Rhodora

Dodova

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by
ALBION REED HODGDON, Editor-in-Chief

ALBERT FREDERICK HILL >)
STUART KIMBALL HARRIS !
RALPH CARLETON BEAN

ROBERT CRICHTON FOSTER | Associate Editors

ROLLA MILTON TRYON
RADCLIFFE BARNES PIKE

VOLUME 64

1962

The New England Botanical Club, fuc.

Botanical Museum, Oxford St., Cambridge 38, Mass.

73
Ad t.
YARLOW REFERENCE LienARY APR 3 1962

Hovora

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by,

ALBION REED HODGDON, Editor-in-Chief

ALBERT FREDERICK HILL
STUART KIMBALL HARRIS
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON
RADCLIFFE BARNES PIKE

Associate Editors

Vol. 64 January-March, 1962 No. 757
CONTENTS:
Change of Editor-in-Chief. Reed C. Rollins .............. re 1

Chromosome Numbers of Dyssodia (Compositae-Tagetinae)
and Phyletic Interpretations.

Marshall C. Johnston and B. L. Turner vicccccsccscccsssssscsscesssscceass 2
The Parasitic Behavior of Melampyrum lineare and a Note

on its Seed Color. Martin A. Piehl ................. eren eene nnne 15
Further Notes on the Distribution of Bidens connata vars.

pinnata and gracilipes. Earl Edward Sherf uoco 23
The Annual Species of Houstonia in Illinois.

Robert H. Mohlenbrock and Leo E. Halbig ......... eet 28
Watercress in the New World. Peter S. Green ...... ce ay"
A New Species of Scirpus in the Northeastern United States.

Be I Per TT MRRRRRPRRRUCL ERREUR S 43
A Survey of the Algae of Lake Quinsigamond. '

Tang Shih-Chen Auyang eee eene 49

Che Nem England Botanical Club, Ine.

Botanical Museum, Oxford St., Cambridge 38, Mass.

CONTENTS: — continued

Nomenclatural Notes on Two Southeastern Rhynchosias.

Robert L. Wilbur coececccccesssssccccscccccccecceccseeseeeceesssnnneceaeeeesausauaeeeeeeees 60
The Occurrence and Hybrid Nature of an Enchanter’s

Nightshade in Ohio. Tom S. Cooperrider ................ esses. 63
The Taxonomy of Setcreasea ovata.

Shamin A. Faruqi, R. P. Celarier and K. L. Mehra ............. 68
The Varieties of Luzula acuminata. John E. Ebinger .............. 74
The Occurrence of Spirodela oligorrhiza in the United States.

Edwin H. Daubs cecccccccccccccccccsscscccsssscccessscssesescscesscceeesseseusesecenseeese 83

Paspalum fimbriatum in the United States.Donald J. Banks.... 85
The Genus Anonymos and its Nomenclatural Survivors.

Daniel B. Ward ..... esee ettet tnter nennen 87
John Bachman and the Happoldt European Journal (Review).
Joseph. EWAN viccececsscsessccessescescceeseccessccesscseessecssecessccessssesseceseecseeeeuss 92

RHODORA.—A quarterly journal of botany, devoted primarily to the
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Rhodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 64 January-March, 1962 No. 757

CHANGE OF EDITOR-IN-CHIEF

Since the founding of Rhodora in 1899, there have been
but three principal editors. B. L. Robinson started the jour-
nal and was Editor-in-Chief for thirty years. M. L. Fernald
succeeded Robinson in 1929 and served until his death in
1950. It might appear to some that this long period,
together with the eleven years I have served in this same
capacity, established a tradition that the head of the Gray
Herbarium also be Editor-in-Chief of Rhodora. However,
there is no inherent reason why this should be the case even
though there has always been a very close association be-
tween the Gray Herbarium and the New England Botanical
Club. This association has existed from the founding of the
Club, and we are hopeful that it will continue undiminished
far into the future.

Over the years there has been a steady expansion of the
geographical area to which Rhodora pertains. At first, New
England and the Maritime area of Canada were uppermost
in the minds of both contributors and receivers of the jour-
nal. Later, Gray’s Manual area was thought of in this con-
nection. Now, we think in terms of the North American
Flora as a frame of reference when considering manuscripts
and there is only a loose adherence to this area. Expansion
in a different direction has also occurred. Systematic
Botany is now widely considered to include most aspects of
evolutionary botany, cytotaxonomy, experimental taxono-
my, cytogeography and genecology and Rhodora has regu-
larly accepted papers that reflect this greater breadth of
view of the subject.

bo

Rhodora [Vol. 64

With this issue, the journal begins as a quarterly. It is
anticipated that the coverage will be roughly what it has
been in recent years. Also, it is expected that the number
of printed pages per volume will be proximately 376, about
the same as recent volumes. To carry the journal forward,
the Council of the New England Botanical Club has named
Professor Albion R. Hodgdon, Chairman of the Department
of Botany, University of New Hampshire, as Editor-in-
Chief. Associate Editors are listed on the front cover.

It is a pleasure for me to turn over the reins of Rhodora
to Professor Hodgdon and to wish him every success in this
new undertaking.

Added to my own efforts in editing Rhodora during the
past eleven years have been those of a number of Club
members acting in the capacity of Associate Editors. To
these men, I should like to express publicly my apprecia-
tion. — REED C. ROLLINS.

CHROMOSOME NUMBERS OF DYSSODIA
(COMPOSITAE-TAGETINAE)
AND PHYLETIC INTERPRETATIONS
MARSHALL C. JOHNSTON AND B. L. TURNER!

The genera Dyssodia Cav., Nicolletia Gray, Leucactinia
Gray, Chrysactinia Gray, Porophyllum Adans., Lescaillea
Griseb., Pectis L., Tagetes L., and Adenopappus Benth.
together constitute a natural group characterized by the pos-
session of oil glands and aromatic oils, and by the arrange-
ment of the principal phyllaries in two subequal series and
often united into a cup. This group, whose naturalness has
not been questioned, has on the one hand been given tribal
status as the Tagetinae or Tageteae (Cassini, 1829, 1834;
Rydberg, 1915) and at the other extreme has been sub-
merged in the Heliantheae in the broader sense (Cronquist,
1955), but it has usually been included in the Helenieae? as
the subtribe Tagetininae (Bentham, 1873; Hoffmann, 1894;
etc.)

Of the genera included, Dyssodia in the very broad sense
of Hoffmann (1894) and of this paper, is the most diverse

‘Supported in part by National Science Foundation Grant 9025.

"That the tribe Helenieae is artificial (polyphyletic) has been suggested so often
it scarcely bears repetition (Bentham, 1873; Small, 1919; Leonhardt, 1949; Cronquist.
1955; ete.). For suggestions as to the true affinities of certain included genera see
Cronquist (1955), Turner (1956), Rock (1957), Turner and Johnston (1961), Turner,
Ellison and King (1961), and Raven and Kyhos (1961) and papers cited by them.

1962] Johnston and Turner — Dyssodia 3

from the standpoint of both morphology and cytology.
Among its species, we find some plants which show combina-
tions of the most specialized traits, and others which show
combinations of the least specialized traits of the entire
group. The ancestry of the other genera might well have
comprised plants which would be placed in the genus Dysso-
dia, if they persisted today. Dyssodia, then, is of primary
interest for an elucidation of phylogeny of the subtribe. The
present paper tabulates the few chromosome counts for the
genus already published along with the numerous original
ones (Table 1). The counts are presented within the frame-
work of a possible classification of the genus proposed by
Rydberg (1915).
METHODS

Chromosome counts newly reported in the present paper
were made from pollen mother cell squashes as outlined by
Turner and Johnston (1961a). The microscopy was partly
the work of Dr. W. L. Ellison and Mr. A. M. Powell, to whom
we are much obliged. Camera lucida drawings were made at
an initial magnification of ca. 2,000 diameters, and selected
drawings are presented in Fig. 1-8. Voucher specimens
(Table 1) are deposited in The University of Texas Her-
barium. The determinations are our own.

OBSERVATIONS

Chromosome counts are reported for 45 populations of
Dyssodia representing 17 species (Table 1). The gaps in our
chromosomal knowledge are obvious. Most of these are due
to the general unavailability of the taxa concerned. How-
ever, we may point out such a glaring omission as D. papposa
which occurs from southern Ontario to southern Mexico, and
is one of the most abundant roadside annuals in the high-
lands of Mexico. The deficiencies stem from the difficulty
encountered in attempting to get usable bud material in this
genus as compared to the ease of obtaining usable material
for some other Helenieae. Upwards of 250 bud-collections of
Dyssodia have been received, from which are derived the 45
new counts presented here. The technical difficulties may
partly be related to an abundance of oil in the buds which
seems to interfere with proper fixation of the nuclear

4 Rhodora [Vol. 64

material, and partly to a propensity of the chromosomes to
clump or fragment at meiosis. Perhaps a really thorough
populational sampling, especially in the critical sections
Thymophylla and Hymenatherum, may have to await the
development of better techniques for working with meiotic
chromosomes.

The section Dyssodia, containing the type species D.
papposa, has a base of x — 13. These are plants of the Mexi-
can highlands showing such unspecialized traits as retention
of a calyculum and separate principal phyllaries. Our con-
cept of D. pinnata includes D. subintegerrima Lag., and our
concept of D. papposa (Johnston, 1956a) is likewise more
inclusive than that of Rydberg (1915).

Syncephalantha, a Central American segregate, is obvi-
ously part of Dyssodia from a morphological point of view
and its chromosome base of x — 13 supports such a treat-
ment.

The section Clomenocoma is primarily southern Mexican
and Central American, with at least two Peruvian species.
The only counts reported are from two species which are
anomalous in their ecogeography in that they occur in the
Sonoran Desert. The base number, x = 13, is what one
would expect when the relatively unspecialized character of
the taxon is considered. It is probably that the whole section
is on a base of x — 13.

Dyssodia porophylla and D. cancellata, both placed in the
genus Lebetina by Rydberg, are morphologically very close
to Dyssodia and also show a base of x — 13. They are wide-
spread annual weeds in Mexico.

For Dyssodia acerosa of the section Aciphyllaea we have
conflicting reports, one from the southwestern United States
of n = 12 (Raven, et al., 1961), four of our own tentative
counts of n = ca. 13, and one clear, unequivocal count of
n = 8. This is a widespread desert species of southwestern
United States and far south into Mexico that is fairly uni-
form morphologically. It may also be added that in an other-
wise very difficult genus, this is one of the species we have
found most difficult to work with cytologically, and that
these preliminary data only serve to whet our curiosity.

1962] Johnston and Turner — Dyssodia 5

Plants of the sections Gymnolaena and Adenophyllum of
southern Mexico show combinations of some of the least
specialized habits and involucres to be found in Dyssodia. It
is perhaps from among these sections that we may find
species most closely resembling the ancestral types for the
entire genus. Unfortunately, we have no chromosomal in-
formation for either of these sections.

Dyssodia tagetoides, endemic to Texas and Oklahoma
(Johnston, 1956a), and placed in the monotypic genus Dyso-
diopsis by Rydberg, remains unknown cytologically in spite
of several attempts to obtain counts.

The section Thymophylla comprises several species of low
perennial herbs and shrublets of the dry areas of Mexico,
Arizona, New Mexico and western Texas, showing certain
specializations such as fusion of the principal phyllaries
into a cup and suppression of the calyculum. The section is
of special interest for several reasons, not the least of these
being that in the vicariad Dyssodia Belenidium — D. Thur-
beri it shows the bicentric geography, northern Argentina
— northern Mexico, which parallels the geographic distribu-
tions of so many other plants (I. M. Johnston, 1940; Garcia,
Soto, and Miranda, 1961; etc.) and which may indicate an
antiquity of perhaps tens of millions of years (Croizat,
1952). Although D. Thurberi is morphically identical to D.
Belenidium (Johnston, 1956a and 1956b), we here list the
names separately because of the report that 2n — 32 in D.
Belenidium, whereas we have found that n = 13 in the single
population of D. Thurberi from which we have been able to
obtain countable chromosomes. The further discovery of the
number n = 13 in three other species of this section leads to
a suspicion that the report for D. Belenidium may be in
error and at least needs confirmation.

Dyssodia pentachaeta, in the inclusive sense (Johnston,
1956a), is rather polymorphic and might be broken into
several species satisfactorily. Extensive population sam-
pling for cytological data is desirable, but the six counts
here reported do not offer much hope that chromosome num-
bers per se will be helpful in an elucidation of the taxonomy.
Moreover, we have found D. pentachaeta one of the most

6 Rhodora [Vol. 64

difficult species from which to obtain countable material.
The preliminary indication of two levels of ploidy in the
somewhat polymorphic taxon D. setifolia (sensu Johnston,
1956a) also indicates that further sampling is desirable.

The section Hymenatherum is closely related to section
Thymophylla. Even so ardent a proponent of segregate
genera as Rydberg (1915) placed them together, in the
genus Thymophylla. The species of section Hymenatherum
show a greater specialization than those of any others of the
genus. They are all annuals of the Texano-Mexican dry
areas, some of them being much reduced ephemeral desert
annuals. The calyculum is entirely suppressed and the in-
volucral fusion proceeds so far that in several species the
individual phyllaries are scarcely discernible. Furthermore
the base chromosome number x — 8 prevails among these
plants. These facts, together with the additional character
that most of the section Thymophylla have strictly opposite
leaves and branches whereas Hymenatherum shows an al-
ternate arrangement, at least in the upper parts of the stems,
might lead to the resurrection of Hymenatherum to the
generic level. But other evidence militates against such a
move, viz. the evidence presented here that in Dyssodia
tenuiloba the group is dibasic with one population clearly
showing n = 13, another n = 8, and observations by the
senior author of a group of plants which were probably the
result of hybridization between D. tenuiloba and D. penta-
chaeta, at one locality just south of Monterrey, Nuevo León
(unfortunately, the station has now been obliterated by en-
largement of a gravel pit).

Dyssodia micropoides, a species of depressed annuals with
a clearly defined distribution largely restricted to the dry
limestone piedmont of Coahuila and Nuevo León (Johnston,
1956a), has been something of an orphan, with no clear line
of descent within the genus. Rydberg (1915) placed it, as
the genus Gnaphaliopsis, immediately following his genus
Thymophylla. The discovery of n = 8 for this taxon sug-
gests that the species might best be placed near or with the
section Hymenatherum.

The rare species, Dyssodia neomexicana, deserves special

1962] Johnston and Turner — Dyssodia 7

comment for several reasons, one being that the present
report extends the known distribution several hundred miles
to the south. The species has previously been reported from
Chihuahua, from the type locality in southwestern New
Mexico, and from Apache County, Arizona. The plants are
ephemeral annuals with opposite leaves below, alternate
above, and an involucre very much like that of D. aurea of
the section Hymenatherum except for larger size and the
constant presence of a calyculum of a few linear bractlets.

v* dq hi N^.

? VY "ANE
e 50 "o E
<> D e
CN ¢ è
7 8 Se
Fic. 1-8. Camera lucida drawings of the meiotic chromosomes of Dyssodia spp.
Fig. 1. D. micropoides (n = 8). Fig. 2. D. neomexicana (n=7).— Fig. 8. D.
Treculii (n — 16). Fig. 4. D. tenuifolia (n = 8).— Fig. 5. D. acerosa (n — 8).
Fig. 6. D. aurea (n = 8). Fig. 7. D. tagetiflora (n = 13). — Fig. 8. D. decipiens

(n = 13). (X ea. 2000).

8 Rhodora [Vol. 64

Rydberg (1915) placed the species in his monotypic genus
Trichaetolepis, next to his segregate genus Dysodiopsis. We
list the species among Hymenatherum largely because of
lack of any other place to put it and because of the discovery
of the low number of n = 7, which stands alone in the genus.
DISCUSSION

The “gap” between x = 8 and x — 12, 13 in this genus has
been mentioned by Turner, ef al. (1961) and compared to
gaps in such genera as Aster, Chrysopsis, Gutierrezia and
Inula. The idea that the present gap in Dyssodia is the result
of inadequate sampling must not be rejected summarily. But
dismissing it momentarily, we might suggest another ex-
planation. The gaps, at least in some genera, not including
Dyssodia, have been explained as the result of aneuploid loss
to fairly low numbers (ca. 5) from an ancestral base of
x = ca. 9 (this hypothesis, or a modification of it, has been
presented by several authors, Stebbins, et al., 1953; Huzi-
wara, 1958; Raven, et al., 1960, etc.). On the other hand,
Turner, et al. (1961) think it more plausible to postulate a
fairly low ancestral base for the family (x = 4 or 5) and
to consider the probability that in various phyletic lines we
may detect aneuploid loss or gain in series both from the
ancestral base and from derivative multiples of it. At any
rate, regardless of the true etiology, we would like to
reiterate the absence of any known mechanism to explain
differential survivorship (i.e., natural selection) of certain
chromosome numbers as compared to other chromosome
numbers in the same phyletic or in the same aneuploid
series. Nor does it appear likely that such a mechanism will
suddenly come to light as a deus ex machina for the
aneuploid-reduction loss hypothesis, or as an explanation
of the situation in Dyssodia.

The gap in the Dyssodia series which includes numbers of
n — 9, 10, 11, is not strictly comparable to the gap of x — 6,
7 and/or 8 in the genera named above. In the latter genera
the counts clustering at 9 and 10 are very nearly double those
clustering at 4 and 5, whereas in Dyssodia the higher bases
of « — 12, 13 are not double the common lower base of
i = 8. If we postulate an ancestral base x = 8 we must ex-

1962] Johnston and Turner — Dyssodia 9

plain away a new “gap” of x — 14, 15, and if we postulate
an ancestral base of x — 7, we must explain the present
rarity of n = 7 and n = 14. Furthermore, in Dyssodia the
gap occurs not simply between species or species groups but
even within groups of plants so similar morphically that they
have been considered to constitute a single species (e.g., D.
Belenidium, D. tenuiloba, and D. acerosa). If one assumes
an ancestral basic chromosome number of x — 4 for the
genus Dyssodia, then much of the difficulty in explaining the

Section or species group:

T LEBETINA X13
Scr SYNCEPHALANTHA x : 13
(genus) z
1h DYSSODIA x. 13
ADENOPHYLLUM Le ?
DYSSODIA CLOMENOCOMA x+13
X*7,8,12,13 m
D.acerosa X: 8,12,13
THYMOPHYLLA LE 8(7),13
HYMENANTHERUM LE 8,13
D. micropoides X: 8
© D. neomexicana 1]
POROPHYLLUM tenus)
LE 12
PECTIS (genus)
x:12
LEUCACTINIA (tenus)
LESCAILLEA (tenus?) i?
1:?
on NICOLLETIA (genus)
x: 10
CHRYSACTINIA (eenus)

14,5 ee T

TAGETINEAN ANCESTOR

Fic. 9. Hypothetical phyletic relationships among genera, sections, and species of
the genus Dyssodia and related genera.

10 Rhodora [Vol. 64

TABLE 1. SPECIES OF DYSSODIA
ARRANGED IN A TENTATIVE CLASSIFICATION TOGETHER WITH
AVAILABLE CHROMOSOME COUNTS

Voucher specimens Chromosome Nos.
Taxa and/or literature reference n 2n
Sect. Dyssodia
D. papposa (Vent.) Hitche."
D. tagetiflora Lag. QUERÉTARO: 8 mi. n. of Queré-
taro, Johnston 4026A. 13 (Fig. 7)
” OAXACA: Oaxaca, King 2942. 13
” VERA CRUZ: 10 mi. sw. of No-
gales, King 2337. 13
D. pinnata (Cav.) Rob. SAN LUIS POTOSI: La Capilla,
Roek M-452. 13
” TAMAULIPAS: 11 mi. s. of Palmil-
las, Johnston 5622. 13
” MEXICO: 10 mi. w. of Toluca,
King 3591. 13
" MEXICO: Rte. 55, 3 mi. from
junction with Rte. 57, Powell
& Edmondson 582. 13
” MEXICO: Rte. 57, 29 mi. s. of toll-

gate, Powell & Edmondson 584. 18
Sect." [Genus Syncephalantha

Bartl.]
D. decipiens (Bartl.) OAXACA: 10 mi. n. of Oaxaca,
M. C. Johnston" King 2490. 13
” OAXACA: 3 mi. se. of Oaxaca,
King 3468. 13
” OAXACA: 9 mi. nw. of Oaxaca,
King 3514. 13
B CHIAPAS: 19 mi. ne. of Las
Cruces, King 3445. 13 (Fig. 8)
Sect. Clomenocoma (Cass.)
O. Hoffm.
D. grandiflora DC."
D. Jelskii Hieron."
D. remota Blake“
D. montana (Benth.) Gray‘
D. squarrosa Gray“
D. speciosa Gray"
D. aurantia (L.) Rob.*
D. Cooperi Gray Raven and Kyhos (1961) 26
D. porophylloides Gray Raven and Kyhos (1961) 13

Sect." [Genus Lebetina Cass.]

1962] Johnston and Turner — Dyssodia 11
Voucher specimens Chromosome Nos.
Taxa and/or literature reference n 2n
D. cancellata (Cass.) Gray AGUASCALIENTES: 4 mi. s. of
Aguascalientes, Rock M-465. 13
d DURANGO: 47 mi. n. of Durango.
Powell & Edmondson 956. ca. 13
H TAMAULIPAS: 5-6 mi. nw. of
Ocampo, King 3957. 13
D. porophylla (Cav.) Cav. GUANAJUATO: 18 mi. w. of Sala-
manca, Johnston 5949. 18
Sect. Gymnolaena DC."
D. oaxacana Greenm.
D. serratifolia DC.
D. integrifolia Gray
D. Seleri Rob. & Greenm.
Sect. Adenophyllum (Pers.)
O. Hoffm."
D. coccinea Lag.
Sect. Aciphyllaea DC.
D. acerosa DC. Raven and Kyhos (1961) 24
2 TEXAS: Crockett Co., 25 mi. w. of
Ozona. Johnston 6407. 13
" TEXAS: Val Verde Co., Del Rio,
Johnston 6489. 13
" TEXAS: Brewster Co., Elephant
Mesa, Johnston 6423. 131
" TEXAS: Brewster Co., 14 mi. e. of
Marathon, Johnston 6454. 8 (Fig. 5)
" COAHUILA: 5 mi. w. of Saltillo.
Powell & Edmondson 517. 13°
Sect." [Genus Dysodiopsis
(A. Gray) Rydb.]*
D. tagetoides T. & G.
Sect. Boeberastrum Gray*
D. littoralis Brandg.
D. anthemidifolia Benth.
D. concinna (A. Gr.) Rob.
Sect." [Genus Urbinella Greenm.]*
D. Palmeri (Greenm.) Macbr.
D. Gentryi M. C. Johnston
Sect. Thymophylla (Lag.)
O. Hoffm. :
D. tephroleuca Blake*
D. setifolia (Lag.) Rob. var. COAHUILA: Carneros, Johnston
setifolia 4186. 26

12 Rhodora [Vol. 64
Voucher specimens Chromosome Nos.
Taxa and/or literature reference n 2n
” SAN LUIS POTOSI: 8 mi. w. of Rio-
verde, Johnston 5649. 26
D. setifolia (Lag.) Rob. var. TAMAULIPAS: 11 mi. s. of Palmil-
setifolia las, Johnston 5629. ca. 26
" NUEVO LEÓN : 41 m. s. of Saltillo,
Rock M-27 4. ca. 26
D. setifolia var. Greggii (Gray)
M. C. Johnston*
D. aurantiaca (Brandg.) Rob.*
D. Belenidium (DC.) Macl. Covas and Schnack (1946) 32
D. Thurberi (Gray) A. Nels. TEXAS: Brewster Co., 1 mi. n. of
Black Gap, Johnston 6443. 13
D. Hartwegii (Gray) Rob. SAN LUIS POTOSI: 11 mi. e. of
San Luis Potosí, Johnston
4040a. 26
” DURANGO: 57 mi. ne. of Durango,
King 3761. ca, 26
D. pentachaeta (DC.) Rob. NUEVO LEON: 30 m. s. of Saltillo,
sensu lato Ellison 51. 13
» COAHUILA: 5 mi. w. of Allende,
Johnston 4169. 13
D. pentachaeta (DC.) Rob. COAHUILA: 5 mi. w. of Saltillo,
sensu lato Powell & Edmondson 516. ca. 13
D. pentachaeta (DC.) Rob. SAN LUIS POTOSI: 8 mi. w. of Rio-
sensu lato verde, Johnston 5648. 13"
D. pentachaeta (DC.) Rob. TEXAS: Brewster Co., 8 mi. w. of
Marathon, Johnston 6449. 13°
» TEXAS: Pecos Co., southern tip
of co., Johnston 6457. ca. 13
" TEXAS: Kinney Co., 8 mi. e. of
Brackettville, Johnston 6490. 13°
Sect. Hymenatherum (Cass.)
O. Hoffm.
D. tenuiloba (DC.) Rob. TEXAS: Webb Co., Laredo, John-
ston 5466. 13e
” TEXAS: Brooks Co., 8 mi. e. of
Falfurrias, Johnston 5467. 8e
D. Wrightii (Gray) Rob. TEXAS: San Patricio Co., 3 mi. n.
of Sinton, Johnston 5468. 8
» TEXAS: Goliad Co., 1 mi. w. of
Refugio Co. line, Johnston 5471. 8

»

TEXAS: Refugio Co., 2 mi. n. of

Aransas River, Johnston 5469. ca. 8

1962] Johnston and Turner — Dyssodia 13
Voucher specimens Chromosome Nos.
Taxa and/or literature reference n 2n
D. Treculii (Gray) Rob. TEXAS: Val Verde Co., Del Rio,
Johnston 6488. 16 (Fig. 3)
D. texana Cory*
D. mutica M. C. Johnston Johnston (1960) 8
D. aurea (Gray) A. Nels. var.
aurea
D. aurea var. polychaeta (Gray) COAHUILA: Bolsón de Mapimí,
M. C. Johnston Rock M-484. 8
a COAHUILA: 10 mi. e. of San
Pedro de las Colonias. Powell &
Edmondson 520. 8
CHIHUAHUA: 14 mi. n. of Parral,
Powell & Edmondson 972. 8
” CHIHUAHUA: just east of Chi-
huahua City, Powell & Edmond-
son 1018. 8 (Fig. 6)
D. anomala (Canby & Rose)
Rob."
D. tenuifolia (Cass.) Loes. SAN LUIS POTOSI: 7 mi. e. of San
(Incl. D. Neaei (DC.) Rob. Luis Potosí, Johnston 4041. 8 (Fig. 4)
and D. diffusa (Gray) Rob.) Same locality, Johnston 5638. 8
D. neomexicana (Gray) Rob. MEXICO: 36 mi. s. of tollgate
north of Mexico City, Powell &
Edmondson 588. Te (Fig. 2)
Sect." [Genus Gnaphaliopsis DC.]
D. micropoides (DC.) Loes. NUEVO LEON: w. of Monterrey,
Rock M-496. 8
" NUEVO LEON: 26 mi. s. of Sa-
binas Hidalgo, Johnston 5455. 8
di COAHUILA: San Lázaro, John-
ston 4184. 8 (Fig. 1)

"No nomenclatural combination has been published for this taxon at the proper
rank. Formal changes in status are not deemed appropriate for this contribution, but
must await a formal taxonomic treatment.

>bDyssodia decipiens (Bartl.) M. C. Johnston, comb. nov., based on Syncephalantha

decipiens Bartl., Ind, Sem. Hort. Goett. 1836: 6.

1836.

*No chromosome counts have been made or reported for this taxon.
4One or two fragments were also seen.
eThis count was quite clear and unequivocal.

obvious gap is removed. Thus taxa with n — 8 would be
tetraploid; n = 12, hexaploid; and taxa with n = 7 and 13
aneuploid derivatives from such taxa. The fact that at least

14 Rhodora [Vol. 64

one species, D. acerosa, has chromosome numbers n = 8, 12,
13 might be explained by the hypothesis of Turner, et al.
(1961) that the Compositae showed a very rapid and ex-
plosive evolution while still at the predominately diploid level
of x — 4 and 5, most of the generic lines being formed at
that time. Surviving diploid taxa, in that they are for the
most part not buffered against mutational change (Stebbins,
1950) and are thus more prone to rapid evolution, should
be relatively more specialized.

The data and some of our speculations are summarized in
Fig. 9. We can do no more at present than to pose the prob-
lems and remark that at the very least the picture must
tantalize the would-be phylogenist. — THE PLANT RESEARCH
INSTITUTE, DEPARTMENT OF BOTANY, THE UNIVERSITY OF

TEXAS, AUSTIN
LITERATURE CITED

BENTHAM, G. 1873. Notes on the classification, history and geograph-
ical distribution of Compositae. Jour. Linn. Soc. Lond. 13: 335-557.
CassINI, H. 1829. Tableau synoptique des Synanthérées. Ann. Sci.
Nat. (Paris) 17: 387-423.
. 1884. Opuscules phytologiques. vol. 3. Paris.

CROIZAT, L. 1952. Manual of Phytogeography. 587p. The Hague:
Uitgerif Dr. Junk.

CRONQUIST, A. 1955. Phylogeny and taxonomy of the Compositae.
Amer. Mid. Nat. 53: 478-511.

GARCIA, E., C. SOTO, AND F. MIRANDA. 1961. Larrea y clima. Anal.
Inst. Biol. Mex. 31: 133-171.

HOFFMANN, O. 1894. Compositae [in] Engler and Prantl, Natürl.
Pflanzenfam. 4(5) : 87-875.

HuziwARA, Y. 1958. Karyotype analysis in some genera of Composi-
tae. V. The ehromosomes of American Aster species. Jap. Jour.
Genetics 33: 129-137.

JOHNSTON, I. M. 1940. The floristic significance of the shrubs com-
mon to the North and South American deserts. Jour. Arnold Arb.
21: 356-363.

JOHNSTON, M. C. 1956a. The Texas species of Dyssodia (Composi-
tae). Field & Lab. 24: 60-69.

.1956b. A further note on Texas species of Dys-

sodia (Compositae). Field & Lab. 24: 104.

. 1960. Dyssodia mutica (Compositae), new species.
Southwest. Natural. 5: 225-226.

LEONHARDT, R. 1949. Phylogenetisch-systematische Betrachtungen.
I. Betrachtungen zur Systematik der Compositen. Oesterr. Bot.
Zeitschr. 96: 295-324.

1962] Piehl — Melampyrum lineare 15

RAVEN, PETER H., O. T. SOLBRIG, D. W. KyHos, AND R. SNOW. 1960.
Chromosome numbers in Compositae. I. Astereae. Amer. Jour.
Bot. 45: 124-132.

, AND D. W. Kvnos. 1961. Chromosome numbers in
Compositae. II. Helenieae. Amer. Jour. Bot. 48: 842-850.

tock, H. F. L. 1957. A revision of the vernal species of Helenium
(Compositae). Rhodora 59: 101-116, 128-158, 168-178, 203-216.

RYDBERG, P. A. 1915-1916. Carduaceae: Tageteae. No. Amer. FI.
34: 147-216.

SMALL, J. 1917-1919. The origin and development of the Com-
positae. New Phytol. 16: 157-177, 198-221, 253-276, 1917. 17: 13-
40, 69-94, 114-142, 200-230, 1918. 18: 1-35, 65-89, 129-176, 201-234,
1919.

STEBBINS, G. L. 1950. Variation and evolution in plants. 643 p. New
York: Columbia University Press.

, J. A. JENKINS. AND M. S. WALTERS. 1953. Chromo-
somes and phylogeny in the Compositae, tribe Cichorieae. Univ.
Calif. Publ. Bot. 26: 401-430.

TURNER, B. L. 1954. A cytotaxonomic study of the genus Hymeno-
pappus (Compositae). Rhodora 58: 163-186, 208-242, 250-308.

, W. L. ELLISON, AND R. M. KiNG. 1961. Chromosome
numbers in the Compositae. IV. North American species, with
phyletie interpretations. Amer. Jour. Bot. 48: 216-223.

, AND M. C. JoHNSTON. 1961. Chromosome numbers in
the Compositae-III. Certain Mexican Species. Brittonia 13: 64-69.

THE PARASITIC BEHAVIOR OF
MELAMPYRUM LINEARE AND A NOTE ON
ITS SEED COLOR'

MARTIN A. PIEHL

Parasitism has been described for several members of the
genus Melampyrum, the cow-wheats (Scrophulariaceae),
beginning with M. arvense of Europe (Decaisne, 1847). To
my knowledge, however, a parasitic existence has not been
substantiated for M. lineare, the single North American rep-
resentative of the genus. Fraysse (1906) ascribes to Hein-
richer (1904) the assertion that all species of the genus are
parasitic; however, the latter actually said (p. 412), “Alle
untersuchen Arten der Gattung Melampyrum sind para-
sitisch (M. arvense, barbatum, nemorosum, silvaticum, pra-
tense)”, and no reference was made to M. lineare. Shreve

1Contribution from the University of Michigan Biological Station.

16 Rhodora [Vol. 64

et al. (1910) failed to find parasitic connections for M.
lineare in Maryland but state that this plant was reportedly
parasitic farther north’, and Pennell (1935) whose studies
included field observations makes no mention of parasitism.
The lack of information concerning this species' parasitism
has received comment previously (Beauverd, 1916; Marie-
Victorin, 1935).

While studying the parasitism of Comandra, Pedicularis,
and other angiosperms, I have had occasion to examine M.
lineare for comparative purposes. The majority of my ob-
servations were made during the spring and summer of
1957, 1958, and 1960 in the Mackinac Straits region, Michi-
gan. Here M. lineare often occurs in dense clumps several
decimeters across, suggesting seeding from one or more
plants which previously grew nearby. Its characteristic
habitat is open woodland, including both sandy, often dry
Sites under conifers and more mesic aspen stands. Rarely it
oecurs in bogs.

This species is an opposite-leaved annual, which in the
study area is usually about 2 dm. tall at flowering, with con-
siderable variation in height. In contrast to achlorophyllous
holoparasites, e.g., Orobanche spp., there is no suggestion of
the parasitic habit in the aerial parts of such semiparasites
— the plants are green and the leaves are not proportion-
ately smaller than those of many autotrophs. The whitish
root system is shallow, consisting of a poorly defined pri-
mary root which is surpassed by some of the several lateral
roots. There are comparatively few root hairs, a condition
which has also been described for the European M. pratense
(Koch, 1887). The excavated root system gives the appear-
ance of extending only a few centimeters into the soil; how-
ever, the slender rootlets break easily, obscuring its full
extent. This breakage explained my failure to find parasitic
connections in the initial excavations. Minute swellings on
the roots were nonetheless detected, and upon close examina-
tion some were found attached to fragments of foreign plant
tissue. Later excavation was facilitated by selecting plants

*Additional information was not given by Shreve et al. as to when, where, and by
whom this observation was made.

1962] Piehl — Melampyrum lineare 17

in loose, dry sand, and after repeated attempts involving
careful brushing away of sand, a thread-like rootlet was
found attached to a host root by a near-microscopic, hemi-
spherical enlargement — the haustorium. Subsequently, it
was determined that haustoria develop on essentially all
plants of M. lineare, and that rhizomes as well as roots are
parasitized. By ordinary excavation with a trowel the
fragile parasite roots leading to connections were severed,
leaving the haustoria attached to the host, and thus destroy-
ing direct evidence of parasitic connections. The delicate-
ness of the attachments has undoubtedly contributed to the
lack of observations on the parasitic behavior of this species.

The haustoria are developed laterally along the parent
root, although at times they appear terminal because root
growth beyond the haustorium has been arrested. A similar
development is known for other semiparasites (Herbert,
1923; other authors). Often several haustoria develop in
close succession from a single parasite root (Plate 1265, C).
Unicellular, epidermal hairs develop from just above the dis-
tal end of some haustoria and may reach down to, or even
extend for some distance over, the surface of the host root
or rhizome (Plate 1265, D). Their development appears to
be associated with the early ontogeny of the haustorium.
Presumably they function as ordinary root hairs for a short
time, and later may aid (very slightly) in superficial attach-
ment of the haustorium to the host. Similar hairs occur on
other species, e.g., M. pratense (Knorz, 1848; Koch, 1887).

Anatomically, the haustorium consists of an outer region
of loosely-organized, sometimes partly-collapsed, parenchy-
matous cells, and a central region of compact, densely-proto-
plasmic cells. Through the latter region pass one or a few
vessels, leading to the distal, invading portion, and eventual-
ly effecting direct continuity between the xylem of the para-
site root and that of the host. I plan to discuss further
anatomical details in a later report.

The haustoria are nearly circular in surface view and are
ca. 0.5 (0.2-1.4) mm. in diameter, depending somewhat on
the size and nature of the root or rhizome parasitized. The
smaller haustoria are produced from fine rootlets with dia-

18 Rhodora [Vol. 64

meters down to 0.1 mm. and are attached to host rootlets of
about the same thickness. In general, host organs are Ca.
1 mm. in diameter, although on occasion those of greater
thickness are attacked. Among the largest observed were
roots of large-toothed aspen (Populus grandidentata) 5 mm.
in diameter, while those of blackberry (Rubus cf. alleghe-
niensis) and the rhizomes of bracken (Pteridium aquilinum)
were nearly as large. A maximum number of 26 haustoria
per linear centimeter of host root was recorded for aspen
roots of this size. Since this count was made from preserved
material, even more haustoria may have been present which
were unavoidably dislodged during collection. On the rough-
ened bark of these relatively large aspen roots, haustoria
tended to form in the furrows rather than on ridges, and
were often insecurely attached. When sectioned, these haus-
toria were found to have penetrated a part of the compara-
tively thick bark but had not reached living tissue.
Apparently the thickness and other characteristics of the
bark of some hosts limit the size of roots which can be suc-
cessfully attacked. When attached to larger host organs,
haustoria have a slightly concave, sucker-like distal surface,
while on smaller organs they are often clasping, as they
partly enclose the host. Haustorial roots tend to grow paral-
lel to large organs (Plate 1265, E), while they often contact
smaller ones at an angle.

Haustorial connections to 12 species have been detected.
These are taxonomically distributed over two phyla, in one
of which three classes and two subclasses are involved. Eight
of the hosts are dicots, but these belong to several generally
diverse families. The species to which attachments have
been found follow:

Sphagnum capillaceum (Weiss) Schrank?
Pteridium aquilinum (L.) Kuhn Amelanchier cf. arborea

Pinus resinosa Ait. (Michx. f.) Fern.

Carex pensylvanica Lam. Rubus cf. allegheniensis Porter
Populus grandidentata Michx. Gaultheria procumbens L.
Betula papyrifera Marsh. Melampyrum lineare Desr.
Quercus rubra L. Diervilla lonicera Mill.

3Although very little material was available for identification, Prof. H. L. Blomquist
believes it belongs to this species.

Plate 1265. Haustorial connections of Melampyrum lineare. A. pem attached
to a Pinus resinosa root, X 14. B. Haustorial connection to Sphagnum capillaccum
(pointer), X 12. C. Closely suecessive haustoria on a Populus grandidenta root, X 12.
D. Single haustorium showing exceptionally strong development of hairs; a few hairs
prolonged over the host root (pointer), X 36. E. Isolated haustoria formed from
Melampyrum roots paralleling an unusually large host root (Populus grandidentata),
X 9. F. Self-parasitic connection (pointer), connections to dead roots and humus
(small black particles), X 12.

20 Rhodora [Vol. 64

Undoubtedly many more hosts can be added with further
observation. The host ranges reported for several European
species of Melampyrum are limited to gymnosperms and
angiosperms (Heinricher, 1924). Heinricher also states that
these cow-wheats differ from each other somewhat in their
ability to grow on certain hosts.

Occasionally, I have found cases of intraspecific para-
sitism, where connections are formed to other individuals of
M. lineare growing nearby. Furthermore, self-parasitism,
involving other roots of a single individual, has been ob-
served (Plate 1265, F). In general, the external features
of attachment and penetration are the same here as in cases
involving other species.

My single observation of Sphagnum as a host was made
at (Inverness) Mud Lake Bog, Cheboygan County, Michi-
gan, where haustoria were found attached to the lower
achlorophyllous portion of the moss some distance below the
mat surface (Plate 1265, B). Whether this should be con-
sidered a case of true parasitism is questionable, for al-
though the upper portion of the moss was alive, the part to
which haustoria were attached may have consisted wholly of
dead cells. Attachments to a bryophytic host have not pre-
viously been recorded for a parasitic angiosperm.

' [In addition I have found numerous cases of what appears
to be saprophytism, for the haustoria were attached to dead
fragments of plant tissue (Plate 1265, F). These fragments
can often be recognized as dead and sometimes partly de-
cayed roots, and occasionally as other organs. In some in-
stances haustorial connections were formed to unidentifiable
humus particles. Those roots still intact, as well as the
Sphagnum mentioned above, could have died subsequent to
invasion ; however, this is not possible for the already much-
decayed material. Thus, living tissue is not a prerequisite
for attachment. Whether substances are obtained from the
dead material is unknown. This observation of attachments
to dead tissues agrees with those of M. pratense (Knorz,
1848; Koch, 1887) ; however, these workers believed this
species formed attachments to dead material only, and de-
scribed it as saprophytic rather than parasitic. Attachments

1962] Piehl — Melampyrum lineare 21

of M. pratense to living plants as well have since been re-
ported (Sperlich, 1902).

I have frequently observed some individuals of M. lineare
in a wilted condition in nature, and as collectors soon learn,
this species is one of the first to wilt, even when kept in a
vasculum. I suspected that those plants which wilted easily
had relatively few haustorial connections, and subsequent
examination of their roots tended to verify this. This pro-
pensity to wilt, and if the drought continues, to succumb, is
consistent with the observation that this species tends to be
morphologically unadapted to xeric conditions; paradoxical-
ly, however, it frequents dry sandy habitats. The partial
heterotrophy of this plant may well be vital to its success
under such conditions.

An observation concerning seed color in M. lineare seems
worthy of inclusion here. Although the seeds have been
described as blackish* (Pennell, 1935; Fernald, 1950), I
have repeatedly noted that they are beige or tan with a
lighter colored tip (elaiosome according to Gislén, 1949)
when released from the capsules. Ungerminated seeds found
at or near the soil surface early the following spring are
similarly colored, suggesting that the blackening is not a
normal after-ripening change. However, seeds from her-
barium specimens from various parts of the range are
almost always blackish, and apparently form the basis for
previous descriptions. Reference to seeds of my own speci-
mens which were recorded as beige in the field showed that
they had blackened after the specimens were dried and
pressed, and germination attempts indicated such seeds
were inviable. Also, some specimens blacken throughout
when dried, a phenomenon which is rather common in both
the Euphrasieae and Buchnereae sensu Pennell. Whether
such a color change occurs in seeds of the European species
is unknown, but if it does, an interesting etymological situa-
tion arises when one considers the generic name.

SUMMARY
Although previously known for European members of the

4Pennell describes them as brown or dark brown to blackish for ssp. pectinatum
which is not known from northern Michigan.

22 Rhodora [Vol. 64

genus, parasitism is substantiated for the first time for
Melampyrum lineare Desr. from studies in Michigan. Sub-
terranean connections of this seemingly wholly autotrophic
angiosperm are established by extremely fine roots bearing
minute haustoria. Invasion of the host root or rhizome
ordinarily effects direct connection between the xylem of
the host and that of the parasite. The hosts are taxonomical-
ly diverse — dicots, monocots, conifers, ferns, and in one
instance, a bryophyte. Melampyrum lineare can also be
self-parasitic. In addition, it forms attachments to dead
plant parts and humus.

Descriptions of the seeds of M. lineare as blackish are
believed based on dead, dried material which has blackened
after collection. Mature seeds in nature have been observed
to be beige or tan.

ACKNOWLEDGMENTS

I wish to acknowledge the helpful suggestions of Profes-
sors W. S. Benninghoff, Rogers McVaugh, W. H. Wagner,
Jr., and E. G. Voss of the University of Michigan in connec-
tion with the preparation of the manuscript. — DEPARTMENT
OF BOTANY, UNIVERSITY OF MICHIGAN, ANN ARBOR.

LITERATURE CITED

BEAUVERD, G. 1916. Monographie du genre Melampyrum L. Mém. Soc.
Phys. Hist. Nat. Genéve 38:291-657.

DECAISNE, J. 1847. Sur le parasitisme des Rhinanthacées. Ann. Sci.
Nat. Bot. III, 8:5-9.

FERNALD, M. 1950. Gray's Manual of Botany, 8th Ed. American Book
Co., New York.

FRAYSSE, A. 1906. Contribution à la biologie des plantes phanéro-
games parasites. Thesis, Univ. Paris, 178 pp. Soc. Anonyme de
l'Imprimerie Générale du Midi.

GISLÉN, T. 1949. Problems concerning the occurrence of Melampy-
rum arvense in Sweden. Oikos 1:208-234.

HEINRICHER, E. 1904. Melampyrum pratense L., ein in gewissen
Grenzen spezialisierter Parasit. Ber. Deutsch. Bot. Ges. 22:411-
414.

. 1924. Methoden der Aufzucht and Kultur der para-
sitischen Samenpflanzen. In Abderhalden, E., Ed., Handbuch der
biologischen Arbeitsmethoden. Abt. 11, Teil 2:237-350.

HERBERT, D. 1923. Phanerogamic parasites. Philip. Agric. 11:17-18.

1962]: Sherff — Distribution of Bidens connata 23

KNoRZ, DR. 1848. Über den von Hrn. Decaisne angegebenen Para-
sitismus der Rhinanthaceen. Bot. Zeit. 6:239-241.

KocH, L. 1887. Über die direkte Ausnützung vegetabilischer Reste
durch bestimmte chlorophyllhaltige Pflanzen. Ber. Deutsch. Bot.

Ges. 5:350-364.
MaRIE-VICTORIN, F. 1935. Flore Laurentienne. Imprimerie de la

Salle, Montréal.

PENNELL, F. 1935. The Scrophulariaceae of eastern temperate North
America. Monogr. Acad. Phila. 1:1-650.

SHREVE, F., M. CHRYSLER, F. BLopGETT, AND F. BESLEY. 1910. The
Plant Life of Maryland. Md. Weather Service, Baltimore.

SPERLICH, A. 1902. Beiträge zur Kenntniss der Inhaltsstoffe in den
Saugorganen der grünen Rhinanthaceen. Beih. Bot. Centr. 11:

437-485.

FURTHER NOTES ON THE DISTRIBUTION OF
BIDENS CONNATA VARS.
PINNATA AND GRACILIPES

EARL EDWARD SHERFF

Fassett (Rhodora 30: 31-35, Plate 160. 1928) listed the six
varieties of Bidens connata Muhl. known to him in the State
of Wisconsin: vars. typica (connata), fallax, petiolata,
ambiversa, anomala, and pinnata. For the last of these, var.
pinnata Wats., he listed but two collections from Wisconsin,
these from Polk County (see also Aldrich & Fassett, Science
70: 45. 1929). Subsequently he collected, either alone or with
some of his students, numerous specimens of var. pinnata
from various other counties in northwestern Wisconsin.
Through the kindness of Dr. H. Iltis of the Department of
Botany of the University of Wisconsin, I have been permit-
ted to examine those specimens critically. Moreover, in early
September of 1961, I made visits to some of the localities
where Fassett had collected and this permitted me to observe
the living specimens in their native habitats. Suites of speci-
mens were obtained to distribute to herbaria. As a contribu-
tion to the study of Wisconsin's flora, there follows here a
condensed list of all Wisconsin specimens of var. pinnata
studied by me. My own collections may be found represented
in the Herbarium of the Chicago Natural History Museum
(F) ; the others are in the Herbarium of the University of
Wisconsin:

24 Rhodora [Vol. 64

CHIPPEWA COUNTY: N. C. Fassett, John Budd, & C. L. Brynildson
22695, sandy shore of Bass Lake, 12 miles north of Bloomer, Oct. 2,
1949. BARRON COUNTY : W. T. McLaughlin 1020, sandy shore of Crystal
Lake, Comstock, Aug. 29, 1929; McLaughlin 1021, sandy shore of
Horseshoe Lake, Turtle Lake, Aug. 28, 1929; McLaughlin 1055, sandy
shore of Silver Lake, Cumberland, Aug. 29, 1929. POLK COUNTY: N. C.
Fassett 7891, sandy shore of Bass Lake, 9 miles west of Luck, Sept. 23,
1928; Fassett & L. R. Wilson 4016, sandy shore of Poplar Lake, Osceo-
la, Sept. 4, 1927 (3 sheets) ; W. T. McLaughlin 1024, sandy shore of
Pine Lake, Star Prairie; McLaughlin 1026, narrow sand strip, Twin
Lake, Amery, Aug. 25, 1929; McLaughlin 1030, grass-invaded shore,
small pond, Star Prairie, Aug. 26, 1929. WASHBURN COUNTY: N. C.
Fassett 7900, sandy shore of Shell Lake, Shell Lake, Sept. 22, 1928;
Fassett 7901, same place and date; Fassett 7906, sandy shore of Silver
Lake, Lampson, Sept. 15, 1928; Fassett 7907, sandy shore of small lake
2 miles northeast of Sand Lake, 12 miles northwest of Minong, Sept. 6,
1928; Fassett 7911, wet, sandy shore of Sand Lake, 12 miles northwest
of Minong, Sept. 6, 1928; Fassett 7912, at water's edge, sandy shore of
Wood Lake, 12 miles northwest of Minong, Sept. 6, 1928; Fassett
11971, sandy shore of same lake, same date; Fassett 17644, bog, Devil's
Lake, Sarona, Sept. 14, 1935 (2 sheets) ; Fassett 17645, sandy shore of
Bass Lake, Sarona, Sept. 14, 1935; Fassett & W. T. McLaughlin 10891,
shore of Mathews Lake, Trego, Sept. 11, 1929; Fassett & McLaughlin
10892, shore of Cable Lake, Spooner, Sept. 12, 1929; Fassett & Mc-
Laughlin 10893, sandy shore of McKinley Lake, Trego, Sept. 12, 1929;
Fassett & McLaughlin 10894, shore of Cable Lake, Spooner, Sept. 12,
1929; Fassett & MeLaughlin 10895, sandy shore of Horseshoe Lake,
Minong, Sept. 12, 1929; Fassett & McLaughlin 10896, shore of McLain
Lake, Minong, Sept. 12, 1929; Earl E. Sherff XY 6002, sandy and stony
east shore of Silver Lake, south of Minong, Sept. 1, 1961; Sherff
XY6002, sandy west shore of Shell Lake, Shell Lake, Sept. 1, 1961;
Sherf XY 6004, sandy shore of Little Long Lake, west of Spooner,
Sept. 1, 1961; Homer Stevens 187, bog beyond “Hogin’s” on Route M,
Aug. 28, 1947. BURNETT COUNTY: N. C. Fassett 1214, sandy shore of
Devil's Lake, Webster, Sept. 6, 1929; Fassett 7898, sandy shore of Long
Lake, Hertel, Sept. 19, 1928; Fassett 7899, sandy shore of Crooked
Lake, Siren, Sept. 19, 1928; Fassett 7902, sandy shore of Clear Lake,
Siren, Sept. 19, 1928; Fassett 7904, shore of Hanscon [Hanscomb]
Lake, Webb Lake, Sept. 21, 1928; Fassett 7905, sandy shores of Long
Lake and Sucker Lake, Webb Lake, Sept. 21, 1928; Fassett & J. W.
Thomson 18715, shore of Fish Lake, south of Webb Lake, Sept. 17,
1957; W. T. McLaughlin 1018, sandy shore of Clear Lake, Siren,
Aug. 21, 1929; McLaughlin 1019, sandy shore of Conners Lake, Web-
ster, Aug. 19, 1929; McLaughlin 1025, sandy shore of Macaboyne Lake,
Webb Lake, Aug. 18, 1929; McLaughlin 1028, sandy shore of Devil's
Lake, Webster, Sept. 1, 1929; McLaughlin 1032, sandy shore of Fish
Lake, Webb Lake, Aug. 17, 1929; McLaughlin 1033, sandy shore of a

1962] Sherff — Distribution of Bidens connata 25

small lake, Webb Lake, Sept. 11, 1929; McLaughlin 1035, sandy shore
of Ham Lake, Webster, Sept. 10, 1929; McLaughlin 1036, sandy shore
of Long Lake, Webb Lake, Sept. 11, 1929; McLaughlin 1037, sandy
shore of Sucker Lake, Webb Lake, Sept. 11, 1929; McLaughlin 1038,
sandy shore of small lake north of Sand Lake, Gaslyn, Sept. 4, 1929;
McLaughlin 1039, sandy shore of Loon Lake, Danbury, Sept. 3, 1929;
McLaughlin 1040, sandy shore of Oak Lake, Gaslyn, Aug. 24, 1929;
McLaughlin 1041, sandy shore of Long Lake, Hertel, Aug. 15, 1929;
McLaughlin 1042, sandy shore of Birch Island Lake, Birch Island,
Aug. 17, 1929; Earl E. Sherff XY 6005, sandy shore of Crooked Lake, 1
mile north of Siren, Sept. 2, 1961, a form approaching in its foliage
var. gracilipes Fern.; Sherf XY 6007, sandy shore of Benach Lake
(N. W. corner of sect. 29, tshp. 40N., R.15W.), +27 miles (by road)
northwest of Spooner, Sept. 2, 1961; Sherf XY 6008, sandy shore of
Mallard Lake, Sept. 2, 1961; Sherff XY 6009, sandy shore of small lake
near Mallard Lake, Sept. 2, 1961; Sherff XY6010, very abundant on
sandy north shore of North Sand Lake, northwest of Gaslyn, Sept. 2,
1961. BAYFIELD COUNTY: N. C. Fassett 7884, sandy shore of McDonald
Lake, Barnes, Sept. 11, 1928; Fassett 7894, same place and date; Fas-
sett 7903, sandy shore of Pigeon Lake, west of Drummond, Sept. 3,
1928; Fassett 7914, sandy shore of Lake Ruth, Iron River, Sept. 3,
1928; Fassett 7915 (1-2 meters from water, sandy shore of) and 7916
(at water's edge of) Island Lake, Barnes, Aug. 29, 1928; Fassett 7946,
muddy shore of Lake Ellison, Barnes, Sept. 2, 1928; W. T. McLaughlin
1017, sandy shore of Lake Ruth, Iron River, Sept. 13, 1930; Earl E.
Sherf XY 6001, sandy south shore of Pigeon Lake, west of Drummond,
Sept. 1, 1961. DOUGLAS COUNTY: G. H. Conklin, sandy shore of White-
fish Lake, Aug. 11, 1929; N. C. Fassett 7892, sandy shore of Wilderness
Lake or Pattangel Lake,* Barnes, Sept. 15, 1929; Fassett 7897, sandy
shore of Murray Lake, 12 miles east of Solon Springs, Sept. 2, 1928;
Fassett 7908, under eaves of shed on sandy shore of Loon Lake,
Wascott, Sept. 6, 1928 (2 sheets, "a" and '"b"); Fassett 7909, wet,
sandy shore of Whitefish Lake, Wascott, Sept. 6, 1928; Fassett 7910,
same place and date; Fassett 11970, sandy shore of Halfway Lake,
Gordon, Aug. 18, 1929; W. T. McLaughlin 1016, sandy shore of Simms
Lake, Sept. 13, 1930; McLaughlin 1027, sandy shore of Halfway Lake,
2 miles south of Gordon, Sept. 13, 1930.

Particular notice was given the var. pinnata specimens in
the live state to see if there were transitions to the less com-
pound types of leaves characteristic of B. connata var.
gracilipes Fern., a variety found in similar habitats, edges of
sandy lake shores, at various localities in Wisconsin and

*Fassett’s label gives the location for the lakes as Sect. 6, T. 45 N. R. 8 W. im
Douglas County, but Barnes itself is across the county border in Bayfield County.

26 Rhodora [Vol. 64

often interspersed with var. pinnata. No definite transitions
of var. pinnata into var. gracilipes were found.

As previously stated in my revisional treatment of Bidens
(Sherff, The Genus Bidens, Bot. Ser. Field Mus. Nat. Hist.
16: 262. 1937), the var. pinnata was known to range geo-
graphically through parts of Minnesota (where the type
had been collected at Richfield in Hennepin County) and
Wisconsin. On several trips in the autumn of 1961 to lakes
in western Michigan, I was unable to find var. pinnata, even
though the lakes possessed sandy shores identical with those
of the Wisconsin habitats. So far, then, as I have been able
to determine, var. pinnata is limited to Minnesota and Wis-
consin.

Special attention was given the same season also to the
occurrence of the var. gracilipes Fern. This variety is ex-
tremely interesting in the matter of its distribution. As was
stated in my above-cited monograph of Bidens (p. 261), the
type of var. gracilipes was a plant collected at Harwich, far
out on Cape Cod, Massachusetts. A few other specimens
were found, ranging from Maine to Connecticut. From the
known facts at that time, therefore, var. gracilipes seemed
to be an endemic variety of the northeastern United States.
The researches of Fassett upon the Wisconsin Flora, how-
ever, revealed the presence of the variety at numerous sites
in Wisconsin. Some specimens were cited by him in the
paper referred to above (Rhodora /oc. cit.) and many addi-
tional ones were collected by him later on, either singly or
in the company of his students, and deposited in the herbari-
um of the University of Wisconsin. It was my privilege to
examine all of these recently and to confirm his determina-
tions.

Subsequent to Fassett's publications, Mr. and Mrs. Hanes,
in their Flora of Kalamazoo County, Michigan (page 259.
1947) cited var. gracilipes for Kalamazoo County, Michigan.
Later, I myself cited (Brittonia 11: 190. 1959) a Hanes
specimen for the same county. Again, in Rhodora of May,
1961, I cited material, C. W. Bazuin 4282, from the "sandy
shore of Wolf Lake, 3-4 miles east of Muskegon, Michigan,"
a habitat "roughly seventy miles farther north and twenty

1962] Sherff — Distribution of Bidens connata 27

miles farther west than the habitat" where the Kalamazoo
County specimen had been collected. Unfortunately the last
two papers had been compiled with the thought of collab-
orating in the production of a new Flora of Michigan, which
authorities at the University of Michigan contemplate pub-
lishing (vide Mich. Flora Newsletter no. 7, Univ. of Mich.,
Nov. 25, 1960) and thus I inadvertently described the Wolf
Lake habitat as being "the farthest point west at which the
variety .... was known to occur." The words “in Michigan"
should have been added after “west” of course, to indicate
a cognizance of the many habitats known to occur for the
variety in Wisconsin. — Having the entire United States in
mind, then, we may put Wisconsin as marking the western
limit of distribution for var. gracilipes. Indeed, from the
comparative abundance of var. gracilipes specimens in Wis-
consin, it seems very probable that Wisconsin was the real
center of origin for the variety, and that specimens found in
New England states had originally grown there from
achenes transported from the west, presumably with the
aid of their retrorsely barbed aristae.

Returning to the occurrence of var. gracilipes in Michigan,
I have been able, in the autumn of 1961, to find specimens of
it growing in a seemingly discontinuous distribution begin-
ning with Kalamazoo County at the south and extending
northeastwardly into Ionia County, thence northwestward
into Kent County and Muskegon County, thence far north-
northeastward into Lake County, to a point (Wolf Lake)
some seven miles north of Baldwin. Ample material was
obtained for herbarium specimens and was turned over to
the Chicago Natural History Museum for accession and dis-
tribution to other institutions. A list of exsiccatae by coun-
ties follows:

KALAMAZOO COUNTY: E. E. Sherff XY6016, sandy north shore of
Eagle Lake,* Oct. 5, 1961: Sherf XY 6017, sandy shore of east side of
Eagle Lake, same date. IONIA COUNTY: Sherf XY 6015, sandy spot on
north shore of Jordan Lake, Lake Odessa, Sept. 29, 1961. KENT COUN-
TY: Sherf XY6012, sandy shore of Long Lake, east of Kent City,
Sept. 18, 1961. MUSKEGON COUNTY: Sherff XY 6013, sandy southwest

*This and the next references are to the Eagle Lake southwest of Oshtemo.

28 Rhodora [Vol. 64

shore of Wolf Lake, Sept. 18, 1961. LAKE COUNTY: Sherf XY6011,
sandy east shore of Wolf Lake, +7 miles north of Baldwin, Sept. 6,
1961.

Throughout my examinations of var. gracilipes in the field
at the above-mentioned Michigan habitats, attention was
paid to the foliar variations in the direction of var. pinnata.
Such variations were found to be rare. In the suite num-
bered XY6016, composed of some twenty or more small,
depauperate plants, two were found to have several leaves
each with two pairs of slenderly lanceolate lateral leaflets
or pinnae, instead of a single pair of broader ones. Ap-
parently this was the maximum extent to which var. gra-
cilipes approached var. pinnata. To summarize, then, while
the var. pinnata, as stated earlier, was not found to inter-
grade into var. gracilipes, the latter variety was found,
though very rarely, to offer a slight approach in leaf-form to
var. pinnata. There was no indication, however, that the two
varieties were not distinct or worthy of being accorded
separate status. — CHICAGO NATURAL HISTORY MUSEUM.

THE ANNUAL SPECIES OF HOUSTONIA IN ILLINOIS
ROBERT H. MOHLENBROCK AND LEO E. HALBIG

The genus Houstonia in northeastern North America is
represented by two well-defined groups. One group contains
delicate, vernal-flowering annuals with single-flowered
peduncles, salverform corollas, and globular seeds. The
other contains coarser, summer-flowering perennials with a
compound inflorescence, funnelform corollas, and rather
flattened seeds.

This paper is concerned with the annual vernal species in
Illinois. A subsequent paper will be devoted to the perennial
species.

Four or five species of the annual bluets occur in north-
eastern North America. (The discrepancy arises from the
status of H. faxonorum Fern. or H. caerulea var. faxonorum
Pease and Moore). Three of these are attributed to Illinois
in varying degrees of abundance by different workers.

1962] Mohlenbrock and Halbig — Houstonia 29

Houstonia serpyllifolia Michx., from the mountains of Penn-
sylvania south to Georgia, does not occur in Illinois.

A check of any manual concerning the Illinois flora would
lead one to believe that H. caerulea, H. pusilla, and H.
minima are more common than is actually the case.

The latest treatment on distribution of Illinois vascular
plants is that by Winterringer and Evers'. These workers
list H. caerulea from ten Illinois counties along the eastern
border and the southern fourth of the state. The present
study reveals that five of the six reports from southern
Illinois are based on misidentifications. The sixth report
from St. Clair County late last century is likely a case of
mixed labels since the collector often was guilty of this.

Winterringer and Evers attribute H. minima to 17 Illinois
counties, including 3 in the southern tip of the state. Our
study shows H. minima to occur in 17 counties, but to be
absent in the extreme southern counties of Jackson and
Saline. Records from these counties are based on misidenti-
fications for H. pusilla.

Houstonia pusilla is limited in Illinois to the southern one-
third of the state. The present study has enabled us to fill in
most of the gaps in its distribution in southern Illinois.

Since all three species are reported to occur in southern
Illinois with seeming regularity, the senior author thought
it strange that on his many field trips and those of his col-
leagues and students, no authentic material of two of these
— H. caerulea and H. minima — showed up in the collec-
tions.

During the spring of 1961, the junior author made vast
collections of annual bluets throughout southern Illinois.
These were studied intensively before being pressed for the
herbarium. In the entire collection of several hundred speci-
mens, there appeared no H. caerulea and no H. minima. All
specimens were unquestionably H. pusilla.

Why, then, are there so many apparently erroneous re-
ports of H. caerulea and H. minima? There is general agree-
ment among field botanists that each species has a distinctive

'Winterringer, G. S. and R. A. Evers. 1960. New Records for Illinois Vascular
Plants. Illinois State Museum, 135 pp.

30 Rhodora [Vol. 64
field recognition character. Houstonia caerulea is distin-
guished by the yellow eye or center in each flower, while
the other species basically have a dark, usually reddish eye.
A hurried glance at the flower either of H. pusilla or H.
minima might reveal a yellow center, due to the presence of
pollen on the mature anther, and thus this may account for
the erroneous reports of H. caerulea in southern Illinois.
Anyone familiar with the larger, pale-flowered H. caerulea,
with its conspicuous yellow center, could never confuse this
species with any other.

Houstonia minima sometimes is recognized in the field by
its tiny stature. In our study, we have found H. pusilla as
dwarfed as 1 cm. tall, during anthesis. Local naturalists, on
being shown this specimen, immediately respond with H.
minima. The only valid criterion to distinguish H. minima
and H. pusilla is not stature, but relative length of calyx to
corolla. Houstonia minima has a calyx equalling the corolla
tube in length, while H. pusilla has a calyx barely half the
length of the corolla tube. Although no specimens of H.
minima were discovered in the field in southern Illinois dur-
ing this study, there exists an authentic specimen in the
University of Illinois herbarium of this species from John-
son County. Efforts to relocate this population have failed.

Since most modern manuals fail to give complete descrip-
tions of H. caerulea, H. pusilla, and H. minima, it seems
desirable to present descriptions in this paper.

KEY TO THE ANNUAL SFECIES OF HOUSTONIA IN ILLINOIS
1. Flowers pale blue or lilac, with a broad, pale yellow center; corolla
tube 5-10 mm. long; capsule 3-4 mm. wide .................. 1. H. caerulea.
1. Flowers purple, with a small dark, usually red, eye; corolla tube
3-5 mm. long; capsule 2-3 mm. wide.

2. Calyx lobes 1.2-1.9 mm. long, half as long as corolla tube ............
:MRMMMMMNMMMMMMMMMMMMMMMMMWMMMMWZMMMMMMMMP 2. H. pusilla.

2. Calyx lobes 2.0-3.5 mm. long, equalling the corolla tube ................
3. H. minima.

1. Houstonia caerulea L. Sp. Pl. 105. 1753.

Slender, tufted annuals; stems erect, flexuous, rather sparsely
branched, to 15 em. tall (in Illinois), glabrous or rarely puberulent ;
rosette leaves obovate to elliptic to spatulate, obtuse to subacute at
apex, 4-15 mm. long, 2-6 mm. broad, glabrous or short hairy; petiole
1-6 mm. long, glabrous or sparsely puberulent; cauline leaves much

1962] Mohlenbrock and Halbig — Houstonia 31

reduced, 2-7 mm. long, 1-3 mm. broad; peduncles filiform, 2-7 em. long,
glabrous or nearly so; bracts resembling the cauline leaves; sepals
oblong, acute, 1.0-2.5 mm. long, glabrous; corolla salverform, pale blue
to lilac to rarely nearly white, with a large yellow center (eye), the
tube 5-10 mm. long, glabrous, the flower 10-15 mm. wide; capsule bi-
lobed, flattened, 3-4 mm. wide, broader than long.

Habitat. — Open woods and fields.

General Range. — Nova Scotia to Wisconsin south to Missouri and
Georgia.

Illinois Range. — Four eastern border counties; absent from the
extreme south.

Date of Flowering in Illinois. — April 7 to May 17.

2. Houstonia pusilla Schoepf, in Reise 2:306. 1788.

Houstonia patens Ell. Bot. S. C. & Ga. 1:191. 1816.

Usually tiny, tufted annuals; stems erect, stiffer than the preceding,
1-12 cm. tall, glabrous; rosette leaves ovate to elliptic, subacute to
acute, 5-10 mm. long, 3-7 mm. broad, glabrous and usually scaberulous
along the margins; cauline leaves smaller and narrower; petiole 1-3
mm. long, glabrous; peduncles 1-4 cm. long, glabrous or nearly so;
bracts resembling the upper cauline leaves; sepals linear to narrowly
oblong, acute, 1-2 mm. long, glabrous; corolla salverform, deep purple,
usually with a dark reddish center (eye), the tube 3-5 mm. long, about
twice the length of the calyx, glabrous, the flower 4-8 mm. wide;
capsule bilobed, flattened, 2-3 mm. wide, slightly longer than broad.

Habitat. — Xeric sandstone bluffs, open woods, fields, and pastures.

General Range. — Virginia to Missouri south to Texas and Florida.

Illinois Range. — The southern one-third of the state, north to
Montgomery County.

Date of Flowering in Illinois. — March 22 to May 4.

Five-merous flowers sometimes occur.

3. Houstonia minima Beck, in Am. Journ. Sci. 10:262. 1826.

Tiny, tufted annuals; stems erect, 1-7 cm. tall, glabrous; rosette
leaves ovate to elliptic, subacute to acute, 5-7 mm. long, 2-5 mm. broad,
glabrous and usually scaberulous along the margins; cauline leaves
smaller and narrower; petiole 1-3 mm. long, glabrous; peduncles 1-4
cm. long, glabrous or nearly so; bracts resembling the upper cauline
leaves; sepals lanceolate to ovate-lanceolate, acute, 2-4 mm. long,
glabrous or scaberulous; corolla salverform, deep purple, usually with
a dark reddish center (eye), the tube 3-5 mm. long, barely longer than
the calyx, glabrous, the flower 3-6 mm. wide; capsule bilobed, flattened,
2-3 mm. wide, slightly longer than broad.

Habitat. — Open woods and fields.

General Range. — Illinois tc Kansas south to Texas and Arkansas.

Illinois Range. — Essentially the western counties, although one
record from Richland County.

Date of Flowering in Illinois. — April 6 to May 19. — SOUTHERN
ILLINOIS UNIVERSITY, CARBONDALE.

32 Rhodora [Vol. 64

WATERCRESS IN THE NEW WORLD
PETER S. GREEN

Fifteen years ago Howard and Manton (1946) put on
record the existence of both a diploid and tetraploid species
of watercress and distinguished them under the names
Nasturtium officinale R. Br. (diploid, 2n = 32) and N. uni-
seriatum Howard & Manton (allotetraploid, 2n — 64). The
following year, Airy Shaw (1948) elucidated the nomencla-
ture and showed that the name N. microphyllum Boenn. ex
Reichb. should be used for the tetraploid on grounds of
priority. Since this date it has become widely accepted that
on a world basis Nasturtium is not distinguishable generical-
ly from Rorippa and the names of the watercresses then
become R. nasturtium-aquaticum (L.) Hayek (N. officinale
R. Br.) and R. microphylla (Boenn.) Hyland, whilst for
simplicity of reference the sterile triploid hybrid between
the two species has been given the name R. X sterilis Airy
Shaw (Shaw 1951).

Both species are natives of Europe, but both have been
introduced into the New World, presumably because of their
use and value as a salad. Airy Shaw (1948) first recorded
the existence of the tetraploid in Canada and the U.S.A.,
and Green (1955, p. 297-8) added further New World records
and noted the existence in the U.S. A. of the sterile hybrid
as well. A recent examination of the material in the Gray
and New England Botanical Club Herbaria however, has
indicated that watercress is far more widely distributed than
had at first been expected, and it became apparent that the
two species have not been distinguished in American litera-
ture. To take the two major Floras in the north-east:
Gleason (1952) continued to describe Nasturtium as mono-
typic in 1952, and the figure he published is clearly that of
the diploid species. Fernald (1950), however, distinguished
a var. microphyllum of N. officinale, but it is clear from his
description, and from annotations he made on specimens in
the two herbaria just mentioned, that whilst his concept of
var. microphyllum is very close to the tetraploid species it
is not entirely coincident, and his figure, labeled as N. offici-

1962] Green — Watercress in the New World 33

nale, shows fruit clearly recognizable as those of Rorippa
microphylla. In the two handbooks for the identification of
North American aquatic plants: Muenscher (1944) has one
species, which he calls N. officinale, and his illustration is
clearly that of the tetraploid. Fassett (1940 and 1957)
recognizes four named forms based on vegetative characters
but he does not mention the name microphyllum. Murley in

Fic. 1. Fruit and inflorescence of: A, Rorippa nasturtium-aquaticum (L.) Hayek;
B, R. microphylla (Boenn.) Hyland. Half natural size,

her review of the “Seeds of the Cruciferae of North-eastern
North America” (1951) only considers the one species, as
Nasturtium officinale, and gives a beautiful illustration of a
seed of the diploid (her fig. 8).

There is no doubt that both Rorippa nasturtium-aquati-
cum and R. microphylla are vegetatively both very variable.

34 Rhodora [Vol. 64

For certain and easy distinction it is necessary to use charac-
ters of the fruit. In fact, unless the plants are in fruit it is
difficult to differentiate between the two species (see Howard
& Lyon 1950). In R. nasturtium-aquaticum the siliquae are
10-15(-20) mm. long and relatively stout, whereas in R.
microphylla they are longer, (14-) 17-26, mm. long, and more
slender (Fig. 1). The fruits are also borne on slightly shorter
pedicels in R. masturtium-aquaticum where they are
6-15(-18) mm. long compared with (6-)10-20 mm. in R.
microphylla. A field character is further provided by the
fact that the seeds appear in R. nasturtium-aquaticum to

Fic. 2. Individual seed of: A, Rorippa nasturtium-aquaticum (L.) Hayek; B, R.
microphylla (Boenn.) Hyland.
lie in two rows in the pod, whereas in R. microphylla they lie
in a single row, (hence the epithet wniseriatum proposed by
Howard & Manton for the tetraploid) but this character has
to be used with caution. A character of sure value, however,
is provided by the reticulations, or areolae, on the seed coat,
which in R. nasturtium-aquaticum are large, and number
about 35 on any one side of the ripe seed, whilst in R. micro-
phylla there are about 150 and these are consequently much
smaller (Fig. 2). It is interesting in this connection that a
photograph of seeds of R. nasturtium-aquaticum is given in
one of the plates bound in the center of the 1961 yearbook of

1962] Green — Watercress in the New World 35

the U. S. Department of Agriculture. If plants to be identi-
fied have not reached the fruiting stage then it is possible to
distinguish between the species on pollen grain size (Green
1955) and stomatal index (Rowson in Howard & Manton
1946) but the methods are somewhat time consuming.

The hybrid Rorippa X sterilis is, as its name suggests,
easily distinguished by its sterility. The fruits do not devel-
op properly and only an occasional, non-viable, seed is pro-
duced. Furthermore, examination of the pollen grains in,
for example, a drop of lactic acid, shows that the pollen too
is sterile, the grains being very variable in size and the
majority of them crumpled and abortive. In fact to be
certain of the hybrid it is usually advisable to examine the
pollen, for occasional plants of the parental species set little
seed from the first flowers to open and, especially in R.
microphylla, the plant may set very little seed at all if grow-
ing under crowded conditions etc.

Because of the variability of vegetative characters the
epithet microphylla is misleading; leaf shape and size are of
no value in distinguishing the species and their hybrid.
Large-leaved specimens are found with fruits of R. micro-
phylla, and depauperate specimens with small leaves are, as
often as not, found to be R. nasturtium-aquaticum. A variety
siifolia has been described within R. nasturtium-aquaticum
and was recognized by Fernald (1950) along with so-called
var. microphylla. In it the leaflets are more or less oblong
and elongate as opposed to roundish to oval, especially in the
case of the terminal leaflet, but such leaves are frequently
found in the diploid, tetraploid and hybrid and have no
taxonomic significance, at least at the rank of variety.

The distribution of watercress in the U.S. A. and im-
mediately adjacent territories as reflected in the collections
of the Gray and New England Botanical Club Herbaria is
shown in Fig. 3 (one or two records have been added from a
hasty examination of the collections in the herbarium of
the New York Botanical Garden made during a recent brief
visit). It will be seen that Rorippa nasturtium-aquaticum is
far more widespread than both R. microphylla and the
hybrid, which, apart from two records in Idaho and Oregon,

36 Rhodora [Vol. 64

are restricted to eastern Canada and northeastern United
States. How many times watercress has been introduced
from Europe is unknown, but in view of the esteem in which
it has long been held as a salad in England, France and
Germany, it is suspected that it has been brought across the
Atlantic many times, the first occasion probably having
taken place more than 200 years ago. Even from Utah there

@ nasturtium-aquaticum, A microphylla

X sterilis, O unidentified watercress

Fic. 3. Distribution map of the watercresses in the U.S.A. and adjacent territories.
In the north-east several closely adjacent records may be represented by a single
symbol.
is a specimen in the Gray Herbarium collected by E. Palmer
as long ago as 1875 which bears a note “introduced from
France by J. E. Johnson” and one may believe that the
introduction to the eastern states took place long before this,
although the earliest specimen I have seen was collected in
1847 from Niagara, New York. The diploid, R. nasturtium-
aquaticum, is far more widespread than the tetraploid
species and is recorded from almost every state. Muenscher
(1944) published a map in which it is recorded from every

1962] Green — Watercress in the New World di

state but six and I have seen material from three of these six
in the course of this investigation. It is debatable whether
watercress was intentionally introduced to every state where
it now occurs and although it frequently has the appearance
of a native plant and is often found remote from any habita-
tion, it must be borne in mind that it commonly grows on
muddy stream banks and seed may be relatively easily trans-
ported over long distances embedded in mud attached to the
feet of birds, and broken portions of the plant can easily
be carried down stream once it is established in a particular
river or stream. The records of R. microphylla and the
hybrid in the west almost certainly represent separate intro-
ductions and it is not without significance perhaps that at
least in recent times the watercresses which are grown
commercially in Britain (see Howard 1947) are the diploid,
R. nasturtium-aquaticum, which is the most widespread in-
troduction, and the hybrid. It is possible too that diploid
R. nasturtium-aquaticum is more able to flourish in the
warmer more southern states, and this is suggested by its
distribution outside the United States, whereas the tetra-
ploid is probably the hardier species and the one more able
to survive the rigorous winters in the north-east. No doubt
the examination of watercress material in other herbaria
and the painstaking measurement of pollen grains in
material showing only flowers would increase our knowledge
of the detailed distribution of the species but it is felt that
sufficient indication of the total range is given by the map
(fig. 3) and by the citation of the specimens below, to draw
the attention of botanists in America to the existence of the
two species and how they may be distinguished.

In Central America, the West Indies and South America
only material of Rorippa nasturtium-aquaticum in the strict
sense has been seen. There are three sheets of apparently
sterile plants from Cuba in the Gray Herbarium, but exami-
nation of the pollen shows that the grains are fully formed
and not largely crumpled and abortive as in the hybrid, R.
X sterilis. However, the fruits seem undeveloped and empty,
yet in each case the whole plant appears small and the leaves
reduced, with few pinnae; it is felt that further comment on

38 Rhodora [Vol. 64

their identity should be reserved until more is known of the
conditions under which they were growing. Schulz (1934,
p. 41) recorded Nasturtium officinale var. microphyllum
from Chile, but this identification was apparently based on
leaf shape alone and in this investigation only the diploid
has been seen with certainty from Chile, or for that matter,
from the whole of South America.

MATERIAL EXAMINED*

Rorippa nasturtium-aquaticum (L.) Hayek
(Nasturtium officinale R. Br.)

CANADA. British Columbia: Comox, July 18, 1915, J. M. Macoun
91854.

UNITED STATES. Maine: Cumberland Co., Yarmouthville, Yar-
mouth, Aug. 1904, Kate Furbish (NEBC). Vermont: Rutland Co., Rut-
land, June 15, 1892, Willard W. Eggleston 1. Massachusetts: Berkshire
Co., Pittsfield, Sept. 23, 1899, R. Hoffmann; Norfolk Co., Dedham,
Purgatory Swamp, June 27, 1897, J. M. Greenman 260; Nantucket
Co., Nantucket, Polpis, June 1, 1900, M. A. Day 2 (GH, NEBC). Rhode
Island: Newport Co., Block Island, June 24, 1917, R. P. Marshall
(NEBC). Connecticut: Fairfield Co., Bridgeport, June 3, 1902, E. H.
Eames (NEBC) ; Hartford Co., Southington, June 30, 1898, L. Andrews
311; Litchfield Co., Woodbury, June 12, 1909, E. B. Harger 5548
(NEBC) ; New Haven Co., North Branford, Aug. 5, 1914, E. B. Harger
6415 (NEBC); New London Co., Franklin, June 18, 29, 1915, R. W.
Woodward. New York: Niagara Co., Niagara Falls near Table Rock,
June 1847, John A. Lowell; Wayne Co., Newark, Aug. 31, 1872, E. L.
Hankenson. Pennsylvania: Berks Co., Hamburg, July 19, 1892, C. D.
Lippincott; Chester Co., 1858-1864, S. P. Sharples; Lebanon Co.,
Miners Village, May 30, 1889, A. A. Heller. Delaware: New Castle
Co., Centreville, May 20, 1865, A. Commons. Maryland: Montgomery
Co., shores of Potomac, June 6, 1881, John Donnell Smith. W. Virginia:
Munroe Co., Sweet Springs, Sept. 15, 1903, E. S. and Mrs. Steele 315;
Pocahontas Co., Minnehaha Spring, July 31, 1930, W. V. U. Botanical
Exped. Virginia: Botetourt Co., E. Buchanan, along the Otter Rd.,
May 17, 1892, John K. Small; Page Co., Luray, Stony Man Mountain
and vicinity in the Blue Ridge, Aug. 12, 1901, E. S. and Mrs. Steele
161. North Carolina: Avery Co., Cranberry, on US 19E, June 17, 1958,
Harry E. Ahles & J. A. Duke 43572; New Hanover Co., Carolina Beach,
April 18, 1938, R. K. Godfrey & M. F. Buell 3542. Florida: Sarasota
Co., Sarasota, March 31, 1943, Anne E. Perkins. Michigan: Wash-
tenaw Co., edge of Fleming Creek, 434 mi. east of Ann Arbor, F. J.
Hermann 6950. Ohio: Portage Co., Garrettsville, July 18, 1896, R. J.
Webb 156. Indiana: Tippecanoe Co., 0.5 mi. s.w. of Lafayette, west

*Specimens in the Gray Herbarium unless otherwise stated.

1962] Green — Watercress in the New World 39

bank of Wabash River valley, June 2, 1945, Ray C. Friesner 18859.
Kentucky: Jefferson Co., near Beuchel, July 5, 1939, M. Seargent 49;
Wayne Co., Beaver Creek, s.w. of Monticello, July 12-14, 1937, L. B.
Smith & A. R. Hodgdon 3868. Tennessee: Davidson Co., Nashville,
June 14, 1960, Howard S. Gentry 18589. Wisconsin: LaCrosse Co.,
Bohemian Creek, Thomas A. Hartley 778 (Ny). Illinois: Kane Co.,
Elgin, June 9, 1911, Earl E. Sherff; McHenry Co., McHenry, June 25,
1860, George Vasey 2105. Minnesota: Dakota Co., railroad near Nicols,
Sept. 19, 1941, J. W. Moore, E. K. Butters & D. Jenkins 15113. Iowa:
Fayette Co., Fayette, Aug. 1894, B. Fink. Missouri: Clay Co., Ran-
dolph, June 20, 1897, Kenneth K. MacKenzie (NY). South Dakota:
Fall River Co., Black Hills, Hot Springs, June 14, 1892, P. A. Rydberg
531. Nebraska: Dundy Co., Rock Creek Park, Aug. 3, 1945, Walter
Kiener 19417; Redwillow Co., 8 miles west of McCook, July 31, 1945,
Walter Kiener 19411. Kansas: Riley Co., 1896, J. B. Norton 613.
Oklahoma: Carter Co., between Ardmore and Springer, April 29, 1961,
Reed C. Rollins 61156; Comanche Co., Fort Sill, June 16, 1916, Mrs.
J. Clemens 11601; Ellis Co., on edge of Wolf Creek near Shattuck,
June 10, 1914, R. L. Clifton 3200K; Love Co., 2 miles s.w. of Bomar,
June 23, 1953, G. J. Goodman & E. L. Rice 5686. Texas: Brewster Co.,
Leoncita Springs, Kokernot ranch, April 30, 1948, Barton H. Warnock
& F. M. Churchill 7725; Gonzales Co., Cottonwood Springs, April 16,
1934, V. L. Cory 8271; Jeff Davis Co., Limpia Canyon 10 miles north
of Ft. Davis, June 13, 1941, R. R. Innes & Brunelle Moon 1095; Kerr
Co., Kerrville, May 14-21, 1894, A. A. Heller 1753; Real Co., Prade
Ranch, headwaters of Rio Frio, April 18, 1959, R. C. Rollins & D. S.
Correll 5940; Travis Co., Austin, May 15, 1872, Elihu Hall 14; Val-
verde Co., along the Rio Grande from Brownsville to El Paso, Devil's
River, April 20, 1919, H. C. Hanson 523. Montana: Gallatin Co., Boze-
man, July 10, 1902, W. W. Jones. Idaho: Ada Co., Boise, Aug. 19, 1911,
June A. Clark 257; Canyon Co., Falk's Store, July 7, 1910, J. F. Mac-
bride 327; Teton Co., Victor, July 11, 1901, E. D. Merrill & E. N. Wil-
cox 1015; Nez Perce Co., Upper Ferry, above Lewiston, Clearwater
River, June 2, 1892, J. H. Sandberg, D. T. MacDougal & A. A. Heller
298. Wyoming: Albany Co., Laramie, Sept. 30, 1984, Aven Nelson
1152; Crook Co., Sand Creek near Beulah, June 21, 1950, C. L. Porter
5349; Platte Co., Sibyllee Creek, July 18, 1945, C. L. Porter 3692.
Colorado: La Plata Co., T. 34 N., R. 8 W., Aug. 14, 1937, Marion Own-
bey 1439. Utah: Cache Co., half mile west of Logan, July 30, 1940,
Bassett Maguire 20099; Salt Lake Co., Salt Lake City, 1869, Sereno
Watson 6; Utah Co., Spring Lake, 1875, E. Palmer. Nevada: Clark
Co., Willow Springs, Aug. 6, 1935, I. W. Clokey 5481; Mineral Co.,
Corey Canon. Wassuk Mts., June 27, 1919, Ivar Tidestrom 10084;
Ormsby Co., King's Canon, June 11, 1902, C. F. Baker 1057. New
Mexico: Colfax Co., vincinty of Ute Park, Sept. 10, 1916, Paul C.
Standley 14611; Grant Co., Fort Bayard Watershed, Stephen's Ranch,
Oct. 22, 1905, J. C. Blumer 112; San Miguel Co., near Pecos, Aug. 20,

40 Rhodora [Vol. 64

1908, Paul C. Standley 5129; Sierra Co., Berendo Creek, May 20, 1904,
O. B. Metcalfe 907; Socorro Co., Mogollon Mts., on or near the west
fork of the Gila River, Aug. 25, 1903, O. B. Metcalfe 606. Arizona:
Coconino Co., Painted Desert, Tuba Oasis, July 15-20, 1920, W. N.
Clute 107; Yavapai Co., 6 miles east of Prescott, June 29, 1928, Carl B.
Wolf 2375. California: Inyo Co., White Mts., Roberts Ranch, Wyman
Creek, July 21, 1931, Victor Duran 3140; Los Angeles Co., Claremont,
May 28, 1910, J. D. Taylor 133; Marin Co., Tomales Point, one mile
west of Indian Beach, May 24, 1941; H. L. Mason 12424; Mono Co.,
Lakeview Spring, 6.5 miles from Bridgeport-Sweetwater Highway,
Aug. 3, 1945, Ira L. Wiggins & R. C. Rollins 546; Monterey Co., Pacific
Grove, July 1, 1905, C. P. Smith 1005; Riverside Co.; Santa Anna
River at Chino Creek, May 28, 1933, L. C. Wheeler 1747; Santa Clara
Co., Palo Alto estate, May 4, 1894; Siskiyou Co., near the Box Canyon
of the Sacramento near Mt. Shasta City, July 12, 1940, W. B. Cooke
15321; Stanislaus Co., Modesto, July 13, 1935, R. F. Hoover 686.
Oregon: Curry Co., Port Orford near Battle Rock, June 4, 1928, J. W.
Thompson 4480; Klamath Co., near Ft. Klamath, Aug. 7, 1894, J. B.
Leiberg 665. Washington: Spokane Co. Clark Springs, Spokane,
July 8, 1902, F. O. Kreager 124; Walla Walla Co., Waitsburg, May 25,
1897, R. M. Horner B54; Whitman Co., Wawawai, June 4, 1892, Lake
& Hull 477. Alaska: Manley Hot Springs, near the Tanana River,
Sept. 15-19, 1949, Edith Scamman 5798.

MEXICO: Chihuahua: vicinity of Chihuahua, May 1-21, 1908,
Edward Palmer 184. Coahuila: Saltillo, June 1898, Edw. Palmer 223;
Durango: Durango, July 25, 1944, C. V. Morton 44191. Hidalgo:
Pachuca, near Zerezo and below Parque Nacional El Chico, May 12,
1947, H. E. Moore 2793. Jalisco: Guadalajara, May 15, 1901, C. G.
Pringle 8494. Michoacan: Zitácuaro, Sept. 5, 1935, Geo. B. Hinton et
al. 11848. Mexico: Vallee de Mexico, June 4, 1865-6, Bourgeau 18.
Puebla: Huauchinango, sand along Rio Necaxa, March 27, 1945, A. J.
Sharp 45369.

CUBA. Santa Clara Prov., Mina Carlota, southeast of Cumanay-
agua, Sierra de San Juan, March 21-23, 1938, H. A. Senn 332. Oriente
Prov., 1856-1857, C. Wright 7.

HAITI. San Michel to Marmelade, Aug. 6, 1905, George V. Nash &
Norman Taylor 1472 (NY).

DOMINICA. Prov. San Juan, El Cercado, Juan Santiago, Hondo
Valle, Sept. 1, 1946, R. A. € E. S. Howard 8743.

PUERTO RICO. Adjuntas, in mont “Cienega”, April 11, 1886, P.
Sintenis 4116.

VENEZUELA. Near Tovar, 1854-1855, A. Fendler 20.

COLOMBIA. Dept. of Cundinamarca, Bogota, Oct. 4-8, 1917,
Francis W. Pennell 2348.

ECUADOR, Prov. Canar, near village of San Marcos, Azogues,
April 1, 1945, Francisco Prieto E2479. Prov. Chimborazo, Canon of
the rio Chanchan near Huigra, May 7-14, 1945, W. H. Camp E3153.

1962] Green — Watercress in the New World 41

PERU. Dept. Lima, Prov. Chancay near Supe, Sept. 4, 1938, Alan
A. Beetle & O. B. Horton 9064. Dept. of Cusco, Pisae, April 30, 1925,
Francis W. Pennell 13718. Dept. of Junin, Oroya, 1919, Margaret
Kalenborn 60.

CHILE. Prov. Aconcagua, Valle de Marga-Marga, 1930-32, Felix
Jaffuel & Anastasio Pirion 3265. Prov. Atacama, Dept. Copiapó, vicini-
ty of Copiapó, Nov. 16, 1925, Ivan M. Johnston 4992. Prov. Coquimbo,
Dept. Elqui, El Colorada, 70 km. on road from Rivadavia to Laguna,
tributary dam of Laguna River, Feb. 16, 1940, R. Wagenknecht 18501.
Prov. Osorno, Puerto Octay, A orillas del lago Llanquihue, Nov. 29,
1939, Hugo Gunckel 9323. Prov. Santiago, Penalobu, Jan. 2, 1927,
G. Looser 79. Prov. Valdivia, Rinihue, Feb. 12, 1933, Hugo Gunckel
5821.

BRAZIL. S. Leopoldo, Oct. 1940, J. Eugenio Leite 1794. S. Rosa,
Sombrie, S. Catarina, Oct. 11, 1944, R. Reitz C767.

BOLIVIA. Dept. Cochabamba, Cuidad de Cochabamba garten,
Dec. 26, 1928, José Steinbach 8788. Dept. La Paz, La Paz, Oct. 10,
1921, Otto Buchtien 244.

URUGUAY. Dept. Motevideo, Cerro Casabo, Sept. 1926, G. Herter
443 (80926).

ARGENTINA. Prov. Buenos Aires, Partido de Tornquist, Sierra de
la Ventana, Albra de la Ventana, Nov. 7, 1938, A. L. Cabrera 4693.
Prov. de Cordoba, Unquillo, 1926, C. Bruch. Prov. of Tucuman: Dept.
Famaillá, Rio Colorado, Sept. 1919, S. Venturi 415. Terr. de Rio Negro,
Region del Lago Nahuel Huapi, Basiloche, Feb. 5, 1940, A. L. Cabrera
5965.

Rorippa microphylla (Boenn.) Hyland. (Nasturtium microphyllum

(Boenn.) Reichb., N. wuniseriatum Howard & Manton)

CANADA. Newfoundland: Waterford River between Waterford
Bridge and St. John's, Aug. 1, 1911, M. L. Fernald & K. M. Wiegand
5482. Ontario: Simcoe Co., Collingwood, Baie Georgienne, Aug. 26,
1933, F. Marie-Victorin, F. Rolland-Germain, & René Meilleur 45069;
Bruce Co., Mouth of Pine River, 6 mi. south of Kincardin, L. Huron,
June 19, 1948, J. H. Soper & H. M. Dale 3952.

UNITED STATES. New Hampshire: Strafford Co., along Oyster
River, near Northwood-Durham turnpike, July 4, 1943, A. R. Hodgdon
4565 (NEBC). Vermont: Bennington Co., Arlington, July 3-4, 1908,
W. W. Eggleston 3229 (NY). Massachusetts: Barnstable Co., Harwich,.
the head of Allen’s Harbor Creek, Aug. 6, 1919, M. L. Fernald &
Bayard Long 18502; Berkshire Co., North Adams, June 25, 1913,
M. L. Fernald & Bayard Long 9553 (GH, NEBC) ; Essex Co., Newbury-
port, “The Gully”, July 31, 1940, R. C. Bean (NEBC); Franklin Co.,
Greenfield, July 25, 1911, C. H. Knowlton (NEBC); Middlesex Co.,
Waltham, 1861, Wm. Boott; Norfolk Co., Wrentham, near north end
Archers Pond, Sept. 18, 1897, F. G. Floyd 374 (NEBC) ; Plymouth Co.,
West Hingham, July 25, 1888, Walter Deane (NEBC); Suffolk Co.,.
Jamaica Plain, Arnold Arboretum, Bussey Brook, June 27, 1927, E. J..

42 Rhodora [Vol. 64

Palmer 28003 (A). Rhode Island: Providence Co., Goosenest Brook,
Wickford Junction, July 31, 1909, Thomas Hope (NEBC). New York:
Tompkins Co., Ithaca, Dwyer's Pond, July 29, 1918, E. L. Palmer 564;
Cortland Co., Inlet to Mud Pond, McLean Wild Life Preserve, July 29,
1932, W. C. Muenscher 17921. Michigan: Emmett Co., Pickerel Lake,
east of Petoskey, July 6, 1933, H. A. Gleason 100. Idaho: Blaine Co.,
Picabo, July 3, 1916, J. F. Macbride & Edwin B. Payson 3009. Oregon:
Hood River, May 28 & July 1885, W. N. Suksdorf 506.

Rorippa X sterilis Airy Shaw

(Rorippa microphylla X R. nasturtiwum-aquaticum)

UNITED STATES. New Hampshire: Strafford Co., Durham, near
Madbury Line, Aug. 1, 1943, A. R. Hodgdon 4569 (NEBC). Connecticut:
Fairfield Co., Bridgeport, June 19, 1896, E. H. Eames; Hartford Co.,
East Hartford, June 3, 1893, C. A. Weatherby (NEBC); New Haven
Co., Oxford, July 21, 1896, E. B. Harger 200 (NEBC). Michigan: Mac-
kinac Co., Mackinac Island, July 30, 1924, F. W. Hunnewell 9308.
Idaho: Owyhee Co., Flint Creek, July 30, 1910, J. F. Macbride 492.

Two final comments are thought to be of value. Airy Shaw (1948)
recorded R. microphylla from California, however, he considered both
of the records he gave to be doubtful, and one, based on Copeland 368
from Jonesville, Butte Co., was found (Green, 1955) on examination
of the pollen grains not to be a watercress at all. Examination of a
duplicate of this in the Gray Herbarium, and comparison there with
the extensive collections from North America shows that it is a rather
depauperate specimen of Cardamine breweri S. Wats.

Finally, whilst not strictly concerning the watercresses of the New
World, the opportunity was taken to examine the material in the Gray
Herbarium from Japan. Previous examination of Japanese material
(Green, 1955) in the herbaria at Edinburgh and Kew showed all the
flowering specimens to be R. X sterilis alone. Three further collections
in the Gray Herbarium have now been seen and they too are each of
them the hybrid. (Hondo: June 2, 1929, K. Shiota 1044; May 19, 1935,
K. Shiota 8437. Shikoku: June 2, 1935, I. Yogo 9557). It is interest-
ing to see, however, that Kitamura and Murata (1961) list Nasturtium
officinale R. Br. and illustrate (their fig. 86 (i)) a typical fruit of the
diploid; their coloured illustration (plate 42) however, shows only
flowers and has no fruits. — ARNOLD ARBORETUM OF HARVARD UNIVER-
SITY.

LiTERATURE CITED
Fassett, N. C. 1940. A Manual of Aquatic Plants: 237.
1957. A Manual of Aquatic Plants, ed. 2: 257.
FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th edn.: 716.
GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora
of the Northeastern United States and Adjacent Canada, 2: 38.
GREEN, P. S. 1955. Pollen Grain Size in Nasturtium and Cakile.
Trans. Proc. Bot. Soc. Edinb. 36: 289-304.

1962] Schuyler — A new Species of Scirpus 43

Howarp, H. W. 1947. Wild and Cultivated Watercress Types. Agri-
culture, 53: 453-456.

HowaRD, H. W. AND A. G. Lyon. 1950. The Identification and Dis-
tribution of the British Watercress Species. Watsonia, 1: 228-233.

HowaARD, H. W. AND I. MANTON. 1946. Autopolyploid and Allopoly-
ploid Watercress with the Description of a New Species. Ann. Bot.
II. 10: 1-13.

KITAMURA AND MURATA. 1961. Coloured Illustrations of Herbaceous
Plants of Japan (Choripetalae): 177.

MUENSCHER, W. C. 1944. Aquatic Plants of the United States: 254.

MURLEYv, MARGARET R. 1951. Seeds of the Cruciferae of Northeastern
North America. Amer. Mid. Nat. 46: 1-81.

ScHULZ, O. E. 1934. Neue Cruciferen aus Südamerica. Notizbl. Bot.
Gart. Berlin, 12: 39-41.

Suaw, H. K. Airy. 1948. The Botanical Name of the Wild Tetraploid
Watercress. Kew Bull. 1947: 39-46.

1951. A Binary Name for the Hybrid Watercress.

Watsonia, 2: 73-75.

A NEW SPECIES OF SCIRPUS IN
THE NORTHEASTERN UNITED STATES

ALFRED E. SCHUYLER

While collecting leafy species of Scirpus in the Connecti-
cut River valley through Vermont, Massachusetts, and
Connecticut during the summer of 1960, I was surprised to
find a population of plants which did not conform to any
descriptions in the standard manuals for this region (Fer-
nald, 1950; Gleason, 1952). Vegetatively the plants ap-
peared very similar to Scirpus atrovirens Willd., which is
widespread in eastern North America. However, an exami-
nation of the comparatively large achenes and rigid perianth
bristles revealed some striking differences from S. atro-
virens, and later, other differences were discovered. Also,
more specimens of this taxon were found in herbarium fold-
ers containing specimens of S. atrovirens and its relative, S.
polyphyllus Vahl. Because of its well-marked distinctions
from previously recognized species of Scirpus, I herein de-
scribe it as a new species.

IT am indebted to the curators of the following herbaria where I visited or obtained
loans: Gray Herbarium, University of Massachusetts, Dartmouth College, New York
State Museum, Wiegand Herbarium, New York Botanical Garden, Pennsylvania State
University, Philadelphia Academy of Natural Sciences, U.S. National Museum, Uni-
versity of Michigan, and Missouri Botanical Garden.

44 Rhodora [Vol. 64

Scirpus ancistrochaetus sp. nov.

Culmi 0.8-1.2 m. alti, foliis inferioribus usque ad 8 mm. (10 mm.)
latis. Spiculae glomeratae; squamae spicularum vix mucronatae, illis
medianis 1.55-1.90 mm. longis. Setae 6, 1.1-1.7 mm. longae, rigidae,
fere usque ad basim dentibus retrorsis crassis acribus armatae. Fruc-
tus 1.10-1.35 mm. longi, fere omnes obovati, parte supra cavum seminis
lenta et crassa.

Plants perennial, producing flowering culms from short, woody,
underground rhizomes; lateral buds of the rhizome producing erect
shoots which break through the subtending leaf sheaths. Flowering
culms 0.8-1.2 m. tall; lower leaves up to 8 mm. (rarely 10 mm.) wide,
40-60 times as long as wide; uppermost leaf 3-5 mm. in width, 30-50
times as long as wide; lowermost involucral bract 3-5 mm. in width,
5.5-17.0 em. in length. Inflorescence rays up to 6.3 em. long, bearing
clusters of brown spikelets. Scales slightly mucronate, those from the
central portion of the spikelet 1.55-1.90 mm. long. Bristles 6, 1.1-1.7
mm. long, rigid, armed almost to the base with thick-walled, sharp-
pointed, retrorse teeth. Stamen number variable, between 0 and 3.
Styles 3-parted, up to 1.35 mm. long. Fruits yellow-brown, 1.10-1.35
mm. long, mostly obovate, the portion above the seed cavity tough and
thickened. “Oil layer" of the seed with cell walls inconspicuous as a
result of abundant pits.

TYPE LOCALITY: VERMONT: Windham Co.: 5 mi. n. of Bellows Falls,
e. side of rt. 5, small population along margin of pool, July 3, 1960,
A. E. Schuyler 3051 (MICH, type) ; Aug. 11, 1960, A. E. Schuyler 3298
(MICH); Aug. 12, 1961, A. E. Schuyler 3445 (MICH).

ADDITIONAL LOCALITIES: PENNSYLVANIA: Lackawanna Co.: 74 mi. w.
of Fleetville, east of Lake Kewanee (Windfall Pond), clump in mud-
hole on margin of wet woods, July 29, 1946, S. L. Glowenke 8052 (NYS,
PAC). [Lackawanna Co.?:] Near Carbondale, Mud Creek, Aug. 1, 1897,
D. L. Topping (us). Blair Co.: Bald Eagle Valley, 1865, J. R. Lowrie
(PAC)?

Vegetatively S. ancistrochaetus may be distinguished
from S. atrovirens by its narrower leaves, more ascending
inflorescence rays, and larger, slightly mucronate scales.
The flowers of S. ancistrochaetus have 6 perianth bristles
which are rigid and barbed almost to the base with thick-
walled, sharp-pointed, retrorse teeth; flowers of S. atro-
virens have a variable number of perianth bristles (usually
4-6) which are delicate and often wrinkled, and the thin-
walled, round-tipped, retrorse teeth are concentrated

2After submitting this manuscript, I found a specimen of S. ancistrochaetus from
New York State: Washington Co.: W. Ft. Ann, Mt. Nebo, 1800 ft., wet place, Sept. 13,
1900, S. H. Burnham (cv).

1962] Schuyler — A new Species of Scirpus 45

ine

BRRBARIOM OF THE UNIVERSITY OF MICHIGAN
FLANTS OF VERRONT

Scirpus ancistrochaetus scivayler TYPE
ES in A eel

PLATE 1266. Type specimen of Scirpus ancistrochaetus.

46 Rhodora [Vol. 64

toward the tip of the bristle. The achenes of S. ancistro-
chaetus are about 1.25 mm. long and the portion above the
locule is tough and thickened; achenes of S. atrovirens are
about 1 mm. long and lack a tough, thickened portion above
the locule. Some of these characteristics of S. ancistrochae-
tus and S. atrovirens are illustrated in figure 1.

Scirpus polyphyllus and S. ancistrochaetus have achenes
which are almost the same length, but those of S. polyphyllus
tend to be wider and less obovate. Also, S. polyphyllus may
be distinguished from S. ancistrochaetus by its more mucro-
nate scales and longer, more delicate bristles.

The bristles of S. ancistrochaetus are almost identical with
those of S. sylvaticus L. of the Old World and its North
American counterpart, S. expansus Fern. However, these
species are easily distinguished from S. ancistrochaetus by
their wider leaves, more branched inflorescences, blacker
scales, and three-angled achenes. Many of the cells outside
of the midrib in the scales of S. ancistrochaetus contain an
orange pigment while these same cells in the scales of 5.
sylvaticus and S. expansus either contain a black pigment or
lack pigment altogether. The achenes of S. ancistrochaetus
are more plano-convex, with the dorsal angle not very con-
spicuous.

Some other interesting specimens have been turned up
which appear to be robust forms of S. ancistrochaetus.
Superficially the seeds appear to be normal, but closer exami-
nation reveals that they do not contain an embryo or an
endosperm. A cytological examination of meiosis might
prove very useful in ascertaining the relationship of this
type to typical S. ancistrochaetus. It is known only from
one locality: PENNSYLVANIA: Clinton Co.: 4 mi. n. of Logan-
ton, in bog, Sept. 13, 1940, H. A. Wahl 876 (GH, CU).

Hybrids between S. ancistrochaetus and S. atrovirens are
morphologically intermediate between the two parents and
have abortive seeds. Some of their characteristics are illus-
trated in figure 1, between those of the two parents. The
criteria of assessing the hybrid nature of these plants are
quite similar to those found in the study of S. peckii (Schuy-
ler, 1961). Hybrids have been found in two localities: VER-

1962] Schuyler — A new Species of Scirpus 47

MONT: Windham Co.: 5 mi. n. of Bellows Falls, e. side of
rt. 5, growing between populations of S. ancistrochaetus and
S. atrovirens along margin of pool, Aug. 12, 1961, A. E.
Schuyler 3453 (MICH). PENNSYLVANIA: Centre Co.: 314 mi.
s. of State College, Musser’s Gap, June 13, 1914, A. R. Bech-
tel (PAC).

The distribution of S. ancistrochaetus and its hybrid with
S. atrovirens in Pennsylvania (see figure 2) appears to be
correlated with the western limits of the ridge and valley
region. Some localities (Bald Eagle Valley and the Connecti-
cut River valley) indicate that this species may occur pri-
marily in lowland habitats in hilly country. The habitats,
themselves, appear to be wetter than those of S. atrovirens.

fi d d
09 09 |

SCIRPUS ATROVIRENS HYBRID SCIRPUS ANCISTROCHAE TUS

Figure 1: 1. Bristle tips, x 47. 2. Bristle teeth, x 190. 3. Seales, x 8.5. 4. Fruits
with inclosed seeds (stipled), x 16.
In the Vermont locality, S. ancistrochaetus occurs along the
margin of a pool and S. atrovirens occurs a short distance
away from the margin. Herbarium labels on the Pennsyl-
vania specimens also suggest a wetter habitat.

Because of the floristic affinity between Asia and North
America in the genus Scirpus (Raymond, 1959), it was con-
sidered possible that S. ancistrochaetus also occurs in Asia.
However, neither Koyama (1958) nor Raymond (1957,
1959) has mentioned or described any plants closely resem-
bling S. ancistrochaetus.

The occurrence of S. ancistrochaetus in isolated localities
in a region where the flora is relatively well known suggests
that it may be a relict species. It is possible that in regions

48 Rhodora [Vol. 64

where it does not now exist, it has made a permanent imprint
on our flora through hybridization. For example, even
though the F, hybrids with S. atrovirens that I have dis-
covered appear to be highly sterile, the possibility still re-
mains that some individuals may be partially fertile and
backcross with either parent. The backcross individuals may
be extremely rare and quite sterile, but fertility may be

e S. ANCISTROCHAETUS
t ROBUST FORM
x S. ANCISTROCHAETUS X S. ATROVIRENS

FIGURE 2: Distribution of Scirpus ancistrochactus, including its robust form and its
hybrid with Scirpus atrovirens.

gradually restored as the backcross individuals acquire more
and more genes from one of the original parents, in this case
S. atrovirens. These fertile individuals which result from
backerossing would make this original parent appear more
polymorphic. I think that most taxonomists would agree

1962] Auyang — Algae of Quinsigamond 49

that S. atrovirens is morphologically variable; and Hicks
(1928) has indicated that there is also substantial cytological
variability. Some of this variability could be due to past
hybridization of S. atrovirens with S. ancistrochaetus. —
DEPARTMENT OF BOTANY, UNIVERSITY OF MICHIGAN, ANN

ARBOR.
ACKNOWLEDGMENTS
I am grateful to P. A. Hyypio and S. J. Smith for their help in
connection with my field work during the past three summers; and to
E. G. Voss and W. H. Wagner, Jr. for their help with the manuscript.
LITERATURE CITED

FERNALD, M. L. 1950. Gray's Manual of Botany. 8th ed. American
Book Co., New York.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora
of the Northeastern United States and Adjacent Canada. The
New York Botanical Garden, New York.

Hicks, G. C. 1928. Chromosome studies in the Cyperaceae with
special reference to Scirpus. Bot. Gaz. 86: 295-317.

KovAMa, T. 1958. ‘Taxonomic study of the genus Scirpus Linné.”
Jour. Fac. Sci. Tokyo Bot. 7: 2771-366.

RAYMOND, M. 1957. Some new cr critical Scirpus from Indo-China.
Nat. Canad. 84: 111-150. (Mém. Jard. Bot. Montréal 48:111-150).

. 1959. Additional notes on some S. E. Asiatic Scirpus.
Nat. Canad. 86: 225-242. (Mém. Jard. Bot. Montréal 54: 225-242).

SCHUYLER, A. E. 1961. Evidence for the hybrid origin of Scirpus
peckii. Rhodora 63: 237-243.

A SURVEY OF THE ALGAE OF
LAKE QUINSIGAMOND'

TANG SHIH-CHEN AUYANG?

This study of the fresh water algae of Lake Quinsigamond,
Worcester, Massachusetts, was undertaken in order to iden-
tify the species, to investigate their distribution, and to
compare the results with those of G. E. Stone in his paper
“Flora of Lake Quinsigamond” published in 1900.

Stone listed 81 genera and 331 species of algae, excluding
Diatoms, in his paper; the Desmids (including placoderm
and saccoderm types) were the largest group, totalling 150

1Portion of a thesis submitted to the Department of Biology, Clark University, in
partial fulfillment of requirements for the degree of Master of Arts. The thesis was
undertaken under the direction of Dr. Vernon Ahmadjian.

*Present address: Department of Natural History, Taiwan Normal University,
Taipei, Republic of China.

50 Rhodora [Vol. 64

species. The author found 86 genera and 217 species of
algae, excluding Diatoms, in which only 66 species of Des-
mids were found. This great difference in the number of
Desmid species can be explained possibly by the fact that
external environment factors have changed during the past
60 years, and have inhibited the growth of these algae.

Probably as a result of these environmental changes, the
author was not successful in collecting any species of Chara
and Nitella, of which Stone found 5 and 6 species, respec-
tively. On the other hand, tremendous quantities of Hydro-
dictyon, which was not listed in Stone’s paper, were found
by the author.

Thirty-nine genera listed by Stone were identified by the
author. However, 42 genera which Stone found were not
found by the author, while 47 genera, excluding Diatoms,
not listed by Stone were collected and identified by the
author. These figures show that the algal flora of Lake Quin-
sigamond has changed greatly due to factors which will
require more detailed investigation by future workers.

The lake was described by Stone as follows: “Lake
Quinsigamond which is situated near Worcester is one of
the largest lakes in Massachusetts, and is about six miles
long, hardly exceeding one-half mile in width, although in
many places it is from fifty to one hundred feet or more in
depth. It runs in a northerly and southerly direction along
the edge of one of the geological dividing lines of the
State which separates the central highlands from the less
elevated areas of our seaboard. The immediate surroundings
on the lake consist geologically of sand and gravel and
probably the greater portion of the basin itself is made up of
this material, over which there is spread a considerable
mass of decomposed matter, the results of centuries of
vegetable and animal decay. While a considerable portion
of the basin of the lake is too deep to give rise to much vege-
table life, there are vast areas of shallow water which are
especially adapted to a profusion of vegetable forms, thus
making it one of the best collecting grounds in Massachu-
setts."

PROCEDURE — The field work for this thesis started in

1962] Auyang — Algae of Quinsigamond 51

January, 1960. At that time, except for the northern inlet
from which water flows into the lake, the whole lake was
frozen with seven inches of ice covered with knee-deep

71°45) |
|
MAP OF LAKE L | NORTH CAUSEWAY
QUINSIGAMOND \ | 7 — BRIDGE
ue BRIDGE A
| $
NS!
|
|
pnmo Te Lr s uM EMEN
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Denim s E a
SCALE OF MILES
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52 Rhodora [Vol. 64

snow. In the beginning of April, ice on the lake gradually
started to melt from north to south. By April, 5th, all of the
ice had melted completely. While the lake was frozen, algae
were collected within the limits of the northern inlet and
samples were taken from a hole about 2 ft. in diameter dug
through the ice in the middle of the lake. During the warm-
er, ice-free months, collections were made periodically either
by walking along the lake shore or using a boat to travel to
various parts of the lake.

DISTRIBUTION — Lake Quinsigamond can easily be divided
into sections with the aid of four bridges; naming them
from north to south, they are North Causeway Bridge,
Lincoln Bridge, Causeway Bridge and Blackstone Bridge
(Fig. 1).

1). At the North Causeway Bridge, Draparnaldia was
very abundant. The water there was clear and running and
its temperature was about 1.5 C° when collections were
made. Draparnaldia was found attached to small stones
under the bridge. In the quiet water near the shore many
specimens of Tetraspora were found. Planktonic collections
in this area contained large numbers of desmids, diatoms
and several species of Pediastrum in early spring.

2). Under Lincoln Bridge there were a few big rocks on
which Ulothrix, in the vegetative stage, was collected in the
early months of spring, when the water was 4.0 C°. The
abundance of Ulothrix decreased gradually, disappearing al-
most entirely in the early summer. Collections in late spring
also contained Zygnema, in the vegetative stages, inter-
mingled with Spirogyra, Tolypothrix, Oscillatoria and
Rhizoclonium.

From late summer to fall the region between these two
bridges was covered by a mat of Hydrodictyon, Spirogyra
and other filamentous algae.

3). On the east side of Causeway Bridge, the clear water
ran slowly and contained Stigeoclonium attached to sub-
merged aquatics growing near the shore in late spring. At
the same time, about two miles south of the bridge and also
on the east shore, Vaucheria was found in fruiting stages in
the shallow, sandy bottom.

1962] Auyang — Algae of Quinsigamond 53

North and south of this bridge as far down as Blackstone
Bridge on the east bank, the red alga Batrachospermum was
found in scattered localities. This alga grew in clusters on
small stones near the shore and produced fruiting bodies in
June.

On the west shore, half a mile south of the bridge, the
water near the shore and farther away from it contained
abundant nets of Hydrodictyon. Collections from the sur-
face water as well as below contained this alga, after
the beginning of June.

4). The water north of the Blackstone Bridge and espe-
cially just beneath the bridge contained abundant specimens
of species of Polycystis, Coelosphaerium and Anabaenopsis.
A small dam south of the bridge retarded the water flow,
and probably caused the great accumulation of these forms
under it. The “water bloom” of this region was observed
in the fall.

Generally speaking, the filamentous green algae such as
Spirogyra, Mougeotia and Oedogonium were widespread and
long-lasting in occurrence. Mostly the vegetative stages
were observed and only few of these algae were in reproduc-
tive stages when collected.

Lyngbya, a filamentous blue-green alga, was collected
from May to autumn in many regions but especially at the
Lincoln Bridge, where great masses were found in the fall.

The many other algae, unicellular, colonial and multicellu-
lar which are recorded in other parts of this paper were of
widespread occurrence, either in the plankton, attached to
submerged objects, or free floating on the surface. The
vegetative stages of these algae and occasionally the repro-
ductive stages were observed in fresh collections.

ALPHABETICAL LIST OF SPECIES

Amphora ovalis Kiitzing A. faleatus var, acicularis
Anabaena limnetica G. M. Smith (A. Braun) G. S. West

A. variabilis Kützing Aphanocapsa grevillei (Hass.)
A. viguieri Denis & Frémy Rabenhorst

Anabaenopsis elenkinii Miller Aphanochaete repens A. Braun

Ankistrodesmus convolutus Corda Aphanothece clathrata G. S. West
A. faleatus (Corda) Ralfs A. microscopica Nageli

54

A. stagnina (Spreng.) A. Braun
Asterionella formosa Hassall

A. gracillima (Hantzsch) Heiberg

Batrachospermum vagum (Roth)
C. A. Agardh

Bulbochaete repanda Wittrock

B. scrobiculata (Tiff.) Tiffany

Bumilleria sicula Borzi

Caloneis silicula (Ehrenb.)

Cleve var, inflata (Grun.) Cleve

Calothrix stellaris Bornet &
Flahault

Ceratium hirundinella (O.F.M.)
Schrank

Characiopsis acuta (A. Braun)
Borzi

Chlamydomonas angulosa Dill

C. mucicola Schmidle

C. polypyrenoideum Prescott

C. snowii Printz

C. sphagnicola Fritsch & Takeda

Chlcrella ellipsoidea Gerneck

C. vulgaris Beijerinck

Chlorococcum humicola (Nàg.)
Rabenhorst

Chroococcus limneticus
Lemmermann

Closterium acerosum (Schrank)

Ehrenberg

. calosporum Wittrock

. dianae Ehrenberg

. didymotocum Ralfs

. ehrenbergii Meneghini

. incurvum de Brébisson

. lagoense Nordstedt

lanceolatum Kützing

Rhodora

maaaaaaaaaana

. leibleinii Kiitzing

. lunula (Müller) Nitzsch

. moniliferum (Bory) Ehrenberg
. turgidum Ehrenberg

. sigmcideum Lagerheim &

Nordstedt

Coelastrum microporum Nägeli
Coelosphaerium dubium Grunow

C

naegelianum Unger

Coleochaete orbicularis

Pringsheim

[Vol. 64

C. scutata de Brébisson

C. soluta (de Bréb.) Pringsheim

Cosmarium biretum de Brébisson
var. trigibberum Nordstedt

C. broomei Thwaites

C. botrytis Meneghini

C. dentiferum Corda

C. formosulum Hoffman

C. formosulum var. nathorstii

(Boldt) W. & G. S. West

C. granatum de Brébisson

C. margaritatum Roy & Biss.

C. monomazum Lundell

C. nitidulum DeNotaris

C. orbiculatum Ralfs

C. panamense Prese

C. protractum (Nàg.) DeBary

C. pseudoconnatum Nordstedt

C. punctulatum de Brébisson

C. quadrum Lundell

C. reinforme (Ralfs) Archer

Cosmocladium hitcheockii (Wolle)

G. M. Smith

Crucigenia truncata G. M. Smith

Cyclotella bodanica Eulenstein

C. bodanica var. stellata
Skvortzow

C. comta (Ehrenb.) Kützing

Cymbella aspera (Ehrenb.) Cleve

C. tumida (de Bréb.) Van Heurck

Desmidium swartzii C. A. Agardh

Diatoma hiemale (Lyngbye)
Heiberg var. mesodon
(Ehrenb.) Grunow

D. vulgare Bory

Dinobryon sertularia Ehrenberg

Dictyosphaerium pulchellum Wood

Draparnaldia glomerata (Vauch.)
C. A. Agardh

D. judayi Prescott

Epithemia turgida (Ehrenb.)
Kützing

Euastrum elegans (de Bréb.)
Kützing

E. pulchellum de Brébisson

E. verrucosum (Ehrenb.) Ralfs

Euglena gracilis Klebs

1962]

E. spirogyra Ehrenberg

Fragilaria capucina Desmazieres

F. construens (Ehrenb.) Grunow

F. crotonensis Kitton

F. crotonensis var. prolongata
Grunow

Gloeocystis ampla (Kiitz.)
Lagerheim

G. gigas (Kütz.) Lagerheim

G. major Gerneck ex
Lemmermann

G. vesiculosa Nägeli

Golenkinia paucispina W. &
G. S. West

Gomphoneis herculeana (Ehrenb.)
Cleve var. robusta (Grun.)
Cleve

Gomphonema acuminatum
Ehrenberg

G. constrictum Ehrenberg

Gonatozygon kinahani (Arch.)
Rabenhorst

Gymnodinium fuscum (Ehrenb.)
Stein

G. palustre Schilling

Hormidiopsis ellipsoideum
Prescott

Hyalotheca dissiliens (J. E.
Smith) de Brébisson

Hydrodictyon reticulatum (L.)
Lagerheim

Kirchneriella lunaris (Kirch.)
Moebius var. irregularis
G. M. Smith

Leptosira mediciana Borzi

Licmophora gracilis (Ehrenb.)
Grunow

L. paradoxa (Lyngbye)
C. A. Agardh

Lyngbya epiphytica Hieronymus

T, latissima Prescott

L. majuscula (Dill.) Harv.

L. wollei Farlow

Merismopedia punctata Meyen

Melosira italica (Ehrenb.)
Kützing

M. juergensii C. A. Agardh

Auyang — Algae of Quinsigamond

55

Micrasterias americana (Ehrenb.)
Ralfs

M. apiculata (Ehrenb.)
Meneghini

M. apiculata var. fimbriata
(Ralfs) Nordstedt f. spinosa
(Bissett) W. & G. S. West

M. radiata Hassall

M. rotata (Grev.) Ralfs

M. truncata (Corda) de Brébisson

Microspora willeana Lagerheim

Microthamnion kuetzingianum
Nägeli

Mougeotia capucina (Bory)
C. A. Agardh

M. floridana Transeau

M. genuflexa (Dillw.)
C. A. Agardh

M. laetevirens (A. Braun)
Wittrock

M. parvula Hassall

M. reinschii Transeau

M. robusta (de Bary) Wittrock

M. scalaris Hassall

Mougeotiopsis calospora Palla

Navicula anglica Ralfs

Netrium digitus (Ehrenb.)
Itsigsohn & Roth

Nostoc muscorum C. A. Agardh

Oedogonium borisianum (Le Cl.)
Wittrock

O. capillare (L.) Kützing

O. crenulatocostatum Wittrock

O. grande Kützing var.
aequatoriale Wittrock

O. laeve Wittrock

O. plusiosporum Wittrock

O. pringsheimii Cramer

Oócystis pusilla Hansgirg

Oscillatoria cortiana Meneghini

O. curviceps C. A, Agardh

O. ornata Kützing

O. prolifica (Grev.) Gomont

O. splendida Greville

O. tenuis C. A. Agardh

Pandorina morum (Muell.) Bory

Pediastrum biradiatum Meyen

56

P. boryanum (Turp.) Meneghini

P. boryanum var. longicorne
Raciborski

P. duplex Meyen

P. duplex var. clathratum
(A. Braun) Lagerheim

P. duplex var. rotundatum Lucks

P. duplex var. rugulosum
Raciborski

P. ehrenbergii A. Braun

P. tetras (Ehrenb.) Ralfs

P. tetras var. tetraodon (Corda)
Rabenhorst

Penium margaritaceum (Ehrenb.)

de Brébisson
P. digitus (Ehrenb.) Itsigsohn &
Roth
Peridinium cinctum (Müll.)
Ehrenberg
Phacus longicauda (Ehrenb.)
Dujardin
Phormidium nareanum Grunow
Pinnularia nobilis Ehrenberg
Planktosphaeria gelatinosa
G. M. Smith
Pleurotaenium coronatum (de
Bréb.) Rabenhorst
P. maximum (Reisch) Lund
P. trabecula (Ehrenb.) Nàgeli
Polycystis aeruginosa Kützing
P. incerta Lemm.
Pleurococcus vulgaris Nägeli
Protoderma viride Kützing
tadiofilum flavescens G. S. West
Rhizochrysis limnetica
G. M. Smith
Rhizoclonium hieroglyphicum
(C. A. Ag.) Kützing
R. hieroglyphicum var. hosfordii
(Wolle) Collins
R. hookeri Kützing
Scenedesmus abundans (Kirch.)
Chodat
S. abundans var. brevicauda
G. M. Smith
S. acutiformis Schroeder

Rhodora

S

S

U UNUNUN UN N

S
S

S

S

S
S

NNNMNN

S.

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S
S

UN N N

[Vol. 64

. arcuatus Lemmermann var.
platydisca G. M. Smith

. armatus (Chod.) G. M. Smith
var. major G. M. Smith

. bijuga (Turp.) Lagerheim

. brasiliensis Bohlin

. denticulatus Lagerheim

. dimorphus (Turp.) Kützing

. longus Meyen

. longus var. minutus
G. M. Smith

. obliquus (Turp.) Kützing

. quadricauda (Turp.) de
Brébisson

. quadricauda var. longispina
(Chod.) G. M. Smith

orastrum americanum (Bohlin)
Schmidle

phaeroceptis schroeteri Chodat

pirogyra cleveana Transeau

. ellipsospora Transeau

. hydrodictya Transeau

. jugalis (Fl. Dan) Kiitzing

. mirabilis (Hass.) Kützing

. nitida (Dillw.) Link

submaxima Transeau

pirotaenia condensata Brébisson

pirulina duplex Wolle

pondylosium pulchellum Arch.

taurastrum alternans de
Brébisson

. avieula de Brébisson

. brevispinum de Brébisson var.
tumidum G. M. Smith

. cerastes Lund

. crenulatum (Delp.) Nägeli

. dickiei Ralfs var. maximum
W. & G. S. West

. dilatatum Ehrenberg

. furcigerum de Brébisson

. gracile Ralfs

. grande Bulnheim

. leptocladum var. denticulatum
G. M. Smith

. odontatum Wolle

. orbieulare (Ehrenb.) Ralfs

. paradoxum Meyen

1962]

S. subgrande Borge var. minus
G. M. Smith

Stauroneis acuta W. Smith

Stigeoclonium flagelliferum
Kützing

S. nanum Kützing

Surirella elegans Ehrenberg

S. splendida (Ehrenb.) Kiitzing

Synechocystis aquatilis
Sauvageau

Synedra ulna (Nitzsch)
Ehrenberg

S. ulna var. aequalis (Kütz.)
Hustedt

Synura uvella Ehrenberg

Tabellaria fenestrata (Lyngb.)
Kützing

T. floceulosa (Roth) Kützing

Tetraédron limneticum Borge

Tetraspora cylindrica (Wahl.)
C. A. Agardh

Auyang — Algae of Quinsigamond

57

T. lubrica (Roth) C. A. Agardh
.etrastrum staurogeniaeforme
(Schroeder) Lemmermann
Volypothrix conglutinata Borzi
T. distorta Kützing
lanata Wartmann
Trachelomonas crebea (Kellicott)
Deflandre
Tribonema bombycinum (C. A.
Ag.) Derbés & Solier
Ulothrix tenerrima Kützing
U. zonata (Weber & Mohr)
Kützing
Vaucheria ornithocephala
C. A. Agardh
V. sessilis (Vauch.) DeCandolle
Westella linearis G. M. Smith
Xanthidium antilopaeum (de
Bréb.) Kützing var. polymazum
Nordstedt
Zygnema insigne (Hass.) Kützing

The following is an alphabetical list which shows the genera found
by both Stone and the author, and those found only by Stone or by the

author.

Stone and author Stone Author
Anabaena Arthrodesmus Anabaenopsis
Aphanochaete Bambusina Ankistrodesmus
Batrachospermum Botrydium Aphanocapsa
Bulbochaete Calocylindrus Aphanothece
Calothrix Chaetophora Bumbilleria
Closterium Chara Ceratium
Coelastrum Characium Characiopsis
Coelosphaerium Chlamydococcus Chlamydomonas
Coleochaete Chroolepus Chlorella
Cosmarium Cladophora Chlorococcum
Desmidium Clathrocystis Chroococcus
Dictyosphaerium Conferva Cosmocladium
Draparnaldia Craterospermum Crucigenia
Euastrum Cylindrocapsa Dinobryon
Euglena Cylindrospermum Golenkinia
Gloeocystis Dimorphococcus Gonatozygon
Hyalotheca Docidium Gymnodinium
Lyngbya Eremosphaera Hormidiopsis
Micrasterias Gloeotrichia Hydrodictyon
Microthamnion Gonium Kirchneriella

58 Rhodora [Vol. 64
Nostoc Hapalosiphon Leptosira
Oedogonium Hydrurus Merismopedia
Oscillatoria Isactis Microspora
Pandorina Leptothrix Mougeotia
Pediastrum Mesocarpus Mougeotiopsis
Penium Mesotaenium Netrium
Pleurococcus Nephrocytium Oócystis
(Protococcus) Nitella Peridinium
Rhizoclonium Ophiocytium Phacus
Scenedesmus Palmella Phormidium
Sorastrum Pleurocarpus Planktosphaeria
Spirogyra Polyedrium Pleurotaenium
Stigeoclonium Porphyridium Protoderma
Staurastrum Raphidium Polycystis
Tetraspora Rivularia Radiofilum
Tolypothrix Schizochlamys Rhizochrysis
Ulothrix Seytonema Sphaerocystis
Vaucheria Sirosiphon Spirotaenia
Xanthidium Sphaerozosma Spirulina
Zygnema Stauraspermum Spondylosium
Tetmemorus Synechocystis
Volvox Synura

Tetraédron
Tetrastrum
Trachelomonas
Tribonema
Westella

Diatoms excluded from above lists.

DISCUSSION — It is possible that some of the differences in
the algae collected by Stone and the author may be due to
the fact that Stone listed algal names now regarded as
synonymous. This may be one explanation as to the great
difference in the number of species listed by the author.
However, the difference in the number of Desmid species,
especially, cannot be due solely to differences in nomencla-
ture. The reason why the number of species, especially
Desmids, decreased between 1900-1960 could be because the
environmental conditions in Lake Quinsigamond have
changed.

The chief environmental factors are light, temperature,
chemical composition and pH of water. The first two factors
were not recorded in Stone's paper. Therefore, there is no
way to compare and discover what changes have occurred

1962] Auyang — Algae of Quinsigamond 59

in these two factors from 1900 to 1960. However, there is
at least one external environmental factor which has been
introduced on the lake during this 60-year period, and that
is the presence of motor boats. Outboard motor boats de-
posit a film of oil on the surface of the lake. This oil is then
moved, by the motion of waves, to the shores or coves. These
coves are the best habitats for the planktonic Desmids. If
the water is covered with an oily film, atmospheric oxygen
cannot dissolve into the water and as a result the respiration
of Desmids will eventually cease. This unfavorable environ-
ment probably inhibited growth of certain Desmids.

Five species of Chara and six species of Nitella were found
by Stone. The author was not successful in collecting any
species of these genera. One possible reason is that “Chara
thrive best in clear, hard water (Smith 1950)." The Health
Department of Worcester stated that in 1945 the water in
Lake Quinsigamond was quite soft, the hardness being 23.
It may be that the water today exhibits an even lower degree
of hardness due to factory pollutions (personal communica-
tion, Health Department, Worcester, Mass.). However, this
fact has not been substantiated either by the Worcester
Health Department or by the author. Another possible
reason is that there are many ducks on the lake and they
feed upon these algae.

The range of pH values of water samples in Lake Quinsig-
amond was 6.5-7.8. However, there was one exception,
namely, the water sample taken from the west shore, one
half mile south of Causeway Bridge. This water in which
tremendous quantities of Hydrodictyon were found has a
pH of 9.0 in June. It is interesting to note that this finding
supports the following statement: 'Hgydrodictyon may be
used as an index organism for a high pH (Prescott, 1951).”

LITERATURE CITED

PRESCOTT, G. W. 1951. Algae of the Western Great Lakes Area.
Cranbrook Inst. of Sci., Bull. No. 31, pp. 1-946.

SMiTH, G. M. 1950. Freshwater Algae of the United States. Mc-
Graw-Hill, N. Y.

STONE, G. E. 1900. Flora of Lake Quinsigamond. F. S. Blanchard
and Co., Printers, Worcester, Mass. pp. 1-12.

60 Rhodora [Vol. 64

NOMENCLATURAL NOTES ON
TWO SOUTHEASTERN RHYNCHOSIAS

ROBERT L. WILBUR!

In 1834 Nuttall described what he considered to be a new
genus of legumes from Florida naming its only species
Pitcheria galactoides. Almost all authors since, including
Bentham (Gen. Pl. 1: 543. 1865) and Taubert (in Engler
and Prantl’s Nat. Pfl. III. 3: 373. 1894), have treated it as a
species of Rhynchosia. Burkart (Darwiniana 11: 268, 269.
1957) also concluded that it did not merit generic rank and
that it had close affinities with species in the Antilles and
South America. He noted that the name most commonly
employed under Rhynchosia was illegitimate, being a later
homonym of R. galactioides (H.B.K.) DC. This later homo-
nym is, of course, illegitimate even though the prior species
with the earlier name has long been considered a member of
the genus Calopogonium. Burkart therefore provided a new
name, R. pitcheria.

However, Bertoloni in an often overlooked paper named
this species Lespedeza cytisoides more than one hundred
years earlier. In spite of Bertoloni's unfortunate decision as
to the generic placement of this as well as many other species
described by him from the Southeast, there can be no doubt
as to the identity of the species as it was fully described and
accompanied by a full-page plate. Asa Gray in his review
(Am. Jour. Sci. II. 14: 114-115. 1852) of Bertoloni's contri-
bution unhesitatingly assigned the name to the synonymy
of Pitcheria galactoides. Therefore the following new combi-
nation is required for this Gulf Coast endemic:

Rhynchosia cytisoides (Bert.) comb. nov.
Pitcheria galactoides Nutt., Jour. Acad. Phil. I. 7: 94. 1834.
Rhynchosia galactioides (Nutt.) Endl. ex Walp., Repert. Bot. Sys-
tem. 1: 790. 1842 but not R. galactioides (H.B.K.) DC., Prodr. 2: 387.
1825.
Lespedeza cytisoides Bert., Mem. Acad. Sci. Bologna 2: 278. 1850.
Rhynchosia pitcheria Burkart, Darwiniana 11: 268. 1957.

1Grateful acknowledgment is made to the National Science Foundation for a grant
of research funds to Duke University (NSF-Grant 5636) which made the present study
possible.

1962] Wilbur — Southeastern Rhynchosias 61

The binomials most recently employed for the erect, uni-
foliolate Rhynchosia of the southeastern Coastal Plain have
been either R. simplicifolia (Walt.) Wood or R. reniformis
(Pursh) DC. Since neither of these names is fully in accord
with the current International Code, a brief discussion seems
desirable.

The most widely used name, R. simplicifolia (Walt.)
Wood, is a later homonym of R. simplicifolia (H.B.K.) DC.,
and hence is not legitimate (Art. 64.2).

Recently Turner (The Legumes of Texas, p. 255. 1959)
has taken up R. reniformis (Pursh) DC. for this species.
However Pursh published this as Glycine reniformis with
Trifolium simplicifolium Walt. listed in synonymy. There
was no presently acceptable reason why Pursh should not
then have taken up Walter’s epithet as Glycine simplicifolia
H.B.K. was not published until 1823 or nine years after
Pursh’s publication. Hence G. reniformis Pursh is an illeg-
itimate name as it was superfluous when published (Art.
64.1).

The desirability of Article 64.1 has been recently chal-
lenged by both Furtado (Taxon 9: 147-150. 1960) and Holt-
tum (Taxon 10: 33-34. 1961). Furtado’s long campaign
against this portion of the Code has now entered its third
decade having been rejected most recently by the Montreal
Botanical Congress. Holttum argues that “a number of well-
known and long used names, which under Art. 64.1 have
been regarded as unusable, could be used" if this portion
of the Code were eliminated and botanists relied instead
upon typification and priority to rule unaided. This is no
doubt true but it is certain also that the elimination of this
provision would change the names of a very large number
of plants to which botanists have become accustomed in the
more than thirty years that the present provision has been
in effect. As Fosberg points out (Taxon 8: 65. 1959) Art.
64.1 is indeed “a two-edged sword." Without wishing to
debate the original merits of our Art. 64.1, I believe it now
would serve stability better to retain the primary strictures
of this provision.

Fortunately however, for the sake of at least some stabili-

62 Rhodora [Vol. 64

ty, Article 72 permits the retention of one of the binomials
by which this unifoliolate Rhynchosia has been known in the
past. There was no epithet available in 1825 that could have
been transferred by DeCandolle into Rhynchosia but the
binomial R. reniformis then published by him need be no
more considered as a new combination than are Desmodium
rotundifolium or D. lineatum. All three of these names were
first published by DeCandolle who cited as synonyms names
which were later homonyms. All are or should be credited
solely to DeCandolle with no parenthetical authority. The
synonymy of the species is listed below.

Rhynchosia reniformis DC.

Trifolium simplicifolium Walt., Fl. Car. 184. 1788.

Glycine tomentosa var. ó monophylla Michx., Fl. Bor. Am. 2: 63. 1803,

Glycine reniformis Pursh, Fl. Am. Sept. 486. 1814. nom. illegit. (Art.
64.1)

Arcyphyllum simplicifolium (Walt.) Ell, Jour. Acad. Phil. I. 1:
371. 1818.

Glycine monophylla (Michx.) Nutt., Gen. N. Am. Pl. 2: 115. 1818,
not Linnaeus, Mant. 1: 101. 1767.

Glycine simplicifolia (Walt.) Ell., Sk. Bot. S. C. & Ga. 2: 234. 1823,
not G. simplicifolia H.B.K., Nov. Gen. et Sp. 6: 419. 1823.

Rhynchosia reniformis DC., Prodr. 2: 384. 1825.

Rhynchosia tomentosa var monophylla (Michx.) T. & G., Fl. N. Am.
1: 284. 1838.

Phaseolus reniformis (Pursh) Eaton & Wright, N. Am. Bot. 353.
1840.

Psoralea alnifolia Bert., Mem. Acad. Sci. Bologna 2: 274. 1850.

Rhynchosia simplicifolia (Walt.) Wood, Class-Book 321. 1861, not
DC., Prodr. 2: 389. 1825.

Dolicholus simplicifolius (Walt.) Vail, Bull. Torrey Club 26: 114.
1899. — DEPARTMENT OF BOTANY, DUKE UNIVERSITY.

1962] Cooperrider — Enchanter’s Nightshade 63

THE OCCURRENCE AND HYBRID NATURE OF AN
ENCHANTER’S NIGHTSHADE IN OHIO

TOM S. COOPERRIDER

Floristic work by the writer! and one of his students in
northeastern Ohio during the summer of 1960 yielded two
collections of Circaea canadensis Hill, making it possible to
add this species with certainty to the known state flora.
Schaffner (1932) noted that this species had been reported
for the state but that he had seen no specimens. The state
is not cited as part of the species range in either of the two
regional manuals (Fernald, 1950; Gleason, 1952), but its
known occurrences in neighboring states of Pennsylvania,
West Virginia, and Michigan make its presence here not un-
expected. Examination of the Circaea specimens in the Her-
barium of The Ohio State University’ disclosed one earlier
collection: Ash Cave, Hocking County, E. A. Albaugh & Len.
Stephenson, Summer, 1930. This is a fragmentary specimen
originally identified as C. lutetiana L., but is undoubtedly C.
canadensis. This station, located in southeastern Ohio, has
not yet been visited by the writer.

The two newly discovered stations are in the extreme
northeastern part of the state. The original collections from
both have been added to the Herbarium of Kent State Uni-
versity. The collection data are:

(1) Open wooded slope; east side of Brewster Road and north of
Chagrin River where the two cross; Bainbridge Township, Geauga
County. W. D. Hawver 1123, July 25, 1960. (2) Moist, rocky, wooded
valley of Mill Creek at “Cassey’s Springs” area, 2 miles NNW of
Mesopotamia, Mesopotamia Twp., Trumbull County. Tom S. Cooper-
rider 7924, Aug. 11, 1960.

The nature of C. canadensis has been the subject of past
conjecture, Gleason (op. cit.) noting that it is often consid-
ered a hybrid between the other two species Circaea in this
area: C. alpina L. and C. quadrisulcata (Maxim.) Franch.

1This field work was supported by a Kent State University Summer Faculty Research
Appointment.

?Appreciation is expressed to Dr. Clara Weishaupt, curator of the herbarium, for
the loan of these specimens.

64 Rhodora [Vol. 64

& Sav. Fernald (1917), however, decided that the “range
of the three species does not coincide and there is little to
suggest hybrid characters”. Field examination of the two
newly discovered Ohio stations for C. canadensis, and meas-
urements in the herbarium on a sample of ten or more
specimens of this taxon from each location suggest that a
re-evaluation of Fernald’s arguments is appropriate.

Both stations in northeastern Ohio were studied in the
summer of 1961. They are remarkably similar. Each is
centered in a deep ravine of the type known locally as a
“hemlock gorge". In each ravine, all three taxa of Circaea
are growing. At the head of both ravines, a small stream
falls over a sandstone cliff about 20 feet in height. C. alpina
grows on or near the cliff in a moist, north-facing situation,
usually on dripping, mossy ledges. Colonies of C. canadensis
are in the rocky, wooded valley a few hundred feet down-
stream from the cliffs. In both valleys, some plants of C.
quadrisulcata grow within a few feet of the C. canadensis
plants. C. quadrisulcata is more abundant, however, in the
dry, upland woods bordering the ravines. The proximity of
the three taxa at these two stations would negate any argu-
ment against hybridization based on spatial separation.

That C. canadensis is generally intermediate in morpho-
logy between the other two species is apparent from casual
observation (Plate 1267, Fig. 1) More specifically, its inter-
mediacy with regard to:

stature, length of corolla lobe,
length of leaf, length of anther,

length of leading raceme, length of fruit pedicel, and
width of calyx lobe, thickness of fruit

has been noted previously (Fernald, 1950; Gleason, 1952).
Cursory measurements on Ohio specimens indicate that it
is also intermediate, both in range of variation and in mean,
with respect to:

number of internodes above roots, length of disk above hypanthium,

position of longest internode, length of calyx lobe,
length of longest internode, w dth of corolla lobe,
width of leaf blade, length of filament, and

length of flower bud, size of pollen grain.

1962] Cooperrider — Enchanter’s Nightshade 65

Stebbins (1950, p. 26) has reasoned on a genetic basis that
“the larger the number of characters in which a particular
plant can be found to be intermediate betweeen its putative
parents, the greater is the probability of its hybrid origin".
The large number of morphologically intermediate charac-

PLATE 1267

Fic. 1. Habit of representative plants. Left, Circaea quadrisulcata; center, C,
canadensis; right, C. alpina (al X 1/6). Fic. 2. Fruiting racemes collected from
Geauga Co., Sept. 9, 1961. Left, C. alpina; center, C. canadensis; right, C. quadrisul-
cata (al X !4). Fic. 3. C. quadrisulcata, six pollen grains that stained with aniline
blve (X 100). Fic. 4. C. canadensis, three pollen grains that did not stain with
aniline blue (X 100). Fic. 5. C. alpina, three pollen grains that stained with aniline
blue, one that did not (X 100).

66 Rhodora [Vol. 64

ters in this case supports the notion of hybrid origin of C.
canadensis.

Failure of fruit to set on C. canadensis is relevant. This
feature is noted by Gleason (op. cit.), but not by Fernald
(1917, 1950). Careful searching of the plants of C. cana-
densis in northeastern Ohio disclosed none with even a
single fruit set in early September, 1961. Nearby plants of
the other two species had set fruit abundantly (Fig. 2).
Plants of all three taxa moved into garden culture in Spring
of 1961 showed a similar pattern, both C. alpina and C.
quadrisulcata set fruit abundantly ; plants of C. canadensis,
observed until killed by frost, set no fruit.

Fresh pollen of plants from each of the two stations in
northeastern Ohio was stained with aniline blue in lacto-
phenol (Fig. 3-5), a technique described by Lawrence (1951,
p. 71) and Long (1959) as a test for fertility and, hence,
possible hybridity. Pollen taken from two to six plants of
each taxon from these two stations gave the following re-
sults:

1. For C. alpina, of 747 grains counted, 592 (79%) stained dark
blue and 155 (21%) took no blue stain.

2. For C. quadvrisulcata, of 1000 grains counted, 791 (79%) stained
dark blue and 209 (21%) took no blue stain.

3. For C. canadensis, of 2036 grains counted, 11 (0.5%) stained
dark blue, 173 (8.5%) showed a dark blue spot on the body of the
grain, and 1852 (91%) took no blue stain.

Assuming only those grains which stained completely to be
“good”, this count reveals that while the amount of good
pollen produced by C. alpina and C. quadrisulcata is 79%,
that of nearby plants of C. canadensis is less than 1%. A
second generalization by Stebbins (1950, p. 27) on the
nature of interspecific hybrids is pertinent. “If the plant is
completely healthy and growing in company with its puta-
tive parents, then the discovery of a high-proportion of
well-filled pollen grains in the latter and 50% or more of
empty ones in the suspected plant is strong evidence that
this plant is a hybrid between the two forms that are partly
isolated from each other genetically."

1962] Cooperrider — Enchanter’s Nightshade 67

On the basis of spatial proximity, morphological inter-
mediacy, and sterility, as indicated by failure of fruit to
set and production of only a small percentage of good pollen,
it seems reasonable to conclude that these plants of C. cana-
densis in northeastern Ohio are indeed interspecific hybrids
of C. alpina x C. quadrisulcata.

In connection with the planning of further study of the
matter, it is noteworthy that a similar, or perhaps the same,
situation exists in Europe in C. intermedia Ehrh., which
Fernald (1917) considered conspecific with C. canadensis.
C. intermedia has been variously considered as a Form of
C. alpina, a Form of C. lutetiana, an interspecific hybrid
between the two, and as a distinct species (Hegi, 1908;
Gagnepain, 1916). — DEPARTMENT OF BIOLOGICAL SCIENCES,
KENT STATE UNIVERSITY, KENT, OHIO.

LITERATURE CITED

FERNALD, M. L. 1917. The identity of Circaea canadensis and C.
intermedia. Rhodora 19:85-88.

. 1950. Gray's Manual of Botany. 8th ed. American
Book Co., New York.

GAGNEPAIN, F. 1916. Revision du genre Circaea. Bull. Soc. Bot.
France 63:39-43.

GLEASON, H. A. 1952. New Britton and Brown Illustrated Flora of
Northeastern United States and Adjacent Canada. Lancaster
Press, Lancaster, Pa. vol. 2.

HEGI, G. 1908. Illustrierte Flora von Mittel-Europa. vol. 5, part II.

LAWRENCE, G. H. M. 1951. Taxonomy of Vascular Plants. Macmillan
Co., New York.

Long, R. W. 1959. Natural and artificial hybrids of Helianthus Maxi-
miliani X H. grosseserratus. Amer. Jour. Bot. 46:687-692.

SCHAFFNER, J. H. 1932. Revised catalog of Ohio vascular plants.
Ohio Biol. Survey, Bull. 25. Ohio State University, Columbus.

STEBBINS, G. L., JR. 1950. "Variation and evolution in plants. Colum-
bia Univ. Press, New York.

68 Rhodora [Vol. 64

THE TAXONOMY OF SETCREASEA OVATA
SHAMIM A. FARUGI, R. P. CELARIER' AND K. L. MEHRA

Setereasea ovata is a variable taxon which has frequent-
ly been confused with S. brevifolia (Torr.) Sch. et Sydow.
Johnston (1944) in an attempt to clarify the differences
between S. brevifolia and S. buckleyi Jtn. mentioned speci-
mens from the Chisos and Davis Mts. of Texas as S. brevi-
folia, but actually referred them to S. ovata. On the other
hand, Rose (1899) included S. ovata as a synonym of S.
brevifolia. This confusion is primarily due to the similarity
of leaf shape of these two species. Also, morphological stud-
ies have shown that S. ovata is very similar to S. pallida
Rose in its floral characters.

S. pallida differs from S. brevifolia in possessing oblong-
elliptic leaves, broad membranous connectives and glabrous
ovaries, whereas S. brevifolia possesses elliptic-ovate leaves,
narrow and thick connectives and pubescent ovaries. For
a clear-cut differentiation of the two species, the following

descriptive material should be helpful.
S. pallida: stem more or less 40 cm. tall; leaves oblong-elliptic;
sepals 3; petals 3; stamens 6, epipetalous, connectives broad and

membranous; ovary glabrous.

S. brevifolia: stem more or less 40 cm. tall; leaves ovate-elliptic;
sepals 3; petals 3; stamens 6, epipetalous, connectives narrow and
thick; ovary pubescent.

S. ovata was originally described by Coulter (1890) as
Tradescantia leiandra var. ovata. S. brevifolia was also des-
cribed as a variety of T. leiandra by Torrey (1859). Rose
(1899) gave generic rank to this group of plants, charac-
terized by having connate corollas and epipetalous stamens,
under a new genus Treleasea. Treleasea being the name
of an already described fungus, Schumann and Sydow
(1899), changed it to Setcreasea. The rank of various spe-
cies described by Rose in the treatment of Schumann and
Sydow remained the same, and, as mentioned before, Rose
had included var. ovata as a synonym under S. brevifolia.

The original description of Torrey (1859) for var. brevi-

IDeceased December 23, 1959.

1962] Faruqi, Celarier and Mehra — Setcreasea ovata 69

folia indicates a pubescent ovary. The colored plate
included by Rose in his publication shows the same. These
facts and others lead us to believe that both Rose and Johns-
ton did not study the internal structures of S. ovata and pre-
sumed it to be S. brevifolia because of the close resemblance
of the leaves in each case.

All of the taxa of Setcreasea that were studied experi-
mentally in our laboratory show a vegetative mode of
reproduction and a high degree of self-incompatibility. How-
ever, inter-specific hybridization is common. Even triploid
hybrids are quite vigorous and are vegetatively normal.
Therefore, the designation of specific rank to a new taxon
should be made only with caution. However, the species
S. ovata was found to be an exception to the generic tenden-
cy toward easy interspecific hybridization. We could not
produce any hybrids involving either S. brevifolia or S.
pallida. A single hybrid was produced by using S. purpurea
as a female parent. The details of the crossing data are
given in Table 1.

S. jawmavensis Matuda according to the type descrip-
tion is another species that comes very close to S. ovata.

TABLEI
Attempted Cross Pollina- Swollen Seed Seedlings Plant
tions ovary Set
S. ovata X S. pallida 17 1 2 none none
S. ovata X S. purpurea 11 9 32 none none
S. pallida X S. ovata 65 48 6 none none
S. purpurea, X S. ovata 22 11 29 1 1
S. brevifolia X S. ovata 12 6 ii none none

2n 28

These species are similar in their floral structures. The
type specimen of S. jaumavensis is a fragment of Rozhyn-
sky’s collection number 260, dated December 14, 1931, and is
supposed to be in the herbarium of the New York Botani-
cal Garden. This specimen has been misplaced, consequent-
ly it is at present unavailable. We came across another
fragment of Rozhynsky’s collection 260 from the herbarium
of the Chicago Natural History Museum determined as
Tradescantia semisomna. A very careful study of floral

70 Rhodora [Vol. 64

parts showed a fused corolla and epipetalous stamens, the
criteria used to separate the genus Setcreasea from Trade-
scantia, This plant is found to be similar to S. ovata both
in floral and vegetative structures, consequently a doubt
arose regarding the validity of S. ovata as a separate
species. On the basis of the type description, the leaves of
S. jaumavensis are broadly lanceolate and pubescent. In
Rozhynsky 260 from the Chicago Natural History Museum
the leaves are ovate and glabrous. The difference in leaf
shape should not be too seriously considered due to the fact
that in this specimen there are only two upper leaves pres-
ent which may not represent a normal mature leaf. But
the absence of leaf pubescence provides a very good reason
to believe that Rozhynsky 260 from the Chicago Natural
History Museum is not S. jawmavensis, rather it appears
to be part of a mixed collection. Dr. Matuda (personal
communication) is in agreement with the authors’ opinion
that S. ovata is distinctly different from S. jawmavensis,
specially in leaf pubescence.

These studies point out that S. ovata is a distinct taxon
which is not easily confused with any of the three species, i.e.
S. pallida, S. brevifolia or S. jawmavensis. It certainly
deserves specific rank.

S. ovata (Coulter) Faruqi, Celarier and Mehra, stat. nov.

Based on Tradescantia leiandra var. ovata Coulter, Contr.
U. S. Nat. Herb. 1: 50, 1890.

Plant perennial with underground rhizome; aerial stem usually
erect, great variation in the height of the plant, 20-30 em. or more;
leaves elliptic or ovate; inflorescence terminal subtended by two leafy
bracts; pedicel pubescent; calyx 3; corolla 3, pink; stamens 6,
epipetalous, connectives broad membranous; ovary glabrous.

TYPE: Texas. Neally 1889 (US).

TEXAS: Chisos Mts. Brewster Co. Hinckley 7-31-1940 (GH) ; Hinckley
1113 (F); Havard 78 (GH, US) ; Warnock T 574 C (GH, US, F); Warnock
643 (US); Sperry 1476 (GH, US); Moore & Steyermark 3228 (GH, MO,
F); Mueller 7854 (F, US, MO, MICH); Young 122 (MO); McDougall 33
(GH) ; Birdmine Hill, East of Alpine, Warnock 21589 (MO, F) ; Railroad
at Rio Grande, Eggert 1737023 (MO); Devils River, Clark 4138 (MO).

MEXICO: Coahuila: North of Cañon de Cienegas, White 1899 (MICH);
Villa Acuna, Wynd and Mueller 624 (GH, MO); Muzquiz, Wynd and
Mueller 335, (GH, US); Soledad, SW of Monclova, Palmer 2014 (GH);

1962] Faruqi, Celarier and Mehra — Setcreasea ovata 71

Sierra de la Encantada, Rancho Buena Vista, Stewart 1438 (GH);
Sierra del Pino, Head of Cañon Ybarra, Stewart 1257 (GH); Sierra de
las Cruces, Stewart 1109 (GH); Cuatro Cienegas, Marsh 2049 (GH).
Nuevo León: Sabinas Hidalgo, Leavenworth 43 (F).

Morphologically this species is very closely related to S.
pallida. The production of a hybrid with S. purpurea also
points out a relationship. The indecisiveness of negative
results from crossing as stated by Rollins (1953) for gen-
era is valuable as well for the species relationship. The
production of a hybrid indicates its relationship with S.
purpurea, while it does not negate a potential relationship
to S. pallida. We may point out that there also exists a close
relationship between S. pallida and S. purpurea. Both S.
ovata and S. purpurea used as parents are tetraploids with
2n = 24 chromosomes, while S. pallida is a diploid. From
the crosses involving this tetraploid, S. purpurea, and S.
pallida it is also concluded (unpublished) that S. pallida has
contributed one genome to the tetraploid S. purpurea. S.
purpurea. has also been reported in diploid condition (Bose,
1959). The relationships between these three morpho-
logically closely related species cannot be fully understood
without more experimental evidence especially involving
diploid S. purpurea. Superficially S. ovata looks very much

tnt

Fic. 1, Leaf shape in S. pallida; Fic. 2-4. Leaf shape variation in S. ovata; Fic.
5-6. S. brevifolia. Figures represent half of the actual leaf size.

72 Rhodora [Vol. 64

like a segmental allotetraploid between two ecotypes of S.
pallida.

S. ovata was once described as a variety of S. leiandra but
morphological data do not provide any basis for a direct
relationship between the two species. Our studies indicate
that S. leiandra is a somewhat specialized species directly
related to one of the more primitive species, S. australis.
The line of evolution here is apparently different from that
of S. purpurea, S. pallida and S. brevifolia. S. ovata and S.
brevifolia were included as varieties of S. leiandra at the time
when the genus Setcreasea did not exist. The early refer-
ence indicates a broad group relationship rather than a
direct close one.

The most characteristic feature of S. ovata is the presence
of a very strong underground rhizome. This provides a
very efficient method of vegetative reproduction. The other
experimentally known species of this genus also possess a
vegetative mode of reproduction. These species, as pointed
out earlier, are easily crossed, but they lack an underground
rhizome. In the latter, only aerial branches in contact with
soil for a period of time are able to produce roots. This
method is by no means efficient enough to colonize an area.

Whether strong vegetative reproduction is a direct out-
come of polyploidy or if it has evolved through selection
cannot be answered without further experimentation and
field study. Absence of a rhizome in herbarium specimens
may not necessarily mean its absence in nature. It may
be due to incomplete collection. It may only be said that a
capacity for vegetative propagation is present in the near
relatives, a condition preliminary to efficient vegetative
reproduction, as pointed out by Darlington (1956).

Within S. ovata a range of variation in leaf shape is found
from narrowly elliptic to ovate. Superficially it looks like
a range of variation that would be expected of intermediates
between S. pallida and S. brevifolia (fig. 1-6). From the
available data it does not seem to be the result of introgres-
sion involving these two species. First of all, no tetraploid
S. pallida has been reported, and S. ovata is not a triploid.
On the basis of experimental evidence from crossing data

1962] Faruqi, Celarier and Mehra — Setcreasea ovata 73

a free exchange of genes with S. brevifolia does not seem to
be probable. One way this variation could be explained is
through mutations as found in Cyrilla (Thomas, 1960).

ACKNOWLEDGEMENTS
Thanks are extended to the curators of the Gray Herbarium
of Harvard University, Cambridge; United States National
Herbarium, Washington, D. C.; Missouri Botanical Garden, St. Louis;
Chicago Natural History Museum, Chicago, and Herbarium of the
University of Michigan, Ann Arbor for the loan of the herbarium
specimens. The authors are also indebted to Mr. Robert M. Ahring, in
charge of U.S.D.A. Seed Testing Laboratory, Stillwater, for the
germination tests on hybrid seeds. A critical review by Professor
Reed C. Rollins, Director, Gray Herbarium, Harvard University,
Cambridge, Mass., and helpful suggestions of Professor Jack R.
Harlan, Department of Agronomy, and Dr. J. M. J. de Wet Depart-
ment of Botany, Oklahoma State University, Stillwater, Oklahoma
are highly appreciated. — DEPARTMENT OF BOTANY AND PLANT
PATHOLOGY, OKLAHOMA STATE UNIVERSITY, STILLWATER, OKLAHOMA.

LITERATURE CITED

BosE, SMRITIMOY. 1959. A cytological study of Setereasea purpurea
Boom and of an aneuploid taxon of S. brevifolia (Torr.) Rose.
Dissertation Abstract. 20 (4): 1132.

COULTER, J. M. 1890. Upon a collection of plants made by G. C.
Nealley, in the region of the Rio Grande, in Texas, from Brazos
Santiago to El Paso County. Contr. U. S. Nat. Herb. 1: 29-61.

DARLINGTON, C. D. 1956. Chromosome Botany. George Allen and
Unwin Ltd. London, pp. 1-186.

JonNsTON, I. M. 1944. Plants of Northern Mexico, III. Jour. Arn.

(AEDs 25: 54.

MATUDA, E. 1955. Las Commelinaceas Mexicanas. Anal. Inst. Biol.
Mex. 26: 303-432.

RoLLiNs, REED C. 1953. Cytogenetieal approach to the study of
genera. Plant genera — their nature and definition. A symposium.
Chronica Botanica. 14: 133-139.

RosE, J. N. 1899. Treleasea, A new genus of Commelinaceae. Contr.
U.S. Nat. Herb. 5: 207-208.

1903. Studies of Mexican and Central American Plants
— No. 3. Contr. U. S. Nat. Herb. 8: 1-55.

1911. Studies of Mexican Central American Plants —
No. 7a. Contr. U. S. Nat. Herb. 13: 291-312.

SCHUMANN, K. ET Sypow (1899) 1901. In Just, Bot Jahrb. 27 (1) : 452.

THomas, J. L. 1960. A monographic study of the Cyrillaceae. Contr.
Gray. Herb. 186. 114 p.

Torrey, J. 1859. Botany of the Boundary. In Emory, U. S. and Mex.
Bound. Survey. 29-270.

74 Rhodora [Vol. 64

THE VARIETIES OF LUZULA ACUMINATA'

JOHN E. EBINGER

The woodrush, Luzula acuminata, is confined to the east-
ern United States and Canada where it is of rather sporadic
occurrence. This early-blooming species is usually restricted
to moist open woodlands but is occasionally found
along roadsides and in other open areas. It was not until
the work of Rafinesque (1840) that the specific distinctions
between this North American species and the European
species L. pilosa were noticed. Even here Rafinesque men-
tions that this North American species, L. acuminata, is
perhaps L. pilosa. Before this time the American species
was combined with L. pilosa, although Schultes and Schultes
(1829) and Hooker (1840) considered it a variety of L.
pilosa.

Watson (1879) also considered that the North American
species was specifically distinct and, apparently unaware of
Rafinesque's earlier name, proposed for it the name Luzula
carolinae. Later, Fernald (1903), also apparently unaware
of the earlier name by Rafinesque, proposed the name L.
saltuensis for the North American species. In contrast,
he considered L. carolinae to represent a local species of
the Carolina mountains, similar to the Asiatic species L.
plumosa.

There has been a difference of opinion concerning the
treatment of the Luzula acuminata complex. Some authors
have treated this complex as being one species, others as two
species and others as varieties of one species. In most
instances the two entities are recognized as varieties and
both Fernald (1950) and Gleason (1952), in the two major
floras of the northeastern United States, treat the species
in this manner. In both cases the two varieties are sepa-
rated on the basis of the number of secondary pedicels in the
inflorescence though Fernald (1950), also mentions that
the color of the perianth may differ.

The author would like to thank Dr. John R. Reeder for his helpful criticisms during
the progress of this study and the preparation of the paper. I am also grateful to
the curators of the Gray Herbarium (GH) and of the United States National
Herbarium (vs) for the loan of herbarium material.

1962] Ebinger — Varieties of Luzula acuminata 75

Jones (1950) used the binomial Luzula saltuensis in his
flora of Illinois since he does not agree with Fernald regard-
ing the name L. acuminata. Jones (1951) suggests that
this name should be rejected as a nomen dubium because,
in his opinion, Rafinesque’s description is ambiguous. He
objects to the fact that Rafinesque described the North
American plant as having glabrous leaves that are less than
three inches long; an inflorescence which is congested and
not exceeding the leaves; and flowers that are small.

When considering fruiting material the objections of
Jones are well founded; however, with respect to flowering
material Rafinesque’s description is correct. The leaves
in young material, particularly the cauline ones, are less
than three inches long and quite glabrous except for the
silky marginal hairs that are usually present in all Luzula
species. Also, some specimens of this species have been
found that are completely glabrous. In young flowering
material the inflorescence is congested and does not exceed
the leaves and the flowers are small. In my opinion the
description by Rafinesque is not correct when fruiting
material is considered, but is when young flowering material
is used. Furthermore, the fact that Rafinesque states that
the inflorescence is corymbose and that the plant is closely
related to L. pilosa, excludes any other Eastern North
American species of Luzula.

Though Rafinesque’s description leaves no doubt as to
which species he was referring, it is impossible to determine
with certainty which of the two entities (acuminata or caro-
linae) he was describing. However, the fact that he men-
tions that the type locality is boreal America indicates he
was referring to the northern variety (var. acuminata).
Furthermore, this same interpretation was made by Fer-
nald (1944) when he considered the northern entity to be
variety acuminata and reduced Luzula carolinae S. Wats. to
a variety of L. acuminata. Along with this confusion con-
cerning the correct name there is some lack of agreement as
to whether one or two species are involved. In this paper
it will be shown that there is only one species in this com-
plex, but that there are two varieties that can be separated

76 Rhodora [Vol. 64

and that each has a distinct geographic distribution.
MATERIALS AND METHODS

The basis of this study was herbarium material, and
more than 200 collections from Northeastern United States
and Canada were studied. The herbarium specimens were
examined for the characters mentioned below and a diligent
search was also made for any additional morphological
criteria which might be of significance. For determination
of anther and filament lengths flowering material was used
since after the pollen is discharged there is a slight decrease
in anther size. For all other measurements and observations
mature fruiting specimens were studied. Finally, the results
obtained were plotted on a map to show the geographic
distribution of the two varieties.

All material used in this study was obtained from the
U.S. National Herbarium (Us), the Gray Herbarium (GH),
and the Yale University herbarium (YU).

DISCUSSION

To separate the two varieties of Luzula acuminata, Fer-
nald (1938) used the relative lengths of the filaments and
anthers, the length of the bracts at the base of the inflores-
cence, the color of the perianth segments, and the forking
rays of the inflorescence. The present study has revealed
that most of these characters are completely unreliable: in
fact, the only usable trait is the forking rays of the inflores-
cence (secondary pedicels that develop just beneath the
flowers to form a compound corymb).

The anther and filament of every flowering specimen was
measured and it was found that there is no appreciable
difference in the two series. In variety carolinae the aver-
age filament length is 0.5 mm and the anther length is 1.2
mm, while in variety acuminata the average length is
0.4 mm for the filament and 1.4 mm for the anther. Not
only are the lengths very similar, but the variation over-
laps so that it is impossible to distinguish between the two
series by using this character.

Measurements were also made of the bract at the base of
the inflorescence. In both varieties the length is nearly the
same and it is always shorter than the fruiting inflores-

1962] Ebinger — Varieties of Luzula acuminata 77

cence. Fernald (1938) mentions that the type of variety
carolinae has the inflorescence overtopped by an erect frond-
ose bract. Upon careful examination of this type specimen
(Gray and Carey, July 1841), it was found that the frondose
bract, mentioned by Fernald, is actually the upper cauline
leaf. Above this leaf is a small bract that does not overtop
the inflorescence.

Fernald (1938) also mentions that castaneous perianth
segments are common in variety carolinae. A number of
specimens of this variety do have castaneous sepals and
petals, but this character is also found in some specimens
of variety acuminata and therefore cannot be used to sepa-
rate the two entities. Also, some specimens of variety
carolinae have stramineous to light brown perianth
segments, a characteristic which is common in variety
acuminata.

The only reliable character found to separate the two
varieties is the number of forking rays, i.e. secondary pedi-
cels, in the inflorescence. In variety acuminata the number
of primary pedicels in the corymbose inflorescence varies
from 7 to 16 with an average of 10 or 11. Secondary pedicels
are usually lacking in this variety, but occasionally one or
two are found. When secondary pedicels occur they are
restricted to one or two of the primary pedicels. Also, they
are usually restricted to one inflorescence of the specimen
while the other inflorescences lack secondary pedicels. In
variety carolinae, in contrast, the inflorescence is always a
compound corymb with the number of primary pedicels
varying from 9 to 18 and with an average of 12 or 13.
In all the inflorescences of a specimen of this variety, many
of the primary pedicels have at least one secondary pedicel
growing from them. Sometimes 2, 3 or 4 secondary pedicels
are found on one or more of the primary pedicels. The
average number of secondary pedicels found in an inflores-
cence of this variety is 8 or 9 but some inflorescence have
as many as 20. Since variety acuminata sometimes has a
few secondary pedicels in the inflorescence the ratio of the
number of secondary pedicels to the number of primary
pedicels was used as a means of separating the two varieties.

78 Rhodora [Vol. 64

In scoring this character the primary and secondary
pedicels in all of the inflorescences of each specimen were
counted. From this count the ratio of the number of sec-
ondary to primary pedicels was found and the results were
then plotted on a map (Fig. 1) to determine if the two

RATIO OF SECONDARY TO PRIMARY PEDICELS
o 0.0 TO OI
e Oo! TO O3

e. ABOVE 0.3
NUMBER OF SECONDARY PEDICELS ON ANY
ONE PRIMARY PEDICEL
b ONE
TWO
ó THREE

Fic. I. Map showing the distribution of Luzula acuminata war. acuminata and
Luzula acuminata var. carolinae. The open circles indicate specimens of variety
acuminata, the darkened circles indicate specimens of variety carolinae and the half
darkened circles indicate specimens that are intermediate between the two varieties
with respect to the ratio of the number of secondary pedicels to primary pedicels,
The bars radiating from the circles indicate the number of secondary pedicels that
are present on any one of the primary pedicels of the inflorescence.

series have different geographic distributions. On this map
an open circle indicates that the ratio is between 0 and 0.1
while a half darkened circle indicates a ratio between 0.1

1962] Ebinger — Varieties of Luzula acuminata 79

and 0.3 and a completely darkened circle indicates that the
ratio is greater than 0.3.

The number of secondary pedicels on any one primary
pedicel is also indicated on the map. This character is shown
by the bars radiating from the circle. A bar radiating from
the left of the circle indicates that at least one primary
pedicel in the inflorescence has one secondary pedicel grow-
ing from it, while a bar radiating from the top of the circle
indicates that at least one primary pedicel has two secondary
pedicels growing from it, and a line radiating from the right
indicates that three secondary pedicels are present on at
least one primary pedicel.

As can be seen from the map, variety carolinae has a
southern distribution which extends from Alabama and
Georgia, west to the Mississippi River, and north to southern
Ohio and Pennsylvania. In contrast, variety acuminata has
a more northern distribution. The southern boundary of this
variety is southern Ohio and Pennsylvania, and in the west
it extends to South Dakota and Manitoba, Canada. It is
also common in southeastern Canada and the New England
States.

In general, the region of high variability between the two
varieties includes the states of Ohio, Pennsylvania, southern
New York, West Virginia, Maryland and New Jersey. In
this region of high variability there are numerous inter-
mediate plants as well as some plants that are typical of both
varieties, Excluding this region of high variability the two
varieties are fairly constant in their characters. Variety
carolinae, which is represented by the darkened circles with
several bars radiating from them, is the southern variety.
Of the 50 specimens studied from this area all but six have
a ratio greater than 0.3. Of these six specimens, five have
a ratio between 0.1 and 0.3 and the remaining specimen has
a ratio of 0.0 and should be referred to variety acuminata.
Also, most of the southern specimens have 2, 3 or more
secondary pedicels growing from at least one primary
pedicel in an inflorescence.

The northern variety (acuminata) is not as constant in
its characters, but there can be no doubt in distinguishing

80 Rhodora [Vol. 64

the two series. Of the 125 specimens studied from this
region, 92 lack secondary pedicels. In the remaining 33
specimens, only eight have a ratio of secondary pedicels to
primary pedicels between 0.1 and 0.3 while the remainder
have a ratio of less than 0.1. No specimens were found with
a ratio greater than 0.3. Also, of the 33 specimens that have
secondary pedicels, only five have more than one secondary
pedicel growing from any of the primary pedicels of the
inflorescence, and in most cases no more than one secondary
pedicel was found in an inflorescence.

The secondary pedicels found in the southern variety do
not seem to be a result of a longer growing season. Even
in young flowering specimens the flowers on the secondary
pedicels are as well developed as the flowers on the primary
pedicels, and both produce seed at nearly the same time.
This indicates that the existence of secondary pedicels is not
a result of the environment but that it is a genetically con-
trolled factor.

TAXONOMIC TREATMENT

Since the results of this paper show that there are two
varieties in this complex, a short taxonomic treatment of the
two will be undertaken. This should help to clarify the
nomenclature of the varieties since in the past there has been
confusion as to the correct name.

Luzula acuminata Raf. Autikon Bot. 193. 1840
var. acuminata
Luzula pilosa (L.) Willd. var. B. Americana Schultes &

Schultes Syst. Veg. 7:262. 1829.

Luzula pilosa (L.) Willd. var. &. Hook. Fl. Bor.-Am. 2: 188.

1840.

Luzula saltuensis Fern. Rhodora 5:195. 1903.
Juncoides saltuense (Fern.) Heller, Muhlenbergia 6:12.

1910.

Juncoides pilosum (L.) Coville var. michiganense Farwell,

Rep. Mich. Acad. Sci. 20:170. 1918.

Juncoides pilosum (L.) Coville var. saltuense (Fern.) Far-

well, Rep. Mich. Acad. Sci. 20:170. 1918.

Luzula carolinae S. Wats. var. saltuensis (Fern.) Fern.

Rhodora 40:404. 1938.

1962] Ebinger — Varieties of Luzula acuminata 81

This is the northern variety and can be separated from
the more southern variety by the lack of secondary pedicels
in the inflorescence. In specimens where secondary pedicels
exist the ratio is always less than 0.3 and rarely is there
more than one secondary pedicel to any one primary pedicel.

Luzula acuminata Raf. var carolinae (S. Wats.)
Fern. Rhodora 46:5. 1944
Luzula carolinae S. Wats. Proc. Am. Acad. 14:302. 1879.
Juncodes carolinae (S. Wats.) O. Ktze. Rev. Gen. Pl. 2:724.

1891.

This variety has a much more southern distribution than
variety acuminata. It can be distinguished from the more
northern entity by the larger number of secondary pedicels
in the inflorescence. Usually some of the primary pedicels
have two or more secondary pedicels growing from them.

CONCLUSIONS

The above results show conclusively that there is a distinct
difference between the two varieties and that each has a
definite geographic range. Variety acuminata has a more
northern distribution than variety carolinae and can easily
be separated from the latter by its lack of secondary pedicels
in the inflorescence. In the specimens of variety acuminata
that do have secondary pedicels the ratio of these to the
primary pedicels is always less than 0.3 and in most cases
there is no more than one secondary pedicel to any of the
primary pedicels. Just the opposite condition exists in the
southern variety (carolinae). Here a ratio above 0.3 is
always found and the ratio is sometimes greater than 1.0.
Also most specimens of this southern variety have two or
more secondary pedicels on some of the primary pedicels
of the inflorescence.

In the region of high variability between the two series
are a number of specimens that are difficult to place tax-
onomically. The large amount of variability in this region
as well as the specimens of the northern variety that have
secondary pedicels show that the two series intergrade into
each other. This variability indicates that only two varieties
are involved and that they are not separate species.

82 Rhodora [Vol. 64

LITERATURE CITED

FERNALD, M. L. 1903. The American Representatives of Luzula
vernalis. Rhodora 5:193-196.

1938. Noteworthy Plants of Southeastern Virginia.

Part I. Rhodora 40:364-424.

1944. Overlooked Species, Transfers and Novelties

in the Flora of Eastern North America. Rhodora 46:1-21.

1950. Gray's Manual of Botany. ed. 8. New York,
American Book Co. Ixvi+ 1632 pp. (Luzula pp. 416-420).

GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora
of the Northeastern United States and Adjacent Canada. 3 Vol.
Lancaster, Lancaster Press. (Luzula 1:400-402).

Hooker, W. J. 1840. Flora Boreali-Americana. 2 Vol. (Luzula
2:187-189).

JoNES, G. N. 1950. Flora of Illinois. Notre Dame, Univ. of Notre
Dame Press. vi4-368 pp. (Luzula p. 96).

1951. On the Nomenclature of Luzula saltuensis.
Rhodora 53:242-244.

RAFINESQUE, C. S. 1840. Autikon Botanikon. (Luzula p. 193).

ScHULTES, J. A. & SCHULTES, J. H. 1829. Systema Vegetabilium.
T:(Luzula pp. 254-279).

WATSON, S. 1879. Contributions to American Botany. II. Descrip-
tion of some new species of North American plants. Proc. Amer.
Acad. 14:288-302.

REPRESENTATIVE SPECIMENS: Luzula acuminata Raf. var. acuminata
UNITED STATES: South Dakota; E. J. Palmer 37562 (GH). Minnesota:
Butters & Abbe 10 (GH); J. W. Moore 21601 (us); J. B. Moyle 387
(us); Pease & Bean 26387 (GH); Rosendahl 4966 (GH); Rosendahl &
Butters 4614 (GH). Iowa: Thorne 10005 (us). Wisconsin: Colby 4472
(us); Fassett 2751 (GH) 2744 (GH). Michigan: Farwell 2559 (GH),
4817 (GH), 4864 (GH), 5421 (GH). New York: A.J. Eames 290 (GH);
Haberer 962 (GH); House 8817 (GH), 9346 (GH), 21354 (GH), 22402
(GH); MeVaugh 4066 (GH); Muenscher & Bachtel 104 (US) ; Phelps
282 (GH, US); Wherry & Muenscher 15308 (GH); Wiegand 1968 (GH).
Pennsylvania: Fogg 12124 (GH). Maryland: Hermann 14936 (US).
Virginia: Hunnewell 18931 (GH). New Jersey: Long 31981 (GH),
37425 (GH), 37478 (GH). Connecticut: Ebinger 90 (vv); Parker
5492 (YU); Weatherby 2403 (vU). Massachusetts: Hill & St. John
1698 (YU). New Hampshire: Bissell 1498 (vU); Edmondson 4189
(GH); A. H. Moore 3408 (us). Maine: Fellows 1768 (US), 2368 (vs);
Fernald 85 (GH, US, YU), 2510 (GH); Ricker 180 (US); St. John &
Nichols 2203 (vv, US). Indiana: Deam 44268 (GH). CANADA: Ontario:
Hosie, Losee & Bannan 1512 (GH); Dutilly & Lepage 36312 (US) ;
Montgomery 1069 (GH); Soper 2652 (GH). Quebec: Chrysler 1181
(Us); Collins, Fernald & Pease 5462 (GH); Dutilly & Lepage 15036
(GH); Rouleau 1689 (GH); Rousseau 2621 ( GH, us); St. John 1824

1962] Daubs — Spirodela oligorrhiza 83

(GH); Marie-Victorin 28491 (GH); Marie-Victorin & Rolland-Germain.
25793 (GH, US), 27117 (GH), 29264 (GH), 45674 (GH), 47459 (GH).
Prince Edward Island: Fernald & St. John 10988 (GH). Nova Scotia:
Bissell & Linder 20733 (GH); Fernald & Long 23583 (GH); Long &
Linder 20732 (GH), 20735 (GH); Nichols 79 (YU), 542 (YU, GH) ; Pease
& Long 20734 (GH, US). Newfoundland: Pease & Edgerton 27216
(GH).

Luzula acuminata Raf. var. carolinae (S. Wats.) Fern. UNITED
STATES: Alabama: Harper 3703 (GH, US), 3956 (US). Georgia: Allard
81 (US), 82 (US); Cronquist 4979 (GH); D. Eyles 6863 (GH); Harper
2056 (GH, US), 2062 (GH, US); Hermann 10186 (GH); Muenscher &
Smith 2954 (GH). South Carolina: House 1847 (us); E. J. Palmer
35405 (GH). North Carolina: Correll 5020 (GH); Godfrey 3414 (GH),
3814 (GH); Godfrey, Campana & Fox 48070 (GH); Godfrey & Fox
50307 (GH); Godfrey, Fox & Woods 49111 (GH); Godfrey & White
7013 (GH, US); Gray & Carey (July, 1841) (GH-HOLOTYPE); House
4130 (US); Hunnewell 10272 (GH), 14188 (GH). Tennessee: Hunine-
well 15158 (GH); Nease 194 (US). Kentucky:Mecinterr & Shacklette
615 (US). Ohio: Leonard 551 (US), 552 (US). West Virginia: Dickey
244 (GH); Fosh 1034 (US). Virginia: Fernald & Long 6958 (GH),
6959 (GH), 6960 (GH), 6961 (GH), 7787 (GH), 7788 (GH), 14526 (an);
Fernald, Long & Abbe 14123 (GH); Fernald, Long & Pease 11657 (GH,
US), 11658 (GH); Grimes 3400 (GH).

THE OCCURRENCE OF SPIRODELA OLIGORRHIZA
IN THE UNITED STATES

EDWIN H. DAUBS!

Spirodela oligorrhiza (Kurz) Hegelm. of the Lemnaceae
was first reported and described by Kurz (1867) from India
under the binomial Lemna oligorrhiza. Shortly thereafter
Hegelmaier (1868) transferred the species to the genus
Spirodela, and reported its further occurrence in Australia
and Java. At the same time he also described and named
four varieties. Later (1896) he gave each of these varieties
species status.

The species remained unreported outside of this Far
Eastern area until Saeger (1934) recorded it from two
locations in Missouri. The first of these was made by him
in Swope Park, Kansas City, and the second by F. H. Woods
from a pond in southwestern Missouri. It is also reported
that this pond contained goldfish, indicating the probability

Department of Botany, University of Illinois, Urbana.

84 Rhodora [Vol. 64

that the plant was introduced from commercial supply
sources for domestic aquaria. More recently, Mason (1957)
has reported the plant as occurring in the vicinity of Ber-
keley, California, but without a specific location, and this
appears to be another instance of the same nature as
reported by Saeger.

Within the past few years however, a considerable number
of new collections from widely separated areas have been
made. The author has collected this species from six loca-
tions in Florida, two in Louisiana, and from one in Illinois.
The voucher specimens for these collections are in the Her-
barium of the University of Illinois. Also from Florida five
collections were reported (personal communication) by
Professor R. K. Godfrey and Richard D. Houk.

The specific citations for my own collections are as follows: FLORIDA.
— PASCO CO., small pond south of Mazaryktown along U.S. highway
41, Feb. 1, 1961, Daubs 651; SARASOTA CO., drainage ditch along Fruit-
ville Road east of Sarasota, Feb. 3, 1961, Daubs 666; HILLSBOROUGH
CO., ditch along U.S. 301 north of Tampa, Feb. 9, 1961, Daubs 670;
PASCO CO., pond north of Dade City along U. S. 301, Feb. 9, 1961, Daubs
671; SUMTER CO., ditch along roadside near junction of U. S. 301 and
Fla. Rd. 42, Feb. 9, 1961, Daubs 672; MARION CO., pond near U. S. 301
between Ocala and Pedro, Feb. 9, 1961, Daubs 673. LOUISIANA. — ST.
CHARLES PARISH, ditch along U. S. 90, 1/2 mile east of Bouette, May 31,
1961, Daubs 782; LA FOURCHE PARISH, ditch along U.S. 90, between
Raceland and Houma, May 31, 1961, Daubs 786. ILLINOIS. — ALEX-
ANDER CO., Horseshoe Lake, Nov. 24, 1960, Daubs 545; same location,
June 2, 1961, Daubs 803.

From California, Dr. L. R. Heckard of the Jepson
Herbarium, University of California, Berkeley, reports
(personal communication) a collection of S. oligorrhiza from
Millerton Lake, Bacigalupi and. Heckard 7693, Apr. 30, 1961,
"most certainly naturally occurring" and further remarks,
“There must be many more localities in California".

This recent plethora of reports is probably not due to any
sudden widespread dissemination of the species but rather
to the failure of potential collectors to identify it from eye
level, confusing it with Lemna minor, to which it is usually
similar in size. It can be easily separated from any of the
Lemna species through the possession of more than one root,

1962] Banks — Paspalum fimbriatum 85

usually 2-4, and from S. polyrrhiza which usually has 5-many
roots, and is orbicular to obovate in shape, as well as usually
much larger in size. It would appear that S. oligorrhiza,
most probably introduced as indicated, has now become well
established in many areas, and will doubtless continue to
spread throughout the area to which it is adapted. The
plant is not known to produce the familiar overwintering
turions found in S. polyrrhiza, but it flowers rather readily
and the seeds may provide an adequate overwintering form
for those areas too cold for vegetative survival.
LITERATURE CITED
HEGELMAIER, F. 1868. Die Lemnaceen. Leipzig.
1896. Systematische Ubersicht der Lemnaceen.
Bot. Jahrb. 21:268-305.
Kurz, S. 1867. Enumeration of Indian Lemnaceae. Jour. Linn.
Soc. Bot. 9:264-268.
Mason, H. L. 1957. A flora of the marshes of California. Univer-
sity of California Press, Berkeley.
SAEGER, A. 1934. Spirodela oligorrhiza collected in Missouri. Bull.
Torr. Bot. Club 61:233-236.

PASPALUM FIMBRIATUM IN THE UNITED STATES
DONALD J. BANKS

While searching for plants of the Setacea group of the
genus Paspalum in southern Florida, I found several plants
of Paspalum fimbriatum HBK. growing under natural
conditions. This annual species is very distinct because of
its winged spikelets. It has not been previously reported as
occurring in the continental United States. According to
Chase (1929), the species has been collected in Panama,
the West Indies, northern South America, and the Hawaiian
Islands.

Erdman West! of the University of Florida reported
receiving recently for identification a sample of fimbria-
tum which had been collected in a lawn near Homestead,
Dade County, Florida by F. C. Craighead on Sept. 7, 1961
(FLAS 82141). West remarked that Paspalum fimbriatum
had been planted in the “old grass garden" at Gainesville
and was collected Aug. 19, 1922 by W. E. Stokes (FLAS 3545)

1Personal communication.

86 Rhodora [Vol. 64

but it apparently had not persisted as it was not collected
again. According to Swallen', Smithsonian Institution, and
Godfrey’, Florida State University, no U.S. specimens are
on file in their herbaria. Thus, these Florida collections are
probably the first of this species for the continental United
States.

My first collection was made on Plantation Key on Sept.
9, 1961. Later that day another collection was made near
Florida City, approximately 39 miles from the first site.
The following day a single plant was collected near Home-
stead approximately 7.2 miles northwest of the second site.

It is possible that this species could be a relict. However,
since it had not been discovered previously even though con-
siderable plant collecting has been done in Florida, it appears
that the species may be a new immigrant. It seems probable
that the light winged seeds could have been carried from
the West Indies by recent hurricanes. Botanists should look
for the species to more adequately determine its distribution.

My collections of Paspalum fimbriatum (all from Florida) are as
follows:

Roadside on dry coral, Plantation Key, Monroe Co. Associated with
Paspalum caespitosum Fliigge and Paspalum ciliatifolium Michx. Sept.
9, 1961. D. J. Banks 1223; Miami oolitic limestone 3.2 miles south of
Florida City on U. S. Highway no 1, Dade Co. Associated with Paspa-
lum caespitosum Flügge. Sept. 9, 1961. D. J. Banks 1242; Miami
oolitic limestone, approximately 2 miles northwest of Homestead, Dade
Co. Associated with Chloris petraea Swartz, Paspalum caespitosum
Flügge, and Paspalum pubescens Muhl. Sept. 10, 1961. D. J. Banks
1260.

Specimens are being deposited in the herbaria of the Chicago
Natural History Museum, Florida State University, University of
Florida, University of Georgia, Harvard University, Missouri Botani-
cal Garden, New York Botanical Garden, and the Smithsonian Institu-
tion. — DEPARTMENT OF BOTANY, UNIVERSITY OF GEORGIA,

LITERATURE CITED

CHASE, AGNES. 1929. The North American Species of Paspalum.
Contrib. U. S. Natl. Herb. 28:235.

1962] Ward — The Genus Anonymos 87

THE GENUS ANONYMOS AND ITS
NOMENCLATURAL SURVIVORS

DANIEL B. WARD

Thomas Walter, in his pioneer descriptive study, Flora
Caroliniana (1788), named and described over 300 species
which he believed to be new to science. A majority of these
species truly were new, and Walter’s names are of familiar
occurrence in articles dealing with the flora of the south-
eastern United States.

Among his many new species Walter detected representa-
tives of what he considered to be undescribed genera. He
recognized 32 of these new genera and placed in them 52
species, about half also new, the rest transfers from other
genera. He named four of these genera, containing eight
species, in a conventional manner, while he modestly labeled
the remaining 28 with the ambiguous pseudonym
Anonymos.

Although there has never been doubt that Walter intended
Anonymos to be merely a provisional generic name, there
has been dispute as to whether the specific epithets published
under this name were available for transfer to more precisely
defined genera. In the United States, following the lead of
M. L. Fernald and S. F. Blake, modern authors have treated
Walter’s epithets as legitimate and have recognized their
transfer to other genera. But in June, 1950, the Nomencla-
ture Section of the VIIth International Botanical Congress
(Stockholm) decreed otherwise, voting that all “binary
combinations of a specific epithet with the word Anonymos"
were illegitimate.

legitimacy of the epithets used by Walter in connection
with the provisional name Anonymos means, of course, that
they are not available for transfer to other genera and
Walter cannot properly appear in the conventional author
citation of the legitimate name.' But illegitimacy does not
mean that the same epithet cannot be used for the same

1The Code (Art. 23) does permit the optional inclusion of Walter's name if enclosed
in brackets. An alternative and perhaps preferable method of full citation is the use
of "ex" beween Walter’s name and that of the publishing author (Rec. 46A).

88 Rhodora [Vol. 64

species by a later author; if an epithet from an illegitimate
name is used again in a different genus it is treated, not as
a transfer, although that may have been the intent of the
later author, but for all practical purposes as a new epithet.

Even preceding the vote against Anonymos it was realized,
as has recently been pointed out again by Wilbur (Rhodora
63:37. 1961), that many of Walter’s epithets were transfer-
red to other genera by J. F. Gmelin in his Systema Naturae,
published in 1791 and 1792, before other authors had occa-
sion to re-describe and rename the same plants. Gmelin, it
is true, often failed to provide a specific description, but this
requirement for valid publication is fulfilled in the case of
monotypic genera by the generic description, abstracted
from Walter, and in all cases by reference to Walter and the
appropriate specific description. Often validation is further
secured by the existence of specimens which may be taken
to be types in the John Fraser collection on deposit at the
British Museum. The familiar epithets of Walter, then,
remain available for those plants where they have been
used in the past, the only changes occurring in the author
citation.

A few of the epithets transferred by Gmelin resulted in
the formation of unfamiliar synonyms such as Mattuschkia
aquatica, not recognized by him to be based on the long
known Saururus cernuus L. But 19 of Walter’s Anonymos
epithets, together with a twentieth omitted by Gmelin but
picked up by Willdenow, survive to the present in the correct
names of members of the Southeastern flora. With the
expectation of providing a convenient source for determining
the proper citation for these names, the following list has
been prepared.

Agalinis setacea (J. F. Gmel.) Raf., New Fl. 2: 64. 1837, replaces
A. setacea (Walt.) Raf. (syn. Gerardia setacea (Walt.) J. F. Gmel.;
Gerardia rejected for conservation, Taxon 10:124. 1961). Basonym
(Gerardia setacea J. F. Gmel, Syst. 2:928. 1792) is inadequately
described (“G. foliis setaceis, caule ramosissimo, floribus terminali-
bus"), but is validated by reference to Walter (Fl. Carol. 170. 1788).

Burmannia capitata (J. F. Gmel.) Mart., Nov. Gen. Sp. 1:12. 1823,
replaces B. capitata (Walt.) Mart. Basonym (Vogelia capitata J. F.
Gmel., Syst. 2: 107. 1791) lacks specific description, but is validated

1962] Ward — The Genus Anonymos 89

by generic description and by reference to Walter (Fl. Carol. 69.
1788).

Crotalaria rotundifolia J. F. Gmel., Syst. 2: 1095. 1792, replaces
C. rotundifolia (Walt.) Poir. in Lam. (syn. C. maritima Chapm.).
Description is inadequate for separation from C. angulata Mill. (*Cr.
foliis rotundatis integris pilosis sparsis, caule subdecumbente"), but is
validated by reference to Walter (Fl. Carol. 181. 1788) and by
Walter's type (photo, Rhodora 50: plate 1107. 1948). C. rotundifolia
Poir. in Lam. is a later homonym; Poiret (Enc. suppl. 2: 402. 1812)
described a quite different plant and made no reference to Walter or
Gmelin.

Cynoctonum sessilifolium J. F. Gmel., Syst., 2: 443. 1791, replaces
C. sessilifolium (Walt.) J. F. Gmel. Description is inadequate (“C.
foliis sessilibus"), but is validated by reference to Walter (Fl. Carol.
108. 1788).

Dioscorea quaternata J. F. Gmel, Syst. 2: 581. 1791, replaces
D. quaternata (Walt.) J. F. Gmel. Description is brief but diagnostic.
Reference is made to Walter (Fl. Carol. 246. 1788).

Elytraria caroliniensis (J. F. Gmel.) Pers., Syn. Pl. 1: 23. 1805,
replaces E. caroliniensis (Walt.) Pers. (syn. E. virgata Michx.).
Basonym (Tubiflora caroliniensis J. F. Gmel., Syst. 2: 27. 1791) lacks
specific description, but is validated by generie description and by
reference to Walter (Fl. Carol. 60. 1788).

Houstonia procumbens (J. F. Gmel.) Standley, N. Am. Flora 32: 26.
1918, replaces H. procumbens (Walt.) Standley (syn. H. rotundifolia
Michx.). Basonym (Poiretia procumbens J. F. Gmel. Syst. 2: 263.
1791) has inadequate description (^P. caule procumbente"), but is
validated by reference to Walter (Fl. Carol. 86. 1788).

Lachnanthes tinctoria (J. F. Gmel.) Ell., Sketch Bot. S. C. & Ga. 1:
47. 1816, replaces L. tinctoria (Walt.) Ell. Basonym (Heritiera tinc-
torium J. F. Gmel, Syst. 2: 113. 1791, “tinctorum”) lacks specific
description, but is validated by generic description and by reference
to Walter (Fl. Carol. 68. 1788).

Liatris graminifolia Willd., Sp. Pl. 3: 1636. 1803, replaces L. gra-
minifolia (Walt.) Willd. Description is adequate. Reference is made
to Walter (Fl. Carol. 197. 1788).

Lithospermum caroliniense (J. F. Gmel.) MacM., Metasp. Minn. Val.
438. 1892, replaces L. caroliniense (Walt.) MacM. (syn. L. gmelinii
(Michx.) Hitche.). Basonym (Batschia caroliniensis J. F. Gmel., Syst.
2: 315. 1791) lacks specific description, but is validated by generic
description and by reference to Walter (Fl. Carol. 91. 1788).

Micranthemum umbrosum (J. F. Gmel.) Blake, Rhodora 17: 131.
1915, replaces M. umbrosum (Walt. Blake (syn. M. orbiculatum
Michx.). Basonym (Globifera wmbrosa J. F. Gmel., Syst. 2: 32. 1791,
*"umbros") lacks specific description, but is validated by generic de-
scription and by reference to Walter (Fl. Carol. 63. 1788).

Nymphoides aquatica (J. F. Gmel.) Kuntze, Rev. Gen. 2: 429. 1891

90 Rhodora [Vol. 64

(as “Nymphodes aquaticum”; Hill (Brit. Herb. 77. 1756) originally
spelled the genus “Nymphoides” and treated it as feminine, the gender
advised by Rec. 75A of the International Code,) replaces N. aquatica,
(Walt.) Kuntze. Basonym (Villarsia aquatica J. F. Gmel., Syst. 2:
447. 1791) lacks specific description, but is validated by generic de-
scription and by reference to Walter (Fl. Carol. 109. 1788).

Petalostemon pinnatum (J. F. Gmel.) Blake, Rhodora 17: 131. 1915,
replaces P. pinnatum (Walt.) Blake (syn. P. corymbosum Michx.).
Basonym (Kuhnia pinnata J. F. Gmel., Syst. 2: 375. 1791) has inade-
quate description (“K foliis pinnatis"), but is validated by reference
to Walter (Fl. Carol. 103. 1788).

Planera aquatica J. F. Gmel., Syst. 2: 150. 1791, replaces P. aquatica
(Walt.) J. F. Gmel. Specific description is lacking, but name is vali-
dated by generic description and by reference to Walter (Fl. Carol.
230. 1788).

Ruellia caroliniensis (J. F. Gmel.) Steud., Nom. Bot. ed. 2. 2: 481.
1841, replaces R. caroliniensis (Walt.) Steud. (syn. R. hybrida Pursh).
Basonym (Pattersonia caroliniensis J. F. Gmel., Syst. 2: 925. 1792)
lacks specific description, but is validated by generic description and
by reference to Walter (Fl. Carol. 168. 1788).

Seymeria cassioides (J. F. Gmel.) Blake, Rhodora 17:134. 1915,
replaces S. cassioides (Walt.) Blake (syn. S. tenuifolia Pursh).
Basonym (Afzelia cassioides J. F. Gmel., Syst. 2: 927. 1792) lacks
specific description, but is validated by generic description and by
reference to Walter (Fl. Carol. 171. 1788).

Trilisa odoratissima (J. F. Gmel.) Cass., Dict. Sci. Nat. 55: 310.
1828, replaces T. odoratissima (Walt.) Cass. (T. odoratissima
(Michx.) Cass.). Basonym (Chrysocoma odoratissima J. F. Gmel.,
Syst. 2: 1204. 1792) is questionably adequate (‘‘Chr. foliis radicalibus
ovato-oblongis; caulinis oblongo-lanceolatis laevibus, floribus corymbo-
sis"), but is validated by reference to Walter (Fl. Carol. 198. 1788).

Trilisa paniculata (J. F. Gmel.) Cass., Dict. Sci. Nat. 55: 310. 1828,
replaces T. paniculata (Walt.) Cass. (T. paniculata (Michx.) Cass.).
Basonym (Chrysocoma paniculata J. F. Gmel., Syst. 2: 1204. 1792) is
recognizably described (“Chr. foliis lanceolatis adscendentibus, caule-
que viseoso simplici pilosis, paniculata terminali"), and is supplement-
ed by reference to Walter (Fl. Carol. 198. 1788).

Uvularia caroliniana (J. F. Gmel.) Wilbur, Rhodora, 63:39. 1961,
replaces U. pudica (Walt.) Fern. (syn. U. puberula Michx.). Basonym
(Erythronium carolinianum J. F. Gmel, Syst. 2: 546. 1791) lacks
specific description, but is validated by reference to Walter (“Anony-
mos pudic", Fl. Carol. 123. 1788).

Zornia bracteata J. F. Gmel., Syst. 2: 1096. 1792, replaces Z. brac-
teata (Walt.) J. F. Gmel. Specific description is lacking, but name is
validated by generic description and by reference to Walter (Fl. Carol.
181. 1788).

1962] Ward — The Genus Anonymos 91

The remaining 24 new names, of the 44 created by Walter
by the combination of an epithet with Anonymos, appear
not to have been used as basonyms by later authors. This
neglect is justified since all are either (a) synonyms of
earlier names, (b) homonyms resulting from the improper
application of an earlier name, or (c) not clearly identifi-
able. They are listed and tentatively identified below, to

complete the disposition of Walter’s Anonymos epithets.

aquatic[a] (p. 127)
aquatic[a] (p. 137)
caroliniensis (p. 164)

caroliniens[is] (p. 188)

caroliniensis (p. 263)
ciliat[a] (p. 197)

corymbos[a] (p. 139)

erect[a] (p. 86)
erect[a] (p. 170)

flava (p. 170)
frondos[a] (p. 139)
frutescens (p. 186)
ligustrin[a] (p. 139)
pedicularis (p. 170)

petiolat[a] (p. 108)
pilos[a] (p. 197)

purpurea (p. 170)
quinat[a] (p. 246)

ramos[a] (p. 198)

repens (p. 110)
repens (p. 160)
sagittalis (p. 181)
sempervirens (p. 98)

Saururus cernuus L.

Decodon. verticillatus (L). Ell.

probably Verbena canadensis (L.) Britt.

Amphicarpa bracteata (L.) Fern.

possibly Hydnum erinaceus Bull.

possibly Liatris although not so recognized
by Gaiser (Rhodora 48:165-183, 216-382,
393-412. 1946)

possibly Vaccinium myrsinites Lam., or V.
tenellum Ait., not Vaccinium corymbosum
L. according to range given by Camp
(Brittonia 5:260. 1945)

Houstonia caerulea L.

identified by Pennell (Scroph. of E. Temp.
N. A., 473. 1935) as either Agalinis
(Gerardia) linifolia (Nutt.) Britt., A.
obtusifolia Raf., or A. tenella Pennell

Aureolaria flava (L.) Farw.

Gaylussacia frondosa (L.) Torr.

Wisteria frutescens (L.) Poir. in Lam.

Lyonia ligustrina (L.) DC.

probably Awreolaria pectinata
Pennell

Cynoctonum mitreola (L.) Britt.

possibly Liatris although not so recognized
by Gaiser (op. cit.)

Agalinis purpurea (L.) Pennell

probably Dioscorea quaternata J. F. Gmel.,
according to Anderson (Rhodora 36:345.
1934)

possibly Liatris although not so recognized
by Gaiser (op. cit.)

Dichondra carolinensis Michx.

Phyla nodiflora (L.) Greene

Crotalaria sagittalis L.

Gelsemium sempervirens (L.) Ait. f.

(Nutt.)

92 Rhodora [Vol. 64

uniflor[a] (p. 198) probably Carphephorus _ bellidifolius
(Michx.) T. & G. or C. tomentosus
(Michx.) T. & G.

— UNIVERSITY OF FLORIDA, GAINESVILLE, FLORIDA.

NOTE: After the manuscript for the above paper was submitted to
the editor, Dr. R. L. Wilbur of Duke University very generously made
available to me his independently prepared manuscript on the same
subject. I have made a few changes and corrections in the present
paper, as suggested by his studies and our subsequent correspondence.
A new combination proposed by Dr. Wilbur has been left for his later
treatment. — D. B. W.

JOHN BACHMAN AND THE HAPPOLDT EUROPEAN JOURNAL.
— Reverend John Bachman is remembered as a close asso-
ciate of Audubon for he ably assisted in the writing of both
Birds of America and Quadrupeds, and the two families were
united by Bachman's two daughters marrying Victor and
John Woodhouse Audubon. Less familiar is Bachman's own
interest in botany. In 1835 he published a fifteen-page check-
list of 1030 species entitled Catalogue of Phanerogamous
plants and. ferns, native or naturalized, found. growing in
the vicinity of Charleston, South-Carolina. When Hooker's
friend, William A. Bromfield, visited America, ne spent
about five weeks in Charleston in 1847 and often saw Bach-
man. On February eleventh Bromfield “went to Rev. Dr.
Bachman’s to look over Elliott's herbarium with Mr. Ravenel
of St. John's who has made an extensive collection of the
species about the place. Dr. Bachman drove me out 8 miles
from Charleston to look for Ulmus alata in flower." Some
of Bachman's specimens are lodged at the Academy of
Natural Sciences, Philadelphia, via the Short Herbarium,
and others are in the Durand Herbarium at Paris, but his
own herbarium, along with a valuable library (including a
copy of Catesby's Natural History), was burned in Colum-
bia, whence it had been moved for safe-keeping, at the time
of Sherman's raid.

Bachman at twenty-five had come to Charleston from
Pennsylvania in 1815 and the following year married Harriet
Martin, daughter of a Charleston parson. Harriet's sister
was Maria Martin who drew many botanical backgrounds

1962] Ewan — The Happoldt European Journal 93

for Audubon’s bird plates. After Harriet’s death the Rev-
erend Bachman married Maria.

Though it is not evident from the title! this “Contribution”
from the Charleston Museum contains much of interest to
any reader with a curiosity about South Carolina natural
history. The index is helpful though not exhaustive. More
valuable in some connections than the journal is the eighty-
nine page biography of Bachman prepared by Professor
Claude H. Neuffer, of the Department of English, University
of South Carolina, who “discovered” and edited the journal.

Christopher Happoldt (1823-1878), soldier-surgeon, was
a protegé of Reverend John Bachman, and the journal kept
by young Christopher, aged fifteen, concerned a six months’
tour of Europe, June 5 to December 27, 1838, while acting as
a travelling companion during Bachman's convalescence on
a recuperative sea voyage. Bachman was nearing the zenith
of an active scientific life, all the while fulfilling his duties
faithfully as minister at St. John’s Lutheran Church. He
had assisted Audubon assiduously during several years past.
There are 134 references to Bachman’s aid in Ornithological
Biography alone. To combat a general debility enhanced in
part by a mild attack of cholera in 1836, it had been decided
that a tour of Europe, including a chance to visit Audubon
then busily working in Edinburgh with Macgillivray, would
be refreshing for the pastor. Leaving Charleston on the
Chicora for Liverpool, young Happoldt and Reverend Bach-
man arrived at No. 4 Wimpole Street, the Audubons’ London
Home, to be greeted by Lucy and the family, and to be off for
Edinburgh within two weeks. So begins what amounts to a
lively if somewhat repetitive diary by a frank young man
viewing the European scene.

Upon completing his medical education abroad, Dr. Hap-
poldt practiced in Charleston and edited the local medical
Journal and Review. However this was not to last long
because, on his wife’s insistence, he moved to Morgantown,
North Carolina, and passed from the professional spotlight
to the obscurity of a country doctor. He later served the

Christopher Happoldt Journal. Edited by Claude Henry Neuffer. Charleston
Museum, Charleston, S. C., 1961. p. 214 [plus 14 unnumbered index pp.] $5.00 ppd.

94 Rhodora [Vol. 64

Confederacy, and then the citizens of Vicksburg when a
yellow fever outbreak struck that river city. There he died
“the victim of the disease he had fought against so valiantly
in order that others might live."

The Happoldt diary is of particular interest for its fleeting
glimpses of personalities and places. There is Dr. Frank
Buckland of Oxford; the Prince of Massena, a devotee of
natural history; “Prof. Braum of Carlsbruche" [Heinrich
Georg Bronn, 1800-1862, who translated Darwin’s Origin] ;
Dr. J. C. Faber of Charleston and James Trudeau of New
Orleans, both then visiting Paris; and such institutions as
the Jardin des Plantes in Paris and Knowsley Hall near
Liverpool where Bachman met the Earl of Derby for whose
zoo he had transported live anhingas across the Atlantic!
Yes, they saw the Queen in London town, attended by the
royal family, and her some 1500 Swiss guards “all mounted
on black horses."

Young Christopher sampled dolphin roe for dinner and
pronounced it “certainly very fine.” At Regent Park Zoo
there were three thousand persons on Sunday, July 8th,
"not," as he observed, “for the purpose of seeing the animals
but the company." In true adolescent form, Christopher
paled at the succession of museums, and when Bachman
visited the Tower, he did not go “because he did not like to
see everything twice." However, his sage verdict finally
was:

“I find it a great use to Travel, the man who remains home
all his life, has no idea of mankind; either of the Improve-
ments of the world." — JOSEPH EWAN, TULANE UNIVERSITY.

Volume 63, No. 756, including pages 327-367, was issued January 22, 1962

ow REFERENCE LIBRARY

Ju. 11 1962

Dodota

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by

ALBION REED HODGDON, Editor-in-Chief
ALBERT FREDERICK HILL
STUART KIMBALL HARRIS
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON
RADCLIFFE BARNES PIKE

Associate Editors

Vol. 64 April-June, 1962 No. 758
CONTENTS:
Comment from the New Editor. A. R. Hodgdon .................... 97
Some Additions to the Flora of Grand Manan Island, New
Brunswick, A. R. Hodgdon and Radcliffe B. Pike ............ 98
Cytological Observations of Polygala in Eastern North
America. Walter H. Lewis and Shirley A. Davis ............ 102
Synthesis of Aster Herveyi. Leonard J. Uttal .................. ei RLS
Three Misidentified So-Called Cordilleran Species in Eastern
North America. Muf Dahl... erase erii EE TM
Additions to the Bryophyte Flora of Keweenaw County,
DEMNM. P. DEGPPIAOM. ...0500.0c0cccosnnsssseabsanusiesesaseninn nese RS 121
Contribution to the Fungus Flora of Northeastern North
America. Howard E. Bigelow and Margaret E. Barr ........ 126

(Continued on Inside Cover)

Che Nem England Botanical Club, Inc.

Botanical Museum, Oxford St., Cambridge 38, Mass.

CONTENTS: — continued
The Noseburn (Tragia, Euphorbiaceae) of Western Texas.

Morahodt OM ohneton 5. 34... dipsa chives peers oer eie eie nee 137
A New Species of Haplopappus, Section Blepharodon.
Zay CI DEBERE. coepere vcevseeseransmeasinngsaboevestines tbe RR erint 142
Agropyron Hybrids and the Status of Agropyron Pseudo-
repens. MONAT WA Pohl 21222 staa iG E eset ss 143

Chromosome Numbers for Some Angiosperms of the South-
ern United States and Mexico. Walter H. Lewis,

H. Larry Stripling and Richard G. Ross ................ eee 147
Glandularity in Rubus Allegheniensis Porter.
A. R. Hodgdon and Frederic Steele ........................... cesses 160

Fifteenth Report of the Committee on Plant Distribution ...... 168
Occurrence of Species of Polycarpaea Lam. (Caryophylla-

ceae) in North America. Olga Lakela ..................... esee 179
A New Station for Pinguicula Vulgaris in Ontario.

Nd OD LAB S N AEE N a rotat lidiqatersn iden caido E noia 182
Hyptis Radiata (Labiatae) an Illegitimate Name.

Lloyd H, SRRQUE LL iki moescnsesssccevincsémparsdiseunizssssinecintendeaealbdlbedews 184
Makers of North American Botany. Joseph Ewan ............... 186

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Rhodora

JOURNAL OF THE
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Vol. 64 April-June, 1962 No. 758

COMMENT FROM THE NEW EDITOR

It should be pointed out that the January-March 1962
number of Rhodora was edited entirely by Professor Reed
Rollins, who volunteered to continue his services during the
critical period of transition. This seems to have been a
wise solution, because much of the editing of this number
had been accomplished when the change in editors was made.
I much appreciate Professor Rollins’ willingness to help out
in this way to bridge the gap.

It is particularly reassuring to me to have nearly all of the
experienced editorial board continuing to serve, for much
of the quality of the journal is attributable to their critical
attention and judgment.

Since Rhodora, for the first time, is being edited at some
distance from Cambridge, it was felt that the selection of a
new associate editor from my colleagues at the University
of New Hampshire would do much to facilitate my work.
Mr. Radcliffe Pike, who has worked with me for many
years in botanical investigations, has consented to serve on
the board.

Professor Rollins, in my opinion, during his twelve years
as editor has very wisely broadened Rhodora, both as to
geographical coverage and the range of subject matter. But
because of this very broadening, the Gray’s Manual Area
and New England should continue to provide abundant
material for new insights and interpretations. It is hoped
that botanists interested in the more local area will have a
resurgence of activity in the years ahead. — A. R. HODGDON

97

98 Rhodora [Vol. 64

SOME ADDITIONS TO THE FLORA OF
GRAND MANAN ISLAND, NEW BRUNSWICK

A. R. HODGDON AND RADCLIFFE B. PIKE

The Vascular Flora of Grand Manan was excellently pre-
sented by Weatherby and Adams in 1945 (1) and has served
as an admirable source of reliable information relating to
our current floristic study of the nearby Wolf Islands and
other islands in the Bay of Fundy.*

In connection with our work a number of questions had
arisen that bore directly on Grand Manan, so in July 1961
we visited Grand Manan and Long Island for parts of two
days and in August returned for three days of active bot-
anizing on the main island as well as on Outer Wood and
White Head Islands. One of our main projects was to
determine whether a considerable number of taxa that we
had found on the Wolf Islands but which Weatherby and
Adams had not reported from Grand Manan, actually were
absent there. This turned out to be true so far as our
observations and collections were concerned, thus confirming
our opinions that a fairly thorough canvass had been made
of the Grand Manan flora. However, as a by-product of
our work, a number of new records were compiled as well
as one somewhat questionable record confirmed. These
additions will now be considered along with a few problems
that still invite further study.

Lycopodium tristachyum Pursh.-—On the evening of
August 21 while on our successful search for Pinus resinosa,
we encountered a dry somewhat sandy area with a sparse
growth of Vaccinium myrtilloides, Kalmia angustifolia,
Rhododendron canadense and associated species. Here we
found a few individuals of this clubmoss with its charac-
teristically deeply embedded rhizomes. Though lacking
strobili there seems to be no question about its identity.

*This research is part of a project entitled ‘‘Floristic and Phytogeographie Investiga-
tions of the Wolf Islands and other islands in, the Bay of Fundy which was supported
by a grant from the Central University Research Fund of the Graduate School of the
University of New Hampshire.

Published with the approval of the Director of the New Hampshire Agricultural
Experiment Station as Scientific Contribution No. 285.

1962] Hodgdon and Pike — Grand Manan Flora 99

Specimens of this and other taxa listed here as additions
to the flora of Grand Manan, unless especially noted, are
deposited in the Gray Herbarium and in the Herbarium of
the University of New Hampshire.

Tsuga canadensis (L.) Carr. — Hemlock was included in
the “Flora” on the strength of Allan Moses’ report of a
single tree in the ravine of Dock Brook. There is little
reason to doubt this record but we were unable to find any
hemlock trees though we scrutinized Dock Brook and its
environs carefully except for a short lower section. It is
perhaps more surprising that hemlocks are not more preva-
lent on Grand Manan. It is of some significance certainly
that, during our search for Red Pine, on August 21 we
collected a branch of Rhododendron canadense with its
leaves conspicuously infected with a rust which has been
identified by Dr. Avery Rich, Plant Pathologist at the
University of New Hampshire as Pucciniastrum myrtilli
(Schum.) Arth. The alternate host of this rust is Tsuga
canadensis. Presumably further search for hemlock should
be carried on in this part of the island. This specimen of
rust is in the mycological herbarium at the University of
New Hampshire.

Pinus resinosa Ait. — Weatherby and Adams loc. cit. p.
77 placed this taxon in their list of “Doubtful and Unveri-
fied Records" attributing it to Perkin's list. They stated of
this record as follows ‘‘very likely correct, though the soils
of Grand Manan are not such as the red pine prefers."

We first heard of the possible occurrence of red pine in
July from Mrs. Earl Green of Grand Harbour. She told
us that her father a Mr. Cheney knew of some “different
pines" and that she could find someone to direct us to
specimens of them. We much appreciate her assistance in
providing us with the services of Terrance Ingalls, a very
energetic and intelligent boy from Grand Harbour, who on
August 21, led us to a number of fine trees of Pinus resinosa
a short way inland from Grand Harbour. There were sev-
eral scattered trees of good size varying from one to two
feet dbh. Seedlings were observed in the vicinity of the

100 Rhodora [Vol. 64

larger trees but no intermediate small specimens were
noted. These pines were scattered over a sufficiently wide
area to lead one to hope that some of them may continue
to survive there for many years to come.

Eleocharis tenuis (Willd.) Schultes var. tenuis. To-
ward the end of our unsuccessful search for hemlocks along
Dock Brook we didn't want to return empty handed so our
attention wandered to other things. In a moist opening along
a wood-road we saw a fine growth of Eleocharis growing
with Juncus articulatus. After very careful examination
our specimens, which are in good fruiting condition, prove
to be E. tenuis var. tenuis. E. elliptica Kunth resembles E.
tenuis in some of its vegetative characteristics and in its
general appearance and it has been collected on Grand
Manan. The achenes of the two species are very distinct,
those of E. elliptica being pronouncedly yellow while those
of E. tenuis are olivaceous and smaller. The tubercule also
is less compressed in E. tenuis. We compared our material
carefully with the specimen of E. elliptica from Grand
Manan and with reliable specimens of both E. elliptica and
E. tenuis in the New England Botanical Club Herbarium.

Betula papyrifera Marsh var. cordifolia (Regel) Fernald.
— Weatherby and Adams loc. cit. p. 41 listed no variety of
this species nor made any comment about its variability on
Grand Manan. Fernald's treatment of the series Albae in
eastern North America (2) appeared at about the same
time that the Grand Manan Flora first came out in the
autumn of 1945. For this reason Mr. Weatherby may not
have been stimulated to study birches very closely during
the years preceding this when he was actively investigating
the flora. Most of the Betula papyrifera of Grand Manan
is whitish barked but in 1955 the junior author noted a
birch of this variety with dark bark and abundant catkins
along the road to the Whistle House. At that time the area
was densely wooded.

On August 23, 1961 we revisited this locality, which had
been lumbered for the conifers in the interim, finding many
of the birches in question with abundant catkins, dark

1962] Hodgdon and Pike — Grand Manan Flora 101

bark and leaves mostly cordate-based. We took specimens
from ten trees which we hope to study later along with
similar population samples which we have from the Wolf
Islands and Lubec, Maine.

Thlaspi arvense L. — A specimen in fruit with no leaves
was collected by us at Dark Harbour on August 22, 1961. We
have noticed in examining the Weatherby and Adams list
that several very common and widespread weeds and intro-
duced plants have not yet been reported. This and the taxon
to follow will help to augment the list of introductions.
Specimens of these are to be found in the herbarium at the
University of New Hampshire.

Raphanus raphanistrum L. forma sulphureus (Babey)
Hayek. — Since Weatherby and Adams often reported con-
spicuous forms in other groups it is appropriate to record
this one which we collected at Dark Harbour.

Rubus canadensis L. — Weatherby and Adams loc. cit.
p. 52 state that R. canadensis L. is “the common high-bush
blackberry of the island.” In August 1955 the junior author
and Dr. Donald L. Craig of Kentville, Nova Scotia (3), who
then was working with R. canadensis at the University of
New Hampshire, spent several days on Grand Manan
searching for good material of that species. Following the
interpretation of Bailey (4) and that of Fernald (5) of
R. canadensis, good material was conceived of as being
essentially without prickles, comparable in this respect to
the usual specimens of R. canadensis that are so common in
northern New England and in cooler mountainous areas
farther to the south. Surprisingly a very small percentage
of the specimens seen in the summer of 1955 qualified as
typical Rubus canadensis. Most of the plants were of vary-
ing degrees of prickliness, in this respect fitting either R.
amicalis Blanch. or R. elegantulus Blanch. as these are in-
terpreted by Fernald (5). But it is to be noted that
Weatherby and Adams, loc. cit. p. 53 relied on Bailey for
the identification of their blackberries. Bailey’s treatment
of R. canadensis in 1944 was a peculiar one particularly in
the light of his usual tendency to split the species as finely

102 Rhodora [Vol. 64

as possible. After a preliminary delineation of R. canadensis
as an essentially smooth blackberry, he combined R. elegan-
tulus with it as only “a small and usually slender phasis”
(see Bailey loc. cit. p. 475). The Grand Manan plants
present a particular problem because they combine the
prickliness of R. elegantulus and R. amicalis with the
features that are commonly associated with F. cana-
densis including large size and strongly racemose inflores-
cences. Except for the degree of prickliness, most of the
bigger plants certainly resemble R. canadensis.

— DEPARTMENT OF BOTANY AND DEPARTMENT OF HORTICUL-
TURE, UNIVERSITY OF NEW HAMPSHIRE, DURHAM, NEW HAMP-
SHIRE

LITERATURE CITED

1. WEATHERBY, C. A. AND JoHN ADAMS. 1945. A list of the Vascular
Plants of Grand Manan, Charlotte County, New Brunswick, Con-
trib Gray Herb. CLVIII.

2. FERNALD, M. L. 1945. Notes on Betula in eastern North America.
Rhodora 47: 303-329.

3. CRAIG, DONALD L. 1959. The Cytology and Breeding Behavior of

Rubus canadensis L., Thesis, University of New Hampshire.

BAILEY, L. H. 1944. Gent. Herb. Vol. V Fasc. VII pp. 465-476.

. FERNALD, M. L. 1950. Gray's Manual of Botany. Eighth ed. p. 832.

-aa

ct

CYTOLOGICAL OBSERVATIONS OF POLYGALA IN
EASTERN NORTH AMERICA

WALTER H. LEWIS AND SHIRLEY A. DAVIS!

For a genus of over 450 species, of which no fewer than
200 are described in North and Central America, only 15
species of Polygala have been studied cytologically. No
New World species are included in this meager total.

Unfortunately, the Polygala material is not particularly
favorable for study. At meiosis the greatest difficulty is
the low number of pollen mother cells formed in each
anther, giving few figures for observation when a satis-
factory stage is obtained. Under these circumstances, we
have found that much more fixed bud material must be

"Undergraduate Research Participant, National Science Foundation G-12059.

1962] Lewis and Davis — Polygala 103

collected for results than is usual for most genera. When
to this need is added the paucity of plants at any one locality,
particularly for those species endemic to the Mexican high-
lands where the greatest number of North American species
occur, it is apparent why this feature limits chromosomal
research. The meiotic process also is a hindrance. Although
prophase I appears to be of unusually long duration, we
agree with Glendinning (1960) that diakinesis is deceptive
in that some chromosomes are still very poorly stained. The
most satisfactory stages between prometaphase I and telo-
phase I, however, are completed in very short periods of
time which markedly decrease the chance of locating satis-
factory figures. Complete restitution nuclei are formed only
to be followed in rapid sequence by the phases of meiosis II.
Finally the chromosomes of most species are very small and
high in number.

A list of reported chromosome numbers in Polygala has
been compiled (Table 1). Five of the 15 species have been
examined by two or more authors, but except for P. myrti-
folia L. none of these has been given the same chromosome
number. This is perhaps the best illustration of the techni-
cal difficulties to be encountered in Polygala. The 3 African
species listed have a basic number of x — 19 which are
according to Larsen (1959) secondary diploid species prob-
ably of tribasic origin from 6 + 6 + 7 stocks. The single
Asian species, P. japonica Houtt., has n — 21 chromosomes
and may be a hexaploid species with a base of x — 7. As
many as 75% of the European species counted have basic
numbers of x — 17. As noted in Table 1, the reports are
not unanimous for species considered in this series, but
the numerous counts from several localities made by Glen-
dinning (1960) outweigh the evidence of earlier authors.
It cannot be overruled, however, that chromosmal races may
exist for a single species, so that P. amara L. may have an
^ — 14 race in eastern Europe (Baskay, 1956; Skalifiska
et al, 1959) and an « — 17 race in western Europe
(Glendinning, 1960). The remaining European species have
basic numbers of x — 7, 8, and 23 (Table 1).

Of other genera in the Polygalaceae records exist for only

104 Rhodora [Vol. 64

4 species, one each in Atroxima, Bredemeyera, Carpalobia,
and Securidaca. These are A. liberica Stapf., 2n = 18
(Mangenot and Mangenot, 1957) B. colletioides Phil., 2n =
14 (Covas and Schnack, 1946), C. lutea G. Don, 2n = 20
(Mangenot and Mangenot, 1957), and S. longipedunculata
Fres., 2n = 32 (Mangenot and Mangenot, 1958; Miege,
1960).
MATERIALS AND METHODS

From 39 collections of immature flower buds made from
localities in the eastern United States and Mexico, chromo-
some numbers for 24 species of Polygala are reported. The
buds were fixed in the field in modified Carnoy’s solution
(4: 3: 1) and shortly thereafter were stored in the refrig-
erator for periods up to 6 months before squashing in 1%
acetic-orcein. The results are mostly from the study of
pollen mother cells or more rarely from premeiotic mitosis.
The chromosome number reported for each species is when
possible based on the study of more than one plant from
several localities. Voucher specimens were obtained for
each collection. A complete set has been filed at the Southern
Methodist University herbarium (SMU), except for a few
which are deposited elsewhere as indicated in Table 2.
Duplicates of most collections are filed at the University of
Texas herbarium (TEX). We wish to thank the curators
of these herbaria for accepting specimens for permanent
accession and we especially appreciate receiving fixed mate-
rial of 3 species from Dr. Robert Kral, Virginia Polytechnic
Institute, Blacksburg.

RESULTS

The gametic chromosome numbers for 24 species of
Polygala are listed by sections (Blake, 1924) in Table 2.
Most of the data are based on distinct figures giving defi-
nite counts, although the numbers for a few species are
only approximations. With rare exceptions (P. praetervisa
Chod., fig. 11), meiotic behavior was regular. The chromo-
somes of all species were found to be generally small with
the smallest in the sections Hebeclada (fig. 5), Monninopsis
(fig. 15), and Timutua (fig. 3, 10, 11, 14), and the largest

1962] Lewis and Davis — Polygala 105

in the section Timutua (fig. 2, 7-9). Chromosomes of indi-
vidual karyotypes often varied from very short to inter-
mediate in length as shown by the mitotic chromosomes of
P. reducta Blake (fig. 13).

The most striking result was the diversity of chromosome
numbers for such a small fraction of species. Numbers
ranging from 2n — 16 for P. scoparia HBK. to 2n — ca. 104
(108) for an undescribed Mexican species related to P. alba
Nutt. represent both the lowest and the highest numbers
known in the genus. Basic numbers were equally as variable.
The species are separable into a primary aneuploid series
with basic numbers of x — 6(12), 7(14), 8, and 10, and a
secondary basic series of x — 15, 17, and 23. Probably the
closely related P. cruciata L. and P. chapmanii T. & G. with
^ — 18 and 36, respectively, represent species in still an-
other series of x — 9, but until further research reveals the
existence of species with 9 haploid chromosomes or until
some other evidence is presented, all species having chro-
mosome multiples of 6 are grouped in the x = 6(12) line.

Common to species in the sections Monninopsis and
Timutua, the basic number of x — 17 includes about 30%
of the species studied. The tetraploid species are slightly
more common than the diploids. The basic number of x = 6
was also shared by species in these sections. Species with
x = T are distributed in 3 sections, Eurhinotropis, Hebe-
clada, and Timutua, which illustrates a greater degree of
morphological diversity than do species of other basic series
in eastern North America. The x — 8 and 10 lines are repre-
sented by one species each in the sections Monninopsis and
Timutua.

Since at least 4 primary basic numbers are reported for
Polygala, the secondary dibasic and tribasic lines may have
arisen in a number of ways. The x — 15 group, common to
2 species in the section Microthrix, could have formed from
species with » — 7 and 8 or, providing stronger evidence
is found for the existence of an x — 9 series, 6 and 9 chro-
mosomes. In view of the chromosomal diversity recorded
by this small sample, it is not inconceivable that these species
are unibasic polyploids at the 6x and 12z levels of an x = 5

106 Rhodora [Vol. 64

series. There is less doubt that the x — 17 series is of
dibasie origin and it may have formed by hybridization of
species with 7 and 10 or 8 and 9 haploid chromosomes. The
single species with x = 23, P. nuttallii T. & G., is most prob-
ably of tribasic origin and numerous combinations exist in
species of the same section to give the necessary chromo-
somal complement. Without experimental evidence and
with only limited counts in the genus as a whole, it is far
too speculative at the present time to suggest definite
origins for the secondary lines.

The levels and frequencies of ploidy exhibited by the 24
North American species are summarized as follows: pri-
mary diploids, 5% ; secondary diploids, 21% ; primary poly-
ploids, 5396; and secondary polyploids, 21%. Since the
secondary diploids are probably all amphidiploids, this gives
a total polyploid frequency of 95% or a markedly higher
one than the average of 30-35% estimated for the angio-
sperms in general (Stebbins, 1938). These data are tabu-
lated in Table 3 by basie chromosome number, except that
species no. 4 (Table 2) is omitted.

DISCUSSION

Although the total number of species studied is only 39,
it is sufficiently large to show the importance of polyploidy
in addition to hybridization and aneuploidy in the phylogeny
of Polygala. The numerical modifications have been so
extensive, however, that they have obscured the original
chromosome number of the genus. The most frequent num-
ber, x = 17, can be derived from several primary lines and
gives no direct evidence of the composition of the original
complement. The numbers of x — 7, 8, 9, and 10, in four

Figures 1-16. Chromosomes of Polygala, drawn with the aid of a camera lucida at
X 2300 reduced by ca. 1/3 in reproduction. Fig. 1. P. alba, n — 12, Lewis 5537; fig.
2. P. chapmanii, n — 36, Lewis 5690; fig. 3. P. cruciata, n — 18, Lewis 5692; fig. 4. P.
curtissii, n —20 (1 side of metaphase II), Kral 13817; fig. 5. P. grandiflora, n = 14,
Lewis 5680; fig. 6. P. scoparia, 2n = 16, Lewis 5753; fig. T. P. mariana, n = 17, Lewis
5625; fig. 8. P. nana, n = 34, Lewis 5654; fig. 9. P. nuttallii, n = 23, Kral 13791; fig.
10. P. polygama var. obtusata, m — 28, Lewis 5621; fig. 11. P. praetervisa, n = 48
(as 46u-+1m1+11), Lewis 5667; fig. 12. P. ramosa, n = 34 (1 side of anaphase I),
Lewis 5691; fig. 13. P. reducta, 2n = 30, Lewis 5757; fig. 14. P. rugelii, n = 34, Lewis
5681; fig. 15. P. scoparioides, n = 17, Lewis 5545; fig. 16. P. verticillata var. isocycla,
n = 17 (1 side of metaphase II), Lewis 5628.

107

Lewis and Davis — Polygala

1962]

108 Rhodora [Vol. 64

other polygalaceous genera are too limited to suggest a
trend. Even though the series with x — 7 includes just 7
species, they are native to Asia, Europe, and North America,
to give in total the most extensive distribution of any series.
In addition, these species are classified in 5 sections and
represent the most diverse morphology of any series. Fur-
ther research may suggest a different conclusion, but the
evidence now available supports « = 7 as the original basic
number for Polygala and probably for the family. The
origin of the x — 6 line can be postulated on the basis of
chromosomal loss and the x — 8, 9(?), and 10 groups by
chromosomal gain from 7.

A suggestion of the antiquity of the primary aneuploid
lines is found in the widespread occurrence of the x = 17
stock in the modern flora. A majority of the European and
southeastern North American species now known have this
number in common. It is reasonable to speculate that the
species with x = 17 had a single origin and that migration
around the North Atlantic, when it was effectively possible
in the early Tertiary (Axelrod, 1960), established the
group in both continents. Alternately, the number could
have arisen independently, but the chance of forming a
derived stock twice with both giving rise to parallel lines
of descent of about equal significance and success in two
continents seems much less probable. If the x = 17 series
had formed in either Europe or North America by the early
Tertiary, then the basic aneuploid differentiation must have
occurred in the Cretaceous or earlier. The great antiquity
of the primary series may account to some extent for the
loss of most of the diploid species (1 out of 39 species) and
for the high level of polyploidy reached by the x = 6 (162)
and 7 (12a) groups. In the secondary aneuploid lines, un-
questionably of more recent origin, about one-half of the
North American species are diploids and all polyploids are
only 4x. — DEPARTMENT OF BIOLOGY, STEPHEN F. AUSTIN
STATE COLLEGE, NACOGDOCHES, TEXAS.

LITERATURE CITED
AxELROD, D. I. 1960. The evolution of flowering plants. Evolution

1962] Lewis and Davis — Polygala 109

After Darwin. Vol. 1, Evolution of Life. The University of
Chicago Press.

Baksay, L. 1956. Cytotaxonomical studies on the flora of Hungary.
Budapest. Magyor Nem Muz. Evkàn 7: 321-334.

BLAKE, S. F. 1924. Polygala, in N. Am. Fl. 25: 305-370.

CONTANDRIOPOULOS, J. 1957. Nouvelle contribution a l'étude caryolo-
gique des endemiques de la Corse. Bull. Soc. Bot. Fr. 104: 533-538.

Covas, G., and B. SCHNACK. 1946. Numero de cromosomas en antó-
fitas de la región de Cuyo (Republica Argentina). Rev. Argent.
Agron. 13: 153-166.

GLENDINNING, D. R. 1955. La cytologie de Polygala chamaeburus L.
Bull. Soc. Neuchatel. Sci. Nat. 78: 161-167.

1960. Cytology of Polygala. Nature 188: 604-

605.

HAGERUP, O. 1932. Uber Polyploidie in Beziehung zu Klima,
Okologie und Phylogenie Chromosomenzahlen aus Timbuktu.
Hereditas 16: 19-40.

LARSEN, K. 1956. Chromosome studies in some Mediterranean and
South European flowering plants. Bot. Not. 109: 293-307.

1959. On the cytological pattern of the genus Polygala.
Bot. Not. 112: 369-371.

LóvE, A., and D. LóvE. 1944. Cyto-taxonomical studies on boreal
plants. III. Some new chromosome numbers of Scandinavian
plants. Ark. Bot. 31A: 1-22.

MANGENOT, S., and G. MANGENOT. 1957. Nombres chromosomiques
nouveaux chez diverses dicotylédones et monocotylédones d'Afrique
occidentale. Bull. Jard. Bot. Bruxelles 27: 639-654.

1958. Deuxième liste de nombres
chromosomiques nouveaux chez diverses dicotylédones et monoco-
tylédones d'Afrique occidentale. Bull. Jard. Bot. Bruxelles 28:
315-329.

MATTICK, T. 1950. in G. Tischler, Die Chromosomenzahlen der
Gefasspflanzen Mitteleuropas. 'S-Gravenhage, The Netherlands.

MiEGE, J. 1960. Nombres chromosomiques de plantes d'Afrique
occidentale. Rev. Cyt. Biol. Vég. 21: 373-384.

SKALINSKA, M., R. CzaPik, M. PioTrROWICZ, et al. 1959. Further
studies in chromosome numbers of Polish angiosperms (dicoty-
ledons). Act. Soc. Bot. Polon. 28: 487-529.

STEBBINS, G. L. 1938. Cytological characteristics associated with the
different growth habits in the dicotyledons. Am. Jour. Bot. 25:
189-198.

SUZUKA, O. 1950. Chromosome numbers in pharmaceutical plants. I.
Rep. Kihara Inst. Biol. Res. 4:57-58.

WurrrF, H. D. 1938. Chromoscmenstudien an der Schleswig-Hol-
steinischen Angiospermen-Flora II. Ber. Deutsch. Bot. Ges. 56:
247-254.

110 Rhodora [Vol. 64

Table 1. Chromosome numbers reported for the Old World Polygala.

Taxon n 2n Author
AFRICA
P. myrtifolia L. 19 is Larsen (1959)
P. myrtifolia L. e. ca. 38 Glendinning (1960)
P. triflora L. 19 "x Hagerup (1932)
P. virgata Thunb. var.

speciosa (Sims) Chod. D. 38 Larsen (1959)
ASIA
P. japonica Houtt. D. 42 Suzuka (1950)
EUROPE
P. alpestris Rchb. 17 ca. 34 Glendinning (1960)
P. alpina Perr. & Song. + ca. 34 Glendinning (1960)
P. amara L. 14 EM Baksay (1956)
P. amara L. subsp.

amarella Cr. 17 34 Glendinning (1960)
P. amara L. subsp.

brachyptera (Chod.) Hay. . . 28 Skalinska et al. (1959)
P. calcarea Schultz 17 34 Glendinning (1960)
P. chamaebuxus L. e. 38 Mattick (1950)
P. chamaebuxus L. 22,23,24 .. Glendinning (1955)
P. chamaebuxus L. e. ca. 46 Glendinning (1960)
P. comosa Schkr. M 28-32 Mattick (1950)
P. comosa Schkr. e. 28 Larsen (1956)
P. comosa Schkr. 17 34 Glendinning (1960)
P. major Jacq. e. 32 Mattick (1950)
P. nicaensis Risso subsp.

corsica, Graebn. s 34 Contandriopoulos (1957)
P. serpyllifolia Hose. 17 34 Glendinning (1960)
P. vayredae Costa ss 28 Glendinning (1960)
P. vulgaris L. 24-28 Js Wulff (1938)
P. vulgaris L. D. ca. 70 Live & Love (1944)
P. vulgaris L. . . 28,32, ca. 56 Mattick (1950)
P. vulgaris L. 34 68 Glendinning (1960)

Table 2. Chromosome numbers for 24 North American species of

Polygala.
Taxon Gametic
Number (n) Voucher
Section Eurhinotropis
P. tweedyi Britt. ca. 14 TEXAS. San Patricio Co., 2.1 miles NW
of Mathis, Lewis & Jones 5583 (1).°
Section Hebeclada

P. grandiflora Walt. 14 FLORIDA. Glades Co., 8.6 miles SE of
Palmdale, Lewis 5680 (3).

1962] Lewis and Davis — Polygala 111

Taxon Gametic
Number (n)

Section Microthrix

P. ovatifolia Gray 28-30"

P. reducta Blake
P. scoparia HBK.

P. scoparioides Chod,

P. sp. (Ll)

Section Timutua
P. alba Nutt.

P. boykinii Nutt. ca.

P. chapmanii T. & G.

P. cruciata L.

P. curtissii Gray

P. lutea L.

P. mariana Mill.

P. nana (Michx.) DC.

15*

8*

17

12"

12

14*

36

18

20

94

Tt

94

Voucher

TEXAS. San Patricio Co., 2.1 miles NW
of Mathis, Lewis & Jones 5586 (1).
NUEVO LEON. 15.2 miles E of Hwys. 57
and 60 junction, Lewis 5757 (2).
NUEVO LEON. 11 miles E of Hwys. 57
and 60 junction, Lewis 5753 (4).
TEXAS. Brewster Co. Big Bend Na-
tional Park, Chisos Mts., Lewis 5545
(1); Panther Junction, Lewis 5466
(1).

NUEVO LEON. 9.9 miles E of Hwys. 57
and 60 junction, Lewis 5746 (1, US).

TEXAS. Brewster Co., Big Bend Na-
tional Park, Lewis & Oliver 5455 (2);
Culberson Co., North McKittrick Can-
yon, Lewis 5537 (2).

FLORIDA. Dade Co., Homestead, Lewis
$676 (1).

MISSISSIPPI. Hancock Co., 0.5 miles W
of St. Louis Bay & Hwy. 90, Lewis
5690 (2).

MISSISSIPPI. Hancock Co., 6.2 miles
ENE of Pearl River and Hwy. 90,
Lewis 5692 (3); viRGINIA. Brunswick
Co. 10.5 miles WNW of Emporia,
Kral 13633 (2, VPI).

VIRGINIA. Southampton Co., 8 miles E
of Emporia, Kral 13817 (2, VPI).
MISSISSIPPI. Hancock Co., 0.5 miles W
of St. Louis Bay and Hwy. 90, Lewis
5689 (1).

TEXAS. Hardin Co., Kountze Lookout
Tower, Lewis 5629 (2); Jasper Co., 50
yds. S of Hwys. 96 and 1004 junction,
Lewis 5625 (1); Polk Co., 3 miles SW
of Barnum, Lewis 5633 (2); San Au-
gustine Co., 0.7 miles SW of Boykin
Spring entrance and Hwy. 68, Lewis
& Oliver 5511 (1).

FLORIDA. Citris Co., 0.4 miles N of
Citris Co.-Hernando Co. line and Hwy.
19, Lewis 5654 (2); TEXAS. Jasper Co.,

112 Rhodora [Vol. 64

Taxon Gametic Voucher
Number (n)
50 yds. S of Hwys. 96 and 1004 junc-
tion, Lewis 5623 (2) ; Newton Co., 4.7
miles S of Newton, Lewis 5619.

P. nuttallii T. & G. 23 VIRGINIA. Nansemond Co., N of Cle-
opus, Kral 13791 (1, VPI).
P. polygama Walt. 28 TENNESSEE. Polk Co. 1 mile N of
var. polygama Hwys. 64 and 30 junction, Lewis 5640
(1).
P. polygama Walt. 28 GEORGIA. Henry Co. 10 miles N of
var. obtusata Chod. Griffen, Lewis 3648 (3); TEXAS. Har-

din Co., 1.4 miles SE of Votaw, Lewis
5631 (2); Newton Co., 2.5 miles N of
Burkeville, Lewis 5616 (1), 2.8 miles
S of Newton, Lewis 5621 (1).

P. praetervisa Chod. 48 FLORIDA. Monroe Co., Big Pine Key,
Lewis 5667 (1).
P. ramosa Ell. 34 MISSISSIPPI. Hancock Co., 6.2 miles

ENE of Pearl River and Hwy. 90,
Lewis 5691 (4).

P. rugelii Shuttlw. 34 FLORIDA. Glades Co., 8.6 miles SE of
Palmdale, Lewis 5681 (1).

P. verticillata L. 17 TEXAS. Hardin Co., Kountze Lookout

var. isocycla Fern. Tower, Lewis 5628 (3); Nocogdoches

Co., 0.6 miles E of Martinsville, Lewis
5723 (1); Panola Co., 2.3 miles NE of
Pinehill, Lewis 5720 (2).

P. sp. (2)” ca. 42" NUEVO LEON. 11 miles E of Hwys. 57
and 60 junction, Lewis 5754 (1, US).

P. sp. (83)* ca. 36 OAXACA. N of Huaguapan de Leon,
Oliver 136 (1, US).

P. sp. (4)" 52-54" COAHUILA. 3.1 miles N of Los Llanos,
Lewis 5728 (2, US).

P. sp. (4) ca. 52° NUEVO LEON. 15.2 miles E of Hwys. 57

and 60 junction, Lewis 5755 (1, US).

aBased on the somatic number of premeiotic cells.
"Probably represent new species to be discussed in a subsequent paper.
*Number of plants examined for the chromosome number.

Table 3. Basic chromosome numbers and levels of ploidy with numbers
of species by sections for 23 North American species of Polygala.

Basic Ploidy Number of Section
number species
x=6 Ax 2 Monninopsis, Timutua

6x 1 Timutua

1962] Uttal — Aster Herveyi 113

12x 2 Timutua
16x 1 Timutua
x= 4a 3 Eurhinotropis, Hebeclada, Timutua
8x 1 Timutua
12x 1 Timutua
2-8 2x 1 Monninopsis
x = 10 4x 1 Timutua
x = 15 2x 1 Microthrix
Ax 1 Microthrix
2=17 2x 8 Monninopsis, Timutua
4x 4 Timutua
x = 23 2x 1 Timutua

SYNTHESIS OF ASTER HERVEYI
LEONARD J. UTTAL

Long suspected to be a hybrid between Aster spectabilis
Ait. and A. macrophyllus L.1, A. Herveyi Gray can now be
reported as so in fact. The putative parents have been
artificially crossed and resulting progeny compare satisfac-
torily with naturally occurring A. Herveyi. Subsequent
reference to this plant will be as A. X Herveyi Gray.

A. spectabilis is a coastal plain species from Massachu-
setts to South Carolina?*. A. macrophyllus is a species of the
eastern North American upland. The two are allopatric
except in a few counties of southeastern New England and
eastern Long Island. Here the two ranges impinge ecologi-
cally and physiographicall, and here occur the hybrid
swarms of A. X Herveyi. A specimen from Plainfield, New
Jersey collected in 1909, now in the New York Botanical
Garden Herbarium, indicates that sympatry between the
parent species existed in parts of the New York metropoli-
tan area before its development.

A. spectabilis and A. macrophyllus are both present in the

1Fernald, M. L. 1950. Gray’s Manual of Botany, 8th. Ed.: 1430.
?Southern limit based on information from North Carolina herbaria. (Harry E.

Ahles, pers. com.)

114 Rhodora [Vol. 64

vicinity of Sag Harbor, Suffolk County, Long Island. A. X
Herveyi occurs here in colonies among which there is a
central clone of robust plants surrounded by smaller clones
of plants of diminishing stature. Apparently introgressive
clones occur consisting of plants intermediate between the
hybrid and either of the parents. These can not be assigned
satisfactorily to any category involved. Marked variation
in vegetative vigor is evidenced by different clones in sim-
ilar sites. The hybrid swarms of Long Island are similar to
those on the mainland as reported by Hervey*.

The intermediate nature of the hybrid is most graphically
manifested by the basal leaves and the similar ones of the
characteristic vegetative tufts. In A. macrophyllus, these
leaves are quite large, ovate, serrate or crenate, cordate, and
long-petioled. In A. spectabilis, comparative leaves are
much smaller, lanceolate to narrowly ovate or obovate, entire
or remotely serrate, the blade tissue gradually tapering to
the short petiole. In A. X Herveyi, these leaves are vari-
ously intermediate in size, broad to narrowly ovate, serrate
or crenate, with the blade rounded to the petiole. The blade
is sometimes subcordate or cordate, in which case, blades
with rounded bases are prevalent on neighboring plants.

Characters of the A. macrophyllus parent seem to pre-
dominate in the hybrid: leaf serration, naked petioles,
usually appressed involucral bracts. Perhaps for this reason,
extremes of A. macrophyllus have, on occasion, been called
A. Herveyi.

Sometimes the squarrosity of the involucral bracts of
A. spectabilis comes through slightly in hybrids. In such
cases, it tends to be accompanied by the deep purple-blue
and long rays characteristic of A. spectabilis.

The A. spectabilis parent used in the artificial crossing
experiment originated in Riverhead, Suffolk County, Long
Island, New York. The A. macrophyllus parent, of the typi-
cal variety, originated from Main Top Mountain, Amherst
County, Virginia. Crossings were made in my garden in
Madison Heights, Virginia. The A. spectabilis was planted

*?Hervey, E. W. 1916. Rhodora 18: 183-184.

1962] Uttal — Aster Herveyi 115

IL

PLATE 1267. Representative leaves from the vegetative tufts of synthetic A. X
Herveyi. The range of naturally occurring variation is artificially duplicated.

116 Rhodora [Vol. 64

in the ground while the A. macrophyllus was kept in a pot
for mobility. Both were kept in individual cheesecloth tents
to prevent entomophilous contamination of heads.

Working on the presumption that both putative parents
are self-incompatible (other Aster species tested are’, and
populations of parent species are characteristic of out-
breeders), flower heads of both were pressed together to
effect pollination. These heads produced abundant plump
achenes. Certain heads not pollinated and left to themselves
produced only shriveled achenes.

Ripe achenes were sown in flats kept outdoors over winter.
Sprouting was sparse in late fall; abundant the following
April. Eight F, plants bloomed the first fall (1961) and
were made into herbarium sheets. Seven of these, with a
sheet each of the parents, were sent to the Gray Herbarium
for preservation. An eighth F, sheet was sent to the New
York Botanical Garden Herbarium. Many additional F,
plants, vegetative in 1961, are expected to bloom in 1962.
Specimens of these can be supplied upon request.

The high degree of variation prevailing in progeny
resulting from multiple crossings of the same parents
indicates highly complicated parental genetic makeup. This
is in keeping with the extensive variability of the parents,
especially of A. macrophyllus.

Tests of F, seed will have to wait another season. Obser-
vations from the field indicate F, fertility is impaired.

For a genus with so many sympatric outbreeding species,
in an era of so much disturbed habitat, Aster has a remark-
able paucity of described hybrids. This is an area meriting
closer inspection; perhaps it is the key to better under-
standing of certain “difficult” taxa.

To the venerable naturalist of Orient, New York, Mr.
Roy Latham, for introducing me to the Long Island hybrid
swarms, and for supplying me with A. spectabilis; to Dr.
Reed C. Rollins, Director of the Gray Herbarium, for
verifying my work, to Mr. Joseph Monachino, of the New

4Fryxell, P. A. 1957. Bot. Review 23(3): 169.

1962] Dahl — Cordilleran Species 117

York Botanical Garden, and Dr. Harry E. Ahles, of the
University of North Carolina Herbarium, for valued assis-
tance, I express my sincere appreciation.

— MADISON HEIGHTS, VA.

THREE MISIDENTIFIED SO-CALLED CORDILLERAN
SPECIES IN EASTERN NORTH AMERICA!

EILIF DAHL

Since the work of Fernald (1925) the cordilleran species
in eastern North America have attracted considerable atten-
tion. With increasing exploration, expecially of the formerly
little-known areas around Hudson Bay and the Canadian
Arctic Archipelago, many species previously considered as
cordilleran disjuncts have been shown to have a more or
less continuous area across the continent (Raymond, 1950).
Many of the remaining cordilleran disjuncts are serpen-
tinicolous species, halophilous species or species with adap-
tations to long-distance dispersal (light wind-dispersed
spores or seeds, hooks on the fruits, etc.). Only a very
limited list of species with no adaptations to long-distance
dispersal remains as cordilleran disjuncts and it will be
shown below that the records of three of them are based
upon misidentifications.

1. Agrostis rossae Vasey was reported from Newfound-
land at Bonne Bay by Fernald (1933 p. 203) based on a
specimen collected by K. P. Jansson. The specimen is kept
in the Gray Herbarium and consists of two culms, 18 and 20
cm. high, and attached basal tufts of leaves, the leaves being
about 1 mm. broad. Dead leaves from the previous year
testify the plant to be perennial thus corresponding to the
Rocky Mountain A. variabilis Rydb., the name A. rossae
Vasey being reserved for an annual plant restricted to hot
springs in Yellowstone Park, Wyoming (Chase 1950 p. 342
and 346). The panicle is narrow, almost spikelike, with

1Contribution from the Herbarium of the University of Colorado Museum.

118 Rhodora [Vol. 64

spikelets near the base of the lower branches. In these
characters the Newfoundland plant compares well with
Rocky Mountain A. variabilis.

However, in some important characters the Newfoundland
plant differs from A. variabilis. Most important is the
presence of a palea a little more than half the length of the
lemma, while in true A. variabilis the palea is minute or
absent. In the Newfoundland plant the ligule on the culm
leaves is 0.3-0.4 mm. long and on the sterile shoots about 0.6
mm. and rather transversely cut. In Rocky Mountain A.
variabilis the ligule of the culm leaves is 1.3-2.0 mm. long,
somewhat acute and continuing downwards as a hyaline
margin of the sheath. Thus it is evident that the Newfound-
land plant neither belongs to A. variabilis nor to A. rossae
s. str.

In the characters of the palea and the ligule the New-
foundland plant compares well with the European A. tenuis
Sibth. and it was found in an area where a European ele-
ment in the flora is strongly pronounced. Lindroth (1957)
has shown how many of these species became introduced
during the early fishing trade between Europe and New-
foundland. A. tenuis is a highly variable species, usually
with a more spreading panicle and higher culms than the
Newfoundland plant, but the Newfoundland specimen comes
well within the range of variation in these respects of the
European A. tenuis.

2. Carex filifolia Nutt. was recorded by Delabarre (1902
p. 192) from Nachvak in Labrador, the identification being
accredited to Fernald. The specimen is kept in the herbar-
ium of Brown University, R. I. The material consists of two
culms, 10 and 13 em. high, with no leaves or rooting parts
attached, and carry spikes about 1 cm. long with an upper
staminate part and a lower pistillate part. The pistillate
flowers are badly infected by smut; no intact perigynia are
present.

Two species come in question for identification of the
scanty material, the cordilleran C. filifolia Nutt. and the
cireumpolar C. rupestris All; the latter is well known to

1962] Dahl — Cordilleran Species 119

occur in Labrador. The culms are slightly higher than
average for C. rupestris in the area, but well within the
range of variation of the species. The best character
separating the two species is that the perigynia are hairy
in C. filifolia and glabrous in C. rupestris. The hairs on the
perigynia may sometimes be seen even in smut-infected
flowers of C. filifolia; no such hairs were observed in the
Nachvak plant, but this is not entirely reliable. There are,
however, other indications that the Labrador plant belongs
to C. rupestris rather than to C. filifolia.

a. In C. filifolia the lower pistillate bracts are gibbous at
base forming a broad sac, and the bracts have a very wide
hyaline margin only the central part being brownish. In
C. rupestris and the Nachvak specimen the lower pistillate
bracts are not gibbous at base and are brown over most of
the area.

b. In C. rupestris and the Nachvak specimen the upper
part of the culm is sharply triangular, often with scabrosely
toothed margins and also in the middle and lower part of the
culm the triangular outline can be seen. In C. filifolia the
culms are round or very obtusely triangular and rarely
scabrosely toothed.

c. The smut on the specimen has been examined by Dr.
D. S. O. Savile, Department of Agriculture, Ottawa, who
writes: “Smut is Cintractia caricis (Pers.) Magn. affin. v.
caricis closely matching many specimens on C. rupestris
including Ungava peninsula. Smut on C. filifolia is quite
distinct."

Thus, it is almost certain that the collection from Nachvak
in Labrador belongs to C. rupestris and can on no account,
with confidence, be assigned to C. filifolia.

9. Polygonum bistortoides Pursh has been recorded from
southwestern Newfoundland by Fernald (1950 p. 580). The
Specimen was collected by A. S. Pease and M. S. Edgerton
on July 10th, 1939 in rills and open meadows in the Cape
Anguille Mts., Millville, and is kept in the Gray Herbarium.
It consists of four basal leaves with no fruiting or flower-
ing stems and does not look very similar to material of

120 Rhodora [Vol. 64

P. bistortoides collected in the Rocky Mountains. The leaves
are long, 16-22 cm. with long stipes, 12-22 cm. Such long
leaves can sometimes be found also in the Rocky Mountains.
A comparison of leaves of the same length from the Rocky
Mountains with the collection from Newfoundland gives a
number of significant differences which can be summarized

as follows:

Polygonum bistortoides
Rocky Mountains

Leaves (2)3-4(4.5) cm. broad.
Central nerve 0.5-2(3) mm. broad,
firm.

First order sidenerves 0.4-1.4 cm.
apart.
Epidermis even, hairy or pruinose
below.

Microscopic circular glands on the
nerves.

Marginal leaf-cells longitudinally
oriented, brown, with narrow lu-

*Polygonum bistortoides"
Newfoundland

Leaves 2-3 cm. broad.

Central nerve 2-3.5 mm. broad,
showing a number of separate
vascular bundles.

First order sidenerves 1.5-2 cm.
apart.

Epidermis scabrosely papillose or
wrinkled, neither hairy nor pru-
inose.

Glands lacking.

Marginal leaf-cells isodiametric,
hyaline, margin finely dented or

mina, margin smooth, curving eroded, plane.

downwards.

Thus it is evident that the Newfoundland collection does
not belong to P. bistortoides. In all characters outlined above
it compares better with a species of Rumex, particularly of
the group Avillares. Especially the characters of the central
nerve and the leaf margin serve to discriminate between the
two genera. It is not easy to assign the material to any
particular species of Rumex although Rumex pallidus might
be a good guess.

I am indebted to the curator of the Gray Herbarium and
to Professor Church, Brown University, R. I. for loan of
specimens, to Dr. D. S. O. Savile, Ottawa for identification
of a smut and to Dr. W. A. Weber, University of Colorado
for placing herbarium facilities at my disposal and for
stimulating discussions. — AGRICULTURAL COLLEGE OF NOR-
WAY, VOLLEBEKK, NORWAY.

1962] Hermann — Bryophyte Flora 121

LITERATURE CITED

CHASE, A. 1950. Manual of the grasses of the United States by A. S.
Hitchcock, 2nd ed. Washington.

DELABARRE, E. B. 1902. Report of the Brown-Harvard expedition to
Nachvak, Labrador in the year 1900. Bull. Geogr. Soc. of Phila-
delphia 3: 65-212.

FERNALD, M. L. 1925. Persistence of plants in unglaciated areas of
Boreal America. Mem. Am. Acad. 15(3) : 237-342.

1933. Recent discoveries in the Newfoundland flora.

Rhodora 35: 1-16, 47-63, 80-107, 120-140, 161-185, 230-247, 265-283.

1950. Gray's Manual of Botany, 8th ed. New York.

LINDROTH, C. H. 1957. The faunal connections between Europe and
North America. New York.

RAYMOND, M. 1950. Esquisse phytogéographique du Québec. Mem.
Montreal Bot. Gard. 5: 1-147.

ADDITIONS TO THE BRYOPHYTE FLORA OF
KEWEENAW COUNTY, MICHIGAN

F. J. HERMANN!

The Keweenaw Peninsula, on Lake Superior in northern-
most Michigan, became botanically renowned with the pub-
lication in 1935 of Fernald’s paper, Critical Plants of the
Upper Great Lakes Region of Ontario and Michigan (2).
In this paper Professor Fernald discussed at length an
impressive assemblage of species among the vascular plants,
known principally from the Rocky Mountains and the Pacific
coast but occurring also in isolated colonies on the Keweenaw
Peninsula and a few other restricted areas, possibly unglaci-
ated during the Pleistocene, far east of their principal range.

In 1937 and 1938 Dr. W. C. Steere published noteworthy
papers on the bryophytes of the Keweenaw Peninsula (10,
11) in which he pointed out that many of the mosses and
hepatics occurring on the Peninsula paralleled the disjunct
distribution of the Cordilleran, Pacific coast and arctic
vascular plants enumerated by Fernald. Western and arctic

1Forest Service Herbarium, Division of Range, Wildlife Habitat and Recreation
Research, U. S. Department of Agriculture, Washington, D. C.

122 Rhodora [Vol. 64

elements in the flora of Isle Royale (which is also part of
Keweenaw County) have likewise been known as a result
of Ruthven’s ecological survey (7) and of a fairly recent
survey of the bryophytes of the island (13), in which
account was taken of earlier reports by Holt (6) and
Cooper (1).

That the bryophyte flora of Keweenaw is still incompletely
known, however, was pointed out by Steere and was attested
by the results of a few days’ collecting by the writer around
Copper Harbor, on the peninsula, and Rock Harbor, on the
island, during August 1960. Because the bryophytes of
Michigan as a whole (12), and of the Isle Royale and the
Keweenaw Peninsula in particular, are comparatively well-
documented it seems worth while to record here the few
additions (which include three genera new to the State)
to the known flora that turned up in 1960.

The writer is grateful to Howard Crum and Margaret
Fulford for help in the identification of critical species, and
to A. LeRoy Andrews he is indebted for determination of
the Sphagnum and Bryum. Robert R. Ireland kindly gave
permission to include the record of his collection of Tricho-
don cylindricus made in 1961.

Species new to the flora of Michigan, based on Steere, 1947
(12), are indicated in the following list with an asterisk.
All of the species listed, with the exception of Andreaea
rupestris, are new to either the Keweenaw Peninsula or Isle
Royale according to the most recent reports ( 12,13). Speci-
mens have been deposited in the U. S. National Herbarium,
Washington, D. C. and duplicates in the herbarium of either
the University of Michigan, Ann Arbor or Michigan State
University, East Lansing, and the National Herbarium of
Canada.

KEWEENAW PENINSULA

Lophozia quadriloba (Lindl.) Evans. On conglomerate shore of Lake
Superior at edge of woods, Copper Harbor, 16354. Although this
species is not included in the most recent list (12) of Michigan bryo-
phytes, it has been subsequently reported from Copper Harbor by
Schuster (8) in his account of Minnesota hepatics.

1962] Hermann — Bryophyte Flora 123

Andreaea rupestris Hedw. On granitic outcrop, Mt. Bohemia, just
west of Lac La Belle, 16218. This moss is not new to either the Kewee-
naw Peninsula or Isle Royale but additional records are noted here
because of its rarity in the State. As pointed out by Steere (10) it has
previously been known in. Michigan from only a single locality on Isle
Royale and from one each in Keweenaw and Marquette Counties.

Encalypta procera Bruch. Abundant on conglomerate bluff at edge
of woods along Mandan Road (US 41), 1 mile south of Copper Harbor,
16236. (Determination verified by Seville Flowers). The geographical
distribution of this species is given by Flowers (3) as “Alaska to
Montana, Idaho and Washington. Common in British Columbia. Also
in Ontario, Greenland, Northern Europe and Asia." It was reported
from Isle Royale by Cooper (1), however, in 1913.

Heterocladium squarrulosum (Voit) Lindb. Base of gneiss outcrop
near shore of Lake Superior, Copper Harbor, 16151; moist conglom-
erate slope, gorge of Manganese River, 1% mile south of Copper Harbor,
16163. Apparently the only previous Michigan record for this species
is from the Porcupine Mountains, Ontonagon County.

*Oreoweisia serrulata (Funck) De Not. On moss-covered crest of
conglomerate shore of Lake Superior, Devil's Washtub, 115 miles west
of Copper Harbor, 16133; crevice in conglomerate shore of Lake Supe-
rior, Fort Wilkins, Copper Harbor, 16225. Also occurring on Isle
Royale as noted below. Although Grout (5) refers to this moss as
being a rare plant in North America (and a calciphile), unlike most
of the other Keweenaw novelties it is not predominantly western or
arctic in its distribution but is known also from eastern North America
and in the mountains as far south as Tennessee and Kentucky.

Orthotrichum sordidum Lesq. & James. On trunk of sugar maple in
woods near Lake Manganese, 1 mile south of Copper Harbor, 16101;
on trunk of ash, Silver Creek, 412 miles east of Eagle Harbor, 16138.

"Trichodon cylindricus (Hedw.) Schimp. (Ditrichum cylindricum
(Hedw.) Grout). On soil along road up Mt. Bohemia, alt. 867 ft.,
just west of Lac La Belle, R. R. Ireland, Jr. 5364, Aug. 21, 1961 (US).
This is another farwestern species which is evidently a glacial relic in
Michigan. Grout (5) gives its range as “western North America,
Yukon to Montana, Nevada and Washington."

"Trichostomum tenuirostre (Hook. & Tayl.) Lindb. (T. cylindricum
(Bruch) C. Muell) Moist crevices in conglomerate shore of Lake
Superior, Fort Wilkins, Copper Harbor, 16118. An earlier, but evi-
dently unreported collection, from Keweenaw County by W. C. Steere
and A. J. Sharp was recently noted in the University of Michigan
Herbarium by Mrs. Jennie Dieterle and called to the writer's atten-
tion. It was collected on Mt. Bohemia, September 1, 1937. Like
Oreoweisia this is a species of widespread distribution, occurring as
far south as the mountains of North Carolina and Arizona.

124 Rhodora [Vol. 64

ISLE ROYALE

Cephaloziella striatula (Jens.) Douin. With Lophozia ventricosa
on vertical face of saturated tussock of Scirpus caespitosus in rock
pool, Scoville Point, 1632542. Not recorded from Michigan in Steere's
list (12), and ascribed only to Greenland, Alaska, British Columbia
and Europe by Frye and Clark (4), but Schuster (8) reports the
species (as C. subdentata Warnst.) from Minnesota, Michigan (Che-
boygan County), Maine, New Hampshire and New York.

Scapania paludicola Loeske & K. Muell. In bed of Empetrum at edge
of rock pool, Scoville Point, 16328%.

Amblystegium serpens (Hedw.) BSG. On conglomerate near shore
of Rock Harbor, 16261.

Andreaea rupestris Hedw. On vertical face of basalt boulder along
trail to Scoville Point, 16325; on basalt boulder along trail to Hidden
Lake, 16333. Previously known on Isle Royale only from Passage
Island (13).

Anomodon attenuatus (Hedw.) Hiiben. In shallow cave below Mon-
ument Rock, 1634512. It seems remarkable that no species of this
usually very common genus has previously been reported from the
island.

Bryum cuspidatum (BSG.) Schimp. Crevice in trap rock shore of
Rock Harbor, 16281. A questioned report of B. affine (Bruch.) Lindb.
in Thorpe and Povah’s list (13) might be referable to B. cuspidatum.

Calliergon giganteum (Schimp.) Kindb. On Thuja log near Hidden
Lake, 16335.

Cynodontium schistii (Wahl.) Lindb. On vertical face of boulder in
shade, Monument Rock, 16349.

Dichodontium pellucidum (Hedw.) Schimp. Crevice in trap rock
shore of Rock Harbor, 16275.

Grimmia alpicola Hedw. On trap rock shore of Rock Harbor, 16262
and 16282. Only var. rivularis Brid. has been previously recorded
from Isle Royale.

Grimmia apocarpa Hedw. var. stricta (Turn.) Mitt. Extensive mat on
trap rock shore of Rock Harbor, 16260. Only the typical form of the
species has been reported from Isle Royale, although G. ambigua and G.
conferta, also reported, are now commonly treated as additional
varieties of the polymorphic G. apocarpa.

Mnium punctatum L, var. elatum Schimp. Crevice in conglomerate
shore of Rock Harbor, 16273. The variety has not been previously
reported from the island.

Oreoweisia serrulata (Funck) De Not. On trap rock shore of Rock
Harbor, 16280.

Orthotrichum obtusifolium Brid. On trunk of aspen, upper shore of
Rock Harbor, 16274 and 16284; on trunk of mountain ash, trail to
Scoville Point, 16324.

1962] Hermann — Bryophyte Flora 125

Orthotrichum sordidum Lesq. & James. On trunk of aspen, shore of
Rock Harbor, 1627312.

Pylaisia selwynii Kindb. On trunk of aspen, trail to Scoville Point,
16317.

Rhytidium rugosum (Hedw.) Kindb. In bed of Empetrum at edge
of rock pool, Scoville Point, 16331.

Sphagnum dusenii C. Jens. In Chamaedaphne bog, Raspberry Island,
16300.

Tortella fragilis (Drumm.) Limpr.? Crevices in trap rock shore of
Rock Harbor, Scoville Point, 16315.

LITERATURE CITED

1. Cooper, W. S. 1913. A list of mosses collected upon Isle Royale,
Lake Superior. Bryologist 16: 3-8.

2. FERNALD, M. L. 1935. Critical plants of the upper Great Lakes
region of Ontario and Michigan. Rhodora 37: 197-222, 238-262,
272-301, 324-341

3. FLOWERS, S. 1938. Encalyptaceae, in A. J. Grout: Moss Flora of
North America 1: 137-145.

4. FRYE, T. C. and L. CLARK. 1937-1947. Hepaticae of North
America. Univ. of Washington Publ. Biol. 6:1-1022. Seattle,

Wash.
5. GRoUT, A. J. 1928-1940. Moss Flora of North America, 3 vols.
Newfane, Vt.

6. Hott, W. P. 1909. Notes on the vegetation of Isle Royale, Michi-
gan. Rep. Mich. Geol. Surv. 1908: 217-248.

7. RUTHEVEN, A. G. 1906. An ecological survey in the Porcupine
Mountains and Isle Royale, Michigan. Ann. Rep. Geol. Survey
Mich. 1905.

8. SCHUSTER, R. M. 1953. Boreal Hepaticae, a manual of liverworts
of Minnesota and adjacent regions. Am. Midl. Nat. 49: 257-684.

9. STEERE, W. C. 1937. Bryophytes of the Porcupine Mountains,
Ontonagon County, Michigan. Pap. Mich. Acad. 22: 183-200.

10. . 1987. Critical bryophytes from the Keweenaw
Peninsula, Michigan. Rhodora 39: 1-14, 33-46.

11. . 1938. Critical bryophytes from the Keweenaw
Peninsula, Michigan, II. Ann. Bryol. 11:145-152.

12. . 1947. The bryophyte flora of Michigan. Pap.

Mich. Acad. I. 31: 33-56.
18. THORPE, F. J. and A. H. PovaH. 1935. The bryophytes of Isle
Royale, Lake Superior. Bryologist 38: 32-46.

*A collection of this species from Alger County (pockets in dry sandstone ledge,
Pictured Rocks, 9 miles northeast of Munising, 1609115) evidently represents the first
recorded from that county.

126 Rhodora [Vol. 64

CONTRIBUTION TO THE FUNGUS FLORA OF
NORTHEASTERN NORTH AMERICA. II

HOWARD E. BIGELOW AND MARGARET E. BARR’

Field work in Massachusetts during 1960 and 1961, and
studies of specimens collected in previous years throughout
the northeast, have revealed some agarics and pyrenomy-
cetes which seem worthy of description. The fruiting of
several other fungi is reported as well in order to extend
knowledge of their distribution. Two new combinations are
proposed: Clitocybe morgani (Peck) Bigelow and Phae-
ostoma sphaerophila (Peck) Barr.

We wish to express our appreciation to Stanley J. Smith,
Senior Curator of Botany, New York State Herbarium, and
to Dr. Clark Rogerson, Curator of Fungi, New York Botan-
ical Garden, for the opportunity of studying the type mate-
rial of several of the species discussed. Dr. Alexander H.
Smith, Director, University of Michigan Herbarium and Dr.
Kenneth A. Harrison, Research Station, Kentville, Nova
Scotia have helped in loan of specimens. We gratefully
acknowledge the financial support of the Faculty Research
Council, University of Massachusetts. Field work in Massa-
chusetts during 1958, 1959, and 1960 would not have been
possible without the assistance of the Council. Our research
in the fall of 1961 has been supported by National Science
Foundation Grant G 19534.

The colors cited in the descriptions of agarics are from
Ridgway, R. 1912. Color standards and color nomencla-
ture. Washington, D. C.

BASIDIOMYCETES

Amanita parcivolvata (Peck) Gilbert, in Bresadola, Icon. Myc. 27,
suppl. 1: 226. 1941.

Amanitopsis parcivolvata Peck, Bull. Torrey Club 27: 610. 1900.

Pileus 5-8.5 em. broad, convex to plane, margin striate, disc shal-
lowly depressed in age, surface with yellow (“maize yellow", “buff
yellow”) coating, powdery or somewhat floccose, glabrescent near dise,

ay. Rhodora 62: 186-198. 1960. Contribution from the Department of Botany, Univer-
sity of Massachusetts, Amherst.
?Mrs. Howard E. Bigelow.

1962] Bigelow and Barr — Fungus Flora 127

margin appendiculate with yellow-floccose patches at first, soon
appressed, ground color orange-yellow (“light orange yellow’) or
darker; flesh thin, firm, concolorous with pileus or paler, odor none,
taste not known.

Lamellae free, distant from stipe, close, moderately broad (up to 1
cm.), light yellow (“massicot yellow”, “straw yellow"), edges dentate
to crenate from particles of partial veil.

Stipe 8-15 em. long, 5-10 mm. in diameter at apex, 1-2.5 cm. at base,
base clavate to subbulbous, fragments of universal veil forming orange-
yellow floccose patches on base of stipe, whitish beneath, apex and
median portion of stipe concolorous with lamellae, fibrillose or fur-
furaceous from remains of partial veil, yellow, powdery to flocculent,
central, stuffed or hollow.

Spores 10-13 x 6-8 y, elliptical, smooth, not amyloid, white in mass;
basidia 32-46 x 8.5-13 u, 4-spored; cystidia: cheilocystidia and pleuro-
cystidia present but scattered, basidioid to saccate, 15-21 y. in diameter,
smooth ; pileus tissue: surface often with cells from veil, broadly fusoid
to cylindrical or saccate, 7.5-50 y, in diameter, walls smooth or slightly
roughened, thin or slightly thickened, cuticle thin, hyphae cylindrical,
3-5 y in diameter, trama thin, hyphae cylindrical to inflated, 1.5-11
u in diameter, clamp connections present, scattered laticiferous hyphae
present; gill trama bilateral, hyphae cylindrical to inflated, 5.5-15.5 y.
in diameter.

Solitary, on soil or humus in open hardwoods of beech, birch, and
maple.

Material examined: Bigelow 9232, D.A.R. State Forest, Goshen,
Massachusetts, Sept. 1, 1960; 9778, Conway, Sept. 5, 1961; 9956,
Conway, Oct. 9, 1961.

A. parcivolvata appears to be typical of a more southern agaric
flora, but does appear in the northern states in small quantity upon
occasion. Hesler (1960) has published a photograph of the species.

Pilát (1954) lists A. parcivolvata as a synonym of Amanita
muscaria. The two species are quite distinct by the nature of the veils.
In A. parcivolvata both veils are powdery and very fragile, never
forming warts or a persistent, membranous annulus as in A. muscaria.

Cantharellus minor Peck, N. Y. State Cab. Rep. 23: 122. 1872. Plate
1268.

Pileus 5-30 mm. broad, convex at first, becoming plane with a nar-
rowly decurved margin and the disc slightly depressed, finally infundi-
buliform with the margin arched and undulate, even, glabrous,
yellow-orange to orange when moist (“orange”), fading to pale
orange-buff or pale orange; flesh concolorous with pileus, thin, no
odor or taste when fresh, at times developing fragrant odor like
Cantharellus cibarius after drying.

Lamellae decurrent (unevenly), distant, very narrow, forked, not

128 Rhodora [Vol. 64

intervenose, edges even, somewhat thickeed, straight, concolorous with
the moist pileus (“orange”).

Stipe 1.5-2.5(-5) em. long, 3-7(-9) mm. thick at the apex, base
attenuated, central, hollow, glabrous, compressed and furrowed at

PLATE 1268. Cantharellus minor Peck X 1.

times, often curved and flexuous, concolorous with moist pileus or
paler (“deep chrome"), not fading, base slightly more yellowish where
embedded in humus.

Spores 6-11.5 x 4-6 u, usually broadly elliptical or elliptical to oblong,
at times bent on apicular side, smooth, not amyloid, light yellowish-
orange in mass; basidia (34-)39-65 x 4.5-8.5 u, 4-spored; cystidia not
differentiated; pileus tissue: light yellow-orange or yellow in KOH,
pigment dilute in cell contents, cuticle thin and indistinct, hyphae
mostly cylindrical, 2.5-5.5 u in diameter, tramal hyphae cylindrical
to inflated, 3-12(-16) u in diameter, clamp connections present; gill
trama interwoven, hyphae mostly cylindrical, 1.5-4.5 y in diameter.

Scattered to gregarious, on humus, in open hardwoods, particularly
of beech and maple. July to August.

Material examined: Bigelow 3755, Madawaska, Maine, Aug. 2, 1956;
7015, Amherst, Massachusetts, July 23, 1958; 7080, Amherst, July 24,
1958; 7229, Amherst, July 30, 1958; 7549, Mt. Tom State Reservation,
Holyoke, Aug. 19, 1958; 8888, Williamsburg, July 14, 1960; 9052,
Conway State Forest, Conway, Aug. 9, 1960; 9524, Amherst, July 14,
1961; 9542, Amherst, July 18, 1961; 9587, Conway State Forest, Con-
way, July 23, 1961; 9622, Pittsfield- State Forest, Pittsfield, July 26,

1962] Bigelow and Barr — Fungus Flora 129

1961; 9632, Conway State Forest, Conway, July 27, 1961; 9661, New
Salem, Aug. 2, 1961; C. H. Peck, Greenbush, New York, July (type of
C. minor).

The position of this fungus, is difficult to determine. It resembles
C. cinnabarinus Schw. in stature, but C. cibarius Fr. in color. On
the other hand, there is a waxy quality which first led me to try
and determine it in Hygrophorus. Possibly, C. minor occupies an
intermediate position between the two genera, but certainly the colored
spore deposit indicates a closer relationship to Cantharellus.

Of uncertain relationship is Cantharellus Friesii Quélet which occurs
in Europe. According to the literature this species seems very close
to C. minor.

Clitocybe morgani (Peck) Bigelow, comb, nov.

Cantharellus morgani Peck, Bot. Gaz. 7: 43. 1882.

Cantharellus olidus Quélet, Ench. Fung. p. 138. 1886.

Cantharellus rosellus Peck, N. Y. State Mus. Rep. 42: 24. 1889.

Clitocybe olida (Quél.) Konrad, Bull. Soc. Myc. Fr. 45: 60. 1929.

Hygrophoropsis olida (Quél.) Métrod, Rev. de Myc. 14, Suppl., p. 15.

1949.

Material examined: Bigelow 3432, near Guerette, Aroostook Co.,
Maine, July 23, 1956; 3569, July 26, 1956; 3594, 3626, July 29, 1956;
3704, Aug. 1, 1956; 4442, Aug. 28, 1956; J. Dearness, 2696, Kilworth
woods, near Toronto, Ontario, Sept. 14, 1928; K. Harrison 191, Heigh-
bury, Nova Scotia, Oct. 10, 1932; 2466, Glenmont, Sept. 28, 1953;3984,
Waternish, Guysborough Co., Sept. 5, 1957; 3985, Cape Split, Kings
Co., Sept. 1, 1958; 4630, Sept. 1954; 4631, Aspen, Antigonish Co.,
Sept. 2, 1956; 4641, Glenmont, Kings Co., Oct. 11, 1952; 4642, Casey's
Corner, Kings Co., Oct. 12, 1952; 4643, Ayleford Lake, Kings Co., Oct.
10, 1952; A. P. Morgan, Vermont (type of Cantharellus morgani) ;
C. H. Peck, North Elba, New York, September (type of Cantharellus
rosellus).

Formerly, this agaric was treated by most investigators under the
name Clitocybe olida. A complete description based on specimens from
Oregon and Washington has been published by Smith (1944). How-
ever, recent study of type material in the Peck Herbarium at Albany
reveals that Cantharellus morgani is identical, necessitating a new
combination in Clitocybe. The dried carpophores of the type agree in
all observable details with specimens determined as Clitocybe olida
from several localities in North America. Peck’s description also com-
pares favorably in most respects with that of Cantharellus olidus
Quélet. Peck did not mention any odor for C. morgani, but this discrep-
ancy does not seem serious in view of the collaborating evidence of
other field characters and microscopic features.

Clitocybe morgani appears confined to a coniferous habitat, particu-
larly spruce and fir in the northeastern region. The distribution of

130 Rhodora [Vol. 64

the species in North America is broader than, previously known though
fruiting seems abundant only in certain restricted localities. Besides
the collections cited above I have had also the opportunity to examine
material deposited at the University of Michigan Herbarium from
Colorado, Idaho, Michigan, Oregon, and Washington.

!
|
D —
|.
i
|

PLATE 1269. Laccaria trullisata (Ellis) Peck. X 1/2.

Laccaria trullisata (Ellis) Peck, N. Y. State Mus. Bull. 157: 90. 1912.
Plate 1269.

Agaricus trullisatus Ellis, Bull. Torrey Club 5: 45. 1874.

Clitocybe trullisata (Ellis) Saccardo, Syll. Fung. 5: 195. 1887.

Pileus 2.5-6 cm. broad, convex to plane with an incurved margin,
even, disc depressed at times, surface dry, matted-fibrillose to scaly,
areolate and cracked at times, color often irregular, streaked and
mottled, with various shades of brown and buff (“vinaceous buff”,
“avellaneous”, “fawn”, “army brown”, “wood brown”, “russet”,
“mikado brown”, “tawny”, “cinnamon brown”, “cinnamon”); flesh
rather thick, soft, concolorous with pileus or lamellae, no odor or
taste.

Lamellae adnate to decurrent (unevenly), subdistant to distant,
broad (up to 12 mm.), not forked or intervenose, color purplish when
young (“vinaceous purple”, “slate purple”, “deep slate purple”, “dark
lavender”), rufous in age (nearest “army brown”, “sorghum brown”),
pruinose at times, edges even, thick, brittle.

Stipe 2-10 cm. long, 5-20 mm. thick at apex, tapering upward from
a long clavate or clavate-bulbous base, deeply embedded in sand, often

1962] Bigelow and Barr — Fungus Flora 131

compressed, solid-stuffed, central, surface fibrillose-streaked above,
reddish brown (“tawny”, *russet", “mars brown”, “cinnamon brown"),
base purplish and concolorous with young lamellae, often with purplish
mycelium.

Spores 11.5-19 x 6-7 y, elliptical to oblong, finely punctate under
oil immersion, not amyloid, white in deposit; basidia 42.5-60 x 10-13 u,
mostly 4-spored, occasionally 2-spored, sterigmata 4.5-7.5 x long;
cheilocystidia present in some sections, basidioid to broadly cylindrical
or broadly fusoid, 14-19 » broad, 31-46 u long, protruding beyond
hymenium 15-30 y; pileus tissue: cuticle brownish in KOH, pigment
dilute in cell contents or in slightly thickened walls, cuticular hyphae
interwoven, numerous end cells protruding beyond surface, cylindri-
cal or subclavate, hyphae mostly cylindrical, 5.5-11.5 » in diameter,
trama hyaline, hyphae 7.5-14.5 y, in diameter, cylindrical to inflated,
clamp connections large and abundant; gill trama regular to sub-
parallel, hyphae mostly cylindrical, 4-8 y, in diameter.

Solitary, scattered, or subcespitose. In sand, near open white pine.
September to October,

Material examined: Bigelow 6314, North Sunderland, Massachu-
setts, Sept. 24, 1957; 7837, Sept. 25, 1958; 9342, Sept. 22, 1960; 9792,
Sept. 6, 1961; 9922, Northfield, Oct. 7, 1961.

Abundant fruitings of L. trullisata are common on certain dunes
along the Connecticut River valley. While associated with white pine
in this region, this agaric occurs under jack pine on the Lake Superior
shore of northern Michigan,

The spores of this species are reported as smooth in the literature.
However, under the oil immersion lens there are very fine punctations
at least on some of the spores. These markings are difficult to observe
and it was not possible to tell if they were protrusions or pits. The
presence of some ornamentation lends support for maintaining this
species in Laccaria.

Lentinus haematopus Berkeley, Grev. 1: 33. 1872.

Material examined: Bigelow 3401, Guerette, Aroostook Co., Maine,
July 20, 1956;4060, Aug. 16, 1956.

This species, unusual because of the curious red coating on the stipe,
has been reported from a variety of locations in the northeast. It
seems widely distributed but fruits seldom and then only in small
numbers. I know of collections from Vermont, New Hampshire,
Ontario, New York, Michigan, and Saskatchewan, as well as the new
records cited above. Solitary specimens were found on birch or maple
logs in both Maine collections. A complete description and photograph
are found in Overholts (1934).

Mycena urania (Fr.) Gillet, Champ. Fr. 1: 279. 1878.

Agaricus uranius Fries, Syst. Myc. 1: 144. 1821.

Material examined: Bigelow 5256 and 5257, Mt. Albert, Gaspé Parc,
Quebec, July 7, 1957.

132 Rhodora [Vol. 64

Numerous carpophores were found on wet moss near a stream by
the Park Headquarters at Mt. Albert, Quebec. This species is noted
as rare by Smith (1947), in his monograph of Mycena. Previous
records from North America are from Michigan, North Carolina, and
Tennessee.

Tricholoma davisiae Peck, Bull. Torrey Club 27:611. 1900.

Melanoleuca davisiae (Peck) Murrill, N. Am. Fl. 10: 14. 1914.

Pileus 10-13 cm. broad, obtuse conic at first, becoming plane with a
slight broad umbo, margin narrowly incurved, even, sinuate or lobed
at times, somewhat elevated and recurved in places when old, surface
dry, radiate-fibrillose on the disc, fibrillose-scaly at the margin, ground
color dull yellow, fibrils smoky gray; flesh whitish, moderately thick
on the disc, thin at the margin, odor and taste farinaceous.

Lamellae sinuate, close, broad, yellow but bruising reddish, edges
uneven, undulate.

Stipe 10-11 cm. long, 2-4 cm. thick at apex, base abruptly tapered,
central, solid-stuffed, surface silky fibrillose, white but bruising reddish.

Spores 6-7.5 x 4.5-5.5 u, subglobose to elliptical, smooth, not amyloid,
white in mass; basidia 21.5-30 x 5.5-7.5 y, 1-, 2-, and 4-spored; cheilo-
cystidia present, clustered or scattered, obtuse or clavate, sometimes
pedicellate, smooth, hyaline or with yellowish contents in KOH, 23-36 y.
long, 6-15 y in largest diameter, two-celled at times, pleurocystidia
also present at times but rare, size and shape same as cheilocystidia ;
pileus tissue: cuticle pale yellowish in KOH, pigment apparently in
slightly thickened walls or as very fine encrustation, hyphae cylindrical
to somewhat inflated, 4.5-18 y in diameter, cells short, 11.5-84.5
v. long, trama hyaline, hyphae cylindrieal or somewhat inflated,
4-13 y in diameter, smooth, walls thin, cells short to long, clamp
connections absent; gill trama regular to subparallel, broad, hyphae
cylindrical to inflated, 3-12 » in diameter.

Gregarious, on humus at roadside in mixed woods. September to
November.

Material examined: Bigelow 9288, Conway State Forest, Conway,
Massachusetts, Sept. 18, 1960; 9381, Sept. 29, 1960; H. C. Davis, Fal-
mouth, Maine, November, 1899 (type of Tricholoma davisiae).

As far as I have been able to determine, this is the first record of
fruiting for this species since the original description based on speci-
mens found in Falmouth, Maine. The Massachusetts collections cited
above were found growing along a roadside in a mixed woods of beech,
birch, maple, pine, and hemlock,

T. davisiae is a striking species by virtue of the colors and size of
carpophore as well as the microscopic characteristics. Particularly
noteworthy is the cuticular structure of the pileus and the presence
of cystidia on the lamellae. The cuticle, composed of very short cells,
cannot be considered “hymeniform”, for the cells are recumbent and
apparently form from the fibrils on the cap surface. The cystidia

1962] Bigelow and Barr — Fungus Flora 133

usually do not protrude beyond the basidia to any great extent, but
they are conspicuous by their diameter and frequent two-celled nature.
One-spored basidia were observed upon occasion but they did not attain
the size of the cystidia. Cheilocystidia were not difficult to locate in
the Massachusetts collections, but pleurocystidia seemed less frequent
than in the type collection.

Xeromphalina kauffmanii A. H. Smith, Pap. Mich. Acad. I. 38:81.
1953.

Material examined: Bigelow 9605, Pittsfield State Forest, Massa-
chusetts, July 27, 1961.

The type collection and additional material of this species were
found on oak stumps during late spring and early summer in Michigan
according to Smith (1953). The Massachusetts collection is small in
number of carpophores and was found on a birch stump.

Figs. 1, 2. Ophiodothella vaccinii Boyd: 1, ascus, 2, two spores, on the right showing
a pseudoseptum in Melzer’s reagent. Figs. 3-5. Phaeostoma sphaerophila (Peck)
Barr, comb. nov.: 3, habit of fungus over black knot, 4, asci and hypha from centrum,
5, ascospores, those on the right showing scattered protrusions. Figs. 1, 2, 4, 5, X 1500;
fe. S x 15.

ASCOMYCETES
Ophiodothella vaccinii Boyd, Mycologia 26: 465. 1934. Figures 1, 2.
Material examined: Barr 2855, Mt. Toby, Leverett, Massachusetts,
Oct. 20, 1960; W. H. Davis 2884, Leverett, Oct. 20, 1922 (as Rhytisma

134 Rhodora [Vol. 64

vaccinii Schw.) ; E. S. Boyd, Athens, Georgia, April, 1933 (cotype of
Ophiodothella vaccinii).

This fungus appears to be predominantly southern in distribution.
Most of the herbarium specimens examined were from Georgia, Mis-
sissippi, and South Carolina, on Vaccinium arboreum. The Davis
specimen cited above was on V. stamineum, and although immature
was identical with my collection on V. corymbosum.

My collection was immature when gathered but after four days in
moist chamber produced asci and spores. It agrees in most respects
with Boyd's (1934) detailed description and with authentic material
of the species. Smaller leaf spots, lack of a conidial stage, and only
partial development of a second beak in the Massachusetts material
were the differences noted. Boyd's description should be correeted in
one respect. There is no amyloid reaction (I+) in the ascus, as she
stated, but the inner perithecial wall turns blue in Melzer's reagent.

Boyd suggested that Ophiodothella should be placed in the Clypeo-
sphaeriaceae, because of the development of a clypeus. However, the
sphaeriaceous structure, immersed perithecia, and stroma of super-
ficial blackened clypeus and light-colored hyphae within the substrate,
all agree with the Polystigmataceae of modern authors. O. vaccinii
and the other species of Ophiodothella which I have examined are
quite closely related to Polystigma, Phyllachora, and Diachora of the
Polystigmataceae, differing chiefly in the filiform or cylindrical spores.
O. vaccinii appears to be particularly closely related to Diachora
onobrychidis (Fr.) J. Mueller, the type of that genus (Arx and
Mueller, 1954). Both fungi have immersed perithecia, blackened crusts
in epidermal layers, hymenium in an equatorial band at the sides, and
inner perithecial wall amyloid in Melzer’s reagent. Diachora appar-
ently has only a single beak, and the spores are ellipsoid and one-
celled. O. vaccinii differs from the other species of Ophiodothella in
the equatorial arrangement of asci and the frequent occurrence of
two beaks on the perithecium.

Fhaeostoma sphaerophila (Peck) Barr, comb. nov. Figures 3-5.

Periconia sphaerophila Peck, N. Y. State Mus. Rep. 34:50. 1881.

Sporocybe sphaerophila (Peck) Saccardo, Syll. Fung. 4: 609. 1886.

Perithecia 200-400 & in diameter, globose, with beaks 200-300 yj.
long (up to 1 mm. at times), 80-100 y. in diameter near base, tapering
to 50-60 y at truncate apex, thickly grouped in patches over surface
of ascostromata of Apiosporina morbosa, superficial on a thin basal
stroma of compacted brown hyphae; perithecial wall 27-40 y wide at
sides, 55-75 y wide at base, composed of numerous layers of slightly
compressed, polygonal brown cells, inner wall tissue of hyaline to
yellowish layers of compressed cells, 10-15 y wide, wall of beak of
blackish-brown, parallel rows of elongate cells, canal periphysate, wall
roughened externally by protruding hyphae and discharged spores.

1962] Bigelow and Barr — Fungus Flora 135

Asci 11-16.5 x 6.5-10 y, ovoid, with a delicate stalk of variable length,
wall single, thin, deliquescent, 2-spored, numerous, arising from
interior of centrum wall, interspersed with short, broad, septate
hyphae.

Spores 6-10 x 4.5-6.5(-7.5) u, oblong to doliform in side view, ovate
in end view, hyaline becoming light brown, pigment concentrated in
a broad band across the middle of cell, leaving ends hyaline, one-celled,
wall double, thick and brown at sides, hyaline over ends, at times,
especially when mounted in Melzer’s reagent, bearing scattered low
hyaline protrusions over surface, usually remaining in pairs after
deliquescence of ascus wall.

Hyperparasitic on ascostromata of Apiosporina morbosa (Schw.)
Arx, on various species of Prunus.

Material examined: Barr 2099, Lake Munroe, Mt. Tremblant Park,
Quebec, July 24, 1957; 2164, July 29, 1957; 2839, Conway State Forest,
Conway, Massachusetts, Sept. 18, 1960; C. H. Peck, Edmond’s Pond,
Adirondack Mts., New York, July, 1880 (type of Periconia sphaero-
phila).

I first became aware of the ascomycetous nature of this fungus on
studying the Quebec and Massachusetts specimens. Type material of
Periconia sphaerophila is likewise an ascomycete with well-developed
perithecia and delicate, deliquescent asci, agreeing in all respects with
my collections. Additional records of the species were obtained from
herbarium collections of Apiosporina morbosa at the Universities of
Massachusetts and Michigan. These are: W. G. Solheim 622, Mycoflora
Saximontanensis Exsiccata, Williams Canyon, Manitou Springs, Colo-
rado, Dec. 22, 1949; V. M. Spaulding, Ann Arbor, Michigan, 1879;
E. Bartholomew 2335, Fungi Columbiani, Long Pine, Nebraska, Oct.
12, 1906; A. H. Povah, Dells, Wisconsin, May, 1929. From the scattered
distribution records thus obtained, it seems that Phaeostoma sphaero-
phila must be of frequent occurrence on the common black knot disease
of plums. The species is readily recognized in the field by the rough,
spiny surface caused by the long protruding beaks over the otherwise
smooth knot.

The superficial perithecia with carbonaceous dark wall, long beak,
thin-walled asci, and one-celled brownish spores are characteristic of
the genus Phaeostoma Arx and Mueller (1954). Doguet (1955) dis-
cussed the fungus, as Melanospora sphaerophila Sacc. He noted that
this name was given on the label of Reliquiae Farlowianae 56, but he
could not discover if Saccardo had actually published the combination.
I was also unable to find valid publication of the combination. Accord-
ing to Doguet's description, Reliquiae Farlowianae 56 is identical with
specimens which I have examined. Doguet observed the similarity in
shape of the ascospores of this species and Melanospora barbata (Fr.)
Dur. and Mont. (cf. Barr, 1959), but recognized that the carbonaceous

136 Rhodora [Vol. 64

walls of the perithecia differed entirely from the soft, thin wall of
Melanospora.

Within the genus Phaeostoma, P. sphaerophila differs from the three
previously described species in habit, in two-spored asci, and in dif-
ferently shaped ascospores. P. vitis (Fckl) Arx and Mueller, the
type species, occurs on rotting roots of various plants such as Vitis
and Humulus, has four-spored asci and spores which are 5-6 x 3-4 y,
ellipsoid to ovoid, with a germ pore at the lower end. P. juniperina
(Elis and Everh.) Arx and Mueller occurs on species of Juniperus
and forms gall-like growths on twigs. It has six- to eight-spored asci,
with ellipsoid to globose spores 3-4 y in diameter, and a small germ
pore. P. lagenarium (Pers. ex Fr.) Munk develops on old fruiting
bodies of Polyporus. The asci are four-spored, the spores 13-16 x 7-10
u with apiculus and germ pore at each end.

The recent publication of Wehmeyer's Monograph of Pleospora and
its segregates (1961) is of great value in the determination of these
fungi. Included here are records of three species of Pleospora, to
extend our knowledge of the distribution of these fungi.

Pleospora calvescens (Fr.) Tul., Sel. Fung. Carp. 2:266. 1863.

Additional synonym: Leptosphaeria eutypoides Peck, N. Y. St.
Mus. Rep. 38:105. 1885. Type material from the Peck Herbarium
at Albany, collected by C. H. Peck, on dead stems of Chenopodium
album, W. Albany, New York, May, 1884, agrees in all respects with
Wehmeyer's description of P. calvescens. The spores are three-septate
and lack a vertical septum, which undoubtedly influenced Peck in his
choice of the generic name.

Pleospora laricina Rehm, Hedwigia 21:121. 1882. var. laricina.

On branches of Cornus florida, Amherst, Mass., Aug. 10, 1959, Barr
2619.

Pleospora straminis Sacc. and Speg., Michelia 1:407. 1879.

On overwintered stalks of Zea mays, North Amherst, Mass., April
29, 1959, Barr 2549. This collection has spores somewhat broader
(10.5-15 u.) than does P. straminis, and approaches P. pulchra Kirschst.
in that respect. However, the spores are shorter than in P. pulchra,
have fewer septa, and are less strongly asymmetrical. Both species
occur on reddish spots on the substrate, as does my collection.

On leaves of Carex sp., Mt. Albert, Gaspé Parc, Quebec, July 11,
1957, Barr 2020. Although the substrate was not reddened, this collec-
tion is otherwise identical with that on Zea, and with Wehmeyer's
description.

LITERATURE CITED

ARX, J. A. VON and MUELLER, E. 1954. Die Gattungen der amero-
sporen Pyrenomyceten. Beitr. Kryptogamenfl. Schweiz 11(1): 1-
434,

1962] Johnston — Noseburn 137

Barr, M. E. 1959. Northern Pyrenomycetes I. Canadian Eastern
Arctic. Contr. Inst. Bot. Univ. Montréal 73: 1-101.

Bovp, E. S. 1934. A developmental study of a new species of Ophio-
dothella. Mycologia 26: 456-468.

DoGuET, G. 1955. Le Genre “Melanospora”. Botaniste 39: 1-313.

HESLER, L. 1960. Mushrooms of the Great Smokies. U. of Tenn.
Press, Knoxville. 289 pp.

OvERHOLTS, L. O. 1934. Mycological notes for 1933. Mycologia 26:
502-515.

PILAT, A. 1954. Mushrooms. H. W. Bijl, Amsterdam. 342 pp.

SMITH, A. H. 1944. Unusual North American agarics. Am. Midl.
Nat. 32: 669-698.

1947. North American Species of Mycena. U. of

Mich. Press, Ann Arbor. 521 pp.

1953. New and rare agarics from the Douglas Lake
region and Tahquamenon Falls State Park, Michigan, and an
account of the North American species of Xeromphalina. Pap.
Mich. Acad. I. 38: 53-87.

WEHMEYER, L. E. 1961. A world monograph of the genus Pleospora
and its segregates. U. of Mich. Press, Ann Arbor. 451 pp.

THE NOSEBURN (TRAGIA, EUPHORBIACEAE)
OF WESTERN TEXAS

MARSHALL C. JOHNSTON

Widespread in the arid grasslands and brush of northern
Mexico, north to Arizona, Colorado, and Kansas, occur the
humble, nondescript, stinging herbs which the English-
speaking natives usually call stinging-nettle, and the
Spanish-speaking natives chichicastle, a term also loosely
applied to several stinging herbs. A more specific name,
applied by some cowmen in western Texas, is noseburn, an
allusion to the effect of the plants on stock. Noseburns are
often quite abundant in grazed grasslands, and are among
that great retinue of noxious, toxic or unpalatable plants
which flourish under the abusive practices of overstocking
which are the rule rather than the exception in the range-
lands of western Texas. Noseburns can be recognized
among our other genera of this family by the weak, trailing
or even vine-like stem, the stinging hair, the serrate leaf,

138 Rhodora [Vol. 64

and especially the placement of the short androgynous
raceme which is actually terminal, but in development is
quickly surpassed by the branch from the axil of the im-
mediately subtending leaf, and thus appears nodal (it has
been described as axillary, though it rises on the opposite
side of the stem from the leaf).

The habit, leaves, inflorescences, and flowers of our nose-
burns are somewhat variable. On the basis of combinations
of various quantitative characters such as texture of style
and narrowness of leaf the German monographers Mueller
Argoviensis (1866: 930-935) and Pax and Hoffman (1919:
33-52), the latter of whom tended to follow the former
slavishly and uncritically, have discerned at least 9 taxa,
species and varieties, among our noseburns. Even more
curiously, they distribute these taxa into two different sec-
tions of the genus. A synoptic key to these sections may
appear more or less as follows, and I say “more or less”
because the species descriptions given by these authors
contradict the characters given for the more inclusive taxa,
and I have had to compile and modify to obtain a clearer
picture: .

Stamens (1—)3(— rarely 4); filaments free or united at the very
base; rudiment very small or rarely absent; staminate calyx 3-
parted or less commonly 4-parted; pistillate calyx 6-parted or
rarely fewer-parted ................ eene Tragia Sect. Tragia.

Stamens (3 —)4 — 6(— 8 or even rarely 20); filaments commonly
coherent; rudiments small or absent; staminate calyx 4- or
5-parted; pistillate calyx 5- or 6-parted .............. eee
NERA Tragia Sect. Leucandra (Klotzsch) Muell. Arg.

Perhaps their haste, governed by the magnitude and ambi-
tiousness of their task, and the paucity of American material
available to them, can explain their acceptance of such weak
categories.

Exomorphic studies of a large amount of material in the
herbarium and in the field in many parts of Mexico and
western and southern Texas permit the following state-
ments. The number of stamens is usually 3, but varies to
4 even in the same raceme. The staminate calyx is usually

1962] Johnston — Noseburn 139

3-lobed, but again varies to 4-lobed even in the same raceme.
Often, after a raceme has produced a number of staminate
flowers, the meristematic tip ceases to produce normal
flowers, and does not elongate further, but produces a
terminal “flower ”, probably representing the monstrous
joining of 2 or more flowers, with 5 or more sepals and 5 to
6 or more stamens (such “flowers” could have been the
basis of species and varieties of the “Sect. Leucandra" in
our area). Our noseburns commonly have the filaments
coherent basally around a minute central rudiment. The
pistillate calyxes are usually 6-lobed but sometimes only 5-
lobed. The various quantitative characters used by the
monographers to distinguish species and varieties occur in
every conceiveable combination and recombination. No
combination of such traits shows the broadscale geographic
restriction upon which a subspecies or variety could be
founded. Furthermore, the variability in our noseburns is
not greater than that encountered in many weedy wide-
Spread species of a number of other genera and families.

Our noseburns belong to a single species. Among the
congeries of names thus thrown together, the one with
priority is Tragia nepetifolia. MeVaugh (1961: 202-203)
says that this name has been “loosely interpreted,” and that
further study is needed. A study of material from near the
type locality, in the state of Hidalgo, convinces me that the
referral of our noseburns to T. nepetifolia by Harrington
(1954) and Shinners (1958 and 1961) and other writers of
floras does not represent a loose or incorrect interpretation.
The synonymy and typification are as follows:

Tragia nepetifolia Cav. Icones Pl. 6: 87, t. 558, f. 1. 1801.

Between Ixmiquilpan and Zimapán, Hidalgo, Sessé and/or Mocino
(MA?, F?). If no authentic herbarium material exists, the original
illustration will serve as type. Cavanilles misspelled the name “nepe-
taefolia." T. ramosa Torr. Ann. Lyc. N. Y. 2: 245. 1828. Sources of
the Canadian River, James exs. 407 (NY?). T. nepetaefolia var. ramosa
(Torr.) Muell. Arg. DC. Prodr. 15(2) : 934. 1866. T. angustifolia Nutt.
Trans. Amer. Phil. Soc. n.s. 5: 172. 1835. On the prairies of Red
River, in arid situations, prebably Choctaw County, Oklahoma, Nuttall,

end of May, 1819 (BM?, apparent isotype seen labeled merely “Red
River, Arkansas," PH). T. teucriifolia Scheele, Linnaea 25: 586. 1853.

140 Rhodora [Vol. 64

New Braunfels, Comal County, Texas, Lindheimer, July-Sept., 1846
(B?). Blankinship (1907: 148) indicates that the type collection has
the exs. No. 522 (Lindheimer field No. 299); but the material which
reached Scheele probably was not numbered the same as Blankinship’s
series at MO. T. nepetaefolia var. teucriifolia (Scheele) Muell. Arg.
DC. Prodr. 15(2) : 934. 1866. T. scutellariaefolia Scheele, Linnaea 25:
587. 1853. New Braunfels, Comal County, Texas, Lindheimer in July
1846 (B?) ; Blankinship (1907: 148) says the type is numbered exs. 521
(Lindheimer 298). T. stylaris var. angustifolia Muell. Arg. Linnaea
34: 180. 1865. The type bore the same exsiccatae number as the last.
T. nepetaefolia var. scutellariaefolia (Scheele) Muell. Arg. DC. Prodr.
15(2): 934. 1866. T. ramosa var. ? leptophylla Torr. Bot. Mex. Bound.
p. 201. 1859. “Near Howard’s Springs; Bigelow; No. 1796, Wright.”
T. stylaris var. leptophylla (Torr.) Muell. Arg. Linnaea 34: 181. 1865.
T. nepetaefolia var. leptophylla (Torr.) Shinners, Southw. Nat. 6: 101.
1961. T. stylaris Muell. Arg. Linnaea 34: 180, 1865. Novo Mexico,
"Wright 776”, according to Mueller, but this is probably Fendler 776,
as listed by Torrey (1859: 201), “On the upper Rio Grande, New
Mexico," and the type also of Tragia stylaris var. (alpha) latifolia
Muell. Arg. Linnaea 34: 180. 1865. T. nepetaefol’a var. latifolia Muell.
Arg. DC. Prodr. 15(2): 934. 1866. Novo Mexico [probably western
Texas], Wright exs. 1794; Mueller also cites three Mexican collecticns,
Aschenborn, Hartweg 82, and Virlet d'Aoust 121 (all G?). T. nepetae-
folia var. amblyodonta Muell. Arg. DC. Prodr. 15(2) : 934. 1866. Novo
Mexico [probably western Texas], Wright exs. 1793 (G?). T. amblyo-
donta (Muell. Arg.) Pax & K. Hoffm. Pflanzenr. IV 147 IX (Heft 68),
p. 51, 1919. T. scandens M. E. Jones, Extr. Contr. West. Bot. 18: 49.
1933, non Muell. Arg. 1865, synonymy fide C. V. Morton, Contr. U. S.
Nat. Herb. 29: 108-109. Guaymas, Sonora, M. E. Jones 23300, Jan. 26,
1927 (POM). [Morton's synonymy must be regarded as tentative.]
Tragia nepetaefolia var. angustifolia Muell. Arg, DC. Prodr. 15(2):
934. 1866. Texas, Berlandier 2542 (G?). The specimen was taken
between Matamoros and Goliad, southern Texas, in April or May, 1834.

Perennial herbs 5-35 cm. tall with short white rigid stinging hairs;
taproot 1-5(-12) mm. thick, brownish-white, often contorted; stems
few to many from the woody crown, the lower parts often buried,
brownish-white, resembling branched rootstocks; lower aerial stems
1-2 mm. thick, 5-50 em. long, often purplish-green basally, green to
grayish-green distally, weak and trailing or decumbent or even erect
(ours never twining), profusely alternately branched; branches not
greatly diverging; internodes 0.6-3(-9) cm. long. Leaves alternate;
blades linear-lanceolate to deltoid or even reniform, with or without
weak hastate lobes basally, or even ovate, 1-4.5 em. long (or slightly
longer in Val Verde County specimens), 0.7-6(-9) times as long as
broad, apically acute, basally truncate to cordate, marginally sharply
serrate, thin, green, pubescent; petioles slender, 2-10(-15) mm. long

1962] Johnston — Noseburn 141

(or to 30 mm. in Val Verde County) on lower leaves and often at an
angle to the blade, 1-2(-4) mm. long on upper leaves, parallel to
blade; stipules lanceolate, acute, entire, ca. 1-2.5 (-3.3) mm. long,
green but persistent and drying brown, glabrate. Flowers monoecious,
in androgynous ascending peduncled racemes 1.5-5 em. long, at the
nodes opposite the leaves (actually terminal and quickly surpassed by
the branch from the axil of the subtending leaf, this branch parallel
to the previous internode, thus appearing as a continuation of a single
main stem); peduncles 1-10 mm. long; the single, lowermost node
pistillate, the remainder 3-20-- nodes staminate, with internodes 0.5-
1.8 mm. long; bracts oblanceolate, subcucullate, ca. 1-2 mm. long,
acute, pubescent, entire. Staminate flowers: pedicels slender, 0.6-2
mm. long, with zone of abscission near middle, the lower part persis-
tent; sepals 3-4 (usually 3 but varying even on same raceme), valvate
in bud, oblanceolate to narrowly obovate-oblong, 1-1.7 mm. long,
glabrous or sparingly pubescent dorsally, spreading or reflexed at
anthesis; petals none; glands none; stamens 3-4, usually 4 but varying
even en same raceme [note that the terminal “flower” (probably rep-
resenting the monstrous joining of 2 or more flowers and not noticed
until the raceme is considerably elongated) sometimes has 5+ sepals
and 5-6+ stamens, as contrasted with the truly axillary flowers, and
as described for T. stylaris]; filaments thickened and fleshy, ca. 0.4-0.5
mm. long, connate basally; rudiment none or a minute central eminence
between the connate portions of the filaments. Pistillate flowers: pedi-
cels ca. 1 mm. long, elongating to 2-3 mm. in fruit; sepals 5-6, usually 6,
imbricate in bud, elliptic-lanceolate, ca. 1-1.5 mm. long at anthesis,
ca. 2-2.5 mm. long in fruit, acute, entire, hispid dorsally; petals
and glands none; ovary nearly round, densely long erect hispid;
styles 3, connate a third their lengths, 2-4 mm. long, entire, papillate-
fungoidal on the adaxial surface, coiling outward apically after
anthesis, often reddish or purplish. Capsules distinctly roundly tri-
coccous, ca. 3.5-4 mm. long, ca. 6-7 mm. broad, densely erect-hispid;
columella slender, whitish, 2-2.4 mm. long, with 3 long interlocular
points apically. Seeds nearly spherical, ca. 2.7-3.5 mm. long, when
mature brownish black with tawny tinge or mottling; surficially smooth
but not shiny; ecarunculate,

These plants occur from Veracruz, Hidalgo, San Luis
Potosí, and Zacatecas, north to Missouri, Kansas, Colorado,
and Arizona; and perhaps much farther south and east (cf.
MeVaugh 1961 : 202-203). They occur in a great variety of
circumstances, but are particularly abundant in grasslands
at moderate elevations and have been collected at elevations
up to 8000 feet in the Guadalupe Mountains of Culberson
County, Texas. — THE PLANT RESEARCH INSTITUTE, THE
UNIVERSITY OF TEXAS, AUSTIN.

142 Rhodora [Vol. 64

LITERATURE CITED

BLANKINSHIP, J. W. 1907. Plantae Lindheimerianae Part III. Misscuri
Bot. Gard. 18th Annual Rep. 123-224.

HARRINGTON, H. D. 1954. Manual of the Plants of Colorado for the
Identification of the Ferns and Flowering Plants of the State.
Denver: Sage Books. x, 666 p.

McVaucu, R. 1961. Euphorbiaceae Novae Novo-Galicianae. Brit-
tonia 13: 145-205.

MUELLER ARGOVIENSIS, J. 1866. Euphorbiaceae (pars). DC., Prodr.
15 (2) : 189-1286.

Pax, F. and K. HOFFMANN. 1919. Euphorbiaceae-Crotonoideae-
Acalypheae-Plukenetiinae. Pflanzenreich IV 147 IX (Heft 68, pars
primus). 108 p.

SHINNERS, L. H. 1958. Spring Flora of the Dallas-Fort Worth Area
Texas. Dallas: Privately published. v, 514 p.

1961. Tragia nepetaefolia var. leptophylla instead
of var. ramosa (Euphorbiaceae). Southwest. Nat. 6:101.

ToRREY,J. 1859. Botany of the Boundary in Emory, Wm. H., Report
on the United States and Mexican Boundary Survey, Washington.
II: 270.

A NEW SPECIES OF HAPLOPAPPUS,
SECTION BLEPHARODON

RAY C. JACKSON

Haplopappus texensis sp. nov.

Herba perennis 3-7 dm. alta; caulibus pluribus (vel unica), erectis,
striatis, tomentosis vel glabratis in maturitate; foliis alternis, sessili-
bus, ad 7 em. longis et 1.3 em. latis, oblanceolatis, lobis dentibusque
ad apeces spinuloso-mucronatis, insuper in maturitate sparse tomen-
tosis vel glabratis, subtuse tomentosis; capitulis pluribus, cymosis;
disco dia. 5-8 mm., receptaculis fimbriatis; phyllariis anguste lanceo-
latis, acuminatis, ad apeces spinulosis, dorsis tomentosis, ca. 5 mm.
longis et 1 mm. latis; radiis 22-28, 1-1.5 cm. longis; disci corollis glabris,
5.5-6 mm. longis; achaeniis turbinatis, pubescentis, 2.4-2.8 mm. longis;
pappis 6-6.8 mm. longis.

Perennial herb, 3-7 dm. tall, stems one to several, erect, striate,
tomentose or glabrate with age; leaves alternate, sessile, up to 7 cm.
long and 1.3 cm. wide, oblanceolate, the basal ones lobed, the median
ones dentate, lobes and teeth spinulose-mucronate, sparsely tomentcse
above or glabrate with age, tomentose below; heads several, cymosely
arranged; disc diam. 5-8 mm., receptacle frimbriate; phyllaries nar-
rowly lanceolate, acuminate, tomentose on the backs, spinulose at the

1962] Pohl — Agropyron Hybrids 143

tips, about 5 mm. long and 1 mm. wide; rays 22-28, 1-1.5 cm. long;
disc corolias glabrous, 5.5-6 mm. long; pappus 6-6.8 mm. long. Chro-
mosome number n = 4.

TYPE: Brooks County, Texas, sandy soil along railroad right-of-
way about 7.5 miles south of Falfurrias, Jackson 2938-1 (KANU),
August 7, 1959.

Haplopappus texensis is thus far known only from the
type locality where several hundred plants were observed.
A number of plants have been under cultivation since the
species was first collected in late August of 1957. Numerous
attempts to cross the species with morphologically related
taxa of the Blepharodon section of Haplopappus have been
unsuccessful.

On gross morphological characters, H. texensis may be
distinguished from related perennial species by its erect
growth habit, greater height, and less deeply divided leaves.
— UNIVERSITY OF KANSAS, LAURENCE, KANSAS

Field work pertinent to this study was supported by National Science Foundation
Grant G-7111.

AGROPYRON HYBRIDS AND THE STATUS OF
AGROPYRON PSEUDOREPENS'

RICHARD W. POHL

A number of specimens of Agropyron from Iowa (listed
in Table 1) have rhizomes similar to those of A. repens (L.)
Beauv. or A. smithii Rydb., but bear narrow spikelets with
enlarged, persistent glumes, resembling those of the cespi-
tose A. trachycaulum (Link) Malte. Examination of these
specimens shows that they have low seed set, ranging from
0-67% in the specimens examined. Pollen from these speci-
mens was mounted in lacto-phenol and cotton blue. It was
found that the pollen of each of these specimens was col-
lapsed, shrunken, and without stainable contents. (See
Figure 1).

Journal paper J-4249 of the Iowa Agricultural and Home Economic Expt. Station,
Ames. Project 1136. The facilities of the Iowa State University Herbarium, supported
by the Industrial Science Research Institute, were used in this study. The author wishes
to acknowledge the loan of specimens from the U. S. National Herbarium, Jason R.
Swallen, Head Curator, and from The Gray Herbarium.

144 Rhodora [Vol. 64

Figure Legend
Fig. 1-2. Pollen of Agropyron hybrids: Fig. 1. MeDill 295 (repens X trachycaulum)
Fig. 2. Rydberg 2018 (Type of A. pseudorepens)

Hubbard (1954) reported male-sterile hybrids of A.
repens with other species in Great Britain. He stated that
A. repens is self-sterile. Gillett and Senn (1960) have also
found self-sterility in the American rhizomatous species, A.
smithii Rydb. Many of our weed infestations of A. repens
probably have originated from single seeds, followed by
extensive vegetative spread. The absence of pollen of other
genetic strains of A. repens near such a colony would make
crossing with other species quite probable. Since A. trachy-
caulum is the only other common species of Agropyron in
Iowa, and since the pollen sterile plants are morphologically
similar to A. trachycaulum, the latter is probably the male
parent of these putative hybrids. Since the hybrids were
detected by the presence of rhizomes, other specimens of
similar origin but lacking rhizomes may have been un-
detected. This cross may have occurred much more fre-
quently than we are able to tell from herbarium records.

These pollen-sterile specimens key to A. pseudorepens
Scribn. and Smith in Hitchcock’s Manual. Fernald (1933)
regarded this “species” as belonging to A. trachycaulum,

1962] Pohl — Agropyron Hybrids 145

var. majus (Vasey) Fern. Beetle (1952) designated it as
A. trachycaulum, var. majus, f. pseudorepens (Scribn. &
Smith) Beetle.

The material of A. pseudorepens in the U. S. National
Herbarium was examined. The type specimen (Rydberg
2018, from Kearney, Nebraska ; U.S. 556663) bears rhizomes
and has leaf blades strongly ridged above. The glumes are
broad, strongly ridged, and nearly as long as the spikelets.
Morphologically, the type is intermediate between A. trachy-
caulum and A. smithii. Pollen taken from the anthers of
the type specimen is collapsed and shrunken (See Fig. 2).
The type of A. pseudorepens is therefore a probable male-
sterile hybrid of A. trachycaulum and A. smithii. Both
species are known from Nebraska. The name A. pseudo-
repens, if used, should be applied only to such hybrids. It
is not applicable to the Iowa hybrids, which are of other
parentage.

Examination of other rhizomatous specimens of “A.
pseudorepens" from the collections of the U. S. National
Herbarium, cited in Table 2, revealed many instances of
total pollen sterility. Such specimens may also represent
hybrids of various rhizomatous species of Agropyron with
the widespread A. trachycaulum.

Table 1. Agropyron repens X A. trachycaulum in Iowa

Specimen % seed set Pollen
Mason City. August 12, 1922 0 sterile
L. H. Pammel,

(ISC 106233)

Dickens, Clay Co. 67 sterile
J. Fults 2908 (ISC)

Dickinson Co., August 1921, 11 sterile
R. I. Cratty,

(ISC 97682)

Hayden Prairie, — sterile

Howard Co., R. C. McDill 295

July 9, 1952 (ISC)

Kossuth Co., Wesley Twp., 33 sterile
July 21, 1951, R. W. Pohl 7136 (ISC)

[Vol. 64

Rhodora

146

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1962] Lewis, Stripling & Ross — Chromosome Numbers 147
LITERATURE CITED

BEETLE, A. A. 1952. New names for Wyoming wheatgrasses.
Rhodora 54: 195-196.

FERNALD, M. L. 1933. Agropyron, Sect. Goularda in Eastern North
America. Rhodora 35: 161-185.

GILLETT, J. M. and H. A. SENN. 1960. Cytotaxonomy and infraspe-
cific variation of Agropyron smithii Rydb. Can. Journ. Bot. 38:
747-760.

HUBBARD, C. E. 1954. Grasses, pp. 77-83. Penguin. Hammonds-
worth.

SCRIBNER, F. LAMSON and J. G. SMITH. 1897. Native and introduced
species in the genera Hordeum and Agropyron. U.S.D.A. Div.
Agrostology Bull. 4:V:23-36

CHROMOSOME NUMBERS FOR SOME ANGIOSPERMS
OF THE SOUTHERN UNITED STATES AND MEXICO

WALTER H. LEWIS, H. LARRY STRIPLING!,
AND RICHARD G. Ross?

Chromosome numbers are reported for 34 families
including 57 genera and 73 species and varieties collected
in the southern United States and Mexico. The chromosomes
of one family, Turneraceae, and of 17 genera are given for
the first time. The new generic reports include those for:
Brunnichia (Polygonaceae), Cnidoscolus (Euphorbiaceae),
Cliftonia (Cyrillaceae), Piriqueta (Turneraceae), Rotala
(Lythraceae), Samolus (Primulaceae), Halesia (Styraceae),
Cynoctonum (Loganiaceae), Sabatia (Gentianaceae), Bon-
amia (Convolvulaceae), Pyenanthemum (Labiatae), Bacopa,
Buchnera, Lindernia, Mecardonia, Trigiola (Scrophulari-
aceae), and Sphenoclea (Campanulaceae). In addition, the
first definite chromosome numbers are recorded for Apios
(Leguminosae) and Utricularia (Lentibulariaceae). Among

1I wish to acknowledge the partial support from grant RG-6305, Division of General
Medical Sciences, Public Health Service, issued to Dr. R. K, Godfrey, Florida State
University, and to thank Dr. M. Y. Menzel for her help during my part of this study.

"Undergraduate Research Participant during 1960-61, Stephen F, Austin State
College, under the direction of the senior author (National Science Foundation,
G-12059).

148 Rhodora [Vol. 64

the 73 species and varieties listed, 53 are new chromosome
reports, 4 conflict with previous results, and 16 are verifi-
cations of earlier counts.

The families are listed according to Cave (1956-60),
except that the Smilacaceae and the Krameriaceae are
separated from the Liliaceae and the Leguminosae, respec-
tively. The genera and species are reported alphabetically
within families. For each taxon the following data are
given: (a) an asterisk after the name if this is the first
reported number or if it is at variance with a previous
record; (b) somatic and/or gametic numbers from pre-
meiotic cells and/or PMCs, unless otherwise indicated;
(c) figure number if the chromosomes are illustrated ;
(d) voucher with locality, collector, and citation of her-
barium where deposited; (e) number of plants studied when
more than one; and (f) name of taxonomist who determined
voucher if not by the authors. When pertinent a brief dis-
cussion and reference to earlier work is also added.

Most of the chromosome numbers were obtained from
PMC meiosis and premeiotic mitosis of immature buds fixed
in the field. Occasionally, root tips, pretreated for 2-3 hours
in paradichlorobenzene, and pollen mitosis served as the
basis for the chromosome reports. For some species plants
from more than one locality were sampled and when possible
more than one from each population. All material was fixed
in modified Carnoy’s (4: 3: 1) and stained in either 1%
acetic-orcein or 1% iron aceto-carmine. Chromosomes were
drawn with the aid of a camera lucida originally at X 2300
and reduced by ca. 2/5 in reproduction.

CYPERACEAE

Eleocharis obtusa (Willd.) Schultes*: n = 5 (pollen mitosis), fig. 1.
Texas. Panola Co., Lake Murvaul, Hull 7 (SMU).

XYRIDACEAE

Xyris elliottii Chapm.*: (1) n= 9. Florida. Citrus Co., 0.4 miles
N of Citrus-Hernando Co. line, Lewis 5653 (VPI). (2) n= 9, fig. 2.
Florida. Glades Co., 8.6 miles SE of Palmdale, Lewis 5684 (vPI).
Determined by Dr. Robert Kral.

1962] Lewis, Stripling & Ross — Chromosome Numbers 149

X. torta Sm.: n = 9, fig. 3. Virginia. Augusta Co., ca. 2.5 miles
SW of Sherando, Kral 13394 (vPI). Determined by the collector.
Verification of Lewis (1961) for a plant allied to X. torta but thought
to represent a new taxon.

COMMELINACEAE

Commelina erecta L. var. angustifolia (Michx.) Fern.*: n = 30, fig.
4. Texas. Nacogdoches Co., 1.4 miles NW of Nacogdoches, Stevens 10
(SMU).

C. diffusa Burm. f.: 2n = 30, fig. 5. Texas. Nacogdoches Co.,
Nacogdoches, Stevens 17 (SMU, TEX). Verification of Simmonds
(1954), but not that of Morton (1956) who found 2n = 28 chromosomes
for west African material.

C. virginica L.*: n = 30. Texas. Nacogdoches Co., Angelina River
and Hwy. 21, Stevens 16 (SMU, TEX).

Tradescantia navicularis Ortgies*: n = 24 (first pollen mitosis),
fig. 6. MEXICO. Nuevo Leon. 15.2 miles E of Hwys. 57 and 60 junction,
Lewis 5756 (SMU). Determined by Dr. Harold E. Moore, Jr. Darl-
ington (1929) reported » — 16 (4x) for this taxcn, which suggests
that our material is part of a hexaploid race or that Darlington's
plant was a second species. This is the first record of T. navicularis
from Nuevo Leon.

T. ohiensis Raf.: n — 12 (first pollen mitosis). Arkansas. St.
Francis Co., 1.5 miles N of Goodwin, Lewis 5611 (SMU). Verification
of the 4x race recorded by Anderson and Sax (1936) under T.
canaliculata Raf.

T. reverchoni Bush: n — 12 (first pollen mitosis). Texas. Nacog-
doches Co. 2 miles NE of Nacogdoches, Stevens 9 (SMU, TEX).
Verification of Brown et al. (1951);

SMILACACEAE
Smilax glauca Walt.*: 2n — 32. Texas. Nacogdoches Co., Angelina
River and Hwy. 21, Ross 70 (SMU, TEX). Jensen (1937) recorded
n = 14 chromosomes for S. glauca which on the basis of this count,
and for others of closely related species, seems to be in error.
S. bona-nox L.*: 2n — 32, fir 7. Texas. Panola Co., 2.3 miles NE of
Pinehill, Lewis 5718 (SMU, TEX); 2 plants studied.

AMARYLLIDACEAE
Ipheion uniflorum (Graham) Raf. 2n = 12 (root tips). Bulbs
received from Dr. L. H. Shinners, originally from the Oakhurst
Gardens, Arcadia, California, Lewis 5434 (SMU). Verification of Saez
(1949) under Brodiaea.
LORANTHACEAE
Phoradendron bolleanum (Seem.) Eichl.*: 2n = 28. Texas. Brewster
Co., Big Bend National Park, Lewis & Oliver 5452 (SMU). Without

150 Rhodora [Vol. 64

pretreatment the metaphase chromosomes vary in length from 13.4 y
to 33.6 y. for the longest pair. Bowden and Speese (1957) noted the
exceptional length of chromosomes for P. flavescens [= P. serotinum
(Raf.) M. C. Johnst.] even after pretreatment for 24 hours.

POLYGONACEAE
Brunnichia cirrhosa Gaertn.*: n = 24, fig. 8. Texas. Nacogdoches
Co., 1 mile E of Martinsville, Oliver 62 (SMU).

PHYTOLACCACEAE
Phytolacca americana L.: n = 18. Texas. Nacogdoches Co., Stephen
F. Austin Experimental Forest, Ross 82 (ASTC). Verification of
2n — 36 by Suzuka (1950).
PORTULACACEAE
Claytonia virginica L.: (1) n — 12, fig. 9. Kentucky. Warren Co.,
0.7 miles NW of Petros, Lewis 5606 (SMU); 2 plants studied. (2)
n — 36, fig. 10. Kentucky. Green Co., 1.4 miles NW of Greensburg,
Lewis 5602 (sMU); 2 plants studied. PMC meioses of both plants
from the first population were regular, but those from the second
population were highly abnormal in behavior. The following config-
urations from two plants were noted at metaphase I: 351 > 21.
341 + 41,831 61, 3211+ 81, 3411 + 1 rr +11, 3380 7 lum FRI
3011 + 1101 + 71, and 3211 + 11v + 41. These variations in chromo-
some number and in meiotic behavior are not surprising in view of the
results of Rothwell (1959) and of Lewis (1962).

RANUNCULACEAE
Aquilegia canadensis L. var. australis (Small) Munz*: 2n = 14,
fig. 11. Florida. Jackson Co., 3 miles N of Marianna, Mitchell 27
(FsU) ; 2 plants studied. This variety is known only from the Marianna
Red Hills region of Florida.

Isopyrum. biternatum (Raf.) T. & G.*: 2n = 14, fig. 12. Florida.
Jackson Co., 3 miles N of Marianna, Mitchell 1 (FSU) ; 2 plants studied.
MENISPERMACEAE

Cocculus carolinus (L.) DC.: n = 39. Texas. Nacogdoches Co.,
Nacogdoches, Ross 42 (SMU). Verification of Bowden's (1945) 2n — 78.
CRUCIFERAE

Cardamine bulbosa (Schreb.) B.S.P.*: 2n — 64. Texas. Shelby Co.,
2 miles SW of Timpson, Lewis 4998 (SMU).

Lesquerella purpurea (Gray) Wats.*: n — 9, 2n — 18, fig. 13. Texas.
Brewster Co., Big Bend National Park, Ross 50 (SMU, TEX).

SAXIFRAGACEAE

Ribes echinellum (Cov.) Rehd.* (Grossularia echinella | Cov.) :
2n — 16, fig. 14. Florida. Leon Co. Tallahassee, cultivated, Stripling
779 (FSU). The Florida gooseberry is endemic to the Lake Miccosukee
region in Jefferson Co.; this plant was studied after transplanting to
Tallahassee.

1962] Lewis, Stripling & Ross — Chromosome Numbers 151

LEGUMINOSAE

Apios americana Medic.*: n — 11, fig. 15. Texas. Nacogdoches Co.,
Nacogdoches, Redus 15 (SMU); 2 plants studied. This number does
not agree with that found by Atchison (1949), and reported under
the synonym A. tuberosa Moench., and undoubtedly her number of
2n — ca. 40 refers to another taxon. The basic number for Apios is
x = 11.

Centrosema virginianum (L.) Benth.*: (1) n = 9, fig. 16. Texas.
Naccgdoches Co., Nacogdoches, Redus 1 (SMU, TEX); 2 plants studied.
(2) n=9. Texas. Nacogdoches Co., Angelina River and Hwy. 21,
Redus 13 (TEX): 2 plants studied. These counts add a new basic
number of « = 9 to the genus, a number rare in the Leguminosae.

Desmodium canescens (L.) DC.: n = 11, fig. 17. Texas. Nacogdoches
Co., Nacogdoches, Redus 9 (SMU, TEX). Verification of Young (1940).

Erythrina herbacea L.: 2n = 42. Texas. Nacogdoches Co., Nacog-
doches, Lewis & Oliver 5234 (ASTC). Verification of Atchison (1947).

Galactia macreei Curtis*: n — 10, fig. 18. Texas. Nacogdoches Co.,
Nacogdoches, Redus 6 (SMU).

KRAMERIACEAE

Krameria lanceolata Torr.: n — 6. Texas. San Patricio Co., 5 miles
SE of Mathis, Lewis & Jones 5589 (SMU). Verification of Turner
(1958). At anaphase I, the longest chromosome measured 24.6 y, the
shortest 10.0 » and the six averaged 17.3 u. As noted by Turner,
these chromosomes are among the longest known in the dicotyledons
although those of Phoradendron bolleanum given. above are even
greater in length.

EUPHORBIACEAE
Cnidoscolus texanus (Muell. Arg.) Small*: n — 18, fig. 19. Texas.
Nacogdoches Co., Nacogdoches, Ross 66 (SMU).
Croton capitatus Michx. var. lindheimeri (Engelm. & Gray) Muell.
Arg.*: 2n —20 (root tips), fig. 20. Texas. Nacogdoches Co., Nacog-
doches, Lewis 4994 (ASTC).

CYRILLACEAE

Cliftonia, monosepala (Lam.) Sarg.*: (1) n = 10, 2n = 20, fig. 21.
Florida. Leon Co., 15 miles SW of Tallahassee, Stripling 100? (FSU);
2 plants studied. (2) 2n — 20. Florida. Jefferson Co., 1.5 miles SW
of Wacissa, Stripling 1004 (FSU); 2 plants studied. (3) n = 10.
Florida. Leon Co., Branch Bay, near Silver Lake, Godfrey 60543
(FSU). (4) 2n — 20. Florida. Wakulla Co., Spring Hill Road, SW of
Tallahassee, Godfrey 60544 (FSU). There are no previous counts for |
this family other than a report of n = 20 in most cells of Cyrilla
racemiflora L. (Thomas, 1960). Thomas noted fast deterioration of
fixed buds and this was found to be true for Cliftonia. Our best
results were obtained no later than 2-4 days after fixation.

152 Rhodora [Vol. 64

GUTTIFERAE

Hypericum sp. *: n = 9, 2n = 18, fig. 22. Florida. Washington Co.,
Adams & Tyson 833 (FSU); 2 plants studied. A new specific name
will be given to this collection by Dr. P. Adams, DePauw University.

H. microsepalum (T. & G.) Gray* [Crookea microsepalum (T. & G.)
Small]: (1) n — 9, 2n — 18, fig. 23. Florida. Wakulla Co., 4 miles N
of Sopchoppy, Stripling 772 (Fsu). (2) 2n —18. Florida. Leon Co.,
6 miles S of Tallahassee, Stripling 786 (FSU).

VIOLACEAE

Viola walteri House: 2n — 20 (root tips), fig. 24. Texas. Nacog-
doches Co., Nacogdoches, Lacey 3 (SMU, ASTC). Verification of Gershoy
(1934).

TURNERACEAE

Piriqueta glabrescens Small*: n = 7, fig. 25 as 611 + 21. Florida.
Collier Co., 7.1 miles E of Napes, Osborne 62 (FSU, SMU); 3 plants
studied, mostly with 711. Determined by Dr. C. M. Rogers.

P. tomentosa HBK.*: n — 7, fig. 26. Florida. Monroe Co., Big Pine
Key, Osborne 66 (FSU); 3 plants studied. Determined by Dr. C. M.
Rogers.

LYTHRACEAE

Rotala ramosior (L.) Koehne*: n — 16, fig. 27. Texas. Angelina

Co., Angelina River and Hwy. 59, Lewis & Oliver 5273 (SMU).

PRIMULACEAE

Samolus parviflorus Raf.*: n = 13, 2n = 26, fig. 28. Louisiana. St.
Mary Par., 8.7 miles NW of Boyeau Vista, Oliver 261 (sMU) ; 2 plants
studied.

STYRACEAE

Halesia diptera Ellis. var diptera*: 2n = 24, fig. 29. Florida. Leon
Co., Tallahassee, cultivated, Stripling 732 (FSU).

H. diptera Ellis. var. magniflora Godfrey*: 2n — 24, fig. 30. Florida.
Leon Co., Tallahassee, cultivated, Stripling 730 (FSU).

H. tetraptera Ellis.* [H. parviflora Michx., not H. carolina (tetrap-
tera) as listed in Darlington & Wylie (1956), cf. Godfrey (1958) ]:
2n —24, fig. 31. Florida. Leon Co., Tallahassee, cultivated, Stripling
716 (FSU).

OLEACEAE

Menodora scabra Gray*: n — 11, fig. 32. Texas. Brewster Co., Big
Bend National Park, Ross 52 (SMU, TEX). This number does not agree
with that of Taylor (1945) who found 2n = 44 chromosomes. Perhaps
2x and 4x races exist for the species.

LOGANIACEAE
Cynoctonum mitreola (L.) Britt.*: n = 10, 2n = 20. Texas. Liberty

1962] Lewis, Stripling & Ross — Chromosome Numbers 153

Co., 4 miles E of Cleveland, Ross 73 (SMU, TEX); 2 plants studied.
Determined by Dr. L. H. Shinners.

GENTIANACEAE
Sabatia campestris Nutt.*: (1) n = 13, fig. 33. Texas. Brazos Co.,
College Station, Lewis 5494 (SMU). (2) 2n = 26. Texas. Nacogdoches
Co., Nacogdoches, Lewis & Oliver 5233 (SMU).

CONVOLVULACEAE

Bonamia humistrata (Walt.) Gray*: n = 14, fig. 34. Texas. Panola
Co., Sabine River and Hwy. 59, Lewis & Oliver 5318 (SMU). Some
PMCs were observed with 13 11 + 21 and still others with a trivalent
at metaphase I.

B. pickeringii (Torr.) Gray*: (1) n —14, fig. 35. Texas. Rusk Co.,
3.1 miles N of Cushing, Lewis & Oliver 5279 (sMU). At least 50%
of the PMCs had 1311 + 21. (2) 2n = 28. Texas. Nacogdoches Co.,
0.5 miles W of Nacogdoches, Lewis & McDaniel 5509 (SMU).

Jacquemontia abutiloides Benth.*: 2n — 18. MEXICO. Baja California
Sur. 3 km. S of Miraflores, Lewis 5339 (SMU); 2 plants studied.
Determined by Dr. L. H. Shinners.

LABIATAE

Pycnanthemum albescens T. & G.*: n = 33-36. Texas. Liberty Co.,
2 miles E of Cleveland, Ross 61 (SMU).

Salvia reflexa, Hornem.*: n — 10, fig. 36. MEXICO. Nuevo Leon. 9.9
miles E of Hwys. 57 and 60 junction, Lewis 5750 (SMU, TEX). Deter-
mined by Dr. L. H. Shinners.

Scutellaria cf. integrifolia L.*: 2n = 32, fig. 37. Florida. Sarasota
Co., 12 miles E of Sarasota, Stripling 754 (FSU); 2 plants studied.

Stachys tenuifolia Willd.*: n = 16, fig. 38. Texas. Nacogdoches Co.,
Stephen F. Austin Experimental Forest, Ross 56 (SMU, TEX).

SOLANACEAE

Nicotiana glauca Graham: 2n = 24. MEXICO. Baja California Sur.
1.4 km. W of San José del Capo, Lewis 5354 (SMU). Verification of
Goodspeed (1923).

SCROPHULARIACEAE

Bacopa monnieri (L.) Pennell*: 2n — 64, fig. 39. Texas. Harris Co.,
2 miles E of Humble, Lewis 5497 (SMU).

Buchnera americana, L.*: 2n — ca. 42. Texas. Jasper Co., 50 yards
S of Hwys. 96 and 1004 junction, Lewis 5623 (SMU).

Gratiola ramosa Walt.*: (1) n — 7, 2n — 14, fig. 40. Florida. Levy
Co., 1 mile SW of Otter Creek, Stripling 734 (FSU) ; 2 plants studied.
(2) n— 7, 2n = 14. Florida. Wakulla Co., 3 miles E of Carabelle,
Stripling 781 (FSU); 2 plants studied.

G. aurea Muhl.*: 2n = 28, fig. 41. Georgia. Cook Co., Little River
bridge and Hwy. 76, Adams 588 (FSU); 3 plants studied.

154 Rhodora [Vol. 64

G. brevifolia Raf.*: (1) n = 14, 2n = 28, fig. 42. Florida. Leon Co.,
7 miles SW of Tallahassee, Stripling 794 (FSU); 3 plants studied.
(2) 2n = 28. Florida. Gadsden Co., 3 miles W of Ochlockonee River
bridge and Hwy. 90, Stripling 777 (Fsu). (3) 2n = 28. Florida.
Jefferson Co., 4.5 miles E of Monticello, Stripling 795 (FSU); 2 plants
studied. (4) n = 14, 2n = 28. Texas. Liberty Co. 4 miles E of
Cleveland, Ross 77 (SMU, FSU); 2 plants studied.

G. virginiana L.*: n —8, 2n — 16, fig. 43. Florida. Leon Co., 6 miles
SE of Tallahassee, Stripling 705 (FSU); 3 plants studied.

These counts represent a part of a detailed study of the Gratiolas
(s. 1.) now in progress. A new basic number of x — 7 is reported for
the genus and includes G. ramosa, G. aurea, and G. brevifolia in the
section Gratiolaria while G. virginiana in the section Nibora belongs
to a second line with x — 8 chromosomes.

Lindernia dubia (L.) Pennell*: n — 16, 2n — 32, fig. 44. Texas. San
Augustine Co., 8.2 miles SE of San Augustine, Lewis et al. 5500 (FSU).

Mecardonia acuminata (Walt) Small*: 2n = 42+ 2. Texas.
Cherokee Co., Angelina River and Hwy. 21, Lewis et al. 5505 (SMU).

Scoparia dulcis L.*: n = 10, fig. 45. Texas. Liberty Co., 2 miles
E of Cleveland, Ross 62 (SMU, TEX). Raghaven & Srinivasan (1940)
recorded a tetraploid number of 2n = 40.

Tragiola pilosa (Michx.) Small & Pennell*: (1) 2n = 22, fig. 46.
Florida. Taylor Co., 9 miles W of Perry, Stripling 883 (FSU) ; 2 plants
studied. (2) 2n — 22, Florida. Levy Co., 11 miles NW of Dunnellon,
Stripling 876 (FSU).

Veronica persica Poiret: 2n — 28, fig. 47. Florida. Leon Co., Talla-
hassee, Stripling 788 (FSU); 3 plants studied. Beatus (1936) recorded
this number for V. persica in Europe, but there is some doubt as to
the exact species he examined. Although not previously examined
cytologically from the United States, our data verify counts from
Canadian (Mulligan, 1959) and Icelandic (Lóve & Love, 1956)
populations.

LENTIBULARIACEAE
Utricularia inflata Walt.*: (1) n — 9, 2n = 18, fig. 48. Florida.
Leon Co., 1 mile E of Ochlockonee River and Hwy. 90, Godfrey 59374

Fic. 1-14. Chromosomes of angiosperms, X 1350. Fig. 1. Eleocharis obtusa, Hull 7,
n = 5, with 3 degenerating nuclei; fig. 2. Xyris elliottii, Lewis 5684, n = 9; fig. 3.
X. torta, Kral 13394, n — 9; fig. 4. Commelina erecta var, angustifolia, Stevens 10,
n = 30; fig. 5. C. diffusa, Stevens 17, 2n = 30; fig. 6. Tradescantia navicularis,
Lewis 5756, n = 24; fig. 7. Smilax bona-now, Lewis 5718, 2n = 82; fig. 8. Brunnichia
cirrhosa, Oliver 62, n = 24; fig. 9. Claytonia virginica, Lewis 5606, n = 12; fig. 10.
C. virginica, Lewis 5603, n = 36 (29 II + 814 2 III) ; fig. 11. Aquilegia canadensis var.
australis, Mitchell 27, 2n = 14; fig. 12. Isopyrum biternatum, Mitchell 1, 2n = 14;
fig. 13. Lesquerella purpurea, Ross 50, 2m — 18; fig. 14. Ribes echinellum, Stripling
779, 2n — 16.

1962] Lewis, Stripling & Ross — Chromosome Numbers 155

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156 Rhodora [Vol. 64

(Fsu); 3 plants studied. (2) 2n = 36, fig. 49. Florida. Leon Co., 4
miles W of Tallahassee, Reinert 2 (FSU); 2 plants studied. The
previous numbers reported for Utricularia (cf. Darlington & Wylie,
1956) are all approximations, viz., 2n — 36-40 or ca. 40 chromosomes.
Our data establish x = 9 as a basic number for the genus and also
record U. inflata as having 2x and 4x races.

ACANTHACEAE

Justicia lanceolata (Chapm.) Small*: (1) n= 14, fig. 50. Texas.
Angelina Co., Angelina-Nacogdoches Co. line and Hwy. 301, Lewis &
Oliver 5320 (SMU). (2) n — 14. Texas. Angelina Co., Angelina River
and Hwy. 7, Lewis & Oliver 5325 (SMU). (3) n= M +1, 2n = 29,
fig. 51. Texas. Panola Co., 0.7 miles NE of Sabine River and Hwy. 59,
Lewis & Oliver 5309 (SMU). The single plant from the third locality
consistently had 1411 + 11 at meiosis I and an extra chromosome in
the mitotie plates. The chromosome was small but not heteropycnotic.

CAMPANULACEAE

Sphenoclea zeylanica Gaertn.*: n = 12. Texas. Nacogdoches Co.,
Angelina River and Hwy. 59, Oliver 357 (SMU, TEX). Determined by
Dr. L. H. Shinners.

COMPOSITAE

Cirsium horridulum Michx.*: (1) 2n = 34, fig. 52. Florida.
Franklin Co., near Alligator Point, Godfrey 59335 (Fsu); 2 plants
studied. (2) 2n = 34. Florida. Leon Co., 21 miles SW of Tallahassee,
Stripling 697 (FSU); 2 plants studied.

Elephantopus caroliniamus Raeusch.: n — 11. Texas. Nacogdoches
Co., Angelina River and Hwy. 21, Ross 72 (SMU, TEX). Verification
of Baldwin & Speese (1955) as 2n = 22.

Fig. 15-38. Chromosomes of angiosperms. X 1350. Fig. 15. Apios americana, Redus
15, n= 11; fig. 16. Centrosema virginianum, Redus 1, n — 9; fig. 17. Desmodium
canescens, Redus 9, m = ll; fig. 18. Galactia macreei, Redus 6, n = 10; fig. 19.
Cnidoscolus texanus, Ross 66, n — 18; fig. 20. Croton capitatus, Lewis 4994, 2n — 20;
fig. 21. Cliftonia monosepala, Stripling 1007, n — 10; fig. 22. Hypericum sp., Adams
& Tyson 833, m —9; fig. 23. H. microsepalum, Stripling 772, n = 9; fig. 24. Viola
walteri, Lacey 3, 2n = 20; fig. 25. Piriqueta glabrescens, Osborne 62, n=T7
(611 + 21); fig. 26. P. tomentosa, Osborne 66, n = T; fig. 27. Rotala ramosior, Lewis
& Oliver 5273, m — 16; fig. 28. Samolus parviflorus, Oliver 261, n — 13; fig. 29.
Halesia diptera var. diptera, Stripling 732, 2m — 24; fig. 30. H. diptera var.
magniflora, Stripling 730, 2m = 24; fig. 31. H. tetraptera, Stripling 716, 2n = 24;
fig. 32. Menodora scabra, Ross 52, n= 11 (1 side anaphase I); fig. 33. Sabatia
campestris, Lewis 5494, n — 13; fig. 34. Bonamia humistrata, Lewis & Oliver 5318,
n = 14; fig. 35. B. pickeringii, Lewis & Oliver 5279, n = 14; fig. 36. Salvia reflexa,
Lewis 5750, n = 10; fig. 87. Scutellaria cf. integrifolia, Stripling 754, 2n = 32; fig.
38. Stachys tenuifolia, Ross 56, n — 16.

1962] Lewis, Stripling & Ross — Chromosome Numbers 157

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158 Rhodora [Vol. 64

Liatris acidota Engelm. & Gray: n — 10. Texas. Liberty Co., 4
miles E of Cleveland, Ross 76 (SMU, TEX). Verification of Gaiser
(1949).

Perityle sp.*: n — 19. MEXICO. Baja California Sur. 14 km. N of
La Paz, Lewis 5351 (SMU).

P. palmeri Wats.*: n — 19, fig. 53. MEXICO. Sonora. 2.8 miles N
of Guaymas, Lewis 5334 (SMU).

Silphium asperrimum Hook.*: (1) n= 8, 2n = 16, fig. 54. Texas.
Smith Co., 1 mile N of Bullard, Stripling 510 (FSU) ; 2 plants studied.
(2) 2n — 16. Texas. Nacogdoches Co., 3 miles S of Nacogdoches,
Stripling 586 (FSU).

S. compositum Michx.*: n = 7, 2n = 14, fig. 55. Florida. Lafayette
Co., between Fletcher and Cross City along Hwy. 351, Stripling 798
(FSU) ; 3 plants studied. — STEPHEN F. AUSTIN STATE COLLEGE, NACOG-
DOCHES, TEXAS, AND FLORIDA STATE UNIVERSITY, TALLAHASSEE.

LITERATURE CITED

ANDERSON, E. AND K. Sax. 1936. A cytological monograph of the
American species of Tradescantia. Bot. Gaz. 97: 433-476.

ATCHISON, E. 1947. Studies in the Leguminosae. I. Chromosome
numbers in Erythrina L. Amer. Jour. Bot. 34: 407-414.

1949. Studies in the Leguminosae. IV. Chromosome

numbers and geographical relationships of miscellaneous

Leguminosae. Jour. Elisha Mitchell Sci. Soc. 65: 118-122.

BALDWIN, J. T. JR. AND B. M. SPEESE. 1955. Chromosomes of
Elephantopus and of Pluchea in the range of Gray’s manual of
botany. Amer. Jour. Bot. 42: 123-125.

1957. Phoradendron

flavescens: chromosomes, seedlings, and hosts. Amer. Jour. Bot.
44: 136-140.

BEATUS, R. 1936. Die Veronica-Gruppe Agrestis der Section Alsinebe
Griseb., ein Beitrag zum Problem der Artbildung 1. Zeitschr.
Indukt. Abst. Ver. 71: 3553-381.

Fics. 39-55. Chromosomes of angiosperms, X 1350. Fig. 39. Bacopa monnieri,
Lewis 5497, 2n = 64; fig. 40. Gratiola ramosa, Stripling 734, 2n = 14; fig. 41. G.
aurea, Adams 588, 2n = 28; fig. 42. G. brevifolia, Stripling 794, n =14; fig. 43. G.
virginiana, Stripling 705, 2n = 16; fig. 44. Lindernia dubia, Lewis et al. 5500, n =
16; fig. 45. Scoparia dulcis, Ross 62, n = 10; fig. 46. Tragiola pilosa, Stripling 883,
2n = 22; fig. 47. Veronica persica, Stripling 788, 2n = 28; fig. 48. Utricularia inflata,
Godfrey 59374, n = 9; fig. 49. U. inflata, Reinert 2, 2n = 36; fig. 50, Justicia lanceola-
ta, Lewis & Oliver 5320, n = 14; fig. 51. J. lanceolata, Lewis & Oliver 5311, n = 14
+1 (14 1 +11); fig. 52. Cirsium horridulum, Godfrey 59335, 2n = 34; fig. 53. Perityle
sp., Lewis 5351, n = 19; fig. 54. Silphium asperrimum, Stripling 510, 2n = 16; fig. 55.
S. compositum, Stripling 798, n = 7.

1962] Lewis, Stripling & Ross — Chromosome Numbers 159

44 45

e 43

160 Rhodora [Vol. 64

Bowben, W. M. 1945. A list of chromosome numbers in higher
plants. II. Menispermaceae to Verbenaceae. Amer. Jour. Bot.
32: 191-201.

Brown, W. V., H. J. PONEWCZYNSKI, AND H. H. SCARBOROUGH, JR. 1951.
A cytological study of Tradescantia reverchoni Bush, Bull. Torrey
Bot. Cl. 78: 66-69.

CAVE, M. S. (editor) 1956-60. Index to Plant Chromosome Numbers
for 1956-60.

DARLINGTON, C. D. 1929. Chromosome behavior and structural
hybridity in the Tradescantiae. Jour. Genet. 21: 207-286.

AND A. P. WYLIE. 1956. Chromosome Atlas
of Flowering Plants. MacMillan Co., New York.

GaAISER, L. O. 1949. Chromosome studies in Liatris. I. Spicatae and
Pycnostachyae. Amer. Jour. Bot. 36: 122-135.

GERSHOY, A. 1934. Studies in North American violets. III. Chromo-
some numbers and species characters. Vt. Agr. Exp. Sta. Bull.
367: 1-91.

GopFREY, R. K. 1958. Some identities in Halesia (Styracaceae).
Rhodora 60: 86-88.

GOODSPEED, T. H. 1923. A preliminary note on the cytology of
Nicotiana species and hybrids. Svensk. Bot. Tidsk. 17:472-478.

JENSEN, H. W. 1937. Meiosis in several species of dioecious mono-
cotyledoneae I. The possibility of sex-chromosomes. Cytologia,
Fujii Jub. Vol. 96-103.

LEWIS, W. H. 1961. Chromosome numbers for three United States
species of Xyris. Southwest. Nat. 6: 99-100.

1962. Aneusomaty in aneuploid populations of Clay-
tonia virginica. Amer. Jour. Bot. (in press)

LóvE, A. AND D. LévE. 1956. Cytotaxonomical conspectus of Icelandic
flora. Act. Hort. Gotob. 20: 65-290.

MoRTON, J. K. 1956. Cytotaxonomic studies on the Gold Coast
species of the genus Commelina Linn. Jour. Linn. Soc. 60: 507-531.

MULLIGAN, G. A. 1959. Chromosome numbers of Canadian weeds.
II. Canad. Jour. Bot. 37: 81-92.

PENNELL, F. W. 1935. The Scrophulariaceae of eastern temperate
North America. Acad. Nat. Sci. Philadelphia, Monographs 1:
1-650.

RAGHAVAN, T. S. AND V. K. SRIMIVASAN. 1940. Studies in the Scro-
phulariaceae. I. The cytology of Angelonia grandiflora C. Morr.
and some related genera. Cytologia 11: 37-54.

ROTHWELL, N. V. 1959. Aneuploidy in Claytonia virginica. Amer.
Jour. Bot. 46: 353-360.

SAEZ, F. A. 1949. Los cromosomas de Brodiaea uniflora. Lilloa 19:
105-110.

SIMMONDS, N. W. 1954. Chromosome behaviour in some tropical
plants. Heredity 8: 139-150.

19692] Hodgdon and Steele — Glandularity in Rubus 161

SUZUKA, O. 1950. Chromosome numbers in pharmaceutical plants I.
Rep. Kihara Inst. Biol. Res. 4:57-58.

TAYLOR, H. 1945. Cyto-taxonomy and phylogeny of the Oleaceae.
Brittonia 5: 337-367.

THoMas, J. L. 1960. A monographic study of the Cyrillaceae. Contr.
Gray Herbarium 186: 1-114.

TURNER, B. L. 1958. Chromosome numbers in the genus Krameria:
evidence for familial status. Rhodora 60: 101-106.

YounG, J. O. 1940. Cytological investigations in Desmodium and
Lespedeza. Bot. Gaz. 101: 839-850.

GLANDULARITY IN RUBUS ALLEGHENIENSIS
PORTER?

A. R. HODGDON AND FREDERIC STEELE

For the past 4 years we have been studying the black-
berries of New Hampshire and nearby parts of Maine and
Vermont. This was started as a floristic study to determine
what taxa occur in New Hampshire and where in the State
each is found.

Fernald (1) quoted in part here, has the following to say
about the classification of Rubus Subgenus Eubatus which
embraces the Blackberries, ‘“‘Taxonomically a most difficult
group. Our few original wide-ranging, essentially unvary-
ing and ancient species have greatly commingled producing
sometimes localized but rapidly spreading offspring . . ."
We have assumed as Fernald apparently did from what he
stated farther on in this long footnote, that certain of the
multitude of species that have been described are of a dif-
ferent order taxonomically from the others.

One of our primary objectives has been to determine
which of our species in New Hampshire may be character-
ized as well-marked. Unquestionably one of the most clearly
defined is the one in question here, Rubus allegheniensis. It
has an extensive range and possesses some taxonomic char-
acters that are recognized readily in the majority of speci-
mens though considerable diversity is displayed in some

1Published with approval of the Director of the New Hampshire Agricultural Experi-
ment Station as Scientific Contribution No. 290.

162 Rhodora [Vol. 64

of its features. We have come to be suspicious of the terms
“unvarying” or “essentially unvarying" as applied to any
wide ranging species of blackberry known to us. A study
based on populations more than on individuals seems to lead
inevitably to a broader and more inclusive interpretation
of the well established taxa in Rubus. Since many of the
less well-marked species are characterized by features that
seem to us to fall within the proper limits of variation of
the better defined ones we suggest that eventually there
should be a marked reduction in the number of accepted
species in the Subgenus Hubatus.

This paper presents evidence to show that Rubus alle-
gheniensis has been interpreted too narrowly in respect to
its glandularity. This should make it possible for more
effective use to be made of Fernald’s key to the Section
Alleghenienses (2) by changing some of the statements
there. At present, with many specimens, the key is quite
unworkable if followed literally, and therefore, in our
opinion, needs to be corrected. The first major division of
the key is concerned with glandularity of the primocanes.
We note the following statements, “axis of primocane
(except sometimes at expanding tip) without or essentially
without stipitate glands on the internodes or among the
prickles” contrasted to “axis of primocane bearing several
to very many long stipitate glands among the prickles.”
Four New England species thus are distinguished from R.
allegheniensis by having glandular primocanes. Of these
only one, R. ravus Bailey, is stated in the key to have long
cylindric racemes of flowers, in this respect resembling R.
allegheniensis. In using the key let us assume that a speci-
men is found to occur in the Section Alleghenienses; if it
has evident glands on the mature primocanes and in addi-
tion has its flowers in well-developed racemes, the specimen
will be identified in the key as R. ravus rather than R. alle-
gheniensis. One would also have trouble getting by the
first statement in the key in L. H. Bailey's treatment of
Rubus (3) which, in part at least, seems to have served
as a model for the key in Gray's Manual.

1962] Hodgdon and Steele — Glandularity in Rubus 163

For many years we have been confused by these keys, a
major difficulty being that much material, at least in North-
ern and Central New England, has glands on the mature
primocanes and yet in other ways fits R. allegheniensis.
R. ravus is an excessively prickly and glandular taxon of
localized occurrence. We don't at present have convincing
evidence that it is a good species. In herbaria it is repre-
sented from only a few localities but recent field observations
and collections by us indicate it to be of occasional occur-
rence, though it may be widespread. The material to be
discussed below resembles R. ravus only in having glands
on the primocanes and by having more or less elongated
racemes. Except for the glandular primocanes, there would
certainly be no hesitancy about calling all of these specimens
R. allegheniensis.

In 1953 the junior author made some observations of a
colony of R. allegheniensis in Sandwich, New Hampshire.
In the middle of June it was noted that all of the primo-
canes in the colony, which had not then reached full growth,
were heavily glandular. Collections made from the colony
at weekly intervals indicated that the glands tended to dis-
appear as the canes matured. A rather dense pubescence
remained on the primocanes of this particular colony. Since
1951 the senior author frequently has collected specimens
which have the characteristics of R. allegheniensis except
that they have noticeable glands on the primocanes. At first
he was inclined to call these R. ravus but glandular plants
occurred so often and these had such varying numbers of
glands that a very different explanation was suggested.
These observations thus impelled us to undertake a syste-
matic study of gland-occurrence on the primocanes of R.
allegheniensis.

Collections of 10 primocanes were made from each of 9
colonies in different localities in New Hampshire and from
one in southern Maine between the dates of July 15 and
November 11, 1959. These colonies were selected entirely
without regard for glandularity. The colonies were all
vigorous, consisting of well-grown primocanes and flori-

164 Rhodora [Vol. 64

canes intermixed. Since the canes in each colony grew in
close proximity, it was thought that in many of the colonies
the canes were genetically similar, having arisen from the
same single rootstock. Sections about 30 cm. long were
collected; these were taken from about the middle of full-
sized primocanes. The attempt was made to distribute the
collecting somewhat uniformly throughout each colony. The
stands ranged in size from 20 or 30 square metres down
to stands that barely yielded 10 primocanes for our study.

The results of counts of glands are shown in Fig. 1; 35%
of the plants fell in the class of less than 100 glands per
decimeter and therefore, compared with heavily glandular
stems, might be considered as essentially without glands
though it is to be noted that they are still fairly obvious
under 10X magnification down to fewer than 50 per deci-
meter. The matter of glandularity then can become im-
portant subjectively, depending on the diligence with which
one searches for glands, also the quality of lens used and
the interpretation of what is meant by the phrase “essen-
tially without glands." Assuming that any condition fewer
than 100 glands per decimeter may fit the requirement of
R. allegheniensis of relative glandlessness, we were left
with 65% of the canes, including at least 2 from each colony
and all of them from one, which had more than 100 glands
per decimeter — certainly enough to notice and to create
doubt and confusion when trying to use the key. In 7 of
the colonies one or more canes had more than 500 glands
per decimeter, a situation which could hardly be ignored
by any critical observer. Of considerable interest is the
variation in glandularity within any particular colony. Some
of the colonies almost certainly were from common root-
stocks. Yet all displayed some differences between canes
and 6 of the colonies had markedly different numbers of
glands on most of their canes. Many questions and unsolved
problems are suggested by these data.

Because of some of the questions raised in this study, a
canvass was made of readily available herbarium specimens
to find out if the results noted in New Hampshire and

1962] Hodgdon and Steele — Glandularity in Rubus 165

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Fig. 1. Frequency distribution of numbers of glands in Rubus allegheniensis.

166 Rhodora [Vol. 64

southern Maine might be expected elsewhere in the range
of the species. Unfortunately, many herbarium specimens
of blackberries are quite useless for critical study, lacking
either floricanes or primocanes and often without growth-
habit notes. In addition, young primocanes collected at the
time of flowering tend to give an unfair impression of
glandularity. Nonetheless a sufficient number of good speci-
mens were available in the Gray Herbarium and the herb-
arium of the New England Botanical Club, as well as the
University of New Hampshire collection and the private
collection of the junior author to give us a more general
picture. Table 1 summarizes the data on primocane glands
as best it could be determined from the aforementioned
herbaria. From outside of New England and particularly
from the southern parts of the range the specimens avail-
able were few and mostly inadequate in that frequently
they lacked good primocanes. Similar field and herbarium
studies of R. allegheniensis should be carried out in areas
other than New England to provide a more complete analysis
of glandularity in this species. A glance at some specimens
from the Maritime Provinces of Canada indicated that we
might expect to find considerable glandularity there.

Table 1. — Glandularity in Herbarium Material of R. allegheniensis
Numbers of primocanes in classes based on
number of glands per decimeter.

Location more than
0-99 100-199 200-299 300-399 400-499 500

Maine 11 3 2 0 1 5

New Hampshire 38 10 6 8 5 10

Other New

England States 5 5 — 4 2 3

States other than

New England 21 2 2 1 3 1

From the table we see that slightly more than half of the
primocanes are relatively glandless with fewer than 100
glands per decimeter, this being considerably more than in
our population samples. It is evident however that, in most
parts of the range, there would be specimens to cause per-
plexity and confusion when using the key. In states other

1962] Hodgdon and Steele — Glandularity in Rubus — 167

than New England most of the heavily glandular specimens
came from the northern tier of states. If diminished glandu-
larity is partly a function of development, it might be that
the longer growing period of the more southern parts of the
range may contribute to a reduction in glands. Though not
shown in the table, entirely glandless specimens constituted
only a minor proportion of the primocanes in herbarium
specimens.

The var. neoscoticus Fernald of Rubus allegheniensis is
interpreted by Bailey (4) as being glandular, though this
character is ignored in Fernald’s handling of it in Gray's
Manual. Our glandular specimens definitely do not fit this
variety.

In the Harvard Herbarium an examination was made of
sheets of the supposedly glandular species of Section Alle-
ghenienses including R. ravus Bailey, R. glandicaulis
Blanch., R. frondisentis Blanch. and R. sceleratus Brainerd.
The typical plants of these were not only heavily glandular
with 1,500 or, more frequently, 2 or 3,000 or more glands
per decimeter but differed also in character of armature, the
prickles in some cases being thin and acicular and in others
broad-based but very abundant.

Bailey in Gentes Herbarium makes a more satisfactory
separation of R. ravus and the other very glandular species
than does Fernald in that he takes the accompanying arma-
ture into consideration. As a substitute for the primary
divisions of the key to the Section Alleghenienses (2), we
suggest that the following be used:

Mature primocanes with none or few glands or only moderately
glandular (0-1200 per dm.) along with few to many stiff broad-based
prickles.

Mature primocanes with numerous glands (1500-5000 per dm.)
along with many bristles or slender prickles or exceptionally broad-
based prickles, the prickles if broad-based, abundant and almost con-
tiguous.

Many of the taxa in the very glandular group appear to
be hybrids with an admixture of R. setosus or to show intro-

gression with that species.

168 Rhodora [Vol. 64

Specimens of primocane and voucher material used in
the population studies have been placed in the University
of New Hampshire Herbarium.

The authors wish to thank the curators of the Harvard
Herbarium and the New England Botanical Club Herbarium
for their permission to consult these collections freely.
— DEPARTMENT OF BOTANY, UNIVERSITY OF NEW HAMPSHIRE,
DURHAM, NEW HAMPSHIRE AND ST. MARY’S-IN-THE-MOUN-
TAINS, LITTLETON, NEW HAMPSHIRE.

LITERATURE CITED

1. FERNALD, M. L. 1950. Gray’s Manual, 8th edition (footnote on p.

822).

2. . 1950. Gray's Manual, 8th edition (footnote on p.
853).

3. BAILEY, L. H. 1944. Gent. Herb. V. 508-512.

4. . 1944. Gent. Herb. V. 509.

FIFTEENTH REPORT OF
THE COMMITTEE ON PLANT DISTRIBUTION

The fourteenth report included the Dicotyledoneae from
Papaveraceae through Platanaceae. The present report
deals with the genus Carex of the Cyperaceae.

The data for these reports have been compiled from the
material found in the herbarium of the New England Botan-
ical Club, in the Gray Herbarium, and in the herbarium of
the Peabody Museum at Salem.

PRELIMINARY LISTS OF NEW ENGLAND PLANTS — XL

The sign + indicates that an herbarium specimen has
been seen, the sign — that a reliable printed record has been
found and the sign * is used for those plants which are not
native in the New England area.

CYPERACEAE
Me. N.H. Vt. Mass. R.I. Conn.
Carex abdita Bickn. + + ao + + +
Carex abscondita Mackenz. de + + +
*Carex acutiformis Ehrh. +

1962] Committee on Plant Distribution 169

Me. N.H. Vt. Mass. R.I. Conn.
Carex adusta Boott +
Carex aenea Fern. +
Carex aestivalis M. A. Curtis
Carex alata T. & G.
Carex albolutescens Schwein.
Carex albursina Sheldon
Carex alopecoidea Tuckerm. +
Carex amphibola Steud. var. rigida
(Bailey) Fern.
Carex amphibola var. turgida Fern.
Carex angustior Mackenz.
Carex annectens Bickn.
Carex annectens var. xanthocarpa
(Bickn.) Wieg.
Carex aquatilis Wahlenb. var. altior
(Rydb.) Fern.
Carex arcta Boott
Carex arctata Boott
Carex argyrantha Tuckerm.
Carex artitecta Meckenz.
Carex artitecta var. subtilirostris
F. J. Herm.
Carex atherodes Spreng.
Carex atlantica Bailey
Carex atratiformis Britt.
Carex aurea Nutt.
Carex Backii Boott
Carex Baileyi Britt.
Carex Barrattii Schwein. & Torr.
Carex Bebbii Olney
Carex Bicknellii Britt.
Carex Bigelowii Torr.
Carex blanda Dew.
Carex brevior (Dew.) Mackenz.
Carex bromoides Schkuhr.
*Carex brunnea Thunb.
Carex brunnescens (Pers.) Poir.
Carex brunnescens var. sphaerostachya
(Tuckerm.) Kükenth.
Carex bullata Schkuhr.
Carex Bushii Mackenz.
Carex Buxbaumii Wahlenb.
Carex canescens L.
Carex canescens var. disjuncta Fern.
Carex canescens var. subloliacea Laestad.
Carex capillaris L.
Carex capillaris var. major Blytt
Carex capitata L.
*Carex caryophyllea Lat.
Carex castanea Wahlenb.
Carex cephalantha (Bailey) Bickn.
Carex cephaloidea Dew.
Carex cephalophora Muhl.
Carex chordorrhiza L.f.
Carex Collinsii Nutt.
Carex communis Bailey

EE
EE

++ ++
++

+++

+ ++++
+++
=H

+++ + + +++ ++

TRY
++++
TERME
+++

-H

TL
|
-H

+++ ++ ttt + + F+t]t+ + +444 ++++++
+++ +
+++ +++

[+ + ++++++ +++++

++ + ++
++ + +
++ ++++

HHHH + HHHH ++ + F4+4+4+4+4+ ++H+++
+ +4+4+4

+ Ft + +++ ++ + ++ +++
4

+ +++++

+ +++ +

++ +/+

++ +444

170 Rhodora

Carex comosa Boott
Carex conoidea Schkuhr.
Carex convoluta Mackenz.
Carex X copulata (Bailey) Mackenz,
Carex Crawei Dew.
Carex Crawfordii Fern.
Carex crinita Lam.
Carex crinita var. brevicrinis Fern.
Carex crinita var. gynandra (Schwein.)
Schwein. & Torr.
Carex crinita var. minor Boott
Carex crinita var. Porteri (Olney) Fern.
Carex crinita var. simulans Fern.
Carex cristatella Britt.
Carex cumulata (Bailey) Mackenz.
Carex Davisii Schwein. & Torr.
Carex debilis Michx.
Carex debilis var. interjecta Bailey
Carex debilis var. Rudgei Bailey
Carex debilis var. strictior Bailey
Carex deflexa Hornem.
Carex demissa Hornem.
Carex Deweyana Schwein.
Carex diandra Schrank
Carex digitalis Willd.
Carex disperma Dew.
Carex eburnea Boott
Carex elachycarpa Fern.
Carex Emmonsii Dew.
Carex exilis Dew.
Carex festucacea Schkuhr.
Carex flaccosperma Dew. var. glaucodea
(Tuckerm.) Kükenth.
Carex flava L.
Carex flava var. fertilis Peck
Carex flava var. gaspensis Fern.
Carex foenea Willd.
Carex folliculata L.
Carex formosa Dew.
Carex Garberi Fern. var. bifaria Fern.
Carex gracilescens Steud.
Carex gracillima Schwein.
Carex granularis Muhl.
Carex granularis var. Haleana (Olney)
Porter
Carex Grayii Carey
Carex Grayii var. hispidula Gray
Carex gynocrates Wormsk.
Carex X Hartii Dew.
Carex Haydenii Dew.
Carex hirsutella Mackenz.
*Carex hirta L.
Carex hirtifolia Mackenz.
Carex Hitchcockiana Dew.
Carex hormathodes Fern.
Carex hormathodes X straminea

+++ +4+4++4+4++4+ + + ++++

+ + + +4444

+ ++ +++

+++ +4445

z

++ +++

+++ ++

[++ +++ +4 +4+4+4+4+4+4+4++4+

++

E

++

++ ++4++4+4++4+4+4+4+/4+ +++++

++ +++ +4+44+44|

++ +++

++ +4+++8

z
m
th
z

+H +++ +++ +++ +++ ++H+++H +++ F444 ++ +++ 4444+

GREGG

[Vol. 64
R. I. Conn.
+ +
"I +
+ T

+
+
+
-+ +
+ +
+ +
+
4-
uu
4-
T +
+
— +
+
+ E
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+ +
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1962]

Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
*Carex
Carex
Carex
Carex
Carex

Committee on Plant Distribution

Houghtonii Torr.

Howei Mackenz.

hystricina Muhl.

incomperta Bickn.

interior Bailey

intumescens Rudge

intumescens var. Fernaldii Bailey
Josselynii (Fern.) Mackenz.
katahdinensis Fern.

X Knieskernii Dew,

lacustris Willd.

laevivaginata (Kükenth.) Mackenz.
lanuginosa Michx.

lasiocarpa Ehrh. var. americana Fern.

laxiculmis Schwein.

laxiflora Lam.

lenticularis Michx.

lenticularis var. albi-montana Dew.
lenticularis var. Blakei Dew.
leporina L.

leptalea Wahlenb.

leptonervia Fern.

limosa L.

livida (Wahlenb.) Willd. var.

Grayana (Dew.) Fern.

Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
Carex
*Carex
Carex
Carex
Carex

Longii Mackenz.

lupuliformis Sartwell

lupulina Muhl.

lupulina var. pedunculata Gray
lurida Wahelenb.

Mackenziei Krecz.

mainensis Porter
Merritt-Fernaldii Mackenz.
mesochorea Mackenz.
Michauxiana Boeckl.
Mitchelliana M. A. Curtis
molesta Mackenz.

Muhlenbergii Schkuhr
Muhlenbergii var. enervis Boott
nigra (L.) Reichard

nigra var. strictiformis (Bailey) Fern.

nigromarginata Schwein.
normalis Mackenz.

novae-angliae Schwein.
oligocarpa Schkuhr

oligosperma Michx.

X Olneyi Boott

ormostachya Wieg.

oronensis Fern.

paleacea Wahlenb.

pallescens L. var. neogaea Fern.
panicea L.

pauciflora Lightf,

paupercula Michx,

paupercula var. irrigua (Wahlenb.)

Fern.

+ otttttt ++ HHHH FF+FF+EHH+4EF+44444448

+

+ ++++4+4++ + ++ ++

D

rA

TER

++t+++++t+44+444

+++

+ +

+ +++++ +++ ++ +444

+++ + +++ + +4+444+4+44

I +++ + +

+ ++4++4+

+ ++ +

+++ BEP

Mass,

++

++++

+++ ++ +t+4+4++ 44444444

+ +++ +Ht44+ +4+4+4+4+44/4

171

R. I. Conn,
+ +
— +
+ +
+ +
+ +
au
+ +
+ +
+ +
+ +
+ +
+ +
+ +
ES
JL
+ +
A
+ +
+ +
+ +
+ +
ES
i
+ +
+ +
zs
JL
+ +
E
A
E
+ +
af
+ +
+ +
-4-
-+

172 Rhodora [Vol. 64

N.H. Vt. Mass. R.I. Conn.

+

Carex paupercula var. pallens Fern.

Carex Peckii Howe

Carex pedunculata Muhl.

Carex pensylvanica Lam.

Carex pensylvanica var. distans Peck

Carex plantagianea Lam.

Carex platyphylla Carey

Carex polymorpha Muhl.

Carex prairea Dew.

Carex prasina Wahlenb.

Carex praticola Rydb.

Carex projecta Mackenz.

Carex Pseudo-Cyperus L.

Carex X pseudohelvola Kihlm.

Carex radiata (Wahlenb.) Dew.

Carex rariflora (Wahlenb.) Sm.

Carex retroflexa Muhl.

Carex retrorsa Schwein.

Carex Richardsonii R. Br.

Carex rosea Schkuhr

Carex rostrata Stokes

Carex rostrata var. ambigens Fern.

Carex rostrata var. utriculata (Boott)
Bailey

Carex salina Wahlenb. var. kattegatensis
(Fries) Almq.

Carex salina X stricta

Carex saxatilis L. var. miliaris (Michx.)
Bailey

Carex saxatilis var. rhomalea Fern.

Carex scabrata Schwein.

Carex scirpoidea Michx.

Carex Schweinitzii Dew.

Carex scoparia Schkuhr

Carex scoparia var. tesselata Fern. & Wieg.

Carex seorsa Howe

Carex X setacea Dew. +

Carex silicea Olney

Carex sparganioides Muhl.

*Carex spicata Huds.

Carex Sprengelii Dew.

Carex squarrosa L.

Carex sterilis Willd.

Carex stipata Muhl.

Carex straminea Willd.

Carex striatula Michx.

Carex stricta Lam.

Carex stricta var. strictior (Dew.) Carey

Carex styloflexa Buckl.

Carex Swanii (Fern.) Mackenz.

Carex tenera Dew.

Carex tenera var. echinodes (Fern.) Wieg.

Carex tenuiflora Wahlenb.

Carex tetanica Schkuhr

Carex tincta Fern,

Carex tonsa (Fern.) Bickn.

++

++ + + ++ + ++++++++

+tt+] + ++ ++ +++++
++ 44

++ ++ + ++ +4+4+4+4+44+4+

—
+

+ + ttt + +tt++t+tt+++++++++3
—
-+

++ + +++ ++ + ++ +4+4+4+4+4+4+4++44+

++++
+

++ +++
+ ++

++ +4 |+

+ +444

4
++ +4+4+4+4++4++4+4++4++4+ ++ +
++ ++ 4+ + 4+ + + +

++ ++
ttt Ftt4+4+t+4t+++t+4+4+44+ ++ +

++ +) 4+ ++

++ + ++ ++
+++ |

++

1962] Committee on Plant Distribution 173

=
E

s. R. I. Conn.

Carex torta Boott

Carex tribuloides Wahlenb.

Carex X trichina Fern.

Carex trichocarpa Muhl.

Carex trisperma Dew.

Carex trisperma var. Billingsii Knight

Carex Tuckermani Boott

Carex typhina Michx.

Carex umbellata Schkuhr

Carex vaginata Tausch

Carex vesicaria L.

Carex vesicaria var. distenta Fries

Carex vesicaria var. jejuna Fern.

Carex vesicaria var. monile (Tuckerm.)
Fern.

Carex vesicaria var. Raeana (Boott) Fern.

Carex vestita Willd.

Carex virescens Muhl.

Carex viridula Michx.

Carex vulpinoidea Michx.

Carex Walteriana Bailey var. brevis Bailey

Carex Wiegandii Mackenz.

Carex Willdenowii Schkuhr

Carex Woodii Dew.

z
Fei

++++++ + + + +++
+ ++++++4++++ MR
[T4
TE + BA +4+4++4++ +4

To HEFEI ete toe TER
lL++4++ + + ++

|+t++t++ + + + F4+4+4+4+4+ ++

++ +++

+

Carex was treated by Prof. M. L. Fernald in Rhodora
4:218-230. Since then the number of specimens available
for study has become very much larger and also there has
been a considerable change in nomenclature. It consequently
seemed desirable to reevaluate the genus in the light of the
more abundant material. At the conclusion of his report
Prof. Fernald added “Suggestions for Special Observations”.
It is interesting to note that in at least half the species cited,
the extension of range that he anticipated has been con-
firmed.

In contrast to the previous reports the number of intro-
duced species is insignificant. There are only seven in our
New England area and of these, two were not included in
Prof. Fernald’s list.

The geographical areas are in general the same as'in the
previous reports.

I. GENERALLY DISTRIBUTED. — Carex angustior, C. canescens var. dis-
juncta, and var. subloliacea, C. cephalantha, C. crinita and var. gynan-
dra, C. debilis var. Rudgei, C. intumescens, C. lasiocarpa var.
americana, C. leptalea, C. limosa, C. pallescens var. neogaea, C. rostrata
and var. utriculata, C. stipata, C. striata, C. tenera, C. trisperma and
var. Billingsii, C. vulpinoidea.

174 Rhodora [Vol. 64

Carex limosa apparently is absent from southern New Hampshire,
southern Connecticut and all of Rhode Island. C. tenera is largely
absent from Aroostook County, Maine with only one station at Fort
Fairfield, is represented by only one specimen each in Plymouth County
at Bridgewater and in Barnstable County at Provincetown, and is
absent from Rhode Island.

Ia. GENERAL, EXCEPT CAPE COD. — Carex arctata, C. bromoides, C.
brunnescens var. sphaerostachya, C. communis, C. conoidea, C. Dewey-
ana, C. disperma, C. flava. and var. fertilis, C. Haydenii, C. hystricina,
C. lacustris, C. leptonervia, C. pedunculata, C. projecta, C. scabrata,
C. tribuloides, C. vesicaria and vars. monile and jejuna.

Carex bromoides is infrequent north of 45°. C. brunnescens var.
sphaerostachya is not yet known from Rhode Island or southern Con-
nectieut. C. conoidea has not been found on. Nantucket. C. Deweyana
has not yet been reported from Rhode Island or southern Connecticut
and only two stations are known in eastern Massachusetts. C. flava,
C. hystricina, C. pedunculata and C. projecta are absent from Plymouth
and Bristol counties in Massachusetts and from Rhode Island. C.
scabrata belongs in this category except for a single specimen from
Sandwich, Massaehusetts. C. tribuloides occurs on Cape Cod but only
at Woods Hole and Falmouth, Massachusetts.

Ib. GENERAL, EXCEPT CAPE COD AND THE MAINE COAST EAST OF THE
KENNEBEC RIVER. — Carex torta belongs here but it seems to avoid all
the coast from the Quinnipiack River in Connecticut northeastward.

Ic. GENERAL, EXCEPT CAPE COD AND WASHINGTON COUNTY, MAINE.
Carex abdita, C. convoluta, C. Buxbaumii, C. gracillima, C. interior,
C. stricta var strictior. Although absent from Cape Cod there is a
specimen of C. abdita from Martha's Vineyard. C. convoluta is rare
north of 45°. C. Buxbaumii is a rather uncommon species with only
one station in New Hampshire at Columbia in Coos County, and three
in Vermont. In southern New England it seems to be concentrated
west and north of Boston. There is one specimen of C. interior from
Nantucket. Carex stricta var. strictior, although a rich woods species,
is represented by two specimens from Martha's Vineyard annotated
by K. K. Mackenzie.

IIa. NORTHERN — NOT OR NOT MUCH SOUTH OF 43°. — Carex aenea,
C. arcta, C. atratiformis, C. Backii, C. canescens, C. crinita var. minor
and var. simulans, C. demissa, C. gynocrates, C. Houghtonii, C. lenti-
cularis and var. Blakei, C. Michauxiana, C. paupercula and var.
irrigua, C. Peckii, C. scirpoidea, C. tenuiflora, C. vesicaria var. Raeana,
C. Wiegandii. C. atratiformis is infrequent and chiefly calcicolous.
Although C. Backii is a northern species it is not represented by any
specimen north of 45^. C. demissa is adventive along the coast in
southern Connecticut. C. gynocrates apparently is restricted to the
caleareous areas of northern Maine with no station south of 45°.

1962] Committee on Plant Distribution 175

C. scirpoidea is found chiefly in mountainous areas. C. tenuiflora is
apparently a species of basic soils and is rare in New England.

IIb. NORTHERN — NUMEROUS STATIONS SOUTH OF 43°, — Carex Craw-
fordii, C. deflexa, C. intumescens var. Fernaldii, C. novae-angliae, C.
oligosperma, C. ormostachya, C. pauciflora, C. paupercula var. pallens,
C. Pseudo-Cyperus, C. retrorsa, C. tincta. C. intumescens var. Fernaldii
is absent from Rhode Island and is known from a single station, Cole-
brook, in northwestern Connecticut. C. ormostachya and C. tincta
have their southernmost stations chiefly in western Massachusetts.
C. pauciflora, south of 43°, occurs at several stations in western Massa-
chusetts and at one in Norfolk in northwestern Connecticut. C.
retrorsa, south of 43°, is confined to and is frequent in western New
England.

III. ARCTIC-ALPINE. — Carex Bigelowii, C. brunnescens, C. capil-
laris, C. capitata, C. lenticularis var. albi-montana, C. katahdinensis,
C. mainensis, C. rariflora, C. saxatilis var. miliaris and var.
rhomalea. C. Bigelowii is found on some of the lower mountains in
northern New England and thus is not exclusively arctic-alpine.

IVa. SOUTHERN — GENERAL SOUTH OF 45?.— Carex artitecta, C.
comosa, C. cumulata, C. Emmonsii, C. folliculata, C. lupulina, C. lurida,
C. Merritt-Fernaldii, C. pensylvanica, var. distans, C. scoparia, C.
tonsa. C. artitecta is very sparsely represented in the three northern
states. C. cumulata is absent from Massachusetts west of the Connecti-
cut River except for stations at Sheffield and Mt. Washington. North
of 43° C. Emmonsii is entirely coastal except for one station at Burling-
ton, Vermont. C. tonsa is very sparsely represented in the three north-
ern states.

IVb. SOUTHERN — GENERAL SOUTH OF 45° BUT NOT ON MAINE COAST
EAST OF THE KENNEBEC RIVER. — Carex annectens and var. xanthocarpa,
C. blanda, C. Howei. C. annectens var. xanthocarpa is local with only
seven stations.

IVe. SOUTHERN — GENERAL SOUTH OF 45° BUT NOT IN WASHINGTON
COUNTY. — Carex atlantica, C. digitalis, C. pensylvanica, C. Swanii,
C. umbellata, C. virescens.

IVd. SOUTHERN — SOUTH OF 45° BUT NOT ON CAPE COD. — Carex
argyrantha.

IVe. SOUTHERN — GENERAL SOUTH OF 45° BUT NEITHER ON CAPE COD
NOR IN WASHINGTON COUNTY.— Carex brevior, C. cephalophora, C.
foenea, C. laxiflora, C. lupulina var. pedunculata, C. normalis, C. platy-
phylla, C. prasina, C. radiata. C. cephalophora is local north of 43^. C.
lupulina var. pedunculata occurs on both Martha's Vineyard and
Nantucket but not on Cape Cod.

V. CHIEFLY THE THREE SOUTHERN STATES. — Carex incomperta, C.
laevigata, C. laxiculmis, C. Muhlenbergii and var. enervis, C. vestita.
C. Muhlenbergii has numerous stations in extreme southwestern Maine

176 Rhodora [Vol. 64

and southeastern New Hampshire and two stations in Vermont. C.
Muhlenbergii var. enervis apparently prefers more basic soils.

Va. CHIEFLY THE THREE SOUTHERN STATES BUT NOT IN WESTERN
MASSACHUSETTS. — Carex alata, C. bullata, C. Longit. C. alata is
confined to eastern Massachusetts and Rhode Island.

Vb. CHIEFLY THE THREE SOUTHERN STATES BUT NOT ON CAPE COD. —
Carex aestivalis, C. flaccosperma var. glaucodea, C. polymorpha, C.
retroflexa, C. seorsa, C. aestivalis is absent from Rhode Island and is
local in northeastern Connecticut,

Vc. CHIEFLY THE THREE SOUTHERN STATES BUT NEITHER CAPE COD NOR
WESTERN MASSACHUSETTS. — Carex abscondita, C. Bicknellii, C. fes-
tucacea, C. straminea, C. striatula. C. festucacea is infrequent. C.
striatula is local with only six stations.

VI. SOUTHWESTERN NEW ENGLAND CHIEFLY.— Carex Bushiü, C.
Davisii, C. nigromarginata.

VII. WESTERN NEW ENGLAND, CHIEFLY WEST OF THE CONNECTICUT
VALLEY. — Carex Baileyi, C. cristatella, C. gracilescens, C. hirsutella,
C. squarrosa. C. Baileyiis largely confined to Vermont, save for one
station at Cheshire in western Massachusetts and one at Franconia,
New Hampshire. C. cristatella occurs chiefly in western New England
but with isolated stations in eastern Massachusetts and New Hamp-
shire. C. gracilescens is occasional on basic soils near Boston and
occurs at a single station in Coos County and one in Grafton County,
New Hampshire. The Boston and Providence stations for C. squarrosa
doubtless represent waifs.

VIIIa. COASTAL PLAIN PROPER. — Carex Mitchelliana, C. Walteriana
var. brevis.

VIIIb. COASTAL PLAIN EXTENSIONS. — Carex albolutescens is a
southern coastal plain species that also occurs at Westwood, Massa-
chusetts and at two stations in southeastern New Hampshire.

IXa. CALCICOLOUS — CHIEFLY WEST OF THE CONNECTICUT RIVER IN
THE SOUTH, IF IN THE EAST MOSTLY NORTH OF 45°. — Carex albursina,
C. alopecoidea, C. amphibola var. rigida and var. turgida, C. capillaris
var. major, C. castanea, C. Crawei, C. eburnea, C. elachycarpa, C. flava
var. gaspensis, C. formosa, C. Garberi var. bifaria, C. granularis, C.
Grayii, C. Hitchcockiana, C. Josselynii, C. lupuliformis, C. oligocarpa,
C. Schweinitzii, C. tetanica, C. trichocarpa, C. typhina, C. vaginata.

The majority of these calciphiles occur only in the limestone area of
western New England. C. alopecoidea has one station at Vassalboro,
Kennebee County, Maine. C. amphibola war. turgida is of more
northern. distribution than var. rigida which has not been reported
north of 43? 30". C. capillaris var. major, C. Garberi var. bifaria, C.
elachycarpa, C. flava var. gaspensis and C. Josselynii are restricted
to the calcareous regions of Aroostook County, Maine. C. castanea
occurs in limy areas in both western and northern New England.

1962] Committee on Plant Distribution 177

C. Crawei has been reported from only four stations, two in the
Aroostook Valley and two in the upper Housatonic Valley in Connecti-
cut. C. trichocarpa is absent from most of Vermont. Apparently the
only station east of the Connecticut River for C. typhina is the one
at Leeds, Maine.

IXb. PLANTS OF BASIC SOILS, NOT NECESSARILY LIMESTONES. — Carex
aquatilis var. altior, C. aurea, C. Bebbii, C. cephaloidea, C. diandra,
C. granularis var. Haleana, C. hirtifolia, C. lanuginosa, C. plantaginea,
C. prairea, C. rosea, C. sparganioides, C. Sprengellii, C. Tuckermani.
C. aurea in our area is not necessarily a cacliphile. C. Bebbii is infre-
quent and is not reported from Rhode Island or eastern Connecticut.
C. cephaloidea, C. hirtifolia and C. sparganioides are rich woods plants
not collected north of 45?. C. plantaginea is an inland species with no
stations on the coast. It is rare north of 45°.

Xa. MARITIME IN VICINITY OF COAST.— Carex hormathodes, C.
Mackenziei, C. paleacea, C. salina var. kattegatensis, C. silicea. A
hybrid of C. hormathodes and C. straminea has been found on Nan-
tucket. C. Mackenziei is northern and has not been reported south of
York County, Maine. C. paleacea is northern extending southward to
Plymouth, Massachusetts. C. salina var. kattegatensis is also northern
and is absent scuth of Boston. A hybrid of C. salina var. kattegatensis
and C. stricta is reported from Arlington and Medford, Massachusetts.

Xb. MARITIME WITH INLAND STATIONS. — Carex nigra and var.
strictiformis. The latter has a similar distribution, but is much less
common.

XI. MISCELLANEOUS. — Carex adusta, C. artitecta var subtilirostris,
C. atheroides, C. Barrattii, C. chordorhiza, C. Collinsii, C.x copulata,
C. crinita var. brevirostris and var. Porteri, C. debilis and var. inter-
jecta and var. strictior, C. exilis, C. Grayii var. hispidula, C.x Harti,
C.x Knieskernii, C. livida var. Grayana, C. mesochorea, C. molesta,
C.x Olneyi, C. oronensis, C. praticola, C.x pseudohelvola, C. Richard-
sonii, C. rostrata var. ambigens, C. salina x stricta, C. scoparia var.
tesselata, C.x setacea, C. sterilis, C. tenera var. echinoides, C.x trichina,
C. vesicaria var. distenta, C. viridula, C. Willdenowii, C. Woodit.

C. adusta occurs only east of the Penobscot River near the ccast.
C. artitecta var. subtilirostris has only one station at Malden in east-
ern Massachusetts. C. atheroides has only one station at Rockland,
Maine and is possibly a waif. C. Barrattii and C. Collinsii are southern
in distribution reaching their northeastern limit in Connecticut and
Rhode Island. C. chordorrhiza is a northern species and very local,
occurring only in eastern Maine and western Vermont. C.x copulata
is reported from four stations, Grafton, New Hampshire; Newfane,
Vermont; Sandisfield, Massachusetts, and Waterford, Connecticut.
C. crinita var. brevirostris occurs only at Cohasset, Massachusetts;
Providence, Rhode Island, and Bridgeport and Danbury, Connecticut.

178 Rhodora [Vol. 64

C. erinita var. Porteri has two stations, Monhegan, Maine, and Han-
cock, Vermont. C. debilis has been found on Martha's Vineyard and
Nantucket only. C. debilis var. interjecta has three stations in New
Hampshire, one at South Burlington, Vermont, seven in Massachusetts
and one in Connecticut. C. debilis var. strictior has one station in
Maine at Brownville, eleven in New Hampshire, one at Willoughby,
Vermont and one at Bellingham, Massachusetts. C. exilis is con-
centrated in northeastern Massachusetts but otherwise has only widely
scattered stations. C. Grayii var. hispidula has two stations, Hartford
and Middletown, Connecticut. C.x Hartii has seven scattered stations
in New England. C.x Knieskernii is represented by only five stations
in our whole area. C. livida var. Grayana, a calcicolous species, is
placed here as in all New England there are only six stations, two in
Aroostook County, Maine, one in western Vermont and three in
northeastern Massachusetts. C. mesochorea is cited in Gray's Manual
from Massachusetts but no specimen has been seen. C. molesta has only
four stations in New England, Concord, Massachusetts; Ferrisburg,
Vermont, and Bethlehem and Southington, Connecticut, C.x Olneyi
has twelve stations in southeastern New England. C. oronensis seems
to be endemic along the lower Penobscot River in Maine. C. praticola
is a northern species at its southern limit with two stations only in
northern Maine, at Island Falls and Masardis, both in Aroostook
County. C.x pseudohelvola has two stations in extreme southeastern
Maine at Moose Island and Pembroke. C. Richardsonii is represented
by a single station at Mt. Equinox in Manchester, Vermont where it
is abundant. C. rostrata var. ambigens has three stations, at St.
Francis, Maine and Monterey and Springfield, Massachusetts. C.
salina x stricta was collected in 1858 at Medford and in 1887 at Arling-
ton in eastern Massachusetts. C. scoparia var. tesselata has three
stations on the Maine coast east of the Penobscot River. C.x setacea
is apparently a hybrid with only four stations in our area. C. sterilis
has one station at Presque Isle in northeastern Maine and three in
Connectieut. C. tenera, var. echinoides has three stations only, one at
South Hero, Vermont and two in Connecticut at East Hampton and
North Canaan. C.x trichina has been collected at two stations in the
Aroostook Valley, Maine at Chapman and Fort Fairfield. C. vesicaria
var. distenta has only ten widely scattered stations. Perhaps this
should be recognized only as a form and not as a good variety. C.
viridula is placed in this category as it is in part coastal and in part
inland chiefly calciphile. Possibly this species requires further study.
C. Willdenowii has four stations only in New England, one at Sudbury
in western Vermont, two at Malden and Randolph in eastern Massachu-
setts and one at East Haven in southern Connecticut. C. Woodii has
two stations only at Salisbury and New Milford in extreme western
Connecticut,

1962] Lakela — Polycarpaea 179

XII. INTRODUCED SPECIES. — Carex acutiformis, C. brunnea, C.
caryophyllea, C. hirta, C. leporina, C. panicea, C. spicata. C. acuti-
formis is reported only from the vicinity of Boston. C. brunnea has
been collected but once — at Salem, Massachusetts. —

R. C. BEAN, A. F. HILL, R. J. EATON

OCCURRENCE OF SPECIES OF POLYCARPAEA LAM.
(CARYOPHYLLACEAE) IN NORTH AMERICA

OLGA LAKELA

Plants unknown to the author were discovered in a margi-
nal area of sandhill off Overlook Drive, adjacent to the
business district of Temple Terrace, Tampa, Florida. The
plants, reminiscent of Paronychieae, were localized in open-
ings of the vegetation, prevailingly of grasses and compo-
sites, and a few dwarf live oak. Their compact clusters
of silver white cymes in late anthesis and in fruit, gave a
touch of luster to the nondescript background. Collection no.
23410 was made October 15, 1960. Subsequently, nos. 24530,
August 7, and 24779, October 18, 1961, were collected from
the same site.

In the meantime, a search of the area revealed a wider
distribution, east of Overlook Drive and 56th Street. The
plant is well established on a grassy terrace adjoining the
grounds of Temple Terrace School southside, and in the
margin of a field about 1 block north of the school. Its
sporadic occurrence was discovered in the area embracing
largely the abandoned Henderson Hillsborough Internation-
al Airport, extending over two miles northwest from Temple
Terrace. There it is surviving in grassy and weedy vege-
tation along the old runways and road boulevards. Collec-
tion no. 24916, March 18, 1962 consists of post-mature
plants with clusters of cymes, faded in everlasting fashion.

Discovery of the novelty raised the question of its specific
identity. Dr. Carroll E. Wood, Jr., referred it tentatively to
Polycarpaea corymbosa (L.) Lam. (Achyranthes corym-

180 Rhodora [Vol. 64

bosa L. Sp. Pl. 205. 1753). The species with pantropic dis-
tribution was based on a Ceylon plant. An intensive study
and investigation ensued. Specimens were sent to the Royal
Botanie Garden, Kew, where they were critically examined
in comparison with Kew materials from the various conti-
nents by Messrs J. P. M. Brennan and Peter Taylor. Accord-
ing to their esteemed report the Florida plant is different
from P. corymbosa, the Asian-African taxon, but it differs
less from P. brasiliensis Camb., the South American taxon.
Since the identities of the above named taxa seem to be in
need of an overall survey, the status of the Florida plant
remains in question. However, it seems appropriate at this
time to report the occurrence of Polycarpaea in North Amer-
ica. Meanwhile a study of the Florida plant is underway
with hopes of gaining a better understanding of characteris-
tics and affinities in relation to these known taxa.

It may suffice to point out the habit and the more promi-
nent characteristics of the South American and the Old
World taxa studied in collections. P. brasiliensis Camb., in
St. Hilaire Fl. Brasil. merid. II. 132. (1829), a plant of
Brazil, described without citations of particular specimens,
is distinguished by thick, fusiform radix giving rise to sev-
eral stems. Chodat et Hassler, in Bulletin de L'Herbier
Boissier (2weSer.) 3:190 (1903) treated Paraguayan plants
as P. corymbosa var. brasiliensis, with a diagnostic state-
ment, “Herba perennis, radix lignosa napiformis —” citing,
Hassler Y-aca, Dec., n. 6691 & Apa. Dec., n. 8123, (F, GH,
US). The leaves in South American specimens are strongly
revolute, appearing filiform, with deeply cleft stipules pro-
longed to long filaments, and a terminal bristle to 3 mm.
long. The radix is often lacking in herbarium specimens
and the plant may be too young for a thickened crown;
however, the leaf character may be an aid in identification.

P. corymbosa (L.) Lam., Rohrbach in Mart. Fl. Bras.
XIV. 2 (1872) p. 254, t. 58, fig. 1, is representative of the
Old World taxon, an annual plant with slender tap root,
stems to 30 cm. tall, with internodes usually longer than the
leaves, strict or more or less diffusely branched. The leaves,

1962] Lakela — Polycarpaea 181

despite the pubescence, with short terminal bristles and
slightly revolute margins, are similar to those of the Florida
plant. The collection of Cooley and Siyambalagstenne from
Dambulla, Ceylon, 14-1-57, (USF), the type region, well
conforms to the description of the species. The ten mounted
plants bear a strong resemblance to those of China, India
and Africa. These, like the South American plants, retain
the cotyledonary fascicles and the basal leaves of the lower
nodes almost through the flowering period. In the Florida
plant they are lost in preanthesis. Therefore a conclusion
of its identity cannot be made before the plant as a whole
is understood. To verify the occurrence of Polycarpaea in
North America the plant is herewith described.

DESCRIPTION OF THE FLORIDA PLANT

Upright annual herb 6-10 (14) cm. tall; taproot to 25 cm. long, with
relatively few secondary roots; stem profusely branched through the
upper half to two-thirds of its length; internodes 5-15 mm. long,
scintillescent and reddish under the loose white pilosity, or glabrate;
nodes with singular swellings of yellowish fatty parenchyma, em-
bracing the axils of leaves with stipules, floral bracts and to a lesser
extent those of the sepals; leaves linear, with revolute margins, 1-2.5
em. long, 1-1.5 mm. wide, opposite, appearing verticillate with axillary
fascicles, glabrous, fleshy, essentially sessile, excepting a few, in the
cotyledonary node, 1-nerved, the nerve prominent below and termi-
nating in a bristle about 1 mm. long, and, like the internodes, scintil-
lescent and becoming reddish; stipules scarious, cleft, the points
prolonged to filaments, 8-5 mm. long; inflorescence leafy, cymes
dichasial, compound, terminating the numerous branches; floral bracts
scarious, entire or cleft, overlapping the pedicels; flowers lustrous-
white, 2.8-3.1 mm. long, perigynous; hypanthium short, crater-like,
reddish within in anthesis; sepals scarious glabrous, lanceolate-acute,
dorsally rounded, margins convex in outline, with a reddish-orange,
deltoid, median basal spot; petals ovate, to 1 mm. long, entire or erose
at tips, rose pink, fading to orange, connate with the stamen bases
around the rim of the hypanthium; filaments subulate, curved inward,
.4 mm. long, anthers white, oblong closely surrounding the stigma in
bud; pistil in anthesis 1 mm. long, the short stipe centered within
the hypanthium base; stigma consistently nearly sessile, or with style
.13 mm. long, minutely papillose, 3-fid; capsule 1.5 mm. long, usually
shed without the stipe, 3-valved, bronzed, the valve margins yellow,
the stigma with style often short-persistent to a strand of inner tissue
of one of the valves; ovules 7, amphitropous, seeds subreniform, com-

182 Rhodora [Vol. 64

pressed, brownish, .3 mm. long when mature; before fully mature, the
thin integuments clearly reveal the transparent endosperm with the
curved embryo around the convex margin; cotyledons accumbent.
Flowers, late August to November; ripe seeds to Jan. Known only

from Tampa.

For citations and loans of herbarium materials, the author
is indebted to Dr. C. Earle Smith, Jr., Chicago Museum of
Natural History (F); Dr. C. E. Kobuski, Gray Herbarium
(GH) ; Dr. J. R. Swallen, United States National Herbarium
(US) ; Dr. James D. Ray, Jr., University of South Florida
(USF); to Drs. G. Taylor and C. L. Hubbard of Roya!
Botanic Garden, Kew, for report of study. — UNIVERSITY
OF SOUTH FLORIDA, TAMPA.

A NEW STATION FOR PINGUICULA VULGARIS IN ONTARIO.
— Pinguicula vulgaris L. is an amphi-atlantic species which
has an uneven distribution both in Europe and North Amer-
ica. In the New World it is a boreal species which extends
across Canada from Newfoundland to British Columbia and
Alaska and south into northern New York, New Hamp-
shire, Vermont, Minnesota, Michigan, Montana, Washing-
ton and Oregon. In Ontario P. vulgaris has been known for
many years from the James Bay lowlands, Lake Nipigon
and the north shore of Lake Superior, and the Bruce Penin-
sula which separates Georgian Bay from Lake Huron. A
map of its distribution as known to Hultén is given in
Kungl. Svenska Vetenskapsakademiens Handlingar, Fjarde
Serien Band 7. Nr.1:231. 1958.

In the southern part of its range in northern New York
State, this species is found only on wet limestone cliffs of
deep cool ravines such as occur at Ithaca and the Finger
Lakes. A similar situation was found in Wellington County,
Ontario, near Elora, where the Grand River cuts through
the Guelph Formation of Silurian limestone (dolomite).

1Contribution No. 181 from the Plant Research Institute, Research Branch, Canada
Department of Agriculture, Ottawa.

1962] Cody — Pinguicula 183

While botanizing on August 16, 1958, with Dr. W. G.
Dore along the floor of the ravine cut out by the river, a
a peculiar green was noted on the cliff on the opposite side
of the river. Fast water along most of the base of the cliff
made it difficult to get near, but it was possible to reach the
water’s edge on the north side of the river by climbing
down a steep well-wooded slope nearby. We were then able
to work along the foot of the slope to the cliff face. From
there we could see the peculiar light yellowish-green leaves
of Pinguicula vulgaris. The plants were very numerous,
rooted in wet moss on what appeared to be a permanently
wet and dripping rock face. By dint of much stretching we
were able to secure eight plants, enough to make one her-
barium sheet. All but one of the plants collected were
sterile. The fertile plant bore an almost mature capsule.
The data are as follows: ONTARIO: Wellington Co., Elora
Glen, Grand River, in wet moss in bright sun, numerous
but inaccessible on southfacing cut of gorge, Cody & Dore,
11046 (DAO).

An examination of specimens in the University of Toron-
to Herbarium revealed another apparently unreported col-
lection from along the Grand River some 30 miles to the
south: Brant Co., Glen Morris, R. F. Cain s.n., 8/6/29
(TRT).

Elora Glen and Glen Morris are approximately half way
between the Bruce Peninsula, where Pinguicula vulgaris
has been collected frequently, and the sites in northern New
York. It is not far from the fault of the Niagara Escarp-
ment which runs from the Bruce Peninsula into New York
at Niagara Falls. This then might be the route by which
this species reached northern New York and it is possible
that other sites for Pinguicula vulgaris might be found in
similar situations along the escarpment or where streams
have cut through the sedimentary rock nearby. Such situa-
tions might possibly be found in the Niagara Gorge, along
the Jordan River, at Credit Forks and along the Rocky
Saugeen River. — W. J. Copy.

184 Rhodora [Vol. 64

HYPTIS RADIATA (LABIATAE) AN ILLEGITIMATE NAME. —
Hyptis radiata Willd., Sp. PI. (ed. 4) 3 pt. 1: 84, 1800, was
published as a new name for Clinopodium rugosum L., Sp.
Pl. (ed. 1) 2: 588, 1753 (Willdenow of course cited ed. 3,
with page no. 822, but the name originates in the first edi-
tion). Because the epithet rugosum was available for use
under Hyptis but was not retained, the binomial Hyptis
radiata is automatically illegitimate by present rules. Be-
cause of the validly published Hyptis rugosa Benth., Lab.
Gen. et Sp. p. 86, 1832, the Linnaean epithet cannot now be
transferred to Hyptis. In Carl Epling’s “Synopsis of the
genus Hyptis in North America” (Fedde Repert. Sp. Nov.
34: 73-130, 1933), the only available name seemingly is H.
latidens Urban, 1926, three other binomials published by
Gandoger in 1918 being listed as “nomina” only (1. c. p.
118). This is not correct, for Gandoger supplied Latin
diagnoses in the form of keys, and cited specimens, so that
all three of his names were validly published (Bull. Soc. Bot.
France 65: 66, 1918). The identity of these will be dis-
cussed later.

The hitherto unidentified Pyenanthemum ? alatum Rafin-
esque, Fl. Ludov. p. 40, 1817, has never been listed in any
synonymy under Hyptis, although Rafinesque himself
thought it might really belong to that genus. Here is his
account:

116. Pyenanthemum ? alatum Raf. — Foliis oblongis,
acutis, dentatis, subtus tomentosis, longe petio-
latis, petiolis alatis, capitulis longe peduneulatis,
involueris 13 phyllis, alternis longioribus. Raf.
— Monarde bouton Rob. p. 388. Stems four feet
high, four angular, grooved and branched, leaves
green above, flowers white, unfolding gradually.
Faintly aromatic and bitter: vulgar name herbe à
bouton; Robin owns it has four stamina: but
says it has all the other characters of Monarda.
It is perhaps a Hyptis.

Anyone who has seen the Gulf Coast Hyptis radiata will
recognize at once that it is exactly what Rafinesque de-

1962] Shinners — Hyptis radiata 185

scribed. Since his specific epithet has not been used under
Hyptis, and is much older than those of Gandoger and
Urban, the correct name for H. radiata becomes H. alata
(Raf.) Shinners, comb. nov., based on Pyenanthemwum ?
alatum Raf., quoted above.

Of the three species published by Gandoger (all on the
same page), the first, H. floridana, was described as ‘‘pubes-
cens, caulis copiose setoso-muricatus, folia basi rotundata
deltoidea, bracteae florales calyce breviores, sepala vix 14
mm. longa. — Florida, ad Jacksonville ( Williamson!)." The
pubescent and muricate stem, broad leaf blades, small floral
bracts, and very short calyx teeth are all perfectly charac-
teristic of H. mutabilis (A. Richard) Briquet, and H. flori-
dana Gandoger is to be listed as a synonym of that species
rather than of H. alata, (H. radiata). Gandoger's other two
species are discussed together separately from H. floridana,
and are plainly forms of H. alata. The third is the common
and widespread type, identical with H. radiata Willd. and
Pycnanthemum ? alatum Raf.: “Involucri phylla capitulum
villosum superantia, caules ad angulos hispiduli, folia late
oblonga. — America sept., Mississippi ad Biloxi (Tracy n.
61!) et ad Mendenhall (ejusdem n. 8751!)." The first num-
ber cited is adopted as lectotype of H. Tracyi. I have seen
only topotype material (Demaree 28392, SMU). :

Gandoger’s second species, H. leiocephala, was a more
glabrous and narrow-leaved form: “Involucri phylla capi-
tulo glabro breviora, caules ad angulos glabri, folia anguste
oblonga. — Florida, ad Jacksonville (Keeler! Leggett!)."
This sounds very similar to H. latidens Urban, Fedde
Repert. Sp. Nov. 22: 368, 1926, from Cuba, considered by
Epling only a form of H. radiata not worthy of recognition.
Because the extremes are so different in appearance, be-
cause the narrow-leaved one is geographically segregated,
and because it is a significant piece of evidence about the
history of the Southeastern flora, I think that the formal
designation of a variety is justified. Not having seen either
Gandoger’s or Urban’s specimens, I prefer to supply an
entirely new name based on material I have examined:

186 Rhodora [Vol. 64

Hyptis alata var. stenophylla Shinners, var. nov. Caulis (nodis excep-
tis) glaber vel subglaber. Folia caulina angusta plerumque laminis
lanceolatis vel lineari-lanceolatis paucidenticulatis vel subintegris.
TYPE: northwest of Loxahatchee in the Corbitt Wildlife Management
area, Palm Beach Co., Florida, R. Kral 5657, 3 Sept. 1957 (holotype
SMU, isotype FSU). “Frequent on moist to wet sandy peat of broad
marsh and grass-sedge meadows.” The leaf blades of these specimens
are linear-lanceolate and subentire. In the following they are lanceo-
late and mostly few-toothed. All are from lower peninsular Florida.
CHARLOTTE CO.: 15 miles north-northwest of Fort Myers, Kral 7502
(FSU). Caloose Experimental Range, southeastern Charlotte County,
William P. Adams 218 (FSU). COLLIER CO. vicinity of Lake Trafford,
about 3 miles west of Immokalee, Adams 208 (FSU). One mile south of
Ochopee, Richard S. Mitchell 630 (FSU). DADE CO.: 5 miles west of
Naranja, F. H. Sargent 6451 (SMU).

As indicated above, H. leiocephala Gandoger and H. lati-
dens Urban are very probably synonyms of var. stenophylla
rather than of var. alata. Unless it is desired to recognize
two species, the precise disposition of these names is of no
consequence.

I am greatly indebted to Dr. R. K. Godfrey for the loan of
herbarium material from Florida State University. —
LLoyp H. SHINNERS, SOUTHERN METHODIST UNIVERSITY,
DALLAS 22, TEXAS.

MAKERS OF NORTH AMERICAN BOTANY’

Who are the Makers of North American Botany ? — When
Dumas Malone and his associates organized the Dictionary
of American Biography they posed the simple criterion when
selecting the persons to be included: Did the botanist infiu-
ence the American scene? Certainly not only the innovators
but those who developed the science should be included.
The DAB includes forty botanists? that are not mentioned
by Humphrey. They are:

1Makers of North American Botany. By Harry Baker Humphrey. Edited by Robert
R. Humphrey. Ronald Press, New Pork. $6.00 xi + 265 pp. 1961.
?See DAB Index volume, 373. 1943 and Supplements I and II.

1962] Ewan — Book Review 187

W. P. C. Barton,’ W. D. Brackenridge, Robert Bridges, S. B. Buckley,
Henry James Clark, John Clayton, Manasseh Cutler, Stephen Elliott,
B. T. Galloway, F. H. Knowlton, Arthur Hollick, Henry Kraemer,
Adam Kuhn, J. G. Lemmon, F. J. Lindheimer, Thomas Meehan, Ezra
Michener, Elisha Mitchell, John Mitchell (d. 1768), Charles T. Mohr,
J. S. Newberry, John Alsop Paine, Andrew Parmentier, F. P. Porcher,
T. C. Porter, John L. Riddell, J. N. Rose, J. T. Rothrock, H. H. Rusby,
P. A. Rydberg, H. P. Sartwell, J. L. E. W. Shecut, C. W. Short,
George Thurber, Edward Tuckerman, L. M. Underwood, Thomas
Walter, David White, E. H. Wilson, Charles Wright.

Certainly no two authors would select the same “makers
of North American botany" but most, if not all, of the fore-
going DAB biographees might be expected in a book
described as “invaluable for biographical reference." Some
other ‘makers’ that come to mind are David Hosack, who
patronized botany and founded the Elgin Botanic Garden ;
Bernard M'Mahon, author of the first gardeners' guide to
be printed in this country ; Eugene Waldemar Hilgard, soil
scientist and pioneer agronomist; John Belling, cytologist
and stain technologist; Edward Palmer, ethnobotanist and
zealous collector in remote regions of Mexico and the West;
Almira Lincoln Phelps, student of Amos Eaton, author of
an influential textbook used in women’s colleges ; David Fair-
child, pioneer in the field of plant introduction, author of
The World was my Garden, one of the best botanical story
books of the century ; Wilhelm Seifriz, geobotanist, physiolo-
gist, pioneer in protoplasmic studies; Edward Wilbur Berry,
productive paleobotanist, whose knowledge of fossil plants
surpassed many botanists' knowledge of living floras! and
Alice Eastwood, whose seeds of enthusiasm for botany were
sown in thousands of minds for over seventy years!

Dr. Humphrey did not live to see his book published and
it was brought to print by his son who added a few sections
to the account of 122 botanists. Regrettably the reference
value is diminished by at least 94 typographical errors in
the citation of names, dates, and bibliographical references

*Absence of Benjamin Smith Barton from the DAB is notable in view of his impor-
tant role in fostering colonial botany. See Jeannette E. Graustein, Penna. Mag. Hist.
and Biog. 85:423-438. Oct. 1961, for the latest "informed judgment" on Professor
Barton,

188 Rhodora [Vol. 64

to render the book unreliable. This is partly failure to read
proofs carefully, but there are many inconsistencies in the
citation of references which were surely in the original
manuscript.

Humphrey was evidently unaware of a number of impor-
tant studies of some of the botanists he considered. How
can he write of Rafinesque and not make use of Merrill’s
essays? Of Bartram, and not use Francis Harper’s? Of
Benjamin Smith Barton and of Muhlenberg and many other
Philadelphia figures, and not use Pennell’s contributions”
He paid too little attention to the formative influences in
the lives of the ‘makers!’ An index would have served as a
guide to the influence of mentor on student, to contempor-
aries, such as John Belling, now lost within accounts. The
outstanding contribution of the botanist is too often
smothered in the recitation of medals and memberships.
Humphrey’s garland of botanical worthies is not a book of
recollections written for the uninitiated reader browsing
for pleasure. He wrote impersonally as if he feared intimacy
even for those colleagues in his old United States Depart-
ment of Agriculture whom he must have met almost daily.
His prose is without relief in the recital of facts.

Only a few factual errors have been noticed. “Rev. Dr.
[Zaccheus] Collins" mentioned as a Philadelphian who
befriended F. A. Michaux was neither a minister nor a
physician but a merchant. The oft-repeated misalliance of
Audubon with Albert Kellogg has been corrected by Profes-
sor Geiser(1).

More numerous are the fallacious interpretations that
appear all through the book. For example, to say E. L.
Greene *had more than a casual interest" in Pittonia and
Erythea, — journals which he had founded — is a singular
understatement. Lewis David von Schweinitz's herbarium
*became the property of the Academy of Natural Sciences
of Philadelphia," it is true, but the lamentable scheme of
von Schweinitz in discarding the collectors' original labels
in favor of his own copied tickets is not mentioned. Hum-
phrey's statement conveys the impression that this collection

1962] Ewan — Book Review 189

of “approximately twenty-three thousand species” is valu-
able when in fact its value was seriously impaired (2). Hum-
phrey infers that Sargent illustrated his Silva with 740
plates by his own hand when they were the work of Charles
E. Faxon, a high point in American botanical illustration.
Again and again the works of botanists are cited as 'solo
flights’ when rather they were co-authored by others who
go unnoticed. Thus some notable events in American botany
are missed altogether such as when Wilfred Robbins and
Richard Holman created a winning text which was to outsell
all competitors for nearly thirty years. It was recently
revised by professors Weier and Stocking. Pursh is said to
have worked on a flora of Canada making “several important
botanical excursions,” and assembling “important collec-
tions." Both are misleading statements, certainly not sup-
ported by the evidence which I presented in the paper that
Humphrey cited. Pursh worked desultorily, frustrated part-
ly by ill health brought on by discouraging lack of support
for western exploration and by alcoholism. The few surviv-
ing Canadian collections (which evidently were thought
unworthy of mounting by Lambert, Pursh’s patron) give
no impression of his former sharp eye for novelties and
overlooked range extensions seen in his Virginia and New
York collections. Nuttall’s trip up the Missouri River with
John Bradbury is misleadingly reported by Humphrey.
Graustein’s papers, overlooked by Humphrey, should be
consulted regarding this relationship (3). William T. Horna-
day, not D. T. MacDougal, wrote Campfires in Desert and
Lava. Humphrey confused facts about Dr. Engelmann. The
collected Botanical Works of Engelmann were a reissue of
his many scattered papers, the original “notes, drawings,
and data” of which are to be found in the sixty volumes
preserved at the Missouri Botanical Garden. Amos Eaton
never held a faculty appointment at Yale; in fact, he himself
wrote that he had been ‘‘a mere learner at Yale.” McAllis-
ter’s Amos Eaton would have set Humphrey right on this
point. Elias Durand’s herbarium presented to the Jardin
des Plantes represented a selection of one or a few sheets

190 Rhodora [Vol. 64

of each species described from North America, — often
removed from the Philadelphia Academy’s collections! — to
make up what Durand called the Herbarium Boreali Ameri-
canum. Agnes Chase and F. W. Pennell have recorded these
facts(4). Jane Colden prepared notable descriptions of New
York state plants, though her drawings were indeed indiffer-
ent. Together they fortify Humphrey's decision to include
this remarkable young woman in the garland of North
American botanists(5). To call T. S. Brandegee “‘one of the
first pioneer plant collectors of the West" is to overlook the
roll of honor drawn up by Alice Eastwood (6) which includes
Collignon, Mocino, Haenke, Menzies, Chamisso, Vosnesen-
sky, Douglas, Hartweg, Jeffrey, etc., — all prior to Brande-
gee in the West! There is no doubt of the importance of
Brandegee in the extent and excellence of his collecting but
Humphrey does not make this clear.

Notable is the considerable number of men who rose from
amateurship to make a permanent mark in American botany.
If an early motivation in the lives of these ‘makers of North
American botany’ were sought in these sketches, it would
be the appeal of the American outdoors. Sense of explora-
tion played an important part in the formative years of
botanists who ultimately were to enter the experimental
laboratory as well as in those who were to leave their
largest contribution in field botany.— JOSEPH EWAN, TULANE
UNIVERSITY.

1. Field and Lab. 27:94. 1959.

2. F. W. Pennell in Bartonia 16:1-8. 1935.

3. Chronica Botanica 14:1-88. pls. 68-79, 1951; Mo. Hist. Soc. Bull.
12: 249-252. 1956.

4. Bartonia 17: 33-39, and 40-45. 1936.

5. Anna Murray Vail in Torreya 7: 21-34. 1907.

6. Calif. Hist. Soc. Quart. 18: 335-346. 1939.

'ARLOW REFERENCE LIBRARY

OCT 4 1962

Hovova

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by

ALBION REED HODGDON, Editor-in-Chief

ALBERT FREDERICK HILL
STUART KIMBALL HARRIS
RALPH CARLETON BEAN
ROBERT CRICHTON FOSTER
ROLLA MILTON TRYON
RADCLIFFE BARNES PIKE

Associate Editors

Vol. 64 July-September, 1962 No. 759
CONTENTS:
Ernest Jesse Palmer. 1875-1962.
Clarence E. Kobusli coo. cccccccscssescovcacsscscocins suanomscsielttsec 0 (eRmeeninas 195
Some Observations on Flowering in Ruellia (Acanthaceae).
Robert W. Long and Leonard J. Uttal sen 200
Dryopteris Dilatata (Hoffm.) A. Gray in North America.
Donald M. Britton ..saaeeusieeceeereesecescoten DERI NER. 207
On Pinguicula Macroceras Link in North America.
ccc scurcoseeveceesncccesssesesecccasacersenpecoiiesessig imma 212
Holosteum Umbellatum L., An Angiosperm New to Michigan.
'sscque CLIQ. VOpUlAe ooo sé sind sacvevsscessoseeceescudecestelsitcensster tonne S Do
Two New Species of Polygala From Northern Mexico.
BED UII ui eceiteseteieesecocetictceecioes ertt HD e A 225

(Continued on Inside Cover)

The Nem England Botanical Club, Ine.

Botanical Museum, Oxford St., Cambridge 38, Mass.

y Se

CONTENTS: — continued

The Aquatic Form of Alisma Subcordatum Raf.
Richard W. Rhoades „csser 227

Mazus Reptans (Scrophulariaceae) in Maryland.
Lloyd H. Shinners cocccccccccccccccccccccccccssccccccceseccceesacccccecescecceeeseceeees 229

Studies in the Guttiferae, II. Taxonomic and Distributional
Observations on North American Taxa.
Preston Adams -oecccccccccccccccsccecssecececcecessessesssecessceeesecsessusecesececeees 231

The Use of Vapor Fractometry in the Analysis of Some New
England Hops. Edward L. Davis and Richard L. Burns ... 243

Chromosome Numbers in the Compositae. VI. Additional
Mexican and Guatemalan Species.

B. L. Turner, M. Powell and R. M. King ....... e 251
Chromosome Numbers in Mexican and Guatemalan Com-

positae.

J. H. Beaman and B. L. Twrner eee ees 271

Chromosome Counts of Two Thymelaeaceae.
Lorin I. Nevling, Jv. |t eene terere etre 277

Zizaniopsis Miliacea from Illinois.
Glen S. Winter ringer eee eee 283

A Note on the Ascription of Elymus Arenarius to Illinois.
G. Neville Jones voicccccecsscccccccceeeeccsseessstscccccsseceessitsesesessertuceceececees 284

wW
sd

ERNEST JESSE PALMER PLATE 1270

Rhodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 64 July-September, 1962 No. 759

ERNEST JESSE PALMER. 1875-1962*

Ernest Jesse Palmer died at Webb City, Missouri Febru-
ary 25, 1962 in his 87th year. This marked the end of a long
and in many ways remarkable career in the fields of botany
and allied sciences.

He was born in Leicester, England April 8, 1875 and in
1878, when he was three years old he and his sister, Louise,
were brought to this country by their parents, Amos and
Annie Palmer, who settled temporarily near Warrensburg,
Missouri in Johnson County.

At the age of 14, his father, responding to the stories of
quick wealth offered by the mining opportunities in the
southwestern part of Missouri, moved his family to Webb
City and built a home for them at 321 South Main Street.
The wealth never materalized! Throughout the rest of his
life, although he spent many of his years away from Webb
City, the Palmer home was maintained and after retirement
Palmer moved with his family and lived out the rest of his
life there.

In this mining area, young Palmer did find a wealth of his
own liking. It was here that he began his studies of natural
phenomena. In the piles of waste material from the mines,
fossils of plants and animals caught his fancy and as in all
boys of his age the collecting instinct came to the fore. He
developed a very keen instinct for unearthing the most inter-
esting artifacts. In later years at the Arnold Arboretum

*This article on the life of Mr. Palmer will be published also in the current volume
of the Journal of the Arnold Arboretum. Included in the Journal will be a bibliogra-
phy of Mr. Palmer’s published papers compiled by Mrs. Lazella Schwarten and Mrs.
Ernest Palmer.

195

196 Rhodora [Vol. 64

he was always “picking up” Indian arrowheads and other
Indian deposits. This was easy for him — but not neces-
sarily for others. I can recall his telling me of a spot in the
Arboretum where arrowheads were in abundance. I hurried
to the exact location but found nothing. The next day we
went out together — and sure enough, he picked up several
more artifacts. Eventually two attractive trays of Palmer’s
finds in the Arboretum were mounted and have been on dis-
play there ever since.

His formal education was broken off suddenly while he
was attending high school. A physical collapse of his father
made it necessary for him to leave school and seek employ-
ment. This was undoubtedly a severe blow to his pride
because he wanted an education above all things. He loved
study and research — and he intended to continue it at all
costs. Just recently I learned from an article by Marcia W.
Kershaw, a Special Correspondent, of the St. Louis Post-
Dispatch that Palmer managed to study Latin and Greek at
old Webb City Baptist College. I had often wondered where
he obtained his profound knowledge of these languages,
especially Latin which he used so ably in his original techni-
cal descriptions of new species of plants. Mrs. Kershaw's
article brought to light many other facets of Mr. Palmer's
career which were previously unknown to me and I am very
thankful to her for her assistance.

All his life he had managed to study not only the Natural
Sciences but also English literature, mathematics, economies
and poetry until he was considered very proficient in each of
them.

The portrait which accompanies this article is very recent
and shows Mr. Palmer at his desk with a copy of his pub-
lished book of poems entitled, “Gathered Leaves, Green,
Gold and Sere." These poems, I understand are truly
gathered leaves, "leaves" which had been lying around in
various places during the years and were gathered together
by his wife and published through her efforts when Palmer
was eighty-three years old.

His scientific papers were noted not only for his Latin des-
criptions but also for the fine phrasing which he always

1962] Kobuski — Ernest Jesse Palmer 197

employed. He wrote so well and so easily that often he
found it necessary to delete whole paragraphs from his
papers because he considered the contents “unscientific.” It
was a pity — because everyone would have enjoyed reading
them.

Although he was dedicated to the life of a naturalist it
was not until his meeting with Benjamin Franklin Bush,
also from Missouri, that he tried his wings and ventured
away from his home setting. When Bush died in 1937, Pal-
mer published a resumé of his life in “The American Mid-
land Naturalist.” On re-reading this biography one cannot
help but be struck by the similarity in the early backgrounds,
the interests and, in fact, the entire careers of the two men.

In 1901, Bush, then a collector for the Arnold Arboretum,
visited the Palmer home and spent a week there using it as
a base for his botanical collections. Bush was especially
interested in obtaining specimens of Crataegus for C. S.
Sargent, then director of the Arnold Arboretum, and when
he departed from Webb City he left with Palmer a supply
of dryers for further collections. Thus at the age of 25
Palmer began his life long interest in Crataegus and his
association with Sargent and the Arnold Arboretum. For
the next twelve years he collected extensively in his home
area concentrating on the flora of Missouri and it was not
until 1913, when he was 38, that he became associated with
the Missouri Botanical Garden. Two years later at the age
of 40 he began his many collecting trips to the southwest
under the aegides of both the Missouri Botanical Garden and
the Arnold Arboretum.

Palmer began publishing as early as 1910, at which time
his first paper, “Flora of the Grand Falls Chert Barrens”
appeared in the Transactions of the Academy of Science,
St. Louis. During the next ten years he published seven
papers, mostly on ferns. In 1920, while still a collector-at-
large he sent in from Webb City to the newly organized
Journal of the Arnold Arboretum his first of many papers
to be published by this Harvard University publication. Two
more papers were published in this journal before he came
to Jamaica Plain in 1921 as a regular staff member. He

198 Rhodora [Vol. 64

retained this status at the Arnold Arboretum for twenty-
seven years, until he retired in 1948.

My personal association with Palmer began in 1927. By
that time he was firmly established at the Arboretum as a
collector-botanist. Since he was away so much of the time
on his many collecting trips and since he was still a bachelor
he maintained personal quarters in a rooming house.

He was a very busy man during this period. He managed
to make a collecting trip nearly every year, usually alone,
occasionally with a single companion. Upon his return from
these trips his time was spent identifying his specimens,
making up sets for distribution, and writing occasional
papers — and getting ready for his next trip. He once men-
tioned to me that he had hopes of collecting in all the states
of the Union. At the time he had nearly accomplished his
intentions. If I recall correctly he said that there were only
two states in which he had not made “official” collections —
and as one might suspect one of them was surprisingly close
by!

The greater portion of his collections were made in the
southwestern United States. However, he always maintained
his interest in the Missouri flora and in 1935 there appeared
in the Annals of the Missouri Botanical Garden the 385 page
article by Palmer & Steyermark," An Annotated Catalogue
of the Flowering Plants of Missouri." This catalogue is more
complete than is usually found in such a publication. The
first seventy-five pages are devoted to the botanical history,
terrain, climate, plant regions etc. of the state. Under the
actual listing of the taxa may be found the habitat and a
record of the counties wherein specimens of each entry had
been collected. A footnote states that an earlier publication
of the ferns and fern allies of Missouri published by the
same authors three years earlier in the American Fern
Journal should be considered supplementary to the 1935
Catalogue.

A recent letter to me from Julian Steyermark, the junior
author of the above two publications mentions that his latest
publication “Flora of Missouri" which has been in press

1962] Kobuski — Ernest Jesse Palmer 199

since 1959 is being dedicated to E. J. Palmer. Palmer had
hoped to see the book and knew of the proposed dedication.

In 1930, when he reached the age of 55 he surprised us
all by getting married. We had thought of him as a con-
firmed bachelor. He married Elizabeth McDougal, a bac-
teriologist at the Massachusetts State Laboratory which is
located along side the Arnold Arboretum. Naturally, a great
change took place in Palmer’s life. He eventually set up a
very fine home in one of the houses belonging to the Arnold
Arboretum in Jamaica Plain. He and his wife had three
children, Ernest Macdougal Palmer born in 1931, Grace
Elizabeth Palmer in 1932, and Theodore Windle Palmer in
1935.

Palmer possessed a physique that defied age. He was the
slender, wiry type and at the time of his retirement and
departure from Jamaica Plain for Webb City could outlast
any of us on a collecting trip. Although I did not see him
again I understand from his long letters and from various
conversations with others that he continued his same brisk
manner in everything which he undertook. His letters show
that when he retired he merely transferred his activities
from Massachusetts to Missouri. In a letter to me in Febru-
ary 1955 he says: “Of course we miss many friends there
[Jamaica Plain], and I miss many of the associations at the
Arboretum. But I have never been sorry that I retired when
I did, for my life on the whole has been quite happy here. I
have never been busier at any time or interested in more
things. In April I will pass another anniversary — and you
can figure it out as the records show that I was born in 1875.
It is hard for me to believe it as I still feel as strong in body
and mind — so far as I can tell — as I did at fifty. I can
easily walk ten miles any day ; and I think I could climb a
good sized tree, if there was anything at the top of it that
I wanted badly enough. . . . Steyermark and I are still
working actively on the Flora of Missouri. . . . Specimens
of Crataegus, Quercus and other genera keep coming to me
for determination or revision. . . . Other collections in my
museum also occupy much time. . . . I have agreed to talk
to a joint meeting of all the Garden Clubs of Carthage at the

200 Rhodora [Vol. 64

public library. While I do not solicit or particularly enjoy
speaking engagements, I fill them occasionally. . . . It is so
fine and warm today that I think I will go fossil hunting this
afternoon.” For a man retired and eighty years old
he certainly was enjoying life to the fullest, doing exactly
what he wanted to do — and he seemed so well equipped to
do it!

During his life time Ernest Palmer joined many scientific
organizations, among them: the American Fern Society, the
American Association for the Advancement of Science, the
American Society of Plant Taxonomists, the Boston Mineral
Club, the Botanica] Society of America, the Missouri Ar-
chaeological Society, the New England Botanical Club (past
president 1944-1945), and the St. Louis Academy of Science.

CLARENCE E. KOBUSKI,
ARNOLD ARBORETUM AND GRAY HERBARIUM,
HARVARD UNIVERSITY

SOME OBSERVATIONS ON FLOWERING IN RUELLIA
(ACANTHACEAE)

ROBERT W. LONG! AND LEONARD J. UTTAL

During the past four years the authors have been making
systematic observations on transplants and natural popu-
lations in Ruellia, commonly referred to as “wild petunia”.
Although primarily tropical and subtropical in distribution,
this genus is represented in eastern United States by 11
species of low, perennial herbs with opposite leaves and
large blue or lavender, funnelform flowers.

Ruellia is a difficult genus for the taxonomist because of
the absence of reliable characters in certain species. Twenty
varieties and forms have been described in R. caroliniensis

*Present address, Department of Biological Sciences, University of South Florida,
Tampa, Florida.

*We wish to thank the following persons for their courtesy in supplying us with
Ruellia materials for greenhouse cultures: Drs. J. T. Baldwin, R. E. Woodson, R. K.
Godfrey, W. H. Duncan, J. D. Ray, G. S. Winterringer, Mr. K. E. Bartel and Mrs.
T. C. Lacey. Voucher specimens for these cultures are deposited in the Herbarium of
the University of South Florida,

1962] Long — Flowering in Ruellia 201

(Walt.) Steud. and R. humilis Nutt. (Fernald, 1945; Tharp
and Barkley, 1949) largely on the basis of minor character
differences. Fernald did much to clarify the nomenclature,
but his classification of eastern species is complex and dif-
ficult to use. It is now apparent that a major cause of this
is a poor understanding of the reproductive biology of these
plants. Monographic treatments of Ruellia have been based
almost entirely on herbarium specimens, and no attempt
made to investigate plants in culture or extensively in the
field.

This report attempts to clarify the relationship of cleis-
togamy to normal flower-production (chasmogamy) and to
describe the reproductive behavior of certain eastern U. S.
species as far as they have been investigated. Four species
are included in this discussion, all except one having been
observed in both garden and greenhouse cultures, and in the
field. In these species the following kinds of breeding
Systems have been observed: obligate and facultative
cleistogamy, the latter condition either coincident with
chasmogamy or sequential; obligate and facultative chas-
mogamy ; chasmogamy involving normal-sized corollas, and
these either long-styled, medium-styled, or short-styled ;
chasmogamy involving small-sized corollas, called here
"miniatures", that are approximately half as long as normal
flowers; facultative proterogyny was regularly observed
in some greenhouse plants, although synanthesis is usual in
Ruellia; finally, polygamy was noted in two species discussed
in this paper, involving apetalous, pistillate flowers in one
and staminate, petaliferous flowers in the other.

Ruellia strepens L. (5 natural populations examined, Ohio
and Virginia; 3 greenhouse cultures, Ohio and Virginia).
This species has been studied the most extensively of all
and is perhaps the best known one to taxonomists because
of its wide distribution in the Middle West. It is readily
separable from other eastern species by means of its lanceo-
late calyx lobes ; other species have linear or setiform sepals.
Ruellia strepens is remarkably uniform between and within
populations. Chasmogamous flowering begins in late April
and continues to mid-July, followed by small, greenish-

202 Rhodora [Vol. 64

yellow cleistogamous flowers produced from late June to
the first frost and borne in sparse to congested subsessile,
axillary glomerules. Plants occasionally do not produce any
flowers for a period between reproductive phases.

Ever since Gray (1878) formalized a varietal status for
cleistogamous plants of R. strepens, differences of opinion
have existed as to the nature of such plants. Chapman
(1884) reported late flowers sometimes set fruit in the bud.
McCoy (1937), after observing a single garden transplant,
reported cleistogamy sequential to chasmogamy in the same
individual. Nevertheless, recent monographers (Fernald,
ibid.; Tharp and Barkeley, ibid.) have continued to describe
cleistogamous specimens as either forma cleistantha (Gray)
S. McCoy or var. cleistantha Gray. We have observed hun-
dreds of plants of R. strepens in a total of 5 populations.
In all instances cleistogamy regularly follows chasmogamy
except that young plants appear to be exclusively cleisto-
gamous in their first year of flowering. In addition, green-
house cultures of plants collected during the chasmogamous
phase later produced cleistogamous flowers. Immediate
environmental factors strongly influence the production of
flower-types. Thus, cleistogamy is precocious in sear, open
habitats, whereas chasmogamous flowering is prolonged by
shade, moisture, and rich soil. Moreover, cleistogamous
plants may revert to chasmogamy as a result of prolonged
cool or cloudy weather. Also, chasmogamy may coincide
with cleistogamy for a period of time. Greenhouse cultures
of this species have never flowered chasmogamously, even
in May and June, but produce only closed flowers; garden
transplants produce both chasmogamous and cleistogamous
flowers.

The earliest flowers of R. strepens apparently are seldom
pollinated and thus rarely form seeds. Observations of
natural populations reveal that, although abundantly pro-
duced, chasmogamous flowers are infrequently visited by
insects. Early seed-production is mostly destroyed by
Lepidopterous larvae, but insect-depredations taper off in
early autumn. Plants thus appear to produce seed abun-
dantly in their autumnal cleistogamous phase. Although

1962] Long — Flowering in Ruellia 203

Fernald noted the low seed-production of chasmogamous
flowers, and the correspondingly high production of cleis-
togamous ones, he failed to explain why his open-flowered
variety appeared to be more common than the closed-
flowered variety.

Three kinds of chasmogamous flower-structures have been
observed in this species: (1) early chasmogenes with anthers
about the throat orifice and with the stigma much exserted
on a long style; (2) a later-appearing chasmogene, but one
which may appear coincidentally among earlier flowers, with
anthers and stigma borne at approximately the same level;
(3) a less common type, with a short style resulting in the
stigma being below the level of anthers. Thus, partial
heterostyly exists in R. strepens which, even in chasmoga-
mous flowers, tends to favor inbreeding. Open-flowers last
only part of the day; ordinarily anthesis occurs soon after
sunrise and corollas fall from the receptacle shortly after
noon. Ruellia strepens apparently is a habitually inbreeding
species with very few biotypes, these hardly dissimilar
morphologically, resulting in homogeneous populations.

Ruellia caroliniensis (Walt.) Steud. (10 natural popula-
tions examined, Florida, North Carolina, South Carolina,
Virginia; 11 greenhouse cultures, Virginia, Kentucky,
Florida, Georgia, North Carolina, South Carolina). This
species has been described as the most perplexing Ruellia
of eastern North America (Fernald, ibid.). Field work has
been carried out in Virginia, North and South Carolina,
and Florida. In this region a number of varieties and forms
have been described by Fernald, none of which is accepted
by Gleason (1952). Judging from the variations in green-
house and garden transplants, R. caroliniensis is apparently
the most racially complex member of the genus in North
America. Few, if any, of these races may deserve taxonomic
recognition, but definite taxonomic evaluations can not be
made at this time. Although the species varies widely,
plant-to-plant variability within populations is negligible.
The interpopulation variability of R. caroliniensis probably
is correlated with the wide range of habitats in which this
species can be found.

204 Rhodora [Vol. 64

Both chasmogamous and cleistogamous flowers are pro-
duced in a manner essentially similar to R. strepens. Chas-
mogamous flowers are produced abundantly in the spring
but apparently rarely set fruits. Cleistogamous flowers,
previously unreported for this species, occur in the summer
and early autumn with abundant seeds produced. In R.
caroliniensis races have been grown in the greenhouse that
form only chasmogamous flowers, but if hand-pollinated,
produce normal fruits with seeds that germinate readily.
Short-styled and long-styled flowers occur in the same indi-
vidual with long-styled flowers predominant. One green-
house race was quite variable, producing staminate flowers,
“miniature” flowers, and exhibiting proterogyny occasion-
ally. Another race produced only cleistogamous flowers.

Ruellia humilis Nutt. (2 natural populations examined,
Ohio and Virginia; 10 greenhouse cultures, Illinois, Texas,
Kansas, and Ohio). This species, occurring primarily in the
prairies and plains of central United States, has been segre-
gated into a number of varieties. Fernald reports cleis-
togamy as rare, but observations show the reverse to be
true. Flowering may be chasmogamous or cleistogamous,
sequentially or coincidentally. Cleistogamous flowers are
produced more abundantly in all races as the season pro-
gresses. Chasmogamous flowers with the long-exserted
stigmas predominate, but one greenhouse culture occasion-
ally produced short-styled ones. This same race regularly
formed chasmogenes and cleistogenes simultaneously rather
than sequentially as is generally true for the species. In R.
humilis greenhouse observations revealed that, although
pollinators were absent, chasmogamous flowers usually
formed fruits. This was owing to the manner in which the
corolla was shed from the receptacle, resulting in the epi-
petalous stamens coming in contact with the exserted stigma.
This same mechanism was observed in other species but not
so regularly as in R. humilis.

Ruellia pedunculata Torr. (6 greenhouse cultures, Texas)
This is a remarkably stable species, morphologically, that
characteristically produces long-stalked blue, lavender or
white flowers. It is distributed widely in Texas, the Ozark-

1962] Long — Flowering in Ruellia 205

ian uplands and into southern Illinois. The reproductive
systems of this species, as reflected by the kinds of flowers
produced, are among the most complex of any investigated
this far.

Cleistogamy occurs regularly in all greenhouse cultures
along with open-flowers, although apparently this has not
been noted before. The closed-flowers measure about 1.0 em.
long, are white with greenish-brown tips, and may occur
coincidentally with open flowers or sequentially with open
flowers. Evidentally, here, as in R. humilis and R. carolini-
ensis there exist at least two races differing in the timing
of flower-production. Partial heterostyly, similar to that in
R. strepens, occurred frequently in one race. Long-styles
measure 2.5-3.0 cm. long and short-styles only 1.0-1.5 cm.
long, well within the throat of the corolla and below the
stamens. Long-styled flowers failed to form fruits when
protected from insects and not hand-pollinated. Short-styled
flowers always formed fruits.

A fourth kind of flower appeared in two cultures. This
was an apetalous, pistillate flower with a short-style. That
this was not merely a cleistogamous flower that had shed
the corolla was established by repeated observations. The
occurrence of these unisexual flowers was sporadic but
coincident with open-flowers. All classes of flowers were
functional and could form normal fruits, but apparently in
nature the species is habitually self-pollinated.

No field work has been possible with R. pedunculata as
yet, but observations reported here are based on 80 green-
house transplants distributed in 11 different races. There
is relatively little morphological variation within and
between these races despite the fact that most of the col-
lections came from regions of sympatry with R. caroliniensis
and R. humilis. Ruella pedunculata can be hybridized with
these two species but natural hybrids are unknown at the
present time.

CONCLUSIONS

Breeding experiments with greenhouse and garden cul-
tures are in early stages and only preliminary results have
been obtained (Long, unpubl.), but apparently all classes

206 Rhodora [Vol. 64

of flowers in these four species are functional and can form
fruits with viable seeds. Field observations, however, indi-
cate that Ruellia species are habitually self-pollinated. The
relatively small amount of morphological variation in R.
strepens and R. pedunculata suggests that outcrossing
seldom takes place. On the other hand, Long (1961) adduced
evidence from herbarium samples and hybridization experi-
ments that crossing of R. humilis and R. caroliniensis prob-
ably does occur and that at least some of the apparent
intergraduation between these species may be attributed
to natural hybridization. More detailed and extensive
observations are needed on species thus far only partially
investigated, such as R. ciliosa, R. heteromorpha and R.
purshiana as well as on numerous North American species
as yet poorly known. Apparently, however, R. strepens, R.
caroliniensis, R. humilis, and R. pedunculata are examples
of plants that are functionally autogamous but with many
of the typical flower structures of allogamous ones.—DEPART-
MENT OF BOTANY AND BACTERIOLOGY, OHIO WESLEYAN UNI-
VERSITY, DELAWARE, OHIO; AND ROUTE 3, MADISON HEIGHTS,
VIRGINIA.

LITERATURE CITED

CHAPMAN, A. W. 1884. Flora of the Southeastern United States.
University Press, Cambridge, Mass.

FERNALD, M. L. 1945. Ruellia in the Eastern United States. Rhodora
47: 1-88; 47-63; 69-90.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated
Flora, Vol. 3, New York.

GRAY, A. 1878. Synoptical Flora of North America. Vol II - Part I.
New York.

Loc, R. W. 1961. Convergent patterns of variation in Ruellia
caroliniensis and R. humilis (Acanthaceae). Bull. Torrey Bot.
Club 88: 387-396,

McCoy, S. 1937. A cleistogamous Ruellia. Am. Bot. 43: 22-23.

THARP, B. C. and F. A. BARKLEY. 1949. Genus Ruella in Texas.
Am. Midl. Nat. 42: 1-86.

1962] Britton — Dryopteris dilatata 207

DRYOPTERIS DILATATA (HOFFM.) A. GRAY IN
NORTH AMERICA

DONALD M. BRITTON

Taxonomic authorities have differed widely in their inter-
pretation of the species of Dryopteris in North America
which resemble the old world Dryopteris dilatata (Hoffm.)
A. Gray. Fernald (1) considered the fern that is found in
the Appalachians north to Quebec and Labrador, and yet
resembles D. dilatata superficially, to be a variety of D. spin-
ulosa (O. F. Miill.) Watt, namely D. spinulosa var. ameri-
cana (Fischer) Fernald, whereas Morton in Gleason (2),
refers to this entity as D. austriaca (Jacq.) Woynar var.
austriaca. There has been similar disagreement concerning
the species found in Montana and Oregon north to Alaska,
although Taylor (3) employs the latter designation for the
common form in British Columbia.

In a recent article in Brittonia, Wagner and Hagenah (4)
have suggested that not only are the eastern and western
entities separate species, but that within the range of Gray's
Manual, D. spinulosa var. americana should be further sep-
arated into two species. Some evidence for the latter separa-
tion is found from Walker (5) who reported that D. spinu-
losa var. americana, (D. campyloptera (Kunze) Clarkson)
from Vermont and also Quebec is tetraploid (4X) (x=41),
whereas Wagner reports that ferns in Michigan previously
referred to as var. americana are diploid. I am in full agree-
ment with these cytological findings because last spring I
studied 12 collections of var. americana from Jefferson
Notch and Mt. Washington, New Hampshire collected by
R. and A. Tryon and all were tetraploids (Tryon 5684, 5689-
5700 June 25, 1961). In 1960, I studied 11 collections of var.
americana from Mile 80, north of Sault Ste. Marie, Ontario
and these were all normal diploids (2X) (B373-383 June 23,
1960).

However, I take strong exception to the authors' sugges-
tion that the Lake Superior diploid cannot be identified with
western D. austriaca var. austriaca although both are di-
ploid. The cytological argument is quite circuitous. Walker

208 Rhodora [Vol. 64

(5) has named the diploid species of dilatata of the old
world, D. assimilis S. Walker, and has considered that mor-
phologically and cytologically D. maderensis Alston is insep-
arable from D. intermedia (Muhl.) A. Gray. (There is nor-
mal meiotic pairing between the chromosomes of maderensis
and intermedia). In the two hybrids studied from a cross
between D. maderensis X D. assimilis, Walker (5) found
that there was also normal pairing but that ca. 40% of the
spores aborted. He concluded on the basis of chromosomal
homologies that intermedia and assimilis represented the
same ancestral genome. Walker further suggested that
"morphological similarity between the Vancouver diploid
and D. assimilis suggests that they represent the same
genome and possibly the same species." This conclusion was
fortified by the fact that the progeny of the cross of the
Vancouver diploid with the United Kingdom tetraploid
dilatata gave the same cytological picture as the cross D.
assimilis (2X) X D. dilatata (4X) namely ca. 41 pairs and
41 single chromosomes in meiosis. Wagner and Hagenah
(4) found one hybrid between the putative parents D. inter-
media and the Lake Superior diploid and report that there
is little or no chromosomal pairing between the genomes of
this plant, although the number of cells studied and the num-
ber of clear meiotic plates analysed is not given. According-
ly, they contrast Walker's cross (two plants) of D. maderen-
sis—intermedia X D. assimilis with their one hybrid of D.
intermedia X Lake Superior diploid, and conclude that if
D. assimilis is identical with the Vancouver diploid then the
Vancouver diploid is unlike the Lake Superior diploid. How-
ever, as yet we have no direct comparisons between these
diploids. What is needed are hybrids between (a) the Lake
Superior diploid and the western diploid (b) the Lake
Superior diploid and D. assimilis and (c) the Western di-
ploid and D. intermedia. It should also be evident that many
more plants should be studied cytologically before reaching
sweeping conclusions from meiotic analysis of one or two
plants, some of which are hybrids of putative parents.
Although a cytotaxonomist is apt to be impressed with
the evidence presented by Walker (5) to consider D. ma-

1962] Britton — Dryopteris dilatata 209

derensis Alston and D. intermedia (Muhl.) A. Gray as con-
specific, there has as yet been no adequate taxonomic treat-
ment to substantiate this conclusion.

It would seem extremely hazardous to put too much
emphasis on pairing in spite of its great usefulness in
genome analysis. D. intermedia and D. assimilis are mor-
phologically separable and yet show good pairing, hence
would be one species using this latter criterion alone. In
fact this would leave D. assimilis Walker, D. maderensis
Alston, D. austriaca var. austriaca in British Columbia, and
D. intermedia as being all the same ancestral genome and
the same species on the basis of genome analysis. The Lake
Superior diploid, however, which so closely resembles the
British Columbia diploid in morphology, and must be closely
related, is then considered to be an entirely different species
merely on its pairing behaviour with intermedia.

Two pieces of cytogenetic evidence should be kept in mind
to temper the use of the criterion of chromosome pairing.
One is the finding in wheat that pairing is under genetic
control and governed by the presence or absence of genes
for synapsis (6). If this should prove to be true in Dryop-
teris, the basis of genome analysis would be invalidated.
The other point may be related, and concerns races of
Asplenium trichomanes in Ontario (7). These races are
morphologically extremely similar and yet one is diploid
and the other is tetraploid without multivalent pairing at
meiosis. If the latter is an autotetraploid with four identical
genomes as it appears on morphological grounds, then why
are only bivalents formed? It would seem that the tetra-
ploid has become effectively “diploidized.” A similar situa-
tion to this was found to be present in Pellaea ternifolia
which had both diploid and tetraploid sexual races which
formed only bivalents at meiosis (8).

It should be stressed that all hybrids found in the wild
are given parentage on the basis of their intermediate
morphology between putative parents. The interesting plant
(Wagner 9434) assigned to D. *dilatata" (Lake Superior
diploid) X D. intermedia is described by Wagner and
Hagenah (4) cursorily as an intermediate, and one has no

210 Rhodora [Vol. 64

way of knowing whether it is the result of an average or
typical Lake Superior diploid X an average or typical inter-
media, or for that matter whether it could not be an atypical
intermedia X an atypical D. marginalis. A further possi-
bility is that this plant (Wagner 9434) is a hybrid of a
glandular Lake Superior “dilatata” X marginalis, since the
illustrations of the authors' look similar. The point is, that
nothing is known of the characteristics of the parents except
as they are represented in the hybrid. In this case, “the leaf
form was intermediate" although “the previous year's leaves
were green and persistent."

The morphological data given by Wagner and Hagenah
(4) for the western diploid is extremely scanty. There is no
mention of the number of specimens examined or where
they might have been from. The findings are summarized in
six lines and concern such variable features as petiole scales
and leaf blade shape.

The authors (4) disregard the plant geographical evi-
dence. One can assemble quite an impressive array of
species in the Lake Superior basin that are of Cordilleran
affinities. Stebbins (9) lists ten species found even further
east in the Bruce Peninsula, Ontario, that may be considered
Rocky Mountain types. Butters and Abbe (10) list four
western American types in a table of 70 rare species found
in Cook County, Minnesota. Among the ferns of the Lake
Superior basin, Cystopteris montana, Woodsia scopulina,
and Cryptogramma crispa are considered to have western
affinities.

Accordingly, when the geographical, morphological and
cytological evidence is reviewed critically, one can only con-
clude that the western diploid, D. dilatata (D. assimilis),
and the Lake Superior diploid, D. dilatata (D. spinulosa var.
americana) have not as yet been shown to be distinct and
separate species.

Concerning the eastern tetraploid, D. campyloptera Clark-
son, Wagner and Hagenah (4) suggest that this might be an
allotetraploid of D. intermedia X Lake Superior diploid. An
alternative suggestion that it is an ancient autotetraploid of
the Lake Superior diploid which in turn is a diploid “D.

1962] Britton — Dryopteris dilatata 211

dilatata" (D. assimilis) cannot be disregarded on the avail-
able evidence. D. campyloptera which now forms only
bivalents could be analagous to the tetraploid race of Asplen-
ium trichomanes.

One is confronted in Dryopteris with the larger problem
of what is a species? Wagner is a proponent of biological
species, but critical criteria for separating species are few
in the Dryopteris spinulosa complex. All the species are
apparently able to cross with one another, consequently this
is of little use as a limiting criterion, although much used
for genome analysis. Morphology by itself, has so far failed
to give a clear picture (e.g. Lake Superior diploid vs. D.
campyloptera of the Appalachians). Instead, the practice is
to return to the morphological evidence after the cytogenetic
analysis is completed. Geographical distribution is of value
to delimit a sexually interbreeding natural population even
though the boundary between species is often obscured by
disjunct ranges such as the amazing one shown by Poly-
stichum scopulinum. Also with some calcicolous species of
ferns, it may well be, that distribution is limited by lack of
suitable habitats e.g. some Woodsias. Cytology has been
extremely useful, but it is felt that too much emphasis on
the criterion of homology, when so little is known about the
true nature of pairing and chiasma formation at zygotene of
meiosis, is apt to lead to false conclusions. — DEPARTMENT
OF BOTANY, ONTARIO AGRICULTURE COLLEGE, GUELPH, ON-

TARIO, CANADA.

LITERATURE CITED

1. FERNALD, M. L. 1950. Gray's Manual of Botany, American Book
Co, New York.

2. GLEASON, H. A. 1952. Illustrated flora of the northeastern United
States and adjacent Canada. New York Botanical Garden, New
York.

3. TAYLOR, T. M. C. 1956. The ferns and fern-allies of British Co-
lumbia. Handbook No. 12, British Columbia Provincial Museum,
Victoria, B. C.

4. WAGNER, W. H. JR. AND DALE J. HAGENAH. 1962. Dryopteris in
the Huron Mountain Club area of Michigan. Brittonia 14: 90-
100.

212 Rhodora [Vol. 64

5. WALKER, S. 1961. Cytogenetic studies in the Dryopteris spinulosa
complex-II. Amer. Jour. Bot. 48:607-614.

6. RILEY, R., AND V. CHAPMAN. 1958. Genetic control of the cyto-
logically diploid behaviour of hexaploid wheat. Nature 182:713-
715.

7. BRITTON, DONALD M. 1953. Chromosome studies on ferns. Amer.
Jour. Bot. 40:575-583.

8. TRYON, ALICE F. AND DONALD M. BRITTON. 1958. Cytotaxonomic
studies on the fern genus Pellaea. Evolution 12:137-145.

9. STEBBINS, G. L. JR. 1935. Some observations on the flora of the
Bruce Peninsula, Ontario. Rhodora 37:63-74.

10. BUTTERS, FRED K., AND ERNST C. ABBE, 1953. A floristic study
of Cook County, Northeastern Minnesota. Rhodora 55:21-55, 63-
101, 116-154. 161-201.

ON PINGUICULA MACROCERAS LINK
IN NORTH AMERICA

S. JosT CASPER

Link, 1820, described a new species of Pinguicula under
the name P. macroceras, based on a sheet collected by Pallas
in Unalaschka and later on preserved under no. 422 in the
herbarium of Willdenow at Berlin. Reichenbach (1823)
gave a picture of P. macroceras in his “Icones” (tab.
LXXXII, fig. 169-170) ; but Chamisso (1831) demonstrated
that it represented his own P. microceras, a plant of Unalas-
chka too.

In Willdenow's herbarium existed two other specimens of
Pinguicula, both collected by Pallas and first mentioned by
Link (1820) under the names P. camtschatica and P. dau-
rica. Ledebour (1847/49) was acquainted with this fact,
but related the two specimens to P. macroceras, which should
be *P. vulgari et leptocerati maxime affinis et vix diversa".
Herder (1871/72) followed him, but asserted that P. macro-
ceras would be only a big-flowering form of P. vulgaris!

Nowadays most botanists treat it as P. vulgaris, the Japa-
nese writing P. vulgaris var. macroceras Herder, while only
a few authors attribute the rank of species to P. macroceras
(Rydberg 1917, 1922, 1954; Komarov 1950, 1951; Boivin
1948).

1962] Casper — Pinguicula macroceras 213

In several papers Hultén (1949, 1958, 1960) discusses the
problem and says: “It is ... hardly possible to separate
them’’— i.e. P. macroceras and P. microceras —“specifically
from the European plant" — P. vulgaris — “although the
length and form of spur and lips are very different in some
cases". (Hultén 1958, 230). Recently Ernst (1961) relates
P. macroceras, P. camtschatica, P. daurica and P. micro-
ceras to P. vulgaris.

In a former paper (Casper, 1962) I first gave an account
of my own views on the point in question. Now further
studies on specimens in American herbaria enable me to
confirm the meaning of Boivin (1948, 220) who wrote: “...
P. macroceras est certainement spécifiquement distinct du
P. vulgaris".

According to Link (1820) P. macroceras is characterized
by the following diagnosis: “scapo foliisque glabris, calcare
turbinato corollae laciniis longiore".

Unfortunately the type-specimen of Pallas was lost dur-
ing World War II at Berlin and therefore, is not available
for comparative studies (cf. Hultén, 1960). But there is no
doubt that the specimens collected later by several botanists
at the same locality are equivalent to those of Pallas: their
flowers are typically long-spurred, and the long spur is the
main characteristic in the diagnosis of P. macroceras given
by Link.

Tables 1 and 2 reproduce the quantitative analysis of the
length of spur (SPL) and corolla (C-L). Dried material of
P. vulgaris from all parts of its area with particular consid-
eration of Scandinavian specimens and of P. macroceras
from the American part of its distribution was measured.
These tables show the exact numerical composition of the
investigated material.

KEY TO P. MACROCERAS AND P. VULGARIS:
Corolla lobes of the lower lip subobovate-oblong, more or less cover-
ing each other, or at least touching; lower lip of the calyx split up to
half of its length.
Length of the corolla (15) 18-27 (36) mm. (spur included), length
of the spur (3) 6-9 (11) mm. .................. eee eder eee eee tn eor tn etna trae eaa s ana enaaS
oio x DESEE PEE DOAHDDR ERU RATEM P. macroceras LINK var. macroceras
Length of the corolla (10) 14-22 (29) mm. (spur included), length

[Vol. 64

Rhodora

214

%00T ro 6*0 AH 69 IST 6‘LZ P92 SCT 0% ro
989 Z G 6 LE L6 OST TFI 28 IT Z SESMA 'q
%00T 8'g 9ST SPE ZZE 8T Ze - vo - - "S's
88% IT LE 66 Z6 68 6 : I - - SRIƏJOIDLU 'q
25001 9% 86 6'8ec 9'ec ecl 9'8 c9 a9 oP rc TS
61P II If OOT 66 TS 9g 92 9c 6I OT Selad019RUL ‘d
euruns OT 6 8 L 9 G i4 € Z I yy) — ww
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6g€ - " Z 9 9% LG v8 CTT LOT 98 LY 6 sues[na 'q
25001 8*0 9‘T eg “Pe 196 2% Or FS ro 30) - - ‘S's
v86 G 6 &% 69 €L 69 86 ST I Z - - $*9192012€UI 'q
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OTP Z 6 cS eL 98 c8 €9 $8 FI LI Z I SBIIIOIIVUL 'q
ZI IT OT 6 8 L 9 g Y g Z I MD
BUIUINS Z€ Ig/0g 62/86 Le/96 €c/vcG 87/77 137/07 61/81 LI/9T SII SIZI M Ur
(1-0) *i[0100 ay} Jo u33uo'T. :[ TIIVL

1962] Casper — Pinguicula macroceras 215

of the spur 1-3 (5) mm.; lobes of the lower lip of the calyx almost
entirely separated. A plant relatively frequent on the Aleutian
Islands P. macroceras var. microceras (CHAM.) CASPER
Corolla lobes of the lower lip oblong, not covering nor touching each
other; lower lip of the calyx split up to 2/3 ef its length; length of
the corolla (10) 14-21 (29) mm. (spur included), length of the spur
ADT BER ETI or iscsereriner étaient sereno ci b ERR P. vulgaris L.

It is at once apparent from a study of figure 1 that there
is a really quantitative difference between P. macroceras and
P. vulgaris with regard to the length of spur and corolla.
Link, therefore, was right in characterizing P. macroceras
as a species very different from P. vulgaris by means of the
length of its spur. In P. macroceras the spur measures (1)
6-9 (11) mm., the corolla (15) 18-27 (86) mm. (spur includ-
ed), in P. vulgaris the spur (1) 3-6 (10) mm,, the corolla
(10) 14-21 (29) mm. (spur included).

Other characteristics also distinguish P. macroceras from
P. vulgaris. The typically obovate large, broad rounded
lobes of the lower lip of the corolla of P. macroceras were
seen already by Ledebour (1847/49), who enlarged Link's
diagnosis by this data. In most cases the lobes more or
less cover one another (fig. 2). As opposed to these facts
the corresponding lobes of P. vulgaris have an oblong form
and are distinctly separated from one another (fig. 3).

In P. macroceras the two lobes of the lower lips of the
calyx are oblong and grown together only to the half of
their length. P. vulgaris has a smaller calyx and the lobes
of the lower lip are grown together to 2/3 of their length.

The geographical distribution of P. macroceras reaches
from Japan and Kamtschatka over the Aleutian Islands to
the Pacific coast regions of North America (Casper, 1962) :
the area of an amphi-pacific plant. The map (fig. 4) shows
exactly the known distribution in North America. The
records are based on the study of specimens in herbaria.
From the Aleutians over Pacific-Alaska, British-Columbia,
Alberta, Washington, Montana, Oregon its area extends
south into Northern California (Del Norte County). The
northern and eastern borders of the area are not sufficiently
known. Further studies are necessary to clear up especially

216

Zot

40

30

20

10

Rhodora [Vol. 64

%4

20

10

——— P vulgaris L.
——-—- P macroceras LINK s.l.
—7—" P. macroceras LINK s.s. 1
A.
- e
A \
l_e. \ j
j^ -— NS ^u
P4 SA, \
Ly \ \
voy NN
fo, NN
z^ J No
. a N
Pus "d
r a
— 37 P 4 4
5 6 8 9 10
Gk— CL
P. vulgaris L.

——— P. macroceras LINK s.l.

—:—" P macroceras LINK ss. 1

1962] Casper — Pinguicula macroceras 217

--——

Figure 2 (above) Figure 3 (below)

218 Rhodora [Vol. 64

the contact-zone with P. vulgaris, which is distributed in
America from Labrador to Nome and Coronation Gulf, but
is not known from the most eastern parts of Asia (Casper,
1962).

A peculiar problem is associated with the name P. micro-
ceras Chamisso. The author described the plant as follows:
*... species unalaschcensis alpina, nana, iam cornu brevi,
laciniis corollae multo breviore . . ." (1831, 568). Mainly it
was the small acute conical spur — the type specimen shows
a spur of 1.5 mm. length — that induced Chamisso to sep-
arate his plant from P. macroceras or P. vulgaris. The other

Pinguicula macroceras LINK

Map of distribution in Northern America

( Attu- Isl. not drawn!)

JOST CASPER 1962

Figure 4

1962] Casper — Pinguicula macroceras 219

characteristics are similar to those in P. macroceras. Figure
1 shows two curves concerning P. macroceras. The first one
— P. macroceras s.s. — excludes all microceras types from
the Aleutians, the second one -— P. macroceras s.l. —
includes these types. It is apparent that in the case of P.
macroceras s.l. the curve is depressed and displaced to the
left by the microceras-material. But the particular charac-
ter of the curve in relation to P. vulgaris is not changed.

Evidently P. microceras is a short-spurred form of P.
macroceras, restricted to “the mountains and . . . arctic
places, while macroceras types prevail more to the south"
(Hultén, 1949: 1425; Casper, 1962). Already Chamisso
(1831) understood P. microceras as "alpin" as opposed to
the "subalpin-montanen" P. macroceras. Hultén (1949:
1425) is right in his statement, that “all the transitions
exist between P. macroceras and P. microceras. But when
he states that “... the same variation in the form and size
of the flower can be observed in material of P. vulgaris from
other parts of its range, for instance from Scandinavia . . ."
he is mistaken as figure 1 shows. The curves confirm
Hultén's own words: “It should be admitted, however, that
plants approximating to the two extreme forms are more
common on the Aleutians than they are in Scandinavia”
(1960: 328). “All the transitions" concern only P. macro-
ceras and P. microceras, but not P. vulgaris! The short-
spurred microceras-type belongs to P. macroceras but never
to P. vulgaris.

P. macroceras Link is a good species, sufficiently distin-
guished from P. vulgaris by its long spur, its obovate round-
ed lobes of the lower lip of the corolla and its deeply
separated lobes of the lower lip of the calyx. Its area is
typical amphi-pacific. Chromosome numbers are not known.
— INSTITUT FÜR SPEZIELLE BOTANIK DER FRIEDERICH-SCHIL-
LER-UNIVERSITAT JENA.

RECORDS’
ALASKA. Aleutians: Adak Id. (CHI 1162067), (DS 374181).
Amchitka Id. (Us 173872). Atka. Id. (MIN 355434, DS 243871). Attu
Id. (MO 1705350), Bassett Creek (MO 1304358), Cape Khlebnikof (Uc

"The list includes only half of the whole material studied.

220 Rhodora [Vol. 64

7303384). Kuiu Id., Washington Bay (MIN 530054). Unalaska (UC
081604), Tliuliuk Lake (vc 081605). Kodiak Id.: Olga Bay (TRT
91769), Red River, Alitak (US 220252). Raspberry Id., Port Vita
(MIN 525347), Eagle River, Juneau (DAO 11226) Hinchinbrook Id.,
Cordova (pS 150701). Glacier Bay (uc 533512). Yakutat (MO
822106). New Metlakatla (Mo 1764214). Baranoff Id. (Mo 1764215).
Muir Glacier (Mo 1764216). Chumagin Ids: Popoff Id. (Mo 1764220).
Windham Bay (Mo 17464221).

BRITISH COLUMBIA. Log Cabin (v 12025). Ogilvie Mts. (V
12225). Prince Rupert (v 6330). SW Gowgaia Bay (v 34881). Goose
Island (v 21541). Vancouver Island: Mt. Splendor (v 11034), Gordon
River (MO 1764210, DS 20540), Port Hardy (UBC 23524), Moat Lake
(v 29750), Forbidden Plateau, Meadow Lake Stream (v 15821), Uclue-
let (V 4586), Henderson Lake (v 4584), Mt. Joan (V 15910), Cowichan
Lake, N.E. shoulder Mt. Landalt (v 23690 + 28691), Mt. Arrowsmith
(v 22882), Strathcona Park, Mt. Rooster Comb (v 10473). Coast,
Cypress Creek (UBC 24652), Lillocet (v 5971). Cheam (V 4588).
Fairmont H. Springs (v 13980). Canal Flat (v 15343). Kootenay,
Gray Creek, Clarke's Pt. (v 12978, UBC 5691). Coast, Hollyburn Ridge
(UBC 62839), between Black Mt. and Hollyburn (DAO 11259). Whistler
Pass (UBC 5692). Dam Mt. Goat Lake (UBC 5693). Kaslo, White
Water Mine (UBC 5694). Haffner Creek, Marble Canyon Campground
(DAO 11261). Hat Creek Valley, Marble Mts. (WTU, herb. J. W.
Thompson 17197). Cassiar, Ingenika River (UBC 51290). Tenquille
Lake (UBC 83843), Finch Ridge (UBC 48506). Atlin District, Nakina
Lake (DAO 11262), Lake Atlin (UBC 51295, UC 494021). Yohs National
Park, Emerald Lake (UBC 65608), Twin Falls (MIN 267961), Sorrent
fan (MIN 241875), Sir Donald Trail (MIN 241874). Selkirk Mts., Low-
er Sandford Morains (MIN 241873). Victor Lake, 11 mi. W. of Revel-
stoke (Ds 352516). Columbia Valley, Albert Canyon (UBC 74339).

ALBERTA. Banff (v 26321, DAO 11188), Middle Springs (DAO
11190), Sulfur Mt., (MIN 241872), Bow River (DAO 11458), Hillsdale
Cabin (VT), between the Mistaya and Bow River Valleys (UC 707445).
Jasper, Lake Beauvert (TRT 124265). Lake Louise (v 26323). Selkirk
Flora, Waterfall at gorge of Columbia (PH 526504).

WASHINGTON. Olympic National Forest: Deer Lake (WTU
110683), Heart Lake (WTU 33020), Mt. Angeles (WTU, herb. J. W.
Thompson 24361), Mt. Colonel Bob Lookout (WTU, herb. J. W. Thomp-
son). Mount Baker National Forest: Mt. Baker, Boulder Glacier
(WTU 32274), Mt. Baker Lodge, Bagley Lake (wTU, herb. J. W.
Thompson 24361), Mt. Hermon (uc), Skagit River Gorge, Newhalem
(MIN 357710), Skagit Pass (WTU 16083), Mt. Higgins (UC 518084),
Baldy Peak (Mo 1764206). Mt. Rainier (MO 1764208), Van Trump
Park (wTU 88707), Comet Falls (wTU 37612), above Main Cowlitz
Glacier (MO 1764204). Kittitas Co.: Mt. Stuart (MIN 191336), North
Fork of Teanaway River, Wenatchee Mts. (BH 4453), Iron Peak (UC

1962] Casper — Pinguicula macroceras 221

931294), Fish Lake (WTU 75350, UC 710875). Cascade Mts., Stevens
Pass (UC 185022). Okanogan Co.s Horse Shoe Basin (vT). Snohomish
Co.: Mt. Dickerman (Mo 1027818), Columbia Peak, N of Monte Cristo
(WTU 35468). Yakima Region (UC 173684).

MONTANA. Glacier National Park: N of Logan Pass (vc 028325),
Grinnell Lake (UCLA 40261), Grinnell Falls (WTU 129482), Iceberg
Lake (CHI 953072, 953073).

OREGON. Wallowa Co.: Ice Lake, bog (WTU, herb. J. W. Thompson
8179), on trail from Wallowa River (ILL, herb. G. N. Jones 7497).
Josephine Co.: near Kerby (UC 76680), other side of Tennessee Pass
(MO 925823).

CALIFORNIA. Del Norte Co.: French Flat, Gasquet, Smith River
(UC 501348), near Camps (MO 1261851), Douglas Park, 9 mi. E of
Crescent City (DS 179461), Rock Creek Lodge (WTU 183910).

LITERATURE CITED

BouviN, B. 1948. In Nat. Canad. 75: 220.

CASPER, J. 1962. “Revision der Gattung Pinguicula in Eurasien” —
Fedde Rep. 66, in press.

CHAMISSO, A. DE. 1831. “De plantis in expeditione Romanzoffiana
observatis . . ." — Linnaea 6: 545-592.

Ernst, A. 1961. “Revision der Gattung Pinguicula” — Bot. Jahrb.
80, 2: 145-194.

HERDER, F. DE. 1855-1859. “Lobeliaceae, Campanulaceae . . . Sero-
phulariaceae a Cl. D-re G. Radde annis in Sibiria orientali collec-
tae" — Acta Horti Petrop. 1 (1871/72): 378.

HULTÉN, E. 1949. Flora Alsaka Yukon IX: 1425.

1958. The Amphi-Atlantic Plants. 230.
1960. Flora of the Aleutian Islands in Tüxen, R., Flora
et Vegetatio Mundi Vol. I. Weinheim: 327-328.

JONES, G. N. 1936. “A Botanical Survey of the Olympic Peninsula,
Washington" — Univ. Wash. Publ. Biol. 5: 233.

LEDEBOUR, C. F. 1847-1849. Flora Rossica. Stuttgart.

Link, H. F. 1820. Jahrb. Gewächs. 1, 3: 54.

Moss, E. H. 1959. Flora of Alberta. London.

RYDBERG, P. A. 1917. Flora of the Rocky Mountains. I ed.; II ed.
1922.

STANDLEY, P. C. 1921. “Flora of Glacier National Park Montana"
— Contr. U. S. Nat. Herb. 22, 5: 410.

222 Rhodora [ Vol. 64

HOLOSTEUM UMBELLATUM L., AN ANGIOSPERM
NEW TO MICHIGAN'

MARTIN A. PIEHL

In May 1958 I collected what I felt was an “unusual chick-
weed” at the then recently acquired 200-acre tract for the
new University of Michigan Botanical Gardens on Dixboro
Road, 4 miles northeast of Ann Arbor. After identifying
this plant as Holosteum umbellatum, the jagged chickweed,
I suspected it to be an interesting find since there were no
Michigan specimens in the University Herbarium.

Several months later my attention was called to a speci-
men of Holosteum being accessioned by the herbarium. This
collection was made at Ann Arbor in 1949 by the late L. H.
Jordal (no. 1498)", and appears to have been the first Michi-
gan record. Whether or not Jordal realized that his collec-
tion was a record has not been determined, but the evidence
at hand suggests he did not. In April 1961 I searched for
Jordal’s locality and found a good stand of Holostewm on
cultivated soil in the general vicinity.

I have checked for records of this species in published
local floras and at other herbaria in the state (see below).
An additional collection was found in the herbarium of
Eastern Michigan University, Ypsilanti. the specimens hav-
ing been taken from the lawn near Pierce Hall on the
campus of that institution by Richard A. Giles, 5 April
1955. I visited this site in May 1961 and found the plant
both in lawn turf and in the adjacent open soil. Also, upon
revisiting my original locality in 1960, 1961, and 1962, I
found this species each year, and although actual numbers
were not recorded, there appeared to be some increase in
the number of plants. Specimens of each of my collections
will be deposited in the University of Michigan Herbarium
(Piehl 5846, 6029, 6101, 6102).

"The observations reported herein were made while the writer was engaged in
doetoral studies in the Department of Botany, University of Michigan, Ann Arbor.

"It is of interest that this collection is from the old botanical gardens, a 40-acre
tract on Iroquois Drive in Ann Arbor. However, it is very unlikely that the Holosteum
was transported from that site to the new locality, since the transfer of the gardens'
plants had not begun at the time of my collection.

1962] Piehl — Holosteum umbellatum 223

Holosteum, a native of Europe, is well distinguished from
our other chickweeds by its pale, generally glaucous foliage,
its non-cleft but often jagged or emarginate petals, and by
its umbellate inflorescence. Because some of the flowers open
before the pedicels have elongated fully, the umbellate na-
ture of the inflorescence may be obscure when flowering
begins. The inflorence become distinctly umbellate later,
however, and as has been recorded (Fernald, 1950), some of
the pedicels actually become reflexed in fruit, although they
are described by Britton and Brown (1897) as erect and are
illustrated as such in the most recent edition of The Illus-
trated Flora (Gleason, 1952). None of the above manuals
mentions the fact that the very delicate, somewhat hyaline,
white or rarely pinkish petals are persistent, although they
become hidden by the more rigid, scarious-margined sepals.
A poorly-defined basal rosette is formed by large specimens,
but it is lacking from the smaller plants. Although the stems
tend to be somewhat decumbent, they are not trailing like
those of some species of Cerastium and Stellaria.

I suspect that Holosteum is often overlooked due to its
rather unusual phenology. It begins to flower in early April
at Ann Arbor, and perhaps earlier depending on the season,
but unlike the common chickweed, Stellaria media, it flowers
and fruits for a relatively short time and is, itself, ephem-
eral. In early May I have observed both dehisced and de-
veloping capsules as well as flowers. By mid-May, however,
the foliage yellows, and the entire plant soon dries and easily
escapes detection. Thus, specimens of Holosteum are likely
to be found over a period of only a few weeks, the plant
apparently passing most of the collecting season in the form
of seeds.

The precociousness of Holosteum caused me to doubt that
the plant was, strictly speaking, an annual, as some of the
manuals indicate (e.g. Gleason, 1952). During the winter
1961-62 I visited the Dixboro Road locality at a time when
most of the snow had melted and found numerous seedlings
which apparently had developed the previous autumn. Holo-
steum thus behaves as a winter annual. Whether some seeds
also germinate the following spring is not known. The leaves

224 Rhodora [Vol. 64

(not meaning the cotyledons) of the seedlings are spatulate
or oblanceolate to oblong, the lamina narrowing in some
cases nearly to the extent of making the leaves petioled, and
some have a tiny black fleck at the very apex. They tend to
have a thicker epidermal covering and to be more succulent
than the leaves formed the ensuing growing season. The
shape of the cauline leaves, in contrast, is from oblong to
ovate, and they are sessile. The over-wintering leaves of the
seedlings, which soon yellow and dry as growth is resumed
the following spring, are not found on the majority of her-
barium specimens, and are omitted from most descriptions.
Elongation of the stem and the appearance of the inflores-
cence occurs rather rapidly and will take place at relatively
low temperatures.*

The occurrence of Holosteum in southern Michigan indi-
cates that its range is still being expanded, and that it
should be looked for by early-ranging botanists elsewhere
in the Midwest. Previous to the Michigan collections report-
ed herein, the jagged chickweed was recorded as far west as
Ohio and Kentucky by Fernald (1950), while earlier it was
known at first only from the vicinity of Lancaster, Penn-
sylvania (Gray, 1856), and later from New Jersey and
Pennsylvania to Georgia (Robinson and Fernald, 1908).

I wish to acknowledge the cooperation received from
curators of the following herbaria: Cranbrook Institute of
Science, Eastern Michigan University, Michigan State Uni-
versity, The University of Michigan, and Wayne State Uni-
versity. — SANTA BARBARA BOTANIC GARDEN, SANTA BARBARA,
CALIFORNIA.

LITERATURE CITED

Britton, N. L., AND A. BROWN. 1897. An illustrated flora of the
northern United States, Canada, and the British Possessions,
Vol. 2. Scribner's, New York.

FERNALD, M. L. 1950. Gray's manual of botany, 8th Ed. American
Book Co., New York.

GLEASON, H. A. 1952. The new Britton and Brown illustrated flora,
Vol. 2. The New York Botanical Garden, New York.

3Specimens kept at a constant 40°F in March before spring growth had begun under-
went elongation of the stems and enlargement of the flower buds.

1962] Lewis — Polygala 225

GRAY, A. 1856. Manual of the botany of the northern United States,
2nd Ed. George Putnam and Co., New York.

RoBINSON, B. L., AND M. L. FERNALD. Gray's new manual of botany,
7th Ed. American Book Co., New York.

TWO NEW SPECIES OF POLYGALA FROM
NORTHERN MEXICO!

WALTER H. LEWIS

In an earlier paper on the cytology of Polygala (Lewis
& Davis, Rhodora 64: 102-113, 1962), reference was made to
five collections thought to represent four new species.
Using the keys and descriptions of Blake (N. Am. Fl. 25:
305-370, 1924), species numbered 3 (Oliver 136) and 4
(Lewis 5728, 5755) could be placed with P. alba Nutt. and
the var. suspecta Wats. The chromosome number of P. alba
in Texas was reported as n — 12 whereas those for number
3 from Oaxaca and number 4 from Coahuila and N uevo Leon
were recorded as n = ca. 36 and n = 52-54, respectively.
Although wide differences in chromosome number are not
unknown for a single species, these counts, when considered
in conjunction with the known heteromorphism of individ-
uals now included under P. alba (Dr. M. C. Johnston,
University of Texas, personal communication), strongly
suggest that more than one species is involved. Additional
cytotaxonomic study should be completed, however, par-
ticularly with the Mexican forms in the complex, before
the naming and describing of new taxa will be purposeful.
Consequently, only those plants represented by numbers
1 and 2 will be presented.

Polygala shinnersii W. H. Lewis, sp. nov.

Caules e radice tenui perenni plures 1.5-2.0 dm. alti. Folia inferiora
mediocriaque verticillata spatulato-obovata, suprema alterna linearia.
Racemi cylindrici acuminati 6-8 mm. diametro; flores candida purpureo-
cristati; sepala lanceo-elliptica 1.0-1.2 mm. longa; alae obovatae 2.9-3.3

"This study was supported by the National Science Foundation G-21818.
I am indebted to Dr. Lloyd H. Shinners, Southern Methodist University, for the
translations to the Latin.

226 Rhodora [Vol. 64

mm. longae. Capsulae ovales 2.6-3.2 mm. longae; semina pilosa 2.4-2.7
mm. longa, arillorum lobae duae linearo-ellipticae 1.6-1.8 mm. altae.

Stems several from slender perennial root, erect, angular, 1.5-2.0
dm. tall; leaves 4-6 in whorls to middle of stem or beyond, usually
shorter than the internodes, spatulate-obovate, 6-13 mm. long, 2-3 mm.
wide, the upper alternate, linear. cuspidate, 13-21 mm. long, 1.5-2.0
mm. wide, glabrous; peduncles 2.2-4.2 cm. long; racemes dense, cy-
lindrical, acuminate, 6-8 mm. thick; pedicels 0.2-0.3 mm. long; flowers
white with purple crests; sepals lance-elliptic, acuminate, 1.0-1.2 mm.
long; wings obovate, 2.9-3.3 mm. long, 1.7-2.0 mm. wide; keels 2.7-3.0
mm. long, the crest of 4 lobes on each side; capsules oval, 2.6-3.2 mm.
long, 2.2-2.6 mm. wide; seeds cylindrical, pilose, 2.4-2.7 mm. long, 1.2-
1.5 mm. wide; arils 1.6-1.8 mm. high, the 2 linear-elliptic lobes ap-
pressed; chromosome number 2n — ca. 84.

Holotype: MEXICO: Nuevo Leon: 11 miles E of junction of High-
ways 57 and 60, Lewis 5754 (US). Rare in open pine woods, 2 Sept.
1961. Known only from the type locality.

Although keying to the section Timutua, P. shinnersii is not closely
related to other species in the section. In habit it has a striking resem-
blance to that of Galium.

Polygala vergrandis W. H. Lewis, sp. nov.

Caules e radice annua plures puberuli 3-8 cm. longi. Folia alterna
linearia. Racemi cylindrici acuminati 2.5-4.0 mm. diametro; flores
candida viridi- et purpureo- venosi; sepala lanceo-elliptica 1.0-1.5 mm.
longa; alae obovatae 1.8-2.2 mm. longae. Capsulae oblongae 2.5-3.0
mm. longae indehiscentes loculo superiore alato inferiore exalato;
semina pilosa 2.0-2.4 mm. longa, arilli 0.6-0.9 mm. alti, lobis duabus
appressis (in seminibus e loculo inferiore deficientibus).

Stems several from a long slender annual root, 3-8 cm. long, erect
or ascending, slender, densely puberulous; leaves alternate, linear, 4-8
mm. long, 1 mm. wide, acuminate at apex, l-nerved below, margins
somewhat revolute; peduncles 3-6 mm. long; racemes cylindric, acu-
minate, dense, 2.5-4.0 mm. thick, the axis 10-15 mm. long; bracts lanceo-
late, deciduous, 1.0-1.3 mm. long; pedicels 0.3-0.5 mm. long; flowers
white, green- and purple- veined; sepals lance-elliptic, subacuminate,
1.0-1.5 mm. long; wings obovate, 1.8-2.2 mm. long, 0.6-0.9 mm. wide,
obtuse at apex; keels 1.6 mm. long, the crest of 2 lobes on each side;
capsules oblong, rounded at both ends, light green, 2.5-3.0 mm. long,
1.2-1.5 mm. wide, the upper cell larger, winged, the lower wingless,
indehiscent; seeds cylindrical, curved, pilose, 2.0-2.4 mm. long, 0.5 mm.
wide, seeds of lower cell somewhat smaller; arils 0.6-0.9 mm. high, 2
robes appressed, wanting in seed from lower cell; chromosome number

n = 24,

Holotype: MEXICO: Nuevo Leon: 9.9 miles E of junction of High-
ways 57 and 60, Lewis 5746 (US). Rare on rocky edge of hill, 2 Sept.
1961. Known only from the type locality.

1962] Rhoades — Alisma subcordatum 227

Characteristics of the section Monninopsis nearest P. scoparioides
Chod., but the resemblance to this species is not close. — STEPHEN F.
AUSTIN STATE COLLEGE, NACOGDOCHES, TEXAS.

THE AQUATIC FORM OF ALISMA SUBCORDATUM
Raf.

RICHARD W. RHOADES

Alisma subcordatum is a common species of marsh or
aquatic herb of temperate North America and usually grows
emersed with ovate or eliptical aerial leaves. In the fall of
1958 a botany class at the University of New Hampshire
collected a specimen growing in 11% feet of water which
had narrow-lanceolate floating leaves. Subsequently other
plants with floating leaves were collected in the vicinity of
Durham, N. H. While a deep water form of the European
Alisma plantago-aquatica is recognized (Arber 1920, 1925),
apparently most authors do not recognize a similar form
in the American species. Muenscher (1944), Fassett (1957 |»
Fernald (1950) and Hendricks (1957) do not mention a
deep water form, but Pierce (In Gleason 1952, vol. 1, p. 88)
states that plants growing in relatively deep water or where
the water level varies have longer, laxer, ovate-lanceolate to
linear leaves.

The following experiments were conducted in the green-
house to test the response of plants to varying depths of
water. Seeds were stratified at 5^ C. for 60 days according
to the recommendations of Crocker and Barton (1952), and
were then germinated in sand under water. On April 1 the
seedlings were transplanted to muck soil in battery
jars, about 20 Seedlings per jar. The water levels were
adjusted to 2 em (10 jars), 12 cm (4 jars) and 25 cm (2
jars). Nothing more was done to the plants except to
maintain the water level. By early July, four to six plants
in each jar with 2 or 12 cm of water had produced aerial
leaves and inflorescences In addition, the plants grown in
12 em of water produced floating leaves. Plants grown in
25 cm of water, however, produced only floating leaves.

228 Rhodora [Vol. 64

During the course of development from seedling to flower-
ing stage, Alisma subcordatum produces three types of
leaves. Seedlings possess a cluster of several gramineous,
primary leaves, 3.5 em in length which are usually sub-
merged. As the plants mature they next produce petioled,
narrow-lanceolate, juvenile leaves. If emersed, the later
juvenile leaves develop into ovate, aerial leaves, but if sub-
mersed, the later juvenile leaves retain their narrow form,
the petioles elongate, and the blades float on the surface
of the water.

The next phase of the experiment was designed to test
the effect of a raised water level on the development of plants
which previously had been grown in shallow water. Four
of the ten jars which had been maintained at the 2 cm
level were used. The water level was raised to 25 cm in
two of these. Inflorescences were removed from the plants
in one of these jars and also from the plants in one of the
jars with 2 cm of water. The status of each jar was then as
follows:

No. 1 25 cm water; inflorescences not removed

No. 2 25 cm water; inflorescences removed

No. 3 2 em water ; inflorescences not removed

No. 4 2 em water; inflorescences removed

A month later the plants in jars 1 and 2 had produced
only floating leaves. Those in jar 2 also produced reduced
inflorescences. The plants in jars 3 and 4 produced new
aerial leaves and new inflorescences.

The plants which had been grown continuously in 12 cm
of water produced aerial leaves, floating leaves and flowers
throughout the summer. Plants growing in 25 cm of water
from the beginning produced only floating leaves during the
entire period of observation.

Similar observations on the growth of Alisma plantago-
aquatica have been made by Arber (loc. cit.). This species
also commonly occurs in shallow water and produces aerial
leaves. The aerial leaves are preceded by submerged, “band-
shaped", primary leaves and usually by “swimming leaves”
as well. In moderately deep water, only submerged and

“swimming leaves" are produced. These “swimming leaves”

1962] Shinners — Mazus reptans 229

appear to be equivalent to the juvenile or floating leaves
observed in the present study.

The plants used in this experiment were the smaller-
flowered, more southern species Alisma subcordatum.
Although not recognized very widely by botanists, the
aquatic form of this species probably occurs rather com-
monly. This experiment has shown that plants of Alisma
have the ability to produce floating leaves if submersed at
any time during their period of development.

Specimens cited: in 1 to 1% feet of water, old reservoir, Durham,
Strafford County, N. H. R. W. Rhoades and A. R. Hodgdon. 16 Oct.
1958 (NH); in 2 feet of water, Hepler's Pond, Madbury, Strafford
County, N. H. R. W. Rhoades. 1 Nov. 1959 (N.H.).

— DEPARTMENT OF BOTANY, DUKE UNIVERSITY, DURHAM,
NORTH CAROLINA.

LITERATURE CITED

ARBER, A. 1920. Water plants. Cambridge, England.
1925. Monocotyledons. University Press, Cambridge.

CROCKER, W. and L. V. BARTON. 1952. Physiology of seeds. Chronica
Botanica Co. Waltham, Mass.

FassETT, N. C. 1957. A manual of aquatic plants (Revision ap-
pendix by E. C. Ogden). University of Wisconsin Press. Madison.

FERNALD, M. L. 1950. Gray's Manual] of Botany, 8th ed. American
Book Co. New York.

HENDRICES, A. J. 1957. A revision of the genus Alisma (Dill) L.
Amer. Midl. Nat. 58: 470-493.

MUENSCHER, W. C. 1944. Aquatic plants of the United States.
Ithaca, New York.

PIERCE, J. H. 1952. Alismaceae. In Illustrated flora of the northern
United States and adjacent Canada by H. A. Gleason. New York
Botanical Garden,

MAZUS REPTANS (SCROPHULARIACEAE) IN
MARYLAND. — Mazus reptans N. E. Brown, Bot. Mag. t.
8554, 1914, is a freely stoloniferous, mat-forming perennial
with flowers half again as large as those of the non-stoloni-
ferous, annual M. japonicus (Thunb.) Kuntze. The latter
Species is the only one hitherto reported as wild in the

230 Rhodora [Vol. 64

United States, first by Small in the second edition of his
Flora of the Southeastern United States (1913, p. 1365, as
M. rugosus), from southern Louisiana. It is not mentioned
in his Manual of the Southeastern Flora (1933). In Pennell’s
Monograph of the Scrophulariaceae of Eastern Temperate
North America (Phila. Acad. Monog. 1: 197, 1935), it is
cited from Pennsylvania, the District of Columbia, and Ore-
gon as well as Louisiana. More recently Fernald (Gray’s
Manual, 8th ed., p. 1275, 1950) reports it from “Pa. to Mo.
and La.; Pacific Slope,” and Gleason (The New Britton &
Brown Illustrated. Flora 3: 215, 1952) says “cultivated for
ornament and tending to escape in lawns,” without locality.
Gleason's description of it as “a ereeping perennial" and
“cultivated for ornament" cannot apply to M. japonicus.
Dr. Arthur Cronquist informs me that the New York Bo-
tanical Garden has a specimen of M. reptans “as a lawn
weed on the Cornell campus in Ithaca," New York, evidently
the basis for Gleason's remarks. It can now be reported that
M. reptans is genuinely naturalized in MARYLAND, Cal-
vert Co.: moist clay cliffs with Liquidambar Styraciflua &
Cryptotaenia canadensis, Scientists’ Cliffs, District No. 1,
Frank C. Seymour 16,697, 24 July 1946 (SMU). *Forming
a mat in dense shade." Plant past flowering. This species
is not treated in Hui-Lin Li’s “The genus Mazus (Scrophu-
lariaceae),” Brittonia 8: 29-38, 1954; his revision actually
treats only species known from China. Brown says that the
plant comes from the Himalayas. Presumably it is confined
to the eastern portion, since the account of the genus in
Pennell’s The Scrophulariaceae of the Western Himalayas
(Phila. Acad. Monog. 5: 33-35, 1943) does not include it.

Plants of M. reptans received from Rocknoll Nursery,
Morrow, Ohio, in late March, 1961 (in bloom on arrival),
survived in my yard in Dallas, Texas, to bloom again in the
spring of 1962, with more deeply colored and showier corol-
las than the previous year. They are growing in calcareous
clay in shade, and require constant watering during our hot,
dry summers. N. E. Brown spoke of it as flowering ‘‘almost
continuously from early spring to late autumn” under culti-
vation in England, but that is not the case under Dallas con-

1962] Adams — Guttiferae 231

ditions. It is surviving a second summer here (July, 1962),
but like last year shows no sign of blooming after the spring
season. The potential area which this species may occupy is
evidently very great. It should be watched for as an escape
elsewhere in the United States and southern Canada.

Mazus japonicus is an annual weed which, like several
others, both alien and native (e.g. Cardamine pennsylvanica,
Stachys floridana, Youngia japonica), is being spread all
through the South in shipments of ornamental shrubs,
especially azaleas and camellias. There are specimens in the
SMU Herbarium from Alabama (Baldwin Co.), Arkansas
(Clark Co.), Florida (Liberty Co.), Louisiana (Lafayette,
Ouachita, and St. Tammany parishes; Pennell knew it from
East Baton Rouge and Orleans parishes), and Texas (Dal-
las, Harrison, and Jefferson counties). The Dallas plants
seem to have been killed out by the abnormally severe winter
of 1961-1962, with repeated severe freezes (down to 17? F.),
but further observation will be needed to confirm this. —
LLovD H. SHINNERS, SOUTHERN METHODIST UNIVERSITY,
DALLAS 22, TEXAS.

STUDIES IN THE GUTTIFERAE. II.
TAXONOMIC AND DISTRIBUTIONAL
OBSERVATIONS ON NORTH AMERICAN TAXA:

PRESTON ADAMS?

This contribution is a miscellany of observations which
have accumulated during recent studies of the Guttiferae in
North America. Included are comments on the delimitation.
of the Guttiferae, a re-evaluation of the generic status of
Sanidophyllum Small, the relegation to synonymy of a few
specific epithets, some notes on geographic distribution of

‘Financial support of these studies was provided by the Fernald Fund for Field
Study in Systematic Botany at Harvard University, established by Mr. F., W. Hunne-
well of Wellesley, Massachussets, and a research grant (RG-6305) to Dr. R. K.
Godfrey of Florida State University from the Division of General Medical Studies,
Public Health Service.

?Present address: Department of Botany and Bacteriology, DePauw University,.
Greencastle, Indiana.

232 Rhodora [Vol. 64

several species, and a brief survey of some of the systematic
problems discernible in the family.

DELIMITATION OF THE GUTTIFERAE

The Englerian concept of the Guttiferae as comprising
five subfamilies is being followed in my studies. Some recent
authors, including Lawrence (1951) and Hutchinson
(1959), have split the group into two families, elevating
Engler's Hypericoideae to the familial level (Hypericaceae).
This restricted concept was used by me in earlier studies
(Adams, 1957, 1959). Recent investigation of the relevant
genera by Dr. Norman K. B. Robson at Kew strongly sug-
gests that a re-evaluation of the claim of the Hypericaceae
to family rank is necessary (Robson, personal communica-
tion). I agree with Dr. Robson that if Engler’s Hypericoi-
deae is elevated to the family level then it would appear
that the other four subfamilies must be made into families
also. This is clearly undesirable because of the close interre-
lationship which these groups appear to have with one
another. The Hypericoideae seem to be related to the Calo-
phylloideae and to the Clusioideae through Hypericum
(Lawrence, 1951; Robson, personal communication). The
other members of the H ypericoideae are T riadenum (eastern
North America and Japan), Vismia (tropical America),
Cratoxylon (Indomalaysia), Eliaea, Psorospermum, Endo-
desmia, and Haronga. (The last three genera are native to
tropical Africa and/or Madagascar.) One of Small’s generic
segregates, Sanidophyllum, of southern Florida, U. S. A., is
best included within Hypericum (see below).

RE-EVALUATION OF THE GENERIC STATUS OF SANIDOPHYLLUM

The genus Sanidophyllum was established by Small in
1924 to accommodate a single species, S. cumulicola Small.
Plants of this taxon are wiry herbaceous perennials which
grow in the Pinus clausa scrub in Highlands Co., Florida.
Commonly, several stems arise from an elongated taproot
near or just below ground level. The stems die back to
ground level each winter but the rootstocks survive for
several years. New growth each spring is due to the forma-

1962] Adams — Guttiferae 233

tion of young shoots from these rootstocks. On the juvenile
stems the leaves are densely crowded and tend to be more
or less closely appressed, the whole presenting a striking
bryoid appearance. As the stems elongate the internodes
become much longer at successively higher levels. Much of
the photosynthetic function is apparently carried on by the
stems since the leaves are greatly reduced in size compared
to other Hypericum species, excepting H. gentianoides. The
slender, sharply ascending stems with their tiny scale-like
leaves suggested to Small a species of Cathartolinum
(Linum).

Recent intensive studies of the floral anatomy and taxono-
my of Hypericum and several of the generic segregates
(Robson, 1956; Adams, 1962) have demonstrated that a
re-evaluation of the status of these groups is necessary.
While the anatomy of the flower in Sanidophyllum cumuli-
cola has not been investigated, comparative morphological
study of these plants as well as their relatives in Hypericum
Sect. Brathys strongly suggests that this species represents
merely an extreme evolutionary development within this
section of Hypericum. Morphological features common to
Sanidophyllum and certain species of Hypericum (especially
many of those in Sect. Brathys) include perennial habit,
much-reduced scale-like leaves which lack an articulation
at the base, clear glands in the leaves, sepals, and young
stems, 5 slightly unequal sepals which are non-articulated
at base, 5 slightly asymmetrical yellow petals whose with-
ered remains persist long after anthesis, long-persistent
stamens, 3 filiform-elongate styles which are seperate to the
base and somewhat spreading at anthesis, capitate stigmas,
3-carpelled gynoecium, parietal placentation, and dark
brown seeds with a finely reticulate testa.

In its morphology as well as its ecology Sanidophyllum
cumulicola seems to be most closely related to H. gentian-
oides (L.) BSP, a plant found over much of eastern United
States. The leaves of both are considerably reduced in
surface area, with those of the latter species being noticeably
smaller than the former. Plants of both species possess
slender wiry stems ; those of H. gentianoides are often much-

234 Rhodora [Vol. 64

branched, presenting a bushy appearance in contrast to the
relatively few-branched aspect of S. cumulicola. The wide-
spread H. gentianoides is an annual, even in peninsular
Florida, while the very local S. eumulicola is definitely peren-
nial. The inflorescence in both species is basically dichasial,
the pattern typical of Hypericum. The flowers of H. gentian-
oides are considerably smaller in size than those of S. cumu-
licola. AM these features suggest a fairly close relationship.

Both of these plants are adapted to dry habitats. Sanido-
phyllum cumulicola grows in the sand of the scrub in asso-
ciation with Pinus clausa, Paronychia pulvinata, Polygonella
basiramia, Prunus geniculata, Ceratiola ericoides, and other
species. It is most abundant in *blow-out" areas where the
moisture is more abundant and the competition from other
sand scrub species is less severe. Hypericum gentianoides
also grows in sandy soil, often becoming very abundant in
turkey oak-pine scrub (especially following clearing by
man), fallow fields, moist hollows in coastal dunes, rock
outcrops, and roadside embankments. Both of these species
are closely related to H. drummondii (Grev. & Hook.) T. &
G., a plant of the southern United States.

Sanidophyllum cumulicola is related, although much less
closely, to Hypericum fuertesii Urb. of Hispaniola. Plants of
this West Indian species have much-reduced leaves which
are strongly ascending and crowded. The inflorescence of
H. fuertesii is much less floriferous, however, and is often
reduced to a 3-flowered dichasium or even a single flower.
It is an inhabitant of high elevations (6,000-8,000 feet)
while S. cumulicola grows at only a few hundred feet. Other
relatives of this species comprise the H. pauciflorum complex
in Mexico and Central America (see below).

The reduction of Sanidophyllum to Hypericum makes
necessary the following combination :

Hypericum (Sect. Brathys) cumulicola (Small) P. Adams,
comb. nov., based on Sanidophyllum cumulicola Small, Bull.
Torrey Club 51:391. 1924.

The limited distributional area of this species is being
diminished at an alarming rate. The sand scrub lands of
central and southern Highlands County, Florida, are rapidly

1962] Adams — Guttiferae 235

succumbing to the bulldozer and the citrus grove. Studies
of the ecology and cytology need to be made before the
species becomes extinct.

TAXONOMIC NOTES

Hypericum aphyllum Lundell, Am. Midl. Nat. 29:477.
1943. TYPE: British Honduras, Toledo District, Monkey
River, near Jenkins Creek, Gentle 4175 (Holotype, MICH,
not seen; isotype, GH).

Lundell distinguished this species from Hypericum gen-
tianoides (L.) BSP, a plant of eastern United States, pri-
marily on seed characters. The seeds of H. aphyllum were
Said to be shorter and the testa smoother than in H. gen-
tianoides. Also, the number of Seeds per capsule is twice
that of H. gentianoides, according to Lundell. Two speci-
mens from Florida were assigned to the new species also.
My examination of many collections of H. gentianoides from
throughout its range indicates that seed characteristics are
highly variable. There is a tendency for the seeds to be
shorter and the testa smoother in specimens from the south-
ern portion of the range, with those of some collections
(e.g. those cited by Lundell and others) much resembling
those of H. aphyllum. Study of an isotype of that species
strongly suggests that, upon consideration of the entire
morphology of the plant, the claim of H. aphyllum to specific
status is highly questionable. Dr. A. J. Sharp wrote upon
the isotype sheet in 1950 that H. aphyllum is "doubtfully
distinct" from H. gentianoides. I think that a thorough
analysis of seed variation within the latter species ought to
be made before recognizing the British Honduran plant as
a separate taxon.

Hypericum pringlei Watson, Proc. Am. Acad. 25:143.
1890. TYPE: Mexico, Nuevo Leon, Sierra Madre, near Mon-
terey, Pringle 3012 (GH).

The type specimen of H. pringlei compares readily with
numerous specimens of H. perforatum L. from western
United States as well as from other regions. In my opinion
Watson's species is conspecific with H. perforatum.

Hypericum terrae-firmae Sprague & Riley, Kew Bull. page

236 Rhodora [Vol. 64

12. 1924. TYPE: British Honduras, Peck 321 (Holotype, K,
not seen; isotype, GH).

Sprague and Riley noted that their Hypericum terrae-
firmae is "the representative on the mainland of the Cuban
H. styphelioides A. Rich." However, they made no mention
of the features which they used to separate the mainland
plant. An isotype of H. terrae-firmae is hardly distinguish-
able, if at all, from many Cuban specimens of H. stypheli-
oides, a species originally described in 1845 (Ess. Fl. Cuba.
237). Although the Cuban plants exhibit considerable varia-
tion, especially in leaf size and shape, I am unable to find
any basis for taxonomic segregation of the Honduran speci-
mens. I first questioned the claim of H. terrae-firmae to
recognition in 1959 while doing casual study of the West
Indian-Central American specimens of the genus at the Gray
Herbarium. Recently, Dr. Louis O. Williams has independ-
ently arrived at the same conclusion, judging from his anno-
tation of the isotype at the Gray Herbarium. It seems clear,
therefore, that the plant of British Honduras can be readily
assigned to the earlier-named H. styphelioides.

NOTES ON GEOGRAPHIC DISTRIBUTION

Since the publication of distribution maps for the species
in the generic segregate Ascyrum (Adams, 1957) some note-
worthy range extensions or corrections have come to my
attention. These species were transferred to Hypericum by
Adams and Robson (1961).

HYPERICUM STRAGULUM Adams & Robson (Ascyrum mul-
ticaule Michx.). Mr. W. E. Buker, an associate of the Carne-
gie Museum in Pittsburg, Pennsylvania, has written me
that plants of this species have been collected in several
counties of southwestern Pennsylvania. These collections
(not seen by me) include the following: Bedford Co., south
of Silver Mills, Henry & Buker 5/19/51; Fayette Co.,
Ohiopyle, Shafer 9/1/01, and Bright 9/6/1915; Greene Co.,
1.5 miles east of Mt. Morris, Beer, Henry & Buker 8/8/51.
All of these specimens are in the Herbarium of the Carnegie
Museum, according to Mr. Buker.

At the time I published the revision of Ascyrum men-

1962] Adams — Guttiferae 201

tioned above, Hypericum stragulum was not known to grow
in South Carolina. In September of 1958 I collected it five
miles northwest of Walhalla in Oconee County (Adams 1 60,
FSU). Diligent search in adjacent counties will do doubt
reveal other stations.

HYPERICUM HYPERICOIDES (L.) Crantz (Ascyrum hyperi-
coides L.). The disjunction in range which was thought to
exist (Adams, 1957) in Mexico between southern Nuevo
Léon and eastern Hidalgo, a distance of some 300 km,
has been virtually eliminated by two recent collections. A
station in the state of San Luis Potosi, about 50 miles east
of the city of that name, was reported by Rzedowski &
Rzedowski (1957). Plants of this species have been collected
also in the state of Tamaulipas about 40 miles north-north-
west of Aldama (the Sierra de Tamaulipas range) by Dr.
R. L. Dressler (2408, GH).

SPECIES COMPLEXES REQUIRING INTENSIVE STUDY

During recent studies of North American Hypericum I
have become aware of several taxa which are in need of
critical evaluation. I believe that a brief discussion of these
groups will be of interest, especially to the workers current-
ly engaged in floristic studies. This is presented with the
hope that other botanists, especially graduate students in
search of thesis problems, will be enticed to study these
plants. In addition, the naming of new taxa, especially in
Mexico, is discouraged until thorough knowledge of the
variation present within these groups is forthcoming.

The HYPERICUM PAUCIFLORUM complex

The Mexican-Central American members of Hypericum
Sect. Brathys comprise a most variable group, judging from
study of the specimens at the Gray Herbarium, the U. S.
National Herbarium, and the Herbarium of the University
of Texas. At least seventeen species have been described but
several of these appear to represent only ecotypic variation.
The great altitudinal diversity (from sea level to at least
12,000 feet) present in Mexico and Central America is most
certainly correlated with much of the observable variability.

238 Rhodora [Vol. 64

The taxa which belong to the so-called H. pauciflorum com-
plex include the following: H. brevistylum Choisy, H. denti-
culatum HBK, H. eastwoodianum Johnston, H. fastigiatum
HBK, H. gnidioides Seem., H. hintonii Bullock, H. hondur-
asense Keller, H. longibracteatum Keller, H. paniculatum
HBK, H. pauciflorum HBK, H. paucifolium S. Wats., H.
pinetorum Standley, H. pratense Cham. & Schl., H. schaff-
neri S. Wats., H. silenoides Juss., H. submontanum Rose,
H. uliginosum HBK, and H. woodsonii Standley. These
plants are related to the United States species H. cumulicola
(Small) P. Adams, H. denticulatum Walt., H. drummondii
(Grev. & Hook.) T. & G., and H. gentianoides (L.) BSP.
Finally, the very distinctive H. setosum L., a plant of the
United States Atlantic and Gulf Coastal Plain, is clearly a
member of this complex also. Its pilose stems and leaves
and ciliate sepals, not present in any other Hypericum in
North America, are probably an independent evolutionary
development.

Research in this complex should include field observations
on growth form, perennation, altitudinal effects upon the
plant body, and chromosome counts, as well as the usual
examination of herbarium specimens. Pending the comple-
tion of at least a synoptic treatment of these taxa I would
strongly advise against the publication of putative hitherto
undescribed taxa. To continue to add to an obviously over-
burdened synonymy seems to be most unwise.

The HYPERICUM PUNCTATUM complex

Another group within Hypericum which is obviously in
need of intensive study is the H. punctatum complex of Sect.
Hypericum, as currently circumscribed (Keller, 1925). At
least six species are recognized, including H. formosum
HBK, H. graveolens Buckley, H. mitchellianum Rydberg, H.
pseudomaculatum Bush, H. punctatum Lam., and H. scouleri
Hook. These taxa are morphologically very similar and,
therefore, presumably closely related. One of these, H. for-
mosum, is a plant of high elevations from British Colombia
southward through western United States and Mexico into
Guatemala. The United States plants of this species are

1962] Adams — Guttiferae 239

classified either as a separate taxon, H. scouleri Hook., or as
a variety. Many specimens of H. formosum, especially from
Mexico and Guatemala, are practically identical with, if not
indistinguishable from, those of H. pseudomaculatum, a
plant of the Ozarkian region of Arkansas. Specimens of
these two species compare readily with those of H. graveo-
lens, a plant endemic in the Blue Ridge mountains of western
North Carolina at high elevations. The Ozarkian H. pseu-
domaculatum has been considered to be a variety of H.
punctatum, a plant widespread over much of eastern North
America (Fernald, 1950). Hypericum mitchellianum, also
endemic in the Blue Ridge mountains, is doubtfully distinct
from H. punctatum. In fact, both possess an unusual ring
formation of the chromosomes during meiosis (see below).
Both H. graveolens and H. mitchellianum grow in close asso-
ciation at Mt. Mitchell, North Carolina.

At least two members of the Hypericum punctatum com-
plex are highly interesting cytologically. During meiosis the
sixteen diploid chromosomes become attached end-to-end,
forming a ring. Such unusual behavior is often indicative of
structural hybridity (Stebbins, 1950). This phenomenon
was first reported by Hoar (1931) in Massachusetts plants.
During my studies I have observed ring formation in plants
of this species from North Carolina and Tennessee. I have
also observed this abnormal cytological condition in several
collections of H. mitchellianum. In the one collection of H.
pseudomaculatum which I have been able to study (courtesy
of Dr. Paul L. Redfearn) no evidence of meiotic irregularity
was noted. This is taken to be further evidence supporting
the specific distinctness of these plants from H. punctatum.
No chromosome studies are available for H. formosum.

For the cytologically-inclined systematist the Hypericum
punctatum complex presents a most intriguing problem.
Several questions arise. For instance: Is the ring formation
found in plants of H. punctatum throughout its geographic
range? Is this condition actually absent in H. pseudomacula-
tum? Would intensive study reveal its presence in H. grave-
olens and H. formosum? What is the relationship between
“true” H. formosum and the western United States H.

240 Rhodora [Vol. 64

scouleri? Would careful study show that perhaps the Ozark-
ian H. pseudomaculatum is really conspecific with the Mexi-
can H. formosum? What is the relationship, if any, between
H. punctatum (a ring-forming plant) and these three taxa?
Is the endemic H. mitchellianum worthy of taxonomic recog-
nition and, if so, what is its relationship to H. punctatum?
Could it be merely a high altitude ecotype of H. punctatum?
These are merely a few of the more obvious questions which
come to mind; doubtless many other problems exist within
the H. punctatum complex.

The HYPERICUM CANADENSE complex

This highly polymorphie group within Sect. Brathys of
eastern North America has long been a puzzle to botanists.
The species limits are not at all clear. At least five are
generally recognized, including H. boreale (Britt.) Bickn.,
H. canadense L., H. gymnanthum Engelm. and Gray, H.
majus (Gray) Britt., and H. mutilum L. The taxonomic
difficulties may well be due to extensive hybridization, as
suggested by Fernald (1950). The principal taxonomic
characters (at least those currently in use) are in the vege-
tative body, especially the leaves. Since these plants fre-
quently grow on soils of varying wetness it is not at all
unlikely that the habitat may affect the commonly-used
characteristics. I have observed one species, H. mutilum, at
many stations in northern Florida and southern Georgia
and have noticed great variation in size and habit of the
plants and in the size of the leaves. The presence of vegeta-
tive reproduction is an additional complicating factor, ren-
dering the collection of poulation samples most difficult.

Numerous problems are suggested by even a cursory
examination of the herbarium material and the literature.
For example: Does hybridization actually occur? If so, is
H. dissimulatum Bickn., as suggested by Fernald (1950),
really “an unusually constant and recurrent hybrid" of H.
boreale or H. mutilum and H. canadense? What is the status
of Fernald's three varieties of H. mutilum? I can detect
little, if any, geographic localization of any of them, except-
ing perhaps var. latisepalum which seems to be restricted

1962] Adams — Guttiferae 241

to the Atlantic and Gulf Coastal Plain. What is the status of
Fernald’s varieties magninsulare and galiiforme of H. cana-
dense? Both appear to me to be merely responses to unusual
habitat situations and perhaps not worthy of taxonomic
recognition.

The HYPERICUM DENTICULATUM complex

An interesting problem involving variation within Hyper-
icum denticulatum Walt. is discernible. According to Fer-
nald and Schubert (1948) this species of the eastern United
States comprises three varieties, each with some degree of
geographic localization. One of these, var. denticulatum,
grows on the Atlantic Coastal Plain from New Jersey south-
ward to at least South Carolina. Fernald and Schubert's
map also shows a station on the Cumberland Plateau in
central Tennessee, a disjunction of at least three hundred
miles from the nearest South Carolina locality. A second
variety, recognitum, is principally a plant of the upland and
mountainous areas from southeastern Virginia, through the
inner Piedmont of North Carolina to northeastern Alabama
and northward into southern Indiana and West Virginia.
Fernald and Schubert's map shows an occasional station
near the Fall Line in South Carolina, Georgia, and Alabama.
The third variety, acutifolium, is scattered about the south-
eastern United States, with occasional stations on the outer
Piedmont of Virginia and North Carolina, the Coastal Plain
of South Carolina and adjacent southeastern North Carolina,
northern Florida, the outer Piedmont of Georgia and adja-
cent Alabama, and the Cumberland Plateau of Tennessee.

Leaf shape was the principal feature used by Fernald and
Schubert (1948) to distinguish these three varieties. In one,
var. acutifolium, all or at least the middle and upper leaves
were described as “narrowly linear or linear-lanceolate"
while in a second, var. denticulatum, the leaves were “nar-
rowly to broadly ovate, oval or oboval." These two extremes
in leaf shape can be recognized fairly easily. The var. recog-
nitum is very obscure, however, the leaves appearing to be
intermediate between the two extremes. Fernald and Schu-
bert acknowledged that “the three varieties may merge" and

242 Rhodora [Vol. 64

that “nondescript individuals can be found.” In fact, two
of these taxa, var. acutifolium and var. recognitum, may
even grow closely associated within the same habitat, judg-
ing from Fernald’s annotations on herbarium material.
Obviously, therefore, H. denticulatum is in need of careful
and intensive study, especially in the field. It is not unlikely
that some differentiation may have occurred, rendering the
plants of the Coastal Plain slightly distinct from those of
the Piedmont and the mountains. However, I think that the
variation picture within this species is not so easily under-
stood as Fernald and Schubert’s analysis would suggest.

LITERATURE CITED

ADAMS, PRESTON. 1957. A revision of the genus Ascyrum (Hyperi-
caceae). Rhodora 59:74-95.

1959. The taxonomy of Hypericum section Myri-

andra (Hypericaceae). Unpublished Ph.D. Thesis, Harvard

University.

1962. Studies in the Guttiferae. I. A. synopsis of
Hypericum section Myriandra. Contributions from the Gray Her-
barium of Harvard University 189:1-51.

and N. K. B. RoBsON. 1961. A re-evaluation of
the generic status of Ascyrum and Crookea (Guttiferae). Rhodora
63:10-16.

FERNALD, M. L. 1950. Gray’s Manual of Botany. Ed. 8. American
Book Co., New York.

and B. G. SCHUBERT. 1948. Studies of American
types in British herbaria. Part IV. Rhodora 50:205-208.

Hoar, C. S. 1931. Meiosis in Hypericum punctatum. Bot. Gaz. 92:
396-406.

HUTCHINSON, J. 1959. The Families of Flowering Plants. Vol. 1.
Ed. 2. Oxford University Press. London.

LAWRENCE, G. H. M. 1951. Taxonomy of Vascular Plants, The
Macmillan Co., New York.

RZEDOWSKI, J. and G. C. RZEDOWSKI. 1957. Notas Sobre la flora y la
vegetación del estado de San Luis Potosí. VI. Acta Cientifica
Potosina 1:199-218.

STEBBINS, G. L. 1950. Variation and Evolution in Plants. Columbia
University Press. New York.

1962] Davis and Burns — Analysis of Hops 243

THE USE OF VAPOR FRACTOMETRY IN THE
ANALYSIS OF SOME NEW ENGLAND HOPS'

EDWARD L. DAVIS AND RICHARD L. BURNS

In recent years a number of articles have been published
on the use of chemical characteristics in taxonomic studies
(e.g. Turner and Alston, 1959; Alston and Irwin, 1961).
These examples have shown that chemical analysis can be
extremely useful in areas where extensive morphological or
taxonomic information has been accumulated. Not only may
the chemical analysis “solve” certain taxonomic problems,
but it may also serve to substantiate conclusions formed
originally on morphological features alone and in this way
give greater credence to studies which have been based
heavily upon qualitative rather than quantitative observa-
tions.

We are reporting here the use of a very sensitive method
of chemical analysis, vapor fractometry, as it is applied
to the problem of subspecific variations in the hops plant,
and its correlation with morphological studies which have
been done previously. The genus Humulus is a small one
consisting of two distinct species, one a perennial and the
other an annual. Several attempts have been made to dis-
tinguish two or more separate species of the perennial hops.
The difficulty of this separation, from herbarium material,
has been considered in detail (Davis, 1957). Within the
perennial species, Humulus lupulus L., it is clear that there
are variants, but these are not distinct enough to justify
specific rank. After a study of cultivated hops from through-
out the world, and wild hops of North America, the perennial
hops were divided into “complexes” which represent approx-
imately varietal differences. The differences between these
complexes were based upon cytological and morphological
features. Such features as the angularity and scar occur-
ring on the cone axis; pubescence, dentation and lobing of
the leaf; venation of the cone bract; and pairing of the sex
chromosomes have been used. It is stil premature to

1This research was supported by a grant from the National Science Foundation,
NSF-G18013.

244 Rhodora [Vol. 64

attempt to delimit these complexes in a definite way. Addi-
tional studies since 1957 have shown the complexity of the
situation. Neve’s work on hops chromosomes (1956, and
personal communications) make it necessary to withhold
final judgement as to chromosome types which might fit
American, Continental and English complexes of hops.
Work is being done at present upon the cytological features
of the hops in the American Rockies and this should lead
to more positive conclusions.

A comparison of morphological features and alpha acid
content among cultivated varieties suggests that there is a
difference in the proportion of humulone and cohumulone
between the different morphological groups. Rigby (1956)
and Howard and Tatchell (1955) classified hops according
to the cohumulone content of the humulone complex and
made 4 groups with approximately 20%, 30%, 40% and over
40% cohumulone. Examination of morphological features
of many of these varieties show those with 20-25% cohumu-
lone to correspond very closely to the Continental complex ;
those with 26-35% cohumulone to the English complex, and
those with cohumulone of 36% and above to the American
type. Considerable caution must be used in drawing these
comparisons. It is necessary to establish that these chemical
differences are not due to environmental conditions. Fortu-
nately, extensive evidence indicates that although the abso-
lute amounts of these substances are changed by the
environment, the relative proportions are not. Commercial
samples of the same variety, grown in very different habitats
and in different seasons have fallen into the same chemical
groups (Howard and Tatchell, 1956). Additional plant by
plant studies over a number of years, already in progress,
will be desirable to confirm this, but present evidence favors
the stability of the chemical characteristics. A more funda-
mental problem is related to drawing conclusions about a
correlation of structural features and chemical tests based
upon the kind of samples which have been examined so far.
Neve and Weston (1958) have very clearly stated this
problem in relation to the resins and oils in hops. The
cultivated varieties of hops are not a random sample. It is

1962] Davis and Burns — Analysis of Hops 245

not surprising that as a result of selection there are certain
common characteristics within each of the three major
groups of cultivated varieties, both in chemical and morpho-
logical features. These cultivated plants represent a very
small fraction of the variation within the species. Also,
within each group some common genetic background must
occur, although their ancestry is very incompletely known.
Working in the experimental fields at the Hops Research
Station at Corvallis, Oregon, with the morphological com-
plexes referred to earlier, the senior author found that
among the many hundred of crosses there were many plants
which would not fit easily into any of the three complexes
suggested. The difficulties were much greater than had been
observed among collections of wild hops throughout North
America. Although this topic will be developed more fully
in a later paper, it is clear that under natural conditions
many of the potential hops types do not survive, and that
there are strong forces limiting the variants within wild
plants. For example, it was shown previously (Davis, 1957)
that the wild hops of the midwestern United States are
remarkably uniform, but that hops in the Rocky Mountains
and New England are extremely variable. Only the Ameri-
can type has so far been found in the midwest, but apparent
variants of Continental and English hops occur in the West-
ern and Eastern parts of North America. Valuable as the
morphological analysis has shown itself to be, we have had
a considerable desire to find some other method of approach
to determine if the heterogeneity in wild hops is real and
related to that occurring among cultivated plants and not
due to a weakness in the morphological classification itself.
Cytological studies are being conducted, but because of the
very limited information on chromosome types in wild
American hops it is impossible to interpret the results at
this time. Another alternative is offered by the humulone
complex of substances which have been shown to be differ-
ent in American and European cultivated hops. We have
indicated that this method of analysis, which is firmly
established in respect to the chemical procedure, still
requires study before its biological and genetical value can

246 Rhodora [Vol. 64

be established beyond any possible doubt, but that there is
much information which justifies its use at present.

The wild hops of New England are particularly fascinat-
ing because in addition to their botanical variation they
present an intriguing problem due to historical circum-
stances. It is not certain that hops are native to New
England. Cultivation in the New England Colonies occurred
so early (probably with English hops) that there is a ques-
tion as to whether wild hops are exclusively escapes from
cultivation or a mixture of wild and escaped plants. Asa
Gray (1886) refers to wild hops scattered on banks of
Streams from Canada west to New Mexico, but he is not
clear about New England where he notes that all cultivated
plants are undoubtedly of European origin. Nuttall (1847 )
is certain that hops are native to the Rocky Mountains, but
did not commit himself on New England. During Revolu-
tionary days, New England and particularly the Connecticut
River Valley, was the center of hops cultivation in the
United States. There can be no doubt that English hops
were grown here and had ample opportunity fifty to two
hundred years ago to become established. Our previous
analysis of New England hops, from herbarium material,
has shown the leaf pubescence pattern typical of English
hops to be by far the most common, but in at least some
characteristics all three complexes are represented. As we
gradually learn more about the hops now wild in New
England we will know whether they are related to English
hops only, or to Continental and American hops as well.
Chemical analysis should contribute significantly to this
picture.

EXPERIMENTAL PROCEDURE

The determination of the relative proportions of humu-
lone, cohumulone and adhumulone in the alpha acids was
first done by the technique of Verzele and Govaert (1955)
by separating the humulone complex on a silica gel column,
followed by the determination of the relative amount of
each compound with a Beckman Du spectrophotometer,
measuring absorbancy at 276 my. This procedure did permit
a separation of the individual components, but it proved

1962] Davis and Burns — Analysis of Hops 247

very difficult to replicate the results quantitatively. A much
more exact method, developed by Rigby, Sihto, and Bars
(1960) involves extracting from the hops cones the alpha
acids, precipitating them as a lead salt, pyrolyzing the salt
to break off isobutyric, 2-methylbutyric, and isovaleric acid
from the molecules of cohumulone, adhumulone and humu-
lone respectively. Rigby has shown that this technique gives
results very similar to that from a separation of the humu-
lone complex by the much more time consuming counter-
current distribution technique. The fatty acids were then
esterified with isopropyl alcohol, and the isopropyl esters
separated on a chromatographic column (type R, Perkin-
Elmer) at 100°C, and a helium gas flow rate of 120 ml/min,
using a Perkin Elmer vapor fractometer, Model 154. The
details of the extraction and preparation of the fatty acid
esters are to be found in Methods of Analysis of Assoc. of
Agric. Chemists (1945) and Rigby, Sihto, and Bars, (1960).
The relative quantity of each component was determined by
calculation of the areas under the curves for each acid, as
recorded by the fractometer recorder, and the results report-
ed as a percentage of the total alpha acid.

RESULTS

Several hops plants were collected from Massachusetts
and Maine and their morphological and chemical character-
istics determined. Other plants from Vermont, Maine and
Massachusetts produced such a poor yield of cones that it
was impossible to analyze the alpha acids and the morpho-
logical analysis of these plants has not been included in this
study. For comparative purposes commercial samples of
Idaho Seedless hops (a Late Clusters type) and cones from
a plant of Late Clusters grown in Amherst are included.
Late Clusters belongs to the American complex of hops.
Table I contains the results of the chemical analysis. Three
tests were run on each hop. Table 2 shows the results of
the examination of morphological features. In each table,
the sample is assigned to a “type” in the last column. The
details of the morphological analysis is found elsewhere
(Davis, 1957).

248 Rhodora [Vol. 64

DISCUSSION

Comparison of the tables for morphological and chemical
analysis shows that when the cohumulone scale is established
as indicated, there is an exact correspondence between
morphological and chemical types. The scale was estab-
lished prior to this study, after comparing the work on
cultivated varieties referred to above. It does not imply,
for example, that all plants cultivated or wild, bearing the
designation American, are of origin within North America.
Such a conclusion could only be drawn at the completion of
an extensive study. Rather, the terms American, Contin-
ental, and English were chosen because, of the wild and
cultivated plants studied, these designations were the most
descriptive of their center of distribution. It may be neces-
sary to add additional designations for other parts of the
world, or to establish several complexes for the large areas
like North America. The more important goal is to establish
a chemical analysis that will support morphological investi-
gations. These preliminary results are very encouraging in
this respect.

The hops from Biddeford, Maine were found growing in
sand near the coast and could not have been in cultivation
at any time. Those from Windsor, Massachusetts lined a
fence near fields long deserted. They certainly had not been
cultivated where they were growing, but might have escaped
from cultivation in this area in recent times. The collection
from Dalton, on the other hand, was of plants which had
been used for brewing within the last 25 years.

Chemical analysis shows the Dalton hops to be strikingly
similar to the cultivated Late Clusters, such as is grown in
Idaho. The sources of its roots are unknown, but it is not
unreasonable to suggest that they were obtained commercial-
ly from a supplier of cultivated hops, and not brought into
cultivation from the wild state. The hops from Maine and
Windsor, which may have been seedlings from cultivated
plants of several generations ago are similar to English
cultivated types. In morphological features they resemble
the oldest herbarium material of New England hops.

With a continued examination of hops about which

1962] Davis and Burns — Analysis of Hops 249

detailed information on their history is available, we should
be able to answer the question as to whether or not American
type hops in New England are only of recent origin with
as much certainty as is possible in this kind of taxonomic
problem. Perhaps most important, these preliminary
chemical tests confirm our previous observations of a
morphological variation in New England wild hops which
corresponds to the variation between cultivated hops
throughout the world, and gives additional support to the
use of morphological evaluation in tracing the evolution of
hops.

__ DEPT. OF BOTANY, UNIV. OF MASSACHUSETTS, AMHERST

TABLE I
CHEMICAL ANALYSIS
Percentage of

cohumulone adhumulone humulone TYPE*
Idaho Seedless 49.2 8.7 42.1
(1961 crop) 48.6 8.4 43.0
46.8 10.4 42.8

average 48.2 9.2 42.6 American

Late Clusters

(Amherst) 36.1 14.9 49.0
37.8 15.7 46.5
37.6 13.2 49.2

average 37.2 14.6 48.2 American
Dalton, Mass. 46.0 10.5 43.5
47.9 8.6 43.5
45.6 12:2 42.2

average 46.5 10.4 43.1 American
Windsor, Mass. 32.2 13.0 54.8
32.5 12.8 54.7
35.7 13.2 51.1

average 33.5 13.0 53.5 English
Biddeford, Maine 29.6 13.1 57.3
24.9 12.9 62.2
24.8 12.8 62.4

average 26.4 12.9 60.6 English

*The designation is based upon a scale for cohumulone of: Continental, 20-25%;
English, 26-35%; American, 36% and over.

[Vol. 64

Rhodora

250

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1962] Turner, Powell & King — Chromosome numbers 251

LITERATURE CITED

ASSOCIATION OF OFFICIAL AGRICULTURAL CHEMISTS. 1945. Official
and Tentative Methods of Analysis. Washington 4, D. C.

ALSTON, R. E. AND H. S. IRwIN. 1961. The comparative extent of
variation of free amino acids and certain “secondary” substances
among Cassia species. Am. Jour. Bot. 48: 35-39.

Davis, E. L. 1957. Morphological complexes in hops (Humulus lupu-
lus L.) with special reference to the American race. Ann. Missouri
Bot. Gard. 44: 271-294.

Gray, ASA. 1886. American Druggist 15: 111.

Howarp, G. A. AND A. R. TATCHELL. 1956. Evaluation of hops. New
approach to the detailed analysis of hop resins. J. Inst. Brew.
62: 20-27.

Neve, R. A. 1958. Sex chromosomes in the hop Humulus lupulus.
Nature 181: 1084-1085.

NEVE, R. A. AND E. W. WESTON. 1958. Resin and oil relationships
in hops. J. Inst. Brew. 64: 247-248.

NUTTALL, T. 1847. Descriptions of plants collected by William
Gambel in the Rocky Mts. and upper California. Jour. Acad. Phil.
N.S. 1:182.

RigBy, F. L. 1956. Recent advances in the chemistry of hop con-
stituents and their significance in brewing technology. Technical
Proceeding Master Brew. Assoc. of Am. 69: 9-16.

RiGBy, F. L., E. SIHTO AND A. Bars. 1960. A rapid method for the
detailed analysis of the alpha acid fraction of hops by gas
chromatography. J. Inst. Brew. 66: 242-249.

TURNER, B. L. AND R. E. ALSTON. 1959. Segregation and recombina-
tion of chemical constituents in a hybrid swarm of Baptisia
laevicaulis X B. viridis and their taxonomic implications. Am.
Jour. Bot. 46: 678-686.

VERZELE, M. AND F. GOVAERT. 1955. Analysis of the *Humulone
Complex" by a chromatographic partition procedure. Wallerstein
Lab. Commun. 18: 181-190.

CHROMOSOME NUMBERS IN THE COMPOSITAE.
VI. ADDITIONAL MEXICAN AND
GUATEMALAN SPECIES.

B. L. TURNER, M. PoWELL AND R. M. KING

The present contribution is essentially a continuation of
several papers, the latest of which (Turner et al., 1961a)

252 Rhodora [Vol. 64

dealt with chromosome counts from species of southern
Mexico and Guatemala.

Chromosome counts were made from pollen-mother-cell
squashes as outlined by Turner and Johnston (1961).
Voucher specimens (table 1) are deposited at the University
of Texas Herbarium; these were collected during the year
1961. The tribal and subtribal arrangements listed in table
1 follow those of Hoffmann (1894).

While most of the identifications are our own we would
like to acknowledge the kind assistance of Dr. Kittie Parker
and Dr. Arthur Cronquist for the identification of certain
difficult taxa.

EUPATORIEAE — Eupatorium (x = 10, 17). Chromosome
counts for the 7 species listed in table 1 are consistent with
the basic numbers obtained for the approximately 40 other
species examined (Turner et al., 1961b). E. pycnocephalum
(n = 10, 20), a widespread, highly variable species, appar-
ently consists of diploid and tetraploid races, some of these
being in close proximity to each other (cf. King 4242 and
4243A, table 1).

Piqueria (x — 12, 11). The 2 species listed in table 1
are diploid with » — 12; Turner and Johnston (1961) have
reported one other species, P. laxiflora, to be diploid with
n = 11.

Mikania cf. gonzalezii (n — 17) is the second species of
the genus to be counted. Other workers have reported the
widespread species, M. scandens, to be diploid with 2n = 38
(Darlington and Wylie, 1956) or 2n — 36 (Mangenot and
Mangenot, 1958).

Chromosome counts for the species of Ageratum (n — 10,
20), Brickellia (n = 9) and Oxylobus (n = 16) are consis-
tent with the basic numbers obtained for these genera by
other workers (Turner et al., 1961a ; Gaiser, 1953; Beaman
et al., 1962). The chromosome count for Trichocoronis
wrightii (n — 15) is a first report for the genus.

ASTEREAE — Aster bimater (n — 5) is related to A. lima
Lindl.; it apparently belongs to the Section Aster (subsec-
tion Homophylli) as treated by Gray (1886). Chromosome

This study was supported by National Scienee Foundation Grant 9025,

1962] Turner, Powell & King — Chromosome numbers 253

numbers on a base of « = 5 occur in the sections Oxytripo-
lum (annuals and biennials) and Aster (perennials) of the
genus Aster. In the former section only counts of n = 5
or 10 are known while the latter has species with both n = 5
and n= 9. A. exilis var. australis (n — 5) belongs to the sec-
tion Oxytripolum; several additional counts of the species
(all n = 5) were reported by Turner et al. (1961b; cited as
A. subulatus Michx.).

Erigeron (x = 9) — The chromosome numbers of the
several species listed in table 1 are consistent with the basic
number obtained for numerous other taxa (Montgomery
and Yang, 1960). E. scaposus with n = 9II, 1811, and
27 (91I and 91) is a widespread, variable species, apparently
consisting of diploid, tetraploid and triploid races.

Psilactis (x — 4, 9, 8?). Chromosome counts for the 2
species listed in table 1 are particularly interesting in that
they parallel the multibasic condition found in Aster (Tur-
ner et al., 1961b). Meiotic figures of P. brevilingulata with
n = 9 appeared to be unequivocal for at least one collection
(King 2939) but for the other collections listed the counts
might have been 8II and 2 fragments as indicated for the
Powell and Edmondson collection (the 2 “fragments” taking
less stain than the well defined bivalents, but occurring
together at meiosis as if they were pairing). As in Chrysop-
sis (x = b, 4 and 9) and in Aster (x — 5 and 8, 9) the
species of Psilactis with the lower basic number has rela-
tively large chromosomes. Psilactis is composed of only 4
or 5 species, these restricted to the southwestern United
States and Mexico. It is a closely knit group of doubtful
affinity ; Gray in his original description of the genus placed
it “between Dieteria [= Machaeranthera] and Aster,
except from the want of pappus in the ray. . . ." Super-
ficially, at least in habit, the species centering about P.
brevilingulata and P. asteroides appear closest to the genus
Aster, section Aster or Oxytripolium, both of which, as
indicated above, have species with n = 5. However, an inclu-
sive morphological study of Psilactis (Turner & Horne, un-
published) shows the relationship of Psilactis to be closer to
Machaeranthera, particularly through P. coulteri which is a

254 Rhodora [Vol. 64

good match for Machaeranthera parviflora Gray, as indi-
cated by Gray in his original description of this latter
species.

Chromosome numbers for the species of Haplopappus,
Heterotheca, Machaeranthera, Solidago and Xanthocepha-
lum (table 1) are consistent with the basic numbers pre-
viously reported for these genera (Turner, 1961b; Darling-
ton and Wylie, 1956; Solbrig, 1961).

INULEAE — Chromosome counts for the species of Gna-
phalium (n = 7, 14) and Pluchea (n = 10) are consistent
with the basic numbers reported for these genera by previ-
ous workers.

HELIANTHEAE — Melampodinae — Polymnia (n= 16, 17).
Mr. J. E. Wells of Ohio State University is currently study-
ing the genus and has found yet other species with n = 15
(personal communication). Chromosome counts for Ber-
landiera (n = 15), Parthenium (n = 18) and Dugesia (n =
18) are consistent with counts reported for these genera by
previous workers (Turner and Johnston, 1956; Turner et al.,
1961a).

ZINNINAE — Heliopsis buphthalmoides (n — 28). A col-
lection of this species from Oaxaca, Mexico was reported to
be diploid with n = 14 by Turner ef al. (1961b). Fisher
(1957) reported counts for 6 other taxa of the genus; all
were diploid with n — 14. Philactis is a genus with perhaps
2 or 3 species. P. nelsonii (n — 28) is probably a tetraploid
on a base of x — 14 since the genus is closely related to
Heliopsis.

Zinnia maritima (n = 13). Previous published counts
for species of Zinnia have been n — 10, 11, 12, 19, and 21
(Torres, 1961). The meiotic figures for Z. maritima. were
particularly clear (Fig. 11). A collection of Z. angustifolia
from San Luis Potosi was reported as diploid with » — 11
by Turner et al. (1961b) ; the Nayarit collection of this
species (table 1) was found to be diploid with n = 12, agree-
ing with counts obtained for the species by Torres (personal
communication). Zinnia is obviously a multibasic taxon
with x = 10, 11, 12, 13, 19 and 21. (The latter two counts
are probably derived through amphiploidy or else through

1962] Turner, Powell & King — Chromosome numbers 255

aneuploid loss and gain respectively from a tetraploid on
a base of z — 10.)

Sanvitalia (x —8). A number of chromosome counts
have been reported for this genus; all have been diploid or
tetraploid on a base of x = 8 (Turner et al., 1961a ; 1961b).

The subtribe Zinninae, as treated by Hoffmann (1894),
includes only 6 genera: Philactis (including Grypocarpha;
Blake, 1930), Heliopsis, Aganippea, Tragoceros, Sanvitalia
and Zinnia. Five of the 6 genera have been counted and
basic chromosome numbers of x = 8, 10, 11, 12, 13 and 14
have been established (excluding the chromosome counts of
a presumed polyploid origin). At least two obvious hypothe-
ses may be tentatively proposed to account for this sequence
of numbers: (1) That the genera have been derived through
progressive aneuploid gain from an ancestral base of # = 8
or else the reverse has occurred from an ancestral base of
z — 14. In either case one would have to assume that, con-
trary to the situation in many other composite genera, the
hypothetical taxa on a base of x = 9 were lost from the
sequence. Another hypothesis may be proposed which avoids
this difficulty, this being that x — 4 is the ancestral basic
number of the subtribe, the numbers n = 8, n = 12 being
tetraploid and hexaploid respectively, chromosome numbers
of n = 11, 10 and 13, 14 being derived through both aneu-
ploid gain and loss.?

VERBESININAE — Aldama dentata Less. (Not Aldama
dentata Llave & Lex.) This species has been treated as
belonging to the genus Sclerocarpus by several workers
where it is called S. schiedianus because of the earlier name,
(S. dentata (Llave & Lex.) B. & H. ex Hemsl.). Morpho-
logically Aldama dentata Less. is quite different from
Sclerocarpus proper, lacking the conspicuous clawed rays of
the latter genus and possessing in addition rather distinct
achenes and involucral bracts. As indicated in table 1,

3Any number of alternate numerical hypctheses could be proposed, one of the most
provocative being that suggested by Sató (1960). He postulates that the ancestral
chromosome number ("protokaryotype") for the plant kingdom might be xz = 2,
presumably the major phyletic lines becoming established on this base. Obviously
with such a low basic number one could assume, on numerical grounds at least, that
ail higher numbers are polyploids from such a base, the odd numbers simply being
aneuploid derivatives,

256 Rhodora [Vol. 64

Aldama dentata. Less. has a chromosome number of m = 17,
while true Sclerocarpus has chromosome numbers of n —11,
12 and 14 (table 1). Aldama dentata Less. appears to be
closest to species of the genus Rhysolepis (the generic name
Aldama, being based on Aldama dentata Llave & Lex., is
synonymous with Sclerocarpus). It differs from Rhysolepis
in being a smaller plant with less pronounced and fewer
involucral bracts and smaller flowers, but they are very
much alike in floral morphology. While the species of
Rhysolepis (only 2 have been described) have not been
examined for chromosome numbers, what appears to be an
undescribed species (King 3645) has a number of n = 17.

In any case, some disposition of Aldama dentata Less.
(— Sclerocarpus schiedianus) should be made other than its
inclusion in Sclerocarpus; in our opinion, this is suggested
by both the morphological and chromosomal evidence.

Sclerocarpus (x — 11, 12, 14). This genus is in much
need of critical revision (Mr. Feddema of the University of
Michigan is currently undertaking such a study). The
species are quite variable, both morphologically and chrom-
osomally. Turner (1960a) previously reported the chromo-
some number of S. uniserialis (n = 12) ; this species is
apparently closely related to S. dentatus which, as indicated
in table 2, has populations and/or individuals with chromo-
some numbers of both n = 11 and 12.

Spilanthes (x — 13). Chromosome counts of the several
collections listed in table 1 indicate that S. americana
(n — 13, 26, 39) consists of diploid, tetraploid and hexaploid
races and/or taxa. One other species, S. decumbens, has
been reported as diploid with n = 13 (Darlington and Wylie,
1956).

Viguiera. (x — 8, 17, 18). Heiser and Smith (1955) and
Heiser (1960) reported counts of n = 18, 17 and 8 for
species of this genus. Turner (1960) and Turner et al.
(1961) reported several counts of V. dentata as n = 17. V.
longifolia (n. = 8), table 1, is closely related to V. multiflora
which Heiser and Smith reported as n = 8. In our opinion
the lower number is probably a relictual feature, perhaps
being the same as or close to the ancestral diploid number

1962] Turner, Powell & King — Chromosome numbers 257

of the phylad which gave rise to Viguiera and related genera.
It will be interesting to see if this chromosomal hiatus holds
as additional species are examined. Blake (1918) recognized
143 species for the genus but to date only 7 species have
been counted.

Wedelia (x = 11, 12). The only previous chromosome
report for this genus has been that of Turner and Irwin
(1960) for the South American W. brasiliensis (Spreng.)
Blake (n= 29 + 1). The genus, as presently circum-
scribed, is multibasic and apparently includes polyploids.

Zexmenia (x — 10, 11, 14). Heiser and Smith (1955)
have reported one other species, Z. frutescens (Mill.) Blake,
to be diploid with » — 11. With only 5 of the approximately
30 species reported to date (table 1), it appears likely that
additional basic numbers will be added to this multibasic
series.

Chromosome counts for Helianthus (x — T), Pery-
menium (x = 15), Tithonia (x =17) and Verbesina
(x = 16, 17, 18) are consistent with the basic numbers
reported for these genera by other workers (Darlington and
Wylie, 1956; Turner et al., 1961a ; 1961b).

The chromosome counts for species of Hymenostephium
(n = 17), Iostephane (n = 17), Notoptera (n = 15), Podo-
chaenium (n = 17), and Salmea (n = 18) are first reports
for these genera. Beaman and Turner (unpublished) have
obtained chromosome numbers of n = 9 for other species of
Jaegeria, thus the two species with n = 18 (table 1) are
tetraploids.

COREOPSIDINAE — Coreopsis mutica (n= ca. 24, 26).
This species is a shrub up to 3 meters high; while the counts
are only approximate they do indicate the species to be
tetraploid since most taxa in the genus are on a base of
x = 12, 13, 14 (Turner, 1960b). Chromosome counts for
the species of Dahlia (n = 18) and Cosmos (n = 12) are
consistent with the basic numbers established by other work-
ers (Darlington and Wylie, 1956).

GALINSOGINAE — Calea integrifolia (n = ca. 16) — Tur-
ner et al. (1961b) reported a chromosome count of n = ca.
17 for this species. No certain count could be made from the

258 Rhodora [Vol. 64

present collection, but the meiotic figures appeared to be
n = 15 or 16, the latter count being obtained more fre-
quently. The chromosome count for C. trichotoma (n — 18)
is the first unequivocal count for the genus.

Sabazia sp. nov. (n = 18) — Turner and Johnston (1961)
have reported the only other species count, S. humilis (n —
4).:

Galinsoga parviflora (n — 8, 16). This species has here-
tofore been reported as diploid with » — 8 (Haskell and
Marks, 1952) ; the 6 collections from southern Mexico and
Guatemala listed in table 1 were tetraploid, while the 2
collections from Central Mexico were diploid. Haskell and
Marks recognized 2 species in the G. parviflora complex in
the British Isles (both introduced) : G. ciliata (Raf.) Blake
(n = 16) and G. parviflora (n — 8). Fernald (1950)
recognized 4 species as belonging to the complex in the
northeastern United States maintaining both G. parviflora
and G. ciliata. Most authors have distinguished between the
latter 2 taxa by a combination of technical features such
as stem pubescence, absence or presence of a pappus on the
ray florets, etc. (Fernald, 1950; Clapham et al.; etc.). The
several characters used to distinguish these 2 taxa are, in
our opinion, slight and while they might hold for the intro-
duced populations in Britain, they do not hold singly or in
combination for the Mexican material (nor apparently for
the material from temperate North and South America,
although there is a tendency for more glabrate forms in
temperate latitudes; specific recognition of such races hard-
ly seems warranted in view of the widespread, weedy nature
of the taxon).

HELENIEAE — Schkuhria anthemoides (n = 20). Chromo-
some counts for 2 South American species, S. pinnata
(n —10) and S. multiflora (n = 11), have been reported by
other workers (Darlington and Wylie, 1956). As indicated
in table 1, the North American populations of S. pinnata
are apparently tetraploid.

Perityle microglossa (n = ca. 46 + 4). The chromosome
number of a Texas collection of this species was reported
as n = 36 by Turner and Ellison (1960).

1962] Turner, Powell & King — Chromosome numbers 259

Galeana pratensis (n — 9). This is the first chromosome
count reported for the genus.

SENECIONEAE — S. imparipinnatus (n = 23). Chromo-
some counts for most species of the genus Senecio have been
en a base of x = 5; however at least 2 other species are
known with counts of n = 23 (Turner et al., 1961b).

Neurolaena lobata (n — 11). Counts for this genus have
not been reported previously.

MUTISIEAE — T'rixis radialis (n — 27). Only 11 of the
approximately 66 genera in the tribe Mutisieae have been
counted. Including Trixis, 4 of the 11 are unibasie with
x — 27; the unusually high basic number is apparently poly-
ploid in origin, presumably from an ancestral base of
x = 9 (the haploid number, n = 9, is not known in those
members of the tribe studied to date, but the Australian
genus T'richocline has been reported as n — 18).

SUMMARY

Chromosome counts for species of Piqueria (n = 12),
Mexican and Guatemalan Compositae representing 133 taxa
(131 species and 2 varieties) distributed in 66 genera.
Counts of approximately 700 of the species are reported for
the first time including the following genera: Calea, x = 18;
Galeana, x —9; Hymenostephium, x — 17; Iostephane,
x = 17; Jaegeria, x — 9; Neurolaena, x = 11; Notoptera,
x = 15; Philactis, x = 28; Podochaenium, x = 19; Psilactis,
€ = 4, 9, (8?) ; Rhysolepis, x = 17; Salmea, x = 18; Trago-
ceros, x = 11; Trichocoronis, x = 15; Trixis, x = 27; and
Wedelia, x — 11, 12.

Chromosome counts for species of Piqueria (m = 12),
Mikania (n = 17), Polymnia (n = 17), Zinnia (n= 13),
Sclerocarpus (n —11, 14), Wedelia (n=11, 12) and
Zexmenia (n — 10, 14) differ from the reported basic num-
bers as determined from other species in these genera.
When appropriate the chromosomal information has been
related to systematic problems. BOTANY DEPARTMENT AND
THE PLANT RESEARCH INSTITUTE, UNIVERSITY OF TEXAS,

AUSTIN.

260 Rhodora [Vol. 64

M $ T ~ #3 9

18 19

Fig. 1-33. Meiotic chromosomes of species of Compositae. — Fig. 1. Ageratum

corymbosum (n = 10 + 6 fragments) — Fig. 2. Eupatorium morifolium (n — 10) —
Fig. 3. Eupatorium petiolare (n — 11) — Fig. 4. Eupatorium pycnocephalum (n = 20)
- Fig. 5. Eupatorium pycnocephalum (n = 10) — Fig. 6. Piqueria pilosa (n — 12) —
Fig. 7. Trichocoronis wrightii (n = 15) — Fig. 8. Machaeranthera gymnocephala
(n = 4) — Fig. 9. Psilactis cf. asteroides (n = 4) — Fig. 10. Polymnia ef. apus (n = 16)
- Fig. 11. Zinnia maritima (n = 13) — Fig. 12. Iostephane heterophylla (n = 11) —

Fig. 13. Podochaenium eminens (n = 19) — Fig. 14. Salmea scandens (m — 18 + 2

á

1962] Turner, Powell & King — Chromosome numbers 261

P
* e 5
ve

23

fragments) — Fig. 15. Sclerocarpus dentatus (n = 11) — Fig. 16. Sclerocarpus dentatus
(n = 12) — Fig. 17. Sclerocarpus cf. divaricatus (n = 11) — Fig. 18. Sclerocarpus cf.
frutescens (n —12) — Fig. 19. Sclerocarpus cf. phyllocephalus (n = 11) — Fig. 20.
Sclerocarpus sessilifolius (n = 14) — Fig. 21. Spilanthes americana (n = 26 + 4 frag-
ments) — Fig. 22. Wedelia filipes (n — 11) — Fig. 23. Zexmenia cf. aurea (m = 10)
— Fig. 24. Zexmenia costaricensis (n = 11) — Fig. 25. Zexmenia pringlei (n = 14) —
Fig. 26. Zezmenia virgulata (n = 11) — Fig. 27. Sabazia sp. nov (n = 8) — Fig. 28.
Galeana pratensis (n = 9) — Fig. 29. Schkuhria anthemoides (m = 20) — Fig. 30.
Neurolaena lobata (n = 11) — Fig. 31. Senecio imparipinnatus (n = ca. 23) — Fig. 32.
Arctotis stoechadifolia (n = 9) — Fig. 33. Triris radialis (n = 27). x ca. 2000.

262

Rhodora

[Vol. 64

TABLE 1. SPECIES OF COMPOSITAE EXAMINED FOR CHROMOSOME NUMBER

VERNONIEAE

Vernonia, karwinskiana Hort.
n=17+1 HIDALGO: 9 m.
ne. of Jacala. King 4210.

EUPATORIEAE

Ageratum corymbosum Zucc.

n — 10 CHIAPAS: 23 m. se of
Comitán. King 3045.

n —10* (Fig. 1) PUEBLA: 4
m. w. of Izücar de Matamoros.
King 2923.

n — 10* OAXACA: 14 m. ne.
of Huajuapan de Leon. King
3544.

Ageratum houstonianum Mill.
n = 10 GUATEMALA: 4 miles
south of Coban. King 3311.
n — 10 MORELOS: 11 miles
south of Cuernavaca. King

4160.

Agera'um latifolium Cav.

n =10 HIDALGO: 12 m. sw.
of the Hidalgo-San Luis Po-
tosi state border along route
85. King 4226.

n —10 PUEBLA: 6 m. sw. of
the Puebla-Veracruz state
border along route 130. King
4140.

Ageratum paleaceum (Gay)
Hemsl. var. nelsonii Rob.

n — 10° CHIAPAS: 7 m. e. of
the Chiapas-Oaxaca border
along route 190. King 2751.

Ageratum cf. paleaceum (Gay)
Hemsl. var. paleaceum

n =11 +1 CHIAPAS: 22 m.
s. of Las Cruces. King 3112.

Ageratum cf. tomentosum
(Benth.) Hemsl.

n — 10 CHIAPAS: 10 m. e. of
the Oaxaca-Chiapas border
along route 190. King 2981.

Brickellia robinsonii Nels.

n —9 SAN LUIS POTOSI: 2

m. w. of Xilitla. King 4292.

Eupatorium cf. aschenbornia-

num Sch.

n =20° GUATEMALA: 7 m.
w. of Quezaltenango. King
3182.

Eupatorium betonicum Hemsl.

n — 10 MORELOS: 4 m. w. of
Izücar de Matamoros. King
2925.

Eupatorium cf. gracilicaule
Sch.-Bip.

n — 10 CHIAPAS: 24 m. se.
cf Comitán. King 3037.

Eupatorium morifolium P. Mil-
ler

n = 10 (Fig. 2) SAN LUIS
POTOSI: 6 m. ne. of Xilitla.
King 4248.

Eupatorium petiolare Moc.

n =17 SAN LUIS POTOSI:
Near waterfall at E] Salto.
King 3919.

n —17 (Fig. 3) HIDALGO:
5 m. n. of Actopan. King
4199.

Eupatorium pycnocephalum
Less.

n = 20 (Fig. 4) HIDALGO:
12 m. sw. of the Hidalgo-San
Luis Potosi state border along
route 85. King 4224.

n-—20 SAN LUIS POTOSI:
6 m. ne. of Xilitla. King 4242.

n —20 SAN LUIS POTOSI:
near waterfall at El Salto.
King 3875.

n = 20 VERACRUZ: 7 m. s.
of Tampico el Alto. King
4099.

n = 20 VERACRUZ: 10 m. s.

“Indicates one to several fragments.

1962]

of Cerro Azul. King 4119.

n = 20 VERACRUZ: 38 m. s.
of Acayucan. King 2452.

Eupatorium cf. pycnocephalum
Less.

7 —10 (Fig. 5) SAN LUIS
POTOSI: 6 m. ne. of Xilitla.
King 42424.

Eupatorium quadrangulare DC.

n=10 SAN LUIS POTOSI:
near waterfall at El Salto.
King 3853.

Mikania cf. gonzalezii Rob. &
Greenm.

n=17 VERACRUZ: 5 m. w.
of Tuxpan. King 4128.

Orylobus glanduliferus | (Sch.-
Bip.) Gray

n=16 GUATEMALA: 4 m.
e. of Totonicapan. King 3216.

Piqueria pilosa H. B. K.

n —512 MEXICO STATE: 8
m. n. of San Francisco Cheje.
King 3581.

n — 12 (Fig. 6) 9 m. ne. of San
Francisco Cheje. King 3569.

Piqueria trinervia Cav. `

n= 12 CHIAPAS: 19 m. e. of
San Cristóbal de Las Casas.
King 2821.

Trichocoronis wrightii Gray

n =15 (Fig. 7 TAMAULI-
PAS: 2 m. ne. of Altamire.
King 4034.

ASTEREAE

Aster bimater Standl. & Stey-
erm.

n —5 GUATEMALA: 6 m. s.
of Huehuetenango. King 3423.

Aster exilis var. australis Gray

* —5 GUATEMALA: 3 m. e.
of Quezaltenango. King 3205.

"n —5 GUATEMALA: 3 m.s.
of Huehuetenango. King
3418.

m=5 SINALOA: 4 m. w. of
El Palmito. Powell & Ed-

Turner, Powell & King — Chromosome numbers 263

mondson 909.

Baccharis glutinosa Pers.
n=9 OAXACA: 1 m. n. of
Tamazulapan. King 2935.

Baccharis serraefolia DC.

m=9 SAN MARCOS: 4 m.
e. of San Marcos. King 3159.

Chrysopsis ef. villosa (Pursh.)
Nutt.

n —9 DURANGO: City limits
Durango. King 3725.

Conyza canadensis L.

n —9 PUEBLA: 20 m. nw. of
Tehuacan. King 2640.

Conyza coronopifolia H. B. K.

n=9 CHIAPAS: 1 m. e. of
San Cristóbal de Las Casas.
King 2838.

Conyza sophiaefolia H. B. K.

n —59 GUATEMALA: 3 m. e.
of Quezaltenango. King 3206.

Erigeron delphinifolius Willd.

"n -— ca. 9 DURANGO: City
limits of Durango. King 3726.

n=9 DURANGO: 4 km. n.
of Durango. King 3780.

Erigeron heteromorphus Rob.

n=9 SAN LUIS POTOSI:
Waterfall near El Meco. M.
C. Johnston 5116.

Erigeron cf. karvinskianus DC.

n = ca. 27 GUATEMALA:
Between Solola and Pana-
jachel. King 3224.

Erigeron repens Greenm.

n =9 VERACRUZ: 10 m. &
of Tampico el Alto. King
4100.

Erigeron scaposus DC.

mnm =9 PUEBLA: 0.5 m. sw. of
Tehuacan. King 2311.

m=18 OAXACA: 6 m. s. of
Tamazulapan. King 2938.

Erigeron cf. scaposus DC.

n - —18 MEXICO STATE: 4
m. ne. of San Francisco
Cheje. King 3575.

264

Erigeron cf. scaposus DC.

n=9II+9I MEXICO STATE:
11 m. e. of the Mexico-Mich-
oacan state border along
route 15. King 93597.

Erigeron cf. scaposus DC.

n=9 OAXACA: 21 m. n. of
the junction with route 190.
King 3498.

Erigeron. sp.

» —9 TAMAULIPAS: 9m. s.
of Ciudad Victoria. King
4537.

Haplopappus stoloniferus DC.

n=4 CHIAPAS: 5 m. e. of
San Cristóbal de Las Casas.
King 2805.

Heterotheca inuloides Cass. var.
inuloides

n=9 OAXACA: 40 m. se. of
Oaxaca. King 2897.

Machaeranthera gymnocephala
(DC.) Shinners

n-—4 DURANGO: 15 m. sw.
of Durango. King 3738.

n= (Fig. 8) MEXICO
STATE: 19 m. w. of Toluca.
King 3594.

Machaeranthera tanacetif olia
(H. B, K.) Nees.

n-—4 CHIHUAHUA: 14 m.
sw. of Chihuahua City. Powell
& Edmondson 976.

n-— 4 DURANGO: 13 m. n. of
Durango. King 3754.

Psilactis asteroides Gray

"n — 4 DURANGO: City limits
of Durango. King 3728.

Psilactis cf. asteroides Gray

n= (Fig. 9) MEXICO
STATE: 22 m. w. of Toluca.
King 8595.

Psilactis brevilingulata Sch.-Bip.

m=ea. 9 (81-2 frags?)
MEXICO STATE: 10 m. w.
of Toluca. Powell & Edmond-
son 799.

Rhodora

[Vol. 64

m=9 OAXACA: 6 m. s. of
Tamazulapan. King 2939.

n — ca. 9 OAXACA: By the
ruins at Monte Alban. King
2899.

n=ca. 9 QUERETARO: 18
m. s. of San Luis de la Paz.
Powell & Edmondson 573.

Solidago velutina DC.

n-—9 DURANGO: 71 m. ne.
of Durango. King 3762.

Solidago wrightii Gray

n=9 DURANGO: 24 m. sw.
of Durango. King 3742.

Xanthocephalum gymnosper-

moides (Gray) Benth. &
Hook.

n-—6 CHIHUAHUA: 0.5 m.
w. of Cuahutemoc. Powell &
Edmondson 1000.

Xanthocephalum hwmile
(H.B.K.) Sch.-Bip.

n=4 MEXICO STATE: 12
m. n. of San Francisco Cheje.
King 3586.

m —4 MEXICO STATE: 42
km. s. of Mexico City. King
2908.

m= 4 MORELOS: 15 m. n. of
Cuernavaca. Powell & Ed-
mondson 736.

INULEAE

Gnaphalium leptophyllum DC.
n = 7 MEXICO STATE: 8m.
e. of the Mexico-Michoacan
state border along route 15.

King 3598.

n=7 GUATEMALA: Just
west of San Marcos. King
3131.

Gnaphalium cf. leptophyllum
DC.

n = ca. 14 GUATEMALA: 1
m. w. of Quezaltenango. King
8186.

Pluchea odorata (L.) Cass.

n —10 HIDALGO: 12 m. sw.

1962]

of the Hidalgo-San Luis Po-
tosi state border along route
85. King 4228.

Pluchea purpurascens (Sw.)
DC,

n =10 NAYARIT: Behind the
beach at San Blas. King 3697.

HELIANTHEAE

Subtribe Melampodinae

Berlandiera lyrata Benth.

n — 15 DURANGO: City limits
of Durango. King 3730.

Dugesia mexicana Gray

m =18 MEXICO STATE:
Mexico City area. Powell &
Edmondson 604.

n =18 PUEBLA: 14 m. w. of
Puebla. King 3561.

. Parthenium tomentosum DC.

"m — 18 OAXACA: 23 m. se. of
Oaxaca. King 3482.

Polymnia cf. apus Blake

n = 16 (Fig. 10) NAYARIT:
1m. n. of Tepic. King 3688.

Polymnia maculata Cav.

n=16 SAN LUIS POTOSI:
7 m. ne. of Xilitla. King 4427.

Polymnia cf. maculata Cav.

n — 17 GUATEMALA: 1m.s.
of Coban. King 3306.

Polymnia oaxacana Sch.-Bip.

n=ca. 16 GUATAMALA: 2
m. w. of Santa Cruz Verapaz.
King 3338.

Subtribe Ambrosinae

Ambrosia cf. cumanensis
H. B. K.

n — ca. 36 MICHOACAN: 21
m. e. of Jiquilpan. King 3643.

Ambrosia peruviana Willd.

n=18 GUATEMALA: 4 m.
s. of Coban. King 3313.

SUBTRIBE Zinninae

Heliopsis buphthalmoides
(Jacq.) Dunal.

n =28 GUATEMALA: 9 m.
s. of Salama. King 3263.

Turner, Powell & King — Chromosome numbers 265

n = ca. 28 CHIAPAS: 10 m.
e. of Teopisca. King 3030.

Philactis nelsonii (Greenm.)
Blake

n = ca. 28 CHIAPAS: 2 m. w.
of Chiapas-Oaxaca border
along route 190. King 2878.

n= 28 CHIAPAS: 21 m. ne.
of Las Cruces. King 3446.

Sanvitalia cf. procumbens Lam.

n—8 CHIAPAS: 12 m. e. of
Cintalapa. King 2984.

Sanvitalia procumbens Lam.

n= 8 PUEBLA: 22 m. ne. of
Tepeaca. Powell & Edmond-
son 639,

Sanvitalia sp. nov.

n-—8 OAXACA: 1 m. n. of
Tamazulapan. King 2929.
Tragoceros americanum (Mill.)

Blake

m —11 JALISCO: 16 m. s. of
Guadalajara. King 3658.

Tragoceros mocinianus Gray

n = ca. 11 JALISCO: 45 m. w.
of Sahuayo. Powell & Ed-
mondson 850.

Tragoceros schiedeanus Less.

n =11 JALISCO: 45 m. w. of
Sahuayo. Powell & Edmond-
son 848.

Zinnia angustifolia H. B. K.

»1-—12 NAYARIT: 8 m. se.
of Tepic. King 3672.

Zinnia elegans Jacq.

"1 —12 GUERRERO: 5 m. n.
of Ocotito on the Chilpan-
cingo Acapulco highway.
Johnston 5988.

Zinnia leucoglossa Blake

n=11 DURANGO: 14 m. sw.
of Durango. King 3739.

Zinnia maritima H. B. K.

n — 13 (Fig. 11) GUERRERO:
10 m. e. of Acapulco. King
4180.

Zinnia peruviana (L.) L.

266

n —12 OAXACA: 1 m. se. of
Oaxaca. King 3466.

Zinnia tenella Rob.

n — 11 or 12 DURANGO: 15
m. sw. of Durango. King
37306.

SUBTRIBE Verbesininae

Aldama dentata Less.

n — 11 OAXACA: 9m. nw. of
Oaxaca. King 3516.

n — 17 NAYARIT: City limits
of Tepic. King 3668.

n — 17 MORELOS: 5 m. se. of
Yautepec. King 2912.

n=17 PUEBLA: Just se. of
the Morelos-Puebla border.
King 2919.

m — 17 VERACRUZ: 19 m. se.
of Poza Rica. King 4137.

n =17 VERACRUZ: 1m. s.
of Cuitlahuac. King 2677.

m — 17 OAXACA: Along route
190, just south of Etla. King
2509.

Helianthus laciniatus Gray

n = 17 DURANGO: City limits
of Durango. King 3756.

Hymenostephium sp.

n = ca. 17 NAYARIT: 11 m.
se. of Tepic. King 3675.

Iostephame heterophylla var.
dicksonii (Lindl.) Sharp

n=17 (Fig. 12) SINALOA:
1-2 m. sw. of the Sinaloa-
Durango border. King 3721.

m — 17 DURANGO: 24 m. sw.
of Durango. King 3741.

Iostephame trilobata Hemsl.

n — 17 OAXACA: 10 m. ne. of
Oaxaca. Powell & Edmond-
son 676.

Jaegeria hirta Less.

m=18 GUATEMALA: Be-
tween Solola and Panajachel.
King 3236.

Jaegeria pedunculata H. & O.

"«-— 18 NAYARIT: 5 m. se. of

Rhodora

[Vol. 64

Tepic. King 3670.

Notoptera tequilana (Gray)
Blake

n — ca. 15 JALISCO: 26 m.
nw. of Tequila. King 3664.

Perymenium ef, asperifolia
Sch.- Bip.

n = ca. 45 OAXACA: 5m. n.
of the junction with route
190. King 3488.

Perymenium cf. chalarolepis
Rob. & Greenm.

n — 15 1 m. w. of San Marcos.
King 3138.

Podochaenium eminens (Lag.)
Sch.-Bip.

n — 19 OAXACA: Along route
175, 10 m. n. of the junction
with route 190. King 3494.

n — 19 (Fig, 13) SINALOA: 5
m. sw. of the Sinaloa-Dur-
ango border along route 40.
King 3720.

Rhysolepis sp. nov.

n-— 17 MICHOACAN: 2 m
e. of Zamora. King 3645.

Salmea scandens (L.) DC.

n — 18" (Fig. 14) SAN LUIS
POTOSI: 2 m. w. of Xilitla.
King 4309.

Sclerocarpus dentatus (Llave &
Lex.) Hemsl.

n —12 COAHUILA: 37 m. s.
of Monclova. Powell & Ed-
mondson 506,

n = 11 (Fig. 15) OAXACA: 49
m. w. of Tehuantepec. King
3458.

n = 12 (Fig. 16) OAXACA: 1
m. ne. of Huajuapan de Leon.
King 3533.

n —12 TAMAULIPAS: 2 m.
ne. of Altamira. King 4086.
m =12 VERACRUZ: 4 m. n.

of Tampico el Alto. King
4094.
n-— 12 VERACRUZ: 3 m. s

1962]

of Naranjos. King 4118.

n=12 SAN LUIS POTOSI:
Just north of the San Luis
Potosi-Hidalgo state border.
King 4233.

Sclerocarpus cf. divaricatus
(Benth.) Hemsl.

^ = 11 (Fig. 17) VERACRUZ:
20 m. s. of Acayucan. King
2733.

Sclerocarpus cf. frutescens
Brandegee

n — 12 (Fig. 18) OAXACA: 1
m. n. of Tamazulapan. King
2936.

Selerocarpus cf. phyllocephalus
Blake

n = 11
km. w.
3372.

n = 11 (Fig. 19) CHIAPAS:
Along the railroad track to
Tapachula at the village of
Soconusco. King 3126.

Sclerocarpus cf. phyllocephalus
Blake

n — 12 CHIAPAS: 5 m. ne. of
Las Cruces. King 2440.

Sclerocarpus sessilifolius
Greenm.

"n — 14 (Fig. 20) NAYARIT:
8 m. se. of Tepic. King 3673.

Spilanthes americana Hieron.

"n —ca. 26 SAN LUIS PO-
TOSI: 6 m. ne. of Xilitla.
King 4245.

n — 25 +1 VERACRUZ: 5m.
n. of Tampico el Alto. King
4092.

n = ea. 13 PUEBLA: 6 m. sw.
of the Puebla-Veracruz state
border along route 130. King
4141.

"n — ca. 26 HIDALGO: 14 m.
ne. of Jacala. King 4222.

»n 250 Xm 201) VERA.
CRUZ: 9 m. sw. of Tuxpan.

GUATEMALA: 4-5
of Escuintla. King

Turner, Powell & King — Chromosome numbers 267

King 4132.

n — ca. 26 VERACRUZ: 25 m.
se. of Poza Rica. King 4139.

n — ca. 26 MORELOS: 5 m.
se. of Yautepec. King 2913.

Spilanthes americana cf. var.
stolonifera (DC.) Moore

n = ca. 39 MICHOACAN: 21
m. e. of Jiquilpan. King 3642.

Tithonia longeradiata (Berl.)
Blake

n =17 GUATEMALA: 10 m.
s. of Quezaltenango. King
3428.

Verbesina erocata (Cav.) Less.

n —18 MORELOS: 11 m. s.
of Cuernavaca, King 4159.

Verbesina hypargyrea Rob. &
Greenm.

m= ca. it, CHIAPAS: I8 m.
se. of Comitán. King 3044.

n —17 OAXACA: 41. m. w.
of Tehuantepec. King 3456.

"n —517 COAHUILA: 14 m. s.
of Saltillo. Powell & Ed-
mondson 538.

Verbesina seatonii Blake

n —517 MEXICO STATE: 9
m. ne. of San Francisco
Cheje. King 3567.

Viguiera grammatoglossa DC.

n —17 OAXACA: 27 m. n.
of the junction along route
190. King 3504.

Viguiera longifolia (Rob. &
Greenm.) Blake
^ —8 CHIAPAS: Wet fields

just west of San Cristóbal de
Las Casas. King 2993,
Wedelia filipes Hemsl.
n —11 GUATEMALA: Near
Panajachel. King 3239.

n —11 (Fig. 22) GUATE-
MALA: Near Panajachel.
King 3240.

Wedelia parviceps Blake
" —12 GUATEMALA: 9 m.

268

n. of Salama. King 3283.
Zexmenia cf. aurea (DC.)
B. & H.
n — 10 (Fig. 23) JALISCO: 17
m. nw. of Tequila. King 3663.
Zexmenia costaricensis Benth.
n-—11 (Fig. 24) ALTA VERA-
PAZ: 11 m. w. of San Cris-
tóbal Verapaz. King 3347.
Zexmenia pringlei Greenm.
n —14 (Fig. 25) OAXACA:
Just s. of the Oaxaca-Puebla
border along route 125. King

3547.
Zermenia virgulata Klatt
n-—11 (Fig. 26) GUATE-

MALA: 10 m. s. of Huehue-
tenango. King 3392.

SUBTRIBE Coreopsidinae

Coreopsis mutica DC,

n — ca. 26 CHIAPAS: 13 m.
e. of San Cristóbal de Las
Casas. King 2813.

m — ea. 24 CHIAPAS: 14 m.
w. of San Cristóbal de Las
Casas. King 3082.

Cosmos diversifolius Otto

n=12 OAXACA: 30 m. nw.
of Oaxaca. King 3525.

n —129 OAXACA: 29 m. nw.
of Oaxaca. King 3523.

Dahlia dissecta S. Wats.

n —18 MEXICO STATE: Sa-
vannah-like forest on route

190 at Llano Grande. Rock
M-351.

Dahlia scapigera (A. Dietr.)
L. & O.

n —18 PUEBLA: 14 m. w. of
Texmelucan. King 3564.

SUBTRIBE Galinsoginae

Calea integrifolia (DC.) Hemsl.

n = ca. 16 PUEBLA: 5 m. ne.
of Villa Juarez. King 4143.

Calea nelsonii Rob. & Greenm.

m = ca. 18 CHIAPAS: 10 m. e.
of the Oaxaca-Chiapas bor-

Rhodora

[Vol. 64

der. King 2982.

Calea cf. trichotoma D. Smith

n —18 CHIAPAS: 23 m. se.
of Comitán. King 3043.

Galinsoga parviflora Cav.

n — ca. 16 CHIAPAS: Just w.
of San Cristóbal de Las
Casas. King 2991.

n-— 16 GUATEMALA: Along

National Route 1, between
Solola and Panajachel. King
3225.

n —16 GUATEMALA: Along
National Route 1, between
Solola and Panajachel. King
3235.

n = 16 MICHOACAN: 2m. n.
of Zitacuaro. King 3603.

n =16 MICHOACAN: 5 m.
w. of Morelia. Powell & Ed-
mondson. 825.

m -— 16 OAXACA: 12 m. ne.
of Oaxaca. Powell & Edmond-
son 684.

n-—8 MEXICO: 10 m. w. of
Toluca. Powell & Edmondson
804.

n=8 PUEBLA: 10 m. e. of
Puebla. Powell & Edmondson
624.

Sabazia sp. nov.

n = 8 (Fig. 27) OAXACA: 21
m. n. of the junction with
route 190. King 3499.

HELENIEAE
Galeana pratensis (H. B. K.)

Rydb.

n = 9 (Fig. 28) JALISCO: 12
m. nw. Guadalajara. King
3661.

n=9 GUATEMALA: 15 m.

s. of Rabinal. King 3363.
Schkuhria anthemoides var.
wislizeni (Gray) Heiser
n = 20 (Fig. 29) OAXACA:
17 m. se. of Nochistlan. King

3527.

1962] Turner, Powell & King — Chromosome numbers 269

Schkuhria pinnata var.
virgata (Llave) Heiser

n=21+1 GUATEMALA: 8
m. s. of Huehuetenango. King
3426.

n —ca. 20 GUATEMALA: 1
m. s. of Huehuetenango. King
3413.

Perityle microglossa Benth.

n — ca. 46 + 4 SAN LUIS PO-
TOSI: Near the waterfall at
El Salto. King 3877.

ANTHEMIDEAE

Chrysanthemum parthenium
Benth.

n=9 OAXACA: 10 m. n. of
the junction with route 190.
King 3495.

SENECIONEAE

Neurolaena lobata (L.) R. Br.

"n —11 (Fig. 30) SAN LUIS
POTOSI: 2 m. w. of Xilitla.
King 4276.

Schistocarpha bicolor Less.

n=8 VERACRUZ: 5 m. ne.
of Villa Juarez. King 4144.

n —8 HIDALGO: 9 m. sw. of
the Hidalgo-San Luis Potosi
state border along route 85.
King 4230.

^ —8 SAN LUIS POTOSI: 6
m. generally ne. of Xilitla.
King 4241.

Senecio cobanensis var.
sublanciniatus Greenm.

n= ca. 65 GUATEMALA: 11
m. s. of Salama. King 3270.

Senecio confusus Britton

n=—45+5 SAN LUIS PO-
TOSI: 2-3 m. w. of El Na-
ranjo. King 3973.

Senecio conzatii Greenm.

n—=20 OAXACA: Along route
175, 12 m. n. of the junction
with route 190. King 3496.

Senecio deformis Klatt

7^-— ca. 20 DISTRITO FED-

ERAL: El Zarco on route 15,
outside Mexico City. Rock
M-412.

m =20 MEXICO STATE: 8
m. n .of San Francisco Cheje.
King 3583.

n —20 MEXICO STATE: 11
m. e. of the Mexico-Michoacan
state border along route 15.
King 3596.

Senecio hirsuticaulis Greenm.

7-— 30 SAN LUIS POTOSI:
17 m. e. of Ciudad del Maiz.
Johnston 5104.

Senecio imparipinnatus Klatt

n = ca. 23 (Fig. 31) TAMAU-
LIPAS: 4 m. s. of Ciudad
Monte. King 3845.

Senecio picridis Schaur.

"n-—20 GUATEMALA: 11 m.
w. of Quezaltenango. King
3191.

m=20 MORELOS: 3 m. s.
of the Morelos-Federal Dis-
trict border along route 95.
King 4151.

Senecio salignus DC.

n = 30 HIDALGO: 1 m. e. of
Pachuca. King 4148.

n= 30 MEXICO STATE: 9
m. sw. of the pyramids at
Teotihuacan. King 4149.

Senecio toluccanus DC.

m = 20 MEXICO STATE: 9
m. ne. of San Francisco
Cheje. King 3574.

Cacalia sp.

n = ca. 30 JALISCO: 21 m.
se. of Guadalajara. King
3656.

Cacalia sinuata Llave & Lex.

n= 30 DURANGO: 6 m. sw.
of Durango. King 3734.

Cacalia cf. tussilaginoides
H. B. K.

m — ca. 25 JALISCO: 3 m. nw.
of Guadalajara. King 3659.

270 Rhodora [Vol. 64
ARCTOTIDEAE 2768.
Arctotis stoechadifolia Berk. n— 27 SAN LUIS POTOSI:
n=9 (Fig. 32) GUATE- Near the waterfall at El
MALA: 6 m. ne. of Quezal- Salto. King 3904.
tenango. King 3210. n = 27 (Fig. 33) SAN LUIS
MUTISIEAE POTOSI: 2 m. w. of Xilitla.
Trixis californica Kellogg King 4291.
n = 27 CHIHUAHUA: 30 m. CICHORIEAE

w. of Chihuahua City. Powell
& Edmondson 1004.

Trixis radialis (L.) Kuntze

n = ca. 27 CHIAPAS: 9 m. e.
of Tuxtla Gutierrez. King

Lactuca pulchella (Pursh) DC.

n -171 GUATEMALA: 9 m.
w. of San Cristóbal Verapaz.
King 3346.

LITERATURE CITED

BEAMAN, J. H., D. C. D. DEJoNG AND W. P. STOUTAMIRE. 1962. Chro-
mosome studies in the Alpine and subalpine floras of Mexico and
Guatemala. Am. Jour. Bot. 49: 41-50.

BLAKE, S. F. 1918. A revision of the genus Viguiera. Contr. Gray
Herb. 54: 1-205.

1930. Notes on certain type specimens of American
Asteraceae in European herbaria. Contr. U. S. Nat. Herb. 26:
227-263.

CLAPHAM, A. R., T. G. TUTIN AND E. F. WARBURG. 1952. Flora of the
British Isles. Cambridge. 1591 p.

DARLINGTON, C. D. AND A. P. WYLIE. 1956. Chromosome atlas of
flowering plants. p. 519. Macmillan Co., New York.

FERNALD, M. L. 1950. Gray's Manual of Botany. Eighth edition.
American Book Co. New York, 632 p.

FisuER, R. T. 1957. Taxonomy of the genus Heliopsis (Compositae).
Ohio Jour. Sci. 57:171-191.

GAISER, L. O. 1953. Chromosome studies in Kuhniinae (Eupatorieae).
I. Brickellia. Rhodora 55: 253-267.

GRAY, A. 1886. Compositae. Synoptical flora of North America. 2nd
ed., 1^: 48-455,

HASKELL, G. AND G. E. Marks. 1952. Chromosome ecology of British
Galinsoga species. New Phytol. 51: 382-387.

HEISER, C. B. AND D. M. SurTH. 1955. New chromosome numbers in
Helianthus and related genera (Compositae). Proc. Indiana Acad.
64 : 250-253.

Heiser, C. B. 1960. Documented chromosome numbers of plants.
Madrono 15: 219-221.

HOFFMANN, O. 1894. Compositae. I» K. Engler and A. Prantl, Die
Natürlichen Pflanzenfam. IV (5):87-391.

MANGENOT, S. AND G. MANGENOT. 1958. Deuxième liste de nombres

1962] Beaman and Turner — Chromosome numbers 271

chromosomíques nouveaux chez diverses dicotylédones et monoco-
tylédones d'Afrique occidentale. Bull. Jard. Bot. Bruxelles 28:
315-329.

MONTGOMERY, F. H. AND S. YANG. 1960. Cytological studies in the
genus Erigeron. Canad. Jour. Bot. 38: 381-386.

Sato, D. 1960. The protokaryotype and phylogeny in plants. Sci. Pap.
Coll. Gen. Educ., Univ. Tokyo 10: 303-327.

SOLBRIG, O. T. 1961. Synopsis of the genus Xanthocephalum. Rhodora
63: 151-164.

Torres, A. M. 1961. Hybridization studies in Zinnia. Am. Jour. Bot.
48:549. (Abstract)

TURNER, B. L. 1960a. Documented chromosome numbers of plants.
Madrono 15: 219-221.

.1960b. Meiotic chromosome numbers in Texas species

of the genus Coreopsis (Compositae-Heliantheae). Southw. Nat.

5: 12-15.

AND W. L. ELLISON. 1960. Chromosome numbers in the
Compositae. I. Tex. Jour, Sci. 12: 146-151.

AND H. S. IRWIN. 1960. Chromosome numbers in the
Compositae. II. Meiotie counts for fourteen species of Brazilian
Compositae. Rhodora 62: 122-126.

, J. BEAMAN AND H. F. L. Rock. 1961a. Chromosome
numbers in the Compositae. V. Mexican and Guatemalan species.
Rhodora 63: 121-129.

, W. L. ELLISON AND R. M. KING. 1961b.. Chromosome
numbers in the Compositae. IV. North American species, with
phyletic interpretations. Am. Jour. Bot. 48: 216-223.

and M. C. JOHNSTON. 1961. Chromosome numbers in
the Compositae. III. Certain Mexican species. Brittonia 13: 64-69.

CHROMOSOME NUMBERS IN MEXICAN AND
GUATEMALAN COMPOSITAE'

J. H. BEAMAN AND B. L. TURNER

The chromosome numbers reported here were obtained
from bud material collected by Beaman in the summer of
1960. The counts were made by Turner (except in Seiges-
beckia which Beaman examined) from pollen mother cell
squashes as outlined by Turner and Ellison (1960). The
voucher specimens were determined by Beaman, except

Supported by National Science Foundation grants G-9025 and G-9045.

272 Rhodora [Vol. 64

Helianthella quinquenervis which was determined by Dr.
C. B. Heiser, Jr., and they are deposited in the Michigan
State University Herbarium. A nearly complete set is also
in the Herbarium of the University of Texas. All material
studied is listed in Table 1.

DISCUSSION

EUPATORIEAE — Piqueria pilosa (n=12). Turner, Powell, and
King (1962) reported two other collections of this species as n=12.

ASTEREAE — Erigeron coronarius (n=18). Diploid counts of n
—9 were obtained by De Jong and Longpre (unpublished) in two
Durango collections of this species. E. coronarius as presently under-
stood is polymorphic, and further study may indicate that the varia-
tions are correlated with chromosomal races.

Xanthocephalum linearifolium (n=4). The count in this collection
was noted by Solbrig (1961) who reported counts for three species of
Xanthocephalum.

HELIANTHEAE — Bidens triplinervia var. macrantha (n=12).
Tetraploid and hexaploid populations of this widely distributed and
variable taxon occur on the Guatemalan volcanoes of Pleistocene or
recent age (Beaman, De Jong, and Stoutamire, 1962). This diploid
material comes from the older, non-volcanie Sierra de los Cuchuma-
tanes.

Jaegeria petiolaris (n—9). Turner, Powell and King (1962)
have found two other species (J. pedunculata Hook. & Arn. and J.
hirta Less.) with n=18. The present report establishes the tetraploid
nature of the other counts.

Calea sp. ? (n—16). Turner et al. (1961 (1962) have reported
approximate counts of n —16, 17, and 18 for several other species, and
they have obtained a definitive count for only one species, C. tricho-
toma with »-—18. The generic determination of our material may
possibly be in error; if it belongs to the genus Calea, it is apparently
undescribed.

Siegesbeckia nudicaulis (n=15), S. repens (n—15), S. triangularis
(n—15). The first two of these species are very restricted endemics,
while the latter is more widespread and may be conspecific with S.
orientalis (sens. lat.) in which Diers (1961) found »—15.

HELENIEAE — Microspermum debile (n=12). This is the first
count reported in this small and little collected genus.

SENECIONEAE — Werneria nubigena (n=50). Counts in 13
species of Werneria, including a count of 2n=212+8 in W. nubigena,
were reported by Diers (1961). All but one species he investigated had
somatic numbers of 100 or higher.

SUMMARY
Meiotic chromosome counts are reported for 25 species of

1962] Beaman and Turner — Chromosome numbers 273

Mexican and Guatemalan Compositae. These include the
first counts published for 16 taxa. The first report is given
for the genus Microspermum (n—12). A count of n=9 was
found for Jaegeria petiolaris, thus establishing the tetra-
ploid nature of previous chromosome reports of »—18 for
this genus. Numbers consistent with established basic num-
bers or previously reported counts were found in Piqueria,
Erigeron, Grindelia, Heterotheca, Xanthocephalum, Heli-
anthella, Sanvitalia, Siegesbeckia, Viguiera, Florestina, Hel-
enium, Cacalia, Senecio, Cirsium, and Hieracium. Numbers
differing with previously reported counts or basic numbers
were obtained in Bidens triplinervia var. macrantha (n=
12), Calea sp. ? (n=16), and Werneria nubigena (n—50).
— DEPARTMENT OF BOTANY AND PLANT PATHOLOGY, MICHI-
GAN STATE UNIVERSITY, EAST LANSING; PLANT RESEARCH
INSTITUTE, THE UNIVERSITY OF TEXAS, AUSTIN.

^e £9388 «Vti Pete

1 2

Figs. 1-4. Meiotie chromosomes of selected species, approximately X 2000. Fig. 1.
Jaegeria petiolaris (n—9). Fig. 2. Calea sp. ? (n—16). Fig. 3. Microspermum debile

(n—12). Fig. 4. Werneria nubigena (n—50).

274 Rhodora [Vol. 64

LITERATURE CITED

BEAMAN, J. H., D. C. D. DE JoNG, AND W. P. STOUTAMIRE. 1962.
Chromosome studies in the alpine and subalpine floras of Mexico
and Guatemala. Amer, Jour. Bot. 49: 41-50.

DrERS, L. 1961. Der Anteil an Polyploiden in den Vegetationsgürteln
der Westkordillere Perus. Zeitsch. für Botanik 49: 437-488.
SoLBRIG, O. T. 1961. Synopsis of the genus Xanthocephalum (Com-

positae). Rhodora 63: 151-164.

TURNER, B. L., AND W. L, ELLISON. 1960. Chromosome numbers in
the Compositae. I. Meiotic chromosome counts for 25 species of
Texas Compositae including 6 new generic reports. Texas Jour.
Sei 12: 146-151.

TURNER, B. L., W. L. ELLISON, AND R. M. KING. 1961. Chromosome
numbers in the Compositae. IV. North American species, with
phyletic interpretations. Amer. Jour. Bot. 48: 216-223.

TURNER, B. L., AND M. C. JoHNSTON. 1961. Chromosome numbers in
the Compositae — III. Certain Mexican species. Brittonia 13: 64-
69.

TURNER, B. L., M. POWELL, AND R. M. KING. 1962. In press. Chrom-
osome numbers in the Compositae. VI. Additional Mexican and
Guatemalan species. Rhodora 64: 251-271.

TABLE 1. Summary of collections studied.

Taxon Locality n chromosome number

EUPATORIEAE
Piqueria pilosa H.B.K. State of Mexico. Nevado de To- 12
luca, northwest side of mtn., ca.
3715 m. alt. Beaman 3471.
ASTEREAE
Erigeron coronarius Greene State of Mexico. 5 kilometers 18
north of Atlacomulco, ca. 2500
m. alt. Beaman 8370.
State of Mexico. 6 kilometers 18
north of Atlacomulco; ca. 2500
m. alt. Beaman 3358a.
Nuevo Leon. Cerro Potosí, sum- 6
mit of mtn., ca. 3650 m, alt. Bea-
man 3340.
State of Mexico. 3 kilometers 9

E. scaposus DC.

Grindelia inuloides Willd.

Heterotheca inuloides Cass.

var. inuloides

Xanthocephalum lineari-
folium (DC.)
Greenman

southeast of Amecameca; ca. 2500
m. alt. Beaman 3474.

Distrito Federal, Mexico. At La
Cima Station between Mexico and
Cuernavaca, 3035 m. alt. Beaman
3653.

4

1962] Beaman and Turner — Chromosome numbers

HELIANTHEAE

Bidens triplinervia var.
macrantha (Wedd.)
Sherff

Helianthella quinquenervis
(Hook.) A. Gray

Jaegeria petiolaris Robins.

Calea sp. ?

Sanvitalia procumbens
Lam.

Siegesbeckia nudicaulis
Standl. & Steyerm.

S. repens Robins. &
Greenm.

S. triangularis Cav.

Viguiera hemsleyana Blake

HELENIEAE

Florestina pedata (Cav.)
Cass.

Helenium hoopesii A. Gray

Guatemala. Dept. of Huehueten-
ango. Between Tojiah and Chem-
al at Km. 319.5 on Ruta Nacional
9 N, ca. 3380 m. alt. Beaman
3743.

Nuevo Leon. Cerro Potosi, near
summit of mtn. on northeast side,
ca. 3600 m. alt. Beaman 3339.
Michoacan. In large llano ca. 4
kms. southeast of Cerro San An-
dres, ca. 10 kms. (straight line
distance) north of Ciudad Hidal-
go, ca. 2930 m. alt. Beaman 4245.
Guatemala. Dept. of Huehuetan-
ango. Between Tojiah and San
Juan Ixcoy at Km. 323.5 on Ruta
Nacional 9 N, ca. 3200 m. alt.
Beaman 3956.

Puebla. 1.5 miles west of Cha-
chapa, ca. 2300 m. alt. Beaman
3615.

Guatemala. Dept. of Huehueten-
ango, between Tojiah and San
Juan Ixcoy at Km. 324.5 on Ruta
Nacional 9 N, ca. 3140 m. alt.
Beaman 3844.

Oaxaca. Llano de las Flores, on
the Oaxaca-Valle Nacional high-
way 20 kilometers east of Ixtlan,
ca. 2870 m. alt. Beaman 3699.
Guatemala. Dept. of Huehueten-
ango, between Tojiah and San
Juan Ixcoy at Km. 323.5 on Ruta
Nacional 9 N, ca. 3200 m. alt.
Beaman 3849.

Oaxaca. On the Oaxaca-Valle
Nacional highway, 8 miles east
of Ixtlan, ca. 3600 m. alt. Beaman
3662.

Puebla. 1.5 miles west of Cha-
chapa, ca. 2300 m. alt. Beaman
3616.

Nuevo Leon. Cerro Potosí, sum-

275

12

9 (Fig. 1)

16 (Fig. 2)

15

15

15

ca
34

10

15

276

Microspermum debile
Benth.

SENECIONEAE
Cacalia peltata H.B.K.

Senecio conzattii Greenm.

S. iodanthus Greenm.

Werneria nubigena H.B.K.

CYNAREAE
Cirsium skutchii Blake

CICHORIEAE
Hieracium mexicanum Less.

H. selerianum Zahn

Rhodora [ Vol.

mit of mtn., ca. 3650 m. alt. Bea-
man 3347.

Oaxaca. On the Oaxaca-Valle
Nacional highway, on Cerro Pe-
long, 25.5 miles east of Ixtlan, ca.
2950 m. alt. Beaman. 3663,

Puebla. 5.3 miles southwest of
San Salvador el Seco; ca. 2500 m.
alt. Beaman 3619.

Oaxaca. Llano de las Flores, on
the Oaxaca-Valle Nacional high-
way 20 kms. east of Ixtlan, ca.
2870 m. alt. Beaman 3700.
Michoacan. Summit of Cerro San
Andres, ca. 12 kms. (straight line
distance) north of Ciudad Hidal-
go, 3589 m. alt. Beaman 4241.
Guatemala. Dept. of Huehueten-
ango. Between Tojiah and Chem-
al at Km. 319.5 on Ruta Nacional
9 N, ca. 3380 m. alt. Beaman

3741.

Guatemala. Dept. of Huehueten-
ango. Between Tojiah and Chem-
al at Km, 319.5 on Ruta Nacional
9 N, ca. 3380 m. alt. Beaman
3742.

State of Mexico. Iztaccihuatl,
south side of mtn. at La Joya, ca.
3990 m. alt. Beaman 3501.

Guatemala. Dept. of Huehueten-
ango. Between Tojiah and Chem-
al at Km. 317.5 on Ruta Nacional
9 N, ca. 3400 m. alt. Beaman

3802,

64

12 (Fig. 3)

30

20

20

50 (Fig. 4)

17

1962] Nevling — Thymelaeaceae 277

CHROMOSOME COUNTS OF TWO THYMELAEACEAE
LORIN I. NEVLING, JR.

The earliest reports of chromosome numbers in the
Thymelaeaceae were published shortly after the turn of this
century, as incidental remarks, in discussions of endosperm
formation, embryo sac development, and parthenogenesis.
Chromosome numbers for nearly forty taxa, in seven genera,
have been reported, a small percentage considering that
several hundred species in fifty-five genera are recognized.
The authors of these reports understandably have had little
interest in the family as a group and, therefore, rarely have
correlated their data with the taxonomic system. Domke’s
(1934) realignment of the classification down to the generic
level has increased taxonomic interest and research of both
the revisionary and floristic types. His broad study is based
primarily on anatomy and morphology and makes no
attempt to include the meager cytological data then avail-
able. There have been few opportunities, since that time, to
profitably apply cytological information to taxonomic prob-
lems because of the nature of the groups under study, i.e.,
revisionary investigations have been limited to genera which
are cytologically unknown.

Although the chromosome counts are restricted, with a
few exceptions, to the larger and more accessible genera,
ie. Daphne, Pimelea, and Wikstroemia, even these few
reports have demonstrated an interesting biological com-
plexity to the family and some of the accompanying taxo-
nomic consequences. Horticulturists, cytologists, and
taxonomists, for example, for considerable time have known
Daphne odora Thunb. to be sterile. Osawa (1913) attempted
to explain this lack of seed development but he misdeter-
mined the chromosome number and succeeded only in pre-
cipitating a series of papers on the subject. Okura and Kono
only recently (1959) elucidated fully the cytological situa-
tion (allotriploidy) which is the basis of the sterility.
Cruickshank (1953) discovered, in the Australasian genus
Pimelea, an interesting polyploid series ranging from diploid
to hexaploid species, some gynodioecious, others dioecious,

278 Rhodora [Vol. 64

which will have to be given serious consideration when that
group is revised. Apogamy has been reported in the taxo-
nomically troublesome genus Wikstroemia, a genus closely
related to, if not congeneric with Daphne. It is suspected in
a number of species on the basis of aborted pollen but has
been demonstrated only in W. indica C. A. Mey. (Fagerlind,
1940). Some of the difficulties in this genus may stem from
this phenomenon. There has been a recent trend toward
reporting chromosome numbers and coórdinately shifting or
erecting additional taxonomic categories to accommodate
newly uncovered chromosome-number patterns. This pro-
cedure, often accomplished with complete disregard for
other factual data and lacking overall perspective of the
taxonomic continuity of the group, while hailed in some
quarters as a “miracle drug" to "cure" taxonomy is, in
reality, a placebo.

It is difficult to obtain living material of this family,
excepting the horticulturally important species of the genus
Daphne. Observations of cytological importance can be
made only on living material, necessitating the accumulation
of this type of data whenever it is available. During the
past year I have been fortunate to obtain living material of
Daphne blagayana Freyer, an Old World species, and
Daphnopsis americana (Mill.) J. R. Johnston, a New World
species. Somatic chromosome numbers for these species are
reported here for the first time.

Daphne blagayana Freyer, 2n — 18 (Nevling 101, A).
This low, spreading, Old World species with handsome
terminal inflorescences of creamy white flowers (see Curtis
Bot. Mag. 124: t. 7579. 1898) is native to subalpine woods
of Roumania, Hungary, Yugoslavia, Albania, and Bulgaria.
It is related to several other species of horticultural interest
such as D. collina Smith, D. sericea Vahl, and D. petraea
Leybold. In Europe it is cultivated and considered to have
two "forms" (Hodgkin, 1961) one of which flowers two
weeks earlier (March in Great Britain) than the other.
During the fall of 1961, the Arnold Arboretum was fortu-
nate to receive six rooted shoots from the Arboretum Volcji
Potok, Radomlje, Yugoslavia. These shoots were collected

1962] Nevling — Thymelaeaceae 279

from wild plants which had layered naturally in a habitat
of perpetual shade on the cool, moist, north-facing slopes of
low hills.

Inasmuch as pollen mother cells were not available for
examination, root tips were selected, placed in a saturated
aquaeous solution of paradichlorobenzene for two hours,
rinsed, fixed in three parts absolute ethyl alcohol and one
part glacial acetic acid for 30 minutes; hydrolized in 1 N.
hydrochloric acid at 55° C. for 20 minutes, rinsed, stained
in leuco-basic fuchsin for two hours and squashed. In such
preparations a somatic chromosome number of 18 was
observed repeatedly. All stages in division were observable.

It may seem unusual that the chromosome number of this
well-known species has not been reported previously as there
are several papers devoted to its embryo-sac development.
These papers, however, are pre-1900, and apparently little
attention has been given this species in recent years, except
by horticulturists. The chromosome number 2n» = 18 has
been reported for nine of ten species of Daphne, the only
exception being the previously mentioned D. odora, a tri-
ploid.

Daphnopsis americana (Mill. J. R. Johnston, 2n = 18
(Nevling 102, A). This New World species of dioecious,
small trees is distinguished from other members of the
genus by the large, dichotomously branching, terminal inflor-
escences. It is the most widespread and complex of the
genus Daphnopsis with seven subspecies (Nevling, 1959).
Daphnopsis americana ssp. caribaea (Griseb.) Nevl. has the
most extensive geographical distribution of any of the sub-
species, ranging from Nicaragua south through Costa Rica
and Panama to Colombia, eastward across northern Vene-
zuela, and north through the Lesser Antilles to Puerto Rico.
Immature fruiting material of this subspecies (R. & E.
Howard 15258, A) was collected, in 1960, from an isolated
tree, on a rockslide, on the north slope of Mt. Eagle, St.
Croix. Cuttings and mature fruiting inflorescences from
this isolated tree were forwarded in February, 1961. The
pure white drupes were ovoid, about 8 mm. long and 6 mm.
in diameter with a fleshy outer layer and an inner fibrous

280 Rhodora [Vol. 64

one. The extensive fleshy layer, containing compound rhom-
boidal crystals of calcium oxalate, is lost in the drying
process of preparing herbarium specimens. The pseudo-
monomeric ovary contains a single bi-integumented ovule;
the inner integument forms a dark, hard, seed coat while
the outer integument forms an outer membranaceous layer.
All cuttings failed to strike but a half-dozen seedlings were
obtained from less than a dozen seeds, the first germinating
in two and one-half months and the last after five months.
The sex of seedling plants has not yet been determined.

Root tips from the seedlings were examined cytologically
after various methods of treatment, including that used for
Daphne blagayana. Except for the treatment outlined below,
the chromosomes were unsuitable for study due to insuffi-
cient chromosomal contraction and poor staining. Root tips
were placed in water at 10° C. for 15 hours, fixed and hydro-
lized as in Daphne blagayana, rinsed, stained and squashed
in aceto-carmine. A somatie chromosome number of 18
was observed repeatedly, with all stages of division being
observable and appearing normal. Since this is the first
chromosome count to be reported for a member of the
Thymelaeaceae from the New World it is interesting to note
that it fits the base number of nine as previously reported
in the family.

I am indebted to Ing. Igo Ora&, Director Emeritus of the
Voleji Potok Arboretum, for plants of Daphne blagayana,
and to Mr. A. J. Oakes, Research Agronomist of St. Croix,
for cuttings and fruits of Daphnopsis americana, ssp. cari-
baea. The editorial suggestions of Drs. C. E. Wood, Jr., and
W. R. Ernst, which have substantially contributed to the
organization of this note, are appreciated by the author.
Special thanks are due to Mr. Alfred Fordham, Propagator
of the Arnold Arboretum, for his sympathetic handling of
the research materials, without which this paper would
not be possible. — ARNOLD ARBORETUM, HARVARD UNIVERSITY.

LITERATURE CITED

CRUICKSHANK, R. H. 1953. Chromosome numbers in the genus
‘Pimelea? Proc. Roy. Soc. Tas. 87: 13-16, figs. 1-4.

1962] Santamour — Chromosome Number in Maples 281

DoMKE, W. 1934. Untersuchungen iiber die systematische und geo-
graphische Gliederung der Thymelaeaceen. Bibliot. Bot. 27(111):
1-151.

FAGERLIND, F. 1940. Zytologie und Gameotophyten bildung in der
Gattung Wikstroemia. Hereditas 26: 23-50.

HopGKIN, E. 1961. Daphnes. Jour. Roy. Hort. Soc. 86: 481-488.

NEVLING, L. L, JR. 1959. A revision of the genus Daphnopsis. Ann.
Missouri Bot. Gard. 46: 257-358.

Okura, E. AND M. Kono. 1959. Cytogenetical consideration of
Daphne odora Thunb. based on its karyotype. Biol. Jour. Okayama
Univ. 5: 51-56.

Osawa, I. 1913. On the development of the pollen grains and
embryo sac of Daphne, with special references to the sterility of
Daphne odora. Jour. Coll. Agric. 4: 237-264, pls. 25-27.

CHROMOSOME NUMBER IN STRIPED AND MOUNTAIN
MAPLES. — Striped maple (Acer pensylvanicum L.) and
mountain maple (Acer spicatum Lam.) are both common
species throughout the Northeast. Their ranges, which ex-
tend west to Minnesota and south in the mountains to north-
ern Georgia, are similar. Although both species may attain
small tree size, they are of little commercial importance for
timber and have attracted only mild interest as ornamentals.
Neither species has been investigated cytologically.

The writer has collected both species for chromosome
studies from several northern localities. Striped maple
collections were from Mifflin and Luzerne Counties in Penn-
sylvania, and Hampshire County, Massachusetts. Mountain
maple was collected in Hennepin County, Minnesota; Dela-
ware County, Pennsylvania; and Middlesex County, Massa-
chusetts.

Entire inflorescences were removed from the trees during
the spring, and were fixed for 24 hours in 3:1 alcohol-acetic
with a few drops of ferric chloride added. Storage was in
75 percent ethyl alcohol at 10°C. Standard aceto-carmine
smear techniques on pollen mother cells were used in the
chromosome studies.

It was found that, for both species, meiosis took place
before the bud scales had fully separated. Chromosome

282 Rhodora [Vol. 64

counts for all material gave n — 13, and no marked meiotic
irregularities were noted. Thus both species appear to be
diploid over the northern portion of their ranges.

Of the 10 species of maple native to the United States
for which chromosome numbers are now available, 8 are
diploid and only red and silver maples are polyploid. —
FRANK S. SANTAMOUR, JR. GENETICIST, NORTHEASTERN
FOREST EXPERIMENT STATION, U. S. FOREST SERVICE, STATIONED
AT MORRIS ARBORETUM, PHILADELPHIA, PA., IN COOPERATION
WITH THE UNIVERSITY OF PENNSYLVANIA.

ILLINOIS FLORA: NOTES ON LIMNOSCIADIUM, DICLIPTERA,
AND IRESINE. — During recent field work I found one of the
Umbelliferae that is new to the Illinois flora, an additional
stand of Dicliptera brachiata (Pursh) Spreng., and one of
Iresine rhizomatosa Standl., both species rare in Illinois.

Limnosciadium pinnatum (DC.) Math. & Const. is a plant
of wet places that ranges from Louisiana and Texas, north
to southeastern Kansas and southwestern Missouri. On
July 14, 1960, I spent some time collecting plants east of
Fayetteville, a small community in southeastern St. Clair
County, about 25 miles southeast of East St. Louis. One
of my collections from a roadside ditch east of the Kaskaskia
River was Limnosciadium pinnatum. This site is almost
300 miles northeast of the range in southwestern Missouri.
Herbarium specimens, in the herbarium of the Illinois
Natural History Survey (ILLS) and the University of
California (UC), have the following label data:

Roadside ditch, east of Fayetteville, St. Clair County,
Illinois, July 14, 1960, R. A. Evers 65864.

In a previous article (Rhodora 58:49-50) I reported the
occurrence of Dicliptera brachiata in southern Illinois, east
of Joppa, Massac County. That collection was made in 1951.
On subsequent trips to this locality I failed to find this
species but did discover a stand of another plant rare in

II wish to thank Dr. Lincoln Constance, University of California, Berkeley, for
identifying the specimen.

1962] Winterringer — Zizaniopsis 283

Illinois, Iresine rhizomatosa. On September 2, 1960,while
botanizing the bluffs and floodplain of the Ohio River east
of Cave-in-Rock, Hardin County, I found a sizable stand of
D. brachiata at the base of the bluff. This locality, the
second for this species in Illinois, lies over 40 miles north-
east of the Massac County site. On October 12, 1961, I
revisited the ravine east of Joppa and once more collected
D. brachiata. The two recent collections have the following
label data:

Base of wooded bluffs, east of Cave-in-Rock, Hardin
County, Illinois, September 2, 1960, R. A. Evers 67398.
Wooded ravine, east of Joppa, Massac County, Illinois, Octo-
ber 12, 1961, R. A. Evers 72015.

The herbarium sheets of Iresine rhizomatosa in the Sur-
vey herbarium have the following data:

Wooded ravine, east of Joppa, Massac County, Illinois,
September 23, 1952, R. A. Evers 35975; September 12, 1961,
R. A. Evers 71286.

This species formed a colony about one meter in diameter
and was very conspicuous along the bank of the intermittent
stream in the ravine. — ROBERT A. EVERS, ILLINOIS NATURAL
HISTORY SURVEY, URBANA. i

ZIZANIOPSIS MILIACEA FROM ILLINOIS. — The robust,
perennial, aquatic grass Zizaniopsis miliacea (Michx.) Döll
& Aschers. has not been previously reported as an estab-
lished species in Illinois according to information available
to the writer. The natural range of this southern wild rice
or water-millet is through most of the southern and south-
eastern states including Tennessee, Kentucky and Missouri.
The present locality of the Illinois station for Zizaniopsis
miliacea is in Montgomery County near Hillsboro. In this
locality the plants grow in marginal patches and isolated
clumps in shallow water around the shore of Lake Hillsboro.
The first specimens were called to the writer’s attention by
Mr. A. C. Lopinot of the Illinois Department of Conserva-
tion. Mr. Lopinot requested identification of the broad-

284 Rhodora [Vol. 64

leaved grass which he found growing with Typha latifolia
and Leersia oryzoides.

Lake Hillsboro is an artificial lake of about 100 acres
created in 1917 as a source of water supply for the town.
Mr. Paul Graden of Hillsboro revealed that the grass, Z.
miliacea, had been growing, to his knowledge, around the
lake for perhaps 20 years. Further inquiry did not reveal
how the original plants or seeds arrived or if they had been
planted; but they are certainly persisting, and continued
survival seems likely. We do not know if this grass arrived
in the locality by natural or other means, and the fact that
it has persisted and become naturalized seems worthy of
reporting. Only a few culms bore inflorescences and, grow-
ing as it does with Typha latifolia, it may have been easily
overlooked. Verification of identity was made by Dr. Jason
Swallen. Specimens are deposited in herbaria of the follow-
ing: Illinois State Museum, Springfield, Illinois Natural
History Survey, Urbana, and the U. S. National Museum,
Washington. Collection data ase are follows: Montgomery
Co., Illinois, near Hillsboro. Shallow water along the margin
of an artificial lake. Oct. 4, 1961, G. S. Winterringer, 18526,
18527 (ISM), 18515 (US), and 18516 (ILLS). — GLEN S.
WINTERRINGER, ILLINOIS STATE MUSEUM, SPRINGFIELD.

A NOTE ON THE ASCRIPTION OF ELYMUS ARENARIUS TO
ILLINOIS. — In recording the recent discovery of Elymus
arenarius in Wisconsin (Rhodora 62: 199-201, 1960), the
authors say “The taxonomic confusion surrounding Elymus
mollis and E. arenarius has been carefully and thoroughly
elucidated by Bowden (1957). All Illinois collections of E.
mollis cited by Jones (1955), who followed Hitchcock &
Chase (1950) are E. arenarius.”

There is, so far as I am aware, no taxonomic confusion
surrounding E. mollis, an American species, and the Eurasi-
an E. arenarius, although it is a fact that some specimens of
the latter species sometimes have been misidentified as E.
mollis. As early as 1918 Miss Edna Mosher had correctly

1962] Jones — Elymus Arenarius 285

named the grass as E. arenarius, and cited what is so far as
I know the first collection of it in Illinois: “Wilmette, Killip,
June 1916.” This report was repeated by Pepoon in 1927.

Of course, like practically everyone else who has written
anything on Gramineae of the United States during the last
quarter of a century, we generally “followed” Hitchcock or
Hitchcock & Chase. In this instance, however, the reference
is not particularly apt, as instead of “following” Hitchcock
& Chase we pointed out the true taxonomic identity of the
European lyme-grass that is establishing itself on sand
dunes around the south end of Lake Michigan. This identity
was abundantly proved and demonstrated by the Canadian
botanist, Dr. W. M. Bowden, two years later. — G. NEVILLE
JONES, UNIVERSITY OF ILLINOIS, URBANA.

Volume 64, No. 758, including pages 97-190, was issued June 27, 1962
Volume 64, No. 759, including pages 191-285, was issued September 28, 1962

ARLOW REFERENCE LIBRARY, 7

JAN 1 1963

Dodora

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by

ALBION REED HODGDON, Editor-in-Chief

ALBERT FREDERICK HILL .

STUART KIMBALL HARRIS |

RALPH CARLETON BEAN
Ln Edit

ROBERT CRICHTON FOSTER a es

ROLLA MILTON TRYON

RADCLIFFE BARNES PIKE

Vol. 64 October-December, 1962 No. 760
CONTENTS:
Studies in the Iridaceae, VII Robert C. Foster ......... eere 291

Chromosomal Races in Eastern North American Species of
Hedyotis (Houstonia) Walter H. Lewis and Edward E.

Terrell osccccccecseseoccesscodcsscescasdesonscdocnaccdccunedcieetssteneecceedh cocuse TT PEDES 313
A new Crucifer from the Great Slave Lake Area of Canada

Reed C, Rollina..... LLL errrreeteskeessssaeeicee meh es cO DEI 324
Two New Varieties and a New Combination in Rudbeckia

Robert E. Perdue, Jr. reru 328
A New Species of Setcreasea from Mexico Shamin A.

F'arugi, K. L. Mehra and R. P. Celarier ...0..0scnssssmnels 329
The Parasitic Behavior of Dasistoma macrophylla

Martin. A. Piehl ..cccsscecccscssarecccssarccossosccesstacsooeetoes e ECT TTE DENEN 331

(Continued on Inside Cover)

The Nem England Botanical Club, Ine.

Botanical Museum, Oxford St., Cambridge 38, Mass.

CONTENTS: — continued

Heteromorphic Pollen Grains in Polymnia T. Richard Fisher

and James R. Wells ......... eese enn 336
Changes in Flora of the Machias Seal Islands Radcliffe B.

Pike and A. R. Hodgdon ............. sse ee 340
The Ferns and Fern Allies of Chihuahua — Book Review

Rolla M. Tryon eese eene eene 347
A note on X Cyperus Weatherbianus Marcel Raymond ........ 349
An Albino Form of Vaccinium macrocarpon

Frank C. MacKeever essem eene eene 350
Juniperus horizontalis in New Hampshire

A. R. Hodgdon and F. L. Steele sss BOL
Cyperus ferruginescens in Vermont Richard J. Eaton ........ 352
An Unusual Rubus Frederick L. Steele ............... 353
Western Plants in Northern Illinois Egbert W. Fell ........ 354

Additions to the Flora of Illinois Robert H. Mohlenbrock,
James E. Ozment and George W. Folkerts ccccccccccccscccceeeees 356

Nostoc parmeliodes in Massachusetts Edgar E. Webber ........ 358

Rhodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 64 October-December, 1962 No. 760

STUDIES IN THE IRIDACEAE, VII
ROBERT C. FOSTER!

INTRODUCTION

In view of the fact that the genera Mastigostyla and Car-
denanthus have been treated piecemeal by me in three dif-
ferent publications, it seems desirable to give them unified
treatments, with complete keys, and adding two more species
to Mastigostyla. A treatment of Anomalostylus, fuller than
that given in the original publication, is included. Finally,
there are some notes on the genus Trimezia, with the descrip-
tion of two new species, some miscellaneous notes, and des-
criptions of three novelties in the genus Sisyrinchium.

Aside from specimens in the Gray Herbarium (GH),
material has been seen from the U. S. National Herbarium
(US), the New York Botanical Garden (NY), the Missouri
Botanical Garden (MBG), the Chicago Natural History
Museum (F), the University of California (UCAL), and
the Academy of Natural Sciences of Philadelphia (PHIL).
To the administrative officers of these herbaria, I am indebt-
ed for the opportunity of seeing this material.

1. MASTIGOSTYLA I. M. JOHNST.

Reasons for believing Mastigostyla distinct from Cypella
were given in detail in Contrib. Gray Herb. no. 155: 22-23
(1945). Additional material, both of Mastigostyla and
Cypella, seen since that time has strengthened the view there

!Gray Herbarium, Harvard University.

291

292 Rhodora [Vol. 64

expressed. This may be recapitulated briefly as follows:
Mastigostyla has a short but definite perianth-tube, with the
base of the flower infundibuliform; the long stamen-fila-
ments are either completely united or nearly so, while the
anthers are free from the style; the style-arms are bifid to a
point well below the stigmas; the inner tepals are greatly
reduced, far more so, proportionately, than in any species
of Cypella.

Mastigostyla I. M. Johnst. in Contrib. Gray Herb. (n. s.) no. 81:
85 (1928); Diels in Engler & Prantl, Nat. Pflanzenfam. (ed. 2) 15a:
498 (1930), as synonym of Cypella; Macbride in Field Mus. Pub. Bot.
13: 716 (1936), as synonym of Cypella; Foster in Contrib. Gray Herb.
no. 155: 22-23 (1945) and no. 161: 15 (1946).

Bulbous, caulescent or acaulescent herbs. Basal leaves few, linear;
cauline leaves 1-2. Stem simple or branched. Spathes firm-textured,
usually subequal, or the outer shorter, 1-4-flowered, the slender pedi-
cels usually not exserted at anthesis. Flowers infundibuliform at the
base, with a short perianth-tube (usually 2 mm. long) ; the two series
of tepals markedly dissimilar in size and shape, the outer larger than
the inner. Stamen-filaments united in a column, the anthers sessile,
or nearly so, on the column, opposite the style-arms. Style as long
as the column, more or less wing-margined above; style-arms bifid
below the transverse stigmas, the crests (2 per style-arm) more or
less petaloid or flagelliform, decurrent on the abaxial faces of the
style-arms. Capsule oblong-ellipsoid; seeds small, angulate.

Type-species: M. cyrtophylla I. M. Johnst.

KEY
a. Length of style, style-arms and -crests less than 15 mm.
b. Spathes immediately subtended by 1-2 cauline leaves.

c. Spathes subtended by 2 cauline leaves; style-crests 0.5 mm.
long; inner tepals oblanceolate, 8 mm. long; Argentina.............
NENRMMMMMRMRMRMMRMWRHRWRMWRMRMRMRWMWMWMWMWMWWWMWWMWWMZMWMMMMWMEMMMMMMEMMMMM 1. M. Cabrerae.

c. Spathes subtended by 1 cauline leaf; style-crests 1-2.5 mm.
long; Bolivia.

d. Style-crests 1 mm. long; inner tepals linear or narrowly

elliptic, 7 mm. long, not over 1 mm. wide. 2. M. brevicaulis.

d. Style-crests about 2.5 mm. long; inner tepals ovate, 5.5 mm.

long, 2.5 mm. wide. .......... eee 3. M. potosina.

b. Spathes not subtended by cauline leaves. ............ 4. M. Johnstonii.
a. Length of style, stvle-arms and -crests more than 15 mm.

e. Spathes subtended by cauline leaves, the plants appearing

acaulescent. ...........sseeeeeeeeee nennen I enne nennen nnns 5. M. Hopp.
e. Spathes not subtended by cauline leaves, the plants obviously
caulescent.

f. Style-crests more or less flagelliform, more than 2 mm. long.

1962] Iridaceae — Foster 293

g. Spathes ventricose; inner tepals to 1.4 em. long. oo... eee
Hooded ETT 6. M. cyrtophylla.

g. Spathes not ventricose; inner tepals shorter.
h. Outer tepals obovate-spatulate, 3 cm. long; inner tepals

oblanceolate.
i. Basal leaves to 50 cm. long, 4 mm. wide; inner tepals
Woo Mr RETIRO ONU 7. M. gracilis.
i. Basal leaves to 29 em. long, 1-2 mm. wide; inner tepals
repro ur o 4 PRENNENT Deu 8. M. Joergensenii.
h. Outer tepals oblong, to 2.2 em. long; inner tepals narrowly
BL EYES CR rt oM NEL OM e 9. M. peruviana.

f. Style-crests elliptic-lanceolate, 2 mm. long. .. 10. M. Cardenasii.

1. Mastigostyla Cabrerae R. C. Foster in Contrib. Gray Herb. no.
171: 25 (1950). Bulb ovoid, about 2.5 em. high and 1.5 em. wide, the
outer tunics thin, dark-brown. Stem for the most part subterranean,
3-7 em. long, simple. Basal leaves mostly reduced to brown sheaths,
rarely 1 produced, filiform, acute, to 10 cm. long and 0.5 mm. wide;
cauline leaves 2, subtending the spathes, the lower one 12-22 cm.
long, 2 mm. wide, linear, long-attenuate, acute, glabrous, the upper
similar but shorter, to 6 cm. long. Spathes subequal or the outer
longer, 2.5-3 cm. long, the outer one acuminate, obtuse, the inner one
suboblong, broad at the apex, obtuse, 1-2-flowered. Ovary oblong-
ellipsoid, glabrous, 7 mm. long. Flowers lilac-colored, obscurely spot-
ted; outer tepals obovate-spatulate, to 2 cm. long and 5 mm. wide,
long-unguiculate, rounded at the apex; inner tepals oblanceolate, 8
mm. long and about 1 mm. wide, the base of the claw glandular-
tumescent. Stamen-column 8 mm. long; anthers linear, 4 mm. long.
Style equal to the column in length; style-arms ca. 3 mm. long, bifid
for about 1.5 mm., the style-crests much reduced, about 0.5 mm. long.
Capsule and seeds not seen,

Known only from ARGENTINA. SALTA: San Antonio de los
Cobres, Jan. 29, 1944, A. L. Cabrera, no. 8255 (type, GH).

The great reduction of the style-crests is a notable feature
of this species. Other species with flowers of about the same
size, M. brevicaulis, for instance, have very much longer
style-crests.

2. M. brevicaulis (Baker) R. C. Foster in Contrib. Gray Herb. no.
161: 16 (1946). Nemastylis brevicaulis Baker, Handbk. Irid. 113
(1892). Bulb ovoid, to 2.5 cm. high, about 1 cm. in diameter, the
tunies brown, membranous. Basal leaves 1-2, 20-30 cm. long, 3-4
mm. wide, linear, plicate, acute, glabrous; cauline leaves 1-2, 8-30 cm.
long, 2-4 mm. wide, the uppermost usually subtending the spathes.
Stem simple, to 20 cm. tall, terete, glabrous. Spathes herbaceous, sub-
equal, to 3 em. long, the outer acute, 1-3-flowered; pedicels filiform, not
exserted at anthesis. Ovary ellipsoid-turbinate, to 5 mm. long, glab-
rous. Flowers blue; outer tepals long-unguiculate, oblanceolate, some-
what acute, to 1.8 cm. long and about 5 mm. wide; inner tepals 7 mm.

294 Rhodora [Vol. 64

long, 0.75-1 mm. wide, glandular-tumescent at the base, linear or
lance-linear, acute. Filament-column 6 mm. long; anthers 4 mm. long,
deeply retuse at the apex. Style 7 mm. long; style-arms 2 mm. long,
bifid for 1 mm. below the stigmas, the style-crests petaloid, rather
than flagellar, 1 mm. long. Immature capsule turbinate, about 8 mm.
long; seeds not seen.

Known only from BOLIVIA. La Paz: LARECAJA: vic. Sorata; colle
Ullontiji, 2700 m. alt., Jan.-Apr., 1859, Mandon 1224 in part (isotype,
NY); MURILLO: near La Paz, 3000 m. alt., Jan. 14, 1907, Buchtien no.
819 (US).

As explained when the transfer was originally made, the
type-number contains a mixture of Cypella and Mastigo-
styla. The isotype at New York agrees well with the original
description and, for that reason, the name has been limited

to the Mastigostyla element.

3. M. potosina, spec. nov. Bulbus ovoideus, 2 cm. altus, 1 cm. latus,
tunicae brunneae, tenues. Folia basalia 2 (in specimine incompleta),
2 mm. lata; folia caulina ad 10 em. longa, 2 mm. lata, acuta, glabra.
Caulis simplex, glaber, 3-6 em. altus. Spathae subaequales vel exterior
parum brevior, ad 3.5 em. longae, acutae, herbaceae, 2-fl., pedicelli
glabri, filiformes, anthesin non exserti. Ovarium oblongum, ad 6 mm.
longum, glabrum. Flores caerulei, perianthii tubus ad 1 mm. longus;
tepala exteriora longe unguiculata, obovata, 1.8 em. longa et 6 mm.
lata, apice truncato-obtusa, breve mucrcnulata; tepala interiora ovata,
5.5 mm. longa et 2.5 mm. lata, acute vel subacuta. Staminum columna
circa 9 mm. longa; antherae 4.5-b mm. longae, connectivus latus et
pertenuis. Stylus columnam longitudine aequans; styli rami et cristae
ad 1.4 em longi, 2.5-3 mm. bifidi. Capsula seminaque ignota.

BOLIVIA: Potosi: between Potosí and Camargo, in sandy, rocky
soil, 3800 m., Feb. 1949, Cárdenas no. 4265 (type, GH).

This species is most closely related to M. brevicaulis but
can be readily distinguished by longer style-crests and short-
er and broader inner tepals.

4. M. Johnstonii R. C. Foster in Contrib. Gray Herb. no. 155: 25
(1945). Bulb ovoid, 1-2.5 em. high, 6-10 mm. in diameter, the tunics
dark-brown, membranous. Basal leaves absent, or 1 rudimentary
sheath at the base of the stem; cauline leaf borne on the stem about
4-8 cm. below the inflorescence, lance-linear, attenuate at base, acute,
plicate, glabrous, at least 15 cm. long and 3-5 mm. wide. Stem simple,
terete, glabrous, 10-20 em. long. Spathes herbaceous, the outer 3.4-
4.4 cm. long, acuminate, acute, the inner 3.2-4 cm. long, rather obtuse,
retuse, 1-3-flowered, the pedicels 2.5-3.5 em. long. Ovary ellipsoid,
glabrous, to 8 mm. long. Flowers violet; outer tepals at least 1.8
cm. long and 6 mm. wide, long-unguiculate, obovate; inner tepals much
reduced, 5-6 mm. long, about 2 mm. wide, ovate-oblanceolate, acute,
the very short claw tumescent, apparently glandular. Stamen-column

1962] Iridaceae — Foster 295

6 mm. long; anthers nearly sessile on the column, 5 mm. long. Style
as long as the column; style-arms and appendages 6 mm. long, the
arms entire for 3 mm. and then bifid. Mature capsule and seeds not
seen.

Known only from ARGENTINA. TUCUMÁN: CHICLIGASTA: Estan-
cia Las Pavas, 3200 m. alt., Dec. 3, 1926, Venturi, no. 4636 (type, GH;
isotype, US); Estancia Santa Rosa, 3600 m. alt, Dec. 13, 1925,
Venturi, no. 6500 (US).

In habit, this species most closely resembles M. brevicaulis,

but by the technical details of the flower it can readily be
separated. The inner tepals are ovate to oblanceolate, rather
than linear to lance-linear, and the style-arms and style-
crests are much longer, with the crests more nearly flagelli-
form. From M. brevicaulis it differs further in that the

spathes are not immediately subtended by cauline leaves.

5. M. Hoppii R. C. Foster in Contrib. Gray Herb. no. 155: 24 (1945).
Cypella Hoppii Diels ex Macbride in Field Mus. Pub. Bot. 13:717
(1936), nomen invalidum (without Latin diagnosis). Bulb ovoid to
subglobose, 1.5-2 cm. high, 1-1.5 em. wide, the tunies dark-brown.
Basal leaf solitary, shorter than the stem, 3.5-5 cm. long, mostly
sheathing; cauline leaves 2, immediately subtending the inflorescence,
the lower to 18 em. long, spathiformly clasping at the base, the linear-
attenuate blade acute, 1-2 mm. wide, the upper leaf similar, 2.5-7 em.
long, 1-1.5 mm. wide. Stem simple, terete, glabrous, 3-7 cm. tall.
Spathes herbaceous, subventricose, the outer to 2 cm. long, acute,
the inner to 2.5 em. long, obtuse, 2-4-flowered, the pedicels shorter
than the spathes, even in fruit. Ovary oblong, glabrous, about 5-6
mm. long. Flowers violet; outer tepals to 2.5 cm. long, the blade
about 8-10 mm. wide, obovate, obtuse, unguiculate; inner tepals re-
duced, much smaller than the outer. Stamen-column to 1 cm. long,
anthers 4-5 mm. long. Style as long as the column; style-arms and
appendages about 7 mm. long. Capsule oblong-ellipsoid, rather sharply
trigonous, about 1 cm. long; seeds dark-brown, angulate, 1.5-2 mm.
long.

Known only from PERU. AREQUIPA: Arequipa, Hopp (type, Berlin,
not seen; photo GH, F); open gravelly soil above Arequipa, 2500-
2600 m. alt, Apr. 7-16, 1925, Pennell, no. 13173 (GH, NY); above
Arequipa, 2500 m. alt., Apr. 7-16, 1925, Pennell, no. 13180 (F, PHIL).

No flowering material has been seen by me. The descrip-
tion given has been drawn up from the excellent photograph
of the type and from fruiting material collected by Pennell.

6. M. cyrtophylla I. M. Johnst. in Contrib. Gray Herb. (n. s.) no.
81: 85 (1928). Cypella cyrtophylla (Y. M. Johnst.) Diels in Engler
& Prantl, Nat. Pflanzenfam. (ed. 2) 15a: 498 (1930); Macbride in
Field Mus. Pub. Bot. 13: 716 (1936). Bulb ovoid, to 3 em. high, 1.5-2
cm. wide, the membranous tunics dark-brown. Basal leaves 1-2, re-

296 Rhodora [Vol. 64

duced to brownish sheaths, or if produced, to 30 cm. long, 1 mm. wide,
linear, acute, glabrous; cauline leaves 1-2, to 30 cm. long, 2-4 mm.
wide, exceeding the inflorescence, the uppermost reduced. Stem occa-
sionally simple, but usually 1-2-branched well above the base, terete,
glabrous, to 30 cm. long. Spathes herbaceous, drying with a purplish
tinge, somewhat ventricose, subequal or the outer shorter, to 4 cm.
long, several-flowered, the filiform pedicels not exserted at anthesis.
Ovary oblong, glabrous, 4 mm. long. Flowers clear blue or violet,
sometimes with darker spots; outer tepals long-unguiculate, the blades
spreading, to 3 em. long, 1.2 em. wide, obovate, the apex obtuse;
inner tepals more or less linear-oblanceolate, to 1.4 em. long, 1.5 mm.
wide, acute. Filaments entirely united, 8 mm. long; anthers to 6 mm.
long. Style as long as or a little longer than the stamen-column; style-
arms 5 mm. long, bifid for about 3 mm. below the stigmas, the flagelli-
form crests to 8 mm. long. Capsule and seeds not seen.

Known only from PERU. AREQUIPA: southern slopes of Chachani
Mt., north of Arequipa, 2440-3660 m. alt., March, 1920, Hinkley &
Hinkley, no. 16 (type, GH); open sandy soil above Arequipa, 2500-
2600 m. alt, Apr. 7-16, 1925, Pennell, no. 13172 (GH, NY), and
Pennell, no. 13173a. (PHIL, F); alrededores de Arequipa, 2400-2500
m. alt., Mar. 31, 1949, Vargas, no. 8102 (GH); cerros de Tura, 2575-
2620 m. alt., Mar. 28, 1949, Vargas, no. 7962 (GH); Yura, 2700 m. alt.,
Mar. 8, 1914, Weberbauer, no. 6838 (GH, F, US, NY).

Although the original description of this species gave
details of capsule and seeds, these details were taken from
Pennell, no. 13173, a specimen which I now consider to
belong to M. Hoppii. The very long flagelliform style-crests
of M. cyrtophylla represent the most extreme development
of that structure in the genus.

rm

i. M. gracilis, spec. nov. Bulbus ovoideus, 2.5 cm. altus, 1.5 cm.
latus, tunieae atrobrunneae, membranaceae. Folia basalia 1-2, ad 50
cm. longa et 2-3 mm. lata, glabra, acuta, plicata; folia caulina 1-2, 4-14
em. longa, 2 mm. lata. Caulis simplex vel 1-ramosus, teres, glaber,
ad 26 em. longus. Spathae subaequales, vel exterior parum brevior,
ad 4 em, longae, exterior acuminata, acuta, herbacea, interior apice
obtusa et membranacea, 2-4-fl., pedicelli filiformes, glabri, anthesin
non exserti. Flores caerulei, perianthii tubus ca. 1.5 mm. longus;
tepala exteriora longe unguiculata, obovato-spathulata, subobtusa, ad
3 cm. longa et 8 mm, lata; tepala interiora oblanceolata, acuta, ad 10
mm. longa et 2 mm. lata. Staminum columna ad 1 em. longa; antherae
ad 6 mm. longae. Stylus columnam longitudine aequans; styli rami et
cristae ad 9-10 mm. longae, rami 4 mm. bifidi. Capsula immatura
oblongo-ovoidea, 1.3 cm. longa; semina non visa.

BOLIVIA: COCHABAMBA: MIZQUE: Chaguarani, in sandy red clay,
2700 m. alt., Mar. 31, 1939, Eyerdam, no. 25081 (type, UCAL; isotype,
F).

1962] Iridaceae — Foster 297

Most closely related, probably, to M. Joergensenii, M.
gracilis has a habit of growth which is much more open than
that of its rather stiff congener. Its basal leaves are longer
and wider, and the inner tepals longer than those of M.

Joergenseni.

8. M. Joergensenii R. C. Foster in Contrib. Gray Herb. no. 171: 26
(1950). Bulb small, ovoid, to 2 cm. high, 9-10 mm. wide, the tunics
brown, membranaceous. Basal leaves usually reduced, with 1 pro-
duced, to 28 cm. long and 1-2 mm. wide; cauline leaves 2, to 15 em.
long and 2 mm. wide, linear, acute, glabrous. Stem 1-2-branched well
above the base, terete, glabrous, to 22 cm. long. Spathes herbaceous,
subequal or the outer more or less shorter, to 4.8 cm. long, 3-4-flowered
the filiform pedicels not exserted at anthesis. Ovary oblong-ellipsoid,
to 5 mm. long, glabrous. Flowers blue or purple; outer tepals long-
unguiculate, to 3 cm. long, the blade about 8 mm. wide, narrowly
obovate or spatulate, obtuse; inner tepals to 7 mm. long, 2 mm. wide,
oblanceolate, subacute, the base glandular-tumescent. Filaments
almost entirely united, 1.2 cm. long; anthers linear, 8 mm. long. Style
about 1.4 em. long; style-arms 5 mm. long, bifid for 2 mm. below the
stigmas; style-crests about 4 mm. long, subflagelliform. Capsule and
seeds not seen.

Known only from ARGENTINA. CATAMARCA: Andalgalá [no fur-
ther data available], Jörgensen (type, US no. 921640). La RIOJA:
Chilecito: Sierra de Famatina, camino a La Mejicana, 2550 m. alt.,
Feb. 5, 1927, Parodi, no. 8012 (GH).

In its size, this species suggests M. Cardenasii, but it is
easily distinguished by its more or less flagelliform style-
crests, narrower outer tepals, and by being branched well

above the base.

9. M. peruviana R. C. Foster in Contrib. Gray Herb. no. 171: 25
(1950). Bulb ovoid, to 2 cm. high and 1.5 cm. wide, the thin smooth
tunies dark-brown or blackish-brown. Basal leaves 1-2, 8-19 cm. long
and 1-3 mm. wide, linear, acute, glabrous; cauline leaf 1, exceeding
the inflorescence, 6-12 cm. long, to 2 mm. wide. Stem simple, terete,
glabrous, 2-9 cm. long. Spathes subequal, 3 cm. long, not ventricose,
2-flowered; pedicels to 2.5 em. long. Ovary obovoid, 8 mm. long,
glabrous. Flowers blue; outer tepals about 2.2 cm. long, 6-7 mm.
wide, long-unguiculate, the blade oblong, the apex obtusely rounded;
inner tepals 8-9 mm, long, 2 mm. wide, narrowly elliptie, acute, the
claw basally tumescent. Filaments united for 9 mm., free for 1 mm.;
anthers 5 mm. long. Style 1 cm. long; style-arms 4 mm. long, bifid
for 2 mm.; style-crests flagelliform, about 6 mm. long. Capsule and
seeds not seen.

Known only from PERU. APURIMAC: ANDAHUAYLAS: quebrada 2
km. east of Andahuaylas, among grass, sand and clay, 3050 m. alt.,
Feb. 23, 1939, Stork & Horton, no. 10726 (type, F; isotype, UCAL).

298 Rhodora [Vol. 64

Since M. peruviana is obviously caulescent and the single
cauline leaf does not immediately subtend the spathes, it
can readily and quickly be distinguished from the dwarf
M. Hoppii. From M. cyrtophylla it is distinguished by its
short unbranched stem, smaller flowers, and nonventricose
spathes.

10. M. Cardenasii R. C. Foster in Contrib. Gray Herb. no. 155: 23
(1945); Foster in Contrib. Gray Herb. no. 161: 16 (1946). Bulb
ovoid, 2-2.5 em. high, 1.5 em. in diameter, the dark-brown membranous
tunies prolonged upwards in a collar around the base of the stem
and leaves. Basal leaves 1-2, to 15 cm. long, 1.5 mm. wide, linear,
acute, glabrous, finely ribbed; cauline leaf 1 (or none), the base
spathiformly sheathing the stem, equaling or exceeding the inflores-
cence. Stem simple or branched near the base, 5-30 cm. long, terete,
glabrous. Spathes herbaceous, firm-textured, to 4.5 cm. long, the
outer with the margins basally united for 6-7 mm., acuminate, acute,
the inner spathes equal to or longer than the outer, with broad hyaline
margins, acute, 2-4-flowered; pedicels filiform, shorter than the spathes
at anthesis. Ovary obovoid-ellipsoid, glabrous, about 8 mm. long.
Flowers dark-blue, with darker spots; outer tepals long-unguiculate,
obovate-spatulate, obtuse, to 3 em. long and 1.5 em. wide; inner tepals
5 mm. long, 1 mm. wide, narrowly elliptic-lanceolate, acute. Stamen-
column to 1.2 cm. long, the filaments then free for 1-2 mm.; anthers
linear, 6 mm. long. Style about as long as the stamen-column; style-
arms and style-crests 1 cm. long, bifid for 5 mm., the stigmatic pro-
jections about 1.5 mm. long, the elliptic-lanceolate petaloid crests about
2 mm. long. Immature capsule ellipsoid-obovoid, to 1 em. long; seeds
not seen.

PERU: Cuzco: Saxaihuaman, 3400 m. alt., March, 1943, Cárdenas,
no. 2337 (type, GH). BOLIVIA: COCHABAMBA: TARATA: Huayrapata,
near Anzaldo, 3900 m. alt., in open sandy loam, April, 1944, Cárdenas,
no. 2491 (US).

On the basis of present knowledge, the other species of
this genus seem to be rather narrowly endemic. Consequent-
ly, it is of interest to find this species occurring in Pert
near Cuzco and in the department of Cochabamba in Bolivia.

2. CARDENANTHUS R. C. FosTER

Although much of the material seen was originally deter-
mined as Nemastylis, Cardenanthus differs from that genus,
as well as from all but two other American bulbous genera,
in having a definite perianth-tube. Usually short in itself,
in relation to the size of the flowers, the tube is a rather
marked element of the floral morphology. From Ewurynotia

1962] Iridaceae — Foster 299

it differs not only in size but also in having divided style-
arms, not simple style-arms as is true of that genus. From
Mastigostyla, which it closely resembles in habit and vege-
tative characters, it differs in having the stigmatic area
apical or lateral downward from the apex, not transverse
at the base of style-crests. The general structure of the style
and stamens associates the genus unmistakably with Tigri-
dia and its allies.

Cardenanthus R. C. Foster in Contrib. Gray Herb. no. 155: 3 (1945)
and no. 161: 13-15 (1946).

Plants small, bulbous, subacaulescent or appearing so. Leaves few,
narrowly linear. Inflorescence subsessile, the spathes terminal, flowers
shortly pedicellate. Flowers basally infundibuliform, perianth-tube
present, short, the tepals very unequal in size and shape, the outer
much larger. Stamens opposite the style-arms, the filaments united
in a column inserted at the apex of the perianth-tube; anthers sessile
or subsessile on the column. Style filiform, three-branched, the branches
bifid, canaliculate, apically stigmatose. Capsule oblong-ellipsoid; seeds
small, numerous.

Type-species: C. boliviensis R. C. Foster.

KEY
a. Inner tepals glandular-tumescent at base or apex of claw; perianth-
tube over 3 mm. long
b. Inner tepals 6 mm. long or less, 1 mm. wide, or less.
c. Perianth-tube 4 mm. long.
d. Inner tepals oblanceolate, 1 mm. wide; style-arms 2 mm.

long, bifid for 1 mm. ................... eee 1. C. orurensis.

d. Inner tepals linear, less than 0.5 mm. wide; style-arms 3.5
mm. long, bifid ea. 2 mm. ............... 2. C. boliviensis.

c. Perianth-tube 7-8 mm. long ................. 3. C. longitubus.

b. Inner tepals more than 6 mm. long, the narrowly linear or oblan-
ceolate blade 1.5 mm. wide.
e. Staminal column ca. 1 cm. long; outer tepals 8 mm. wide, ........
mE 4. C. peruvianus.
e. Staminal column ca. 7 mm. long; outer tepals ca. 4 mm. wide.
une TERRIER. 5. C. Shepardae.
a. Inner tepals not glandular-tumescent; perianth-tube not over 2 mm.
long.
f. Inner tepals to 3.5 mm. long, widest at the base. ................ e
er T P———— ÁHORREREREUNEE 1 6. C. tunariensis.
f. Inner tepals to 8 mm. long, widest above the middle.
g. Inner tepals narrowly oblanceolate, 1 mm. wide. 7. C. Venturi.
g. Inner tepals obovate-spatulate, 3 mm. wide. ...... 8. C. Vargasii.
1. Cardenanthus orurensis R. C. Foster in Contrib. Gray Herb. no.
161: 14 (1946). Plant nearly acaulescent, the bulb ovoid, 2-2.5 cm.

300 Rhodora [Vol. 64

high, 1-2 cm. wide, the tunics brown or blackish-brown, rather thin.
Basal leaves 1-2, much reduced, 2-4 cm. long, little more than fuscous
or straw-colored sheaths, rarely 1 produced basal leaf present, to 12
em. long, 1 mm. wide; cauline leaves 2, subtending the spathes, the
lower one 13-15 em. long, 1.5-2 mm. wide, linear, acute, several-nerved,
glabrous, the upper leaf similar, 3.5-5 cm. long, 1 mm. wide. Outer
spathe to 2 cm. long, herbaceous or submembranaceous, acute, the
inner spathe to 2.4 cm. long, apically truncate, 1- (rarely 2-) flowered;
pedicels to 6 mm. long. Ovary oblong, glabrous, 5-7 mm. long. Flowers
blue or blue-purple; perianth-tube 4 mm. long; outer tepals to 1.2 cm.
long and 6 mm. wide, shortly unguiculate, obovate, apically somewhat
cbtuse; inner tepals to 4-4.5 mm. long, 1 mm. wide, narrowly elliptic
or oblanceolate, acute, the base of the claw glandular-tumescent.
Staminal column 5 mm. long; anthers linear, 3 mm. long. Style as
long as the column; style-arms 2 mm. long, bifid for 1 mm. Capsule
and seeds not seen.

Known only from BOLIVIA. ORURO: CERCADO: Hacienda Huanca-
roma, near Eucaliptus, 3000 m. alt., Feb. 19-27, 1934, Hammarlund,
no. 94 (type, NY).

A second specimen, in fruit, from which the details of
capsule and seeds have been drawn for the generic charac-
terization, may belong here or with the next species. It is
Asplund, no. 6362 (US), from Uyuni, in the Department of
Potosí, collected at 3700 m. alt., Mar. 25, 1921.

2. C. boliviensis R. C. Foster in Contrib. Gray Herb. no. 155: 4
(1945) and no. 161: 14 (1946). Plant seemingly acaulescent, but with
a 2-4 cm. underground stem, the bulb ovoid, 2 cm. high, 1 cm. wide.
Basal leaves absent; cauline leaves 2, immediately subtending the
spathes, the lower to 9 em. long, 1 mm. wide, the upper to 4 cm. long,
1 mm. wide, linear, acute, glabrous. Spathes herbaceous, the outer
1.5-2 em. long, the inner 2-2.5 cm. long, obtuse, retuse, 2-3-flowered,
the flowers subsessile. Ovary ellipsoid, glabrous, about 3 mm. long.
Flowers blue; perianth-tube to 4 mm. long; outer tepals to 1.4 cm.
long, 4-5 mm. wide, ovate, acuminate to the subobtuse apex; inner
tepals to 5 mm. long, linear, the base of the claw glandular-tumescent,
the enlarged portion to 0.75 mm. long, the blade about 0.33 mm. wide,
not acuminate, blunt at the apex. Staminal column 5 mm. long;
anthers 5 mm. long. Style about 8-9 mm. long; style-arms 3.5 mm.
long, bifid for more than half their length, the stigmatic area ciliate.
Capsule and seeds unknown.

Known only from BOLIVIA. Potosi: Potosí, 4000 m. alt., Jan. 1932,
Cárdenas, no. 124 (type, GH).

Unlike other members of the genus, in this species the

blade of the inner tepals seems to be completely linear, with
a blunt apex.

3. C. longitubus R. C. Foster in Contrib. Gray Herb. no. 171: 23

1962] Iridaceae — Foster 301

(1950). Bulb ovoid, to 1.5 em. high, 1 cm. wide, the thin tunics dark-
brown. Leaves basal, several, the outermost reduced to ventricose
cataphylls, the inner 3-4 long-sheathing, the narrowly linear, strongly
recurved, glabrous, acute blades to 10 cm. long, 1 mm. wide (or less),
the midrib and edges thickened. Stem mainly subterranean, simple,
terete, glabrous, to 3 em. long. Spathes equal or unequal, the outer
to 2.2 em. long, abruptly acuminate, the inner nearly as long, or some-
what shorter, obtuse, 1-2-flowered, pedicel filiform, glabrous, to 6 mm.
long. Ovary narrowly ellipsoid, 6 mm. long, glabrous, trigonous.
Flowers light purple, sometimes with darker streaks; perianth-tube
to 7-8 mm. long, narrowly cylindrical, abruptly ampliate at the apex;
outer tepals long-clawed, obovate, to 1.5 cm. long, the somewhat acute
blade to 6 mm. wide; inner tepals linear, acute, to 6 mm. long,
slightly less than 1 mm. wide, with a tumescent patch about 1.5 m
above the base. Staminal column 5 mm. long; anthers linear, 4-5 m
long. Style 1.4 cm. long; style-arms 1 mm, long, bifid for 0.5 m
Mature capsule and seeds not seen.

Known only from BOLIVIA. La PAZ: PACAJES: Rosario, 4100 m.
alt., Jan. 13, 1921, Mrs. R. B. Shepard, no. 235 (type, GH; isotype,
US).

Growing in wet sandy soil, this species can be distin-
guished from its congeners by the longest perianth-tube and
the longest style in the genus.

4. C. peruvianus R. C. Foster in Contrib. Gray Herb. no. 171: 24
(1950). Bulb ovoid, to 2 cm. high, 1.5 em. wide, the thin tunics dark-
brown. Basal leaves 1-2, to 28 cm. long and 0.5-1.5 mm. wide, plicate,
glabrous, very acute; cauline leaves 1-2, to 25 cm. long, 1.75 mm. wide,
immediately subtending the spathes. Stem simple, mostly subterran-
ean, 2.5-4 em. long. Spathes about equal, or the outer somewhat
longer, to 3 em. long, obtuse, 1-2-flowered, the glabrous pedicels to 7
mm. long. Ovary truncately ellipsoid, glabrous, to 7 mm. long. Flowers
pale violet with darker spots; perianth-tube 4 mm. long; outer tepals
to 1.8 em. long, 8 mm. wide, obovate-spatulate, somewhat obtuse; inner
tepals to 8 mm. long, 1.5 mm. wide, oblanceolate, acute, the claw short
and glandular-tumescent. Staminal column to 10 mm. long; anthers
3.3.5 mm. long. Style to 1.1 cm. long; style-arms 2.5 mm. long, bifid
for 1.5 mm., wide, conduplicate. Capsule and seeds unknown.

Known only from PERU. TACNA: TARATA: Candarave, 2900-3000
m. alt., Mar. 11-13, 1925, Weberbauer, no. 7387 (type, F).

Growing in open shrubbery, this close relative of C. Shep-
ardae can be distinguished from it by the longer style, longer
staminal column, less bifid style-arms, and broader outer
tepals.

5. C. Shepardae R. C. Foster in Contrib. Gray Herb. no. 155: 5

(1945). Bulb ovoid, 1.5 cm. high, 0.5-1 cm. wide. Basal leaves 1-2,
to 4 em. long, 1 mm. wide, glabrous; cauline leaf single, subtending

m.
m.
m.

302 Rhodora [Vol. 64

the spathes, itself somewhat spathiform, to 2.7 em. long, 1 mm. wide,
subfaleate. Stem mostly underground, 1-1.5 em. long. Spathes her-
baceous, the outer to 2.2 cm. long, acuminate, acute, the inner to 2.5
cm. long, tapering to a blunt retuse apex, 2-flowered, the pedicels about
4 mm. long at anthesis. Ovary about 5 mm. long, ellipsoid, glabrous.
Flowers light purple; perianth-tube to 4 mm. long; outer tepals ca.
1.5 em. long, ea. 4 mm. wide, oblanceolate-spatulate; inner tepals ca.
1 em. long,1.5 mm, wide, linear or narrowly oblanceolate, with an oval
glandular patch on the claw 4 mm. above the base. Staminal column
ca. 7 mm. long; anthers 4 mm. long. Style as long as the column;
style-arms 2.3-4 mm. long, bifid nearly to the base. Capsule and seeds
unknown.

Known only from eastern PERU. PUNO: HUANCANE: Umuchi, in
sandy places, 3125 m. alt., Dec. 10, 1919, Mrs. R. S. Shepard, no. 99
(type, GH).

Through an unfortunate error, presumably in typing the
manuscript, the original description stated that the length
of the staminal column was 1.1 cm. ; this has been corrected
in the description given above.

6. C. tunariensis R. C. Foster in Contrib. Gray Herb. no. 155: 5
(1945) and in no. 161: 15 (1946). Nemastylis nana sensu Rusby, non
S. Watson, in Mem. Torr. Bot. Club, 6(1) : 125 (1896), in part. Plant
subacaulescent, the ovoid bulb to 2 em. high, about 1 em. wide. Basal
leaf solitary, 7-12 cm. long, 1 mm. wide; cauline leaf solitary, sub-
tending the spathes, 4-8 cm. long, 1 mm. wide, acute, glabrous.
Spathes herbaceous, the outer to 3 cm. long, acute, acuminate, the
inner 1.7-2.6 cm. long, subobtuse, 2-flowered, the pedicels to 1.3 em.
long. Ovary ellipsoid, glabrous, 4 mm, long. Flowers blue or purple;
perianth-tube to 2 mm. long; outer tepals 1 em. long, the blade 4 mm.
wide, oblanceolate; inner tepals to 3.5 mm. long, 1 mm. wide at the
base, narrowly lance-deltoid, acuminate, acute, eglandular. Staminal
column 5 mm. long; anthers 3-3.5 mm. long, nearly sessile on the
column. Style as long as the column; style-arms 1.5 mm. long, bifid
almost to the base. Capsule and seeds unknown.

Known only from BOLIVIA. CocHABAMBA: CERCADO: near snow-
line, Mt. Tunari, 1891, Bang, no. 1042 (type, GH; co-types US, MBG).

Although the available material of this species is scanty
and poor, it seems established that the inner tepals lack the
glandular-tumescent area and are broadest at the base.

7. C. Venturii R, C. Foster in Contrib. Gray Herb. no. 155: 6 (1945).
Bulb ovoid, 1.5-2 cm. high, 1-1.5 cm. wide. Basal leaves 1-2, to 10 em.
long, 1 mm. wide, acute, glabrous; cauline leaf solitary, subtending the
spathes, to 6.5 cm. long, 1 mm. wide. Stem mostly underground, 1-2
em. long. Spathes herbaceous, to 2.5 cm. long, subequal or the inner
somewhat shorter, the outer acute, the inner obtuse, 2-flowered, the
pedicels to 5 mm. long at anthesis. Ovary ellipsoid, glabrous, about

1962] Iridaceae — Foster 303

4 mm. long. Flowers blue; perianth-tube 2 mm. long; outer tepals to
1.5 em. long, 3 mm. wide, oblanceolate-spatulate, subobtuse; inner te-
pals 8 mm. long, 1 mm. wide, narrowly oblanceolate, acute, not glandu-
lar-tumescent. Staminal column 7 mm. long; anthers sessile on the
column, 2.5 mm. long. Style as long as the column; style-arms 1.5 mm.
long, bifid for 0.5 mm. Capsule and seeds unknown.

Known only from northern ARGENTINA. Jusuy: HUMAHUACA:
Cerro La Soledad, 3500 m. alt., Jan. 25, 1928, Venturi, no. 9025 (type,
GH; isotype, US). TUCUMÁN: CHICLIGASTA: Estancia Santa Rosa,
4600 m. alt., Dec. 13, 1925, Venturi, no. 4065 (US).

Since the original publication of this species, another col-
lection has been seen, which strengthens my feeling of its

distinctness.

8. C. Vargasii R. C. Foster in Contrib. Gray Herb. no. 171: 24
(1950). Bulb small, more or less globose, about 1 cm. in diameter, the
brown, thin membranous coats prolonged upward in a collar. Basal
leaves several, basally long-sheathing, the sheaths reaching nearly to
the middle of the spathes, the blades sharply reflexed outward above
the sheaths, to 6 cm. long, 2 mm. wide, plicate, lance-linear, acute,
glabrous. Stem mostly subterranean, simple, to 1.5 cm. long, cauline
leaves apparently absent. Spathes subequal, to 1.8 cm. long, acute,
or the inner subobtuse, 1- (rarely 2-) flowered, the flowers shortly
pedicellate, the pedicels only a few mm. long. Ovary oblong-ellipsoid,
glabrous, to 5 mm. long. Flowers white, with violet lines; perianth-
tube to 1.5-2 mm. long; outer tepals spatulate, to 1.2 cm. long, the
obtuse, retuse blade 6 mm, wide, the apical indentation crenulate; inner
tepals unguiculate, the claw 2 mm. long, not glandular-tumescent, the
obovate-spatulate blade 6 mm. long, 3 mm. wide, obtuse. Filaments
completely united, the tube 4 mm. long; anthers 2.3 mm. long. Style
6 mm. long; style-arms 1.5 mm. long, bifid almost to the base, broad,
conduplieate, with an internal central ridge and a minute tubercle at
each secondary sinus. Capsule and seeds not seen.

Known only from PERU. PuNo: CARABAYA; pampa de Lacka,
Macusani, 4360 m. alt., Feb. 15, 1948, Vargas (leg. E. P.), no. 7135
(type, GH).

The broad inner tepals and the lack of cauline leaves serve
to distinguish this species from its relatives. The most note-
worthy difference, aside from the color of the flower, is the
presence of small tubercles at the secondary sinuses of the
style-arms. These have not been noted in other members of

the genus.
3. ANOMALOSTYLUS R. C. FOSTER

Until recently, the species treated here as belonging to
Anomalostylus have been considered to belong to Cypella.

304 Rhodora [Vol. 64

The principal difference between the two genera is the
strange development of the style in Anomalostylus. Not
only are the style-arms suppressed, as is rarely true in
Cypella, but the style-crests are fused in such a manner that
three apical lobes, alternate with the stamens, are produced.
Actually, however, the stamens are opposite the transverse
stigmatic areas, so that there can be no question of the rela-
tionship of Anomalostylus and Cypella.

Anomalostylus R. C. Foster in Contrib. Gray Herb. no. 165: 110
(1947).

Bulbous, perennial, caulescent herbs, the bulbs probably rather
large, Basal leaves several, cauline leaves few. Inflorescences terminal
on the main axis and branches, the flowers long-pedicellate in herba-
ceous spathes. Flowers crateriform, the tepals basally connate, but
a perianth-tube absent, the two series of tepals dissimilar in size and
shape. Filaments free or slightly united at the base; anthers apically
and adaxially adherent to the style-wings below the stigmatic areas.
Style long-cyathiform, strongly trialate, style-arms completely obso-
lete, the three sides of the style produced above the stigmas as low,
semi-orbicular, or truncately deltoid, retuse lobes, these alternate with
the stamens; stigmatic tongues two above each stamen, transverse,
small, faleate, fimbriate-papillate, the stigmatic area apparently con-
tinued along the inner edge of each lobe to the retuse apex. Capsule
and seeds unknown.

Type-species: A. crateriformis R. C. Foster.

KEY
a. Leaves and spathes densely pruinose. ........... ees 1. A. grandis.
a. Leaves and spathes not pruinose.

b. Basal leaves to 80 cm. long and 1.8 cm. wide; pedicels shorter
than inner spathe at anthesis; filaments 5-6 mm. long. ................
:NEMRMMRRMMMMNMMMWMMWMWMMWMWMWWMMKMKKKAHMWWWWWWNNNNNNNN 2. A. erateriformis.

b. Basal leaves to 40 em. long and 4-5 mm. wide; pedicels equaling
the inner spathe at anthesis; filaments 3-4 mm. long ................

1. Anomalostylus grandis (Kranzl.), comb. nov. Cypella grandis

Kranzl. in Fedde, Rep. Spec. Nov. 14: 295 (1916). Bulb large, ovoid,
7-8 em. high and 4 em. wide, the tunics verrucose (fide Kranzlin).
Basal leaves about 4, linear-attenuate, acute, stiff, with numerous fine
veins, pruinose, to 60 em. (80 em. fide Kranzlin) long, to 1.8 cm. wide;
cauline leaves 3-4, appressed against the stem, overlapping or subim-
bricate, pruinose, acute, thick, very firm-textured, the lowest to 20 em.
long (45 em. fide Kránzlin), two-thirds sheathing, the up-
per leaves reduced to more or less sheathing, spathiform, cari-
nate bracts. Stem terete, glabrous, branched, nearly 70 cm. tall.
Spathes pruinose, nearly equal, the outer a few millimeters shorter

1962] Iridaceae — Foster 305

than the inner, to 6 cm. long (8 em. fide Kranzlin), broad, rather
abruptly acute, carinate, rigid, finely veined, the inner similar but
truncate at the somewhat membranous apex, 2-3-flowered; pedicels
stcutish, glabrous, well-exserted from the spathes at anthesis. Ovary
oblong-clavate, to 2 cm. long, glabrous. Flowers yellow (fide Krünz-
lin); outer tepals oblong-obovate, without marked differentiation be-
tween blade and claw, the inner face of the lower portion rather
sparsely glandular-pubescent, about 4 cm. long and 1.8 cm. wide
(6 cm. x 2.5 cm., fide Kranzlin), the rather blunt apex somewhat
penicillate; inner tepals about 2 cm. long and 1 em. wide (3 cm. x 8
mm., fide Kranzlin), with a well-marked, long, spatulate-naviculate
claw, the margins more or less densely glandular-cilate, somewhat
erect or spreading, but inflexed and with a broad pubescent arc at
that point, the oblong-ovate blade strongly deflexed, not glandular-
ciliate, with a broad pubescent patch in the center. Filaments very
slightiy united at the base, 4 mm. long; anthers linear, 8-9 mm. long,
adnate to the style about 2 mm. below the sinuses. Style 1.5 cm. long,
filiform for about 4 mm., then dilating and becoming oblong-cyathi-
form upward, the three apical lobes rounded, about 5 mm. broad at
the base and 2 mm. high.

Known only from PARAGUAY. Sierra de Amambay, January, Hass-
ler (coll. Rojas) no. 10,001 (type, not seen; photo, GH, F); in the
neighborhood of the Río Alto Paraná, 1909/10, Fiebrig, no. 6273 (GH,
US); Cordillera de Altos, Oct. 8, 1902, Fiebrig, no. 224 (F).

This stiff erect plant with firm-textured leaves is easily
distinguished from the other two species by the heavily
glaucous or pruinose leaves and spathes, by the sheathing
cauline leaves, which conceal most of the upper portion of
the stem, and by the very thick texture of the flowers.

2. A. crateriformis R. C. Foster in Contrib. Gray Herb. no. 165:
111 (1947). Bulb incompletely known, probably large. Basal leaves
several, to 80 cm. long and 1.8 cm. wide, linear, acute, glabrous, not
prvinose, striate with numerous fine veins, the central vein distinctly
more prominent than the rest; cauline leaves 2-4, the lowermost 12-42
cm. long, basally sheathing, the upper ones successively reduced, but
not concealing the stem. Stem 1-3-branched, terete, glabrous. Spathes
herbaceous, rather naviculate, the outer striate, carinate, abruptly
acute, 3.5-4.5 em. long, the inner slightly longer, truncate, several-
flowered; pedicels shorter than the inner spathe at anthesis, rather
slender, glabrous, ultimately elongating. Flowers apparently yellow;
outer tepals obovate, to 3.5 em. long and 2.2 em, wide, without well-
marked differentiation between blade and claw, the basal portion
with 8-9 ridges and very sparse puberulence on the inner surface, the
apex of the blade retuse with a short, blunt, penicillate apiculus;
inner tepals to 2.5 cm. long and 1 em. wide, the spatulate-naviculate
claw flushed and streaked with blue or purple, much darker than the
outer tepals (at least in dried material), densely glandular-ciliate,

306 Rhodora [Vol. 64

the iuner face glandular-pubescent, erect or somewhat spreading,
somewhat inflexed at the apex, the strongly reflexed blade yellow, with
a broad U-shaped pubescent patch and 2 round or oval pubescent
patehes in the center, these sometimes confluent, the apex rather blunt
and penicillate. Filaments united at the base for 1 mm., 5-6 mm.
long; anthers 7-11 mm. long, adherent to the style 1-2 mm, below the
sinuses. Style enlarged upward almost from the base, 1.4-1.5 em.
long, the low, rounded or subdeltoid, apieal lobes 3-4 mm. wide at
the base and 1.5 mm, high.

BRAZIL: PARANA: Serrinha, ad marg. viae ferreae, Oct. 14, 1909,
Dusén, no. 8705 (type, US; isotype, NY); MINAS GERAIS: 1865-66,
Regnell, III, no. 1214 (US); Arceira, Feb. 24, 1951, A. Macedo, no.
3212 (US); Rio GRANDE DO SUL: Estancia L. Gomez (Neu-Württem-
berg), 500 m. alt., Oct. 18, 1904, Bornmüller, no. 302 (GH). PARA-
GUAY: in the campo Estancia Primera, January, 1932, Jorgensen, no.
4537 in part (as to NY, PHIL).

As was pointed out in the original description, this plant
is almost certainly the one figured by Larranaga, Escritos,
Atlas, 1: t. XII (1927). It has been sufficiently distinguished
from A. grandis, and from A. coriifolius it may be separated
by its larger size, thicker leaves, pedicels shorter than the
spathes at anthesis, longer filaments, and by the much thick-
er texture of the tepals and style.

3. A. coriifolius (Baker), comb. nov. Cypella coriifolia Baker in
Bull. Herb. Boiss. (ser. 2) 3: 1103 (1903). Bulb incompletely known
but apparently large. Basal leaves several, to 40 em. long, 4-5 mm.
wide, linear, acute, glabrous, not pruinose, rather thick and rigid,
densely and finely nerved, the mid-nerves hardly more prominent than
the rest; cauline leaves 2-3, the lower-most partly sheathing, 7-20
cm. long, 3-7 mm. wide, the upper ones mostly sheathing, spathiform,
but not concealing the upper portion of the stem. Stem 2-3-branched,
terete, glabrous, to 66 em. tall. Spathes herbaceous, finely veined,
rather stiff, unequal, the outer to 4 cm. long, long-acute, the inner
somewhat longer, truncate at the apex, several-flowered; pedicels
slender, equaling the inner spathe at anthesis, glabrous. Ovary
oblong-ellipsoid, to 7 mm. long, glabrous. Flower pale yellow, tinged or
spotted with blue at the base within; outer tepals not well-differen-
tiated into claw and blade, obovate-spatulate, the apex retuse with a
very short penicillate mucro, to 3.2 cm. long and 1.5 em. wide, the
basal portion sparsely glandular-puberulent and about 8-ridged on the
inner face; inner tepals somewhat shorter, the margin, except the
upper half of the blade, glandular-ciliate, the claw narrow at the
base, then widening considerably, becoming spatulate-naviculate, the
inner face with about 5 ridges and sparsely puberulent, erect or
somewhat spreading but inflexed at the apex, with a dense, broadly
U-shaped, pubescent blotch at the base of the strongly reflexed,

1962] Iridaceae — Foster 307

broadly ovate (8-9 mm. wide) blade, with a semi-lunate patch in the
center of the blade, the apex rather blunt and shortly penicillate.
Filaments slightly united at the base, 3-4 mm. long; anthers 6-7 mm.
long, adnate to the style about 2 mm. below the sinuses. Style 1.2 cm.
long, filiform at the base but soon becoming cyathiform, the 3 apical
lobes semiorbicular or truncately deltoid, about 2.5 mm. wide at the
base and 1.5 mm. high.

Known only from PARAGUAY. In the campo Hiaty, January, 1930,
Jorgensen, no. 4532 (US, NY, MBG, F, PHIL); in the campo Tapytu,
March, 1931, Jorgensen, no. 4537 in part (as to US, MBG).

This species is admittedly close to the preceding, but
seems distinct. To the points of differentiation mentioned
in the discussion of A. crateriformis, the increased glandu-
lar-ciliation of the inner tepals of A. coriifolius may be

added.

4. NOTES ON TRIMEZIA

For a number of years, the binomial Tritonia riparia Cor-
demoy, Fl. lle Réunion, 161 (1895), has seemed to me to be
a “spook.” It seemed highly unlikely that a true Tritonia
should be found native on the island of Réunion. Dr. E. P.
Phillips, Gen. S. Afr. Fl. Pls. (ed. 2) 218 (1951), says of the
generic distribution: “between 40 and 50 species in South
Africa, found mostly in the southwestern districts of the
Cape Province, but extending from Namaqualand through
the coastal area into Natal; found also in the Transvaal.”
But that is a far cry from Réunion. There is in the Gray
Herbarium a sheet from Réunion, collected by G. de L’Isle,
no. 562, from one of the localities cited in the original publi-
cation of the species, “Ravine séche, St. Benoit." The plant
seemed strangely familiar in appearance and dissection soon
showed that it is the widespread and common Caribbean
irid, Trimezia martinicensis (Jacq.) Herb. As a result, the
binomial Tritonia riparia becomes an additional synonym of
Trimezia martinicensis, a name already well-burdened with
synonyms.

Within the past few years I have had more than one
inquiry as to the correct spelling of the generic name Tri-
mezia, my correspondents pointing out that the original
spelling was Trimeza. This is correct. The generic name
was spelled Trimeza when it was published by Salisbury in

308 Rhodora [Vol. 64

Trans. Hort. Soc. 1: 308 (1812). However, this name was
published as a nomen nudum and must, therefore, be disre-
garded. The first valid use of the name known to me is
Herbert’s in Bot. Reg. 30: Misc. p. 88 (1844), where he
used the spelling Trimezia. Since Herbert gave a generic
character, there can be no question of the validity and legiti-
macy of the publication. Therefore, Herbert’s spelling, Tri-
mezia, rather than Salisbury’s spelling, Trimeza, must be
used.

Again in response to inquiry, it should be noted that the
binomial Trimezia martinicensis (Jack.) Herb. was made
in the description of T. meridensis Herb. in Bot. Reg. 30:
Misc. p. 88 (1844), where Herbert wrote “Caetera ut in T.
Martinieensi." In the brief English discussion which fol-
lowed, he referred to “This plant, handsomer than the old
species, long called Iris Martinicensis. . . ." Consequently,
there can be no doubt concerning the basionym of T. mar-
tinicensis. Since the two binomials, T. martinicensis and T.
meridensis, were thus published simultaneously, as well as
simultaneously with the validation of the generic name, it
becomes necessary to select one of them as the type-species
of the genus. Since T. martinicensis is based on the much
older and well-understood name of Jacquin, I am designating
it as the type-species. The second species, T. meridensis, is
not well-understood (indeed, there is considerable question
as to what it is, as well as to whether it represents merely a
local variant of T. martinicensis or is a distinct species) , and
it seems to me ill-advised to select a doubtful concept as the
type of the genus.

Trimezia lutea (Klatt), comb. nov. Cypella lutea Klatt in Mart.
Fl. Bras. 3(1): 522 (1871). Marica bulbosa Klatt in Abh. Naturf.
Ges. Halle, 16: 374 (1882). Neomarica bulbosa (Klatt) Sprague in
Kew Bull. 1928: 281 (1928).

It is obvious from an excellent photograph of Martius, no.
437, from Minas Gerais, Brazil, which may be chosen as the
lectotype, that Cypella lutea Klatt is unquestionably a Tri-
mezia. It differs from T. martinicensis in having the single
cauline leaf close to the apex of the stem, with the inflores-
cence appearing pseudolateral in some instances. The root-
stock, however, is completely characteristic of Trimezia,

1962] Iridaceae — Foster 309

resembling a small bulb or corm, with the outer tunics split
into parallel fibers. No other genus of New World irids
possesses such a rootstock, and for that reason Cypella lutea
Klatt is here transferred to Trimezia. The name Marica
bulbosa was merely a renaming by Klatt when he trans-
ferred the plant to Marica, because of the existence of
Marica lutea Herb. But Herbert's name was apparently a
nomen provisorium and must be disregarded. In any case,
it could not bar the use of the epithet lutea in Trimezia.

BRAZIL: MINAS GERAIS: Ouro Preto, Martius, no. 437 (lectotype;
photo GH). COLOMBIA: Meta: Rancho Menegua, 100 km. SE of
Villavicencio, Helen Schieffer, no. 816 (GH).

Trimezia Martii (Baker), comb. nov. Marica Martii Baker,
Handbk. Irid. 63 (1892). Neomarica Martii (Baker) Sprague in Kew
Bull. 1928: 281 (1928).

From the description of the rootstock, it seems clear that
Marica Martii is a Trimezia. Since it seems distinct from
other species of the martinicensis-complex, I am retaining
it as a separate species, at least for the present.

Trimezia fistulosa, spec. nov. Perennis, breve rhizomatosa, rhi-
zoma ad 3-4 em. crassum, foliis vetustioribus vestitum, haec deinde
in fibris parallelibus. Folia basalia plura, perfistulosa et tubulosa, per-
recurvata, ad 20 cm. longa et 6-8 mm. in diametro, glabra. Scapus ad
15 em. altus, glaber, folia caulina absentia. Spathae plures, terminales,
spatha infima ad 2 cm. longa, acuta, spatha suprema ad 3.5 cm. longa,
obtusa, apice hyalina et subcrenulata. Pedicelli 3-4, spathas anthesin
aequantes. Ovarium glabrum, oblongum, ad 8 mm. longum. Flores
fructus seminaque ignota. BRAZIL: MINAS GERAIS: Serra do Cipó,
1800 m., Aug. 5, 1936, W. A. Archer, no. 3675 (type in U. S. Nat.
Herb.; isotype in herb. U. S. Nat. Arb.) ; Serra do Cipó, Santa Luzia,
km. 118, Aug. 23, 1933, Mello Barreto, no. 3040 (F); Diamantina,
Glaziou (photo, GH).

To describe a new irid without flowers, fruit or seeds may
appear somewhat rash. Yet the rootstock is that of a Tri-
mezia (in the Lansburgia group), and the leaves are unique
in American irids. Thick, firm-textured, strongly recurved
and tubular-fistulose, they leave no alternative to describing
the plant as a new species. According to a handwritten note
on the Glaziou specimen, the flowers are yellow, which
accords with the attribution to Trimezia. When flowers are

available for study, it is possible that this plant may prove

310 Rhodora [Vol. 64

to represent an undescribed genus. For the time being,
however, I prefer to leave it in Trimezia.

Trimezia Steyermarkii, spec. nov. A T. martinicensi foliis longis
et latis, caule ramoso, et rhizomate crasso magnoque differt. Rhizome
to 4 em. wide and 4 cm. long. Leaves ensiform, thin-textured, the
midrib prominent, to 60 cm. long and 2 cm. wide, occasionally a little
wider. Spathes obviously terminal and not pseudolateral, several-
flowered, herbaceous, thin-textured, unequal, the outer spathe about
2 em. long, the inner to 3 cm. long, or a little longer. Pedicels
glabrous, equaling the spathes at anthesis. Ovary glabrous, more or
less clavate, less than 1 em. long. Flowers yellow with purple or
brown-purple bands, probably not over 3 cm. in diameter. Style and
stamens those of Trimezia, Fruit and seeds unknown. GUATEMALA:
ALTA VERAPAZ: Pansamala, 4100 ft., J. D. Smith, Pl. Guat. no. 1786
(US); along Quiché highway about 12 km. west of San Cristóbal, 1100
m., in dense mixed forest, Standley, no. 89752 (F, GH); wooded slope
near Río Ievolay, near Hacienda Yaxcabnal, 5 mi. NW. of Cubilgüitz,
250-300 m., Steyermark, no. 44690 (F, GH) ; between Hacienda Yax-
cabnal alonga Río Ievolay and Rio Apia, 6-8 mi. NW of Cubilgüitz,
210-250 m., Steyermark, no. 45011 (F, GH); vicinity of Rio Dolores,
near Dolores, 1-2 mi. NE. of Cubilgüitz, 290-300 m., Steyermark, no.
44827 (F, GH); Coban, 1500 m., von Tuerckheim in J. D. Smith, Pl.
Guat. no. 96 (US); Coban, 1400 m., von Tuerckheim II, no. 760 (US);
von Tuerckheim (US no. 933767) ; Huehuetenango: cafetal of Finca
Soledad, 5 mi. SE. of Barillas, Sierra de los Cuchumatanes, 1150 m.,
Steyermark, no. 49539 (type, GH; isotype, F).

No satisfactory material is available at the moment for
dissection, and so floral details cannot be given. Bud-dissec-
tion shows, however, that the plant is a Trimezia. Most of
the material cited was originally determined as Neomarica.

In addition to the Guatemalan material cited, there are
two additional collections from southern Mexico which prob-

ably belong here.

Oaxaca: Lalana, 1000 m., Chinantla, Teotalcingo, Galeotti no. 5380
(US). CmiaPAS: Chicharras, 1000-2000 m., E. W. Nelson, no. 3756
(US).

5. MISCELLANEOUS NOTES AND NOVELTIES

In the treatment of the family Iridaceae in Macbride's
Flora of Peru (Field Mus. Pub. Bot. 13(1) : 707-717 [1936]),
the genus Eleutherine is not included. Consequently, it was
a matter of great interest to discover a sheet of Eleutherine
bulbosa (Mill. Urban in a small but important suite of

1962] Iridaceae — Foster Bil

specimens sent to the Gray Herbarium by Dr. César Vargas
C. The data for the specimen are as follows: PERU: Cuzco:
CONVENCION: Quillabamba, Granja de Misiones, 1040 m.
alt., enero-marzo, 1947, Vargas, no. 6347.

Through the kindness of Dr. Vargas, an isotype of Cypella
Goodspeediana Vargas (Vargas, no. 2509) has been pre-
sented to the Gray Herbarium. Its appearance is not typical
of Cypella and a bud-dissection showed unmistakably that
it is a Cipura. The following transfer is thus necessary.

Cipura Goodspeediana (Vargas), comb. nov. Cypella Goodspeediana
Vargas in Revist. Univ. Cuzco, 33 (no. 87) : 171 (1945).

During determinative work, over a period of years, three
apparently undescribed species of Sisyrinchium have been
noted. Their descriptions follow.

Sisyrinchium bromelioides, spec. nov. Herba perennis, rhizomatosa,
rhizoma subcrassum, breve. Folia basalia ad 110 cm. longa et 2 cm.
lata, minute denseque papillosa, marginibus tenuibus; folia caulina
nulla. Caulis ad 60 cm. altus (ad 2 m. fide Rambo), ancipitus, bractea
terminzlis ad 8-9 cm. longa. Inflorescentia terminalis non pseudolat-
eralis, ad 30 em. alta, multiramosa, bracteis acutis, firmis obtecta,
bracteae usque ad 2 cm. longae; spathae firmae, acutae, ad 1.5 cm.
longae, raro longiores, pluriflorae. Pedicelli tenues, spathas anthesin
subexcedentes, glabri. Ovarium turbinatum, glabrum, ad 2 cm.
longum. Perianthium flavum, atrolineatum; tepala subaequalia, ellip-
tica, acuta, ad 9 mm. longa et 2.5 mm. lata. Filamenta ad 3 mm.
longa, 1 mm. basi coalita; antherae lineares, 2.5 mm. longae. Stylus
1.5 mm. longus, styli rami 3 mm. longi. Capsula subglobosa, ad 6 mm.
alta; semina complanato-globosa, ad 2.5 mm. lata, brunneo-nigra,
sublucentia, irregulariter foveolata. BRAZIL: Rio GRANDE DO SUL:
S. Leopoldo, in wet or marshy places, 17 Dec. 1948, Rambo, no. 38856
(GH); S. Leopoldo, Nov. 1941, Leite, no. 2032 (type, Arnold Arbore-
tum); Vila Oliva, pr. Caxias, 2 Jan. 1946, Rambo, no. 33939 (GH).

The closest relatives of this new species are S. macroce-
phalum R. Grah. and S. Wettsteinii Hand.-Mazz. From both
it differs in its larger size, smaller flowers, and, above all,
in the inflorescence, which is very large, open, diffusely
branched, with bracts covering most of the branches. The
spathe-clusters are extremely numerous, appressed against
the branches when young and becoming more divergent in
maturity. It is probably the tallest species in the genus,
since Father Rambo noted that the flowering stem may reach
2 meters in height. The specific epithet was chosen to show
the striking resemblance to some bromeliad inflorescences.

312 Rhodora [Vol. 64

Sisyrinchium deflexum, spec. nov. Herba perennis, tota glabra,
rhizomatosa, rhizoma tenue, breve. Folia basalia nulla; folia caulina
ad bracteas acutas, spathiformes, convolutas reducta, ad 2.5 cm.
longa. Caulis simplex, ad 3.5 dm. altus. Spathae ad bracteas caulinas
persimiles, ad 2.5 cm. longae, acutae, 1-florae. Pedicelli spathas anthe-
sin excedentes, in fructu multo elongati et perdeflexi. Ovarium ad 2
mm. longum, turbinatum vel oblongum. Perianthium pallide flavum,
non lineatum; tepala subaequalia, elliptica, subacuta, ad 8 mm. longa
et 2 mm. lata. Filamenta circa 2.5 mm. longa, ad basin 1 mm. coalita;
antherae lineares, 2-2.5 mm. longae. Stylus ad 2 mm. longus, styli
rami circa 2 mm. longi. Capsula subglobosa, 3 mm. in diametro;
semina immatura 1 mm. lata. PARAGUAY: in campo, Paso Yobay,
Dec. 1938, F. Schade (type, GH).

Beyond saying that this species is apparently a member of
the Marchio-vaginatum complex, it is rather difficult to make
any statement about relationships. It is distinctive in the
complete similarity of cauline leaves and spathes, the latter
being 1-flowered. Nowhere else in the genus have I seen
solitary pedicels so strongly deflexed at maturity.

Sisyrinchium Reitzii, spec. nov. Herba perennis, tota dense pubes-
cens, rhizoma tenue, breve vel breve substoloniferum. Folia basalia
nulla; folia caulina ad bracteas acutas, spathiformes, convolutas
reducta, ad 2.5 em. longa. Caulis simplex, ad 30 cm. altus. Spathae
ad bracteas caulinas persimiles, ad 1.5 cm. longae, acutae, 1-florae,
vel raro 2-florae. Pedicelli spathas anthesin aequantes, glabri, in
fructu nec elongati nec deflexi. Ovarium glabrum, subglobosum, ad
1.5 mm. longum. Perianthium flavum, non lineatum; tepala subae-
qualia, ad 8 mm. longa et 3 mm. lata, elliptica, apice breve mucronata.
Filamenta ad 2.5 mm. longa, basi 1.5 mm. coalita; antherae lineares,
2 mm. longae. Stylus circa 2 mm. longus, styli rami circa 2 mm. longi.
Capsula matura seminaque ignota. BRAZIL: Sâo PAULO: Santo
Amaro, 15 Apr. 1932, Hauff, no. 18 (GH) ; SANTA CATARINA: Morro do
Iquererim, Campo Alegre, 1400 m., 5 Sept. 1957, Reitz & Klein, no.
4872 (type, US), 1500 m., 18 Oct. 1957, Reitz & Klein, no. 5228 (US).

Like S. deflexum, S. Reitzii is a member of the Marchio-
vaginatum complex. Superficially, it bears a strong resem-
blance to S. deflexwm. It differs, however, in the dense,
short, grayish pubescence which covers leaves, stems and
spathes. Sometimes the spathes are 2-flowered, rather than
1-flowered, but the pedicels are neither elongate nor deflexed
in fruit. It has been named in honor of Father Raulino Reitz,
the leading student of the flora of the state of Santa Cata-
rina in Brazil.

1962] Hedyotis — Lewis and Terrell 313

CHROMOSOMAL RACES IN
EASTERN NORTH AMERICAN SPECIES
OF HEDYOTIS (HOUSTONIA)!

WALTER H. LEWIS AND EDWARD E. TERRELL

In a phylogenetic study of the North American species of
Hedyotis (including Houstonia) (Rubiaceae), Lewis (1962
b) reported diploid and tetraploid races in populations of H.
caerulea, H. polypremoides (Gray) Shinners, and H. pur-
purea. Reference was made to a more extensive study of
the occurrence of infraspecific polyploidy. The purpose of
this paper is to report these findings.

Chromosome numbers of 116 collections of species in
the H. caerulea and H. purpurea groups were determined.
P. M. C. meiosis or somewhat less frequently mitosis was
studied in immature flower buds fixed in the field. The
methods used to prepare slides were identical to those des-
cribed for Hedyotis (Lewis, 1962b). In addition, represent-
ative permanent slides were mounted in euparal. Voucher
specimens for each collection are filed in the herbaria of the
U.S. National Museum (US), the Southern Methodist Uni-
versity (SMU), or Indiana University (IND; one collec-
tion).

The nomenclature of the H. purpurea complex follows the
revision by Terrell (1959) except that the taxa are treated
under Hedyotis rather than Houstonia (Lewis, 1961). This
does not imply acceptance of Hedyotis by Terrell.

RESULTS

The chromosome numbers of Hedyotis purpurea, H. longi-
folia, H. canadensis, H. nuttalliana ( — Houstonia tenuifolia
Nutt.), H. caerulea, and H. michauxii ( = Houstonia serpyl-
lifolia Michx.), are listed with voucher data and, in
parentheses, the number of plants examined. Although the
chromosome numbers of several collections were published
by Lewis (1962b), as indicated by an asterisk, they are in-
cluded for completeness and as a basis for mapping.

"This study was supported by a grant to the first author from the National Science
Foundation (G-9800).

314 Rhodora [Vol. 64

1. H. purpurea (L.) T. & G.

la. H. purpurea var. purpurea

n — 6. SOUTH CAROLINA: Edgefield Co., 4 miles WSW of Owdoms,
Terrell & Barclay 3430 (3). TEXAS: Newton Co., 2.8 miles S of
Newton, Lewis 5620 (5)*.

n = 6, n —12. TENNESSEE: Polk Co., 0.5 mile N of Hwys. 64 &
30 junction, Lewis 5638 (4 2n, 2 4n).

n = 12. GEORGIA: Gilmer Co., 3.9 miles NE of Ellijay,Lewis 5645
(3). MARYLAND: Montgomery Co., Wheaton, Terrell 3525 (4). NORTH
CAROLINA: Guilford Co., Guilford College, Terrell 3244 (1)*: Jackson
Co., summit of Whitesides Mountain, Terrell & Barclay 3472 (3);
Macon Co., SE of Highlands, Terrell & Barclay 3463 (2); Standing
Indian Wildlife Management Area, Terrell & Barclay 3478 (2).
TENNESSEE: Cheatham Co., near Kingston Springs, Terrell & Barclay
3360; Cumberland Co., 1.5 miles W of Ozone, Terrell & Barclay 3516
(3); Polk Co., 2.5 miles E of Hwys. 64 & 30 junction, Lewis 5643
(atypical) (4); Sullivan Co., near Sullivan-Carter Co. line & Hwy.
19E, Terrell & Barclay 3524 (8). VIRGINIA: Sussex Co., along Notto-
way River W of Homeville, Terrell 3636 (2).
lb. H. purpurea var. calycosa (Gray) Fosberg

n — 6. TENNESSEE: Wilson Co., Cedars of Lebanon State Park,
Terrell & Barclay 3339 (5); W border of Lebanon, Terrell & Barclay
3340 (6).

n — 12. ALABAMA: Franklin Co., 7 miles E of Russellville, Terrell
& Barclay 3499 (3); just E of Russellville, Terrell & Barclay 3361
(2) ; Jackson Co., 3 miles W of Scottsboro, Terrell & Barclay 3485 (3).
MISSISSIPPI: Carroll Co., 1.5 miles E of Carrolton, Lewis 5634 (atypi-
cal) (3). TENNESSEE: Maury Co., 16 miles NW of Lewisburg, Terrell
& Barclay 3351 (4)*; Meigs Co. 7 miles S of Decatur, Terrell &
De Selm 3246 (1).
le. H. purpurea var. montana (Small) Fosberg

n = 6. TENNESSEE-NORTH CAROLINA: Carter Co.-Mitchell Co.,
summit of Roan Mountain, Terrell & Barclay 3615 (2).

2. H. longifolia (Gaertn.) Hook.

2a. H. longifolia var. longifolia

n = 6. MASSACHUSETTS: Middlesex Co. Horn Pond, Woburn,
E & B Terrell 3595 (3).

n-—6, n= 12. MINNESOTA: Chisago Co. St. Croix River at
Taylors Falls, Likhite, July 1961 (2 2n, 5 4n).

n = 12. ONTARIO: Norfolk Co., Turkey Point, C & M Heimburger,
15 July 1961 (1).
2b. H. longifolia var. compacta (Terrell) Lewis

n —6. GEORGIA: Meriwether Co. 5 miles W of Greenville,
Terrell & Barclay 3410 (2). NORTH CAROLINA: Alexander Co., base
of Rocky Face Mountain, Terrell 3257 (1)*. SOUTH CAROLINA: Abbe-
ville Co., E side of Savannah River, Terrell & Barclay 3424 (atypical)
(1); Anderson Co., ca. 11 miles NNE of Anderson, Terrell & Barclay

1962] Hedyotis — Lewis and Terrell 315

3451 (2); Lexington Co., ca. 10 miles S of Columbia, Terrell & Barclay
3444 (atypical) (3). VIRGINIA: Augusta Co., 11 miles NE of Hwys.
42 & 250 junction, E & B Terrell 3569 (3); Frederick Co., 2 miles
WNW of Gore, E & B Terrell 3539 (2); Page Co.-Madison Co. line,
Milepost 47, Skyline Drive, Shenandoah Natl. Pk., E & B Terrell 3577
(3); Rappahannock Co., Hazel Mountain Overlook, Shenandoah Natl.
Pk., Terrell 60-11 (2) (no voucher); Shenandoah Co., 4.5 miles SW
of Jerome, E & B Terrell 3567 (3). Another collection is quite atypi-
cal: GEORGIA: Dade Co., Lookout Mountain, Terrell & De Selm 3254

(1).

n — 12. MARYLAND: Frederick Co., Boonesboro Mountain Rd. &
Hwy. 40 junction, E & B Terrell 3533 (4). SOUTH CAROLINA: Edge-
field Co., near Hwy. 378 & 67 junction, Terrell & Barclay 3429 (3).
VIRGINIA: Augusta Co.-Albemarle Co., 0.4 miles S of Milepost 97,
Skyline Drive, Shenandoah Natl. Pk, E & B Terrell 3570 (3);
Bedford Co., Milepost 90, Blue Ridge Parkway, Terrell 3250 (2);
Reckingham Co.-Greene Co., Sandy Bottom Overlook, Skyline Drive,
Shenandoah Natl. Pk., E & B Terrell 3573 (2). WEST VIRGINIA:
Pendleton Co., 5 miles N of Hwy. 33 & Spruce Knob rd. junction,
E & B Terrell 3545 (3).
2c. H. longifolia var. glabra (Terrell) Lewis

n — 6. NORTH CAROLINA: Jackson Co., summit of Whitesides
Mountain (type locality), Terrell & Barclay 3473 (3); Macon Co.,
near Highlands, Terrell 3255 (1)*; Slick Rock, SE of Highlands,
Terrell & Barclay 3464 (2).

3. H. canadensis (Willd. ex R. & S.) Fosberg

n = 6. TENNESSEE: Cheatham Co., near Kingston Springs, Ter-
rell & Barclay 3358 (4).

n — 12. INDIANA: Clark Co., 4 miles NW of Henryville, Terrell
3251 (1)*. TENNESSEE: Marion Co., below Hales Bar Dam, Terrell &
Barclay 3480 (3). ONTARIO: Bruce Co., Bruce Peninsula, Red Bay,
Heimburger & Price, 11 June 1961 (2).

4. H. nuttalliana Fosberg

n — 6. ALABAMA: Randolph Co., ca. 5 miles W of Wedowee, Ter-
rell & Barclay 3397 (4); Saint Clair Co., 0.5 mile W of Hwys. 35 & 53
junction, Terrell & Barclay 3391 (3). ARKANSAS: Montgomery Co.,
11.5 miles WSW of Norman, Lewis 5169 (2)*. GEORGIA: Heard Co.,
3 miles SW of Franklin, Terrell & Barclay 3401 (5). NORTH CAROLINA:
Stokes Co., Hanging Rock State Park, Terrell 3243 (1)*. OKLAHOMA:
Leflore Co., Lake Wister State Park, Lewis 5161 (2)*. SOUTH CAR-
OLINA: Lexington Co., ca. 4 miles E of Lexington Co.-Saluda Co. line,
Terrell & Barclay 3445 (3). VIRGINIA: Shenandoah Co., 2.6 miles E
of Liberty Furnace, E & B Terrell 3560 (3).

5. H.caerulea (L.) Hook.
5a. H. caerulea var. caerulea
n= 8. ALABAMA: Blount Co., ca. 3 miles E of Cullman, Terrell

316 Rhodora [Vol. 64

& Barclay 3387 (3); Randolph Co., ca. 5 miles W of Wedowee, Terrell
& Barclay 3398 (8); Saint Clair Co., 0.5 mile W of Hwys. 85 & 53
junction, Terrell & Barclay 3392 (8); Talladega Co., 2-3 miles W of
Talladega Co.-Clay Co. line & Hwy. 77, Terrell & Barclay 3396 (2).
ARKANSAS: Clark Co., 2.4 miles SE of Alpine, Lewis 5171 (3) *: Dallas
Co., 1.9 miles S of Ouachita, Lewis 5172 (3)*; Hot Spring Co., 1.5
miles NE of Donaldson, Lewis 5613 (1). GEORGIA: Gilmer Co., 3.9
miles NE of Ellijay, Lewis 5646 (1). INDIANA: Brown Co., near
Trevlae, Heiser & Ellis 4763 (IND) (3). KENTUCKY: Logan Co., 1.3
miles SW of Auburn, Lewis 5602 (2); Metcalf Co., 3.1 miles E of
Wisdom, Lewis 5605 (4). NORTH CAROLINA: Alleghany Co., 2 miles
NE of Blue Ridge Parkway & Hwy. 21 junction, Terrell 3242 (2);
Jackson Co., ca. 4 miles E of Cashiers, Terrell & Barclay 3455 (3);
Macon Co., Nantahala River & Hwy. 64, Terrell & Barclay 3479 (3).
SOUTH CAROLINA: Lexington Co., ca. 4 miles E of Lexington Co.-
Saluda Co. line, Terrell & Barclay 3446 (3). TENNESSEE: Cheatham
Co., near Kingston Springs, Terrell & Barclay 3359 (3); Stewart Co.,
12.9 miles WSW of Dover, Lewis 5601 (2); Kentucky Lake & Hwy.
79, Lewis 5610 (2).

n — 9. ARKANSAS: Hot Spring Co., 3.6 miles NNE of Hwys. 67
& 210 junction, Lewis 5612 (2).

n — 16. CONNECTICUT: Litchfield Co., 1-1/2 miles S of junction
112 & 7, Salisbury Twp., Terrell 3662 (1); Tolland Co., 3 miles S of
Rockville, Terrell 3663 (2); near State Line Pond, Terrell 3664 (3).
MAINE: York Co., North Berwick, E & B Terrell 3591 (1). MARYLAND:
Frederick Co., Catoctin Creek & Hwy. 17, E & B Terrell 3538 (1);
Montgomery Co., Wheaton Regional Park, Terrell 3526 (2). MASSA-
CHUSETTS: Franklin Co., 3 miles S of Northfield, Terrell 3667 (3);
Worcester Co., 1/2 mile N of Hardwick, Terrell 3665 (3). NEW HAMP-
SHIRE: Cheshire Co., 3 miles E of Winchester, Terrell 3666 (1) ; Coos
Co., lower slopes of Mount Washington (alt. 2500 ft.), E & B Terrell
3589 (2); Grafton Co., Hanover, Fosberg 39838 (2). NORTH CAROLINA:
Guilford Co., Guilford College, Terrell 3231 (5)*; Stokes Co., NW of
Francisco, Terrell 3239 (1)*. PENNSYLVANIA: Franklin Co., 0.5 mile
N of Richmond Furnace, Terrell 3629 (2) ; Juniata Co., 5 miles NE of
East Waterford, Terrell 3628 (4); Perry Co., 1 mile SE of Sherman-
dale, Terrell 3627 (3). SOUTH CAROLINA: Edgefield Co., ca. 4 miles
WSW of Owdoms, Terrell & Barclay 3431 (2). TENNESSEE: Cumber-
land Co., 7 miles SE of Mayland, Terrell & Barclay 3514 (3); Knox
Co., Knoxville, De Selm, 1 May 1962 (1). VERMONT: Windham Co., 2
miles W of Brattleboro, Terrell 3668 (2). VIRGINIA: Hanover Co., 2
miles NW of Doswell, Terrell 3631 (2); Page Co.-Madison Co., Big
Meadows, Shenandoah Natl. Pk., E & B Terrell 3575 (2). wEST VIR-
GINIA: Tucker Co., 8 miles S of Davis, E & B Terrell 3544 (2); 1-2
miles N of Thomas, E & B Terrell 3540 (1). ONTARIO: Toronto (intro-
duced), Heimburger, 5 May 1961 (4) (omitted from map in Fig. 2).

n = 24. VIRGINIA: Sussex Co., along Nottoway River W of Home-
ville, Terrell 3634 (1).

1962] Hedyotis — Lewis and Terrell 317

5b. H. caerulea var. faxonorum (Pease & Moore) Fosberg
n — 16. NEW HAMPSHIRE: Coos Co, below summit of Mount
Washington (alt. 6000 ft.), E & B Terrell 3586 (4).

6. H. michauxii Fosberg

n — 16. NORTH CAROLINA: Macon Co., Standing Indian Wildlife
Area, Terrell & Barclay 3477 (3); Transylvania Co., 0.5 mile N of
Davidson River & Hwy 64 junction, Terrell & Barclay 3460 (4).
TENNESSEE: Monroe Co., Cherokee National Forest, De Selm, 1 May
1962 (3). TENNESSEE-NORTH CAROLINA: Carter Co.-Mitchell Co., sum-
mit of Roan Mountain, Terrell & Barclay 3520 (5).

n — 24. NORTH CAROLINA: Alleghany Co., 2 miles NE of Blue
Ridge Parkway & Hwy. 21 junction, Terrell 3241 (1)*.

7. Putative hybrids or intergrading collections
"a. H. longifolia—H. purpurea

n — 6. SOUTH CAROLINA: McCormick Co., 2.5 miles NW of Mount
Carmel, Terrell & Barclay 3425 (1) (< longifolia).

n = 12. GEORGIA: Elbert Co., 2 miles W of Hwys. 72 & 79 junc-
tion, Terrell & Barclay 3423 (2) (< longifolia); Rabun Co., ca. 7
miles S of North Carolina-Georgia line & Hwy. 28, Terrell 3147 (1);
Walton Co., 4 miles NE of Monroe, Terrell & Barclay 3420 (2)
(< longifolia). TENNESSEE: Cumberland Co., 2 miles NW of Mayland,
Terrell & Barclay 3513 (3) (< purpurea).

Tb. H. canadensis > — H. purpurea

n — 12. INDIANA: Clark Co., 4 miles NW of Henryville, T'errell
$252 (1).

Te. H. caerulea—H. michauxii

n — 16. WEST VIRGINIA: Tucker Co., just above Blackwater Falls,
E & B Terrell 3542 (5).

DISCUSSION

The basic chromosome number of Hedyotis purpurea, H.
longifolia, H. canadensis, and H. nuttalliana is x — 6. The
basic chromosome number of H. caerulea and H. michauxii
is x = 8. These data agree with the morphological evidence:
the first four species are closely related and were included
by Terrell (1959) in the H. purpurea group; the last two
species are so alike that herbarium material sometimes is
difficult to distinguish.

A dash, —, is used here to indicate morphological intergradation. The second author
prefers to use this symbol instead of X, as in this case additional knowledge about
the species is needed to attribute their intermediacy to hybridization. He believes
that botanists should restrict the use of X to instances in which evidence of hybridi-
zation has been thoroughly studied. 'The symbols > and < were used by others (cf.
Li, 1957; Hardin, 1958). For example, H. canadensis > —H. purpurea denotes speci-
mens which intergrade but are more similar to the former species.

318 Rhodora [Vol. 64

In H. purpurea (Fig. 1) our sampling indicates that tetra-
ploid populations are in the great majority. The only known
2n populations are in the southern part of the range. One
population included both 2n and 4n individuals. The sam-
pling is not sufficient to show any geographic separation of
chromosomal races, if such exists. Var. montana, the only
known stations for which are on Roan Mountain and Grand-
father Mountain, is diploid at the former station (not shown
in Fig. 1).

In contrast to H. purpurea, H. longifolia has almost twice
as many 2n populations as 4n. These are found in the north

- Kens
(ia)
76

e
/ O
| O
O FIG. |
P4
\
\ ^ H PURPUREA & @
-—t- vor. purpurea @
-— —— var. calycosa O O

Q 4 / - x
L phy Y o
^ ? | A

Figure 1. General'zed range (enclosed by dashed line) of H. purpurea and locations
of diploid and tetraploid collections. Var. montana not shown. New England distribu-
tion of var. calycosa not shown.

\

1962] Hedyotis — Lewis and Terrell 319

as well as in the south, and one population in Minnesota con-
tains individuals with both chromosome numbers. Var. gla-
bra, restricted to the southern Appalachians, is diploid on
the basis of three collections.

Plants of H. canadensis from only four localities were
studied but these included samples from the extremes of the
species range in Tennessee and Ontario. On the basis of the
small sample, the tetraploids prevail over the diploids in a
proportion similar to that for H. purpurea. With both 2n
and 4n races in Tennessee, no geographic separation of the
forms is apparent. Only 2n individuals were found among
plants of H. nuttalliana from 8 stations in 7 states, from
Virginia to Oklahoma. This sample is sufficiently large and
widespread to indicate that tetraploid individuals must be
rare in H. nuttalliana if they exist. Infraspecific polyploidy
is probably unimportant in the recent evolution of the spe-
cies. On the basis of the frequency of diploidy and tetra-
ploidy, with the high frequency of tetraploidy representing
the higher stage of evolution, the species in the x = 6 group
can be ranked from the simplest to the most specialized as:
H. nuttalliana (2n only), H. longifolia (2n:4n = 2:1), H.
purpurea and H. canadensis (2n:4n = 1:3). Using this
criterion, H. nuttalliana is the least evolved species of the
series, a position corroborating the results of Lewis (1962b)
based chiefly on morphological evidence.

Among 45 collections of H. caerulea var. caerulea sampled
from a wide area of the species range, 18 were diploid (n =
8), 25 were tetraploid (n = 16), one was aneuploid (n =
9), and one was hexaploid (n = 24). Thus, this species con-
sists predominantly of 2n and 4n populations, with the latter
slightly outnumbering the former in our sample. The base
number is x — 8. The chromosomal races appear to have
partly distinct distributions: the diploid race is widely dis-
tributed in the southern United States (Fig. 2) and in our
sample occurs only south of the glacial boundary. Only
tetraploids are known in New England, but they do occur
also in certain parts of the South. The single collection of
var. faxonorum, a variety restricted primarily to higher
altitudes in the White Mountains, was tetraploid.

On the basis of a single population of H. michauxii, Lewis

320 Rhodora [Vol. 64

(1962b) reported the species as hexaploid. Additional sam-
pling has shown, however, that 4n and 6n races exist, the
former occurring much more frequently. The species is
limited in distribution to the higher regions of the southern
and central Appalachians, a distribution interestingly asso-
ciated with the finding of two polyploid races and no diploid
forms in a subgenus where “pure” polyploid species are
infrequent.

In the P. M. C.’s of all plants examined, bivalent formation
and separation by and large were regular. Occasionally 1 to
3 quadrivalents were observed at metaphase I in the 4n
plants (e.g., H. purpurea, Terrell & Barclay 3351; H. caeru-

H, CAERULEA

@ 2n

e. eo^
° T 6n
e.
A @

- —
———-d—————l——

C ls UN EN,

Figure 2, Generalized range (enclosed by dashed line) of H. caerulea and locations
of diploid and polyploid collections. The single aneuploid (n = 9) collection not shown.

1962] Hedyotis — Lewis and Terrell 321

lea, Terrell 3627), but these were exceptional. Spindle dis-
turbances were not observed. In one cell, a plant of H.
caerulea (Lewis 5610) had 2n —15, but all other mitotic
cells had 2n = 16 and in the P. M.C.'s & —8. Although
statistical data are not available, plump, well-stained pollen
grains were noted for most diploids and tetraploids. If the
fertility of the tetraploids was decreased markedly in com-
parison with the diploids it was not apparent from observa-
tions of pollen.

Seven collections were of intermediate or intergrading
morphology. Certain of these are putative hybrids. As no
definite evidence of hybridization is available, we are using
a dash (—) to connect the two names. This is intended to
indicate intergradation without necessarily suggesting hy-
bridization as its cause. In the seven collections diploid and
tetraploid populations were found, with the latter more
frequent. Of the five collections of H. purpurea — H. longi-
folia, a diploid (Terrell & Barclay 3425) had regular pairing
at metaphase I as did all P. M. C.'s examined from the tetra-
ploid individuals of T. & B. 3420 and 3513. Only mitosis was
observed in the other two collections of H. longifolia — H.
purpurea. and in H. canadensis — H. purpurea. Meiosis in
H. caerulea — H. michauxii was regular; this population is
not thought to be of hybrid origin. Pollen was more or less
normal in individuals of the seven collections. In general,
the chromosome data indicate that chromosome number per
se is no barrier to hybridization in these species within each
ploidy level, i.e. 2 = 6 and x = 8. The chromosome numbers
in the H. purpurea group are entirely compatible with the
data from morphology expressed in the taxonomic treatment
of this group by Terrell (1959) but the few admittedly pre-
liminary chromosome counts listed in the same paper should
be ignored.

As more cyto-population research is completed, it becomes
apparent that varying chromosome numbers for a species
are much more common than hitherto suspected. In the
Rubiaceae alone, where very few cytological studies have
been completed, infraspecific polyploidy has been recorded
for species of 13 genera, particularly in the Rubieae (Fager-
lind, 1937; Ehrendorfer, 1961), Coffea and Coprosma (cf.

322 Rhodora [Vol. 64

Darlington & Wylie, 1956), Bouvardia (Lewis, 1962a), and
Hedyotis subg. Oldenlandia (Lewis, 1959, 1962b).

In Hedyotis subg. Edrisia, in which the species under dis-
cussion are classified, chromosome numbers of 25 species are
known (Lewis, 1962b). Of these, seventeen species are
diploid, four species are diploid and tetraploid, one species
is diploid, tetraploid, and hexaploid, two species are tetra-
ploid, and one species is tetraploid and hexaploid. Twenty-
three species are euploid and two have recorded aneuploid
individuals. Thus six species or 24% of the North American
species studied have infraspecific polyploidy, 4% exhibit
infraspecific aneuploidy, and 4% exhibit both infraspecific
polyploidy and aneuploidy. All of these species are hetero-
stylous perennials and with one exception are herbaceous.
No chromosomal plasticity is known among the eight annual
species of the subgenus or among the species having only
short-styled flowers. This suggests a certain correlation in
the subgenus between the heterostylous species constructed
for outbreeding and the perennial habit with the occurrence
of infraspecific polyploidy and aneuploidy.

The distributions of the diploid and tetraploid races of
H. caerulea, H. purpurea, and H. longifolia may be grouped
into two categories, viz., those with different geographic dis-
tributions and those without such tendencies. As an example
of the first kind, the Zn race of H. caerulea occupies a south-
ern range while the 4n race has a more northern distribu-
tion. Possibly the tetraploid race colonized areas drastically
altered by glaciation in the northern region of the species’
present range, while the ancestral race remained predomi-
nantly south of the glacial boundary (cf. Stebbins, 1950).
It also appears that the 4n race is now successfully compet-
ing with the 2m race somewhat south of the southern limits
of the last glaciation. The high altitude distribution and the
polyploid races of H. michauxii also suggest the colonization
of less favorable environments by a polyploid species. That
this tendency is not universal, even for species in the same
subgenus, is suggested by the infraspecific distributions for
H. purpurea and H. longifolia. Their diploid and tetraploid
races are about equally distributed and do not appear to have

1962] Hedyotis — Lewis and Terrell 323

either peripheral or central concentrations although we did
not sample their entire ranges.

When herbarium specimens from all of these collections
were compared, tetraploid and diploid (and hexaploid)
plants of each taxon looked just alike; there were no observ-
able gross morphological differences in plants of different
ploidy levels and it is clear that these are autopolyploids.
Five out of the six species had polyploid populations. Auto-
polyploidy is an essential feature of the evolution of the H.
caerulea and H. purpurea groups.

ACKNOWLEDGEMENTS

We appreciate the assistance of those who contributed material for
cytological study, viz., Drs. H. R. De Selm, F. R. Fosberg, M.
Heimburger, C. B. Heiser, and V. N. Likhite. Special appreciation
is due Dr. A. S. Barclay for assistance in the field. — STEPHEN F.
AUSTIN STATE COLLEGE, NACOGDOCHES, TEXAS, AND CROPS RESEARCH
DIVISION, AGRICULTURAL RESEARCH SERVICE, U. S. DEPARTMENT OF AGRI-
CULTURE, BELTSVILLE, MARYLAND.

LITERATURE CITED

DARLINGTON, C. D. and A. P. WYLIE. 1956. Chromosome Atlas of
Flowering Plants. The Macmillan Co., New York.

EHRENDORFER, F. 1961. Evolution of the Galium multiflorum com-
plex in western North America. I. Diploids and polyploids in this
dioecious group. Madrono 16: 109-122.

FAGERLIND, F. 1937. Embryologische, Zytologische und Bestaéu-
bungsexperimentele Studien in der Familie Rubiaceae nebst
Bemerkungen über einige Polyploiditátsprobleme. Acta Hort.
Berg. 11: 195-470.

HARDIN, J. W. 1958. The annotation of introgressants. Taxon 7
(2) : 52-53.

Lewis, W. H. 1959. Chromosomes of east Texas Hedyotis (Rubi-
aceae). Southwest. Nat. 3: 204-207.

. 1961. Merger of the North American Houstonia and

Oldenlandia under Hedyotis. Rhodora 63: 216-223.

1962a. Chromosome numbers in North American

Rubiaceae. Brittonia 14: 285-290.

1962b. Phylogenetic study of Hedyotis (Rubiaceae)
in North America, Amer. Jour. Bot. 49: 855-865.

Li, HurLiN. 1957. A symbol for introgressants. Taxon 6(8):
216-218.

STEBBINS, G. L. 1950. Variation and Evolution in Plants. Colum-
bia University Press, New York.

TERRELL, E. E. 1959. A revision of the Houstonia purpurea group
(Rubiaceae). Rhodora 61: 157-180, 188-207.

324 Rhodora [Vol. 64

A NEW CRUCIFER FROM
THE GREAT SLAVE LAKE AREA OF CANADA

REED C. ROLLINS

A general problem of classification in the Cruciferae re-
sults from the lack of sharp definitive boundaries between
the genera in some sections of the family. This problem has
long been recognized. The situation is readily understood
and should perhaps even be expected because of the rela-
tively short evolutionary history of the family. However, an
understanding of the situation does not alleviate the torment
that ensues whenever an unknown species is found that does
not quite fit well-established genera. The uncertainty as to
which genus might be involved requires that all possible
genera be carefully checked and studied to determine wheth-
er or not the species has been described in any one of them.
This in itself is a much longer and more laborious process
than if the species were clearly referable to a well known
and well defined genus. Furthermore, the unknown species
is usually not closely related or readily comparable to a
known species and thus the comparative procedures usually
employed in fitting such a species into a classificatory scheme
are not applicable.

For over a year, we have been periodically concerning
ourselves with material of just such an unknown species.
Four collections of it were made by John W. Thieret and
Robert J. Reich, along the Yellowknife Highway near Great
Slave Lake of northwestern Canada, during June and July
of 1961. A fifth collection was made in August, 1962, by
Thieret. The plants grow in marshy to wet situations and
develop relatively large lateral roots. The petiolate basal
leaves with more or less crenate margins are reminiscent of
Armoracia aquatica or possibly horseradish, Armoracia
rusticana, but there the similarity stops. The long, slender,
wide-spreading pedicels suggest Rorippa nasturtium-aqua-
ticum. However, other features, such as the undivided
leaves and uniseriate seeds, make a close association of the
plant in question with watercress untenable. All charac-
teristics considered, the new species does fall more nearly
within the group of species making up Rorippa than any

1962] New Crucifer — Rollins 325

Plate 1271. Silhouette photograph of two fertile specimens and one sterile specimen
of Rorippa crystallina, X 2/5. The specimen at right and the sterile specimen make
up a part of the holotype sheet.

326 Rhodora [Vol. 64

other genus known to me, even though no single species can
readily be singled out for easily handled comparative pur-
poses.

An interesting feature of the new species is the basis for
the specific epithet chosen, crystallina.! Relatively large
crystals of calcium oxalate are found in the inner tissues of
the fleshy leaves, the stems, pedicels and fruits. These crys-
tals occupy several adjacent cells and are large enough to
erupt the leaf-surface when the plants are pressed flat in
drying them for specimen purposes. According to Metcalfe
and Chalk? calcium oxalate crystals are rare in the Cruci-
ferae. These authors indicate that crystals are present in
Crambe and Sisymbrium but Rorippa is not mentioned as a
genus in which calcium oxalate crystals have been observed.
The number and distribution of crystals in Rorippa crystal-
lina is variable. These crystals may be clustered or widely
spaced, associated with or near vascular bundles, or remote
from them. There is no evident association with vascular
trace endings in the leaves. In the larger basal leaves, the
crystals are more prominent and more numerous on the
lower side of the leaf than on the upper. They are more
numerous per unit of area in the smaller cauline leaves than
in the basal leaves and give a pustular appearance to the
leaf-surface, both above and below. Generally, they are
more prominent on the lower surface than on the upper.

Rorippa crystallina Rollins, sp. nov. Plate 1271

Perennial, glabrous throughout; roots thick, often well-developed
laterally; stems one to several, erect to decumbent, glabrous, 1-4 dm.
long, arising below an active fascicle of leaves, unbranched except in
the inflorescence; basal leaves borne on short shoots,fascicled, petiolate,
ovate to narrower, obtuse, shallowly and coarsely dentate, blade 4-10
(-20) em. long, 1.5-4 (-6) cm. wide, cuneate at base, petioles 3-8
(-12) em. long, winged above; lower cauline leaves petiolate, narrowly
oblanceolate, obtuse, 4-6 cm. long, 4-10 mm. wide, dentate, toothed
or lobed, petiole winged; upper cauline leaves sessile, overlapping,
lanceolate to nearly oblong, narrowed at base, entire to sparsely
dentate, thick when fresh; all leaves more or less pustular on both

1I am indebted to Professors I. W. Bailey and Adriance S. Foster for cleared leaves,
tests for calcium oxalate and shared observations on the material. My appreciation
goes to Dr. John W. Thieret for providing ample material upon which this study is
based.

2Anatomy of the Diocotyledons, Vol. 1, p. 82. 1950.

1962] New Crucifer — Rollins 327

surfaces from the presence of relatively large crystals present in the
mesophyll; individual stems terminated by a short raceme 5-10 (-15)
cm, long, occasional flowering branches just below main inflorescence;
sepals yellowish-green, non-saccate, broadly oblong, 4-5 mm. long;
petals spatulate, whitish, sometimes tinged below with light lavender,
6-8 mm. long, ca. 3 mm. wide; stamens tetradynamous. anthers ob-
long, ca. 1 mm. long; fruiting pedicels slender, spreading at right
angles to somewhat ascending, 12-18 mm. long, slightly enlarged at
apex; siliques terete, 1.5-2.5 cm. long, widely spreading to somewhat
ascending, nearly sessile or with a short thick gynophore less than
0.5 mm. long; valves with a central inconspicuous branching nerve;
styles ca. 1 mm. long; stigmas slightly bilobed with lobes over the
replum; ovules 10-15 in each loculus; funiculi free, rather spongy;
septum with a prominent central nerve-like area extending full length
down the middle, cells of the septum prominent, usually hexagonal;
seeds plump, broadly oblong, wingless, 1.5-2 mm. long, ca. 1.2 mm.
broad; seed-coat finely reticulate-colliculate, covered with a mucilagi-
nous sheath, buff-colored when dry, beak curved into a short hook,
funiculus detached at hilum leaving a circular scar, funicular append-
age absent; cotyledons accumbent.

Herba perennis caespitosa glabra, caulibus erectis vel decumbentibus
1-4 dm. longis; foliis pustulatis, basilaribus fasciculatis petiolatis,
laminis ad basim cuneatis ovatis vel anguste-ovatis dentatis 4-10 cm.
longis, 1.5-4 em. latis; foliis caulinis inferne crassis petiolatis anguste
oblanceolatis dentatis vel lobatis; foliis caulinis superne sessilibus
crassis integris vel sinuato-dentatis obtusis; sepalis nonsaccatis
obtusis 4-5 mm. longis; petalis spathulatis albis vel albido-lilacinis
6.5-8 mm. longis; pedicellis in fructu divaricatis tenuis 12-18 mm.
longis; siliquis teretibus divaricatis 1.5-2.5 cm. longis; stylis ca. 1 mm.
longis; seminibus oblongis immarginatis 1.5-2 mm. longis; cotyledoni-
bus accumbentibus.

Type in the Gray Herbarium collected in a Carex marsh at mile
23 N., along the Mackenzie River - Yellowknife Highway, northwest
of Great Slave Lake, District of Mackenzie, Northwest Territories,
Canada, July 10, 1961, John W. Thieret and Robert J. Reich 7512.

Other specimens studied, all from the same general area, and
deposited in the Gray Herbarium: mile 23.8 N., June 14, 1961,
Thieret and Reich 6637; mile 35 N., July 7, 1961, Thieret and Reich
7413; mile 16.5 N., July 9, 1961, Thieret and Reich 7484; mile 23.5 N.,
Aug. 9, 1962, Thieret 9085. — GRAY HERBARIUM, HARVARD UNIVERSITY.

328 Rhodora [Vol. 64

TWO NEW VARIETIES AND A NEW COMBINATION
IN RUDBECKIA

ROBERT E. PERDUE, JR.!

Rudbeckia nitida Nutt. var. texana Perdue var. nov. Folia integra,
crenata, dentata, vel serrata, dentibus (si adsint) numerosis crebris
10 vel pluribus quoque in margine; folia basalia anguste vel late ellip-
tica vel spatulata. Discus 2.0-4.5 cm. altus, achaenia 5.0-7.5 mm.
longa.

Leaves entire, crenate, dentate, or serrate, teeth abundant if present,
basal leaves narrowly to broadly elliptic, or spatulate, blade 10-15 em.
long, 3-6 cm. wide, sharply or broadly acute, attenuate below to a long
slender petiole; disk conical to conic-cylindrical, 2.0-4.5 cm. long;
achenes 5.0-7.5 mm. long.

TYPE: In prairie along railroad, 6 miles S. of Stowell, Chambers Co.,
Texas. L. H. Shinners 7710, May 15, 1945 (SMU).

DISTRIBUTION: Low open areas along roadsides and in fields; south-
eastern Texas to central Louisiana.

This is a morphologically distinct variety separated geo-
graphically from R. nitida var. nitida of eastern Georgia and
northeastern Florida by more than 400 miles. The abun-
dently toothed, crenate, dentate, or serrate leaves of R.
nitida var. texana readily distinguish it from the eastern
variety. The latter characteristically bears leaves with 4 to
8, distantly and more or less equally spaced small dentate
teeth on each margin.

Considerable justification could be offered to support the
description of this variety as a new species. On the other
hand, the conservative treatment I have applied to this and
related Rudbeckias provides a better arrangement for indi-
cating the relationships of the taxa.

Rudbeckia laciniata L. var. bipinnata Perdue var. nov. Folia caulina
inferiora et mediocria bipinnatifida vel pinnatisecta, segmentis vel
foliolis 5-7, pinnatifidis, segmetis ultimis lanceolatis. Discus globosus
vel ovoideus 1.5-1.9 cm. altus.

Stems 0.6-1.5 m. high, mostly slender, the peduncles rarely more
than 1.5 mm. in diameter; basal leaves not seen, lower and middle
stem leaves petiolate, bi-pinnatifid or pinnately divided into 5 or 7
irregularly pinnatifid leaflets or segments, the ultimate divisions
lanceolate, entire or coarsely toothed, upper stem leaves similar but
with fewer divisions, sessile, the uppermost merely 3-lobed or entire

— "Robert E. Perdue, Jr. Botanist, Crops Research Division, Agricultural Research
Service, U. S. Department of Agriculture, Beltsville, Maryland.

1926] Setcreasea — Faruqi, Mehra and Celarier 329

and ovate; heads mostly 8-15; rays 4.0 (3.5)-5.0 cm. long; disks glo-
bose to ovoid, 1.5-1.9 em. long; achenes 4-6 mm. high.

TYPE: Swampy open woods along Mud Creek, valley of Fall Creek,
Dryden, Tompkins Co, New York, MacDaniels & Eames 1293,

August 10, 1913 (GH).
DISTRIBUTION: Moist open or partly shaded sites: stream banks,

meadows, borders of woodland, open woods, etc.; Massachusetts and
New Hampshire to eastern Pennsylvania and Maryland.

R. laciniata var. bipinnata is intermediate between the
common R. laciniata var. laciniata of the eastern United
States and Canada, with which it intergrades imperceptibly,
and R. laciniata var. digitata (Mill.) Fiori of the northern
half of the Atlantic coastal plain. From var. laciniata the
new variety is distinguished by its bipinnatifid leaves with
narrowly lanceolate or linear-lanceolate ultimate segments.
From var. digitata it is distinguished by its larger disks
(1.5 cm. or more high) and longer rays (3.5 to 6.0 cm.
long).

Rudbeckia occidentalis Nutt. var. montana (Gray) Perdue, comb.
nov. Based on R. montana Gray, Proc, Amer. Acad. 17: 217 (1881-

1882).

A NEW SPECIES OF SETCREASEA FROM MEXICO

SHAMIM A. FARUQGI,' K. L. MEHRA,’ AND R. P. CELARIER*

The authors came across a new species of the genus Set-
ereasea from Mexico, collected by Edward Palmer, first
identified as S. pallida, but later as “S. palmeri” probably by
Rose, who, it seems, did not get a chance to describe it. Rose
(1891) did report another new species, Tradescantia pal-
meri, from Edward Palmer's collections of Mexico. Though
the genus Setcreasea was separated from Tradescantia at a
later date (Rose 1899, 1903, Schumann et Sydow 1899) on
the basis of fused corolla and epipetalous stamens, the
species T. palmeri should not be confused with this new
species, because 7'. palmeri is in no way similar to it.

1At present: Cameron State Agricultural College, Lawton, Oklahoma.

7At present: Division of Botany, Indian Agricultural Research Institute, New Delhi,
India.

Deceased December 23, 1959.

330 Rhodora [Vol. 64

On the strength of its dense-woolly trichomes on leaf, leaf
sheath, and pedicel, in combination with thick connective,
and pubescent ovary, we believe that the present taxon is a
new species. The name “S. palmeri” is not taken into con-
sideration to avoid a nomenclatural confusion.

Setcreasea lanceolata Faruqi, Mehra & Celarier, sp. nov.

Herba perennis, caulibus subramosis, erectis, glabratis, foliis lanceo-.
latis, pilosis, vaginis pilosis, usque 1.8 cm longis, calycibus pilosis,
ovario piloso.

Perennial herbs, 20-25 em high; rhizomes creeping; roots arising
from the underground nodes, fleshy and slender; aerial stems few,
succulent, erect or slightly decumbent, glabrous, striate when dry, the
internodes 5.5-8 cm long, those bearing inflorescences 6.8-10.2 em long;
lamina sessile, lanceolate, oblong or rarely elliptic, 10-15 cm long,
2.5-3 em wide, acute at the apex, sheath prominent, up to 1.8 em long,
both the lamina and the sheath dense-woolly; inflorescences terminal
subtended by 2 leaf-like bracts, bracts lanceolate, the outer 4.0 em
long, 2.0 cm wide, the inner 1.8 em long, 2.0 cm wide; flowers several,
pedicellate, the pedicels 6.0 mm long, woolly; sepals subequal, elliptic,
5.5 mm long, 2.0 mm wide; corolla "purple white with a bar of purple
color in the center of the limb’, gamopetalous, the tube conspicuous,
2.0 mm long; stamens 6, epipetalous, subequal, filaments barbate, the
connective thick and broad; ovary tricarpellary and pubescent; stigma
capitate, 3-lobed; fruit a three-chambered capsule; seeds dark gray 1-2
in each locule.

Mexico: San Louis Potosi, Edward Palmer 135 (GH type, US).

RELATIONSHIPS

Setcreasea lanceolata may show a superficial resemblance
to S. hirsuta Markgraf (1952). It differs from S. hirsuta
in its densely pubescent leaves, sepals and ovary. According
to Markgraf's description S. hirsuta is pubescent only at leaf
bases while its calyx and ovary are glabrous.

In spite of the differences in size and shape of leaf, and
pubescence of leaf and calyx, S. lanceolata shares two very
important characteristics with S. brevifolia, i.e., highly
pubescent ovary and thick connectives. The evolutionary
pattern indicates, that in the genus Setcreasea leaf size and
shape have undergone many more changes than ovary and
stamen. Thus, similarities of connectives and ovary may
indicate a closer ancestral relationship of S. lanceolata and
S. brevifolia than any other species of this genus.

^Annotation on the herbarium sheet.

1962] Dasistoma — Piehl 331

ACKNOWLEDGEMENTS
The authors are thankful to the curators of the Gray Herbarium
of Harvard University and United States National Herbarium for the
loan of the herbarium specimens, and to Dr. Shiu Ying Hu of Arnold
Arboretum, Harvard University, for reading the manuscript. —
DEPARTMENT OF BOTANY AND PLANT PATHOLOGY, OKLAHOMA STATE
UNIVERSITY, STILLWATER, OKLAHOMA.

LITERATURE CITED
MARKGRAF, F. 1952. Eine neu Setereasea aus Mexico. Mitt Bot.
Staatssam München. 5: 166-167.
Rose, J. N. 1891. List of plants collected by Dr. Edward Palmer
in Western Mexico and Arizona in 1890. Contr. U. S. Nat. Herb.

1: 91-127.
1899. Treleasea, A new genus of Commelinaceae, Contr.

U. S. Nat. Herb. 5: 207-208.
19083. Studies of Mexican and Central American Plants

No. 3. Contr. U. S. Nat. Herb. 8:1-55.
SCHUMANN, K. ET Sypow (1899) 1901. In Just, Bot. Jahrb. 27(1) : 452.

THE PARASITIC BEHAVIOR OF
DASISTOMA MACROPHYLLA'

MARTIN A. PIEHL

The first and perhaps only reference to the parasitic
behavior of Dasistoma macrophylla (Nutt.) Raf. appears
to be that of Pennell (1928), who in the course of his pri-
marily taxonomic studies of the Scrophulariaceae observed
parasitic attachments to buckeye, Aesculus glabra, Willd., in
Indiana. In the same paper he lists Dasistoma as one of the
genera which may be restricted to a single host. That he
had probably not learned of additional hosts in the years
shortly after this observation is indicated by the fact that
he stated simply that Dasistoma “is parasitic upon the roots
of Aesculus" (Pennell, 1935, p. 405).

Further information on this point has been recently re-
ceived from Professor Edgar T. Wherry of the University
of Pennsylvania who relates an incident when Pennell noted
disagreement in the ranges of Dasistoma and Aesculus

1Facilities for this study were provided by the Department of Botany, University of
Michigan, Ann Arbor, where the author was engaged in doctoral studies.

332 Rhodora [Vol. 64

glabra in Indiana, he communicated with C. C. Deam, saying
that he felt further study would show that Aesculus was
present wherever Dasistoma was. After further considera-
tion of the range of Aesculus in that state, Deam agreed that
this was true. Thus, it would appear that Pennell felt this
species was an obligate parasite on Aesculus. A parallel situ-
ation has been carried in the literature for many years with
regard to the host specificity of Buckleya distichophylla
Torr., which is claimed to be restricted to hemlock, Tsuga
(e.g., see Fernald, 1950). Such (presumed) highly special-
ized host-parasite relationships are to be doubted when one
considers that all other chlorophyllous parasites? of the
Scrophulariaceae and Santalaceae that have been studied in
some detail are known to parasitize a number of hosts.

The present study was made to determine if Dasistoma
does in fact parasitize other plants as well, thereby making
it possible (if it is an obligate parasite) for it to grow in
areas where Aesculus is absent. Further, within the Scro-
phulariaceae, observations on parasitism are lacking for the
tribe Buchneriae, to which Dasistoma belongs, except for
some species of Gerardia L. (Agalinis Raf.). Most previ-
ous observations are for the members of the Euphrasiae.
None of the several manuals consulted (e.g., Fernald, 1950;
Gleason, 1952) which cover the range of this species have
included the fact that this relatively little known plant is
parasitic.

As Pennell (1928, 1935) has indicated, there have been
long-term differences of opinion and confusion concerning
the placement of this species as to genus. It was given the
name Dasistoma (aurea) by Rafinesque (1819), who later
(1837) apparently gave it a new name Dasistema (auricu-
lata). Bentham (1846) altered the spelling to Dasystoma,
but misidentified Rafinesque’s description, applying the
name not to this plant but to Aureolaria Raf. Nuttall (1818)
placed it in his genus Seymeria, where it has been main-
tained in some recent works (e.g., Fernald, 1950). I have

*Such photosynthetic parasites which obtain only a portion of their total nutrition
from hosts are often referred to as semi-, half-, or partial parasites. Present knowledge
concerning the role of parasitism in the life of such plants indicates that although
they may persist for a limited time without hosts, parasitism is necessary for optimum
development and sexual reproduction.

1962] Dasistoma — Piehl 333

followed Pennell who treated it as a monotypic genus, Dasis-
toma.

Dasistoma macrophylla is a robust, yellow-flowered herb
becoming 1-2 m. tall, and bearing large, dissected or pinna-
tifid lower leaves and narrower upper leaves which are mere-
ly toothed. Although it is described as an annual (Pennell,
1928; Gleason, 1952), the plants studied appeared to be
biennial, or perhaps even perennial. Its roots are much
larger than those I have examined for any of the truly
annual parasitic Scrophulariaceae. Young, rosette-forming
plants were found in the autumn that would presumably
develop into flowering individuals the following summer, as
I have noted for the related Awreolaria pedicularia (L.)
Raf. The common name bestowed upon Dasistoma, mullein-
foxglove (Fernald, 1950), is probably less well known and
certainly more confusing than the scientific name.

This species is limited to the central states, ranging from
northwestern Georgia to northeastern Texas, north to east-
ern Nebraska, southern Wisconsin, and northern Ohio. The
observations discussed here were made in October, 1959 in
Mercer County, Kentucky, 5 miles northeast of Shakertown
near U.S. Route 68, in hilly, conifer-hardwood woodland
over limestone. The associated species apparent at that sea-
son were Juniperus virginiana L., Ostrya virginiana (Mill.)
K. Koch, Quercus cf. muehlenbergu Engelm., Q. rubra L.,
Carya sp., Ulmus americana L., Celtis tenuifolia Nutt., Cer-
cis canadensis L., Acer saccharum Marsh., Fraxinus quad-
rangulata Michx., Panicum sp., and Aster shortii Lindl.

Parasitic connections were detected to the roots of two
species: American elm (Ulmus americana) and sugar maple
(Acer saccharum), with additional connections to what was
probably a third, unidentified, woody host. Since numerous
haustoria are formed by a single plant, simultaneous connec-
tions to two or more different hosts is the rule (Figs. A and
C). Self-parasitic connections to the roots of the same or
other individuals of Dasistoma were almost as frequent as
those to foreign roots. In addition to their surprisingly high
frequency (other root parasites have far fewer), some of
the self-parasitic connections were unusual in that they
lacked a distinct haustorium, the connection being suggestive

334 Rhodora [Vol. 64

of simple root grafting. Perhaps further observations will
determine whether such occurrences, which may be of con-
siderable ecological significance, are common in such plants.

The root system of Dasistoma resembles that of many
parasitic Scrophulariaceae in that it is whitish or light tan
when fresh, but will blacken quickly upon drying. The usu-

7
B trimit

8
|

9
|l

|
HH

Plate 1272

Figures A-C. Parasitic connections of Dasistoma macrophylla. A. Haustoria of a
single plant on roots of American elm (e) and sugar maple (m). Both large and
relatively small haustoria are visible at the right on maple (pointers); near the
center, three haustoria produced in rather close successsion on elm are indicated. At
the right the crown and large roots of the parasite are concentrated about roots of
maple. B. Dasistoma root (above) paralleling a maple root to which primarily single
haustoria have attached (pointer). Another haustorium is visible at the extreme right
(parasite root has been severed just beyond). C. Haustoria of a single Dasistoma root
attached both to elm and maple roots. Distal to the point of attachment, a marked
decrease in the diameter of the maple root is visible, X 2.5

1962] Dasistoma — Piehl 335

ally disc-like haustoria, which are commonly hemispherical
in vertical outline and from circular to broadly oblong in
surface view, ranged from 0.5-6.0 mm. in width.

Dasistoma haustoria differ rather markedly from those
of such members of the Euphrasiae as Melampyrum lineare
Desr. (Piehl, 1962) and Pedicularis canadensis L. (Piehl,
in press) in the following ways. They are larger on the
average and lack the epidermal hairs found on some haus-
toria of these species. Also, they are produced from large
as well as fine roots, and more frequently form attachments
to larger (diameters up to 7 cm.) host organs. The larger
haustoria-bearing roots, particularly, remain intact during
excavation, making the observation of connections consider-
ably easier. Another distinctive feature involving an abrupt
decrease in the diameter of maple roots distal to the haus-
torium was noted occasionally (Fig. C) ; in an extreme case
the development of the distal portion of roots 2.5 mm. in
diameter had been arrested completely so that the host root
appeared to terminate at the haustorium. The crown and
large roots of Dasistoma differ from the above Euphrasiae
in being concentrated near rather large host roots (Fig. A),
the parasite roots sometimes tending to partly surround
them in a manner which has its most extreme expression
(in temperate North America) in certain non-green para-
sites, e.g., Epifagus. The haustoria tend to simulate those
of the Euphrasiae in the mode of attachment to and pene-
tration of host organs, and also in being produced both
singly and in close succession.

It is the author’s opinion that further study would show
that a number of other species, the majority of them woody,
serve as hosts as well. It remains to be determined if her-
baceous plants also act as hosts, and if so, whether the
parasite would grow as well if restricted to such plants as
grasses, a situation which would involve smaller, but perhaps
more numerous haustoria. — SANTA BARBARA BOTANIC GAR-
DEN, SANTA BARBARA, CALIFORNIA.

LITERATURE CITED
BENTHAM, G. 1846. In De Candolle, Prodromus . . ., Vol 10. Paris.
FERNALD, M. L. 1950. Gray's Manual of Botany, 8th Ed. American
Book Co., New York.

336 Rhodora [Vol. 64

GLEASON, H. A. 1952. The New Britton and Brown Illustrated
Flora, Vol. 3. The New York Botanical Garden, New York.
NuTTALL, T. 1818. The Genera of North American Plants, Vol. 1.
The author, Philadelphia.

PENNELL, F. W. 1928. Agalinis and allies in North America, I.
Proc. Acad. Phila. 80: 339-449.

1935. The Scrophulariaceae of eastern temperate
North America. Monogr. Acad. Phila. 1: 1-650.

PrEHL, M. A. 1962. The parasitic behavior of Melampyrum lineare
and a note on its seed color. Rhodora 64: 15-25.

RAFINESQUE (-SCHMALTZ), C. S. 1819. Jour. de Phys. 89: 99.

1836. New Flora and Botany of

North America ..., Vol. 2.

HETEROMORPHIC POLLEN GRAINS IN POLYMNIA'

T. RICHARD FISHER AND JAMES R. WELLS

During the course of a biosystematic study of the genus
Polymnia, it was discovered that one population (Wells 254)
of P. laevigata Beadle contained an extremely high percent-
age of different pollen shapes and sizes. This collection was
taken from a population 5 miles southeast of Monteagle,
Tennessee on U. S. Route 41. The plant was transplanted to
the greenhouse and all pollen samples were taken from the
living plant. This station was made known from previous
collections of Ford and Russell for the University of Ten-
nessee in 1946. Flowering heads were preserved in 3 parts
95% ethyl alcohol and 1 part acetic acid. The immature
pollen was stained with aceto-carmine and studied for over-
all shape and number of nuclei. Aniline blue in lactophenol
was used to study stainability of the mature pollen grains.
Voucher slides are on deposit at The Ohio State University.

RESULTS

From observation of several hundred pollen grains, five
morphological types were easily discerned as follows:

1Publication 675, The Department of Botany and Plant Pathology, The Ohio State
University, Columbus 10, Ohio.

1962] Polymnia — Fisher and Wells 337

1. uninucleate and spherical (fig. 2, 3, 5)

2. binucleate and elliptical (fig. 4)

3. trinucleate and kidney bean-shaped (fig. 3)

4. trinucleate and ovoid (fig. 4)

5. quadrinucleate and spherical (fig. 2, 5)

The above morphological forms were placed in four
groups based on the number of nuclei in each pollen grain
(table 1). In a sample of 500 randomly selected pollen
grains, the uninucleate condition had a frequency of 68%

Plate 1273

Figures 1-6. Photographs of Polymnia laevigata showing plant and various pollen
forms. Fig, 1. Immature plant. Figs. 2-5. "Variable nuclear condition and mor-
phologic forms of pollen. Fig. 6. Mature pollen, large and small spheres. Large
spherical pollen grains in figures 2, 5, and 6 are 25g in diameter; small uninucleate
spherical forms are 15u in diameter.

338 Rhodora [Vol. 64

and the grains were about 15, in diameter. This is the con-
dition normally observed in other populations of this species.
The trinucleate condition was the least observed condition,
only 1.4% of the sample.

It is a common practice in taxonomic studies to use pollen
stainability using aniline blue and lactophenol, as an index
of pollen viability. Since this stain often obscures nuclear
detail, rendering an exact count of nuclei uncertain, the pol-
len was grouped into two categories; namely, “large” and
“small.” The “large” group included quadrinucleate and
trinucleate forms. The “small” group included the uninucle-
ate and binucleate forms. There were a few instances in
which the size of the pollen and its orientation made group-
ing difficult but familiarity with the material was thought to
have avoided such miscalculation. The results of this treat-
ment are shown in table 2.

From a random sample of 500 pollen grains, 77.5% fell
into the “small” group and proved to be 94% stainable. The
large pollen grains represented by 22.5% of the total sample
were 88% stainable. In other words, the frequency of stain-
ability among the smaller pollen grains was higher than for
the larger. Ordinarily, when such a stainability test is
applied to large and small pollen, the smaller pollen usually
fails to become stained.

DISCUSSION

Dimorphie pollen has been reported previously. Lee
(1961) in Tripogandra grandiflora observed two types of
pollen from the same flower; namely, hemispherical, fertile
grains and elongate, sterile grains. Erdtman (1952) noted
two pollen shapes in two different plants of Primula farin-
osa. Punnett (1923) reported round pollen versus long
pollen in different plants of Lathryus odoratus. Clark
(1940) observed variation in the number of nuclei in pollen
of Zea mays.

In P. laevigata, both anomalies are combined within the
same plant: that is, variation in the number of nuclei and
the shape of pollen grains. Moreover, there are not just 2
shapes of pollen (dimorphism) but 4 distinct shapes.

1962] Polymnia — Fisher and Wells 339

This assemblage of pollen forms coupled with variation
in number of nuclei is termed “heteromorphic pollen.” To
the authors’ knowledge, this term has not been used previ-
ously. No meiotic abnormalities were observed using the
aceto-carmine squash technique. The gametic chromosome
number is n — 15. First generation hybrids have been ob-
tained involving P. laevigata and P. canadensis but only one
hybrid resulted from 38 attempted crosses using this plant
(Wells 254) as the pollen parent.

Two reasonable explanations can be proposed to account
for this heteromorphic pollen. First, failure of wall forma-
tion at the completion of meiosis II would account for a
pollen grain with four nuclei. Second, and we believe more
plausible, an erratic precocious division of the microspore
nucleus. This population of Tennessee plants will be revisit-
ed and an attempt made to secure more material for a
thorough developmental study.

SUMMARY

One population representative of P. laevigata was found
to contain 4 kinds of pollen grains in respect to number of
nuclei and four kinds in respect to shape. If large and small
spheres are considered separately, there are 5 kinds of pollen
grains. The smaller sized pollen were uninucleate ; the larger
quadrinucleate. A term suggested for this type of pollen
found on the same plant is “heteromorphic pollen." Although
no explanation for this condition is known, two theories are
proposed ; namely, failure of wall formation at the termina-
tion of meiosis II and erratic precocious divisions of the
microspore nucleus.

LITERATURE CITED

CLARK, F. J. 1940. Cytogenetic studies of divergent meiotic spindle
formation in Zea mays. Amer, Jour. Bot. 27: 547-558.

ERDTMAN, G. 1952. Pollen morphology and plant taxonomy. Chron-
ica Botanica Co. 539 p.

LEE, R. E. 1961. Pollen dimorphism in Tripogandra grandiflora.
Baileya. 9(2) : 53-55.

PUNNETT, R. C. 1923. Linkage in sweet pea (Lathyrus odoratus).
Jour. Gen, 13: 101-125,

340 Rhodora [Vol. 64

Table 1. The number and frequency of nuclei per pollen
grain from a random sample of 500 grains.

number frequency
BEEN Loss 940. (68% )
Dc ccccsvssesussussussusivsnesuesucsuciesnsstesussussucecsuesusssecseesesuesuecteanenentens 58 (11.6%)
"NNNM 7 (14%)
A cseescessenncoveacsassstssusspuntnssisesutvapussinsnosserpnsseassoeansescaneastatoseensis 95 (19%)
Total 500 (100%)

Table 2. The stainability and frequency of small and large
pollen from a random sample of 500 grains.

stainabi lity f requ ency
small pollen m MEM

+ stain ..eeseeee emen ne nennen nnne nennen nenne 366 (94% )

— stain essen nme Henne en nn nnn nennen nnn nnne ens 23 ( 6%)

Total 389 (100% )

large pollen

+ stain .eeeee emen nmn nennen n hhnneeen enne innen 98 (88.3% )

— Stain eene eene nnn nnne nennt enne nennen nnns 13 (11.7%)

Total 111 (100% )

CHANGES IN FLORA OF THE MACHIAS SEAL
ISLANDS

RADCLIFFE B. PIKE AND ALBION R. HODGDON' *

Machias Seal Islands, at the entrance to the Bay of Fundy,
have occupied a prominent place in ornithological literature
ever since 1603 when they were named Les Isles des Perro-
quets or the Parrot Islands by Champlain. A number of
studies were made here by Audubon, and at the present time,
a bird sanctuary is maintained by the United States Govern-

IThis research is part of a project entitled “Floristic and Phytogeographic Investi-
gation of the Wolf Islands and Other Islands in the Bay of Funly," which was sup-
ported in part by a grant from the Central University Research Fund of the
Graduate School of the University of New Hampshire and in part by a grant from
the Society of the Sigma Xi.

Published with the approval of the Director of the New Hampshire Agricultural
Experiment Station, as Scientific Contribution No. 300.

?Department of Horticulture and Botany, University of New Hampshire, Durham,
N. H.

1926] Machias Seal Islands — Pike and Hodgdon 341

ment. Observations of plant life on these islands, when
made, seem to have been buried in ornithological writings
or have gone unpublished, as practically nothing appears
in botanical journals or records.

As part of a phytogeographic survey of Islands of the
Bay of Fundy, a short visit was made by the senior author
to the Machias Seal Islands on 29 August 1960. The islands
are hardly more than ledges, with only the larger one hav-
ing any real soil or obvious terrestrial vegetation other than
lichens and a few halophytic plants of rock-crevices.

These islands lie about ten miles off Cutler, Maine, and
can be reached by small boats when the “Bay” is reasonably
calm. Botanizing is more pleasant after the nesting season
is over and the birds have departed sometime in August.
Only the larger island was visited, the smaller one, called
Gull Rock, being practically inaccessible; however, observa-
tions were made from the boat with binoculars. No trees or
evidence of any trees nor any woody plants of any sort oc-
cur on these islands, not even the matted or prostrate shrubs
such as Juniperus horizontalis, Empetrum nigrum, or Vac-
cinium Vitis-Idaea. Herbaceous plants in great vigor, how-
ever, occupied every available niche in 1960. Asters in spec-
tacular profusion, in full bloom at the time, dominated the
flora of the island and were easily the most conspicuous ele-
ment of the vegetation. Many of the plants stood breast-
high and individual clones varied from six to twenty feet
across. The flowers were mostly in shades of blue and violet
with some plants varying to pinkish or nearly white. Sen-
tinels of matured Coelopleurum and Rumex obtusifolius
were scattered throughout, with clumps of yarrow, the foli-
age so gigantic as to be almost unrecognizable. Plants of
Iris versicolor with leaves six feet tall were noted. The extra-
ordinary vigor and lushness of the herbage doubtless was
due to the continual rain of guano during the early growing
season, combined with the high humidity of the Fundian
atmosphere. The only other conspicuous kind of vegetation
was the well-kept lawn near the lighthouse which, at the
time of the visit, was carpeted with euphrasias in full flow-
er. During the short stay on the island on August 29, a total

342 Rhodora [Vol. 64

of 34 species was collected as shown in the accompanying
list.

The following year, a chance clue led to an exchange of
communication with Dr. R. T. Clausen, who called our at-
tention to a record of collections and a survey made in the
years 1947-48 by Oscar Hawksley (Bird-Banding XXVIII
No. 2 April 1957) as part of an investigation of the nesting
habits of the Arctic Tern. In two years, 41 species of plants
were collected. Hawksley stated on page 4, “an attempt was
made to collect all the species of vascular plants on the is-
land." Since his list was remarkably different in many re-
spects from our list of 1960, a further survey was made to
reconcile the two. On September 17, 1962, therefore, the
senior author, this time with two experienced assistants,
carefully combed the island. At the same time, a complete
sampling of the aster clones was made for a later population
analysis.

Sixty-two species as well as other minor taxa were col-
lected, a marked increase over either previous list, but
twelve species of Hawksley’s list were still missing. The
obvious procedure at this point would be to carefully study
Hawksley’s herbarium specimens, which we were informed
were deposited in the Wiegand Herbarium at Cornell Uni-
versity. Unfortunately, the Curator, Dr. Clausen, reports
that these are missing or perhaps were not left there. Not
wanting to postpone this paper indefinitely, we prefer to
proceed with our discussion of the Machias Seal Islands now,
and in the event that this collection does come to light,
make any necessary corrections in the list at that time.

The most noteworthy fact about the Machias Seal Island
flora is the remarkable difference over an eighteen year pe-
riod, which may be explained by any of the following rea-
sons. The lists may be imperfect, resulting from fragmen-
tary collecting or collecting only at one season. On the other
hand, weed species may have come and gone or new ones
arrived recently on an island that is so much visited. The
climatic factors of one year may provide different growing
conditions from another. The effect of domestic animals
may have been most drastic, particularly in earlier years.

1926] Machias Seal Islands — Pike and Hodgdon 343

The interaction of the plants and the usually heavy breeding
population of birds may result in periodic fluctuations of
dominance of certain plant species.

Hawksley lists twelve species that we have been unable
to find, while we are reporting 36 species not found by him.
Several of Hawksley’s twelve are species that might be ex-
pected, Carex brunnescens, C. silicea, Urtica dioica, Iris
Hookeri, Rumex crispus, Lathyrus palustris, and Euphrasia
canadensis, for example. But it is to be noted that a speci-
men of Urtica viridis Rydb. collected by Mrs. A. H. Norton
is present in the Herbarium of the New England Botanical
Club. Certainly these two nettles might easily be confused.
It is also of interest that A. H. Norton (Rhodora 15:138,
1913) in recording observations of Iris Hookeri, made the
following comment: "this plant ..... abounds on most of
the islands (excepting Machias Seal Island) east of Petit
Manan Point" A particular search for this species was
made both in 1960 and 1962, but without success. Of
course, it may exist in a precarious state of balance with
other more vigorous vegetation. Rumex mexicanus may
easily be confused with R. pallidus. Our material in good
fruit proves to be the latter. Cerastium vulgatum was col-
lected on both recent visits and occurs there in some abun-
ance, and it could be that C. viscosum was confused with it.
Although similar in general appearance, the common plant
of seacoasts and low marine islands is Potentilla Egedei var.
groenlandica rather than P. anserina. Coelopleurum luci-
dum, when immature, might be confused with Heracleum;
we have not seen it on other small islands in the Bay of
Fundy where Coelopleurum is common. The much greater
list of 1962 may be explained in part by the advantages of
getting more species in maturity and at a time when birds
did not interfere with collecting and with the growth and
development of plants.

The phenomena of plant succession and vigor were close-
ly observed by Hawksley as they seemed to have direct bear-
ing on his population study of Artic terns. He noted a num-
ber of vegetational changes, some of the more violent of
which were influenced by man and his domestic animals.

344 Rhodora [Vol. 64

In 1944 there were 40 sheep on the island. In Hawksley’s
words (p. 85), “the vegetation ..... was close-cropped and
much of it was destroyed completely." Recovery was ap-
parent by 1946, and the following year the plant growth
reached a height of 40 inches. A decline started in 1947
again with the aid of sheep (only four) and continued into
1949 with Rumex Acetosella and grasses replacing much
yarrow.

Curiously enough, little mention is made of asters, which
eleven and thirteen years later were the dominant herbage
on much of the island. Even though asters would not have
been in bloom during the months of Hawksley’s visits, yet
it would have been hardly possible to have ignored them
when preparing a detailed vegetation map such as he made
(Fig. 2, p. 63). He collected only one aster, and this species
appears to be the minor one in the aster complex which was
such a major feature in 1960 and 1962. One can only specu-
late as to what led to the appearance and rapid dominance
of these asters.

Certain conclusions from this study bearing on island
flora seem to be warranted. Even on a limited and isolated
island, the composition of the flora under some conditions
can apparently change to a considerable degree in a matter
of only a few years. As a result of this work, one is led to
wonder just what constitutes a definitive flora. The dynam-
ics of floristic and vegetative change along with the uncer-
tainty of making entirely complete collections show the dan-
ger involved in assuming a “flora” to be finished.

All specimens collected on the two visits in 1960 and 1962
are to be found in the Herbarium of the University of New
Hampshire.

CONSOLIDATED FLORA OF MACHIAS SEAL ISLAND HAWKSLEY PIKE PIKE
1947-48 1960 1962

Osmunda cinnamomea L. X X
Festuca rubra L. var. rubra X X
Puccinellia paupercula (Holm) Fern. & Weath.

var. alaskana (Scribn. & Merr.) Fern. & Weath. X
Poa annua L. X X
Agropyron repens (L.) Beauv. var. subulatum

(Schreb.) Reichenb. X X

1926] Machias Seal Islands — Pike and Hodgdon 345

Agropyron repens var. subulatum forma
Vaillantianum (Wulf. & Schreb.) Fern. X
Agropyron repens var. subulatum forma
setiferum Fern.
Elymus arenarius L. var. villosus Mey. X
Calamagrostis canadensis (Michx.) Nutt. var.
canadensis
Calamagrostis canadensis var. robusta Vasey
Agrostis alba L.
Agrostis alba var. palustris (Huds.) Pers.
Phleum pratense L.
Carex canescens L. var. canescens
Carex brunnescens (Pers.) Poir.
Carex scoparia Schkuhr
Carex silicea Olney
Carex hormathodes Fern.
Carex paleacea Wahlenb.
Juncus bufonius L.
Juncus bufonius var. halophilus Buchenau & Fern.
Sisyrinchium montanum Greene
Iris Hookeri Penny
Iris versicolor L.
Urtica viridis Rydb. collected by
Mrs. A. H. Norton, July 27, 1902 (NEBC).
Urtica dioica L.
Rumex mexicanus Meisn.
Rumex pallidus Bigel. X
Rumex orbiculatus Gray X
X

a

bd bM Mx
bd bd be
bd bébé bd bd bd db bd bd bet pd pd

p P rn

p P

Rumex crispus L.

Rumex Acetosella L.

Polygonum aviculare L. var. littorale (Link)
W. D. J. Koch X

Atriplex patula L. var. patula

Atriplex glabriuscula Edmondston X

Spergularia marina (L.) Griseb.
var. leiosperma (Kindb.) Gürke

Arenaria lateriflora L.

Stellaria media (L.) Cyrillo

Stellaria graminea L.

Cerastium vulgatum L. X

Cerastium viscosum L.

Ranunculus repens L. var. villosus Lamotte

Ranunculus acris L.

Thalictrum polygamum Muhl.

Capsella Bursa-pastoris (L.) Medic.

Sedum Rosea (L.) Scop.

Potentilla norvegica L.

p v

pA pr

PA PA
pA pA pA pA pA pd pA pA pA rA pA pA po pdo PA A

pP do XM

346 Rhodora [Vol. 64

Potentilla anserina L. X
Potentilla Egedei Wormsk.

var. groenlandica (Tratt.) Polunin X X
Trifolium pratense L. X
Trifolium repens L. X X X
Trifolium hybridum L. X
Vicia augustifolia Reichard X X
Vicia Cracca L. X X
Lathyrus palustris L. X
Callitriche verna L. X
Impatiens capensis Meerb. X X X
Viola Mackloskii Lloyd subsp. pallens

(Banks) Baker X X
Carum Carvi L. X X
Ligusticum scothicum L. X X X
Coelopleurum lucidum (L.) Fern. X X
Heracleum maximum Bartr. X
Lycopus uniflorus Michx. X X
Euphrasia Randii Robins. X
Euphrasia canadensis Townsend X
Euphrasia americana Wettst. X X
Rhinanthus Crista-galli L. X X X
Plantago major L. X X
Plantago juncoides Lam. var, decipiens

(Barneoud) Fern. X X
Galium tinctorium L. X X
Aster foliaceus L. (complex) X X
Aster johannensis Fern. X X X
Aster umbellatus Mill. X
Gnaphalium uliginosum L. X
Achillea borealis Bong. X
Achillea lanulosa Nutt. X X
Matricaria matricarioides (Less.) Porter X X
Cirsium arvense (L.) Scop. X X
Cirsium arvense forma albiflorum (Rand & Redf.)

R. Hoffman X
Hypochoeris radicata L. X
Leontodon autumnalis L. X
Taraxacum erythrospermum Andrz. X
Taraxacum officinale Weber X X
Sonchus asper (L.) Hill X

1962] Book Review — Tryon 347

THE FERNS AND FERN ALLIES OF CHIHUAHUA.!, — This
attractively prepared volume merits special attention for it
is the first comprehensive modern treatment of the rich
pteridophyte flora of Mexico. The authors have both collect-
ed extensively in the state of Chihuahua, with the particular
purpose of augmenting the previous collections and phyto-
geographic knowledge. Together, they have collected all but
18 of the species. They are to be congratulated on the
thoroughness of their work and the accurate presentation of
their results.

Although 120 species are known from Chihuahua, and an
additional 6 from adjacent areas are treated, the authors
recognize that the fern flora is still incompletely known.
Additional species will be discovered as collecting continues
in many less accessible areas, especially the isolated mesic
barrancas of the western Sierra Madre Occidental. How-
ever, the basic explorations have been completed. The
authors are to be commended in bringing out their work at
this time so that it may fill a long-standing need.

The introduction surveys the collecting activities in the
state of Chihuahua, the vegetation, geology, physiography,
climate and the ecology and distribution of the pteridophy-
tes. There is also a discussion of the typical structure and
life-history of ferns, their cultivation and economic uses. A
list of the known chromosome numbers of Chihuahuan
species is a useful supplement. The principal text presents
the pertinent nomenclature, a careful and complete descrip-
tion and the ecology and distribution of the 138 taxa recog-
nized. Each species is illustrated; a number of them
adequately for the first time. Keys are provided as an aid
in identification and these, in combination with the excellent
illustrations and the frequent discussions of characters make
the book one of unusual utility. The appendix is devoted to
comments on the species to be expected in Chihuahua, a

JFerns and Fern Allies of Chihuahua, Mexico, by Irving W. Knobloch and Donovan
S. Correll (illustrated by Phoebejane Horning and Jane Roller) i-xiv, 1-198, t. 1-57.
(Contrib, Texas Res, Found, vol 3) Texas Research Foundation, Renner, Texas.
1962. $10.00,

348 Rhodora [Vol. 64

gazetteer, a glossary of terms, a bibliography and an index.
The gazetteer will be useful to many botanists concerned
with the location of often obscure place-names in Chihuahua.

From my own point of view, it is refreshing that the
classification adopted is a highly realistic one. I refer par-
ticularly to the recognition of the Polypodiaceae in the broad
sense, to the generic treatment of the Cheilanthoid ferns and
to the use of the category variety (rather than species) for
important but subordinate taxa. Such practice is a depar-
ture from the tendency of many authors to accept, uncritic-
ally, some of the more recent but unsubstantiated views on
classification.

The pteridophyte flora of Chihuahua, as one expects, is
primarily a xeric one. This is emphasized by the predomi-
nance of the genera characteristic of arid lands in the
Americas. Cheilanthes is represented by 23 species, Notho-
laena by 15, Selaginella by 11 xeric ones and Pellaea by 9.
The truly mesic element consists of about 12 species and it
is confined to the barrancas of the Sierra Madre Occidental.
This is a tropical element which mostly reaches its north-
western limit in Chihuahua. Representative species of this
element are: Dennstaedtia distenta, Trichomanes radicans,
Hymenophyllum tunbridgense, Adiantum Poiretii, Thelyp-
teris rudis and Dryopteris parallelogramma. This book will
be useful for a considerably larger area than the single state
it covers. Fifty-nine of the species also occur in southern
California, Arizona or New Mexico and 110 of them occur
elsewhere in Mexico. Of these, 85 extend southward to
central Mexico and 61 to southern Mexico or beyond. Eleven
species are known in Mexico only from Chihuahua. Five of
these are endemic and the remainder occur in the United
States.

THE FERNS AND FERN ALLIES OF CHIHUAHUA is an authori-
tative treatment and it can be recommended to all botanists
with an interest in ferns or with a broader interest in the
flora of arid regions. It will also be an important reference
for those concerned with any portion of the flora of Mexico
or the southwestern United States. — ROLLA M. TRYON, GRAY
HERBARIUM, HARVARD UNIVERSITY.

1962] X Cyperus Weatherbianus — Raymond 349

A NOTE ON X CYPERUS WEATHERBIANUS — Early in
1962, Mr. Richard J. Eaton, a member of the Club Commit-
tee on Plant Distribution which is preparing reports on the
distribution of Cyperaceae in New England, asked me while
I was at the Gray Herbarium to examine the type of X Cy-
perus Weatherbianus Fern., a putative intergeneric hybrid
between Cyperus dentatus Torr. and Rhynchospora capitel-
lata (Michx.) Vahl. This plant had been puzzling me for a
long time, for Cyperus and Rhynchospora belong to different
tribes. Other cyperologists had some doubt too. For in-
stance, in his monograph of the genus Cyperus (Pflanzen-
reich IV. 20(Heft 101): 251. 1936) the late G. Kükenthal
commented as follows:

*Nota. C. Weatherbianus Fernald in Rhodora XX. (1918) 190, t.
CXXV, fig. 1-5 a cl. Fernald pro prole hybrida bigenerea, nempe
Cyperus dentatus Torr. X Rhynchospora capitellata Vahl sumptus
mihi dubius esse apparet, parentibus indicatis in systemate Cy-
peracearum admodum distantibus. Sed specimina originalia non vidi.

Crescit in Massachusetts: Sandy shore of Simmons Pond (C. A.
Weatherby, Fernald und Long n. 16287) ."

A close look at the type of X Cyperus Weatherbianus
Fern. shows that this plant has very little, if any, Cyperus
influence in its make-up. First, the stem is nodose, and the
scales of the fascicled spikelets are three ranked. These
two characters eliminate the genus Cyperus, which has only
basal leaves and two-ranked spikelets. After studying all
the possibilities, I am convinced that the plant is the result
of a cross between Rhynchospora capitellata (Michx.) Vahl
and Dulichium arundinaceum (L.) Britton, Rhynchospora
having provided the three-ranked colored scales, whereas
Dulichium contributed the shape and the distribution of the
fascicles emerging from the leaf-sheaths of the stem, as well
as the subulate tubercle of the achene.

This finding is of great phylogenetic importance, for the
real position of the genus Dulichium in the family Cyper-
aceae is still a debatable point. Whereas Kiikenthal (Bot.
Jahrb. 75: 485-488. 1952) placed it amongst the Rhyncho-
sporoideae, Fernald (Gray’s Manual of Botany, ed. 8. 248)
squeezed it between Cyperus and Eleocharis, and Schultze-
Motel (Willdenowia 2(2): 173. 1959) created a new tribe,
Dulichieae, to accommodate this unusual monotypic genus.

550 Rhodora [Vol. 64

This new evidence shows that Dulichium rightly belongs to
the Rhynchosporoideae. It is strange that the late Professor
Fernald after having originally described his intergeneric
hybrid in 1918 with the scales spirally arranged, changed
his mind in 1950 and in the Gray’s Manual stated clearly,
"the spikelets with many distichous scales." They are de-
cidedly three ranked.

I do not think it necessary to transfer X Cyperus Weath-
erbianus to any other genus, for only one tussock has been
found, that from Cape Cod. It is sufficient that we assume
the probable correct identity of this odd individual and
emphasize its phylogenetic importance. It is a chance, sterile,
intergeneric hybrid that will probably never duplicate itself,
so the problem of giving it a distinct name does not arise. —
MARCEL RAYMOND, MONTREAL BOTANICAL GARDEN.

AN ALBINO FRUITED FORM OF VACCINIUM MAC-
ROCARPON. — A colony of albino berried plants was
found on the west shore of Tom Nevers Pond, Nantucket
Island, Massachusetts, on September 11, 1958. Two sets of
specimens were collected of this plant, which were assigned
the collection number N2. Because of the difficulty of getting
into the area, and due to a previous coronary, I have never
returned to it. The small colony extended from the shore
into a large area covered with plants bearing very dark red
berries of large size. The albino berries were about three
quarters of the size of the red berries. The set of fruit was
about equal. There was a definite contrast between the plants
due to the lighter foliage and the white berries of the albino
plants. Thus, from high ground, these plants appeared to be
massed in such a way as to resemble a slice of pie, the apex
being furthest from the shore. The albino berries were near-
ly spherical, whereas the red berries were decidedly oblong.
A few of the larger and older berries had a tendency to form
“a blush" on their surfaces which were more exposed to the
sun.

When I was certain that this was definitely a colony of
albinos, I envisioned marketing the red and white berries

1962] Albino Vaccinium — MacKeever 351

packaged together. An eye pleasing and picturesque product,
to say the least. However, sauce made from the berries was
of an amber color, insipid, and much like the flavor of a sour
blackberry. (And — I know how to make a delicious cran-
berry sauce!)

On August 31, 1962 I had the good fortune to find another
colony of these albinos. Some of the fruits were almost pure
white but the majority were of a pale, cream color. The
fruits on the uppermost portions of the plants, unlike those
found previously, rarely showed any signs of a “blush.” The
prominent remains of the calyces were persistent on the
somewhat constricted blossom ends of the berries — not
fugacious as in most cranberries. Specimens from this colony
will be distributed under my collection number N695.

This white berried plant is an albino fruited form of
Vaccinium macrocarpon Aiton, and it should not be confused
with the white berried V. oxycoccos L., forma leucocarpum
Ascherson and Magnus.

Vaccinium macrocarpon Aiton, forma eburnea MacKeever. Differt ab
planta typica in fructis eburneis. This form is similar to the species
except for its albino fruits. The type specimen, N2, was collected by
Frank C. MacKeever, at Tom Nevers Pond, Nantucket Island, Mass.,
on September 11, 1958, and is deposited in the herbarium of the New
York Botanical Garden. An isotype is in the herbarium of the Museum
of Natural Science, Nantucket, Mass. — FRANK C. MACKEEVER, NEW

YORK BOTANICAL GARDEN.

JUNIPERUS HORIZONTALIS IN NEW HAMPSHIRE.

A colony of this plant occurs at Holt’s Ledge in Lyme,
Grafton County. This was called to our attention by Pro-
fessor James Poole of Dartmouth College with whom we
had the pleasure of visiting the area on June 23, 1959.
Professor Poole does not know who discovered this colony.
He collected specimens from it at the time of his first visit
to the station on October 21, 1937, but he informs us that
there are records of collections by earlier botany classes
as far back as 1930. On our June, 1959, visit we were able
to find the juniper again after brief searching.

Only 2 small mats of it were seen; neither one more than

352 Rhodora [Vol. 64

3 or 4 feet across, and each probably consisting of only a
single plant. They were growing on shelves near the crest
of the high and steep ledge. It is possible that there were
other remnants of this meager colony farther down on more
inaccessible shelves of the ledge, but we were satisfied, for
the time being at least, to find these small plants. This
colony has more than passing interest because, aside from
the well known station on Mt. Equinox in southwestern Ver-
mont, it is the only known inland station in New England,
but unlike the Vermont locality there seemed to be no
calcareous rock in the Holt’s Ledge area, nor any plants
present that might be thought of as of calcareous affinity.

Specimens have been deposited in the Herbaria of the
University of New Hampshire, Dartmouth College and the
New England Botanical Club.

This is the first verified record of this plant from New
Hampshire. Bean, Hill and Eaton (Rhodora 63: 348) cor-
rectly excluded it from the state on the basis of the then
available published information. — A. R. HODGDON and F.
L. STEELE, UNIVERSITY OF NEW HAMPSHIRE, DURHAM AND ST.
MARY'S-IN-THE-MOUNTAINS, LITTLETON, NEW HAMPSHIRE.

CYPERUS FERRUGINESCENS IN VERMONT. In
early September, 1961, I collected a Cyperus on the grassy
shore of the Connecticut River in Vernon, Vermont, at the
southeast corner of the state. Subsequently I referred it to
C. ferruginescens Boeckl. This species is rare in New Eng-
land and hitherto known in New England only from three
riparian stations near Hartford, Connecticut. There were
no specimens in the Club Herbarium (N.E.B.C.) and only
four sheets in the Gray Herbarium. My collection, Eaton
5088, at Vernon, Windham County, Vermont, September 7,
1961, represents a significant extension of range, north ward.
Mr. F. C. Seymour and Dr. Marcel Raymond have kindly
examined it and concur in the determination. It has been
placed in the herbarium of the New England Botanical Club.

There is some question whether C. ferruginescens deserves

1962] Cyperus Ferruginescens — Eaton 353

specific rank or is better treated as a variety of C. odoratus
L. In New England it seems distinct morphologically as
well as in respect to range and habitat preference. Here it
appears to be confined to alluvial soils well removed from
salt water, whereas C. odoratus is generally found in saline
or brackish situations, and only along the coast as far
northeastward as Essex County, Massachusetts. Elsewhere
in North America the former is conspicuously inland in its
distribution, whereas the latter is primarily coastal as far
west as the mouth of the Mississippi River. There is some
overlap in range northward along the edges of the Missis-
Sippi River valley and particularly beyond its western side
(South Dakota, Nebraska, Oklahoma, Texas, and thence
westward to Southern California). Furthermore, C. odor-
atus is semi-cosmopolitan (North and South America, east-
ern Asia), whereas C. ferruginescens is confined, I believe,
to North America. Unless close study has demonstrated
the existence of intergrading forms in the areas of overlap,
it seems better to maintain them as separate species. RICH-
ARD J. EATON, LINCOLN, MASSACHUSETTS.

AN UNUSUAL RUBUS. The subgenus Eubatus of the genus
Rubus contains such a variable assemblage of entities that
it is not unusual to find a plant that cannot be identified.
However, the blackberry to be described in this note pre-
sents some unusual features. On July 21, 1961 in the town
of Altamount, New York, I found a Rubus with an unusual
inflorescence in an old field near the edge of a bog. The
flowers were small, about the size of a blossom of R. hispi-
dus, but many were of a type often referred to as “double”,
with 10-15 petals of various sizes, apparently representing
altered stamens. This plant did not fit any blackberry des-
cribed in Gray’s Manual, Eighth Edition, but seemed to fall
in the section Tholiformes, and in the key came closest to the
group that included R. biformispinus, R. grandidens and R.
arcuans. However, it differed from all of these plants in
several ways. It seemed more probable that it was of hybrid
origin.

354 Rhodora [Vol. 64

In an effort to determine the putative parents, eight
samples of primocanes and floricanes were collected and
growth habit notes made for each one individually. Later
various measurements were made. The following character-
istics were noted: growth habit from trailing to doming to
erect up to 214’; armature consisting of stiff prickles, acicu-
lar prickles, bristles and glands, length of longest prickle
up to 5 mm., number of all types of armature combined from
200 to 800 per dm.; primocane leaflets uniformly dull, sub-
coriaceous, length from 5.5 to 7.0 cm., number of leaflets
ranging from 3 to 5, ratio of width to length from .65 to .8,
broadest part of leaf at the middle or well above the middle;
inflorescence resembling R. hispidus or R. setosus.

On the basis of these characters it was not possible to
form a positive conclusion as to the origin of the blackberry.
The leaflet shape and number seemed to be derived from R.
hispidus, the armature from R. elegantulus and R. setosus.
Other characters could have come from one or more of these
three species, all of which were in the vicinity. Extensive
field observations have convinced me that these species
hybridize frequently.

A specimen of this blackberry has been deposted in the
Herbarium of the University of New Hampshire.

FREDERICK L. STEELE, ST. MARY’S-IN-THE-MOUNTAINS,
LITTLETON, NEW HAMPSHIRE.

WESTERN PLAINS PLANTS IN NORTHERN ILLINOIS! — A
number of western plants are established on a gravel bluff
prairie, known locally as Bell Bowl prairie, in southern
Winnebago County, in Greater Rockford Airport a few miles
south of Rockford, Illinois. They can be divided into two
groups.

One group is composed of annual or perennial plants that
are being spread by seed and are found over the county and
over the state. Among them are such more or less common
species as Agropyron smithii Rydb., Froelichia gracilis Moq.,
Solanum rostratum Dunal, Artemisia dracunculoides Pursh

1Contribution of the Evelyn I. Fernald Memorial Herbarium of Rockford College.

1962] Plains Plants — Fell 355

and Ratibida columnifera (Nutt.) Wooton & Standl. They
are now widespread and their presence here does not require
any special comment.

The other group is made up of the following perennials
which have a number of peculiarities in common. Poa arach-
nifera Torr. is found elsewhere in the county in connection
with highway shoulder seeding (Fell, 1955) but on Bell
Bowl prairie it does not have the appearance of having been
planted nor of spreading except by stolons. Buchloé dacty-
loides (Nutt.) Engelm. (buffalo grass) is reported by Jones
(1958) as having been collected recently by V. H. Chase in
Peoria County 110 miles south of us. In Bell Bowl it occurs
in several patches each about 10 feet in diameter. It is
spreading by stolons. Chloris verticillata Nutt. (windmill
grass) is credited to the same collector from the same area
by Jones & Fuller (1955). With us it grows in a number of
patches none of which is more than two feet in diameter. It
produces viable seed abundantly but does not seem to be
spreading by this means. Carex praegracilis W. Boott and
C. stenophylla Wahl. var. enervis (C. A. Mey.) Kiikenth.
(C. eleocharis Bailey) have been previously commented
upon (Fell, 1958). They produce seed sparingly and seem
to reproduce only vegetatively. Schrankia nuttallii (DC.)
Standl. (sensitive briar) was collected by Francis McDonald
in the Peoria area in 1903 according to Jones & Fuller (l.c.).
On Bell Bowl prairie it produces flowers but no seeds were
found.

None of the above plants are recorded otherwise than as
mentioned from Illinois so the presence of all of them here
in a small area, a gravel bluff one half mile long, is a matter
of interest. This is particularly true because of their absence
from the numerous other prairie patches in the county that
furnish the same type of habitat (Fell and Fell, 1956), dry
gravel hills or bluffs exposed to the prevailing westerly
winds, which are from the north in winter and from the
south in summer. If they are natives they are relics of dying
populations. This is suggested somewhat by their poor re-
production by seed. It seems unlikely that all these plants
could have been growing in this vicinity a hundred years
ago and some of them not have been found by so active a

356 Rhodora [Vol. 64

collector as M. S. Bebb. Another possible explanation of
their being here is the fact that this area was within Camp
Grant during World War I and this particular tract was an
important remount station and trench warfare training cen-
ter. The numerous horses and mules were shipped here from
the west of the Mississippi River as was also hay and straw
in bales for use as bedding and in making ramparts for the
trenches and effigies for bayonet practice. These shipments
may have served as transports for the seed of these plains
plants. The ones that reproduce readily by seed here have
become disseminated, the others have stayed where they
were originally introduced. — EGBERT W. FELL.’

LITERATURE CITED
FELL, E. W. 1955. Flora of Winnebago County, Illinois. Nature
Conservancy, Washington, D. C.
1958. New Illinois Carex records. Rhodora 60:115-

116.
and GEORGE B. FELL. 1956. The gravel hill prairies of
Rock River valley in Illinois. Ill. Acad. Sci. Trans. 49: 47-62.
Jones, G. NEVILLE. 1958. Buchloé dactyloides in Illinois. Rhodora
60: 259-260.
and GEORGE D. FULLER et al. 1955. Vascular plants
of Illinois. Univ. of Ill. Press and Ill. State Museum.

?Dr, Egbert W. Fell died on July 16, 1960.

ADDITIONS TO THE FLORA OF ILLINOIS. — Field and her-
barium studies in Illinois continue to add to the number of
species known from that State. As has been the situation
for the past six or seven years, many of the additions are
native species rather than adventives or incidental waifs.

All specimens cited are deposited in SIU, except that of
Agrimonia microcarpa which is in MO.

Paspalum lentiferum Lam. A large colony of this southern grass
occurs in a semi-swampy roadside ditch near Karnak in Pulaski
County. Considerable similarity exists between this species and the
southeastern P. boscianum. The stramineous fruit distinguishes P.
lentiferum. The fact that the southern Illinois station is several
hundred miles from the nearest known station would seem to indicate
an adventive status for this species, although the marshy situation
simulates its natural habitat,

1962] Flora Illinois— Mohlenbrock, Ozment & Folkerts 357

Pulaski Co.: low roadside diteh, near Karnak, S. Boyce s.n.

Rhynchospora macrostachya Torr, This giant horned rush, which
closely resembles R. corniculata and under which name it was dis-
tributed, has its only station in a swamp north of Pulaski in Pulaski
County. The long bristles which greatly exceed the achenes readily
distinguish this species from R. corniculata. The nearest station to
the Illinois locality is in southern Missouri. Some of our specimens
attain a height of nearly one meter. Rhynchospora corniculata is
usually a taller species in Illinois.

Pulaski Co.: north of Pulaski, July 12, 1955, R. Mohlenbrock and
J. Voigt 5542.

Carex nigromarginata Schwein. This species belongs to the diffi-
cult $ Montanae, which is represented in Illinois by seven species.
Of these, Carex emmonsii, C. physorhyncha, and the present species
have been found within the last four years. Carex migromarginata,
because of its partially hidden spikelets and its persistent, shreddy
leaves, is very similar to C. umbellata and C. abdita. It is distin-
guished by its much narrower perigynia.

Union Co.: cherty soil, Pine Hills, W. Ashby & R. Kelting s.n.

Cydonia oblonga L. The quince is an Asiatic shrub which rarely
escapes from cultivation. On the top of Atwood Ridge in Union
County occur two shrubs of this species along a trail. Although no
other indication is evident of previous habitation, the area must have
been the site of an old homestead. The finding of these shrubs in an
otherwise wild area is surprising.

Union Co.: along trail, Atwood Ridge, May 27, 1957, Steyermark,
Voigt, & Mohlenbrock s.n.

Agrimonia microcarpa, Wallr. A collection of this species (labeled
A. platycarpa) from near Cahokia in the latter part of the nineteenth
century is the sole record of this species from Illinois. This specimen
apparently has been overlooked in the past. The Illinois station marks
the northwest limit of the range of this species. It closely resembles
A. parviflora, but is distinguished by its smaller fruits (up to 3 mm
long) and its fewer leaflets (5).

St. Clair Co.: near Cahokia, in 1877, H. Eggert s.n. (MO).

Polygala sanguinea L. f. albiflora (Wheelock) Millsp. The white-
flowered form of the common Polygala sanguinea occurs in a rather
large colony in a railroad prairie near Elkville. The "typical" form
is abundant in the same area.

Jackson Co.: railroad prairie, one-fourth mile south of Elkville
along Route 51, July 2, 1962, G. Folkerts s.n.

Euphorbia dictyosperma Fisch. & Mey. This western species was
found on a limestone ledge at the edge of a hill prairie near Fults in
Monroe County. Only a few specimens were observed. The beautiful,
intricately reticulated fruit distinguishes this species from the similar
E. obtusata.

Monroe Co.: limestone ledge, near Fults, J. Ozment s.n.

Vaccinium stamineum L. This species has been sought in southern

358 Rhodora [Vol. 64

Illinois for several years since it is known from Indiana and Missouri.
It finally has been found on a xeric sandstone bluff at Hayes Creek
Canyon, where it occurs with the very abundant Vaccinium arboreum.
The senior author was attracted to the two shrubs of V. stamineum
because of the much paler color of the foliage. Our specimens, with
the branchlets and leaves glabrous, belong to var. neglectum (Small)
Deam.

Pope Co.: sandstone blufftop, Hayes Creek Canyon, May 19, 1962,
R. Mohlenbrock 14914.

Infrequently Collected Southern Illinois Plants

Phragmites communis Trin. The most southern records of this
species in Illinois previously are from St. Clair County (Eggert in
1877) and Effingham County (Voigt in 1960). These stations are
over one hundred miles from the locality reported here. At this
Franklin County station, a huge colony exists.

Franklin Co.: around pond, near Christopher, August 1, 1962, C.
Bollwinkel & J. Richardson s.n.

Poa palustris L. A collection of this primarily northern species
from a low woodland in the Pine Hills of Union County marks the
first record from southern Illinois. The nearest station is in Fulton
County, over two hundred miles distant.

Union Co.: low woodland, Pine Hills, August, 1961, W. Ashby &
R. Kelting s.m.

Potentilla millegrana Engelm. This species, which is distinguished
from P. norvegica var. monspeliensis by possessing 10-15 stamens, has
been found in a disturbed woods in Jackson County. The only previous
record of this species from Illinois is a collection (H. Eggert in 1877)
from National Stockyards, St. Clair County.

Jackson Co.: open woods, Thompson’s Woods, Carbondale, August
3, 1962, G. Folkerts s.n.

Tragia cordata Michx. This semi-twining plant had been collected
twice on the banks of the Ohio River at Golconda, Pope County, by
S. A. Forbes and E. J. Palmer. Now Hardin County can be added to
its distribution.

Hardin Co.: limestone bluff, near Shetlerville, J. Ozment s.n.—
ROBERT H. MoHLENBROCK, JAMES E. OZMENT & GEORGE W. FOLKERTS,
SOUTHERN ILLINOIS UNIVERSITY, CARBONDALE.

NOSTOC PARMELIOIDES IN MASSACHUSETTS — Studies of
the algae of Worcester County are few, the most recent
being those of Auyang (1) and the author (6). Particular
interest is aroused, therefore, when a rather erratically
occurring alga (3) is located in the county.

In September, 1961 Dr. Burton Gates of Clark University

1962] Nostoc Parmelioides — Webber 359

collected growths of Nostoc from granite rocks in Keyes
Brook, East Princeton, Mass. While instructing in the sum-
mer session at Clark, I became interested in learning more
about this alga, for it appeared to be restricted to this habi-
tat and location. The water temperature at several collecting
trips was 22? to 23? C., and the pH a constant 8.3. Exposure
ranged from full sun to partial shade. Specimens were sent
to Mr. William Daily (2), and the alga was determined to
be Nostoc parmelioides Kuetz.

Tilden (5) records collections of this alga made between
1851 and 1898 from five states, Connecticut, Pennsylvania,
Wyoming, Nevada, New York, and from Canada.

Drouet (3) considers it to have “a peculiar distribution
in North America." He has records of it from the following
stations: Connecticut (Goshen, 1895; Mt. Carmel, 1891;
Hamden, 1932; Quinebaug River, 1893), Quebec in 1930,
New Jersey (Delaware River, 1941; Saxton Falls, 1953,
1958), Indiana in 1934, Nebraska in 1948, and then numer-
ous locations from the Rocky Mountains westward.

Fralick (4) at the Farlow Herbarium has seven collec-
tions of this species from Connecticut, Wisconsin, and fur-
ther west.

The collections by Gates in 1961, then, appear to be a first
for this species not only in Worcester County, but also in
Massachusetts.

Several trips to East Princeton were made by the author
in July, 1962, and specimens of Nostoc parmelioides are on
deposit at the following institutions: Academy of Natural
Sciences, Philadelphia, Pa., Butler University, Indianapolis,
Ind., Cornell University, Ithaca, N. Y., Dartmouth College,
Hanover, N. H., Farlow Herbarium, Harvard, Cambridge,
Mass., Michigan State University, E. Lansing, Mich., Uni-
versity of Massachusetts, Amherst, Mass. — EDGAR E. WEB-
BER, DEPARTMENT OF BOTANY, UNIVERSITY OF MASSACHU-
SETTS, AMHERST.

LITERATURE CITED
1 AUYANG, S. C. 1960. Survey and Taxonomy of the Algae of Lake
Quinsigamond. M.A. Thesis. Clark Univ.; Rhodora 64: 49-59.
1962.
2 DaiLvy, W. A. Dept. of Botany. Butler Univ. Indianapolis. Per-
sonal Communication.

360 Rhodora [Vol. 64

3 Drouet, F. Academy of Natural Sciences. Philadelphia. Personal

Communication.
4 FRALICK, R. A. Farlow Herbarium, Harvard Univ. Personal Com-
munication.

5 TILDEN, J. 1910. Minnesota Algae. Minneapolis.
6 WEBBER, E. E. 1961. The Ecology of Some Attached Algae in
Worcester County, Massachusetts. M.S. Thesis. Cornell Univ.

Volume 64, No. 760 including pages 291-379, was issued December 31, 1962

Errata for Rhodora Volume 64

p. 33. Caption to Fig. 1.
Transpose Rorippa nasturtium — aquaticum and R. microphylla

p. 40 line 38, for DOMINICA read DOMINICAN REPUBLIC

p. 172 Carex plantagianea read Carex plantaginea

p. 173 line 45, C. striata read C. stricta

p. 175 line 44, C. laevigata read C. laevivaginata

p. 177 line 9, C. Sprengellii read C. Sprengelii

p. 177 line 27, C. chordorhiza read C. chordorrhiza

p. 177 line 28, C. crinita var. brevirostris read C. crinita var. brevi-
crinis

1962]

Index to Volume 64

361

INDEX TO VOLUME 64

New scientific names and combinations are printed in bold face type

Acer pensylvanicum 281; sac-
charum 333; spicatum 281
Achillea borealis 346; lanulosa
346

Achyranthes corymbosa 179

Aciphyllaea 4, 11

Adams, Preston, Studies in the
Guttiferae, II Taxonomic and
distributional observations on
North American taxa 231

Additions to the Bryophyte flora
of Keweenaw County, Michigan
121

Additions to the flora of Grand
Manan Island, New Brunswick,
Some 98

Adenopappus 2

Adenophyllum 9, 11

Adiantum Poiretii 348

Aesculus glabra 331

Afzelia cassioides 90

Agalinis linifolia 91; obtusifolia
91; purpurea 91; setacea 88;
tenella 91

Agaricus trullisatus 130; uranius
131

Ageratum corymbosum 260, 262;
houstonianum 262; latifolium
262; paleaceum var. nelsonii
262; cf. paleaceum var. palea-
ceum 262; cf. tomentosum 262

Agrimonia microcarpa 356, 357;
parviflora 357; platycarpa 357

Agropyron Hybrids and the Sta-
tus of Agropyron pseudorepens
143

Agropyron pseudorepens 143, 144,
145, 146; repens 143, 144, 145;
repens var. subulatum 344; re-
pens var. subulatum forma
setiferum 345; repens var. sub-
ulatum forma Vaillantianum
345; smithii 143, 145, 354; tra-

chycaulum 143, 144, 145; tra-
chycaulum var. majus 144, f.
pseudorepens 145

Agrostis alba 345; alba var. pa-
lustris, 345; tenuis 118; varia-
bilis 117, 118; rossae 117, 118

Aguilegia canadensis var. austra-
lis 150, 154

Aldama dentata 255, 256, 266

Algae, of Lake Quinsigamond, A
survey of the 49

Alisma plantago-aquatica 227,
228; subcordatum 227, 228,
229

Amanita muscaria 127; parcivol-
vata 126, 127

Amanitopsis parcivolvata 126

Amblystegium serpens 124

Ambrosia cumanensis 265; peru-
viana 265

Amelanchier arborea 18

Armoracia aquatica 324;
cana 324

Amphicarpa bracteata 91

Amphora ovalis 53

Anabaena limnetica 53; variabi-
lis 53; viguieri 53

Anabaenopsis elenkinii 53

Andreaea rupestris 122, 123, 124

Ankistrodesmus convolutus 53;
faleatus 53; var. acicularis 53

Annual species of Houstonia in
Illinois, The 28

Anomalostylus coriifolius — 304,
306, 307; crateriformis 304,
305, 307; grandis (comb. nov.)
304, 306

Anomodon attenuatus 124

Aphanocapsa grevillei 53

Aphanochaete repens 53; clathra-
ta 53; microscopica 53; stag-
nina 54

Apios americana 151, 156; tuber-

rusti-

362

osa 151

Apiosporina morbosa 134, 135

Aquatic form of Alisna subcorda-
tum Raf., The 227

Arctotis stoecladifolia 261, 270

Arcyphyllum simplicifolium 62

Arenaria lateriflora 345

Artemisia dracunculoides 354

Ascyrum hypericoides 237; multi-
caule 236

Asplenium trichomanes 209

Aster bimater 252, 263; exilis var.
australis 253, 263; foliaceus
346; Herveyi 113, 114; X Her-
veyi, 113, 114, 155 Pl. 1267;
johannensis 346; lima 252; ma-
crophyllus 113, 114, 116; shortii
333; spectabilis 113, 114, 116;
subulatus 253; umbellatus 346

Asterionella formosa 54; gracil-
lima 54

Atriplex glabriuscula 345; patula
var. patula 345

Atroxina liberica 104

Aureolaria flava 91; pectinata 91;
pedicularia 333

Auyang, Tang Shih-Chen, A sur-
vey of the Algae of Lake Quin-
sigamond 49

Baccharis glutinosa 263; serrae-
folia 263

Bachman, John and the Happoldt
European Journal 92

Bacopa monnieri 153, 158

Banks, Donald J., Paspalum fim-
briatum in the United States
85

Barr, Margaret E. and Howard
E. Bigelow, Contribution to the
Fungus Flora of Northeastern
North America. II 126

Batrachospermum vagum 54

Batischia caroliniensis 89

Beaman, J. H. and B. L. Turner,
Chromosome Numbers in Mexi-
can and Guatemalan Composi-
tae 271

Rhodora

[Vol. 64

Bean, R. C., A. F. Hill and R. J.
Eaton, Fifteenth Report of the
Committee on Plant Distribu-
tion 168

Berlandiera lyrata 265

Betula papyrifera 18; var. cordi-
folia 100

Bidens connata var. ambiversa
23; var. anomala 23; var. fallax
23; var. gracilipes 25, 26, 27,
28; var. petiolata 23; var. pin-
nata 23, 25, 26, 28; var. typica
(connata) 23; triplinervia var.
macrantha 272, 273, 275

Bigelow, Howard E. and Margar-
et E. Barr, Contributions to the
Fungus Flora of Northeastern
North America. II 126

Boeberastrum 11

Bonamia humistrata 153, 156;
pickeringii 153, 156

Bredemeyera colletioides 104

Brickellia robinsonii 262

Britton, Donald M., Dryopteris
dilatata (Hoffm.) A. Gray in
North America 207

Brunnichia cirrhosa 150, 154

Bryophyte Flora of Keweenaw
County, Michigan, Additions to
the 121

Bryum affine 124; cuspidatum 124

Buchloé dactyloides 355

Buchnera americana 153

Buckleya distichophylla 332

Bulbochaete repanda 54; scrobi-
culata 54

Bumilleria sicula 54

Burmannia capitata 88

Burns, Richard L. and Edward L.
Davis, The Use of Vapor Frac-
tometry in the Analysis of
Some New England Hops 243

Cacalia peltata 276; sinuata 269;
tussilaginoides 269

Calamagrostis canadensis
var. robusta 345

Calea integrifolia 257, 268; nel-

345;

1962] Index to Volume 64 363

sonii 268; trichotoma 272, 258,
268

Calliergon giganteum 124

Caloneis silicula var. inflata 54

Calothrix stellaris 54

Cantharellus cibarius 127; cin-
nabarinus 129; Friesii 129;
minor 127, 128 Pl. 1268, 129;
morgani 129; olidus 129; rosel-
lus 129

Capsella Bursa-pastoris 345

Cardamine bulbosa 150; pennsyl-
vanica 231

Cardenanthus boliviensis 299, 300;
longitubus 299, 300; orurensis
299; peruvianus 299, 301; Shep-
ardae 299, 301; tunariensis 299,
302; Vargasii 299, 303; Ventu-
rii 299, 302

Carex abdita 168, 174, 357;
abscondita 168, 176; acutifomis
168, 179; adusta 169, 177; aenea
169, 174; aestivalis 169, 176;
alata 169, 176; albolutescens
169, 176; albursina 169, 176; al-
opecoidea 169, 176; amphibola
var. rigida 169, 176; var. tur-
gida 169, 176; angustior 169,
173; annectens 169, 175; var.
xanthocarpa 169, 175; aquatilis
var. altior 169, 177; arcta 169,
174; arctata 169, 174; argyran-
tha 169, 175; artitecta 169, 175;
var. subtilirostris 169, 177; ath-
erodes 169, 177; atlantica 169,
175; atratiformis 169, 174;
aurea 169, 177; Backii 169, 174;
Baileyi 169, 176; Barrattii 169,
177; Bebbii 169, 177; Bicknellii
169, 176; Bigelowii 169, 175;
blanda 169, 175; brevior 169,
175; bromoides 169, 174; brun-
nea 169, 179; brunnescens 169,
175, 343, 345; var. sphaeros-
tachya 169, 174; bullata 169,
176; Bushii 169, 176; Buxbau-
mii 169, 174; canescens 169,
174; var. canescens 345; var.

disjuncta 169, 173; var. subloli-
acea 169, 173; capillaris 169,
175; var. major 169, 176; capi-
tata 169, 175; caryophyllea 169,
179; castanea 169, 176; cepha-
lantha 169, 173; cephaloidea
169, 177; cephalophora 169, 175;
chordorrhiza 169, 177; Collinsii
169, 177; communis 169, 174;
comosa 170, 175; conoidea 170,
174; convoluta 170, 174; X co-
pulata 170, 177; Crawei 170,
176; Crawfordii 170, 175; crini-
ta 170, 173; var. brevicrinis
170, 177; var. gynandra 170,
173; var. minor 170, 174; var.
Porteri 170, 177, 178; var. si-
mulans 170, 174; cristatella 170,
176; cumulata 170, 175; Davisii
170, 176; debilis 170, 177, 178;
var. interjecta 170, 177, 178;
var. Rudgei 170, 173; var.
strictior 170, 177, 178; deflexa
170, 175; demissa 170, 174; De-
weyana 170, 174; diandra 170,
177; digitalis 170, 175; disper-
ma 170, 174; eburnea 170, 176;
elachycarpa 170, 176; eleocharis
355; Emmonsii 170, 175, 357;
exilis 170, 177, 178; festucacea
170, 176; filifolia 118, 119; flac-
cosperma var. glaucodea 170,
176; flava 170, 174; var. fertilis
170, 174; var. gaspensis 170,
176; foenea 170, 175; folliculata
170, 175; formosa 170, 176;
Garberi var. bifaria 170, 176;
gracilescens 170, 176; gracil-
lima 170, 174; granularis 170,
176; var. Haleana 170, 177;
Grayii 170, 176; var. hispidula
170, 177, 178; gynocrates 170,
174; X Hartii 170, 177, 178;
Haydenii 170, 174; hirsutella
170, 176; hirta 170, 179; hirti-
folia 170, 177; Hitcheockiana
170, 176; hormathodes 170, 177,
345; hormathodes X straminea

364

170; Houghtonii 171, 174;
Howei 171, 175; hystricina 171,
174; incomperta 171, 175; in-
terior 171, 174; intumescens
171, 173; var. Fernaldii 171,
175; Josselynii 171, 176; katah-
dinensis 171, 175; X Kniesker-
nii 171, 177, 178; lacustris 171,
174; laevivaginata 171, 175; la-
nuginosa 171, 177; lasiocarpa
var. americana 171, 173; laxi-
culmis 171, 175; laxiflora 171,
175; lenticularis 171, 174; var.
albi-montana 171, 175; var.
Blakei 171, 174; leporina 171,
179; leptalea 171, 173; lepto-
nervia 171, 174; limosa 171,
173, 174; livida var. Grayana
171, 177, 178; Longii 171, 176;
lupuliformis 171, 176; lupulina
171, 175; var. pedunculata 171,
175; lurida 171, 175; Macken-
ziei 171, 177; mainensis 171,
175; Merritt-Fernaldii 171, 175;
mesochorea 171, 177, 178; Mich-
auxiana 171, 174; Mitchelliana
171, 176; molesta 171, 177, 178;
Muhlenbergii 171, 175; var. en-
ervis 171, 175; nigra 171, 177;
var. strictiformis 171, 177; nig-
romarginata 171, 176, 357;
normalis 171, 175; novae-
angliae 171, 175; oligocarpa
171, 176; oligosperma 171, 175;
X Olneyi 171, 177, 178; ormo-
stachya 171, 175; oronensis 171,
177, 178; paleacea 171, 177, 545;
pallescens var. neogaea 171,
173; panicea 171, 179; pauci-
flora 171, 175; paupercula 171,
174; var. irrigua 171, 174; var.
pallens 172, 175; Peckii 172,
174; pedunculata 172, 174; pen-
sylvanica 18, 172, 175; var. dis-
tans 172, 175; physorhyncha
357; plantaginea 172, 177;
platyphylla 172, 175; polymor-
pha 172, 176; praegracilis 355;

Rhodora

[Vol. 64

prairea 172, 177; prasina 172,
175; praticola 172, 177, 178;
projecta 172, 174; Pseudo-
Cyperus 172, 175; X pseudo-
helvola 172, 177, 178; radiata
172, 175; rariflora 172, 175;
retroflexa 172, 176; retrorsa
172, 175; Richardsonii 172,
177, 178; rosea 172, 177; rostra-
ta 172, 173; var. ambigens 172,
177, 178; var. utriculata 172,
173; rupestris 118, 119; salina
var. kattegatensis 172, 177;
salina X stricta 172, 177, 178;
saxatilis var. miliaris 172, 175;
var. rhomalea 172, 175; scab-
rata 172, 174; Schweinitzii 172,
176; scirpoidea 172, 174; sco-
paria 172, 175, 345; var. tessel-
ata 172, 177, 178; seorsa 172,
176; X setacea 172, 177, 178;
silicea 172, 177, 343, 345; spar-
ganioides 172, 177; spicata 172,
179; Sprengelii 172, 177; squar-
rosa 172, 176; stenophylla var.
enervis 355; sterilis 172, 177,
178; stipata 172, 173; straminea
172, 176; striatula 172, 176;
stricta 172, 173; var. strictior
172, 174; styloflexa 172; Swanii
172, 115; tenera 172, 173, 174;
var. echinodes 172, 177, 178;
tenuiflora 172, 174; tetanica
172, 176; tincta 172, 175; tonsa
172, 175; torta 173, 174; tribu-
loides 173, 174; X trichina 173,
177, 178; trichocarpa 173, 176;
trisperma 173; var. Billingsii
173; Tuckermanni 173, 177;
typhina 173, 176; umbellata
173, 175, 357; vaginata 173,
176; vesicaria 173, 174; var.
distenta 173, 177, 178; var.
jejuna 173, 174; var. monile
173, 174; var. Raeana 173,
174; vestita 173, 175; virescens
173, 175; viridula 173, 177, 178;
vulpinoidea 173; Walteriana

1962]

var. brevis 173, 176; Wiegandii
173, 174; Willdenowii 173, 177,
178; Woodii 173, 177, 178;

Carpalobia lutea 104

Carphephorus bellidifolius 92; to-
mentosus 92

Carum Carvi 346

Casper, S. Jost, On Pinguicula
macroceras Link in North
America 212

Celarier, R. P., Shamim A. Fa-
ruqi and K. In Mehra, A New
Species of Bel irenaea from
Mexico 329

Celarier, R. P., K. L. Mehra and
Shamim Faruqi, The Taxonomy
of Setcreasea ovata 68

Celtis tenuifolia 333

Centrosema virginianum 156, 157

Cephaloziella striatula 124; sub-
dentata 124

Cerastium viscosum 343, 345; vul-
gatum 343, 345

Ceratiola ericoides 234

Ceratium hirundinella 54

Cercis canadensis 333

Characiopsis acuta 54

Chenopodium album 136

Chlamydomonas angulosa 54;
mucicola 54; polypyrenoideum
54; snowii 54; sphagnicola 54

Chlorella ellipsoidea 54; vulgaris
54

Chloris petraea 86;
355

Chlorococcum humicola 54

Chromosomal Races in Eastern
North American Species of
Hedyotis (Houstonia) 313

Chromosome Counts of Two Thy-
melaeaceae 277

Chromosome Numbers of Dysso-
dia (Compositae-Tagetinae)
and Phyletic Interpretations 2

Chromosome Numbers for Some
Angiosperms of the Southern
United States and Mexico 147

Chromosome Numbers in the

verticillata

Index to Volume 64

365

Compositae. VI Additional
Mexican and Guatemalan Spe-
cies 251

Chromosome Numbers in Mexican
and Guatemalan Compositae
271

Chromosome Numbers in Striped
and Mountain Maples 281

Chroococcus limneticus 54

Chrysactinia 2, 9

Chrysanthemum parthenium 269

Chrysocoma odoratissima 90; pa-
niculata 90

Chrysopsis villosa 263

Cintractia caricis var. caricis 119

Cipura Goodspeediana comb. nov.
311

Circaea alpina 63, 64, 65, 66, 67;
canadensis 63, 64, 65, 66, 67;
intermedia 67; lutetiana 63, 67;
quadrisulcata 63, 64, 65, 66, 67;
Pl. 1267

Cirsium arvense 346; forma albi-
florum 346; horridulum 156,
158; skutchii 276

Claytonia virginica 150, 154

Cliftonia monosepala 151, 156

Clinopodium rugosum 184

Clitocybe morgani 126, 129; olida
129; trullisata 130

Clomenocoma 4, 9, 10

Closterium acerosum 54; calos-
porum 54; dianae 54; didymoto-
cum 54; ehrenbergii 54; incur-
vum 54; lagoense 54; lanceolat-
um 54; liebleinii 54; lunula 54;
moniliferum 54; turgidum 54;
sigmoideum 54

Cnidoscolus texanus 151, 156

Cocculus carolinus 150

Cody, W. J., A New Station for
Pinguicula vulgaris in Ontario
182

Coelastrum microporum 54

Coelosphaerium dubium 54; nae-
gelianum 54

Coleochaete orbicularis 54; scuta-
ta 54; soluta 54

366

Coelopleurum lucidum 343, 346

Commelina diffusa 149, 154; erec-
ta var. angustifolia 149, 154;
virginica 149

Contribution to the Fungus Flora
of Northeastern North Ameri-
ca. II 126

Conyza canadensis 263; coronopi-
folia 263; sophiaefolia 263

Cooperrider, Tom S., The Occur-
rence and Hybrid Nature of an
Enchanters Nightshade in
Ohio 63

Coreopsis mutica 257, 268

Cornus florida 136

Cosmarium biretum var. trigib-
berum 54; botrytis 54; broomei
54; dentiferum 54; formosulum
54; var. nathorstii 54; grana-
tum 54; margaritatum 54; mo-
nomazum 54; nitidulum 54;
orbiculatum 54; panamense 54;
protractum 54; pseudo conna-
tum 54; punctulatum 54; quad-
um 54; reinforme 54

Cosmocladium hitchcockii 54

Cosmos diversifolius 268

Crookea microsepalum 152

Crotalaria angulata 89; maritima
89; rotundifolia 89; sagittalis
91

Croton capitatus var. lindheimeri
151, 156

Crucifer from the Great Slave
Lake Area of Canada, A New
324

Crucigenia truncata 54

Cryptozramma crispa 210

Cryptotaenia canadensis 230

Cyclotella bodanica 54; var. stel-
lata 54; comta 54

Cydonia oblonga 357

Cymbella aspera 54; tumida 54

Cynoctonum mitreola 91, 152;
sessilifolium 89

Cynodontium schistii 124

Cypella coriifolia 306; cyrtophyl-
la 295; Goodspeediana 311;

Rhodora

[Vol. 64

grandis 304; lutea 308, 309

Cyperus dentatus 349; ferrugines-
cens 352, 353

Cyperus ferruginescens in Ver-
mont 352

Cyperus odoratus 353; X Cyper-
us Weatherbianus 349, 350

X Cyperus Weatherbianus, A
note on 349

Cyrilla racemiflora 151

Cystopteris montana 210

Cytological Observations of Poly-
gala in Eastern North America
102

Dahl, Eilif, Three Misidentified
So-called Cordilleran Species
in Eastern North America 117

Dahlia dissecta 268; scapigera
268

Daphne blagayana 278, 280; col-
lina 278; odora 277, 279; pet-
raea 278; sericea 278

Daphnopsis americana 278; ssp.
caribaea 279, 280

Dasistoma macrophylla 331, 388,
334 Pl. 1272

Dasistoma macrophylla, The Par-
asitic Behavior of, 331

Daubs, Edwin H., The Occurrence
of Spirodela oligorrhiza in the
United States 83

Davis, Edward L. and Richard L.
Burns, The Use of Vapor Frac-
tometry in the Analysis of
Some New England Hops 243

Davis, Shirley A. and Walter H.
Lewis, Cytological Observations
of Polygala in Eastern North
America 102

Decodon verticillatus 91

Dennstaedtia distenta 348

Desmidium swartzii 54

Desmodium canescens 151, 156;
lineatum 62; rotundifolium 62

Diachora onobrychidis 134

Diatoma hiemale, var. mesodon
54; vulgare 54

1962]

Dichodontium pellucidum 124
Dichondra carolinensis 91
Dicliptera brachiata 282, 283
Dictyosphaerium pulchellum 54
Diervilla lonicera 18
Dinobryon sertularia 54
Dioscorea quaternata 89, 91
Ditrichum cylindricum 123
Dolicholus simplicifolius 62
Draparnaldia glomerata 54; ju-
dayi 54
Dryopteris assimilis 208, 209, 210,
211; austriaca, var. austriaca
207, 209; campyloptera 210,
211; dilatata 207, 208, 209, 210,
211; intermedia 208, 209, 210;
maderensis 208, 209; marginal-
is 210; spinulosa 207; var.
americana 207, 210; parallelo-
gramma 348
Dryopteris dilatata (Hoffm.) A.
Gray in North America 207
Dyssodia acerosa 4, 7, 9, 11, 14;
anomala 13; anthemidifolia 11;
aurea 7; var. aurea 13; var.
polychaeta 13; aurantia 10;
aurantiaca 12; Belenidium 5, 9,
12; cancellata 4, 11; coccinea
11; concinna 11; Cooperi 10;
decipiens 7, 10, 13; diffusa 13;
Gentryi 11; grandiflora 10;
Hartwegii 12; integrifolia 11;
Jelskii 10; littoralis 11; micro-
poides 6, 7, 9, 13; montana 10;
mutica 13; Neaei 13; neomexi-
cana 6, 7, 9, 13; oaxacana 11;
Palmeri 11; papposa 3, 4, 10;
pentachaeta 5, 6, 12; pinnata 4,
10; porophylla 4, 11; porophyl-
loides 10; remota 10; Seleri 11;
serratifolia 11; setifolia 6; var.
Gregii 12; var. setifolia 11, 12;
speciosa 10; squarrosa 10; sub-
integerrima 4; tagetiflora 7, 10;
tagetoides 5, 11; tenuifolia 7,
13; tenuiloba 6, 9, 12; tephro-
leuca 11; texana 13; Thurberi
5, 12; Treculii 7, 13; Wrightii

Index to Volume 64

367

12
Dysodiopsis 5, 8, 11
Dugesia mexicana 265
Dulichium arundinaceum 349

Eaton, R. J., R. C. Bean and A.
F. Hill, Fifteenth Report of the
Committee on Plant Distribu-
tion 168

Eaton, Richard J., Cyperus Fer-
ruginescens in Vermont 352

Ebinger, Jchn E., The Varieties of
Luzula Acuminata 74

Eleocharis elliptica 100; obtusa
148, 154; tenuis, var. tenuis 100

Elephantopus carolinianus 156

Eleutherine bulbosa 310

Elymus arenarius 284, 285; var.
villosus 345; mollis 284

Elytraria caroliniensis 89; virgata
89

Empetrum nigrum 341

Encalypta procera 123

Enchanter’s Nightshade, The Oc-
currence and Hybrid Nature of,
in Ohio 63

Epithemia turgida 54

Erigeron coronarius 272, 274;
delphinifolius 263; heteromor-
phus 263; karvinskianus 263;
repens 263; scaposus 253, 263,
264, 274

Erythrina herbacea 151

Erythronium carolinianum. 90

Euastrum elegans 54; pulchellum
54; verrucosum 54

Euglena gracilis 54; spirogyra 55

Eupatorium aschenbornianum
262; betonicum 262; gracili-
caule 262; morifolium 260, 262;
petiolare 260, 262; pycnocephal-
um 252, 260, 262, 263; quad-
rangulare 263

Euphorbia dictyosperma 357; ob-
tusata 357

Euphrasia americana 346; cana-
densis 343, 346; Randii 346

Evers, Robert A., Illinois Flora:

368

Notes on Limnosciadium, Dic-
liptera and Iresine 282

Ewan, Joseph, John Bachman and
the Happoldt European Journal
92; Makers of North American
Botany (Review) 186

Faruqi, Shamim A., K. L. Mehra
and R. P. Celarier, A New Spe-
cies of Setcreasea From Mexi-
co 829; The Taxonomy of Set-
creasea ovata 68

Fell, Egbert W., Western Plains
Plants in Northern Illinois 354

Ferns and Fern Allies of Chihua-
hua 347

Festuca rubra, var. rubra 344

Fifteenth Report of the Commit-
tee on Plant Distribution 168

Fisher, T. Richard and James R.
Wells, Heteromorphic Pollen
Grains in Polymnia 336

Flora of Illinois, Additions to the
356

Flora of the Machias Seal Islands,
Changes in 340

Florestina pedata 275

Folkerts, George W., Robert H.
Mohlenbrock and James E. Oz-
ment, Additions to the Flora of
Illinois 356

Foster, Robert C., Studies in the
Iridaceae, VII 291

Froelichia gracilis 354

Fragilaria capucina 55; constru-
ens 55; crotonensis 55; var.
prolongata 55

Fraxinus quadrangulata 333

Further Notes on the Distribution
of Bidens connata vars. pin-
nata and gracilipes 23

Galactia macreei 151, 156

Galeana pratensis 259, 261

Galinsoga ciliata 258; parviflora
258, 268

Galium tinctorium 346

Gaultheria procumbens 18

Rhodora

[Vol. 64

Gaylussacia frondosa 91

Genus Anonymous and its nom-
enclatural Survivors, The 87

Gelsemium sempervirens 91

Gerardia setacea 88

Glandularity in Rubus Allegheni-
ensis Porter 161

Globifera umbrosa 89

Gloeocystis ampla 55; gigas 55;
major 55; vesiculosa 55

Glycine monophylla 62; renifor-
mis 61, 62; simplicifolia 62; to-
mentosa 62

Gnaphaliopsis 6, 13

Gnaphalium leptophyllum
uliginosum 346

Golenkinia paucispina 55

Gomphoneis herculeana, var. ro-
busta 55

Gomphonema acuminatum 55;
constrictum 55

Gonatozygon kinahani 55

Grand Manan Island, Some Addi-
tions to the Flora of 98

Gratiola aurea 153, 158; brevifolia
154, 158; ramosa 153, 158; vir-
giniana 154, 158

Green, Peter S., Watercress in the
New World 32

Grimmia alpicola 124; apocarpa,
var. stricta 124

Grindelia inuloides 274

Grossularia echinella 150

Gymnodinium fuscum 55; palus-
tre 55

Gymnolaena 5, 11

264;

Halbig, Leo E. and Mohlenbrock,
Robert H., The Annual Species
of Houstonia in Illinois 28

Halesia carolina 152; diptera, var.
diptera 152, 156; var. magni-
flora 152, 156; tetraptera 152,
156

Haplopappus stoloniferus
texensis (sp. nov.) 142, 143

Happoldt European Journal, John
Bachman and the (Review) 92

264;

1962]

Hedyotis caerulea 313, 315, 317,
320, 321, 322, 323; var. caerulea
315, 319; var. faxonorum 317,
319; canadensis 313, 315, 317,
319, 321; longifolia 313, 314,
317, 318, 319, 321, 322; var.
compacta 314; var. glabra 315,
319; var. longifolia 314; mich-
auxii 318, 317, 319, 321, 322;
nuttaliana 318, 315, 317, 319;
polypremoides 313; purpurea
313, 314, 317, 318, 319, 320, 321,
322, 323; var. calycosa 314,
318; var. montana 314, 318;
var. purpurea 314

Hedyotis (Houstonia), Chromoso-
mal Races in Eastern North
American Species of 313

Helenium hoopesii 275

Helianthella quinquenervis 272,
275

Helianthus laciniatus 266

Heliopsis buphthalmoides 254, 265

Heracleum maximum 346

Heritiera tinctorium 89

Hermann, F. J., Additions to the
Bryophyte Flora of Keweenaw
County, Michigan 121

Heterocladium squarrulosum 123

Heteromorphie Pollen Grains in
Polymnia 336

Heterotheca inuloides, var. inu-
loides 264, 274

Hieracium mexicanum 276; seleri-
anum 276

Hill, A. F., R. J. Eaton and R. C.
Bean, Fifteenth Report of the
Committee on Plant Distribu-
tion 168

Holosteum umbellatum 222

Holosteum Umbellatum L., an
Angiosperm New to Michigan
222

Hodgdon, A. R. and Radcliffe B.
Pike, Changes in Flora of the
Machias Seal Islands 340; Some
Additions to the Flora of Grand
Manan Island, New Brunswick

Index to Volume 64

369

98

Hodgdon, A. R. and F. L. Steele,
Glandularity in Rubus Alle-
gheniensis Porter 161, Juniper-
us Horizontalis in New Hamp-
shire 351

Hops, The Use of Vapor Frac-
tometry in the Analysis of
Some New England 243

Hormidiopsis ellipsoideum 55

Houstonia caerulea 29, 30, 91;
var. faxonorum 28; faxonorum
28; minima 29, 30, 31; procum-
bens 89; pusilla 29, 30, 31; ro-
tundifolia 89; serpyllifolia 29,
313; tenuifolia 313

Humulus lupulus 243

Hyalotheca dissiliens 55

Hydnum erinaceus 91

Hydrodictyon reticulatum 55

Hygrophoropsis olida 129

Hymenatherum 6, 8, 9, 12

Hymenophyllum tunbridgense 348

Hypochoeris radicata 346

Hypericum aphyllum 235; boreale
240; brevistylum 238; cana-
dense, 240; var. galiiforme 241;
var. magninsulare 241; cumuli-
cola 234, 238; denticulatum, 238,
241, 242; var. recognitum 241,
242; var. acutifolium 241, 242;
var. denticulatum 241; dissimu-
latum 240; drummondii 284,
238; eastwoodianum 238; fasti-
giatum 238; formosum 238, 239,
240; fuertesii 234; gentianoides
233, 234, 235, 238; gnidioides
238; mraveolens 238. 239; gym-
nanthum 240; hintonii 238;
hondurasense 238; hypericoides
237; longibracteatum 238; ma-
jus 240; microsepalum 152, 156;
mitchellianum 238, 239, 240;
mutilum 240; var. latisepalum
240; paniculatum 238; pauci-
florum 234, 238; paucifolium
238; perforatum 235; pinetor-
um 238; pringlei 235; pratense

370

238; pseudomaculatum 238,
239, 240; punctatum 238, 239,
240; schaffneri 238; scouleri
238, 239, 240; setosum 238;
silenoides 238; stragulum 236,
237; styphelioides 236; sub-
montanum 238; terrae-firmae
235, 236; uliginosum 238;

woodsonii 238

Hyptis alata 185; var. stenophylla
186; floridana 185; latidens 184,
185, 186; leiocephala 185, 186;
mutabilis 185; radiata 184, 185;
rugosa 184

Hyptis Radiata (Labiatae) an
Illegitimate Name 184

Illinois Flora: Notes on Limnosci-
adium, Dicliptera and Iresine
282

Iostephane heterophylla 260; var.
dicksonii 266; trilobata 266

Ipheion uniflorum 149

Iresiine rhizomatosa 282, 283

Iridaceae, Studies in the, VII 291

Iris Hookeri 343, 345; versicolor
341, 345

Isopyrum biternatum 150, 154

Jackson, Ray C., A New Species
of Haplopappus, Section Ble-
pharodon 142

Jacquemontia abutiloides 153

Jaegeria hirta 266, 272; peduncu-
lata 266, 272; petiolaris 272,
273, 275

Johnston, Marshall C., The Nose-
burn (Tragia, Euphorbiaceae)
of Western Texas 137

Johnston, Marshall C. and B. L.
Turner, Chromosome Numbers
of Dyssodia (Compositae-Tage-
tinae) and Phyletic Interpreta-
tions 2

Jones, Neville G., A note on the
Ascription of Elymus Arenari-
us to Illinois 284

Juncus articulatus 100; bufonius

Rhodora

[Vol. 64

345; var. halophilus 345

Juncoides carolinae 81; pilosum
var, michiganense 80; saltuense
80; var. saltuense 80

Juniperus horizontalis 341; vir-
giniana 333

Juniperus Horizontalis in New
Hampshire 351

Justicia lanceolata 156, 158

Kalmia angustifolia 98

Keweenaw County, Michigan,
Additions to the Bryophyte

Flora of 121

King, R. M., B. L. Turner and M.
Powell, Chromosome Numbers
in the Compositae. VI Addi-
tional Mexican and Guatemalan
Species 251

Kirchneriella lunaris, var. irregu-
laris 55

Kobuski, Clarence E., Ernest
Jesse Palmer, 1875-1962 195

Krameria lanceolata 151

Kuhnia pinnata 90

Laccaria trullisata 130 Pl. 1269,
131

Lachnanthes tinctoria 89

Lathryus odoratus 338; palustris
343

Lactuca pulchella 270

Lakela, Olga, Occurrence of Spe-

cies of Polycarpaea Lam.
(Caryophyllaceae) in North
America 179

Labetina 4, 9, 10
Leersia oryzoides 284
Lemna minor 84; oligorrhiza 83
Lentinus haematopus 131
Leontodon autumnalis 346
Leptosira mediciana 55
Leptosphaeria eutypoides 136
Lespedeza cytisoides 60
Lesquerella purpurea 150, 154
Lewis, Walter H., Two New Spe-
cies of Polygala from Northern
Mexico 225

1962]

Lewis, Walter H. and Edward E.
Terrell, Chromosomal Races in
Eastern North American Spe-
cies of Hedyotis (Houstonia)
313

Lewis, Walter H., H. Larry Strip-
ling and Richard G. Ross,
Chromosome Numbers for Some
Angiosperms of the Southern
United States and Mexico 147

Lewis, Walter H. and Shirley A.
Davis, Cytological Observations
of Polygala in Eastern North
America 102

Liatris acidota 158; graminifolia
89

Licmophora gracilis 55; paradoxa
55

Limnosciadium pinnatum 282

Lindernia dubia 154, 158

Liquidambar Styraciflua 230

Ligusticum scothicum 346

Lithospermum caroliniense 89;
gmelinii 89

Long, Robert W. and Leonard J.
Uttal, Some Observations on
Flowering in Ruellia (Acantha-
ceae) 200

Lophozia quadriloba 122; ventri-
cosa 124

Luzula acuminata 74, 75; var.
acuminata 76, 77, 78, 79, 80, 81,
82; var. carolinae 76, 77, 78, 79,
81, 83; carolinae 74, 75, 81; var.
saltuensis 80; pilosa 74, 75; var.
B americana 80; plumosa 74;
saltuensis 74, 75, 80

Lycopodium tristachyum 98

Lycopus uniflorus 346

Lyngbya epiphytica 55; latissima
55; majuscula 55; wollei 55

Lyonia ligustrina 91

Machaeranthera gymnocephala
260, 264; parviflora 254; tenace-
tifolia 264

Machias Seal Islands, Changes in
Flora of the 340

Index to Volume 64

371

Mackeever, Frank C., An Albino
Fruited Form of Vaccinium
macrocarpon 350

Makers of North
Botany 186

Marica bulbosa 308, 309; lutea
309; Martii 309

Mastigostyla brevicaulis 292, 293,
294, 295; Cabrerae 292, 293;
Cardenasii 293, 297, 298; cyrto-
phylla 292, 293, 295, 296, 298;
gracilis 293, 296, 297; Hoppii
292, 295, 296, 298; Joergensenii
293, 297; Johnstonii 292, 294;
peruviana 293, 297, 298; poto-
sina 292, 294

Matricaria matricarioides 346

Mattuschkia aquatica 88

Mazus japonicus 229, 230, 231;
reptans 229, 230; rugosus 230

Mazus reptans (Scrophularia-
ceae) in Maryland 229

Mecardonia acuminata 154

Mehra, K. L., R. P. Celarier, and
Shamim A. Faruqi, A New Spe-
cies of Setcreasea From Mexico
329; The Taxonomy of Setere-
asea ovata 68

Melanoleuca davisiae 132

Melanospora barbata 135; sphae-
rophila 135

Melampyrum arvense 15; barba-
tum 15; lineare 15, 16, 17, 18,
20, 21, 22, 335, Pl. 1265; nemo-
rosum 15; ssp. pectinatum 21;
pratense 15, 16, 17, 20, 21; sil-
vaticum 15

Melosira italica 55; juergensii 55

Menodora scabra 152, 156

Merismopedia punctata 55

Mexico, A New Species of Setcre-
asea from 329; Two New Spe-
cies of Polygala from North-
ern 225

Michigan, Holosteum umbellatum
L. An Angiosperm New to 222

Micranthemum orbiculatum 89;
umbrosum 89

American

372

Micrasterias americana 55; apicu-
lata 55; var. fimbriata f. spin-
osa 55; radiata 55; rotata 55;
truncata 55

Microspermum debile 272, 273,
276

Microspora willeana 55

Microthamnion kuetzingianum 55

Mikania gonzalezii 252, 263; scan-
dens 252

Mnium punctatum var. elatum 124

Mohlenbrock, Robert H., James
E. Ozment, and George W. Fol-
kerts, Additions to the Flora of
Illinois 356

Mohlenbrock, Robert H. and Leo
E. Halbig, The Annual Species
of Houstonia in Illinois 28

Mougeotia capucina 55; floridana
55; genuflexa 55; laetevirens
55; parvula 55; reinschii 55;
robusta 55; scalaris 55

Mougeotiopsis calospora 55

Mycena urania 131

Nasturtium microphyllum 32, 35,
41; officinale 32, 33, 38; var.
microphyllum; 32, 38; uniseri-
atum 32, 34, 41

Navicula anglica 55

Neomarica bulbosa 308; Martii
309

Netrium digitus 55

Neurolaena 259; lobata 261, 269

Nevling, Lorin I., Jr., Chromo-
some Counts of Two Thyme-
laeaceae 277

New Species of Scirpus in the
Northeastern United States, A
43

New Species of Haplopappus, Sec-
tion Blepharodon, A 142

New Station for Pinquicula vul-
garis in Ontario, A 182

Nicotiana glauca 153

Nostoc muscorum 55;
lioides 359

Nostoc parmelioides in Massachu-

parme-

Rhodora

[Vol. 64

setts 358
Nomenclatural Notes on Two
Southeastern Rhynchosias 60
Noseburn (Tragia, Euphorbi-
aceae) of Western Texas, The
137
Note on the Ascription of Elymus
arenarius to Illinois, 284
Notoptera tequilana 266
Nymphoides aquatica 89

Occurrence and Hybrid Nature of
an Enchanter's Nightshade in
Ohio, The 63

Occurrence of Species of Poly-
corpaea Lam. (Caryophylla-
ceae) in North America 179

Occurrence of Spirodela oligorr-
hiza in the United States, The
83

Oedogonium borisianum 55; capil-
lare 55; crenulatocostatum 55;
grande var. aequatoriale 55;
laeve 55; plusiosporum 55;
pringsheimii 55

Oócystis pusilla 55

Ophiodothella vaccinii 133, 134

Oreoweisia serrulata 123, 124

Orthotrichum obtusifolium 124;
sordidum 123, 125

Oscillatoria cortiana 55; curviceps
55; ornata 55; prolifica 55;
splendida 55; tenuis 55

Osmunda cinnamomea 344

Ostrya virginiana 333

Oxylobus glanduliferis 263

Ozment, James E., George W.
Folkerts and Robert H. Mohlen-
brock, Additions to the Flora
of Illinois 356

Palmer, Ernest Jesse, 1875-1962
195

Pandorina morum 55

Parasitic Behavior of Dasistoma
Macrophylla, The 331

Parasitic Behavior of Melampy-
rum Lineare and a Note on its

1962]

Seed Color, The 15

Paronychia pulvinata 234

Parthenium tomentosum 265

Paspalum boscianum 356; caespi-
tosum 86; ciliatifolium 86; fim-
briatum 85, 86; lentiferum 356;
pubescens 86

Paspalum fimbriatum in the
United States 85

Pattersonia caroliniensis 90

Pediastrum biradiatum 55; bory-
anum 56; var. longicorne 56;
duplex 56; var. clathratum 56;
var. rotundatum 56; var. rugu-
losum 56; ehrenbergii 56; tetras
56; var. tetraodon 56

Pedicularis canadensis 335

Pellaea ternifolia 209

Penium digitus 56; margaritace-
um 56

Perdue, Robert E., Jr., Two New
Varieties and a New Combina-
tion in Rudbeckia 328

Periconia sphaerophila 134, 135

Peridinium cinctum 56

Perityle microglossa 258, 269;
palmeri 158

Perymenium asperifolia 266;
chalarolepis 266

Petalostemon corymbosum 90;

pinnatum 90

Phacus longicauda 56

Phaeostoma juniperina 136; lage-
narium 136; sphaerophila 126,
133, 134, 135, 136; vitis 136

Phaseolus reniformis 62

Philactis nelsonii 254, 265

Phleum pratense 345

Phoradendron bolleanum 149, 151

Phormidium nareanum 56

Phragmites communis 358

Phyla nodiflora 91

Phytolacca americana 150

Piehl, Martin A., Holosteum um-
bellatum L., An Angiosperm
New to Michigan 222; The Par-
asitic Behavior of Dasistoma
macrophylla 331; The Parasitic

Index to Volume 64

373

Behavior of Melampyrum line-
are and a Note on its Seed
Color 15

Pike, Radcliffe B. and A. R. Hodg-
don, Changes in Flora of the
Machias Seal Islands 340; Some
Additions to the Flora of Grand
Manan Island, New Brunswick
98

Pinus clausa 232, 234; resinosa
18, 19, 98, 99

Pinguicula camtschatica 212, 213;
daurica 212, 213; macroceras
212, 218, 214, 215, 216, 218, 219;
var. macroceras 212, 213; var.
microceras 215; microceras 212,
213, 218, 219; vulgaris 182, 183.
212, 213, 214, 215, 216, 218, 219

Pinguicula Macroceras Link in
North America 212

Pinnularia nobilis 56

Piqueria laxiflora 252; pilosa 260,
263, 272, 274; trinervia 263

Piriqueta glabrescens 152,
tomentosa 152, 156

Pitcheria galactoides 60

Planera aquatica 90

Planktosphaeria gelatinosa 56

Plant Distribution, Fifteenth Re-
port of the Committee on 168

Plantago juncoides, var. decipiens
346; major 346

Pleospora calvescens 136; larici-
na, var. laricina 136; pulchra
136; straminis 136

Pleurococcus vulgaris 56

Pleurotaenium coronatum 56;
maximum 56; trabecula 56

Pluchea odorata 264; purpuras-
cens 265

Poa annua 344; arachnifera 355;
palustris 358

Podochaenium eminens 260, 266

Pohl, Richard W. Agropyron Hy-
brids and the Status of Agropy-
ron pseudorepens 143

Poiretia procumbens 89

156;

374

Pollen, Heteromorphic Grains in
Polymnia 336

Polycarpaea brasiliensis — 180;
corymbosa 179, 180; var. brasi-
liensis 180

Polyeystis aeruginosa 56; incerta
56

Polygala alba 105, 106, 111; var.
suspecta 225; alpestris 110; al-
pina 110; amara 103, 110; ssp.
amarella 110; ssp. brachyptera
110; boykinii 111; calcarea 110;
chamaebuxus 110; chapmanii
105, 106, 111; comosa 110; cru-
ciata 105, 106, 111; curtissii
106, 111; grandiflora 106, 110;
japonica 103, 110; lutea 111;
major 110; mariana 106, 111;
myrtifolia 103, 110; nana 106,
111; nicaensis, ssp. corsica 110;
nutallii 106, 112; ovatifolia 111;
polygama, var. obtusata 106,
112; var. polygama 112; prae-
tervisa 104, 106, 112; ramosa
106, 111; reducta 105, 111; ru-
geli 106, 112; sanguinea f.
albiflora 357; scoparia 105, 106,
111; scoparioides 106, 111, 227;
serpyllifolia 110; —shinnersii
225, 226; triflora 110; tweedyi
110; vayredae 110; vergrandis
226; verticillata 106; var. iso-
cycla 112; virgata, var. speciosa
110; vulgaris 110

Polygonella basiramia 234

Polygonum aviculare, var. littor-
ale 345; bistortoides 119, 120

Polymnia apus 260, 265; canaden-
sis 339; laevigata 336, 337 Pi.
1273, 338, 339; maculata 265;
oaxacana 265

Polymnia, Heteromorphie Pollen
Grains in 336

Polystichium scopulinum 211

Populus grandidentata 18, 19

Potentilla anserina 343, 346;
Egedei. var. groenlandica 343,
346; millegrana 358; norvegica

Rhodora

[Vol. 64

345; var. monspeliensis 358

Powell, M., R. M. King, and B.
L. Turner, Chromosome Num-
bers in the Compositae. VI Ad-
ditional Mexican and Guate-
malan Species 251

Primula farinosa 338

Protoderma viride 56

Prunus geniculata 234

Psilactis asteroides 253, 260, 264;
brevilingulata 253, 264; coul-
teri 253

Psoralea alnifolia 62

Pteridium aquilinum 18

Puccinellia paupercula, var. alas-
kana 344

Pucciniastrum myrtilli 99

Pycnanthemum ? alatum 184,
185; albescens 153

Pylaisia selwynii 125

Quercus muehlenbergii 333
Quercus rubra 18, 333

Radiofilum flavescens 56

Ranunculus acris 345; repens var.
villosus 345

Raphanus raphanistrum forma
sulphureus 101

Ratibida columnifera 355

Raymond, Marcel, A Note on X
Cyperus Weatherbianus 349

Rhinanthus Crista-galli 346

Rhizoclonium hieroglyphicum 56;
var. hosfordii 56; hookeri 56

Rhoades, Richard W., The Aqua-
tic Form of Alisma subcorda-
tum Raf. 227

Rhododendron canadense 98, 99

Rhynchosia galactoides 60; pitch-
eria 60; reniformis 61, 62; sim-
plicifolia 61, 62; tomentosa var.
monophylla 62

Rhynchospora capitellata 349;
corniculata 357; macrostachya
357

Rhytidium rugosum 125

Rhytisma vaccinii 133

1962]

Ribes echinellum 150, 154

Rollins, Reed C., A New Crucifer
from the Great Slave Lake
Area of Canada 324

Rorippa crystallina sp. nov. 325
Pl. 1271, 326; microphylla 32,
33, 34, 35, 37, 41, 42; nasturti-
um-aquaticum 32, 33, 34, 35, 36,
37, 38, 42, 324; var. siifolia 35;
X sterilis 32, 35, 37, 42

Ross, Richard G., Walter H. Lewis
and H, Larry Stripling, Chro-
mosome Numbers for Some An-
giosperus of the Southern
United States and Mexico 147

Rotala ramosior 152, 156

Rubus, An Unusual 353

Rubus allegheniensis 18, 161, 162,
163, 164, 165, 166; var. neosco-
ticus 167; amicalis 101, 102;
arcuans 353; biformispinus 353;
canadensis 101, 102; elegantu-
lus 101, 102, 354; frondisentis
167; glandicaulis 167; grandi-
dens 353; hispidus 353, 354;
ravus 162, 163, 167; sceleratus
167; setosus 167, 354

Rudbeckia laciniata var. bipinnata
328, 329; var. digitata 329; var.
laciniata 329; var. montana
329; nitida var. nitida 328; var.
texana 328

Rudbeckia, Two New Varieties
and a New Combination in 328

Ruellia (Acanthaceae), Some Ob-
servations on Flowering in 200

Ruellia caroliniensis 90, 200, 203,
204, 205, 206; ciliosa 206; hete-
romorpha 206; humilis 201, 204,
205, 206; hybrida 90; peduncu-
lata 204, 205, 206; purshiana
206; strepens 201, 202, 203, 204,
205, 206; var. cleistantha 202;
forma cleistantha 202

Rumex Acetosella 344, 345; cris-
pus 343, 345; mexicanus 343,
345; obtusifolius 341; orbicu-
latus 345; pallidus 343, 345

Index to Volume 64

375

Sabatia campestris 153, 156

Sabazia humilis 258

Salmea scandens 260, 266

Salvia reflexa 153, 156

Samolus parviflorus 152, 156

Sanidophyllum cumulicola 232,
233, 234

Santamour, Frank S., Jr., Chro-
mosome number in striped and
mountain maples 281

Sanvitalia 265; procumbens 265;
275

Saururus cernuus 88, 91

Scapania paludicola 124

Scenedesmus abundans 56

Scenedesmus abundans var. brevi-
cauda 56; acutiformis 56; ar-
cuatus var. platydisca 56; arm-
atus var. major 56; bijuga 56;
brasiliensis 56; denticulatus 56;
dimorphus 56; longus 56; lon-
gus var. minutus 56; obliquus
56; quadricauda 56; quadri-
cauda var. longispina 56

Schistocarpha bicolor 269

Schkuhria anthemoides 258, 261;
anthemoides var. wislizeni 268;
multiflora 258; pinnata 258, 269

Schrankia nuttallii 355

Schuyler, Alfred E., A New Spe-
cies of Scirpus in the North
Eastern United States 43

Scirpus ancistrochaetus 44, 45. 46,
47, 48, 49, Pl. 1266; atrovirens
43, 44, 46, 47, 48, 49; caespito-
sus 124; expansus 46; peckii 46;
polyphyllus 43, 46; sylvaticus
46

Sclerocarpus dentatus 256, 261,
267; divaricatus 261, 267; fru-
tescens 261, 267; phyllocephalus
261, 267; schiedianus 255, 256;
sessilifolius 261, 267; uniserialis
256

Scoparia dulcis 154, 158

Scutellaria integrifolia 153, 156

Securidaca longipedunculata 104

Sedum Rosea 345

376

Senecio cobanensis var. sublanci-
niatus 269; confusus 269; con-
zatii 269, 276; deformis 269;
hirsuticaulis 269; imparipinnat-
us 259, 261, 269; iodanthus 276;
picridis 269; salignus 269; to-
luccanus 269

Setcreasea, A new species from
Mexico 329

Setcreasea australis 72; brevifolia
68, 69, 70, 71, 72, 13, 330; buck-
leyi 68; hirsuta 330; jaumaven-
sis 69, 70; lanceolata 330; leian-
dra 72; ovata 68, 69, 70, 71, 72;
pallida 68, 69, 70, 71, 72, 329;
palmeri 329, 330; purpurea 69,
71, 72

Setcreasea ovata, The Taxonomy
of 68

Seymeria cassioides 90; tenuifolia
90

Sherff, Earl Edward, Further
notes on the Distribution of
Bidens connata vars. pinnata
and gracilipes 23

Shinners, Lloyd H., Hyptis Radi-
ata (Labiatae) an Illegitimate
Name 184; Mazus reptans
(Scropulariaceae) in Maryland
229

Siegesbeckia nudicaulis 272, 275;
orientalis 272; repens 272, 275;
triangularis 272, 275

Silphium asperrimum 158; com-
positum 158

Sisyrinchium bromelioides 311;
deflexum 312; macrocephalum
311; montanum 2345; Reitzii
312; Wettsteinii 311

Smilax bona-nox 149, 165; glauca
149

Solanum rostratum 354

Solidago velutina 264; wrightii
264

Some Observations on Flowering
in Ruellia (Acanthaceae) 200

Sonchus asper 346

Sorastrum americanum 56

Rhodora

[Vol. 64

Spergularia marina, var. leiosper-
ma 345

Sphaeroceptis schroeteri 56

Sphagnum capillaceum 18, 19;
dusenii 125

Sphenoclea zeylanica 156

Spilanthes americana 256, 261,
267; var. stolonifera 267; de-
cumbens 256

Spirodela oligorrhiza 83, 84, 85;
polyrrhiza 85

Spirogyra cleveana 56; ellipsos-
pora 56; hydrodictya 56; ju-
galis 56; mirabilis 56; nitida
56; submaxima 56
Spirotaenia condensata 56

Spirulina duplex 56

Spondylosium pulchellum 56

Sporocybe sphaerophila 134

Stachys floridana 231; tenuifolia
153, 156

Staurastrum alternans 56; avicula
56; brevispinum, var. tumidum
56; cerastes 56; crenulatum 56;
dickiei, var. maximum 56; dila-
tatum 56; furcigerum 56; gra-
cile 56; grande 56; leptocladi-
um, var. denticulatum 56; odon-
tatum 56; orbiculare 56; para-
doxum 56; subgrande, var.
minus 57

Stauroneis acuta 57

Steele, Frederic L., An Unusual
Rubus 354

Steele, F. L. and A. R. Hodgdon,

Glandularity in Rubus Alle-
gheniensis Porter 161; Juni-
perus Horizontalis in New
Hampshire 351

Stellaria graminea 345; media

223, 845

Stigeoclonium flagelliferum 57;
nanum 57

Stripling, H. Larry, Richard G.
Ross and Walter H. Lewis,
Chromosome Numbers for Some
Angiosperms of the Southern
United States and Mexico 147

1962]

Studies in the Guttiferae. II Tax-
onomic and Distributional Ob-
servations on North American
Taxa 231

Surirella elegans 57; splendida 57

Survey of the Algae of Lake
Quinsigamond, A 49

Syncephalantha 4, 9, 10

Synechocystis aquatilis 57

Synedra ulna 57; var. aequalis 57

Synthesis of Aster Herveyi 113

Synura uvella 57

Tabellaria fenestrata 57; floccu-
losa 57

Taraxacum erythrospermum 546;
officinale 346

Taxonomy of Setcreasea ovata,
The 68

Terrell, Edward E. and Walter
H. Lewis, Chromosomal Races
in Eastern North American
Species of Hedyotis (Housto-
nia) 313

Tetraédron limneticum 57

Tetraspora cylindrica 57; lubrica
57

Tetrastrum staurogeniaeforme 57

Thalictrum polygamum 345

Thelypteris rudis 348

Thlaspi arvense 101

Three Misidentified so-called Cor-

dilleran Species in Eastern
North America 117
Thymelaeaceae, Chromosome

Counts of Two 277

Thymophylla 5, 6, 9, 11

Tithonia longeradiata 267

Tolypothrix conglutinata 57; dis-
torta 57; lanata 57

Tortella fragilis 125

Trachelomonas crebea 57

Tradescantia canaliculata 149;
ohiensis 149; leiandra var.
ovata 68, 70; navicularis 149,
154; palmeri 329; reverchoni
149; semisomna 69

Tragia amblyodonta 140; angus-

Index to Volume 64

377

tifolia 139; cordata 358; nepe-
taefolia var. amblyodonta 140;
nepetaefolia var. angustifolia
140; nepetaefolia var. latifolia
140; nepetaefolia var. scutel-
lariaefolia 140; nepetaefolia
var. teucriifolia 140; nepetifolia
var. ramosa 139; ramosa 139;
ramosa var. ? leptophylla 140;
scandens 140; scutellariaefolia
140; stylaris 140, 141; stylaris
var. angustifolia 140; stylaris
var. (alpha) latifolia 140; sty-
laris var. leptophylla 140; teu-
criifolia 139

Tragiola pilsoa 154, 158

Tragoceros americanum 265; mo-
cinianus 265; schiedeanus 265

Tribonema bombycinum 57

Trichaetolepis 8

Trichocoronis wrightii 252, 260,
263

Trichodon cylindricus 122, 123

Tricholoma davisiae 132

Trichomanes radicans 348

Trichostomum cylindricum 123;
tenuirostre 123

Trifolium hybridum 346; pratense
346; repens 346; simplicifolium
61, 62

Trilisa odoratissima 90; panicu-
lata 90

Trimezia fistulosa 309; lutea 308;
Martii 309; martinicensis 307,
308, 310; meridensis 308; Stey-
ermarkii 310

Tripogandra grandiflora 338

Tritonia riparia 307

Trixis californica 270;
259, 261, 270

Tryon, Rolla M., The ferns and
fern allies of Chihuahua 347

Tsuga canadensis 99

Tubiflora caroliniensis 89

Turner, B. L. and J. H. Beaman,
Chromosome numbers in Mexi-
can and Guatemalan Composi-
tae 271

radialis

378

Turner, B. L., M. Powell and R.
M. King, Chromosome numbers
in the Compositae — VI Addi-
tional Mexican and Guatemalan
species 251

Turner, B. L. and Marshall C.
Johnston, Chromosome num-
bers of Dyssodia (Compositae
— Tagetinae) and Phyletic In-
terpretations 2

Two new species of polygala from
Northern Mexico 225

Typha latifolia 284

Ulmus americana 333

Ulothrix tenerrima 57;
57

Urtica dioica 348, 345

Urtica viridis 343, 345

Use of Vapor Fractometry in the
Analysis of Some New England
Hops, The 243

Uttal, Leonard J., Synthesis of
Aster Herveyi 113

Uttal, Leonard J. and Robert W.
Long, Some Observations on
Flowering in Ruellia (Acan-
thaceae) 200

Utricularia inflata 154, 158

zonata

Uvularia caroliniana 90; pube-
rula 90; pudica 90
Vaccinium arboreum 134, 358;

corymbosum 134; macrocarpon
351; macrocarpon forma ebur-
nea 351; macrocarpon, An Al-
bino Fruited Form of 350;
myrsinites 91; myrtilloides 98;
oxycoccos forma leucocarpum
351; stamineum 134, 357, 358;
stamineum var. neglectum 358

Vaccinium tenellum 91; Vitis-
Idaea 341

Vapor Fractometry, The Use of
in The Analysis of Some New
England Hops 243

Varieties of Luzula acuminata,
The 74

Rhodora

[Vol. 64

Vaucheria ornithocephala 57; ses-
silis 57

Verbena canadensis 91

Verbesina crocata 267;
gyrea 267; seatonii 267

Vernonia karwinskiana 262

Veronica persica 154, 158

Vicia augustifolia 346

Vicia Cracca 346

Viguiera dentata 256; grammato-
glossa 267; hemsleyana 275;
longifolia 256, 267; multiflora
256

Villarsia aquatica 90

Viola walteri 152, 156

Vogelia capitata 88

hypar-

Ward, Daniel B., The Genus
Anonymous and its Nomencla-
tural Survivors 87

Water Cress in the New World 32

Weber, Edgar E., Nostoc parme-
lioides in Massachusetts 358

Wedelia brasiliensis 257; filipes
261, 267; parviceps 267

Wells, James R. and T. Richard
Fisher, Heteromorphic Pollen
Grains in Polymnia 336

Werneria nubigena 272, 273, 276

Westella linearis 57

Western Plains Plants in north-
ern Illinois 354

Wikstroemia indica 278

Wilbur, Robert L., Nomenclatural
notes on two Southeastern
Rhynchosias 60

Winterringer, Glen S., Zizaniop-
sis miliacea from Illinois 283

Wisteria frutescens 91

Woodsia scopulina 210

Xanthidium antilopaeum var.
polymazum 57
Xanthocephalum gymnosper-

moides 264; humile 264; line-
arifolium 272, 274
Xeromphalina kauffmanii 133

LOT iil

00341 3868
1962] Index to Volume 64 379

Xyris elliottii 148, 154; torta 149, Zinnia angustifolia 254, 265; ele-
154 gans 265; leucoglossa 265;
maritima 254, 260, 265; peruvi-
ana 265; tenella 266
Zea Mays 136, 338 Zizaniopsis miliacea 283, 284
Zexmenia aurea 261, 268; cos- Zizaniopsis miliacea from Illinois
taricensis 261, 268; fratescens 283
257; pringlei 261, 268; virgu- Zornia bracteata 90
lata 261, 268 Zygnema insigne 57

Youngia japonica 231