S| D CONTRIBUTIONS
TO BOTANY

Volume |

1962-1964

SIDA Contributions to Botany volume 1 (in 6 numbers)
copyright 1962, 1963, 1964
by Lloyd H. Shinners, SMU Box 473, Dallas, Texas 75222, U.S.A.

DATES OF PUBLICATION

No. 1, pp. 1—62: 23 November No. 4, pp. 187—256: 25 June 1964.
ae No. 5, pp. 257—298: 25 June 1964.
No. 2, pp. 683—108: 23 November

1962. No. 6, pp. 299—417: 30 October
No. 3, pp. 109—186: 12 December 1964.

1963.

For Contents, see the unnumbered pages which form the front covers
of the separate numbers.

ERRATA GRAVIORA

p. 29 note 4. Dr. L. O. Williams informs me that the Chicago Museum
acquired the Coulter Herbarium not directly from Coulter but from
the University of Chicago, first on indefinite loan, later as gift.

p. 31 note 19. By egregious oversight the name of Andrew Denny Rodg-
ers III was omitted.

p. 61 par. 3. Second synonym should be Tephrosia purpurea var. angus-
tissima.

p. 78, line 7 from bottom. Add to synonymy Micheliella anisata (Sims)
Briquet in Engler & Prantl, Nat. Pflanzenfam. Teil IV. Abt. 3a: 325.
1897.

p. 100. The three species included under Aneilema become Murdannia
nudiflora (L.) Brenan, M. Keisak (Hasskarl) Handel-Mazzetti, and
Gibasis linearis (Bentham) Rafinesque in the improved classification
adopted by Rohweder, Die Farinosae in der Vegetation von El Salva-

dor, 1956 (Commelinaceae pp. 98—178).

p. 105. For a different opinion on Warea, see “Nomenclatural and taxo-
nomic corrections in Warea (Cruciferae)” by R. B. Channell and C. W.
James, Rhodora 66: 18—26, 1964.

p. 173, key lead 6a. should read as follows:
6a. Lower leaves with flat segments 1 mm. or more wide; upper
leaves with progressively more slender segments . ,
G. rigidula var. rigidula

S | DA CONTRIBUTIONS
TO BOTANY

VOLUME 1 NUMBER 1 NOVEMBER 1962

CONTENTS
Evolution of the Gray’s and Small’s manual ranges
Annual Sisyrinchiums (lridaceae) in the United States

Chromosome numbers of Heals (lridaceae)
in the eastern United State

New names in Arenaria (Caryophyllaceae)

Drosera (Droseraceae) in the Southeastern United States:
an interim report

Key to Southeastern glabrous-styled Tephrosia (Leguminosae)

53

60

SIDA is privately published by Lloyd H. Shinners, SMU Box 473,
Dallas 22, Texas, U.S.A. Subscription price $6 (U.S.) per volume of about
360—400 pages, parts issued at irregular intervals.

©
SIDA Contributions to Botany volume 1 number 1 pages 1—62
copyright 1962
by Lloyd H. Shinners

EVOLUTION OF THE GRAY’S AND
SMALL’S MANUAL RANGES

LLOYD H. SHINNERS
Herbarium, Southern Methodist University, Dallas 22, Texas

“Those who are ignorant of history are condemned to repeat it.” To
suggest that botanists, and especially systematic botanists, are ignorant
of history may seem surprising, exercised as they are with precedents,
priorities, author-citations, and bibliographic rummaging. But their his-
ory does not go beyond the merely chronological or anecdotal. Critical
evaluation, interpretation, explanation, discernment of patterns, prece-
dents for positive actions — there is scarcely even an awareness that
such remoter intellectual levels exist. As for introducing sociology,
philosophy, psychology — a scientists is above such things. He clings
to his scientific purity in his day-to-day work, certain that science can-
not help but progress in the soundest possible way.

The sad truth is that science does not advance purely or even chiefly
by scientific means. It is, after all, simply one form of human cultural
activity, and a victim of the same shortcomings and influences as an
other such activity. What seems to me extraordinary is that systematic
botanists, the nature of whose work should keep them from the extremes
of narrow-minded dogma, remain so stubbornly ignorant of their own
condition. Instead of making a broad-ranging, critical scrutiny of them-
selves, they clutch at devices which will make them respectable in the
eyes of true scientists. But salvation does not lie in cytotax xonomy or

master in his house and not the witless object of accidents and outside
forces. And this requires stepping outside the narrow cultural limits
within which American botany is confined.

In reviewing the history of the familiar “Manual ranges” into which
the eastern third of North America has been divided for over a centu ury,
I wish to point out that the division does not have a scientific basis, has

the progress of systematic botany; that a knowledge of all of them,
changing and unchanging, can enable a botanist to organize his efforts
in a way to extract the most from his opportunities and suffer the least
from his handicaps. I reject the assertion that “the only thing History
has to teach us is that it can teach us nothing.”

SIDA 1 (1): 1—31. 1962.

HISTORICAL RECORD

In 1817 there appeared A Manual of Botany for the Northern States...
to the North of Virginia, by members of the botanical class in Williams
College, Massachusetts. A year later appeared a second edition, Amos
Eaton now acknowledging authorship. The title for this and the two
following editions (3rd in 1822, 4th in 1824) is slightly altered: Manual
of Botany for the Northern and Middle States, but the specification to
the North of Virginia (then including West Virginia) remains. With the
5th edition in 1829 the title becomes Manual of Botany for North
America ... North of the Gulf of Mexico. What this really meant is
revealed in the introductory notice on botanical districts. “The Northern
and Southern districts are separated by a line drawn fom the mouth of
the Delaware River... in a direction to intersect the south end of
Lake Michigan ... leaving all Pennsylvania and the north part of
Delaware, of Maryland and of Ohio, in the Northern district. This
direction of the division line is required, because southern plants extend
to higher latitudes on the western side of the Allegany range, than on
the eastern side.” These two districts are further broken down into
Eastern and Western divisions. “The Allegany mountain is the division
line in the Southern district; a line drawn from the intersection of the
Allegany mountain and the river Potomack, in the direction of Cayuga
lake, is the division line in the Northern district.” For the following
two editions the title and subtitle are unchanged, but this is not true
of the botanical districts. In the 6th edition (1833) no menion is made
of them, the shock of the arrival of Torrey’s edition of Lindley’s Natural
System being so great that Eaton devotes nearly four pages to de-
nouncing it and similar works, omitting the geographic section. With
the 7th edition (1836) passion had only partially subsided. Eaton quotes
with great satisfaction the remarks of W. J. Hooker in praise of the
Linnaean system (in the latter’s British botany). Parts of Hooker’s
Flora Boreali-Americana (a flora of Canada), had appeared, and ac-
cording to Eaton, all the information was incorporated in his revised
Manual. But the only change in botanical districts was a minor one
suggested by Western botanists. “J. L. Riddell, of Cincinnati, has been
his chief guide in drawing the line between the Eastern and Western
regions. The value of the Catalogue of Mr. Gibbs, of Columbia, 5S. C.,
kindly sent by the author, was in great measure lost on account of its
being received too late” (Preface, p. v.). On page 9, under “Location of
Species,” we are told that “S. at the end of a specific description indi-
cates that it grows South of the North line of Virginia, as well as North.
W. (capital) within the parenthesis after a species, indicates, that it
grows West of the Allegany range and its continuation through Cayuga
Lake, &c. — also East of the West line of Missouri and Arkansas.” In
a footnote we are told of this last statement “This limit is authorised
by Drs. Short, Peter, Riddell and Lock.” These comments are repeated

in the 8th edition (1840), now entitled North American Botany; Com-
prising the Native and Common Cultivated Plants, North of Mexico,
prepared by Eaton and Dr. John Wright. There are three new abbrevia-
tions: A. for Alpine, L. (Littoribus) for seashore, O. (Omnibus locis) for
“throughout the Northern and Southern States.” There is a further brief
section headed “Arctic, Rocky Mt., and Oregon Species,” with a con-
fusing second A. for Arctic, R. for “On the Rocky Mt. or west of it; or
between the Mt. and the States of Missouri and Arkansas,” and Cal. for
California. In a footnote in the preface (p. vi), Eaton reports with evident
satisfaction, “These five last editions extended to two thousand copies
each — and one of them to two thousand five hundre

Eaton was a teacher and popularizer of botany, not a botanist by
virtue of original studies or researches. He was not a notable collector,
nor did he attempt to accumulate a good herbarium. Indeed, his final
word (p. 16 of the 7th edition) showed that he never even considered
careful documentation as a method, for he declares that the only way
to assemble adequate geographic data is for every natural history society
“to devote a secure place to the preservation of manuscript catalogues
of all collecting botanists.” He did not travel widely, and his notions of
Western and Southern geography were decidedly naive, as the preceding
quotations show. Despite the inflated title used for editions 5 through
8 of his Manual, he never seriously intended it for use much outside
the area of the first four. Thus his comment in edition 8 (p. 16), after
explaining the abbreviations for Arctic and Rocky Mountain: “These
distant ere Uani le will not embarrass the student; because a solitary R.

pected elsewhere.” It seemed not to trouble him at all that his early,
simple separation into North and South and subdivision into East and
West had become utterly incongruous; he kept on using them until the
last. One suspects that the “distant localities” were thrown in for pos-
sible benefit to sales, a suspicion strengthened by his complacent foot-
note about the number of copies in the earlier editions.

As a compiler rather than an investigator, a rather superficial popular-
izer instead of a critical student, and finally as a reactionary violently
opposing the newer approaches to classification, Eaton did not attract
followers of high calibre. Although Mrs. Lincoln’s Familiar Lectures on
Botany, patterned on his own, continued to be a best seller for years
after his death, he had no real botanical successor in direct line. But
history did repeat itself, though with a difference, in the work of
Alphonso. Wood, whose A Class-Book of Botany first appeared in 1845,
and in a revised edition only a year later. As Wood himself tells us
(preface to the 1860 edition), “It was originally prepared with immediate
reference to the wants of the author’s own pupils, with scarcely a hope
of approval from the community beyond.” It was Williams College all
over again, this time at small Kimball Union Academy near Hanover,

4

New Hampshire. The flora which comprised a major part of the book
covered “that section of the United States which lies north of the
Capitol, that is, of the 39th parallel, including essentially the states
lying north of the Ohio River and Maryland.” Then, with an eye to
sales, it is added, “With some exceptions, therefore, this Flora will
answer for the adjacent states of Delaware, Maryland, Virginia, Ken-
tucky, Missouri, and the Canadas.” Like Eaton, Wood was encouraged
by the commercial success of his book, and, again like Eaton, became
expansionist, with the added stimulus of the desire to keep ahead of

e new rival, Gray’s Manual. For the 1860 edition (3rd copyright date;
unrevised new printings of the 1846 editions were confusingly numbered
as new editions), “The limit of our Flora in this new series has been
much extended. It now embraces the territory lying east of the Missis-
sippi River with the exception of the Southern Peninsula of Florida, and
South of the Great Lakes and the River St. Lawrence... . This Class-
Book is, therefore, now professedly adapted to the student’s use from
Quebec to New Orleans and from St. Pauls (sic) to St. Augustine.”
Unlike Eaton, Wood had actually traveled through much of this large
area: “Therefore, into nearly every section of this territory, from the
St. Lawrence and the Lakes to the Gulf, and from the Sea-Coast to the
Great River, the author has made repeated excursions in delighted con-
verse with the vegetable world.” But like Eaton he was not a notable

g
pected, though he seems to have done more in this regard than did his
predecessor. He too was a compiler rather than an investigator; his
primary aim likewise was teaching and popularizing, not research.
And, once more like Eaton, he had no botanical successor.

In 1824 John Torrey published the first volume of what was to remain
an unfinished work, A Flora of the Northern and Middle Sections of the
United States, covering the same area as Eaton’s early editions: the states
north of Virginia. In 1826 appeared the more condensed A Compendium
of the Flora of the Northern and Middle States, for the same area, this
time described as “north of the Potomac.” Torrey was to set the pattern
for future progress by corresponding and exchanging with European
botanists in order to have critical identifications, and by championing
the Natural System against the Linnaean. His efforts were to come to
full flower in the never-finished Flora of North America (1838-1843),
undertaken jointly with Asa Gray, and in numerous reports on the

in 1848 had the slightly modified title Botany of the United States North
of Virginia. In the same year appeared the famous first edition of Gray’s
Manual of the Botany of the Northern United States, from New England
to Wisconsin and South to Ohio and Pennsylvania. It was destined to

be the last manual for the area that may be called the Old North — that
it, the area north of the Mason-Dixon line and the Ohio River. Scientifi-
cally it carried on the Torreyan traditions of critical identifications and
use of the Natural System. But it had still more important reasons for
being: it would offset Wood’s odious popularity, and affirm Gray’s
position as leader for the critical botanists, and, not least, it would
make money. Prestige, rivalry, and commercialism were to dominate
the subsequent history of the Manual until the 7th edition and, inevit-
ably, of the later Southern floras as well.

We may never know with certainty all the reasons that led Gray to
abandon a regional boundary of more than forty years’ standing, adopted
first by Eaton, and accepted without question by Torrey, Beck, Wood,
and Gray himself. External chance played a part. As late as May, 1855,
in a letter to Darwin, he speaks of “this moderate area (bounded by
the Atlantic Coast, New Brunswick, St. Lawrence, Great Lakes, Missis-
sippi, and Potomac or Chesapeake Bay).” Darwin’s request for informa-
tion about plant distribution seems to have set Gray thinking. When
the 2nd edition of his Manual appeared in 1856 (foreword dated June
30), it had been expanded to include “Kentucky, Virginia, and all east
of the Mississippi,’ an area retained for the three remaining editions
prepared by Gray himself: 3rd (1857), 4th (1862), and 5th (1867), the
last with the range re-worded to “east of the Mississippi and north of
North Carolina and Tennessee.” (There is some confusion because of
various reprintings, the earlier merely as “r evised edition” without

2 a ol i

Northern United States,’ published (1856-1857) just after the appear-
ance of the revised Manual, Gray discusses the botanical reasons for
the change. “The work, which forms the basis of the following statistics
of the botany of the Northern United States, has now been extended in
geographical area beyond the limits of the Northern States, politically
so-called; inasmuch as this area includes Virginia and Kentucky, and
stretches westward to the Mississippi River. The south boundary of 36°
30’ has been adopted (instead of Mason and Dixon’s line) because it
coincides better than any other direct geographical line with the natural
division between the cooler-temperate and the warm-temperate vegeta-
tion, — between the flora of the northern and of the southern Atlantic
states. Few characteristically southern plants advance to the north of it,
and those chiefly on the coast of the low south-eastern corner of Virginia,
in the Dismal Swamp, and the environs of Norfolk. Could we vary the
line where it intersects the longitude of Washington, carrying it north
until it reaches the James River, and thence due east again, the small
quadrangle thus excluded would exclude nearly all the properly south-
ern indigenous plants now comprised in the volume, and mark the true
division eastward between our southern and northern botanical regions,
namely, at the northern limit of the Live Oak, the Long-leaved Pine,

and the Black Moss (Tillandsia unseoides) . .. On the Mississippi, the
plant most southern in character which crosses the parallel is Jussiaea
repens. This sparingly extends up the Ohio to lat. 38°, where also the
Taxodium reaches about as far north as on the Atlantic Coast.” And
prophetically he remarks, “Probably a good many more southern species
inhabit this (southeastern) corner of Virginia, of which I have as yet
no indications.” It was in effect an ante-bellum act of cultural aggression
against the South. It passed unchallenged, among other reasons, because
there were few botanists in the South (and many of these were, like
Darby and Chapman, immigrants from the North), because knowledge
of the details of distribution was still very inadequate, and because the
whole matter was viewed from a strictly Northern standpoint. State
boundaries or parallels of latitude and longitude are hardly ideal for
sean botanical regions. They were matters of convenience, especial-

in view of the limited knowledge of the day. Despite the listing of
species ae plausible sound of his remarks, Gray’s new boundary can-
not be said to have a solid botanical basis. It did, of course, supply more
ample data for answering Darwin’s queries, but that is not the same
thing. Small, with vastly more field experience than Gray, later con-
sidered the Mason-Dixon line a better floristic dividing point.’

Its convenience was certainly reinforced by decidedly non-botanical
considerations. A major purpose in putting out the Manual was to make
money,” and the enlargement of the area covered could be expected to
mean more sales. Whether Gray had by this time gotten wind of Wood’s
plan to annex the whole South in his next edition I do not know. M. A.
Curtis in 1857 warned Gray about Wood’s travels in the South,? and
other friends may well have done so early enough to influence Gray’s
decision on the new boundary. Or perhaps the strategic and commercial
benefits to his continuing war with Alphonso Wood were in this case
pure serendipity growing out of his efforts to give Darwin a satisfactory
answer. I doubt that his motives were pure. Whatever they were, they
rigidly fixed the botanical boundary between North and South from
that day to this. Although he was too cautious to sweep in the whole
South, as Eaton had done before and Wood was to do again in 1860,
he eventually felt compelled to stand up to his rival. In 1869 ee
dated 1868), in his Field, Forest and Garden Botany (decidedly a com
mercial venture), he too annexed the rest of the country east of the
Mississippi River. Meantime he had induced Chapman to write a South-
ern flora, published in 1860, about which more later.

Before taking up the short and simple history of the strictly Southern
floras, we must follow the subsequent history of the Manual and its rivals,
the Britton and Brown Illustrated Flora and Britton’s Manual. Other than
the minor instance of Darby, Southern botany had no independent
existence, but was merely a pawn in the rivalry between Gray and

1 All notes are at end of article.

Wood and thereafter between the Gray Herbarium and the New York
Botanical Garden.

In 1890, two years after Gray’s death, there appeared a 6th edition of
his Manual (copyright date 1889), “revised and extended westward to
the 100th Meridian,’ by Sereno Watson and John M. Coulter. It has
been suggested that the extension was due to the fact that both men
were Westerners (actually Midwesterners by origin, but both had been
active in the Far West). I doubt if the retiring, scholarly Watson would
have made such a departure from precedent had he worked alone. It is
a thoroughly characteristic action of the aggressive Coulter, ever a
schemer, promoter, opportunist, and in general the first major politician
in American botany.* It was really repeating Gray’s 1856 move, this
time toward the West instead of the South. One wonders what might
have happened if, in 1890, two botanists from the South had put out
a new edition of the Manual. I suspect the results would have been
exactly parallel with those of the 1890 edition of Watson and Coulter:
the annexed territory would prove to have included too much that was
foreign to the Old North, and the boundary would have been moved
back. This happened with the 7th edition of the Manual (1908), by
Benjamin Lincoln Robinson (from. Illinois) and Merritt Lyndon Fernald
(from Maine). “To cover a more natural area... some alterations have
been made in the geographic limits adopted in the sixth edition. ... (1)
the exclusion of the territory at the west between the 96th and 100th
meridians, . . . (2) the inclusion of the Canadian provinces of Nova
Scotia, Prince Edward Island, New Brunswick, and the greater part of
Quebec and Ontario.” The more strongly northeastward slant continued
with the 8th edition (1950) by Fernald alone: “The northern limit of
range now includes the area south of the Straits of Belle Isle and from
Anticosti Island westward along the 49th parallel of latitude in Quebec
to the northwestern corner of Minnesota. The western and southern
limits are unchanged.”

There are curious parallels, and even more curious contradictions,
between the events of 1820—1860 and those of 1890—1935. Nathaniel
Lord Britton, Ph.D. (in geology), was the first non-medical doctor to
write a manual, and the first to adopt the metric system. His one-volume

from the Atlantic Ocean Westward to the 102nd Meridian, whose first
volume appeared in 1896. With its crude drawings, slovenly taxonomy,
and outrageous nomenclature, this was a repudiation of everything for
which Torrey and Gray had striven. It was a naked act of imperialist
aggression, not only covering the over-extended range of the 6th edition
of Gray’s Manual, but annexing still more territory to the west and
north. “For convenience,” says the introduction, “the whole of Nebraska
has been included ... a manual of the whole Flora of the northeastern

8

part of the continent, with the exception of that of Greenland and the
Arctic Circle.” A second edition in 1913 took in the same area. This
dreadful production was still being reprinted and sold as late as 1950.
companion work in one volume, evidently intended to displace Gray’s,
and similarly titled (Manual of the Flora of the Northern States and
Canada), appeared in 1901, followed by a 2nd edition in 1905 and a 3rd
in 1907. The area for all was much the same as that of the large
Illustrated Flora: “from Newfoundland and Labrador to Manitoba, the
southern boundary of Virginia, Kentucky and Kansas, and the western
boundary of Kansas and Nebraska.” That Britton’s Manual failed to
displace Gray’s is chiefly owing, I believe, to the lack of illustrations.
The long popularity of the Illustrated Flora was certainly owing to the
pictures, which made it a commercial success at the same time that it
was a scientific failure. Those not wishing to invest in the expensive
3-volume work would naturally purchase instead that 1-volume work
which did have illustrations. There is irony in the fact that the very
ing Britton introduced so lavishly in his Illustrated Flora was also
responsible for Gray’s Manual defeating his own. Certainly it cannot
be said that the general public, whose purchases determined commercial
success, gave much scrutiny to scientific merit. Illustrations and com-
mercial success were the only things the Illustrated Flora and the il-
lustrated Gray’s Manual had in common.
ther it was really necessary to endure the two editions of the
Illustrated Flora for the sake of progress is a matter of conjecture.
Taxonomically speaking they represent the most backward steps ever
taken in American botany. But Britton’s real contribution was the
assembling of the rich library and herbarium resources of the New York
Botanical Garden. It may be that from a crudely practical standpoint a
popular commercial success had to be produced to accomplish this, quite
apart from any question of merit or ethics. In any case, it was possible
for Henry A. Gleason in 1952 to put out a work that repudiated nearly
everything that Britton stood for. The label “3rd edition” is rightly
eschewed for The New Britton and Brown Illustrated Flora of the
Northeastern United States and Adjacent Canada, which for quality
stands in extreme contrast with its predecessors. Brittonian imperialist
aggression is abandoned, though not that of Gray: “Its southern bound-
ary from east to west follows the southern lines of Virginia, Kentucky,
and Missouri. To the west, it extends to the west boundary of Missouri,
Iowa, and Minnesota, and to the north, it follows the northern bound-
aries of Minnesota and Michigan. From the eastern end of aoe ee

it eee See forty-seventh parallel of latitude across On . to
. Lawrence River. . It excludes Anticosti, Newfo er Sabie
aa St. ae and WMichelon: ” A companion 1l-volume Manual by

rthur Cronquist is in press (autumn 1962); presumably its se
area will be the same.
The history of the Southern floras is shorter and simpler, and as has

9

been indicated already, it mainly follows and is subordinate to that of
the Northern ones. Only three authors are involved, all of them North-
erners by origin, only two actually residing in the South. The first was

rby

that of his contemporary, Alphonso Wood. His A Manual of Botany
Adapted to the Productions of the Southern States was published in
1841 at Macon, Georgia, where the author taught at the Wesleyan Female
College. This was reprinted at Savannah in 1847. A new edition appeared
in 1855 (reprinted in 1869) as Botany of the Southern States, published
in New York, with preface dated Auburn, Alabama. Just what was
meant by “Southern States” is nowhere explained. In the text there
are frequent references to the Carolinas, Georgia, Florida, and Alabama,
with occasional ones to “S, Western states,” to Louisiana, and rarely
to Texas. About the author Gray said “he would probably claim to
have a good general, but no very profound acquaintance” with systematic
botany.’ After thus damning with faint praise, Gray proceeded to damn
quite brutally by completely ignoring the systematic section of the book
in his review, devoting his attention solely to the morphological and
physiological portion. Darby’s specimens were lost in shipment — the
same calamity that befell his contemporary S. B. Buckley a few years
later — and it is difficult now to tell whether Gray’s contempt for him
was altogether justified. The hundreds of localities cited by Darby
constituted a really impressive addition to the knowledge of the dis-
tribution of Southern plants at that time, and some comment on this

as surely in order. Later Chapman, dutifully following Gray’s lead,
also pointedly ignored Darby’s work. Darby himself moved to Kentucky
in 1869, after that state had been annexed to the Gray’s Manual range,
and he seems to have given up systematic botany altogether.

r

Sevag took a degree at Louisville, Kentucky (according to Trelease;

art’s footnote in Kimball’s reminiscences does not mention this),
sae spent most of his life practising in northern Florida. In Dupree’s
life of Asa Gray, surprisingly little is told of the relations between the
two men. It is known that Chapman corresponded with Gray, and that
the latter encouraged him at length to write a Flora of the Southern
United States, itemized as Tennessee, North and South Carolina, Georgia,
Alabama, Mississippi, and Florida, published in New York in 1860. In
his preface Chapman remarks, “My original design did not contemplate
so wide a field; but was limited to an enumeration of the plants of the
Carolinas, Georgia, and Florida, — to which, chiefly, my attention has
been directed during the past thirty years. But, influenced by the solici-
tation of friends, and by the apparent need of a more general work, I
have extended my plan, so as to embrace all the States south of Virginia
and Kentucky, and east of the Mississippi River.” He adds that “The
plan of the work is nearly the same as that adopted by Professor Gray,

10

in his excellent Manual of the Botany of the Northern United States.”
He also strongly recommends Gray’s textbooks to his readers. Like the
Manual and his own Flora, these were all published by the same New
York company. It is obvious that Chapman’s boundaries were chosen
to match those of Gray’s Manual. Except for his brief comment about
having been persuaded to extend west to the Mississippi River, he offers
no explanations, botanical or otherwise, for the stated limits. These re-
main the same for the 2nd edition (1883; reprint of the first with sup-
plement) and 3rd edition (1897). Trelease, who visited Chapman the
winter before the latter’s death, quotes these remarks by him regarding
new species: “But, you know, even if I were not at the end of my work,
I should prfer someone else to name them. I never did care to name
species, as so many others do.” In the preface to the 2nd edition of his
flora he had said, “And now, since the different sections of all the
States which are included have been pretty thoroughly explored, and
future acquisitions will, probably, be comparatively few in number,”
indicating that he did not much believe in new species anyway.

It is an interesting example of the role of chance or coincidence in
history that both Chapman and Small were color-blind to shades of
red.® But their ideas about new species were in violent contrast. John
Kunkel Small, native of Pennsylvania (his surname was an Ameri-
canization of Pennsylvania Dutch Schmal), never lived in the South,
though he made numerous and sometimes extended visits there, chiefly
in the Atlantic states, especially Florida. Employed by Britton primarily
as curator, he was also expected to implement Brittonian imperialism,
with the South (and later the Southwest) as his special territory. Legend
has it that Britton, fearful of a destructive fire (it was in the days of
gas lights), allowed no one to work at the New York Botanical Garden
after dark. He also expected Small to devote his daylight hours to
curatorial duties. Small had a large and musical family (he himself had
once been flute-player in the New York Philharmonic), and after supper
there was a performance by a family orchestra until bed-time for the
children. Work on his Flora of the Southeastern United States (pub-
lished by the author, 1903; followed, like Chapman’s, by a reprint with
supplement as 2nd edition, in 1913) is said to have been carried on
during midnight hours, and on the basis of no more than one specimen
of each species or sometimes even genus. Those who have used the
book will find the story wholly believable. Its taxonomy and nomencla-
ture are thoroughly typical of the Britton school. It was another weapon
of political warfare, not a work of careful scholarship. Understandably,
its geographic area extends west to parallel that of the Illustrated Flora,
but for some reason the 102nd meridian was given up for the 100th,
thus matching the limits of the 6th edition of Gray’s Manual. No ex-
planation for the choice is given; neither line makes any sense botani-
cally. It seems to have become clear eventually that the area was over-
extended and instead of a 3rd edition there appeared in 1933 the Manual

11

of the Southeastern Flora (reprinted in 1953 by the University of North
Carolina Press, the first production of a manual in the South since
Darby’s of 1847), reverting to exactly the limits of Chapman’s floras.
The parallel with the similar reversion to earlier and narrower limits
for the 7th edition of Gray’s Manual is striking. But imperialism was
not being abandoned. There was to be a Manual of the South-Central
Flora to keep the states west of the Mississippi River in the Brittonian
fold, and parts of it were actually written.’? But the driving hand of
Britton had been removed with his retirement in 1929, Small himself
was nearing the end of his life, and his successor-designate, E. J.
Alexander, belonged to a generation among whom writers of regional
manuals were virtually extinct, and authors even of whole state floras
very rare.®

We may round out the account of Southern floras with brief notes on
another abortive one, and three recently initiated. Less powerful and
aggressive than Britton, but like him a representative of the Age of
Empire Builders, William Trelease early sought to make the Missouri
Botanical Garden a Gray Herbarium of the West, specifically preparing
the ground for a Manual of the Southwestern Flora, which was to have
been written by J. M. Greenman. Trelease himself collected in Louisiana,
Arkansas, Oklahoma, and Texas, and made taxonomic studies of such
typically Southwestern groups as Agave and Yucca. He hired numerous
collectors, and bought up all available private collections, especially in
Texas. Greenman’s students were put to work on revisions of genera
prominent in the region, and until his retirement he used to state in his
annual reports that “progress has been made toward a Flora of the
Southwest.”” Unlike the rival manual of Small, not a page of it appears
ever to have been written. Seemingly all the ingredients for effective
results were there: herbarium and library facilities, institutional back-
ing, a trained taxonomist, abundant help, advance planning. Yet the
result was complete failure.

History is once more repeating itself, though not exactly; how great
the difference will be remains to be seen. Once more broad-scale plans
have been made, and facilities and personnel accumulated, this time for
an innovation and an anomaly in American botanical history; a regional
flora for the Southeast carried down to genera only. It is an innovation
and an anomaly on several other counts: it is being done at Harvard,
which had never before attempted anything like it; and its geographic
limits, taking the Chapman and later Small area with the addition of
Arkansas and Louisiana, match neither a predecessor nor a natural area.
When first announced in the AIBS Bulletin for April, 1956 (p. 26), this
was to do more than any previous flora. “This massive project, planned
to locate and identify every kind of vascular plant in a thousand mile
square area, will have the cooperation of botanists in several southern
universities. The study will cover the plants of Alabama, Arkansas,

12

Florida, Georgia, Louisiana, Mississippi, North Carolina, South Carolina
and Tennessee. C. E. Wood, Jr., of the Arnold Arboretum and Reed C.
Rollins of the Gray Herbarium will conduct the study. The project,
which is expected to take 20 years, was first proposed and supported
by George R. Cooley, a retired Albany, N. Y. banker who has devoted
years to the study of the southern flora. The project will now also have
financial assistance from the National Science Foundation, Three separ-
rate treatises on the vascular flora of the southeast are planned,” two
volumes to be devoted to the wild flora, a third to the cultivated plants.
Twelve cooperating botanists are listed, seven of them permanent resi-
dents in the South. For immediate results, it was decided to prepare a
generic flora, published as a series of articles in the Journal of the
Arnold Arboretum. In my understanding of the term, a “seneric flora”
is not a flora at all, that word properly signifying an account of species.”°
In any case, the “generic flora” (currently appearing at a speed which,
if maintained, will require well in excess of another half century to
complete) is not comparable with the other manuals and floras here
discussed, That it is also incongruous from a historical-cultural view-
point will be very evident from my following accounts of backgrounds
and perspectives.

Lastly there are two regional floras now being worked on by myself.
The first in conception (but likely to be last in execution) was a Flora
of the Gulf Southwest, intended more or less to take the place of the
abortive manuals of Greenman and Small, covering Arkansas, Oklahoma,
Louisiana west of the Mississippi River, and Texas east of the Pecos. The
boundaries, mostly artificial, were adopted in the belief that it was best

urgent to include parts of the states covered by them. In other words,
the old method of arbitrary lines was followed out of precedent, utility,
and convenience. Purely botanical reasons were involved only in ex-
cluding Trans-Pecos Texas; otherwise science had nothing to do with it.

It becomes obvious quite early that a Flora of the Gulf Southwest
could not be completed without a great deal of study in the states east
of the Mississippi River. After some tentative probings in the form of
field trips and synoptical studies of wide-ranging groups, a number of
ideas gradually took shape. Many of these have been summarized and
documented in my synopsis of Bonamia, written more for that purpose
than for taxonomic reasons. The major conclusions to be stated are
first, that a generic flora fills no real need — what is urgently required
is a working manual of the species, in the Torrey and Gray tradition
instead of the Brittonian, with nomenclature according to present rules;
second, that we have progressed sufficiently to begin using botanical
boundaries for the Southeast instead of the traditional arbitrary ones;
third, that we have now developed library and herbarium facilities that

13

make it possible to write Southern floras with little dependence on
outside resources; fourth, that floras get written because one person
makes up his mind to do it, not because of mass-planning. I am there-
fore actively working on a concise Flora of the Southeast as well as a
Flora of the Gulf Southwest.

HISTORICAL BACKGROUND: CULTURAL

Perhaps the first thing that strikes one about the historical record is
the abundance of authors and floras in the North, and their paucity in
the South — and the futher strange fact that all three authors of
Southern floras came from the North. This is exactly contrary to what
the much greater richness and diversity of the Southern flora would
lead one to expect. Surely, one would think, the stimulation offered by
that flora, and the long blooming season, ought to have resulted in far
more interest in and study of the plant life by those born in the South.
Plainly the reasons are not botanical. That social, cultural, and economic

state of affairs be readily explained in terms of general history, but
even small details can be traced to non-botanical origins.

To begin with, American scientists have been overwhelmingly of lower
and middle class origins. The rich and aristocratic rarely have cultivated
science to the extent of making significant scientific contributions, though
some have been financial patrons. We can see at once a major reason
why the North alone supplied all the authors of regional floras. Although
there existed a landed aristocracy in the Old North, there was never
the social cleavage that existed in the South. It was a region of grass-
roots democracy, with a comparatively homogeneous population of

amid such cultural conditions? Especially when the shrewd Yankee
knew there would be plenty of buyers for his books. There seems to have
been an ideal level of urbanization and commercial growth at which
the countryside was still familiar, and the pursuit of Linnaeus’s harm-
less science was among the many little luxuries now widely available.
Not until the next century would extreme urbanization make inroads
into the serious pursuit of botany by non-professionals, and it would
then become difficult to disentangle the internal complications, arising
from specialization and the rise of more technical aspects of botany, from
external blighting influences,

After the Civil War two new factors strongly influenced the develop-
ment of American botany. One was the appearance of the newly-rich

14

‘“malefactors of great wealth” with the great increase in Northern in-
dustrialization, The Gray Herbarium was to benefit modestly from the
new Northern wealth; Britton was to exploit is as no botanist before
or since. He used it to build the New York Botanical Garden and to
hire Small; much of Small’s field work in Florida was directly spon-
sored by wealthy individuals. In the South, only the short-lived Bilt-
more Herbarium near Asheville, North Carolina, came out of the post-
war Northern wealth, but this was hardly more than a plaything of the
Vanderbilt family.” It had scarcely begun to function when it was
largely destroyed by a flood, never to be revived; the surviving remnants
were eventually turned over to the U. S. National Herbarium.

The second post-war influence was to be very slow in taking effect.
This was the rise of a powerful, centralized, Federal government. Ironi-
cally, despite long and violent (and still continuing) opposition to cen-
tralized authority, the South was to benefit far more than the North
from the activities of Federal agencies. In terms of botany, this meant

worth noting here that every one of the regional floras and manuals
for both the Northeast and the Southeast was entirely a private venture.

In contrasting Southern with Northern conditions, it must be kept
in mind that there were really two Souths, or rather three, if Texas is
taken into account, This has fateful consequences in the history of
Southern botany. The semi-fictional Old South is the Plantation or
Lowland South, whose earliest flowering was in Tidewater Virginia.
This was dominated by a landed, slave-holding aristocracy which was
itself slave to climatic conditions, “King Cotton,” and the British textile
industry. Before the one-crop economic system had developed a strangle-
hold (which it did, ironically, with notable help from a Connecticut
Yankee: Eli Whitney, inventor of the cotton gin), this aristocracy could

this was increasingly as part of the social graces, and fitted more and
more to the conservative outlook of a leisure class. One did not stoop
to the kind of vulgar grubbing that Yankees did; one did not write text-
books, nor get into squabbles about new systems of classification. That
before the Civil War Henry William Ravenel of South Carolina, owner
of 32 slaves, should study the local flora and put out notable exsiccatae
of fungi, was extraordinary — indeed, unique. But Ravenel was only
modestly rich, and after the war it was dire need that led him to resume
collecting botanical specimens for sale. The deep shock of defeat and
ruin did not induce the planter aristocracy to turn to such things as the
pursuit of botany for its own sake. They bided their time, eventually
regained political control (fossilizing it in the style that until today has
kept large city populations under control of rural counties from Georgia

15

to Texas), and promoted a romanticized image of themselves as the
true and only Old South.

The intellectual history of these people is among the most absorbing
and perplexing subjects confronting the historians. Because it is so
vividly illustrated in the history of Southern botany, it deserves extend-
ed comment here. The blighting influence of a slave economy and a
dominating oligarchy, so different from Northern conditions, seems
obvious. But it is not that simple. The ancient Greeks had a slave
economy and ruling oligarchies, but cultural blight was conspicuously
not a result. And indeed the Old South in its earlier period did not dis-
play the intellectual sterility’* that characterized it during the height

wer h

Stephen Elliott’s Sketch of the Botany of South-Carolina and Georgia
(1816—1824, 2 volumes) stands even today as one of the finest local
floras ever written in the United States. It was the work of a South
Carolina banker who had no training in botany. That the Old South
never again showed itself capable of producing anything of the kind
was but one facet of a growing mental rigidity’ whose roots were
certainly in the South’s “Peculiar Institution”: Negro slavery. This was
a very different thing from slavery among the Greeks. Slaves with
them were often prisoners of war, not men born to slavery, and not
irrevocably doomed to remain slaves; racism was not involved. And
this was slavery in the midst of bustling commercialism, centered in
city-states; with an atmosphere quite unlike that of the rural Old
South, which remained culturally a frontier region until quite recent
times, and was antipathetic toward the rude commercialism so typical
of Yankees. The social rigidity imposed by slavery and racism en-
gendered mental rigidity, which was greatly intensified in resisting the
forces that were undermining the entire slave economy. ough many
even in the Old South believed that slavery would ultimately die a
natural death, very few there were eager to see a way of life that was
good to them disappear.

Despite the very successful efforts in politics and propaganda by the
Plantation South, there was and is another South: the Upland South,
or the South of small farmers ake owned no slaves. These people were

were to the Plantation South. That they differed so greatly in intel-
lectual interests and activities (not in abilities; the falsity of the hill-
billy legend has been demonstrated by the events of World War II
and after) may be explained in part by the fact that they still lived
under frontier conditions for decades after the North had become

tivities of the Federal government during Reconstruction was the

16

establishment of public schools and the importation of teachers. But
when the men of the Old South resumed control, they showed little
enthusiasm for such things.’® Illiteracy was in their view a desirable
thing for the non-slave-holding whites and the former slaves both.
Simple lack of education made it impossible for botanical manuals to
be written or even read by the very people in the South who, judging
by what had happened in the North, were most apt to have produced
them.

The men of the Upland South were intensely loyal to their home
states, most of them supporting the Confederacy without question in a
war that was not in their own best interests, and falling in with the
political schemes and racism that dominated Southern history after-
ward. But there were notable exceptions. Eastern Tennessee made one
unsuccessful attempt to set up a new state, and “Tennessee Johnson,”

leaders of the Confederacy, who did not belong to his South. West
Virginia, of course, is the conspicuous example of the split between
Upland and Lowland South, carried to successful political outcome for
the former. North Carolina was never so dominated by the plantation
aristocracy as were her eee states, and Florida, which for long
was sparsely settled, likewise was never fully a part of the legendary

ld South espite the at catch-phrase ‘‘the Solid South,” the
South is not and never was such a unity.

We shall review the significance of all this in the current and future
history of Southern botany shortly. But first we must take up the
special case of Texas, which, although a member of the Confederacy,
is not a typically Southern state, ee or otherwise. It is as
ae Western as Sou n, but most of all it is just Texas, never for-
getting that for i years before ee one of the United States it
was a sovereign nation. The Plantation South and negro slavery did
spread into the southeastern part of the state, but never attained the
level they did in Louisiana and states east of the Mississippi River.
Another landed aristocracy, the Cattle Barons, did develop farther
west, but by the nature of things it was not numerous nor old, and
without slavery did not develop into anything like the Old South. The

of the Edwards Plateau with its many springs. The settlers before the
Civil War were diverse: many from the Upland South, some from the
Middle West, great numbers from Germany and other European coun-
tries. In social and economic terms, these people belong with those of
the Old North and the Upland South; despite geographic proximity and
political association with the Plantation South, they are not an integral
part of it. Noteworthy is the fact that of the seven original (pre-

17

Sumter) states of the Confederacy, only Texas submitted the Ordinance
of Secession to popular vote; in the other six it was passed by the ruling
oligarchy.

ll of the South remained impoverished for long after the Civil
War, although Texas did not suffer as much as most. Nevertheless the
“Big Rich” so much talked about nowadays are a quite recent phe-
nomenon, as well as a minority. They do, however, mark the first

it was a complex and varied group of developments in the direction of
urbanization and industrialization, given enormous impetus by World
War II. We need not go into details. It is enough to point out that
although far from having equalled the North, the South is now well
along in an economic boom very similar to that experienced in the
North a little before and more especially after the Civil War

When we look back at the history of the various botany manuals, it
is easy to see how much of that history is only a manifestation of the
general history of the times: of social conditions, economic develop-
ments, intellectual climate. Purely botanical matters are of extra-
ordinarily little consequence. Eaton’s manuals were popula r and in-

the time and the region. But they did not establish either the profes-
sional practice or the reputation of American botany. Torrey and Gray
did successfully introduce the Natural System, and accomplished much
good work; yet the spirit of the age which followed them was such that
their efforts to establish a tradition of sound scholarship failed, and
their ee of accomplishing the basic task of writing a complete flora
of the country were never realized. The Northeast, thanks to its long
aaa tradition, devotion to public education, sympathetic intel-
lectual climate, and earlier and greater economic development, is now
supplied with good, recent floras which reflect the benefits of repeated
revisions and the prolonged and intensive work of many hands. The
South, because of internal social and cultural conditions, had to have
its few and very inadequate floras written by outsiders. But social,
cultural, and economic conditions are not static, and we have now
reached a stage at which we can begin to see the shape of things to
come, and the reasons for them. But before proceeding to diagnosis and
prognosis, we must briefly fill in the pertinent scientific background.

HISTORICAL BACKGROUND: SCIENTIFIC

The entire history of the manual ranges is one of inadequate progress
with the rather elementary job of compiling a catalogue. There is
virtually nothing that could be called intellectual development in it,
such changes in concepts or techniques as appeared being of external
origin, the majority coming from Europe. Following is a list of those
developments which were mainly scientific in origin or nature and

18

which influenced the manuals. Not that they were purely scientific, of
course. All had in greater or lesser degree contributing cause in con-
temporary social and economic conditions, but are most conveniently
discussed in tems of their manifestations in the scientific world.

he Natural System.—Linnaeus himself considered his simple pro-
cedure of counting stamens and pistils no more than a convenience, to
be replaced some day by an approach using many characters. Many of
his followers with more limited mental horizons (Eaton among them)
would have preferred to keep things simple forever. But the Linnaean
System became more and more obviously unworkable as knowledge of
the world’s flora increased. The organization of that flora into orders
and families based on many features, a work almost entirely carried
out by Europeans, required much more in the way of critical study and
evaluation than before. Torrey’s promotion of the Natural System in
American botany was but one aspect of his efforts to create a truly
critical science on this side of the Atlantic. With its establishment
American botany for the first time acquired intellectual substance,
however modest.

Darwinism.—Although Asa Gray himself was a leading champion of
the theory of evolution, acceptance of it brought no change in his ap-
proach to classification. A species was treated as a morphological type
which for all practical purposes was constant. Not until the new sciences
of genetics and ecology had been born and made some growth could
there develop the concept of a species as a population which might
include considerable variation. All the manuals and floras that have
been published for the two ranges so far have been quite uninfluenced
by evolutionary theory. While this was largely by default (only Gleason
among the various authors had experience with intensive work in
ecology, or had given attention to the philosophical bases in his re-
search), it is as things should be, for the proper function of a flora is
to record facts and make them accessible, not to theorize. This point will
be elaborated further under Neo-Darwinism.

Nomenclatural Codes.—This of course means chiefly the American
Code, which represented neither profound thought nor a deep desire
to aid science. In its extremism with regard to priority, going outside
the genus to find the earliest names for species and creating unnecessary
new combinations (in contrast with what Fernald would later refer to
acidly as “the sensible and therefore discarded Kew Rule,” under which
only names already existing within the genus had to be considered),
it followed the preachings of a few Europeans lke Otto Kuntze in
Germany. In its use of undesignated trinomials, it borowed from zoo-
logical practise. Its type method, commonly held up as a eieal Ameri-
can contribution, was simply a refinement of the “preuves” of Alphonse
de Candolle’s La Phytographie.

Far more significant than its content were the concealed purpose of
the Code and the manner in which it was promulgated. It was here

19

that Britton” stooped his lowest to get ahead of the Gray Herbarium.

Club, American Association for the Advancement of Science,” a hitherto
nonexistent group, made up of henchmen rather suddenly and mys-
teriously appointed, to put out a List of Pteridophyta and Spermatophyta
rowing without Cultivation in Northeastern North America employing
the hundreds of unfamiliar names required by the American Code. In
the preface it is stated with bland mendacity that “the general rules on
which the list has been compiled are in accordance with the views of
the great majority of North American students of systematic botany.”
Britton was chairman of the committee, which included no. members

addition of the States of Kansas and Nebraska, and the Canadian
Provinces from Manitoba to Newfoundland.” This of course was the
territory to be covered by the Illustrated Flora whose first volume

of the new would have to buy the Illustrated Flora and later Britton’s
nual. The American Code was a Brittonian device for achieving
political power and commercial advantage.

In a period of great corruption in public life, this kind of thing was
not unusual. The growing imperialist sentiment of the time, soon to
erupt in the war with Spain and to be personified in Teddy Roosevelt,
found added attraction in something specifically named the American
Code. Britton very successfully exploited attitudes and techniques of a
society whose ideals were far removed from those of Torrey and Gray.
Eventually there were compromises; the International Code of today
includes some features from the American one, and is the only code in
use. The two most recent Northern floras are essentially alike as to
nomenclature, but differ considerably from any predecessors. The South
still suffers with Small’s Manual, nominally following the American
Code but not consistent in that respect, and very different from the
current Northern counterparts.

The Rise of Technical Botany.—During Britton’s own lifetime there
occurred a great development in non-taxonomic fields of botany, es-
pecially those employing the microscope and the laboratory. These were
not involved in disgraceful nomenclatural squabbles, had the attrac-
tion of newness, and the appearance of being more truly scientific than
taxonomy seemed to be. There is supreme irony in the fact that before
he reached retirement, the very science in which Britton had tried to
make himself supreme had fallen into disgrace, in great part as a direct
result of his own actions. He had made himself the leader not of Ameri-
can botany, but only of a discredited segment of it.

Much of the newer technical botany had little to do with the prepara-

20

tion of manuals, and botany became more and more a study for special-
ists. But at least three fields were to have much to do with the attitudes
and methods of future writers of manuals: genetics, cytology, and
ecology. Only with the aid of these would it become possible to develop
the concept of species as population rather than morphological type. But
one still of necessity recognizes a population as belonging to a secies
more by the visible morphology than anything else. Despite much
ballyhoo about the ew Systematics,” sound taxonomy is not so very
far removed from what Torrey and Gray tried to do. This is especially
true when progress has hardly advanced out of the primitive stage, as is
true of our knowledge of the Southern flora.

Neo-Darwinism.—This exists in its most virulent form among modern
vertebrate zoologists, who by virtue of having simple and limited

measure of the weakness of American systematic botany, reinforced by
modern mass culture and the urge to conform, that the botanists have
accepted ideas and assertions which do great harm to the sound progress
of plant taxonomy. Just as Brittonism had exploited taxonomy as an
instrument of political and commercial warfare, so Neo-Darwinism has
exploited it as a vehicle for the pseudo-science of phylogeny. Indeed,
it has even come to be taught as dogma that the purpose of taxonomy
is to construct a family tree, not to be taxonomy at all. Basic taxonomy,
as I conceive it, comprises a body of factual information, and serves
as the basis for many other kinds of study. Phylogeny does neither; it
only diverts taxonomy into a bastard activity between science and
fiction. After the twin calamities of Brittonism and phylogeny, it is a
marvel that any honest taxonomy still exists.

An important difference between Brittonism and Neo-Darwinism is
that while the former still took into account the general public, the latter
is meant only for the professionals. It is doubly hostile to the preparation
of floras, and ought to be rigidly excluded from consideration by any-
one engaged in such work

Over-all Scientific Background.—It has been said that the 19th was
a Biological Century, while the 20th is a Century of the Physical Sci-
ences. There is no question but that writing floras was a more common,
more acceptable, and much easier activity during the 1800’s than now.
In today’s intellectual climate, so heavily dominated by the physical
sciences and mathematics, with confused overtones of war and space
travel, the peaceful writing of mundane floras finds little encourage-
ment. The weak science of botany seeks to borrow strength by adopting
techniques from those more powerful. I think it would gain more
strength if it sought a clearer understanding of its condition and a
firmer grasp on its proper business. At this stage in our progress, I be-
lieve the writing of floras is an ideal means of doing both.

21

Incidentals.—Though I have gone to some lengths to demonstate how
cultural and economic conditions lay behind the greater success of
Northern botany, there is at least one contributing factor in the nature
of the materials being studied. The flora of the Northeast is smaller
and simpler than that of the South. It also has much more in common
with that of Europe than does the Southern flora, and the work of
European botanists certainly helped to make easier the task of those
writing floras for the North. On the other hand, today’s Southern
botanists have benefits from rapid transportation and communication
that facilitate their work in a way never before possible.

HISTORICAL PERSPECTIVES

History is the product of forces. Enough has been said, I think, to
demonstrate my thesis that botanical history—specifically illustrated by
the evolution of the two manual ranges—is only a minor part of cul-
tural, social, and economic history, and its progress is subject to a host
of influences quite remote from itself. Yet the history of a science ought
to differ from general history, since science has (at least in short-range
view) concrete, specific objectives. Why then has American botany been
so completely the victim of circumstances? The answer I believe lies
in e failure of American botanists to view themselves and their
activities scientifically.

Surely the first task for American botany was to get the country’s
flora written up and made accessible to everyone, and this task is still

practical convenience. But after three generations of Southern floras
and more of Northern ones, we have surely progressed far enough to
demand something better. We particularly need to have the dividing
line scrutinized from the Southern side, not just the Northern.

The imposing array of authors and manuals for the North is matched
by an even more imposing array of large herbarium and library collec-
tions. The South still lags, but not nearly so much as before. Good and
recent herbarium collections are available in quantity at institutions
within the South, and although library facilities are less adequate, there
is no longer the complete monopoly once enjoyed by the North. Facilities
are not a barrier to progress at present,

Personnel is another matter entirely. Not because of small numbers;
never before have there been so many Ph.D.’s. But it is precisely here,
ironically, that most of the trouble lies. The character, attitudes, and
instilled beliefs of the modern American professional botanist are largely
inimical to the writing of floras. This means that the most immediate
and influential intellectual atmosphere is likewise inimical to such
work. We have come back to cultural and social influences, this time
at a more immediately personal level.

22

Must we at this point give up hope of any further scientific evaluation?
No; if human history is indeed the product of forces, we have some
possibility of the detection and objective evaluation of those forces. In
fact we have already done so in recounting the past history of the two
manual ranges. One might be led to predict then that their future his-
tory will simply continue to reflect the functioning of largely irrelevant
forces, and nothing scientific can be done about it. Nothing purely
scientific, perhaps, but having demonstrated to what an extent botany
is a cultural pursuit, we need not be thus restricted. Let us try to
discern some of the influences that are harmful, and need to be resisted,
and some that are favorable and need to be aided.

In speaking of Britton and the Age of Empire Builders, I briefly
indicated the non-botanical context: the Robber Baron era of corrup-
tion, monopoly, power politics, and expansionism, Their work repre-
sented a direct importation into “pure” science of contemporary cultural
attitudes and activities such as had never before taken place. The next
period, overlapping in time with the Age of Empire Builders, I have
called the Age of Dilettantes. This was in part a reaction against the
immediately preceding period, in part an acceptance of it. The Empire
Builders went in for regional and even continental floras, dabbled in
all groups, did work of generally poor quality, championed the Ameri-
can Code and all the needless nomenclatural upsets that went with it,
and left to their heirs immense herbarium and library facilities — along
with a ruined reputation for plant taxonomy as a science. Their heyday
saw the great rise of other fields of botany, so strikingly illustrated in
the career of that great opportunist, John Merle Coulter. Laboratory
botany benefited hugely from the loss of sound scientific standing by
taxonomy. e Dilettantes who succeeded the Empire Builders no
longer wrote general floras, but specialized (sometimes very narrowly),
did work of generally high quality, supported compromises and the In-
ternational Code, took their rich working facilities pretty much for
granted, and — being themselves all Ph.D.’s — went on grinding out
Ph.D.’s more numerous than distinguished. The trend toward more
careful work received an opportune boost in the form of cytological
methods, a European innovation which fit in perfectly with the Ameri-
ean flair for push-button gadgets and had altogether phenomenal suc-
cess in the United States. Taxonomy began to be restored to favor in
the eyes of the laboratory scientists.

Now another cultural trend is beginning to be prominent: the drift to
stereoptyed mass-culture, in part arising from the constant increase in
population. In terms of botany, it means more and more Ph.D.’s being
ground out according to remarkably uniform pattern — taught out of
mass-produced, standardized textbooks, in stereotyped courses. Tax-
onomists grow up with dogmas (myths, really) floating in the air,
never set down in clear terms and never critically examined or even

23

questioned: taxonomy had gone bad; one must lean over backward to
avoid publishing unnecessary names or describing too many species; one
had to specialize in order to be really scientific; one did not waste time
on introduced plants; phylogeny was the real thing, work on general
floras was bad. And in line with the growing American tradition, one
mustn’t criticize. The Age of Dilettantes is passing into the Age of
Conformists, in which often excellent but specialized work goes hand
in hand with uncritical conformity and intellectual shallowness.

What I have described is really the cultural history of Northern
botany. For the South there was the long stagnation from the 1830’s to
the 1940’s. But the illustrious early period with Stephen Elliott, and
the immense spurt of activity since World War II, make it plain that
the long stagnation was something forced by special factors, not evi-
dence of lack of capacity. By reason of this very gap in its past, the
New South has a lively history ahead of it, and we can not only see that
history beginning to take shape; we can perceive the details and the
reasons for them. It is no accident that two of the four largest her-
bariums in the South are in Texas and two in North Carolina; that
those in Texas began their current growth earlier; that there are more
native sons among the taxonomists in those two states than in any
others in the South; that a descriptive flora of West Virginia is nearly
completed, while the Old Dominion has nothing comparable now or
in prospect; that Florida, North Carolina, Tennessee, and Texas lead
in the number of active botanical centers. These are wholly expectable
consequences of economic progress’* in areas least dominated by the
Planter Aristocracy. But the whole South is becoming urbanized and
industrialized, and the pattern of change keeps spreading and intensi-
fying. Historians of the future may find the current progress of botany
at the University of Georgia even more notable than the examples
just cited.

The South’s needs botanically are decidedly regional, something out
of step with the national trend to uniformity. The South has also been
a stronghold of individualism as well as states’ rights, again out of
step with the national trend to mass-culture. And more than any other
section, it has a great awareness of history.!® These ingredients in the
intellectual atmosphere are added sources of strength to those who
have the special advantages, in performing belatedly a necessary sci-
entific task, of modern facilities and, not least, the lessons of history.

In so applying the lessons of history, something new is introduced
into American science, Plainly the opportunity before Southern botany
is not simply to imitate what has been done before, or to fall in with
the Age of Conformists. The more independent it is, the more it draws
on those regional cultural peculiarities that offer strength and en-
couragement, the greater its scientific achievement will be

APPENDIX ONE
CALAMITY AND SOUTHERN BOTANY

Chance undoubtedly plays a part in history, and in the history of
Southern botany, ill chance has figured more largely than for any other
section of the country. Jones and Meadows, in tabulating American
institutional herbaria, note that seven had been partly or wholly de-

these may be added the Biltmore Herbarium in North Carolina, partly
destroyed by flood, and never restored. For one half-century period
alone, a quarter of the country was victim of more than half the major
catastrophes, In the 19th Century, important collections made by Darby
and Buckley were lost while being shipped north. The deliberate de-
struction of most of Rafinesque’s herbarium must also be counted as
a Southern calamity, for Rafinesque had made particular effort to obtain
Southern collections, and had named many species from the region.
Having at all times much less than the North or the Pacific states, the
South could not afford to lose even an equal amount with them, but
it lost much more.

The premature deaths of men of great promise has darkened the
history of botany in all sections, and the South had its full share —
Thomas Walter, Hardy Croom, W. B. Fox, to mention but one for each
of the three centuries in its history. As with the lost collections, the small
numbers of those who pursued botany in the South made the loss of
each one greater than if it had occurred elsewhere.

In commenting on Reinhold Niebuhr’s thesis that the American
people have had too happy a history, and are therefore not really pre-
pared for world leadership, C. Vann Woodward points out that the South,
having suffered defeat and ruin in the Civil War, is different and unique
in just this respect. It is a strange coincidence that the history of South-
ern botany, quite apart from the episode of the War, should be more
touched with tragedy than that of any other section.

APPENDIX TWO

THE PATTERN OF ALIENS AND FRONTIERSMEN
IN SOUTHERN BOTANY

What is a Southern botanist? The best answer I think lies in Oswald
Spengler’s conception of race, as the manifestation of a distinctive cul-
ture, not a matter of blood. “It is what one has, not what one is... . The
one is ethos; the other — zoology.” This hardly fits the dogma of Anglo-
Saxon supremacy and the preoccupation with family ties so typical of
the Planter Aristocracy of the Old South. As a matter of historical
record, even they did not practice what they proclaimed. There was

25

the embarrassing French origin of the old Huguenot families who were
numerous and prominent in South Carolina high society; but at least
they were old, and the definition of Anglo-Saxon could be stretched
just enough to include them. General Beauregard, handsome and with
polished manners, in charge when Fort Sumter was fired upon, popular
hero and darling of Charleston society, was a Creole from Louisiana.
Jefferson Davis, President of the Confederacy, came of an upstart West-
ern family, only one generation settled in Mississippi, and while re-
garded with condescension and mistrust by the older aristocracy, none-
theless was made President, later to become not only apologist but
prime symbol of the Plantation Aristocracy.

Such unsettled social conditions of course reflect the frontier status
in which the South remained so long. The pattern in political life was
duplicated in botany, and has been consistently followed for two
hundred years. Thomas Walter was an English immigrant, Stephen
Elliott, a native son; Darby and Chapman were Yankees, while Rugel,
Gattinger, and Mohr came from Germany, and the two Michaux from

an is special attraction for botanists in frontier country
with still ae plants, of course. But there is also a non-scientific
factor in the attraction of frontier conditions for individualists, and tax-
onomic botany has been overwhelmingly carried on by men who were
strongly individualistic. A running theme in Geiser’s eanatee of
the Frontier is that of gifted men crushed by unfavorable frontier con-
ditions, men who in more civilized surroundings might have risen high.
But with civilized society they might have proved temperamentally in-
compatible; it was in their character to choose the frontier, and their
achievements there as likely as not greater than they might have been
in different circumstances.

Texas inevitably offers the most spectacular illustration of the
pattern of aliens and frontiersmen, and the changes now under way.
It had no native botanists until well into the 20th Century. Berlandier
came from Switzerland (born just across the border in France), Drum-
mond from Scotland, Lindheimer and Roemer from Germany, Reverchon
from France; Dr. Edwin James, Charles Wright, Buckley, and Riddell
were from the Northeastern United States, and Gideon Lincecum was
born in Georgia. After nearly a century and a half of botanical explora-
tion, the picture today is in part one of extreme contrast, in part one
of the continuation of old patterns. Texas today has more native-born
taxonomic botanists than any other state in the South, almost more
than all the rest combined, even if we count only those who have re-
cently published books about or described new species in its flora
(M. C. Johnston, Fred Jones, C. L. Lundell, Ellen Schulz Quillen, Chester
Rowell, B. C. Tharp, B. L. Turner, Barton Warnock, Eula Whitehouse).
In a remarkable illustration of “curious chances,” the alien element is
represented by three Canadian-born botanists (Walter Lewis, Lloyd
Shinners, Alfred Traverse”), one from each of the three major sections

26

of Canada (British Columbia, Prairie Provinces, Eastern Canada), work-
ing respectively at a state college, a private university, and a private
research corporation, and pursuing three different aspects of systematic
botany (cytotaxonomy, classical taxonomy, paleobotany). The out-of-
state Americans are represented by a North Carolinian (D. S. Correll)
and a Louisianan (R. A. Vines) in the Lincecum tradition, while a
Yankee from Indiana (N. C. Henderson) is the most recent addition to
the roster of Texas systematic botanists.

The pattern in the rest of the South is less extreme, but similar. There
are fewer native-born botanists, and almost no aliens, while immigrants
from the North are a large contingent, continuing the pattern that goes
back nearly a century and a half. In time we may expect the native sons
to become a majority,” but it is unlikely that there will ever be an era
of exclusively home-grown botanists. Even Britain, the world’s most
prolific producer of botanists, in her Golden Age, had such men as
the Germans Seemann and Stapf. Although Rafinesque complained” of
“not having been able to explore as yet the Southern States, deterred
by the bad roads, unhealthy climate, scanty fare, heavy expenses and
state of society. A pedestrian Botanist is not always very welcome there,”
the South has traditionally been hospitable to botanists, even enduring
the rabid Unionist and Yankee A. W. Chapman all through the Civil
War. It is a tradition likely to continue.

After this brief additional sampling of the historical record, we may
answer the question posed at the start. A Southern botanist is one who
lives in the South and devotes himself to botany. Origins may be inci-
dentally interesting, but are not involved in the definition.

ommentary on aliens and frontiersmen would be incomplete without
some mention of those whose families have been in America several
generations longer than the family of the present President of the United
States or the families of most of the nation’s citizens, but who scarcely
enjoy the usual status of early settlers. The blighting influence of the
slave economy on Southern whites has already been noted. For the
negroes it was of course much more extreme, and made worse by racist
attitudes on both sides. We have seen how, in terms of botanical history,
the release of the Upland White from long cultural suppression has
begun to have positive results. We may well expect that the longer
and more severe cultural suppression of negroes will be followed by
a longer period of recovery. But recovery is visibly under way, and like
so much else in the New South, at a quickening pace.

I vividly recall an incident that took place a year or two after I came
to Texas. I was showing a Northern visitor some native weeds in a
vacant lot in Dallas. While we were on hands and knees digging speci-
mens (as I recall, the plant was our attractive though small-flowered
native bindweed, Convolvulus hermannioides), a young colored lady
stopped to watch, and at length asked what we were doing. We ex-
plained as well as we could. “Oh,” she exclaimed, “I just love flowers

27

and wish I could know more about them!” Such had been my own
feelings from earliest childhood. But for her there was no chance; for
her children, perhaps.

Another incident, a year or two later, in St. Louis, on a Sunday after-
noon. I had gone out North Broadway Street to collect bees and wasps
in waste areas fringing the Mississippi River bottoms. Waiting at the
street-car stop to go back into town was a colored man of indeterminate
but considerable age. He could hardly wait for me to come up before
breaking into angry speech about a miserable patch of corn across the
street underneath a huge cottonwood tree. He was up from Mississippi
to visit relatives; back home they knew how corn ought to be grown,
and planting it under a cottonwood was all wrong. “It cain’t ’cwmulate,”
he insisted repeatedly, indignant that anybody should treat corn like
that, “it cain’t ’cumulate!” Illiterate he might have been; unintelligent
or incompetent he certainly was not. And again the natural love of
plants was plain to see.

Just as this paper was beginning to be written, there was showing
close to campus the motion-picture version of the prize-winning Broad-
way play, A Raisin in the Sun, written by and about negroes. Some
weeks earlier I had heard its author, Laraine Hansberry, in a radio
interview, telling with bitter intelligence some of the personal beliefs
that had been voiced in the play (“Why give God the credit for the
things man does for himself?’’). Memorable in the picture is the scene
in which the matriarch, defending a bedraggled house plant in their
Chicago flat from the cynical daughter, exclaims, “It expresses me!”

On my latest visit to Montgomery, Alabama, first capital of the Con-
federacy, a local newspaper had this to report (19 July 1962): “Two
years ago, Bullock County had around 2,200 white and five Negro voters.
It now has about 2,100 white and around 1,000 Negro voters, ”

The interest is there, the ability is there; the opportunity has been

anting, but is plainly on the way. The next Southeastern flora will
still be hardly past the pioneer stage. It may well be that the first
definitive Southern flora will be written a generation (or two, or three)
hence by a colored botanist, quite possibly one born in Mississippi.

REFERENCES
Sin a titles and dates for the various manuals and floras are given in the text, they
are not repeated here. Many of the titles in this list are not Peres cited in einer the
text or ne otes, they were among the most helpful to me in trying to understand
South ult and history. To those ae te) os ow th h, and to th h
think that they do, I strongly recommend t ery recent books dealing with the South

and the Civil War listed below. Today’s S eae in Piers al ae moved far beyond the
rather naive and limited viewpoint of the Twelve Southerners who in 1930 published Vil
Take My Stand. But he belongs to a small minority; the South in general still is
behind. Botanists, being themselves a small minority, should feel special kinship with the
historians, sociologists, and men of letters

GBY, PHILIP. 1958 feign and iSRore Longmans, Green & Co., London. (Repr.
1959, avec of a ia i Berkeley.)
BARNHART, J. H 23. Hardy Bryan “Croom (1797—1837). Journ. New York Bot.
Gard. 24: 7. (Esenaas in ae by John K. Small.)

28

BROOKS, VAN WYCK. 1936. The Flowering of New England 1815—186
Dutton & Co., New York. (Repr. 1946, The World Publishing Co., Cleveland oe a
York,

BROWN, eee GARROTT. 1902 (repr. 1930). The Lower South in American
pees Peter Smith, New York.
TTON, ee 1961. ane Coming Fury. (Centennial History of the Civil War vol.
1.) Dosbledi & Co., New Y
CLARK, THOMAS D. 1961. au Emerging South. Oxford University Press, New York.
DANIELS, JONATHAN. 1938. A Southerner Discovers the South. Macmillan Co., New
Yor

am A. HUNTER. 1959. Asa Gray 1810—1888. The Belknap Press of Harvard
oe ey Cambridge
AN, HOWARD R. 1958. The South in Northern Eyes 1831 to 1861. University of
Texas Press, Austin.
GEISER, SAMUEL WOOD. 1948. Naturalists of the Frontier (2nd ed.). Southern
ete ae se Dallas
AY, ASA Coa of Botany of the Southern States by John Darby.) Amer.
ae Sci. 70: Cae

185 x 18:57; se ny of the flora of the Northern United States. Amer.
Journ. Sci, 72: 204—232; 73: 62—84, 369-403

JONES, GEORGE NEVILLE, AND EDNA epee: aoe Principal institutional
herbaria be the United States. Amer. Midl. Nat. 40: 724—

KIMBALL, eee 1921. Reminiscences of Alvan GHEE Chapman. Journ. New
York Bot. Gard.

LYON, Pees 7 1939. A 50 edition best seller. Dartmouth Alumni Magazine 31: 18,
81—83. (I have seen only a reprint without page numbers; these obtained from Merrill,
1948.)

MERRILL, E. D. ore The Amos Eaton Herbarium. Rhodora
48. Unlisted new names in Alphonso oat botanical publuaions

Rhodora 50: ean
snes C. WRIGHT. 1956. The Power Elite. Oxford University Press, New York.
RISON, SAMUEL ELIOT, AND HENRY STEELE COMMAGER. 1950. The Growth
of ee American Republic (4th ed.), vol. 2. Oxford University Press, New York.
NICHOLLS, WILLIAM H. 1960. Southern Tradition and Regional Progress. University
of North Carolina Press, iow Hill.
ae CHO ge te F, 1961, The Stakes of Power 1845—1877. Hill and Wang, New York.
UM, ARD a. 1936. Southern Regions of the United States. University of
a Le ae De Chapel Hill.
PENNELL, FRANCIS W. 1941. Scrophulariaceae of Trans-Pecos Texas. Proc. Acad. Nat.
Sci. Phila. 92: 289—308.
PRATT, FLETCHER. 1948. Ordeal by Fire (2nd ed.). William Sloane Associates, New
or ia
eee Oa J. 1954. Americans Interpret Their Civil War. Princeton University
Pes chem
AFIN FSU, C. S. 1836. New Flora of North America. First Part. The Author,
Palade hia
RAVENEL, HENRY WILLIAM. (Edited by Arney Robinson Childs.) The Private
Journal of Henry William Ravenel 1859—1887, ee of South Carolina Press,
Columbia.

GERS, ANDREW ae 1944. John Merle Coulter, Missionary in Science.
Princeton University Press, Princet
RUBIN, LOUIS D., JR.. AND ROBERT D. JACOBS (editors). 1953. Southern Ren-
ascence. The rete of the Modern cane The Johns Hopkins Press, Baltimore.
SASS, HERBERT RAVENEL. 1935. Adventures in Green Places. G. P. Putnam’s Sons,
New York

29

NNERS, LLOYD H. 1962. haa oe United States Bonamia, including Breweria
A ‘Silane (Convolvulaceae). Castanea 27
MPSON, GEORGE LEE. 1956. “i ue of Carolina. University of North Carolina
ee eae Hill.
SMALL, JOHN K. 1923. Green deserts and dead gardens. Journ. New York Bot. Gard.
24: fey ee
Daa WILLIAM. 1899. Alvin (sic) Wentworth Chapman. Amer. Nat. 33: 643—

re SOUTHERNERS. 1930. I'll Take My Stand. Harper & Brothers, New York
and London

UNITED. STATES DEPARTMENT OF COMMERCE. BUREAU OF THE CENSUS.
918. Negro Population 1790—1815. Government Printing Office, Washington.

VANCE, RUPERT B., AND NADIA DANILEVSKY. 1945. All These People. The
Nation’s Human Resources in the South. University of North Carolina Pie Chapel Hill.

WARREN, ROBERT PENN. 1961. The Legacy of the Civil War. Random House, New
ork

WHERRY, ascent T. 1957. Reminiscences of John K. Small. Castanea 22: 126—129.
WHYTE, H. he Organization Man. Simon & Schuster, New York.
(Repr. 1957, eens Ree See
WARD, C. VANN. 1960. The Burden of Southern History. Louisiana State Uni-
versity Press, Baton Rouge

NOTES
1“The Mason-Dixon Line is, i way, biological as well as political. Its vicinity is the
dividing line between early neatone Seen eee ae nd later seasons northward, in the first half
of the year and vice-versa in eee econd half.” (Small, 1923, p. 194.)
2 For numerous indications of this, see Dupree’s biography

gers’ biograph Digind to ke a shining hero out of Coulter, but fortunately
ug

5
for work he never did, and which went into disrepute with Scribner. Rose was his pupil,
mnie ; : ale

a

of sitting once in a while with Rose while Rose explained the work he had done on the
ra. Coulter’s chief part in the deliberations was smoking rank cigars and stinking up
the room with tobacco smoke. It was the same method which produced Coulter’s Flora of
Te ut to date no one has ever claimed being the clerk who got it out. Coulter’s
Botan os VAG sie ae U.S. Nat. Herb. vol. a 1891—1894) is such a poor
that ned to believe it was ane | his alone, and tee on what

ece

ie Pie a his ae as ae he did not have more talented associates to exploit. On

the basis of the ee it eae that his famous last words CI should like to a a serv-
ice’) omitted two ones: “to Coulter.’

See his ee ai eee and geniali ae Gray could at times be ruthless and ev
vindictive. His treatment of S. B. Buckle oe is clear case in point. oe. Scan eet
that with the many collections from T n his possession cee of Berlandier, Drum-
mond, Leavenworth, Lindheimer, Wright, pee Pree ), he had al complete representa-
tio e state’s flora was eas when Buckley published numerous new species
without consulting hi supplying a good set of specimens, so much so that
i pa in the haarrens of ce Philadelphia Academy he penis Spee ay
h species as wo s, and a the Academy no an

is new sp

Buckley’s work. The Civil War supervened, and it was some years before Sones was Sane
to defend himself. He pointed out that the main shipment of his specimens had been lost
in transit, so that he had to get along with mere scraps that he had kept with him for

30

tudy; this was the reason for the poor quality of his fies, biloase Gray had on

ee noted, and for his failure to distribute duplicate We add today that Gra
was mistaken about the Texas flora; a large proportion i 3 "s new species were ee
fectly good, and are recognized today. One ot h seas if by also was

un-
fairly condemned, but leer arate we do not have even poor fragments = his specimens.

° See the reminiscences of Chapman by Kimball, and of "Small by Wher

™See the introdu uctory remarks in Pennell’ s “Scrophulariaceae of Vesna reas Texas.”

In ick i aa I find oe eight living American botanists who have authored

a je rs nual of a or region: oe vis (Idaho), Harrington (Colorado),
Munz Sacmae ra nee (Neth Dakota), Strausbaugh and Core as hi ginia), Glea-
son (The New Britton & Brown), Cronquist (its companion Manual). of these men
are now an (The concluding volume ee the West Virginia flora, and ae s Manual,
are reported in press as J write bas Rees 1962.

® Personal communication from Dr. Robert E. Woodson, Jr.

19 See also my acid remarks on ce generic flora in Gane aan with Drosera, this journal
pp. 33-59,

™ Julia Hale’s The Peterkin Papers is the classical work depicting the mania for learning.
See feces in Van Wyck Brook’s ed Flowering - New ae

™ Biltmore Botanical Studies ran o ju ust two numbers, published in 1901 and 190
detailed eae of the very a institution which ae ae is greatly to be ats
almost no information re it is available
3In the first 21 volum ns Cantribation from a U.S. edie meals two es rt
papers oe E. 8. Steele ie vo the only that chiefly

fs) There q
devoted to ts of the South ee ing x Texas), plus three whole Noa (Botany of
West a a Life of Alabama, Flora of the District of Columbia). addition, a
number of monographs (that of Panicum, for example) are of groups most prevalent in
the South. Later volumes of the Contributions have come to deal almost a with

s made for a society gay and polished, even brilliant . The ee ar is
that this society was so sterile Sealeotaliys (Flet etcher Beste Ordeal by Fir
“The South, like most aristocracies, was at in education, oe of ae a re y
and - the individual member.” (Thid., 280.) “They were kind, ee ae one =
serena But it also noted exe their roe were trivial, mipsel a a mport
than the concoction of mint juleps. “They visit eee other, eat, drin re merry, se
that is all. They have excellent qualities,, but no occasion ealls them ae >” (Review
J Pendleton Kennedy’s Swallow Barn, or ee ojourn in the Old Dominion, summarized

and quoted in Howard Floan’s The South in ne Eyes, p. 91
«Thus when the frontier period passed . . . the Old ee hod had as ee a
considerable number of gifted and vigorous workers. The South, sat had become pre-
Seu movement. rom this time ational movements
in the South dwindled.” (S. — Niswoies of the Frontier, ee 13, Notes on
Scientists of the First a p. 26 An interesting and rather vag illustration of
the tenacious survival of the Old South S mental rigidit ty is to be fou erbe rt Ravenel

y ut t
name of William eels one that it is “a ca Huguenot name and Saat to i
syne ors se ” but nothing is said of Dr. Francis Peyre Porcher who, to help th

Con d Resources of the Southern Fields and Forests, Medical, ee
and eee Suliche d at Cha rleston in ae eas

ife. At th
inserted a defense of Calhoun! ‘tht a man Al pretensions a cholarliness should, in a
book of local natural history, omit mention of the most illustrious naturalists in the
alc an come to ied defen ae ee frivolities and Calhounism, is striking evidence of how

tad
—

the of the Old South could restrict ion gee
on new lee of the Sout o be ef gisunas less cultured, but
cautious and tight-fisted. On the ee | hand, ae me did n are the ante-bellu

Democrats’ opposition to “Federal appropriations. . . . They Slee press steadily for the

31

segregation of the Negro and his elimination from politics, and they were not sais
intereste oy in providing improved educational facilities.” (Nichols, "The Stakes of Pow
pp. 212

anf ee is an even more signific ant figur re in American botanical history than Asa Gr ray,

cussion (se mple “Darwin and Botany,” a ee York Acad. Sci. 19: 28—33,
1909) ee a eens commonplace mind. He a power politi ician first and fore-
most, who by some strange chance seized on coe as an ideal field in which to exercise
his talents

apoE economic conditions are important, and I refer to them repeatedly, they are
not everything. I quite agree with the views of Fletcher Pratt Se ei introduction to his
Ordeal a Fire); other ahs may outwei igh economic considera We have

a
ranked lower) it accepted the political and inevitably the cultu goraarien of the
Plantation Aristocracy, with its blighting influence on the inte ies zi a . In explaining
West Virginia’s achievements ere handicaps, we must certainly ihclude the role of
active individuals as a major fac

t is pes to note ae of the tiny handful of living historians of American
botany (Dupree, Ewan, Geiser, McKelvey, McVaugh, and—on the strength of this paper—
Shinners), one e half now reside in se South (Ewan, Geiser, Shinners), and tw

Georgia). When add the eee awareness of its Ea to Western
with Baek inevi ae we find Texas again has had major attraction. It can hardly be
pure chance. However su aa the eee of ee climate may function, we have
visible evidence that they
ewis is cu ne on extended leave in Europe and Africa. Dr. Traverse has ve

recently qtiearae 1962) left the Shell nee Company to train for the Epi ea

try.

xpo rtation of Southern brain- ee has been one reason for the low number of
Southern botanists who are Sout ern born. The present Head Curator of the New York
Botanical ne is a native of Geo

22 See his New Flora of North a verica (First Part), p. Had Rafinesque come on

horseback, he might have been somewhat more kindly ee ie the South, a gentleman
did not ee on foot.

aa

ANNUAL SISYRINCHIUMS (IRIDACEAE)
IN THE UNITED STATES

LLOYD H. SHINNERS

Only three annual species of Sisyrinchium occur in the United States.
It is thoroughly characteristic of the elusiveness of taxonomic characters
in the genus that one of them (S. rosulatum) is sometimes perennial.
The major area for all three is in eastern Texas and Louisiana. There
one of them (S. minus) is native, but it has been introduced into Cali-
fornia and North Carolina, and abroad into Argentina and Uruguay.
The other two are natives of temperate South America which were
introduced into the Southern United States (and elsewhere around the
world) beginning in the middle or latter part of the 19th Century, but
did not become well established and common until relatively recent
times. These two hybridize readily in their new home, as apparently is
true also in the areas where they occur together as natives in South

erica. Nevertheless they appear to be maintaining their identities
as separate species. There has pice no indication of crossing between
the two aliens and the native S. minus, a fact now made readily under-
standable by Oliver and Lewis’s eke (1962) of the haploid chromo-
some numbers: 16 in each of the two aliens, 5 in S. minus.

The species and hybrids are all easily recognized from the colors of
the fresh flowers, but such information is grievously lacking with most
herbarium specimens. This account is based primarily on my own ex-
tensive field observations and collections made from Texas to northern
Florida, in large part (1956-1961) under a grant from the National
Science Foundation for preliminary field work toward a flora of the
Gulf Southwest. An extended tour of the Northeastern States during
the winter of 1945-1946, and shorter trips at intervals since, have
=e me to examine types and other specimens at the Chicago Natural

Texas, Texas A. . College, Tulane WAiveecias the United States

National ciate eae and the United States National Arboretum Her-

barium, as well as Southern Methodist University. I am indebted to the
if

made their facilities available during my visits. Most recently I have
to thank Dr. Robert L. Wilbur for the loan of collections from the Duke
University Herbarium; Dr. George B. Van Schaack, of the Missouri
Botanical Garden, for a copy of the original ae of S. va
vianum; and Prof. J. Leandri, of the Paris Museum, for notes ce a
photograph of the type of S. micranthum.

Despite the sometimes perennial habit of S. rosulatum the three an-

SIDA 1 (1): 32—42. 1962.

33

nual species are generally easily recognized as such, and it takes but
little experience to be able to distinguish them on sight from the more
numerous perennial ones. Apart from intangible features of general ap-
pearance, their gamut of flower colors is almost completely different
from that of the perennials, excepting a few Far-Western ones. Only the
uncommon albino form of S. minus (white with yellow eye) duplicates
what may appear in the perennials. The latter (all those in the So

and East, a majority of those in the West) have medium to large
perianth ranging from white to light blue, deep violet-blue, or even
purplish blue (but still decidedly on the blue side), with yellow eye.
The annuals never have a distinctly blue perianth (though often bluish
or greenish in withering), the colors ranging from pinkish lavender to
rosy purple (often partly or largely white with eye-ring and stripes)
to yellow, and in the hybrids to various shades of brownish purple or
purple-red (see key below and notes on hybrids at end). Taxonomically
these form an artificial group, but it is convenient to treat them together.

KEY TO THE ANNUAL SPECIES

la. Ovary and capsule oblong-ellipsoid or oblong-pyriform, more than
1% times as long as broad; stamens well exserted, about half the
length of the perianth; perianth variously lavender-pink to purple-
rose, white with yellow eye, or all yellow............. . S. minus

. Ovary and capsule globose or subglobose, shorter to barely longer
than broad; stamens barely or not exserted, 1/6—1/3 as long as the
perianth; perianth variously colored (see next couplet), but never
just as in the preceding (following two species hybridize freely;
see remarks at end of text)

. Perianth yellow with brown-red eye ring and often a single brown-
red center line on each lobe; length (half-width) of perianth 5—10
mm.; diameter of capsule 2.7—3.5 mm................... 2. S. exile

. Perianth white to lavender-pink or bluish-purple-tinged, with yel-
low eye circled by rose-purple eye ring and commonly three (but

sometimes one, or none, or more) lines or stripes down each lobe;
length aa width) of perianth 9—16 mm.; diameter of capsule
3 vial @ BY 0 ed ce ete PO oOo. ie cee 3. S. rosulatum

a
ion

bo
ish)

nN
om

1. S. MINUS Engelmann & Gray, Pl. Lindh. p. 55 (Boston Journ. Nat.
Hist. 5: 263). 1845. “Margin of pools, &c. in the prairie west of San
Felipe,” Austin Co., Texas, Lindheimer Fl. Tex. Exs. 313, April, 1844
(holotype ae isotypes MO, SMU). — S. flexuosum Rafinesque, Aut.
Bot. p. 65. 0. “Arkanzas and Texas.” Not S. flexuosum (L.) Sprengel,
Syst. 1: a aoe — S. Bermudiana var. minus (Englemann & Gray)
Klatt, Linnaea 31:69. 1861. — S. Thurowii Coulter & Fisher, Bot. Gaz.

: ‘ um
Canby 238, 25 March 1900 (NY). To my eye the perianth on the type is

34

rosy lavender, not “bright purplish blue” as described by Bicknell. —
The name S. geniculatum Herbert is given as a synonym of this by
Baker and Johnston, but I do not believe this is correct (see remarks in
list of doubtful or excluded names at end).

The usual perianth color in this species is a distinctive purple-rose
(in my notes I find I most often called it magenta-rose) which is quite
uniform, except for being occasionally lighter or darker than usual;
rarely it is pale enough to be called lavender-pink. There is not a con-
tinuous series of intermediates between this and the other two color
forms, which are even more uniform. White is occasional and wide-
spread, usually few individuals amony many of the typical form, rarely
in reverse proportions, or by itself. The yellow form I have found only
once (at Needville, Fort Bend Co., Texas), growing with and less com-
mon than the typical form. The type of S. Thurowii is the only other
record of the yellow known to me.

These color forms are highly significant in relation to the evolution

Such mutation elieve is occurring now and has long occurred, and
is sufficient by itself to explain the origin of many of our existing
species and varieties. It would be of great interest to know the precise
details of the inheritance of color forms, and in particular why there
are no intermediates. As for survival value, to try to read anything of the
sort into these variations is to indulge in wild fantasy. At least two of
the forms are successful in invading new territory. The third is so rare
that it has so far given no indication of spreading, but this proves
nothing about its future.

Because it was first found in Texas in natural habitats, and for long
was known only from there and adjacent Oklahoma (Rafinesque’s

indubitably native. From its habitats, and the recency of the records, I
believe that in Louisiana it is largely introduced. It is unquestionably an
introduction in California and North Carolina, and I have no doubt that
the same is true of its occurrence in Argentina and Uruguay (reported
by Johnston, 1938).

Rafinesque reported this species from both Texas and “Arkanzas.”
Like Nuttall’s ‘“Arkansa,” the latter term referred to the old Arkansas
Territory, which included eastern Oklahoma. Although Waterfall does
not include S. minus in his catalogue of the Oklahoma flora (1952), it
is not at all unlikely that the species once occurred there; it has long
been common as far north as Dallas, Texas. All the United States
collections I have seen from outside Texas are cited below. For the
latter state I have merely listed the counties from which it is known.
Since the publication of my 1948 map, it has been found as a roadside
weed in Montgomery and Tyler counties, and introduced with St.

35

Augustine grass sod in Nacogdoches County, all east of the area shown
on the ma

CALIFORNIA. Los Angeles Co.: in grassy field dominated by Phalaris
Lemmoni; Sepulveda Blvd., northwest of Los Angeles airport, Frank W.
Gould 2287, 15 April 1944 (SMU). “Flowers white.” (Distributed as
S. bellum var.) LOUISIANA. Grant Parish: 4.6 miles southeast of
Colfax (from road junction on U.S. Highway 71), road shoulder,
Shinners 29,510, 18 April 1962 (SMU). “Perianth magenta-rose (white
on one plant).” Madison Parish: 2.8 miles west of Waverly, foot of road
fill by creek, Shinners 28,221, 18 April 1960 (SMU). “Perianth white
with yellow eye.” Same locality and date, Shinners 28,240 (SMU). “One
plant with magenta-rose perianth (all others seen white). Natchitoches
Parish: Natchitoches, swampy open ground, E. J. Palmer 7486, 3 May
1915 (MO). Red River Parish: 1.4 miles west of Grand Bayou, road fill,
Shinners 27,227, 22 April 1958 (SMU). “Perianth purple-rose; lobes
several-striped on back.” St. Martin Parish: 1.8 miles south of Parks,
dried-up ditch, Shinners 28,128, 16 April 1960 (SMU). “One plant only.”
Tensas Parish: 3 miles north of Helens, margin of swamp forest, hard-
woods, J. Ewan 19054, 20 April 1957 (NO). “Flowers very pale pink,
mostly past.” NORTH CAROLINA. Durham Co.: Duke campus, waste
places, W. B. Davis 819, 17 May 1932 (DUKE). TEXAS. Aransas, Atas-
cosa, Austin, Bastrop, Bee, Bell, Brazoria, Brazos, Dallas, Dimmit, Falls,
Fort Bend, Harris, Karnes, Kleberg, Liberty, Llano, Matagorda, Mont-

omery, Nacogdoches, Navarro, Nueces, Robertson, San Patricio, Travis
Trinity, Tyler, Williamson; also “Seguin—Lavernia (Guadalupe or Wil-
son Co.), “Victoria—Goliad”’ (counties with same names

2. S. EXILE Bicknell, Bull. Torr, Bot. Club 28: 573574, 1901. “Sandy
sea shores at Galveston,” Galveston Co., Texas, J. E. Bodin, 25 February
1890, “herb. Univ. of Minn. and U.S. Nat, herb.” (latter specimen ex-
amined). — S. Brownii (sphalm. Brownei) Small, ex Small & Alexander,
Bot Siouseore Iridaceous Pl. Gulf Ae (Contrib. New York Bot. Gard.
3917): 330. 1931. (“Excerpt from the forthcoming Manual of the Flora of
the Southeastern United States.”) Not designated in the list of new
binomials (unnumbered page at end), and no type indicated; noted
only as “S E La.” It was in fact named in honor of Prof. Clair A. Brown
of Louisiana State University. For unknown reasons I find no notes
on the type from my New York visit in 1946, but the description leaves
no doubt as to the identity of the plant. I did examine the following
later collection at New York. LOUISIANA, Livingston Parish: roadside,
pine land near Hammond, C. A. Brown 3846, 9 April 1932. “Flowers
yellow, purple brown line on inside of petals.” — This is S. micranthum
of many authors (see doubtful and excluded names at end).

In flower color this is rather uniform, the chief variation being the
extent to which the brown-red eye ring extends as a thin line down
each perianth segment. Predominantly the perianth is medium yellow,
very rarely pale or sulfur yellow. The species is now a common and

36

often abundant weed of sandy road shoulders and damp sandy ground
along highways in southeastern Texas and Louisiana, and has spread
north into Arkansas. East of the Mississippi River it is still largely
restricted to areas near the Gulf, chiefly in northern Florida, but it
occurs as a lawn weed as far aR as Statesboro, Bulloch Co.,
Georgia (Gordon P. DeWolf, in letter). I was surprised in tabulating
the records to find none for Alabama. I feel certain that it occurs there,
but I failed to collect it in several trips to the three southernmost
counties made with a eel particularly in mind. State and county
(parish) records are as follow

ARKANSAS. Bradley, ae (Also Drew, according to Moore, 1958.)
FLORIDA. Clay, Jackson, Polk, St. Johns. (Also Washington, on basis
of hybrids; see detailed notes on these at end.) LOUISIANA. Acadia,
Allen, Beauregard, Bienville, Calcasieu, Jackson, Lafayette, La Salle,
Livingston, Natchitoches, Rapides, Sabine, St. Helena, St. Tammany,
Vermilion, Vernon, West Feliciana, Winn. (Also Evangeline, Jefferson
Davis, on basis of hybrids.) MISSISSIPPI. a River. TEXAS. Angelina,
Austin, Chambers, Galveston, Hardin, Harris, Jasper, Jefferson, Liberty,
Montgomery, Newton, Panola, Polk, Rains, Robertson, Sabine, Shelby,
Trinity.

Apparently the first collection of this cae from the United States
was made in Texas by Elihu Hal, probably in 2. There is a specimen
at the Gray Herbarium with no data except oe yellow,” the col-
lector’s name, and that of the state. Hall collected at various central
Texas localities from the coast inland to Austin and Dallas. He may
well have found the plant at Galveston, where Bodin later collected
the type of S. exile. One possible means of introduction for both this
species and S. rosulatwm is suggested by an incident reported in Wini-
red Kimball’s reminiscences of Chapman, occurring some time after
1887. “When a South American ship brought up clay from the ‘Rio de
la Plata’ as ballast, and my father had it spread over the garden, Doctor
Chapman’s interest grew apace. He watched over each new ‘weed’
that cropped up.” There are specimens in the Gray Herbarium from
Easter Island (collected in 1904), Fiji (Viti Levu, 1927), Hawaii (Hawaii
National Park, 1943) and Australia (Queensland, 1943). The original
home of this now very widespread weed seems to have been in the

1820) are worth quoting. “We find no account of this plant but what
has been derived from the description and figure above quoted, which
were taken from a solitary dried specimen in Jussieu’s herbarium, col-
lected in Peru. Communicated in July last by Mr. Anderson, of the
Botanic Garden at Chelsea; to whom it was sent by Mr. Otto, from
the Royal Botanical Garden at Berlin.”

37

3. S. ROSULATUM Bicknell, Bull. Torr. Bot. Club 26: 228—229. 1899.
“Dry open places in sandy soil, coast of South Carolina and Alabama.
South jee Sullivan’s Island (Charleston Co.), May 8, 1852, Pro-
fessor Lewis B. Gibbes. Alabama: Mobile, April 6, 1896, May 5, 1896.
Dr. Charles Mohr.” (Mohr specimens examined, US.) Bicknell states
“flowers not seen, reported to me by Dr. Mohr as being of a reddish

names at end). een of my collection down to 1962 were all dis-
tributed under this name.

Apart from the evident hybrids discussed below, there is great varia-
tion in flower color in this species, especially in pattern, which I be-
lieve indicates spontaneous genetic diversity rather than introgression.
The perianth is rather large and showy, commonly white with varying
amounts of rose-purple in the form of an eye ring and lines down the
segments, but occasionally colored throughout. It is also more variable

”

embarrassing combinations of character.
Florida, Louisiana, and Texas. S. rosulatum in the United States oc-
cupies much the same area as S. exile. In Texas it is much less common,
but in Alabama and Mississippi it is more so, than the yellow-flowered
species.
ALABAMA. Baldwin, Mobile, Washington. ARKANSAS. Union. (Also
E

nkin. R
OUTH CAROLINA. Charleston. (Syntype of the species; no recent
A j ogd

Newton, Tyler. (Also Polk, Trinity, on basis of hybrids

The oldest collections from the United States were those from South
Carolina (1852) and Alabama (1896) on which Bicknell based the
species. It was found in “open fields, Richland,’ presumably St. Mary
Parish, Louisiana, by R. S. Cocks in June, 1908 (NO). But most of its
North American range has been attained more recently. When I began
field work in the Gulf States in 1945, it was rare in southeastern Texas
(I found it only in Jefferson County); now it is frequent there. In

so; certainly it is much more conspicuous. Its original home was nearly
the same as that of S. exile, from southern Brazil to Argentina. Whether
Johnston’s report of S. laxwm from Europe refers to this species or the
true S. laxum I do not know.

38

2 X 3. S. EXILE X ROSULATUM. The binomial S. Metae Herter
probably was based on a nothomorph of this cross (see under doubt-
ful or excluded names at end). In Louisiana intermediates between the
species are rather common (elsewhere they are much less so), nearly
always in association with the parents, the majority appearing to be
first-generation hybrids, while apparent back-crosses or second-genera-
tion segregates are rather uncommon. Below ae cited 15 collections
representing such intermediates, with notes on perianth color. All are
deposited at SMU, and all but the last one were collected by myself.

ORIDA. Washington Co.: 1.8 miles east of Shipley, 27,009. ‘““White
with chocolate-red central ring; lobes with slender dorsal central line.”

central stripe.’ Same locality, 23,068. “Medium large, light yellow.
Growing with S. micranthum (i.e. exile), 2 color forms of S. laxum
(i.e. rosulatum), unidentified intermediate type, 1 plant with large,
pale yellow, lined perianth.” Same locality, 23,069. “Large, pale yellow
with dark lines,” growing with preceding. Beauregard Parish: 0.5 mile
north of Ragley, 23,665. “Garnet, tepals with darker base and central
line.” — 3.7 miles south of Longville, 23,543. “With yellow eye, tepals
scarlet-mauve at base and center.” Calcasieu Parish: 4 miles south of
Gillis, 23,153. “Smaller than in S. laxum (i.e. rosulatum), mauve with
scarlet tinge, lobes with single dark central stripe.” Evangeline Parish:
8.5 miles east-southeast of Oakdale, 27,970. “Withered (2 P.M.), tube
yellow.” Jefferson Davis Parish: 5.1 miles southeast of Jennings, 23,140.
“Light scarlet-mauve, tepals with dark central line.” Rapides Parish:
2.5 miles northeast of Glenmora, 23,255. “Medium large, mauve with
scarlet tinge.” Vernon Parish: 2.3 miles northwest of Leesville, 22,753.
“Smaller than in S. laxum (ie. rosulatum), white with mauve ring
around yellow eye.” — 2.2 miles northwest of Anacoco, 23,673. “Brown-
ish mauve, tepals darker at base and down center.” Winn Parish: 7.5
miles north of Winnfield, 23,342. “Scarlet-mauve.” TEXAS. Polk Co.:
2 miles east of Livingston, 23,488. ‘“Perianth 1/3 larger than in associated
S. micranthum (i.e. exile), tepals brownish mauve, darker down center
and at base.” Trinity Co.: 1 mile west of Neches River, R. L. Oliver 312.
“Light yellow with yellow center; outside base mauve-brown and
along veins’; perianth as large as in typical S. rosulatwm.

During April and May of 1956, when a majority of the above collec-
tions were made, the presumed first-generation hybrids (with perianth
of intermediate size, of distinctive scarlet-mauve or brownish-mauve
color unlike any forms of the seer with a single central line on each
segment) were very common and remarkably uniform. In subsequent
years they have been much less aa There is no indication that
the two species are becoming completely mongrelized. On the contrary,
they appear to be retaining their separate identities to an astonishing
degree. Whatever mechanism or mechanisms served to maintain the

39

two in their native area evidently continues to operate in their new
home. I have seen no evidence of crossing between an annual and a
native perennial species, though there is evident hybridization among
several of the latter (see comments in my Spring Flora of the Dallas-
Fort Worth Area, Texas, 1958).

I have made no attempt to grow these plants or try artificial crosses.
Much intensive genetical and doubtless chemotaxonomic work could be
done on them, with a large staff of assistants, numerous graduate stu-
dents (possibilities of several Ph.D. theses at least), and a succession of
research grants, each larger than the one I received for work on the
entire flora of the Gulf Southwest. Having committed myself to general
flora work in an enormous area where it is desperately needed, I have
no time for such things. To anyone trying to view the development of
American botany in some reasonable scientific and historical perspec-
tive, it is all food for some very melancholy thoughts.

DOUBTFUL OR EXCLUDED NAMES

All the botanists who have done revisionary work including the three
annuals (Klatt, Baker, Bicknell, Johnston, Foster) knew the plants
only from herbarium specimens. But Sisyrinchium simply is not a genus
that can be worked out solely in the herbarium. I have no acquaintance
with the South American species other than the two introduced ones,
but feel sufficiently well acquainted with the two to know the limits of
their variation, and to reject all of the names referred to them in
Johnston’s really very creditable revisio

S. geniculatum Herbert, Edward’s a Reg. 1843 Misc. p. 84. Placed
under the heading “Columnea staminea cylindrica,’ the entire descrip-
tion is as follows. ‘5. Geniculatum, mihi; ex prov. Texas dicto, parva
caule geniculato perianthio limbo laete coeruleo.” This is listed by
Johnston as “nomen” only, in the synonymy of S. minus. Earlier Baker
had indicated like identity by citing it under S. Bermudiana L. “Var.
3. S. GENICULATUM Herb.,” with S. minus as synonym. There is
enough description in the Suen publication so that it must be re-
garded as validly published, even though what was said is, for this
genus, all but useless for eee If indeed identical with S.
minus, its name would have to be adopted for that species, being two
years older, unless its still older appearance as nomen nudum in as-
sociation with a Brazilian plant can be taken as grounds for rejecting it
as a nomen confusum. The description of the perianth as “limbo laete
coeruleo” certainly does not fit any of the known color forms of §S.
minus, but would apply very well to S. pruinosum and other native
perennials of Texas. It is very probable that Herbert had a Drummond
collection, and Drummond unhappily worked in a veritable hot-bed
of complex forms, introgressive hybrids, and what not involving the
abundant native perennials of south-central Texas. Whether S. genicu-
latum could be satisfactorily identified even with a type specimen at

40

hand is doubtful at best. Things are complicated by the fact that the
name did appear in print a year before the species was described, and
it then associated with a Brazilian species which according to
Johnston was S. laxum (i.e. S. rosulatum). This first appearance was in
“Contributions towards a flora of Brazil,’ by Gardner, in London Journ.
Bot. 1: 538, 1842. The account there is as follows: “217. Sisyrinchium
geniculatum. Herbert Mss. in Herb. Hook. HAB. In moist sandy places
at Tejuca. Fl. Nov.” The use of the plural “Mss.” implies that Herbert
annotated several specimens with this name, but as it was published
the following year for a Texas plant, Brazilian material was by implica-
tion excluded by Herbert himself.

S. laxum Otto ex Sims, Bot. Mag. 49: 2312. 1882. “This new species
of SISYRINCHIUM was introduced into the Chelsea garden in 1820, by
Mr. Otto, curator of the Berlin Botanical Garden, under the name which
we have adopte . Communicated by Mr. Anderson of the Chelsea
garden, who has ereated it as an alpine, and it has survived the last
winter exposed to the open air. Native country unknown.” The plate,
which must stand as the type for this species, shows the top of a plant
only. The flowers appear relatively small in proportion to the very

rge capsules; the perianth is whitish with dark purple lines (the
description says only “white-streaked”), quite similar to S. rosulatum
except that the lines are uniformly very thin all the way down to the
very narrow eye ring (in S. rosulatum they widen toward base, and the
eye ring is usually very prominent). Light was shed on the identity of
the plant figured by Sims when I received an unidentified specimen
from New Zealand, though I did not at first realize it. The specimen
is M. B. Ashwin 530, from Lower Hutt, Wellington, North Id., 3 Nov.
1958 (SMU). “Forming small tufts in dry ground by ond de Intro-

dried perianth shows the uniformly thin lines exactly as in Sims’s
plate. In other respects, especially the very large capsules, the speci-
men is an almost perfect match for the plate, which evidently repre-
sented the plant exactly in life size. The larger capsules on the specimen
are 6 mm. in diameter (the larger one in the plate is 8 mm.), much
too large for S. rosulatum. The perianth in the New Zealand specimen
is 15 mm. long as pressed, the whole plants (two on the sheet) coarser

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laxum
still unknown, though presumably it is South American.

S. Metae Herter, Revista Sudamericana de Botanica 5: 28. 1987.
Johnston places this in the synonymy of S. laxum (1938, p. 391), but
Herter describes it as having white flowers, the tepals 3—5 mm. long
(too small for either S. larum or S. rosulatum), and lacking the purple
coloring of those species. I strongly suspect that it is a hybrid form,
quite probably involving S. exile, but since still other species not

41

familiar to me occur in Uruguay and might hybridize with S. exile or
S. rosulatum, I cannot decide this point. Anyone wishing to adopt a
bionomial for the hybrids would have to settle the identity of S. Metae
and other names placed in synonymy by Johnston.

S. micranthum Cavanilles, 6ta Dissertatio Botanica p. 345; pl. 191, fig.
2. 1788. “Habitat in Peru: examinatum ibi a D. Josepho de Jussieu. V. S.
unicum exemplar apud eius nepotem.” Neither description nor figure
is conclusive. Flower color unfortunately is not mentioned. A photo-
graph of the type reveals that the drawing was crudely made, but the
specimen itself is immature, and no great help either. It is erect and
densely leafy. Prof. Leandri very kindly compared with it some small
plants of S. exile which I had collected in St. Helena Parish, Louisiana,
and reports that they seem to belong to the same species, but that on
the Jussieu specimen “les racines sont toutefois un peu plus fortes.”
This is exactly a distinction just noted between S. laxum and S. rosu-
latum. Chiefly on the basis of general appearance, nature of the roots,
and geographic location, I believe that the true S. micranthum is not
the now cosmopolitan annual which has so long passed under that name,
but a closely related species, perhaps perennial, of highland regions in
western and northern South America. I believe Johnston was correct
in making S. iridifolium H.B.K. (from Venezuela) a synonym of S.
micranthum; probably S. scabrum Schlechtendal & Chamisso is also
the same. These plants will have to be studied in the field by someone
who also knows S. exile in a living state.

S. Pearcei Philippi, Linnaea 33: 251. 1864—1865. Listed by Johnston
with query as synonym of S. laxum. The original description states that
the perianth is yellow, so that it cannot be either S. laxum or S. rosu-
latum; it may well belong to the true S. micranthum.

S. uniflorum Gay ex Philippi, Linnaea 29: 63. 1857. Listed by Johnston
as synonym of S. lawum. In the original description the word “caerulei’”
(sky-blue) is applied to the perianth, and the plant is compared with
the blue-flowered S. chilense, differing in having scabrous stem. Neither
color nor stem indument apply to S. laxum or S. rosulatum.

valdivianum Philippi, Anal. Univ. Chile 91: 616—617. 1895. Despite
f t

capsule is described as 4—5 mm. in diameter, and on this basis the name
canont apply to S. rosulatum.

REFERENCES
KNELL, EUGENE. 1899. ee in ee Sixteen new species from the
ee States. Bull. Torr. Bot. Club 2 17—231.
. 1901. Studies in CR ne The species of Texas and the South-

west. Bull. Torr. Bee Club 28: 570—592.,

Rete ROBERT G. 1948. Studies in the flora of Bolivia,—II. Contrib. Gray Herb.
166; 28 . (Devoted entirely to a synopsis of Sisyrinchium.)

JOHN Coy IVAN M. 1938. ae seis of Sisyrinchium in Uruguay, Paraguay and
Brazil. Journ. Arnold Arb. 19: 376—

42
KIMBALL, WINIFRED. 1921. ear of Alvan Wentworth Chapman. Jou
New York Bot. Gard. 22: 1—12. (See for weed introduction via dirt ballast Ae
Argentina or Uruguay.)

MOOR ew ease for the Arkansas flora. IV. Proc. Ark. Acad. Sci.

9—16, Soman, Pe 10—11, .
EWIS. oe ee numbers of
> 43—

OLIVER, CE AND
pa eee in eastern can Aeaerica, Sida
NERS, an ae Pe Geographic limits of some ae weeds in Texas. Texas
isyrinchium, pp.

Eats Mag. en (Si
958. ae ne of the Dallas-Fort Worth Area, Texas. (Sisyrinchium,

pp. 100—102.)

SMALL, JOHN K., AND EDWARD J. ALEXANDER.
of the itidaceous plants of the Gulf States. Contrib. New York Bot. Ga
g Manual of the Flora of the Southeastern United isa *) (Sisyrin-

1931. sor ia interpretation
rd, 327. (“Excerpt

from the

chium, pp. e

ea ae U. T. 1952 A Catalogue of the Flora of Oklahoma. (Sisyrinchinm, pp.
£96)

CHROMOSOME NUMBERS OF SISYRINCHIUM
(IRIDACEAE) IN EASTERN NORTH AMERICA’

ROYCE L. OLIVER AND WALTER H. LEWIS
Stephen F. Austin State College, Nacogdoches, Texas

Chromosome numbers for 18 species of Sisyrinchiwm with basic com-

plements of x = 8, 9, and 17, have been reported (Vilmorin and Simonet,
927; Maude, 1940; Bowden, 1945; Covas and Schnack, 1946; Sermonti,

1948; Skottsberg, 1953; Lewis and Oliver, 1961). The 8 and 9 series con-
tain both diploid and high polyploid species, but only diploid species are
found in the secondary x = 17 series.

MATERIALS AND METHODS.—Immature flower buds were collect-
ed in the field and were fixed in modified Carnoy’s solution (4: 3: 1).
As soon as possible after fixation the buds were stored at refrigerator
temperatures for periods up to 8 months; only after 7 months was deteri-
oration noted in some cells. Buds were squashed in 1% acetic-orcein and

S or more occasionally somatic cells were studied. Representative
chromosomes were drawn with the aid of a camera lucida at X2300. The
first set of voucher specimens are filed at the Southern Methodist Uni-
versity herbarium (SMU). Field work for this study was supported in
part by the National Science Foundation, G-9800.

For one collection seeds were soaked in water until the radicle appear-
ed and then sown on a culture medium (Lewis and Elvin-Lewis, 1961)
to facilitate more rapid growth. After pretreatment for 1-2 hours in
supersaturated paradichlorobenzene, the seedlings were fixed and stain-
ed following the procedure used for the buds and mitosis was studied in
root tips and apical meristems.

OBSERVATIONS.—Fifteen species and one hybrid of Sisyrinchium
from a total of 50 populations are listed in Table 1 with their chromo-
some numbers and voucher data. For 11 species the collections represent
a wide range of morphological variation including atypical individuals,
but in each case the chromosome numbers were found to be consistent.
Chromosome numbers for 9 species are reported for the first time. These
include S. minus Engelm. & Gray, n = 5; S. fibrosum Bickn., n = 8; S.
campestre Bickn., n = 16; S. intermedium Bickn., n = 16; S. laxum Otto,
n = 16; S. mucronatum Michx., n = 16; S. sagittiferum Bickn., n = 16;
S. arizonicum Roth., 2n = 34-36; S. atlanticum Bickn., n — 48. Those
for the remaining species, S. albidum Raf., S. bermudiana L. (as S.
angustifolium Mill.), S. ensigerum Bickn., S. langloisii Greene, S. mi-

7 EDITOR’S NOT was not until after this paper went to press that I was able to
reach a conclusion ae the correct names of the introduced annuals. I believe that S.
ee of Nie! pa oe is correctly ‘S. rosulatum Bicknell, and S. micranthum is rather S. exile

ion ell. See ‘Annual Sisyrinchiums (Iridaceae) in the United States,” this issue, pp. 32—42

. A. Bere

SED AGN (Ly a43= 4B 19 62%

cranthum Cav., and S. pruinosum Bickn., agree with counts by Bowden
(1945) and Lewis and Oliver (1961).

The number for S. minus Engelm. & Gray adds a new basic number
of x = 5 to the genus (Fig. 1, 2). Although an undetermined diploid
species in the x = 8 series has been reported from South America
(Bowden, 1945), the number of n = 8 for S. fibrosum Bickn. (Fig. 3)
is the first report of a diploid North American species in this series. All
other pues studied, with the exception of S. arizonicum Roth., occur
in the x = 8 series at either the tetraploid or the dodecaploid love Only
a eee count of 2n = 34 or 36 is reported for S. arizonicum Roth.

here is little difference in chromosome size for most species of
Sisyrinchium (Fig. 1-9), although S. bermudiana L. (Fig. 11) has larger
chromosomes than S. atlanticum Bickn. (Fig. 10) in the same basic series.

Meiotic “irregularities” were rarely observed except for the extreme
bunching of chromosomes, This phenomenon was observed for most col-
lections and consequently only a small proportion of the meiotic meta-
phase and anaphase plates could be accurately interpreted. Despite this,
pollen were usually normal in appearance except for one collection from
1.5 miles west of the Neches River and Highway 94, Trinity Co., Texas
(Oliver, 312). In a sample of 100 pollen grains from each of several
plants, pollen was non-staining, appeared shriveled, and micropollen
were frequent. Meiosis was not observed, but the number of microspores
per PMC at the tetrad stage, and frequency based on a random sample
of 100 PMCs, was 4 microspores (6%), 5 microspores (26%), 6 micro-
spores (36%), 7 microspores (12%), 8 microspores (12%), 9 microspores
(4%), and 10 microspores (4%). With only 6% normal tetrad forma-
tion, meiosis was probably highly irregular. These plants are morphologi-
cally intermediate between S. larwm Otto and S. micranthum Cav.,
which were both growing in the immediate vicinity, and are assumed
to be hybrids between these species.

Recast any CITED
BOCHER, T. AND K. RSEN Lah Chromosome numbers of some arctic or
boreal oe oe Medd. espe 14
BOWDEN, W. M. 1945. A list es chromosomn numbers in higher plants. I. Acanthaceae
to Myrtaceae. Amer. Journ. Bot. 32
OVAS, G., AND B. SC HNACK, ae Numero de cromosomas en Antofitas de region
de cuyo (Republica Argentina). Rev. Argent. Agron. 13: 6.
LEWIS, W. H., AND M. ELVIN-LEWIS. 1961. ene for growing small rubiaceous
seeds from herbarium material. Castanea 26; 146-155.
ND OLIVER. 1961. Meiotic chromosomes in six Texas and
Mexican Nemastylis and Sisyrinchium (Iridaceae). Southwest. Nat. 6: 45-4
i P. F. 1940. Chromosome numbers in some British lanes New Phytol. 39:

17

SERMONTI G. 1948. Osservazioni sul cariogramma di Sisyrinchium striatum. Caryologia
1:

SKOTTSBERG, C. 1953. Chromosome numbers in Hawaiian flowering plants. Ark. f.
Bot. 3: 63-7

VIL MORIN, R., AND M. SIMONET. 1927. Nombre des chromosomes dans les genres
ioe Linum, et chez quelques autres especes vegetales. Compt. Rend. Soc. Biol. 96:
166-1

Table 1. CHROMOSOME NUMBERS OF SISYRINCHIUM IN
EASTERN NORTH AMERICA.

Taxon Gametic No.

S. minus Engel. & Gray

S. fibrosum Bickn.

S. albidum Raf.

S. campestre Bickn.

S. ensigerum Bickn.

S. intermedum Bickn.

S. langloisii Greene

S. laxum Otto

5

16

Voucher
TEXAS. Nacogdoches Co., Nacog-
doches, Oliver 317 (tepals purple),
Oliver 318 (tepals white); Brazos Co.,
10.7 miles SE of College Station, Oliver
2965"

GEORGIA. Pike Co., 0.5 miles N of
Pike Co.-Upson Co. line on Hwy. 19,
Oliver 327.

LOUISIANA. Bossier Par., 5.6 miles E
of Bossier City, Oliver 272+. TEXAS.
Nacogdoches Co., Fern Lake Fire
Tower, Oliver 243+, 0.6 miles S of
Nacogdoches, Oliver 241}.

ARKANSAS. Ouachita Co., 7 miles

of Wayne, Morley, 12 May 1961; Re-
public Co., 1 mile E and 0.7 miles N
of Belleville, Morley, 13 May 1961.

TEXAS. Hays Co., 1 mile S of San
Marcus, Osborne 39; Lampasas Co., 8.9
miles S of Lampasas, Osborne 42; Wil-
son Co., 10 miles S of Stockdale, Os-
borne 38.

TENNESSEE. Polk Co., 1 mile N of
Hwys. 64 and 30 junction, Oliver 325*.

TEXAS. Brazoria Co., 3.7 miles S
of Freeport, Lewis 5571, 4 miles S of
Freeport, Lewis 5569B*; Chambers Co.,
2.3 miles SW of Hwys. 121 and 87
Oliver 252; Panola Co., Lake
Murvaul, Oliver 276; San Augustine
Co., 3 miles E of Attoyac River on
Hwy. 21, Oliver 248.

TEXAS. Angelina Co., 5 miles S of
Lufkin, Oliver 310; Jasper Co., Hwys.
96 and 1004 junction, Lewis 5626;
Nacogdoches Co., Stripling Island,
Oliver 304; Newton Co., 4.7 miles S of
Newton, Lewis 5618.

&
“sO 2?
a? Ya a3
x. YW o\
| 2 ) 3

a re 4) Jat W/ ‘ : es

8% r) of ad z o> BE,
° 478 _Y 4
Pees 4a gu 90
fd

4 sa
periee, yo "36
a 90% 4° 6 e o
F N62. of § O,°7 § %,

Figs. 1-11. Meiotic tie a of ee eed drawn at ae 00 and reduced
by ca. 2% in reproduction. Fig. 1. S. 5, Oliver 296. Fi . S. minus, n = 5,
Oliver 318. Fig. 3. 8. eee —_— 8, ‘Oho "327, Fig. i 4 i n = 16 (one side
of ene Il), Oliver 302. Fig. 5. S. campestre, n = 16, Mor en 13 May 1961. Fig. 6.
S. pruinosum, n = 16, Lewis 5615. Fig. 7. S. cai icv: n = 16, Oliver 241, Fig. 8.
S. sagittiferum, a = 16, Oliver 269. Fig. 9. S. intermedum, » = 16, Oliver 325. Fig. 10.
S. atlanticum, n = 48, Chie 253. Fig. 11. S. bermudiana, n = = One 292

S. micranthum Cav. 16
S. mucronatum Michx. 16+
S. pruinosum Bickn. 16
S. sagittiferum Bickn. 16

S. lacum X micranthum 16+

S. arizonicum Roth. 17-184

S. atlanticum Bickn. 48

47

TEXAS. Angelina Co., 5 miles S of
Lufkin, Oliver 309; Hardin Co. 1.4
miles SW of Votaw, Lewis 5630; Trinity
Co., 1 mile N of Neches River, Oliver
Sie

CANADA. ONTARIO. Bruce Co., Oli-
phant, Heimburger, 17 June 1961.

ARKANSAS. Hempstead Co., 1 mile
SW of Hope, Lewis 5615+. TEXAS.
Aransas Co., 4.2 miles SW of Aransas
Pass, Lewis 5591, 0.2 miles S of Aransas
Wildlife Refuge Headquarters, Lewis
5597, 1 mile W of Aransas Wildlife
Refuge Headquarters, Lewis ;
mile NE of Rockport, Lewis 5575+;
Ellis Co., 1 mile N of Italy, Oliver 281;
Henderson Co., 5.3 miles W of Hwys.
175 and 31 junction, Osborne 48; Lam-
pasas Co., 4 miles W of Lampasas,
Osborne 45; Matagorda Co., 4.7 miles
NE of Markham, Lewis "5572; San
Patricio Co., 0.2 miles W of Welder
Wildlife Foundation Headquarters,
Lewis 5580+; Van Zandt Co., 3 miles
SE of Wills Point, Oliver 278+: Whar-
ton Co., Louise, Lewis 5599.
LOUISIANA. La Salle Par., 2.5 miles
SE of Gene, Oliver 269. TEXAS.
Nacogdoches Co., 10 miles S of Nacog-
doches, Oliver 237+; Orange Co., 3.8
miles SW of Orange, Oliver 255.
TEXAS. Trinity Co., 1 mile N of Neches
River and Hwy. 94, Oliver 312.
MEXICO. DURANGO. 19 miles SE of
Durango, Waterfall 15541 (SMU).
TEXAS. Chambers Co., 1 mile NE of
Hwys. 121 and 87 junction, Oliver 253*,
5 miles NE of Ferry Landing, Oliver
ZoLe

omosomes of 3 foes examined; otherwise the number is based on the study of 1 plant.

Chro
‘ ne collectior
+ Chromosome ne from somatic cells.

48

S. bermudiana L,

48

LOUISIANA. West Feliciana Par., 10
miles S of La.-Miss. state line on Hwy.
61, Oliver 266. TEXAS. Nacogdoches
Co., Goodman’s Bridge over Angelina
River, Oliver 308, 1 mile NE of Nacog-
doches, Oliver 316, 10 miles S of
Nacogdoches, Oliver 292,

NEW NAMES IN ARENARIA
(CARYOPHYLLACEAE)
LLOYD H. SHINNERS

Arenaria as it occurs in the Southeastern and Gulf Southwestern
United States is a very heterogeneous assemblage which for convenience
is best treated as a single genus, in the broad sense of Fernald (1919,
1950) and Maguire (1951, 1952). To it may be added a few species from
related genera in which they are anomalous or transitional toward
Arenaria. Mattfeld (1929) rejected Fernald’s union of Alsine (Minuartia)
with Arenaria, but his version of Minuartia does not carry conviction.
It too is extremely heterogeneous both morphologically and geographic-
ally, and one wonders what gain there is in maintaining two large and
poorly differentiated genera instead of one. Fernald’s arguments are
answered only in part. That the difference between a capsule dehiscing
by three valves and one dehiscing by six teeth is not great is shown for
example by Arenaria Benthamii, in which the capsule dehisces somewhat
unevenly, appearing at times to have three partly divided valves, though
ordinarily it is about equally 6-parted. In A. Drummondii, the three
valves commonly are slightly but distinctly notched at a Surely it
is better to keep these in a single admittedly diverse gen

About one point I must agree with Mattfeld. Like Sail ee (under
Alsinopsis), he associates the Texan Stellaria Nuttalltii (Arenaria Drum-
mondiz) with Arenaria patula, both species with notched petals much like
those of Cerastium, and approaching (but less extreme than) those of
Stellaria. If notched petals may occur in Arenaria, then it is plain that
other North American species of Stellaria with merely notched rather
than deeply bifid petals must also be placed there. Furthermore, since
capsule shape varies greatly within Arenaria, there is no good reason to
leave under Cerastium the handful of anomolous species which have three
styles, an essential feature of Arenaria. These changes will leave both
Cerastium and Stellaria more homogeneous, while the circumscription of
the already very heterogeneous Arenaria is not significantly altered.

The necessary new names (and one previously published but not men-
tioned in Maguire’s revision), together with a new species and a routine
new combination, may be grouped as follows.

1. SPECIES FROM STELLARIA WITH MERELY NOTCHED PETALS

II Shinners, Field & Lab. 17: 89. 1949. Stellaria Nut-
tallii T. & G., Fl. N.A, 1: 183—184. 1838. (Not Arenaria Nuttallii Pax,
1893.) Peculiar in having pedicels which become reflexed in age. Minu-
artia Nuttallii (T. & G.) Mattfeld, 1921, is illegitimate, being a later
homonym of M. Nuttallii (Pax) Briquet, 1911. Despite the similarity in
petals, this does not seem to be closely related to A. patula

SIDA 1 (1): 49—52. 1962.

50

A. Jamesiana (Torrey) Shinners, comb. nov. Stellaria Jamesiana Tor-
rey, Ann. Lyc. N.Y. 2: 169. 1827. (The spelling Jamesii was adopted
later.) There is startlingly close resemblance between this and the
Asiatic A. holosteoides (C. A. Meyer) Edgeworth in Hooker f., Fl. British
India 1: 241, 1874. The following collection extends the range of A.
Jamesiana to TEXAS. Culberson Co.: infrequent annual (sic) beneath
pines, South McKittrick Canyon, top of Guadalupe Mts., alt. 8000 ft.,
J.C. Hunter Ranch, Barton H. Warnock 12025, 5 Sept. 1954 (SMU).

A. Stephaniana (Willdenow) Shinners, comb. nov. Stellaria Stephaniana
Willd. in Schlecht., Berlin Mag. 1816 p. 194. (Not seen; reference taken
from DC., Prodr. 1: 399, 1824, and Ledebour, Fl. Ross. 1: 379—380, 1842.)
Stellaria dichotoma L., Sp. Pl. 2: 603. 1753. (Not Arenaria dichotoma
Krock, 1798, nor Moench, 1794.) “Petals divided only %2” (Popov, Flora
Srednei Sibiri 1: 406, 1957, in description of Stellaria Series Dichotomae).
Represented in North America by the following.

A. STEPHANIANA var. americana (Porter) Shinners, comb. nov.

. N.Y. ard. 1
(Porter) ae FI. seal Nat. ae (Contrib. U.S. Nat. Herb. 22
Hi. 5D) te ese, 1021,
2. SPECIES FROM CERASTIUM WITH THREE STYLES

A. anomala (Waldstein & Kitaibel) Shinners, comb. nov. (nom. nov.
by Code recommendation, the basinym being a later homonym).
Cerastium anomalum Waldst. & Kit. ex Willd., Sp. Pl. (ed. 4) 2: 812.
1799. (Not C. anomalum Schrank, Briefe ueber den Donaumoor p. 75.
1795. This reference not seen; taken from Schwarz, cited under Cerastium
dubium, below.) Stellaria viscida Bieberstein, Fl. Taur.-Cauc. 1: 342.
1808. (Substitute name for Cerastium anomalum, proposed without refer-
ence to C. anomalum Schrank, hence illegitimate.) Stellaria dubia Bas-
tard, Suppl. Fl. Maine-et-Loire p. 24. 1812. (Not seen; taken from
Schwarz, lc. Not Arenaria dubia Suter, Fl. Helv. 1: 266—267. 1802. This
itself is illegitimate, being a substitute name for A. hybrida Villars,
Prosp. p. 48, 1779, but it nevertheless prevents transfer of Stellaria dubia
to Arenaria.) Cerastium dubium (Bastard) Schwarz, Mitt. Thuering. Bot.
Ges. 1: 98. 1949

Soellner (1954) reports that C. anomalum and C. cerastoides (Arenaria
trigyna, below) are clearly allied and differ from Cerastiwm proper
cytologically.

argaea (Boissier & Balansa) Shinners, comb. nov. Cerastium

argaeum Boiss. & Bal. ex Boiss., Diagn. Ser. II. 6: 38. 1849.

A. iranica Shinners, nom. nov. Cerastiuwm persicum Boiss., Diagn. Ser.
I, 1:54. 1842. (Not Arenaria persica Boiss., 1842.)

A. Kotschyi (Boissier) Shinners, comb. nov. Cerastium Kotschyi Boiss.,
el, Oro Le LO. 2e61.

ol

A. trigyna (Villars) Shinners, comb. nov. Cerastium trigynum Vill,
Prosp. p. 48. 1779. (Not seen. Published also in Hist. Pl. Dauph. 1: 269,
1786, and 3: 645, 1789.) Stellaria cerastoides L., Sp. Pl. 1: 422. 1753. (Not
Arenaria cerastioides Poiret, 1789, nor Persoon, 1805.) Cerastiwm ceras-
toides (L.) Britton, Mem. Torr. Bot. Club 5: 150 1894. (As cerastioides.)
Cerastium lapponicum Crantz, Inst. 2: 402. 1766. (Substitute name for
Stellaria cerastoides L., hence illegitimate.) C. refractum Allioni FI.
Pedem. 2: 117. 1785

3. SPECIES FROM STELLARIA WITH SLIGHTLY EMARGINATE
R ENTIRE PETALS OR NONE

A. Fassettii Shinners, nom. nov. Stellaria muscorum Fassett, Rhodora
39: 460. (Not Arenaria muscorum Fischer ex DC., 1824.) Closely related
to the next species, but with well-developed petals exceeding the sepals.

A. fontinalis (Short & Peter) Shinners, comb. nov. Sagina fontinalis
Short & Peter, Transylv. Journ. Med. 7: 600. 1836. Shoinne fontinalis
(Short & Peter) B. L. Robinson, Proc. Amer. Acad. 29: 286. 1894. Alsine
fontinalis (Short & Peter) Britton, Mem. Torr. Bot. Club 5: 356. 1894

A. Godfreyi Shinners, nom. nov. Stellaria paludicola Fernald & Schu-
bert, Rhodora 50: 197. 1948. (Not Arenaria plaudicola B. L, Robinson,
Proc. Amer. Acad. 29: 298. 1894.) Stellaria uniflora of authors, not Walter.

4. MISCELLANEOUS

A. ludens Shinners, sp. nov. Perennis? (radix deest) tenella 14—16 cm.
alta aspectu Drymariae leptophyllae (D. tenellae). Caulis ad lineas
puberulus, apicem versus solum ramosus. Folia parva internodiis
breviora subciliata cuspidata basi subconnata, inferiora obovato-lanceo-
lata subpetiolata 6—7 mm. longa 2—3 mm. lata, superiora lineari-lanceo-
mm. longa 0.5—1.0 mm. lata. Inflorescentia laxe divaricato-
cymosa 9—13 cm. lata sat pauciflora parvibracteata. Pedicelli 3—7 mm.
longi. Sepala 2.8 mm. longa ovato-lanceolata subacuminata glabra sub-
scariosa cum costa unica viridi. Petala alba obtusiuscula sepalis quartam
partem breviora. Capsula oblongo-ovoidea calyce paulum brevior denti-
bus nunc 4 nunc 6 dehiscens. Semina matura non visa. HOLOTYPE: in-
frequent above upper spring, igneous soil, Madera Canyon on Mt.
Livermore, Davis Mts., alt. 7900 feet, Jeff Davis Co., Texas, Barton H.
Warnock 7419, 11 Sept. 1947 (SMU). Two stems, presumably from two
plants, are on the type sheet; the one at the left has 4-toothed capsules,
that on the right 6-toothed. The latter is assumed to be the normal form,

since the occurrence of 4-parted capsules is exceptional in the genus.
A. LANUGINOSA (Michaux) Rohrbach var. cinerascens (B. L. Robin-
son) Shinners, comb. nov. A. saxosa var. cinerascens B. L. Robin-
son, Proc. Amer. Acad. 29: 293. 1894. If the variable western forms are
treated as a variety rather than a subspecies, the earliest available epi-
thet in the rank is cinerascens. Not credited to Texas by Maguire, al-
though Robinson (1897, p. 240) cites A. saxosa from the Guadalupe

52

Mountains, Texas, collected by Havard. Its occurrence there is confirm-
ed by the following specimen: Guadalupe Mountains 2.7 miles north of
Pine Springs Camp near Highway 62, Culberson Co., Eula Whitehouse
17044, 22 Sept. 1946 (SMU).

REFERENCES
PHAM, A. R., T. G. TUTIN, oe oe F. WARBURG. 1962. Flora of the British
ae (2nd ban, Caryopttic pp. 211
RNALD, M. L. 1919. The unity of a genus Arenaria. Rhodora 21: 1—7. (Contrib.
Gray Herb. 57/1.)
- 1950. Gray’s Manual of Botany (8th ed.). Caryophyllaceae, pp. 611—
636.
LUDWIG, WOLFGANG. 1954. Cerastium dubium (Bast -) Schwarz (C. anomalum
Waldst. et Kit. non Schrank), ee n ueber Nomenklatur, ieee und Vorkommen
m ein, 2: 1—3.

MAGUIRE, BASSETT. 1951. Sead in the aa aaden) Arenaria in America
north of Mexico. ee » Midl. Nat
—_———_.. 952, Coeyophylacen. ia H. “A. Gleason, The New Britton & Brown
ses ae 2: ee

D, J. 1922. es zur Kenntnis der systematischen Gliederung und geo-

eeaphischen ee der Gattung Minuartia. Bot. Jahrb. 57 Beibl. 127: 13—63. (Co

nts on rs punk 1919 paper, p )

PAX ND MANN. 1934. Pea aaa In Engler & Prantl, Die
Nee baeeaiee (2nd ed.) 16c:

ROBINSON, por a aah 1897. eens In Gray, Synoptical Flora
of North America

MALL, JO UNKEL. oe ene in 2nd ed., 1913). Flora of the Southeastern
United Stace Aine, pp. 416—42
—————___—.. 19 etanval of oe Southeastern Flora. Alsinaceae, pp. 496—504

SOELLNER, ROLAND, ae Recherches cytotaxinomiques sur le genre Cerastium. Bull.
Soc. Bot. Suisse 64:

nis "FREDERIC 1898. A revision of the genus Arenaria, Linn. Journ. Linn.
Soc. Bot. 33;

DROSERA (DROSERACEAE) IN THE
SOUTHEASTERN UNITED STATES:
AN INTERIM REPORT
LLOYD H. SHINNERS

After cursory study, I concluded that E. L. Reed had been correct in.
describing the common Texas species of Drosera as new and distinct
from the eastern D. brevifolia (1915). It accordingly appears as D. annua
E. L. Reed in my Spring Flora of the Dallas-Fort Worth Area, Texas
(1958). In the same month that the book appeared (April), a trip to
Florida enabled me to see and collect the eastern species, which proved
conspicuously different from the Texas plant. It was a distinct surprise
when two years later Dr. Carroll E. Wood, Jr., in a commentary on
Southeastern Drosera, dismissed D. annua as not worthy of recognition.
Further study in both field and herbarium leaves me completely satis-
fied not only that D. annua is a distinct species from D. brevifolia, but
that what has long passed as D. brevifolia is in fact an undescribed
species and not what Pursh named at all. This provokes skeptical
thoughts about Harvard’s “massive project” for a generic flora of the
Southeastern United States, of which Dr. Wood’s paper is a part. Before
making detailed comments, let me offer my synopsis of the South-
eastern species of Drosera.

This account is based primarily on collections in the SMU Herbarium,
and my own field observations in Texas, Louisiana, Alabama, and
Florida. I had hoped to examine the material used by Miss Wynne, whose
revision Dr. Wood mainly followed (the most notable difference being
that she did not even mention D. annua). Because the New York Botani-
cal Garden was closed for remodeling at the time, I bo ae instead
the Southeastern material (except of D. rotundifolia) from
National Herbarium. After the study was essentially finished, - became
possible to borrow from New York, but only a few specimens (kindly
selected for me by Dr. Arthur Cronquist) were examined, and as it turned
out, none had been annotated by Miss Wynne. I am grateful to the
various curators for the use of their material,

KEY TO SOUTHEASTERN DROSERA
la. Leaf blades no wider than the petioles
a. Petals 7-10 mm. long; plant 12-40 cm. tall while in flower; leaves
with gland-tipped hairs only, or (in a few specimens from north-
ern New Jersey and New York) also minutely and inconspicuously
gland-dotted; blooming July-August; southern Maryland (intro-
duced), New Jersey to Massachusetts. ............ 1. D. filiformis

SIDA 1 (1): 53—59. 1962.

54

2b. Petals 10-20 mm. long; plant 25-60 cm. tall while flower; leaves
with gland-tipped hairs and at least dorsally with numerous ses-
sile glands; blooming April-May; Southeastern Coastal Plain in
Georgia, Florida, and Mississippi (reportedly extending to South
Carolina and Louisiana)................. 0.000.000. 2. D. Tracyt

lb. Leaf blades much wider than the petioles

3a. Scape glabrous or with inconspicuous, sessile glands; stipules

prominent
4a. Petioles glabrous or with inconspicuous, sessile glands.
3. D. intermedia
4b. Petioles with few to many, moderately long hairs lacking gland-
tips
5a. Leaf blades suborbicular to reniform-orbicular, the largest
broader than long; calyx cylindrical-ovoid just before and
after flowering, the sepals thin, narrowly oblong-oblanceolate
or oblong, mostly united less than 4....... 4. D. rotundifolia
5b. Leaf blades narrowly obovate to suborbicular, the largest as
broad as long or narrower, calyx cup-shaped for funnelform-
campanulate just before and after flowering, the sepals firm,
oblong-elliptic to ovate-elliptic, united about %
5. D. capillaris
3b. Scape with gland-tipped hairs except toward base; stipules absent
or vestigial
6a. Petals 1-5, light to deep lavender-pink, 2.5-8.0 mm. long; sepals
2.5-4.0 mm. long; frequent to rather common west of Mississippi
River, rare in S. Alabama and E. Tennessee........ 6. D. annua
6b. Petals 5, pure white, 5-10 mm. long; sepals 3.0-4.5 mm. long;
Gulf and Southe astern Coastal Plain, southern Louisiana to
lorida, north to Virginia...................... . D. leucantha

1. D. FILIFORMIS se eee The Maryland record for this species
is as follows. Prince Georges Co.: Suitland Bog, E. H. Walker 4162, 5
Aug. 1947 (US). “Flowers pinkish or purple, closed. In seepage bog.
Introduced by A. V. Smith. Growing successfully and spreading. New
plants sprout from fallen leaves.” Wood reports it from North Carolina
and doubtfully from South Carolina and Georgia, but I have seen no
material from so far south.

2. D. TRACYI Macfarlane in L. H. Bailey, Standard Cyclop. Hort. 2:
1077. 1914. D filiformis var. Tracyi Diels, Pflanzenreich IV. 112: 92.
1906. For nomenclatural quibblers, it should be pointed out that Drosera
Tracyi Macfarlane is a new name, not a new combination based on
Diel’s earlier publication of it as variety (this in turn based on the then
only manuscript binomial). Wynne (1944) stated that this differed “only
in its pale green pubescence, larger size, and more robust habit” from
D. filiformis, and in 1952 apparently did not consider it worth even
varietal status. Wood speaks of it as “a very distinct taxon,” which it
certainly is. The marked difference in flowering period cannot be ex-

55

plained as due merely to difference in latitude, and this added to the
other features given in the key seems to me to warrant treating it as a
species. Wynne reports it (1952) from “S.C. to Fla. and La.” (incorrectly

definite, leafy stems, and in the long petioles and narrow leaf blades.
Said by both Wynne and Wood to extend west to Texas in the Gulf
States, but I have seen no specimens from west of Mississippi. ALA-

New Hanover, Onslow. SOUTH CAROLINA. Aiken ae ae Ker-
shaw, Lexington, VIRGINIA. Princess Anne. momen July-August.
Petals white (color noted on only one specimen out of 41 examined).

4, D ROTUNDIFOLIA L. No material of this species was borrowed.
The following two specimens at SMU are the ace ones seen from the

Southeast. NORTH CAROLINA. Henderson Co:: peat bog
at East Flat Rock, Don Correll 3321, 27 Taig Ges a be cae as
“Drosera capillaris Poir.?”). Macon Co.: wet sphagnous pockets on

ledges of cliff; Horse Cove, near Highlands, W. B. Schofield 9071, 7
July 1958. Said by Wynne and Wood to extend south to Georgia and
South Carolina (latter queried by Wood). Color not noted on any speci-
mens at hand (32 sheets, mostly Northeastern); described by Fernald
as eae rarely pin

. CAPILLARIS eorer Encycl. Meth. Bot. 6: 299. 1804. D. brevifolia
pe Fl. Am, Sept. 1: 211. 1813 (‘1814’). Not D. brevifolia of authors
from Chapman (1860) on, which is mostly D. leucantha, below. Pursh’s
entire description follows:
3. D. pusilla; scapis radicatis simplicibus, foliis brevibus brevifolia

In sandy swamps of Georgia, Enslen. June. v.s.

in Herb. Enslen. The smallest of all the species
known; flowers rose-coloured.

Without seeing the type (location unknown; possibly not in existence;
Diels cites another Enslen specimen bearing no locality beyond Southern
U.S.) there may be a little doubt as to what Pursh had, since the
description says nothing about pubescence on the scape (though failure
to mention it suggests there was none) or presence of stipules. Pursh
listed only four species, the other three being D. rotundifolia, D. long-
vfolia, and D. filiformis; he makes no mention of D. capillaris. But his
statements “smallest of all the species known; flowers rose-coloured”
apply exactly to D. capillaris among the Southeastern species. The words
cannot possibly refer to the plant with large, white flowers which

56

Chapman and later authors mistakenly have called D. brevifolia. The
phrase “foliis brevibus cuneatis vix petiolatis” of course excludes
filiformis, and makes both the very long-petioled D. intermedia and the
round-leaved D. rotundifolia very unlikely candidates. Enslen collected
in Lower Georgia, which I take to mean the Coastal Plain. The only
species in this area to which Pursh’s description reasonably applies is
D. capillaris. The amplified descriptions of D. brevifolia given by
Nuttall and by Torrey & Gray indicate that they applied the name
chiefly to D. capillaris (which name they also do not mention), but in-
cluded forms of other species under it. Occasional plants with rather
long, narrow leaves superficially resemble D. intermedia. I have seen
specimens from the following states and counties (parishes). AL LABAMA.
Mobile. FLORIDA. Duval, Escambia, Glades, Highlands, Hillsborough,
n

GE Baker, Su : A. Beauregard, Rapides, St
Tammany. MISSISSIPPI. Jackson; also “Avondale” (county not de-
termined). SOUTH C I Berkeley, Charleston, Georgetown

February-June, and less freely July-September. Petals pale lavender-
pink or almost white.

My number 23,514 from Beauregard Parish, Louisiana, distributed as
D. annua, is D. capillaris.

6. D. ANNUA E. L. Reed, Torreya 15: 246—247, 1915. This is very
closely related to D. maritima St. Hil. of southern Brazil and Uruguay,
of which I have seen only five specimens, one of them sterile. In D.
maritima the naked portion of the scape is 1.5—2.5 (rarely —4) times
as long as the leaves, the sepals are obtuse or subacute, and the lowest
pedicel is 1.0—3.5 mm. long. In D. annua the naked portion of the
scape is 2.5—7.0 times as long as the leaves, the sepals are acute or
subacute, and the lowest pedicel is 1—5 mm. long. These differences
may appear slight. But considering how closely herbarium specimens
of D. annua and D. leucantha may resemble each other when well-
ae flowers and color data are lacking, while live plants could not
possibly be confused, I prefer to treat the North and South American
plants as two species. Some rather robust specimens collected by Dr.
B. C. Tharp on Padre Island, Texas, greatly resemble the South Ameri-
can species. At the other extreme, collections sees by Dr. H. K. Sven-
son in Coffee and Franklin Counties, Tenn e, are exceptionally
small. In addition to the records given re pte (1938) reports
D. annua from Latimer, Le Flore, and McCurtain counties in eastern
Oklahom

TN Mobile. ARKANSAS. Ashley, Bradley, Calhoun, Drew
Hot Springs, Jefferson, Miller, Pope. LOUISIANA. Allen, Clean,
Vernon, Winn. TENNESSEE. Coffee, Franklin. TEXAS. Anderson,
Bastrop, Brazos, Cherokee, Galveston, Hardin, Harris, Jefferson, San

57

Patricio, Smith; also Padre Island (county not determined). Flowering
late February—early June.

7. D. leucantha Shinners, sp. nov. Annua (interdum perennans?) parva
grandiflora exstipulata foliis obovatis petiolatis scapo glanduloso-pub-
escente petalis candidis 5—10 mm. longis. HOLOTYPE: St. Simon’s
Island, near Brunswick, Glynn Co., Georgia, Arthur Conquist 5255, 11
April 1947 (SMU; isotypes NY, US). “Among cabbage palmetto is moist
pale gray sand that is blackened with organic matter. Delicate perennial
(sic!). Fls. white, just beginning to open about 8 A.M. Closing again
shortly after noon.” is is D. brevifolia in the sense of Chapman,
Small, and Fernald, and in part of Wynne and Wood; not of Pursh,
which is D. capillaris, as explained under no. 5. The following additional
specimens have been seen. FLORIDA. Alachua Co.: Gainesville, Gerrit
S. Miller Jr. 438 (US). Brevard Co.: Titusville, R. E. Earle (US). Collier
Co.: East Henson Marsh, L. J. Brass 15970 (US). Duval Co.: near Jack-
sonville, A. H. Curtiss 4554 (US). Escambia Co.: 5.4 miles south of
McDavid, Shinners 29,702 (SMU). Jackson Co.: just east of Grandridge,
Shinners 26,990 (SMU). Lake Co.: vicinity of Eustis, G. V. Nash 10
(US). (Petals noted as pink, but I believe this an error; Nash also col-
lected 3 numbers of the pink-flowered D. capillaris at the same locality.)
Manatee Co.: Manatee, J. H. Simpson (US). St. Johns Co.: without lo-
cality, Miss Reynolds (Herb. J. D. Smith) (US). (Mixed collection: 6
plants of D. leucantha, 1 of D. capillaris.) Washington Co.: Caryville,
Shinners 27,018 (SMU). GEORGIA. Chatham Co.: 0.5 erie south of
Savannah, Gilbert G. Rossignol (US). LOUISIANA. Calcasieu Parish:
Dequincy, F. W. Pennell 10236 (NY). (Mixed collection, partly D. an-
nua.) St. Tammany Parish: vicinity of Covington, Bro. Anect 29 (US).
Tangipahoa Parish: Hammond, Lewena Gallup 11 (US). MISSISSIPPI.

Harrison (US); Gerrit S. Miller Jr. (US). Nansemond Co.: south of
South Quay, Fernald & Long 12089 (US). Sussex Co.: about 4 miles
northwest of Homeville, Fernald & Long 9940 (US). Flowering Feb-
ruary—June,

This very striking species was first described by Stephen Elliott, who
mistakenly called it D. rotundifolia. He stated that it is annual, an

scribes it as annual without rhizomes. Small’s Flora follows Chapman’s
nomenclature, but says that it is “biennial, or perennial by short root-
stocks.” In his Manual, there is no mention of rootstocks or duration.
Fernald, also following Chapman’s nomenclature, describes the whole

58

genus as consisting of “low perennials or biennials” but says nothing
further about the life-span of this species; he states “petals white, 5—
mm. long” and “the large flower closing at noon.” Wynne makes no
mention of duration. Her description of the petals as “white to pink,
45 mm. long” is obviously based on a mixture, and does not apply to
most of the plants belonging to D. lewcantha. Wood considers the species
basically perennial, but “apparently behaves as an annual in many areas
and has been so described (D. annua Reed).” Like Wynne he of course
was referring to a mixture, but nothing he included under the name is
typically perennial.

DROSERA AND THE SOUTHEASTERN GENERIC FLORA

A flora is an account of the species of plants of a given area. A generic
flora is a contradiction in terms. It is not a flora, but a device for
evading the trying job of writing one. In the case of the Droseraceae,
in which Small both failed to splinter the genera and overlooked the
fact that Rafinesque had done so, a discussion of genera seems par-
ticularly pointless. When in the course of it we are given a rehash of
an unsatisfactory previous account of the species with the addition of
fresh error, one may well ask just what good it all is.

The completeness of information in a flora depends on the amount
of direct study that has been done on the plants and in the area con-
cerned. When such study has been done by many persons over periods
of hundreds of years, it is possible to produce a work like Clapham,
Tutin & Warburg’s recent Flora of the British Isles, including details
on habitat preferences, pollination, chromosome number, and so on.
The phrase “biologically oriented flora” is to me altogether meaning-
less. When we consider Drosera in particular, we find that out of 33
references cited by Wood, not one relates to the Southeast as delimited
for the proposed generic flora, and a previous bibliography of 3% pages
to which reference is made likewise contains not a single title reporting
work done on the plants in that area. To offer all this as material for
a “biologically oriented” flora of the Southeast is as preposterous as it
is pretentious.

What is needed first and foremost is direct study of the Southeastern
plants themselves. The best of the insufficient authentic information
we have is to be found in the floras of Elliott, Chapman, and Small,
and in the publications of Roland M. Harper — men who lived in the
South or did extensive field work there. But this is not enough for a
good, up-to-date flora of the area. Much activity is now under way
there, but it is hampered by the very lack of a manual. The greatest
contribution that could be made toward a definitive flora of the South-
east is a concise interim guide as complete as present knowledge permits.
It is far too early to think of an encyclopedic treatise.

59

An erudite compilation from publications largely only tangential or
irrelevant may be of interest to some, but it is not the material out of
which a Southeastern Flora of any kind can be made

REFERENCE

S
BARCLAY, HARRIET G. 1938. A Breanne report of the ecology of a Drosera
meadow. Proc. Okla. Acad. Sci. 18: 22—

Sane A. W. 1860. Drosera. Fl. is ‘US. pp. 36—

HAM, A. R., T. G. TUTIN, AND E. F. oa 1962. Flora of the British
Hee ae ed.).

DIELS, L. 1906. Drosera. Pflanzenreich IV. 112: 61—128.

ELLIOTT, STEPHEN. 1817. Drosera. Sketch Bot. S.C. & Ga. 1: 375—376.

a lrecer MERRITT LYNDON. 1950. Drosera. Gray’s Man. Bot. (8th ed.) pp. 729—
73

are THOMAS. 1818. Drosera. Gen. N. Amer. Pl. 1: 141—142.
PURSH, FREDERICK. 1813 ee Drosera. Fl. Am. ie 1: 210—211.
REED, E. L. 1915. Drosera annua sp. . Torreya 15:
1916. Ecologic notes on eee annua. ae 5G 125—130.
SHINNERS, LLOYD H. 1958. Drosera. Spring Fl. Dallas-Fort Worth Area p. 169.
SMALL, JOHN KUNKEL. 1903. Drosera. Fl. S.E. ae

1933. Drosera. Man
eat JOHN, AND ASA GRAY. 1838. ae e N.A. 1: 145—147.

OD, LL E., The genera of i aaa pe Droseraceae of the
Seen ce States. Bre Soun Arnold A

eee foes E. 1944. Drosera in eastern ee America. aa Torr. Bot. Club
71: 166—

. 1952. Drosera. In Gleason, New Britton & Brown Ill. Fl. 2: 252—253,

KEY TO SOUTHEASTERN GLABROUS-STYLED
TEPHROSIA (LEGUMINOSAE)
LLOYD H. SHINNERS

The thorough study of the American barbistyled species of Tephrosia
by Wood (1949) included all the Southeastern members of the genus
but a handful which in the latest available accounts (Rydberg, 1923;
Small, 1933) are placed under Cracca. Necessary corrections of nomen-
clature under Tephrosia (nomen conservandum) are summarized below,
together with one new species which has come to light. Grateful ac-
knowledgement is made for the loan of material from the University of
Florida and the New York Botanical Garden.

KEY TO THE SPECIES
la. Stem densely spreading-pubescent................... 1. T. corallicola
lb. Stem appressed-pubescent (sometimes with a few spreading hairs
also) or glabrate.
2a. Leaflets 2.8—9.0 mm. wide, mostly over 4 mm.
3a. Stipules 1—3 mm. long; flowering in fall.......... 2, 1, Curtissit
3b. Stipules 4—8 mm. long; flowering in spring....... 3. T. Semino
2b. Leaflets 1.0—4.5 mm. wide, mostly under 3.5 mm..4. T. angustissima
1. T. CORALLICOLA (Small) Leon, Fl. de Cuba 2 (Contrib. Ocas.
Mus. Nac. De La Salle, Habana, 10): 304. 1951. (According to Gray
Herbarium Card Index.) Cracca corallicola Small, Bull. Torr. Bot. Club
36: 160. 1909. Known only from Dade County, Florida; type “in pinelands
between Cocoanut Grove and Cutler,” Small 2112 (isotypes examined,
FLAS, NY). Flowering in November
2. T. Curtissili (Small) Shinners, comb. nov. Cracca Curtissii Small ex
Rydberg, N. Amer. Fl. 24 (3): 179. 1923. Type from “sand ridges near
Cape Malabar,” Brevard County, Florida, Curtiss 584* (isotype ex-
amined, NY). A second collection has been seen, from beaches near
Jupiter Inlet, Palm Beach County, Florida, Curtiss 5561 (FLAS, NY).
Flowering in September.
3. T Seminole Shinners, sp. nov. Videtur perennis parva subdecumbens,
ne strigosa cum pilis paucis patentibus. Stipulae deltoideo-setaceae
oblo at

laxi. Pedicelli 6—7 mm. longi. Calyx 5 mm. longus laxe appresso-pilosus
dentibus tubae longitudine sesquilongis. Corolla 8—9 mm. longa. HOLO-
TYPE: Godden’s Mission, Big Cypress (Collier County?), Florida, Perley
Poore Sheehan, 12 March 1919 (NY). “Medicinal Plants of the Seminole
Indians. Devil’s shoe-string—(cracca purpurea): Used as a specific for

SIDA 1 (1): 60—62. 1962.

61

nose-bleed. The whole plant is steeped in cold water, and the decoction
is then used as a wash.—Plant a short-stemmed vine with compound
leaves and small but handsome flowers; the fruit is a narrow pod. ty
grows on prairies.’ Noted by Wood in 1948 as “not JT. purpurea.”
PARATYPE: East Florida, Dr. Leavenworth, no other date (sterile;
determined as Cracca Curtissii) (NY).

Known only from the above two fragmentary specimens. Stem rather
thinly strigose and with some spreading hairs. Stipules deltoid-
setaceous, 4—8 mm. long, persistent. Leaflets 9—15, oblong-oblanceolate,
18—34 mm. long, 2.8—5.0 mm. wide, obtuse and mucronate, rather thinly
strigose beneath, glabrouse above, the veins prominent beneath and
rufous. Racemes rather short-peduncled, both axillary and terminal,
slender and loose, the lower nodes (seen on only 1 sheet in early flower)

2—32 mm. apart, a reduced leaf sometimes present at lst or 2nd node.
Pedicels 6—7 mm. long in flower. Calyx 5 mm. long, rather loosely ap-
pressed-pilose, the acuminate teeth 1.5 times as long as the tube. Corolla
8—9 mm. long (as pressed). Fruit not seen.

4. T. ANGUSTISSIMA Shuttleworth ex Chapman, Fl. S. U.S. p. 96.
1860. Cracca angustissima (Shuttleworth) Kuntze, Rev. Gen. 1: 174. 1891.
Tephrosia purpurea (Shuttleworth) B. L. Robinson, Bot. Gaz. 28: 201.
1899. Frequent in Dade County, Florida; one collection seen from Bre-
vard County (near Eau Gallie; NY). Flowering May—early December.

EXCLUDED SPECIES
T. CINEREA (L.) Persoon, Syn. Pl. 2: 327. 1807. Cracca cinerea (L.)
Morong, Ann. N.Y. Acad. Sci. 7: 79. 1892. This is known only from an
old record as a waif in ballast ground at Mobile, Alabama, in June,
1888 (? last numeral not clear on label), collector not named but pre-
sumably C. Mohr, No, 19 (NY). There is no evidence that it has per-
sisted. It may be distinguished by emending the above key as follows.
3b. Stipules 3—8 mm. long.

4a. Corolla 8—9 mm. long; leaflets rather thinly strigose beneath.
T. Seminole

4b. Corolla 11—13 mm. long; leaflets densely strigose beneath.
T. cinerea

ADDENDA ON BARBISTYLED SPECIES
T. FLORIDA (F. G. Dietrich) C. Wood var. gracillima (B. L. Robin-
son) Shinners, comb. nov. T. ambigua var. gracillima B. L. Robinson,
Bot. Gaz. 28: 201. 1899. I believe that there is sufficient morphological
difference combined with geographic separation to justify recognition
of this variety.
T. MOHRII (Rydberg) Godfrey, Brittonia 10: 169. 1958. Cracca Mohrii
Rydberg, N. Amer. Fl]. 24 (3): 164. 1923. Reasons for maintaining this
as distinct from T. virginiana are given by Godfrey, 1. c.

REFERENCES

RYDBERG, PER AXEL. 1923. Fabaceae-Galegeae, Cracca. N. Amer, Fl. 24 (3): 157—
183.
SMALL, JOHN KUNKEL. 1933. Cracca. Man. S.E. Fl. pp. 704—708.

WOOD, CARROLL E., JR. 1949. The American barbistyled species of Tephrosia (Legu-
minosae). Rhodora 51: 193—231, 233—302, 305—364, 369—384, (Reprinted as Contrib.
Gray Herb. 170.)

S | Dd A CONTRIBUTIONS
TO BOTANY

VOLUME 1 NUMBER 2 NOVEMBER 1962
CONTENTS

Chromosome numbers of Linum ({Linaceae)

from the Southern United States and Mexico 63
Calamintha (Labiatae) in the Southern United States 69
Synopsis of Collinsonia (Labiatae) 76
Synopsis of Conradina (Labiatae) 84
Synopsis of Dicerandra (Libiatae) 89
Vegetative key to woody Labiatae

of the Southeastern Coastal Plain 92
Micromeria Brownei and its allies (Labiatae) 94
NOTES: Weed tl eh in St. Augustine grass sod in ve xas 9. Spa eae oldies azurea
(Pontederiaceae) in Texas Coastal Bend: new to the ted States. 99.— eae
(Commelinaceae) in ie Southern United States. 100. * siphonychia Ba hes a Par
chia (Caryophyllaceae). 101.—Stellaria Corei Shinners, no (Car oe PEN
103.—Ranunculus trachycarpus (Ranunculaceae) in south cent Ha ‘eats iana: a to North
America. 104,.—Warea auriculata instead of amplexifolia of Small (Cruciferae). 105.—
gala pereacl folia instead of W. macrantha es ceae): nomenclatural corrections.
106. ana DeLoachii Oe ibe Lemmon) Shi s, airs nov. (Gentianaceae). 107.—
Agesteche ase flora (Gray) Epling var. Haverdi "Gra y) Shinners, comb. nov. (Labiatae).
107.—Scutellaria laevis AH a another endemic in Trans-Pecos Texas. 107.—Solanum
puree Annee nom. nov. (Solanaceae). 108.

SIDA is privately published by Lloyd H. Shinners, SMU Box 473,
Dallas 22, Texas, U.S.A. Subscription price $6 (U.S.) per volume of about
360—400 pages, parts issued at irregular intervals.

©
SIDA volume 1 number 2 pages 63—108
copyright 1962
by Lloyd H. Shinners

CHROMOSOME NUMBERS OF LINUM FROM
THE SOUTHERN UNITED STATES
AND MEXICO

W. PHILLIP OSBORNE AND WALTER H. LEWIS
Stephen F. Austin State College, Nacogdoches, Texas

About 150 species of Linum are recognized throughout a predomi-
nantly tropical and subtropical distribution. In North America, Small
(1907) included 48 species in Cathartolinum and only 5 in Linum,
although Winkler (1931) transferred all species to Linum. A total of
36 species have been studied cytologically of which 12 are found in
North America

MATERIALS AND METHODS.—Plants were collected in the field
in Alabama, Floriad, Georgia, Mississippi, New Mexico, Texas, and
Mexico, Immature flower buds were fixed following the procedure of
Lewis and Oliver (1961) from which PMCs are more rarely somatic
cells were examined. Occasionally, seeds obtained from commercial
sources and herbarium sheets were cultured on a medium developed
by Lewis and Elvin-Lewis (1961). Seeds were first soaked in water
for 1-2 days and 1 day after germination mitosis was observed in root
tip cells. All chromosome drawings were made wi the aid of a
camera lucida at X 2300. Vouchers for all collections are filed in the
Southern Methodist University Herbarium and duplicates have been
distributed elsewhere. We appreciate the verifications of some collec-
tions by Dr. C. Marvin Rogers, Wayne State University. Field work was
in part aided by a grant from the National Science Foundation, G-9800.

RESULTS AND DISCUSSION.—Fourteen species and varieties have
been studied from 31 localities as listed in Table 1. Three species, L.
grandiflorum (n=8, Fig. 1), L. perenne L, (2n=18), and L. usitatissi-
mum (n=15, Fig. 15), are introduced into North America and our re-
sults verify those counts by previous workers. Of the remaining species,
all indigenous to this continent, the numbers for L. lewisii (n=9, Fig. 2)

nd L. rigidum (n=15, Fig. 15) verify those of Kikuchi (1926, 1929),
een (1933), and Ray (1944). However, Ray (1944) reported L.
medium as n—15, but from 22 plants collected in Florida, Mississippi,
and Texas, only n=18 (Fig. 8-9) and 2n=36 were found. From plants
collected in North Carolina, Ray also reported L. virginianum with
n—15, but from Georgia we found 2 plants with n=18 (Fig. 11). Un-
fortunately Ray does not appear to have preserved voucher specimens
so that it is not possible to check his determinations. The first chromo-
some numbers are reported for L. arenicola (n=18, Fig. 4), L. flori-
danum var. chrysocarpum (n=18, Fig. 5), L. greggii (n=18, Fig. 6-7),
L. imbricatum (n=15, Fig. 12), and L. pratense (n=9, Fig. 3)

SIDA I (2): 63—68. 1962.

65

Although meiosis was generally found to be regular, a somewhat lower
chiasmata frequency than is normal often resulted in incomplete bi-
valent formation. Consequently the chromosome numbers of PMCs at
diakenesis (Fig. 8) and prometaphase (Fig. 12) were usually obscure
and difficult to determine.

The North American species of Linum were grouped by Winkler
(1931) into two sections, Linum (Eulinum) and Cathartolinum. The
latter, considered by Small (1907) of generic rank, includes a majority
of the North American species and was divided by Small into nu-
merous sections. Those species having styles united one-half or more,
large yellow petals, alternate leaves, and, so far as known, a chromo-
some number of x=15, form a natural group in Linwm which Small
recognized by his sections Rigida, Multicaulia, and possibly Sulcata
under Cathartolinum. Thus LINUM, section MULTICAULIA (Small)

imbricatum (type
encries) and L. rigidum. Additional research may prove that other
species particularly those from the sections Rigida and Sulcata may be
included in this section.

bo
ol
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REFERENCES
Eee A. C. 1933. Chromosome numbers in flax (Linum). Science 78 (2027): 409.
KIKUCHI, M. 1926. Studies on the difference of chromosome numbers in Iinum species.
Jour. Agric. and For (Sapporo). 81: 26-37.
1929. Cytological studies of the genus Linum. Jap. Jour. Genet. 4:

202-212.
WIS, W. H. and M. ELVIN-LEWIS. 1961. Medium for growing small rabiaceous seeds
from herbarium material. Castanea 26: 146-1
and . oo Gee 1961. Cytogeosraphy and phylogeny of the North
American species of Verbena. Amer. Jour. Bot. 48: 638-64
AY, . JR. 1944, Goren Hae on the flax genus, Linum. Amer, Jour. Bot.
31:241-24
SMA am i K. 1907. Linaceae, in N. Amer. Fl. 25
WINKLER, H. 1931. Linaceae, in A. Eaeice tig o eee Die Nat. Pflanzenfam. 19a:
82-120

Figs. 1-15. Chromosomes of Linum drawn with the aid of a camera lucida originally at

X 2300 reduced by 21% in reproduction. Fig. 1. L. grandiflorum, n = 8, Osborne 70.
ig. 2. L. lewissi, n = 9, Osborne 78 B Fs . pratense, n = 9, Osborne sh Fig. 4.
L. arenicola, n = 18, Osborne b4. Fig. L. floridanum var. cbr ysoeer rpum, = 36

Osborne 69. Fig. 6.01. sue teh eee II with two poles), Lewis as hit 7.
L. greggii, n = 18, Lewis 5752

j gf f ip

67

Table 1. CHROMOSOME NUMBERS FOR 13 SPECIES OF LINUM

Taxon n

an

Voucher

Section Linum
L. grandiflorum Desf. 8

L. lewisit Pursh 9

L. perenne L.
L. pratense (Norton)
Small

9
L. usitatissimum L. 15
Section Cathartolinum
L. arenicola (Small)
Winkler 18

L. floridanum
(Planchon) 18
Trelease var.
chrysocarpum Rogers
L. greggi (Engelm.)
Small 18

. 8. L. medium var. texanum, n =
. 10

Fig
1 = ‘1s, Osborne 55. Fi
CAS) i

i i
(anaphase II with ae poles), ee en) Fig

TEXAS. Harrison Co., Karnack (culti-
ania Osborne 70 (4).*

AS. Howard Co., 9.5 miles
a Spring on Hwy. 80, Osborne 78 (3).
Rowe’s Seed (2)

TEXAS. Dallas Co., Kiest Park, Dallas,
Osborne 47 (2), 51 (4); Hill Co, 8.7
miles N of Hillsboro, Oliver 284 (2);
Somervell Co., 9.4 miles N of Brazos
River on Hwy. 50, Osborne 50 (4).
TEXAS. Harrison Co., Karnack (culti-
vated), Osborne 80 (4).

FLORIDA. Monroe Co., Big Pine Key,
Osborne 65 (2), Park Key, Osborne
64 (3).

MISSISSIPPI. Hancock Co., 6.2 miles
ENE of Pearl River on Hwy. 90, Os-
borne 69 (2)

MEXICO. COAHUILA. 3.1 miles N of
Los Llanos, Lewis 5743 (2). NUEVO
LEON. 11 miles E of junction of Hwys.
o7 and 60, Lewis 5752 (5).

, Osborne 54, Fig. 9. L. medium var. texanum,

= 18, Osborne 75. Fig. 11. L. virginia-
2. L. imbricatum, n = 15, Lewis 5588. Fig. 13. L. ri-
r 285. Fig. 14 idum var. rigidum, n == 15

g. 15. L. usitatissimum, n = 15, Oboe 80.

68

L. medium (Planchon) 18 36

Britton var.
texanum
(Planchon) Fern.

L. schiedeanum S.&C. 18

L. virginianum L. 18

Section Multicaulia
L. imbricatum (Raf.)
Shinners 15

L. rigidum Pursh var. 15
rigidum

L. rigidum Pursh var. 15
berlandieri (Hook.)

3

* Number of plants examined.

oOo

FLORIDA. Collier Co., 7.1 miles E of
Naples, Osborne 63 (2); Sarasota Co.,
1 mile W of junction of Hwys.

Til, ‘Ostorne 61 (2), ae ccroat
Hancock Co., 0.5 miles W of St. Louis
Bay on Hwy. 90 Osborne 68 (2).
TEXAS. Hardin Co., Kountze Fire
Lookout Tower, Lewis 5627 (1); Marion
Co., Jefferson, Osborne 57 (3), Lake
of the Pines, nr Jefferson, Osborne 55
(6); Nacogdoches Co., Stephen F. Aus-
tin oe eal Torest, Osborne 53
(4); 2).

ae Brewster Co., Bib Bend Na-
tional Park, Chisos Mountains, Os-
borne 73° (2). Far2)e 775 5) 7S).
MEXICO. COAHUILA. 3.1 miles N of
Los Llanos, Lewis 5730 (1).
GEORGIA. Pike Co., 0.5 miles N of
Pike-Upson Co. line and Hwy. 19,
Osborne 56 (2).

TEXAS. San Patricio Co., 5 miles SE
of Mathis, Lewis 5588 (4).

TEXAS, Bee Co., 0.9 miles N of Pettus,
Osborne 33 (2); San Patricio Co., Port
Aransas, Lewis 5590 (3).

TEXAS. Bee Co., 0.5 miles N of Tuleta,
Osborne 32 (2); Hill Co., 8.7 miles NE
of Hillsboro, Oliver 285 (3).

CALAMINTHA (LABIATAE) IN THE
SOUTHERN UNITED STATES
LLOYD H. SHINNERS

The last world-wide account of Satureja (by Briquet in Engler &
Prantl, 1897) defines the genus very broadly, merging with it Micro-
meria and Calamintha, which the generally conservative Bentham (in
DC., 1848) had retained as distinct. DeWolf, in summarizing the culti-
vated species (1954), observed that Briquet “enlarged Satureja to such
an extent that it was almost undefinable.”’ He follows several recent
European authors by recognizing five genera instead of one. The largest
of those involving wild plants of the South is Calamintha, including the
species treated in Small’s Manual as Clinopodium groups Herbacea and
Fruticosa. Only half of the six native species which I consider valid in
the two groups have had proper combinations published for them
under Calamintha. It is primarily to supply the missing ones, and inci-
dentally to comment on their synon and taxonomy, that this brief
account has been prepared. The species sometimes referred to Satureja
but not included in Calamintha, and known to occur wild in the South
(broadly delimited), are as follows.

ACINOS ARVENSIS (Lamarck) Dandy, Journ. Ecology 33: 326. 1946.
Satureja Acinos (L.) Scheele. Clinopodium Acinos (L.) Kuntze. Neither
Fernald nor Gleason reports this European weed from south of the
Mason-Dixon Line; the following collection thus extends its range (see
also Strausbaugh & Core, 1958). WEST VIRGINIA. Hampshire Co.:
plentiful in old peach orchard near Hanging Rock, Wilbert M. Frye

95, 25 June 1949 (SMU).

CLINOPODIUM VULGARE L., Satureja vulgaris (L.) Fritsch, includ-
ing var. neogaea Fernald, Rhodora 46: 388. 1944. The species is repre-
sented at SMU by 19 sheets from North America (D.C., Michigan, New
Jersey, North Carolina, Quebec, Vermont, Virginia, West Virginia,
Wisconsin) and 11 from Europe (British Isles, Czechoslovakia, France,
Germany, Italy, Netherlands, Sweden). All have leaves pilose on both
surfaces, the European ones more variable as to density, but not separ-
able from the North American plants; most are exactly the same. On
the basis of this limited but diversified sample, it does not appear that
Fernald’s variety (supposed to differ in having leaves glabrous or only
sparsely strigose above, and to represent the native American race) is
at all tenable. The species is primarily northern, extending south in
the mountains to western North Carolina,

MICROMERIA BROWNEI (Swartz) Bentham var. PILOSIUSCULA
Gray. M. pilosiuscula (Gray) Small. M. xalapensis (HBK) Bentham.

SIDA 1 (2): 69 — 75. 1962.

70

Satureja Brownei (Swartz) Briquet var. pilosiuscula (Gray) Briquet.
Central and northern Florida, adjacent Georgia (Decatur Co.), south-
ern Louisiana (St. Bernard Parish), southern Texas (Brazoria to
Cameron counties near the coast, locally inland in Bexar and Colorado
counties); through Mexico to Guatemala. M. Brownei var. Brownei is
confined to Jamaica; another variety is found in Cuba (rare) and
Mexico (Yucatan Peninsula), and closely related species in the Bahama
Islands and Hispaniola (Shinners, 1962).

PILOBLEPHIS RIGIDA (Bartram) Rafinesque, New Fl. N.A. 3:
52—53. 1838 (“1836”). Satureja ? rigida Bartram ex Bentham, Lab. Gen.
et Sp. p. 354. 1832—1836. (Not seen; quoted in DC., Prodr. 12: 211.
1848.) Pycnothymus rigidus (Bartram) Small, Fl. S.E. U.S. (ed. 1) p.
1042. 1903. This is one of the many cases in which Small needlessly
supplied a new name, overlooking the much older one of Rafinesque.
The species occurs almost throughout peninsular Florida.

The following key and notes on Calamintha are based chiefly on
collections at Southern Methodist University, Florida State University,
and the University of Florida. Material of the introduced species was
borrowed from the Gray Herbarium, and a few sheets of C. arkansana
were examined on a visit to the University of Texas. Grateful acknowl-
edgment is made to the several curators for their courtesies.

KEY TO THE SPECIES
la. Herbaceous perennial (may flower the first year, appearing annual)
2a. Stem pubescent; leaf blades elliptic to ovate

3a. Corolla 22—35 mm. long; calyx 10—13 mm. long; leaf blades
sharply and rather coarsely toothed.......... 1. C. grandiflora
3b. Corolla 7—18 mm. long; calyx 2.8—10.2 mm. long; leaf blades

entire or with shallow, rounded or incurved teeth
4a, Calyx 6.0—10.2 mm. long, the hairs inside the throat barely
or not exserted; blades of larger stem leaves 2— - cm. long
2. officinalis
4b. Calyx 2.8—6.0 mm. long, the hairs inside the een exserted,
prominent; blades of larger stem leaves 0.8—2.4 cm. long
3. C. Nepeta
2b. Stem glabrous or pubescent only at nodes; leaf blades linear to
elliptic lanceolate on flowering stems (broader on sterile shoots)
5a. Nodes glabrous or inconspicuously pubescent; middle stem
leaves 1—5 mm. wide, entire or occasionally with 1 or 2 teeth
on each margin; plant developing stolons (commonly wanting
at main flowering period), their leaves with orbicular-ovate or

orbicular-rhombic blades; corolla 7—12 mm. 1

4. C. arkansana
5b. Nodes moderately to densely pubescent; middle stem leaves
3—12 mm. wide, with 1—4 teeth on each margin; plant not
developing stolons, sometimes with ascending leafy shoots, their

71

leaves with oblanceolate to oblong-obovate blades; corolla
LS ASTI eee Old Curae a cet ae con ee ee 5. C. glabella
lb. Shrubby perennial

6a. Corolla aes mm. long, bright red or rarely yellow; calyx

(oie—oullte) te mbeghy ke) ay =h tu ee inaeneley Aa mee ane mnEUReLSTR.O “ie- 1y eam 6. C. coccinea
6b. Corolla ions mm. long, light lavender or lavender pink with
ark dots; calyx 6.0—7.5 mm. long

7a. Leaves subsessile, minutely and densely pubescent
8a. Leaf blades linear- to oblong-lanceolate, widest about middle,
not strongly tapered at base, with entire, revolute margins
. C. Asher
8b. Leaf blades oblanceolate or obovate, tapered at base, the
lower more or less toothed, the upper often entire and with
VOLUPCISI AR SINS es oo pu kok ate eee eee ee eae ~ CG. -dentata
filgy, oe PEloOled slabrOus 2 ts sion ee 9. C. georgiana

1. C. GRANDIFLORA Moench. Satureja grandiflora (Moench) Scheele.
There is no reliable record of the occurrence of this as a wild plant.
In the Gray Herbarium there is one specimen from the herbarium of
H. P. Sartwell, “Legit Curtiss,” the habitat given as “Carol.” followed
by an illegible word, seemingly ‘Lenten’ but possibly ‘“Septen.” was
meant. To this has been added, in Asa Gray’s handwriting, ‘wild?
Calamintha officinalis L.” The plant, native of Europe, is cultivated
in the United States. It has been in my garden in Dallas for two years,
barely surviving, and showing no inclination to bloom. Unless fresh
evidence is forthcoming, this species should be excluded from the
Southern flora

Ce OFFICINALIS Moench. Satureja Calamintha (L.) Scheele. The
only North American specimen I have seen is the following, at the
Gray Herbarium, VIRGINIA. Isle of Wight Co.: rich calcareous slopes
along James River, west of old Fort Boykin, Fernald & Long 13739,
8 Sept. 1941. (Flowers past: calyx 6 mm. long, villous.)

3. C. NEPETA (L.) Savi. Clinopodium Nepeta (L.) Kuntze. Satureja
Nepeta (L.) Scheele. S. Calamintha var. Nepeta (L.) Briquet, var.
nepetoides (Jordan) Briquet, and var. glandulosa (Requien) Briquet.
The C. Nepeta-C. officinalis complex is represented at SMU by 21
sheets from Europe and 10 from North America; an additional 46 from
North America were borrowed. I am unable to follow with this ma-
terial the very detailed account given by Briquet in Les Labiées des
Alpes Maritimes or the very brief one given by Fernald. In one notable
case, I found myself trying to call duplicates of Ahles 17630, from
Granville County, North Carolina, by two different names, though
when placed side by side they obviously represented only one moder-
ately variable entity. There does seem to be a fairly good break be-
tween C. Nepeta and C. officinalis, and they are accordingly accepted
as distinct though closely related species. C. Nepeta was found by Asa

72

Gray and J. Carey in Virginia and North Carolina in July, 1841, and
had appeared at scattered localities elsewhere before the end of the
19th Century. It seems to have continued to spread gradually, but
avoids the Coastal Plain. The record for Mobile County, Alabama, listed
below, presumably represents a waif only; there are no recent collec-

s Be
Botetourt, Caeaabell, Caroline, Chesterfield, Giles, Goochland, Hanover,
James City, Rockbridge, Rockingham, Shenandoah, Smyth, Spotsylvania,
Wythe, York. (Rather surprisingly, Strausbaugh & Core, 1958, do not
report it from West Virginia

4. C. arkansana (Nuttall) enna: comb, nov. Hedeoma arkansana
Nuttall, Trans. Amer. Phil. Soc. n.s. 5: 186. 1834. “In moist and rocky
prairies near the sources of the Kiamesha river. Flowering in May and
June.” (The locality is in eastern Oklahoma, at that time part of
Arkansas Territory; see Geiser, 1956.) Calamintha Nuttalhi Bentham
in DC., Prodr. 12: 230. 1848. (Bentham cites “Micromeria Nuttallit
Tor t Gr. ms.,” which presumably is the real original for the name.
se ane by present rules, since the epithet arkansana was avail-

C. glabella var. Nuttallii (Bentham) Gray, Man. (ed. 2) p. 307—

308. 1857. (Doubly illegitimate, since both the preceding and the ine
ing are cited.) Micromeria glabella var. angustifolia Torrey, Fl. N.-Y.
2: 67. 1843. Satureja glabella var. angustifolia (Torrey) Svenson, Rho-
dora 42: 7—8. 1940. Calamintha glabella var. angustifolia (Torrey)
DeWolf, Baileya 2: 150. 1954. (Basinym incorrectly given as Satureja
glabella var. angustifolia (Torrey) Svenson.) The name Hedeoma glab-
rum was used by Nuttall (Genera 1: 16, 1818) for this species and the
next which he did not at first separate; it is nomenclaturally an
illegitimate new name for the next species, proposed by Persoon.

Despite the great similarity in general appearance between this and
the next, I agree with Fernald that the two are to be regarded as dis-
tinct although closely related. The peculiarity of leafy stolons (seldom
present on herbarium specimens) needs to be studied during periods
when the plant is not in flower. C. arkansana blooms from late May to
early August. It is a plant of limestone areas, extending southwest
from the Ozark region into central Texas, where it is rare. ARKANSAS
Baxter, Fulton, Izard, Lawrence, Randolph, Sharp, Stone. OKLAHOMA
Murray, Pontotoc, Rogers, Seqouyah. TENNESSEE. Wilson. TEXAS.

is.

. GLABELLA (Michaux) Bentham, in DC., Prodr. 12: 230, 1848.
Cunita glabella Michaux, Fl. Bor.-Am. 1: 13. 1803. “In rupibus riparlis
fluvii Tennassee, juxta Nashville.” Hedeoma glabrum Persoon, Syn. Pl.

73

2: 131. 1807. (The change in form of the specific epithet was probably
only a slip of the pen, reference being made to ‘‘Michx. sub Cunila.”
Pursh and Nuttall follow Persoon but change the gender to feminine,
which under present rules is not permissible.) Satureja glabella
(Michaux) Briquet in Engler & Prantl, Nat. Pflanzenfam. Teil IV
Abt. 3a: 302. 1897.

Flowering late May—July. Said by Fernald to occur in Kentucky,
Tennessee, southern Missouri, and Arkansas. I have seen specimens
from the following states and counties. ARKANSAS. Benton, Garland,
Logan, Newton, Saline, Washington. OKLAHOMA. McCurtain.

6. C. COCCINEA (Nuttall) Bentham in DC., Prodr. 12: 229. 1848.
Cunila coccinea Nuttall ex Hooker, Exotic Flora 2: t. 163. 1825. (There
is a second t. 163 opening volume 3; Bentham erroneously quotes the
latter. He aes cites “Melissa coccinea Spreng., Syst. 2: 229,” 1825, but
that page is devoted to Acer; the precise citation is Syst. 4 pt. 2: 224,
1827, where Cunila coccinea is given as synonym.) Type (not seen):
Florida, Ware (ANSP). Description pears by Hooker from culti-
vated specimens, grown by Mr. H. Shepherd from seed from the type
collection. Satureja coccinea (Nuttall) Bertoloni, Misc. Bot. 8: 23. (Not

en. Published in 24 parts, 1842—1863

ium coccineum (Nuttall) Kuntze, Rev. Gen. Pl. 2: : . Clinopo-

dry pine cae Indian River, Florida, Curtiss 2012, August (isotype
examined, AS). Satureja macrocalyx (Small) Druce, Rept. Bot.
Exch. Club ; canes 644. 1917.

In flower from April to October. There is a tendency for flowers to
be larger in the southeastern part of the range of this species( peninsular
Florida), but the range of variation is continuous from one extreme to
the other. The isotype of Clinopodium macrocalyx which I examined
has calyx only 13 mm. long, and on this basis would run to C. coccineum
in Small’s own key; it does, however, have a very large corolla (46
mm. long). ALABAMA. Baldwin, Mobile. FLORIDA. Bay, Citrus, Es-
cambia, Franklin, Gulf, Hernando, olmes, Indian River, Souk
Orange, Osceola, Pasco, Santa Rosa, Seminole, Wakulla.

Chandler, Emmanuel, Toombs, Wheeler, MISSISSIPPI. Harrison, pace
son, Stone

7. C. Ashei (Weatherby) Shinners, comb. nov. Satureja Ashei Weather:

ida, W. W. Ash
e Co., Florida. Ashe, April 1823 (GH). (These specimens not ex-
amined. The detailed original description, key, and localities, leave no
doubt as to the identity of the plant.) Clinopodium Ashei (Weatherby)
Small, Bull. Torr. Bot. Club 51: 385. 1924,
Flowering late January to October. Another of the many endemics
centering in the lake region of peninsular Florida; specimens seen from

74

Highlands, Marion, Polk, and Volusia counties. Dried plants of the next
species lacking lower leaves are superficially very similar to this.

8. C. DENTATA Chapman, FI. S. U.S. p. 318. 1860. “Sand ridges near
Aspalaga, Florida” (Gadsden Co.) ‘Sept. and Oct.” Clinopodium denta-

Abt. 3a: 302. 1897. In flower from April to October. Confined to north-
ern Florida and southeastern Georgia. FLORIDA. Bay, Gadsden, Lib-
erty, Wakulla, Walton. GEORGIA. Tattnall.

9. C. georgiana (Harper) Shinners, comb. nov. Clinopodium georgianum
R. M. Harper, Bull. Torr. Bot. Club 33: 243. 1906. New name for
Thymus carolinianus (non ‘“Walter’) Michaux, Fl. Bor.-Am. 2: 9. 1803.
Michaux described this species, but based his name on Thymbra ?
caroliniana Walter, Fl. Carol. 1 1788, which in reality was the
quite different Macbridea iis Elliott, correctly to be called M.
caroliniana (Walter) Blake, Rhodora 17: 132, 1915. DeWolf (1954)
perpetrates several errors by referring this to “Calamintha caroliniana
(Nuttall) Bentham”; neither of those authors is to be credited with the
combination. The relevant sensu synonymy is as follows. Calamintha
caroliniana (non ‘“‘Walter”) Sweet, Hort. Brit. (ed. 2) p. 408. 1830. Sweet
quotes only Thymus grandiflorus “B. M.” (i.e., Sims, Bot. Mag. 25: t.
997, 1807). Sims gives both Thymus ee gas Michaux and Thymbra
? caroliniana Walter as synonyms, with query, and goes on to say
“We have very little doubt that Walter’s plant is the same with ours,
but are not sufficiently certain with respect to Michaux’s, to dare to
adopt his specific name.” It would be extremely difficult to decide
whether Sims’s name was legitimate or not on the basis of his state-

(L.) Scopoli, Fl. Carniolica (ed. 2) 1: 424, 1772. Similarly Calamintha
grandiflora (Sims) Pursh, Fl. Am. Sept. 2: 414, 1813 (“1814”), is illegiti-
mate as a later homonym of C. grandiflora (L.) Moench, 1794. (In fact
it is doubly illegitimate because Pursh cites all three of the binomials
mentioned by Sims without query, but failed to adopt the oldest one.)
Satureja carolinana (non “Michaux”) Briquet in Engler & Prantl,
Nat. Pflanzenfam. Teil IV. Abt. 3a: 302. 1897. All the binomials with
the epithet caroliniana are of course synonyms of Macbridea caroliniana
(Walter) Blake on the basis of Walter’s type, even though Michaux,
Sweet, and Briquet all meant Calamintha georgiana on the basis of
their descriptions,

Flowering from May to October. The most widespread (but not most

Anson. SOUTH CAROLINA. Abbeville, Anderson, Chesterfield.

REFERENCES
BENTHAM, G. 1848. Labiatae: Satureia, Micromeria, Calamintha. In DC., Prodr. 12:
208—23
_ PRIQUET, JOHN. 1895. Satureia Calamintha. Les Labiees des Alpes Maritimes 3: 430—

. 1897. Satureia. In Engler & Prantl, Nat. Pflanzenfam. Teil IV. Abt.
3a: 296—303.

DEWOLF, shiver ie 1954, Notes on cultivated Labiates. 4. Satureja and some related
genera. Baileya

Pees MERRITT LYNDON. 1950. Satureja. In Gray’s Man. (8th ed.) pp. 1239—

12

Sage S. ue oe Thomas Nuttall’s botanical collecting trip to the Red River, 1819.
Field & Lab. 24:

me AY, aes Ten ye ed. 1886). Satureia, Micromeria, Calamintha. Syn. Fl. N.A. 2

358
Gai iieNae H. 1962. Micromeria Brownei and its allies (Labiatae). Sida 1:
94-97,
SMALL, JOHN KUNKEL. 1933. Pycnothymus, Micromeria, Clinopodium. Man. S.E. FI.
pp. 1067—1069
STRAUSBAUGH, P. D., AND EARL L. CORE. 1958. Satureja FI.
SVENSSON, H. K. 1940. Plants of Southern United States. II. nent ace. Hie

SYNOPSIS OF COLLINSONIA (LABIATAE)
LLOYD H. SHINNERS

The difference between two stamens and four in the Labiatae often
separates genera and whole tribes. Briquet in 1897 concluded that
Collinsonia, which had accumulated species with both numbers, should

e divi . Accordingly he segregated two species having four stamens
as Ce st overlooking the fact that Rafinesque exactly eighty years
before had published the genus Hypogon, likewise differing in having
four stamens. Although each author recognized two species in his
segregate genus, they were not all the same ones. Briquet’s Micheliella
verticillata is indeed very distinct from the others, as indicated in the
key below, and is here considered sole representative of a new subgenus.
Rafinesque’s Hypogon verticillatum, despite the confusing identity of
epithets, is an entirely different species, the long misunderstood Col-
linsonia serotina Walter (C. punctata Elliott), which may have either
two or four functional stamens, with intermediate forms having greater
or lesser development of rudiments or filaments. Even without this
awkward variation. Hypogon is too similar in all other respects to Col-
linsonia proper to justify segregation. Likewise Micheliella, separating
M. anisata (which also is in reality C. serotina) from the species it
greatly resembles and associating it with the quite dissimilar C. verti-
cillata, is obviously an unnatural genus, Collinsonia is here recognized
with its pre-1897 limits, including the peculiar C. verticillata as sole
representative of a subgenus defined on the basis of features of in-
florescence and secondarily of leaves rather than number of functional
stamens.

In addition to the material at Southern Methodist University, I have
had the use of collections from Florida State University, the University
of Florida, the Gray Herbarium (not including those referred to
Micheliella, which I carelessly forgot to explain were to be included
in Collinsonia when requesting the loan), North Carolina State College,
and the University of North Carolina. My thanks are due the various
curators for their courtesies.

COLLINSONIA L., Sp. Pl. 1: 28, 1753, and Gen. Pl. (ed. 5) p. 16, 1754.
Only original species and automatic type: C. CANADENSIS L.
Hypogon Rafinesque, Fl. Ludov. p. 148. 1817. (Also on p. 41 as
nomen provisorium.) Lectotype species: H. verticillatum Raf, =
COLLINSONIA SEROTINA Walt.

Diallosteira Rafinesque, Neogenyton p. 2, 1825. (Reference for this
and the next not seen; taken from Merrill, Index Rafinesquianus
p. 206 and p. 209, 1949.) Type (only) species: D. punctata (Ell.)

Raf. ex Jackson = COLLINSONIA SEROTINA Walt.

SOA ts (2s e293, 1862.

77

Pleuradenia Rafinesque, Neogenyton p. 2. 1825. Two species: P. prae-
cox (Walt.) Raf. ex Jackson (“precox”’’), ee uncertain (either
Collinsonia serotina Walt. or C. tuberosa Michx.); P. scabra
(“Pers.”) Raf. ex Jackson = COL agSOITES au vnar Walt.

Micheliella Briquet in Engler & Prantl, Nat. Pflanzenfam. Teil IV.
Abt. 3a: 325. 1897. Lectotype species: M. verticillata eae
Briquet = COLLINSONIA VERTICILLATA Baldwin. This is
here recognized as constituting Collinsonia subg. Micheliella
(Briquet) Shinners, stat. nov., differing from subg. Collinsonia
in having flowers subverticillate or alternate instead of opposite,
lacking floral bracts, and having pedicels with enlarged flattened
bases. (See also first couplet in key below.) The number of
functional stamens is variable in subg. Collinsonia as here de-
fined and is not a usable basis for separating the subgenera.

KEY TO THE SPECIES
la. Flowers close-set in groups of 3—6 in lower and middle part of the
simple inflorescence; floral bracts absent, the pedicels with widen-
ed, flat bases; leaves 4 (rarely 2), closely crowded (subopposite or
subverticillate) in upper part of stem, all rather long-petioled;
flowering spring—early summer.................. 1. C. verticillata
lb. Flowers 2 at all nodes of the simple or branched inflorescence;
floral bracts present, minute to large, the pedicels not enlarged at
base; leaves 6 or more, opposite, the uppermost greatly reduced,
short-petioled or sessile; flowering late summer—fall.
2a. Blades of larger stem leaves 8—25 cm. long, with 11—42 teeth on
each margin, glabrous or variously pubescent beneath; plant with
large, woody, irregular, more or less elongate, rhizome-like crown
4—15 cm. long; stem 2—7 mm. thick near base
3a. Flowering calyx 3.2—7.6 mm. long, the lower (narrower) teeth
subacute to acuminate, the midvein barely or not exserted;
stamens variously 2 with 2 rudiments, or 2 with 2 short to full-
length empty filaments, or 4 and all anther-bearing; leaf blades
pubescent or pilosulous over the surface beneath; Coastal Plain,
North Carolina to southeastern Louisiana, rare inland
2. C. serotina
3b. Flowering calyx 2.3—4.3 mm. long, the lower teeth with elon-
gate, subulate tips partly formed by extension of midvein;
stamens 2 with 2 rudiments, or rarely anther-bearing stamens
none; leaf blades glabrous or variously hispidulous or pubes-
cent beneath; interior and northern (except for one outlying
locality, In anortnem. MlOmda) iin .50 6s oe eee 3. C. canadensis
2b. Blades of larger stem leaves 4.0—10.5 cm. long, with 5—15 teeth
on each margin, glabrous or hispidulous on the main veins be-
neath; plant with usually small, rounded, lobed, or elongate tuber-

like crown (up to 5 cm. across or 6 cm. long); stem 1—4 mm. thick
Nea, She Gc eee a a Ee ea 4. C. tuberosa
1 e. en Baldwin ex Elliott, Sketch Bot. S.C. & Ga.
1: 36—87. 1817. “Var. a. grows near Milledgeville, between the Oconee
and Oakmulgee rivers (Baldwin Co.), Georgia.” According to Weatherby
(1942) the type of this species, collected by Dr. Boykin, is preserved at
Charleston. — Micheliella verticillata (Baldwin) Briquet in Engler &
Prantt, Nat. Pflanzenfam. Teil IV. Abt. 3a: 325. 1897. — Hypogon verti-
cillatum (Baldwin) House, N.Y. State Mus. Bull. 233-234: 67. 1922. (Not
Hypogon verticillatum Rafinesque, which is a synonym of C. serotina;
see under no. 2, below.)
Flowering April—June. Seana ao Richmond, Walker. NORTH
CAROLINA. Polk, TENNE McMin
EROTINA Walter, 7 ee p. 7 1788. According to Fernald
agi Schubert (1948), there is a specimen in the Walter Herbarium which
is the same as C. punctata Elliott. I do not consider this or any other
specimen at the British Museum a holotype. The material preserved
there was sent to Fraser by Walter. It may include isotypes or topo-
types, or merely supplementary material, but it is not the primary
Walter collection. Because of lack of labels and mixups among those
preserved, we have no means of knowing precisely what the nature of
the specimens is. In this case the type region plus the scanty description
are In agreement with the evidence from the specimen, and I am willing
to accept the latter as added justification for adopting Walter’s name. —
C. scabriuscula Aiton, Hort. Kew. (ed. 1) 1: 47. 1789. “East Florida. Mr.
John Bartram.” The description and type locality are sufficient to estab-
lish the identity of this species. Why the name is not even mentioned in
all’s floras is a mystery; it was accepted by Chapman and by Gray
in the Synoptical Flora, though misunderstood by them. — C. anisata
Sims, Bot. Mag. 30: t. 1213. 1809. Described from cultivated material.
“Native of S.C.” C. serotina Walter is cited as doubtful synonym. The
plate, which must be taken as the type, is quite distinctive, and repre-
sents an extreme form with very short, wide calyx teeth which at first
I thought separable from C. serotina. — C. ovalis Pursh, Fl. Am. Sept.
1: 21. 1813 (“1814”). “In South Carolina, Fraser.” Referred to C. can-
adensis by Bentham, but the phrase “calycis dentibus brevissimis” ex-
cludes that ee and quite definitely applies to this one. — C. scabra
Pursh, l. c. p. 20. Illegitimate substitute name for C. scabriuscula Aiton
and C. aes Walter, both placed in synonymy without query. —
Hypogon anisatum (Sims) Rafinesque, Fl. Ludov. p. 148. 1817. — Col-
linsonia punctata Elliott, Sketch Bot. S.C. & Ga. 1: 36. 1817. “In rich
soils. Frequent.” There is a phototype at the Gray Herbarium on which
Mr. Weatherby has noted “Leaves almost tomentose is rather strong.” —
C. verticillaris Rafinesque, Fl. Ludov. p. 41. 1817. — Hypogon verticil-
latum Rafinesque, l.c. p. 142. This may have been merely a slip of the
pen, but as published is an ene substitute name for Collinsonia

79

verticillaris. Merrill (Index Rafinesquianus p. 206) mistakenly equates
this with Micheliella verticillata (Baldwin) Briquet. — Collinsonia
canadensis var. ? puberula Bentham in DC., Prodr. 12: 253. 1848. “In
Louisiana (Drumm.!) in Alabama (Rugel!).”—C. canadensis var. punctata
(Elliott) Gray, Syn. Fl. N.A. 2 pt. 1: 351. 1878. (By Fernald this name
was extended to apply to pubescent-leaved forms of C. canadensis.) —
Diallosteira punctata (Elliott) Rafinesque ex Jackson, Index Kewensis

1: 741. 1893. — Pleuradenia scabra (“Pers.”) Rafinesque ex Jackson,
l.c. 2: 562. 1894. (This properly goes back to Collinsonia scabriuscula
Aiton; Persoon merely misspelled the name.) — Hypogon verticillare

(Rafinesque) Nieuwland, Amer. Midl. Nat. 3: 178. 1914.

This is the only Collinsonia that I have seen in the field, and a merry
indoor chase it has led me. My lone collection (no. 28,956) was made
9.8 miles northwest of Loxley in Baldwin County, Alabama, 26 October
1960. The plants were past flowering, but in a few withered corollas
it was possible to find four well-developed filaments. After much ran-
sacking of synonymy, I concluded that it was the long forgotten C.
verticillaris Rafinesque. The real answer to the puzzle came with the
loan from Florida State University. Godfrey & Kral 54288, from Wacissa
Springs, Jefferson County, Florida, 20 October 1955, showed flowers in
the same inflorescence with two and with four stamens. Once stamen-

number was rejected as a taxonomic character, it became possible to
delimit Collonsonia serotina more satisfactorily. It is a variable species,
especially as to width of calyx teeth and density of leaf pubescence.
Some North Carolina specimens proved difficult to determine, closely
approaching C. canadensis. There may be introgressive hybridization in
that state, but on the whole I think the variation in the two species
is a matter of homologous mutations. The ranges of the two are almost
entirely separate. The hairs in C. serotina are rather long and slender;

or sessile glands, so that the key character used by Small is quite worth-
less. One specimen of C. serotina from Decatur County, Georgia (just
west of Jim Woodruff Dam, Richard S. Mitchell 1319, FSU) is notable
in having a simple inflorescence and only 3 pairs of stem leaves, so
that in aspect it suggests C. verticillata. The same specimen shows ex-
ceptionally broad though not short calyx teeth.

Flowering September—October. Except for a few puzzling collections
from interior North Carolina, and one unmistakable one from De Kalb
County, Georgia, this is a Coastal Plain species, from the Carolinas

Cc
Holmes, Jackson, Jefferson, Leon. GEORGIA. Decatur, De Kalb, Meri-
wether. LOU A. Washington. MISSISSIPPI. Forrest. NORTH

R
CAROLINA. Richmond, Rockingham, Stanly. SOUTH CAROLINA.
Beaufort, Lexington.

80

It is this species which William Bartram found in or near Baldwin
County, Alabama, and described in his travels. Harper, following
Mohr, identified it as C. anisata. Mohr states incorrectly (1903, p. 15)
that Bartram named it C. anisata. Bartram did not give it a specific
name, speaking of it merely as a Collinsonia.

DENSIS L., Sp. Pl, 1: 28. 1753. “Habitat in Virginiae,

Canadae sylvis.” There is no description, but reference is made to

description and figure in the Hortus Cliffortianus, and to Colden. A

Hortus Cliffortianus specimen in the British Museum was designated

lectotype by Epling, Journ. Bot. 67: 6, 1929 (phototype in Gray Her-
—C,

barium examined). canadensis var, cordata and var. ovata Pursh,
Fl. Am. Sept. 1: 20. 1813 (“1814”). — C. angustifolia Rafinesque, Med.
Fl. 1: 114, 1828. “Kentucky, Ohio, &c.”’ — The name C. canadensis var.

punctata is used by Fernald for pubescent-leaved forms of this species,
but on the basis of type is a synonym of C. serotina.

As already indicated under the preceding, this species varies greatly
in pubescence. If varieties are recognized, there really should be more
than two. On the basis of the material I have seen, I prefer to regard
these as minor variations not worthy of nomenclatural recognition. I have
seen specimens from as far north and west as Vermont, Ontario, Mich-
igan, and Indiana; it is reported in literature from Quebec (Gleason),
Wisconsin _ (Fernald), llinois (Jones & Fuller), Missouri (Palmer &

not been able to verify, is discussed at the end. Except for an outlying
station in northern Florida, it barely enters the Coastal Plain.

Flowering August—October. ALABAMA. (S. B. Buckley, without
locality.) D.C. (“Near Great Falls, Maryland.”) FLORIDA. Jackson.
G

Greenville, Lancaster, McCormick, Pickens. TENNESSEE. Carroll,
Grundy, Knox, Roane. VIRGINIA. Appomattox, Bedford, Botetourt,
Clarke, Craig, Fairfax, Fauquier, Giles, Montgomery, Rockbridge,
Smythe, Spotsylvania, Surry. WEST VIRGINIA. Sih Greenbrier,
Hampshire, Pocahontas, Summers, Tucker

In the en Flora, Gray reports C. scabriuscula as extending
west to “E. Ark.” used that name, rather strangely, for C. tuberosa,
which does extend phon canes to Memphis, Tennessee, just across the
Mississippi River from Arkansas (or did extend; it was collected there
in 1853). Unfortunately there is no voucher for the Arkansas record at
the Gray Herbarium. In later manuals only C. canadensis is credited

81

to that state. ] have seen no specimens to verify this either, although the
occurrence of the species in several counties in the Missouri Ozarks
makes it seem likely.

4. C. TUBEROSA Michaux, Fl. Bor.-Am. 1: 17. 1803. “In umbrosis
humidisque sylvarum Carolinae.” Phototype, GH. — As noted just above,
Gray misapplied the name C. scabriuscula Aiton (properly a synonym
of C. serotina) to this species. C. praecox Walter is possibly but by no
means certainly the same (see remarks under doubtful or excluded
names at end).

This is really much more distinct than may appear from the key. The
seemingly inconclusive feature of leaf pubescence was inserted to help
separate this from C. serotina, whose geographic range it shares in large
part.

Flowering September. ALABAMA. Blount, Tuscaloosa. GEORGIA.
eel LOUISIANA. East Feliciana, Orleans. MISSISSIPPI. Harri-
H

LIN Berkeley, Calhoun, Chester, Dorchester, Orangeburg, Sumter.
TENNESSEE. Shelby

DOUBTFUL OR EXCLUDED NAMES

The following list does not include the very numerous misapplica-
tions of names, of which the two most noteworthy have been mentioned
in the text (C. canadensis var. punctata for forms of C. canadensis in-
stead of C. serotina; C. scabriuscula for C. tuberosa instead of C. sero-
tina). The precise disposition of most of these names could probably be
determined. Since in all cases but one there are older valid names for
the species to which they might possibly belong, their identity is of
academic interest only. I prefer to devote my inadequate time to mat-
ters that really require it.

In addition to the following, there are five nomina nuda listed in
Merrill’s Index Rafinesquianus (C. bicolor, C. cordata, C. grandiflora,
C. heterophyla, C. longiflora) which need not be considered. From the
type localities, the first four are to be referred to C. canadensis

on ATA var. MAJOR Bentham in DC., Prodr. io 254, ee Abo
Georgia (Nutt.!) et Florida (herb. Hook.!).” The Georgia specimen may

2° 249, 1831. Cited by Bentham as doubtful synonyn of C. canadensis.
I have not seen the original description. All recognized species of the
genus have older names.

C. DECUSSATA Moench, Meth. p. 379. 1794. Said by Bentham and
Gray to be a synonym of C. canadensis.

C. HETEROPHYLLA Graham. Merrill (Index Raf. p. 206) states that
C. bicolor Rafinesque ex M’Murtrie is probably this. I can find no other
mention of such a binomial.

82

C. PRAECOX Walter, Fl. Carol. p. 65. 1788. This may be C. tuberosa,
but it also may be a form of C. serotina. No specimen is preserved, and
the brief description is inconclusive. Because only C. tuberosa and C.
serotina occur in Walter’s home area, it is extremely tempting to equate
C. praecox with C. tuberosa. The comment that it has flowers both
terminal and lateral would certainly apply to normal forms of C
tuberosa, but C. serotina may rarely have lateral flowering branches
also. As Gray noted, C. tuberosa is not early-flowering. In the interest
of nomenclatural mite I believe that C. praecox Walter should not
displace C. tubero

C, PURPUREA Goi ex Elliott, Sketch Bot. S.C. & Ga. 1: 35. 1817.
Appears only as a synonym of C. scabra (i.e., C. serotina).

C. SCABRA “Persoon,” Syn. Pl. 1: 29, 1805, and C. SCABRA “Elliott,”
Sketch Bot. S.C. & Ga. 1: 35, 1817. Both authors credit the name to
Aiton, so that these are simple errors for C. scabriuscula and have no
nomenclatural standing. But C. scabra Pursh was apparently a deliber-
ate substitute name (see under C. serotina).

CIFOLIA Salisbury, Prodr. p. 75. 1796. Listed in Index
Kewensis as a synonym of C. canadensis.

C. VERTICILLATA var. PURPURASCENS Elliott, Sketch Bot. S.C.
& Ga. 1: 37. 1817. “Near Crooked river bridge, Camden county, Georgia.”
The species occurs in the interior. The unusual specimen of C. serotina
from Decatur County, Georgia, noted in the text under that species,
makes me wonder if Elliott did not have something like it (with
simple inflorescence and a few leaves, stimulating C. verticillata). Cam-
den County is on the coast in extreme southeastern Georgia, where C.
serotina rather than C. verticillata is to be expected. According to
Weatherby, no specimen is preserved in the Elliott Herbarium in
Charleston.

REFERENCES

ARTRAM, WILLIAM (ed. Francis Harper). 1958. The Travels (Naturalist’s edition).
rans pp. 260—261 (orig. 411-412), 477.
BENTHAM, G. 1848. Collinsonia. In DC., Prodr. 12:
BRIQUET, J. 1897. Labiatae-Stachyoideae- Perillinaes iio pdictla. Coleone: In Engler &
ao Nat. Pflanzenfam. Teil IV. Abt. 3a: 325—32
RNALD, MERRITT LYNDON. 1950. oe Micheliella. Gray’s Man. Bot. (8th
ay pp. ee

FERN M. L. AND BERNICE G. SCHUBERT. 1948. Studies of American types
in ae Tae Sori e species of Thomas Walter. ce esciae ee serotina. Rhodora 50: 223.
Sr oe HENRY A. 1952. Collinsonia, New Britton & Brown Ill. Flora 3: 189.

pe ig Syn. Fl. N.A. 2 pt. i. DES

Bue pres NEV er ore GEORGE DAMON FULLER. 1955. Collinsonia
sgicaisele Vasc. Pl. i. p.
MERRILL, ELMER D. rs Place Diallosteira, Pleuradenia. Index Raf. pp. 206,

09

MOHR, CHARLES. 1901. Collinsonia. Plant Life of Alabama (Contrib. U.S. Nat. Herb.
6) pp. 15, 696—697

PALMER, ERNEST J, AND JULIAN A. STEYERMARK. 1935. An annotated cata-
logue of the flowering plants of Missouri, Collinsonia canadensis. Ann. Mo. Bot. Garden

83

SMALL, ee KUNKEL. 1903 (2nd ed. 1913). Micheliella, Collinsonia. Fl. S.E. U.S.
pp. 1050—

933, ne: eS Man. S.E. Fl. pp. 1178—
WEATHERBY, a A. specimens in Elliott’s herbarium. Calmont
Rhodora 44: 254 a. ee aa

SYNOPSIS OF CONRADINA (LABIATAE)
LLOYD H. SHINNERS

The account of the small Southeastern genus Conradina in Small’s
Manual of the Southeastern Flora illustrates several of that author’s
shortcomings. Four species are keyed and named; an anomalous plant
is briefly described in addition. Of the four named species, C. puberula
Small is merely a form of C. canescens, in fact the same form that
Gray considered typical of the latter; the types came from nearly the
same locality. C. montana, described as new, had been named earlier
in the same year as C. verticillata Jennison. Small knew almost nine
months before publishing that the other name was already in press
(etter from Jennison to Small, dated 10 March 1933, accompanied by

to prepare his floras solely on the basis of what he had at New York,
ignoring important collections at Cambridge, Washington, and St. Louis.
Recently activity in building up herbarium collections in the South-
ern States underscores the limitations of the older centers. Of the two
new species of Conradina described in this synopsis, New York has
no material at all of one (of which the Gray Herbarium has only a
sterile specimen), and no normal flowering material of the other
(which is represented at New York and the Gray Herbarium only by
duplicates of the same collection).

The genus Conradina comprises small shrubs with entire leaves which
have dense, appressed or matted pubescence on the lower surface, In
other woody Labiatae of the Southeast which without flowers might be
mistaken for Conradina, the leaves are glabrous or have spreading hairs.
The chief generic peculiarity is in the corolla tube, which is abruptly
bent up above the middle, rather than straight or gradually curved.
What benefit this may be to the plant is difficult to imagine. Add to this
the peculiarity that each of the species occupies a geographic area en-
tirely separate from all the others, one of them quite remote from the
rest (C. verticillata on the Interior Low Plateau, the others on the outer
Coastal Plain), and we have a rich subject for useless speculation that
the phylogenists have so far overlooked.

In preparing this synopsis I have had the use of collections at Florida
State University, the University of Florida, the Gray Herbarium, the
New York Botanical Garden, the University of North Carolina, and
Southern Methodist University. My thanks are extended to the various
curators for their help.

SIDA 1 (2): 84—88. 1962.

85

CONRADINA Gray, Proc. Amer. Acad. 8: 294—295. 1870 (Dec. 31).
C. canescens (T. & G.) Gray, the only original species, is automatically
the type.

KEY TO THE SPECIES

la. Calyx tube glabrous or very minutely and inconspicuously puber-

SCENUOUTSIC Cl erase ans tanita oi a Wea Renae gh ae 1. C. glabra
lb. Calyx tube densely short-pubescent or both pubescent and pilose
outside

2a. Calyx tube densely short-pubescent and also pilose with moderately
long (0.3—0.8 mm.) gland-tipped hairs (the teeth with longer
hispid hairs); stem or main branches partly decumbent and root-
ing; plants of Kentucky and Tennessee, flowering May—early
DUNC pee eerie Gone wo. A 2 ce ee . C. verticillata

2b. Calyx tube densely short-pubescent, often also pilose with long
(mostly 1—2 mm.) glandless hairs like those on the teeth, rarely
also short-pilose with gland-tipped hairs less than half as long

as the glandless ones; stems erect or short-decumbent, not freely
rooting; plants of Florida and coastal Alabama, flowering all year

3a. Lower lip of corolla 4—9 mm. long; leaf blades with midrib
moderately to very densely pubescent beneath, the inrolled
margins usually concealing all or nearly all the surface of the
blade; peduncles absent or very short, each axil with 1—6

4a. Larger leaves on well-developed flowering branches 7—20
mm. long, mostly equalling or exceeding the internodes;
middle lobe of lower lip of corolla 3.2—5.5 mm. long; flowers

1—3 per axil; coastal Alabama and northwestern Florida
3. C. canescens
4b. Larger leaves on well-developed flowering branches 6.0—8.2
mm. long, mostly shorter than their internodes; middle lobe
of lower lp of corolla 2—4 mm. long; flowers 1—6 per axil;
interior peninsula Florida (Highlands Co.)....4. C. brevifolia
3b. Lower lip of corolla 9—16 mm. long; leaf blades with midrib
glabrous to moderately densely pubescent beneath, contrasting
with the very densely pubescent surface of the blade (latter
usually only partly concealed by the inrolled margin); peduncles
evident, with 1—12 flowers each.............. 0. C. grandiflora
1. C. glabra Shinners, sp. nov. Frutex parva ramosa maxima parte
glabra. Folia sessilia vel subsessilia subtus cana ramulorum juniorum
lineari-oblanceolata 18—23 mm. long 1.6—3.0 mm. lata subacuta, ramu-
lorum veterum minora sublinearia obtusa. Pedunculi brevissimi pub-
escentes 1—3-flori. Pedicelli pubescentes calycis longitudinem 1/3—2/3
attingentes. Calyx 6.2—7.0 mm. longus, tube glabra vel minutissime
puberula, dentes ciliati, faux hispido-pilosa. Corolla extus pilosula
12—15 mm. longa. HOLOTYPE (with normal stamens): steep, sandy,

86

wooded bluff 14% miles west of junction of Rt. 12 and road to Torreya
State Park, Liberty Co., Florida, F. H. Sargent 6219, 29 April 1952 (SMU).
PARATYPES (some with aborted anthers, as noted) (all from Florida):
Gadsden or Liberty Co.: between Bristol and Greensboro, FE. West &
H. H. Hume, 28 March 1936 (FLAS, 2 sheets with normal anthers, 1 with
aborted anthers, the filament tips enlarged, malformed). Liberty Co.:

irregularly expanded and ae ). Common, along

bayhead, 7 miles south of Wilma, A. A. Will, 1 April 1961 (FLAS,
anthers small or absent). Santa Rosa Co.: dry sand, edge of Swamp, Rt.
90, at roadside park, west of Milton, S. C. Hood 1868, 8 April 1949
(FLAS). County not known: one twig at bottom center of mixed sheet
with fragmentary specimens of C. canescens, from Herb. John A. Lowell
(“Transferred from the Boston Society of Natural History to the Gray
Herbarium... Oct. 2, 1941”), data uncertain (there are several labels
plus pencilled notes, but it could not be determined which are to be
associated with this particular fragment) (GH).

Much branched shrub under 1 m. tall, largely glabrous. Leaves sub-
sessile or sessile, closely canescent beneath, those of young shoots
linear-oblanceolate, 18—23 mm. long, 1.6—3.0 mm. wide, subacute, with
narrowly revolute margins; those on older growth almost linear, 7—
mm. long, obtuse, with strongly revolute margins. Peduncles very short,
minutely pubescent, 1—3 flowered. Pedicels about 1/3—2/3 as long as
the calyx, minutely pubescent. Calyx 6.2—7.0 mm. long, resin-dotted,
glabrous or very minutely and inconspicuously pubescent except for
the hispid-pilose throat and margins of the teeth. Corolla pilosulous out-
side, 12—-15 mm. long (as pressed), the lower lip 4—7 mm. long; color
unknown but apparently very pale lavender or white.

This is the anomalous plant briefly noted by Small, but not named.
Because the Palmer specimens (the only ones seen by Small) do not
show normal anthers, they are not suitable for designation as holotype
and isotypes

2. C, VERTICIL LLATA anes Journ. Elisha Mitchell Sci. Soc. 48:
268—269. 1933 (April). LOTYPE: on sandy beach, north bank of
the Clearfolk River near a Fentress Co., Tennessee, Jennison &
Sharp 3-432, 16 May 1931 (TENN., presumably destroyed in the 1934
fire; a isotype, lacking number but with other label data the
same, GH). C. montana Small, Man. S.E. Fl. p. 1167. 1933 (after Nov.
30). paar near Rugby, Tennessee (NY, not seen; sterile topo-
type, Mrs. Ferriss (Herb. Albert Ruth), July, 1903, NY). Corolla said
to be lavender in the original description, pink on the only specimen
seen bearing color data (Braun s.n., cited below). The following collec-
tions have been examined.

KENTUCKY. McCreary Co.: river bank, South Fork Cumberland
River, E. Lucy Braun s.n., 18 June 1935 (GH). (Also sterile specimens,

87

same locality and collector, GH, NY.) TENNESSEE. Fentress Co.: Rugby,
sandy soil, Albert Ruth, 27 June 1894 (sterile) (GH). About 1 mile
north of Rugby, in sand on beach along north bank of Clear Fork River,
Jennison 33-124, 28 May 1933 (NY; apparent duplicate but lacking
number, GH). Sandy beach along Clear Fork River near Rugby, Jenni-
son 1111, 5 May 1934 (NY, SMU). Morgan Co.: in sand banks of Clear
Fork of Cumberland River, Rugby, Jennison s.n., 4 June 1931 (GH).
Sandy bars along stream and gravelly slopes, Rugby, H. K. Svenson
4085, 19 Aug. 1930 (sterile) (GH). Scott Co.: mouth of No Business Creek
on South Fork River, rocky and sandy soil of Big Island, A. J. Sharp,
R. E. Shanks, E, Clebsch 3835, 20 June 1947 (flowers past) (NY). Also
CULTIVATED, TENNESSEE, Knox Co.: Sanford Arboretum, Knox-
ville, A. F. Sanford, 10 May 1935 (GH).

3. C. CANESCENS (T. & G.) Gray, Proc. Amer. Acad. 8: 295. 1870.
Calamintha canescens T. & G. ex Bentham in DC., Prodr, 12: 229. 1848.
“In Florida ad Tampa Bay (h. Gray!) ad Apalachicola (Drumm.!).”
For reasons given below, the cited specimens have been disregarded
and a lectotype designated in the Gray Herbarium, on a mixed sheet, at
left, with blue label “Calamintha canescens Fla.” on which has been
added “T. & Gr.” in Gray’s hand-writing, and above which ‘‘

“Florida: Apalachicola, old specimen, collector uncertain; later speci-
men, A. H. Curtiss, no. 2014. Also two specimens collected by Dr. Chap-
man.” (Curtiss 2014 and one Chapman specimen from NY examined.)

freely in late winter (February). Common in sand along the coast from
Mobile and Baldwin counties, Alabama, east to Franklin County, Florida;
rarely slightly inland (in Okaloosa and Walton counties, Florida). There
is a decidedly unscientific local legend that this is the true rosemary
of Europe (Rosmarinus officinalis), brought to Florida by early British
settlers and naturalized (clipping from St. Andrews Bay News, 8 April
1924, filled with specimen, NY; see also Hepburn, 1956, p. 7, under
Panama City).

Neither this species nor C. grandiflora (which for long was not
separated from it) occurs about or near Tampa Bay, and there is no
material in the Gray Herbarium so designated. I suspect some mixup
or error of labelling, and therefore consider the first collection cited by
Bentham unsuitable for lectotype. It is uncertain whether Torrey and
Gray (particularly Gray, who usually handled the Sympetalae) saw
the Drummond material before naming the species. There is some indi-
cation that they did not until later. Several labels in the Gray Herbar-
ium read “Keithia sp. Bentham,” suggesting that Torrey and Gray had
provided the specific epithet first, and that Bentham’s acceptance of

88

it came later. Hence Bentham’s second collection is Bae over

This creates no problem, fortunately, for Drummond 23 (GH) a et
to the very same form and is from about the same locality as the desig-
nated lectotype. I have selected from the material at the Gray Her-
barium the sample which from the nature of the label appears to be
oldest, and which conforms to what Gray in his 1870 description indi-
cated as the typical form, the one with calyx tube short-pubescent ae
(“calyce . .. dentibus (rarius tubo) pilis patentissimis hirsutis”’). Thi

is ides aah what Small later described as the alleged species .
puberula. The two are merely genetic forms of one, and are not geo-
graphically segregated. My no. 29,264 from 10 miles south of Foley,
Baldwin Co., Alabama, 7 April 1961 (SMU), has two branches from.
different plants, one showing calyx short-pubescent only, the other both
short-pubescent and pilose.

4. C. brevifolia Shinners, sp. nov. C. canescenti persimilis, foliis
brevioribus plerumque remotis cum fasciculis axillaribus quasi verticil-
latis; floribus ad 6 in quaque axilla, florum labio inferiore minus pro-
funde diviso (loba media 2—4 mm. longa). HOLOTYPE: scrub, south-
west of Avon Park, Highlands Co., Florida, Ray Garrett 41, 16 Feb.
1948 (FLAS, acc. no. 50231). PARATYPES (also Highlands Co.): Clay
pit, Avon Park, J. B. McFarlin 10135, 6 Feb. 1935 (FLAS, acc. no. 49469).
In very sandy soil along road 567 about 7 miles east of Avon Park,
Chas. C. Dean 64200, 13 Feb. 1946 (NCU). — In additiont to these, a sterile
specimen from the same county at first thought to be C. grandiflora
doubtless is to be referred to C. brevifolia instead (the loan had been
returned before I saw flowering material of the new species and the
specimen has not been reexamined): in the scrub north of the Botanic

Ey);

FLORA Small, Bull. Torr. Bot. Club 51: 386—387, 1924.
“The type specimens were collected by the writer on the ancient sand-
dunes near Sebastian, Florida, April, 1921.” (Topotype specimens col-
lected by Small et al., Indian River Co., FLAS, GH, NY, NCU.)
Corolla color (noted on only a few specimens) lavender, pinkish, or
pink-purple. Confined to eastern peninsula Florida, from Volusia County
south to Dade County.

REFERENCES
HEPBURN, ae aaa 1956. Complete Guide to Florida. Houghton-Mifflin American
Travel Series No.
SMALL, JOHN KUNKEL. 1933. Conradina. Man. S.E. Fl. pp. 1166—1167.

SYNOPSIS OF DICERANDRA (LABIATAE)
LLOYD H. SHINNERS

Like Conradina, Dicerandra is a small genus confined to the South-
eastern United States. Long known to comprise three annual species,
it must now be redefined to include a woody-based perennial. The
primary generic character is the presence of a spur-like or horn-like
appendage on the anther sacs. All the known species are also character-
ized by being largely glabrous and having the calyx white or roseate
at the mouth. All are typically fall-blooming (but see special notes on
numbers 1 and 3). The summit of the taproot and base of stem are
often swollen, presumably the effect of some parasitic organism, but
the cause is not known.

Grateful acknowledgment is made for the use of collections from
Florida State University, the University of Florida, the Gray Herbarium,
the New York Botanical Garden (one sheet only, received unidentified
with loan of Conradina), the University of North Carolina, and South-
ern Methodist University.

DICERANDRA Bentham, Bot. Reg. 15: t.1300. 1830. (Not seen; taken

from Bentham
Elliott, Sketch Bot. S.C. & Ga. 2: 1822. (Not Ceranthera Beauvois,
1807, nor Rafinesque, 1818.) Elliott’s only species, C. linearifolia, is
automatically the type.
KEY TO THE SPECIES
la. Stamens prominently exserted; lower lip of calyx 1.5—3.0 mm. long,
divided %4—% into two wide-based, acuminate teeth
2a. Perennial, woody below, with numerous erect to ascending
branches from near base; corolla white or yellowish white with
ULE CLO LS eon ten hc Shy e Adak eg 1. D. frutescens
2b. Annual, unbranched or with few spreading to ascending branches
well above base; corolla pale to deep lavender or reddish purple
with darker do
3a. Peduncles (except at uppermost nodes) evident, usually well-
developed, few-flowered, the inflorescence rather loose; anther
WGOGNSBACUIMIN ALC Meee ll cco ts. «ncaa coc iE . D. linearifolia
Peduncles absent or very short, the flowers numerous and
crowded in the axils of leaves or leafy bracts; anther horns
Obluseworsbarely aCuten coe hae clowi eae oe 3. D. densiflora
lb. Stamens included; lower lip of calyx 2.5—4.0 mm. long, divided
nearly to base into two narrow, long-acuminate teeth
4. D. odoratissima
' 1. D. frutescens Shinners, sp. nov. Suffrutex parvula ad 50 cm. alta
glabra nisi ad nodos superiores, pedunculos, pedicellos, calyces (basin

Se)
CF

SIDA 1 (2): 89—91. 1962.

90

versus solum) corollas (extus solum). Folia sessilia vel subsessilia,
oblongo-linearia (in novellis linearia), integerrima, obtusa, subcoriacea,
punctata, inferiora 12—17 mm. longa 1.8—2.6 mm. lata, superiora minora.
Pedunculi breves vel subnulli bracteati uniflori. Pedicelli dimidium

limbo albido. Corolla extus pilosula alba vel luteola labio inferiore
rubro-punctato; tuba apicem versus reflexa modo generis Conradinae.
Stamina exserta; antherae rubro-purpureae valde aristatae. Stylus ex-
sertus pilosulus bifidus. HOLOTYPE: sandy scrub, 20 miles south of

1945 (GH). “Gregarious locally in sandscrub; aromatic perennial about
90 cm. high; calyx lobes white; corolla white or yellow, purple-spotted.”
In scrub near Lake Placid, J. B. McFarlin 10559, 7 October 1936 (FLAS,
GH). Route 8-A, Hicoria, Brass 14646, 16 February 1945 (half-dead
stems with new sprouts, both leafy and flowering) (GH). “Very abun-
dant on roadsides in sandscrub; bushy, woody herb to 50 cm.; aromatic;
flowers white, prominently marked with purple.” Sandhills near Lake
Stearns, John K. Small & Paul Matthaus 11613, 8 January 1925 (long
past flowering) (NY). PARATYPES from Sumter Co., Florida: sandy
scrub, 5 miles west of Wildwood, E. West, 22 September 1938 (FLAS).
Dry sandy road bank 7 miles west of Wildwood, West & Arnold, 8
November 1946 (in fruit) (FLAS).

The Small & Matthaus collection had been labelled Dicerandra, but
later filed with Conradina, and was received in the loan of material
of that genus from New York. The glabrous leaves made Conradina
seem unlikely, and in the course of working over Dicerandra, an
anomalous specimen received as Ceranthera linearifolia (now the holo-
type of the new species) revealed its true identity. The long-exserted
stamens with conspicuously horned anthers, the white-lipped calyx,
and the glabrous leaves are all characteristic of Dicerandra. The shrubby
habit and the bent corolla-tube suggest Conradina, but hardly call for
a merger of the two genera. D. frutescens adds still another to the many
restricted endemics of the Florida Lake Region. .

2. D. LINEARIFOLIA (Elliott) Bentham, Bot. Reg. 15: t. 1300. 1830.
Ceranthera linearifolia Elliott, Sketch Bot. S.C. & Ga. 2: 94. 1822. “Grows
abundantly in the high pine barren ridges between the Flint and Chata-
houchie rivers” (in extreme southwestern Georgia, where it is re-
ported as “frequent” by Thorne, 1954). “Flowers September and Octo-

Bentham, Lab. Gen. et Sp. p. 413. 1832— . (Not seen. Apparently
only a slp of the pen; Bentham a oe this as synonym of D.
linearifolia in DC., Prodr. 12: 243 (1848.)

This is the ost widespread and most frequently collected member
of the genus. The rather outlying record cited below from Palm Beach

91

Co., Florida, is based on an A. H. Curtiss collection (without number)
dated 20 May 1895 (GH). I suspect an error in labelling; both locality
and date are questionable. Coastal Alabama through northern Florida
to southern Georgia. Specimens seen from the following states and
counties. ALABAMA. Baldwin. FLORIDA. Alachua, Duval, Escambia,
Gadsden, Holmes, Jackson, Leon, Liberty, Madison, Okaloosa, Palm
Beach, Santa Rosa, Taylor, Wakulla, Walton, Washington. GEORGIA.
Baker, Berrien, Jenkins, Taylor.

D. DENSIFLORA Bentham in DC., Prodr. 12: 243. 1848. “In Florida
orientali (h. Torr.!)” (specimen not seen). Ceranthera densiflora

ntham) Gray, Syn. Fl. N.A. 2 pt. 1: 365. 1878. Northern peninsular
Florida; doubtfully South Carolina (see remarks below), and said by
Small (1933) to extend into Georgia. FLORIDA. Alachua, Columbia,
Dixie, Gilchrist, Lafayette, mate Suwanee, Volusia, SOUTH CARO-
LINA. Locality not given, Curt “ex Herb. H. P. Sartwell” (GH).
Possibly an error in labelling. aa the Jasper County record of Ahles
et al., see under next species.

4. D. ODORATISSIMA Harper, Bull. Torr. Bot. Club 28: 479. 1901.
TYPE: dry sand-hills near Seventeen-Mile Creek, Coffee Co., Georgia,
Roland M. Harper 695 (isotype, GH). Additional specimens seen from
Berrien Co., Georgia, and Jasper Co., South Carolina. The latter were
reported as D. densiflora by Ahles, Bell and Radford (1958); they differ
from the others in having calyx roseate at the mouth instead of white.
The corolla in this species is white.

REFERENCES
AHLES, HARRY E., C. RITCHIE BELL, AND A RT E. RADFORD. ne aoe
new to the flora of North or South Carolina. ee ae Rhodora 6
BENTHAM, G. ea Dicerandra. In DC., eer ee ete
SMALL, JOHN KUNKEL. 1933. Dicerandra.

THORNE, ROBERT F. 1954. The vascular Sree of Taueeee brs ‘Dicerandva
linearifolia. Amer. Mid]. Nat. 52: 31

W HERBY, C. A. 1942. A aes of type specimens in Elliott’s herbarium. Rhodora 44:
249—262. (Contrib. Gray Herb. 141.)

VEGETATIVE KEY TO WOODY LABIATAE OF
THE SOUTHEASTERN COASTAL PLAIN
LLOYD H. SHINNERS

Conradina is distinguished technically by the bent corolla tube,
Dicerandra by the appendaged anthers, Trichostema by the abbreviated
upper corolla lip. Without flowers it is hardly possible to key them. In
the course of working up Conradina, Dicerandra, and Calamintha, a
number of sterile or fruiting specimens were encountered, most often
guessed to be Conradina. Actually all the woody species can be readily
determined by vegetative characters or a combination of vegetative
characters and geography, as shown in the following key.

la. Lower surface of leaf blades densely gray-pubescent with appressed
or matted (very rarely loose or somewhat spreading) hairs (midrib
glabrous or glabrate in some species, remainder of lower surface
often partly or wholly concealed by inrolled margins)
2a. Midrib densely gray-pubescent beneath
3a. Plants of coastal Alabama and northwestern Florida
Conradina canescens
3b. Plants of interior peninsular Florida...... Conradina brevifolia
2b. Midrib glabrous to moderately densely pubescent beneath, con-
trasting with the very densely pubescent surface of the blade
4a. Plants of northwestern Florida............... Conradina glabra
4b. Plants of eastern peninsular Florida...... Conradina grandiflora
lb. Lower surface of leaf blades glabrous or finely pubescent with erect
or spreading hairs or sparingly hispid-pilose
5a. Upper surface of leaf blades sparself to moderately densely hispid
with rather long hairs from swollen bases (with or without very
SHOVT Maire BS Wel) sea os 2cckd deo ee cena ees Piloblephis rigida
5b. Upper surface of leaf blades glabrous or minutely pubescent
6a. Stem and branches glabrous except at nodes
Dicerandra frutescens
6b. Stem (at least younger parts) and branches densely and minute-
ly pubescent
7a. Hairs curled or bent upward or downward (sometimes short,
straight, glandular hairs also present)
8a. Hairs stiff, straight, very short; leaf blades obtuse to
rounded or slightly indented at apex, the larger on petioles
less than 2 mm. long; central and southern peninsular
TORIOR- 4 hob Gudea ae ered oo taee es Trichostema suffrutescens
8b. Hairs loosely curled or wavy; leaf blades obtuse to subacute,

SIDA’ 1. (2)= 9293: 1962:

93

the larger on petioles up to 10 mm. long, northern Florida
to southeastern Louisiana and South Carolina
Calamintha georgiana
7b. Hairs spreading at right angles, straight
9a. Leaf blades widest about at middle (linear to oblong-
Calamintha Ashei

anceolaten easy eet aie oe eee ieee
9b. Leaf blades widest above middle (oblong-oblanceolate to
obovate)
10a. Internodes of twigs uniformly pubescent with very short

ai

10b. Internodes of twigs with mixed pubescence: densely cov-

ered with very short glandular hairs (usually of uneven

lengths), sparsely pilose with long glandless hairs, at least
near nodes; larger leaves toothed above middle

Calamintha dentata

REFERENCES
LEWIS, HARLAN. 1945. A revision of the genus Trichostenta. Brittonia 5: 276—303.
(T. suffrutescens Kearney, p. 300.
SHINNERS, LLOYD H. 1962a. an (Labiatae) in the Southern United States.
Sida 1: 69-75. (Including nomenclatural note on Piloblepbis rigida (Bartram) Rafinesque,
p. 70.)

. 1962b. Synopsis of Conradina (Labiatae). Sida 1: 84-88.
. 1962c. Synopsis of Dicerandra (Labiatae). Sida 1: 89-91.

MICROMERIA BROWNEI AND
ITS ALLIES (LABIATAE)
LLOYD H. SHINNERS

In 1903, Small recognized two species of Micromeria in the Southern
United States: M. pilosiuscula in Texas (and Mexico) and M. Brownei
in Florida (and the West Indies). In 1933, he recognized only one for
the continental United States, the Florida plants being placed with the
Texas ones under M. pilosiuscula. In attempting to evaluate Small’s
conclusions, I found that some West Indian material (from, the Bahamas
and Hispaniola) is indeed very distinct from pilosiuscula, but the true
M. Brownei (from Jamaica) does not differ significantly except in
average size and geography. Some future monographer may prefer to
establish a hierarchy of subspecies and varieties to accommodate all of
these under one species name. I prefer to recognize separate, though
admittedly closely related, species, one of them with three varieties.

y immediate purpose has been merely to settle the nomenclature of
the plant of the United States. As an interesting sidelight, it is possible
to point out that in Florida it represents not a primarily West Indian
element (it does not occur in the southern part of the state, and the
plants of the Bahamas and Cuba are not the same), but a Texano-
Mexican one,

All members of the group are small, commonly mat-forming weeds
of damp places, with lavender or partly white corolla. This brief ac-
count is based on collections at Florida State University, the University
of Florida, the Gray Herbarium (U.S, and West Indian only), the New
York Botanical Garden, and Southern Methodist University, to whose
several curators grateful acknowledgement is made.

KEY TO SPECIES AND VARIETIES
la. Shortest calyx teeth 1/5—1/6 as long as entire calyx; margins of
calyx teeth either glabrous or hispid-ciliate; larger leaf blades
—20 mm. wide; plants of Cuba, Jamaica, Guatemala, Mexico, and
the United States

2a. Margins of calyx teeth glabrous or rarely (one Jamaican collec-

tion) inconspicuously ciliate with hairs about 0.1 mm. lon
3a. Plants of Jamaica; larger leaf blades 5—12 mm. wide, glabrous;
pedicels 3—7 mm. long............ la. M. Brownei var. Brownei
3b. Plants of continental North America; larger leaf blades up
to 20 mm. wide, glabrous or oe pedicels 3—15 mm. long
Brownei var. pilosiuscula
2b. Margins of calyx teeth esi with hairs 0.3—0.6 mm. long;
plants of Cuba (rare) and Yucatan....1le. M. Brownei var. ludens

SIDA 1 (2): 94—97. 1962.

95

lb. Shortest calyx teeth 1/3 as long as entire calyx; margins of calyx
teeth prominently hispid-ciliate; larger leaf blades 4—6 (rarely

— 11) mm. wide; plants of the Bahamas and Hispaniola
4a. Pedicels (except uppermost ones) 1.0—3.5 mm. long, shorter than
the adjacent petioles; marginal hairs of calyx teeth spreading;
JSR obs aalsteheng ec rig Gee oes ea thie ae ec 2. M. bahamensis
4b. Pedicels 3.0—6.5 mm. long, longer than the adjacent petioles;
marginal hairs of calyx teeth incurved-ascending, subappressed;

3

A'S TT Oe es ee Me A as GON ee: Se he prone ena omingensis

1. M. BROWNEI (Swartz) Bentham, Lab. et Sp. p. 372. 1832—
1836. (Not seen; quoted in DC., Prodr. 12; 223. es Thymus Brownei
Swartz, Pro p 1788. Type loealiey Jamaica. Satureia Brownei

(Swartz) Briquet in Engler & Prantl, Nat. Pflanzenfam. Teil IV. Abt.
3a: 300. 1897. (As “S. Brownii,’ but this was not the spelling adopted
by Swartz.)

la. M. BROWNEI var BROWNEI. Confined to Jamaica, at various
altitudes, from 200 feet up to 2200 feet; 13 sheets examined (FLAS, GH,
NY). All have glabrous calyx teeth except the following, on which
these are more or less short-hispid-ciliate. Bluefields and vicinity; on
wall, Bluefields Mountain, 700 m. alt., N. L. Britton & Arthur Hollick
1994, 6—7 March 1908 (NY). Sat eie collections dated January, Feb-
ruary, March, July, August, Septem

lb. M. BROWNEI var. eres Gray, Syn. Fl. N.A. 2 pt.
1: 359. 1878. Type locality: near San Antonio, Bexar Co., Texas. Satureia
Brownei var. pilosiuscula (Gray) Briquet in Engler & Prantl, Nat.
Pflanzenfam. Teil IV. Abt. 3a: 300. 1897. Micromeria pilosiuscula (Gray)
Small, Fl. S.E. U.S. pp. 1042, 1337. 1903. Thymus xalapensis H.B.K.,
Nov. Gen. (4to ed.) 2: 316—317. 1818. “In declivitate montium Mexi-
canorum soli oriente obversorum inter villam Pileta et urbem Xalapa,
alt. 760 hex.” (Vera Cruz, Mexico). Micromeria xalapensis (H.B.K.)
Bentham, Lab. Gen. et Sp. p. 372. 1832—1836. (Not seen; quoted in DC.,
Prodr. 12: 223. 1848.)

Very variable as to stature, dwarf plants with glabrous leaves being
hardly distinguishable from var. Brownei except by geography. Pubes-
cent-leaved forms (intergrading with glabrous ones) occur in Texas
and Mexico. At the northeastern extremity of its range (Florida, Georgia,
Louisiana) only the glabrous form is found. This is evidently the end-
product of selective migration from the west and southwest. Since var.
pilosiuscula does not occur in the West Indies, and since it is absent
from the southern part of Florida (where West Indian plants steel
normally be expected), it must be regarded as an unusual Tex
Mexican element in the flora of Florida. The general ee eee of
this variety, on the basis of specimens examined, is summarized by
country and department or state, and for the cae States by county.
GUATEMALA. Alta Verapaz, Huehuetenango, Quiche. MEXICO, Jalisco,
Nuevo Leon, San Luis Potosi, Sinaloa, Tabasco, Tamaulipas, Vera Cruz.

96

UNITED STATES. FLORIDA. Alachua, Bradford, Brevard, Clay, Flag-
ler, Gadsden, Gilchrist, Hillsborough, Jackson, Lake, Levy, Orange,
Osceola, St. Johns, Seminole, Volusia. GEORGIA. Decatur. LOUISIANA.
St. Bernard. TEXAS. Aransas, Bexar, Brazoria, Cameron, Colorado. In
flower in the United States from February to September, farther south
apparently nearly all year.

le. M. BROWNEI var. ludens Shinners, var. nov. A var. Brownei
recedit calycis dentibus hispido-ciliatis. HOLOTYPE: wayside, near
Sabicu, Rangel, Sierra del Rosario, Pinar del Rio Province, Cuba, Bro.
Alain 6137, January, 1957 (NY). PARATYPES (all NY): Cultivated, Est.
Agronomica, Santiago de Vegas, Habana, Cuba, J. Acuna 19539, 5 Feb.
1955. Near Merida, Yucatan, Mexico, Porfirio Valdez 35, in 1896. Lo-
cality illegible (“Coym. ... Is.”?), Yucatan, Mexico, Gaumer 109, ex
Herb. Gray. The disjunct distribution between Cuba and Mexico sug-
gests that of Egletes viscosa, especially f. bipinnatifida (cf. Shinners,
1949)

2, M. bahamensis Shinners, sp. nov. M. Brownei affinis. Caulis ad vel
infra nodos pilosula. Folia perparva, petiolis puberulis, laminis glabris
vel (imprimis facie superiore) parce hispidulis varie subintegris vel
argute denticulatis, majoribus 4—6 mm. latis (rarissime 11 mm.). Pedi-
celli 1.0—3.5 mm. longi, plerumque petiolis breviores. Calyx 3.5—4.1
mm. longus profunde lobatus (dentibus minoribus calycis tertiam par-
tem aequantibus), dentes longe patenter hispido-ciliati. Corolla extus
pilosula. HOLOTYPE: Berry Islands, Lignum Vitae Cay, Bahama Is-
lands, N. L. Britton & C. F. Millspaugh 2302, 2 February 1905 (cn
isotype, NY). A total of nineteen sheets has been examined (GH, NY).
The species is apparently common and widespread in the Bahamas, in
flower December to May and in August. With a single exception the
specimens are remarkably uniform, with very small leaves. e ex-
ception, with blades up to 11 mm. wide, is Britton & Millspaugh 3114,
sink holes near Georgetown, Great Exuma, 22—28 February 1905 (NY).

3. M. domingensis Shinners, sp. nov. Praecedenti persimilis, differt
pedicellis 3.0—6.5 mm. longis petiolos excedentibus, calycis dentibus
subappresse hispido-ciliatis. HOLOTYPE: Civ. Santo Domingo, Cordil-
lera Central, prov. de Azua, San Juan, El Varanjo, edge of brook, c.
800 m., Hispaniola, E. L. Ekman (Mus. Bot. Stockholm Pl. Ind. Occ.
N:O H.13505), 8 September 1929 (GH).PARATYPE: Santo Domingo,
Cordillera Central, prov. de la Vega, Constanza, edge of thickets in
swamps, ca. 1200 m. E. L. Ekman (Mus. Bot. Stockholm Pl. Ind. Occ.
N:O H. 13915), 25 October iene (NY). Wiese are the only specimens
seen. The altitude and flowering dates suggest physiological differences
between this and M. bahamensis.

NAME OF DOUBTFUL APPLICATION

Micromeria stolonifera Bentham, Lab. Gen. et Sp. p. 371. 1832—1836.
(Not seen; quoted in DC., Prodr, 12: 223. 1848.) Type locality: Mexico

97

(‘Nova Hispania”), Mocino et Sesse. “A. M. Brownei differt caulibus
subfirmioribus acute tetragonis et foliis latioribus sessilibus, nec petiolo
limbum aequante donatis. Rami et folia Gratiolam referunt. An M. xala-

pensis var. ? (v. s. olim in herb. Lamb.)” I have seen no collections

bearing this name,
Although petiole-length varies in M.
leaves are never really sessile. I am unable to identify Bentham’s plant.

REFERENCES

SHINNERS, LLOYD ae ae Beer ee
ee) eee 12: 239— (E. viscosa

oar KUNKEL, S508 ee Fl.

1933. Micromeria. Man. S.E. FL. p. ge

nus Egletes Cassini north of South
rat hy

dt: bipinnatfie, pp. 244—
1042, 133

NOTES

WEED TRANSPORT IN ST. AUGUSTINE GRASS SOD IN TEXAS.
— St. Augustine grass (Stenotaphrum secundatum) is commonly recom-
mended for lawns because its dense growth is supposed to choke out
weeds. There is only limited truth in the claim. The grass is established
by plugs or sods rather than by seeding. A spectacular illustration of
the transport of weed seeds with the sod was provided by the land-
scaping of the new Science Information Center at Southern Methodist
University in Dallas, Texas. Built on a moderate slope on what was
originally rolling prairie of calcareous, black clay, it was supplied
with a small, enclosed courtyard on the west side. Because of the slope
of the land, it was necessary to remove earth to a depth of over six
feet at the north and almost three at the south. A small live oak
(Quercus virginiana) near the southwest corner was preserved by being
boxed in on a sort of island, but even here the top soil was removed.
The main part of the court was left completely bare, but almost within
days a few sprouts of Johnson grass (Sorghum halepense) appeared.
Whether these represented deep penetration of rhizomes of plants
growing in the area before excavation, or whether bits of rhizome or

ot known. The new

until planted. The pudding was dedicated the afternoon of 3 November
1961, and in characteristic Texas fashion, the landscape plantings were
all put in on the morning of the same day. For the courtyard, these
comprised chiefly 10-foot yaupon (Ilex vomitoria) against the retaining
wall at the north, and a red oak (perhaps Quercus Shumardii) and a
live oak (Q. virginiana) of similar size near the middle. Most of the area
was carpeted with small squares of St. Augustine grass sod. The en-
suing winter was exceptional for the number of severe freezing spells,
lasting several days each, with temperatures down to 17° F. several
times, and once as low as 8°. This is not ideal weather for St. Augustine
grass, and much of it died. In the spring the courtyard was a weed
patch gratifying to botanical eyes. In April and May, five weedy grasses
and nine miscellaneous weeds were found in bloom:

Poa annua Sisyrinchinm minus

Hordeum pusillun Cerastium brachypodum

Alopecurus Dw Sibara virginica

Agrostis byemalis Ranunculus muricatus

Cynodon oe Euphorbia spathulata
a os eins ds
Core

is
Senecio im eats
Krigia gracilis

In June Paspalum dilatatum (Dallis grass) flowered. Smilax Bona-
nox (cat brier) was present next to the red oak, but may have been

SIDA 1 (2): 98—99. 1962.

99

brought with it rather than in the sods. It is characteristic of the Texas
flora that of the 17 weedy species present (I consider St. Augustine grass
another weed; certainly in my own yard, intended for flowers, it is a
pest), all but four are native to Texas. The exceptions are Poa annua

Asian), Paspalum dilatatum (Dallis grass, South American), and Ran-
unculus muricatus (buttercup, European). The last-named was new
to Dallas County, the nearest previously known occurrence being in
Gregg County, 125 miles to the east. It is fairly common in the south-
eastern third of the state and in Louisiana. The precise source of the
sod could not be determined, but according to an employe of the
landscape company which handled the work, it probably came from
Houston, That city is about 270 airline miles south and slightly east of
Dallas. All the species mentioned are known to occur there. It may well
have been the source of the Dallas occurrence of Melochia pyramidata
mentioned in my Spring none of the Dallas—Fort Worth Area (1958,

>

100 miles farther north as a weed in St. Augustine grass lawn in
Highland Park, Dallas: in front of Christian Science Church, Shinners
13,358, 14 May 1952 (SMU). “A few small patches.” Still another ex-
ample of such weed transport is provided by a specimen of Sisyrinchium
minus from the campus of Stephen F. Austin State College in Nacog-
doches (R. L. Oliver 317, 3 May 1961; SMU), which is noted as “ap-
parently introduced in the San Augustine grass (from Brazos River
bottom, Houston area) which was set out in fall of 1960.” — Lloyd H.
Shinners.

EICHHORNIA AZUREA (PONTEDERIACEAE) IN THE TEXAS
COASTAL BEND: NEW TO THE UNITED STATES. — The common
water hyacinth, Eichhornia crassipes (Martius) Solms, with short, greatly
inflated petioles, was collected in Texas as early as 1903 by Reverchon
(east of La Porte, Harris County), and has been found as far north
as Dallas. The following is apparently the first United States record
of the related E. azurea (Swartz) Kunth, with elongate petioles thicker
toward base but not abruptly inflated. TEXAS. San Patricio Co.: grow-
ing around shores of Lake Corpus Christi, Fred B. Jones 1160, 6 July
1955 (SMU). The species is reported by Alexander (N. Amer. Flora 19:
56—57, 1937) from Mexico, Central America, the West Indies, and South
America. It is not included in the recent (undated; 1961?) Flowering
Plants and Ferns of the Texas Coastal Bend Counties by Fred B. Jones,
Chester M. Rowell, Jr., and Marshall C. Johnston, which lists E. cras-
sipes only, as occurring in “ditches, lakes, etc.” — Lloyd H. Shinners.

SIDA 1 (2): 99. 1962.

100

ANEILEMA (COMMELINACEAE) IN THE UNITED STATES. —
Two Asiatic species of Aneilema have become established in the South-
eastern United States within the present Century, but as yet no Ameri-
can flora treats both of them. A Mexican species has been found very
close to the Texas border, and is worth noting as something to watch

but the uppermost with short but distinctly differentiated sheath;
southeastern Texas east to Atlantic Coast
2a. Sepals 2—3 mm. long, glabrous; flowers early becoming exserted
on long, naked peduncles, several or many in each inflorescence
udiflorum
2b. Sepals 5—6 mm. long, hispid-pilose on back, at aes near tip;
flowers tardily exserted on short, usually 1-flowered peduncles
A. Keisak
1b. Perennial with tuberous-thickened roots, the erect stem with 1—3
leaves, only the lowest with well defined sheath; Mexico, possibly
PPAR ARC OBS LEMS y ac i oe bis Baa te bk eee eee 3. A. lineare
1. A. NUDIFLORUM (L.) R. Brown ex Kunth, Syn. Pl. 4: 66. 1843.
(This combination is merely implied by R. Brown in Prodr. Fl. Nov.
Holl. p. 271, 1810: “Hujus generis sunt Commelina virgata, nudiflora,
spirata, medica, Vahl. enum.” Vahl, Enum. Pl. 2: 176, 1785, credits Com-
melina nudiflora to Linnaeus.) Commelina nudiflora L., Sp. Pl. 1: 41—42.
1753. (Linnaeus cites his own Flora Zeylanica and Plukenet’s Almages-
tum. In his Mantissa Plantarum Altera, p. 177, 1771, he gives a new
description evidently based on later material than was available in
unth, in validating the transfer to Aneilema, specifies the
Piers illustration as representative, in effect making it the type.
Fortunately this agrees with Linnaeus’s own emendation. C. B. Clarke,
in DC., Mono, Phan. 3: 144, 1881, retains the name Commelina nudiflora
L. “Sp. Pl. 1, p. 61, nec Mant. 177, nec auct.” in place of C. diffusa Bur-
man, citing a Ceylon specimen in Burman’s herbarium. Presumably he
thought this the same as the plant described in Linnaeus’s Flora Zey-
lanica. At the same time he recognizes Aneilema nudiflorum on p. 210,
which is nomenclaturally impossible by present rules, since the latter

“Commelina nudiflora, Linn.! Mant. p. 177 (nec Sp. Pl. nee Fl. Zeyl.),”
reversing the typification established by Kunth and implied by Linnaeus
himself. Since Linnaeus, in the citation from the Flora Zeylanica,
specifies “involucro nullo” (and presumably because of that very
peculiarity chose the epithet nudiflora), it is difficult to understand
why Clarke should have equated Commelina nudiflora with the spathe-
bearing C. diffusa. In any case his rejection of Kunth’s typification can-

SIDA I (2): 100—101, 1962.

101

not be accepted. The double listing of the Linnaean binomial in Index
Kewensis is to be disregarded; only the first entry, as synonym of
Aneilema nudiflorum, should stand.)

This species was first reported from the United States by Small in
1910 (Bull. Torr. Bot. Club 37: 513—514) as established around Braiden-
town (Bradenton, Manatee Co.), Florida, where it had been observed
for several years. In the second edition of his Flora of the Southeastern
United States (p. 1328, 1913), the range is given as southern Georgia and
Florida. In his Manual of the Southeastern Flora (p. 263, 1933), he says
‘Fla. to Ga.” The plant has spread to the western Gulf region, as shown
by the following collection. TEXAS. Jefferson Co.: nursery garden of
P. A. Winkler, Voth Road, Beaumont, V. L. Cory 49974, 3 October 1945
(SMU). “Introduced from further east, and is a pest and hard to eradi-
cate from gardens.”

2. A. KEISAK Hasskarl, Commelinaceae Indicae pp. 32—34. 1870. With
var. (alpha) nutans, pp. 33—34, and var. (beta) erectum, p. 34. First
reported from the United States by Neil Hotchkiss in 1940 (Rhodora
42: 21) from Minim Island, Georgetown County, South Carolina, as A.
nudiflorum. This he corrected to A. Keisak in reporting it from numer-
ous localities from Virginia to Georgia in 1951 (Rhodora 53: 92—93).
Radford in the latter year also reported it from several localities in
North Carolina (ibid. p. 25). Fernald considered it a native species o
bicentric distribution between Virginia and eastern Asia (Rhodora 42:
392, 441442; 1940; Gray’s Manual, 8th ed. p. 393, 1950). In view of
the other records, there is little doubt of its being introduced in North
America.

3. A. LINEARE (Bentham) Woodson (as linearis), Ann. Mo. Bot. Gard.
29: 148. 1942. Tradescantia linearis Bentham, Pl. Hartw. p. 27. 1839.
Superficially this has considerable resemblance to Tradescantia Wrightit,
differing conspicuously in the branched, open inflorescence, quite un-
like the condensed, umbel-like one typical of Tradescantia. The follow-
ing collection comes from just south of the Big Bend region of Texas.
COAHUILA. Frequent in shade on north, igneous slopes of Picache del
Centinela, Del Carmen Mts., alt. 6000 ft., Barton H. Warnock 11624, 24
August 1953 (SMU). Other specimens have been seen from Durango and
Jalisco.

In publishing the generic name Aneilema, Robert Brown treated it
as feminine. It may be argued that he was entitled to do so, since it was
a manufactured word. The Jast three syllables are a Greek word mean-
ing veil or covering, and the word in Greek is neuter. Since such Lin-
naen generic names as Erigeron have been altered from the gender as-
signed by Linnaeus to the etymologically correct one, consistency re-
quires that the same be done with Brown’s. — Lloyd H. Shinners.

SIPHONYCHIA TRANSFERRED TO PARONYCHIA (CARYOPHYL-
LACEAE). — The small Southeastern genus Siphonychia has been
maintained as distinct from Paronychia on the basis of having perianth

SIDA 1 (2): 101—I103. 1962.

102

parts united. But in a majority of the species these are joined much
less than half way, while in S. Rugelii the joining is very weak, the

firmly united. On the other hand, in Paronychia Drummondii the seg-
ments are united about a fourth their length at maturity. The generic
difference is thus one of degree only, not of basis structure, and the
degree is virtually non-existent in the case of Paronychia Drummondii.
The most striking thing about the species of Siphonychia is their over-
whelming similarity in nearly all respects to Southeastern representa-
tives of Paronychia. I can distinguish only four species rather than the
seven accepted by Earl L. Core in “A taxonomic revision of the genus
Siphonychia,” Journ. Elisha Mitchell Sci. Soe. 55: 339—345, 1939. Their
names under Paronychia are as follows.

P. patula Shinners, nom. nov. Siphonychia diffusa Chapman, FI.
U.S. (ed. 1) p. 47. 1860. Not Paronychia diffusa A. Nelson, Bull. Torr.
Bot. Club 26: 237, 1889

P. AMERICANA (Nuttall) Fenzl ex Walpers, Rep. 1: 262. 1842.
Herniaria americana Nuttall, Amer. Journ. Sci. 5: 291. 1822. Siphonychia

flora Small, Fl. S.E. U.S. (ed. 1) pp. 402, 1330. 1903. Nuttall’s original
description speaks of the plant as “paucifloris” and goes on to specify
that “the clusters of flowers contain from about eee to five.” Torrey
and Gray, in a description based in part on the same material as
Nuttall’s, say “Bracts very small, similar to the leaves.” Thus by Small’s
key the type material of S. americana must be placed in S. pauciflora.
In reality the distinctions do not hold, and I can make out only a single
somewhat variable species.

P. erecta (Chapman) Shinners, comb. nov. Siphonychia erecta Chap-
man, l.c. p. 47 S. corymbosa Small, Bull. Torr. Bot. Club 24: 337. 1897.
Odontonychia corymbosa (Small) Small, Fl. S.E. U.S. (ed. 1) pp. 402,
1330. 1903. It is possible to recognize three and possibly more pubescence
types in this species. The original S. erecta is a form with stem largely
glabrous, but short-pubescent above. More common are forms with
stem either short-pubescent or relatively long-pubescent throughout.
They do not seem to be so much variations in degree as a group of
distinct Mendelian characters. Core, following Small, describes the
perianth segments of S. corymbosa as ovate, which is not correct; his
illustration accurately depicts them as long and narrow, precisely as
in S. erecta. As with the stem, there is variation in pubescence on the
perianth tube. Again I can recognize only one somewhat variable species.

P. RUGELII Shuttleworth ex Chapman, Fl. S. U.S. (ed. 3) p. 397.
1897. (Published earlier as synonym only, under the next.) Siphonychia
Rugelii (Shuttleworth, ined.) Chapman, Fl. S. U.S. (ed. 1) p. 47. 1860.
Forcipella Rugelii (Shuttleworth) Small, Bull. Torr. Bot. Club 25: 150.

103

1898. Gibbesia Rugelii (Shuttleworth) Small, ibid. p. 621. Odontonychia
interior Small, Man. S.E. Fl. pp. 483, 1504. 1933. Siphonychia interior
m

for his segregate genus Forcipella (Gibbesia). Here is what Small origi-
nally had to say: “Flowers .. . 2—3 in an involucre composed of two
bracts and their broad 2-parted stipules, each, or only 2 seated in a
hard, clamp-like involucel, whose two lobes are notched. Calyx of 5
linear-subulate distinct (sometimes cohering at the base) sepals.” His
illustration in the Manual shows only the bracts and stipules, not the
clamp-like involucel. In Godfrey & Houwk 60289 (SMU), from Lafayette
County, Florida, many of the cymes have abortive (diseased?) flower
buds around the bases of open flowers. Conceivably these were what
Small took to be an involucel; I have been unable otherwise to find any
such structure. The perianth tube is scarcely differentiated, the seg-
ments merely having appressed margins toward base rather than being
firmly united. In other specimens there is a definite tube, the segments
being distinctly united. It is understandable that Chapman should at
different times have put this species in both Siphonychia and Paronychia.
I concur in his second choice, restoring it to the genus in which Shuttle-
worth had first placed it.

To the above may be added a few supplementary notes on species
included in Core’s “The North American species of Paronychia,” Amer.
Midl. Nat. 26: 369—397, 1941.

P. DRUMMONDII T. & G. Hitherto known only from Texas, this can
now be added to the flora of LOUISIANA. Allen Parish: 7.2 miles west
of Kinder, Shinners 21,489, 8 September 1955 (SMU). “Sandy fencerow,
pine land. Perianth white.”

P. BALDWINII (T. & G.) Fenzl. Including P. riparia Chapman. Those
familiar with the Gulf States know how uncertain the difference be-
tween annual and perennial often is. I am unable to distinguish two
species on this basis, and pubescence varies quite independently of ap-
oie ae

rarennGus (Rafinesque) Fernald var. NUTTALLII (Wood)
one Formerly known only from three counties in Pennsylvania,
this occurs also in VIRGINIA. Giles Co.: on dry rocky open exposed
path to the top of Bald Knob, % mile s.e. of Mountain Lake P.O., alt.
4300 feet, Hugh H. Iltis 2011, 1 August 1943 (SMU). The perianth is
pubescent and the segments rather prominently short-awned. — Lloyd
H. Shinners.

STELLARIA COREI SHINNERS, NOM. NOV. (CARYOPHYL-
LACEAE). — Based on S. pubera ssp. silvatica Beguinot, Nuov. Giorn.
Bot. Ital. n.s. 17: 385. 1910. S. pubera var. silvatica (Beguinot) Weather-
by, Rhodora 26: 171. 1924. S. silvatica (Beguinot) Maguire ex Gleason,
Phytologia 4: 23. 1952 (March). (Not S. sylvatica (Maxim.) Maxim. ex
Regel, 1862.) S. tennesseensis (non Mohr) Strausbaugh & Core, Castanea

104

17: 165. 1952 (December). (Basinym cited as Alsine tennesseensis Small,

A. puber co
ee Sane to a form of var. pubera. Small mistakenly applied
the name to var. silvatica, as did Strausbaugh and Core.) I believe that
this plant is properly treated as a species distinct from, although closely
related to, S. pubera. But neither of the names it has received in that
rank can be retained. It seems altogether fitting to associate with it the
name of the man who as botanist, editor, and administrator has done
so much for Appalachian and Southeastern botany

I wish to thank Dr. G, B. Van Schaack of the Missouri Botanical Gar-
den for confirming the validity of Stellaria sylvatica (Maxim.) Maxim.
ex Regel. — Lloyd H. Shinners.

RANUNCULUS TRACHYCARPUS (RANUNCULACEAE) IN SOUTH-
CENTRAL LOUISIANA: NEW TO NORTH AMERICA. — The occur-
rence of the Mediterranean Ranunculus trilobus Desf. in Pointe Coupee
Parish, Louisiana, was reported in 1960 (S.W. Nat. 5: 170). On seeing
quite similar plants in April, 1962, in nearby Avoyelles Parish, I as-
sumed that they represented a range extension for the new introduc-
tion. But on comparison of herbarium specimens they proved to be
quite different, having decidedly spiny instead of merely papillose-
roughened achenes, and the larger leaves were not pinnately divided.
In the key in Lyman Benson’s “A Treatise of the North American
Ranunculi” (Amer, Midl. Nat. 40: 1—261, 1948: see especially p. 110)
they ran to R. muricatus L. which they obviously were not, having
larger petals and smaller achenes, more or less pilose stems, leaves, and
sepals, and long-pedicelled flowers. They could not be determined with
any of the recent standard North American or European floras, but

h. & Mey. in G E

Orientalis 1: 55—56, 1867. iors was a single apceienen so named in the
SMU Herbarium, from Sharon Plain, Kabbara Marshes, Israel, A. Grizi
s.n., 24 May 1954. This consists of two mowed plants which had put
up new branches with smaller, more divided leaves than those of the
Louisiana specimens, and have more densely spiny achenes, but they
evidently belong to the same species, which is described as a variable
one. Ranunculus trachycarpus Fisch. & Mey. may therefore be recorded
from LOUISIANA, Avoyelles Parish: 7.2 miles southeast of Bunkie,
Shinners 29,519, 18 April 1962 (SMU). “Roadside ditch, wet silty clay.
Petals yellow. Locally abundant from Bunkie to here.”

In reporting R. trilobus, I said that the manner of introduction was
entirely unknown. In the case of R. trachycarpus there is a clue from
some associated weeds: Persian clover (Trifolium resupinatum) and

SIDA 1 (2): 104—105. 1962.

105

crimson clover (T. incarnatum), both of which are widely planted on
road shoulders almost throughout Louisiana and which maintain them-
selves in such habitats. T. resupinatum is the more persistent of the
two, and shows more tendency to spread, especially to ditches or along
ditch banks. Both clovers are native to the Mediterranean region. If
commercial seed imported from that area had been used for some of
the road-shoulder plantings, it may have been the vehicle for the
transport of the two species of Ranunculus. — Lloyd H. Shinners.

WAREA AURICULATA INSTEAD OF W. AMPLEXIFOLIA OF
SMALL (CRUCIFERAE). — In his Manual of the Southeastern Flora
(pp. 573—574, 1933), Small uses the name Warea amplexifolia (Nuttall)
Small for a plant with auricled-clasping upper leaves. The combination
is nomenclaturally impossible, since it is identical with one already
made by Nuttall himself. Small supposed that Nuttall had had a mix-
ture, part of his material being W. sessilifolia Nash. In such case he
ought to have written W. amplexifolia (Nuttall) Nuttall emed. Small.
Payson (Ann. Mo. Bot. Gard. 9: 312—312, 1922) also believed that
Nuttall had had a mixture, agreeing with Small except for the author-
citation. Neither author checked Nuttall’s type material, the belief that
it was mixed resting only on two pieces of circumstantial evidence. The
epithet amplexifolia is inappropriate for a plant with merely sessile or
barely clasping upper leaves, although this is what was illustrated by
Nuttall himself in transferring Stanleya ? amplexifolia to Warea. In
his first publication, under Stanleya, he gave “‘East Florida” as the place
of origin, but in the second this becomes “West Florida.” Neither item
can be accepted as proving that Nuttall had a mixture. In evaluating
the epithet amplexifolia it has mistakenly been assumed that the ques-
tion is which of two Florida plants it fits better. This is the wrong frame
of reference. Nuttall was thinking in terms of a Florida plant and the
only previously described Stanleya, S. pinnatifida (S. pinnata), which
has petioled leaves, and the epithet amplexifolia is therefore to be
taken as relative to a petioled-leaved condition. This is not so inap-
propriate for Warea sessilifolia with sessile to slightly clasping upper
leaves; furthermore the lower leaves of this species may be distinctly
clasping. The reference to “West Florida” in the second publication
dealing with Nuttall’s species may have been merely a slip of the pen
on his part, or it may have been a deliberate correction or clarification
of the very vague older usage of the terms “East Florida” (primarily
northeastern as now delimited, but sometimes loosely applied to more
than half of northern Florida) and “West Florida” (originally meaning
only the Pensacola area, but at times extended to include the entire
panhandle section). Florida had then been only recently added to the
United States, and these terms were both subject to change. Hence

SIDA 1 (2): 105—106. 1962.

106

neither of the inferential arguments about a Nuttallian mixture can be
accepted.

Much more important than supposition is the question of what Nut-
tall actually had. Dr. Walter M. Benner kindly checked material at the
Philadelphia Academy for me. He reports that there is only one speci-
men named by Nuttall, and that it has sessile leaves. In other words,
the only concrete evidence we have indicates that Nuttall did not have
a mixture, and the only thing he did have was the plant shown in his
illustration of Warea amplexifolia. This is identital with W. sessilifolia
Nash, and the plant thought to be W. amplexifolia by Nash, Small, and
Payson becomes

W. auriculata Shinners, sp. nov. W. amplexifoliae affinis, sed foliis
superioribus profunde auriculato-amplexicaulibus. ee ee sandy
soil, Lake Wilson Road, Loughman, Kissimee, Osc Co., Florida,
Mary L. Singeltary, 25 September 1937 (DUKE, no. sare PARATYPE:
High hill near Lake Apopka (Orange Co: ?), Florida, Ralph McWilliams,
22 September 1935 (SMU; Schallert Herb. No. 19696). This is Warea
amplexifolia in the sense of recent authors, not of Nuttall, for whose
plant the following synonymy may be given.

W. AMPLEXIFOLIA (Nuttall) Nuttall, Journ. Phila. Acad. 7: 88, with

inadmissible combination W. amplezifolia ‘“(Nuttall) Small,” Bull. Torr.
Bot. Clu b 23: 409, 1896, belongs here on the basis of type, but was

woo : :
Godfrey en 22 SoM: 1955 (DUKE, SMU). “Lower leaves on
non-flowering plants clasping.”

I am grateful to Dr. Benner for the information concerning Nuttall’s
specimen at Philadelphia, and to Dr. R. L. Wilbur for the loan of ma-
terial from the Duke University Herbarium. — Lloyd H. Shinners.

WISSADULA GRANDIFOLIA INSTEAD OF W. MACRANTHA
(MALVACEAE): NOMENCLATURAL CORRECTIONS. — In his “En-
twurf einer Monographie der Gattungen Wissadula und Pseudabutilon”
(Kungl. Svenska Vetenskapsak. Handl. Bd. 43 No. 4, 1908), Rob. E. Fries
described Wissadula macrantha as a new species, comprising three varie-
ties. One of the three, var. grandifolia, was based on a species named a
year before. Because it represented only a localized race, Fries con-
sidered it undesirable to retain it as a species and subordinate the most
widespread race as a variety. Under present rules of course the pre-
sumed phylogenetic status has no bearing on the nomenclature: the first
name in the rank of species is the one that must be used for the whole
ensemble. Though more than half a century has elapsed since the

107

monograph was published, the necessary nomenclatural emendations
have not been made. They are supplied herewith.

WISSADULA GRANDIFOLIA E. G. Baker ex Rusby, Bull. N.Y. Bot.
Gard. 4: 328. 1907. W. macrantha var. grandifolia (E. G. Baker) R. E.
Fries, Kungl. Sv. Vet.-Akad. Handl. 43 no. 4: 68. 1908.

W. GRANDIFOLIA var. brevipedunculata (R. E. Fries) Shinners,
comb. nov. W. macrantha var. brevipedunculata R. E. Fries, l.c. p.

W. GRANDIFOLIA var. macrantha (R. E. Fries) Shinners, comb.
nov. W. macrantha R. E. Fries, l.c. pp. 67—68. (Illegitimate name, since
the earlier W. grandifolia was included within the circumscription of
the new species.) — Lloyd H. Shinners.

GENTIANA DELOACHII (W. P. LEMMON) SHINNERS, COMB.

n dry acid

Statesboro, Bulloch Co.” Described as having separate anthers. The
article “Connate anthers in Gentiana (Gentianaceae)” by Wilbur H.
Duncan and Clayd L. Brown (Rhodora 56: 133—136, 1954) does not
mention this species. — Lloyd H. Shinners.

AGASTACHE BREVIFLORA (GRAY) EPLING VAR. HAVARDII
(GRAY) SHINNERS, COMB. NOV. (LABIATAE). — Based on Cedro-
nella breviflora var. Havardi Gray, Proc. Amer. Acad. 20: 309. 1885.
Agastache pallidiflora Hie aaa ssp. Havardi (Gray) Lint &
Epling, Amer, Midl. N 33: 1945. Found in the Chinati, Chisos,
Davis, and Guadalupe Remee Trans-Pecos Texas. The distinctions
between A. breviflora and the variable A. pallidiflora, as treated in
Harold Lint and Carl Epling’s “A revision of Agastache” (Amer. Midl.
Nat. 33: 207—230, 1945) are not convincing. I prefer Gray’s earlier,
broader version, which under Agastache requires this new combination.
— Lloyd H. Shinners.

SCUTELLARIA LAEVIS (LABIATAE), ANOTHER ENDEMIC IN
TRANS-PECOS TEXAS. — Scutellaria is an extremely difficult genus,
especially in western and southwestern Texas. It is gratifying to be
able to name a new species which is very distinct and easily recognized.
It is in most respects a perfectly typical member of the Section Resinosae
as defined in Carl Elping’s “The American species of Scutellaria’”
(Univ. California Pub. Bot. 20 no. 1: 57—69, 1942), but is unique in that
group in having a glabrous stem. Because of this peculiarity it is here
named,

S. laevis Shinners, sp. nov. Perennis multicaulis humilis (20—-35 cm.
alta) sublignosa e radice crasso lignoso. Caules glabri. Folia brevissime
petiolata integra parce minutissimeque scabro-puberula vel glabrius-
cula, inferiora elliptico-ovata penninervia obtusiuscula 11—21 mm. long

108

6—11 mm. lata, superiora gradatim minora oblongo-lanceolata. Flores
spicato-racemosi brevipedicellati pedicellis scabro-puberulis. Calyx
3.0—3.5 mm. longus minutissime scabro-puberulus vel glabriusculus.
Corolla pallida 14—17 mm. longa extus pilosula. Nuculi crebre obtuse
muriculati. HOLOTYPE: ten miles north of Van Horn, north slope of
Beach Mountain, Culberson Co., Texas, D. S. Correll 13973, 19 August
1946 (SMU). “Flowers white dino with lavender, talus.” PARA-
TYPES (both SMU): on red sandstone along arroyo and base of canyon
in southwestern slopes of Beach Mtn., 6% miles northwest of Van Horn,
Culberson Co., Texas, U. T. Waterfall 5087, 14 July 1943. Numerous a
the bed of ie creek in the head of Victoria Canyon, alt. 5600
Hudspeth Co., Texas, L. C. & Leon Hinckley 102, 18 June 1949 (in ae

The light- eoloied corolla is also distinctive in this group, if con-
sistent, but color is known only from the type collection, Other species
of Resinosae have corolla deeply colored (purple-blue or lavender-
blue) except for a white spot on the palate. — Lloyd H. Shinners.

SOLANUM GODFREYI SHINNERS, NOM. NOV. (SOLANACEAE).
— Based on S. floridanum Shuttleworth ex Dunal in DC., Prodr. 13 (1):
306. 1852. (Not S. floridanum Rafinesque, Fl. Tell. p. 107. 1840.) “In
Floridae apricis, prope S. Marks,” Rugel (type not seen). S. carolinense
var. floridanum (Shuttleworth) Chapman, Fl. S. U.S. p. 349. 1860. I
agree with Asa Gray (Syn. ‘Fl. N.A. 2 pt. 1: 22%, 1878) that this should

basi stellatis pilosus.’ The only specimen I have seen has entirely
glabrous stem, but stellate-pubescent leaf blades. It is in fruit only, and
apart from the stem seems to fit quite well the rather lengthy descrip-
tion given by Dunal. It comes from the general region of the type lo-
cality, but slightly farther east, in Taylor County: clearings of swampy
woodland, vicinity of Nuttal’s Rise, along the Aucilla River, R. K.
Godfrey 60037, 7 July 1960 (SMU). — Lloyd H. Shinners.

D CONTRIBUTIONS
S TO BOTANY

VOLUME 1 NUMBER 3 DECEMBER 1963

CONTENTS

The taxonomic significance of evolutionary parallelism.
Arthur Cronquist. 109

Botanical Survey along the Yellowknife Highway, Northwest
Territories, Canada. |. Catalogue of the Flora.
John W. Thieret. 117

Gilia and Ipomopsis (Polemoniaceae) in Texas.
Lloyd H. Shinners. 17]

Leptoglossis and Nierembergia (Solanaceae) in Texas.
Lloyd H. Shinners. 180

NOTES. Monanthochloe littoralis (Gramineae) in Louisiana. 182.—Cenchrus longisetus M. C.
Johnston, nom. nov. (Gramineae). 182.—Centrosema floridanum (Britton) Lakela, comb. nov.
(Leguminosae). 182.—The varieties of Teucrium canadense (Labiatae). 182.—Dicerandra

)
) in

immcaulata Lakela, sp. nov. (Labiatae). 184.—Wahlenbergia linarioides (C
Florida: a second adventive species for the United States. 185

SIDA is privately published by Lloyd H. Shinners, SMU Box 473,
Dallas 22, Texas, U.S.A. Subscription price $6 (U.S.) per volume of about

360—400 pages, parts issued at irregular intervals.

©
SIDA Contributions to Botany volume 1 number 3 pages 109—186
copyright 1963
by Lloyd H. Shinners

THE TAXONOMIC SIGNIFICANCE OF
EVOLUTIONARY PARALLELISM’
ARTHUR CRONQUIST
New York Botanical Garden, New York 58

The recognition by Darwin that the natural system of classification
is in essence an evolutionary system is a major landmark in taxonomic
history. Let me read you a passage from Chapter 14 of Origin of Species:
“... The Natural System is founded on descent with modification . . the
characters which naturalists consider as showing true affinity between
any two or more species, are those which have been inherited from a
common parent, all true classification being genealogical . . community
of descent is the hidden bond which naturalists have unconsciously been
seeking... .” A corollary of this principle is that a natural taxonomic
group is per se monophyletic, and that polyphyletic groups are in con-
trast artificial and should be rejected.

Those concepts have become so firmly entrenched in taxonomic think-
ing as to approach the status of dogma. However, as we learn more about
phylogeny, a strict and literal application of the monophyletic require-
ment in taxonomy has gotten us into more and more trouble. More and
more groups that have been considered to be natural are turning out not
to be strictly monophyletic. They may be natural in the sense that all
the included subgroups are evolutionarily closely related and have many
features in common, but they are not monophyletic in the sense of
being descended from an original species of the group which has all}
the essential characters of the group.

The mammals are an outstanding example, with a well documented
fossil record, of a natural taxonomic group which is clearly not mono-
phyletic in the strictest sense. We shall return to this fact in a few
minutes.

ture of angiosperms, can be observed among the living members of the

primitive order Ranales. Within the Ranales one can also see all stages.
’ Presidential address delivered to the American Society of Plant Taxonomists at Corvallis, .

Oregon, 29 August 1962.

SIDA 1 (3): 109—116. 1963.

110

with filament and anther took place in several related evolutionary
lines within the Ranales, in parallel fashion, rather than being strictly
monophyletic. Differentiation of the perianth into calyx and corolla
has likewise taken place independently in various families, as has
also the origin of petals from staminodes.

Double fertilization and the extreme reduction of the female gameto-

of the group.

This same problem with the monophyletic requirement, often in an
even more severe form, permeates our whole system of classification.
The tribes of the Compositae do not in general represent strictly mono-
phyletic groups; rather they are constellations of genera which show
certain evolutionary trends in common ana are to some extent linked

would be a Conyza, and the common ancestor to all species of Conyza
would be an Erigeron. The most primitive existing species of Erigeron,
in turn would on morphological grounds be just as well referred to
Aster, and in fact it was first described as Aster peregrinus Pursh. It
is now referred to Erigeron mainly because of its obvious relationship
to species which are necessarily included in Erigeron.

The difficulties attendant on a strict application of the monophyletic
concept in classification have led a few taxonomists in recent years to
exclude the concept entirely from taxonomic theory and practice, to

111

the conflict, rather than to dissolve what has been a most fruitful union.

I believe the conflict is resolvable, and the terms of the resolution
have already been laid down, now 17 years ago, by George Gaylord
Simpson. In 1945 he pointed out that the monophyletic requirement must
be interpreted broadly in order to be useful taxonomically (see “The
principles of classification and a classification of mammals,” American
Museum of Natural History Bulletin, vol. 86). Again, let me quote:
“The condition that classification must be consistent with phylogeny
has as its most important corollary the requirement that all the animals
within a given group, whatever its rank, must have had a common
origin. . . . The principle that the units of classification must have a

history of life and that certainly cannot be demonstrated in any case.
The rule that a group, to be considered monophyletic, must be derived

edge is so far from complete that adherence to such a rule would lay
classification open, to an unnecessary and undesirable degree, to the
caprices of shifting theory and individual opinion. It is not useful to
set up a classification in which groups with different names cannot be
distinguished morphologically, but this does happen if theoretical mono-

most practical and, at least for the present, the most desirable additional
requirement seems to be not that it should be derived from one im-
mediately antecedent genus or species, but, with intentional vagueness,
that its immediate ancestry should be included within a group of lower
rank than itself. For instance, it is not probable on the basis of present
knowledge that all the animals here included in the Mammalia arose
from the Reptilia as a single species, genus, or even family, but it is not
suggested on this account that some of them should be returned to the
Reptilia or that another class should be created for them, They certainly
arose from a unified group of reptiles of much smaller scope than a
class, perhaps a family or perhaps a superfamily, and for practical pur-
poses this is an adequate fulfillment of the requirement of monophyly.”

112

To recapitulate, if a taxonomic group of a particular rank is derived
wholly from another group of lower rank, that is a sufficient degree of
monophylesis for taxonomic purposes. Dr. Simpson has very recently
(1961, in his book Principles of Animal Taxonomy) modified this con-
cept to read “of the same or lower rank,” and here I feel he may have
gone a bit too far. His point, however, is well taken. Monophylesis and
polyphylesis are not such ae distinct things as the terms would
suggest. There is a continuous gradation from the strictest monophylesis
to the most utter polyphylesis in proposed taxonomic groups. In order
to be natural and acceptable, a taxonomic group must fall somewhere
toward the monophyletic end of this scale, rather than toward the
polyphyletic end.

It now appears that a workable taxonomic system cannot provide a
perfect reflection of evolution, no matter how abundant the evidence
on which it is based. Furthermore, conclusions must usually be based
on more or less inadequate evidence; none of us has witnessed the
events of geologic time. But the phylogenetic concept still provides the
underlying rationale for the natural system. Taxonomy can provide only
a somewhat muddy reflection of evolution, but a reflection all the same.

Once we admit the broad interpretation of the monophyletic require-
ment, as I believe we must do in order to have a workable system, then
we are committed to the position that similarities due to evolutionary

weight should be given to parallelism is another question, to which we
will return later.

As long ago as 1912 Wernham pointed out that “critical tendencies
are no less important than critical characters” (see the final paper in
his series on floral evolution in the New Phytologist, vol. 11). He
further pointed out that “the general relation between the significant
features of the ancestry and those of the descendants is, that in the former
the characters in question are not constant throughout the group, nor
may they be completely evolved. In other words, we are dealing with
tendencies to characters, and not with the critical characters themselves,

unites ancestor and descendants represents the transition between the
tendencies and their realization.” He goes on to provide several ex-
amples. Probably most of us here can provide equally good examples
from our own experience.

The proposition that similarities due to common descent, to inherit-
ance from a common ancestor, indicate relationship is of course self-
evident. We have pointed out that taxonomists have been forced, on a
pragmatic basis, to accept also the proposition that similarities due to

113

parallelism also provide some evidence of relationship. Let us now

proceed to an inquiry into why this should be so, or how it can be so.
A concise expression of the most generally accepted present concept

of the mechanism of evolution would be something like this: Random

thinking. To me, and I think to many other people, random implies
that any one thing is as likely as any other thing. Thus in a table of

control or predict them individually; they are certainly not at ran-
dom in the sense of one mutation being as likely as any other. It is
abundantly clear that the different mutations which can occur in a
particular gene do not occur with equal frequency, nor is the mutation
rate from one allele to another the same in opposite directions. There
are even genes which influence the rate and dirction of mutation in
other genes. All this is well known to geneticists. As long ago as 1940
Dobzhansky stated that mutability, like other characteristics, is under
partial genetic control.

one group, and that different groups will have different evolutionary
potentialities. At the grosser levels this is of course immediately obvious

114

that the occurrence of diffuse centromeres in these two families repre-
sents a case of parallelism rather than of inheritance from a common
ancestor. I should add, of course, that no one character by itself pro-
vides proof positive of relationship; diffuse centromeres also occur in
Spirogyra. Nevertheless, the occurrence of this rare character in the
Juncaceae and Cyperaceae suggests that these two families have in com-
mon an unusually high potentiality to evolve in this direction. Some
of the foregoing information on the occurrence of diffuse centromeres
was provided for me by John Ebinger.

Clarkia and Oenothera provide another example at a lower taxonomic
level. Here I get my information, as you might guess, from Harlan
Lewis. Clarkia ea occurs in dry country, bordering
deserts, but not actually in them. Individual colonies at the drier margins
of the range lead a precarious existence, and are subject to being wiped
out in bad years. The plants are annual, and the seeds germinate as
soon as the moisture and temperature conditions are right. If the mois-
ture supply then fails before the seeds are mature, no seeds are left
over to begin again the following year. It appears that the one thing
Clarkia lacks to be a potentially successful desert annual is a variable
period of dormancy, so that some seeds would last over until the sec-
ond or third year, regardless of how favorable conditions may be in
the interim. In the many thousands, perhaps millions of years that
Clarkia has occupied a habitat in which such a change would have a
strong survival value, it appears to have been unable to evolve such a
feature. Presumably the proper mutations just have not occurred. The
closely related genus Oenothera, on the other hand, occurring in similar
habitats, has in several lines given rise to desert species with a variable
period of seed dormancy. There is nothing in the obvious character-
istics of the plants to suggest that the evolution of desert species shoul
be easier for the one genus than the other; the difference instead ap-
pears to lie in the mutative potentialities.

Now let us return to theoretical considerations. The existence of the
same character in two different groups may be due either to their having
inherited the character directly from a common ancestor, or to their hav-
ing developed the character independently by parallel evolution. We are
concerned here only with the second situation, not the first. If the char-
acter was independently developed in the two groups, it is still true, at
the very least, that their respective ancestors had similar evolutionary
potentialities with regard to that character, and thus were genetically
similar—not necessarily identical—in this particular respect. us,
evolutionary parallelism with regard to a particular character is one
straw in the wind, indicating a degree of similarity and possible re-
lationship among the ancestors. Now if we add another character in
which these same two groups resemble each other, by parallel evolu-
tion, we have two straws in the wind, and so on. Thus, the more char-

115

Therefore, we come to the general principle that evolutionary par-
allelism tends to indicate relationship, and that it should be given due
weight, along with other factors, in arriving at taxonomic conclusions.

e
allelism, and vice versa. If the control is essentially selective, then

oJ

frequency than other mutations, so that the mutation pressure, operating
along with the other causes of speciation, can cause parallel evolutionary
developments.

n theoretical grounds, therefore, characters which are not closely
correlated with survival value and ecologic niches are likely to be more
important taxonomically, in many instances, than characters which are
directly influenced by selection. This is especially true in the angio-
sperms, in which the evolutionary barriers between different ecological

niches
On purely pragmatic grounds, plant taxonomists over the last two

116

riers between different ecologic niches are formidable, and within each
general niche the selective pressures operating on different but related
taxa tend to be similar, and different from the selective pressures OP~

groups are recognized in the vertebrates—but that is another story.
It is interesting to note that Wernham, with no knowledge of genes
and not very much knowledge of heredity, also came to the conclusion
that nonadaptive characters are more likely to be important than adap-
tive ones. He spoke of “biological” characters, which are directly re-

which have no relation to the environment nor to any biological func-
tion. He concluded that “A group of plants may share a number of
biological characters in common without being therefore nearly re-

closer the affinity.” To this I would add simply that this evidence is
significant even if the similarities are due to parallel evolution instead
of direct inheritance from a common ancestor.

BOTANICAL SURVEY ALONG
THE YELLOWKNIFE HIGHWAY,
NORTHWEST TERRITORIES, CANADA
I. CATALOGUE OF THE FLORA
JOHN W. THIERET
University of Southwestern Louisiana, Lafayette
The Yellowknife Highway, begun in 1957 and officially opened in
1961, makes accessible a previously remote area in the District of Mac-
kenzie, Northwest Territories. The area was botanically almost un-
known. Thus the highway offered an unparalleled opportunity for

1s paper is one of two that will present the data obtained. It con-
tains a catalogue of the plants collected, a total of 4160 numbers. The
concluding paper will contain descriptions of the region and of its
major plant communities.

Southwestern District of Mackenzie, that portion of Mackenzie in
which the highway is located, is about the size of Louisiana, Mississippi,
Alabama, and Georgia combined. Prior to the work of Porsild (1945)
and Raup (1947), little serious field study of plants had been done there.

portion of the Canol Road. Raup’s Botany of Southwestern Mackenzie

SIDA 1 (3): 117—170. 1963.

118

118°

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Fig. 1. The Yellowknife Highway Region, Northwest Territories, Canada

119

knife airport; and a mile number alone indicates the Enterprise-Mac-
kenzie River section and refers to miles from Enterprise. For each

120

species, a specimen from each of the three sections is cited when such
specimens are available. That a species is not ascribed to a particular
section does not necessarily mean that it does not occur there. The
few species seemingly limited to any one section are so designated in
the text. I have included in the catalogue a few species that were not
found along the Yellowknife Highway itself but were collected along
the adjacent Mackenzie Highway in the Northwest Territories. Unless
otherwise noted, all specimens cited have been deposited in the her-
barium of Chicago Natural History Museum (F), the repository for
most of the material collected during the survey.

The catalogue includes 5 algae, 35 lichens, 38 bryophytes, and 452
vascular plants. The seed plants are represented by 61 families, among
which the largest are Compositae (55 species), Cyperaceae (51), Grami-
neae (35), Cruciferae (23), and Rosaceae (22). Among the seed plants
the genera represented by 10 or more species are Carex (37), Salix
(18), and Potamogeton (10), The genera Poa and Calamagrostis, which
are incompletely treated, are to be made the subject of separate papers
at a later date.

PLS Pele
CHARACEAE

Chara aspera Willd. var. macounit Allen. In shallow water of road-
side pool, mile 110.5 N, 7728, and of marly stream, mile 39.7 N, 8420.

Chara contraria A. Br. In shallow water of marly lakes, mile 40.5,
5426, mile 37, 6138.

Chara contraria A. Br. var. hispidula A. Br. In shallow water of marly
lake, mile 37, 6139.

Chara globularis Thuill. In shallow water of stream, mile 75 N, 7375.

Tolypella prolifera Leonh. In shallow water, Kakisa Lake, 5674.

LICHENS
PELTIGERACEAE

Peltigera aphthosa (L.) Willd. Infrequent, usually growing in moss
mats in rich spruce forests. Number 4519 is var. aphthosa; 8028 is var.
variolosa (Mass.) Thoms. Kakisa Road, 4519; mile 125.1 N, 8028.

Peltigera malacea (Ach.) Funck. Jack pine forest on sandy knoll,
mile 123.4 N, 8071.

CLADONIACEAE

Cladonia alpestris (L.) Rabh. Frequent to abundant as a ground cover
in spruce or pine forests; common on crystalline and limestone out-
crops. In the open jack pine forests around mile 120 N, the white sand
is almost completely covered with this and other lichens, and the ground
appears yellow. Mile 28.5, 4932; mile 123.4 N, 8053; mile 39.8 S, 8326.

121

Cladonia alpicola (Flot.) Vainio. Crystalline outcrop, mile 39.8 S,
8325 (p.p.).

Grane amaurocraea (Flk.) Schaer. Black aie Ronee mile 53,
4954; jack pine forest on sandy knoll, mile 123.4 N,

Cladonia coccifera (L.) Willd. Jack pine forest on ee knoll, mile
123.4 N, 8067.

Cladonia cornuta (L.) Schaer. Spruce forest, mile 125.1 N, 8038;
crystalline outcrop, mile 39.8 S, 8325 (p.p

Cladonia degenerans (Flk.) Spreng. Crystalline outcrop, mile 39.8 S,
8325 (p.p

Clanenig gonecha (Ach.) Asahina. Spruce forest, mile 125.1 N, 8039;
jack pine forest on sandy knoll, mile 123.4 N, 8068.

Cladonia gracilis (L.) Willd. var. pita (Hoffm.) Schaer. Jack pine
forest on sandy knoll, mile 123.4 N, 8069.

Cladonia metacorallifera ae Crystalline outcrop, mile 39.8 5,
8325 (p.p.).

Cladonia mitis Sandst. Frequent in spruce or pine forests and on
limestone and crystalline outcrops. Mile 12, 4869; mile 123.4 N, 8062;
mile 43.7 S, 8315.

Cladonia pyxidata (L.) Hoffm. var. neglecta (Flk.) Mass. Jack pine
forest on sandy knoll, mile 123.4 N, 8069; crystalline outcrop, mile 39.8
S. 8325 (p.p

Cladonia Paneenne (L.) Web. Frequent to common as a ground
cover in spruce or pine forests; frequent on limestone and crystalline
outcrops. Mile 12, 4868; mile 125.1 N, 8051; mile 39.8 S, 8327.

Cladonia sylvatica (L.) Harm. Abundant as a ground cover in a spruce
forest, Kakisa Road, 4516.

Cladonia uncialis (L.) Web. Jack pine ae on sandy knoll, mile
123.4 N, 8063; crystalline outcrop, mile 43.7 S, 8316.

Cladonia verticillata (Hoffm.) Schaer. Crystalline outcrop, mile 39.8
S, 8325 (p.p.).

eva sa tomentosum E. Fr. Spruce forest, mile 125.1 N, 8024.

GYROPHORACEAE

Actinogyra muhlenbergii (Ach.) Schol. Common on crystalline out-
crops. Mile 43.7 S, 831

Lasallia pensylvanica (Hoffm.) Llano. Common on crystalline out-
crops. Mile 43.7 S, 831

LECANORACEAE

ee ae inaequatula (Nyl.) Zahlbr. Limestone outcrop, mile 127

N, 829
PARMELIACEAE

Cetraria crispa (Ach.) Nyl. Black spruce ee mile 112.2 N, 8125;

jack pine forest on sandy knoll, mile 123.4 N, 8066.

122

Cetraria culcullata (Bell.) Ach. Spruce forest, mile 125.1 N, 8030.

Cetraria islandica (L.) Ach. Spruce forest, mile 125.1 N, 8040.

Cetraria nivalis (L.) Ach. Frequent to common in spruce or pine
forests and on limestone and crystalline outcrops. Mile 12, 4870; mile
123.4 N, 8055; mile 39.8 S, 8317.

Cetraria tilesii Ach. Rare on limestone outcrops. Wile 22:7, 5176; s0ule
127 N, 8290.

Parmelia centrifuga (L.) Ach. Crystalline outcrop, mile 43.7 S, 8320.

Parmelia stenophylla (Ach.) Hueg. Crystalline outcrop, mile 39.8 S,
8323.

Parmelia sulcata Tayl. Crystalline outcrop, mile 39.8 S, 8322.

USNEACEAE

Evernia mesomorpha Nyl. Common on bark of birches in spruce forest,
mile 125.1 N, 8027.

Thamnolia vermicularis (Sw.) Ach. Limestone outcrop, mile 127 N,
8296.

Usnea comosa (Ach.) R6Ohl. Common on bark of birches in spruce
forest, mile 125.1 N. 8026.

CALOPLACACEAE
Caloplaca elegans (Link) T. Fr. Limestone outcrop, mile 127 N, 8299.
BUELLIACEAE

Buellia papillata (Somerf.) Tuck. Limestone outcrop, mile 127 N,
8294
PHYSCIACEAE
Physcia muscigena (Ach.) Nyl. Spruce forest, mile 125.1 N, 8025;
limestone outcrop, mile 127 N, 8291.

BRYOPHYTES
SPHAGNACEAE

Sphagnum balticum Russ. On sedge mats around muck bottom lakes.
Mile 57.6 5S, 7247.

Sphagnum capillaceum (Weiss) Schrank var. tenellum (Schimp.)
Andrews. Black spruce forest, mile 59.8 S, 6763.

Sphagnum cuspidatum Ehrh, In shallow water on sedge mats and in
forest pools. Mile 59.8 S, 6764.

Sphagnum fuscum (Schimp.) H. Klinggr. Apparently the most com-
mon Sphagnum in the region. Usually found in hummocks in forests,
principally of black spruce, where it may be the main ground cover.
Mile 53, 4951; mile 59.8 S, 6762; mile 36 N, 7541.

Sphagnum girgensohnii Russ. Black spruce forest, mile 126 N, 9141a.

Sphagnum riparium Angstr. In Ledum muskeg, mile 44.8 S, 7222.

123

Sphagnum squarrosum Pers. ex. Crome. Black spruce forest, mile 126
N, 9141.

Sphagnum warnstorfianum DuRietz. Occasional to common, forming
mounds or mats in black spruce forests and in wet muskegs. Mile 52,
5370; mile 44.8 S, 7223; mile 36 N, 7540.

POLYTRICHACEAE

Polytrichum formosum Hedw. In depressions on crystalline outcrops,
where it forms mats on thin soil. Mile 39 S, 7934.

Polytrichum juniperinum Hedw. Disturbed soil, Kakisa Road, 5359;
crystalline outcrop, mile 11.5 S, 6713.

Polytrichum juniperinum Hedw. var. alpestre (Hoppe) BSG. Spruce
forest, Kakisa Road, 4557; disturbed soil, mile 81.5 N, 6887.

Polytrichum piliferum Hedw. In peaty depressions on crystalline out-
crops. Mile 14.5 S, 7152.

DITRICHACEAE

Ceratodon purpureus (Hedw.) Brid. Disturbed soil, mile 4.2 N, 6625.

Ditrichum flexicaule (Schwaegr.) Hampe. Limestone outcrops, mile
22.7, 5176a, mile 127 N, 8297; spruce forest, Kakisa Road, 4763; shrub
zone around marly lake, mile 64, 6051.

DICRANACEAE

Dicranum bergeri Bland. Shrub zone around a marly lake, mile 64,
6052; spruce forest, mile 125.1 N, 8034.

GRIMMIACEAE
Hedwigia ciliata (Hedw.) P.-B. Crystalline outcrop, mile 27.7 S, 7204.

AULACOMNIACEAE

Aulacomnium acuminatum (Lindb. et Arn.) Par. Frequent to common
as a ground cover in spruce forests, Kakisa Road, 5351; mile 65.8 S,
7990

Aulacomnium palustre (Hedw.) Schwaegr. Frequent in spruce forests,
where it may be important as a ground cover. Mile 27.7, 4936.

Aulacomnium turgidum (Wahl. ex Web. et Mohr) Schwaegr. Lime-
stone outcrop, mile 128 N, 6788; crystalline outcrop, mile 8.3 S, 7134.

MEESIACEAE

Meesia tristicha BSG. In sedge mat around lake, mile 47.2 S, 4957.
Paludella squarrosa (Hedw.) Brid. White spruce forest, mile 62.8 S,
9.

BRYACEAE
Brynum lacustre Bland. Marl deposit, mile 110.5 N, 6837.
Leptobryum pyriforme (Hedw.) Schimp. Disturbed soil, mile 2.5 S,
(C®

124

HYPNACEAE

Campylium stellatum (Hedw.) Lange et C. Jens. Black spruce forests,
mile 44.5, 5122, mile 115 N, 8284; sedge mat and shallow water of marly
lake, mile 63, 5299.

Drepanocladus aduncus (Hedw.) Warnst. var. polycarpus (Bland. ex
Voit) Warnst, Sedge meadow, mile 23.2 S, 7906.

Drepanocladus capillifolius (Warnst.) Warnst. In muck and shallow
water of lake, mile 20.5 S, 7175; sedge meadow, mile 11.4 S, 7833.

Drepanocladus exannulatus (BSG) Warnst. In sedge mats and in
shallow water of muck bottom lakes. Mile 44.8 S, 7218.

Drepanocladus fluitans (Hedw.) Warnst. Shallow pool in black spruce
forest, mile 59.8 S, 6765; among rocks in shallow water, Mackenzie River,
7460.

Drepanocladus vernicosus (Lindb.) Warnst. Shallow water of marly
lake, mile 40.5, 5425.

Hylocomium splendens (Hedw.) BSG. The common “feather moss”
of the region. Especially abundant in rich white spruce forests, where
it forms a dense carpet on the floor, as it may do also in certain black
spruce forests: Characteristic of more mesophytic forests of jack pine,
where it occurs in scattered patches. Mile 33.5, 5990; mile 42.5 N, 6918;
mile 65.8 S, 7986.

Hypnum bambergeri Schimp. Black spruce forest, mile 72 N, 7651.

Scorpidium scorpioides (Hedw.) Limpr. Shallow water of marly lake,
mile 54, 5079.

Tomenthypnum nitens (Hedw.) Loeske. Frequent to common in white
or black spruce forests, where it is a characteristic ground cover. Mile
66, 5268; mile 42.5 N, 6923; mile 62.8 S, 7979.

THUIDIACEAE

Abietinella abietina (Hedw.) Fl. In moss mat on limestone slab, mile
128 N, 6779.

FONTINALACEAE

Fontinalis duriaei Schimp. On rocks in boulder rapids, Kakisa River,
5701.

PLAGIOCHILACEAE
Mylia anomala (Hook.) S. F, Gray. Among Sphagnum fuscum in
forests, mile 113.5 N, 6822, mile 45 S, 7941.
PTILIDIACEAE

Ptilidium ciliare (L.) Nees. Among Dicranum bergeri in spruce forests,
mile 125.1 N, 8035.

MARCHANTIACEAE
Marchantia polymorpha L. Edge of roadside pool, mile 61.2 N, 7604.

125

RICCIACEAE
Ricciocarpus natans (L.) Corda. In shallow water among Carex, Mac-
kenzie River, 5995, and among Lemna minor in small pond, mile 26 Ss,
9276.

VASCULAR PLANTS
EQUISETACEAE

Equisetum arvense L. Infrequent along shores and in moist forests;
locally common in disturbed areas. Kakisa River, 4539; mile 2.5 S, 6673;
mile 24 N, 7437,

Equisetum fluviatile L. Infrequent in shallow water, in marshes, in
sedge mats, and on shores. Mackenzie River, 4239; Kakisa Lake, 4898;
mile 85 N, 6880; mile 3.6 S, 7798.

Equisetum palustre L. Infrequent along shores, in shallow water, and
in black spruce forests; locally frequent in disturbed areas. Mile 44,
4299; mile 119 N, 7747a; mile 66 S, 8019.

Equisetum scirpoides Michx. Infrequent in sandy soil or moss mats in
forests of pine or spruce; locally common in disturbed soil. Mile 123.4 N,
7294; Enterprise, 9051.

Equisetum sylvaticum L. Infrequent in peaty soil or in moss mats in
moist forests, especially of black spruce; locally frequent in disturbed
soil. Kakisa Road, 4508; mile 2.5 S, 6668.

Equisetum variegatum Schleich. Seen only once, sandy-gravelly beach
of Kakisa Lake near outlet, 6096.

LYCOPODIACEAE
Lycopodium complanatum L. Rare in dry sandy soil in pine or spruce
woods. Mile 11.5, 4835; mile 59.3 S, 6747; Prelude Lake, 9221.
SELAGINELLACEAE

Selaginella selaginoides (L.) Link. Rare in muskeg forests and in
shrub zones around marly lakes. Mile 57.5, 5217; mile 119 N, 7740

ISOETACEAE

Isoetes echinospora Dur. var. braunii (Dur.) Engelm. Seen only once,

in shallow water, sandy bouldery bottom, Prelude Lake, 9216.
POLYPODIACEAE

Cryptogramma crispa (L.) R. Br. var. acrostichoides (R. Br.) Clarke.
Infrequent in shallow soil or in crevices on rock outcrops, mainly
crystalline but occasionally limestone. Mile 66 S, 7257; Horseshoe Island,
Yellowknife Bay, 9492.

Cystopteris fragilis (L.) Bernh. Infrequent in crevices on limestone
outcrops, Mile 15.5, 5153; mile 66 S, 6778

Cystopteris montana (Lam.) Bernh. Seen only once, in a spruce-

126

feather moss forest on slope above Kakisa River 1.5 miles below Lady
Evelyn Falls, 5350.

Dryopteris robertiana (Hoffm.) Christensen. Seen only once, in crevices
on face of limestone escarpment in white spruce forest, mile 15.5, 5156.

Dryopteris fragrans (L.) Schott. Infrequent in crevices on crystalline
outcrops. Of the collections made of this species, only 9178, with its
overlapping pinnae, approaches var. fragrans; the others are clearly var.
remotiuscula Kom. Yellowknife, 9178; mile 16 S, 9296.

Polypodium virginianum L. Infrequent in crevices on rock outcrops,
mainly crystalline but occasionally limestone. Mile 2.7 S, 6692; mile
66 S, 7258.

Woodsia glabella R. Br. Infrequent in crevices on limestone outcrops.
Mile 26, 4336; mile 66 S, 6775.

Woodsia ilvensis (L.) R. Br. Frequent in rock crevices, especially on
crystalline outcrops, but also on limestone. Mile 2.7 S, 6688; mile 66 S,
7259.

PINACEAE

cline for about half their length and then ascend. The reclining portion
of the branches usually roots. This plant is most characteristic, perhaps,
of jack pine forests on sandy knolls and limestone outcrops, where it
forms a distinct low-shrub stratum. Mile 70, 4246; mile 2.7 S, 6686;
mile 96.7 N, 7699.

Juniperus horizontalis Moench. (Ch) Frequent in a variety of habitats,
including rocky slopes, forests of most kinds, and, most commonly, rock
outcrops (both crystalline and limestone). Stems prostrate, to at least
20 feet long and 1.7 inch in diameter. Creeping juniper exists along the
highway in two strikingly distinct forms, glaucous and green. These
forms are most effectively contrasted where they grow together, as on
the limestone outcrops at mile 26-28. Here, the elongate branches of
many plants criss-cross each other, forming in many spots a distinctive
reticulate pattern on the rock, Mile 36, 4610; mile 96.5 N, 6868; Yellow-
knife, 9187.

Larix laricina (Du Roi) K. Koch. (Ms) Frequent in muskeg forests,
although seldom forming pure stands of any extent. Larch is most com-
monly found in moist to wet areas, but it occasionally occurs in seeming-
ly xeric habitats, such as atop crystalline outcrops, where it grows in
crevices. The maturing cones are magenta and glaucous. Larch is the
least common of the gymnospermous trees in the highway region. Trees
between 26 and 32 feet tall and 3.3 to 4.9 inches DBH were found to

127
have 24 to 29 annual rings. Mile 64, 6059; mile 2.5 S, 6676; mile 26.5 N,
7519.

Picea glauca (Moench) Voss. (Mg) Common to abundant in white

height and 15.7 inches DBH and did not show more than 91 annual
rings; most were appreciably smaller and younger than this. Mile 65,
6048; mile 74 N, 6894; Yellowknife, 9183.

Picea mariana (Mill.) BSP. (Ms) Common to abundant in black spruce
forests; occasional on crystalline outcrops and on sand plains and
ridges. Black spruce, the commonest tree in hydric sites, seldom ex-

31 feet tall showed about 190; one 4.3 inches DBH and 27 feet tall
showed about 178; and one 3.3 inches DBH and 18 feet tall showed 56.
Kakisa Road, 4576; mile 2.5 S, 6677.

Pinus banksiana Lamb. (Ms) Characteristic tree of sandy knolls, sand
plains, and rock outcrops (both crystalline and limestone). Although
most commonly found in these rather xeric habitats, jack pine can oc-

TYPHACEAE
Typha latifolia L. (HH) Infrequent in wet places or shallow water.
Most commonly observed in disturbed areas, especially roadside ditches;
only infrequently seen in undisturbed areas. In a few sites cattail is a
contributor to the mat around lakes in the Canadian Shield section. Mile
1, 5553; mile 61.2 N, 7594; mile 6.8 S, 7815.

SPARGANIACEAE
Sparganium angustifolium Michx. (incl. S. multipeduncalatum [Mor-
ong] Rydb.) (HH) infrequent in mud or muck of shores or road-
side ditches or in still or flowing water to about 2 feet e s

128

mm. wide; generally the plants growing in the deepest water have the
narrowest leaves. Leaves are commonly flat in the upper half and
convex on the back in the lower half. Foliar characteristics and others,
including size and number of pistillate heads and length of stigma,
frequently used to distinguish S. augustifolium from S. multipeduncula-
tum, appear to break down in our northern material, a conclusion
reached also by Hulten (1941-1950) and Anderson (1959). Kakisa River,
5710; mile 7.7 S, 7818; mile 12 N, 8253.

Sparganium minimum (Hartm.) Fries. (HH) Infrequent in mud or
muck of shores or roadside ditches or in still or flowing water to about
1.5 feet deep. The material I refer to this species is rather variable but it
surely represents only one taxon. Most specimens have 1, 2, or 3 pistillate
heads, but a few have 4 or even 5. The mature heads vary in diameter
from 8 to 12 mm. In about half the plants observed, all the pistillate
heads are sessile, but in the others the lowest head(s) are peduncled,
with peduncles to 23 mm. long. The peduncles are usually axillary, but
occasionally one is clearly supra-axillary. The peduncle of the stami-
nate head varies from 3 to 18 mm. long. Mile 50, 5717; mile 30 over:

ZOSTERACEAE

Potamogeton alpinus Balbis var. tenuifolius (Raf.) Ogden. (HH) Rare
in still water to at least 1.5 feet deep, clay or muck bottom. Noted only
in Canadian Shield section. Mile 9.7 S, 7823.

Potamogeton filiformis Pers. (HH) Infrequent in still or flowing water
to at least 1.5 feet deep, gravel, marl, or clay bottom. Numbers 7637 and
7814, cited below, represent var. borealis (Raf.) St. John; 9061, with
leaves about 1 mm. wide, is best referred to var. macountt Morong.
Mile 23.5, 9061; mile 68.2 N, 7637; mile 6.8 5, 7814.

Potamogeton foliosus Raf. var. macellus Fern. (HH) Rare in shallow
still water, clay or muck bottom. Noted only in Canadian Shield sec-
tion. Mile 38.3 S, 8356.

Potamogeton friesii Rupr. (HH) Rare in shallow still water, clay or
muck bottom. Noted only in Canadian Shield section. Mile 12.7 S, 7858.

Potamogeton gramineus L. (HH) Frequent in still or flowing water
to at least 2 feet deep, gravel, clay, or muck bottom. On the specimens
cited below, 5702 clearly is var. maximus Morong; 8372 is var. grami-
neus; and 8307 is intermediate in leaf characteristics between these var-
ieties. Kakisa River, 5702; Stagg River, 8307; mile 54.4 S, 8372.

Potamogeton pectinatus L. (H Seen only once, among boulders in
shallow water of Mackenzie River at Fort Providence, 4232,

Potamogeton pusillus L. (HH) Infrequent in still or flowing water
to at least 2 feet deep, clay or muck bottom. Noted only in Canadian
Shield section. Mile 7.7 S, 7816.

Potamogeton richardsonii (Benn.) Rydb. (HH) Infrequent in still or
flowing water to at least 4 feet deep, gravel, clay, or muck bottom.

129

Kakisa Lake, 5633; mile 12 N, 8252; mile 49 S, 8306.

Potamogeton vaginatus Turcz. (HH) Rare in still or flowing water
to at least 1.5 feet deep, gravel or sand bottom. Kakisa Lake, 4615;
Yellowknife, 9172.

Potamogeton zosteriformis Fern. (HH) Rare in still water to at least
3 feet deep, muck bottom. Noted only in Canadian Shield section. Mile
36.4 S, 8351.

JUNCAGINACEAE

Triglochin maritima L. (Hrr) Infrequent along rocky or sandy shores,
in marshy areas, in marl deposits, and in sedge mats and shallow water
around marly lakes. Mackenzie River, 4134; mile 66, 4260; mile 72 N,
7033; Prosperous Lake, 9196.

Triglochin palustris L. (Hrr) Infrequent along rocky or sandy shores,
in marshy areas, in marl deposits, in sedge mats and shallow water
around marly lakes, and in spruce forests. Kakisa River, 4276; mile 72
N, 7036; Yellowknife, 8342.

ALISMATACEAE

Kakisa Lake, 5659; mile 49 S, 7959.
GRAMINEAE

Agropyron cristatum Gaertn. (Hs) Seen only once, in disturbed soil,
Fort Providence, 9075a.

Agropyron latiglume (Scribn. et Sm.) Rydb. (Hs) Seen only once, in
disturbed soil, mile 59, 6073

Agropyron repens (L.) Beauv. (Grh) Seen only once, in weedy area,
Yellowknife, 7777

Agropyron trachycaulum (Link) Malte. (Hs) Infrequent to common
along sandy or clay shores, in shallow residual soil on limestone out-

* : . . .

crops, and in disturbed soil. Dominant, with Muhlenbergia richardsonis,

angliae (Scribn.) Fern.; 4137, 7992, and 8224 are var. glaucum (Pease
et Moore) Malte; and 7471 is best referred to var. unilaterale (Cassidy)
Malte. Mile 17 N, 4136, 4137; Fort Providence, 4229; mile 41, 4308; mile

130

24, 5631, 5632; Kakisa River, 5747, 6093; mile 86.7 N, 7063; mile 10 N,
7471: mile 66 S, 7992; Mackenzie River, 8224

Agrostis scabra Willd. (Hs) Infrequent on outcrops, both crystalline
and limestone, in marl deposits, and along mucky, marly, or sandy
shores, becoming frequent to common in disturbed areas. Mile 66, 5235;
mile 66 S, 7262; mile 110 N, 7299

Alopecrus aequalis Sobol. (Hs) Rare in marshes, in sedge mats
around muck-bottom lakes, and along clay, gravelly, sandy, or mucky
shores; becoming frequent in disturbed areas. Fort Providence, 5041;
mile 23.5, 5777; mile 82 N, 7053; mile 16.5 S, 7878.

Arctagrostis latifolia (R. Br.) Griseb. (Incl. A. arundinacea [Trin.]
Beal) (Hsr) Local in disturbed soil along highway; seen only once in
an undisturbed habitat, a white spruce forest above shore of Great
Slave Lake, mile 62.8 S, 7978. Mile 7.5 N, 4215; mile 33, 4319.

Beckmannia syzigachne (Steud.) Fern. (Th) Rare on gravelly, muddy,
or sandy shores and in marshes, becoming frequent in disturbed areas.
Mile 11 N, 4223; Kakisa River, 5210; Prosperous Lake, 9223.

Bromus inermis L. (Hsr) Seen only once, along Mackenzie Highway
4 miles south of Hay River, 5536.

Bromus pumpellianus Scribn. (Hsr) Rare in dry sandy soil in pine or
spruce woods and along rocky shores. Numbers 4234 and 7309 are
notable for the length (up to 4.5 cm.) of their cna Fort Providence,
4234; mile 46, 5404; mile 66 S, 7287; mile 110 N,

Calamagrostis. My collections of eee from the highway

C. lapponica (Wahl.) Hartm., C. neglecta (Ehrh.) Gaertn., or C. pur-
purascens R. Br. Many others cannot be convincingly referred to any of
these species but appear to be intermediates. The specimens are dis-
couragingly variable in those characteristics commonly regarded as
diagnostic in Calamagrostis. I plan to present, in a separate paper, an
analysis of this variation.

Deschampsia cespitosa (L.) Beauv. (Hs) Infrequent on rocky or sandy
shores, in shallow residual soil over limestone, and in marl deposits.
Kakisa River, 4014; mile 110 N, 7306; mile 64.6 S, 8379.

Elymus canadensis L. (Hs) Seen only once, on gravelly shore of Hay
River near mile 49 of Mackenzie Highway, 6167

Elymus innovatus Beal. (Hsr) Infrequent to rare in sand or in moss
mats in white spruce or jack pine forests, becoming frequent in dis-
turbed areas. Fort Providence, 5045; mile 52, 5099; mile 103.2 N, 7342.

Festuca rubra L. (Hsr) Seen only once, in disturbed clay, mile 21.5
N, 6969.

Festuca saximontana Rydb. (Hs) Infrequent in dry sandy soil, es-
pecially in pine or spruce woods, in shallow residual soil over limestone,

131

and on crystalline outcrops, becoming frequent in disturbed areas. Mile
22.7, 5174; mile 70.5 N, 6906; mile 6.1 Su Zoe

Glyceria borealis (Nash) Batch. (Hsr) Rare on muddy or sandy shores
or in shallow water. Seen only in Canadian Shield section. Mile 20.5 S,

Glyceria grandis Wats. (Hsr) Rare along muddy, sandy, or mucky
shores or in shallow water. Kakisa River, 5711; mile 20.5 S, 7900; mile
126 N, 9139.

Glyceria pulchella (Torr.) Trin. (Hsr) Infrequent along muddy, sandy,
or mucky shores; locally abundant in marshes. Four miles northeast of
Fort Providence, 4123; mile 56, 5011; mile 26.9 S, 7911.

Glyceria striata (Lam.) Hitche. (Hsr) Rare along muddy, sandy, or
peaty shores. Mile 13, 5588; mile 66 S, 8021.

Helictotrichon hookeri (Scribn.) Henr. (Hs) Rare in shallow residual
soil over limestone. Noted only south of the Mackenzie River. Mile 22.7,
5161.

Hierochloe odorata (L.) Beauv. (Hsr) Infrequent along shores or in
shallow residual soil on limestone outcrops, Kakisa River, 4730; mile
39.7 N, 6993a.

Hordeum jubatum L. (Hs) Infrequent in drier grasslands, on rocky
shores, and in marl deposits, becoming locally frequent to common in
disturbed areas. Mile 17 N, 4143; mile 66, 5256; mile 23.5 S, 7190.

Koeleria cristata (L.) Pers. (Hs) Infrequent to rare in shallow residual
soil over limestone, Noted only south of the Mackenzie River. Mile 28.5,

Muhlenbergia glomerata (Willd.) Trin. var. cinnoides (Link) Herm.
(Hsr) Rare in marl deposits or in sedge mats around marly lakes, usually
growing on ant hills. Noted only south of the Mackenzie River. Mile 50,
9718

Muhlenbergia richardsonis (Trin.) Rydb. (Hsr) Rare on rocky shores,
in shrub zones around marly lakes, and in limestone crevices; common
in prairies northeast of Fort Providence, where it may be co-dominant
with Agropyron trachycaulum. Mile 17 N, 4191; mile 50, 5397.

Oryzopsis asperifolia Michx. (Hs) Rare in dry peaty or sandy soil in
woods, especially jack pine or white spruce. Noted only south of the
Mackenzie River. Kakisa River, 4741.

limestone. Mile 33, 4318; mile 103.5 N, 6850.

Phalaris arundinacea L. (Grh) Infrequent along rocky or marly
shores and in marl deposits. Number 9450 is notable for the length—up
to 29.5 cm.—of its panicles. Mackenzie River, 4233; mile 59.5 N, 7378;
Kakisa River, 9450.

Phalaris canariensis L. (Th) Seen only once, disturbed soil, mile 3 N,
9434 (in herb. DAO).

132

Phleum pratense L. (Hs) Rare in disturbed soil. Kakisa River, 5474;
mile 21.5 N. 7503.

a. The genus Poa is represented in the highway region by at least
the ee species: P. alpina L., P. glauca Vahl, P. interior Rydb.,
P. leptocoma Trin., P. palustris L., P. pratensis L., and P. stenantha Rydb.
My collections total 114 numbers. They will be reported on in a separate
paper

De omneiie distans (L.) Parl. (Hs) Seen only twice, disturbed soil,
mile 16 S, 7154, and sandy shore of Prosperous Lake, 9198

Puccinellia nuttalliana (Schultes) Hitche. (Hs) Rare in marl deposits,
becoming frequent in disturbed areas. Enterprise, 3898; mile 39.7 N,

Scolochloa festucacea (Willd.) Link. (Hsr) Locally frequent to com-
mon or even dominant in marshes, in marl deposits, and on marly shores.
Four miles northeast of Fort Providence, 4065; mile 39.7 Ni o7b50; mile
Gi I, der.

Sphenopholis intermedia (Rydb.) Rydb. (Hs) Rare along sandy rocky
shores and in disturbed areas. Mile 0.5, 5939; Mackenzie River, 6001.

Trisetum spicatum (L.) Richt. (Hs) Rare in shallow residual soil or
in crevices on limestone outcrops, and in sandy soil in pine woods.
Mile 23.5, 5134; mile 110 N, 7304

CYPERACEAE

Carex aenea Fern. (Hs) Frequent to common in disturbed soils,
especially sand; rare in seemingly undisturbed sand in jack pine woods
and in peaty depressions on crystalline outcrops. Mile oy 5111; mile 110
N, 7298; mile 1.8 S, 7784.

Carex aquatilis Wahl. (Grh ‘HH}) Frequent to abundant in shallow
water and on shores, in marl deposits, and in sedge mats; rare in moss
and lichen mats on limestone and in spruce forests. This, the commonest
sedge of the highway region, in co-dominant with Carex atherodes and
Scolochloa festucacea in the extensive marshes north of Fort Provi-
dence. It is an important contributor to the sedge mats around both
marly and muck bottom lakes. Four miles northeast of Fort Providence,
4029; Kakisa Lake, 4651; mile 59.5 N, 7377; mile 7 S, 9248.

Carex atherodes Spreng. (Grh [HH]) Frequent to abundant in shal-
low water and on shores in marl deposits, and in prairies. Carex
atherodes is a co-dominant species, with Carex aquatilis and Scolochloa,
in the extensive marshes north of Fort Providence, and, in the same
area, is co-dominant in prairies with Calamagrostis neglecta and
Agropyron trachycaulum. Four miles northeast of Fort Providence,
4027; mile 72 N, 8269.

Carex aurea Nutt. (Grh) Local, mostly in disturbed damp sandy,
peaty, or clay soil; occurring also on marly shores and in grasslands.
Kakisa River, 5339; mile 65.6 N, 7615

133

Carex bebbii Olney. (Hs) Infrequent in residual soil over limestone
and in disturbed sandy or peaty soil. Kakisa River, 5332.

Carex buxbaumii Wahl. (Hsr [HH]) Infrequent to abundant in sedge
mats, in marl deposits, and in disturbed peaty soil; noted once in a
black spruce-Hylocomium splendens forest. Carex buxbaumii is, in
places, an important contributor to the mat around marly lakes. Its
rhizomes often extend 2 to 3 feet into open water beyond the edge of
the mat. Mile 66, 4258; mile 28.5 N, 7430.

Carex canescens L. (Hs) Local in disturbed soil at roadside, on sedge
mats and shores of muck bottom lakes, and on crystalline and limestone
outcrops. Much of the material I refer here is seemingly transitional to
Carex brunnescens (Pers.) Poir. Mile 46.6 S, 7225; mile 35 N, 7537.

Carex capillaris L. (Hs) Infrequent to rare on hummocks in marshes,
in marl deposits, in sedge mats, in moss mats in black spruce forests,
and in disturbed soil. Mile 56, 4789; mile 110 N, 7318

Carex capitata L. (Hsr) Infrequent to rare in spruce-feather moss
forests and in disturbed sandy, peaty, or clayey soil. Kakisa River,
4545; mile 42.5 N, 6920; mile 66 S, 7267.

these “stolons” are up to 6 cm. long, and from the nodes arise leaves
and flowering culms—but no roots. The “stolons” survive the winter
and continue growth in length the following spring, evidencing typi-
cally chamaephytic behavior. Mile 4.7 , 6702.

Carex concinna R. Br. (Hsr) Tere quent to rare in moss mats in spruce
forests, in shallow residual soil over limestone, in marl deposits, in
grassy openings in woods, and in disturbed sandy soil. The plants are
most robust in disturbed areas, The culms, especially of more vigorous

eaten River 4522; mile inet

Carex crawfordii Fern. (Hs) Seen ae once, in disturbed sandy soil
along road to ford over Kakisa River, 4287.

Carex deflexa Hornem. (Hsr) Rare; observed only in disturbed situa-
tions, either ead or peaty soil. Kakisa Road, 4514; mile 8.3 S, 7129;
mile 110 N,

Carex ae scan (Hs) Infrequent in sedge mats around both marly
and muck bottom lakes, and in marl and muck deposits. Mile 63, 0292;
mile 16.2 S, 7171.

Carex disperma Dewey. (Grh) Infrequent to rare in mars shy areas
and in disturbed sandy, peaty, or clay soil. Kakisa Lake, 4627; mile 32
N, 6975; mile 46.6 S, 7227.

Carex eburnea Boott. (Grh) Rare in crevices of shaded limestone out-
crops. Mile 26, 4340; mile 124.5 N, 7289

Carex foenea Willd. (Grh) Local in sand in dry situations, usually in

134

pine woods; also in disturbed sand at roadside. Mile 110 N, 7301; Yellow-
knife, 7775.

Carex garberi Fern. (Grh) Infrequent along rocky or marly shores,
in marl deposits, and in disturbed loam or peat soils. Mile 7, 4807;
Mackenzie River, 7458; mile 110.5 N, 7735.

Carex glacialis Mack. (Hs) Seen only once, in crevices in limestone
cliff, mile 66 S, 7286

Carex gynocrates Wormsk. (Grh) Infrequent in moss mats in spruce
forests and in disturbed peaty soil adjacent to these forests. Kakisa
River, 5343; mile 72 N, 7027.

Carex interior Bailey. (Hsr) Seen only once, in sedge mat around
marly lake, mile 61, 5276.

Carex lasiocarpa Ehrh. (Grh [HH]) Infrequent to common along
shores, in sedge mats around both marly and muck bottom lakes, in
shallow water, and in disturbed peaty soil. Carex lasiocarpa is an im-

portant contributor, in places, to sedge mats. Of the collections cited
below, 7459 represents var. latifolia (Béck.) Gleason (Carex lanuginosa
Michx.); the others are var. americana Fern. Mile 44, 4303; Mackenzie
River, 7459; mile 6 S, 7810a.

Carex leptalea Wahl. (Hsr) Infrequent to rare in hummocks in
marshes, persisting and becoming locally frequent in disturbed gravelly
or peaty soil. Mile 56, 4790; mile 35 N, 7539.

Carex limosa L. (Grh [HH]) Rare to frequent in sedge mats around
both marly and muck bottom lakes; sometimes also in shallow water
beyond edge of mat. Mile 61, 5280; mile 44.8 S, 7219.

Carex media R. Br. (Hs) Rare in spruce-feather moss forests, be-
coming somewhat more common in disturbed soil. Mile 66, 5245; mile
32 N, 6976, mile 4.8 S, 7117.

Carex paupercula Michx. var. pallens Fern, (Hs) Seen only once, on
muck bottom of drained lake, mile 20.5 S, 7174.

Carex physocarpa Presl. (Grh [HH}) Infrequent in marl deposits, in
shallow water of marly lakes, and in marshes. Mile 66, 4256; mile 41.3
N, 7003.

Carex praticola Rydb. (Hs) Infrequent in Calamagrostis or Agrophyron-
Muhlenbergia prairies northeast of Fort Providence, spreading to dis-
turbed soil at roadside. Mile 17 N, 4202

Carex raymondii Calder. (Hs) Rare; seen only in disturbed soil. Mile
0.5, 5548; mile 26.5 N, 7521.

Carex rossii Boott. (Hs) Rare; seen only in disturbed soil. Mile 28.5,
4918; mile 41.3 N, 7001.

Carex rostrata Stokes. (Grh [HH]}) Frequent to abundant in shallow
water, marshes, along shores, in sedge mats, and in roadside ditches.

135

This species is occasionally dominant in marshes and sedge mats. Mile
44, 4304; mile 24 N, 7434; mile 9.6 S, 7821

Carex sartwellii Dewey. (Grh) Seen at only three stations: in disturbed
peaty soil, mile 18, 5600; in marl deposit, mile 39.7 N, 7398; in disturbed
clay, mile 21.5 N, 7496

Carex scirpoidea Michx. (Grh) Rare to frequent in moss mats in
spruce forests, in shallow residual soil over limestone, in marl deposits,
and on marly shores, persisting in disturbed areas. Not seen in Canadian
Shield section. Mile 70, 4255; mile 42.5 N, 6924.

Carex supina Wahl. (Grh) Seen only twice: in peaty-sandy soil atop
crystalline outcrop, 3.3 miles east northeast of Yellowknife, 9188; in
sand among jack pines, Yellowknife, 9240.

Carex tenuiflora Wahl. (Hs) Rare along mucky shores and in dis-
turbed soil in Canadian Shield section. Mile 14 S, 7142.

Carex vaginata Tausch. (Grh) Infrequent to rare in spruce-feather
moss forests and in birch thickets. Mile 70, 4250; mile 42.5

Carex viridula Michx. (Hs) Rare in marl deposits or on marly or sandy
shores. Mile 100.5 N, 7310; mile 62, 9480; Prelude Lake, 9220.

Eleocharis acicularis (L.) R. et S. (Grh [HH]) Local, forming mats
in shallow water or on wet shores, in clay, sand, or muck. Kakisa Lake,
5667; mile 49 S, 7240; Mackenzie River, 8217.

Eleocharis palustris (L.) R. et S. (HH) Local in shallow water up to
1 foot deep or on wet shores. Kakisa Lake, 5663; mile 73.7 N, 7662; mile
5.7 8, 7806.

Eleocharis pauciflora (Lightf.) Link var. fernaldii Svenson. (Gst)
Local in wet marly soil or in sedge mats around marly lakes. Seen only
along the Mackenzie River-Frank Channel section of the highway. Mile
65.6 N, 7613.

Eriophorum angustifolium Honck. (Grh [HH]) Local and infrequent
in marshes, in sedge mats and shallow water of marly lakes, and in
peaty soil in depressions on crystalline outcrops. Kakisa Lake, 4679;
mile 4.2 N, 6613; mile 40 S, 7215.

Eriophorum brachyantherum Trautv. (Hs) Local in marshes, in wet
spruce forests, and in peaty depressions on crystalline outcrops. Mile
43, 4307; mile 4.7 S, 6699; mile 78.3 N, 6892.

Eriophorum chamissonis C. A. Mey. (Grh [HH]) Locally frequent to
common in shallow water or sedge mats of marl or muck-bottom lakes,
in marshes, and in disturbed wet soil. Mile 52, 5372; mile 57.6 S, 7248.

Eriophorum gracile Koch. (Grh [HH]}) Rare on sedge mats or hum-
mocks around muck-bcottom lakes or in shallow water of these lakes.
Seen only in the Canadian Shield section. Mile 35 S, 6741.

Eriophorum spissum Fern. (Hs) Material seemingly best referrable
here was collected once, in a peaty depression on crystalline outcrop,
mile 47.9 S, 7235.

136

Eriophorum viridi-carinatum (Engelm.) Fern. (Grh [HH]) Seen only
once, in shallow water and sedge mat, marly lake, mile 50, 5385

Scirpus cespitosus L. var. callosus Bigel. (Hsr) Common in sedge mats
and thickets around marly lakes; occasional in low spots in black spruce
forests. This plant is frequently the dominant species in sedge mats,
especially in firmer portions of the mat. Mile 63, 5298; mile 113.5 N,

Scirpus hudsonianus (Michx.) Fern. (Hs) Rare in marshes and birch
thickets, Mile 86 N, 6878

Scirpus microcarpus Presl. (Hsr) Seen only once, at edge of sedge
dominated island in Kakisa River 4 miles below highway bridge, 5678.

Scirpus pumilus Vahl ssp. rollandii (Fern.) Raymond. (Grh) Seen only
once, in marl deposit, mile 110.5 N, 7091.

Scirpus validus Vahl. (HH) Infrequent along shores or in shallow
water. The identity of our northern bulrush seems by no means cer-
tain. Many specimens are fairly “typical” S. acutus; many are fairly
“typical” S. validus except that the scales may be conspicuously red-
spotted. Between these extremes occur various intermediates. Until a
more thorough study can be made of Scirpus section Pterolepis in the
north, I prefer to call-our material S. validus, the name by which all
Mackenzie material has been known. Kakisa Lake, 5670; mile 16.5 S,
7877; mile 39.7 N, 8422.

ARACEAE

Acorus calamus L. (HH) Seen only once, with Potentilla palustris,
Calla palustris, and Menyanthes trifoliata, in the mat around a small
lake, along road 3 miles south of Fort Rae, 9484.

Calla palustris L. (HH) Frequent to common in shallow water as a
contributor to the mat around lakes in the Canadian Shield section.
The rhizomes may extend, just under the surface of the water, 2 to 3
feet beyond the inner edge of the mat. Calla palustris ranks in im-
portance with Menyanthes trifoliata and Potentilla palustris as a mat
builder. Mile 20.1 S, 7170.

LEMNACEAE

Lemna minor L. (HH) Local in shallow still water or on mud. Noted
only in the Canadian Shield section. In one pond, Lemna minor was
growing among vast numbers of achenes of Ranunculus gmelinii. Mile
49 S, 7242

Lemna trisulca L. (HH) Locally frequent or even abundant in water
to 2 feet deep, either floating just beneath the surface or forming masses,
sometimes large, on the bottom. Mackenzie River, 5996; mile 38.3 S, 8358.

J UNCACEAE
Juncus albescens (Lange) Fern. (Hs) Rare in moss mats or peaty soil

137

in spruce forests, and in sandy soil along streams. Kakisa River, 5320;
mile 66 S, 8015; mile 122.6 N, 8074

Juncus alpinus Vill. (Grh) Infrequent in marshes, along sandy shores,
in marl deposits, in moss mats and residual soil over limestone, and in
disturbed soil, Most of our specimens of Juncus alpinus seem referrable
to var. alpinus; several have some long-pedicelled flowers in the heads
and so are best called var. rariflorus Hartm. Our material shows all
degrees of intergradation, however, between these varieties. Mile 40.5,
0424; Mackenzie River, 5999; mile 39.7 N, 7555; mile 1.8 S, 7783.

Juncus balticus Willd. var. littoralis Engelm. (Grh) Locally frequent
to abundant on gravelly or sandy shores, in residual soil in low places
on limestone outcrops, and in gypsum and marl deposits; rare in peaty
soil in muskeg forests. Kakisa River, 5509; mile 82 N, 7674; Great Slave
Lake, mile 64.6 S, 8377

Juncus bufonius L. (Th) Infrequent on wet clay or sand, or in shallow
water, becoming frequent in disturbed areas. Kakisa River, 4273; Mac-
kenzie River, 8226; Yellowknife, 8340

Juncus castaneus Sm. (Hsr) Rare in wet clay or sandy soil. Mile 122.6

, 8081.

Juncus filiformis L. (Grh) Seen only once, in wet sand on shore of
Prosperous Lake, 9227

Juncus nodosus L. (Gst) Rare on sandy shores. Kakisa Lake, 5656;
Mackenzie River, 8233.

Juncus stygius L. var. americanus Buch. (Hs) Seen only once, in marl
and shallow water at edge of lake, mile 44.5, 6114.

Juncus vaseyi Engelm. (Hs) Rare along mucky or sandy shores. Yel-
lowknife, 8344

LILIACEAE
Allium schoenoprasum L. var, sibiricum (L.) Hartm. (Gb) Rare along
rocky shores and in residual soil over limestone. The plants are usually

me
in disturbed areas than in adjacent undisturbed areas. Collected in full
bloom July 11-21; in young fruit August 10. Mackenzie River, 4131;
mile 22.5, 5623.

Smilacina stellata (L.) Desf. (Grh) Rare along rocky shores, usually
among shrubs. Collected in flower on June 25, in immature fruit July
26-28. Kakisa River, 4731; Mackenzie River, 8236.

Smilacina trifolia (L.) Desf. (Hsr) Rare in Sphagnum mounds or mats
of feather moss in spruce forests, in marshy spots in spruce forests, and
in peaty depressions in crystalline outcrops. In flower June 17-July 1;
mature fruit in mid-August. Mile 64, 4269; mile 36 N, 7544; mile 39 S,
7937.

Tofieldia glutinosa (Michx.) Pers. (Hsr) Rare in sedge mats around
marly lakes and in wet peaty soil and moss mats in black spruce woods.

138

Collected with flower buds on June 26, in full flower on July 14-16,
and with mature fruit on August 14. Mile 36, 4314; mile 72 N, 7031.

Tofieldia pusilla (Michx.) Pers. (Hsr) Infrequent in feather moss or
Sphagnum mats in spruce or larch forests or in sedge mats around marly
lakes. Collected in flower June 23-July 10; in mature fruit August 13.
Mile 70, 4252; mile 42.5 N, 6922.

Zygadenus elegans Pursh. (Gb) Infrequent in loamy or sandy soil in
pine, spruce, or poplar woods, in residual soil over limestone, and in
disturbed peaty or sandy soil. The plants are commonly more robust
and taller and have longer and more branched panicles in disturbed areas
than in adjacent undisturbed ones. Collected in flower from June 26 to
July 15, in maturing fruit on August 13. Mile 64, 4267; mile 103 N, 7721.

IRIDACEAE

Sisyrinchium montanum Greene. (Hs) Rare to locally frequent in
Agropyron-Muhlenbergia or Calamagrostis grassland, in residual soil
over limestone, and on sandy shores. Collected in flower from June 24
to July 8, in fruit from July 19 to August 11. Mile 17 N, 4178; Kakisa
Lake, 5647; Mackenzie River, 8238.

ORCHIDACEAE

Calypso bulbosa (L.) Oakes. (Gst) Rare in moist rich forests. Found
in bloom and with half grown fruits as early as June 14 and with nearly
mature fruits on July 13. Kakisa River, 4554; mile 26.5 N, 6647; mile
66.6 S, 6808

Corallorhiza trifida Chat. (Grh) Rare in moderately rich to rich
forests. Found in bloom as early as June 19 and with half mature fruits
on July 18. Mile 80, 5022; mile 42.5 N, 6929.

Cypripedium calceolus L. var. parviflorum (Salisb.) Fern. (Grh) Rare
in rich woods or boggy areas, in peaty or marly soil. Collected in flower
from June 19 to June 27. Kakisa Road, 4712; mile 119.4 N, 7102

Cypripedium guttatum Swartz. (Grh) Rare in rich spruce forests.
Collected in flower from July 4 to July 15. Kakisa River, 5353; mile
66 S, 7274.

Cypripedium passerinum Rich. (Grh) Rare in rich spruce forests.
Collected with flowers si half mature fruits in mid-July. Kakisa
River, 5354; mile 126 N, 7

Habenaria hyperborea i R. Br. (Grt) Rare in rich woods, musKegs,
sedge meadows, and marl deposits. In flower from June 27 to July 16,
and with young fruit at the latter date. Mile 52, 5368; mile 72 N, 7642.

Habenaria obtusata (Pursh) Rich. (Grt) Infrequent in rich woods
and muskegs. In flower from mid-June to mid-July; with immature
fruit in mid-July. With Orchis rotundifolia, the most often encountered
orchid of the region. Mile 80, 5014; mile 53.8 S, 6746; mile 42.5 N, 6916.

Orchis rotundifolia Banks. (Grt) Infrequent in moderately rich to

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rich woods, usually of spruce. Collected in flower from June 23 to July
24, in immature fruit on July 24. This and Habernaria obtusata are
the most often encountered orchids of the region. Mile 74, 3066; mile
60.5 N, 7016; mile 66 S, 7998.

Spiranthes romanzoffiana Cham. (Grt) Rare in rich woods and mus-
kegs. In flower during July; with immature fruit on July 24, Kakisa
River, 5344; mile 35 N, 7533; mile 66 S, 8001.

SALICACEAE

where balsam poplars 12 to 14 inches DBH are found. Satisfactory incre-
ment borings could not be obtained from these trees because of heartrot.
With half mature fruit on June 21. Mile 11, 4826; mile 70.5 N, 6901.
Populus tremuloides Michx. (Ms) Frequent to common in woods,
especially on drier uplands. With Populus balsamifera, it is character-
istic of burned over areas. A tree 51 feet high and 8.7 inches DBH show-

kenzie River, 9047; mile 34 S, 6740; mile 101 N, 7084
Salix brachycarpa Nutt. (N) Occasional in marl deposits and on marly
shores. Shrub to 3 feet tall. Fruit matures in July. Mile 110.5 N, 6838.
Salix calcicola Fern. et Wieg. (N) Seen only once, in spruce-larch
27

140

Salix glauca L. (M) Infrequent to frequent in pine or spruce forests,
in marshes, at edge of grasslands, in marl deposits, along shores, in
thickets, and in shallow residual soil over limestone. The most com-
mon willow of the highway region, Salix glauca is also the willow most
persistent in disturbed areas. It usually grows 3 to 6 feet tall, although
specimens 10 feet high are characteristic at edges of sinkholes in a white
spruce-jack pine forest at mile 121.3 N. Fruit matures from mid-June
through mid-August. Mile 951, 4294: mile 2.5 S, 6662; mile 110.5 N, 6827.

Salix lasiandra Benth. var. lancifolia (Anderss.) Bebb. (M) Seen only
on the beach of Kakisa Lake near the outlet into Kakisa River, 4612,
4613, where locally it is a characteristic woody plant, and on a gravel
bar in the Kakisa River, 5183. At the latter site, this willow grows about
25 feet tall. In young fruit and with old staminate flowers on June 21.

Salix maccalliana Rowlee. (M) Infrequent at edge of grasslands, in
marshes, in thickets, and along shores. Attains 10 feet in height. With
mature fruit from mid-June to mid-July. Mile 54, 4720; mile 72 N, 8266.

Salix myrtillifolia Anderss. (N) Infrequent in forests, especially of
black spruce, along shores, in marshes, and at edges of grasslands.
Typically a depressed shrub, or even prostrate, less than 16 inches tall,
put occasionally attaining 6 feet in height. Fruit matures mid and late
June. Mile 11, 4831; mile 23.8 N, 6643; mile 66 5, 6798.

Salix padophylla Rydb. (S. pseudomonticola Ball) (M) Seen only
once, on peaty shore of lake, mile 70.5 N, 6899. With dehisced fruit on
June 22.

Salix pedicellaris Pursh. (N) Infrequent to rare in shrub zones around
marly lakes, in sedge mats around both marly and muck bottom lakes,
and in birch-willow thickets. Mature fruit in late June and early July.
Mile 54, 5072 (var. tenuescens Pursh); mile 86.5 N, 6877 (var. hypoglauca
Fern.); mile 57.6 S, 7246 (var. hypoglauca Fern.).

Salix petiolaris J. E. Sm. (N) Rare at edge of grasslands. Mature
fruit in mid-June. Mile 16.5 N, 6633.

Salix planifolia Pursh. (M) Infrequent to locally common on shores
and in marshes. Especially characteristic on the flood plain of Kakisa
River below Lady Evelyn Falls where, with other willows and with
Alnus tenuifolia, it forms the dominant vegetation, attaining 25 feet in
height and 4 inches in stem diameter. Fruit matures mid-June. Kakisa
River, 4527; mile 4.2 N, 6603; mile 2.5 S, 6661a.

Salix pyrifolia Anderss. (N) Rare in peaty soil on crystalline out-
crops, in black spruce forests, and in birch-willow thickets. Shrub to
6 feet high. Fruit matures in late June. Mile 4.2 N, 6616; mile 4.7 S, 6703.

Salix reticulata L. (Ch) Local, usually growing in moss mats or in
peat, in spruce forests. Mature fruit in July. Kakisa River, 4604; mile
126 N, 7770.

Salix scouleriana Barratt. (N) Seen only once, in marsh along Stagg

141

River, 7963. With dehisced fruits on July 23.

Salix serissima (Bailey) Fern. (N) Rare to locally frequent at edges
of grasslands, along shores, and in shrub zones and sedge mats around
marly lakes. Fruit matures mid-July to mid-August. Mile 41.5, 5418;
mile 72 N, 8270; mile 46 S, 9261.

MYRICACEAE

Myrica gale L. (N) Frequent to rare along shores, in Sphagnum or
feather moss hummocks in spruce forests, in sedge and shrub zones
around marly lakes, and in marshes. Flowers appear in mid-June, and
fruit ripens in mid-August. Mile 64, 4268; mile 23.8 N, 6641; mile 2.5 S,
6663.

BETULACEAE

Alnus crispa (Ait.) Pursh. (M) Characteristic understory plant in
white spruce and more mesic jack pine forests; frequent in peaty de-
pressions on crystalline outcrops; infrequent to dominant locally along
peaty or gravelly shores. Shrub to 10 feet tall. Kakisa River, 4524; mile
2.4 8S, 6664.

Betula occidentalis Hook. (M) Rare at edge of grasslands and in
birch-willow thickets, Attains 10 feet in height and 2 inches in stem
diameter. Mile 16 N, 5035.

Betula papyrifera Marsh. (Ms) Infrequent to frequent on limestone

142

brown bark, is perhaps var. commutata (Regel) Fern. Mile 25, 4942;
mile 19.5, 5603; mile 2.5 S, 6685.

URTICACEAE

Urtica dioica L. var. procera Wedd. (Hpr) Rare in disturbed soil
Mile 30, 5525; mile 78.3 N, 6893.

SANTALACEAE

Geocaulon lividum (Rich.) Fern. (Grh) Rare, though sometimes com-
mon locally, in sandy soil or in lichen or moss mats in pine or spruce
forests and in peaty depressions on crystalline outcrops. The flowers,
which are greenish yellow and often purple tinged, bloom in mid-June;
the orange or orange-red fruits mature in August. Mile 80, 5018; mile
2.7 S, 6693; mile 125.1 N, 8042.

POLYGONACEAE

Polygonum achoreum Blake. (Th) Local in disturbed soil. Enter-
prise, 4003; Mackenzie River, 6010; mile 8.6 N, 8249.

Polygonum amphibium L. (HH [Grh}) Local in marshes, along shores,
and in shallow water. Four miles northeast of Fort Providence, 4036
(var. stipulaceum [Coleman] Fern.); Kakisa Lake, 5660 (var. stipula-
ceum forma fluitans [Eat.] Fern.); Kakisa Lake, 5668 (var. stipulaceum
forma hirtuosum [Farw.] Fern.); mile 35 N, 7417 (var. stipulaceum forma
simile Fern.); mile 20.7 8, 7904 (var. stipulaceum).

Polygonum aviculare L. (Th) Local in disturbed soil and along rocky
and sandy shores. Enterprise, 4001; mile 17.5 S, 7887; mile 62 N, 8188.

Polygonum coccineum Muhl. (HH [Grh]) Seen only once, shore of
Hay River, near mile 49, Mackenzie Highway, 4362.

Polygonum convolvulus L. (Th) Seen only once, in weedy area, Fort
Providence, 9075.

Polygonum lapathifolium L. (Th) Local in disturbed soil or along
sandy and rocky shores. Number 9451 is best referred to var. lapathi-
folium; all other collections are var. salicifolium Sibth. Kakisa Lake,
5638; Mackenzie River, 6002; Prosperous Lake, 9451

Polygonum viviparum L. (Gst) Local in moss mats or peaty soil in
spruce forests, mostly black spruce but occasionally white. In flower in
mid-July. Mile 57.5, 5214; mile 119 N, 7746a; mile 66 5, 8002.

Rumex maritimus L. var. fueginus (Phil.) Dusen. (Th) Rare in
marshes, in mar] deposits, and along shores. Kakisa River, 5501; Yellow-
knife, 8343; mile 39.7 N, 8412.

Rumex mexicanus Meisn. (Hs) Rare in disturbed soil. Seen only at
Enterprise, 9049a, and Fort Providence, 9073.

Rumex occidentalis Wats. (Hs) Infrequent in marshes and in dis-
turbed moist soil. Mile 53, 5743; mile 93.3 N, 8258; mile 47 5, 8312.

143

CHENOPODIACEAE

Atriplex patula L. (Th) Local in disturbed soil, All collections made
along the highway are of var. patula. Mile 60 N, 8407.

xyris amaranthoides L. (Th) Seen only once, in disturbed sand,
Enterprise, 9042a (in herb. DAO).

Chenopodium bdberlandieri Moq. var. zschackei (Murr) Murr. (Th)
Local in disturbed soil. Mile 21, 6152; mile 42 S, 7938.

Chenopodium capitatum (L.) Asch. (Th) Local in disturbed soil;
seen also once on gravelly shore of Kakisa River. Kakisa River, 5746;
mile 32 N, 6983; mile 11.4 S, 7836.

Chenopodium glaucum L. var. salinum (Standl.) Boivin. (Th) Local
in disturbed areas and along gravelly shores. Kakisa River, 5488; mile
32 S, 8350.

Chenopodium hybridum L. var. gigantospermum (Aellen) Rouleau.
(Th) Seen only once, in gravel of road bed, mile 60, 6069.

Chenopodium rubrum L. (Th) Local in disturbed soil. Mile 26.9 S,
7910; mile 71.5 N, 8177.

CARYOPHYLLACEAE

Arenaria capillaris Poir. (Ch) Rare in shallow residual soil over
limestone or in sand in pine woods, becoming locally frequent in dis-
turbed sand. Our plants are glabrous in the inflorescence and so are
var. capillaris. Comes into bloom in late J une; fruit matures from mid-
July to late August. Mile 96.5 N, 6872.

Arenaria dawsonensis Britton. (Ch) Rare in sandy soil in upland
woods, along gravelly shores, in mar] deposits, and in shallow residual
soil over limestone, becoming locally frequent in disturbed soil. In
flower the latter half of June and in early July; fruit matures from
early July into August. Mile 10, 4815; mile 96.5 N, 8162.

Arenaria humifusa Wahl. (Ch) Seen at only 3 sites, at each of which
only a few plants could be found. In marl deposit, mile 110.5 N, 7096;
in black spruce-feather moss forest, mile 72 N, 7647; disturbed moist
sand, mile 110 N, 9426. Collected as early as June 27 with both flowers
and mature fruits.

Arenaria lateriflora L. (Hpr) Rare along gravelly shores, in marl
deposits, and in disturbed soil. Begins to flower in mid-June; fruits
mature about 2 weeks after flowering. Kakisa River, 5485; mile 4.2 N,
6621.

Arenaria rubella (Wahl.) Sm. (Ch) Rare in shallow residual soil over
limestone. In flower mid-June to early July; fruits mature as early as
July 5. Mile 12, 4843; mile 66 S, 6776; mile 96.5 N, 6870.

144

Cerastium nutans Raf. (Th) Rare in disturbed sandy soil. Collected
with both flowers and mature fruits on July 1. Kakisa River, 4900.

Melandrium ostenfeldii Porsild. (Hs) Rare in marl deposits and on
rock outcrops, both limestone and crystalline, where it grows in crevices
and in shallow soil. Flowering begins in early June; some fruits have
matured by mid-June. Mile 20, 4885; mile 8.8 S, 6711; mile 110.5 N, 6830.

Silene menziesii Hook. (Hpr) Rare along rocky shores and in dis-
turbed soil. Comes into flower in late June and continues until about
August 1; fruits mature from July 10 until frost. Kakisa River, 4799;
mile 110 N, 7305; Mackenzie River, 8201.

Stellaria calycantha (Ledeb.) Bong. (Hpr) Locally frequent in dis-
turbed peaty or sandy soil, where it forms conspicuous yellowish-green
loose mats. Begins to flower about mid-June, and to fruit in late June,
but continues much of the summer. Mile 6, 5951; mile 2.5 S, 6667; mile
32 N, 6980.

Stellaria crassifolia Ehrh. (Hpr) Local along gravelly or mucky shores,
in marshes, and in sedge mats; somewhat more common in disturbed
areas. In flower from mid-June until early or mid-August; mature
fruit by mid-July. Mile 11, 4822; mile 93.8 N, 6638; mile 20.5 5S, 7177.

Stellaria longifolia Muhl. (Ch) According to Gleason (1958), Stellaria
longifolia is “. . . closely related to the . . . northern and montane S.
longipes, from which it was probably derived and with which it may be
conspecific. The only character by which the two may be finally dis-
tinguished is the inflorescence, branched divaricately in the former
and ascendingly in the latter.” Of my 39 collections from the highway
region that are referrable to the S. longifolia-S. longipes complex,
many are intermediate between these two taxa in all characters and
could as well be placed in one as the other. Perhaps, as Gleason sug-
gests, S. longifolia and S. longipes may indeed be conspecific. The speci-
mens cited below are considered to be “typical” S. longifolia but in
reality are the extremes at one end of a series, at the other end of
which is “typical” S. longipes. Rare in moist grassy places and thickets.
In flower in late June and in fruit soon thereafter, continuing through
much of the summer. Kakisa River, 5328; mile 39.9 N, 8261; mile 48.1 S,
8370.

Stellaria longipes Goldie. (Ch) The taxonomy of the S. longipes com-
plex is in a state of monumental uncertainty. Porsild (1955) wrote: “It
seems doubtful if the taxonomy of this complex .. . group can be satis-
factorily cleared up except by close study, under controlled conditions,
of material grown from seed or from transplants.” Until such elucida-
tion comes about, it seems futile to try to recognize segregates from
this complex, at least in the material collected by me along the Yellow-
knife Highway. Of these specimens, some (e.g., 5362) have sepals that
are ciliate along nearly all the margin and thus appear to be
ciliatosepala Trautv.; others have a few to very few cilia on some of

145

the sepals of some of the flowers; and still others have eciliate sepals.
Rare to locally frequent in grasslands; otherwise seen only in disturbed
soil. Flowers throughout much of the summer, beginning in mid-June;
mature fruits appear as early as late June. Kakisa River, 4567; mile 16.9
S, 6717; mile 96.5 N, 6874.

Stellaria media (L.) Cyr. (Th) Seen at only one site, garden weed,
Yellowknife, 9181.

NYMPHAEACEAE

Nuphar variegatum Engelm. (HH) Common in lakes in the Canadian
Shield section. This species occupies a distinct zone, the “floating stage,”
in succession around these lakes. It is by far the dominant plant of the
“floating stage,” although species of Potamogeton with floating leaves
and Polygonum amphibium may occasionally occur with it. In full
flower in late June to early July; in almost mature fruit by mid-
August. Mile 12.9 S, 7141.

Nymphaea tetragona Georgi ssp. leibergii (Morong) Porsild. (HH)
Seen only once, in a muck bottom lake, among Nuphar, mile 35.5 Ss,
8328. In full flower on July 30.

CERATOPHYLLACEAE

Ceratophyllum demersum L. (HH) Not seen by me, but collected by
Ray Murdy, United States Fish and Wildlife Service, in a 40 acre pond,
mile 38.5 S, 132A (in Herb. LAF).

RANUNCULACEAE

Actdea rubra (Ait.) Willd. (Grh) Rare in white spruce forests along
the Kakisa and Mackenzie rivers. The red fruit matures in late August.
Mackenzie River, 5023; Kakisa River, 5444,

Anemone multifida Poir. (Hs) Infrequent to rare in prairies, in

in marl deposits. Flowering begins in late June and continues through-
out July; earliest fruits mature in late July. In our material the glossy

146

white sepals are sometimes, but not always, bluish tinged at the base
outside. Mile 20, 4892; mile 66 S, 6794; mile 44 N, 6912.

Anemone patens L. var. wolfgangiana (Bess.) Koch. (Hs) Local in
open jack pine woods or in shallow residual soil over limestone. Flowers
not seen by us (except one that bloomed, abnormally, on August 9);
fruits mature as early as June 25. Mile 103.5 N, 6844.

Aquilegia brevistyla Hook. (Hs) Rare in jack pine and in white spruce
woods; becoming more common in disturbed soil. Main flowering sea-
son from about June 24 to mid-July; fruits mature as early as July 5.
The petals are yellowish-white; the spurs and sepals are purple-blue.
Mile 54, 4722; mile 110 N, 7316.

Caltha natans Pall. (HH [Hs]) Local in shallow water or in mud or
muck on shores, more common in disturbed situations. Most frequently
seen in Canadian Shield section. Flowering begins in mid-June and
continues until mid-August; fruits mature as early as mid-July. Mile
10.2, 5577; mile 16.9 S, 6716.

Caltha palustris L. (Hs) Seen only once, in marshy spot, wooded,
along cold swift stream, mile 16, 4866. In full bloom on June 28.

Ranunculus aquatilis L. var. eradicatus Laestad. (HH) Seen only
once, in mud and shallow water, Stagg River, mile 49 S, 7241. In flower
on July 3.

Ranunculus circinatus Sibth. var. subrigidus (Drew) L. Benson. (HH)
Rare in still shallow water, to 10 inches deep, muddy or sandy bottom.
With flowers and mature fruits on July 20. Kakisa River, 5658; mile
11.4 S, 7837.

Ranunculus cymbalaria Pursh. (Hsr) Seen only once, beach of Pros-
perous Lake, 9194. Mature fruit on August 14.

Ranunculus gmeliniti DC. (HH) Locally frequent in shallow water
or on muddy, peaty, or sandy shores, becoming more common in dis-
turbed areas. Flowering starts in mid-June and continues until early
August; fruits mature as early as July 3. Achenes produced in great
abundance. Three varieties of this species, as noted below, occur along
the highway. Mackenzie River, 6018 (var. limosus [Nutt.] Hara); mile
16.9 S, 6715 (var. hookeri [D. Don] Benson); mile 78.3 N, 689la (var.
hookeri [D. Don] Benson); mile 57.6 S, 7250 (var. gmelinii).

Ranunculus lapponicus L. (Grh) Rare in moss mats in shrub zones
around marly lakes, in Ledum thickets, or in spruce forests. With
flowers and half mature fruit on June 28; fully mature fruit on July 23.
Mile 50, 5396; mile 127 N, 7108.

Ranunculus macounii Britton, (Hsr) Local along gravelly, sandy,
muddy, or peaty shores, or in marshes; more common in disturbed soil.
With flowers and half mature fruits in early July; mature fruits from
mid-July on. Kakisa River, 4991; Mackenzie River, 5052; mile 25.5 N,
7518; Prosperous Lake, 9237.

Ranunculus reptans L. (Hsr) Rare along muddy or sandy shores.

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Comes into flower the first week of July; with mature fruit on July 22.
Of the collections cited below, 8216 and 9197 are var. reptans; the others
are var. ovalis (Bigel.) Benson. Kakisa River, 5002, 5635; Mackenzie
River, 8216; Prosperous Lake, 9197.

Ranunculus sceleratus L. (Th) Infrequent to rare in shallow water
or along gravelly, sandy, or muddy shores, in marl deposits, and in
marshes; more common in disturbed areas. Flowering begins in early
June; earliest achenes mature in mid-June. Mile 7, 4811; Mackenzie
River, 5053; mile 35 N, 6653; mile 25.7 S, 7194.

Thalictrum venulosum Trel. (Hsr) Rare to locally common in prairies
and along gravelly shores. In flower in late June and early July; fruit
matures the latter half of July. A common forb in Calamagrostis and in
Agropyron-Muhlenbergia grasslands at mile 14 N to 17 N. Kakisa River,
4729; mile 16.5 N, 5037.

FUMARIACEAE

Corydalis aurea Willd. (Hs) Seen only in disturbed soil; sometimes
common where found. The golden yellow flowers begin to appear in
early June and bloom throughout the summer. Fruits mature as early
as the end of June. Mile 9, 4012; mile 35 N, 6654

Corydalis sempervirens (L.) Pers. (Hs) Locally frequent in disturbed
sandy or loamy soil; seen in seemingly undisturbed situations only in
peaty depressions on crystalline outcrops, where it is rare, The pink
flowers, orange-yellow tipped, begin to bloom the second week in June;
fruits mature as early as July 1. Mile 66, 0250; mile 35 N, 6660; mile
17 8, 7164

CRUCIFERAE

Arabis divaricarpa A. Nels. (Hs) Infrequent in disturbed clay soil;
rare in shallow residual soil over limestone. Collected in flower and
young fruit on June 28. Mile 12, 4859; mile 17 S,

Arabis drummondii Gray. (Hs) Rare in Calamagrostis grassland. In
flower and nearly mature fruit on July 8. Mile 13-14 N, 5029

Arabis hirsuta (L.) Scop. var. pycnocarpa (Hopkins) Rollins. (Hs) Rare
in prairies, on gravelly shores, and in marl deposits; becoming more
common in disturbed soil. In flower from mid-June to mid-July; earliest
fruits mature about July 20. Mile 17 N, 4188; mile 5, 5145; mile 5.5 S, 7805

some of the material is close to var. holboellii. Enterprise, 3899; mile
69.6 S, 6817; mile 96.5 N, 8166.

Barbcnee orthoceras Ledeb. (Hs) Rare along muddy or mucky shores,
more frequent in disturbed areas. Comes into flower in mid- June;

148

fruits mature as early as mid-July. Noted only in Canadian Shield
section. Mile 14 S, 7143.

Brassica campestris L. (Th) Rare in disturbed soil. With flowers and
half mature fruits on July 28. Kakisa Road, 6103; mile 39.5 N, 8260.

Braya humilis (C. A. Meyer) Robinson. (Hs) Rare in marl deposits.
Collected with flowers and half mature fruits on July 15. Sometimes ap-
parently triennial instead of biennial. Mile 110 N, 7732.

Capsella bursa-pastoris (L.) Medic. (Th) Rare in disturbed soil. Fruits
mature as early as July 9. Enterprise, 4004; Fort Providence, 5039

Cardamine parviflora L. var. arenicola (Britt.) O. E. Schulz. (Hs)
Rare in disturbed soil. With flowers and young fruits on June 18. Mile
15.1 S, 6714.

Cardamine pensylvanica Muhl. Infrequent on gravelly shores
Kakisa River; noted nowhere else. Flowering over by mid-July; fruits
mature as early as July 12. Kakisa River, 5198.

Descurainia richardsonii (Sweet) O. E. Schulz. (Th) Rare in disturbed
soil. With flowers and half mature fruit on July 16. Mile 45, 5399; mile
88.5 N, 7362; mile 53.9 5, 7970.

Descurainia sophia (L.) Webb. (Th) Locally frequent in disturbed soil.
With mature fruit as early as July 13. Enterprise, 3897; mile 3 S,

Draba cinerea Adams. (Ch) Rare in crevices on limestone Butanne
Flowers almost gone, and some fruits half mature, on June 19. Mile 66 S,
Gri

Draba lanceolata Royle. (Ch) Rare in shallow residual soil over lime-
stone; becoming locally more common in disturbed areas. With flowers
and half mature fruits on June 29; fruits mature as early as July 15.
Mile 20, 4884; mile 96.5 N, 8163

Erysimum cheiranthoides L. (Hs) Rare along gravelly and sandy
shores; more common in disturbed soil. Comes into flower in late June;
fruits mature in late July. Kakisa River, 5456; Mackenzie River, 8214;
mile 30 S, 8337.

Erysimum inconspicuum (Wats.) MacM. (Hs) Seen only once, on
open, steep slope above Kakisa River below ford, 5206. With half mature
fruits on July 12.

Lepidium bourgeauanum Thell. (Hs) Rare in disturbed soil. Comes
into flower in late June; fruits mature beginning mid-July. Mile 51,
5380; mile 16 S, 7155; mile 39.7 N, 7408

Lepidium densiflorum Schrad. (Hs) Rare in disturbed soil. Collected
with flowers and half mature fruits as early as June 24. Kakisa Road,
4972; mile 59.2 N, 7389

Lesquerella arctica (Wormsk.) Wats. var. scammanae Rollins. (Ch)
Rare in crevices in limestone outcrops. Flowering begins in mid-June;
fruits mature as early as July 4. Mile 66 S, 6773, 7284

Rorippa crystallina Rollins. (Hs) Rare in Carex marshes; found only

149

from mile 16.5 N to mile 35 N. The collections from the highway region
formed the basis of the description of Rorippa crystallina as a new
species by Dr. Rollins in Rhodora 64:324-327, 1962. Mile 23.8 N, 6637.
Rorippa islandica (Oeder) Borbas. (Hs) Infrequent along sandy,
gravelly, muddy, or mucky shores; becoming more common in dis-
turbed soil, especially in low areas at roadside. Comes into flower about
mid-June; flowering plants can be found as late as mid-August; fruits
ture as early as mid-July. Our plants have the upper leaves merely
dentate (sometimes rather deeply so) rather than pinnatifid and so do

these two extremes make the recognition of fernaldiana and hispida
rather arbitrary and of little significance, at least in the highway region.
Mile 30, 5526; mile 82 N, 7054; mile 4.8 S, 7118.

Subularia aquatica L. (Th) Rare in shallow, clear, still water of lakes,
sandy bottom. Noted only at Prelude Lake, 9215, and Prosperous Lake,
9232. In Prelude Lake it was associated with Eleocharis acicularis and
Isoetes echinospora var. braunii; in Prosperous Lake, with Eleocharis
acicularis and Limosella aquatica. With late flowers and all stages of
fruit on August 15

Thlaspi arvense L. (Th) Rare in disturbed soil. In mature fruit as
early as July 9. Enterprise, 4000; Fort Providence, 5040.

DROSERACEAE

Drosera anglica Huds. (Hr) Rare in marl in shallow water and among
sedges at edge of marly lakes. Comes into flower in mid-July; collected
with some mature fruits on August 8. Mile 50, 5393.

Drosera rotundifolia L. (Hr) Rare on Sphagnum mounds in black
spruce forests and in marly bogs. Comes into flower in mid-July; not
collected in fruit. Mile 64, 4264; mile 66 S, 8012

SAXIFRAGACEAE

Mitella nuda L. (Hsr) Infrequent in mats of feather moss in spruce
forests. The yellowish-green flowers appear in mid-June; flowers may
be found until about July 10, when early fruits are mature. Mile 80.5,
5012; mile 42.5 N, 6932.

Parnassia multiseta (Ledeb.) Fern. (Hs) Infrequent on rocky or

e

n
spruce woods. Comes into flower about mid-July and continues until
at least mid-August; fruits oop as early as August 10. Mile 70, 4248;
mile 11.4 S, 7835; mile 65.6 N,

Ribes glandulosum Grauer. na Rare along rocky shores, in peaty
depressions in crystalline outcrops, and in thickets. In full flower in

150

mid-June; the petals are rose, and the sepals are white or yellow-white
with a rose tinge. The bright red fruits, in more or less erect racemes,

mature in early to mid-July. This is a sprawling or prostrate plant
whose leaves, crushed, have the odor of skunk. Mile 6, 5946; mile 2.7 5S,
6690; mile 36 N,

Ribes hudsonianum Rich. (N) Infrequent to frequent along rocky
shores, in thickets, and in black spruce forests. In flower the latter half
of June; the glaucous blue-black, unpleasant tasting fruits, in erect
to drooping racemes, mature late in July and early in August. Ribes
hudsonianum is the most common species of the genus along the high-
way; it is the one most likely to be seen in disturbed soil. Number 4560,
cited below, with its glabrous floral bracts, bud scales, leaves, and
inflorescence, has been described by me as a new form (in Canad.
Field.-Nat. 75:117. 1961). Kakisa River, 4560 (forma glabrum); mile
5.7 5S, 7809.

Ribes lacustre (Pers.) Poir. (N) Rare in rich spruce or pine forests
and along rocky shores. The leaves have a pronounced and distinctive
sheen both above and below that was seen on no other member of the
genus in the highway region. The flowers, with their rose madder sepals
and yellow-white petals, are in full bloom in the latter half of June;
the fruits mature in late July and early August. Mile 10, 4820; mile
66 S, 8018.

Ribes oxyacanthoides L. (N) Infrequent along gravelly shores, in
shallow residual soil on limestone outcrops, in prairies, and in peaty
depressions on crystalline outcrops. In flower the second half of June.
The blue-black berries mature in late July and early August and are
the best tasting of any Ribes fruit along the highway. The flowers have
greenish white sepals and white (sometimes pink tinged) petals. Kakisa
River, 4505; mile 21.3 S, 6721; mile 80.8 N, 7046.

Ribes triste Pall. (N) Infrequent to rare in Populus or spruce forests
or Salix-Alnus thickets along shores, or in spruce forests on limestone

sistent; the translucent bright red fruits mature in late July and early
August. Kakisa River, 4533; mile 61.8 S, 6766.

Saxifraga aizoides L. (Ch) Seen at only one site, in seepage area
around cold spring, mile 122.6 N, 8076. In bloom and with half mature
fruits on July 25. The petals are yellow, but dotted with orange.

Saxifraga tricuspidata Ro (Ch) Infrequent to locally common,
forming mats to eet across, on rock outcrops, both limestone and
crystalline. In flower from mid-June to early July; fruits mature by
mid-August. The petals are white, and have either orange-red dots,
orange dots and red dots in about equal numbers. Mile 12, 4853; mile
2.8 S, 6683; mile 103.5 N, 6860.

151

ROSACEAE

Amelanchier alnifolia Nutt. (N) Infrequent to frequent on rocky
shores and ridges, in shallow residual soil over limestone, and in white
spruce or pine woods. Comes into flower about June 20 and continues for
about 7 to 10 days; the dark red fruits, “saskatoons,” prized by Indians
and Whites alike, mature from mid-July to early August. Fort Provi-
dence, 4225; mile 12, 4854; mile 101 N, 7085; Stagg River, 7239.

Dryas drummondii Rich. (Ch) Local in crevices in crystalline and
limestone outcrops. Coilected in flower on June 19; past fruiting on
August 8. The flowers have erect yellow petals and a green calyx that
is covered with purple gland-tipped hairs. Mile 25.1, 6150; mile 66 S,
6770; Yellowknife, 9186.

Dryas integrifolia Vahl. (Ch) Local in crevices on limestone outcrops,
in marl deposits, and in sandy or peaty soil in spruce or pine woods.
Collected in flower on June 19-24, but plants past flowering and in
young fruit were also collected on June 19; fruits mature during the
second half of July. Kakisa Road, 4695; mile 66 S, 6774; mile 110.5 N,
6829.

Fragaria virginiana Duch. var. terrae-novae (Rydb.) Fern. et Wieg.
(Hrr) Infrequent to locally frequent in shallow residual soil over lime-
stone, in pine or white spruce forests, and along shores, becoming more
common in disturbed soil. Main flowering season from about June 15
to July 1; fruits mature in late July and early August. The plants are
more vigorous and show much greater development of runners in dis-
turbed areas. Kakisa River, 4550; mile 31.9 S, 6735; mile 103.5 N, 6859.

Geum macrophyllum Willd. var. perincisum (Rydb.) Raup. (Hs) Local
in grasslands, in willow thickets, and along sandy and muddy shores.
Flowers from late June to late July; with mature fruit on August 4.
Mile 13-14 N, 5028; Kakisa River, 5196; Stagg River, 7967.

Geum triflorum Pursh. (Hrr) Locally frequent in shallow residual
soil over limestone. Collected in flower and young fruit June ; in
nearly mature fruit on July 21; past fruiting in early August. Not noted
north of the Mackenzie River. Mile 20, 4880.

Potentilla anserina L. (Hrr) Local on sandy, gravelly, or marly
shores. Collected with flowers from June 25 to July 19; past flowering
and with mature fruit on August 8. Most Potentilla anserina in the high-
way region is forma anserina, but forma sericea (Hayne) Hayek is oc-
casional, Kakisa River, 4897; Mackenzie River, 7448; mile 65.6 N, 7618.

Potentilla arguta Pursh. (Hs) Rare in grasslands and in shallow resi-
dual soil over limestone. Collected in fruit August 3-9. Four miles north-
east of Fort Providence, 4113; mile 25.2, 5974.

Potentilla fruticosa L. (N) A common shrub, generally distributed
except in truly aquatic habitats. In bloom from mid-June to frost;
fruits mature as early as August 5. One of the most conspicuous road-

152

side wild flowers. Mile 28.5, 4929; mile 8.8 S, 6710; mile 119 N, 7746.

Potentilla multifida L. (Ch) Seen only once, in crevices in crystalline
outcrops, Horseshoe Island, Yellowknife Bay, 9486. In fruit on August 30.

Potentilla nivea L. ssp. hookeriana (Lehm.) Hiitonen. (Ch) Infrequent
in crevices in limestone and crystalline outcrops. Collected in flower
from June 21 to July 2; past fruiting on August 13. Mile 20, 4879; mile
103.5 N, 6847; Yellowknife, 9175.

Potentilla norvegica L. (Hs) Infrequent along gravelly, sandy, and
marly shores; rare on peeiine outcrops; locally common in disturbed
soil at roadside. Collected with flowers and young fruits on June 30;
with flowers and mature fruits on July 20; flowering past by August 1.
Kakisa River, 5212; mile 6.1 S, 7123; mile 98.5 N. 8167.

Potentilla palustris (L.) Scop. (HH) Rare to locally common in
marshes and in sedge mats around marly and muck bottom lakes. Col-
lected in flower from June 30 to August 4; Mature fruit on August 7.
‘This species is one of the main contributors to the mat around many
muck bottom lakes in the Canadian Shield section. Four miles north-
east of Fort Providence, 4048; mile 6, 5562; mile 47 S, 8313.

Potentilla pensylvanica L. (Hs) Local in mar] deposits, in drier grass-
lands, and on crystalline and limestone outcrops. With late flowers and
mature fruits on July 21. the material seen is best referred to var.
pensylvanica, although it is somewhat variable. Mile 12, 5581; mile
39.7 N, 7567; Yellowknife, 9189.

Prunus pensylvanica L.f. (N) Local on limestone outcrops and dry
rocky slopes. In flower in late June and early July; the tasty, translu-
cent, glossy red fruits mature in late August and are much appreci-
ated by humans and bears alike. At mile 28 we observed a bear cub
who stripped several Prunus pensylvanica, Ribes oxacanthoides, and
Amelanchier alnifiolia bushes of their fruit, but who ignored completely
the abundant bear-berries (Artostaphylos uva-ursi) there. Prunus
pensylvanica, in the highway region, grows to 6 feet tall and has stems
up to 3/4 inch thick. It was observed only south of the Mackenzie River.
Mile 12, 4852.

Prunus virginiana L. (N) Local on limestone outcrops. The fragrant

northernmost known part of its range, grows to 5 feet tall. It was seen
only between mile 20 and 25. Mile 6.

Rosa acicularis Lindl. (N) common shrub, almost generally dis-
tributed in mesic and xeric habitats. Collected in flower from June
until July 10; the glossy red or orange-red fruits mature from late
July on. Mile 24, 5125; mile 11.4 S, 7830; mile 95.5 N, 8169

Rubus acaulis Michx. (Hpr) Local in grasslands, on peaty hummocks
in marshy areas, in moss mats or litter in spruce forests, and on gravelly
or marly shores. Collected in flower from June 13 to July 12; the trans-

d

153
lucent bright red fruits are ripe in early August. Mile 16, 4864; mile 61.2

Rubus chamaemorus L. (Hpr) Infrequent in moss mats or peaty soil
in black spruce forests and in peaty depressions in crystalline out-
crops. Collected in flower only from June 13 to June 17; the pale orange
fruits ae in mid-August. Kakisa Road, 4511; mile 4.2 N, 6618; mile
30 S, 833

Rubus ee L, var. canadensis Rich. (N) Infrequent along shores, on
crystalline outcrops, and in white spruce-balsam poplar woods. Col-
lected in flower from June 17 to July 18; the bright red fruits mature
as early as July 24. Mile 66, 5263; mile 2.5 S, 6684; mile 73 N, 6895

Rubus paracaulis Bailey. (Hpr) Specimens referrable here were seen
only once, along stream in pine-spruce woods, mile 16, 4863. In flower
on June 28.

Rubus pubescens Raf. (Ch) Infrequent along rocky shores, in poplar
or spruce woods, and in alder-willow thickets along rivers. In flower
from about June 20 to July 10; fruits still small and green on July 18.
Kakisa River, 4735; Mackenzie River, 7449.

LEGUMINOSAE

Astragalus alpinus L. (Ch) Observed only in disturbed soil, where
it may be common locally. The plants are prostrate and form mats up
to 3 feet across. Collected with both flowers and mature fruits in late
June. Mile 73 N, 6896; mile 66 S, 8003.

Astragalus americanus (Hook.) Jones. (Hp) Infrequent to rare in
white spruce, pine, larch, or poplar woods, becoming more common in
disturbed areas. Comes into flower in early July and continues for

about 3 weeks; collected with mature fruits as early as July 18. Kakisa
River, 5471; mile 4.7 S, 7801; mile 122.6 N, 8072.

Astragalus dasyglottis Fisch. (A. goniatus Nutt.) (Hpr) Rare in grass-
lands and along gravelly shores. The fragrant purple-blue flowers
begin to bloom in mid-June and continue for about 2 months; fruits
mature in mid-August and later. Mile 17 N, 4169.

Astragalus eucosmus Robinson. (Hp) Seen only once, in disturbed
clay at roadside, mile 4.7 S, 7800. Mature fruits on July 18.

Astragalus striatus Nutt. (Hp) Seen only once, in sandy barrens by
Yellowknife airport, 9242. With old flowers and maturing fruits on
August 16.

Astragalus tenellus Pursh. (Hp) Seen only once, gravelly shore just
south of ae River town, 6129. With flowers and almost mature fruits
on Augus

poaaes yukonis Jones. (Ch) Seen only in disturbed soil, where it
is rare. With flowers and half mature fruits in very late June. Mile 72
N, 7035; mile 8.3 S, 7138

Hedysarum alpinum L. var. americanum Michx. (Hp) Infrequent in

154

thickets along shores and in spruce or pine forests. Begins to flower in
mid-June and continues until about July 25; mature fruits collected in
early July. Kakisa River, 4994; mile 66 S, 6806.

Hedysarum mackenzii Rich. (Hp) Infrequent in pine or spruce woods,
in shallow residual soil over limestone, and in semi-open grasslands.
In flower by June 13; some fruits mature as early as July 10. Mile 54,
4724; mile 32 N, 6985.

Lathyrus ochroleucus Hook. (Hpr) Infrequent in pine or spruce
forests. In flower by June 18, continuing until mid-July, when fruits
are about half mature. Kakisa Road,

Melilotus alba Desr. (Hs) Rare in disturbed soil. Comes into flower
in very late June; mature fruits by July 27. Mile 66, 5241; mile 70.5 N,
7376; mile 5.5 S, 7804.

Melilotus officinalis (L.) Lam. (Hs) Rare in disturbed soil. Comes
into flower in very late June; mature fruits by July 27. Mile 52.5, 5366;
mile 22 N, 7438; mile 2 S, 7790.

Oxytropis campestris (L.) DC. var. varians (Rydb.) Barneby. (Ch)
Rare in crevices on limestone outcrops. Collected in mature fruit on
August 3. Mile 23.5, 5781.

Oxytropis deflexa (Pall.) DC. var. sericea T. et G. (Hs) Rare in
white spruce forests; local in disturbed soil at roadside. Beginning to
flower in late June; some fruits mature by mid-July. Kakisa River, 5472;
mile 74.5 N, 7040; mile 4.8 S, 7112.

Oxytropis splendens Dougl. (Hr) Rare in shallow residual soil over
limestone; local in disturbed soil at roadside. Collected in early flower
on July 5; in mature fruit by late July. Mile 107.5 N, 7330

Oxytropis viscida Nutt. var. hudsonica (Greene) Barneby. (Hr) Seen
only once, in disturbed sandy soil at epg mile 122.6 N, 7106. With
old flowers and maturing fruits on Jun

Vicia americana Muhl. (Hpr) eect in videkets and along shores;
local in disturbed sand or clay soil. Flowering begins in mid-June and
continues until late July; collected with dehisced fruits on July 9. Mile
50, 5113; mile 35 N, 6657

GERANIACEAE

Geranium bicknellit Britton. (Hs) Infrequent to rare on gravelly
shores and in disturbed soil in waste places. Comes into bloom about
mid-June, and late flowers may be found until mid-August; fruits
mature from mid-July on. Geranium bicknellii behaves as a typical
biennial in the highway region. Mile 511, 4293; mile 107.9 N, 8385.

LINACEAE

Linum lewisii Pursh. (Hp) Infrequent to rare in shallow residual
soil over limestone and in clayey gravelly soil along shores. Comes into
flower in early July, and by July 20, flowering is over; collected

155

in mature fruit as early as July 21 and as late as August 28. The blue
petals, on cloudless days, begin to drop at about 1:30 p.m.; on cloudy
days they persist somewhat longer. Mile 28.5, 4928.

CALLITRICHACEAE
Callitriche hermaphroditica L. (HH) In shallow, still or flowing water
or along muddy or sandy shores. Fruit maturing in mid-August and
9

Callitriche palustris L. (HH) In shallow, still or flowing water or
along muddy or sandy shores, Fruit maturing in mid-August and later.
Mackenzie River, 4241; mile 16 S, 7871,

EMPETRACEAE
Empetrum nigrum L. (Ch) Infrequent in feather moss or Sphagnum
mats in spruce forests, in peaty depressions and crevices on crystalline

outcrops, and in sand in jack pine forests or sandy barrens. In the

were half grown; the fruits mature in early August, turning glossy
black. Mile 44, 4297; mile 2.7 S, 6687; mile 72 N, 7649,
ELATINACEAE

Elatine triandra Schk. (Th) Seen only once, in mud and shallow water
at edge of roadside excavation, mile 25 S, 9286. Our plants are var.
triandra and had mature fruits in late August.

CISTACEAE

Hudsonia tomentosa Nutt. (Ch) Rare to locally common in sand in
Open pine forests or in sand barrens. In flower in late June. Hudsonia
tomentosa is especially common in the sand barrens near the Yellow-
knife airport, were locally it is common and may be the only plant or
may be associated with Vaccinium vitis-idaea and Arctostaphylos uva-
urst. Mile 95.5 N, 7069; Yellowknife, 7778.

VIOLACEAE

Viola adunca Sm. (Hsr) Seen only in disturbed gravelly or sandy
soil. Fruits mature early in July; not collected in flower. Mile 107.5

5 (bey

Viola nephrophylla Greene. (Hrr) Rare along rocky or Peaty shores
or in black spruce woods. Collected in flower June 20-24 in fruit
July 7. Kakisa Road, 4709; mile 66.6 S, 6812; Mackenzie River, 7461,

Viola renifolia Gray. (Hrr) Seen only once, in rich white spruce forest
in gorge of Kakisa River just below Lady Evelyn Falls, 4535. In flower
on June 16.

ELAEAGNACEAE

Elaeagnus commutata Bernh, (N) Infrequent to rare on rocky shores,
river bluffs, and beaches. The flowers appear in early July; they are.

156

heavily fragrant and have tan anthers, cream filaments, and sepals
that are light yellow inside and silvery-cream outside. The silvery fruits
mature in the first half of August. Fort Providence, 4226; Kakisa River,
4784.

Shepherdia canadensis (L.) Nutt. (N) Frequent in poplar, spruce, Or
pine forests, in shallow residual soil over limestone, in marl deposits,
and in peaty depressions on crystalline outcrops. The light brown-
yellow flowers begin to open in mid-June; the nauseous translucent
red fruits mature in early August. Kakisa River, 4532; mile 21.3 5,
6720; mile 110.5 N, 6824.

ONAGRACEAE
Epilobium angustifolium L. (Hpr) Rare in spruce or pine forests on
sand and in peaty depressions on crystalline outcrops, but frequent to

7165.

Epilobium glandulosum Lehm. var. adenocaulon (Haussk.) Fern.
(Hp) Infrequent in wetter grasslands, and on muddy, marly, or sandy
shores, becoming locally common in damp disturbed areas. Coming
into flower in mid-July and continuing until mid-August; fruits mature
as early as July 26. When growing en masse, the plants are conspicuous
because of their red coloration. Enterprise, 4010; Mackenzie River,
3229; mile 59.2 N, 7390.

Epilobium palustre L. var. oliganthum (Michx.) Fern. (Hpr) Rare to
infrequent in disturbed moist soil. In flower from late June until late
July; fruits mature as early as July 12. Mile 76, 6029; mile 2.5 S, 7110;
mile 61.2 N, 7598.

HALORAGACEAE

Myriophyllum exalbescens Fern. (HH) Local in shallow, still or
flowing water. Beginning to flower in mid-July and continuing to at

Myriophyllum verticillatum L. var. pectinatum Wallr. (HH) Local in
shallow, still or flowing water. With flowers and young fruits on July
6; some fruits mature by August 4. Mile 75 N, 7374; mile 7.7 S, 7819.

HIPPURIDACEAE
Hippuris vulgaris L. (HH) Infrequent in shallow, still or flowing

157

water (to about 18 inches deep) or along shores. The flowers, with their
glistening white stigmas and purple anthers, begin to open in early
June; fruits mature starting in late June. Mile 44, 4298; mile 35 S, 6743;
mile 23.5 N, 6970.

UMBELLIFERAE

Cicuta bulbifera L. (Hs) Rare to infrequent in sedge mats around
muck bottom lakes, in sedge marshes, and in low Salix thickets. Col-
lected in flower between July 20 and August 2. Mile 12.7 S, 7859; mile
61 N, 8192,

Cicuta douglassii (DC.) Coult. et Rose. (Hs) Rare in marshes and along
gravelly, muddy, or marly shores. Fruit not fully mature by August 9.
Four miles northeast of Fort Providence, 4075; Kakisa River, 6094;
mile 16 S, 7874.

Heracleum lanatum Michx. (Hs) Seen only once along the Yellow-
knife Highway, one plant, basal leaves only, on grassy muddy shore of
Great Slave Lake, mile 62.7 S, 8374. This species is frequent along shores
at the town of Hay River and along the Mackenzie Highway south of
Hay River.

Sium suave Walt. (HH) Rare in shallow, still to flowing water. Col-
lected in flower and very young fruit on July 21; with mature fruit on
August 15. Mile 33.5, 9994; mile 20.5 S, 7894.

CORNACEAE

60.5 N, 7021.
Cornus stolonifera Michx. (M) Infrequent along rocky shores, in

PYROLACEAE
Moneses uniflora (L.) Gray. (Hsr) Rare in moss mats in spruce for-
ests. Comes into flower about July 8 and continues until at least early
August; mature fruits not seen. Mile 80-81, 5024; mile 36 N, 7545.

Pyrola grandiflora Radius. (Hsr) Infrequent to rare in moss mats in:

158

spruce forests and in peaty soil in thickets. Flowering begins about June
13 and is nearly over by mid-July; collected in fruit on August 16.
Kakisa River, 4530; mile 4.3 S, 6695; mile 42.5 N, 6930.

Pyrola secunda L. (Hsr) Local in moss mats, sand, or litter in spruce
or pine forests. Collected in flower and with some mature fruits on July
10. Mile 64, 4263; mile 33 N, 7527.

for about a month; collected with mature fruit on August 14. Mile 37,
4309; mile 83.5 N, 70957.

ERICACEAE
Andromeda polifolia L. (Ch (N]) Infrequent to rare among sedges,

Arctostaphylos rubra (Reha. et Wils.) Fern. (Ch) Infrequent in moss
mats, peaty soil, or litter in spruce or poplar woods. The inconspicuous
and the translucent bright red fruits
mature in late July and early August and persist through the winter.
Mile 70, 4253; mile 66 S, 6797.

Arctostaphylos uva-ursi (L.) Spreng. (Ch) Common in undisturbed
mesic and xeric habitats throughout the region, but especially char-
acteristic as a ground cover plant on limestone and crystalline out-
crops and in spruce or pine woods on sand. In flower from mid-June
until very early July; fruits mature in August. Specimens readily re-
ferrable to var. wvd-ursi, var. adenotricha Fern. et Macbr., and var.
coactilis Fern. et Macbr. can be found growing in close proximity, and,
with them, specimens seemingly intermediate between these varieties.
Mile 17 N, 4206; mile 11.5, 4836; mile 2.5 S, 668

by July 22. Mile 64, 4270; mile 4.2 N, 6615; mile 145 5S, 7151.

Kalmia polifolia Wang. (Ch) Rare in open black spruce forests be-
tween mile 112 N and 119 N, where it grows among lichens (especially
Cladonia and Cetraria), Sphagnum and other mosses, and other low
ericads. Past flowering and with mature fruit by July 15. Mile 119 N,
7742

Ledum decumbens (Ait.) Lodd. (N) Frequent to common in peat or
moss mounds (frequently Sphagnum) in spruce forests, in peaty de-
pressions on crystalline outcrops, on peaty hummocks in boggy areas,
and in litter-covered or bare sand in open pine or spruce woods. In

159

full bloom by June 12; mature fruit by July 14. Ledwum decumbens comes
into full bloom while the flowers of L. groenlandicum are still in bud
(although nearly ready to open). Kakisa Road, 4507; mile 4.7 S, 6698b;
mile 67.8 N, 7024.

in the habitat preferences of the two species of Ledum, which frequently
can be found growing together. Ledum groenlandicum comes into full

Rhododendron lapponicum plants are notable for their stature; they
attain 30 inches (76 em) in height (see note in Canad. Field-Nat. 76: 123.
1962). Mile 66 S, 6790.

Oxycoccus microcarpus Turcz. (Ch) Rare on Sphagnum mounds in
bogs and in spruce or spruce-larch forests. Collected in flower on June
24-26, in mature fruit on August 13. Mile 64, 4271; mile 67.8 N, 7022;
mile 45 S, 7943a.

Vaccinium uliginosum L. (N) Infrequent to frequent in moss mats
or peaty soil in spruce woods, in litter-covered or bare sand in open
pine or spruce woods, in birch-willow thickets, and in peaty depressions

in moss mounds or peaty soil in spruce woods, in litter-covered or bare
sand in open pine or spruce woods, in sand barrens, and in peaty de-

brown purple the next spring. A characteristic ground-cover plant in
pine and spruce woods on sand. Mile 64, 4265; mile 2.5 S, 6678.

PRIMULACEAE
Androsace septentrionalis L. (Hr) Rare in shallow residual soil or in

mossy crevices on limestone outcrops, becoming more common and con-

August; first fruits mature in early July. In the highway region, An-
drosace septentrionalis behaves as a typical biennial, Mile 25, 4945; mile
103.5 N, 6842.

160
Dodecatheon pulchellum (Raf.) Merrill. (D. radicatum Greene) (Hr)

soil over limestone, and in sedge meadows on shores of marly lakes.
Collected in flower on June 24, and with some mature fruits as early
as July 2. Kakisa Road, 4688; mile 67.7 N, 7631.

Naumburgia thyrsiflora (L.) Duby. (HH) Local in shallow water or
along rocky or muddy shores and in Carex and Calamagrostis meadows.
Comes into flower in late June; collected in mature fruit on August

8.

Primula egaliksensis Wormskj. (Hr) Seen only once, in Betula-Myrica
thicket around marly pond, mile 113.5 N, 8120. Past flowering, and with
nearly mature fruit, on July PAS

Primula incana Jones. (Hr) Rare in grasslands, on gravelly-marly
shores, and in sedge meadows and willow copses around marly lakes.
In flower in mid-July, some flowers appearing until early August;
fruits mature from early August on. In some colonies of this species,
certain plants may have only efarinose leaves. Mackenzie River, 6000;
mile 67.7 N, 7633.

Primula mistassinica Michx. (Hr) Rare in mossy soil along streams,
in peaty soil in spruce forests, and in alder-willow thickets on lake
shores. Collected in flower from June 91 to June 28. Mile 16, 4867.

Primula stricta Hornem. (Hr) Rare in mossy clay soil in seepage
areas and in disturbed peaty or clay soil in wet places. Collected in

GENTIANACEAE

Gentianella amarella (L.) Born. ssp. acuta (Michx.) Gillett. (Gentiana
amarella L. var.) (Hs) Infrequent in disturbed soil. Flowers appear in
late July; fruits mature as early as August 18. Mile 0.5, 5941; mile 121.3

Gentianella crinita (Froel.) G. Don ssp. macounti (Th. Holm) Gillett.
(Gentiana macounti Th. Holm) (Th) Rare along gravelly sandy shores.
Comes into flower in early August. Mackenzie River, 4132; Kakisa
River, 6092.

Gentianella crinita (Froel.) G. Don ssp. raupii (Porsild) Gillett. (Gen-
tiana raupii Porsild) (Th) Rare along gravelly or marly shores. Comes

Salix-Betula thickets, and on gravelly-marly lake shores. Flowers begin
to appear in late July. The corollas vary from white to light blue, with
all intermediates between. Mile 65.6 N, 8399.

161

Menyanthes trifoliata L. (HH) Rare in marl in shallow water of marly

later; nearly full-sized fruits seen on only one plant, on July 26. Mile 42,
5407; mile 95.5 N, 7072; Yellowknife, 7780.
POLEMONIACEAE
Collomia linearis Nutt. (Th) Rare along rocky shores; locally frequent
in disturbed sandy soil. Collected in early flower in mid-July, in flower
and mature fruit in late July to mid-August. Fort Providence, 4224;
mile 26, 5782.

HYDROPHYLLACEAE

Phacelia franklinii (R. Br.) Gray. (Hs) Local in disturbed soil. Begins
to flower in mid-June, continuing through much of the summer, Some

BORAGINACEAE
Lappula echinata Gilib. (Th) Seen only once, in disturbed sand, En-
terprise, 9051a. Collected in mature fruit on August 7.
Lappula redowskii (Hornem.) Greene var. occidentalis (Wats.) Rydb.
Rare in disturbed soil. Kakisa River, 5487; mile 33 N,. 7525.

Yellowknife Highway, in spruce-poplar woods, mile 1, 7803. In full
flower on June 27, Frequent in much the same habitat along the NWT
section of the Mackenzie Highway.
LABIATAE

Galeopis tetrahit L. var. bifida (Boenn.) Lej. et Court. (Th) Seen
only once, in disturbed sand, Enterprise, 4007,

Mentha arvensis L. (Hpr) Local along gravelly or muddy shores, in
marl deposits, and in marshes, becoming more frequent in disturbed

39.7 N, 7404

Moldavica parviflora (Nutt.) Britton. (Hs) Infrequent to common in
disturbed areas. Comes into flower in mid-June; with some mature
fruits by July 10, Mile 52, 5082; mile 35 N, 6656.

162

Scutellaria galericulata L. var. epilobiifolia (Hamilt.) Jordal. (Hpr)
Infrequent to rare in marshes, along rocky or muddy shores, and in
marl deposits. Comes into flower about June 25 and continues until
late July; fruits mature from about July 11 on. Four miles northeast
of Fort Providence, 4043; mile 9.5, 5576; mile 72 N, 7025; mile 20.5 5S,
7897.

Stachys palustris L. var. nipigonensis Jennings. (Gst) Rare on gravelly
or sandy shores and in marshes. Comes into flower in late June and con-
tinues at least until early August; collected with mature fruit on August
9 Kakisa River, 5708; mile 39.7 N, 6990.

SCROPHULARIACEAE

Castilleja raupii Pennell. (Hp) Infrequent in grasslands, in shallow
residual soil over limestone, and in thickets. Collected with flowers and
nearly mature fruits on June 30, but flowers may be found throughout
the summer. Mile 17 N, 4144; mile 10, 4816; mile 31.9 S, 6738.

Euphrasia aff. subarctica Raup. (Th) Seen only once, on rocky slope
above inlet on island in Mackenzie River 1 mile west of Fort Provi-
dence, 4130. Specimens of this collection have been examined by Messrs.
Yeo and Sell (Cambridge University) who comment as follows: “Speci-
men 4130... has the general characters of E. subarctica Raup, but all
the plants in the series seen lack glandular hairs, which usually densely
cover the leaves in this species, and if they do not, are present on at

bottom, Prosperous Lake, 9233. With mature fruits on August 15.

Pedicularis labradorica Wirsing. (Hs) Infrequent to rare in moss mats
or litter in spruce or pine woods and in thickets. Comes into flower
about June 22 and continues for about 2 months; mature fruits by early
August. Mile 56, 5010; mile 125.1 N, 8045.

Rhinanthus crista-galli L. (Th) Rare in shallow residual soil over
limestone, in thickets, and along rocky shores. Flowering begins in mid-
July; mature fruits found on August 8. Mile 12, 5580; Mackenzie River,
6008.

Veronica peregrina L. var. axalapensis (HBK) Pennell. (Th) Rare on
gravelly or sandy shores. Mature fruits on July 26. Kakisa River, 5715;
Prosperous Lake, 9234.

Veronica scutellata L. (Hpr) Rare along shores. Collected with flowers
and young fruits on July 1; with mature fruits on July 9. All our plants
are forma villosa (Schum.) Pennell. Kakisa River, 5186; mile 20.5 5,
7179; mile 35 N, 7411.

OROBANCHACEAE
Boschniakia rossica (Cham. et Schl) Fedtsch. (Gp) Seen only once, in

163

white spruce forest, growing under Alnus crispa in deep mats of Hylo-
comium splendens, along Kakisa River below Lady Evelyn Falls, 4520.
Found in flower on July 16 and in mature fruit on August 27.

LENTIBULARIACEAE

Pinguicula villosa L. (Hr) Seen only once, on a Sphagnum mound in
a black spruce-larch forest, Kakisa Road, 4701. In flower on June 24.

Pinguicula vulgaris L. (Hr) Rare in moss mats in spruce forests, in
marl or in sedge mats around marly lakes, and in wet clay soil over
limestone. Collected in flower from June 26 to July 15; mature fruit
in early August. Mile 44.5, 0121; mile 72 N, 7028.

Utricularia intermedia Hayne. (HH) Infrequent in shallow water.

orm as early as July 1. This species is most often seen in marly lakes.
Mile 54, 5080; Mackenzie River, 6006; Stagg River, 7958.

Utricularia minor L. (HH) Rare in shallow water, often growing
among Utricularia vulgaris and U. intermedia. Not seen in flower or
fruit. This bladderwort is easily overlooked. We were able to find it
only in the Canadian Shield section. Stagg River, 8309.

Utricularia vulgaris L. (HH) Local in shallow, still or flowing water.
Comes into flower in late June, continuing until at least August 10; not
seen in mature fruit. Winter buds not noted until very late July and
early August. Mile 50, 5105; mile 12.9 S, 7140; mile 75 N, 7373.

PLANTAGINACEAE

Plantago major L. (Hr) Infrequent along rocky shores, becoming fre-
quent in disturbed soil. The plant of shores and the plant of waste

late June and early July; fruits mature starting in mid-August. Fort
Providence, 4228; Kakisa River, 5496; mile 60 N, 8408a.

Plantago septata Morris. (Hr) Infrequent in shallow residual soil over
limestone; in disturbed soil the plants are twice as large as in undis-
turbed situations. In full flower June 21; fruits mature in late July and
early August, Mile 28.5, 4907; mile 80.8 N, 7047.

RUBIACEAE

Galium labradoricum Wieg. (Hpr) Rare in moss mats in spruce forests
and among sedges at shores of marly lakes. Flowering begins in early
July; fruits mature starting in early August. Mile 53, 4958; mile 36 N,
(Env e

Galium septentrionale R. et. S. (Hpr) Infrequent to frequent along
rocky shores, in drier grasslands, in shallow residual soil over lime-
stone, and in pine, spruce, or larch woods, becoming somewhat more
common in disturbed areas. Flowering season extends from about June

164

24 to early August; fruits mature starting early August. This plant is
locally abundant on anthills in the Fort Providence grasslands. Enter-
prise, 3891; mile 17 N, 4184.

Galium trifidum L. (Hpr) Infrequent on gravelly and mucky shores
and in wetter grasslands, becoming more common in disturbed soil.
Collected in flower from June 24 to July 27; in fruit as early as mid-
July. Kakisa River, 4992; mile 23.5 N, 6971; mile 14 §, 7147.

CAPRIFOLIACEAE

Linnaea borealis L. var. americana (Forbes) Rehd. (Ch) Frequent in
spruce or pine forests, growing either upon moss mats, in litter, or in
sand. Comes into flower late in June; mature fruits appear in late
August. Mile 80, 5017; mile 60.5 N, 7020; Yellowknife, 9244.

Lonicera dioica L. var. glaucescens (Rydb.) Butt. (N) Rare in shallow
residual soil over limestone, usually in pine-dominated areas. The yel-

region, is a twiggy erect shrub 1 to 2 feet tall, with none of the twining
tendencies it exhibits in areas further south. Mile 56, 5010a; mile 103.5

: Ti

Symphoricarpos occidentalis Hook. (N) Seen only once, in grassland,
mile 17 N, 4141. Collected in flower on August ala

Viburnum edule Raf. (N) Frequent along gravelly shores, in poplar,
spruce, or pine woods, and in peaty depressions in crystalline outcrops.
Begins to flower about June 20, and flowering is over by mid-July; the
translucent red fruits mature in early August and are valued by some
Yellowknife people for preserves. This species may attain 5 feet in
height in the highway region. Mile 11.5, 4837; mile 21.3 S, 6724; mile
126 N, 7760.

CAMPANULACEAE

Campanula rotundifolia L, (Hsr) Rare to infrequent in shallow resi-
dual soil over limestone and in pine woods, becoming frequent to com-
mon in disturbed soils, where it is one of the conspicuous roadside wild-
flowers. Comes into bloom in very late June and continues into late
August; fruits mature as early as August 1. Enterprise, 4009; mile 66 S,
7263; mile 110 N, 7303.

Lobelia kalmii L. (Hs) Rare in wet marl among sedges at edge of
marly lakes or depressions. Comes into flower in very late July and
early August. Mile 20, 5764.

COMPOSITAE

Achillea lanulosa Nutt. (Hsr) Frequent to infrequent in drier grass-
lands, on limestone outcrops, along rocky shores, and in sand or moss
and lichen mats in spruce or pine forests, becoming common in dis-

165

turbed soil. Collected in flower from late June until mid-August;
achenes mature from mid-July on. A pink-flowered form is occasional.
This species is generally much more robust and common on disturbed
sites than in adjacent undisturbed ones. One of the conspicuous road-
side wildflowers. Enterprise, 3892; mile 82 N, 7049.

Achillea sibirica Ledeb. (Hs) Rare along rocky and sandy shores and
in disturbed soil at roadside, Collected in flower from July 9 to August
5; with maturing achenes in mid-August. Kakisa Lake, 6097; mile 16.5
N, 7483; mile 31.5 S, 7933.

Antennaria parvifolia Nutt. (Ch) Local in grasslands, on limestone
outcrops, on gravelly-marly shores, and in disturbed soil at roadside.
Flowering in July. Mile 32 N, 7424.

Antennaria pulcherrima (Hook.) Greene. (Ch) Rare in peaty soil in
thickets and spruce forests; somewhat more common in disturbed peaty
or sandy soil at roadside. In flower during late June and July. Mile 44.5,
0727; mile 119 N, 7741.

Antennaria rosea (Eat.) Greene. (Ch) Rare along sandy or muddy
shores and in disturbed soil at roadside. In flower during July. Kakisa
River, 4966; mile 88.5 N, 7361.

Antennaria subviscosa Fern. (Ch) Seen only once, in prairie, mile
17 N, 4194a. Past flowering on August 11.

Arnica lonchophylla Greene, (Hsr) Infrequent on limestone outcrops,

h
typical A. lonchophylla; one collection (mile 42.5 N, 6933) is transitional
to ssp. arnoglossa (Rydb.) Maguire. Mile 16, 4873; mile 4.3 S, 6694;
mile 96.8 N, 7078.

Artemisia biennis Willd. (Hs) Local in disturbed soil. Does not come
into flower until mid-August. Mile 66, 6106; mile 34 N, 8425.

Artemisia campestris L. ssp. borealis (Pall.) Hall et Clem. (Hs) In-
frequent to rare in shallow residual soil over limestone. In flower during
the second half of July; collected with mature achenes on August 28.
Mile 13.5, 5569; mile 96.7 N, 7965.

Artemisia frigida Willd. (Ch) Seen only once, in disturbed sand along
Kakisa Road, 9448. Several small sterile plants were all that could be
found.

Artemisia ludoviciana Nutt. var. gnaphalodes (Nutt.) T. et G. (Hsr)
Seen only once, a large clump at roadside, mile 9.5, 6117. Not yet in
flower on August 7. Collected here in 1959; observed here in 1961 and
1962

Artemisia tilesii Ledeb. ssp. unalaschensis (Bess.) Hulten. (Hpr) Local

166

in disturbed soil, especially pote shores. Comes into flower in late July.
Mile 79.5, 6021; mile 62.7 S,

Aster alpinus L. var. ae (Onno) Cronq. (Hsr) Infrequent in
shallow residual soil or in crevices on limestone outcrops. Comes into
flower in late June; mature achenes by July 21. Mile 26, 4334; mile
80.8 N, 6889.

Aster brachyactis Blake. (Th) Local along shores and in marl de-
posits; locally common in disturbed soil at roadside. Comes into flower
in very late July and early August; mature achenes in August. Mile 75
N, 8395; Prosperous Lake, 9193.

Aster ciliolatus Lindl. (Hsr) Rare in shallow residual soil on lime-
stone outcrops, in grasslands, in sandy pine-spruce woods, on marly
shores, and in peaty soil in black spruce woods; common locally in dis-
turbed soil at roadside. This species is a conspicuous roadside wild-
flower. Comes into flower in mid-July, continuing until mid-August;
with mature achenes late in August. Flowering specimens are infre-
quent and small in undisturbed situations: thought sterile plants may
abound; in disturbed soil, flowering is abundant and the plants are large
and vigorous. Mile 51, 5734; mile 13 N, 7480.

Aster falcatus Lindl. (Hpr) Seen only once, in grassland, mile 17 N,
4149. In flower on August 11.

Aster hesperius Gray var. laetevirens (Greene) Cronq. (Hpr) Seen
only once, limestone crevices, Alexandra Falls, 4365. In flower on August

ee johannensis Fern. (Hpr) Rare in marshes on shore of Great
Slave Lake, mile 62.7 S, 8373, and mile 64.6 S, 8376. In flower on
August 3.

Aster junciformis Rydb. (Hpr) Local in drier grasslands, on gravelly
or sandy shores, in marl deposits, and in sedge meadows. In flower from
mid-July to mid-August. Four miles northeast of Fort Providence,
4045; mile 6, 5953; Stagg River, 7961.

Aster pansus (Blake) Cronq. (Hp) Local in drier grasslands, in marl
deposits, and in disturbed soil at roadside. Flowers appear in late July
and early August; achenes mature from late August until frost. Four
miles northeast of Fort Providence, 4087.

Aster sibiricus L. (Hpr) Rare in sandy soil in spruce or pine woods;
frequent to common in disturbed soil at roadside. Comes into flower
in late June and continues into August; achenes mature in August.
Kakisa River, 4018; mile 107.5 N, 7331.

Bidens cernua L. (Th) Seen only once, along muddy shore of Stagg
River, 8308. Early flowers on July 30.

Crepis elegans Hook. (Hs) Seen only once, along road to Louise Falls
on Hay River, 6153. In flower and mature fruit on August

Crepis tectorum L. (Th) Rare in disturbed soil at ponies With
flowers and mature fruit on August 3. Mile 30, 6116.

167

Erigeron acris L. (Hs) Rare in marl deposits and willow thickets;
frequent in disturbed soil at roadside. In flower from June 25 to early
August; mature achenes collected August 3. Two varieties grow in about
equal numbers along the highway; var. asteroides (Andrz.) DC. (3893,
7441, 7803) and var. elatus (Hook.) Cronq. (5308, 6999, 7114). Enter-
prise, 3893; mile 16 N, 7441; mile 4.7 S, 7803; Kakisa River, 5308; mile
41.3 N, 6999; mile 4.8 S, 7114.

Erigeron compositus Pursh var. glabratus Macoun. (Ch) Rare in shal-
low residual soil or in crevices on limestone outcrops. In flower on June
21; past fruiting on July 14. Mile 23.5, 5961; mile 103.5 N, 6846.

Erigeron glabellus Nutt. var. pubescens Hook. (Hs) Rare in shallow
residual soil on limestone outcrops and in sandy pine woods. In flower
from late June until mid-July; mature achenes on July 27. Mile 28.5,
4911; mile 28 N, 9096.

Erigeron hyssopifolius Michx, (Hp) Infrequent in peaty soil or moss
or lichen mats in spruce forests and in marl deposits. Comes into
flower in mid-June; some fruits are mature by mid-July. Kakisa Road,
4569; mile 35 N, 6659.

Erigeron lonchophyllus Hook. (Hs) Infrequent in grasslands, on
gravelly-marly shores, and in marl deposits. Begins to flower in early
July; mature achenes by August 9. Some plants of E. lonchophyllus may
be only 1.8 cm. high and bear only 1 head. Kakisa River, 6081; mile
110.5 N, 7733.

Erigeron philadelphicus L. (Hs) Rare on gravelly-sandy shores. In
flower in mid-August. Kakisa River, 5307.

naphalium uliginosum L. (Th) Seen only once, in wet sand and
shallow water of Prosperous Lake, 9238. Mature achenes on August 15.

Helenium autumnale L. (Hp) Seen only in limestone crevices, Al-
exandra Falls, 4378, and in roadside ditch, 12.5 miles south of Hay River
on Mackenzie Highway, 5937. In flower and with maturing achenes on
August 15.

Mieracium umbellatum L. (Hp) Infrequent in shallow residual soil
or in crevices on limestone outcrops, in sandy pine forests, and along
rocky shores; more common in disturbed soil at roadside. The earliest
collection made, July 14, has flowers and mature fruits. Mile 24, 5629;
mile 93.5 N, 7687; mile 17.5 S, 7888.

Lactuca pulchella (Pursh) DC. (Hsr) Rare along rocky shores and in
marl deposits. Collected in flower July 7-11; in fruit July 27. Mac-
kenzie River, 7443; mile 39.7 N, 7570.

Matricaria maritima L. var. agrestis (Knaf) Wilmott. (Th) Seen only
once, disturbed soil at roadside, mile 57 N, 9105. In flower and with some
mature achenes on August 9

Matricaria matricaroides (Less.) Porter. (Th) Local in disturbed soil.
Enterprise, 3896; Yellowknife, 7855.

Petasites frigidus (L.) Fries var. nivalis (Greene) Cronq. (P. vitifolius

168

Greene) (Grh) Rare in moist disturbed gravelly soil at roadside and in
wet woods, Past flowering and with immature fruit on June 15 and June
27. Mile 7, 4810; mile 110 N, 9428.

Petasites frigidus (L.) Fries var. palmatus (Ait.) Crong. (P. palmatus
{Ait.] Gray) (Grh) Rare in sand in pine woods and in disturbed soil
at roadside. Past flowering and with immature fruit on June 18. Kakisa
River, 4562; mile 110 N, 7319.

Petasites sagittatus (Pursh) Gray. (Grh) Rare to locally frequent in
grasslands, in marshes, in birch-willow thickets, and in disturbed soil
at roadside. Past flowering and with immature fruit on June 18. Mile
17 N, 4165; Kakisa River, 4565; mile 23.6 S, 6727.

Senecio congestus (R. Br.) DC. (Hs) Infrequent to locally common
along shores, in marl deposits, in wet meadows, and in disturbed soil
of roadside ditches. Collected in flower June 30; with early mature
achenes on August 14. Mile 30, 4332; mile 14 S, 7144; mile 61.2 N, 7610.

Senecio eremophilus Rich. (Hp) Seen only once, in disturbed sandy
soil along road to ford over Kakisa River, 5327. In flower on July 15.

Senecio indecorus Greene. (Hs) Infrequent in sedge meadows, in marl
deposits, and in disturbed soil at roadside. Flowers in mid-July; nearly
mature fruit by August 4. Mile 9, 5567; mile 39.7 N, 7565.

Senecio lugens Rich. (Hsr) Local in moss mats or peaty soil in black
spruce woods; somewhat more common in disturbed peaty or sandy soil
at roadside. Collected in flower on June 23; in nearly mature fruit on
July 15. Mile 56, 4785; mile 66 S, 8010; mile 28 N, 9094.

Senecio pauperculus Michx. (Hs) Rare in grasslands; locally frequent
in moist disturbed soil at roadside. In flower during very late June and
July; immature fruits on July 27. Two of our specimens (5259, 5265),
collected at mile 66, are hybrids between S. pauperculus and S. indecorus,
according to Dr. T. M. Barkley. Kakisa Road, 5326; mile $1.5 N, 737i.

Senecio plattensis Nutt. (Hs) Collected twice, in grassland, mile 13 N-
14 N, 5031, and in disturbed soil at roadside, mile 52, 5091. In flower
July 8-10.

Senecio tridenticulatus Rydb. (Hs) Local in shallow residual soil
over limestone and in litter or sand in jack pine forests; local in dis-
turbed soil at roadside. In flower June 21; in flower and fruit July 4.
Our 1959 collections (4674, 4844, 4875, 4912, 5009, 5091) of this species
were identified by T. M. Barkley. Dr. A. E. Porsild (in litt.) considers
the Mackenzie plants to be Senecio cymbalarioides Nutt. var. borealis
T. et. G. Mile 12, 4844; mile 103.5 N, 6865.

Senecio vulgaris L. (Th) Seen only once, garden weed, Yellowknife,
9301. With mature fruit on August 1

Solidago canadensis L. var. salebrosa (Piper) Jones. (Hpr) Infrequent
to rare in grasslands and along sandy or rocky shores; locally common
in disturbed soil at roadside. In flower in late July and early August;

169

achenes not yet mature on August 11. Mile 17 N, 4190; mile 0.5, 6118;
mile 64.6 S, 8378

Solidago multiradiata Ait. (Hsr) Rare in moss mats or litter in spruce
forests; local in disturbed soil at roadside. In flower during July;
achenes nearly mature on August 3. Mile 31.5 S, 7929

Solidago spathulata DC. var. neomexicana (Gray) Cronq. (S. decum
bens Greene var. oreophila [Rydb.] Fern) (Hsr) Rare in shallow residual
soil over limestone and in sand in pine woods; considerably more common
in disturbed soil at roadside, Begins to flower in mid-J uly; with nearly
mature achenes on August 3. Mile 51, 4292; mile 98. 8 N, 7713; Yellow-
knife, 8347.

Sonchus arvensis L. var. glabrescens Guenth., Grab. et Wimm. (Hsr)
Seen only once, at roadside, mile 33 N, 9439. With flowers and very
immature fruits on August 27.

Tanacetum vulgare L. (Hs) Seen only once, waste place, mile 5 N,
8428. In flower on August 4.

Taraxacum ceratophorum (Ledeb.) DC. (Incl. T. lacerum Greene)
(Hr) Frequent in disturbed soil at roadside; rare in crevices in limestone
outcrops and in sandy pine woods. In early flower mid-June; matur
achenes by June 22. Mile 23.5, 5133; mile 35 N, 6658; mile 6.2 S, 6709.

Taraxacum officinale Wiggers. (Hr) Common in disturbed soil, es-
pecially about settlements. With mature fruits by June 24. Fort Provi-
dence, 6937.

ACKNOWLEDGEMENTS
The Yellowknife Highway sales was begun when I was on the staff of Chicago
Natural History Museum; the um financed the 1958 and 1959 field work. The

project was completed after I eel Res the staff of the University of Southwestern
Louisiana, A grant (G15904) from the National Scie Foundation supported the 1961
ork. To the Fou i

s
ment and for his efforts in behalf of the project; to Mr. an rs. B. Cooper, formerly
of Yellowknife, for their mek meme: and to ao following botanists, for their help
with the ae of various taxa: Faye K. Daily (Characeae); John W. Thomson
(lichens); H. L. Blomaise oan Sy peenene Bod Lange (certain Sphagnum);
Crum (bryophytes); George W. Argus Cage E. Porsild (Melandrium)

e

Benso r D
crystallina) ; Russell ee Basse Maguire (certain Arnica); Arthur Cronquist
(certain Aster oe Beene and T. M. Barkley (certain Senecio).

EFERENCES
ANDERSON, i ins 1959. Flora a Alaska and adjacent parts of Canada. Iowa State
Tae see
CODY i Aer plant records for northern Alberta and southern Mackenzie
District. Hee ae Nat. 70:
60. Plants Be the ae! Ase Norman Wells, Mackenzie District,
Northwest Temes Canad. Field-N 0.
New eae ee ee the upper eas River valley,
Mackenzie District, Nonthere: poe Canad. Field-Nat. 75:
——————_. 1963. A contribution to the knowledge of ae ree of southwestern
Mackenzie District N. W. T. Canad. Field- Nat. 77: 108-123.

170

GLEASON, H. A. The New eee and Brown Illustrated Flora of vie North-
a Sie States ana a cent Canada. New York Botanical Garden, New
ULTEN, E. 1941-1950. Flora of Alaska and Yukon. Lunds Universitets Peres N.F.
Pei es ue 37- a
JEFFR Ww.
Nat. ane nee al ae
MOSS, E. H. 1953. Forest pee in northwestern Alberta. Canad. Jour. Bot. 32:

212-252.
Canad.

Se on plant occurrence along lower Liard River, N.W.T.

. 1953a. Marsh and bog vegetation in northwestern Alberta.
Jour. Bot. 31: 448-470.
1955. The eee ee Alberta. Bot. ae 21: 493-567
1959. Flora of Alber University of T o Press, Tor

PORSILD, A. E. er Materials for a oe ra of the Set rE | eee Teer macs of

Canada. Sargentia 4: a
ss “The ee mie _ the east slope of Mackenzie Mountains, North-
da

west Territories. Nee Mus. Cana
1951. Botany a. ees Yukon adjacent to the Canol Road.

Nat. Mus. Canada ee 121.
e a vascular plants of the western Canadian Arctic Archipelago.

ae Mus. Canada bul
UP, H. M. 1935. a eal investigations in Wood Buffalo Park. Nat. Mus. Canada

Bull 74,
ae SN se studies in the Athabaska-Great Slave Lake region.

I. Catalogue of the ie plants. Jour. Arn. Arb. 17: 180-315.
eh peace studies in he: Athabaska-Great Slave Lake region, II.

Jour. Arn. Arb. 27: 1-

es Botany of southwestern Mackenzie. Sargentia #
neepessiceen H. i 1957. Flora of Manitoba. Nat. Mus. Canada Bull.
THIERET, J. W. 1959. ae ei vegetation near Fort ae Sees Terri-
tories. a Field- vee 73: 161
961. ew aan records for southwestern District of Mackenzie.

Canad. Field-Nat. i: Ree 12
ew elas records from District of Mackenzie, Northwest Terri-

.N
tories. Canad. Field- ae 76: 2
a. Life- penne in the plains flora of southern Mackenzie, North-

3
west Territories. Rhodora 65: 149-
963b. adieone. to the flora of the Northwest Territories. Canad.

Field-Nat. 77: 126.

GILIA AND IPOMOPSIS (POLEMONIACEAE)
IN TEXAS
LLOYD H. SHINNERS
Southern Methodist University, Dallas 22, Texas

two new nomenclatural combinations under Ipomopsis. It is primarily to
publish these and make them available for Dr. Wherry’s use (or rejection,
as the case may be) that this brief paper has been prepared. Since no
keys to the Texas representatives as currently understood are available,
and since the listings in Gould’s recent Texas Plants leave much to be
desired, it seems worth while to include my key to the genera and notes
on the species (other than Phlox). It has been my usual procedure to
work up various groups on the basis of collections at SMU to the extent
of revising the nomenclature, checking relevant publications, and prepar-
ing utility keys for identification of flowering material; then put the
manuscript notes aside until additional material has accumulated, or

TO TEXAS GENERA OF POLEMONIACEAE
la. Leaves pinnately compound, with distinct, broad leaflets
2. Polemonium
lb. Leaves simple, but in some species cut into narrowly lanceolate to
thread-like segments
2a. Lower or all leaves opposite
3a. Calyx enclosed by large, spiny-toothed, net-veined bracts
1. Loeselia
3b. Calyx with entire, 1-ribbed bracts or none (subtended by upper
leaves in some species)
4a. Leaf blades linear to oblong-ovate or obovate, entire, more than
1.3 mm. wide; annuals or perennials, general distribution .
3. Phlox

SIDA I (3): 171—179. 1963,

172

4b. Leaf blades linear to thread-like or cut into thread-like segments
less than 1.3 mm. wide; small annuals of desert areas, Trans-
REGS cs pig bce PERE EAL DSR RSS eee 7, Linanthus
2b. Leaves alternate or basal

5a. Corolla appearing rotate (tube very short), or narrowly funnel-

form with tube flaring upward...........-.--5--++++55> 4, Gilia

5b. Corolla more or less salverform, with nearly cylindrical tube
longer than the lobes

Ga. Calyx and upper leaves sparsely woolly, or without woolly hairs

5. Ipomopsis

6b. Calyx and base of upper leaves largely hidden by matted, wooly

WAS oo ca ees Oaa a eee RE EOLA EELS POS Ss . Eriastrum

1. LOESELIA. The Mexican L. SCARIOSA (Martens & Galeotti)
Walpers has been reported by Standley (1937) from the Chisos Moun-
tains, southern Brewster County. I have not seen specimens. That cited
by Standley was collected in August.

2. POLEMONIUM. P. PAUCIFLORUM S. Watson (including P. Hinck-
leyi Standley, Amer. Midl. Nat. 18: 684, 1937; according to Verne Grant,
1959) occurs in the Davis Mountains, Jeff Davis County. Flowering in
August.

3. PHLOX. P. bifida var. induta Shinners, S.W.Nat. 6: 50—51, 1961, is
referred by Dr. Wherry to his P. oklahomensis. This latter is regarded by
Marsh (1960) as specifically distinct from P. bifida. Marsh’s paper ap-
peared while my note was in press. By a similar curious coincidence,
Phlox Johnstonii Wherry (1961) was published after Erbe and Turner’s

the study of Erbe and Turner is more superficial than that of Dr. Eula
Whitehouse (1945), which remains the classical work on the annual
species. The genus is both large and very difficult in Texas, and I have
not yet completed even a preliminary account.

4, GILIA and 5. IPOMOPSIS. See below.

6. ERIASTRUM. E. DIFFUSUM (Gray) H. L. Mason, Madrono 8: 76.
1945. Gilia filifolia var. diffusa Gray, Proc. Amer. Acad. 8: 272. 1870.
“Fort Mohave and Nevada to New Mexico and the borders of Texas.”
Cited by Craig (1934) from El] Paso and Fort Bliss, both in El Paso
County. The following collections are at SMU. EL PASO CoO.: Frontera
(El Paso), hills and rocky places, Charles Wright, 30 March 1852. Lower
slopes of Mt. Franklin, El Paso, sandy soil, Barton H. Warnock 10349, 19
April 1952. HUDSPETH CO.: steep rocky (igneous) slopes, north end of
Quitman Mts., 8 miles W. of Sierra Blanca, Rogers McVaugh 8026, 21
April 1947.

7. LINANTHUS, the white-flowered L. BIGELOVII (Gray) Greene is

173

known from EL PASO Co.: Frontera (El Paso), Charles Wright, April
1852 (SMU). The yellow-flowered L. AUREUS (Nuttall) Green is said
to extend east to Texas by Kearney & Peebles (1960); I have not seen
specimens from the state.

KEY TO TEXAS GILIA

la. Corolla funnelform, with well-developed, gradually flaring tube
pA CCCCin Ete 1ODeSiampe twig oe ee . G. mexicana
1b. Corolla appearing rotate, the inconspicuous tube much shorter than

2a. Lower stem leaves with long, naked petioles and wide, flat blades
3a. Blades of lower stem leaves slightly to much longer than broad,

taproot); Trans-Pecos mountains............____ 3. G. perennans
2b. Lower stem leaves (not basal ones) sessile or subsessile or with
tappered, winged-petiolar basal portion, commonly with mall,

4a. Calyx in flower divided about half way
da. Basal leaves numerous, persistent; lower stem leaves deeply
pinnatifid, the 5—7 pairs of segments mostly coarsely toothed
or deeply lobed; Rio Grande Plain 4. G. ludens
5b. Basal leaves mostly absent at flowering time; lower stem

da. G. rigidula var. rigidula.
6b. Lower leaves with stiff, almost needle-like segments less than

1 mm. wide; upper leaves similar to lower
db. G. rigidula var. acerosa

4b. Calyx in flower divided about 3/4 or more
7a. Calyx 10—12 mm. long, chiefly of white, scarious tissue with
narrow green bands extending from base up through center of

ENA MO) 1S. alo 3 oar re ec 6. G. insignis
7b. Calyx 4.0—5.5 mm. long, chiefly of green tissue, the lobes with
narrow, white, scarious margins............___. 7. G. Stewartii

1. G. MEXICANA A. & V. Grant, Aliso 3: 255—9257. 1956. Type from.
Arizona; cited also from New Mexico (Grant and Dona Ana counties):

174

and Chihuahua. There are three specimens at SMU from westernmost
Texas EL PASO CO.: Frontera (El Paso), Charles Wright, April, 1852.
Infrequent in limestone soil, McKelligon Canyon, Franklin Mts., El Paso,
Warnock 7664, 26 March 1948. On west, lower, limestone slopes of Frank-
lin Mts. about 2 miles west of El Paso, Warnock 10308, 10 April 1952.

2. G. INCISA Bentham in DC., Prodr. 9: 312. 1845. “In Texas (Drum-
mond coll. 3a n. 463!).” Type material not seen. Drummond collected in
the area from Galveston to Victoria and Gonzales counties, and inland
as far as Milam County. This area touches the southeastern limits of the
range of this species as shown by collections at SMU (county names in
parentheses): Eastern Rio Grande Plain (Cameron, Kleberg, Live Oak),
up the coast to Aransas County, and on the Edwards Plateau (north to
Travis and San Saba, west to Val Verde), at elevations between sea level
and 2,000 feet. Flowering late March—early June,

3. G. perennans Shinners, sp. nov.. G. incisae peraffinis, sed perennans
demum cum radice crasso lignoso, foliorum inferiorum caulinorum la-
minis brevioribus minus divisis; species monticola. HOLOTYPE: rock
crevice, north fork, Guadalupe Mountains, north McKittrick Canyon,
Culberson Co., Texas, D. S. Correll 13958, 18 August 1946 (SMU). “Flow-
ers blue.” The following additional specimens have been seen, all from
Trans-Pecos Texas at elevations of 5,000 feet or above, all deposited at
SMU. CULBERSON CO.: numerous, in a spot growing in the almost solid
rock of the creek bed in north McKittrick Canyon, Guadalupe Mountains,
L. C. Hinckley & Leon Hinckley 29, 3 June 1949. In limestone soil above
Hunter Lodge, in south McKittrick Canyon of Guadalupe Mts., Warnock
9536, 31 August 1950. Same locality, Warnock 10950, 3 August 1952. JEFF
DAVIS CO.: infrequent in Limpia Canyon at mouth of Wild Rose Pass,
Kokernot Ranch, Davis Mts., Warnock & F. M. Churchill 7742, 6 April
1948.

This differs from G. incisa more in its biology than in its morphology,
and it is only after considerable hesitation that it is named as a distinct
species. In addition to the points noted in the key, the calyx averages
slightly larger (4.0—4.7 mm. long vs. 3.5—4.5 mm. in G. incisa). The fact
that all but one of the collections were made in summer, long after the
normal flowering time for G. incisa, is partly due to accidents of col-
lecting, since most botanists have visited during the summer, but in part
it doubtless reflects the higher altitude.

4. G. ludens Shinners, sp. nov. Perennis? humilis (ca. 12-21 em.) pler-

silt loam, railroad right-of-way, 4 miles west of Alice, Jim Wells Co.,
Texas, Shinners 19581, 10 April 1955. (SMU). “Corolla blue (flowers

175

mostly past).” PARATYPES (all SMU). DUVAL CO.:: in Pleistocene

March 1963. WEBB CoO.: in light red sand, State Highway 359, 2 miles
west of Bruni, Elvira G. Garcia 137, 16 March 1963.

Because of the prominent, flat-bladed basal leaves, I at first mistook
this for G. incisa, despite the relatively short-pedicelled flowers. On later
comparison in the herbarium, it seemed closer to G. rigidula, but that
species lacks the prominent basal leaves and is confined to areas from
the Edwards Plateau north and west. Bentham’s description of the leaves
of G. incisa (he apparently had only upper ones) as “cuneatis lanceolatis
linearibus subintegris v. saepius acutissime incisis” makes it certain that
he did not have the species here described.

9. G. RIGIDULA Bentham in DC., Prodr. 9: 312. 1845. “In Texas pr.
Bejar (Berlandier).” This often-mentioned locality is the present San
Antonio, Bexar County. The description and locality are quite sufficient
to establish the identity of Bentham’s plant. I follow Asa Gray in recog-
nizing two varieties,

da. G. RIGIDULA var. RIGIDULA. Edwards Plateau, north to Travis,
Burnet, Jones, Taylor and Mitchell counties, west into eastern Trans-
Pecos (Pecos, Reeves, Val Verde). Flowering late March—May, sporadi-
cally to October.

db. G. RIGIDULA var. ACEROSA Gray, Proc. Amer. Acad. 8: 280.
1870. “North New Mexico to Arizona, Fendler, Gordon, Wright, &c.”
(Specimens not seen.) In the Synoptical Flora the range is extended to
“borders of Texas” (p. 149). Northern Trans-Pecos (Culberson, Huds-
peth, Jeff Davis) and Panhandle, southeast to Taylor County. Flowering
late March (rarely) or April to September,

6 G. INSIGNIS (Brand) Cory & Parks, Cat. Fl. Texas (Texas Agr.
Exp. Sta. Bull. 550) p. 85. 1938 (“1937”). G. rigidula ssp. insignis Brand,
Pflanzenreich IV. 250. p. 149. 1907. “So nur in Nord-Mexiko: Coahuila

abundant especially along road; stony flats, creosote shrub association,
about 3 miles S. of Persimmon Gap, McVaugh 7831, 5 April 1947 (SMU).
Frequent along roadside near Dog Flat, Warnock 21485, 2 April 1938.

7. G. STEWARTII I. M. Johnston, Journ. Arnold Arb. 24: 94, 1943.
Cited by Johnston from Brewster and Hudspeth counties, and from
numerous localities in Mexico. I have seen the following Texas collec-

176

tions. BREWSTER CO.: frequent on limestone hills between Lone Moun-
tain and Nugent Mountain, Big Bend National Park, Warnock 16h, 3
Sept. 1947. On alluvial fan, 6 miles north of Hot Springs, Big Bend Park,
Rose-Innes & Warnock 546, 20 March 1941 (cited by Johnston). PRE-
O CO.: caleareous gravel hillsides, south end of Van Horn Mts.
near Porvenir, U. T. Waterfall 4754, 26 June 1943 (det. I. M. Johnston).
Along rocky ledge e. Van Horn Creek at north end crossing, Porvenir-
Chispa road in western end of county, L. C. Hinckley 2230, 25 Oct. 1941.
Excluded Species
G. ophthalmoides ssp. australis A. & V. Grant, Aliso 3: 263. 1956. De-
scribed from southern Arizona and southwestern New Mexico; er
by Gould from Trans-Pecos Texas, but I have seen no specim
G. ophthalmoides ssp. flavocincta (A. Nelson) A. & V. ve i c. 262.
Cited by the Grants only from Arizona; reported by Gould for the entire
western half of Texas, but I have seen no specimens.
KEY TO TEXAS IPOMOPSIS
la. Corolla tube 0.3—1.0 cm. long
Da. Tea veetal ENE Rs 64 peu nade ee oe ee ESS 8. I. Wrightia
2b. Leaves (except uppermost) toothed or divided into linear or
thread-like segments
3a. Corolla lobes 5—7 mm. long, more than half as long as the tube
1.J. Havardu
3b. Corolla lobes 1.0—4.5 mm. long, less than half as long as the tube
4a. Annual with slender taproot, flowering April—early July;
desert flats
5a. Corolla tube 3—5 mm. long; upper leaves deeply toothed or
pinnatifid, the segments short and wide... .6. I. polycladon
5b. Corolla tube 6—8 mm. heey upper leaves entire or cut into
8-5 thread-like segments...........--....---. 7.
4b. Perennial with rather sae pee flowering J ig-Oeroben:
mountains
6a. Primary axis of inflorescence with about 1—10 nodes, the
lower flowering branchlets mostly subtended by reduced un-
divided leaves (see excluded species at end).. I. multiflora
6b. Primary axis of inflorescence with about 10—35 nodes, the
lower flowering branchlets mostly subtended by pinnatisect
leaves (see excluded species at end)............ I. pinnata
1b. Corolla tube 1.2—4.5 cm. long
Ta. Inflorescence loose, open, broad; stem freely and widely branch-
ing; corolla lavender-blue to white
8a. Corolla tube 1.2—2.0 cm. long; corolla lobes 3—6 mm. long, 1.0—
Si AOE, «4 4a Kee REED eee PAY On 2. I. laxiflora
8b. Corolla tube 2.2—4.5 cm. long; corolla lobes 6.5—12.0 mm. long,
(Hil. WIGS: i.e ok esos eee ee ede neo Ree 3. I. longiflora

177

7b. Inflorescence narrow, dense, elongate; stem normally simple or
with few, elongate branches; corolla red or rarely yellow
Ya. Calyx lobes about as long as the tube; Trans-Pecos
4. I. aggregata
9b. Calyx lobes nearly twice as long as the tube: central and eastern
he ig. apr cecnd io te ARCnet RA LEO Leave» do. I. rubra

1. I, HAVARDII (Gray) V. Grant, Aliso 3: 357. 1956. Loeselia Havardi
Gray, Proc. Amer. Acad. 19: 87, 1883. “W. Texas, on the Rio Grande near
Presidio del Norte, Dr. N. (sic) Havard, 1881.” Gilia Havardi Gray, Syn.
F]. 2 pt. 1 (suppl.) p. 411. 1886. I have seen only the following specimens.
BREWSTER CoO.: limestone soil in Avery Canyon, Big Bend National
Park, Warnock 9141, 23 July 1950. PRESIDIO CO.: about 1 mile south-
east of Greenwood ranchhouse some 50 miles south of Marfa, L. C.
Hinckley 3578, 4 April 1946. Common along washes, rolling Rio Grande
Plain, 4 miles SE of Ruidosa, C. H. Muller 8440, 28 July 1945.

2. J. LAXIFLORA (Coulter) V. Grant, Aliso 3: 361. 1956. Gilia Ma-

combit var. laxiflora Coulter, Contrib. U.S. Nat. Herb. 1: 44, 1890. “Camp
eee (Ixion county),” (i. e. Irion County), G. C. Nealley 311 (US;
not seen). Gilia laxiflora (Coulter) Osterhout, Bull. Torr. Bot. Club 24:
51. 1897. Trans-Pecos (Jeff Davis, Presidio, Reeves), east in South Plains
to Lubbock and Mitchell counties. Flowering late April—July.

3. I. LONGIFLORA (Torrey)V. Grant, Aliso 3: 361. 1956. Cantua longi-
flora Torrey, Ann. Lyc. N.-Y. 2: 221. 1826. “On the Canadian” (probably
in the Texas Panhandle). Gilia Tenaiflera (Torrey) G. Don, Gen. Hist.
Dichlam, Pl. 4: 45. 1838. Widespread in Trans-Pecos, Panhandle, South
Plains and Red Plains, east to Wilbarger and Stonewall counties. Flow-
ering late March (Big Bend area) or April to October.

4. I. AGGREGATA (Pursh) V. Grant var. texana (Greene) Shinners,
comb. nov. Callisteris texana Greene, Leafl. Bot. Obs. & Crit. 1: 160.
1905. “Guadalupe Mountains, western Texas, V. Havard, Sept., 1881. Type
in U. S. Nat. Herb.” (not seen). Gilia aggregata var. texana (Greene) I.

Gould, as Gilia texana and as Ipomopsis aggregata. Common in the
Trans-Pecos mountains in Brewster, Culberson, Jeff Davis and Presidio
counties. Flowering July—September.

5. I. RUBRA (L.) Wherry, Bartonia 18: 56. 1936. Polemonium rubrum
L., Sp. Pl. 1: 163. 1753. “Habitat in Carolinae citerioris arenosis. B.

eastern counties in the Pine Belt; frequent and locally abundant on var-
ious substrates, often on dry, rocky, limestone slopes, in central Texas
west to Taylor and Scurry counties and southwest to Caldwell County.
Also frequent in cultivation. Flowering late May—early July, and spora-

178

dically to September. A semi-albino with yellow flowers has been ob-
served in Rockwall County.

This is an example of a species first described from the Southeastern
United States but relatively uncommon there, having its center of abun-
dance west of the Mississippi River. Mirabilis albida and Hedyotis nigri-
cans are other examples of such distribution. These are all of South-
western and Mexican relationship; I would not agree with Verne Grant
in decribing Ipomopsis rubra as characteristic of the Southeast.

6. I. POLYCLADON (Torrey) V. Grant, Aliso 3: 361. 1956. Gilia poly-
cladon Torrey in Emory, Rept. U.S. & Mex. Boundary Survey 2: 146.
1859. “Stony hills near El Paso, March.” No collector is named; the fol-
lowing is the only specimen seen. EL PASO CO.: Frontera (Hl Paso),
Gravelly places, Charles Wright, April 1852.

7. I. PUMILA (Nuttall) V. Grant, Aliso 3: 361. 1956. Gilia pumila
Nuttall, Journ. Acad. Nat. Sci. Phila. (ser. 2) 1: 156, 1848. “Near the first
range of the Rocky Mountains of the Platte. Flowering in May. (Nut-
tall.)” In Texas confined to the Trans-Pecos. CULBERSON CO.: near
Salt Lake on Highway 54, about 30 miles north of Van Horn, Eula White-
house 11455, 5 July 1931. HUDSPETH CO.: only four plants found in
barpit at roadside about 5 miles S. Sierra Blanca, Hinckley & Hinckley
68, 12 June 1949. PRESIDIO CO.: sandy desert flats, 2 miles north of
Porvenir; abundant, McVaugh 7995, 17 April 1947.

8. I. Wrightii (Gray) Shinners, comb. nov. (This appears as a nomen
nudum without indication of combining author in Gould’s Texas Plants,
1963.) Gilia Wrightii Gray, Proc. Amer. Acad. 8: 273. 1870. “Western
frontiers of Texas, on the Rio Grande forty miles below El Paso, C.
Wright, n. 496.” Two collections seen, the first from the general area of
the type locality. EL PASO CO.: in sandy soil along road about 15 miles
north of Ysleta, W. J. Tebeaux, 13 Aug. 1951. Rather infrequent in deep
sand along Carlsbad highway about 16 miles east of El Paso, Warnock
10902, 28 July 1952. Annual developing a stout taproot; Gray described
the species as doubtfully perennial.

Excluded species
I. multiflora (Nuttall) V. Grant, Aliso 3: 357. 1956. Gilia multiflora
tall. Gray in 1870 (Proc. Amer. Acad. 8: 260) stated that Collomia
Cavanillesiana occurred on ‘borders of W. Texas, New Mexico, Arizona
and adjacent Mexico.” In the first edition of the Synoptical Flora ee
2 pt. 1: 136, 1878), he reports it from “New Mexico and W. Texas
Arizona.” In the Supplement added to the 2nd edition (p. 411, 1886), .
name is given as a synonym “in part” of Gilia multiflora, which is stated
to be “common in New Mexico and Arizona.” Presumably the earlier re-
port of Texas was found to be in error. I have seen specimens from
New Mexico and Arizona, but not from Texas.

179

I. pinnata (Cavanilles) V, Grant, Aliso 3: 357. 1956. Phlox pinnata
Cavanilles, Icones 6: 17, t. 528 fig. 1. 1801. “Habitat in Montevideo vici-
niis, . . . Phlox haec pinnata crescit etiam passim in Nova Hispania
prope Real del Monte.” Bentham and Gray were certainly correct in be-
lieving that the locality Montevideo was an error (cf. Gray, 1870, p. 260,
where it is erroneously given as “Buenos Ayres”). Collomia Cavanillesi-

reich IV. 250. p. 112. 1907. Specimens seen from Chihuahua, Durango and
Zacatecas. It is not unlikely that this species may be found in the Big
Bend area of Texas

DV IN C Bas

AIG, THOMAS. 1934. A revision of the See Pee of the genus Gilia (Pole-
ne Bull. Torr. Bot. Club 61: 385—396, 411—
ERBE, LAWRENCE, AND B. L. TUR ean 1962. i Sbosystemati study of the Phlox
cus pidata— Phlox ieee complex. Amer. Midl. Nat.
GOULD, F. W. (January; title page aie a Tecan Blancs (Polemoniaceae,
Dez
ALVA, AND VERNE eee ee: Genetic and taxonomic studies in Gilia.
on The Cobwebby G ae Aliso 3: 203—2
RANT, VERNE 6. A synopsis mee oe ue 3: 351—362.
oie tural eee of the Phlox Fami ie Volum
GR ane oe Tons eee of the North rea peas ae Amer. Acad.
8: 247—
- 1886. Polemoniaceae. Syn, Fl. N.A. (2nd ed.) 2 pt. 1: 128—151, suppl.

Hye
KEA ARTES THOMAS H., AND ROBERT H. PEEBLES. 1960. Arizona Flora. (Pole-
eee Ppp. 678—696.
ARSH, DANIEL L. 1960. ao en bee ox oklahomensis to Fags Nae complex:
d,

eee a new subspecies of Phlox bifida. Trans. Kansas Aca
MASON, HERBERT L. 1945. The genus Frist as the ee of Benham and
Gray upon the problem of generic confusion in Polem 8:

aceae. Madrono
SHINNERS, LLOYD H. 1961. Phlox te a var. ae (ieee he a new erdcinic
in north central Texas. $.W. Nat. 6: 50—5
STANDLEY, PAUL C. 1937. Three new ae from aoe and Mexico and some recent
additions to the Texas flora. pet Midl. Nat. 18: 683—
WHERRY, re a 195 eee Genus Phlox. Morris ee Mono. III.
LA annual Phlox. Wrigheia 2: 198—199,
WHITEHOUSE, ou 1945, Annual Phlox species. Amer. Midl. Nat. 34: 388—401.

LEPTOGLOSSIS AND NIEREMBERGIA
(SOLANACEAE) IN TEXAS

LLOYD H. SHINNERS
Southern Methodist University, Dallas 22, Texas

Two genera of Texas Solanaceae with salverform corollas are so
strikingly similar to Polemoniaceae that at first glance they are apt to
be mistaken for members of that family. Technically of course they dif-
fer in having simple style and stigma. They also differ in the conforma-
tion of the calyx, which is narrowly funnelform, so that there is con-
siderable open space between it and the narrowly cylindrical corolla
tube. In the Polemoniaceae which they resemble the calyx is appressed
to the corolla tube or nearly so. It is an interesting example of recom-
binations of characters that Bouchetia erecta DC., another representative
of Texas Solanaceae, has a flaring calyx closely resembling that found
in the two genera here discussed, but its corolla tube also narrowly fun-
nelform, so that the calyx fits closely against it. Leptoglossis and Nierem-
bergia are represented in Texas by a single species each, the former
native but currently passing under an incorrect name, the latter intro-
duced and not previously reported as a wild plant for the state. They
may be distinguished as follows.

Corolla tube abruptly swollen toward one side at summit; anther-bearing
stamens 4, in two very unequal pairs, barely exserted; native (en-
demic in Val Verde County)............0.0 00002 e sees eptoglossis

Corolla tube narrowly cylindrical to summit; anther-bearing stamens 5,

equal or subequal, prominently exserted; cultivated and escaped
Nierembergia

LEPTOGLOSSIS TEXANA (Torrey) Gray, Proc. Amer. Acad. 12: 164.
1877. Browallia texana Torrey in Emory, Rept. U.S. & Mexican Boundary
Survey 2: 156. 1859. “Near the mouth of the Pecos, September-October;
Bigelow, Pope. (No. 535, Wright.)” (These sient not seen, but sev-
eral approximate topotypes examined: SM U.) Nierembergia viscosa

preceding.) Leptoglossis viscosa (Torrey) Millan, Darwiniana 5:489.
1941. Probably it was the influence of the American Code, which ac-
cepted page priority as binding, that led to the revival of the name
Nierembergia viscosa, by which this species has recently been called.
Gray, in uniting the two Torreyan species, deliberately chose the epithet
texana in preference to vicosa (L¢., p. 165), and under current rules, his
choice must be followed. Millan evidently had not seen Gray’s com-

SIDA 1 (3): 180—181. 1963.

181

ments when he excluded the species from the genus Nierembergia.

Leptoglossis texana is a perennial which, like so many in Texas, may
flower the first year from seed. It suggests a Phlox with light purple or
purple-pink corolla, blooming late March—early May and rarely in
October. All collections examined were from Val Verde County, on lime-
stone substrates.

NIEREMBERGIA HIPPOMANICA Miers var. CAERULEA (Miers)
Millan, Darwiniana 5: 521. 1941. N. caerulea Miers. (For full synonymy

REFERENCES
GRAY, ASA. 1877. Characters of some new or little known genera of plants. Leptoglossis
subg. Brachyglossis. Proc. Amer. Acad. 12: 164—165.
MILLAN, ROBERTO. 1941. Revision de las especies del genero Nierembergia. Darwiniana
3: 487—547.
ORTON, C. V. 1944. Taxonomic studies of tropical American plants. Notes on
Bouchetia. Contrib. U.S. Nat. Herb. 29: 72—73. (Considers B. erecta of Texas distinct
from B. anomala of Argentina and Uruaguay.)

NOTES

MONANTHOCHLOE LITTORALIS (GRAMINEAE) IN LOUISIANA.
— The following collection is apparently the first record of the grass
Monanthochloe littoralis Engelm. from Louisiana: '% mile east of mouth
of Mermentau River at Hackberry Beach, Cameron Parish, Wayne G.
Harris 55, July 17, 1962. The species was found to be common locally,
with Spartina patens and Distichlis spicata, on the slope behind the shell
beach, about 100 yards north of the gulf shore. Voucher specimens have
been deposited in the herbarium of the University of Southwestern
Louisiana, in the United States National Herbarium, and in the herbar-
ium of Southern Methodist University—A. G. Owens, Jr. and Sam
Riche, Louisiana Mosquito Control Association, Lafayette, Louisiana.

CENCHRUS LONGISETUS M. C. JOHNSTON, NOM. NOV. (GRA-
MINEAE).— Based on Pennisetum villosum R. Brown ex Fresenius,
Mus. Senckenb. Abh. 2: 134, 1837 (fide Hitchcock, Man. Grasses ed. 2 p.
934). Not Cenchrus villosus (Sprengel) Sprengel, Syst, 1: 300, 1829.-—
Marshall C. Johnston, University of Texas, Austin 12:

NTROSEMA FLORIDANUM (BRITTON) LAKELA, COMB. NOV.
ere. — Based on Bradburya floridana N. L. Britton, Tor-
reya 4: 142. 1904. Since Centrosema has been made nomen conser-
vandum under the International Code of Botanical les this
transfer is required—Olga Lakela. (Contribution No. 4, Botanical
Laboratories, University of South Florida, Tampa.)

THE VARIETIES OF TEUCRIUM CANADENSE (LABIATAE). —
Elizabeth McClintock and Carl Epling, in “A revision of Teucrium in
the New World, with observations on its variation, geographical dis-
tribution and _ history” (Brittonia 5: 491—510, 1946), recognize three

as follows.
la. Calyx without glandular hairs
2a. Leaf blades medium to moderately dark green above, grayish green
to gray beneath, midrib on lower surface with hairs pointing for-
ward, widely spreading, or very loosely retrorse; plants of wide
distribution, outside the area of the next.......... var. canadense
2b. Leaf blades very dark green or blackish green above (medium
green in a few Mississippi coast specimens), silvery beneath,

SIDA 1 (3): 182—183. 1963.

183

midrib on lower surface with closely appressed retrore hairs in
basal portion (rarely loose or spreading ni north Florida speci-

TOTS) ee a ey ih selon oto, cde Wh ne a eee var. Nashii

lb. T. CANADENSE var. Nashii (Kearney) Shinners, comb. nov. T.
Nashii Kearney, Bull. Torr. Bot. Club 21: 484, 1894, “Collected in middle
Florida in 1836, by Dr. Chapman; in Duval county, Florida, by Mr.
A. H. Curtiss (No. 1975) and near Eustis, Florida, in 1894, by Mr. George
V. Nash (Numbers 1496, 1505 and 1516).” I have not seen the syntype

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closely appressed hairs, those on the basal portion of the midrib pointing
toward the petiole. Rarely the hairs on the midrib are loose (one speci-

medium green upper surface on the leaves, approaching var. canadense.
In all cases the leaf blades are relatively narrow, but I do not consider
this justification for putting the Florida populations with the Arizona
ones, as was done by McClintock and Epling. The pubescence of the
latter is loose and gray, not silvery, and the blades are not strongly
discolorous.

lc. T. CANADENSE var. boreale (Bicknell) Shinners, comb. nov. Pe
boreale Bicknell, Bull. Torr. Bot. Club 28: 171. 1901. “New Hampshire
to northern New York.” Type from Vermont, in Herb. N. Y. Bot.
Garden.” (Not examined.) T. occidentale var. boreale (Bicknell) Fernald,
Rhodora 10: 85. 1908. T. occidentale Gray, Syn. Fl. N.A. 2 pt. 1: 349. 1878.
T. canadense var. occidentale (Gray) McClintock & Epling, Brittonia 5:

184

DICERANDRA IMMACULATA LAKELA, SP. NOV. (LABIATAE).—
D. frutescenti peraffinis differt floribus paulum brevioribus, corollis
rubris vel purpurascentibus immaculatis tuba minus exserta labio super-
iori cristato lato claviformi, antherarum appendiculis puberulis. HOLO-

PE: numerous plants in flower; on disturbed sandscrub, west of U.S.
1, near south boundary of Indian River County, Florida, Lakela 25440,
30 September 1962 (USF). The species occurs also in adjacent northern
St. Lucie County. The site of the type specimen is a transition yellow
and white sandscrub, with remnants of Pinus clausa, Zanthoxylum Clava-
Herculis, Carya floridana and scrub species of Quercus. Post-mature
fruiting cymes (Lakela 25613, 19 January 1963) contained only a few
nutlets; mildew growing on the nectary gland destroys the fruits. Plants
in preanthesis (Lakela 25221, 29 July 1962) were collected in white
sandscrub with overhead of tall Pinus clausa, in St. Lucie County. Fruit-
ing cymes from this colony (Lakela 25614, 19 January 1963) were also
infected with mildew.

Dicerandra immaculata differs from D. frutescens Shinners in slightly
shorter flowers, color of the spotless corolla, less exserted tube, broadly
clavate upper lip as outlined by dorsal crests, and puberulent anther

“Synopsis of Dicerandra (Labiatae)” (SIDA 1: 89—91, 1962) may be
emended to accommodate the new species by deleting the reference to
flower color in his couplet 2a and inserting the following couplet after
it; his text may be supplemented by adding the full description of the
new species given below.
Corolla white or yellowish white with purple dots; interior peninsular
D

Florida (Highlands and Sumter COUNTIES) «heck: . frutescens
Corolla peach-red or purplish or white, without dots; eastern peninsular
Florida (Indian River and St. Luie counties)....... D. immaculata

DICERANDRA IMMACULATA Lakela. Aromatic, frutescent peren-
nial 4—5 dm. tall from woody, wide-spreading roots. Stems several,
branching from the base, glabrous or minutely puberulent, especially at
nodes; internodes alternatingly sulcate between decurrent lines, green
and sparingly punctate above, woody below with brownish exfoliating
periderm with latent buds and persistent petiole bases. Leaves 2—3 cm.
long, short-petioled; blades oblanceolate, essentially glabrous or min-
utely puberulent at base, copiously punctate, uninerved, passing into
floral bracts; fascicular leaves similar but smaller. Floral axes puberu-
lent; cymes axillary, usually 3-flowered; peduncles 3—5 mm. long, hori-
zontal, about equalling the ascending or erect pedicels. Cayx in full
anthesis 7—8 mm. long, punctate, green or often purplish, becoming
whitish at the ciliate, bilabiate apex, glabrous or puberulent without,
with a circle of cilia just below sinus level within the throat; upper lip

SIDA 1 (3): 184—185. 1963.

185

serted, anterior pair 10—12 mm. long, posterior 7—8 mm. long; anthers
purple, horizontally oblique with subulate, puberulent horns. Style to
19 mm. long, pilose above, with equal stigmas. Fruiting calyx (dry)
8.2 mm. long; nutlets ovoid, 1.0—1.2 mm. g£

26,573, 11 October 1963 (USF).—Olga Lakela. (Contribution No. 5, Botani-
cal Laboratories, University of South Florida, Tampa.)

WAHLENBERGIA LINARIOIDES (CAMPANULACEAE) IN FLOR-
IDA: A SECOND ADVENTIVE SPECIES FOR THE UNITED STATES,

from central Louisiana (Shinners, S.W. Nat. 2: 44, 1957). There are also
unreported collections at SMU from southern Mississippi made in 1960
and 1962 (Shinners, in letter).

A second species, W. linarioides (Lam.) A. DC., has appeared in the
weed flora of FLORIDA, Escambia Co.: sandy vacant lot, West Pensa-

bergia, I sent material of this one to Dr. Carroll E. Wood, Jr., Harvard

University, who generously made the determination and stated that there

are specimens in the Gray Herbarium from Argentina, Chile, Bolivia and

Brazil. The principal differences between it and W. marginata may be

summarized as follows:

Hypanthium in flower 3.5—4.5 mm. long, narrowly obconical to sub-
cylindrical, in fruit up to 12 mm. long, subcylindrical; seeds broadly

SIDA 1 (3): 185—186. 1963.

186

AOC civ soo Sied ewan o wea ae eee ee W. linarioides
Hypanthium in flower 1.53.0 mm. long, ellipsoid or ovoid, in fruit up
to 7.5 mm. long, obconic-obovoid; seeds oblong........ W. marginata

Wahlenbergia linarioides is a perennial herb, usually with several
erect, glabrous stems up to about 5 dm. tall, simple to the inflorescence.
Leaves glabrous, sessile, the lower linear-oblanceolate, 2—3 cm. long,

glands and/or the barest suggestion of teeth. Inflorescence open-cymose,
about 2 dm. long and 1.5 dm. broad; branches alternate, elongate, each
terminated by a cymule of about 3—10 flowers. Hypanthium in flower
narrowly obconical, glabrous, in fruit obconical to subcylindrical, rib-

linear-oblong anthers. Style slender-elongate at base, dilated abruptly
above to an obconical, truncate stigma. Ovary 2-locular, producing
numerous, broadly elliptic, lenticular, lustrous, amber seeds about 0.3
mm. long.

Expenses incurred in this study were met in part by research grant
GM-06305 from the Division of General Medical Sciences, Public Health
Service—R. K. Godfrey, Department of Biological Sciences, Florida
State University, Tallahassee.

CONTRIBUTIONS
TO BOTANY

VOLUME 1 NUMBER 4 JUNE 1964

CONTENTS

Botanical Survey along the Yellowknife Highway, Northwest
Territories, Canada. II. Vegetation.

* John W. Thieret. 187

Systematic status of Ammopursus Ohlingerae (Compositae).
Olga Lakela.

NOTES. Fatoua villosa (Moraceae) in Louisiana: new to North America. 248.—Amaranthus
microphyllus Shinners, sp. nov. (Amaranthaceae). 248.—Lysimachia japonica (Primulaceae)
and Clinopedium gracile (Labiatae) in Louisiana: new to the United States. 249.—New
combinations in Texas Polemoniaceae. 250.—Scutellaria Thieretii (Labiatae), a new species
from coastal Louisiana. 251.—Micranthemum glomeratum (Chapman) Shinners, comb, nov.
(Scrophulariaceae). 252.—Texas Evax transferred to Filago (Compositae). 252.—Verbesina
Walteri Shinners, nom. nov. (Compositae). 253.

REVIEW. Flora of Henry County, Illinois, by Dobbs. 254,

SIDA is privately published by Lloyd H. Shinners, SMU Box 473,
Dallas, Texas 75222, U.S.A. Subscription price $6 (U.S.) per volume of

about 360—400 pages, parts issued at irregular intervals.

SIDA Contributions to Botany volume 1 number 4 pages 187—256
copyright 1964
by Lloyd H. Shinners

BOTANICAL SURVEY ALONG
THE YELLOWKNIFE HIGHWAY,
NORTHWEST TERRITORIES, CANADA

II. VEGETATION
JOHN W. THIERET
University of Southwestern Louisiana, Lafayette

During field work along the Yellowknife Highway in 1958, 1959, 1961,
and 1962 I was engaged in two major procedures: (1) the making of a
plant collection to document the flora, and the recording of distribu-
tional, phenological, life-form, and other data; and (2) the carrying out
of a primary survey of the vegetation, that is, recognizing and describing
the major plant communities and listing their floristic composition. Part
I of Botanical Survey along the Yellowknife Highway, also published
in SIDA (Thieret, 1963), contains a brief introduction, a map of the
highway region, and an annotated catalogue of the flora. The present
paper contains (1) additional data to characterize the region and (2)
descriptions of the following: forest vegetation; vegetation of rock out-
crops; vegetation of lakes and rivers; vegetation of marl and gypsite
deposits; vegetation of strands and islands; vegetation of sand plains;
and vegetation of disturbed soil. I have already described grassland
vegetation of the region (Thieret, 1959). Descriptions of vegetation in
areas adjacent to the highway region have been published by Cody,
1960; Jeffrey, 1961; Moss, 1953a, 1953b, 1955; Porsild, 1945, 1951; Raup,
1935, 1946, 1947; and Thieret, 1961. The reader is referred to these papers
for comparison with the present account.

Climatological data are given in Table 1. The climate of the highway
region can best be described as northern continental, with short, dry,
relatively warm summers and long intensely cold winters. The growing
season is short and hazardous. Freezing and below freezing tempera-
tures may occur even in July and August. Nevertheless, there is an
average frost-free period ranging from 83 days at Fort Providence to
113 days at Yellowknife. Snowfall is heavy, but annual precipitation is
low, averaging about 10 inches, of which somewhat more than half
(about 5.5 inches) falls as rain. Much of the rain (about 3.6 inches) falls
in July and August. More rain falls in autumn than in spring. The low
annual precipitation is compensated for in part by the presence of
permafrost—which retards drainage, accounting for some of the vast
amount of surface water in the region—and by the relatively low sum-
mer evaporation.

SIDA 1 (4): 187—239. 1964.

188

Day length data for Yellowknife are given in Table 2. The length of
the summer days is notable from the standpoint of plant growth. During
the longest days, there is twilight illumination during the night because
the sun remains just below the horizon.

Data and numerous references concerning the geology of the highway
region are given by Douglas (1959) and Raup (1946, 1947) and need

mately the first 66 miles (from Enterprise) the road runs along the
northern edge of the Alberta Plateau (a division of the Great Plains).
Elevation ranges from about 650 to 875 feet. On the great descending
steps of the plateau—and also on the risers—considerable limestone of
Palaeozoic age is exposed. Marly lakes are frequent, as are extensive
peat and lake marl deposits and swift, cold, clear streams with rocky
or sandy beds. Sandy or gravelly soils mantle many of the ridges. One
major river, the Kakisa, averaging 200 feet wide, crosses the highway
at about mile 54. The Kakisa bridge is welcome as affording the only
bit of paved road along the highway outside of Yellowknife. Lady
Evelyn Falls on the river is 47 feet high. It and Kakisa Lake (elevation
729 feet) are accessible via “Kakisa Road,’ which leaves the highway
at mile 53 and goes 10 miles to the lake. The lake, about 25 miles long
and 8 miles wide, has, in the few places I investigated it, a low, rocky,
Table 1. SELECTED CLIMATOLOGICAL DATA FOR YELLOWKNIFE,
Y RIVER, AND FORT PROVIDENCE, N.W.T.*
Yellowknife Hay River
Providence
Mean annual precipitation, 8.45 12.02 9.63
inches (10 years) (29 years) (10 years)
Mean annual rainfall, inches 5.00 7.34 5.35
(10 years) (29 years) (10 years)
Mean rainfall, June, July, 2.90 4.06 4.13
August, inches (10 years) (29 years) (10 years)
Mean temperature, January, F -18 -12 -17
(10 years) (29 years) (9 years)
Mean temperature, July, F 60 60 60
(10 years) (29 years) (9 years)
Extreme high temperature, F 86 96 97
(1941-1960) (1893-1959) (1943-1959)
Extreme low temperature, F -60 -62 -60
(1941-1960) (1893-1959) (1943-1959)
Average frost-free period 113 days 88 days 83 days
(10 years) (53 years) (7 years)

* Data in lines 1 through 5 from Anonymous, 1954; in eae 6 and 7 from Seige
Branch, Department of Transport, Toronto, in lift line 8 from Anonymous, 1956
Lengths of time on which the data are based are piven ars each entry.

189

and sandy shore. Several hundred feet from its southern shore is a
high escarpment in which considerable limestone is exposed.

At mile 66 the highway leaves the plateau and descends to the
Mackenzie Lowlands (also a division of the a Plains), in which it
remains until it enters the Canadian Shield at Frank Channel. The
Mackenzie River, crossed by a ferry, is reached at mile 82: here it is
slightly more than a mile wide and is 513 feet in elevation. After the
crossing, the highway runs atop the river bluff for 4 miles and then
turns northeast. At about mile 4 N, a branch road nearly 4 miles long
runs to Fort Providence.

The Mackenzie Lowlands, an area of low relief, is generally poorly
drained and muskeg covered. The watercourses, for the most part, are
sluggish and have muddy beds. Many of the lakes have marly bottoms,
and there are many small to large deposits of marl. Maximum elevation
along the highway, about 900 feet, is in the area between mile 90 N and
105 N, where sand deposits, outcroppings of limestone, and numerous
sinkholes occur. Gypsum is near the surface, and there are several small
lakes in which gypsite has been deposited. Perhaps the most scenic
portion of the highway is at about mile 120 N, where the road descends
into the valley of Mosquito Creek. Here can be seen a splendid panorama
of massive limestone escarpments, dense forest, and, in the distance, the
waters of Great Slave Lake. Extensive sand deposits are found between
Mosquito Creek and Frank Channel.

Several great Pre-Cambrian outcrops, not easily accessible from the
highway, represent outliers of the Canadian Shield and can be seen
from the road several miles before it enters the shield at Frank Channel.
The 58 miles from the channel to Yellowknife are totally different in
aspect from the remaining 222 miles of the highway. Generally low
rounded outcrops of Pre-Cambrian rock with intervening lakes or
marshes are the dominant feature of the landscape. In the vicinity of
Yellowknife occur large sand deposits.

The Yellowknife Highway lies wholly within the boreal forest al-
though certain limited areas along it are strikingly reminiscent cf th
lichen woodland of the boreal forest-tundra ecotone. The predomi-
nating vegetation is, of course, forest of various types. However, the

Table 2, APPROXIMATE TIMES OF SUNRISE AND SUNSET, AND
DAY LENGTH FOR SELECTED DATES, YELLOWKNIFE, N.W.T.*

Sunset Day length
Date Mountain Standard Time Hrs.-Mins
May 25 3:14 AM 9:45 PM 16 - 48
June 25 2:40 AM 10:38 PM 19 - 58
July 25 3:37 AM 9:47 PM 18 - 10
August 25 5:03 AM 8:13 PM 15 - 10
September 25 6:26 AM 6:27 PM 12 - O01

* Data supplied by H. W. Murdy, United States Bureau of Sport Fisheries and Wildlife.

190

forest cover is by no means continuous but is broken by grasslands, rock
outcrops, shrub communities, numerous lakes and watercourses, and

The fowenae season is brief, lasting only about 2% months. The
first herbaceous species to bloom appear to be Calypso bulbosa and
Anemone patens var. wolfgangiana. By the time of my earliest arrival
at the highway, June 13, Calypso had half-mature-size fruits as well
as flowers; the pasque flower was through blooming, and its fruits were
about half grown. By this time, most willows are past the height of
flowering, as are Arctostaphylos rubra and Rhododendron lapponicum.
The peak of the flowering season is during the last week of June and
the first two weeks of July. Goldenrods i. beginning to bloom
in mid-July, are the harbingers of a fleeting fall, which is well upon
the region by August 1, when the fringed gentians (Gentianella crinita
ssp. macounii and ssp. rawpii) and Lomatogonium rotatum open their
flowers. After these, only two species of Artemisia, unlikely stragglers
(they may well be introductions from areas to the south), are left to
bloom; they come into flower in mid-August.

During the course of the field work, and after I had gained some
familiarity with the local vegetation, attention was concentrated upon
what might be called the “typical” stands of certain eure types,
and notes were made on each such stand observed. It must be empha-
sized that data were gathered only for readily ener vegetation
types and that little serious attention—because of time limitations—was
paid to the many stands that were not of these types and that were
seemingly difficult of classification into definable communities. It is one
thing to recognize and describe “typical” jack pine forest or marly-
lake vegetation and to refer these to community type(s); it is another
altogether to aie to refer to community type a stand in which all
tree species known in the highway region grow in close association or
a stand that is a ae black spruce-Sphagnum forest in all respects
except one: the trees are not black spruces but are jack pines. Such
stands as these actually exist and, with many others that are of differ-
ent nature but are equally or more perplexing, constitute a large per-
centage of the vegetation along the highway. The vegetation types de-
scribed in the following pages make up perhaps a smaller percentage
of the total plant cover than do the vegetation types I have not attempted
to describe. The former are obviously no more important than the
latter—they are merely easier to characterize and categorize.

FOREST VEGETATION

BLACK SPRUCE FOREST
The commonest type of forest in the highway region is that dominated
by Picea mariana. Black spruce forests occupy the wettest of the forest-
dominated areas, ranging from deep, peat-filled (or peat-and marl-

194

filled) depressions to shallow depressions, slopes, and level land. The
forests vary considerably in development and floristic structure. Where
they occur in deep, peat-filled depressions they represent an advanced
stage in hydrarch succession and may be designated as bog (or muskeg)
forests; elsewhere, ey develop where moisture relations favor black
spruce over more mesic tree species.

The bog forest—in its wetter phase—is characterized particularly by
an abundance of Sphagnum (especially S. fuscum and S. warnstorfia-
num; also A. capillaceum var. tenellum), which forms great hummocks
that serve as substratum for the trees and other plants of the forest. In
the wettest places Sphagnum girgensohni and S. squarrosum May occur.

(below which is marl in the case of succession from a marly lake).
Usually a distinct low shrub stratum, composed principally of Ledum
decumbens or L. groenlandicum or both, is present. In the depressions
between the Sphagnum hummocks (where there may be standing water)
can occur various plants (e.g., Triglochin maritima, Carex aquatilis,
Eriophorum chamissonis, Scirpus cespitosus var. callosus, Menyanthes
trifoliata, Utricularia spp.) that do not “belong” to the forest community
but are remnants of preceding successional stages.

A common associate of black spruce in the bog forest is Larix laricina,
larch, which occurs usually as isolated trees (rarely in stands of limited
extent). Among other especially common or characteristic plants of the
forest are: Equisetum palustre, Selaginella selaginoides, Smilacina
trifolia, Tofieldia glutinosa, Ranunculus lapponicus, Drosera rotundi-
folia, Parnassia multiseta, Rubus chamaemorus, Oxycoccus microcarpus,
Vaccinium vitis-idaea var. minus, Pinguicula vulgaris, and Senecio lugens.

The bog forest, in its drier phase, shows ascendancy of woodland
mosses, especially Hylocomium splendens, over the Sphagnum. Hylo-
comium does not grow in hummocks like Sphagnum but produces a
relatively even carpet over the forest floor. Such forests, where Hylo-
comium has become the dominant ground cover and Sphagnum has dis-
appeared altogether or is restricted to isolated hummocks, may be
floristically almost identical with black spruce forests that have de-
veloped on sufficiently wet shallow depressions, slopes, or level land, as
described below.

Black spruce forests that develop on shallow depressions, slopes, or
level land that are sufficiently wet to preclude other forest types are
distinct from the wetter phase of the bog forest in their history and
floristic composition; with the drier phase of the bog forest, as mentioned
in the preceding paragraph, they intergrade floristically. They are char-
acterized particularly by a continuous carpet of Hylocomium splendens
and other woodland mosses. In some black spruce-Hylocomium forests,
Sphagnum is absent; in others it occurs in scattered hummocks. The

192

forest is underlain by a shallow layer of peat, as little as 4 inches, in
contrast with the deep peat under the bog forest. An obvious low shrub
stratum is generally lacking or poorly developed. The plants that are
especially common or characteristic of the bog forest are uncommon or
lacking in the black spruce-Hylocomium forest, which is floristically

thosa, Juniperus communis var. depressa, Zygadenus elegans, Mitella
nuda, Rosa acicularis, Rubus pubescens, Hedysarum spp., Moneses uni-
flora, and Linnaea borealis var. americana.

The many stands of black spruce along the highway include not only

stand that floristically is a fine example of a bog forest—but which has
not developed through hydrarch succession: the Sphagnum hummocks
rest upon only about 6 inches of peat that, in turn, overlies white sand
(the sand here is at least 5 feet deep, as can be seen in the nearby road
cut). The site is on a slight rise, which only adds to the perplexity.
Higher plants (in addition to Picea mariana) observed in black spruce
forests are listed here (those followed by (S) were seen only in bog
forests; those followed by (H) were seen only in black spruce-Hylo-
comium forests; the others were observed in both types): Equisetum
arvense, E. palustre (S), E. scirpoides, E. sylvaticum, Selaginella selagi-
noides (S), Juniperus communis var. depressa (H), J. horizontalis (H),
Larix laricina, Triglochin maritima (S), Calamagrostis neglecta (H),
Oryzopsis pungens (H), Carex buxbaumii (H), C. scirpoidea (H), C.
vaginata (H), Eriophorum chamissonis (S), Scirpus cespitosus var. cal-
losus (S), Smilacina trifolia (S), Tofieldia glutinosa (S), T. pusilla,
Zygadenus elegans (H), Calypso bulbosa (H), Corallorhiza trifida (H),
Cypripedium calceolus var. parviflorum (S), C. guttatum (H), Habenaria

cosa, Rosa acicularis (H), Rubus acaulis (H), R. pubescens (H), Hedysar-
um alpinum var. americanum (H), H. mackenzti (H), Empetrum nigrum,
Viola nephrophylla (H), Shepherdia canadensis, Cornus canadensis, Mon-
eses uniflora (H), Pyrola asarifolia, P. grandiflora (H), P. virens (H), An-
dromeda polifolai, Arctostaphylos rubra, A. wva-ursi (H), Chamaedaphne
calyculata, Kalmia polifolia (H), Leduwm decumbens, L. groenlandicum,
Rhododendron lapponicum, Oxycoccus microcarpus (S), Vaccinium wligi-

193

nosum, V. vitis-idaea var. minus, Castilleja raupt (H), Pedicularis
labradorica, Pinguicula vulgaris (S), Galiwm labradoricum (H), Linnaea
borealis var. americana (H), Viburnum edule (H), Antennaria pul-
cherrima (S), Erigeron hyssopifolius, Senecio lugens (S), and Solidago
multiradiata.

WHITE SPRUCE FOREST

White spruce (Picea glauca) forests occur generally on the mesic
forest-occupied sites, and especially on moist well-drained uplands and
along watercourses. In young stands that have arisen following fire,
the trees are small and closely spaced, the floor is shallowly covered
with litter, and the associated plants are few in species and numbers,
e.g., scattered lichens, Rosa OCleUWIaris, Shepherdia canadensis, Epilobium

cana. Travel through such stands is difficult because of the dense
growth and the brule.

Quite another picture is presented by a mature white spruce forest,
of which the finest example to be seen along the highway is on the
steep slope above Kakisa River about % mile below Lady Evelyn Falls.
The trees are large, the biggest seen being 28.1 inches DBH, about 129
feet tall, and about 183 years old. They cast a dense even shade. The
floor is deeply carpeted with Hylocomiwm splendens into which the
walker sinks 3 or 4 inches at every step. The moss and peat layer is
thick enough so that, when I was trying to dig through it in several
places, I struck frozen peat (about 16 inches down in mid-July) before
I was able to reach mineral soil. Reproduction of the spruce is good.
Cladonia rangiferina occurs in small scattered patches, and Peltigera
apthosa is common. Alnus crispa, to about 8 feet tall, forms a more or
less definite understory. Parasitic on the roots of the alder, and very
rare, is Boschniakia rossica, the only Orobanchacea known in the region.
Other plants in the forest are much scattered and include the shrubs
Juniperus communis var. depressa, Rosa acicularis, Shepherdia can-
adensis, Arctostaphylos rubra, Ledum groenlandicum, Vaccinium vitis-
idaea var. minus, Linnaea borealis var. americana, and Viburnum edule;
and the herbs Cystopteris montana, Carex concinna, Corallorhiza trifida,
Cypripedium guttatum, Geocaulon lividum, Actaea rubra, Hedysarum
mackenzii, Mitella nuda, Moneses uniflora, Pyrola asarifolia, and P.
grandiflora.

On the slope above the south shore of the Mackenzie River (mile
80-81) the white spruce forest is younger than the Kakisa stand, being
composed of trees 45-55 years old. Balsam poplar is frequent. About
75 per cent of the floor is covered with deep litter (3 to 5 inches), the
rest with mats of Hylocomium splendens about 5 inches thick and a few
small patches of Cladonia. The moss cover is more complete in the most

194

densely shaded part of the forest, where Hylocomium and Cladonia are
the only plants other than the trees. In addition to the plants seen in
the Kakisa stand( but with the exception of Boschniakia rossica), the
folowing were observed here: Calypso bulbosa, Habenaria obtusata,
Salix myrtillifolia, Ribes lacustre, Fragaria virginiana var. terraenovae,
Rubus acaulis, Lathyrus ochroleucus, Empetrum nigrum, Pyrola virens,
and Pedicularis labradorica.

The white spruce forest richest in secondary species, and the most
open one seen, is on the slope above Mosquito Creek. e floor is densely
carpeted with Hylocomium splendens; Tomenthypnum nitens is frequent,
as are Peltigera aphthosa and Cladonia. Sphagnum hummocks are oc-
casional. Considerable brule, well decayed, is present. The following sec-
ondary species, none common, were observed here: Equisetum arvense,
E. scirpoides, Juniperus communis var. depressa, J. horizontalis, Carex
capillaris, C. capitata, C. gynocrates, C. vaginata, C. scirpoidea, Tofieldia
pusilla, Zygadenus elegans, Calypso bulbosa, Corallorhiza trifida, Cypri-
pedium passerinum, C. guttatum, Habenaria obtusata, Orchis rotundi-
folia, Salix glauca, S. myrtillifolia, S. reticulata, Myrica gale, Geocaulon
lividum, Anemone parviflora, Dryas Hg Moe Potentilla fruticosa,

Rosa acicularis, Hedysarum alpinum var. americanum, Empetrum nig-
TUM, eae canadensis, Moneses uniflora, ae grandiflora, P.
virens, Andromeda polifolia, Arctostaphylos rubra, Ledum groenlandi-

cum, xycoccus microcarpus, Rhododendron lapponicum, Vaccinium
uliginosum, Vaccinium vitis-idaea var. minus, Castilleja raupii, Pedi-
cularis labradorica, Linnaea borealis var. americana, Viburnum edule,
and Solidago multiradiata.

The Kakisa and Mackenzie stands of white spruce are typical of most
stands of white spruce of similar age that are to be seen along the
highway; the Mosquito Creek stand is distinctly atypical because of its
openness and relatively large number of secondary species. Many stands
of white spruce are closely similar to mature stands of jack pine except
for the presence or relatively greater abundance of Hylocomium splen-
dens. White spruce forests, then, are characterized typically by a ground
cover of Hylocomiuwm splendens or of this moss and thick litter; by
relatively dense shade; and by a low number of second i
and of individuals of these species. Many of them have scattered balsam
poplars. In stands of white spruce other than those previously men-
tioned, the folowing additional species were found: Rhytidiwm rugosum
(common as ground cover, with Hylocomiwm splendens, at mile 22.7:
this species, inadvertently omitted from the Catalogue, was determined
by William D. Reese), Cladonia alpestris, Elymus innovatus, Ribes triste,
Astragalus americanus, Arctostaphylos uwva-ursi, Lonicera dioica var.

_—

glaucescens, and Galium septentrionale.
xamples of mature white spruce forest examined by me ap-
peared to be self-perpetuating, i.e., good reproduction of spruce was

195

shown and no other tree species appeared as invaders. Following fire,
the spruce may regenerate itself, or its place may be taken by jack pine
or aspen. White spruce may form pure stands, or forests of this species
may contain few to many individuals of other tree species, notably jack
pine, aspen, and poplar, but also larch and black spruce. Similarly, white
spruce may occur in forests that are primarily of other species.

JACK PINE FOREST

Dry sandy or gravelly ridges and flatlands are typically covered with
forests in which Pinus banksiana, jack pine, is the sole or by far the
commonest tree species. Many acres, nas recently burned, are
dominated by a scrubby growth of pine on mineral soil. Here, walking
may be exceedingly onerous because of the brule and the dense growth
of young trees. Such sites may be barren of plants except for the pines
or there may be occasional Rosa acicularis, Shepherdia canadensis,
Epilobium angustifolium, Arctostaphylos uva-ursi, Vaccinium vitis-idaea
var. minus, and Linnaea borealis var. americana. As such forests age,
there is gradually increasing mesophytism. Woodland lichens, herbaceous
plants, and shrubs return and increase in abundance; a litter and

and are characterized by rather widely spaced trees that are 35-55 feet
tall, at least 40 years old, and, for the most part, even-aged. The under-
growth in such forests is scanty. The floor may be covered largely with
litter or with litter and humus or it may show extensive development
of lichens, especially fruticose Cetraria and Cladonia. The most common
and characteristic low woody plants are Rosa acicularis, Shepherdia
canadenis, Arctostaphylos uva-ursi, and Vaccinium vitis-idaea var.
minus. The Arctostaphylos and Vaccinium are common to abundant as
ground-cover plants. Calamagrostis purpurascens and Elymus innovatus
are characteristic grasses. In places, Alnus crispa, to about 8 feet tall,
forms a definite understory. Jack pine appears not to reproduce well in
the shade, so young pines are not to be found in the forest.

Jack pine forest of this type occupies the upper parts of ridges and
the most xerophytic of the flatlands. In such sites it may be regarded as
an edaphic or pyric climax, being maintained not only by the xero-
phytism of the habitat but also by recurrent fires. In such a habitat I
found no jack pine forest more than 70 years old. It is not uncommon
to see, on one part of a sandy ridge, a mature pine forest as just
described and, on an adjacent part of the ridge, a burned-over area
with still-standing dead trees and a dense growth of small young pines.
These trees may bear cones when they are only 3 feet tall.

On more mesophytic sites (e.g., the lower flanks of ridges) the suc-
cessional trend is obviously toward the replacement of jack pine by
white spruce. As the pine forest matures, the litter and humus layer

196

deepens, woodland mosses (especially Hylocomium splendens) appear in
scattered patches, and Picea glauca becomes established. In such a
forest there are no young pines, but a lower stratum of young white
spruces is a frequent sight. Barring fire, one may expect ascendancy of
the spruce at the expense of the pine. In a well developed white spruce
forest, it is not unusual to see large, old, and plainly dying pines—the
remnant of the pine forest that once occupied the site. Young pine forests
that are burned appear most commonly to be succeeded by an abundant
and immediate regrowth (from seed) of pine; burned mature pine
forests appear to be succeeded commonly by aspen.

The following plants were collected or observed in jack pine forests:
Equisetum scirpoides, Lycopodium complanatum, Juniperus communis
var. depressa, J. horizontalis, Bromus puwmpellianus, Calamagrostis pur-
purascens, Elymus innovatus, Festuca saximontana, Oryzopsis asperi-
folia, O. pungens, Carex daenea, C. foenea, Zygadenus elegans, Calypso
bulbosa, Corallorhiza trifida, Populus balsamifera, P. tremuloides, Salix
bebbiana, S. glauca, Alnus crispa, Betula glandulosa, B. papyrifera,
Geocaulon lividum, Arenaria capillaris, Anemone multifida, A. patens
var. wolfgangiana, A. parviflora, Aquilegia brevistyla, Ribes lacustre,
Amelanchier alnifolia, Fragaria virginiana var. terrae-novae, Potentilla
fruticosa, Rosa acicularis, Astragalus americanus, Hedysarum alpinum
var. americanum, H. mackenzti, Lathyrus ochroleucus, Oxytropis splen-
dens, Empetrum nigrum, Hudsonia tomentosa, Shepherdia canadensis,
Epilobium angustifolium, Cornus canadensis, Pyrola asarifolia, P. sec-
unda, P. virens, Arctostaphylos uva-ursi, Ledum groenlandicum, Vac-
cinium vitis-idaea var. minus, Apocynum androsaemifolium, Pedicularis
labradorica, Galiwm septentrionale, Linnaea borealis var. americana,
Lonicera dioica r. glaucescens, Viburnum edule, Campanula rotundi-
folia, Arnica lonchophylla, Aster ciliolatus, A. sibiricus, Erigeron glabel-
lus var. pubescens, Hieracium uwmbellatum, Senecio tridenticulatus, and
Solidago spathulata var. neomexicana.

DECIDUOUS FORESTS

Three species of deciduous trees occur in the highway region: Betula
papyrifera, Populus balsamifera, and P. tremuloides. The first of these,
Betula papyrifera (paper birch), typically occurs scattered among Picea
glauca and Pinus banksiana; locally on sand plains it may form small
pure stands. In contrast, the two species of Populus occur not only as
scattered individuals among other trees but they also may form exten-
sive pure stands.

Populus tremuloides (aspen) stands appear to arise primarily follow-
ing burning of more or less mature jack pine or white spruce forests. It
is not at all unusual to find an island of unburned pine or spruce that
is surrounded by burned forest in which a vigorous stand of young
aspens is developing. Among the aspens the secondary flora may con-.

197

tain many of the species found in the adjacent unburned forest, e.g.,
Elymus innovatus, Geocaulon lividum, Fragaria virginiana var. terrae-
novae, Rosa ene ne Rubus pubescens, Shepherdia canadensis, Cornus
canadensis, Vaccinium vitis-idaea var. minus, and Viburnum edule.
Epilobium angustifollum is generally frequent to comm as

shrubby willows, especially Salix bebbiana. Typically ae is ae
reproduction of aspen, but young white spruces—frequently with a
mixture of Populus balsamifera—may be common. A mature aspen
woods is floristically similar to a mature jack pine woods. It may con-
tain la old pines, which apparently are trees that survived the fire
that destroyed the original pine forest. Populus balsamifera, poplar,
forms pure stands on flood plains, where a poplar stage immediately
precedes white spruce. Such stages can be seen along the Hay River,
which parallels the Mackenzie Highway in northern Alberta and in the
Northwest Territories. Along the Yellowknife Highway a “typical”
flood plain (but one of quite limited area) was seen only along the
Kakisa River just below Lady Evelyn Falls. Otherwise, poplar occurs
generally in mixture with white spruce; it may locally be dominant or

nated areas is similar to the flora of the adjacent areas that are domi-
nated by white spruce.

VEGETATION OF ROCK OUTCROPS
LIMESTONE OUTCROPS

Along the Enterprise-Frank Channel section of the highway many
outcroppings of limestone occur. The exposures may be vertical only—
as on the faces of the gorge immediately below Lady Evelyn Falls—
or both vertical and horizontal—as in many places between miles 10 and
28 where the descending great limestone “stairs” of the northern edge
of the Alberta Plateau are exposed over large areas. Outcrops occur also
between miles 80 N and 107 N—here the exposures are largely horizontal
or gently inclined—and in the cliffs near mile 127 N, where the slope
below the limestone cliff is littered with small to huge limestone blocks
and slabs. My notes on the flora of limestone outcrops were derived
principally from investigation of horizontal exposures at miles 20 to
28, 80.8 N, 96.5 N, 103.5 N, and 107 N

Limestone outcrop areas are characterized by much exposed rock and,
in places, by the presence of a fine, dark brown, residual, non-calcareous
soil. The soil may be confined to crevices or may exist in a shallow
layer (typically 1 to 4 inches deep) over the rock. A large percentage
of such soil is typically bare of plant cover. In some areas, a litter layer
up to about 1 inch deep may accumulate. Lichens are common to
abundant in outcrop areas, both on bare rock and on soil. The most
apparent species are the fruticose Cladonia alpestris, C. mitis, C. rangi-

198

ferina, and Cetraria nivalis. An occasional patch of Cetraria tilesii is
conspicuous because of its bright yellow color.

rees, when present, are typically widely spaced. Pinus banksiana and
Populus tremuloides are the commonest species; Picea glauca is less
common; and Picea mariana, Populus balsamifera, and Betula papy-
rifera are uncommon. Many outcrop areas have no trees on them.
Especially characteristic of the outcrops is the conspicuousness of
the chamaephytes Juniperus horizontalis, Saxifraga tricuspidata, and
Arctostaphylos uva-ursi. In many places the long (up to eet)

h

forming a distinctive network on the rock. Saxifraga tricuspidata forms
circular mats up to 2 feet across, often with the innermost portion of
the mat dead. The prostrate branches of Arctostaphylos wva-ursi reach
6 feet in length. Of the erect shrubs on limestone outcrop areas,
Juniperus communis is the most common; it forms on many sites a
distinct low-shrub stratum. Other common shrubs are Amelanchier
alnifolia, Potentilla fruticosa, Prunus pensylvanica, Rosa acicularis, and
Shepherdia canadensis. Prunus virginiana, notable because it is here
at the northermost known portion of its range, is local, forming thickets
or occurring as isolated shrubs.

On the limestone cliffs at Lady Evelyn Falls, and on the moist faces
of limestone escarpments between miles 10 and 28, the ferns Cystopteris
fragilis, Dryopteris robertiana (seen only at mile 15.5) and Woodsia
glabella are found and may be common locally. The Cystopteris and the
Woodsia occur also at ee meets Creek cliffs, along with Crypto-
gramma crispa va olypodium virginianum, and Woodsia
ilvensis (the ee ae three, heen are much more common on

Pre-Cambrian rocks). On the Mosquito Creek cliffs the few trees are
Picea glauca, P. mariana, and Betula papyrifera. Salix glauca, to about
8 feet tall, is present. The most common plant there is Dryas integrifolia,
which forms mats on both vertical and horizontal rock faces. Two
crucifers, Draba cinerea and Lesquerella arctica var. scammanae, and
a sedge, Carex glacialis, found in crevices, were collected only at
Mosquito Creek.

Vascular plants collected or observed on limestone outcrops are:
Cystopteris fragilis, Dryopteris robertiana, Polypodium virginianum,

oodsia glabella, W. ilvensis, Juniperus communis var. depressa, J.
horizontalis, Picea glauca, P. mariana, Agropyron trachycaulum, Agrostis
scabra, Calamagrostis inexpansa, C. neglecta, C. purpurascens, Des-
champsia cespitosa, Elymus innovatus, Festuca saximontana, Helicto-
trichon hookeri, Koeleria cristata, Oryzopsis pungens, Poa alpina, P.
canbyi, P. glauca, P. interior, Carex bebbi, C. concinna, C. glacialis,
C. scirpoidea, Juncus alpinus, J. balticus var. littoralis, Allium schoeno-
prasum var. sibiricum, Zygadenus elegans, Sisyrinchium montanum,
Populus tremuloides, Salix arbusculoides, S. bebbiana, S. glauca, Betula

199

papyrifera, Arenaria capillaris, A. dawsonensis. A. rubella, Cerastium
arvense, Melandrium ostenfeldii, Anemone multifida, A. parviflora, A.
patens var. wolfgangiana, Arabis divaricarpa, A. holboellii, Draba cin-
erea, D. lanceolata, Lesquerella arctica var. scammanae, Ribes oxyacan-
thoides, Saxifraga tricuspidata, Amelanchier alnifolia, Dryas drummondii,
D. integrifolia, Fragaria virginiana var. terrae-novae, Geum triflorum,
Potentilla arguta, P. fruticosa, P. nivea ssp. hookeriana, P. pensylvanica,
Prunus pensylvanica, P. virginiana, Rosa acicularis, Hedysarum mac-
kenzii, Oxytropis campestris var. varians, O. splendens, Linum lewisit,
Shepherdia canadensis, Cornus canadensis, C. stolonifera, Arctostaphylos
uva-ursi, Androsace septentrionalis, Dodecatheon pulchellum, Gentian-
ella amarella ssp. acuta, Castilleja raupii, Rhinanthus crista-galli, Plan-
tago septata, Galium septentrionale, Lonicera dioica var. glaucescens,
Campanula rotundifolia, Achillea lanulosa, Antennaria parvifolia, Arnica
Artemisia campestris ssp. borealis, Aster alpinus var.
vierhapperi, A. ciliolatus, A. hesperius var. laetevirens, Erigeron com-
positus var. glabratus, E. glabellus var. pubescens, Helenium autumnale,
Hieracium umbellatum, Senecio tridenticulatus, Solidago spathulata var.
neomexicana, and Taraxacum ceratophorum.

PRE-CAMBRIAN OUTCROPS

Along the Frank Channel-Yellowknife section of the highway Pre-
Cambrian outcrops become a dominant feature of the landscape. These
outcrops, with perpendicular to gently sloping” sides and with rounded

ascend in a series of stair-like levels. The bases of the outcrops are over-
lain with mineral soil (principally silt and clay, but also some sand and

mariana, Populus tremuloides, and Betula papyrifera, and thickets of
Salix spp., Alnus crispa, Rosa acicularis, Shepherdia canadensis, and
Viburnum edule.

The rock composing the outcrops is largely granite-gneiss, granodio-
rite, and granite except in the Yellowknife area, where volcanic, sedi-
mentary, and metasedimentary rock types occur. The outcrops show a
wide range of colors, including pink, dark green, dark grey, and almost
black. The natural color of the rock is, however, obscured by weathering
and, even more important, by an abundance of saxicolous lichens. Of
these the pale dull greens and yellows of Cladonia and Cetraria, the
ashy grey of Parmelia, and the blacks of Actinogyra and Lasallia are
the most characteristic. The rock tripes (i.e., Actinogyra and Lasallia),
black when dry, become a dark olive when they get wet. Outcrops
covered with rock tripe become markedly different in aspect during the
first few minutes of a rainfall: they change slowly—but most percepti-
bly—from black to dark olive as the lichens absorb moisture. On out-

200

crops covered with other lichens, the colors become more intense during
a rain. Lichens collected on Pre-Cambrian outcrops include: Cladonia
alpicola, C. cornuta, C. degenerans, C. metacorallifera, C. mitis, C. pyxi-
data, C. rangiferina, C. uncialis, C. verticillata, Actinogyra muhlenbergii,
Lasallia pensylvanica, Cetraria nivalis, Parmelia centrifuga, P. steno-
phylla, and P. sulcata

Crustose lichens and rock tripe are best represented on cliff faces,
slopes, and the most exposed level places; fruticose lichens tend to be
most abundant in more sheltered places, especially shallow depressions,
where a few inches to several square feet may be covered by an un-
broken mat of Cladonia and Cetraria. In somewhat deeper depressions,
xerophytic matted moses (especially Polytrichum formosum, P. juni-
perinum, P. piliferum, and Hedwigia ciliata) tend to be dominant over

under mosses appears to be greater than under lichens. Such soil serves
as a seed bed for higher plants.

The vascular flora of Pre-Cambrian outcrops is poor in species and,
for the most part, in individuals. The outcrops offer two types of
habitats for vascular plants: (1) rock crevices, and (2) depressions. The
commonest trees are Pinus banksiana, Picea mariana, and Betula papy-
rifera; Picea glauca and Larix laricina are infrequent. These trees grow
either in crevices (where jack pine, white spruce, and white birch are

6 and 20 feet tall, although larger specimens can be found (e.g., the
largest jack pine found by me on a Pre-Camprian outcrop was about
40 feet tall and 13 inches DBH; it had 135 annual rings). In the most
exposed situations, only jack pine and black spruce are found; here
they are gnarled, sometimes almost prostrate or with all but the lowest
branches killed.

In rock crevices wherein a little soil has accumulated, the commonest
vascular plants are Cryptogramma crispa var. stichoides, Dryopteris
fragrans, Polypodium virginianum, Woodsia i vensis, Juniperus com-
munis var. depressa, J. onigonial = Saxifraga tricuspidata, Empetrum
nigrum, Arctostaphylos uva-ursi, and Vaccinium vitis-idaea var. minus;
less common to rare are Festuca saximontana, Carex canescens, C.
supina, Melandrium ostenfeldii, Potentilla multifida, P. nivea ssp. hook-
eriana, and P. pensylvanica. These are also the first of the higher plants
to become established in lichen and moss mats in depressions. It is not
unusual to see a moss- or lichen-dominated depression (with a well-
formed soil layer under the mat) in which just one or two species of
herbaceous vascular plants (and often only one or two individuals) are
growing, e.g., Agrostis scabra, Calamagrostis canadensis, Poa interior,
Carex aenea, C. chordorrhiza, Eriophorum, angustifolium, E. brachy-

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antherum, Smilacina trifolia, Geocaulon lividum, Corydalis sempervirens,
and Arabis holboellii. A considerable layer of peaty soil (2 feet in one

calyculata. Less common to rare shrubs are Salix bebbiana, S. pyrifola,
Alnus crispa, Ribes glandulosum, R. oxyacanthoides, Amelanchier alni-
folia, Dryas drummondii, Potentilla fruticosa, Rosa acicularis, Rubus
idaeus var. canadensis, Shepherdia canadensis, Vaccinium wuliginosum,
and Viburnum edule. The most common tree is Picea mariana. Among
other plants found in such depressions are Equisetum sylvaticum, Rubus
chamaemorus, Epilobium angustifolium, Artemisia campestris ssp. bore-
alis, and Solidago spathulata var. neomexicana,

VEGETATION OF LAKES AND RIVERS
MARLY LAKES

Along several sections of the Great Plains portion of the highway
are many small lakes that are notable because of the large amount of
lake marl (CaCO,) precipitated from their water and because of the
consequent deposit of marl that develops on the bottom. Many of these
marly lakes were studied between miles 40 and 66 and between miles
40 N and 90 N. The marly lakes occur in areas where calcareous bed-
rock is exposed. Carbonates dissolved from the rock by streams passing
over it are carried into the lakes. Precipitation in the lakes is brought
about by plant life, which uses carbon dioxide, causing supersaturation.
A sample of marl from a small drained lake at mile 66 was sent to
the Geological Survey of Canada for analysis. Dr. J. Terasme in-
formed me that the sample was 90% or a little more acid soluble
and that it contained a rich diatom flora, including, among the many
genera and species, Navicula tuscula, Achantidium flexellum, Pinnularia,

and many kinds of pollen and spores. Pinus banksiana was the pre-
dominating pollen (80-90%); others included Picea glauca (15-20%),
P. mariana (1-3%), various Cyperaceae, and Shepherdia canadensis.
Spores included Selaginella, Lycopodium, Equisetum, and Sphagnum
and other mosses.

For the purpose of this discussion, I recognize two types of marly
lakes: (1) those around which a sedge mat develops; and (2) those
with gravelly-marly beaches.

The marly lakes found between miles 40 and 66 exhibit the develop-
ment of a sedge mat. Open water in these lakes varies from only a few
square feet in the nearly obliterated lakes to several acres in the larger
ones. The water is crystal clear, and, even in the largest lakes, only a

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foot or less of this water overlies the marl on the “bottom.” Actually,
the marl on the “bottom” is far from solid; it is, in contrast, loosely
suspended in the water. It offers little resistance to the passage of objects
(stick, stone, hand, etc.) through it. In some lakes, the deposit of marl
is about 3 feet thick; in others, I was unable to reach the underlying
layer with an 8 foot pole. Underlying the marl (at least those deposits
whose bottom I was able to reach with a pole) is a hard layer, probably
limestone.

The vegetation around marly lakes can be grouped into four more
or less concentric zones: (1) vegetation of open water; (2) a sedge zone,
Le., the foremat, which is dominated solely by sedges; (3) a shrub zone,
the next older portion of the mat, which is dominated by various low
shrubs; and (4) a typical muskeg forest of black spruce.

In the marly lakes the following plants were found free-floating in
shallow water beyond the edge of the mat: the mosses Campylium stel-
latum and Scorpidium scorpioides; Utricularia intermedia, and U. vul-
garis. Plants rooted, albeit loosely, in marl in shallow water beyond the
mat were: Equisetum pdlustre, Sparganium minimum, Potamogeton
filiformis var. borealis, Triglochin maritima, T. palustris, Carex aquatilis,
C. lasiocarpa, C. limosa, C. physocarpa, Eriophorum angustifolium, E.
chamissonis, E. viridi-carinatum, Scirpus validus, Juncus stygius var.
americanus, Drosera anglica, and Menyanthes trifoliata.

The mat is built up primarily through the activity of rhizomatous
species of Carex, especially C. aquatilis, C. buxbaumii, and C. lasi iocarpa.
Carex limosa is important in places. The rhizomes of these species may
extend 2 to 3 feet into open water beyond the edge of the mat. The
mats may extend themselves quite rapidly over open water: in August
of 1959, the inner edge of a mat at mile 44 was 37 feet from a marked
small tree of Larix laricina: in August of 1961 it was 40 feet away from
the tree. In most marly lakes it is possible to walk—or rather to wade—
to within 4 or 5 feet of the leading edge of the mat without breaking
through the hummocky tangle of rhizomes, roots, culms, and ves
comprising the mat. The mat becomes considerably firmer in its older
portions because of the build-up of organic matter and the continued

infrequent to rare, are Triglochin maritima, T. palustris, Muhlenbergia
glomerata var. cinnoides (on anthills), Carex capillaris, C. diandra, C.
interior, C. rostrata, Eriophorum angustifolium, E. viridi- carinatum,
Tofieldia glutinosa, T. pusilla, Parnassia multiseta, Potentilla palustris,

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Andromeda polifolia, Pinguicula vulgaris, Utricularia intermedia, U.
vulgaris, Galium labradoricum, and Lobelia kalmit.

The woody plants that dominate the shrub zone typically begin to
appear as isolated individuals in the sedge zone, especially toward the
outer edge of this zone. Myrica gale is generally the most common in-
vader, although Salix athabascensis, S. candida, and S. serissima occur
also. In the shrub zone itself, the commonest species are Myrica gale,
Betula glandulosa, and Potentilla fruticosa; less common are the willows
Salix athabascensis, S. candida, S. pedicellaris, and S. serissima. The
sedges of the preceding zone persist to some degree, but the plants are
for the most part scattered and not vigorous, especially in the older
parts of the shrub zone. The shrubs grow on peaty hummocks (which
were, of course, formed by the sedges), and the hummocks themselves
are underlain by several inches of additional peat; below the peat is
marl, Other than the shrubs, relatively few plants grow in the shrub
zone. Probably the most common are mosses, of which I collected four
species: Ditrichum flexicaule, Dicanum bergeri, Campylium stellatum,
and Tomenthypnum nitens. Occasional clumps of Sphagnum occur in the
older parts of the zone, as do occasional small trees of Larix laricina
and Picea mariana.

Succeeding the shrub zone is eae a muskeg forest of Picea
mariana, perhaps with some Larix laricina. The transition between the
shrub zone and the muskeg forest is generally a rather abrupt one. At
the outer edge of the shrub zone the moss mounds of the forest rise 1
o 2 feet. One literally steps up into the muskeg forest. The mounds may
be largely of Sphagnum, especially S. fuscwm and S. warnstorfianum,
or of other mosses, of which Tomenthypnum nitens appears to be
common.

At about mile 65 N and miles 37 and 64 are marly lakes that do not
exhibit the formation of a sedge mat, but have, in contrast, wide marly-
gravelly beaches. The water in these lakes is deeper and the bottom is
firmer than in the sedge-mat lakes. In shallow water along the shore
of these lakes the plants noted were Chara contraria, C. contraria var.
hispidula, Potamogeton filiformis var. borealis, and Scirpus validus. On
the lower beach grew Triglochin maritima, T. palustris, Scolochloa
festucacea, Carex aquatilis, C. atherodes, Eleocharis pauciflora var.
fernaldii, and Scirpus validus. On the upper beach were collected
Calamagrostis neglecta, Carex aurea, C. garberi, C. scirpoidea, C. viri-
dula, Salix brachycarpa, S. candida, Betula glandulosa, Rumex mariti-
mus var. fueginus, Ranunculus sceleratus, Parnassia multiseta, Potentilla
anserina, Rubus idaeus var. canadensis, Cicuta douglasti, Epilobium glan-
dulosum var. adenocaulon, Dodecatheon pulchellum, Primula incana,
Gentianella amarella ssp. acuta, Gentianella crinita ssp. raupit, Lomato-
gonium rotatum, Mentha arvensis var. villosa, Castilleja raupii, Aster

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brachyactis, A ciliolatus, A. junciformis, Erigeron lonchophyllus, Sene-
cio congestus, and S. pauperculus.

MUCK-BOTTOM LAKES

The commonest type of lake along the Canadian Shield section of
the highway is characterized by a bottom of muck (i.e., organic

panocladus spp., Calla palustris, Potentilla palustris, and Menyanthes
trifoliata are important components, and by the abundance of Nuphar
variegatum.

In open water beyond the edge of the mat the most obvious plant is
Nuphar variegatum, whose relatively large leaves, floating on the
surface, may cover a significant percentage of the water. Nuphar grows
in water up to 5 feet deep. Nymphaea tetragona var. porsildii was found
by me among Nuphar in only one lake, although Ray Murdy tells me
that in 1963 he observed the species “here and there” in almost all of
the medium to large lakes he studied along the highway from mile 10 S
to 39 S. Other rooted aquatics with floating leaves are infrequent but
may be locally prominent: Polygonum amphibium, Caltha natans, and
Potamogeton gramineus. Submerged aquatics, inconspicuous but some-
times abundant, include Potamogeton alpinus var. tenuifolius, P. foliosus,
P. friesii, P. pusillus, P. richardsonii, P. zosteriformis, Ceratophyllum
demersum, Myriophyllum exalbescens, M. verticillatum var. pectinatum,
Utricularia intermedia, U. minor, and U. vulgaris. The mosses Drepano-
cladus capillaceus and D. exannulatus are common to abundant on the
mucky bottom. Locally, Lemna trisulea forms great masses just below
the surface of the water, and Lemna minor is common. Emergent
aquatics, generally found in the shallow water just beyond the edge
of the mat, are few, the most common being Equisetum fluviatile,
Sparganium angustifolium, S. minimum, Hippuris vulgaris, and Meny-
anthes trifoliata. Others are Sagittaria cuneata, Alopecurus aequalis,
Scirpus validus, Cicuta douglasti, and Senecio congestus.

The mat around muck bottom lakes is composed largely of rhizomatous
species of Carex, especially C. aquatilis, C. lasiocarpa, C. limosa, and C.
rostrata. The mosses Drepanocladus aduncus, D. capillifolium, D. exan-
nulatus, and Meesia tristicha are also common to abundant. In many
sites, a conspicuous zone of Calla palustris, Potentilla palustris, or
Menyanthes trifoliata occurs at the leading edge of the mat and is
sharply distinct from sedges of the mat because of its different color
and coarser foliage. These three species, where they occur, are most
important contributors to the mat. Their thick tangled rhizomes extend
2 to 3 feet into open water, as do those of the sedges. Calamagrostis
canadensis is common, especially but not solely in the firmer portions
of the mat. Locally, certain other rhizomatous plants are important:
Equisetum fluviatile, Typha latifolia, Eriophorum angustifolium, Eleoch-

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aris palustris, Glycerta grandi, G. pulchella, Eriophorum angustifolium,

canescens, C. diandra, Polygonum amphibium, Rumex occidentalis, Stel-
laria crassifolia, Ranunculus sceleratus, R. gmelinii, Parnassia multiseta,
Cicuta bulbifera, C. douglasii, Epilobium glandulosum var. adenocaulon,
Naumburgia thyrsiflora, Scutellaria galericulata var. epilobifolia, Aster
junciformis, and Senecio congestus. In at least some of the lakes, perma-
frost is only a foot or so below the mat, even in late summer.

Succession beyond the sedge mat appears to culminate in either
a white spruce-Hylocomium forest (infrequently) or a black spruce-
Sphagnum forest (commonly). In succession to white spruce, the sedge
mat is invaded first by shrubs, commonly Myrica gale, Chamaedaphne
calyculata, or willows (Salix bebbiana, S. glauca, S. planifolia, S. scoul-
eriana, or a mixture of these.) Alnus tenuifolia occurs occasionally.
Thickets of these shrubs may occupy extensive areas around shield lakes.
The shrub stage is succeeded by or intermingled with a stage in which
Betula papyrifera is dominant but in which some Populus tremuloides
may occur. In the Betula stage (and sometimes in the shrub stage),
young white spruces appear. The succeeding white spruce stage is char-
acterized by an abundance of woodland mosses (especially Hylocomium
splendens) and by the presence of such “typical” plants as Equisetum
arvense, Alnus crispa, Salix glauca, Mitella nuda, Ribes triste, Empet-
rum nigrum, Cornus canadensis, Moneses uniflora, and Pyrola grandi-
flora. In succession to white spruce, bog mosses (Sphagnum) do not
appear to enter into the sere as they do in succession to black spruce.

Succession to black spruce is characterized mainly by the invasion of
the mat by Myrica gale, Chamaedaphne calyculata, Ledum groenlandi-
cum, and sometimes Betula glandulosa, and, in addition, usually by
Sphagnum spp. Encroaching upon this shrub stage are mounds of
Sphagnum that support “typical” black spruce forest vegetation.

Almost every conceivable variant of the above seres can be observed.
One of the commonest is the invasion of the mat by willows and the
encroachment upon the willows by the black spruce-Sphagnum com-
munity. Sphagnum does not appear here until it is seen in the Sphagnum
mounds of the forest stage.

The water level of many of the lakes adjacent to the highway has
been lowered by ditching; some lakes have been drained completely. In
addition to the submerged plants that are thus left stranded to die, and
to the emergent plants that may persist, the following were observed on
such exposed and drying muck: Agrostis scabra, Carex canescens, Cc.
diandra, C. paupercula, Eleocharis acicularis, Juncus vaseyt, Salix sp.
(seedlings), Rumex maritimus var. fueginus, R. occidentalis, Ranunculus
gmelinii, R. sceleratus, Stellaria crassifolia, Barbarea orthoceras, Rorippa
islandica, Potentilla norvegica, Potentilla palustris, and Galium trifidum.

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Many muck-bottom lakes in the Mackenzie River-Frank Channel
section of the highway show vegetation similar to that of muck-bottom
lakes in the Canadian Shield section, i.e, a sedge mat followed by a
shrub stage and finally a black spruce-Sphagnum forest. Conspicuously
lacking, however, are Calla palustris and Nuphar variegatum: Potentilla
palustris and Menyanthes trifoliata, moreover, are much less important
than they are in lakes on the shield.

SANDY- OR MUDDY-BOTTOM LAKES AND RIVERS

In the highway region, aquatic vegetation is especially luxuriant in
the Stagg River (mile 49 S). Here, in still or slowly flowing clear water,
the following plants were observed: Potamogeton gramineus, P. richard-
soni (in water to 4 feet deep), Sagittaria cuneata, Sparganium angusti-
folium, Eleocharis acicularis, Lemna minor, L. trisulea (forming great
masses on the bottom), Ranunculus aquatilis var. eradicatus, Callitriche
hermaphroditica, C. palustris, Naumburgia thyrsiflora, Utricularia inter-
media, U. minor, U. vulgaris, and Bidens cernuda.

In still, protected areas along the shore of Kakisa Lake and Kakisa
River, Potamogeton gr amineus, P. richardsonii, P. vaginatus, and Hip-

Sparganium eke cta. Sagittaria seis Phalaris pn ee

gonum amphibium, Ranunculus circinatus var. subrigidus, Caiericns
hermaphroditica, C. palustris, Myriophyllum exalbescens, Sium suave,
and Utricularia vulgaris. At the outlet of Kakisa Lake into Kansas River
the water is choked with masses of Potamogeton.

Three aquatics were collected only in the vicinity of Yellowknife:
Isoetes echinospora var. braunii (in Prelude Lake), Subularia aquatica
(in Prelude and Prosperous lakes), and Limosella aquatica (in Prosper-
ous Lake). Additional aquatics observed in the highway region in sandy
or muddy substrata include Sparganium minimum, Glyceria borealis,
G. pulchella, G. grandis, G. striata, Scolochloa festucacea, Juncus bu-
fonius, J. filiformis, Polygonum lapathifolium, Potentilla palustris, and
Myriophyllum verticillatum var. pectinatum.

MAN-MADE EXCAVATIONS

In roadside ditches and man-made pools, all of which are yet rela-
tively new, only a few species can be found, and generally only a few
individuals of each. Observed in such habitats were Typha latifolia,
Sparganium angustifolium, S. minimum, Potamogeton alpinus var. tenui-
folius, P. filiformis, P. pusillus, Sagittaria cuneata, Arctagrostis latifolia,
Lemna minor, Ranunculus circinatus var. subrigidus, R. gmelinu, Caltha
natans, Polygonum amphibium, Callitriche hermaphroditica, C. palustris,
Elatine triandra, Myriophyllum exalbescens, M. verticillatum var. pecti-

207

natum, Hippuris vulgaris, and Senecio congestus. Of these, Typha lati-
folia is the commonest. Many roadside pools have nothing but a colony
of cattails in them. Most of the excavations that contain water have
bottoms of mud—exceedingly soft mud.

VEGETATION OF MARL AND GYPSITE DEPOSITS

During the course of construction of the highway, a number of marly
lakes, both south of the Mackenzie River and in the Mackenzie River-
Frank Channel section, were partly or completely drained, exposing
sizeable marl deposits. Certain marl deposits, such as the extensive one
at mile 39.7 N, apparently were exposed by natural lowering of lake
water level. A drained lake above the highway at mile 119 N has in
its bed a deposit of gypsite at least 3 feet thick. The deposit is surrounded
by a fine example of bog forest; in the gypsite itself, only one plant
(Juncus balticus var. littoralis) was growing, probably because of the
short time the lake had been drained.

In marl] deposits the plants occur in very open associations. In the
deposit at mile 30, recently exposed by draining, a rather weedy flora
has developed and is confined to the periphery of the deposit where the
marl is driest. Toward the center of the deposit is standing water.
Walking on the drier marl is much like walking on a firm, thick, sponge-
rubber mattress; the walker sinks in the wetter marl. The following
plants were observed here: Calamagrostis canadensis, Hordeum jubatum,
Carex diandra, C. media, Eleocharis palustris, Scirpus validus, Juncus
alpinus, Urtica dioica var. procera, Arenaria dawsonensis, A. lateriflora,
Ranunculus gmelinii, R. sceleratus, Arabis hirsuta var. pycnocarpa, Des-
curainia sophia, Rorippa islandica. Potentilla norvegica, Hippuris vul-
garis, Epilobium glanduloswm var. adenocaulon, Phacelia franklinii,
Campanula rotundifolia, and Senecio congestus. Of these, the Epilobium
was by far the commonest plant, giving to the band of vegetation rim-
ming the deposit a reddish cast because of its anthocyanous stems and
leaves.

As an example of a mari deposit in a long-naturally-drained lake bed,
the one at mile 110.5 N may be offered. The marl is much fissured, and
large areas are bare of plants. Occasional pieces of gypsum occur in
the marl. The most obvious plants are low, scattered shrubs: Salix
brachycarpa, S. candida, S. glauca, Betula glandulosa, Potentilla fruti-
cosa, Shepherdia canadensis, and Arctostaphylos wva-ursi. Other plants
are relatively many in species but few in numbers of individuals of
each species: Bryum lacustre. Equisetum arvense, Triglochin maritima.
T. palustris, Deschampsia cespitosa, Poa interior, Carex aquatilis, C.
atherodes, C. capillaris, C. concinna, C. garberi, C. scirpoidea, C. viri-
dula, Scirpus cespitosus var. callosus, S. pumilus var. rollandiu, Habe-
naria hyperborea, Arenaria humifusa, Melandrium ostenfeldit, Anemone
parviflora, Braya humilis, Dryas integrifolia, Castilleja raupu, Erigeron

208
hyssopifolius, E. lonchophyllus, and Solidago spathulata var. neomexi-

na.

Additional plants observed in other marl deposits are: Calamagrostis
neglecta, Phalaris arundinacea, Poa glauca, P. pratensis, Puccinellia
nuttalliana, Scolochloa festucacea, Carex physocarpa, C. sartwellii, Atri-
plex patula, Rumex maritimus var. fueginus, Lepidium bourgeauanum,
Potentilla norvegica, P. pensylvanica, Lomatogonium rotatum, Mentha
arvensis var. villosa, Scutellaria galericulata var. epilobiifolia, Achillea
lanulosa, Antennaria parvifolia, Aster brachyactis, A. ciliolatus, A. junci-
formis, A. pansus, Lactuca pulchella, Senecio indecorus, S. pauperculus,
and Taraxacum ceratophorum.

VEGETATION OF STRANDS AND ISLANDS
KAKISA RIVER

Along the rocky shore of the Kakisa River and on gravel bars and
islands in the river, four zones of vegetation may be distinguished: (1)
a fore zone, characterized by the absence of woody plants and by local
dominance of sedges (especially Carex aquatilis), Calamagrostis, or
Phalaris arundinacea, and by scattered other herbaceous plants: (2) a
shrub zone, characterized by dominance of Salix, Cornus stolonifera,
and Alnus tenuifolia; (3) a deciduous-tree zone, characterized by domi-
nance of Populus balsamifera; and (4) an evergreen-tree zone, char-
acterized by Picea glauca and developing into a typical white spruce
forest community. These zones intergrade somewhat, and all of them
are not necessarily present at every site.

Detailed study was made of three low rocky islands in the Kakisa
River about 3 miles below Lady Evelyn Falls. One of these islands
supported only herbaceous vegetation of the “fore zone”; one, some-
what higher, had a central zone of shrubs in addition to the fore zone:
the third, the highest, had a fore zone and a central shrub zone and,
in this, a central band of Populus balsamifera in which several young
white spruces (ca. 3 feet tall) were growing.

On island I, and at the periphery (fore zone) of the other two islands,
were collected or noted all of the taxa of herbaceous plants that I found
elsewhere in similar habitats along the river. Locally, in the fore zone
vegetation, Carex aquatilis, Calamagrostis canadensis, C. neglecta, and
Phalaris arundicnacea grew in large patches. Other plants of this zone,
typically much scattered and in several cases represented by only one
or two individuals, were: Equisetum fluviatile, E. palustre, Typha lati-
folia, Agropyron trachycaulum, Beckmannia syzigachne, Deschampsia
cespitosa, Poa glauca, Carex rostrata, Juncus balticus var. littoralis,
Chenopodium capitatum, C. glaucum var. salinum, Polygonum aviculare,
P. lapathifolium, Rumex maritimus var. fueginus, Arenaria dawsonensis,
Stellaria crassifolia, S. longipes, Ranunculus macounii, R. reptans, R.
sceleratus, Corydalis aurea, Arabis hirsuta var. pycnocarpa, Cardamine

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pensylvania, Erysimum cheiranthoides, Rorippa islandica, Potentilla an-
serina, P. norvegica, P. palustris, Geranium bicknellii, Cicuta douglassi,
Sium suave, Epilobium glandulosum var. adenocaulon, Hippuris vulgaris,
Naumburgia thyrsiflora, Gentianella crinita ssp. macounu, Mentha arven-
sig var. villosa, Stachys palustris var. nipigonensis, Veronica peregrina,
V. scutellata, Plantago major, and Galium trifidum.

The shrub zones on islands II and III were dominated by a dense
growth of Salix bebbiana, S. candida, S. lasiandra, S. planifolia, Alnus
tenuifolia, and Cornus stolonifera. Among the shrubs grew only a few
herbs: Zygadenus elegans, Arenaria lateriflora, Parnassia multiseta,
Geum macrophyllum var. perincisum, Hedysarum alpinum var. ameri-
canum, Cornus canadensis, Castilleja raupii, and Aster junciformis. Rosa
acicularis occurred on island III. The shrubs formed a dense thicket and
ranged from 4 to 25 feet tall (the latter height being reached only by
Salix lasiandra and S. planifolia). The poplars on island III were
growing among tall willows and were about 35 feet high and 8 inches

Similar stages of vegetation can be observed in many places along the
Kakisa River. Just below Lady Evelyn Falls, the lower part of the
“flood plain” is dominated by willows and alder; in this zone can be
found small balsam poplars. The upper part of the plain is occupied by
a typical white spruce-Hylocomium forest. The transition area between
the willow-alder zone and the spruce forest exhibits particularly lush
growth of shrubs and herbs, including Actaea rubra, Aquilegia brevi-
styla, three species of Ribes, Amelanchier alnifolia, Cornus canadensis,
Cornus stolonifera (to 8 feet tall), Pyrola grandiflora, and Viburnum
edule (to 5 feet tall).

MACKENZIE RIVER

The bank of the Mackenzie River at Fort PE OviIocnee and at mile 4 N

which is sandy silt. At the lower ee of the bank, the silt is damp and
may be moss covered; higher up it is drier and may exhibit cracks. The
habitat offered for plants is a rigorous one subject to much ice and
water erosion; yet a surprising variety of plants grows on the river
banks. In places, seemingly every available bit of space between the
boulders is occupied by plants. The most obvious plants are shrubs, of
which the following were noted: Salix myrtillifolia, S. serissima, S.
bebbiana, S. planifolia, Alnus tenuifolia, Ribes oxyacanthoides, Amelan-
chier alnifolia, Potentilla fruticosa, Rosa acicularis, Rubus pubescens,
Elaeagnus commutata, and Cornus stolonifera. Some small Populus bal-
samifera occur on the bank. The herbaceous flora, richer toward the
base of the bank, contained the following taxa: Equisetum arvense,
Agropyron trachycaulum, Agrostis scabra, Beckmannia syzigachne, Cal-

210

amagrostis canadensis, C. inexpansa. Hordeum jubatum, Poa leptocoma,
P. palustris, Sphenopholis intermedia, Carex aquatilis, C. atherodes, C.
garberi, Eleocharis acicularis, Juncus balticus var. littoralis, J. alpinus,
J. bufonius, Allium schoenoprasum var. sibiricum. Smilacina stellata,
Sisyrinchium montanum, Arenaria lateriflora, Silene menziesii, Poly-
gonum aviculare, Rumex maritimus var. fueginus, Anemone canadensis,
A. multifida, Ranunculus macounti, R. reptans, R. sceleratus, Thalietrum
venulosum, Erysimum cheiranthoides, Capsella bursa-pastoris, Rorippa
islandica, Fragaria virginiana var. terrae-novae, Potentilla anserina, P.
norvegica, Vicia americana, Viola nephrophylla, Epilobiwm glandulosuwm
var. adenocaulon, Naumburgia thyrsiflora, Gentianella crinita var. raupii,
Collomia linearis, Stachys palustris var. nipigonensis, Ewphrasia aff.
subarctica, Rhinanthus crista-galli, Plantago major, Galium septentrion-
ale, Achillea lanulosa, A. sibirica, Arnica chamissois, Aster junciformis,
Hieracium umbellatum, Lactuca pulchella, and Solidago canadensis var.
salebrosa.
SANDY OR GRAVELLY BEACHES

Sandy gravelly beaches were observed only at Kakisa Lake and in
the vicinity of YelNowknife. Such habitats are hazardous ones for plants
because of ice accumulation and movement and because they are sub-
ject to flooding during high water in spring and to drying out later in
the growing season.

At Kakisa Lake no woody plants were noted on the fore beach: the
following herbaceous plants were seen: Equisetum fluviatile, E. variega-
tum, Triglochin maritima, Calamagrostis inexpansa, Deschampsia cespi-
tosa, Eleocharis acicularis, E. palustris, Scirpus validus, Juncus alpinus,
J. balticus var. littoralis, J. bufonius, J. nodosus, Sisyrinchium montanum,
Chenopodium glaucum var. salinum, Polygonum amphibium (stranded),

P. lapathifolium, Ranunculus gmelinii, R. reptans. R. sceleratus, Erysi-
mum cheiranthoides, Rorippa islandica, Potentilla anserina, P. norvegica,
Epilobium glandulosum var. adenocaulon, Sium suave, Gentianella crinita
var. macounii, Veronica scutellata. Plantago major, Achillea sibirica,
Aster junciformis, and Erigeron philadelphicus. These plants are all
much scattered, except for the rhizomatous grasses and sedges, which
form small colonies. The upper beach is characterized by the presence
of low willows, Salita lasiandra and S. planifolia, and by a more con-
tinuous cover of grasses and sedges, especially Carex aquatilis and
Calamagrostis. Above the upper beach is a zone of tall willows and
Alnus tenuifolia in which an occasional poplar occurs and in which
Smilacina stellata, Zygadenus elegans, Thalictrum venulosum, Hedy-
sarum alpinum var. americanum, and woodland mosses were noted,

On sandy beaches of Prosperous and Prelude lakes, Frame Lake, and
Yellowknife Beach (all near Yellowknife) the following taxa were
noted: Sparganium minimum (stranded), Triglochin maritima, T. palus-
tris, Sagittaria cuneata, Eleocharis acicularis, Scirpus validus, Juncus

211

alpinus, J. bufonius, J. filiformis, J. vaseyi, Alopecurus aequalis, Beck-
mannia syzigachne, Glyceria borealis, G. grandis, Puccinellia distans,
Carex aquatilis, C. rostrata, Polygonum lapathifolium, Rumex fueginus,
Ranunculus cymbalaria, R. macounii, R. reptans, Sium suave, Veronica
peregrina, Plantago major, Aster brachyactis, and Gnaphalium uligi-
nosum.
VEGETATION OF SAND PLAINS

Occurring in the vicinity of Yellowknife and also just west of the
shield margin at Frank Channel are areas that may be designated “sand
plains.” Here, on white or tan sand, the trees are Pinus banksiana, Picea
glauca, P. mariana, and Betula papyrifera. In places, these species may
all grow together; in other places, each may form pure stands. The trees
are widely spaced and low branched, producing a parklike aspect. Con-

covered with litter or, more commonly, with a dense greenish yellow
mat of fruticose Cladonia and Cetraria. The plains that are dominated
by such lichen woodland are strongly reminiscent of the parklike timber
nearer to the continental limit of trees to the northeast.

Shrubby and herbaceous plants on the sand plains are few. The most
common and characteristic shrubs are Arctostaphylos uva-ursi, Ledum
decumbens, L. groenlandicum, and Vaccinium vitis-idaea var. minus.
These may occur together or, more frequently, as isolated plants or small
colonies. Vaccinium vitis-idaea var. minus appears to be the first woody
plant to become established in blowouts. Other shrubs collected on
sand plains are Salix glauca, Rosa acicularis, Empetrum nigrum, Shep-
herdia canadensis, Hudsonia tomentosa, and Vaccinium uliginosum. Of
these, Empetrum nigrum and Hudsonia tomentosa are the most fre-
quently encountered. Empetrum forms mats up to 3 feet across; Hudsonia
is especially eae of the sand plains near the Yellowknife air-
port, where locally it is common, growing alone in otherwise bare sand
or with en uva-ursi or Vaccinium vitis-idaea var. minus.

Herbaceous plants collected on sand plains are Equisetum sylvaticum,
Lycopodium annotinum, Calamagrostis purpurascens, Festuca saximon-
tana, Geocaulon lividum, Cornus canadensis, Astragalus striatus, Carex
foenea, C. supina, Apocynum androsaemifolium, and Linnaea borealis
var. americana.

VEGETATION OF DISTURBED SOIL
In the highway region, areas disturbed through the activity of man
are mainly in the vicinity of the settlements (Enterprise, Fort Provi-
dence, Fort Rae, and Yellowknife) and along the highway right-of-way.
To me, the outstanding characteristics of the vegetation of these sites
are two: (1) the small number of introduced plants present; and (2)
the persistence of native plants after disturbance.

212

Of all the species collected in disturbed habitats, only the following
are certainly introduced: Agropyron cristatum, A. repens, Bromus
mermis, Phalaris canariensis, Phleum pratense, Puccinellia distans, Poly-
gonum aviculare, Polygonum convolvulus, Axyris amaranthoides, Chen-
podium glaucum var. salinum, Stellaria media, Brassica campestris,
Capsella bursa-pastoris, Descurainia sophia, Erysimum cheiranthoides,
Thlaspi arvense, Melilotus alba, M. officinalis, Lappula echinata, Galeop-
sis tetrahit var. bifida, Crepis tectorum, Matricaria maritima var. agrestis,
Senecio vulgaris, Sonchus arvensis var. glabrescens, Tanacetum vulgare,
and Taraxacum officinale. These taxa, comprising about 5.5% of the
total vascular flora, are decidedly uncommon along the highway right-
of-way; indeed, finding one of them there is somewhat of an “event.”
Some of them, notably Polygonum aviculare and Taraxacum officinale,
become more frequent in the vicinity of settlements. With the exception
of Puccinellia distans, Polygonum aviculare, Chenopodium glaucum var.
salinum, and Erysimum cheiranthoides, which grow sometimes in un-
disturbed habitats and appear native, all of them were found only in
disturbed soil. It is to be expected, of course, that as time passes, these
weeds will become more common along the highway (the oldest sections
of which had been completed only five years when I last worked along
them). Many of them are common to abundant along the older Mac-
kenzie Highway.

What might be called the “settlement” flora is well represented at
Enterprise (mile 0). Here, in disturbed sandy soil, occur many species
of weedy plants, some native, some introduced. A similar flora can be
found in waste places around each of the settlements. At Enterprise the
species observed were: Equisetum scirpoides, Agropyron trachycaulum,
Agrostis scabra Hordeum jubatum, Oryzopsis pungens, Phleum pratense,
Poa pratensis, Puccinellia nuttalliana, Carex aenea, Zygadenus elegans,
Axyris amaranthoides, Chenopodium berlandieri var. zschackei, C. cap-
itatum, Polygonum achoreum, P. aviculare, Rumex mexicanus, Cerastium
nutans, Silene menziesu, Stellaria longipes, Aquilegia brevistyla, Cory-
dalis aurea, Arabis holboelu, Capsella bursa-pastoris, Descurainia sophia,
Erysimum cheiranthoides, Lepidium, densiflorum, Thlaspi arvense, Frag-
aria virginiana var. terrae-novae, Potentilla norvegica, Melilotus alba,
Vicia americana, Geranium bicknellu. Epilobium angustifolium, E. gland-
ulosum var. adenocaulon, Phacelia franklinii, Lappula echinata, L. red-
owski var. occidentalis, Galeopsis tetrahit var. bifida, Moldavica parvi-
flora, Campanula rotundifolia, Galium septentrionale, Achillea lanulosa,
Aster ciliolatus. Aster sibiricus, Matricaria matricarioides, Crepis tecto-
rum, Solidago canadensis var. salebrosa, and Taraxacum officinale.

To list all the native plants that persist—and often do well indeed—in
disturbed areas would require considerable space, for such a list would
contain perhaps 90% of the species. Some of these just “manage to
survive,” being of reduced vigor in disturbed soil, e.g., Smilacina trifolia.

213

Calypso bulbosa, and Spiranthes romanzoffiana, which can be found on
mounds of disturbed peat in clearings. Many others, in contrast, do
exceedingly well and respond to their altered habitat with vigor. Out-
standing examples are: Allium schoenoprasum var. sibiricum, Zygadenus
elegans, Arabis divaricarpa, Fragaria virginiana var. terrae-novae, Poten-
tilla norvegica, Androsace septentrionalis, Campanula rotundifolia, Achil-
lea lanulosa, Aster ciliolatus, Solidago multiradiata, and S. spathulata
var. neomexicana. These species are generally more to much more
robust and abundant in disturbed areas than in adjacent undisturbed
ones, where they may be rare and depauperate by comparison. The
Potentilla, Campanula, Achillea, Aster, and two species of Solidago are
in many places locally abundant in the right-of-way and are quite con-
spicuous when they are in flower.

ACKNOWLEDGEMENTS
Work along the Yellowknife Highway during 1961 and 1962 was ae ee by a
grant (G15904) from the National Science Foundation. The 1958 and 59 work was
upported by Chicag atur i chester Hansen, Gerald I ae and

James Delahoussaye gave especially valuable service in

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REFERENCES

ANONYMOUS. 1954. Addendum to Volume 1 of Climatic Sumpinatics for Selected
Meteorological Stations in Canada. Average oS of opeeeree and Precipitation. Me-
teorological Division, Deparument of Transport, TVoron

ANON OUS. 1956. Climatic Summaries for ee Meteorological ag hae in a
Volume 3. Frost Data. ee Division, Department of Transport, Toror

CODY, W. J. 1960. Plants of the vicinity of Norman Wells, Mackenzie District, ak
west Territories. a Field-Nat. 74: 71-100

DOUGLAS, R. J. W. 1959. oe Slave i Trout River map-areas, Northwest Terri-
tories. Geol. Surv. Canada Paper 58-11

JEFFREY, W. W. 1961. — on plan occurrence along lower Liard River, N.W.T.
Nat. Mus. Canada Bull. 171: -115

MOSS, E. H. 1953a. Forest communities in northwestern Alberta. Canad. Jour. Bot.

252

19536. Marsh and bog vegetation in northwestern Alberta. Canad.
Jour. Bot. 31: 448-470.

1955, The vegetation . Alberta Rev 4

ORSILD, A. E. 1945. The alpine flora of the east oe of eee a ee North-
west Territories. Nat. Mus. Canada Bull.
—______—. 1951. Botany of een Yukon adjacent to the Canol Road. Nat.
Mus. Canada Bull. 121.

] , H. M. 1935. Botanical investigations in Wood Buffalo Park. Nat. Mus. Canada
Bull. 74

——________—. 1946. er ee studies in the Athabaska-Great Slave Lake Re-
gion, II. Jour. Arn. Arb. 27: 1-8

—_________—.. 1947. Bo nope of southwestern Mackenzie. Sargentia

THIERET, J. W. 1959. oe vegetation near Fort Providence, ee est Territories.
Canad. Field-Nat. 73: 161-167.

—________——. 1961. A election of Eee from the Horn Plateau, District of Mack-
enzie, Northwest Se eano ss. Cana i Field 75%. 77-83%

ee, 3. Botanical survey eae the oo Highway, Northwest
Territories, Canada. I. Catalogue "of the flora. Sida 1: 117-170

PLATE. J

(Upper) The Yellowknife Highway, mile 11. Forest of Pinus banksiana
and Picea glauca. Limestone is just below the surface here, as is evi-
denced by tripod telephone poles (holes could not be dug to set poles).

(Lower) Picea mariana-Hylocomium forest along Kakisa Road, with
road right-of-way in foreground.

215

PLATE I

(Upper) Picea glauca-Hylocomium forest with scattered Populus
tremuloides, mile 76. Highway in foreground.

(Lower) Interior of above forest, showing thick carpet of Hylocomium
splendens.

217

PLATE JI]

(Upper) Young Pinus banksiana forest, mile 30. Highway right-of-
way in foreground.

(Lower) Mature Pinus banksiana forest, mile 34. Elymus innovatus,
Calamagrostis purpurascens, and lichens (Cladonia, Cetraria) common.

219

220

PLATE -1Y

(Upper) Edge of limestone outcrop area, mile 24. Juniperus communis,
J. horizontalis, and Arctostaphylos uva-ursi common to abundant. Picea
glauca-Pinus banksiana forest in background.

(Lower) Juniperus horizontalis on limestone, mile 24.

222

PLATE V

(Upper) Limestone cliff, with talus, mile 66 S. Trees mainly Picea
glauca,

(Lower) Pre-Cambrian outcrop about 10 miles east-northeast of Yel-
lowknife. Trees are Picea glauca, P. mariana. Pinus banksiana, and
Betula papyrifera. Lichens abundant.

223

PLATE VI

(Upper) Pre-Cambrian outcrop, mile 37 S. Top of outcrop darkened
by lichens (Actinogyra and Lasallia).

(Lower) Upper portion of above outcrop. Pinus banksiana on left,
Betula papyrifera to left of center. Lichens (especially Cladonia, Cet-
raria, Actinogyra, and Lasallia) abundant.

226

PLALE. Vi)

Marly lake, mile 44, showing floating islands. Muskeg forest in back-
ground,

LOG

PLATE VIII

(Upper) Muck-bottom lake adjacent to Canadian Shield section of
highway. Nuphar variegatum common and in flower. Photo courtesy H.
W. Murdy, United States Fish and Wildlife Service.

(Lower) Extensive sedge mat around muck-bottom lake, Canadian
Shield section of highway. Photo courtesy H. W. Mudry, United States
Fish and Wildlife Service.

229

PLATE IX
Low island in Kakisa River just above highway bridge. Dominant
plants are Carex aquatilis, Calamagrostis canadensis, and Phalaris arundi-
nacea.

PLATE X
Sandy gravelly beach of Kakisa Lake near outlet into Kakisa River.
On mid-beach are willows; behind these is dense growth of taller willows
and Alnus tenwifolia.

PLATE x1

(Upper) Bouldery shore of Mackenzie River 4 miles east of Fort
Providence. Shrubs are mainly willows and Cornus stolonifera.

(Lower) Extensive deposit of lake marl, mile 39.7 N.

PLATE XI

(Upper) “Sand plain,’ mile 128 N. Pinus banksiana on right, Betula
papyrifera on left. The white sand floor is carpeted with mounds of
lichens, especially Cladonia and Cetraria.

(Lower) Close-up of lichen carpet on the “sand plain” pictured above.

237

238

PLATE XIII

(Upper) Edge of “sand plain” near Yellowknife. Low plant on sand

in foreground is Hudsonia tomentosa; trees in background are Pinus
banksiana and Picea glauca.

(Lower) Calamagrostis grassland (prairie) with adjacent Salix-
Populus woods, mile 16 N

SYSTEMATIC STATUS OF
AMMOPURSUS OHLINGERAE (COMPOSITAE) '

OLGA LAKELA
University of South Florida, Tampa

Lacinaria Ohlingerae was described by Dr. Blake (Bull. Torr. Bot.
Club 50: 203-204, pl. 9, 1923) from sand-scrub of central Florida. Dr.
Small, upholding the endemic as generically distinct, defined its limits
under the binomial Ammopursus Ohlingerae (1. c. 51: 392-393, 1924).
Dr. Robinson, following adoption of Liatris as a nomen conservandum
under the International Code, reinstated sand-torch in the large genus
of the blazing-stars as Liatris Ohlingerae (Contrib. Gray Herb. 104: 49,
1934). This treatment was followed by Dr. L. O. Gaiser in her scholarly
work on the genus Liatris (Rhodora 48: 373-375, 1946).

The underground parts of Liatris. regardless of whether stems or roots,

stipe-like tapering base and the intact pappus. The seedlings with
oblanceolate cotyledons and a single encircling band of roothairs just
below the soil line, were potted in deeper sand and placed under glass
without artificial hea

The period of aonmeney was surprisingly short. An effort was made
to a. some evidence of correlation in natural environment. On the
14th of January 1962, the white sand-scrub of Josephine Creek, Highlands
Co., was thoroughly searched for viable fruits for a chromosome recor
The tufts of faded florets clinging to receptacles were sterile; in fhe
fallen ones caught in the lichens a few ripe achenes were found. During
hours of search over a large area, three seedlings with expanded coty-
ledons were observed. They were somewhat younger than the ones
under cultivation, in which the tip of the first radical leaf was appearing
between the cotyledons.

boas No. 9, Botanical Laboratories, University of South Florida,
SIDA 1 (4): 240-—247. 1964

241

The growth of the planted seedlings was continuous, without well
defined cluster of radical leaves. Caulescence appeared early; the plants
in May were returned to the slath house open to rains, but reduced
light intensity. By September the tallest one had attained the height of
1.6 dm., including the panicle of 12 heads. The plants, six in all, flowered
the first year.

Fruits of Liatris Garberi were discovered on the 4th of January 1962,

cal leaves at the crown of the taproot. Six became caulescent and
flowered the first year; one remained in rosette stage developing a
terminal bud in true Liatris fashion; its underground parts consisted of
9 thickened secondary roots. Cross sections revealed a pentarch xylem,
and 2 rings of growth in the secondary xylem. The plants in each series,
were transferred to the greenhouse garden, where they flowered through
October.

Dr. Small described the root of Ammopursus as “long perpendicular,
fleshy tap-root.” (Man. S. E. Fl. 1933). Actually, it is a chain of seg-
ments formed by thickening of the secondary roots in succession, singly
or in multiples. Two-three segments were formed during the first year
of growth. The crisp, fleshy segments are fragile at joints. The root of
an older plant with two flowering stems, as depicted in Pl. 1, is seldom
seen intact: two scars indicate the places of chains of segments lost in
collecting. One’s groping fingers may feel the whole, suspended in the
mesh of rootlets of Sabal Etonia, in porous, sun-heated sand, but on
exposure only detached parts may be at hand.

The root featured in Pl. 1 was collected in white sand-scrub near
Frostproof, Polk County, 23rd of November 1961. Dr. J. B. Carlson, Uni-
versity of Minnesota, Duluth, executed the illustrations with interpreta-
tion. The freehand sections were stained in safranin and traced under
microprojector.

Section A represents an unthickened root with tetrarch primary

xylem; in section B, the primary tissues become disorganized; the sec-
ondary phloem, mostly of parenchyma, includes patches of sieve tubes.
Similarly, the secondary xylem consists mostly of parenchyma with

essentially the same as to structural content. No rings of growth are
apparent. Section G is a stem, the base of the hypocotyl.

The floral structures have been adequately described in previous
works. Annotations with photographs may emphasize differentiating
characters. Up to 25 florets are borne on the truncate, clavate receptacle.
The 4-5 -seriate inner phyllaries are crowded on an axis barely 2 mm.
long, between the glandular, deeply alveolate apex and the persistent

ho
ro
Nw

outer phylaries (Pl. 2 A & B). Plane and thin, barely thickened at the
base, they readily spread and dissever from the slender connections
with the axis. The loose-appearing head is in sharp contrast with com-
pact heads of L. cymosa and L. cylindracea. In these the outer phyllar-
les are convex on the back, thick and gibbous at base with broad
rhomboidal attachment. They are persistent, strongly imbricated and
appressed against the axis, 4-5 mm. long. The receptacle is convex
with florets on more than one plane. The two above species with L.
Ohlingerde constitute Series Cylindraceae as defined by Gaiser, In
character of the receptacle, they are fundamentally unlike. So is L.
scariosa with a broadly convex receptacle which in Dr. Blake’s concept
is closely related to L. Ohlingerae.

The generic limits as defined by Dr. Small include floral character-
istics. The photographs (PI. 3 and Pl. 4A) bring out clearly zygomorphy
of the urceolate corolla and included stamens. The blue-violet anthers,
only 2 mm. long on subulate filaments about the same length, are
exceedingly short for a floret up to 30 mm. long in life. The pale apices
of anther connectives are 0.1 mm. long. The proportionately longer
anthers, often with conspicuous connectives, exsert in Liatris, or at

a
bundle at the bottom of an alveolus (Pl. 4B). The fruit coat with
ribs tapers to the point and remains attached through germination. In
Liatris the achene base tapers less with looser attachment of the fruit

pappus is barbellate, as described by Dr. Small. Moreover, fleshiness
or succulence of the roots and the foliage were manifestly conclusive
characters of Ammopursus; he regarded the genus on a par with
Garberia and Carphephorus. Short period of dormancy, continuous
growth of the seedling without notable formation of rosette leaves
and the distinctive structure of the achene are herewith added as de-
limiting characters for upholding the systematic status of Ammopursus.

The chromosome pattern, 2n=20 is recorded in a root tip cell. (PI.
4C). The same number has been recorded for species of Trilisa, Eupator-
vum and Liatris

ACKNOWLEDGEMENTS
The cn _ to extend thanks to Dr. J. B. Carlson for preparing the figures in
I, and to Dr. Peter Raven for determining the chromosome record, which appear here

a

cecua, Dee courtesy

PLATES

I. Root of Ammopursus Ohlingerae. All cross sections are X 4.44.
(See text, p. 239.)

II. Head of Ammopursus Ohlingerae. A Receptacle and phyllaries. B.
Flowering head. (Slightly enlarged.)

Ill. Flowers of Ammopursus Ohlingerae. (Slightly enlarged.)

IV. Ammopursus Ohlingerae. A. Single corolla and stamens. B. Achene.
(About four times natural size.) C. Chromosomes, from root-tip cell.

244

245

LVG

NOTES

FATOUA VILLOSA (MORACEAE) IN LOUISIANA: NEW TO NORTH
AMERICA.—In habit, Fatoua villosa (Thunb.) Nakai (F. pilosa Gaud.),
an eastern Asiatic species, is disturbingly reminiscent of the Urticaceae.
In the Lafayette area the plant behaves as an annual and occurs as a
weed in greenhouses and in flower and vegetable gardens. I first noted
it in flower beds and among shrubs on the campus of the University of
Southwestern Louisiana. Seedlings were frequent on the campus this
past spring, even following the severe winter of 1962-63, when the

New Orleans as a wee or at least 15 years. I am indebted to Dr.
Carroll E. Wood, Jr., of the Arnold Arboretum, for the identification
of our material and for various data concerning it. Fatoua villosa is a
most aberrant member of the Moraceae in being herbaceous, in having
one style arm so reduced that it is easily overlooked, and in having a
rather cymose pistillate inflorescences that looks urticaceous. It has in its
leaves punctate cystoliths that are especially prominent in dried speci-
mens. Some Urticaceae have such cystoliths; apparently only punctate
cystoliths are found in the Moraceae. Voucher specimens, bearing my
field numbers 10219, 10221, and 1 ave been cas in the
herbaria of the University of Southw exter Louisiana, o
Methodist University, and of Harvard University.—John W. oie ret,
University of Southwestern Louisiana, Lafayette.

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3

AMARANTHUS MICROPHYLLUS SHINNERS, SP. NOV. (AMAR-
ANTHACEAE).—A. album floribus seminibusque, A. blitoidewm habitu
prostrato accedit; ab ambobus recedit magnitudine minore, foliis per-
parvis brevissime petiolatis, tepalis enervibus. Monoica parva prostrata
ramosa caulibus 5—16 cm. longis glabris vel parce minute puberulis.
Folia maiora 1.0—1.6 em. longa glabra subsessilia vel brevipetiolata
petiolis 1—5 mm. longis laminis obovatis vel elliptico-oblanceolatis
albonervosis obtusis mucronatis. Flores axillares crebri parvi bracteis
1.3—2.5 mm. longis lanceolatis subscariosis vel scarioso-marginatis aris-
tato-acuminatis. Florum marium tepala 3 hyalina ad 2 mm. longa ovata
concava enervia breviter aristato-acuminata; antherae 1 mm. longae.
Florum foemineorum tepala 1 vel 2 hyalina 0.6—1.1 mm. longa lanceo-
lata vel ovata aristato-acuminata; styli 3. Fructus compressus laevis
circumscissilis. Semina nigra 1.0—1.2 mm. longa. HOLOTYPE: silty soil,
dried bottom and edges of pond, Hueco Tanks, Hueco Mountains, El
Paso Co., Texas, U. T. Waterfall 6620, Aug. 20, 1946 (SMU). PARATYPE:
edge of gravel pit north of Waterton River, Transition zone, elev.
4250 feet, Waterton Lakes National Park, Alberta, Canada, August J.
Breitung 17588, Aug. 16, 1953 (SMU). Both collections had been referred

SIDA 1 (4): 248. 1964.

249

to A. blitoides S. Watson (for which Fernald, followed by most American
authors, mistakenly adopted the name A. graecizans L., which properly
belongs to a Mediterranean species). In its prostrate habit A. micro-
phyllus suggests a small form of that species, but it does not have the
expectable 5 tepals, short floral bracts, or large seeds. In floral features
it is very close to A. albus, in which the pistillate flowers have 3 tepals,
at least the two larger with midrib, and the floral bracts usually greatly
exceeding the tepals. The new species is distinctive in having only 1
or 2 ribless' tepals in the pistillate flowers, and seeds intermediate in
size between those of the other two (0.7—1.0 mm. in A. albus, 1.3—1.8
mm. in A. blitoides) —Lloyd H. Shinners.

LYSIMACHIA JAPONICA ee AND CLINOPODIUM
ISIANA: O TH T

recorded before from the United States or, indeed, from the New World.
Voucher specimens of each have been placed in the herbaria of the
University of Southwestern Louisiana and of Southern Methodist Uni-
versity.

Lysimachia japonica Thunb. St. Landry Parish: common for about 200
feet in narrow strip of ground between edge of dirt road and road-
side ditch, in woods just southeast of Washington, John W. Thieret
16120, 3 July 1963. The plant was first collected here in November 1962
by Mr. Edward L. McWilliams. For the specific determination we are
indebted to Dr. Lloyd H. Shinners. A second collection extends the
range of this species to Iberia Parish: rare in disturbed area along oil
pipeline right-of-way through upland woods, Avery Island, Thieret
16380, 27 Sept. 1963. According to Bentvelzen (in Flora Malesiana, Ser.
I, vol. 6 (2): 183, 1962), Lysimachia japonica is known from “Kashm
to Yunnan, Siam, Eastern China, Formosa, Ryukyu Is., Japan, also in

W

field notes for no. 16120 are as follows: “Corolla rotate, bright yellow,
ca. 8—9 mm. wide when fully open. Petals as long as sepals. Capsule
globose, light brown. Pedicels reflexed in fruit. Some stems erect, some
elongate prostrate.” The plant suggests a small, pubescent form of the
well-known Lysimachia nummularia L.

Clinopodium gracile (Benth.) O. Kuntze. Lafayette Parish: weed

29 April 1963. Iberia Parish: in center of little used dirt road through
woods, Jefferson Island, Thieret 15943, 23 May 1963. St. Martin Parish:
at edge of willow dominated depression along Highway 90 about 7
miles south of Broussard, Thieret 17063, 3 April 1964. This Hedeoma-
like plant is frequent on the campus of the University of Southwestern
Louisiana and in gardens and open woodlands in the Lafayette area.

SIDA 1 (4): 249—250. 1964.

250

It was determined by Dr. Lloyd H. Shinners; our material compares
favorably with specimens of C. gracile that we received on loan from
the Gray Herbarium. Dr. Shinners’ determination was verified (as
Calamintha gracilis Benth.) by Dr. A. Borissova of the Botanical Insti-
tute of the Academy of Sciences of the U.S.S.R., Leningrad. An illustra-

Calyx 3.0—3.5 mm. long; corolla light pinkish, with a red-purple
blotch at base of each of the 3 lower lobes, 3.5—4.0 mm. long; corolla
lobes directed forward.” This Asiatic species is quite inodorous.—John
W. Thieret, University of Southwestern Louisiana, Lafayette.

NEW COMBINATIONS IN TEXAS POLEMONIACEAE.—In the
preparation of a treatment of the family Polemoniaceae in Texas, nomen-
clatorial changes of three taxa have proved desirable. The category of
subspecies, though interpreted in diverse ways by different workers, and
even discarded by some, is deemed by the writer useful for taxa which
are moderately distinct in morphology and geography, yet not suffi-
ciently so to justify species segregation. On this basis the following
combinations are proposed:

GILIA RIGIDULA Benth. subsp. acerosa (Gray) Wherry, stat. nov.
Gilia rigidula var, acerosa Gray, Proc. Amer. Acad. 8: 280, 1870, bas-
lonym,; (G. acerosa (Gray) Britt., Man. Bot. NE. St. 761, 1901; Giliastrwm
acerosum (Gray) Rydb., Fl. Rocky Mts. 699, 1917.) This taxon inter-
grades too freely with the species-type to accept the Britton-Rydberg
view of species independence, but in northern Texas and adjacent states
the reduction of its leaf-segments to subacerose filiform outline does
become consistently extreme.

X DRUMMONDII Hook, subsp. johnstonii (Wherry) Wherry,
Stat. nov. Phlox johnstonii Wherry, Wrightia 2: 198, 1961, basionym.
While the copious glandularity and elongate corolla-tube distinguish
this taxon from all the other annual Phloxes, and it is endemic in a
small area in northwest Texas—having been recognized in two counties
adjacent to Kent since its original discovery—it seems after all insuffi-
ciently differentiated to merit species independence, so reduction in
status is here proposed.
DRUMMONDII Hook. subsp. tharpii (Whiteh.) Wherry,

comb. nov. Phlox tharpit Whitehouse, Amer. Midl. Nat. 34: 399, 1945,
ees (Phlox glabriflora (Brand) Whiteh., subsp. tharpii (Whiteh.)
Wherry, Gen. Phlox 62, 1955: Phlox drummondii Hook. subsp, drum-
mondit var. tharptti (Whiteh.) Erbe, Amer. Midl. Nat. 67: 280, 1962).
Recent study of some hundreds of herbarium sheets of annual Phloxes
in several herbaria has led the writer to ange his view as to the
relationship of this taxon, and instead of ee Whitehouse in group-
ing it close to taxon glabriflora, the Erbe and Turner plan of placing

SIDA 1 (4): 250. 1964,

201

it under taxon drummondii is favored. At the same time, it is deemed
to deserve the higher status previously assigned, so is herewith pub-
lished in the corresponding new combination.—Edgar T. Wherry, Univer-
sity of Pennsylvania, Philadelphia.

SCUTELLARIA THIERETII (LABIATAE), A NEW SPECIES FROM
COASTAL LOUISIANA.—Among some Louisiana collections sent by
Dr. John W. Thieret for determination was a Scutellaria evidently allied
to S. Drummondii Bentham, a very common and variable species in
Texas (though not in the easternmost counties), extending into Okla-
homa, New Mexico, and northern Mexico. Additional material supphed
by Dr. Thieret showed that the Louisiana plant also was rather variable
despite its restriction to a very small geographic area. Although th
variations make it extremely difficult to find usable key differences,
individuals of similar size and age of the Louisiana plant and of S. Drum-
mondii could easily be distinguished. The most tangible differences were
in the smaller and rather long-petioled lower leaves of S. Drummondti
(unfortunately not present except early in the season), and the marked
reduction of the uppermost ones to floral bracts much shorter than the
flowers (not plainly evident until fairly late in the season). There was
no difference in nutlets, such as distinguishes S. muriculata Epling.
The Louisiana plant is considered to be one more in a group of very
closely related species, and in honor of an energetic and productive
collector it is named

LLARIA Thieretii Shinners, sp. nov. Annua ex affinitate S.
Drummondii, differt caule crassiore, foliis inferioribus majoribus sed
brevipetiolatis, foliis superioribus minus reductis, supremis flores ex-

Os.

cedentibus vel eis paulum brevioribus. HOLOTYPE: roadside, in shell
sand, Pecan Island, Vermilion Parish, Louisiana, J Thieret 16162,
18 July 1963 (SMU; isotype, USL). P apg a all ui LOUISIANA
CAMERON PARISH: grazed meadow-li ar ighway at

of Ww

Grand Chenier, Thieret 8774, 6 July 1962 ne aa on PARISH:
few plants on shell ridge in brackish marsh, vic. of U.S.L. Biology Lab,
Redfish Point, west side of Vermilion Bay, William D. Reese 2296, 29
July 1959 (USL); same locality, Reese 4167, 4187, 30 April 1961 (both
USL). Roadside, Pecan Island, Thieret 8682, 23 June 1962 (USL). Shell
ridges in vicinity of USL field station, south side of Redfish Point, west-
ern shore of Vermilion Bay, Dr. Norden’s Estuarine Biology Class, 15
July 1962 (SMU, USL).

Annual with a taproot. Stems solitary or much less commonly several,
simple or freely branched, 7—65 cm. tall, rather stouter than in S.
Drummondii of equivalent size, densely pubescent with mixed short
to medium long (0.2—1.0 mm.), glandless or partly inconspicuously
gland-tipped hairs, these either straight and spreading at right angles

SIDA 1 (4): 251—252. 1964.

202

or slightly retrorsely curved. Lower leaves (soon withering) relatively
short-petioled, the petioles 3—10 mm. long, shorter than the blades;
blades oblong-ovate to oblong-elliptic, 6—12 mm. wide by 12—25 mm
long, obtuse, shallowly crenate or subentire, the base widely tapered
or subtruncate but with wide V-attachment to the petiole, rather strongly
pinnately veined, rather densely pubescent on both surfaces, the hairs
on the upper surface erect, those on the lower suberect to low-spreading
or subappressed. Middle and upper leaves (floral bracts) progressively
shorter-petioled to sessile, with gradually reduced, narrower, more
entire blades, the uppermost prominent, extending to 3/4 the length of
the flowers or beyond. Flowers borne in upper 1/2—5/6 of plant. Calyx
and corolla densely pubescent outside with short, erect hairs. Calyx in
flower 2.5—4.0 mm. long. Corolla 7—14 mm. long; tube white, lmb
purple-blue, lower lip with purple-blotched white center.

Scutellaria Thieretii is the easternmost representative of the South-
western an exican Section Resinosae, separated by a small gap from
the main area of the group. It is also notable as an addition to the ex-
tremely small number of endemics in the Louisiana flora. I am indebted
to Dr. Thieret for supplying the two SMU collections, and for the loan
of mounted specimens from the University of Southwestern Louisiana.
—Lloyd H. Shinners.

MICRANTHEMUM GLOMERATUM (CHAPMAN) SHINNERS,
COMB. NOV. (SCROPHULARIACEAE).— Based on Micranthemum
Nuttallii var. ? glomeratum Chapman, Fl. S. U.S. ed. 2 (2nd _ issue)
Suppl. 2 p. 690. 1892. Hemianthus glomeratus (Chapman) Pennell, Proc.
Acad. Nat. Sci. Phila. 71: 248. 1920. The second issue of the second
edition of Chapman’s Flora is evidently very rare, there being no copy
even at the Library of Congress. It is of considerable importance since
the Second Supplement occupies pages 675—703 inclusive and contains
a number of new names as well as many new records. A copy was kindly
loaned by the Library of the University of Virginia, for which I am
very grateful—tLloyd H. Shinners.

TEXAS EVAX TRANSFERRED TO FILAGO (COMPOSITAE).—It

cluded species and source of the name (adopted by Linnaeus from
Loefling), Filago belongs to those species later segregated by Gaertner
under the name Evax (Josf Holub and Jindrich Chrtek, Zur Nomen-
klatur des Gattungsnames Filago L. 1753. TAKON 11: 195—201, 1962).
All the plants treated in my brief account of the Texas species of Evax
(Field & Lab, 19: 125—126, 1951) must have new names under Filago,
as follows.

FILAGO candida (T. & G.) Shinners, comb. nov. Calymmandra can-

SIDA I (4): 252—-253. 1964,

253

dida T. & G., Fl. N.A. 2: 262—263. 1842. Evax candida (T. & G.) Gray,
Syn. Fl. N.A. 2 pt. 1: 230. 1878.

AGO Nuttallii Shinners, nom. nov. Evax prolifera Nuttall ex DC.,
Prodr. 5: 459. 1836. Diaperia prolifera Nuttall, Trans. Amer. Philos. Soc.

esque, Atlantic Journal 1: 178, 1833 Gaewiae to Merrill, Index
Rafinesquianus; not seen); also Herbarium Rafinesquianum p. 24. 1833.
Evax multicaulis DC., Prodr. 5: 459. 1836. Diaperia ? multicaulis (DC.)
Nuttall, Trans. Amer. Philos. Soc. ns. 7: 338, 1840. Filago multicaulis
DC.) H

S. Texas) 1: 102. 1894. (Not Filago multicaulis Lamarck, 1778.) Filago
nivea Small, Bull. Torr. Bot. Club 24: 333. 1897. Evax nivea (Small)
936

FILAGO VERNA var. Drummondii (T. & G.) Shinners, comb.
Filaginopsis Drummondii T. & G., Fl. N.A. 2: 263—264. - cee
multicaulis var. Drummondii (T. & G.) Gray, Syn. Fl. N.A. 2 pt. 1: 229.
1878.—Lloyd H. Shinners, Southern Methodist University, ee 22,
Texas.

VERBESINA WALTERI SHINNERS, NOM. NOV. (COMPOSITAE) .—
Based on Athanasia paniculata Walter, Fl. Carol. p. 201. 1788, (Not Ver-
besina paniculata Poiret, 1808.) V. Coreopsis var. alba Michaux, FI.
Bor.-Am. 2: 135. 1803. “In Carolina maritima.” Actinomeris alba (Mich-
aux) T. & G., Fl N.A. 2: 335. 1842. (Not Verbesina alba L., 1753.)
Actinomeris paniculata (Walter) Small, Fl. S.E. U.S. pp. 1271, 1340.
1903. Ridan paniculatum (Walter) Small, Man, S.E. Fl. pp. 1445 (as
paniculata), 1 . 1933. I prefer to treat Actinomeris as part of the
diversified genus Verbesina, under which Walter’s plant requires a new
name.—Lloyd H. Shinners.

REVIEW

FLORA OF HENRY COUNTY, ILLINOIS, AN ANNOTATED LIST
OF THE VASCULAR PLANTS. Raymond J. Dobbs. 350 pp., 2 pl., map
(folded, inside back cover). Natural Land Institute, 303 Penfield Place,
Rockford, Illinois. 1963. $5.00

Henry County, Illinois, scarcely ranks as a botanical paradise. Located
in the northwestern part of the state (close to but not on the Mississippi
River, three counties south from the Wisconsin border), it is a typical
glaciated prairie county, largely under cultivation. The difference in
elevation between the lowest and highest points within its 826 square
miles is just 315 feet. The only major physical feature is the Rock
River, and this only for a short distance at the northwest corner, where
it forms the boundary for some 20 miles. The few woodlands have mostly
been cleared; marshes and bogs have been drained. The author tells us
that he worked for some years as a naturalist in California, including
a tour of duty in Yosemite National Park, and traveled extensively
throughout the United States and Canada to observe plants. One would
think that any leisure hours back in Geneseo would be spent looking
at pictures of more interesting places. Instead, for 22 years, Mr. Dobbs
made systematic and intensive botanical trips up and down his home
county, both by car and on foot. He says of his Flora, “It was made
solely as a scientific contribution, at the personal expense of the author,
and under no official auspices.” One would like to devise a special
medal to award those who do not demand inflated N.S.F. grants, a pack
of assistants, and the ballyhoo and flummery that have become a routine
part of American science, but who calmly, objectively, and conscien-
ticusly devote themselves to straightforward accomplishment of straight-
forward scientific tasks. Its recipients would be rare birds indeed.

Mr. Dobbs speaks in the introduction of his friendship with C. C.
Deam, and the first title in his list of references is Deam’s Flora of
Indiana, Perhaps the best compliment one can give the Flora of Henry
County is that Deam would certainly have approved. It immediately
invites comparison with Egbert W. Fell’s recent (1955) Flora of Winne-
bago County, Illinois, another excellent local flora which evidently
served as a model. Both are annotated catalogues without keys, re-
produced from typescript with non-justified margins, and bound in hard
covers. For the Dobbs book, the type has not been reduced in size, and
scientific names are in capital letters rather than lower case underlined;
the resulting page appearance is, I think, more pleasing. Mr. Dobbs gives
a rather full account of glacial history and soils, but in contrast with
the Winnebago Flora, says very little about vegetation types. But then
there is hardly a great deal to be said about natural vegetation in the

SIDA I (4): 254—256. 1964.

295

county described as “Hog Capital of the World.” There are 1,073 species
accepted as members of the flora (Winnebago County has 1,210). Notes
on medicinal plants and those poisonous to livestock have been supplied
from other publications. More valuable to the taxonomist are notes
on the distinctness of different varieties as represented in the area.
There are two full-page illustrations: a photo of Pedicularis canadensis
var. Dobbsii Fernald as frontispiece, and on page 1 a drawing of
Rubus Dobbsii L. H. Bailey, here newly described from manuscript
supplied by Dr. Bailey. Nomenclature mainly follows that of Fernald’s
8th edition of Gray’s Manual; the author makes numerous acknowledg-
ments of assistance with determinations given by Fernald. Some collec-
tions have been deposited at the Gray Herbarium; the principal set is
at the State Natural History Survey in Urbana.

In reviewing Jones and Fuller’s Vascular Plants of Illinois several
years ago (S.W. Nat. 1: 4447, 1956), I commented on the impressive
progress the state has made toward the kind of complete coverage which
exists for the British Isles. It is of interest to make a few comparisons
between the Flora of Henry County and a recent English county flora,
the privately published A Flora of Nottinghamshire, by R. C. L. Howitt
and B. M. Howitt (1963). Nearly identical in size with Henry County
(844 square miles), Nottinghamshire is described by the authors as
almost “the average English county. ... There is nothing spectacular,
no mountains or torrents, no seaboards, no natural lakes.” The most con-
spicuous difference between it and the Illinois county is the presence
of a large industrial city and of coal mines e most conspicuous
botanical difference is in the original eestor: which was forest. In

fame, still exist. There are 79 persons listed as recorders who have con-
tributed information about the flora (in addition to the two authors),
over a period of more than 300 years, starting about 1650—different
indeed from the strictly modern, one-man performance of Mr. Do

The Nottinghamshire Flora suggests some comforting things about the
American scene. Two very familiar cliches applied to European settle-
ment in America are “taming the wilderness” and “the raw frontier.”
The two reveal a subconscious awareness that crude destruction is not
exactly the same as refined civilization. The land is to be lived in rather
than off, and familiarly known and loved. In Winnebago County
there are forest preserves, and the Flora was sponsored by The Nature
Conservancy—hopeful omens for the future. Henry County of course
cannot boast any remnants of a Sherwood Forest, but enough survives
to have kept an energetic naturalist occupied for 22 years. Now we
have something that records what is there, and in so doing makes us
more aware of it. We can hope that those who live in Henry County

256

will acquire greater appreciation of the land and the plants around
them

Mr. Dobbs has given us a book that is not only a solid contribution
to science, but a significant step in cultural progress as well—Lloyd H.
Shinners.

S D CONTRIBUTIONS
| A TO BOTANY

VOLUME 1 NUMBER 5 JUNE 1964

CONTENTS

How to study the Florida flora.

Lloyd H. Shinners. 257
Notes on the vegetation of the Mexican state of Morelos.
Chester M. Rowell. 262
The identity of Sagittaria isoetiformis (Alismataceae).
R. K. Godfrey and Preston Adams. 269
Meiotic chromosomes in African Commelinaceae.
274

Walter H. Lewis.

NOTES. More additions to the Louisiana flora. 294.—Tigridia purpurea (Herbert) Shinners,
comb. nov. (Iridaceae). 295.—Machaeranthera pinnatifida (Hooker) Shinners, comb. nov.

(Compositae). 295.

REVIEW. Roadside Flowers of Texas, by Wills and Irwin. 296.

SIDA is privately published by Lloyd H. Shinners, SMU Box 473,
Dallas, Texas 75222, U.S.A. Subscription price $6 (U.S.) per volume
of about 360—400 pages, parts issued at irregular intervals.

©)
SIDA Contributions to Botany volume 1 number 5 pages 257—298
copyright 1964
by Lloyd H. Shinners

HOW TO STUDY THE FLORIDA FLORA’
LLOYD H. SHINNERS
Herbarium, Southern Methodist University, Dallas, Texas 75222

The British Isles are the most thoroughly botanized area in the world,
and may well be taken as an ideal model. The new edition of Clapham,
Tutin and Warburg’s Flora of the British Isles (1962) is the culmination
of nearly 450 years of activity by British botanists. In addition to the
usual keys, recent synonymy, and full descriptions, the book contains a
wealth of geographical and biological information, including chromo-
some numbers and pollinating agents. Add to this fine flora the com-
panion series of Illustrations (three parts so far published), H. Godwin’s
The History of the British Flora (1956), and the Atlas of the British
Flora by F. H. Perring and S. M. Walters (1962), and we are indeed
well supplied with information about the British flora.

Although the combined area of the British Isles is a little more than
twice that of Florida, their flora is much smaller, owing to the far
northern latitude and history of total glaciation. For purposes of com-
parison we may take the total of 1,511 “good” (or Linnaean) native species
given in J. E. Dandy’s List of British Vascular Plants (1958). We have
no accurate information as to the total in Florida except that it is
very large, including on the one hand members of the cool-temperate
Appalachian flora and tropical species on the other, together with one of
the most notable concentrations of endemics in the continental United
States. Considering all this floral wealth, and the rate at which new
native species are still being found, I think it reasonable to estimate
hat the total native flora will run close to three times that of the
British Isles, or not far from the totals known for Texas and for
California.

Botanical knowledge cannot be accumulated without botanists, whether

well as manpower, and even though we deduct from the indicated 1,350
years the past 180 (stretching things some, for many of those years saw
little or no progress at all with work on Florida plants), we are still
left with more than a millennium of time as our second need. Floras are
always tacitly assumed to refer to living plants, but as a practical work-
ing matter, they are primarily based on herbarium specimens. With these
Britain is abundantly supplied: Kew reports 6 million, the British
Museum has released no figures but is also huge, and Edinburgh claims

‘All notes are at end - article,
964.

SIDA 1 (5): 257—261.

258

2 million. A combined total of more than 10 million would be a low
estimate for the three principal institutions alone. A majority of these,
to be sure, came from outside the British Isles. But it was necessary to
have a world sample in order to delimit and organize the orders and
families, and to establish the correct identity of species, while the
constant influx of aliens means an ever-growing need for large collec-
tions of exotics. Since the organization of the world flora into orders
and families has been reasonably well done for us, we may need much
less than the indicated 30 million for the preparation of our ideal Florida
Flora. But we still require vastly more than the combined total of less
than 1/3 of a million currently held by the three principal herbariums
in the state.

My calculations may seem unrealistic, but the facts behind them are
worth citing to emphasize how very much needs to be done. We do not
have even a simple list, still less an adequate manual for identification,
while an encyclopedic reference work with everything down to the latest
chemocytophylesis is hardly to be thought of at this stage. Where to
begin? We may turn again to the British Isles for illumination.”

In the 1520’s William Turner, a young medical student at Cambridge,
undertook to describe wild herbs from direct observation, rather than
accepting the authority of the Continental herbalists. Thus began the
scientific study of the British flora. A century later, in the summer of
1629, a group of men led by Thomas Johnson made excursions from Lon-
don to observe wild plants. Though most of them had some professional
connection with medical practice, their trips had no specifically utilitar-
lan purpose, but were made rather for the satisfaction of idle curiosity.
Botany as a pure science had been born in Britain. By 1670 John Ray,
Puritan divine and gifted amateur naturalist, was able to publish his
See Plantarum Angliae et Insularum Adjacentium. The posthumous

ird edition of its successor, Synopsis Methodica Stirpium Britanni-
carum, in 1724, represents the high-water mark of pre-Linnaean botany.
In 1762 William Hudson’s Flora Anglica introduced Linnaean classifica-
tion and binomial nomenclature. These were made availa the
English language in 1776 in A Botanical Arrangement of All the Vegeta-
bles Growing Naturally in Britain, by William Withering.? James
Sowerby’s English eile in 386 volumes (1790—1814; text by J. E.
Smith, whose name does not appear in the earliest volumes), provided
the first completely eee an In 1829 John Lindley’s A Synopsis
of the British Flora established the Natural System. For over a century
afterward British botanists were to have available to them at all times
a choice of several current floras, prepared by such men as the two
Hookers, Bentham, and Babington, running through as many as eleven
editions. With the publication of H. C. Watson’s Outline of the Geo-
graphical Distribution of British Plants in 1832, particular attention
was focussed on local studies, largely carried out by amateurs. In 1873

299

Watson’s work was to appear in expanded form as his famous Typo-
graphical Botany, which in 1932 was replaced by G. C. Druce’s Comital
Flora of the British Isles. In 1928 E. J. Salisbury proposed a biological
flora of Britain, but publication did not begin until 1941, as a series of
papers in the Journal of Ecology, contributed by numerous authors.
This is still in continuation and far from completion. The epoch-making
first edition of Clapham, Tutin and Warburg’s Flora of the British Isles
in 1952 provided not only an up-to-date manual in the established tra-
dition, but (so far as the limits of a compact, 1-volume work allowed)
a biological flora as well, with details of distribution, habitat, life form,
abundance, pollination, and chromosome number. An
abridged companion work, Excursion Flora of the British Isles, was
published in 1959.

Most noteworthy is the fact that British botany begins with direct
field study, and this remains a major feature of it down to the present.
Not merely collecting specimens, but continuing observation, checking
what is stated in books and filling in gaps in knowledge. This broad and
solid factual foundation is something greatly to be envied—and a lesson
to be taken to heart. Also enviable is the gradual and easy progress
from simple to very complex and detailed investigation. In this respect
we are not fortunate: we are confronted with the entire gamut of methods
from the most elementary to the most advanced all at once. But there
is something of a lesson for us here too. The best advanced work must
have a solid basis on which to rest; until we have that basis, we must
concentrate on producing it. So-called “alpha taxonomy” should come
first; more specialized studies can follow. (I do not use the term “bio-
systematics” for the latter, since it by no means includes all the possi-
bilities; furthermore it is a term offensively misused by too many
present-day American botanists who imagine that only those who use
cytology or cytogenetics really know the biology of plants. The ‘‘old-
fashioned” taxonomist employing data from ecology, geography, and
phenology along with phenotype variations is in fact taking into account
a broader range of biological information than many biosystematists do.)
Historically suggestive also is the way in which one thing leads to

; r example, the floras of Hudson and Withering provided easy
tools for local botanists to study the flora. If one looks over the lst of
British county and local floras, it is immediately clear that they do not
begin to be produced in numbers until these pioneer floras have opened
the door, and they later appear in flood proportions when a variety of
manuals is available, with the added stimulus of H. C. Watson’s and
Druce’s attention to geographical distribution. To be sure, economic and
social factors were also involved—industrialization, rise in population,
increasing wealth, greater ease of travel—but the existence of utility
manuals was certainly a key factor in the great increase in local studies,
which in turn made possible better general floras, along with promoting

260

an atmosphere favorable to giving both public and private support to
professional botanists.

Perhaps at this point we ought to say something about what a flora
is, or ought to be. It is first, and above all, a record of facts. What plant
grows where, and when, is information wanted and need by agricul-
turists, hort cultures field zoologists, conservationists, nature-lovers,
geologists, biochemists, and a host of others. Floras do not exist merely
for the attention of professional botanists. Their purpose is not to provide
material for the diversion of phylogenists, or to gratify the disdain of
biosystematists. It is a basic reference which serves a great variety of
needs.

Writing a flora is a tedious and laborious job. We have seen what a
stupendous amount of background activity precedes the writing of a
really good one. Taxonomists are prone to seek relief from the strain,
A recent illustration is C. W.
James’s paper, “Endemism in Florida” (Brittonia 13: 225—244, 1961).
On the basis of selected, very incomplete evidence, the author seeks
to explain Florida endemics and relics' as immigrants from Appalachia.
This was Fernald’s naive theory of the origin of the Coastal Plain flora.
Anyone who has studied the Texas flora realizes very quickly that such
an explanation is quite inapplicable in the Gulf Southwest, and Florida
and Texas have too much in common to suppose that an explanation un-
workable for the one can be accepted for the other.“ One can only
regret that the time spent by Dr. James in developing ill-founded
speculations was not devoted instead to developing more foundations.
His strictly taxonomic papers show that he is well able to do this. He
has been too influenced in his thinking by the eastern Manual Ranges
and the limited outlook of their dominating authors.

We may conclude that the best way to study the Florida flora is to
begin by being a Texas botanist.

NOTES
' Based on a talk given to the Life Sciences Seminar of the University of South Florida,

bravely invited me to epee and heroically refrained from imposing any restrictions.
wish also to express my appreciation to Dr. Vatgs O. as who made a very spirited
and able defense of Dr. James. It was a pleasure to have lively audience participa tion
? There are very readable summaries of British botanical pee in British Botanists, by
John Gilmour (1946), a in Wild Flowers: Botanising in Britain, ps John Gilmour and
W yi

Max Walters (new edition, 1955; see especially Cas 2, “Ho Yur Flora Was Dis-
covered”’). See also Nicholas Polunin’s “British Floras Ancient ae lies Rhodora 55:
209—224, 1953

>To whom, it is said, this punning tribute was paid while he lay dying: ‘The Flower
of Feat ea! is Withering.

4A in good ae means a widow; in Clementsian ecological gobbledygook it is
a later oe for

°Tt has been nee ‘ak isolated colonies of Coastal Plain species within the Appa-
lachian highland represent ancestral stocks. I think a more likely explanation is just the
reverse See are late emigrants from the Coastal Plain. We know from geological evidence
that he close of the Pleistocene was a period of vast flooding and erosion, with the develop-

261

ment of flood plains and evilge aie the glaciated area. It seems to me highly probable

that the Dea coe wee tal Plain species were rapid invaders, spreading far out
of the Coastal Plai self cc. hand se Coastal Plain species about the western Great
Lakes are al on, illustrations), t replaced more gradually in succession by the
Appalachian flora. The endemic Co serine, capes to a few spots on eroding

ore ancient occurrence o

> m

stream banks in Kentucky and Tennessee, may represent

the same thing. It is one of five shrubby species comprising hee genus, all the rest of which
are confined to the outer Coastal Plain.

_ have been shore ni and hence coastal plains,
nd land have been differentiated. I see no reason to
plants ever exist anyw ae else. In Te e
genera Vaseyochloa (Gramineae) and ee. (Compositae) mus
they occur on the very yo — oan, and there is nothing to relate them to the
Appalachian or Ozarkian or Sierra Madrean or any other upland flora. There are exactly
comparable aio in Florida, as well as a striking disjuncts as Bonamia villosa, oa
ring both states. I cons res the so-called Orange Island hypothesis, espoused by Wood-

son, not ae possible and logical, but necessary

however small, for as long as
e that typic

—
=
»
fon

NOTES ON THE VEGETATION
OF THE MEXICAN STATE OF MORELOS
CHESTER M. ROWELL
Department of Biology, Texas Technological College, Lubbock'

During the summers of 1949 and 1950 the author and his students were
privileged to make extensive observations and collections in the Mexican
State of Morelos. The work was carried on as part of the summer field
courses under the joint sponsorship of the departments of Wildlife
Management and Biology of the Agricultural and Mechanical College of
Texas. Some additional collections were made by the author in August
of 1960.

The state of Morelos is located south of the Federal District in central
Mexico. It is bounded on the north by the state of Mexico, and the
Federal District, on the east by Pueblo, by Oaxaca on the south and
Guerrero on the west. There is a variation of altitude within the state
from approximately 17,800 ft. in the northeast to approximately 2,700
ft. in the southwest. The state boundary extends from the peak of the
volcano Popocatepet] to the edge of the Balsas Basin.

The vegetation zones of Mexico as presented by Leopold (1959), and
his terminology will be followed in general in this paper.

The flora of the state is divisible into two major types, temperate and
tropical. Four of Leopold’s vegetation types are found within the political
limits of Morelos.

h emperate zone includes vegetational types: Pine-Oak Forest,
Boreal Forest, and pine-Meadow. These vegetational types are found
in the northern one-third of the state.

The Mixed Forest type or Pine-Oak Forest is characterized by open
scattered stands of woodlands dominated by pines or oaks. Apparently
dependent on the elevation and available moisture, either the pines or
the oaks may assume dominance in any particular area and occasionally
they are distributed almost equally. It is interesting to note that no
distinct zone of oak-shrub was observed by the author in Morelos where
this type merged with some of the more xeric types at lower elevations
of approximately 9,000 and 6,500 ft.

The dominant members of the flora of the Pine-Oak Forest are: Pinus
montezumae, Pinus lawsonti, Pinus teocote, Quercus spp. Other important
members of the flora included in this type are: Arbutus glandulosa,
Ceanothus azureus, Buddleia americana.

The Boreal Forest type is found in rather limited areas in the north
portion of the state. It is characterized by coniferous forests with a

1 - . . - x
Current temporary address: Department of Botany, Oklahoma State University, Still-

water.
SIDA 1 (5): 262—-268. 1964.

263

bunch grass or sacaton (Festuca tolucensis) ground cover. A humid
environment is associated with this vegetational type and limits it in
Morelos between the elevations of approximately 9,000 and 13,000 ft.
Canyons at lower elevations which are subject to frequent fog condi-
tions also exhibit this type of vegetation.

The more important species of this vegetational type include: Pinus
lawsonti, Pinus ayacahuite, Pinus leiophylla, Pinus teocote, Pinus monte-
zumae, Pinus hartwegii, Abies religiosa, Festuca tolucensis.

The third temperate vegetational type is the Alpine Meadow. To my
knowldge it is found in Morelos only in the small area where the state
line extends up the side of the volcano Popocatepetl. In this area, which
is normally above timberline, there are extensive meadows of sacaton
(Festuca spp.) with robust herbs such as Lupinus and Castilleja.

The tropical vegetation type found in the state is the Tropical Deci-
duous Forest with some variations. This type covers approximately the
southern two-thirds of the state and is usually found below elevations
of 6,500 ft.

This Tropical Deciduous Forest as observed in Morelos exhibits two
rather distinct forms. One form is typical of that outlined by Leopold
and the other, a more arid type, seems related to his Arid Tropical
Shrub.

The typical Tropical Deciduous Forest type is found on the lower
mountain slopes from approximately 6,500 to 4,000 ft. and in canyons
at lower altitudes that have higher available moisture.

This type is predominantly composed of low, shrub-like trees and
some of the larger cacti. The trees are usually leafless during the dry
season, but are more abundant and larger than those of the more arid
type.

The more important members of this type include the following:
Ipomea arborea, Juliana adstrungens, Bursera spp., Pseudosmodingium
spp., Comocladia spp., Parchycereus marginata, Cephalocereus spp.

The more arid form of the Tropical Deciduous Forest is usually found
at elevations below 4,000 ft. It is characterized by sparse, thorny, low
shrubs and cacti. The more important members of this type include the
following: Acacia farnesiana Acacia spp., Pithecolobium spp., Opuntia
spp., Cephalocereus spp., Crescentia spp., Dodonaea viscosa, Lippia spp.,
Lantana spp.

In the riparian associations along stream beds and in marsh lands the
vegetation exhibits marked variations and includes such forms. as:
Persea americana, Taxodium distichum, Ficus spp., Astianthus viminalis,
Guazuma ulmifolia.

Due to the extremes in ecological conditions as expressed through both
altitudinal and moisture variations, Morelos exhibits remarkable varia-
tion in vegetation and as such presents a very interesting problem in
floristics.

264

The following list of species is based on specimens collected in
Morelos by the author and his students. Most of the determinations were
made by the author using the facilities at the University of Michigan
Herbarium. All species of the Verbenaceae were verified or determined
by Dr. H. N. Moldenke. Dr. Rogers McVaugh verified the author’s
determinations of many species in the Leguminosae. It is hoped that
this lst will contribute to the sparse knowledge of the distribution of
the Mexican flora.

For simplicity the species are arranged alphabetically by families,
genera, and species. After each species the town nearest the collection
site is listed plus a number in parenthesis indicating vegetation type.
1. Alpine Meadow; 2. Boreal Forest; 3. Pine-Oak Forest; 4. Tropical
Deciduous Forest. Specimens on which these determinations were made
are deposited at one or more of the following herbaria: S. M. Tracy
Herbarium, Texas A. and M. College; The Herbarium, the University of
Michigan; The Herbarium, Southern Methodist University.

ALISMATACEAE
Sagittaria lancifolia L. Cuautla (4)
AMARYLLIDACEAE
Allium glandulosum Link and Otto Cuautla (4)
Nothoscordum fragrans (Vent.) Kunth Tres Cumbres (2)
Pancratium littorale Jacq. Axochiapan, Cuernavaca (1)
APOCYNACEAE
Haplophytum cimicidum A. DC. Yautepec (1)
ASCLEPIADACEAE
Asclepias grandiflora Fourn. Yautepec (1)
BIGNONIACEAE
Astianthus viminalis (H.B.K.) Baill. Amacusac (1)
BOMBACACEAE
Waltheria americana L. Cuautla (1)
CARYOPHYLLACEAE
Arenaria decussata Willd. Lagunas de Zempoala (2)
COMMELINACEAE
Commelina coelestis Willd. Cuautla, Tlacotepec, Tres
Cumbres, L. de Zempoala
(2, 3)
Tradescantia commelinoides R. & S. Lagunas de Zempoala (2)
COMPOSITAE
Achillea millefolium L. Lagunas de Zempoala (2)
Ageratum corymbosum Zucc. Yautepec, Cuautla, Axochia-
pan (4)
Aphanostephus pachyrrhizus Shinners Cuautla (4)
Aster Lima Lindl. Axochiapan (4)

Conyza filaginoides (DC.) Hieron. Cuautla (4)

265

Conyza sophiaefolia H.B.K. Cuautla (4)

Dyssodia pinnata (Cav.) Robinson Cuautla (4)

Erigeron maximus Link & Otto Lagunas de Zempoala (2)

Erigeron scaposus DC. Huitzilac, Cuautla, Tres

umbres (2,

Florestina pedata (Cav.) Cass, Cuautla, Yautepec (4)

Florestina trifida DC. Cuautla

Florestina tripteris DC. Cuautla, Yautepec,
Axochiapan (4)

Galeana hastata Llave & Lex. Tiacotepec (4)

Galinsoga aristulata Bicknell Progreso (4)

Heterotheca inuloides Cass. Cuautla (4)

Lagascea rubra H.B.K. Cuautla a

Melampodium oblongifolium DC. Axochiapan, Yautepec (4)

Melampodium paludosum H.B.K. oe Cuautla (4)

Otopappus robustus Hemsley Axochiapan (4)

Pectis latisquama Schultz Cuautla (4)

Pinaropappus roseus Less. Cuautla (4)

Sanvitalia procumbens Lam. Axochiapan, Tlacotepec,

Cuautla (4), Yautepec (4)
Sclerocarpus divaricatus (Bentham)
Yautepec, Axochiapan (4)

Sclerocarpus uniserialis B. & H. Axochiapan (4)
Spilanthes americana var. parvula
(Rob.) A. H. Moore Axochiapan (4)

Stevia serrata Cav. Cuautla (4)

Tagetes filifolia Lag. Cuautla (4)

Tagetes jaliscana Greenm. Cuautla (4)

Tagetes lucida Cav. Cuautla (4)

Tridax coronopifolia Hemsley Cuautla Yautepec (4)

Tridax procumbens L. Axochiapan (4)

Zexmenia aurea B. & H. Tres Cumbres (2)

Zexmenia crocea Gray Yautepec (4)

Zexmenia helianthoides (DC.) Gray Yautepec (4)

Zinnia multiflora L. Yautepec, Axochiapan,

Cuautepec (4)

CRUCIFERAE

Eruca sativa Mill. Cuautla (3, 4)
CYPERACEAE

Eleocharis nodulosa (Roth) Schultes Cuautla (3)
ERICACEAE

Vaccinium leucanthum Schlecht. Lagunas de Zempoala (2)
HYPERICACEAE

Hypericum pauciflorum H. B. K. Cuautla (3)

266

LEGUMINOSAE

Acacia angustissima (Muill.) Kuntze

Acacia farnesiana (L.) Willd.

Acacia paniculata Willd.

Aeschynomene virginica (L.) B.S. P.

Astragalus strigulosus H. B. K.

Brongniartia podalyrioides H. B. K.

Calliandra grandiflora (L’Her.)
Bentham

Calliandra houstoniana (Mil.) Standley

Calliandra penduliflora Rose

Cassia occidentalis L.

Cassia uniflora Mill.

Cologania procumbens Kunth

Crotalaria mollicula H. B. K.

Crotalaria pumila Ortega

Crotalaria vitellina Ker

Eysenhardtia polystachya (Ortega)

Sarg.
Lupinus elegans H. B. K.
Mimosa albida H. & B.
Mimosa benthami Macbride
Mimosa caerulea Rose
Nissolia fruticosa Jacq.
Nissolia hirsuta DC.
Pachyrrhizus erosus (L.) Urban
Phaseolus atropurpureus Bentham
Phaseolus coccineus L.
Phaseolus heterophyllus Willd.
Pisum sativum L.
Rhynchosia pyramidalis (Lam.) Urban
Tephrosis nicaraguensis Oerst.
Trifolium amabile H. B. K.
Zornia diphylla (L.) Pers.

LILIACEAE

Anthericum aurantiacum J. G. Baker
Bessera elegans Schult.
Milla biflora Cav.

LOASACEAE

Mentzelia aspera L.

MALPIGHIACEAE

Bunchosia palmeri 5. Watson

Yautepec (4)
Axochiapan (4)
Axochiapan (4)

Yautepec (4)

Cuautla (4)

Axochiapan, Cuautla (4)
Axochiapan, Yautepec (4)
Axochiapan (4)
Yautepec (4)

Cuautla, Progresso (4)
Cuautla, Axochiapan (4)

Huitzilac, Cuautla (3)
Lagunas de Zempoala (2)
Yautepec (4)

Yautepec, Axochiapan (4)
Yautepec (4)
Cuautla (4)

Cuautla, Tres Cumbres (2,3)
Cuautla (3)

Yautepec (3, 4)
Yautepec (4)
Cuautla (4)

Yautepec, Cuautla (4

Yautepec, Axochiapan,
Cuautla (4)

MALVACEAE
Anoda cristata (L.) Schl.

Anoda hastata Cav.

Malvastrum coromandelianum (L.)
Garcke

Sida procumbens Sw.

Dorstenia drakena L.
ONAGRACEAE
zia mexicana Jacq.
POLYPODIACEAE
Adiantum concinnum H. & B.

Adiantum kaulfussi Kunze

Adiantum poireti Wikstr.

Bommeria pedata (Swartz) Fournier

Cheilanthes angustifolia H. B. K

Cheilanthes cucullans Fee

Cheiloplecton rigidum (Swartz) Fee

Notholaena aurea (Poir) Desv.

Pellaea skinneri Hooker

Polypodium polypodioides (L.) A. S.

Hitchcock var. aciculare Weatherby

PONTEDERIACEAE

Heteranthera limosa (Sw.) Willd.
RANUNCULACEAE

Clematis drummondii T. & G.
RHAMNACEAE

Karwinskia umbellata (Cav.) Schlecht.

RUBIACEAE
Bouvardia ternifolia (Cav.) Schlecht.
Cephalanthus salicifolia H. & B.
Diodia tetracocca Hemsley
Galium asperrimum Gray
Paederia pringlei Greenman
Spermacoce haenkeana Hemsley
Spermacoce patula M. & G

SAPINDACEAE
Cardiospermum halicacabum L.
Dodonaea viscosa Jacq.

Cuautla, Axochiapan,
Yautepec (4)
Progresso (4)

Yautepec, Axochiapan (4)
Yautepec (4

Yautepec (3, 4)
Yautepec (4)
Cuautla (4)

Cuautla, Yautepec,
Axochiapan (4)

Axochiapan (4)
Cuautla (3)
Axochiapan (4)
Cuautla (3)
Amacusac (4)

Cuautla (3)

Yautepec (topotype) (4)
Cuautla (3)

Cuautla, Axochiapan,
Huitzilae (3, 4

Axochiapan (4)
Yautepec (4)

268

SCROPHULARIACEAE

Castilleja pringlei Fern. Lagunas de Zempoala
Tres Cumbres (2)
Castilleja scorzoneraefolia H. B. K. Cuautla, Lagunas de
Zempoala (2, 3)
Castilleja tenuiflora Bentham Cuautla, Huitzilac (3)
Mimulus glabratus (L.) Wettst. Lagunas de Zempoala (2)
Pedicularis mexicana Zu Lagunas de Zempoala (2)
Penstemon campanulatus Willa, Yautepec, Cuautla, Tres
Cumbres (3, 4)
Veronica americana (Raf.) Schwein. Lagunas de Zempoala (2, 3)
SELAGINEL
Selaginella pallescens (Pres.) Spring Yautepec (4)
Nicotiana glauca Graham Cuautla (3, 4)
Solanum bicolor Willd. Cuautla (3)
num nigrum L. Lagunas de Zempoala (2)
STERCULIACEAE
Ayenia montana Rose Yautepec (4)
Melochia pyramidata L. Cuautla (3, 4)
Physodium dubium Hemsley Yautepec, Axochiapan (3, 4)
TURNERACEAE
Turnera ulmifolia L. Cuautla (4)
VERBANACEAE
Bouchea prismatica (L.) Ktze. Axochiapan (3)
Lantana achyranthifolia Desf. Yautepec, Axochiapan,
Cuautla (3, 4)
Lantana camara L. Cuautla, Yautepec,
Axochiapan (3, 4)
Lantana hispida H. B. K. Yautepec (3, 4)
Lantana velutina H. B. K. Cuautla, Axochiapan,

Tlacotepec (3, 4)

Lippia berlandieri Schauer Yautepec (4)
Vitex mollis H. B. K. Cuautla (4)
VITACEAE
Cissus subtruncata Rose Yautepec (4)
REFERENCES
1. LEOPOLD, A. er R. 1950, Vegetation Zones of Mexico. Ecology 31:4.
2, ———_——__ 1959. Wildlife of Nexen: University of California ben Berkeley.

3. MART Z, is 1937, Catalogo de Nombres Vulgares y Cientificos de Plantas Mexi-
canas. Mexicc
: 1950. Personal Communicatio
rr Los Pinos Mexicanos. Mex XICO
i VIVO, J. A. iene eae de Mexico. Fondo de tee Economica. Mexico, D. F.

THE IDENTITY OF SAGITTARIA ISOETIFORMIS
(ALISMATACEAE)'
R. K. GODFREY AND PRESTON ADAMS

Department of Biological Sciences. Florida State University, Tallahassee,
and Department of Botany, Depauw University, Greencastle, Indiana

Shortly after publishing a revision of the North American species of

named S. isoetiformis (1859b), were said to be common along sandy
lake margins where they formed extensive patches in the shallow water,
their slender interlacing stolons bearing tufts of leaves and rooting at
the nodes. The type specimen (Fla., Lake Co., Nash, March 22, 1894, MO)
possesses slender, attentuate to only slightly dilated phyllodia about 1-2

wide—a distinctive feature which apparently suggested to Smith
the leaves of Isoetes (hence the specific epithet). Smith (1895b) con-
sidered S. isoetiformis to be more closely related to S. graminea Michx.
than to any other species.

The identity and specific distinctness of Sagittaria isoetiformis long
went unquestioned. Recently, however, Bogin (1955) interpreted this
taxon merely as an ecological variant of S. graminea var. graminea, the
variant said to occur in lakes having a marked seasonal drop in water
level. Beal (1960), in treating the Alismataceae of the Carolinas, made
no reference to S. isoetiformis, but he distinguished material from the
Carolinas as S. teres S. Watson and considered it specifically distinct from
S. graminea. Bogin (op. cit.) had considered S. teres as a variety of S.
graminea with a range from Cape Cod to southern New Jersey. We
identify the Carolina plants called S. teres by Beal with S. isoetiformis,
the latter ranging from southeastern North Carolina to peninsular
Florida and southern Alabama, and consider S. teres distinct from both
S. isoetiformis and S. graminea var. graminea.

Fernald (1950) says in part of Sagittaria teres: :
all represented by terete, attenuate, often nodose ‘arfliodic: those of
terrestrial plants slender and elongate (up to 6 dm long), those of deep
water shorter, very thick, spongy and digit-like;... achenes... with
strongly rounded crenate dorsal keel, the faces (when fully ripe) rugose
and irregularly 2-4 (or more) keele

The terete, attenuate, nodose chviloais characteristic of the New
England Sagittaria teres specimens are unlike the phyllodia of material
from the Carolinas southward. Note particularly Fernald’s statement
that those of S. teres are erect, slender and elongate if the plants are

“ce

leaves erect,

! This gal aes was sp ean (in part) by a research grant, GM-06305, to the senior
author from the Division of General Medical oh caoed "Public Health eae
SIDA 1 ne 269—273. 1964.

270

I
i
I}
|
| 7 |
DP | / |
VW a /
fo iJ i
; f | ¥ | |
| ff Lf of
| ff
| /
ee F
| fy
| i
4 FE POS
ei LIA

i

€

a-c: Sagittaria teres. a. Habit, deeper water form. b. Habit, on
Habit, form with

Fig. 1.
shore form. c. Achene. d-e: Sagittaria graminea. d.
submersed winter rosette and emersed leaves. e. Achenes.

271

terrestrial (Fig. 1b), shorter, very thick, spongy and digit-like in deep
water (Fig. la). In lakes and ponds of the Southeast, terrestrial (on
shore) plants of S. isoetiformis have short (0.5-1.0 dm) phyllodia,
flattened dorsally, some, at least, of the phyllodial tips slightly dilated
and laminar (Fig. 2b). In water the phyllodes are lax and very much
longer (to at least 4-5 dm), flattened and strap-like, and with gradually
attenuate tips (Fig. 2a). Rarely the phyllodia of submersed plants are
slightly dilated at their apices (Fig. 2c). Both S. teres and S. isoetiformis
have slender rhizomes (Figs. la, b and 2a, b). The surface of the achene
of S. teres has an irregularly crenate dorsal keel and 2-4 (or more)
prominent, rugose or irregularly knobby facial keels with no oil glands
apparent in the facial view (Fig. lc). The surface of the achene of S.
isoetiformis has a somewhat irregularly crenate to entire dorsal keel
and three or more low, non-rugose or non-knobby facial keels between
which the oil glands are conspicuous (Fig. 2f).

Sagittaria graminea var. graminea forms stout, horizontal rhizomes
from which shoots of the season emerge. If the rhizomes are submersed
(in Florida, at least), prominent rosettes of broad flat phyllodia occur
during winter. In spring, at about the time inflorescence scapes are
produced, new leaves arise which have elongate petioles and emersed
laminae (Fig. 1d). If the rhizomes are not submersed during winter,
rosettes of flat phyllodia are not produced and the spring leaves are of
the same type as the spring leaves of submersed plants. Both S. teres
and S. isoetiformis, as indicated above, have very slender, elongate
rhizomes. In regard to the winter rosette phyllodia of S. graminea var.
graminea, it is important to emphasize that they exhibit much varia-
bility in size, particularly length. They range from a few centimeters
long in shallow water to about 6 dm long in deeper water. Fluctuation
of water depth in places inhabited by this plant frequently varies
markedly in short periods of time. Thus plants which formed rosettes
in shallow water may have short phyllodia at a given time even
though the water may have recently become fairly deep owing to recent
rains. On the other hand, plants which have been submersed in fairly
deep water all winter have long phyllodia.

The achenes of S. isoetiformis and S. graminea var. graminea are much
alike with respect to keels and oil glands. We have not attempted to
examine and compare large numbers of them to ascertain whether or
not thy have subtle distinctive features of systematic value.

In conclusion, Sagittaria isoetiformis is considered specifically distinct
from S. graminea, var. graminea and from S. teres. The former occurs
in the coastal plain from southeastern North Carolina to peninsular
Florida and westward to southern Alabama. The latter occurs) from
eastern Massachusetts to southern New Jersey (to eastern Maryland
according to Fernald, op. cit.), S. graminea var. graminea is widespread
in eastern and central North America.

272

submersed plant. b. Habit, on

s. a. Habit,

Fig. 2. Sagittaria isoetiformi

rsed plant. e. Enlargement of fruiting in-

sections of phyllode of eme

florescence. f. Achene.

REFERENCES
BEAL, E. O. 1960. The Alismataceae of the Carolinas. Jour. Elisha Mitch. Sci. Soc. 76:
68-79.
eee C. 1955. Revision of the genus Sagittaria (Alismataceae). Mem. N. Y. Bot. Gard.
9: 179-2
na Gan , M. L. 1950. Gray's Manual of Botany. Am. Book Co., New Y

ork.
H, J. G. 1895a. A revision . - pe American species of See ices and Lopho-

I
tocar pus. Mo. Bot. ae 6th Ann.
95b. Notes ps Sbservacons on new or little known species. Mo. Bot.

Gard. 6th Ann. 115-116. Plat

MEIOTIC CHROMOSOMES
IN AFRICAN COMMELINACEAE
WALTER H. LEWIS
Stephen F. Austin State College, Nacogdoches, Texas

Following a limited study of Ethiopian Commelinaceae' an op-
portunity existed to expand the chromosomal survey to other parts
of Africa. Such an attempt seemed warranted for a number of reasons.
Early evidence indicated that basic numbers of genera had been mis-
interpreted, that polyploidy and aneuploidy, but rarely both, were char-
acteristic of different genera, and that infraspecific polyploidy and
aneuploidy were widespread and also typical of certain genera. I sup-
posed that a study of these features might lead to a clearer understand-
ing of their roles in speciation and significance in phylogeny. Not least
among my considerations were the varying definitions of commelinace-
ous subdivisions, perhaps best illustrated by Brenan’s? discussion on
assigning his newly described Triceratella to a tribe. Preliminary counts
in Africa in conjunction with existing data disclosed a marked similarity
of basic chromosome numbers for associated genera which in some
degree corresponded to major subdivisions of the Commelinaceae. All
these trends needed exploration and to this end the study was under-
taken.

MATERIALS AND METHODS

Immature flower buds and herbarium specimens of Cyanotis (Trades-
cantieae), and Aneilema, Commelina, and Murdannia (Commelineae),
were collected in east, central, and south Africa during September-
December, 1962. Buds were fixed in 4 parts chloroform, 3 parts absolute
ethanol, and 1 part glacial acetic acid; as soon as possible thereafter,
usually up to 10 days, the vials were airmailed to England for storage
at —40°C. Buds were examined for PMC meiosis in 2% acetic-orcein
and satisfactory squash preparations were mounted in euparal for future
reference. No difficulty in staining was experienced even after 9 months
of fixation and presumably if needed buds could be kept satisfactorally
at this temperature for longer periods of time. Whenever possible col-
lections from more than 1 plant were examined and these results are
indicated in parenthesis following my collection numbers in tables list-
ing the chromosome numbers. It is regrettable, particularly in view of
the marked frequency of infraspecific aneuploid and polyploid races
in most genera, that this procedure is not followed elsewhere. The
importance of knowing how many plants have a particular number
under these circumstances can not be overemphasized.

* Supe
SIDA 1 (5)

rscript numerals refer to list of references at end of. article.
I): 274—293. 1964.

279

Although useful meiotic plates were found for most collections, about
20% of those collected failed to show meiosis even when a wide range
of buds had been fixed. By referring to my field notes, I found that by
and large such buds had been fixed between 11 a.m. and 3 p.m. on clear
days in more or less exposed localities. On the other hand, meiosis was
found rapidly in buds fixed from 8-11 a.m. and from 3-5 p.m. on clear
days without shade or at any time during the day if cloudy or if the
plants were growing in the shade providing the ‘correct’ size had been

during mid-day under hot, exposed, often dry conditions, whereas this
decrease was not demonstrable either earlier or later in the day. Similar
daily ‘meiotic cycles’ have been noted in collections of Linaceae, Poly-
galaceae, and Rubiaceae from Mexico and the southwestern U. 5S.

A complete set of voucher specimens has been deposited at the Royal
Botanic Gardens, Kew (K); duplicates are in the U. S. National Museum
(US) and the Missouri Botanical Garden (MO). As the systematic study
of the African Commelinaceae proceeds at Kew, the unnamed collections
listed here will eventually be associated with binomials.

CYANOTIS

For 5 species listed in Table 1, basic numbers of x=11, 12 and 13
are reported with the 2=13 line new to Cyanotis. When these data
are combined with other African reports, °:'! 2=12 is the most common
basic number for species native to that continent.

Generally the meiotic process was regular. An exception was the
nondisjunction noted in about 20% of the anaphase plates of 1 plant of
C. sp. (Fig. 4) giving cells with 11+13 rather than the normal comple-
ment of 12 chromosomes. Otherwise the plates were normal in appear-
ance. Unequal distribution of chromosomes during anaphase has re-
cently been reported for Setcreasea."" From a casual observation of
pollen, I found only a small number of hollow and shrivelled grains,
no more than for those plants with normal disjunction, suggesting that
the loss or gain of 1 chromosome had no deleterious effect on the new
haploid cells.

Even the infrequent occurrence of nondisjunction in Cyanotis could
explain, at least in part, the fairly high incidence of aneuploidy within
species populations. For example, among a sample of 6 plants from the
Transvaal (Table 1), the homomorphic C. speciosa was found with 3
cytotypes: typical plants with 13,, (Fig. 5) as well as those with
13,,+1, and 15,, (Fig. 6). The anticipated trivalent and quadrivalent

ceivable that these hyperaneuploids had their origins from nondisjunc-
tion forming aneuploid races without, as yet, recordable morphological

276

TABLE 1.

Species n Voucher & locality :

Basic number x=11
C. barbata D. Don ll KENYA: Rift Valley Prov., Navasha
Dist., 2 miles W of west entrance to
Aberdare National Park, Lewis 5927
(1), Figs. 1-2.
Basic number x=12

C. longifolia Benth. 12 N. RHODESIA: N W Prov., Mwinilunga

(dwarf form) Dist., Mujileshi River, ca.4 miles E of
Angola-N.R. border, Lewis 6133 (2).

(tall form) 12 N. RHODESIA: N W Prov., Mwinilunga

Dist., Zambesi River, 4 miles N of
Kalene mission, Lewis 6206 (2).

border & 1-4 miles SW of Mujileshi
River, Lewis 6147 (2). Figs. 3-4.
Basic number x=13

C. foecunda Hassk. 13 KENYA: Rift Valley Prov., Trans Nzoia
Dist., ENE slope of Mt. Elgon, Lewis
0964 (1).

C. speciosa (L.f.) Hassk. 13° N. RHODESIA: N W Prov., Mwinilunga

Dist., 1-4 miles E of Angola-N.R.
border, Lewis 6134 (1); 3-4 miles SE
of Angola-N.R. border & 1-4 miles SW
of Mujileshi River, Lewis 6157 (1). S.
AFRICA: Natal, Hlabisa Dist., Charters
reek, Lewis 6304 (2).
13,+1,,2,, S. AFRICA: Transvaal, Pretoria Dist.,
retoria, Wonderboom, Lewis, 6344
(3,13,,3 2,13,,+1,;1,15,p, Figs. 5-6.

differences. Other examples of infraspecific aneuploidy have been
observed in more heteromorphic species than C. speciosa. The present
count of n=11 for C. barbata from Kenya confirms the report from
Ethiopian populations,'’ but Sharma and Sharma?! found n=12 for an
Indian collection. It would be interesting to know how widespread the
n=12 race is in Asia and whether or not C. barbata is multibasic ac-
cording to a continental distribution. It is well worth noting that the
interpretation of meiosis at diakinesis is often confusing in Cyanotis
and other Commelianaceae and this factor can not be overlooked in

paral
explaining some of the diversity in recorded chromosome numbers. As

resembles a bivalent having 2 chiasmata during mid-diakinesis. Thus
the meiotic number for the PMC illustrated in Fig. 2 might be given as
nm=12 (and was by a cytologist who examined the cell), but later
diakinesis on the same slide clearly shows PMCs with only 11 bivalents

stages of meiosis; it does, however, stress the danger of drawing too
hasty a conclusion from diakinesis alone. Kammathy and Rolla”
found a n=11 race for C. arachnoidea C.B.Cl., a species typically having
n=12'*>-25 and ‘fragments’ were noted by Islam and Baten’ for C.
cristata Schult.f.*

When the results for 18 Afro-Asian species with known chromosome
numbers are summarized, we find: 1 species with n=8; 2 species with
n=10; 1 species with n=11: 2 species with n=11 and 12; 9 species or
50% with n=12, 24, 36; 2 species with n=13; and 1 species with
n=14. By far the most frequent basic number is therefore x= pre-
dominantly at the diploid level but including the only known polyploids
(C. adescendens, 4x7! and C. tuberosa, 2x, 4x, 6x'*'*5), This trend is
not likely to be altered as more species are studied and x=12 should
remain the central focus of chromosomal differentiation in the genus.

How does this number fit those of other genera usually grouped with
Cyanotis? If one follows Briickner,* who included Cyanotis in the
Subfamily Tradescantieae, Tribe Hexandrae, then Tradescantia, Leptor-
rhoeo, Setcreasea, and Zebrina are the 4 genera most closely associated.
The most frequent basic number for Tradescantia is x=6, for Setcreasea
x—6, for Zebrina x=12, and none is known for Leptorrhoeo. If one follows
Clarke,* who placed Cyanotis in the Tribe Tradescantieae, the only definite
count known for those genera listed near Cyanotis is n=12 for Floscopa.”
Although the data are incomplete, the evidence reveals basic numbers

f x—6 and 12 for genera associated with Cyanotis by Bruckner and
Clarke. It also supports the inclusion by Clarke and Woodson” of
Floscopa in the Tradescantieae since these numbers are very rare in the
Commelineae where Briickner placed Floscopa. This suggests that x=6
is an original basic complement for these genera, and perhaps for the
Tradescantieae, and that the x=12 is a polyploid derived from such a
prototype having become widespread as a basic number for several
genera in this circle of affinity. A significant example would be Cyanotis,
the largest genus in the Tradescantieae.

wo species of “Cy anotis” considered by Islam and Baten need clarifying. Reference is
made to Cyanotis zenonii of Darlington’? which Darlington’? long ago corrected to Cam-
soli zanonia ae Secondly they refer to “C. spironema a presumably from the
legend on p. of Darlington,“ which is sim ply “C.” for the third figure of the plate
illustrating not the chromosomes of Cyanotis but rather of ions fragrans (= Callisia

fragrans (Lindl. : "Woodso n).

278

Five minor lines of descent, x=8, 10, 11, 13, and 14, are each repre-
sented in Cyanotis by only 1 or 2 species. These could have formed by
a gain or loss of chromosomes from x=12 in a similar way to the ex-
amples of infraspecific hypo- and hyper-aneuploidy outlined above
except that they have reached a morphological differentiation recogniza-
ble at the rank of species. Probably C. somaliensis C.B.Cl. with n=14°
has not yet attained such a level. According to Brenan (oral communi-
cation), this species may represent but a part of the C. foecunda complex
(n=13) and as such C. somaliensis may eventually be recognized merely
as an infraspecific aneuploid race.

Based on chromosome numbers and frequencies, a hypothetical evolu-
tion of Cyanotis has been constructed (Fig. 7). Some alterations in the
figure are anticipated as the cytotaxonomic analysis of the genus pro-
ceeds (e.g., changing the x=14 basic line represented by C. somaliensis
to x=13+1), but these are not expected to alter the principal features
illustrated.

I have ignored chromosome size largely because meiotic chromosomes
are inexact for comparative purposes and most research has been con-
fined to meiosis. Exceptions are 2 photomicrographs of pretreated
somatic cells illustrated by Shetty and Subramanyan.”* From these I
estimate the chromosomes of C. axillaris (L.) R. & S. to be 3.2-5.5
microns in length and for C. arachnoidea C.B.Cl. only 1.8-2.8 microns.
In the same paper Shetty and Subramanyan described bivalents of C.
papilionacea as “larger” than and those of C. arachnoidea as “smaller”
than the other species studied which included C. awillaris, C. cristata
(L.) D. Don, C. fasciculata R. & S., C. tuberosa R. & S., and C. villosa
R. & S. Their bivalents would be considered as more or less inter-
mediate in size. On comparing these results with the meiotic plates of
C. barbata, C. speciosa, and an undescribed species (Fig. 1-6), I find the
bivalents and chromosomes approximately intermediate in size and quite
comparable with the majority illustrated by Shetty and Subramanyan.
These sketchy data suggest that mitotic chromosomes and bivalents of
Cyanotis species are predominantly of an intermediate size (e.g., 3.2-
5.9 microns in C. axillaris) with a few species having smaller (e@.£.,
C. arachnoidea) or larger (e.g., C. papilionacea) chromosomes.

When the anaphase I chromosomes of C. sp. (Fig. 3-4), Commelina
benghalensis (Fig. 10-11), and C. diffusa (Fig. 14) are measured, the
meiotic chromosomes of Cyanotis average 3.0 microns while those of
the Commelina species are smaller at 2.6 and 2.1 microns, respectively.
Apparently Cyanotis chromosomes on the average are somewhat larger
than are those of Commelina and also Murdannia (see further discus-
sion below).

ANEILEMA
The 5 species of Aneilema examined (Table 2, Fig. 8-9) are grouped

279

TABLE 2.
GAMETIC CHROMOSOME NUMBERS IN AFRICAN ANEILEMA

Species n Voucher & locality

Basic number x=9
A. sp. aff. pedunculosum 9 KENYA: Rift Valley Prov., Trans Nzoia
C.B.Cl. Dist., ENE slope of Mt. Elgon, Lewis
5973 (2), Fig. 8.
Basic number x=13
A. tacazzeanum Hochst. 13 UGANDA: E Prov., Teso Dist., 1.8 miles
W of Wera, Lewis 5999 (2), Fig. 9.
A. welwitschii C.B.Cl. 26(+1?) CONGO: Katanga Prov., Lualaba Dist.,
15 miles NNW of Kalene mission, Lewis
6229 (1); N. RHODESIA: N W Prov.,
Mwinilunga Dist., Mujileshi River, 4.5
miles E of Angola-N.R. border, Lewis

6143 (1).
Basic number x=15 or 10
A. aequinoctiale 30 KENYA: Central Prov., Meru Dist., 8
(P.Beauv.) Kunth miles NE of Runyenje’s, Lewis 5911

(1); S. AFRICA: Natal, Durban Dist.,
Durban, Lewis 6279 (1)
Basic number x=16 or 8

A. johnstonii K. Sch. 16 N. RHODESIA: N Prov., Abercorn Dist.,
Chilunoma River, nr. Abercorn, Lewis
Obie: (3);

under 4 newly reported basic numbers, x=9, 13, 15 (or 10), and 16
(or 8). To these can be added the counts of n=14 for A. montanum
Wight!? 6 giving 5 basic complements for a sample of only 6 species.
(Many species with established chromosome numbers have been pub-
lished under Aneilema, but all are referrable to Murdannia.) The genus
is a rather large one and until more data are accumulated, little can
be noted regarding chromosomal trends other than that aneuploidy and
polyploidy have apparently played significant roles in the evolution of
Aneilema giving rise to a multibasic group of species at several levels
of ploidy.

COMMELINA
From a sample of 37 populations involving at least 26 taxa, basic
numbers of x=11, 13, 14, and 15 are reported for Commelina (Table 3,
Fig. 10-16). Those species with x=15 are in the majority, about 70%
of the total; species with x=14 and 13 are infrequent, and the x=11
series is represented solely by C. benghalensis. Infraspecific polyploidy
is reported for C. africana with 2x, 4x, and 8x races and for C. beng-

280

halensis with 2x (Fig. 10) and 4x (Fig. 11) races. The results also add
a diploid race (Fig. 12) to the report of n=30 (4x) for C. imberbis."*
No infraspecific aneuploid is recorded and regular melosis was char-
acteristic throughout (ignoring clumping and various adhesions attri-
buted to fixation).

In a strict sense, x=13 is a newly reported basic number. The re-
lated Commelinantia is known with n=13' and its transfer to Com-
melina by Woodson* is now supported by the existence of similar com-
plements in typical commelinas. However, Rowley” reported Com-
melinantia as having pollen with 3 colpi rather than the single colpus
ound for all other Commelinaceae studied: hence it might be argued
that this unique micromorphological feature, together with certain gross
characters, is worthy of generic recognition.

Seven species listed in Table 3 have been examined previously, all
but one by Morton’ from west African material. His results for these
species are summarized in Table 4 together with those for the present
study and for others. Morton’s counts are based on x—14 in contrast
to mine and most others which are characteristically x=15. The excep-
tion is x=11 for C. benghalensis having diploids widely distributed in
India and both diploids and tetraploids frequent in Africa. The n—ca.24
count by Anderson and Sax! and 2n=ca.68 by Darlington® suggest 4a
and 6x races; unfortunately original localities were not given. But
Morton’s data are not similar. Possibly infraspecific aneuploidy exists
for all these species, yet I think it peculiar that this mechanism should
be largely confined to west African populations. It is clearly infrequent
elsewhere. Regrettably my collections based on x—14 must for the
present remain unnamed; among these a verification of some of Mor-
ton’s numbers may be possible.

At least three suggestions regarding the original basic number of
Commelina have been proposed. Certainly the oddest is found in an
abstract by Deodikar* in which no evidence is given to corroborate the
statement that “there are two polyploid series in primary and secondary
chromosomal balance with 8 and 16 as their respective monoploid num-
ber.” Not only does this quotation lack meaning to me, but n=8 has yet
to be found in the genus. Unquestionably this abstract is to be ignored
until some results are published to support the conclusions. On the
basis of associations of groups of bivalents, Sharma*’ has suggested
x=4 as the basic number of Commelina. I find the evidence incon-
clusive based as » is on the very questionable premise of bivalent
association and then for only 2 species. Perhaps it is noteworthy that
in a later paper Sharma and Sharma”! fail to make further use of such
associations in deriving basic numbers and evolutionary groups in the
family. Morton'* has proposed x=7, but I have shown that his results
are not characteristic of Commelina as known today. In short, I find
little evidence to support x=4, 7, or 8 as basic numbers.

281

TABLE 3.
CAMETIC CHROMOSOME NUMBERS IN AFRICAN COMMELINA

Species n

Voucher & locality

Basic number x=11
C. benghalensis L. 1

any

Nw
iw)

Basic number x=13

C. eckloniana Kunth 13
C. cf. eckloniana Kunth 13
13(+1?)
Basic number x=14
Cy sp. 1 14
C. sp. 2 14
C. sp. 3 14
C. sp. 4 28
Basic number x=15
C. africana L. 15

var. africana

i

var. 1 3

KENYA: Rift Valley Prov., Trans Nzoia
Dist., ENE slope of Mt. Elgon, Lewis
5961 (1), Fig. 10.
UGANDA: N Prov., Karamoja Dist.,
base of Mt. Moroto, nr. Moroto, Lewis
5996 (1), Fig. 11.

N. RHODESIA: N Prov., Abercorn Dist.,
Chilunoma River, nr. Abercorn, Lewis
6112 (3).

N. RHODESIA: N W Prov., Mwinilunga
Dist., Mujileshi River, 4.5 miles E of

19 miles SSW of Mutschatsha, Lewis
6142 (2).

N. RHODESIA: N W Prov., Mwinilunga
Dist., 1 mile E of Ikelengi, Lewis 6189

(4).
N. RHODESIA: N W Prov., Mwinilunga
Dist., Mujileshi River, 5-6 miles SE of

Dist., Zambesi River, 4 miles N o
Kalene mission, Lewis 6196 (3), Fig. 13.
N. RHODESIA: N W Prov., Mwinilunga
Dist., 3-4 miles SE of Angola-N.R.
border & 1-4 miles SW of Mujileshi
River, Lewis 6146 (2).

S. AFRICA: Natal, Hlabisa Dist., Chart-
ers Creek, Lewis 6305 (1). UGANDA:
W Prov., Toro Dist., Queen Elizabeth
National Park, Lewis 6011 (2).

S. AFRICA: Natal, Hlabisa Dist., 4.

miles W of Charters Creek, Lewis 6309
Cs

282

TABLE 8 (cont.)
Commelina africana
var. 2 30 S. AFRICA: Natal, Estcourt Dist.,

Drakensberg Mts., base of Mt. Cham-

pagne, Lewis 6266 (1).

S. AFRICA: Transvaal, Pretoria Dist.,

Pretoria, Wonderboom, Lewis 6345 (2).

S. RHODESIA: Wankie Dist., Victoria

Falls, Lewis 6247 (1), Fig. 14. UGAN-

DA: W Prov., Bunyoro Dist., 12 miles

S of Victoria Nile on road to Masindi,
Lewis 6004 (2).

C. imberbis Ehrenb. ex 1 ANYIKA: Tanga Region, Tanga

Hassk. Area, 6.3 miles W of Tanga, Lewis 6062

(2), Fig. 12.

KENYA: Rift Valley Prov., Trans Nzoia

Dist., ENE slope of Mt. Elgon, Lewis

5960 (1).

N. RHODESIA: N W Prov., Mwinilunga

Dist., 2 miles W of Ikelengi, Lewis 6193

(1), Fig. 15.

C. sp. 5 15 N. RHODESIA: N W Prov., Mwinilunga
Dist., 3-4 miles SE of Angola-N.R.
border & 1-4 miles SW of Mujileshi
River, Lewis 6148 (3).

C. sp. 6 15 N. RHODESIA: N W Prov., Mwinilunga
Dist., Zambesi River, 4 miles N_ of
Kalene mission, Lewis 6197 (3).

©

var. 3 6

Q
on

diffusa Burm.f. 1

on

on

C. purpurea C.B.Cl. 1

QQ
on

. scaposa C.B.Cl. 1

C. sp. 7 lo S. AFRICA: Transvaal, Pretoria Dist.,
Pretoria, Wonderboom, Lewis 6347 (1).

C. elgonensis Bullock 30 KENYA: Rift Valley Prov., Trans Nzoia
Dist., ENE slope of Mt. Elgon, Lewis
9974 (1).

C. gerrardii C.B.Cl. 30 S. AFRICA: Natal, Durban Dist., Dur-

ban, Lewis 6281 (1); Isipingo Beach,
Lewis 6283 (1) & 6285 (1): Hlabisa
Dist., Charters Creek, Lewis 6300 (1).
S. AFRICA: Natal, Hlabisa Dist.,
Charters Creek, Lewis 6296 (2); Trans-
vaal, Pretoria Dist., Pretoria, Wonder-
boom, Lewis 6346 (1). S. RHODESIA:
Salisbury Dist., Salisbury, Lewis 6259

Q
S

livingstonii C.B.Cl. 3

(1).
S. RHODESIA: Salisbury Dist., Salis-
bury, Cranborne, Lewis 6252 (1).

w
Oo

C. welwitschii C.B.Cl.

283

C. sp. 8 ca.30 KENYA: Rift Valley Prov., Trans Nzoia
ist., Kitale, Lewis 5981 (1).
C. sp. 9 30 N. RHODESIA: N W Prov., Mwinilunga

Dist., nr. Mujileshi River, 4 miles E of
Angola-N.R. border, Lewis 6185 (1); 7
miles NW of Kalene mission, Lewis
6226 (3), Fig. 1

C. sp. 10 30 TANGANYIKA: 's Highlands Region,
Mbeya Area, Mbeya Range (ca.8 miles
NE of Mbeya), Lewis 6085 (2).

C. sp. 11 ca.45 N. RHODESIA: N W Prov., Mwinilunga

Gonien & 1-4 miles SW of Mujileshi
River, Lewis 6145 (1).

In an attempt to offer an alternate suggestion, I have summarized
the gametic numbers known for Commelina in Table 5 together with
frequencies of cytotypes. Each cytotype corresponds to a species except-
ing the infraspecific polyploids and rarely aneuploids. These are repre-
sented once for each race. The most obvious fact is the predominance
of the x—15 line totaling 75% of all cytotypes. For this group almost
twice as many polyploids as diploids are known and these are to a
level of 10x. Minor lines of descent account for the remaining cyto-
types with basic numbers of x= 11, 12, 13, 14, and 16, chiefly of diploids,
infrequently of tetraploids, and of only one questionable hexaploid race.
The table does not reveal the infraspecific polyploidy now known for
8 species or 20% of those species studied. Nearly one-half of these, such
as C. africana, C. benghalensis, and C. obliqua Ham., each have 3 races
at different levels of ploidy. When one considers that many species have
only 1 recorded number from a single plant, the incidence of infraspecific
polyploidy becomes even more remarkable and obviously this percentage
can be expected to increase as more counts are accumulated. As I have
noted above the definite examples of infraspecific aneuploidy are much
less notable. Collections originally published under C. nudiflora L. were
found with primarily n=15 and 2n=30** with 2n=28 and 56,” all from
India, and with 2n=56 by Darlington® without indication of source —_
by Anderson and Sax! from North American material as Morton™ re-
ported). These give C. nudiflora 2 primary numbers of n=15 and 14
with the latter having a tetraploid race. If C. diffusa is considered a
part of this complex, then, as I recorded in Table 4, the n=15 comple-
ment is by far the most common and probably the n=14 line arose by
hypoaneuploidy from plants with n=15. This is the only verified example
of aneuploidy within a species of Commelina; I expect that when the
reports of Morton can be more fully explored and when my collections
based on x=14 are named more cases with a pattern similar to C.

TABLE 4.
PRESENT CHROMOSOME NUMBERS IN COMMELINA AND
PREVIOUS COUNTS

Species Lewis Morton" Others

C. africana n= 15,30,60 2n=28 mlb?

C. diffusa i eel 5) Pn 28 Visors Dee ae

C. forskalaei n=15 2n=28 m= 14% napa,
n—=15 & 2n= 30°??"?

C. imberbis n=15 n=30"

C. gerrardii n=30 2n=56

C. livingstonii n= 30 2n=56

C. benghalensis (2x)n=11 2n=28 ade es 1 1B
a1) 2 2h eee

(polyploid) n=22 2n—=56 n=22"); n=ca.24;

2n=ca.68"’

nudiflora may come to light. Undoubtedly, however, the contribution of
aneuploidy to speciation in Commelina is secondary to the role of poly-
ploidy.

Do these facts and trends suggest an original chromosome number for
Commelina? The x=15 line is certainly of secondary origin and as such
gives no direct answer to this question. But consider Commelina in rela-
tionship with Murdannia (Table 6), a genus having a characteristic basic
number of x=10. Except for Aneilema, these are the only large genera
in the Commelineae and they are probably indicative of the tribe as
a whole. The dominance of 2x=10 and 15 in the tribe suggests an ancient
basic number of 5 giving rise by polyploidy to the widespread occurrence
of multiples of this number today.

In referring briefly to chromosome size in Commelina, a topic already
introduced under Cyanotis, I should mention again that a discussion
based solely on meiotic figures is not satisfactory. However, Anderson

Sax' noted small chromosomes for C. benghalensis and mitotic
chromosomes illustrated for C. diffusa’ measure 1.9-3.7 microns in
length. In meiosis there is not much difference between the anaphase I
chromosomes of C. benghalensis (Fig. 10-11) and C. diffusa (Fig. 14).
These chromosomes as well as the bivalents of C. imberbis (Fig. 12), C.
scaposa (Fig. 15), and C. sp. (Fig. 16) would all be described as more
or less small. Yet the bivalents of an unnamed Commelina with n=14
(Fig. 13) are of intermediate size and not unlike the majority figured
for Cyanotis.

TABLE 5.
CYTOTYPES REPORTED IN COMMELINA WITH FREQUENCY OF
PL i

n= Ploidy Total
22x 4x 6x 8x 10x
11 1 1 1(?) 3
12 1 1
13 3 3
14 4 2 6
15 12 18 9 2 1 42
16 1 1

* Excluding results of Morton,'® several cirea counts, and the meiotically irregular Sx

C. salicifolia.”®

MURDANNIA

All plants from 5 populations of M. simplex (Vahl) Brenan were
found with n=20 and regular meiosis. These include collections from:
N. RHODESIA—N.W. Prov., Mwinilunga Dist., 3-4 miles SE of Angola-
N.R. border & 1-4 miles S.W. of Mujileshi River, Lewis 6156 (1), 5-6
miles SE of Angola-N.R. border, Lewis 6172 (2), 4-5 miles SE of Angola-
N.R. border, Lewis 6179 (1) (Fig. 17); TANGANYIKA—E Region

more widespread than are the hexaploid'**! and octoploid?® races which
are known only from Asia.

A total of 24 cytotypes have been published for 15 species of Mur-
dannia (see excellent review by Shetty and Subramanyam). Six species
are known to have 2 or 3 levels of ploidy although several of these may
be attributed to misidentification, but at most a fraction. When the n
numbers are summarized according to frequency and level of ploidy
(Table 6), the most obvious feature is the high frequency (71%) of the
basic complement of x=—10. This frequency is comparable to the 75%
for x=15 in Commelina. In addition more than one-half of all cytotypes
are polyploids, multiplied to the 8x level in 1 or possibly 2 lines, and
this too is parallel to the situation in Commelina. Although the range

footnote in Table 6. The most frequent number in these species is
usually questionable as for M. semiteres (Dalz.) Santapau with n—7,”!
10,7" 12,% and 20,*? but possibly multiples of 10 will be found most
frequently for this species. For the present discussion, it is clear that

286

CYTOTYPES REPORTED IN MURDANNIA WITH FREQUENCY OF
PLOIDY

n= . Ploidy Total

2x 4x 6x 8x
Ce 1 1
8(?)* 1(?) 1
9 1 1
10 6 6 4 1 17
11 1 1 2
12' 1 1
15 1 1

* Known only as infraspecific aneuploids.

n=T7 and 12 can not represent lines of descent or basic numbers. This
leaves x=9, 11, and 15, each having 1 species. The origin of z=9 by the
loss of a chromosome pair and of x=11 by a similar increase from x=10
are reasonable speculations, but the origin of w=15 offers several pos-
sibilities. Assuming that the number of 2n=30 for M. keisak (Hassk.)
Hand. Mazz." is not that of a naturally occurring triploid, the species
could have formed by a succession of chromosomal gains from n=10,
11, etc., but this is a long route beset with many gaps. Alternately M.
keisak could have arisen from stocks of 10 and 5 or 5, 5, and 5. This
hypothesis also has major drawbacks (not least among them is the ab-
sence of a species with n=5); even so the known chromosome numbers
in the genus, the high frequency of polyploidy, and the low frequency
of aneuploidy all strongly suggest this origin. I might note that the
relationship of Murdannia and other Commelineae to the Tribe Pollieae
needs further study and it should not be overlooked that Pollia is known
with n=5."

In summary, I propose that x=5 is an ancient, probably extinct, basic
number for Murdannia represented today by a dominant x=10 line com-
posed of species with n—10, 15, 20, 30, and 40.

In regard to chromosome size, the bivalents of M. simplex (Fig. 17)
are smaller than those illustrated for Cyanotis and Aneilema and about
equal or smaller in size than those of Commelina. Bivalents of M. elata
(Vahl) Briickn.’” are similar to M. simplex.

CONCLUSIONS
1. Basic numbers of x=8, 10, 11, 12, 13, and tentatively 14 are known
for Cyanotis. This large genus is thought to have arisen from a x=6
prototype and to have had its chromosomal! differentiation from a n=12
stock foremost by aneuploidy and secondarily by polyploidy. Infra-

287

specific aneuploidy is widespread while infraspecific polyploidy is re-
stricted

2. The most common basic numbers for Cyanotis and the Tradescan-
tieae as a whole are x=6 or 12

3. Basic numbers of r=9, 13, 14(7), 15(10), and 16(8) have been found
for Aneilema. Results from the few species examined suggest a fertile
area for additional study.

4. Species with a basic number of x=—15 form the dominant line of
descent in Commelina. Based on frequency, all other lines are of minor
importance as is infraspecific aneuploidy. In contrast, the majority of
known species are polyploids and many have polyploid races.

A line of descent for Murdannia is x=10. All others
are of lesser significance and, as for Commelina, most species are poly-
ploids with infraspecific polyploidy more common than _ infraspecific
aneuploidy. This parallel chromosomal pattern of speciation in Mur-
dannia and Commelina is opposed to the major role of aneuploidy and
minor contribution of polyploidy in the speciation of Cyanotis.

6. Although the common basic numbers of Commelina and Murdannia
differ, i.e, x=10 and 15, prototypes of x=5 are suggested for both
genera.

7. Small chromosomes typify Commelina and Murdannia, those of
Aneilema are somewhat larger, while those of Cyanotis range from
small to large with a majority intermediate in size.

8. The most frequent basic number encountered in the Tradescantieae
is x=6 or multiples of it, while the most frequent numbers found in
the Commelineae are multiples of 5. Probably the prototypes of the
tribes differed, the Tradescantieae from a stock based on x=6 and the
Commelineae on x=5. A consideration of typical basic chrom
numbers in the classification of the Commelinaceae will undo ener
contribute to a more natural grouping of genera than has been hereto-
fore possible

ACKNOWLEDGEMENTS

This study was pena by grant number G-21818 from the National ae Founda-

tion and completed at the Royal Botanic Gardens, Kew. To the Director of Kew, Keeper of

rbarium, and ‘Dr. Keith Jones of the Jodrell L praeeee I am grateful te the many
facilities placed at my disposal. Special appreciation is due Mr. J. P. M. Brenan of Kew for
his determinatio y lh ium = specimens. | ‘adait my sincere
thanks to th y botanists ek Kenya to South Africa for t assistance in the field
and herbarium and to Mr. S. R. J. White who accompanied me throughout the tr rip.

REFERENCES

ANDERSON, E. AND K. SAX. 1936. A cytological monograph of the American
species of Tradescantia. ae be 97: 433-476.

RENAN, J. = M. Taepdiella, a new genus of Commelinaceae from Southern
Redes ae I: ae.

KNER, iC 1930. Commelinaceae, #7 Engl. & Prantl, Nat. Pflanzenfam. ed. 2,
ISa: ae ee
4. CL KE, C. B. 1901. Commelinaceae, iz Flora Trop. Afri 8:
5. DA co C, D. 1929. Chromosome behaviour and Sees Tybadig in the

Tradescantieae. J. Genet. 21: 207-286.

288

6. DARLINGTON, C. D. 1937. Recent Advances in Cytolo ogy. ed.
7. seo Cc. D. 1937. Chromosome cee viour and ede | year in
igs inl J. Genet. 35:
re G. B. 1950. A pelminae note on the cytogenetic survey of the genus
as in India. Proc. 37 Ind. . Cong. 1950, pt. 7
9. GANGULY, J. K. 1946. The somatic and meiotic Ae reer of Commelina bengha-
lensis Linn. Curr. Sci. 15: 112.
SLAM, A. S., AND A. BATEN. 1952. Cytology of ee Nature 169: 457-458
KAMMATHY, R. V., AND S. R. ROLLA. 1962.
II: eres ehseeiations: Bot. Surv. India 3 (1961): ‘167-1
12, KAMMATHY, R. V., AND S. R. ROLLA. 1962. ey on Inia Commelinaceae—
IIT: ee observa ie a Bot. Surv. India a fe
H., H. L. STRIPLING, AND R. nd a eee numbers
for some nigel Aine of ~ southern United States Mexico. a oder ra 147-161
14. LE , W.H., AND TADDESSE E. 1964. Chromosome numbers in te Cone:
melinaceae, tka 4 (in press
{

LIK, C. P. ote Chromosome number in some Indian angiosperms: monocotyle-

oe MEHRA, K. L., S. ry ae UQI, AND ‘ — ee 1961. A cytotaxonomical

study five species ee Setcreasea. Phyton 17:

17: MITSUKURL Y. 1947. Cytological serve in “Commein aceae. I. Chromosome num-
bers of Japanese Ete Jap. J. Genet. 22: 18-

18. oo J. K. 1956, Cytotaxonomic ig on oe Gold Coast species of the genus
Connie 4 Linn. J. Lin oe Bot. 55: 505-

9, ANIGR AHL, G., “AND R. V. KAMMATI TY. 1961. Studies in the evytomorpnalogy
of ae *ma sensu lato in eastern India. Proc. 48 Ind. Sci. Cong. 1961, pt. es

20. PANIGRAHI, G., AND R. VY. KAMMATHY, ee a seus in certain
species of Commelina in eastern India. Proc. 49 Ind. . Cong. 1962, pt. 3: 30.

: 329-

1 RAGHAVAN, R. S., AND S. R. ROLLA. ee Cytological pe note on the
Indian species of Commelinaceae. Curr. Sci. 30: 310-311.

22. ROWLEY, J. R. 1959. The fine structure of the pollen wall in the Commelinaceae.
Grana cy 2: 3-31.

23. SHARMA, A. K. 1955. Cytology of some of ne mabe Zi ee Commelinaceae and
its beatae on the wea of phylogeny. Ger 27

24. SHARMA, A. AND SHARMA. 1958 Baee: investigations on cytology of
members of Cormelinucese ee spacial reference to the role of polyploidy and the origin

63- ae

25. SHETTY, B. V., AND eon 1961. Cytology of some taxa of Com-
melinaceae. Proc. 48 Ind. Sci - ong. 1961, pt. 3: 299.

26. SHETTY, B. V., AND K. ea teas 1962. Cytological studies in Com-

27. SIMMONDS, N. WV. Los, Chromosome behaviour in some tropical plants. Heredity
8: 139- 146.

28. ODSON, R. E. 1942. Commentary on the North American genera of Com-
melinaceae. Ann. Mo. Bot. Gard. 29: 141-154.

Fig. 1-6. Meiosis in Cyanotis. 1300X%. Fig. 1-2. C. barbata, 11,, with the
nucleolus in Fig. 2 resembling a bivalent, 5927. Figs. 3-4. C. sp.,, 12412
and 11+13, 6147. Fig. 5-6. C. speciosa, 13,, and 15,,, 6344.

Fig. 7. Hypothetical representation based on chromosome number of the
evolution of Cyanotis from a x=6 prototype to a dominant extant group
with x=12. From this stock it is suggested that cytotypes have evolved
by hypoaneuploidy, hyperaneuploidy, and euploidy giving rise to a
genus multibasic at the 2a” level and (as presently known) unibasic at
the 4a and 6x levels. Euploidy is vertically illustrated, aneuploidy hori-
zontally as interspecific (solid circles) or infraspecific (hollow circles).
The smallest circle represents one cytotype and is equivalent to one
species excepting the infraspecific aneuploids (small circle for each
cytotype). Other circles to scale.

HYPOANEUPLOIDY
-2

EUPLOIDY

EUPLOIDY

PROTOTYPE

HYPERANEUPLOIDY

6X

4X

2x

292

2 13

Fig. 8-13. Meiosis in Aneilema and Commelina. 1300X, 1550X% for Fig.
10-11. Fig. 8. A. sp. aff. pedunculosum, 9,,, 5973. Fig. 9. A tacazzeanum,
13,,, 9999. Fig. 10-11. C. benghalensis, 11411, 5961, and 22422, 5996.
Fig. 12. C. imberbis, 15,,, 6062. Fig. 13. C. sp., 14,,, 6196.

eee ae

293

Fig. 14-17. Meiosis in Commelina and Murdannia. 1550X%. Fig. 14. C.
diffusa, 15,,, 6247. Fig. 15. C. scaposa, 15,,, 6193. Fig. 16. C. sp., 30,,, 6226.
Fig. 17. M. simplex, 20,,, 6179.

NOTES

MORE ADDITIONS TO THE LOUISIANA FLORA.—The following
taxa apparently represent new records for the state of Louisiana. Voucher
specimens are deposited in the herbaria of the University of South-
wescern Louisiana (LAF) and (except for Cyperus uniflorus) of South-
ern Methodist University (SMU).

Bulbostylis barbata (Rottb.) C. B. Clarke. Iberia Parish; locally com-
mon in sandy soil in sugarcane field, Weeks Island, John W. Thieret and
William D. Reese 9703, 15 October 1962.

Cyperus difformis L. Lafayette Parish: in fallow rice field 13 miles
west southwest of Lafayette near Acadia Parish border, A. J. Delahous-
saye 71, 23 October 1963.

Cyperus Oxylepis Nees. Cameron Parish: back ridge along Gulf of
Mexico about 2 miles east of Holly Beach, Thieret 8818, 7 July 1962.
Lafayette Parish: disturbed soil at edge of swamp about 3 miles north-
east of Broussard, Thieret 16728, 20 November 1963. St. Mary Parish:
disturbed soil in clearing in upland woods, Cote Blanche Island, Thieret
16316, 14 September 1963.

Cyperus uniflorus Torr. et Hook. Grant Parish: Pollock, John Lynch
s.n., 9 September 1938.

- Cypselea humifusa Turp. St. Landry Parish: in drying mud of cow
path in clearing about 4 miles northwest of Grand Coteau, Thieret 16462,
5 October 1963.

Drymaria cordata (L.) Willd. Lafayette Parish: among grasses at edge
of grazed field, southwest side of Lafayette, Thieret 16594, 28 October
1963.

Phyllanthus urimaria L. St. Mary Parish: locally common i ed of
little-used dirt road through upland woods, Cote Blanche Island, Thieret
16712, 15 November 1963.

Abutilon hulseanum (Torr. et Gray) Torr. ex Chapm., St. Mary Parish:
disturbed soil near salt mine, Cote Blanche Island, Thieret 15876, 7 May
1963.

Bowlesia incana Ruiz et Pavon. St. Martin Parish: locally common at
edge of willow dominated depression along highway 90, about 6.8 miles
south of Broussard, Thieret 17063, 3 April 1964. Although credited to
Louisiana in Small’s Manual ,Bowlesia incana is cited as occurring “from
Texas to California” by Mathias and Constance in their treatment of
Umbelliferae in North American Flora.

Facelis retusa (Lam.) Sch.-Bip. East Feliciana Parish: disturbed soil
at roadside just west of Jackson, Thieret 17255, 10 May 1964. St. Helena
Parish: roadside 3 miles east of Pine Grove, Thieret 17318, 15 May 1964.
Vermilion Parish: shell ridge in vicinity of University of Southwestern
Louisiana Field Station, Redfish Point, west side of Vermilion Bay,
Reese 4183, 30 April 1961.

295

Mikania cordifolia (L.) Willd. Iberia Parish: along dirt road in upland
woods, Weeks Island, Thieret 16566, 25 October 1963. Lafayette Parish:
abundant in clearing in woods near Vermilion River, south side of La-
fayette, Reese 6804, 7 October 1963. St. Landry Parish: edge of clearing in
woods about 4 miles northwest of Grand Coteau, Thieret 16494, 14 October
1963. St. Mary Parish: at roadside in upland woods, Cote Blanche Island,
Thieret 16703, 15 November 1963. This species, first recognized in the
Lafayette area by Dr. Robert Kral, is locally common here.—John W.
Thieret, University of Southwestern Louisiana, Lafayette, Louisiana

TIGRIDIA haar (HERBERT) SHINNERS, COMB. NOV.
(IRIDACEAE). — Based on Nemostylis? purpurea Herbert, Bot. Mag.
66: t. 3779 text, p. 3). 1840. Eustylis purpurea (Herbert) Engelmann
& Gray, Boston Journ. Nat. Hist. 5: 236 (Pl. Lindh. I, repr. p. 27). 1845.
Tigridia buccifera S. Watson, Garden & Forest 2: 412. 1889. Watson’s
generic assignment for this species was surely correct. He erroneously
gave the origin of his supposedly new species as Jalisco, Mexico, culti-
vated at Cambridge from corms sent by Pringle. Pringle in his diary
(ed. Helen Burns Davis, 1936, p. 65) records a trip made expressly to
collect quantities of the plant from August 9 to 12, 1889. It was obtained
at Pena station, 70 miles east of Laredo, Texas. The plant’s name is
misspelled as Tigridia vaccata in the entry for Aug. 9, but appears
correctly in that for Aug. 11. Its identity with Eustylis purpurea was
noted by Small in 1937 (Addisonia 20: 13). Foster excludes it from
Nemastylis (Contrib. Gray Herb. 155: 44, 1945), leaving it in Eustylis,

d inne Ss
quite satisfactorily to Tigridia. It is the northernmost representative of
that genus, cccurring from extreme southern Texas northeastward to
central Louisiana and extreme southern Arkansas (Union Co.: Norphlet,
A. J. Hoiberg 496, 13 June 1954 (SMU); “open sandy pine woods’).
—Lloyd H. Shinners.

MACHAERANTHERA PINNATIFIDA (HOOKER) SHINNERS,
COMB. NOV. ae ae er Faas ed on Diplopappus pinnatifidus
Am

Sept. 2: 564. ne (“1814”). (Not Machderanthera spinulosa Greene,
1899.) Starkea ? pinnata Nuttall, Gen. 2: 169. 1818. (Illegitimate since
Amellus spinulosus Pursh is given as synonym, but the specific epithet
not adopted.) Machaeranthera pinnata (Nuttall) Shinners, Field & Lab.
18: 41. 1950. Since Hooker’s epithet is both legitimate and available, it
must be adopted in place of Nuttali’s. I am indebted to Dr. Marshall C.
Johnsten for bringing this to my attention. — Lloyd H. Shinners.

REVIEW

ROADSIDE FLOWERS OF TEXAS. Paintings by Mary Motz Wills,
text by Howard S. Irwin. 295 pp. University of Texas Press, Austin.
1961. $5.75

Texas’ botanical wealth has never received the attention it deserves,
aesthetically or scientifically or even commercially. But there are signs
of improvement. This book has a fly-leaf announcement signed by the
President of the University of Texas, and the Humble Oil and Refining
Company recently offered it at a reduced price to credit-card holders.
Like several predecessors (Whitehouse’s Texas Flowers in Natural
Colors, Schulz’s rs. Quillin’s) Texas Wild owers, Casey’s 101
American Wild Flowers), it provides illustrations of a selection of the
many flowering plants found wild in the state. It goes beyond any pre-
decessor in having a text supplied by a botanist who makes a serious
effort to provide notes on a broad sample and includes keys to those
illustrated.

Primary basis for the book is a group of watercolor paintings by Mary
Motz Wills of Abilene. Most of them do not depict whole plants, but
small portions, such as one might have gathered for a miscellaneous
bouquet on a casual walk. The impression of fragmentariness is accentu-
ated by their having been reduced to fit four on a page. The scale varies
greatly, at times to a misleading degree. The huge trumpets of Datura
Wrightii appear little larger than the blossoms of Heliotropium con-
volvulaceum on the facing page, whereas the former are actually nearly
ten times as large as the latter. The medium of watercolor does not lend
itself well to depicting such botanically significant features as hairs
on stem or leaves; the illustration labelled Astragalus mollissimus is
hardly recognizable as that densely hairy plant. But the paintings were
not made with the expectation of publishing them in a book, especially
in association with technical botanical information, so criticism from
such a technical viewpoint is really not fair. The pictures will be quite
serviceable aids to the recognition of many common wild flowers of
Texas, and that after all is the chief purpose of the book.

Following the 64 pages of illustrations are 185 pages of keys and
descriptions, with brief notes on additional species related to those
illustrated, information on distribution within the state, and items of
special interest. Compiling all this for a state which had no complete
published flora and not even an up-to-date checklist was no small task.
When one recalls that the author of the text was at the time only a
graduate student, and a newcomer to Texas to boot, it must be acknowl-
edged a really amazing performance. Three pages of line drawings to
illustrate botanical terms, a glossary, and separate indexes for common
and scientific names conclude the book.

297

In an introductory “Note to the Amateur Botanist,” Dr. Irwin addresses
hopefully “the intelligent lay botanist who is interested in enlarging
his knowledge of the flora around him,” and who is willing to tackle
botanical keys and botanical terminology. If my experience with my
Spring Flora of the Dallas-Fort Worth Area is any indication, he is
addressing some exceedingly rare animals.’ The self-discipline and
patient effort needed for the pursuit of Linnaeus’s “harmless science” are

attractive colored illustrations and a well-done, serious, technically
respectable (but still quite elementary) text should prove a boon.

With no desire whatever to belittle Dr. Irwin’s achievement, it has
to be stated that some of the illustrations cannot be positively identified,
because too incomplete or not showing certain important technical
details, and a number of others have names attached to them which
are definitely not the correct ones. Perhaps the originals are sufficiently
superior to the reproductions to allow more confident identifications.
To my eye, at least, the illustrations designated as Iris hexagona, Mira-
bilis nyctaginea (as nyctagineus), Lepidium virginicum, Brassica juncea,
Draba platycarpa, Astragalus mollissimus, Castilleja latebracteata, C.
indivisa, Plantago Helleri, Liatris punctata, Gutierrezia dracunculoides,
Solidago nemoralis, S. altissima, Aster oblongifolius, A. praealtus (as
prealtus), A. subulatus var. ligulatus, Senecio plattensis, and Pyrrhop-
appus multicaulis are not positively identifiable as those species, and I
am unable to state with certainty what they are. The names listed below
are definitely not correct; when possible I have given what I believe
to be the correct names in capital letters.

PLATE 1. Sagittaria latifolia. S. LONGILOBA. Projecting leaf-bases
distinctly longer than the apical portion.

PLATE 3. Yucca Treculeana. The whole plant at left may be this
species, though the leaves seem too narrow. The portion of inflorescence
at right definitely is not; it may be Y. ARKANSANA or Y. ANGUSTI-

™Not a single book store within the area of the Spring Flora regularly carries it in stock,
and only three have ever ordered it, one of these just one copy in the more than 6 years
since its publication. When we den i of a eae newspaper asked that a copy
purchased for the paper’s reference libr the est was refused. In all athe it
should be mentioned that the Dallas Public oe ry ave purchased a total of 17 copies. But
aca eee for the book in an area making loud claims to cultural ee. is hardly
impressive. Even sales for use in teaching go overwhelmingly to cities outside the area, some
of ae sever a handed miles away. No copy has every been sold for this purpose in

F
Worth, and the only such sales in Dallas in more than a year have been at the high ae
el.

lev

298

FOLIA.—Yucca arkansana. The whole plant at left is probably Y.
PALLIDA or its close relative ¥. RUPICOLA. The portion of inflor-
escence at right I do not recogn

Plate 11. Clematis reticulata. Ce PITCHERI. The former has thick,
st:ff, heavily veiny leaf blades.

PLATE 26. Viola missouriensis. V. VILLOSA. The leaf-blades of V.
missouriensis are triangular-pointed and its flowers are definitely on
the blue side.

PLATE 33. Asclepias oenotheroides. A. LATIFOLIA. A. oenotheroides
has petioled leaves.

PLATE 39. Verbena pumila. V. TENUISECTA. The flowers of V.
pumila are extremely small and are not elevated on naked peduncles

in the manner shown.

PLATE 41. Brazoria scutellarioides. SCUTELLARIA RESINOSA. The
Erazoria is annual, with a very slender taproot, and the flowers are
smaller and paler.

TE 42. Salvia farinacea. S. AZUREA var. GRANDIFLORA. In
S. jarinacea the calyx is white-woolly

PLATE 55. Aster laevis. Not identifiable. This species is northern and
does not occur in or near Texas.

PLATE 56. Aster sagittifolius. A. TEXANUS. The former also is a
northern species not known from Tex

PLATE 59. Thelesperma eee ern Probably HELENIUM BAD-
IUM. The Thelesperma has a rather flat, yellow center to the head.

PLATE 61. Helenium latifolium. H. FLEXUOSUM. The former (usually
referred to H. autumnale) also has heads with yellow center.

PLATE 64. Krigia virginica. Not too accurately depicted but almost
certainly K. OCCIDENTALIS, with shorter pappus in proportion to the
body of the achene (which is partially cbscured by the pappus scales
in the illustration).

In one case there is discord between the common name and the Latin
one: “Old Plainsman” for Hymenopappus artemisiaefolius. This is
mainly an East Texas species, not one of the prairies or plains, though
it has relatives occurring there.—Lloyd H. Shinners.

S| D CONTRIBUTIONS
TO BOTANY

VOLUME 1 NUMBER 6 OCTOBER 1964

CONTENTS

Fewer Florida rarities: changing flora of the Pineola Grotto,
Citrus County. Olga Lakela. 299

Cytological studies in Paspalum, group Setacea (Gramineae).
Donald J. Banks. 306

Chromosome numbers of some North American species of Astragalus
(Leguminosae). G. F. Ledingham and M. D. Fahselt. 313

Yellow-flowered Linum (Linaceae) in Texas. C. M. Rogers. 328

Calylophus (Oenothera in part: Onagraceae) in Texas.

Lloyd H. Shinners. 337
Taxonomy and heterostyly of North American Gelsemium

(Loganiaceae). Wilbur H. Duncan and Donald W. DeJong. 346
Texas Asclepiadaceae other than Asclepias. Lloyd H. Shinners. 358

New or otherwise interesting Coreopsidinae (Compositae) from
northwestern South America. Earl Edward Sherff. 368

New names and records for Texas Compositae. Lloyd H. Shinners. 373

NOTES. Digitaria Ischaemum (Gramineae) in Mississippi and Texas. 380.—Chromosomes of

two Moraea (lridaceae) from eae nae Africa. 381.—Eriogonum annuum (Polygonaceae)
biennial in eu 382.—A deceiving aquatic Neptunia (Leguminosae) in Central
America. .—A hexaploid Linum riers from eastern Ethiopia, 383.—Cayratia alan
(Vitaceae) in southeastern Louisiana: new to the United States. 384.—Three new varietal

names in Sphaeralcea (Malvaceae). | varietal names for New World Ludwigia
(Onagraceae).,
Youngias (‘‘Crepis japonica”: ees introduced in the southern United aise, 386.

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VIEW. — Third New International Dictionary of the English Language, Un-
een 38

INDEX TO VOLUME 1. 393.

SIDA is privately published by Lloyd H. Shinners, SMU Box 473,
Dallas, Texas 75222, U.S.A. Subscription price $6 (U.S.) per volume
of about 360—400 pages, parts issued at irregular intervals.

©)
SIDA Contributions to Botany volume 1 number 6 pages 299—417
copyright 1964
by Lloyd H. Shinners

FEWER FLORIDA RARITIES: CHANGING FLORA
OF PINEOLA GROTTO, CITRUS COUNTY
OLGA LAKELA
University of South Florida, Tampa, Florida 33620

The limestone grotto near Pineola has been searched for ferns ever
since its discovery by A. H. Curtiss in 1881. Roland M. Harper (1916)
graphically recounted this eventful discovery of an unspoiled fern flora,
thitherto unknown to botanists. Curtiss’ discovery of previously unre-
ported species in the United States, aroused interest among his con-
temporary botanists. J. D. Smith, 1883, journeyed to Pineola, finding a
few species not noticed by Curtiss. Harper’s own expedition, 1915, re-
sulted in an annotated list of 12 species, inclusive of all the previously
recorded ones which he observed and 2 of his own finding. The list was
increased to 14 species by J. K. Small (1920A) who journeyed to the area
in 1918.

The physical aspects and the vegetation of the grotto are eloquently
narrated by Curtiss, Harper and Small. In Small’s own words, “we found
ourselves in a veritable amphitheater, surrounding a cypress swamp. On
entering through rocky wall we found ourselves among boulders, chasms,
canons, natural bridge, and caves of eroded limestone. Everything was
partly or completely covered with fern growth of at least a dozen differ-
ent kinds of ferns... an enhancing grove of both conifers and broad-
leaved trees over-shadowed the grotto, altogether a grove and a grotto
that would have been a delight to the devotees of the worship of Baal;
but this sanctuary had already been profaned.... After nature has built
and adorned this grotto beyond power of words to describe, man has
recently started the task of utterly destroying it.” At the edge of the
grotto a mill was pulverizing the rock for commercial liming of culti-
vated fie

he ner site, an extensive pit overgrown with weeds and intro-
duced grasses, remains. Just south of it, the part that escaped utter
destruction, curves its crescentic contour toward Withlacoochee River
marsh. This—the present day Pineola Fern Grotto—features no caves

utmost precaution in maintaining a stable foothold. Boulders, cliffs and
ledges, are all more or less covered with mats of lichens, mosses, ferns
and flowering herbs, beneath high-flung canopy of the hammock tree
association of primeval times. Even this remnant of a natural area is
being despoiled by grazing herds. As a part of a fenced and posted ranch-
land for decades, the natural vegetation along hammock margins and on
the lower rocks of the grotto has been replaced by invading forbs and
local weeds.

ety ag a Botanical Laboratories, University of South Florida. -

6): —305. 1964.

SID+

300

To this grotto, the author, aided by Mr. and Mrs. James A. Lassiter of
Tampa, has made several collecting and study trips during the past two
years. It is hoped that the appended enumeration of species of an area
whose botanical history dates back to 1881, is deemed worthy of placing
on record,

Although the number of species of ferns in the present list is greater
than that of the previous ones, the populations in a given area are less
concentrated and widely dispersed. The rarest of the rare spleenworts,
the species most sought, have become decimated to the verge of extinc-
tion. In 1962 Asplenium abscissum was no longer in evidence; A. x
Curtissti was represented only by three known plants. All the species of
trees previously reported in hammock associations have been observed
with exception of Acer rubrum, Persea borbonia and Cornus florida.
They are frequently encountered off the immediate area; it is assumed
that the record refers to the destroyed hammock ae In addi-
tion, vigorous and prolific Sapindus marginatus has been observed in
various stages of growt

loss of a natural area in the keeping of man is regrettable. Even
after decades of misuse, if the causes of destruction could be removed,
restoration of the grotto by nature’s creative forces lies within possibility.
The Pineola Fern Grotto is worthy of preservation.
PSILOTACEAE
Psilotum nudum (L.) Beauv. Sight record of 1 plant; epiphyte.
HIZAEACEAE
Anemia adiantifolia (L.) Sw. Sight record of 1 sterile colony, on low
rocks

OPHIOGLOSSACEAE
Botrychium dissectum Spreng. var. tenuifolium (Underw.) Farw. 26404.
ew plants noted on a rocky margin of hammock.
OSMUNDACEAE
Osmunda regalis L. 26416. A sterile plant noted in margin of swamp,
Withlacoochee River, One leaf collected.
YPODIACEAE
Adiantum tenerum Sw. 25462. Occasio
Asplenium abscissum Willd, 20468. a noted since 1962. Apparently
exterminated.
A. X Curtissti Underw. 25470. Almost completely decimated.
A. heterochroum Kunze, 25468. Very rare.
A. resiliens Kunze, 26812. Rare, low rocks.
verecundum Chapm. 26021. Rare
Dryopteris ludoviciana (Kunze) Small. 26043. Vigorous growth.
Polypodium dispersum ined. 26808. Rare.
P. pectinatum L. 26499. Rare.
P. plumula HBK. 26446. Rare.
P. polypodioides (L.) Watt. Sight record, epiphytic.

S

Pteris cretica L, 27395. Two plants noted. Rare.

P, vittata L, 25469. Local; rare.

Tectaria heracleifolia os Underw. 25460. Shady limestone slopes;
young ferns noted; e.

Thelypteris dentata ae E. St. John, 25471. Rare.

T. normalis (C. Chr.) Mox. 25476. Occasional.

T. reptans (J. F. Gmel.) Morton. 25461. Frequent on shady sloping lime-

stone.
T. Torresiana (Gaud.) Alston. 25467. (T. setigera (Blume) Kuntze).
T. tetragona (Sw.) Small, 26394. Rare.
TAXODIACEAE
Taxodium distichum (L.) Rich. Sight record. The swamp-facing side of
the grotto.

MINEAE

Oplismenus setarius (Lam.) R. & §. 25455. Frequent in thin soil over
limestone

Panicum anceps Michx. 25456. Few tufts noted, in gravelly soil of grotto
margin.

P. joorii Vasey, 26032. Few plants in vernal phase; among marginal
grasses.

Paspalum conjugatum Berg. 26443. Among grasses, escaped from cultiva-

ang rn. 393.
rite sessiliflora Poir. 26444, Few plants with wood ferns, grotto bottom.
PALMAE
Rhapidophyllum Hystrix (Fraser) Wendl. 25443. Few over rocks.
Sabal minor (Jacq.) Pers. 26421. Marginal area of grotto facing the river.
Serenoa repens (Bartr.) Small, sight record; margin of hammock.

ACEAE
Arisaema Dracontium (L.) Schott, 26423A. Grotto bottom between rock
walls. Occasional.
ROMELIACEAE

Tillandsia simulata Small. 26050. Occasional.
T. usneoides L. Sight record. Frequent

COMMELINACEAE
Callisia cordifolia (Sw.) Anders. & Woods. 25473. Over low mossy rocks.

LILIACEAE

Smilax auriculata Walt. 26053. Frequent.

AMARYLLIDACEAE
Crinum americanum L. 26420. Soft soil of grotto margin, facing the river

IRIDACEAE
Sisyrinchium rosulatum Bicknell. 26048. Crevices of low rocks; uncom-
mon,

302
ORCHIDACEAE
Habenaria quinqueseta (Michx.) Sw. 26389. Several plants along fence
of pasture adjoining the grotto hammock.
PIPERACEAE
Peperomia leptostachya (Nutt.) Chapm. 26392. Thriving colony over
crumpled lichen-coated rocks.
CORYLACEAE
Carpinus caroliniana Walt. Frequent; sight record.
FAGACEAE
Quercus Michauxii Nutt. 26040. Very large trees in grotto hammock;
specimen from fallen branch,
Q. Shumardii Buckl. 26391. Spee. from fallen branch. Very large trees
over grotto.
Q. virginiana Mill. Sight record. Very large trees with expansive crowns
over grotto and in hammock.
ULMACEAE
Celtis laevigata Willd. 26027. Embankment of the excavated grotto site.
Ulmus floridana Chapm. Sight record. Large trees in hammock and
margin of grotto.

URTICACEAE
Boehmeria cylindrica (L.) Sw. 25925, Frequent in grotto.
Parietaria floridana Nutt. 25933. Frequent in moist shady hollows of
grotto.
Urtica chamdedryoides Pursh, 25452. Persisting in rock clefts, shady
moist soil.

PHYTOLACCACEAE
Petiveria alliacea L. 25453. Few plants in shade over grotto rocks.
Rivina humilis L. 26034. Slender shrubs in rock clefts. Infrequent.
YLLACEAE
Arenaria lanuginosa (Michx.) Rohrb, 25474. Local. Stems trailing from
Drymaria cordata (L.) Willd. Sight record. Low moist hollows with
Stellaria.
Stellaria media (L.) Cyrill. Low moist clefts, floor of grotto. Anthers 5,
red.
RANUNCULACEAE
Clematis reticulata Walt. 26041, One vine noted base of grotto wall.
HAMAMELIDACEAE
Liquidambar Styraciflua L. 26051. Frequent over rocks and in hammock.
ACEAE
Decumaria barbara L. 26396. Stems creeping in mosses and lichen at
grotto walls, and twining high on tallest trees. Local, abundant.
ROSACEAE
Rubus trivialis Michx. Sight record; on low rocks of grotto.

303

LEGUMINOSAE

Amphicarpa bracteata (L.) Fern. 26422. In soil-filled clefts of low rocks;
flowers bluish-lavender; uncommon.

Cassia occidentalis L. 25441. Oak and palm grove adjoining grotto ter-
race. Infrequent.

Desmodium cuspidatum (Muhl.) Loud. 1 plant noted.

D. paniculatum (L.) DC. 26386. In shade over low rocks, and swamp
margin adjoining the grotto.

Galactia Macreei M. A. Curtis. 26036. Grassy margin of trail to grotto
entry.

RUTACEAE

Citrus aurantium L. 26806. Several trees noted; eastern part of hammock.

Zanthoxylum Fagara (L.) Sarg. 26409. Small, sterile shrubs; few on high
rocks.

MELIACEAE
Melia Azedarach L. 26027. Few trees on the rim of the old pit.
ANACARDIACEAE
Rhus Toxicodendron L. Sight record. Fairly general in grotto floor.
ACERACEAE

Acer Negundo L. 25916, Frequent over grotto rocks; trees large with full,
leafy crown.
SAPINDACEAE
Sapindus marginatus Willd. 26039. Frequent over rocks in hammock.
RHAMNACEAE
Sageretia minutiflora (Michx.) Trel. 26023. Young shrubs at base of rock
wall

VITACEAE
Parthenocissus quinquefolia (L.) Planch. 26026. High climbing over tree
on rim of the old pit.
Vitis rotundifolia Michx. High climbing; in oaks. Sight record.
TILIACEAE
Tilia floridana Small. 26045. Large tree, margin of grotto hammock, on
trail.

Pavonia spinifex (L.) Cav. 25475. Locally frequent over grotto on rocks.
VIOLACEAE
Viola floridana Brainerd. 26803. In moist soil of shady hammock and
grotto rocks.
LYTHRACEAE
Decodon verticillatus (L.) Ell. 26441. Shallow water on cypress pond,
margin of grotto hammock.
ACEAE
Nyssa biflora Walt. 26054. Young shrubs in abundant fruit, swampy tract
adjoining the grotto hammock.

304

MELASTOMACEAE

Rhexia mariana L. var. exalbida Michx. 26047. Swamp margin with

Decodon

UMBELLIFERAE

Ptilimnium capillaceum (Michx.) Raf. 25923. In moist rock clefts of

grotto.
Sanicula canadensis L. 25934. Common on grotto rocks.

RIMULACEAE

Samolus parviflorus Raf. 25922. Wet soil of swamp, grotto margin. Few.

LEPIADACEAE
Cynanchum palustre (Pursh) Heller. 25928. Frequent over grotto trees.
CONVOLVULACEAE

Ipomoea trichocarpa Ell, 26438. Luxuriant colony in grotto and ham-
mock margin.
VERBENACEAE
Callicarpa americana L. 26025. Occasional. Plants rooted in clefts of rocks.

I
Hyptis mutabilis (A. Rich.) Briq. 26024. Few plants noted in grassy area.
Leonotis nepetaefolia R. Br. 26423. Open shade of Carya along trail to

Salvia coccinea Juss. 26022. Frequent in grotto over rocks.
S. lyrata L. 26804. Rosettes only noted. Openings of vegetation along
trail,
SOLANACEAE
Capsicum frutescens L. 25447. One small shrub in abundant fruit, in
grotto.
Solanum floridanum Shuttlw. 26415. Occasional in rock clefts.
SCROPHULARIACEAE
Bacopa coroliniana (Walt.) Robins. 26388. Shallow water of cypress
pond, adjoining grotto hammock.
BIGNONIACEAE
Campsis radicans (L.) Seem. Sight record.

Dicliptera assurgens (L.) Kuntze. 26439. Over high rocks, scattered. In-
frequent.

Dyschoriste humistrata (Michx.) Kuntze. 25919. Wet soil of swamp
order along grotto

Ruellia carolinensis (Walt.) Steud. 25918. Frequent, in thin soil of rock
ledges.

RUBIACEAE

Galium pilosum Ait. var. laevicaule (Weath.) Blake. 25932. Occasional
among shady boulders.

Mitchella repens L. 25926. Few plants noted, among ferns.

Psychotria nervosa Sw. 26037. Occasional in rocky holes and on top of
rocks,

305

VALERIANACEAE
Valeriana scandens L. 25924. Well established in low fissures of lime-

stone
CUCURBITACEAE
Melothria pendula L. 26922. Frequent over grotto herbs and grasses.
AMPANUL E
Lobelia homophylla F. E. Wimmer. 26319. Small colony in shade, low
rocks.
COMPOSITAE

Ambrosia elatior L. 26319. Weedy.

Aster pinifolius Alexander, 26400.

Bidens bipinnata L. 25921. Weedy throughout low rocks.

Bidens pilosa L. 26399. Few plants noted, in grassy areas, among rocks.

Cirsium sp. Sight record of rosettes only

Bilephantopis carolinianus Willd. 25459. Well established in under-
growth.

Eupatorium coelestinum (L.) DC. 26031. One colony no

Haplopappus divaricatus (Nutt.) Gray. 26397. Open ranchland adjoining
the grotto

Pyrrhopappus carolinianus (Walt.) DC. 26030. One plant noted in ham-
mock adjoining grotto.

REFERENCES
HARPER, R. M. 1916. Fern grottoes of Citrus County, Florida. American Fern Journal
» pl. J.
SMALL, _ K. 1920A. A journey to the fern grottoes. Journ. N. Y. Bot. Gard. 21 (242):
p. 45.
20B. Of grottoes and ancient dunes. Journ. N. Y. Bot. Gard. 21 (243):

45—52, pl. 244. aes y the plate applies to Pineola.)
TT . Ferns of the Southeastern States.

CYTOLOGICAL STUDIES IN PASPALUM,
GROUP SETACEA (GRAMINEAE) '
DONALD J. BANKS
Stephen F, Austin State College, Nacogdoches, Texas

The Setacea group of the genus Paspalum consists of a network of
closely related taxa which are taxonomically difficult. Chase (1929)
recognized 10 species in the group but acknowledged that they are poorly
defined and appear to intergrade. This study was made to obtain infor-
mation to be used in a toxonomic revision of the group. The names used
for the taxa are according to Chase’s concept. My concept of the taxa is
to be published in another paper.

CHROMOSOMES

Immature inflorescences were collected in the field or from greenhouse
transplants, killed in a 3:1 alcohol-glacial acetic acid solution, and the
anthers squashed in aceto-carmine. The chromosome counts were usually
made at diakinesis in pollen mother cells. Photomicrographs were made

of microsporocytes with chromosomes distributed so that they were
countable. Some slides representing each taxon were made permanent
by McClintock’s (1929) method. Voucher specimens are deposited in the
University of Georgia and the Stephen F. Austin State College herbaria.

A summary of chromosome counts made during this study and by
previous investigators is given in Table 1. In all the plants that I studied
the microspores contained 10 chromosomes and meiosis appeared normal.
Differences in chromosome size and morphology within or between taxa
were slight. Drawings made by tracing photomicrographs of chromo-
somes of each species (sensu Chase, 1929) are shown in Figures 1 to 10.

time. Somatic counts previously reported for P. ciliatifolium Michx.,
P. pubescens Muhl., P. setaceum Michx., and P. supinum Bosc varied
from 20 to 80 with different counts for the same species (see Table 1).
Darlington and Wylie (1955) reported the basic chromosome numbers
10 and 12 for Paspalum. Forbes and Burton (1961) suggested the base
number 10 may have been derived from some lower ancestral base
number, possibly 5 or 6, because of strong secondary associations of
bivalents noted in their investigations with P. almum Chase. Some of

1 Contribution No. 58 = the Stephen F. Austin State College — of Biology
This paper is based partly on a aos submitted to the Graduate Faculty the Oa
versity of Georgia. The ees has been aided by two National Scienc toca tion
Summer Fe lowships for Graduate Teaching Assistants, an NSF eee nares
Project for College Teachers at the University of Texas, Department - Botany, and a facul-
ty research grant, Stephen F. Austin State College. * oe are bo a . Wilbur H. Duncan,
who served as my major professor during portions of this study.

SIDA. 7 (6): 306-312. 1964

307

my preparations suggested secondary associations. Additional studies are
required to confirm this suggestion. This study disclosed no chromosomal
data that I consider to be useful in distinguishing between the Setacea
taxa.

EMBRYO SAC DEVELOPMENT

A better concept of the taxa might be possible if the mode of reproduc-
tion could be determined. Apomixis, which occurs in some Paspalum
species, was suspected in Setacea because of morphological uniformity
within its component taxa where they grow sympatrically and because
of similarity of progeny to the maternal parents in some progeny tests.
The following study was conducted to determine whether or not the
Setacea taxa reproduce by apomixis.

Immature inflorescences collected in the field or greenhouse were
killed and fixed in a 3:1 absolute alcohol-glacial acetic acid solution.
The material was stored in 70% alcohol at 5°C until dehydration was
begun. Dehydration was completed with a tertiary butyl alcohol series.
The material was infiltrated with paraffin, sectioned at thicknesses of
15 to 17 microns with a rotary microtome, and stained with safranin-fast
green. Twelve to twenty spikelets per plant were sectioned. Twenty-two
plants representing the ten taxa were studied. The slides were examined
microscopically to ascertain whether or not multiple embryo sacs were
present.

None of the material studied showed more than one embryo sac.
Embyro sac development in Setacea is the Polygonum type except the
antipodals usually form several cells rather than three.

pomixis was reported in Paspalum by Burton (1948), Smith (1948),
Bashaw and Holt (1958), Brown and Emery (1958), Forbes (1960), and
Snyder (1957, 1961). Brown and Emery (1958) reported normal embryo
sac development for P. pubescens, the only Setacea Paspalum which
apparently had been investigated prior to this study.

The type of apomixis detected in Paspalum thus far is somatic apos-
pory followed by pseudogamy. Usually one or more nucellar cells adja-
cent to the megaspore mother cell begins to enlarge and one of the cells
usually develops into a functional embryo sac. A nucellar embryo sac
may consist of an egg, two synergids, two polars, and several antipodals
as in P. secans Hitche. and Chase (Snyder, 1957) or the synergids and
antipodals may be absent as in P. dilatatum Poir. (Bashaw and Holt,
1958). Fertilization of the polar nuclei is believed to be necessary for
the development of endosperm, but the embryo develops from an un-
reduced, unfertilized egg.

e failure to detect multiple embryo sacs in Setacea suggest that
they reproduce sexually, although stages representing actual fertiliza-
tion of the egg or polars were not observed. The formation of a single
embryo sac by generative apospory, however, is not precluded by my
data; but since the chromosome studies indicated meiosis to be normal,

308

and since the taxa appear to be diploid, sexual reproduction, rather than
apomixis, seems likely. If apomixis is dismissed as a possible mode of
reproduction, the similarities of offspring to maternal parents in some
progeny tests are best explained by suggesting self-fertilization as the
usual method of reproduction. Further investigations are needed to estab-
lish the method of reproduction.

REFERENCES
BASHAW, FE. C. AND E. C. HOLT. 1958. Megasporogenesis, embryo sac development,
and embryogenesis in Ree pour eae Poir. Agron. Jour. 50: 753-756.
_— WN, W. V. . A cytological study in the Gramineae. Am. Jour. Bot. 35: 382-

————— AND W. H. P. EMERY. 1958. Apomixis in the Gramineae: Panicoideae.
Am. as Bot. 45: 253-263
3URTO

I N, . W. 1940. A cytological study of some species in the genus Paspalum. Jour.
Agr. Res. 60: - 198,
————. 1942. A cytological study of some species in the tribe Paniceae. Am.

Jour. Bot. 29: 355-359,

——————.. 1948. The method of reproduction in common bahiagrass, Paspalum no-
fatum., Agron. Jour. 40: 443-452.

CHASE, A. 1929. The North American species of Paspaluir. Contr. U. S. Natl. Herb.

28(1): 1-310.
CHURCH, G. L. 1929. eae Les in certain Gramineae. Bot. Gaz -84,
DARLINGTON, C. D. AND . WYLIE. 1955. Chromosome atlas of flowering “planes:

L a4
FORBES, L, Jr. 1960. A rapid enzyme- smear technique for the detection and study of
plural embryo sacs in mature ovaries in several Paspalum Pee es. Agron. Jour. 52: 300-301.
——————— AND G. W. BURTON. 1961. Cytology ae iploids, natural and induced
tetraploids and intraspecies hybrids of bahiagrass, ase ee Fligge. Crop Sci.
02-406.
Sica FLW. 1958. C

—

Aromosome numbers in southwestern grasses. Am. Jour. Bot. 45:
757- :
Cc evan: N. 1940. Untersuchungen zur Cytologee und Systematik der Grami-
neen. Beith Bot. Centralbl. : 1-56.

McCLINTOCK, B. 1929. A method for making acetocarmine smears permanent. Stain
a 4: 53-56,

AURA, F. 1941. Cariologia de algun as i ay 2 genera Paspalum. Facultad de agro-

nomica Y veterinaria. Institute de genetica. 41-

SMITH, B.S. 1948. Hybridity ma apomixis in ss perennial grass, Paspalum dilatatune.
Genetics. ne 628-629.

SNYDER, L. A. 1957. Apomixis in Paspalum secans. Am. Jour. Bot. 44: 318-324

ee, 1961 Asyndesis and meiotic non-reduction in microsporogenesis of apo-
mictic Paspalum secans. Cytologia. 26: 50-61.

4 ye
| ee |

Fig. 1-10. Chromosomes of Setacea species (sensu Chase, 1929) at diaki-
nesis.—Fig. 1. P. ciliatifolium, Banks 1339.—Fig. 2. P. debile, Banks
1625.—Fig. 3. P. longepedunculatum, Banks 891.—Fig. 4. P. propinquum,
Banks 1726.—Fig. 5. P. psammophilum, Banks 1507.—Fig. 6. P. pubescens,
Banks 1525.—Fig. 7. P. rigidifolium, Banks 1314.—Fig. 8. P. setaceum,
Banks 1451.—Fig. 9. P. stramineum, Dwyer 16 Aug. 1961.—Fig. 10. P.
supinum, Banks 906.

TABLE 1

SUMMARY OF CHROMOSOME NUMBERS OF THE
SETACEA PASPALUMS

Species Plants examined for this study!
(Sensu Chase, 1929) Gametic Somatic Investigator All gametic chromosome numbers were 10
ciliatifolium? 20 Burton (1940) L-65, Glynn Co., Ga.; 933, 934, Jefferson Co., Fla.;
20 Brown (1948) 954, Taylor Co., Fla.; 964, Alachua Co., Fla.; 984,
Wheeler Co., Ga.; 1339, Columbia Co., Fla.; 1412,
Cooke Co., Tex.; 1413, Warren Co., Miss.; 1570,
Taylor Co., Ga.; 1630, Santa Rosa Co. Fla.; W. H.
Duncan 21888, iennos Co, Plas Dd, G. Randolph
R-2, Jack Co., Tex.
debile 832, Santa Rosa Co., Fla.; 912, Leon Co., Fla.; 1160,

Hardee Co., Fla.; 1440, Camden Co., N. C.; 1616A,
1619, 1625, Santa Rosa Co., Fla.; 1713, Bay Co.,
Fla.; 1747, Levy Co., Fla.; 1885, Travis Co., Tex.;
3670, Brooks Co., Tex.

‘Numbers are my collections except where otherwise indicated.
* Darlington and Wylie (1955) listed P. aN without author, as 2n=80, as determined by Saura (1941). P. epile Nash is a synonym
of P. ciliatifolium., The plant determined by Saura was P. epile Parodi (P. parodianum Hennr.) and is not synonymous with P. atten.

OLE

TABLE 1 (Continued)

Species
(Sensu Chase, 1929) Gametic Somatic Investigator

Plants examined for this study!
All gametic chromosome numbers were 10

longepedunculatum

891, Bay Co., Fla.; 1145, 1146, Lake Co., Fla.; W. H.
Duncan, 21844, Collier Co., Fla.

propinquum

1726, Taylor Co., Fla.; 1733, Dixie Co., Fla.

psammophilum

1459, Burlington Co., N. J.; 1507, 1516, Camden
Co., N. J.

10 Church (1929)

pubescens?
10 Gould (1958)

L-25, L-31, L-35, Laurens Co., Ga.; 974, Putnam
Co., Fla.; 994, Oglethorpe Co., Ga.; 1017, Clarke
Co., Ga.; 1437, Martin Co., N. C.; 1439, Camden Co.,
N. C.; 1452, Gloucester Co., N. J.; 1525, Dinwiddie
Co., Va.; 1531, Nash Co., N. C.; 1898, Nacogdoches

Co,,

rigidifolium

1158, Polk Co., Fla.; 1314, Marion Co., Fla.; 1746,
Levy Co., Fla.; 1773, Pinellas Co., Fla.

3 Brown (1948) reported 2
of Texas, is really P. longipilum Nash.

n=60 for P. pubescens. His voucher specimen (2605), which I examined in the Herbarium of the University

w
_
lanl

TABLE 1 (Continued)

Species
(Sensu Chase, 1929) Gametic

Somatic Investigator

Plants examined for this study!
All gametic chromosome numbers were 10

setaceum

L-19, Wilkinson Co., Ga.; L-24, Laurens Co., Ga.;

L-146, Taylor Co., Ga.; 770, Pike Co., Ala.; 870,
Okaloosa Co., Fla.; 908, Leon Co., Fla.; 1430, On-
slow Co., N. C.; 1451, Gloucester Co., N. J.; 1460,
Burlington Co., N. J.; 1524, Dinwiddie Co., Va.;
1536, Baldwin Co., Ga.; 1682, Baldwin Co., Ala.;
1790, Putnam Co., Fla.

stramineum

1411, 1815, Payne Co., Okla.; 1886, 1889, Travis Co.,
Tex.; 1895, Burnet Co., Tex.; D. Dwyer Aug. 16,
1961, Payne Co., Okla.

supinum

50 Kirshnaswamy
(1940)

40 Brown (1948)

20 Burton (1942)

906, Leon Co., Fla.; 953, Taylor Co., Fla.; 1620,
iS)

anta Rosa Co., Fla.; 1668 Jackson Co., Miss.

rats

CHROMOSOME NUMBERS OF
SOME NORTH AMERICAN SPECIES OF
ASTRAGALUS (LEGUMINOSAE)

G. F. LEDINGHAM and M. D. FAHSELT

University of Saskatchewan, Regina Campus, Regina, Saskatchewan

Astragalus, with nearly two thousand species, is one of the largest
genera of the flowering plants. It is widespread in both Old and New
World but it is most abundant in the northern hemisphere. In the Old
World, where there are some 1,600 species, the greatest number of taxa
occur in southwest Asia with gradual decrease westward around the
Mediterranean in both Europe and Africa and north and east through
Asia to the Bering Strait. In the New World there are some 300 species
concentrated in the western United States with a few extending into
Mexico or into Canada and Alaska. There are no species in tropical
America but there are about ninety species in the high western parts of
South America.

Ledingham (1960) speculates that Astragalus is at least biphyletic
since New and Old World species have different chromosome numbers.
Old World species have a basic haploid chromosome number of eight
(Senn, 1938; Darlington and Wylie, 1955, Love and Love, 1961) and show
a high percentage of polyploidy. New World species have haploid chromo-
some numbers of 11, 12 or 13 (Vilkomerson, 1943; Head, 1957; Turner,
1956) and show less than one per cent of polyploidy. The nine South
American species of Astragalus so far counted have either 11 or 13 as
gametophyte chromosome number (Ledingham, 1960) so they seem to be
a part of the New World phylogenetic line.

Since some species of Astragalus and Oxytropis are circumpolar, oc-
curring in both New and Old World (Yurtsev, 1963), it would seem that
the geographic barrier in the Bering Strait region is relatively recent.
The circumpolar species are all 8-chromosome species or polyploids of
these so evidently it was the Old World group which was able to use
the migration route and invade the New World. There are no known
cases of 11- or 12-chromosome species which have spread into the Old
World. The decreasing abundance of 8-chromosome species as one moves
from Alaska south and east supports the idea that a number of Old
World species entered the New World by this route. Some of these
naturally spread farther than others.

The North and South American species of Astragalus are now sepa-
rated by a wide tropical zone which acts as an effective geographic
barrier. The species are closely related but none of them occurs in both
regions (Johnston, 1947). The absence of common species would indicate
that the barrier has been present relatively longer than the barrier in

SIDA 1 (6): 313—327. 1964.

314

the Bering Strait region, giving time for the evolution of different forms.
The fact that there still is considerable morphological and chromosome
number similarity would indicate that Astragalus species in North and
South America, though separated for a considerable length of time, are
actually a part of the same phylogenetic line and they are more closely
related than New World species are to the 8-chromosome Old World
species.

Since Old World species of Astragalus and Oxytropis, including a few

species have n=11, 12 or 13, they must be two different phylogenetic
lines (Ledingham and Rever, 1963). If the divergence occurred after the
origin of the genus then we may still find some species with chromosome
numbers which will explain how the evolution took place. If the diver-
gence occurred before the evolution of the ancestral Astragalus then
the explanation of the relationship may be found in other genera of
the Leguminosae. Turner and Fearing (1959) suggest that there was a
split in the Caesalpinoideae giving evolutionary lines with higher or
lower chromosome numbers and that these provided two origins for
species and genera of the Papilionoideae. If this theory proves tenable
then the Papilionoideae, and Astragalus in particular, provide a remark-
able example of parallel evolution, and the origin of the divergence otf
the two phylogenetic lines of Astragalus would have to be sought in
the Caesalpinoideae. This hypothesis seems improbable.

Ledingham (1960), finding that Astragalus somalensis Taub. ex Harms

flowering or fruiting material we germinated more seeds. The material
had been obtained from the Grassland Research Station, Kitale, Kenya,
East Africa. The count 2n=20 was confirmed (Ledingham and Rever,
1963) and the voucher plants were watched as they grew. The leaflets
were conspicuously veiny and the arrangement of the leaflets in the bud
was not like other Astragalus species we had seen. We were not surprised
when Gillett (1963) revised this species, along with two others from
East Africa, and placed them in the genus Galeg

This paper reports 69 counts on 49 species including 28 species and

that the chromosome meee in this complex of species is stable an

Astragalus belong in different phylogenetic lines.
MATERIALS AND METHODS
The species reported in this paper, except for five Old World species
which came as seed from botanical gardens, were collected in their

315

native habitats by people interested in the Leguminosae. The specimens,
most of which are available in the University of Saskatchewan, Regina
Campus herbarium, have in many cases been identified by Mr. R. C.
Barneby, New York Botanical Garden. Mature seeds were removed and
germinated in petri dishes to give root tips for chromosome counts. Some
seeds have also been germinated to give additional study material and
voucher specimens. Since these species are mostly slow growing peren-

treated with 8-hydroxyquinoline before fixation and staining. The
standard procedure for the Feulgen stain was used in making the root
tips squashes.
RESULTS

Chromosome counts for species of Astragalus and Oxytropis studied
here during the summer of 1963 are presented in Tables I and II. Table I
reports somatic chromosome numbers (22, 24 and in two cases 26) and
collection data for 39 collections representing 28 species of New World
Astragalus. Table II reports somatic chromosome numbers (16, 32, 48
and 42) for 30 collections (15 of which were made in North America) of
21 Old World species of Astragalus and Oxytropis.

The typical New World species of Astragalus listed in Table I were
collected in their natural habitats in the western parts of United States,
except for one species which was collected in Canada. There are voucher

Saskatchewan, Regina Campus, for each of the collections. One collec-
tion. A. coccineus, is represented only by fruits and a single dried
flower, but the long crimson petals provide convincing evidence of the
identity of the species. Some of the collections were badly parasitized
and contained only a few viable seeds so that a sufficient number of
good counts could not always be made; the chromosome number is
reported in four cases with some reservation. Examination of Fig. 12,
A. calycosus, 2n=22, shows that it is not easy to be sure whether the
count is 2n=22 or 24. A quick count of this group may give 24, for there
is a conspicuous nonstaining area, eg. lowest chromosome on the left, in

johannis-howellu. Fig. 9, A. whitneyi, 2n=22, also shows this appear-
ance of an extra pair of small chromosomes which may lead to some
disagreement between counts of New World species of Astragalus.

Three species counts of 2n=24 confirm previous counts. A. kentrophyta,
2n—24 was reported by Ledingham (1960), but that report gives the
wrong author for the species. A. preussii, 2n=24, was reported in 1943
by Vilkomerson. The earlier reports on these two species did not identify
the variety counted. A. spatulalus, 2n=—24, was reported by Ledingham
(1957)

TABLE I. SOMATIC CHROMOSOME COUNTS FOR NEW WORLD ASTRAGALI

Species Seed no, Collection no. Origin 2nchr.no. Fig.
A. amphioxys Gray
var. amphioxys 6424 Rever 72 Arches Nat. Monument, Utah 22 2
fe 6439 Rever 67 Crystal Geyser, Utah 22
A. argophyllus Nutt.
var. martini Jones 6430 Rever 54 Castlegate, Utah 22
A. calycosus Torr. 6402 DeDecker 1522 White Mts. California 22 12
A. ceramicus Sheld.
var. ceramicus Sheld. 6304 Barneby 13,121 Escalante, Utah 22
var. imperfectus Sheld., 6455 Porter 3954 Gillette, Wyoming 22 7
A. coccineus Bdg. 6386 DeDecker 22/7/62 Inyo Mts., California 22 6
A. cymboides Jones 6429 Rever 58 Wellington, Utah 24 10
” 6453 Rever 83 Huntington, Utah 24
A. desperatus Jones
var. desperatus Jones 6418 Rever 78 oab, Utah 24 1
» 6423 Rever 71 Arches Nat. Monument, Utah 24
A. eastwoodae Jones 6299 Barneby 13,064 Gypsum Gap, Colorado 26
A. flavus Nutt.
var. flavus 6416 Rever 81 Moab, Utah 26

OTE

bob

Sd

>

. inyoensis Sheld.
. johannis-howellii Barn,

. kentrophyta Gray

var. coloradensis Jones

. lentiginosus Doug.

var. palans Jones

var. palans Jones

var. fremontii (Gray) Wats.
. miguelensis Greene

. miser Dougl. var.

serotinus (Gray) Barn.

. mollissimus Torr. var.

thompsonae (Wats.) Barn.

. nuttallianus DC. var.

micranthiformis Barn.

. oophorus Wats.

. plattensis Nutt.

6403
6404

6308

6499

6417
6426

6302
6390
6289

* These species counted previously; details in text.

DeDecker 1519
DeDecker 1505

Barneby 13,115

Rever 73
Rever 76
DeDecker 1484

Raven 18,012

Turner 11,180

Rever 80
Rever 75

Barneby 13,110
DeDecker 488
Porter 8385

Inyo Mts., California

Crowley Lake, California

Glen Canyon City, Utah

Arches Nat. Monument, Utah
Moab, Utah
Santa Rita Flat, California

San Clemente Island, California

Pocahontas, Alberta

Moab, Utah
Devil’s Garden, Utah

Coconino Co., Arizona
Badger Flat, California

Cook Co., Wyoming

LTE

. preussit Gray

var. preussii 6419 Rever 66 Green River, Utah
7 6428 Rever 79 Moab, Utah
"A. purshii Dougl. var.
glareosus (Dougl.) Barn. 6414 Rever 111 Boise, Idaho
i 6440 Rever 112 Boise, Idaho
lectulus Jones 6406 DeDecker 1,525 Coyote Ridge, California
longilobus Jones 6393 DeDecker 1,472 Harkless Flat, California
A. ravenii Barn. 6394 DeDecker 1,112 Sawmill Pass, California
A. serenot (Kuntze) Sheld. 6389 DeDecker 1,469 Harkless Flat, California
A. shortianus Nutt. 6285 Porter 8,460 Albany Co., Wyoming
“A. spatulatus Sheld. 6307 Barneby 13,234 Biddle, Montana
A. utahensis (Torr.) T.&G. 6457 Rever 35 Pocatello, Idaho
A. whitneyi Gray 6407 DeDecker 1,535 Coyote Flat, California

A. zionis Jones 6298 Barneby 13,111 Coconino Co., Arizona

* These species counted previously; details in text.

8Té

319

The previous report of chromosome number of A. miser var. serotinus
gives 2n=24 (Ledingham, 1958). Since our 1963 study shows 2n=22, we
suspect that our original report is in error. Ledingham (1960) reported

n=22 for A. nuttallianus var. nuttallianus but the present study shows
2n=24 for var. micranthiformis. The varieties of A. nuttallianus are both
quick growing annuals and it is possible that both counts are correct.

In Astragalus lentiginosus there have been previous reports of 2n=22
reported for var. palans (Vilkomerson, 1943), var. lentiginosus (Head,
1957) and var. variabilis (Ledingham, 1960). This paper confirms the
previous count of var. palans (two collections) and reports 2n=22 for
var. fremontii.

In A. purshii, Head (1957) reported counts of 2n=22 for var. purshii
and var. glareosus. This paper confirms the count for var. glareosus and
reports 2n=22 for var. lectulws and var. longilobus. The chromosome
number seems stable for the many varieties of the highly variable
species A. purshii and lentiginosus.

The remaining species, for which there was a previous chromosome
count, is A. mollissimus. The present report for var. thompsonae agrees
with Ledingham’s (1960) report for var. earlei. Previously Head (1957)
had reported 2n=24 for var. earlei.

No attempt was made to obtain Old World material for the 1963
studies but some seed samples did become available and routine chromo-
some counts were made. These counts are reported in Table II even
though voucher specimens are not available for every collection. EKac é
these Old World species, except one, obviously belongs to an 8-chromo-
some series in which diploids are common but tetraploids and hexaploids
frequently occur. Different chromosome numbers may be present in the
same species. Table II includes some species of Old World Astragalus
and Oxytropis which have migrated into northwestern North America.
It should be pointed out that eleven of the fifteen North American col-
lections reported in Table II were made in Canada or Alaska whereas in
Table I only one of the 39 collections was made north of the Canada-
U.S.A. border.

The chromosome numbers of 13 of the 21 taxa listed in Table II have
been reported before. In all cases except two the present report agrees
with previous reports and details need not be given again here. Astragalus
odoratus was previously reported as having 2n=—64 by Ledingham (1960).
When this material was finally grown out it proved to be A. cicer. The
seed packet had been wrongly identified and Ledingham had made his
report before verifying the identity of his material. A hamosus) is re-
ported (Darlington and Wylie, 1955) as having 2n=48. We are con-
vineed that this number is not correct but we found these chromosomes
very difficult to separate and count and give a tentative count of 2n=ca

Table II gives chromosome counts of four samples of A. alpinus, two
identified as var. brunetianus. The collection from Churchill, Manitoba

TABLE II. SOMATIC CHROMOSOME COUNTS FOR OLD WORLD ASTRAGALI
Species Seed no. Collection no. Origin 2nchr.no. Fig.
*A. alpinus L. 6230 cultivated Acad. Science, Leningrad 16
" 6243b Churchill, Manitoba 32
var. brunetianus Fern. 6399 A. Dechamplain Rimouski, Quebec 16
2/7/58
” 6400 Dutilly & Lepage Missinaibi, Ontario 16
30/7/58
A. angustifolius Lam. 6442 O. Tosun Ankara, Turkey via USDA. 32 16
*A. eucosmus Robins 6396 J. G. Dickson Big Delta, Alaska 32
11/8/56
*4A. hamosus L. 6002 V. Tackholm ca 42
a 6273 N. Feinbrun Peleponese, Greece ca 42
_ (as A. Bucesas) 6438 cultivated Madrid Botanical Garden ca 42 18
A. hololeios Bornum 6493 cultivated 4309 Universitatis Bergensis, Norway 16 19
A. micropteris Fisch. 6446 O. Tosun Ankara, Turkey via USDA. 32 14
*A. monspessulanus L. 6361 E. Muller 22/8/61 Tiefenkastel, Switzerland 16
*A. odoratus Lam. 6445 O. Tosun Ankara, Turkey via USDA. 16 13
A. ovalis Boiss. 6443 O. Tosun Ankara, Turkey via USDA. 16 17

* These species previously counted; cases of disagreement discussed in the text.

OZ&

. sesameus L.

. spinosus Muschl

. uliginosus L.

. umbellatus Bunge

. vulpinus Willd.

*Oxytropis campestris (L.) DC

*O.

o

5

var. gracilis (A. Nels.) Barn.

deflexa (Pall.) DC.
var. sericea T. & G.

”

. halleri Bunge
. monticola A. Gray

. multiceps Nutt.

parryi Gray
”

. sericea Nutt.

var. spicata (Hook.) Barn.

cultivated

J. Mandaville
20/4/63

cultivated

J. G. Dickson
26/8/56

cultivated

G. H. Turner 15/8/60

cultivated
G. H. Turner 24/7/60
G. H. Turner 10/8/60

cultivated
cultivated

C. L. Porter 8322
M. DeDecker 1523
M. DeDecker 1534
C. L. Porter 8458
G. H. Turner 11071

* These species previously counted; cases of disagreement discussed in the text.

Madrid Botanical Garden
Saudi Arabia

Vladivostok, USSR.

Glenn Highway, Alaska

Russia

Fort Saskatchewan, Alberta
Udaipur, India

Fort Saskatchewan, Alberta
Fort Saskatchewan, Alberta

Inst. Alpin du Lautatet, France

Kamploops Expt. Farm, B.C.
Albany Co., Wyoming

Mono Co., California
Inyo Co., California

Albany Co., Wyoming
Jasper Nat. Park, Alberta

IGé

322

has 2n=32 while the collections from Ontario, Quebec and Russia have

n=16. Professor C. Favarger in personal conversation at the IXth Inter-
national Botanical Congress in Montreal in 1959 said that he had found
both 2n=16 and 2n=32 in some Switzerland collections of A. alpinus.
Ledingham (1960) reported on four collections, three from Alberta,
Manitoba and Saskatchewan having 2n=16 and one from Yukon having
2n=32. The count of 2n=ca 56 listed in Darlington and Wylie (1955) is
considered erroneous.

It should be pointed out that although the present chromosome count
2n=32 for O. campestris var. gracilis agrees with some of the previous
reports for this species there are also reports of 2n=48. Ledingham
(1957, 1960) reports 2n=32 for six Saskatchewan and one Alberta col-
lection. Jalas (1950) for ssp. sordida and Ledingham (1960) for one Old
World and Ontario and British Columbia material report 2n—48.

Plates I and II give some camera lucida drawings of chromosomes of
representative species of New World and Old World Astragalus. These
drawings are all done with the same apparatus and at the same mag-
nification. Although it seemed as if New World species had smaller
chromosomes which were more difficult to separate and count this is not
supported by the drawings. Plate I illustrates chromosomes of New
World species and Plate II shows chromosomes of Old World species of
Astragalus.

DISCUSSION

This paper reports chromosome counts for 28 species of Astragalus
not previously reported. Twently of these counts are of New World
species (Table I and Figs. 1-12) and they form an aneuploid series with
n=11, 12, or 18. Eight new counts are reported for Old World species,
5 in Astragalus and 3 in Oxytropis, (Table II and Figs. 13-22) and these
have 2n=16, 32 or 48, ie. are diploids, tetraploids or hexaploids of the
8-series,

These 28 counts together with additional counts for 21 other species
for which there have been previous reports further support the con-
clusion (Ledingham, 1960) that there are two main phylogenetic lines
in Astragalus. There are now counts for 109 New World species and all
have n=11 (53 species), 12 (38), 13 (14), 14 (3) or 22 (A. grayi, the only
New World tetraploid). There are counts for 202 Old World species of
Astragalus and Oxytropis and they have n=8 (146 species), 16 (21),
24 (18), 32 (8), 40 (1), 48 (2), ca 80 (1) and others (5).

In an earlier paper Ledingham (1960) gave a count of n=10 for
Astragalus somalensis and suggested that species with the intermediate
chromosome numbers n=9 and 10 might form part of the aneuploid
series which includes both Old and New World Astragalus. No further
evidence has been found to support this idea, and since Gillett (1963) has
removed A. somalensis to Galega, it seems less likely. There are now
no known species with n=9 or 10 in the Astragalus complex.

323

The suggestion has been made (Turner, 1959) that the n=14, 138, 12
and 11 of New World species of Astragalus are derived hypoploids from
an ancestral n—16 tetraploid species. There are, however, no tetraploids
in North America except for a few circumpolar species which are
actually a part of the Old World phylogenetic line. There are several
species in the Old World which may be hypoploids, e.g. the annual con-
spicuously self-fertile species A. boeticus, 2n=30.

Our counts of 2n=ca 42 for A. hamosus make us suspicious of the pre-
vious report of 2n=48 for this species. Our suggestion for the moment
is that A. hamosus does not really belong in Astragalus. It seems likely
that A. hamosus is a hexaploid in some n=7 phylogenetic line. Tetra-
ploids in this line may include A. pentaglottis, 2n=—28 (Senn, 1938; Led-
ingham, 1960) and A. bubaloceras, n=ca. 14-15 (Senn, 1938). Diploids
of this n=7 line are unknown. There is, then, little evidence of hypo-
ploids in Old World Astragalus and it seems very unlikely that New
World Astragalus has arisen by chromosome loss from tetraploid
(2n=32) plants of the Old World line.

Although we do not yet have enough information to reconstruct the
evolutionary history of the Leguminosae, or more specifically of Astra-
galus and Oxytropis, it is clear that chromosome numbers can be used
with considerable confidence to show true relationship in this family.
This paper further establishes that New World and Old World species
of Astragalus have had a different evolutionary history and must be
considered as different subgenera or genera if taxonomy is to re-
flect true relationships. The chromosome evidence would indicate that
Oxytropis is closely related to, or is a part of, the Old World Astragalus.
It is now clearly established that chromosome number can be used as
a significant character in this family. The relationships of species with
anomalous counts should be studied critically.

ACKNOWLEDGMENTS

It would be impossible to list all of a many cae ns and institutions who have helped
We Id like er, nks

ith our research. wou ike, howe e special tha the following who
provided material for this study: Mr. R. ee Ss Miss M. Belcher, Mrs. E. Beckett
Mrs. M. DeDecker f Davis, Dr. A. Dutilly inbrun, ptre epage,

: Au Cy LE,
oe and Dr. G. H. Turner. We would like, also, to acknow ee our debe to Mr.
as Barneby who has eed identified our herbarium specimens.
are grateful to the tional Research Council, Ottawa, Canada for the financial
aid a has been given to a proje

=

REFERENCES
ARLINGTON, C. D. and A. P. WYLIE. 1955. Chromosome atlas of the flowering

aie Macmillan, New York.

GILLETT, J. B. 1963. Galega L. (Leguminosae) in tropical Africa. Kew Bull. 17: 81-85.

HEAD, S. C. 1957. Mitotic chromosome studies in the genus Asfragalus. Madrono 14:
95-106

JOHNSTON, I. M. 1947. Asfragalus in Argentina, Bolivia and Chile. Jour, Arn. Arb.
28: 336-40

324

DINGHAM, G. F. 1957. Chromosome numbers of some Saskatchewan Be as
aa particular reference to Astragalus and O xytropis. Canad. Jour. Bot. 35 666.
————_——.. 1958. Chromosome numbers in Asiisealis. Proc. Genet. pi Cink 3:

15-18.

—————.. 1960. Chromosome numbers in Astragalus and Oxyfropis. Canad. Jour.
Genet. Cytol. 2: 119-128

——_____——. and B. M. REVER. 1963. Chrom - numbers of some as hee
species of Nees and Owytropis eon anad. Jour. Genet. Cytol

OVE, A. and D. LOVE. 1961. Chromosome ieee of central and ae aa
pean plant species. Opera Botanica 5: 1-581.
SENN, H. - 1938. Chromosome number relationships in the Leguminosae. Bibliogr.
Genet. 12: 175-336.
Sa R, B L. 1956. Chromosome numbers in the Leguminosae. Am. Jour. Bot. 43:
- 1959. The Legumes of Texas. University of Texas Press, Austin.
————— and O. S. FEARING. 1959, Chromosome numbers in aca
II: African species, including phyletic interpretations. Am. Jour. Bot. 46: 49
oe H. 1943. Chromosomes of Asfragalus. Bull. Torr. cs Club 70:
430-
YURTSEY B. A. 1963. On the floristic relations between steppes and prairies. Bot, Not.
116: 396-40

PLATE I
Figs. 1-12—Somatic chromosomes of New World species of Astragalus
drawn with the aid of a camera lucida originally at x 2250 reduced by
50% in reproduction.

Fig. 1 A. desperatus var. desperatus, 2n=24, Rever 78.

Fig. 2. A. amphioxys var. amphioxys, 2n=22, Rever 72.
Fig. 3. A. inyoensis, 2n=22, DeDecker 1519.

Fig. 4. A. lentiginosus var. fremontii, 2n=22, DeDecker, 1484.
Fig. 5. A. johannis-howellii, 2n=22, DeDecker, 1505.

Fig. 6. A. coccineus, 2n=22, DeDecker, July 22, 1962.

Fig. 7. A. ceramicus var. imperfectus, 2n= 22, Porter 3954.
Fig. 8. A. purshii var. glareosus, 2n=22, Rever 111.

Fig. 9. A. whitneyi, 2n=22, DeDecker 1535.

Fig. 10. A. cymboides, 2n=24, Rever 58.

Fig. 11. A. miguelensis, 2n=22, Raven 18012.

Fig. 12 A. calycosus, 2n=22, DeDecker 1522.

Origin of material given in Table I.

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PLATE II

Figs. 13-22. Somatic chromosomes of Old World species of Astragalus and
Oxytropis drawn with the aid of a camera lucida originally at x 2250
reduced by 50% in reproduction.

Fig. 13.
Fig. 14.
Fig. 15.

A.

odoratus, 2n=16, O. Tosun.

A. micropteris, 2n=32, O. Tosun.

A. spinosus, 2n=16, J. P. Mandaville, April 20, 1963.
A. angustifolius, 2n=32, O. Tosun.

A. ovals, 2n—16,.O- Tosun,

. Az
A
A
O
O

hamosus, 2n=ca. 42, Ledingham 2805.

. hololeios, 2n=16, Universitatis Bergensis 4309.
. vulpinus, 2n=16, Russia.

. parryi, 2n=16, DeDecker 1523.

. monticola, 2n=48, Kamloops 1939.

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YELLOW-FLOWERED LINUM (LINACEAE)
IN TEXAS
C. MARVIN ROGERS
Department of Biology, Wayne State University, Detroit, Michigan

During the course of some studies of the genus Linum, I have had
the opportunity to examine a large number of specimens from Texas.
Since the state is of interest in having far more species than any other
and since, henceforth, attention will not be given particularly to the
Texas taxa, it seems profitable to bring together at this time some of
the accumulated data. Some collections from the state have not yet been
seen and these no doubt would reveal some more county records, but
it seems unlikely that the distribution patterns which have emerged
will be greatly changed.

The features used to distinguish the various taxa, especially in the
L. rigidum group, will be discussed further at a later date. The characters
used in the key are certainly not necessarily the most significant. Living
material of most of the Texas species is now at hand and it may be that
cytological and genetic studies of these plants will result in some modi-
fication of the species and varieties as treated here, but a sufficient
amount of material has been examined that the present interpretation
should provide a basic framework for the field identification and recog-
nition of the Texas taxa.

For the ranges of some of the plants described here, collections from
about 45 herbaria were examined, but for species of the L. rigidum group
(L. alatum, L. aristatum L. australe, L. puberulum, L. rigidum and L.
vernale) distributional data comes principally from the collections of
the United States Museum, the Gray Herbarium, the New York Botanical
Garden, the Missouri Botanical Garden, the Chicago Museum of Natura
History, Southern Methodist University, the University of Texas and the
Lundell Herbarium, while for L. rupestre and L. schiedeanum the records
come mainly from the last three. To the curators of all of these collec-
tions thanks are gratefully given.

The features of the genus as found in the state (excluding the two or
three blue flowered species) may be summarized as follows:

labrous or occasionally pubescent annual or perennial herbs; leaves
simple, sessile, entire or the upper glandular-toothed, alternate, opposite
or rarely whorled on the lower part of the stem, alternate above; stipular
glands present or none; inflorescence a terminal scorpioid cyme; flowers
regular; sepals 5, imbricate, all or only the inner commonly with glandu-
lar-toothed margins; petals 5, convolute, separate, yellow, fugacious:

‘Contribution No. 121 from the Department of Biology, Wayne State University, De-
troit, Michigan

SIDA 1 an 328—336. 1964.

329

stamens 5, united basally, with or without diminutive intervening
staminodia; ovary superior, 5-carpelled, but becoming more or less com-
pletely 10-locular through the intrusion of false septa, dehiscing into 5
or 10 segments; seeds 10; styles 5, separate or united; stigmas capitate.

KEY TO THE SPECIES AND VARIETIES
la. Styles separate or nearly so; fruit ultimately dehiscing into 10 one-

3a. Fruit pyriform, longer than broad;
pollen about 10-colpate . . Ll. L. floridanum
3b. Fruit spheroidal, as broad or noua en ee pollen 3-colpate.
4a. Margins of inner sepals with conspicuous stalked glands; mature
fruit in dried specimens usually adhering to the plant; leaves
narrowly lanceolate or oblanceolate 2. L. medium var. texanum
4b. Margins of inner sepals glandless or with very inconspicuous
glands; mature fruit in dried specimens usually soon shattering;
leaves elliptic to oblanceolate or obovate . . . 3. L. striatum
2b. Sepals all with glandular teeth
5a. Perennial; styles completely separate; pollen 3-colpate
6a. Leaves lanceolate or oblanceolate or atc some of the lower
ones in whorls of four . . L. schiedeanum
6a. Leaves linear, the lower ones sien or pene
5. L. rupestre
5b. Annual; styles united at the base; pollen aoe: about 20 germ
pores 6. L. suleatum
lb. Styles united to above the middle; fruit achiveue along the false
septa into 5 two-seeded segments
7a. Sepals entire or fringed, not glandular-toothed
8a. Upper leaves and bracts sparsely, but conspicuously ciliate-
margined; cartilaginous portion of false septa conspicuously wider
toward the base of carpel . 7, L. imbricatum
8b. Upper leaves and bracts not Aiiate: anayeined: cartilaginous por-
tion of false septa uniformly narrow or absent throughout
8. L. hudsonioides
7b. Sepals glandular-toothed
9a. Plants grayish puberulent throughout . . . 9. L. puberulum
9b. Plants glabrous or nearly so throughout
10a. Outer sepals ovate, the broad, scarious margins irregularly
crenate, each of the coarse teeth bearing a delicate gland
0 L. alatum
10b. Outer sepals lanceolate or narrower, the margins not scarious
or narrowly so, regularly, though sometimes sparsely, serrate
with gland-tipped teeth
lla. False septa incomplete, the inner c adaoes terminating in a
loose fringe; sepals persistent in fru : 11. L. vernale

llb. False septa complete; sepals usually deciduous in fruit
12a. Leaves small, the lower tending to be hidden among the
branches; plant broomlike, bushy with long, slender stiffly
spreading-ascending, few-flowered branches
. 12. L. aristatum
12b. Leaves quite evident; plants not ‘proomaliice: rather few-
branched at the base or in the inflorescence
13a. Stipular glands absent
14a. Styles 6—10 mm. long
15a. Stigmas pale; sepals green
16a. Fruit thin-walled (dark seeds commonly evident through the
wall), elliptic, the base rounded.
13a. L. aioe var. rigidum
16a. Fruit thick walled, opaque, broadly ovoid, tapering abruptly
at the flattened base . . . 13b, L. rigidum var. berlandieri
15b. Stigmas black: sepals sieioielh: or purplish. .
: e Ll. dane: var. filifolium
14b. Styles 3—4 mm. long .. . “13d. L. rigidum var. compactum
13b. Stipular glands present (sometimes on the lower part of plant only)
17a. Styles more than 6 mm. long; petals more than 10 mm. long
18a. Sepals green; stigmas pale . 13b. L. rigidum var. berlandieri
18b. Sepals grayish or purplish; stigmas black.
13c. L. Paae var. . filifolium
17b. Styles less than 6 mm. ign: Ee less than 10 mm. lon
19a. Stipular glands present only near the base of the plant
. 14a. L. australe var. iia
19b. Stipular glands present ate ieoninent throughout .
14b. L. australe var. Si inaiosun

1. LINUM FLORIDANUM (Pianchs Trel. var. FLORIDANUM. This,
with the next two, is a part of a series of seven perennials, all confined
to eastern North America. (See Brittonia 15: 47-122, 1963, for further dis-
cussion of these two species.) Though differing from the next in perhaps
a dozen qualitative and quantitative characters, this variety is closely
related to L. medium var. texanum, and is thought to hybridize with it.
Several collections from eastern Texas appear to involve L. floridanum
as a parent and it is possible that additional collecting in that part of
the state will show it to be more widespread there. It is a fairly common
plant in pine and pine-palmetto woodlands throughout much of the
southern Atlantic and Gulf Coastal Plains, but is presently known in
Texas from a single specimen from Hardin County.

2. LINUM MEDIUM (Planch.) Britton var. TEXANUM (Planch.) Fern.
The typical variety of this species is confined to Ontario, but variety
texanum ranges throughout most of eastern United States, westward to
southeastern Iowa and eastern Texas, where it is a species of open woods,
meadows and grassy roadsides.

eee eee
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6.L.sulcotum | jst
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332

3. LINUM STRIATUM Walt. This species is also widely distributed in
eastern United States, mostly in the nonglaciated regions in moist situa-
tions along the borders of ponds, streams and roadside ditches. Perhaps
two dozen collections from scattered localities in 15 counties in eastern
Texas have been seen.

4. LINUM SCHIEDEANUM Schlecht. and Cham. This and the next
are part of a complex of taxa, principally Mexican, of which the identity
and relationships are far from completely clear. It seems fairly certain
that the Texas plants included here belong to a very widely distributed
species, correctly interpreted as L. schiedeanum. As such, it is found
mostly in calcareous soil from southern Mexico northward to the Chisos,
Del Norte, Glass and Guadalupe Mountains of western Texas and south-
ern New Mexico.

5. LINUM RUPESTRE (A. Gray) Engelm. This variable species occu-
pies about the same range and habitat as the last and the two are found
together over much of their ranges. Linum rupestre is found throughout
most of Mexico, with central and western Texas constituting the north-
ernmost limit of its range.

6. LINUM SULCATUM Riddell. This species is found in prairies and
prairie like areas throughout central and northeastern United States and
southern Canada, but is infrequently collected in the southern part of
the range. It is of special interest in the genus, since it combines traits
of the primitive L. rupestre and the highly specialized L, rigidum groups.

7. LINUM IMBRICATUM (Raf.) Shinners. This and the next species
resemble one another and have not generally been separated. (See
Rhodons 65: 50-55, 1963, for further discussion of these three species.)
They are small plants of distinctive habit with many, small imbricate
leaves and few-flowered inflorescences. Though specimens have come
from along both the Oklahoma and Mexico boundaries, the known range
lies entirely within Texas, where it is a plant | of sandy soil
throughout much of the east central part of the state.

8. LINUM HUDSONIOIDES Planch. Along with L. wmbricatum, this is
closely allied to the L. rigidum group which follows. It is found princi-
pally in sandy or gravelly, sometimes calcareous soil in the west central
part of the state, with outlying stations in the trans-Pecos region of west
Texas and southern New Mexico, the Wichita Mountains and perhaps in
central Kansas.

9. LINUM PUBERULUM (Engelm.) Heller. This is easily recognized
as the only densely pubescent species in the region. It is closely related
to L. australe and occupies about the same range as that species at low
and medium elevations in the mountains from southeastern Wyoming to
Utah, south into northern Mexico and eastward through the trans-Pecos
region of western Texas where it is found in rocky, sandy or occasionally
calcareous situations.

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334

10. LINUM ALATUM (Small) Winkler. Like the previous species, this
may be quite readily recognized, in this case by the unique sepals.
Though it differs in other ways from L. rigidum var. berlandieri, it is
surely closely related to that plant. The ranges of the two overlap, but
there does not appear, in the specimens examined, evidence of hybridism.
Linum alatum is found in sandy soil, sometimes along the beach, in south-
eastern Texas, with a collection or two from Tamaulipas, Mexico, near
the Rio Grande and from western Louisiana.

11. LUNUM VERNALE Wooton. The poorly developed false septa and
the tendency for the sepals to persist, together with several other fea-
tures such as fruit shape and texture, indicate that this may be one of
the more primitive species of the L. rigidum group. The species is found
on stony, commonly limestone hills in the trans-Pecos region of Texas,
adjacent New Mexico and northern Mexico.

12, LINUM ARISTATUM Engelm. This is a distinctive species, though
certain of its features do not lend themselves to precise description. In
addition to the characters indicated in the key, it has unique pale, nar-
rowly elliptic, thin walled, easily crushed capsules. It is a species of
sandy soil, ranging from eastern Utah and western Colorado south to
northern Mexico and eastward into west Texas.

13a. LINUM RIGIDUM Pursh var. RIGIDUM. Linum rigidum ranges
from southern Canada to central Mexico, with the Mexican populations,
partly because of the scarcity of good collections, being poorly known.
Long a source of confusion, the complex of which this is a part is the
subject of some studies now in progress. In Texas the species appears to
consist of four varieties which, though tending to intergrade somewhat in
some areas, are sufficiently distinct that nearly every plant can be
readily named. What is usually interpreted as the typical variety (the
type has not been located) is a plant of the plains, ranging from
north central Texas (one specimen from Aransas County) northward to
Alberta and Manitoba. It is the tallest variety (average about 30 cm.)
with an open, few flowered inflorescence, relatively large floral parts
and no stipular glands.

13b. L. RIGIDUM var. BERLANDIERI (Hook.) T. & G. This is a showy
plant, with some reason often considered a separate species, It is gen-
erally shorter (average 15-20 cm.) and more compact than var. rigidum,
with leaves averaging twice as wide (2 mm.) and with sepals and floral
bracts tending to be coarser and 3-nerved rather than 1-nerved. In
the northern part of the range, where it overlaps that of var. rigidum,
a number of collections appear to be intermediate. In southern Texas

short, bushy branched, leafy variation which probably warrants further
study. While the key is designed to include plants without stipular glands,
they are present in nearly 90% of the specimens examined of this and

i i
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rr ba ol Anyang ei
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I3a. L.rigidum — ~ corpus Christi 4 I Sb. be rigidu NS Gh
var. rigidum — PLaredo ar. perigndieg :
ace
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t i
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1 i ‘Lubba U i
pt Dallas, \ i ots re
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itd yes e eee ees <
Ayes ad San/Angelo, a ee , 5am Angelo, ‘ Maco, i
a 1 : . es aN aan
\ Lg . — . . LVN
i \ : i eee c)
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a 4 Soe, r uston?
San ANTOD, i “J i \ — Antonio,
I3c. L. rigidum \ pong I3d.L. rigidum %: Corpus Christi
ili i Laredo.
var. filifolium e 0 ar. compactum ani
‘Re aon
— ~
ee eee ‘aied es WEE
a4 ror
i i | i
| Amarillo a i Amarillo j
i eel.
so ie , i hi Fas? se ete yg
il lo wichita ae ry] igs : ichita Fatts =
{ ~ ‘Lubbdck” ae i i Lubbdc tals LY 1
+ + T o t we
i - Fort Worth? 2 ; H Fort Worth woe
i RE ce nen : N.
SET Paso TOTS! rE | | Mago me ac San Angelo Waco, : \
if San‘Angelgg ” ‘ : BEMo Tx EN {
T 2 x | if
7 i : Sd é
“S Me 4 Austin )
\ siete Beaumonto , \ .. g. Beaumonto ¢”
* ., \ . —se 7
\ aoe A “ ‘San Antonio,
Oi ae San Antonio, Seat \ poo
~

i4qa. L. australe
vor. australe

X Corpus Christi
“Ylarede
\

\
\.
I4b.L. australe Corpus Christi
pitsreo
\

var. glandulosum*.

336

of variety filifolium. Variety berlandieri is found pretty much through-
out the state and as far north as southeastern Colorado and central

ansas

13c. L. RIGIDUM var. FILIFOLIUM Shinners. As interpreted here, var.
filifolium is a rather variable population. Most collections have come
from sandy, rocky or sometimes calcareous soil in west Texas, but a few
puzzling plants included here come from the southern plains country,
while southeastward near the Mexico boundary, there is a gradual tran-
sition toward plants which are often perennial and possess long, slender
sepals. These have been called L. elongatum. More study is necessary to
determine the relationship of the west Texas plants to those found from
Webb to Hidalgo Counties, as well as to some anomalous collections from
northern Mexico.

13d. L. RIGIDUM var. compactum (Nelson) Rogers, comb. nov. (L.
compactum, Nelson, Bull. Torr. Bot. Club 31: 241, 1904). This

small floral parts, no stipular glands and with rather coarse foliage and
fruit somewhat as in var. berlandieri. Like var. rigidum, this is a plant
of the plains, being found from northern Texas to southern Canada.

14a. LINUM AUSTRALE Heller var. AUSTRALE. Linum australe is a
plant of low and medium elevations in the Rocky Mountains, most
closely allied to L. puberulum, but also very likely related to L. rigidum.
In Texas two varieties may be recognized. The typical variety, which has
been collected in the Chisos and Davis Mountains, is found from north-
ern Mexico to Alberta.

14b, L. AUSTRALE var. glandulosum Rogers, var. nov. Differt a var.
auStrali,quod stipulas glandulosas atque clarissimas, flores paulo maiores,
fructum paulo minorem hebet. HOLOTYPE: Palmer 465, Otinapa,
Durango, Mexico (US; isotypes F, GH, MO,

This differs from var. australe in the possession of very conspicuous
stipular glands, these often being found at the base of the sepals as well
as the bracts and leaves, and in the somewhat larger floral parts and
smaller fruits. It is found from southern Arizona and the Davis Moun-
tains in western Texas, south to Pueblo, Mexico.

CALYLOPHUS (OENOTHERA IN PART:
ONAGRACEAE) IN TEXAS
LLOYD H. SHINNERS
Herbarium, Southern Methodist University, Dallas, Texas 75222

Dr. Raven’s new look at Oenothera and its allies is sure to win
adherents, though it will take some time to become adjusted to the
unfamiliar nomenclature. Among the species conservatively included
in Oenothera, those with entire or subentire stigma and widely flaring
summit on the calyx tube form a very well-marked group. Users of the
floras of Small and Rydberg have known them as Meriolix and Galpinsia,
but the first generic name published with description was Calylophus.
All the species recognized variously by Munz and Raven occur in Texas,
as do most of the infraspecific taxons. Because a monographic treatment
by Dr. Raven is not likely to be ready in time for one local and two
state floras now in active preparation, this brief review has been pre-
pared. It is based on 308 specimens in the S.M.U. Herbarium (205 of
them from Texas), plus original descriptions of species and varieties
previously published, and a few notes on type specimens given by Munz
(1929)

Just as among the genera there are almost no differentiating char-
acters that hold throughout, so every promising character for separating

pattern. For example, extremely short sepal-tips on the flower buds are
typical of C. Hartwegii var. lavandulaefolius, but rarely a plant typical
in other respects may have long sepal-tips. This conceivably could result
from introgression with var. Hartwegii in the rather large area in which
both occur. But in southern Trans-Pecos Texas the reverse variation is
found, var. Hartwegii rarely showing extremely short sepal-tips, here
outside the range of var. lavandulaefolius. I believe that the basic varia-
tion pattern, and chief cause of taxonomic difficulty, is one of homo-
logous mutations, appearing especially in pubescence and leaf-form. In-
trogression also occurs (to a limited extent among the varieties of C.
Hartwegii, more extensively between those of C. serrulatus), and along
with seasonal variation, helps to complicate the picture. All this makes
construction of dichotomous keys most difficult. Those given here are
intended for identification, not definition, and are for use with plants in
flower. When leaf dimensions are used, they are those of leaves on
flowering stems. In the text are mentioned additional features which,
though commonly present and helping to define the varieties, break
down too often to be really useful in a working key.

SIDA 1 (6): 337-345. 1964,

338

Additional synonymy is to be found in the two publications of Munz
(1929, 1944). I have cited only what was necessary to establish the
nomenclature here adopted and to indicate the principal departures from
the treatments of Munz. It would be most easy to add a long, specula-
tive discussion of the history and relationships of the recognized taxons,
but I do not feel that present knowledge justifies it.

KEY TO SPECIES

la. Sepals with prominent raised midrib or low keel, foe buds 4-
ribbed or 4-ridged, especially toward summit. . C. serrulatus

lb. Sepals without prominent midrib or keel, flower 7 smooth
2a. Calyx tube funnelform in upper 2/3 or more, 6—30 mm. long
(above ovary) . . C. tubicula
2b. Calyx tube esas in upper 1/2 or igs! 1b 55 mm. long . .
3. C. Hartwegii
1. C. SERRULATUS (Nuttall) Raven, Brittonia 16: 286. 1964. Two var-
leties are recognized for Texas, with intermediates due partly to intro-
gression, but also due partly to variation trends not related to introgres-
sion. More intensive study may justify recognition of the large-flowered
southern phase of var. serrulatus, occurring mainly outside the range
of the even larger-flowered var. spinulosus. Both varieties show more or
less clinal variation, var. serrulatus in flower size, var. spinulosus in
leaf-width, but the extremes are not considered worthy of nomenclatural
recognition. A single specimen from Arizona combines the small flowers,
short leaves, and pubescence of var. serrulatus with the narrow leaf-
dimensions of var. spinulosus, but is outside the range of both. I consider
this as belonging to a third variety, illustrating independent mutation or

recombination of characters derived from a remote ancestor.!

KEY TO VARIETIES OF C. SERRULATUS
la. Leaves 3'2—9 times as long as wide . . var. serrulatus
lb. Leaves (except lowest) 9—40 times as one as wie a oe as
var. spinulosus.
la. C. SERRULATUS var. SERRULATUS. Oenothera serrulata Nuttall,
Genera 1: 246—247. 1818. rom the river Platte to the mountains, on
dry hills; flowering in June. ... Stem simple, slender, 8 to 12 inches
high, foliose; leaves a little more than an inch long, 2 to 3 lines wide,
attenuated downwards, distinctly serrulate, not toothed... .” O. serrulata
var. Nuttallu T. & G., Fl. N. A. 1: 501. 1840. (Based on the preceding,
equivalent to what we now call var. serrulata.) O. serrulata var. Drum-
mondti T. &. G., lc. 502. “Low, minutely puberulent; stems simple:

~ 1 CALYL OPHUS SERRULATUS var. arizonicus swe var. nov. Ad var. serrula-
‘um statura minore, foliis et floribus parvis, pubescentia cinerea praecipue partium juniorum,
ad var. spinulosum ate perangustis (e.g. x 2.2 mm. 28 BC 2.8 ele heey . HOLO-
TYPE: 4 miles upstream Pa White River on the White River, Nav Co., Arizona,

S. J. Preece, Jr. & B. L. Turner 2692, 25 June 1951 (SMU). “Dry sandy river bank; silty
sandy soil. Plant 1 foot or less heh. Petals yellow. Tap-root woody.”

339

leaves linear-spatulate or spatulate-oblong ... ; flowers larger; capsules
puberulent ... Texas, Drummond!” (Based on Calylophus Drummondit
Spach, whose description I have not seen, but the different form of the
epithet makes it a new name rather than a new combination, and the
citation of a specimen seen by T. & G. could also justify treating it as an
entirely new and independent though synonymous name.) This name
has been used by Munz chiefly for var. spinulosus, but the original
description clearly indicates the larger-flowered phase of var. serrulatus
common in southern Texas, while var. spinulosus is absent from the
counties botanized by Drummond.

Plant low-growing, more or less gray pubescent, with short, subentire
or rather bluntly and inconspicuously toothed leaves and small to
moderately large flowers. Variations in Texas which may be due to
introgression with var. spinulosus include plants with tall stems, or
largely glabrate, or with large, prominently spinulose-toothed leaves,
or in some cases large flowers, or combinations of these features. Proba-
bly most of the variation in toothing of leaves represents spontaneous
mutations. At least in the southern part of the range, I believe that

a

outside the area of var. spinulosus. In
flowered plants were treated under var. Drummondii, but flower size
does not correlate well with leaf-dimension, which I believe permits a
better geographic separation.

Panhandle to Grand Prairie (Denton and Tarrant counties), south
and southeast to the lower Rio Grande Plain and Coastal Bend (east to

ilam, Brazos, Jackson, and San Patricio counties), southwest to
Loving, Ward, Val Verde, and Webb counties; absent from the Trans-

are considered aberrant forms of this variety on grounds of locality (well
outside the range of var. spinulosus), dwarf stature, and short leaves.
KENEDY CO.: Yturria Ranch near Willacy Co. line, Lundell & Lundell
8735, 6 May 1940. (Leaves 31 X 3.1 mm., 27 X 3 mm.,, 26 X 3.2 mm.;
plant apparently normal. Other collections from the region very similar
to it have slightly wider leaves.) KLEBERG CO.: 12 miles southwest
of Riviera, infrequent on sandy roadside, Cory 55259, 31 March 1949.
(Leaves 30 X 1.1 mm., 25 X 2 mm.; an injury form, with small, slender
shoots from apparently mowed and perhaps burned plants.)
lb. C. SERRULATUS var. spinulosus (Nuttall, ined.; ex T. & G.)
Shinners, comb. nov. Oenothera serrulata var. spinulosa (Nutt., ined.)
= A. 1: 502. 1840. “Taller, often branching, almost glabrous;
leaves linear, elongated, acute (sometimes obtuse), spinulose-serrate;
flowers rather large; capsules minutely pubescent.—OE. spinulosa, Nutt.!
ined. ... Arkansas, Nuttall! Dr. Leavenworth!” The locality refers to
the Arkansas Territory of that time, including eastern Oklahoma whic

340

almost certainly was the actual type of locality; I have seen no speci-
mens from Arkansas. Reference is made to Hooker, Exotic Flora 2 t.
140, 1825, which shows the upper portion of a plant of this variety as
here understood, the form with pure yellow flowers. As mentioned above,
the plants treated by Munz as Oenothera serrulata var. Drummondii
chiefly belong here.—Oenothera serrulata var. pinifolia Engelm. ex
Gray, Boston Journ. Nat. Hist. 6 (Pl. Lindh. 2): 189. 1850. “Rocky prairies,
New Braunfels. April.—This is just the OE. serrulata var. spinulosa,
except that the leaves are extremely narrow.” (See also Heller’s com-
ment, te under the next.)—Meriolix melanoglottis Rydb. ex Small,

1. . U.S. 846 and 1335. 1903. Type collection Heller 1600, about Kerr-
ville. — Co., Texas, 12—19 June 1894. There are two sheets of this
number at SMU, apparently belonging to what had been two different
collections, later combined under one distribution number. One is a
whole plant 24 cm. tall, with root, in early flower, with linear-
oblanceolate leaves, a representative one measuring 45 «x 4.5 mm. The
other has a piece of stem almost 11 cm. long with 7 branches up to 40 cm.
long, bearing a few flower buds, and lance-linear leaves, a representa-
tive one measuring 43 x 2.5 mm. The original description covers both
forms. Heller himself gave these comments in his Botanical Explora-
tions in Southern Texas (Contrib. Herb. Franklin & Marshall College
1: 71, 1895): “Plentiful about Kerrville, especially along the Guadalupe
and Town Creek, usually in gravelly or stony ground. Not only the throat
of the calyx and the disk-shaped stigma are dark black-purple, but also
the throat of the corolla. Of the hundreds of flowers seen, hardly half
a dozen were without this marking. The variety pinifolia is merely a
very narrow leaved form of this species. Both forms grow together and
there is no other character to distinguish them.”

Plant rather tall, glabrate, with long, very slender, spinulose-toothed
leaves and moderately to very large flowers. Variations perhaps due to
introgression with var. serrulatus are lower, or with more pubescence,
or shorter leaves with less prominent teeth, or smaller flowers, or
combinations of these features. Variation plainly due to spontaneous
mutation is that in leaf form, running to the extreme which was named
var. pinifolia, concentrated about the Edwards Plateau. I believe that

ark pigmentation in the center of the flower, found in the southern
part of the range, also represents spontaneous mutation.

Edwards Plateau, Grand Prairie and Blackland Prairie, extending
west to Taylor and Val Verde counties, east in the northern part of its
range to Prairie Border (Van Zandt Co.). Found north and northeast

(where perhaps introduced; the one specimen seen is from along rail-
road in Waukesha Co.). On the basis of the description given in Steyer-
mark’s Flora of Missouri (1963, p. 1102), it is this variety which is re-
ported from that state as Oenothera serrulata. It occurs also in Coahuila.

341

Caroline Dormon, in Flowers Native to the Deep South (1958, pp.
84—85), speaks of Meriolix melanoglottis as “entirely distinct” (from
Oenothera spinulosa), ‘“‘and in the author’s humble opinion it deserves
specific rank. A perfect rock-garden plant, it has spreading wiry stems
and linear leaves with a few scattered teeth. The lovely bright yellow
flowers, about 14% inches across, have many tiny folds, giving them a
‘crepy’ look. They open out flat and remain open all day, as do those of
Cream-cups. The most distinctive feature of the flower is the litle black
‘tongue’ (stigma) ...In the Deep South it is really a tiny shrublet, which
remains green all winter. . . . Texas and southwestern Louisiana.” I
have not seen Louisiana material, but the supposedly distinguishing fea-
tures described by Miss Dormon can be found in Texas in various combi-
nations among plants of what I would consider perfectly typical spinu-
losus. The variability at the type locality for Meriolix melanoglottis ob-
served by its original collector has already been mentioned. At Dallas
the plant puts out slender, trailing to ascending, almost vine-like shoots
with small leaves that are green over winter, just as Miss Dormon
describes.

2. C. TUBICULA (Gray) Raven, Brittonia 16: 286. 1964. (The epithet
is not in available dictionaries. Presumably it is an atypical diminutive
of tuba, but the case of radicula and radula, words unrelated to each
other, eee doubts.) oe tubicula Gray, Smithsonian Contrib.
3 art. 5 (Pl. Wright. 1): 71. 1852. “Prairies beyond the Pecos; Aug.,”
Wright 197. “Also sores in much larger and much better pecans
in the collection of 1851.” Including var. demissa Gray, ibid. 71—72.
the Guadalupe Mountains; Oct.,” (Wright 197 (partim)).

Northeastern Trans-Pecos and adjacent counties just east of the Pecos;
specimens seen from Brewster, Culberson, Pecos, Presidio, Reeves,

or short-lived perennial with rather short, broad leaves; large-flowered
forms are extremely similar superficially to C. Hartwegii var. Hartwegi
(particularly those forms of the latter which have been treated as
Oenothera Hartwegii var. Fendlert) ).

3. C. HARTWEGII (Bentham) Raven, Brittonia 16: 286. 1964. A trouble-
some assemblage of forms, most of them wide-ranging and overlapping
geographically. The extremes appear quite distinct, but there is so much
variation that races cannot be sharply defined. I recognize five varieties
in Texas; a sixth occurs in Arizona.’

All have an extended blooming season from spring to fall. Two con-

trasting patterns of variation are shown by those in Texas, two varieties

having prevailingly broader, shorter leaves than var. Hartwegii, while
two are consistently very narrow-leaved.

ARTWEGII var. Toumeyi (Small) Shinners, comb. nov. Galpinsia oe

1)

or ET Torr. Bot. Club 25: 317. 1898. Ocnothera Heres var. Toumey: (Smal
Munz, Amer. Journ. Bot. 16: 708.

342

KEY TO VARIETIES OF C. HARTWEGII
la. Leaves (except lowest) abruptly narrowed to truncate or slightly
clasping at base, narrowly ovate or ovate-oblong to oblong or oblong-
elliptic 3c. var. pubescens
lb. Leaves ease ener | at Ge or senteniciy narrow throughout,
filiform or linear to lanceolate, oblanceolate, or elliptic-lanceolate
2a. Leaves 2%—5 times as long as wide, widely spreading to slightly
reflexed or occasionally ascending, sharply dentate or occasionally
sub-entire; plants of lower Rio Grande Plain (also northeastern
Mexico) b. var. Maccartii
2b. Leaves 540 ae as Sone as wide seeehainicsn or rarely spreading,
entire or occasionally sharply dentate (frequently so in Mexico);
plants found west and north of lower Rio Grande Plain
3a. Leaves linear to oblanceolate or elliptic-lanceolate, 1.3—13.0 mm.
wide, 5—20 times as long as wide (narrowest dimensions on sum-
mer or fall shoots of var. lavandulaefolius with dense, gray pubes-
cence)
4a. Leaves linear-lanceolate to oblanceolate or elliptic-lanceolate,
glabrous to sparsely pubescent (rarely rather densely and min-
utely pubescent with hairs up to 0.3 mm. long in plants from

outside Texas) . . da. var. Hartwegii
4b. Leaves linear or ree inecolete:. gray with dense, mostly
appressed hairs up to 0.6 mm. long . 3d. var. sae es tomee

3b. Leaves filiform to linear, 0.5—2.0 mm. wide, 12—40 times as
ong as wide, green, inconspicuously puberulent and glandular-
viscid; plants of gypsum outcrops, northern Trans-Pecos (also
adjacent New Mexico) . . ; . . 8e, var. filifolius
3a. C. HARTWEGII var. HARTWEGII. Oenothera Hartwegii Bentham,
Pl. Hartw. 5—6. 1839. From central Mexico; precise locality not known.
The entire original description is quoted below; the first two lines ap-
peared on em 5, the rest on p. 6

wo

OENOTHERA Hartwegii, sp. n., suffruticosa, humilis, de-
Saaiene: ‘oliis line Sar me us, Saas integris Vv. sinuato- Henratls
glabrius culis, calycix e libera ovario cylindrico sub-4-ies

longiore apice infunabeliform ae laciniis ovato-lanceolatis

glabri usculis longiuscule subulato-acuminatis, antheris stigma

4-partitum sequantibys.—Caly ix tubus ultra 2 poll. longus.

Corolla purpur

Oenother ceo Gray, Mem. Amer. Acad. 4 pt. 1 (Pl. Fendl.): 46. 1849.
Fruticulosa, ramossissima, erecta, undique minutissime viscido-puberula;:
foliis parvis (3—6 lin. longis) spathulatis vel oblanceolatis integerrimis
sessilibus seu in petiolum pl. m. attenuatis; floribus parvulis; tubo calycis
apice breviter obconico filiformi ovario sextuplo laciniisque triangulari-
lanceolatis cuspidatis petala rhomboidea subaequantibus quintuplo long-
iore; capsula sessili oblongo-prismatica.—Hill southeast of Pelayo, in
Chihuahua, Dr. Gregg; May, 1847. ‘A very small semi-shrub; flower yel-
low.’ The specimens are about 8 inches high, very bushy; the petals turn

343

to rose-color in drying, as in the allied species, and are one third of an
inch in length. Capsules scarcely half an inch long.” Munz (1929, pp.
709—710) makes these comments. “Gray’s type is the smallest and most
glabrate plant that I have seen, his variety pubescens being based on a
type more like the other plants I have included under Greggii var.
typica. But his var. pubescens is not worth varietal rank, the type of
the variety typica being pubescent but more minutely so. O. Greggii
var. typica intergrades freely with var. lampasana.”’ On the basis of leaf
shape, Greggti definitely belongs with var. Hartwegii, while pubescens
and lampasana belong together but not with var. Hartwegii.—Oenothera
Greggii var. Pringlei Munz, Amer. Journ. Bot. 16: 711. 1929. “Leaves and
stems strigose-canescent; leaves 1—3(4) cm. long, 1—3(4) mm. wide,
wavy-margined and denticulate.’ Type (not seen) from Bachimba

this with var. Hartwegii as I understand it.—Oenothera Fendleri Gray,
Mem. Amer. Acad. 4 pt. 1 (Pl. Fendl.): 45—46. 1849. ‘“Muinutissime
pulverulento-glandulifera, glabra; caulibus e radice lignosa decumben-
tibus; ramis brevibus adsurgentibus; foliis lanceolatis oblongisve sessili-
bus subintegerrimis; calycis tubo apice infundibulari-inflato ovario
prismatico sessili laciniisque triangulari-lanceolatis cuspidatis 3—4-plo
longiore; petalis rhombei-obovatis stylo paulo longioribus.—Sunny hill-
sides at Santa Fe, and on the Rio del Norte; also (chiefly a narrow-
leaved form) from Rock Creek eastward to the Cimarron River; May to
August,” Fendler 230. O. Hartwegit var. Fendleri Gray, Smithsonian
Contrib. 5 art. 6 (Pl. Wright. 2): 58. 1853.

This is the most widespread and the most heterogeneous variety. Un-
common in Texas; known from the Panhandle, east in the Red Plains
to Wilbarger Co., and from the Trans-Pecos, east to Uvalde and Val
Verde counties. Until late in my study I attempted to maintain var.
Fendleri as a more northern, more glabrous race with broader and more
entire leaves (despite the inclusion of narrow-leaved forms in the orig-
inal), but the separation proved far too weak to maintain. Stem glabrous
or variously pubescent with short hairs only. One specimen from Reeves
Co. (Lake Toyah, Cory 52099) has stems with short, erect hairs and some
medium long ones, approaching var. pubescens, possibly due to intro-
gression with the latter. A form with finely gray-pubescent leaves,
superficially resembling var. lavandulaefolius but with shorter hairs,
occurs in Mexico, well south of the range of the latter; I believe it should
be regarded as a homologous mutation. I have not seen Texas specimens
of this form.

3b. C. HARTWEGII var. Maccartii Shinners, var. nov. Folia plerumque
patentia vel subreflexa, petiolata vel basi angustata, laminis denticulatis
(rarius subintegris) oblanceolatis vel oblongo-lanceolatis pro ratione
brevibus (ca. 10—38 mm. longis x 2—8 mm. latis). HOLOTYPE: U.S.
Highway 83, 6 miles northwest of Rio Grande (City), Starr Co., Texas,

344

Rosa Ena Benavides 91, 24 March 1963. ‘In mesquite savannah.” Two
additional U.S. collections seen, both from Duval Co.: State Highway
44, 7 miles east of Freer, Rebecca M. Rodriguez 104, 18 March 1962.
State Highway 359, 10 miles southwest of Benavides, Elvira G. Garcia
113, 22 March 1963. Found also in nearby parts of Mexico. NUEVO LEON.
108 km. (65 mi.) no. (sic! ie. south) of Nuevo Laredo, on road to
Monterrey, I’. C. & E. M. Frye 2369, 19 April 1939. Highway 85, 45 miles
south of Nuevo Laredo, Juan G. Rivas, Platon Ostos & Wm. L. McCart
8133, 17 March 1962. Villaldama Road, 16 km. west of Sabinas Hidalgo,
Martha Dominguez M. & Wm. McCart 8255, 7 April 1962. Highway 85,
17 miles northwest of Sabinas, Juan Jorge Rodriguez 70, 20 March 1963.
TAMAULIPAS. 20 miles east of the International Highway, by the
Riberena Road, Lorenzo Escalante 55, 24 April 1962

Named in honor of William Larrey McCart, Head of the Science De-
partment, Laredo Junior College, for his long and continuing services
as an energetic collector of the Texas flora. Lest some hasty pedant
accuse me of misspelling, it should be stated that the extra a has been
added deliberately because it makes a better Latin form of the name.
Records of Oenothera Greggii var. Pringlei from the Rio Grande Plain
given by Munz (1944) probably belong here. The leaf dimensions he
gives, apparently quoted from the original description, do not apply,
but he notes that the Texas plants “are not quite so closely strigose, nor
so narrow-leaved as plants from Coahuila and farther south, but they do
approach the latter.”

3c. C. HARTWEGII var. pubescens (Gray) Shinners, comb. nov.
Oenothera Greggii var. pubescens Gray, Smithsonian Contrib. 3 art. 5
(Pl. Wright. 1): 72. 1852. “Pilis aoe patentibus villosa—Dry hills
beyond the Pecos; Aug.,” Wright 199. “Leaves oblong, 2—4 lines long.
Though Munz refers to this in his discussion of O. Greggii, already
quoted, he nowhere cites it in his lists of synonyms.—O. lampasana
Buckley, Proc. Acad. Sci. Phila. 13 (1861): 454—455. 1862. “Caule sub-
prostrato, glanduloso-pilosa; foliis numerosis, ovato-lanceolatis, integris,
muDSeSSNIDUS, acutis, glanduloso-pubescentibus . .. Prairies, Lampasas
County.” O. Greggu var. lampasana (Buckley) Munz, Amer. Journ. Bot.
16: 710. 1929

Stems with long, widely spreading hairs together with short, erect,
gland-tipped or glandular hairs and very short, incurved, glandless ones.
Occasional plants have few or no glandular hairs, or only short, erect
hairs; one specimen from Taylor Co, (east edge of Abilene, Norlan
Henderson 63-376) has every dense pubescence of short to medium-long,
mostly non-glandular hairs. Leaves usually short and rather wide; nar-
row-leaved forms might indicate introgression with other varieties, but
the others are so uncommon or localized within the area of var. pubescens
that spontaneous mutation seems a more likely explanation. This is much
the most common variety in Texas, from the Panhandle and Red Plains

345

to the West Cross Timbers (Erath Co.), south to the northern Edwards
Plateau, and southwest through the Trans-Pecos.

3d. C. HARTWEGII var. lavandulaefolius (T. & G.) Shinners, comb.
nov. Oenothera lavandulaefolia T. & G., Fl. N.A. 1: 501. 1840. “Plains of
the Platte, Dr. James! Nuttall! (‘near Scott’s Bluffs.’) ... Very nearly
allied to the Mexican OE. Hartwegii, Benth., which is a more glabrous
plant, with narrower leaves, a more slender calyx tube, and ee
acuminate segments.” O. Hartwegii var. lavandulaefolia (T. G.) S.
Watson, Proc. Amer. Acad. 8: 590. 1873. O. Hartwegii var. ae te
Munz, Amer. Journ. Bot. 16: 705. 1929. Calylophus lavandulifolius (T.

are actually hyphenated words from which the hyphen had been dropped;
they are not exactly the same as compound words, which the epithet here
would become if spelled lavanduwlifolius instead of lavandulaefolius.)

Relatively uniform in having very narrow, densely gray-pubescent
leaves (narrower than a majority of plants of var. Hartwegit, contrary
to the statement in the original description), and nearly always with
extremely short free tips to the sepals in bud, but these features are
hardly sufficient to justify regarding it as a distinct species. The most
northerly of the recognized varieties; as already noted under var.
Hartwegti, occasional gray-pubescent forms from Mexico are better re-
garded as parallel mutations under that variety and not properly placed
here. Frequent in the Texas Panhandle, south to Garza Co.; one record
from the northern Trans-Pecos (northern Culberson Co.).

3e. C. HARTWEGII var. filifolius (Eastwood) Shinners, comb. nov.
Oenothera tubicula var. filifolia Eastwood, Proc. Calif. Acad. (ser. 3) 1:
72. 1897. (This reference not seen; taken from Munz and the Gray
Herbarium Card Index.) Type from White Sands, New Mexico. O.
Hartwegii var. filifolia (Eastwood) Munz, Amer. Journ. Bot. 16: 707.

The most uniform of the accepted varieties, confined to gypsum out-
crops in the Trans-Pecos and immediately adjacent counties; specimens
seen from Culberson, Hudspeth, and Ward counties

REFERENCES
MUNZ, PHILIP A. 1929. Studies in Onagraceae IV. A_ revision a the subgenera Sal-
pingia and cia of the genus Oenothera, Amer. Journ. Bot. 16: 70
———————.. 1944. Onagraceae, Oenothera. In Pande 211, Flora of es 3 ae 221—

34
RAVE ae base . H. 1964. The generic subdivision of Onagraceae, tribe Onagreae, Brit-
tonia

TAXONOMY AND HETEROSTYLY OF
NORTH AMERICAN GELSEMIUM
(LOGANIACEAE)

WILBUR H. DUNCAN AND DONALD W. DEJONG
Department of Botany, University of Georgia, Athens

According to available floras the genus Gelsemium is represented in
North America by two species. One is G. rankinii Small which occurs
within 120 miles of the seacoast from North Carolina to Louisiana and
the other G. sempervirens (L) Ait. f., which is known in the United
States from Virginia to Arkansas and eastern Texas, Mexico, and Guate-
mala (Standley, 1924). Both species are woody evergreen vines with
attractive yellow funnelform corollas. Another species occurs in eastern

sia.

Although G. sempervirens was known by Linneaus (1753) and to
earlier authors as well (e.g., Gronovius, 1739), G. rankinii apparently
was not recognized in literature until Nuttall (1818) referred to a variety
of Gelsemium with inodorus flowers. Much later Small (1928) described
it as a species. The description and accompanying illustrations clearly
indicate the dimorphic nature of its flowers. Dimorphism in G. semper-
virens 18 well illustrated by Alexander (1929). The first reference to the
dimorphic nature of Gelsemium flowers that is known to us is that by
Walter (1788). In his description of G. sempervirens he states, “Varie-
tates, staminibus longioribus; stylo longiore.’”’ This may be the earliest
report of heterostyly for any species. The earliest observation of
heterostyly reported by Darwin (1884) is that by Persoon in 1794.

Some aspects of the dimorphism and the apparent intergradation of
certain diagnostic characters prompted the present study which included
extensive field research and breeding experiments as well as studies of
herbarium specimens. Problems attacked included the possible corre-
lation of certain morphological and cytological features with the species
and heterostylic type involved, some genetical aspects of the heterostyly,
the value of characters reported to be of diagnostic value, the distribu-
tion of the species, and the extent of any regional variation in the
species. Although some questions have not been answered it is appro-
priate to report what has been learned.

STUDIES OF GROSS CHARACTERISTICS
According to the descriptions and keys given by Small (1933) the two
species differ in several gross characteristics, i.e., those that ordinarily
are used in manuals and in descriptions of species. Our observations have
led to the following conclusions. The flowering shoots of G. sempervirens
are more likely to be green and those of the G. rankinii red-tinged, than

SIDA 1 (6): 346—357. 1964.

347

the reverse which is given by Small. The dilation of corolla tubes of the
two species is so frequently similar that this character is of little use
diagnostically. The body of the capsules of both species is generally
veined, in G. sempervirens slightly more veined, this apparently being
somewhat correlated with the size of the capsules. The species have
nearly the same range in length of corolla and corolla tube instead of
those of G. rankinii being clearly the shorter. The lengths of stamens
and pistils in pin types, and also in thrum types of flowers, are essentially
the same for both species.

Observations on many fresh flowers over a wide geographic range in
the field east of Mississippi indicate that G. sempervirens always has
odorous flowers and that G. rankinii usually does not. Several popula-
tions of the latter, however, have been found to have faintly to strongly
odorous flowers, a condition apparently not reported in literature. In
these populations the possibility of introgression with G. sempervirens
was considered but was concluded to be improbable because features
characteristic of G. sempervirens were mostly lacking on those plants
with odorous flowers. More information about the possibility of intro-
gression will be found later in our discussion of leaf base angles, pollen
sizes, pollen fertility, and chromosomal studies.

Leaf bases in G. sempervirens are reported by Small (1933) to be
narrowed at the base and those of G. rankinii rounded. Our data show
that this is only generally true and that the species often cannot be
separated by angles of the leaf bases. Angles were measured from special
collections from the field and supplemented by others from herbarium
specimens. The widest angle and the narrowest angle encountered on

for G. sempervirens was from 70° to 150° and for G. rankinii from

respectively. Those collections of G. sempervirens with any leaf base
over 110° were examined for other characters typical of G. rankinii.
None was found, the other characters definitely being those of G.
sempervirens. In the case of those collections of G. rankintw having a
maximum leaf base angle under 150° examination was made for other
characteristics of G. sempervirens. None was found except that three
collections with angles of 120°, 130°, and 135°, respectively, had odorous
flowers. This is of no great significance, however, for there are more
collections of otherwise typical G. rankinii having as small or smaller
angles of the leaf base and there were five odorous flowered collec-
tions having larger angles (to 180°).

Leaf length and width for the two species were also studied. Data
from over a hundred leaves of each species when averaged, plotted in
a scatter diagram, and otherwise analyzed, show that the leaves of
G. sempervirens average less in width and more in length, but only
generally have a larger length-width ratio than those of G. rankinii.

348

Ratios for the former were from 2.9 to 5.1 and for the latter 1.0 to 3.9.

Observations were made in the field during anthesis on the colors of
the corollas of fresh flowers of the two species. The two species could
often be distinguished on this basis, the colors of the flowers of G
rankinii usually being the darker. The colors for G. sempervirens were:
brilliant yellow (2.5Y-9/9 of the Nickerson Color Fan, published by the
Munsell Color Co., 1957) to moderate orange yellow (10YR-8/10). For
G. rankinit they were: vivid yellow (2.5Y-8/12) to strong orange yellow
(7.5YR-7/11). The inner part of the corolla tube of both species, and
especially of G. rankinii, was darker than the remainder of the corolla.

On the basis of the characters discussed above many herbarium speci-
mens or plants in the field, especially those without flowers or fruits,
would be difficult to place to a taxon. There also might be some doubt
about maintaining two species. Such doubt is dispelled by other gross
characters, these of the flower and fruit. The sepals, which appear not to

united, are obtuse to broadly pointed in G. sempervirens and acute
to usually acuminate in G. rankinit. Pedicels in the former are scaly
throughout (an occasional one may be partially naked), whereas in the
latter the upper part of the pedicel is naked (an occasional pedicel will
have a single scale reaching the base of the calyx, this being less fre-
quent for fruits and mature flowers than for flowers just having
opened). The body of the mature fruit of the former is 14.0 to 23.0 mm
long and 8.0 to 11.0 mm wide, the beak being 1.3 to 3.0 (5.4) mm long.
For the latter the data are 9.0 to 12.5 mm, 5.5 to 8.0 mm, and (2.4) 3.0
to 4.3 mm, respectively. Beak measurements for G. rankinii under 3.0
mm were fruits of the previous year, the shortness apparently being
due to disintegration during the winter. For G. sempervirens measure-
ments over 3.0 mm are uncommon and include a part or all of the upper
portion of the style. This portion apparently falls off later as a unit,
for beaks were either under 3.1 mm or over 5 mm long. In the latter
case a region of dehiscense was usually evident. The seeds readily dis-
tinguish the two species for those of G. sempervu ens are winged and
those of G. rankinii not.

The possibility of correlation of pin and thrum flower types with gross
characters was also studied. The characters included length of corolla
and corolla tube, color of corolla, leaf dimensions, and fruit dimensions.
No correlation was

DISTRIBUTIONAL STUDIES

Observations in the field and data accompanying herbarium specimens
show that G. rankinit grows in moist to wet situations and is confined
to the Coastal Plain of the southeastern United States (Fig. 1). Although
G. sempervirens occasionally occurs in moist situations, it only rarely
grows in wet habitats. In the southern United States it is found abund-
antly in drier situations, including shallow soils of granite outcrops and
being associated with scrub oak vegetation of sandhills as far inland

G. RANKINII

Fig. 1. TOP. Distribution of Gelsemiuwm rankinii as indicated by her-
barium specimens. Insert: Drawings of somatic metaphase chromosomes
in a root cell of G. rankinii. BOTTOM. Distribution of G. sempervirens
as indicated by herbarium specimens.

350

as the Fall Line. This species has a much wider geographic distribution
than the former. The known distribution except for Mexico and Guate-
mala is also given in Fig. 1. Records for Alabama and Virginia include
those reported by Harper (1928) and Massey (1961), respectively. G.
sempervirens has been reported for Oklahoma by Vines (1960). We
made no special effort to locate specimens to authenticate this report,
but Gelsemiwm is not included in Waterfall’s (1960) flora of Oklahoma.
We believe, therefore, that the genus does not occur in that state. Coulter
(1891) includes this species in his Botany of West Texas (west of 97th
meridian), but we find no specimens to verify its occurrence in that
region. Data from Standley (1924) and Martinez (1959) place G.
sempervirens in Mexico (states of Chiapas, Oaxaca, Puebla, Veracruz)
and Guatemala. The northernmost station appears to be Hanover Co.,
Virginia,

EPIDERMAL STUDIES

Microscopic studies were made of the lower epidermis of the two
species by using herbarium mounting plastic in a technique very similar
to that described by Sinclair and Dunn (1961) in their method A. The
number of stomates per .0926 sq mm varied from 20 to 34 in G. semper-
virens and 34 to 46 in G. rankinii. These data were based on the average
of two counts made from peels from a leaf from each of twenty speci-
mens of each species. Stomatal size was evaluated by measuring ten
imprints on a plastic peel from a leaf from each of twenty specimens of
each species. Distances measured for length and width values were be-
tween the ends of the guard cells and between their distant sides. These
data are presented in Fig. 2, a scatter diagram. It is obvious that the
measurements of stomates of G. sempervirens are usually the larger.
There was no significant difference in respect to measurements of size
and density of stomates between plants bearing pin and thrum type
flowers.

Larger stomatal size and a lesser number of stomates per unit of area
have been shown by Celarier and Mehra (1958), Stone (1961), and others
to be associated with a higher degree of polyploidy when comparing
species of a given genus. Our guard cell data, therefore, suggest that
G. rankinit has a lesser number of chromosomes than G. sempervirens.

The surface of the lower epidermis, much more abundantly in G.
rankini than G. sempervirens, appears to be finely grooved in lines
oriented along the sides of the stomates. These lines and other features
of the epidermis need further studies which should include cross sec-
tions of leaves.

STUDIES OF POLLEN
The sizes of pollen grains have been known to differ in pin and thrum
types of flowers at least since Darwin’s (1862) studies of dimorphism in
Primula. Also, larger sizes of pollen grains have been shown by many
to be associated frequently, although not invariably, with a higher degree

e a
@ e ms)
mM
. L
e ee &
e
e ee oO
rm =
e e e &
-
e ©
@ ° :
e e L
fe) Zz
e e fe) - wo
@ 6 fe} nN
a Li
Oo Oo
O 9 O —I
° o J
(e) e) —
(e) (@) o) - o
ce) re) i
fe)
ant Coane | T qT qT T T qT qT T T qT ‘ T LJ
w (e) wo
~N N —

Ww | D TH

Fig. 2. Scatter diagram of measurements of stomatal size in Gelsemium
rankinii (circles) and G. sempervirens (dots).

352

of polyploidy. A recent study describing such a situation is that of Stone
(1963) in a study of Carya species. A study of pollen sizes in Gelsemium,
therefore, seemed desirable.

The pollen studied was obtained from dried specimens from herbaria
and supplemented by specimens collected for this purpose in the field and
dried in presses over electric driers along with other herbarium speci-
mens. A single anther was removed from a flower of each collection to
be studied, dissected in water, and then observed under a microscope at
430X. The diameters of ten apparently normal pollen grains were re-
corded for each collection.

The data obtained on pollen size are summarized in Table 1.

TABLE 1. AVERAGE POLLEN SIZESa
a

Flower Type G. rankin G. sempervirens
Pin

No. of samples 21 30

Minimum 31.3 34.0

Maximum 36.7 42.2,

Mean 34.3 38.1
Thrum

No. of samples 20 29

Minimum 34.4 35.7

Maximum 43.9 45.6

Mean 40.8 41.5

“Expressed as diameter in microns, Based on measurements of ten pollen grains from each
collection.

It may be seen that the average diameter measurements of pollen of
thrum type flowers are larger than those of pin type, a situation re-
ported by Ray and Chisaki (1957) for two species of Amsinckia, Trans-
lating the measurements into volume of the pollen, that of the thrum
would be 67% larger than that of the pin in G. rankinii and 29% in G.
sempervirens. These volume differences between thrum and pin types
might be associated with the different distances the derived pollen tubes
would need to grow to allow fertilization of the eggs.

A comparison of average measurements of pollen for the two species
(Table 1) shows that they are larger for G. sempervirens in both the
pin and the thrum types of flower. Pollen size data, therefore, as well
as stomatal size and density, indicate a lower number of chromosomes
for G. rankinii,

Shape differences in pollen grains also have been known to be associ-
ated with heterostyly, e.g., Johnston (1952). In both species of Gelsemium
the pollens seemed to be of uniform size except for shrunken grains.
Counts were made of the number of these per hundred grains. The
numbers varied from 0 to 70 and were not associated with style type.
Since shrunken grains are indicative of abortion and abortion is often
present in hybrids, we investigated the possibility of relation of the

353

numbers of such grains to possible hybridization between the two species.
The specimen with 70% ‘aborted” pollen was of G. sempervirens. Al-
though this was from a locality only a few miles from where G. rankinii
was known to grow, the specimen exhibited no other evidences of
hybridization between the two species. It was also noted that relatively
high percentages of ‘aborted’ pollen were frequently found on speci-
mens of G. sempervirens that were from localities a hundred miles or
more from where G. rankinii is known to occur. The evidence from our
pollen studies, therefore, does not indicate that hybridization occurs be-
tween the two species.
PHENOLOGICAL STUDIES

There is evidence that the flowering periods of the two species of
Gelsemium differ. Small (1928) stated that Mr. H. A. Rankin wrote that
in the vicinity of Hallsboro, N. C., G. rankinii bloomed 20 days later
than G. sempervirens. On 30 Mar. 1963 in Echols and nearby counties in
Georgia the senior author observed the latter species to be nearly past
flowering while the former was just approaching maximum flowering.
During the Spring of 1964 G. rankinii plants transplanted to the senior
author’s premises did not begin to blossom until after those of the other
species planted there had ceased flowering.

When data from herbarium specimens throughout the entire range
of both species were analyzed no appreciable difference in flowering
time was discernable. When data were confined to specimens from areas
where both species occur (omitting the small disjunct area in North
Carolina), it was found that the Spring flowering period of G. semper-
virens was from 1 Feb. to 1 Apr. and G. rankinii essentially from 16 Feb.
to 19 Apr. One specimen of the latter species in flower on 29 Jan. was
collected about 10 miles south of St. Augustine, St. Johns Co., Fla. This
early flowering date may be associated with the close prone of the
plants to the ocean. Both species occasionally flowered in October and
there is one record for G. rankinii on 27 Sept. A plant of G. sempervirens
brought in from the woods nearby and cultivated in Chapel Hill, N. C. is
recorded (specimen No. 31472 in U.N.C. Herbarium) as blooming every
Fall, almost as abundantly as in the Spring. Herbarium records indicate
that from 30° 30’ southward, flowering of G. sempervirens begins 1 Feb.
and is essentially completed by 1 Apr. In the Coastal Plain of Ga. and
Ala. the period is from 19 Feb. through 14 Apr. In S. C., N. C., and Va.
and in the Piedmont of Ga. and Ala. flowering occured almost entirely
between 16 Mar. and 1 May.

Attention was also given to the possibility of differences in flowering
time between pin and thrum types of flowers. Field observations and
analysis of data on herbarium specimens gave no evidence of any differ-
ence in the flowering time of these two types.

GENETICAL STUDIES
Several years ago plants of G. sempervirens from two widely separated

354

colonies in Clarke Co. and from one colony in Oglethorpe Co., Georgia
were transplanted for observation to the senior author’s premises east
of Athens, Georgia. Plants from all three colonies grew in close proximity
and have flowered yearly since 1955. All had pin type flowers. No other
Gelsemium plants are known nearer than four miles. None of the plants
planted east of Athens produced fruits for eight years. On 1 April 1963
pollen from thrum type flowers was placed on the stigmas of about 50
pin type flowers of these plants. That Fall many more than 50 fruits
were harvested, several of these from plants whose flowers had received
no introducd pollen from the hands of the senior author. Presumably
insects had carried the introduced thrum type pollen from stigma to
stigma. No artificial pollinations were attempted during 1964 and again
no fruits developed.

Also on 1 April 1963 branches bearing thrum type flowers in a colony
of G. sempervirens south of Athens were carefully pruned of all opened
flowers and developing fruits and enclosed in plastic bags. On the follow-
ing day bags were removed and opened flowers were tagged and treated
as follows: five were self pollinated, and ten received pollen from pin
flowers of the isolated plants east of Athens. The bags were replaced
and left on the branches until the corollas of the pollinated flowers had
dropped off. That Fall six fruits had developed on flowers that had re-
ceived pollen from pin type flowers and no fruits on the self-pollinated
thrum type flowers.

We believe that the above data indicate that pin type plants of G.
sempervirens are self sterile in nature and cross fertile to pollen from
thrum type flowers, and possibly that thrum type flowers are self sterile
and cross fertile. The latter two conclusions need to be checked b
further experiments. The senior author plans to grow progeny of the
isolated plants that had received pollen from the thrum type flowers in
order to determine ratios of the F, generation for that cross. It is hoped
to make additional reciprocal crosses and successful selfing experiments
in order to determine the genetics of heterostyly in the two species of
Gelsemium.

Some studies were also made of the ratio in nature of plants bearing
pin and thrum type flowers. Several large population samples taken
in limited areas gave quite varied results, the flowers sometimes being
largely of one type or the other. Since Gelsemium reproduces abundantly
vegetatively, a sampling was made in the field at broader intervals. No
sample was taken nearer than a mile to another. They were made 27-30
March 1963 from McDuffie and Baldwin Cos., Ga. south to Duval and
Suwanee Cos., Fla. For G. sempervirens there were 34 pin type and 38
thrum type flowers and for G. rankinii 10 pin and 9 thrum type. These
indicate a 1:1 ratio.

Counts from herbarium specimens strongly indicate the same ratio.
Specimens borrowed from other herbaria were utilized and all duplicates

399

eliminated. All flowering specimens of G. rankinii were tabulated as
to type. The results were 21 pin and 20 thrum type (see Table 1). Counts
from herbarium specimens of G. sempervirens gave 111 pin and 108
thrum type. Nine pin type specimens were omitted from the above num-
ber. They were the nine available collections from Orange Co. NC. all
of which had pin type corollas. After studying the accompanying her-
barium labels and discussing the data with someone familiar with the
localities involved, we concluded that all nine specimens were most likely
propagated from the same clone.

As has been pointed out by Crosby (1949), Ray and Chisaki (1957),
and others, a 1:1 ratio, or close to it, indicates self sterility of pin and
thrum type flowers and an entirely outcrossing population. Our studies
of ratios, therefore, confirm the conclusions of our breeding studies that
both pin and thrum type flowers are self sterile and are cross fertile.

CHROMOSOMAL STUDIES

A chromosome number of 2n=16 has been reported for G. semper-
virens by Moore (1947). After vain attempts to repeat Moore’s method
using leaf tips, root tips were taken of rooting sections of stems and the
2n number of 16 was verified for this species by using a Feulgen squash
method.

The chromosome number for G. rankinii is unreported in literature.
The Feulgen squash method was tried with inconclusive results on root
tips from roots of layered stems of this species. It was, however, deter-
mined that active mitotic division occurred about 11:00 P.M. At this
point in the studies we turned to cross sectioned root tips. Root tips were
placed in a modification of Navashin’s Fluid Fixitive described by Sass
(1958), sectioned, stained by a modification of Newton’s Gentian Violet-
Iodine method described by Johansen (1940), and mounted in balsam.

From root tip material collected during the latter part of June, 1964,
mitotic metaphase counts of 2n=8 were obtained. A voucher specimen
(Duncan 22020) is on deposit in the University of Georgia Herbarium
(sheet No, 74279). A sample chromosome plate is shown in the inset of
Fig. 1. The chromosomes are approximately 1 to 3 wu in length. Occasion-

ally cells with approximately 16 (in the outer three rows of root cells)
and rarely with 9 or 10 chromosomes were seen. Moore (1947) reported
for G. sempervirens chromosomes 1.3-3 jb in length and occasional poly-
ploid cells and cells with intermediate numbers in leaf smears.

Indications from stomatal and pollen data presented ane that G.
rankinii had a smaller number of chromosomes are thus confirmed by
the determination of a 2n number of 8

CONCLUSION

Our studies show that there are two distinct species of Gelsemium in
North America and that there is probably no introgression between them.
Hybrids were not detected and should not be expected in abundance
since one species is diploid and the other is tetraploid.

356

The determination of the chromosome number of 2n=8 for G. rankinii
lowers the known n numbers for the Longaniaceae and the basic number
for Gelsemium to 4. Darlington and Wylie (1955) had reported the basic
number for Gelsemium as 8. Moore (1947) had previously cited a report
of n=6 for two species of Fagraea which belongs to the same subfamily
as Gelsemium. He suggested on the basis of cytological evidence that
Gelsemium might have a genetic link with one branch of the Apocynaceae
which are characterized by a haploid number of 8. Moore also suggested
that Gelsemium may have diverged from an ancient line which pro-
duced the Apocynaceae. The basic chromosome number of 4 for Gelsem-
ium seems to support Moore’s suggestion that divergence may have
occurred at such a distant time that Gelsemium can not be regarded, on
morphological grounds, as a true member of the Apocynaceae.

Our cytological evidence indicates that G. sempervirens was derived
more recently and probably from G. rankinii. Distributional data in the
southern United States also indicate such a relationship. G. rankinii is
confined to wet habitats of a limited area of the Coastal Plain (Fig. 1),
while G. sempervirens, being a tetraploid and probably more vigorous,
occurs in a variety of habitats (wet to very dry) and occupies a much
wider area (Fig. 2). The reported occurrence of G. sempervirens in the
disjunct Mexican-Guatemalan region and the apparent absence of G.
rankint from there could be taken to indicate that the former species
is the ancestral one. Before serious conclusions involving distributions
in the Mexican-Guatemalan region are made, however, the identity of
all collections of Gelsemiwm from there should be checked and the
absence of G. rankinii verified by additional studies. Most persons deal-
ing with the flora of that region are probably unfamiliar with G. rankinii.

Although pin type plants are generally heterozygous and the thrum
type homozygous recessive (Ray and Chisaki, 1957; et al.), we have no
evidence that this is the case in Gelsemium. Additional breeding experi-
ments are needed to determine the genetic makeup of the two forms as
well as the extent of self sterility, especially for thrum type plants.

Our studies also have added to those known situations in which
stomatal size and density, and size of pollen are correlated with ploidal
levels.

We wish to thank those curators who permitted examination of collec-
tions at his herbarium, loaned specimens to us, and provided information
for the study. The junior author, who worked on this study for ten
weeks during the summer, 1962, received support from National Science
Foundation Grant (G-20468). Martha Gordon and William L. Cleeg, who
made studies of the epidermis and pollen, received support from National
Science Foundation Grant (G-20296). The help of Wesley Walraven in
making chromosomal studies of G. rankinii is appreciated.

357

REFERENCES
ALEXANDER, EDWARD J. 1929. Gelsemium sempervirens, Addisonia 14: 8.
CELARIER, ROBERT P. AND K. L. aes 1958. Determination of polyploidy
from acini Pa Rhodora 60: 89-
HN M. 1891. Botany of ok Texas. Contr. U. S. Nat. Herb. 11: 271.
Sea l ‘4 1949. Selection of an unfavorable gene complex. Evolution 3: 212-230.
D. A

ee mregae ON, C. D. AND A. P. WYLIE. 1955. Chromosome Atlas
N, CHARLES D. 1862. On the — or sae aes condition in the species
of aa and on their eomigelable sexual aes s. J. Linn. Soc. Bot. 77.

DARWIN, CHARLES D. 1884. The oe Forms pe Flowers on te of the Same
Species.

GRONOVIWUS, J. F. 1739. Flora Virginica,

HARPER, ROLAND M. 1928. Economic Botany of Alabama. Part 2 Catalogue of oe
trees, ae and vines of Alabama. Geol. Sur. of ‘Alabama Monogr. 9. University, Alabam

J SEN, D. A. 1940. Plant Mice

ON, IVAN M. 1952. a in oe Boraginaceae, XXIII. A survey of the genus

aonen J. Arnold Arboretum 33

LINNAE . 1753. Species eee

MARTINEZ MAXIMINO. 1959. Las Plantas Medicinales de Mexico.

MASSEY, A. B. . Virginia Flora. Tech. Bull. 155. Va. Agr. Exp. Sta.
RE, R o J. 1947. Cytotaxonomic as in the Loganiaceae. | Chromo-
some nuwbers and phylogeny in the Loganiacea r. J. . 34: 527-538

NUTTAL HOMAS. 1818. The Genera of ak American Plants and a catalogue of

the sae ce the year 1817.
PETER M. AND HARU F. CHISAKI. 1957. Studies on pie I. A synopsis
of the genus with a study - heterostyly in it. Amer. J. Bot. 44:
ASS, . E. 1958. a ee Microtechnique, p. 18.

ee , CL B. AND DAVID B. DUNN. lade Surface printing of plant
leaves ae lowenevic ee Stain Technol. 36:

SMALL, J. K. 1928. Gelsemium Rankinii, Addisonia 13: 37-38.

SMALL, J. K. 1933. Manual of the Southeastern Flor

Sree PAUL C. 1924. Trees and Shrubs of Mexico. Contrib, U. S. Nat. Herb.
23: 1141-114

STONE, D NALD E. 1961. Affinities of a Mexican endemic, Carya palmeri, with
American and Asian hickories. Amer. J. Bot. 49: 199-212

STONE, DONALD E. 1963. Pollen size in hickories (Carya). Brittonia 15: 208-214.

VINES, ROBERT A. 1960. Trees, Shrubs and Woody Vines of the Southwest.

WALTER, THOMAS. 1788. Flora Caroliniana.

WATERFALL, U. T. 1960. Keys to the Flora of Oklahoma.

TEXAS ASCLEPIADACEAE OTHER
THAN ASCLEPIAS
LLOYD H. SHINNERS
Herbarium, Southern Methodist University, Dallas, Texas 75222

Woodson’s drastic reduction in the number of genera of Asclepiadaceae
in 1941 was certainly welcome. But for the botanist having to make
routine identifications or attempting local or regional studies within the
United States, the years since have been a time of continuing frustra-
tion. Only the original, highly technical key to the revised genera
using inconvenient pollen characters, has been available; the genera
themselves were listed without descriptions, and only two (Asclepias
and Sarcostemma) have been monographically treated; several needed
nomenclatural changes have remained unpublished: and, for Texas in
particular, a new species and a new variety have gone undescribed. This
paper has been prepared in order to clear up some of the loose ends and
to provide a working guide for identification. It is based chiefly on
collections in the S.M.U. Herbarium and in that of the University of
Texas; for the loan of the latter I am indebted to Drs. B. L. Turner and
Marshall C. Johnston. Some additional material was examined on visits
to the Missouri Botanical Garden, and several critical specimens were
kindly loaned by Mr. Fred B. Jones of Corpus Christi, Texas, from his
personal herbarium.

Supplementing the account of the Texas representatives, I have added
some routine new combinations for plants found outside the state, a key
to Southeastern Cynanchum (owing to lack of adequate flowering ma-
terial I have not yet completed one for Matelea in the same region;
Sarcostemma clausum (Jacquin) Roemer & Schultes in southern Florida
is the only Southeastern member of that genus), and a finding-list of
generic names. The key to genera given below can also be used for
the Southeast except that the introduced Cryptostegia grandiflora R.
Brown, established in southern Florida, must be added. This is immedi-
ately separable from all the other genera by its extremely large flowers,
with funnelform corolla 4—6 cm. long; in the rest the corolla (or its
lobes, if spreading or reflexed) is 0.2—2.0 cm. long.

KEY TO GENERA
la. Stamen column or its base surrounded by 5 separate, fleshy-inflated
or fleshy-thickened, erect or spreading appendages

2a. Stems prostrate to erect, not twining; base of corolla not with

fleshy disk under the separate appendages
3a. Leaves not both cordate and petioled; corolla green to white,
yellow, orange, red, brown, or purple; wild or cultivated
1. Asclepias

SIDA 1 (6): 358—367. 1694,

359

3b. Leaves both cordate and petioled; corolla lavender-blue; culti-

vated F i 2. Oxypetalum
2b. Stems iene: ee tea jowara dips: eororl oh. a fleshy disk at
base under the appendages . 4. Sarcostemma

lb. Stamen column or its base with 1 or 9 rows of flat, thin appendages,
or a single, entire or lobed, fleshy disk or cup
4a. Appendages thin and flat, in 2 rows, or a single, entire or lobed,
fleshy disk or cup ie ee a . oO. Matelea
4b. Appendages thin ad flat, in 1 row
5a. Corolla funnelform or campanulate, 2.0—6.2 mm. long; wild her-

baceous vines (sometimes weeds in gardens) . . 3. Cynanchum
5b. Corolla rotate, its narrowly oblong lobes about 10 mm. or more
ong; cultivated woody vine . . . . . 6. Periploca

1. ASCLEPIAS, with about 32 species, will not be discussed further
here; no new names are required for Texas representatives, so far as
known. 2. OXYPETALUM has only one infrequently cultivated species
in the state, O. caeruleum Decaisne, with densely soft-pubescent leaves;
native of Argentina.

3. CYNANCHUM.

Small to large twining vines. Corolla white to yellowish or yellow-
green, rather small. Five species.
la. Leaf blades with cordate base

2a. Appendages nearly as long as the corolla, deeply divided into

linear segments (resembling staminodes) . . . 1. C. laeve
2b. Appendages iess than 2/3 as long as the seule Guaacis oblong
with toothed or lobed summit . 2. C unifarium

lb. Leaf blades with narrowed to sounded: Gane ase

3a. Flowers rather numerous, terminating naked peduncles longer

than the pedicels; corolla lobes glabrous within; leaf blades linear-
lanceolate, the larger 4—9 cm. long . . 3. C. palustre
3b. Flowers solitary or few, peduncles very short or absent; corolla
lobes pilose or pubescent within; leaf blades lanceolate to oblong-
elliptic, 1—4 cm. lon
4a. Corolla lobes conspicuously pilose within; appendages lance-
linear to linear-filiform, 112—2 times as long as the stamen column
5a. Corolla 3.6—5.2 mm. long . 4a.C. barbigerwm var. barbigerum
5b. Corolla 2.8—3.2 mm. long . 4b. C. barbigerum var. breviflorum
4b. Corolla lobes rather minutely pubescent within; appendages nar-
rowly lanceolate to ovate-acuminate, slightly longer than the
stamen column . . 5. C. Maccartit

1. C. LAEVE naa Persoon. Enslenia - albida Nuttall. Ampelamus
albidus (Nuttall) Britton. In a north-south belt a little east of the cen-
ter of the state, from Clay, Cooke, and Grayson counties south to
Matagorda County. Flowering August—September. In Gould’s Texas
Plants this is listed both as Ampelamus and as Cynanchum, and assigned

360

two entirely different distributions. It is absent from the extreme eastern
part of the state, where limestone is absent. Its preferred natural habitat
is low ground in limestone areas; it is frequently a weed in flower beds.

2. C. UNIFARIUM (Scheele) Woodson. Rouliniella unifaria (Scheele)
Vail. Including Roulinia Palmeri S. Watson, Cynanchum Watsonianum
Woodson. The slight difference in size and toothing of the appendages
hardly justifies recognition of a second species. Very similar in general
appearance to the preceding. Edwards Plateau to Trans-Pecos and Rio
Grande Plain, north to Parker and Taylor counties, southeast locally to
Brazos (in shrubbery on Texas A. & M. campus, possibly introduced),
Bastrop, and San Patricio counties; also in northeastern Mexico. Flower-
ing mid May—October. In alluvial habitats, like C. laeve, but also in
drier ground, often in rocky or sandy soils.

TRE (Pursh) Heller. Lyonia palustris (Pursh) Small.
ee palustris (Pursh) Vail. Local along the Gulf Coast; specimens
seen from Aransas, Galveston, and Kenedy counties. Flowering April—
September.

4. C. BARBIGERUM (Scheele) Shinners, Field & Lab. 19: 65. 1951.
Metastelma barbigerum Scheele. Type from New Braunfels, Comal Co.

a. C. BARBIGERUM var. BARBIGERUM. Common on the southern
Edwards Plateau from Travis, Llano, Mason, and Terrell counties south,
and on the Rio Grande Plain, east to Karnes and Refugio counties; ap-
parently rare in the Trans-Pecos (Brewster Co.), but the two specimens
seen from that area, with somewhat small flowers (corolla 3.6 and 3.7
mm. long), may prove to be only exceptional forms of the next variety.
Also in northeastern Mexico. Flowering March (in extreme south) or
April—September.

4b. C. BARBIGERUM var. breviflorum Shinners, var. nov. Corolla

r 2.8—3.2 mm. longa (vice 3.6—5.2 mm.). HOLOTYPE: Big Bend
National Park, Chisos Mountains, granite peak in center of Basin, alt.
5500 ft.; common, twining over low shrubbery; corolla white, Grady L.
Webster 4340, 15—19 July 1952 (SMU). Largely if not wholly replacing
var. barbigerum in the Trans-Pecos, mainly in igneous rock areas; also
in Chihuahua. Flowering June—August.

9. C. Maccartii Shinners, nom. nov. Based on Metastelma Palmeri S.
Watson, Proc. Amer. Acad. 18: 115. 1883. Cynanchum Palmeri (S. Watson)
Shinners, Field & Lab. 19:65. 1951. (Not C. Palmeri (S. Watson) Blake,
1917, based on Pattalias Palmeri S. Watson.) Very similar in general
appearance to C. barbigerum, especially the small-flowered var. brevi-
florum. Type collected “at Laredo on the Rio Grande.” Rather rare, Rio
Grande Plain (Duval Co.) north and west to Uvalde and Val Verde
counties; also in northeastern Mexico. Named for William Larrey McCart,
Head of the Science Department, Laredo Junior College

. SARCOSTEMMA.
Small to moderately ince twining vines (small plants twining only

361

at tips of stems). Flowers umbellate, terminating naked peduncles.
Corolla shallowly campanulate, medium large (lobes 6—11 mm. long),
greenish to creamy white, pink, purple-green or purple. Three species.
la. Sepals narrowly lanceolate, more than 3 times as long as wide

1. S. erispum
1b. Sepals lanceolate to ovate, less than 3 times as eae as wide

2a. Sepals 4—6 mm. long, pubescent on both surfaces; stems usually

densely pubescent . 2.8. Torreyi
2b. Sepals 2—3 mm. ina ubeceent on pee ene stems globrous or
sparsely pubescent
3a. Leaf blades (except saan 1—3 times as long as wide
_ S. eynanchoides var. cynanchoides
3b. Leaf blades 312 times as tone as wide
.S. cynachoides var. Hartwegu

1. S. CRISPUM Bentham. ae S. lobatum Waterfall, Rhodora
51: 58. 1949. tinge crispa (Bentham) Vail. Fundstrum crispum
(Bentham) Schlech labrous or inconspicuously pubescent. Leaf
blades narrowly aan lanceolate with deeply cordate base, varying
to linear-lanceolate with abruptly narrowed base, the margin usually
(but not aways) ruffled or crisped. Frequent in Trans-Pecos, occasional
east and northeast on Edwards Plateau to Travis and McLennan coun-
ties, in the Panhandle, Red Plains, and West Cross Timbers (Callahan
and Palo Pinto counties); collected at West Dallas by Reverchon, noted
as “local and very rare,” not found there recently. Flowering late April—
early August.

2. S. TORREYI (Gray) Woodson. Philibertella Torreyi (Gray) Vail.
Funastrum Torreyi (Gray) Schlechter. Trans-Pecos, rather rare; known
from Brewster and Presidio counties, Flowering June—August. Very
similar in general appearance to S. cynanchoides var. cynanchoides, with
slightly larger flowers.

3 S CYNANCHOIDES Decaisne. Philibertella cynanchoides (Decaisne)
Vail. Funastrum cynanchoides (Decaisne) Schlechter. The commonest
species, with two intergrading varieties.

3a. S. CYNANCHOIDES var. CYNANCHOIDES. Leaf blades tri-
angular-ovate with cordate base. Frequent from Trans-Pecos to lower
Rio Grande Plain, Edwards Plateau, Panhandle, and Red Plains, rare in
West Cross Timbers (Parker) and along Red River to Grayson County.
Flowering June—September.

3b. S. CYNANCHOIDES var. Hartwegii (Vail) Shinners, comb.
Philibertella Hartwegii Vail, Bull. Torr. Bot. Club 24: 308. ne
Sarcostemma cynanchoides ssp. Hartwegiit (Vail) R. Holm, Ann. Mo.
Bot. Gard. 37: 530. 1950. The epithet heterophyllum has been applied to
this plant, in various combinations; according to Dr. Holm, its type
specimen is actually a form of S. crispum. Leaf blades lanceolate to

362

linear with an abruptly wider hastate or cordate base, or without wider
base. Frequent in the Trans-Pecos. Flowering April—September.

5». MATELEA.

Plants herbaceous, prostrate to suberect and rather small, or small to
large twining vines, nearly glabrous or variously pubescent or pilose.
Flowers small to medium large; corolla green to yellowish, brown-red,
or purple-brown. Eleven species.
la. Stems prostrate to suberect, not at all twining

2a. Peduncles absent (pedicels attached directly in leaf axils): stamen

column with a single, lobed, fleshy disk around base
3a. Pedicels shorter than or equalling the adjacent petioles
. |. M. biflora
3b, Pedicels (except lowest) exceeding the scaacest petioles
. cynanchoides
2b. Peduncles well-developed; stamen olen ete by a double
row of thin appendages
4a. Outer appendages wider than long, slightly shorter to slightly
longer than the stamen column, truncate to shallowly 3-lobed
at summit . a 5 3. M. brevicoro
4b. Outer ee ee ienieee fan ide Sn tie to much longer me
the stamen column, prominently 2-pronged or 2- pointed at summit
arely some of them single-pointed . . . . 4. M. parviflora
lb. Stems twining, at least toward tips
da. Flowers at middle and upper leaf axils on peduncles shorter than
the pedicels, or without peduncles; plants small, semi-trailing or
low-climbing
6a. Corolla lobes 3—4 mm. long . ... . . . 5. M. parvifolia
6b. Corolla lobes 7—12 mm. long
7a. Peduncles absent; flowers solitary or paired, short-pedicelled
8a. Crown (appendage around stamen column) saucer-shaped,
entire; Trans-Pecos mountains (Jeff Davis Co.)
. 6. M. sagittifolia
8b. Crown cup-shaped or short- inane is margin 5-parted;
Rio Grande Plain west to Val Verde Co. . . 7. M. Woodsonii
7b. Peduncles present except in uppermost leaf axils, 1—5 flowered
8. M. producta
ob. Flowers all on elongate peduncles; mein: to saaaee climbing vines
9a. Corolla lobes oblong-lanceolate to linear, not reticulate-veined
10a. Sepals glabrous or sparsely hispid . . . . 9. M. gonocarpa
10b. Sepals both hispid and short-pubescent . . 10. M. decipiens
9b. Corolla lobes ovate, finely reticulate-veined on upper surface
: . M. reticulata

1. M. BIFLORA (Rafinesque) Woodson, ieee Guillen inn (Rafi-
nesque) Heller. Common on the Blackland Prairie of north central
Texas, west and south to Lubbock, Sutton, Travis, Gonzales, and Bastrop

363

counties; on clayey, rocky, or less often sandy soils. This is another of
the species originally described from “Arkanzas,” meaning the Arkansas
Territory, actually collected in present Oklahoma, persistently credited
to the present state of Arkansas, where so far as I know it does not
occur. Flowering April—June, rarely September.

2. M. CYNANCHOIDES (Engelmann) Woodson. Vincetoxicum cynan-
choides (Engelmann) Heller. In northern and central Texas, from west-
ern part of Pine Belt (Upshur Co.) west to West Cross Timbers (Young
Co.), south to Goliad Co., in sandy soil; frequent. Flowering April—
August.

3. M. BaEvE ‘ORONATA (B. L. Robinson) Woodson. Gonolobus par-
viflorus var. brevicoronatus B. L. Robinson, Vincetoxicum brevicoro-
natum (B. L. Robinson) Vail. Type collected at Laredo by Pringle. Rare,
in lower Rio Grande Plain, in sandy or gravelly soils; specimens seen
from Hidalgo, Kenedy, and Webb counties. Flowering March—Septem-
ber. Found wholly within the range of the next species, and distinguisha-
ble from it only by the appendages within the flower. Robinson says
that it also differs in having a corolla that is not reflexed, but he must
have seen flowers that were not yet fully developed; at full maturity
the corolla is distinctly reflexed.

4. M. PARVIFLORA (Torrey) Woodson. Vincetoxicum parviflorum
(Torrey) Heller. Frequent in Rio Grande Plain, northeast to Karnes
County, west to Webb County, in sandy or gravelly soils. Flowering late
March—October. A pathological plant from Dimmit Co. (west of Artesia
Wells, Harold Gentry 1479; SMU) has much-branched inflorescences
with mostly malformed flowers, some proliferous, a pedicel or branch
arising from the flower center.

PARVIFOLIA (Torrey) Woodson. Gonolobus parvifolius Torrey
in Emory, Rept. U.S. & Mex. Bound. Surv. 2 (Botany): 166. 1859. “Sides
of hills, cafion of the Rio Grande, below Mt. Carmel, October; Parry.”
(A second specimen cited from “near the Limpia,’ Wright; this was later
referred by Gray to the next species.) The Sierra del Carmen is in
Coahuila, and it is most probable that Parry collected this plant on the
Mexican side, as he did the type of Chaetopappa Parryi; it was merely
Gray’s ignorance of local geography that led him to specify “Texas” in
the Synoptical Flora. I have seen no specimens of this, either from Texas
or elsewhere; it is included in the Kearney & Peebles Arizona Flora and
the Munz & Keck California Flora.

6. M. sagittifolia (Gray) Woodson in herb., ined. Gonolobus sagitti-
folius Gray, Proc. Amer. Acad. 12: 77. 1876. Type from “Rio Limpio,”
Jeff Davis Co., Texas, Wright. Described as having single, saucer-shaped,
entire crown. As long ago as 1942 Dr. Woodson used the binomial Matelea
sagittifolia in identifying plants from the Rio Grande Plain, geographi-
cally remote from the type locality and differing in critical details of
the crown. The name is not among the numerous transfers made by him

364

in 1941, and it has remained unpublished until now. The species is evi-
dently very rare; I have seen no specimens, but the original description
is quite clear.

M. Woodsonii Shinners, sp. nov. E descriptione M. sagittifoliae
peraffinis sed corona cupulata vel brevicylindrica margine 5-fida. HOLO-
TYPE: 8 miles northeast of Rio Grande City, Starr Co., Texas, Lundell
& Lundell 9926, 3 April 1941 (SMU). “Herbaceous vine, corolla green.
In scrub on sand.” Two other collections seen, both from Texas. Kleberg
Co.: about 5 miles southeast of Ricardo, Fred B. Jones 2816, 9 March
1959 (in private herb. Fred B. Jones). “On sandy slope near ravine.
Twining on Castela. Fls. greenish yellow.” Val Verde Co.: rocky (lime-
stone) hills above dam at foot of Devils Lake, about 20 miles N.N.W. of
Del Rio, Rogers McVaugh 7727, 31 March 1947 (SMU, TEX). “Scarce;
woody vine; corolla yellow-green.” It is this species which is reported
as M. producta in Flowering Plants and Ferns of the Texas Coastal Bend
Counties by Jones, Rowell and Johnston (1961, pp. 10—11).

8. M. PRODUCTA (Torrey) Woodson, Vincetoxicum productum (Tor-
rey) Vail. Leaf blades triangular-ovate, deeply cordate, soft-pubescent,
mostly 2—7 cm. long (two to four times as long as those of the two
preceding species). Rocky slopes, confined to the Trans-Pecos; specimens
seen from Brewster, El Paso, and Jeff Davis counties. Flowering April—
August.

9. M. GONOCARPA (Walter) Shinners, Field & Lab. 18: 73. 1950.
Vincetoxicum gonocarpos Walter. Gonolobus gonocarpos (Walter) Perry.
In a nearly north-south belt a little east of the center of the state, from
Cooke, Parker, and Dallas counties south to Comal, Karnes, and Brazos
counties, in steam-bottom thickets. Flowering late May—August.

10. M. DECIPIENS (Alexander) Woodson, Odontostephana decipiens
Alexander. Gonolobus decipiens (Alexander) Perry. Occasional in east-
ern part of north central Texas (specimens seen from Grayson, Hender-
son, Hunt, and Wood counties), in sandy woods. Flowering April—May.

11, M. RETICULATA (Engelmann) Woodson. Vincetoxicum reticula-
tum (Engelmann) Heller. Rather common from West Cross Timbers
(Palo Pinto Co.) south through the Edwards Plateau and Rio Grande
Plain, east in the middle parts of its range to Bastrop and San Patricio
counties, west to the eastern Trans-Pecos (Brewster and Pecos counties):
also in northeastern Mexico. In thickets or brush, rocky or silty ground.
Flowering March (far south), April, or May (at northern limits) to
October.

6. PERIPLOCA.

P. GRACEA L. is rarely cultivated and may persist, as indicated by
the following collection. Dallas Co.: from yard in White Rock area,
Dallas. Plant originally found on fence line near house at an old farm
on Gus Thomasson Road, now real estate development. Blackland soil,
flowers purple. Anne Estelle Orr 297, 4 May 1958. (SMU). Fernald in the

365

8th edition of Gray’s Manual reports this as escaped in the northeastern
United States and as far southwest as Oklahoma

SUPPLEMENTARY TRANSFERS AND NOTES
CYNANCHUM arizonicum (Gray) Shinners, comb, nov. Metastelma
arizonicum Gray, Proc. Amer. Acad. 19: 85. 1883.
ANCHUM Blodgettii (Gray) Shinners, comb. nov. Metastelma
Blodgettii Gray, Proc. Amer. Acad. 12: 73. 1877
CYNANCHUM Wigginsii Shinners, nom. nov. Metastelma ? angusti-
folium Torrey in Emory, Rept. U.S. & Mex. Bound. Surv. 2 (Botany):
159. 1859, Melinia angustifolia (Torrey) Gray, Proc. Amer. Acad. 12:

in honor of Dr. Ira L. Wiggins, indefatigable student of the flora of the
Sonoran Desert.
KEY TO SOUTHEASTERN CYNANCHUM

la. Leaf blades (at least middle and lower) with cordate base

2a. Appendages deeply divided into linear segments nearly equalling

the corolla. . C. laeve

2b. Appendages foriae < a noe crown + ath rounded ibe
ie C. cubense
lb. Leaf blades tapered to rounded-truncate at base

4a. Calyx lobes triangular-ovate, obtuse . .. . C. scoparium
4b. Calyx lobes lanceolate, acute . . . . « « C. palustre
3b. Corolla lobes pubescent or pilose ean

5a. Corolla about 3 mm. long, the lobes pilose within toward tip;
leaf blades linear-lanceolate, 1—4 mm. wide . . C. Blodgettu

5b. Corolla about 4 mm. long, the lobes densely pubescent within;
leaf blades oblong-lanceolate or oblong-elliptic, 6—18 mm. wide

C. Northropiae

C. laeve and C. palustre are included with fie meas species; the new
combination C. Blodgettii (Gray) Shinners is published above; authori-
ties for the other species are C. cubense (Grisebach) Woodson, C. North-
ropiae (Schlechter) Alain, C. scoparium Nuttall.

SARCOSTEMMA BILOBUM Hooker var. Lindenianum (Decaisne)
Shinners, comb. nov. S. Lindenianum Decaisne in DC., Prodr. 8: 541.
1844. S. bilobum ssp. Lindenianum (Decaisne) R Holm, Ann, Mo. Bot.
Gard. 37: 519. 1950

MATELEA albomarginata (Pittier) Shinners, comb. nov. Exolobus
albomarginatas Pittier, Contrib. U.S. Nat. Herb. 13: 108. 1910. Gonolobus
Albomarginatus (Pittier) Woodson, Ann. Mo. ard.

MATELEA aristolochiaefolia (Brandegee) cee comb. nov. Fisch-
eria aristolochiaefolia Brandegee, Univ. Calif. Publ. Bot. 6: 190. 1915.
Gonolobus aristolochiaefolius (Brandegee) Woodson, l.c.

366

MATELEA arizonica (Gray) Shinners, comb. nov. Lachnostoma ari-
zonicum Gray, Proc. Amer. Acad. 20: 296. 1885. Gonolobus arizonicus
(Gray) Woodson l.c. 243.

MATELEA calycosa (J. D. Smith) Shinners, comb. nov. Fimbristemma
calycosa J. D. Smith, Bot. Gaz. 16: 196. 1891. Gonolobus calycosus (J. D.
Smith) Woodson, l.c. 242

MATELEA chiapensis (Brandegee) Shinners, comb nov. Vincetoxicum
chiapense Brandegee, Univ. Calif. Publ. Bot. 6: 190. 1915. Gonolobus
chiapensis (Brandegee) Woodson, Ic.

MATELEA cteniophora (Blake) Shinners, comb. nov, Vincetoxicum
cteniophorum Blake, Contrib. Gray Herb. 52: 84. 1917. Gonolobus ctenio-
phorus (Blake) Woodson, l.c. 243.

MATELEA Greenmanii Shinners, nom. nov. Lachnostoma gonoloboides
Greenman, Proc. Amer, Acad. 39: 84. 1903. Gonolobus gonoloboides
(Greenman) Woodson, lec. 243. Not Matelea gonolobides (Robinson &
Greenman) Woodson, 1941.

MATELEA Johnstonii Shinners, nom. nov. Gonolobus stenopetalus
Gray, Proc. Amer. Acad. 21: 398. 1886. Matelea stenopetala (Gray)
Woodson, l.c. 231. Not M. stenopetala Sandwith, Kew Bull. 1931: 485.
The type of Gray’s species was collected by Pringle at Chihuahua, but
the epithets obviously suggested by collector and locality are both al-
ready used in the genus. I have therefore renamed it in honor of Dr.
Marshall C. Johnston, aia collector and keen student of the floras
of both Texas and Mexi

MATELEA tt eee (Hemsley) Shinners, comb, nov. Lachnostoma
lasiostemma Hemsley, Biol. Centr.-Am. Bot. 2: 335. 1882. Gonolobus
Lasiostemma (Hemsley) Woodson (sic), lc. 243

MATELEA oblongifolia (J.D. Smith) Shinners, comb, nov. Trichos-
telma oblongifolium J. D. Smith, Bot. Gaz. 48: 296. 1909. Gonolobus
oblongifolius (J. D. Smith) Woodson, Le. 243.

MATELEA Smithii Shinners, nom. nov. Fimbristemma stenosepala
J. D. Smith, Bot. Gaz. 18: 208— 209, 1893. Gonolobus stenosepalus (J. D.
Smith) Woodson, l.c. 243. Not Matelea stenosepala Lundell, 1942.

MATELEA stenantha (Standley) Shinners, comb. nov. Vincetoxicum
stenanthum Standley, Field Mus. Publ. Bot. Ser. 4: 255. 1929. Gonolobus
stenanthus (Standley) Woodson, l.c. 243.

FINDING-LIST OF GENERIC NAMES
The following list is of the generic names used in Gray’s Synoptical
Flora, Small’s Flora of the Southeastern United States and Manual of
the Southeastern Flora, and Kearney & Peebles’ Arizona Flora. A few
of the names are only in the sense used in one or more of these floras,
not as to proper type. The names in CAPITALS are those finally adopted
by Woodson, with the minor emendation of reducing Gonolobus to

367

another synonym of Matelea. The introduced Old World genera Crypto-
stegia and Periploca were not among those discussed by Woodson.

Acerates= ASCLEPIAS Lyonia=CYNANCHUM
Ampelamus=CYNANCHUM MATELEA (incl. Cyclodon,
Amphistelma=CYNANCHUM Edisonia, Gonolobus,
Anantherix=ASCLEPIAS Himantostemma, Lachnostoma,
ASCLEPIAS (incl. Acerates, Odontostephana, Pherotrichis,
Anantherix, Asclepiodella, Rothrockia, Vincetoxicum
Asclepiodora, Biventraria, in part
Gomphocarpa, Oxypterix, Melinia=CYNANCHUM
Podostigma, Schizonotus) Mellichampia=CYNANCHUM
Asclepiodella= LEPIAS Metalepis=CYNANCHUM
Asclepiodora=A PIAS Metastelma=CYNANCHUM
Astephanus=CYNANCHUM Odontostephana=MATELEA
Basistelma=CYNANCHUM OXYPETALUM
Biventraria=ASCLEPIAS Oxypteris=ASCLEPIAS
CRYPTOSTEGIA PERIPLOCA
Cyclodon=MATELEA Pherotrichis=MATELEA
CYNANCHUM (incl. Philibertella=SARCOSTEMMA
Amphistelma, Astephanus, Philibertia=SARCOSTE A
Basistelma, Epicion, Lyonia, Podostigma=ASCLEPIAS
Melinia, Mellichampia, Rothrockia=MATELEA
Metalepis, Metastelma, Roulinia=CYN HUM
Roulinia, Rouliniella, Seutera, Rouliniella= CYNANCHUM
Vincetoxicum in part) AR TEMMA (incl.
Edisonia=MATELEA Funastrum, Philibertia,
Epicion=CYNANCHUM Philibertella)
Funastrum=SARCOSTEMMA Schizonotus= ASCLEPIAS
Gomphocarpa=ASCLEPIAS Seutera=CYNANCHUM

Vincetoxicum =CYNANCHUM
(Gray’s species), MATELEA
Lachnostoma=MATELEA (Small’s species)

REFERENCES

GRAY, ASA. 1886. Asclepiadaceaec. Syn. Fl. N.A.

HOLM, RICHARD W. 1950. The American species of Sarcostemma R.
j -560.

(ed. 2) 2 pt. 1: 85-106, 401-404.
Br. (Asclepiada-

PERRY, LILY M. 1938. Gonolobus within the Gray’s Manual range. Rhodora 40: 281-
288. (Contrib. Gray Herb. /1.)

SHINNERS, LLOYD H. 1950. The species oF ues (including Gonolobus) in North
Central Texas (Asclepiadaceae). Field & Lab. 18

WOODSON, eve E., JR. 1941. The Nosh American Asclepiadaceae I. Perspective
of the general An Mo. Bot. Gard. 28: 193-244.

——————- ee The North egies eae species of Asclepias L. Ann. Mo. Bot. Gard.

41: 1-211.

NEW OR OTHERWISE INTERESTING
COREOPSIDINAE (COMPOSITAE) FROM
NORTHWESTERN SOUTH AMERICA

EARL EDWARD SHERFF

Department of Botany, Chicago Natural History Museum,
hicago 5, Illinois

Recently I was sent for examination a small lot of herbarium speci-
mens from the Smithsonian Institution, Washington, D.C. The lot was
found to contain one new species and one new variety of Bidens L. and
one new species and one new variety of Coreopsis L., also specimens
representing a range extension of one previously described species of
Coreopsis, and one specimen representing an isotype (or type?) of C.
holodasya Blake. For the privilege of examining these specimens, I must
express here my indebtedness to Dr. Lyman B. Smith, Curator of the Di-
vision of Phanerogams at the Smithsonian Institute. To assist me in sur-
veying all the known taxa in the genus Coreopsis for South America, I
have been particularly fortunate in being permitted to borrow all the
mounted photographs (79 herbarium sheets in all) belonging to the
Chicago Natural History Museum and representing South American taxa
in Coreopsis. For this privilege, I take pleasure in expressing my grati-
tude to Mr. E. Leland Webber, Director of the Chicago Natural History
Museum, and to Mr. J. R. Millar of the Department of Botany in that
institution.

BIDENS HOLWAYI Blake & Sherff, var. colombiana var. nov.—
Frutex, scandens, magnus, demum probabiliter saltem 5-10 m. al-
tus; caule ramisque tetragonis sulculatisque, glabratis. Folia petiolata
petiolis gracilibus 1.5-4.5 cm. longis basi connatis et hispido-ciliatis

branaceis lanceolatis rarius ovato-lanceolatis apice attenuato-acuminatis,
lateralibus subsessilibus vel _tenuiter petiolulatis petiolulis usque ad 7

(pedunculos tenues glabros sulculatos usque ad 2 dm. longos) adgregata,
radiata, pensa ad anthesin, +4 cm. lata et 2-2.3 em. alta. Involucri
patellati viridisque basaliter glabrati vel subdense reflexo-hispidi brac-
teae exteriores patentes vel demum reflexae 10-14, elongatae, 1-2.2 cm.
longae, oblonge lineares basim versus sensim angustatae apice acutae,
longitudinaliter 1- vel 3-7-nerviae marginibus ciliatae faciebus sub-
glabratae; interiores fere dimidio breviores, oblongo-lanceolatae, extus
glabrae vel apice pubescentes, multistriatae. Flores ligulati +3 (sine

SIDA 1 (6): 368—372. 1964.

369

dubio saepius circ. 5), flavi, ligula lineares, circ. 10- vel 12-nervii, +1.5
cm. longi, apice acriter angusteque circ. 3-denticulati. Paleae angustae,
lineares, tenerrimae, striatae, sub 1 cm. longae. Achaenia obcompressa,
anguste linearia, corpore 16-22 mm. longa et 1-1.3 mm. lata, atra, facie-
bus glabra et angustissime sulculata marginibus acriter setuloso-ciliata,
apice recte setosa et biaristata aristis gracilibus divergentibus circ. 5-6
mm. longis, fulvis, retrorsum hamosis.

Specimens examined: José Cuatrecasas & R. Romero Castaneda 25156,
“climber, leaves dark green, involucre green,’ Andean forest and bushes,
2700-2800 m. alt., Quebrada de Floridablanca, Sierra de Perija, Magda-
lena, Colombia, Nov. 9, 1959 (2 topotype sheets, US, where label on one
sheet reads “corolla lilac,’ surely an error; rays, which doubtless are
meant, are lacking, but cf. description for no. 25223 sequ.); Cuatrecasas
& Castaneda 25223, climbing shrub; leaves green above, grayish beneath;
ligules yellow, florets yellow; Andean forest and bushes, 2700-2800 m.
alt., same locality, Nov. 11, 1959 (1st type sheet, herb. no, 2339578, US;
2nd type sheet, herb. no. 2339577, US).

The first type specimen bears four flowering heads, these slenderly
and elongately pedunculate (peduncles 12-20 cm. long). The ligules ap-
pear to have been indeed yellow when fresh, as stated on the label for
the second type sheet, not “lilac’’ as recorded for one of the two topotypes
examined.

Appears to differ from the var. holwayi of southwestern Guatemala
(Quezaltenango) in its smaller flowering heads, these about 4 cm., not
about 6 ecm. across; outer phyllaries 10-14, not 8 or 9, and measuring
longer, about 1-2.2 cm. instead of 9-15 mm. long; ligules only about
1.5 not 2.3-3 em. long, etc. In my revision of the genus Bidens (Bot. Ser.
Field Mus. Nat. Hist. 16: 1-709. 1937), this variety would trace at once to
the fourth letter g on page 56. From the two species included there-
under, Bidens segetum and B. squarrosa, var. colombiana differs sharply
in its longer achenial bodies, these 16-22 mm. not 6-13 mm. long, also in
in larger flowering heads, with outer phyllaries 1-2.2 cm. not 3-7 mm
long, etc.

BIDENS pusilla sp. nov.—Herba pusilla, perennis, omnino hispidula,
caulibus patentibus numerosisque vix 6-10 cm. longis internodiis gracil-
limis tantum 4-16 mm. longis, usque ad circ. 1 mm. crassis, apicem versus
lateraliter subantrorsumque ramosis ramis terminaliter in pedunculos
gracillimos monocephalicosque productis. Folia opposita, minima, petiolo
adjecto tantum 4-7 mm. longa, petiolo subplano +1- 2 mm. longo, lamina
plus minusve tripartita, lobis (foliolis) lateralibus 1 jugo, sessilibus,
cuneato-obovatis apice irregulariter acriterque 2- vel 3-dentatis, lobo
(foliolo) terminali tripartito segmentis varie simplicibus vel rursus sectis.
cuneato-oblongis setis magis conspicuis sed paucioribus. Capitula radi-
ata, pansa ad anthesin 1.5-2 cm. lata et circ. 5 mm. alta. Involucri bracteae
exteriores circ. 5 vel 6, oblongo-lineares, circ. 3 mm. longae, apice

370

calloso subobtusae, extus subglabrescentes sed marginibus conspicue
ciliatae; interiores purpureo-atrae, lanceolato-oblongae vel elliptico-
oblongae, extus conspicue sed subsparsim elongato-setulosae, quam ex-
teriores usque ad duplo longiores. Flores ligulati 7 vel 8, flavi, sub 1 cm
longi, hgula anguste oblongi apice minutissime denticulati longitudinaliter
lineati. Paleae tenerrimae anguste lineari-oblongae apice atro-auranti-
acae acutaeque circ. 6-7 mm. longae. Achaenia obcompressa, anguste
linearia, exalata, inferne sensim angustata, utraque facie 2-sulculatis,
corpore atra +4.3 mm. longa, basi apiceque flavida, glabra vel apicem
versus aegre suberecteque setulosa, apice biaristata aristis +1 mm.
longis rectisque, apicem versus retrorsum paucibarbatis.

Specimens examined: Harriet G. Barclay & Pedro Juajibioy 7986, herb
with spreading stems and finely divided leaves; heads with wide, yellow

Prov. Cotopaxi, Ecuador, July 15-16, 1959 (US, type, herb. sheet no.
2372755)

In the entire genus Bidens L., this species is equaled by no other
species in the miniature size of its remarkably small leaves. The entire
plant on the type sheet bears a superficial resemblance to Bidens an-
themoides (DC.) Sherff of southern Mexico, a species with more decom-
pound leaves, measuring over all 1-5 or even to 7.5 em. in length. In the
key given in my revision of the genus Bidens (Bot. Ser. Field Mus. Nat.
Hist. 16: 57. 1937) this species would trace to the final letter g on page
57, except that its ligulate florets are scarcely 1 cm. long. From all taxa
included under the final g, however, B. pusilla differs at once in its
diminutive leaves.

COREOPSIS piurana sp, nov.—Frutex parvus, erectus, sine dubio sub
1 m. altus, supra corymbose ramosus et +25-capitulatus, caule sub-
angulato et sulculato demum atro-brunnescente et+3.5 mm. crasso, in-
ternodiis glabris et +3.5 cm. longis; ramis conspicue suberectis, tetra-
gonis, sub 1.5 mm. crassis, internodiis plerumque 2-3 cm. longis, glabris
vel inferne medianeque adpresse obsoletissimeque setulosis, superne in
corymbos parvos pauci-capitulatos abeuntibus; pedicellis tenuibus per-
spicue irregulariterque albo-tomentosis 2-3 (-5) cm. longis. Capitula
radiata, erecta, pansa ad anthesin circ. 2.5-3 cm. lata et vix 1 ecm. alta.
Involucrum obconico-hemisphaericum, superne glabrum inferne mani-
feste albo-tomentosum, bracteis exterioribus 8-10, ovatis vel oblongis vel
etiam oblanceolato-oblongis, adpressis, +4 mm. longis, l-nerviis, apice
subacutis vel subobtusis; interioribus oblongo-ovatis, exsiccatis subatris,
margine anguste diaphanis, apice irregulariter acutis vel obtusis 8-9
mm. longis, numerosissime striatis. Flores ligulati circ. 8 vel 9, aurei,
cire. 1.5 cm. longi; ligula elliptico-oblonga, numerose striata, apice con-
stricta et plus minusve denticulata. Paleae lineari-oblongae, tenuissimae,

371

striatae, dorso mediane sursum setosae, +6 mm. longae. Achaenia ob-
compressa, exalata, corpore oblongo-oblanceolato, nigro +4.5 mm. longo
et (superne) vix 2 mm. lato, tergo glabrato, facie ventrali marginibusque
sursum albo-setoso (setis summis plus minusve fasciculatis); apice bia-
ristato aristis rectis +1.5 mm. longis atro-stramineis, densissime antrorso-
setosis.

Specimens examined: J. Soukup H662, near Huancabamba, Dept. of

sima. 37. C. Pickeringii.” That species, however, is a plant of very differ-
ent habit, having solitary heads on long and slender peduncles measur-
ing up to 1.5 (more rarely to 2.3) dm. long. In C. piurana, by contrast,
the peduncles are clustered in groups of three or so, and are so short
and slender that they might better be termed pedicels, as given in the
above description.

COREOPSIS SUAVEOLENS var. ecuadoriensis var. nov.—Frutex forsi-
tan altior, usque ad 2 m. altus, similiter odore Covilleae divaricatae
(Cav.) Vail suaveolens. Capitula (non solitaria) +3-adgregata ad fines
ramulorum pedicellis gracilibus circ. 1.2-2.5 em. longis, sparsim brevis-
simeque glanduloso- hispidulis.

Specimens examined: Harriet G. Barclay and Pedro Juajibioy 8563,
shrub to 2 m. tall, growing up through other shrubs. Leaves finely divided
into linear segments. Involucral bracts green with darker vein in cen-

dental y Cordillera Oriental, Paramos de Silvan, Prov. Azuay, Ecuador,
July 30, Aug. 3, 1959 (type, US, herb. sheet no. 23728

Coreopsis suaveolens Sherff (Bot. Gaz. 89: 369. 1930; Revision of the
Genus Coreopsis, Bot. Ser. Field Mus. Nat. Hist. 11: 336. 1936) was
founded upon a single collection, Erich Werdermann 1114, at alt. of about
3,800 meters, Cordillera de Lallinca, Prov. Tarapaca, Dept. Tarapaca,
Chile, March, 1926, represented by four specimens, GH (type, my photo-
graph no. 2152) and (isotypes) F, S (my photograph no. 2029), and UC.
As remarked at the time, the species possessed a habital appearance
strongly similar to that of C. fasciculata Wedd. An easily distinguishing
character, however, was the pleasant odor of the dried herbarium speci-
mens, these having the fragrance characteristic of the well known
creosote-bush of the Southwestern United States, Covillea divaricata
(Cav.) Vail. The same agreeable odor is pronounced in the lone specimen,
Barclay & Juajibioy 8563, cited above from Ecuador for the type of a
new variety.

372

In C. suaveolens proper (1.e., var. suaveolens) the heads are solitary,
terminating the supernally naked branches (slender peduncles). In var.
ecuadoriensis, they are clustered mostly in threes on slender pedicels
about 1.2-2.5 cm. long.

COREOPSIS WOYTKOWSKII Sherff, Revision of the Genus Coreop-
sis, Bot. Ser. Field Mus. Nat. Hist. 11: 326. 1936—An extension of
range.—This species was based upon a single collection: Felix Woytkow-
ski 24, on rocky hills at altitude of 2625 m., vicinity of Celendin, Dept.
Cajamarca, Peru, June 5, 1936 (type, F, my photograph no. 3179;
Chicago Natural History Museum photograph no. 49160, of two frag-
ments of type). Shortly afterwards a collection was made in the De-
partment of Chachapoyas, Peru: Christopher Sandeman Lt, ALG. TOO" TE
Chachapoyas, Peru, August, 1938 (K, my photograph no. 3454). Among
the specimens of Coreopsis and related genera recently sent me from
the Smithsonian Institution (US) is an excellent specimen from still a
third collection: J. J. Wurdack 1147, shrub 1 m. tall, on dry cliff face.
Rays and disk yellow; summit of Puma-urcu southeast of Chachapoyas,
alt. 3100-3200 m., Prov. of Chachapoyas, Dept. of Amazonas, Peru. It is
seen then, that Sandeman 17 and Wurdack 1147 extend the geographic
range of C. woytkowskii, known heretofore only from the Department
of Cajamarca, into the Department of Amazonas, immediately to the
northeast of the Department of Cajamarca.

COREOPSIS HOLODASYA Blake, Kew Bull, 15: 373. 1962.—In my
above cited Revision of the Genus Coreopsis, this little-known species,
unknown till two years ago, would trace in the analytical key there
given to the first letter c on page 290, “Folia 0.5-2 em. lata.” Under-
neath that step are listed three species, the first two, C. oblanceolata and
C. woytkowskii grouped under the step d, and having the leaves oblan-
ceolate and 0.5-1.5 cm. wide: the third species, C. irmscheriana, standing
under the second d, and having the leaves narrowly ovate-lanceolate,
and 1-2 cm. wide.

In C. holodasya, the leaves are neither oblanceolate nor narrowly
ovate-lanceolate, but instead narrowly elliptic-oblong, and under 1 em.
wide. In my recently published Annotated List of My Botanical Writings
(Ilinois Wesleyan Univ., Bloomington, IIL, May, 1964), no mention
unfortunately was made of this species, which at the time was unknown
to me. It should have been provided for, of course, on page 18 of that
work, with the direction to include its mention on page 290 of the
Revision of the Genus Coreopsis, as above provided for. (In passing, we
may note a misprint of the word breviora in line 28 of page 290.)

—

NEW NAMES AND RECORDS FOR
TEXAS COMPOSITAE

LLOYD H. SHINNERS
Herbarium, Southern Methodist University, Dallas, Texas 75222

When I agreed to contribute a summary of the Compositae for F. W.
Gould’s Texas Plants —a Checklist and Ecological Summary (cover page
date June 1962; actually published January 1963), I fully expected to
have time to get the new names validly published in good time, and even
dreamed of throwing in keys to at least some of the genera for good
measure. But the distractions of moving with a large herbarium to a
new building, and subsequently of ill health, shattered the dreams and
delayed publication until now. This brief paper is intended primarily
to validate several names that appeared in the Checklist as nomina nuda;
some other name changes and new species or new records which have
turned up since completion of the Checklist account are also included.
They do not, alas, represent a final word. New names in Echinacea and
Thelesperma await publication of revisions of those genera which have
been completed by others. After becoming better acquainted with
Brickellia, I am satisfied that it cannot be maintained as a genus distinct
from Kuhnia, but the Herculean task of providing more than 100 new
names in the latter genus is temporarily postponed. Southeastern species
of Eupatorium badly need revision; only two of a number of inevitable
name changes are indicated here. Solidago needs much more collecting
in eastern Texas. Somed puzzling, localized forms of Aster cannot be
disposed of until better material is available. And so on, But despite its
shortcomings, I believe the Checklist summary represents very sub-
stantial progress with our knowledge of the largest family of plants in
Texas.

The peo ane are entirely new records to be added: eee ccc
scabridum, Machaeranthera brevilingulata, M. tenuis, M. texensis, Not
calais canada. Solidago petiolaris var. petiolaris, Thelesperma curiv-
carpum. The following should replace the Checklist names given in
parentheses: Bahia dissecta (B. biternata), Cirsium Engelmanii (C.
terrae-nigrae), Erigeron superbus (E. speciosas var. australis), E. lobatus

confused status of the names Evax and Filago is noted under the latter.
ACHILLEA MILLEFOLIUM L. var. OCCIDENTALIS (Rafinesque,
ined.) DC., Prodr, 6: 24. 1837. “Frequens a Pensylvania ad reg. Illinoen-

SIDA 1 (6): 373—379. 1964.

374

sem.” Including A. gracilis Rafinesque, Herb. Raf. p. 22. 1833. Locality
not specified, but this is species no. 12 under the heading ‘“Florula
Texensis .. . New Dicotyle Plants of Texas and Arkansas.” A. ille-
foltum var. gracilis (Rafinesque) DC., Lc. “Agro Kentuckiensi.” Though
A. gracilis was published earlier, in the rank of variety the two epithets
appeared simultaneously, and I have chosen occidentalis as the more
appropriate. This earlier name for what is commonly known as A.
lanulosa or A. Millefolium var. lanulosa has generally been overlooked.

ARTEMISIA LUDOVICIANA Nuttall var. albula (Wooton) Shinners,
comb. nov. A. albula Wooton, Contrib. U.S. Nat. Herb. 16: 193. 1913. A.
ludoviciana ssp. albula (Wooton) Keck, Proc. Calif. Acad. Sci. (ser. 4)
25: 446. 1946

ARTEMISIA LUDOVICIANA var. redolens (Gray) Shinners, comb.
nov. A. redolens Gray, Proc. Amer. Acad, 21: 393. 1886. A. vulgaris ssp.
redolens (Gray) Hall & Clements, The oe Method in Tax-
onomy (Carnegie Inst. Washington Publ. 326): 1923. A. ludoviciana
ssp. redolens (Gray) Keck, Proc. Calif. Acad. Sh ee 4) 25: 454. 1946,

BAHIA DISSECTA (Gray) Britton. This name should replace B.
biternata in the list; see Ellison, Rhodora 60: 190—199, 201—204, 1964.

BAHIA NEOMEXICANA Gray. After seeing specimens of the South
American Schkuhria multiflora Hooker & Arnott, I agree with Heiser’s
view that the North American plant is the same (see Ann. Mo. Bot. Gard.

: 274—275, 1945). Ellison agrees with Heiser in leaving the species
under Schkuhria, but I am not wholly persuaded. If retained under
Bahia, a new combination based on the Hooker & Arnott name is
required.

BAHIA WOODHOUSII Gray. At different times Gray treated this
under three different generic names, always spelling the eponymous
epithet with the double i, as here given. He obviously thought it a better
Latin form than Woodhousei. He did not misspell the name, as Warnock
implies (Wrightia 2: 74, 1960). Ellison also uses the illegitimately “cor-
rected” spelling in his revisicn.

CENTAUREA SOLSTITIALIS L. Long known from California and
more recently from Oklahoma, this Old World species had not previously
been reported from Texas. DALLAS CO.: Belt Line Road 0.3 mile west
of U.S. Highway 67, Cedar Hill, David Flyr, 6 June 1962 (SMU). “Also
seen about one mile north of Cedar Hill.”

CHRYSOTHAMNUS VISCIDIFLORUS (Hooker) Nuttall var. ludens
Shinners, var. nov. Folia glabra vel scabro-puberula lincari-oblanceo-
lata, majora 2.0—2.5 cm. longa, 2—3 mm. lata. Capitula pauca congesta
ramos erectos terminantia. Involucra 5.0—5.5 mm. alta. Corollae pro-
funde divisae lobis angustis 2—3 mm. longis. Achaenia glabra vel ad
angulos parcissime appresse hirsutula. HOLOTYPE: Guadalupe Mts.
above Pine Springs Station, Culberson Co., Texas, Shinners 9063, 15 Aug.

375

1946 (SMU). “On higher slopes, elev. about 6000 ft. Shrubs 42—1 m. tall,
virgate; bark blackish to gray-brown.” PARATYPE: Infrequent peren-
nial in limestone soil above Hunter Lodge, in South McKittrick Canyon,
Guadalupe Mts., alt. 9499 feet (sic! collection number by error repeated
for altitude; highest point in the range is 8751 ft.), Culberson Co., Barton
H. Warnock 9499, 13 Aug. 1949 (SMU). This somewhat isolated repre-
sentative of the C. viscidiflorus complex is named in allusion to the
fact that it will not fit any of the possible choices in the key to sub-
species given in the Hall & Clements monograph (The Phylogenetic
Method in Taxonomy, Carnegie Inst. Washington Publ. 326: 181, 1923),
and because of the glabrous or glabrate achenes will hardly even fit in
the key to species (l.c. 175). A similar but judging from the description
distinct plant has just been described from New Mexico as C. spathulatus
L. C. Anderson, Madrono 17: 226—227, 1964.

CIRSIUM ENGELMANNII Rydberg, Fl. Rocky Mts. 1013 and 1069.
1917. Based on C. virginianum var. filipendulum Engelmann ex Gray,
Man. ed. 2 p. 233. 1856. (Not C. filipendulum Lange, 1861.) C. terrae-
nigrae Shinners, Field & Lab. 17: 27—29. 1949. Purposely based on a
different type, but taxonomically identical with the preceding. Ryd-
berg’s name is as to type, not as to plant described; the Blackland Prairie
thistle does not extend even as far as the High Plains, let alone the
Rocky Mountains. It is confined to a narrow belt from south central
Oklahoma to central Texas, with an outlying southern station in Harris
County, Texas. I am indebted to Dr. R. J. Moore of the Plant Research
Institute, Canada Department of Agriculture, Ottawa, for calling my
attention to my oversight in providing another name for the species.

ENCELIA SCAPOSA Gray var. stenophylla Shinners, var. nov. A var.
scaposa recedit foliis angustissimis 1.0—3.5 mm. latis (vice 3—7 mm.).
HOLOTYPE: 91/3 miles east of Dryden, Terrell Co., Texas, V. L. Cory
43870, 28 March 1944 (SMU). A second sheet, probably a duplicate but
numbered 43869 (it was Mr. Cory’s practice at that time to number
every sheet rather than every collection), is designated PARATYPE,
same place and date (SMU). E. scaposa var. scaposa occurs farther west
at higher elevations, in Hudspeth and Jeff Davis counties.

ERIGERON SUPERBUS Greene. This name should replace E. speciosus
var. australis in the list. The only Texas specimen seen (Davis Mts., Jeff
Davis Co., M. S. Young, 13 Sept. 1918, TEX) has distinctly ciliate leaves
as in E. speciosus, but otherwise seems definitely to belong with E
superbus, which was reported from the same locality by Cronquist
(Brittonia 6: 150—151, 1947).

ERIGERON TENELLUS DC., Prodr. 5: 288. 1836. “In Mexico circa
Tamaulipas in campis Matamoros legit cl. Berlandier martio flor.” This
species seems to have been completely overlooked since its original
description. In above-ground parts it greatly resembles E. tenuis T. & G,,
and Texas collections have been referred to that species. But E. tenellus

376

is an annual with a slender taproot, while E. tenuis is perennial with
fibrous roots from a stubby crown. The following three collections from
Cameron Co., close to the type locality just over the border in Mexico,
may be cited (all SMU). About 8 miles west of Boca Chica, Lundell &
Lundell 10778, 17 March 1942. Yard in Brownsville, J. F. Brenckle 47-325,
3 April 1947. Along Highway 106 E. of Harlingen at Harlingen Air Force
Base, Alfred Traverse 1018, 21 April 1959.

ERIGERON Traversii Shinners, sp. nov. (Sect. Phalacroloma.) E.
strigoso peraffinis, sed foliis infimis saepe pinnatim dentato-lobulatis
vel sublyratis sicut in E. tenui, praecox (Marte-Maio florens), formosior,
ligulis latioribus (0.8—1.2 mm., vice 0.5—1.0 mm.) candidis vel rarissime
carneis. HOLOTYPE: Off U.S. 59, about 8 miless south of Nacogdoches,
Nacogdoches Co., Texas, Lundell & Lundell 11093, 11 April 1942 (SMU).
Pine Belt of eastern Texas and adjacent Louisiana, flowering two to
four weeks ahead of E. strigosus, a showier plant easily distinguished in
the field though not in the herbarium. The following additional collec-
tions have been seen. TEXAS. Jasper Co.: 6 miles southeast of Jasper,
Shinners 18,402, 9 April 1954 (SMU). Nacogdoches Co.: 15 miles south
of Nacogdoches, B. L. Turner 4377, 12 April 1958 (TEX). “Dark pink-
flowered form among a population of white-flowered types. Only plant
of this color seen in the vicinity.” Newton Co.: 3 miles west of Newton,
Shinners 18,387, 9 April 1954 (SMU). Panola Co.: 4.3 miles southeast of
Tatum, Shinners 18,503, 9 April 1954 (SMU). Sabine Co.: 12 miles south-
east of Patroon, Eula Whitehouse 20,861, 18 March 1949 (SMU). 4 miles
south of San Augustine, Shinners 18,450, 9 April 1954 (SMU). Shelby
Co.: 10 miles southeast of Center, Shinners 7618, 10 May 1945 (with
many empty receptacles, the flowers fallen) (SMU). LOUISIANA.
Sabine Parish: 4.8 miles south of Many, Shinners 22,772, 23 April 1956

MU)

When Dr. Traverse brought me specimens of Erigeron tenellus for
identification, I at first intended to name that species for him, recog-
nizing it as different from any previously known from the United
States. When it proved to have been named from Mexico, this species
was used instead, in appreciation for the many excellent collections
made by him in the Gulf States from Texas to Florida.

ERIGERON LOBATUS A. Nelson var. Warnockii Shinners, var. no
A var. lobato differt pedunculis strigosis subeglandulosis. eee
Brewster Co., Texas (without precise locality), Warnock 424, 15—23

EUPATORIUM GLAUCESCENS Elliott, Sketch Bot. S.C. & Ga. 2:
297. 1822. FE. cuneifolium Willdenow, Sp. Pl. (ed. 4) 3 pt. 3: 1753. 1803.
(Illegitimate name: the earlier E. Marrubium Walter is cited as synonym
without qualification, but not adopted.) E. semiserratum DC... Prodr..5:

377

177. 1836. E. cuneifolium var. semiserratum (DC.) Fernald & Griscom,
Rhodora 37: 179. 1935. E. parviflorum var. lancifolium T. & G., Fl. N.A. 2:
85. 1841. E. semiserratum var. lancifolium (T. & G.) Gray, Syn. Fl. N.A.
1 pt. 2: 98—99. 1884. The complex to which these names relate is a most
difficult one. Size of involucre, uesd by both Fernald and Cronquist to
distinguish this from E. linearifoltum and related plants, is not a reliable
character. I have adopted the oldest valid name as species. Unless var.
lancifolium and var. semtserratum can be shown to be taxonomically
separable, the former name must replace the latter. My studies have
not progressed sufficiently for me to state any conclusions.
EUPATORIUM SCABRIDUM Elliott, Sketch Bot. S.C. & Ga. 2: 299—
300. 1822. E. rotundifolium var. scabridum (Elliott) Gray, Syn. FL N.A.
1 pt. 2: 99. 1884. This is another of those species which, although originally
described from the Southeast, is rather rare there, but is widespread
and common west of the Mississippi River, especially in Arkansas and
Louisiana. Local in TEXAS. Newton Co.: State Forest No. 1, 5 miles east
of Kirbyville, Cory 49,775, 30 Sept. 1945 (SMU). Smith Co.: Swan, J.

description. Leaves smaller than in E. rotundifolium, with distinctly
cuneate bases.

FILAGO. The following new names were recently published for the
Texas plants previously listed under Evazx: F. candida (T. & G.) Shinners,
F. Nuttallii Shinners (Evax prolifera Nuttall, not Filago prolifera Pomel),
F. verna (Rafinesque) Shinners (Evax multicaulis DC., a later name
than E. verna Rafinesque), and F. verna var. Drummondii (T. & G.)
Shinners. There was a belated attempt to conserve the name Filago in
the previous sense by the questionable device of typifying it with a
species added in the Appendix to Species Plantarum rather than one
given in the main text. Presumably the proposal has been acted upon
by the Edinburgh Congress, but as this goes to press I do not know
what was decided. If Filago is thus conserved, there will have to be
another new combination for the last-mentioned variety under Evax
vernda.

HELIANTHUS ANNUUS L. var. texanus (Heiser) Shinners, comb.
nov. H. annuus ssp. texanus Heiser, Amer. Mid], Nat. 51: 299. 1954.

HELIANTHUS ludens Shinners, sp. nov. Annua? (radix deest) parva
erecta 32-43 cm. alta hispidulo-pubescens suprene corymboso-ramosa.
Folia petiolata laminis lanceolatis integris vel leviter sinuato-dentatis
triplinervibus. Capitula pauca mediocria involucris 7 mm. altis phyl-
lariis lanceo-linearibus sub-3-seriatis subaequalibus subappressis discum
vix aequantibus. Flores radii et disci flavi. HOLOTYPE: Lobo Flat, 19
miles east of Van Horn, Culberson Co., Texas, Turner, Tharp & Warnock
53-543, 28 Aug. 1953 (SMU). “Ditch beside cotton field.” In aspect more

378

like a Verbesina or Viguiera, but the achenes and pappus are definitely
those of Helianthus.

IVA AUGUSTIFOLIA Nuttall var. latior Shinners, var. nov. Folia
caulina laminis lanceolatis ad 50 x 12 mm., suprema laminis anguste
lanceolatis nec lineari- filiformibus. HOLOTPYE: south of Falfurrias, in

tinctly lanceolate, not “linear to linear-filiform” as described in R. C.
Jackson’s revision (Univ. Kansas Sci. Bull. 41: 805, 1960) and as found
in specimens of var. angustifolia. Stem leaves of the latter, as described
by Jackson, are 5—10 times as long as wide. A second specimen referred
to the new variety is divided at base into three stems, the central one
with a branch just above base, and has lost the middle and lower leaves;
the upper ones are 5—6 times as long as wide. Goliad Co.: 9.5 miles south
of Goliad, Shinners 25,206, 13 Oct. 1956 (SMU). Both collections are
from southwest of the range of var. angustifolia as understood e

MACHAERANTHERA annua (Rydberg) Shinners, comb. nov. Sider-
anthus annuus Rydberg, Bull. Torr. Bot. Club 31: 653. 1904. Haplopappus
phyllocephalus ssp. annuus (Rydberg) Hall, The Genus Haplopappus
(Carnegie Inst. Washington Publ. 389): 58. 1928. Machaeranthera phyl-
locephala var. annua (Rydberg) Shinners, Field & Lab. 18: 40. 1950
An erect annual, resembling a small Prionopsis ciliata, quite distinct
from the coastal M. phyllocephala.

MACHAERANTHERA BOLTONIAE (Greene) Turner & Horne, Brit-
tonia 16: 328. 1964. This name should replace Psilactis asteroides in the
list; transfer of the latter is prevented by Machaeranthera asteroides

reene, a different species.

MACHAERANTHERA BREVILINGULATA (Schultz-Bipontinus)
Turner & Horne, l.c. 324, Psilactis brevilingulata Schultz-Bipontinus ex
Hemsley. This species should be added to the list as NAW from Region
10.

MACHAERANTHERA PINNATIFIDA (Hooker) ee Sida 1: 295.
1964. This name should replace M. pinnata in the li

MACHAERANTHERA TENUIS (S. Watson) 6 & Horne, Brit-
tonia 16: 326. 1964. This species should be added to the list as NAW from
regions 6 and 1]

MACHAERANTHERA texensis (R. C. Jackson) Shinners, comb.
Haplopappus texensis R. C. Jackson, Rhodora 64: 142143. 1962. es
species should be added to the list as NPW from Region 6.

NOTHOCALIS CUSPIDATA (Pursh) Greene. Troximon cuspidatum
Pursh. Agoseris cuspidata (Pursh) Steudel. Microseris cuspidata (Pursh)
Schultz-Bipontinus. This Great Plains species has been known from as
far south as Oklahoma; it occurs also in the Texas Panhandle, in Lips-
comb, Ochiltree, and Roberts counties, where it was collected by
Charles S. Wallis in 1960. The nomenclature follows that of Kenton L.

379

Chambers (see Contrib. Dudley Herb. 5: 66—67, 1957). It should be added
to the list as NPC from Region 9.

SENECIO SPARTIOIDES T. & G. var. Parksii (Cory) Shinners,
comb. nov. S. Riddellii var. Parksii Cory, Rhodora 45: 164. 1943.

SENECIO Warnockii Shinners, sp. nov. Species gypsogena S. spartiodeo
affinis. Perennis sublignosa humilis 10—-30 cm. alta multicaulis plus
minusve floccoso-albescens. Folia crebra angustissime linearia carnosa
3—7 cm. longa ca. 1 mm. lata integerrima. Capitula, involucra, floresque

t in S. spartioideo. HOLOTYPE: 40 miles north of Van Horn, alt. 4000
ft., Culberson Co., Texas, Turner & Warnock 202, 16 Sept. 1948 (SMU).
When the troublesome S. spartioides complex is revised this may be re-
duced in status, but with its dwarf stature and crowded, entire leaves,
it is a much more extreme departure from the type than any of the other
variants included under that binomial. Three additional collections have
been seen. TEXAS. Culberson Co.: gyp soil along pipeline between Tex-
line and Orla, Warnock 10,276, 7 Oct. 1951 (SMU). County not deter-
mined: 2 miles south of Rustlers Springs, Parks & Cory 30830, 20 Oct.
1938 (SMU). NEW MEXICO. Eddy Co.: 13 miles S.W. of White City (S.
of Carlsbad), David B. Dunn 8732, 12 Oct. 1952 (SMU). “Arid alkaline
grassland. Caliche beds exposed. El. 3800 ft.”

SOLIDAGO PETIOLARIS Aiton var. PETIOLARIS. This was inad-
Merion! omitted from the lst. It should be included as NPW from
Region
| eee CURVICARPUM Melchert, S.W. Nat. 8: 179. 1963.
This should be added to the list as NAC from Region 7.

rateful acknowledgment is due Dr. B. L. Turner for the long-term
loan of critical specimens from the University of Texas, and to the
National Science Foundation, whose 5-year grant (1956—1960) in sup-
port of field work preliminary to a Flora of the Gulf Southwest per-
mitted much additional collecting and field observation of Compositae
as well as other groups in the region.

NOTES

DIGITARIA ISCHAEMUM (GRAMINEAE) IN MISSISSIPPI AND
TEXAS.—Although the second edition of Hitchcock’s Manual of Grasses
of the United States (1952) assigns the introduced Digitaria Ischaemum
Schreber an all-inclusive range (“Quebec to Georgia, west to Washing-
ton and California’), the map (p. 578) shows no records for most of the
southernmost states. The obviously expectable spread of the weed to two
more of those states can now be reported. MISSISSIPPI. Clarke Co.:
south side of Quitman, Shinners 29,074, 26 Oct. 1960. Coahoma Co::
Clarksdale, Shinners 25,587, 29 Oct. 1956. Wayne Co.: 5 miles southeast
of Waynesboro, Shinners 29,062, 26 Oct. 1960. TEXAS. Camp Co.: 4.4
miles north of Pittsburg, Shinners 16,140, 16 Sept. 1953. Cass Co.: 4%

Shelby Co.: 2 miles west-northwest of Joaquin, Shinners 22,351, 10 Oct.
1955. (All collections at SMU.) This more or less northern species is
exceedingly similar to the pantropical (believed to be originally Asian)
D. violascens Link, which is widespread and common in the Gulf states.
Descriptions and keys treating these two in Hitchcock’s Manual and in
Henrard’s Monograph of the Genus Digitaria (1950) are partly contra-
dictory and not reliable. After much effort, the best separation I can
make for them is as foilows, based on 57 specimens of D. Ischaemum
(47 U.S., 9 European, 1 Asian) and 31 of D. violascens (30 U.S., 1 West
Indian).

Width of racemes 1.3—2.0 mm. (smallest measurements on short race-
mes of small plants); spikelets 1.7—2.3 mm. long, mostly attached in
2’s or 3’s (singly near base and tip of raceme) . . . Ischaemum

Width of racemes 0.8—1.5 mm. (largest Srercimren ani on long racemes
of large plants); spikelets 1.3—1.8 mm. long, mostly attached singly or
in 2’s, but often in 3’s near middle of racemes . . D. violascens

Much is made by Henrard of difference in type of hairs on the spike-
lets, D. Ischaemum being characterized by capitellate hairs, D. violascens
by verrucose ones. With the usual magnifications of up to 10 diameters
it is impossible to make out this difference, and even with magnifica-
tions up to 30 diameters I could not make a satisfactory separation of
available material. Spikelet pubescence varies in abundance and length
in both species, and according to Henrard himself, even in kind in D.
violascens (he notes that some spikelets on the type specimen have
ordinary, non-verrucose hairs). This feature seems to me to be a matter
of minor genetic variation, not a character so fundamental that it can
be used to define entire sections of the genus. In his comments under
D. violascens, he states that D. Ischaemum (which he places in a differ-
ent section) differs in “longer, thicker spikelets, about 214 mm. long,

SIDA 1 (6): 380—381, 1964.

381

glabrous pedicels... .”’ But under the latter species he accepts as valid
a var. asiatica Ohwi with spikelets only 1.5—1.9 mm. long. I have seen
no spikelets as large as 2.5 mm. among the 57 sheets of D. Ischaemum
examined, nor are the pedicels always glabrous, but commonly variously
scabrous or puberulent at summit or throughout. In D. violascens, ac-
cording to Henrard, the spikelets are “scarcely 2 mm. long, mostly
1.6—1.8 mm. ... with scabrous pedicels.” The range in spikelet size for
the 31 specimens examined was 1.3—1.8 mm., as stated in the key, and
the pedicels are variously scabrous or puberulent as in D. Ischaemum.
In Hitchcock’s Manual, D. Ischaemum is keyed as having spikelets 2 mm.
long, 1 mm. wide, the hairs “or most of them” capitellate, while D.
floridana Hitchcock and D. violascens are separated on the basis of
spikelets 1.5 to 1.7 mm. long, about 0.6 mm. wide, the hairs not capitel-
late. The two latter are then differentiated as “Sterile lemma with 9
distinct nerves; spikelets sparingly pubescent, 1.7 mm. long” (but in
the description stated to be 1.5 to 1.7 mm.); “fertile lemma light brown;
racemes, if more than 2, not digitate” for D. floridana, “Sterile lemma
with 3 distinct nerves; spikelets distinctly pubescent, 1.5 mm. long; fertile
lemma dark brown, racemes usually all digitate” for D. violascens. For
the 31 sheets of D. violascens examined, none of these characters will
stand up. Henrard, who saw fragments of the type and only known
collection of D. floridana (from Hernando Co., Florida), adds that it
shows only non-verrucose hairs, and refers it to still another section of
the genus. I strongly suspect that D. floridana is merely a form of D.
violascens in which the non-verrucose hairs, conceded by Henrard him-
self to be present with the verrucose ones, are the predominant or ex-
clusive type.

Grateful acknowledgment is due the National Science Foundation for
two grants in support of field work in the Gulf Southwest, under which
many of the specimens used in this study were collected.—Lloyd H.
Shinners, Southern Methodist University, Dallas, Texas 75222.

CHROMOSOMES OF TWO MORAEA (IRIDACEAE) FROM SOUTH-
ERN AFRICA.—A new basic number of x=6 in Moraea has recently
been reported for 2 South African species by Riley (Canad. J. Genet. &
Cytol. 4: 50-55, 1962). Two additional species can now be assigned to
this line.

M. erici-rosenii Fries — n=—6, 2n= 12 (from 6 plants). N. RHODESIA:
Mwinilunga Dist., Zambesi River rapids, 4 miles N of Kalene mission,
10 Nov. 1962, Lewis 6224 (K, US, MO). “Collected at the base of massive
granite outcrops among islands of grasses and sedges in black, shallow,
wet soil; almost indistinguishable among other monocots until tepals
open daily at 4 p.m. till dark.” The species has been found sporadically
throughout southern Africa, but its rarity can be at least partially attri-
buted to late afternoon flowering for at other times of the day plants
are very difficult to locate. Mitotic chromosomes from untreated cells

382

of immature flower buds have submedian to subterminal centromeres
and vary from 10.3 be for the shortest pair to 18.2 pL for the longest pair.

M. setacea Ker. — 2n=12 (from 2 plants). S. AFRICA: Natal, Hlabisa
Dist., Charters Creek, 5 Dec. 1962, Lewis 6306 (K, US, MO). ‘Sloping
grass field in sandy soil just above ocean.’ The chromosomes found in
untreated root-tip cells are comparable with those of M. erici-rosenii.
viz., Ssubmedian and subterminal ranging from 10.8-19.9 je in length.

On measuring the chromosomes from pretreated cells of M. polystachya
illustrated by Riley, I estimate their lengths to vary from 12.1-15.4
while the chromosomes of M. spathulata appear to be only about one-
half as long. Riley noted that the chromosomes of M. polystachya had
subterminal and submedian centromeres. Thus in both chromosome
length and centromere position the chromosomes of M. polystachya are
similar to those of M. erici-rosenii and M. setacea.

Plants from both collections are in cultivation at the Royal Botanic
Gardens, Kew. I appreciate the help of Mrs. Susan Holmes of Kew in
determining these species.—Walter H. Lewis, Missouri Botanical Garden,
and Department of Botany, Washington University, St. Louis, Missouri.

ERIOGONUM ANNUUM (POLYGONACEAE) BIENNIAL IN NE-
BRASKA. The life-form of Eriogonum annuum. occurring in the United
States from North Dakota and Montana south to Texas and New Mexico,
is commonly described as therophyte (annual). I have observed this
species over a two year period in Holt and McPherson counties, Ne-
braska, where it behaves as a typical biennial. Its seeds germinate in the
spring, and a rosette is produced. The rosette overwinters, and the fol-
lowing year a leafy, flowering shoot develops, seeds are matured, and
the plant dies. At least in parts of Nebraska, then, Eriogonum annuum
is not a therophyte but is a hemicryptophyte of the semi-rosette type.
—John W. Thieret, University of Southwestern Louisiana, Lafayette.

A DECEIVING AQUATIC NEPTUNIA (LEGUMINOSAE) IN CEN-
TRAL AMERICA.—Neptunia prostrata is a distinctive and fantastic
species, particularly as one would scarcely expect to find a strictly
aquatic plant among the Mimoseae. The prostrate stems, lying just below
the surface in warm pools, are jointed and spongy-thickened, white
(one might use Vachel Lindsay’s term “fish-belly white”), soft and
fleshy, reminding one of a great worm: the leaves are held up in the air
and are sensitive, folding when touched: the flowers resemble those of
Mimosa. It is rather unexpected, then, to find another species of Neptunia,
usually terrestrial, invading the water and so closely simulating N.
prostrata as to masquerade frequently under that name in the herbarium.

NEPTUNIA PLENA (L.) Bentham f. lumbricoides Fassett, f. nov.
Planta aquatica caulibus incrassatis spongiosis prostratis submersis, eis
N. prostratae simulantibus. EL SALVADOR: Dept. La Paz, floating in
Laguna Nahualapa, 6 km. S.W. of El Rosario de la Paz, Fassett 28323,

383

21 October 1950 (HOLOTYPE F; ISOTYPES GH, MO, US, WIS). Corn-

spot in Crescentia savanna, Choluteca, 31 October to 9 November 1949,
Standley 24588 (F.). MEXICO: Acapulco, Guerrero, October 1894 to
March 1895, Palmer 284 (GH). In a pond-llano, Gutzalama, Cuyuca
District, Guerrero, 25 August 1943, Hinton 6495 (F, GH). BRAZIL: in
shallow water and on margins of Acude Columinjuba, Municipio de
Maranguapa, 9 October 1935, Drouet 2580 (GH). Lagoa Mecejana, Muni-
cipio de Fortaleza, 18 July 1935, Drowet 2143 (GH).

The two collections from E] Salvador are extreme, and in appearance
exactly simulate N. prostrata. They are distinguished from that species
by the longer fruit with sometimes as many as 18 seeds (4—8 in N.
prostrata), and by the gland at the summit of the petiole. (See Standley
& Steyermark, Flora of Guatemala, Fieldiana: Botany 24 pt. 5: 65, 1946.)
The aquatic phase of N. plena has recently been discussed in relation to
its occurrence in Texas (B. L. Turner, Revision of United States species
of Neptunia, Amer. Midl. Nat. 46: 84, 1951). The one collection of that
species from Texas appears to be f. lumbricoides.—Norman C. Fassett,
University of Wisconsin, Madison.

DITOR’S NOTE. The above was one of the last manuscripts com-
pleted by Dr. Fassett before his untimely death in 1954. Evidently in-
tended to be part of a series, it was originally titled “Studies of aquatic
plants in Central America. 2. A deceiving Neptunia.” It has been sub-
mitted by Dr. Hugh H. Iltis in order to make the herbarium name
available for use by another botanist now monographing the genus.

A HEXAPLOID LINUM (LINACEAE) FROM EASTERN ETHIOPIA.—
In Africa south of the Sahara, Linum is scarcely represented. No species,
for example, is listed for the Flora of West Tropical Africa and only 2 are
recorded from the region of the Flora Zambesiaca (by Robson, 2: 91—99,
1963). During a recent trip in Ethiopia, I was able to collect 1 species
listed in the latter flora, L. holstii Engler ex Wilczek. Plants were found
infrequently in Harar Prov., 7.4 km E of Giggiga (Lewis 5889, 24 Aug.
1962) on a short grass plateau at 5000 ft. Immature flower buds were
fixed and air mailed to England for storage at -40°C. At the same time,
herbarium vouchers were collected and these are deposited at K, US,

Seven months later, whole buds were squashed in 2% acetic-orcein.
Diakinesis in PMCs of 2 plants of L. holstii showed n=27 with the 27
bivalents illustrating a strong tendency for early terminalization of
chiasmata. The average size of chromosomes at diakinesis was
The species is thus a hexaploid in the x=9 series, a series common to
the eastern North American species of Linwm, but then only to the
tetraploid level (Osborne & Lewis, Sida 1: 63—68, 1962). The number
is unique to the genus and the species is to my knowledge the first ex-

384

ample of an indigenous hexaploid flax—Walter H. Lewis, Missouri
Botanical Garden, and Department of Botany, Washington University,
St. Louis, Missouri.

CAYRATIA JAPONICA (VITACEAE) IN SOUTHEASTERN LOUISI-
ANA: NEW TO THE UNITED STATES.—Among some collections made
at the Delta Regional Primate Research Center of Tulane University by
Michael Kent Rylander and sent to me for determination was a strange-
looking plant obviously in the Vitaceae, with pedately compound leaves,
unlike any North American species known to me. The tetramerous
flowers in short, wide, long-peduncled cymes indicated Cissus, and the
plant was first tentatively identified as C. japonica (Thunb.) Willd.
(included in Bailey’s The Standard Cyclopedia of Horticulture, but not
in his Manual of Cultivated Plants), then more positively as Cayratia
japonica (Thunb.) Gagnepain, Notulae Systematicae 1: 349, 1911 (more
fully treated by that author, with description and figures of flower
details, in Lecomte’s Flore Générale de V’Indo-Chine 1: 983—984 and PI.
xX , 1912). There are illustrations of the plant in Makino’s An Illus-
trated Flora of Japan (enlarged edition), p. 341, 1956 (as Cissus), and
Steward’s Manual of Vascular Plants of the Lower Yangtze Valley,
China, p. 233 (text account, p. 240), 1958. Both show rather obtuse
terminal leaflets. In the specimen these are acute, and Gagnepain’s

mate Research Center, Covington, St. Tammany Parish, Louisiana
(SMU). It possibly represents an escape from cultivation.—Lloy
Shinners.

THREE NEW VARIETAL NAMES IN SPHAERALCEA (MALVA-
CEAE).—In Thomas H. Kearney’s “The North American species of
Sphaeralcea subgenus Eusphaeralcea” (Univ. Calif. Publ. Bot. 19: 1—128,
1935), the author follows the American Code usage of undesignated tri-
nomials which are subspecies; he so refers to them repeatedly in the text.
Later, in a joint paper with Robert H. Peebles publishing new names
. Arizona plants, he included a paragraph replacing the oo

new combinations as varieties (Journ. Washington Acad. Sci. :
Hs 1939). In three cases the epithet used for a variety is not the ere
available in that rank, The correct combinations are supplied herewith.

S. E I var. californica (Parish) Shinners, comb. nov. S. Fendleri
var. californica Parish, Zoe 5: 71—72. 1900. S. Emoryi ssp. variabilis
(Cockerell) Kearney, Univ. Calif. Publ. Bot. 19: 39. 1935. S. Emoryi var.
variabilis (Cockerell) Kearney, Journ. Washington Acad. Sci. 29: 486.
1939

S. ANGUSTIFOLIA var. oblongifolia (Gray) Shinners, comb. nov. S.

385

incana var. oblongifolia Gray, Smithsonian Contrib. 5 art. 6 (Pl. Wright.
pt. 2): 21. 1853. S. angustifolia var. lobata S. Watson, Smithsonian Misc.
Coll. 15 (Bibl. Index): 1438. 1878. (Illegitimate new name based on the

angustifolia var. lobata S. Watson; the use of the epithet was evidently
a coincidence.) S. angustifolia ssp. lobata (Wooton) Kearney, Univ. Calif.
Publ. Bot. 19: 69. 1935. S. angustifolia var. lobata (Wooton) Kearney,
Journ. Washington Acad. Sci. 29: 486. 1939. (Illegitimate as a later
homonym of S. angustifolia var. lobata S. Watson.)

var. angustiloba Gray, Proc. Amer. Acad. 22: 292. 1887. S. tenuipes
Wooton & Standley, Contrib. U.S. Nat. Herb. 16: 148. 1913. S. digitata
ssp. — (Wooton & Standley) Kearney, Univ. Calif. Publ. Bot. 19:
91. 1935. S. digitata var. tenuipes (Wooton & Standley) Kearney, Journ.
iui cae Acad. Sci. 29: 486. 1939.

It should be noted that although not designated as new and not
entered in the Gray Herbarium Card Index, S. angustifolia ssp. cuspi-
data (Gray) Kearney, Univ. Calif. Publ. Bot. 19: 67, 1935, was a new
combination based on S. angustifolia var. cuspidata Gray. In the 1939
list of new varietal combinations, S. axillaris var. violacea (Rose)
Kearney appears by a slip of the pen as var. rosacea.—Lloyd H. Shinners.

NEW VARIETAL esen FOR NEW WORLD LUDWIGIA (ON-
AGRACEAE).—For the sake of uniformity in my several projected
floras, new ee in varietal rank are needed for plants recently
treated by Peter Raven as subspecies. For completeness all those occur-
ring in the New World are included. I see no benefit whatever in dis-
carding the rank of variety in favor of that of subspecies. Indeed, such
a proceeding is not in accord with the present International Code of
Botanical Nomenclature, for the two are not identical in status. It is also
highly impractical, for it will require an astronomical number of new
names. I prefer the lesser by far of two evils.

LUDWIGIA OCTOVALVIS (Jacquin) Raven, Kew Bull. 15: 476. 1962.
The automatic var. octovalvis applies to those plants treated by Munz as
Jussiaea suffruticosa (including var. ligustrifolia and var. octofila) and
by Hara as Ludwigia pubescens.

L. OCTOVALVIS var. macropoda (Presl) Shinners, comb. nov. Jus-
siaea macropoda Presl, Rel. Haenk, 2: 35. 1835. J. suffruticosa var.
macropoda (Presl) Munz, Darwiniana 4: 239. 1942. Ludwigia octovalvis
ssp. macropoda (Presl) Raven, Kew Bull. 15: 476. 1962.

OVALVIS var. sessiliflora (Micheli) Shinners, comb. nov.

Fissinen octonervia f. sessiliflora Micheli in Martius, Fl. Bras. 13 (2):

J. octonervia var. sessiliflora Micheli, ibid. 180 and pl. 35.

Ludwigia octovalvis ssp. sessiliflora (Micheli) Raven, Kew Bull. 15: 476.
2

386

LUDWIGIA PEPLOIDES (H.B.K.) Raven, Reinwardtia 6: 393. 1964.
The automatic var. peploides applies to those plants treated as Jussiaea
repens var. peploides by Munz and as Ludwigia adscendens var. pep-
loides by Hara

L. PEPLOIDES var. glabrescens (Kuntze) Shinners, comb. nov. Jus-
siaea repens var. glabrescens Kuntze, Rev. Gen. Pl. 1: 251. 1891. Lud-
wigia peploides ssp. glabrescens (Kuntze) Raven, Reinwardtia 6: 394.
1964

L. PEPLOIDES var. montevidensis (Sprengel) Shinners, comb. nov.
Jussiaead montevidensis Sprengel, Syst. 2: 232. 1825. J. repens var.
montevidensis (Sprengel) Munz, Darwiniana 4: 276. 1942. Ludwigia
peploides ssp. montevidensis (Sprengel) Raven, Reinwardtia 6: 395. 1964.

Further synonymy is supplied by P. A. Munz, “Studies in Onagraceae
XII. A Revision of the New World Species of Jussiaea,”’ Darwiniana 4:
179—284, 1942; Hiroshi Hara, “Ludwigia versus Jussiaea,” Journ. Jap.
Bot. 28 (10): 289—294, 1953: Peter H. Raven, “The Old World Species of
Ludwigia (Including Jussiaea), with a Synopsis of the Genus (Ona-
graceae),” Reinwardtia 6: 327—427, 1964.—Lloyd H. Shinners.

NOTES ON CALYSTEGIA (CONVOLVULACEAE) IN THE CARO-
LINAS.—In the forthcoming “Guide to the Vascular Flora of the
Carolinas” two species of Calystegia are included on the basis of single
collections, Calystegia sericata (House) Bell, comb. nov. based on Con-
volvulus sericatus House (Torreya 6:150, 1906), was collected in June
1940, by H. L. Blomquist, “about 8 miles north of Salem, Oconee Co.,
S. C.” (Duke No. 61054). This area, just across the border from the area
in Georgia which is the type locality for this species, was visited in
June 1964, but no trace of the plant could be found. A second species,
Calystegia soldanella (L.) R. Br., previously known in North America
only from west coast collections, was collected in May 1963 by Sue F.
Moore (No. 268) on the Atlantic side of the sand dunes between Kill
Devil Hill and Duck, in Dare Co., N. C. Both of these species represent
additions to the flora of the Carolinas as treated by previous manuals.
—C, Ritchie Bell, University of North Carolina, Chapel Hill, N. C

TWO YOUNGIAS (“CREPIS JAPONICA”: COMPOSITAE) INTRO-
DUCED IN THE SOUTHEASTERN UNITED STATES.—Under the name
Crepis japonica (L.) Bentham, a common annual weed of tropical to
warm-temperate regions, originally from southeastern Asia, was first
reported from the United States in Small’s Manual of the Southeastern
Flora in 1933 (p. 1495) as follows: “‘Roadsides, waste places, and meadows,
S La. Nat. of Japan.—(W.I.)—All year.” Fernald’s 8th edition of Gray’s

mentioned in the New Britton & Brown Illustrated Flora (1952). As
Youngia japonica (L.) DC., it is given incidental mention in the com-

387

panion Manual of Gleason and Cronquist (1963, p. 759), apparently
quoting Fernald: “reported to be locally established from Pa. to Va.”
There are a dozen sheets from the United States in the S.M.U. Herbarium
referred to this species, but representing two strikingly different forms.
In The Genus Youngia by Babcock and Stebbins (Carnegie Inst. Wash-
ington Publ. 484: 94—100, 1937) they are identifiable as Y. japonica ssp.
genuina and Y. japonica ssp. Elstonii. Despite the lack of obvious tech-
nical characters to separate them, and the scantiness of the material seen,
I am inclined to regard them as distinct species. Because the synonymy
of Babcock and Stebbins disregards the type method, the correct name
for the second subspecies in the rank of species is wrongly listed under
ssp. genuina. If treated as varieties, a new combination will be needed.
Partly to avoid this, the two are discussed below under their valid
binomials.

YOUNGIA JAPONICA (L.) DC., Prodr. 7: 194. 1838. Y. lyrata Cassini,
Ann. Sci. Nat. (ser. 1) 23: 88. 1831. Type from the island of Mauritius.

Y. Thunbergiana DC. was indeed validly named.) Plant rather small
(8—75 cm. tall), usually with only 1 or 2 (rarely 3—5) well-developed
stem leaves; ligules about 1 mm. wide; another-tube 1.75 mm. long
(measurements from Babcock and Stebbins). Chiefly in gardens or parks
in cities, Florida to Texas. The following collections have been seen.
FLORIDA. Lake Co.: weed around buildings, at Alexan der Ue as
R. K. Godfrey & Richard D. Houk 62791, 11 May 1963.

escape in our garden, Altamonte Springs, Dr. Paul O. Sc aL rt or
10 May 1958. Garden weed, same locahty, Schaller 8262 (bis), 1 Mare

1956. St. Tammany Parish: old field, Primate Research Center, ‘Coving-
ton, Michael Kent Rylander 215, 16 May 1964. MISSISSIPPI. Jackson
Co.: courthouse lawn, Pascagoula, F. H. Sargent 7844, 17 Oct. 1961.
TEXAS. Dallas Co.: Northaven gardens, northern Dallas, weed under
lathe, Shinners 29,153, 6 Nov. 1960. (On later visits, after an abnormally
severe winter, it was not found.) Galveston Co.: Friendswood, Bales
Nursery, in plant bed under lathing cover, Alfred Traverse 2622, 1 Nov.
1961. The height of the involucre in these specimens varies from 4.0 to
5.7 mm.; in 5 Asiatic specimens also seen (1 from Ceylon, 4 from Pakis-
tan) it varied from 4.2 to 6.0 mm., considerably exceeding the limit of
5 mm. given by Babcock and Stebbins and others.

YOUNGIA THUNBERGIANA DC., Prodr. 7: 192. 1838. Based on Pre-

388

nanthes lyrata Thunberg, Fl. Jap. p. 303. 1784. (Not Youngia lyrata
Cassini, 1831.) According to Babcock and Stebbins, type material in the
Thunberg Herbarium “is apparently the same as certain slender forms”
of their Y. japonica ssp. Elstonii, but because specimens in the De
Candolle Herbarium named Y. Thunbergiana were actually Y. japonica
ssp. genuina, the binomial is placed in synonymy under the latter with-
out the qualification that it was as to plant described, not as to type,
which of course is that of Thunberg’s species.—Chondrilla lyrata (Thun-
berg) Poiret, Encycl. Meth. Bot. Suppl. 2: 332. 1811. Also misleadingly
cited by Babcock and Stebbins as a synonym of Y. japonica ssp. gen-
uina—Crepis japonica var. Elstonit Hochreutiner, Candollea 5: 340—341.
1934.—Youngia japonica ssp. Elstonii (Hochreutiner) Babcock & Steb-
bins, Carnegie Inst. Washington Publ. 484: 98. 1937. Plant generally much
more robust than Y. japonica, half to over a meter in height, with leafy
stem, the leaves gradually reduced upward; ligules 0.5—0.6 mm. wide;
anther-tube 0.75—1.0 mm. long. Having seen only two specimens, I am
unable to say how well the measurements given by Babcock and Steb-
bins stand up. It is curious that the more robust plant should have
markedly smaller flower parts. Hochreutiner found this and Y. japonica
growing together in Hawaii, but with no intermediates. Only the follow-
ing two collections have been seen. DISTRICT OF COLUMBIA. Wash-
ington, Kensington at Connecticut Ave. and Rock Creek, along wood
road, apparently an old dumping ground, F. H. Sargent s. n., 19 May
1951. (Plant 70 cm. tall, in early flower; involucre 5.5 mm. high.) NORTH
CAROLINA. Pasquotank Co.: roadside, 1.3 miles north of Knobbs Creek
on U.S. 17-158 then 0.1 mile east (north of Elizabeth City), Harry E.
Ahles (With R. P. Ashworth) 40082, 10 May 1958. (Plant 80 cm. tall,
fruiting; involucre 5.3 mm. high.) In both specimens the leaves are pin-
natifid with rather sharply pointed lobes and teeth. In those of Y.
japonica the leaves are more strongly lyrate, the terminal lobe much
larger than the fewer lateral ones, commonly rounded or blunt but
occasionally rather sharp-pointed.

The restriction of Y. japonica to the Gulf States, and its apparent
inability to withstand the continental winter climate of Dallas, suggest
that it is less cold-resistant than Y. Thunbergiana. If that is the case I
would expect the reports of Crepis japonica from Pennsylvania and
Virginia to belong rather to Youngia Thunbergiana. Lloyd H. Shinners.

REVIEW

WEBSTER’S THIRD NEW INTERNATIONAL DICTIONARY OF THE
ENGLISH LANGUAGE, UNABRIDGED. C. & L. Merriam Co., Spring-
field, Massachusetts. 1961.

Many have been the reviews, not all of them friendly, of Webster’s
Third New International Dictionary, Unabridged. The present general,
but not exhaustive, survey of the way in which botany fares in this tome
does not inspire supreme confidence. (At least the work is vastly better
than the cheap pocket dictionary I once saw, which defined “electricity”
merely as “a subtle force”; one assumes that its editors were never
struck by lightning!)

The advertisement at the very end of the dictionary emphasizes its
easy “crisp” definitions, whereas I would say that its major scientific,
if not literary, fault is a very lack of crispness. Instead of the clear,
concise, definitive statements which one would expect in a dictionary,
there is too often a marked tendency to stray deep into the citation of
examples and other non-definitive material. If the editors desired to
make brief entries of an encyclopedic nature, this material might better
have been placed in separate sentences. (Sentences, however, are scrupu-
lously avoided by the editors, who do not even end the entries with a
period.)

As one of the best (worst) examples, the entry for “enzyme” may be
cited: “any of a very large class of complex proteinaceous substances (as
amylases or pepsin) that are produced by living cells, that are essential
to life by acting like catalysts in promoting at the cell temperature usu.
reversible reactions (as hydrolysis and oxidation) without themselves
undergoing marked destruction in the process but frequently requiring
the presence of activators (as metal ions) or of coenzymes, and that can
act also outside of living organisms and therefore are useful in many
industrial processes (as fermentation, tanning of leather, and production
of cheese).” In this example, the problem is not conveying of misinforma-
tion; the statement at its beginning is a basically good and comprehen-
sive one, albeit a complex one to read (“‘crisp’?). But why tack onto a
definition two more clauses of non-definitive matter and a list of indus-
trial processes which are made possible by the fact that enzymes may
act outside of living organisms?

Another fine example of “definition” by confusing example or use
with directness is found under “diastase”
tained usu. as a yellowish white nice enous powder from malt and used
chiefly in desizing textiles and converting starch to maltose.” Similarly,
the treatment of “neurospora”’: “a genus of ascomycetous fungi (family
Sphaeriaceae) used extensively in genetic research, having black peri-

SIDA 1 (6): 389—392. 1964.

390

theciae [sic; the singular perithecium is properly listed elsewhere in the
volume] and persistent asci, and including some forms that have salmon
pink or orange spore masses and cause severe damage in bakeries.”
Apart from the fact that the nature of the damage (whether to the
bakery furniture or to its products) is left entirely to the imagination,
we find no use of the word “conidia” nor implication that they are
characteristic of the genus. Note also that the countless names of genera
are not capitalized as main entries, necessitating insertion of “cap.” each
time; when a word is always begun with an initial capital it might
have saved space and promoted clarity and good usage to enter it that
way.

A matter of style which is grammatically careless as well as potentially
misleading, especially to persons outside the relevant field—those who
most need the dictionary, is the very frequent fuzziness in relative pro-
nouns (“that” seems for some reason decidedly preferred to Ridecsui,
These often do not follow their antecedents; e.g., under “ovary”
“... basal portion of the pistil or gynoecium of an angiospermous Bien
that bears the ovules...” [is it the portion, the pistil, the gynoecium, or
the plant which bears the ovules?]. Or, under “chondriosome”: “any of
a class of . . . lipoprotein complexes in the cytoplasm of most cells that
are thought to function .. .” [what is thought to function? }.

In the rapidly growing areas of cellular and physiological biology, the
editors had reasonable success in keeping up with new words, although
they sometimes seem to have been unable to crystallize the primary
definitive features and hence there is much ‘ ‘beating around the bush.”

hile “respiration” is quite well treated, “digest” and ‘‘digestion” are
rather too much defined in terms of each other, and with no clear
indication of any applicability in the plant kingdom. “Ribosome” is not
included at all (although the prefix “ribo-” is ), and mitochondrion is
defined as a granular or globular (rather than the more frequent rodlike)
chondriosome, the primary entry being the much more archaic latter
word. The definition of auxin does not make clear that it is a naturally
occurring substance in | plants (merely that it promotes growth or causes

enzymatic, comparing them to viruses (rather than the other way
around), and making no mention of nucleic acid. The discussion of
omits the important new idea of a duplex molecule but
does devote half its words to material irrelevant to the definition. (The
hydrolysis products of polynucleotides would better be discussed in a
separate sentence. )

The common word “mold” is not very lucidly treated. The first defini-
tion is exceedingly broad (“a superficial often woolly growth .. .”)
with no oS that the growth causes the decay on which it is
esp. und; the second definition attempts a narrowing down “esp.”

391

to the order Mucorales—thus omitting a great many of the plants usually
called molds, a term without much taxonomic significance.

Names of families and orders of plants are freely listed, and the editors
cannot be blamed for the inherent problem of assigning families to
orders when there is so little agreement among botanists on definitions
“Amentiferae”’ but not with “Parie-
in
some classifications.” In a basically modern approach, the dictionary
accepts the widely used Tippo classification of Tracheophyta and its sub-
divisions. It is to be hoped that good practice will be promoted among
users of the dictionary by its clear indication that the names of higher
categories are plural in form. Good botanical usage does not accept un-
designated trinomials (lacking insertion of the ultimate rank, whether
variety or subspecies) and it is unfortunate that such trinomials regu-
larly appear when such taxa are cited. It is welcome to see made the
distinction between preferred usage of “phylum” in the animal kingdom
and ‘‘division” in the plant kingdom. Overall, the editors are to be con-
gratulated upon freeing themselves from the influences of the merican
Code” of nomenclature, which permeated the Second Edition. Tautonyms
are apparently avoided, and family names are more generally acceptable.

The scientific names of plant species referred to seem reasonably up
to date, a conspicuous exception being “Rhus toxicodendron” for the
poison ivy “common in the eastern and central U.S.” At least it is stated
how one contracts poison ivy, while the statement under poison hemlock
and many other poisonous plants makes no reference to the part of the
plant which is poisonous nor to defining the nature of ‘‘poisonous”—
whether upon mere contact or only actual ingestion. Nor is there refer-
ence to the colored plate (not identical with that in the Second Edition)
of poisonous plants (some 17 pages after the “poison” entries). Skunk
cabbage is pictured on this plate, but no mention of any sort of poison
(unless one counts “offensive-smelling”’) is given in the definition of
skunk cabbage—which is seldom if ever considered an important poison-
ous plant. The function of the plate of so-called poisonous plants thus
seems chiefly ornamental, for correlation with definitions is minimal.
Another lack of correlation between text and illustration is under
“nasturtium,” which is properly considered in the light of its two widely
differing applications (Tropaeolum and a cruciferous genus). However,
the drawing of “Nasturtium” does not state which of the two definitions
it illustrates and hence is rather useless.

“Adder’s-tongue” comes in for confusion almost as bad as_ that
described by Fernald (Rhodora 46: 313—314.. 1944) in reviewing another
dictionary. After the first definition (Ophioglossum), the new Webster’s
ee in Achillea, Arum, Erythronium, Geranium, Orchis, and Peram-

m, “having leaves or flower or fruiting spikes suggesting the fruiting
ke of adder’s-tongue fern.” Botanists familiar with these plants will

392

raise their eyebrows; there is no need to elaborate on the general lack of
resemblance of these plants to the fertile frond of Ophioglossum nor on
the fact that of the plants named only Erythronium is commonly called
adder’s-tongue.

General words with precise biological applications sometimes fare very
well; “nomenclature” and “publication,” for example, are given their
specialized meanings. The new and exceedingly popular word “taxon” is
duly included, its origin indicated as “ISV” [International Scientific

ocabulary”—words with no positive evidence that they were coined
in English]. An acceptable definition of “polygamous” in its botanical
sense is included. Inconsistently, “polygamodioecious” is given a very
poor definition (“having some plants polygamous and some dioecious in
the same species’), while “polygamomonoecious” is not listed at all.
“Species” is given a modern definition in that there is emphasis on rela-
tionship to evolutionary process, but there is too much stress on sexual
reproduction and no reference to the possibility of asexual species (which
are not uncommon in the plant kingdom).

Attention should perhaps be called to the fact that this edition omits
both the gazetteer and biographical portions which had considerable
usefulness in its predecessor.

The only actual typographical error I happen to have encountered
(unless “peritheciae,” mentioned above, is considered one) is “Araman-
thaceae” (for Amaranthaceae) under “Caryophyllales.”

In summary, the dictionary has been generally successful in including
new words, but has regressed in often including both definitive and
supplementary material (examples, etc.) in a single, complex, decidedly
“uncrisp” statement. To persons outside a field, wading through termi-
nology which may be unfamiliar, this practice is likely to lead to further
confusion in selecting the really essential definitive points—a matter in
which the editors themselves seem sometimes confused. All this is not
to deny that there are many excellent, fully acceptable, and helpful
definitions (‘flower” is a good example). But one would hope for a
higher percentage of such definitions in a work which has gone to con-
siderable trouble to include the words.

I am indebted to Dr. A. S. Sussman, chairman of the Department of
Botany, University of Michigan, for his helpful advice in evaluating the
treatment of words in the areas of physiological and cellular biology.
—Edward G. Voss, Herbarium, University of Michigan, Ann Arbor.

INDEX

Names of contributing authors are in capital letters. New scientific

names are in boldface followed first by page on which formally pub-

lished (in some cases this is not the first page on which mentioned).

Synonyms are in italics. Optional capitalization of specific names is uni-

formly adopted here though not used by most of the contributing au-

thors. Illustrations are indexed only by captions, in some cases not on

the same page.

Abies religiosa 263
Abietinella abietina 124
Abutilon Hulseanum 294
Acacia angustissima 263, 266;
farnesiana 263, 266; paniculata

Acanthaceae 304

Acer Negundo 303; rubrum 300
Aceraceae 303

Acerates 367

Achaetogeron 110

Achantidium flexellum 201
Achillea gracilis 374; lanulosa
164, 199, 208, 210, 212, 374; Mille-
folium 264; var. gracilis 374,
var. lanulosa 374, var. occiden-
talis 373; Se 165, 210

Actinogyra 199, 224; Munienbers
gil 121, 200
Actinomeris alba 253; paniculata

ADAMS, PRESTON 269
Adiantum concinnum 267; Kaul-
fussii 267; Poiretii 267; tenerum

Aeschynomene virginica 266
Africa 274, 38

Agastache breviflora ssp. Havar-
di 107, Havardii 107

Agastache breviflora (Gray) Ep-
ling var. Havardii (Gray) Shin-
ners, comb. nov. (Labiatae) 107
Age of Conformists 23; Dilettan-
tes 22, 23; Empire Builders 22

Ageratum corymbosum 264

Ze
trachycaulum 129, 198, 209,

Agrostis hyemalis 98; scabra 130,
198, 200, 205, 209, 212

Algae 120

Alismataceae 129, 264 (see also
Sagittaria)

Allium glandulosum 264; Schoe-
noprasum var. sibiricum 137,

Alnus crispa 141, 193, 195, 199,
201, 205; incana 141, 205, 208,
209, 210, 232

Alopecurus aequalis 130, 204, 211;
carolinianus 98

Alsine 49; americana 50; tennes-

Amaranthaceae (see Amaran-
thus)

Amaranthus albus 248, 249; bli-
toides 248, 249; graecizans 249;
microphyllus 248, 249
Amaranthus microphyllus Shin-
ners, sp. nov. (Amaranthaceae)

Amaryllidaceae 264, 301
Ambrosia elatior 305
Amelanchier alnifolia 151, 196,
198, 199, 201, 20

Amellus ee caloeis 295
Ammopursus Ohlingerae 240, 241,
242, 243

Ampelamus 367; albidus 359

394

Amphicarpa bracteata 303
Amphistelma 367

Andromeda _ polifolia 158, 192
194, 203
Androsace

199, 213
Aneilema 274, 278, 279, 284, 286,

septentrionalis 159,

101; lineare 100; montanum 279;
nudiflorum 100, 101; peduncu-
losum 279, 292;
279, 292; Welwitschii 279
Aneilema (Commelinaceae) in
the southern United States 100
Anemia adiantifolia aoe

207; patens var. Wolfgangiana
146, 196,

Annual Sisyrinchiums (Iridaceae)
in the United States 32

Anoda cristata 267; hastata 267
Antennaria parvifolia 165, 199,
208; pulcherrima 165, 193; rosea
165; subviscosa 165

Anthericum aurantiacum 266
Aphanostephus pachyrrhizus 264
Apocynaceae 161,

Apocynum androsaemifolium 161,
196, 211

Aquilegia brevistyla 146, 196,
209,

Arabis divaricarpa 147, 199, 213;
Drummondii 147; hirsuta var.
pycnocarpa 147, 207; Holboellii
147, 199, 201, 212

Araceae 136, 301

Arbutus glandulosa 262
Archibaccharis 110

Arctagrostis latifolia 130, 206
Arctostaphylos rubra 158, 192,
93, 194; Uva-ursi 158, 192, 193,
94, 195, 196, 198, 199, 200, 207,
211, 220

Arenaria 48; americana 50; ano-
mala 50; argaea 50: Benthamii
saa capillaris 143, 196, 199; ceras-

nm pe

00; lanuginosa 302, var. cineras-
cens 51; lateriflora 143, 207, 209,
210; ludens 51; muscorum 51:

tula 49; persica 50; rubella 143,
199; saxosa 51, var. cinerascens
d1; Stephaniana 50, var. ameri-
cana 50; trigyna 51

Arisaema Dracontium 301
Arnica Chamissonis 165, 210; lon-
chophylla 165, 196, 199

Arnold Arboretum 12

Artemisia albula 374; biennis 165;

albula 374, var. albula 374, var.
gnaphalodes 165, ssp. redolens
375, var. redolens 374; redolens
374; Tilesii ssp. unalaschensis
165; vulgaris ssp. redolens 374
Asclepiadaceae 264, 304, 358

Asclepias 358, 359; grandiflora

Asclepiodella 367

Asclepiodora 367

Asplenium abscissum 300; Curtis-
sii 300; heterochroum 300; resili-
ens 300; verecundum 300
Astephanus 367

Aster alpinus var. Vierhapperi

199; johannensis 166;
mis 166, 204, 205, 208, 209, 210;
Lima 264; pansus 166, 208;
peregrinus 110; pinifolius 305;
sibiricus 166, 196, 212
Astianthus viminalis 263, 264

var. ceramicus 316, var. imper-
fectus 316, 324; coccineus 3165,
316, 324; cymboides 316, 324;
dasyglottis 153; desperatus 316,

leios 320, 326; inyoensis 317, 324

Johannis-Howellii 315, 317, 324:
Kentrophyta 315, var. colora-
densis 317; lentiginosus 319, var.
Fremontii 317, 319, 324, var.
palans 317, 319, var. variabilis

Earlei 319, var. Thompsonae 317,
319: monspessulanus 320; Nut-

tallianus var. micranthiformis

323; plattensis 317; Preussii 315,
318; Purshii var. glareosus 318,
319, 324, var. lectulus 318, 319,

395

var. longilobus 318, 319, var.
Purshii 319; Ravenii 318; Sere-
noi 318; sesameus 321; Shorti-
anus 318; somalensis 314, 3

153; uliginosus 321; umbellatus
321; utahensis 318; vulpinus 321,
326; Whitneyi 315, 318, 324; yu-
konis 153; zionis 318

Athanasia paniculata 253

Atlas of the British Flora 257
Atriplex patula 143, 208
Aulacomniaceae 123
Aulacomnium acuminatum 123;
palustre 123; turgidum 123
Axyris amaranthoides 143, 212
Ayenia montana

Bacopa caroliniana 304

Bahia biternata 373, 374; dissecta
373, 374; mneomexicana 374;
Woodhousii 374

BANKS, DONALD J. 306
Barbarea orthoceras 147,
Basistelma 367; angustifolium
365

Beck, Lewis C. 4, 5

Beckmannia Syzigachne 130, 208,
209, 1

BELLI Cy RITCHIE 360
Bentham 258

Bessera elegans 266

Berlandier 25

Betula glandulosa 141, 192, 196,
203, 205, 207; occidentalis 141;
papyrifera 141, 196, 198, 199, 200,
205, 211, 222, 224, 236
Betulaceae 141 (see also Coryla-
ceae)

Bidens anthemoides 370; bipinna-
ta 305; cernua 166, 206; Holwayi
var. colombiana 369; pilosa 305;

396

pusilla 369; segetum 369; squar-
rosa

Bignoniaceae 264, 304

Biltmore Herbarium 24, 30
Biographical Index of Deceased
British and Irish Botanists 257
Biventraria 367

Black spruce forest 190
Boehmeria cylindrica 302

Boraginaceae 161

Boschniakia rossica 162, 193, 194
Botanical Arrangement of All the
Vegetables Growing Naturally
in Britain 258

Territories, Canada. I. Catalogue
of the Flora 117. II. Vegetation
187.

Botrychium dissectum var. tenu-
ifolium 300

Bouchea prismatica 268
Bouvardia ternifolia 267
Bowlesia incana 294

Bradburya floridana 182

Brassica campestris 148, 212
Braya humilis 148, 207

British Isles 257

British Museum 257

Britten and Boulger, Biographi-
cal Index 257

Britton, Nathaniel Lord 7, 8, 14,
19 20. 22.

Britton and Brown Illustrated
Flora 7, 8, 19

Britton’s Manual 7, 8, 19
Bromeliaceae 301

Bromus inermis 130, 212; Pump-
ellianus 180,

Brongniartia podalyrioides 266
Bryaceae 123

Bryophytes 122

Bryum lacustre 123, 207
Buckley 25, 29, 30

Buddleia americana 262

Buellia papillata 122

Buelliaceae 122

Bulbostylis barbata 294
Bunchosia Palmeri 266

Bursera 263

Calamagrostis 130, 208, 210, 238:

198, 208; purpurascens 195, 196,
198, 211, 21
Calamintha 69; arkansana 72, 70;

Ashei 73, 71, 93; canescens 87:
carolintiana 74; coccinea 71, 73

71; Nuttallii 72; officinalis 70, 71
Calamintha (Labiatae) in the
southern United States 69

Calla palustris 136, 204, 206
Calliandra grandiflora 266; Hous-
toniana 266; penduliflora 266
Callicarpa americana 304
Callisia cordifolio 301; fragrans

Calisteris texana 177
Callitrichaceae 155

Callitriche hermaphroditica 155,
206; palustris 155, 206
Caloplaca elegans 122
Caloplacaceae 122

Caltha natans 146, 204, 206; pa-
lustris 14

Calylophus 337; Drummondii 339;
Hartwegii 337, 338, 341, var.
filifolius 342, var. Hartwegii 337,
342, 345, var. lavandulaefolius
345, 337, 342, var. Maccartii 343,

342, var. pubescens 344, 342, var.
Toumeyi 341; lavandulifolius
345; serrulatus 337, 338, var.
arizonicus 338, var. serrulatus
338, 340, var. spinulosus 339, 338;
tubicula 338, 341

Calylophus (Oenothera in part:

da 252
Calypso bulbosa 138, 192, 194, 196,
213

Calyptocarpus vialis 99
Calystegia sericata 386, Soldanel-
la 386

Campanula rotundifolia 164, 196,
199, 207, 212,

Campanulaceae 164, 305 (see also
Wahlenbergia)

Campsis radicans 304
Campylium stellatum 124, 202,
203

Candolle, Alphonse de 18
Caprifoliaceae 164

Capsella Bursa-pastoris 148, 210,
212

Capsicum frutescens 304

Cardamine parviflora var. areni-
cola 148; pensylvanica 148, 208
Cardiosperum Halicacabum 267

aurea 132, 203; Bebbii 133, 198;
brunnescens 133; Buxbaumli
133, 192, 202; canescens 133, 200,

211; Garberi 134,
203, 207, 210: glacialis 134, 198;

397

gynocrates 134, 194; interior 134,

flora 135; vaginata 135, 192, 194;
viridula 135, 203, 207

Carolinas 386

Caryophyllaceae 143, 264, 302 (see

Cassia occidentalis ae 303; uni-

flora 266

Castilleja 263; Pringlei 268; Rau-

pili 162, 193, 194, 199, 203, 207,

209; scorzoneraefolia 268; tenui-

flora 268

Catalogus Plantarum Angliae et
nsularum Adjacentium 258

Cathartolinum 65, sect. Multicau-

lia 65

Cayratia japonica 384

Cayratia japonica (Vitaceae) in

anothus azureus 262
Cedronella breviflora var. Ha-
vardi 1
Celtis laevigata 302
Cenchrus longisetus 182; villosus

Cenchrus longisetus M. C. John-
ston, nom. nov. (Gramineae) 182
Centaurea solstitialis 374

Central America 382

Centrosema floridanum (Britton)
Lakela, comb. nov. (Legumino-
sae) 182

Cephalanthus salicifolia 267

398

Cephalocereus 263
Ceranthera 89:
linearifolia 90
Cerastium 49, 50; anomalum 50:
argaeum 50; arvense 143,
brachypodum 98; cerastoides 50,
D1; dubiwm 50; Kotschyi 50; lap-
ponicum 51; nutans 144, 212:
persicum 50; refractum 51; tri-
gynum 51

Ceratodon purpureus 123
Ceratophyllaceae 145
Ceratophyllum demersum_ 145
204

densiflora 91;

’

Cetraria 191, 195, 199, 200, 211,

197, 200; Tilesii 122, 197
Chaetopappa Parryi 363
Chamaedaphne_ calyculata 158,

192, 201, 205
Chapman, A. W. 6, 9, 10, 25, 26
Chapman, Flora of the Southern
United States 9, 10, 11
Chara aspera var. Macounii 120:
contraria 120, 203, var. hispidula
120, 203; globularis 120
Characeae 120
Cheilanthes angustifolia 267: cu-
cullans 267
Cheiloplecton rigidum 267
Chenopodiaceae 143
Chenopodium Berlandieri_ var.
Zschackei 143, 212; capitatum
143, 208, 212; glaucum var. sali-
num 143, 208, 210, 212; hybri-
dum var. gigantospermum 143;
rubrum 143
Chondrilla lyrata 388
Chromosome numbers 43, 63, 242,
274, 306, 313, 355, 381, 383
Chromosome numbers of Linum
from the southern United States
and Mexico 63

Chromosome numbers of Sisyrin-
chium (Iridaceae) in the eastern
United States 43

Chromosome numbers in some
North American species of Ast-
ragalus (Leguminosae) 313

Chromosomes of two Moraea
(Iridaceae) from southern Africa

Chrysothamnus spathulatus 375:
viscidiflorus 375, var. ludens 374
Cicuta bulbifera 157, 205; Doug-
lasii 157, 203, 204, 205, 209
Cirsium Engelmannii 373, 375:
filpendulum 375; sp.
rae-nigrae 373, 375; virginianum
var. filtpendulum 375

Cissus japonica 384: subtruncata
268

Cistaceae 155

Citrus aurantium 303

Cladonia 191, 193, 194, 195, 199,
200, 211, 218, 224, 236: alpestris
120, 194, 197; alpicola 121, 200;
amaurocraea 121; coccifera 121:
cornuta 121, 200; degenerans 121,
200; gonecha 121; gracilis 121:
metacorallifera 121, 200: mitis

rangiferina 121, 193, 197, 200:
sylvatica 121; uncialis 121, 200:
verticillata 121, 200
Cladoniaceae 120

Clapham, Tutin and Warburg,
Excursion Flora 259; Flora of
the British Isles 257, 259
Clarkia 114

Clematis Drummondii 267; retic-
ulata 302

Clinopodium Acinos 69; coccine-
um 73; dentatum 74; georgian-
um 74; gracile 249; macrocalyx
73; vulgare 69

Coastal Plain 92, 260, 261

Collinsonia 76; angustifolia 80;
anisata 78, 80, var. major 81;
canadensis 76, 77, 79, 80, 81, var.
cordata 80, var. ovata 80, var.
puberula 79, var. punctata 79,

, 81; subg. Collinsonia 77;
cuneata 81; decussata 81; heter-
ophylla 81; subg. Micheliella 77;
ovalis 78; praecox 78, 81, 82;
punctata 76, 78: purpurea 82;
scabra 78, 82; scabriuscula 78,
80, 81; serotina 76, 77, 78, 81;
bees a 77, 78, 80, 81; urticifolia
82; verticillaris 78, 79; verticil-
lata 76, 77, 78, var. PUTPUTascens

Collomia Cavanillesiana 178, 179;
linearis 161,

Cologania procumbens 266
Comital Flora of the British Isles

Commelina 274, 278, 279, 280, 281,

282: Forskalaei 284: Gerrardii

flora 100, 283, 284; obliqua 283;
purpurea 282; scaposa 282, 284,
293; spp. 281, 282, 283, 293; Wel-
witschii 282

Commelinaceae 264, 274 (see also
Aneilema)

Commelinantia 280

Comocladia 263

Compositae 164, 264, 305, 368, 373
(see also Ammopursus, Crepis,
Evax, Filago, Machaeranthera,
Verbesina, Youngia)

Conradina 84, 89, 93; brevifolia

399

88, 85, 93: canescens 85, 87, 93
glabra 85, 93; grandiflora 85, 87,
88, 93: montana 84, 86; puberula
84, 87: verticillata 84, 85, 86, 261
Convolvulaceae 304 (see also

Calystegia)
Convolvulus sericatus 386
Conyza 110; filaginoides 264;

sophiaefolia 265

Cooley, George R. 12
Corallorhiza trifida 138, 192, 198,
194, 196

Coreopsis 368; holodasya 372;
Irmscheriana 372; oblanceolata
372; Pickeringil 371; piurana
370; suaveolens 371, 372, var.
ecuadoriensis 371, var. suaveo-
lens 372; tinctoria 98; Woytkow-
skii 372

Cornaceae 157

Cornus canadensis 157, 193, 196,
197, 199, 205, 209, 211; florida
300; stolonifera 157, 199, 208,
209, 234

Correll, D. S. 26

Corydalis aurea 147, 208, 212;
SsCilipel virens 147, 201
Corylaceae 302 (see also Betu-

laceae)

Coulter, John Merle 7, 29

Cracca cinerea 61; corallicola 60;
Curtissu 60: Mohrii 61

Crepis elegans 166; japonica 386,
var. Elstoni 388: tectorum 166,
212

Crescentia 263

Crinum americanum 301
CRONQUIST, ARTHUR 109
Cronquist, Arthur 8

Croom, Hardy 24

Crotalaria mollicula 266; pumila
266; vitellina 266

Cruciferae 147, 265 (see also
Warea)

400

Cryptogramma crispa var. acrosti-
choides 125, 198, 200

Cryptostegia 367; grandiflora 358
ucurbitaceae 305

Cunila coccinea 73; glabella 72

Curtis, M.

Cyanotis 274, 210, 00, ely 2b,

somaliensis 278; sp. 275, 276, 278,
289; speciosa 276, 289; fabeross
277, 278; villosa 278; Zenonii 277
Cyclodon 367
Cymbella 201
Cynanchum 358, 359, 365, 367;

bigerum 359, 360, var. brevi-
florum 360, 359; Blodgettii 365;
cubense 365; laeve 359, 360;
Maccartii 360, 359; Northropiae
365; Palmeri 360; palustre 304,
359, 360, 365; scoparium 365;
unifarium 359, 360; Watsonianum
360; Wigginsii 365

Cynodon dactylon 98, 99
Cyperaceae 113, 114, 132, 201
Cyperus difformis 294; oxylepis
294; uniflorus 294

Cypripedium Calceolus var. par-
viflorum 138, 192; guttatum 138,
192, 198, 194; passerinum 138,
194

Cypselea humifusa 294
Cystopteris fragilis 125, 198;
montana 125, 193

Cytological studies in Paspalum,
group Setacea (Gramineae) 306
Dandy, J. E., List of British Vas-
cular Plants 257

Darby, John 6, 9, 25

Darwin, Charles 5, 109
Darwinism 1

Dasystephana DeLoachii 107
Deam, C. C., Flora of Indiana 254
Deceiving aquatic Neptunia (Le-
guminosae) from Central Amer-
ica 382

Deciduous forests 196

Decodon verticillatus 303

Deschampsia caespitosa 130, 198,
207, 208, 210

Descurainia Richardsonii 148; So-
phia 148, 207, 212

Desmodium cuspidatum 303; pan-
iculatum 303

Diallosteira 76; punctata 76, 79
Diaperia multicaulis 253; proli-
fera 253

Dicerandra 89, 92; cerisOrs 89,

90; linearis 90; odoratissima 89,
9

Dicerandra immaculata Lakela,
sp. nov. (Labiatae) 184
Dicliptera assurgens 304
Dicranaceae 123

Dicranum Bergeri 123, 203
Digitaria floridana 381; Ischae-
mum 380, 381; violascens 380,

Digitaria Ischaemum (Gramin-
eae) in Mississippi and Texas

Diodia tetracocca 267
Diplopappus pinnatifidus 295
Ditrichaceae 123

Ditrichum flexicaule 123, 203
Dobbs, Raymond J., Flora of
Henry County, Illinois (review)

Dodecatheon pulchellum 160, 199,
03; radicatum 160

Dodenaea viscosa 263, 267

Dorstenia Drakena 267

Draba cinerea 148, 198, 199; lan-
ceolata 148

Drepanocladus 204; aduncus 124,

204; capillifolius 124, 204; exan-
nulatus 124, 204; fluitans 124;
vernicosus 124, 204

Drosera 53, 58; anglica a 202;

(Droseraceae) in the
Southeastern United States: an
interim report 53

Drosera

Droseraceae 149

Druce, G. C., Comital Flora 259
Drummond 25

Dryas Drummondii 151, 199, 201;
integrifolia 151, 192, 194, 198
199, 207

Drymaria cordata 294, 302
Dryopteris fragrans 126, 200; lu-
doviciana 300; Robertiana 126,
198

DUNCAN, WILBUR H. 346
Dupree 31

Dyschoriste humistrata 304
Dyssodia pinnata 265

Eaton, Amos 2, 3, 4, 5, 6, 17
Edinburgh 257

Edisonia 367

Egletes viscosa f. bipinnatifida 96
Eichhornia azurea 99; crassipes
Fichhornia azurea (Pontederia-
ceae) in the Texas Coastal

401
Bend: new to the United States

Elaeagnaceae 155

Elaeagnus commutata 155, 209
Elatinaceae 155

Elatine triandra 155, 206
Eleocharis acicularis 135, 205, 206,
210; nodulosa 265; palustris 135,
204, 206, 207, 210; pauciflora var.
Fernaldii 1385, 20

Elephantopus carolinianus 305
Elliott, Stephen 15, 25; Sketch of
outh Carolina

Elymus canadensis 130; innova-
tus 130, 194, 195, 196, 197, 198,

Empetraceae 155

Empetrum nigrum 155, 192, 194,
196, 200, 205, 211

Encelia scaposa var. scaposa 375,
var. stenophylla 375

Endemism in Florida 260
English Botany 258

Enslenia albida 359

Epicion 367

Epilobium angustifolium 156, 193,
194, 196, 197, 201, 212; glandu-
losum var. adenocaulon 156, 203,
205, 207, 209, 210, 212; palustre
var. oliganthum 156
Equisetaceae 125

Equisetum 201; arvense 125, 192,

192, 194, 196, 212;
, 211; variegatum

Svivalieui

Eriastrum 172; diffusum 172
Ericaceae 158, 265

Erigeron 110; acris 167; composi-
tus var. glabratus 157, 199; gla-
bellus var. pubescens 167,

>

402

199; hyssopifolius 167, 193, 207;
lobatus var. Warnockii 376, 373;
lonchophyllus 167, 204, 208: max-
imus 265; philadelphicus 167,
210; Sta v0sue 265; speciosus var.

Traversii 376; Warnockii 373
Eriogonum annuum (Polygonace-
ae) biennial in Nebraska 382
Eriophorum angustifolium 135,
200, 202, 204, 205; brachyanther-

205; spissum 135; viridi-carinat-
um 136, 202

Eruca sativa 265

Erysimum cheiranthoides 148,
209, 210, 212; inconspicuum 148

Eupatorium coelestinum — 305;
cuneifolium 376, var. semiser-
ratum 373, 377; glaucescens 373,
376; Marrubium 376; parviflo-
rum var. lancifoliwm 377; rotun-
scabridum
377; scabridum 373, 377; semi-
serratum 376, var. lancifolium

Euphorbia spathulata 98
Euphrasia subarctica 162, 210
Eustylis purpurea 295
Evax 252, 377: candida 253: mul-
ticaulis 253, 377, var. Drummon-
uw 253; nivea 253; prolifera 253,
377; verna 253, 377
Evernia mesomorpha 122
Evolution of the Gray’s and
Small’s manual ranges 1
Evolutionary parallelism 109
Ewan 31

Excursion Flora of the British
Isles 259

Exolobus albomarginatus 365
Eysenhardtia polystachya 266
Facelis retusa 294

FASSETT, NORMAN C. 382
Fatoua pilosa 248: villosa 248
Fatoua_ villosa (Moraceae) in
Louisiana: new to North Ameri-
ca 248

Fell, Egbert W., Flora of Winne-
bago County, aes 254

Festuca rubra 130: saximontana
130, 196, 198, 200, 211: tolucensis

Fewer Florida rarities: changing
flora of the Pineola Grotto, Cit-

Filaginopsis Drummondii 253
Filago 252, 377; candida 252, 377:
multicaulis 253; nivea 253; Nut-
tallii 253, 377; prolifera 253, 377;
verna 253, var. Drummondii 253,
Sri

Fimbristemma calycosa 366: ste-
nosepala 366

Fischeria aristolochiaefolia 365
Flora, Florida 257; Anglica 258;
of Henry County, Illinois (re-

Nottinghamshire 255; of the Bri-
tish Isles 257, 259: of the South-
eastern United States 10, 101; of
the Southern United States 10,
11; of Winnebago County, Illi-
nois 254

Florestina pedata 265; trifida 265;
tripteris 265

Florida 185, 257, 299

Floscopa 277

Flowering Plants and Ferns of
the Texas Coastal Bend Coun-
ties 99, 364

Fontinalaceae 124

Fontinalis Duriaei 124
Forcipella 103; Rugelii 102

Fox, W. B. 24

Fragaria virginiana var. terrae-
novae 151, 194, 196, 197, i 210,
212, 213

Fragilaria lapponica 201
Fumariaceae 147

Funastrum 367; crispum 361; cy-
nanchoides 361; Torreyi 361
Galactia Macreei 303

Galeana hastata 265

Galega 314, 322

Galeopsis Tetrahit var. bifida 161,
212

Galinsoga aristulata 265

199, 210, 212; trifidum 164, 205,

Galpinsia 337; Toumeyi 341

Gattinger 25

Geiser, Samuel Wood 25, 30, 31

Gelsemium 346; Rankiniu 346,

3 348, 349, 350, 351, 352, 353,
6,

5, 356: sempervirens 34
347, 348, 349, 350, 351, 352, 353,
354, 355, 35

Generic flora of the Southeast 11,
12, 30, 58

Gentiana amarella var. acuta 160;
DeLoachii 107; Macounii 160;

Gentiana DeLoachii (W. P. Lem-
mon Shinners, comb. nov.
(Gentianaceae) 1
Gentianaceae 160
Gentianella amarella ssp. acuta
160, 199, 203; crinita ssp. Ma-

403

counii 160, 209, 210; ssp. Raupii
160, 203, 21

Geocaulon lividum 142, 192, 193,
194, 197, 201, 211

Geraniaceae 154

Geranium Bicknellii 154, 209, 212
Geum macrophyllum 151, 209;
triflorum 151, 199

Gibbesia 103; Rugelu 103

Gilia 171, 172, 173; aggregata var.
texana 177; pono var. ait iuse

flora 177; longiflora 177; ludens
174, 173; Macombii var. laxiflora
177; mexicana 173; multiflora

cladon 178; pumila 178; rigidula
175, ssp. acerosa 250, var. acero-
sa 173, 175, ssp. insignis 175, var.
rigidula 173, 175; rubra 177;
Stewartii 173, 175; texana 177;
Wrightii 178

Gilia and Ipomopsis (Polemoni-
aceae) in Texas 171

Gleason, Henry A. 8, 18, 113
Glyceria borealis 131, 206, 211;
grandis 131, 205, 206, 211; pul-
chella 131, 205, 206; striata 131,

Gnaphalium uliginosum 167, 211
GODFREY, R. K. 185, 269
Godwin, H., History of the Bri-
tish Flora 257

Gomphocarpa 367

Gonolobus 367; albomarginatus
365; aristolochiaefolius 365;
arizonicus 366: calycosus 366;
chiapensis) 366; cteniophorus 366;

gonoloboides 366; laevis 359;
Lasiostemma 366; oblongifolius
366; parvifolius 363; sagittifolius

404

363; stenanthus 366; stenopetalus
366; stenosepalus 366
Gramineae 129, 301 (see also Cen-
chrus, Digitaria, Monanthochloe,
Paspalum)

Gray, Asa 4, 5, 6, 9, 18, 29, 30, 31
Gray Herbarium 7, 11, 12, 14, 19
Gray’s Manual 4, 5, 6, 7, 8, 10
Greenman, J. M. 11
Grimmiaceae 123

Guazuma ulmifolia 263
Gyrophoraceae 121

Habenaria hyperborea 138, 192,
207; obtusata 138, 192, 194; quin-
queseta 302

Haloragaceae 156
Hamamelidaceae 302
Haplopappus _ divaricatus 305;
texensis 378

Haplophytum cimicidum 264
Hedeoma arkansana 72; glabrum

Hedwigia ciliata 123, 200
Hedysarum 192: a
americanum 153, 192, 194, 196,
209, 210; Mackenzii 154, 192, 193,
196, 199
Helenium autumnale 167, 199
Helianthus annuus ssp. texanus
77, var. texanus 377: ludens
377
Helictotrichon Hookeri 131, 198
Hemianthus glomeratus 252
Henderson, N. C. 26
Heracleum lanatum 157
Herniaria americana 102
Heteranthera limosa 267
Heterotheca inuloides 265
Hexaploid Linum (Linaceae)
from eastern Ethiopia 383
Hieracium umbellatum 167, 196,
199, 210
Hierochloe odorata 131
Himantostemma 367

Hippuridaceae 156

Hippuris vulgaris 156, 204, 205,
206, 207, 209

History of the British Flora 257

Hookers 258

Hordeum jubatum 131, 207, 210,
212; pusillum 98

How to study the Florida flora
257

Howitt, R. C. L. and B. M., Flora

of Nottinghamshire 255

Hudson, William, Flora Anglica
8

Hudsonia tomentosa 155, 196, 211,

Hydrophyllaceae 161
Hylocomium 191, 192, 194, 214,
216; splendens 124, 191, 193, 194,
196, 205, 216

Hypericaceae 265

Hypericum pauciflorum 265
Hypnum Bambergeri 124
Hypogon 76; anisatum 78; verti-
cillare 79; verticillatum 76, 78
Hyptis mutabilis 304

Identity of Sagittaria isoetiformis
(Alismataceae) 269

Ipomoea arborea 263; trichocarpa
304

Ipomopsis 171, 172, 176; aggre-
gata var. texana 177; Havardii
176, 177; laxiflora 176, 177; lon-
giflora 176, 177; multiflora 178;
pinnata 179; polycladon 176, 178:
pumila 176, 178; rubra 177;
Wrightii 178, 176

Iridaceae 138, 301 (see also Mo-
raea, Sisyrinchium, Tigridia)
Irwin, Howard S., Roadside
Flowers of Texas (review) 296
Isoetaceae 125

Isoetes echinospora bar. Braunii
125, 206

Iva angustifolia var. angustifolia
378, var. lata 378
Jack pine forest 195
James, C. W., Endemism in Flori-
da 260
James, Edwin 25
Johnson, Thomas 258
Johnston, M. C. 25
HNSTON, MARSHALL C. 182

the Texas Coastal Bend Coun-
ties 99, 364

Juliana adstringens 263
Juncaceae 113, 114, 136
Juncaginaceae 129

Juncus 113; albescens 136; alpin-

stygius var. americanus 137, 202;
Vaseyi 137, 205, 211

Juniperus communis var. depres-
sa 126, 192, 193, 194, 196, 198,
200, 220; horizontalis 126, 192,
194, 196, 198, 200, 220

Jussiaea macropoda 385; monte-
vidensis 386; octonervia f. sessi-
liflora 385, var. sessiliflora 385;
repens var. glabrescens 386, var.
montevidensis 386, var. pep-
loides 386; suffruticosa 385, var.
ligustrifolia 385, var. octofila
3

5
Kalmia polifolia 158, 192
Karwinskia umbellata 267
Keithia 87

Key to southeastern glabrous-
styled Tephrosia (Leguminosae)

Koeleria cis 131, 198

(see also
Acinos, Calamintha, Clinopodi-

phis, Scutellaria, Teucrium, Tri-
chostema)

Lachnostoma 367; arizonicum
366; gonoloboides 366; lasio-
stemma 366

Lacinaria Ohlingerae 240
Lactuca pulchella 167, 208, 210
Lagascea rubra 265

LAKELA, OLGA 182, 184, 240,

Lantana 263; achyranthifolia 268;
Camara 268; hispida 68; velutina

Lappula echinata 161, 212; Re-
dowskii var. occidentalis 161,
Diz

Larix laricina 126, 191, 192, 200,
202, 203
Lasallia 199, 224; pensylvanica
121, 200
Lathyrus ochroleucus 154, 194,

Lecanoraceae 121
LEDINGHAM, G. F. 313

Ledum decumbens 158, 191, 192,
201, 211; groenlandicum 159,
191, 192, 193, 194, 196, 201, 205,
211

Leguminosae 153, 266, 303 (see
also Astragalus, Centrosema,
Neptunia, Tephrosia)

Lemna minor 136, 204, 206; tri-
sulea 136, 204, 206

Lemnaceae 136

Lentibulariaceae 163

Leonotis nepetaefolia 304
Lepidium Bourgeauanum 148,
208; densiflorum 148, 212

406

Leptobryum pyriforme 123
Leptoglossis 180; texana 180
viscosa 180

Leptoglossis and Nierembergia
(Solanaceae) in Texas 180
Leptorrhoeo 277
Lesquerella arctica var.
manae 148, 198, 199
LEWIS, WALTER H. 43, 63, 274,
381, 383

Lewis, Walter 25

Liatris cylindracea 242; cymosa
242; Garberi 240, 241: Ohlin-
gerae 240

Lichens 121, 220, 222, 224, 236
Liliaceae 137,

Limnosciadium pinnatum 98
Limosella aquatica 162, 206
Linaceae 154 (see also Linum)
Linanthus 172; aureus 173; Bige-
lovii 172

Lincecum, Gideon 25, 26
Lincoln, Mrs. 3

Lindheimer 25

Lindley, John, Synopsis of the
British Flora 258

Linnaea borealis var. americana
164, 192, 193, 194, 195, 211
Linnaean System 2, 5,

Linum 63, 328: alatum an 329,
333, 334; arenicola 63, 65, 67;
aristatum 328, 330, 333, 334:
australe 330, 335, 336, var. glan-
dulosum 336, 330, 335; compac-
tum 336; floridanum 329, 330,
var. chrysocarpum 63, 65, 67:
grandiflorum 63, 65, 67; Greggii

,

Scam-

; perenne 63, 67; pra-
tense 63, 65, 67; puberulum 328,

329, 332, 333; rigidum 63, var

68
Lippia 263; Berlandieri 268
Liquidambar Styraciflua 303
List of British Vascular Plants

Loasaceae 266
Lobelia homophylla 305; Kalmii
64, 203

Loeselia 171, 172; scariosa 172
Loganiaceae (see Gelsemium)
Lomatogonium rotatum 160, 203,

Lonicera dioica var. glaucescens
164, 194, 196, 199

Lopezia mexicana 267

Louisiana 104, 182, 248, 249, 251,
294, 384

Ludwigia 385; adscendens var.

pepioides 386: octovalvis ssp.

macropoda 385, var. macropoda
octovalvis 385, ssp.

Socal tions 385, var. sessiliflora

385; peploides 386, ssp. glabres-

cens 386, var. glabrescens 386,

ssp. montevidensis 386, var.

montevidensis 386; pubescens

Lundell, C. L. 25

Lupinus 263; elegans 266

Luzula 113

Lycopodiaceae 125

Lycopodium 201; annotinum 211:

complanatum 125, 1

Lyonia 367

Lysimachia Japonica 249
Lysimachia japonica (Primula-
linopodium gracile
(Labiatae) in Louisiana: new to
the United States 249
Lythraceae 303

Macbridea caroliniana 74; pul-
chra 74

Machaeranthera annua 378; Bol-
toniae 378; brevilingulata 378;
pinnata 295, 378; pinnatifida 295,
378; spinulosa 295; tenuis 378;

pinnatifida
(Hooker) Shinners, comb. nov.

Malvaceae 267, 303 (see also
Sphaeralcea, Wissadula)
Malvastrum coromandelianum
Manual of the Southeastern Flo-
ra 10

Manual of the Southwestern Flo-
ra ll

Marchantia polymorpha 124
Marchantiaceae 124

Martynia annua 267
Martyniaceae 267

Mason-Dixon Line 29

; oblongifolia 366;
parviflora 362, 363; parvifolia

culata 362, 364; sagittifolia 362,
363; Smithii 366; stenantha 366;

407

stenopetala 366; stenosepala 366;
Woodsonii 364,

Matricaria maritima var. agrestis
167, 212; matricarioides 167

Meesia tristicha 123, 204

Meesiaceae 123

Meiotic chromosomes in African
ommelinaceae 274

Melampodium oblongifolium 265;

padudosum 265

Melandrium Ostenfeldii 144, 199,

200, 207

Melastomaceae 304

Melilotus alba 154, 212; officinalis
154,

Melinia 367; angustifolia 365
Melissa coccinea 73

Mellichampia 367

Melochia pyramidata 99, 268
Melothria pendula 305

Mentha arvensis 161, var. arven-
sis 161, var. villosa 161, 203, 208,

Mentzelia aspera 266
Menyanthes trifoliata 166, 191,
202, 204, 206

Meriolix 337; melanoglottis 340,
341

Mertensia paniculata 161

Metastelma 367; angustifolium
365; arizonicum 365; barbigerum
360; Blodgettii 365; Palmeri 360
Mexico 262

Michaux 25

Micheliella 76, 77; anisata 76;
verticillata 77, 78, 79
Micranthemum glomeratum 252;
Nuttallii var. glomeratum 252

Micranthemum glomeratum

408

(Chapman) Shinners, comb. nov.
(Serophulariaceae) 252
Micromeria bahamensis 96, 95:
Brownell 94, var. Brownei 70,
94, 95, var. ludens 96, 94, var.
pllosiuscula 69, 94,
gensis 96, 95: glabella var. an-
gustifolia 72; Nuttallii 72: pilo-
stuscula 69, 95: stolonifera 96;
xalapensis 69, 95

Micromeria Brownei and its al-
lies (Labiatae) 94

Mikania cordifolia 295

Milla biflora 266

Mimosa albida 266; Benthamii
266; caerulea 266

Mimulus glabratus 268
Minuartia 49; Nuttallii 49
Mississippi 380

Missouri Botanical Garden 11
Mitchella repens 304

Mitella nuda 149, 192, 193, 205
Mohr 25

Moldavica parviflora 161, 212
Monanthochloe littoralis (Gra-
mineae) in Louisiana 182
Moneses uniflora 157, 192, 193,
194, 205

Moraceae 267 (see also Fatoua)
Moraea 381; Erici-Rosenii 381:
setacea 382

More additions to the Louisiana

Muhlenbergia glomerata var. cin-
noides 131, 202; Richardsonis 131
Murdannia 274, 278, 285, 286, 287,
293; elata 286; Keisak 286; semi-
teres 285; simplex 285, 286, 293:
stinicum 285

Myla anomala 124

Myrica Gale 141, 192, 194, 203, 205
Myricaceae 141

Myriophyllum exalbescens 156,

204, 206; verticillatum var. pec-
tinatum 156, 204,
National Science Foundation 12,

Natural System 2, 4 ,5, 18
Naumburgia thyrsiflora 160, 205,
206, 209, 210

Navicula tuscula 201

Nebraska 382

Nemastyls 295

Nemostylis purpurea 295
Neo-Darwinism 18, 20

Neptunia plena 383, f. lumbri-
coides 382; prostrata 382

New combinations in Texas Pole-
moniaceae 250

New names and records for Texas
Compositae 373

New names in Arenaria (Caryo-
phyllaceae) 49

New or otherwise interesting
Coreopsidinae (Conpositae)
from northwestern South Amer-
ica 368

New varietal names for New
World Ludwigia (Onagraceae)

New York Botanical Garden 7,
Nicotiana glauca 268
Nierembergia 180; hippomanica
var. caerulea 181; viscosa 180
Nissolia fruticosa 266; hirsuta 266
Nitzschia angustata 201
Nomenclatural codes 18
Northwest Territories, Canada
117, 187

Notes on Calystegia (Convolvu-
laceae) in the Carolinas 386
Notes on the flora of the Mexican
state of Morelos 262

Nothocalais cuspidata 373, 378
Notholaena aurea 267
Nothoscordum fragrans 264

Nuphar variegatum 145, 204, 206,
228

Nymphaea tetragona ssp. Libergli
145, var. Porsildii 204
Nymphaeaceae 145

Nyssa biflora 303

Nyssaceae 303

Ochrolichia inaequatula 121

Odontonychia corymbosa 102; in-
terior 103

Odontostephana 367; decipitens

men ounOhe 114, 337; Fendleri 343;

ar. filifolia 345, var. glandulosa
45, var. lavandulaefolia 345,
var. Toumeyi 341; lampasana
344: lavandulaefolia 345; serru-
lata 338, 340 ’
388, 340, var. Nuttallii 338, var.
pinifolia 340, var. serrulata 338,
var. spinulosa 339; spinulosa 339;
tubicula 341, var. pr 345
OLIVER, ROYCE
Onagraceae 156, 267, if (see also
Calylophus, Ludwigia, Oeno-
thera)
Ophioglossaceae 300
Oplismenus setarius 301
Opuntia 26
Orchidaceae 138, 302
Orchis rotundifolia 138, 194
Orobanchaceae 162
Oryzopsis asperifolia 131, 196;
pungens 131, 192, 196, 198, 212
Osmunda regalis 300
Osmundaceae 300
Otopappus robustus 265
Outline of the Geographical Dis-
tribution of British Plants 258
OWENS, A. G., Jr. 182

409

Oxycoccus microcarpus 159, 191,
192, 194

Oxypetalum 359, 367; caeruleum
359

Oxypteris 367

Oxytropis 313, 314, 315, 319, 322;
campestris var. gracilis 321, 322,
ssp. sordida 322, var. varians 154,
199; deflexa var. sericea 154,
321; Halleri 321; monticola 321,
; multiceps 321; Parryi 321,
326; sericea var. spicata 321;
splendens 154, 196, 199; viscida
154

Pachycereus marginata 263
Pachyrrhizus erosus 266
Paederia Pringlei 267

Palmae 301

ww
i)
aD

Paludella squarrosa 123
Pancratium littorale 264
Panicum anceps 301; Joorii 301
Parietaria floridana 302

Parmelia 199; centrifuga 122, 200;
stenophylla 122, 200; sulcata 122,
200

Parmeliaceae 121

Parnassia multiseta 149, 191, 192,
202, 203, 209

Paronychia 101; americana 102;
Baldwinii 103; diffusa 102;
Drummondii 102, 103; erecta
102; fastigiata var. Nuttallii 103;
patula 102; Rugelii 102

Parthenocissus quinquefolia 303

Paspalum 306; ciliatifolium 306,
309, 310: conjugatum 301; debile
306, 309, 310; dilatatum 98, 99,
307; epile 310; ;
longepedunculatum 306, 309,
311; longipilum 311; Pariodia-
num 310; propinquum 306, 309,
311; psammophilum 306, 309,
311; pubescens 306, 307, 311;
rigidifolium 306, 309, 311; seta-

410

ceum 306, 309, 312; stramineum
306, 309, 312; supinum 306, 309,

Pattalias Palmeri 360

5)
Pedicularis labradorica 162, 198,
194, 196; mexicana 268

Peltigera aphthosa 120, 192, 193,
; malaca 120

Peltigeraceae 120

Pennisetum villosum 182

Penstemon campanulatus 268

Peperomia leptostachya 302

Periploca 359, 364, 367; graeca

Perring, F. H., and S. M. Walters,
Atlas of the British Flora 257
Persea americana 263; Borbonia
300

Petasites frigidus var. nivalis 167,
var. palmatus 168; palmatus 168;
sagittatus 168; vitifolius 167

Petiveria alliacea 302

Phacelia Franklinii 161, 207, 213

Phalaris arundinacea 131, 206,
208, 212, 230; canariensis 131,
yaw

Phaseolus stropurpureus 266;
coccineus 266; heterophyllus 266

Pherotrichis 367

Phleum pratense 132, 212

Tharpii 250; Johnstonii 172, 250;
oklahomensis 172; pinnata 179;
Tharpiit 250

Philibertella 367; crispa 361;
cynanchoides 361;

361; Torreyi 361

Philibertia 367

Phyllanthus Urinaria 294

Hartwegii

phylogeny 109—116
Physcia muscigera 122
Physciaceae 122

238; mariana 127, 192, 198, 199,
200, 201, 203, 211, 214, 222
Pinaceae 126

Pinaropappus roseus 265

Pineola Grotto 299

Pinguicula villosa 163; vulgaris

Pinus ayacahuite 263; Banksiana
127, 195, 198, 200, 211, 214, 218,
222, 224, 236, 238; Hartwegii 263;
Lawsonii 262; Montezumae 262;
teocote 262

Piperaceae 302

Pisum sativum 266
Pithecolobium 263
Plagiochilaceae 124
Plantaginaceae 163

Plantago major 163, 209, 210, 211;
septata 163, 199

Pleuradenia 77; praecox 17;
scabra 77, 79

Poa 132; alpina 132, 198; annua
98, 99; Canbyi 198; glauca 132
198, 208; interior 132, 198, 200,
207; leptocoma 132, 210; palus-
tris 132, 210; pratensis 132, 208,
212; stenantha 132

Podostigma 367

Polemoniaceae 161, 171, 250
Polemonium 171, 82; Hinckleyi
172; pauciflorum 172; rubrum
177

?

Polygonaceae 142 (see also Erio-
gonum
Polygonum achoreum 142, 212;

amphibium 142, 204, 205, 206,
210; aviculare 142, 208, 210, 212;
coccineum 142; Convolvulus 142,
212; lapathifolium 142, 206, 208,
210, 211; viviparum 142, 192
Polypodiaceae 125, 300
Polypodium dispersum 300; pec-
tinatum 300; plumula 300; poly-
podioides 300, var. aciculare 267;
virginianum 126,
Polytrichaceae 123

?

Polytrichum formosum 123, 200;
juniperinum 123, 200, var. alpes-
tre 123; piliferum 123, 200
Pontederiaceae 267 (see also
Eichhornia)

Populus 238; balsamifera 139, 196,
197, 198, 208, 209; tremuloides

formis 128, 206, var. borealis 128,
, ’ . Macouni 128;
foliosus 128, 204; Friesii 128, 204;
gramineus 128, 204, 206; pusillus
128, 204, 206; Richardsonii 128,
204, 206; vaginatus 129, 206;
vaginatus 129, 206; zosteriformis
129, 204
Potentilla anserina 151, 203, 209,

Prenanthes lyrata 387, 388
Primula egaliksensis 160; incana
160, 203; mistassinica 160; stricta

Primulaceae 159, 304 (see also
Lysimachia)

Pringle, C. G. 295
Prunus pensylvanica 152

Psilotum nudum 300
Psychotria nervosa 304
Pteris cretica 301; vittata 301

Pycnothymus rigidus 70

Pyrola asarifolia 192, 193, 196,
var. purpurea 157; grandiflora
157, 192, 193, 194, 205; ;
secunda 158, 193, 196; virens 158,
192, 194, 196

Pyrolaceae 157

Pyrrhopappus carolinianus 305
Quercus 262; Michauxii 302; Shu-
mardii 302; virginiana 302
Quillin, Ellen Schulz 25
Rafinesque 24, 26, 31
Ranunculaceae 145, 267, 302 (see
also Ranunculus)

Ranunculus aquatilis var. eradi-
catus 146, 206; circinatus var.
subrigidus 146, 206; Cymbalaria

205, 207, 208, 210; trachycarpus
104; trilobus 104

Ranunculus trachyearpus (Ran-
unculaceae) in south central
Louisiana: new to North Ameri-
ca 104

Raven, Peter 337

Ravenel, Henry William 14, 30
Ray, John, Catalogus Plantarum

412

258; Synopsis Methodica 258
Rendle, A. B., Biographical Index
of Deceased British and _ Irish

reviews 254, 296, 389
Rhamnaceae 267

Rhapidophyllum Hystrix 301
Rhexia mariana var. exalbida 304
Rhinanthus Crista-galli 162, 199,

Rhododendron lapponicum 159,
192, 194

Rhus Toxicodendron 303
Rhynchosia pyramidalis 266
Rhytidium rugosum 194

Ribes glandulosum 149, 201; hud-
sonianum 150, 192; lacustre 150,
194, 196; oxyacanthoides 150,
199, 201, 209; triste 150, 194, 205
Ricciaceae 125

Ricciocarpus natans 125

RICHE, SAM 182

Ridan paniculatum 253

Riddell 2

Rivina humilis 302

Roadside Flowers of Texas (re-
view) 296

Robinson, Benjamin Lincoln 7
Roemer 25

ROGERS, C. M. 328

Rollins, Reed C. 12

Rorippa crystallina 149; islandica
149, 205, 207, 209, 210

Rosa acicularis 152, 192, 193, 194,
195, 196, 197, 198, 199, 201, 209,
211

Rosaceae 152

Rosmarinus officinalis 87
Rothrockia 367

Rowlinia 367; Palmeri 360

Rouliniella 367; unifaria 360

ROWELL, CHESTER M. 262

Rowell, Chester 25

Rubiaceae 163, 267, 304

Rubus acaulis 152, 192, 194:
Chamaemorus 153, 191, 201:
idaeus var. canadensis 153, 201,
203; paracaulis 153; pubescens
153, 192, 197, 209; trivialis 302
Rugel 25

Rumex maritimus var. fueginus
142, 208, 205, 208, 210, 211;
mexicanus 142, 212: occidentalis

Rutaceae 303

Rydberg 31; Flora of the Prairies
and Plains 12

Sabal Etonia 241; minor 301
Sageretia minutiflora 303

Sagina fontinalis 51

Sagittaria cuneata 129, 204, 206,
210; graminea 269, 270, 27
isoetiformis 269, O71. 272; evict
folia 264; teres 269, 270, 271
Salicaceae 139

Salisbury, E. J. 259

Salix 205, 232, 288: arbusculoides
139, 198; athabascensis 139, 202;
Bebbiana 139, 196, 197, 198, 201,
205, 209; brachycarpa 139, 203,
207; calcicola 139; candida 139,

liana 140; myrtillifolia 140, 192,
194, 209; padophylla 140; pedi-
cellaris 140, 203; petiolaris 140;
planifolia 140, 205, 209, 210:
pseudomonticola 140; pyrifolia
201; reticulata 140, 194; Scou-
leriana 140, 205; serissima 141,
Salvia coccinea 304; lyrata 304
Samolus parviflorus 304
Sanicula canadensis 304
Santalaceae 142

Sanvitalia procumbens 265

Sapindaceae 267, 303

Sapindus marginatus 300, 303
Sarcostemma 358, 359, 360, 367;
bilobum ssp. Lindenianum 365,

wegiti 361, var. Hartwegii 361;
Lindenianum 365; lobatum 361;
Torreyl 361

Sass, Herbert Ravenel 30
Satureja 69; Acinos 69; Ashei 73;
Brownei 95, var. pilosiuscula 70,
95; Calamintha 71; caroliniana

grandiflora 71; macrocalyx 73;
Nepeta 71: rigida 70

Saxifraga aizoides 150; tricuspi-
data 150, 198, 199, 200
Saxifragaceae 149, 302
Schizaeaceae 300
Schizonotus 367
Schkuhria multiflora 374
Scirpus acutus 136; ce

var. callosus 136, 191, 192, 202,

divaricatus 265;

Scolochloa festucacea 132, 203,
206, 208

Scorpidium scorpioides 201
Scrophulariaceae 113, 162, 268
304 (see also Micranthemum)
Seutellaria Drummondii 251; ga-
lericulata var. epilobiifolia 162,
205, 208; laevis 107; muriculata
251; Thieretii 251

Scutellaria laevis (Labiatae), an-
other endemic in Trans-Pecos
Texas 107

2

413

Scutellaria Thieretii (Labiatae),
a new species from coastal Lou-
isiana 251
Selaginella 201;
selaginoides 125,
Selaginellaceae 125, 26

Senecio congestus 168, 204, 205,
207; eremophilus 168; imparip-
innatus 98; indecorus 168, 208;
lugens 168, 191, 193; pauperculus
168, 204, 208; plattensis 168:
Riddellit var. Parksii 379; spar-
tioides 379, var. Parksii 379;
tridenticulatus 168, 196, 199;
vulgaris 168, 212; Warnockii 379
Serenoa repens 301

Setcreasea 277

Seutera 367: palustris 360
Shepherdia canadensis 156, 192,
193, 194, 195, 196, 197, 198, 199,
201, 207,

SHERFF, EARL EDWARD 368

pallescens 268;

Shinners, Lloyd H. 12, 13, 25, 31;
Spring Flora 53, 99, 297

Sibara virginica 98

Sida procumbens 267

Silene Menziesii 144, 210, 212
Simpson, George Gaylord 111, 112
Siphonychia 101; americana 102:
corymbosa 102; diffusa 102;
erecta 102; interior 103; pauci-
flora 102; Rugeli 102
Siphonychia transferred to Paro-
nychia (Caryophylaceae) 101
Sisyrinchium 32, 43; albidum 43;
angustifoltwm 43, 45; arizonicum
43, 47; atlanticum 438, 46, 47;
Bermudiana 43, 46, 48, var. mi-

414

nus 33; campestre 43, 45, 46;
Canbyi 33; ensigerum 43, 45;
exile (see also micranthum) 33,

medium 43, 45, 46; Langloisii 43,
45; laxum (see also rosulatum)

45, 301; sagittiferum 43, 45, 46,
47; Thurowii 33, 36; uniflorum
41, valdivianum 41

Sium suave 157, 206, 209, 210, 211
Small, John Kunkel 10, 31; Flora
of the Southeastern United
States 10, 12; Manual of the
South Central Flora 11; Manual
of the Southeastern Flora 10, 11
Smilacina stellata 137, 210; tri-
folia 137, 191, 192, 201, 210, 212
Smilax auriculata 301; Bona-nox

Smith, J. E., English Botany 258
Solanaceae 113, 268, 304 (see also
Leptoglossis, Nierembergia, So-
lanum)

Solanum bicolor 268; floridanum
108, 304; Godfreyi 108; nigrum
Solanum Godfreyi Shinners,
nom. nov. (Solanaceae) 108
Solidago 213; canadensis var.
salebrosa 168, 210, 212; multi-
radiata 169, 193, 194, 213; peti-
olaris 379; spathulata var. neo-
mexicana 169, 196, 199, 201, 208,

3

Sonchus arvensis var. glabrescens
169, 212
Sowerby, James, English Botany

Sparganiaceae 127

Sparganium angustifolium 127,
, 206; minimum 128, 202, 204,

206, 210

Spermacoce Haenkeana 267; pa-

tula 267

Sphaeralcea 384; angustifolia ssp.

var. violacea 385; digitata var.
angustiloba 385, ssp. tenwipes
385, var. tenuipes 385; Emoryi
var. californica 384, ssp. vari-
abilis 384, var. variabilis 384;
Fendleri var. californica 384:
pedata var. angustiloba 385;
tenuipes 385

Sphagnaceae 122

Sphagnum 191, 192, 194, 201, 203,
205; balticum 122: capillaceum
, 191; cuspidatum 122; fus-
cum 122, 191, 203; Girgensohnii
122, 191; riparium 122; squarro-
sum 123, 191; Warnstorfianum

Sphenopholis intermedia 132, 210
Spilanthes americana var. par-
vula 265

Spiranthes Romanzoffiana 139,
192, 213

Spirogyra 114

Spironema fragrans 277

Spring Flora of the Dallas-Fort
Worth Area, Texas 53, 99, 297
St. Augustine grass sod 98
Stachys palustris var. nipigonen-
sis 162, 209, 210

Stanleya amplexifolia 106
Starkea pinnata 295

Stauroneis 201

Stellaria 49, 50; calycantha 144;
cerastoides 51; Corei 103; cras-
sifolia 144, 205, 208; dichotoma
50, var. americana 50; dubia 50;
fontinalis 51; Jamesiana 50;

corum 951; Nuttatlii “49: paludi-
cola 51; pubera 104, ssp. stlvati-
ca 103, var. silvatica 103, var.
tennesseensis 104; silwatica 103;
sylvatica 103; tennessensis 103;
uniflora 51; viscida 50
Stellaria Corei Shinners, nom.
nov. (Caryophyllaceae) 103
Stenotaphrum secundatum 98
Sterculiaceae 268

Stereocaulon tomentosum 121
Stevia serrata 265

Subularia aquatica 149, 206
Symphoricarpos occidentalis 164
Synopsis Methodica
Britannicarum 258

Stirpium

Synopsis of Collinsonia (Labia-
tae) 76; Conradina (Labiatae)
84; Dicerandra (Labiatae) 89;
the British Flora 258
Systematic status of Ammopur-
sus Ohlingerae (Compositae)
240

Tagetes filifolia 265;
265; lucida 265
Tanacetum vulgare 169, 212
Taraxacum ceratophorum 169,
199, 208; officinale 169, 212
Taxodiaceae 301

Taxodium distichum 263, 301
Taxonomic significance of evo-
lutionary parallelism 109
Taxonomy and_ heterostyly of
Gelsemium

jaliscana

North American
(Loganiaceae) 346
Tectaria heracleifolia 301

415

Tephrosia 60; ambigua var. gra-
cilima 61; angustissima 60, 61;
cinerea 61; corallicola 60; Cur-
tissii 60; florida var. gracillima
61; Mohrii 61; nicaraguensis 266;
Seminole 60, 61

Texas 25, 98, 99, 107, 171, 180,
250, 252, 296, 328, 337, 358, 373,

Texas Asclepiadaceae other than
Asclepias 358

Texas Evax transferred to Filago
(Compositae) 252

Teucrium canadense 182, var.
boreale 183, var. canadense 183,
var. Nashii 183, var. occidentale
183; Nashii 183; occidentale 183,
var. boreale 183

Thalictrum venulosum 147, 210
Thamnolia vermicularis 122
Tharp, B. C

Thelesperma_ curivearpum — 373,

Thelypteris dentata 301; normalis
301; reptans 301; tetragona 301;

Thlaspi arvense 149, 212

Three new varietal names in
Sphaeralcea (Malvaceae) 384
Thuidiaceae 124

Thurovia 261

Thymbra caroliniana 74

Thymus carolinianus 74; grandi-
florus 74

Tigridia buccifera 295; purpurea
295; vaccata 295

Tigridia purpurea (Herbert)
Shinners, comb. nov. (Iridaceae)
295

Tilia floridana 303

Tiliaceae 303

416

Tilandsia simuata 301; usneoides
301
Tofiedia gutinosa 137, 191, 192,

Topographic Botany 259
Torrey, John 4; Torrey and Gray

Tradescantia 277; commelinoides
264; linearis 101; Wrightii 101
Traverse, Alfred 25

Trelease, William 10, 11
Triceratella 274

Trichostelma oblongifolium 366
Trichostema suffrutescens 92
Tridax coronopifolia 265; pro-
cumbens 265

Trientalis europaea var. arctica
160

Trifolium amabile 266; incarna-
tum 105; resupinatum 104, 105
Triglochin maritima 129, 191, 192,
202, 203, 207, 210; palustris 129,
202, 203, 207, 210

Trisetum spicatum 132

Turner, William 258

Turnera ulmifolia 268
Turneraceae 268

Two Youngias (‘“Crepis japoni-
ca”: Compositae) introduced in
the southern United States 386
Typha latifolia 127, 204, 206, 207,

Typhaceae 127

Umbelliferae 157, 304

Uniola sessiliflora 301

United States National Herbari-
um 14

Urtica chamaedry oides 302; diol-

ca var. procera 142, 207
Urticaceae 142, 302

Usnea carnosa 122

Usneaceae 122

Utricularia 191; intermedia 163,
202, 203, 204, 206; minor 163,
204, 206; vulgaris 163, 202, 203
204, 206

Vaccinium leucanthum 265; uli-

Valeriana scandens 305
Valerianaceae 305

Vaseyochloa 261

Vegetation of disturbed soil 211;
of lakes and rivers 201; of marl
and gypsite deposits 207; of rock
outcrops 197; of strands and is-
lands 208

Verbenaceae 268, 304

Verbesina alba 253; Coreopsis
var. alba 253; paniculata 253:

Veronica americana 268; pere-

grina 209, 211, var. xalapensis
162; scutellata 162, 209, 210

Vincetoxicum 367; biflorum 362;

brevicoronatum 363; chiapense
366; cteniophorum 366; cynan-
choides 363: gonocarpos 364:
parviflorum 363; productum 364;
reticulatum 364; stenanthum 366
Vines, R. A. 26

Viola adunca 155; floridana 303:
nephrophylla 155, 192, 210: reni-
folia 155

Violaceae 155, 303

Vitaceae 268, 303 (see also Cay-
ratia, Cissus)

Vitex mollis 268

Vitis rotundifolia 303

VOSS, EDWARD G. 389
Wahlenbergia linarioides 185,
186; marginata 185, 186
Wahlenbergia linarioides (Camp-
anulaceae) in Florida: a second
adventive species for the United
States 185

Walter, Thomas 24, 25

Waltheria americana 264

Warea amplexifolia 105, 106;
auriculata 106, 105; sessilifolia
105, 106

Warea auriculata instead of W.
amplexifolia of Small (Cruci-
ferae) 105

Warnock, Barton 25

Watson, H. C., Outline of the
Geographical Distributions of
British Plants 358; Topographic
Botany 259

Watson, Sereno 7

Webster’s Third New Interna-
tional Dictionary of the English
Language (review) 389
Wernham 112, 116

WHERRY, EDGAR T. 250

White spruce forest 193
Whitehouse, Eula 25

Wills, Mary Motz, Roadside
Flowers of Texas (review) 296

417

Wissadula grandifolia 106, 107,
var. brevipedunculata 107, var.
macrantha 107; macranse 106,

Wissadula grandifolia instead of
W. macrantha (Malvaceae): no-
menclatural corrections 106
Withering, William 260; Botani-
cal Arrangement 258

Wood, pong 3, 4, 5, 6, 7, 9
Wood, C. 12

Woodsia ee 126, 198; ilven-
sis 126, 198, 200

Wooton and Standley, Flora of
New Mexico

Wright, Charles 25, 99
Yellow-flowered Linum = (Lin-
aceae) in Texas 328

Yellowknife Highway 117, 187
Youngia 386; japonica 386, 387,
388, ssp. Elstonii 388; lyrata 387,
388: Thunbergiana 387, 388
Zanthoxylum Fagara 303
Zebrina 277

Zexmenia aurea 265; crocea 265;

Zygadenus elegans 138, 192, 194,
196, 198, 209, 210, 212, 213