; connatum 195;
copiosum 176; coronopifolium
51, 53; cornutum 184, 185; cos-
taricense 7, 18, 181, 183, 210;
cupulatum 7, 17, 20, 56, 59, 60,
64, 65, 210; dichothomum 202;
dicoelocarpum 4, 7, 10, 18, 30,
172, 182, 184, 210; digynum
204; diffusum 6, 8, 15, 25, 27,
210; var. lancelatum 26; divari-
catum 5, 7, 8, 10, 12, 18, 64,
69, 172, 175, 178, 195, 202,
210; var. macranthum 176;
diversifolium 202; dombeyanum
204; durandi 204; flaccidum
176; geminatum 204; glabri-
bracteatum 7, 16, 23, 67, 210;
Index to Volume 74 547
glabrum 7, 11, 19, 30, 199-201,
211; gracile 7, 14, 20, 164-166,
168, 210; var. oblongifolium
164; heterophyllum 21; hidal-
goa 204; hirsutum 205; hispi-
dum 20, 47, 53; humile 205;
kunthianum 22; lanceolatum
25, 167; laxum 169, leucanthum
5, 6, 9, 11, 13, 34-36, 40, 41, 46,
172, 210; var. argophyllum 45;
liebmanii 190; linearilobum 6,
11, 15, 21, 31-33, 202, 210;
longicorne 6, 19, 20, 54, 56, 210;
longifolium 7, 11, 12, 19, 27,
161, 210; longipes 6, 16, 29-32,
202, 210; longipilum 7, 11, 18,
20, 27, 68, 69, 185, 210; manil-
lense 25; microcarpum 164; mi-
crocephalum 7, 15, 20, 165-168,
210; mimulifolium 7, 20, 27,
163, 164, 210; minimum 205;
minutiflorum 205; montanum
4, 7, 9, 12, 189, 190, 193, 202,
211; var montanum 7, 17, 190,
191, 211; var. viridulum 7, 17,
191, 211; nayaritense 6, 15, 58,
59, 210; nelsonii 22; nutans 7,
15, 30, 200, 201, 211; oblongi-
folium 164; ovatifolium 176;
paludicola 205; paludosum 176;
panamense 177; paniculatum 7,
14, 166, 168-172, 210; parvulum
162; perfoliatum 3, 7, 8, 12, 18,
194-197, 200-202, 210; pilosum
6, 15, 27, 28, 210; pinnatum
202; pringlei 6, 16, 20, 23, 50,
51, 210; pumilum 176; ramosis-
simum 39, 43; repens 7, 14, 59,
188, 211; rhomboideum 162;
rhombifolium 177, 202; rosei 7,
18, 59, 64, 65, 67, 210; var. sub-
integrum 64; ruderale 205;
sericeum 6, 16, 20-22, 46-48,
51, 53, 210; var. brevipes 47;
var. exappendiculatum 47; var.
longipes 29; sinaloense 7, 19,
52, 186, 211; sinuatum 7, 9, 17,
548 Rhodora
20, 56, 59, 63, 64, 210; strigo-
sum 6, 19, 51, 52, 55, 56, 210;
suffruticosum 206; tenellum 7,
17, 59, 66, 67, 69, 210; var.
flaccidum 176; tepicense 7, 19,
52, 175, 184, 185, 211; terna-
tum 204; villicaule 68
Melampodium canescens 31, 32
Mellinger, Marie, see Duncan,
Wilbur H.
Melobesia lejolisii 424
Melosira juergensii 480
Membranoptera alata 424
Mendezia bicolor 515
Mertensia Fendleri 83
Microcoleus tenerrimus 484
Microlonchus salmanticus 339
Mimulus alatus, f. albiflorus 386
Mish, Lawrence B., List of the
Cladoniae of Southeastern Mas-
sachusetts with Special Refer-
ence to the Collection of
Charles Albert Robbins 254
Mitchella repens 437
Mohlenbrock, Robert H. and Dan
K. Evans, Illinois Field and
Herbarium Studies 142
Moore, Mary L, New Form of
Thuja occidentalis Resembling
Known Cultivars 352
Moore, R. J., Distribution of Na-
tive and Introduced Knapweeds
(Centaurea) in Canada and the
United States 331
Monostroma fuscum 414; f. blytii
266; grevillei 414; leptodermum
414; oxyspermum 411, 414,
427, 428, 482, 485-487; pulch-
rum 297, 414
Montanoa affinis 510, 511; graci-
lis 510, 511; karvinskii 510,
511; olivae 510, 511; subtrun-
cata 510, 511
Morton, J. K., Spontaneous Hy-
brids between Cerastium to-
[Vol. 74
mentosum Linn. and C. arvense
Linn. 519
Myriophyllum alterniflorum 139;
exalbescens 150; heterophyllum
139; humile 136; spicatum, var.
exalbescens 132; verticillatum,
var. pectinatum 137
Najas flexilis 133
Nemalion helminthoides 424
Neomirandea allenii 273, 274; A
New Epiphytic Composite of
the American Rain Forest 272;
angularis 272; araliaefolia 272;
costaricencis 273; eximia 274;
sciaphila 272, 274
Nephelea pubescens 445
New Hampshire, The Algal Veg-
etation of the Hampton-Sea-
brook Estuary and the Open
Coast near Hampton 406
New Hampshire Flora, Recent
Discoveries in the 291
New Hampshire, Snow Cover and
the Diapensia lapponica Habi-
tat in the White Mountains of
358
Nigella damascena 143
North Dakota Plants, New Rec-
ords of 389
Nothites ovatifolia 504-506
Nuphar variegatum 137
Nymphaea alba, f. rosea 188, 139;
odorata 137, 138; var. gigantea
138; f. rubra 139; tuberosa 135-
139
Ontario, À White-flowered Form
of Iris lacustris from 271
Ophryosporus ovatifolius 504-
506; petraeus 504, 505; scabrel-
lus 505
Oreocarya fulvocanescens 83;
multicaulis 83
Orobanche multiflora 387
1972]
Oryzopsis asperifolia 271; pun-
gens 271
Osmorhiza claytoni 393
Oxypappus scaber 511
Panicum philadelphicum 294, 378;
virgatum 478, 483
Paronychia depressa 380
Paspalum distichum 378
Pectis seemannii 511
Pellaea atropurpurea 220-225,
228-233, 237-240; Spores,
Chromosomes and Relations of
the Fern 220; longimucronata
232; notabilis 220-225, 229-234,
237-240; ovata 240; ternifolia
220, 226, 234, 237, 239; var.
ternifolia 221, 224, 226, 227,
229-232, 234, 235, 2371-239; var.
Wrightiana 221, 224, 226-228,
232, 234-239; truncata 232, 239
Peltandra undulata 516-518; vir-
ginica 517, 518; (L.) Schott &
Endlicher, The Combination
516
Penstemon procera 393
Percursaria percursa 411, 415,
421, 482, 485
Percome caudata 109; glandulosa
109
Petalonia fascia 409, 419, 420,
480, 481
Petrocelis middendorfii 424
Peyssonelia rosenvingii 424
Phacelia constancei 451-454; cor-
rugata 456, 459, 468; crenulata
451; Group (Hydrophyllaceae),
New Species in the 451; how-
elliana 456, 460, 461; integri-
folia 464; palmeri 451; pinnati-
fida 464; rafaelensis 454, 456-
458; utahensis 454, 468; vossii
462-464; welshii 465-468
Phlox alyssifolia 393
Phoradendron juniperinus 83
Phryma leptostachya 292
Index to Volume 74
549
Phycodrys rubens 270, 424, 477
Phyllophora brodiaei 269, 424;
membranifolia 424
Phymatolithon compactum 422;
laevigatum 424, 427; lenor-
mandi 425
Physalis Fendleri 83; pendula
150; similas 83
Physostegia formosior 393; parvi-
flora 393; virginiana 393
Pike, R. B., see Hehre, Edw.
Pilaiella littoralis 419 see also
Pylaiella
Pinus banksiana 389; leiophylla
350; ponderosa 350
Plumaria elegans 425, 428
Poa alpigena 367; Bigelovii 83;
Fedleriana 82
Polygala senega 392
Polygonum sawatchense 380
Polyides rotundus 411, 425
Polymnia perfoliata 194; revo-
luta 195
Polypogon monspeliense 389
Polysiphonia denudata 425, 480;
elongata 425; fibrillosa 425,
427, 428; harveyi 270; lanosa
270, 298, 425, 480; nigra 425,
480; nigrescens 270, 425, 480;
novae-angliae 426; subtilissima
426; urceolata 270, 426, 481
Porphyra miniata 426; umbili-
calis 268, 411, 426; f. epiphy-
tica 426
Potamogeton Berchtoldii 135; var.
tenuissimus 135; capillaceus
135; confervoides 135, 138, 139;
crispus 132; epihydrus, var.
ramosus 135, 136; filiformis
131; var. borealis 131; var.
Macounii 131; Friesii 133;
gemmiparus 135; gramineus,
var. gramineus 135; var. my-
riophyllus 135; Hillii 134;
longiligulatus 133, 134; natans
135; pectinatus 133; praelongus
550
137; Spirillus 135; strictifo-
lius, var. rutiloides 133; zosteri-
formis 134, 137
Potentilla palustris 391; propin-
qua 83; tridentata 367, 368,
370, 374
Prasiola stipitata 266
Prenanthes boottii 367
Protoderma marinum 415
Prunus serotina 321
Pseudendoclonium marinum 409,
415, 478, 485-487
Pseudolithodermum
419, 427
Pterophyton heterophyllum 99,
100; pauciflorum 99
Ptilimnium costatum 148
Ptilota serrata 269, 426
extensum
Punctaria latifolia 267, 298;
plantaginea 298
Pyenanthemum incanum 292;
tenuifolium 292; Torrei 292;
verticillatum 292; virginianum
292
Pylaiella littoralis 266, 298
Pyrola chlorantha 396, 397;
Priority of the Binomial 396;
virens 396, 397
Quercus ellipsoidalis 390; nove-
mexicana $83; palustris 148;
reticulatus 350; robur 529
Ralfsia borneti 419, 420, 428;
clavata 420, 428, 479-481, 486;
fungiformis 267, 420; verrucosa
409, 420, 479-481; 486
Ranunculus cardiophyllus 391;
carolinianus 149; repens, var.
pleniflorus 148; septentrionalis
149
Redfearn, Paul L., Jr., The Yel-
low Mandarin, A Notable Addi-
tion to the Flora of Kansas
523
Reveal, James L. and William J.
Rhodora
[Vol. 74
Hess, Variation within Calo-
chortus venustulus (Liliaceae)
350
Reynolds, Don R. and Ken Cal-
houn, Equisetum from Penin-
sula Florida 426
Rhamnus davurica 392
Rhapidophyllum hystrix 525; in
Mississsippi 525
Rhizoclonium implexum 482, 484,
485; riparium 411, 415, 484; f.
polyrhizum 482; f. riparium
482; f. validum 485; tortuosum
266, 415
Rhodochorton purpureum 426
Rhododendron lapponicum 367,
368, 370, 374
Rhododermis elegans 426; Rho-
domela confervoides 270, 426
Rhodymenia palmata 269, 427
Rhodophyllis dichotoma 427
Rhodophysema elegans 426, 427
Ribes cereum 391
Robinia viscosa 321
Robbins, Charles Albert, List of
the Cladoniae of Southeastern
Massachusetts with Special
Reference to the Collection of
254
Robinson, H., see King, R. M.
Rogers, Ken E., see Watson, J.
Ray, Jr.
Rollins, Reed C. and E. A. Shaw,
Nomenclatural Changes in Les-
querella 76
Rorippa islandica, var. fernald-
iana 144; var. islandica 144;
sessiliflora 144
Rubus idaeus, var. strigosus 144;
occidentalis 144; procerus 144
Rudbeckia hirta 108; var. angus-
tifolia 108; var. corymbifera
108; var. floridana 108; var.
hirta 108; var. pulcherrima
108; serotina 109
Rumex stenophyllus 379
1972]
Ruppia maritima 479
Sabatia campestris, f. albiflora
385
Saccorhiza dermatodea 420; poly-
Schides 129
Salix Fendleriana 83; irrorata
83; pedicellata 391; uva-ursi
367, 370, 374
Sambucus pubens 394
Santamour, Frank J., Jr., Chro-
mosome Numbers in Liquidam-
bar 287
Sargassum fluitans 295; natans
298
Saxifraga occidentalis, var. occi-
dentalis 382
Sciaromium lescurii 524
Scirpus americanus 476, 483;
cespitosus, var. callosus 367;
debilis 398; maritimus, var.
Fernaldii 291; olneyi 329;
purshianus 398, 400, 401; and
S. smithii, Chromosome Num-
bers of 398; var. purshianus
398, 399, 401; f. williamsii 398,
399, 401; robustus 291; smithii
398, 400, 401; f. setosus 398,
399, 401; f. levisetus 398, 399,
401; f. smithii 398, 399, 401
Schizothrix calcicola 487
Schoenoxiphium kobresioides
327; rufum 323
Schuyler, Alfred E., Chromo-
some Numbers of Scirpus
purshianus and Scirpus smithii
398
Schweinfurth, Charles, In Memo-
riam 71
Schwetschkeopsis fabronia 524
Sclerocarpus columbianus 203
Scutellaria brittonii 386; ovata,
var. ovata 148, 149; var. rugosa
148, 149; var. versicolor 148,
149
Index to Volume 74
551
Scytosiphon lomentaria 267, 298,
409, 420, 479, 480
Sedum alamosamum 311; cocker-
ellii 302-306, 309, 311, 312, 314;
diffusum 311; glaucophyllum
312; griffithsii 302, 303, 308,
311; havardii 302, 308, 311,
320; in the Southwestern
United States, Intraspecific
Variation in Chromosomes of
301; lanceolatum 3811, 312;
niveum 302, 305, 317; spathuli-
folium 312; stelliforme 302,
305, 306, 308, 309, 311, 312,
318; ternatum 306, 312;
wrightii 302, 305, 306, 308, 310,
311, 318
Selaginella Underwoodei 82
Senecio filifolius 111; longilobus
111; vulgaris 395
Shaw, Elizabeth A., see Rollins,
Reed C.
Sida spinosa 378, 384; f. albiflora
383
Sigesbeckia flosculosa 204
Skog, Laurence E., The Genus
Coriaria (Coriariaceae) in the
Western. Hemisphere 242
Smith, Earle C., Jr., Redefinition
of Cedrela oaxacensis C. DC.
& Rose 124
Spartina patens 411, 476, 483,
484; alterniflora 411, 413, 414,
417, 418; var. glabra 476, 482,
483
Sphacelaria plumosa 420, 427;
radicans 420, 428
Sphaeralcea Fendleri 83
Sphaeropteris atahuallpa 442-
444; horrida 443; marginalis
442; sipapoensis 441, 442, 444
Sphaerotrichia divaricata 267
Sphagnum capillaceum, var. tene-
rum 524
Spilanthes guatamalensis 177
552
Spongomorpha arcta 266, 297,
415; spinescens 266, 415
Spongonema tomentosum 421
Sporobolus airoides 390
Solanum sarrachoides 386
Solidago cutleri 367; flexicaulis
107; multiradiata 107; var.
scopulorum 107; sempervirens
108, 478; var. mexicana 108;
var. sempervirens 108
Sorapion kjellmani 420, 427
Stachys palustris 393; scopulo-
rum 393
Steele, Frederic L., see Hodgdon,
A, R.
Stellaria aquatica 381
Stevens, O. A., New Records of
North Dakota Plants 389
Stuessy, Tod F., Revision of the
Genus Melampodium (Com-
positae: Heliantheae) 1, 161
Suckleya suckleyana 380
Svenson, Henry K., The Rachilla
in Cape Cod Species of Carex
with Notes on the History of
the Perigynium and Rachilla
321; Rediscovery of Tilia neg-
lecta Spach 469
Swan, Emery F., see Mathieson,
A. C.
Synthlipsis berlandieri 78, 79;
var. hispida 78, 79; hetero-
chroma 78
Takhtajan, A., XII International
Botanical Congress, 528
Thelesperma marginatum 395
Thuja occidentalis 352-354, 356;
f. prostrata 354; Resembling
Known Cultivars, A New
Form of 352; var. ericoides 354
Thymelaea passerina 384
Tiffney, Wesley N., Jr, Snow
Cover and the Diapensia lap-
poncica Habitat in the White
Rhodora
[Vol. 74
Mountains of New Hampshire
358
Tilia americana 472, 473; var.
heterophylla | 473; argentea
473; caroliniana 473; glabra
473; heterophylla 470, 472,
473; Michauxii 472, 473; neg-
lecta 469-474; Spach, Redis-
covery of 469; pubescens 472
Tolypothrix tenuis 485
Tragoceros americanus 511, 512;
microglossus 511, 512; schie-
deanum 203, 511, 512; zin-
nioides 512
Tribulus cistoides 117, 120
Trichipteris cordata 446; cyclo-
dium 446-449; Maguirei 447-
449; saggittifolia 446; Steyer-
markii 446, 448; Williamsii
447, 449
Trifolium pratense, f. leucochra-
ceum 150
Trigzonospermum adenostemmoi-
des 513
Triodanis biflora 387
Triphora trianthophora 141;
(Sw.) Rydb. in Maine, A Sec-
ond Occurrence for 141
Triplasis purpurea 390
Tryon, Alice F., Spores, Chro-
mosomes and Relations of the
Fern Pellaea atropurpurea 220
Tryon, Rolla, Taxonomie Fern
Notes, VI. New Species of
American Cyatheaceae 441
Tussilago farfara 403, 404; L.
(Senecioneae-Asteraceae) in
Virginia 403
Uhl, Charles H., Intraspecific
Variation in Chromosomes of.
Sedum in the Southwestern
United States 301
Ulmus procera 143
Ulothrix flacca 265, 415;
flaccida 485
sub-
1972]
Ulva gigantea 479; lactuca 266,
409, 411, 416; rigida 480
Uncinia uncinoides 323
Unxia camphorata 204, 205;
digyna 204; hirsuta 205; suf-
fruticosa 206
Urococcus foslieanus 487
Urospora collabens 416; penicilli-
formis 416; speciosa 416
Utricularia geminiscapa 136
Vaccinium angustifolium 367;
uliginosum 365, 367, 368, 370-
375; vitis-idaea 365, 367, 370,
374
Vallisneria americana 133
Vaucheria arcassonensis 484;
compacta, var. koksoakensis
486; intermedia 482, 484, 485
Verbena hastata X urticifolia
150
Verbesina chapmanii 100; (Com-
positae) Endemic to Florida,
Nomenclatural Clarification of
Two Species of 92; heterophyl-
la 98, 99; pauciflora 99, 100;
warei 98-100
Veronica barbinervis 513; chaca-
lana 513; palmeri 513; triflos-
culosa 513, 514
Veronicastrum virginicum 394
Vesicaria lasiocarpa 77
Viguiera blakei 514; multiflora
109; puruana 514
Villanova achillaeoides 203
Viola pedata 293
Virginia, Tussilago farfara L.
(Senecioneae-Asteraceae) in
403
Index to Volume 74
553
Voluntarella muricata 339
Watson, J. Ray, Rhapidophyllum
hystrix in Mississippi 525; and
Ken E. Rogers, Eriogonum
harperi Goodman in Tennessee
154
Webber, E. E. and R. T. Wilce,
The Ecology of Benthic Salt-
marsh Algae at Ipswich, Mas-
sachusetts I. Zonation and Dis-
tribution of Algal Species 475
Wedelia minor 175; ovatifolia
175; perfoliata 194
Weedon, Ronald R., see Mac-
gregor Ronald L.
Wilce, R. T., see Webber, E. E.
Wind and the Winter-exposed
Plant, 528
Wisconsin, Records on the Flora
of 156
Woelkerling, Wm., Some Algal
Invaders of the Northwestern
Fringes of the Sargasso Sea
295; Studies on the Audouinella
microscopica (Naeg.) Woelk.
Complex (Rhodophyta) 85
Wolffia punctata 136; columbiana
136
Wood, R. D., see Koetzner, Ken-
neth
Xanthocephalum conoideum 514,
515; seriocarpum 514
Zarabellia rhomboidea 12, 162
Zinnia bicolor 515, elegans 515,
516; tenella 515; violacea 515,
516
Volume 74, No. 800, including pages 441-553, was issued December 29, 1972.
INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA
Manuscripts must be double-spaced or preferably triple-
spaced (not on corrasable bond), and a list of legends for
figures and maps provided on a separate page. Footnotes
should be used sparingly, as they are usually not necessary.
Do not indicate the style of type through the use of capitals
or underscoring, particularly in the citations of specimens,
except that the names of species and genera may be under-
lined to indicate italics in discussions. Specimen citations
should be selected critically especially for common species
of broad distribution. Systematic revisions and similar
papers should be prepared in the format of “The System-
atics and Ecology of Poison-Ivy and the Poison-Oaks,"
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu-
larly with reference to the indentation of keys and syno-
nyms. Papers of a floristic nature should follow, as far as
possible, the format of “Contribution to the Fungus Flora
of Northeastern North America. V.," H. E. Bigelow & M. E.
Barr, Rhodora 71: 177-203. 1969. For bibliographic cita-
tions, a recommended list of standard journal abbreviations
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot.
Club 85: 277-300. 1958.
vmm
CONTENTS: — continued
Lupinus mexicanus Cerv. ex. Lag.
IEE AIMÉ AAE E E E E 489
Nomenclatural and Taxonomic Notes on Mexican Compos-
itae.
Rogers MeVOWughy EE E E E enotas 495
The Combination Peltandra virginica (L.) Schott &
Endlicher.
Will H. Blackwell, Jv. ..ccccccccccssssccssesssccsssseccccccessssascesssssssessesssnes 516
Spontaneous Hybrids Between Cerastium tomentosum Linn.
and C. arvense Linn.
JURE IVE OTE os. oo de ct taea Cerere evoss coe donsensseceacuscesdtwasnsviervetesseetess da 519
Amphiprora ornata Bailey — A Second Station in New Eng-
land.
Ts CS CH LOU ROTE EA S ONDE MC ante eee o t e teer 521
The Yellow Mandarin, A Notable Addition to the Flora of
Arkansas.
tuos DB Pd E BIT PAA I A EE A E a a UU: ERES 523
Rhapidophyllum hystrix in Mississippi.
PANELA Ta 1010: O E E E a A E E E 525
Equisetum from Peninsula Florida.
Don R. Reynolds and Ken Calhoun ................. eee 526
Wind and the Winter-exposed Plant.
EEG EU ERO! USE CERRAR D E DEUM REC 528
XII International Botanical Congress.
ELT E N a E T inii MM nL 532
ee E OEO EE aana ieser antaiaheuuiees 583
Index to Volume 74 oi..c.ccccccccccccccccceccssscccecsscscceseeeseetaceceesteceeceeeens 5387
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